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Lansade, L., Bertrand, M., & Bouissou, M. - F. (2005). Effects of neonatal handling on subsequent manageability, reactivity and learning ability of foals. Appl. Anim. Behav. Sci., 92(1-2), 143–158.
Abstract: Behaviour is an important factor to be taken into account in the various uses of horses. Today horses are mainly used for sport and leisure activities. They should therefore be easy to manage, calm and not fearful. Early handling is known to improve manageability and learning ability and to reduce fearfulness in various species. It has become fashionable in the horse industry to use an early training procedure, referred to as “imprint training”, which is said to produce durable if not permanent effects. However, no studies concerning the long-term effects of such neonatal handling have been carried out in horses. The present study examines the short- and long-term effects of neonatal handling on manageability, general reactivity and learning ability of foals. Twenty-six Welsh foals were studied: 13 were handled daily for 14 days from birth and 13 were non-handled controls. The handling procedure consisted of fitting a halter, gently patting all parts of each foal's body, picking up feet and leading over 40 m. Two days, 3 months, 6 months and 1 year after the end of the handling period, foals underwent behavioural tests to measure their manageability and various aspects of their reactivity. The results showed that neonatal handling has only short-term effects on manageability: 2 days after the handling period, handled animals were significantly easier to handle than controls for the four parameters measured during this test (time to fit a halter, time to pick up feet, walk ratio that is time during which foal walks under constraint/total time measured during leading and number of defensive reactions). Two parameters (time to fit a halter and walk ratio) were still lower in handled foals than in non-handled foals 3 months later and only one 6 months later (walk ratio). One year later there was no difference between groups. In addition, there was no effect of handling on reactivity at any time of testing or in any of the tests (reaction to isolation from conspecifics, presence of a human, presence of a novel object and to a surprise effect). Finally, neonatal handling did not improve the spatial or discriminative learning abilities measured at 14 months of age. To conclude, the effects of neonatal handling are only temporary.
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McGreevy, P. D., & Rogers, L. J. (2005). Motor and sensory laterality in thoroughbred horses. Appl. Anim. Behav. Sci., 92(4), 337–352.
Abstract: We investigated lateralisation in horses because it is likely to be important in training and athletic performance. Thoroughbred horses (n = 106) were observed every 60 s for 2 h, when they were at pasture, and the position of the forelimbs in relation to one another was recorded. There was a population bias skewed to standing with the left forelimb advanced over the right (i.e. directional lateralisation). Using the first 50 observations, the distribution of preferences was 43 significantly left, 10 significantly right with 53 being non-significant (i.e. ambidextextrous). The strength of motor bias increased with age, suggesting maturation or an influence of training. The horses were also presented with an olfactory stimulus (stallion faeces) to score the tendency to use one nostril rather than the other. A significant preference to use the right nostril first was shown in horses under 4 years of age (n = 61) but not in older horses. Of the 157 horses tested for nostril bias, 76 had been assessed for motor bias and so were used for analysis of the relationship between laterality in the two modalities. There was no significant relationship between direction of foreleg motor bias and first nostril used, total number of inhalations or laterality index of nostril use. The absence of a correlation between laterality of nostril use and motor bias indicates that lateralisation of the equine brain occurs on at least two levels of neural organisation--sensory and motor--a finding that is consistent with other examples of lateralisation in species that have been examined in more detail.
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Dumont, B., Boissy, A., Achard, C., Sibbald, A. M., & Erhard, H. W. (2005). Consistency of animal order in spontaneous group movements allows the measurement of leadership in a group of grazing heifers. Appl. Anim. Behav. Sci., 95(1-2), 55–66.
Abstract: The term `leadership' has been used in several different senses, resulting in very different ways of identifying leaders and apparently inconsistent conclusions on how leadership is determined in herbivores. We therefore propose the following definitions: (i) a leader is the individual that is consistently the one who initiates long-distance, spontaneous group movements toward a new feeding site and (ii) long-distance spontaneous group movements are movements which happen when an animal changes activity and location and is immediately followed by a similar change in activity and location by other members of the group. Using these definitions, we tested for consistency of movement order across time and situation within a group of fifteen 2-year-old heifers. We found that the same individual was recorded as the very first animal in 48% of movements toward a new feeding site and could therefore be identified as the `leader'. We also showed that movement order when the animals entered an experimental plot, or progressed slowly through the field during a grazing bout, did not produce the same result. This method, which enables us to identify leaders in groups of animals at pasture, should improve our knowledge of how leadership is determined in grazing herbivores.
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Pompilio, L., & Kacelnik, A. (2005). State-dependent learning and suboptimal choice: when starlings prefer long over short delays to food. Anim. Behav., 70(3), 571–578.
Abstract: Recent studies have used labels such as `work ethics', `sunk costs' and `state-dependent preferences' for apparent anomalies in animals' choices. They suggest that preference between options relates to the options' history, rather than depending exclusively on the expected payoffs. For instance, European starlings, Sturnus vulgaris, trained to obtain identical food rewards from two sources while in two levels of hunger preferred the food source previously associated with higher hunger, regardless of the birds' state at the time of testing. We extended this experimentally and theoretically by studying starlings choosing between sources that differed not only in history but also in the objective properties (delay until reward) of the payoffs they delivered. Two options (PF and H) were initially presented in single-option sessions when subjects were, respectively, prefed or hungry. While option PF offered a delay until reward of 10 s in all treatments, option H delivered delays of 10, 12.5, 15 and 17.5 s in four treatments. When training was completed, we tested preference between the options. When delays in both options were equal (10 s), the birds strongly preferred H. When delay in H was 17.5 s, the birds were indifferent, with intermediate results for intermediate treatments. Preference was not mediated by disrupted knowledge of the delays. Thus, preferences were driven by past state-dependent gains, rather than by the joint effect of the birds' state at the time of choice and knowledge of the absolute properties of each alternative, as assumed in state-dependent, path-independent models of optimal choice.
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Fernández-Juricic, E., Smith, R., & Kacelnik, A. (2005). Increasing the costs of conspecific scanning in socially foraging starlings affects vigilance and foraging behaviour. Anim. Behav., 69(1), 73–81.
Abstract: Social foragers receive and use information both about companions (social information) and about events external to the group, such as presence of potential predators. Scanning behaviour is often incorporated in theoretical models using simplifying assumptions in relation to the trade-off in information gathering between body postures (head-up versus head-down); however, some avian visual systems may allow individuals to scan in both body postures. We studied these issues experimentally, using starlings, Sturnus vulgaris, foraging in enclosures on natural fields. We varied the availability of information from conspecifics by placing visual barriers that blocked the view when the subjects were in head-down position and by manipulating the distance between group members. We found that as social information was reduced, starlings spent more time scanning (on and off the ground) and head-up scanning was mainly oriented towards conspecifics. The visual-obstruction effects imply that some information about conspecifics is normally gathered while starlings are foraging head-down. Neighbour distance and visual obstruction negatively affected food-searching rates and intake rates in two ways: (1) the effect of obstruction was mediated mostly through time competition between foraging and scanning on the ground, and (2) the effect of distance was due to a reduction in the rate of prey returns per searching effort while the birds were head-down. We conclude that the head-up posture is only one component of scanning, that the effects of head-down scanning should also be considered in species with ample visual fields, and that scanning in starlings is strongly connected to monitoring other flock members.
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Robbins, M. M., Robbins, A. M., Gerald-Steklis, N., & Steklis, H. D. (2005). Long-term dominance relationships in female mountain gorillas: strength, stability and determinants of rank. Behaviour, 142(6), 779–809.
Abstract: A common practice in studies of social animals is to rank individuals according to dominance status, which has been shown to influence access to limited resources and stability of social relationships, and may in turn correlate with reproductive success. According to the socioecological model for primates, most female dominance relationships are either nepotistic or virtually undetectable (egalitarian), with nepotistic species being philopatric, and dispersing females being egalitarian. Female mountain gorillas (Gorilla beringei beringei) disperse, and they have been characterized as being egalitarian, but previous studies have not examined their dominance relationships from a long-term perspective. We evaluated 15 matrices of displacement/supplantation interactions that spanned 30 years of observations in the Virunga Volcanoes region, and included 51 female mountain gorillas in six groups. Only 4% of displacements were directed against higher ranking females, and when matrices had less than 5% unknown dyads, linearity indices were consistently greater than 0.95. Therefore, previous results suggesting undetectable dominance relationships may have reflected an insufficient quantity of data for this species, rather than actual nonlinearity in its hierarchies. Dominance depended on age and group tenure rather than nepotism, yet some females maintained a high ranking for most of adulthood (15-25 years). Most rank shifts occurred through changes in group composition, rather than switches in established relationships. These results fit within growing evidence for linear individualistic hierarchies in some primates, often coupled with dispersal, as commonly found in ungulates. In light of these results, we propose that the dominance relationships of female mountain gorilla are best characterized as “Dispersal-Individualistic” instead of the previously suggested “Dispersal-Egalitarian”.
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Stevens, J., Vervaecke, H., de Vries, H., & Van Elsacker, L. (2005). The influence of the steepness of dominance hierarchies on reciprocity and interchange in captive groups of bonobos (Pan paniscus). Behaviour, 142(7), 941–960.
Abstract: Biological market models explain variability in reciprocity and interchange between groups. In groups with a shallow dominance gradient, grooming will be mostly exchanged for itself (i.e. exchange will occur). In groups with steep dominance hierarchies, interchange is expected: individuals will groom higher ranking individuals to get access to limited resources or commodities such as support in conflicts, and grooming will be traded for these commodities. We examine patterns of reciprocity in grooming and support, and of interchange of grooming for support or for tolerance in six captive groups of bonobos. We test whether differences between groups in patterns of reciprocity and interchange can be attributed to differences in a measure of steepness of dominance hierarchies, which is based on dyadic agonistic interactions. We found that grooming was reciprocal in some, but not all groups. Support was highly reciprocal, but this was a side effect of dominance in most groups. Interchange between grooming and support was observed in some groups. Corroborating earlier findings, this was a side effect of individuals preferring high ranking individuals as grooming and support partners, possibly because these high-ranking individuals provide more efficient support in conflicts. There was no evidence for interchange of grooming for tolerance. Variability in grooming reciprocity was explained by differences in steepness of dominance hierarchies, as predicted by the biological market models. In groups with a shallow dominance hierarchy, grooming was more reciprocal. This was not true for reciprocity in support. There was some evidence that individuals groomed dominants more frequently in groups with a steep dominance hierarchy. The variation in interchange relations between grooming and support did not depend on the steepness of dominance hierarchies. We suggest that grooming in itself is a valuable commodity in bonobos, especially under captive conditions, which can be exchanged reciprocally. Bonobos may interchange grooming for another value equivalent, with food sharing as a very likely candidate. This interchange effects seem more dependent on potential to monopolise food than on steepness of dominance hierarchies per se.
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Christensen, J. W., Keeling, L. J., & Nielsen, B. L. (2005). Responses of horses to novel visual, olfactory and auditory stimuli. Appl. Anim. Behav. Sci., 93(1-2), 53–65.
Abstract: Responses of horses towards fear-eliciting stimuli can have important consequences for both human and horse safety. This experiment was designed to investigate behavioural and heart rate (HR) responses of horses to novel visual, auditory and olfactory stimuli. Twenty-four 2-year-old, previously unhandled, stallions were habituated to receive a food reward from a container in a test arena. Each horse was exposed to three 2 min tests in a balanced design where in addition to the feed container, either a traffic cone (visual test), white noise (auditory test) or eucalyptus oil applied to the inside of the container (olfactory test) were used as the novel stimuli. Compared to the control, less time was spent eating during all tests. There was no difference in locomotion activity in the different test situations, but presentation of the novel visual and auditory stimuli elicited significantly increased HR responses in the horses, compared to their response to the arena without novel stimuli (control), whereas there was no increase in HR response to the olfactory stimulus. However, during the olfactory test, the horses had an increased number of eating bouts and became more vigilant towards their surroundings, whereas during the visual and auditory tests, more time was spent alert towards the stimulus. The horses also took significantly more steps backwards in response to the auditory test. The heart rate responses correlated between tests and reflect a non-differentiated activation of the sympathetic nervous system, while the behavioural responses were linked to the type of stimulus.
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Kruger, K., Gaillard, C., Stranzinger, G., & Rieder, S. (2005). Phylogenetic analysis and species allocation of individual equids using microsatellite data. Journal of Animal Breeding and Genetics, 122(s1), 78–86.
Abstract: Summary The taxonomic status of all equid species is not completely unravelled. This is of practical relevance for conservation initiatives of endangered, fragmented equid populations, such as the Asiatic wild asses (in particular Equus hemionus onager and E. hemionus kulan). In this study, a marker panel consisting of 31 microsatellite loci was used to assess species demarcation and phylogeny, as well as allocation of individuals (n = 120) to specific populations of origin (n = 11). Phylogenetic analysis revealed coalescence times comparable with those previously published from fossil records and mtDNA data. Using Bayesian approaches, it was possible to distinguish between the studied equids, although individual assignment levels varied. The observed results support the maintenance of separate captive conservation herds for E. hemionus onager and E. hemionus kulan. The first molecular genetic results for E. hemionus luteus remained contradictory, as they unexpectedly indicated a closer genetic relationship between E. hemionus luteus and E. kiang holderi compared with the other hemiones.
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Lamoot, I., Callebaut, J., Demeulenaere, E., Vandenberghe, C., & Hoffmann, M. (2005). Foraging behaviour of donkeys grazing in a coastal dune area in temperate climate conditions. Appl. Anim. Behav. Sci., 92(1-2), 93–112.
Abstract: A small herd of donkeys was introduced in a coastal dune reserve `Houtsaegerduinen' (ca. 80 ha) in Belgium, in order to slow down expansion of dominant grass and shrub species. The Houtsaegerduinen is a nutrient poor scrub-dominated dune system with a spatially heterogeneous vegetation pattern. Different aspects of the grazing behaviour (grazing time, bite rate, habitat use, diet composition) of the free-ranging donkeys are described and analysed. Behavioural data (of maximum six adult mares) were collected through continuous focal animal observation in three consecutive years (1998-2001). Temporal variation in grazing time, habitat use and diet composition was determined. During daylight, donkeys spent most of their time on grazing (56%). In all 3 years, grazing time was significantly shorter in summer (45% of their time), longest grazing times were achieved in spring (64%). In spring, the donkeys also achieved the highest bite rate (21.5 bites/min). The grassy habitat was preferred for foraging in all seasons, while the use of scrub and woodland was variable over time. Averaged over the four seasons, the general diet consisted for 80% of graminoids, 10% of forbs and 10% of woody plants. However, diet composition varied not only among seasons and years, but depended also on the foraged habitat type. We discuss the possible role of the donkeys in nature management.
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