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Waran, N. K., Clarke, N., & Farnworth, M. (2008). The effects of weaning on the domestic horse (Equus caballus). Appl. Anim. Behav. Sci., 110(1-2), 42–57.
Abstract: For free-living or feral horses weaning takes place naturally at around 8-9 months [Gill, E.L., 1988. Factors affecting body condition of New Forest Ponies. Ph.D. Thesis. Department of Biology, University of Southampton]. Some mares will continue to suckle their foal until shortly before the arrival of their next foal, gestation being approximately 342 days depending upon the breed of the horse [Ropiha, R.T., Mathews, G., Butterfield, R.M., 1969. The duration of pregnancy in Thoroughbred mares. Vet. Rec. 84, 552-555]. Under domestic conditions, weaning tends to take place earlier, typically between 4 and 6 months of age. The weaning process has been identified as associated with potential psychological, physical and nutritional stressors that are of welfare concern. Following a review of the literature it is evident that there is a need for detailed research into what should constitute best practice with respect to foal and mare welfare. In addition, there is a need to understand the potential long-term impact of weaning on, for example, trainability and later maternal behaviour, and whether the stresses associated with early weaning have detrimental effects on the performance horse. There is also a lack of clear information concerning the most frequently observed weaning practices and the reasons why certain weaning methods are chosen. Some variables should be closely managed during weaning in order to minimise stress responses. These include: early creep feeding to familiarise the young animal with the food it will be exposed to during weaning, feeding a high fibre diet and keeping the animal in extensive conditions using a gradual approach to weaning. However, we conclude that there may not be one best method for weaning, since the chosen method must take into account a number of factors including: available resources, the housing environment, the individual foal's stage of development, the strength of the mare-foal attachment, the foal's ability to cope with changes in social conditions and the ability of the horse owner to implement the chosen method. We do however suggest that the fewest stress responses appear to occur where foals are weaned gradually and allowed to have social contact either with other foals or with older horses.
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Lansade, L., Bouissou, M. - F., & Erhard, H. W. (2008). Reactivity to isolation and association with conspecifics: A temperament trait stable across time and situations. Appl. Anim. Behav. Sci., 106(2-4), 355–373.
Abstract: A temperament trait is generally defined as individual differences in behaviour that are present early in life and relatively stable across situations and over time. The aim of this study was to test the existence of a trait <<gregariousness>> in horses, by testing the stability across situations and over time of the responses to different social events. Sixty-six Welsh ponies and 44 Anglo-Arab horses were successively tested at 8 months and 1.5 years of age. Among them, 33 ponies and 21 horses were also tested at 2.5 years of age. They were submitted to four test situations: isolation and separation from, attraction towards and passing conspecifics. We carried out the analysis using each of four test groups as a unit (e.g. 33 Welsh ponies born in 2001, tested in isolation). Isolation and separation stood out as tests that showed a high consistency within test, across tests and across time. The most interesting behavioural parameter was the frequency of neighing, which was well correlated with other parameters measured in the same tests, such as defecation, locomotion and vigilance, as well as across the 3 years (e.g. for separation test: 0.41 < R < 0.61). Therefore, the behaviour of neighing observed in separation or isolation tests as early as 8 months of age appears to be a good indicator of similar behaviour in similar situations later in life, but also of other behaviours which can render the horse difficult to use. No parameter recorded during the attraction test presented stability across situations and time: the reactions to this test were not the expression of a stable characteristic of the individual and did not reflect the same characteristic as the three other tests. Of the different parameters recorded during the passing conspecifics test, the time to cross the arrival line near conspecifics showed good stability across years (0.35 < R < 0.68). This parameter was also correlated with many others recorded during the same test, and also, to a certain extent, to the frequency of neighing in the isolation and separation tests. This stability across responses expressed in various social contexts, and this stability over time, from 8 months to 2.5 years of age suggest the existence of a trait of gregariousness in the horse. From a practical point of view, that means it is possible to estimate the level of gregariousness of a horse as early as 8 months of age. Furthermore, additional analysis shows that gregariousness decreases with age.
Keywords: Horse; Equus caballus; Temperament; Trait; Isolation; Conspecific association; Stability
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Schmied, C., Waiblinger, S., Scharl, T., Leisch, F., & Boivin, X. (2008). Stroking of different body regions by a human: Effects on behaviour and heart rate of dairy cows. Appl. Anim. Behav. Sci., 109(1), 25–38.
Abstract: From observations of intra-specific social grooming in cattle and studies on human stroking in other species, we hypothesised that cows' reactions to human stroking differ depending on the body regions being stroked. Moreover, we tested, whether cows `reactions to stroking change with the animals' experience of stroking. Sixty dairy cows were stroked in three different body regions, i.e. the withers, W, neck ventral, NV (both licked often in social grooming) and the lateral chest, LC (licked rarely), in a balanced order during 10-min sessions. Behavioural reactions and heart rate during stroking as well as reactions to the human just after stroking were recorded. Two test sessions were carried out with 3 weeks of treatment in-between. During this period, the cows were randomly allocated to four treatment groups: three groups received 5 min of daily stroking in either W, NV or LC and the last one (control group) was exposed to simple human presence. During stroking W and NV, cows showed longer neck stretching and ear hanging than during stroking LC (P < 0.001). Moreover, ear hanging was shown longer when W was stroked as compared to NV (P < 0.001), but neck stretching was observed longer during stroking NV as compared to W only after the treatment period (P < 0.01). In the first test session, more animals showed head shaking and head throwing during stroking W and NV than LC (P < 0.01), whereas in the second test session these behaviours were observed only in few animals. In the first test session heart rate was higher during stroking W than the other body regions (P <= 0.01). In the second test session, the lowest heart rate was found during stroking NV (P < 0.05). Finally, contact with the experimenter following stroking differed for the three body regions stroked in both test sessions (first: P = 0.06, second: P < 0.01); contact was shortest for LC. When comparing both test sessions, stretching the neck and ear hanging increased (P < 0.001), while head shaking and head throwing decreased (P < 0.05); there was no difference for heart rate and contact with the experimenter. The four treatments had no influence on reactions to stroking. To conclude, stroking of body regions often licked during social grooming led to more responses than stroking the one licked rarely. Some reactions, such as stretching the neck, are also observed during social licking. This suggests that cows may in part perceive human stroking of body regions often licked similarly to social licking. This knowledge could be of interest for an improvement in quality of human-cattle interactions.
Keywords: Cattle; Tactile stimulation; Human-animal relationship; Social licking
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Newberry, R. C., & Swanson, J. C. (2008). Implications of breaking mother-young social bonds. Appl. Anim. Behav. Sci., 110(1-2), 3–23.
Abstract: Whereas mammalian mothers and young may retain long-term social affiliations in nature, the management of animals in captivity typically dictates that offspring are abruptly and permanently separated from their mothers at a relatively early age, often prior to the time of natural weaning. For animal breeders, this strategy can enhance the yield of offspring from a breeding population. Morbidity and mortality can also precipitate severance of mother-young bonds. Although it is recognized that early weaning provides nutritional challenges for the young, relatively little attention has been paid to the psychological consequences and long-term impacts of breaking the mother-young bond in non-human mammals. Furthermore, whereas great strides are being made in our understanding of the neurobiological and genetic underpinnings of social bonding, the mechanisms underlying the process of detachment following establishment of a mother-young bond remain relatively unexplored, although parallels can be drawn with processes involved in withdrawal from addictive substances. In this review, we outline mechanisms involved in social bonding. We consider the diversity in extent and duration of mother-young attachment across mammalian lineages and implications for predicting the outcome of severing ties between mothers and young at different times post-partum. We identify characteristics signalling emotional distress resulting from separation of mothers and young and discuss strategies for mitigating separation-induced distress. These include postponement of separation, ensuring high-quality maternal care of young prior to separation, providing bonded individuals with opportunities to separate voluntarily for brief periods prior to permanent separation, use of anti-suck devices prior to separation, allowing a period of partial (fence line) contact prior to full separation, providing substitutes for stimuli previously exchanged between mother and young, providing social buffers, gradual introduction to new housing arrangements, and pharmacological intervention. Areas for future research are proposed, including the use of functional neuroimaging technologies and functional genomics approaches, in combination with behavioural assessments of reinstatement motivation, individual recognition memory and long-term consequences of early separation, to shed further light on the nature of mother-young bonding and detachment in animals.
Keywords: Animal welfare; Emotion; Separation; Social attachment; Weaning
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Smith, J. E., Kolowski, J. M., Graham, K. E., Dawes, S. E., & Holekamp, K. E. (2008). Social and ecological determinants of fission-fusion dynamics in the spotted hyaena. Anim. Behav., 76(3), 619–636.
Abstract: Theory predicts that individuals living in fission-fusion societies, in which group members frequently change subgroups, should modify grouping patterns in response to varying social and environmental conditions. Spotted hyaenas, Crocuta crocuta, are long-lived carnivores that reside in permanent social groups called clans. Clans are complex, fission-fusion societies in which individual members travel, rest and forage in subgroups that frequently change composition. We studied two clans in Kenya to provide the first detailed description of fission-fusion dynamics in this species. Because social and ecological circumstances can influence the cohesiveness of animal societies, we evaluated the extent to which specific circumstances promote the formation of subgroups of various sizes. We found that cooperative defence of shared resources during interclan competition and protection from lions were cohesive forces that promoted formation of large subgroups. We also tested hypotheses suggesting factors limiting subgroup size. Mothers with small cubs avoided conspecifics, thereby reducing infanticide risk. Victims of aggression either reconciled fights or separated from former opponents to reduce the immediate costs of escalated aggression in the absence of food. As predicted by the ecological constraints hypothesis, hyaenas adjusted their grouping patterns over both short and long time scales in response to feeding competition. Crocuta were most gregarious during periods of abundant prey, joined clanmates at ephemeral kills in numbers that correlated with the energetic value of the prey and gained the most energy when foraging alone because cooperative hunting attracted numerous competitors. Overall, our findings indicate that resource limitation constrains grouping in this species.
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Patris, B., Perrier, G., Schaal, B., & Coureaud, G. (2008). Early development of filial preferences in the rabbit: implications of nursing- and pheromone-induced odour learning? Anim. Behav., 76(2), 305–314.
Abstract: Newborn rabbits, Oryctolagus cuniculus, discriminate between different categories of adult conspecifics on the basis of their abdominal odour cues. Whether these cues can support the development of filial preferences has not been adequately tested. Using a two-choice paradigm, we assessed the ability of 3-8-day-old pups to orient selectively to the mother versus an unfamiliar female, either spontaneously or after odour conditioning. In experiment 1, nonconditioned pups roamed indifferently over the mother and an unfamiliar female. In experiment 2, pups conditioned to a neutral odorant while nursing or with the mammary pheromone became attracted by the odorant. In experiment 3, pups that had learned the odorant while nursing oriented for longer to any female carrying it, but the unscented mother and a scented unfamiliar female were equally attractive. Finally, in experiment 4, pups that had learned the odorant paired with the mammary pheromone showed a preference for their scented mother, but not systematically for a scented unfamiliar female; furthermore, they were equally attracted by the unscented mother and a scented unfamiliar female. In sum, pups did not spontaneously evince an olfactory preference for the mother when opposed to an unfamiliar female, although they seemed able to detect individual maternal odours. In fact, they appeared to react to both species-specific cues and individual cues that they had learned, and their responses depended on their degree of familiarity with the cues and on the context. The mammary pheromone by itself might act as both a releasing and a reinforcing signal in these early socially oriented behaviours.
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Wey, T., Blumstein, D. T., Shen, W., & Jordán, F. (2008). Social network analysis of animal behaviour: a promising tool for the study of sociality. Anim. Behav., 75(2), 333–344.
Abstract: Social animals live and interact together, forming complex relationships and social structure. These relationships can have important fitness consequences, but most studies do not explicitly measure those relationships. An approach that explicitly measures relationships will further our understanding of social complexity and the consequences of both direct and indirect interactions. Social network analysis is the study of social groups as networks of nodes connected by social ties. This approach examines individuals and groups in the context of relationships between group members. Application of social network analysis to animal behaviour can advance the field by identifying and quantifying specific attributes of social relationships, many of which are not captured by more common measures of sociality, such as group size. Sophisticated methods for network construction and analysis exist in other fields, but until recently, have seen relatively little application to animal systems. We present a brief history of social network analysis, a description of basic concepts and previous applications to animal behaviour. We then highlight relevance and constraints of some network measures, including results from an original study of the effect of sampling on network parameter estimates, and we end with promising directions for research. By doing so, we provide a prospective overview of social network analysis' general utility for the study of animal social behaviour.
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Whitehead, H. (2008). Precision and power in the analysis of social structure using associations. Anim. Behav., 75(3), 1093–1099.
Abstract: I develop guidelines for assessing the precision and power of statistical techniques that are frequently used to study nonhuman social systems using observed dyadic associations. Association indexes estimate the proportion of time that two individuals are associated. Binomial approximation and nonparametric bootstrap methods produce similar estimates of the precision of association indexes. For a mid-range (0.4-0.9) association index to have a standard error of less than 0.1 requires about 15 observations of the pair associated, and for it to be less than 0.05, this rises to 50 observations. The coefficient of variation among dyads of the proportion of time that pairs of individuals are actually associated describes social differentiation (S), and this may be estimated from association data using maximum likelihood. With a poorly differentiated population (S~0.2), a data set needs about five observed associations per dyad to achieve a correlation between true and estimated association indexes of r=~0.4. It requires about 10 times as much data to achieve a representation with r=~0.8. Permutation tests usually reject the null hypothesis that individuals have no preferred associates when S2H>5, where H is the mean number of observed associations per individual. Thus most situations require substantial numbers of observations of associations to give useful portrayals of social systems, and sparse association data inform only when social differentiation is high.
Keywords: association; precision; social structure; statistical power
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Palagi, E., Antonacci, D., & Norscia, I. (2008). Peacemaking on treetops: first evidence of reconciliation from a wild prosimian (Propithecus verreauxi). Anim. Behav., 76(3), 737–747.
Abstract: Reconciliation is defined as the first postconflict affinitive contact between former opponents. While reconciliation in anthropoid primates has been widely investigated, few studies have focused on postconflict mechanisms in prosimians, and only in captivity. Unlike anthropoids, Malagasy prosimians show female dominance, lack of sexual dimorphism and seasonal breeding. However, they share features with anthropoids such as cohesive societies, female philopatry and individual recognition. Comparing social prosimians with anthropoids is crucial for understanding the evolution of reconciliation dynamics. Here we present the first study on reconciliation in a wild prosimian. We focused on the Propithecus verreauxi (sifaka) of the Berenty forest (southern Madagascar). We examined postconflict behaviour in the light of theoretical expectations based on potential costs and benefits of the individuals involved. Our results indicate that P. verreauxi can evaluate possible risks and benefits of engaging in postconflict reunions. Victims were most likely to interact affinitively with the aggressor after low-intensity aggression. Moreover, only the conflicts occurring outside the feeding context were reconciled. Such results are consonant with the fact that, in P. verreauxi, social dominance is translated more into feeding priority than into a framework of despotic relationships. In agreement with the valuable relationship hypothesis, P. verreauxi were more likely to reconcile with valuable partners: reconciliation preferentially occurred between subordinates and top-ranking individuals, and between animals sharing good relationships (high levels of affinitive behaviours). Over the short term, reconciliation in P. verreauxi seems to have an important role in reducing the probability of further attacks by the aggressor.
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
Keywords: Equus caballus; feral horse; maternal investment; play
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