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Smith, L.A.; Wells, K.L.; Marion, G.; Swain, D.L.; Hutchings, M.R. |
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Title |
Effects of group composition on the grazing behaviour of herbivores |
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Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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In Press, Corrected Proof |
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grazing; group composition; herbivore; individual behaviour; physiological state |
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Animal behaviour is often a function of the animal's physiological state. Groups of animals will often contain individuals with a range of physiological states and the grazing behaviour of herbivores is affected by their physiological state. This study compared the grazing decisions of animals in groups of single and mixed physiological states. Using a grazing model that simulated individual herbivore behaviour in relation to environmental distributions of forage resource (grass) and parasites (faeces), we tested the hypothesis that an animal's level of parasite exposure via the faecal-oral route is affected by the composition of physiological states in the group. Four physiological states were considered: parasite-naïve, parasitized, lactating and parasite-immune animals. Baseline parasite exposure levels for each state were generated by simulating single-state groups and were compared to simulations of each of the six two-state combinations. In single-state groups parasitized animals had the least and lactating animals had the greatest levels of parasite exposure. When co-grazing with lactating animals, parasitized, immune and naïve animals increased their parasite exposure, relative to single-state groups. When co-grazing with parasitized animals, lactating, immune and naïve animals reduced their parasite exposure, relative to single-state groups. There was no difference in parasite exposure of the immune or naïve animals co-grazing together when compared to the single-state groups. These results highlight the need to recognize the impact of the individual when studying group-living animals. |
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0003-3472 |
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Equine Behaviour @ team @ |
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5155 |
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Author |
Seyfarth, R.M. |
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Title |
A model of social grooming among adult female monkeys |
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Journal Article |
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Year |
1977 |
Publication |
Journal of Theoretical Biology |
Abbreviated Journal |
J. Theor. Biol. |
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Volume |
65 |
Issue |
4 |
Pages |
671-698 |
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Keywords |
Animals; Behavior, Animal; Female; *Grooming; Haplorhini/*physiology; *Models, Biological; Reproduction; Social Dominance; Time Factors |
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Abstract |
Grooming networks among adult female monkeys exhibit two similar features across a number of different species. High-ranking animals receive more grooming than others, and the majority of grooming occurs between females of adjacent rank. A theoretical model which duplicates these features is presented, and the properties of the model are used to explain the possible causation and function of female grooming behaviour. The model illustrates how relatively simple principles governing the behaviour of individuals may be used to explain more complex aspects of the social structure of non-human primate groups. |
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0022-5193 |
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PMID:406485 |
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Equine Behaviour @ team @ |
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5259 |
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Newton-Fisher, N.E.; Lee, P.C. |
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Title |
Grooming reciprocity in wild male chimpanzees |
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Journal Article |
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Year |
2011 |
Publication |
Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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81 |
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2 |
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439-446 |
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Keywords |
altruism; biological market theory; Budongo; chimpanzee; grooming; Pan troglodytes |
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Understanding cooperation between unrelated individuals remains a central problem in animal behaviour; evolutionary mechanisms are debated, and the importance of reciprocity has been questioned. Biological market theory makes specific predictions about the occurrence of reciprocity in social groups; applied to the social grooming of mammals, it predicts reciprocity in the absence of other benefits for which grooming can be exchanged. Considerable effort has been made to test this grooming trade model in nonhuman primates; such studies show mixed results, but may be confounded by kin effects. We examined patterns of reciprocity within and across bouts, and tested predictions of the grooming trade model, among wild male chimpanzees, Pan troglodytes: a system with negligible kin effects. In accord with the model's expectations, we found that some grooming was directed by lower- to higher-ranked individuals, and that, on average, higher-ranked individuals groomed more reciprocally. We found no support, however, for a prediction that more reciprocity should occur between individuals close in rank. For most dyads, reciprocity of effort occurred through unbalanced participation in grooming bouts, but reciprocity varied considerably between dyads and only a small proportion showed strongly reciprocal grooming. Despite this, each male had at least one reciprocal grooming relationship. In bouts where both individuals groomed, effort was matched through mutual grooming, not alternating roles. Our results provide mixed support for the current grooming trade, biological market model, and suggest that it needs to incorporate risks of currency inflation and cheating for species where reciprocity can be achieved through repeated dyadic interactions. |
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0003-3472 |
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Equine Behaviour @ team @ |
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5329 |
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Author |
Lee, R.D. |
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Title |
Rethinking the evolutionary theory of aging: transfers, not births, shape senescence in social species |
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Journal Article |
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Year |
2003 |
Publication |
Proceedings of the National Academy of Sciences of the United States of America |
Abbreviated Journal |
Proc Natl Acad Sci U S A |
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100 |
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16 |
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9637-9642 |
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Keywords |
Adaptation, Physiological; *Aging; Animals; *Biological Evolution; Demography; Economics; Environment; Fertility; Humans; Life Expectancy; Longevity; Models, Theoretical; Parturition; Population Dynamics; Population Growth; Reproduction |
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The classic evolutionary theory of aging explains why mortality rises with age: as individuals grow older, less lifetime fertility remains, so continued survival contributes less to reproductive fitness. However, successful reproduction often involves intergenerational transfers as well as fertility. In the formal theory offered here, age-specific selective pressure on mortality depends on a weighted average of remaining fertility (the classic effect) and remaining intergenerational transfers to be made to others. For species at the optimal quantity-investment tradeoff for offspring, only the transfer effect shapes mortality, explaining postreproductive survival and why juvenile mortality declines with age. It also explains the evolution of lower fertility, longer life, and increased investments in offspring. |
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Department of Demography, University of California, 2232 Piedmont Avenue, Berkeley, CA 94720-2120, USA. rlee@demog.berkeley.edu |
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0027-8424 |
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PMID:12878733 |
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no |
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Call Number |
Equine Behaviour @ team @ |
Serial |
5465 |
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Author |
Sloet van Oldruitenborgh-Oosterbaan, M.M.; Blok, M.B.; Begeman, L.; Kamphuis, M.C.D.; Lameris, M.C.; Spierenburg, A.J.; Lashley, M.J.J.O. |
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Title |
Workload and stress in horses: comparison in horses ridden deep and round ('rollkur') with a draw rein and horses ridden in a natural frame with only light rein contact |
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Journal Article |
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Year |
2006 |
Publication |
Tijdschrift Voor Diergeneeskunde |
Abbreviated Journal |
Tijdschr Diergeneeskd |
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131 |
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5 |
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152-157 |
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Animal Husbandry/methods; Animals; Biomechanics; Blood Glucose/analysis; Female; Heart Rate/physiology; Hematocrit/veterinary; Horses/blood/*physiology; Hydrocortisone/blood; Lactic Acid/blood; Physical Conditioning, Animal/adverse effects/*physiology; Stress, Physiological/blood/etiology/veterinary |
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'Rollkur' or 'overbending' is the low and deep riding of a dressage horse during training or warming up. Lately, this technique has been criticized, and not necessarily objectively, on welfare grounds. To be able to evaluate these criticisms, more needs to be known about the workload and stress of horses being ridden 'rollkur'. The aim of the present study was to compare the workload of eight riding-school horses when being ridden deep and round with a draw rein ('rollkur') and when being ridden in a natural frame with only light rein contact ('free'). Workload (as measured by heart rate and blood lactate concentration) was slightly higher when horses were ridden 'rollkur' than when they were ridden 'free'. There were no differences in packed cell volume, or glucose and cortisol concentrations. No signs of uneasiness or stress could be determined when the horses were ridden 'rollkur'. Subjectively, all horses improved their way of moving during 'rollkur' and were more responsive to their rider. |
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Department of Equine Sciences, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 12, 3584 CM Utrecht, The Netherlands. m.sloet@vet.uu.nl |
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0040-7453 |
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PMID:16532786 |
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Equine Behaviour @ team @ |
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5638 |
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Author |
Palme, R. |
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Title |
Non-invasive measurement of glucocorticoids: Advances and problems |
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Journal Article |
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Year |
2019 |
Publication |
Physiology & Behavior |
Abbreviated Journal |
Physiol. Behav. |
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199 |
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229-243 |
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Glucocorticoids; Non-invasive; Faecal cortisol/corticosterone metabolites; Immunoassays; Physiological/biological validation |
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Glucocorticoids (GCs; i.e. cortisol/corticosterone) are a central component of the stress response and thus their measurement is frequently used to evaluate the impact of stressful situations. Their metabolites from faeces of various animal species are more and more taken as a non-invasive aid to assess GC release and thus adrenocortical activity. The current literature review includes an extensive collection (1327 papers) and evaluation (see also Supplementary Tables) of the literature on faecal cortisol/corticosterone metabolite (FCM) analysis published to date. It aims at giving reference for researchers interested in implementing FCM analysis into their study or seeking to improve such methods by providing background knowledge on GC metabolism and excretion, conveying insights into methodological issues and stating caveats of FCM analysis and by highlighting prerequisites for and some examples of a successful application of such methods. Collecting faecal samples and analysing FCMs may appear simple and straightforward, but researchers have to select and apply methods correctly. They also need to be aware of the many pitfalls and potentially confounding factors and, last but not least, have to carefully interpret results. Applied properly, measurement of FCMs is a powerful non-invasive tool in a variety of research areas, such as (stress) biology, ethology, ecology, animal conservation and welfare, but also biomedicine. |
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0031-9384 |
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Palme2019_attachment.pdf |
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Equine Behaviour @ team @ |
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6517 |
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Bussières, G.; Jacques, C.; Lainay, O.; Beauchamp, G.; Leblond, A.; Cadoré, J.-L.; Desmaizières, L.-M.; Cuvelliez, S.G.; Troncy, E. |
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Development of a composite orthopaedic pain scale in horses |
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2008 |
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Research in Veterinary Science |
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85 |
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2 |
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294-306 |
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Horse; Acute orthopaedic pain; Experimental model; Behaviour; Physiological parameters; Validation |
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This study addresses development and validation of a composite multifactorial pain scale (CPS) in an experimental equine model of acute orthopaedic pain. Eighteen horses were allocated to control (sedation with/without epidural analgesia – mixture of morphine, ropivacaine, detomidine and ketamine) and experimental groups: amphotericin-B injection in the tarsocrural joint induced pain and analgesia was either i.v. phenylbutazone administered post-induction of synovitis, or pre-emptive epidural mixture, or a pre-emptive combination of the 2. Inter- and intra-observer reproducibility was good (0.8<K<1). The key specific and sensitive behavioural indices were response to palpation of the painful area, posture, and, of lesser value, pawing on the floor, kicking at abdomen and head movement. Of particular interest was the statistical correlation observed between the CPS and both non-invasive blood pressure (P<0.0001) and blood cortisol (P<0.002). This study established the value of some behavioural and physiological criteria in determining equine orthopaedic pain intensity and clearly demonstrated that pre-emptive, multimodal analgesia provided better management than the two other protocols tested. |
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0034-5288 |
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Equine Behaviour @ team @ |
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6707 |
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Massen, J.; Sterck, E.; de Vos, H. |
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Close social associations in animals and humans: functions and mechanisms of friendship |
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2010 |
Publication |
Behaviour |
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147 |
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11 |
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1379 |
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Both humans and group-living animals associate and behave affiliatively more with some individuals than others. Human friendship has long been acknowledged, and recently scientists studying animal behaviour have started using the term friendship for close social associates in animals. Yet, while biologists describe friends as social tools to enhance fitness, social scientists describe human friendship as unconditional. We investigate whether these different descriptions reflect true differences in human friendship and animal close social associations or are a by-product of different research approaches: namely social scientists focussing on proximate and biologists on ultimate explanations. We first stress the importance of similar measures to determine close social associations, thereafter examine their ultimate benefits and proximate motivations, and discuss the latest findings on the central-neural regulation of social bonds. We conclude that both human friendship and animal close social associations are ultimately beneficial. On the proximate level, motivations for friendship in humans and for close social associations in animals are not necessarily based on benefits and are often unconditional. Moreover, humans share with many animals a similar physiological basis of sociality. Therefore, biologists and social scientist describe the same phenomenon, and the use of the term friendship for animals seems justified. |
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Both humans and group-living animals associate and behave affiliatively more with some individuals than others. Human friendship has long been acknowledged, and recently scientists studying animal behaviour have started using the term friendship for close social associates in animals. Yet, while biologists describe friends as social tools to enhance fitness, social scientists describe human friendship as unconditional. We investigate whether these different descriptions reflect true differences in human friendship and animal close social associations or are a by-product of different research approaches: namely social scientists focussing on proximate and biologists on ultimate explanations. We first stress the importance of similar measures to determine close social associations, thereafter examine their ultimate benefits and proximate motivations, and discuss the latest findings on the central-neural regulation of social bonds. We conclude that both human friendship and animal close social associations are ultimately beneficial. On the proximate level, motivations for friendship in humans and for close social associations in animals are not necessarily based on benefits and are often unconditional. Moreover, humans share with many animals a similar physiological basis of sociality. Therefore, biologists and social scientist describe the same phenomenon, and the use of the term friendship for animals seems justified. |
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Equine Behaviour @ team @ |
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5813 |
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Author |
VanDierendonck, M.C. |
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Social relationships in a group of horses without a mature stallion |
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2006 |
Publication |
The Importance of Social Relationships in Horses |
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Chapter 4 |
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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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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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Universiteit Utrecht |
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Equine Behaviour @ team @ |
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2369 |
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Author |
VanDierendonck, M.C. |
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Diff erences in social behaviour between late pregnant, post-partum and barren mares in a herd of Icelandic horses |
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2006 |
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The Importance of Social Relationships in Horses |
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Chapter 5 |
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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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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. |
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Ph.D. thesis |
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Universiteit Utrecht |
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Equine Behaviour @ team @ |
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2370 |
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