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Weckerly, F. W. (1999). Social bonding and aggression in female Roosevelt elk. Can J Zool, 77(9), 1379–1384.
Abstract: Abstract: The relationship between degree of social bonding (extent of association among individuals) and level of aggression in ruminants is unclear. I examined social bonding and aggression in three groups of female Roosevelt elk (Cervus elaphus roosevelti) over 2 years. I hypothesized that when animals are socially bonded, bouts of aggression will be won by the individual initiating the aggression, occur quickly, and involve little physical contact, and the level of aggression does not correlate with group size. The degree of social bonding was high among individuals in all groups. Dyads of known individuals were together >80% of the time. A permutation analysis indicated that groups with the observed sizes had <0.001 chance of random association, except on one occasion when the probability was 0.72 for one group. Using focal-animal sampling, aggressive interactions were won 72% of the time by the initiator, occurred quickly (<5 s), and involved little physical contact, and the level of aggression was not correlated with group size. The level of aggression was, however, significantly lower in one of the groups. This group may have had access to more abundant food resources than the other groups. Socially bonded elk conducted aggressive interactions in a fashion that did not disrupt social stability. Résumé : La relation entre le degré de liaison sociale (importance des associations entre individus) et l`agressivité n`est pas claire chez les ruminants. J`ai étudié les liaisons sociales et l`agressivité chez trois groupes de femelles du Cerf de Roosevelt (Cervus elaphus roosevelti) pendant 2 ans. J`ai posé en hypothèse que, chez les animaux liés socialement, la victoire devrait être emportée par l`individu qui entreprend l`agression, l`agression devrait être de courte durée, se faire avec peu de contacts physiques et la fréquence des agressions ne devrait pas être liée à la taille du groupe. Des paires d`individus passaient plus de 80% de leur temps ensemble. Une analyse des permutations a démontré que, chez les groupes des tailles observées, la probabilité d`une association aléatoire était de moins de 0,001, sauf en un cas où cette probabilité a été évaluée à 0,72 chez un groupe. Par échantillonnage directionnel, j`ai observé que les interactions agressives étaient gagnées par l`individu attaquant 72% du temps, étaient de courte durée (<5 s), se faisaient avec peu de contacts physiques et leur fréquence n`était pas reliée à la taille du groupe. Il y avait cependant moins d`agressivité chez l`un des groupes. Il se peut que ce groupe ait eu accès à plus de ressources alimentaires que les autres. Chez les cerfs liés par des liens sociaux, l`agressivité ne se manifeste pas de façon à déséquilibrer la stabilité sociale.
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Hogan, J. (2005). Causation: the study of behavioural mechanisms. Animal Biology (formerly Netherlands Journal of Zoology), 55(4), 323–341.
Abstract: This paper describes current work on the causal analysis of behaviour systems. It is noted that while causal work investigating the neural, hormonal, and genetic bases of behaviour is flourishing, work being conducted at a strictly behavioural level of analysis has declined greatly over the past 40 years. Nonetheless, most recent research on animal cognition and applied ethology is still being carried out at a behavioural level of analysis and examples of both types of research are presented: memory mechanisms of food-storing birds and decisions of spider-eating jumping spiders, as well as feather pecking in fowl and animal welfare issues, are all briefly discussed. Finally, I discuss the similarities between neural network modelling and early ethological models of motivation, and then show how a modern version of Lorenz's model of motivation can account for current research findings on dustbathing in chickens and sleep in humans. I conclude that valuable information can still be obtained by research at a behavioural level of analysis.
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Bolhuis, J. (2005). Function and mechanism in neuroecology: looking for clues. Animal Biology (formerly Netherlands Journal of Zoology), 55(4), 457–490.
Abstract: The four questions that Niko Tinbergen identified for behavioural biology ? evolution, function, development and causation ? are all important and should be studied in their own right. Recently, there has been a debate as to whether these four questions should be investigated separately or whether they should be integrated. Integration of the four questions has been attempted in novel research disciplines such as cognitive ecology, evolutionary psychology and neuroecology. Euan Macphail and I have criticised these integrative approaches, suggesting that they are fundamentally flawed as they confound function and mechanism. Investigating the function or evolutionary history of a behaviour or cognitive system is important and entirely legitimate. However, such investigations cannot provide us with answers to questions about the mechanisms underlying behaviour or cognition. At most, functional or evolutionary considerations can provide clues that may be useful for a causal analysis of the underlying mechanisms. However, these clues can be misleading and are often wrong, as is illustrated with examples from song learning and food storing in birds. After summarising the main issues in the neuroecology debate, I discuss some misunderstandings that were apparent in the responses to our critique, as well as some recent relevant data. Recent results do not support the neuroecological approach. Finally, I suggest that the way forward is a cautious and critical use of functional and evolutionary clues in the study of the mechanisms of behaviour.
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Reader, S. M. (2003). Innovation and social learning: individual variation and brain evolution. Anim. Biol. Leiden., 53(2), 147–158.
Abstract: This paper reviews behavioural, neurological and cognitive correlates of innovation at the individual, population and species level, focusing on birds and primates. Innovation, new or modified learned behaviour not previously found in the population, is the first stage in many instances of cultural transmission and may play an important role in the lives of animals with generalist or opportunistic lifestyles. Within-species, innovation is associated with low neophobia, high neophilia, and with high social learning propensities. Indices of innovatory propensities can be calculated for taxonomic groups by counting the frequency of reports of innovation in published literature. These innovation rate data provide a useful comparative measure for studies of behavioural flexibility and cognition. Innovation rate is positively correlated with the relative size of association areas in the brain, namely the hyperstriatum ventrale and neostriatum in birds, and the neocortex and striatum in primates. Innovation rate is also positively correlated with the reported variety of tool use, as well as interspecific differences in learning. Current evidence thus suggests similar patterns of cognitive evolution in primates and birds.
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VanDierendonck, M. C., de Vries, H., & Schilder, M. B. H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic orses in Captivity. Netherl. J. Zool., 45(3-4), 362–385.
Abstract: Th e applicability of the concept of dominance was investigated in a captive herd of  Icelandic
horses and  ponies of diff erent breeds. Eight out of  behaviours possibly related to dominance occurred frequently enough to be investigated in detail. For these eight agonistic behaviours the coverage, the unidirectionality in the exchange, and the degree of transitivity (Landau`s linearity index) were calculated. Four off ensive behaviours, together with avoidance, were suitable for further analysis with regard to dominance. Th e patterns of asymmetries with which these behaviours were exchanged were suffi ciently similar as to justify the application of the dominance concept and to construct a (nearly) linear dominance hierarchy. Th e rank order of the castrated stallions was completely linear, the hierarchy of the mares was almost completely linear. Th e results suggest that off ensive and defensive aggressive behaviours should be treated separately and that the concept of dominance is applicable. However, ritualized formal dominance signals between adult horses appear to be (almost) absent. Th e rank positions of the individuals were correlated with age and residency in the herd but not with height. Middle ranking horses tended to be more frequently in the close vicinity of another horse than high ranking or low ranking horses. Over and above this correlation at the individual level, it was found that pairs of horses close in rank to each other were more often also spatially close to each other. Being in oestrus did not infl uence the dominance relationships between mares. For castrated stallions the rank positions were correlated with the age at which they were castrated. Th is suggests that in male horses experience prior to neutering infl uences the behaviour afterwards. Keywords: Dominance; rank order; horses; Icelandic horses.
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VanDierendonck, M. C., de Vries, H., Schilder, M.B.H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic horses in captivity. Netherl. J. Zool., 45(3-4), 362–385.
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.
Keywords: Dominance; rank order; horses; Icelandic horses.
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Bennett Dk,. (1980). Stripes do not a zebra make, Part I: A cladistic analysis of Equus. Syst Zool, 29(2), 272–287.
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Smuts, M. M. S., & Penzhorn, B. L. (1988). Descriptions of antomical differences between skulls and mandibles of Equus zebra and E. burchelli from southern Africa. South African Journal of Zoology, 23((4)3), 328–336.
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Briefer, E. F., Mandel, R., Maigrot, A. - L., Briefer Freymond, S., Bachmann, I., & Hillmann, E. (2017). Perception of emotional valence in horse whinnies. Frontiers in Zoology, 14(1), 8.
Abstract: Non-human animals often produce different types of vocalisations in negative and positive contexts (i.e. different valence), similar to humans, in which crying is associated with negative emotions and laughter is associated with positive ones. However, some types of vocalisations (e.g. contact calls, human speech) can be produced in both negative and positive contexts, and changes in valence are only accompanied by slight structural differences. Although such acoustically graded signals associated with opposite valence have been highlighted in some species, it is not known if conspecifics discriminate them, and if contagion of emotional valence occurs as a result. We tested whether domestic horses perceive, and are affected by, the emotional valence of whinnies produced by both familiar and unfamiliar conspecifics. We measured physiological and behavioural reactions to whinnies recorded during emotionally negative (social separation) and positive (social reunion) situations.
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Kaiser, S., Hennessy, M. B., & Sachser, N. (2015). Domestication affects the structure, development and stability of biobehavioural profiles. Frontiers in Zoology, 12(1), 1–11.
Abstract: Domestication is an evolutionary process during which the biobehavioural profile (comprising e.g. social and emotional behaviour, cognitive abilities, as well as hormonal stress responses) is substantially reshaped. Using a comparative approach, and focusing mainly on the domestic and wild guinea pig, an established model system for the study of domestication, we review (a) how wild and domestic animals of the same species differ in behaviour, emotion, cognition, and hormonal stress responses, (b) during which phases of life differences in biobehavioural profiles emerge and (c) whether or not animal personalities exist in both the wild and domestic form. Concerning (a), typical changes with domestication include increased courtship, sociopositive and maternal behaviours as well as decreased aggression and attentive behaviour. In addition, domestic animals display more anxiety-like and less risk-taking and exploratory behaviour than the wild form and they show distinctly lower endocrine stress responsiveness. There are no indications, however, that domestic animals have diminished cognitive abilities relative to the wild form. The different biobehavioural profiles of the wild and domestic animals can be regarded as adaptations to the different environmental conditions under which they live, i.e., the natural habitat and artificial man-made housing conditions, respectively. Concerning (b), the comparison of infantile, adolescent and adult wild and domestic guinea pigs shows that the typical biobehavioural profile of the domestic form is already present during early phases of life, that is, during early adolescence and weaning. Thus, differences between the domestic and the wild form can be attributed to genetic alterations resulting from artificial selection, and likely to environmental influences during the pre- and perinatal phase. Interestingly, the frequency of play behaviour does not differ between the domestic and wild form early in life, but is significantly higher in domesticated guinea pigs at later ages. Concerning (c), there is some evidence that personalities occur in both wild and domestic animals. However, there may be differences in which behavioural domains – social and sexual behaviour, emotionality, stress-responsiveness – are consistent over time. These differences are probably due to changing selection pressures during domestication.
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