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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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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.
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King, S. R. B., & Gurnell, J. (2007). Scent-marking behaviour by stallions: an assessment of function in a reintroduced population of Przewalski horses (Equus ferus przewalskii). J Zool, 272(1), 30–36.
Abstract: Abstract Scent marking is a common form of intraspecific communication in mammal species, and using faeces or urine is a cost-effective way of signalling competitive ability and resource holding power. Marking is ritually performed by male equids, and here we assess the function of male scent-marking behaviour in a recently introduced population of Przewalski horses Equus ferus przewalskii in Mongolia. Two forms of scent marking were observed: defecation on stud piles formed from repeated dunging in the same place, and overmarking of faeces and urine of mares. Stud piles were marked with dung by the harem holder and sniffed before and after dung was deposited. They were not found specifically at the periphery of harem ranges but occurred for the most part along routes used by the horses, and were more common in the core parts of harem ranges or where harem ranges overlapped. Thus, rather than being used to defend range boundaries, stud piles were placed predominantly where they would be encountered by male intruders. Mare excreta were covered with urine by the stallion, but were only sniffed before they were marked, not after. These marks appear to advertise to the mare and other, intruding stallions that the harem holder was the mare's consort and that the interloper should not risk trying to steal the mare or sneak a mating. Thus, the two forms of marking by harem holders appear to combine as first and second lines of defence of paternity rights in male intrasexual competition.
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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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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.
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Alexander, R., MCN et al. (1977). Fast locomotion of some african ungulates. J Zool, 183(3), 291–300.
Abstract: ABSTRACT
Ten species of ungulate were filmed, galloping in their natural habitat. They ranged in size from Thomson's gazelle (about 20 kg) to giraffe (about 1000 kg). They were pursued to make them run as fast as possible. The films have been analysed to determine speed, stride frequency, stride and step lengths, and duty factors. The dependence of these quantities on body size is discussed.
Summary:
Fast locomotion of zebra, giraffe, warthog and seven species of Bovidae has been studied. The animals were filmed from a pursuing vehicle while galloping in their natural habitat.
Stride frequency was more closely correlated with limb length (represented by hip height) than with body mass. Mean stride frequency was proportional to (hip height)-0·51 and maximum stride frequency to (hip height) -0·63.
Maximum speed was between 10 and 14 m s -1 for all species except buffalo (7 m s -1). It was not significantly correlated with body mass.
Since the small species ran at least as fast as the large ones they attained higher Froude numbers. Relative stride length was approximately 1·8 (Froude number)0·39 for all species, irrespective of size. Relative step length was approximately 0·65 (Froude number)0·2, both for the fore feet and for the hind ones. The vertical forces exerted by the feet are proportional to (body weight)×(Froude number)0·2 so the forces at maximum speed are larger multiples of body weight for small species than for large ones.
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