Abstract: We analyzed the leadership behavior of breeding and nonbreeding gray wolves (Canis lupus) in three packs during winter in 1997-1999. Scent-marking, frontal leadership (time and frequency in the lead while traveling), initiation of activity, and nonfrontal leadership were recorded during 499 h of ground-based observations in Yellowstone National Park. All observed scent-marking (N = 158) was done by breeding wolves, primarily dominant individuals. Dominant breeding pairs provided most leadership, consistent with a trend in social mammals for leadership to correlate with dominance. Dominant breeding wolves led traveling packs during 64% of recorded behavior bouts (N = 591) and 71% of observed travel time (N = 64 h). During travel, breeding males and females led packs approximately equally, which probably reflects high parental investment by both breeding male and female wolves. Newly initiated behaviors (N = 104) were prompted almost 3 times more often by dominant breeders (70%) than by nonbreeders (25%). Dominant breeding females initiated pack activities almost 4 times more often than subordinate breeding females (30 vs. 8 times). Although one subordinate breeding female led more often than individual nonbreeders in one pack in one season, more commonly this was not the case. In 12 cases breeding wolves exhibited nonfrontal leadership. Among subordinate wolves, leadership behavior was observed in subordinate breeding females and other individuals just prior to their dispersal from natal packs. Subordinate wolves were more often found leading packs that were large and contained many subordinate adults.
Nous avons analysé le comportement de commandement chez des loups gris (Canis lupus) reproducteurs et non reproducteurs appartenant à  trois meutes durant les hivers de 1997-1999. Le marquage d'odeurs, la position en tête de meute (la durée et la fréquence au cours des déplacements), l'initiation des activités et la prise de décisions ailleurs qu'en tête du groupe ont été notés pendant 499 h d'observations au sol dans le Parc national de Yellowstone. Tous les marquages (N = 158) ont été faits par des loups reproducteurs, surtout des individus dominants. Ce sont surtout les couples dominants qui assurent le commandement, en accord avec une tendance chez les mammifères sociaux chez lesquels la fonction de chef est en corrélation avec la dominance. Les loups reproducteurs dominants ont conduit les meutes en déplacement pendant 64 % (N = 591) des épisodes de comportement et pendant 71 % des épisodes de déplacement (N = 64 h). Les mâles et les femelles reproducteurs ont dirigé les meutes en déplacement à peu près également, ce qui reflète probablement l'investissement parental important aussi bien de la part des reproducteurs mâles que des femelles. Les comportements nouveaux (N = 104) ont été adoptés presque trois fois plus souvent par des reproducteurs dominants (70 %) que par des individus non reproducteurs (25 %). Des femelles reproductrices dominantes ont été instigatrices des activités de leur meute environ quatre fois plus souvent que les femelles reproductrices subordonnées (30 vs. 8 fois). Bien qu'une femelle reproductrice subordonnée ait pris la direction de sa meute plus souvent que les individus non reproducteurs au cours d'une saison, cela n'est pas habituel. Dans 12 cas, des loups reproducteurs ont pris la direction de leur meute sans être en tête. Chez les individus subordonnés, le comportement de commandement a été observé chez des femelles reproductrices et chez d'autres individus juste avant qu'ils ne quittent leur meute d'origine au moment de la dispersion. Les loups subordonnés mènent surtout de grands troupeaux qui comptent beaucoup d'individus subordonnés.[Traduit par la Rédaction]

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.

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.

Abstract: In many social species, relationships within groups seem to be non-random but related to variables such as rank, kinship or sexual attractiveness. The endangered African wild dog Lycaon pictus is a social carnivore that lives in large, stable packs, and intra-pack associations might be expected to display similar patterns. We investigated patterns of coalition formation (support during dominance interactions, and partnership interactions) and resting associations between members of a captive pack of 19 wild dogs. The social organization of the captive pack was similar to that of free-ranging packs in many respects. Polyadic (group) incidents of coalition support were also observed in a free-ranging pack. Patterns of coalition formation in the captive pack were related to rank. Most aggressive interactions involved high-ranking individuals (particularly the alpha, beta and third-ranking males) and coalitionary support tended to reinforce the existing hierarchy. However, there was at least one example of support influencing a successful rank challenge. Support was affected by potential risks and benefits, the latter including dominance through association and revolutionary alliances. An even stronger pattern overlaid associations between pack members: coalitions and resting associations were strongest between members of the same age–sex cohort, and may have enabled the eventual dominance of younger pack members over adults. Among adults, coalitionary associations were sometimes overridden by intersexual relationships. The results from this captive pack suggest that wild dogs are sensitive to differences in competitive ability. This information, in conjunction with strong affiliative bonds between littermates, is used to manoeuvre for position in the social hierarchy. It may also be important during dispersal, in encounters with other dispersing groups of the same sex. Although most features of the social structure of the captive pack were comparable to those of free-ranging packs, aspects such as the influence of relatedness on coalition formation still need to be explored.

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.