Abstract: In studies of animal behaviour investigators correlate dominance with all kinds of behavioural
variables, such as reproductive success and foraging success. Many methods are used to
produce a dominance hierarchy from a matrix reflecting the frequency of winning dominance
interactions. These different methods produce different hierarchies. However, it is difficult to
decide which ranking method is best. In this paper, we offer a new procedure for this decision:
we use an individual-based model, called DomWorld, as a test-environment. We choose this
model, because it provides access to both the internal dominance values of artificial agents
(which reflects their fighting power) and the matrix of winning and losing among them and,
in addition, because its behavioural rules are biologically inspired and its group-level patterns
resemble those of real primates. We compare statistically the dominance hierarchy based on
the internal dominance values of the artificial agents with the dominance hierarchy produced
by ranking individuals by (a) their total frequency of winning, (b) their average dominance
index, (c) a refined dominance index, the David`s score, (d) the number of subordinates each
individual has and (e) a ranking method based on maximizing the linear order of the hierarchy.
Because dominance hierarchies may differ depending on group size, type of society, and the
interval of study, we compare these ranking methods for these conditions.We study complete
samples as well as samples randomly chosen to resemble the limitations of observing real
animals. It appears that two methods of medium complexity (the average dominance index
and David`s score) lead to hierarchical orders that come closest to the hierarchy based on
internal dominance values of the agents. We advocate usage of the average dominance index,
because of its computational simplicity.

Abstract: We spent two winters studying the social behaviour of wild woodland caribou (Rangifer tarandus caribou) at a time when their main food (ground lichens; Cladina sp.) is available only at snow craters dug by the animals. The competition for access to such craters was severe, the animals constantly trying to take over the craters of others. During a two-month period when a group maintained a constant size (20) and composition (all age-sex classes represented), we could rank the animals in a rather linear dominance hierarchy (Landau's index = 0.87). Rank was correlated with access to resources, percent of time spent active, and percent of time feeding in craters. It was also correlated with age and antler size. However, rank is not an attribute of individuals, but of a relationship between individuals. As such it is only an intervening variable between physical attributes and access to resources, a variable whose value has meaning only within a given group. Among the three attributes studied (age, sex, antler size), the latter was by far the best predictor of the occurrence and outcome of interactions. Between two individuals within any of the three age-sex classes studied (adult and yearling males and adult females), the one with larger antlers initiated significantly more often, escalated its aggression (to the point of hitting the target) less often, and enjoyed a higher success rate in obtaining resources. When their antlers were larger than those of an adult male target (i.e. males that had shed their antlers), adult females won almost all their interactions with adult males even though they escalated only one fourth of them. This clarifies the long-standing speculation that female caribou have antlers and shed them later than males, in order to overcome their sexual handicap in competition for food in the winter. We conclude that the link between rank and dominance of an individual on one hand, and some of its attributes on the other (e.g. sex, age, weight, antler size) is fundamentally realized by the animal itself through its active preference for targets it is likely to beat, i.e. targets with smaller antlers.

Abstract: This study examines the role of observation during the formation of triads in female domestic hens. Results indicate that during hierarchy formation, a hen observing agonistic interactions and conflict settlement between its former dominant and a stranger uses this information when in turn confronted by the latter. Under a first condition (E, N = 15 triads), bystanders witnessed their prior dominant being defeated by a stranger before being introduced to them. In a second condition (C1, N = 16 triads), bystanders witnessed the victory of their prior dominant over a stranger. In a third condition (C2, N = 15 triads), bystanders witnessed two strangers establishing a dominance relationship before being introduced to their prior dominant and to a stranger the former had just defeated. The behavioural strategies of bystanders depended on the issue of the conflict they had witnessed. Bystanders of the E condition behaved as having no chance of defeating the stranger. They never initiated an attack against it, and upon being attacked, readily submitted in turn to the stranger. On the contrary, bystanders of the C1 condition behaved as having some chances against the stranger. They initiated attacks in 50% of cases, and won 50% of conflicts against the stranger. Under condition C2, bystanders first initiated contact with the strangers in only 27% of cases, which approximates the average of their chances for defeating the stranger. However, bystanders finally defeated the strangers in 40% of cases. These results suggest that bystanders of conditions E and C1 gained some information on the relationship existing between their prior dominant and the stranger and that they used it coherently, perhaps through transitive inference, thus contributing to the existence of transitive relationships within the triads. Alternate explanations are examined.

Abstract: Worm-running is behaviour in which a chick runs carrying a worm-like object while flock mates follow and attempt to grab the object from its beak. We hypothesised that social ranks based on worm-running frequency are stable over time and are positively correlated with social ranks based on success in aggressive interactions when older. At 8-12 days of age, we scored worm-running in 17 groups of 12 female White Leghorn chicks during three 10-min tests. Based on instantaneous scans at 5-s intervals, the bird carrying the `worm' most often was placed in rank one and so on down the rank order. These tests were repeated at 68-70 days of age. An aggression index for each bird was calculated as the number of aggressive acts given, divided by the number given and received, during three 1-h observation periods when the birds were 68-70 days. Ranks obtained in worm-running tests were positively correlated over the two age periods (P<0.05) but were not correlated with ranks based on the aggression index (P>0.05). Our results indicate that worm-running ranks are not predictive of success in aggressive interactions. Instead, worm-running fits some criteria for play.

Abstract: This simulation study explores some conditions leading to transitivity within dominance orders. Combinations of three parameters were varied to study their consequences upon hierarchy formation and upon the degree of linearity of resultant structures. The factors studied were: (1) the importance of initial resource holding potentials (RHPs); (2) changes brought in RHPs by successive victories and defeats; and (3) accuracy of RHP assessment made by opponents. Results show that initial differences in RHP always lead to perfectly transitive chains whose rank order reflects the importance of initial differences. Even when simulated animals make important errors while assessing each other during round robin tournaments, emerging dominance structures are perfectly linear and ranks obtained in the structure are highly correlated with initial values in RHPs. Moreover, accumulated experiences of victory and/or defeat alone always lead to perfectly linear hierarchies. Their combination with initial individual differences in RHP led to the same conclusion. Even when assessment was far from being perfect, not only perfect chains were formed but initial values in RHPs significantly influenced rank order when the contribution of victory and defeat to RHP was relatively unimportant. The higher the importance of victory and defeat to RHP as compared to that of initial RHP values, the lower was the correlation between initial RHP values and the ranks order reached by individuals in the resultant hierarchies. In general also, the lower the variation within initial RHPs, the lower was the correlation between initial RHPs and ranks in the hierarchy. At a given level of initial RHP dispersion, increasing the contribution of victory and defeat to RHP diminished the correlation between initial RHP values and obtained ranks. In addition, inaccurate assessment reduced the overall correlation, especially when dispersion of initial RHP values was low and the contribution of victory and defeat relatively unimportant. These results shed some light on the controversy about the respective roles of initial individual attributes and that of patterns of resolution in the formation of animal hierarchies. We present the emergence of social order within closed systems as those simulated here as a case of self-organization.