Abstract: I review literature on four different approaches to the study of traditions in animals: observation of free-living animals, laboratory experiment, armchair analysis, and field experiment. Because, by definition, a tradition entails social learning of some kind, it is difficult, perhaps impossible, to establish that a behavior is in fact traditional without knowledge of how it develops. Observations of free-living animals often provide strong circumstantial evidence of a tradition. However, even in the view of several researchers who have studied possibly traditional behaviors in natural populations, observation alone has not proven sufficient to show that social learning contributes to development of behaviors of interest. The relevance of laboratory experiments to the understanding of the development of behaviors in free-living animals is always open to challenge. Armchair analyses of field data can produce interesting hypotheses but cannot test them. Field experiments to determine how behaviors of interest develop in population members provide a promising way forward.

Abstract: Theoretical models predict that the cumulative number of individuals displaying a socially learned novel behavior will follow an accelerating pattern over time, whereas asocial processes have been associated with linear or decelerating functions. This raises the possibility that the shape of the diffusion curve may reveal something about the learning processes involved. If true, this would be particularly useful for identifying social transmission in observational field studies. Published data are reviewed and are found to provide limited support for this view. The use of accelerating curves as a diagnostic is challenging because (1) alternative theoretical models make similar predictions, (2) clear supporting empirical data are lacking, and (3) practical considerations frequently make accurate construction of the diffusion curve difficult.

Abstract: Evidence from both field and laboratory is consistent with the hypothesis that animals can acquire mate preferences by observing the mating behavior of others. It is difficult, however, to distinguish social learning about mates from a host of other social effects on mating that do not produce changes in preferences. Examples are drawn from laboratory studies on mate choice in female and male Japanese quail that illustrate ways in which social cues influence mating decisions. Quail of both sexes use social cues to modify their mate choices, but the sexes use the information to serve different purposes. Female quail gain preferences for males seen mating with other females, whereas males avoid females that they had observed mating with other males. This sex difference in social learning provides an example of how costs and benefits of sexual behavior can shape decision-making processes. Implications of the influence of social learning on sexual selection are briefly discussed.

Abstract: nderstanding herd organization is important when considering management alternatives designed to benefit or manipulate elk (Cervus elaphus) populations. We studied the seasonal and annual herd organization of cow elk in Custer State Park, South Dakota from 1993-1997 by examining seasonal subherd range size, spatial arrangement, overlap, and site fidelity. Based on social interaction analyses, we combined locations of radiocol-lared cow elk to delineate subherds. We computed 95% kernel home ranges with least-squares cross validation for each subherd by season and year. Subherd overlap and fidelity by season and year were computed using the Volume of Intersection Index (VI) statistic. We identified 5 relatively discrete, resident cow-calf subherds. We observed little overlap in utilization distributions of adjacent subherds. The mean VI score across all subherds and time points (n=140) was 0.06 (SE=0.009), indicating an average 6% overlap in subherd area utilization. Subherd overlap between pairs was 0.08 in fall (SE= 0.021), 0.06 in winter (SE=0.018), 0.06 in spring (SE=0.2), and 0.05 in summer (SE= 0.016). Range sizes were not different between any pairs of seasons or years (F13,52=0.7, P=0.75). Subherd fidelity ranged from 0.41 (SE=0.033) to 0.60 (SE=0.018) overall, indicating differential use within the subherd boundary across years. The ability to distinguish discrete cow-calf subherd units is consistent with other studies and may aid elk management in Custer State Park. However, use patterns within subherd boundaries were inconsistent across years and may reflect human disturbances (e.g., hunting and logging activities), differences in our sampling approach, or changes in matriarchal leadership. Further evaluation into factors affecting space-use patterns is necessary to predict changes in range use within the subherd boundary.

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.