Dora Biro

Hierarchical group dynamics in pigeon flocks.

Animals that travel together in groups display a variety of fascinating motion patterns thought to be the result of delicate local interactions among group members. Although the most informative way of investigating and interpreting collective movement phenomena would be afforded by the collection of high-resolution spatiotemporal data from moving individuals, such data are scarce and are virtually non-existent for long-distance group motion within a natural setting because of the associated technological difficulties. Here we present results of experiments in which track logs of homing pigeons flying in flocks of up to 10 individuals have been obtained by high-resolution lightweight GPS devices and analysed using a variety of correlation functions inspired by approaches common in statistical physics. We find a well-defined hierarchy among flock members from data concerning leading roles in pairwise interactions, defined on the basis of characteristic delay times between birds’ directional choices. The average spatial position of a pigeon within the flock strongly correlates with its place in the hierarchy, and birds respond more quickly to conspecifics perceived primarily through the left eye—both results revealing differential roles for birds that assume different positions with respect to flock-mates. From an evolutionary perspective, our results suggest that hierarchical organization of group flight may be more efficient than an egalitarian one, at least for those flock sizes that permit regular pairwise interactions among group members, during which leader–follower relationships are consistently manifested.

Emergence of Culture in Wild Chimpanzees: Education by Master-Apprenticeship

This chapter describes a series of field experiments aimed at investigating aspects of emergence of cultural traditions in wild chimpanzee communities. Long-term research at a number of sites in Africa has revealed that each community of chimpanzees has developed its unique set of cultural traditions (Boesch and Boesch-Achermann 2000; Goodall 1986; McGrew 1992; Nishida 1990; Whiten et al. 1999). The evidence poses an intriguing question: How did these unique cultures come into existence?.

Chimpanzees share forbidden fruit.

The sharing of wild plant foods is infrequent in chimpanzees, but in chimpanzee communities that engage in hunting, meat is frequently used as a ‘social tool’ for nurturing alliances and social bonds. Here we report the only recorded example of regular sharing of plant foods by unrelated, non-provisioned wild chimpanzees, and the contexts in which these sharing behaviours occur. From direct observations, adult chimpanzees at Bossou (Republic of Guinea, West Africa) very rarely transferred wild plant foods. In contrast, they shared cultivated plant foods much more frequently (58 out of 59 food sharing events). Sharing primarily consists of adult males allowing reproductively cycling females to take food that they possess. We propose that hypotheses focussing on ‘food-for-sex and -grooming’ and ‘showing-off’ strategies plausibly account for observed sharing behaviours. A changing human-dominated landscape presents chimpanzees with fresh challenges, and our observations suggest that crop-raiding provides adult male chimpanzees at Bossou with highly desirable food commodities that may be traded for other currencies.

Information transfer in moving animal groups

Moving animal groups provide some of the most intriguing and difficult to characterise examples of collective behaviour. We review some recent (and not so recent) empirical research on the motion of animal groups, including fish, locusts and homing pigeons. An important concept which unifies our understanding of these groups is that of transfer of directional information. Individuals which change their direction of travel in response to the direction taken by their near neighbours can quickly transfer information about the presence of a predatory threat or food source. We show that such information transfer is optimised when the density of individuals in a group is close to that at which a phase transition occurs between random and ordered motion. Similarly, we show that even relatively small differences in information possessed by group members can lead to strong collective-level decisions for one of two options. By combining the use of self-propelled particle and social force models of collective motion with thinking about the evolution of flocking we aim to better understand how complexity arises within these groups.

NUMERICAL ORDERING IN A CHIMPANZEE (PAN TROGLODYTES) : PLANNING, EXECUTING, AND MONITORING

Perceptual and cognitive processes underlying the skill of ordering numerals were assessed in a female chimpanzee (Pan troglodytes) with previous experience in computer-assisted numerical competence tasks. The subject was required to order 3 numerals from the range of 0-9 into an ascending series, with occasional probe trials (referred to as switch trials) in which the positions of the 2nd and 3rd numerals were exchanged immediately after the selection of the 1st. On these trials, errors were scored frequently, whereas correct responses to the intermediate numeral became reliably slower. These and other data indicated that the subject had already established, before making the 1st choice, (a) the correct sequence in which she was to select the numerals and (b) the motor sequence leading to a.correct answer. These findings show that a 3-unit ordering task is supported in the chimpanzee, much as it is in humans, by planning, executing, and monitoring phases.

Experimental identification of social learning in wild animals

Field experiments can provide compelling demonstrations of social learning in wild populations. Social learning has been experimentally demonstrated in at least 23 field experiments, in 20 species, covering a range of contexts, such as foraging preferences and techniques, habitat choice, and predator avoidance. We review experimental approaches taken in the field and with wild animals brought into captivity and note how these approaches can be extended. Relocating individuals, introducing trained individual demonstrators or novel behaviors into a population, or providing demonstrator-manipulated artifacts can establish whether and how a particular act can be socially transmitted in the wild and can help elucidate the benefits of social learning. The type, strength, and consistency of presented social information can be varied, and the provision of conditions favoring the performance of an act can both establish individual discovery rates and help determine whether social information is needed for acquisition. By blocking particular avenues of social transmission or removing key individuals, routes of transmission in wild populations can be investigated. Manipulation of conditions proposed to favor social learning can test mathematical models of the evolution of social learning. We illustrate how field experiments are a viable, vital, and informative approach to the study of social learning.

Homing pigeons develop local route stereotypy.

The mechanisms used by homing pigeons (Columba livia) to navigate homeward from distant sites have been well studied, yet the mechanisms underlying navigation within, and mapping of, the local familiar area have been largely neglected. In the local area pigeons potentially have access to a powerful navigational aid--a memorized landscape map. Current opinion suggests that landmarks are used only to recognize a familiar start position and that the goalward route is then achieved solely using compass orientation. We used high-resolution global positioning system (GPS) loggers to track homing pigeons as they became progressively familiar with a local homing task. Here, we demonstrate that birds develop highly stereotyped yet individually distinctive routes over the landscape, which remain substantially inefficient. Precise aerial route recapitulation implies close control by localized geocentric cues. Magnetic cues are unlikely to have been used, since recapitulation remains despite magnetic disruption treatment, and olfactory cues would have been positionally unstable under the variable wind conditions, making visual landmarks the most likely cues used.

Pigeons combine compass and landmark guidance in familiar route navigation

How do birds orient over familiar terrain? In the best studied avian species, the homing pigeon (Columba livia), two apparently independent primary mechanisms are currently debated: either memorized visual landmarks provide homeward guidance directly, or birds rely on a compass to home from familiar locations. Using miniature Global Positioning System tracking technology and clock-shift procedures, we set sun-compass and landmark information in conflict, showing that experienced birds can accurately complete their memorized routes by using landmarks alone. Nevertheless, we also find that route following is often consistently offset in the expected compass direction, faithfully reproducing the shape of the track, but in parallel. Thus, we demonstrate conditions under which compass orientation and landmark guidance must be combined into a system of simultaneous or oscillating dual control.

What are leaders made of? The role of individual experience in determining leader–follower relations in homing pigeons

Negotiating joint routes during group travel is one of the challenges faced by collectively moving animals, on spatial scales ranging from daily foraging trips to long-distance migrations. Homing pigeons, Columba livia, provide a useful model system for studying the mechanisms of group decision making in the context of navigation, owing to the combination of their gregarious nature and the depth of our understanding of their individual orientational strategies. Previous work has shown that during paired flight, if two birds’ individually preferred routes are sufficiently different, one bird will emerge as leader whom the other follows. What determines the identity of a leader has important implications for the efficiency of a moving collective, since leaders with higher navigational certainty can increase the accuracy of the group. We examined factors contributing to the establishment of leadership/followership, focusing on the role of previous navigational experience. We tested, on a homing task, pairs of pigeons in which the two partners had relatively greater and lesser prior experience, generated through individual training. Analysis of the GPS-tracked routes taken by such pairs revealed a negative correlation between homing experience and the probability that a pigeon would follow a co-navigating partner. Thus, the larger the difference in experience between two partners, the higher the likelihood the more experienced bird would emerge as leader. Our results contribute to a better understanding of the mechanisms and potential payoffs of collective navigational decision making in species that travel in mixed-experience groups.

Context-dependent hierarchies in pigeons

Hierarchical organization is widespread in the societies of humans and other animals, both in social structure and in decision-making contexts. In the case of collective motion, the majority of case studies report that dominant individuals lead group movements, in agreement with the common conflation of the terms “dominance” and “leadership.” From a theoretical perspective, if social relationships influence interactions during collective motion, then social structure could also affect leadership in large, swarm-like groups, such as fish shoals and bird flocks. Here we use computer-vision–based methods and miniature GPS tracking to study, respectively, social dominance and in-flight leader–follower relations in pigeons. In both types of behavior we find hierarchically structured networks of directed interactions. However, instead of being conflated, dominance and leadership hierarchies are completely independent of each other. Although dominance is an important aspect of variation among pigeons, correlated with aggression and access to food, our results imply that the stable leadership hierarchies in the air must be based on a different set of individual competences. In addition to confirming the existence of independent and context-specific hierarchies in pigeons, we succeed in setting out a robust, scalable method for the automated analysis of dominance relationships, and thus of social structure, applicable to many species. Our results, as well as our methods, will help to incorporate the broader context of animal social organization into the study of collective behavior.