No abstract

The evolution of group living transformed the history of animal life on earth, yielding substantial selective benefits. Yet, without overcoming fundamental challenges such as how to coordinate movements with conspecifics, animals cannot maintain cohesion, and coordination is thus a prerequisite for the evolution of sociality. Although it has been considered that animal groups must coordinate the timing and direction of movements, coordinating speed is also essential to prevent the group from splitting. We investigated speed consensus in homing pigeon, Columba livia, flocks using high-resolution GPS. Despite observable differences in average solo speed (which was positively correlated with bird mass) compromises of up to 6% from the preferred solo speed were made to reach consensus in flocks. These results match theory which suggests that groups fly at an intermediate of solo speeds, which suggests speed averaging. By virtue of minimizing extreme compromises, speed averaging can maximize selective benefits across the group, suggesting shared consensus for group speed could be ubiquitous across taxa. Nevertheless, despite group-wide advantages, contemporary flight models have suggested unequal energetic costs in favour of individuals with intermediate body mass/preferred speed (hence the 'Goldilocks principle').

Bird flocks under predation demonstrate complex patterns of collective escape. These patterns may emerge by self-organization from local interactions among group-members. Computational models have been shown to be valuable for identifying what behavioral rules may govern such interactions among individuals during collective motion. However, our knowledge of such rules for collective escape is limited by the lack of quantitative data on bird flocks under predation in the field. In the present study, we analyze the first GPS trajectories of pigeons in airborne flocks attacked by a robotic falcon in order to build a species-specific model of collective escape. We use our model to examine a recently identified distance-dependent pattern of collective behavior: the closer the prey is to the predator, the higher the frequency with which flock members turn away from it. We first extract from the empirical data of pigeon flocks the characteristics of their shape and internal structure (bearing angle and distance to nearest neighbors). Combining these with information on their coordination from the literature, we build an agent-based model adjusted to pigeons’ collective escape. We show that the pattern of turning away from the predator with increased frequency when the predator is closer arises without prey prioritizing escape when the predator is near. Instead, it emerges through self-organization from a behavioral rule to avoid the predator independently of their distance to it. During this self-organization process, we show how flock members increase their consensus over which direction to escape and turn collectively as the predator gets closer. Our results suggest that coordination among flock members, combined with simple escape rules, reduces the cognitive costs of tracking the predator while flocking. Such escape rules that are independent of the distance to the predator can now be investigated in other species. Our study showcases the important role of computational models in the interpretation of empirical findings of collective behavior.

Complex patterns of collective behaviour may emerge through self-organization, from local interactions among individuals in a group. To understand what behavioural rules underlie these patterns, computational models are often necessary. These rules have not yet been systematically studied for bird flocks under predation. Here, we study airborne flocks of homing pigeons attacked by a robotic falcon, combining empirical data with a species-specific computational model of collective escape. By analysing GPS trajectories of flocking individuals, we identify two new patterns of collective escape: early splits and collective turns, occurring even at large distances from the predator. To examine their formation, we extend an agent-based model of pigeons with a ‘discrete’ escape manoeuvre by a single initiator, namely a sudden turn interrupting the continuous coordinated motion of the group. Both splits and collective turns emerge from this rule. Their relative frequency depends on the angular velocity and position of the initiator in the flock: sharp turns by individuals at the periphery lead to more splits than collective turns. We confirm this association in the empirical data. Our study highlights the importance of discrete and uncoordinated manoeuvres in the collective escape of bird flocks and advocates the systematic study of their patterns across species.