Flow interactions between uncoordinated flapping swimmers give rise to group cohesion.

Many species of fish and birds travel in groups, yet the role of fluid-mediated interactions in schools and flocks is not fully understood. Previous fluid-dynamical models of these collective behaviors assume that all individuals flap identically, whereas animal groups involve variations across members as well as active modifications of wing or fin motions. To study the roles of flapping kinematics and flow interactions, we design a minimal robotic "school" of two hydrofoils swimming in tandem. The flapping kinematics of each foil are independently prescribed and systematically varied, while the forward swimming motions are free and result from the fluid forces. Surprisingly, a pair of uncoordinated foils with dissimilar kinematics can swim together cohesively-without separating or colliding-due to the interaction of the follower with the wake left by the leader. For equal flapping frequencies, the follower experiences stable positions in the leader's wake, with locations that can be controlled by flapping amplitude and phase. Further, a follower with lower flapping speed can defy expectation and keep up with the leader, whereas a faster-flapping follower can be buffered from collision and oscillate in the leader's wake. We formulate a reduced-order model which produces remarkable agreement with all experimentally observed modes by relating the follower's thrust to its flapping speed relative to the wake flow. These results show how flapping kinematics can be used to control locomotion within wakes, and that flow interactions provide a mechanism which promotes group cohesion.

Flow interactions lead to self-organized flight formations disrupted by self-amplifying waves.

Collectively locomoting animals are often viewed as analogous to states of matter in that group-level phenomena emerge from individual-level interactions. Applying this framework to fish schools and bird flocks must account for visco-inertial flows as mediators of the physical interactions. Motivated by linear flight formations, here we show that pairwise flow interactions tend to promote crystalline or lattice-like arrangements, but such order is disrupted by unstably growing positional waves. Using robotic experiments on "mock flocks" of flapping wings in forward flight, we find that followers tend to lock into position behind a leader, but larger groups display flow-induced oscillatory modes - "flonons" - that grow in amplitude down the group and cause collisions. Force measurements and applied perturbations inform a wake interaction model that explains the self-ordering as mediated by spring-like forces and the self-amplification of disturbances as a resonance cascade. We further show that larger groups may be stabilized by introducing variability among individuals, which induces positional disorder while suppressing flonon amplification. These results derive from generic features including locomotor-flow phasing and nonreciprocal interactions with memory, and hence these phenomena may arise more generally in macroscale, flow-mediated collectives.

Hydrodynamic schooling of flapping swimmers.

Fish schools and bird flocks are fascinating examples of collective behaviours in which many individuals generate and interact with complex flows. Motivated by animal groups on the move, here we explore how the locomotion of many bodies emerges from their flow-mediated interactions. Through experiments and simulations of arrays of flapping wings that propel within a collective wake, we discover distinct modes characterized by the group swimming speed and the spatial phase shift between trajectories of neighbouring wings. For identical flapping motions, slow and fast modes coexist and correspond to constructive and destructive wing-wake interactions. Simulations show that swimming in a group can enhance speed and save power, and we capture the key phenomena in a mathematical model based on memory or the storage and recollection of information in the flow field. These results also show that fluid dynamic interactions alone are sufficient to generate coherent collective locomotion, and thus might suggest new ways to characterize the role of flows in animal groups.