Abstract
The evolution of costly cooperation, where cooperators pay a personal cost to benefit others, requires that cooperators interact more frequently with other cooperators. This condition, called positive assortment, is known to occur in spatially-structured viscous populations, where individuals typically have low mobility and limited dispersal. However many social organisms across taxa, from cells and bacteria, to birds, fish and ungulates, are mobile, and live in populations with considerable inter-group mixing. In the absence of information regarding others’ traits or conditional strategies, such mixing may inhibit assortment and limit the potential for cooperation to evolve. Here we employ spatially-explicit individual-based evolutionary simulations to incorporate costs and benefits of two coevolving costly traits: cooperative and local cohesive tendencies. We demonstrate that, despite possessing no information about others’ traits or payoffs, mobility (via self-propulsion or environmental forcing) facilitates assortment of cooperators via a dynamically evolving difference in the cohesive tendencies of cooperators and defectors. We show analytically that this assortment can also be viewed in a multilevel selection framework, where selection for cooperation among emergent groups can overcome selection against cooperators within the groups. As a result of these dynamics, we find an oscillatory pattern of cooperation and defection that maintains cooperation even in the absence of well known mechanisms such as kin interactions, reciprocity, local dispersal or conditional strategies that require information on others’ strategies or payoffs. Our results offer insights into differential adhesion based mechanisms for positive assortment and reveal the possibility of cooperative aggregations in dynamic fission-fusion populations.
Author Summary Cooperation among animals is ubiquitous. In a cooperative interaction, the cooperator confers a benefit to its partner at a personal cost. How does natural selection favour such a costly behaviour? Classical theories argue that cooperative interactions among genetic relatives, reciprocal cooperators, or among individuals within groups in viscous population structures are necessary to maintain cooperation. However, many organisms are mobile, and live in dynamic (fission-fusion) groups that constantly merge and split. In such populations, the above mechanisms may be inadequate to explain cooperation. Here, we develop a minimal model that explicitly accounts for mobility and cohesion among organisms. We find that mobility can support cooperation via emergent dynamic groups, even in the absence of previously known mechanisms. Our results may offer insights into the evolution of cooperation in animals that live in fission fusion groups, such as birds, fish or mammals, or microbes living in turbulent media, such as in oceans or in the bloodstreams of animal hosts.
