Abstract

The evolution of cooperation among non-kin poses a major theoretical puzzle: why should natural selection favor individuals who help unrelated conspecifics at a cost to themselves? The relevance of architecture to this question has rarely been considered. Here I report cooperation among non-kin in social hermit crabs (Coenobita compressus), where unrelated conspecifics work together to evict larger individuals from a housing market of architecturally remodeled shells. I present (1) the first detailed description of natural coalitions in the wild and (2) a theoretical framework, which examines the evolutionary benefits to each coalition member and predicts when forming a coalition will be successful. In the wild, important ecological and social constraints exist, which are built into the model. Based on these constraints, I show that coalitions can be a successful strategy if several key criteria hold: the coalition is necessary, effective, stable dyadically, and stable polyadically. Notably, the “splitting the spoils” problem—which often undermines non-kin cooperation—is eliminated via architecture: a small individual (C) who helps a medium individual (B) to evict a large individual (A) will ultimately benefit, since C will get B’s left behind shell after B moves into A’s shell. Coalitions, however, can break down due to added layers of social complexity involving third-party “free riders” and “cheaters,” which strategically butt in the architectural queue and thereby steal incentives from the smaller coalition member. Overall, therefore, substantial scope exists for both cooperation and conflict within nature’s housing market of architecture. Experiments are now needed to directly test the impact on coalitions of architecture, from the interior of homes up to whole housing markets.

Highlights

  • Cooperation poses one of the major theoretical puzzles in evolutionary biology

  • Unlike prior explanations for non-kin cooperation, where “splitting the spoils” can undermine the cooperative relationship, this novel architectural solution predicts that the evolutionary interests of non-kin can coincide, on an architectural staircase with discrete reusable resources, like shells

  • Coalitions among non-kin can arise even as genetically unrelated individuals all selfishly pursue their own evolutionary interests, competing fiercely over scarce architectural resources. Future work in this area can profitably wed theory with experiment in an interdisciplinary approach spanning biology, engineering, and mathematics to better understand the coalitions that form in nature

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Summary

Introduction

Cooperation poses one of the major theoretical puzzles in evolutionary biology. Why should natural selection favor individuals who help others at a cost to themselves? Fifty years of theoretical and empirical work has demonstrated the importance of shared genetic material in promoting cooperation among kin: kin share genes in common, and these genes can be indirectly passed on through a genetic relative, so helping kin can be evolutionarily favored (Hamilton, 1964a,b).Architecture of CoalitionsAn elegant theoretical distillation for kin selection and inclusive fitness is Hamilton’s rule (rB > C), which has been supported by an abundance of empirical studies on cooperation among kin (reviewed in Bourke, 2011). Even if a temporary alignment of interests enables a coalition to attain its cooperative goal, a major dilemma exists once this goal is successfully realized: splitting the spoils that are the rewards of the joint cooperative effort (Harcourt and de Waal, 1992). This so-called “splitting the spoils” problem often arises because one member of the coalition, the more powerful one, may monopolize the resulting spoils, thereby eliminating any evolutionary incentive for the other, less powerful coalition member to have even cooperated at all. Insight into potential solutions to this mystery might be had by considering coalitions in the context of architecture

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