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JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. Online enhancements: appendixes.abstract: In eusocial species, some individuals sacrifice their own reproduction for the benefit of others. It has been argued that the evolution of sterile helpers in eusocial insects requires synergistic efficiency gains through cooperation that are uncommon in cooperatively breeding vertebrates and that this precludes a universal ecological explanation of social systems with alloparental care. In contrast, using a model that incorporates realistic ecological mechanisms of population regulation, we show here that constraints on independent breeding (through nest-site limitation and dispersal mortality) eliminate any need for synergistic efficiency gains: sterile helpers may evolve even if they are relatively inefficient at rearing siblings, reducing their colony's per-capita productivity. Our approach connects research fields by using hypotheses developed for cooperative breeding to explain the evolution of eusociality. The results suggest that these hypotheses may apply more generally than previously thought.

Models of sex‐allocation conflict are central to evolutionary biology but have mostly assumed static decisions, where resource allocation strategies are constant over colony lifespan. Here, we develop a model to study how the evolution of dynamic resource allocation strategies is affected by the queen‐worker conflict in annual eusocial insects. We demonstrate that the time of dispersal of sexuals affects the sex‐allocation ratio through sexual selection on males. Furthermore, our model provides three predictions that depart from established results of classic static allocation models. First, we find that the queen wins the sex‐allocation conflict, while the workers determine the maximum colony size and colony productivity. Second, male‐biased sex allocation and protandry evolve if sexuals disperse directly after eclosion. Third, when workers are more related to new queens, then the proportional investment into queens is expected to be lower, which results from the interacting effect of sexual selection (selecting for protandry) and sex‐allocation conflict (selecting for earlier switch to producing sexuals). Overall, we find that colony ontogeny crucially affects the outcome of sex‐allocation conflict because of the evolution of distinct colony growth phases, which decouples how queens and workers affect allocation decisions and can result in asymmetric control.

Evolutionary game theory and the adaptive dynamics approach have made invaluable contributions to understanding how gradual evolution leads to adaptation when individuals interact. Here, we review some of the basic tools that have come out of these contributions to model the evolution of quantitative traits in complex populations. We collect together mathematical expressions that describe directional and disruptive selection in class- and group-structured populations in terms of individual fitness, with the aims of bridging different models and interpreting selection. In particular, our review of disruptive selection suggests there are two main paths that can lead to diversity: (i) when individual fitness increases more than linearly with trait expression; (ii) when trait expression simultaneously increases the probability that an individual is in a certain context (e.g. a given age, sex, habitat, size or social environment) and fitness in that context. We provide various examples of these and more broadly argue that population structure lays the ground for the emergence of polymorphism with unique characteristics. Beyond this, we hope that the descriptions of selection we present here help see the tight links among fundamental branches of evolutionary biology, from life history to social evolution through evolutionary ecology, and thus favour further their integration. This article is part of the theme issue ‘Half a century of evolutionary games: a synthesis of theory, application and future directions’.