Queens of eusocial species live extraordinarily long compared to their workers. So far, it has been argued that these lifespan divergences are readily explained by the classical evolutionary theory of ageing. As workers predominantly perform risky tasks, such as foraging and nest defense, and queens stay in the well‐protected nests, selection against harmful genetic mutations expressed in old age should be weaker in workers than in queens due to caste differences in extrinsic mortality risk, and thus, lead to the evolution of longer queen and shorter worker lifespans. However, these arguments have not been supported by formal models. Here, we present a model for the evolution of caste‐specific ageing in social insects, based on Williams’ antagonistic pleiotropy theory of ageing. In individual‐based simulations, we assume that mutations with antagonistic fitness effects can act within castes, that is, mutations in early life are accompanied by an antagonistic effect acting in later life, or between castes, where antagonistic effects emerge due to caste antagonism or indirect genetic effects between castes. In monogynous social insect species with sterile workers, large lifespan divergences between castes evolved under all different scenarios of antagonistic effects, but regardless of the degree of caste‐specific extrinsic mortality. Mutations with antagonistic fitness effects within castes reduced lifespans of both castes, while mutations with between‐caste antagonistic effects decreased worker lifespans more than queen lifespans, and consequently increased lifespan divergences. Our results challenge the central explanatory role of extrinsic mortality for caste‐specific ageing in eusocial organisms and suggest that antagonistic pleiotropy affects castes differently due to reproductive monopolization by queens, hence, reproductive division of labor. Finally, these findings provide new insights into the evolution of tissue‐specific ageing in multicellular organisms in general.

The evolution of eusociality has long been recognized as an evolutionary paradox. We identify three different but complementary approaches to the study of the evolution of eusociality. Kin selection models explain why individuals can evolve to have distinct reproductive or non-reproductive roles. Division of labour models elucidate how specialisation can emerge and evolve. Conceptual models from evolutionary developmental biology propose hypotheses for the origin and evolutionary fixation of a helper-phenotype. However, we argue that none of these three approaches is a sufficient explanation for the evolution of eusociality on its own. Consequently, we plead for novel unifying explanations and formal models to better understand the interrelation of reproductive altruism, task specialisation and phenotypic plasticity.

The conversion of natural ecosystems to agricultural land is one of the most important drivers of biodiversity decline worldwide, particularly in the tropics. Species loss is typically trait‐associated, leading to filtering of disturbance‐resistant species during community assembly, which affects ecosystem functioning and evolutionary potential of communities. To understand the ecological and phylogenetic impact of rainforest conversion to agricultural systems, we combine analysis of nesting habit, feeding habit, colony size, and body size of canopy ants (Formicidae) with a phylogenetic analysis of species collected in four land‐use systems in Sumatra, Indonesia: (1) lowland tropical rainforest, (2) jungle rubber (extensive rubber agroforest), and smallholder plantations of (3) rubber and (4) oil palm. Canopy ant communities in these land‐use systems differed in trait composition, with a larger proportion of generalist nesting and generalist‐omnivore feeding species in oil palm compared to rainforest and a larger proportion of generalist nesters and species with large colonies (>1000 individuals) in rubber than in rainforest. Traits of canopy ant communities in jungle rubber were more similar to those in rainforest than to those in rubber and oil palm plantations. In rainforest, mean pairwise phylogenetic distance was lower than expected for random community assembly, but did not differ from random in the other land‐use systems. Of the traits nesting habit, feeding habit, and colony size, only feeding habit exhibited phylogenetic signal. Our results show that rainforest conversion to agricultural systems is accompanied by shifts in trait composition of canopy ant communities. Further, our results argue against environmental filtering of closely related canopy ant species as the major community assembly mechanism in plantation systems, but suggest that the Sumatran lowland rainforests harbor recently diverged endemic ant species that are particularly vulnerable to rainforest conversion to agricultural systems. Given the importance of ants for tropical ecosystems, the ecological differences among ant communities along the land‐use gradient might have important consequences for ecosystem functioning and services in plantation systems.

Cooperatively breeding animals live longer than their solitary counterparts. The traditional explanation for this is that cooperative breeding evolves more readily in long-lived species.Here, we reverse this argument and show that long lifespans are an evolutionary consequence of cooperative breeding. Natural selection favours a delayed onset of senescence in cooperative breeders, relative to solitary breeders, because cooperative breeders have a delayed age of first reproduction due to reproductive queueing. Especially long lifespans evolve in cooperative breeders with age-dependent reproductive queueing. Finally, we show that lower genetic relatedness among group members leads to the evolution of longer lifespans. This is because selection against higher mortality is weaker when mortality reduces competition between relatives. Our results link the evolutionary theory of ageing with kin selection theory, demonstrating that the evolution of ageing in cooperative breeders is driven by the timing of reproduction and kin structure within breeding territories. Results

Division of labour occurs in a broad range of organisms. Yet, how division of labour can emerge in the absence of pre-existing interindividual differences is poorly understood. Using a simple but realistic model, we show that in a group of initially identical individuals, division of labour emerges spontaneously if returning foragers share part of their resources with other group members. In the absence of resource sharing, individuals follow an activity schedule of alternating between foraging and other tasks. If non-foraging individuals are fed by other individuals, their alternating activity schedule becomes interrupted, leading to task specialisation and the emergence of division of labour. Furthermore, nutritional differences between individuals reinforce division of labour. Such differences can be caused by increased metabolic rates during foraging or by dominance interactions during resource sharing. Our model proposes a plausible mechanism for the self-organised emergence of division of labour in animal groups of initially identical individuals. This mechanism could also play a role for the emergence of division of labour during the major evolutionary transitions to eusociality and multicellularity.