The role of per-capita productivity in the evolution of small colony sizes in ants

The evolution of colony size in social insects is influenced by both extrinsic and colony-intrinsic factors. An important intrinsic trait, per-capita productivity, often declines in larger colonies. This pattern, known as Michener's paradox, can limit the growth of insect societies. In this study, we first describe this problem, survey its occurrence across different ant species, and present a case study of eight cavity-dwelling ants with very small colony sizes. In these species, colonies might never reach sizes at which per-capita productivity decreases. However, in six out of the eight focal species, per-capita productivity did decline with increasing size, in accordance with other studies on per-capita productivity in ants. Several mechanisms, such as resource availability or nest-site limitation, may explain the decrease in per-capita productivity with increases in colony size in our focal species. In these central-place foragers, the individual foraging mode is expected to lead to an increase in travel time as colonies grow. We suggest that polydomy, the concomitant occupation of several nest sites, could serve as a potential strategy to overcome this limitation. Indeed, for one species, we show that polydomy can help to circumvent the reduction in productivity with increasing colony size, suggesting that limited resource availability causes the observed decrease in per-capita productivity. Finally, we discuss the influence of other factors, such as the nesting ecology and colony homeostasis, on the evolution of colony size in these cavity-dwelling ants.