Experimentally measured group direct benefits according to worker density explain group living of the termite Reticulitermes chinensis

The evolution of cooperation requires more benefits of group living than solitary lifestyle. However, to some degree, our understanding about the benefits is hindered by abstract debates over theoretical and experimental evidences of individual selection or group selection because it is difficult to examine the actual benefits at the group level. Moreover, group density is a crucial ecological factor which deeply affects group reproduction and survival, few studies have been performed in social insects. Here, we study the effects of worker density on group direct benefits in the termite species Reticulitermes chinensis. The termite R. chinensis is an ideal model which lives with a high worker density in wood. We used the quantity of eggs and the total biomass (biomass of all group members) accumulation as two components of group benefits. We investigated the group benefits in the context of worker density according to eleven worker densities, and we measured the group benefits and the resource consumption with the same group members in two types of artificial nest areas. Moreover, we counted the stomodeal trophallaxis occurrences from any workers to queens under three worker densities to explore the degree of cooperation according to worker density. We found that both the number of eggs and the total biomass accumulation significantly increased with increasing worker density in groups. Furthermore, the consumption of resources was similar between groups with the same number of individuals gathered in small or large nest areas, but the production of eggs and the biomass accumulation were higher in groups of small nest areas than in large nest areas. Additionally, we found the stomodeal trophallaxis behavior significantly increased in higher worker density groups. Our results suggest that the group benefits influenced by the high worker density may at least partially explain the group living of eusocial insects in ecology.