Self-complexification through integral feedback in eusocial paper wasps of various levels of sociality.

We investigate how simple physical interactions can generate remarkable diversity in the life history of social agents using data of social wasps, yielding complex scalable task partitioning. We built and analyzed a computational model to investigate how diverse task allocation patterns found in nature can emerge from the same behavioral blueprint. Self-organizing mechanisms of interwoven behavioral feedback loops, task-dependent time delays and simple material flows between interacting individuals yield an emergent homeostatic self-regulation while keeping the global colony performance scalable. Task allocation mechanisms based on implicitly honest signaling via material flows are not only very robust but are also highly evolvable due to their simplicity and reliability. We find that task partitioning has evolved to be scalable and adaptable to life history traits, such as expected colony size or temporal bottlenecks in the available workforce or materials. By tuning solely the total number of agents and a social connectivity-related parameter in the model, our simulations yield the whole range of emergent patterns in task allocation and task fidelity akin to observed field data. Our model suggests that the material exchange ("common stomach mechanism") found in many paper wasps provides a common functional "core" across these genera, which not only provides self-regulation of the colony, but also provides a scalable mechanism allowing natural selection to yield complex social integration in larger colonies over the course of their evolutionary trajectory.