Us, them, and the others: Testing for discrimination amongst outgroups in a single-piece nesting termite, .

Recognition of group members is an important adaptation in social organisms because it allows help to be directed toward kin or individuals that are likely to reciprocate, and harm to be directed toward members of competing groups. Evidence in a wide range of animals shows that responses to outgroups vary with context, suggesting that cues to group membership also depend on the social or environmental context. In termites, intergroup encounters are frequent and their outcomes highly variable, ranging from destruction of a colony to colony fusion. As well as genetic factors, nestmate recognition in social insects commonly relies on cues that are mediated by environmental factors such as food source. However, single-piece nesting termite colonies share nesting material and food source with rival colonies (their wood substrate serves as both). In principle, the shared environment of single-piece nesting termite colonies could constrain their ability to identify non-nestmates, contributing to some of the variation seen in encounters, but this has not been investigated. In this study, we raised incipient colonies of a single-piece nesting termite, , on two different wood types and conducted behavioral assays to test whether nestmate discrimination can be constrained by common environmental conditions. We found that non-nestmates elicited higher rates of identity checking and defense behavior compared to nestmates, but there was no effect of wood type on the strength of behavioral responses to non-nestmates. We also found that one key cooperative behavior (allogrooming) was performed equally toward both nestmates and non-nestmates. These findings offer no support for the hypothesis that common wood type constrains the nestmate recognition system of single piece nesting termites. We suggest that where groups encounter each other frequently in a common environment, selection will favor discrimination based on genetic and/or higher resolution environmentally mediated cues.