When and why do territorial coalitions occur? Experimental evidence from a fiddler crab

Neighboring territory owners are often less aggressive toward each other than to strangers ("dear enemy" effect). There is, however, little evidence for territorial defense coalitions whereby a neighbor will temporarily leave his/her own territory, enter that of a neighbor, and cooperate in repelling a conspecific intruder. This is surprising, as theoreticians have long posited the existence of such coalitions and the circumstances under which they should evolve. Here we document territorial defense coalitions in the African fiddler crab Uca annulipes, which lives in large colonies wherein each male defends a burrow and its surrounding area against neighbors and "floaters" (burrowless males). Fights between a resident and a floater sometimes involve another male who has left his territory to fight the floater challenging his neighbor. Using simple experiments, we provide the first evidence of the rules determining when territorial coalitions form. Our results support recent models that suggest that these coalitions arise from by-product mutualism.