Pup guarding by greater spear-nosed bats

Alloparental care poses an evolutionary dilemma because effort is expended on non-filial offspring. Thus, instances of alloparental care have been attributed to either mistaken identity, (i.e., recognition errors) or active cooperation. In greater spear-nosed bats (Phyllostomus hastatus), reproductive females roost together in stable long-term social groups in cave ceilings. Non-volant pups frequently fall from roost sites to the cave floor where they can die unless retrieved by an adult. In this study, we examined the function of adult female visits to non-filial young and tested whether visits were attributable to recognition errors or to cooperation. We found that females visited non-filial pups from their own social group more than expected. Females from different social groups attacked and sometimes killed pups, and male pups were attacked more frequently than female pups. Visits by group mates benefited fallen pups by reducing the likelihood of attack by females from other groups. In contrast to the mistaken identity hypothesis, we found that some females leave their own pups to approach and remain with group mates' pups. We used microsatellite markers to estimate relatedness and test whether kinship could explain this alloparental care and found that females were unrelated to the pups they guard. We conclude that females who reside in highly stable social groups exhibit cooperative behavior that cannot be explained by kinship and is unlikely to be due to direct or generalized reciprocity. Instead, our data suggest that alloparental care likely involves a complex interplay between group membership and cooperative foraging.