Living with roommates in a shared den: Spatial and temporal segregation among semifossorial mammals.

Positive interspecific interactions in animal communities (i.e. den sharing) have long been overlooked in animal ecology. The assessment of spatiotemporal overlap among species living within the same burrow system is paramount to explain their strategies of interspecific coexistence. We studied spatiotemporal behavioural patterns of coexistence among four den-sharing mammal species (i.e. the crested porcupine Hystrix cristata, the Eurasian badger Meles meles, the red fox Vulpes vulpes and the European pine marten Martes martes), inhabiting a hilly area of central Italy. Intensive camera trapping (September 2015-September 2018) was used to estimate the interspecific overlap of both temporal and spatial activity patterns for all species combinations. An extensive nocturnal temporal overlap was recorded among all the species, except the diurnal pine marten. However, crested porcupines were mostly active in the darkest nights, whereas bright moonlight enhanced the hunting success of the red fox. Activity of badgers was limited in bright nights only during cold months, when predation pressure and poaching risk were the highest. Crested porcupines avoided spatial sharing outside the den with both nocturnal carnivores, particularly during the winter, when its cubs are in the den. Overlap in ranging areas and activity rhythms between the red fox and the Eurasian badger may be promoted by a remarkable food niche partitioning. Conversely, spatiotemporal overlap between red foxes and pine martens suggested a significant interspecific spatial partitioning, due to the overlap in feeding habits. Den-sharing represents a form of positive interspecific interaction which may limit energy waste and increase local species diversity and densities. Species using the same burrow system may show both spatial and temporal niche partitioning throughout the year, thus allowing a non-competitive coexistence.