Fruit-frugivore dependencies are important in Ebolavirus outbreaks in Sub-Saharan Africa

Ebolaviruses have the ability to infect a wide variety of species, with many African mammals potentially serving either as primary reservoirs or secondary amplifying hosts. Previous work has shown that frugivorous bats and primates are often associated with spillover and outbreaks. Yet the role that patterns of biodiversity, either of mammalian hosts or of common fruiting species such as Ficus (figs, fruit resources used by a wide variety of species), play in driving outbreak risk remains unclear. We investigated what factors most directly influence Ebolavirus outbreak risk in Sub-Saharan Africa by using a phylogenetically informed path analysis to compare a wide array of potential models (path diagrams) of spatial dynamics. We considered mammalian frugivore richness, cercopithecid and hominid primate richness, richness of pteropodid (fruit) bats, the spatial distribution of species that have tested positive for Ebolavirus antibodies in the wild, Ficus habitat suitability, and environmental conditions (mean annual and variability in temperature and rainfall). The proximate factors that most influenced whether a given host species range contained a site of a previous outbreak event were 1) habitat suitability for Ficus and 2) the diversity of cercopithecid primates. Frugivore richness overall (including bats, primates, and a few other mammals) and the richness of bats in the family Pteropodidae had a strong effect on which species tested positive for Ebolavirus antibodies, but did not influence outbreak risk directly in pathways explored. We interpret this as evidence that foraging around Ficus and frugivorous mammals (such as cercopithecid primates which are commonly hunted for food) play a prominent role in driving outbreaks into human communities, relative to other factors we considered which influence outbreak risk more indirectly.