Ungulate co-occurrence in a landscape of antagonisms.

Protected areas largely now exist as coupled natural-human ecosystems where human activities are increasingly forcing wildlife to adjust behaviors. For many ungulate species that rely on protected areas for their persistence, they must balance these anthropogenic pressures amid natural regulators. Here, we investigated the pressures exerted from humans and livestock, apex predators, and within guild competitors on ungulate co-occurrence patterns in a fragile protected area complex in West Africa. Specifically, we used multi-species occupancy modeling to quantify co-occurrence among four ungulates (Tragelaphus scriptus, Redunca redunca, Kobus kob, Phacochoerus africanus) and applied structural equation models to discern the relative contributions of pressures on co-occurrence patterns. We observed a strong spatial gradient across with higher co-occurrence in the wetter western portion of our ~13,000 km study area. Co-occurrence patterns among ungulate dyads ranged from 0.15 to 0.49 with the smallest body sized pair showing highest levels of sympatry, warthog and reedbuck. We found that anthropogenic pressures, namely cattle had the greatest effect in reducing sympatry among wild ungulates more strongly than the presence of African lions that also exhibited negative effects. Humans, hyenas, and competitors showed positive effects on ungulate co-occurrence. In a region of the world ongoing rapid socio-ecological change with increasing threats from climate and environmental instability, protected areas in West Africa represent a major safeguard for wildlife and human livelihoods alike. Our findings highlight the need for effective interventions that focus on large carnivore conservation, habitat restoration, and containment of livestock grazing to promote the coexistence of biodiversity and socio-economic goals within the region.