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

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