Using joint species distribution modelling to identify climatic and non-climatic drivers of Afrotropical ungulate distributions

The relative importance of the different processes that determine the distribution of species and the assembly of communities is a key question in ecology. The distribution of any individual species is affected by a wide range of environmental variables as well as through interactions with other species; the resulting distributions determine the pool of species available to form local communities at fine spatial scales. A challenge in community ecology is that these interactions (e.g. competition, facilitation, etc.) often are not directly measurable. Here, we used hierarchical modelling of species communities (HMSC), a recently developed framework for joint species distribution modelling, to estimate the role of biotic effects alongside environmental factors using latent variables. We investigate the role of these factors determining species distributions in communities of Artiodactyla, Perissodactyla and Proboscidea in the Afrotropics, an area of peak species richness for hoofed mammals. We also calculate pairwise trait dissimilarity between these species, from a mixture of morphological and behavioural traits, and investigate the relationship between dissimilarity and estimated residual co-occurrence in the model. We find that while ungulate distributions appear to be predominantly determined (~ 70%) by climatic variables, such as precipitation, a substantial proportion of the variance in ungulate species distributions (~ 30%) can also be attributed to modelled latent variables that likely represent a combination of dispersal barriers and biotic factors. Although we find only a weak relationship between residual co-occurrence and trait dissimilarity, we suggest that our results may show evidence that biotic factors, likely influenced by historical barriers to species dispersal, are important in determining species communities over a continental area. The HMSC framework can be used to provide insight into factors affecting community assembly at broad scales, and to make more powerful predictions about future species distributions as we enter an era of increasing impacts from anthropogenic change.