“…In contrast, others propose using more direct (also termed "proximal") variables whenever possible (Austin, 2007(Austin, , 2002. However, studies that separate direct versus indirect associations in species distribution modelling are rare (Aguilera et al, 2011;Chen and Pollino, 2012;Gieder et al, 2014;Hamilton et al, 2015). Our study shows that path models such as Gaussian Bayesian Networks can be used to retrieve meaningful direct and indirect associations between physiographic variables, environmental variables and species distribution.…”