Effects of ungulate density and sociality on landscape heterogeneity: a mechanistic modeling approach

Animals can be important vectors of nutrient transfer within and across landscapes, with important implications for ecosystem productivity and composition. While it is presumed large ungulates are agents of nutrient dispersal via movement and activity, research analyzing their net effects on landscapes remains scarce. We present an individual‐based model that investigates how caribou affect the distribution of nutrients on a landscape through consumption only, as well through the cumulative effects of consumption and nutrient deposition (i.e. fecal waste and carcass deposition). We explored these dynamics in simulations that altered the context of environments, either initially containing heterogeneous or homogeneous nutrient distributions, animal densities and sociality of caribou behavior. In the consumption‐only simulations, caribou density and sociality created different patterns of heterogeneity at both the landscape and local scale depending on the initial landscape conditions. In these simulations, caribou populations crashed at high densities because the lack of animal deposition resulted in low resources across the landscape. This was not the case when considering the cumulative effects of consumption and deposition, indicating the return of nutrients from animals may be important for population stability. Additionally, in simulations considering the cumulative effects of caribou, increasing caribou density increased landscape heterogeneity irrespective of the initial condition (i.e. heterogeneous and homogeneous landscapes), and maintained or increased local heterogeneity in heterogeneous and homogenous landscape, respectively. Importantly, in all simulations the net impact of caribou at the individual patch level was extremely variable, suggesting that animal inputs are highly varied throughout the landscape. Our results indicate the movement of large ungulates such as caribou can increase the heterogeneity of available nutrients within a landscape and provide an important feedback for population stability. Thus, the loss of large ungulates from natural ecosystems via anthropogenic activity is likely to result in less heterogeneous natural landscapes.