Formation of spatial vegetation patterns in heterogeneous environments.

Functioning of many resource-limited ecosystems is facilitated through spatial patterns. Patterns can indicate ecosystems productivity and resilience, but the interpretation of a pattern requires good understanding of its structure and underlying biophysical processes. Regular patterns are understood to form autogenously through self-organization, for which exogenous heterogeneities are negligible. This has been corroborated by reaction-diffusion models which generate highly regular patterns in idealized homogeneous environments. However, such model-generated patterns are considerably more regular than natural patterns, which indicates that the concept of autogenous pattern formation is incomplete. Models can generate patterns which appear more natural when they incorporate exogenous random spatial heterogeneities (noise), such as microtopography or spatially varying soil properties. However, the mechanism through which noise influences the pattern formation has not been explained so far. Recalling that irregular patterns can form through stochastic processes, we propose that regular patterns can form through stochastic processes as well, where spatial noise is filtered through scale-dependent biophysical feedbacks. First, we demonstrate that the pattern formation in nonlinear reaction-diffusion models is highly sensitive to noise. We then propose simple stochastic processes which can explain why and how random exogenous heterogeneity influences the formation of regular and irregular patterns. Finally, we derive linear filters which reproduce the spatial structure and visual appearance of natural patterns well. Our work contributes to a more holistic understanding of spatial pattern formation in self-organizing ecosystems.