Spatial variation in population densities across a landscape is a feature of many ecological systems, from
self-organised patterns on mussel beds to spatially restricted insect outbreaks. It occurs as a result of
environmental variation in abiotic factors and/or biotic factors structuring the spatial distribution of
populations. However the ways in which abiotic and biotic factors interact to determine the existence
and nature of spatial patterns in population density remain poorly understood. Here we present a new
approach to studying this question by analysing a predator–prey patch-model in a heterogenous
landscape. We use analytical and numerical methods originally developed for studying nearest-
neighbour (juxtacrine) signalling in epithelia to explore whether and under which conditions patterns
emerge. We find that abiotic and biotic factors interact to promote pattern formation. In fact, we find a
rich and highly complex array of coexisting stable patterns, located within an enormous number of
unstable patterns. Our simulation results indicate that many of the stable patterns have appreciable
basins of attraction, making them significant in applications. We are able to identify mechanisms for
these patterns based on the classical ideas of long-range inhibition and short-range activation, whereby
landscape heterogeneity can modulate the spatial scales at which these processes operate to structure
the populations
