The finding that regular spatial patterns can emerge in nature from local interactions between organisms has prompted a search for the ecological importance of these patterns. Theoretical models have predicted that patterning may have positive emergent effects on fundamental ecosystem functions, such as productivity. We provide empirical support for this prediction. In dryland ecosystems, termite mounds are often hotspots of plant growth (primary productivity). Using detailed observations and manipulative experiments in an African savanna, we show that these mounds are also local hotspots of animal abundance (secondary and tertiary productivity): insect abundance and biomass decreased with distance from the nearest termite mound, as did the abundance, biomass, and reproductive output of insect-eating predators. Null-model analyses indicated that at the landscape scale, the evenly spaced distribution of termite mounds produced dramatically greater abundance, biomass, and reproductive output of consumers across trophic levels than would be obtained in landscapes with randomly distributed mounds. These emergent properties of spatial pattern arose because the average distance from an arbitrarily chosen point to the nearest feature in a landscape is minimized in landscapes where the features are hyper-dispersed (i.e., uniformly spaced). This suggests that the linkage between patterning and ecosystem functioning will be common to systems spanning the range of human management intensities. The centrality of spatial pattern to system-wide biomass accumulation underscores the need to conserve pattern-generating organisms and mechanisms, and to incorporate landscape patterning in efforts to restore degraded habitats and maximize the delivery of ecosystem services.
