• Assembly rules;
  • co-occurrence;
  • extinction;
  • fragmentation;
  • habitat suitability;
  • piscivory;
  • predation;
  • winter kill


Diamond [Assembly of species communities. In: Ecology and Evolution of Communities (eds Cody, M.L. & Diamond, J.M.). Harvard University Press, Cambridge, pp. 342–444] proposed that resource competition leads to checkerboard-like distributions of competing species. This proposal prompted research that revealed checkerboard patterns within a wide range of communities, but the mechanisms that generate such patterns are still poorly understood. Here we present whole-lake natural experiments and analyses of species–environment relationships in small coastal lake fish communities that were fragmented when land uplift isolated these lakes from the Baltic Sea, showing that a combination of predation and habitat suitability generated checkerboard distributions. Checkerboard patterns developed because two piscivores, northern pike and Eurasian perch, caused the extinction of several prey species in deep lakes. Conversely, low oxygen levels in shallow lakes caused extinction of the piscivores, and these areas served as a refuge for tolerant prey species. Based on these findings, we suggest that habitat suitability and biotic interactions should be viewed simultaneously in null models of assembly rules.