Ecosystem size determines food-chain length in lakes

Food-chain length is an important characteristic of ecological communities(1) : it influences community structure(2), ecosystem functions(1-4) and contaminant concentrations in top predators(5,6). Since Elton(7) first noted that food-chain length was variable among natural systems, ecologists have considered many explanatory hypotheses(1,4,8,9), but few are supported by empirical evidence(4,10,11). Here we test three hypotheses that predict food-chain length to be determined by productivity alone (productivity hypothesis)(4,10,12,13), ecosystem size alone (ecosystem-size hypothesis)(14,15) or a combination of productivity and ecosystem size (productive-space hypothesis)(7,16-18). The productivity and productive-space hypotheses propose that food-chain length should increase with increasing resource availability; however, the productivity hypothesis does not include ecosystem size as a determinant of resource availability. The ecosystem-size hypothesis is based on the relationship between ecosystem size and species diversity, habitat availability and habitat heterogeneity(14,15). We find that food-chain length increases with ecosystem size, but that the length of the food chain is not related to productivity. Our results support the hypothesis that ecosystem size, and not resource availability, determines food-chain length in these natural ecosystems.