Historical contingency and productivity effects on food-chain length.

Food-chain length (FCL) is a fundamental ecosystem attribute, integrating information on both food web composition and ecosystem processes. It remains untested whether FCL also reflects the history of community assembly known to affect community composition and ecosystem functioning. Here, we performed microcosm experiments with a copepod (top predator), two ciliate species (intermediate consumers), and bacteria (producers), and modified the sequence of species introduction into the microcosm at four productivity levels to jointly test the effects of historical contingency and productivity on FCL. FCL increased when the top predator was introduced last; thus, the trophic position of the copepod reflected assembly history. A shorter FCL occurred at the highest productivity level, probably because the predator switched to feeding at the lower trophic levels because of the abundant basal resource. Thus, we present empirical evidence that FCL was determined by historical contingency, likely caused by priority effects, and by productivity.