FISHES AS INTEGRATORS OF BENTHIC AND PELAGIC FOOD WEBS IN LAKES

Studies of lake ecosystems generally focus on pelagic food chains and processes. Recently, there has been an emerging recognition of the importance of benthic production and processes to whole-lake ecosystems. To examine the extent to which zoobenthos contribute to higher trophic level production in lakes, we synthesized diet data from 470 fish populations (15 species) and stable isotope data from 90 fish populations (11 species), all of which are common inhabitants of north-temperate lakes. Across all species considered, zoobenthos averaged 50% of total prey consumption. Indirect consumption of zoobenthos (i.e., feeding on zoobenthos-supported fishes) contributed another 15%, for a total of 65% reliance on benthic secondary production. Stable isotopes provided estimates of mean zoobenthivory ranging from 43% to 59%. For most fish species, consumption of zoobenthos was highly variable among populations. The overwhelming concern of ecologists with pelagic food chains and processes contrasts sharply with our finding that benthic secondary production plays a central role in supporting higher trophic level production. This extensive zoobenthivory can subsidize fish populations, leading to apparent competition and otherwise altering trophic dynamics and ecosystem processes in the pelagic zone. We argue for a more integrated view of lake ecosystems that recognizes the duality of benthic and pelagic production pathways. Food web models that explicitly consider energy flow from pelagic and benthic sources will provide a more realistic energy flow template for understanding the regulation of lake ecosystem functioning.