River food webs: an integrative approach to bottom-up flow webs, top-down impact webs, and trophic position.

The majority of food web studies are based on connectivity, top-down impacts, bottom-up flows, or trophic position (TP), and ecologists have argued for decades which is best. Rarely have any two been considered simultaneously. The present study uses a procedure that integrates the last three approaches based on taxon-specific secondary production and gut analyses. Ingestion flows are quantified to create a flow web and the same data are used to quantify TP for all taxa. An individual predator's impacts also are estimated using the ratio of its ingestion (I) of each prey to prey production (P) to create an I/P web. This procedure was applied to 41 invertebrate taxa inhabiting submerged woody habitat in a southeastern U.S. river. A complex flow web starting with five basal food resources had 462 flows >1 mg·m-2 ·yr-1 , providing far more information than a connectivity web. Total flows from basal resources to primary consumers/omnivores were dominated by allochthonous amorphous detritus and ranged from 1 to >50,000 mg·m-2 ·yr-1 . Most predator-prey flows were much lower (<50 mg·m-2 ·yr-1 ), but some were >1,000  mg·m-2 ·yr-1 . The I/P web showed that 83% of individual predator impacts were weak (<10%), whereas total predator impacts were often strong (e.g., 35% of prey sustained an impact >90%). Quantitative estimates of TP ranged from 2 to 3.7, contrasting sharply with seven integer-based trophic levels based on longest feeding chain. Traditional omnivores (TP = 2.4-2.9) played an important role by consuming more prey and exerting higher impacts on primary consumers than strict predators (TP ≥ 3). This study illustrates how simultaneous quantification of flow pathways, predator impacts, and TP together provide an integrated characterization of natural food webs.