Food Web Complexity and Community Dynamics

Food webs in nature have multiple, reticulate connections between a diversity of consumers and resources. Such complexity affects web dynamics: it first spreads the direct effects of consumption and productivity throughout the web rather than focusing them at particular "trophic levels." Second, consumer densities are often donor controlled with food from across the trophic spectrum, the herbivore and detrital channels, other habitats, life-history omnivory, and even trophic mutualism. Although consumers usually do not affect these resources, increased numbers often allow consumers to depress other resources to levels lower than if donor-controlled resources were absent. We propose that such donor-controlled and "multichannel" omnivory is a general feature of consumer control and central to food web dynamics. This observation is contrary to the normal practice of inferring dynamics by simplifying webs into a few linear "trophic levels," as per "green world" theories. Such theories do not accommodate common and dynamically important features of real webs such as the ubiquity of donor control and the importance and dynamics of detritus, omnivory, resources crossing habitats, life history, nutrients (as opposed to energy), pathogens, resource defenses, and trophic symbioses. We conclude that trophic cascades and top-down community regulation as envisioned by trophic-level theories are relatively uncommon in nature.