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224 Ecological Networks Beyond Food Webs


A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Ecologists can broadly define all life forms as either autotrophs or heterotrophs, based on their trophic levels, the position that they occupy in the food web. To maintain their bodies, grow, develop, and to reproduce, autotrophs produce organic matter from inorganic substances, including both minerals and gases such as carbon dioxide. These trophic levels are not binary, but form a gradient that includes complete autotrophs, which obtain their sole source of carbon from the atmosphere, mixotrophs (such as carnivorous plants), which are autotrophic organisms that partially obtain organic matter from sources other than the atmosphere, and complete heterotrophs that must feed to obtain organic matter. The linkages in a food web illustrate the feeding pathways, such as where heterotrophs obtain organic matter by feeding on autotrophs and other heterotrophs. The food web is a simplified illustration of the various methods of feeding that link an ecosystem into a unified system of exchange.


There are different kinds of consumer-resource interactions that can be roughly divided into herbivory, carnivory, scavenging, and parasitism. Some of the organic matter eaten by heterotrophs, such as sugars, provides energy. Autotrophs and heterotrophs come in all sizes, from microscopic to many tonnes - from cyanobacteria to giant redwoods, and from viruses and bdellovibrio to blue whales. Charles Elton pioneered the concept of food cycles, food chains, and food size in his classical 1927 book "Animal Ecology"; Elton's 'food cycle' was replaced by 'food web' in a subsequent ecological text. Elton organized species into functional groups, which was the basis for Raymond Lindeman's classic and landmark paper in 1942 on trophic dynamics. Lindeman emphasized the important role of decomposer organisms in a trophic system of classification. The notion of a food web has a historical foothold in the writings of Charles Darwin and his terminology, including an "entangled bank", "web of life", "web of complex relations", and in reference to the decomposition actions of earthworms he talked about "the continued movement of the particles of earth".


Even earlier, in 1768 John Bruckner described nature as "one continued web of life". Food webs are limited representations of real ecosystems as they necessarily aggregate many species into trophic species, which are functional groups of species that have the same predators and prey in a food web. Ecologists use these simplifications in quantitative (or mathematical representation) models of trophic or consumer-resource systems dynamics. Using these models they can measure and test for generalized patterns in the structure of real food web networks. Ecologists have identified non-random properties in the topological structure of food webs. Published examples that are used in meta analysis are of variable quality with omissions. However, the number of empirical studies on community webs is on the rise and the mathematical treatment of food webs using network theory had identified patterns that are common to all. Scaling laws, for example, predict a relationship between the topology of food web predator-prey linkages and levels of species richness.


Food webs are the road-maps through Darwin's famous 'entangled bank' and have a long history in ecology. Like maps of unfamiliar ground, food webs appear bewilderingly complex. They were often published to make just that point. Yet recent studies have shown that food webs from a wide range of terrestrial, freshwater, and marine communities share a remarkable list of patterns. Links in food webs map the feeding connections (who eats whom) in an ecological community. Food cycle is an obsolete term that is synonymous with food web. Ecologists can broadly group all life forms into one of two trophic layers, the autotrophs and the heterotrophs. Autotrophs produce more biomass energy, either chemically without the sun's energy or by capturing the sun's energy in photosynthesis, than they use during metabolic respiration. Heterotrophs consume rather than produce biomass energy as they metabolize, grow, and add to levels of secondary production. A food web depicts a collection of polyphagous heterotrophic consumers that network and cycle the flow of energy and nutrients from a productive base of self-feeding autotrophs.


The base or basal species in a food web are those species without prey and can include autotrophs or saprophytic detritivores (i.e., the community of decomposers in soil, biofilms, and periphyton). Feeding connections in the web are called trophic links. The number of trophic links per consumer is a measure of food web connectance. Food chains are nested within the trophic links of food webs. Food chains are linear (noncyclic) feeding pathways that trace monophagous consumers from a base species up to the top consumer, which is usually a larger predatory carnivore. Linkages connect to nodes in a food web, which are aggregates of biological taxa called trophic species. Trophic species are functional groups that have the same predators and prey in a food web. Common examples of an aggregated node in a food web might include parasites, microbes, decomposers, saprotrophs, consumers, or predators, each containing many species in a web that can otherwise be connected to other trophic species.



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