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Food webs are complex. Complexity is a measure of an increasing number of permutations and it is also a metaphorical term that conveys the mental intractability or limits concerning unlimited algorithmic possibilities. In food web terminology, complexity is a product of the number of species and connectance. Connectance is the fraction of all possible links that are realized in a network. These concepts were derived and stimulated through the suggestion that complexity leads to stability in food webs, such as increasing the number of trophic levels in more species rich ecosystems. This hypothesis was challenged through mathematical models suggesting otherwise, but subsequent studies have shown that the premise holds in real systems.
At different levels in the hierarchy of life, such as the stability of a food web, the same overall structure is maintained in spite of an ongoing flow and change of components. The farther a living system (e.g., ecosystem) sways from equilibrium, the greater its complexity. Complexity has multiple meanings in the life sciences and in the public sphere that confuse its application as a precise term for analytical purposes in science. Complexity in the life sciences (or biocomplexity) is defined by the properties emerging from the interplay of behavioral, biological, physical, and social interactions that affect, sustain, or are modified by living organisms, including humans.
Several concepts have emerged from the study of complexity in food webs. Complexity explains many principals pertaining to self-organization, non-linearity, interaction, cybernetic feedback, discontinuity, emergence, and stability in food webs. Nestedness, for example, is defined as a pattern of interaction in which specialists interact with species that form perfect subsets of the species with which generalists interact—that is, the diet of the most specialized species is a subset of the diet of the next more generalized species, and its diet a subset of the next more generalized, and so on. Until recently, it was thought that food webs had little nested structure, but empirical evidence shows that many published webs have nested subwebs in their assembly.
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