where, S(S-1)/2 is the maximum number of binary connections among S species. "Connectance (C) is the fraction of all possible links that are realized (L/S2) and represents a standard measure of food web complexity..." The distance (d) between every species pair in a web is averaged to compute the mean distance between all nodes in a web (D) and multiplied by the total number of links (L) to obtain link-density (LD), which is influenced by scale dependent variables such as species richness. These formulas are the basis for comparing and investigating the nature of non-random patterns in the structure of food web networks among many different types of ecosystems.