Host-parasite nestedness: A result of co-evolving trait-values

Nestedness is an intriguing feature of ecological networks, where those species found in species-depauperate communities are subsets of those found in communities with greater species richness. For bipartite interaction networks, a " community" of species may be thought of as all those pollinating a particular plant, or infecting a particular host, for example. While there is much clear evidence for nestedness in mutualistic webs, host-parasite webs have proven more contentious. There have been a number of suggested causes for nestedness, including an association between the abundance of individuals and the resulting number of species interactions, and the matching of phenotypic traits between species. Questions remain as to the relative importance of these driving factors, especially as host-parasite and mutualistic webs contain completely different interaction types.We propose a model motivated by both of the above factors, considering a trade-off in resources that a species faces in optimizing its transmission or defense. We construct a multi-species model in which both hosts and parasites have limited resources with which to attack or defend themselves from each other. We analyze the evolution of the manner in which they use these resources using adaptive dynamics, to arrive at a final species interaction matrix, which we then test for nestedness. A general model with m hosts and n microparasite species is described here, but results are given for m= n= 5, chosen to be a large enough system for patterns to be identified, but not so large that computational time becomes prohibitive.Our results demonstrate that this co-evolution leads to an unusual amount of nestedness when the trade-offs in transmission for parasites are concave, and an unusual amount of anti-nestedness when they are convex. This enables us to predict the circumstances under which we would expect to observe nestedness in real networks.