Concepts Of Coevolution

Coevolution refers to several processes. One possible form of coevolution is cospeciation, the coordinated branching (speciation) of interacting species (such as host and parasite). To the extent that this has occurred, concordant (or matching) phylogenies of host and parasite clades (or evolutionary lines) would be expected. Cospeciation might be caused by the interaction between species, but it could also result from a joint history of geographic isolation, assuming that divergence and reproductive isolation evolve at similar rates in the two groups. Concordance of the two phylogenies implies a longer history of association, and of opportunity for reciprocal adaptation, than, for example, when parasites or symbionts have frequently switched from one host to another. Host switching can be inferred from certain patterns of discordance between host and symbiont phylogenies. Both cospeciation and host switching have been revealed in herbivorous insects, symbiotic bacteria, and parasites. For example, lice associated with gophers and with certain seabirds appear to have cospeciated to a considerable extent, and endosymbiotic, mutualistic bacteria (Buchnera) display almost complete phylogenetic concordance with their aphid hosts, from the family level down through relationships among conspecific populations.

In its most frequent usage, coevolution refers to genetic changes in the characteristics of interacting species resulting from natural selection imposed by each on the other—i.e., reciprocal adaptation of lineages to each other. Such changes are referred to as specific or pairwise coevolution if the evolutionary responses of two species to each other have no impact on their interactions with other species. Diffuse or guild coevolution occurs when the genetic change in at least one species affects its interaction with two or more other species. For example, cucumber genotypes with high levels of the chemical cucurbitacin have enhanced resistance to mites but also enhanced attractiveness to cucumber beetles; this is an instance of a negative genetic correlation in resistance. Early season attack by flea beetles makes sumac plants more susceptible to stem-boring cerambycid beetles, and so resistance to the former would also reduce the impact of the latter.

In one of the seminal papers on coevolution, Ehrlich and Raven postulated in 1964 what has since been named "escape and radiate" coevolution—a process in which evolutionary changes temporarily reduce or eliminate the ecological interactions between species. Applying this concept to plants and herbivorous insects, Ehrlich and Raven postulated that in response to selection by herbivores, a plant species may evolve new defenses that enable it to escape herbivory and to flourish so well that it gives rise to a clade of descendant species with similar defenses. At some later time, one or more species of herbivores adapt to the defenses and give rise to an adaptive radiation of species that feed on the plant clade. In this scenario, the evolutionary diversification of both herbivores and plants is enhanced by their interactions.

Despite a common misconception, coevolution need not promote stable coexistence of species, and it certainly need not enhance mutual harmony. For example, parasites may evolve to become more virulent or less, depending on their life history. The Darwinian fitness of a genotype of parasite is measured by the average reproductive success of an individual of that genotype. Extracting more resource from a host, thereby reducing its chance of survival, often enhances the parasite's reproductive success, as long as the parasite individual, or its offspring, can escape to new hosts before the current host dies. Evolution of the parasite, by individual selection, may result in such high virulence that the prey or host population is extinguished. Extinction of prey populations does not alter the relative fitnesses of individual parasite genotypes and so does not select for reduced virulence. However, group selection may favor lower virulence or proficiency. If populations of more virulent parasites suffer higher extinction rates than less virulent populations, the species as a whole might evolve lower virulence. Although individual selection is likely to be stronger than group selection in most species, the population structure of some parasites may provide an opportunity for group selection to affect their evolution.

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