Evidence for ecological speciation via a host shift in the holly leaf miner, Phytomyza glabricola (Diptera: Agromyzidae).

Evolutionary radiations have been well documented in plants and insects, and natural selection may often underly these radiations. If radiations are adaptive, the diversity of species could be due to ecological speciation in these lineages. Agromyzid flies exhibit patterns of repeated host-associated radiations. We investigated whether host-associated population divergence and evidence of divergent selection exist in the leaf miner Phytomyza glabricola on its sympatric host plants, the holly species, Ilex coriacea and I. glabra. Using AFLPs and nuclear sequence data, we found substantial genetic divergence between host-associated populations of these flies throughout their geographic range. Genome scans using the AFLP data identified 13 loci under divergent selection, consistent with processes of ecological speciation. EF-1α data suggest that I. glabra is the original host of P. glabricola and that I. coriacea is the novel host, but the AFLP data are ambiguous with regard to directionality of the host shift.

Reproductive isolation between host races of Phytomyza glabricola on Ilex coriacea and I. glabra.

Recently diverged taxa often show discordance in genetic divergence among genomic loci, where some loci show strong divergence and others show none at all. Genetic studies alone cannot distinguish among the possible mechanisms but experimental studies on other aspects of divergence may provide guidance in the inference of causes of observed discordances. In this study, we used no-choice mating trials to test for the presence of reproductive isolation between host races of the leaf-mining fly, Phytomyza glabricola on its two holly host species, Ilex coriacea and I. glabra. These trials inform our effort to determine the cause of significant differences in the degree of divergence of nuclear and mitochondrial loci of flies collected from the two host plants. We present evidence of reproductive isolation between host races in a controlled greenhouse setting: significantly more mate pairs consisting of flies from the same host plant species produced offspring than inter-host mate pairs, which produced no offspring. We also tested whether the presence of the natal or non-natal host plant affects reproductive success. Flies collected from I. coriacea were more likely to produce offspring when in the presence of the natal host, whereas the presence or absence of either the natal or non-natal host had no effect on flies collected from I. glabra. The results indicate discordant patterns of nuclear and mitochondrial divergence among host races of P. glabricola are likely due to incomplete lineage sorting, and the host races may be well on their way to becoming biological species.

Herbivores affect natural selection for floral-sex ratio in a field population of horsenettle, Solanum carolinense.

Although aspects of a plant's breeding system are generally believed to have evolved in response to selection for effective pollination, herbivores may also play a selective role. Here we report on a field experiment involving 960 transplanted ramets of the andromonoecious herb Solanum carolinense in which the pattern of natural selection for an important breeding-system trait was influenced by naturally occurring herbivores. As the level of flower and fruit herbivory increased, the pattern of selection on floral-sex ratio went from stabilizing (with an optimum of 29% male flowers), to directional toward a lower proportion of males, with the optimum bottoming at 0% male. This pattern of selection likely helped generate the broad-sense genetic correlation (r = 0.42) between floral-sex ratio and resistance to herbivory. These results contribute to the growing awareness that herbivores can be important influences not only on plant resistance traits, but also on the evolution of their hosts' breeding system.