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A central but little-tested prediction of ''escape and radiation'' coevolution is that colonization of novel, chemically defended host plant clades accelerates insect herbivore diversification. That theory, in turn, exemplifies one side of a broader debate about the relative influence on clade dynamics of intrinsic (biotic) vs. extrinsic (physical-environmental) forces. Here, we use a fossil-calibrated molecular chronogram to compare the effects of a major biotic factor (repeated shift to a chemically divergent host plant clade) and a major abiotic factor (global climate change) on the macroevolutionary dynamics of a large Cenozoic radiation of phytophagous insects, the leaf-mining fly genus Phytomyza (Diptera: Agromyzidae). We find one of the first statistically supported examples of consistently elevated net diversification accompanying shift to new plant clades. In contrast, we detect no significant direct effect on diversification of major global climate events in the early and late Oligocene. The broader paleoclimatic context strongly suggests, however, that climate change has at times had a strong indirect influence through its effect on the biotic environment. Repeated rapid Miocene radiation of these flies on temperate herbaceous asterids closely corresponds to the dramatic, climatedriven expansion of seasonal, open habitats.adaptive radiation ͉ coevolution ͉ climate change ͉ macroevolution ͉ Agromyzidae

The diversity of tropical herbivorous insects has been explained as a direct function of plant species diversity. Testing that explanation, we reared 2857 flies from flowers and seeds of 24 species of plants from 34 neotropical sites. Samples yielded 52 morphologically similar species of flies and documented highly conserved patterns of specificity to host taxa and host parts. Widespread species of plants can support 13 species of flies. Within single populations of plants, we typically found one or more fly species specific to female flowers and multiple specialists on male flowers. We suggest that neotropical herbivorous insect diversity is not simply a function of plant taxonomic and architectural diversity, but also reflects the geographic distribution of hosts and the age and area of the neotropics.

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Phylogenetic relationships among populations of the polyphagous pea leafminer, Liriomyza huidobrensis (Blanchard), were investigated using DNA sequence data. Maximum parsimony analysis of 941 bp of mitochondrial cytochrome oxidase I and II genes showed that L. huidobrensis contains two well-defined monophyletic groups, one composed of specimens from California and Hawaii and one composed of specimens from South and Central America together with populations that have been recently introduced into other parts of the world. The differentiation between the two clades within L. huidobrensis is equivalent to that seen between other agromyzid species, suggesting that L. huidobrensis as currently defined contains two cryptic species. This finding is consistent with field observations of differences in pest status and insecticide resistance between L. huidobrensis populations. Until additional studies are complete, no changes in L. huidobrensis taxonomy are proposed. However, researchers and quarantine officials may wish to consider the findings of the current study in designing research, pest management, and quarantine programs for L. huidobrensis.

We ask whether patterns of genetic variation in a phytophagous insect's responses to potential host plants shed light on the phylogenetic history of host association. Ophraella communa feeds chiefly, and in eastern North America exclusively, on Ambrosia (Asteraceae: Ambrosiinae). Using mostly half‐sib breeding designs, we screened for genetic variation in feeding responses to and larval survival on its own host and on seven other plants that are hosts (or, on one case, closely related to the host) of other species of Ophraella. We found evidence for genetic variation in feeding responses to five of the seven test plants, other than the natural host. We found no evidence of genetic variation in feeding responses to two plant species, nor in capacity for larval survival on six. These results imply constraints on the availability of genetic variation; however, little evidence for constraints in the form of negative genetic correlations was found. These results are interpreted in the context of a provisional phylogeny of, and a history of host shifts within, the genus. Ophraella communa does not present evidence of genetic variation in its ability to feed and/or survive on Solidago, even though it is probably descended from a lineage that fed on Solidago or related plants, possibly as recently as 1.9 million years ago. Genetic variation in performance on this plant may have been lost. Based on evidence for genetic variation and on mean performance, by far the greatest potentiality for adaptation to a congener's host was evinced in responses to Iva frutescens, which not only is related and chemically similar to Ambrosia, but also is the host of a closely related species of Ophraella that may have been derived from an Ambrosia‐associated ancestor. Genetic variation in O. communa's capacity to feed and/or survive on its congeners' hosts is less evident for plants that do not represent historically realized host shifts (with one exception) than for those that may (but see Note Added in Proof). The results offer some support for the hypothesis that the evolution of host shifts has been guided in part by constrained genetic variation.

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