Abstract. Studies of patterns of molecular variation in natural populations can provide important insights into a number of evolutionary problems. Among these, the question of whether geographic factors are more important than ecological factors in promoting population differentiation and ultimately speciation has been an important and contentious area in evolutionary biology. Systems involving herbivorous insects have played a leading role in this discussion. This study examined the distribution of molecular variation in a highly specialized gall-forming insect, grape phylloxera (Daktulosphaira vitifoliae Fitch), that is found on both sympatric and allopatric host-plant species of the genus Vitis. In addition, the relationship of insects in the introduced range in the United States to ancestral populations in the native range was examined. Evidence for differentiation along host-plant lines from both nuclear (RAPD) and mitochondrial (COI) DNA was confounded with the effect of geography. Differentiation was found where hosts were allopatric or parapatric, but no evidence was found for such differentiation on two hosts, V. vulpina and V. aestivalis, that are broadly sympatric. The question of population differentiation onto these sympatric hosts can be considered to be resolved-it has not occurred in spite of a long history of association. Evidence was equivocal, but suggestive of a period of divergence in allopatry prior to reestablishment of contact, for insects associated with another host plant species, V. cinerea, found in both sympatric and parapatric populations. A low level of diversity and placement of samples collected from the grape species V. riparia at the tip of a phylogenetic tree supports the hypothesis that this host has been recently colonized from populations from the Mississippi Valley. A polyphyletic origin for biotype B grape phylloxera was supported: Although most samples collected from vineyards in the introduced range in California had similar haplotypes, they were closely related to natives on V. vulpina from the Atlantic CoastPiedmont region. All samples collected from vineyards in Oregon and Washington were closely related to natives on V. riparia in the northern United States.Key words. Biotypes, cytochrome oxidase I, geographic distribution, grape phylloxera, herbivorous insects, hostplant variation, sympatric divergence, Vitis. Geographical features of the environment of organisms that prevent or reduce gene flow are widely considered to be better predictors of genetic differentiation of populations than are ecological factors (Dobzansky 1937;Mayr 1963;Futuyma and Mayer 1980;Patterson 1981;Templeton 1981;Carson 1982;Barton and Charlesworth 1984;Lynch 1989;Allmon 1992;Avise 2000;Barraclough and Vogler 2000), but the consensus on this issue is far from complete. Increasing theoretical support for the sympatric model of speciation has accumulated over the years and many evolutionary biologists now feel quite comfortable with this model (Maynard