Speciation in many host specific parasites may be initiated in sympatry when populations shift and adapt to new hosts. The recent shift of the apple maggot fly, Rhagoletis pomonella (Diptera: Tephritidae) from its native host plant hawthorn (Crataegus spp.) to introduced, domesticated apple (Malus pumila), provides a direct test of the "sympatric speciation" hypothesis by indicating whether partially reproductively isolated "host races" can evolve in the absence of geographic isolation. We report finding significant allele frequency differences for six allozymes between paired apple and hawthorn infesting populations of R. pomonella from across the midwestern United States. Latitudinal allele frequency dines exist among both apple and hawthorn populations, however, for a majority of the loci displaying racial differences. Inter-host genetic differentiation is therefore superimposed on clinal patterns of variation within the races such that the magnitude of host associated divergence is a function of latitude. The results indicate that host associated races can form in sympatry and implicate differences in host plant recognition and developmental timing (related to ambient temperature) as key factors restricting gene flow between apple and hawthorn populations. However, some of the same processes differentiating apple and hawthorn populations at sympatric sites also appear to be occurring within the two host races across their respective ranges. R. pomonella populations are therefore diverging with respect to both their host plant affiliations and local environmental conditions.
INTRODUCTIONSpeciation, the rendering of one closed genetic system into two, is a fundamental process responsible for the evolution of life on earth. Unfortunately, because speciation is a relatively rare, often time consuming event, much of our understanding of the process is based on population genetics models and indirect analyses of closely related species. We therefore have very few examples directly documenting how populations become reproductively isolated from one another.The apple maggot fly, Rhagoletis pomonella, is an exception, however, having a biology and natural history which makes it ideal for the empirical study of speciation. R. pomonella is part of an endemic, sibling species complex of true fruit flies whose members overlap broadly in their geographic distributions across North America (Bush, 1966). Rhagoletis larvae are internal parasites in the fruit of their hosts, with each species in the group being monophagous or oligophagous for a different set of host plants (Bush, 1966). Adult females lay their eggs directly into the host fruit which they identify by specific visual, tactile and olfactory cues (Prokopy, 1968a;Prokopy et a!., 1973Prokopy et a!., , 1987Prokopy et a!., , 1988Moericke et a!., 1975;Fein et a!., 1982;Owens and Prokopy, 1986;Papaj and Prokopy, 1986). Males are attracted by the same cues and courtship and mating occurs almost exclusively on or near the fruits of the host plant (Prokopy et a!., 1971(Pr...