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Hierarchical Analysis of Genetic Differentiation

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Hierarchical Analysis of Genetic Differentiation A Hierarchical Analysis of Genetic Differentiation in a Montane Leaf Beetle Chrysomela aeneicollis (Coleoptera: Chrysomelidae) Author(s): Nathan Egan Rank Source: Evolution, Vol. 46, No. 4 (Aug., 1992), pp. 1097-1111 Published by: Society for the Study of Evolution Stable URL: http://www.jstor.org/stable/2409759 Accessed: 01/02/2010 19:13 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. 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Society for the Study of Evolution is collaborating with JSTOR to digitize, preserve and extend access to Evolution. http://www.jstor.org Evolution, 46(4), 1992, pp. 1097-1111 A HIERARCHICAL ANALYSIS OF GENETIC DIFFERENTIATION IN A MONTANE LEAF BEETLE CHRYSOMELA AENEICOLLIS (COLEOPTERA: CHRYSOMELIDAE) NATHAN EGAN RANK' Department of Zoology, University of California, Davis, CA 95616 USA Abstract. -Herbivorous insects that use the same host plants as larvae and adults can have a subdivided population structure that corresponds to the distribution of their hosts. Having a subdivided population structure favors local adaptation of subpopulations to small-scale environ- mental differences and it may promote their genetic divergence. In this paper, I present the results of a hierarchical study of population structure in a montane willow leaf beetle, Chrysomela aenei- collis (Coleoptera: Chrysomelidae). This species spends its entire life associated with the larval host (Salix spp.), which occurs in patches along high-elevation streams and in montane bogs. I analyzed the genetic differentiation of C. aeneicollis populations along three drainages in the Sierra Nevada mountains of California at five enzyme loci: ak-l, idh-2, mpi-J, pgi-J, and pgm-J, using recent modifications of Wright's F-statistics. My results demonstrated significant differentiation (FST = 0.043) among drainages that are less than 40 kilometers apart. One locus, pgi-1, showed much greater differentiation than the other four (FST = 0.412), suggesting that it is under natural selection. C. aeneicollis populations were also subdivided within drainages, with significant dif- ferentiation 1) among patches of willows (spanning less than three kilometers) and 2) in some cases, among trees within a willow patch. My results demonstrate that this species has the capacity to adapt to local environmental variation at small spatial scales. Key words. -Allozymes, Chrysomela aeneicollis, F-statistics, gene flow, genetic differentiation, insect population structure, natural selection, phosphoglucose isomerase. Received February 4, 1991. Accepted November 20, 1991. All speciesconsist of populationsthat vary ample, if a herbivorous insect lives on trees in size and degree of isolation from other that grow in distinct patches separated by populations. Local population sizes, migra- geographical barriers, a study of that insect's tion rates, and the patternsof matings with- population structure should include sam- in a population all affect a species' popu- ples from individual trees from several lation structure (Hartl and Clark, 1989; host-tree patches. The advantage of a hi- McCauleyand Eanes, 1987; Slatkin, 1987). erarchical study is that one can distinguish In many cases, local environmental varia- evolutionary processes such as isolation due tion causes naturalselection to operate dif- to geographic barriers, isolation by distance, ferently among local populations, and pop- and isolation resulting from the organism's ulations may differ genetically in response behavior itself. Until recently however, few to this naturalselection. Even in the absence studies have attempted to measure popu- of selection, genetic drift among small-sized lation structure in this way. populationsincreases their likelihood of be- Willow leaf beetles (Family Chrysomeli- coming genetically differentiated. On the dae) occur on willow shrubs or trees that other hand, gene flow tends to make pop- commonly grow in separated patches of ulations more genetically homogeneous boggy or moist habitat. These beetles feed (Slatkin, 1987). To measure the genetic on the same host species as adults and as structureof populations,Wright (1978) pro- larvae, and they usually overwinter below posed that studies should use a hierarchical their host tree or under the bark of dead sampling design that correspondsto spatial branches (Brown, 1956; Raupp and Denno, subdivisions of naturalpopulations. For ex- 1983; Rowell-Rahier, 1984; Pasteels et al., 1988; Smiley and Rank, 1986). Although ' Present address: Institute fuirterrestrische okologie, they can fly, most willow leaf beetles do so ETH-Zirich, Grabenstr. 3, CH-8952 Schlieren, SWIT- rarely (Brown, 1956). ZERLAND. Fax 41-1-731-0783. Because of their low vagility and because 1097 1098 NATHAN EGAN RANK C. aeneicollis populations are potentially RC ~ ~ ~ R subdivided at several levels. First, popula- tions are localized in isolated drainages that are separated by high-elevation ridges. Sec- ond, C. aeneicollis populations occur on SL000m separated patches of willows in bogs, on ta- 5km. 1. MapofthelBP B lus slopes, or along streams. Finally, within N LA these patches, the willows grow as distinct shrubs supporting populations of C. aenei- collis larvae and adults. Thus, for this spe- cies, which has a limited vagility, physical SL BP G barriers and the patchiness of its host plants may favor genetic subdivision at each spa- tial scale. I tested this hypothesis by sam- D E pling C. aeneicollis populations hierarchi- cally 1) among isolated drainages, 2) among patches of willows within a drainage (lo- calities), and 3) among individual willows within a locality. FIG. 1. Maps of the localities. Top /eft-locations MATERIALSAND METHODS of the three drainages in the eastern Sierra Nevada. The areas shaded in black indicate regions higher than Collection. - Chrysomela aeneicollis 3500 meters. Top right-locations of the three locali- adults from the previous year can be found ties in the Rock Creek drainage. Bottom left -locations in May and June, but adults from the pres- of the five localities near South Lake in the Bishop Creek drainage. Bottom right-locations of the seven ent generation do not emerge from their pu- localities in the Big Pine Creek drainage. pae until late August or early September. For this study, I collected newly emerged adult beetles on September 3-7, 1988. The they occur in distinct patches of host plants, beetles were collected along three drainages: one may predictthat willow leaf beetles have Big Pine Creek (37?7'N/118?29'W), Bishop a subdivided population structure. This Creek (370 1'N/118?32'W, populations subdivision may enhance their potential for sampled near South Lake), and Rock Creek local adaptationto small scale environmen- (37025'N/1 18044'W). These drainages are tal variation. Indeed, an electrophoretic isolated by barren ridges that constitute part study of a willow leaf beetle, Plagioderaver- of the Sierra crest. Big Pine Creek is sepa- sicolor, which was introduced to North rated from Bishop Creek by about 10 ki- America at the turn of the century, found lometers and one ridge, but populations in significantdifferentiation among patches of both of these drainages are separated from willows separated by several kilometers the Rock Creek populations by several ridg- (McCauley et al., 1988). This study also es and about 40 kilometers. I sampled bee- demonstrated significant variation among tles in a stepping stone fashion within each patches in average relatedness of individ- drainage at localities that were 0.3 to 5 ki- uals within larval feeding groups, which lometers apart (Fig. 1). In each locality, I suggests that genetic differentiationin av- sampled beetles from two to eight willows erage relatedness has occurred. separated by 2 to 150 meters (Table 1). Be- In this study, I used gel electrophoresisto cause these willows are shrubs, all of the quantify the population structureof a wil- branches could be sampled. All of the bee- low leaf beetle, Chrysomela aeneicollis, tles were collected from a single, salicin-rich which occurs at high elevations in the east- willow species, S. orestera,except for those ern Sierra Nevada of central California collected from six S. boothi individuals at (Smiley and Rank, 1986). Chrysomela locality BPf. The two willow species grew aeneicollisis native to westernNorth Amer- together at this locality. ica from the Yukon to California (Brown, Electrophoresis. -After collection, I 1956). At this southernextreme of its range, brought the beetles to the University of Cal- LEAF BEETLE
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