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High Levels of Gene Flow in the California Vole (Microtus Californicus) Are Consistent Across Spatial Scales, Western North American Naturalist Volume 70 Number 3 Article 3 10-11-2010 High levels of gene flow in the California olev (Microtus californicus) are consistent across spatial scales Rachel I. Adams Stanford University, Stanford, California, [email protected] Elizabeth A. Hadly Stanford University, Stanford, California, [email protected] Follow this and additional works at: https://scholarsarchive.byu.edu/wnan Recommended Citation Adams, Rachel I. and Hadly, Elizabeth A. (2010) "High levels of gene flow in the California vole (Microtus californicus) are consistent across spatial scales," Western North American Naturalist: Vol. 70 : No. 3 , Article 3. Available at: https://scholarsarchive.byu.edu/wnan/vol70/iss3/3 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Western North American Naturalist 70(3), © 2010, pp. 296–311 HIGH LEVELS OF GENE FLOW IN THE CALIFORNIA VOLE (MICROTUS CALIFORNICUS) ARE CONSISTENT ACROSS SPATIAL SCALES Rachel I. Adams1,2 and Elizabeth A. Hadly1 ABSTRACT.—Gene flow links the genetic and demographic structures of species. Despite the fact that similar genetic and demographic patterns shape both local population structure and regional phylogeography, the 2 levels of population connectivity are rarely studied simultaneously. Here, we studied gene flow in the California vole (Microtus californicus), a small-bodied rodent with limited vagility but high local abundance. Within a 4.86-km2 preserve in central California, genetic diversity in 6 microsatellites was high, and Bayesian methods indicated a single genetic cluster. However, indi- vidual-based genetic analysis detected a clear signal for isolation-by-distance (IBD) and fine-scale population structure. Mitochondrial cytochrome b sequencing revealed 11 unique haplotypes from the one local area where we sequenced 62 individuals. Phylogeographic analysis of these individuals combined with those sampled from the northern geographic range of the species (the range of the species spans western North America from southern Oregon to northern Mexico and is centered geographically within the state of California) again indicated a lack of structure but a signal for IBD. Pat- terns of gene flow thus are consistent across spatial scales: while dispersal of the California vole is limited across geo- graphic distance, there is nonetheless considerable movement across the landscape. We conclude that in this species, high local population abundances overcome the potential genetic and demographic effects of limited dispersal. Key words: fine-scale genetic structure, population genetics, phylogeography, Jasper Ridge Biological Preserve, cytochrome b, Bayesian analysis, SIMCOAL. The geographic distribution of genetic diver- individuals across space and time—the demo- sity in natural populations harbors a wealth of graphic characteristics at the different scales information on species’ evolutionary and eco - vary, and consequently the patterns may be logical histories, demographic parameters, and quite distinct. Nevertheless, the 2 levels of gene conservation statuses. Gene flow, mediated by flow are rarely studied concurrently. For a spe - the movement of reproductive individuals and cies with limited vagility and strong philopatry therefore genes across space, is a prominent but high abundance, expectations for signals determinant of the genetic composition and of gene flow may differ depending on spatial diversity within a species and ultimately of the scale considered. A strong signal of genetic species’ survival. The role of gene flow in shap- structure is expected on both a microgeographic ing the pattern of gene frequencies depends and macrogeographic scale for organisms with on ecological, historical, and demographic fac- limited vagility, high fidelity to natal site, and tors. Therefore, measurements of gene flow specific habitat requirements (Lowe et al. 2004). give insight into the behavior, migration, and However, species with high abundances have mating patterns of natural populations. Gene the potential to override factors limiting gene flow is traditionally considered at 2 different flow through sheer force of numbers, and temporal scales: for short distances and times, therefore, genetic differentiation on a microgeo- gene flow is a process, along with genetic drift, graphic scale, but not necessarily on a macro- mutation, and natural selection, that shapes geographic scale, would be lacking in these population genetic structure (Hartl and Clark species. We set out to test whether signals of 1997); at longer distances and times, gene flow, gene flow would be similar at different spatial and the lack thereof, determine a species’ phy- scales for a species characterized by low disper- logeographic pattern. sal, high philopatry, and large population sizes. While both local population structure and By using 2 genetic markers of variable muta- phylogeography ultimately study the same tion rates (microsatellites and cytochrome b) evolutionary process—that of movement of from the same Microtus californicus individuals 1Department of Biology, Stanford University, Stanford, CA 94305-5020. 2E-mail: [email protected] 296 2010] GENE FLOW ACROSS SPATIAL SCALES 297 in one locality and linking these data to a study Heske 1987, Salvioni and Lidicker 1995), and of the phylogeography of the species that used this variation is known to affect the genetic cytochrome b (Conroy and Neuwald 2008), we diversity and structure of populations. Using were able to examine genetic structure across allozyme data from M. californicus, Bowen a small scale and place the locality into the phy - (1982) found significant genetic differentiation logeographic structure of the species—effec- between populations separated by only 50–200 tively synthesizing patterns of gene flow at 2 m when population density was low. This differ- spatial scales. ential disappeared when density rebounded. A The California vole (Microtus californicus) is a similar recovery of connectivity during peri- small rodent (<100 g) whose demographic and ods of high density has been observed in another ecological features have been well characterized. vole, Arvicola terrestris (Berthier et al. 2006). Individual dispersal distances are estimated to However, other vole species show increased be small, at 50 m or less (e.g., Lidicker 1973, gene flow during periods of low density (Andrea - Bowen 1982), and philopatry high, with male ssen and Ims 2001, Hadly et al. 2004). home ranges slightly larger than female home On a larger spatial scale, Microtus californicus ranges (e.g., 180 m2 and 120 m2, respectively; has 17 distinct subspecies (Hall 1981) that span Heske 1987). California voles, like many other the range of the species and are based on pel - microtine rodents, undergo 2- to 4-year popula- age, skull shape, and body size (Kellogg 1918). tion cycles (Krebs 1966). In addition, the Cali- However, further work based on distinct mor- fornia vole is limited to areas of heavy ground phological differences, decreased fertility in cover, predominantly grasslands and oak wood- crosses, and allozyme variation has shown a lands, where it lives in underground burrows break between only 2 broad regions—northern but travels aboveground to forage for grasses, and southern California (Gill 1980). Recently, a herbs, and sedges. It breeds throughout the year, phylogeographic study of the California vole although reproduction peaks whenever food and corroborated this geographic barrier, reporting cover are most abundant. Litter size averages strong genetic differentiation between popula- 4 individuals but ranges from 1 to 9, and fe - tions on the northern and southern sides of the males produce 2–5 litters per year. Females Transverse Mountain range in southern Cali- reach sexual maturity at an average of 29 days. fornia (Conroy and Neuwald 2008). The California vole’s circadian activity, along Thus, previous work with California voles with its widespread and abundant distribution, indicates that barriers to gene flow occur on make it an important prey for carnivores. Al - temporal scales as well as local and regional though it has a broad range, from Baja Califor- spatial scales, but never before have the same nia throughout western and central California individuals been used to determine gene flow to southern Oregon, the vole’s populations are at the 2 spatial scales simultaneously. For this naturally fragmented into suitable habitat, which goal, our study sampled heavily in one locality, is patchily distributed across this range. the Jasper Ridge Biological Preserve (JRBP), The particular biology of the California vole San Mateo County, located in the San Francisco and other voles leads to the presence of popu- Bay region of California. The preserve itself lation structure on a small spatial scale due to represents habitat within an increasingly de - habitat patchiness and landscape features (Neu- veloped area, and JRBP is varied in particular wald 2010, Berthier et al. 2005, Gauffre et al. landscape features. The preserve contains both 2008), as well as demography (Bowen 1982, native and nonnative grasslands. Nonnative Berthier et al. 2006, Gauffre et al. 2008). In grasses, predominantly of Mediterranean origin, southern California,
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