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Genetic Sampling of Palmer's Chipmunks in the Spring Mountains, Nevada

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Genetic Sampling of Palmer's Chipmunks in the Spring Mountains, Nevada Western North American Naturalist Volume 73 Number 2 Article 8 7-5-2013 Genetic sampling of Palmer's chipmunks in the Spring Mountains, Nevada Kevin S. McKelvey USDA Forest Service, Missoula, MT, [email protected] Jennifer E. Ramirez USDA Forest Service, Las Vegas, NV, [email protected] Kristine L. Pilgrim USDA Forest Service, Missoula, MT, [email protected] Samuel A. Cushman USDA Forest Service, Flagstaff, AZ, [email protected] Michael K. Schwartz USDA Forest Service, Missoula, MT, [email protected] Follow this and additional works at: https://scholarsarchive.byu.edu/wnan Part of the Anatomy Commons, Botany Commons, Physiology Commons, and the Zoology Commons Recommended Citation McKelvey, Kevin S.; Ramirez, Jennifer E.; Pilgrim, Kristine L.; Cushman, Samuel A.; and Schwartz, Michael K. (2013) "Genetic sampling of Palmer's chipmunks in the Spring Mountains, Nevada," Western North American Naturalist: Vol. 73 : No. 2 , Article 8. Available at: https://scholarsarchive.byu.edu/wnan/vol73/iss2/8 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 73(2), © 2013, pp. 198–210 GENETIC SAMPLING OF PALMER’S CHIPMUNKS IN THE SPRING MOUNTAINS, NEVADA Kevin S. McKelvey1,4, Jennifer E. Ramirez2, Kristine L. Pilgrim1, Samuel A. Cushman3, and Michael K. Schwartz1 ABSTRACT.—Palmer’s chipmunk (Neotamias palmeri) is a medium-sized chipmunk whose range is limited to the higher- elevation areas of the Spring Mountain Range, Nevada. A second chipmunk species, the Panamint chipmunk (Neotamias panamintinus), is more broadly distributed and lives in lower-elevation, primarily pinyon-juniper (Pinus monophylla–Junipe- rus osteosperma) habitat types. Panamint chipmunks are not closely related to Palmer’s, but field identification of the 2 species is unreliable. Palmer’s chipmunk is a species of concern in the state of Nevada and is listed by the International Union for Conservation of Nature (IUCN) as endangered. As such, conservation of Palmer’s chipmunks is a priority in the Spring Mountains National Recreation Area. We sampled putative Palmer’s chipmunks from 13 sites distributed across the Spring Mountains during 2010–2011. We removed Panamint chipmunks by using DNA-based identifications and then ana- lyzed the genetic population structure of Palmer’s chipmunks by using a panel of 9 microsatellites. Of the 228 samples that were genotyped, 186 were Palmer’s; there was no evidence of hybridization between species. Four sites had exclusively Panamint chipmunks, 5 had exclusively Palmer’s chipmunks, and 3 had a mixture of the 2 species. In this study, Palmer’s chipmunks were exclusively captured at sites above 2400 m elevation, and Panamint chipmunks were exclusively captured at sites below 2200 m. Panamint chipmunks were trapped in areas typed as pinyon-juniper, but they were also trapped at sites typed as ponderosa pine (Pinus ponderosa) and mixed conifer. Both species were trapped at 3 sites; at all 3 sites, the lower- elevation traps contained Panamint chipmunks and the higher ones Palmer’s chipmunks. Population structure within Palmer’s chipmunks was minimal: heterozygosity was relatively high, and the populations displayed no signs of recent bot- tlenecks. Indications are that the distribution of Palmer’s chipmunk is limited to higher-elevation areas in the Spring Moun- tains, but within this area, Palmer’s chipmunk occurs as a single, large, well-connected, and stable population. RESUMEN.—Neotamias palmeri es una ardilla de tamaño mediano cuyo hábitat se limita a las áreas más elevadas de Spring Mountain Range, Nevada. Una segunda especie de ardilla, Neotamias panamintinus, tiene una distribución más amplia y vive en áreas más bajas, principalmente en ambientes de pino piñonero (Pinus monophylla–Juniperus osteosperma). Si bien Neotamias panamintinus no se relaciona con Neotamias palmeri, la identificación de ambas especies en el terreno no es confiable. Neotamias palmeri es una especie en riesgo en el estado de Nevada y está incluida en la lista de especies en peligro de extinción de la Unión Internacional para la Conservación de la Naturaleza (IUCN), por lo tanto, la conservación de esta especie es una prioridad en Spring Mountains National Recreation Area. Entre el 2010 y el 2011 tomamos muestras de supuestas Neotamias palmeri en 13 lugares ubicados en Spring Mountains. Eliminamos Neotamias panamintinus identifi- cando con muestras de ADN y analizamos la estructura genética de la población de Neotamias palmeri utilizando un panel de 9 microsatélites. De los genotipos de las 228 muestras que tomamos, 186 correspondían a Neotamias palmeri y no se encontraron rastros de hibridación entre las especies. En cuatro lugares se encontraban exclusivamente Neotamias pana - mintinus, en cinco exclusivamente Neotamias palmeri y en tres se encontró una mezcla de las dos especies. En este estudio, se capturaron únicamente Neotamias palmeri en lugares con una elevación >2400 m y se capturaron únicamente Neotamias panamintinus en lugares con una elevación <2200 m. La especie Neotamias panamintinus se atrapó en ambientes de pino piñonero, en ambientes de pino ponderosa (Pinus ponderosa) y también en bosque mixto de coníferas. Ambas especies se encontraron en tres lugares, en los cuales las trampas ubicadas en zonas menos elevadas contenían Neotamias panamintinus y las ubicadas en zonas más elevadas contenían Neotamias palmeri. La estructura de la población en la especie Neotamias palmeri era mínima, la heterocigosidad era relativamente elevada y las poblaciones no mostraron signos de cuellos de botella recientes. Se observó que Neotamias palmeri se limita a áreas de mayor elevación en Spring Mountains, dentro de dicha área, esta especie constituye una población única, de tamaño considerable, estable y sin aislamiento entre sus miembros. Palmer’s chipmunk (Neotamias palmeri) is Uinta chipmunk (N. umbrinus) and the grey- a medium-sized chipmunk in the “Dorsalis” collared chipmunk (N. cinereicollis), which col - clade (Banbury and Spicer 2007). This clade lectively are the Palmer’s chipmunk’s closest also contains the cliff chipmunk (N. dorsalis), relatives (Piaggio and Spicer 2001). Several of 1USDA Forest Service, Rocky Mountain Research Station, 800 East Beckwith, Missoula, MT 59801. 2USDA Forest Service, Spring Mountains National Recreation Area, Humboldt–Toiyabe National Forest, 4701 North Torrey Pines Drive, Las Vegas, NV 89130-2301. 3USDA Forest Service, Rocky Mountain Research Station, 2500 S. Pine Knoll Rd., Flagstaff, AZ 86001. 4E-mail: [email protected] 198 2013] PALMER’S CHIPMUNK GENETICS 199 these species are widely distributed in the west- isms, describing hybridization patterns, evalu- ern United States, yet the only chipmunk other ating population structure and connectivity, than Palmer’s found in the Spring Moun tain and evaluating the likelihood of recent popula- Range, Clark County, Nevada, is the Pana mint tion contractions (bottlenecks). Interpretation chipmunk (N. panamintinus), a smaller and dis- of genetic results is greatly simplified for iso- tantly related (Piaggio and Spicer 2001) chip- lated species where the organism’s range is munk. Few published studies discuss Palmer’s well understood. Thus, a species like Palmer’s chipmunk, but it has been re ported as limited chipmunk that can be confused with a sym- to higher elevation (2250–3600 m) areas in the patric species and has likely been isolated for Spring Mountains (Hall and Kelson 1959); Wil- thousands of years is particularly amenable to son and Ruff (1999) re port the lower elevation genetic analyses. limit as 2100 m. A recent unpublished study Assuming that Palmer’s chipmunk is en - (Lowrey and Longshore 2011) reported the demic to the higher-elevation areas in the capture of Palmer’s chipmunks at elevations Spring Mountains, its vulnerability will largely ranging from 2080 m to 3290 m. Panamint be related to its population size and connectivity chipmunks are associated with pinyon-juniper across the Spring Mountains. The area where (Pinus monophylla–Juniperus osteosperma) for - Palmer’s chipmunks are known to occur is ex - ests (Lowrey and Longshore 2011) and, in the tensive (>80 km2) and contains potential bar- Spring Mountains, are associated with lower- riers, such as alpine ridges. If local popula- elevation sites (1600–2400 m; Hall and Kelson tions within specific canyons are isolated, then 1959). Lowrey and Longshore (2011) caught genetic drift and lack of gene flow will produce Panamint chipmunks at elevations up to 2643 m. genetic structure at neutral loci when evaluated The marked differences in niche characteris- across the extent of the range. Even with moder- tics be tween these 2 species may be due to ate levels of connectivity, we would expect to see adaptive differentiation. For ex ample, in labo- some genetic population structure and “isola- ratory experiments, Hirshfeld and Bradley tion by distance,” meaning that the population is (1977) found that Palmer’s chip munks exhibit not structured by geographic barriers, but rather faster growth than do Pana mint chipmunks. simply by geographic distance (Wright 1943). The authors attributed this difference in growth Because Palmer’s chipmunk is isolated in a to a longer period of hibernation
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