COLUMBIA SPOTTED luteiventris Thompson, 1913 (Ranidae)

State rank S2? (1999-09-30) Global rank G4 (1997-07-11) State rank reasons Restricted range in Southeast . Population status unknown, although suspected low. Needs more study. Major threats include loss/degradation due to development and glacial rebound.

isolation and habitat differences, including disparate biological, chemical, and thermal Formerly included in Rana pretiosa. Green et al. characteristics of occupied springs and wetlands (1996) examined allozyme and morphometric (Hovingh 1993, USFWS 1993). In addition, due to variation in R. pretiosa from 26 and 38 localities, the dependence of spotted on aquatic respectively, and concluded that at least two (Bos and Sites 2001) and population were represented, although isolation (Toline and Seitz 1999), there is likely no morphometrically, the two species are "almost gene flow existing between the Wasatch Front indistinguishable." and West Desert DPSs.

Green et al. (1997) determined that frogs from the Spotted frogs on Mitkof Island near Petersburg, vicinity of the type locality of R. pretiosa are Alaska, may exhibit a distinct phenotype of heavy conspecific with the species residing in south- dusky gray ventral coloring (MacDonald 2003). central and the Cascade Mountains of Oregon. Hence, they concluded that General description populations from southwestern British Columbia, A bumpy-skinned, medium-sized frog with western Washington, western and central relatively short hind legs, inconspicuous dorsal Oregon, and northeastern California are R. folds, and fully webbed toes. Individuals vary from pretiosa and that spotted frogs from the light to dark brown above with a scattering of remainder of the range are R. luteiventris. R. large black spots often with light centers. luteiventris was regarded as possibly comprising Underside is creamy or mottled gray, with a multiple weakly-differentiated species. covering of bright salmon or red on lower

abdomen and undersurfaces of hind legs in Further analyses of taxonomic relationships adults. Light stripe on the upper jaw, and the eyes among range-wide spotted frog populations were are upturned. Adults to 7.6 cm. This species is performed by Bos and Sites (2001); this study larger than the wood frog (R. sylvatica), has bright revealed four genetically distinct lineages. Two of salmon color over ventral surfaces, and lacks a these lineages are represented in : (1) the dark eye mask or light vertebral stripe. Deep Creek lineage (Deep Creek-Ibapah Distinguished from the introduced red-legged frog population in the West Desert distinct population (R. aurora) by its shorter legs (heel of hind leg segment or DPS), and (2) the Bonneville lineage when extended forward falling short of snout), (all other populations in Utah, including the with greater webbing, rougher skin, upturned Wasatch Front and the remainder of the West rather than out-turned eyes, shorter jaw stripe, Desert DPSs). The Wasatch Front DPS appears and lack of mottling on the groin (MacDonald to have originated from the West Desert 2003). populations in relatively recent evolutionary time, Length (cm) 7.6 during the recession of Lake Bonneville (Toline and Seitz 1999, Bos and Sites 2001). Therefore, Reproduction genetic differences between these populations Breeds in February at sea level in British have not yet been established. However, Columbia, mid-March at 1,395 m in Utah, May- separation of the West Desert and Wasatch Front June at 2,377 m in , mid-April through DPSs is supported by ecological and mid-May in Southeast Alaska; generally as early demographic distinctiveness due to geographic as winter thaw permits. In northeastern Oregon, eggs were not deposited on days when maximum Though movements of up to 6.5 km have been water temperature was below 9.4° C; at 18 sites, recorded, these frogs generally stay in wetlands duration of egg deposition ranged from 1 to 20 and along streams within 1 km of their breeding days (Bull and Shepherd 2003). Females may lay (Turner 1960, Hollenbeck 1974, Bull and egg masses in communal clusters. Eggs hatch in Hayes 2001, Pilliod et al. 2002). Frogs in isolated 3-21 days (12-21 days in northeastern Oregon; may not leave those sites (Bull and Hayes Bull and Shepherd 2003), depending on 2001). temperature. Metamorphosis occurs before fall or tadpoles may over-winter and metamorphose the Food following spring. Sexually mature in 2-6 years, Opportunistic. Eats a wide variety of terrestrial depending on location and elevation (matures and aquatic insects as well as different mollusks, later at high elevations). In Wyoming, individual , and . Larvae eat algae, females breed yearly at low elevations, every 2-3 organic debris, plant tissue, and minute aquatic years at high elevations (Nussbaum et al. 1983). organisms.

Ecology Phenology In the Toiyabe Range in , Reaser (2000) May move overland in spring and summer after captured 887 individuals over three years, with breeding. Inactive in winter in north. average mid-season density ranging from 2 to 24 frogs per 150 m of habitat. Columbia spotted Global habitat frogs may pass the winter in a torpid state Highly aquatic; rarely found far from permanent underwater or under soil or rocks, especially at quiet water; usually occurs at the grassy/sedge higher latitudes (James 1998). margins of streams, lakes, ponds, springs, and marshes (Hodge 1976, Licht 1986). May disperse Migration into , grassland, and brushland during wet May make short movements between breeding weather. Uses stream-side small mammal and non-breeding habitats. Males arrive in burrows as shelter (Blomquist and Tull 2002). breeding areas prior to females. Overwintering sites in the Great Basin include undercut stream banks and spring heads (K. In central , frogs moved up to 1030 m to Hatch, pers. comm., cited by Blomquist and Tull reach summer habitats; females (the more mobile 2002). Breeds usually in shallow water in ponds sex) moved less than 500 m on average from or other quiet waters. See Munger et al. (1998) for breeding or over-wintering sites to summer quantitative information on habitat in foraging areas (Pilliod et al. 2002). Pilliod et al. (in southwestern Idaho. Koch et al. 1997) reported that individual high mountain lake populations of R. luteiventris in State habitat Idaho are actually interdependent and part of a Closely associated with permanent water. Found larger contiguous metapopulation that includes all predominantly in outwash ponds and backwater the lakes in the basin. In Nevada, Reaser (1996 in lakes, beaver ponds, muskeg ponds, river Koch et al. 1997) determined that one individual channels, and streams (Waters 1992, MacDonald of R. luteiventris traveled over 5 km in a year. 2003).

In a three-year study of R. luteiventris movement Global range within the Owyhee Mountain subpopulation of the Extreme southeastern Alaska, southwestern Great Basin population in southwestern Idaho, Yukon (Slough 2002), northern British Columbia, Engle (2000) PIT-tagged over 1800 individuals and western Alberta south through Washington but documented only five (of 468 recaptures) over east of the Cascades, eastern Oregon, Idaho, and 1,000 m from their original capture point. All western Montana to Nevada (disjunct; Mary's, recaptures were along riparian corridors and the Reese, and Owyhee river systems), southwestern longest distance between capture points was Idaho (disjunct), Utah (disjunct; Wasatch 1,765 m. Although gender differences were Mountains and west desert), and western and observed, 88 % of all movement documented was north-central (disjunct) Wyoming (Stebbins 1985, less than 300 m from the original capture point. Green et al. 1996, 1997). Disjunct populations Engle (2001) found a two-year-old individual 6.5 occur on isolated mountains and in arid-land km downstream from its natal pond (a year after springs. Elevation range is from sea level to about being marked and released). 10,000 ft (Stebbins 1985). West Desert

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(Bonneville) population occurs mainly in two large Global short term trend spring complexes, with several additional Significant declines have occurred in some areas concentrations in smaller nearby springs; of Utah and Wyoming. Possibly has declined in extirpated from the northern portions of the range. Idaho, but numbers still apparently are high The Wasatch Front population occurs in isolated (Phillips 1990, Groves pers. comm. 1992). springs or riparian wetlands in Juab, Sanpete, Summit, Utah, Tooele, and Wasatch counties. Declines have been reported for disjunct Extirpated from the Salt Lake Valley and populations in the Wasatch Front in Utah, but the tributaries to the Jordan River and Great Salt recent trend is toward more secure populations, Lake (USFWS 2002). Currently, there are seven reduced threats, and improved habitat conditions localized populations that comprise the Wasatch (USFWS 2002). Front population or DPS. The largest known concentration is currently in the Heber Valley; the The West Desert (Bonneville) population has remaining six locations are Jordanelle/Francis, declined in range and abundance. Springville Hatchery, Holladay Springs, Mona Springs Complex/Burraston Ponds, Fairview, and Recent intensive surveys indicate severe declines Vernon (USFWS 2002). in the Great Basin populations. See Federal Register, 7 May 1993 (USFWS 1993) and 2 April State range 1998 (USFWS 1998). In the Toiyabe Range in Present in coastal of Southeast Alaska, central Nevada, demographic parameters although range limits are not precisely known exhibited significant spatial and temporal (Hodge 1976, MacDonald 2003). Present variation, some of which likely was due to distribution confined to coastal transboundary extreme variations in annual weather patterns river corridors of continental mainland in (Reaser 2000). Recent reports suggest Alberta Southeast, such as Salmon, Taku, Stikine and populations may be in decline (James 1998). Unuk rivers and the Agassiz Peninsula (MacDonald 2003). Have been documented on Global long term trend Mitkof, Sergief and Vank Islands within the Relatively stable in most of the range, but adjacent Alexander Archipelago (Waters 1992, populations in the arid southern portion of the Lindell and Grossman 1998). Mitkof Island range have declined. population (in and near the city of Wrangell) possibly introduced. Carstensen et al. (2003) State trend reported frogs at one location along the Juneau Population status in Alaska is unknown. road system, suspected this was also an introduction. Also reported but not confirmed in Global protection Haines area (MacDonald 2003). Somewhat protected in several federal and state parks and refuges, though management usually Global abundance ignores this species. Total adult population size is unknown but likely exceeds 100,000. Numerous in many areas in A considerable portion of the range of the West Canada and the Rocky Mountains. Desert population is under management of the Bureau of Land Management. Conservation West Desert population: over 6,000 spotted frog activities implemented for the least chub egg masses were observed in 1993 (D.Ross, (Iotichthys phlegethontis) should also benefit the Herpetologist, Utah Division of Wildlife). West Desert population.

State abundance Wasatch Front population occurs mainly on Current population size unknown. Population private land, with some federal ownership along studies conducted at isolated ponds in the Taku, Jordanelle Dam and the Provo River. Habitat Stikine and Unuk River corridors estimated local acquisition and protection are in progress for the populations ranged from 7 to 594 Columbia Jordanelle/Francis, Heber Valley, spotted frogs per pond and densities ranging from Mona/Burraston, and Fairview populations. 0.07 to 1.49 spotted frogs per m2 of pond habitat Current ventures are focused on acquiring habitat (Lindell and Grossman 1998). easements along approximately 9.7 kilometers (6 miles) above Jordanelle Dam, including occupied and suitable spotted frog habitats. Easements are

Columbia spotted frog 3 currently being pursued with 7 Fairview reproduction. Additional threats include continued landowners to protect approximately 162 hectares development along the Wasatch Front, water (400 acres) of occupied spotted frog habitat and diversions for irrigation, cattle grazing, timber migration corridors from potential water and harvesting, and construction of roads and trails. residential development. The remaining 15 Introduced bullfrogs and fishes may have an percent of the Provo River corridor in the Heber adverse impact on this population, but the current Valley is projected to be purchased and protected degree of threat is unknown. Mosquito control by 2004. In the Mona/Burraston population, fee- agents also pose a potential threat. However, title purchase or conservation easements are USFWS (2002) concluded the following: The currently being negotiated for 7.9 hectares (19.5 overall level of threats to the long-term acres) which would allow for protection of all persistence of the Wasatch Front population has spring and potential spotted frog habitat on this decreased in recent years, particularly since site. 1998. Although most of the human activities that contributed to these threats still occur to some Great Basin population occurs primarily on lands extent throughout the Wasatch Front, there is no managed by the U.S. Forest Service and Bureau longer the same level of impacts on the frog that of Land Management. resulted from past wide-spread and loss of populations. Much of the occupied State protection habitat is under state or federal ownership, and Most areas of occurrence in Alaska are federally ongoing management of these lands emphasizes owned, administered by the USDA Forest the long-term persistence of R. luteiventris. Service. These include portions of the Tongass Threats still exist in localized areas, but National Forest and Misty Fiords Monument mechanisms are in place through federal, state, (which are managed as no or low development and local conservation and land-use plans to areas) (Lindell and Grossman 1998). In Alaska, identify and correct the problems, and protect are managed by Alaska Department spotted frog populations. To date, these actions of Fish and Game under statute 16.05.030, in have been successful at reducing threats to which amphibians are legally included in the extant populations, largely by acquiring important definition of “fish.” This statute makes it illegal for habitats and implementing management actions anyone to “hold, transport or release” any native that improve habitat conditions. Success is amphibians without a valid permit. evidenced by the stable to improving status of the spotted frog throughout the Wasatch Front in the Global threats most recent time period evaluated. Populations in western Utah are limited by scarce habitat (springs) and are potentially threatened by Great Basin population has been adversely habitat degradation from cattle grazing and affected by habitat degradation resulting from agricultural activities. Oil and gas exploration is an mining, livestock grazing, road construction, increasing threat. Water development could lower agriculture, and direct by bullfrogs and water tables and adversely impact spring habitats. non-native fishes. In central Nevada, introduction Introduced bullfrogs and fishes may have an of exotic trout and cattle are likely the most adverse impact, but the current degree of threat is important anthropogenic factors limiting the unknown. Mosquito control agents pose a distribution and persistence of R. luteiventris potential threat. A recent conservation agreement (Reaser 2000). among the state of Utah and other agencies has significantly reduced the level of threat to the This species is not likely to be at risk from present West Desert population (Federal Register, 2 April acidification inputs in the Rocky Mountains (Corn 1998). and Vertucci 1992). Global climate changes are possibly a factor (Hayes and Jennings 1986). UV- Wasatch Front population is facing serious threats B radiation to frog embryos does not currently from habitat loss and modification, especially seem to be contributing to population declines water development associated with the Central (Blaustein et al. 1999). Utah Project; current and imminent threats include the Provo River Restoration Project and State threats Wasatch County Efficiency Project; wetlands Human-caused direct impacts are low due to created as mitigation for the Central Utah Project protection throughout a large part of its range in have contributed only minimally to spotted frog the Tongass National Forest and Misty Fiords

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National Monument. Filling or draining of wetland ecological redundancy in ecological function and habitat and alterations to ground or surface water genetic and demographic stochasticity. There are flow from development are potential hazards. several habitats already identified which may Chronic or acute water contamination from provide suitable reintroduction sites. Future existing and proposed mining activities on the conservation should include reestablishment of Canadian reaches of the Taku and Stikine Rivers Columbia spotted frog populations, and may be of concern. Existing wetlands in the Taku associated research and land management and Stikine River corridors may be significantly necessary to maintain new populations in: (a) reduced as postglacial rebound continues (Lindell Areas where populations previously occurred if and Grossman 1998). suitable habitat remains and (b) other suitable habitat within the natural range of the species. (3) State research needs Some Wasatch Front spotted frog populations are Identify threats or limiting factors. Establish notably small in size and vulnerable to risks of monitoring programs to track population trends. detrimental genetic processes (inbreeding, loss of genetic diversity) and demographic uncertainty. Global inventory needs Springville Hatchery/T-Bone Bottom population is Range-wide population inventories are needed to particularly vulnerable based on its current size determine abundance and distribution. and decreasing trend. Actions should be taken to augment or, through some other process, State inventory needs increase the size of this population. Furthermore, Additional inventory to precisely determine the the current trend should be evaluated to species' range is needed; possible locations of determine if specific land or water use activities occurrence include the vicinity of Haines, are exacerbating the decrease. If specific threats Skagway, Whiting and Taiya Rivers. Population are identified, priority should be placed on estimates for each area of occurrence are reducing these threats such that the population needed; establish programs to monitor population would remain secure into the future. trends. Periodic population estimate surveys should continue where baseline estimates have State conservation and management needs been established (Lindell and Grossman 1998). Prevent habitat loss in known breeding areas. This species is disappearing from many areas of Global conservation and management needs its range, but still considered common in British See Mullen (1999) for a brief description of a draft Columbia. The current population trend should be conservation strategy and agreement for spotted evaluated, and threats to populations identified. If frogs in northeastern Nevada. specific threats are identified, priority should be placed on reducing these threats such that the For the Wasatch Front population, USFWS (2002) population will remain secure into the future. concluded that the focus of conservation efforts can reasonably shift to acquisition of additional occupied and unoccupied suitable habitats and LITERATURE CITED range expansion efforts, including: (1) Land Blaustein, Andrew R.; Hays, John B.; Hoffman, protection mechanisms, such as conservation Peter D.; Chivers, Douglas P.; Kiesecker, easements and fee-title acquisitions which Joseph M.; Leonard, William P.; Marco, generally provide the most long-term benefits for Adolfo; Olson, Deanna H.; Reaser, Jamie sensitive species. Voluntary conservation actions K.; Anthony, Robert G. DNA repair and on parcels of private land may provide site- resistance to UV-B radiation in western specific benefits to the frog. Future conservation spotted frogs. Ecological Applications, 9(3): should continue to focus on land acquisition and 1100-1105. easements that include buffer zones sufficient to minimize direct and indirect impacts from land use Blomquist, S.M., and J.C. Tull. 2002. Rana as well as protection and maintenance of luteiventris: burrow use. Herpetological dispersal or migration corridors. Furthermore, Review 33:131. steps should be taken to protect water sources (i.e. Juab Valley) where potential threats are Bos, D.H. and J.W. Sites. 2001. Phylogeography identified. (2) Although there is no specific and conservation genetics of the Columbia number of populations necessary to prevent spotted frog (Rana luteiventris; Amphibia, extinction, reintroduced populations provide Ranidae). Molecular Ecol 10: 1499-1513.

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are bullfrogs (Rana catesbeiana) Bull, E.L., and M.P. Hayes. 2001. Post-breeding responsible? J. Herpetol. 20(4):490-509. season movements of Columbia spotted frogs (Rana luteiventris) in northeastern Oregon. Hodge, R.P. 1976. Amphibians & reptiles in Western North American Naturalist 61:119- Alaska, the Yukon, & Northwest Territories. 123. Alaska Northwest Publishing Company, Anchorage, AK. 89 pp. Bull, E.L., and J.F. Shepherd. 2003. Water temperature at oviposition sites of Rana Hollenbeck, R.R. 1974. Growth rates and luteiventris in northeastern Oregon. Western movements within a population of Rana North American Naturalist 63:108-113. pretiosa pretiosa Baird and Girard in south central Montana. M.A. thesis, Montana State Carstensen, R., M. Willson and R. Armstrong. University, Bozeman. 2003. Habitat use of amphibians in northern southeast Alaska. Unpublished report to Hovingh, P. 1993. Aquatic habitats, life history Alaska Department of Fish and Game. observations and zoogeographic Juneau, AK: Discovery Southeast. considerations of the spotted frog (Rana pretiosa) in Tule Valley, Utah. Great Basin Corn, P. S., and F. A. Vertucci. 1992. Descriptive Naturalist. 53:168-179. risk assessment of the effects of acidic deposition on Rocky Mountain amphibians. J. James, J.D. 1998. Status of the Columbia spotted Herpetol. 26:361-369. frog (Rana luteiventris) in Alberta. Wildlife Status Report No. 17. Edmonton, AB: Alberta Engle, J.C. 2000. Columbia spotted frog Great Environmental Protection, Fisheries & Wildlife Basin population (Owyhee Mountains Management Division, and Alberta subpopulation) long-term monitoring plan. Conservation Association. Year 200 Results. (draft). Boise, ID. Koch, E.D., G. Williams, C.R. Peterson, and P.S. Engle, J.C. 2001. Population biology and natural Corn. 1997. A summary of the conference on history of Columbia spotted frogs (Rana declining and sensitive amphibians in the luteiventris) in the Owyhee Uplands of Rocky Mountains and Pacific Northwest. southwest Idaho: implications for monitoring November 7-8, 1996, Boise Idaho. Meeting and management. M.Sc. Boise State sponsored by U.S. Fish and Wildlife Service, University, Boise, ID. U.S. Forest Service, U.S. Bureau of Land Management, Idaho Herpetological Society, Green, D.M., T.F. Sharbel, J. Kearsley, and H. Declining Population Task Force Kaiser. 1996. Postglacial range fluctuation, Rocky Mountain Working Group, U.S. genetic subdivision and speciation in the Geological Survey - Biological Resources western North American spotted frog Division, Idaho Museum of Natural History, complex, Rana pretiosa. Evolution 50:374- Idaho State University. 390. Licht, L.E. 1986. Food and feeding behavior of Green, D.M., H. Kaiser, T.F. Sharbel, J. Kearsley, sympatric red-legged frogs, Rana aurora, and and K.R. McAllister. 1997. Cryptic species of spotted frogs, Rana pretiosa, in southwestern spotted frogs, Rana pretiosa complex, in British Columbia. Canad. Field-Naturalist western North America. Copeia 1997:1-8. 100:22-31.

Groves, Craig. 1992. Phone call regarding the Lindell, J.R. and E.M. Grossman. 1998. Columbia status of the spotted frog. Idaho Natural spotted frog (Rana luteiventris) distribution Heritage Program, Dept. of Fish and Game, and local abundance in southeast Alaska: 600 S. Walnut St., Box 25, Boise, ID 83707. Juneau, Alaska: U.S. Fish and Wildlife (208) 334-3402. Service, Southeast Alaska Ecological Services. Hayes, M.P. and M.R. Jennings. 1986. Decline of Ranid frog species in western North America: MacDonald, S.O. 2003.The amphibians and reptiles of Alaska. A Field Handbook.

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Unpublished report to U.S. Fish and Wildlife petition to list the spotted frog. Fed. Register Service, Juneau, AK. 58:27260-27263.

Mullen, C. 1999. Community-based stewardship U.S. Fish and Wildlife Service. 1998. Endangered in Nevada. Endangered Species Bulletin 24:9. and threatened wildlife and plants: new 12- month finding for a petition to list the Utah Munger, J.C., M. Gerber, K. Madrid, M.-A. Carroll, Wasatch Front and west desert populations of W. Petersen, and L. Heberger. 1998. U.S. spotted frog. 30 April 2, 1998. Federal National Wetland Inventory classifications as Register 63:16218-16220. predictors of the occurrence of Columbia spotted frogs (Rana luteiventris) and Pacific U.S. Fish and Wildlife Service. 2002. 12-month tree frogs (Hyla regilla). Conservation Biology finding for a petition To list the Wasatch Front 12:320-330. Columbia spotted frog as threatened throughout its range. 30 August 2002 Federal Nussbaum, R.A., E.D. Brodie, Jr., and R.M. Register 67(169):55758-55767. Storm. 1983. Amphibians and reptiles of the Pacific Northwest. Univ. Press of Idaho. 332 Waters, D.L. 1992. Habitat associations, pp. phenology, and biogeography of amphibians in the Stikine River basin and southeast Phillips, K. 1990. Where have all the frogs and Alaska. Unpubl. rep. of the 1991 pilot project. toads gone? BioScience 40:422-424. U.S. Dept. Interior, Fish and Wildlife Service, California Cooperative Fishery Research Unit, Pilliod, D.S., C.R. Peterson, and P.I. Ritson. 2002. Humboldt State University, Arcata, CA. 61 pp. Seasonal migration of Columbia spotted frogs (Rana luteiventris) among complementary resources in a high mountain basin. Canadian Acknowledgements Journal of Zoology 80:1849-1862. State Conservation Status, Reaser, J.K. 2000. Demographic analysis of the Element Ecology & Life History Columbia spotted frog (Rana luteiventris): Author(s): Gotthardt, T.A. case study in spatiotemporal variation. State Conservation Status, Element Ecology & Canadian Journal of Zoology 78:1158-1167. Life History Edition Date: 22Mar2005

Slough, B.G. 2002. Geographic distribution: Rana Reviewer(s): Stephen MacDonald, University of luteiventris. Herpetological Review 33:146. New Mexico; Blain Anderson, National Park Service. Stebbins, R.C. 1985. A Field Guide to Western Reptiles and Amphibians. Second Edition. Life history and Global level information were Houghton Mifflin Company, Boston, obtained from the on-line database, NatureServe Massachusetts. xiv + 336 pp. Explorer (www.natureserve.org/explorer). In many cases, life history and Global information were Toline, C.A. and A.M. Seitz. 1999. Mitochondrial updated for this species account by Alaska and nuclear DNA variation within and among Natural Heritage Program zoologist, Tracey populations of Columbia spotted frog (Rana Gotthardt. All Global level modifications will be luteiventris) in Utah. Unpublished report sent to NatureServe to update the on-line version. prepared for the Utah Division of Wildlife NatureServe Conservation Status Factors Resources. Salt Lake City, UT. Edition Date: 27Jan2003 NatureServe Conservation Status Factors Turner, F.B. 1960. Population structure and Author: Hammerson, G. dynamics of the western spotted frog, Rana p. Global Element Ecology & Life History Edition pretiosa Aaird and Girard, in Yellowstone Date: 05Jan2004 Park, Wyoming. Ecol. Monogr. 30:251-178. Global Element Ecology & Life History Author(s): Hammerson, G. U.S. Fish and Wildlife Service. 1993. Endangered and threatened wildlife and plants: finding on

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