Overview

Comprehensive Description

Description

This frog is a member of the mountain yellow-legged frog complex which is comprised of two species: Rana muscosa and Rana sierrae. Both species are highly aquatic and are always found within a meter or two from the edge of water. Rana sierrae is yellowish or reddish brown from above, with black or brown spots or lichen-like markings. Toe tips are usually dusky. Underside of hind legs and sometimes entire belly is yellow or slightly orange, usually more opaque than in foothill yellow-legged frog, Rana boylii. Yellow often extends forward to level of forelimbs. Dorsolateral folds present but frequently indistinct. The tadpoles are black or dark brown and are large (total length often exceeds 10 cm) and metamorphose in 1-4 years depending on the elevation. Rana sierrae differs from Rana muscosa in having relatively shorter legs. When a leg is folded against the body the tibio-tarsal joint typically falls short of the external nares. The mating call of R. sierrae is significantly different from that of R. muscosa in having transitions between pulsed and noted sounds. Both species call underwater. Males can be heard above water but only from a short distance away (<2 meters). The two species also differ in mitochondrial DNA. The mitochondrial DNA, male advertisement calls, and morphology datasets are geographically concordant (Vredenburg et al. 2007).

  • Wright, A. H. and Wright, A. A. (1949). Handbook of Frogs and Toads of the United States and Canada. Comstock Publishing Company, Inc., Ithaca, New York.
  • Stebbins, R.C. (1951). Amphibians of Western North America. University of California Press, Berkeley.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Berger, L., Speare, R., Daszak, P., Green, D. E., Cunningham, A. A., Goggin, C. L., Slocombe, R., Ragan, M. A., Hyatt, A. D., McDonald, K. R., Hines, H. B., Lips, K. R., Marantelli, G., and Parkes, H. (1998). "Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America." Proceedings of the National Academy of Sciences of the United States of America, 95(15), 9031-9036.
  • Storer, T. I. (1925). "A synopsis of the amphibia of California." University of California Publications in Zoology, 27, 1-342.
  • Jennings, M. R., and Hayes, M. P. (1994). ''Amphibian and reptile species of special concern in California.'' Final Report #8023 Submitted to the California Department of Fish and Game. California Department of Fish and Game, Sacramento, California..
  • Drost, C. A., and Fellers, G. M. (1996). "Collapse of a regional frog fauna in the Yosemite area of the California Sierra Nevada, USA." Conservation Biology, 10(2), 414-425.
  • Bradford, D. F. (1989). "Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California: implication of the negative effect of fish introductions." Copeia, 1989, 775-778.
  • Bradford, D. F. (1989). ''Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California USA: Implication of the negative effect of fish introductions.'' Copeia, 1989(3), 775-778.
  • Bradford, D. F., Tabatabai, F., and Graber, D. M. (1993). ''Isolation of remaining populations of the native frog, Rana muscosa, by introduced fishes in Sequoia and Kings Canyon National Parks, California.'' Conservation Biology, 7, 882-888.
  • Briggs, C. J., Knapp, R. A., and Vredenburg, V. T. (2010). ''Enzootic and epizootic dynamics of the chytrid fungal pathogen of amphibians.'' Proceedings of the National Academy of Sciences, 107(21), 9695-9700 .
  • Davidson, C. (2004). ''Declining downwind: Amphibian population declines in California and historical pesticide use.'' Ecological Applications, 14, 1892-1902.
  • Davidson, C., Shaffer, H. B., and Jennings, M. R. (2002). ''Spatial tests of the pesticide drift, habitat destruction, UV-B, and climate-change hypotheses for California amphibian declines.'' Conservation Biology, 16, 1588-1601.
  • Finlay, J. and Vredenburg, V. T. (2007). ''Introduced trout sever trophic connections between lakes and watersheds: consequences for a declining montane frog.'' Ecology, 88(9), 2187-2198.
  • Grinnell, J., and Storer, T. I. (1924). Animal Life in the Yosemite. University of California Press, Berkeley, California.
  • Knapp, R. A. and Matthews, F. (2000). ''Non-native fish introductions and the decline of the Mountain Yellow-legged Frog from within protected areas.'' Conservation Biology, 14(2), 428-439.
  • Knapp, R. A., Boiano, D. M., Vredenburg, V. T. (2007). ''Recovery of a declining amphibian (Mountain Yellow-legged Frog, Rana muscosa) following removal of non-native fish.'' Biological Conservation, 135, 11-20.
  • Knapp, R.A. (1996). ''Non-native trout in the natural lakes of the Sierra Nevada: an analysis of their distribution and impacts on native aquatic biota.'' Sierra Nevada Ecosystem Project, Final Report to Congress, Center for Water and Wildland Resources, University of California (Davis), Davis, California, 363-390.
  • Livezey, R. L., and Wright, A. H. (1945). ''Descriptions of four salientian eggs.'' American Midland Naturalist, 34, 701-706.
  • Rachowicz, L. J., Knapp, R. A., Morgan, J. A. T., Stice, M. J., Vredenburg, V. T., Parker, J. M., and Briggs, C. J. (2006). ''Emerging infectious disease as a proximate cause of amphibian mass mortality.'' Ecology, 87, 1671-1683.
  • Vredenburg, V. T. (2004). ''Reversing introduced species effects: Experimental removal of introduced fish leads to rapid recovery of a declining frog.'' Proceedings of the National Academy of Sciences of the United States of America, 101, 7646-7650.
  • Vredenburg, V. T., (2007). ''Concordant molecular and phenotypic data delineate new taxonomy and conservation priorities for the endangered mountain yellow-legged frog (Ranidae: Rana muscosa).'' Journal of Zoology, 271, 361-374.
  • Vredenburg, V. T., Fellers, G., and Davidson, C. (2005). ''The mountain yellow-legged frog Rana muscosa (Camp 1917).'' Status and conservation of U.S. Amphibians. M. Lannoo, eds., University of California Press, Berkeley, 563-566.
  • Vredenburg, V. T., Knapp, R. A., Tunstall, T. S., and Briggs, C. J. (2010). ''Dynamics of an emerging disease drive large-scale amphibian population extinctions.'' Proceedings of the National Academy of Sciences, 107(21), 9689-9694.
  • Zweifel, R. G. (1955). ''Ecology, distribution, and systematics of frogs of the Rana boylei group.'' University of California Publications in Zoology, 54, 207-292.
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Distribution

Global Range: (5000-20,000 square km (about 2000-8000 square miles)) Historical range extended from the Diamond Mountains northeast of the Sierra Nevada in Plumas County, and from just north of the Feather River in the extreme northwest region of the Sierra Nevada, California, south through the Sierra Nevada to Inyo County, California, and east to Mt. Rose, northeast of Lake Tahoe, in Washoe County, Nevada (Vredenburg et al. 2007). West of the Sierra Nevada crest, the southern part of the range is bordered by ridges that divide the Middle and South Fork of the Kings River, ranging from Mather Pass to the Monarch Divide; east of the Sierra Nevada crest, R. sierrae occurs in the Glass Mountains just south of Mono Lake (Mono County) and along the east slope of the Sierra Nevada south to the type locality at Matlock Lake (Inyo County) (Vredenburg et al. 2007). Rana sierrae is now extirpated from Nevada and from large portions of the historical range in the Sierra Nevada of California. Elevational range is 1,370-3,690 meters (Fellers et al. 2013).

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endemic to a single nation

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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Range Description

Historical range extended from the Diamond Mountains northeast of the Sierra Nevada in Plumas County, and from just north of the Feather River in the extreme northwest region of the Sierra Nevada, California, south through the Sierra Nevada to Inyo County, California, and east to Mt. Rose, northeast of Lake Tahoe, in Washoe County, Nevada (Vredenburg et al. 2007). West of the Sierra Nevada crest, the southern part of the range is bordered by ridges that divide the Middle and South Fork of the Kings River, ranging from Mather Pass to the Monarch Divide; east of the Sierra Nevada crest, R. sierrae occurs in the Glass Mountains just south of Mono Lake (Mono County) and along the east slope of the Sierra Nevada south to the type locality at Matlock Lake (Inyo County) (Vredenburg et al. 2007). Rana sierrae is now extirpated from Nevada and from large portions of the historical range in the Sierra Nevada of California.
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Distribution and Habitat

This montane species once occurred in California and Nevada, USA but is now extinct in the state of Nevada. Rana sierrae ranges from the Diamond Mountains northeast of the Sierra Nevada in Plumas County, California, south through the Sierra Nevada to the type locality, the southern-most locality at Matlock Lake just east of Kearsarge Pass (Inyo County, California). In the extreme northwest region of the Sierra Nevada, several populations occur just north of the Feather River, and to the east, there was a population on Mt. Rose, northeast of Lake Tahoe in Washoe County, Nevada, but, as mentioned above, it is now extinct. West of the Sierra Nevada crest, the southern part of the R. sierrae range is bordered by ridges that divide the Middle and South Fork of the Kings River, ranging from Mather Pass on the John Muir Trail east to the Monarch Divide. East of the Sierra Nevada crest, R. sierrae occurs in the Glass Mountains just south of Mono Lake (Mono County, CA) and along the east slope of the Sierra Nevada south to the type locality at Matlock Lake (Inyo County, CA).

  • Wright, A. H. and Wright, A. A. (1949). Handbook of Frogs and Toads of the United States and Canada. Comstock Publishing Company, Inc., Ithaca, New York.
  • Stebbins, R.C. (1951). Amphibians of Western North America. University of California Press, Berkeley.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Berger, L., Speare, R., Daszak, P., Green, D. E., Cunningham, A. A., Goggin, C. L., Slocombe, R., Ragan, M. A., Hyatt, A. D., McDonald, K. R., Hines, H. B., Lips, K. R., Marantelli, G., and Parkes, H. (1998). "Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America." Proceedings of the National Academy of Sciences of the United States of America, 95(15), 9031-9036.
  • Storer, T. I. (1925). "A synopsis of the amphibia of California." University of California Publications in Zoology, 27, 1-342.
  • Jennings, M. R., and Hayes, M. P. (1994). ''Amphibian and reptile species of special concern in California.'' Final Report #8023 Submitted to the California Department of Fish and Game. California Department of Fish and Game, Sacramento, California..
  • Drost, C. A., and Fellers, G. M. (1996). "Collapse of a regional frog fauna in the Yosemite area of the California Sierra Nevada, USA." Conservation Biology, 10(2), 414-425.
  • Bradford, D. F. (1989). "Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California: implication of the negative effect of fish introductions." Copeia, 1989, 775-778.
  • Bradford, D. F. (1989). ''Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California USA: Implication of the negative effect of fish introductions.'' Copeia, 1989(3), 775-778.
  • Bradford, D. F., Tabatabai, F., and Graber, D. M. (1993). ''Isolation of remaining populations of the native frog, Rana muscosa, by introduced fishes in Sequoia and Kings Canyon National Parks, California.'' Conservation Biology, 7, 882-888.
  • Briggs, C. J., Knapp, R. A., and Vredenburg, V. T. (2010). ''Enzootic and epizootic dynamics of the chytrid fungal pathogen of amphibians.'' Proceedings of the National Academy of Sciences, 107(21), 9695-9700 .
  • Davidson, C. (2004). ''Declining downwind: Amphibian population declines in California and historical pesticide use.'' Ecological Applications, 14, 1892-1902.
  • Davidson, C., Shaffer, H. B., and Jennings, M. R. (2002). ''Spatial tests of the pesticide drift, habitat destruction, UV-B, and climate-change hypotheses for California amphibian declines.'' Conservation Biology, 16, 1588-1601.
  • Finlay, J. and Vredenburg, V. T. (2007). ''Introduced trout sever trophic connections between lakes and watersheds: consequences for a declining montane frog.'' Ecology, 88(9), 2187-2198.
  • Grinnell, J., and Storer, T. I. (1924). Animal Life in the Yosemite. University of California Press, Berkeley, California.
  • Knapp, R. A. and Matthews, F. (2000). ''Non-native fish introductions and the decline of the Mountain Yellow-legged Frog from within protected areas.'' Conservation Biology, 14(2), 428-439.
  • Knapp, R. A., Boiano, D. M., Vredenburg, V. T. (2007). ''Recovery of a declining amphibian (Mountain Yellow-legged Frog, Rana muscosa) following removal of non-native fish.'' Biological Conservation, 135, 11-20.
  • Knapp, R.A. (1996). ''Non-native trout in the natural lakes of the Sierra Nevada: an analysis of their distribution and impacts on native aquatic biota.'' Sierra Nevada Ecosystem Project, Final Report to Congress, Center for Water and Wildland Resources, University of California (Davis), Davis, California, 363-390.
  • Livezey, R. L., and Wright, A. H. (1945). ''Descriptions of four salientian eggs.'' American Midland Naturalist, 34, 701-706.
  • Rachowicz, L. J., Knapp, R. A., Morgan, J. A. T., Stice, M. J., Vredenburg, V. T., Parker, J. M., and Briggs, C. J. (2006). ''Emerging infectious disease as a proximate cause of amphibian mass mortality.'' Ecology, 87, 1671-1683.
  • Vredenburg, V. T. (2004). ''Reversing introduced species effects: Experimental removal of introduced fish leads to rapid recovery of a declining frog.'' Proceedings of the National Academy of Sciences of the United States of America, 101, 7646-7650.
  • Vredenburg, V. T., (2007). ''Concordant molecular and phenotypic data delineate new taxonomy and conservation priorities for the endangered mountain yellow-legged frog (Ranidae: Rana muscosa).'' Journal of Zoology, 271, 361-374.
  • Vredenburg, V. T., Fellers, G., and Davidson, C. (2005). ''The mountain yellow-legged frog Rana muscosa (Camp 1917).'' Status and conservation of U.S. Amphibians. M. Lannoo, eds., University of California Press, Berkeley, 563-566.
  • Vredenburg, V. T., Knapp, R. A., Tunstall, T. S., and Briggs, C. J. (2010). ''Dynamics of an emerging disease drive large-scale amphibian population extinctions.'' Proceedings of the National Academy of Sciences, 107(21), 9689-9694.
  • Zweifel, R. G. (1955). ''Ecology, distribution, and systematics of frogs of the Rana boylei group.'' University of California Publications in Zoology, 54, 207-292.
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Ecology

Habitat

Comments: The habitat of frogs of the Rana muscosa/Rana sierrae complex includes sunny river margins, meadow streams, isolated pools, and lake borders in the Sierra Nevada. Sierran frogs are most abundant in high elevation lakes and slow-moving portions of streams. They seldom are found away from water but may cross upland areas in moving between summer and winter habitats (Matthews and Pope 1999). Breeding success depands on perennial bodies of water because larave require mutliple years of development before metamorphosis. Wintering sites include areas near shore under ledges and in deep underwater crevices (Matthews and Pope 1999).

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Habitat and Ecology

Habitat and Ecology
The habitat of frogs of the Rana muscosa/Rana sierrae complex includes sunny river margins, meadow streams, isolated pools, and lake borders in the Sierra Nevada. Sierran frogs are most abundant in high elevation lakes and slow-moving portions of streams. They seldom are found away from water but may cross upland areas in moving between summer and winter habitats (Matthews and Pope 1999). Wintering sites include areas near shore under ledges and in deep underwater crevices (Matthews and Pope 1999).

Systems
  • Terrestrial
  • Freshwater
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Migration

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

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Trophic Strategy

Comments: Adults of the Rana muscosa/Rana sierrae complex eat aquatic and terrestrial invertebrates and anuran larvae; availability of larval anuran prey may be an important factor in distribution, body condition, and survival of adults (Pope and Matthews 2002). Larvae eat algae, organic debris, plant tissue, and minute organisms in water.

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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 - 300

Comments: Extensive surveys between 1995 and 2005 yielded only 11 occupied sites (Vredenburg et al. 2007).

California Department of Fish and Game (2011) determined that Rana sierrae currently occupies 1,199 sites across 94 USGS HU12 watersheds (a site was defined as a discrete pond, lake, reservoir, meadow, marsh, spring, or stream).

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Global Abundance

1000 - 10,000 individuals

Comments: Total adult population size is unknown but may not exceed 10,000. Most populations are very small (fewer than 100 post-metamorphic frogs (not all of these are adults), but ten HU12 watershed units include 501-2,500 post-metamorphic individuals and two watershed untis include more than 2,500 (California Department of Fish and Game 2011).

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Life History and Behavior

Cyclicity

Comments: Inactive in cold temperatures. Primarily diurnal. Inactive 7-9 months each year at high elevations (Pope and Matthews 2002).

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Reproduction

At high elevations larvae overwinter up to 3-4 times before metamorphosis.

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Conservation

Conservation Status

National NatureServe Conservation Status

United States

Rounded National Status Rank: N2 - Imperiled

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NatureServe Conservation Status

Rounded Global Status Rank: G2 - Imperiled

Reasons: Occurs in the Sierra Nevada of California and (formerly) extreme western Nevada; numerous population declines and local extirpations have occurred and are ongoing; introduced trouts and chytrid fungal disease are major factors in the decline; vulnerable to further declines from the effects of climate change.

Intrinsic Vulnerability: Moderately vulnerable

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IUCN Red List Assessment


Red List Category
EN
Endangered

Red List Criteria
B2ab(ii,iv,v)

Version
3.1

Year Assessed
2008

Assessor/s
Geoffrey Hammerson

Reviewer/s
Global Amphibian Assessment Coordinating Team (Simon Stuart, Janice Chanson and Neil Cox)

Contributor/s

Justification
Listed as Endangered in view of severe recent declines that likely are continuing (rate of decline over past 10 years unknown) and that has left only a small number of extant populations occupying a small fragmented area.
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Global Short Term Trend: Decline of 30-50%

Comments: Area of occupancy, number of subpopulations and locations, and population size appear to have undergone a substantial decline over the past three generations (around three decades, since the early 1980s) (California Department of Fish and Game 2011).

Most populations appear to have declined, but a population in a low-elevation site in Yosemite National Park was relatively stable (45-115 adults) during the period 2003-2011 (Fellers et al. 2013).

Global Long Term Trend: Decline of 30-70%

Comments: A precipitous decline in Rana muscosa/Rana sierrae appears to have occurred over the past 3-4 decades (Bradford 1991, USFWS 1999). For the Rana muscosa/Rana sierrae complex as a whole, Jennings and Hayes (1994) mapped many more extirpated populations than extant populations. Rana sierrae has declined greatly in the Yosemite area of the Sierra Nevada, California (Drost and Fellers 1996). In the Sierra Nevada, recent surveys indicate that Rana muscosa/Rana sierrae has been reduced to a small number of widely scattered, mostly very small populations (fewer than 20 adults) (Knapp and Matthews 2000). Surveys in the 1990s indicated that the rangewide decline in distribution may be as much as 70-90 percent (USFWS 2000).

Of the 146 historical R. sierrae sites studied by Vredenburg et al. (2007), only 11 sites contained frogs when revisited between 1995 and 2005 (92 percent extirpation rate).

California Department of Fish and Game (2011) determined that Rana sierrae has been extirpated from 220 (69 percent) of 318 historical localities analyzed (i.e., with sufficient data) and from 55 (44 percent) of 124 historically occupied HU12 watersheds.

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Population

Population
Extensive surveys between 1995 and 2005 yielded only 11 occupied sites (Vredenburg et al. 2007). Total adult population size is unknown but may not exceed a couple thousand (generously assuming 20 sites each with 100 adults).

A precipitous decline in Rana muscosa/Rana sierrae appears to have occurred over the past 3-4 decades (Bradford 1991, USFWS 1999). For the Rana muscosa/Rana sierrae complex as a whole, Jennings and Hayes (1994) mapped many more extirpated populations than extant populations. Rana sierrae has declined greatly in the Yosemite area of the Sierra Nevada, California (Drost and Fellers 1996). In the Sierra Nevada, recent surveys indicate that Rana muscosa/Rana sierrae has been reduced to a small number of widely scattered, mostly very small populations (fewer than 20 adults) (Knapp and Matthews 2000). Surveys in the 1990s indicated that the rangewide decline in distribution may be as much as 70-90 percent (USFWS 2000).

Of the 146 historical R. sierrae sites studied by Vredenburg et al. (2007), only 11 sites contained frogs when revisited between 1995 and 2005 (92 percent extirpation rate).

The current trend (past 10 years) is unknown, but probably the decline is ongoing.

Population Trend
Decreasing
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Life History, Abundance, Activity, and Special Behaviors

Similar to R. muscosa, breeding begins soon after ice-melt or early in spring and can range from April at lower elevations to June and July in higher elevations (Wright and Wright 1949; Stebbins 1951; Zweifel 1955). Eggs are deposited underwater in clusters attached to rocks, gravel, and under banks, or to vegetation in streams or lakes (Wright and Wright 1949; Stebbins 1951; Zweifel 1955). Livezey and Wright (1945) report an average of 233 eggs per mass(n=6, range 100-350). Eggs contain a vitelline capsule, and three gelatinous envelopes, all clear and transparent (see illustrations in: Stebbins 2003). In laboratory breeding experiments egg hatching times ranged from 18-21+ days at temperatures ranging from 5-13.5 °C (Zweifel 1955). The length of the larval stage depends upon the elevation. At lower elevations where the summers are longer, tadpoles are able to grow to metamorphosis in a single season (Storer 1925). At higher elevations where the growing season can be as short as three months, tadpoles must overwinter at least once and may take 2 or 4 years of growth before they are large enough to transform (Wright and Wright 1949; Zweifel 1955).

  • Wright, A. H. and Wright, A. A. (1949). Handbook of Frogs and Toads of the United States and Canada. Comstock Publishing Company, Inc., Ithaca, New York.
  • Stebbins, R.C. (1951). Amphibians of Western North America. University of California Press, Berkeley.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Berger, L., Speare, R., Daszak, P., Green, D. E., Cunningham, A. A., Goggin, C. L., Slocombe, R., Ragan, M. A., Hyatt, A. D., McDonald, K. R., Hines, H. B., Lips, K. R., Marantelli, G., and Parkes, H. (1998). "Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America." Proceedings of the National Academy of Sciences of the United States of America, 95(15), 9031-9036.
  • Storer, T. I. (1925). "A synopsis of the amphibia of California." University of California Publications in Zoology, 27, 1-342.
  • Jennings, M. R., and Hayes, M. P. (1994). ''Amphibian and reptile species of special concern in California.'' Final Report #8023 Submitted to the California Department of Fish and Game. California Department of Fish and Game, Sacramento, California..
  • Drost, C. A., and Fellers, G. M. (1996). "Collapse of a regional frog fauna in the Yosemite area of the California Sierra Nevada, USA." Conservation Biology, 10(2), 414-425.
  • Bradford, D. F. (1989). "Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California: implication of the negative effect of fish introductions." Copeia, 1989, 775-778.
  • Bradford, D. F. (1989). ''Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California USA: Implication of the negative effect of fish introductions.'' Copeia, 1989(3), 775-778.
  • Bradford, D. F., Tabatabai, F., and Graber, D. M. (1993). ''Isolation of remaining populations of the native frog, Rana muscosa, by introduced fishes in Sequoia and Kings Canyon National Parks, California.'' Conservation Biology, 7, 882-888.
  • Briggs, C. J., Knapp, R. A., and Vredenburg, V. T. (2010). ''Enzootic and epizootic dynamics of the chytrid fungal pathogen of amphibians.'' Proceedings of the National Academy of Sciences, 107(21), 9695-9700 .
  • Davidson, C. (2004). ''Declining downwind: Amphibian population declines in California and historical pesticide use.'' Ecological Applications, 14, 1892-1902.
  • Davidson, C., Shaffer, H. B., and Jennings, M. R. (2002). ''Spatial tests of the pesticide drift, habitat destruction, UV-B, and climate-change hypotheses for California amphibian declines.'' Conservation Biology, 16, 1588-1601.
  • Finlay, J. and Vredenburg, V. T. (2007). ''Introduced trout sever trophic connections between lakes and watersheds: consequences for a declining montane frog.'' Ecology, 88(9), 2187-2198.
  • Grinnell, J., and Storer, T. I. (1924). Animal Life in the Yosemite. University of California Press, Berkeley, California.
  • Knapp, R. A. and Matthews, F. (2000). ''Non-native fish introductions and the decline of the Mountain Yellow-legged Frog from within protected areas.'' Conservation Biology, 14(2), 428-439.
  • Knapp, R. A., Boiano, D. M., Vredenburg, V. T. (2007). ''Recovery of a declining amphibian (Mountain Yellow-legged Frog, Rana muscosa) following removal of non-native fish.'' Biological Conservation, 135, 11-20.
  • Knapp, R.A. (1996). ''Non-native trout in the natural lakes of the Sierra Nevada: an analysis of their distribution and impacts on native aquatic biota.'' Sierra Nevada Ecosystem Project, Final Report to Congress, Center for Water and Wildland Resources, University of California (Davis), Davis, California, 363-390.
  • Livezey, R. L., and Wright, A. H. (1945). ''Descriptions of four salientian eggs.'' American Midland Naturalist, 34, 701-706.
  • Rachowicz, L. J., Knapp, R. A., Morgan, J. A. T., Stice, M. J., Vredenburg, V. T., Parker, J. M., and Briggs, C. J. (2006). ''Emerging infectious disease as a proximate cause of amphibian mass mortality.'' Ecology, 87, 1671-1683.
  • Vredenburg, V. T. (2004). ''Reversing introduced species effects: Experimental removal of introduced fish leads to rapid recovery of a declining frog.'' Proceedings of the National Academy of Sciences of the United States of America, 101, 7646-7650.
  • Vredenburg, V. T., (2007). ''Concordant molecular and phenotypic data delineate new taxonomy and conservation priorities for the endangered mountain yellow-legged frog (Ranidae: Rana muscosa).'' Journal of Zoology, 271, 361-374.
  • Vredenburg, V. T., Fellers, G., and Davidson, C. (2005). ''The mountain yellow-legged frog Rana muscosa (Camp 1917).'' Status and conservation of U.S. Amphibians. M. Lannoo, eds., University of California Press, Berkeley, 563-566.
  • Vredenburg, V. T., Knapp, R. A., Tunstall, T. S., and Briggs, C. J. (2010). ''Dynamics of an emerging disease drive large-scale amphibian population extinctions.'' Proceedings of the National Academy of Sciences, 107(21), 9689-9694.
  • Zweifel, R. G. (1955). ''Ecology, distribution, and systematics of frogs of the Rana boylei group.'' University of California Publications in Zoology, 54, 207-292.
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Threats

Degree of Threat: High

Comments: Decline has been attributed to the effects of the amphibian chytrid fungus, introduced trouts, pesticide exposure, or a combination of factors (e.g., Bradford 1989; Knapp and Matthews 2000; Fellers et al. 2001, 2013; Davidson et al. 2002; Davidson and Knapp 2007; Sodhi et al. 2008; Sparling and Fellers 2009; Briggs et al. 2010; Vredenburg et al. 2010; California Department of Fish and Game 2011).

Extensive surveys in the Sierra Nevada clearly demonstrate the strong detrimental impact of introduced trouts on R. muscosa/Rana sierrae populations (Bradford 1989, Knapp and Matthews 2000). Removal of non-native fishes (relatively easy in some Sierra Nevada lakes) might easily reverse the decline (Knapp and Matthews 2000).

Some declines have been associated with amphibian chytrid fungus (Vredenburg et al. 2010), but at least one population has remained relatively stable despite the presence of the fungus for many years (Fellers et al. 2013).

Davidson et al. (2002) found support for the hypothesis that airborne agrochemicals have played a significant role in the decline of frogs of the Rana muscosa/Rana sierrae complex, but subsequent study of pesticide presence versus yellow-legged frog population trends (Bradford et al. 2011) found no support for the hypothesis that pesticides have contributed to the population declines of R. muscosa and R. sierrae in the alpine zone of the southern Sierra Nevada.

Climate change may detrimentally affect this species if drought increases.

Wildfires, which may increase in extent, frequency, or intensity with climate change, also may result in habitat loss and degradation and presumably could result in extirpation or decreases of small localized yellow-legged frog populations.

See Bradford (1991) for information on mass mortality and extinction of a population due at least in part to red-leg disease and predation on metamorphics by Brewer's blackbird; reestablishment of the extirpated population probably will be prevented through predation by introduced fishes.

Frogs of the Rana muscosa/Rana sierrae complex are possibly but probably not threatened by sublethal effects of low pH and elevated levels of dissolved aluminum (Bradford et al. 1992).

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Major Threats
A petition to list the Rana muscosa/Rana sierrae complex as endangered cited the following threats: non-native fish introductions, contaminant introductions, livestock grazing, acidification from atmospheric deposition, nitrate deposition, ultraviolet radiation, drought, disease, and other factors (see USFWS 2000).

Extensive surveys in the Sierra Nevada clearly demonstrate the strong detrimental impact of introduced trouts on R. muscosa/Rana sierrae populations (Bradford 1989, Knapp and Matthews 2000). Removal of non-native fishes (relatively easy in some Sierra Nevada lakes) might easily reverse the decline (Knapp and Matthews 2000).

See Bradford (1991) for information on mass mortality and extinction of a population due at least in part to red-leg disease and predation on metamorphics by Brewer's blackbird; reestablishment of the extirpated population probably will be prevented through predation by introduced fishes.

Frogs of the Rana muscosa/Rana sierrae complex are possibly but probably not threatened by sublethal effects of low pH and elevated levels of dissolved aluminum (Bradford et al. 1992).

Fellers et al. (2001) documented oral chytridiomycosis (often indicated by oral disc abnormalities) in larvae and recently metamorphosed individuals of the Rana muscosa/Rana sierrae complex in the Sierra Nevada, where recent declines have occurred. However, loss of pigmentation of larval mouthparts does not always indicate chytridiomycosis (Batrachochytrium infection) (Rachowicz 2002).

Davidson et al. (2002) found support for the hypothesis that airborne agrochemicals have played a significant role in the decline of frogs of the Rana muscosa/Rana sierrae complex.
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Life History, Abundance, Activity, and Special Behaviors

Rana sierrae is critically endangered, along with its sister species Rana muscosa. These frogs have declined dramatically despite the fact that most of the habitat is protected in National Parks and National Forest lands. A study that compares recent surveys (1995-2005) to historical localities (1899-1994; specimens from the Museum of Vertebrate Zoology and the California Academy of Sciences) found that 92.5% of populations have gone extinct (11 remaining out of 146 sites; Vredenburg et al. 2007).

The two most important factors leading to declines in R. sierrae and R. muscosa are disease and introduced predators.

Introduced trout prey on R. sierrae (Needham and Vestal 1938; Mullally and Cunningham 1956)and have been implicated in a number of studies as one of the sources of decline (Bradford 1989; Bradford et al. 1993; Jennings 1994; Knapp 1996; Drost and Fellers 1996; Knapp and Matthews 2000). In fact, as early as 1915 Joseph Grinnell and his field crews (Grinnell and Storer 1924) noticed that Rana sierrae rarely survived in lakes where trout were planted. Whole lake field experiments have shown that when non-native trout are removed, both Rana sierrae and Rana muscosa populations rebound (Vredenburg, 2004; Knapp et al. 2007). While it is clear that introduced trout negatively affect R. sierrae and R. muscosa mainly through predation on tadpoles, trout also compete for resources with adult frogs. A food web study that used stable isotopes to trace energy through the Sierran lake food webs concluded that introduced trout are superior competitors and suppress the availability of large aquatic insects that make up a major portion of the diets of adult frogs (Finlay and Vredenburg 2007).

A lethal disease, chytridiomycosis, caused by an aquatic fungal pathogen Batrachochytrium dendrobatidis, or Bd (Berger et al. 1998) has caused population extinctions in R. muscosa and R. sierrae in the Sierra Nevada (Rachowicz et al. 2006). Long-term studies reveal that infection intensity is key; once a critical threshold of Bd fungal infection is reached, death ensues (Vredenburg et al. 2010). Population extirpation is the most common outcome, but a few mountain yellow-legged frog (Rana sierrae and Rana muscosa) populations have survived in low numbers. Modeling shows that chytriodiomycosis outcome at the population level (extirpation vs. persistence) can result solely from density-dependent host-pathogen dynamics, which may hold for other wildlife diseases as well (Briggs et al. 2010). In an effort to rescue the last surviving frogs, the Vredenburg lab is treating adult Rana sierrae in the field with anti-fungal medication; frogs are bathed for five minutes daily over the course of a week (Lubick 2010).

Other possible causes for decline in R. sierrae include air pollution (pesticide drift; Davidson et al. 2002; Davidson 2004), UV-B radiation, and long term changes in weather patterns, especially concerning the severity and duration of droughts. Acidification from atmospheric deposition has been suggested as another cause, but Bradford et al. (1994) found no evidence to support this hypothesis.

For more information on active research on this species please see:

Vance Vredenburg

Cherie Briggs

Roland Knapp

  • Wright, A. H. and Wright, A. A. (1949). Handbook of Frogs and Toads of the United States and Canada. Comstock Publishing Company, Inc., Ithaca, New York.
  • Stebbins, R.C. (1951). Amphibians of Western North America. University of California Press, Berkeley.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Berger, L., Speare, R., Daszak, P., Green, D. E., Cunningham, A. A., Goggin, C. L., Slocombe, R., Ragan, M. A., Hyatt, A. D., McDonald, K. R., Hines, H. B., Lips, K. R., Marantelli, G., and Parkes, H. (1998). "Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America." Proceedings of the National Academy of Sciences of the United States of America, 95(15), 9031-9036.
  • Storer, T. I. (1925). "A synopsis of the amphibia of California." University of California Publications in Zoology, 27, 1-342.
  • Jennings, M. R., and Hayes, M. P. (1994). ''Amphibian and reptile species of special concern in California.'' Final Report #8023 Submitted to the California Department of Fish and Game. California Department of Fish and Game, Sacramento, California..
  • Drost, C. A., and Fellers, G. M. (1996). "Collapse of a regional frog fauna in the Yosemite area of the California Sierra Nevada, USA." Conservation Biology, 10(2), 414-425.
  • Bradford, D. F. (1989). "Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California: implication of the negative effect of fish introductions." Copeia, 1989, 775-778.
  • Bradford, D. F. (1989). ''Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California USA: Implication of the negative effect of fish introductions.'' Copeia, 1989(3), 775-778.
  • Bradford, D. F., Tabatabai, F., and Graber, D. M. (1993). ''Isolation of remaining populations of the native frog, Rana muscosa, by introduced fishes in Sequoia and Kings Canyon National Parks, California.'' Conservation Biology, 7, 882-888.
  • Briggs, C. J., Knapp, R. A., and Vredenburg, V. T. (2010). ''Enzootic and epizootic dynamics of the chytrid fungal pathogen of amphibians.'' Proceedings of the National Academy of Sciences, 107(21), 9695-9700 .
  • Davidson, C. (2004). ''Declining downwind: Amphibian population declines in California and historical pesticide use.'' Ecological Applications, 14, 1892-1902.
  • Davidson, C., Shaffer, H. B., and Jennings, M. R. (2002). ''Spatial tests of the pesticide drift, habitat destruction, UV-B, and climate-change hypotheses for California amphibian declines.'' Conservation Biology, 16, 1588-1601.
  • Finlay, J. and Vredenburg, V. T. (2007). ''Introduced trout sever trophic connections between lakes and watersheds: consequences for a declining montane frog.'' Ecology, 88(9), 2187-2198.
  • Grinnell, J., and Storer, T. I. (1924). Animal Life in the Yosemite. University of California Press, Berkeley, California.
  • Knapp, R. A. and Matthews, F. (2000). ''Non-native fish introductions and the decline of the Mountain Yellow-legged Frog from within protected areas.'' Conservation Biology, 14(2), 428-439.
  • Knapp, R. A., Boiano, D. M., Vredenburg, V. T. (2007). ''Recovery of a declining amphibian (Mountain Yellow-legged Frog, Rana muscosa) following removal of non-native fish.'' Biological Conservation, 135, 11-20.
  • Knapp, R.A. (1996). ''Non-native trout in the natural lakes of the Sierra Nevada: an analysis of their distribution and impacts on native aquatic biota.'' Sierra Nevada Ecosystem Project, Final Report to Congress, Center for Water and Wildland Resources, University of California (Davis), Davis, California, 363-390.
  • Livezey, R. L., and Wright, A. H. (1945). ''Descriptions of four salientian eggs.'' American Midland Naturalist, 34, 701-706.
  • Rachowicz, L. J., Knapp, R. A., Morgan, J. A. T., Stice, M. J., Vredenburg, V. T., Parker, J. M., and Briggs, C. J. (2006). ''Emerging infectious disease as a proximate cause of amphibian mass mortality.'' Ecology, 87, 1671-1683.
  • Vredenburg, V. T. (2004). ''Reversing introduced species effects: Experimental removal of introduced fish leads to rapid recovery of a declining frog.'' Proceedings of the National Academy of Sciences of the United States of America, 101, 7646-7650.
  • Vredenburg, V. T., (2007). ''Concordant molecular and phenotypic data delineate new taxonomy and conservation priorities for the endangered mountain yellow-legged frog (Ranidae: Rana muscosa).'' Journal of Zoology, 271, 361-374.
  • Vredenburg, V. T., Fellers, G., and Davidson, C. (2005). ''The mountain yellow-legged frog Rana muscosa (Camp 1917).'' Status and conservation of U.S. Amphibians. M. Lannoo, eds., University of California Press, Berkeley, 563-566.
  • Vredenburg, V. T., Knapp, R. A., Tunstall, T. S., and Briggs, C. J. (2010). ''Dynamics of an emerging disease drive large-scale amphibian population extinctions.'' Proceedings of the National Academy of Sciences, 107(21), 9689-9694.
  • Zweifel, R. G. (1955). ''Ecology, distribution, and systematics of frogs of the Rana boylei group.'' University of California Publications in Zoology, 54, 207-292.
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Management

Restoration Potential: Rana muscosa/Rana sierrae populations that have been extirpated or reduced as a result of fish introduction can recover to predisturbance levels after fish disappear, if a nearby source population of frogs exists (Knapp et al. 2001). Several agencies (National Park Service, CDFG and U.S. Forest Service) have begun and/or planned recovery efforts involving removal of introduced fishes, and a number of populations have recovered (Vredenburg 2004).

Preserve Selection and Design Considerations: Basins with a variety of deep lakes and shallow ponds may be the most appropriate reserves for this declining species (Pope and Matthews 2001).

Management Requirements: Conservation recommendations for the southern California population include: (1) Installation of signage along trails adjacent to occupied areas to encourage the public to remain on designated trails; (2) removal of picnic equipment or campsites (barbeque pits, picnic tables) adjacent to occupied areas; (3) organization of workshops to educate campground permittees about this population; (4) acquisition of habitat within private inholdings; (5) assignment of additional patrols to prevent illegal suction dredge mining within the Sheep Mountain Wilderness Area of Angeles National Forest; and (6) relocation of a trail adjacent to an area within Little Rock Canyon (USFWS 2002).

Translocation of adults in the Rana muscosa/Rana sierrae complex may not be an effective conservation tool; frogs may return to original capture site and are stressed by translocation (Matthews 2003).

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Needs: Introductions of non-native fishes in lakes and ponds should be avoided.

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Conservation Actions

Conservation Actions
Most occurrences are on lands administered by the U.S. Forest Service or U.S. National Park Service. However, occurrence in protected, pristine areas does not ensure population persistence.
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Relevance to Humans and Ecosystems

Risks

Stewardship Overview: California Department of Fish and Game (2011) made the following conservation recommendations for Rana sierrae and Rana muscosa:

Continue resource assessment efforts, throughout the historical range, to discover previously unknown populations. Special focus should be given those watersheds that have not been surveyed within the past 10 years but have historical occurrences, or high probability of occurrence as determined by the species distribution model.

Continue to implement and support projects that stabilize existing populations and/or expand distribution within each of the six genetic clades, such as: removing non-native trout from targeted water bodies to benefit frog populations; identifying water bodies whose non-native fish population will likely expire naturally and provide fish free habitat for frog translocations; removing non-native trout from targeted water bodies to provide fish free habitat for frog translocations.

Continue to manage fisheries within the historical range of frogs in a manner that does not conflict with frog conservation goals, such as: halting fish stocking in areas harboring existing frog populations or in areas identified for native species management in an Aquatic Biodiversity Management Plan' evaluating current stocking techniques and protocols to minimize stocking related impacts on frogs; evaluating potential impacts to non-game species, via the PSEP process, as mandated by the Hatchery and Stocking Program Environmental Impact Report/Environmental Impact Statement.

Continue monitoring frog population trends and Bd infection levels to establish long-term population baselines and evaluate conservation efforts.

Continue activities that support research directed at frog conservation goals, with special focus given research directed at translocating frogs in a Bd positive environment and captive breeding and rearing, such as: sharing California Department of Fish and Game data sets and analyses with the research community; collaborating with scientists to implement translocations in a manner appropriate to test a hypothesis; continuing to issue scientific collecting permits for research critical to the conservation and recovery of the frogs.

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Relation to Humans

Mountain yellow-legged frogs (the amphibian species complex including both Rana muscosa and Rana sierrae) were once the most common vertebrates in the high elevation Sierra Nevada. Documented historical accounts go back to the turn of the last century (1915) from surveys conducted by Joseph Grinnell and Tracy Storer (published in 1924) from the University of California's Museum of Vertebrate Zoology. Joseph Grinnell was instrumental in the foundation of Yosemite National Park, one of the jewels of the American National Park Service.

  • Wright, A. H. and Wright, A. A. (1949). Handbook of Frogs and Toads of the United States and Canada. Comstock Publishing Company, Inc., Ithaca, New York.
  • Stebbins, R.C. (1951). Amphibians of Western North America. University of California Press, Berkeley.
  • Stebbins, R. C. (2003). Western Reptiles and Amphibians, Third Edition. Houghton Mifflin, Boston.
  • Berger, L., Speare, R., Daszak, P., Green, D. E., Cunningham, A. A., Goggin, C. L., Slocombe, R., Ragan, M. A., Hyatt, A. D., McDonald, K. R., Hines, H. B., Lips, K. R., Marantelli, G., and Parkes, H. (1998). "Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America." Proceedings of the National Academy of Sciences of the United States of America, 95(15), 9031-9036.
  • Storer, T. I. (1925). "A synopsis of the amphibia of California." University of California Publications in Zoology, 27, 1-342.
  • Jennings, M. R., and Hayes, M. P. (1994). ''Amphibian and reptile species of special concern in California.'' Final Report #8023 Submitted to the California Department of Fish and Game. California Department of Fish and Game, Sacramento, California..
  • Drost, C. A., and Fellers, G. M. (1996). "Collapse of a regional frog fauna in the Yosemite area of the California Sierra Nevada, USA." Conservation Biology, 10(2), 414-425.
  • Bradford, D. F. (1989). "Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California: implication of the negative effect of fish introductions." Copeia, 1989, 775-778.
  • Bradford, D. F. (1989). ''Allotopic distribution of native frogs and introduced fishes in high Sierra Nevada lakes of California USA: Implication of the negative effect of fish introductions.'' Copeia, 1989(3), 775-778.
  • Bradford, D. F., Tabatabai, F., and Graber, D. M. (1993). ''Isolation of remaining populations of the native frog, Rana muscosa, by introduced fishes in Sequoia and Kings Canyon National Parks, California.'' Conservation Biology, 7, 882-888.
  • Briggs, C. J., Knapp, R. A., and Vredenburg, V. T. (2010). ''Enzootic and epizootic dynamics of the chytrid fungal pathogen of amphibians.'' Proceedings of the National Academy of Sciences, 107(21), 9695-9700 .
  • Davidson, C. (2004). ''Declining downwind: Amphibian population declines in California and historical pesticide use.'' Ecological Applications, 14, 1892-1902.
  • Davidson, C., Shaffer, H. B., and Jennings, M. R. (2002). ''Spatial tests of the pesticide drift, habitat destruction, UV-B, and climate-change hypotheses for California amphibian declines.'' Conservation Biology, 16, 1588-1601.
  • Finlay, J. and Vredenburg, V. T. (2007). ''Introduced trout sever trophic connections between lakes and watersheds: consequences for a declining montane frog.'' Ecology, 88(9), 2187-2198.
  • Grinnell, J., and Storer, T. I. (1924). Animal Life in the Yosemite. University of California Press, Berkeley, California.
  • Knapp, R. A. and Matthews, F. (2000). ''Non-native fish introductions and the decline of the Mountain Yellow-legged Frog from within protected areas.'' Conservation Biology, 14(2), 428-439.
  • Knapp, R. A., Boiano, D. M., Vredenburg, V. T. (2007). ''Recovery of a declining amphibian (Mountain Yellow-legged Frog, Rana muscosa) following removal of non-native fish.'' Biological Conservation, 135, 11-20.
  • Knapp, R.A. (1996). ''Non-native trout in the natural lakes of the Sierra Nevada: an analysis of their distribution and impacts on native aquatic biota.'' Sierra Nevada Ecosystem Project, Final Report to Congress, Center for Water and Wildland Resources, University of California (Davis), Davis, California, 363-390.
  • Livezey, R. L., and Wright, A. H. (1945). ''Descriptions of four salientian eggs.'' American Midland Naturalist, 34, 701-706.
  • Rachowicz, L. J., Knapp, R. A., Morgan, J. A. T., Stice, M. J., Vredenburg, V. T., Parker, J. M., and Briggs, C. J. (2006). ''Emerging infectious disease as a proximate cause of amphibian mass mortality.'' Ecology, 87, 1671-1683.
  • Vredenburg, V. T. (2004). ''Reversing introduced species effects: Experimental removal of introduced fish leads to rapid recovery of a declining frog.'' Proceedings of the National Academy of Sciences of the United States of America, 101, 7646-7650.
  • Vredenburg, V. T., (2007). ''Concordant molecular and phenotypic data delineate new taxonomy and conservation priorities for the endangered mountain yellow-legged frog (Ranidae: Rana muscosa).'' Journal of Zoology, 271, 361-374.
  • Vredenburg, V. T., Fellers, G., and Davidson, C. (2005). ''The mountain yellow-legged frog Rana muscosa (Camp 1917).'' Status and conservation of U.S. Amphibians. M. Lannoo, eds., University of California Press, Berkeley, 563-566.
  • Vredenburg, V. T., Knapp, R. A., Tunstall, T. S., and Briggs, C. J. (2010). ''Dynamics of an emerging disease drive large-scale amphibian population extinctions.'' Proceedings of the National Academy of Sciences, 107(21), 9689-9694.
  • Zweifel, R. G. (1955). ''Ecology, distribution, and systematics of frogs of the Rana boylei group.'' University of California Publications in Zoology, 54, 207-292.
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Wikipedia

Sierra Nevada yellow-legged frog

The Sierra Nevada yellow-legged frog[2] or Sierra Nevada Mountain yellow-legged frog, Rana sierrae, is a true frog endemic to the Sierra Nevada mountains (in California and Nevada).

Contents

Physical description[edit]

This species is very similar in appearance to the southern mountain yellow-legged frog.[2]

Conservation status[edit]

Shallow lakes that dry in summer endanger this high-elevation frog.[2] Predation by non-native trout also plays a large role in limiting breeding and tadpole development.[2] Many factors endangering the southern mountain yellow-legged frog also affect this species.

See also[edit]

Notes[edit]

  1. ^ Geoffrey Hammerson (2008). Rana sierrae. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 12 Dec 2008.
  2. ^ a b c d Igor Lacan, Kathleen Matthews, Krishna Feldman. "Interaction of an introduced predator with future effects of climate change in the recruitment dynamics of the imperiled Sierra Nevada yellow-legged frog (rana sierrae)" (PDF). Herpetological Conservation and Biology 3 (2): 211–223. 
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Names and Taxonomy

Taxonomy

Comments: Yellow-legged frog populations now recognized as Rana sierrae formerly were included in Rana muscosa. Vredenburg et al. (2007) examined phylogeography of Rana muscosa as defined by Stebbins (2003) and determined that R. muscosa occurs in the southern Sierra Nevada and in mountains to the south and that populations in the Sierra Nevada north of this range comprise a distinct species (Rana sierrae).

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