9—
Amphibians

J. Robert Macey and Theodore J. Papenfuss

Introduction

Amphibians are cold-blooded vertebrates with moist, scaleless skin. These animals either occur in close association with permanent water or are active only during wet times of the year. There are approximately 3,200 species in the world, and amphibians are present on all continents except Antarctica.

Amphibians are divided into three groups: caecilians, salamanders, and frogs and toads. Caecilians are elongate, "wormlike," burrowing or aquatic forms that lack limbs and functional eyes. The approximately 165 species are restricted to the tropical regions of the world; no species occur in the United States. Most caecilians are less than 2 ft (0.6m) long; however, one species in South America reaches a length of 5 ft (1.5m).

Most salamanders have a lizardlike shape with four legs and a tail. They have smooth, moist skin that lacks scales. In the New World there are three areas with high species diversity. One is the mountainous region of southern Mexico and northern Central America, with some 50 species. Another is the Appalachian Mountains of the eastern United States, with about 45 species. The third region comprises the Pacific states of California, Oregon, and Washington, with about 35 species.

The species that are present in California have two very different developmental patterns. Some breed in water, and the adults return to streams and ponds during the rainy season to lay eggs. Gilled larvae develop in the water and later transform into terrestrial adult salamanders. Other salamanders are completely terrestrial. The eggs are laid on land in moist places. The larval stage takes place in the egg, and small, fully transformed salamanders emerge. All western fully terrestrial salamanders lack lungs, and respiration takes place through the skin.

Three species of salamanders are known to occur in the California portion of the White-Inyo mountains region. The only state with no native salamanders is Nevada. There have been several reports of salamanders in Nevada, but none have been confirmed. Although the species accounts included here for salamanders have been brought up to date, they are probably incomplete and may even be lacking a species. We would appreciate receiving any additional information, such as a new distribution or a salamander that does not fit the descriptions.

The frogs and toads are the largest group of amphibians. Approximately 2,600 species of frogs and toads are known, and they occur on every continent except Antarctica. The greatest species diversity is in the tropics, but frogs and toads are

Map 9.1

Map 9.2

well represented in temperate regions. These amphibians occur in a great variety of habitats. Some species occur in arid regions, where they spend most of their lives underground, coming to the surface only during occasional rains. Others are fully aquatic and never leave the water. A few species in tropical regions spend their entire lives in the tree canopy. There are 23 families of frogs and toads worldwide, of which 5 occur in California and Nevada. The families that occur in the area under study are the True Toads (Family Bufonidae), Treefrogs (Hylidae), Spadefoot Toads (Pelobatidae), and True Frogs (Ranidae).

Species Accounts

The following species accounts cover the three salamanders and seven frogs and toads that occur in the White-Inyo mountains region. Accounts are ordered by main group, then alphabetically by family name, and finally alphabetically by species name. Each account consists of a brief identification, information on habitat, and a remarks section, which treats natural history. A brief summary of the range and a list of the exact localities corresponding to the dots on the range maps are also included. When the locality is vague, it is not plotted on a map. In these lists the first reference to a place name is listed in its entirety, and all other references to that place name follow it as separate localities. These localities are based on museum specimens. The majority of specimens from this region are deposited in the Museum of Vertebrate Zoology, University of California at Berkeley (MVZ). Other museums represented in the localities sections are: American Museum of Natural History, New York (AMNH); California Academy of Sciences, San Francisco (CAS); Carnegie Museum of Natural History, Pittsburgh (CMNH); Field Museum of Natural History, Chicago (FMNH); University of Kansas (KU); Natural History Museum of Los Angeles County (LACM); Museum of Comparative Zoology, Harvard University (MCZ); San Diego Natural History Museum (SDSNH); University of Michigan Museum of Zoology (UMMZ); University of Nevada, Reno (UN); and United States National Museum, Washington, D.C. (USNM). The acronym of the museum follows each locality. Absence of an acronym indicates that the locality is based on a specimen from the Museum of Vertebrate Zoology. For accuracy, elevations and distances of localities are cited as recorded by the collector and have not been converted to or from the English or metric system.

The color plates of species will aid in identification. All amphibians in the color plates are from the White-Inyo mountains region except the Northern Leopard Frog (Rana pipiens ), which is from Churchill County, Nevada. For each species, pertinent literature references are listed; the complete citations are at the end of the chapter. General references to the amphibians of the region are: Macey (1986), Papenfuss (1986), Stebbins (1951), and Stebbins (1985). All scientific names are after Collins (1990). In addition, the following chapter of this book, on reptiles, has two sections that contain general information on amphibians of the White-Inyo mountains region.

Lungless Salamanders (Family Plethodontidae)

Kern Plateau Slender Salamander, Batrachosepssp.(no author, not yet described). (Plate 9.1) 3–4 in (7.5–10 cm); dorsal pattern variable; black with a brown middorsal stripe or black with an overlay of silver specks; only four toes on each foot. Habitat: This species occurs around permanent springs and creeks with riparian vegetation. The salamanders live in the daytime under cover objects such as rocks and logs. Remarks: This salamander is related to the Inyo Mountains Slender Salamander (B. campi ) and was the second salamander to be discovered in the White-Inyo mountains region. The Kern Plateau Slender Salamander has a narrower head and shorter legs than the Inyo Mountains Slender Salamander. The ranges of the two species do not overlap. Species of the genus Batrachoseps are the only salamanders in California that have just four toes on each hind foot. Other species of Batrachoseps , known as Slender Salamanders, are widely distributed throughout California. They commonly occur in urban areas under rocks and boards. This species is currently known from the Kern Plateau and the southeastern slopes of the Sierra Nevada, but it may have a more extensive range. Range: Kern Plateau and the eastern slopes of the Sierra Nevada south and west of Owens Lake. Reference: Stebbins (1985).

Inyo Mountains Slender Salamander,Batrachoseps campi(Marlow, Brode & Wake, 1979). (Plates 9.2 [from Hunter Canyon] and 9.3 [from French Spring], Map 9.3) 2–3 in (5–7.5 cm); dorsal pattern variable; background color dark brown to black; in some populations scattered green, lichenlike spots on body; in others the background color is completely overlain with silver flecks; only four toes on each foot. Habitat: Occurs only around permanent springs and seeps that provide a riparian habitat. Inyo Mountains Slender Salamanders are active at night. During the day they take shelter under moist rocks or in damp crevices. They have been found at springs as low as 1,800 ft (550 m) and as high as 8,600 ft (2,600 m). Remarks: The discovery in 1973 of these salamanders in the arid Inyo Mountains was unexpected. In the past, numerous biologists had visited many of the springs where these salamanders live without detecting them, because salamanders are not considered desert animals. Doubtless, additional populations will be found. These salamanders are relicts that entered the Inyo Mountains prior to the rise of the Sierra Nevada and the formation of the Mojave and Great Basin deserts. In some instances these isolated populations may consist of only a few hundred individuals. They are protected by state law and should not be collected. At the present time, capping of springs is the major threat to these populations. Range: Both east and west slopes of the Inyo Mountains. References: Marlow, Brode, and Wake (1979); Yanev and Wake (1981).

Localities: California, Inyo Co.: 6,800–7,000, 8,000–8,300 ft, Addie Canyon; 6,400 ft, Barrel Springs; 6,400 ft, Cove Springs; 3,500 ft, Craig Canyon; 6,000 ft, French Spring (LACM, MVZ); 1,800 ft, 2,200 ft, Hunter Canyon; 4,000 ft, Keynot Canyon; 6,500 ft, Lead Canyon (LACM, MVZ); 5,600 ft, Long John Canyon (LACM, MVZ); 3,500 ft, McElvey Canyon; 7,500 ft, top of ridge between Lead and Addie canyons; 7,200 ft, Waucoba Canyon; 4,700 ft, Willow Creek Canyon.

Map 9.3

Map 9.4

Owens Valley Web-toed Salamander, Hydromantessp. (no author, not yet described). (Plate 9.4) 3 1/2–4 1/2 in (9–11.5 cm); body and head very flattened; dorsal pattern variable; greenish brown to silver with scattered black spots; scattered lichenlike silver spots on ventral surface; feet with extensive webbing; four toes on front feet and five toes on hind feet. Habitat: The Owens Valley Web-toed Salamander occurs in the vicinity of permanent springs and mountain streams with riparian vegetation. Although this salamander is nocturnal, it can be encountered under wood and rocks in areas with moist soil. Remarks: The Genus Hydromantes is distributed in California, Italy, and Sardinia. The species in Europe are the only members of the lungless salamander family, Plethodontidae, in the Old World. The Owens Valley Web-toed Salamander was discovered in 1985 during field work in preparation for this chapter.

Range: Eastern slopes of the Sierra Nevada, at least from the area around Owens Lake to Big Pine. Reference: Wake, Maxson, and Wurst (1978).

True Toads (Family Bufonidae)

Western Toad,Bufo boreas(Baird & Girard, 1852). (Plate 9.5, Map 9.4) 3–5 1/2 in (7.5–13.75 cm); marbled dorsal pattern of brown, gray, and green in equal proportions; distinct white or light yellow line down middle of back; ventral background color cream with black spots; small warts scattered over back; oblong gland (parotoid gland) behind eye is longer than upper eyelid. Habitat: In White-Inyo Range, occurs around permanent ponds and slow-moving streams. Western Toads are generally nocturnal; however, recently discovered populations in the northern White Mountains (see Map 9.4) are also active during the day, at least in spring and summer. Remarks: Western Toads are a common, wide-ranging species in much of the western United States, occurring from sea level to above 9,000 ft (2,740 m). They are often seen at night on roads and in yards in rural areas. In the White-Inyo mountains region, this species is generally restricted to valleys, but it has been found above 7,000 ft (2,130 m) in the Pinyon-juniper Woodland of the northern White Mountains. A Creosote Bush Scrub outpost for this species is Darwin Falls Canyon in the northern Argus Mountains. Here the Western Toad and the Red-spotted Toad (B. punctatus ) coexist and occasionally hybridize (see Red-spotted Toad account). At a second isolated locality, Fish Lake in Fish Lake Valley, Western Toads may be extinct due to the introduction of Bullfrogs (Rana catesbeiana ) (see Bullfrog account). Range: Darwin Falls Canyon, northern Argus Mountains; Owens Valley; vicinity of Fish Lake, Fish Lake Valley; extreme northern White Mountains. Reference: Karlstrom (1962).

Localities: California, Inyo Co.: Alvord (USNM); 3.0 mi S Bartlet; Batchelder Spring, Westgard Pass; Big Pine; 1.5 mi SW; 4 mi NW (JACM); Bishop; 0.5 mi NW (UMMZ); 5 mi E (LACM); 4000 ft, Bishop Creek (USNM); Darwin Falls, Argus Mtns.; Diaz Lake, Owens Valley (CAS); Fish Lake Spring (CAS); Independence; Laws (CAS, MVZ); near (UMNZ); Lone Pine (MCZ, MVZ, USNM); 2.9 mi S (CAS). Mono Co.: Benton; 5 mi N (SDSNH); 3 mi from Nevada State line, spring, Taylor Ranch, 5 mi N Benton (UMMZ). Nevada, Esmeralda Co.: Fish Lake (MVZ, UMMZ); 7,820 ft, Buffalo Canyon. Mineral Co.: Orchard Spring, Buffalo Canyon; Queen Canyon.

Black Toad,Bufo exsul(Myers, 1942). (Plate 9.6, Map 9.5) 1 1/2–2 1/2 in (3.75–6.25 cm); dorsal color predominantly black; faint white line down middle of back; ventral color black, with white mottling becoming more extensive on chin; small warts scattered over back; oblong gland (parotoid gland) behind eye is longer than upper eyelid. Habitat: This species is the most aquatic toad in the region. It never occurs far from permanent water. Black Toads prefer marshy areas around pools or slow-moving streams. They are generally diurnal but may be active at night during the summer. Remarks: The Black Toad along with the three salamanders are the only amphibians endemic to the White-Inyo mountains region. It is native only to the Deep Springs Valley but has been introduced to Batchelder Spring on the west side of

Map 9.5

Map 9.6

Westgard Pass. However, no toads have been seen at Batchelder Spring in recent years, and that population may be extinct. The Black Toad is closely related to the Western Toad (B. boreas ). It has been isolated from the Western Toad for at least 12,000 years, since the last Pleistocene moist period. The Black Toad, and the Amargosa Toad (B. nelsoni ) in Nye County, Nevada, have commonly been regarded as subspecies of the Western Toad. However, a recent genetic study suggests that these two forms be recognized as full species. The only other amphibian in Deep Springs Valley is the Great Basin Spadefoot (Spea intermontana ). Spadefoots have light-colored, smooth skin and spades on the hind feet (see Great Basin Spadefoot Toad account). Black Toads are protected by California law and should not be collected. Range: Areas of permanent

water in the Deep Springs Valley; introduced to Batchelder Spring, Westgard Pass, but may be extinct. References: Myers (1942), Schuierer (1962).

Localities: California, Inyo Co.: Antelope Spring; Deep Springs Valley (CAS, LACM); Batchelder Spring, Westgard Pass; Bog Mounds Spring, Deep Springs Valley; Buckhorn Spring, Deep Springs Valley (CAS, MVZ); Deep Springs (CAS, CMNH, FMNH, SDSNH, USNM, KU, MCZ); 6 mi S (LACM, MVZ); 7 mi S; 7.5 mi S (AMNH, UMMZ); 8 mi S (LACM); Deep Springs Lake (CAS, MCZ, USNM); 0.5 mi E; Warm Spring at Deep Springs Lake.

Red-spotted Toad,Bufo punctatus(Baird & Girard, 1852). (Plate 9.7, Map 9.6) 2–3 in (5–7.5 cm); greenish gray background color; small warts scattered over back, arms, and legs; each wart is reddish brown and surrounded by a black circle; round gland (parotoid gland) slightly smaller than upper eyelid, located behind each eye. Habitat: This toad occurs near springs and semipermanent streams in and around Creosote Bush Scrub. Red-spotted Toads are nocturnal but may be found during the day under rocks adjacent to streams. Remarks: This species can be distinguished from other toads in the area by the small red spots and the round parotoid gland that is smaller than the eyelid. It is a wide-ranging, predominantly Creosote Bush Scrub species that reaches the northern limits of its distribution in our area. At Darwin Falls in the northern Argus Mountains (see Map 9.6), Red-spotted Toads and Western Toads (B. boreas ) occur together. Locally, hybrids between the two species have been found; this is the only reported case in California of hybridization between Red-spotted Toads and other toads. Range: East side of the Argus and Inyo mountains in canyons draining into the Panamint and Saline valleys. Reference: Feder (1979).

Localities: California, Inyo Co.: Darwin Falls, Argus Mtns. (CAS, MVZ); Grapevine Canyon, Nelson Mtns.; Hunter Creek, Saline Valley (CAS, MVZ); below Jackass Spring, Panamint Mtns. (E of study area) (USNM); Pat Keyes Canyon, Saline Valley; Willow Creek, Saline Valley.

Treefrogs (Family Hylidae)

Pacific Treefrog,Pseudacris regilla (Baird & Girard, 1852). (Plate 9.8, Map 9.7) 1–2 in (2.5–5 cm); dorsal pattern variable, gray with brown blotches or uniform green; individual frogs can change color; dark stripe extends from nose through eye and beyond jaw; smooth skin; toe tips enlarged to form small adhesive pads. Habitat: In the White-Inyo Range this species is restricted to permanent streams and marshy areas. Treefrogs are mainly nocturnal and form breeding choruses during the spring and early summer. Some live during the day next to streams and on vegetation in marshes. The adhesive pads on their toes allow them to climb. Remarks: The Pacific Treefrog is a widespread and common frog in California. However, in the White-Inyo mountains region its distribution is limited. It appears to be restricted to the southern Owens Valley from Owens Lake to Independence, and it is present in the Queen Valley. It seems to be absent not only from the rest of the Owens Valley, but also from the White-Inyo Range. Suitable habitat is present in both areas, but

Map 9.7

Map 9.8

no specimens have been reported. It is common, however, in several canyons in the Panamint Mountains. The 1891 U.S. Department of Agriculture Death Valley expedition collected two specimens in Cottonwood Canyon, in the northern Panamint Mountains. This locality is just east of the area covered in this chapter, and it is possible that Pacific Treefrogs may occur in the Panamint Mountain portion (see ? on Map 9.7). Range: Owens Lake to Independence in the southern Owens Valley; Queen Valley. Expected in the northwestern Panamint Mountains. References: Hedges (1986); Jameson, Mackey, and Richmond (1966).

Localities: California, Inyo Co.: 3,800 ft, Cottonwood Canyon, below Jackass Spring, Panamint Mtns. (E of study area) (USNM); 3 mi E Independence; 3 mi N; 0.8 mi S Lone Pine. Mono Co.: 40 N Bishop [ = Bramlett-Taylor Ranch Springs, 8 mi N Benton] (UMMZ).

Spadefoot Toads (Family Pelobatidae)

Great Basin Spadefoot Toad,Spea intermontana(Cope, 1883). (Plate 9.9, Map 9.8) 1–2 1/2 in (2.5–6.25 cm); dorsal background light gray with dark gray spots; skin smooth; a jet black oval protrudes from the bottom of each hind foot; eyes relatively large; fleshy bump; known as a boss, between eyes. Habitat: This species is the only amphibian in the area that is not restricted to the vicinity of permanent water. Typical habitat in our area is Great Basin Scrub below 6,500 ft (1,980 m). However, on the east side of Owens Lake they have been found in Creosote Bush Scrub (see Map 9.8). Spadefoots are strictly nocturnal. They are most active in the fall, in the spring, and during summer rains. Remarks: The smooth skin, black spades, and boss between the eyes distinguish this toad from all others in the region. Spadefoots may be observed on roads at night, most commonly during or after rains. In the day and during dry times of the year, they use their spades to dig backward into the soil. These toads are very resistant to water loss and may remain underground for long periods of time. A related species in southeastern California, the Couch's Spadefoot Toad (S. couchii ), has been known to remain underground for two years waiting for rains. The Great Basin Spadefoot Toad usually breeds during the spring in temporary ponds. The hatching of eggs into tadpoles and the transformation of tadpoles into toads may take place in a few weeks. Although Great Basin Spadefoots have not been observed in the White-Inyo mountains region above 6,500 ft (1,980 m), they have been found at an elevation of 9,200 ft (2,800 m) in the Bodie Hills north of Mono Lake. Range: West side of the White-Inyo Range in the entire Owens Valley, Chalfant Valley, Hammil Valley, Benton Valley, and Queen Valley; south and east side of the White Mountains in the Deep Springs and Fish Lake valleys. Reference: Mayhew (1962).

Localities: California, Inyo Co.: Big Pine; 0.5 mi NE; 5.6 mi NE; 27.8 mi NE; 0.3 mi E; Bishop; 3.0 mi N; Deep Springs (CAS); 7 mi S; Deep Springs Lake; NE end (LACM); Diaz Lake (CAS); 0.5 mi SE of Hwy. 168 on Eureka Valley Rd.; 1.4 mi W Independence; 2.0 mi SE; Lone Pine; 8.1 mi SE; 10.0 mi SE; 1.0 mi S Mono County line on Hwy. 168. Mono Co.: Benton; 3.2 mi N (LACM); 1.6 mi W (LACM). Nevada, Esmeralda Co.: 1 mi W Hwy. 264 along Chiatovich Creek, Fish Lake Valley; 5,000 ft, Chiatovich Ranch, Fish Lake Valley; Dyer; 1.8 mi S; 4.0 mi N; Fish Lake (MVZ, UMMZ); 1 mi S.

True Frogs (Family Ranidae)

Bullfrog,Rana catesbeiana(Shaw, 1802). (Plate 9.10, Map 9.9) 4–8 in (10–20 cm); background color light to dark green, uniform on back with crossbars on legs; conspicuous large eardrum behind eye. Habitat: This frog prefers slow-moving streams and rivers. It is also common in marshes associated with permanent ponds and lakes. Bullfrogs are active both during the day and at night. They are very wary, and when approached they commonly escape by jumping from the bank into the water. Remarks: Bullfrogs are not native to the area. They were originally introduced from the eastern United States to parts of western North America to provide a fresh source of meat. Natural expansion of this frog's range and further introductions have greatly

Map 9.9

Map 9.10

increased the area in which Bullfrogs live. Unfortunately, they have had a disastrous effect on native amphibians because they prey on them. A large Bullfrog is capable of eating the young and even adults of the native frogs and toads. Red-legged Frogs (R. aurora ) and Foothill Yellow-legged Frogs (R. boylei ) are now extinct in many areas of central California due to the presence of Bullfrogs. In our area, they were recently introduced into the Fish Lake Valley, where Western Toads (Bufo boreas ) were formerly common. During the preparation of this chapter, extensive field work was conducted in the Fish Lake Valley, but no Western Toads were seen. Range: Introduced into the Owens Valley; Fish Lake, Fish Lake Valley; marshes around Salt Works, Saline Valley. References: Moyle (1973).

Localities: California, Inyo Co.: Owens River, 2 mi S Independence. Mono Co.: Fish Slough, 0.4 mi N Inyo County line. Nevada, Esmeralda Co.: Fish Lake.

Northern Leopard Frog,Rana pipiens (Schreber, 1782). (Plate 9.11, Map 9.10) 2 1/2–4 in (6.25–11.25 cm); background color gray or tan; distinct dark-brown or black spots over back and sides; spots on legs forming bars; pale-colored ridge runs down each side of the back from top of eyelid to hind leg; ventral surface uniformly pale. Habitat: Northern Leopard Frogs occur around permanent ponds and streams. They are active both during the day and at night. This is a wary species that usually jumps into the water when approached. Remarks: Leopard Frogs, which range from Panama to the Northwest Territories, Canada, are the most widely distributed group of frogs in North America. Until recently they were all considered one species. The Northern Leopard Frog is the widest-ranging species currently recognized in this group; it occurs from New Foundland to the western Great Basin. This species' wide range is presumed to be due to major expansions since the recession of the last glacial period, which started 18,000 years ago. Leopard Frogs are commonly used in biology class dissections and laboratories around the country. This practice has led to many introductions of this species where they did not formerly occur. Range: Northern Owens Valley from Big Pine to north of Bishop; may occur further south in the Owens Valley (see ? on Map 9.10); reported from the east side of the White Mountains below Boundary Peak. References: Hillis, Frost, and Wright (1983); Sage and Selander (1979).

Localities: California, Inyo Co.: 1 mi W Big Pine; 4 mi W Bishop (W of study area); 3.2 mi N (UMMZ); 5 mi N (LACM, MVZ); 8 mi S. Nevada, Esmeralda Co.: 7,100 ft, east side of White Mountains below Boundary Peak (UN).

References

Collins, J. T. 1990. Standard common and current scientific names for North American amphibians and reptiles, 3d ed. Society for the Study of Amphibians and Reptiles, Herpetological Circular no. 19.

Feder, J. H. 1979. Natural hybridization and genetic divergence between the toads Bufo boreas and Bufo punctatus. Evolution 33:1089–1097.

Hedges, B. S. 1986. An electrophoretic analysis of Holarctic hylid frog evolution. Systematic Zoology 35:1–21.

Hillis, D. M., J. S. Frost, and D. A. Wright. 1983. Phylogeny and biogeography of the Rana pipiens complex: A biochemical evaluation. Systematic Zoology 32:132–143.

Jameson, D. L., J. P. Mackey, and R. C. Richmond. 1966. The systematics of the pacific tree frog, Hyla regilla. Proceedings of the California Academy of Sciences 19:551–620.

Karlstrom, E. L. 1962. The toad genus Bufo in the Sierra Nevada of California. University of California Publications in Zoology 62:1–104.

Macey, J. R. 1986. The biogeography of a herpetofaunal transition between the Great Basin and Mojave deserts. In C. A. Hall, Jr. and D. J. Young (eds.), Natural history of the White-Inyo Range, eastern California and western Nevada, and high altitude physiology, pp. 119–128. University of California White Mountain Research Station Symposium, August 23–25, 1985, Bishop, Calif. vol. 1.

Marlow, R. W., J. M. Brode, and D. B. Wake. 1979. A new salamander, genus Batrachoseps, from the Inyo Mountains of California with a discussion of relationships in the genus. Natural History Museum of Los Angeles County, Contributions in Science, no 308.

Mayhew, W. W. 1962. Scaphiopus couchi in California's Colorado Desert. Herpetologica 18:153–61.

Moyle, D. B. 1973. Effects of introduced Bullfrogs, Rana catesbeiana, on the native frogs of the San Joaquin Valley, California. Copeia 1973:18–22.

Myers, G. S. 1942. The Black Toad of Deep Springs Valley, Inyo County, California. Occasional Papers, Museum of Zoology, University of Michigan, no. 469.

Papenfuss, T. J. 1986. Amphibian and reptile diversity along elevational transects in the White-Inyo Range. In C. A. Hall, Jr. and D. J. Young (eds.), Natural history of the White-Inyo Range, eastern California and western Nevada and high altitude physiology, pp. 129–136. University of California White Mountain Research Station Symposium, August 23–25, 1985, Bishop, Calif. vol. 1.

Sage, R. D., and R. K. Selander. 1979. Hybridization between species of the Rana pipiens complex in central Texas. Evolution 33:1069–1088.

Schuierer, F. W. 1962. Remarks upon the natural history of Bufo exsul Myers, the endemic toad of Deep Springs Valley, Inyo County, California. Herpetologica 17:260–266.

Stebbins, R. C. 1951. Amphibians of western North America . University of California Press, Berkeley and Los Angeles.

Stebbins, R. C. 1985. A field guide to western amphibians and reptiles . Houghton Mifflin, Boston.

Wake, D. B., L. R. Maxson, and G. Z. Wurst. 1978. Genetic differentiation, albumin evolution, and their biogeographic implications in Plethodontid salamanders of California and southern Europe. Evolution 32:529–539.

Yanev, K. P., and D. B. Wake. 1981. Genic differentiation in a relic desert salamander, Batrachoseps campi. Herpetologica 37:16–28.