Order Anura - Frogs and Toads

Family Bufonidae - Toads



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Bufo americanus Holbrook - American Toad
Bufo americanus

Description: Bufo americanus is a medium-sized toad, with adult head-body lengths ranging from 5.1 to 9.0 cm. Ground color may be gray, brown, or reddish and, when present, dorsal dark spots usually possess only one or two large warts. The venter is usually light, with chest and upper abdomen dark spotted. Parotoid glands are not in direct contact with postorbital ridges, but are usually connected to them by a spur.

Distribution and Habitat: The American toad is most often encountered during its early spring breeding season and is found statewide. It occurs in a wide variety of woodland and openland habitats that provide either permanent or temporary shallow water areas for breeding. Mathews and Echternacht (1984) reported this toad from above 1650 m in Great Smoky Mountains National Park.

Taxonomy: Two subspecies are recognized in Tennessee. Bufo a. americanus ranges over most of the state (Conant and Collins, 1991), with B. a. charlesmithi Bragg occurring in extreme northwestern Tennessee (Conant and Collins, 1991; Lynch, 1964; Smith, 1961). Collins (1991a) recommends further study of these two subspecies and suggests they may represent two distinct species. Evidence of hybridization of B. americanus with B. woodhousei fowleri has been reported in eastern Tennessee by Johnson (1968), in Montgomery County by Scott and Snyder (1968), in Stewart County by Snyder (1972), and in Hardeman County by Norton and Harvey (1975). In contrast, King (1939) mentions no interbreeding in Great Smoky Mountains National Park where he found both species breeding in the same pond.


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Bufo woodhousei Girard - Woodhouse's Toad
Bufo woodhousei

Description: Mature individuals range from 5.1 to 7.5 cm in head-body length. Ground color is variable and ranges from light gray to brick red. When present, each large dorsal dark spot usually possesses three or more small warts. Venter is usually light, but breast may have a single, central dark spot. Anterior edge of parotoids is usually in direct contact with interorbital crests.

Distribution and Habitat: Bufo woodhousei is a very common species that occurs in a wide array of rural and urban habitats throughout the state. The species may occur as high as 1494 m in the Blue Ridge Mountains of eastern Tennessee (Stevenson, 1959). Breeding typically occurs in temporary or permanent aquatic sites, including ponds, sloughs, rivers, and reservoirs.

Taxonomy: Only one subspecies, B. w. fowleri Hinckley has been reported from Tennessee (Conant and Collins, 1991).


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Family Hylidae - Treefrogs


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Acris crepitans Baird - Northern Cricket Frog
Acris crepitans

Description: This species is a small hylid whose adult head-body length varies from 1.6 to 3.8 cm. Dorsal ground color is highly variable and ranges from light gray to dark brown. A dorsal median green stripe may extend from head to rump. Snout is blunt and a dark triangle usually occurs between the eyes. A dark longitudinal stripe with ragged edges is present on rear of thigh. There is typically a pair of prominent anal warts. Tips of toes are only slightly expanded. Fourth toe on hind foot has 1.5 to 2 phalanges free of webbing.

Distribution and Habitat: Acris crepitans is known to occur throughout most of Tennessee, but, based on current data, may be absent from the northeastern corner of the state. Also, the species is probably absent from elevations above 335 m in the Great Smoky Mountains (Huheey and Stupka, 1967). The northern cricket frog is most often found near permanent bodies of water such as ponds, reservoirs, sloughs, and streams.

Taxonomy: Two subspecies have been listed from Tennessee. According to Conant and Collins (1991), A. c. crepitans ranges in the southeastern two-thirds of Tennessee, while A. c. blanchardi Harper occupies the northwestern one-third.


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Acris gryllus Le Conte - Northern Cricket Frog
Acris gryllus

Description: Acris gryllus is a small frog very similar to A. crepitans. Head-body length for adults ranges from 1.6 to 3.2 cm. Dorsal ground color varies from gray to almost black. A dorsal median green, yellow, or brown stripe may extend from head to rump. As compared to A. crepitans, the snout of A. gryllus is more pointed and the body more slender. A dark triangle may occur between the eyes. A distinct dark longitudinal stripe with smooth edges is present on rear of thigh. Scattered warts in anal region, but no prominent pair of warts near vent. Tips of toes are only slightly expanded. Fourth toe on hind foot has at least 2.5 phalanges free of webbing.

Distribution and Habitat: The southern cricket frog is known from five counties in extreme southwestern Tennessee. Like A. crepitans, A. gryllus occurs near permanent aquatic sites and may occur sympatrically with A. crepitans. However, A. gryllus may also utilize temporary pools. Norton and Harvey (1975) noted that where they occurred together, A. crepitans was usually found near the shoreline of a reservoir while A. gryllus typically occurred in well-drained areas and near roadside pools.

Taxonomy: As mapped by Conant and Collins (1991), only the nominate subspecies occurs in Tennessee. Mount (1975) indicated hybridization with A. crepitans in Alabama. No evidence of hybridization has been reported from Tennessee populations.


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Hyla avivoca Viosca - Bird-Voiced Treefrog
Hyla avivoca

Description: Hyla avivoca is a typical treefrog, with ends of digits expanded into adhesive discs. Head-body length in mature specimens ranges from 2.8 to 4.4 cm. Dorsal ground color may be green, various shades of gray, or nearly black. A dark, irregularly shaped blotch is usually present on dorsum. Dark markings are usually present between the eyes, and limbs are usually marked with dark crossbars. A small, quadrangular light spot is present on each upper jaw below the eyes. Dorsal surface of skin is mostly smooth. Inner surfaces of thighs are washed with light green or pale yellow.

Distribution and Habitat: The bird-voiced treefrog is known from the Coastal Plain of West Tennessee and along the lower Cumberland River in Middle Tennessee to near Ashland City. Based on a preserved specimen, Dunn (1927a) reported H. phaeocrypta Cope from Nashville. The specimen was sent to the Museum of Comparative Zoology (MCZ) and has apparently been lost. Before Viosca (1928) described H. avivoca, H. phaeocrypta was the name applied to what are now known as two species, H. avivoca and H. versicolor. Thus, Dunn's record from Nashville remains questionable. In Tennessee, this hylid occurs in bottomland sloughs and swamps along major rivers and large creeks. It is especially abundant around Reelfoot Lake.

Taxonomy: Only the nominate subspecies occurs in Tennessee (Smith, 1966). Mount (1975) and, to a lesser degree, Smith (1966) question the validity of subspecific designations for this species. Conant and Collins (1991) did not recognize subspecies.


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Hyla cinerea (Schneider) - Green Treefrog
Hyla cinerea

Description: Adults range from 3.2 to 5.7 cm in head-body length. Tips of toes are expanded to form adhesive discs. Dorsal surface is smooth. Dorsal color ranges from light to dark green and may have a few scattered small gold flecks. A lateral, sharply defined light stripe (usually white or cream) extends from front of upper lip to about mid-body. Body form is slender.

Distribution and Habitat: The green treefrog inhabits bottomland swamps and sloughs, primarily in the Coastal Plain of West Tennessee where it is especially common around Reelfoot Lake. Hyla cinerea and H. gratiosa have often been confused in literature reports from Tennessee. Jacob (1980) provided a new distribution record for H. gratiosa based on a Hardeman County specimen in the Memphis State University Museum of Zoology (No. A2142) that was originally labeled and reported by Norton and Harvey (1975) as H. cinerea. Burt (1938) reported H. cinerea from Clarksville, Montgomery County, citing a specimen taken by Howell in 1910 that was deposited in the United States National Museum (USNM) (Burt,1937). Our study of USNM holdings and catalogue listings revealed an individual (No. 48194) of H. gratiosa, collected at Clarksville in 1910, that was incorrectly listed as H. cinerea. Thus Burt's (1938) record is probably invalid. Another museum specimen of H. cinerea (Illinois State Natural History Survey No. 9527) supposedly from Montgomery County is correctly identified; however, the locality data are suspect. According to the INHS catalogue, this specimen was taken near Clarksville during the summer of 1960 by Floyd Ford of Austin Peay State University. Dr. Ford, now deceased, was questioned by Floyd Scott about this record. According to Scott, Dr. Ford did not recall where he collected the specimen. However, he did recall conducting fieldwork at Reelfoot Lake during the summer of 1960 and believed the specimen was probably taken there. Scott has nearly 30 years of field experience in Montgomery County and has never observed H. cinerea there. However, he and a colleague (Scott and Koons, 1993) recently reported the species along the eastern shore of Kentucky Lake in nearby Stewart County. Gentry (1955-1956) reported H. cinerea from temporary sinkhole lakes in Warren County. However, considering the close similarity between H. cinerea and H. gratiosa and that subsequently only H. gratiosa has been documented in Warren and adjacent counties, it seems likely that Gentry's record actually represents H. gratiosa.

Taxonomy: No subspecies are recognized (Conant and Collins, 1991). Hybridization with H. gratiosa has been reported in Florida by Lee (1968) and in Alabama by Mount (1975). No evidence of this hybrid cross was observed in Tennessee.


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Hyla gratiosa Le Conte - Barking Treefrog
Hyla gratiosa

Description:Hyla gratiosa is sometimes confused with H. cinerea. Adult head-body length ranges from 5.1 to 6.7 cm. Tips of toes are expanded into adhesive discs. Dorsal surface is more rugose and body form stockier than in H. cinerea. Dorsal ground color ranges from light to dark green. Round dark dorsal spots may be distinct or barely detectable, and are often lost in preservative. A lateral light line may extend from front of upper lip to mid-body but, in contrast to H. cinerea, its borders are broken and irregular.

Distribution and Habitat: In Tennessee, the distribution and habitat requirements of H. gratiosa are poorly known. The species is currently known from only three possibly disjunct geographic units. Jacob (1980), Heineke and Heineke (1984), Miller and Campbell (1995), and Alan Peterson (pers. comm.) reported specimens from the Coastal Plain in West Tennessee. Jacob's record from Hardeman County was based on a specimen taken by Norton (1971) and incorrectly identified as H. cinerea by Norton and Harvey (1975). Heineke and Heineke found a specimen on a patio in suburban Bartlett, Shelby County. Miller and Campbell found the species in Carroll County and Peterson's daughters found a specimen on a wall outside their house in Chester County. Coastal Plain populations are tentatively considered continuous with those from northwestern Alabama (Mount, 1975). The presence of H. gratiosa in limestone sinkponds on the Pennyroyal Plain (Western Highland Rim) of north-central Tennessee has been well documented by Scott and Harker (1968), Scott and Snyder (1967, 1968), and VanNorman and Scott (1987). VanNorman's and Scott's study indicates that populations of H. gratiosa in north-central Tennessee and south-central Kentucky form a continuous geographic unit that is probably disjunct from the southern part of the species' range. Rossman (1958) and Jordan and Howard (1990) reported the species from two localities in White County. These localities are near the transition from Eastern Highland Rim to Cumberland Plateau. Miller and Miller (1992) and Miller, Casey, and Pritts (1993) discovered barking treefrogs on the Eastern Highland rim in Warren County. On the Cumberland Plateau, populations are known from upland swamps and strip-mine ponds in Van Buren County and from a limestone sinkpond in Franklin County. These Cumberland Plateau and Eastern Highland Rim populations are regarded as continuous with those reported from northeastern Alabama by Mount (1975). The Tennessee Wildlife Resources Agency (1994a) lists H. gratiosa as a species in need of management.

Taxonomy: No subspecies are recognized (Caldwell, 1982).


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Hyla versicolor Le Conte - Gray Treefrog
Hyla chrysoscelis Cope - Cope's Gray Treefrog
Hyla versicolor and Hyla chrysoscelis

Description:Because these two species cannot be reliably separated based on external morphology, and the range of each has not been fully determined in Tennessee, they are treated here as a sibling species pair. In addition, members of this species pair are also easily confused with H. avivoca. The following characteristics are shared by both members of the species pair. Adult head-body lengths range from 3.2 to 5.1 cm. Tips of toes have adhesive discs. Dorsal ground color varies from light gray or light green to dark brown. Large dark blotches of irregular size and shape may occur on dorsum. Limbs are usually marked with dark crossbars. A quadrangular light spot is present on the upper jaw below the eyes. Dorsal skin surface is more rugose than in H. avivoca, and the inner surfaces of thighs are washed with bright yellow or orange (in contrast to the greenish or pale yellowish present in H. avivoca).

Distribution and Habitat: The composite range of the species pair is statewide. Most literature references and museum specimens fail to distinguish the two species. However, recent studies have begun to provide specific data. Burkett (1989) provided tentative distribution maps for both species indicating H. chrysoscelis occurs statewide, while H. versicolor has a very limited, disjunct distribution that includes small areas in extreme southwestern and northeastern Tennessee. Based on nucleoli counts, he identified H. chrysoscelis from Bledsoe, Cheatham, Chester, Crockett, Davidson, Dickson, Franklin, Grundy, Hamilton, Hardeman, Hardin, Knox, Lewis, Marion, McMinn, Monroe, Montgomery, Obion, Perry, Pickett, Polk, Robertson, Stewart, Van Buren, and White counties. Based on call rate, he also identified H. chrysoscelis from McNairy and Shelby counties. Other authors have verified H. chrysoscelis from Reelfoot Lake (Bushnell et al., 1939; Wasserman, 1970), Cumberland and Wilson counties (Wiley, 1982), Warren County (Miller, 1991), and Shelby County (Ritke and Beck, 1991). The most recent study of the species complex (Pritts, 1995) examined chromosome squashes prepared from samples of tadpoles taken in 18 Middle Tennessee counties (Bedford, Cannon, Coffee, Davidson, DeKalb, Franklin, Giles, Jackson, Lincoln, Macon, Marshall, Maury, Moore, Robertson, Rutherford, Trousdale, Williamson, and Wilson). All populations studied had the 2n = 24 chromosome number of H. chrysoscelis. None exhibited the chromosome number characteristic of H. versicolor (4n = 48) or H. chrysoscelis X H. versicolor hybrids (3n = 36). Currently, H. versicolor has only been verified from Shelby County (Ritke et al., 1988; Ritke and Beck, 1991). Its presence at high elevations in northeastern Tennessee is based on speculation (Burkett, 1989). During this study, individuals (not verified to species, but probably H. chrysoscelis) were observed in a wide variety of aquatic


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Hyla versicolor Le Conte - Gray Treefrog
and
Hyla chrysoscelis Cope - Cope's Gray Treefrog

(continued)

habitats in both woodland and open areas. Breeding was observed in swamps, small ponds along the edges of large reservoirs, and in flooded fields and roadside ditches. Ritke and Beck (1991) noted that H. versicolor seemed to prefer bottomland as opposed to upland habitats. Individuals not verified to species have been reported from above 1650 m in the mountains of extreme eastern Tennessee (Mathews and Echternacht, 1984).

Taxonomy: Based on breeding experiments that revealed a high degree of incompatibility between the two species and the existence of different mating call trill rates, Johnson (1966) recognized H. chrysoscelis as a cryptic species. Johnson described a fast trill rate for the mating call of H. chrysoscelis and a slower trill rate for H. versicolor. Later studies have reinforced Johnson's conclusions. Additional characters found useful in separating the two species include chromosome number, cell size, and cell nucleus size and composition. Bogart and Wasserman (1972) showed that H. versicolor is tetraploid (4n = 48) while H. chrysoscelis is diploid (2n = 24). They also noted that blood and sperm cells were larger in H. versicolor. Green (1984) reported larger epidermal cells in the toe pads of H. versicolor. The cell nuclei of H. versicolor were found to be larger and contain more nucleoli than the nuclei of H. chrysoscelis (Cash and Bogart, 1978).









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Pseudacris brachyphona (Cope) - Mountain Chorus Frog
Pseudacris brachyphona

Description: The mountain chorus frog is a small stocky hylid, with an adult head-body length of 2.5 to 3.2 cm. Toe tips are slightly expanded to form adhesive discs. Dorsal ground color is usually brown or gray. A dark triangle typically occurs between the eyes. Dark bars on dorsum may form a reverse parenthesis or H-shaped pattern, but these markings may be broken into irregularly shaped spots or be completely absent. A light stripe is present on the upper lip.

Distribution and Habitat: This small hylid is seldom encountered except during its early spring breeding season. On the Cumberland Plateau, P. brachyphona and Bufo americanus often utilize the same breeding sites. Breeding activity typically occurs in wooded seepage pools, shallow flooded ditches along roads and railroads, small puddles, and shallow ponds. In Tennessee, the mountain chorus frog is known from the Cumberland Mountains, Cumberland Plateau, and Blue Ridge Mountains in extreme northeastern and southeastern Tennessee.

Taxonomy: Hoffman (1980) did not recognize any subspecific variation in this species.


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Pseudacris crucifer (Wied) - Spring Peeper
Pseudacris crucifer

Description: The spring peeper is a small hylid, with an average adult head-body length of 1.9 to 3.2 cm. Tips of toes possess adhesive discs. Dorsal skin surface is smooth. Dorsal ground color ranges from light tan with a pinkish tinge to dark brown. Distinct dark brown markings in the form of an X are usually present on dorsum, and a dark transverse bar is typically present between the eyes.

Distribution and Habitat: Pseudacris crucifer is an early spring breeder that occurs statewide. In Tennessee, the species is very common near almost any type of woodland or brushland aquatic habitat, and breeding individuals especially favor sites bordered by dense vegetation. The species does not appear to be limited by elevation in Tennessee and has been reported from above 1650 m in Great Smoky Mountains National Park (Mathews and Echternacht, 1984).

Taxonomy: Conant and Collins (1991) considered all populations in Tennessee as P. c. crucifer.


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Pseudacris triseriata (Wied) - Upland Chorus Frog
Pseudacris triseriata

Description: This species is similar in size to P. brachyphona, but is more slender. Mature individuals have head-body lengths ranging from 1.9 to 3.5 cm. Toe tips are slightly expanded to form adhesive discs. Light stripe is present on upper lip. A lateral dark line originates on snout and extends through the eye to the groin. Dorsal ground color varies from gray to dark brown. A dark triangle generally occurs between the eyes. Dorsal markings are variable but usually consist of one mid-dorsal and two lateral dark stripes.

Distribution and Habitat: Pseudacris triseriata occurs statewide and occupies woodland and openland habitats that provide suitable breeding sites. Preferred breeding sites include shallow water ponds, flooded woodlands and pastures, and roadside ditches.

Taxonomy: Of the four subspecies of P. triseriata recognized by Conant and Collins (1991), only P. t. feriarum (Baird) occurs in Tennessee. Based on electrophoretic, morphological, and call analyses, Pseudacris t. feriarum is considered by some (Ralin, 1970; Hedges, 1986; Platz and Forester, 1988) as a distinct species.


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Family Microhylidae - Narrowmouth Toads


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Gastrophryne carolinensis (Holbrook) - Eastern Narrowmouth Toad
Gastrophryne carolinensis

Description: Gastrophryne carolinensis is a small stocky anuran, with a small head and distinctly pointed snout. Adult head-body lengths range from 2.2 to 3.2 cm. Legs are short, and there is no webbing between toes. Skin is smooth and dorsal ground color may be gray, brown, or rust. Broad, light dorsolateral stripes are usually present.

Distribution and Habitat: Gastrophryne carolinensis is widespread in Tennessee, but is apparently absent at high elevations of the Blue Ridge Mountains. Martof et al. (1980) and Nelson (1972) provide distribution maps that show G. carolinensis absent from most of the Blue Ridge Mountains in Virginia, North Carolina, Tennessee, and Georgia. Available records for Tennessee indicate the species occurs as high as 549 m in Cades Cove in Great Smoky Mountains National Park (Huheey and Stupka, 1967). This burrowing species is found near reservoirs, ponds, drainage ditches, and sloughs.

Taxonomy: According to Nelson (1972), no subspecies are recognized.


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Family Pelobatidae - Spadefoot Toads


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Scaphiopus holbrookii (Harlan) - Eastern Spadefoot
Scaphiopus holbrookii

Description: The eastern spadefoot is toad-like in appearance. Average adult head-body length varies from 4.4 to 5.7 cm. Cranial crests are absent. Pupils of eyes are vertically elliptical. A dark, elongate, horny spade is present on heel of each foot. Dorsal ground color varies from yellowish brown to dark brown. Lyre-shaped light markings are usually present on dorsum.

Distribution and Habitat: Museum records for Tennessee and the range map for North Carolina provided by Martof et al. (1980), indicate that S. holbrookii occurs throughout Tennessee, with the exception of the Blue Ridge Mountains. Wasserman's (1968) map shows the species as absent from the Cumberland Plateau of Tennessee and Alabama, and Conant and Collins (1991) indicate its absence in central and extreme eastern Tennessee. Museum records, however, indicate that the species is present in all these areas except extreme eastern Tennessee. The eastern spadefoot is a secretive, burrowing species that breeds in temporary pools formed by heavy rains.

Taxonomy: Only the nominate subspecies is known to occur in Tennessee (Wasserman, 1968).


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Family Ranidae - True Frogs


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Rana areolata Baird and Girard - Crawfish Frog
Rana areolata

Description: The crawfish frog is a stocky ranid, with an adult head-body length varying from 5.7 to 7.5 cm. A dorsolateral fold is present along each side of body. Snout is conical and upper jaw is mottled with dark and light markings. Dorsal ground color varies from light gray to off-white. Dorsal markings are profuse, consisting of many round dark spots surrounded by light borders. These are interspersed with smaller dark markings of various shapes. Venter is usually white and immaculate.

Distribution and Habitat: Rana areolata is found in the Coastal Plain of western Tennessee. Although records are lacking for a large region in the Hardeman-McNairy county area, the species is known from just across the state line near Corinth, Mississippi (George Folkerts, pers. comm.). The crawfish frog is wary and difficult to approach. It breeds in flooded pastures and woodlands, farm ponds, and small reservoirs. It often occupies abandoned crayfish burrows.

Taxonomy: Two subspecies are recognized, but only one, R. a. circulosa Rice and Davis, is known from Tennessee (Conant and Collins, 1991; Collins, 1991b).


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Rana capito Le Conte - Gopher Frog
Rana capito

Description: Rana capito resembles R. areolata but averages slightly larger. Adult head-body length ranges from 6.4 to 9.0 cm. A dorsolateral fold is present along each side of body. Dorsal ground color is variable, but is usually brown, dark brown, or nearly black. Dorsal markings normally consist of numerous round dark spots that lack light borders. Venter usually spotted from chin to midbody.

Distribution and Habitat: The species was discovered in Tennessee by Dr. Brian Miller, Middle Tennessee State University, in 1993 in Coffee County. Miller and student Tim Casey found a live individual on a gravel road during a night rain (Brian Miller, pers. comm.). The specimen (MTSU 72A) remains the only known individual reported from the state. Habitat use in Tennessee is unknown, but in adjacent states the species usually breeds in small ponds and flooded areas of fields and woodlands during and after heavy rains. The U. S. Fish and Wildlife Service (1994) listed R.. a. sevosa and R. a. capito, respectively, as Category 1 and Category 2 candidates for federal listing. However, the most recent list of federal candidate species (U.S. Fish and Wildlife Service, 1996) does not include either of these subspecies.

Taxonomy: Currently recognized as constituting a separate species (Collins, 1991b; Conant and Collins, 1991), the three subspecies of gopher frogs (R. c. capito, R. c. sevosa, and R. c. aesopus) were previously considered subspecies of the crawfish frog (R. areolata) (Altig and Lohoefener, 1983). With only one specimen from the state to work with, the subspecific status of the Tennessee population remains uncertain (Brian Miller, pers. comm.).


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Rana catesbeiana Shaw - Bullfrog
Rana catesbeiana

Description: The bullfrog is a large ranid, with mature specimens averaging 9.0 to 15.2 cm in head-body length. Dorsolateral ridges are absent. Dorsal color is typically light to dark green with a highly variable pattern of faint dark markings. Mottling is not present on the upper lip. The tympanic fold is well developed.

Distribution and Habitat: The bullfrog is common throughout Tennessee and occurs in most permanent aquatic habitats including creeks, rivers, ponds, reservoirs, swamps, sloughs, and drainage ditches.

Taxonomy: No subspecies are recognized (Conant and Collins, 1991).

 


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Rana clamitans Latreille - Green Frog
Rana clamitans

Description: Rana clamitans is a medium size frog, with adult head-body lengths ranging from 5.4 to 9.0 cm. Dorsolateral folds are present, but distinct only from head to mid-body. Dorsal ground color may be green, brown, or bronze. Distinct dark dorsal markings are usually absent, but indistinct spots, blotches, or worm-like markings may be present.

Distribution and Habitat:The species is a common inhabitant of springs, creeks, rivers, swamps, sloughs, reservoirs, and ponds; it occurs throughout Tennessee.

Taxonomy: According to Stewart (1983), R. c. melanota (Rafinesque) occurs in the eastern two-thirds of Tennessee and R. c. clamitans in the Gulf Coastal Plain of western Tennessee. There appears to be a broad zone of intergradation between these two subspecies, and in some areas subspecific variation is poorly defined (Stewart, 1983; Mount, 1975; Scott and Snyder, 1968; Ferguson, 1961).


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Rana palustris Le Conte - Pickerel Frog
Rana palustris

Description: The pickerel frog is slightly smaller than R. clamitans. Head-body lengths of adults range from 4.4 to 7.5 cm. Dorsolateral ridges are well developed and extend from just behind the eye to groin. Dorsal ground color ranges from light gray to light brown, with distinct quadrangular, paired, dark markings. In a few individuals these markings are fused to form longitudinal bars. A dark spot is typically present on snout. Inner surfaces of hind legs and groin are tinged with yellow.

Distribution and Habitat: Although usually considered to occur statewide, R. palustris is uncommon in the Coastal Plain of West Tennessee. Conant and Collins (1991) show populations along the Mississippi River Valley as disjunct from those to the east along the Tennessee River in West Tennessee. Pickerel frogs are usually found in or near woodlands where they inhabit springs, creeks, ponds, reservoirs, and the twilight zone of caves.

Taxonomy: No subspecies are recognized (Schaff and Smith, 1971).


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Rana sylvatica Le Conte - Wood Frog
Rana sylvatica

Description: The wood frog is a medium-sized ranid, with an adult head-body length of 3.5 to 7.0 cm. Dorsolateral folds are present and extend from just behind eyes to groin. Dorsal coloration varies from light tan to brown. Scattered dark markings may occur on dorsum. A light stripe is present on upper lip. Lateral brown to blackish markings extend from snout to behind tympanum and form a distinct facial mask.

Distribution and Habitat: A species usually found near upland woodland streams and flooded depressions, R. sylvatica is presently known from essentially all of eastern Tennessee and across north central Tennessee westward to Dickson County.

Taxonomy: No subspecific designations are recognized (Martof, 1970).


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Rana utricularia Harlan - Southern Leopard Frog
Rana utricularia

Description: Rana utricularia is a medium-sized frog that as an adult ranges from 5.1 to 9.0 cm in head-body length. Dorsal ground color varies from light green to brown. Dorsal spotting is highly variable, but usually includes scattered, distinctly rounded, large dark spots. In some individuals, the dorsal spots may be elongate, indistinct, or absent. Prominent dorsolateral folds extend from just behind eyes to groin. Lateral surfaces of trunk usually have a few dark spots. A light line is present on upper lip. Venter is typically white, and a white spot usually occurs in center of tympanum.

Distribution and Habitat: Like Gastrophryne carolinensis, R. utricularia is common throughout most of Tennessee, but is apparently absent from the higher elevations of the Blue Ridge Mountains. Rana utricularia may also be absent from a small area of upper northeastern Tennessee. Conant and Collins (1991) provide a distribution map that shows the species absent from the Blue Ridge Mountains of Virginia, North Carolina, and eastern Tennessee. However, in the Great Smoky Mountains of Tennessee, Huheey and Stupka (1967) recorded the species from Cades Cove near 549 m elevation. Southern leopard frogs are common near farm ponds, reservoirs, creeks, rivers, sloughs, and swamps.

Taxonomy: The Rana pipiens species complex has a complicated taxonomic history. Following the most recent treatment (Pace, 1974), populations in Tennessee are designated R. u. utricularia. This species is often referred to as R. sphenocephala Cope or R. pipiens sphenocephala in the literature.


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Order Caudata - Salamanders

Family Ambystomatidae - Mole Salamanders



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Ambystoma maculatum Shaw - Spotted Salamander
Ambystoma maculatum

Description: Ambystoma maculatum is a large, burrowing species, with adults reaching total lengths of 11.2 to 19.7 cm. Dorsal coloration ranges from gray to dark brown. Several pairs of rounded, yellow to orange spots form two irregular rows from eyes to near the end of the tail. Adults typically have 12 costal grooves.

Distribution and Habitat: The spotted salamander, which occurs statewide, is most often found in hardwood or mixed pine-hardwood forests, near both permanent and temporary pools.

Taxonomy: Anderson (1967a) does not list subspecies.


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Ambystoma opacum (Gravenhurst) - Marbled Salamander
Ambystoma opacum

Description: Adult marbled salamanders are stout and medium-sized, ranging in total length from 9 to 10.7 cm. Dorsal ground color is typically dark gray to black. Light gray or white dorsal markings form crossbands that are usually complete, but sometimes broken. Costal groove counts range from 11 to 13.

Distribution and Habitat: Though distribution records are lacking for much of northeastern Tennessee, A. opacum probably occurs statewide. This assumption is based on distributional information provided from Kentucky by Barbour (1971) and from North Carolina and Virginia by Martof et al. (1980). However, Conant's and Collins' (1991) distribution map indicates the species is absent in northeastern Tennessee. The marbled salamander is known from a wide variety of habitats ranging from bottomland hardwood forests to relatively dry, upland pine forests.

Taxonomy: No subspecies are recognized (Anderson, 1967b).


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Ambystoma talpoideum (Holbrook) - Mole Salamander
Ambystoma talpoideum

Description: Ambystoma talpoideum is a medium-sized, short, stocky salamander, with relatively large head and legs. Adults reach total lengths of 7.5 to 10.0 cm. Dorsal ground color ranges from gray to black. Dorsal markings, if present, consist of light colored flecks. There are typically 11 costal grooves.

Distribution and Habitat: The mole salamander is known from swamps and flooded depressions from the Coastal Plain, northern portions of the Western Highland Rim, the Eastern Highland Rim, the Cumberland Plateau west of the Sequatchie Valley, and the extreme southeastern Blue Ridge. Distributional boundaries were provided by Redmond et al. (1982) and Conant and Collins (1991). Populations in the Blue Ridge Mountains of extreme southeastern Tennessee, and those reported by Braswell and Murdock (1979) from southwestern North Carolina, are considered part of one continuous geographic unit that is disjunct from other other portions of the species' range. Likewise, the Cumberland Plateau and Eastern Highland Rim populations in Tennessee and Alabama (Mount, 1975) are regarded as a continuous and disjunct unit. Populations in the northern Western Highland Rim appear to be continuous with those in the Coastal Plain in West Tennessee. The Tennessee Wildlife Resources Agency (1994a) lists the mole salamander as a species in need of management.

Taxonomy: No subspecific variation has been recognized for this species (Shoop, 1964).


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Ambystoma texanum (Matthes) - Smallmouth Salamander
Ambystoma texanum

Description: Compared to other Tennessee ambystomatids, A. texanum is more elongate and has a narrower head and smaller mouth. Adults attain total lengths ranging from 10.0 to 14.0 cm. Dorsal coloration ranges from dark gray to black. Distinct dorsal markings are usually absent, but when present consist of scattered, light flecks. Costal groove counts ranges from 13 to 15.

Distribution and Habitat: Ambystoma texanum occurs in the western half of the state. Its distribution outside the Coastal Plain, especially in south-central Tennessee, is uncertain and needs further study. The species is usually found in bottomland forests near swamps, ponds, and small sluggish streams. However, in Middle Tennessee it has been found along faster flowing woodland creeks and rivers.

Taxonomy: According to Anderson (1967c), there are no subspecies. Petranka (1982) provided evidence that the species as traditionally defined may include a pair of sibling species, including a pond form and a stream form. He found only the pond form in Tennessee.


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Ambystoma tigrinum (Green) - Tiger Salamander
Ambystoma tigrinum

Description: The tiger salamander is the largest ambystomatid in Tennessee, with adults ranging from 18.0 to 21.0 cm in total length. Dorsal ground color is gray to black, prominently marked with irregularly shaped yellow spots or blotches. Costal groove count is typically 12 to 13.

Distribution and Habitat: Except for north-central Tennessee, occurrence records for A. tigrinum are few. Many specimens have been taken in chance encounters during or just after heavy rains in late winter or early spring. For example, Parker (1948) reported a specimen found on a sidewalk in Memphis. Two records from Knox County were reported by residents who found specimens roaming their premises following heavy rains. The species may be absent from the Blue Ridge Mountains and a large area of northeastern Tennessee. Breeding habitats in Tennessee include flooded woodlands (Taylor, 1938), farm ponds (Gentry, 1955-1956; Ashton, 1966), shallow temporary ponds (Snyder, 1972), and limestone quarry ponds (Owen and Yeatman, 1954).

Taxonomy: Only the nominate subspecies has been reported from Tennessee (Gehlbach, 1967).


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Family Amphiumidae - Conger Eels


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Amphiuma tridactylum Culver - Three-toed Amphiuma
Amphiuma tridactylum

Description: This large, aquatic, eel-like salamander attains total lengths of 45.7 to 76.0 cm. External gills are absent. Three toes are typically present on each of four diminutive limbs. The dark gray to black dorsal coloration, separated sharply from a light gray venter, gives the species a distinct bicolored appearance.

Distribution and Habitat: The three-toed amphiuma is an inhabitant of sluggish Coastal Plain streams, oxbow lakes, and flooded ditches in the Mississippi River drainages of West Tennessee. Parker (1948) reported a sight record from the Tennessee River drainage in Benton County, but we considered that record questionable.

Taxonomy: No subspecies are recognized (Salthe, 1973).


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Family Cryptobranchidae - Hellbenders


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Cryptobranchus alleganiensis (Daudin) - Hellbender
Cryptobranchus alleganiensis

Description: The hellbender is a very large aquatic salamander that may reach a total length of 74 cm, although most adults are between 29.2 and 51.0 cm in total length. Trunk and head are dorso-ventrally compressed, and the tail is muscular, well developed, and laterally compressed. Between front and hind limbs are prominent, extensively vascularized lateral skin folds. Adults lack external gills, but have a single pair of inconspicuous gill slits on sides of neck. Eyes are small and without eyelids. Ground color ranges from olive brown to rusty orange. Irregularly shaped gray to black spots may occur on dorsum.

Distribution and Habitat: Although a few specimens have been reported from reservoirs, C. alleganiensis occurs primarily in medium-sized to large free-flowing streams in the Tennessee and Cumberland River drainages. Inhabited streams possess large rocks or logs that provide shelter and breeding sites. No Tennessee records exist from west of the Tennessee River. The Tennessee Wildlife Resources Agency (1994a) listed the species as in need of management and the U.S. Fish and Wildlife Service (1994) listed it as a Category 2 candidate for federal listing. However, the most recent federal list of candidate species (U.S. Fish and Wildlife Service, 1996) does not include the hellbender.

Taxonomy: According to Dundee (1971), only C. a. alleganiensis occurs in Tennessee.


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Family Necturidae - Mudpuppies


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Necturus maculosus (Rafinesque) - Mudpuppy
Necturus maculosus

Description: The mudpuppy is an aquatic species that vaguely resembles the hellbender. Adults range from 20.0 to 33.0 cm in total length, and have well developed external gills. Four toes are present on each of four well developed limbs. Dorsal ground color ranges from pink to brown. Scattered dark dorsal blotches are typically evident. Venter may be immaculate or possess several large dark spots.

Distribution and Habitat: This salamander probably occurs statewide in streams, reservoirs, and other permanent bodies of water. No specimens are available from the Obion River drainage, although Parker (1939) reported specimens caught by commercial fishermen in the Obion River.

Taxonomy: Relationships within the genus Necturus are poorly understood, and there is serious conflict among the various taxonomic schemes that have been proposed by several authors (Hecht, 1958; Neill, 1963a; Brode, 1969; and Mount, 1975). Most of these conflicts involve populations from areas south of Tennessee however, and there is general agreement that Tennessee populations are all assignable to N. maculosus. Conant and Collins (1991) recognize only one subspecies, N. m. maculosus, as occurring in Tennessee. However, N. m. louisianensis Viosca, which Collins (1991a, 1991b) believes may be a distinct species, may occur in the Coastal Plain drainages of West Tennessee.


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Family Plethodontidae - Lungless Salamanders


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Aneides aeneus (Cope and Packard) - Green Salamander
Aneides aeneus

Description: Adults attain total lengths of 8.3 to 12.5 cm. Toe tips are expanded to form adhesive discs. The dorsum is dark brown ,with profuse green to greenish yellow, lichen-shaped blotches.

Distribution and Habitat: Aneides aeneus is found primarily in the Cumberland Mountains, Cumberland Plateau, and Eastern Highland Rim. Presumably disjunct populations occur in the Bays Mountains, on Clinch Mountain, in scattered areas of the Appalachian Ridge and Valley, and in a cedar glade in the Inner Central Basin. Weller (1931) reported a specimen from the eastern slope of Mt. LeConte in Great Smoky Mountains National Park. King (1939) verified the identification of this specimen. Since 1931 the herpetofauna of the Great Smoky Mountains has been studied extensively by numerous scientists and Weller's report remains the only record of the species from that area. The occurrence of A. aeneus in the Great Smoky Mountains today is therefore considered doubtful. Suitable habitats include damp rock crevices of shaded sandstone outcrops and the space beneath loose bark on trees in mesic upland hardwood forests.

Taxonomy: No subspecific variation is recognized (Gordon, 1967). Type locality is mouth of Nickajack Cave, Marion County, Tennessee.


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Desmognathus aeneus Bishop and Brown - Seepage Salamander
Desmognathus aeneus

Description: Desmognathus aeneus is a small, slender salamander, with adult total lengths ranging from 4.4 to 5.7 cm. A light line extends from just behind the eye to angle of jaw. Tail is rounded, without a keel. Hind limbs are noticeably larger than forelimbs. Dorsal ground color is usually reddish brown or bronze, and irregularly shaped dark spots, sometimes forming a mid-dorsal dark stripe, are typically present. The sides of the trunk usually have dark mottling that forms wide, irregularly bounded dorsolateral stripes. These may extend from the forelimbs to the tip of tail. Dorsal surfaces of thighs usually have a light reddish or tan spot. Venter immaculate or lightly mottled, with dark melanophores.

Distribution and Habitat: In Tennessee the seepage salamander is restricted to the Blue Ridge Mountains, specifically the Unicoi Mountains, in extreme southeastern Tennessee. Jones (1982a) studied the ecology and distribution of the species in Tennessee and characterized its habitat as leaf litter near small streams and seepage areas between 280 and 1000 m elevation. The Tennessee Wildlife Resources Agency (1994a) listed the species as in need of management and the U.S. Fish and Wildlife Service (1994) listed it as a Category 2 candidate for federal listing. However, the most recent list of federal candidate species (U.S. Fish and Wildlife Service, 1996) does not include the seepage salamander.

Taxonomy: No subspecies are recognized (Harrison, 1992).


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Desmognathus fuscus (Rafinesque) - Dusky Salamander
Desmognathus fuscus

Description: Desmognathus fuscus is a medium-sized salamander that exhibits extremely variable color patterns. Adult total length ranges from 6.4 to 12.7 cm. A light line extends from just behind the eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is triangular in cross section and is moderately keeled. Jaw line of mature individuals is slightly sinuous. Toes lack dark friction pads. Jaw teeth have blunt crowns. Dorsal ground color ranges from light gray to dark brown. Dorsal color pattern is highly variable. Dorsal dark markings may be indistinct, randomly arranged, or consist of several pairs of light tan, yellowish, or reddish spots bordered by wavy (or sometimes straight) dark dorsolateral stripes. Dorsal color blends gradually with ventral color. Venter is usually mottled with dark melanophores. Older individuals may be melanistic. Desmognathus fuscus and D. santeetlah can often be distinguished by external characteristics, but for some populations a biochemical analysis is required (Tilley, 1981, 1988).

Distribution and Habitat: Excluding the Mississippi River Lowlands and Loess Plain of western Tennessee and high elevations in extreme eastern Tennessee, D. fuscus is common along small to large streams. An apparently isolated population occurs on the Mississippi River Bluffs near Ripley, Tennessee (Brandon and Huheey, 1979; Brandon, pers. comm.). Its occurrence in Great Smoky Mountains National Park and at high elevations along the Tennessee-North Carolina border has been the subject of debate and confusion. King (1939) reported D. fuscus up to 1677 m in the park. Martof and Rose (1963) noted that D. ochrophaeus was morphologically similar to D. fuscus in the Great Smoky Mountains and that D. fuscus was actually rare in the southern Appalachian Mountains. Huheey (1966) and Huheey and Stupka (1967) believed D. fuscus was absent in the park and that previous reports were based on incorrect identifications. Tilley (1981) described D. santeetlah from high elevations along the Tennessee-North Carolina border and stated that past reports of D. fuscusin the park probably referred to D. santeetlah. Tilley (1985) identified D. fuscus from Whiteoak Sinks in Great Smoky Mountains National Park. The status of D. fuscus at low elevations in the park needs further study. Tilley's studies are based primarily on electrophoretic analysis of proteins, a technique useless in the identification of preserved specimens. Because of the confusion that exists regarding the status of putative D. fuscus from the park, we did not include literature records and museum locality data from the park on our distribution map for this species. In the Blue Ridge Mountains south of the park, the occurrence of D. fuscus has been documented by Tilley (1981) and Jones (1982b). Both authors note that D. fuscus and D. santeetlah are essentially parapatric, with D. santeetlah replacing D. fuscus at high elevations along the Tennessee-North Carolina state line. The distributions of D. fuscus and D. santeetlah north of the park are virtually unknown (Tilley, 1981; Tilley, pers. comm.).


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Desmognathus fuscus (Rafinesque) - Dusky Salamander
(continued)

Taxonomy: In Tennessee, Conant and Collins (1991) depicted a relatively wide zone of intergradation between D. f. fuscus and D. f. conanti Rossman. The zone enters the state at its southeastern corner and extends northwestward into western Kentucky. Collins (1991a) speculates these subspecies may represent two separate species. Hybridization with D. santeetlah has been reported by Tilley (1981, 1988) and Jones (1982b).

















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Desmognathus imitator Dunn - Imitator Salamander
Desmognathus imitator

Description: A medium-sized species, D. imitator is morphologically very similar to D. ochrophaeus. Although morphological characteristics may be useful in distinguishing the two species, the only sure method involves an electrophoretic analysis of proteins. Adult D. imitator range from 6.4 to 10.0 cm in total length. A light line extends from eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is round in cross section and lacks a keel. Jaw line of mature individuals is strongly sinuous. Many individuals have yellow, orange, or red cheek patches. Melanistic specimens with red cheek patches mimic the red-cheeked Jordan's salamander, Plethodon jordani. In contrast to the usually straight-edged dorsolateral dark bands of D. ochrophaeus, D. imitator typically has wavy dorsolateral bands that may be broken, and that extend onto the dorsum to enclose irregularly shaped light spots. Venter is usually gray.

Distribution and Habitat: Because of the likelihood of confusing preserved specimens of D. imitator and D. ochrophaeus, the locality data presented in our maps were taken exclusively from Tilley et al. (1978) and Tilley (1985). Desmognathus imitator is restricted to Great Smoky Mountains National Park where it occurs at or above 900 m along small creeks and seepages, in moist leaf litter, and on wet rock faces (Tilley, 1985).

Taxonomy: This form was originally described by Dunn (1927b) as a subspecies of D. fuscus. Most subsequent authors considered it a color morph of D. ochrophaeus. Based on genetic studies using electrophoretic techniques, Tilley et al. (1978) provided evidence that D. imitator deserved species status. According to Tilley (1985), D. imitator is a monotypic species. Type locality is Indian Pass, Great Smoky Mountains, Sevier County, Tennessee.


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Desmognathus monticola Dunn - Seal Salamander
Desmognathus monticola

Description: Adult D. monticola range from 8.3 to 12.5 cm in total length. A light line extends from just behind the eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is triangular in cross section and is moderately keeled. Dark friction pads may be present on tips of toes. Jaw teeth have pointed crowns. Dorsal ground color ranges from light tan to dark brown. Dorsal dark markings are often distinct and form vermiculate blotches. Blotches may enclose several pairs of light tan or reddish brown light spots. Old individuals may be completely dark brown. The transition from dorsal to ventral color on the sides is abrupt. Venter may be immaculate or lightly pigmented with melanophores.

Distribution and Habitat: The range of D. monticola in Tennessee includes approximately the eastern quarter of the state. Its presence in the Blue Ridge and Cumberland mountains is well documented, but its presence in the Appalachian Ridge and Valley and Cumberland Plateau has been documented from only a few widely scattered localities. Seal salamanders occur along permanent, small to medium-sized rocky woodland streams. The species seems to prefer streams with a moderate to steep gradient. Mathews and Echternacht (1984) reported D. monticola from above 1305 m in Great Smoky Mountains National Park.

Taxonomy: No subspecies are recognized (Conant and Collins, 1991).


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Desmognathus ochrophaeus Cope - Mountain Dusky Salamander
Desmognathus ochrophaeus

Description: Desmognathus ochrophaeus is a medium-sized desmognathine, with adults ranging from 7.0 to 10.0 cm in total length. A light line extends from eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is round in cross section and keel is absent. Jaw line of mature individuals is strongly sinuous. Dorsal ground color ranges from light gray to dark brown. Some individuals are melanistic. Dorsal markings are highly variable. Dorsum may be relatively plain with only a few scattered small dark spots or flecks, or dark markings may be concentrated to form a mid-dorsal line. Dark pigment on sides forms dorsolateral bands that may have wavy or straight dorsal edges, or lateral dark pigment may extend onto dorsum to enclose several light, irregularly shaped spots. Ventral color ranges from light gray to brown, with dark melanophores usually present.

Distribution and Habitat: The mountain dusky salamander is known from the Blue Ridge Mountains, Cumberland Mountains, Cumberland Plateau, and from Bays Mountain in the Appalachian Ridge and Valley. It has recently been found on the Eastern Highland Rim (Miller, 1991). For reasons discussed in the account for D. imitator, locality data for D. ochrophaeus from Great Smoky Mountains National Park was taken exclusively from Tilley et al. (1978). At high elevations in the Blue Ridge Mountains, D. ochrophaeus inhabits mesic forests where it may be found in leaf litter or under rocks and logs. At lower elevations and elsewhere in Tennessee the species occurs along small streams, seepage areas, and on moist cliff faces.

Taxonomy: No subspecies are recognized (Martof and Rose, 1963; Tilley, 1973). Desmognathus ocoee Nicholls, a species described from Ocoee Gorge, Polk County (Nicholls, 1949), is considered a local variant of D. ochrophaeus (Martof and Rose, 1963). As described in the account for D. imitator, in the Great Smoky Mountains, D. ochrophaeus and D. imitator are often similar in morphology and color pattern, and a biochemical analysis is often necessary to separate the two.


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Desmognathus quadramaculatus (Holbrook) - Blackbelly Salamander
Desmognathus quadramaculatus

Description: This is the largest species of Desmognathus in Tennessee. Adults attain total lengths of 10.0 to 17.5 cm. A light line extends from eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is triangular in cross section and is strongly keeled. Internal nares are round and distinct. Dorsal color is usually dark brown or black, with lighter brown or rusty brown blotches. A double row of light spots normally occurs on the sides between front and hind limbs. Venter of adults is heavily pigmented, and may be completely black.

Distribution and Habitat: Desmognathus quadramaculatus is found along permanent, rocky woodland streams in the Blue Ridge Mountains and in the Bays Mountain area in the Appalachian Ridge and Valley. Inhabited streams usually have a moderate to steep gradient. The species has been reported above 1650 m in Great Smoky Mountains National Park (Mathews and Echternacht, 1984). The Tennessee Wildlife Resources Agency (1994a) lists the blackbelly salamander as in need of management.

Taxonomy: Valentine (1974) did not recognize subspecies, but he did note color pattern differences between northern and southern populations. Hinderstein (1971) noted these color differences and described biochemical differences. He found two variants; one from north and one from south of the French Broad River. He suggested these may represent two separate forms, but he did not assign taxonomic ranks.


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Desmognathus santeetlah Tilley - Santeetlah Dusky Salamander
Desmognathus santeetlah

Description: This species is closely related to and resembles D. fuscus. Adults attain total lengths of 6.4 to 9.5 cm. A light line extends from just behind the eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is triangular in cross section and is moderately keeled. Jaw line of mature individuals is slightly sinuous. Dark friction pads on tips of toes are absent. Body form is smaller and more slender, with a shorter tail than D. fuscus. Dorsal coloration is usually duller and more indistinct than in D. fuscus. Dorsal ground color may be light brown or greenish brown. Typical dorsal color patterns include (1) dark markings coalescing to enclose light spots, (2) scattered dark markings forming worm-like blotches, and (3) indistinct, small dark flecks widely scattered over dorsum. Lateral surfaces and venter usually have scattered patches of melanophores and may have a yellowish tint. A row of light spots is usually present on lower sides between front and hind limbs. Morphological and color characteristics do not always allow separation of D. fuscus and D. santeetlah (Tilley, pers. comm.). As described by Tilley (1981), the most reliable method of distinguishing the two is by electrophoretic analysis of proteins.

Distribution and Habitat: As shown by Tilley (1981, 1988) and Jones (1982b), the distribution of D. santeetlah includes high elevation seepage areas in the Unicoi and Great Smoky Mountain ranges in eastern Tennessee. As discussed for D. fuscus, many previous reports from the Great Smoky Mountains of D. fuscus probably refer to D. santeetlah.

Taxonomy: Type locality is near crest of Unicoi Mountains in Monroe County, Tennessee (Tilley, 1981). Desmognathus santeetlah hybridizes with D. fuscus (see account for D. fuscus).


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Desmognathus welteri Barbour - Black Mountain Salamander
Desmognathus welteri

Description: Desmognathus welteri is a large salamander similar in appearance to both D. fuscus and D. monticola. Adults attain total lengths of 7.5 to 12.5 cm. A light line usually extends from just behind the eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is triangular in cross section and is strongly keeled. Dark friction pads are present on tips of toes. Jaw teeth possess blunt crowns. Dorsal ground color ranges from light to dark brown. Dorsal dark markings usually consist of numerous dark flecks or small spots that seldom form a distinct pattern. Dark markings on sides may be concentrated to form wide, indistinct dorsolateral stripes. Dorsal ground color blends gradually with ventral color. Venter is usually mottled with dark melanophores. Old individuals may be melanistic.

Distribution and Habitat: Redmond (1980) determined the distribution of D. welteri to include the Cumberland Mountains and northern half of the Cumberland Plateau. The species typically occurs along small to medium-sized permanent streams in mesic upland hardwood forests. Strongly aquatic, its apparent absence on the southern Cumberland Plateau may be due to the seasonal nature of most small streams in that region. The Tennessee Wildlife Resources Agency (1994a) lists the species as in need of management.

Taxonomy: The species, originally described as a subspecies of D. fuscus (Barbour, 1950), was later elevated to species rank (Barbour 1971). Subsequent studies by Caldwell (1977, 1980), Caldwell and Trauth (1979), and Juterbock (1975, 1978, 1984) support Barbour's proposal. No subspecies have been described (Conant and Collins, 1991).


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Desmognathus wrighti King - Pygmy Salamander
Desmognathus wrighti

Description: The pygmy salamander is a small salamander, similar in body size and form to D. aeneus. Adults may reach total lengths of 3.8 to 5.1 cm. A light line extends from just behind the eye to angle of jaw. Tail is rounded in cross section and lacks a keel. Hind limbs are noticeably larger than forelimbs. Dorsal ground color ranges from light gray to rusty brown. Dorsal markings typically consist of narrow dark lines forming a herringbone pattern. Dark markings and scattered silver flecks on sides combine to form dorsolateral bands. Dorsal surface of head and snout is rugose. Venter is usually immaculate.

Distribution and Habitat: Desmognathus wrighti is restricted to high elevations in the Blue Ridge Mountains along the Tennessee-North Carolina border. Most authorities (Huheey, 1966; Huheey and Stupka, 1967; Tilley and Harrison, 1969; Mathews and Echternacht, 1984) have regarded this species as characteristic of spruce-fir forests, although populations are known from as low as 838 m (Huheey, 1966). Tilley and Harrison (1969) believe these lower elevation populations in hardwood forest habitats represent relicts from the past when spruce-fir habitats were more widespread in the southern Appalachians. The pygmy salamander is the most terrestrial of all desmognathine species, and may occur at great distances from streams and seepages. Adults are found under and within rotting logs, beneath rocks, and just beneath leaf litter. The Tennessee Wildlife Resources Agency (1994a) considers the species as in need of management.

Taxonomy: No subspecies have been reported (Conant and Collins, 1991). King (1936) described D. wrighti from Mt. LeConte, Sevier County, Tennessee.


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Eurycea cirrigera (Green) - Southern Two-lined Salamander
Eurycea wilderae Dunn - Blue Ridge Two-lined Salamander
Eurycea cirrigera and Eurycea wilderae

Description: Both species of two-lined salamanders are slender. Adults reach total lengths of 6.4 to 10.7 cm. Ground color ranges from yellow through orange to, occasionally, light brown. A dark lateral stripe occurs on each side of the body and extends from eye to middle of tail or beyond, sometimes reaching tail tip. Small black or brown spots may occur on dorsum between lateral dark stripes.

Distribution and Habitat: Eurycea cirrigera and E. wilderae are usually difficult to distinguish using external characteristics (Jacobs, 1987). In Tennessee, Jacobs indicated the two species have allopatric distributions, with E. wilderae occurring in the Blue Ridge Mountains and E. cirrigera over the remainder of the state. Since specimens on which literature and museum records are based were not analyzed by us to determine specific status, the distribution map above should be regarded as representing a composite range for both species. The dashed line on the map delineates the eastern edge of the Blue Ridge Mountains and, presumably, the approximate boundary between E. wilderae and E. cirrigera. Two-lined salamanders are common in moist-to-wet habitats along woodland streams throughout Tennessee. Populations are known from bottomland habitats in West Tennessee eastward to the forests and balds along the crest of the Blue Ridge Mountains.

Taxonomy: Revising Mittleman (1966), Jacobs (1987) studied biochemical variation in the Eurycea bislineata group. He recommended elevating each subspecies, E. b. bislineata (Green), E. b. cirrigera (Green), and E. b. wilderae Dunn, to species status. According to Jacob's (1987) range map, only two species, E. cirrigera and E. wilderae, occur in Tennessee, where they are mutually allopatric. For the purposes of this Atlas, E. aquatica Rose and Bush is included in synonymy with E. cirrigera. Rose and Bush (1963) indicated E. aquatica might occur in Tennessee, and Ashton (1966) reported the species from Davidson County, Tennessee. In Alabama, Mount (1975) concluded E. aquatica was merely an ecotype of E. bislineata. Wallace (1975) reached the same conclusion for populations in Davidson County, Tennessee. Jacobs (1987) also questioned the taxonomic validity of E. aquatica.


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Eurycea junaluska Sever, Dundee, and Sullivan - Junaluska Salamander
Eurycea cirrigera and Eurycea wilderae

Description:This recently described species is morphologically very similar to E. wilderae and E. cirrigera. Adults attain total lengths of 7.5 to 10.0 cm. In comparison with E. wilderae and E. cirrigera, E. junaluska has a relatively shorter tail and longer limbs. Dorsal coloration is usually a light yellow, with dorsolateral brown stripes absent or broken into narrow wavy lines. Scattered small dark spots or flecks may occur on dorsum.

Distribution and Habitat: Eurycea junaluska is known from medium to large streams in a small area of eastern Tennessee. Sever (1976) reported an individual from Fighting Creek in Sevier County; later he found the species along the Tellico River in Monroe County (Sever, 1983b). He collected individuals from beneath rocks along streams and from roads near streams during or just after rain. During the night of 12 September 1976, one of us (W. H. R.) found several individuals perched on boulders in the Little Tennessee River along the Blount-Monroe county line. This area has since been inundated to form Tellico Reservoir. The Tennessee Wildlife Resources Agency (1994a) lists this species as being in need of management.

Taxonomy: No subspecies have been reported (Sever, 1983a). Jacobs (1987) questioned the taxonomic status of E. junaluska.


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Eurycea longicauda (Green) - Longtail Salamander
Eurycea longicauda

Description: Eurycea longicauda has a slender body and long tail. Adults range from 10.0 to 15.9 cm in total length. Dorsal ground color varies from light yellow to yellowish brown. In Tennessee, two distinct dorsal color patterns occur, representing two subspecies. One subspecies has a single mid-dorsal and two dorsolateral dark stripes. The mid-dorsal stripe originates near the eyes and extends to base of tail, whereas the dorsolateral stripes begin just behind the eyes and may extend to tip of tail. This form, commonly called the three-lined salamander, also possesses dark spots or a mottled pattern on the venter. The other form typically has numerous irregularly shaped dark spots on its back and sides, the markings on the sides sometimes forming indistinct dorsolateral stripes. Venter is usually immaculate. Sides of tail have vertical dark markings that form a distinctive herringbone pattern.

Distribution and Habitat: Eurycea longicauda occurs statewide, but may be absent from higher elevations in the Blue Ridge Mountains. Suitable habitats include streams banks and mesic slopes in woodlands, and the twilight zone of caves.

Taxonomy: Two subspecies are found in Tennessee (Ireland, 1979). Eurycea l. guttolineata (Holbrook), the three-lined salamander, occurs in the Coastal Plain of West Tennessee and has been found in a few scattered localities in the mountains of East Tennessee. Eurycea l. longicauda ranges from the Tennessee River in West Tennessee eastward throughout the state. Parker (1937, 1939) reported Eurycea l. longicauda from the Coastal Plain of West Tennessee in the hills east of Reelfoot Lake. Ireland (1979) comments that along the Blue Ridge escarpment these two subspecies appear to be reproductively isolated. Examination of over 650 specimens during this study revealed no evidence of interbreeding between these two forms in extreme East Tennessee. An examination of specimens from populations along both sides of the Tennessee River in West Tennessee (i.e., Stewart, Henry, Perry, Henderson, Hardin, and Lawrence counties) revealed color patterns suggesting some interbreeding. These intermediate specimens typically had a distinct mid-dorsal dark stripe, a characteristic of E. l. guttolineata, but had reduced amounts of dark pigmentation on the venter, a characteristic of E. l. longicauda. A few of these specimens had a mid-dorsal stripe that was broken and indistinct. Further studies are needed to quantify the extent of intergradation between these two subspecies in Tennessee and elsewhere (Collins, 1991a).


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Eurycea lucifuga Rafinesque - Cave Salamander
Eurycea lucifuga

Description: Like E. longicauda, the cave salamander is slender and has a long tail. Total length in adults ranges from 10.0 to 15.2 cm. Dorsal ground color may be yellowish orange, orange, or reddish orange. Markings include numerous, irregular dark spots over the entire dorsal surface, including the tail. No herringbone pattern is present on sides of tail.

Distribution and Habitat: The range of E. lucifuga in Tennessee extends from the Tennessee River in the west to the western edge of the Blue Ridge Mountains in the east, where the species is known from only two cave localities. Eurycea lucifuga occurs in the twilight zone of caves and near cave entrances. It also inhabits mesic woodlands, especially near bluffs and limestone outcrops.

Taxonomy: Although no subspecies are recognized (Hutchison 1966), Grobman (1943) and Sinclair (1965) reported unusually large, dark, dusky specimens from several localities within and near the Nashville Basin of central Tennessee. Neither author proposed taxonomic recognition for these aberrant individuals. Merkle and Guttman (1977) studied genetic variation using electrophoretic techniques and found allelic differences between populations in the Nashville Basin and elsewhere, but did not recommend naming subspecies.


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Gyrinophilus palleucus McCrady - Tennessee Cave Salamander
Gyrinophilus palleucus

Description: The Tennessee cave salamander is a pale troglobite. Adults range from 7.5 to 18.4 cm in total length. External gills normally persist throughout life. Eyes are small and poorly developed. Snout is flat and head broad. Dorsal ground color varies from pale pink to brown. Dark dorsal spots may occur, and a dark stripe may be present on throat. Occasional individuals undergo metamorphosis, losing their gills. In Tennessee, naturally metamorphosed individuals have been reported from Knox County (Simmons, 1976), Franklin County (Yeatman and Miller, 1985; Brandon et al., 1986), and Warren County (Miller, 1995).

Distribution and Habitat: In Tennessee, this troglobite is known from subterranean waters of the Tennessee River drainage in Knox, Roane, McMinn, Hamilton, Marion, Grundy, and Franklin counties, and from the Cumberland River drainage in Rutherford, Warren, and Wilson counties. The Wilson (Miller and Walther, 1994) and Warren (Miller, 1995) county records are recent discoveries. Little is known about the habitat requirements of this species. Caldwell and Copeland (1992) proposed that inflow (sinkhole) caves, as opposed to outflow caves, provide the best habitat. The Tennessee Wildlife Resources Agency (1994b) listed the species as threatened, and the U.S. Fish and Wildlife Service (1994) listed it as a Category 2 candidate for federal listing. However, the most recent federal list of candidate species (U.S. Fish and Wildlife Service, 1996) does not include the Tennessee cave salamander.

Taxonomy: Three subspecies have been reported for Tennessee. These include G. p. palleucus, G. p. necturoides Lazell and Brandon, and G. p. gulolineatus Brandon (Brandon, 1967a; Conant and Collins, 1991). Type locality for the species is Sinking Cove Cave in Franklin County, Tennessee. Using electrophoretic techniques, Addison Wynn and Jeremy Jacob (pers. comm.) studied biochemical variation and have found evidence of a new species in Tennessee and possible hybridization of G. palleucus and G. porphyriticus. Collins (1991b) proposed that G. p. gulolineatus be recognized as a separate species.


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Gyrinophilus porphyriticus (Green) - Spring Salamander
Gyrinophilus porphyriticus

Description: The spring salamander is a large species, with total lengths of adults ranging from 12.1 to 19.0 cm. The canthus rostralis, a light line from each eye to nostril, may be indistinct or distinctly bordered with black pigment. Ground color is usually yellowish pink, red, reddish brown, or tan. Dorsal dark markings are extremely variable, ranging from virtually absent (a few small black spots or flecks) to heavily mottled with dark reticulations, sometimes forming chevron-shaped markings. Venter may be plain or have numerous melanophores.

Distribution and Habitat: The spring salamander occurs east of the Outer Central Basin in the eastern third of the state. It occurs along shaded, small to medium-sized streams. In areas of karst topography where permanent surface aquatic habitats are scarce, the species is known to occur in cave streams and pools.

Taxonomy: Four subspecies, with extensive zones of intergradation, are recognized. According to Brandon (1962, 1967b), populations in Tennessee from the Eastern Highland Rim to the western edge of the Blue Ridge Mountains are intergradient between G. p. porphyriticus and G. p. duryi Mittleman and Jopson. Populations in the Blue Ridge Mountains are designated as G. p. danielsi (Blatchley). Sinclair (1953, 1955) proposed the recognition of G. warnerias a new species from Middle Tennessee. His comments, available in abstracts of papers presented at meetings of the Tennessee Academy of Science, were never published as a formal description. Brandon (1962) studied specimens made available by Sinclair and concluded that they were not members of the genus Gyrinophilus, but were probably Pseudotriton montanus.


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Hemidactylium scutatum (Temminck and Schlegel) - Four-toed Salamander
Hemidactylium scutatum

Description: The four-toed salamander is a small species, adults ranging from 5.1 to 9.0 cm in total length. Four toes are present on each hind foot. There is a distinct constriction at the base of the tail. Dorsal coloration varies from gray to rusty brown, with indistinct small dark markings. Lateral surfaces are often heavily mottled with black or dark brown markings. Venter is bright white, with numerous distinct scattered black spots.

Distribution and Habitat: The distribution of H. scutatum in Tennessee is poorly known. The range probably extends from the Western Highland Rim eastward to North Carolina. Habitats include woodland swamps, shallow ponds, and sphagnum bogs. The Tennessee Wildlife Resources Agency (1994a) lists the species as in need of management.

Taxonomy: As reported by Neill (1963b), no subspecies have been designated. The type locality is listed as Nashville, Davidson County, Tennessee.


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Leurognathus marmoratus Moore - Shovelnose Salamander
Leurognathus marmoratus

Description: Leurognathus marmoratus, a permanently aquatic species, is often confused with D. quadramaculatus. Adults vary in total length from 9.0 to 12.5 cm. Although often obscure, a light line extends from eye to angle of jaw. Hind limbs are noticeably larger than forelimbs. Tail is laterally compressed and strongly keeled. Snout appears flatter than in D. quadramaculatus. Internal nares are slit-like and obscure. Dorsum is typically dark brown or black, with two rows of irregularly shaped light blotches. Venter is usually dark gray, and may possess a lighter center.

Distribution and Habitat: The shovelnose salamander is found in drainages of the Blue Ridge Mountains north of the Little Tennessee River. The species is typically found in small to medium-sized rocky woodland streams, with steep to moderate gradient. Mathews and Echternacht (1984) recorded L. marmoratus above 1650 m elevation, and Huheey and Stupka (1967) noted its apparent absence below 457 m.

Taxonomy: No subspecies are recognized (Martof, 1963).


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Plethodon aureolus Highton - Tellico Salamander
Plethodon aureolus

Description: The recently described Tellico salamander, a large species very similar in its morphology to P. glutinosus and P. teyahalee (Highton, 1983), can be reliably distinguished from those two species only on the basis of biochemical characters. Adult P. aureolus, ranging from 10.0 to 15.1 cm in total length, are, on average, smaller than both P. glutinosus and P. teyahalee adults (Highton, 1983). Dorsal and lateral ground color is dark gray to black, with many large, brassy spots. Venter is dark gray to black, and chin is typically lighter in color than venter.

Distribution and Habitat: All locality data plotted were taken from Highton (1983), who defined the species' range to include the western slopes of the Unicoi Mountains and adjacent lowlands between the Little Tennessee and Hiwassee rivers. Typical habitat probably includes both upland and stream valley woodlands.

Taxonomy: No subspecies are recognized (Highton, 1983, 1986a). According to Highton (1983), P. aureolus is sympatric with typical P. glutinosus on the western edge of the Unicoi Mountains and sympatric throughout its range with P. teyahalee (see account for P. teyahalee). Evidence exists of hybridization between P. aureolus and P. jordani (Highton,1983). Type locality is Farr Gap, Unicoi Mountains, Monroe County, Tennessee.


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Plethodon cinereus (Green) - Redback Salamander
Plethodon cinereus

Description: Plethodon cinereus is small, adults ranging in total length from 5.7 to 10.0 cm. It is similar to P. dorsalis and morphologically virtually indistinguishable from P. serratus. Dorsal coloration usually consists of a broad, straight-edged light red stripe extending from neck well onto tail, where it becomes conspicuously narrower. In individuals without the dorsal stripe, the dorsum is dark brown or black, with scattered light flecks and no red pigment. Venter is mottled with equal amounts of black and white, but typically lacks red markings. Costal groove counts range from 18 to 20.

Distribution and Habitat: As determined by Highton and Webster (1976), the range of P. cinereus in Tennessee includes the Blue Ridge Mountains north of the French Broad River. The redback salamander is found under logs, rocks, and leaf litter in upland forests. A Tennessee record of P. cinereus from outside the Blue Ridge Mountains was determined invalid by Grobman (1944), who provided substantial evidence that a specimen in the U.S. National Museum (USNM No. 57106), listed from Franklin County, Tennessee, was actually taken in Franklin County, Missouri.

Taxonomy: No subspecies are recognized (Highton and Webster, 1976). Prior to Highton's and Webster's study, all populations of redback salamanders in the Blue Ridge Mountains of Tennessee were considered part of the same species, P. cinereus (Smith, 1963). In their study, they used biochemical differences to recognize part of the P. cinereus complex (including populations in the southern half of Tennessee's Blue Ridge Mountains) as a new species, P. serratus.


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Plethodon dorsalis Cope - Zigzag Salamander
Plethodon dorsalis

Description: Total lengths of adults of this salamander range from 6.4 to 9.0 cm. Many individuals have a dorsal light red, brown, or yellowish stripe, with lobed or wavy margins, extending from neck well onto the tail. Stripe may appear to widen at base of tail. Other individuals have a uniformly dark brown or black dorsum. In the Central Basin of Middle Tennessee, a red shoulder patch is present in most individuals (Brian Miller, per. com.). Ventral surfaces are light, with profuse black or black and reddish mottling. Costal groove count is usually 18.

Distribution and Habitat: The species appears to be absent from elevations above 762 m in the Blue Ridge Mountains (King, 1939) and from most of the Coastal Plain of West Tennessee, where it is known from only two localities. Parker (1939) described one of the West Tennessee sites (in Obion County) as wooded hills east of Walnut Log and Reelfoot Lake, where he found specimens in leaf mats and near springs. Thurow (1966) characterized the same area as bluffs composed of consolidated loess that provided rock shelter habitats. Habitat data for the other Coastal Plain record (a Henry County locality) are lacking. Two specimens were taken from the Obion River area, Highway 69, north of Jones Mill (NLU Nos. 45756-45757). Elsewhere in Tennessee the species is usually found beneath leaf litter, rocks, and logs in mesic upland forests. The status of P. dorsalis in the Cumberland Mountains is poorly known and needs further study.

Taxonomy: According to Thurow (1966) and Conant and Collins (1991), only the nominate subspecies occurs in Tennessee.


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Plethodon glutinosus (Green) - Northern Slimy Salamander,
Plethodon cylindraceus (Harlan) - White-spotted Slimy Salamander,
Plethodon mississippi Highton - Mississippi Slimy Salamander
Plethodon glutinosus, Plethodon cylindraceus, and 

Plethodon mississippi

Description: Like P. aureolus, P. kentucki, and P. teyahalee (treated elsewhere in separate accounts), these three species are also considered members of the P. glutinosus complex as defined by Highton et al. (1989). The three species treated in this account are large plethodontids, with adult total lengths ranging from 12.1 to 17.2 cm. The following descriptions are from Highton et al. (1989). Of the three, P. mississippi tends to be the smallest. Dorsal and lateral ground color is usually brown or black. Plethodon glutinosus and P. mississippi normally possesses large brassy colored dorsal spots, while P. cylindraceus has large white spots on the dorsum. Plethodon cylindraceus has abundant lateral white spots while P. glutinosus and P. mississippi may have both white and yellow lateral spotting. Plethodon cylindraceus is light chinned. Conant and Collins (1991) state that geography is an important criterion in identification of individuals of this species complex.

Distribution and Habitat: As depicted by Highton et al. (1989) and Conant and Collins (1991), the distributions of P. glutinosus, P. cylindraceus, and P. mississippi appear to be allopatric or with very narrow zones of sympatry. The literature and museum data presented on the map above are not based on biochemical analysis, and therefore could not be accurately assigned to any one species. The map should be viewed as a composite range map for all three species. The dark boundary lines approximate the boundaries of species' ranges, with P. mississippi in the Coastal Plain of West Tennessee, P. glutinosusin most of the eastern two-thirds of the state, and P. cylindraceus in the Blue Ridge Mountains and Ridge and Valley in northeastern Tennessee (Highton et al., 1989; Conant and Collins, 1991). Slimy salamanders exploit a wide variety of woodland habitats ranging from mesic bottomland hardwood to relatively dry hillside forests.

Taxonomy: No subspecific designations are recognized for any of these species. All three species, having been defined by biochemical characteristics, are similar in appearance and difficult to distinguish using traditional external taxonomic characteristics (Highton et al., 1989; Conant and Collins, 1991).


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Plethodon jordani Blatchley - Jordan's Salamander
Plethodon jordani

Description: Jordan's salamander is a large plethodontid. Adult total lengths range from 9.0 to 12.5 cm. In most adults dorsal ground color is dark gray or black, with no white or brassy markings. However, individuals from the Unicoi Mountains in southeastern Tennessee typically have lateral white spots and flecks. Populations from the Great Smoky Mountains usually possess red cheek patches, whereas other Tennessee populations have cheeks essentially the same color as the dorsum. Venter is usually lighter than dorsum, and chin is usually lighter than rest of venter.

Distribution and Habitat: The Tennessee range of P. jordani includes high elevation habitats in the Blue Ridge Mountains along the Tennessee-North Carolina border. It occurs in moist woodlands from mountain summits down to 762 m elevation (Huheey and Stupka, 1967). Highton (1983) noted that what appeared to be P. jordani from several localities north of Jones Knob in the Unicoi Mountains in Monroe County was actually P. aureolus, and that Johns Knob was the northernmost locality for P. jordani in the Unicoi Mountains.

Taxonomy: Two races were formerly recognized from Tennessee. These were the uniformly black metcalfi race, and the red-cheeked jordani race (Conant, 1958). Highton (1962, 1973) studied variation in these forms and concluded that subspecific recognition was unwarranted. Highton and Henry (1970) reported some evidence of hybridization between P. jordani and P. glutinosus in the Great Smoky Mountains, but found substantial indications of hybridization with P. glutinosus in the Unicoi Mountains. Highton (1983) later described P. aureolus and P. teyahalee, two biochemically defined cryptic species of P. glutinosus, from the Unicoi Mountains of Tennessee and adjacent western North Carolina, and presented evidence of hybridization of both with P. jordani. Highton (1971) found no indication of hybridization of P. glutinosus and P. jordani east of the French Broad River. The type locality of P. jordani is near the divide along the Tennessee-North Carolina border in Great Smoky Mountains National Park.


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Plethodon kentucki Mittleman - Cumberland Plateau Woodland Salamander
Plethodon kentucki

Description: Plethodon kentucki is a large plethodontid that is very similar to P. glutinosus. Though in areas of sympatry biochemical characteristics may be the only reliable basis for separating the two (Highton, 1986b), the following contrasting features may help distinguish the species. Adult P. kentucki, with total lengths ranging from 9.8 to 16.8 cm, are typically smaller than P. glutinosus. Dorsal and lateral color is black, with scattered white spots that are smaller and less numerous than those of P. glutinosus. Light spots on P. kentucki are not as brassy as those on P. glutinosus. Adult male P. kentucki have larger mental glands than adult males of P. glutinosus. Finally, the chin of P. kentucki is noticeably lighter than the very dark venter (a condition not usually true for P. glutinosus).

Distribution and Habitat: John MacGregor (pers. comm.) collected the only specimen known from Tennessee, and described its habitat as a shale outcrop bordering a gravel road. MacGregor suggests the species probably occurs elsewhere in the Cumberland Mountains in Tennessee. In Kentucky P. kentucki and P. glutinosus are often sympatric (MacGregor, pers. comm.). Optimum habitat for P. kentucki is mature hardwood forests on steep slopes underlain by sandstone or shale (MacGregor, pers. comm.).

Taxonomy: Plethodon kentucki was originally described by Mittleman (1951) from eastern Kentucky. Clay et al. (1955) reduced P. kentucki to synonymy with P. glutinosus. After an analysis of both morphological and biochemical characteristics, Highton and MacGregor (1983) restored P. kentucki to species rank. No subspecies are recognized (Highton, 1986b).


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Plethodon richmondi Netting and Mittleman - Ravine Salamander
Plethodon richmondi

Description: The ravine salamander is a small, worm-like plethodontid, with relatively short limbs. Adults attain total lengths of 7.5 to 11.5 cm. Dorsal and lateral surfaces are dark brown or black, with scattered silver, white, or brassy flecks. Plethodon richmondi differs from other small plethodontids in possessing a predominately dark brown or black venter. Costal groove counts range from 19 to 22.

Distribution and Habitat: Plethodon richmondi is known from northern portions of the Blue Ridge Mountains, Appalachian Ridge and Valley, and Cumberland Mountains. Typical habitat is mesic upland forests where individuals are usually found under rocks, logs, and leaf litter.

Taxonomy: No subspecies are recognized (Conant and Collins, 1991). Thurow (1969) reported evidence of hybridization with P. cinereus on Iron and Houston mountains in northeastern Tennessee.


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Plethodon serratus Grobman - Southern Redback Salamander
Plethodon serratus

Description: Plethodon serratus is a small plethodontid nearly identical morphologically to P. cinereus. Total length of adults ranges from 8.1 to 10.5 cm. Dorsal color pattern includes a straight-edged light red stripe that extends from neck well onto the tail, where it becomes conspicuously narrower. Rare individuals lack the dorsal stripe and are dark brown or black above, with scattered light flecks. Venter is mottled with equal amounts of black and white and usually has some red pigment. Costal groove counts range from 18 to 20.

Distribution and Habitat: Highton and Webster (1976) indicated that in Tennessee P. serratus occurs in the Blue Ridge Mountains south of the French Broad River. Like P. cinereus, P. serratus is found in terrestrial habits in upland forests.

Taxonomy: No subspecies are recognized (Highton and Webster, 1976; Highton, 1986c). Highton and Webster (1976) elevated this form to species status based on biochemical characteristics (see account for P. cinereus).


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Plethodon teyahalee Hairston - Southern Appalachian Slimy Salamander
Plethodon teyahalee

Description: The southern Appalachian slimy salamander is a large plethodontid that attains total lengths of 12.1 to 17.2 cm. Based on external characteristics, it is sometimes difficult to distinguish from its sympatric congeners, P. aureolus and P. glutinosus (Highton et al., 1989). The usually black dorsal ground color is punctuated with scattered small white spots. White spots on lateral surfaces are typically larger than those on dorsum. Small red spots may occur on legs. Chin is lighter than the normally gray venter. Conant and Collins (1991) note that geography is a critical criterion in field identification of individuals of the P. glutinosus complex.

Distribution and Habitat: Because of the difficulty of distinguishing specimens of P. Teyahalee, P. glutinosus, and P. aureolus, locality data shown on the above map are exclusively from Highton (1983, 1987a). In Tennessee, P. teyahalee is known from mesic woodland habitats in the Unicoi Mountains in Monroe and Polk counties, and from the Great Smoky Mountains in Sevier and Cocke counties.

Taxonomy: Highton (1973) considered P. Jordani teyahalee Hairston synonymous with P. glutinosus. Based on biochemical data, he subsequently determined that the teyahalee morph represented a separate species(Highton, 1983) provided numerous localities for P. teyahalee in the Unicoi Mountains in Monroe and Polk counties, and later(Highton, 1987a) provided a detailed account and range map for the species. Hairston (1993) pointed out that the name P. teyahalee was invalid due to the hybrid nature of the population where the type specimen was taken. He proposed a new name P. oconaluftee and designated a new type. Hybridization with P. jordani was reported by Highton (1983).


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Plethodon wehrlei Fowler and Dunn - Wehrle's Salamander
Plethodon wehrlei

Description: Only two specimens (UTKVZC 06250 and 06985), one adult and one juvenile, are currently available from Tennessee (Redmond and Jones, 1985). The adult has a total length of 9.1 cm and a snout-vent length of 4.8 cm. The juvenile is 5.7 cm long overall, with a snout-vent length of 3.1 cm. Distinct webbing is present between all toes of both. When alive, the venter was gray and the dorsum was dark brown, with 8 to 10 irregularly shaped yellow spots.

Distribution and Habitat: The two Tennessee specimens of Wehrle's salamander were found at the same locality, a gorge in a mesic hardwood forest in the Cumberland Mountains. The adult was found in a crevice in a rock shelter on a shaded sandstone cliff face, and the juvenile from a path approximately 20 m from where the adult was taken. Both specimens were collected on warm, misty nights. The Tennessee Wildlife Resources Agency (1994a) lists this species as in need of management.

Taxonomy: According to Highton (1987b), no subspecies are recognized. Throughout the range of P. wehrlei, the yellow-spotted morph is rare and has been reported from only three localities (Cupp and Towles, 1983; Redmond and Jones, 1985). Richard Highton (pers. comm.) does not believe these populations deserve formal taxonomic recognition.


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Plethodon welleri Walker - Weller's Salamander
Plethodon welleri

Description: Weller's salamander is a small plethodontid, with adults ranging from 6.4 to 7.9 cm in total length. The dorsum is black, washed with gold or brassy, irregular blotches. Venter is usually black, with numerous small white flecks or spots.

Distribution and Habitat: Plethodon welleri is restricted to the Blue Ridge Mountains in northeastern Tennessee. Populations are usually found above 762 m elevation on forested mountain summits, mesic woodland talus slopes, and in cove hardwood forests. Thurow (1963) reported a population at 700 to 732 m in a limestone cove forest dominated by hemlock and yellow birch. The Tennessee Wildlife Resources Agency (1994a) lists Weller's salamander as in need of management.

Taxonomy: Thurow (1964) lists one subspecies, P. w. ventromaculatus Thurow, from Tennessee. Conant and Collins (1991) do not recognize subspecies.


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Plethodon yonahlossee Dunn - Yonahlossee Salamander
Plethodon yonahlossee

Description: Plethodon yonahlossee is a large plethodontid whose adult total length ranges from 11.5 to 18.0 cm. A wide, irregularly bordered dorsal red stripe extends from near the head onto the tail. This stripe may be partially or completely interrupted by black spots or blotches. Lateral surfaces are heavily marked with white or light gray. Venter is dark gray, usually with numerous scattered light spots. Throat is lighter.

Distribution and Habitat: In Tennessee, the range of P. yonahlossee is strikingly coincident with that of P. welleri. Both are found only in the Blue Ridge Mountains along the state's northeastern border. Yonahlossee salamanders occur in mature woodlands at elevations ranging from 732 to 1433 m.

Taxonomy: No subspecies are recognized (Pope, 1965; Conant and Collins, 1991).


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Pseudotriton montanus Baird - Mud Salamander
Pseudotriton montanus

Description: The mud salamander is large, with a more slender body than its Tennessee congener, the red salamander (P. ruber). Adults attain total lengths of 7.5 to 16.5 cm. Dorsal and ventral ground color may be coral-pink, red, or reddish brown. A few well defined, rounded black spots are usually present on dorsum. Venter is usually immaculate.

Distribution and Habitat: Excluding the high elevations of the Blue Ridge Mountains, the range of P. montanus includes the eastern half of Tennessee. Highest reported locality in Great Smoky Mountains National Park is 477 m, and several localities in the Cumberland Mountains and Cumberland Plateau are above 550 m. The mud salamander inhabits mud and silt-laden areas of floodplain woodland streams, swamps, and seepage areas. Miller (1990) reported the species from an intermittent stream in a cedar glade in Wilson County.

Taxonomy: Martof (1975a) followed the recommendation of Bruce (1968) and did not recognize subspecies. Conant and Collins (1991) disagreed and recognized four subspecies, one of which, P. m. diastictus Bishop, they show occurring in Tennessee. Collins (1991a, 1991b) suggested that P. m. diastictus is a distinct species.


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Pseudotriton ruber (Sonnini) - Red Salamander
Pseudotriton ruber

Description: The red salamander is a large, stout-bodied species that reaches adult total lengths of 7.0 to 15.2 cm. It is similar to P. montanus, but has a stockier body and smaller head. Dorsal and ventral ground colors range from bright red to a dull purplish brown. Dorsal markings typically consist of many small, irregularly shaped dark spots that may be fused in older individuals. Ventral surface of chin may be lightly flecked or heavily pigmented with black. Venter may be immaculate or spotted with dark markings.

Distribution and Habitat: Pseudotriton ruber is found throughout Tennessee east of the Loess Plain of West Tennessee. Available data indicate the species may be rare in the Inner and Outer Central basins. It has been reported from above 1524 m in Great Smoky Mountains National Park (Huheey and Stupka, 1967). Red salamanders occur near many woodland aquatic habitats including creeks, springs and spring runs, and seepage areas. It may occasionally be found in mesic to relatively dry woodlands.

Taxonomy: Disagreement exists regarding the existence of valid subspecies. Martof (1975b) cites Bruce (1968) and did not recognize subspecific designations, but Conant and Collins (1991) list four subspecies, all of which occur in Tennessee. Pseudotriton r. vioscai Bishop occurs in the eastern two-thirds of the Coastal Plain in West Tennessee, P. r. ruber from the Tennessee River in West Tennessee to the western edge of the Blue Ridge Mountains, P. r. nitidus Dunn in the northern half of the Blue Ridge Mountains, and P. r. schencki (Brimley) in the southern half of the Blue Ridge Mountains.


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Family Salamandridae - Newts


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Notophthalmus viridescens (Rafinesque) - Eastern Newt
Notophthalmus viridescens

Description: Notophthalmus viridescens has a distinct terrestrial larval form and an aquatic adult form. Neither form typically has external gills. The terrestrial stage is commonly called an eft and is bright red or orange, with dorsal red or black spots. Total lengths of efts range from 3.5 to 8.6 cm. Skin of eft is very spinose. Adults are aquatic and attain total lengths of 5.7 to 12.2 cm. Adult dorsal coloration ranges from yellowish green to brown, with either numerous red spots bordered by black or with only small black spots.

Distribution and Habitat: The eastern newt occurs throughout the state. Adults inhabit ponds, pools along and within streams, oxbows, and flooded ditches. Efts are most often encountered under rocks and logs in upland forests.

Taxonomy: Two subspecies occur in Tennessee. Notophthalmus v. viridescens occurs in the eastern two-thirds of the state and N. v. louisianensis (Walterstorff) occurs in the Coastal Plain in the western one-third (Mecham, 1967). Page and Scott (1990) reported evidence of intergradation between the two subspecies in Land Between The Lakes, Stewart County.


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Family Sirenidae - Sirens


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Siren intermedia Le Conte - Lesser Siren
Siren intermedia

Description: The lesser siren is a permanently aquatic, eel-like salamander that possesses well-developed external gills. Adults reach total lengths of 18.0 to 68.6 cm. Tiny front limbs are present, hind limbs absent. Dorsal coloration varies from gray to brown to black and may include small diffuse light spots. Lateral body surfaces and venter may have light flecks.

Distribution and Habitat: The lesser siren is known from sluggish streams, oxbows, and flooded ditches throughout most of the Coastal Plain of West Tennessee and from swamps and marshes in the Cumberland River floodplain in Stewart, Montgomery, and Davidson counties.

Taxonomy: Only one subspecies, S. i. nettingi Goin is found in Tennessee (Martof, 1973).


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