Amphibians of North Carolina
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NC Records

Eurycea longicauda - Long-tailed Salamander

Taxonomy
Class: Amphibia Order: Caudata Family: Plethodontidae Subfamily: Spelerpinae Other Common Name(s): Longtail Salamander
Taxonomic Comments: Eurycea longicauda and E. guttolineata were at one time treated as members of a single polytypic species with three subspecies. These include the Dark-sided Salamander (E. l. melanopleura) that is associated with the Ozark Highlands of Missouri, the Long-tailed Salamander (E. l. longicauda) that is found from New York southwestward to Kentucky, Tennessee, and the southern Appalachian region, and the Three-lined Salamander (E. l. guttolineata) that is primarily found in non-mountainous regions in the Southeast. The taxonomic status of Eurycea l. longicauda and E. l. guttolineata have been debated because the two subspecies coexist in close contact in scattered populations in the southern Appalachians with little evidence of interbreeding (Ireland 1979). The high peaks of the Appalachian Mountains form a natural barrier between E. l. longicauda to the west and E. l. guttolineata to the east. Near the southern extremity of the Appalachians this barrier disappears, and a broad zone of intergradation was reported to occur in northwestern Georgia and northern Alabama that continues northwestward (Mount 1975, Valentine 1962). Subsequent molecular studies of populations in this region by Carlin (1997) revealed that they are distinct species that do not interbreed. Specimens that were thought to be intergrades were found to fall within the range of variation seen within the two species and show no evidence of being hybrids. As currently recognized, E. longicauda contains two subspecies that intergrade in Missouri, and E. guttolineata constitutes a separate species.
Species Comments:
Identification
Description: Eurycea longicauda is geographically variable and contains two subspecies that intergrade in Missouri. The Dark-sided Salamander (E. l. melanopleura) is associated with the Ozark Highlands and has a yellow to yellowish brown dorsum that is marked with numerous, irregular black marks. A broad dark stripe occurs along each side that is marked with numerous light flecks. The belly is pale yellow and marked with indistinct light brown to gray spots (Petranka, 1998). The Long-tailed Salamander (E. l. longicauda) is found farther east, including in the southern Appalachians.

The Long-tailed Salamander, (E. l. longicauda) is a medium-sized salamander with a conspicuously long tail that proportionately increases with age and comprises 60-65% of the total length of adults. The juveniles and adults have a yellowish orange to yellowish brown dorsal ground color (rarely reddish) that in overlain with numerous irregular black spots. The spots are typically less dense and smaller on the head and tail relative to the body, and tend to coalesce on the sides to form broken lateral bands or bars. The tail is laterally compressed and the sides have elongated black marks that tend to form vertical chevrons and a herringbone pattern. The belly is light cream to yellowish and lacks spots (Beane et al. 2010, Petranka 1998). The adults vary from around 10-20 cm TL and there are typically 13 or 14 costal grooves between the limbs. Females from Ohio, Virginia and West Virginia average 5-9% larger in SVL than males (Petranka 1998). Sexually active males have prominent cirri, swollen cloacae, and papillose cloacal lips (smooth in females). They also have hedonic glands on the tail, a round mental gland, and elongated maxillary teeth (Niemiller and Reynolds 2011, Petranka 1998). Adult females also have nasal cirri, but they are far less developed than those of sexually active males.

The hatchlings average around 10-12 mm SVL and 17-19 mm TL. They have a uniformly cream colored to light brownish dorsum that contrasts sharply with the immaculate belly. The larvae are the stream-type with streamlined bodies and a narrow, dorsal fin that does not extend forward beyond the rear limbs. The gills are reddish to reddish brown and are often more elongated in specimens collected from sluggish or standing water. As larvae mature they gradually develop a color pattern that tends towards those of the adults. Within 1-2 months after hatching, larvae develop a dark broad band or heavy dark mottling on either side of the body. A more narrow and less conspicuous mid-dorsal stripe often develops on the back. Recently transformed animals have black sides, a dark chin, and an olive-gray dorsum (Petranka 1998). Long-tailed Salamander larvae lack the conspicuous paired light spots on the dorsum that are characteristic of members of the E. bislineata complex, including E. wilderae.
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Observation Methods: Individuals are occasionally seen crossing roads on rainy nights, but are otherwise best obtained by searching cover objects in or near streams or other breeding sites.
AmphibiaWeb Account
Distribution in North Carolina
Distribution Comments: Eurycea l. longicauda occurs from southern New York westward through Pennsylvania, Ohio, Indiana, and southern Illinois to eastern Missouri where it contacts and hybridizes with E. l. melanopleura. The range extends south and southwestward through Kentucky, West Virginia, western Virginia, central and eastern Tennessee, and western North Carolina before ending in northwestern Georgia, northern Alabama, and extreme northeastern Mississippi. This species is uncommon in North Carolina and patchily distributed in the Blue Ridge Mountains. Populations are currently known mostly from lower elevation sites in the larger river valleys and their tributaries.
Distribution Reference: Beane et al. (2010), Petranka (1998)
County Map: Clicking on a county returns the records for the species in that county.
GBIF Global Distribution
Key Habitat Requirements
Habitat: The Long-tailed Salamander has a biphasic life cycle and requires both aquatic sites for larval development and suitable terrestrial sites for the juveniles and adults. The adults prefer mesic hardwood forests and have frequently been found where shale, limestone, or other bedrocks that readily fissure allow access to underground springs and caverns. Females lay eggs in a wide variety of aquatic habitats, including cisterns and springhouses, vernal ponds, spring-fed ponds, springs and seepages, and headwater and lower-order streams (Petranka 1998). The larval period usually last less than a year, so seasonal habitats such as the ephemeral headwater sections of streams and vernal ponds often provide suitable habitats for the larvae. Long-tailed salamanders in the Ozark Uplift and in the Appalachian region are often found in or near abandoned mine shafts and caves (Petranka 1998). Anderson and Martino (1966) found them to commonly use limestone sink ponds in New Jersey and Niemiller and Reynolds (2011) reported them using ponds in central Tennessee. Populations in North Carolina are generally associated with running water habitats such as small to medium-sized, rocky streams and their associated springs and smaller tributaries. Specimens have also been found beneath rocks along a lakeshore and in a cliff face fissure.
See also Habitat Account for Montane Mesic Forests with Seepages/Headwater Streams
Life History and Autecology
Breeding and Courtship: The terrestrial adults undergo seasonal migrations to breeding and overwintering sites, although in some populations the adults remain close to water year-round (McDowell and Shepherd 2003). Nazdrowicz (2015) found that sexually active males in springhouse populations that he studied in Pennsylvania and Delaware begin moving into springhouses during July and were followed by females several weeks later. Larger individuals of both sexes tended to arrive first. Individuals in springhouses mated before most moved deep within the springs to overwinter. They reappeared again in springhouses during April and May, then dispersed to surrounding terrestrial habitats where they used cover that provided subterranean daytime retreats.

The seasonal patterns observed by Nazdrowicz (2015) appear to be generally similar to those observed elsewhere. Based on examination of the spermatogenic cycle, the adults in many populations appear to mate during the autumn and possibly through the winter and early spring (Ireland 1974, McDowell and Shepherd 2003, Siegel et al. 2014). Nazdrowicz (2015) made direct observations of four courting pairs in September in Pennsylvania and Delaware, which indicates that some populations begin mating during the summer and continue for one or more months thereafter.

Detailed descriptions of courtship are not available. Mating and oviposition appear to mostly occur in underground stream and spring passages and have rarely been observed by naturalists. Cooper (1960) observed partial courtship of a pair of E. l. longicauda in a mine in Maryland on 18 October that entailed elements of the persuasion phase in which the male chased a female and rubbed his head around her cloaca and snout. Nazdrowicz (2015) observed partial courtship sequences on four occasions in September. In one case the male moved his head over the dorsal surface of the female’s pelvic region and stroked his chin and throat across the female’s back towards her head several times in a slow and deliberate fashion. This behavior was accompanied by occasional nudging and rubbing of his snout along the female’s venter. He also observed the male make repetitive, posteriolateral head jerks and stroke the female's back with the left and right forelimbs. On a separate occasion he observed a female engaged in a tail straddle walk with a male in which the female straddled the male’s tail and placed her head over the male’s pelvic region. The male began undulating his tail with waves originating from the pelvic region and dampening towards the tail tip. After several seconds of undulations, the male lifted the base of his tail and deposited a spermatophore on the rock surface. However, the female did not successfully pick up the spermatophore.

The sex ratios among adults varies among populations. Sex ratios of males to females were about 1:1 in populations studied by Anderson and Martino (1966). In most populations that were monitored by Nazdrowicz (2015) the males outnumbered females by a ratio of two or three to one or more.
Reproductive Mode: Many females appear to begin laying eggs within a month or two after mating. Nazdrowicz (2015) made numerous observations of freshly laid eggs in a series of springhouses that he monitored in Pennsylvania and Delaware. He observed newly deposited eggs as early as 24 October and as late as 21 February, but the majority of eggs were deposited within a 2-4 week period in November. The eggs were laid singly on rocks, brick, cement, iron pipes, fibrous tree roots, wood, and artificial fences that were installed to observe egg laying. Females appeared to take up to a week to lay their entire clutch and abandoned their eggs after ovipositing. Oviposition occurred primarily on vertical and ceiling surfaces that were exposed to open water and were well-oxygenated. Cryptic microhabitats such as narrow passages between objects or the undersides of rocks were generally avoided. Nazdrowicz (2015) noted that ovipositing females occasionally consumed a nearby egg that was presumably not their own.

Franz (1964) found a female on 23 November in Maryland that was coiled about five eggs in early stages that were suspended singly just above a pool in a cave in Maryland, while Mohr (1943) found freshly laid eggs in a mine shaft in Pennsylvania on 2 January. The eggs were attached singly in running water to the tops and sides of rocks, and to the edges of submerged boards. Based on the examination of female reproductive tracts, McDowell and Shepherd (2003) concluded that oviposition occurs mostly in November and December in populations in Illinois, while Ireland (1974) found that E. l. melanopleura oviposits from December through March in Arkansas. Two clutches of eggs found in December were in a single row on the undersurface of a rock. Given the small number of observations of eggs in the wild, it is very likely that females in most populations oviposit in cryptic underground recesses such as those found in springheads, caves, and the banks of streams.

The number of eggs that are laid is difficult to determine because females do not brood and scatter their eggs widely. Ovarian egg counts for E. l. longicauda include a mean of 91 (range = 61-106) and a mean of 76 (57-120) for specimens examined by Hutchison (1956) and Nazdrowicz (2015). Clutch size was positively correlated with female SVL in the latter study. The freshly laid ova are yellowish, about 2.5-3.0 mm in diameter, and are surrounded by two jelly envelopes (Petranka 1998).

Embryonic developmental rates are temperature-dependent and depend on local site conditions. The embryos can take two or more months to reach the hatching stages (Mohr 1943, Nazdrowicz 2015, Petranka 1998). Hatchling E. l. melanopleura have been found in springs in winter and early spring (Rudolph 1978). Hatching in most Illinois populations appears to occur primarily from mid-February through April, but in a few cases as late as May or June, which indicates that some females may not oviposit until late winter (McDowell and Shepherd 2003). Anderson and Martino (1966) first found hatchlings in their pond populations in March, while Nazdrowicz (2015) estimated the embryonic period to last about 7-9 weeks in springhouse populations that he monitored, with hatching first occurring from early December through early March.

Aquatic Life History: Hatchlings from eggs that are laid in underground recesses and caves tend to move downstream and eventually enter surface streams where temperatures are warmer and food is more abundant. Nazdrowicz (2015) found that almost all larvae had moved from springhouses to surface streams by the end of April or late May, approximately 2-4 months after hatching. There was little drift downstream and larvae were abundant within the first 25 m of springhouses during the larval period. Anderson and Martino (1966) noted that the larvae are secretive and hide in leaf litter or emergent vegetation in shallow water during the day. Larvae in springs in Oklahoma fed primarily on ostracods, copepods, snails, fly larvae, isopods, beetles, and mayfly nymphs (Rudolph 1978).

The larval period is brief relative to many Eurycea species and typically lasts < 1 year, although a small percentage of animals in some populations may overwinter and transform the following summer (Petranka 1998). Larvae in limestone sink ponds in New Jersey that were studied by Anderson and Martino (1966) began hatching in March and grew 4-5 mm SVL per month before metamorphosing an estimated 65-70 days later in mid-June through July when they average 20-21 mm SVL and 41 mm TL. Growth rates of E. l. melanopleura in Arkansas increased between January and May, and larvae transformed in June and July when 23-28 mm SVL and 4-7 months old (Ireland 1974). Most E. l. melanopleura larvae studied by Rudolph (1978) transformed in late summer and autumn, but some overwintered and transformed the following year when > 1-year old. Metamorphosing larvae collected in Maryland in early July were 18-21 mm SVL (Franz and Harris 1965), while one large larva (51 mm TL) that was collected by Franz (1967) on 20 March presumably overwintered. Larvae in Nazdrowicz's (2015) study populations metamorphosed from May through mid-August with a peak in July, with an estimated larval period of 5-8 months depending on the population and year. We currently have no information on the larval life history of North Carolina populations.
Terrestrial Life History: Limited observations indicate that recently metamorphosed animals dispersed away from breeding sites within a few weeks after metamorphosing and live in the surrounding woods (Anderson and Martino (1966). The older juveniles and adults use terrestrial habitats that surround the breeding sites during the warmer months. Individuals hide beneath litter, logs or rocks during the day and emerge after dark during periods of favorable weather to feed on small invertebrates. Individuals tend to be most active during the first few hours after dark and feed on a wide variety of prey (Petranka 1998). The prey of New Jersey specimens included annelids, isopods, millipedes, centipedes, pseudoscorpions, phalangids, mites, ticks, spiders, homopterans, beetles, flies, ants, lepidopteran larvae, crickets, and thysanurans (Anderson and Martino 1967, Petranka 1998). The major prey of specimens from caves in Virginia were flies, orthopterans, and beetles (Hutchison 1958).

The young metamorphs typically require two years to reach sexual maturity, but Ireland (1974) reported that E. l. melanopleura in Arkansas can become sexually mature in 1-2 years after transforming when the males and females are 31-43 mm and 33-43 mm SVL, respectively. Individuals in other populations typically require two years to mature. McDowell and Shepherd (2003) found that males in Illinois became sexually maturity when > 45 mm SVL and females when > 48 mm SVL after their second year of growth. Similarly, Anderson and Martino (1966) estimated that males reach sexual maturity when > 43 mm SVL and females when > 46 mm SVL in New Jersey, with both sexes requiring around 2 years to mature. Nazdrowicz's (2015) also estimated that most individuals require two years to reach a SVL > 45 mm, with females perhaps requiring additional time to first reproduce.

The juveniles and adults in most populations undertake seasonal migrations to and from subsurface breeding and overwintering sites to the surrounding forests. The adults in many populations migrate to breeding and overwintering sites such as springs, cisterns and caves in late summer or early autumn. They do not resurface and disperse into the terrestrial habitats until the spring warm-up and often move on rainy nights (Guttman 1989, Nazdrowicz 2015, Petranka 1998). Mohr (1944) found large numbers of adults in an old mine shaft in Pennsylvania that remained there for about 8 months out of the year. They dispersed to the surrounding forest in late April and early May, then returned about 4 months later. Franz (1967) observed similar seasonal migrations to and from two Maryland caves. Minton (1972) noted that Indiana adults are active on the ground surface from April to October. Populations that McDowell and Shepherd (2003) studied in Illinois spent less time in underground retreats and were active on the ground surface in the immediate vicinity of breeding sites from late February through late November.

Anderson and Martino (1966) found that the juveniles and adults in their pond-breeding populations overwintered below ground in forests within 30 m of water rather than in aquatic sites. They then moved to the margins of ponds within a few weeks after they emergence in late April or early May. They spent the warmer months near the pond margins, then moved back to the forests and went underground with the onset of colder weather in October.
General Ecology
Population Ecology: We know very little about factors that regulate local population size. This is one of the few plethodontid salamanders tested to date that does not appear to be territorial (Jaeger 1988). The adults have been found in large aggregates beneath logs, in mines, and in springhouses (Petranka 1998). The larvae are palatable to fish, and fish predation may in part confine many local populations primarily to fish-free sections of headwater streams and springs (Rudolph 1978).
Community Ecology: The larvae often co-occur with other salamander larvae in both ponds and streams, but very little information is known about ecological interactions with other community members.
Adverse Environmental Impacts
Effects of Pollution: Brady (2016) surveyed salamander communities on 45–65 year-old forests that had become established on abandoned coal surface mine sites in Ohio and compared them with unmined reference sites. Stream-breeding species such as E. longicauda, E. bislineata, and Desmognathus fuscus were well-represented in reference forests but were missing from all reforested mine sites. Persistent acid drainage from the mined sites were the presumed cause of the absence of stream-breeding plethodontids from these forests.
Status in North Carolina
NHP State Rank: S1S2
Global Rank: G5
Status in North Carolina: T
Status Comments: Populations in North Carolina are on the easternmost periphery of the range and there is concern that many local populations may be in decline (e.g., Decker and Robinson 2009). Most of our records are more than 20 years old, with only a few relatively recent records. This species is now listed as threatened within the state.

Photo Gallery for Eurycea longicauda - Long-tailed Salamander

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Recorded by: Max Ramey
Watauga Co.		Recorded by: Max Ramey
Watauga Co.