Amphibians of North Carolina

Scientific Name:	Common Name:		Family (Alpha):

NC Records
Plethodontidae Members:
Gyrinophilus porphyriticus - Spring Salamander

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Taxonomy

Class: Amphibia

Order: Caudata

Family: Plethodontidae

Subfamily: Spelerpinae


Taxonomic Comments: As currently recognized, Gyrinophilus porphyriticus is a geographically variable species that appears to be a species complex with several undescribed forms. Brandon (1966, 1967) recognized four subspecies, and found numerous intergrades that occurred over broad geographic areas that were difficult to assign to subspecies. These include the Northern Spring Salamander (G. p. porphyriticus) that has a mottled or reticulate pattern on the dorsum and lacks distinct spots or flecks, the Kentucky Spring Salamander (G. porphyriticus duryi) that is salmon colored and has small black spots and flecks on the back, the Carolina Spring Salamander (G. porphyriticus dunni) that has profuse flecking on the back and sides and a pronounced light stripe that is bordered by black, and the Blue Ridge Spring Salamander (G. porphyriticus danielsi) that is similar to G. p. dunni but has distinct spots rather than flecking on the dorsum. Specimens from high elevations usually have conspicuous black and white mottling along the lower jaw. Kuchta et al. (2016) generated molecular sequence data for specimens collected from throughout the range of G. porphyriticus, as well as cave-dwelling Gyrinophilus from Tennessee that are currently recognized as separate species. They found four major clades (evolutionary lineages) with several genetically distinct subclades. One of the clades included the Tennessee cave species that were nested within G. porphyriticus populations. The data indicate that the G. porphyriticus complex originated prior to the beginning of the Pleistocene glacial period, with the major clades being 1-3 million years old or older. Historical river drainages at that time explain a significant portion of the phylogenetic diversity that is seen today. Kuchta et al. (2016) found that the major clades and subclades do not correspond well with the subspecies or intergrades recognized by Brandon (1966), and constitute a species complex. Several species will likely be described in the future, with perhaps two or three species from western North Carolina. Until then, we continue to recognize a single species within the state and do not recognize subspecies.
Species Comments:

Identification

Description: The adults are large, stout-bodied plethodontids that have a salmon to pinkish orange ground color that is overlain with diffuse black streaks or spots. These can sometimes form a vague reticulate pattern. One of the most distinctive features is a light line that extends from each eye to the tip of the snout along a raised ridge, the canthus rostralis. The white line is often paralleled by a faint gray to bold black line below (Petranka 1998). The chin and sides of the lower jaw of southern Appalachian forms are often brightly mottled with black and white. There is no conspicuous sexual dimorphism and the males lack a well organized mental gland. The adults are 11-21 cm TL and there are 17-19 costal grooves.

The hatchlings have dorsal fins that terminate near the rear limbs and range from 18-22 mm TL in the southern Appalachians (Bruce 1978). The larvae have long truncated snouts and small eyes relative to head size. The dorsal ground color varies from light yellowish brown to light gray or lavender and frequently has fine reticulations or flecking that are most conspicuous on older animals (Petranka 1998). They lack the conspicuous dorsal light spots seen in many Eurycea and Desmognathus species, as well as the dorsal dark dots that are typical of older Pseudotriton larvae. The juveniles resemble the adults, but are brighter colored.

Online Photos: Google iNaturalist

Observation Methods: The adults are far less abundant than those of most other stream-breeding salamanders, but can be found beneath cover objects and on the ground surface at night. The larvae are best obtained by searching in debris or beneath cover objects in streams or seepages.

AmphibiaWeb Account

Distribution in North Carolina

Distribution Comments: The Spring Salamander occurs from southern Maine and Quebec to central Alabama in formations associated with the Appalachian uplift. It is generally associated with mountainous regions, but has been found as low as 100 m in elevation in the southern portion of the range. In North Carolina Gyrinophilus porphyriticus occurs throughout the mountains and less commonly in the western Piedmont where it is restricted to cool, shaded streams and ravines.
Distribution Reference: Beane et al. (2010), Dodd (2013)
County Map: Clicking on a county returns the records for the species in that county.


GBIF Global Distribution

Key Habitat Requirements

Habitat: The Spring Salamander typically inhabits cool, heavily shaded habitats with small streams or other aquatic habitats that lack predatory fishes, although the larvae can be found in permanent sections of streams with fishes. The larval period last for more than one year, and populations are restricted to sites with permanent water flow and mesic forests. The larvae are commonly found is springs, seepages, caves, the headwater sections of small tributaries, and smaller streams downstream from the headwaters (Petranka 1998). Populations in the Piedmont of South Carolina are restricted to springs and small streams in deep ravines with mature hardwood forests (Bruce 1972). In more mountainous regions in western North Carolina populations can be found in or near springs, seepages and small to medium-sized streams. The adults reside in stream and seepages, in streamside forested habitats, and occasionally farther from streams where they feed on the forest floor during the warmer months of the year.

See also Habitat Account for Spring Runs and Brooks

Life History and Autecology

Breeding and Courtship: Males examined by Bruce (1972, 1978) from several populations in North Carolina contained sperm in their vas deferentia from October through June, which implies that there is both an autumn and spring mating season in the southern Appalachians. Most females in the southern Appalachians and vicinity oviposit annually and sex ratios are near 1:1 (Bruce 1969, 1972). Detailed observations of courtship have not been published.

Reproductive Mode: Most females probably lay their eggs in deep underground recesses in streams or seeps given that so few nests have been found. Females attach their eggs in monolayers to the undersides of rocks or other objects. When depositing eggs, a female turns upside down and uses her arched body and tail to brace against the substrate (Petranka 1998). The ova are light yellow, about 3.5-4.0 mm in diameter, and are surrounded by three jelly coats (Bishop 1941, Bruce 1969, 1972).

Oviposition typically occurs during the summer and the embryos hatch in late summer or autumn (Petranka 1998). In Pennsylvania, 15 eggs and an attending adult were found on 8 August. The eggs were in early developmental stages and were attached singly to the underside of a submerged stone in a spring. Two egg masses and attending females were found in a small spring in a spruce-fir forest in southwestern Virginia. One clutch found on 14 July contained 41 eggs in intermediate stages of development, while the second found on 6 August contained 66 eggs in advanced developmental stages. In both instances the eggs were attached singly in a monolayer to the undersides of large rocks that were partially embedded in the stream bank. The first clutch covered a circular area about 18 cm in diameter, and the second an oblong area that measured 8 x 20 cm (Petranka 1998). In North Carolina 24 eggs at the tail bud stage were found with an attending female in a small rivulet on 13 July (Bruce 1978). The eggs were attached singly to the underside of a rock buried about 25 cm below the stream surface. A second clutch along with a female was found near Mt. Mitchell on 22 October (Bishop 1924). The eggs were near hatching and were attached singly to the underside of a large rock in relatively still water.

Bruce (1969, 1972) documented marked variation in the clutch sizes of populations in the Piedmont and Blue Ridge of South and North Carolina which were in part related to differences in female body size. The mean number of mature ovarian eggs in populations in western North Carolina varied from 39-63, while individual clutches contained 16-106 ova. Clutch size was positively correlated with female SVL. At a common SVL, females in high elevation populations tend to have lower fecundities than those at lower elevations (Petranka 1998). In a related study, Bruce (1978) reported differences in adult body size, age at maturity, and clutch size in regional populations in the Cowee Mountains in North Carolina.

Aquatic Life History: The larvae live in gravel beds and beneath stones, logs, and other cover in springs, seepages and streams. They are secretive during the day and often reside in subterranean channels and pockets below the surface of stream beds (Bruce 1980). At night individuals often emerge from cover and subsurface retreats and forage on the stream bed (Resetarits 1991). The larvae are gape-limited predators that incorporate larger prey into their diets with age. Small larvae feed primarily on small invertebrates, but larger ones will take salamanders and other large prey such as crayfishes, along with smaller invertebrates. Stomach contents of larvae from a North Carolina population included oligochaetes, spiders, isopods, crayfish, centipedes, odonates, mayflies, stoneflies, caddisflies, flies, salamander eggs, adult and larval Eurycea wilderae, and adult D. ocoee (Bruce 1979). Gyrinophilus larvae in New Hampshire populations feed on two-lined salamander larvae and a variety of aquatic invertebrates including beetles, caddisflies, and dipterans (Petranka 1998).

In North and South Carolina there is no relationship between the seasonal surface activity of larvae and the elevation of local populations (Bruce 1972). Specimens can be found in aquatic sites during every month of the year. Although the larvae are generally far less abundant than those of other plethodontids such as Desmognathus and Eurycea, densities in some Virginia streams may reach 5-10 larvae/m2 of stream bed (Resetarits 1991, 1995). Estimating length of the larval period is challenging because the larvae have variable growth rates and are difficult to collect in large numbers. Bruce (1980) estimated a modal length of around 4 years for a population in western North Carolina. Most larvae transform in late June through late August when 55-65 mm SVL in low elevation (< 1200 m) populations and when 61-70 mm SVL in montane populations at elevations >1200 m (Bruce 1972, 1978, 1980).

Terrestrial Life History: The juveniles and adults live beneath surface or subsurface objects in or near springs or seepages and can occasionally be found under cover objects or foraging at night on the adjoining forest floor. Bishop (1941) found that adults move to subsurface retreats in or near springs and seepages during the winter, as well as in summer when surface flow is greatly reduced.

The seasonal duration and extent of surface activity in North and South Carolina increases with elevation. Bruce (1972) found that proportionately few juveniles and adults occur in lowland samples from the Piedmont where individuals are active on the ground surface from about mid October to late March. At intermediate elevations in the Blue Ridge Embayment, metamorphosed individuals comprise a greater proportion of samples, and juveniles and adults are active on or near the ground surface between mid-September through late June. In high elevation populations in the Nantahala Mountains, metamorphosed salamanders tend to be active near the surface during all months of the year.

Males at low to intermediate elevations in the southern Appalachians become sexually mature immediately after metamorphosing, while those at higher elevations require as much as a year longer to mature after metamorphosing (Bruce 1972). The minimum size at maturity varies substantially among populations. In a group of lowland populations all males > 55 mm were sexually mature. In a second group from higher elevations, some juveniles did not mature until they exceeded 81 mm SVL. Females have similar trends and have delayed development in high elevation populations. Some females in low elevation populations mature shortly after transforming, but those in high elevation populations may not mature for 1 year or more after metamorphosing. Depending on the locality, females may mature when as small as 61 mm SVL or as large as 82 mm SVL.

In northern populations the adults feed primarily on invertebrates, but they regularly feed on other salamanders in the southern Appalachians where the overall surface density of salamanders is often very high (Petranka 1998). Bruce (1972) reported that about half of the Gyrinophilus with identifiable remains that he examined from montane populations in North Carolina contained salamanders, including Eurycea wilderae, Desmognathus ocoee, Plethodon jordani, P. serratus, Pseudotriton ruber, and a conspecific. The remainder of the diet consisted of invertebrates, including large earthworms, beetle larvae, slugs, and snails. In a related study the adults fed on oligochaetes, isopods, spiders, centipedes, insects, adult and larval E. wilderae, adult D. ocoee and Plethodon species. Salamanders comprised a larger component of the diet in metamorphosed Gyrinophilus than in larvae (Bruce 1979).

General Ecology

Population Ecology: Local populations appear to be smaller than those of most co-occurring stream-breeding species based on surveys for the larvae and adults. In North Carolina, this likely reflects the fact that the adults and larvae frequently prey on other salamanders. Populations tend to be mostly restricted to lower-order stream systems and it is uncertain to what extend local populations are connected through occasional dispersal between demes. The downstream drift and washout of larvae during flood events may be important in facilitating dispersal between local populations, but more information is needed on the genetic structure of populations at different spatial scales.

Community Ecology: Several researchers have examined the interactions of Gyrinophilus larvae with other community members. Although Gyrinophilus larvae reach their highest densities in stream sections without fish, they often coexist with predatory fish and appear to lack chemical or behavioral defenses against fish. Resetarits (1991, 1995) found that Brook Trout decreased the survival and growth of larvae in artificial stream systems. Gustafson (1994) examined ecological interactions between E. bislineata larvae and small and large Gyrinophilus larvae in experimental streams and pools. Both sizes of Gyrinophilus larvae preyed on Eurycea, and Eurycea were less active on the surface when large Gyrinophilus were present as compared to small Gyrinophilus. Beachy (1994) found that Eurycea experienced lower survival in the presence of predatory G. porphyriticus and D. amphileucus larvae, and that Gyrinophilus was the more effective predator. Although both species share a common resource, there is no evidence that Gyrinophilus and D. amphileucus compete for food. The adults in the southern Appalachians regularly feed on the diverse array of salamanders that inhabit streamside and woodland communities, but very little is known about the extent to which they influence the community composition and structure of streamside salamander communities.

Adverse Environmental Impacts

Status in North Carolina

NHP State Rank: S5

Global Rank: G5

Photo Gallery for Gyrinophilus porphyriticus - Spring Salamander		21 photos are shown.
	Recorded by: B. Bockhahn United States Co.	Recorded by: Andrew W. Jones Polk Co.
	Recorded by: Andrew W. Jones Polk Co.	Recorded by: tom ward Buncombe Co. Comment: An individual feeding on a Seal Salamander.
	Recorded by: tom ward Buncombe Co.	Recorded by: tom ward Buncombe Co.
	Recorded by: B. Bockhahn, J. Thomson Yancey Co.	Recorded by: B. Bockhahn, J. Thomson Yancey Co.
	Recorded by: Max Ramey Watauga Co.	Recorded by: Max Ramey Watauga Co.
	Recorded by: tom ward & ben urfer Buncombe Co.	Recorded by: tom ward & ben urfer Buncombe Co.
	Recorded by: tom ward Buncombe Co.	Recorded by: tom ward Buncombe Co.
	Recorded by: tom ward & ben urfer Buncombe Co.	Recorded by: B. Bockhahn, CHRO staff Rutherford Co.
	Recorded by: tom ward Buncombe Co.	Recorded by: Steve Hall, NHP Staff Avery Co.
	Recorded by: Owen McConnell Graham Co.	Recorded by: Owen McConnell Graham Co.
	Recorded by: Jim Petranka Buncombe Co.

Gyrinophilus porphyriticus - Spring Salamander

Photo Gallery for Gyrinophilus porphyriticus - Spring Salamander

21 photos are shown.