Amphibians of North Carolina
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NC Records

Plethodon jordani - Red-cheeked Salamander

Taxonomy
Class: Amphibia Order: Caudata Family: Plethodontidae Subfamily: Plethodontinae Other Common Name(s): Jordan's Salamander
Taxonomic Comments: The Plethodon jordani complex is a group of closely related forms that mostly are found at higher elevations in the southern Appalachian Mountains. Because they are largely restricted to mid to higher elevations and are absent from valleys, they form a series of geographic isolates that have diverged from one another to varying degrees. Depending on geographic location, the adults may be unmarked, or have red cheeks, red legs, or brassy frosting on their backs. Members of this group were traditionally treated as a single, geographically variable species known as Jordan's Salamander (Plethodon jordani). Hairston (1950) recognized seven subspecies of P. jordani, but Highton (1962) rejected these because some characters do not vary concordantly among the recognized taxa. In later papers, Highton (1970, 1972) recognized 12 geographic isolates in southern populations and argued that recognizing subspecies would serve no useful purpose.

Molecular studies (Highton and Peabody 2000, Weisrock and Larson 2006) have since revealed the presence of seven major evolutionary lineages that are now recognized as separate species (P. amplus, P. cheoah, P. jordani (sensu stricto), P. meridianus, P. metcalfi, P. montanus, and P. shermani). Some of the recognized species within the Plethodon jordani complex show both historical and current-day genetic influence from one or more other species, and mating studies indicate that none of these forms are completely reproductive isolated due to mating barriers (Reagan 1992, Wiens et al. 2006). Where adjoining forms come into contact, they generally tend to hybridize to varying degrees. In addition, certain members of this group also hybridize with members of the Plethodon glutinosus complex (Weisrock et al. 2005, Wiens et al. 2006). Discordance between mtDNA and nuclear data are well-documented (Weisrock and Larson 2006) and reflect various levels of gene-mixing. This not only is occurring today, but likely occurred historically during the Pleistocene as ranges expanded during glacial periods and previously isolated forms came into contact. Here, we recognize all of these species and discuss issues with hybridization in the individual species accounts.
Species Comments:
Identification
Description: This is a large Plethodon species that is dark gray to bluish black above with a grayish venter. A conspicuous red cheek patch is present on each side of the head of the adults. The tail of adults is slightly longer than the body and is rounded in cross-section. Sexually-active males have conspicuous, rounded mental glands. The adults vary from 8.5-18.5 cm TL and there are usually 16 costal grooves. The young juveniles lack red cheeks but often have red spots on the dorsum. Morphs of Desmognathus imitator with red cheeks superficially resemble this species, but can be distinguished by their smaller size, head shape, relatively large back legs, dorsal patterning, and a light diagonal line that extends from the eye to the angle of the jaw.

Plethodon jordani hybridizes with P. metcalfi on Balsam Mountain and Hyatt Ridge where the Great Smokies meet the Balsam Mountains. A hybrid zone is present where hybrids have reduced amounts of reddish pigmentation on the cheeks (Hairston et al. 1992, Hatfield et al. 2010). Plethodon jordani also hybridizes with P. teyahalee at intermediate elevations in the Great Smoky Mountains National Park (Highton and Peabody 2000). Hybrids between these two typically have reduced red pigmentation on the cheeks and fine white speckling or spotting on the body. Pure P. jordani specimens from outside of the hybrid zone completely lack white speckling, while pure P. teyahalee specimens from outside of the hybrid zone completely lack red pigmentation on the cheeks.
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Observation Methods: The juveniles and adults can be easily observed after they emerge with the onset of darkness and forage and mate on the forest floor. They also reside beneath surface cover during the warmer months of the year.
AmphibiaWeb Account
Distribution in North Carolina
Distribution Comments: The Red-cheeked Salamander is only found in the Great Smoky Mountains in southwestern North Carolina and adjoining areas of Tennessee.
Distribution Reference: Highton and Peabody (2000)
County Map: Clicking on a county returns the records for the species in that county.
GBIF Global Distribution
Key Habitat Requirements
Habitat: Plethodon jordani inhabits mesic hardwood forests and spruce-fir forests at mid to high-elevation sites. It is generally found above 2800' in elevation. Mature forests with moist microhabitats, abundant leaf-litter, bark slabs, and coarse woody debris provide ideal conditions.
Biotic Relationships: Plethodon jordani and Desmognathus imitator are elements of a Batesian mimicry complex (Petranka 1998). Jordan's Salamander has a slimy tail that produces glutinous secretions that are noxious to potential avian predators. This species serves as the model and its red cheek patches presumably function as aposematic (warning) coloration. Desmognathus imitator lacks comparable chemical defenses and is the mimic. Feeding trials show that birds will often release captured red-cheeked D. imitator after several encounters with noxious P. jordani, but almost always kill and eat D. imitator that lack red cheeks (Brodie and Howard 1973, Brodie et al. 1979). Typically, only about 15-20% of individuals in local populations of D. imitator have red cheeks.
See also Habitat Account for General Montane Mesic Forests
Life History and Autecology
Breeding and Courtship: Arnold (1976) observed courtship of members of the P. jordani complex from four geographic isolates and found them all to be indistinguishable. The following is a summary of the major aspects of breeding and courtship as summarized by Petranka (1998). These presumably apply to all seven recognized species in this complex.

The adults appear to court from July through early October based on field observations. A female normally lays a clutch of eggs every other year, and only mates during the year when she nest. Arnold (1976) noted that males will not aggressively court sexually mature females that have small ovarian eggs, but will actively court females with large ovarian eggs.

When courting, a male approaches a female and begins nudging, nosing, or tapping her with his snout. He then places his mental gland and nasolabial grooves in contact with the back, sides, or tail of the female and engages in a 'foot dance' in which the limbs are raised and lowered off the substrate one at a time. The male eventually moves forward and presses his mental gland along the side of the female's head and over her nasolabial grooves. The male then turns his head under the female's chin and lifts. Next, the male circles under the female's chin and laterally undulates his tail as he passes. If the female is responsive, she places her chin on his tail and moves forward to the base of the tail. The pair then engages in a tail-straddle walk that may last for 1 hour. During the walk the male may turn and slap the female across her nasolabial region with his mental gland. This introduces pheromones via the nasolabial grooves.

The male eventually stops moving and begins a series of lateral rocking movements of his sacrum. The female begins a series of synchronous lateral head movements counter to the lateral movements of the sacral region of the male. The male then presses his vent to the substratum and deposits a spermatophore. Immediately after, he flexes the tail to one side and leads the female forward. She stops when her vent is over the spermatophore, then lowers her sacrum and picks up the sperm cap. During this process the male arches the sacral region and does a series of pushup motions with the rear limbs. The pair usually splits up and terminates courtship shortly after spermatophore deposition, even if the female is unsuccessful in picking up the spermatophore.
Reproductive Mode: The nests of P. jordani have never been found. Gravid females with large ova move underground in late spring or early summer and presumably oviposit in deep underground recesses (Hairston 1983a). Females oviposit in May in the southern Appalachians based on the presence or absence of gravid females in samples. Specific data on ovarian complements have not been published, but most females likely produce clutches of 10-15 eggs (Hairston 1983a). Hatching presumably occurs underground in late summer or early autumn, about 2-3 months after the eggs are deposited.
Aquatic Life History: This is a fully terrestrial species and there is no aquatic larval stage.
Terrestrial Life History: Hatchlings in western North Carolina stay beneath ground for 10-12 months before moving to the ground surface (Hairston 1983a). Small individuals that are about 1-year old first appear on the surface in May or June. Hairston (1983a) estimated that males mature sexually about 3 years after hatching when > 43 mm SVL. An estimated 25% of the females oviposit for the first time 4 years after hatching, while another 25% oviposit for the first time 1 year later. The remaining animals first oviposit 6 years or more after hatching, then every other year for the remainder of their lives. The smallest gravid females are 46 mm SVL.

During the warmer months of the year individuals live in burrows and other underground passageways or beneath rocks, logs, or other surface cover (Petranka 1998). Juveniles and adults emerge from cover at dusk and forage on the forest floor where they feed on small invertebrates. On moist or rainy nights a small percentage of individuals can often be observed perched on ferns or other low-lying vegetation. Individuals may also climb short distances up tree trunks (Hairston 1987). The adaptive value of climbing is not fully understood, but could potentially function for feeding, sensing the presence of mates using chemical cues, thermoregulation, or avoiding predators. Individuals retreat to underground overwintering sites with the onset of cold weather in the fall, and emerge soon after the spring leaf-out.

The Red-cheeked Salamander appears to be an opportunistic, gape-limited predator. Powders and Tietjen (1974) found a diverse array of prey in the diet, including annelids, snails, millipedes, centipedes, isopods, harvestmen, pseudoscorpions, mites, spiders, and a variety of insects. Millipedes, annelids, beetles, and insect larvae were the most volumetrically important prey. Millipedes were more important during the spring, and insect larvae were more important during the autumn months. In addition, collembolans and annelids tended to increase in importance with elevation.
General Ecology
Population Ecology: Members of the P. jordani complex that inhabit mature forests have relatively stable populations that likely reflect density-dependent regulation associated with territoriality (Hairston 1987). The adults will aggressively defend their small territories from both conspecific and heterospecific intruders. This likely sets an upper limit on local population size and mediates competition with congenerics such as P. teyahalee (Hairston 1987, Nishikawa 1985, 1990, Petranka 1998). Merchant (1972) estimated the average home ranges for males, females, and juveniles as 11.4, 2.8, and 1.7 m2, respectively.
Community Ecology: Hairston (1980a, 1983b) conducted field experiments to examine competitive interactions between members of the P. jordani complex and P. teyahalee, and documented the responses of each species to the experimental removal of the other. Experiments were conducted both in the Great Smoky Mountains with P. jordani where the elevational overlap is slight, and in the Balsam Mountains with P. metcalfi where the overlap is much more extensive. In both cases the removal of either P. jordani or P. metcalfi increased the surface densities of P. teyahalee. Removal of P. teyahalee did not affect the densities of P. jordani, but did increase the proportion of juveniles in the population. Hairston concluded that competition occurs between the two species, but was unable to pinpoint the resources that were in short supply. Later studies (see Hairston 1987, Nishikawa 1985, 1990, Petranka 1998) suggest that aggressive interactions related to interference competition for space is the primary factor mediating interactions between these species.

Gifford and Kozak (2011) also examined interactions between P. teyahalee and P. jordani in the Great Smoky Mountains and concluded that P. jordani is restricted to higher elevations because of physiological constraints, while competition with P. jordani prevents P. teyahalee from expanding its range to include higher-elevation habitats.
Adverse Environmental Impacts
Status in North Carolina
NHP State Rank: S3
Global Rank: G4
Status in North Carolina: W3
Protected Lands: Great Smoky Mountains National Park
Environmental Threats: The range of the Red-cheeked Salamander is entirely within protected federal lands and has few environmental threats. The long-term effects of acid deposition on high-elevation populations of salamanders have not been examined, but could potentially alter the composition of salamander assemblages and affect species such as P. jordani. Climate change could ultimately affect this species as average temperatures rise with time.
Status Comments: The Red-cheeked Salamander is a narrow endemic that is found only in the Great Smoky Mountains of eastern Tennessee and western North Carolina. It is a very abundant and widespread species where it occurs. Long-term studies indicate that populations have not declined since the 1960's, even though other large Plethodon in the region do show evidence of decline (Caruso and Lips 2012).

Photo Gallery for Plethodon jordani - Red-cheeked Salamander

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Recorded by: Todd Pierson
Swain Co.