Plethodon montanus Highton & Peabody, 2000
Northern Gray-cheeked Salamander, Northern Graycheek Salamander
Highton, R., and R. B. Peabody. (2000). Geographic protein variation and speciation in salamanders of the Plethodon jordani and Plethodon glutinosus complexes in the southern Appalachian Mountains with the description of four new species. Bruce, R. C., R. G. Jaeger, and L. D. Houck eds., The Biology of Plethodontid Salamanders: 31–93. New York, Kluwer Academic/Plenum Publishers.
© 2016 Dr. Joachim Nerz (1 of 20)
Plethodon montanus Highton and Peabody, 2000
David A. Beamer1
1. Historical versus Current Distribution. Northern gray-cheeked salamanders (Plethodon montanus) are recently described members of the P. jordani complex (Highton and Peabody, 2000). They tend to occur at higher elevations on Flat Top, Buckhorn, Burkes Garden, Knob, Clinch, and Brumley mountains in the Valley and Ridge Province of Virginia, and on Roan, Bald, Black, Max Patch, and Sandymush mountains in the Blue Ridge Province of Virginia, North Carolina, and Tennessee. There is no evidence to suggest that the current distribution differs from the historical distribution. A history of the nomenclature of this species is given in Highton and Peabody (2000). Much of the literature that deals with Plethodon metcalfi (southern gray-cheeked salamanders) refers to this species.
2. Historical versus Current Abundance. Highton (2003) surveyed 12 northern gray-cheeked salamander populations prior to 1988 and again after 1994, and found decreased numbers of animals (under similar sampling conditions and with similar effort) in 10 populations. Highton (2003) stresses that animals remain abundant, but these populations should continue to be monitored.
A possible decline in the numbers of northern gray-cheeked salamanders was noted after lumbering (Gordon et al., 1962). By comparing a sample made by Dunn 44 yr earlier, the authors determined that the post-lumbering population of salamanders was only 21% of the pre-lumbering population.
Clearcuts negatively affect populations of northern gray-cheeked salamanders. Petranka et al. (1993, 1994) found that these salamanders occur in much higher numbers in mature forests than in clearcuts. Petranka (1998) states that deforestation and the conversion of mixed hardwood forests to pine monocultures has eliminated or reduced a large number of eastern Plethodon populations. However, Ash (1988, 1997) and Ash and Bruce (1994) have strongly disagreed with Petranka's estimates and do not consider forestry practices to be having as strong an impact on native salamanders.
3. Life History Features.
A. Breeding. Reproduction is terrestrial. Courtship was observed on 3 August at Whitetop Mountain, Smyth County, Virginia (Organ, 1958), and on 31 July near Linville, Avery County, North Carolina (MacMahon, 1964).
Courtship is initiated when a male approaches a female and nudges her with his snout. The male then places his mental gland and nasolabial grooves in contact with the body of the female and begins a foot dance in which the limbs are raised and lowered off the substrate one at a time. The male moves forward to the female’s head, at which point he presses his mental gland on her head and nasolabial grooves. The male then circles beneath the female's chin and laterally undulates his tail. The female then places her chin on his tail and moves forward to straddle the tail. The pair moves forward in a tail-straddle walk until the male stops and begins to make lateral sacral movements. The female makes synchronous lateral head movements counter to the sacral movements of the male. The male then lowers his vent to the substrate and deposits a spermatophore. He then flexes his tail to one side and leads the female forward until her vent is over the spermatophore. The female then picks up the cap from the spermatophore with her cloacal lips (Organ, 1958; MacMahon, 1964).
i. Breeding migrations. Unlikely; breeding migrations are unknown in any Plethodon species.
ii. Breeding habitat. At a North Carolina site, courtship occurred within 1 m of a road in area almost covered with jewelweed (Impatiens sp.; MacMahon, 1964). The surrounding area is described as a second-growth mixed mesophytic woods at approximately 1,250 m.
i. Egg deposition sites. Unknown, but likely to be in underground cavities.
ii. Clutch size. Unknown.
C. Direct Development.
i. Brood sites. Unknown.
ii. Parental care. Unknown, but it is likely that females brood, as with other species of Plethodon.
D. Juvenile Habitat. Unknown, but presumably similar to adult habitat.
E. Adult Habitat. Northern gray-cheeked salamanders have been recorded from both virgin and second-growth forest (Dunn, 1917a; Gordon et al., 1962). Northern gray-cheeked salamanders in Avery County, North Carolina, are found in mixed mesophytic forest with a canopy consisting of read and white oak, maples, buckeyes, ironwood, gum, tulip trees, and a few scattered hemlocks. The understory at this site is sparse and composed of jewelweed, nettle shield fern, pipsissewa, goldenrod, twayblade, bellflowers, and pinesap (Gordon et al., 1962).
Northern gray-cheeked salamanders are reported from high elevations in spruce-fir forest (Walker, 1931). Here they are found under logs but not in stumps (Hoffman and Kleinpeter, 1948).
F. Home Range Size. Unknown, but small home ranges are typical for Plethodon species.
G. Territories. Selby et al. (1996) observed agonistic behavior in northern gray-cheeked salamanders.
H. Aestivation/Avoiding Dessication. Undocumented, but Plethodon species will move from forest floor habitats to underground sites in response to desiccating surface conditions.
I. Seasonal Migrations. Unknown, other than vertical movements from the forest floor to underground sites to avoid dry summer and cold winter conditions.
J. Torpor (Hibernation). Unknown, but animals presumably move to warmer underground sites to escape cold weather.
K. Interspecific Associations/Exclusions. In Avery County, North Carolina, northern gray-cheeked salamanders were found with eastern newts (Notophthalmus viridescens), Blue Ridge dusky salamanders (Desmognathus orestes), seal salamanders (D. monticola), red-backed salamanders (Plethodon cinereus), Yonahlossee salamanders (P. yonahlossee), white-spotted slimy salamanders (P. cylindraceus), and Blue Ridge two-lined salamanders (Eurycea wilderae; Gordon et al., 1962).
On Grandfather Mountain and in the Whitetop Mountain-Mount Rogers area, the following salamander species are reported in association with northern gray-cheeked salamanders: Weller’s salamanders (P. welleri), red-backed salamanders, Blue Ridge two-lined salamanders, Blue Ridge dusky salamanders, and pygmy salamanders (D. wrighti; Walker, 1931; Hoffman and Kleinpeter, 1948; Bogert, 1952; Thurow, 1963).
Hairston (1949) reports the following salamanders along with northern gray-cheeked salamanders from the Black Mountains area, Yancey County, North Carolina: spring salamanders (Gyrinophilus porphyriticus), red salamanders (Pseudotriton ruber), Blue Ridge two-lined salamanders, black-bellied salamanders (D. quadramaculatus), seal salamanders, Carolina mountain dusky salamanders (D. carolinensis), pygmy salamanders, shovel-nosed salamanders (D. marmoratus), Yonahlossee salamanders, and white-spotted slimy salamanders.
Northern gray-cheeked salamanders are widely sympatric with white-spotted slimy salamanders in the Blue Ridge province in the Black and Bald isolates, and most of the Roan isolate. There is no evidence of hybridization between these species (Highton and Peabody, 2000).
Southern Appalachian salamanders (P. teyahalee) are widely sympatric with the Max Patch and Sandymush isolates of northern gray-cheeked salamanders. There is no known morphological or genetic evidence of hybridization (Highton and Peabody, 2000).
The six isolates of northern gray-cheeked salamanders in the Valley and Ridge Province are widely sympatric with northern slimy salamanders (P. glutinosus). The two species also are sympatric in a small section of the Roan isolate of northern gray-cheeked salamanders. There is evidence of occasional hybridization (Highton and Peabody, 2000).
Northern gray-cheeked salamanders are widely sympatric with Yonahlossee salamanders at the eastern edge of the Blue Ridge isolate. There is no evidence of hybridization (Highton and Peabody, 2000).
The ranges of northern gray-cheeked salamanders and Cumberland Plateau salamanders (P. kentucki) overlap in Russell, Smyth, and/or Washington counties in Virginia, but they have not yet been collected at the same sites (Highton and Peabody, 2000).
In the Black Mountains, Yancey County, North Carolina, northern gray-cheeked salamanders and white-spotted slimy salamanders generally replace each other altitudinally. On four of five transects, there was no overlap of these species occurrences; at a fifth transect, the two occur together over an elevational range of 61 m (Hairston, 1951).
L. Age/Size at Reproductive Maturity. Unknown.
M. Longevity. Unknown.
N. Feeding Behavior. Northern gray-cheeked salamanders sometimes forage above ground. Many have been observed on trunks and low branches of shrubs ≤ 1 m from the ground. Groups were observed feeding on fungal gnats around decaying fungi, or on fruit flies and other insects at the base of trees from which sap flowed (Gordon et al., 1962).
The following food items are reported from specimens from Grandfather Mountain, Avery County, North Carolina: Formicidae, Araneida, lepidopteran larvae, coleopteran larvae, Collembola, Diplopoda, Chilopoda, Acarina, Stylommatophora, dipteran larvae, Annelida, Tipulidae, Lepidoptera, shed skin, Ichneumonidae, Diptera, Gryllacrididae, Cicadellidae, Coleoptera, Miridae, Cantharidae, Chrysomelidae, Stylommatophora, Isopoda, Cynipodea, Hymenoptera, Phoridae, Psychodidae, Chironomidae, Cydnidae, Fulgoridae, Scarabaeidae, Reduviidae, Carabidae, Curculionidae, Chelonethida, Vespidae, Gryllidae, Simuliidae, Histeridae, Pentatomidae, Mecoptera, Tingidae, Elateridae, Otitidae, Mycetophilidae, Cryptophagidae, Culicidae, Aphididae, Phalacridae, Dolichopodidae, Staphylinidae, Fungivoridae, and Hemiptera. Ants, mites, and springtails were eaten less frequently by larger individuals while millipedes, earthworms, craneflies, spiders, and centipedes were eaten more frequently. In general, the number of different food items increases in larger individuals (Whitaker and Rubin, 1971).
O. Predators. Undocumented, but probably include forest-dwelling snakes, birds, and small mammals.
P. Anti-Predator Mechanisms. Unknown specifically, but members of the P. jordani complex are known to assume defensive postures and to produce slimy tail secretions that predators find noxious (Brodie et al., 1979; Petranka, 1998). Members of the P. jordani complex frequently become immobile when initially contacted. Northern gray-cheeked salamanders were included in a field study on immobility; however, it is not possible to separate their behavior from the other members of this complex in this published data set. Immobility may increase survival by making the salamander less likely to be detected, especially by visually oriented predators (Dodd, 1989).
Q. Diseases. Unknown.
R. Parasites. Rankin (1937) lists parasites from at least two species of the P. jordani complex. It is not possible to determine which parasites were found in which salamander species, but some of these salamanders were northern gray-cheeked salamanders. The following parasites are noted: Crytobia borrelis, Eutrichomastix batrachorum, Hexamitus intestinalis, Karotomorpha swezi, Prowasekella longifilis, Tritrichomonas augusta, Brachycoelium hospitale and Crepidobothrium cryptobranchi.
4. Conservation. Northern gray-cheeked salamanders are not protected by any state. Among members of the P. jordani complex, northern gray-cheeked salamanders have one of the widest distributions. While relatively wide ranging, northern gray-cheeked salamanders are restricted to higher elevations. Suitable habitat at these elevations may be separated by stretches of lower uninhabited areas, and populations often are not continuous. Within this range there are many federal and state properties that contain suitable habitat for these salamanders.
Northern gray-cheeked salamanders are relatively resilient to disturbances, such as those associated with timbering operations, and frequently are found in second-growth forests and relatively small, fragmented woodlots (Gordon et al., 1962; D.A.B., personal observation).
As with all species of Plethodon, northern gray-cheeked salamanders do not migrate and have small home ranges. Thus, they can exist in habitats of smaller size than many other amphibian species. Conservation activities that promote mature closed-canopy forests should benefit this species.
Acknowledgments. Thanks to Richard Highton, who reviewed this account and gave us the benefit of his insight and experience.
1David A. Beamer
2Michael J. Lannoo
Literature references for Amphibian Declines: The Conservation Status of United States Species, edited by Michael Lannoo, are here.
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