Description
Adult males are slightly smaller than the females, ranging from 58-91 mm in total length and averaging 73 mm. Adult females range from 64-90 mm and average 78 mm. The largest individual on record is 122 mm (Bishop 1943).
The red-backed phase of this species is characterized by a broad, dorsal band running down the midline from the head onto the tail. The color of the stripe varies from light gray or dull yellow to pink, brick-red, and bright red. There are often small flecks of black within the band. The sides are dark gray or black, becoming lighter and mottled toward the belly, which is strongly mottled with white and gray. In contrast, the lead-backed phase lacks the dorsal band and is uniformly dark gray to almost black, with the head and legs usually lighter (Bishop 1943). There is also an erythristic color phase that is mostly red, apparently to mimic juvenile Notophthalmus viridescens (Tilley et al. 1982). Juveniles of the red-backed phase have a well developed dorsal band and the upper sides are strongly pigmented (Bishop 1943)
The body is long and fairly slender, is slightly flattened dorsally, and is well rounded on the sides. The cross section of the tail is nearly circular throughout its length. Regenerating tails are flattened laterally and are usually uniform dark gray. Number of costal grooves ranges from 17 to 20, but there are usually 18 or 19. The gular fold is prominent. The legs are small with short, thick toes. There are four fingers, which in order from longest to shortest are 3-2-4-1. The five toes are slightly webbed, and are 3-4-2-5-1 in order from longest to shortest. The vomerine teeth form two backward-curving lines of 5-7 teeth separated from each other and from the parasphenoid teeth, which are in two imperfectly separated patches. The mouth is fairly large, with the angle of the jaw behind the eye. The small tongue does not fill the floor of the mouth. Males can be identified when in breeding condition by swollen snout, enlarged premaxillary teeth, and proportionally longer legs (Bishop 1943). Black testes can also be seen through the abdominal wall when transiluminated by a strong light (Jaeger et al. 2002a).
As is the case for all members of the genus Plethodon, eggs are laid in terrestrial cavities attended by the female. The larval stage is passed within the egg capsule. The broad, flat, leaf-like gills rise from a common base, are often fully developed at hatching, and then persist for only a few days (Bishop 1943). Embryos average about 19 mm upon hatching and individuals less than 32 mm in snout-vent length are considered to be juveniles (Bishop 1943; Jaeger et al. 2002a). Juveniles have proportionately broad heads, which allows them to forage on a wide range of prey (Maglia 1996). The fingers and toes of the juveniles are well indicated, the inner and outer short (Bishop 1943).
While populations from the formerly glaciated part of the range are very uniform, allozyme studies show that when its entire range is considered, P. cinereus consists of four genetically differentiated geographic groups with within-group D-values ranging from 0-0.15 and between-group D-values ranging from 0.02-0.24. This indicates that the groups living in the unglaciated localities have been isolated from each other for 1.5-2.7 million years, and that populations from formerly glaciated areas are all descended from the same group. Despite their long divergence, there is still extensive gene flow between the groups at the points where they contact one another (Highton and Webster 1976; Highton 2000).
- Highton, R. (2000). ''Detecting cryptic species using allozyme data.'' The Biology of Plethodontid Salamanders. R.C. Bruce, R.G. Jaeger, and L.D. Houck, eds., Kluwer Academic/Plenum Publishers, New York, New York, 215-241.
- Conant, R. and Collins, J.T. (1998). A Field Guide to Reptiles and Amphibians of Eastern and Central North America. 3rd Edition. Houghton Mifflin Company, Boston, Massachusetts.
- Bishop, S.C. (1943). Handbook of Salamanders. Comstock Publishing Company, Inc., Ithaca, New York.
- Alford, R.A., and Richards, S.J. (1999). ''Global amphibian declines: A problem in applied ecology.'' Annual Review of Ecology and Systematics, 30, 133-65.
- Anthony, C. D., Venesky, M. D., and Hickerson, C. A. M. (2008). ''Ecological separation in a polymorphic terrestrial salamander.'' Journal of Animal Ecology, 77, 646-653.
- Bazar, M. A., Quinn, M. J., Jr., Mozzachio, K., Bleiler, J. A., Archer, C. R., Phillips, C. T., and Johnson, M. S. (2008). ''Toxicological responses of red-backed salamanders (Plethodon cinereus) to soil exposures of copper.'' Archives of Environmental Contamination and Toxicology, 57, 116-122.
- Becker, M. H., and Harris, R. N. (2010). ''Cutaneous bacteria of the redback salamander prevent mortality associated with a lethal disease.'' PLoS One, 5(6), e10957.
- Bergeron, C. M., Bodinof, C. M., Unrine, J. M., and Hopkins, W. A. (2010). ''Mercury accumulation along a contamination gradient and nondestructive indices of bioaccumulation in amphibians.'' Environmental Toxicology and Chemistry, 29, 980-988.
- Brodie, E. D., Jr., and Brodie, E. D. III (1980). ''Differential avoidance of mimetic salamanders by free-ranging birds.'' Science, 208, 181-182.
- Bruce, R. C. (2008). ''Intraguild interactions and population regulation in plethodontid salamanders.'' Herpetological Monographs, 2008, 31-53.
- Brucker, R. M., Baylor, C. M., Walters, R. L., Lauer, A., Harris, R. N., and Minbiole, K. P. C. (2008). ''The identification of 2,4-diacetylphloroglucinol as an antifungal metabolite produced by cutaneous bacteria of the salamander Plethodon cinereus.'' Journal of Chemical Ecology, 43, 39-43.
- Brucker, R. M., Harris, R. N., Schwantes, C. R., Gallaher, T. N., Flaherty, D. C., Lam, B. A., and Minbiole, K. B. C. (2008). ''Amphibian chemical defense: Antifungal metabolites of the microsymbiont Janthinobacterium lividum on the salamander Plethodon cinereus.'' Journal of Chemical Ecology, 34, 1422-1429.
- Burton, T.M. (1975). ''Salamander populations and biomass in the Hubbard Brook Experimental Forest, New Hampshire.'' Copeia, 1975(3), 541-546.
- Cabe, P. R., Page, R. B., Hanlon, T. J., Aldrich, M. E., Connors, L., and Marsh, D. M. (2007). ''Fine-scale population differentiation and gene flow in a terrestrial salamander (Plethodon cinereus) living in a continuous habitat.'' Heredity, 98, 53-60.
- Cassell, R. W., and Jones, M. P. (2005). ''Syntopic occurrence of the erythristic morph of Plethodon cinereus and Notophthalmus viridescens in Pennsylvania.'' Northeastern Naturalist, 12, 169-172.
- Davis, A. K. (2010). ''Lead-phase and red-stripe color morphs of red-backed salamanders Plethodon cinereus differ in hematological stress indices: A consequence of differential predation pressure?'' Current Zoology, 56, 238-243.
- Deitloff, J., Adams, D. C., Olechnowski, B. F. M., and Jaeger, R. G. (2008). ''Interspecific aggression in Ohio Plethodon: implications for competition.'' Herpetologica, 64, 180-188.
- Ducey, P.K., Schramm, K., and Cambry, N. (1994). ''Interspecific aggression between the sympatric salamanders, Ambystoma maculatum and Plethodon cinereus.'' American Midland Naturalist, 131, 320-329.
- Fitzpatrick, B. M., Shook, K., and Izally, R. (2009). ''Frequency-dependent selection by wild birds promotes polymorphism in model salamanders.'' BMC Ecology, 9, 12.
- Gabor, C.R., and Jaeger, R.G. (1995). ''Resource quality affects the agonistic bevariour of territorial salamanders.'' Animal Behavior, 49, 71-79.
- Gabor, C.R., and Jaeger, R.G. (1999). ''When salamanders misrepresent threat signals.'' Copeia, 1999(4), 1123-1126.
- Gillette, J. R., Jaeger, R.G., and Peterson, M.G. (2000). ''Social monogamy in a territorial salamander.'' Animal Behavior, 59, 1241-1250.
- Griffis, M. R., and Jaeger, R. G. (1998). ''Competition leads to an extinction-prone species of salamander: interspecific territoriality in a metapopulation.'' Ecology, 79, 2494-2502.
- Grover, M. C. (2000). ''Determinants of salamander distributions along moisture gradients.'' Copeia, 2000, 156-168.
- Grover, M. C., and Wilbur, H. M. (2002). ''Ecology of ecotones: interactions between salamanders on a complex environmental gradient.'' Ecology , 83, 2112-2123.
- Grover, M.C. (1998). ''Influence of cover and moisture on abundances of the terrestrial salamanders Plethodon cinereus and Plethodon glutinosis.'' Journal of Herpetology, 32, 489-497.
- Grover, M.C. and Wilbur, H.M. (2002). ''Ecology of ectones: interactions between salamanders on a complex environmental gradient.'' Ecology, 83, 2112-2123.
- Hairston, N.G. (1996). Long-term studies of vertebrate communities. Academic Press, New York, NY.
- Harris, R. N., Brucker, R. M., Walke, J. B., Becker, M. H., Schwantes, C. R., Flaherty, D. C., Lam, B. A., Woodhams, D. C., Briggs, C. J., Vredenburg, V. T., and Minbiole, K. P. C. (2009). ''Skin microbes on frogs prevent morbidity and mortality caused by a lethal skin fungus.'' The ISME Journal, 3, 818-824.
- Harris, R. N., Lauer, A., and Simon, M. A. (2009). '' Addition of antifungal skin bacteria to salamanders ameliorates the effects of chytridiomycosis.'' Diseases of Aquatic Organisms, 83, 11-16.
- Highton, R. (1959). ''The inheritance of the color phases of Plethodon cinereus.'' Copeia, 1959(1), 33-37.
- Highton, R. (1975). ''Geographic variation in genetic dominance of the color morphs of the red-backed salamander, Plethodon cinereus.'' Genetics, 80, 363-374.
- Highton, R. (1995). ''Speciation in eastern North American salamanders of the genus Plethodon.'' Annual Review of Ecology and Systematics, 26, 579-600.
- Highton, R. (1999). ''Geographic protein variation and speciation in the salamanders of the Plethodon cinereus group with the description of two new species.'' Herpetologica, 55, 43-90.
- Highton, R., and Webster, T.P. (1976). ''Geographic protein variation and divergence in populations of the salamander Plethodon cinereus.'' Evolution, 30, 33-45.
- Jaeger, R.G. (1978). ''Plant climbing by salamanders periodic availability of plant dwelling prey.'' Copeia, 1978(4), 686-691.
- Jaeger, R.G. (1979). ''Season spatial distribution of the terrestrial salamander Plethodon cinereus.'' Herpetologica, 35, 90-93.
- Jaeger, R.G. (1980). ''Density-dependent and density-independent causes of extinction of a salamander population.'' Evolution, 34, 617-621.
- Jaeger, R.G. (1980). ''Fluctuations in prey availability and food limitation for a terrestrial salamander Plethodon cinereus.'' Oecologia, 44, 335-341.
- Jaeger, R.G. (1981). ''Dear enemy recognition and the costs of aggression between salamanders.'' Animal Behavior, 29, 1100-1105.
- Jaeger, R.G. (1981). ''Foraging tactics of a terrestrial salamander Plethodon cinereus choice of diet in structurally simple environments.'' American Naturalist, 117, 639-664.
- Jaeger, R.G. (1984). ''Agonistic behavior of the Red-Backed Salamander Plethodon cinereus.'' Copeia, 1984(2), 309-314.
- Jaeger, R.G. and M.G. Peterson (2002). ''Familiarity affects agonistic interactions between female Red-Backed Salamanders.'' Copeia, 2002(3), 865-869.
- Jaeger, R.G., Gabor, R.C., and Wilbur, H.M. (1998). ''An assemblage of salamanders in the southern Appalachian mountains: competitive and predatory behavior.'' Behavior, 135, 795-821.
- Jaeger, R.G., Gillette, J.R., and Cooper, R.C. (2002). ''Sexual coercion in a territorial salamander: males punish socially polyandrous female partners.'' Animal Behavior, 63, 871-877.
- Jaeger, R.G., Goy, J., Tarver, M., and Marqez, C. (1986). ''Salamander Plethodon cinereus territoriality pheromonal markers as advertisement by males.'' Animal Behavior, 34, 860-864.
- Jaeger, R.G., Kalvarsky, D., and Shimizu, N. (1982). ''Territorial behavior of the Red-Backed Salamander Plethodon cinereus expulsion of intruders.'' Animal Behavior, 30, 490-496.
- Jaeger, R.G., Prosen, E.D., and Adams, D.C. (2002). ''Character displacement and aggression in two species of terrestrial salamanders.'' Copeia, 2002(2), 391-401.
- Jaeger, R.G., Schwarz, J., and Wise, S.E. (1995). ''Territorial male salamanders have foraging tactics attractive to gravid females.'' Animal Behavior, 49, 633-639.
- Jaeger, R.G., Wicknick, J.A., Griffis, M.R., and Anthony, C.D. (1995). ''Socioecology of a terrestrial salamander: juveniles enter adult territories during stressful foraging periods.'' Ecology, 76, 533-543.
- Jaeger, R.G., and Rubin, A.M. (1982). ''Foraging tactics of a terrestrial salamander judging prey profitability.'' Journal of Animal Ecology, 51, 167-176.
- Kleeberger, S.R. and Werner, J.K. (1982). ''Home range and homing behavior of Plethodon cinereus in northern Michigan.'' Copeia, 1982(2), 409-415.
- Lang, C. and R.G. Jaeger (2000). ''Defense of territories by male-female pairs in the Red-Backed Salamander (Plethodon cinereus).'' Copeia, 2000(1), 169-177.
- Lauer, A., Simon, M. A., Banning, J. L., André, E., Duncan, K., and Harris, R. N. (2007). ''Common cutaneous bacteria from the eastern red-backed salamander can inhibit pathogenic fungi.'' Copeia, 2007, 630-640.
- Maerz, J. C., Nuzzo, V. A., and Blossey, B. (2009). ''Declines in woodland salamander abundance associated with non-native earthworm and plant invasions.'' Conservation Biology, 23, 975-981.
- Maglia, A.M. (1996). ''Ontogeny and feeding ecology of the Red-Backed Salamander Plethodon cinereus.'' Copeia, 1996(3), 576-586.
- Mathis, A. (1990). ''Territoriality in a terrestrial salamander the influence of resource quality and body size.'' Behavior, 112, 162-175.
- Mitchell, J.C. and Woolcott, W.S. (1985). ''Observations of the microdistribution diet and predator-prey size relationships in the salamander Plethodon cinereus from the Virginia Piedmont.'' Virginia Journal of Science, 36, 281-288.
- Moore, J.-D., and Wyman, R. L. (2010). ''Eastern red-backed salamanders (Plethodon cinereus) in a highly acid forest soil.'' The American Midland Naturalist, 163, 95-105.
- Moreno, G. (1989). ''Behavioral and physiological differentiation between the color morphs of the salamander Plethodon cinereus.'' Journal of Herpetology, 23, 335-341.
- Ng, M.V. and Wilbur, H.M. (1995). ''The cost of brooding in Plethodon cinereus.'' Herpetologica, 51, 1-8.
- Ousterhout, B. H., and Liebgold, E. B. (2010). ''Dispersal versus site tenacity of adult and juvenile red-backed salamanders (Plethodon cinereus).'' Herpetologica, 66, 269-275.
- Petruzzi, E. E., Niewiarowski, P. H., and Moore, F. B.-G. (2006). ''The role of thermal niche selection in maintenance of a colour polymorphism in redback salamanders (Plethodon cinereus).'' Frontiers in Zoology, 3, 10, doi:10.1186/1742-9994-3-10.
- Rankin, J. S. (1937). ''An ecological study of parasites of some North Carolina salamanders.'' Ecological Monographs, 7, 169-269.
- Ransom, T. S. (2010). ''Earthworms, as ecosystem engineers, influence multiple aspects of a salamanders ecology.'' Oecologia, published online 17 September 2010, DOI: 10.1007/s00442-010-1775-1.
- Ransom, T. S., and Jaeger, R. G. (2006). ''An assemblage of salamanders in the southern Appalachian Mountains revisited: competitive and predatory behavior.'' Behaviour, 143, 1357-1382.
- Robertson, J. (2010). ''Mimicry: experimental design and scientific logic.'' An Introduction to Methods and Models in Ecology, Evolution, and Conservation Biology. S. Braude and B. S. Low, eds., Princeton University Press, Princeton, NJ.
- Simons, R.R., Gelgenhauer, B.E., and Jaeger, R.G. (1994). ''Salamander scent marks: site of production and their role in territorial defence.'' Animal Behavior, 48(11), 97-103.
- Simons, R.R., Jaeger, R.G., and Gelgenhauer, B.E. (1997). ''Competitor assessment and area defense by territorial salamanders.'' Copeia, 1997(1), 70-76.
- Takahashi, M. K. and Pauley, T. K. (2010). ''Resource allocation and life history traits of Plethodon cinereus at different elevations.'' The American Midland Naturalist, 163, 87-94.
- Tilley, S.G., Lundrigan, B.G., and Brower, L.P. (1982). ''Erythrism and mimicry in the salamander Plethodon cinereus.'' Herpetologica, 38, 409-417.
- Venesky, M. D., and Anthony, C. D. (2007). ''Antipredator adaptations and predator avoidance by two color morphs of the eastern red-backed salamander Plethodon cinereus.'' Herpetologica, 63, 450-458.
- Woodhams, D. C., Vredenburg, V. T., Simon, M. A., Billheimer, D., Shakhtour B., et al. (2007). ''Symbiotic bacteria contribute to innate immune defenses of the threatened mountain yellow-legged frog, Rana muscosa.'' Biological Conservation, 138, 390-398.
