Red-backed salamander

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Red-backed salamander
Plethodon cinereus1.jpg
"Redback" phase
Plethodon cinereus.jpg
"Leadback" phase

Least Concern (IUCN 3.1)[1]
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Urodela
Family: Plethodontidae
Subfamily: Plethodontinae
Genus: Plethodon
Species:
P. cinereus
Binomial name
Plethodon cinereus
(Green, 1818)
Plethodon cinereus map.svg
Range of P. cinereus
Synonyms[2]
  • Salamandra cinerea Green, 1818
  • Plethodon cinereus — Tschudi, 1838

The red-backed salamander (Plethodon cinereus) is a small, hardy woodland salamander species in the family Plethodontidae. The species inhabits wooded slopes in eastern North America, west to Missouri, south to North Carolina, and north from southern Quebec and the Maritime provinces in Canada to Minnesota.[3] It is also known as the redback salamander,[4] eastern red-backed salamander,[4] or the northern red-backed salamander to distinguish it from the southern red-backed salamander (Plethodon serratus). It is one of 56 species in the genus Plethodon. Red-backed salamanders are notable for their color polymorphism and primarily display two color morph varieties ("red-backed" and "lead-backed"), which differ in physiology and anti-predator behavior.[5][6][7]

Description and ecology[]

Red-backed salamander in its habitat

The red-backed salamander is a small terrestrial salamander, 5.7–10.0 cm (2.2–3.9 in) in total length (including tail), which usually lives in forested areas under rocks, logs, bark, and other debris.[3] It is one of the most numerous salamanders throughout its range.[3] Red-backed salamanders are mostly insectivorous, but prey on a wide assortment of other small invertebrates including isopods, millipedes, centipedes, pseudoscorpions, harvestmen, spiders, and gastropods.[8] On one occasion an individual was found to have cannibalized a P. cinereus embryo.[8] Predators of the red-backed salamander are many and varied, and include small mammals, snakes, ground-foraging birds, and larger salamanders.[9]

As with all amphibians, the red-backed salamander has permeable skin. They also lack lungs, a condition which is an ancestral trait of the Plethodontidae.[10] Red-backed salamanders are thus entirely reliant on cutaneous respiration for gas exchange. Permeable skin is susceptible to desiccation and must be kept moist in order to facilitate cutaneous respiration; as a result much of the ecology and behavior of the red-backed salamander is restricted by climatic and microclimatic variables, particularly dryness and temperature.[11] Individuals confine themselves to moist microhabitats (beneath rocks, woody debris, etc. as well as beneath the soil) for long periods of time in order to maintain hydration when surface conditions are inhospitably dry or hot, and are only active on the surface to travel, forage, or reproduce for short periods.[11] The duration of surface activity is directly limited by the rate of cutaneous water loss to the environment, which is influenced by environmental variables such as altitude, forest canopy cover, and the amount of recent precipitation.[11][12] Hydric limitations are thus a major constraint on red-backed salamander growth and reproductive success as well as the genetic interconnectedness of red-backed salamander populations.[11][12][13]

The skin of red-backed salamanders was found to contain Lysobacter gummosus, an epibiotic bacterium that produces the chemical 2,4-diacetylphloroglucinol and inhibits the growth of certain pathogenic fungi.[14]

Polymorphism[]

The red-backed salamander exhibits color polymorphism and two color variations are common: the "red-backed" or "red-stripe" variety has a red dorsal stripe that tapers towards the tail, and the darker variety, known as the "lead-backed" (or simply "lead") phase, lacks most or all of the red pigmentation.[3] The red-backed phase is not always red, but may actually be various other colors (e.g., yellow-backed, orange-backed, white-backed, or a rare erythristic morph in which the body is completely red).[3] Both morphs have speckled black and white bellies.[3] Additional color anomalies of this species also exist, including iridistic, albino, leucistic, amelanistic, and melanistic anomalies.[15] These color morphs are rarer than the red-backed, lead-backed, and erythristic morphs, but still have been reported with consistency among varying populations of this species.[15]

Lead-backed phase Redback Salamander - Plethodon cinereus
Lead-backed phase Redback Salamander - Plethodon cinereus

P. cinereus color morph frequencies are correlated with climatic variables, suggesting habitat temperature and more broadly climate to be potential sources of selective pressure on P. cinereus polymorphism.[5][6] The red-backed form is found with greater frequency in colder regions at more northerly latitudes and easterly longitudes throughout its range, whereas the opposite is true of the lead-backed form.[5][6] Additionally, lead-backed morphs withdraw from surface activity earlier in the autumn than red-backed morphs, presumably to avoid cooling temperatures.[5][16][17] Standard metabolic rate has also been found to differ between the morphs at certain temperatures, with significantly lower metabolic rates being displayed by the lead-backed form at 15 °C;[16] in the same study, lead-backed individuals were also more active on the ground surface at this temperature.[16] These findings suggest that the lead-backed color variant is less tolerant of cool temperatures than the red-backed color variant, and that the two color forms differ physiologically and behaviorally at certain temperatures.[5][6][16][17]

An alternate explanation for the uneven geographic distribution of the red-backed and lead-backed P. cinereus color morphs involves phenotypic plasticity responding to developmental temperature. Although the genetic origins of the P. cinereus polymorphic condition are not fully understood, initial studies indicate that color morph dominance is likely subject to epistasis, and that multiple loci may interact to determine an individual's morph condition.[18][19] However, more recent research indicates that a plastic response to thermal conditions during development also contributes to color morph determination; in one study, P. cinereus eggs incubated at a higher temperature hatched a greater proportion of lead-backed morphs than eggs incubated at a lower temperature.[20] Temperature-dependent color morph determination may therefore also potentially influence the spatial distribution of P. cinereus color morphs.[20]

The two primary P. cinereus color morphs also differ in diet.[17][21] The prevalence of certain prey taxa and the overall diversity and quality of prey items have been observed to differ seasonally between the two morphs in the spring and autumn when surface activity is greatest.[17][21] In one study, the autumn diet of red-backed morphs was more diverse and of higher quality, and found to be dominated by mites, springtails, and ants, whereas the most important prey for lead-backed morphs were ants, mites, and isopods.[17]

Lead-backed phase

Behavior[]

Antipredator behavior of P. cinereus was found to differ between the two color phases; the lead-backed phase has a tendency to run away from predators, whereas the red-backed phase often stays immobile and possibly exhibits aposematic coloration.[7] Stress levels of each color phase were estimated by determining the ratio of neutrophil to lymphocyte cells in the blood, and the results suggest stress levels are higher in the lead-backed phase than in the red-backed phase.[22] This may be a consequence of a higher predation risk experienced in the wild by the lead-backed phase, and may also mean lead-phase salamanders could be more vulnerable in captivity settings.[22]

Plethodon cinereus, like many plethodon species, exhibit homing behavior, with homing of females to their nests, as well as non-attending females and males to a home range.[23] This allows for essential contact between a female and her eggs in order to ensure their survival, as well as for non-attending females and males to explore beyond the home range when under predation pressures, or searching for food or cover, and return to their home range if a more favorable microhabitat is not found.[23]

Reproduction and biomass[]

Males and females of P. cinereus typically establish separate feeding and/or mating territories underneath rocks and logs. However, some red-backed salamanders are thought to engage in social monogamy, and may maintain co-defended territories throughout their active periods. Breeding occurs in June and July. Females produce from four to 17 eggs in a year. The eggs hatch in 6 to 8 weeks. Not much is known about the dispersal of neonates, although neonates and juveniles are thought to be philopatric. The species largely consumes invertebrates and other detritus dwellers. In some areas with good habitat, these salamanders are so numerous, their population densities may surpass 1,000 individuals per acre (2,471 individuals per hectare). Pokagon State Park in Indiana is one such place.[24]

As in many Plethodon species, female red-backed salamanders have the ability to store sperm as spermatophore, and have been evidenced in doing so up to eight months prior to the oviposition period in June and July.[25] Sperm or spermatophores are not retained following the oviposition period.[25]

References[]

  1. ^ IUCN SSC Amphibian Specialist Group (2015). "Plethodon cinereus". IUCN Red List of Threatened Species. 2015: e.T59334A78907687. doi:10.2305/IUCN.UK.2015-4.RLTS.T59334A78907687.en. Retrieved 19 November 2021.
  2. ^ Stejneger L, Barbour T (1917). A Check List of North American Amphibians and Reptiles. Cambridge, Massachusetts: Harvard University Press. 125 pp. (Plethodon cinereus, p. 15).
  3. ^ a b c d e f Conant R, Collins JT (1998). A Field Guide to Reptiles and Amphibians of Eastern and Central North America, Third Edition. Boston: Houghton Mifflin. 616 pp. ISBN 0-395-90452-8.
  4. ^ a b Integrated Taxonomic Information System [Internet] 2012. [updated 2012 Sept; cited 2012 Nov 26] Available from: www.itis.gov
  5. ^ a b c d e Lotter, Fred; Scott, Norman J. (1977). "Correlation between Climate and Distribution of the Color Morphs of the Salamander Plethodon cinereus". Copeia. 1977 (4): 681–690. doi:10.2307/1443166. ISSN 0045-8511. JSTOR 1443166.
  6. ^ a b c d Gibbs, James P.; Karraker, Nancy E. (2006). "Effects of Warming Conditions in Eastern North American Forests on Red-Backed Salamander Morphology". Conservation Biology. 20 (3): 913–917. doi:10.1111/j.1523-1739.2006.00375.x. ISSN 1523-1739. PMID 16909583.
  7. ^ a b Venesky, Matthew D.; Anthony, Carl D. (2007). "Antipredator adaptations and predator avoidance by two color morphs of the eastern red-backed salamander, Plethodon cinereus". Herpetologica. 63 (4): 450–458. doi:10.1655/0018-0831(2007)63[450:AAAPAB]2.0.CO;2.
  8. ^ a b Burton, Thomas M. (1976). "An Analysis of the Feeding Ecology of the Salamanders (Amphibia, Urodela) of the Hubbard Brook Experimental Forest, New Hampshire". Journal of Herpetology. 10 (3): 187–204. doi:10.2307/1562980. ISSN 0022-1511. JSTOR 1562980.
  9. ^ Grant, Alexa H.; Ransom, Tami S.; Liebgold, Eric B. (April 2018). "Differential Survival and the Effects of Predation on a Color Polymorphic Species, the Red-Backed Salamander (Plethodon cinereus)". Journal of Herpetology. 52 (2): 127–135. doi:10.1670/16-185. ISSN 0022-1511. S2CID 90207320.
  10. ^ Noble, G. Kingsley (1931). The biology of the amphibia (1st ed.). New York: McGraw-Hill. doi:10.5962/bhl.title.82448.
  11. ^ a b c d Feder, Martin E. (1983). "Integrating the Ecology and Physiology of Plethodontid Salamanders". Herpetologica. 39 (3): 291–310. ISSN 0018-0831. JSTOR 3892572.
  12. ^ a b Peterman, W. E.; Semlitsch, R. D. (2014-10-01). "Spatial variation in water loss predicts terrestrial salamander distribution and population dynamics". Oecologia. 176 (2): 357–369. Bibcode:2014Oecol.176..357P. doi:10.1007/s00442-014-3041-4. ISSN 1432-1939. PMID 25154754. S2CID 11041103.
  13. ^ Peterman, William E.; Semlitsch, Raymond D. (2013-05-06). "Fine-Scale Habitat Associations of a Terrestrial Salamander: The Role of Environmental Gradients and Implications for Population Dynamics". PLOS ONE. 8 (5): e62184. Bibcode:2013PLoSO...862184P. doi:10.1371/journal.pone.0062184. ISSN 1932-6203. PMC 3646024. PMID 23671586.
  14. ^ Brucker, Robert M.; Baylor, Cambria M.; Walters, Robert L.; Lauer, Antje; Harris, Reid N.; Minbiole, Kevin 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. 34 (1): 39–43. doi:10.1007/s10886-007-9352-8. PMID 18058176. S2CID 27149357.
  15. ^ a b Moore, Jean-David; Ouellet, Martin (2014-10-16). "A review of colour phenotypes of the Eastern Red-backed Salamander, Plethodon cinereus, in North America". The Canadian Field-Naturalist. 128 (3): 250–259. doi:10.22621/cfn.v128i3.1603. ISSN 0008-3550.
  16. ^ a b c d Moreno, Gabriel (1989). "Behavioral and Physiological Differentiation between the Color Morphs of the Salamander, Plethodon cinereus". Journal of Herpetology. 23 (4): 335–341. doi:10.2307/1564043. ISSN 0022-1511. JSTOR 1564043.
  17. ^ a b c d e Anthony, Carl D.; Venesky, Matthew D.; Hickerson, Cari-Ann M. (2008). "Ecological separation in a polymorphic terrestrial salamander". Journal of Animal Ecology. 77 (4): 646–653. doi:10.1111/j.1365-2656.2008.01398.x. ISSN 1365-2656. PMID 18479343.
  18. ^ Highton, Richard (1959). "The Inheritance of the Color Phases of Plethodon cinereus". Copeia. 1959 (1): 33–37. doi:10.2307/1440097. ISSN 0045-8511. JSTOR 1440097.
  19. ^ Highton, Richard (1975-06-25). "Geographic Variation in Genetic Dominance of the Color Morphs of the Red-Backed Salamander, Plethodon Cinereus". Genetics. 80 (2): 363–374. doi:10.1093/genetics/80.2.363. ISSN 1943-2631. PMC 1213333. PMID 17248684.
  20. ^ a b Evans, Annette E.; Urban, Mark C.; Jockusch, Elizabeth L. (2020-04-01). "Developmental temperature influences color polymorphism but not hatchling size in a woodland salamander". Oecologia. 192 (4): 909–918. Bibcode:2020Oecol.192..909E. doi:10.1007/s00442-020-04630-y. ISSN 1432-1939. PMID 32162072. S2CID 212669210.
  21. ^ a b Stuczka, Angela; Hickerson, Cari-Ann; Anthony, Carl (2016-01-01). "Niche partitioning along the diet axis in a colour polymorphic population of Eastern Red-backed Salamanders, Plethodon cinereus". Amphibia-Reptilia. 37 (3): 283–290. doi:10.1163/15685381-00003055. ISSN 1568-5381.
  22. ^ a b Davis AK, Milanovich JR (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 (2): 238–243.
  23. ^ a b Kleeberger, Steven R.; Werner, J. Kirwin (1982). "Home range and homing behavior of Plethodon cinereus in northern Michigan". Copeia. 1982 (2): 409–415. doi:10.2307/1444622. ISSN 0045-8511. JSTOR 1444622.
  24. ^ APPALACHIAN NATURE: An Entrée of Salamanders
  25. ^ a b Sayler, Anne (1966). "The reproductive ecology of the red-backed salamander, Plethodon cinereus, in Maryland". Copeia. 1966 (2): 183–193. doi:10.2307/1441125. ISSN 0045-8511. JSTOR 1441125.

Further reading[]

  • Behler JL, (1979). The Audubon Society Field Guide to North American Reptiles and Amphibians. New York: Alfred A. Knopf. 743 pp., 657 color plates. ISBN 0-394-50824-6. (Plethodon cinereus, pp. 336–337 + Plates 71, 117).
  • Green R (1818). "Descriptions of several species of North American Amphibia, accompanied with observations". Journal of the Academy of Natural Sciences of Philadelphia 1 (2): 348-359. (Salamandra cinerea, new species, pp. 356–357). (in English and Latin).
  • Powell R, Conant R, Collins JT (2016). Peterson Field Guide to Reptiles and Amphibians of Eastern North America, Fourth Edition. Boston and New York: Houghton Mifflin Harcourt. 494 pp., 47 color plates, 207 Figures. ISBN 978-0-544-12997-9. (Plethodon cinereus, pp. 78–80 + Plate 5 + Figures 33, 36, 37).
  • Zim HS, Smith HM (1956). Reptiles and Amphibians: A Guide to Familiar American Species: A Golden Nature Guide. Revised Edition. New York: Simon and Schuster. 160 pp. (Plethodon cinereus, pp. 147, 157).

External links[]

Data related to Plethodon cinereus at Wikispecies

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