Plethodontidae

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Lungless salamander
Temporal range: Oligocene–Present
Kaldari Batrachoseps attenuatus 02.jpg
California slender salamander (Batrachoseps attenuatus)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Urodela
Suborder: Salamandroidea
Family: Plethodontidae
Gray, 1850
Subgroups
DIstribution of Plethodontidae.png
Native distribution of plethodontids (in green)

Plethodontidae, or lungless salamanders, are a family of salamanders. [1] [2] With over 500 species, lungless salamanders are by far the largest family of salamanders in terms of their diversity. Most species are native to the Western Hemisphere, from British Columbia to Brazil. Only two extant genera occur in the Eastern Hemisphere: Speleomantes (native to Sardinia and mainland Europe south of the Alps) and Karsenia (native to South Korea). [3]

Contents

Biology

Palaeoplethodon hispaniolae preserved in amber Salamander in amber.jpg
Palaeoplethodon hispaniolae preserved in amber

Adult lungless salamanders have four limbs, with four toes on the fore limbs, and usually with five on the hind limbs. Within many species, mating and reproduction occur solely on land. Accordingly, many species also lack an aquatic larval stage, a phenomenon known as direct development in which the offspring hatch as fully-formed, miniature adults. Direct development is correlated with changes in the developmental characteristics of plethodontids compared to other families of salamanders including increases in egg size and duration of embryonic development. Additionally, the evolutionary loss of the aquatic larval stage is related to a diminishing dependence on aquatic habitats for reproduction. The lift of this constraint allowed widespread colonization and diversification within a broad number of terrestrial habitats which is a testament to the high success and proliferation of Plethodontidae. [4]

Despite the absence of lungs, some can grow rather large. The largest species of lungless salamanders, Bell's false brook salamander, can reach lengths of 36 cm (14 in). [5]

Many species have a projectile tongue and hyoid apparatus, which they can fire almost a body length at high speed to capture prey.

Measured in individual numbers, they are very successful animals where they occur. In some places, they make up the dominant biomass of vertebrates. [6] An estimated 1.88 billion individuals of the southern redback salamander inhabit just one district of Mark Twain National Forest alone, about 1,400 tons of biomass. [7] Due to their modest size and low metabolism, they are able to feed on prey such as springtails, which are usually too small for other terrestrial vertebrates. This gives them access to a whole ecological niche with minimal competition from other groups.

Courtship and mating

Plethodontids exhibit highly stereotyped and complex mating behaviors and courtship rituals that are not present in any other salamander family. Mating behavior tends to be uniform among all plethodontids and typically involves a tail-straddle walk in which the female orients her head at the base of the male's tail while also straddling the tail with her body. The male will twist his body around and deposit a sperm capsule, known as the spermatophore, on the substrate in front of the female's snout. As the male leads the female over the spermatophore with his tail, the female lowers her cloaca onto the spermatophore and lodges the sperm mass inside while leaving the base of the spermatophore on the ground. [8]

Within many species of plethodontidae, the courtship ritual is often accompanied by transfer of male pheromones during the tail-straddling walk. During the breeding period, males will grow enlarged anterior teeth used to scratch the female's skin on her head as a part of the courtship ritual. Subsequently, the male will rub pheromones onto the abraded spot which are secreted from a pad of tissue called the mental gland located underneath the male's chin. [8] [9] [10] [11]

Courtship pheromones greatly increase male mating success for a variety of reasons. Overall, the pheromone secretions increase female receptivity to courtship and sperm transfer. This not only increases the likelihood of successful mating with a specific female, but also shortens the duration of courtship which is important because it minimizes the chance of the male being interrupted by other competing males. [12]

In scientific literature discussing the variations between the mental glands of plethodontid salamanders, it was discovered that male plethodontids had minor variations in height  and diameter of the simple tubular glands, and major variation was found in the diameter of the secretory granules. This is attributed to the fact that males can mate throughout all months of the year, while females oviposit seasonally.

Respiration

A number of features distinguish the plethodontids from other salamanders. Most significantly, they lack lungs, conducting respiration through their skin, and the tissues lining their mouths. [3] Some species of cave salamanders are neotenic, and keep their larval gills even as adults. Gills are absent in all other adult plethodontids. [13]

Plethodontid salamanders are almost entirely reliant on cutaneous respiration. [14] Approximately 83%–93% of oxygen uptake is through this method. [15] Plethodontid salamander respiration rates are constrained by their SA:V, and higher SA:Vs are correlated to warmer, wetter climates. [16]

Plethodontids are constantly exposed to air or water, which allows for constant gas exchange that is not limited by ventilation. [14] Oxygen uptake is identical in water and air, assuming the partial pressure of oxygen is the same. [17] Oxygenated and non-oxygenated blood are mixed together in the venous system, which causes the partial pressure of oxygen within cardiac blood to typically be low. [14]

Plethodontids can tolerate hypoxia for prolonged periods by reducing their metabolic rate instead of by relying on anaerobic cutaneous respiration, as initially theorized. [17]

Plethodontids have been observed to develop rudimentary lungs as embryos. [18] The lung rudiment develops similarly to that of non-plethodontid salamanders for the first three weeks of development and then begins to regress through apoptosis. [18]

The nasiolabal groove of a dusky salamander Nasiolabial groove of Desmognathus fuscus.jpg
The nasiolabal groove of a dusky salamander

Chemoreception

Another distinctive feature is the presence of a vertical slit between the nostril and upper lip, known as the "nasolabial groove". The groove is lined with glands, and enhances the salamander's chemoreception which is correlated with a higher degree of olfactory lobe and nasal mucous membrane development in plethodontids. [3] [19] The presence of this specialized structure is likely related to the absence of lungs in these salamanders. Though some lunged salamanders do exhibit similar structures, they are reduced in size and are not arranged near the nostrils (i.e. nares) in the same fashion as plethodontids. Due to the fact that plethodontids cannot generate air pressure via expulsion of air from the lungs and through the nares, they are presented with the challenge of removing water and debris from the nasal passages which has the potential to significantly limit olfactory processes. As such, the nasolabial grooves are structured in a way that maximizes drainage from the nose. The groove is deeper and more narrow directly around the nares and the orifices of the glands are slightly elevated both of which aid in the gravitational flow of fluid from the nares and nasal depression. Additionally, the nasolabial glands around the margins of the nares secrete a fatty film which further encourages the removal of water from the nasal passages due to differences in polarity between water and the lipid secretions. [19]

Evolutionary history

Plethodontidae are estimated to have split from their sister group Amphiumidae around the K-Pg boundary, and to have diversified during the Paleogene. [20] The origin region of the family is North America, with oldest of the European members of the family known from the Middle Miocene of Slovakia. [21]

Subfamilies and genera

The family Plethodontidae consists of two extant subfamilies and about 516 [2] to 520 [1] species divided among these genera, making up the majority of known salamander species: [2]

SubfamilyGenus, scientific name, and authorCommon nameSpecies
Hemidactyliinae
Hallowell, 1856
Aquiloeurycea Rovito, Parra-Olea, Recuero, and Wake, 2015
6 - 7
Batrachoseps Bonaparte, 1839Slender salamanders
23
Bolitoglossa Duméril, Bibron & Duméril, 1854Tropical climbing salamanders
139 - 140
Bradytriton Wake & Elias, 1983Finca Chiblac salamander
1
Chiropterotriton Taylor, 1944Splay-foot salamanders
23
Cryptotriton García-París & Wake, 2000Hidden salamanders
7
Dendrotriton Wake & Elias, 1983Bromeliad salamanders
8
Eurycea Rafinesque, 1822North American brook salamanders
34
Gyrinophilus Cope, 1869Spring salamanders
4
Hemidactylium Tschudi, 1838Four-toed salamanders
1
Isthmura Dubois and Raffaelli, 2012
7
Ixalotriton Wake and Johnson, 1989Jumping salamanders
2
Nototriton Wake & Elias, 1983Moss salamanders
22
Nyctanolis Elias & Wake, 1983Long-limbed salamanders
1
Oedipina Keferstein, 1868Worm salamanders
40
Parvimolge Taylor, 1944Tropical dwarf salamanders
1
Pseudoeurycea Taylor, 1944False brook salamanders
41
Pseudotriton Tschudi, 1838Mud and red salamanders
2 - 4
Stereochilus Cope, 1869Many-lined salamanders
1
Thorius Cope, 1869Minute salamanders
29
Urspelerpes Camp, Peterman, Milanovich, Lamb, Maerz, and Wake, 2009Patch-nosed salamanders
1
Plethodontinae
Gray, 1850
Aneides Baird, 1851Climbing salamanders
10
Desmognathus Baird, 1850Dusky salamanders
39
Ensatina Gray, 1850Ensatinas
1
Hydromantes Gistel, 1848Web-toed salamanders
5
Karsenia Min, Yang, Bonett, Vieites, Brandon & Wake, 2005Korean crevice salamanders
1
Phaeognathus Highton, 1961Red Hills salamanders
1
Plethodon Tschudi, 1838Slimy and mountain salamanders
58
Speleomantes Dubois, 1984European cave salamanders
8

Following a major revision in 2006, the genus Haideotriton was found to be a synonym of Eurycea , while the genus Lineatriton were made synonyms of Pseudoeurycea . [22]

A single hemidactyliine ( Palaeoplethodon ) is known from Miocene fossil remains preserved in Dominican amber, marking the only record of salamanders in the Caribbean. [23]

Conservation status

Conservation Status of Plethodontidae According to IUCN Redlist (2020)
StatusNumber of Species
Least Concern94
Near Threatened39
Vulnerable60
Endangered88
Critically Endangered68
Extinct1
Data Deficient40

[24]

Related Research Articles

<span class="mw-page-title-main">Salamander</span> Order of amphibians

Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten extant salamander families are grouped together under the order Urodela from the group Caudata. Salamander diversity is highest in eastern North America, especially in the Appalachian Mountains; most species are found in the Holarctic realm, with some species present in the Neotropical realm.

<i>Amphiuma</i> Genus of amphibians

Amphiuma is a genus of aquatic salamanders from the United States, the only extant genus within the family Amphiumidae. They are colloquially known as amphiumas. They are also known to fishermen as "conger eels" or "Congo snakes", which are zoologically incorrect designations or misnomers, since amphiumas are actually salamanders, and not fish, nor reptiles and are not from Congo. Amphiuma exhibits one of the largest complements of DNA in the living world, around 25 times more than a human.

<span class="mw-page-title-main">Salamandridae</span> Family of amphibians

Salamandridae is a family of salamanders consisting of true salamanders and newts. Salamandrids are distinguished from other salamanders by the lack of rib or costal grooves along the sides of their bodies and by their rough skin. Their skin is very granular because of the number of poison glands. They also lack nasolabial grooves. Most species of Salamandridae have moveable eyelids but lack lacrimal glands.

<span class="mw-page-title-main">Ambystomatidae</span> Family of amphibians

Ambystomatidae is a family of salamanders belonging to the Suborder Salamandroidea in the class Amphibia. It contains two genera, Ambystoma and Dicamptodon. Ambystoma contains 32 species and are distributed widely across North America, while Dicamptodon contains four species restricted to the Pacific Northwest. These salamanders are mostly terrestrial and eat invertebrates, although some species are known to eat smaller salamanders. They can be found throughout the US and some areas of Canada in damp forests or plains. This family contains some of the largest terrestrial salamanders in the world, the tiger salamander and the coastal giant salamander. Some species are toxic and can secrete poison from their bodies as protection against predators or infraspecific competition. Neoteny has been observed in several species in Ambystomatidae, and some of them like the axolotl live all of their lives under water in their larval stage.

<span class="mw-page-title-main">Texas blind salamander</span> Species of amphibian

The Texas blind salamander is a rare and endangered cave-dwelling troglobite amphibian native to San Marcos, Hays County, Texas, specifically the San Marcos Pool of the Edwards Aquifer. This species resembles the olm, another stygofaunal salamander from Europe. Unlike the olm, this amphibian's body is not as elongated, and also has less reduced digits on its limbs.

<i>Bolitoglossa</i> Genus of amphibians

Bolitoglossa is a genus of lungless salamanders, also called mushroom-tongued salamanders, tropical climbing salamanders, or web-footed salamanders, in the family Plethodontidae. Their range is between northern Mexico through Central America to Colombia, Venezuela, Ecuador, Peru, northeastern Brazil, and central Bolivia. Neotropical salamanders of the Bolitoglossa make up the largest genus in the order Caudata, consisting of approximately one-fifth of all known species of salamanders. Adult salamanders range anywhere from 45mm to 200mm in length depending on their specific species. They are notorious for their ability to project their tongue at prey items, as indicated from their name. They are also known for their webbed feet, having significantly more webbing than any other species outside their genus with the exception of the cave-dwelling Mexican bolitoglossine Chiropterotriton magnipes. Although webbed feet are a common characteristic of these salamanders, only about half of the species in this genus contain webbed feet.

The northern banana salamander, also known as common dwarf salamander or rufescent salamander, is a species of salamander in the family Plethodontidae. It is found in the Atlantic slopes of Meso-America from San Luis Potosi, Veracruz, and northern Chiapas in Mexico continuing on to the southern part of Guatemala, Belize, and northern Honduras. However, its range south of Mexico is uncertain because the records may refer to other species.

<span class="mw-page-title-main">Seepage salamander</span> Species of amphibian

The seepage salamander is a small, terrestrial species of salamander in the family Plethodontidae. It is endemic to the United States. They are found in small areas of Tennessee, North Carolina, Georgia, and Alabama. Its natural habitats are temperate forests, intermittent rivers, and freshwater springs. It gets its name from the seepages around which it lives. It is very similar in its appearance and life history to the pygmy salamander. These two species differ greatly from the other Desmognathus species. They are the smallest salamanders in the genus, measuring only 3–5 cm (1–2 in) in length. They are also the only two terrestrial, direct-developing Desmognathus species. However, the two species are not often seen to coexist, differing in distribution by elevation; although there are exceptions. The seepage salamander is currently listed as Near Threatened, with its numbers declining in most of states in which it is found. It is threatened by habitat loss, with logging having a major effect.

<i>Desmognathus fuscus</i> Species of amphibian

Desmognathus fuscus is a species of amphibian in the family Plethodontidae. The species is commonly called the dusky salamander or northern dusky salamander to distinguish it from populations in the southern United States which form several distinct species, the southern dusky salamanders. The northern dusky salamander is the most widespread representative of its genus in Canada. It can be found in eastern North America from extreme eastern Canada in New Brunswick south to South Carolina. The size of the species' total population is unknown, but is assumed to easily exceed 100,000. The species' habitat differs somewhat geographically; dusky salamanders in the northern part of the range prefer rocky woodland streams, seepages, and springs, while those in the south favor floodplains, sloughs, and muddy places along upland streams. They are most common where water is running or trickling. They hide under various objects, such as leaves or rocks, either in or near water. Alternatively, they may enter burrows for protection. The dusky salamander lays its eggs close to water under moss or rocks, in logs, or in stream-bank cavities. The larval stage which follows is normally aquatic.

<span class="mw-page-title-main">Ocoee salamander</span> Species of amphibian

The ocoee salamander is a species of salamander in the family Plethodontidae. This salamander has a variety of colors and patterns, and got its name from Tennessee state wildflower. Its natural habitats are temperate forests, rivers, intermittent rivers, freshwater springs and wet rocks in mountainous areas of the Southeastern United States. It was first described by Nicholls in 1949. They are territorial and feed on small invertebrates. It is widely distributed in the southeastern United States and is listed as "Least Concern" by the International Union for Conservation of Nature.

<span class="mw-page-title-main">Northern two-lined salamander</span> Species of amphibian

The northern two-lined salamander is a species of salamander in the family Plethodontidae found in Canada and the United States. Its natural habitats are temperate forests, temperate shrubland, rivers, intermittent rivers, freshwater marshes, freshwater springs, arable land, and urban areas. It is more water-oriented than the related northern redback salamander, and can often be found in and around water such as rain puddles, streams, swamps, and damp stream beds, whereas the northern redback tends to be found in damp ground, but usually not near open water.

<span class="mw-page-title-main">Chamberlain's dwarf salamander</span> Species of amphibian

Chamberlain's dwarf salamander is a species of salamander in the family Plethodontidae, endemic to the United States. It is only found in the states of North Carolina and South Carolina. It was previously thought to be a color morph of the southeastern dwarf salamander, but was described in 2003 as a distinct species based on distinct morphology and behavior. A 2017 study reaffirmed it as being a distinct species. It is estimated that E. chamberlaini diverged from E. quadridigitata anywhere from 27 to 15 million years ago.

<span class="mw-page-title-main">Blue Ridge two-lined salamander</span> Species of amphibian

The Blue Ridge two-lined salamander is a species of salamander in the family Plethodontidae, endemic to the United States. This species is found in the southern Appalachian Mountains, mostly south of Virginia. To the north is a similar salamander, Eurycea bislineata, or the northern two-lined salamander. Its genus, Eurycea contains 33 species and includes taxa that have either a metamorphic life cycle or larval-form paedomorphosis. In species that metamorphose, there can be within-and among-population variation in larval life-history characteristics, e.g., duration of the larval period and size at metamorphosis. Intraspecific geographic variation in species of Eurycea has been attributed to several factors: temperature, stream order and productivity of the larval habitat.

<span class="mw-page-title-main">Spring salamander</span> Species of amphibian

The spring salamander is a species of salamander in the family Plethodontidae. It is found in Canada and the United States. The genus, Gyrinophilus, means "tadpole lover" and refers to the long period of time it spends as a gilled larva before maturing. The specific epithet, porphyriticus, is Latin from Greek, meaning the color of porphyry, a purple stone, and this salamander has also been called the purple salamander.

<span class="mw-page-title-main">Red salamander</span> Species of amphibian

The red salamander is a species of salamander in the family [[Plethodontidae) endemic to the eastern United States. Its skin is orange/red with random black spots. Its habitats are temperate forests, small creeks, ponds, forests, temperate shrubland, rivers, intermittent rivers, freshwater, trees springs. Overall this species is common and widespread, but locally it has declined because of habitat loss and it is considered threatened in Indiana and New York. Red salamanders eat insects, earthworms, spiders, small crustaceans, snails, and smaller salamanders. To eat, they extend their tongue to capture prey on the tip of it and retract it back into their mouths. The red salamander, as a member of the family Plethodontidae, lacks lungs and respires through its skin.

<i>Speleomantes</i> Genus of amphibians

Speleomantes, or European cave salamanders, are a genus of salamander in the family Plethodontidae, or lungless salamanders. It is one of two genera in the family to inhabit the Old World, with the remaining 250 or so species being found in North, Central and South America. The genus is endemic to Italy and a few nearby areas.

Thorius aureus, the golden thorius, is a species of salamander in the genus Thorius, the Mexican pigmy salamanders, part of the lungless salamander family. It is endemic to mountainous areas of north central Oaxaca State in Mexico. It is one of the largest Thorius species.

<span class="mw-page-title-main">Sexual selection in amphibians</span> Choice of and competition for mates

Sexual selection in amphibians involves sexual selection processes in amphibians, including frogs, salamanders and newts. Prolonged breeders, the majority of frog species, have breeding seasons at regular intervals where male-male competition occurs with males arriving at the waters edge first in large number and producing a wide range of vocalizations, with variations in depth of calls the speed of calls and other complex behaviours to attract mates. The fittest males will have the deepest croaks and the best territories, with females making their mate choices at least partly based on the males depth of croaking. This has led to sexual dimorphism, with females being larger than males in 90% of species, males in 10% and males fighting for groups of females.

The Ocmulgee slimy salamander is a species of salamander in the family Plethodontidae. It is endemic to the state of Georgia in the United States, where it is found in regions of the coastal plain and Piedmont that are associated with the Ocmulgee River drainage system. It is only known from a few counties, and due to this restricted range, it is at high risk of extinction. Many populations of this species are already experiencing precipitous declines, with some even possibly being extirpated.

A mental gland is a part of the body found in many species of amphibians and reptiles. Mental glands produce chemicals that conspecific animals use to communicate.

References

  1. 1 2 Frost, Darrel R. (2024). "Plethodontidae Gray, 1850". Amphibian Species of the World: An Online Reference. Version 6.2. American Museum of Natural History. doi:10.5531/db.vz.0001 . Retrieved 4 March 2024.
  2. 1 2 3 "Plethodontidae". AmphibiaWeb. University of California, Berkeley. 2024. Retrieved 4 March 2024.
  3. 1 2 3 Lanza, B.; Vanni, S.; Nistri, A. (1998). Cogger, H.G.; Zweifel, R.G. (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. pp. 74–75. ISBN   0-12-178560-2.
  4. ↑ Lewis, Zachary R.; Kerney, Ryan; Hanken, James (2011). "Lung development in lungless salamanders!". Developmental Biology. 356 (1): 250–251. doi: 10.1016/j.ydbio.2011.05.560 .
  5. ↑ Bell's False Brook Salamander - World Land Trust
  6. ↑ Hairston, N.G., Sr. 1987. Community ecology and salamander guilds. Cambridge University Press. Cambridge.
  7. ↑ "Salamanders a more abundant food source in forest ecosystems than previously thought". ScienceDaily. 18 November 2014.
  8. 1 2 Sever, David M. (2003). Reproductive biology and phylogeny of Urodela. Enfield, NH: Science Publishers. pp. 383–424. ISBN   978-1-57808-645-0. OCLC   427511083.
  9. ↑ Sever, David M.; Dustin S. Siegel; Michael S. Taylor; Christopher K. Beachy1 (March 17, 2016). "Phylogeny of Mental Glands, Revisited". Copeia. 104 (1): 83–93. doi:10.1643/CH-14-210. PMC   6054469 . PMID   30034038.{{cite journal}}: CS1 maint: numeric names: authors list (link)
  10. ↑ David M. Sever (January 18, 2016). "Ultrastructure of the mental gland of the Red-Backed Salamander, Plethodon cinereus (Amphibia: Plethodontidae)". Acta Zoologica. 98 (2): 154–162. doi:10.1111/azo.12158.
  11. ↑ Ariana E. Rupp; David M. Sever (February 14, 2017). "Histology of mental and caudal courtship glands in three genera of plethodontid salamanders (Amphibia: Plethodontidae)" (PDF). Acta Zoologica. 98 (2): 154–162. doi:10.1111/azo.12188 . Retrieved July 5, 2022.
  12. ↑ Houck, Lynne D.; Reagan, Nancy L. (1990). "Male courtship pheromones increase female receptivity in a plethodontid salamander". Animal Behaviour. 39 (4): 729–734. doi:10.1016/s0003-3472(05)80384-7. S2CID   53185123.
  13. ↑ Holman, J. Alan (2006). Fossil Salamanders of North America. Indiana University Press. ISBN   0253347327.
  14. 1 2 3 Gatz, Randall N.; Crawford, Eugene C.; Piiper, Johannes (1974-02-01). "Respiratory properties of the blood of a lungless and gill-less salamander, Desmognathus fuscus". Respiration Physiology. 20 (1): 33–41. doi:10.1016/0034-5687(74)90016-4. PMID   4821655.
  15. ↑ Whitford, Walter G. (1973). "The effects of temperature on respiration in the Amphibia". American Zoologist. 13 (2): 505–512. doi:10.1093/icb/13.2.505.
  16. ↑ Baken, Erica K.; Mellenthin, Lauren E.; Adams, Dean C. (2020). "Macroevolution of desiccation‐related morphology in plethodontid salamanders as inferred from a novel surface area to volume ratio estimation approach". Evolution. 74 (2): 476–486. doi: 10.1111/evo.13898 . PMID   31849047. S2CID   209407983.
  17. 1 2 Gatz, Randall N.; Crawford, Eugene C.; Piiper, Johannes (1974-02-01). "Metabolic and heart rate response of the plethodontid salamander Desmognathus fuscus to hypoxia". Respiration Physiology. 20 (1): 43–49. doi:10.1016/0034-5687(74)90017-6. PMID   4821656.
  18. 1 2 Lewis, Zachary R.; Kerney, Ryan; Hanken, James (2022-08-19). "Developmental basis of evolutionary lung loss in plethodontid salamanders". Science Advances. 8 (33): eabo6108. Bibcode:2022SciA....8O6108L. doi:10.1126/sciadv.abo6108. PMC   9385146 . PMID   35977024.
  19. 1 2 Brown, Charles E.; Martof, Bernard S. (1966-10-01). "The Function of the Naso-Labial Groove of Plethodontid Salamanders". Physiological Zoology. 39 (4): 357–367. doi:10.1086/physzool.39.4.30152358. S2CID   87787255.
  20. ↑ Shen, Xing-Xing; Liang, Dan; Chen, Meng-Yun; Mao, Rong-Li; Wake, David B.; Zhang, Peng (January 2016). "Enlarged Multilocus Data set Provides Surprisingly Younger Time of Origin for the Plethodontidae, the Largest Family of Salamanders". Systematic Biology. 65 (1): 66–81. doi: 10.1093/sysbio/syv061 . PMID   26385618.
  21. ↑ Sanchíz, Borja; Venczel, Márton (2005). "A fossil plethodontid salamander from the Middle Miocene of Slovakia (Caudata, Plethodontidae)". Amphibia-Reptilia. 26 (3): 408–411. doi: 10.1163/156853805774408586 .
  22. ↑ Frost, D. R.; Grant, T.; Faivovich, J. N.; Bain, R. H.; Haas, A.; Haddad, C. F. B.; De Sá, R. O.; Channing, A.; Wilkinson, M.; Donnellan, S. C.; Raxworthy, C. J.; Campbell, J. A.; Blotto, B. L.; Moler, P.; Drewes, R. C.; Nussbaum, R. A.; Lynch, J. D.; Green, D. M. & Wheeler, W. C. (2006). "The amphibian tree of life". Bulletin of the American Museum of Natural History. 297: 1–291. doi: 10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2 . hdl:2246/5781.
  23. ↑ Poinar Jr., G.; Wake, David B. (2015). "Palaeoplethodon hispaniolae gen. n., sp. n. (Amphibia: Caudata), a fossil salamander from the Caribbean" (PDF). Palaeodiversity. 8: 21–29.
  24. ↑ IUCN RedList, (2020). Plethodontidae. Retrieved from https://www.iucnredlist.org/search?taxonomies=101246&searchType=species
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