# Mole salamander

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The **mole salamanders** ([genus](/source/Genus) ***Ambystoma***) are a group of [advanced salamanders](/source/Salamandroidea) endemic to North America. The group has become famous due to the study of the [axolotl](/source/Axolotl) (*A. mexicanum*) in research on [paedomorphosis](/source/Neoteny), and the [tiger salamander](/source/Tiger_salamander) (*A. tigrinum, A. mavortium*) which is often sold as a pet, and is the [official amphibian of four US states](/source/List_of_U.S._state_amphibians).

## General description

[Terrestrial](/source/Terrestrial_animal) mole salamanders are identified by having wide, protruding eyes, prominent [costal grooves](/source/Costal_groove), and thick arms. Most have vivid patterning on dark backgrounds, with marks ranging from deep blue spots to large yellow bars depending on the species. Terrestrial adults spend most of their lives underground in [burrows](/source/Burrow), either of their own making or abandoned by other animals. Some northern species may [hibernate](/source/Hibernation) in these [burrows](/source/Burrow) throughout the winter. They live alone and feed on any available [invertebrate](/source/Invertebrate). Adults spend little time in the water, only returning to the ponds of their birth to breed.

All mole salamanders are [oviparous](/source/Oviparous) and lay large eggs in clumps in the water. Their fully aquatic [larvae](/source/Larva) are branchiate, with three pairs of external [gills](/source/Gills) behind their heads and above their [gill slits](/source/Gill_slit). Larvae have large [caudal](/source/Caudal_(anatomical_term)) [fins](/source/Fin), which extend from the back of their heads to their tails and to their [cloacae](/source/Cloaca). Larvae grow limbs soon after hatching, with four toes on the fore arms, and five toes on the hind legs. Their eyes are wide-set and lack true eyelids.

The larvae of some species (especially those in the south, and tiger salamanders) can reach their adult size before undergoing [metamorphosis](/source/Metamorphosis_(biology)). During metamorphosis, the gills of the larvae disappear, as do the fins. Their tails, skin, and limbs become thicker, and the eyes develop lids. Their lungs become fully developed, allowing for a fully terrestrial existence.

Some species of mole salamanders (as well as populations of normally terrestrial species) are [neotenic](/source/Neoteny) (retaining their larval form into adulthood). The most famous example is the [axolotl](/source/Axolotl). They cannot produce [thyroxine](/source/Thyroxine), so their only means of metamorphosis is mainly through the outside injection of it. This usually shortens the lifespan of the salamander.

## Tiger salamander complex

Morphologically, tiger salamanders (*Ambystoma tigrinum* complex) have large heads, small eyes, and thick bodies.[1] This basic morphology is similar across most mole salamanders (genus *Ambystoma*), though tiger salamanders are among the largest of the mole salamanders,[2] and have relatively large larvae.

Tiger salamanders inhabit a wide variety of ecosystems across North America. Given this geographic diversity, subpopulations of tiger salamanders exhibit morphological and behavioral diversity.[3] Whether subpopulations constitute independent species or subspecies within the *Ambystoma tigrinum* complex, as well as the driving forces behind diversification, remains an active area of research as of 2024. Several subspecies within the *Ambystoma tigrinum* complex have been reclassified as an independent species. For example:

1. *Ambystoma mavortium* ([barred tiger salamander](/source/Barred_tiger_salamander)) comprises former subspecies *A. t. diaboli*, *A. t. mavortium*, *A. t. melanostictum*, *A. t. nebulosum*, and *A. t. stebbinsi.*[4][5][6]
1. *Ambystoma californiense* ([California tiger salamander](/source/California_tiger_salamander))
1. *Ambystoma velasci* ([Plateau tiger salamander](/source/Plateau_tiger_salamander)),[7] which may be paraphyletic and shares habitats with [axolotl](/source/Axolotl) (*A. mexicanum*)

## Hybrid all-female populations

Unisexual (all-female) populations of ambystomatid salamanders are widely distributed across the Great Lakes region and northeastern North America. The females require sperm from a co‑occurring, related species to fertilize their eggs and initiate development. Usually[8] the eggs then discard the sperm genome and develop asexually (i.e., [gynogenesis](/source/Gynogenesis), with [premeiotic doubling](/source/Parthenogenesis#Types_and_mechanisms)); however, they may incorporate the genome from the sperm into the resulting offspring.[9] Sperm incorporation commonly[8] takes the form of genome addition (resulting in [ploidy](/source/Ploidy) elevation in the offspring), or genome replacement, wherein one of the maternal genomes is discarded. This unique mode of reproduction has been termed [kleptogenesis](/source/Kleptogenesis) by Bogart and colleagues.[9] This is in contrast to [hybridogenesis](/source/Hybridogenesis), where the maternal genomes are passed hemiclonally and the paternal genome is discarded every generation before the egg matures and reacquired from the sperm of another species.

The nuclear DNA of the unisexuals generally comprises genomes from up to five species:[10] the [blue-spotted salamander](/source/Blue-spotted_salamander) (*A. laterale*), [Jefferson salamander](/source/Jefferson_salamander) (*A. jeffersonianum*), [small-mouthed salamander](/source/Small-mouthed_salamander) (*A. texanum*), [streamside salamander](/source/Streamside_salamander) (*A. barbouri*), and tiger salamander (*A. tigrinum*), denoted respectively as L, J, Tx, B, and Ti. This flexibility results in a large number of possible nuclear biotypes (genome combinations) in the unisexuals. For example, an LJJ individual would be a triploid with one *A. laterale* genome and two *A. jeffersonianum* genomes, while an LTxJTi individual would be a tetraploid with genomes from four species. Because they have hybrid genomes, unisexual salamanders are a cryptic species with morphology similar to coexisting species. For example, LLJs look like blue-spotted salamanders and LJJs look like Jefferson salamanders. Silvery salamanders LJJ (*A. platineum*), Tremblay's salamanders LLJ (*A. tremblayi*), and Kelly's Island salamanders LTxTx and LTxTi (*A. nothagenes*) were initially described as species. Species names were later dropped for all unisexual salamanders because of the complexity of their genomes. The offspring of a single mother may have different genome complements;[9] for example, a single egg mass may have both LLJJ and LJJ larvae.

Despite the complexity of the nuclear genome, all unisexuals form a [monophyletic](/source/Monophyletic) group based on their [mitochondrial DNA](/source/Mitochondrial_DNA). The maternal ancestor of the unisexual ambystomatids was most closely related to the [streamside salamander](/source/Streamside_salamander), with the original hybridization likely occurring 2.4~3.9 million years ago,[9] making it the oldest known lineage of all-female vertebrates.[11] The hybridization was most probably with an *A. laterale*. All known unisexuals have at least one *A. laterale* genome[10] and this is thought to be essential for unisexuality. However, the *A. laterale* genome has been replaced several times, independently, in each of the lineages by matings with *A. laterale*.

## Limb regeneration

*Ambystoma mexicanum*, a [neotenic](/source/Neoteny) [salamander](/source/Salamander) with exceptional [regenerative capabilities](/source/Regeneration_(biology)) is one of the principal models for studying limb regeneration.[12] Limb regeneration involves the propagation of a mass of low differentiated and highly proliferative cells termed the [blastema](/source/Blastema).[12][13] During limb regeneration, blastema cells experience [DNA double-strand breaks](/source/DNA_damage_(naturally_occurring)) and thus require [homologous recombination](/source/Homologous_recombination), a form of [DNA repair](/source/DNA_repair) that deals with double-strand breaks.[13]

## Taxonomy

*Rhyacosiredon* was previously considered a separate genus within the family Ambystomatidae. However, cladistic analysis of the mole salamanders found the existence of *Rhyacosiredon* makes *Ambystoma* [paraphyletic](/source/Paraphyletic), since the species are more closely related to some *Ambystoma* species than those species are to others in *Ambystoma*. The stream-type morphology of these salamanders (which includes larvae and [neotenes](/source/Neoteny) with short gills and thicker [gular folds](/source/Gular_fold)) may have led to their misclassification as a different genus.

The genus name *Ambystoma* was given by [Johann Jakob von Tschudi](/source/Johann_Jakob_von_Tschudi) in 1839.[14] Tschudi did not provide a derivation for the name, and many thought that he intended the name *Amblystoma*, "blunt-mouth". Occasionally, old specimens and documents use the name *Amblystoma*. Writing in 1907, [Leonhard Stejneger](/source/Leonhard_Stejneger) offered a derivation of *Ambystoma* based on a contraction of a Greek phrase meaning "to cram into the mouth",[15][16] but others have not found this explanation convincing.[17] In the absence of clear evidence that Tschudi committed a [lapsus](/source/Lapsus), the name given in 1839 stands.

## Species

The genus *Ambystoma* contains 32 extant [species](/source/Species), listed below, the most recently-named being *A. bishopi*. Some species are [Terrestrial](/source/Terrestrial_animal), others are [neotenic](/source/Neoteny#Neoteny_in_Other_Species), and some species have established populations of both neotenic and terrestrial forms.

Image Species and author Common name Distribution Lifestyle IUCN status A. altamirani Dugès, 1895 Mountain stream salamander, Achoque Central Mexico, west and south of the Valley of Mexico Terrestrial and neotenic A. amblycephalum Taylor, 1940 Blunt-headed salamander West-central Mexico (Michoacán state), near Morelia Terrestrial and neotenic A. andersoni (Brandon and Krebs, 1984) Anderson's salamander West-central Mexico (Michoacán state), Laguna de Zacapu Neotenic A. annulatum Cope, 1886 Ringed salamander South-central United States (Arkansas, Illinois, Missouri, Oklahoma), Ozark Plateau and Ouachita Mountains Terrestrial A. barbouri Kraus & Petranka, 1989 Streamside salamander South-midwest United States (Indiana, Kentucky, Ohio, Tennessee, West Virginia) Terrestrial A. bishopi Pauly, Piskurek & Shaffer, 2007 Reticulated flatwoods salamander Southeast United States (Florida Panhandle and southernmost Georgia), west of the Apalachicola-Flint River Terrestrial A. bombypellum (Taylor, 1940) Delicate-skinned salamander Central Mexico (State of Mexico) near Jilotepec Terrestrial A. californiense Gray, 1853 California tiger salamander Central Valley of California Terrestrial A. cingulatum Cope, 1868 Frosted flatwoods salamander Southeast United States (southern South Carolina and Georgia south to northern Florida) Terrestrial A. dumerilii (Dugès, 1870) Lake Pátzcuaro salamander, Achoque West-central Mexico (Michoacán state), Lake Pátzcuaro Neotenic A. flavipiperatum Dixon, 1963 Yellow-peppered salamander, Ajolote de Chapala West-central Mexico (Jalisco) Terrestrial A. gracile (Baird, 1859) Northwestern salamander Northwest North America (southernmost Alaska to northern California) Terrestrial A. granulosum Taylor, 1944 Granular salamander, Ajolote Central Mexico (State of Mexico) near Toluca Terrestrial A. jeffersonianum (Green, 1827) Jefferson salamander Northeastern North America (Ontario south to Virginia and west to Illinois) Terrestrial A. laterale Hallowell, 1856 Blue-spotted salamander Northeastern North America (Nova Scotia west to Manitoba and Minnesota and south to Indiana and New Jersey) Terrestrial A. leorae Taylor, 1943 Leora's stream salamander, Ajolote Central Mexico (Mexico state - Puebla border), Mount Tlaloc Terrestrial A. lermaense (Taylor, 1940) Lake Lerma salamander Central Mexico (State of Mexico), Lake Lerma near Toluca Terrestrial and neotenic A. mabeei Bishop, 1928 Mabee's salamander Coastal southeast United States (southeast Virginia to South Carolina) Terrestrial A. macrodactylum Baird, 1950 Long-toed salamander Northwest North America (Alaska south to northern California and east to Alberta and Montana) Terrestrial A. maculatum (Shaw, 1802) Spotted salamander Eastern North America (Nova Scotia west to Wisconsin and south to eastern Texas and Georgia) Terrestrial A. mavortium Baird, 1850 Barred tiger salamander Western North America (Manitoba south to Texas and west to Washington and California) Terrestrial and neotenic A. mexicanum (Shaw and Nodder, 1798) Axolotl Central Mexico (State of Mexico), Lake Xochimilco Neotenic A. opacum (Gravenhorst, 1807) Marbled salamander Eastern United States (New Hampshire south to northern Florida and east to Missouri and Texas) Terrestrial A. ordinarium Taylor, 1940 Puerto Hondo stream salamander West-central Mexico (Michoacán state), Puerto Hondo stream Terrestrial and neotenic A. rivulare Taylor, 1940 Michoacan stream salamander Central Mexico (western State of Mexico) Terrestrial and neotenic A. rosaceum Taylor, 1941 Tarahumara salamander Northwest Mexico, Sierra Madre Occidental Terrestrial and neotenic A. silvense Webb, 2004 Durango salamander Northwest Mexico (Durango and Chihuahua), Sierra Madre Occidental Terrestrial and neotenic A. talpoideum Holbrook, 1838 Mole salamander Southeast United States (Virginia west to Oklahoma and south to northern Florida) Terrestrial and neotenic A. taylori Brandon, Maruska, and Rumph, 1982 Taylor's salamander Southeast Mexico (Puebla), Laguna Alchichica Neotenic A. texanum Matthes, 1855 Small-mouth salamander South-central United States (Ohio west to Nebraska and south to Texas and Alabama) Terrestrial A. tigrinum (Green, 1825) Eastern tiger salamander Eastern North America (New York northwest to Manitoba and south to Texas and northern Florida) Terrestrial and neotenic A. velasci (Dugès, 1888) Plateau tiger salamander Mexican Plateau Terrestrial and neotenic

In addition, two groups of unisexual hybrid populations are sometimes named under their own species:

- [Silvery salamander](/source/Silvery_salamander) (*A. platineum*)
- [Tremblay's salamander](/source/Tremblay's_salamander) (*A. tremblayi*)

### Fossil species

Several extinct species of *Ambystoma* are known from fossils found throughout North America. These include:[18]

- *[Ambystoma kansense](/source/Ambystoma_kansense)* — late [Pliocene](/source/Pliocene), Kansas, USA
- *[Ambystoma hibbardi](/source/Ambystoma_hibbardi)* — late [Pliocene](/source/Pliocene), Kansas, USA
- *[Ambystoma minshalli](/source/Ambystoma_minshalli)* — late [Miocene](/source/Miocene)-early Pliocene, Nebraka, USA
- *[Ambystoma tiheni](/source/Ambystoma_tiheni)* — late Eocene, Saskatchewan, Canada
- *[Ambystoma quetzalcoatli](/source/Ambystoma_quetzalcoatli)* — late Pliocene, Mexico

## See also

- *[Oophila amblystomatis](/source/Oophila_amblystomatis)*

## References

1. Irschick, Duncan J. & Shaffer, H. Bradley (1997). "The Polytypic Species Revisited: Morphological Differentiation among Tiger Salamanders (Ambystoma tigrinum) (Amphibia: Caudata)". *Herpetologica*. **53** (1): 30–49. [ISSN 0018-0831](https://www.worldcat.org/issn/0018-0831). [JSTOR 3893241](https://www.jstor.org/stable/3893241)

1. Romano, Joseph. ["Ambystoma mavortium (Barred Tiger Salamander)"](https://animaldiversity.org/accounts/Ambystoma_mavortium/). *Animal Diversity Web*. Retrieved 2024-12-15.

1. Everson, Kathryn M.; Gray, Levi N.; Jones, Angela G.; Lawrence, Nicolette M.; Foley, Mary E.; Sovacool, Kelly L.; Kratovil, Justin D.; Hotaling, Scott; Hime, Paul M.; Storfer, Andrew; Parra-Olea, Gabriela; Percino-Daniel, Ruth; Aguilar-Miguel, X.; O'Neill, Eric M.; Zambrano, Luis (2021-04-27). "Geography is more important than life history in the recent diversification of the tiger salamander complex". *Proceedings of the National Academy of Sciences*. **118** (17). [Bibcode:2021PNAS..11814719E](https://ui.adsabs.harvard.edu/abs/2021PNAS..11814719E). [doi:10.1073/pnas.2014719118](https://doi.org/10.1073/pnas.2014719118). [PMC 8092605](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8092605). [PMID 33888580](https://pubmed.ncbi.nlm.nih.gov/33888580)

1. Tighe, Kenneth (2022-12-09). [*Catalog of Type Specimens of Recent Caudata and Gymnophiona in the National Museum of Natural History, Smithsonian Institution*](https://smithsonian.figshare.com/articles/book/Catalog_of_Type_Specimens_of_Recent_Caudata_and_Gymnophiona_in_the_National_Museum_of_Natural_History_Smithsonian_Institution/21699749?file=38489195). Vol. 654. Smithsonian Contributions to Zoology. Smithsonian Institution Scholarly Press. [doi:10.5479/si.21699749](https://doi.org/10.5479/si.21699749)

1. ["AmphibiaWeb - Ambystoma mavortium"](https://amphibiaweb.org/species/5887). *amphibiaweb.org*. Retrieved 2024-12-15.

1. Irschick, Duncan J. & Shaffer, H. Bradley (1997). "The Polytypic Species Revisited: Morphological Differentiation among Tiger Salamanders (Ambystoma tigrinum) (Amphibia: Caudata)". *Herpetologica*. **53** (1): 30–49. [ISSN 0018-0831](https://www.worldcat.org/issn/0018-0831). [JSTOR 3893241](https://www.jstor.org/stable/3893241)

1. ["AmphibiaWeb - Ambystoma velasci"](https://amphibiaweb.org/species/5886). *amphibiaweb.org*. Retrieved 2024-12-15.

1. Bi, Ke; Bogart, James P.; Fu, Jinzhong (2008). "The prevalence of genome replacement in unisexual salamanders of the genus Ambystoma (Amphibia, Caudata) revealed by nuclear gene genealogy". *[BMC Evolutionary Biology](/source/BMC_Evolutionary_Biology)*. **8** (1): 158. [Bibcode:2008BMCEE...8..158B](https://ui.adsabs.harvard.edu/abs/2008BMCEE...8..158B). [doi:10.1186/1471-2148-8-158](https://doi.org/10.1186/1471-2148-8-158). [ISSN 1471-2148](https://www.worldcat.org/issn/1471-2148). [PMC 2413238](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2413238). [PMID 18498635](https://pubmed.ncbi.nlm.nih.gov/18498635)

1. Bogart, James P.; Bi, Ke; Jinzong Fu; Noble, Daniel W.A.; Niedzwiecki, John (February 2007). "Unisexual salamanders (genus *Ambystoma*) present a new reproductive mode for eukaryotes". *[Genome](/source/Genome_(journal))*. **50** (2): 119–136. [doi:10.1139/g06-152](https://doi.org/10.1139/g06-152). [ISSN 0831-2796](https://www.worldcat.org/issn/0831-2796). [PMID 17546077](https://pubmed.ncbi.nlm.nih.gov/17546077)

1. Bogart, J.P.; Bartoszek, J.; Noble, D.W.A.; Bi, K. (2009-07-29). "Sex in unisexual salamanders: discovery of a new sperm donor with ancient affinities". *[Heredity](/source/Heredity_(journal))*. **103** (6): 483–493. [Bibcode:2009Hered.103..483B](https://ui.adsabs.harvard.edu/abs/2009Hered.103..483B). [doi:10.1038/hdy.2009.83](https://doi.org/10.1038/hdy.2009.83). [ISSN 0018-067X](https://www.worldcat.org/issn/0018-067X). [PMID 19639004](https://pubmed.ncbi.nlm.nih.gov/19639004)

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1. García-Lepe UO, Torres-Dimas E, Espinal-Centeno A, Cruz-Ramírez A, Bermúdez-Cruz RM. Evidence of requirement for homologous-mediated DNA repair during Ambystoma mexicanum limb regeneration. Dev Dyn. 2022 Jun;251(6):1035-1053. doi: 10.1002/dvdy.455. Epub 2022 Jan 25. PMID 35040539

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1. Lyon Jr., M. W. (30 June 1916). "*Ambystoma* not *Amblystoma*". *Science*. **43** (1122): 929–930. [Bibcode:1916Sci....43..929L](https://ui.adsabs.harvard.edu/abs/1916Sci....43..929L). [doi:10.1126/science.43.1122.929.a](https://doi.org/10.1126/science.43.1122.929.a). [JSTOR 1639383](https://www.jstor.org/stable/1639383). [PMID 17793100](https://pubmed.ncbi.nlm.nih.gov/17793100)

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1. Herrera-Flores, Jorge A. & Velasco-de León, María Patricia (May 2026). ["*Ambystoma quetzalcoatli* sp. nov., a new species of fossil axolotl (Caudata: Ambystomatidae) from the Pliocene of Santa María Amajac, Hidalgo, Mexico"](https://palaeo-electronica.org/content/2026/5636-ambystoma-quetzalcoatli-a-fossil-axolotl-from-mexico). *[Palaeontologia Electronica](/source/Palaeontologia_Electronica)*. **29** (2). [doi:10.26879/1644](https://doi.org/10.26879/1644)

- [http://amphibiaweb.org/lists/Ambystomatidae.shtml](http://amphibiaweb.org/lists/Ambystomatidae.shtml)

## External links

- [Tree of Life: Ambystomatidae](http://tolweb.org/Ambystomatidae/15448)
- [Herpseeker.dk](https://web.archive.org/web/20050529085120/http://www.herpseeker.com/worldspecies/Caudata/ambystomatidae/ambystoma.htm)
- [IUCN redlist of threatened Ambystomatidae](http://www.iucnredlist.org/search/search.php?freetext=ambystoma&modifier=phrase&criteria=wholedb&taxa_species=1&taxa_subspc=1&taxa_stock=1&redlistCategory%5B%5D=allex&redlistAssessyear%5B%5D=all&country%5B%5D=all&aquatic%5B%5D=all&regions%5B%5D=all&habitats%5B%5D=all&threats%5B%5D=all&Submit.x=33&Submit.y=8)

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Adapted from the Wikipedia article [Mole salamander](https://en.wikipedia.org/wiki/Mole_salamander) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Mole_salamander?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.