Cryptobranchus alleganiensis (Sonnini de Manoncourt and Latreille, 1801)

Class: Amphibia > Order: Caudata > Family: Cryptobranchidae > Genus: Cryptobranchus > Species: Cryptobranchus alleganiensis

Salamandra alleganiensis Sonnini de Manoncourt and Latreille, 1801 "An. X", Hist. Nat. Rept., 4: 406 (description in Sonnini de Manoncourt and Latreille, 1801 "An. X", Hist. Nat. Rept., 2: 253). Holotype: animal figured in Sonnini de Manoncourt and Latreille, 1801 "An. X", Hist. Nat. Rept., 2: figure preceding p. 253, of a specimen in the MNHNP, now lost. Type locality: "Virginie sur les montagnes Alléganis"; corrected to "vicinity of Davenport's Plantation", North Toe River, 1 mile south of the mouth of the Bushy Creek and 4 miles east-northeast of the Spruce Pine Creek, Mitchell County, North Carolina", USA, by Harper, 1940, Am. Midl. Nat., 23: 721. See Dundee, 1971, Cat. Am. Amph. Rept., 101: 3, for discussion of the type locality. See Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 49, for discussion of the original description, the publication date of which they give as 1802, although Harper, 1940, Am. Midl. Nat., 23: 692–723, and Frétey and Raffaëlli, 2021, Bionomina, 25: 47, suggests that it was published in 1801. 

Salamandra alleghaniensis Daudin, 1803 "An. XI", Hist. Nat. Gen. Part. Rept., 8: 231. 

Salamandra horrida Barton, 1807, Philadelphia Med. Phys. J., Suppl., 2 (2): 197. Type(s):  Not stated or known to exist although the animal figured by Barton, 1814 "1812", Mem. Animal Class Rept. Amph.: 15, is presumably of the type. Type locality: "in the lakes Ontario, Erie, & c., and also in the waters of the Ohio and Susquehanna . It seems, how ever, to be especially confined to the lakes , and to the waters of the Missisippi , including those which empty themselves into the Ohio", USA. One of three original names proposed for this taxon, of which Salamandra horrida Barton, 1807, was selected to have precedence by Barton, 1807, Discourse on Some Principal Desiderata Nat. Hist.: 23.  

Salamandra gigantea Barton, 1807, Philadelphia Med. Phys. J., Suppl., 2 (2): 197. Type(s):  Not stated or known to exist although the animal figured by Barton, 1814 "1812", Mem. Animal Class Rept. Amph.: 15, is presumably of the type. Type locality: "in the lakes Ontario, Erie, & c., and also in the waters of the Ohio and Susquehanna . It seems, how ever, to be especially confined to the lakes , and to the waters of the Missisippi , including those which empty themselves into the Ohio", USA. One of three original names proposed for this taxon, of which Salamandra horrida Barton, 1807, was selected to have precedence by Barton, 1807, Discourse on Some Principal Desiderata Nat. Hist.: 23. 

Salamandra maxima Barton, 1807, Philadelphia Med. Phys. J., Suppl., 2 (2): 197. Type(s):  Not stated or known to exist although the animal figured by Barton, 1814 "1812", Mem. Animal Class Rept. Amph.: 15, is presumably of the type. Type locality: "in the lakes Ontario, Erie, & c., and also in the waters of the Ohio and Susquehanna . It seems, how ever, to be especially confined to the lakes , and to the waters of the Missisippi , including those which empty themselves into the Ohio", USA. One of three original names proposed for this taxon. One of three original names proposed for this taxon, of which Salamandra horrida Barton, 1807, was selected to have precedence by Barton, 1807, Discourse on Some Principal Desiderata Nat. Hist.: 23.    

Salamandra horrida Barton, 1808, Some Account of Siren lacertina: 8. A subsequent use of Salamandra horrida Barton, 1807, but not so recognized until Frétey and Raffaëlli, 2021, Bionomina, 25: 35–51. When considered a first use it had a type locality of "in the great lakes of our country, in the waters of the Ohio, and Susquehanna [rivers], and other parts of the United States", USA; restricted to the "Muskingum River, Ohio", USA, by Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 11. This restriction deemed invalid by Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 48, as not being based on a neotype designation or evidence. Synonymy by Harlan, 1827, J. Acad. Nat. Sci. Philadelphia, 5: 320.

Triton alleganiensisOppel, 1811, Ordn. Fam. Gatt. Rept.: 81. New combination. 

Molge giganteaMerrem, 1820, Tent. Syst. Amph.: 187. New combination. 

Salamandra alleganensis — Leuckart, 1821, Isis von Oken, 9: 253. Incorrect subsequent spelling of species name. 

Cryptobranchus salamandroides Leuckart, 1821, Isis von Oken, 9: 260. Substitute name for Salamandra gigantea Barton, 1807, although Leuckart cited only Cryptobranchus gigantea Barton, 1814.  

Urotropis mucronata Rafinesque, 1822, Kentucky Gazette, Lexington, N.S.,, 1: 3. Type(s): Not designated or known to exist. Type locality: "the Kentucky river", Kentucky, USA. New combination. Synonymy by Brame, 1972, Checklist Living & Fossil Salamand. World (Unpubl. MS): 28.

Triton alleghaniensis — James, 1823, Account Exped. Pittsburgh–Rocky Mts., 3: 5. Incorrect subsequent spelling of the species name. 

Abranchus alleghaniensisHarlan, 1825, Ann. Lyc. Nat. Hist. New York, 1: 233. Incorrect subsequent spelling of species name.

Menopoma alleghaniensisHarlan, 1825, Ann. Lyc. Nat. Hist. New York, 1: 271. Incorrect subsequent spelling of species name.

Salamandra alleganensis — Gray, 1825, Ann. Philos., London, Ser. 2, 10: 217. Incorrect subsequent spelling of species name. 

Abranchus alleganensis — Gray, 1825, Ann. Philos., London, Ser. 2, 10: 217. Incorrect subsequent spelling of species name. 

Protonopsis horrida — Barnes, 1826, Am. J. Sci. Arts, 11: 278. New combination. 

Salamandrops giganteusWagler, 1830, Nat. Syst. Amph.: 209; Leunis, 1844, Synops. Drei Naturr., Zool., Ed. 1: 148.

Abranchus horridaGray, 1831, in Cuvier, Animal Kingdom (Griffith), 9—Appendix: 109.

Salamandra gigantia — Griffith and Pidgeon, 1831, Cuvier's Animal Kingdom, 9 (1): 410. Incorrect subsequent spelling of species name. 

Eurycea mucronataRafinesque, 1832, Atlantic. J. and Friend of Knowledge, Philadelphia, 1: 121.

Amphiuma (Menopoma) giganteaVan der Hoeven, 1833, Handb. Dierkd., 2: 304, by implication.

Cryptobranchus alleghaniensisVan der Hoeven, 1838, Tijdschr. Natuurl. Geschied., 4: 384; Cope, 1889, Bull. U.S. Natl. Mus., 34: 38. Incorrect subsequent spelling of the species name. 

Menopoma giganteaTschudi, 1838, Classif. Batr.: 96.

Salamandra alleghaniensisTschudi, 1838, Classif. Batr.: 96. Incorrect subsequent spelling of species name. 

Menopoma fusca Holbrook, 1842, N. Am. Herpetol., Ed. 2, 5: 99. Types: Specimen figured on pl. 33 of the original publication. Type locality: "waters of the mountainous regions of North Carolina and Georgia” and "waters of French Broad . . . of Ashville, Buncomb county, North Carolina", USA. Synonymy by Dundee, 1971, Cat. Am. Amph. Rept., 101: 1.

Salamandrops giganteusFitzinger, 1843, Syst. Rept.Fitzinger 1843: 34. New combination.

Triton alleghaniensisGray, 1850, Cat. Spec. Amph. Coll. Brit. Mus., Batr. Grad.: 54. Error.

Protonopsis horrida — Gray, 1850, Cat. Spec. Amph. Coll. Brit. Mus., Batr. Grad.: 53.

Protonopsis fuscaGray, 1850, Cat. Spec. Amph. Coll. Brit. Mus., Batr. Grad.: 54.

Salamandra alleghaniensisGray, 1850, Cat. Spec. Amph. Coll. Brit. Mus., Batr. Grad.: 53. Incorrect subsequent spelling in synonymy of species name.

Protonophis horrida — Duméril, Bibron, and Duméril, 1854, Erp. Gen., 9: 54. New combination. 

Salamandra (Menopoma) giganteaSchlegel, 1858, Handl. Dierkd., 2: 61. New combination. 

Menopoma alleghaniense Knauer, 1878, Naturgesch. Lurche: 96. New combination and incorrect subsequent spelling of the species name.  

Menopoma fuscumYarrow, 1882, Bull. U.S. Natl. Mus., 24: 20; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 82. 

Menopoma alleghaniense — Davis and Rice, 1883, Bull. Chicago Acad. Sci., 1: 26. Incorrect subsequent spelling of the species name. 

Cryptobranchus alleganiensisGarman, 1884, Bull. Essex Inst., 16: 36, by implication; Morse, 1901, Ohio Nat., 1: 114; Fowler and Dunn, 1917, Proc. Acad. Nat. Sci. Philadelphia, 69: 8;  Bishop, 1943, Handb. Salamanders: 59..

Cryptobranchus fuscusGarman, 1884, Bull. Essex Inst., 16: 36, by implication; Cope, 1889, Bull. U.S. Natl. Mus., 34: 43. New combination.

Salamandrops gigantea —  Cope, 1889, Bull. U.S. Natl. Mus., 34: 39. 

Cryptobranchus terassodactylos Wellborn, 1936, Zool. Anz., 114: 63–64. Holotype: ZMB 9639, according to Bauer, Good, and Günther, 1993, Mitt. Zool. Mus. Berlin, 69: 290. Type locality: "Nordamerika". Restricted to "Allegheny Mountain in Virginia", USA by Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 11; this restriction invalid according to Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 50. Synonymy by Grobman, 1943, Occas. Pap. Mus. Zool. Univ. Michigan, 470: 5. See comments by Frétey and Raffaëlli, 2021, Bionomina, 25: 42. 

Cryptobranchus alleganiensis alleganiensisSchmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 11.

Cryptobranchus bishopi Grobman, 1943, Occas. Pap. Mus. Zool. Univ. Michigan, 470: 6. Holotype: UMMZ 68930, by original designation. Type locality: "Current River at Big Spring Park, Carter County, Missouri", USA. Synonymy by Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 12.

Cryptobranchus alleganiensis bishopi — Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 12; Dundee and Dundee, 1965, Copeia, 1965: 169-170.

Cryptobranchus guildayi Holman, 1977, Ann. Carnegie Mus., 46: 157–172. Holotype: CM 20470, by original designation. Type locality: Cumberland Cave, Allegany County, Maryland, USA. Synonymy by Bredehoeft and Schubert, 2015, J. Herpetol., 49: 157. 

Cryptobranchus bishopi — Collins, 1991, Herpetol. Rev., 22: 43.

English Names

Allegany Hell-bender (Menopoma alleghaniensis: De Kay, 1842, Zool. New York, 1(3): 89).

Alleghany Hell-bender (Menopoma allegheniense: Yarrow, 1876, List Skeletons and Crania: 40).

Large Hellbender (Abranchus horrida: Gray, 1831, in Cuvier, Animal Kingdom (Griffith), 9—Appendix: 108).

Ground-puppy (Harlan, 1825, Ann. Lyc. Nat. Hist. New York, 1: 271 [as Menopoma alleghaniensis; Wood, 1863, Illust. Nat. Hist., 3: 185 [as Protonopsis horridus]).

Menopome (Protonopsis horrida: Wood, 1863, Illust. Nat. Hist., 3: 185).

Tweeg (Protonopsis horrida: Wood, 1863, Illust. Nat. Hist., 3: 185).

Mud Devil (Protonopsis horrida: Wood, 1863, Illust. Nat. Hist., 3: 185).

Tennessee Hellbender (Menopoma fuscum [no longer recognized]: Yarrow, 1882, Bull. U.S. Natl. Mus., 24: 20).

Brown Hellbender (Cryptobranchus fuscus [no longer recognized]; Rhoads, 1895, Proc. Acad. Nat. Sci. Philadelphia, 47: 406).

Giant Salamander (Cryptobranchus alleganiensis: Bishop, 1943, Handb. Salamanders: 59).

Hellbender (Cryptobranchus alleganiensis: Harlan, 1825, Ann. Lyc. Nat. Hist. New York, 1: 271; Cuvier, 1831, Animal Kingdom (M'Murtrie), 2: 88; Jordan, 1878, Man. Vert. North. U.S., Ed. 2: 196; Yarrow, 1882, Bull. U.S. Natl. Mus., 24: 20; Davis and Rice, 1883, Bull. Chicago Acad. Sci., 1: 26; Hay, 1892, Annu. Rep. Dept. Geol. Nat. Res. Indiana for 1891: 423; Rhoads, 1895, Proc. Acad. Nat. Sci. Philadelphia, 47: 403; Brimley, 1907, J. Elisha Mitchell Sci. Soc., 23: 151; Bishop, 1943, Handb. Salamanders: 59; Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 11; Conant, Cagle, Goin, Lowe, Neill, Netting, Schmidt, Shaw, Stebbins, and Bogert, 1956, Copeia, 1956: 174; Collins, Huheey, Knight, and Smith, 1978, Herpetol. Circ., 7: 5; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 28; Collins, 1997, Herpetol. Circ., 25: 6; Crother, Boundy, Campbell, de Queiroz, Frost, Highton, Iverson, Meylan, Reeder, Seidel, Sites, Taggart, Tilley, and Wake, 2001 "2000", Herpetol. Circ., 29: 20; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 15; Collins and Taggart, 2009, Standard Common Curr. Sci. Names N. Am. Amph. Turtles Rept. Crocodil., ed. 6: 10; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 25).

Alleghany Salamander (Cryptobranchus alleganiensis: Hay, 1892, Annu. Rep. Dept. Geol. Nat. Res. Indiana for 1891: 423).

Hellbender (Cryptobranchus alleganiensis alleganiensis: Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 11; Conant, Cagle, Goin, Lowe, Neill, Netting, Schmidt, Shaw, Stebbins, and Bogert, 1956, Copeia, 1956: 174; Conant, 1975, Field Guide Rept. Amph. E. Cent. N. Am., Ed. 2: 240),

Eastern Hellbender (Cryptobranchus alleganiensis alleganiensis: Collins, Huheey, Knight, and Smith, 1978, Herpetol. Circ., 7: 5; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 28; Crother, Boundy, Campbell, de Queiroz, Frost, Highton, Iverson, Meylan, Reeder, Seidel, Sites, Taggart, Tilley, and Wake, 2001 "2000", Herpetol. Circ., 29: 20; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 15; Collins and Taggart, 2009, Standard Common Curr. Sci. Names N. Am. Amph. Turtles Rept. Crocodil., ed. 6: 10; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 25; Powell, Conant, and Collins, 2016, Field Guide Rept. Amph. E. North Am., 4th ed.: 38; Highton, Bonett, and Jockusch, 2017, in Crother (ed.), Herpetol. Circ., 43: 25).

Eastern Hellbender (Cryptobranchus alleganiensisPowell, Conant, and Collins, 2016, Field Guide Rept. Amph. E. North Am., 4th ed.: 38). 

Ozark Hellbender (Cryptobranchus alleganiensis bishopiBishop, 1943, Handb. Salamanders: 63; Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 11; Conant, Cagle, Goin, Lowe, Neill, Netting, Schmidt, Shaw, Stebbins, and Bogert, 1956, Copeia, 1956: 174; Conant, 1975, Field Guide Rept. Amph. E. Cent. N. Am., Ed. 2: 241; Collins, Huheey, Knight, and Smith, 1978, Herpetol. Circ., 7: 5; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 28; Crother, Boundy, Campbell, de Queiroz, Frost, Highton, Iverson, Meylan, Reeder, Seidel, Sites, Taggart, Tilley, and Wake, 2001 "2000", Herpetol. Circ., 29: 20; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 15; Collins and Taggart, 2009, Standard Common Curr. Sci. Names N. Am. Amph. Turtles Rept. Crocodil., ed. 6: 10; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 25).

Ozark Hellbender (Cryptobranchus bishopiPowell, Conant, and Collins, 2016, Field Guide Rept. Amph. E. North Am., 4th ed.: 38). 

Distribution

Central and southwestern New York south to northern Maryland and central Ohio and western Virginia to westernmost South Carolina, northern Georgia, northern Alabama, northeastern Mississippi, Tennessee, and Ohio River drainage of southeastern Illinois, southern Indiana, USA; isolated populations in the Ozarks of Missouri and northern Arkansas, USA. See comment for discussion of the species complex and its nomenclature (DRF).

Comment

See accounts (all of which included Cryptobranchus bishopi as a subspecies or in synonymy) by Dundee, 1971, Cat. Am. Amph. Rept., 101: 1–4; Petranka, 1998, Salamand. U.S. Canada: 140–144; Nickerson and Mays, 1972, Publ. Biol. Geol. Milwaukee Public Mus., 1: 1–106, and Phillips and Humphries, 2005, in Lannoo (ed.), Amph. Declines: 648–651 Dubois and Raffaëlli, 2012, Alytes, 28: 77–161, also commented on the systematics of this species. Raffaëlli, 2013, Urodeles du Monde, 2nd ed.: 83–84, provided a brief account, photo, and map. Altig and McDiarmid, 2015, Handb. Larval Amph. US and Canada: 100–101, provided an account of larval morphology. Cryptobranchus bishopi was resurrected from the synonymy of Cryptobranchus alleganiensis by Collins, 1991, Herpetol. Rev., 22: 43, on the basis of its allopatry and diagnosability. Sabatino and Routman, 2009, Conserv. Genetics, 10: 1235–1246, reported on molecular phylogeography and conservation genetics, and suggested on the basis of mtDNA  analysis that nominal bishopi is phylogenetically imbedded within alleganiensis and that bishopi forms a paraphyletic grouping, with the North Fork of the White River and the Spring River populations in Missouri, more clearly related to populations of alleganiensis in the northern Ozarks, Ohio, and Tennessee, than to the bishopi population of the Current and Eleven Point River drainages of Missouri. Crowhurst, Faries, Collantes, Briggler, Koppelman, and Eggert, 2011, Conserv. Genetics, 12: 637–646, reporting on statistical similarity of microstellite DNA across drainages in the Ozarks of Missouri provided a more nuanced view: with Cryptobranchus bishopi populations sharing morphological characters but being composed of two genetically distinctive lineages associated with a) the North Fork and Bryant Creek drainages, and b) Current River and Eleven Point River drainages, with the North Fork and Bryant Creek populations more similar to alleganiensisFreake, O'Neill, Unger, Spear, and Routman, 2018, Conserv. Genetics, 19: 571–585, reported on mtDNA phylogeography in the Tennessee Valley, finding strong drainage loyalty and recommending that four populations be treated as conservation units. Hime, 2017, Unpubl. PhD Diss.: i–xv + 1–257, in his unpublished dissertation provided substantial evidence for the existence of five species within nominal Cryptobranchus alleganiensis (sensu lato):(1)  the White and Black River drainages in the Ozarks of south-central Missouri, USA; (2) Kanawha and New River drainages of southern and central West Virginia, western Virginia, and extreme northeastern Tennessee; (3) Tennessee River drainages of northeastern Mississippi, west-central Tennessee, northern Alabama, and northern Georgia, northeast to southeastern Kentucky and extreme southwestern Virginia; (4) Ohio, Allegheny, and Susquehanna River drainages of southern and southeastern Illinois southeast to central Tennessee, most of Kentucky, southern Indiana, central and eastern Ohio, extreme western Maryland, central and western Pennsylvania, and southwestern and southcentral New York; and (5) Missouri, Mississippi, and Green River drainages of central Missouri, USA. Hime applied the names Cryptobranchus bishopi to lineage #1 and Cryptobranchus alleganiensis to #3. Of the remaining three lineages, he suggested that substantial nomenclatural work was needed to find any available names that may be available. Frétey and Raffaëlli, 2021, Bionomina, 25: 35–51, provided a detailed discussion of the nomenclature of the complex and noted that the name Salamandra horrida Barton, 1807, is available for Hime's widespread species #4, but that Hime's #2 and #5 currently lack names. DRF is leaving this complex as a single taxon for the moment inasmuch as the evidence upon which this taxonomy is built is not currently formally published. 

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