Desmognathus marmoratus (Moore, 1899)

Class: Amphibia > Order: Caudata > Family: Plethodontidae > Subfamily: Plethodontinae > Genus: Desmognathus > Species: Desmognathus marmoratus

Leurognathus marmorata Moore, 1899, Proc. Acad. Nat. Sci. Philadelphia, 51: 316. Holotype: ANSP 19610, according to Fowler and Dunn, 1917, Proc. Acad. Nat. Sci. Philadelphia, 69: 21; Malnate, 1971, Proc. Acad. Nat. Sci. Philadelphia, 123: 348. Type locality: "large clear rocky pool on the south flank of Grandfather Mt., N.C. [Avery County, North Carolina, USA], and at an elevation of about 3,500 feet".

Leurognathus marmoratus — Brimley, 1907, J. Elisha Mitchell Sci. Soc., 23: 154.

Leurognathus marmorata marmorata — Pope, 1928, Am. Mus. Novit., 306: 14.

Leurognathus marmoratus marmoratus — Conant, 1958, Field Guide Rept. Amph. E. Cent. N. Am.: 227.

Leurognathus marmoratus intermedius — Conant, 1958, Field Guide Rept. Amph. E. Cent. N. Am.: 228.

Leurognathus marmoratus — Martof, 1962, Am. Midl. Nat., 67: 30. Rejects status of subspecies.

Desmognathus marmoratus — Bernardo, 1994, Am. Nat., 143: 15; Titus and Larson, 1996, Syst. Biol., 45: 451.

Desmognathus (Leurognathus) marmoratus — Dubois and Raffaëlli, 2012, Alytes, 28: 144. See comment under Desmognathus regarding the status of the subgenus.

Desmognathus marmortus — Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 176. Incorrect subsequent spelling. 

Common Names

Moore's Triton (Desmognathus marmorata: Brimley, 1907, J. Elisha Mitchell Sci. Soc., 23: 154).

Shovel-nosed Salamander (Desmognathus marmorata: Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 32; 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: 270; Crother, Boundy, Campbell, de Queiroz, Frost, Highton, Iverson, Meylan, Reeder, Seidel, Sites, Taggart, Tilley, and Wake, 2001 "2000", Herpetol. Circ., 29: 21; Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39: 16; 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.: 47; Highton, Bonett, and Jockusch, 2017, in Crother (ed.), Herpetol. Circ., 43: 26).

Shovelnose Salamander (Desmognathus marmorata: Collins, Huheey, Knight, and Smith, 1978, Herpetol. Circ., 7: 7; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 32; Collins, 1997, Herpetol. Circ., 25: 6; Collins and Taggart, 2009, Standard Common Curr. Sci. Names N. Am. Amph. Turtles Rept. Crocodil., ed. 6: 12).

Moore's Salamander (Leurognathus marmorata marmorata) [no longer recognized]: Bishop, 1943, Handb. Salamanders: 220).

Northern Shovel-nosed Salamander (Leurognathus marmorata marmorata) [no longer recognized]: Schmidt, 1953, Check List N. Am. Amph. Rept., Ed. 6: 32; Conant, Cagle, Goin, Lowe, Neill, Netting, Schmidt, Shaw, Stebbins, and Bogert, 1956, Copeia, 1956: 174).

Distribution

Southwestern Virginia and southward east of the Appalachian divide through North Carolina to eastern Tennessee, USA, 300 to 1700 m elevation. See comment. 

Geographic Occurrence

Natural Resident: United States of America, United States of America - North Carolina, United States of America - Tennessee, United States of America - Virginia, United States of America - West Virginia

Endemic: United States of America

Comment

This species name has undergone an enormous transformation from applying to nearly all of the Desmognathus marmoratus/quadramaculatus complex prior to the revisionary work of Beamer and Lamb, 2020, Zootaxa, 4734: 1–61, Pyron, O'Connell, Lemmon, Lemmon, and Beamer, 2020, Mol. Phylogenet. Evol., 146 (106751): 1–13, Pyron and Beamer, 2022, Bionomina, 27: 1–43, to the (maybe) final nomenclatural resolution by Raffaëlli, 2022, Salamanders & Newts of the World: 1012, with the restriction of the name to Desmognathus marmoratus E of Beamer and Lamb, 2020, Zootaxa, 4734: 54. All literature prior to 2020 should be employed with caution. 

See detailed accounts by Martof, 1963, Cat. Am. Amph. Rept., 3: 1–2 (as Leurognathus marmoratus) and Petranka, 1998, Salamand. U.S. Canada: 184–187. Voss, Smith, Beachy, and Heckel, 1995, J. Herpetol., 29: 493–497, reported on geographic allozyme variation and suggested that taxonomic revision (likely recognition of additional species) is warranted to recognize the distinctiveness of three populations: 1) the population on the west side of the Appalachian divide, drained by the Nanantahala River; 2) the population in the drainage of the Chattahoochee River; and 3) the populations of the drainage of the Tallulah and Chattooga rivers, which they referred to as in the Savannah drainage. They did not examine specimens from Virginia. Camp and Tilley, 2005, in Lannoo (ed.), Amph. Declines: 711–713, provided an account containing a detailed summary of the literature and range. Kozak, Larson, Bonett, and Harmon, 2005, Evolution, 59: 2000–2016, found multiple clades composed of intercalated populations of both Desmognathus marmoratus and Desmognathus quadramaculatus, suggesting that multiple species under both names or, alternatively, the possibility of very complicated hybridization and introgression among populations of Desmognathus marmoratus and Desmognathus quadramaculatus given that the genes samples were all mitochondrial and therefore subject to cytoplasmic inheritance and the morphological distinctiveness of these nominal species has not previously been questioned. Subsequently, but apparently unaware of the publication of Kozak et al. (2005) Jones, Voss, Ptacek, Weisrock, and Tonkyn, 2006, Mol. Phylogenet. Evol., 38: 280–287 (supported subsequently by mtDNA evidence published by Wooten and Rissler, 2011, Acta Herpetol., Firenze, 6: 175–208), suggested on the basis of mtDNA that neither Desmognathus marmoratus nor Desmognathus quadramaculatus are monophyletic, instead being composed of multiple inter-related species and suggested strongly that the population restricted to the Chattahoochee, Tallulah, and Chattooga River basins of northeastern Georgia and South Carolina, although there was some ambiguity in their statement, presumably due to the background knowledge of the common occurrence of mtDNA introgression in desmognathines. Dubois and Raffaëlli, 2009, Alytes, 26: 145, expanded on this, recognizing Desmognathus aureatus (the Chattahoochee drainage), but also applied the name Desmognathus melianus to the population on the west side of the continental divide in the Appalachian Nantahala drainage of eastern Tennessee and North Carolina. Substantial thought by people who know desmognathines has focused on this problem with at least the salamander subcommittee of the SSAR Common and Standard Names List (Tilley, Highton, and Wake, 2012, in Crother (ed.), Herpetol. Circ., 39), deciding to not recognize Desmognathus aureatus and Desmognathus melanius due to the practical lack of decisiveness on results based solely on mtDNA in a group famous for mtDNA introgression (although the allozymic evidence of Voss et al., 1995, corroborates these results). While I certainly respect the decision of that group, my inclination here is to recognize Desmognathus aureatus and Desmognathus melanius (now in the synonymy of Desmognathus intermedius) pending publication of counter-evidence. Raffaëlli, 2013, Urodeles du Monde, 2nd ed.: 431–432, provided a brief account, photograph, and range map. Altig and McDiarmid, 2015, Handb. Larval Amph. US and Canada: 103–104, provided an account of larval morphology. Beamer and Lamb, 2020, Zootaxa, 4734: 1–61, in their discussion of Desmognathus mtDNA phylogenetics, confirmed the intercalation of apparent cryptic species under this name with apparent cryptic species of Desmognatus quadramaculatus. Pyron, O'Connell, Lemmon, Lemmon, and Beamer, 2020, Mol. Phylogenet. Evol., 146 (106751): 1–13, suggested on molecular grounds that this nominal species is a complex of lineage-species. See Pyron and Beamer, 2022, Bionomina, 27: 1–43, for an extensive discussion of the complexity of this array of lineages, all of which do not appear to be closest relatives, being imbedded within various places with the phylogeny of Desmognathus. Raffaëlli, 2022, Salamanders & Newts of the World: 1012, provided an account summarizing systematics, morphology, life history, population status, and distribution (including a polygon map).  

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