identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
C8746C4AFFE70A17FF1BF979FECBFE64.text	C8746C4AFFE70A17FF1BF979FECBFE64.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Desmognathus marmoratus (Moore 1899)	<div><p>Desmognathus marmoratus (Moore, 1899)</p> <p>Leurognathus marmorata Moore, 1899</p> <p>Syntypes and Type Locality: Moore (1899) ’s description was explicitly based on both a male and female syntype along with an additional female specimen which was skeletonized for illustration. These were taken from “a large clear rocky pool beneath a waterfall of a stream on the south flank of Grandfather Mt., N. C., and at an elevation of about 3,500 feet.” A site matching this description is found at Green Mountain Creek Falls (36.114308, -81.778430; 1182m) on U.S. Hwy 221 near Blowing Rock (NC: Watauga), which we suggest is the type locality. The male syntype was deposited by Moore in 1916 at the Academy of Natural Sciences of Philadelphia as ANSP 19610. The two females (the alcoholic syntype and the skeleton) were said by Dunn (1917) to have been examined by himself at the University of Pennsylvania, presumably still in the collection of Moore, who was still a professor there at the time. Fowler and Dunn (1917) listed ANSP 19610 as a paratype, while Malnate (1971) listed it as the holotype. It is neither, but rather a syntype as per the original publication. We here designate it as the lectotype of Leurognathus marmorata Moore, 1899 to remove any ambiguity. This renders the second, female syntype a paralectotype. This specimen is of unknown disposition; it was originally in the P.J. Moore collection at the University of Pennsylvania, but inquiries to the university and the ANSP yielded no information. The female skeleton is also not known to exist but is not part of the type series and carries no name-bearing function.</p> <p>Description: After Moore (1899) and Martof (1956), a large (SVL = ~ 30–75mm), highly aquatic, fin-tailed salamander with slit-like internal nares; an indistinct dorsal color pattern consisting of a dark-brown background color and numerous mottled markings lacking light-colored centers; darker or grey venter; adult males lacking vomerine teeth, females with very few; large, prominent eyes; and snout usually not darker than the remainder of body (Fig. 5, 6). Can putatively be differentiated from the other two described shovel-nosed species based on the original characters given in the other accounts below, though these have not been re-assessed based on modern understandings of species limits and are of unknown diagnostic utility. Based on our size-corrected linear morphometric analyses, this species can be distinguished from Desmognathus aureatus and D. intermedius by a shorter head (SG in original measurements = 4.3–16.4mm) and from D. intermedius by a shorter trunk (AG in original measurements = 9.3–41.7mm). Of the 10 specimens observed in West Virginia, 8 were adults and the largest (a gravid female) was only ~ 60mm SVL, suggesting that this population may represent a dwarf race since the median SVL of 45 metamorphosed individuals sampled from southern populations is ~ 62mm. Ironically, little is known about the natural history of this species despite it being the earliest-described species as most studies have focused on D. aureatus and D. intermedius. Pope (1924) found a clutch of 28 eggs on 6 August 1924, and Bishop (1924) sampled females in October with 26–34 ovarian eggs. Previous reports suggest that larvae typically metamorphose between 26–33mm SVL (see Camp and Tilley in Lannoo 2005 and references therein).</p> <p>Range: In high-gradient mountain streams of the Appalachians, on the Blue Ridge northeast of the Pigeon River to southwestern Virginia. Populations are known from the South Fork Holston, Watauga, and Nolichucky (Upper Tennessee) and Upper Broad and Upper Catawba (Santee) River drainages. Also known from a handful of riffle zones along a 2–3km stretch of Glade Creek near Beckley, Raleigh County, West Virginia in the Lower New (Kanawha) River drainage. We are confident that this species is absent from several other similar creeks in the New River Gorge in which we exerted similar sampling effort (Fig. 2). In contrast, other nearby streams with apparently similar geological and hydrological characteristics may harbor populations of this species. This population is widely (~120 airline km) disjunct from the nearest known populations in the Upper Tennessee drainage in southwestern Virginia (Smyth and Washington Counties). Elevational range ~ 450–1600m (see Fig. 1 for localities).</p> <p>Habitat: Occurs almost exclusively in the rocky riffle zones of a fast-flowing, high-gradient mountain stream of the “typical” character associated with other Shovel-nosed Salamanders farther south in the Appalachian Mountains (Martof 1962).</p> <p>Etymology: The specific epithet is a Latin singular adjective in the nominative case meaning “marbled,” in reference to the dorsal color-pattern.</p> <p>Standard English Names: Moore’s Triton (Brimley 1907), Moore’s Salamander (Bishop 1943), Northern Shovel-nosed Salamander (Schmidt 1953), Shovelnose Salamander (Collins et al. 1978). We suggest that “Northern Shovel-nosed Salamander” is most appropriate.</p> <p>Conservation: The re-described concept of this species occupies a smaller range than the previous, more inclusive definition but is nevertheless widespread and abundant at numerous historical localities. Consequently, we suggest that it retain its “LC—Least Concern” status on the IUCN Red List based on the known information (see Maes et al. 2015). Special attention should be paid to the isolated population in the New River Gorge of West Virginia.</p> <p>Notes: First assigned to Desmognathus by Titus and Larson (1996) after the unpublished results of Titus (1992) and the informal suggestion of Bernardo (1994). Genetic variation in the widespread former concept of this species was first reported by Voss et al. (1995) and expanded by Jackson (2005), Kozak et al. (2005), and Jones et al. (2006), who confirmed the existence of multiple species-level taxa based on phylogenetic analysis of mitochondrial and nuclear DNA. This echoed earlier morphological work (Pope 1928; Martof 1956), though Martof (1962) had later downplayed the significance of this phenotypic variation.</p> <p>Accordingly, we offer revised accounts of the other known species of shovel-nosed salamander. These occur in two distantly related clades, Nantahala (Desmognathus aureatus) and Pisgah (D. intermedius). Beamer and Lamb (2020) and Pyron et al. (2020, 2022c) sampled the type locality of Leurognathus m. aureata Martof, 1956 and populations geographically proximate (5.5 airline km) to the type locality of L. m. roborata Martof, 1956, confirming their inclusion in marmoratus B. Acting as the First Reviser (Article 24.2.1), Raffaëlli (2013) had previously selected the former as the senior subjective synonym when the two taxa are considered conspecific. Consequently, this lineage should be referred to as Desmognathus aureatus (Martof, 1956) and include D. roboratus (Martof, 1956) in synonymy.</p> </div>	https://treatment.plazi.org/id/C8746C4AFFE70A17FF1BF979FECBFE64	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Pyron, R. Alexander;Beamer, David A.	Pyron, R. Alexander, Beamer, David A. (2023): A systematic revision of the Shovel-nosed Salamander (Plethodontidae: Desmognathus marmoratus), with re-description of the related D. aureatus and D. intermedius. Zootaxa 5270 (2): 262-280, DOI: 10.11646/zootaxa.5270.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5270.2.5
C8746C4AFFE40A10FF1BFE62FCEFF9B7.text	C8746C4AFFE40A10FF1BFE62FCEFF9B7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Desmognathus aureatus (Martof 1956)	<div><p>Desmognathus aureatus (Martof, 1956)</p> <p>Leurognathus marmorata aureata Martof, 1956</p> <p>Leurognathus marmorata roborata Martof, 1956, holotype UMMZ 111568 by original designation, collected 18 September 1954 by “the Bernard Martof family,” type locality “Reed Creek, along Burrell Ford Road, 0.5 of a mile from its junction with Glade School Road, about 3.5 miles northwest of Pine Mountain Community, Rabun County, Georgia; 2,350 feet elevation.” Paratypes UMMZ 111569 (lot of 26 specimens); same data. Informally synonymized with L. marmorata by Martof (1962) and resurrected by Dubois and Raffaëlli (2012) based on a suggestion by Jones et al. (2006). Here designated a junior subjective synonym of D. aureatus.</p> <p>Holotype: UMMZ 111566 by original designation, collected 11 September 1954 by “ Stephen and Andrew Martof,” type locality “ Jarrard’s Creek, 0.2 of a mile below its crossing of U.S. Route 19, about 9 air miles north-northeast of Dahlonega, Lumpkin County, Georgia; 1,550 feet elevation.”</p> <p>Paratypes: UMMZ 111567 (lot of 20 specimens); “data as for the holotype.”</p> <p>Description: After Martof (1956), adults of this species have no vomerine teeth (versus presence in Desmognathus intermedius and D. marmoratus [some females]); ventral color-pattern pale or mottled (versus dark or black in other Blue Ridge lineages); and dark brown to black background of dorsum with dorsal color-pattern consisting of paired, golden, marbled or lichenous blotches (albeit less distinct in northeastern populations previously assigned to L. m. roborata Martof, 1956). These characters should be re-evaluated for diagnostic consistency. In our sample, this is the smallest shovel-nosed species, with metamorphosed SVL = 34–73mm. Based on our size-corrected linear morphometric analyses, this species can be distinguished from D. intermedius by a shorter trunk (AG in original measurements = 17–40mm) and from D. marmoratus by a longer head (SG in original measurements = 8.1–17.1mm). From Martof (1962), females at sites in Georgia develop sexual maturity around 55–59mm SVL, typically laying 27– 54 eggs from May to July, with clutch size dependent on body size. Hatchlings are as small as 10–12mm SVL and metamorphose at 25–38mm (usually ~30) after 10–20 months, though up to three distinct size classes at some suggest possibly longer periods up to ~36 months. Larvae and adults are primarily insectivorous, likely preying opportunistically on available aquatic invertebrates (Martof and Scott 1957). Adults in some Georgia populations are occasionally bright yellow, a characteristic that apparently shifts rapidly in frequency within populations over short timescales (Martof and Walton 1965).</p> <p>Range: The Blue Ridge Mountains of northeastern Georgia, northwestern South Carolina, and small portions of the nearby border regions of North Carolina, in headwater streams of the Chattahoochee, Chattooga, and Tallulah River drainages (Fig. 1; Voss et al. 1995; Jones et al. 2006). Elevational range ~ 400–1100m.</p> <p>Habitat: As with all shovel-nosed salamanders, occurs almost exclusively in the rocky riffle zones of a fast-flowing, high-gradient mountain streams.</p> <p>Etymology: The specific epithet is a Latin singular adjective in the nominative case meaning “adorned with gold,” in reference to the dorsal coloration (Fig. 7).</p> <p>Standard English Names: For Desmognathus aureatus: Golden Shovel-nosed Salamander (Conant 1958); for Leurognathus marmoratus roboratus Martof, 1956: Husky Shovel-nosed Salamander (Conant 1958). We suggest that “Southern” Shovel-nosed Salamander is more appropriate.</p> <p>Conservation: This species is apparently widespread and abundant at numerous historical and recent sites across its range, a significant proportion of which is protected by state, federal, and private entities. Consequently, we suggest that it be considered “LC—Least Concern” based on available data (Maes et al. 2015).</p> <p>Notes: Implicitly synonymized with Leurognathus marmoratus (Moore, 1899) by Martof (1962). Indirectly suggested for resurrection by Jones et al. (2006), though this was essentially ignored. Later explicitly resurrected by Dubois and Raffaëlli (2012), though few subsequent authorities followed this arrangement. Part of the Nantahala clade (Jones and Weisrock 2018; Pyron et al. 2020, 2022c) and the sister lineage of the Dwarf Black-bellied Salamander Desmognathus folkertsi Camp, Tilley, Austin, &amp; Marshall, 2002. The presence of the shovel-nosed phenotype in this lineage may have been transmitted to or from the Pisgah clade via an episode of ancient adaptive introgression (Pyron et al. 2020).</p> <p>Within the Pisgah clade, Beamer and Lamb (2020) and Pyron et al. (2020, 2022c) sampled specimens from the type localities of Leurognathus marmorata intermedia Pope, 1928 and L. m. melania Martof, 1956, which were highly similar genetically and belonged to the marmoratus C lineage. Therefore, we conclude that marmoratus C should be referred to as Desmognathus intermedius (Pope, 1928), including D. melanius (Martof 1956) in synonymy. Consequently, the species D. melanius recognized by Frost (2019) following Dubois and Raffaëlli (2012) and Raffaëlli (2013) is not valid but is instead preoccupied by:</p> </div>	https://treatment.plazi.org/id/C8746C4AFFE40A10FF1BFE62FCEFF9B7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Pyron, R. Alexander;Beamer, David A.	Pyron, R. Alexander, Beamer, David A. (2023): A systematic revision of the Shovel-nosed Salamander (Plethodontidae: Desmognathus marmoratus), with re-description of the related D. aureatus and D. intermedius. Zootaxa 5270 (2): 262-280, DOI: 10.11646/zootaxa.5270.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5270.2.5
C8746C4AFFE30A12FF1BF953FDFBFA27.text	C8746C4AFFE30A12FF1BF953FDFBFA27.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Desmognathus intermedius (Pope 1928)	<div><p>Desmognathus intermedius (Pope, 1928)</p> <p>Leurognathus marmorata intermedia Pope, 1928</p> <p>Leurognathus marmorata melania Martof, 1956 type specimen UMMZ 111564 by original designation, collected 4 Sepember 1954 by “the Bernard Martof family,” type locality “Otter Creek, tributary of the Nantahala River, 0.5 of a mile west of Tellico Gap, Macon County, North Carolina; 3,600 feet elevation.” Paratypes UMMZ 111565 (20 specimens); same data. Informally synonymized with L. marmoratus by Martof (1962) and resurrected by Dubois and Raffaëlli (2012) and Raffaëlli (2013) as D. “melianus” in error, though few subsequent workers followed this recommendation. Designated a junior subjective synonym of D. intermedius by Raffaëlli (2022).</p> <p>Holotype: AMNH 25557 (a male) by original designation, collected 17 July 1927 by C.H. Pope, type locality “ Davis Gap, Waynesville, North Carolina ” and emended by Bishop (1943) to “ Davis Farm, about 2 miles east and a little north of Waynesville, (Haywood County) on Highway No. 276.”</p> <p>Paratypes: AMNH 25544–25556 &amp; 25558–25580 from the original type collection are listed as paratypes in the AMNH catalogue, but not explicitly designated as such by Pope (1928).</p> <p>Description: Among shovel-nosed salamanders, Desmognathus intermedius can be diagnosed from D. aureatus by the presence of vomerine teeth in adults of both sexes, and from D. marmoratus by a more scattered and indistinct dorsal pattern in adults (after Pope 1928; Pope and Hairston 1947; Fig. 8, 9). These characters should be re-evaluated for diagnostic consistency. Ironically given the original intent of the name “ intermedius,” this is by far the largest shovel-nosed species in our sample with metamorphosed SVL = 28–85mm. Based on our size-corrected linear morphometric analyses, this species can be distinguished from both D. aureatus and D. marmoratus by a longer trunk (AG in original measurements = 14–45mm) and from D. marmoratus by a longer head (SG in original measurements = 6.5–20.0mm). After Martof (1962) and Bruce (1985), larvae are generally 15–35mm SVL, with a 3-year period until metamorphosis and oviposition and transformation both occurring in summer.</p> <p>Range: In the Blue Ridge Mountains of western North Carolina and eastern Tennessee, confirmed in the Tennessee drainages of the Lower Little Tennessee, Upper Little Tennessee, Tuckasegee, Lower French Broad, Pigeon, and Upper French Broad Rivers (Fig. 1). Curiously rare in or absent from the upper reaches of the Little Tennessee and Tuckasegee Rivers (Martof 1962, 1963). An isolated population is known from the Black Mountains east of the French Broad River in the Upper French Broad drainage (O’Connell et al., unpubl. data). Elevational range ~ 300–1500m.</p> <p>Habitat: As with all Shovel-nosed Salamanders, occurs almost exclusively in the rocky riffle zones of fast-flowing, high-gradient mountain streams.</p> <p>Etymology: The specific epithet is a Latin singular adjective in the nominative case meaning “intermediate,” apparently referring to the species’ supposedly smaller size compared to D. marmoratus as known to Pope (1928) based on relatively few specimens.</p> <p>Standard English Names: For Leurognathus marmorata intermedia: Pope’s Salamander (Bishop 1943), Southern Shovel-nosed Salamander (Schmidt 1953); for Leurognathus m. melanius: Black Shovel-nosed Salamander (Conant 1958), Otter Creek Dusky Salamander (Fouquette and Dubois 2014). We suggest that “Western” Shovel-nosed Salamander is most appropriate.</p> <p>Conservation: This species is apparently widespread and abundant at numerous historical and recent sites across its range, a significant proportion of which is protected by state, federal, and private entities. Consequently, we suggest that it be considered “LC—Least Concern” based on available data (Maes et al. 2015).</p> <p>Notes: This lineage was previously recognized as a distinct species Leurognathus intermedia by Pope and Hairston (1947) before being demoted to a subspecies L. m. intermedia by Schmidt (1953) and informally synonymized with L. marmoratus by Martof (1962).</p> </div>	https://treatment.plazi.org/id/C8746C4AFFE30A12FF1BF953FDFBFA27	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Pyron, R. Alexander;Beamer, David A.	Pyron, R. Alexander, Beamer, David A. (2023): A systematic revision of the Shovel-nosed Salamander (Plethodontidae: Desmognathus marmoratus), with re-description of the related D. aureatus and D. intermedius. Zootaxa 5270 (2): 262-280, DOI: 10.11646/zootaxa.5270.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5270.2.5
