taxonID	type	description	language	source
039087A5C5023F4FC4BA1F2770890916.taxon	materials_examined	Holotype: USNM 596065 (RAP 0892; Figs. 5 – 8), collected 14 August 2019 by RAP and T. W. Pierson on Cataloochee Balsam in Great Smoky Mountains National Park (NC: Swain; Fig. 9), ~ 1650 m ASL. Paratypes: USNM 596063 – 4 (RAP 0890 – 1; Figs. 6 – 8, 10, 11), same collection as the holotype. NCSM 108356 (RAP 0909), collected 15 August 2019 by RAP and T. W. Pierson on Rough Ridge (NC: Haywood; 35.543, - 83.156), ~ 1610 m ASL.	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5023F4FC4BA1F2770890916.taxon	diagnosis	Diagnosis: Several phenotypic characters exhibit varying combinations of states which are purported to be diagnostic and differentiate D. adatsihi from related or similar taxa. This species is a small (~ 23 – 50 mm SVL; n = 15), primarily terrestrial mountain dusky salamander with a tail shorter than the body (up to ~ 90 % SVL) that is round in cross section along its length. It is differentiated from D. santeetlah by the lack of any keeling on the dorsal surface of the tail (vs. presence thereof). It is sometimes differentiated from D. imitator by a morph with the prominent remnants of paired yellowish, reddish, or orangish dorsal spots connected by a wash of lighter pigments on a lighter ground color (Figs. 6 – 8, 11; vs. more distinct and separated spots on a darker ground color in some D. imitator), or in uniformly dark older specimens, a lack of red cheek patches (Fig. 12 a; vs. the presence of such coloration in some uniformly dark D. imitator). Many individuals can be uniquely diagnosed by the presence of an exceptionally straight and unadorned dorsal stripe of brownish or yellowish color (Figs. 5 – 8, 13 a), a characteristic most common in Smokies populations. This trait is never observed in D. imitator, other D. ocoee lineages, or D. santeetlah, which if exhibiting dorsal color pattern, all typically possess paired spots with varying degrees of intercalated melanophore pigmentation, sometimes yielding relatively unadorned stripes with irregular edges. Some individuals from the paratype locality exhibit yellowish dorsal stripes with irregular edges and indistinct black patterning on the mid-dorsal line (Fig. 13 b), an arrangement almost never observed in D. apalachicolae and very rarely in any other D. ocoee lineages. Very rarely, other color patterns include characteristic erythristic or xanthic morphs that have never been observed in other species (Figs. 6 – 8, 10, 14 a) and appear to be uniquely diagnostic when present.	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5023F4FC4BA1F2770890916.taxon	distribution	Distribution: In the Great Smoky Mountains of Tennessee and North Carolina and the Plott Balsam Mountains of North Carolina at high elevations. All genetically verified samples illustrated here (Fig. 4) originated> 1500 m ASL. Tilley et al. (1978) tentatively suggested ~ 1200 m ASL as the approximate lower limit for this species and reported Starkey Gap as the westernmost limit of its range. An informal transect sampling several points at the paratype locality along the western slope of Rough Ridge below Strawberry Knob (NC: Haywood) revealed only D. imitator at ~ 1340 m, ~ 1460 m, and ~ 1490 m, and only D. adatsihi at ~ 1540 m, ~ 1570 m, and ~ 1610 – 1630 m; previous surveys from near this locality (Mashie Stomp Creek) also found only D. imitator at ~ 1350 m (Tilley 2000).	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5023F4FC4BA1F2770890916.taxon	etymology	Etymology: From a Tsalagi (Cherokee) word (in syllabary:) that translates in a rough sense to “ the mother of all, ” in reference to the matrilineal inheritance of the mitochondrion, which has an unusual hybrid ancestry in this species along with ocoee B – D. Name is a non-Latin singular noun used in apposition. We suggest the common names “ Cherokee ” or “ Great Smokies ” Mountain Dusky Salamander. Notes: Comprises the ocoee A lineage first tentatively identified by Tilley et al. (1978) and defined by Kozak et al. (2005), Beamer & Lamb (2020), and Pyron et al. (2020, 2022 c). Little is known of the biology of this species. Like many other mountain duskies, it is highly terrestrial and appears to spend most of its adult life on the forest floor significant distances away from any permanent flowing water. Some populations from the Smokies were briefly described by Huheey (1966 b) and Dodd (2004). An aberrant, possibly leucistic morph has been observed at the paratype locality (Fig. 15). The third species accounted for here is the ocoee B lineage, the second member of the Balsam clade of Pyron et al. (2022 c). This taxon is distributed throughout the Great Balsam Mountains of western North Carolina, southeast of Balsam Gap. It shares hybridized non-lineal mitochondrial ancestry with Desmognathus adatsihi and ocoee C / D but is otherwise genealogically exclusive and exhibits no apparent admixture with any other lineage. Consequently, we recognize it here as:	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5183F4EC4BA1EC072920E26.taxon	materials_examined	Holotype: AMNH A- 193885 (RAP 0650; Fig. 16), collected 12 April 2019 by RAP from Bubbling Spring Branch near Mount Hardy Gap (NC: Haywood; 35.308, - 82.908, ~ 1530 m ASL). Paratypes: AMNH A- 193884 & A- 193886 (RAP 0649 & 0651), as from the type. MNHN 2021.0115 & BMNH 2021.7526 (RAP 0644 – 5), collected 12 April 2019 by RAP on Green Ridge (NC: Haywood; 35.353, - 82.917), ~ 1110 m ASL. NCSM 108355 (RAP 0647), collected 12 April 2019 by RAP near Sunburst Falls (NC: Haywood; 35.338, - 82.904), ~ 1260 m ASL.	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5183F4EC4BA1EC072920E26.taxon	diagnosis	Diagnosis: Several phenotypic characters exhibit varying combinations of states which are purported to be diagnostic and differentiate D. balsameus from related or similar taxa. This species is a small (~ 19 – 47 mm SVL; n = 22), primarily terrestrial mountain dusky salamander with a tail shorter than the body (up to ~ 90 % SVL) that is round in cross section along its length. It is differentiated from D. santeetlah by the lack of any keeling on the dorsal surface of the tail (vs. presence thereof), which is the only species in its range with which it is likely to be confused. Exhibits a wide range of color patterns, with most individuals possessing a partially or mostly complete dorsal stripe with wavy or undulating edges (vs. usually straight in D. adatsihi), formed from the irregular invasion of melanophore patches between the remnants of paired larval spots, with a brownish, yellowish, or reddish coloration against a darker lateral ground color, and a general lack of any regular patterning along the mid-dorsal line (Figs. 17 – 18). Ontogenetic darkening seems to be relatively more common and at smaller sizes than in related species, with individuals as small as ~ 33 mm SVL exhibiting overall dark brown to blackish coloration obscuring the dorsal pattern (Fig. 12 b, 14 b, 18 b, 19 a).	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5183F4EC4BA1EC072920E26.taxon	distribution	Distribution: In the Great Balsam Mountains of western North Carolina at elevations above ~ 1000 m ASL, primarily in Haywood, Jackson, and Transylvania Counties, but possibly extending into western Buncombe and Henderson Counties (Fig. 4).	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5183F4EC4BA1EC072920E26.taxon	etymology	Etymology: From the Latin balsamum for “ balm, ” after the Fraser fir trees (Abies fraseri), known in the past as the “ she-balsam ” or “ balsam fir ” for which the Great Balsam Mountains are also named (Fig. 19 b), though now treated as distinct from the Balsam fir A. balsameus. Name is a Latin singular adjective in the nominative case, masculine gender. We suggest the common name “ Great Balsams ” Mountain Dusky Salamander. Notes: Comprises the ocoee B lineage defined by Kozak et al. (2005), Beamer & Lamb (2020), and Pyron et al. (2020, 2022 c). The fourth species in our taxonomy represents the resurrection of Desmognathus perlapsus Neill, 1950 for the ocoee C / D lineage, which shares hybrid non-lineal mitochondrial ancestry with the Balsam clade but is otherwise genealogically exclusive and topologically distinct from D. adatsihi and D. balsameus (see Pyron et al. 2022 c). The primary expansion of this taxon concept here is the inclusion of populations previously referred to D. conanti from the Chattahoochee River drainage in the Piedmont of Georgia and Alabama, as reported by Beamer & Lamb (2020) and corroborated by Pyron et al. (2022 c). The ocoee C lineage reported by Kozak et al. (2005) is a member of this species and exhibits reciprocal admixture with adjacent ocoee E and possibly ocoee F / G / H populations in the southern Nantahala Mountains (Pyron et al. 2022 c). The ocoee D lineage is otherwise genetically cohesive and comprises northern and southern phylogeographic sublineages with a broad hybrid zone across the Blue Ridge / Piedmont transition. Correspondingly, we offer a re-description of:	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5193F4AC4BA191F72C40EB6.taxon	materials_examined	Holotype: ERA-WTN 14150 (Fig. 2), collected 4 August 1950 by Wilfred T. Neill, type locality on a “ rocky outcropping on the western wall of Tallulah Gorge, near the town of Tallulah Falls, Rabun County, Georgia. ” The locality described is southeast of Tallulah Falls in Habersham County (Huheey in Valentine 1964), approximate coordinates 34.730, - 83.385. Type specimen not known to exist; Neill (1950) included a note that it was to be deposited in the FLMNH, but it was not listed by Christman in Gilbert (1974) or among the current holdings. Because the identity of the taxon is not in dispute, a neotype is not warranted. Paratypes: ERA-WTN 14151 – 66, from the type collection. Not known to exist.	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5193F4AC4BA191F72C40EB6.taxon	description	Description: After Neill (1950) and Valentine (1961), a robustly proportioned salamander with a round tail (or weakly keeled on the distal portion), four pairs of dorsal spots, and a well-defined color-pattern with wide black margins around the larval spots, white flecking on the upper surfaces, and whitish mottling on the sides and venter, putatively diagnosable from Desmognathus ocoee by larger adult body size (~ 22 – 56 mm SVL; n = 77), a longer and wider head, shorter limbs, vomerine teeth present in adult males, long slender parasphenoid tooth-patches, lessdefined dorsal color-pattern, and 4 vs. 5 – 6 pairs of spots between the axilla and groin. However, Valentine (1961) questioned the diagnostic utility of these characters; for instance, many specimens have 5 – 6 pairs of larval spots, and the size ranges overlap substantially. Additional work is needed to establish the morphometric description of this taxon. Some populations in the Cowee Mountains and Highlands Plateau also occasionally exhibit red, orange, or yellow cheek patches (Brimley 1928; Huheey 1966 a; Labanick 1983), differentiating them from all lineages other than D. ocoee E and D. imitator. Specimens from the southern part of the range in the Piedmont and adjacent Coastal Plain of Alabama and Georgia are typically considerably more melanic with an overall dark coloring and obscured pattern of paired larval spots, often with a partial keel on the tail.	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5193F4AC4BA191F72C40EB6.taxon	distribution	Distribution: Based on expanded genetic sampling, this species ranges from the Alarka and Cowee Mountains in western North Carolina between the Tuckasegee and Little Tennessee Rivers, the western headwater mountain streams of the Savannah River drainage in northwestern South Carolina, northeastern Georgia, and adjacent North Carolina, and the Chattahoochee River drainage in the Piedmont of Georgia and Alabama to the Fall Line and Uchee Creek (AL: Russell) in the adjacent Coastal Plain (Fig. 4).	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C5193F4AC4BA191F72C40EB6.taxon	etymology	Etymology: From the perfect passive participle of the Latin perlabor for “ slipping ” or “ gliding through ” as an adjective in the nominative singular, in reference to the lithe nature of the species when active in rock crevices, or in hand once captured. Previous common names include Tallulah Salamander (Schmidt 1953) and Cliffside Salamander (Conant et al. 1956). Neither is particularly appropriate; we suggest “ Chattooga Dusky Salamander. ” Notes: Comprises the ocoee C / D lineage defined by Kozak et al. (2005), Beamer & Lamb (2020), and Pyron et al. (2020, 2022 c). Synonymized with Desmognathus ocoee by Valentine (1961) and resurrected here with an expanded geographic range. Voluminous data on the biology of this species exist under the name D. ocoee but will need to be carefully disambiguated by reference to the originating populations; see extensive bibliographies in Valentine (1964) and Camp & Tilley (2005). The fifth and final species corresponds to a slight restriction of Valentine (1961) ’ s concept of D. ocoee (see Valentine 1964), including five geographically distinct genetic population segments. These are the ocoee E – H lineages of Kozak et al. (2005), along with the apalachicolae A 2 populations from the southernmost Blue Ridge first identified by Beamer & Lamb (2008) and named by Pyron et al. (2022 c). The ocoee H lineage occupies a restricted portion of the southern Cumberland Plateau in Tennessee; the G lineage ranges along an extensive portion of Sand Mountain / Walden Ridge in northeastern Alabama, northwestern Georgia, and central Tennessee (genetically identified as D. ocoee by Anderson & Tilley 2003); and the F lineage occurs in the Unicoi Mountains and adjacent ridges in eastern Tennessee and extreme western North Carolina. Together, these form a single genetic unit, the ocoee F / G / H lineage (Beamer & Lamb 2020; Pyron et al. 2020, 2022 c). The ocoee E lineage occurs in the Nantahala Mountains of western North Carolina and northern Georgia, and the apalachicolae A 2 lineage occupies the southern terminus of the Blue Ridge escarpment in north-central Georgia. These three groups are parapatric and replace each other geographically in the mountains of Georgia, North Carolina, and Tennessee, while also exhibiting admixture across their contact zones. Therefore, we conservatively treat them as a single species with at least three major phylogeographic lineages. Phylogenetic estimates suggest that D. apalachicolae is nested within this species, though population-genetic analyses indicate that it is genealogically exclusive, and the nested topological position may be an artifact of the complex history of diversification and reticulation in the group (Chan et al. 2020; Dolinay et al. 2021). Given its geographic and genetic distinctiveness, we treat D. apalachicolae as distinct from apalachicolae A 2, which we consider part of D. ocoee. We re-describe D. ocoee here as:	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C51D3F45C4BA19A272EB0F56.taxon	materials_examined	Holotype: USNM 128007 (field tag J. C. N. 1001; Fig. 1) by original designation; type locality “ on the surface and in crevices of cliffs at Ship’s Prow Rock, in Ocoee Gorge, beside U. S. Highway 64, nine miles airline west of Ducktown, in Polk County, Tennessee, ” collected 14 November 1948 by J. C. Nicholls, Jr. Paratypes: A series of 28 specimens including the holotype was reported, so ostensibly 27, including untraceable specimens in the “ J. C. Nicholls, Jr. collection ” and the “ S. C. Bishop collection, Rochester, New York, ” AMNH A- 54385 – 6, CM 29290 – 1, MCZ A- 26589 – 90 & A- 28307 (MCZ catalog notes received in “ Exch. Chicago N. H. Mus. ” — original number CNHM [now FMNH] 90035), FMNH (originally given as “ Chicago Natural History Museum, Chicago, Ill. ” — CNHM) 57313 – 4 & 90034 – 5 (FMNH 90035 now MCZ A- 28307), and untraceable specimens at “ Emory University, Emory University, Ga. ”	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C51D3F45C4BA19A272EB0F56.taxon	description	Description: A small (~ 15 – 54 mm SVL; n = 122) mountain dusky salamander exhibiting a wide variety of color patterns and ecomorphologies throughout its range. Diagnosable from Desmognathus abditus in the Cumberland Plateau by the presence of a modest keel on the distal portion of the tail and keratinized toe tips (Fig. 20; vs. absence of both, see Drukker et al. 2018). Usually diagnosable from D. adatsihi and D. balsameus by generally lacking a solid dorsal stripe (vs. often striped with straight, wavy, or undulating borders). Some individuals of the ocoee E lineage are diagnosable from all other species by the presence of red, yellow, or orange patches on the legs or very rarely both the legs and cheeks, and all species other than D. imitator and D. perlapsus by the presence of such patches on the cheeks. Neill (1950) suggested seven diagnostic characters differentiating D. ocoee from D. perlapsus, with D. ocoee being smaller, having a shorter and narrower head, longer limbs, no vomerine teeth in adult males, parasphenoid teeth in oblong patches, more-defined dorsal color pattern, and 5 – 6 pairs of spots (vs. 4) on the dorsum between the limb insertions. However, Valentine (1961) demonstrated exceptional variability in these traits in both species and questioned this diagnosis. Additional work is needed to diagnose this taxon morphometrically. Range: Three disjunct population segments (Fig. 4); one restricted to a small portion of the Cumberland Plateau between Sewanee and Orme, Tennessee in Franklin and Marion Counties; a second occurring broadly on the Cumberland Escarpment (Walden Ridge and Sand Mountain) of south-central Tennessee, extreme northwestern Georgia, and northeastern Alabama (see Anderson & Tilley 2003); and a third in the Nantahala, Unicoi, and southern Blue Ridge Mountains of western North Carolina, southeastern Tennessee, and north-central Georgia. Additional populations probably referable to this species likely remain to be discovered and sampled. We examined a historical collection (MCZ A- 143377 – 86) from Horse Cove, Rich Mountains (GA: Gilmer) which appears to be this species based on morphological gestalt and collected a single recent individual ourselves (Fig. 21). We have also received anecdotal reports from the Cohutta Mountains in Georgia (Fig. 4), due south of the type locality along the same general mountain range (S. P. Graham, pers. comm.), and their identity is of great interest.	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
039087A5C51D3F45C4BA19A272EB0F56.taxon	etymology	Etymology: Named after the type locality in Ocoee Gorge. Various informal sources report “ ocoee ” as a Cherokee name for the Passionflower, Passiflora incarnata (e. g., the state symbol of Tennessee; https: // www. tn. gov / about-tn / state-symbols. html). Name is a non-Latin noun used in apposition. The widely-used common name is “ Ocoee Salamander ” (Schmidt 1953). Notes: Consists of the D. ocoee concept of Tilley & Mahoney (1996) and Anderson & Tilley (2003) in part, the ocoee E – H lineages of Kozak et al. (2005) and Beamer and Lamb (2020), and the apalachicolae A 2 lineage of Beamer & Lamb (2008) as defined by Pyron et al. (2022 c). Future work may reach more subtle and complex taxonomic conclusions regarding the specific identity of the ocoee E and apalachicolae A 2 lineages. Extensive data on the biology of this species exist but will need to be carefully disambiguated from D. perlapsus by reference to geography; see bibliographies in Valentine (1964) and Camp & Tilley (2005).	en	Pyron, R. Alexander, Beamer, David A. (2022): Systematics of the Ocoee Salamander (Plethodontidae: Desmognathus ocoee), with description of two new species from the southern Blue Ridge Mountains. Zootaxa 5190 (2): 207-240, DOI: https://doi.org/10.11646/zootaxa.5190.2.3
