1 AMPHIBIA: CAUDATA: PLETHODONTIDAE Desmognathus monticola

Catalogue of American Amphibians and Incorrect attribution. See Comments. Reptiles 925 Salamandra phoca: Dunn 1923:39. Desmognathus phoca: Dunn 1926:73. Regester, K. J., C. G. Hoffman, E. R. Patter- Desmognathus fuscus fuscus: Netting son, and P. C. Timashenka. 2020. 1933a:105. Incorrect attribution. See Desmognathus monticola. Comments. Desmognathus phoca: Netting 1935:43. Incor- Desmognathus monticola Dunn rect attribution. See Comments. Seal Salamander Desmognathus phoca: Hibbard 1936:279. In- correct attribution. See Comments. Salamandra phoca Matthes 1855:273. Lapsus. Desmognathus phoca: Scharlinski 1939:57. See Comments. Incorrect attribution. See Comments. Desmognathus monticola Dunn 1916:73. Desmognathus phoca: Yoder 1940:91. Incor- Type locality, “Elk Lodge Lake, near Bre- rect attribution. See Comments. vard, [Transylvania County,] North Car- Desmognathus monticola: Grobman 1945:39. olina; altitude about 3000 feet.” Holotype, Desmognathus monticola jeffersoni: Hoffman National Museum of Natural History 1951:250. Type locality, “Saddle Hollow (USNM) 38313, an adult male, collected on Jarman’s Mountain, 2 miles west of 13 July 1908 by R. Tipping and E. Tipping Crozet, Albemarle County, Virginia (el- (not examined by authors). evation 1600 feet).” Holotype, USNM Desmognathus monticola Dunn 1918b:463. 126891, an adult, collected 11 March 1945

Figure 1. An adult Desmognathus monticola from Macon County, North Carolina. The blunt snout and bulging eyes that characterize the head give the salamander a seal-like appearance. Photo by Todd Pierson. 2

Map. Geographic distribution of Desmognathus monticola in the eastern United States. Gray shading indicates the range with a periphery defined by watershed boundaries. The white circle marks the type locality, the "×" marks the two closely adjacent fossil localities, and the black dots represent voucher specimens in major museum collections (n = 9921). Coordinates associated with specimens were obtained from iDigBio (idigbio.org; accessed 30 June 2017) and VertNet (vertnet.org; accessed 12 December 2017), and then verified using county-level distribution maps in state and regional publications (see Distribution and Comments sections). Note the isolated population in the western panhandle of Florida. A red dot in the inset map marks an introduced population in the Ozark Plateau of Arkansas. Account 925 3

by R. L. Hoffman (not examined by au- CONTENT. No subspecies are currently rec- thors). ognized, however, two subspecies were de- Desmognathus monticola monticola: Hoffman scribed (see Nomenclatural History). 1951:251. Desmognathus monticolus: Spight 1967:128. DESCRIPTION. Ova are nonpigmented Lapsus. and deposited in clusters of 13–40. Egg sus- Desmognathus monticola: Hulse et al. 2001:81. pension cords are relatively long and their Incorrect attribution. See Comments. attachment points are often visible. Four epi- Desmognathus monticola: Rissler and Tay- branchial placodes are associated with the lor 2003:201. Incorrect attribution. See embryonic pharyngeal arches. Hatchling lar- Comments. vae range from 11–20 mm total length (TL), Desmognathus monticola: Graziano and Reid lack balancers, and have narrow, cylindrical, 2006:6. Incorrect attribution. See Com- pointed digits (four on forelimbs, five on rear ments. limbs). Mandibular grooves bisect the lower Desmognathus monticola: Beamer and Lamb lip anteriorly, and the tail fin originates at the 2008:147. Incorrect attribution. See Com- tail-body junction. Gill rami of larvae are rel- ments. atively short and mound-like with 16–17 fili- Desmognathus (Desmognathus) monticola: form fimbriae bilaterally. The medium brown Dubois and Raffaëlli 2012:144. dorsum has four to five pairs of prominent, red dorsolateral spots between the limb in-

Figure 2. A juvenile Desmognathus monticola from Indiana County, Pennsylvania. The dorsum of a typical juvenile has four or five pairs of orange, reddish-orange, or chestnut-colored spots bordered by darker pigments. A pale line extends from the eye to the angle of the jaw, but this characteristic is often obscured by dark pigments in older individuals. Photo by Kurt Regester. 4 sertions. The underside of the tail is diffusely are obscured as the background color of the blotched. Larvae have keratinized toe tips and dorsum becomes darker with age. Mottling metamorphose at 35–50 mm TL. In young ju- and color patterns are highly variable among veniles, the four or five pairs of dorsal spots individuals, and are markedly reduced or ab- between the limb insertions are orange, red- sent in many populations. Albino specimens dish-orange, or chestnut-colored, and the have been reported (Brame 1962; Houtcoo- venter is whitish or pale. Dorsal spots are per 1979, 1981). invaded by darker pigment, and the ventral Desmognathus monticola is a moderately surface becomes increasingly gray with age. long and heavy-bodied salamander (45–80 The head of both adult and juvenile sal- mm snout–vent length [SVL], 75–150 mm amanders is proportionally large relative TL). The maximum size of adult males com- to body size. The head is characterized by a monly exceeds that of females. Adult males blunt snout, bulging eyes, and mottling with can be distinguished from females by the small brown and black spots on the dorsal presence of enlarged premaxillary teeth, pa- surface. A pale line extends from the eye to pillose cloacal lips, a slightly more flexuous the angle of the jaw. A tubercle in the anterior outline of the jaw, and an inconspicuous angle of the eye and a gular fold are present. mental gland. The testes are unpigmented. The rear limbs are proportionally longer and Vomerine teeth are not lost by males follow- more robust than the forelimbs. The relative ing sexual maturity. The vomerine teeth form lengths of the forelimb digits are Ieggs deposited by Desmognathus whereas three intercostal folds are typical folkertsi, Desmognathus marmoratus, and for adults. The proximal portion of the tail is Desmognathus quadramaculatus have short round. The distal two-thirds of the tail are lat- suspension cords and they appear tightly at- erally compressed and distinctly keeled. tached to the substrate. Published sources Dorsal and ventral colorations are clearly lack sufficient information for the positive demarcated. The venter has uniformly spaced identification of eggs from those of sympatric melanophores, appears pale, and lacks con- congeners. The identification of an attending spicuous mottling. A line of small, irregu- parent can serve for field indentification. -In larly shaped white spots is often present on formation on species-specific distributions the lower sides of the body between the fore and microhabitat preferences will generally and rear limbs. The dorsum is light brown or facilitate diagnoses of all life stages. grayish in background color and may have Larval Desmognathus monticola have ke- reticulated patterning. Dark brown or black ratinized toe tips and four or five dorsolateral mottling and worm-like markings are typical- spots are present on the dorsum. Larval life ly present; in most individuals, these mark- stages of Desmognathus abditus, Desmog- ings extend to the dorsolateral surfaces of the nathus carolinensis, Desmognathus imitator, body and dorsal surface of the tail. In the Blue Desmognathus ochrophaeus, Desmognathus Ridge Mountains physiographic province, ocoee, and Desmognathus orestes do not have dark dorsal markings may be reduced to scat- keratinized toe tips; the dorsum of those tered, round spots. Patterns of pigmentation species may be uniformly reddish to brown, Account 925 5

A

B

Figure 3. Larval and recently metamorphosed Desmognathus monticola from Oconee County, South Carolina. (A) The dorsum of a typical larva has four or five pairs of prominent dorsolateral spots between the limb insertions. Larvae possess relatively short gill rami and have cylindrical, pointed digits with ke- ratinized toe tips. (B) The spots of a recently metamorphosed individual are typically bordered by dark pigments. Photos by Todd Pierson. 6 patterned with 2–7 dorsolateral spots, or pat- tails that are round or oval in cross section terned with dorsolateral spots and a lateral for their entire length (Desmognathus aeneus, stripe. Larvae of several congeners typically Desmognathus abditus, Desmognathus car- have a greater number of dorsolateral spots, olinensis, Desmognathus imitator, Desmog- including Desmognathus auriculatus (7–9), nathus ochrophaeus, Desmognathus ocoee, Desmognathus conanti and Desmognathus Desmognathus orestes, Desmognathus organi, fuscus (5–13), Desmognathus folkertsi and and Desmognathus wrighti). Among conge- Desmognathus quadramaculatus (6–8), and ners with keeled tails, Desmognathus montico- Desmognathus welteri (5–8). Additional char- la is characterized by internal nasal openings acteristics can be used to diagnose larvae of that are circular. The head is not distinctly Desmognathus monticola from those of sym- flattened, possessing a gradual slope starting patric congeners. Desmognathus monticola at the eyes. The venter is uniformly colored does not have black stripes along the lateral white to light gray, and gray-black friction margins of the dorsolateral spots (present in pads cover the tips of the digits. Desmog- Desmognathus conanti and Desmognathus nathus marmoratus has slit-like internal nasal fuscus) or white spots associated with the lat- openings, a distinctly flattened head (slope eral line pores (present in Desmognathus folk- starting behind the eyes), and a dark gray to ertsi and Desmognathus quadramaculatus). black venter. Desmognathus quadramaculatus Desmognathus monticola has fewer costal and Desmognathus folkertsi also have darkly grooves (14) than Desmognathus santeetlah colored toe tips. However, they can be diag- (16), and a less prominent white line from nosed by their black bellies and two rows of the eye to angle of the jaw. Unlike the wide light, distinctly round spots along the lower head of Desmognathus monticola that appears lateral surface between the limbs. The tips of rounded in dorsal view, the relatively narrow the digits in Desmognathus auriculatus, Des- head of Desmognathus marmoratus appears mognathus conanti, Desmognathus fuscus, strongly pointed. Further, the gills of Desmog- and Desmognathus santeetlah lack friction nathus monticola are colored similarly to its pads and thus appear white or light gray; the brown body whereas the pale gills of Desmog- ventral surfaces of those species range from nathus marmoratus contrast markedly with light gray to gray-brown to black. Desmog- its darker body color. The gills ofDesmog - nathus welteri lacks bold markings on the nathus monticola possess 16–17 gill fimbriae dorsum and a distinct demarcation between per side, compared to a greater number of the dorsal and ventral colorations. The venter gill fimbriae inDesmognathus welteri (17–27) of Desmognathus welteri lacks uniformity, be- and Desmognathus auriculatus (22–40). In ing slightly mottled in juveniles and heavily contrast to the medium brown body color mottled with brown in older individuals. The and prominent red-orange spots on the dor- proximal third of the tail is evenly rounded sum of larval Desmognathus monticola, the in Desmognathus monticola whereas the base body of Desmognathus welteri is uniformly of the tail in Desmognathus welteri is laterally green to brown and the rows of dorsolateral compressed. dots are faint; the body of Desmognathus au- riculatus is darkly pigmented and the dorso- PHYLOGENETIC RELATIONSHIPS. Most lateral spots are pale. Desmognathus, including Desmognathus Adult Desmognathus monticola are rela- monticola, have an aquatic larval stage. How- tively large (7.5–15 cm TL) and have keeled ever, they are nested in a clade of direct-de- tails. Adults of several sympatric congeners veloping species indicating a major life-his- are generally smaller (≤10 cm TL) and have tory reversal from direct development to an Account 925 7

Figure 4. An adult Desmognathus monticola from Johnson County, Tennessee with eggs. The rear limbs are proportionally longer and more robust than the forelimbs. The proximal portion of the tail is round; the distal two-thirds of the tail are laterally compressed and distinctly keeled. Photo by Scott Bolick. aquatic larval stage (Chippindale et al. 2004). legheny Plateau range are relatively uniform Phaeognathus is the sister taxon to Desmog- in haplotype diversity and represent a lineage nathus, and Pygmy salamanders (Desmog- associated with a northward post-Pleistocene nathus wrighti and Desmognathus organi) are range expansion from the southern highlands sister to the clade including all other members of the Appalachian Mountains (Beamer and of Desmognathus. At least three sister taxa to Lamb 2008; Kozak et al. 2005; Rissler and Desmognathus monticola have been proposed Taylor 2003). Several southern populations based on mitochondrial DNA, nuclear DNA, occur in historical refugia, where greater than morphological, and reproductive character 50% of haplotype diversity is restricted to less data: Desmognathus apalachicolae (Titus and than 3% of the current distribution (Bonett Larson 1996), Desmognathus carolinensis et al. 2007). Haplotypes from an introduced (Pyron and Wiens 2011; Rissler and Taylor population in Arkansas are identical to those 2003), and Desmognathus ocoee (Beamer and from Lumpkin, Towns, Union, and White Lamb 2008; Kozak et al. 2005; Martin et al. counties in northeastern Georgia (Bonett et 2016; Shepard et al. 2016). al. 2007). Populations assigned to Desmognathus monticola typically form a well-supported PUBLISHED DESCRIPTIONS. Descrip- clade and have relatively low genetic variation tions were provided by Anonymous (1987), across much of their distribution. Populations Ashton and Ashton (1988), Barbour (1971), within the Appalachian Highlands and Al- Bartlett (1989), Bartlett and Bartlett (1999), 8

Beane et al. (2010), Behler and King (1979, Coborn n. d., 1996; H. Collins 1959, 1981; 1985, 1988), Bishop (1943, 1947, 1962, 1967, Dodd 2004; Folkerts 1986; D. Green et al. 1969, 1994), Camp (2008), Camp and Tilley 2013; N. Green and Pauley 1987; Hennessy (2005), Catlin (1984, 1992), Conant (1958, 2016; Herrmann 2000, 2001; Hulse et al. 1975), Conant and Collins (1991a, 1991b, 2001; J. Mitchell 1977a; J. Mitchell and Gib- 1998), Dodd (2004), Dunn (1916, 1917a, bons 2010; J. Mitchell et al. 2006; Pauley 2004; 1917b, 1918a, 1926, 1972), Folkerts (1986), Petranka 1998; Pingleton and Holbrook 2019; Freytag (1960, 1961, 1967), N. Green and Pau- Powell et al. 2016; Rimpp 1985; Shaffer 1991, ley (1987), Houtcooper (1979), Huheey and 1995; Shupe 2018; H. Smith 1978; Stanisze- Stupka (1967), Hulse et al. (2001), Klingel- wski 1995; Wyckoff and Niemiller 2011). Ad- höffer (1956), Martof et al. (1980), Matsui ditional relevant photographs documented (1993), Means (1978, 1992), J. Mitchell and habitat (King 1939) and predation (Bernar- Gibbons (2010), Mount (1975, 1996), Mount do and Yoke 2006; Lawhorn et al. 2017). et al. (1984), Murray (2018), Nerz (2011), Pe- Photographs of several cellular and ana- tranka (1998), Powell et al. (2016), Raffaëlli tomical characteristics have been published: (2007, 2013), Tilley and Huheey (2001), Val- femur histology (Laurin et al. 2009), mental entine (1963, 1967), Wallays (2006) and Wy- gland histology (Sever 1976), nasal gland ckoff and Niemiller (2011). Descriptions of morphology (Dawley 2017), nuclear mate- eggs and larvae were provided by Altig and rials (Hally et al. 1986), spermatozoa mor- McDiarmid (2015). phology (Wortham et al. 1982), toe mor- phology and histology (Caldwell and Trauth ILLUSTRATIONS. Various life stages of 1979), and tooth morphology (Matson et al. Desmognathus monticola have been photo- 2010; Means 1974). graphed or illustrated: eggs (Petranka 1998), Species characteristics have been illus- larvae (Altig and McDiarmid 2015; Ashton trated in line drawings of the adult dorsal and Ashton 1988; Camp 2008; Dodd 2003; pattern (Allyn 1952, 1956, 1961, 1977; N. J. Mitchell and Gibbons 2010; Noble 1931, Green 1969), feeding (Lombard and Wake 1954; Petranka 1998; Wyckoff and Niemiller 1977; Pough et al. 1998, 2016); skull (Cald- 2011), juveniles (Conant 1958, 1975; Conant well 1977; Means 1974; Rubenstein 1971), and Collins 1991a, 1991b, 1998; Powell et al. spermatophore (Noble and Weber 1929; Or- 2016), adults in black-and-white (Freytag gan and Lowenthal 1963), tail cross section 1967; Harrison 1950a, 1950b; Hoffman 1951; (Powell et al. 1998, 2012, 2019), and teeth Krysko et al. 2011; Mansueti 1947; Means (Caldwell 1977; Caldwell and Trauth 1979; 1978, 1992; Mount 1975, 1996; Nerz 2011; Noble 1927). Obst et al. 1984, 1988; Petranka 1998), and adults in color (AmphibiaWeb 2019; Ash- DISTRIBUTION. Desmognathus montico- ton and Ashton 1988; Barbour 1971; Bartlett la has a broad geographic range through the and Bartlett 1999, 2006; Beane et al. 2010; W. central and southern Appalachian Mountains Brown et al. 2003; Camp 2008; Cassie 1999; of North America. Most populations occur

Figure 5 (facing page). Variation in coloration and pattern in metamorphosed Desmognathus monti- cola: (A) Graham County, North Carolina, (B) Macon County, North Carolina, (C) Macon County, North Carolina, (D) Caldwell County, North Carolina, (E), Indiana County, Pennsylvania, (F) Graham County, North Carolina, (G) Indiana County, Pennsylvania, and (H) Oconee County, South Carolina. The dorsal pigmentation pattern of the typical adult is increasingly obscured with age by a dark background color- ation (G–H). Photos by Carter Ricks (A, B), Todd Pierson (C, D, H), Chris Bortz (E), Stephen Tilley (F), and Edwin Patterson (G). Account 925 9

A B

C D

E F

G H 10 at elevations between ~400–1400 m, but as objects than juveniles. Small individuals are high as 1700 m. The continuous portion of frequently found in the interstitial spaces of the range extends from western Pennsyl- streambed substrates and among layers of vania southward to West Virginia, western leaf litter. Areas adjacent to seeps and streams Maryland and western Virginia, eastern Ken- are also used by salamanders; these include tucky and eastern Tennessee, western North streamside burrows, crevices on the faces of Carolina and northwestern South Carolina, moist cliffs, and forest leaf litter. During and northern and central Alabama, and northern following periods of precipitation, individu- Georgia, USA. Disjunct populations occur on als have been observed 1–2 m above ground, the coastal plain of southern Alabama and positioned upon the trunks of trees. the westernmost region (Escambia County) of the panhandle of Florida, USA. An intro- FOSSIL RECORD. Fossils dated to the Pleis- duced population occurs on the Ozark Pla- tocene Epoch have been reported from two teau of Arkansas, USA (Bonett et al. 2007), caves in Pendleton County, West Virginia where salamanders were most likely relocated (Holman 1995, 2006). A single vertebra from for use as fish bait. Hamilton Cave was described and associated Distribution maps were provided by with the Irvingtonian North American Land Ashton (1978), Barbour (1971), Bishop Mammal Age (NALMA) (Holman and Grady (1943, 1947, 1962, 1967, 1969, 1994), Cald- 1989). Two fossils of 13 and 5 vertebrae were well (1977), Camp (2008), Camp and Tilley described from New Trout Cave and associ- (2005), Conant (1958, 1975), Conant and ated with the Rancholabrean NALMA (Hol- Collins (1991a, 1991b, 1998), Dunn (1917b, man and Grady 1987). The convex or flat- 1926, 1972), Freytag (1974, 1984), D. Green et tened posterior end of the vertebral neural al. (2013), Grobman (1945), Hoffman (1951), arch was used for species-level identification J. Mitchell and Gibbons (2010), Petranka (Holman 2006). (1998), Powell et al. (2016), H. Smith (1978), and Stevenson (1976). Regional maps were PERTINENT LITERATURE. Published ref- provided by Beane et al. (2010), Camp (2008), erences to the species are listed by topic: acid- Folkerts (1986) Franz (1972), N. Green and ification (Little et al. 1991; J. Mitchell 1999; Pauley (1987), Harris (1969, 1975), Huheey Roudebush 1988, 1989), adult habitat (Bick- (1966), Hulse et al. (2001), McCoy (1982), ert 1969; Camp 1997a; Hairston 1949; Klee- Means (1978, 1992), J. Mitchell (1977a), J. berger 1985; Krzysik 1979; Organ 1961; Tilley Mitchell and Gibbons (2010), J. Mitchell and 1973, 1980), aggression (Camp 2003; Jackson Reay (1999), Shaffer (1991, 1995), Tobey and Cupp 1997; Keen and Sharp 1982, 1984; (1985), Williamson and Moulis (1979), and Nelson 1979; Southerland 1986a), albinism Wyckoff and Niemiller (2011). A map of re- (Brame 1962), antipredator behavior (Bro- cords in the Great Smoky Mountain National die 1978; Dodd and Brodie 1976; Fauth 1996; Park, Tennessee was given by Huheey (1966). Hensel and Brodie 1976; Howard and Brodie Desmognathus monticola inhabits mesic 1973), bait trade (Cogger et al. 2002, 2004; hardwood forests. Habitats are characterized Freytag 1974, 1984; Jensen and Waters 1999, by seeps and low order streams providing 2000; Martof 1953), behavior (Forester 1978; cool, well-aerated water and abundant cover. Gordon et al. 1962; Mabry and Verrell 2004; Metamorphosed individuals are found un- Mathis et al. 1995; Rissler et al. 2004; Shupe der woody debris and stones located outside 2018; Soler 1950), biochemical genetics the main flow of water, and typically below (Beamer 2015; Beamer and Lamb 2008; Ber- headwaters of seeps. Adults use larger cover nardo et al. 2007; Bonett et al. 2007; Casey Account 925 11

2002a; Chippindale et al. 2004; Goin and Walls 1989; Martof and Rose 1962; Pough et Goin 1971; Integrated Taxonomic Informa- al. 1998; Resetarits 1991; Schwartz 1954, tion System 2017; Kozak 2005; Kozak et al. 1957; Sever et al. 1976; Southerland 1985; 2005; Larson et al. 2003; Martin 2015; NCBI Spohn and Sattler 1990), competition (Bran- 2017; Pyron and Wiens 2011; Rissler 2000; don and Huheey 1971; Hairston 1981; Hof- Rissler and Taylor 2003; Rissler et al. 2004; facker et al. 2018; Kleeberger 1983, 1984), Rovito 2009; Tilley 1997, 2016; Tilley et al. conservation (Bailey et al. 2006; Millsap et al. 2008a; Titus and Larson 1996), biogeogra- 1990), costal grooves (Highton 1957), criti- phy (Redmond 1991), biomass (Crawford cal thermal maxima (Hutchison 1961; Layne and Peterman 2013; Hairston 1987; Woodall 1984; Layne and Claussen 1987; Markle and and Wallace 1972), blood (Davic and Gallati Kozak 2018; Sealander and Adolph 1966; 1979; Reynolds and Pickard 1973; Vernberg Werber and Benton 1967), density (Bruce 1955; Villolobos et al. 1968), body size 1995; Kleeberger 1984), diel activity (Bran- (Beachy and Bruce 1999; Bruce 1987; Davic don and Huheey 1975; Gordon et al. 1962; 1983), body temperature (Brattstrom 1963), Hairston 1949, 1986; Hall and Taylor 1988; breeding season (Organ 1961; Tilley 1968, Holland and Taylor 1989; Keen 1982; Keen 1973, 1977), cell size (Roth et al. 1994), chem- and Reed 1984a, 1984b; 1985a, 1985b, 1985c; ical cues (Brown and Martof 1966; Forester et Lynnette and Andreadis 2000; Shealy 1975; al. 2000; Jacobs and Taylor 1992; Kats and Sievert and Andreadis 2002), diet and forag- Dill 1998; Moberly 2012; Roudebush and ing (Bernardo 2002; Brandon 1965; Brandon Taylor 1986a, 1987b; Southerland 1986a; Stef- and Huheey 1971; W. Brown et al. 2003; fen 2015), citation for original description Camp 1997b; Donavan 1972; Duncan 1970; (Beltz 1995), climate change (Bernardo and Felix and Pauley 2006; Franz 1966; Gorsuch Spotila 2006; Bernardo et al. 2007; Dodd and Owen 2014; Hairston 1949, 1986; Huheey 1997; Grant et al. 2010; Grover 2000; Hamed 1966; Kleeberger 1984, 1985; Krzysik 1979; 2014; Hoffacker et al. 2018; Keitzer and Go- Kuzmin 1992; Pough et al. 1998, 2001, 2004, forth 2013; Markle and Kozak 2018; Mila- 2016; Shealy 1975), dispersal (Bruce 1986; novich et al. 2015), cloacal anatomy (Sever Grant et al. 2010), distribution (Adams et al. 1994a; Sever and Trauth 1989, 1990), clutch 1996; Ashton and Ashton 1988; Ballinger and attendance (Bannikov 1969; Bruce 1996; Lynch 1983; Bartlett and Bartlett 1999; Bish- Camp 1997a; Nussbaum 1985, 1987, 2003; op 1924, 1925, 1926, 1928; Blair et al. 1957, Organ 1961; Pope 1924), clutch size (Bruce 1968; Breder and Breder 1923; Chamberlain 1990b, 1996; Camp 1997a; Folkerts 1968; Or- 1928; H. Collins 1959; Cooper 1948, 1960a, gan 1961; Pope 1924), common and scientif- 1960b, 1961, 1965; Duellman and Sweet 1999; ic name checklists (Banks et al. 1987; J. Col- Dury and Williams 1933; Franz 1972; Gentry lins 1990, 1997; J. Collins and Taggart 2002, 1955; N. Green 1969; Harris 1969, 1975; Hoff- 2009; J. Collins et al. 1978, 1982; Comité sur man 1945; Hoffman and Kleinpeter 1948; les Français Standardisés 2012; Conant et al. Huheey 1966; Jackson 1944; Jopson 1984; 1956a, 1956b; Crother et al. 2001; Fouquette King 1939; Manis et al. 2005; Martof and Rose and Dubois 2014; Freytag 1974, 1984; High- 1962; McCauley and East 1940; McClure ton et al. 2001, 2017; Hutchins et al. 2003; 1931; Means and Longden 1970; Meinz 1975; Schmidt 1953, 1954; Stejneger and Barbour Meshaka et al. 2008; J. Mitchell and Reay 1917, 1923, 1933, 1939, 1943; Tilley et al. 1999; Mount 1975, 1996; Neill 1941, 1954; 2008b, 2012), communities (Beachy 1994; Netting 1932c; Nicholls 1950; Patterson and Bishop 1926; Bruce 1972, 1985, 1986, 2011; Regester 2016; Penney 1952; Pickens 1927; Dunn 1924; Hairston 1980a, 1987; Jaeger and Pope 1928; Redmond 1980; Rose and Dobie 12

1963; Sever 1984; Thompson 1982; Thurow (Bourne 2015; Crawford 2007; Dodd 1997; 1957; Tilley and Harrison 1969; Tobey 1985; Gingerich 2009; Gordon et al. 1962; L. Green Welter and Carr 1939; Williamson and Mou- 2006; Hamilton 2002; S. Jones 2010; Kong lis 1979, 1994b; Wilson and Friddle 1950; Zell 2006; Matthews et al. 2010; McDonald 2001; 2017), eggs (Altig and McDiarmid 2015; Roudebush 1988, 1989; Southerland and Brady 1924; Bruce 1989a, 1989b, 1990a, 1996; Stranko 2008; Walz 2002; Ward 2005; Wat- Camp 1997a; Dodd 2003; Folkerts 1968; kins-Colwell and Leenders 2006; Whiteleath- Nussbaum 1987, 2003; Organ 1961; Pope er 2001; Wood and Williams 2013a), habitat 1924), electrophoresis (Hinderstein 1969, selection (Hairston 1987; Keen 1985), home 1971b; Shontz 1968), evolution (Camp et al. range (Hardin et al. 1969; Kleeberger 1984, 2000; Collazo 1990; Eastman 2010; Morescal- 1985; MacGregor 1982b), integument chi 1975; Noble 1927; Sever et al. 2016; Shep- (Houck and Sever 1994; McMahon and Pav ard et al. 2016; Tipton-Jones and Pyles 1994; 1982; Means and Karlin 1989), interspecific Wake 1964, 1966; White et. al. 2000), field interactions (Bruce 2008; Carr and Taylor guides (Ballinger and Lynch 1983; Bartlett 1983, 1985; Davic 1979; Fauth 1998; Grover and Bartlett 1999, 2006; Behler and King 2000; Hairston 1983, 1986; Juterbock 1977a; 1979, 1985, 1988; Bishop 1943, 1947, 1962, Keen 1982, 1985; Keen and Sharp 1984; Klee- 1967, 1969, 1994; Cassie 1999; Cochran and berger 1983, 1984; Mathis et al. 1995; Organ Goin 1970; Conant 1958, 1975; Conant and 1961; Roudebush and Taylor 1986b, 1987a; Collins 1991a, 1991b, 1998; Hipes et al. 2001; Southerland 1984, 1986a, 1986b, 1986c; Til- Pauley 2004, 2013; Powell et al. 2016; H. ley 1973; White et al. 2000), interspecific Smith 1978; Thompson 1982), genome spacing (Fauth et al. 1992; Folkerts 1968; (Bachmann 1970; Hally et al. 1985, 1986; Lar- Grover 2000; Hairston 1949; Karlin and Gut- son 1984; Morescalchi 1975; NCBI 2017; Ses- tman 1979; Keen 1982; Keen and Sharp 1984; sions and Larson 1987), general ecology Krzysik 1979; Organ 1961; Shealy 1968, 1975; (AmphibiaWeb 2019; Arnold 2000; Barbour Southerland 1986a), intraspecific spacing 1953; Bond 2007; Brandon 1966; Brown 1992; (Anonymous 1989; Camp and Lee 1996; Col- Cochran 1961a, 1968; Cogger et al. 2002, ley et al. 1989; Hairston 1986; Keen and Col- 2004; H. Collins 1959; Crockett 2001; Dodd ley 1987; Krzysik and Miller 1979; Souther- 2003; Duellman and Trueb 1986, 1994; Gib- land 1986a, 1986b, 1986c), introduced bons and Buhlmann 2001; D. Green et al. populations (Bonett et al. 2007; C. Bush et al. 2013; N. Green 1937; Hairston 1973; Leviton 2017; Connior et al. 2013), inventories 1972; Mattison 2005; McCoy 1982; Rimpp (Anonymous 1981, 1987, 1998; Buhlmann 1985; Schneider 2010; Svancara 2010; Tilley 1987; F. Bush 1959; Colwell 1983; Dodd 2003; 1973; Vitt and Caldwell 2009, 2014; Wallace Frantz 1992; Jung et al. 2000, 2005; Neff 2017; et al. 1992; Wells 2007; Zug 1993; Zug et al. Netting and Wilson 1940; Rosenberg 1978; 2001), growth rate (Bruce 1989a, 2016a, Sattler and Gibson 2018; Simpson 2013; Tu- 2016b; Castanet et al. 1996; Hally et al. 1985), berville et al. 2005; Windsor 1931, 1932), ju- gut fungus (Tills 1977), habitat (Bailey et al. venile habitat (Anonymous 1989; Brandon 2006; Bruce 1987; H. Collins 1981; Cooper and Huheey 1971; Colley et al. 1989; Folkerts 1960b, 1961; Eaton 1954; Ford et al. 2002b; 1968; Hairston 1986; Keen and Colley 1987; Grant et al. 2005; Grover 1996; Jackson 1944; Krzysik and Miller 1979; Shealy 1975), keys Means 2000; Metts et al. 2001; J. Mitchell et al. to adults (Ashton 1978; Barbour 1971; Blair 2006; Newman 1955; Noble and Evans 1932; 1957, 1968; Brimley 1926, 1944; Chermock Petranka and Smith 2005; Resetarits 1997; 1952; Dunn 1917b, 1918a, 1926, 1972; Feuer Sattler and Gibson 2018), habitat change 1967; N. Green 1951, 1969; MacGregor Account 925 13

1982b; Martof 1956; Netting 1935; Powell et (Brame 1972; Dubois and Raffaëlli 2012; al. 1998, 2012, 2019; Stevenson 1976; Thomp- Dunn 1916, 1923; Grobman 1945; Sokolov son 1982), keys to larvae (Altig and Ireland 1988; Zhao et al. 1993, 1998), numeric codes 1984; Altig and McDiarmid 2015; Ireland (Brame et al. 1978), nutrient cycling (Drum- 1981), key to skull (Caldwell 1977), land use heller 1979; Keitzer and Goforth 2013; Mila- (Ash and Bruce 1994; Brannon and Purvis novich et al. 2015; Stiffler 1993, 2005; Wood- 2008; Crawford and Semlitsch 2007, 2008; all 1972), oocyte counts (Bruce 1995, 1996; Ford et al. 2002a; Knapp et al. 2003; Mackey Bruce and Hairston 1990; Tilley 1968), ooph- et al. 2014; Matthews et al. 2010; Moseley et agy (Camp 1997a), osteology (Caldwell 1977; al. 2008; W. Peterman 2008; Peterman et al. Castanet et al. 2003; Juterbock 1977b, 1978a; 2011; Petranka and Smith 2005; Petranka et Laurin et al. 2004, 2009; Wake 1964, 1966), al. 1993; Price et al. 2016, 2018; Stranko et al. pathogens (Davidson and Chambers 2011; 2010; Surasinghe 2013; Surasinghe and Bald- Hossack et al. 2010; Keitzer et al. 2011; win 2015; Ward et al. 2008; Williams 2003; McAlpine 1996; Rothermel et al. 2008; Williams et al. 2017; Wood and Williams Spaulding et al. 2018; Timpe et al. 2008; 2013b), larvae (Altig and McDiarmid 2015; Vazquez and Rothermel 2007; Vazquez et al. Eaton 1956), larval behavior (Orr and Maple 2009), parasites (Baker 1987; Baker et al. 1978), larval development (Bruce 1989b, 1987; Cheng 1958; Dunbar and Moore 1979; 1990b, 1995, 2019; Bruce and Hairston 1990; Dyer and Brandon 1973; Goater 2000; Goater Bruce and Hally 1986; Dodd 2004; Folkerts et al. 1987; Jones 1987; Joy et al. 1993; Rankin 1968; Juterbock 1984; Organ 1961; Ruben- 1937a, 1937b, 1938; Sellers et al. 1981), phy- stein 1969, 1971), life history evolution (Ar- logeography (Bernardo et al. 2007; Bonett et nold 2000; Arnold et al. 2017; Bonett and al. 2007; Casey 2002a, 2002b; Mabry and Var- Blair 2017; Bruce 1989b, 1996, 2005, 2007, rell 1997; Martin 2015; Martin et al. 2016; 2009, 2010, 2013, 2019; Bruce and Hairston Spight 1967), physiology (Beckenbach 1975; 1990; Bruce and Hally 1986; Chippindale et Bernardo et al. 2007; Brown et al. 1979; Lanza al. 2004; Duellman 2007; Hairston 1973; 1959; Spotila 1972; Vernberg 1955; Whitford Houtcooper 1981; MacGregor 1982a; Steb- and Hutchison 1965; Wittle 1983), popular bins and Cohen 1995; Tilley 1980; Tilley and accounts (Allyn 1952, 1956, 1961, 1977; Bernardo 1993), locomotion (Edwards Anonymous 1976b; N. Green 1943; Mansueti 1976), longevity (Bowler 1977; Bruce 1990a; 1947; J. Mitchell 1977a; Morris 1945, 1957, Bruce and Hairston 1990; Bruce et al. 2000, 1974; Netting 1933b), population sampling 2002; Castanet et al. 1996; Snider and Bowler (Bailey et al. 2004a; Graeter et al. 2013; Gro- 1992), marking (Kinkead et al. 2006; S. ver 2006; Hyde and Simons 2001; Jaeger et al. Mitchell et al. 2017; Perry et al. 2011), moni- 2016; Mackey et al. 2010; Marsh 2009; Pauley toring (Dodd et al. 2001; Grover 2006; Welsh and Little 1998; Rubio 1968), population and Droege 2001), morphology (Caldwell trends (Dixon and Pechmann 2005; Hairston and Trauth 1979; Dawley 2017; Dunn 1924; 1996; Hairston and Wiley 1993; Tipton-Jones Fowler and Dunn 1917; Highton 1961; Hin- and Pyles 1994), predation (Bernardo and derstein 1971a; Means 1972, 1974; Pingleton Yoke 2006; Bruce 1979; Hairston 1980b, 1986; and Holbrook 2019; Rehorek et al. 2013; Roth Jaeger et al. 1998; Peterman 2007; Resetarits and Walkowiak 2015; Sever 1974, 1983; Wake 1991; Tilley 1968), regulations on collection et al. 1988), museum collections (Cochran (Bartlett 1986), reproduction (Arnold 1972, 1961b; Dunn 1918b; N. Green 1949; Hoffman 1977; Brady 1924; Bruce 1989b, 1990b, 1995, and Mitchell 1994; [Wiesenberg] 1976; Wil- 1996, 2018; Bruce and Hairston 1990; Dun- liamson and Moulis 1994a), nomenclature can 1970; Organ 1961; Pope 1924; Tilley 1968; 14

Tilley and Tinkle 1968), seasonal activity Scott 1996; Redmond et al. 1990; Richmond (Shealy 1975), sexual behavior (Arnold 1972; and Boggess 1941; Richmond and McCoy Arnold et al. 2017; Brock and Verrell 1994; 1965; Roble and Hobson 1995; Rubenstein Keen and Reed 1985a; Mabry 2004; Mabry 1968; Sattler 1995; Scott and Redmond 2002; and Verrell 2003; Noble 1931, 1954; Noble A. Smith 1945; Tilley et al. 2000; Virginia and Brady 1930; Organ 1961; Pough et al. Herpetological Society 1968; Witt 1993; Wolf 2001, 2004; Verrell 1999; Verrell and Mabry 1981; Young 1993), subspecies (Hoffman 2000), sexual dimorphism (Bruce 1993; Cas- 1951), survivorship (Bruce 1990a, 1995; Du- tanet et al. 1996, Noble 1929, 1931, 1954), ellman and Trueb 1986, 1994; Organ 1961; sexual maturity (Bruce 1989b, 1990b, 1995; Porter 1972; Tilley 1968), systematics (Cald- Bruce et al. 2000, 2002; Castanet et al. 1996; well and Folkerts 1976; Hess 2016; Hess and Juterbock 1978b; Means and Longden 1970; Townsend 2017), tail autonomy (Wake and Organ 1961), spermatheca (Sever 1994b), Dresner 1967), taxonomic references spermatophore (Noble and Weber 1929; Or- (BoldSystems 2019; Frank and Ramus 1995; gan and Lowenthal 1963), spermatozoa Frost 1985, 2017; Frost et al. 2006; Harding (Barker and Baker 1970; Goin and Goin 1962; 1983; Integrated Taxonomic Information Sys- Scheltinga and Jamieson 2003; Sever 2000; tem 2017; NCBI 2017), territories (Keen and Sever and Hamlett 1998; Wortham et al. Reed 1984a, 1984b, 1985a, 1985b; Keen and 1977), state and provincial publications Sharp 1984), toxicology (Raimondo et al. (Anonymous 1976a, 2017; Camp et al. n.d.; 2003), vision (Caldwell 1980, Roth et al. Hoffman 1983, 1985; Hoffman and Klein- 1998), and worldwide checklists (Brame peter 1948; Hutchison 1956; J. Mitchell 1981; 1967; Gorham 1974; D. Mitchell 2017). Moriarty and Bauer 2000), state and regional checklists and bibliographies (Arment 2005; ETYMOLOGY. Desmognathus is derived Bailey et al. 2004b; Barbour 1950, 1957; Beltz from the Greek words desmos, δεσμός, (‘lig- 1989; Brimley 1926, 1939, 1944; Burger ament’) and gnathos, γνάθος, (‘jaw’). The 1958a, 1958b, 1959; Chambers 2006; DePoe generic name is descriptive of the large at- et al. 1961; Driver 1942; Dunn 1920; Echter- lanto-mandibular ligaments that are man- nacht 1980; Echternacht and Harris 1993; ifest, characteristic bilateral protuberances Enge and Dodd 1986, 1992; Fergus 2000, over the adult mandibular joint. The specif- 2003; Garriock et al. 1996; Garton et al. 1993; ic epithet monticola is a Latin word meaning Gentry 1955; N. Green 1936, 1937, 1948, “mountain-dwelling” and refers to the gener- 1954, 1961, 1963, 1964; N. Green and Brant al geographic range of the species. 1966; N. Green and Dowler 1966; Hayslett 1992; Johnson 1970; Lovich 1997; MacGregor NOMENCLATURAL HISTORY. Desmog- 1974, 1982b; McCoy 1986, 1989a, 1989b, nathus monticola was the first species de- 1992; Meshaka et al. 2012; J. Mitchell 1977b, scribed by zoologist Emmett R. Dunn and 1981; J. Mitchell and Pague 1984; Moler 1988, appears in his research between 1916 and 1990, 1991, 1999; Montanucci 2006; Neill 1920. Dunn (1923) later recognized that Mat- 1949; Netting 1931, 1932a, 1932b, 1933a, thes (1885) had named a similar salamander 1934a, 1934b, 1936a, 1936b, 1939a, 1939b, Salamandra phoca and suggested the name 1946a, 1946b, 1947, 1949, 1950; Netting and Desmognathus phoca. Authors used Desmog- Orton 1950; Netting and Richmond n.d., nathus phoca until 1943. Grobman (1945:p. 1955, 1970, 1974; Penney 1952; Pinder and 40) clarified the identity, distribution, and Greenlee 1999; Rappole 2007; Redmond and nomenclature of the species; he considered Account 925 15

“Salamandra phoca Matthes a synonym of a typically colored specimen of [Desmog- Desmognathus fuscus fuscus (Rafinesque)” nathus] welteri, and is not [Desmognathus] and “Desmognathus monticola Dunn as the monticola” (Juterbock 1984). valid name for the salamander currently recognized under the name Desmognathus COMMENTS. Invalid distribution records phoca (Matthes)”. Several authors continued included those associated with Desmog- to use Desmognathus phoca in subsequent nathus conanti in Monroe County, Alabama articles (Cheng 1958; Nicholls 1950; Penney (Beamer and Lamb 2008; Rissler and Tay- 1952; Vernberg 1955). lor 2003), Desmognathus fuscus in Blair and Additional English names and scientific Clearfield counties, Pennsylvania (Hulse et names included in literature are Appalachian al. 2001; Netting 1933a, 1935; Yoder 1940), Seal Salamander (Desmognathus monticola and Desmognathus fuscus in Adams County, monticola: Barbour 1971; Behler and King Ohio (Graziano and Reid 2006; Matson et al. 1979, 1985, 1988; Brame et al. 1978; Burger 2010). A specimen from Edmonson County, 1958a; Chermock 1952; J. Collins et al. 1978, Kentucky (MCZ [Museum of Comparative 1982; Conant 1975; Conant et al. 1956a, Zoology] 2230) was noted by Dunn (1918b) 1956b; Cooper 1960a, 1965; DePoe et al. likely in error. The same specimen was later 1961; Franz 1966; N. Green 1969; N. Green included in a list of salamanders examined and Dowler 1966; Harris 1969, 1975; Hutchi- by Dunn (1926, 1972), along with reference son 1961; McCoy 1982, 1986, 1992; Organ to the type specimen of Desmognathus phoca 1961; Organ and Lowenthal 1963; Ruben- (=Desmognathus fuscus) from Taylor’s Creek stein 1969; Seehorn 1982; Shaffer 1991, 1995; (Hamilton County, Ohio) near Newport, Virginia Herpetological Society 1968; Wil- Kentucky (Matthes 1855). When Grobman liamson and Moulis 1979; and Desmognathus (1945) clarified the identity of Desmognathus monticola: Gingerich 2009; Joy et al. 1993; monticola, he noted a second invalid record Kong 2006; Ward 2005; Ward et al. 2008), from Edmonson County, Kentucky (Hibbard Blue Ridge Seal Salamander (Desmognathus 1936) and rejected an additional record from monticola jeffersoni: Schmidt 1953), Moun- Norfolk County, Virginia (Scharlinski 1939). tain Desmognath (Desmognathus phoca: Desmognathus monticola is relative- Brimley 1926), Seal Salamander (Desmog- ly abundant throughout most of its range. nathus monticola monticola: Barbour 1957; However, this species is adversely affected Martof 1956; D. Mitchell 2017; Moler 1988; by factors associated with the degradation Schmidt 1953; Wrobel 2004), Virginia Seal of lotic systems and adjacent terrestrial hab- Salamander (Desmognathus monticola jef- itats. Factors that have been examined in fersoni: Behler and King 1979, 1985, 1988; this context include decreased water quali- Brame et al. 1978; Burger 1958a; J. Collins ty, increased sedimentation, edge and road et al. 1978, 1982; Conant 1975; Conant et al. effects, mountaintop removal mining, and 1956a, 1956b; Harris 1969, 1975; Seehorn valley filling (Price et al. 2016, 2018; Ward 1982; Virginia Herpetological Society 1968), et al. 2008). Terrestrial habitat quality is ad- and Wood Puppy (Desmognathus monticola: versely impacted by clear-cut, even-aged, and Breder and Breder 1923). partial cut timber harvests in riparian zones and forest stands (Ash and Bruce 1994; Mo- REMARKS. The illustration of Desmog- seley et al. 2008; Petranka et al. 1993; Suras- nathus phoca (=monticola) used by Bishop inghe and Baldwin 2015). For example, ter- (1943, 1947, 1962, 1967, 1969, 1994) “is of restrial habitat use by salamanders decreased 16 following a riparian timber harvest; further, ly narrow range of environmental tempera- relative abundance of salamanders remained tures, so it is among amphibians least likely depressed in even-aged forest stands for up to be impacted by effects of climate change to 40 years (Crawford and Semlitsch 2008; (Bernardo et al. 2007). Larval life stages of Peterman et al. 2011). Management plans re- salamanders are a large standing stock of ni- quire an overall stream buffer of at least 100 trogen, phosphorus, and calcium in headwa- m to retain essential habitat for terrestrial life ter Appalachian streams, where they excrete stages using riparian zones (Crawford and limiting nutrients in stream nutrient cycles Semlitsch 2007). (Keitzer and Goforth 2013; Milanovich et al. Desmognathus monticola is found in the 2015). Desmognathine salamanders such as salamander bait trade. Sixty-seven percent of Desmognathus monticola are dominant verte- ‘spring lizards’ sold in northern Georgia bait brate consumers in most headwater streams shops were Desmognathus monticola (Jensen and are an important component of lotic sys- and Waters 1999, 2000). The relocation of tems and adjacent habitats. these salamanders to uninhabited areas has resulted in at least one introduced population ACKNOWLEDGMENTS. We thank Ginger in Arkansas (Bonett et al. 2007). Surveillance McGiffin for her valuable assistance in locat- efforts and detailed studies on the ecopath- ing many of the source materials, and Ronald ology of fungal (Batrachochytrium dendro- Bonett for his helpful comments on phylo- batidis, Batrachochytrium salamandrivorans) genetics. We thank Taylor Braunagel, Emma and viral pathogens (Ranavirus) are need- Carter, and Jenna Stragand for retrieving and ed. Batrachochytrium dendrobatidis was not organizing documents. Addison Wynn pro- detected in populations in Georgia, North vided images of the original ledgers associat- Carolina, and Tennessee (Keitzer et al. 2011; ed with holotype specimens. Rothermel et al. 2008; Timpe et al. 2008), and prevalence in the central Appalachians LITERATURE CITED of Maryland and Virginia was relatively low Adams, H. S., M. S. Hayslett, and C. Hob- (Kozak et al. 2005). Salamanders experimen- son. 1996. Salamander diversity and tally infected with Batrachochytrium dendro- abundance along Buck Run in the Laurel batidis exhibited histological signs of disease Fork Area of Highland County, Virginia. but no mortality occurred and few clinical Catesbeiana. Bulletin of the Virginia Her- signs of disease were observed (Vasquez et al. petological Society 16:35–43. 2009). Allyn, R. 1952. A Dictionary of Reptiles. First Species-level responses to climate change Edition. The Great Outdoors Association, have been examined in the context of several St. Petersburg, Florida. 85 + [2] pp. characteristics of amphibians, including vari- Allyn, R. 1956. A Dictionary of Reptiles and ation in physiological tolerance, interspecific Amphibians. Great Outdoors Publishing competition, population size, habitat pref- Co., St. Petersburg, Florida. 87 pp. erence, and range size (Bernardo and Spo- Allyn, R. 1961 [1956]. A Dictionary of Rep- tila 2006; Bernardo et al. 2007; Dodd 1997; tiles. Fourth Edition. Great Outdoors Grant et al. 2010; Grover 2000; Hamed 2014; Publishing Co., St. Petersburg, Florida. 86 Hoffacker et al. 2018; Markle and Kozak pp. 2018). Desmognathus monticola is a broadly Allyn, R. 1977 [1956]. A Dictionary of Rep- distributed, eurythermal generalist strongly tiles and Amphibians. Reprinted. Great associated with lotic systems. The species is Outdoors Publishing Co., St. Petersburg, not physiologically constrained to a relative- Florida. 87 pp. Account 925 17

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Bartlett, R. D. 1989. Salamandermania part Beane, J. C., A. L. Braswell, J. C. Mitchell, W. III. The Herptile. Journal of the Interna- M. Palmer, and J. R. Harrison III. 2010. tional Herpetological Society 14:9–13. Amphibians and Reptiles of the Carolinas [Reprinted from Notes from NOAH and Virginia. Second Edition. Revised 14(11):2–6; cited above under Anony- and Updated. The University of North mous, 1987]. Carolina Press, Chapel Hill, North Caro- Bartlett, R. D. and P. P. Bartlett. 1999. A Field lina. 274 pp. Guide to Florida Reptiles and Amphibi- Beckenbach, A. T. 1975. Influence of body ans. Gulf Publishing Company, Houston, size and temperature on the critical oxy- Texas. xvi + 280 pp. gen tension Bartlett, R. D. and P. P. Bartlett. 2006. Guide of some plethodontid salamanders. Phys- and Reference to the Amphibians of East- iological Zoology 48:338–347. ern and Central North America (North Behler, J. L. and F. W. King. 1979. The Audu- of Mexico). University Press of Florida, bon Society Field Guide to North Ameri- Gainesville, Florida. xviii + 284 pp. can Reptiles and Amphibians. First Print- Beachy, C. K. 1994. Community ecology in ing. Alfred A. Knopf, New York, New streams: Effects of two species of predato- York. 719 pp. ry salamanders on a prey species of sala- Behler, J. L. and F. W. King. 1985. The Audu- mander. Herpetologica 50:129–136. bon Society Field Guide to North Amer- Beachy, C. and R. Bruce. 1999. Miniaturiza- ican Reptiles and Amphibians. Fourth tion in Desmognathus quadramaculatus Printing. Alfred A. Knopf, New York, is due to accelerated maturation. P. 64 in New York. 743 pp. [Pagination increased Program Book and Abstracts. Hosted by between 1st and 4th printing because in- the Pennsylvania State University. State dex changed from three columns to single College, Pennsylvania, USA. June 24–30, column]. 1999. Joint meeting of the American So- Behler, J. L. and F. W. King. 1988. The Audu- ciety of Ichthyologists & Herpetologists, bon Society Field Guide to North Ameri- 79th Annual Meeting. American Elas- can Reptiles and Amphibians. Fifth Print- mobranch Society, 15th Annual Meet- ing. Alfred A. Knopf, New York, New ing. Herpetologists’ League, 47th Annual York. 743 pp. [Thirty printings between Meeting. Society for the Study of Amphib- 1988 and 2018]. ians and Reptiles, 42nd Annual Meeting. Beltz, E. 1989. List of scientific names and Beamer, D. A. 2015. Rectifying Limitations original citations for the reptiles and am- on Species Delineation in Dusky Sala- phibians of the continental United States manders: Lineage Detection Using an and Canada part II: Salamanders. Bulle- Ecoregion-Drainage Sampling Regime. tin of the Chicago Herpetological Society Unpublished Ph.D. Dissertation, East 24:93–97. Carolina University, Greenville, North Beltz, E. 1995. Citations for the Original De- Carolina. [2] + [8] + 168 + [1] pp. scriptions of North American Amphibi- Beamer, D. A. and T. Lamb. 2008. Dusky Sal- ans and Reptiles. Society for the Study of amanders (Desmognathus, Plethodon- Amphibians and Reptiles Herpetological tidae) from the Coastal Plain: Multiple Circular 24. iii + 44 pp. independent lineages and their bearing Bernardo, J. 2002. Desmognathus carolinen- on the molecular phylogeny of the genus. sis (Carolina Mountain Dusky Salaman- Molecular Phylogenetics and Evolution der) and Plethodon welleri (Weller’s Sala- 47:143–153. mander); Desmognathus monticola (Seal 20

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