Pristimantis nelsongalloi, Valencia & Valladares-Suntasig & Tipantiza-Tuguminago & Dueñas, 2019

Valencia, Jorge H., Valladares-Suntasig, Francisco, Tipantiza-Tuguminago, Luis & Dueñas, Manuel R., 2019, A new species of terrestrial-breeding frog of the genus Pristimantis (Anura Terrarana: Craugastoridae) from the eastern Andean slopes of the southern Ecuador, Zootaxa 4658 (3), pp. 509-525 : 511-520

Pristimantis nelsongalloi sp. nov. ( Figs. 1–2 )

Suggested English name: Nelson Gallo’s Rainfrog

Suggested Spanish name: Cutín de Nelson Gallo

Holotype. FHGO 13018 ( Figs. 1–2 ) an adult male from 9 de Octubre (2°14’52” S, 78°16’37” W, 1778 m elevation), Morona , province of Morona Santiago, Ecuador, collected on 29 Octubre 2017 by Jorge H. Valencia and Julio Venegas.

Paratypes (n = 13). Two adult female ( FHGO 12997 , 13240 ), seven adult males ( FHGO 13016–17 , 13241–43 ), and a juvenile male ( FHGO 13009 ) from the same locality collected between October 6 and 9, 2017 by Jorge H. Valencia and Julio Venegas, and between December 17 and 23, 2017 by Luis Tipantiza-Tuguminago and Javier Naja- ndey. Two adult males ( DHMECN 5223 , 5226 ), an adult male ( DHMECN 5224 ); and a juvenile female ( DHMECN 5225 ) from Reserva Río Zuñac (1°24’03” S, 78°11’12” W, 1697 m elevation), Baños , province of Tungurahua, Ecuador, collected on 15 May 2008 by Mario Yánez-Muñoz , Miguel Urgilés, and Andrés Laguna .

Diagnosis. Pristimantis nelsongalloi sp. nov. is characterized by: (1) skin texture of dorsum finely shagreen, with two subconical scapular tubercles and several scattered pustular tubercles, dorsolateral folds thin anteriorly and discontinuous posteriorly; venter areolate; discoidal fold weak; thoracic fold present; (2) tympanic membrane and tympanic annulus round, well differentiated; horizontal diameter of tympanic annulus 70–93% (mean = 78.5%) the diameter of the eye and separated from the eye by a distance of approximately one sixth of the length of the eye; (3) snout short, subacuminate in dorsal view, rounded in profile, canthus rostralis distinct, rounded in dorsal view and slightly sharp in lateral profile, loreal region slightly concave to flat; nostrils slightly protuberant, directed dorsolaterally; area between nostril slightly convex; (4) upper eyelid bearing four or five small, flat and low supraocular tubercles, approximately 90% of eye diameter; cranial crests absent; (5) choanae large and rounded, concealed by palatal shelf of maxillary arch; dentigerous processes of vomers low, and oblique, slightly smaller than choanae, arranged in V-shaped, posterior and medial to choanae, each bearing three to five teeth; (6) males lacking vocal sacs and vocal slits, nuptial pads absent; (7) Finger I shorter than Finger II, discs on Finger I and II not expanded or slightly expanded, discs on Finger III and IV expanded, all fingers bearing lanceolate discs ( Fig. 3A ); (8) fingers lacking lateral fringes; (9) small ulnar tubercles present; (10) inner edge of tarsus bearing subconical tubercles, heel bearing one subconical tubercle and several smaller tubercles; tarsal fold absent; (11) inner metatarsal tubercle oval and slightly elevated, about 1.7–2.1x the size of the small, round subconical outer tubercle; supernumerary plantar tubercles present; (12) toes without lateral fringes; webbing absent; Toe V equal or slightly longer than Toe III; discs on Toes III to IV slightly longer than those outer fingers, all toes bearing lanceolate discs ( Fig. 3B ); (13) dorsum yellowish tan to pale brown with darker markings, throat and venter bearing minute brown spots, iris yellowish bronze with median, horizontal reddish streak, all reticulated with black; (14) SVL in adult males 12.0–17.0 mm (mean = 14.8, SD = ±1.6, n = 8), females 18.5–21.7 mm (n = 2) ( Table 1 ).

Similar species. Pristimantis nelsongalloi is distinguished from most species in the genus Pristimantis by the combination of the following characters: Toe V equal or slightly longer than Toe III, Toe V reaches about to the level of the penultimate subarticular tubercle on Toe IV, fingers and toes slightly expanded and bearing lanceolate discs, vocal slits and sac absent, and large tympanum (70–93% the diameter of the eye). Pristimantis nelsongalloi is most similar to eastern Andean frogs of the Pristimantis myersi species-group ( Table 2 , Fig. 4 ): Pristimantis bicantus Guayasamin & Funk, 2009 , P. festae ( Peracca, 1904) , P. gladiator ( Lynch, 1976) , P. leoni ( Lynch, 1976) , P. myersi ( Goin & Cochran, 1963) , and P. ocreatus ( Lynch, 1981) (characters of the new species in parentheses). Pristimantis bicantus has a tympanum diameter 45–50% of eye diameter, ulnar tubercles absent, heel, inner and outer edges of tarsus lacking tubercles (tympanum diameter 70–93% of eye diameter, small ulnar tubercles present, heel bearing one large tubercle and several smaller tubercles, inner edge of tarsus bearing indistinct tubercles); Pristimantis festae has a small tympanum, about half eye length, venter black with yellow or reddish blotches, heel and tarsus lacking tubercles, toes bearing narrow lateral fringes (tympanum 70–93% of eye diameter, venter cream or yellowish, heel bearing one large tubercle and several smaller tubercles, lateral fringes absent on toes); Pristimantis gladiator has a dorsum with short ridges, snout slightly pointed, male with vocal slits and subgular vocal sac, venter gray and yellow-gray (dorsolateral folds thin anteriorly and discontinuous posteriorly, snout lacking papilla at tip, males lacking vocal silts and sac, venter cream or yellowish); Pristimantis leoni has a dorsum with many short ridges and scattered large tubercles, tympanum diameter about 30–40% of eye diameter, males with vocal slits and subgular vocal sac, venter gray with black flecks (dorsolateral folds thin anteriorly and discontinuous posteriorly, tympanum diameter 70–93% of eye diameter, males lacking vocal silts and sac, venter cream or yellowish); Pristimantis myersi has paravertebral folds and sinuate dorsolateral folds, venter strongly areolate, snout bearing papilla at tip, dentigerous process of vomer absent, males with vocal slits and nuptial pads, toes bearing lateral fringes (dorsolateral folds thin anteriorly and discontinuous posteriorly, venter areolate, snout lacking papilla at tip, dentigerous process of vomers present, vocal slits and nuptial pads absent in males, lateral fringes on toes absent); Pristimantis ocreatus has short vocal slits, internal vocal sac, vomers lacking dentigerous process, fingers lacking pads, tympanum concealed (vocal sacs and slits absent, dentigerous process of vomer present, fingers bearing pads, tympanum present, diameter 70–93% of eye diameter).

Pristimantis albujai Brito, Batallas & Yánez-Muñoz, 2017 , and Pristimantis sambalan Brito, Batallas & Yánez- Muñoz, 2017 two species recently described from the Upper Upano Valley ( Brito et al. 2017) are also similar to Pristimantis nelsongalloi but differs by the following characters ( Table 2 , Fig. 4 ): Pristimantis albujai has a dorsum with dorsolateral folds, upper eyelid with a prominent conical tubercle, Toe V much longer than Toe III, Toe V reaching the proximal edge of distal subarticular tubercle on Toe IV, venter dark brown, groin reddish (dorsolateral folds thin anteriorly and discontinuous posteriorly, upper eyelid bearing four or five small tubercles, Toe IV slightly longer than Toe III, Toe V does not extend beyond the distal edge of distal subarticular tubercle on Toe IV, venter cream, yellowish or pale brown, groin orange); Pristimantis sambalan has a dorsum with a row of dorsolateral subconical tubercles, eyelid bearing one or two conical tubercles, Toe V longer than Toe III, Toe V not extended beyond the proximal edge of distal subarticular tubercle on Toe IV, venter reddish with brown spots or blotches (dorsolateral folds thin anteriorly and discontinuous posteriorly, upper eyelid bearing four or five small tubercles, Toe IV slightly longer than Toe III, Toe V no extended beyond the distal edge of distal subarticular tubercle on Toe IV, venter cream, yellowish or pale brown).

In addition, Pristimantis nelsongalloi eventually could be confused with Niceforonia elassodisca ( Lynch, 1973) due to similarities in body shape and dorsal coloration. Niceforonia elassodisca has venter smooth, tympanum diameter about 40–50% of eye diameter, Finger I longer than Finger II, rounded discs on fingers and toes, terminal discs not expanded, Toe III longer than Toe IV, and terminal phalanges narrow T-shaped (venter areolate, tympanum diameter 70–93% of eye diameter, Finger I slightly smaller than Finger II, lanceolate discs on finger and toes, terminal discs on digits expanded bearing well-defined circumferential grooves, Toe V equal or slightly longer than Toe III, terminal phalanges wide T-shaped) ( Fig. 5 ). Both species are distributed in the eastern Andean slopes of Ecuador. However, N. elassodisca is known from elevations of 2300–2900 m in upper humid montane forest ( Lynch 1981), and P. nelsongalloi occurs in elevations of 1627–1800 m in montane cloud forest.

Description of holotype. Adult male (16.9 mm SVL; Figs. 2 A–D); head slightly wider than body; head barely wider than long; head length 37.8% of SVL. Canthus rostralis sharp, loreal region slightly concave to flat, sloping abruptly; nostrils slightly protuberant, directed laterally; eye–to–nostril distance 86% of eye diameter; lips rounded; upper eyelid width 70.8% of interorbital distance, bearing several small and flat tubercles; snout subacuminate in dorsal view and rounded in lateral profile; tympanic membrane and tympanic annulus evident, round, its upper and posterodorsal part obscured by rounded supratympanic fold; supratympanic fold thick and slightly warty; several low postrictal tubercles situated posterodorsally to tympanic annulus; choanae small, rounded concealed by palatal shelf of maxillary arch; dentigerous processes of vomers small and oblique each bearing three teeth; tongue longer than wide, bilobate, posterior ½ not adherent to floor of mouth; vocal slits and sac absent.

Skin of dorsum finely shagreen and bearing scattered small tubercles; dorsolateral folds short and low on middle of dorsum, and granular posteriorly; ventral and lateral skin coarsely areolate, that on throat shagreen; interocular folds absent; discoidal fold present; thoracic fold present (more conspicuous in life); small ulnar tubercles present without forming a distinct fold; thenar tubercle smaller than divided palmar tubercle; supernumerary palmar tubercles barely evident; subarticular tubercles slightly prominent and rounded, including the most distal tubercle; fingers short and moderately slender, lacking narrow lateral fringes; disc on Finger I not expanded, on Finger II slightly expanded, those of Fingers III and IV expanded; discs on Fingers I and IV lanceolate; disc pads on Fingers I–II triangular, those of Fingers III and IV lanceolate ( Fig. 3A ); tip of Finger IV reaches beyond the distal subarticular tubercle on Finger III; tip of Finger II does not reach the distal subarticular tubercle of Finger III; knee bearing low tubercles.

Hind limbs moderately short; tibia length 46.5% of SVL; foot length 39.4% of SVL; Toe V as long as Toe III; tip of Toe III reaching the distal edge of the penultimate subarticular tubercle on Toe IV; Toe V does not extend beyond the distal edge of distal subarticular tubercle on Toe IV ( Fig. 3B ); discs on inner toes slightly smaller than those on outer fingers; tips of digits lanceolate; webbing absent; subarticular tubercles on toes distinct, barely projecting and ovoid; supernumerary plantar tubercles at base of toes present; inner metatarsal tubercle ovoid and slightly elevated, about 2/3 the length of Toe I; lateral fringes absent; outer metatarsal tubercle small, subconical; tarsus without fold but small subconical tubercles along outer border, indistinct on inner border; heel bearing one small, subconical tubercle and two or three flat, smaller tubercles; cloacal sheath and tubercles present, cloacal opening direct posteroventrally at level of thighs.

Coloration of the holotype in life. Dorsum brown-orange with a distinct dark brown interorbital bar, pale brown-grayish chevron markings, and two distinct dark brown scapular spots ( Fig. 1A ); dark brown cloacal region; distinct dark brown canthal and postocular stripes are present; canthal region pale yellow, and upper lips with brown labial bars and brown-orange spots; scapular region with two black spots. Dorsal surface of limbs brown–orange with pale brown-grayish transversal bars. Skin dorsum on hands and feet with brown blotches, Finger I and II pale yellowish. Flanks brown-orange with brown markings extending towards groin. Groin, hidden surfaces of thighs, and ventral surface of arms bright orange. Venter and ventral surface of thighs translucent with white spots; throat with indistinct brown spots ( Fig. 1B ); iris yellowish bronze with median, horizontal reddish streak, all reticulated with black.

Coloration of the holotype in preservative. Dorsum pale brown with a dark brown interorbital bar, dark brown chevron markings, and two distinct dark brown scapular spots; dark brown cloacal region with yellowish spots; dark brown canthal and postocular stripes; canthal region pale yellow, and upper lips with dark brown labials bars. Dorsal surface of limbs yellow with dark brown transversal bars. Flanks yellowish finely speckled with brown. Groin and hidden surfaces of thighs pale reddish. Venter and throat pale yellowish with white spots ( Fig. 2 A–B).

Measurements of holotype (in mm). SVL 17.0; HL 6.4; HW 6.6; ED 2.1; END 1.8; EW 1.9; IOD 2.1; IND 1.7; ETD 0.4; TYD 1.6; TL 7.4; FL 6.8; FoL 6.7; HaL 3.5.

Variation. Morphometric variation in adult males and females is described in Table 1 . Color variation in life is shown in Figure 6 . Dorsum dark brown or dark gray in females FHGO 12997, 13240; dorsum with a longitudinal yellow stripe in juvenile male FHGO 13009; dorsum bearing several dark brown blotches in male DHMECN 5223; chevron marks and bars on dorsal surface of thighs darker in males DHMECN 5225, FHGO 13243; orange-brown blotches on dorsum, arms, and thighs in male FHGO 13017; two white spots on dorsum in male FHGO 13017; head darker in female FHGO 12997; canthal and postocular pale brown stripes barely evident in female FHGO 13240; interorbital bar barely evident in female DHMECN 5224, and juvenile male DHMECN 5225; two postocular dark brown subconical tubercles in female DHMECN 5224; labial bars slightly differentiated in females DHMECN 5224, FHGO 12997, 13240; two males FHGO 13009, 13243 have a slightly smaller tympanum diameter, between 61–65% of their eye diameter; venter strongly mottling with brown spots in females DHMECN 5224, FHGO 12997, and males DHMECN 5226, FHGO 13243; venter cream with orange and white blotches in juvenile males DHMECN 5225, FHGO 13009; female DHMECN 5224 with orange groin; throat with white blotches in female FHGO 12997; throat strongly dark brown mottling in female DHMECN 5524, and male DHMECN 5226; groin and anterior surface of thighs yellowish; preanal cream stripe in male FHGO 13243, two preanal white spots in female FHGO 12997; disc on Finger III and IV rounded in males DHMECN 5223, FHGO 13017, 13241; feet lacking brown bars in female FHGO 12997.

Advertisement call. Pristimantis nelsongalloi has a distinctly pulsed note produced in very short series of modulate frequency. Twelve calls of the holotype ( FHGO 13018) and nine calls of the paratype ( FHGO 13016) were recorded at an air temperature of 13.5–15.1° C. Duration of notes of the holotype ranged 1.001 –1.250 seconds (mean = 1.120, SD = 0.06) and paratype ranged 1.030 –1.233 seconds (mean = 1.125, SD = 0.05) and consist of three short pulses. The dominant frequency of the holotype was 11771–12722 Hz (mean = 12389.9, SD = 364.1), and of the paratype was 11346–12237 Hz (mean = 11773.7, SD = 318.6). Waveforms and spectrograms of calls are presented in Figure. 7 A–B.

Etymology. The specific epithet nelsongalloi is a patronym honoring Dr. Nelson Gallo. Dr. Gallo, Ecuadorian biologist who was the Director of the Laboratory of Zoology and Professor of Zoology at the Biology College of the Universidad Central del Ecuador. He formed several generations of biologist (including the first author of this work). We name this new species in his honor for his indefatigable teaching work, support, friendship, and guidance.

Distribution and natural history. Pristimantis nelsongalloi is known from two localities ( Fig. 8 ). The locality of 9 de Octubre, buffer zone of the Parque Nacional Sangay, in the province of Morona Santiago, and Reserva Biológica Río Zuñac, from the province of Tungurahua, Ecuador. Both localities are in evergreen montane forest (based on Ministerio del Ambiente de Ecuador 2013, ecosystem classification system), located in the eastern Andean slopes of Ecuador (elevations ranges is 1627–1800 m) ( Fig. 6 ). Pristimantis nelsongalloi is a nocturnal and secretive species associated to riverine vegetation. Specimens were found in Andean cloud forest during a cloudy and drizzly night. Males were observed calling partially hidden in the leaf litter. Six to nine males were hearing in transects of 100 m. Some specimens were found under rocks or trunks by day. A gravid female contained 13 white eggs of 2–3 mm. Sympatric congeners include Pristimantis bicantus , P. churuwiai , P. peruvianus , P. serendipitus , P. trachyblepharis , P. ventrimarmoratus and other three undescribed species.

Conservation status. We consider Pristimantis nelsongalloi to be Data Deficient following IUCN (2001) guidelines criteria because it is known only from two localities, separated from each other by a distance of ca. 96 km., suggest that it might as well be preset in other localities along of the eastern Andean slopes. The habitat of P. nelsongalloi is highly fragmented surrounded by grassland suggesting tolerance to disturbance in the type locality ( Fig. 9 A–B). Population size is unknown, but the scant evidence suggests moderate abundance in 9 de Octubre and Río Zuñac (M. Yánez-Muñoz com. pers.). Therefore, we consider there is inadequate information to make a direct, or indirect, assessment of its extinction risk based on its scarce distribution data.

Figures

FIGURE 1. Holotype of Pristimantis nelsongalloi sp. nov. in life, FHGO 13018, adult male, SVL 17.0 mm: A) dorsolateral view, and B) ventral view.

FIGURE 2. Holotype of Pristimantis nelsongalloi sp. nov. (FHGO 13018, adult male, SVL 17.0 mm) in preservative:A) dorsal view, B) ventral view, C) head, lateral view, and D) head, dorsal view.

FIGURE 3. Hand and foot of the holotype of Pristimantis nelsongalloi sp. nov. in preservative FHGO 13018, adult female:A) palmar view of hand, B) plantar view of foot.

FIGURE 4. Coloration in life of new species and similar Pristimantis species from eastern Andes slopes of Ecuador. A) Pristimantis nelsongalloi sp. nov., FHGO 13018, from 9 de Octubre, province of Morona Santiago, B) P. albujai, DHMECN 12245, from Sardinayacu, province of Morona Santiago, C) P. bicantus, QCAZ 57262, from Napo Wildlife Center, province of Napo, D) P. festae, QCAZ 39664, from San Marcos lagoon, province of Pichincha, E) P. gladiator, QCAZ 46264, from Las Carmelas, province of Napo, F) P. leoni, DHMECN 6434, from La Bonita, province of Sucumbios, G) P. myersi, QCAZ 46605, from Santa Barbara, province of Sucumbios, H) P. ocreatus, QCAZ 70061, from Parque Nacional Llanganates, province of Morona Santiago, I) P. sambalan, DHMECN 12251, from Cerro Sambalan, province of Morona Santiago. Pictures are not scaled.

FIGURE 5. Drawings showing disc pad (gray) and disc cover on fingers (left), and terminal phalange (right): (A) Pristimantis nelsongalloi sp. nov., adult female, paratype DHMECN 5224: lanceolate disc cover and disc pad, and wide T-shaped terminal phalange; B) P. bicantus, adult male, FHGO 13324: round disc cover and disc pad, and wide T-shaped terminal phalange; and C) Niceforonia elassodisca, adult female, FHGO 13291: round pad, disc cover absent, and narrow T-shaped terminal phalange in. Illustrations are not scaled.

FIGURE 6. Live specimens of Pristimantis nelsongalloi sp. nov. showing variation in dorsal and ventral coloration: A) adult female FHGO 13240, SVL = 21.7 mm, B) juvenile male, FHGO 13009, SVL = 12.0 mm), C) adult male, FHGO 13016, SVL = 16.0 mm, D) adult male, FHGO 13017, SVL = 14.7 mm. All specimens are shown in the same scale.

FIGURE 7. Vocalizations of Pristimantis nelsongalloi sp. nov. adult male, FHGO 13018: A) Waveform (upper) and Spectrogram (lower) of a single, pulsed call; B) Waveform (upper) and Spectrogram (lower) of a call series with three calls.

FIGURE 8. Type locality of Pristimantis nelsongalloi sp. nov., locality of 9 de Octubre province of Morona Santiago, Ecuador (star). Circle represents the locality of Río Zuñac, province of Tungurahua, other locality where the new species was found.

FIGURE 9. Panoramic view of the fragmented habitat (A) and microhabitat (B) of Pristimantis nelsongalloi sp. nov. within the type locality.

Tables

TABLE 2. Selected diagnostic characters and elevational range of Pristimantis species similar to P. nelsongalloi from eastern Andean slopes of Ecuador. Information was taken from references and specimens examined.

Characters	P. nelsongalloi	P. albujai	P. bicantus	P. festae	P. gladiator	P. leoni	P. myersi	P. ocreatus	P. sambalan
Reference	This work	Brito et al. 2017	Guayasamin & Funk 2009	Lynch & Duellman 1980	Lynch & Duellman 1997	Lynch & Duellman 1997	Lynch 1981	Lynch 1981	Brito et al. 2017
SVL females (mm)	12.0–17.0	20.6–21.9	17.0–21.7	17.3–25.5	15.8	19.7–25.0	17.5–23.2	19.6–20.2	20.6–21.9
SVL males (mm)	18.5–21.7	15.6	12.0–15.8	14.4–21.3	14.9–15.6	13.0–18.3	13.7–17.5	12.4–17.2	15.6
Skin on dorsum	Finely shagreen	Finely shagreen with low and flat warts	Shagreen with small scattered, rounded warts	Smooth with small pointed warts and short ridges	Smooth with short ridges	Shagreen with many short ridges, scattered large tubercles	Shagreen and pustulate	Finely areolate bearing subconical warts on flanks	Shagreen with several subconical tubercles
Dorsolateral folds	Thin anteriorly and discontinuous posteriorly	Present	Usually absent	Absent	Absent	Absent	Present	Present	Thin and bearing a row of subconical tubercles
Scapular tubercles	Present	Present	Present	Absent	Absent	Absent	Absent	Present	Absent
Sacral tubercles	Absent	Absent	Absent	Absent	Absent	Absent	Absent	Absent	Absent
Shape of snout	Subacuminate in dorsal view, rounded in profile	Subacuminate in dorsal view, rounded in lateral profile	Round in dorsal and lateral views	Round in dorsal and lateral views	Acuminate in dorsal view, round in lateral profile	Subacuminate in dorsal view, round in lateral profile	Subacuminate in dorsal view, round in lateral profile	Round in dorsal and lateral views	Subacuminate in dorsal view, round in lateral profile One to two
Upper eyelid tubercles	Four or five small and flat tubercles	A conical tubercle	Several low tubercles	Small and flat tubercles	Absent	Two to four conical tubercles	Numerous low tubercles	Three and five conical tubercles	conical tubercles and several round small tubercles

TABLE 2. (Continued)

Characters	P. nelsongalloi	P. albujai	P. bicantus	P. festae	P. gladiator	P. leoni	P. myersi	P. ocreatus	P. sambalan
Skin on venter	Areolate	Slightly areolate with warts	Areolate	Coarsely areolate	Areolate	Areolate	Areolate	Areolate	Areolate
Males characters present	Vocal sacs, vocal slits and nuptial pads absent	Unknown	Vocal slits present, nuptial pads absent	Vocal sacs and vocal slits present, nuptial pads absent	Vocal sacs and vocal slits present, nuptial pads absent	Vocal sacs and vocal slits present, nuptial pads absent	Vocal sacs, vocal slits, and nuptial pads present	Short vocal slits, and internal vocal sac, nuptial pads absent	Vocal sacs and vocal slits present, nuptial pads absent
Tympanic membrane and annulus condition and length	Present, 70–93% of eye diameter	Present, 40–45% of eye diameter	Present, 40– 50% of eye diameter	Present, 33– 50% of eye diameter	Present, 40– 50% of eye diameter	Partially concealed by skin, 33–40% of eye diameter	Present, 40–67% of eye diameter	Concealed	Present, 40–45% of eye diameter
Discs on finger and toes	Lanceolate	Truncate	Round	Round	Round	Round	Round	Round, pads absent	Round
Tarsal tubercles	A row of subconical tubercles	A conical tubercle and tree rounded tubercles	Absent	Absent	Absent	A row of small tubercles	A row of non-conical tubercles	A row of conical tubercles	A row of conical tubercles
Heel tubercles	One subconical tubercle	A prominent conical tubercle	Absent	Absent	Small non- conical tubercles	Absent	Several low tubercles	Absent	Several small conical tubercles
Elevational ranges (m)	1627–1800	1400	1750–2400	2630–3650	2350–2910	1960–3400	2800–3470	3500–4150	2851