Pristimantis verrucolatus , Paez, Nadia B. & Ron, Santiago R., 2019
Paez, Nadia B. & Ron, Santiago R., 2019, Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species, ZooKeys 868, pp. 1-112: 1
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Pristimantis verrucolatus sp. nov.
English: Warty Flank Rain Frog Spanish: Cutín de flancos verrugosos
(17: 15 males, 1 female, 1 juvenile). Ecuador: Azuay Province: QCAZ 45174, adult female, QCAZ 45175, QCAZ 45195, adult males, from San Antonio, Cajas National Park border (2.8612S, 79.3787W, 2943 m), collected by Elicio E. Tapia and Juan Carlos Morocho in August 2009; QCAZ 46981, QCAZ 46984-989, adult males, collected with the holotype; QCAZ 46993-995, adult males, QCAZ 46992, juvenile, from Cochapamba (2.8020S, 79.4139W, 3662 m), collected by Silvia Aldás and Freddy Velásquez in January 2010; QCAZ 46999-7001, adult males, from Cochapamba (2.7971S, 79.4156W, 3548 m), collected by Elicio E. Tapia in January 2010.
A member of the Pristimantis phoxocephalus group characterized by the following combination of characters: (1) skin on dorsum shagreen with or without scattered tubercles, larger on posterior dorsum; with or without a faint middorsal fold; head with or without a middorsal row of two inconspicuous tubercles; flanks with warts and larger tubercles than those on dorsum; dorsolateral folds absent; thick, continuous or fragmented lateral folds on anterior flanks; skin on venter coarsely areolate; discoidal fold present or absent; (2) tympanic membrane and tympanic annulus prominent, its upper and posterior margin concealed by supratympanic fold; (3) snout moderately long, acuminate with a fleshy keel in dorsal view, protruding in profile; (4) upper eyelid with several low rounded tubercles; cranial crests absent; (5) dentigerous processes of vomers prominent, oblique, moderately separated, posteromedial to choanae; (6) males bearing vocal slits, vocal sac, and white nuptial pads; (7) Finger I shorter than Finger II; discs of digits broadly expanded, elliptical to truncate; (8) fingers with lateral fringes; (9) ulnar tubercles low and round sometimes connected by a weak fold; (10) heel bearing one or more small, prominent, subconical tubercles surrounded or not by some lower tubercles; outer and inner edge of tarsus bearing a row of low, rounded tubercles, inner tarsal fold present; (11) inner metatarsal tubercle ovoid, elevated, four times the size of round outer metatarsal tubercle; supernumerary tubercles numerous; (12) toes with broad lateral fringes; basal webbing present; Toe V longer or much longer than Toe III (disc on Toe III reaches the proximal to distal edge of penultimate subarticular tubercle on Toe IV, disc on Toe V reaches the proximal to distal edge of distal subarticular tubercle on Toe IV); toe discs smaller than those on fingers, elliptical to truncate ( Fig. 9DView Figure 9); (13) in life, dorsum light to dark brown; head with an interorbital stripe or band and a dark supratympanic stripe; venter cream to white with or without brown reticulations; groins and posterior surfaces of the thighs reddish brown with small light brown, orangey brown or yellow spots; iris coppery brown ( Fig. 36View Figure 36); (14) average SVL in adult females: 44.0 ± 4.5 mm (40.4-47.5 mm; n = 2); in adult males: 29.4 ± 2.7 mm (25.1-34.5 mm; n = 15).
Comparison with other species.
Pristimantis verrucolatus is most similar to P. atillo , P. jimenezi , P. phoxocephalus , P. teslai , P. torresi , and P. totoroi . It differs from all of them by having thick lateral folds. Except for P. teslai , it is the only having large tubercles or warts on flanks; though tubercles on flanks are also present in P. teslai , they are not as large as the ones of P. verrucolatus ; furthermore, dorsum of P. teslai is tuberculate, while the dorsum P. verrucolatus is shagreen, with or without scattered tubercles. Pristimantis verrucolatus is further distinguished from P. atillo by the coloration of its groins (orange in P. atillo ; reddish brown with light brown to yellow spots in P. verrucolatus ), and its larger size (males Z = 2.35, p = 0.0188, SVL = 23.4-27.3 mm in P. teslai , 25.1-34.5 mm in P. verrucolatus ). Pristimantis jimenezi , which has an adjacent distribution at lower elevations, is smaller than P. verrucolatus (males Z = 3.58643, p = 0.0003, SVL = 21.8-27.0 mm in P. jimenezi , 25.1-34.5 mm in P. verrucolatus ; females Z = 2.00347, p = 0.0451, SVL = 31.1-37.4 mm in P. jimenezi , 40.4-46.8 mm in P. verrucolatus ). Additionally, the advertisement call of P. verrucolatus is very distinctive, distinguishing it from all species with available calls: P. jimenezi , P. phoxocephalus , and P. totoroi . The call of P. verrucolatus has a single note, longer than those of the calls of the other species. It also has the lowest dominant frequency and the highest variation in frequency from the beginning to the end of the note (Table 6). Pristimantis sp. (CCS1) is the closest species to P. verrucolatus (average p genetic distance = 3.2%), they are not morphologically alike. CCS1 has a chubbier body, lacks a keel at the tip of the snout (present in P. verrucolatus ), has slightly expanded discs on fingers and toes (broadly expanded in P. verrucolatus ), and lacks lateral folds (present in P. verrucolatus ).
Description of the holotype.
QCAZ 46982, SC29952 adult male. Measurements in mm: SVL 28.4; TL 13.3; FL 12.9; HL 8.8; HW 9.8; ED 3.3; TD 1.4; IOD 2.7; EW 2.5; IND 2.2; EN 2.6; TED 1.0. Head wider than body, wider than long; snout moderately long, acuminate with a fleshy keel in dorsal view, protruding in profile; nostrils slightly protuberant, ovoid, directed dorsolaterally; canthus rostralis weakly concave in dorsal view, angular in cross section; loreal region concave; cranial crests absent; upper eyelid bearing several low rounded tubercles; tympanic membrane and annulus prominent, its upper and posterior margins concealed by supratympanic fold; two enlarged, prominent and rounded postrictal tubercles surrounded by smaller tubercles. Choanae large, rounded, not concealed by palatal shelf of maxillae; dentigerous processes of vomers prominent, oblique, moderately separated, posteromedial to choanae; each vomer bearing several teeth; tongue slightly longer than wide, posterior border notched; posterior three fifths free; vocal slits slightly curved, located at posterior half of mouth floor in between tongue and margin of jaw; vocal sac present.
Skin on dorsum shagreen with scattered low tubercles; thin middorsal fold; thick lateral folds; flanks with large warts; skin on venter coarsely areolate. Chest, throat, and ventral surfaces of thighs areolate; discoidal fold absent. Low and round ulnar tubercles connected by an ill-defined fold; white nuptial pads present; palmar tubercles prominent, outer palmar tubercle bifid, slightly bigger than ovoid thenar tubercle; subarticular tubercles prominent, rounded; supernumerary tubercles at base of fingers prominent, smaller than subarticular tubercles; fingers with lateral fringes; Finger I shorter than Finger II; discs on fingers broadly expanded, elliptical; ventral pads on fingers surrounded by circumferential grooves ( Fig. 9DView Figure 9).
Hindlimbs slender; dorsal surfaces of hindlimbs with scattered low tubercles; posterior surfaces of thighs smooth, ventral surfaces of thighs areolate; heel bearing a small subconical tubercle surrounded by some lower rounded tubercles; outer edge of tarsus bearing large low subconical tubercles; inner tarsal fold present followed by small distinct round tubercles on inner edge of tarsus; inner metatarsal tubercle ovoid, elevated, 4 × the size of round outer metatarsal tubercle; distinct, round supernumerary tubercles at the base of toes, indistinct ones on the rest of plantar surface; subarticular tubercles prominent, rounded; toes with broad lateral fringes; basal webbing present; discs on toes expanded, elliptical, smaller than those on fingers; all toes having ventral pads surrounded by circumferential grooves; relative lengths of toes: I < II < III < V < IV; Toe V much longer than Toe III (disc on Toe III reaches distal edge of the penultimate subarticular tubercle on Toe IV, disc on Toe V reaches distal edge of distal subarticular tubercle on Toe IV; Fig. 9DView Figure 9). Coloration in life and preservative is shown in Figures 33BView Figure 33 and 36AView Figure 36.
Coloration of holotype in preservative. Dorsum gray with light gray reticulations bordered by black lines; black interorbital and supratympanic stripes, and black canthal and labial bars; dorsal surfaces of limbs light gray with dark transversal bands bearing scattered black flecks; flanks with oblique light gray reticulations bordered by black spots; groins and posterior surfaces of thighs dark brown with cream spots; venter white; ventral surfaces of limbs dusty cream; throat dusty cream with dark brown mottling near the lips ( Fig. 33BView Figure 33).
Coloration of holotype in life. Based on studio photographs. Dorsum brown with light brown reticulations bordered by black lines; black interorbital and supratympanic stripes, and black canthal and labial bars; dorsal surfaces of limbs light brown bearing dark brown transverse bands with scattered black flecks; flanks with oblique light brown reticulations bordered by black spots; groins and posterior surfaces of thighs dark brown with small cream spots; venter white; throat yellowish cream, vocal sac and ventral surfaces of limbs dusty pinkish cream; iris coppery brown with thin black reticulations; white sclera ( Fig. 36AView Figure 36).
Data are based on 18 preserved specimens and photographs from six living individuals. Variation in life and preservative is shown in Figures 36View Figure 36, 37View Figure 37. Coloration in life is given in parenthesis. Dorsal coloration varies from light gray to dark gray or brown (light to dark brown). Dorsal pattern can be uniformly colored, with irregular reticulations or with a middorsal band. Light colored individuals bear a pale stripe on lips. Head usually bears a dark canthal stripe that continues along the upper eyelid and above the tympanum, and an interorbital stripe. Most specimens have labial bars. Flanks bear or not pale reticulations or black flecks. Groins and posterior surfaces of thighs are brown with cream dots (reddish brown with small light brown or orangey brown spots). Ventral surfaces of limbs are dusty cream to brown; venter is cream or white with or without brown reticulations; throat is cream to dusty brown with or without dark mottling. Iris is coppery brown with thin black reticulations; white sclera. Flanks have medium sized to large tubercles and warts, arranged or not in rows; lateral folds can be thick and continuous or be present as a row of tubercles.
Based on recordings of QCAZ 46981 (September 21, 2010; 19h00; 8.8 °C) and QCAZ 46984 (January 20, 2010; 21h10). Advertisement calls consist of a single sharp whistle ( Fig. 6DView Figure 6); each lasting on average 0.43 s (range 0.41-0.46 s). The peak time of each note occurs exactly in the middle of its duration. The dominant and fundamental frequencies are the same, on average 2144 Hz (range 1927-2302 Hz). The note increases its frequency from beginning to end (2231-1838 Hz on average). Descriptive statistics for bioacoustic parameters are shown in Table 6.
Distribution, natural history, and conservation status.
This species is known from Western Andean slopes of Azuay Province, between 2943-3662 m a.s.l ( Fig. 2View Figure 2). It inhabits Paramo and Western Montane Forest regions. Individuals collected during the day were found beneath rocks or inside arboreal bromeliads up to 2 m above ground; at night they were found active on branches of low shrubs, up to 80 cm above the ground, or inside bromeliads. Gravid females were found in August and January; calling males in January.
Pristimantis verrucolatus is known from only two localities (sensu IUCN) and has a very restricted distribution. These places are adjacent to Parque Nacional Cajas, a protected area with unexplored regions that represent potential distribution for the species. Thus, following the IUCN (2017) guidelines, we assign this species to the Data Deficient Red List category.
The specific epithet is a noun in apposition with masculine gender. It is derived from the Latin words verruca meaning wart, and latus meaning flanks. The name refers to the large and low tubercles or warts on flanks that characterize this species.
Pristimantis cryptomelas species group new taxon
Definition. Monophyly of the P. cryptomelas species group is strongly supported in our phylogeny. Members of this group are characterized by: (i) postocular folds present; (ii) dorsolateral folds absent; (iii) cranial crests absent (except for low crests in P. spinosus ); (iv) tympanic membrane and tympanic annulus prominent; (v) eyes bearing prominent tubercles; (vi) dentigerous processes of vomer present; (vii) prominent tubercles on heel and tarsus; (viii) fingers and toes with lateral fringes; (ix) broadly expanded discs on fingers and toes; (x) basal webbing between toes present; (xi) in life, groins and concealed surfaces of thighs have distinctive coloration patterns, including flash colors, and light or bright colored flecks or spots on a darker background; colors, shapes, and sizes of these ornaments are variable among species; (xii) SVL females 25.9-46.1 mm; SVL males 16.1-30.3 mm.
Content. The P. cryptomelas species group comprises four described species, P. cryptomelas , P. gagliardoi , P. muscosus , and P. spinosus , and the newly described P. nangaritza sp. nov. We suspect that the species content of the group will increase with collections and genetic studies of populations from northern Peru.
Distribution. Members of the P. cryptomelas group occur in the eastern Andean slopes of southern Ecuador and northern Peru. In Ecuador, they inhabit the Eastern Foothill and Montane Forest of Cañar, Loja, Morona Santiago, and Zamora Chinchipe Provinces, between elevations of 1800 and 3500 m a.s.l. In Peru, they live between 1770 and 2820 m a.s.l. Distribution based on QCAZ collection specimens, Duellman and Lehr (2009), and Yánez-Muñoz et al. (2012).
Remarks. The P. muscosus specimen included in our phylogeny is from the same and only population of P. muscosus reported for Ecuador ( Yánez et al. 2012). The population is 216 km from the type locality in Peru (east slope of Abra Pardo de Miguel). Our results indicate that most species of Huicundomantis have small geographic ranges. The large geographic distance from the type locality of P. muscosus suggests that the Ecuadorian population may represent an undescribed species. The same applies to Peruvian populations of P. cryptomelas , which are widely separated from the type locality in Ecuador.
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