Two new species of frogs of the genus Pristimantis from Llanganates National Park in Ecuador with comments on the regional diversity of Ecuadorian Pristimantis (Anura, Craugastoridae) Author Navarrete, Maria J. Author Venegas, Pablo J. Author Ron, Santiago R. text ZooKeys 2016 593 139 162 http://dx.doi.org/10.3897/zookeys.593.8063 journal article http://dx.doi.org/10.3897/zookeys.593.8063 1313-2970-593-139 C9B3FBF2B1E34C44B87ECBF83349BDCA C9B3FBF2B1E34C44B87ECBF83349BDCA Taxon classification Animalia Anura Craugastoridae Pristimantis llanganati sp. n. Common name. English: Llanganates Rain Frog. Spanish: Cutin de los Llanganates. Holotype QCAZ 46140 (field no. SC-PUCE 29720; Figs 2 C-D , 6 A-B ), adult male from Ecuador, Provincia Napo, Canton Tena, "La Cueva" at the confluence of the Mulatos and Langoa rivers ( 0.9663° S , 78.2224° W ), 2483 m above the sea, collected by Elicio E. Tapia and Fernando Nunez on 15 November 2009. Figure 6. Preserved holotype of Pristimantis llanganati sp. n., QCAZ 46140, adult male, SVL = 27.1 mm. Dorsal (A), ventral (B). Paratopotypes (3 specimens). QCAZ 46221 adult female, 46141 and 46142 juveniles, collected with the holotype. Paratypes (2 specimens). Ecuador, Provincia Napo, QCAZ 46227 adult male, Salcedo-Tena road ( 0.9847°S , 78.1928°W ), 2253 m, collected by Elicio E. Tapia and Fernando Nunez on 16 November 2009; QCAZ 46217, juvenile, Salcedo-Tena road ( 0.9670°S , 78.2484°W ), 2883 m, collected by Elicio E. Tapia and Fernando Nunez on 14 November 2009 Diagnosis . The new species is assigned to the genus Pristimantis . Although morphological synapomorphies are unknown for Pristimantis , the new species has the characteristic morphology of most Pristimantis including T-shaped terminal phalanges, toes without membranes, and Toe V longer than Toe III. Pristimantis llanganati is characterized by the following combination of characters: (1) Skin on dorsum covered by minute conical tubercles, skin on venter areolate with scattered warts; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus present, covered by supratympanic fold on its upper and posterior margins; (3) snout short, rounded in dorsal and lateral view; (4) upper eyelid with a low conical tubercle and some low indistinct tubercles posteriorly; EW 82% of IOD; cranial crests absent; (5) dentigerous processes of vomers varying from low and indistinct to high and evident, oblique, moderately separated, posteromedial to choanae; (6) vocals slits present, nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits broadly expanded, truncate; (8) fingers bearing narrow lateral fringes; (9) ulnar and tarsal tubercles present, conical and low; (10) heel with two or three low conical tubercles; inner tarsal fold present, long and ill defined; (11) inner metatarsal tubercle elliptical, low, 2X as large as outer metatarsal tubercle; outer metatarsal tubercle small, ovoid; supernumerary plantar tubercles low and indistinct; (12) toes bearing narrow lateral fringes; toe webbing absent; Toe V longer than Toe III (disc on Toe III reaches the proximal edge to the proximal subarticular tubercle on Toe IV, disc on Toe V extends to the proximal edge of distal subarticular tubercle on Toe IV); toe discs about as large as those on fingers; (13) in life, dorsum olive green with X-shaped or rhomboidal dark brown mark on scapular region, and scattered brown flecks or blotches; flanks, dorsal and posterior surfaces of thighs and ventral surfaces of shanks dirty white or white with dark brown diagonal stripes; groins white or tan with distinct black or dark brown diagonal stripes; venter dirty cream with brown mottling in the throat and chest and brown flecks in belly and ventral surfaces of thighs. Iris coppery with a reddish horizontal stripe; (14) SVL in one adult female 29.8 mm, in adult males 24.0-27.0 mm (n = 2). Comparisons with other species. (Fig. 1). In this section, coloration refers to live individuals unless otherwise noted. Pristimantis llanganati can be easily distinguished from other congeners from the Andes of Ecuador and Colombia, except Pristimantis chloronotus , Pristimantis colonensis , and Pristimantis eriphus , by the presence of the following traits: dorsum green, greenish brown or mossy; spiny appearance bearing distinct conical tubercles on eyelids, heels and outer edge of tarsus; groins white or tan with distinct black or dark brown diagonal stripes; posterior surfaces of thighs and concealed surfaces of shanks with oblique white and brown or black bars. Pristimantis llanganati is most similar to Pristimantis eriphus . Both species have greenish or mossy coloration and tuberculate skin. However, Pristimantis eriphus can be readily distinguished from Pristimantis llanganati in having a brown vertebral band or dark brown chevrons on the scapular regions (brown or reddish brown X-shaped or rhomboidal mark in Pristimantis llanganati ) and greenish coopery or red iris without reticulations (copper iris with dark brown reticulations in Pristimantis llanganati ). Pristimantis chloronotus and Pristimantis colonensis can be easily distinguished from Pristimantis llanganati in having sinuous paravertebral folds (absent in Pristimantis llanganati ; Fig. 1). In addition, Pristimantis chloronotus upper eyelids are covered with small conical tubercles (one distinct conical tubercle surrounded by several lower conical tubercles in Pristimantis llanganati ). Pristimantis colonensis also differs from Pristimantis llanganati in having the posterior surfaces of thighs dark brown with white or cream oblique lines (white with brown bars in Pristimantis llanganati ), and iris yellowish gold (coppery in Pristimantis llanganati ). Pristimantis incanus is also similar to Pristimantis llanganati in coloration and disposition of tubercles on eyelids, heels, and tarsus. However, males of Pristimantis incanus haven vocal slits (absent in Pristimantis llanganati ). Moreover, Pristimantis incanus have glossy white points on groins, posterior surfaces of thighs and concealed surfaces of shanks (those areas are white and black or have dark brown stripes in Pristimantis llanganati ). Description of the holotype. Adult male. Measurements (in mm): SVL 27.0; tibia length 18.8; foot length 22.8; head length 7.9; head width 9.8; eye diameter 3.2; tympanum diameter 1.3; interorbital distance 3.4; upper eyelid width 2.7; internarial distance 2.3; eye-nostril distance 2.8; tympanum-eye distance 1.3. Slender body; head as wide as body, wider than long; head width 36% of SVL; head length 29% of SVL; snout short, rounded in dorsal and lateral view; eye-nostril distance 89% of eye diameter; nostrils narrow, higher than long, directed dorsolaterally; canthus rostralis curved in dorsal view, slightly curved in profile; loreal region concave; lips rounded; upper eyelid bearing one small conical tubercle on its center and some low rounded tubercles posteriorly; upper eyelid width 82% of IOD; tympanic annulus barely visible, with upper and posterior margins covered by supratympanic fold; tympanic membrane present, distinct; tympanum diameter 39% of eye diameter, tympanum-eye distance 102% of tympanum diameter; few indistinct postrictal tubercles present. Choanae small, semicircular, not concealed by palatal shelf of maxilla; dentigerous processes of vomers low, indistinct, oblique, moderately separated, posteromedial to choanae; right vomer bearing two teeth and left vomer one tooth; vocal slits present; tongue slightly wider than long, notched behind, free posteriorly along one third of its length. Skin on dorsum covered by minute conical tubercles, head bearing two large tubercles, one between the orbits and the other half way between the interorbital line and the tip of the snout; dorsolateral folds absent; skin on flanks with minute conical tubercles; skin on throat and chest weakly areolate, belly areolate with scattered enlarged warts, and ventral surfaces of thighs weakly areolate; discoidal fold absent; cloacal sheath short; skin in cloacal region tuberculate with two enlarged tubercles, on each side, below the cloacal sheath. Ulnar tubercles present, subconical and low; elbow bearing one low subconical tubercle; nuptial pads absent; palmar tubercles low, weakly defined, outer palmar tubercle bifid, approximately twice the size of ovoid thenar tubercle; subarticular tubercles low, well defined, round in ventral and lateral view; supernumerary tubercles at base of fingers present, indistinct; fingers bearing narrow lateral fringes; Finger I shorter than Finger II; disc on Finger I rounded and on Finger II expanded, disc on Finger III and IV broadly expanded and truncate; pads on fingers well defined by circumferential grooves on all fingers (Fig. 2). Hindlimbs slender, tibia length 70% of SVL; foot length 84% of SVL; upper surfaces of hindlimbs with minute conical and subconical tubercles; posterior surfaces of thighs smooth, ventral surfaces weakly areolate; knee bearing low conical tubercles; heel bearing low conical tubercles; inner tarsal fold present, long and weakly defined; inner metatarsal tubercle low, elliptical, rounded, two times the size of smaller, oval, rounded outer metatarsal tubercle; plantar surface weakly tuberculate; subarticular tubercles well defined, round in ventral and lateral view; toes bearing narrow lateral fringes; basal webbing between toes absent; discs nearly as large as those on fingers, most prominent on Toe IV and V; discs on toes expanded, rounded; all toes having ventral pads well defined by circumferential grooves; relative lengths of toes: 1 <2 <3 <5 <4 (Fig. 2); Toe V longer than Toe III (disc on Toe III reaches the proximal edge of the proximal subarticular tubercle on Toe IV, disc on Toe V reaches the proximal edge of distal subarticular tubercle on Toe IV). Color of holotype in life is unknown. Color of holotype in ethanol 70% (Fig. 6): dorsal background color is greenish gray with a U-shaped brown mark on the top of head, a broad reddish-brown interorbital bar, an X-shaped reddish brown mark on the mid-dorsum, and scattered brown blotches; sides of head with a broad dark brown canthal stripe, two brown vertical labial bars, brown supratympanic fold; flanks white with dark brown diagonal stripes; groins white with dark brown diagonal stripes; forelimbs with brown transversal bars, hindlimbs with dark brown (nearly black) diagonal transversal bars; posterior surfaces of thighs and concealed surfaces of shanks whitish cream with broad dark brown bars. Venter creamy brown with brown mottling on throat, chest, and forelimbs, dark brown flecks on belly and ventral surfaces of thighs; palms and soles brownish cream. Variation. In this section, coloration refers to preserved individuals. In the type series, adult males (24.0-27.0 mm) (n = 2) are smaller than the single known female (SVL = 29.8 mm) (n = 1). See Table 2 for measurements and proportions of the type specimens and see Figure 7 for photographs of preserved individuals. The single adult male paratype (QCAZ 46227) differs from the holotype in having less conspicuous dorsal marks. The only known female (QCAZ 46221) has the dorsum grayish cream. Three juvenile specimens (QCAZ 46217, SVL 19.3 mm; QCAZ 46141, SVL 12.5 mm and QCAZ 46142, SVL 10.7 mm) are identical in coloration to the holotype but lack well defined dorsal marks. The juvenile QCAZ 46142 has a dark brown venter. Figure 7. Preserved individuals of Pristimantis llanganati sp. n. showing dorsal and ventral variation. A dorsal view B ventral view. From left to right: QCAZ 46221, adult female, SVL = 29.8 mm; QCAZ 46227, adult male, SVL = 24.0 mm. Specimens are shown at the same scale. Table 2. Measurements (in mm) and proportions of the type series of Pristimantis llanganati sp. n. All specimens are adults, range and average (in parenthesis) show for two males.

Characters	Females (n = 1)	Males (n = 2)
SVL
TL
FL
HL
HW
ED
IOD
EW
IND
TLSVL
FLSVL
HLSVL
HWSVL
HWHL
ED
EWIOD
ED

Coloration in life (Fig. 8): Based on digital photographs of adult male (QCAZ 46227) and of adult female (QCAZ 46221). Dorsum olive green with an X-shaped mark in the mid-dorsum (QCAZ 46221) or with a light brown blotch on the scapular region with a rhomboidal dark brown mark and dark brown flecks (QCAZ 46227) limbs bearing transversal broad dark brown bars, posterior and anterior surfaces of thighs and concealed surfaces of shanks white with dark brown bars; top of head light brown to brown with a broad dark brown interorbital bar; sides of head greenish brown with a dark brown canthal stripe, two broad dark brown labial bars and dark brown supratympanic fold; flanks, including the groins white to whitish cream with broad dark brown diagonal stripes; anterior and posterior surfaces of thighs, concealed surfaces of shanks and dorsal surfaces of limbs are white with dark brown transversal bars. Ventrally, throat and chest are dirty cream with brown mottling, belly and ventral surfaces of thighs are dirty cream with brown flecks; ventral surfaces of tarsus, forelimbs, palms, and soles brown with minute cream flecks. Iris cooper with dark brown reticulations and a reddish midhorizontal band. Sexual dimorphism in morphology could not be evaluated due to the small sample of adult individuals (n = 3; one female and two males). Figure 8. Coloration in life of Pristimantis llanganati sp. n. A dorsal view B ventral view. From left to right: QCAZ 46227, adult male, SVL = 24.0 mm; QCAZ 46221, adult female, SVL = 29.8 mm. Pictures are not scaled. Distribution , natural history, and conservation status. Pristimantis llanganati is known from three localities (elevation range is 2253-2883 m) from Provincia del Napo, Parque Nacional Llanganates, along the Salcedo-Tena road. Maximum airline distance between localities is 6.5 km. Ecosystem type is Evergreen Montane Forest of the Eastern Andean Cordillera (as defined by Ministerio de Ambiente del Ecuador 2013 ) or Eastern Montane Forest (as defined by Ron et al. 2016 ). All specimens were collected at night on vegetation. Four of them were in a flooded area in forest border. Two were in primary forest. A deforestation map by Ministerio de Ambiente (2013) shows continuous forest at the known localities. Because we lack population data and most of the Llanganates region is unexplored, we assign Pristimantis llanganati to the Data Deficient Red List category (based on IUCN 2001 guidelines). Figure 9. Map showing known localities for Pristimantis llanganati sp. n. and Pristimantis yanezi sp. n. Circles are for Pristimantis llanganati sp. n. and triangles for Pristimantis yanezi sp. n. Localities are based on type specimens deposited at the QCAZ collection (see Suppl. material 1 for a list). The limit of Llanganates National Park is shown in red. Etymology . The species name llanganati is a noun that refers to the kichwa word "Llanganati" that means "beautiful hill". This word also gives name to Llanganates National Park, the area where the species was discovered. Many areas in the park are difficult to access and are biologically unexplored. This inaccessibility has protected large areas of its paramos and montane forests, a valuable asset for the conservation and Andean biodiversity. Remarks. As in Pristimantis yanezi , Pristimantis llanganati is not assigned to a species group until genetic data allows determining its phylogenetic position.