AnimaliaAnuraStrabomantidaePáezNadia B.RonSantiago R.Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new speciesZookeys0182019868111210.3897/zookeys.868.267664FFD270148D35548960B3C83B72FEC13 Pristimantis torresi http://zoobank.org/1E92D590-2072-4530-88C6-2019A0A6D8F9 sp. nov.Common name.

English: Torres’ Rain Frog. Spanish: Cutín de Torres.

Holotype.

QCAZ 47342, adult female from Guanchanamá, military antennas, Loja Province, Ecuador (4.0511S, 79.8693W, 2729 m), collected by Elicio E. Tapia and Margarita Baquero on February 21, 2010. Figure 28B.

Paratypes

(46: 16 males, 6 females, 24 juveniles). Ecuador: Loja Province: QCAZ 47343, QCAZ 47354, adult females, QCAZ 47344, adult male, QCAZ 47340–341, QCAZ 47345–351, juveniles, collected with the holotype; QCAZ 47397, QCAZ 47405, QCAZ 47409, QCAZ 47414, QCAZ 47416–418, QCAZ 47503, adult males, from Celica-Alamor road (4.0990S, 79.9799W, 2101 m), collected by Elicio E. Tapia and Freddy Velásquez on February 27, 2010; QCAZ 64524, adult male, QCAZ 64525, QCAZ 64527–529, QCAZ 64555, juveniles, from Guanchanamá, military antennas (4.0511S, 79.8693W, 2729 m), collected by Nadia Páez, Fernando Ayala, Pablo Sandoval and Ricardo Gavilanes in August 2016; QCAZ 64533, QCAZ 64538, juveniles, from Celica-Guachanamá road (4.0907S, 79.9509W, 2348 m), collected by Nadia Páez, Fernando Ayala, Pablo Sandoval and Ricardo Gavilanes in August 2016; QCAZ 64544, juvenile from Guachanamá-La Tolera road (4.0321S, 79.8836W, 2736 m), collected by Nadia Páez, Fernando Ayala, Pablo Sandoval and Ricardo Gavilanes in August 2016; QCAZ 64549, adult male, QCAZ 64551, QCAZ 64556, juveniles, from Celica–Alamor road (4.0974S, 79.9796W, 2117 m), collected by Nadia Páez, Fernando Ayala, Pablo Sandoval and Ricardo Gavilanes in August 2016. Nearby Guachanamá, collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Santiago Guamán and Guadalupe Calle in November 2016: QCAZ 65762, adult male, (4.0441S, 79.8814W, 2664 m); QCAZ 65766–767, adult females (4.0410S, 79.8827W, 2770); QCAZ 65769, adult male (4.0408S, 79.8829W, 2787 m); QCAZ 65771, adult male (4.0431S, 79.8817W, 2738 m); QCAZ 65774, QCAZ 65776, adult males (4.0441S, 79.8814W, 2646 m). Nearby La Tolera, collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Santiago Guamán and Guadalupe Calle in November 2016: QCAZ 65779, adult female (4.0317S, 79.8824W, 2737 m); QCAZ 65782, adult female, QCAZ 65780, juvenile (4.0326S, 79.8836W, 2752 m); QCAZ 65783–786, juveniles (4.0395S, 79.8848W, 2833 m).

Diagnosis.

A member of the Pristimantis phoxocephalus group characterized by the following combination of characters: (1) skin on dorsum shagreen; thin middorsal fold; head with a middorsal small tubercle or row of tubercles; dorsolateral folds absent; flanks with longitudinal lateral folds on anterior half; skin on venter areolate to weakly 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 a small and rounded tubercle, surrounded by several lower tubercles; cranial crests absent; (5) dentigerous processes of vomers prominent, oblique, moderately to broadly separated, posteromedial to choanae; (6) vocals slits, vocal sac and nuptial pads present; (7) Finger I shorter than Finger II; discs of digits broadly expanded, elliptical to truncate; (8) fingers with lateral fringes; (9) ulnar tubercles ill-defined; (10) heel bearing a small, rounded tubercle surrounded or not by smaller tubercles; outer and inner tarsal tubercles absent or ill-defined; inner tarsal fold present; (11) inner metatarsal tubercle ovoid, rounded, elevated, six 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 middle or exceeds distal edge of penultimate subarticular tubercle on Toe IV; disc on Toe V reaches the middle or exceeds distal edge of distal tubercle on Toe IV); toe discs as large or slightly smaller than those on fingers (Fig. 9B); (13) in life, dorsum cream, brown or orangey brown; head bearing a dark supratympanic stripe and interorbital band or stripe; coloration of groins and concealed surfaces of thighs have sexual dimorphism: in females (and juveniles), ground color varies from light purplish brown to medium brown with or without minute cream flecks on; males have the same ground color with yellow, cream or orangey-yellow small to big spots; venter white to dusty cream; iris golden to beige with black reticulations and a red or reddish brown medial streak (Fig. 31); (14) average SVL in adult females: 34.7 ± 3.7 mm (30.1–39.5 mm; n = 5); in adult males: 25.9 ± 2.1 mm (23.3–30.0 mm; n = 18).

10.3897/zookeys.868.26766.figure31a2676c2d-3254-55f8-9b85-92894294e9cc

Color variation in live individuals of Pristimantis torresi sp. nov. AQCAZ 64524 (male, SVL 26.7 mm) BQCAZ 65767 (female, SVL 35.8 mm) CQCAZ 65771 (male, SVL 28.7 mm) DQCAZ 65779 (male, SVL 26.4 mm). Dorsolateral view on the left, ventral view on the right.

https://binary.pensoft.net/fig/322423
Comparison with other species.

Pristimantis torresi is similar to P. atillo, P. jimenezi, P. phoxocephalus, P. teslai, P. totoroi sp. nov., and P. verrucolatus sp. nov., which also have an acuminate snout with a fleshy keel. Pristimantis atillo usually has orange groins and black dots on the flanks, instead, P. torresi has brown groins with or without yellow spots, and its flanks lack black dots. The most similar species to P. torresi is P. jimenezi, whose iris varies from copper to red (golden to beige with a red medial streak in P. torresi). Pristimantis torresi differs from P. phoxocephalus in having a golden to beige iris (copper in P. phoxocephalus), brown groins with or without light brown to yellow spots (yellow with black reticulations in P. phoxocephalus), and a wider head relative to its body (males Z = -2.56285, p = 0.0104, HW/SVL = 33.4–34.4% in P. phoxocephalus, 34.2–37.8% in P. torresi; females Z = -2.08893, p = 0.0367, HW/SVL = 33.1–37.3% in P. phoxocephalus, 36.7–38.5% in P. torresi). Pristimantis torresi is easy to distinguish from P. teslai by having shagreen dorsal skin, lateral folds, and golden to beige iris with a red to reddish-brown medial streak (tuberculate dorsal skin, lateral folds absent and copper iris in P. teslai). Pristimantis totoroi sp. nov. has more prominent tubercles and folds than those of P. torresi, and its head is longer (males Z = 3.84623, p = 0.0001, HL/HW = 95.1–102.3% in P. torresi, 99.7–104% in P. totoroi sp. nov; females Z = -2.76079, p = 0.0058, HL/HW = 90.6–94.7% in P. torresi, 95.6–103.2% in P. totoroi sp. nov), relative to head width. Pristimantis torresi differs from Pristimantis verrucolatus sp. nov. in lacking large tubercles and warts on the flanks (present in P. verrucolatus) and having a golden to beige iris (coppery brown in P. verrucolatus).

Description of the holotype.

Adult female (QCAZ 47342, SC32171). Measurements (in mm): SVL 35.8; TL 17.5; FL 16.4; HL 12.4; HW 13.2; ED 3.7; TD 1.8; IOD 3.8; EW 3.1; IND 2.8; EN 3.5; TED 0.8. Head wider than long, narrower than body; snout moderately long, acuminate with a fleshy keel in dorsal view, protruding in profile; nostrils slightly protuberant, directed anterolaterally; canthus rostralis distinct, curved in dorsal view, rounded in cross-section; loreal region concave; upper eyelid bearing a small rounded tubercle surrounded by several lower tubercles; tympanic membrane and tympanic annulus distinct, its upper and posterior margin covered by supratympanic fold; one enlarged subconical postrictal tubercle surrounded by several smaller tubercles. Choanae median, semicircular, not concealed by palatal shelf of maxilla; dentigerous processes of vomers prominent, oblique, moderately separated, posteromedial to choanae; each vomer bearing several indistinct teeth; tongue as wide as long, posteriorly notched, free on posterior half of its length.

Skin on dorsum shagreen; faint middorsal fold; head with a middorsal row of two small tubercles; dorsolateral folds absent; flanks areolate, with thin lateral folds on anterior half; skin on belly and chest areolate, skin on throat, chest and ventral surfaces of limbs smooth; discoidal fold present. Ulnar tubercles present, indistinct; palmar tubercles prominent, outer palmar tubercle bifid, almost twice size of ovoid thenar tubercle; subarticular tubercles prominent, rounded; supernumerary tubercles at base of fingers distinct; fingers with broad lateral fringes; Finger I shorter than Finger II; discs broadly expanded and rounded; pads on fingers surrounded by circumferential grooves on all fingers (Fig. 9B).

Hindlimbs slender; dorsal surfaces of hindlimbs smooth; posterior surfaces of thighs smooth, ventral surfaces of thighs areolate; heel bearing a low rounded tubercle surrounded by some smaller tubercles; outer tarsal tubercles absent; inner tarsal fold extend to half of length of tarsus; inner metatarsal tubercle ovoid, rounded, elevated, six times the size of rounded, ill-defined outer metatarsal tubercle; plantar surface with supernumerary tubercles, those on the base of toes low but distinct; subarticular tubercles prominent, rounded; toes with broad lateral fringes; basal webbing present; discs nearly as large as those on fingers, elliptical; all toes having 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 the middle of the penultimate subarticular tubercle on Toe IV, disc on Toe V exceeds distal edge of distal subarticular tubercle on Toe IV; Fig. 9B). Coloration of the holotype in preservative is shown in Figure 28B; coloration in life is unknown.

Coloration of holotype in preservative. Dorsal surface of body light gray; head with cream interorbital band outlined with black, dark brown supratympanic stripes, and faint gray labial bars; limbs with transversal bands slightly darker than background; dorsolateral surfaces with oblique reticulations extending to the flanks, slightly darker than background; groins and concealed surfaces of thighs gray with small cream spots; ventral surfaces of body white, venter with faint gray flecks and midventral longitudinal line; plants and palms dusty cream (Fig. 28B).

Coloration of holotype in life. Unknown.

Variation.

Based on the 47 preserved specimens of the type series and photographs for 24 individuals. Variation in living and preserved individuals is shown in Figures 31, 32. Coloration in life is given in parenthesis. Dorsum coloration varies between cream, light gray and brown (cream, brown or orangey-brown). Some individuals present dark brown reticulations, chevrons, flecks, a middorsal band, an hourglass-shaped band or a thin white middorsal stripe on dorsum. Head always bears brown supratympanic stripes and interorbital stripes or bands; sometimes, it bears brown canthal stripes or labial bars. Limbs present faint or well-defined transversal bands. Flanks may have diagonal bars. Coloration of groins and concealed surfaces of hindlimbs has sexual dimorphism: both sexes have cream, gray or brown backgrounds (light purplish brown to medium brown); females have none or little pale (cream) flecks; males have small or big pale (yellow, cream or orangey yellow) spots, usually more conspicuous in larger males. Venter varies from white to dusty cream, usually with gray or light brown markings and a midline (in males, throat cream, light or bright yellow). Iris is golden to beige with black reticulations and a red or reddish-brown medial streak; sclera varies from cream to light blue. Coloration in life of each individual may change; the ground color of the hidden surfaces of thighs and other markings as the supratympanic stripe can vary from light purplish brown to medium brown in the same individual.

10.3897/zookeys.868.26766.figure329b6851e3-2478-59ac-b00b-3c87abbd63fb

Color variation in preserved individuals of Pristimantis torresi sp. nov. A Dorsal view of (from left to right): QCAZ 65767 (female), QCAZ 65762 (male), QCAZ 65769 (male), QCAZ 64524 (male) B Dorsal view of: QCAZ 64549 (male), QCAZ 65782 (male), QCAZ 47397 (male), QCAZ 47416 (male) C Ventral view of specimens in (A) D Ventral view of specimens in B. See Suppl. material 2 for locality data. All specimens are shown at the same scale.

https://binary.pensoft.net/fig/322424
Distribution, natural history, and conservation status.

This species is known from the surroundings of Celica, Guachanamá and La Tolera, towns in the western Andean slopes of Loja Province in Ecuador (Fig. 2). It inhabits Inter-Andean Shrub region between 2101 and 2833 m a.s.l. Most individuals were found at night inside terrestrial or arboreal bromeliads up to 3 m above the ground, inside patches of native vegetation, or on low vegetation up to 60 cm above ground. During the day they were only found inside bromeliads. Calling males have been found on low vegetation, 30 cm above the ground in February.

We propose assigning P. torresi to the Critically Endangered Red List category following the B1ab(iii) IUCN criteria. Available records come from two localities (sensu IUCN 2017) whose habitat is being degraded by human settlements, cattle raising and agriculture; the Extent of Occurrence of the species < 100 km2 (21 km2).

Etymology.

The specific epithet is a noun in the genitive case and is a patronym for Omar Torres-Carvajal, curator of reptiles of Museo de Zoología at Pontificia Universidad Católica del Ecuador. The species name is in recognition of his significant contributions to herpetological research in Ecuador and the development of collections at the QCAZ museum.

10.3897/zookeys.868.26766.figure28a75c2fae-d27e-5dd6-b9ba-a7166e19c1f1

Holotypes of Pristimantis teslai sp. nov. and P. torresi sp. nov. Photographs of preserved holotypes of AP. teslai (QCAZ 46213, male) and BP. torresi (QCAZ 47342, female). Dorsal view on the left, ventral view on the right. Specimens are shown at the same scale.

https://binary.pensoft.net/fig/322420
10.3897/zookeys.868.26766.figure9241560b7-f74e-5107-aa85-844e6ba2c761

Palmar and plantar surfaces of Pristimantis teslai sp. nov., Pristimantis torresi sp. nov., Pristimantis totoroi sp. nov., and Pristimantis verrucolatus sp. nov. Photographs of hand and foot of the holotypes of the following species: APristimantis teslai sp. nov.: QCAZ 46213, male BPristimantis torresi sp. nov.: QCAZ 47342, female CPristimantis totoroi sp. nov.: QCAZ 25105, male DPristimantis verrucolatus sp. nov.: QCAZ 46982, male. All specimens are shown at the same scale.

https://binary.pensoft.net/fig/322401
10.3897/zookeys.868.26766.suppl28e002865-cf8c-568c-8d26-9cca25ce0347

Examined specimens

Data type: occurrence

Explanation note: Museum specimens analyzed in this taxonomic review. *: analyzed only with molecular information. ** = analyzed only with photographic material.

https://binary.pensoft.net/file/322433This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Nadia B. Páez, Santiago R. Ron
10.3897/zookeys.868.26766.figure2434c89fd-e411-50ef-a562-c9b8a72a3f27

Distribution of Pristimantis, subgenus Huicundomantis. Records are based on specimens deposited at the Museum of Zoology, Pontificia Universidad Católica del Ecuador (QCAZ). Candidate species numbers correspond to those of the text and Figure 3. Locality information is shown in Suppl. material 2. Abbreviations: CCS = confirmed candidate species, UCS = unconfirmed candidate species.

https://binary.pensoft.net/fig/322394
10.3897/zookeys.868.26766.figure3d26edbc5-acfb-5bd0-afec-e670e6536748

Phylogenetic relationships of Pristimantis, sugenus Huicundomantis. Maximum likelihood tree for genes 16S, ND1 and RAG1. Bootstrap values (%) followed by Bayesian posterior probabilities are shown under the corresponding branches. Asterisks indicate support values of 100 (bootstrap) or 1 (posterior probabilities); missing values indicate values below 50 (bootstrap) or 0.5 (posterior probability). The collection number, identification, province and locality of the samples are shown next to each terminal; all samples are from Ecuador. Outgroup is not shown. Abbreviations: CCS = confirmed candidate species, NP = national park, PF = protected forest, UCS = unconfirmed candidate species.

https://binary.pensoft.net/fig/322395
IUCN Standards and Petitions Subcommittee (2017) Guidelines for Using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. http://www.iucnredlist.org/documents/RedListGuidelines.pdf. [Accessed on: 2017-12-15]