Pristimantis gloria, 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

Pristimantis gloria sp. nov.

Common name.

English: Gloria’s Rain Frog. Spanish: Cutín de Gloria.

Holotype.

QCAZ 57201, a gravid female from Cochapata, Azuay Province, Ecuador (3.4284S, 79.0699W, 3100 m), collected by Edwin Carrillo on May 1, 2014. Figures 15A , 16A .

Paratypes

(59: 14 females, 24 males, 21 juveniles). Ecuador: Azuay Province: QCAZ 3051-052, QCAZ 3054, adult males, QCAZ 3053, juvenile, from 7 km E Sigsig (3.0973S, 78.7970W, 2890 m); QCAZ 3055, juvenile, from 8 km E Sigsig (3.1025S, 78.7929W, 2920 m); QCAZ 3056, juvenile, from 3 km Sigsig, Sigsig-Las Minas road (3.0750S, 78.8000W, 2460 m), collected by Luis Coloma, Oscar Delgado and Luis Eduardo López on July 25, 1989; QCAZ 3452, male, QCAZ 3451, juvenile, from 23 km SE Sigsig (3.1794S, 78.7993W, 3318 m), collected by Stella de la Torre, Luis Coloma, and Felipe Campos on November 27, 1992; QCAZ 16449, adult female, QCAZ 16452, adult male, QCAZ 16447-448, juveniles, collected by Luis Eduardo López on September 2001; QCAZ 45289, adult male, collected by Luis Eduardo López on August 28, 2009; QCAZ 56533, QCAZ 56536, adult females, QCAZ 56534, juvenile, collected by Luis Eduardo López on December 6, 2013; QCAZ 62779, QCAZ 62783, QCAZ 62788, QCAZ 62791, adult females, QCAZ 62778, QCAZ 62782, QCAZ 62787, adult male, QCAZ 62780, QCAZ 62789, juveniles, collected by Luis Eduardo López on December 13, 2015, from Patococha (3.0040S, 78.6658W, 3416 m); QCAZ 29133, juvenile, from Matanga (3.1779S, 78.8000W, 3325 m), collected by Luis Coloma on December 1, 1991; QCAZ 30737, female, from 11.1 km W Azuay-Morona Santiago border (2.9480S, 78.7120W, 3110 m), collected by Luis Coloma and John Wiens on April 20, 1990; QCAZ 30738, adult male, QCAZ 30739, juvenile, from 8.1 km W Azuay-Morona Santiago border (2.9640S, 78.7020W, 3140 m), collected by John Wiens on April 20, 1990; QCAZ 30740-745, juveniles from 10 km S Cutchil (3.1340S, 78.8130W, 2900 m), collected by Luis Coloma and John Wiens on April 29, 1990; QCAZ 56463, adult male from Chiguinda, Páramo de Matanga (3.1885S, 78.7922W, 3327 m), collected by Jorge Brito on October, 2013. Loja Province: QCAZ 31455, QCAZ 31457, adult females, from Fierro Urcu (3.7104S, 79.3049W, 3439 m), collected by Ítalo G. Tapia on March 2006. Morona Santiago Province: QCAZ 17143, juvenile female, from Gualaceo-Macas road (3.0097S, 78.6650W, 3462 m), collected by Omar Torres, Jennifer Pramuk and Lena Echelle on September 14, 2001; QCAZ 23933, adult female, QCAZ 23929-932, adult males, from 52 km N El Ideal (3.1831S, 78.7961W, 3525 m), collected by Santiago Ron and Giovanna Romero on April 10, 2003; QCAZ 26367, QCAZ 26370, adult females, QCAZ 26368-369, QCAZ 26372-373, adult males, QCAZ 26371, QCAZ 26375, juveniles, from Gualaceo-Plan de Milagro road (3.0007S, 78.6616W, 3372 m), collected by Andrés Merino, Ítalo Tapia and Erik Wild on August 10, 2003; QCAZ 27280, adult female, QCAZ 27276-279, QCAZ 27281, adult males, from Gualaceo-Plan de Milagro road (3.0029S, 78.6572W, 3096 m), collected by David A. Kizirian and Juan Carlos on November 2, 2000; QCAZ 40784, juvenile, from Sigsig-Gualaquiza road (3.1850S, 78.7962W, 3322 m), collected by Omar Torres, Ernesto Arbeláez, Amaranta Carvajal and David Salazar on June 17, 2008.

Referred specimens.

KU 218035 from 8.1 km W Morona Santiago border, Gualaceo-Limón road (2.9645S, 78.7016W, 3145 m) Azuay Province, Ecuador.

Diagnosis.

A species of Pristimantis having the following combination of characters: (1) skin on dorsum tuberculate to warty; tubercles and warts are more prominent in posterior dorsum; middorsal fold present; dorsolateral folds absent; skin on flanks bears large warts and more prominent tubercles than those on dorsum; skin on venter coarsely areolate; (2) tympanic membrane and tympanic annulus prominent, its upper and posterolateral margin concealed by supratympanic fold; (3) snout moderately long, subacuminate in dorsal view, rounded in profile; with or without a small papilla at the tip; (4) upper eyelid without conspicuous tubercles; cranial crests absent; (5) dentigerous processes of vomers low to prominent, oblique, moderately separated, posteromedial to choanae; (6) vocal slits, vocal sac and nuptial pads present in adult males; (7) Finger I shorter than Finger II; discs of digits expanded, truncate to elliptical; (8) fingers with broad lateral fringes; (9) indistinct ulnar tubercles; (10) heel bearing a low rounded tubercle, surrounded by several smaller; outer edge of tarsus bearing inconspicuous tubercles; inner edge of tarsus bearing a medium to long fold followed by small tubercles; (11) inner metatarsal tubercle ovoid, elevated, six times the size of round outer metatarsal tubercle; supernumerary tubercles low, numerous; (12) toes with broad lateral fringes; basal webbing present; Toe V longer to much longer than Toe III (disc on Toe III reaches or exceeds distal edge of penultimate subarticular tubercle on Toe IV, disc on Toe V reaches middle to distal edge of distal subarticular tubercle on Toe IV); discs on toes smaller than those on fingers, truncate to elliptical ( Fig. 7C ); (13) in life, dorsum light to dark brown with or without shades of cream, yellow or orange; groins and concealed surfaces of thighs brown, creamish brown, pinkish brown, or purplish brown with irregular cream to brown flecks or spots; background color looks like thin reticulations when flecking is dense; ventral surfaces of body cream to brownish cream; iris cream to light silver bearing wide black reticulations and a red medial streak ( Fig. 16 ); (14) average SVL in adult females: 30.1 ± 3.0 mm (26.7-35.8 mm; n = 15); in adult males: 21.7 ± 2.3 mm (16.8-24.7 mm; n = 24).

Comparison with other species.

Similar species are P. balionotus , P. chomskyi , P. lutzae sp. nov., P. multicolor sp. nov., P. percultus , and Pristimantis sp. (CCS2). Pristimantis gloria is readily distinguished from them (except P. percultus ) by the presence of wide black reticulations in its iris (thin black reticulations in the other species). Pristimantis gloria can be further distinguished from P. balionotus by having basal webbing between toes (absent in P. balionotus ) and warty flanks (tuberculate in P. balionotus ). Pristimantis chomskyi is different from P. gloria by having shagreen dorsal skin (tuberculate to warty in P. gloria ), lacking a middorsal fold (present in P. gloria ), having a larger body (males Z = -2.31490, p = 0.0206, SVL = 23.98-32.38 mm in P. chomskyi , 16.7-24.74 mm in P. gloria ), a smaller tympanum (males Z = 2.66173, p = 0.0078, TD/SVL = 4.5-4.7% in P. chomskyi , 5.1-6.2% in P. gloria ) and larger eye (males Z = -2.50743, p = 0.0122, ED/SVL = 11.1-12% in P. chomskyi , 11-13.8% in P. gloria sp. nov.), relative to body length. The dorsum and flanks of Pristimantis lutzae sp. nov. are predominantly covered with tubercles, while it is mostly covered with warts in P. gloria ; the ratio between the length and width of the head is larger in P. gloria than P. lutzae sp. nov. (males Z = -5.00826, p <0.0001, HL/HW = 96.8-114.5% in P. gloria , 90-97% in P. lutzae sp. nov.; females Z = -3.77517, p = 0.0002, HL/HW = 92-105% in P. gloria , 90-96% in P. lutzae sp. nov.). Morphometrically, P. multicolor sp. nov. is larger than P. gloria (males Z = 3.07054, p = 0.0021, SVL 16.7-24.7 mm in P. gloria , 19.7-29.7 mm in P. multicolor sp. nov.; females Z = 2.85671, p = 0.0043, SVL 26.7-35.8 mm in P. gloria , 29.3-40.5 mm in P. multicolor sp. nov.), and has a wider head (males Z = 2.23159, p = 0.0256, HW/SVL = 36.1-40.7% in P. gloria , 36.3-40.9% in P. multicolor sp. nov.; females Z = 4.13252, p <0.0001, HW/SVL = 36.1-39% in P. gloria , 39.6-42.2% in P. multicolor sp. nov.) relative to body length; furthermore, P. multicolor sp. nov. lacks a middorsal fold. Pristimantis percultus , like P. gloria , has wide black reticulations on the iris, and warts and tubercles on flanks. They can be readily recognized because P. percultus has a keel on the snout (absent in P. gloria ), cranial crests (absent in P. gloria ), and is larger (Table 5). Pristimantis sp. (CCS2) is the closest species to P. gloria ( Fig. 3 , Table 2). CCS2 has a red dorsum, a golden iris with a red medial streak and thin reticulations, and broadly expanded discs on finger ( Fig. 4M ); while the dorsum of P. gloria is light to dark brown, its iris is cream to light silver with a red medial streak and wide black reticulations. Though there is only one female available for CCS2, it has a larger body and smaller tympanum than any individual of P. gloria (Table 5). Pristimantis gloria is also similar to the Peruvian species Pristimantis pataikos (Duellman & Pramuk, 1999) from which can be distinguished by the presence of nuptial pads in adult males (absent in P. pataikos ), the presence of lateral fringes on fingers and toes (absent in P. pataikos ), the presence of vomerine odontophores (absent in P. pataikos ), and the presence of pale spots on the groins and concealed surfaces of thighs (uniformly brown in P. pataikos ) ( Duellman and Pramuk 1999).

Description of the holotype.

An adult female (QCAZ 57201). Measurements (in mm): SVL 35.8; TL 15.6; FL 16.5; HL 12.4; HW 13.5; ED 3.6; TD 1.8; IOD 3.5; EW 3.0; IND 2.5; EN 3.5; TED 1.7. Head wider than long, slightly narrower than body; snout moderately long, subacuminate in dorsal view, rounded in profile; cranial crests absent; nostrils slightly protuberant, narrow, directed anterolaterally; canthus rostralis slightly concave in dorsal view, angular in cross section; loreal region slightly concave; upper eyelid with indistinct tubercles, smaller than those on dorsum; tympanic annulus prominent, upper and posterolateral edge concealed by supratympanic fold; tympanic membrane distinct; two large, low and rounded postrictal tubercles surrounded by smaller tubercles. Choanae median, ovoid, not concealed by palatal shelf of maxillae; dentigerous processes of vomers prominent, large, oblique, moderately separated, positioned posteromedial to choanae; each vomer bearing several teeth; tongue slightly longer than wide, posteriorly notched, posterior half free.

Dorsal surfaces of body tuberculate, dorsum bearing median low rounded tubercles and scattered warts, more prominent posteriorly; skin on head shagreen; middorsal fold present; dorsolateral folds absent; skin on flanks bearing more prominent and larger tubercles and warts than dorsum; skin on chest and belly coarsely areolate, that on throat shagreen; discoidal fold present; ventral surfaces of limbs smooth, ventral surfaces of thighs coarsely areolate. Ulnar tubercles indistinct except for low antebrachial tubercle; outer palmar tubercle bifid, twice the size of ovoid thenar tubercle; subarticular tubercles prominent, rounded; median low supernumerary tubercles at base of fingers and palms; fingers bearing broad lateral fringes; Finger I shorter than Finger II; discs on fingers expanded and truncate; pads on fingers surrounded by circumferential grooves on all fingers ( Fig. 7C ).

Dorsal surfaces of hindlimbs shagreen with scattered tubercles; posterior surfaces of thighs smooth, ventral surfaces of thighs coarsely areolate; heel bearing a median, low and rounded tubercle surrounded by indistinct ones; outer edge of tarsus bearing indistinct tubercles; inner edge of tarsus bearing a median fold ending in a row of small tubercles; inner metatarsal tubercle elevated, ovoid, six times the size of round outer metatarsal tubercle; plantar surface with several small, indistinct supernumerary tubercles; subarticular tubercles prominent, rounded; toes bearing broad lateral fringes; basal webbing between toes IV and V present; discs on toes smaller than those on fingers, expanded, elliptical; all toes having pads surrounded by circumferential grooves; relative lengths of toes: I <II <III <V <IV; Toe V longer than Toe III (disc on Toe III reaches distal edge of penultimate subarticular tubercle on Toe IV, disc on Toe V reaches the middle of distal subarticular tubercle on Toe IV; Fig. 7C ). Color of holotype in life and preservative is shown in Figures 15A , 16A , respectively.

Variation.

This section is based in 60 individuals of the type series and photographs available from 15 individuals. Variation in life and preservative is shown in Figures 16 , 17 . In preservative, dorsal coloration varies from light to dark brown with or without shades of gray. Dorsum uniformly colored or bearing a pattern of parallel longitudinal stripes, irregular reticulations or pale blotches. Most individuals have scattered black spots on dorsum, especially on dorsolateral surfaces; three individuals (QCAZ 57201, QCAZ 26369, QCAZ30741) display scattered white spots. Black or brown canthal bars, supratympanic stripes, labial bars, and interorbital bars or stripes can be present in this species. Flanks have the same or slightly lighter hue than dorsum. Groins and posterior surfaces of thighs bear pale irregularly shaped small spots or flecks on a brown background; when flecking is very dense the background color resembles brown reticulations. One individual lacks these flecks and spots (QCAZ62778). Limbs bear or not transversal bars and black spots. Ventral coloration varies from cream to dirty cream. In life, dorsal coloration varies from light to dark brown with or without shades of cream, yellow or orange. When present, reticulations and longitudinal stripes are brighter colored than the rest of dorsum and bordered by thin black stripes. Dorsum usually bears black spots, rarely, white spots. When present, canthal and supratympanic bars or stripes are brown or black, interorbital bars or stripes are black, brown or yellow. Flanks are the same color as dorsum or lighter. Groins and posterior surfaces of thighs are brown, creamish brown, pinkish brown or purplish brown with irregular flecks or spots; when flecking is dense, the background color resembles thin reticulations (QCAZ 62778 is the only individual that do not present these spots). Venter varies from cream to brownish cream. Iris is cream or light silver with wide black reticulations that can cover almost completely the iris; a faint red to reddish brown medial streak is present ( Fig. 16D ); sclera varies from cream to light blue.

Coloration of holotype in preservative. Dorsum dark brown with abundant scattered black medium-sized spots, more concentrated on dorsolateral surfaces and a few scattered white spots; flanks slightly lighter than dorsum; black supratympanic stripes; dorsal surface of limbs with same background coloration as dorsum and scattered black spots; groins and concealed surfaces of thighs brown with cream flecks; ventral surface of body pale cream ( Fig. 15A ).

Coloration of holotype in life. Based on studio photographs ( Fig. 16A ). Dorsal surfaces of body grayish brown with scattered black medium-sized spots, more abundant on dorsolateral surfaces; black supratympanic stripe; few scattered white spots on dorsum and dorsal surfaces of thighs; flanks lighter than dorsum with scattered black spots; groins and concealed surfaces of thighs reddish brown with cream flecks; venter and ventral surfaces of thighs pinkish cream with cream spots coinciding with areolate pattern; throat cream; ventral surfaces of shanks, tarsus, feet and hands pinkish cream; iris silver with wide black reticulations and a reddish brown medial streak.

Distribution, natural history, and conservation status.

Pristimantis gloria is known from Inter-Andean Shrub and Paramo regions in Azuay, Loja, and Morona Santiago Provinces, between 2460 and 3525 m a.s.l. ( Fig. 1 ). Most individuals were found underneath rocks in paramos or pastures near the roadside, during day or night. Two couples in amplexus were found underneath rocks at night in April. A calling male was found in low vegetation at 19h00 on October 16, 2013.

We consider P. gloria as an Endangered species following B1ab(iii) + 2ab(iii) IUCN criteria because: (i) it is only known from five localities (sensu IUCN 2017) in non-protected areas, partly affected by human settlements, agriculture and cattle raising; (ii) its Extent of Occurrence is estimated to be less than 5000 km2 (617 km2); and (iii) its Area of Occupancy <500 km2 (44 km2).

Etymology.

The specific epithet is a patronym for Gloria Lorena Rosales Narváez and Gloria María Esmeralda Narváez, mother and grandmother of the leading author. This species is named after them in gratitude for all their love and support.

Remarks.

Because of the previous lack of molecular data, this species has been mistakenly identified as P. riveti (e.g., collections at the QCAZ museum). Here, we recognize it as a different species and assign it to the P. phoxocephalus species group. It has the lowest genetic divergence with its closest species, Pristimantis sp. CCS2; the average uncorrected p distance between them is 2%. The differences in morphology between them are so evident that we consider 2.0% as our reference value to delimit candidate species.

Figures

Figure 1. Distribution of Pristimantis, subgenus Huicundomantis. Records are based on specimens deposited at the Museum of Zoology, Pontificia Universidad Catolica del Ecuador (QCAZ). Locality information is shown in Suppl. material 2.

Figure 3. Phylogenetic relationships of Pristimantis, sugenus Huicundomantis. Maximum likelihood tree for genes 16 S, ND 1 and RAG 1. 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.

Figure 4. Morphological variation within Pristimantis, subgenus Huicundomantis. A Pristimantis philipi. Pristimantis cryptomelas group (*): B Pristimantis gagliardoi C Pristimantis muscosus D Pristimantis cryptomelas. Pristimantis phoxocephalus group (**): E Pristimantis torresi sp. nov. F Pristimantis hampatusami G Pristimantis jimenezi sp. nov. H Pristimantis phoxocephalus I Pristimantis totoroi sp. nov. J Pristimantis atratus K Pristimantis chomskyi sp. nov. L Pristimantis multicolor sp. nov. M Pristimantis sp. (CCS 2) N Pristimantis gloria sp. nov. O Pristimantis lutzae sp. nov. P Pristimantis tinguichaca Q Pristimantis verrucolatus sp. nov. R Pristimantis sp. (UCS 1) S Pristimantis teslai sp. nov. T Pristimantis versicolor U Pristimantis atillo sp. nov. Red branches are for clades without available photographs of live individuals. Not shown at the same scale.

Figure 7. Palmar and plantar surfaces of Pristimantis atillo sp. nov., Pristimantis chomskyi sp. nov., Pristimantis gloria sp. nov., and Pristimantis jimenezi sp. nov. Photographs of left hand and foot of the holotypes A Pristimantis atillo sp. nov.: QCAZ 42500, male B Pristimantis chomskyi sp. nov.: QCAZ 47515, male C Pristimantis gloria sp. nov.: QCAZ 57201, female D Pristimantis jimenezi sp. nov.: QCAZ 45170, male. All specimens are shown at the same scale.

Figure 15. Holotypes of Pristimantis gloria sp. nov. and P. jimenezi sp. nov. Photographs of preserved holotypes of A P. gloria (QCAZ 57201, female) and B P. jimenezi (QCAZ 45170, male). Dorsal view on the left, ventral view on the right. Specimens are shown at the same scale.

Figure 16. Color variation in live individuals of Pristimantis gloria sp. nov. A QCAZ 57201 (holotype, female, SVL 35.8 mm) B QCAZ 62778 (male, SVL 19.7 mm) C QCAZ 62780 (juvenile female, SVL 23.5 mm) D QCAZ 62789 (juvenile female, SVL 21.8 mm) For A-C dorsolateral view on the left, ventral view on the right. For D dorsolateral view on the left, close up of the eye in the right.

Figure 17. Color variation in preserved individuals of Pristimantis gloria sp. nov. A Dorsal view of (from left to right): QCAZ 31457 (female), QCAZ 31455 (female), QCAZ 16449 (female) B Dorsal view of: QCAZ 16448 (juvenile female), QCAZ 56463 (male), QCAZ 16452 (male), QCAZ 16447 (juvenile female), QCAZ 31461 (juvenile female) 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.