Pristimantis multicolor , 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 multicolor sp. nov.
English: Multicolored Rain Frog. Spanish: Cutín multicolor .
QCAZ 47213, an adult male from Yacuri National Park, Laguna Negra, Loja Province, Ecuador (4.7081S, 79.4322W, 3400 m), collected by Carlos Antonio Rodríguez on December 10, 2009. Figure 20BView Figure 20.
(59: 35 males, 23 females, 1 juvenile). All individuals from Yacuri National Park, nearby the type locality. Ecuador: Loja Province: QCAZ 47214, adult female, collected with the holotype; QCAZ 47215, adult male, from Laguna Negra (4.7095S, 79.4356W, 3348 m), collected by Carlos Antonio Rodríguez on December 10, 2009; QCAZ 61279, QCAZ 61390-391, QCAZ 61404, adult females, QCAZ 61392, QCAZ 61394, QCAZ 61396, adult males, from Jimbura-San Andrés road (4.7325S, 79.4352W, 3300 m); QCAZ 61283, QCAZ 61287, adult females, QCAZ 61284, QCAZ 61286, QCAZ 61297, adult males, from Laguna Negra (4.7121S, 79.4308W, 3186 m); QCAZ 61293, adult male, from Loja-Zamora border (4.7436S, 79.4234W, 3515 m); QCAZ 61299-300, QCAZ 61302-304, adult females, QCAZ 61301, QCAZ 61306, adult males, from entrance to Lagunas Coloradas, Jimbura-San Andrés road (4.7349S, 79.4283W, 3390 m); QCAZ 61310, adult female, QCAZ 61314, QCAZ 61316-317, adult males, from Laguna de los Patos (4.7204S, 79.4286W, 3247 m); QCAZ 61344, QCAZ 61346, QCAZ 61348, adult females, QCAZ 61343, adult male, from Loja-Zamora border (4.7433S, 79.4244W, 3506 m); QCAZ 61349, QCAZ 61352, QCAZ 61361, adult females, QCAZ 61362, QCAZ 61367-368, adult males, QCAZ 61351, juvenile, from Jimbura-San Andrés road (4.7277S, 79.4333W, 3411 m); QCAZ 61358, adult male, from surroundings of refuge (4.7126S, 79.4399W, 3245 m); QCAZ 61359, QCAZ 61363, QCAZ 61369-370, adult males, from Los Picachus (4.7121S, 79.4308W, 3186 m); QCAZ 61373, adult female, from Laguna Negra (4.7154S, 79.4212W, 3242 m); QCAZ 61377, QCAZ 61379-380, adult females, from Jimbura-Zumba road (4.7252S, 79.4353W, 3333 m); QCAZ 61383-384, adult females, QCAZ 61381, adult male, from Jimbura-San Andrés road (4.7309S, 79.4340W, 3332 m), collected by Francy Mora, Keyko Cruz, David Velalcázar, Andrés Calahorrano, Darwin Núñez, Juan Sánchez, Javier Pinto and Daniel Rivadeneira between April 21 and May 2, 2015. Zamora Chinchipe Province: QCAZ 61407, QCAZ 61424-425, adult females, QCAZ 61399, QCAZ 61410, QCAZ 61413, QCAZ 61427, QCAZ 61432, QCAZ 61435-436, adult males, from Laguna Colorada (4.7379S, 79.3958W, 3318 m), collected by Francy Mora, Keyko Cruz, David Velalcázar, Andrés Calahorrano, Darwin Núñez, Juan Sánchez, Javier Pinto and Daniel Rivadeneira between April 21 and May 2, 2015.
A species of Pristimantis having the following combination of characters: (1) skin on dorsum shagreen to warty; dorsolateral folds absent; skin on flanks bearing low warts, bigger than those of dorsum; skin on venter coarsely areolate; discoidal fold absent; (2) tympanic membrane and tympanic annulus prominent, its upper and posterior margin concealed by thick supratympanic fold; (3) snout moderately long, subacuminate in dorsal view, rounded in profile, with or without a papilla at the tip; (4) upper eyelid lacking tubercles; cranial crests absent; (5) dentigerous processes of vomers low to prominent, oblique, narrowly to moderately separated, posteromedial to choanae; (6) vocals slits, vocal sac, and nuptial pads present in adult males; (7) Finger I shorter than Finger II; discs of digits expanded to broadly expanded, elliptical to truncate; (8) fingers with lateral fringes; (9) ulnar tubercles present; (10) heel bearing one rounded tubercle, surrounded or not by several smaller tubercles; inner and outer edge of tarsus bearing small and low tubercles; (11) inner metatarsal tubercle ovoid, elevated six times the size of elliptical, elevated outer metatarsal tubercle; supernumerary tubercles numerous; (12) toes with lateral fringes; basal webbing present; Toe V 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 or exceeds distal edge of distal subarticular tubercle on Toe IV); toe discs smaller than those on fingers, elliptical ( Fig. 8BView Figure 8); (13) in life, dorsal surfaces vary from yellow to dark brown with or without olive or orange tones; black or brown canthal and supratympanic stripe usually present; groins and posterior surfaces of thighs cream, orange, brown, or black with or without cream to yellow spots; venter varies from cream to dark brown with a few to many small beige to yellow spots; iris copper to creamy yellow with or without a dark brown to red medial horizontal streak, with black reticulations ( Fig. 24View Figure 24); (14) average SVL in adult females: 35.3 ± 3.5 mm (29.4-40.5 mm; n = 27); in adult males: 26.2 ± 3.5 mm (19.7-29.7 mm; n = 32).
Comparison with other species.
It is most similar to P. balionotus , P. chomskyi , P. gloria , P. lutzae , and P. percultus . Pristimantis multicolor is different from P. balionotus by having basal webbing between toes (absent in P. balionotus ), larger discs on fingers and toes, and larger tubercles and warts on flanks. Morphometrically, P. chomskyi has a smaller tympanum (males Z = 2.38157, p = 0.0172, TD/SVL = 4.5-4.7% in P. chomskyi , 4.9-6.0% in P. multicolor ). Pristimantis multicolor differs from P. gloria by its larger size (males Z = 3.07054, p = 0.0021, SVL 16.7-24.7 mm in P. gloria , 19.7-29.7 mm in P. multicolor ; females Z = 2.85671, p = 0.0043, SVL 26.7-35.8 mm in P. gloria , 29.3-40.5 mm in P. multicolor ), by lacking a middorsal longitudinal fold (present in P. gloria ), and having thinner black reticulations on the iris; additionally, P. multicolor has a proportionally 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 ; females Z = 4.13252, p < 0.0001, HW/SVL = 36.1-39% in P. gloria , 39.6-42.2% in P. multicolor ). Compared to P. lutzae , Pristimantis multicolor sp. nov. has a longer head (males Z = 3.67756, p = 0.0002, HL/SVL = 33.4-37% in P. lutzae , 34.1-40.4% in P. multicolor ; females Z 3.9524, p < 0.0001, HL/SVL = 35-37.5% in P. lutzae , 36.5-40.6% in P. multicolor ), larger tympanum (males Z = 3.57469, p = 0.0004, TD/SVL = 5-5.4% in P. lutzae , 4.9-6% in P. multicolor ; females Z = 3.9524, p < 0.0001, TD/SVL = 4.9-5.6% in P. lutzae , 5.5-6.7% in P. multicolor ) and larger eyes (males Z = 2.75174, p = 0.0059, ED/SVL = 10.3-12.1% in P. lutzae , 10.3-13.2% in P. multicolor ; females Z = 3.3083, p = 0.0009, ED/SVL = 9.9-12.1% in P. lutzae , 10.7-12.4% in P. multicolor ), relative to body size. Pristimantis percultus is easily distinguished from P. multicolor by having low cranial crests (absent in P. multicolor ) and a red stripe on the upper lip.
Description of the holotype.
An adult male ( QCAZ 47213, SC29908). Measurements (in mm): SVL 29.3; TL 14.8; FL 14.5; HL 11.0; HW 11.5; ED 3.5; TD 1.8; IOD 3.7; EW 2.8; IND 2.5; EN 3.1; TED 1.9. Head wider than long, wider than body; snout moderately long, subacuminate in dorsal view, rounded in profile; cranial crests absent; nostrils slightly protuberant, directed anterolaterally; canthus rostralis concave in dorsal view, prominent and rounded in cross section; loreal region concave; upper eyelid with indistinct tubercles; tympanic membrane distinct; tympanic annulus prominent, upper and posterior edge concealed by thick, heavy supratympanic fold; three large rounded postrictal tubercles. Choanae median ovoid, not concealed by palatal shelf of maxillae; dentigerous processes of vomers prominent, oblique, moderately separated, positioned posteromedial to choanae; each vomer bearing several teeth; tongue slightly longer than wide, posterior border notched, posterior three fifths not adherent to floor of mouth; vocal slits slightly curved, located at posterior half of mouth floor in between tongue and margin of jaw; vocal sac present.
Dorsal surfaces of body shagreen with scattered low warts; dorsolateral folds absent; skin on flanks bearing rounded warts, larger than those on dorsum; skin on venter coarsely areolate, ventral surfaces of limbs smooth, ventral surfaces of thighs coarsely areolate; discoidal fold absent. Low, round ulnar tubercles, antebrachial tubercle prominent; white nuptial pads present; outer palmar tubercle bifid, almost twice the size of ovoid thenar tubercle; subarticular tubercles prominent, rounded; ill-defined supernumerary tubercles; fingers bearing lateral fringes; Finger I shorter than Finger II; discs on fingers broadly expanded, truncate; pads on fingers surrounded by circumferential grooves on all fingers ( Fig. 8BView Figure 8).
Hindlimbs slender; dorsal surfaces of hindlimbs shagreen; posterior surfaces of thighs smooth, ventral surfaces of thighs coarsely areolate; heel bearing a medium sized, low, rounded tubercle surrounded by several smaller tubercles; outer and inner edge of tarsus bearing low rounded tubercles; inner metatarsal tubercle ovoid, elevated, six times the size of elliptical, elevated outer metatarsal tubercle; plantar surface bearing numerous ill-defined supernumerary tubercles; subarticular tubercles prominent, subacuminate; toes bearing lateral fringes; basal webbing between toes IV and V present; discs on toes smaller than those on fingers, expanded and 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 distal edge of penultimate subarticular tubercle on Toe IV, disc on Toe V reaches the distal edge of distal subarticular tubercle on Toe IV; Fig. 8BView Figure 8). Coloration of the holotype in preservative is shown in Figure 20BView Figure 20; coloration in life, unknown.
Coloration of holotype in preservative. Dorsum dark brown covered with minute light brown irregular spots; dorsal surfaces of forelimbs light brown with scattered minute dark brown flecking; dorsal surfaces of hindlimbs dark brown with light brown spots becoming larger and more abundant posteriorly; groins and concealed surfaces of thighs, shanks and tarsus brown; ventral surfaces of body dusty brown; chest with cream irregular blotches; vocal sac and ventral surfaces of thighs cream; ventral surfaces of fingers and toes pale cream ( Fig. 20BView Figure 20).
Coloration of holotype in life. Unknown.
Based on 60 preserved specimens and photographs from 57 individuals. Variation of live and preserved individuals is shown in Figures 24View Figure 24, 25View Figure 25. Texture of skin varies among individuals. Dorsum varies from shagreen to warty, with very low warts (not tuberculate); flanks vary from tuberculate to warty, warts on flanks are larger than those of dorsum; tubercles on ulna, shanks, and tarsus can be distinct or inconspicuous. Coloration in life is highly variable ( Fig. 24View Figure 24). Dorsal surfaces can be yellow, cream, orange, or pale to dark brown with or without olive or orange tones. They can have few to abundant black spots on dorsum, flanks, and dorsal surfaces of limbs. Dorsum occasionally bears pale scapular or sacral blotches, a middorsal stripe, or white spots. Head usually bears black or brown canthal and supratympanic stripes, sometimes an interorbital bar. Groins and posterior surfaces of thighs are cream, orange, reddish brown, orangey brown, brown, dark brown, or black with or without cream to yellow spots. The background color of the venter varies from cream to dark brown with different levels of transparency; small beige to yellow blotches can be present on the belly, chest, and throat, but in some individuals are restricted to the chest. Iris is copper, golden, creamy yellow or creamy brown with or without a dark brown to red medial horizontal streak; black reticulations are present; sclera can be light brown, cream, white, or light blue.
Distribution, natural history, and conservation status.
Pristimantis multicolor is known from Parque Nacional Yacuri, near the border with Peru, at Loja and Zamora Chinchipe Provinces ( Fig. 1View Figure 1). It inhabits Paramo and Eastern Montane Forest between 3186 and 3515 m. During the late afternoon and night, individuals were commonly found beneath or over moss, mat-forming plants, tussock grasses and rocks, perched on branches and leaves of shrubs up to 1.75 m above ground, or inside bromeliads, terrestrial or epiphytes, up to 1 m above the ground. During the day they were found inside bromeliads. Calling males and gravid females were found in December, April, and May; males were calling on low branches of shrubs.
According to available data, P. multicolor has a very restricted distribution (Extent of Occurrence = 10 km2) but is locally abundant. However, we propose assigning it to the Data Deficient Red List Category ( IUCN 2017) because extensive areas in Yacuri National Park remain unexplored and represent potential distribution for this species.
The specific epithet comes from the Latin words multus meaning many, and color meaning color. It refers to the wide range of color variation in this species ( Fig. 24View Figure 24).
This species was collected for the first time in 2015 by personnel from the QCAZ museum during expeditions to Yacuri National Park. It was identified as Pristimantis aff. riveti . Here, we recognize it as a different species and assign it to the P. phoxocephalus species group.
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