The Pristimantis trachyblepharis species group, a clade of miniaturized frogs: description of four new species and insights into the evolution of body size in the genus Author Zumel, Daniel Author Buckley, David Author Ron, Santiago R text Zoological Journal of the Linnean Society 2022 2022-05-01 195 1 315 354 https://academic.oup.com/zoolinnean/article/195/1/315/6359884 journal article 55607 10.1093/zoolinnean/zlab044 76944e29-2322-4e8e-aa72-ac66ea956837 0024-4082 6530571 PRISTIMANTIS UJUCAMI SP. NOV. ( FIGS 15–19 ; TABLES 2 AND 3 ) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 0FCACB6D-7A6F-45C8-8D5A-D35AE2AD51DD Holotype ( Figs 15 , 16 ): QCAZ 49030 (field no. YSNA-130), adult female from Ecuador , Provincia Morona Santiago , Cantón Morona , Parroquia Proaño , on the eastern border of Abanico Protected Forest , near the Hidroabanico Hydroelectric Power Plant ( 2.2590º S , 78.1950º W ), 1720 m a.s.l. , collected by Yerka Sagredo and Rubén Jarrín on 25 July 2010 . Paratypes (30: nine males , 21 females ): Ecuador : Provincia Morona Santiago : Abanico Protected Forest, QCAZ 49026, adult female, and QCAZ 49031, subadult female ( 2.25901º S , 78.1950º W ), 1720 m a.s.l.; QCAZ 56998, adult male ( 2.2576º S , 78.1986º W ), 1542 m a.s.l.; QCAZ 57001–57002, adult males ( 2.2579º S , 78.1985º W ), 1555 m a.s.l.; QCAZ 57003, adult male ( 2.2581º S , 78.1986º W ), 1526 m a.s.l.; QCAZ 57004– 57005, subadult females ( 2.2582º S , 78.1984º W ), 1564 m a.s.l.; QCAZ 57008, adult female ( 2.2582º S , 78.1984º W ), 1573 m a.s.l.; QCAZ 57010, subadult female ( 2.2583º S , 78.1984º W ), 1550 m a.s.l. Collected by Y. Sagredo, R. Jarrín, F. Ayala, S. Arroyo, S. Valverde and L. Cedeño on 23–25 July 2010 and 2 March 2014 ; Volcano River, Sangay National Park. QCAZ 58865, adult female, and QCAZ 58874, subadult female ( 2.0983º S , 78.5554º W ), 1406 m a.s.l.; QCAZ 58866, subadult female ( 2.0987º S , 78.1560º W ), 1403 m a.s.l.; QCAZ 58868, subadult female ( 2.0903º S , 78.1897º W ), 1551 m a.s.l.; QCAZ 58870, adult male ( 2.0971º S , 78.1553º W ), 1411 m a.s.l. Collected by D. Rivadeneira, F. Mora, J.C. Sánchez, and A. Correa, on 3 February 2015 . Provincia Zamora Chinchipe : Numbami Reserve, QCAZ 57646, adult male ( 4.1672º S , 78.9489º W ), 1458 m a.s.l. Collected by S. Ron, D. Paucar, P. Venegas, D. Almeida, D. Velalcázar, M.J. Navarrete, S. Arroyo, N. Páez and Z. Lange, on 8 July 2014 ; Bombuscaro River, Podocarpus National Park. QCAZ 60194, adult female ( 4.1146º S , 78.9670º W ), 981 m a.s.l.; QCAZ 60195–60196, adult females ( 4.1250º S , 78.9788º W ), 1164 m a.s.l.; QCAZ 60200, subadult male, and QCAZ 60202, adult female ( 4.1146º S , 78.9633º W ), 1112 m a.s.l.; QCAZ 60205, QCAZ 60612 and QCAZ 60214, adult females ( 4.1344º S , 78.9938º W ), 1443 m a.s.l.; QCAZ 60206–60207, subadult females ( 4.1344º S , 78.9938º W ), 1443 m a.s.l.; QCAZ 60216, adult female ( 4.1333º S , 78.9854º W ), 1281 m a.s.l.; QCAZ 60218, subadult female ( 4.1333º S , 78.9854º W ), 1281 m a.s.l.; QCAZ 60621 and QCAZ 60629, adult males ( 4.1333º S , 78.9854º W ), 1281 m a.s.l. Collected by D. Rivadeneira, F. Mora, J.C. Sánchez, D. Velalcázar, D. Núñez, J. Pinto, K. Cruz and L. Tipantiza on 8 April, 2015. Figure 15. Photographs in live of the holotype of Pristimantis ujucami (QCAZ 49030; adult female, SVL 21.16 mm). A, dorsolateral view; B, ventral view; C, dorsolateral and inguinal region view. Suggested common name: English: Ujukam rain frog. Spanish: cutín de Ujukam . Diagnosis ( Figs 15–19 ): A species of Pristimantis characterized by the following combination of characters:(1) dorsal skin slightly shagreen to shagreen, bedecked with pustules and/or spicules; scapular folds distinct,> <, \ / or) (-shaped, with scapular subconical or conical tubercles along them; scapular folds may join posteriorly and extend as diffuse paravertebral folds; ventral skin weakly areolate on throat and chest, areolate on belly and coarsely areolate on posteroventral surface of thighs; discoidal fold distinct; (2) tympanic annulus distinct and externally visible through skin; tympanic membrane present but hidden by skin; upper and posterior margins of tympanic annulus covered by low supratympanic fold; two distinct postrictal subconical tubercles on each side of the head; (3) snout moderate in length with a small rostral papilla at tip, subacuminate to acuminate in dorsal view, rounded in lateral view; canthus rostralis very distinct, rounded in lateral view and concave or straight in dorsal view (4) upper eyelid with several subconical tubercles; (5) cranial crests absent; (6) vocal slits and nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits expanded, rounded to truncate; (8) narrow lateral fringes on fingers; (9) low, rounded and subconical ulnar tubercles; (10) heel bearing one distinct subconical tubercle and a few nearby lower pustules; inner tarsal fold distinct; low and subconical outer tarsal tubercles; (12) narrow lateral fringes on toes; basal toe webbing, most distinctive between Toe IV and Toe V; Toe V slightly longer than Toe III; toe discs nearly as large as those on fingers; (13) coloration in life highly variable, dorsum generally pale brown or yellowish; scapular and supratympanic folds, and scapular tubercles darker than background; dark labial bars on both sides of head; flanks and dorsal surfaces of limbs usually with the same colour of the dorsum, showing several darker diagonal stripes; slightly translucent venter with dark flecks distributed on throat, chest, belly and ventral surfaces of limbs, but usually more numerous on throat, chest and ventral surfaces of hindlimbs; in males, groins and venter with same colour; in females groins are salmon reddish; bright golden iris; and (14) SVL in adult females from 18.92 to 23.76 mm (mean = 21.46; N = 12), in adult males SVL from 14.11 to 16.42 mm (mean = 15.19; N = 8; Tables 2 and 3 ). Figure 16. Photographs of the holotype of Pristimantis ujucami (QCAZ 49030; adult female, SVL 21.16 mm) in preservative. A, dorsal view; B, ventral view; C, ventral view of the left hand; D, ventral view of the right foot. Comparisons with other species: In this section, coloration refers to live individuals unless otherwise noticed. Among the species of the P. trachyblepharis species group, P. ujucami is most similar to P. aquilonaris by having a moderately long snout, usually subacuminate in dorsal view and rounded in profile. However, P. aquilonaris has an angular canthus rostralis in lateral view, which is rounded and more distinct in P. ujucami ( Fig. 17 ). They are also similar because both show red groins and a golden iris. Nevertheless, in P. aquilonaris , the reddish colours are usually arranged in rounded blotches, which are absent in P. ujucami . Additionally, the iris is usually more brilliant in P. ujucami . Pristimantis ujucami is similar to a few species outside the P. trachyblepharis group, such as P. bicantus ( Guayasamin & Funk, 2009 ) , P. prolatus ( Lynch & Duellman, 1980 ) and P. nelsongalloi ( Valencia et al. , 2019 ), as they also have a distinct canthus rostralis and some individuals have similar snouts in profile. In addition, Pristimantis ujucami is like P. bicantus and P.nelsongalloi in having red groins, and like P.prolatus in having a golden iris. However, P. bicantus has a rounded snout in dorsal view (subacuminate in P. ujucami ), have vocal slits and lacks ulnar, outer tarsal and heel tubercles (unlike P. ujucami ). Pristimantis prolatus has vocal slits too and does not show red groins and hindlimbs. Finally, P. nelsongalloi differs from P. ujucami by having clearly distinguishable dorsolateral folds and by lacking inner tarsal fold. Figure 17. Colour variation in life individuals of Pristimantis ujucami in dorsolateral view. A, QCAZ 49031 (subadult female, SVL 16.19 mm); B, QCAZ 57002 (adult male, SVL 14.39 mm); C, QCAZ 57010 (subadult female, 18.95 mm); D, QCAZ 57003 (adult male, SVL 15.47 mm); E, QCAZ 57004 (subadult female, SVL 18.91 mm); F, QCAZ 57646 (adult male, SVL 14.2 mm); G, QCAZ 60212 (adult female, SVL 22.33 mm); H, QCAZ 58865 (adult female, SVL 20.62 mm); I, QCAZ 60214 (adult female, SVL 23.76 mm); J, QCAZ 60194 (adult female, SVL 22.43 mm); K, QCAZ 60196 (adult female, SVL 20.45 mm); L, QCAZ 58874 (subadult female, SVL 16.74 mm). Description of the holotype : Adult female (QCAZ 49030). Measurements (in mm): SVL 21.10; tibia length 11.37; foot length 9.54; head length 6.55; head width 7.74; interorbital distance 2.13; width of upper eyelid 2.13; internarial distance 1.93; eye–nostril distance 2.17; eye diameter 2.72; tympanum diameter 1.05. Colour of the holotype in life and preservative is shown in Figures 15 and 16 , respectively. Head ( Figs 15 , 16A, B ): Wider than long, less wide than body; snout with a small rostral papilla at the tip, subacuminate in dorsal view, rounded in profile; canthus rostralis distinct, concave in dorsal view, angular in lateral view; loreal region concave; tympanic annulus distinct and externally visible through skin; tympanic membrane present but hidden by skin; upper margin of tympanic annulus covered by low supratympanic fold; two subconical postrictal tubercles on each side of head; four subconical tubercles and several pustules and spicules on upper eyelid; skin on throat weakly areolate; dentigerous processes of vomer rounded, slightly oblique, each vomer bearing four or five teeth, widely separated and posterior to choanae; choanae not concealed by palatal shelf of maxilla; tongue longer than wide, not notched posteriorly, posterior two thirds not adherent to the mouth floor. Figure 18. Colour variation in life individuals of Pristimantis ujucami in ventral view. A, QCAZ 49031 (subadult female, SVL 16.19 mm); B, QCAZ 57002 (adult male, SVL 14.39 mm); C, QCAZ 57010 (subadult female, 18.95 mm); D, QCAZ 57003 (adult male, SVL 15.47 mm); E, QCAZ 57004 (subadult female, SVL 18.91 mm); F, QCAZ 57646 (adult male, SVL 14.2 mm); G, QCAZ 60212 (adult female, SVL 22.33 mm); H, QCAZ 58865 (adult female, SVL 20.62 mm); I, QCAZ 60214 (adult female, SVL 23.76 mm); J, QCAZ 60194 (adult female, SVL 22.43 mm); K, QCAZ 60196 (adult female, SVL 20.45 mm); L, QCAZ 58874 (subadult female, SVL 16.74 mm). Figure 19. Colour variation in preserved individuals of Pristimantis ujucami : 1, dorsal views; 2, ventral views. A, QCAZ 49026 (adult female, SVL 22.62 mm); B, QCAZ 60194 (adult female, SVL 22.43 mm); C, QCAZ 60212 (adult female, SVL 22.33 mm); D, QCAZ 60214 (adult female, SVL 23.76 mm); E, QCAZ 58874 (subadult female, SVL 16.74 mm); F, QCAZ 60206 (subadult female, SVL 16.61 mm); G, QCAZ 57010 (subadult female, SVL 18.95 mm); H, QCAZ 57003 (adult male, SVL 15.47 mm); I, QCAZ 60621 (adult male, SVL 16.42 mm); J, QCAZ 57002 (adult male, 14.39 mm). Dorsum and venter ( Figs 15 , 16A, B ): Dorsal skin shagreen bedecked with some pustules; scapular folds distinct, \ /-shaped, with a pair of subconical scapular tubercles towards its posterior end, and another pair outside the fold and posterolateral to it; small pustules grouped in rows to form very low, almost negligible paravertebral and dorsolateral folds on dorsum, and lateral folds on flanks; one pair of subconical sacral tubercles; several subconical tubercles and pustules on dorsal surface of cloacal sheath; skin on chest weakly areolate, belly areolate; discoidal fold distinct. Forelimbs ( Fig. 16C ): Low ulnar tubercles; Finger I shorter than Finger II; discs expanded, rounded; all fingers having ventral pads surrounded by circumferential grooves; palmar tubercles well defined, outer palmar tubercle bifid, approximately 1.5 times the size of the ovoid thenar tubercle; subarticular tubercles prominent; hyperdistal subarticular tubercles low; distinct supernumerary tubercles at base of fingers; fingers bearing narrow lateral fringes; basal webbing slightly noticeable. Hindlimbs ( Fig. 16D ): Heel bearing several pustules and one low subconical tubercle; three subconical outer tarsal tubercles; inner tarsal fold distinct; Toe V slightly longer than Toe III; discs expanded, rounded; all toes having ventral pads surrounded by circumferential grooves; inner metatarsal tubercle enlarged, approximately five times the size of the rounded outer metatarsal tubercle; subarticular tubercles prominent, well defined; hyperdistal subarticular tubercles low; distinct supernumerary tubercles at base of toes; toes bearing broad lateral fringes; basal webbing, most distinctive between Toes IV and V; discs nearly as large as those on fingers; skin on chest and anteroventral surfaces of thighs weakly areolate, posteroventral surface of thighs coarsely areolate. Variation: The 14 individuals of P. ujucami with available genetic material show a mean intraspecific uncorrected genetic p- distance of 1.6% in the 16S gene sequence. The highest distances are those between populations in Morona Santiago vs. Zamora Chinchipe Provinces. In the type series, adult males (SVL = 14.11– 16.42 mm ) are between 13.22 and 40.52% smaller than adult females (SVL = 18.92–23.76 mm ). Below we list character states distinct from those described in the holotype , followed by an example. Colour variation in life and in preservative is shown in Figures 17 to 19. Head ( Figs 17–19 ): Shagreen and highly spiculate (QCAZ 58865); smooth with scattered pustules (QCAZ 60196). Head wider than body (QCAZ 60621). Snout acuminate in dorsal view (QCAZ 60212). Canthus rostralis straight in dorsal view (QCAZ 60194); rounded in profile (QCAZ 60612). Postrictal tubercles on each side of head: two conical (QCAZ 60196); one conical and two subconical (QCAZ 60195); one conical and one subconical (QCAZ 60212). Tubercles on upper eyelid: two subconical (QCAZ 58874); two conical and several subconical (QCAZ 49031). Interorbital tubercle: one conical (QCAZ 58866); two subconical (QCAZ 60207). Low interocular fold (QCAZ 60195). Subconical occipital tubercle (QCAZ 56998). Dentigerous processes of vomer narrowly separated (QCAZ 60194). Throat smooth (QCAZ 60196). Dorsum and venter ( Figs 17–19 ): Individuals of both Morona Santiago and Zamora Chinchipe Provinces show a similar morphology despite being at a distance of 220 km . A frequent polymorphism (16% of the paratypes , N = 5) that appears only on individuals of Morona Santiago Province is the presence of a pale broad stripe that starts at the back of the head and narrows posteriorly down to the cloacal region ( QCAZ 57010 ). Two males ( QCAZ 56998 , QCAZ 57001 ), out of six adult males with photographs in life, show a small diffuse reddish blotch on groins. Other males do not show reddish colours on any part of their body. Among females with photographs in life ( 11 adult females; nine subadult females), all of them show reddish colours on groins. Therefore , there seems to be a certain sexual dimorphism in colour. Dorsum smooth with scattered pustules ( QCAZ 60196 ); shagreen, highly spiculated with scattered conical and subconical tubercles ( QCAZ 60207 ). Scapular folds:) (-shaped ( QCAZ 60194 );> <-shaped ( QCAZ 49031 ), low and diffuse ( QCAZ 57646 ); several subconical scapular tubercles along the scapular fold ( QCAZ 57010 ). Dorsolateral folds formed by rows of tubercles and pustules ( QCAZ 58874 ). Distinct paravertebral folds ( QCAZ 58868 ). Chest and belly smooth (60196). Forelimbs ( Fig. 19 ): In females, reddish colours on shoulders (QCAZ 58865). Fingers without lateral fringes (QCAZ 60205). Basal webbing absent (QCAZ 49026). Posteroventral surface of thighs areolate (QCAZ 60196). Discs on fingers truncate (QCAZ 57010). Hindlimbs ( Fig. 19 ): In females, reddish colours may appear on inner surface of shanks, anterior, posterior and dorsal surfaces of thighs, and inner surfaces of tarsus (QCAZ 58865). One conical and one subconical tubercle on dorsal surface of shanks (QCAZ 58865). Low outer tarsal subconical tubercles (QCAZ 58870). Toes bearing narrow lateral fringes (QCAZ 60205). High inner metatarsal tubercle (QCAZ 60205). Discs on toes truncate (QCAZ 60194). Distribution, natural history and conservation status: Pristimantis ujucami is known from four localities in Morona Santiago and Zamora Chinchipe Provinces, in Ecuador ( Fig. 3 ). In Morona Santiago , it was found at the Abanico Protected Forest near the Hidroabanico Hydroelectric Power Plant, and near the Volcano River in Sangay National Park. In Zamora Chinchipe , it was reported near the Bombuscaro River, in Podocarpus National Park, and at the Numbami Reserve. Elevation range is 981 to 1720 m a.s.l. Frogs were found perching from 10 to 180 cm above the ground level, mainly on leaves and ferns by both day and night. Their natural region is Eastern Montane Forest and Eastern Foothills. Occurrences were near the forest edge in areas covered by native woodland, shrubby vegetation or even pasturelands ( Fig. 4 ). Proximity to pastureland suggests a certain tolerance to anthropogenic habitat disturbances. Extent of Occurrence of 1489.5 km 2 and Area of Occupancy of 28 km 2 . Zamora Chinchipe populations were found within Podocarpus National Park, Bombuscaro sector, and a private protected area, Reserva Numbami.Airline distance between populations from Zamora Chinchipe Province and those of Morona Santiago is 220 km . Therefore, because intermediate areas are largely unexplored, we assign P. ujucami to the Data Deficient Red List Category ( IUCN 2021 ). Etymology: The specific epithet ujucami is a noun in the genitive case and is a patronym for Martín Ujukam, known as the last Shuar warrior of Zamora, who led the struggles to prevent the Shuar people, a native American group, from being displaced from its territory. He lived at the confluence of the Zamora and Jamboé rivers. The Jamboé valley is inhabited by this species.