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Through the combination of morphological and phylogenetic evidence, we describe two species of Pristimantis from the upper basin of the Napo River. Both species have well-defined dorsolateral folds, a conical tubercle on the eyelid, a papilla on the tip of the snout, weakly expanded discs, and small size (female SVL < 28.2 mm). Pristimantis omarrhynchus sp. nov. differs from its sister species, Pristimantis miltongallardoi sp. nov., by the absence of iridophores on the belly, subacuminate snout in dorsal view, and narrow digits. Our phylogeny and morphological evidence, are conclusive in assigning them to the Pristimantis boulengeri species group, closely related to P. boulengeri, P. cryptopictus, P. dorspictus, and P. brevifrons. The new species are the first reported for the P. boulengeri group in Ecuador and the Amazon basin. We also comment on the correct identity of GenBank sequences previously assigned to P. thymelensis and P. myersi., A través de la combinación de evidencia morfológica y filogenética describimos dos especies de Pristimantis de la cuenca alta del río Napo. Las dos especies presentan pliegues dorsolaterales bien definidos, un tubérculo cónico en el párpado, una papila en la punta del hocico, discos poco dilatados y tamaño pequeño (hembras LRC < 28,2 mm). Pristimantis omarrhynchus sp. nov. se diferencia de su especie hermana, Pristimantis miltongallardoi sp. nov., por la ausencia de iridóforos en el vientre, hocico subacuminado en vista dorsal y dígitos estrechos. Nuestra filogenia, en combinación con la evidencia morfológica, son concluyentes para asignarlas al grupo de especies Pristimantis boulengeri y cercanamente relacionadas a P. boulengeri, P. cryptopictus, P. dorspictus y P. brevifrons. Las nuevas especies son las primeras reportadas para el grupo P. boulengeri en el Ecuador y la cuenca amazónica. También comentamos la identidad correcta de secuencias GenBank previamente asignadas a P. thymelensis y P. myersi., Asociación Herpetológica Argentina

Pinheiro, Paulo D P, Blotto, Boris L, Ron, Santiago R, Stanley, Edward L, Garcia, Paulo C A, Haddad, Célio F B, Grant, Taran, Faivovich, Julián (2022): Prepollex diversity and evolution in Cophomantini (Anura: Hylidae: Hylinae). Zoological Journal of the Linnean Society 195 (3): 995-1021, DOI: 10.1093/zoolinnean/zlab079, URL: https://academic.oup.com/zoolinnean/article/195/3/995/6384856

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. 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,> 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). 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. 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. 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);> 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.

PRISTIMANTIS VENTRISTELLATUS SP. NOV. (FIGS 20–24; 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: AB88F2AD-924D-4739-B841-561C 09292737 Holotype (Figs 20, 21): QCAZ 69240 (field no. SC-PUCE 59773), adult male from Ecuador, Provincia Morona Santiago, Cantón Santiago de Méndez, Parroquia San Francisco de Chinimbimi, Puchimi (2.7843º S, 78.1426º W), 1905 m a.s.l., collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia and Jefferson Mora on 10 September 2017. Paratypes (26: 12 males, 13 females, one juvenile): Ecuador: Provincia Morona Santiago: Puchimi, QCAZ 69263, adult female (2.7845º S, 78.1400º W), 1930 m a.s.l.; QCAZ 69318, adult female (2.7880º S, 78.1299º W), 2283 m a.s.l.; QCAZ 69209, adult female (2.7844º S, 78.1409º W), 1915 m a.s.l.; QCAZ 69211, adult male (2.7845º S, 78.1400º W), 1930 m a.s.l.; QCAZ 69243, adult male (2.7844º S, 78.1419º W), 1908 m a.s.l.; QCAZ 69245, adult male (2.7843º S, 78.1412º W), 1926 m a.s.l.; QCAZ 69264, adult female (2.7857º S, 78.1388º W), 1979 m a.s.l.; QCAZ 69265, adult male (2.7857º S, 78.1376º W), 1919 m a.s.l.; QCAZ 69267, QCAZ 69273, adult males (2.7858º S, 78.1372º W), 2032 m a.s.l.; QCAZ 69291, adult male (2.7848º S, 78.1391º W), 1969 m a.s.l.; QCAZ 69296, adult female (2.7843º S, 78.1403º W), 1938 m a.s.l.; QCAZ 69319, adult female, and QCAZ 69320, adult male (2.7880º S, 78.1299º W), 2283 m a.s.l.; QCAZ 69338, adult female (2.7866º S, 78.1336º W), 2125 m a.s.l.; QCAZ 69348, juvenile, and QCAZ 69349, adult male (2.7870º S, 78.1320º W), 2180 m a.s.l.; QCAZ 69351, adult male (2.7871º S, 78.1319º W), 2193 m a.s.l.; QCAZ 69357–69359, adult females, and QCAZ 69362, adult male (2.7873º S, 78.1317º W), 2203 m a.s.l.; QCAZ 69365, adult female (2.7874º S, 78.1314º W), 2211 m a.s.l.; QCAZ 69368, adult male (2.7875º S, 78.1312º W), 2218 m a.s.l.; QCAZ 69395, adult female (2.7858º S, 78.1340º W), 2043 m a.s.l.; QCAZ 69397, adult female (2.7860º S, 78.1338º W), 2029 m a.s.l. Collected by D. Almeida, D. Núñez, E. Nusirquia and J. Mora on 9–13 September 2017. Suggested common name: English: stellated venter rain frog. Spanish: cutín de vientre estrellado. Diagnosis (Figs 20–24): A species of Pristimantis characterized by the following combination of characters: (1) dorsal skin shagreen with scattered tuberclesandpustules; scapularregionwithW-shaped scapular folds and two pairs of anterolateral and posteromedial conical or subconical tubercles along them; low interocular fold, usually with one or two low subconical interocular tubercles; dorsolateral folds usually absent; ventral skin weakly areolate on throat and chest, and coarsely areolate on belly and posteroventral surface of thighs; discoidal fold distinct; (2) tympanic annulus present, its upper margin covered by low supratympanic fold; tympanic membrane present but hidden by skin; one or two postrictal conical or subconical tubercles; (3) snout short to moderate in length with a rostral papilla at tip, subacuminate in dorsal view and rounded to protruding in profile; (4) upper eyelid with one distinct conical tubercle on the posterolateral quadrant, and several subconical tubercles spread over all its surface; (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) fingers without lateral fringes; (9) low and subconical ulnar tubercles; (10) heel bearing one to few low subconical tubercles; inner tarsal fold illdefined; low and subconical outer tarsal tubercles; (12) toes with narrow lateral fringes; basal toe webbing, most distinctive between Toe IV and Toe V; Toe V slightly longer than Toe III; discs expanded and rounded, nearly as large as those on fingers; (13) in life, dorsum brown to dark brown with a darker W-shaped scapular mark posterior to the scapular fold; dark brown interorbital bar; sides of head brown bearing darker labial bars; flanks showing the same colour of the dorsum, with or without several diffuse dark brown diagonal stripes; venter dark brown to brown with white spots, which may reach the lower halves of flanks and head; anterior surface of thighs and inner surface of shanks usually bearing some inconspicuous pale red blotches; brown groins with or without inconspicuous red blotches; iris bronze to reddish copper; and (14) SVL range in adult females from 18.62 to 22.97 mm (mean = 20.67; N = 13), in adult males 14.51 to 16.69 mm (mean = 15.87, N = 13; Tables 2 and 3). Comparisons with other species: In this section, coloration refers to live individuals unless otherwise mentioned. Pristimantis ventristellatus can be confounded with P. pramukae by having a dark brown dorsal coloration and a dark venter with scattered clear spots and pale flecks (Figs 13, 23). However, dorsal surfaces of P. ventristellatus are usually more tuberculate, the clear ventral spots are more numerous and its venter is darker. They also differ in the snout shape, which is subacuminate in dorsal view and protruding in profile in P. ventristellatus (Fig. 20A, B), and usually rounded in both views in P. pramukae (Fig. 10A, B). The shape of the snout does not allow to clearly distinguish P. ventristellatus from P. trachyblepharis. However, they differ in dorsal and ventral coloration, lighter and generally creamy-yellow in P. trachyblepharis, whose venter is also slightly translucent (unlike in P. ventristellatus). Finally, the most tuberculate individuals of P. ventristellatus could be mistakenly identified as P. albujai, since both species are small, dark brown dorsally and ventrally, have a subacuminate snout in dorsal view and have prominent scapular folds. They can be easily differentiated by looking at their groins and hidden surfaces of the hindlimbs, which are usually red in P. albujai and brown with or without inconspicuous red blotches in P. ventristellatus (Fig. 23). Pristimantis ventristellatus also differs from P. albujai in lacking distinct dorsolateral folds, which are prominent in P. albujai. Description of the holotype: Adult male (QCAZ 69240). Measurements (in mm): SVL 15.67; tibia length 9.02; foot length 7.75; head length 6.03; head width 5.41; interorbital distance 2.03; width of upper eyelid 2.06; internarial distance 0.9; eye–nostril distance 1.78; eye diameter 2.53; tympanum diameter 0.97. Colour of the holotype in life and preservative is shown in Figures 20 and 21, respectively. Head (Figs 20, 21A, B): Longer than wide, wide as body; snout moderate in length with a rostral papilla at the tip, subacuminate in dorsal view, protruding in profile; canthus rostralis distinct, concave in dorsal view, slightly rounded in profile; loreal region concave; tympanic annulus distinct, its upper margin covered by low supratympanic fold; tympanic membrane present but hidden by skin; two postrictal tubercles, one conical and one smaller and subconical; low interocular fold with a subconical interocular tubercle; upper eyelid with several subconical tubercles and three conical tubercles, of which the largest is posterolateral; many pustules on the occipital region surround a pair of subconical tubercles; skin on throat weakly areolate; dentigerous processes of vomer indistinct; choanae concealed by palatal shelf of maxilla; tongue as wide as long, posteriorly notched, posterior half not adherent to the mouth floor. Dorsum and venter (Figs 20, 21A, B): Dorsal skin shagreen bearing numerous pustules, spicules and tubercles; W-shaped scapular folds; two pairs of conical scapular tubercles, anterolateral and posteromedial along the scapular fold; several subconical cloacal tubercles; low lateral folds formed by rows of pustules on flanks; skin on chest weakly areolate, belly coarsely areolate; discoidal fold ill-defined. Forelimbs (Fig. 21C): Low and subconical ulnar tubercles; Finger I shorter than Finger II; discs narrowly expanded, rounded; all fingers having ventral pads surrounded by circumferential grooves; palmar tubercles not well defined, barely visible; fingers lacking lateral fringes; basal webbing smaller than that on toes, slightly noticeable between Fingers III and IV. Hindlimbs (Fig. 21D): Heel bearing several pustules and one low subconical tubercle; distinct subconical tubercles on outer surface of tarsus; inner tarsal fold diffuse; Toe V slightly longer than Toe III; discs expanded, rounded, nearly as large as those on fingers; all toes having ventral pads surrounded by circumferential grooves; inner metatarsal tubercle enlarged, approximately three times the size of the outer metatarsal tubercle; hyperdistal subarticular tubercle of Toe IV distinct; narrow lateral fringes; basal webbing more noticeable between Toe IV and V; skin on posteroventral surfaces of thighs coarsely areolate. Variation: Four individuals have available DNA sequences; mean intraspecific uncorrected genetic p- distance is 0.6% for the 16S gene. In the type series, adult males (SVL range 14.51–16.69 mm) are between 10.37 and 36.84% smaller than adult females (SVL range 18.62–22.97 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 22 to 24. Head (Figs 22–24): Snout short (QCAZ 69395). Snout rounded in profile (QCAZ 69264). Canthus rostralis angular in profile (QCAZ 69359). Three postrictal tubercles, one conical and two subconical (QCAZ 69263); one conical tubercle (QCAZ 69243); two subconical tubercles (QCAZ 69245). Interocular region without interocular fold or interocular tubercles (QCAZ 69368); two subconical interocular tubercles (QCAZ 69319). Upper eyelid with one elongate tubercle, two conical tubercles and several subconical tubercles (QCAZ 69319); several subconical tubercles (QCAZ 69211); one conical tubercle and several subconical tubercles (QCAZ 69243). Dentigerous processes of vomer posterior to the choana, oblique and widely separated, each vomer bearing two or three teeth (QCAZ 69357). Dorsum and venter (Figs 22–24): Dorsal skin finely shagreen with scattered pustules and tubercles specially in the posterior half (QCAZ 69357); shagreen, highly tuberculate (QCAZ 69267). Low scapular folds (QCAZ 69362). Two pairs of subconical scapular tubercles (QCAZ 69358); conical scapular tubercles (QCAZ 69338). Inconspicuous dorsolateral folds (QCAZ 69245). Two lateral folds on each flank (QCAZ 69245). Forelimbs (Fig. 24): Supernumerary tubercles at base of fingers (QCAZ 69318). Subarticular tubercles distinct, well defined (QCAZ 69318). Low hyperdistal tubercles (QCAZ 69318). Outer palmar tubercle lower than thenar tubercle (QCAZ 69357). Outer palmar tubercle bifid, approximately 1.5 times the size of the thenar tubercle (QCAZ 69209). Narrow lateral fringes (QCAZ 69623). Basal webbing distinct (QCAZ 69209). Discs truncate (QCAZ 69368). Hindlimbs (Fig. 24): Two conical tubercles and a few subconical tubercles on outer surface of tarsus (QCAZ 69267). Inner tarsal fold distinct (QCAZ 69273) or absent (QCAZ 69351). Low hyperdistal tubercles (QCAZ 69318). Outer metatarsal tubercle lower than inner metatarsal tubercle, approximately twice its size (QCAZ 69349). Discs truncate (QCAZ 69264). Distribution, natural history and conservation status: This species is only known from the type locality, Puchimi, in the Cutucú Mountain Range, in Morona Santiago Province, Ecuador, between 1908 and 2283 m a.s.l. (Fig. 3). The natural region is Eastern Montane Forest. Individuals were found from 10 to 100 cm above the ground level on leaves in forest (Fig. 4) characterized by low shrubs and abundant mosses and terrestrial bromeliads. All observations were made at night. Individuals were collected down to 1 km from the nearest disturbed area (pastureland), suggesting that P. ventristellatus can persist at least under moderate anthropogenic habitat disturbances. The type locality is within a protected area, the Kutukú-Shaimi Protection Forest. According to the available data, this species has a restricted distribution (Extent of Occurrence 0.122 km 2, Area of Occupancy 12 km 2). However, adjacent areas remain unexplored, so we assign P. ventristellatus to the Data Deficient Red List Category (IUCN 2021). Etymology: The specific name ventristellatus refers to the dark colour with white spots on the venter of this species, which resembles a starred (stellate) night sky.

PRISTIMANTIS PRAMUKAE SP. NOV. (FIGS 10–14; 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: 82CD48C8-6AB1-4BEA-91E2-AD3E5AD6CDF3 Holotype (Figs 10, 11 ): QCAZ 68549 (field no. SC-PUCE 59107), adult female from Ecuador, Provincia Morona Santiago, Cantón Gualaquiza, Parroquia Bomboiza, on the buffer zone of El Quimi Biological Reserve, tepui plateau on the eastern side of the Quimi River (3.5184º S, 78.3919º W), 1991 m a.s.l., collected by Darwin Núñez, Eloy Nusirquia, Alex Achig, and Ricardo Gavilanes on 4 July 2017. Paratypes (34: 19 males, 13 females, two juveniles): Ecuador: Provincia Morona Santiago: Buffer zone of El Quimi Biological Reserve, tepui plateau on the eastern side of the Quimi River, QCAZ 68547 and QCAZ 68580 adult males (3.5184º S, 78.3919º W), 1991 m a.s.l.; QCAZ 68559, subadult female (3.5180º S, 78.3980º W), 2055 m a.s.l.; QCAZ 68572, subadult male (3.5191º S, 78.3893º W), 2003 m a.s.l.; QCAZ 68574, subadult male (3.5196º S, 78.3844º W), 2074 m a.s.l.; QCAZ 68579, subadult female (3.5197º S, 78.3849º W), 2074 m a.s.l.; QCAZ 68591, subadult male (3.5188º S, 78.3802º W), 2132 m a.s.l.; QCAZ 68595, adult female (3.5189º S, 78.3839º W), 2035 m a.s.l.; QCAZ 68610, adult male (3.5181º S, 78.3975º W), 2032 m a.s.l.; QCAZ 68613, adult male (3.5185º S, 78.3921º W), 1980 m a.s.l.; QCAZ 72588 and QCAZ 72592, juveniles (3.5182º S, 78.3913º W), 1994 m a.s.l.; QCAZ 72589, QCAZ 72593, QCAZ 72605 and QCAZ 72610–72612, adult males (3.5182º S, 78.3913º W), 1994 m a.s.l.; QCAZ 72590–72591, QCAZ 72606–72608 and QCAZ 72620, adult females (3.5182º S, 78.3913º W), 1994 m a.s.l.; QCAZ 72617, subadult male (3.5182º S, 78.3913º W), 1994 m a.s.l.; QCAZ 72600–72601, subadult females, and QCAZ 72602, adult male (3.5182º S, 68.3969º W), 2010 m a.s.l.; QCAZ 72604, adult female (3.5187º S, 78.3729º W), 2165 m a.s.l.; QCAZ 72613, subadult male (3.5198º S, 78.3902º W), 2059 m a.s.l.; QCAZ 72614, subadult male, and QCAZ 72615, adult female (3.5197º S, 78.3904º W), 1995 m a.s.l.; QCAZ 72618, adult male (3.5157º S, 78.4106º W), 2017 m a.s.l.; QCAZ 72619, adult male (3.5193º S, 78.3869º W), 2028 m a.s.l. Collected by D. Almeida, D. Núñez, E. Nusirquia, A. Achig, R. Gavilanes and M. Moretta on 12–20 April 2018 and 4–10 July 2017. Suggested common name: English: Bomboiza rain frog. Spanish: cutín de Bomboiza. Diagnosis (Figs 10–14): A species of Pristimantis characterized by the following combination of characters: (1) dorsal skin smooth to shagreen; scapular region usually with two scapular subconical tubercles and a low \ /-shaped scapular fold; dorsolateral folds absent; ventral skin weakly areolate on throat and chest, coarsely areolate on belly and posteroventral surface of thighs; discoidal fold distinct; (2) tympanic annulus distinct and externally visible through skin, its upper margin covered by low supratympanic fold; tympanic membrane present but hidden by skin; one or two postrictal conical tubercles; (3) snout short to moderate in length, rounded to subacuminate in dorsal view and rounded to protruding in profile; (4) upper eyelid with one distinct subconical tubercle on its posterolateral quadrant, and several pustules spread over all their surface; (5) cranial crests absent; (6) vocal slits and nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits expanded, rounded; (8) fingers without lateral fringes; (9) low and subconical ulnar tubercles; (10) heel bearing one to few low subconical tubercles; inner tarsal fold diffuse; 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 as large as those on fingers; (13) in life, dorsum brown to dark brown usually with a W-shaped paler mark in the scapular region; head bearing a pale cream interorbital bar and sides of head brown with darker labial bars; flanks with the same colour of the dorsum, with or without several diffuse dark brown diagonal stripes, which also extend along dorsal surfaces of limbs; surfaces adjoining the outline of these diagonal stripes are generally paler and show white spots or flecks; venter brown with pale flecks and a few white spots, which may reach lower halves of flanks and head; a pale midventral stripe extends from the lower lip to the belly in those individuals having middorsal stripe; inconspicuous small red blotches may appear on upper groin and dorsal surface of thighs; groins pale brown with or without inconspicuous red blotches; iris copper; and (14) SVL in adult females from 18.27 to 21.03 mm (mean = 19.43 mm; N = 9), in adult males from 14.09 to 16.80 mm (mean = 15.09 mm; N = 13; Tables 2 and 3). Comparisons with other species: In this section, coloration refers to live individuals unless otherwise mentioned. Pristimantis pramukae is most similar to some congeneric species of the P. trachyblepharis species group, especially to P. nanus in its short snout and the W- shaped pale marks in the scapular region. Pristimantis ventristellatus can also be confounded with P. pramukae because both have similar dorsal colorations and a dark venter with scattered clear spots and pale flecks (Figs 13, 23). However, P. nanus (SVL = 11.04–12.46 mm in adult males; 14.08– 14.82 mm in adult females) is smaller than P. pramukae (SVL = 14.09–16.80 mm in adult males; 18.27–21.03 mm in adult females; Tables 2 and 3) and has less expanded discs on fingers and toes. On P. ventristellatus, the clear ventral spots are more numerous than in P. pramukae, and the venter is noticeably darker. In addition, dorsal surfaces of P. pramukae are usually less tuberculate than that on P. ventristellatus. Outside the P. trachyblepharis group, P. pramukae is similar to P. exoristus (Duellman & Pramuk, 1999) by having a small size and a W- shaped mark in the scapular region. However, in P. exoristus, that mark is dark, and in P. pramukae it is paler than the background (Fig. 10A). Additionally, in P. exoristus the snout is moderately long (in P. pramukae it is short) and Toe V is much longer than Toe III (Toe V slightly longer than III in P. pramukae). Description of the holotype: Adult female (QCAZ 68549). Measurements (in mm): SVL 19.83; tibia length 10.22; foot length 8.69; head length 7.34; head width 6.41; interorbital distance 2.03; width of upper eyelid 1.77; internarial distance 1.78; eye–nostril distance 2.37; eye diameter 2.57; tympanum diameter 0.75. Colour of the holotype in life and in preservative is shown in Figures 10 and 11, respectively. Head (Figs 10, 11A, B): Longer than wide, wide as body; snout short, rounded in dorsal and lateral views; canthus rostralis distinct, concave in dorsal view, rounded in profile; loreal region concave; tympanic annulus distinct, its upper margin covered by low supratympanic fold; tympanic membrane present but hidden by skin; one conical postrictal tubercle; posterolateral quadrant of eyelid bearing a single subconical tubercle; skin on throat areolate; dentigerous processes of vomer oblique, posterior to choanae, broadly separated from each other; choanae not concealed by palatal shelf of maxilla; tongue longer than wide, not notched posteriorly, posterior threefifths not adherent to the mouth floor. Dorsum and venter (Figs 10, 11A, B): Dorsal skin smooth; scapular region with low \ /-shaped scapular folds and a pair of posteromedial subconical tubercles; dorsolateral folds absent; many pustules on cloacal region; skin on chest weakly areolate, belly coarsely areolate; discoidal fold distinct. Forelimbs (Fig. 11C): Low and subconical 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 twice the size of the ovoid thenar tubercle; high subarticular tubercles; low hyperdistal subarticular tubercles; supernumerary tubercles at base of fingers; fingers lacking lateral fringes; basal webbing slightly noticeable. Hindlimbs (Fig. 11D): Heel bearing one low subconical tubercle; outer surface of tarsus with low subconical tubercles; inner tarsal fold absent; Toe V slightly longer than Toe III; discs expanded and rounded, except those on Toe III and Toe IV of the right foot, which are expanded and elongate acuminate; discs nearly as large as those on fingers; all toes having ventral pads surrounded by circumferential grooves; subarticular tubercles well defined, as high as those on hands; low hyperdistal subarticular tubercles; numerous indistinct supernumerary tubercles; enlarged inner metatarsal tubercle, approximately five times the size of the small, rounded and high outer metatarsal tubercle; narrow lateral fringes; basal webbing, most distinctive between Toe IV and Toe V; skin on posteroventral surfaces of thighs coarsely areolate. Variation: There is no appreciable mean uncorrected genetic p- distance between the three individuals of P. pramukae with available sequences for the 16S gene. In the type series, adult males (SVL = 14.09– 16.80 mm) are between 8.05 and 33.01% smaller than adult females (SVL = 18.27–21.03 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 12 to 14. Head (Figs 12–14): Snout moderate in length (QCAZ 72591). Snout subacuminate in dorsal view, protruding in profile (QCAZ 68572). Canthus rostralis distinct (QCAZ 68613), angular in profile (QCAZ 72607). One subconical postrictal tubercle (QCAZ 68580); two subconical tubercles (QCAZ 72614). Upper eyelid with two subconical tubercles (QCAZ 68579); one conical tubercle and three subconical tubercles (QCAZ 72590). Dentigerous processes of vomer barely noticeable (QCAZ 68595). Dorsum and venter (Figs 12–14): Cream middorsal stripe extending from snout to cloacal region in 40% of the paratypes (N = 14). Dorsal skin shagreen (QCAZ 72611); shagreen with scattered pustules (QCAZ 72591). Scapular folds diffuse (QCAZ 72615). Two pairs of scapular subconical tubercles along the scapular fold, anterolateral and posteromedial in the scapular region (QCAZ 72606); two posteromedial conical tubercles in the scapular region (QCAZ 68591). Cloacal subconical tubercles (QCAZ 72620). Two lateral folds on each flank (QCAZ 72605). Diffuse dorsolateral folds (QCAZ 68613). Forelimbs (Fig. 14): Antebrachial tubercle distinct (QCAZ 68613). Narrow lateral fringes (QCAZ 72610). Hindlimbs (Fig. 14): Heel tubercle and outer tarsal tubercles prominent (QCAZ 72592). Distribution, natural history and conservation status: This species is only known from a single locality in the buffer zone of El Quimi Biological Reserve, in Morona Santiago Province, Ecuador, between 1991 and 2132 m a.s.l. (Fig. 3). Its natural region is Eastern Montane Forest. Both localities are on a limestone tepui plateau on the eastern side of the Quimi Valley. The new species lives in forest (Fig. 4) composed of thin, scattered trees 10–15 m tall, shrubby vegetation up to 5 m sometimes mixed with a few palms and soils with cushioned consistency, covered by mosses, roots and terrestrial bromeliads. This type of soil is locally known as bamba and is characteristic of limestone tepui formations. The easternmost locality was at a distance of about 1 km from the nearest disturbed area (pastureland), suggesting that P. pramukae may have some level of resistance to human-generated environmental disturbances. By day, individuals were found at ground level. At night, they occur perching up to a height of 300 cm on leaves, branches, bromeliads and mosses. One amplectant couple was found on a branch 130 cm above the ground, near a stream, at 9: 36 p. m. on 16 April 2018. The height above the ground and the substrate on which the amplecting couple was found is similar to several individuals found at night, compared to those collected during the day, which may suggest that it is a species with nocturnal activity. The Extent of Occurrence of this species is 0.601 km 2, and the Area of Occupancy 12 km 2. However, there are still unexplored adjacent zones that could represent potential distribution areas for this species. Therefore, we assign P. pramukae to the Data Deficient Red List Category (IUCN 2021). The type locality is Etymology: The specific epithet pramukae is a noun in the genitive case and is a matronym dedicated to Jennifer B. Pramuk, who, in collaboration with William E. Duellman, made one of the most influential reviews of Peruvian Pristimantis, including the description of 18 new species. Several of those species inhabit the Cordillera del Cóndor, near the type locality of P. pramukae.

PRISTIMANTIS NANUS SP. NOV. (FIGS 5–9; 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: 594B77F6-9A58-49EC-8088-3B79E98891F2 Holotype (Figs 5, 6): QCAZ 72596 (field no. SC-PUCE 63140), adult female from Ecuador, Provincia Morona Santiago, Cantón Gualaquiza, Parroquia Bomboiza, in the buffer zone of El Quimi Biological Reserve, tepui plateau on the eastern side of the Quimi River (3.5182º S, 78.3913º W), 1994 m a.s.l., collected by Diego Almeida, Darwin Núñez, Eloy Nusirquia, Alex Achig and María del Mar Moretta on 23 April 2018. Paratypes (six: five males, one female): Ecuador: Provincia Morona Santiago: buffer zone of El Quimi Biological Reserve, tepui plateau on the eastern side of the Quimi River, QCAZ 68569, adult male (3.5187º S, 78.3920º W), 1978 m a.s.l.; QCAZ 68616, adult male (3.5189º S, 78.3689º W), 2209 m a.s.l.; QCAZ 68618, adult female (3.5188º S, 78.3697º W), 2202 m a.s.l.; QCAZ 72595, QCAZ 72598 and QCAZ 72616, adult males (3.5182º S, 78.3913º W), 1994 m a.s.l. Collected by D. Almeida, D. Núñez, E. Nusirquia, A. Achig, R. Gavilanes and M. Moretta on 5, 11 July 2017 and 23 April 2018. Suggested common name: English: dwarf rain frog. Spanish: cutín enano. Diagnosis (Figs 5–9): A species of Pristimantis characterized by the following combination of characters: (1) dorsal skin smooth with scattered pustules; low scapular folds, \ /-shaped, with two pairs of low scapular subconical tubercles along them, anterolateral and posteromedial; ventral skin weakly areolate on throat and chest, areolate on belly and on posteroventral surface of thighs; discoidal fold distinct; (2) tympanic annulus distinct; tympanic membrane present but hidden by skin and muscle; upper and posterior margins of tympanic annulus covered by low supratympanic fold; one subconical tubercle and several pustules on postrictal region; (3) snout short with a very small rostral papilla at tip, rounded in both dorsal and lateral views; canthus rostralis poorly defined, concave in dorsal view, rounded in lateral view; (4) one to few subconical tubercles and pustules on upper eyelid; (5) cranial crests absent; (6) vocal slits and nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits narrowly expanded, elongately acuminate; (8) fingers bearing narrow lateral fringes; (9) very low subconical ulnar tubercles; (10) one low subconical tubercle on heel; low and subconical outer tarsal tubercles; inner tarsal fold present; (12) toes bearing lateral fringes; basal toe webbing, most distinctive between Toe IV and Toe V; Toe V slightly longer than Toe III; toe discs slightly smaller as those on fingers; (13) in life, dorsal coloration highly variable, generally reddish brown with or without greenish areas; W- shaped paler mark in the scapular region; supratympanic folds darker than background; broad interorbital bar, pale cream anteriorly and dark posteriorly; sides of head usually with diffuse dark labial bars; flanks and dorsal surfaces of limbs with similar colour patterns and several darker diagonal stripes; surfaces adjoining the outline of these stripes are generally paler and have white spots or flecks; pale brown to dark brown venter with some white spots and dark flecks or mottling on throat, chest and belly; dark flecks and white spots of venter may reach the lower halves of flanks and head; throat darker than chest, belly and ventral surfaces of limbs; pale brown groins; dark red to reddish copper iris; and (14) SVL in adult females from 14.08 to 14.82 mm (mean = 14.45 mm; N = 2), adult males from 11.04 to 12.46 mm (mean = 11.67 mm; N = 5; Tables 2 and 3). Comparisons with other species: In this section, coloration refers to live individuals unless otherwise noticed. Pristimantis nanus can be differentiated from the other species of the P. trachyblepharis species group, except P. minimus and P. trachyblepharis, by its tiny size (Tables 2 and 3). Beyond that, P. nanus is similar to P. pramukae because both have W- shaped pale marks in the scapular region and a short snout. However, P. nanus has a chubbier appearance and less expanded, elongate-acuminate discs. Discs of P. pramukae may also be elongate-acuminate in some digits of several specimens, but this condition is not common. Moreover, P. pramukae usually have darker dorsal and ventral colours as compared with reddish, greenish or pale brown colours of P. nanus. Pristimantis nanus cannot be clearly distinguished from P. minimus (SVL range = 9.5–13.7 mm in adult males; 15.3– 18.9 in adult females) and P. trachyblepharis (SVL range = 12.37–15.47 mm in adult males; 15.53–22.24 in adult females) by size (Tables 2 and 3); however, P. minimus is much less tuberculate, lacks tympanic membrane and males have vocal slits, unlike P. nanus males. P. trachyblepharis has a longer snout, a slimmer body and usually shows lighter colorations. Pristimantis nanus can be confounded with other small congeneric species such as P. coronatus (SVL = 15.3 mm in a single female; Lehr & Duellman, 2007) and P. andinognomus (SVL = 10.0– 14.5 mm in adult males; 12.6–17.9 mm in adult females; Lehr & Coloma, 2008). Whereas P. nanus does not show distinctive colours on groins, P. coronatus show orange reddish colours. Pristimantis andinognomus has a dorsally acuminate snout as compared with the rounded snout of P. nanus and has white or yellow distinctive coloration in the groins (groins pale brown in P. nanus) Additionally, males of P. andinognomus have vocal slits, which are absent on P. nanus males. Pristimantis nanus is also similar to P. exoristus (Duellman & Pramuk, 1999) in having a small size (SVL = 15.0– 16.9 mm in adult males; 21.3–23.5 mm in adult females) and a W- shaped mark in the scapular region. However, in P. exoristus that mark is dark, and in P. nanus is paler than the background. Additionally, in P. exoristus the snout is moderately long, while in P. nanus it is short. Description of the holotype: Adult female (QCAZ 72596). Measurements (in mm): SVL 14.82; tibia length 7.06; foot length 6.16; head length 5.21; head width 4.41; interorbital distance 1.65; width of upper eyelid 1.24; internarial distance 1.10; eye–nostril distance 1.36; eye diameter 1.78; tympanum diameter 0.65. Colour of the holotype in life and in preservative is shown in Figures 5 and 6, respectively. Head (Figs 5, 6A, B): Longer than wide, wide as body; snout short with a very small rostral papilla at tip, rounded or slightly subacuminate in dorsal view, and rounded in profile; canthus rostralis poorly defined, concave in dorsal view, rounded in profile; loreal region concave; several pustules on dorsal surface of snout, and one low subconical tubercle anteromedial to the interocular region; tympanic annulus distinct, its upper margin covered by low supratympanic fold; tympanic membrane present but hidden by skin and muscle; one subconical postrictal tubercle and some pustules on each side of head; upper eyelid with three low subconical tubercles; skin on throat weakly areolate; dentigerous processes of vomer slightly posterior to choanae, triangular, widely separated, each vomer bearing four or five teeth; choanae not concealed by palatal shelf of maxilla; tongue slightly longer than wide, posteriorly notched, posterior half not adherent to the mouth floor. Dorsum and venter (Figs 5, 6A, B): Dorsal skin smooth bedecked with several pustules; low \ /-shaped scapular folds, bearing one pair of subconical tubercles on its anterior part, and another pair near its posterior end; scapular folds continue posteriorly with rows of small pustules to form almost negligible dorsolateral folds; cloacal sheath without tubercles; skin on chest weakly areolate, belly areolate; discoidal fold distinct. Forelimbs (Fig. 6C): Very low ulnar tubercles; Finger I slightly shorter than Finger II; discs narrowly expanded, elongate acuminate, although slightly more rounded than discs on toes; all fingers having ventral pads surrounded by circumferential grooves; palmar tubercles not well defined, outer palmar tubercle bifid, approximately twice the size of the ovoid thenar tubercle; distinct subarticular tubercles; low hyperdistal subarticular tubercles; indistinct supernumerary tubercles; fingers bearing narrow lateral fringes; basal webbing slightly noticeable. Hindlimbs (Fig. 6D): Heel bearing one low subconical tubercle; several subconical tubercles and some pustules on outer surface of tarsus; inner tarsal fold distinct; Toe V slightly longer than Toe III; discs narrowly expanded, elongate acuminate, nearly as large as those on fingers; all toes having ventral pads surrounded by circumferential grooves; ovoid inner metatarsal tubercle, approximately four times the size of the small and rounded outer metatarsal tubercle; subarticular tubercles distinct; hyperdistal subarticular tubercles low; supernumerary tubercles indistinct; toes bearing narrow lateral fringes; basal webbing, most distinctive between Toe IV and Toe V. Skin on anteroventral surface of thighs weakly areolate, posteroventral surface of thighs areolate. Variation: The four individuals of P. nanus with available sequences show a mean intraspecific uncorrected genetic p- distance of 0.3% in the 16S gene. In the type series, adult males (SVL = 11.04– 12.46 mm) are between 11.51 and 25.51% smaller than adult females (SVL = 14.08–14.82 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 7 to 9. H e a d (F i g s 7 – 9): T h r e e s u b c o n i c a l postrictal tubercles (QCAZ 72595); two subconical postrictal tubercles (QCAZ 72616). Upper eyelid with several pustules (QCAZ 68616); one subconical tubercle (QCAZ 72595). Occipital subconical tubercle (QCAZ 72616). Dorsum and venter (Figs 7–9): Scapular folds almost undistinguishable (QCAZ 68616); prominent (QCAZ 72616). Pair of high subconical scapular tubercles towards the posterior end of the scapular folds (QCAZ 72616). Dorsolateral folds absent (QCAZ 72598). Distinct subconical sacral tubercles (QCAZ 72616). Dorsal skin uniformly smooth (QCAZ 68616). Belly and posteroventral surface of thighs coarsely areolate (QCAZ 68569). Forelimbs (Fig. 9): Fingers without lateral fringes (QCAZ 72598). Basal webbing absent (QCAZ 68618). Outer palmar tubercle completely divided (QCAZ 72598). Supernumerary tubercles at base of fingers (QCAZ 72598). Hindlimbs (Fig. 9): Dorsal surface of shanks highly pustulated (QCAZ 68569). Inner tarsal fold inconspicuous (QCAZ 72616). Toe V longer than Toe III (QCAZ 68618). Basal webbing absent (QCAZ 72598). Distribution, natural history and conservation status: This species is only known from a single locality in the buffer zone of El Quimi Biological Reserve, in Morona Santiago Province, Ecuador, between 1978 and 2209 m a.s.l. (Fig. 3). The natural region is Eastern Montane Forest. The type locality is on a limestone tepui plateau on the eastern side of the Quimi Valley. It was found on mosses, leaves, branches and bromeliads from the ground up to a height of 100 cm. The new species lives on forest (Fig. 4) composed of thin, scattered trees 10–15 m high, shrubby vegetation 1.5 m high and soils with cushioned consistency, covered by mosses, roots and terrestrial bromeliads. This type of soil is locally known as bamba and is characteristic of limestone tepui formations. Individuals were collected about 3 km or more from the nearest disturbed area (pastureland). All observations were made at night. According to available data, this species has a restricted distribution (Extent of Occurrence 0.078 km 2, Area of Occupancy 12 km 2). However, adjacent areas remain unexplored, so we assign P. nanus to the Data Deficient Red List Category (IUCN, 2021). The type locality is Etymology: The specific name is derived from the Latin word nanus, meaning dwarf or small.

PRISTIMANTIS TRACHYBLEPHARIS SPECIES GROUP Definition: The Pristimantis trachyblepharis species group is strongly supported in our phylogeny. Members of this group share the following morphological traits: (1) tympanic membrane present (except for P. minimus) and tympanic annulus distinct; (2) cranial crests absent; (3) dentigerous processes of vomer usually present; (4) males without vocal slits (except for P. minimus) and nuptial pads; (5) dorsal skin smooth to weakly tuberculate; (6) scapular folds distinct (except for P. minimus); (7) discs of fingers and toes narrowly expanded to expanded; (8) Finger I shorter than Finger II; (9) toes with narrow lateral fringes (except for P. minimus); (10) Toe V slightly longer to longer than Toe III; and (11) small size (SVL range in adult females from 14.08 to 23.76 mm, in adult males 11.04 to 16.80 mm). Content: The Pristimantis trachyblepharis species group comprises eight described species (four of them are described below): P. albujai Brito et al., 2017, P. aquilonaris Lehr et al., 2007, P. minimus Terán-Valdez & Guayasamin, 2010, P. nanus, P. pramukae, P. trachyblepharis (Boulenger, 1918), P. ujucami and P. ventristellatus. Distribution (Figs 3, 4): Eastern Andean slopes of central and southern Ecuador, and northern Peru, in five provinces: Morona Santiago, Pastaza, Tungurahua and Zamora Chinchipe in Ecuador and Huancabamba (Piura Department) in Peru. Species inhabit Eastern Montane Forest and Eastern Foothill Forest (Ron et al., 2019), between 981 and 2567 m a.s.l. They are found on low vegetation from the ground up to a height of 300 cm in forested areas with low vegetation. Remarks: According to our phylogeny, the P. trachyblepharis species group is strongly supported as sister to a clade composed of 18 species (Supporting Information, Figs S3, S 4) that have been included in several different groups (e.g. P. chalceus group, P. myersi group and P. unistrigatus group) or have not been assigned to any group (Hedges et al., 2008; Padial et al., 2014)., Published as part of Zumel, Daniel, Buckley, David & Ron, Santiago R, 2022, 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, pp. 315-354 in Zoological Journal of the Linnean Society 195 (1) on pages 323-325, DOI: 10.1093/zoolinnean/zlab044, http://zenodo.org/record/6530571, {"references":["Brito J, Batallas D, Yanez-Munoz MH. 2017. Ranas terrestres Pristimantis (Anura: Craugastoridae) de los bosques montanos del rio Upano, Ecuador: lista anotada, patrones de diversidad y descripcion de cuatro especies nuevas. Neotropical Biodiversity 3: 125 - 156.","Boulenger GA. 1918. Descriptions of new South American batrachians. Annals and Magazine of Natural History 2: 427 - 433.","Ron SR, Merino-Viteri A, Ortiz DA. 2019. Anfibios del Ecuador, v. 2019.0. Museo de Zoologia, Pontificia Universidad Catolica del Ecuador. Electronic database accessible. Available at https: // bioweb. bio / faunaweb / amphibiaweb / (accessed 20 January 2020).","Hedges SB, Duellman WE, Heinicke MP. 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa 1737: 1 - 182.","Padial JM, Grant T, Frost DR. 2014. Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa 3825: 1 - 132."]}