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In this study we describe for the first time the calls of Pristimantis galdi and Pristimantis katoptroides. Recordings were obtained in Sangay National Park, Ecuador. We highlight the importance of recording P. galdi since its call has been recorded after 153 years of having been described as a species. The call of P. galdi consists of 7 to 9 short notes, the sounds of which are similar to a hammer hitting a nail, with a mean dominant frequency of 2.39 kHz. In turn, the call of P. katoptroides consists of a single note, the sound of which is similar to a metallic “tic”, with a mean dominant frequency of 1.74 kHz. We compared the advertisement calls of P. galdi, P. katoptroides and P. roni as these species share similar morphological characteristics and are grouped in the spiny green frog’s ecotype. Despite these morphological similarities, their advertisement calls are different. Obtaining calls of Pristimantis species in Ecuador might prove difficult with short-term studies due to the great sampling efforts that may be needed to get these recordings. Therefore, implementing active and passive monitoring could help improve our knowledge of acoustic signals in Ecuador’s rainfrogs.

The rainfrogs of the genus Pristimantis are one of the most diverse groups of vertebrates, with outstanding reproductive modes and strategies driving their success in colonizing new habitats. The rate of Pristimantis species discovered annually has increased continuously during the last 50 years, establishing the remarkable diversity found in this genus. In this paper the specifics of publications describing new species in the group are examined, including authorship, author gender, year, language, journal, scientific collections, and other details. Detailed information on the descriptions of 591 species of Pristimantis published to date (June 2022) were analyzed and extracted. John D. Lynch and William E. Duellman are the most prolific authors, yet Latin American researchers have scaled up and continued the description processes since the 1990s. The most common language used for descriptions is English, followed by Spanish. The great majority of authors have described only one species. The largest proportion of authors who have participated in the descriptions is of Ecuadorian nationality. Ecuador is the country with the highest description rate per year (3.9% growth rate). Only 20% of the contributions have included women and only 2% have featured women as principal authors. 36.8% of the species described are in the Not Evaluated or Data Deficient categories under the IUCN global red list. The importance of enhancing the descriptions in Spanish is emphasized and the inclusion based on equal access to opportunities for female researchers in Pristimantis taxonomy is encouraged. In general, if the current trends in Pristimantis descriptions continue, in ten years, a total of 770 or more species described could be expected.

Phrynopus sancristobali sp. nov. Figures 3–7, Table 1 urn:lsid:zoobank.org:act: 9B5329CC-7045-4EE5-B71F-A6DD3ACC0DFF Holotype PFAUNA 552, adult male, from Escalerapampa, District of Ayahuanco, Province of Huanta, Department of Ayacucho, Peru (12°32’50”S, 74°15’07”W, 3910 m a.s.l.) collected on 16 January 2020 by Vladimir Díaz and Randy Llantoy. Figures 3–5. Paratypes Two adults from near the type locality: female PFAUNA 550 (12°33’8”S, 74°15’02”W, 3910 m a.s.l.) and female PFAUNA 551 (12°32’50”S, 74°15’07”W, 3910 m a.s.l.) collected on 16 January 2020 by V. Díaz and R. Llantoy. Figure 6–7. Description Phrynopus sancristobali sp. nov. is distinguished by a combination of the following characters: (1) skin on dorsum and flanks with large, round and irregular pustules, forming irregular ridges; skin on the belly strongly areolate; discoidal fold absent, thoracic fold absent; dorsolateral folds absent; postocular fold prominent; (2) tympanic membrane and tympanic annulus absent; (3) snout short, round in dorsal view, curved antero-ventrally in lateral view; (4) upper eyelid without tubercles; width of upper eyelid slightly narrower than IOD, cranial ridges absent; (5) dentigerous processes of vomers absent; (6) vocal slits and nuptial pads absent; (7) Finger I shorter than Finger II; rounded and bulbous fingertips, lacking discs or grooves; (8) fingers without basal webbing and without lateral fringes; (9) ulnar tubercles absent, tarsal tubercle absent; (10) thenar and palmar tubercles small, internal tarsal fold absent; (11) internal metatarsal tubercle rounded, larger than external, ovoid metatarsal tubercle; subarticular tubercles entire on all fingers, supernumerary plantar tubercles absent; (12) toes without lateral fringes, basal webbing absent, Toe V slightly longer than Toe III, bulbous fingertips, no discs; (13) in life, dorsum and dorsal parts of extremities light brown, with dark brown reticulations that appear to divide the pustules; flanks golden brown with gray reticulations and dark brown; palms of hands orange, Toes IV and V dark brown, plantar surface of foot of a more intense orange color; belly cream to yellow with gray to light brown crosslinks, groin and throat orange; iris silvery with dark brown marks; (14) SVL 20.7 and 22.2 mm in two females and 19.5 in one male. Diagnosis Phrynopus sancristobali differs from P. auriculatus, P. montium, P. mariellaleo and P. peruanus by lacking tympanic membrane and annulus; from P. auriculatus, P. remotum, P. inti, P. bracki, P. dagmarae, P. interstinctus, P. lechriorhynchus, P. kauneorum, P. kotosh, P. peruanus and P. vestigiatus by lacking dentigerous processes of vomers; from P. juninensis, P. kauneorum, P. inti, and P. apumantarum by the presence of prominent postocular folds; from P. chaparroi, P. thompsoni, P. paucari, P. heimorum, P. remotum, P. anancites, P. miroslawae, P. juninensis, P. kauneorum, P. tautzorum, and P. apumantarum by the presence of dorsal pustules with large, round and irregular warts; from P. remotum, P. anancites, P. miroslawae, P. juninensis, P. kauneorum, P. tautzorum, and P. apumantarum by the presence of pustules on dorsum; from P. inti, P. tribulosus, P. valquii, P. oblivius P. auriculatus, P. kauneorum, P. bracki, P. mariellaleo and P. tribulosus by having ventral skin strongly areolated. Phrynopus sancristobali most resembles P. bufoides, from which it differs by having irregular pustules on the dorsal part of the fore and hind limbs, maximum SVL 22.2 mm in females, ventral coloration in females cream with irregular yellow spots, and throat and ventral surface of arm and leg mostly orange (pustules arranged in longitudinal rows, maximum SVL 32.9 mm in females, venter and throat coloration in females gray with dark-brown spots irregularly distributed, and ventral surface of leg mostly dark-brown in P. bufoides). Description of holotype Head nearly as wide as body, wider than long; head width 107% of head length; head width 37% of snout-vent length; head length 35% of SVL. Snout short, rounded in dorsal view, curved anteroventrally in lateral view (Figs. 3–5), eye-to-nostril distance 48% of eye diameter; nostrils not protuberant; canthus rostralis slightly curved in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid lacking tubercles, narrower than interorbital distance (EW 66% of IOD); postocular and tympanic region with prominent folds (Fig. 3); tympanic membrane and tympanic annulus absent, tympanic region with discontinuous pustules; choanae small, ovoid, partly concealed by palatal of maxilla, dentigerous processes of vomers absent; tongue broad, about 1.7 as long as wide, notched posteriorly, posterior half free; vocal slits absent. Skin on dorsum with circular pustules of varying dimensions (Fig. 4); skin on flanks shagreened with scattered rounded pustules; skin on throat, chest and belly strongly areolate (Fig. 3B); discoidal and thoracic folds absent; cloacal sheath absent; cloacal region with prominent pustules. Outer surface of forearm with prominent pustules; outer palmar tubercle barely visible, low, ovoid, smaller than ovoid inner palmar tubercle; supernumerary tubercles absent; subarticular tubercles absent; fingers without lateral fringes; Finger I slightly shorter than Finger II (Fig. 5); tips of digits rounded, bulbous, lacking circumferential grooves. Hind limbs short and slender, tibia length 34% of SVL; foot length 37% of SVL; dorsal surface of hind limbs with irregular pustules; anterior surfaces of thighs smooth, posterior surfaces of thighs pustulate; heel with low rounded, irregular tubercles; outer surface of tarsus bearing pustules like those on dorsum; outer metatarsal tubercle rounded and prominent, slightly larger than the ovoid inner metatarsal tubercle; supernumerary plantar tubercles absent; subarticular tubercles low, not distinguishable; toes without lateral fringes; basal webbing absent; toe tips bulbous, rounded, lacking circumferential grooves, about as large as those on fingers; relative lengths of toes: 1 Measurements of holotype (in mm). SVL, 19.5; TL, 6.6; HL, 6.8; IND, 1.7; IOD, 2.3; EW, 1.6; HW, 7.3; FL, 7.3; E-N, 1.1; ED, 2.2 (Table 1). Variation Female PFAUNA 550 is larger (SVL = 22.2 mm) than both the holotype and male PFAUNA 551 (SVL = 20.7 mm; Table 1). Furthermore, the female has the first subarticular tubercle entire in Finger I and IV, and divided in Fingers II and III (all subarticular tubercles entire in both males; Fig. 7). Coloration of holotype in life The holotype had a light brown to golden dorsum, with dark brown reticulations that appear to divide the pustules, and similar coloration on the dorsal part of the limbs; the flanks were golden brown with gray and dark brown reticulations; the belly was cream to yellow with gray to light brown reticulations, the groin and throat were orange; there is a dark brown halo on the nose and on the eye, extending as a post ocular stripe that ends in the tympanic palms and soles were orange, but fingers IV and V were dark brown in dorsal and ventral view, fading to orange at about half of palms (Fig. 3). Coloration of holotype in preservative The holotype in preservation maintains the same general color patterns described above, only decreasing in intensity (Fig. 4). Distribution and natural history Phrynopus sancristobali sp. nov. is only known from the type locality, which is characterized by the transition from humid puna to montane forest at elevations from 3796 to 3910 m a.s.l., in the upper watershed of the Caballuyoc river, tributary of the Mantaro river, within the Huanta province in the Ayacucho Department. The frogs were found under rocks in grasslands (Fig. 8) dominated by Stipa ichu, Stipa obtusa, Senecio cf. bukatii, Oreithales integrifolia, Halenia umbellata, Azorella creanata, and Chaptalia similis. Phrynopus sancristobali sp. nov. is sympatric with frog species Gastrotheca sp. and Pleurodema marmoratum, as well as the lizard Wilsonosaura josyi. Etymology The specific epithet honors the Universidad Nacional San Cristóbal de Huamanga, alma mater of the first author, in recognition of its commitment to forming professional biologists with a holistic approach to biodiversity conservation., Published as part of Díaz, Vladimir, Mamani, Luis & Catenazzi, Alessandro, 2023, A new species of Andean frog of the genus Phrynopus (Anura: Strabomantidae) from southeastern Peru, pp. 333-348 in Zootaxa 5293 (2) on pages 338-343, DOI: 10.11646/zootaxa.5293.2.7, http://zenodo.org/record/7960244

The Cordillera de los Andes is one of the most important regions for biodiversity. Among amphibians, many endemic species of terrestrial-breeding frogs have recently been discovered. Herein we describe Phrynopus sancristobali from the Andes of southeastern Peru based on molecular and morphological data. The new species is known from the ecotone between humid puna and montane forest at 3910 m a.s.l. on the left side of the Apurímac Valley in the Department of Ayacucho. The new species differs from congeners by having dorsum bearing pustules and light brown with dark brown reticulations surrounding the areolas, and coloration consisting of flanks golden brown with gray or dark brown marks, palms pale orange, soles deep orange, toes IV and V dark brown in dorsal and ventral view, belly cream to yellow with gray to light brown marks, and groin and throat deep orange. The snout-vent length (SVL) is 20.7 and 22.2 in two females, and 19.5 mm in one male. According to our phylogeny inferred using Maximum Likelihood with a concatenated dataset of three mitochondrial and two nuclear genes, P. sancristobali is sister taxon of P. apumantarum, recently described from Department Huancavelica. Our description extends the known geographic range of Phrynopus 73 km to the south, and P. sancristobali is the only species in the genus known to occur south of the Mantaro River, whose deep valley is hypothesized to be a biogeographic barrier for high-Andean organisms. The discovery of P. sancristobali confirms the high levels of endemism and beta diversity of Phrynopus in the moist puna grasslands and montane forests of the high Andes of Peru, and suggests that further work will reveal the presence of additional species in southern Peru.

The taxonomic status of the strabomantid frog species Pristimantis nebulosus (Henle, 1992), originating from the southern Cordillera Azul in central Peru, is investigated based on examination of the holotype and its comparison with freshly collected topotypic material. Following current standards, we provide a redescription of the holotype. It is in a rather poor state and exhibits certain damages and preservation artifacts, conditions that have hampered an allocation of this nominal taxon to any known living population of Pristimantis in the past. Our detailed specimen-to-specimen comparison provided morphological evidence for the conspecifity of the holotype and freshly collected topotypes. Molecular phylogenetic analysis, based on the mitochondrial 16S gene fragment places P. nebulosus in the P. conspicillatus species group, being closely related to P. bipunctatus and an undescribed candidate species from the Cordillera de Carpish. From both, P. nebulosus differs by considerable divergence in the 16S gene (p-distance 4.1–6.2%). Based on the specimens available, we provide an updated diagnosis for P. nebulosus, compare it to other species in the P. conspicillatus group and describe its advertisement call. In addition, we describe and name the closely related candidate species from the Cordillera de Carpish. It is sister to P. bipunctatus and P. nebulosus, but differs from both mainly by its tuberculate dorsal skin (versus shagreen) and divergence in the 16S gene (3.3–4.1%). We briefly discuss cryptic species diversity in the P. conspicillatus and P. danae species groups and provide justification for the description of a singleton species.

Pristimantis is already the most speciose genus among vertebrates, yet the current number of species remains largely underestimated. A member of the P. unistrigatus species group from the Guiana Shield has been historically misidentified as P. ockendeni, a species described from southern Peru. We combined mitochondrial (16S and COI) and nuclear (RAG1) loci, external morphology, skull osteology (μ-CT scan), vocalization (advertisement and courtship calls), geographic distribution and natural history data to differentiate the Guiana Shield populations from P. ockendeni, and describe them as a new species. The new species is crepuscular and nocturnal and inhabits the understory of unflooded (terra firme) forests in Brazil, Guyana and Suriname. It is phylogenetically related to P. arda­lonychus, P. martiae and undescribed species from Brazilian Amazonia. The new species notably differs from P. ockendeni and its congeners in the P. unistrigatus species group occurring in the Guiana Shield by the combination of the following characters: absence of dentigerous processes of vomers, presence of vocal slits in males, body size (SVL 16.2–20.7 mm in males and 21.4–25.7 mm in females), advertisement call (call with 4–6 notes, call duration of 158–371 ms and dominant frequency of 3,466–4,521 Hz) and translucent groin coloration in life. To facilitate the recognition and description of cryptic species previously hidden under the name P. ockendeni, we provide an amended diagnosis of this taxon based on external morphology and advertisement call of specimens recently collected nearby the type locality and additional localities in southwestern Amazonia.

With 566 species, the neotropical genus Pristimantis is the most speciose vertebrate genus. As a result of its striking diversity, taxonomic reviews remain a challenge. Herein, we present an updated phylogeny of the Pristimantis lacrimosus group and describe a new species from Llanganates and Sangay National Parks. We also report, for the first time, the phylogenetic position of Pristimantis degener, P. eugeniae, P. katoptroides, and P. petersi. Based on our phylogeny, we add two species to the Pristimantis lacrimosus group. Through the integration of molecular and bioacoustic evidence, we describe a new species which was hidden under “Pristimantis petersi”. Pristimantis petersioidessp. nov. is most closely related to Pristimantis petersi and an undescribed species from Peru. It can be distinguished from P. petersi by its advertisement call and large genetic differences (uncorrected p-genetic distances 7.9% to 8.4% for gene 16S). Moreover, the new species and P. petersi are not sister species. We suggest assigning the new species to the Endangered Red List category because it has a small distribution range with deforestation as result of agriculture and other anthropogenic influences.

Family Craugastoridae Hedges, Duellman & Heinicke, 2008 Pristimantis marmoratus (Boulenger, 1900) is currently distributed in the western portion of the Guiana Shield, through Venezuela and Guyana, and its sister clade has a more eastern distribution, occurring in French Guiana and Amapá (Kok et al. 2018). Fouquet et al. (2022b) recently described this clade as Pristimantis crepitaculus Fouquet, Peloso, Jairam, Lima, Mônico, Ernst & Kok, 2022. We considered four out of our five records of P. marmoratus (Lima 2008; 2018; Queiroz et al. 2011; Benício & Lima 2017) as P. crepitaculus. Pristimantis ockendeni (Boulenger 1912) was once thought to occur all over the upper Amazon basin, from southern Peru to Colombia, but it is currently thought to be a species complex with the nominal P. ockendeni currently known only from the type locality, in Peru (Elmer et al. 2007; Elmer & Canatella 2008). Silva e Silva & Costa-Campos (2018) recorded a species with overall morphology similar to P. ockendeni which they identified as P. cf. ockendeni , but they also recorded P. marmoratus. Besides P. crepitaculus, Fouquet et al. (2022b) also mention P. grandoculis from the state of Amapá, a species they revalidated in their study. The main morphological difference between these two species is the tympanum, which is present in the first species and absent in the latter. Thus, we consider the records of Silva e Silva & Costa-Campos (2018) as P. crepitaculus (their P. cf. ockendeni) and P. grandoculis (their P. marmoratus).

We herein present the first annotated anuran checklist for the Brazilian state of Amapá, eastern Amazonia, based on a thorough literature review. We recorded the occurrence of 111 species belonging to 13 anuran families distributed across 48 localities throughout Amapá, within two biomes. Among these species, 62.5% occur exclusively in the Tropical Moist Broadleaf Forest biome, ~8% occur exclusively in the Tropical Savanna biome, and ~29% occur in both. Two species were considered endemic to Amapá and were registered only in the central portion of the state. Regarding the conservation status, only one species (Dendropsophus amicorum) is classified as threatened, assigned to the “critically endangered” category. The other species are categorized as either “least concern” or “data deficient” (85 and 8, respectively), whereas 21 are not evaluated. The current annotated list contributes to the incipient knowledge on anuran species richness in Amapá and, despite the research regarding anuran taxonomy has considerably progressed over the past 20 years, there is still much to do. Our data highlight the need for trained taxonomists to develop research in the state.

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., 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 325-330, DOI: 10.1093/zoolinnean/zlab044, http://zenodo.org/record/6530571, {"references":["Lehr E, Duellman W. 2007. A diminutive new species of Pristimantis (Amphibia: Anura: Leptodactylidae) from northern Peru. Salamandra 43: 165 - 171.","Lehr E, Coloma LA. 2008. A minute new ecuadorian Andean frog (Anura: Strabomantidae, Pristimantis). Herpetologica 64: 354 - 367.","Duellman WE, Pramuk JB. 1999. Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers Natural History Museum the University of Kansas 13: 1 - 78.","IUCN. 2021. The IUCN Red List of Threatened Species. Available from: https: // www. iucnredlist. org (accessed 18 May 2021)."]}

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."]}

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.

Craugastor portilloensis sp. nov. Holotype. — UTA A-62393 (field ID: JAC 21431; Fig. 32), subadult female collected by E.N. Smith and colleagues in Portillo del Rayo, Distrito San Pedro Pochutla, Sierra Madre del Sur, Oaxaca, Mexico, 15.9794448N, 96.5166678W, 1550 m, 1 October 2001. Paratypes (4). —MZFC-HE-35580 and UTA A- 66095, juvenile males. MZFC-HE- 35581 subadult female, and UTA A- 66096 subadult male. All collected at the type locality between 1550 and 1585 mon 24 September 2001 by E.N. Smith and colleagues. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 11.4 mm); (2) reduced ossification of skeleton in adults relative to other members of the series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank darker than dorsum, faintly barred lips, snout–nostril– canthal–supratympanic stripe; (8) one or two postrictal tubercles; (9) gular region with pale spotting; (10) dorsal surface unicolored pale, diffuse interorbital bar, sometimes with two dark rump spots; (11) with or without a middorsal ridge; (12) dorsum smooth or with scattered fine tubercles; (13) body flank has dark supratympanic stripe extending toward lower mid-flank, smooth, very few small tubercles; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and slightly past snout; (16) outer tarsal ridge smooth or with 1–3 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not expanded, toe tips slightly expanded; (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor portilloensis can be differentiated from C. candelariensis, C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by the presence of vomerine odontophores (absent in C. portilloensis). It can be differentiated from C. bitonium, C. hobartsmithi, and C. pygameus by the absence of a posterolateral projection of the frontoparietal (present in C. portilloensis). It can be further differentiated from C. pygmaeus by the absence of a canthal mask (present in C. portilloensis). It can be differentiated from C. cueyatl and C. polaclavus by metatarsal tubercles of unequal size (equal in C. portilloensis). It can be differentiated from C. rubinus by relative finger lengths of IV ¼ II (IV> II in C. portilloensis). Description of holotype. — Holotype small female (SVL ¼ 11.4 mm); snout rounded and short (0.8 mm naris–snout; 6% SVL); short eye–nostril distance (1.1 mm; 9.8% SVL); tympanum 0.8 mm (6.8% SVL); no supratympanic fold; small shoulder tubercle; finger length formula III Variations in paratypes. —Body sizes in SVL 8.7 mm (MZFC-HE-35580), 8.4 mm (UTA A-66095), 12.1 mm (MZFC-HE-35581), 12.6 mm (UTA A-66096); eye–nostril distance 10–11% SVL; tympanic ratios 7%. Etymology. —The name is an abbreviated allusion to the type locality near the town of Portillo del Rayo and the Latin suffix - ensis meaning place. Distribution. —Intermediate elevations in the foothills of the Sierra Madre del Sur in Oaxaca 1550–1585 m (Fig. 8). These habitats consist of mixed tropical dry and temperate sierra forests. Phylogenetics. —In the concatenated analysis, Craugastor portilloensis is the sister taxon to a clade of small-bodied species (C. bitonium, C. candelariensis, C. cueyatl, C. hobartsmithi, C. polaclavus, C. pygmaeus, and C. rubinus), but with low branch support (ML ¼ 53, BAYES ¼ 66; Fig. 3). It occupies the same phylogenetic placement in the mtDNA-only data set (Fig. 4), but in the nDNA-only data set is the sister taxon of a clade including (C. bitonium, C. cueyatl, C. hobartsmithi, C. pygmaeus, and C. rubinus) with moderate support in the Bayesian analysis (ML ¼ 43, BAYES ¼ 0.78; Fig. 5). In terms of genetic distances (Table 4), C. portilloensis was most similar to C. montanus (5.7%), followed by similarity to C. polaclavus (5.8%). Remarks. —The skull of C. portilloensis is similar to C. mexicanus, C. omiltemanus, and C. saltator, with a more anteriorly placed anterior suture of frontoparietal and prootic than in other species. We dissected all specimens of C. portilloensis; males possess pigmented testes and females unpigmented ovaries. This species likely co-occurs with C. candelariensis, C. polaclavus, and C. pygmaeus in southcentral Oaxaca (Figs. 6 and 8). It was collected in sympatry with C. polaclavus at the type locality of Portillo del Rayo, Oaxaca, Mexico (Fig. 8)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 33-34, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastormexicanus (Brocchi 1877) Leiuperus mexicanus Brocchi 1877:184. Holotype unsexed (MNHNP 6218) from ‘‘ Mexico ’’ (¼ Cerro San Felipe, Oaxaca, Mexico [Smith and Taylor 1950]). [Examined]. Paludicola mexicana (Brocchi): Boulenger 1882:237; Neiden 1923:513. Pleurodema mexicana (Brocchi): Parker 1927:475. Microbatrachylus oaxacae Taylor 1940:505. Holotype male (FMNH 100001) from ‘‘Cerro San Flipe, near Oaxaca, Oaxaca, Mexico.’’ [Examined]. Microbatrachylus lineatissimus Taylor 1941:87. Holotype male (FMNH 1000036) from ‘‘Cerro San Flipe, near Oaxaca, Oaxaca, Mexico.’’ [Examined]. Microbatrachylus fuscatus Davis and Dixon 1957:146. Holotype female (TCWC 12171) from ‘‘ 20 miles east of Tulancingo, Hidalgo, Mexico.’’ [Not examined; neartopotypic specimen from Hidalgo examined (UTA A-66138)]. Eleutherodactylus oaxacae (Taylor): Lynch 1965:3. Eleutherodactylus lineatissimus (Taylor): Lynch 1965:3. Eleutherodactylus mexicanus (Brocchi): Gorham 1966:86. Craugastor mexicanus (Brocchi): Crawford and Smith 2005:536. Diagnosis. —Based on 26 specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) large adult size (maximum SVL ¼ 40.5 mm); (2) full ossification of the skeleton in adults; (3) presence of posterolateral projection of frontoparietal (Fig. 26B); (4) presence of vomerine odontophores (in larger individuals); (5) presence or absence of raised tubercles on eyelids, Comparisons. — Craugastor mexicanus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by the equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. mexicanus). It can be differentiated from C. bitonium and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. mexicanus). It can be differentiated from C. polaclavus and C. rubinus by the absence of vomerine odontophores (present in C. mexicanus). It can be differentiated from C. montanus by the condition of supratympanic folds in adults; developed in C. mexicanus versus moderate to poorly developed in C. montanus. It can be differentiated from C. omiltemanus by ventral skin texture in life; smooth to granular in C. mexicanus versus areolate in C. omiltemanus. Craugastor mexicanus is most similar to C. saltator. We were unable to identify any reliable morphological characters to differentiate C. mexicanus and C. saltator; however, they have nonoverlapping geographic distributions, with C. saltator only occurring in western Guerrero (Fig. 6). Description. —In previous literature, described as largebodied, long-legged with row of small tubercles on outer edge of the tarsus (Taylor 1941); variable palmar tubercle arrangements (palmar tubercle divided, single, or evaginated; Lynch 1965); inner metatarsal tubercle larger than outer metatarsal tubercle; lack of tarsal tubercles (Lynch 2000). Holotype (MNHNP 6318) is large (~ 40 mm SVL); owing to poor preservation, columella of holotype partially ruptured the tympana on both sides (Fig. 1B); finger lengths III> IV> II> I; toe length IV> V ¼ III> II> I. Distribution. —This species is widespread throughout eastern Mexico in high-elevation habitats (1554–2700 m) of Oaxaca, Puebla, Hidalgo, and Veracruz (Fig. 6). These habitats span the Sierra Madre Oriental and Sierra Madre del Sur. Canseco Márquez and Gutiérrez Mayén (2010) report that C. mexicanus occurs in the forests adjacent to the Tehuacán-Cuicatlán Valley in Puebla and Oaxaca. It is likely this species occurs in Guerrero; however, most specimens resembling C. mexicanus we examined from Guerrero are referrable to C. saltator. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Phylogenetics. —The concatenated data set placed C. mexicanus as the sister taxon to a clade of C. omiltemanus þ C. saltator with high support in the BAYES analysis (0.98) but moderate support in the ML analysis (75; Fig. 3). This appears to be a relationship supported exclusively by the mtDNA data set because separate analysis of the nDNA markers did not confidently infer a sister taxon of C. mexicanus (Figs. 4 and 5). Although BAYES analyses strongly supported the monophyly of C. mexicanus (> 0.90), the ML analyses recovered variable support ranging from 79 (concatenated analysis) to 54 (nDNA-only analysis). In terms of genetic distances (Table 4), C. mexicanus was most similar to C. omiltemanus (4.9%), followed by similarity to C. saltator (5.1%). Intraspecific variation. —We examined over 220 specimens of C. mexicanus for this revision (Appendix I), and briefly provide some patterns of intraspecific variation that we observed. Many populations have substantial colorpattern polymorphism (Fig. 27), similar to what is observed in the C. rhodopis series (Lynch 1966; Streicher et al. 2014). In some northern populations (Hidalgo and Puebla), the canthal mask is broken into a black spot posterior to the tympanum (often with a thin yet distinctive white margin). Specimens throughout Oaxaca varied in whether they had a single or divided palmar tubercle (as reported by Lynch 1965, see his Fig. 2); in six specimens we examined there was asymmetry with a single palmar tubercle on one hand and divided on the other. Patterns of dorsal ridging and coloration often coincide, with observations of the following morphs: (1) straight raised ridges that are tubercular and darker in coloration than the rest of the dorsum; (2) straight raised dorsal ridges that are tubercular but of the same color as the rest of the dorsum; (3) straight lines of color that are dark but with no tubercular raised ridges; (4) ridges that are not straight but form an hourglass shape and are tubercular; (5) ridges that are not straight but form an hourglass shape and are not tubercular; (6) scattered tubercles on the dorsum, which are sometimes dark and sometimes same color as the background; (7) pale or dark unicolored dorsum; (8) a unicolored dorsum with a pale median stripe that can be ridged or smooth; (9) a wide dark band on the dorsum, which usually co-occurs with a pale upper labium; and, (10) white hills of coloration and a peppered grayish dorsum sometimes with a diffused lighter cream color, a pattern that may mimic bird droppings (appearing superficially as a smear of uric acid and digestive waste). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Remarks. —The skull of C. mexicanus is similar to that of C. omiltemanus and C. saltator, with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. We noticed that the ‘‘ M. lineatissimus ’’ morphotype (raised and parallel ridges on the dorsum) occurs widely throughout the range of C. mexicanus. Interestingly, the examined paratype of ‘‘ M. lineatissimus ’’ has a unique skull shape for C. mexicanus. The shape is similar to that of C. hobartsmithi, C. montanus, and C. pygmaeus, with a more posteriorly placed anterior suture of the frontoparietal and prootic than in other species, coupled with a narrower back of the skull than in any other species (Fig. 27A). This unique condition likely explains the specimen as an outlier in several of our statistical analyses (Figs. 9, 11, and 12). However, we were unable to CT-scan other individuals with the ‘‘ M. lineatissimus ’’ morphotype (or the holotype of M. lineatissmus), so future investigation is necessary to determine whether the examined paratype (FMNH 104548) is an aberrant individual of C. mexicanus or represents a valid taxon. Lynch (1965) reports that ‘‘ E. oaxacae ’’ has parotoid glands. However, we saw no evidence of these glands in the two type specimens that we examined (FMNH 100001 and FMNH 126638), nor are we aware of any species of Craugastor that possess parotoid glands (¼ large poison glands on the nuchal region). Craugastor mexicanus likely co-occurs with C. omiltemanus at high-elevation localities of central Oaxaca (Figs. 6 and 8). At intermediate elevation localities in Veracruz and Oaxaca, it may overlap with C. pygmaeus (Fig. 7). Male C. mexicanus have significantly larger tympana than do female C. mexicanus (Fig. 13)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 27-29, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Brocchi, P. 1877. Sur quelques batraciens raniformes et bufoniformes de l'Amerique Centrale. Bulletin de La Societe Philomathique de Paris 1: 175 - 179. [In French.]","Smith, H. M., and E. H. Taylor. 1950. Type localities of Mexican reptiles and amphibians. The University of Kansas Science Bulletin 33: 313 - 339.","Boulenger, G. A. 1882. Catalogue of the Batrachia, Salientia and Ecaudata in the Collection of the British Museum. Natural History Museum Publications, UK.","Neiden, F. 1923. Anura. I. Subordo Aglossa und Phaneroglossa, Sectio 1. Arcifera. Das Tierreich 46: 32 - 584. [In German.]","Parker, H. W. 1927. A revision of the frogs of the genera Pseudopaludicola, Physalaemus, and Pleurodema. Journal of Natural History 20: 450 - 478.","Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571.","Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94.","Davis, W. B., and J. R. Dixon. 1957. Notes on Mexican amphibians, with description of a new Microbatrachylus. Herpetologica 13: 145 - 147.","Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Canseco Marquez, L., and M. G. Gutierrez Mayen. 2010. Anfibios y Reptiles del Valle Tehuacan-Cuicatlan. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Mexico.","Lynch, J. D. 1966. Multiple morphotypy and parallel polymorphism in some neotropical frogs. Systematic Biology 15: 18 - 23.","Streicher, J. W., U. O. Garcia-Vazquez, P. Ponce-Campos, O. Flores-Villela, J. A. Campbell, and E. N. Smith. 2014. Evolutionary relationships amongst polymorphic direct-developing frogs in the Craugastor rhodopis Species Group (Anura: Craugastoridae). Systematics and Biodiversity 12: 1 - 22."]}

Craugastor candelariensis sp. nov. Holotype. — UTA A-64253 (field ID: JAC 21885), male collected by E.N. Smith and colleagues Nof Candelaria on the road to Oaxaca; Sierra Madre del Sur, Oaxaca, Mexico, 15.94960°N, 96.47110°W, 668 m, on 21 January 2002 between 1130 and 1200 h, near stream bordering coffee plantation and secondary forest. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Paratypes (3). —MZFC-HE-35617 (formerly UTA A-64252; field ID: JAC 21873; Fig. 23), male with heavily pigmented testes, same data as holotype except collected 1.2 mi on rough road toward Pluma Hidalgo on the Candelaria – Portillo road, 15.95610°N, 96.44930°W, 1051 m, on 21 January 2002 at 1000 h in leaf litter of coffee plantation. UTA A- 66116 (Field ID: JAC 21851), male with pigmented testes collected by E.N. Smith and colleagues from San Gabriel Mixtepec, Puente de Hamaca, Oaxaca, Mexico, 16.10510°N, 97.06310°W, 710 m, on 20 January 2002 at 1520 h on forest floor. UTA A- 55247 (Field ID: ENS 9698), female with unpigmented gonads and extended oviducts collected by Karin S. Castaneda along the Carretera San Gabriel Mixtepec–Miahuatlán of the Sierra Madre del Sur, Oaxaca, Mexico, 16.160556°N, 97.00111°W, 1270–1350 m, on 15 March 1998 at 1630 h from pine forest habitat. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 18.6 mm); (2) full ossification of most skeletal elements in adults, lacking ossification only of Stage 6 (Table 3); (3) absence of posterolateral projection of frontoparietal; (4) presence of vomerine odontophores; (5) absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank with nostril– canthal–supratympanic stripe, lips colored as dorsum; (8) two postrictal tubercles; (9) gular region uniformly pale to slightly evenly peppered with melanocytes; (10) dorsal surface unicolored pale; (11) pale middorsal ridge, sometimes with few tiny spots; (12) evenly fine tubercles on dorsum; (13) body flank unicolored pale, shagreened with fine tuberculation; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and tip of snout; (16) outer tarsal ridge with 3–8 tiny and pointed tubercles on slightly raised fringe; (17) finger and toe tips lanceolate to mucronate (toes and outer two fingers); (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor candelariensis can be differentiated from C. bitonium, C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. pygmaeus, C. rubinus, and C. saltator by a larger innermetatarsal tubercle (inner and outer metatarsal tubercles are similar sizes in C. candelariensis). Craugastor candelariensis can be differentiated from C. cueyatl and C. hobartsmithi by the absence of vomerine odontophores (present in C. candelariensis). It can be differentiated from C. portilloensis by the presence of posterolateral projections of the frontoparietal (absent in C. candelariensis). Description of holotype. — Holotype small male (SVL ¼ 13.3 mm); snout rounded and short (0.5 mm naris–snout; 4% SVL); long eye–nostril distance (1.7 mm; 13% SVL); tympanum 1.2 mm (7.6% SVL); no supratympanic fold and no shoulder tubercle; finger length formula III ¼ I; single palmar tubercle; single prepollical tubercle; subarticular tubercles present on all fingers; supernumerary tubercles present on Finger III; toe length formula IV Variations in paratypes. —Body sizes (SVL) 12.4 mm (MZFC-HE-35617), 14.3 mm (UTA A-66116), 18.6 mm (UTA A-55247); eye–nostril distance 10–13% SVL (males), 9% SVL (female); tympanic ratios 7–10%. Etymology. —The name is an abbreviated allusion to the municipality of Candelaria Loxicha (near the type locality) and the Latin suffix - ensis meaning place. It is simultaneously a reference to the Latin noun candēla meaning a fire or light made of wax, given the translucent yellow appearance of several type specimens in preservative, as if someone were shining a candle through them. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Distribution. —This species is known from intermediate elevations of southern Oaxaca (668–1350 m), an area that mostly consists of Sierra Madre del Sur pine–oak forest habitat. Phylogenetics. — Craugastor candelariensis was strongly supported as monophyletic in the concatenated analysis (ML ¼ 100; BAYES ¼ 1.0; Fig. 3). In this analysis, the sister taxon of C. candelariensis was inferred to be C. polaclavus (ML ¼ 80; BAYES ¼ 0.99). We also observed this sister relationship in the nDNA-only analysis (Fig. 5); however, in the mtDNAonly analysis C. candelariensis was inferred to be the sister taxon of a clade containing C. bitonium þ C. pygmaeus (Fig. 4). In terms of genetic distances, Craugastor candelariensis is most similar to C. polaclavus and C. pygmaeus (both 6.4%; Table 4). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Remarks. —The skull of C. candelariensis is similar to C. bitonium, C. hobartsmithi, C. montanus, and C. pygmaeus, with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. Two type specimens appear white-yellowish in preservative (possibly having been saponified). This species likely co-occurs with C. pygmaeus, C. polaclavus, and C. portilloensis in southcentral Oaxaca (Figs. 6 and 8). In terms of body size and ossification level it is the smallest member of the C. mexicanus series to complete Stage 5 of our ontogenetic sequence., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 17-23, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastor pygmaeus (Taylor 1936) Eleutherodactylus pygmaeus Taylor 1936:352. Holotype female (UIMNH 16125) from ‘‘one mile north of Rodriguez Clara, Veracrux, Mexico.’’ [Examined]. Microbatrachylus albolabris Taylor 1940:502. Holotype female (FMNH 100071) from ‘‘two miles west of Córdoba, Veracruz, Mexico.’’ [Examined]. Microbatrachylus pygmaeus (Taylor): Taylor 1940:500. Microbatrachylus minimus Taylor 1940:507. Holotype male (FMNH 100323) from ‘‘Agua del Obispo, Guerrero, Mexico.’’ [Examined]. Microbatrachylus imitator Taylor 1942:70. Holotype female (USNM 115508) from ‘‘La Esperanza, Chiapas, Mexico.’’ [Examined]. Craugastor pygmaeus (Taylor): Crawford and Smith 2005:536. Diagnosis. —Based on 37 specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) small adult size (mean SVL ¼ 13.5 mm [SD ¼ 1.77], n ¼ 29); (2) full ossification of skeletal elements in adults; (3) lack of posterolateral projection of frontoparietal; (4) lack of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred canthus and jaws (rarely dark and blotched), no canthal stripe; (8) one (or two fused) postrictal tubercles; (9) gular region pigmentation present or absent; (10) dorsal surface two-toned usually with dark suprascapular ^^shape, or striped and with pale middorsal stripe; (11) variable middorsal ridge; (12) dorsum smooth or with only some large scattered tubercles; (13) body flank barred darker anteriorly, slightly shagreened to smooth; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between eye and slightly beyond tip of snout; (16) outer tarsal ridge smooth, no raised fringe; (17) finger and toe pads round, expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor pygmaeus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. pygmaeus). It can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. rubinus, and C. saltator by the presence of a posterolateral projection of the frontoparietal (absent in C. pygmaeus ; Fig. 26C). Craugastor pygmaeus is most similar to C. bitonium (in morphology, osteology, and genetic distance), but may be differentiated from this taxon by the condition of the outer tarsal ridge, which is smooth in C. pygmaeus versus 1–6 small tubercles in C. bitonium. Description. —In previous literature, described as smallbodied (‘‘diminutive’’) and short-limbed, with unequal inner and outer metatarsal tubercle sizes; distinct subarticular tubercles; barely visible supernumerary tubercles; no vomerine odontophores (Taylor 1936); rounded canthus; two palmar tubercles (Taylor 1942). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Holotype (UIMNH 16125) ~ 18 mm SVL. Among select specimens that we examined, we observed short, rounded snout (naris–snout ¼ 0.78 mm, SD ¼ 0.14, n ¼ 29; 5.8% SVL); long eye–nostril distance (1.19 mm, SD ¼ 0.24, n ¼ 29; 8.9% SVL); some specimens with single palmar tubercle; relative finger lengths III> IV> II ¼ I; relative toe lengths IV Distribution. —Widely distributed throughout lowland to intermediate localities in the states of Chiapas, Puebla, Oaxaca, and Veracruz (also possibly Tabasco, Mexico, and western Guatemala) from near sea level to 2000 m (Fig. 6). Canseco Márquez and Gutiérrez Mayén (2010) report C. pygmaeus occurs in the forests adjacent to the Tehuacán– Cuicatlán Valley in Puebla and Oaxaca. The western range edge of C. pygmaeus is uncertain; we examined one specimen (UTA A-66131) a male with pigmented testes, from San Vicente de Benitez (17.290618N, 100.279558W, 951 m), Guerrero, collected 17 June 2004, that appears referable to C. pygmaeus. Diet. —One male specimen (UTA A-64263, determined by the presence of pigmented testes), was found to contain ants (Formicidae) in its stomach. Phylogenetics. — Craugastor pygmaeus was inferred to be the sister taxon of C. bitonium with high support in the concatenated analyses (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). This sister relationship was also recovered in both mtDNA and nDNA analyses, although with lower support in the nDNAonly analyses (ML ¼ 54, BAYES ¼ 0.67; Figs. 4 and 5). Craugastor pygmaeus is separated from C. bitonium by a P - distance of 4.7% (Table 4). Remarks. —The skull of C. pygmaeus is similar to C. hobartsmithi and C. montanus, with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. Despite how common this species is in most museum collections, we know very little about its natural history, including reproductive behavior, call, and diet. This species likely co-occurs with C. candelariensis, C. polaclavus and C. portilloensis in southcentral Oaxaca and (possibly) C. bitonium in central Guerrero. It may overlap with (1) C. montanus in Chiapas, (2) C. mexicanus at intermediate elevations of the Sierra Madre del Sur and Sierra Madre Oriental (Fig. 7), and (3) C. hobartsmithi in central and western Guerrero. Males likely have larger tympana than do females (Figs. 13 and 33)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 34-35, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1936 [1937]. New species of amphibia from Mexico. Transactions of the Kansas Academy of Science 39: 349 - 363.","Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571.","Taylor, E. H. 1942. New tailless Amphibia from Mexico. University of Kansas Science Bulletin 28: 67 - 89.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Canseco Marquez, L., and M. G. Gutierrez Mayen. 2010. Anfibios y Reptiles del Valle Tehuacan-Cuicatlan. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Mexico."]}

Craugastor rubinus sp. nov. Holotype. — UTA A-62345 (field ID: JAC 30720; Fig. 34A), male collected by J.W. Streicher, C.L. Cox, C.M. Sheehy, III, R.U. Tovar, and M.J. Ingrasci on the road between Talpa de Allende and El Cuale, Jalisco, Mexico, 20.377078N, 105.047938W, 1771 m, 8 July 2009. Paratypes (2). — UTA A- 62347 (Fig. 34 C) and MZFC-HE-35616 (formerly UTA A- 62346; Fig. 34 B), same collection data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size; (2) reduced ossification of the skeleton in adults relative to other members of series, lacking ossification of any skeletal elements beyond Stage 2 (Table 3); (3) presence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred, with snout–nostril–canthal–supratympanic stripe, sometimes broken canthaly and postocularly; (8) one postrictal tubercle; (9) gular region with pale spotting; (10) dorsal surface blotched or unicolored pale; diffuse interorbital bar, small suprascapular spots, sometimes two dark rump spots; (11) middorsal ridge present; (12) dorsum smooth with no tubercles; (13) body flank dark supratympanic stripe extending toward lower midflank, broken, paler toward groin, smooth to finely shagreened; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches nostril; (16) outer tarsal ridge with 0–4 extremely small, flat, and round tubercles, no raised fringe; (17) finger and toe pads round, finger tips not or just barely expanded, toe tips slightly lanceolate and barely expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor rubinus can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. rubinus). It can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, and C. saltator by the presence of vomerine odontophores (absent in C. rubinus). It can be differentiated from C. bitonium and C. pygmaeus by the absence of a posterolateral projection of the frontoparietal (present in C. rubinus). Description of holotype. — Holotype small male (SVL ¼ 12.6 mm); snout rounded and short (0.8 mm naris–snout; 7% SVL); short eye–nostril distance (1.3 mm; 10% SVL); tympanum 1.2 mm (9.1% SVL); no supratympanic fold or shoulder tubercle; finger length formula III C. mexicanus series (Fig. 35); grey in preservative; many bands present on arms and legs. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Variations in paratypes. —Body sizes (SVL) 10.8 mm (MZFC-HE-35616), 11.5 mm (UTA A-62347); eye–nostril distance both 10% SVL; tympanic ratios both 9%; dorsal color pattern mottled (MZFC-HE-35616; Fig. 34B) or with dark stripe extending from snout to flank (UTA A-62347, Fig. 34C). Distribution. —Known only from the type locality in Jalisco (Talpa de Allende), habitat of pine–oak forest at the southern extent of the Sierra Madre Occidental. Etymology. —The specific epithet is derived from the Latin rubinus, which means ruby. This name is inspired by the garnet mines found near the type locality of Talpa de Allende in the Sierra Madre Occidental of Jalisco. Phylogenetics. — Craugastor rubinus was strongly supported as monophyletic in the concatenated analysis (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). The new species is also strongly supported as the sister taxon of C. cf. hobartsmithi (ML ¼ 100; BAYES ¼ 1.0). Craugastor rubinus is separated from C. cf. hobartsmithi by a P -distance of 3.4% (Table 4). Remarks. —The skull of C. rubinus is similar to that of C. hobartsmithi, C. montanus, and C. pygmaeus with more posteriorly placed anterior suture of the frontoparietal and prootic than in other species. The type series was collected from the leaf litter surrounding a mountain stream. The male holotype has slightly pigmented testes. Afemale paratype, UTA A-62345, has large, pigmented ovaries. This species may co-occur with C. hobartsmithi in Jalisco. Despite the small genetic distances separating C. rubinus and C. cf. hobartsmithi , there are many skeletal and morphological differences between these two species, including relative metatarsal tubercle sizes, condition of the posterolateral projection of frontoparietal, and relative snout length (Table 6)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 35-37, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Craugastor hobartsmithi (Taylor 1936) Eleutherodactylus hobartsmithi Taylor 1936:355. Holotype male (FMNH 100114) from ‘‘Uruapan, Michoacán, Mexico.’’ [Examined]. Microbatrachylus hobartsmithi (Taylor): Taylor 1940:501. Craugastor hobartsmithi (Taylor): Crawford and Smith 2005:536. Microbatrachylus pygmaeus Duellman 1961:34. [Misidentification]. Craugastor pygmaeus Ahumada-Carrillo et al. 2013:1338. [Misidentification]. Diagnosis. —Based on six specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) small adult sizes (males 11.3–15.2 mm, females ~ 16.7 mm; Table 5); (2) full ossification of most skeletal elements in adults, lacking ossification beyond Stage 4 (Table 3); (3) absence of vomerine odontophores; (4) absence of posterolateral projection of frontoparietal; (5) presence of a row of 4–6 rounded tubercles along outer edge, and 1–3 in mid upper eyelid; (6) supratympanic fold poorly developed; (7) face flank barred with or without distinctive canthal stripe; (8) one or two postrictal tubercles; (9) gular region peppered with melanocytes; (10) dorsal surfaces finely blotched, usually with dark interorbital bar and suprascapular ^-shape, some individuals with pale dorsal color and four stripes, paravertebral and lateral, originating at corners of eyes and ending above groin (lateral more prominent); (11) middorsal ridge (dark or background color); (12) mostly smooth dorsum or with just fine tubercles or folds toward back; (11) body flank darker anteriorly, around axilla, slightly tubercular; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches between anterior corner of eye and snout; (16) outer tarsal ridge with 4–6 rounded to slightly pointed tubercles, no raised fringe; (17) finger and toe tips round, finger tips slightly expanded, toe tips expanded; (18) similar sizes of inner and outer metatarsal tubercles. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Comparisons. — Craugastor hobartsmithi can be differentiated from C. bitonium, C. mexicanus, C. montanus, C. omiltemanus, C. polaclavus, C. pygmaeus, and C. rubinus by an inner metatarsal tubercle that is twice the size of the outer (these are similar sizes in C. hobartsmithi). It can be differentiated from C. candelariensis and C. saltator by the presence of vomerine odontophores (absent in C. hobartsmithi). It can be differentiated from C. cueyatl and C. portilloensis by the presence of posterolateral projections of the frontoparietal (absent in C. hobartsmithi). Description. —In previous literature, C. hobartsmithi has been described as small-bodied with pigmented gonads (Taylor 1936, 1940); presence of tubercles on the tarsus (Duellman 1961); two palmar tubercles (Lynch 1965); tarsus bearing a row of tubercles along its outer edge (Lynch 1970). Holotype (FMNH 100114) small male (13.5 mm); snout rounded and short (0.9 mm naris–snout; 6% SVL); short eye–nostril distance (1.18 mm; 8.7% SVL); tympanum 1.9 mm (14% SVL). We further examined two specimens of C. cf. hobartsmithi from coastal Michoacán (UTA A-66133–34; Fig. 25B and C) and noted the following characteristics: supratympanic fold terminating in two postrictal tubercles; finger length formula III ¼ II Distribution. — Craugastor hobartsmithi occurs in the pine–oak forest of Michoacán. Craugastor cf. hobartsmithi occurs throughout western Mexico in low to intermediate habitats of Jalisco, Nayarit, Michoacán, Guerrero, and Sinaloa (Fig. 8; Hardy and McDiarmid 1969). FloresCobarrubias et al. (2012) reported C. hobartsmithi from Hostotipaquillo, Jalisco. García and Ceballos (1994) reported C. hobartsmithi from coastal Jalisco. The records of C. pygmaeus reported in Duellman (1961) and AhumadaCarrillo et al. (2013) are all likely C. hobartsmithi or C. cf. hobartsmithi because our molecular results indicate that C. pygmaeus does not occur west of Guerrero. Similarly, many iNaturalist (https://www.inautralist.org, accessed June 2019) accounts of C. cf. hobartsmithi are listed under C. pygmaeus —these accounts also suggest that C. cf. hobartsmithi is much more widely distributed in western Mexico than museum collections indicate. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Phylogenetics. — Craugastor cf. hobartsmithi was found to be the sister taxon of C. rubinus with strong support (ML ¼ 100; BAYES ¼ 1.0) in both the concatenated and separate mtDNA and nDNA analyses (Figs. 3, 4, and 5). The pairwise P -distances between C. cf. hobartsmithi and C. rubinus is 3.4% (Table 4); this is the smallest genetic distance between any species of the C. mexicanus series, suggesting recent divergence. Remarks. —The skull of C. hobartsmithi is similar to C. montanus and C. pygmaeus, with more posteriorly placed anterior suture of frontoparietal and prootic than in other species. We tentatively referred several museum collections to C. hobartsmithi (Fig. 8) as C. cf. hobartsmithi , but these should be further examined. The specimens of C. pygmaeus reported by Duellman (1961) from Arteaga, Michoacán, are referred to C. cf. hobartsmithi because we examined several C. pygmaeus from Oaxaca with tubercles on the outer edge of the tarsus rendering Duellman’s (1961) apomorphic character for C. hobartsmithi unreliable. Craugastor hobartsmithi may co-occur with C. pygmaeus in southcentral Guerrero (Figs. 6 and 8). The tissues of C. cf. hobartsmithi used in our phylogenetic analysis originated from Colima. Although we lack a voucher specimen for the tissue, the collector of the tissue (J. Reyes-Velasco) provided us with photographs of C. cf. hobartsmithi from Montitlan, near where the tissue was collected (Fig. 25E and F). One female specimen of C. hobartsmithi from near Uruapan (UMMZ 94230) had several intradermal trombiculid mites on its venter (Fig. 15D)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 25-27, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1936 [1937]. New species of amphibia from Mexico. Transactions of the Kansas Academy of Science 39: 349 - 363.","Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Duellman, W. E. 1961. The amphibians of Michoacan, Mexico. Miscellaneous Publications of the University of Kansas Museum of Natural History 15: 1 - 148.","Ahumada-Carrillo, I. T., J. C. Arenas-Monroy, F. A. Fernandez-Nava, and O. Vazquez-Huizar. 2013. New distributional records for the pygmy robber frog Craugastor pygmaeus (Terrarana: Craugastoridae) in western Mexico. Revista Mexicana de Biodiversidad 84: 1338 - 1342.","Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA.","Lynch, J. D. 1970. Taxonomic notes on some Mexican frogs (Eleutherodactylus: Leptodactylidae). Herpetologica 26: 172 - 180.","Hardy, L., and R. McDiarmid. 1969. The amphibians and reptiles of Sinaloa, Mexico. University of Kansas Publications, Museum of Natural History 18: 39 - 252.","Garcia, A., and G. Ceballos. 1994. Guia de Campo de los Reptiles y Anfibios de la Costa de Jalisco, Mexico. Fundacion Ecologica de Cuixmala, A. C., Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico. [In Spanish.]"]}

Craugastor bitonium sp. nov. Holotype. — UTA A-64254 (field ID: JAC 22117; Fig. 22A), adult female from road between Yerba Santa and Yextla (HWY 196), Guerrero, Mexico, 17.52666°N, 99.9579°W, 2071 m, collected by J.A. Campbell and colleagues on 10 June 2002. Paratypes (5). —MZFC-HE-35600–01 and UTA A- 66117– 18 adult females, and UTA A-66119 adult male (Fig. 22B–D), all same collection data as holotype. Diagnosis. —A species of Craugastor distinguished by the following combination of characters: (1) small adult size (maximum SVL ¼ 16.7 mm); (2) full ossification of skeletal elements in adults; (3) absence of posterolateral projection of frontoparietal; (4) absence of vomerine odontophores; (5) presence of raised tubercles on eyelids; (6) supratympanic fold absent or poorly developed; (7) face flank barred or with supralabial pale stripe, and with or without dark canthal stripe; (8) single postrictal tubercle; (9) gular region peppered with melanocytes; (10) dorsal surface two-toned, usually with a dark suprascapular ^ shape, or almost unicolored; (11) pale or ground color middorsal ridge; (12) scattered fine tubercles on dorsum; (13) body flank barred darker anteriorly, slightly shagreened to smooth; (14) inguinal glands present and axillary glands absent in adults; (15) when leg adpressed to body, heel reaches middle of eye to slightly beyond snout; (16) outer tarsal ridge with 1–6 small mostly round tubercles, no raised fringe; (17) finger and toe tips round, slightly lanceolated, slightly expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor bitonium can be differentiated from C. mexicanus, C. montanus, C. omiltemanus, and C. saltator by their large adult body sizes of SVL> 20 mm (C. bitonium), the presence of vomerine odontophores (absent in C. bitonium), and the presence of three palmar tubercles (one palmar tubercle in C. bitonium). Craugastor bitonium can be differentiated from C. candelariensis, C. cueyatl, C. hobartsmithi, and C. portilloensis by the presence of metatarsal tubercles of similar sizes (different sizes in C. bitonium). Craugastor bitonium can be differentiated from C. rubinus by the presence of posterolateral projections of the frontoparietal (absent in C. bitonium). Craugastor bitonium can be differentiated from C. polaclavus by its shorter eye–nostril distance with an eye–nostril distance 9–10% SVL in C. bitonium and 10–13% SVL in C. polaclavus. Craugastor bitonium is most similar to C. pygmaeus (in morphology, osteology, and genetic distance), but may be differentiated from this taxon by the condition of the outer tarsal ridge; 1–6 small tubercles in C. bitonium versus no tubercles (¼ smooth) in C. pygmaeus. Holotype description. — Holotype small female with unpigmented developing ova (SVL ¼ 15.8 mm); snout rounded and short (0.9 mm naris–snout; 6% SVL); short eye–nostril distance (1.42 mm; 9% SVL); tympanum 1.4 mm (8.9% SVL); small supratympanic fold terminating in shoulder tubercle; finger length formula III Variations in paratypes. —Body sizes (SVL) 15.8 mm (MZFC-HE-35600), 15.2 mm (MZFC-HE-35601), 16.9 mm (UTA A- 66117), 16.7 mm (UTA A- 66118), 12.3 mm (UTA A- 66119); eye–nostril distance 9–10% SVL; tympanic ratios 7– 9%; dorsal color patterns variable, often with two distinctive patches of differing ground coloration ranging from orange to tan. Etymology. —The specific epithet is a combination of the Latin prefix bi- meaning two and tonium meaning tone. It is a reference to the two distinctive patches of color found on the holotype and several paratypes that create the appearance of a ‘‘two-tone’’ dorsal coloration. Distribution. —This species is known only from the Sierra Madre del Sur of central Guerrero (~ 2071 m). The closest named places to the type locality are Izotepec to the north and Los Bajos to the southwest. The habitat at the type locality is montane pine–oak forest. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Diet. —CT-scan of the holotype revealed the presence of a small millipede (Diplopoda) in the stomach. We also noted the presence of a small red ant (Formicidae) in the mouth of the holotype. Phylogenetics. — Craugastor bitonium was inferred to be the sister taxon of C. pygmaeus, with high support in the concatenated analyses (ML ¼ 99; BAYES ¼ 1.0; Fig. 3). This sister relationship was also recovered in both mtDNA and nDNA analyses, although with lower support in the nDNAonly analyses (ML ¼ 54, BAYES ¼ 0.67; Figs. 4 and 5). Craugastor bitonium is separated from C. pygmaeus by a P - distance of 4.7% (Table 4). Remarks. —The skull of C. bitonium is similar to C. hobartsmithi, C. montanus, and C. pygmaeus. Craugastor bitonium displays a developmental pattern similar to C. pygmaeus (with high levels of ossification at small sizes), suggesting small adult body sizes (Fig. 11). This species likely co-occurs with C. pygmaeus, C. omiltemanus, and C. saltator in central Guerrero (Figs. 6 and 8). The specimen UTA A-66132 is referred with some hesitation because it was collected from a lower elevation than the type locality and has a Finger Ilength nearing C. pygmaeus (which also occurs in Guerrero). Six of the specimens were adult females containing unpigmented ovaries with yolked eggs and thick oviducts, the seventh (UTA A-66119; Fig. 22C, far left) was an adult male with pigmented testes., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 13-16, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587

Jameson, Tom J.M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J., Smith, Eric N. (2022): Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species. Herpetological Monographs 36 (1): 1-48, DOI: 10.1655/0733-1347-36.1.1, URL: http://dx.doi.org/10.1655/0733-1347-36.1.1

Craugastor omiltemanus (Günther 1900a) Syrrhaphus omiltemanus Günther 1900a:213. Holotype unsexed (BMNH 1901.12.19.7) from ‘‘Omilteme, Guerrero, Mexico.’’ [Examined]. Hylodes calcitrans Günther 1900b:229. Holotype unsexed (BMNH 1901.12.19.25) from ‘‘Omilteme, Guerrero, Mexico.’’ [Examined]. Eleutherodactylus omiltemanus (Günther): Lynch 1970:175. Craugastor omiltemanus (Günther): Crawford and Smith 2005:536. Diagnosis. —Based on Hylodes calcitrans type series of 25 individuals and 1 additional specimen. Aspecies of Craugastor distinguished by the following combination of characters: (1) large adult size (maximum SVL ¼ 38.8 mm); (2) reduced ossification of skeleton in adults relative to other members of series, lacking ossification of any elements beyond Stage 2 (Table 3) except for the sphenethmoid; (3) presence of posterolateral projection of frontoparietal; (4) presence of vomerine odontophore; (5) presence or absence of raised tubercles on eyelids, smooth, six round and only slightly protruding tubercles, sometimes a few aligned at the outer edge; (6) supratympanic fold developed; (7) face flank barred or dark; canthus dark, pale, or spotted; canthus with or without a stripe (complete or broken); (8) one or two postrictal tubercles; (9) gular region from evenly scattered fine pigmentation, to densely pigmented with a mid-pale stripe; (10) dorsal surface unicolored pale brown or grey; unicolored to lightly spotted above the scapular and/or rump areas, or tubercles and ridges (if present), interorbital bar; (11) with or without a middorsal ridge; (12) dorsum smooth with only few fine tubercles toward flanks and urostyle or with ridges forming hourglass patterns and medium tuberculation; (13) body flank darker anteriorly (postaxillary) due to posterolateral expansion of supratympanic stripe; sometimes with contrasting white and dark blotching inguinal area, otherwise pale colored; shagreened; (14) inguinal gland present and axillary gland present in adults; (15) when leg adpressed to body, heel reaches middle of eye to mid-canthal area; (16) outer tarsal ridge 0–5 rounded and only slightly raised tubercles, no raised fringe; (17) finger and toe pads round, fingertips slightly or not expanded, toe tips expanded; (18) inner metatarsal tubercle larger than outer metatarsaltubercle. Comparisons. — Craugastor omiltemanus can be differentiated from all other members of the C. mexicanus series by the combination of rough (and often raised) areolate skin on its venter and a massive inner metatarsal tubercle (Figs. 21G and 29; Lynch 1970, 2000). Description. —In previous literature described as largebodied, short-legged, with Toe III C. omiltemanus, BMNH 1901.12.19.7 (reregistered as BMNH 1947.2.16.62; Fig. 1 J) and BMNH 1901.12.19.8 (reregistered as BMNH 1947.2.16.63). We designate the former as the lectotype and the latter as the paralectotype of this species. Both specimens appear to have partially desiccated at some point in their history, whereas the type series of Hylodes calcitrans is in much better condition (Fig. 1K; Fig. 23A and B). We designate BMNH 1901.12.19.29 (reregistered as BMNH 1947.2.16.47) as the lectotype of H. calcitrans. Relative finger lengthsof the typesare III> IV> II> Iandrelative toe lengths are IV Distribution. —Prior to our study, C. omiltemanus was known only from Guerrero (Günther 1900a; Crawford and Smith 2005). Our discovery of a specimen from Oaxaca (UTA A-64264) that is both phylogenetically and morphologically assignable with C. omiltemanus extends the distribution of this species eastward. This species occurs in high-elevation pine–oak forest habitats of the Sierra Madre del Sur in the states of Guerrero and Oaxaca (Fig. 8; Fig. 30E and F). Phylogenetics. —In all analyses, C. omiltemanus was found to be monophyletic with strong support (ML> 90; BAYES> 0.90; Figs. 3 and 4). In the concatenated analysis C. omiltemanus was found to be the sister taxon of C. saltator, although with limited support (ML ¼ 44; BAYES ¼ 0.74; Fig. 3). There was less support for the sister relationship with C. saltator in the mtDNA-only analysis (ML ¼ 33; BAYES ¼ 0.82; Fig. 4). In the nDNA-only analysis (which did not include C. saltator), C. omiltemanus was placed in basal polytomy with two other clades: (1) C. mexicanus and (2) all other species of the C. mexicanus series (Fig. 5). In terms of genetic distances (Table 4), C. omiltemanus was most similar to C. mexicanus (4.9%), followed by similarity to C. saltator (5.6%). Remarks. —The skull of C. omiltemanus is similar to C. mexicanus and C. saltator with a more anteriorly placed anterior suture of frontoparietal and prootic than that in other species. Skull examination also was conducted on a smaller subadult individual (UTA A-55240, SVL ¼ 16.5 mm) that had only Stage 1 of the ontogenetic sequence complete and only two features of Stage 2 present. Unlike the adult specimen, the nasals are completely unossified and the frontoparietal greatly reduced, and while vomerine odontophores are present the posterolateral projection of the frontoparietal is absent. Craugastor omiltemanus likely shares a similar distribution with other Craugastor species endemic to the Sierra Madre del Sur (e.g., C. uno; Streicher et al. 2011). Lynch (2000) reports that C. omiltemanus has white testes, but we observed pigmented testes in this species (Table 5). Throughout its range, C. omiltemanus likely co-occurs with C. bitonium, C. mexicanus, C. pygmaeus, and C. saltator. Male C. omiltemanus have significantly larger tympana than do female C. omiltemanus (Lynch 2000; this study)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 30-31, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Gunther, A. C. L. G. 1900 a. Reptilia and Batrachia, part 155. Pp. 213 - 220 in Biologia Centrali Americana 7 (O. Salvin and F. D. Godman, eds.). R. H. Porter and Dulau & Co., UK.","Gunther, A. C. L. G. 1900 b. Reptilia and Batrachia, part 157. Pp. 229 - 236 in Biologia Centrali Americana 7 (O. Salvin and F. D. Godman, eds.). R. H. Porter and Dulau & Co., UK.","Lynch, J. D. 1970. Taxonomic notes on some Mexican frogs (Eleutherodactylus: Leptodactylidae). Herpetologica 26: 172 - 180.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94.","Streicher, J. W., J. M. Meik, E. N. Smith, and J. A. Campbell. 2011. Low levels of genetic diversity among morphologically distinct populations of an enigmatic montane frog from Mexico (Craugastor uno: Craugastoridae). Amphibia-Reptilia 32: 125 - 131."]}

Craugastor saltator (Taylor 1941) Eleutherodactylus saltator Taylor 1941:89. Holotype female (FMNH 100116) from ‘‘Omilteme, Guerrero, Mexico.’’ [Examined]. Eleutherodactylus mexicanus: Lynch 2000:134. [Misidentification]. Craugastor saltator (Taylor): Crawford and Smith 2005:536. Diagnosis. —Based on holotype and three additional specimens. Aspecies of Craugastor distinguished by the following combination of characters: (1) large adult size (maximum SVL ¼ 44 mm); (2) full ossification of skeleton in adults but under a different ontogenetic sequence than other members of the series (Table 3), where the sphenethmoid, humeral interior epiphyses, and tibiofibular epiphyses do not ossify during stage 3, the fontopareital-prootic suture does not offset posteriorly during stage 4, and the epicorocoids do not ossify during stage 6; (3) presence of posterolateral projection of the frontoparietal; (4) presence of vomerine odontophores; (5) presence or absence of raised tubercles on eyelids; (6) supratympanic fold developed; (7) face flank, labium barred or dark with a cream stripe above; canthal stripe complete or broken; (8) one or two postrictal tubercles; (9) gular region with trace of mid-pale stripe; (10) dorsal surface unicolored, blotched, or with wide middorsal stripe bordered by cream-colored stripes, dark interorbital bar, sometimes with small suprascapular and/or rump spots; (11) middorsal ridge present; (12) dorsum smooth or slightly tuberculate; (13) body flank unicolored, rarely supratympanic stripe extending to area behind insertion of arm, making anterior area darker; finely shagreened; (14) inguinal gland present and axillary gland present in adults; (15) when leg adpressed to body, heel reaches beyond snout; (16) outer tarsal ridge with 0–5 extremely small, flat, and round tubercles, no raised fringe or ridge; (17) finger and toe pads round and expanded; (18) inner metatarsal tubercle larger than outer metatarsal tubercle. Comparisons. — Craugastor saltator can be differentiated from C. bitonium, C. cueyatl, C. hobartsmithi, C. pygmaeus, and C. rubinus by the absence of vomerine odontophores (present in C. saltator). It can be differentiated from C. candelariensis and C. portilloensis by equal sizes of the inner and outer metatarsal tubercles (unequal sizes in C. saltator). It can be differentiated from C. omiltemanus by ventral skin texture in life; smooth to granular in C. saltator versus areolate in C. omiltemanus. It can be differentiated from C. montanus and C. polaclavus by shorter relative leg sizes with a crus ratio of 50–58% SVL (long relative leg sizes of 62–73% SVL in C. saltator). Craugastor saltator is most similar to C. mexicanus; however, they do not overlap in geographic range (see C. mexicanus account for additional information). Description. —Detailed description in Taylor (1942). Described as large-bodied, long-legged, with pigmented testes, unequal inner and outer metatarsal tubercle sizes, large vomerine odontophores, and generally smooth dorsal skin (Taylor 1941); with less population-level chromatic variation than its relative C. mexicanus (Lynch 2000). Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022 Holotype (FMNH 100166) large female (SVL ¼ 44 mm; Fig. 36) with relative finger lengths III> IV> II> Iand relative toe lengths IV> III> V> II> I. Asubadult individual used in the molecular analysis, UTA A- 66120 (Fig. 36) had the following characteristics: SVL ¼ 14.5 mm; tympanum width ¼ 1.2 mm; naris–snout length ¼ 0.7 mm (4.7% SVL); eye–nostril distance ¼ 1.5 mm (10.3% SVL); relative finger lengths III> IV> II> I; relative toe lengths IV> III> V> II> I; unlike adult specimens, inner metatarsal tubercle and outer metatarsal tubercle near equal in length. Distribution. —Known only from the high-elevation pine–oak forests of Guerrero in the Sierra Madre del Sur (Fig. 6). Phylogenetics. —We were only able to sequence mtDNA data for C. saltator, so only it was included in the concatenated and mtDNA analyses. In the concatenated analysis, it was recovered as the sister taxon of C. omiltemanus with limited support (ML ¼ 44; BAYES ¼ 0.74; Fig. 3). In the mtDNA-only analysis, support for this relationship was lower in the ML analysis but higher in the BAYES analysis (ML ¼ 33, BAYES ¼ 0.82; Fig. 4). In terms of genetic distances (Table 4), C. saltator was most similar to C. mexicanus (5.1%), followed by similarity to C. omiltemanus (5.6%). Remarks. —The skull of C. saltator is similar to that of C. mexicanus and C. omiltemanus, with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. Taylor (1941:91) makes multiple references to C. saltator and C. omiltemanus (as Eleutherodactylus calcitrans) being similar and states that these two species can be differentiated by the ‘‘very long limb, and the reduced inner metatarsaltubercle [of C. saltator].’’ Lynch (2000) noted that C. saltator is actually far more similar to C. mexicanus from adjacent Oaxaca and subsequently synonymized C. saltator with C. mexicanus (see Methods, Taxonomic History). Despite the finding of Crawford and Smith (2005), which were used to revalidate C. saltator, at one point in our study one of us (JWS) was convinced by Lynch’s (2000) argument that C. saltator should be a junior synonym of C. mexicanus. The basis for this suspicion was (1) a specimen collected by J.D. Godman from Omilteme, Guerrero (the type locality) identified by J.D. Lynch as C. saltator on 13 January 1972 (BMNH 1901.12.19.24) is similar in morphology to C. mexicanus; (2) the nDNA analysis of Crawford and Smith (2005) found C. mexicanus þ C. saltator to be monophyletic; and, (3) we did not observe range overlap between C. mexicanus and C. saltator (Fig. 6), which may be consistent with a single large-bodied, long-legged species inhabiting the Sierra Madre del Sur. However, as in Crawford and Smith (2005), our mtDNA phylogenetic results did not recover C. mexicanus þ C. saltator as monophyletic (Fig. 3) and the specimens of C. saltator we examined had on average larger body sizes and differing toe length formulae than did C. mexicanus (Figs. 10 and 11; Table 6). As such, we continue to recognize C. saltator as a distinct species pending further taxonomic investigation. Craugastor saltator likely co-occurs with C. bitonium, C. pygmaeus, and C. omiltemanus in the Sierra Madre del Sur of Guerrero. It may overlap with C. mexicanus in eastern Guerrero and western Oaxaca (Fig. 6)., Published as part of Jameson, Tom J. M., Streicher, Jeffrey W., Manuelli, Luigi, Head, Jason J. & Smith, Eric N., 2022, Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species, pp. 1-48 in Herpetological Monographs 36 (1) on pages 37-38, DOI: 10.1655/0733-1347-36.1.1, http://zenodo.org/record/6518587, {"references":["Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94.","Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156.","Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555.","Taylor, E. H. 1942. New tailless Amphibia from Mexico. University of Kansas Science Bulletin 28: 67 - 89."]}

Knowledge of the biodiversity of amphibians and reptiles is essential in developing conservation for these taxa. We collected data from visual encounters and bibliographic review to determine the diversity of amphibians and reptiles in different habitats in the Putumayo–Napo–Amazonas interfluvium. This interfluvium presents 140 species of amphibians and 108 reptiles. The hill forest had the highest species richness, with 58 species of amphibians and 40 species of reptiles. The richest families were Hylidae and Colubridae. The Hilids presented a greater number of species in flooded ecosystems, while the Craugastoridae and Bufonidae families had greater richness in non-flooded ones. The Colubridae family was present in both ecosystems, although with a greater presence in flooded habitats. A community similarity analysis showed two groups: one formed by the amphibians of the high terrace, middle terrace and hill forest forests, and a second group from the low terrace, palm swamp, and peatland pole forest. In the reptiles, two groups were also formed: one by the reptiles of the palm swamp, hill forest and high terrace, and the second group made up of the reptiles of the low terrace forest, middle terrace and peatland pole forest. This diversity is threatened by deforestation, agriculture, caiman hunting and turtle egg collection, if intensified, could cause drastic population declines in these species.

Studies on spatial occupation are fundamental to understand amphibian communities. The aim of this study was to record information on the spatial distribution of anurans in the Tejipió forest, state of Pernambuco, Northeastern Brazil. Fieldwork was carried out weekly between October 2011 and April 2012, with daytime and night-time excursions for time-constrained active searching, in forested and open areas, military construction area and water bodies. Pitfall traps and accidental sightings were also used as alternative collection methods. Data were used to calculate richness, rarefaction curves and richness estimators. A total of 21 species were recorded, distributed in six families: Bufonidae (2 spp.); Craugastoridae (1 sp.); Hylidae (8 spp.); Leptodactylidae (8 spp.); Microhylidae (1 sp.) and Phyllomedusidae (1 sp.). Only the species Rhinella jimi was found occupying all sampled habitats in the research area. Adenomera hylaedactyla and Pristimantis ramagii deserve special care in the area because they are specialists, occupying a smaller number of habitats and microhabitats. The community of anurans of the Tejipió forest is similar to those recorded in other areas of the Atlantic Forest at the Pernambuco State, and its knowledge is essential as a basis for conservation of the area. The gradual recovery of this Atlantic Forest remnant would favor the recolonization of fauna and flora and the conservation of local biodiversity.

Una de las principales hipótesis para explicar el origen de la diversidad amazónica invoca el efecto de barrera de los ríos para explicar los patrones de diversidad. Esa hipótesis propone que algunos ríos pueden separar poblaciones continuas conduciendo a la diferenciación y la especiación. En ese sentido nos propusimos estudiar la estructura genética de Pristimantis latro, una especie recién descrita que habita una región bajo fuerte presión antrópica producto de la ganadería expansiva, la minería ilegal y la construcción de represas hidroeléctricas. El ADN se extrajo de 52 individuos de P. latro y se amplificó mediante reacción en cadena de la polimerasa (PCR) usando los marcadores mitocondriales 16S rRNA y COI. Para inferir el tiempo de divergencia entre las localidades de P. latro, construimos un árbol de especies e hicimos análisis de varianza molecular (AMOVA) para inferir la diferenciación genética entre y dentro de las poblaciones de P. latro. Encontramos que P. latro presenta una estructuración genética en las poblaciones de las márgenes derecha e izquierda del río Xingu y dentro de las regiones interfluviales deTapajós-Xingu y Xingu-Tocantins. Asimismo, el tiempo de divergencia entre las poblaciones de las márgenes derecha e izquierda del río Xingu aconteció hace aproximadamente 380.000 años. El patrón de estructura genética que se encontró se corresponde con el indicado en artículos recientes para el género Pristimantis, el cual revela que las especies sin renacuajos exhiben una estructura genética que responde a la hipótesis de los ríos como barreras. One of the main hypotheses to explain the origin of Amazonian diversity is the barrier effect of the rivers known as the riverine-barrier hypothesis, which suggests that riverine barriers isolated once continuous populations leading to differentiation and speciation. In this context, we studied the genetic structure of Pristimantis latro, a newly described species that inhabits a region under marked anthropic pressure due to expansive livestock, illegal mining, and hydroelectric dam construction. The DNA was extracted from 52 P. latro individuals and then amplified via polymerase chain reaction (PCR) using the mitochondrial 16S rRNA and COI markers. To infer the time of divergence between the P. latro localities, we built a species tree and performed an analysis of molecular variance (AMOVA) to infer the genetic differentiation between and within the P. latro populations. We found that P. latro has a marked genetic structure in the populations of the right and left margins of the Xingu River and within the Tapajós-Xingu and Xingu-Tocantins interfluvial regions and that the time of divergence between the populations of the East and West banks of the Xingu River occurred approximately 380,000 years ago. This pattern of genetic structure corresponds to that reported in recent articles for the Pristimantis genus evidencing that species without tadpoles exhibit a genetic structure explained by the hypothesis of rivers as barriers.

A new species of Pristimantis (Craugastoridae, subgenus Pristimantis) is described from a relict and unexplored cloud forest in the western slope from Cordillera Oriental of the Colombian Andes. The specific name was chosen by consensus expert scientists and local people. Pristimantis chamezensissp. nov. is easily distinguished from congeneric species by having a gray iris with black reticulations in life, subconical tubercles on the upper eyelid, the chin edged with irregular, dark-brown blotches, and conical heel tubercles. The phylogenetic analyses suggest that the origin and radiation of its clade may have occurred in the highlands. With the description of P. chamezensissp. nov., we identify 14 species distributed throughout the eastern slope of the Andes that are associated with the Orinoco Basin.

Pristimantis bacchus is an endangered terrestrial frog, endemic to cloud forests of the northern part of the Cordillera Oriental in the Colombian Andes. The knowledge of its reproductive biology and microhabitat use can offer important information for future conservation programs. We evaluated the reproductive activity and microhabitat use for both sexes, and selection of calling sites by males during rainy and dry seasons. Individuals of all reproductive conditions were observed throughout the sampled months; however, we observed a higher abundance of frogs during the first rainy season, after the driest months of the year. Thus, this population seems to have continuous reproductive activity, with an apparent peak at the onset of the first rainy season of the year. There were differences in the microhabitat use between sexes, between adults and juveniles, and between non-reproductive and reproductive females. Males use a wider variety of substrates at higher perches, while females were mainly found on leaf litter and over leaves at low heights; however, ovigerous females mostly occupied substrates above 120 cm. Juveniles were less abundant than adults, and occupy low substrates, mostly at forest litter. Two selection models evaluated the preference for calling sites, related to physiological requirements of humidity and for vocal sound dispersion. Males preferred to vocalize on perches at heights among 41-120 cm and with a vegetal structure that maintains high humidity, avoiding desiccation, and reducing exposure to predators. Therefore, intrapopulation variation in microhabitat use in this species is related to age, sex, reproductive condition, and physiological requirements.

We describe a new glandular terrestrial frog of the Pristimantis (Anura: Craugastoridae) genus. The new species inhabits the high montane forests of the Cañar province in southern Ecuador. It differs from other terrestrial frogs by having a striking dark gray and red dorsal coloration, dorsal skin with a corrugated texture, a small tubercle on the eyelid and heel, absence of vomerine odontophores, and large glandular patches on the supratympanic area, arms and limbs. Our molecular analyses show that the new species is in a clade with P. orcesi and P. erythros. Based on our results, we redefine the orcesi group and suggest that it contains only three species (P. orcesi, P. erythros, and the new species); the group is diagnosed mainly by exhibiting conspicuous dermal macroglands on the head, body and legs.

Pristimantis relictus sp. nov. Hylodes ramagii Rocha, 1948 Eleutherodactylus sp Lima-Verde & Cascon (1990) Eleutherodactylus cf. ramagii Borges-Nojosa & Lima (2001) Ischnocnema gr. ramagii Carnaval & Bates (2007) Pristimantis sp Loebmann & Haddad (2010) Pristimantis sp 2. Canedo & Haddad (2012) Pristimantis sp Roberto & Loebmann (2016) Pristimantis sp (cf. ramagii)— Castro et al. (2019) Pristimantis sp (Northern Ceará clade)— Trevisan et al. (2020) Holotype. URCA-H 13344 (adult male) collected at Parque Nacional de Ubajara, municipality of Ubajara, state of Ceará, Brazil (03°50’18.5”S, 40°54’37.9” W; 833 m above sea level), on April 10, 2013 by Igor J. Roberto (Figure 2). Paratypes. 17 males, 33 females and 3 juveniles of undetermined sex: URCA-H 13343 (adult male) and 13342 (adult female) collected with the holotype. CFBH 20304 (adult male), CFBH 20306–08, 20310–11, 20313 (adult females), collected at Parque Nacional de Ubajara, municipality of Ubajara, state of Ceará, Brazil (03°50’31.57” S, 40°53’56” W, 850 m a.s.l.), on June 29, 2008 by Daniel Loebmann. CFBH 15899 –00, 15904 (adult males), collected at Mata das Bromélias, Parque Nacional de Ubajara, municipality of Ubajara, state of Ceará, Brazil (03°51’16” S, 40°55’16” W, 820 m a.s.l), on March 24, 2007 by Daniel Loebmann. URCA-H 1546 (adult male), 1543–45, 1547 (adult females) collected at municipality of Guaramiranga, state of Ceará, Brazil (04°15’21”S, 38°58’17” W, 830 m a.s.l), on February 12, 2012 by Robson W. Ávila. CFBH 20316–17, 25415–16, 25419 (adult females), CFBH 25417–18, 25420–21 (adult males), collected at Parque das Trilhas, municipality of Guaramiranga, state of Ceará, Brazil (04°15’48”S, 38°55’59” W, 853 m asl), on January 23, 2009 by Daniel Loebmann. URCA-H 6988 (adult male), 6979–80, 6984, 6987 (adult females) collected on February 3–5, 2013. URCA-H 9214 (adult male) collected on April 7, 2014 at Área de Proteção Ambiental Bica do Ipu, municipality of Ipu, state of Ceará, Brazil (41°7’21.78”S, 40°42’47.69”W), by Robson W. Ávila. URCA-H 2373 (adult female) collected on March 23, 2011 by Igor J. Roberto, 9828–30 (adult females) and 9633, 9837–38 (juveniles) collected by Herivelto F. Oliveira on August 14–20, 2014 at Serra de Maranguape, municipality of Maranguape, state of Ceará, Brazil (35°3’59.23”S, 38°43’0.32”W, 800 m a.s.l). CFBH 24531 (adult female), collected at Serra de Maranguape, municipality of Maranguape, state of Ceará, Brazil (35°3’59.23”S, 38°43’0.32”W, 770 m a.s.l), on December 11, 2008 by Daniel Loebmann. CFBH 25887 (adult female), CFBH 25888 (adult male) collected at Serra da Aratanha, municipality of Pacatuba, state of Ceará, Brazil (03°58’60”S, 38°38’00” W, 700 m a.s.l), on March 29, 2010 by Daniel Loebmann. CFBH 25406–12 (adult females) collected at Fazenda Gameleira, municipality of Tianguá, state of Ceará, Brazil (03°43’16”S, 40°55’46” W 800 m a.s.l), on October 28, 2008 by Daniel Loebmann. CFBH 25413–14 (adult males) collected at Estrada do Retiro, municipality of Uruburetama, state of Ceará, Brazil (03°35’53.4”S, 39°34’50” W), on January 23, 2009 by Daniel Loebmann. URCA-H 16276–77 (adult males) collected at Serra da Meruoca, municipality of Meruoca, state of Ceará, Brazil (03°37’03.3”S, 40°21’46.7” W, 700 m a.s.l), on February 20, 2019 by Kassio C. Araújo. Referred specimens: URCA-H 9835-9836 (juveniles) collected at Serra de Maranguape, municipality of Maranguape, state of Ceará, Brazil; URCA-H 6981-6983, 12287 collected at municipality of Ipu, state of Ceará; MNRJ 55882-55883 collected at Serra da Aratanha, municipality of Pacatuba, state of Ceará, Brazil; MNRJ 55884 collected at Serra da Ubatuba, municipality of Granja, state of Ceará, Brazil; CHFURG 2119-23 collected at Sítio São José, municipality of Guaramiranga, state of Ceará, Brazil. Definition. Pristimantis relictus sp. nov. is diagnosed by the following combination of characters: (1) dorsal skin shagreen with small scattered tubercles; (2) ventral skin areolate; (3) dorsolateral fold absent; (4) discoidal fold present; (5) tarsal fold present; (6) lateral fringes absent on fingers, narrow on toes; (7) vocal slits present in males; (8) advertisement call composed of 1–8 pulsed notes (2–5 pulses per note), note rate of 0.5–2.2 notes per second, call duration 20–660 ms, fundamental frequency located at the first energy band between 1895–2153 Hz and dominant frequency located at the second energy band between 3617–4220 Hz; (9) in life dorsum coloration dark brown to yellowish-brown, with presence of dark brown chevrons and black spots in some individuals, transversal brown bars on legs and arms; (10) immaculate cream-colored pattern of concealed surfaces of thighs; (11) presence of a dark brown stripe from tip of snout to eye, including inter-nostril area, and a dark brown interorbital bar; (12) supratympanic fold bold, black; (13) gular region yellowish; (14) iris color cooper with brown reticulations. Morphological comparisons with other species within the Pristimantis conspicillatus species group. The shagreen texture of the dorsal skin differentiates Pristimantis relictus sp. nov. from P. avicuporum (Duellman & Pramuk, 1999), P. metabates (Duellman & Pramuk, 1999), P. johannesdei (Rivero & Serna, 1988), P. phalaroinguinis (Duellman & Lehr, 2007), and P. pictus (all have smooth dorsal skin). Nineteen species have smooth ventral skin, differing from the areolate ventral skin of P. relictus sp. nov.: P. achatinus (Boulenger, 1898), P. buccinator (Rodriguez, 1994), P. charlottevillensis (Kaiser, Dwyer, Feichtinger & Schmid, 1995), P. chiastonotus (Lynch & Hoogmoed 1977), P. citriogaster (Duellman, 1992), P. condor (Lynch & Duellman, 1980), P. conspicillatus (Günther, 1858), P. fenestratus (Steindachner 1864), P. gaigei, P. gutturalis, P. johannesdei P. latro, P. lymani (Barbour & Noble, 1920), P. malkini (Lynch, 1980), P. metabates, P. peruvianus (Melin, 1941), P. phalaroinguinis, P. samaipatae (Köhler & Jungfer 1995), and P. vilarsi (Melin, 1941). The areolate ventral skin also distinguishes P. relictus sp. nov. from P. dundeei (Heyer & Muñoz, 1999), P. giorgii, P. iiap Padial, Gagliardi-Urrutia, Chaparro & Gutiérrez, 2016, P. incertus, and P. ventrigranulosus Maciel, Vaz-Silva, Oliveira & Padial, 2012 (ventral skin granular or slightly granular). The absence of dorsolateral folds distinguishes P. relictus sp. nov from Pristimantis adiastolus (Duellman & Hedges, 2007), P. ardilae, P. avicuporum, P. bipunctatus (Duellman & Hedges, 2005), P. buccinator, P. chiastonotus, P. conspicillatus, P. iiap, P. latro, P. malkini, P. meridionalis (Lehr & Duellman, 2007), P. peruvianus, P. pluvian, P. skydmainos, and P. zeuctotylus (presence of dorsolateral folds) The subovoid snout in dorsal view of P. relictus sp. nov. distinguishes it from P. carranguerorum (Lynch, 1994) (obtuse) and P. medemi (Lynch, 1994), P. latro, and P. zeuctotylus (subacuminate). The immaculate cream-colored pattern of concealed surfaces of thighs differentiates P. relictus sp. nov. from P. pluvian (red), P. bipunctatus, P. condor, P. conspicillatus, P. malkini, P. peruvianus, and P. pictus (with well-defined spots). In addition to the new species, five other species of Pristimantis are distributed in Northeast Brazil: P. moa, P. paulodutrai, P. ramagii, P. rupicola and P. vinhai (see Table 1). While Pristimantis relictus sp. nov. is distributed in highland marshes of Ceará, P. moa is known from transition areas between the Cerrado and Amazon Forest (Oliveira et al. 2020), P. rupicola occurs in Campos Rupestres of Chapada Diamantina (Taucce et al. 2020), and the remaining three species inhabit the Atlantic Forest domain (Trevisan et al. 2020). Besides geographic distribution, P. relictus sp. nov. can be distinguished from P. moa by areolate ventral skin (smooth in P. moa), snout subovoid in dorsal view (truncate in P. moa), immaculate cream-colored pattern of concealed surfaces of thighs (strongly stained yellow on a dark background in P. moa), finger fringes absent (present in P. moa), and webbing on toes absent (present in P. moa) (Oliveira et al. 2020). The new species differs from P. rupicola by possessing shagreen dorsal skin (granular in P. rupicola), and snout subovoid in dorsal view (rounded or truncate in P. rupicola) (Taucce et al. 2020). The new species can be distinguished from the Atlantic Forest species as follows: from P. paulodutrai by the presence of nuptial pads (absent in P. paulodutrai), ventral skin areolate (granular in P. paulodutrai), and immaculate creamcolored pattern of concealed surfaces of thighs (with red blotches in P. paulodutrai) (Bokermann, 1975); from P. ramagii by snout subovoid in dorsal view (subacuminate in P. ramagii), presence of nuptial pads and toe fringes (absent in P. ramagii) (Boulenger, 1888); and from P. vinhai by ventral skin areolate (slightly granular in P. vinhai), snout subovoid in dorsal view (acuminate in P. vinhai) (Bokermann, 1975). Bioacustical comparisons with other species within the Pristimantis conspicillatus species group. The shorter note duration (20–50 ms) differentiates P. relictus sp. nov. from P. samaipatae (59–141 ms) and P. fenestratus (50–91 ms) (Padial & De La Riva 2009). The lower note rate (0.5–2.2 notes/s) distinguishes P. relictus from P. fenestratus (7.7–12.7), P. koehleri (11.8–17.3), P. samaipatae (2.7–14.9), P. dundeei (13.7–20.7) (Heyer & Muñoz 1999; Giaretta et al. 2018), and P. vilarsi (14) (Heyer & Barrio-Amorós 2009). The lower number of pulses per note of P. relictus sp. nov. (2–5) also differentiates the new species from P. fenestratus (9–17), P. samaipatae (11–23), P. latro (6–9), and P. pluvian (8–16). The higher fundamental frequency of P. relictus sp. nov. (1894.9–2153.3 Hz) distinguishes the new species from P. samaipatae (1535–1834 Hz), P. latro (1342–1448.6 Hz), P. moa (1320.8–1660.2 Hz), and P. giorgii (663.2–1872.3 Hz). The higher peak frequency of P. relictus sp. nov. (3617.6–4220.5 Hz) differentiates the new species from P. fenestratus (1710–3591 Hz), P. latro (2635.9–3272 Hz), P. moa (2657.1–3400 Hz), and P. pictus (2487.4–3272.2 Hz) (Padial & De La Riva 2009; Oliveira et al. 2017, 2020). Also, advertisement call differentiates P. relictus from P. rupicola. The new species has a higher peak of frequency (3617.6–4220.5 Hz), located in the second band of energy, [peak frequency (2410–3490 Hz) in the first band of energy of the call; Taucce et al. (2020)]. The advertisement call of P. relictus is very similar to that of P. ramagii but the species can be distinguished by the number of pulses per note of 2–5 (7–53 in P. ramagii; Octaven et al. 2017). Description of the holotype. Adult male (URCA-H 13344; Figure 2). Head longer than wide; head width 37% of SVL; head length 40% of SVL; snout subovoid in dorsal view, rounded in lateral view; nostrils not protuberant, directed anterolaterally; internarial distance less than eye-nostril distance; canthus rostralis angular, weakly concave, loreal region slightly concave; eye large, with horizontally elliptical pupil; eye diameter slightly larger than interorbital distance; cranial crests absent; upper eyelid smooth, 74% of eye diameter; tympanum large, with distinct rounded tympanic annulus; tympanic membrane present, visible; supratympanic fold present; labial bars absent; vocal sac subgular, small; tongue ovoid covering the entire floor of mouth, free and notched behind; vocal slits present, lateral to tongue; choana small, rounded; odontophores oblique and widely separated posterior to the choanae, each one bearing six teeth; skin on dorsum shagreen with scattered low tubercles; dorsolateral fold absent; skin on venter areolate, weakly spotted, discoidal fold present; hands large, 29% of SVL; relative lengths of fingers II Measurements of the holotype (mm). SVL 24.4, HL 9.7, HW 8.9, ED 3.4, SL 4.4, IND 2.0, END 2.8, IOD 3.3, UEW 2.5, TD 2.0, FAL 5.4, HAL 7.0, FIII 1.4, FIV 1.3, THL 12.4, TL 13.1, TAL 7.1, FL 10.7, and TIV 1.4. Color in preservative. Dorsal and lateral surfaces pale cream, snout light brown, brown interocular band in dorsal view, distinct dark brown stripe from tip of snout to posterior region of supratympanic fold. Irregular brown marks in lower orbital region. Two brown chevrons on dorsum with irregular brown dots in inguinal region, light brown stripes on dorsal surfaces of limbs. Ventral surface background pale cream in color. Advertisement call. The call of Pristimantis relictus sp. nov. is described based on three individuals (MNRJ 55582 and two unvouchered individuals) (see Table 2;). The vocalization is composed of 1–8 multi pulsed notes (2–5 pulses per note), with an ascendant amplitude modulation. The call duration ranges 0.02– 0.66 s (0.06 ± 0.08; n = 181), with intercall interval of 0.8– 9.6 s (2.12 ± 1.5; n = 74), note duration ranges 0.02– 0.05 s (0.03 ± 0.01; n = 150), with a note rate of 0.52–2.20 notes per second (Figure 3. The call has three visible bands. The fundamental frequency is in the first band, ranging 1894.9–2153.3 Hz, followed by a second band located in the dominant frequency, ranging 3617.6–4220.5 Hz, and a third weak band with a higher frequency, ranging 4565–6287.7 Hz. Variation among paratypes. Adult females (SVL 26.8–32.8 mm) are slightly larger than adult males (SVL 23.1–29.7 mm) (Table 3). Variation in coloration of fixed specimens is mostly related to the presence of chevrons on the dorsum (URCA-H 6980, 2373, 1544, 13342, 6988, and 9828). In life, the dorsal coloration is highly polymorphic: dark brown, reddish to yellowish-brown, or grey (Figure 4), with the presence of dark brown chevrons and black spots in some individuals. Other variations in coloration among paratypes are: brown bars on the leg and arm; dark brown stripe from the tip of the nose to the eye, including the internarial area; dark brown bar in the interorbital region; labial region yellowish to white; supratympanic fold bold black; gular region yellowish; and iris cooper-colored with brown reticulations. Geographic distribution. Pristimantis relictus is endemic to the “Brejos de Altitude” of the state of Ceará, Northeast Brazil. This species occurs in the Planalto da Ibiapaba (municipalities of Ipu, Tianguá, Ubajara, Granja, Viçosa do Ceará), Serra de Maranguape (municipality of Maranguape), Serra da Aratanha (municipality of Pacatuba), Serra de Baturité (municipalities of Guaramiranga, Pacoti, Aratuba, Mulungu), Serra da Uruburetama (Roberto & Loebmann 2016), and Serra da Meruoca (municipality of Meruoca) (Figure 5). Natural history. Pristimantis relictus occurs in dry forests and humid forests (Loebmann & Haddad 2010; Roberto & Loebmann 2016; Castro et al. 2019), from 100 to 900 m.a.s.l. The species has a prolonged breeding period throughout the raining season from December to April. Vocalization occurs from 5 pm throughout the night, with a peak of individuals vocalizing during 6–9 pm. Individuals were found vocalizing on leaves, tree trunks and twigs, mostly facing downwards. Amplexus is axillary, and the eggs are deposited on humid surfaces of rock and wooden debris. Mature females range 28–33 mm and 1.45–3.27 g (n = 13), and clutches have 21– 53 eggs (n = 13). We observed territorial behavior involving agonistic combat and the emission of territorial and release calls; unfortunately, these calls were not recorded. At Serra de Baturité mountain range we recorded the following species occurring in sympatry with P. relictus: Boana raniceps, Dendropsophus minusculus, D. minutus, D. tapacuarensis, Pithecopus gonzagai, Scinax tropicalia, Trachycephalus typhonius, Adelophryne baturitensis, Odontophrynus carvalhoi, Rhinella casconi, R. dapsilis, R. diptycha, R. granulosa, Adenomera juikitam, Leptodactylus macrosternum, L. natalensis, L. pustulatus, L. mystaceus and Physalaemus cuvieri. See Loebmann & Haddad (2010), and Castro et al. (2019)

Pristimantis relictus sp. nov. Hylodes ramagii Rocha, 1948 Eleutherodactylus sp Lima-Verde & Cascon (1990) Eleutherodactylus cf. ramagii Borges-Nojosa & Lima (2001) Ischnocnema gr. ramagii Carnaval & Bates (2007) Pristimantis sp Loebmann & Haddad (2010) Pristimantis sp 2. Canedo & Haddad (2012) Pristimantis sp Roberto & Loebmann (2016) Pristimantis sp (cf. ramagii)��� Castro et al. (2019) Pristimantis sp (Northern Cear�� clade)��� Trevisan et al. (2020) Holotype. URCA-H 13344 (adult male) collected at Parque Nacional de Ubajara, municipality of Ubajara, state of Cear��, Brazil (03��50���18.5���S, 40��54���37.9��� W; 833 m above sea level), on April 10, 2013 by Igor J. Roberto (Figure 2). Paratypes. 17 males, 33 females and 3 juveniles of undetermined sex: URCA-H 13343 (adult male) and 13342 (adult female) collected with the holotype. CFBH 20304 (adult male), CFBH 20306���08, 20310���11, 20313 (adult females), collected at Parque Nacional de Ubajara, municipality of Ubajara, state of Cear��, Brazil (03��50���31.57��� S, 40��53���56��� W, 850 m a.s.l.), on June 29, 2008 by Daniel Loebmann. CFBH 15899 ���00, 15904 (adult males), collected at Mata das Brom��lias, Parque Nacional de Ubajara, municipality of Ubajara, state of Cear��, Brazil (03��51���16��� S, 40��55���16��� W, 820 m a.s.l), on March 24, 2007 by Daniel Loebmann. URCA-H 1546 (adult male), 1543���45, 1547 (adult females) collected at municipality of Guaramiranga, state of Cear��, Brazil (04��15���21���S, 38��58���17��� W, 830 m a.s.l), on February 12, 2012 by Robson W. ��vila. CFBH 20316���17, 25415���16, 25419 (adult females), CFBH 25417���18, 25420���21 (adult males), collected at Parque das Trilhas, municipality of Guaramiranga, state of Cear��, Brazil (04��15���48���S, 38��55���59��� W, 853 m asl), on January 23, 2009 by Daniel Loebmann. URCA-H 6988 (adult male), 6979���80, 6984, 6987 (adult females) collected on February 3���5, 2013. URCA-H 9214 (adult male) collected on April 7, 2014 at ��rea de Prote����o Ambiental Bica do Ipu, municipality of Ipu, state of Cear��, Brazil (41��7���21.78���S, 40��42���47.69���W), by Robson W. ��vila. URCA-H 2373 (adult female) collected on March 23, 2011 by Igor J. Roberto, 9828���30 (adult females) and 9633, 9837���38 (juveniles) collected by Herivelto F. Oliveira on August 14���20, 2014 at Serra de Maranguape, municipality of Maranguape, state of Cear��, Brazil (35��3���59.23���S, 38��43���0.32���W, 800 m a.s.l). CFBH 24531 (adult female), collected at Serra de Maranguape, municipality of Maranguape, state of Cear��, Brazil (35��3���59.23���S, 38��43���0.32���W, 770 m a.s.l), on December 11, 2008 by Daniel Loebmann. CFBH 25887 (adult female), CFBH 25888 (adult male) collected at Serra da Aratanha, municipality of Pacatuba, state of Cear��, Brazil (03��58���60���S, 38��38���00��� W, 700 m a.s.l), on March 29, 2010 by Daniel Loebmann. CFBH 25406���12 (adult females) collected at Fazenda Gameleira, municipality of Tiangu��, state of Cear��, Brazil (03��43���16���S, 40��55���46��� W 800 m a.s.l), on October 28, 2008 by Daniel Loebmann. CFBH 25413���14 (adult males) collected at Estrada do Retiro, municipality of Uruburetama, state of Cear��, Brazil (03��35���53.4���S, 39��34���50��� W), on January 23, 2009 by Daniel Loebmann. URCA-H 16276���77 (adult males) collected at Serra da Meruoca, municipality of Meruoca, state of Cear��, Brazil (03��37���03.3���S, 40��21���46.7��� W, 700 m a.s.l), on February 20, 2019 by Kassio C. Ara��jo. Referred specimens: URCA-H 9835-9836 (juveniles) collected at Serra de Maranguape, municipality of Maranguape, state of Cear��, Brazil; URCA-H 6981-6983, 12287 collected at municipality of Ipu, state of Cear��; MNRJ 55882-55883 collected at Serra da Aratanha, municipality of Pacatuba, state of Cear��, Brazil; MNRJ 55884 collected at Serra da Ubatuba, municipality of Granja, state of Cear��, Brazil; CHFURG 2119-23 collected at S��tio S��o Jos��, municipality of Guaramiranga, state of Cear��, Brazil. Definition. Pristimantis relictus sp. nov. is diagnosed by the following combination of characters: (1) dorsal skin shagreen with small scattered tubercles; (2) ventral skin areolate; (3) dorsolateral fold absent; (4) discoidal fold present; (5) tarsal fold present; (6) lateral fringes absent on fingers, narrow on toes; (7) vocal slits present in males; (8) advertisement call composed of 1���8 pulsed notes (2���5 pulses per note), note rate of 0.5���2.2 notes per second, call duration 20���660 ms, fundamental frequency located at the first energy band between 1895���2153 Hz and dominant frequency located at the second energy band between 3617���4220 Hz; (9) in life dorsum coloration dark brown to yellowish-brown, with presence of dark brown chevrons and black spots in some individuals, transversal brown bars on legs and arms; (10) immaculate cream-colored pattern of concealed surfaces of thighs; (11) presence of a dark brown stripe from tip of snout to eye, including inter-nostril area, and a dark brown interorbital bar; (12) supratympanic fold bold, black; (13) gular region yellowish; (14) iris color cooper with brown reticulations. Morphological comparisons with other species within the Pristimantis conspicillatus species group. The shagreen texture of the dorsal skin differentiates Pristimantis relictus sp. nov. from P. avicuporum (Duellman & Pramuk, 1999), P. metabates (Duellman & Pramuk, 1999), P. johannesdei (Rivero & Serna, 1988), P. phalaroinguinis (Duellman & Lehr, 2007), and P. pictus (all have smooth dorsal skin). Nineteen species have smooth ventral skin, differing from the areolate ventral skin of P. relictus sp. nov.: P. achatinus (Boulenger, 1898), P. buccinator (Rodriguez, 1994), P. charlottevillensis (Kaiser, Dwyer, Feichtinger & Schmid, 1995), P. chiastonotus (Lynch & Hoogmoed 1977), P. citriogaster (Duellman, 1992), P. condor (Lynch & Duellman, 1980), P. conspicillatus (G��nther, 1858), P. fenestratus (Steindachner 1864), P. gaigei, P. gutturalis, P. johannesdei P. latro, P. lymani (Barbour & Noble, 1920), P. malkini (Lynch, 1980), P. metabates, P. peruvianus (Melin, 1941), P. phalaroinguinis, P. samaipatae (K��hler & Jungfer 1995), and P. vilarsi (Melin, 1941). The areolate ventral skin also distinguishes P. relictus sp. nov. from P. dundeei (Heyer & Mu��oz, 1999), P. giorgii, P. iiap Padial, Gagliardi-Urrutia, Chaparro & Guti��rrez, 2016, P. incertus, and P. ventrigranulosus Maciel, Vaz-Silva, Oliveira & Padial, 2012 (ventral skin granular or slightly granular). The absence of dorsolateral folds distinguishes P. relictus sp. nov from Pristimantis adiastolus (Duellman & Hedges, 2007), P. ardilae, P. avicuporum, P. bipunctatus (Duellman & Hedges, 2005), P. buccinator, P. chiastonotus, P. conspicillatus, P. iiap, P. latro, P. malkini, P. meridionalis (Lehr & Duellman, 2007), P. peruvianus, P. pluvian, P. skydmainos, and P. zeuctotylus (presence of dorsolateral folds) The subovoid snout in dorsal view of P. relictus sp. nov. distinguishes it from P. carranguerorum (Lynch, 1994) (obtuse) and P. medemi (Lynch, 1994), P. latro, and P. zeuctotylus (subacuminate). The immaculate cream-colored pattern of concealed surfaces of thighs differentiates P. relictus sp. nov. from P. pluvian (red), P. bipunctatus, P. condor, P. conspicillatus, P. malkini, P. peruvianus, and P. pictus (with well-defined spots). In addition to the new species, five other species of Pristimantis are distributed in Northeast Brazil: P. moa, P. paulodutrai, P. ramagii, P. rupicola and P. vinhai (see Table 1). While Pristimantis relictus sp. nov. is distributed in highland marshes of Cear��, P. moa is known from transition areas between the Cerrado and Amazon Forest (Oliveira et al. 2020), P. rupicola occurs in Campos Rupestres of Chapada Diamantina (Taucce et al. 2020), and the remaining three species inhabit the Atlantic Forest domain (Trevisan et al. 2020). Besides geographic distribution, P. relictus sp. nov. can be distinguished from P. moa by areolate ventral skin (smooth in P. moa), snout subovoid in dorsal view (truncate in P. moa), immaculate cream-colored pattern of concealed surfaces of thighs (strongly stained yellow on a dark background in P. moa), finger fringes absent (present in P. moa), and webbing on toes absent (present in P. moa) (Oliveira et al. 2020). The new species differs from P. rupicola by possessing shagreen dorsal skin (granular in P. rupicola), and snout subovoid in dorsal view (rounded or truncate in P. rupicola) (Taucce et al. 2020). The new species can be distinguished from the Atlantic Forest species as follows: from P. paulodutrai by the presence of nuptial pads (absent in P. paulodutrai), ventral skin areolate (granular in P. paulodutrai), and immaculate creamcolored pattern of concealed surfaces of thighs (with red blotches in P. paulodutrai) (Bokermann, 1975); from P. ramagii by snout subovoid in dorsal view (subacuminate in P. ramagii), presence of nuptial pads and toe fringes (absent in P. ramagii) (Boulenger, 1888); and from P. vinhai by ventral skin areolate (slightly granular in P. vinhai), snout subovoid in dorsal view (acuminate in P. vinhai) (Bokermann, 1975). Bioacustical comparisons with other species within the Pristimantis conspicillatus species group. The shorter note duration (20���50 ms) differentiates P. relictus sp. nov. from P. samaipatae (59���141 ms) and P. fenestratus (50���91 ms) (Padial & De La Riva 2009). The lower note rate (0.5���2.2 notes/s) distinguishes P. relictus from P. fenestratus (7.7���12.7), P. koehleri (11.8���17.3), P. samaipatae (2.7���14.9), P. dundeei (13.7���20.7) (Heyer & Mu��oz 1999; Giaretta et al. 2018), and P. vilarsi (14) (Heyer & Barrio-Amor��s 2009). The lower number of pulses per note of P. relictus sp. nov. (2���5) also differentiates the new species from P. fenestratus (9���17), P. samaipatae (11���23), P. latro (6���9), and P. pluvian (8���16). The higher fundamental frequency of P. relictus sp. nov. (1894.9���2153.3 Hz) distinguishes the new species from P. samaipatae (1535���1834 Hz), P. latro (1342���1448.6 Hz), P. moa (1320.8���1660.2 Hz), and P. giorgii (663.2���1872.3 Hz). The higher peak frequency of P. relictus sp. nov. (3617.6���4220.5 Hz) differentiates the new species from P. fenestratus (1710���3591 Hz), P. latro (2635.9���3272 Hz), P. moa (2657.1���3400 Hz), and P. pictus (2487.4���3272.2 Hz) (Padial & De La Riva 2009; Oliveira et al. 2017, 2020). Also, advertisement call differentiates P. relictus from P. rupicola. The new species has a higher peak of frequency (3617.6���4220.5 Hz), located in the second band of energy, [peak frequency (2410���3490 Hz) in the first band of energy of the call; Taucce et al. (2020)]. The advertisement call of P. relictus is very similar to that of P. ramagii but the species can be distinguished by the number of pulses per note of 2���5 (7���53 in P. ramagii; Octaven et al. 2017). Description of the holotype. Adult male (URCA-H 13344; Figure 2). Head longer than wide; head width 37% of SVL; head length 40% of SVL; snout subovoid in dorsal view, rounded in lateral view; nostrils not protuberant, directed anterolaterally; internarial distance less than eye-nostril distance; canthus rostralis angular, weakly concave, loreal region slightly concave; eye large, with horizontally elliptical pupil; eye diameter slightly larger than interorbital distance; cranial crests absent; upper eyelid smooth, 74% of eye diameter; tympanum large, with distinct rounded tympanic annulus; tympanic membrane present, visible; supratympanic fold present; labial bars absent; vocal sac subgular, small; tongue ovoid covering the entire floor of mouth, free and notched behind; vocal slits present, lateral to tongue; choana small, rounded; odontophores oblique and widely separated posterior to the choanae, each one bearing six teeth; skin on dorsum shagreen with scattered low tubercles; dorsolateral fold absent; skin on venter areolate, weakly spotted, discoidal fold present; hands large, 29% of SVL; relative lengths of fingers II Measurements of the holotype (mm). SVL 24.4, HL 9.7, HW 8.9, ED 3.4, SL 4.4, IND 2.0, END 2.8, IOD 3.3, UEW 2.5, TD 2.0, FAL 5.4, HAL 7.0, FIII 1.4, FIV 1.3, THL 12.4, TL 13.1, TAL 7.1, FL 10.7, and TIV 1.4. Color in preservative. Dorsal and lateral surfaces pale cream, snout light brown, brown interocular band in dorsal view, distinct dark brown stripe from tip of snout to posterior region of supratympanic fold. Irregular brown marks in lower orbital region. Two brown chevrons on dorsum with irregular brown dots in inguinal region, light brown stripes on dorsal surfaces of limbs. Ventral surface background pale cream in color. Advertisement call. The call of Pristimantis relictus sp. nov. is described based on three individuals (MNRJ 55582 and two unvouchered individuals) (see Table 2;). The vocalization is composed of 1���8 multi pulsed notes (2���5 pulses per note), with an ascendant amplitude modulation. The call duration ranges 0.02��� 0.66 s (0.06 �� 0.08; n = 181), with intercall interval of 0.8��� 9.6 s (2.12 �� 1.5; n = 74), note duration ranges 0.02��� 0.05 s (0.03 �� 0.01; n = 150), with a note rate of 0.52���2.20 notes per second (Figure 3. The call has three visible bands. The fundamental frequency is in the first band, ranging 1894.9���2153.3 Hz, followed by a second band located in the dominant frequency, ranging 3617.6���4220.5 Hz, and a third weak band with a higher frequency, ranging 4565���6287.7 Hz. Variation among paratypes. Adult females (SVL 26.8���32.8 mm) are slightly larger than adult males (SVL 23.1���29.7 mm) (Table 3). Variation in coloration of fixed specimens is mostly related to the presence of chevrons on the dorsum (URCA-H 6980, 2373, 1544, 13342, 6988, and 9828). In life, the dorsal coloration is highly polymorphic: dark brown, reddish to yellowish-brown, or grey (Figure 4), with the presence of dark brown chevrons and black spots in some individuals. Other variations in coloration among paratypes are: brown bars on the leg and arm; dark brown stripe from the tip of the nose to the eye, including the internarial area; dark brown bar in the interorbital region; labial region yellowish to white; supratympanic fold bold black; gular region yellowish; and iris cooper-colored with brown reticulations. Geographic distribution. Pristimantis relictus is endemic to the ���Brejos de Altitude��� of the state of Cear��, Northeast Brazil. This species occurs in the Planalto da Ibiapaba (municipalities of Ipu, Tiangu��, Ubajara, Granja, Vi��osa do Cear��), Serra de Maranguape (municipality of Maranguape), Serra da Aratanha (municipality of Pacatuba), Serra de Baturit�� (municipalities of Guaramiranga, Pacoti, Aratuba, Mulungu), Serra da Uruburetama (Roberto & Loebmann 2016), and Serra da Meruoca (municipality of Meruoca) (Figure 5). Natural history. Pristimantis relictus occurs in dry forests and humid forests (Loebmann & Haddad 2010; Roberto & Loebmann 2016; Castro et al. 2019), from 100 to 900 m.a.s.l. The species has a prolonged breeding period throughout the raining season from December to April. Vocalization occurs from 5 pm throughout the night, with a peak of individuals vocalizing during 6���9 pm. Individuals were found vocalizing on leaves, tree trunks and twigs, mostly facing downwards. Amplexus is axillary, and the eggs are deposited on humid surfaces of rock and wooden debris. Mature females range 28���33 mm and 1.45���3.27 g (n = 13), and clutches have 21��� 53 eggs (n = 13). We observed territorial behavior involving agonistic combat and the emission of territorial and release calls; unfortunately, these calls were not recorded. At Serra de Baturit�� mountain range we recorded the following species occurring in sympatry with P. relictus: Boana raniceps, Dendropsophus minusculus, D. minutus, D. tapacuarensis, Pithecopus gonzagai, Scinax tropicalia, Trachycephalus typhonius, Adelophryne baturitensis, Odontophrynus carvalhoi, Rhinella casconi, R. dapsilis, R. diptycha, R. granulosa, Adenomera juikitam, Leptodactylus macrosternum, L. natalensis, L. pustulatus, L. mystaceus and Physalaemus cuvieri. See Loebmann & Haddad (2010), and Castro et al. (2019), Published as part of Roberto, Igor Joventino, Loebmann, Daniel, Lyra, Mariana L. & ��vila, Robson Waldemar, 2022, A new species of Pristimantis Jim��nez de la Espada, 1870 (Anura: Strabomantidae) from the " Brejos de Altitude " in Northeast Brazil, pp. 521-540 in Zootaxa 5100 (4) on pages 524-532, DOI: 10.11646/zootaxa.5100.4.4, http://zenodo.org/record/6224832, {"references":["Rocha, D. 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C., Batalha-Filho, H., Garda, A. A., Menezes, L., Dias, I. R., Sol, M., Canedo, C., Junc, F. A. & Napoli, M. F. (2020) Cryptic diversity and ancient diversification in the northern Atlantic Forest Pristimantis (Amphibia, Anura, Craugastoridae). Molecular Phylogenetics and Evolution, 148, 1055 - 7903. https: // doi. org / 10.1016 / j. ympev. 2020.106811","Duellman, W. E. & Pramuk, J. B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers, Natural History Museum, The University of Kansas, 13, 1 - 78.","Rivero, J. A. & Serna, M. A. (1988) Tres nuevas especies de Eleutherodactylus (Amphibia, Leptodactylidae) de Antioquia, Colombia. Caribbean Journal of Science, 23, 386 - 399.","Duellman, W. E. & Lehr, E. (2007) Frogs of the genus Eleutherodactylus (Leptodactylidae) in the Cordillera Occidental in Peru with descriptions of three new species. Scientific Papers, Natural History Museum, The University of Kansas, 39, 1 - 13.","Boulenger, G. A. (1898) An account of the reptiles and batrachians collected by Mr. F. H. Rosenberg in western Ecuador. Proceedings of the Zoological Society of London, 1898, 107 - 126.","Rodriguez, L. (1994) A new species of the Eleutherodactylus conspicillatus Group from Peru, with comments on its call. Alytes, 12, 49 - 63.","Kaiser, H., Dwyer, C. M., Feichtinger, W. & Schmid, M. (1995) A new species of Eleutherodactylus (Anura: Leptodactylidae) from Tobago, West Indies, and its morphometric and cytogenetic characterization. Herpetological Natural History, 3 (2), 151 - 163.","Hoogmoed, M. S., Lynch, J. D. & Lescure, J. (1977) A new species of Eleutherodactylus from Guiana (Leptodactylidae, Anura). Zoologische Mededelingen, 51, 33 - 42.","Duellman, W. E. (1992) A new species of the Eleutherodactylus conspicillatus group (Anura: Leptodactylidae) from northeastern Peru. Revista Espanola de Herpetologia, 6, 23 - 29.","Lynch, J. D. & Duellman, W. E. (1980) The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 69, 1 - 86. https: // doi. org / 10.5962 / bhl. title. 16222","Gunther, A. (1858) Neue Batrachier in der Sammlung des Britischen Museums. Archiv fur Naturgeschichte, 24, 319 - 328.","Steindachner, F. (1864) Batrachologische Mittheilungen. Verhandlungen des Zoologisch-Botanischen Vereins in Wien, 14, 239 - 288.","Barbour, T. & Noble, G. K. (1920) Some amphibians from northwestern Peru, with a revision of the genera Phyllobates and Telmatobius. Bulletin of the Museum of Comparative Zoology at Harvard, 63, 395 - 427.","Melin, D. E. (1941) Contributions to the knowledge of the Amphibia of South America. Goteborgs Kungl. Vetenskaps-och Vitterhets-samhalles. Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1 - 71.","Kohler, J. & Jungfer, K. H. (1995) Eine neue Art und ein Erstnachweis von Froschen der Gattung Eleutherodactylus aus Bolivien. Salamandra, 31 (3), 149 - 156.","Heyer, W. R. & Munoz, A. M. (1999) Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on the types and type locality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and a description of a new species of Eleutherodactylus from Mato Grosso, Brazil. Proceedings of the Biological Society of Washington, 112, 1 - 18.","Padial, J. M., Gagliardi-Urrutia, G., Chaparro, J. C. & Gutierrez, R. C. (2016) A new species of the Pristimantis conspicillatus Group from the Peruvian Amazon (Anura: Craugastoridae). Annals of Carnegie Museum, 83 (3), 207 - 218. https: // doi. org / 10.2992 / 007.083.0302","Maciel, N. M., Vaz-Silva, W., de Oliveira, R. M. & Padial, J. M. (2012) A new species of Pristimantis (Anura: Strabomantidae) from the Brazilian Cerrado. Zootaxa, 3265 (1), 43 - 56. https: // doi. org / 10.11646 / zootaxa. 3265.1.3","Duellman, W. E. & Hedges, S. B. (2007) Three new species of Pristimantis (Lissamphibia, Anura) from montane forests of the Cordillera Yanachaga in Central Peru. Phyllomedusa, 6 (2), 119 - 135. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 6 i 2 p 119 - 135","Duellman, W. E. & Hedges, S. B. (2005) Eleutherodactyline frogs (Anura: Leptodactylidae) from the Cordillera Yanachaga in Central Peru. Copeia, 2005 (3), 526 - 538. https: // doi. org / 10.1643 / CH- 05 - 019 R","Lynch, J. D. (1994) Two new species of the Eleutherodactylus conspicillatus group (Amphibia: Leptodactylidae) from the Cordillera Oriental of Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 19 (72), 187 - 193.","Oliveira, E. A., Silva, L. A., Guimaraes, K. L. A., Penhacek, M., Martinez, J. G., Rodrigues, L. R. R., Santana, D. J. & Hernandez- Ruz, E. J. (2020) Four new species of Pristimantis Jimenez de la Espada, 1870 (Anura: Craugastoridae) in the eastern Amazon. PLoS One, 15, 1 - 28. https: // doi. org / 10.1371 / journal. pone. 0229971","Taucce, P. P. G., Nascimento, J. S., Trevisan, C. C., Leite, F. S. F., Santana, D. J., Haddad, C. F. B. & Napoli, M. F. (2020) A new rupicolous species of the Pristimantis conspicillatus group (Anura: Brachycephaloidea: Craugastoridae) from central Bahia, Brazil. Journal of Herpetology, 54 (2), 245 - 257. https: // doi. org / 10.1670 / 19 - 114","Bokermann, W. C. A. (1975) Tres especies novas de Eleutherodactylus do sudeste da Bahia, Brasil (Anura, Leptodactylidae). Revista Brasileira de Biologia, 34 (1), 11 - 18.","Boulenger, G. A. (1888) On some Reptiles and Batrachians from Iguarasse, Pernambuco. The Annals and Magazine of Natural History, Series 6, 2 (7), 40 - 43. https: // doi. org / 10.1080 / 00222938809460873","Padial, J. M. & de la Riva, I. (2009) Integrative taxonomy reveals cryptic Amazonian species of Pristimantis (Anura). Zoological Journal of the Linnean Society, 155 (1), 97 - 122. https: // doi. org / 10.1111 / j. 1096 - 3642.2008.00424. x","Giaretta, A. A., Teixeira, B. F. V., Bernardes, C. S. & Marinho, P. (2018) Distribution, call and habitat of Pristimantis dundeei (Anura, Craugastoridae). Neotropical Biodiversity, 4 (1), 134 - 137.","Heyer, W. R. & Barrio-Amoros, C. L. (2009) The advertisement calls of two sympatric frogs, Leptodactylus lithonaetes (Amphibia: Anura: Leptodactylidae) and Pristimantis vilarsi (Amphibia: Anura: Strabomantidae). Proceedings of the Biological Society of Washington, 122 (3), 282 - 291. https: // doi. org / 10.2988 / 09 - 02.1","Oliveira, E. A., Rodrigues, L. R., Kaefer, I. L., Pinto, K. C., Hernandez-Ruiz, E. J. (2017) A new species of Pristimantis from eastern Brazilian Amazonia (Anura, Craugastoridae). ZooKeys, 687, 101 - 129. https: // doi. org / 10.3897 / zookeys. 687.13221","Oitaven, L. P. C., Santos, J. R. O., Silva, A. D. O., Gambale, P. G. & Moura, J. G. B. (2017) Description of vocalisations and analysis of intra and inter-individual variation in Pristimantis ramagii (Boulenger, 1888) in an upland swamp, northeast Brazil. Herpetology Notes, 10, 197 - 203.","Heyer, W. R. & De Carvalho, C. M. (2000) The enigmatic advertisement call of Eleutherodactylus ramagii (Amphibia: Anura: Leptodactylidae). Amphibia-Reptilia, 21, 117 - 121."]}

Pristimantis blasi sp. nov. (Figs. 2���5). Holotype. ARUQ1291 (Figs. 2���3), adult female collected ESMA Alto Amurrup��, Santa Cecilia, municipality of Pueblo Rico, department of Risaralda, Colombia (5.31444 N, 76.15361 W, 1119 m a.s.l, WGS84), on December 27, 2019, by Manuela Montoya-Mar��n, Jackeline Rivera-G��mez and Sebasti��n Duarte-Mar��n. Paratypes. (N=8), ARUQ1025���1029, CUS-A 145���147 (Figs. 4, 5), male adults collected with the holotype. Referenced specimens. One subadult female ARUQ1030, and two subadult males ARUQ1031���1032, collected with holotype. Definition. Pristimantis blasi sp. nov. is diagnosed by the following combination of characters: (1) skin on dorsum shagreen (with scattered tubercles), becoming finely shagreened laterally and coarsely on the flanks; dorsolateral folds absent; lateral folds present; ventral skin areolate; discoidal fold present, well anterior to groin. (2) Tympanic membrane partially translucent; tympanic annulus evident, ovoid, corresponding to 1/3 (30%���45%) of eye length; supratympanic fold distinguished, which extends from posterior corner of orbit along upper edge of temporal region and curved toward the insertion of the arm. (3) Snout subacuminate in dorsal view, rounded in lateral profile and lacking of papilla; canthus rostralis straight in dorsal view, rounded in profile; two and three subconicals postrictal tubercles. (4) IOD wider than upper eyelid; craneal crest absent; frontoparietals concave at the proximal edge in contact with the exoccipital in lateral view; upper eyelid bearing small low tubercle at the posterior margin. (5) Choanae small, ovoid; not concealed by palatal shelf of maxillary arch; dentigerous processes of vomers prominent, positioned posterior to level of choanae, moderately separated, each dentigerous process of vomers bearing 5���7 teeth. (6) Males with vocal slits and single, median and subgular vocal sac; double nuptial pads glandular and white, present on the dorsomedial surface on the base of the thumb. (7) Finger I shorter than II; discs truncated. (8) Lateral fringes on fingers; palmar tubercle in heart-shape (not divided); thenar tubercle oval, slightly smaller than palmar tubercle; supernumerary tubercles low, distributed on all fleshy parts of palm; subarticular tubercles low, with rounded base and larger than supernumerary tubercles, hyperdistal tubercle present. (9) Antebrachial tubercle present; two and three ulnar tubercles small and subconical, not coalesced. (10) Subconical tubercle on heel and small on outer edge of tarsus, small inner tarsal fold. (11) Oval inner metatarsal tubercles, representing three times the size of outer tubercle; supernumerary plantar tubercles restricted to the proximal segments of the digits subarticular tubercles rounded, prominent. (12) Toes with narrow lateral fringes, toe webbing absent; toe III shorter than toe V; toe III extending to distal edge of the penultimate subarticular tubercle of toe IV; toe V reaching distal edge of distal subarticular tubercle of toe IV; discs and circumferential grooves present on all toes. (13) Two cloacal tubercles present, anterior to cloaca. (14) Polymorphic dorsal pattern (Fig. 4), dorsum darker brown with longitudinal stripes or blotches pale brown, labial bars cream, longitudinal stripe from the posterior edge of the eye, covering 30���50% of the tympanum, which extends to the flanks; flanks darker brown with cream, in anterior surface with some dark blotches; groin, anterior and posterior surfaces of thighs black with yellow light blotches; throat and belly surfaces with black markings on cream or pale yellow; concealed surfaces of tibia with contrasting pattern consisting of black mottling on cream or yellow pale background; pale band across heel; iris red wine in females, copper red in males. (15) SVL adult males 19.2���31.0 mm (mean = 22.4 �� SD = 3.93 mm; n = 8) and female 40.0 mm. Description of the holotype. An adult female with head wider than body; head width 38.9% of SVL; HL 43.7% of SVL; snout subacuminated in dorsal view and rounded in lateral profile; snout lacking papilla at its tip. Canthus rostralis concave in dorsal view, in transverse cross section is rounded; loreal region straight, lips no flared.Anterior part of nostrils is directed laterally; internarial distance 60.4% of eye-to-nostril distance; eye-to-nostril distance equal to of eye diameter. Upper eyelid bearing small subconical tubercle at the posterior margin, upper eyelid width is 92.2% of IOD and eye diameter 118.4% of IOD. Craneal crest absent. Frontoparietals region concave at the proximal edge in contact with the exoccipital in lateral view. Upper edge of tympanum covered by supratympanic fold, which extends from posterior corner of orbit along upper edge of temporal region and curved toward the insertion of the arm; tympanum size is 38.4% of eye diameter; tympanum superficial, ovoid, and partially translucent; tympanic annulus evident, eye to tympanum distance is 58.9% of tympanum diameter; two postrictal subconical tubercles. Choanae small, ovoid, not concealed by palatal shelf of maxillary arch. Dentigerous processes of vomers prominent, positioned posterior to level of choanae, moderately separated, bearing 5 vomerine teeth. Tongue longer than wide, posterior edge notched, posterior 2/3 not adherent to the floor of the mouth. Skin on dorsum shagreen, with scattered tubercles, laterally finely shagreen and coarsely on the flanks; dorsolateral folds absent. The groin and throat skin are smooth. Venter areolate. Discoidal fold well anterior to groin. Upper surface of limbs is smooth, with absent transversal folds over tibia. Radio-ulna length is 22.5% of SVL and hand length is 144.8% of radio-ulna length. Forearm on outer edge bearing three cream small subconical ulnar tubercles. Elbow fold present. Palmar tubercle in heart-shape (not divided), twice wider than the oval thenar tubercle. Supernumerary tubercles low, distributed on all fleshy parts of palm (among six and seven). Subarticular tubercles are rounded and low. Fingers bearing lateral fringes and truncate disks with rounded borders. Pad on thumb slightly wider than digit above the pad. The pads on fingers II-IV broadly wider than digits, the fingers III and IV the disks nearly as large as tympanum, ventral pads on fingers are broader than long. First finger is shorter than second. Tibia length is 51.0% of SVL; subconical tubercle on heel, three cream small tubercles on outer edge of tarsus. Small inner tarsal fold. Foot length is 50.0% of SVL. Metatarsal tubercles are subconical. Inner metatarsal tubercles oval twice as long as wide, 3 times the size of outer metatarsal rounded conical tubercle. Supernumerary plantar tubercles are low (four), restricted to the proximal segments of the digits. Subarticular tubercles rounded, prominent. Toes bearing narrow lateral fringes, toes lack webbing. The pads on toes wider than digits, the pads on toes size order from highest to lowest is 4>5=3>2>1; pads on toes III, IV and V twice wider than digits; ventral pads truncate, wider than long. Outer pads on toes slight narrower than the pads on outer fingers. Tip of V reaching distal edge of distal subarticular tubercle of toe IV, tip of toe III extending to distal edge of the penultimate subarticular tubercle of toe IV. Coloration of the holotype. In life, darker brown with irregular longitudinal stripe pale brown dorsally; flanks darker brown with cream, in anterior surface with some dark blotches. Labial bars, nostrils and longitudinal stripe from the posterior edge of the eye cream. Groin, anterior and posterior surfaces of thighs black with yellow light blotches. Throat and belly surfaces with black markings on cream or pale yellow; concealed surfaces of tibia with contrasting pattern consisting of black mottling on yellow pale background; pale band across heel. Cloacal triangle absent. Iris red wine with a black transversal band. In ethanol, the patterns remain, but the dorsum turns gray and ventral surfaces turn cream. Variation. Measurements and ratios of males and female are shown in Table 1. Some males (ARUQ1025, 1028; CUS-A146) have two small suprascapular tubercles. Male ARUQ1025 have W-shaped scapular fold. Some males (ARUQ1029; CUS-A145���146) have four ulnar tubercles. The dorsal pattern varies from longitudinal stripes or blotches (Fig. 4). The throat may be dark with irregular cream blotches (ARUQ1027���1029, 1031���1032; CUSA146���147). Groin, anterior and posterior surfaces of thighs black with pale blotches (ARUQ1029, 1031���1032; CUSA146���147). Ventrally, finger I, II and concealed surfaces of tibia pale orange with black markings (ARUQ1025, 1027; CUS-A145���146, Fig. 5), Iris copper red in males. Distribution and Natural history. Pristimantis blasi inhabits primary and secondary in tropical humid forests of the western slope of the Cordillera Occidental, from 1000 to 1350 m a.s.l (Fig. 6). The individuals of this species are nocturnal, being active from 2100 to 2300 h; however, it is possible to find inactive individuals during the day by removing the fallen leaves. The individuals are found in ferns, giant ear plant (Xanthosoma sp.) and trunks at a height from 0.1 to 1.5 m. The advertisement call of P. blasi consists in a pulsed note similar as �����ec��� to the human ear, like P. erythropleura and P. cruentus, unfortunately we were unable to record the vocalizing males. The individuals of this species generally have a strong smell in their groin, similar to the macerated herbs. When finding and handling these frogs, no defensive behaviors were observed other than fleeing quickly or staying immobile in the vegetation when found. Sympatric species include Diasporus quidditus, Oophaga histrionica, Phyllobates bicolor, Pristimantis orpacobates and P. labiosus. Conservation status of the new species. Pristimantis blasi was found within ESMA Alto Amurrup�� (type locality) and NNP Tatam�� (Gonz��lez-Dur��n coms. pers.). In these localities there are secondary and primary forests, and the presence of agriculture, which includes livestock, mining activities and logging. Although P. blasi was not as abundant, 5 to 10 individuals could be found in a few hours of sampling. We suggest that this species should be ���Endangered��� according to the IUCN category, by criterion ���B1(a)��� (extent of occurrence less than 5000 km �� and a number of localities Etymology. We proposed the etymology blasi in honor to Blas Antonio C��rdenas for his valuable work in the conservation of Santa Cecilia biodiversity, for his personal contributions to the ecotourism and the protection of cloud forests in the municipality of Pueblo Rico, which directly benefited the species we describe here (Fig. 6C). Comparative diagnosis. Pristimantis blasi sp nov. (Fig. 7A) is most similar to five species inhabiting northern of Cordillera Occidental (P. aemulatus, P. cruentus, P. mars, P. orpacobates and P. ruedai), and three species of Cordillera Central and Occidental (P. bicolor, P. erythropleura, and P. penelopus) by having nuptial pad in males, digital discs expanded, digital pads with circumferential grooves, heel tubercle present, dorsolateral fold present (present in some morph of P. erythropleura), venter areolate, inner tarsal fold present, cranial crests absent, toe V reaches distal subaticular tubercle of toe IV and bright flash color in the inguinal region (Fig. 7). It differs from those species by dorsum skin shagreen with low tubercles (smooth in P. erythropleura, P. bicolor, P. ruedai and P. cruentus), cranial crest absent (present in P. orpacobates), frontoparietals concave at the proximal edge in contact with the exoccipital in lateral view, a small subconical on the upper eyelid, vocal slits and vocal sac (absent in P. cruentus, P. orpacobates and P. mars, P. penelopus), iris red wine in females, and groin of thighs black with yellow light blotches (Fig. 8). Additional differences among species are summarized in Table 2., Published as part of Duarte-Mar��n, Sebasti��n, Montoya-Mar��n, Manuela & Rivera-G��mez, Jackeline, 2022, A New Species of Red-eyed frog of the genus Pristimantis (Anura: Strabomantidae) from the Western Slope of the Cordillera Occidental, Risaralda, Colombia, pp. 218-232 in Zootaxa 5093 (2) on pages 219-227, DOI: 10.11646/zootaxa.5093.2.5, http://zenodo.org/record/5905199, {"references":["Lynch, J. D. (1992) Distribution and variation in a Colombian frog, Eleutherodactylus erythropleura (Amphibia: Leptodactylidae). Studies on neotropical fauna and environment, 27 (4), 211 - 226. https: // doi. org / 10.1080 / 01650529209360880","Ruiz-Carranza, P. M., Lynch, J. D. & Ardila-Robayo, M. C. (1997) Seis nuevas especies de Eleutherodactylus Dumeril, Bibron, 1841 (Amphibia: Leptodactylidae) del Norte de la Cordillera Occidental de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 21, 155 - 174.","Lynch, J. D., Ruiz-Carranza, P. M. & Ardila-Robayo, M. C. (1994) The identities of the Colombian frogs confused with Eleutherodactylus latidiscus (Boulenger) (Amphibia: Anura: Leptodactylidae). Natural History Museum, the University of Kansas, 170, 1 - 42.","Savage, J. M. (2002) The amphibians and reptiles of Costa Rica: a herpetofauna between two continents, between two seas. University of Chicago press, Chicago, Illinois, 934 pp."]}

We describe a new species of red-eyed frog of the genus Pristimantis from the tropical forest at elevations of 1000–1350 m on western slope of the Cordillera Occidental, department of Risaralda, Colombia. The new species differs from congeners by having dorsum skin shagreened with low tubercles; vocal sac single, median and subgular in males, iris red wine in females copper red in males, and by presenting black groin coloration with yellow light blotches. In addition, the new species has concave frontoparietals, an unusual character described in Brachycephaloidea frog species.

Pristimantis postducheminorum sp. nov. Figures 3 –, 4, 5; Table 2. Holotype. MHUA-A11550 (Figs. 3 and 4). An adult female, at the Mesenia-Paramillo Nature Reserve (5.496º N, 75.889º W), about 1.5 km south of the field station at 2800 m.a.s.l., 14 km southwest of the municipality of Jardín, vereda La Mesenia, municipality of Andes, department of Antioquia, Colombia; collected on July 2, 2018, by Luis A. Mazariegos-H. and Ubiel Rendón. Paratypes. (Figs. 3, 5) Three males (ANDES-A4425, MHUA-A11551, MHUA-A12715), and five females (ANDES-A4427, ANDES-A4428, MHUA-A11549, MHUA-A12715), collected at the type locality, between June 2017 and October 2019 by Luis A. Mazariegos-H. and Ubiel Rendón. An unsexed specimen (MHUA-A11313), El Centello Nature Reserve (5.504º N, 75.849º W, 2400 m.a.s.l.), vereda Macanas, municipality of Jardín, department of Antioquia, Colombia less than 5 km northeast (bearing 68º) from the type locality, collected on September 5, 2018 by Fredy A. Grisales. Etymology. We propose the specific epithet postducheminorum as a patronym to honour Dr. Gerald Post, veterinary oncologist and conservation biologist, and his husband David Duchemin. Both are avid conservationists and supporters of non-profit organisations dedicated to raise awareness and take conservation actions to help biodiversity. Definition and diagnosis. We assign the new species to the genus Pristimantis on the basis of the phylogenetic results (Fig. 1). Pristimantis postducheminorum sp. nov. is characterised by (1) body dorsal surface of the skin with sparse conical tubercles, much coarser towards the posterior end of the dorsum, without any apparent distribution pattern; discoidal fold barely visible; dorsolateral folds absent; paravertebral folds absent; postocular folds short, extending posteromedially from the eyes, barely reaching the line connecting the axillae, often fragmented into pustules; lateral surfaces of the body intermediate between dorsum and belly, albeit more similar to the dorsum; all the ventral surfaces are areolate, including belly, proximal surface of the thighs, throat and chest; (2) tympanic membrane and tympanic annulus prominent; supratympanic folds present, arc-shaped, diagonal, and ventrally bordered by a diagonal black band; horizontal diameter of tympanum 40–50% of eye diameter (Table 1); (3) snout rounded from both dorsal and lateral views (Fig. 4); snout tubercle absent; interocular fold absent; (4) upper eyelid tubercle present and conical, often with 1–3 smaller subconical tubercles on the eyelid; 1–2 postrictal tubercles, subconical and low, ovoid at the base; cranial crests absent; (5) vomerine odontophores triangular in shape, oblique, and widely separated; choanae ovoid; (6) males with barely visible vocal slits; nuptial pads absent, very small and whitish testis; (7) finger I slightly shorter than finger II (Fig. 4); webbing short and basal among the fingers; finger discs with pads, expanded and truncate (wider than long); (8) lateral fringes of fingers present; one subarticular tubercle in fingers I (thumb) and II, two on fingers III and IV; all subarticular tubercles low and rounded, and slightly oval on the base; thenar tubercle oval or somehow elongate; palmar tubercle oval; supernumerary tubercles in hand present, and small (Fig. 4); (9) between 1–5 ulnar tubercles, subconical, not forming a fold, notorious in larger individuals, without a distinctive coloration; (10) heel tubercles present, as long as wide; (11) inner metatarsal tubercle longer than wide, and oval; outer metatarsal tubercle about half in area compared to inner metatarsal tubercle, and oval in shape; between 1–3 supernumerary tubercles at the base of toes II–IV (Fig. 4); (12) toe V longer than toe III; toes without fringes but with barely noticeable basal webbing; toes I and II with a single subarticular tubercle; toes III and V with two subarticular tubercles; and toe IV with three subarticular tubercles; toe discs with pads, expanded and truncate (Fig. 4); (13) in life (Fig. 3) the dorsum is predominantly dark brown, with black marks of variable shape; throat cream, yellow, or orange, with V-shaped longitudinal black bars or dark blotches; belly and often legs conspicuously coloured, in shades of orange, yellow to reddish, sometimes lighter than throat; limbs brown, often with dark spots or traversed by dark bars; uniform and distinctive dark grey coloration in the inguinal area and in the concealed surfaces of the thighs; iris bronze to dark brown; (14) adults medium in size (Table 1), SVL in males 22.5–23.2 mm (n = 3), SVL in females 27.5–30.3 mm (n = 5). Comparison with related species. The new species is phylogenetically closest to P. satagius (Lynch 1995), to the sympatric P. ferwerdai (Amézquita et al. 2019), as well as to the clade formed by P. kelephus (Lynch 1998) and P. calcaratus (Boulenger 1908) (Table 2). It is also phenotypically similar to P. orpacobates (Lynch et al. 1994), P. quicato (Ospina-Sarria et al. 2011), P. cremnobates (Lynch & Duellman 1980), and P. jubatus (García & Lynch 2006). It can be distinguished externally from all these species but P. jubatus by the conspicuous orange, yellow or reddish cream ventral coloration (versus black, brown or cream in other species); ventral coloration is cream or yellow golden with brown or gray spotting in P. jubatus, but this species exhibits dorsolateral folds and finger lateral fringes, both of which are absent in P. postducheminorum sp. nov. It can also be distinguished by the uniformly dark grey coloration of groin and concealed surface of the thighs (versus brown with cream in P. satagius, black with white blotches in P. ferwerdai, with many pale spots in P. calcaratus, uniformly brown in P. kelephus, alternating dark and brown bars in P. orpacobates, white with dark bars in P. quicato, and pale yellow or yellow-cream with brown spotting in P. jubatus). No other species of Pristimantis with conspicuous ventral coloration has been found at or near the study area. For a comparison of character states among similar species see Table 3. . Description of the holotype (Figs. 3, 4).An adult female, 28.7 mm in snout-vent length (SVL). Head width 41% of SVL; 9% wider than long, head length 34.8% of SVL. Snout rounded in dorsal and lateral views; boreal region concave with very small tubercles; interorbital distance 94.1% of eye-to-nostril distance; canthus rostralis concave in dorsal view and angular in cross-section; eye-to-nostril distance 97.1% of eye-length. Upper eyelid width slightly narrower than IOD, 76.5% of IOD; upper eyelid with one distinctive large, conical tubercle and four smaller ones, cranial crest absent. Tympanic annulus prominent, its diameter 37.1% of eye diameter, supratympanic fold arcshaped, diagonal, and ventrally bordered by a diagonal black band, one postrictal tubercle subconical and low, ovoid at the base; choanae ovoid; vomerine odontophores triangular in shape, oblique, and widely separated; tongue longer than wide. Skin on dorsum granular with some conical tubercles sparsely and irregularly distributed; dorsolateral folds absent; ventral skin areolate, discoidal fold visible, throat with similar pattern as venter but slightly darker and more redish, with V-shaped dark bands; ulnar tubercles present, without forming a distinct fold; thenar tubercle elongate; supernumerary palmar tubercles present; subarticular tubercles visible; fingers with lateral fringes; finger I slightly shorter than finger II; finger discs with pads, expanded and truncate (wider than long), except for finger I, which is rounded; inner digits bearing much narrower discs; hind limbs moderate; tibia long 46.9% of SVL; foot long 41.1% of SVL; relative length of adpressed toes IV> V> III> II> I; tip of toe V extending to base of distal subarticular tubercle of toe IV; tip of toe III extending beyond distal border of penultimate subarticular tubercle on toe IV, but before last subarticular tubercle of toe IV; discs of toes smaller than discs on outer fingers, without lateral fringes; webbing basal; heel bearing small tubercles, one slightly larger and conical; subconical tubercles along outer border of tarsus; inner metatarsal tubercle longer than wide, and oval; outer metatarsal tubercle about half in area compared to inner metatarsal tubercle, and oval in shape; between 1–3 supernumerary plantar tubercles present, at bases of toes II–V; prominent subarticular tubercles. Coloration of the holotype. (Fig. 3). In life, the holotype dorsum was predominantly dark brown, with a mid dorsal yellowish line, and black marks of variable shape; the throat exhibited essentially the same pattern as the belly, but darker, more reddish, and with V-shaped longitudinal black bars; the belly and legs conspicuously coloured in white to reddish, with a mid dorsal ventral line; limbs with essentially the same pattern as the dorsum, except by the distinctive dark grey coloration in the concealed surfaces of the thighs; the flanks darker than the dorsum; the iris bronze to dark brown. Variation of type series. (Figs. 3, 5). Morphometric variation among individuals of the type series is depicted in Table 1. There is strong sexual dimorphism in body length, with males (SVL = 22.8 ± 0.3 mm, N = 3 individuals) attaining 79% % of females’ body size (28.8 ± 1.0 mm, N = 5 individuals). No other measurement appears to differ among the sexes, once allometry is considered, but the low number of individuals preclude any formal statistical comparison. Among standard morphological traits, we found remarkable variation between 1–4 in the number of minor tubercles besides the eyelid tubercle, as well as in the shape of the thenar tubercle: elongate in the holotype (MHUA-A11550), and a second female (MHUA-A11549), but oval in other females and in all males. Regarding coloration, the dorsum of most individuals in the type series is predominantly dark but light brown in male MHUA- A12715, and female MHUA-A12716, always with irregular black marks. The ventral surface was yellow to orange in ANDES-A4428 and ANDES-A4427, light orange in MHUA-A 1549, and white to reddish in MHUA-A11550 (holotype) and MHUA-A11551. Three out of the eight individuals exhibit a mid-ventral and only two a mid-dorsal whitish line; the holotype MHUA-A11550 and MHUA-A11551 exhibited both lines. The throat of three individuals was crossed by the oblique V-shaped anteromedial dark bars (two of them in Fig. 3). We found no sign of sexual dichromatism, but the sample size is too low at this point to infer any generalisation in this regard. In preservative (1–4 years in 70% ethanol), the dorsum looks dark grey with irregular black marks (Fig. 5). Limbs are dark dorsally, and black with creamy areas ventrally. Fingers 1 and 2 are creamy yellow. The belly looks light orange, yellow or cream, with black blotches or reticulation. Both patterning and medial lines remain recognisable in preservative. The gular area looks lighter than in life. The concealed surfaces of the thighs and groin remain uniformly dark grey. The forelimbs they have a greyish green dorsal coloration and a creamy yellow ventral coloration; the limbs posteriors have a creamy yellow coloration on the ventral part of the femur. Natural history, distribution, and conservation status. The specimens of Pristimantis postducheminorum sp. nov. were found sitting on the moss covering tree trunks, branches and roots, and on the outer surface of epiphyte bromeliads (Fig. 6). No individual was seen calling, although a single-note, barely modulated, call was heard in the capture area and often led us to find males of this species. Individuals were far from any pool, puddle, or stream, and only syntopic with Pristimantis peraticus and another unidentified species of Pristimantis. To the best of our knowledge, the species is known only from two localities separated by less than 4.5 km in straight line, both situated within protected forests, yet surrounded by severely fragmented habitats. The species seems also to be rare despite four intensive searches by three people, throughout one week each, at the type locality; the reserve has been also searched for this and other Pristimantis species during the rainy and dry seasons on five different years. Therefore, based on the known area of occupancy of less than 500 km 2, the number of known locations being less than five, and the low number of encountered mature individuals, we proposed to declare this new species as Endangered [IUCN; B2a, 2c. (IV)] until new information is gathered.

Pristimantis postducheminorum sp. nov. Figures 3 ���, 4, 5; Table 2. Holotype. MHUA-A11550 (Figs. 3 and 4). An adult female, at the Mesenia-Paramillo Nature Reserve (5.496�� N, 75.889�� W), about 1.5 km south of the field station at 2800 m.a.s.l., 14 km southwest of the municipality of Jard��n, vereda La Mesenia, municipality of Andes, department of Antioquia, Colombia; collected on July 2, 2018, by Luis A. Mazariegos-H. and Ubiel Rend��n. Paratypes. (Figs. 3, 5) Three males (ANDES-A4425, MHUA-A11551, MHUA-A12715), and five females (ANDES-A4427, ANDES-A4428, MHUA-A11549, MHUA-A12715), collected at the type locality, between June 2017 and October 2019 by Luis A. Mazariegos-H. and Ubiel Rend��n. An unsexed specimen (MHUA-A11313), El Centello Nature Reserve (5.504�� N, 75.849�� W, 2400 m.a.s.l.), vereda Macanas, municipality of Jard��n, department of Antioquia, Colombia less than 5 km northeast (bearing 68��) from the type locality, collected on September 5, 2018 by Fredy A. Grisales. Etymology. We propose the specific epithet postducheminorum as a patronym to honour Dr. Gerald Post, veterinary oncologist and conservation biologist, and his husband David Duchemin. Both are avid conservationists and supporters of non-profit organisations dedicated to raise awareness and take conservation actions to help biodiversity. Definition and diagnosis. We assign the new species to the genus Pristimantis on the basis of the phylogenetic results (Fig. 1). Pristimantis postducheminorum sp. nov. is characterised by (1) body dorsal surface of the skin with sparse conical tubercles, much coarser towards the posterior end of the dorsum, without any apparent distribution pattern; discoidal fold barely visible; dorsolateral folds absent; paravertebral folds absent; postocular folds short, extending posteromedially from the eyes, barely reaching the line connecting the axillae, often fragmented into pustules; lateral surfaces of the body intermediate between dorsum and belly, albeit more similar to the dorsum; all the ventral surfaces are areolate, including belly, proximal surface of the thighs, throat and chest; (2) tympanic membrane and tympanic annulus prominent; supratympanic folds present, arc-shaped, diagonal, and ventrally bordered by a diagonal black band; horizontal diameter of tympanum 40���50% of eye diameter (Table 1); (3) snout rounded from both dorsal and lateral views (Fig. 4); snout tubercle absent; interocular fold absent; (4) upper eyelid tubercle present and conical, often with 1���3 smaller subconical tubercles on the eyelid; 1���2 postrictal tubercles, subconical and low, ovoid at the base; cranial crests absent; (5) vomerine odontophores triangular in shape, oblique, and widely separated; choanae ovoid; (6) males with barely visible vocal slits; nuptial pads absent, very small and whitish testis; (7) finger I slightly shorter than finger II (Fig. 4); webbing short and basal among the fingers; finger discs with pads, expanded and truncate (wider than long); (8) lateral fringes of fingers present; one subarticular tubercle in fingers I (thumb) and II, two on fingers III and IV; all subarticular tubercles low and rounded, and slightly oval on the base; thenar tubercle oval or somehow elongate; palmar tubercle oval; supernumerary tubercles in hand present, and small (Fig. 4); (9) between 1���5 ulnar tubercles, subconical, not forming a fold, notorious in larger individuals, without a distinctive coloration; (10) heel tubercles present, as long as wide; (11) inner metatarsal tubercle longer than wide, and oval; outer metatarsal tubercle about half in area compared to inner metatarsal tubercle, and oval in shape; between 1���3 supernumerary tubercles at the base of toes II���IV (Fig. 4); (12) toe V longer than toe III; toes without fringes but with barely noticeable basal webbing; toes I and II with a single subarticular tubercle; toes III and V with two subarticular tubercles; and toe IV with three subarticular tubercles; toe discs with pads, expanded and truncate (Fig. 4); (13) in life (Fig. 3) the dorsum is predominantly dark brown, with black marks of variable shape; throat cream, yellow, or orange, with V-shaped longitudinal black bars or dark blotches; belly and often legs conspicuously coloured, in shades of orange, yellow to reddish, sometimes lighter than throat; limbs brown, often with dark spots or traversed by dark bars; uniform and distinctive dark grey coloration in the inguinal area and in the concealed surfaces of the thighs; iris bronze to dark brown; (14) adults medium in size (Table 1), SVL in males 22.5���23.2 mm (n = 3), SVL in females 27.5���30.3 mm (n = 5). Comparison with related species. The new species is phylogenetically closest to P. satagius (Lynch 1995), to the sympatric P. ferwerdai (Am��zquita et al. 2019), as well as to the clade formed by P. kelephus (Lynch 1998) and P. calcaratus (Boulenger 1908) (Table 2). It is also phenotypically similar to P. orpacobates (Lynch et al. 1994), P. quicato (Ospina-Sarria et al. 2011), P. cremnobates (Lynch & Duellman 1980), and P. jubatus (Garc��a & Lynch 2006). It can be distinguished externally from all these species but P. jubatus by the conspicuous orange, yellow or reddish cream ventral coloration (versus black, brown or cream in other species); ventral coloration is cream or yellow golden with brown or gray spotting in P. jubatus, but this species exhibits dorsolateral folds and finger lateral fringes, both of which are absent in P. postducheminorum sp. nov. It can also be distinguished by the uniformly dark grey coloration of groin and concealed surface of the thighs (versus brown with cream in P. satagius, black with white blotches in P. ferwerdai, with many pale spots in P. calcaratus, uniformly brown in P. kelephus, alternating dark and brown bars in P. orpacobates, white with dark bars in P. quicato, and pale yellow or yellow-cream with brown spotting in P. jubatus). No other species of Pristimantis with conspicuous ventral coloration has been found at or near the study area. For a comparison of character states among similar species see Table 3. . Description of the holotype (Figs. 3, 4).An adult female, 28.7 mm in snout-vent length (SVL). Head width 41% of SVL; 9% wider than long, head length 34.8% of SVL. Snout rounded in dorsal and lateral views; boreal region concave with very small tubercles; interorbital distance 94.1% of eye-to-nostril distance; canthus rostralis concave in dorsal view and angular in cross-section; eye-to-nostril distance 97.1% of eye-length. Upper eyelid width slightly narrower than IOD, 76.5% of IOD; upper eyelid with one distinctive large, conical tubercle and four smaller ones, cranial crest absent. Tympanic annulus prominent, its diameter 37.1% of eye diameter, supratympanic fold arcshaped, diagonal, and ventrally bordered by a diagonal black band, one postrictal tubercle subconical and low, ovoid at the base; choanae ovoid; vomerine odontophores triangular in shape, oblique, and widely separated; tongue longer than wide. Skin on dorsum granular with some conical tubercles sparsely and irregularly distributed; dorsolateral folds absent; ventral skin areolate, discoidal fold visible, throat with similar pattern as venter but slightly darker and more redish, with V-shaped dark bands; ulnar tubercles present, without forming a distinct fold; thenar tubercle elongate; supernumerary palmar tubercles present; subarticular tubercles visible; fingers with lateral fringes; finger I slightly shorter than finger II; finger discs with pads, expanded and truncate (wider than long), except for finger I, which is rounded; inner digits bearing much narrower discs; hind limbs moderate; tibia long 46.9% of SVL; foot long 41.1% of SVL; relative length of adpressed toes IV> V> III> II> I; tip of toe V extending to base of distal subarticular tubercle of toe IV; tip of toe III extending beyond distal border of penultimate subarticular tubercle on toe IV, but before last subarticular tubercle of toe IV; discs of toes smaller than discs on outer fingers, without lateral fringes; webbing basal; heel bearing small tubercles, one slightly larger and conical; subconical tubercles along outer border of tarsus; inner metatarsal tubercle longer than wide, and oval; outer metatarsal tubercle about half in area compared to inner metatarsal tubercle, and oval in shape; between 1���3 supernumerary plantar tubercles present, at bases of toes II���V; prominent subarticular tubercles. Coloration of the holotype. (Fig. 3). In life, the holotype dorsum was predominantly dark brown, with a mid dorsal yellowish line, and black marks of variable shape; the throat exhibited essentially the same pattern as the belly, but darker, more reddish, and with V-shaped longitudinal black bars; the belly and legs conspicuously coloured in white to reddish, with a mid dorsal ventral line; limbs with essentially the same pattern as the dorsum, except by the distinctive dark grey coloration in the concealed surfaces of the thighs; the flanks darker than the dorsum; the iris bronze to dark brown. Variation of type series. (Figs. 3, 5). Morphometric variation among individuals of the type series is depicted in Table 1. There is strong sexual dimorphism in body length, with males (SVL = 22.8 �� 0.3 mm, N = 3 individuals) attaining 79% % of females��� body size (28.8 �� 1.0 mm, N = 5 individuals). No other measurement appears to differ among the sexes, once allometry is considered, but the low number of individuals preclude any formal statistical comparison. Among standard morphological traits, we found remarkable variation between 1���4 in the number of minor tubercles besides the eyelid tubercle, as well as in the shape of the thenar tubercle: elongate in the holotype (MHUA-A11550), and a second female (MHUA-A11549), but oval in other females and in all males. Regarding coloration, the dorsum of most individuals in the type series is predominantly dark but light brown in male MHUA- A12715, and female MHUA-A12716, always with irregular black marks. The ventral surface was yellow to orange in ANDES-A4428 and ANDES-A4427, light orange in MHUA-A 1549, and white to reddish in MHUA-A11550 (holotype) and MHUA-A11551. Three out of the eight individuals exhibit a mid-ventral and only two a mid-dorsal whitish line; the holotype MHUA-A11550 and MHUA-A11551 exhibited both lines. The throat of three individuals was crossed by the oblique V-shaped anteromedial dark bars (two of them in Fig. 3). We found no sign of sexual dichromatism, but the sample size is too low at this point to infer any generalisation in this regard. In preservative (1���4 years in 70% ethanol), the dorsum looks dark grey with irregular black marks (Fig. 5). Limbs are dark dorsally, and black with creamy areas ventrally. Fingers 1 and 2 are creamy yellow. The belly looks light orange, yellow or cream, with black blotches or reticulation. Both patterning and medial lines remain recognisable in preservative. The gular area looks lighter than in life. The concealed surfaces of the thighs and groin remain uniformly dark grey. The forelimbs they have a greyish green dorsal coloration and a creamy yellow ventral coloration; the limbs posteriors have a creamy yellow coloration on the ventral part of the femur. Natural history, distribution, and conservation status. The specimens of Pristimantis postducheminorum sp. nov. were found sitting on the moss covering tree trunks, branches and roots, and on the outer surface of epiphyte bromeliads (Fig. 6). No individual was seen calling, although a single-note, barely modulated, call was heard in the capture area and often led us to find males of this species. Individuals were far from any pool, puddle, or stream, and only syntopic with Pristimantis peraticus and another unidentified species of Pristimantis. To the best of our knowledge, the species is known only from two localities separated by less than 4.5 km in straight line, both situated within protected forests, yet surrounded by severely fragmented habitats. The species seems also to be rare despite four intensive searches by three people, throughout one week each, at the type locality; the reserve has been also searched for this and other Pristimantis species during the rainy and dry seasons on five different years. Therefore, based on the known area of occupancy of less than 500 km 2, the number of known locations being less than five, and the low number of encountered mature individuals, we proposed to declare this new species as Endangered [IUCN; B2a, 2c. (IV)] until new information is gathered., Published as part of Palacios-Rodr��guez, Pablo, Daza, Juan M., Mazariegos-H, Luis A., Rend��n, Ubiel & Am��zquita, Adolfo, 2022, A new species of Pristimantis (Anura: Strabomantidae) with a colourful venter from the cloud forests of Colombian western Andes, pp. 67-84 in Zootaxa 5092 (1) on pages 69-79, DOI: 10.11646/zootaxa.5092.1.3, http://zenodo.org/record/5869529, {"references":["Lynch, J. D. (1995) Three new species of Eleutherodactylus (Amphibia: Leptodactylidae) from paramos of the Cordillera Occidental of Colombia. Herpetologica, 29, 513 - 521. https: // doi. org / 10.2307 / 1564734","Amezquita, A., Suarez, G., Palacios-Rodriguez, P., Beltran, I., Rodriguez, C., Barrientos, L. S., Daza, J. M. & Mazariegos, L. (2019) A new species of Pristimantis (Anura: Craugastoridae) from the cloud forests of Colombian western Andes. Zootaxa, 4648 (3), 537 - 548. https: // doi. org / 10.11646 / zootaxa. 4648.3.8","Lynch, J. D. (1998) New species of Eleutherodactylus from the Cordillera Occidental of western Colombia with a synopsis of the distributions of species in western Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 22, 117 - 148.","Boulenger, G. A. (1908) Descriptions of new batrachians and reptiles discoverd by Mr. M. G. Palmer in south-western Colombia. Annals and Magazine of Natural History, Series 8, 2, 515 - 522. https: // doi. org / 10.1080 / 00222930808692531","Lynch, J. D., Ruiz-Carranza, P. M. & Ardila-Robayo, M. C. (1994) The identities of the Colombian frogs confused with Eleutherodactylus latidiscus (Boulenger) (Amphibia: Anura: Leptodactylidae). Occasional Papers of the Museum of Natural History, University of Kansas, 170, 1 - 42.","Ospina-Sarria, J. J., Mendez-Narvaez, J., Burbano-Yandi, C. E. & Bolivar-Garcia, W. (2011) A new species of Pristimantis (Amphibia: Craugastoridae) with cranial crests from the Colombian Andes. Zootaxa, 3111, 37 - 48.","Lynch, J. D. & Duellman, W. E. (1980) The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous Publication. Museum of Natural History, University of Kansas, 69, 1 - 86. https: // doi. org / 10.5962 / bhl. title. 16222","Cuellar-Valencia, O. M. & Bernal-Rivera, A. (2020) Pristimantis kelephus. Catalogo de Anfibios y Reptiles de Colombia, 6, 51 - 56."]}

The Terrarana frogs of the genus Pristimantis are acknowledged for their direct development into froglets and for their astonishing species richness, which renders it the anuran genus with the highest number of species. We describe a new species of Pristimantis from the northwestern Andes of Colombia. The species is distributed from an area between 2750-2900 m.a.s.l. in the Mesenia-Paramillo Nature Reserve. Pristimantis postducheminorum sp. nov. differs from other, phylogenetically related, or similar resembling Pristimantis taxa by a striking yellow coloration in the ventral area, dark grey coloration in the concealed surfaces of the thighs and groin, absence of nuptial pads, presence of lateral fringes on fingers, presence of vomerine odontophores triangular in shape from the ventral view, and absence of dorsolateral folds. Molecular phylogenetics place this new species close to P. satagius and therefore within the P. ridens species group. The new species is also phylogenetically close and sympatric with the recently described P. ferwerdai, which further indicates that the Pristimantis fauna has been notoriously underestimated in the Colombian western cloud forests, a fact that should be considered in assessments of environmental impact.

Species richness in the genus Pristimantis is underestimated due to the existence of morphologically cryptic species. This is worsened by the low sampling effort and the lack of studies using genetic markers. Here, we use molecular and morphological data to determine the phylogenetic relationships of a clade of Pristimantis distributed throughout montane tropical forests in the eastern Andes, from central Ecuador to northern Perú. We name this clade the Pristimantis trachyblepharis species group. Our results show that it comprises nine species, of which four are formally described and five are new. Four of these undescribed species are formally described here. The group is composed of miniaturized species, such as Pristimantis nanus sp. nov., currently the smallest known species of the genus and the smallest vertebrate in Ecuador. As a first approach to understanding the evolutionary origin and implications of body-size reduction in Pristimantis, we here study the phylogenetic signal and evolutionary trends of body size within the genus. We also provide the first record of P. aquilonaris in Ecuador and we show, for the first time, the phylogenetic position of P. albujai, P. aquilonaris, P. minimus and P. trachyblepharis, which are also members of the P. trachyblepharis species group.

Pristimantis daquilemai sp. nov. Fig. 2���8 urn:lsid:zoobank.org:act: 0B81B35F-F891-48C9-90CF-80D066920FD2 Common names. English: Daquilema���s Rain Frog. Spanish: Cut��n de Daquilema. Holotype. ZSFQ 1206, adult female collected near the community of R��o Blanco (3.907196��S, 78.485674��W, 2054 m), Paquisha parish, Paquisha county, province of Zamora Chinchipe, Republic of Ecuador (Fig. 4), collected on 10 February 2018 by David Brito-Zapata and Juan Hurtado. Paratypes (22 ♀, 22 ♂ ). Adult females collected at the type locality by David Brito-Zapata and Juan Hurtado: ZSFQ 1201 (3.906969��S, 78.485297��W, 2044 m); ZSFQ 1202���1203, ZSFQ 1207 (3.907441��S, 78.485835��W, 2041 m); ZSFQ 1204���1205, ZSFQ 1210 (3.906679��S, 78.484964��W, 2059 m); ZSFQ 1208 (3.906851��S, 78.485045��W, 2050 m), on 10 February, 2018; DHMECN 13055 (3.906607��S, 78.484857��W, 2073 m), on 12 January, 2016.Adult females collected at La Herradura (4.054202��S, 78.556270��W, 1750 m), Chinapintza, Paquisha county, province of Zamora Chinchipe, Ecuador, by Manuel A. Morales on 30 June, 2003: QCAZ 30842���30843, QCAZ 30847��� 30848, QCAZ 30852, QCAZ 30857���30859, QCAZ 30861���30863, QCAZ 30868���30869. Adult males collected at the type locality by David Brito-Zapata y Juan Hurtado: ZSFQ 1209 (3.907441��S, 78.485835��W, 2041 m); ZSFQ 1211 (3.906516��S, 78.484839��W, 2054 m); ZSFQ 1212 (3.906589��S, 78.484893��W, 2067 m), on 10 February, 2018; DHMECN 13066 (3.910324��S, 78.500900��W, 1740 m), DHMECN 13071, (3.892927��S, 78.516816��W, 1554 m), DHMECN 13080���13081, 13059 (3.906607��S, 78.484857��W, 2073 m), on 12 January, 2016. Adult males collected at La Herradura, by Manuel A. Morales on 30 June, 2003: QCAZ 30844���30846, QCAZ 30849���30851, QCAZ30853���30856, QCAZ 30860; QCAZ 30865���30867. Etymology. The specific epithet ���daquilemai��� is a noun in the genitive case and a patronymic for Fernando Daquilema Guam��n, an indigenous of the Puruh�� culture from Ecuador, who led an insurrection in December 1871, considered as one of the most important indigenous rebellions in 19 th century, Ecuador. Known as the Yaruqu��es insurrection, it was caused by high church taxes and forced labor for national roadworks. The movement mobilized thousands of people against the tactics of the Ecuadorian state-making model carried forward by the presidency of Gabriel Garc��a Moreno (Ibarra, 2018; L��pez-Oc��n, 1986; Mutlak, 2009). Diagnosis. Pristimantis daquilemai differs from its congeners by the following combination of characters: (1) Dorsal skin shagreen with scattered small tubercles, flanks densely tuberculated; ������ ������-shaped scapular folds, with two subconical tubercles on the medial and posterior sections of folds; with thin dorsolateral folds extending from scapular folds to post-sacral region; venter areolate; discoidal fold weakly defined; (2) tympanic membrane not differentiated from surrounding skin, tympanic annulus not visible externally, hidden by muscle and skin; supratympanic region with a weakly defined fold, conspicuous conical or subconical postrictal tubercles; (3) snout subacuminate in dorsal view, rounded in profile, with a prominent rostral papilla at tip of snout; (4) upper eyelid with one elongated conical tubercle and several conical and subconical smaller tubercles, inter-ocular distance wider than upper eyelid; cranial crests absent; (5) dentigerous processes of vomers present, oblique, with two or three teeth; (6) males with weakly defined nuptial pads on dorsum of Finger I; vocal slits and vocal sac absent; (7) Finger I shorter than Finger II; emarginated discs of fingers broadly expanded on fingers II���IV; (8) fingers with lateral fringes; (9) ulnar tubercles present, conspicuous, conical; (10) heels with a conical tubercle; two or three outer tarsal subconical tubercles; indistinct inner tarsal fold; (11) inner metatarsal tubercle ovoid, about 3���4x the size of subconical, rounded, outer metatarsal tubercle; (12) toes with lateral fringes; supernumerary plantar tubercles present, weakly defined basal membrane; expanded discs on all toes, expanded in the same proportion as discs of fingers; Toe V longer than Toe III (disc of Toe V not reaching distal subarticular tubercle on Toe IV; (13) in life, dorsal background orange-brown to dark brown with numerous little orange spots with brown markings; venter and hidden surfaces of posterior legs from greenish-yellow to dark brown with small white dots; groin with orange or yellow spots, with highest intensity and size in females; iris golden with black reticulations and a horizontal reddish brown bar (14) SVL in females 17.3�� 1.1 mm (15.5���19.0 mm, n = 22), in males 13.2�� 0.9 mm (12.0���15.0 mm, n = 22). Comparisons. Detailed comparisons with similar species are presented in Table 1. Pristimantis daquilemai is morphologically similar to P. trachyblepharis (Boulenger, 1918), P. aquilonaris (Edgar Lehr, Aguilar, Siu-Ting, & Carlos Jord��n, 2007), P. minimus Ter��n-Valdez & Guayasamin, 2009, P. altamnis Elmer & Cannatella, 2008, and P. kichwarum Elmer & Cannatella, 2008. P. daquilemai and P. aquilonaris have their upper eyelids with a conical tubercle and some smaller tubercles and colored pattern in inner surfaces of legs. Males of P. daquilemai, P. trachyblepharis and P. aquilonaris lacking vocal slits. All four species are easily differentiated from P. daquilemai by having a visible tympanum, which is not visible in P. daquilemai. Pristimantis minimus resembles the new species by its small size and by lacking an external visible tympanum. However, both species can be distinguished by the presence of a prominent rostral papilla at the tip of the snout in P. daquilemai (absent in P. minimus). Pristimantis aquilonaris (females SVL = 19.4���2.0 mm and males SVL = 13.7���17.6 mm), P. kichwarum (females SVL = 24.4���27.0 mm and males SVL = 17.5���21.9 mm) and P. altamnis (females SVL = 26.9���27.6 mm and females SVL = 16.9���19.9 mm) are larger than P. daquilemai (females SVL = 15.5���19 mm and males SVL = 12.0���15.0 mm). Pristimantis aquilonaris, P. kichwarum and P. altamnis have W scapular folds, while P. daquilemai has ������ ������ scapular folds. Pristimantis trachyblepharis present low ulnar tubercles and P. altamnis lack ulnar tubercles (conical and conspicuous in P. daquilemai). In P. trachyblepharis the rostral papilla is much less evident, no conspicuous colors on groin or other hidden surfaces, contrary to the new species. P. daquilemai has groin with orange or yellow spots, which may extend to the inner surfaces of legs and flanks, while P. aquilionaris has groin, anterior and posterior surfaces of thighs, concealed surfaces of tarsus, and axilla blackish brown with yellowish-orange or reddish-orange flecks (P. trachyblepharis, P. kichwarum and P. altamnis without colored pattern in inner surfaces). Other species closely related are in a clade including P. parvillus (Lynch, 1976), P. nietoi Arteaga, Pyron, Pe��afiel, Romero, Culebras, Bustamante, Y��nez-Mu��oz and Guayasamin, 2016, P. walkeri (Lynch, 1974), P. luteolatralis (Lynch, 1976) and P. ceuthospilus (Duellman & Wild, 1993). Pristimantis daquilemai and P. ceuthospilus have blotches on groins and inner surfaces of legs, P. ceuthospilus lacks scapular folds, evident rostral papilla and subacuminate snout (present in P. daquilemai), also bears evident tympanic annulus and membrane (completely covered by skin in P. daquilemai), and has larger SVL: females SVL = 25.1 mm, 23.5���26.7 mm, and males SVL = 22.4 mm, 19.0��� 25.8 mm (Pristimantis daquilemai: females SVL = 17.3�� 1.10 mm, 15.5���19 mm, males SVL = 13.19�� 0.87 mm, 12.0���15.0 mm). For the remaining species (P. parvillus, P. nietoi, P. luteolateralis and P. walkeri) the main characters that allow to differentiate P. daquilemai from these species are tympanic membrane not differentiated from surrounding skin, tympanic annulus not visible externally and presence of an elongated conical tubercle on the upper eyelid (tympanic annulus evident and lack of elongated conical tubercle on the upper eyelid in the aforementioned species). Another species that could be similar to P. daquilemai due to its small body size is P. andignomus Lehr & Coloma, 2008, however, the latter does not present orange-yellow spots in the groin, unlike P. daquilemai. Description of holotype. Adult female (16.8 mm SVL, Fig. 2���3), head as long as wide (40.6% of SVL); snout long (EN 15.6 % of SVL, eye-nostril distance equal to eye diameter), subacuminate in dorsal view, rounded with a prominent rostral papilla on the tip in profile (Fig. 3); nostrils slightly protuberant, directed dorsolaterally (Fig. 3); canthus rostralis weakly concave in dorsal and lateral view; loreal area slightly concave, lips rounded; upper eyelid with one enlarged conical tubercle surrounded by several smaller, conical and subconical tubercles (Fig. 3); interorbital area flat (EW 80% IOD); cranial crests absent; tympanic membrane not differentiated from surrounding skin, tympanic annulus not visible externally, hidden by muscle and skin; supratympanic fold weak, conspicuous conical or subconical postrictal tubercles. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomer small, ovoid, separated, posteromedial to choanae, with 3 teeth. Tongue longer than wide, notched posteriorly, posterior 30% not adhered to mouth floor. Dorsal skin shagreen with scattered small tubercles, ������ ������-shaped scapular fold present, with two subconical tubercles on medial and posterior sections of folds, thin dorsolateral folds extending from scapular folds to the post-sacral region; flanks densely tuberculated; skin on chest and venter areolate; discoidal fold weakly defined; cloacal region with two large rounded tubercles. Two conical outer ulnar tubercles; outer palmar tubercle bifid, approximately two times the size of elongate inner palmar tubercle; supernumerary tubercles present; subarticular tubercles well defined, rounded in lateral view. Fingers with narrow lateral fringes; Finger I shorter than Finger II; emarginated discs of fingers broadly expanded, most prominent on Fingers II���IV, about 2�� the size of the digit; discs rounded terminally, all with ventral pads well defined by circumferential grooves (Fig. 5). Hind limbs slender (TL 56.3% of SVL; FL 43.7% of SVL); heel with one conical tubercle, inner edge of tarsus with two or three subconical tubercles; inner metatarsal fold indistinct; toes with narrow lateral fringes, with basal membrane, distinctive between toes IV and V; subarticular tubercles round; inner metatarsal tubercle ovoid, about four times the size of subconical outer metatarsal tubercle; plantar supernumerary tubercles present (Fig. 5); discs of toes expanded in all discs, slightly narrower than those on fingers; toes with ventral pads well-defined by circumferential grooves; Toe V longer than Toe III, disc of Toe V not reaching the base of distal subarticular tubercle on Toe IV (Fig. 5). Coloration of holotype in preservative. Dorsum grayish-cream with irregular brown blotches, head with post-occipital W- shaped cream mark, ������ ������-shaped scapulars black marks; flanks and dorsal surfaces of the limbs light brown with transversal bars dark brown; dorsal surfaces of hands and feet thickly dotted with grayish-brown; throat and venter cream with scattered irregular dark brown blotches, aggregated in the throat; little white spots concentered in the venter; groins and inner surfaces of thighs with pale pink blotches delimited by dark brown, extending towards the flanks; ventral surfaces of limbs cream with dark brown blotches; outer edge of the tarsus with a dark brown longitudinal stripe; dark brown cloacal mark (Fig. 2, 7). Coloration of holotype in life. Dorsum light brown with irregular dark brown blotches, dorsolateral folds orange, head with ������ ������-shaped black scapular folds delimited by orange; flanks with brown bars separated by greenish-yellow, scattered tubercles pigmented with orange; venter light brown with minute dark brown spots and white small low warts, throat light brown with scattered irregular dark brown blotches; ventral surfaces of limbs light brown with minute dark brown and orange spots, ventral surfaces of hands and feet light brown, supernumerary and subarticular tubercles pigmented with orange; groin and inner surfaces of thighs and calves with distinctive orange blotches; snout light brown with weakly defined light brown subocular bars; golden iris with black reticulations and a horizontal reddish brown bar; ()-shaped dark brown cloacal mark (Fig. 6). Measurements (in mm) of holotype: SVL= 16; E���N= 2.5; HL= 6.5; HW= 6.5; IOD= 2.5; IND= 2.5; TL= 9; FL= 7; HL= 45; ED= 2.5; EW= 2.0. Variations. Measurement ranges and proportions are showed in Table 2. Snout-vent length in females from 15.5 to 19.0 mm and males from 12.0 to 15.0 mm. Tubercles on the scapular fold and dorsolateral folds visible in life and reduced in preservative. Dorsum varies from cream (e.g., ZSFQ 1211), grayish cream (e.g., ZSFQ 1201), grayish-brown (e.g., ZSFQ 1206), gray (e.g., ZSFQ 1204), light brown (e.g., QCAZ 30858) to dark brown (e.g., ZSFQ 1212), with irregular dark brown marks. Males with black ������ ������-shaped scapular folds except one (DHMECN 13071) in which they are depigmented. In addition, male QCAZ 30855 has a dorsal dark brown stripe, and the male DHMECN 13059 has a dark brown facial mask extending from the nostril to the scapular region. Males with weakly defined white nuptial pads on dorsum of Finger I. Females are more variable; some of them with a dorsal grayish-brown stripe delimited by dark brown (e.g., DHMECN 13055), others with ��������-shaped grayish-brown mark in the sacral region (e.g., ZSFQ 1201). Female QCAZ 30852 has the same facial mask as male DHMECN 13059. Meanwhile, the female QCAZ 30863 has a longitudinal brown stripe on dorsum and a gray helmet-shaped mark in the post occipital region. All females have black scapular folds except QCAZ 30858 in which they are semi pigmented. Ventral coloration varies from cream (e.g., ZSFQ 1211), grayish cream (e.g., ZSFQ 1204) to light brown (e.g., QCAZ 30852). Ventral pattern varies from dotted with dark brown (e.g., DHMECN 13071), to densely mottled (e.g ZSFQ 1204, QCAZ 30852). In preservative, all females have groins with pale pink marks delimited by dark brown. These marks are inconspicuous in males (Fig. 7; Fig. 8). Distribution and natural history. This species is known from three localities at elevations between 1554 and 2067 m, on the Cordillera del Condor, southern Ecuador. All localities are geopolitically in the Paquisha county, province of Zamora-Chinchipe. The ecosystem at type locality belongs to Montane Evergreen Forest of the Cordilleras del C��ndor and Kutuk�� (BsMa02) (MAE, 2013). It is considered a peculiar dwarf forest having plant species of foothills and lower montane forest of the same mountain range but smaller in size. This ecosystem shows a high abundance of epiphytes, bryophytes, and leaflitter, understory is dominated by Chusquea and canopy reach up to 12 m (Morales et al. 2013). Pristimantis daquilemai was found active at night on leaflitter and in low vegetation, up to 0.6 m above ground. Stomach contents of specimen ZSFQ 1203 consisted of one ant, subfamily Dolichoderinae (Hymenoptera), one wasp (Hymenoptera), and remnants of one fly (Diptera). Syntopic anurans included P. quaquaversus, P. ledzeppelin, and four unidentified species of Pristimantis. Extinction risk We follow IUCN criteria (G��rdenfors, Hilton-Taylor, Mace, & Rodr��guez, 2001). All known records of Pristimantis daquilemai are from the Cordillera del Condor. Habitat change and loss across this mountain range are increasing due to mining activity and two of the three known localities are within mining concessions. These concessions are already registered and are in an active process of exploration and, based on their practices, correspond to largescale mining concessions (IGF 2019). We make the following extinction risk assessment for P. daquilemai: (1) We suspect that P. daquilemai will suffer population declines due to continuous decline in habitat quality across its distribution, however, we refrain from use criterion A until more ecological data are available. (2) P. daquilemai has an estimated extent of occurrence EOO of 35.3 km 2, within the threshold for Critically Endangered (P. daquilemai may be relatively high and we propose that it should be classified under the IUCN threatened category Endangered (EN), based on criteria B1ab(iii) + 2ab(iii). The species have not been found in protected areas, but it may inhabit El Quimi, El Zarza, and El Condor biological reserves, in Cordillera del Condor. Other areas in the region are protected forests, but they are not national protected areas and receive little or no real protection (Prieto-Albuja et al. 2019). Research and monitoring actions should be established to study the ecology of P. daquilemai and other endemic species of the Cordillera del Condor., Published as part of Brito-Zapata, David, Reyes-Puig, Carolina, Cisneros-Heredia, Diego, Zumel, Daniel & Ron, Santiago R., 2021, Description of a new minute frog of the genus Pristimantis (Anura: Strabomantidae) from Cordillera del Condor, Ecuador, pp. 351-372 in Zootaxa 5072 (4) on pages 355-365, DOI: 10.11646/zootaxa.5072.4.3, http://zenodo.org/record/5748784, {"references":["Ibarra, H. G. R. (2018) La rebelion de Daquilema (Yaruquies-Chimborazo, 1871). Coleccion Testigos de la Historia. Cooperativa Fernando Daquilema, Instituto Nacional de Patrimonio Cultural, Quito, 84 pp.","Lopez-Ocon, L. (1986) Etnogenesis y rebeldia andina: la sublevacion de Fernando Daquilema en la provincia del Chimborazo en 1871. Boletin americanista, 36, 113 - 133.","Mutlak, N. D. (2009) Daquilema, Fernando (d. 1872) and the 1871 uprising, Ecuador. Available from: https: // doi. org / 10.1002 / 9781405198073. wbierp 0437 (accessed 16 November 2021)","Boulenger, G. A. (1918) Descriptions of new South American batrachians. Annals and Magazine of Natural History, Series 9, 427 - 433.","Elmer, K. R. & Cannatella, D. C. (2008) Three new species of leaflitter frogs from the upper Amazon forests: cryptic diversity within Pristimantis \" ockendeni \" (Anura: Strabomantidae) in Ecuador. Zootaxa, 1784 (1), 11 - 38. https: // doi. org / 10.11646 / zootaxa. 1784.1.2","Lynch, J. D. (1976) New species of frogs (Leptodactylidae: Eleutherodactylus) from the Pacific versant of Ecuador. Occasional Papers of the Museum of Natural History, University of Kansas, 55, 1 - 33.","Lynch, J. D. (1974) A new species of Eleutherodactylus (Amphibia: Leptodactylidae) from the Pacific lowlands of Ecuador. Proceedings of the Biological Society of Washington, 87, 381 - 388.","Duellman, W. E. & Wild, E. R. (1993) Anuran amphibians from the Cordillera de Huancabamba, northern Peru: Systematics, ecology, and biogeography. Occasional Papers of the Museum of Natural History, University of Kansas, 157, 1 - 53.","Lehr, E. & Coloma, L. A. (2008) A minute new ecuadorian Andean frog (Anura: Strabomantidae, Pristimantis). Herpetologica, 64, 354 - 367.","Lehr, E., Aguilar, C., Siu-Ting, K. & Carlos Jordan, J. (2007) Three New Species of Pristimantis (Anura: Leptodactylidae) from the Cordillera De Huancabamba in Northern Peru. Herpetologica, 63 (4), 519 - 536. https: // doi. org / 10.1655 / 0018 - 0831 (2007) 63 [519: TNSOPA] 2.0. CO; 2","Lynch, J. D. & Duellman, W. E. (1980) The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). The University of Kansas, Museum of Natural History, Miscellaneous Publications, 69, 1 - 86.","Morales, C., Jadan, O. & Aguirre, Z. (2013) Bosque siempreverde montano de las cordilleras del Condor-Kutuku. In: Sistema de Clasificacion de Ecosistemas del Ecuador Continental, Quito, 2013, pp. 205 - 206."]}

We describe a new species of Pristimantis from southern Ecuador, province of Zamora Chinchipe. The new species is closely related to an undescribed species of Pristimantis from Reserva Tapichalaca, Ecuador and with species of a clade historically assigned to the P. unistrigatus species group, such as P. parvillus, P. luteolateralis, P. walkeri, among others. The new species of Pristimantis is a miniaturized new frog (females 17.1±1.1 mm; males 13.2±0.9 mm), characterized by the presence of “› ‹”-shaped scapular folds, with two subconical tubercles on the medial and posterior regions of folds; tympanic membrane and tympanic annulus present but not externally visible; a prominent rostral papilla present; upper eyelid with one elongated conical tubercle; a conical tubercle on heels; groin with orange or yellow spots. The new species of Pristimantis is distributed in a restricted area in the Cordillera del Condor, a highly-diverse mountain range threatened by multiple anthropogenic activities. We recommend assigning the new species to the Endangered IUCN threatened category because it is only known from three nearby localities within mining concessions.

First record of the vocalizations of the anuran species Ecnomiohyla sukia (Hylidae) and Craugastor megacephalus (Craugastoridae) in Costa Rica.