Systematics of minute strabomantid frogs allocated to the genus Noblella (Amphibia: Anura) with description of a new genus, seven new species, and insights into historical biogeography Author Ortega, Jhael A. Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Apartado 17 - 01 - 2184, Quito, Ecuador Author Cisneros-Heredia, Diego F. Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Instituto de Biodiversidad Tropical IBIOTROP, Museo de Zoología / Laboratorio de Zoología Terrestre, Quito, Ecuador & Instituto Nacional de Biodiversidad INABIO, División de Herpetología, Quito, Ecuador Author Camper, Jeffrey D. Department of Biology, Francis Marion University, Florence, SC 29506, USA Author Romero-Carvajal, Andrés Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Apartado 17 - 01 - 2184, Quito, Ecuador Author Negrete, Leonardo Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Apartado 17 - 01 - 2184, Quito, Ecuador Author Ron, Santiago R. Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Apartado 17 - 01 - 2184, Quito, Ecuador santiago.r.ron@gmail.com text Zoological Journal of the Linnean Society 2025 2025-01-13 203 1 1 60 https://doi.org/10.1093/zoolinnean/zlae162 journal article 10.1093/zoolinnean/zlae162 0024-4082 14832689 7-01-2184 Phyllonastes ecuadoriensis sp.nov. LSID: urn:lsid:zoobank.org:act: 2F984FAA-F04B-4623-9E30- 7E29E32D659D Noblella lochites — Hedges et al. (2008) . Noblella lochites — Padial et al. (2014) . Noblella lochites — Catenazzi and Ttito (2016) . Noblella lochites — Reyes-Puig et al. (2019) . Noblella lochites — Santa-Cruz et al. (2019) . Noblella lochites — Condori et al. (2020) . Noblella lochites — Reyes-Puig et al. (2020) . Noblella lochites — Catenazzi et al. (2020) . Noblella lochites — Motta et al. (2021) . Noblella lochites — Reyes-Puig et al. (2021) . Noblella lochites — Portik et al. (2023) . Holotype ( Figs 12 , 13 , 14A , 15A ): QCAZ 57064 (field no. SC-PUCE 45767) adult female collected in Republic of Ecuador , province of Napo , Archidona canton , Pacto Sumaco parish, Wildsumaco Wildlife Sanctuary , Las Cascadas trail ( 0.6847°S , 77.6002°W ), 1417 m a.s.l. , by Fernando Ayala , Paloma Lima , Nadia Páez , Javier Pinto , and Santiago R. Ron on 3 April 2014 . Paratypes (N = 4; Fig. 12 ): All collected in Ecuador . Province of Napo : QCAZ 48916 subadult female , province of Napo, Wildsumaco Wildlife Sanctuary , Benavides trail ( 0.67744°S , 77.60133°W ), 1465 m a.s.l. , Jeffrey Camper and D. J. Zart , 15 July 2010 ; ZSFQ 346–347 , adult females , ZSFQ 348 , adult male , Archidona canton , Reserva Narupa ( 0.682°S , 77,74°W ), 1150– 1170 m a.s.l. , David Brito-Zapata , Jose Vieira , Malki Bustos , 21 June 2018 . Proposed standard English name: Ecuadorian leaflitter frog. Proposed standard Spanish name: Cutín de Hojarasca ecuatoriano. Definition ( Figs 12 , 13 , 14A , 15A ; Tables 2 and 3 ): We assign the new species to the genus Phyllonastes based on its phylogenetic relationships. The new species is characterized by: (1) skin on dorsum and flanks shagreen, dorsolateral folds absent, ventral surfaces smooth, discoidal fold present; (2) tympanic membrane and tympanic annulus distinct, supratympanic fold present, postrictal tubercles present or absent; (3) snout truncate in dorsal view and rounded in lateral view; (4) upper eyelid without tubercles, cranial crests absent; (5) vomerine teeth absent; (6) vocal slits present, nuptial pads absent; (7) fingers slightly expanded distally, with fingertips acuminate, bearing papillae, finger I shorter than finger II, supernumerary tubercles absent; (8) fingers without lateral fringes or circumferential grooves; (9) distal phalanges blunt or T-shaped, phalangeal formula of hand 2-2-3-2; (10) ulnar tubercles absent; (11) knees and heels without tubercles, outer edge of tarsus without tubercles, inner edge of tarsus bearing one subconical tubercle; (12) inner metatarsal tubercle large, elongate in ventral view and rounded in lateral view, bigger than knob of outer metatarsal tubercle; (13) toes expanded distally, with acuminate toe tips and bearing papillae, supernumerary tubercles present, toes bearing narrow lateral fringes, toe basal webbing absent, all toes bearing ill-defined pads and circumferential grooves, toe V shorter than toe III; (14) in life, dorsum brown, with faint dark marks and chevrons in some specimens, throat brown, contrasting with paler belly, dark mask extending onto flanks; and (15) SVL in adult males 14.0 mm ( N = 1) and SVL in adult females 16.76 mm ( N = 3) ( Table 4 ). Diagnosis: Phyllonastes ecuadoriensis resembles the closely related P. coloma and P. personinus by the presence of a tympanic membrane and supratympanic fold, acuminate fingertips and toe tips, and dark brown mask. It differs from them by having only two phalanges in finger IV ( P. coloma and P. personinus bear three phalanges in finger IV), truncate snout in dorsal view (round in P. coloma and P. personinus ), fingers slightly expanded distally (fingers not expanded in P. coloma nor P. personinus ), and toes expanded distally (toes slightly expanded in P. coloma nor P. personinus ); by the absence of supernumerary palmar tubercles (present in P. coloma and P. personinus ) and by the presence of supernumerary plantar tubercles (absent in P. coloma and P. personinus ). Phyllonastes ecuadoriensis also differs from P. coloma by the presence of papillae on fingertips and toe tips (both absent in P. coloma ), and from P. personinus by having dorsum and flanks shagreen (dorsum smooth with few low pustules and flanks smooth in P. personinus ). For a comparison with other Phyllonastes species that are more phylogenetically distant, see Tables 2 and 3 . Figure 12. Live and preserved specimens of Phyllonastes ecuadoriensis . Lateral, dorsal, and ventral views of: A, B, holotype QCAZ 57064 , adult female, SVL = 18.07 mm; and C, paratype QCAZ 48916, subadult female. A, in life. B, C, in preservative. A scale bar is given only for preserved adults. The depigmentation visible in both specimens is attributable to preservation defects. Figure 13. Palmar (A) and plantar (B) surfaces of Phyllonastes ecuadoriensis . Photographs of left hand and right foot of the holotype QCAZ 57064 . Description of holotype ( Figs 12 , 13 ): Adult female ( QCAZ 57064 ). Measurements (in millimetres): SVL, 18.1; tibia length, 9.3; foot length, 8.1; head length, 5.8; head width, 6.2; eye diameter, 2.2; tympanum diameter, 1.3; interorbital distance, 2.7; upper eyelid width, 1.5; internarial distance, 2.09; eye–nostril distance, 1.6. Body robust, head is not distinct from body, head slightly wider than long. Snout truncate in dorsal view and rounded in lateral view, without rostral papilla; canthus rostralis slightly concave; interorbital space flat, with no cranial crests; tympanic membrane distinct, tympanic annulus prominent on anterior and ventral borders, and a broad and low supratympanic fold resembling a bulge is present dorsally; postrictal tubercles absent; vomerine teeth absent; tongue large, lanceolate, and with a straight posterior border; vocal slits and nuptial pads absent; lips are not flared; nostrils are elongate and slightly protuberant. Skin of dorsum and flanks shagreen, top of the head and upper eyelid smooth; dorsolateral folds absent; ventral surfaces smooth; skin in cloacal region areolate; discoidal fold present. Ulnar tubercles absent, palmar tubercles distinct, low; outer palmar tubercle elongate, proximally wider and about twice the size of the elongate thenar tubercle; subarticular tubercles ill-defined, elongate, and low; supernumerary tubercles absent; dermal fringes on fingers absent; discs narrow, bearing round and inconspicuous pads without circumferential grooves. Phalangeal formula is 2-2-3-2 and relative length of fingers is I <II <IV <III. Fingers are slightly expanded distally, with acuminate tips that are dorsoventrally flattened, papilla present, and antebrachial ornamentation absent. Hindlimbs robust; knee and heel without tubercles; tarsal folds (inner and outer) absent. Outer tarsal tubercles absent; inner tarsal tubercle present, prominent, round in ventral view and subconical in lateral view, and about half the size of the outer metatarsal tubercle and in line with the inner metatarsal tubercle. Inner metatarsal tubercle elongate, ~1.5 × length of outer metatarsal tubercle. The latter is knob like and protrudes posteriorly. Subarticular tubercles low and rounded, supernumerary tubercles minute and ill defined. Toes with narrow lateral dermal fringes; basal webbing between toes absent; discs on toes ill defined, toes expanded distally bearing papilla; ill-defined circumferential grooves; relative lengths of toes I <II <V <III <IV; toe III longer than toe V (toe III reaches the distal border of the second subarticular tubercle of toe IV; toe V almost reaches the half of the second tubercle of toe IV). Colour of holotype in life (based on digital photographs) ( Fig. 12 ): Dorsum is light greyish orange, becoming more orange towards the cloacal region, bearing dark brown blotches fading posteriorly and a pair of dark sacral lateral patches that do not touch medially. Facial mask dark brown, almost black; the mask extends posteriorly on the flanks halfway to the hindlimb insertion; the dark facial pigment extends dorsally to the eyelid; posterior to the mask extension, flanks are medium brown with light flecking. Lips dark brown, with few irregular white flecks. Forearms are brown, with two pale bands on the left and 1.5 on the right. Dorsal surfaces of hindlimbs orange–brown, with dark brown transversal bars bordered by lighter colour. Throat uniformly dark brown, with an orange flecking anteriorly. Ventral surfaces of upper arms, thighs, and belly bright orange, with scattered dark brown flecks. Ventral surfaces of forearms and shanks dark brown, with small bright orange blotches. Cloacal region dark brown. Figure 14. Skulls of Phyllonastes species. A , Phyllonastes ecuadoriensis QCAZ 57064 . B, Phyllonastes heyeri QCAZ 24875. C, Phyllonastes macuma QCAZ 40180 . Figure 15. Skeletons of Phyllonastes species. A , Phyllonastes ecuadoriensis QCAZ 57064 , SVL = 18.07 mm. B, Phyllonastes heyeri QCAZ 24875, SVL = 14.11 mm. C, Phyllonastes macuma QCAZ 40180 , SVL = 11.92 mm. D, Phyllonastes myrmecoides QCAZ 53583, SVL = 11.13 mm. Colour of holotype in preservative ( Fig. 12 ): Dorsal surfaces dark brown; dorsum bearing a pair of darker sacral lateral patches; limbs with darker transversal bars. Throat, ventral surfaces of forearms, shanks, and feet dark brown, bearing cream flecks densely grouped on the posterior half of shanks. Chest, belly, and ventral surfaces of upper arms and thighs cream, with faint dark brown flecks. Posterior surfaces of thighs dark brown. Cloacal region dark brown, almost black. Variation ( Fig. 12 ): In this section, traits refer to preserved individuals unless otherwise mentioned. Tympanic annulus can be completely differentiated as in QCAZ 48916. The same specimen also presents an inconspicuous supratympanic fold (might be a preservation effect), dorsal surfaces dark brown, with white big irregular blotches in the lower back (might be only an effect of preservation of the individual), limbs without differentiated transversal bars, ventral surfaces of forearms, shanks, and posterior surfaces of thighs cream, bearing dark brown flecks densely grouped. Three specimens have a row of postrictal tubercles (ZSFQ 346–48). One male (ZSFQ 348) has a much paler throat, similar in colour to the venter. Three specimens have the suprainguinal spots elongated (ZSFQ 346–48), two of them merging into an hourglass-shaped large mark on dorsum (ZSFQ 347–48). One specimen (ZSFQ 347) has short papillae on most fingers, except for fingers III and IV on the left hand, which show long papillae. Two specimens (ZSFQ 346 and 348) have short papillae in all their fingers, sometimes ill defined. The male (ZSFQ 348) has vocal slits. Morphometric variation is detailed in Table 4 . Distribution, natural history, and conservation status ( Fig. 3 ): Phyllonastes ecuadoriensis is known from three localities on the eastern slopes of the Andes, in the provinces of Napo and Pastaza , between 1150 and 1465 m a.s.l. in Eastern Foothill Forest. Individuals were collected at night (20:00–00:45 h) on leaf litter in terra firme forest. We propose assigning P. ecuadoriensis to the Data Deficient IUCN Red List Category owing to the lack of information about its population size and geographical range. Etymology: The specific epithet refers to the country where the species is distributed, ecuadoriensis , meaning from Ecuador , and is masculine in gender. Remarks: The identification of P. lochites in our study is based on the redescription provided by Harvey et al. (2013) . Our phylogeny includes specimens from that publication (QCAZ51766/ MEPN14255 and QCAZ51767/MEPN14253). According to Harvey’s work and our phylogenetic results, the specimen KU177356, previously identified as P. lochites (e.g. Hedges et al. 2008 , Catenazzi and Ttito 2016 , Reyes-Puig et al. 2019 , 2020 , 2021 , Santa-Cruz et al. 2019 , Catenazzi et al. 2020 , Condori et al. 2020 , Portik et al. 2023 ), is, in fact, P. ecuadoriensis .