Niceforonia philippi ( Jiménez de la Espada, 1875 ) Sánchez-Vialas & Miñarro & Padial & Riva, 2023

Sánchez-Vialas, Alberto, Miñarro, Marta, Padial, José M. & Riva, Ignacio De La, 2023, Taxonomic reassessment and nomenclatural status of Niceforonia dolops and Hylodes philippi (Amphibia: Anura: Strabomantidae), Zootaxa 5330 (1), pp. 117-125 : 119-122

Niceforonia philippi ( Jiménez de la Espada, 1875) , comb. nov.

Niceforonia dolops — Acosta-Galvis, Streicher, Manuelli, Cuddy & de Sá, 2018, Zootaxa, 4514: 494.

Hypodactylus dolops — Hedges, Duellman & Heinicke, 2008, Zootaxa, 1795: 67–68.

Isodactylus dolops — Hedges, Duellman & Heinicke, 2008, Zootaxa, 1737: 108.

Eleutherodactylus dolops Lynch & Duellman, 1980: 26 . Terra typica: “ Salto de Agua , 2.5 km NNE Río Reventador, Provincia Napo, Ecuador, 1660 m ”.

Hylodes verrucosus Jiménez de la Espada, 1875: 215 , plate 3, figures 1 and 1a (https://www.biodiversitylibrary.org/item/ 44709#page/225/mode/1up). Terra typica: originally not stated. Specimen label mentioned “San José de Moti”.

Hylodes philippi Jiménez de la Espada, 1875: 215 , plate 3, figures 2 and 2a (https://www.biodiversitylibrary.org/item/ 44709#page/225/mode/1up). Terra typica: originally not stated. Specimen label mentioned “San José de Moti”.

Diagnosis: (1) large size (maximum recorded: 57.6 mm); (2) dorsal skin pustulate, with conical and keratinized granules; no dorsolateral folds; ventral skin smooth ( Figure 2A–B ); (3) tympanic membrane and tympanic annulus distinct, about 1/3 (females and juveniles) and 1/2 (males) of eye length, supratympanic fold present ( Figures 3A, 3D ); (4) snout short, subacuminate in dorsal view ( Figures 2A, 2C ), round in lateral profile ( Figures 3A, 3A ); (5) cranial crests absent; upper eyelid broader than interocular distance ( Figure 2A ); (6) dentigerous process of vomers prominent, triangular, located posteromedial to choanae, bearing a transverse row of 8–13 vomerine teeth along posterior border ( Figure 4A ); (7) fingers long and slender, relative length of fingers III> IV = I> II; subarticular tubercles round, prominent, simple: a single one in fingers I and II, two in fingers III and IV; supernumerary tubercles absent; tips of fingers only slightly enlarged, round (as long as broad), with weak circumferential grooves; no lateral fringes; thenar tubercle large, elongated, slightly longer than palmar tubercle; palmar tubercle rounded, bifid, with the outer portion slightly larger than the inner one ( Figure 3B, 3E ); no ulnar tubercles; (8) toes long and slender, lacking lateral fringes and webbing; toe tips enlarged, round (specially enlarged in toes I, II, and III), with barely appreciable disc structure; no tubercles on heel or tarsus; supernumerary tubercles absent; subarticular tubercles longer than wide, prominent, simple: a single one in toes I and II, two in toes III and V, three in toe IV; two metatarsal tubercles, the inner elongate, prominent, and five times larger than the outer, which is relatively low and flat; relative length of toes IV> V> III> II> I ( Figure 3C, 3F ); (9) males with vocal slits; subgular vocal sac.

Niceforonia philippi differs from most of its congeneric species by lacking the general morphological appearance based on stocky bodies, short legs, and not expanded digital tips (“phrynopoid” ecomorph; see De la Riva, 2020). In contrast, N. philippi , presents a slender body shape, longer legs and digits, and larger size (reaching up to 57.6 mm) than those stocky-bodied congeneric species. The most morphologically similar species to N. philippi is N. babax ( Lynch, 1989) (appendix 1), the only other large species of the genus (maximum size recorded: 48.7 mm), which is included in the subgroup I or N. dolops species group fide Lynch (1989) and Lynch & Duellman (1997). The presence of very large nasal bones was considered a synapomorphy for this species group ( Lynch, 1989). Additionally, males of N. babax show vocal slits as in N. philippi ( Lynch, 1989; Lynch & Duellman, 1980). However, molecular data have never been employed to examine the monophyly of this group. The most conspicuous diagnostic traits to differentiate both species are detailed by Lynch (1989) as follows: N. babax presents low and flattened warts, sometimes forming dorsolateral ridges, whereas N. philippi (as N. dolops ) has the skin pustulated; digital discs on the hands more expanded in N. philippi than in N. babax . Niceforonia philippi and N. babax occur allopatrically, being N. babax (type locality: “Finca La Planada”, 5 km by road from caserio Chucunés, Municipio Ricaurte, Departamento de Nariño, Colombia ”) restricted to the pacific versants of the Andes of Ecuador and Colombia, at elevations between 1200 and 2200 m asl. Apart from N. philippi , only four out of the 14 species of Niceforonia occur in the Amazonian slopes of the Andes: N. nana Goin and Cochran, 1963 (recorded from Cordillera Oriental, Colombia), N. columbiana (Werner, 1899) [only known from the type locality at Monte Redondo, Buenavista, Departamento de Cundinamarca or Meta, Colombia; however, its taxonomic validity is questioned (see Acosta-Galvis, 2021)], N. nigrovittata (Andersson, 1945) (ranging across the Amazonian versants of the Andes from Peru, Ecuador and Colombia), and N. elassodiscus (Lynch, 1973) (distributed over the Amazonian slopes of the Andes from Sucumbíos and Napo in Ecuador to Pasto in Colombia) ( Acosta-Galvis et al. 2018). These species can readily distinguished from N. philippi as they are characterized by having phrynopoid morphology, small size, and the lack of vocal slits in males.

Variation: Males differ from females by the presence of vocal slits and by the size and shape of the tympanic annulus, rounded in males and relatively higher than long in females ( Lynch & Duellman, 1980).

The color pattern illustrated by Jiménez de la Espada (1875) is no longer distinguishable in the MNCN type specimens, but it is still well preserved in the types of Eleutherodactylus dolops . The types of both H. philippi and H. verrucosus showed a pattern of transversal dark bars on the hindlimbs and, to a lesser extent, on the forelimbs; they had irregular dark blotches on the dorsum, especially marked in the type of H. verrucosus ( Jiménez de la Espada, 1875) . The colouration pattern of Niceforonia dolops is congruent with that illustrated by Jiménez de la Espada (1875) for H. verrucosus and H. philippi . The ventral coloration of the type series of E. dolops shows a notable variability. Specimen KU 168811 presents a cream ventral surface with small brown spots, whereas the extension of the brown pigmentation on throat and chest in the other type specimens cover a great portion of the ventral surface, with dispersed cream dots (isolated or contacting with each other). The posterior half of the venter is uniformly cream, lacking dark spots in most of the specimens (KU 168811, KU 165866, KU 165867, KU 165868, KU 143505), except for KU 143504, which shows a reticulated pattern.

The types of H. philippi and H. verrucosus and those of E. dolops exhibit the same morphology and relative location of the vomerine processes, sharing a row of 8–13 vomerine teeth along the posterior margin of the vomerine process ( Figure 3 ). No marked differences between the well preserved types of H. verrucosus and E. dolops were found. The type of H. verrucosus , a female, lacks vocal slits and shows a small tympanic annulus, about 1/3 of the eye diameter, such as in the holotype of E. dolops . The holotype of H. verrucosus is smaller than that of E. dolops (48,9 mm vs. 57, 6 mm respectively).

Distribution: Niceforonia philippi is known from a few scattered localities across the eastern Andean foothills of northern Ecuador and southern Colombia at elevations of 940–1950 m ( Jiménez de la Espada, 1875; Lynch & Duellman, 1980; Ruiz-Carranza et al., 1996; Suárez-Mayorga, 1999; Mueses-Cisneros, 2005; Acosta Galvis, 2021) ( Figure 1 ).

Figures

FIGURE 1. Distribution map of Niceforonia philippi. Unambiguous records are represented by blue dots. San José de Mote and San José de Suno are depicted by an orange square and a black triangle, respectively. Dubious localities are not included.

FIGURE 2. Dorsal and ventral view of: A–B holotype of Eleutherodactylus dolops (KU143505) and, C–D holotype of Hylodes verrucosus (MNCN1599).

FIGURE 3. From top to bottom: lateral view of the head, palmar tubercles (left hand), and plantar tubercles (right and left foot, respectively) of:A–C holotype of Eleutherodactylus dolops (KU143505) and, D–F holotype Hylodes verrucosus (MNCN1599). The tip of the left fourth finger of the holotype of E. dolops is framed.

FIGURE 4. Mouth roof showing the vomerine teeth of: A holotype of Eleutherodactylus dolops (KU143505), B paratype of E. dolops (KU165868), C holotype of Hylodes verrucosus (MNCN1599) and D holotype of H. philippi (MNCN1600).