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Pristimantis sagedunneae, P. paladines, P. numbala P. Székely, D. Székely, Armijos-Ojeda, Hualpa-Vega & Vörös, 2023 |
Abstract
We review the species content of the Pristimantis orestes group and, by integrating genetic and morphological data, we distinguish and describe three new cryptic species from the subpáramos of southern Ecuadorian Andes. Genetically, these new species are most related to P. matildae and the other species of the P. colodactylus subgroup; however, in morphology, they are strikingly different from them, especially in terms of the shape of the body, having less-compressed heads and bodies. Additionally, they differ by their ecology, having arboreal habits in contrast with their bromeliad specialist sister species of the subgroup. The new species are also easily distinguished from the species of the P. orestes and P. simonbolivari subgroups, mainly by the lack of tympanum, longer snouts, and longer fingers and toes, and coloration. Pristimantis sagedunneae has a shagreen dorsum, subacuminate to rounded snout in dorsal view, Toe V longer than Toe III, and a brownish gray coloration of the dorsum with blackish or dark brown bars and an intense red coloration on the dorsal surfaces of thighs, groin, and concealed limb surfaces; P. paladines has a tuberculate dorsum, acuminate to subacuminate snout, Toe V longer than Toe III, and dorsum of various shades of brown, reddish brown, greenish brown, green, or reddish orange with dark, blackish bars and orange or reddish brown coloration on the dorsal surfaces of thighs, groin, and concealed limb surfaces; P. numbala has a shagreen dorsum, rounded to broadly rounded snout, Toe V much longer than Toe III, reddish brown coloration of the dorsum with dark brown bars and reddish brown coloration on the dorsal surfaces of thighs, groin, and concealed limb surfaces. All three species have very small distribution ranges of less than 20 km2 located in the western limit of Parque Nacional Podocarpus, at altitudes between 2,800–3,100 m. Finally, we briefly discuss the current species composition of the group, the role that natural history might have in determining the shape of the bodies, and the cryptic diversity and distribution pattern of some of the species in southern Ecuador.
KEYWORDS: Abra de Zamora, amphibians, direct-developing frogs, DNA, Reserva Numbala, Reserva Tapichalaca, tropical Andes
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| Color variation in life for individuals of Pristimantis sagedunneae, in dorsolateral or lateral, dorsal, and ventral views. (A, D, G) MUTPL 597, SVL = 21.7 mm, adult female paratype from the Cajanuma entrance to the Parque Nacional Podocarpus, Loja Province, 2,991 m; (B, E, H) MUTPL 1095, SVL = 23.8 mm, adult female paratype from Abra de Zamora, Loja Province, 2,812 m; (C, F, I) MUTPL 1096, SVL = 14.7 mm, juvenile paratype from Abra de Zamora, 2,812 m. |
Pristimantis sagedunneae sp. nov.
Common English name.—Sage Dunne's Rain Frog.
Common Spanish name.—Cutín de Sage Dunne.
Diagnosis.—We assign this species to Pristimantis based on phylogenetic evidence (Fig. 3) and on the general morphological similarity to other members of the genus. Pristimantis sagedunneae is a medium-sized species (among the P. orestes group; Table 2), distinguished by the following combination of traits: (1) skin on dorsum shagreen with some scattered tubercles (feature more evident in life); skin on venter areolate; discoidal fold present; dorsolateral folds absent; flanks usually with longitudinal lateral folds on anterior half; low middorsal fold present; (2) tympanic annulus and tympanic membrane absent; supratympanic fold present; (3) snout subacuminate to rounded in dorsal view, rounded in profile; canthus rostralis concave in dorsal view, rounded in profile; (4) upper eyelid bearing one larger tubercle and several small tubercles (feature more evident in life), it's width about 68% of interorbital distance; cranial crests absent; (5) dentigerous processes of vomers concealed in buccal mucosa; each process bearing three to five teeth; (6) condition of vocal sacs, vocal slits, and nuptial pads unknown; (7) Finger I shorter than Finger II; discs on fingers broadly expanded, truncate; circumferential grooves present; (8) fingers bearing lateral fringes; subarticular tubercles prominent; hyperdistal subarticular tubercles present; supernumerary palmar tubercles present; palmar tubercle divided into a larger (inner) and a smaller (outer) tubercle(s); thenar tubercle elliptical, same size with the inner palmar tubercle; (9) ulnar tubercles present (feature more evident in life); (10) heel with one to three large tubercles and several small tubercles (feature more evident in life); outer edge of tarsus with row of small tubercles (feature more evident in life); inner tarsal fold present; (11) inner metatarsal tubercle broadly ovoid, about 3× to 4× the size of subconical (in profile) outer metatarsal tubercle; subarticular tubercles prominent; hyperdistal subarticular tubercles present; supernumerary plantar tubercles present; (12) toes bearing broad lateral fringes; webbing basal; Toe V longer than Toe III; discs on toes broadly expanded, truncate, about same size as those on fingers; circumferential grooves present; (13) in life, dorsum brownish gray, with blackish or dark brown bars that on the middle of the back conjoin in >X or XXX-like shapes; flanks, dorsal surfaces of arms, and of hindlimbs with dark transverse bars; large areas of dorsal surfaces of thighs, groin, and concealed limb surfaces red or intense red; head with a wide, black-bordered light interorbital bar, dark labial bars, and dark supratympanic stripes; venter and ventral surfaces of hindlimbs and arms pinkish white or pinkish gray, with or without dark markings on the throat; iris bronze with black reticulations and a median, wide, horizontal dark red streak; (14) SVL 21.4–24.3 mm in adult females (22.5 ± 1.41, n = 5); males unknown.
Etymology.—The specific name sagedunneae is a noun in the genitive case and honors Sage Dunne, in recognition of her passion for Andean wildlife and her family's invaluable support of conservation work in Ecuador. Of particular importance is their contribution to the amphibian conservation in the Sangay-Podocarpus connectivity corridor, Ecuador's first ecological corridor, which protects 567,067 ha of high-elevation páramo grasslands and cloud forest ecosystems, as well as chains of lakes and wetlands, with unique biological diversity and endemism.
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Color variation in life for females of Pristimantis paladines, in lateral or dorsolateral, dorsal, and ventral views. Paratypes from Parque Nacional Podocarpus in the vicinity of Reserva Tapichalaca, Zamora Chinchipe Province, 2,880 m: (A, D, G) MUTPL 1151, SVL = 19.5 mm; (B, E, H) MUTPL 1160, SVL = 19.8 mm. Color variation in life for males of Pristimantis paladines, in lateral or dorsolateral, dorsal, and ventral views. Paratype from the Cerro Toledo sector of Parque Nacional Podocarpus, Loja Province, 3,080 m: (A, D, G) MUTPL 125, SVL = 16.7 mm. Paratypes from the Cerro Toledo sector of Parque Nacional Podocarpus, Zamora Chinchipe Province, 3,104 m: (B, E, H) MUTPL 1107, SVL = 18.0 mm. |
Pristimantis paladines sp. nov.
Common English name.—Paladines Rain Frog.
Common Spanish name.—Cutín de Paladines.
Diagnosis.—We assign this species to Pristimantis based on phylogenetic evidence (Fig. 3) and on the general morphological similarity to other members of the genus. Pristimantis paladines is a small species (among the P. orestes group; Table 2), distinguished by the following combination of traits: (1) skin on dorsum tuberculate with numerous scattered large tubercles (in life the tuberculated texture of the skin is more evident); skin on venter areolate; discoidal fold present; dorsolateral folds absent; flanks usually with longitudinal lateral folds on anterior half; middorsal fold evident, tall, usually with several larger tubercles scattered along its length (feature more evident in life); (2) tympanic annulus and tympanic membrane absent; supratympanic fold present; (3) snout acuminate to subacuminate in dorsal view, rounded in profile; canthus rostralis weakly concave to straight in dorsal view, rounded in profile; (4) upper eyelid bearing two to three large tubercles and several small tubercles (feature more evident in life), its width about 72% of interorbital distance in females and 81% of interorbital distance in males; cranial crests absent; (5) dentigerous processes of vomers inconspicuous, oblique, ovoid or triangular, separated medially by a distance smaller than the width of processes; each process bearing two to five teeth; (6) males without vocal sacs, vocal slits, and nuptial pads; (7) Finger I shorter than Finger II; discs on fingers broadly expanded, truncate; circumferential grooves present; (8) fingers bearing lateral fringes; subarticular tubercles prominent; hyperdistal subarticular tubercles present; supernumerary palmar tubercles present; palmar tubercle usually not divided or only partially divided into a larger (inner) and a smaller (outer) tubercle(s); thenar tubercle elliptical, larger than inner palmar tubercle; (9) ulnar tubercles present (feature more evident in life); (10) heel with two to three large tubercles and several small tubercles (feature more evident in life); outer edge of tarsus with row of large, conical tubercles (feature more evident in life); inner tarsal fold present; (11) inner metatarsal tubercle broadly ovoid, about 2× to 3× the size of subconical (in profile) outer metatarsal tubercle; subarticular tubercles prominent; hyperdistal subarticular tubercles present; supernumerary plantar tubercles present; (12) toes bearing broad lateral fringes; webbing basal; Toe V longer than Toe III; discs on toes broadly expanded, truncate, slightly smaller than those on fingers; circumferential grooves present; (13) in life, dorsum of various shades of brown, reddish brown, greenish brown, green or reddish orange with, dark, blackish bars, with or without X or W markings on the back; flanks, dorsal surfaces of arms and of hindlimbs with dark transverse bars; large areas of dorsal surfaces of thighs, groin, concealed limb surfaces and sometimes of dorsum orange or reddish brown; head usually with dark, blackish interorbital markings, dark labial bars and dark supratympanic stripes; venter and ventral surfaces of hindlimbs and arms pinkish white or pinkish gray, with or without dark markings on the throat; iris bronze with fine black reticulations and a median, wide, horizontal dark red streak; (14) SVL 19.5–21.4 mm in adult females (20.5 ± 0.82, n = 5) and 16.7–18.3 mm in adult males (17.4 ± 0.73, n = 5).
Etymology.—The specific name paladines is a noun in apposition and honors the Paladines family from the city of Loja, in particular Félix Humberto Paladines Paladines (1938–2022), for his valuable contribution to the academic and cultural fields and for safeguarding the history and identity of southern Ecuadorian people. In addition, it constitutes recognition of the remarkable work carried out by his children, Renzo, Bruno, Pedro, and Maria Gabriela, who created the nongovernmental organization Naturaleza y Cultura Internacional (NCI), a consolidated organization dedicated to the protection of wildlife in Latin America where it has managed to protect, to date, more than 8.5 million hectares of tropical forest in various countries.
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| Color variation in life for paratypes of Pristimantis numbala, in lateral or dorsolateral, dorsal, and ventral views. Adult females from Reserva Numbala, Zamora Chinchipe Province, 2,861 m: (A, D, G) MUTPL 1178, SVL = 23.27 mm; (B, E, H) MUTPL 1179, SVL = 20.4 mm. Juvenile from Reserva Numbala, Zamora Chinchipe Province, 2,875 m:(C, F, I) MUTPL 1191, SVL = 14.9 mm. |
Pristimantis numbala sp. nov.
Common English name.—Numbala Rain Frog.
Common Spanish name.—Cutín de Numbala.
Diagnosis.—We assign this species to Pristimantis based on phylogenetic evidence (Fig. 3) and on the general morphological similarity to other members of the genus. Pristimantis numbala is a medium-sized species (among the P. orestes group; Table 2), distinguished by the following combination of traits: (1) skin on dorsum shagreen with some scattered tubercles (feature more evident in life); skin on venter areolate; discoidal fold present; dorsolateral folds absent; flanks usually with longitudinal lateral folds on anterior half; low middorsal fold present (feature more evident in life); (2) tympanic annulus and tympanic membrane absent; supratympanic fold present; (3) snout rounded to broadly rounded in dorsal view, rounded in profile; canthus rostralis concave in dorsal view, rounded in profile; (4) upper eyelid bearing one larger tubercle and several small tubercles (feature more evident in life), it's width about 83% of interorbital distance; cranial crests absent; (5) dentigerous processes of vomers evident, oblique, ovoid, separated medially by distance equal or slightly smaller than the width of processes; each process bearing four to six teeth; (6) condition of vocal sacs, vocal slits, and nuptial pads unknown; (7) Finger I shorter than Finger II; discs on fingers broadly expanded, truncate; circumferential grooves present; (8) fingers bearing lateral fringes; subarticular tubercles prominent; hyperdistal subarticular tubercles present; supernumerary palmar tubercles present; palmar tubercle usually not divided or only partially divided into a larger (inner) and a smaller (outer) tubercle(s); thenar tubercle elliptical, larger than inner palmar tubercle; (9) ulnar tubercles present (feature more evident in life); (10) heel without or with one larger tubercle and several small tubercles (feature more evident in life); outer edge of tarsus with or without row of small tubercles (feature more evident in life); inner tarsal fold absent or present; (11) inner metatarsal tubercle broadly ovoid, about 2× to 3× the size of subconical (in profile) outer metatarsal tubercle; subarticular tubercles prominent; hyperdistal subarticular tubercles present; supernumerary plantar tubercles present; (12) toes bearing lateral fringes; webbing basal; Toe V much longer than Toe III; discs on toes broadly expanded, truncate, about same size or slightly smaller as those on fingers; circumferential grooves present; (13) in life, dorsum reddish brown, with dark brown bars and X markings on the back; flanks grey, pinkish grey, or green, with wide, dark transverse bars; dorsal surfaces of arms and of hindlimbs with dark transverse bars; large areas of dorsal surfaces of thighs, groin, and concealed limb surfaces reddish brown; head with black interorbital markings, dark labial bars and black supratympanic stripes; venter, throat, and ventral surfaces of hindlimbs and arms pinkish white or pinkish gray with blackish markings; iris bronze with fine black reticulations and a median, wide, horizontal dark red streak; (14) SVL 20.4–23.2 mm in adult females (21.9 ± 1.43, n = 3); males unknown.
Etymology.—The specific name numbala is a noun in apposition and refers to Reserva Numbala, an important private protected area managed by the NGO Naturaleza y Cultura Internacional. The reserve was established in 2006, with the main aim of conserving the last remnants of romerillo (Retrophyllum rospigliosii and Prumnopitys harmsiana, Podocarpaceae) forests, which are represented here by some of the last, giant trees of these species. The reserve protects 1,800 ha of subpáramo and montane cloud forest and is home to an important diversity of birds, amphibians, mammals, and plants. It is located between the two isolated extensions of the southern part of Parque Nacional Podocarpus, guaranteeing the connectivity needed for the preservation of the biological diversity of the national park and its area of influence.
Paul Székely, Diana Székely, Diego Armijos-Ojeda, Santiago Hualpa-Vega and Judit Vörös. 2023. Molecular and Morphological Assessment of Rain Frogs in the Pristimantis orestes Species Group (Amphibia: Anura: Strabomantidae) with the Description of Three New Cryptic Species from Southern Ecuador. Herpetological Monographs. 37(1); 41-69. DOI: 10.1655/HERPMONOGRAPHS-D-22-00002
www.natureandculture.org/directory/three-new-frog-species-discovered-in-southern-ecuador/
