Research Article |
Corresponding author: Juan C. Sánchez-Nivicela ( juan.sanchezn13@gmail.com ) Academic editor: Angelica Crottini
© 2018 Juan C. Sánchez-Nivicela, Elvis Celi-Piedra, Valentina Posse-Sarmiento, Verónica L. Urgilés, Mario Yánez-Muñoz, Diego F. Cisneros-Heredia.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sánchez-Nivicela JC, Celi-Piedra E, Posse-Sarmiento V, Urgiles VL, Yánez-Muñoz M, Cisneros-Heredia DF (2018) A new species of Pristimantis (Anura, Craugastoridae) from the Cajas Massif, southern Ecuador. ZooKeys 751: 113-128. https://doi.org/10.3897/zookeys.751.20541
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A new species of Pristimantis is described from the highland paramos on the eastern slopes of the Cajas Massif, southern Andes of Ecuador, at 3400 m. This new species is characterized by having a distinctive reddish color, cutaneous macroglands in suprascapular region and surfaces of arm and legs, and by lacking dentigerous processes of vomers. The cutaneous macroglands are similar to those exhibited by several species of the Pristimantis orcesi group, and may suggest a close phylogenetic relationship. The new species could be a latitudinal substitution of Pristimantis orcesi in the southern Andes of Ecuador.
Describimos una nueva especie de Pristimantis desde las laderas orientales del macizo El Cajas en los páramos andinos del sur de Ecuador a 3400 m.s.n.m. Esta nueva especie tiene un color rojizo distintivo y se caracteriza por tener macroglándulas cutáneas en varias regiones del cuerpo, de la siguiente manera: la región supraescapular, las superficies del antebrazo, parte superior del brazo, las manos y el borde de las piernas. Además, carece de procesos dentígeros en los vomerinos. Las macroglándulas cutáneas son similares a las presentes en el grupo de Pristimantis orcesi, y podrían representar una posición filogenética cercana. La nueva especie puede constituir una sustitución latitudinal de Pristimantis orcesi en los Andes sur de Ecuador.
Andes, glandular frog, paramo, Pristimantis erythros sp. n., taxonomy, Terrarana
Andes, rana glandular, páramo, Pristimantis erythros sp. n., taxonomía, Terrarana
The Andes are one of the major physiographic features on our planet. A heterogeneous mountain system, three geographical separations have been identified in the Andes, based on their different geological, geographical, climatic, and ecosystemic characteristics: northern, central, and southern Andes (
Evolution of paramo biodiversity is strongly linked to orogeny and geomorphology, and complex and rich biotas are known to occur across the northern Andes due to their heterogeneous history and topography (
Although amphibian species richness decreases with higher altitude (
Collections were made at Chanlud hydroelectric project (Fig.
Definitions and terminology follows proposals by
English: Blood Rain Frog. Spanish: Cutín de Sangre
DHMECN 12103 (field series JCS.317); (Figs
DHMECN 12102,
Pristimantis erythros differs from other species of the genus by the combination of the following characters: (1) Skin on head and dorsum granular, flanks and venter areolate with low warts; dorsolateral folds absent; discoidal fold weakly defined; (2) tympanic membrane and annulus present and visible, rounded, ca. 50% of eye diameter, upper half covered by parotoid macrogland; (3) snout short, rounded in dorsal and lateral views; (4) upper eyelid without tubercles, interorbital distance wider than width of upper eyelid (40%); cranial crests absent; (5) dentigerous process of vomers absent; (6) vocal slits and sacs present in males, nuptial pads absent; (7) Finger I shorter than II; discs laterally expanded with dilated pads and narrow fringes, (8) fingers with coarse lateral cutaneous fringes; (9) low ulnar warts in ventral view; radioulnar macroglands covering the upper surfaces of forearm; (10) heel and tarsus lacking tubercles; paracnemid macroglands on upper surfaces of legs, tarsi, and Toes IV and V; (11) inner metatarsal tubercle oval, not prominent, twice as large as outer metatarsal tubercle, outer metatarsal tubercle rounded and low, supernumerary tubercles low and indistinct; Toe V longer than III, disc of Toe III reaches distal border of penultimate subarticular tubercle on Toe IV, disc on Toe V reaches distal border of distal subarticular tubercle on Toe IV; (12) toes with conspicuous lateral fringes, extend to base of fingers, webbing absent; toe pads as large as or slight larger than those on fingers; (13) in life, dorsum uniformly burgundy, red to orange-red (reddish brown to burgundy in preserved) ; flanks, posterior surfaces of legs, groin, throat and venter crimson (dark reddish brown in preserved); iris dark brown with thin golden reticulations; ventral surfaces of hands and feet pinkish cream; (14) SVL in adult females 38.8–42.6 mm (x̄ = 40.3, n = 4), in adult males 36.8–37.1 mm (x̄ = 36.7, n = 2).
(Fig.
Adult female (Fig.
Skin of dorsum granular without tubercles; dorsolateral folds absent; ventral surface areolate. Discoidal fold weakly defined in ventral view; cloacal region short, and covered by small and pronounced warts (Fig.
Hind limbs robust, tibia length 44% SVL; heel and external border of tarsus without tubercles, covered dorsally and ventrally by paracnemid macroglands; inner tarsal fold absent; inner metatarsal tubercles oval, twice as larger than the external metatarsal tubercle; supernumerary tubercles present, rounded and flattened; toes with lateral cutaneous fringes; basal membrane absent between toes; foot disks same size as those of hand, laterally expanded from fingers I–IV; relative length of toes 1<2<3<4>5; Toe IV larger than Toe III (Fig.
Snout-vent length 39.1; head length 10.8; head width 13.8; eye diameter 3.4; eye-nostril distance 3.5; interorbital distance 5.8; internarial distance 3.5; tympanum diameter 1.9; upper eyelid width 2.8; tibia length 17.5; foot length 20.7; hand length 14.5.
Dorsum dark red with slightly lighter shades on head and limbs; dark red on venter. Tips of fingers and toes pinkish cream in dorsal view; ventral surfaces of hands and feet, creamy pink. Iris homogeneously dark brown, with thin golden reticulations (Fig.
Dorsum reddish brown, flanks, posterior surfaces of thighs, venter, and throat dark reddish brown. Dorsal surfaces of fingers pinkish cream; ventral surfaces of hands and feet, creamy pink (Fig.
Morphometric variations of the type series are presented in Table
The specific epithet erythros is derived from the Greek word for red, in allusion to the distinctive coloration of this species.
Measurements (in mm) of the type series of Pristimantis erythros sp. n. All specimens are adults, range is followed by mean ± stander deviation in parentheses. Abbreviations: SVL = snout vent length, HL = head length, HW = head width, ED = eye diameter, EN = eye-nostril distance, IOD = interorbital distance, IND = internarinal distance, UEW = upper eyelid width; TD = tympanum diameter, HAL = hand length, Finger IV disk width = Fin4DW, TL = tibia length, FL = foot length, Toe IV disk width = Toe4DW.
Measurement | Adult Female | Adult Male |
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N = 4 | N = 2 | |
SVL | 38.8–42.6 (40.2 ± 1.7) | 36.7–37.0 (36.9 ± 0.2) |
EN | 2.7–3.5 (3.2 ± 0.3) | 3.1–3.4 (3.3 ± 0.2) |
HL | 10.6–13.7 (11.5 ± 1.4) | 11.5–11.8 (3.3 ± 0.2) |
HW | 13.2–14.7 (13.7 ± 0.6) | 12.6–13.3 (12.9 ± 0.5) |
IOD | 4.7–5.8 (5.1 ± 0.5) | 4.2–5.4 (4.8 ± 0.9) |
IND | 3.1–3.5 (3.3 ± 0.9) | 3.3–3.4 (3.3 ± 0.1) |
TL | 16.8–17.5 (17.1 ± 0.3) | 15.9–16.6 (16.3 ± 0.5) |
FL | 19.5–21.1 (20.2 ± 0.8) | 18.4–18.5 (18.4 ± 0.1) |
HAL | 13.0–14.4 (13.8 ± 0.6) | 12.3–12.8 (12.5 ± 0.4) |
TD | 1.7–1.9 (1.8 ± 0.1) | 1.7–1.8 (1.8 ± 0.1) |
ED | 3.4–4.2 (3.7 ± 0.4) | 2.9–3.8 (3.3 ± 0.7) |
UEW | 2.6–3.5 (3.0 ± 0.4) | 3.1–3.3 (3.2 ± 0.1) |
Fin4DW | 2.3–3.5 (2.6 ± 0.3) | 2.2–2.4 (2.3 ± 0.2) |
Toe4DW | 2.2–2.7 (2.5 ± 0.2) | 2.1–2.3 (2.2 ± 0.1) |
Pristimantis erythros is only known from its type locality in the Cajas Massif. The area is covered by paramos dominated by grassland and shrubs, between 3450 and 3500 m (Fig.
The Paramos on the Cajas Massif (221000 h. approx.) appear well preserved. Part of its extension includes the Cajas National Park (28544 h). However, the continued changes on land cover and land use occurring in several areas over the massif on the buffer area of the national park and not protected nearest regions are leading to habitat loss (
At least 50 species of anurans, including Pristimantis erythros, are currently known to inhabit the paramos of Ecuador (Table
Amphibians of the paramos from the Andes of Ecuador (above 3000 m). Abbreviations: N = northern section, S = southern section (sections are approximately divided by 1.5°S latitude). Nominal species that may be complexes (including more than one cryptic species) are marked with an asterisk.
Species | Cordillera Occidental | Cordillera Oriental | Source | ||
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N | S | N | S | ||
Atelopus bomolochos |
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A. exiguus |
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A. ignescens |
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A. nanay |
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A. pastuso |
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A. petersi |
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A. podocarpus |
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Osornophryne angel |
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O. antisana |
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O. talipes |
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Centrolene buckleyi * |
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Hypodactylus brunneus |
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H. peraccai |
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Lynchius flavomaculatus |
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Pristimantis buckleyi * |
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P. cajamarcensis |
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P. cryophilius * |
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P. cryptomelas |
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P. curtipes * |
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P. devillei * |
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P. erythros | This work | ||||
P. festae |
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P. gentryi |
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P. gualacenio |
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P. huicundo |
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P. leoni * |
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P. lymani |
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P. mazar |
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P. modipeplus |
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P. myersi |
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P. ocreatus |
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P. orcesi * |
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P. orestes * |
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P. ortizi |
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P. philipi | Lynch and Duellman (1995) | ||||
P. phoxocephalus * |
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P. pichincha |
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P. pycnodermis |
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P. riveti * |
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P. thymelensis |
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P. unistrigatus * |
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Hyloxalus anthracinus |
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H. jacobuspetersi |
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H. vertebralis |
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Gastrotheca espeletia |
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G. litonedis * |
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G. pseustes * |
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Hyloscirtus larinopygion |
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Telmatobius niger |
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T. vellardi |
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The Cajas Massif has one of the most particular landscapes in the Ecuadorian Andes. The massif was glaciated during the Pleistocene (
Pristimantis erythros share conspicuous cutaneous macroglands on its body and extremities with P. orcesi, P. pycnodermis, and P. loujosti. Pristimantis erythros is most similar to P. orcesi, from which it differs by its coloration and morphology, and has a significant biogeographic separation due to the isolation of the Cajas Massif from other paramos. Phylogenetic relationships of P. erythros are still uncertain, and due to the lack of additional evidence (e.g., molecular data), we refrain to assign P. erythros to any species group. Although P. erythros and P. orcesi may be related, the Pristimantis orcesi species-group is not a monophyletic group (
We express our gratitude to ETAPA and ElecAustro companies for facilities provided during our field work; to our good friend and colleague Cristian Nieves, for his participation during surveys; to Santiago Ron (Pontificia Universidad Católica de Quito, QCAZ) and Carolina Reyes-Puig (Instituto Nacional de Biodiversidad, INABIO) for access to review specimens under their care. Carolina Reyes-Puig provided photographic material of the holotype, Santiago Ron by P. orcesi photographs and Stalin Cáceres by P. loujosti photographs. JCSN thanks Lorena Orellana for her extreme patience. Work by MYM is part of the Red Terrana Ecuador research program backed by the Instituto Nacional de la Biodiversidad. Work by DFCH was supported by Universidad San Francisco de Quito USFQ (Taxonomía, Biogeografía y Conservación de Anfibios y Reptiles, project ID 48) and Programa “Becas de Excelencia”, Secretaría de Educación Superior, Ciencia, Tecnología e Innovación (SENESCYT), Ecuador. Work by VLU is supported by the Fulbright Foreign Student Program.