Corresponding author: Dan Cogălniceanu (
Academic editor: F. Andreone
A new species of
Describimos una nueva especie de
Székely P, Cogălniceanu D, Székely D, Páez N, Ron SR (2016) A new species of
The Neotropics have the highest amphibian species diversity in the world, housing almost half the number of known species (
Most craugastorids belong to
Field work was carried out between July and September in 2014 and March, April, and July to September in 2015 at several sites in Reserva Buenaventura. The reserve is private and belongs to the Jocotoco Conservation Foundation. The protected area has an altitudinal range between 400 and 1200 m a.s.l. and occurs in a transition zone between Deciduous Costa Forest and Western Montane Forest ( Museo de Zoología de la
For the description of qualitative and quantitative morphological characters snout-vent length head width head length interorbital distance internarial distance upper eyelid width eye diameter eye-nostril distance tympanum diameter femur length tibia length foot length hand length
DNA was extracted from muscle or liver tissue preserved in 96% ethanol or tissue storage buffer, using standard phenol–chloroform extraction protocols ( polymerase chain reaction
Voucher and GenBank accession numbers for specimens used in the phylogenetic analysis.
Voucher number | Species | 16S | RAG1 | 12S |
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MVZ203844 |
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USNM314179 |
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USNM327822 |
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USNM207945 |
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MZUSP131872 |
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- | - |
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ICNMNH23809 |
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- | - |
- |
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USNM318165 |
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USNM286919 |
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- |
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AJC0573 |
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- |
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- |
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QCAZ48309 |
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- |
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- |
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- |
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- |
NRPS0055 |
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NRPS0009 |
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- |
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- |
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MTD45080 |
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- |
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AJC1753 |
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AJC1860 |
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NRPS0048 |
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- |
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MHNSM9267 |
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- |
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AJC0594 |
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QCAZ58040 |
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QCAZ58042 |
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- |
QCAZ58043 |
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- |
QCAZ58044 |
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- |
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- |
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- |
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NRPS0085 |
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- |
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- |
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NRPS0067 |
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NRPS0001 |
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USNM336098 |
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ROM43978 |
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NRPS0060 |
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CVULA7073 |
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SIUC7062 |
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- | - |
The combined DNA matrix had up to 2914 bp. Preliminary sequence alignment was done with MAFFT 7.2 software with the L-INS-i algorithm ( Akaike information criterion
Phylogenetic trees were obtained using maximum likelihood searches with software GARLI 2.0 ( non-parametric bootstraps
The best partitioning scheme consisted of three partitions with their models of evolution in parenthesis: 12S and 16S (GTR + I + G), RAG 1st and 2nd position (HKY + G), and RAG 3rd position (TrNef + G). The phylogeny shows that the new species is most closely related to
Maximum likelihood phylogram depicting phylogenetic relationships of
(Figs
Holotype of
Holotype of
Hand and feet of the holotype of
Color variation of
Juveniles,
This species is placed in the genus
Comparisons are based on molecular evidence to compare
Among the few morphologically similar congeners from southern Ecuador,
Adult female (Fig.
Skin on dorsum shagreen with numerous small tubercles; a low W-shaped dermal ridge is present in the scapular region, with 4 larger warts defining its corners (this trait is more visible in life, Fig.
Ulnar tubercles present, coalescing into low ulnar fold; outer palmar tubercle partially divided distally; thenar tubercle ovoid; subarticular tubercles prominent, round; supernumerary palmar tubercles rounded, smaller than subarticular tubercles; fingers bearing broad lateral fringes; Finger I shorter than Finger II; discs on fingers broadly expanded, elliptical; all fingers bearing pads well defined by circumferential grooves (Fig.
Hind limbs moderately robust; tibia length 46.5% of
In life: dorsal background coloration tan (Drab–19), with dirty white spots and blotches of various sizes; flanks cream (Pale Buff–1) with darker reticulum; venter and throat cream (Pale Buff–1) with dark flecks and blotches; dorsal surface of hind limbs with faint darker transverse bars; ventral surfaces of hind limbs salmon (Light Flesh Color–250); groin, anterior and posterior surfaces of thighs with faint yellow (Light Sulphur Yellow–93) blotches; iris bronze with fine black reticulations and a median, horizontal red (Poppy Red–63) streak which is wider at the edges of the eye.
In preservative: dorsal background coloration grayish brown; the white dorsal spots and blotches become more contrasting than in life; venter and throat dirty white with brown flecks and blotches; ventral surfaces of hind limbs brown with white flecks and blotches; the yellow blotches on the groin and anterior and posterior surfaces of thighs visible in life disappear in preservative.
Measurements (in mm) and morphological proportions (in percentages) of adult males and females of , snout–vent length , head width , head length , interorbital distance , internarial distance , upper eyelid width , eye diameter , eye-nostril distance , tympanum diameter , femur length , tibia length , foot length , hand length
Character | females ( |
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29.9–37.6 (32.7 ± 2.91) | 20.4–24.9 (22.4 ± 1.86) |
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10.8–13.6 (12.1 ± 0.99) | 7.8–8.5 (8.1 ± 0.33) |
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9.2–12.1 (10.7 ± 1.07) | 5.6–7.8 (6.4 ± 0.99) |
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3.1–3.4 (3.2 ± 0.11) | 2.2–2.5 (2.3 ± 0.15) |
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2.2–2.9 (2.5 ± 0.29) | 1.3–1.4 (1.4 ± 0.05) |
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2.8–3.2 (3.0 ± 0.16) | 1.7–2.3 (2.0 ± 0.25) |
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3.9–4.4 (4.1 ± 0.19) | 2.5–3.1 (2.8 ± 0.25) |
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2.9–3.9 (3.5 ± 0.37) | 2.4–2.9 (2.6 ± 0.21) |
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1.4–1.8 (1.6 ± 0.15) | 1.0–1.1 (1.1 ± 0.05) |
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14.7–16.8 (15.7 ± 0.77) | 10.3–11.3 (10.8 ± 0.41) |
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15.3–17.5 (16.7 ± 1.01) | 10.8–12.8 (11.9 ± 0.82) |
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14.2–15.3 (14.7 ± 0.41) | 10.5–11.2 (10.8 ± 0.29) |
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8.8–10.6 (9.7 ± 0.66) | 5.4–6.9 (6.3 ± 0.65) |
36.1–38.0 | 34.1–38.2 | |
30.8–33.7 | 26.2–31.3 | |
85.2–90.9 | 71.6–91.8 | |
31.2–35.3 | 37.2–44.8 | |
36.4–42.4 | 39.7–48.3 | |
90.3–94.1 | 77.3–92.0 | |
74.4–88.6 | 92.9–96.3 | |
35.9–41.5 | 35.5–40.7 | |
44.7–49.2 | 45.4–50.5 | |
46.5–53.6 | 51.4–53.9 | |
40.7–47.4 | 44.9–51.6 |
Some females also have the W-shaped scapular dermal ridge dubbed by dark brown coloration and/or labial bars, canthal and supratympanic stripes like the males (Fig.
The degree of tuberculation and development of dermal ridges on the dorsum and flanks is usually more evident in males than females. However, the tubercles and dermal folds are difficult to observe in preservative. The low W-shaped dermal ridge in the scapular region, the thin mid dorsal fold, the incomplete longitudinal lateral fold and the thoracic fold are easily observable in life but can be very difficult to notice in the preserved specimens.
The specific name is a noun in the genitive case and refers to the Prometeo program of Secretaría de Educación Superior, Ciencia, Tecnología e Innovación, Republic of Ecuador (SENESCYT) through which Dan Cogălniceanu and Paul Székely received funding for their research in southern Ecuador.
Habitat at the type locality of
Distribution of
Our phylogenetic analysis indicates that
The Reserva Buenaventura was created in 1999 for the protection of two endemic species of birds, and despite its rather small size (about 2400 ha) is an important area for conservation in Southwestern Ecuador. Actually, the reserve is one of the most diverse sites in El Oro province hosting more than 60 species of amphibians and reptiles and 320 bird species (
Dan Cogălniceanu and Paul Székely received funding from Secretaría de Educación Superior, Ciencia, Tecnología e Innovación, Republic of Ecuador (SENESCYT) through the Prometeo Project. Laboratory work was funded by a grant from SENESCYT (Arca de Noé Initiative; Santiago R. Ron principal investigator). The authors gratefully acknowledge the support of SYNTHESYS grants: SE-TAF-4807 to Paul Székely and GB-TAF-4710 to Dan Cogălniceanu. The SYNTHESYS Project is financed by the European Community–Research Infrastructure Action under the FP7 “Capacities” Specific Programme. D. B. Provete and an anonymous reviewer provided helpful comments to the manuscript. The authors are also grateful to Jocotoco Foundation for providing access to Reserva Buenaventura and to the staff working at the Reserve, especially to Marco Gálvez, for his support during our stay. Collecting permits were granted by Ministerio de Ambiente del Ecuador (Permit no. 005-14 IC-FAU-DNB/MA).
Ecuador–Azuay: Patacocha, vía Gualaceo-Macas (
Ecuador–Cotopaxi: Pilaló (
Ecuador–Azuay: Parque Nacional El Cajas (
Ecuador–Zamora Chinchipe: Villa Nueva (
Ecuador–Loja: 5-10 km de Loja (
Ecuador–Loja: Reserva Ecológica El Madrigal, Unidad Educativa Amauta (
Ecuador–Azuay: Recinto La López (