Corresponding author: Edgar Lehr (
Academic editor: A. Crottini
We describe a new species of
Lehr E, von May R (2017) A new species of terrestrial-breeding frog (Amphibia, Craugastoridae,
The Pui Pui Protected Forest (Bosque de Protección Pui Pui, hereafter PPPF, Fig.
In 2012–2014, we conducted herpetological surveys in montane forests and Puna of the PPPF to catalog the amphibian and reptile species and to evaluate their conservation status. As a result, we found several new species of frogs (
Map of Peru with the Pui Pui Protected Forest indicated in red.
GenBank accession numbers for taxa and genes sampled in this study.
Taxon | 16S | 12S | Voucher_Nbr | Reference |
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na | MHNC5999 |
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MHNC6022 |
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MHNC6023 |
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MHNC6007 |
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MHNC6009 |
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MHNC6019 |
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VUB 0052 |
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na |
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na |
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MUSM19914 | |
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MHNC6975 |
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MNCN_IDlR5024 |
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MHNC6687 |
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MHNC3396 |
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USNM286919 |
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MUSM19860 |
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MUSM20595 |
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MUSM20613 |
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na | QCAZ19664 |
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na | RVM41_14 | This study |
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na | RVM42_14 | This study |
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na | RVM527 | This study |
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na |
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Mahecha et al., unpublished |
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na | MUSM31186 | This study | |
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NMP6V75522 | This study | |
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NMP6V75524 | This study | |
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NMP6V75525 | This study | |
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NMP6V75528 | This study | |
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na | NMP6V75529 | This study | |
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na | VUB3748 |
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na | MUSM31179 | This study |
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na | MUSM31157 | This study |
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na | MUSM31154 | This study |
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na | NMP6V75535 | This study |
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na | MNCN44234 | Padial and De la Riva 2009 |
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na | MNCNDNA7314 |
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na |
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na | MNCNDNA3943 |
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na | MNCNDNA4138 |
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na | MNCNDNA6377 |
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na | CORBIDI_11862 |
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na | CORBIDI_16695 |
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MUSM9267 |
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na |
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MUSM10071 |
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na |
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na | MNCN43246 | Padial and De la Riva 2009 |
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Extraction, amplification, and sequencing of DNA followed protocols previously used for Neotropical terrestrial breeding frogs (
Geneious R6, version 6.1.8 (Biomatters 2013;
Maximum Likelihood (
Uncorrected p-distances of the 16s mitochondrial rRNA gene for six specimens of
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | |||
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1 |
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2 | 0.000 | ||||||||||
3 | 0.000 | 0.000 | |||||||||
4 | 0.000 | 0.000 | 0.000 | ||||||||
5 |
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6 |
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0.000 | ||||||
7 |
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0.000 | 0.000 | |||||
8 |
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0.000 | 0.000 | 0.000 | ||||
9 |
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0.000 | 0.000 | 0.000 | 0.000 | |||
10 |
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0.000 | 0.000 | 0.000 | 0.000 | 0.000 | ||
11 |
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0.056 | 0.059 | 0.056 | 0.059 |
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12 |
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0.041 | 0.043 | 0.041 | 0.043 |
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13 |
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0.056 | 0.059 | 0.056 | 0.059 |
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14 |
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0.056 | 0.059 | 0.056 | 0.059 |
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15 |
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0.093 | 0.097 | 0.093 | 0.097 |
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16 |
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0.110 | 0.115 | 0.110 | 0.115 |
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17 | 0.105 | 0.110 | 0.105 | 0.110 |
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18 |
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0.093 | 0.097 | 0.093 | 0.099 |
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19 | 0.116 | 0.121 | 0.116 | 0.122 |
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20 | 0.132 | 0.135 | 0.132 | 0.136 |
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10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | ||
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12 |
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0.037 | ||||||||
13 |
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0.048 | 0.049 | |||||||
14 |
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0.048 | 0.049 | 0.000 | ||||||
15 |
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0.082 | 0.076 | 0.074 | 0.074 | |||||
16 |
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0.091 | 0.091 | 0.083 | 0.083 | 0.070 | ||||
17 |
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0.099 | 0.088 | 0.082 | 0.082 | 0.074 | 0.055 | ||||
18 |
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0.084 | 0.080 | 0.068 | 0.068 | 0.066 | 0.078 | 0.095 | ||
19 |
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0.107 | 0.107 | 0.100 | 0.100 | 0.082 | 0.101 | 0.100 | 0.083 | ||
20 |
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0.124 | 0.113 | 0.117 | 0.117 | 0.103 | 0.126 | 0.114 | 0.117 | 0.113 |
English: Attenborough’s Rubber Frog. Spanish: Rana cutín Attenborough.
Life male holotype (
Ventral views of right hand (
Variation of male paratypes of
Variation of female paratypes of
Variation of juvenile paratypes of
We assign this species to
Members of the
Among the three other new species of
The new species shares narrow digits without circumferential grooves and the absence of a tympanic annulus and tympanic membrane with the Andean genera
Head about as long as wide; head length 39.7% of
Skin on dorsum shagreen with low scattered tubercles, skin on flanks tuberculate, irregularly shaped, discontinuous dorsolateral folds present extending from posterior level of tympanic area to level of hind limb insertion; skin on throat, chest, and belly areolate; discoidal fold absent, thoracic fold present; cloacal sheath short.
Outer ulnar surface each with a row of four minute low tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles well defined, most prominent on base of fingers, round in ventral view, subconical in lateral view; supernumerary tubercles indistinct; fingers short and stout lacking lateral fringes, Finger I shorter than Finger II; tips of digits of fingers narrow, round, lacking circumferential grooves (Fig.
Hind limbs short, slender, tibia length 40.2% of
Measurements (in mm) of selected adult type specimens of
Characters | |||||||||
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sex | M | M | M | M | F | F | F | F | F |
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14.6 | 15.9 | 18.6 | 19.2 | 20.1 | 21.5 | 21.9 | 22.9 | 23.0 |
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6.0 | 6.2 | 7.3 | 6.8 | 8.3 | 8.4 | 8.1 | 8.3 | 8.8 |
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5.8 | 6.1 | 7.7 | 7.3 | 9.4 | 8.8 | 8.8 | 9.2 | 10.2 |
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5.3 | 6.2 | 6.2 | 6.8 | 7.5 | 7.6 | 7.3 | 8.4 | 7.1 |
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5.0 | 5.7 | 6.3 | 6.6 | 7.4 | 7.8 | 7.8 | 7.9 | 7.9 |
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1.6 | 1.7 | 1.9 | 1.9 | 2.0 | 2.2 | 2.4 | 2.4 | 2.2 |
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1.8 | 2.1 | 2.4 | 2.1 | 2.7 | 2.5 | 2.3 | 2.6 | 2.9 |
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0.9 | 1.4 | 1.2 | 1.3 | 1.6 | 1.6 | 1.6 | 1.6 | 1.3 |
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1.3 | 1.5 | 1.7 | 2.0 | 2.0 | 1.9 | 2.1 | 2.3 | 2.1 |
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1.1 | 1.0 | 1.3 | 1.3 | 1.3 | 1.7 | 1.5 | 1.8 | 1.7 |
The dorsal ground coloration is pale brown with few dark brown flecks; narrow dark brown canthal and supratympanic stripes; flanks pale brown with many dark brown flecks forming irregularly shaped diagonal bars; groin and anterior surfaces of thighs brown with dark brown flecks; chest, belly, and ventral surfaces of thighs dark brown, throat pale brown and pale gray mottled; palmar and plantar surfaces, and fingers and toes dark brown; iris pale gray.
Measurements (in mm) and proportions of adult male and adult female type specimens of
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14.6–19.2 (17.1 ± 1.2) | 19.2–23.0 (21.6 ± 1.1) |
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5.8–7.6 (6.7 ± 0.5) | 8.0–8.8 (8.4 ± 0.2) |
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5.8–7.8 (7.0 ± 0.5) | 8.8–10.2 (9.3 ± 0.4) |
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5.3–7.3 (6.3 ± 0.5) | 7.1–8.4 (7.6 ± 0.4) |
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5.0–6.9 (6.0 ± 0.5) | 7.3–8.3 (7.9 ± 0.3) |
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1.6–2.1 (1.9 ± 0.2) | 1.8–2.4 (2.1 ± 0.2) |
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1.8–2.5 (2.1 ± 0.1) | 2.3–2.9 (2.7 ± 0.2) |
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0.9–1.9 (1.3 ± 0.2) | 1.3–1.7 (1.5 ± 0.1) |
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1.3–2.1 (1.6 ± 0.2) | 1.9–2.3 (2.1 ± 0.1) |
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0.8–1.4 (1.2 ± 0.1) | 1.3–1.8 (1.5 ± 0.2) |
0.34–0.44 | 0.36–0.42 | |
0.35–0.46 | 0.40–0.47 | |
0.33–0.41 | 0.31–0.39 | |
0.31–0.38 | 0.34–0.39 | |
0.84–1.02 | 0.94–1.11 | |
0.47–0.71 | 0.62–0.89 | |
0.45–0.70 | 0.45–0.70 |
The dorsal coloration ranges from pale gray (
Juveniles (
We dedicate this species to Sir David Frederick Attenborough in honor for his educational documentaries on wildlife, especially on amphibians (e.g.,
Habitats of
A female
The
Distribution of
Given that the known distribution of
When we encountered the first specimen of
With
Additional new species of terrestrial-breeding frogs from montane forests and Puna of the PPPF will be described in the near future.
We thank the reviewers J.C. Chaparro and J.M. Guayasamin for their helpful comments that improved our manuscript. We are grateful to J.C. Cusi for designing the maps. The chief of the community Toldopampa V. Avellaneda helped us to find qualified guides, to rent horses, and allowed us to camp in the community house. We thank the director of the PPPF biologist J. Ríos, park guards H. Llantoy Cárdenas, L.F. Zevallos García, and J.M. Doñe Sánchez, and three local guides E. Bórquez Quintana, B. Porras Bórquez, and C. Avellaneda Solano. We thank J.H. Córdova (