Research Article |
Corresponding author: Edgar Lehr ( elehr@iwu.edu ) Academic editor: Angelica Crottini
© 2017 Edgar Lehr, Rudolf von May.
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:
Lehr E, von May R (2017) A new species of terrestrial-breeding frog (Amphibia, Craugastoridae, Pristimantis) from high elevations of the Pui Pui Protected Forest in central Peru. ZooKeys 660: 17-42. https://doi.org/10.3897/zookeys.660.11394
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We describe a new species of Pristimantis from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest and its close surroundings, Región Junín, central Peru. The description of the new species is based on 34 specimens found at elevations between 3400 and 3936 m a.s.l. Pristimantis attenboroughi sp. n. is characterized by a snout–vent length of 14.6–19.2 mm in adult males (n = 21), 19.2–23.0 mm in adult females (n = 10), and is compared morphologically and genetically with other taxonomically and biogeographically relevant species of Pristimantis. The new species is characterized by having narrow digits that lack circumferential grooves, irregularly shaped, discontinuous dorsolateral folds, and absence of both tympanic membrane and tympanic annulus. The high similarity in morphology between P. attenboroughi sp. n. and members of the Andean genera Phrynopus and Bryophryne provides an example for convergent evolution, and highlights the importance of using molecular data to justify generic assignment. Pristimantis attenboroughi sp. n. is most similar to Phrynopus chaparroi from the Región Junín, suggesting that the generic placement of this species needs to be revised. Phylogenetically the new species belongs to the Pristimantis danae species Group, a clade that includes several Pristimantis species distributed in the montane forests of central Peru, including P. albertus, P. aniptopalmatus, P. ornatus, and P. stictogaster.
Andes, DNA barcoding, frogs, molecular phylogeny, montane forest, Pristimantis attenboroughi new species, Puna
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 (Craugastoridae) as well as new species of lizards (Gymnophthalmidae). All new species were compared morphologically and genetically with other taxonomically and biogeographically relevant taxa mostly from Ecuador, Peru, and Bolivia. Herein we describe a new species of Pristimantis from upper montane and Puna habitats collected between 2012 and 2013.
Fieldwork. Because of its remote location, the PPPF is difficult to reach and is only accessible through a few entrances located ca. 1–2 days of walking distance from the nearest villages. The upper montane forests and Puna of the PPPF were reached from Toldopampa (11°30'15.4"S, 74°55'32.7"W, 3670 m a.s.l., ca. 45 km SW from Satipo) with the help of local guides by walking in 1.5 days (ca. 11 km airline). In 2012 fieldwork was conducted between May 8 and 21 by EL and RvM, and in 2013 between June 21 and July 8 by EL, J. Moravec, and J.C. Cusi. Amphibians were preserved in 96% ethanol and stored in 70% ethanol. Deposited eggs were stored in 70% ethanol.
Morphological characters. The format for the description follows
Molecular phylogenetic analysis. The phylogenetic position of the new species with respect to other morphologically similar species was determined through analysis of DNA sequence data. This analysis included two mitochondrial genes, 16S rRNA (16S) and 12S rRNA (12S). We used tissue samples from specimens collected in central Peru (Región Junín) to obtain DNA sequences for the new species and several other Pristimantis species (Table
Taxon | 16S | 12S | Voucher_Nbr | Reference |
---|---|---|---|---|
Bryophryne bakersfield | KT276289 | na | MHNC5999 |
|
Bryophryne bakersfield | KT276287 | KT276281 | MHNC6022 |
|
Bryophryne bakersfield | KT276290 | KT276282 | MHNC6023 |
|
Bryophryne bakersfield | KT276291 | KT276283 | MHNC6007 |
|
Bryophryne bakersfield | KT276288 | KT276284 | MHNC6009 |
|
Bryophryne bustamantei | KT276293 | KT276286 | MHNC6019 |
|
Bryophryne cophites | EF493537 | EF493537 | KU173497 |
|
Bufo melanostictus | FJ882791 | FJ882791 | VUB 0052 |
|
Hamptophryne boliviana | DQ283438 | DQ283438 | na |
|
Ischnocnema guentheri | EF493533 | EF493533 | na |
|
Lynchius flavomaculatus | EU186667 | EU186667 | KU218210 |
|
Lynchius nebulanastes | EU186704 | EU186704 | KU181408 |
|
Lynchius oblitus | AM039640 | AM039708 | MUSM19914 |
|
Lynchius oblitus | AM039639 | AM039707 | MTD45954 |
|
Lynchius parkeri | EU186705 | EU186705 | KU181307 |
|
Lynchius simmonsi | JF810004 | JF809940 | QZ41639 |
|
Oreobates amarakaeri | JF809996 | JF809934 | MHNC6975 |
|
Oreobates ayacucho | JF809970 | JF809933 | MNCN_IDlR5024 |
|
Oreobates cruralis | EU186666 | EU186666 | KU215462 |
|
Oreobates gemcare | JF809960 | JF809930 | MHNC6687 |
|
Oreobates granulosus | EU368897 | JF809929 | MHNC3396 |
|
Phrynopus auriculatus | EF493708 | EF493708 | KU291634 |
|
Phrynopus barthlenae | AM039653 | AM039721 | SMF81720 |
|
Phrynopus bracki | EF493709 | EF493709 | USNM286919 |
|
Phrynopus bufoides | AM039645 | AM039713 | MUSM19860 |
|
Phrynopus heimorum | AM039635 | AM039703 | MTD45621 |
|
Phrynopus heimorum | AM039636 | AM039704 | MTD45622 |
|
Phrynopus horstpauli | AM039651 | AM039719 | MTD44333 |
|
Phrynopus horstpauli | AM039647 | AM039715 | MTD44334 |
|
Phrynopus kauneorum | AM039650 | AM039718 | MTD44332 |
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Phrynopus kauneorum | AM039655 | AM039723 | MUSM20595 |
|
Phrynopus pesantesi | AM039656 | AM039724 | MTD45072 |
|
Phrynopus tautzorum | AM039652 | AM039720 | MUSM20613 |
|
Phrynopus tribulosus | EU186725 | EU186707 | KU291630 |
|
Pristimantis acuminatus | EU130579 | na | QCAZ19664 |
|
Pristimantis albertus | EU186695 | EU186695 | KU291675 |
|
Pristimantis albertus | KY594749 | na | RVM41_14 | This study |
Pristimantis albertus | KY594750 | na | RVM42_14 | This study |
Pristimantis albertus | KY594751 | na | RVM527 | This study |
Pristimantis altamazonicus | EF493670 | EF493670 | KU215460 |
|
Pristimantis altamazonicus | DQ195449 | na | MC11717 | Mahecha et al., unpublished |
Pristimantis aniptopalmatus | EF493390 | EF493390 | KU291627 |
|
Pristimantis aniptopalmatus | EU186694 | EU186694 | KU291666 |
|
Pristimantis attenboroughi sp. n. | KY594752 | na | MUSM31186 | This study |
Pristimantis attenboroughi sp. n. | KY594753 | KY594761 | NMP6V75522 | This study |
Pristimantis attenboroughi sp. n. | KY594754 | KY594762 | NMP6V75524 | This study |
Pristimantis attenboroughi sp. n. | KY594755 | KY594763 | NMP6V75525 | This study |
Pristimantis attenboroughi sp. n. | KY594756 | KY594764 | NMP6V75528 | This study |
Pristimantis attenboroughi sp. n. | KY594757 | na | NMP6V75529 | This study |
Pristimantis aureoventris | JQ742152 | na | VUB3748 |
|
Pristimantis bipunctatus | EF493702 | EF493702 | KU291638 |
|
Pristimantis bipunctatus | KY594758 | na | MUSM31179 | This study |
Pristimantis cf. mendax | KY628996 | na | MUSM31157 | This study |
Pristimantis cf. mendax | EU186659 | na | MTD45080 |
|
Pristimantis croceoinguinis | KY594759 | na | MUSM31154 | This study |
Pristimantis cruciocularis | EU186656 | EU186656 | KU291673 |
|
Pristimantis cruciocularis | KY594760 | na | NMP6V75535 | This study |
Pristimantis danae | EU192270 | na | MNCN44234 | Padial and De la Riva 2009 |
Pristimantis diadematus | EU186668 | EU186668 | KU221999 |
|
Pristimantis llojsintuta | EU712641 | na | MNCNDNA7314 |
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Pristimantis melanogaster | EF493664 | EF493826 | na |
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Pristimantis orestes | EF493388 | EF493388 | KU218257 |
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Pristimantis ornatus | EU186660 | EU186660 | MTD45073 |
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Pristimantis petrobardus | EF493367 | EF493825 | KU212293 |
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Pristimantis platydactylus | EU712653 | na | MNCNDNA3943 |
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Pristimantis platydactylus | EU712671 | na | MNCNDNA4138 |
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Pristimantis platydactylus | EU712718 | na | MNCNDNA6377 |
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Pristimantis pluvialis | KX155577 | na | CORBIDI_11862 |
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Pristimantis pluvialis | KX155578 | na | CORBIDI_16695 |
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Pristimantis reichlei | EF493707 | EF493707 | MUSM9267 |
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Pristimantis rhabdocnemus | EU186706 | EU186724 | KU291651 |
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Pristimantis rhabdolaemus | EF493706 | EF493706 | KU173492 |
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Pristimantis sagittulus | EF493705 | EF493705 | KU291635 |
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Pristimantis schultei | EF493681 | na | KU212220 |
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Pristimantis simonbolivari | EF493671 | EF493671 | KU218254 |
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Pristimantis simonsii | EU186665 | EU186665 | KU212350 |
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Pristimantis skydmainos | EF493393 | EF493393 | MUSM10071 |
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Pristimantis sp. | AM039658 | na | MTD45201 |
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Pristimantis stictogaster | EF493704 | EF493704 | KU291659 |
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Pristimantis toftae | EF493353 | EF493353 | KU215493 |
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Pristimantis toftae | EU192294 | na | MNCN43246 | Padial and De la Riva 2009 |
Pristimantis wiensi | EF493668 | EF493377 | KU219796 |
|
Extraction, amplification, and sequencing of DNA followed protocols previously used for Neotropical terrestrial breeding frogs (
Geneious R6, version 6.1.8 (Biomatters 2013; http://www.geneious.com/) was used to align the sequences. Within Geneious, we used the MAFFT, version 7.017 (
Molecular phylogenetic analysis. The Maximum Likelihood (ML) tree (Fig.
Uncorrected p-distances of the 16s mitochondrial rRNA gene for six specimens of Pristimantis attenboroughi sp. n. (in bold) and other Pristimantis species from GenBank.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | |||
---|---|---|---|---|---|---|---|---|---|---|---|
1 | Pristimantis albertus KU291675 | ||||||||||
2 | Pristimantis albertus RvM41_14 | 0.000 | |||||||||
3 | Pristimantis albertus RvM42_14 | 0.000 | 0.000 | ||||||||
4 | Pristimantis albertus RvM527 | 0.000 | 0.000 | 0.000 | |||||||
5 |
Pristimantis attenboroughi sp. n. |
0.062 | 0.065 | 0.062 | 0.066 | ||||||
6 |
Pristimantis attenboroughi sp. n. |
0.062 | 0.065 | 0.062 | 0.066 | 0.000 | |||||
7 |
Pristimantis attenboroughi sp. n. |
0.062 | 0.065 | 0.062 | 0.066 | 0.000 | 0.000 | ||||
8 |
Pristimantis attenboroughi sp. n. |
0.062 | 0.065 | 0.062 | 0.066 | 0.000 | 0.000 | 0.000 | |||
9 |
Pristimantis attenboroughi sp. n. |
0.062 | 0.065 | 0.062 | 0.066 | 0.000 | 0.000 | 0.000 | 0.000 | ||
10 |
Pristimantis attenboroughi sp. n. |
0.062 | 0.065 | 0.062 | 0.066 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | |
11 | Pristimantis ornatus MTD45073 | 0.056 | 0.059 | 0.056 | 0.059 | 0.052 | 0.052 | 0.052 | 0.052 | 0.052 | |
12 | Pristimantis stictogaster KU291659 | 0.041 | 0.043 | 0.041 | 0.043 | 0.049 | 0.049 | 0.049 | 0.049 | 0.049 | |
13 | Pristimantis aniptopalmatus KU291627 | 0.056 | 0.059 | 0.056 | 0.059 | 0.043 | 0.043 | 0.043 | 0.043 | 0.043 | |
14 | Pristimantis aniptopalmatus KU291666 | 0.056 | 0.059 | 0.056 | 0.059 | 0.043 | 0.043 | 0.043 | 0.043 | 0.043 | |
15 | Pristimantis rhabdolaemus KU173492 | 0.093 | 0.097 | 0.093 | 0.097 | 0.058 | 0.058 | 0.058 | 0.058 | 0.058 | |
16 | Pristimantis toftae KU215493 | 0.110 | 0.115 | 0.110 | 0.115 | 0.074 | 0.074 | 0.074 | 0.074 | 0.074 | |
17 | Pristimantis toftae MNCN43246 | 0.105 | 0.110 | 0.105 | 0.110 | 0.070 | 0.070 | 0.070 | 0.070 | 0.070 | |
18 | Pristimantis sagittulus KU291635 | 0.093 | 0.097 | 0.093 | 0.099 | 0.066 | 0.066 | 0.066 | 0.066 | 0.066 | |
19 | Pristimantis danae MNCN44234 | 0.116 | 0.121 | 0.116 | 0.122 | 0.094 | 0.094 | 0.094 | 0.094 | 0.094 | |
20 | Pristimantis reichlei MHNSM9267 | 0.132 | 0.135 | 0.132 | 0.136 | 0.118 | 0.118 | 0.118 | 0.118 | 0.118 | |
10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | ||
1 | Pristimantis albertus KU291675 | ||||||||||
2 | Pristimantis albertus RvM41_14 | ||||||||||
3 | Pristimantis albertus RvM42_14 | ||||||||||
4 | Pristimantis albertus RvM527 | ||||||||||
5 |
Pristimantis attenboroughi sp. n. |
||||||||||
6 |
Pristimantis attenboroughi sp. n. |
||||||||||
7 |
Pristimantis attenboroughi sp. n. |
||||||||||
8 |
Pristimantis attenboroughi sp. n. |
||||||||||
9 |
Pristimantis attenboroughi sp. n. |
||||||||||
10 |
Pristimantis attenboroughi sp. n. |
||||||||||
11 | Pristimantis ornatus MTD45073 | 0.052 | |||||||||
12 | Pristimantis stictogaster KU291659 | 0.049 | 0.037 | ||||||||
13 | Pristimantis aniptopalmatus KU291627 | 0.043 | 0.048 | 0.049 | |||||||
14 | Pristimantis aniptopalmatus KU291666 | 0.043 | 0.048 | 0.049 | 0.000 | ||||||
15 | Pristimantis rhabdolaemus KU173492 | 0.058 | 0.082 | 0.076 | 0.074 | 0.074 | |||||
16 | Pristimantis toftae KU215493 | 0.074 | 0.091 | 0.091 | 0.083 | 0.083 | 0.070 | ||||
17 | Pristimantis toftae MNCN43246 | 0.070 | 0.099 | 0.088 | 0.082 | 0.082 | 0.074 | 0.055 | |||
18 | Pristimantis sagittulus KU291635 | 0.066 | 0.084 | 0.080 | 0.068 | 0.068 | 0.066 | 0.078 | 0.095 | ||
19 | Pristimantis danae MNCN44234 | 0.094 | 0.107 | 0.107 | 0.100 | 0.100 | 0.082 | 0.101 | 0.100 | 0.083 | |
20 | Pristimantis reichlei MHNSM9267 | 0.118 | 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.
Holotype.
Paratypes. A total of 33 (Figs
We assign this species to Pristimantis based on our molecular data (Fig.
Diagnosis. A new species of Pristimantis assigned to the danae species Group having the following combination of characters: (1) Skin on dorsum shagreen with low scattered tubercles, skin on flanks tuberculate, skin on venter areolate; discoidal fold absent, thoracic fold present; irregularly shaped, discontinuous dorsolateral folds present; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal and in lateral views; (4) upper eyelid without enlarged conical tubercles; EW shorter than IOD; cranial crests absent; (5) dentigerous processes of vomers present; (6) males without vocal slits, nuptial pads absent; (7) Finger I shorter than Finger II; tips of digits narrow, rounded, lacking circumferential grooves; (8) fingers without lateral fringes; (9) small conical ulnar and tarsal tubercles present; (10) heel with a small conical tubercle; inner tarsal fold usually absent; (11) inner metatarsal tubercle ovoid, 1.5 times as large as outer; outer metatarsal tubercle small, rounded; vie low supernumerary plantar tubercles; (12) toes without lateral fringes; basal toe webbing absent; Toe V longer than Toe III; tips of digits narrow, rounded, lacking circumferential grooves, toe tips slightly smaller than those on fingers; (13) in life, dorsal ground coloration pale or dark gray, reddish brown or brownish olive with dark gray scattered flecks, some with X-shaped mark on scapular and ill-defined diagonal bars on flanks; dark grayish-brown canthal and supratympanic stripes usually present; groin dark gray or pale reddish brown with a pale red to pink tint in some; venter dark gray, pale gray, grayish brown or pale grayish green and in some dark gray mottled; iris pale grayish green with fine black vermiculation and brownish-orange horizontal streak across pupil and lower half of iris; (14) SVL in adult males 14.6–19.2 mm (n = 21), in adult females 19.2–23.0 mm (n = 10).
Comparisons. Pristimantis attenboroughi is readily distinguished from its congeners in Ecuador (176 species,
Members of the Pristimantis orestes species Group are terrestrial and inhabit high elevations in southern Ecuador and in Peru (Duellman and Lehr, 2009) and have narrow digits, and only one of the 17 species (
Among the three other new species of Pristimantis from the upper montane forests and Puna of the PPPF, only Pristimantis sp. n. E lacks circumferential grooves and a tympanum. However, P. attenboroughi and P. sp. n. E both differ regarding other morphological traits, coloration, and genetically.
Pristimantis attenboroughi shares with P. stipa Venegas and Duellman, 2012 from the Puna of northern Peru (
The new species shares narrow digits without circumferential grooves and the absence of a tympanic annulus and tympanic membrane with the Andean genera Phrynopus Peters, 1873 (except for Phrynopus auriculatus Duellman and Hedges, 2008, and P. peruanus Peters, 1873), 28 species from elevations between 2200 and 4400 m a.s.l. in central and northern Peru, Duellman and Lehr, 2009) and Bryophryne Hedges, Duellman, and Heinicke, 2008 (8 species from elevations between 2900 and 4120 m a.s.l. in southern Peru,
Head about as long as wide; head length 39.7% of SVL; head width 38.6% of SVL; cranial crests absent; snout short, rounded in dorsal view, rounded in lateral view (Fig.
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 SVL; foot length 41.3% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs tuberculate, ventral surfaces of thighs areolate; heels each with a small conical tubercle; outer surface of tarsus with few scattered minute low tubercles; inner tarsal fold absent, but small tubercle proximal to metatarsal tubercle; inner metatarsal tubercle ovoid, one and a half times the size of round outer metatarsal tubercle; subarticular tubercles well defined, round in ventral view, subconical in lateral view; few plantar supernumerary tubercles, about one third the size of subarticular tubercles; toes without lateral fringes; basal webbing absent; tips of digits narrow, round, less expanded than those on fingers, lacking circumferential grooves; relative length of toes: 1<2<5<3<4; Toe V slightly longer than Toe III (tip of digit of Toe III and Toe V not reaching distal subarticular tubercle on Toe IV; Fig.
SVL 18.9; tibia length 7.6; foot length 7.8; head length 7.5; head width 7.3; eye diameter 2.0; inter orbital distance 2.7; upper eyelid width 1.4; internarial distance 1.9; eye–nostril distance 1.4.
(Fig.
Measurements (in mm) of selected adult type specimens of Pristimantis attenboroughi sp. n. M = male, F = female. For other abbreviations see methods.
Characters |
31988 |
31992 |
31186 |
244727 |
75523 |
31980 |
31977 |
75076 |
31987 |
---|---|---|---|---|---|---|---|---|---|
sex | M | M | M | M | F | F | F | F | F |
SVL | 14.6 | 15.9 | 18.6 | 19.2 | 20.1 | 21.5 | 21.9 | 22.9 | 23.0 |
TL | 6.0 | 6.2 | 7.3 | 6.8 | 8.3 | 8.4 | 8.1 | 8.3 | 8.8 |
FL | 5.8 | 6.1 | 7.7 | 7.3 | 9.4 | 8.8 | 8.8 | 9.2 | 10.2 |
HL | 5.3 | 6.2 | 6.2 | 6.8 | 7.5 | 7.6 | 7.3 | 8.4 | 7.1 |
HW | 5.0 | 5.7 | 6.3 | 6.6 | 7.4 | 7.8 | 7.8 | 7.9 | 7.9 |
ED | 1.6 | 1.7 | 1.9 | 1.9 | 2.0 | 2.2 | 2.4 | 2.4 | 2.2 |
IOD | 1.8 | 2.1 | 2.4 | 2.1 | 2.7 | 2.5 | 2.3 | 2.6 | 2.9 |
EW | 0.9 | 1.4 | 1.2 | 1.3 | 1.6 | 1.6 | 1.6 | 1.6 | 1.3 |
IND | 1.3 | 1.5 | 1.7 | 2.0 | 2.0 | 1.9 | 2.1 | 2.3 | 2.1 |
E-N | 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 Pristimantis attenboroughi sp. n.; ranges followed by means and one standard deviation in parentheses. For abbreviations see methods.
Characters | Males (n = 21) | Females (n = 10) |
SVL | 14.6–19.2 (17.1 ± 1.2) | 19.2–23.0 (21.6 ± 1.1) |
TL | 5.8–7.6 (6.7 ± 0.5) | 8.0–8.8 (8.4 ± 0.2) |
FL | 5.8–7.8 (7.0 ± 0.5) | 8.8–10.2 (9.3 ± 0.4) |
HL | 5.3–7.3 (6.3 ± 0.5) | 7.1–8.4 (7.6 ± 0.4) |
HW | 5.0–6.9 (6.0 ± 0.5) | 7.3–8.3 (7.9 ± 0.3) |
ED | 1.6–2.1 (1.9 ± 0.2) | 1.8–2.4 (2.1 ± 0.2) |
IOD | 1.8–2.5 (2.1 ± 0.1) | 2.3–2.9 (2.7 ± 0.2) |
EW | 0.9–1.9 (1.3 ± 0.2) | 1.3–1.7 (1.5 ± 0.1) |
IND | 1.3–2.1 (1.6 ± 0.2) | 1.9–2.3 (2.1 ± 0.1) |
E-N | 0.8–1.4 (1.2 ± 0.1) | 1.3–1.8 (1.5 ± 0.2) |
TL/SVL | 0.34–0.44 | 0.36–0.42 |
FL/SVL | 0.35–0.46 | 0.40–0.47 |
HL/SVL | 0.33–0.41 | 0.31–0.39 |
HW/SVL | 0.31–0.38 | 0.34–0.39 |
HW/HL | 0.84–1.02 | 0.94–1.11 |
E-N/ED | 0.47–0.71 | 0.62–0.89 |
EW/IOD | 0.45–0.70 | 0.45–0.70 |
Variation. All paratypes (Figs
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., Life in Cold Blood, Fabulous Frogs), and for raising awareness about the importance of wildlife conservation. The specific epithet is used as noun in apposition.
Pristimantis attenboroughi is known from six localities inside the PPPF (Puna of Quebrada Tarhuish at Laguna Udrecocha, Fig.
Habitats of Pristimantis attenboroughi sp. n. in the PPPF: A type locality in the upper Tarhuish valley at Laguna Udrecocha, Puna at 3936 m a.s.l., 17 May 2012 B upper montane forest at 3550 m a.s.l. where P. attenboroughi sp. n. was found in moss pads C female P. attenboroughi sp. n. (
A female Pristimantis attenboroughi (
The IUCN Red List criteria (
Distribution of Pristimantis attenboroughi sp. n. in the PPPF and its surroundings: type locality: Laguna Udrecocha, 3936 m a.s.l.; 1 Upper part of Quebrada Tasta, “Laguna Luichococha”, 3708 m a.s.l. 2 near trail from Tasta to Tarhuish (first mountain peak), Polylepis forest patch, 3886 m a.s.l. 3 Quebrada Tarhuish, left bank of Antuyo River, “Shiusha”, 3414 m a.s.l. 4 Antuyo, 3700 m a.s.l. 5 Antuyo Bajo, 3400 m a.s.l. 6 Hatunpata, 3710 m a.s.l. 7 Trancapampa, 3550 m a.s.l.
Given that the known distribution of Pristimantis attenboroughi overlaps with the PPPF, a substantial portion of the habitat of this species is formally protected. However, other factors such as fungal infections, climate change, pollution, and man-made fires (used to expand grazing areas for livestock) continue to be threats for many Andean amphibians even inside protected areas (
When we encountered the first specimen of Pristimantis attenboroughi in the field both of us were sure that we had found a new species of Phrynopus because of its overall morphological appearance: most species in the genus Phrynopus usually lack tympanum, have narrow digits without circumferential grooves and are distributed at high elevations. However, following an integrative taxonomy approach that included molecular and morphological data, we realized that Pristimantis attenboroughi is not a Phrynopus species. Our analysis also revealed that Pristimantis attenboroughi is not closely related to other Pristimantis species that have narrow digits (e.g., members of the P. orestes species group), an assumption that could have been made if only morphological data were available. In other words, Pristimantis attenboroughi displays convergence that easily could have led to an incorrect generic assignment. Pristimantis attenboroughi is morphologically most similar to Phrynopus chaparroi (
With Pristimantis attenboroughi, seven species of Pristimantis are known from the Puna (> 3000 m a.s.l.) of Peru. Of these, six occur in northern Peru (P. atrabracus [Duellman and Pramuk, 1999], 2963–3330 m a.s.l.; P. bellator Lehr, Aguilar, Siu-Ting, Jordán, 2007, 1900–3100 m a.s.l.; P. cordovae [Lehr and Duellman, 2007b], 3400–4100 m a.s.l.; P. mariaelenae Venegas and Duellman, 2012, 3596 m a.s.l.; P. pinguis[Duellman and Pramuk, 1999], 3000–3916 m a.s.l.; P. stipa Venegas and Duellman, 2012, 3596 m a.s.l.), and only one species in central Peru (P. attenboroughi, 3400–3936 m a.s.l.),
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 (
Comparative specimens examined
Pristimantis mariaelenae: Peru: Lambayeque: Cañaris, 3406–3494 m:
Pristimantis simonsii: Peru: Cajamarca: 23.5 km NE Encanada, 3510 m:
Pristimantis stipa: Peru: Peru: Lambayeque: Cañaris, 3406–3494 m:
Phrynopus sp. n. A: Peru: Junín: Pui Pui Protected Forest: near trail from Tasta to Tarhuish (first mountain peak), Polylepis forest patch, 3886 m:
Pristimantis sp. n. C: Peru: Junín: Pui Pui Protected Forest: Quebrada Tarhuish on the left bank „Shiusha“ of Antuyo River, 3414 m:
Pristimantis sp. n. D: Peru: Junín: Pui Pui Protected Forest: Quebrada Tasta, Runda, 3463 m:
Pristimantis sp. n. E: Peru: Junín: Peru: Junín: Pui Pui Protected Forest: Laguna Sinchon, 3890 m: