Research Article |
Corresponding author: César Aguilar-Puntriano ( caguilarp@unmsm.edu.pe ) Academic editor: Angelica Crottini
© 2023 Valia Herrera-Alva, Alessandro Catenazzi, César Aguilar-Puntriano.
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:
Herrera-Alva V, Catenazzi A, Aguilar-Puntriano C (2023) A new cryptic species of terrestrial breeding frog of the Pristimantis danae Group (Anura, Strabomantidae) from montane forests in Ayacucho, Peru. ZooKeys 1187: 1-29. https://doi.org/10.3897/zookeys.1187.104536
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Based on morphological and molecular characters, we describe a new species of terrestrial breeding frog of the Pristimantis danae Group from montane forests of La Mar Province, Ayacucho Department in southern Peru, at elevations from 1200 to 2000 m a.s.l. The phylogenetic analysis, based on concatenated sequences of gene fragments of 16S rRNA, RAG1, COI and TYR suggests that the new species is a sister taxon of a clade that includes one undescribed species of Pristimantis from Cusco, Pristimantis pharangobates and Pristimantis rhabdolaemus. The new species is most similar to P. rhabdolaemus, which differs by lacking scapular tubercules and by its smaller size (17.0–18.6 mm in males [n = 5], 20.8–25.2 mm in females [n = 5] in the new species vs. 22.8–26.3 mm in males [n = 19], 26.0–31.9 mm in females [n = 30] of P. rhabdolaemus). Additionally, we report the prevalence of Batrachochytrium dendrobatidis (Bd) in this species.
Describimos una nueva especie de rana terrestre de desarrollo directo del grupo Pristimantis danae de bosques montanos procedentes de la provincia de La Mar, departamento de Ayacucho al sur de Perú con rango de distribución altitudinal entre los 1200–2000 msnm, en base a caracteres morfológicos y moleculares. El análisis filogenético basado en las secuencias concatenadas de los fragmentos de genes ARNr 16S, COI, RAG1 y TYR sugiere que la nueva especie es un taxón hermano del clado que incluye a una especie de Pristimantis no descrita de Cusco, Pristimantis pharangobates y Pristimantis rhabdolaemus. La nueva especie se asemeja más a P. rhabdolaemus; de la cual difiere por la ausencia de tubérculos escapulares y su menor tamaño corporal (17.0–18.6 mm en machos [n=5], 20.8–25.2 mm en hembras [n=5] en la nueva especie vs 22.8–26.3 mm en machos [n=19], 26.0–31.9 mm en hembras [n=30] de P. rhabdolaemus). Adicionalmente, reportamos la prevalencia de Batrachochytrium dendrobatidis (Bd) en esta especie de Terrarana.
Chytridiomycosis, cryptic species, montane forests, morphology, phylogeny
Bosques montanos, especies crípticas, filogenia, morfología, quitridiomicosis
Pristimantis (Terrarana, Strabomantidae) is an amphibian genus that comprises more than 600 species and occurs thoughout the Americas (
One group with cryptic species includes Pristimantis rhabdolaemus. The taxonomic history is complex because
Therefore, as part of a study of Pristimantis rhabdolaemus species boundaries, we collected direct development frogs from the montane forests of La Mar Province, Ayacucho Department. Molecular and morphological analyses revealed that the collected specimens belong to an undescribed species. Our goals are to describe the new species and infer its relationships with other species of the Pristimantis danae species Group, as well as provide information about infection by the fungus Batrachochytrium dendrobatidis (Bd).
VHA conducted field research in the montane forest of two small valleys (Fig.
We extracted liver tissues by pulling the liver out of a small abdominal incision. We fixed specimens in 10% formaldehyde and stored them in 70% ethanol in the Department of Herpetology of the Museo de Historia Natural Universidad Nacional Mayor de San Marcos (MUSM), Lima, Perú. The Ministry of Agriculture issued research, collecting and genetic resources permits (000063-2018-GRA/GG-GRDE-DRAA-DFFS-D, 029-2016-SERFOR-DGGSPFFS and D000012-2022-MIDAGRI-SERFOR-DGGSPFFS-DGSPFS).
We follow
We performed a phylogenetic analysis to infer relationships of the new species with other species of the Pristimantis danae Group (
Primers employed in this study for PCR and DNA sequencing. F = forward, R = reverse.
Locus | Primer | Sequence (5’ – 3’) | Reference | |
---|---|---|---|---|
16S | 16SAR | F | CGCCTGTTTATCAAAAACAT |
|
16SBR | R | CCGGTCTGAACTCAGATCACGT | ||
COI | dgLCO1490 | F | GGTCAACAAATCATAAAGAYATYGG |
|
dgHCO2198 | R | TAAACTTCAGGGTGACCAAARAAYCA | ||
RAG1 | R182 | F | GCCATAACTGCTGGAGCATYAT |
|
R270 | R | AGYAGATGTTGCCTGGGTCTTC | ||
TYR | Tyr1C | F | GGCAGAGGAWCRTGCCAAGATGT |
|
Tyr1G | R | TGCTGGGCRTCTCTCCARTCCCA |
We follow
We used Geneious Prime version 2023.0.1 (Biomatters, http://www.geneious.com/) to assemble pair-end reads, generate a consensus sequence for each gene and align our novel and GenBank sequences with MAFFT included in Geneious as a plug-in. Posteriorly, we concatenated the four genes (16S, COI, RAG1 and TYR) using the default settings in Geneious. We trimmed aligned sequences to a length of 591 bp for 16S, 685 bp for COI, 624 bp for RAG1 and 545 bp for TYR (Fasta file included in doi: 10.5281/zenodo.8278333). To obtain the nucleotide substitution model for each gene, we used PartitionFinder with Python v. 2.7 + Anaconda2 (
Finally, we compare uncorrected p-distances of 591 bp (including gaps) of 16S mithocondrial rRNA gene of Pristimantis included in our analysis (included as a separated file in: doi: 10.5281/zenodo.8278333). To estimate p-uncorrected genetic distances, we used MEGA 11 (
Maximum Likelihood tree of concatenated genes 16S rRNA, COI, RAG1 and TYR taken from GenBank and novel sequences. Numbers on nodes are bootstrap values (see Materials and Methods section for details). Green shadow corresponds to the ingroup. Pristimantis similaris sp. nov. in red, Pristimantis sp. 3 from Bolivia in purple and Pristimantis sp. from Cusco in blue. Maximum Likelihood optimal tree with bootstrap node values from the analysis of a concatenated dataset of 591 bp (16S), 685 bp (COI), 624 bp (RAG1) and 545 bp (TYR) of 128 species aligned by MAFFT and node support assessed using 10,000 replicates in IQTREE.
During fieldwork in 2018, 2019 and 2021, we swabbed live frogs of the new species with a synthetic dry swab (Medical Wire & Equipment #113) to quantify infection by Batrachochytrium dendrobatidis (Bd). We stroked swabs across the skin of juveniles and adults a total of 30 times per individual: five strokes on each side of the abdominal mid-line, five strokes on the inner thighs of each hind leg and five strokes on the foot webbing of each hind leg. We used a standard quantitative Polymerase Chain Reaction (qPCR) assay using DNA extracted from swabs to quantify the level of infection (
The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (ICZ) and, hence, the new name contained in the electronic version is effectively published under that Code from the electronic edition alone. This published work and its nomenclatural acts have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information is viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication is urn: urn:lsid:zoobank.org:pub:226A24AB-B4BE-4EFD-BF11-D6CA719AB601.
Our Maximum Likelihood phylogeny, based on four concatenated gene fragments (Fig.
Pristimantis scitulus is within the danae Group and sister to a clade consisting of P. aniptopalmatus, P. humboldti and P. bounides, but with low support. Both our ML and BI phylogenies recover P. danae as paraphyletic, with individuals from the type locality in Cosñipata (Cusco, Peru) forming part of a clade that includes P. danae specimens from Bolivia and P. reichlei, albeit with low support.
Genetic distances (uncorrected p-distances) for the rRNA 16S gene between P. similaris sp. nov. and species of the P. danae species Group vary from 5.6–6.9% for P. rhabdolaemus, 5.9–6.3% for P. pharangobates, 6.1–6.7% for Pristimantis sp., 6.5–7.5% for P. scitulus, 7.3–7.9% for Pristimantis sp. 3 and 7.7–9.3% for P. toftae (see Suppl. material
We found six out of 18 specimens of P. similaris (30%) infected by the fungus Batrachochytrium dendrobatidis (Bd), with levels of infection varying from 11.5 to 8889.3 zoospore genomic equivalents. Our finding confirms that species of Pristimantis can be infected with the fungus (
Our phylogenetic tree shows a candidate species from Ayacucho with high support and having high genetic distances with closely-related phylogenetic species (see Fig.
English: Similar Rubber Frog. Spanish: Rana cutín similar.
We assign this species to the genus Pristimantis, based on morphology and molecular data (Figs
Holotype. MUSM 41030, adult male (Figs
Paratypes. Nine specimens. Five adult females (MUSM 41031, 41032, 41035, 41036 and MUSM 41037). Four adult males (MUSM 41029, 41028, 41033 and MUSM 41034). All the specimens were collected at the type locality, except MUSM 41037, which was collected in Comunidad Machente (12°41'31.70"S, 73°51'0.30"W, 1640 m a.s.l.), Distrito Ayna, Provincia La Mar, Departamento Ayacucho, Peru, on 11 November 2021 by V. Herrera-Alva, E. Castillo-Urbina, V. Díaz and K. Ñaccha.
A new species of Pristimantis assigned to the P. danae species Group having the following combination of characters: (1) Skin on dorsum shagreen, skin on venter areolate; discoidal and dorsolateral folds present, weak; thoracic fold present; (2) tympanic membrane and tympanic annulus present, distinct, visible externally; (3) snout subaccuminated in dorsal view, round in lateral view; (4) upper eyelid lacking tubercles; EW smaller than IOD; cranial crest absent; two small and flat tubercles above the snout near the eyes; (5) dentigerous processes of vomers low, oblique in five of the paratypes, absent in four paratypes and the holotype; (6) males with vocal slits, subgular vocal sac large extending on to chest and without nuptial pads; (7) Finger I slightly shorter than Finger II; discs of digits expanded, flat and truncated; (8) fingers without lateral fringes; (9) ulnar tubercles present, but diffuse; (10) heel with two to three small and flat tubercles; inner tarsal fold present, small; (11) inner metatarsal tubercle ovoid, 2–3 times larger than outer one; outer metatarsal tubercle small, ovoid; numerous and flat supernumerary tubercles; (12) toes without lateral fringes; basal toe webbing absent; toe V is slightly longer than toe III; toe discs about as large as those on fingers; (13) in life, dorsum varies from blackish to dark brown with three conspicuous chevrons (not very visible in some cases) (Fig.
Pristimantis similaris is distinguished from its congeners in Peru and Bolivia by the following combination of characters: skin on dorsum areolate, tympanum and tympanic annulus distinct, weakly-defined discoidal and dorsolateral folds, two small and flat tubercles above the snout near the eyes (not conspicuous in preservative), dorsum dark brown with three darker chevrons, anterior surface of thighs usually orange-reddish and posterior surface of thighs dark brown. Pristimantis similaris can be distinguished from P. rhabdolaemus and P. pharangobates by the following characters (characters in parenthesis): smaller SVL of 20.8–25.8 mm in ten females and 15.2–18.9 mm in eight males (P. pharangobates 23.1–27.8 mm in females and 15.2–18.2 mm in males; P. rhabdolaemus 25.5–31.9 mm in females and 24.1–26.3 mm in males); absence of scapular tubercles (present in both species); presence of conspicuous longitudinal black and white or yellow marks on the throat and chest (less conspicuous in P. pharangobates and P. rhabdolaemus).
Other species in the Pristimantis danae species Group that are similar to P. similaris include P. danae, P. reichlei, P. scitulus and P. toftae. Pristimantis danae and Pristimantis reichlei also have brown chevrons in the dorsum and differ from the new species by the combination of the following characters: males nuptial pads absent (present in P. danae and P. reichlei), dorsolateral folds present (weak in P. danae and absent P. reichlei), small pale spots in the posterior surfaces of the thighs absent (present in P. danae and P. reichlei) and smaller size in P. similaris. Pristimantis scitulus is morphologically similar to P. similaris and has a parapatric distribution (Yuraccyacu, in the Piene Valley, Ayacucho Region). Pristimantis similaris can be distinguished by lacking a conical tubercle in the upper eyelid and heels (present in P. scitulus), mid-ventral line absent (present in P. scitulus) and absence of marks in the groin or thighs (conspicuous dark spots in the groin that is continuous as marks on the posterior surfaces of the thighs in P. scitulus). Pristimantis toftae is a smaller species that is superficially similar to P. similaris. The new species can be distinghished by the absence of coloured marks or spots in the groin or other parts of its body (yellow spot in the groins of P. toftae), absence of labial bar (presence of a white labial bar in P. toftae).
Pristimantis similaris is also similar to some species in the Pristimantis conspicillatus species Group, which includes P. bipunctatus, P. skydmainos and P. iiap. The parapatric Pristimantis bipunctatus (found in Calicanto at 1940 m. a.s.l. in the Piene Valley, Ayacucho Region), has dorsum and ventral skin shagreen and areolate, snout long, upper eyelids without tubercules similar to P. similaris, but the latter differs by having finger I slightly shorter than finger II (finger I and finger II about equal length in P. bipunctatus), discs on outer fingers truncated (broadly rounded in P. bipunctatus), scapular tubercules absent (present in P. bipunctatus) and by its smaller size (22.6–28.8 mm in males and 32.4–41.5 mm in females in P. bipunctatus). P. similaris can be distinguished from P. skydmainos by the absence of a mid-dorsal tubercle (present in P. skydmainos), absence of nuptial pads (present in P. skydmainos), finger I smaller than finger II (finger I longer than finger II in P. skydmainos), absence of spots or marks in the posterior surfaces of the thighs (minute cream flecks on the posterior surfaces of the thighs in P. skydmainos) and the absence of W-shaped marks (present in the scapular region in P. skydmainos). Pristimantis similaris differs from P. iiap from the lowland Amazon of the Ucayali Region by lacking large granules on flanks (present in P. iiap), lacking granules on the upper eyelids (present in P. iiap) and by having finger I shorter than finger II (finger I and II about the same length in P. iiap).
Another species with some resemblance (mainly on the throat in ventral view) to the new species is Pristimantis tanyrhynchus. Pristimantis similaris can be distinguished from P. tanyrhynchus by the absence of nuptial pads in males (present in P. tanyrhynchus) and absence of tubercles on the heel (heel with a conical and large tubercle in P. tanyrhynchus).
Head longer than wide; head length 43% of SVL; head width 35% of SVL; cranial crests absent; snout subaccuminated in dorsal view and in lateral view (Fig.
Skin on dorsum and flanks shagreen, continuous 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 present; thoracic fold present.
Outer ulnar surface without tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles well defined, most prominent on base of fingers, ovoid in ventral view, subconical in lateral view; supernumerary tubercles indistinct; fingers long and thin lacking lateral fringes, Finger I shorter than Finger II; tips of digits of fingers expanded, truncated, with circumferential grooves; nuptial pads absent (Fig.
Hind limbs long, slender, tibia length 58% of SVL; foot length 49% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs shagreen, ventral surfaces of thighs smooth; heels each with three small conical tubercles; outer surface of tarsus with one minute low tubercle; inner tarsal fold present; inner metatarsal tubercle ovoid, two times the size of round outer metatarsal tubercle; subarticular tubercles well defined, ovoid in ventral view, subconical in lateral view; few plantar supernumerary tubercles, about one quarter the size of subarticular tubercles; toes without lateral fringes; basal webbing absent; tips of digits expanded, truncated, less expanded than those on fingers, with circumferential grooves; relative length of toes: 1 < 2 < 3 < 5 < 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 17.0; tibia length 9.9; foot length 8.4; head length 7.3; head width 5.9; eye diameter 2.3; inter orbital distance 1.9; upper eyelid width 1.7; internarial distance 2.0; eye-nostril distance 2.3; tympanum length 1.0; tympanum height 1.1; forearm length 4.3.
(Fig.
The dorsal ground colouration is pale brown with three browner chevrons; narrow blackish canthal and supratympanic stripes; flanks pale brown with dark brown and cream flecks forming irregularly-shaped diagonal bars; groin and anterior surfaces of thighs brown with dark brown flecks; chest, belly and ventral surfaces of thighs pale cream, throat pale cream and grey-striped; palmar and plantar surfaces and fingers and toes dark brown; iris pale grey.
All specimens have the same general appearance, with three chevrons on the dorsum. MUSM 41029 is completely yellow and lacks marks on the chest or throat. MUSM 41032 has two brown-yellowish longitudinal bars on the dorsolateral folds. MUSM 41341 is blackish-brown and the three chevrons are not very visible (Fig.
Morphological measurements (mm) of nine paratype specimens of Pristimantis similaris sp. nov. For abbreviations, see Materials and methods.
Character | MUSM 41028 | MUSM 41029 | MUSM 41031 | MUSM 41032 | MUSM 41033 | MUSM 41034 | MUSM 41035 | MUSM 41036 | MUSM 41037 |
---|---|---|---|---|---|---|---|---|---|
Sex | Male | Male | Female | Female | Male | Male | Female | Female | Female |
SVL | 18.1 | 17.7 | 25.2 | 22.7 | 18.8 | 18.6 | 24.7 | 23.4 | 20.8 |
TL | 10.3 | 10.4 | 14.8 | 13.5 | 10.5 | 10.8 | 14.0 | 14.1 | 12.1 |
FL | 9.0 | 8.4 | 12.4 | 11.3 | 9.0 | 9.5 | 11.6 | 12.4 | 9.4 |
HL | 7.2 | 7.2 | 9.6 | 8.9 | 7.3 | 7.9 | 9.8 | 9.2 | 8.9 |
HW | 5.9 | 6.5 | 8.5 | 7.5 | 6.6 | 6.9 | 8.2 | 8.0 | 7.5 |
ED | 1.9 | 2.4 | 2.8 | 2.5 | 2.3 | 2.3 | 2.9 | 2.6 | 2.4 |
TY | 0.9 | 1.0 | 1.1 | 1.1 | 1.0 | 1.1 | 1.3 | 1.1 | 1.2 |
IOD | 2.1 | 2.0 | 2.4 | 2.2 | 2.1 | 2.0 | 2.4 | 2.3 | 2.2 |
EW | 1.8 | 1.7 | 2.3 | 1.9 | 1.8 | 1.8 | 2.3 | 2.2 | 2.1 |
IND | 2.0 | 2.3 | 2.9 | 2.6 | 2.2 | 2.2 | 2.7 | 2.5 | 2.5 |
EN | 2.3 | 2.3 | 3.1 | 2.8 | 2.5 | 2.6 | 3.0 | 2.7 | 2.8 |
F4 | 0.7 | 0.7 | 1.0 | 0.8 | 0.9 | 0.7 | 0.9 | 1.0 | 0.8 |
Measurements (in mm) and proportions of adult male and female type specimens of Pristimantis similaris sp. nov.; ranges followed by means and one standard deviation in parentheses. For abbreviations, see Materials and methods.
Character | Males (n = 5) | Females (n = 5) |
---|---|---|
SVL | 17.0–18.6 (18.1 ± 0.7) | 20.8–25.2 (23.4 ± 1.8) |
TL | 9.9–10.8 (10.4 ± 0.3) | 12.1–14.8 (13.7 ± 1.0) |
FL | 8.4–9.5 (8.9 ± 0.5) | 9.4–12.4 (11.4 ± 1.2) |
HL | 7.2–7.9 (7.4 ± 0.7) | 8.9–9.8 (9.3 ± 0.4) |
HW | 5.9–6.9 (6.5 ± 1.0) | 7.5–8.5 (7.9 ± 0.4) |
ED | 1.9–2.4 (2.2 ± 0.5) | 2.4–2.9 (2.6 ± 0.2) |
TY | 1.0–1.1 (1.0 ± 0.1) | 1.0–1.3 (1.2 ± 0.1) |
IOD | 1.9–2.1 (2.0 ± 0.1) | 2.2–2.4 (2.3 ± 0.1) |
EW | 1.7–1.8 (1.75 ± 0.05) | 1.9–2.3 (2.1 ± 0.2) |
IND | 2.0–2.3 (2.1 ± 0.1) | 2.5–2.9 (2.6 ± 0.2) |
EN | 2.3–2.6 (2.4 ± 0.3) | 2.8–3.1 (2.9 ± 0.2) |
F4 | 0.7–0.9 (0.8 ± 0.1) | 0.8–1.0 (0.9 ± 0.1) |
TL/SVL | 0.56–0.59 | 0.57–0.60 |
HL/SVL | 0.39–0.43 | 0.38–0.43 |
EN/HL | 0.32–0.34 | 0.30–0.33 |
The specific name corresponds to the Latin word “similar”. This refers to the similarity of the new species and its close phylogenetic relationship with P. rhabdolaemus and P. pharangobates.
The new species is only known from montane forests of Ayna and Anco in Departamento Ayacucho at elevations from 1200–2000 m a.s.l. in secondary forests (Figs
We describe Pristimantis similaris, a new species morphologically similar and phylogenetically related to P. rhabdolaemus and P. pharangobates. Despite their confusing taxonomic history (see Introduction), our phylogenetic analyses show that P. rhabdolaemus and P. pharangobates are distinct evolutionary lineages.
Pristimantis rhabdolaemus was described from mid-altitude montane forests restricted to Ayacucho and Cusco Departments (
We also found another candidate species from Cosñipata and Alfamayo in Cusco, morphologically similar and phylogenetically related to P. pharangobates and P. rhabdolaemus (in blue, Fig.
The taxonomy of other species of the P. danae species Group requires further work. For instance, specimens identified as P. danae or P. reichlei tend to form paraphyletic groups in phylogenies. We suggest that both species might benefit from future studies clarifying the phylogenetic relationships of their assigned populations. Such studies might include the use of genomic data for these species (including P. toftae) because the use of four genes (three nuclear) in this study was not sufficiently informative to infer with confidence phylogenetic relationships between the most inclusive clades.
Furthermore, our phylogenies include for the first time sequences of P. scitulus from Chungui, Ayacucho previously known only from two type specimens in Yuraccyacu, Ayacucho (at 2600 m a.s.l) and supports the assignment of this species in the P. danae species Group. We also included sequences for the first time of P. iiap (outgroup) and it is recovered in the P. conspicillatus species Group.
According to
We would like to highlight the areas surroundings the type locality of P. similaris and closely-related species in south-eastern Peru. The Departments of Ayacucho and Cusco have biologically “irreplaceable areas” due to the configuration of the western Andes, the eastern Cordillera de Vilcabamba and the Apurimac River (
We also provide information about infection by the fungus Batrachochytrium dendrobatidis (Bd). Chytridiomycosis, caused by the Bd fungus, has negatively affected amphibian communities in the montane forests of Central America and South America (
Fieldwork was possible by the Vicerrectorado de Investigación y Posgrado de la Universidad Nacional Mayor de San Marcos (award project code B22100451 to CAP and VHA) and Prociencia - Concytec for their financial support under contract PE501081904-2022 to VHA. Furthermore, we thank Ernesto Castillo-Urbina, Vladimir Díaz, Kimberly Ñaccha, Maura Fernandez and Juan Gamboa for their support in the field collecting the specimens; and to Sebastián Riva-Regalado for the final editions in the photos of this manuscript. We thank Anna Motta and Rafe Brown from
The authors have declared that no competing interests exist.
No ethical statement was reported.
Funding from Vicerrectorado de Investigación y Posgrado (UNMSM) B22100451 was provided to CAP and VHA, and Prociencia – Concytec PE501081904-2022 to VHA.
Writing - original draft: VHA. Writing - review and editing: VHA, AC, CAP. Investigation: VHA, AC, CAP. Methodology: VHA, AC, CAP. Funding acquisition: CAP, VHA. Data curation: VHA. Formal analysis: VHA, AC. DNA Sequencing: AC. Thesis advice: CAP, AC.
Valia Herrera-Alva https://orcid.org/0000-0001-7858-8279
Alessandro Catenazzi https://orcid.org/0000-0002-3650-4783
Cesar Aguilar-Puntriano https://orcid.org/0000-0001-6372-7926
All of the data that support the findings of this study are available in the main text.
Specimens analysed from museums. See acronyms in Materials and methods. Underlined codes correspond to type material. Coordinates and altitude when available.
Pristimantis pharangobates
: AMNH 6099, 153089: Between Abra Accanaco and Pillahuata, Paucartambo, Cusco;
Pristimantis rhabdolaemus
: CORBIDI 10813, 10815–16: Chiquintirca, La Mar, Ayacucho [-13.0350; -73.6786; altitude: 2635 m a.s.l.];
Pristimantis scitulus
:
Pristimantis sp.: AMNH 153088, 153090: Between Accanaco and Pillahuata, Paucartambo, Cusco;
Pristimantis sp. 4.:
N° | Species | Voucher | Group | Locality | Source | 16 S | COI | RAG1 | TYR |
---|---|---|---|---|---|---|---|---|---|
1 | Pristimantis albertus |
|
Ingroup | Peru: Pasco, 0.9 km N, 2.1 km E Oxapampa |
|
EU186695 | – | – | – |
2 | Pristimantis albertus | RvM 527 | Ingroup | Peru: Junin, Provincia Chanchamayo, Puyu Sacha |
|
KY594751 | – | – | – |
3 | Pristimantis albertus | MUSM 33299 | Ingroup | Peru: Junin, Provincia Chanchamayo, Cerro San Pedro |
|
KY594750 | – | – | – |
4 | Pristimantis albertus | MUSM 33298 | Ingroup | Peru: Junin, Provincia Chanchamayo, Cerro San Pedro |
|
KY594749 | – | – | – |
5 | Pristimantis aniptopalmatus |
|
Ingroup | Peru: Pasco, 2.9 km N, 5.5 km E Oxapampa |
|
EF493390 | – | – | – |
6 | Pristimantis aniptopalmatus |
|
Ingroup | Peru: Pasco, 2.9 km N, 5.9 km E Oxapampa |
|
EU186694 | – | – | – |
7 | Pristimantis aniptopalmatus | NMP6V 75053 | Ingroup | Peru: Junin, Pui Pui |
|
KY006087 | – | – | – |
8 | Pristimantis aniptopalmatus | NMP6V 75051 | Ingroup | Peru: Junin, Pui Pui |
|
KY006086 | – | – | – |
9 | Pristimantis aniptopalmatus | MUSM 31151 | Ingroup | Peru: Pasco, Yanachaga-Chemillen |
|
KY006085 | – | – | – |
10 | Pristimantis aniptopalmatus | MUSM 31130 | Ingroup | Peru: Pasco, Yanachaga-Chemillen |
|
KY006084 | – | – | – |
11 | Pristimantis aniptopalmatus | MUSM 31128 | Ingroup | Peru: Pasco, Yanachaga-Chemillen |
|
KY006083 | – | – | – |
12 | Pristimantis aniptopalmatus | MUSM 31111 | Ingroup | Peru: Pasco, Yanachaga-Chemillen |
|
KY006082 | – | – | – |
13 | Pristimantis attenboroughi | NMP6V 75529 | Ingroup | Peru: Junin, near trail from Tasta to Tarhuish, Polylepis forest patch |
|
KY594757 | KY962784 | KY962764 | – |
14 | Pristimantis attenboroughi | NMP6V 75528 | Ingroup | Peru: Junin, near trail from Tasta to Tarhuish, Polylepis forest patch |
|
KY594756 | KY962783 | KY962763 | – |
15 | Pristimantis attenboroughi | NMP6V 75524 | Ingroup | Peru: Junin, Upper part of Quebrada Tarhuish, ‘Laguna Udrecocha’ |
|
KY594754 | KY962781 | KY962761 | – |
16 | Pristimantis attenboroughi | NMP6V 75522 | Ingroup | Peru: Junin, Quebrada Tarhuish, left bank of Antuyo River |
|
KY594753 | KY962780 | KY962760 | – |
17 | Pristimantis attenboroughi | MUSM 31186 | Ingroup | Peru: Junin, Quebrada Tarhuish, left bank of Antuyo River |
|
KY594752 | KY962779 | KY962759 | – |
18 | Pristimantis attenboroughi | NMP6V 75525 | Ingroup | Peru: Junin, Upper part of Quebrada Tarhuish, ‘Laguna Udrecocha’ |
|
KY594755 | KY962782 | KY962762 | – |
19 | Pristimantis bounides | NMP6V 75097 | Ingroup | Peru: Junin, Quebrada Tasta, ‘’Runda’’ |
|
KY962797 | KY962790 | KY962774 | – |
20 | Pristimantis bounides | NMP6V 75066 | Ingroup | “Peru: Junin, Sector Carrizal, Carrtera Satipo-Toldopampa, km 134” |
|
KY962796 | – | KY962773 | – |
21 | Pristimantis bounides | NMP6V 75540 | Ingroup | “Peru: Junin, Sector Carrizal, Carrtera Satipo-Toldopampa, km 134” |
|
KY962795 | – | KY962772 | – |
22 | Pristimantis bounides | MUSM 31198 | Ingroup | Peru: Junin, Quebrada Tasta, ‘’Runda’’ |
|
KY962794 | KY962789 | KY962771 | – |
23 | Pristimantis cf aniptopalmatus | VG-2017 | Ingroup | Peru: Pasco, Yanachaga-Chemillen |
|
KY006088 | – | – | – |
24 | Pristimantis danae | MNCN 44234 | Ingroup | Peru: Cusco, Union, Valle de Kosnipata |
|
EU192270 | – | – | – |
25 | Pristimantis danae | IDLR 4001 | Ingroup | Bolivia: La Paz: Santa Cruz de Valle Ameno |
|
EU192260 | – | – | – |
26 | Pristimantis danae | MNK-A 7182 | Ingroup | Bolivia: La Paz, Huairuro, senda San Jose - Apolo |
|
EU192261 | – | – | – |
27 | Pristimantis danae | MNCN 43062 | Ingroup | Bolivia: La Paz, Huairuro, senda San Jose - Apolo |
|
EU192262 | – | – | – |
28 | Pristimantis danae | MNCN 43069 | Ingroup | Bolivia: La Paz: Arroyo Huacataya. senda San José y Apolo |
|
EU192263 | – | – | – |
29 | Pristimantis danae | MNK-A 7190 | Ingroup | Bolivia: La Paz: Arroyo Huacataya. senda San José y Apolo |
|
EU192264 | – | – | – |
30 | Pristimantis danae | MNK-A 7273 | Ingroup | Bolivia: La Paz: Serranía Bella Vista |
|
EU192265 | – | – | – |
31 | Pristimantis danae | IDLR 4815 | Ingroup | Peru: Cusco: Unión, Valle de Kosñipata |
|
EU192266 | – | – | – |
32 | Pristimantis danae | MNCN 44232 | Ingroup | Peru: Cusco: Unión, Valle de Kosñipata |
|
EU192267 | – | – | – |
33 | Pristimantis danae | MNCN 44233 | Ingroup | Peru: Cusco: Unión, Valle de Kosñipata |
|
EU192268 | – | – | – |
34 | Pristimantis danae | IDLR 4822 | Ingroup | Peru: Cusco: Unión, Valle de Kosñipata |
|
EU192269 | – | – | – |
35 | Pristimantis danae | IDLR 4824 | Ingroup | Peru: Cusco: Unión, Valle de Kosñipata |
|
EU192271 | – | – | – |
36 | Pristimantis danae | IDLR 4825 | Ingroup | Peru: Cusco: Unión, Valle de Kosñipata |
|
EU192272 | – | – | – |
37 | Pristimantis danae | MVZ 272358 | Ingroup | Peru: Cusco: Valle de Kosñipata |
|
KY652652 | KY672984 | KY672968 | KY681073 |
38 | Pristimantis danae | AC 141.09 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR469891 | – | – | OR542804 |
39 | Pristimantis danae | BL 37.13 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR469893 | OR478457 | OR542831 | OR542792 |
40 | Pristimantis danae | BL 36.13 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR469905 | OR478456 | OR542830 | OR542791 |
41 | Pristimantis humboldti | NMP6V 75538 | Ingroup | Peru: Junin, Quebrada Tarhuish, left bank of Antuyo River, Shiusha |
|
KY962799 | KY962792 | KY962776 | – |
42 | Pristimantis humboldti | MUSM 31194 | Ingroup | Peru: Junin, Quebrada Tarhuish, left bank of Antuyo River, Shiusha |
|
KY962798 | KY962791 | KY962775 | – |
43 | Pristimantis ornatus | MTD 45073 | Ingroup | Perú: Pasco: Oxapampa: Chinche: Cerca de Aquimarca |
|
EU186660 | – | – | – |
44 | Pristimantis pharangobates |
|
Ingroup | Peru: Cusco: Buenos aires |
|
EF493706 | – | – | – |
45 | Pristimantis pharangobates | MNCN 9494 | Ingroup | Peru: Cusco: Valle de Kosñipata |
|
FJ438802 | – | – | – |
46 | Pristimantis pharangobates | MHNC 11451 | Ingroup | Peru: Cusco: La Convención: Echarate: Urusayhua | This paper | OR470757 | – | – | – |
47 | Pristimantis pharangobates | MHNC 11452 | Ingroup | Peru: Cusco: La Convención: Echarate: Urusayhua | This paper | OR471343 | – | – | – |
48 | Pristimantis pharangobates | AC 96.13 | Ingroup | Peru: Cusco: Buenos Aires | This paper | OR471423 | OR478463 | – | OR542798 |
49 | Pristimantis pharangobates | AC 9.13 | Ingroup | Peru: Cusco: Buenos Aires | This paper | – | OR478451 | OR542824 | OR542787 |
50 | Pristimantis puipui | NMP6V 75542 | Ingroup | Peru: Junin, Pui Pui Protected Forest, Laguna Sinchon | von May and Lehr (2017b) | KY962800 | – | KY962777 | – |
51 | Pristimantis reichlei | MUSM 9267 | Ingroup | Perú |
|
EF493707 | – | EF493436 | EF493498 |
52 | Pristimantis reichlei | MNCN 43012 | Ingroup | Bolivia: Cochabamba: Los Guácharos |
|
EU192287 | – | – | – |
53 | Pristimantis reichlei | MNK-A 6621 | Ingroup | Bolivia: Cochabamba: Los Guácharos |
|
EU192286 | – | – | – |
54 | Pristimantis reichlei | MNCN 43249 | Ingroup | Peru: Cusco: 5 km from San Lorenzo hacia Quince Mil |
|
EU192288 | – | – | – |
55 | Pristimantis reichlei | IDLR 4779 | Ingroup | Peru: Puno: Entre Puerto Leguia y San Gabán |
|
EU192285 | – | – | – |
56 | Pristimantis reichlei | MUSM 26931 | Ingroup | Peru: Huanuco: Panguana | Pinto-Sanchez et. al (2012) | JN991461 | JN991392 | – | – |
57 | Pristimantis reichlei | CORBIDI 16219 | Ingroup | Peru: Cusco: Valle de Kosñipata |
|
KY652657 | KY672989 | KY672972 | KY681078 |
58 | Pristimantis reichlei | AC 38.15 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR471610 | OR478458 | OR542832 | – |
59 | Pristimantis reichlei | AC 113.12 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR471611 | OR478467 | OR542839 | OR542801 |
60 | Pristimantis reichlei | AC 20.15 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR471619 | OR478454 | OR542828 | – |
61 | Pristimantis reichlei | AC 138.17 | Ingroup | Peru: Quispicanchi: Soqtapata: Quincemil | This paper | OR471620 | OR478469 | OR542841 | OR542803 |
62 | Pristimantis reichlei | AC 75.17 | Ingroup | Peru: Puno: PN Bahuaja-Sonene: Punto 4 | This paper | OR471646 | OR478460 | OR542833 | – |
63 | Pristimantis reichlei | AC 94.17 | Ingroup | Peru: Puno: PN Bahuaja-Sonene: Punto 4 | This paper | OR471650 | OR478462 | OR542835 | – |
64 | Pristimantis reichlei | AC 16.17 | Ingroup | Peru: Puno, Inambari, Santo Domingo | This paper | OR471653 | – | OR542826 | – |
65 | Pristimantis reichlei | AC 33.17 | Ingroup | Peru: Puno, Inambari, Santo Domingo | This paper | OR475320 | – | OR542829 | – |
66 | Pristimantis reichlei | AC 147.17 | Ingroup | Peru: Puno: carretera San Gaban-Ollachea: Casahuiri | This paper | OR471655 | OR478472 | OR542844 | OR542807 |
67 | Pristimantis reichlei | AC 10.14 | Ingroup | Peru: Cusco: Pilcopata: Villa Carmen | This paper | OR471656 | OR478452 | OR542825 | – |
68 | Pristimantis reichlei | AC 106.17 | Ingroup | Peru: Puno: Isilluni: Valle Limbani | This paper | OR472333 | OR478465 | OR542837 | OR542799 |
69 | Pristimantis reichlei | AC 109.17 | Ingroup | Peru: Puno: Isilluni: Valle Limbani | This paper | OR472388 | – | – | OR542800 |
70 | Pristimantis reichlei | MNCN 4482 | Ingroup | Peru: Cusco, Pantiacolla |
|
EU712720 | – | – | – |
71 | Pristimantis reichlei | NMP6V 72578 | Ingroup | Bolivia: Pando, Bioceanica |
|
EU712719 | – | – | – |
72 | Pristimantis rhabdolaemus | FOCAM 34 | Ingroup | Peru: Ayacucho: La Mar: Chiquintirca | This paper | OR472495 | OR478455 | OR478455 | OR542790 |
73 | Pristimantis rhabdolaemus | FOCAM 53 | Ingroup | Peru: Ayacucho: La Mar: Toccate | This paper | OR472500 | – | – | – |
74 | Pristimantis rhabdolaemus | FOCAM 76 | Ingroup | Peru: Ayacucho: La Mar: Toccate | This paper | OR472501 | OR478461 | OR542834 | OR542797 |
75 | Pristimantis rhabdolaemus | FOCAM 145 | Ingroup | Peru: Ayacucho: La Mar: Toccate | This paper | OR472507 | – | – | – |
76 | Pristimantis rhabdolaemus | FOCAM 153 | Ingroup | Peru: Ayacucho: La Mar: Toccate | This paper | OR472508 | – | – | – |
77 | Pristimantis rhabdolaemus | FOCAM 361 | Ingroup | Peru: Ayacucho: La Mar: Toccate | This paper | OR472509 | – | OR542849 | OR542815 |
78 | Pristimantis rhabdolaemus | KNC 13 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472494 | – | – | OR542788 |
79 | Pristimantis rhabdolaemus | KNC 47 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472498 | – | – | OR542795 |
80 | Pristimantis rhabdolaemus | KNC 48 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472499 | – | – | OR542796 |
81 | Pristimantis rhabdolaemus | KNC 103 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472502 | OR478464 | OR542836 | – |
82 | Pristimantis rhabdolaemus | KNC 106 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472503 | OR478466 | OR542838 | – |
83 | Pristimantis rhabdolaemus | KNC 118 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472505 | OR478468 | OR542840 | – |
84 | Pristimantis rhabdolaemus | KNC 119 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472506 | – | – | – |
85 | Pristimantis rhabdolaemus | KNC 116 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472504 | – | – | OR542802 |
86 | Pristimantis rhabdolaemus | KNC 44 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472496 | – | – | OR542793 |
87 | Pristimantis rhabdolaemus | KNC 45 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR472497 | – | – | OR542794 |
88 | Pristimantis rhabdolaemus | MHNC 17542 | Ingroup | Peru: Cusco: La Convención: Echarate: Aendoshari | This paper | OR472510 | OR478479 | OR542852 | – |
89 | Pristimantis rhabdolaemus | MUBI 17552 | Ingroup | Peru: Cusco: La Convención: Echarate: Aendoshari | This paper | OR472511 | – | – | OR542819 |
90 | Pristimantis sagittulus |
|
Ingroup | Peru: Pasco, 0.9 km N, 2.1 km E Oxapampa |
|
EF493705 | – | EF493439 | EF493501 |
91 | Pristimantis scitulus | MUSM 41310 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR469744 | – | OR542823 | – |
92 | Pristimantis scitulus | MUSM 41309 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR469328 | – | – | – |
93 | Pristimantis scitulus | MUSM 41312 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR469801 | – | – | – |
94 | Pristimantis scitulus | MUSM 41313 | Ingroup | Peru: Ayacucho: Chungui: Chaupichaca | This paper | OR469804 | – | – | – |
95 | Pristimantis similaris | MUSM 41031 | Ingroup | Peru: Ayacucho: La Mar: Cajadela | This paper | OR478195 | OR478473 | OR542845 | OR542808 |
96 | Pristimantis similaris | MUSM 41030 | Ingroup | Peru: Ayacucho: La Mar: Cajadela | This paper | OR478198 | OR478475 | OR542847 | OR542812 |
97 | Pristimantis similaris | MUSM 41035 | Ingroup | Peru: Ayacucho: La Mar: Cajadela | This paper | OR478199 | OR478476 | OR542848 | OR542813 |
98 | Pristimantis similaris | MUSM 41036 | Ingroup | Peru: Ayacucho: La Mar: Cajadela | This paper | OR478200 | – | OR542851 | OR542817 |
99 | Pristimantis similaris | MUSM 41037 | Ingroup | Peru: Ayacucho: Ayna: Machente | This paper | OR478196 | – | – | OR542810 |
100 | Pristimantis similaris | MUSM 41323 | Ingroup | Peru: Ayacucho: Ayna: Machente | This paper | OR478197 | – | – | OR542811 |
101 | Pristimantis sp. | MVZ 272360 | Ingroup | Peru: Cusco |
|
KY652655 | KY672987 | KY681088 | KY681076 |
102 | Pristimantis sp. | MUSM 30433 | Ingroup | Peru: Cusco | This paper | OR478204 | – | OR542855 | OR542822 |
103 | Pristimantis sp. | MUSM 30418 | Ingroup | Peru: Cusco | This paper | OR478203 | – | OR542854 | OR542821 |
104 | Pristimantis sp. | AC 179.19 | Ingroup | Peru: Cusco | This paper | – | – | OR542846 | – |
105 | Pristimantis sp. | MUSM 27912 | Ingroup | Peru: Cusco | This paper | OR478202 | – | OR542853 | OR542820 |
106 | Pristimantis sp. | AC 365.07 | Ingroup | Peru: Cusco | This paper | OR478201 | OR478478 | OR542850 | OR542816 |
107 | Pristimantis sp3 | AMNH-A 165195 | Ingroup | Bolivia: Santa Cruz: Caballero: Canton San José: Parque Nacional Amboro |
|
AY843586 | – | – | – |
108 | Pristimantis sp3 | MNK-A 6628 | Ingroup | Bolivia: Santa Cruz: Serranía de la Siberia |
|
EU192258 | – | – | – |
109 | Pristimantis sp3 | MNCN 43036 | Ingroup | Bolivia: Santa Cruz: La Yunga de Mairana |
|
EU192257 | – | – | – |
110 | Pristimantis stictogaster |
|
Ingroup | Peru: Pasco, 2.9 km N, 5.5 km E Oxapampa |
|
EF493704 | – | EF493445 | EF493506 |
111 | Pristimantis toftae | KST 208 | Ingroup | Peru: Huanuco, Puerto Inca, Panguana, Rio Yuyapichis (AKA Rio Llullapiches) near Rio Pachitea | Pinto-Sanchez et. al (2012) | JN991439 | – | – | JN991566 |
112 | Pristimantis toftae | KST 318 | Ingroup | Peru: Huanuco, Puerto Inca, Panguana, Rio Yuyapichis (AKA Rio Llullapiches) near Rio Pachitea | This paper | OR538542 | – | – | – |
113 | Pristimantis toftae |
|
Ingroup | Peru: Madre de Dios: Cuzco Amazonico: 15 km E de Puerto Maldonado |
|
EF493353 | – | – | – |
114 | Pristimantis toftae | MNCN 43025 | Ingroup | Bolivia: Cochabamba: Los Guácharos |
|
EU192293 | – | – | – |
115 | Pristimantis toftae | MNCN 43246 | Ingroup | Peru: Cusco: San Pedro, Valle de Marcapata |
|
EU192294 | – | – | – |
116 | Pristimantis toftae | AC 107.07 | Ingroup | Peru: Cusco: Valle de Kosñipata |
|
KY652659 | KY672991 | KY672974 | KY681080 |
117 | Pristimantis toftae | AC 19.15 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | OR472575 | OR478453 | OR542827 | OR542789 |
118 | Pristimantis toftae | CORBIDI 11889 | Ingroup | Peru: Cusco: Valle de Kosñipata | This paper | – | – | – | OR542818 |
119 | Pristimantis toftae | AC 144.16 | Ingroup | Peru: Puno: Inambari: Santo Domingo | This paper | OR472576 | OR478470 | OR542842 | OR542805 |
120 | Pristimantis toftae | AC 147.16 | Ingroup | Peru: Puno: Inambari: Santo Domingo | This paper | OR472577 | OR478471 | OR542843 | OR542806 |
121 | Pristimantis bipunctatus | MUSM 31120 | Outgroup | Peru: Pasco, Yanachaga-Chemillen |
|
KY006089 | – | – | – |
122 | Pristimantis bipunctatus | MUSM 31179 | Outgroup | Peru: Junin, Pui Pui Protected Forest, Hito 3, Entrada del parque |
|
KY594758 | KY962785 | KY962765 | – |
123 | Pristimantis bipunctatus |
|
Outgroup | Peru: Pasco, 0.7 km S, 4.5 km E Oxapampa |
|
EF493702 | – | EF493430 | EF493492 |
124 | Pristimantis iiap | MUSM 40783 | Outgroup | Peru: Ucayali: Pucallpa: Curimana | This paper | OR470750 | OR478474 | – | OR542809 |
125 | Pristimantis iiap | MUSM 40841 | Outgroup | Peru: Ucayali: Pucallpa: Curimana | This paper | OR470749 | OR478477 | – | OR542814 |
126 | Pristimantis prolatus |
|
Outgroup | Ecuador: Napo: Rio Salado |
|
EU186701 | – | – | – |
127 | Pristimantis skydmainos | MUSM 29286 | Outgroup | Peru: Cusco: La Convención: Echarate | This paper | OR469849 | – | – | – |
128 | Pristimantis skydmainos | 448895 | Outgroup | Peru |
|
EF493393 | – | – | – |
Maximum Likelihood tree non-collapsed of concatenated genes 16S rRNA, COI, RAG1 and TYR taken from GenBank and novel sequences. Numbers on nodes are bootstrap values (see Materials and Methods section for details). Green shadow corresponds to the ingroup. Pristimantis similaris sp. nov. in red, Pristimantis sp. 3 from Bolivia in purple and Pristimantis sp. from Cusco in blue.
Bayesian Tree phylogeny collapsed of concatenated genes 16S rRNA, COI, RAG1 and TYR. Numbers on nodes are posterior probabilities (see Materials and Methods section for details). Orange shadow corresponds to the ingroup. Pristimantis similaris sp. nov. in red, Pristimantis sp. 3 from Bolivia in purple and Pristimantis sp. from Cusco in blue.
p-uncorrected distances of 591 pb including gaps of rRNA 16s gene
Data type: xls