Research Article |
Corresponding author: Juan C. Sánchez-Nivicela ( juan.sanchezn13@gmail.com ) Academic editor: Pedro Taucce
© 2024 Juan C. Sánchez-Nivicela, Diana Székely, Luis A. Salagaje M., Nicolás Astudillo-Abad, Jaime Culebras, Ernesto Arbeláez Ortiz, Paul Székely.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sánchez-Nivicela JC, Székely D, Salagaje M. LA, Astudillo-Abad N, Culebras J, Ortiz EA, Székely P (2024) One hundred years of solitude: The rediscovery of Pristimantis ruidus (Anura, Strabomantidae) in the southern Andes, Ecuador and its phylogenetic relationships. Zoosystematics and Evolution 100(3): 1107-1120. https://doi.org/10.3897/zse.100.119662
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We report the rediscovery, a century after its last record, of two specimens of Pristimantis ruidus from approximately 6 km northeast of its type locality, in a montane forest from the western Andes of southern Ecuador. This species, described by John D. Lynch in 1979, based on specimens collected in 1922 by George H. Tate, was previously known only from the type series, all specimens originating from a single locality (Molleturo, Provincia Azuay, Ecuador, 2317 m elev.). The rediscovery occurred in November 2022 in the Reserva de Conservación Quitahuaycu, Molleturo Parish, Azuay Province. The two specimens, both females, are characterised by rough skin with numerous rounded low warts and tubercles on their dorsum, a W-shaped occipital fold, tympanum and tympanic ring not externally visible and by the lack of cranial crests, characters that correspond with the description of the holotype. We evaluate the phylogenetic relationships, based on two mitochondrial genes (12S and 16S rRNA) and one nuclear gene (RAG-1). These analyses reveal Pristimantis ruidus as a member of the subgenus Huicundomantis, being most closely related to P. philipi, a species with a distribution restricted to the paramo of the same mountain region (Macizo del Cajas). Pristimantis ruidus co-exists with Pristimantis jimenezi, three additional species of Pristimantis and one Noblella with uncertain taxonomic status. This rediscovery underscores the urgency of implementing effective conservation and monitoring measures for the few remnants of the original ecosystems from western Ecuador, which face ongoing anthropogenic threats and harbour a diversity that remains insufficiently studied.
Reportamos el redescubrimiento, un siglo después de su último registro, de dos especímenes de Pristimantis ruidus en aproximadamente 6 km al noreste de su localidad tipo, en un bosque montano en los Andes occidentales del sur de Ecuador. Esta especie, descrita por John D. Lynch en 1979, con base en especímenes recolectados en 1922 por George H. Tate, se conocía solo por la serie tipo con todos los especímenes originarios de la misma localidad (Molleturo, Provincia Azuay, Ecuador, 2317 m.). El redescubrimiento ocurrió en noviembre de 2022 en la Reserva de Conservación Quitahuaycu, parroquia Molleturo, provincia de Azuay. Los dos ejemplares son hembras y se caracterizan por presentar una piel rugosa con numerosas verrugas bajas y tubérculos redondeados en su textura dérmica dorsal, un pliegue occipital en forma de W, tímpano y anillo timpánico no visibles externamente y carecen de crestas craneales, caracteres que concuerdan con el holotipo. Evaluamos relaciones filogenéticas basadas en dos genes mitocondriales (12S y 16S rRNA) y un gen nuclear (RAG-1). Estos análisis revelan a Pristimantis ruidus como miembro del subgénero Huicundomantis, estando más estrechamente relacionado con P. philipi, especie de distribución restringida que habita el páramo de la misma región montañosa (Macizo del Cajas). Pristimantis ruidus coexiste con Pristimantis jimenezi, tres especies adicionales de Pristimantis y una Noblella con estatus taxonómico incierto. Este redescubrimiento subraya la urgencia de implementar planes efectivos de conservación y monitoreo para todos los remanentes de los ecosistemas del oeste de Ecuador, que enfrentan amenazas antropogénicas continuas y albergan una diversidad que aún no está suficientemente estudiada.
Amphibia, biodiversity, conservation, Huicundomantis, molecular genetics, morphology, new record, phylogeny, systematics, taxonomy
Anfibios, biodiversidad, conservación, filogenia, genética molecular, Huicundomantis, morfología, nuevo registro, sistemática, taxonomía
The direct-developing frogs of the genus Pristimantis Jiménez de la Espada, 1870 are the most species-rich group of all amphibians, with more than 600 formally recognised species (
Pristimantis ruidus (Lynch, 1979) has eluded field researchers for over a hundred years. The 16 known specimens used in the description were collected by the naturalist George Henry Hamilton Tate (1894–1953) during his 1922 expedition for the American Museum of Natural History. The specimens were collected in south-western Ecuador, in Molleturo, Azuay Province, on the Pacific slopes of the Cordillera Occidental, at an elevation of 2317 m (
In 2022, during an expedition to the Reserva de Conservación Quitahuaycu, led by Fundación y Bioparque AMARU, members of the exploration team collected two specimens of Pristimantis, which, after comparison with the original description and holotype photographs, were identified as P. ruidus. Based on the newly-acquired specimens, we present a re-description of this species, provide for the first time images of living specimens, as well as a genetic assessment and evaluation of phylogenetic relationships of P. ruidus.
Fieldwork was carried out at Reserva de Conservación Quitahuaycu (-2.74047, -79.33532 [WGS84]), Molleturo Parish, Azuay Province, Ecuador (Fig.
We used the complete species inventory methodology (
The collected specimens were identified, based on the original description (
Morphological measurements were taken with a precision caliper and rounded to the nearest 0.1 mm. The abbreviations used follow the format of
Colour patterns in life and other important external characteristics were based on field notes and photographs taken in the field and laboratory. The colour identification in life follows
Genomic extraction, amplification and sequencing were done as described in
The sequences were edited, assembled and aligned (MAFFT algorithm with the G-INS-i iterative refinement method;
Phylogenetic relationships were inferred using both Bayesian Inference (BI) and Maximum Likelihood (ML). We used PartitionFinder v. 2.1.1 (
Using the diagnosis and description from
Additionally, the photographs of holotype (
The Bayesian Inference and the four Maximum Likelihood phylogenetic trees showed almost the same topology, with only minor differences in the position of some of the unresolved branches; the BI tree presented fewer unresolved branches and relatively higher branch support (Fig.
Bayesian phylogenetic tree of the Huicundomantis subgenus of Pristimantis. The analysis is based on 2413 base-pairs of concatenated mitochondrial DNA from 12S and 16S and nuclear DNA RAG-1 gene fragments. Outgroup is not shown. The catalogue number, species name, province and short locality name of the vouchers (in the case of Ecuadorian specimens) are shown next to each terminal (associated data are listed in Appendix
Similar to
The initial exploratory FastTree, as well as the BI and ML phylogenetic trees, recovered P. ruidus as a species in the subgenus Huicundomantis. Pristimantis ruidus is revealed as the sister species of P. philipi and both together form a basal group in Huicundomantis (Fig.
Latin, meaning rough, in reference to the skin texture (
Holotype
(
Two females: DHMECN 19106 (field series: JCS-2081) and MUTPL 1613 (field series: JCS-2084). The first individual was found amongst the remains of a felled tree (75 cm from the ground), along a stream edge (-2.73185, -79.34213, elevation 2677 m); the second individual, was on the leaf of a low shrub (40 cm from the ground), also by a stream edge (-2.73387, -79.34182, elevation 2525 m); both specimens were collected in the Reserva de Conservación Quitahuaycu, Molleturo Parish, Azuay Province, Ecuador, by Luis Salagaje, Ernesto Arbeláez, Jaime Culebras and Nicolás Astudillo, on 27 November 2022 (Fig.
A moderate-sized species of Pristimantis in the subgenus Huicundomantis as inferred from molecular phylogenetic relationships, characterised by the following combination of traits: (1) skin on head, dorsum, flanks and extremities shagreen with scattered tubercles, pustules and W-shaped scapular fold; venter coarsely areolate; discoidal fold absent; ventral texture of thighs areolate; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal and profile views; (4) upper eyelid with small rounded tubercles, 85.2% of the IOD; interocular space with rounded tubercle; cranial crests absent; (5) dentigerous processes of vomers rounded, with four and five teeth each, triangular in contour; (6) males lacking vocal sac and slits; (7) Finger I shorter than Finger II; disc expanded, elliptical; (8) fingers with narrow lateral fringes; (9) ulnar tubercles present, rounded, on the outer border of the forearm; (10) heel with several small rounded tubercles; tarsal tubercles rounded; inner tarsal fold present; (11) inner metatarsal tubercle oval, twice the size of round outer metatarsal tubercle; supernumerary plantar tubercles few in number; (12) toes with narrow lateral fringes; webbing absent; Toe V longer than Toe III; discs slightly shorter in size than those on outer fingers; (13) dorsum dark olive or brown with dark brown spots; dark W-shaped mark in scapular region, with orange spots in the higher portions; interorbital blotch, post ocular stripe and labial bars dark brown; venter dusty brown with white spots; (14) SVL females 20.1 and 30.0 mm (type series: males = 25.8–31.1 mm, females = 37.1–39.8 mm).
Pristimantis ruidus is distinguished from its congeners in the subgenus Huicundomantis by lacking a tympanic membrane and a tympanic annulus, head wider than long, short snout, cranial crest absent, thick supratympanic fold and by the presence of subacuminated postrictal tubercles. Pristimantis philipi is the only species lacking the middle ear and vocal apparatus and cranial crests (
DHMECN 19106. Head wider than long, short snout (HL = 33% of SVL), rounded in dorsal and lateral profile, nostrils orientated posterodorsally; canthus rostralis concave; cranial crest absent; one rounded interocular tubercle; upper eyelid with rounded tubercles (three and four each) and few lower; supratympanic fold thick and discontinuous; four large subacuminated postrictal tubercles; tympanic annulus and membrane absent; few small and subconical maxillary tubercles; choana barely visible in the palatal shelf, posteriorly to level of choana and broadly separated, dentigerous process of vomers conspicuous, triangular, with four and five vomerine teeth each; tongue wider than long, 30% adhering to the floor mouth; scapular fold W-shaped. Dorsum skin shagreen with numerous low warts; dorsolateral and medial fold absent; flanks shagreen with low warts and scarcely subconical and low tubercles; venter areolate with large and scattered warts; discoidal and pectoral fold absent; cloacal vent with rounded warts. Arms with low and oblique ridges and small subconical tubercles; subconical ulnar tubercles; U-shaped palmar tubercle, same length to thenar tubercle; thenar tubercle oval; low and few palmar supernumerary tubercles; subarticular tubercles rounded; fingers with lateral fringes, without interdigital membranes; Finger I shorter than Finger II; all digits with ventral pads, narrow and rounded in Finger I, barely expanded and elliptical in Finger II, expanded and elliptical in Fingers III and IV; discs in Fingers II and IV with circumferential grooves. Legs with low and oblique ridges, more noticeable in hind limbs and foot and small rounded tubercles; TL = 51% of SVL; heel and tarsus with subconical tubercles; inner tarsal fold low and extending along the tarsus; inner metatarsal tubercle oval, three times larger than external which is round; few low plantar supernumerary tubercles; subarticular tubercles rounded; toes with lateral fringes, without interdigital membranes; all digits with ventral pads and circumferential grooves, barely expanded and elliptical in Toes I–V; relative length of Toes I < II < III < V < IV (Figs
In preservative, dorsum brown with dark brown blotches, interocular bar dark brown, scapular fold dark brown with yellowish-cream blotches in the higher portions. Flanks greyish-cream with white spots in tubercles tip or warts, postocular stripe black, labial bars dark brown with thin white lines or blotches. Venter yellowish-cream with white spots in warts. Extremities yellowish-cream in concealed surfaces, dorsally brown with diagonal dark brown bars with white spots in tubercles tip, ventrally yellowish-cream with some ash shading and white spots in tip tubercles or warts (Fig.
In life, dorsally Olive Clay Colour (85), with Verona Brown (37) blotches including scapular fold, and diagonal bars in extremities; snout Olive Sulphur Yellow (90), tip of snout and upper eyelids Lime Green (105); tip of tubercles in scapular fold Brunt Orange (10). In lateral view, flanks Olive Clay Colour (85) with Russet (44) botches in the upper middle portion, change to Medium Greenish-Yellow (88) and ashen Ferruginous (35) in the lower middle portion; russet labial bars, canthal blotch and postocular stripe; Smoky White (261) blotches and lines in labials. Ventrally, ashen Cream Colour (12) with Smoky White (261) spots. Interior surfaces of digits, Spectrum Yellow (79) with Verona Brown (37) splashes. Iris Light Chrome Orange (76) with black reticulations and sclera (Figs
In the case of the MUTPL 1613 specimen, the Olive Clay Colour (85) of the dorsum and other olive or green colour tones are replaced with Salmon Colour (83) (Fig.
Morphological measurements (in mm) of the two newly-collected Pristimantis ruidus specimens and for the holotype as provided by
Museum Code | SVL | TL | FL | HL | HW | IOD | EW | IND | EN | ED | Sex | State |
---|---|---|---|---|---|---|---|---|---|---|---|---|
DHMECN 19106 | 30.0 | 15.5 | 15.3 | 10.0 | 10.9 | 3.0 | 2.6 | 2.5 | 2.6 | 3.3 | Female | adult |
MUTPL 1613 | 20.1 | 10.2 | 9.6 | 6.7 | 7.5 | 2.0 | 1.8 | 1.7 | 1.6 | 2.3 | Female | adult |
|
38.4 | 16.3 | -- | 11.4 | 13.4 | 2.9 | 3.6 | -- | 3.2 | 4.5 | Female | adult |
Pristimantis ruidus was known only from its type locality Molleturo (
Both individuals were found during the night, with full moon lighting (21:58 h–00:09 h), close to the ground (< 70 cm), in forested areas with abundant shrubby and herbaceous vegetation, soil covered by leaf litter and close to streams (Fig.
Currently, the forests on the Andean foothills in western Ecuador, which include the Molleturo Forest, retain only 30% of their original extent, this being one of the most degraded and fragmented ecosystems in Ecuador (
Various significant threats have been identified in the region, mostly linked to activities of anthropogenic origin, such as deforestation for the expansion of grazing areas and the extraction of wood for fire. The situation is aggravated by the presence of numerous mining concessions throughout the area, as well as the proliferation of illegal mining. The Molleturo Parish has one of the highest rates of poverty and inequality in the country (
The Reserva de Conservación Quitahuaycu comprises approximately 900 hectares (0.6% of the total area of Molleturo-Mollepungo AVBP), protecting ecosystems of high montane and montane evergreen forests of the western mountain of the Andean cordillera (0.14%) and the Páramo grassland (0.02%) (MAE 2012). This Reserve is the result of conservation efforts made by the community coalition of Arquillo, San José de Huigra, San Antonio and Migüir, together with the Fundación y Bioparque AMARU, aiming to preserve threatened species of amphibians which are present here, such as Atelopus nanay Coloma, 2002 (CR), Pristimantis verrucolatus Páez & Ron, 2019 (CR), Pristimantis jimenezi (CR), in addition to species with taxonomic uncertainty that are probably new species to science.
Pristimantis ruidus has a very low detectability. Despite intense search efforts carried out by different research groups in the area, its presence was missed over the last century and we were only able to encounter two individuals. We suspect that it only inhabits densely-forested areas with ravines and humid areas, so that the species is restricted to the best conserved parts of the ecosystem, Bosque siempreverde montano de Cordillera Occidental de Los Andes (MAE 2012), between elevations of 2317 and 2677 m.
Pristimantis ruidus is categorised as “Critically Endangered (Possibly Extinct) D” according to a recent assessment, because “given survey efforts and lack of records since 1922, it was inferred that it is either extinct or if there is still an extant population, that it is very small (< 50 mature individuals)” (
We identified contradictions between the Diagnosis and Description sections from
The rediscovery of P. ruidus after one hundred years shows that there still exist montane forests that preserve conditions suitable for the survival of rare and threatened species. Despite several unsuccessful efforts to record the species in the type locality, leading to an evaluation of possibly extinct (IUCN 2023), we found that a population still persists in the area, emphasising the importance of conserving this habitat, one of the last remnants of West Andean Montane Forest in southern Ecuador (
In the original description of this species,
The phylogenetic closeness between P. ruidus and P. philipi and their basal position within the entire clade of the subgenus Huicundomantis (
Our comprehensive description of the morphological characters of P. ruidus, along with the additional information regarding its phylogenetic position, distribution and natural history, enhances our understanding of a species that has eluded researchers for decades. This will enable future taxonomic and systematic research of this group of frogs and assist in conservation efforts for a threatened fauna.
We express our gratitude to the colleagues and partners of the Fundación Cóndor Andino for being a part of the field exploration team. Additionally, we extend our thanks to the friends and associates from Fundación y Bioparque AMARU for their invaluable assistance during research field trips and collaboration for the development of this research. Likewise, we are indebted to the Instituto de Biodiversidad del Ecuador (INABIO) for accepting the valuable specimen in their collections. The phylogenetic work was partially funded by Universidad Técnica Particular de Loja, through the project “Descripción de nuevas especies de anfibios y reptiles del sur de Ecuador a partir de las colecciones del Museo de Zoología, UTPL” (PROY_INV_CCBIO_2020_2794), Rainforest Trust and Naturaleza y Cultura Internacional through the “Investigación científica, para la protección y monitoreo de especies de anfibios en el Área Clave de Biodiversidad Abra de Zamora” project (PROY_INV_BA_2022_3502 PY3502). We are kindly grateful to David A. Kizirian, Lauren Vonnahme and Jesse Delia of the American Museum of Natural History for the holotype photographs. The financial support and advice of Byron Swift from ReWild and Fabián Rodas from Naturaleza y Cultura Internacional were essential to promote conservation and biodiversity research of Reserva de Conservación Quitahuaycu.
Voucher number, GenBank accession number and locality for the Pristimantis (Huicundomantis) specimens used in the phylogenetic analysis. Bold letters mark the sequences generated by the present study.
Species | Voucher number | GenBank accession no. | Locality | ||
---|---|---|---|---|---|
12S | 16S | RAG1 | |||
Pristimantis andinogigas | MUTPL359 | MT764339 | MT756022 | MT810305 | Ecuador: Loja, Parque Nacional Podocarpus, Cajanuma |
Pristimantis atillo | QCAZ42488 | - | MK881440 | MK881340 | Ecuador: Morona Santiago, Parque Nacional Sangay, Lagunas de Atillo |
Pristimantis atillo | QCAZ42498 | - | MK881444 | MK881344 | Ecuador: Morona Santiago, Parque Nacional Sangay |
Pristimantis atratus | QCAZ45580 | - | MK881471 | MK881364 | Ecuador: Loja, Cajanuma |
Pristimantis atratus | QCAZ45645 | - | MK881473 | MK881366 | Ecuador: Loja, Parque Nacional Podocarpus, Cajanuma |
Pristimantis balionotus | MUTPL180 | MT778069 | MT756023 | MT810306 | Ecuador: Loja, Reserva Madrigal del Podocarpus |
Pristimantis balionotus | MUTPL392 | MT778071 | MT756025 | MT810308 | Ecuador: Loja, Abra de Zamora |
Pristimantis ceuthospilus | KU212216 | EF493520 | EF493520 | - | Peru: Cajamarca, Chota, 12 km W Llama |
Pristimantis chomskyi | MUTPL524 | MZ678943 | MZ678934 | MZ700220 | Ecuador: Loja, Parque Nacional Podocarpus, Cerro Toledo |
Pristimantis chomskyi | QCAZ45666 | - | MK881476 | MK881369 | Ecuador: Zamora Chinchipe, Reserva Tapichalaca |
Pristimantis colodactylus | MUTPL388 | MT778072 | MT762200 | MT810310 | Loja, Abra de Zamora |
Pristimantis cryptomelas | MUTPL135 | MT778073 | MT756026 | MT810311 | Ecuador: Loja, Abra de Zamora |
Pristimantis cryptomelas | MUTPL168 | MT778075 | MT756028 | MT810313 | Ecuador: Loja, Bosque Protector Washapamba |
Pristimantis diadematus | KU221999 | EU186668 | EU186668 | - | Peru: Loreto, Teniente Lopez |
Pristimantis gagliardoi | QCAZ42575 | - | MK881456 | MK881355 | Ecuador: Morona Santiago, Parque Nacional Sangay, Ranger Station, Tinguichaca river |
Pristimantis gagliardoi | QCAZ46738 | - | MK881480 | MK881372 | Ecuador: Cañar, Reserva Mazar |
Pristimantis galdi | QCAZ32368 | EU186670 | EU186670 | EU186746 | Ecuador: Zamora Chinchipe, El Pangui |
Pristimantis gloria | KU218035 | EF493348 | EF493348 | - | Ecuador: Azuay, 8.1 km W Morona Santiago border, Gualaceo-Limón road |
Pristimantis gloria | MUTPL223 | MT778079 | MT756032 | MT810317 | Ecuador: Loja, 21 km E Urdaneta |
Pristimantis gloria | QCAZ16448 | - | MK881402 | MK881316 | Ecuador: Azuay, Gualaceo-Macas road |
Pristimantis hampatusami | QCAZ58042 | - | MK881504 | MK881387 | Ecuador: El Oro, Reserva Buenaventura |
Pristimantis hampatusami | QCAZ58044 | - | KX525478 | KX525472 | Ecuador: El Oro, Reserva Buenaventura |
Pristimantis imitatrix | KU215476 | EF493824 | EF493667 | - | Peru: Madre de Dios, Cuzco Amazonico, 15 km E Puerto Maldonado |
Pristimantis jimenezi | QCAZ45178 | - | MK881468 | MK881362 | Ecuador: Azuay, San Antonio, Parque Nacional Cajas border |
Pristimantis jimenezi | QCAZ46978 | - | MK881482 | MK881374 | Ecuador: Azuay, Molleturo, Zadracay river |
Pristimantis lojanus | MUTPL178 | MZ678945 | MZ678936 | MZ700222 | Ecuador: Loja, Loja, Quebrada San Simon |
Pristimantis lojanus | MUTPL191 | MZ678946 | MZ678937 | MZ700223 | Ecuador: Loja, Cristal |
Pristimantis lutzae | QCAZ32785 | - | MK881421 | MK881326 | Ecuador: Azuay, Bosque Protector Yanuncay-Irquis, Páramo de Quimsacocha |
Pristimantis lutzae | QCAZ53728 | - | MK881495 | - | Ecuador: Azuay, Parque Nacional Cajas, El Capo, Laguna Toreadora |
Pristimantis mallii | QCAZ45744 | MZ330729 | MZ241492 | - | Ecuador: Pastaza, Reserva Comunitaria Ankaku |
Pristimantis melanogaster | MHNSM56846 | EF493826 | EF493664 | - | Peru: Amazonas, N. Slobe Abra Barro Negro, 28 km SSW Leimebambe |
Pristimantis mallii | QCAZ45770 | MZ330730 | MZ241496 | MZ332932 | Ecuador: Pastaza, Reserva Comunitaria Ankaku |
Pristimantis miktos | GGU807 | - | KP064151 | - | Peru: Loreto, Nanay, Lote 123 |
Pristimantis miktos | QCAZ55445 | - | MZ241510 | MK391383 | Ecuador: Orellana, Parque Nacional Yasuní, Tambococha |
Pristimantis multicolor | QCAZ47213 | - | MK881488 | - | Ecuador: Loja, Parque Nacional Yacuri, Laguna Negra |
Pristimantis multicolor | QCAZ47214 | - | MK881489 | - | Ecuador: Loja, Parque Nacional Yacuri, Laguna Negra |
Pristimantis muscosus | QCAZ54857 | - | MK881501 | MK881386 | Ecuador: Zamora Chinchipe, Reserva Tapichalaca |
Pristimantis nangaritza | QCAZ41710 | - | MK881436 | MK881336 | Ecuador: Zamora Chinchipe, Alto Nangaritza PF, Las Orquídeas, Tepuy forest |
Pristimantis orcesi | KU218021 | EF493679 | EF493679 | - | Ecuador: Pichincha, Bosque de Pasochoa |
Pristimantis orestes | MUTPL242 | MT778087 | MK604538 | MK602185 | Ecuador: Loja, 11 km NE Urdaneta |
Pristimantis percultus | MUTPL810 | MT778088 | MT756034 | MT810325 | Ecuador: Loja, Parque Nacional Podocarpus, Cajanuma |
Pristimantis percultus | MUTPL812 | MT778089 | MT756035 | MT810326 | Ecuador: Loja, Parque Nacional Podocarpus, Cajanuma |
Pristimantis philipi | KU217863 | EF493672 | EF493672 | - | Ecuador: Azuay, 4 km W Laguna Toreadora, nearby Parque Nacional Cajas |
Pristimantis philipi | QCAZ37537 | - | MK881426 | MK881331 | Ecuador: Azuay, Parque Nacional Cajas |
Pristimantis phoxocephalus | QCAZ58463 | - | MK881507 | MK881390 | Ecuador: Cotopaxi, Pilaló surroundings |
Pristimantis rhodoplichus | KU219788 | EF493674 | EF493674 | - | Peru: Piura, El Tambo |
Pristimantis ruidus | DHMECN19106 | PP725379 | PP723736 | PP731015 | Ecuador: Azuay, Quitahuaycu reserve |
Pristimantis ruidus | MUTPL1613 | PP725380 | PP723737 | PP731016 | Ecuador: Azuay, Quitahuaycu reserve |
Pristimantis simonsii | KU212350 | EU186665 | EU186665 | - | Peru: Cajamarca, S slope Abra Quilsh, 28 km NNW Cajamarca |
Pristimantis spinosus | KU218052 | EF493673 | EF493673 | - | Ecuador: Morona Santiago, 10.6 km W Plan de Milagro |
Pristimantis tamia | QCAZ59630 | MZ330735 | MZ241526 | MZ332958 | Ecuador: Pastaza, Parque Nacional Llanganates, Comunidad Zarentza |
Pristimantis tamia | QCAZ59643 | MZ330736 | MZ241528 | MZ332960 | Ecuador: Pastaza, Parque Nacional Llanganates, Comunidad Zarentza |
Pristimantis teslai | QCAZ46213 | - | MK881478 | - | Ecuador: Tungurahua, Llanganatillo, Parque Nacional Llanganates border |
Pristimantis tinguichaca | QCAZ31945 | - | MK881418 | MK881323 | Ecuador: Morona Santiago, Parque Nacional Sangay, San Vicente |
Pristimantis tinguichaca | QCAZ40582 | - | MK881433 | MK881334 | Ecuador: Morona Santiago, Parque Nacional Sangay, Lagunas de Atillo |
Pristimantis torresi | MUTPL996 | MZ678947 | MZ678938 | MZ700224 | Ecuador: Loja, Guachanamá, El Apretadero |
Pristimantis torresi | QCAZ47397 | - | MK881492 | MK881380 | Ecuador: Loja, Celica-Alamor road |
Pristimantis totoroi | KU218025 | EF493349 | EF493349 | - | Ecuador: Chimborazo, 70 km W Riobamba via Pallatanga |
Pristimantis totoroi | QCAZ58425 | - | MK881505 | MK881388 | Ecuador: Cotopaxi, Pilaló surroundings |
Pristimantis unistrigatus | KU218057 | EF493387 | EF493387 | EF493444 | Ecuador: Imbabura, 35 km E Pquela |
Pristimantis verrucolatus | QCAZ46982 | - | MK881483 | MK881375 | Ecuador: Azuay, Yumate, Shoupshe |
Pristimantis verrucolatus | QCAZ46993 | - | MK881485 | MK881377 | Ecuador: Azuay, Cochapamba |
Pristimantis versicolor | KU218096 | EF493389 | EF493389 | EF493431 | Ecuador: Loja, Abra de Zamora |
Pristimantis versicolor | MUTPL494 | MT778095 | MT756038 | MT810332 | Ecuador: Loja, Reserva Madrigal del Podocarpus |
Pristimantis wiensi | KU219796 | EF493377 | EF493668 | - | Peru: Piura, 12.7 km E Canchaque |
Pristimantis sp. | QCAZ26642 | - | MK881409 | - | Ecuador: Azuay, San Antonio de Chaucha |
Pristimantis sp. | QCAZ32790 | - | MK881423 | MK881328 | Ecuador: Azuay, Bosque Protector Yanuncay-Irquis, Páramo de Quimsacocha |
Pristimantis sp. | QCAZ45029 | - | MK881461 | - | Ecuador: Morona Santiago, Parque Nacional Sangay, Etén, Rio Culebrillas |
Pristimantis sp. | QCAZ45129 | - | MK881462 | MK881358 | Ecuador: El Oro, Chillacocha |
Pristimantis sp. | QCAZ45720 | - | MZ241490 | MZ332927 | Ecuador: Pastaza, Reserva Comunitaria Ankaku |
Pristimantis sp. | QCAZ45945 | - | MZ241502 | MZ332938 | Ecuador: Pastaza, Reserva Comunitaria Ankaku |
Pristimantis sp. | QCAZ53999 | - | MK881496 | - | Ecuador: Zamora Chinchipe, Yacuambi, Romerillos |
Pristimantis sp. | QCAZ58855 | - | MZ241514 | MZ332949 | Ecuador: Morona Santiago, Parque Nacional Sangay, Sardinayacu |