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Four new species of terrestrial-breeding frogs of the genus Phrynopus (Anura: Terrarana: Craugastoridae) from Río Abiseo National Park, Peru

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  • Centro de Conservación, Investigación, Manejo, CIMA

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We describe four new species of terrestrial-breeding frogs belonging to the genus Phrynopus from specimens collected on the eastern slopes of the Cordillera Oriental (2800–3850 m) near and within Río Abiseo National Park, Provincia Mariscal Cáceres, Departments of San Martín and La Libertad, northeastern Peru. All four species lack a visible tympanum and inhabit the upper ridges and slopes within or adjacent to the Park. Phrynopus anancites sp. nov. and P. capitalis sp. nov. inhabit the wet montane grasslands on the upper ridges and valleys from 3600 to 3850 m. Phrynopus anancites (SVL = 25.3 mm) has coarsely aerolated skin and olive green coloration and has small vomerine teeth, while P. capitalis (female SVL = 35.6 mm) is characterized by a large head, short limbs, and distinctive dorsal pattern. Phrynopus dumicola sp. nov. (female SVL = 25.3 mm) has a short head and dark colored body with granular skin on the flanks, and is known only from forest patches along the treeline from 3225 to 3550 m, whereas P. personatus sp. nov. (female SVL = 28.2 mm) has a dark facemask and bright yellow groin spots (possibly aposematic), and inhabits a narrow band of continuous tropical montane rain forest from 2890 to 3110 m. We report infection with Batrachochytrium dendrobatidis from one specimen of P. dumicola collected in July of 1988. With the addition of these four new species, Phrynopus now includes 32 nominal species.
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Accepted by J. Padial: 10 Apr. 2017; published: 6 Jun. 2017
ZOOTAXA
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ISSN
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Copyright © 2017 Magnolia Press
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https://doi.org/10.11646/zootaxa.4273.3.4
http://zoobank.org/urn:lsid:zoobank.org:pub:ADC167F6-C3A0-43EC-8636-F65F5ABE85AF
Four new species of terrestrial-breeding frogs of the genus Phrynopus
(Anura: Terrarana: Craugastoridae) from Río Abiseo National Park, Peru
LILY O. RODRIGUEZ
1,3
& ALESSANDRO CATENAZZI
2
1
ILR, Uni-Bonn, Nussallee 21, 53115 Bonn, Germany
2
Department of Zoology, Southern Illinois University, Carbondale, IL 62901, USA
3
Corresponding author. E-mail: lily.rodriguez@ilr.uni-bonn.de
Abstract
We describe four new species of terrestrial-breeding frogs belonging to the genus Phrynopus from specimens collected on
the eastern slopes of the Cordillera Oriental (2800–3850 m) near and within Río Abiseo National Park, Provincia Mariscal
Cáceres, Departments of San Martín and La Libertad, northeastern Peru. All four species lack a visible tympanum and
inhabit the upper ridges and slopes within or adjacent to the Park.
Phrynopus anancites
sp. nov. and P. capitalis sp. nov.
inhabit the wet montane grasslands on the upper ridges and valleys from 3600 to 3850 m. Phrynopus anancites (SVL =
25.3 mm) has coarsely aerolated skin and olive green coloration and has small vomerine teeth, while P. c a pi ta li s (female
SVL = 35.6 mm) is characterized by a large head, short limbs, and distinctive dorsal pattern. Phrynopus dumicola sp. nov.
(female SVL = 25.3 mm) has a short head and dark colored body with granular skin on the flanks, and is known only from
forest patches along the treeline from 3225 to 3550 m, whereas P. personatus sp. nov. (female SVL = 28.2 mm) has a dark
facemask and bright yellow groin spots (possibly aposematic), and inhabits a narrow band of continuous tropical montane
rain forest from 2890 to 3110 m. We report infection with Batrachochytrium dendrobatidis from one specimen of P. du mi -
cola collected in July of 1988. With the addition of these four new species, Phrynopus now includes 32 nominal species.
Key words: amphibians, Andes, Batrachochytrium dendrobatidis, chytridiomycosis, direct development, montane trop-
ical forest, puna, systematics, taxonomy
Resumen
Describimos cuatro especies nuevas de anuros de reproducción terrestre pertenecientes al género Phrynopus, a partir de
especímenes colectados en el flanco Oriental de la Cordillera Oriental (2800–3850 m), Parque Nacional del Río Abiseo,
Provincia Mariscal Cáceres, Departamentos de San Martín y La Libertad, noroeste del Perú. Las cuatro especies carecen
de un tímpano visible y viven en las cimas y flancos más altos del parque. Phrynopus anancites sp. nov. y P. ca pi t al i s sp.
nov. viven en los pajonales húmedos altoandinos, en las alturas del parque entre 3550 y 3850. Phrynopus anancites (LHC=
25.3 mm) se caracteriza por una piel gruesa y granulada, coloración verde oliva, y por poseer dientes vomerinos pequeños,
mientras que P. capitalis (LHC= 35.6 mm en una hembra) se caracteriza por su cabeza grande, miembros cortos y un pa-
trón dorsal distintivo. Phrynopus dumicola sp. nov. (LHC= 25.3 mm en hembras) se caracteriza por su talla pequeña y
coloración oscura, es conocida sólo de parches de bosque a lo largo de la línea de árboles, entre 3225 y 3550 m; P. perso-
natus sp. nov. (LHC= 28.2 mm en hembras) caracterizada por una máscara rostral, y manchas brillantes posiblemente
aposemáticas en las ingles, vive en una banda angosta del bosque continuo, entre 2890 y 3110 m. Reportamos infección
por Batrachochytrium dendrobatidis en un espécimen de P. dumicola colectado en julio de 1988. Con la adición de estas
cuatro nuevas especies, Phrynopus incluye ahora 32 especies nominales.
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Introduction
The genus Phrynopus is endemic to Peru and currently contains 28 species distributed in the Cordillera Oriental
from the Huancabamba depression in northern Peru to the Mantaro-Ene drainages in central Peru (Duellman &
Lehr 2009; Frost 2016), with one species known to occur in the Cordillera Occidental (Duellman 2000). Similarly
to other genera of high-elevation terrestrial-breeding frogs such as Bryophryne and Psychrophrynella (Catenazzi &
Ttito 2016; Lehr & Catenazzi 2010), including species which had been assigned to Phrynopus prior to the splitting
of this genus (Hedges et al. 2008; Padial et al. 2014), species of Phrynopus often have very small geographic
ranges: more than half the species (16 of 28) are only known from the type locality (Duellman & Lehr 2009; Frost
2016).
Species richness is high in central Peru (Figure 1), but habitats occupied by species of Phrynopus such as cloud
forests and high elevation grasslands (puna and wet montane grasslands) are distributed continuously along the
eastern slopes of the Andes as far north as the Huancabamba depression. One could thus expect to find similar
levels of species richness of Phrynopus in northern Peru, where many regions have been poorly surveyed
(Catenazzi & von May 2014). One such region was the Río Abiseo National Park (Río Abiseo NP), on the
Amazonian slopes of the eastern slopes of the Cordillera Oriental, until teams of biologists started surveying
vertebrate communities and discovered several new species (Gardner & Romo 1993; Leo & Gardner 1993).
Río Abiseo NP is located in the Provincia Mariscal Cáceres, Department of San Martín, and protects the entire
watershed of the río Abiseo (274,520 ha) from 600 to 4200 m (Figure 2). This watershed includes the Río
Montecristo to the north, the Río Tumac in the middle, and the Río Abiseo to the south, flooding in a north-eastern
direction into the Huallaga valley. The park lays south of the xeric Huancabamba depression, the most prominent
physiographic feature of the so called cis-Andean region, often considered as a major geographical barrier for the
distribution of Andean herpetofauna (Cadle 1991; Duellman 1979; Duellman & Wild 1993; Lynch 1986). A better
knowledge of the herpetofauna of Río Abiseo NP improves our understanding of the biogeographic relevance of
the Huancabamba depression (Lehr & Catenazzi 2011, 2013).
We surveyed several localities within and adjacent to the upper part of Río Abiseo NP in July 1987, July 1988,
July–August 1989, June–July 1999 and July 2000. This remote area was accessed after 1.5 days by mule and foot
from Pataz (2600 m, Department of La Libertad), in the Marañon Valley. The survey area contains cloud and elfin
forest, wet montane grasslands (grasslands covered by plant assemblages dominated by Loricaria sp., found from
3350 to 3850 m), and puna (general term for high-elevation grasslands) interspersed with glacial lakes which are at
least 10,000 years old (Rodbell et al. 1992; Figure 3). Elfin forests occur within grasslands on slopes of small
glacial valleys at lower elevations (3200–3550 m). Runoff from these valleys feed streams that descend sharply
over steep slopes where continuous forest begins at approximately 3200 m.
Along this altitudinal gradient ranging from 2200 to 3850 m, we collected 22 species of amphibians and
reptiles. Among squamates, we collected an undescribed lizard of the genus Proctoporus sp., Sternocercus
melanopygus Boulenger, and two snakes Erythrolamprus cf. taeniurus Tschudi and Chironius monticola Roze. We
found more species of anurans: Telmatobius atahualpai Wiens, Gastrotheca phelloderma Lehr & Catenazzi,
Hyloscirtus diabolus Rivera-Correa, García-Burneo & Grant, Pristimantis corrugatus (Duellman, Lehr &
Ve ne g a s) , P. d e y i Lehr, Gregory & Catenazzi, P. wagteri (Venegas), and Phrynopus valquii, as well as undescribed
species of Rhinella, Gastrotheca, Hypodactylus, and Pristimantis.
Among the anuran specimens collected were four species of small, short-legged frogs inhabiting high-
elevation montane forest and grasslands. All of these frogs are characterized by having tips of digits narrow,
rounded or bulbous but always lacking circumferential grooves. Herein, we describe and name these four species
and tentatively assign them to the Andean genus Phrynopus sensu Hedges et al. (2008), placed in the Terrarana
taxon, and family Craugastoridae (Padial et al. 2014), based on an overall morphological similarity with other
species of this genus.
Materials and methods
Specimens were collected during day and night while conducting visual transect surveys throughout the habitats.
Mass of living frogs was measured to the nearest 0.5 g with spring scales (Pesola AG, Switzerland) prior to
preservation. Then, specimens were euthanized with benzocaine 20%, preserved in 10% formalin, and stored in
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FIGURE 1. Map of the Andes of central and northern Peru, with type localities of new species Phrynopus capitalis sp. nov., P.
dumicola sp. nov., P. personatus sp. nov. (star) and P. anancites sp. nov. (asterisk) and of congeneric forms: P. valquii
(asterisk), P. thompsoni (1), P. d a e m o n (2), P. lechriorhynchus and P. vestigiatus (3), P. horstpauli (4), P. k a u n e o r u m and P.
kotosh (5), P. dagmarae and P. interstinctus (6), P. heimorum (7), P. barthlenae and P. tautzorum (8), P. miroslawae and P.
nicoleae (9), P. badius and P. curator (10), P. auriculatus, P. bracki and P. tribulosus (11), P. pesantesi (12), P. bufoides and P.
paucari (13), P. juninensis and P. montium (14), P. oblivious and P. peruanus (15), and P. chaparroi (16).
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FIGURE 2. Study area within and around Río Abiseo National Park (bold continuous line) in the Departaments (Regiones)
San Martín and La Libertad, eastern slopes of the Cordillera Oriental of the Andes, Peru, containing type localities of (1)
Phrynopus personatus, sp. nov., El Mirador; (2) P. dumicola, sp. nov., Pampa del Cuy; (3) P. capitalis sp. nov.., Manachaqui
lake; and (4) P. anancites, sp. nov., Puesto de Vigilancia Ventanas.
70% ethanol. Preserved specimens were measured to the nearest 0.1 mm using dial calipers or a micrometer in a
stereoscope when appropriate. We follow Lynch & Duellman (1997) for the format and Duellman & Lehr (2009)
for diagnosing characters. Measurements were taken following Lynch and Duellman (1980) except that head length
was measured from the tip of the snout to the angle of the jaw. Abbreviations for measurements are as follows:
SVL = snout–vent length, HL = head length, HW = head width, TL = tibia length, THL= thigh length, FL = foot
length, IOD = interorbital distance, EW = upper eyelid width, E-N = eye-nostril distance, ED = eye diameter; IND
= internarial distance. Skeletons of Phrynopus dumicola sp. nov. (AMNH 134151 and 134167), P. valquii (MUSM
7467 and 9085), and Pristimantis simonsii (MUSM 1160) were cleared and stained following standard protocol
(Dingerkus & Uhler 1977). When type series were too small to allow clearing and staining, X-ray plates (MUSM
3820, P. d u m i c o l a sp. nov.; MUSM 8959, P. capitalis sp.nov.; MUSM 3813, P. personatus sp. nov.; MUSM 3827,
P. v a l q u i i ) were made at the National Museum of Natural History of the Smithsonian Institution and at the
American Museum of Natural History. At least one specimen of each new species (except for P. anancites sp. nov.)
was dissected and checked for the existence of a tympanic annulus. We also examined and produced X-ray plates
of P. valquii (MUSM 3824) collected near Río Abiseo NP to compare osteological characteristics with those of
three of the new species. Photographs of the preserved types taken by A. Catenazzi and M. Flecks are available at
the Calphoto online database (http://calphotos.berkeley.edu). Format of the description follows that of Duellman &
Lehr (2009).
In November 2015 we swabbed nine ethanol-preserved museum specimens of P. capitalis (n = 1), P. dumicola
(n = 4), P. montium (n = 1), P. personatus (n = 1), and P. valquii (n = 2) collected from 1964 to 1989 to detect
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infection by Batrachochytrium dendrobatidis (Bd) (Appendix 2). We swabbed specimens with a synthetic dry swab
(Medical Wire & Equipment, #113) using a standardized swabbing protocol (Catenazzi et al. 2011; Catenazzi &
Ttito 2016). Specifically, we stroked the swab across the skin a total of 30 times: 5 strokes on each side of the
abdominal midline, 5 strokes on the inner thighs of each hind leg, and 5 strokes on the foot webbing of each hind
leg (total of 30 strokes/frog). We used a real-time Polymerase Chain Reaction with a Life Technologies StepOne
Plus qPCR instrument assay on material collected on swabs to quantify the level of infection (Boyle et al. 2004;
Hyatt et al. 2007). We used PrepMan Ultra to extract DNA from swabs, and analyzed each sample once. We
calculated ZE, the genomic equivalent for Bd zoospores by comparing the qPCR results to a set of standards, and
considered any sample with ZE > 1 to be infected.
We deposited specimens in the herpetological collection of the Museo de Historia Natural, Universidad
Nacional Mayor de San Marcos (MUSM) in Lima, Peru, the American Museum of Natural History of New York
(AMNH), USA, and the Natural History Museum of The University of Kansas (KU) at Lawrence, USA. For
specimens examined, see Appendix 1.
FIGURE 3. Habitats of species of Phrynopus in the Río Abiseo NP and surrounding area. (A, B) habitat of P. capitalis sp. nov.
and P. valquii in the Manachaqui valley, 3500 m, Department of La Libertad, near the border of Río Abiseo NP. (C) Pampa del
Cuy valley, showing forest patches on the slopes; habitat of P. dumicola sp. nov.. (D) View over the Montecristo Valley, from
El Mirador, habitat of P. personatus sp. nov. Photographs by A. Catenazzi.
Generic assignment
We currently lack evidence of any synapomorphic phenotypic trait for Phrynopus that allows distinguishing
species in this genus from Pristimantis and other Terrarana with similar body shape and anatomical structures. The
overall body shape of Phrynopus and similar frogs presumably converged in response to the conditions of high-
elevation environments. In absence of molecular data, we tentatively assign the four new species to Phrynopus on
the basis of the structure of the digital discs, lack of circumferential groves, the absence of differentiated tympanic
membrane and tympanic annulus, overall body morphology with narrow head, wide body and small size, and
Finger I shorter or equal in length than Finger II (Hedges et al. 2008; Duellman & Lehr 2009). However, the
affinities of these four species should be revisited using DNA sequences.
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Results
Phrynopus anancites new species
(Figures 4, 5, 6A)
Holotype. MUSM 33168, an adult male (Figures 4, 5, 6A) collected at Ventanas (8°01´53.54”S, 77°24´28.06” W,
3820 m), Distrito Parcoy, Provincia Pataz, Departamento (Región) La Libertad, Peru on 29 June 1999 by A.
Catenazzi and L. O. Rodriguez (Figure 1).
Diagnosis. A medium size species of Phrynopus characterized by (1) a coarsely aerolate thick dorsal and
ventral skin; thoracic fold present, discoidal fold absent; (2) tympanic membrane and tympanic annulus absent,
well defined supratympanic fold; (3) snout short, rounded in dorsal view, slightly blunt in profile; (4) upper eyelid
without tubercles; width of upper eyelid narrower than IOD, cranial crests absent; (5) dentigerous processes of
vomers minute; (6) male lacking nuptial pads, vocal sac and slits; (7) Finger I as long as Finger II; thenar tubercle
ovoid, 1.5 times larger than outer palmar tubercle which is hart-shaped, tip of digits rounded, bulbous, (8) fingers
without lateral fringes, (9) ulnar tubercle absent, minute tarsal tubercle; (10) heels lacking tubercles, inner tarsal
fold absent; (11) inner metatarsal tubercle ovoid; outer metatarsal tubercle slightly smaller, round, protruding; (12)
no fringes or webbing between toes, Toe V slightly longer than III; (13) in life, dorsum bluish-olive-green, venter
gray, iris yellow with inner orange ring; in preservative, uniform dark brown, lighter brown on upper ventral
surfaces; (14) SVL 25.3 mm in single male.
No other species of Phrynopus has the combination of lack of tympanun, distinctive supratympanic fold, short
snout, small vomerine teeth, and coarsely areolated skin with uniform dorsal and ventral coloration. Of the species
lacking a visible tympanum and bearing dentigerous processes of vomers (character condition for P. anancites in
parentheses), P. bracki, P. dagmarae, P. insterstinctus, P. nicoleae, and P. vestigiatus have bright spots or coloration
in the groin (absent). Furthermore, P. dagmarae, P. insterstinctus, and P. nicoleae have either smooth or shagreened
dorsal skin (coarsely areolate); P. bracki is smaller with male SVL 13.2–16.2 mm, has a smooth venter (coarsely
areolate), ridges forming an “M” on the scapular region (no scapular ridges) and bears lateral fringes on fingers
(absent); P. dagmarae bears dorsolateral folds, lateral fringes on all digits, and the first finger is much shorter than
the second one; P. insterstinctus is a smaller frog with longer limbs than P. anancites, and a smooth venter with a
distinctive spotted pattern (uniformly gray); P. vestigiatus has lateral fringes on digits, bears ridges and dorsolateral
folds (absent). Among species of Phrynopus bearing dentigerous processes of vomers and sharing a supratympanic
fold, P. kauneorum has a shorter head (HL/SVL= 28–33 % in kauneorum, 37% in anancites), with smooth dorsal
and ventral skin (coarsely areolate) and dark blotches over a pale brown dorsum and a lighter ventral color with
some spots (uniform gray in dorsum and venter); P. k o t o s h has widely separated dentigerous processes of vomers
bearing two teeth (minute), dorsal skin shagreened with scattered tubercles, aerolate skin on venter (coarsely
aerolate dorsally and ventrally); and P. lechriorhynchus, has a distinctly spatulate snout (rounded and short).
Phrynopus capitalis has also a thick supratympanic fold, but lacks small vomerine teeth, bears fringes on digits and
basal webbing (absent), and its dorsal coloration forms a pattern (absent). Phrynopus bufoides shares most
characters with P. anancites, including the texture of the dorsal skin, but differs by bearing large elongated warts on
dorsum and flanks over a darker ground color (rounded and continuous), by having first finger longer than second
(shorter, or equal in size), and by lacking the dentigerous processes of vomers (present). Phrynopus daemon also
lacks a tympanum and has uniform dorsum coloration, but lacks vomerine teeth, bears a prominent thoracic fold,
has a shorter and less pronounced supratympanic fold, subacuminate snout in lateral view (blunt), and lateral
fringes in toes (absent). Phrynopus thompsoni lacks a tympanum and has first and second fingers equal in length,
but it bears lateral fringes on toes (absent), has pustular dorsal skin (coarsely areolate), and lacks the dentigerous
processes of vomers (present). Phrynopus chaparroi shares most characters with P. anancites, but differs by having
the skin of the tympanic area covered by elongate, round subconical tubercles instead of a supratympanic fold (fold
present), by having a rounded snout in profile (blunt), by lacking the dentigerous processes of vomers (present),
and by bearing a brownish-black dorsal coloration scattered with white to yellow spots (uniform olive), and a
brown reticulated iris (yellow ring and bronze in P. anancites). Phrynopus auriculatus and P. peruanus have a
visible tympanic annulus (absent), whereas P. montium has a dark dorsum and areolate dorsal and ventral skin, but
differs by lacking dentigerous processes of vomers, by having a visible tympanic annulus and vocal slits and
nuptial pads on swollen Finger I (absent). Most species of the Pristimantis orestes group superficially resemble
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Phrynopus anancites; those are Pristimantis atrabracus, P. chimu, P. cordovae, P mariaelenae, P. melanogaster, P.
pataikos, P. pinguis, P. seorsus, P. simonsii, P. stictoboubonus, P. stipa; all of which, except P. simonsii and P.
attenboroughi, can easily be distinguished from Phrynopus anancites by having a visible tympanum (P.
atrabracus, P. chimu, P. cordovae, P. mariaelenae, P. pinguis), or the tympanic annulus weakly defined and only
visible beneath the skin (P. melanogaster, P. pataikos, P. stictoboubonus, P. seorsus, P. stipa). Pristimantis simonsii
lacks the dentigerous processes of vomers (present), and bears lateral fringes and basal webbing on digits (absent),
while P. attenbouroughi shares with anancites the lack of tympanic membrane and annulus, and lateral fringes on
all digits, and the presence of dentigerous processes of vomers, but differs by bearing dorsal skin shagreened with
low scattered tubercles (coarsely areolate with large rounded warts), by having a dark canthal and supratympanic
stripe (well defined supratympanic fold, no color stripe), and by being smaller in size, males SVL 14.6–19.2 mm
(Lehr and von May, 2017).
FIGURE 4. Holotype of Phrynopus anancites sp. nov., male MUSM 33168 (SVL = 25.3 mm), showing (A) head in lateral
view and (B, C) body in dorsal and ventral views. Specimen collected at Ventanas, 3820 m, near the entrance of Río Abiseo
National Park, San Martín, Peru. Photographs by Morris Flecks.
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FIGURE 5. Holotype of Phrynopus anancites sp. nov., male MUSM 33168 (SVL= 25.3 mm), in life. Specimen from
Ventanas, 3820 m, near the entrance of Río Abiseo National Park, San Martín, Peru. Photograph by A. Catenazzi.
FIGURE 6. Palmar and plantar surfaces of the new species of Phrynopus: (A) P. anancites sp. nov. (MUSM 33168), (B) P.
capitalis sp. nov. (AMNH 134158), (C) P. dumicola sp. nov. (MUSM 33102), and (D) P. personatus sp. nov. (MUSM 33096).
Photographs by M. Flecks (A, C and D) and A. Catenazzi (B).
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Description of the holotype. Head narrower than body, wider than long; HW 107% of HL, HW 41% of SVL;
HL 37% of SVL; snout short, rounded in dorsal view and slightly blunt in lateral view, E-N distance around half
the size of eye; nostrils slightly protruding laterally, canthus rostralis rounded; loreal region bearing a few large
postrictal tubercles under the supratympanic fold; no tubercles on upper eyelid; choanae small, round, well
separated and anterolaterally situated; dentigerous processes of vomers minute, posterior to choanae, slightly
oblique, broadly separated; cranial crests absent; upper eyelid smaller than interorbital distance, EW 88% of IOD;
tympanic membrane and tympanic annulus absent; supratympanic fold thick, extending from edge of eye to the end
of postrictal area; tongue oval, not notched posteriorly, posterior two-third free. Vocal sac and vocal slits absent.
Skin on dorsum coarsely areolate, with large rounded warts, without lateral folds; outside of arms bearing low
rounded tubercles; discoidal fold absent, a thin thoracic fold present; skin on venter and throat coarsely areolated
bearing distinctive rounded warts, extending onto the lower parts of arms and legs, and the femoral and perianal
areas. Outer surface of arms lacking tubercles; first finger and second fingers equal in length; thenar tubercle ovoid,
1.5 times larger than palmar tubercle which is hart-shaped; fingers lacking lateral fringes, tip of digits rounded,
bulbous (Figure 6A); TL 34% of SVL; tarsus bearing a small tubercle, no tarsal fold; two metatarsal tubercles,
inner slightly elongated, outer slightly smaller, rounded, protruding; toes without lateral fringes, lacking basal
webbing; toe tips bulbous, as large as those on fingers; Toe V slightly longer than III (Figure 6A).
Measurements (in mm). SVL 25.3, HL 9.4, HW 10.1, TL 8.5, THL 9.7, FL 9.9, IOD 2.8, EW 2.5, E-N 2.7,
IND 3.1, ED 3.2.
Color in life. The dorsal and lateral surfaces are olive-green (Figure 5), whereas the ventral parts are pale gray
with darker flecks. The iris is uniformly bronze with a yellow ring.
Color in preservative. In alcohol (Figure 4), the dorsal surfaces are uniformly dark grayish-brown; chest and
throat areas are light brown, the remaining areas are as dorsal surfaces.
Etymology. The specific name anancites is a Latin noun for a type of diamond used to drive sadness away,
given in reference to the frog’s rare dorsal coloration in life.
Distribution and ecology. The species is known from a single specimen that was found near the Park rangers’
station and entrance to Rio Abiseo NP at Ventanas, in wet montane grassland covered with mosses at 3820 m of
altitude. Nothing is known about its natural history.
Phrynopus capitalis new species
(Figures 6B, 7, 8, 9A)
Holotype. AMNH 134158, an adult male (Figures 6B, 7) obtained at Lake Manachaqui (7°41’47” S, 77°30’56” W,
3600 m), ca. 14 km airline NE Pataz, Provincia Pataz, Departamento (Región) La Libertad, Peru (Figure 1),
collected on 3 July 1989 by L.O. Rodríguez.
Paratopotypes. MUSM 8959, a gravid female (Figure 8), and MUSM 8957, a juvenile, collected with the
holotype.
Diagnosis. A moderately-sized Phrynopus with the following characteristics: (1) skin on dorsum and flanks
fleshy and tuberculated, densely covered with minute round tubercles, bearing low warts especially above the
insertion of the forelimbs and dorsolaterally; skin on throat, chest and belly coarsely areolate, ventral surface of
thighs coarsely areolate; discoidal fold absent, thoracic fold evident; dorsolateral folds absent; (2) tympanic
membrane and tympanic annulus absent; supratympanic fold swollen (similar to a parotoid gland); (3) snout short,
nearly rounded in dorsal view, sloping and rounded in profile; nasals large, frontoparietals in contact; (4) upper
eyelid without tubercles; width of upper eyelid narrower than IOD; cranial crests absent; (5) dentigerous processes
of vomers absent; (6) males with gray nuptial pad on thumb, lacking vocal slits and vocal sac; (7) Finger I slightly
shorter than Finger II; tips of digits bulbous, terminal phalanges T-shaped (Figure 9A); (8) fingers bearing basal
webbing and fleshy lateral fringes; (9) ulnar and tarsal tubercles absent; (10) heels lacking tubercles; inner tarsal
fold absent; (11) inner metatarsal tubercle rounded, about 1.5 times larger than rounded outer metatarsal tubercle;
supernumerary plantar tubercles absent (12) toes bearing lateral fringes; basal webbing present; Toe V longer than
Toe III; toe tips rounded, bulbous, about as large as those on fingers; terminal phalanges with lateral processes (as
in Figure 9A) (13) in life, dorsal surfaces are brownish tan with small, dark brown reticulate spots having the
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appearance of a labyrinth; areolation on ventral surfaces are grayish cream in a darker ground color; in
preservative, flanks are blueish black or black; (14) SVL in single female 35.6 mm, in single male 23.2 mm.
The new species cannot be confused with any other described species of Phrynopus because of its very robust
and prominent head (HW/SVL = 40 %), swollen tympanic area, absence of tympanic annulus, thick lateral fringes
on all digits, and its distinctive dorsal coloration pattern with small brown reticulated spots. Phrynopus capitalis
differs from P. auriculatus, P. montium and P. peruanus by lacking a visible tympanic annulus. Phrynopus
anancites, P. auriculatus, P. bracki, P. dagmarae, P. interstinctus, P. kauneorum, P. kotosh, P. lechriorhynchus, P.
nicoleae, P. peruanus, P. vestigiatus all differ (character condition for P. capitalis in parentheses) by having the
dentigerous processes of vomers (absent). Phrynopus badius, P. curator, P. horstpauli, and P. thompsoni have
discontinuous or ridge-like dorsolateral folds (absent). Of the species lacking a tympanum, the dentigerous
processes of vomers and dorsolateral folds or ridges, P. barthlenae has similar size, coarsely tuberculate dorsal
skin, and lateral fringes on digits, but differs from P. capitalis by lacking warts (present), by bearing a dark canthal
and postorbital stripe (absent), by having small nasals (large) and widely separated frontoparietals (large nasals,
frontoparietals in contact), and distinctive black dorsal blotches (labyrinth-like short spots on dorsal surfaces).
Phrynopus bufoides is most similar to P. capitalis, but differs by having a discoidal fold (absent), a broad
supratympanic fold (swollen), by lacking lateral fringes on toes (present), and by bearing prominent dorsal isolated
and elongated warts forming discontinuous ridges (coarsely areolate dorsal skin with warts and minute tubercles),
and black mottling on venter (uniform ventral coloration). Phrynopus chaparroi lacks lateral fringes on digits
(present), has a much darker coloration without dorsal pattern (tan with brown pattern); P. d a e m o n is readily
distinguished by having subacuminate snout in lateral view (rounded), short supratympanic fold (swollen), no
dorsal pattern (short labyrinth-like lines) and males having prominent subgular vocal sac (absent), and by lacking
nuptial pads (present). Phrynopus heimorum has a slightly tuberculate dorsum (fleshy and tuberculate) and light
reticulations especially on flanks and venter (dorsum with labyrinth-like small spots, venter uniform), and lacks
lateral fringes on all digits (present); P. juninensis has smooth dorsal and ventral skin (coarsely areolate).
Phrynopus miroslawae differs by lacking lateral fringes on digits and basal webbing on toes (present), by bearing
dorsolateral, occipital and supratympanic folds (absent, supratympanic fold swollen), by having a row of
subconical tubercles on outer edge of tarsus (absent), and by being smaller with the single known female having
SVL = 29.2 mm (SVL = 35.6 mm). Phrynopus pesantesi lacks lateral fringes on digits (present), has a blunt snout
(sloping to rounded in lateral profile) and smaller head with HW/SVL = 36% (40% in P. capitalis), a tuberculate
dorsum (tuberculated bearing low warts), and olive and gray mottled and yellow blotches on groin, ventral surfaces
of throat, upper arms, forearms, thighs, shanks and venter. Phrynopus valquii differs from P. capitalis by lacking
lateral fringes or basal webbing in toes and fingers, by bearing canthal and supratympanic stripes (absent), and by
having dorsal skin shagreen with scattered tubercles (coarsely areolate). Phrynopus tautzorum has a smooth
dorsum with scattered tubercles (fleshy and tuberculate), lacks lateral fringes on digits (present), and has Toe V
slightly shorter than Toe III (Toe V longer than Toe III). The following species of the Pristimantis orestes group
with reduced finger tips may externally resemble Phrynopus capitalis: Pristimantis atrabracus, P. chimu, P.
cordovae, P mariaelenae, P. melanogaster, P. pataikos, P. pinguis, P. seorsus, P. simonsii, P. stictoboubonus, P.
stipa. All of these except P. simonsii and P. attenbouroughi, have a visible tympanum (P. atrabracus, P. chimu, P.
cordovae, P. mariaelenae, P. pinguis), or the tympanic annulus is weakly defined or concealed beneath the skin (P.
melanogaster, P. pataikos, P. stictoboubonus, P. seorsus, P. stipa). The other two members of the group occurring in
Peru, P. corrugatus and P. ventriguttatus have well developed pads on all digits and cannot be confused with
Phrynopus capitalis. Pristimantis simonsii lacks a tympanum, dentigerous processes of vomers, and bears lateral
fringes on digits, as does Phrynopus capitalis, but differs by having relatively larger nasal bones, by lacking the
swollen supratympanic fold, by having smoother dorsal skin (coarsely tuberculate with warts) and by bearing some
dorsal dark blotches (absent). Pristimantis attenbouroughi differs by bearing the dentigerous processes of vomers
(absent), by having dorsal skin shagreened with low scattered tubercles (coarsely areolate with large rounded
warts), a dark canthal and supratympanic stripe (absent, supratympanic fold swollen), and by being smaller with
males SVL = 14.6–19.2 mm (single male SVL = 23.2 mm; Lehr and von May 2017).
Description of holotype. Head distinct, narrower than body, wider than long, (HW 133% of HL; HW 40% of
SVL; snout nearly rounded in dorsal view, sloping to rounded tip in lateral profile; snout moderately short; E-N
70% of eye diameter; nostrils small, weakly protuberant, dorsolateraly directed; canthus rostralis slightly curved in
dorsal view, rounded in profile; loreal region slightly concave, lips weakly flared; interorbital area slightly convex,
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no cranial crests; upper eyelid without enlarged tubercles; EW 90% of IOD in male specimen. Tympanic
membrane and annulus absent; tympanic region with several irregular fused tubercles or warts, some of which are
arranged in a swollen fold (similar to a parotoid gland), extending from the posterior margin of the eye to the
shoulders. Oval to elongate, large tongue, weakly notched; choanae small, round, well separated and anterolaterally
situated; dentigerous processes of vomers absent. Vocal sac and vocal slits absent.
FIGURE 7. Holotype of Phrynopus capitalis sp. nov., male AMNH 134158 (SVL= 23.2 mm), showing (A) head in lateral
view and (B, C) body in dorsal and ventral views. Specimen collected near Lake Manachaqui, 3550 m, near the border of Río
Abiseo National Park, San Martín, Peru. Photographs by A. Catenazzi.
Skin of dorsum irregularly granular bearing some warts and small ridges, especially in the scapular area and on
the lower back, bearing scattered very small tubercles; skin of venter and throat coarsely areolate; two large
postrictal tubercles under the swollen supratympanic fold; a low elongate tubercle on the interorbital area;
metacarpal tubercles large, oval and low; outer areas of forelimbs bearing some elongate warts; inner palmar
tubercle nearly rhomboidal, thenar round, small slightly shorter than palmar; fingers bearing thick lateral fringes,
finger tips rounded and slightly bulbous; first finger slightly shorter than second; digital pads present; subarticular
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tubercles round, conical in profile (Figures 6B, 9A); THL 38% of SVL; two metatarsal tubercles, round and large,
inner one and a half the size of outer; some supernumerary plantar tubercles (Figure 6B). Toes bearing lateral
fringes and basal webbing; toe tips bulbous, emarginated; Toe V longer than Toe III.
FIGURE 8. Female paratopotype of Phrynopus capitalis sp. nov. in life (MUSM 8959, SVL = 35.6 mm) from around Lake
Manachaqui, 3600 m, near Rio Abiseo National Park, San Martín, Peru. Photograph by L. O. Rodriguez.
Color in life. Dorsal surfaces were brownish-gray on the scapular region on the holotype and brownish-tan
bluish black on the flanks; the venter was yellowish gray. Iris cocoa brown. Nuptial pads black.
Color in preservative. Dorsal surfaces are brownish-tan with small dark brown spots appearing like a
labyrinth; flanks are black with some brown on the warts (Figure 7). Groin with black reticulations; posterior thigh
black or brownish-tan, with the same dorsal pattern. Ventral surfaces grayish cream on the areoles with darker
grayish ground color.
Measurements in mm (data for male holotype followed by female paratopotype in parentheses). SVL
23.2 (35.9), HL 7.0 (13.6), HW 9.3 (14.3), TL (9.0 (12.0), THL 9.1 (13.0), FL 9.4 (13.9), IOD 2.9 (3.6), EW 2.6
(3.3), E-N 2.1 (2.8), IND 1.6 (2.7), ED 3.0 (3.7).
Vari a t i o n . Female MUSM 8959 is larger in size (35.6 mm), and EW is narrower than the IOD (wider in the
holotype). The two postrictal tubercles are also present. Coloration is the same as in the holotype. In life (Figure 8),
bluish black coloration under arms and above groin, more conspicuous than in male holotype.
Etymology. The specific name capitalis is a Latin adjective meaning “pre-eminent” and is used in reference to
the distinctive large head of the species
Distribution and ecology. Phrynopus capitalis is known only from the type locality in the Manachaqui Valley,
where it co-occurs with Phrynopus valquii and an undescribed species of Pristimantis. This valley is covered with
wet montane grasslands and contains a glacial lake that feeds a tributary of the Marañon river (Figures 2, 3C). The
three specimens were found within a 100 m
2
area in the wettest flat area of this valley, among dense moss
vegetation. This area lays outside of the Río Abiseo NP, but P. capitalis might be present in areas of the same
habitat within the park.
Phrynopus dumicola new species
(Figures 6C, 9B, 10, 11)
Holotype. MUSM 33102, a gravid female (Figures 6C, 10) obtained at Pampa del Cuy (7°39'49"S, 77°28'43"W;
3470–3550 m), ca. 18 km airline NE Pataz, Parque Nacional del Río Abiseo, Distrito Pataz, Provincia Mariscal
Cáceres, Departamento (Región) San Martín, Peru, collected on 19 July 2000, by A. Catenazzi and R. von May.
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FIGURE 9. Condition of terminal phalanges of Finger III, left hand: T-shaped in (A) Phrynopus capitalis sp. nov. (MUSM
8959), and knob-shaped in (B) P. dumicola sp. nov. (USNM 299788), (C) P. personatus sp. nov. (MUSM 3813), (D) P. valquii
(MUSM 3823). Drawings by F. Chang.
Paratopotypes. Eleven adult males, one adult female and one juvenile female: males MUSM 33092 and
33118, collected on 18 and 19 July 2000 by A. Catenazzi and R. von May; female MUSM 3819, collected on 1 July
1987; males MUSM 3815, 3817, AMNH 134167 (cleared and stained), and juvenile female MUSM 3820,
collected on 19 July 1987; males AMNH 134149–50 and 134151 (cleared & stained), collected on 29 July 1988;
males MUSM 9078–79, collected on 3 August 1990, all collected by L. O. Rodríguez.
Paratypes. Two adult males, one adult female and one juvenile male: adult male AMNH 134152 collected in
elfin forest between Pampa del Cuy and Puerta del Monte (ca. 7°40'S,77° 27'W; 3350 m) on 19 July 1988; adult
female MUSM 3818, adult male KU 220917, and juvenile male USNM 299788, collected at Puerta del Monte, ca.
20 Km airline NE Pataz, (7°39'29.64"S, 77°28'12.06"W; 3225 m) on 18 July 1987 by L. O. Rodríguez.
Diagnosis. This species of Phrynopus has: (1) skin on dorsum shagreen or rugose with low scattered tubercles,
skin on flanks areolate, skin on throat smooth, that of chest and venter areolate but not as coarsely as on flanks,
ventral surface of thighs coarsely areolate; discoidal fold absent, thoracic fold present; dorsolateral ridges present ;
(2) tympanic membrane and tympanic annulus absent, moderate supratympanic fold present; (3) snout short,
rounded in dorsal and lateral views; (4) upper eyelid without conical or rounded tubercles; EW narrower than IOD;
cranial crests absent; (5) dentigerous processes of vomers absent; (6) males lacking vocal slits and nuptial pads; (7)
Finger I slightly shorter or equal in length than Finger II; tips of digits rounded, terminal phalanges knob-shaped;
(8) fingers without lateral fringes; (9) ulnar and tarsal tubercles absent; (10) heels lacking tubercles; inner tarsal
fold absent; (11) inner metatarsal tubercle ovoid, about 1.5 times larger than rounded outer metatarsal tubercle;
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supernumerary plantar tubercles absent; (12) toes without lateral fringes; basal webbing absent; Toe III longer than
Toe V; toe tips rounded, about as large as those on fingers; (13) in life, dorsum dark brown, reddish brown or olive
green with irregular darker blotches, dark brown post orbital stripe present; throat varying from pearly white to pale
salmon or creamy yellow, chest and belly dull brown to creamy white or bluish white with or without brown
irregular blotches; ventral surfaces of forearms and thighs brown or salmon, ventral surface of hands and feet dark
brown with irregular creamy white flecks or blotches; groin dark brown, or reddish brown with pearly white flecks;
(14) SVL in females 22.3–25.3 mm (n = 5), in males 18.3–22.55 mm (n = 6).
FIGURE 10. Holotype of Phrynopus dumicola sp. nov., female MUSM 33102 (SVL = 23.34 mm), showing (A) head in lateral
view and (B, C) body in dorsal and ventral views. Specimen collected at Pampa del Cuy, 3470 m, within Río Abiseo National
Park, San Martín, Peru. Photographs by M. Flecks.
Seven other species of Phrynopus have aerolate skin ventrally, lack the tympanic annulus and dentigerous
processes of vomers, but can be distinguished from P. dumicola by a combination of the following characters
(character condition for P. d u m i c o l a in parentheses). Phrynopus barthlenae bears lateral fringes on all digits
(absent), has basal webbing on toes and nuptial pads in males (absent), and a dorsal pattern (absent). Phrynopus
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bufoides is larger with females up to 33.6 mm SVL (25.2 mm) and wears large warts on dorsum (absent), whereas
P. heimorum has a distinctive red coloration on ventral side of limbs (ventral coloration not red) and T-shaped
terminal phalanges (not expanded). Phrynopus daemon has a weakly areolate skin (areolate), wears lateral fringes
on toes (absent), and males have a subgular vocal sac (absent). Phrynopus horstpauli is larger with females up to
39.7 mm SVL, possesses a longer snout, toe V>III (toe III>V), and Finger I is much shorter than II. Phrynopus
oblivius has a longer snout (E-N/ED 68-69% in P. oblivious, 60-61% in P. dumicola) smooth dorsal skin, bears
ulnar tubercles (absent), has dark labial stripes (absent) and light small spots in venter (absent). Phrynopus valquii
also lacks a tympanic annulus and the dentigerous processes of vomers, but differs by having a shorter and wider
head, by lacking dorsolateral folds (discontinuous), by having shorter digits and bulbous digital tips (round tips),
by bearing irregularly T-shaped terminal phalanges (knob-shaped; see Fig. 9B and 9D), and by having lighter
dorsal coloration, and frontoparietals completely in contact (moderately separated).
Phrynopus chaparroi, P. paucari and P. thompsoni also share most characters with P. dumicola, but P.
chaparroi is larger, with females up to 32.2 mm in SVL, males wear nuptial pads (absent), bulbous toe tips (round),
and a dark venter with irregular, diffuse white to gray blotches (ventral coloration light and lacking a pattern).
Phrynopus paucari possesses a head that is longer than wide (HL < HW), has a discoidal fold (absent) and is bright
greenish-yellow on the hands and the inner side of limbs and flanks, with fine reticulations on venter in live
specimens (dull coloration ventrally, sometimes with salmon or dull pink on inner side of limbs). Phrynopus
thompsoni has a longer snout, wears longitudinal dorsal rows of low tubercles with pustules (dorsum shagreen with
low tubercles), lateral fringes on toes (absent), has Toe III = V (III > V) and many distinct brown spots on creamy
tan venter (venter without any spots).
Other species superficially resemble Phrynopus dumicola, but differ by having the dentigerous processes of
vomers. Of these, P. kotosh further differs by bearing small tubercles on heels (absent) and by having transversal
bars on thighs and lower frank (absent), whereas P. bracki differs by having a concealed tympanum (absent) and a
very short first toe (longer than wide).
FIGURE 11. Live Phrynopus dumicola sp. nov. (MUSM 33127) from Pampa del Cuy, 3470 m, Rio Abiseo National Park, San
Martín, Peru, in (A) dorsolateral and (B) ventral views. Note skin cysts caused by infestation with trombiculid mites.
Photographs A. Catenazzi.
Species of the Pristimantis orestes group with reduced digital pads may externally resemble Phrynopus
dumicola; those are P. atrabracus, P. chimu, P. cordovae, P mariaelenae, P. melanogaster, P. pataikos, P. pinguis, P.
seorsus, P. simonsii, P. stictoboubonus, and P. stipa. All of those species, except P. s i m o n s i i , have a visible
tympanum (P. atrabracus, P. chimu, P. cordovae, P. mariaelenae, P. pinguis), or the tympanic annulus is weakly
defined or concealed beneath the skin (P. melanogaster, P. pataikos, P. stictoboubonus, P. seorsus, P. stipa). The
other two members of the orestes group occurring in Peru, P. corrugatus and P. ventriguttatus bear well developed
pads on all digits and cannot be confused with Phrynopus dumicola. Pristimantis simonsii lacks a tympanum and
dentigerous processes of vomers, as in Phrynopus dumicola, but differs by bearing lateral fringes and basal
webbing (absent), by having relatively larger nasal bones, a shorter head and presenting black blotches on groins
(no marks). Pristimantis attenbouroughi is slightly smaller than dumicola, males SVL to 19.2 mm, females SVL to
23.0 (males SVL to 22.6mm, females SVL to 25.3 mm), bears the dentigerous processes of vomers (absent), has a
relatively longer snout with EN 70% of ED (EN 60% of ED), heel with a small conical ulnar and tarsal tubercles
(absent), a short cloacal sheat (absent) and lighter dorsal coloration with stripes on low flanks (absent). Pristimantis
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stipa externally resembles Phrynopus dumicola, but differs by bearing a supratympanic fold and tympanic annulus
(absent in P. dumicola), by bearing the dentigerous processes of vomers, bulbous digital tips (round), prominent
subarticular tubercles (weakly defined), and by having white mottling on venter, and white nuptial pads in dorsal
and medial surfaces of thumbs in males (absent).
Description of holotype. Head narrower than body, wider than long; HW 103% of HL; HW 37% of SVL;
snout nearly rounded in dorsal view, rounded in lateral profile; canthus rostralis rounded to moderately sharp in
lateral view; nostrils small, slightly protuberant, laterally directed; loreal region concave, sloping; snout short, E-N
86% of eye length; interorbital region flat, no cranial crests; EW 83% of IOD; tympanic membrane and annulus
absent; supratympanic fold short, not prominent bordered by a dark stripe; tongue oval, not notched and one-third
free posteriorly; choanae small, round, not hidden by palatal shelf of maxillae; dentigerous processes of vomers
absent; skin of dorsum shagreen with low scattered tubercles, low postocular folds, discontinuous dorsolateral
folds; some low small supraorbital tubercles; skin of venter coarsely areolate extending to the flanks and postrictal
area; skin of throat areolate; moderate thoracic fold present, no discoidal fold; no ulnar tubercles; weakly defined
palmar tubercle, palmar slightly smaller than thenar tubercle; fingers lacking lateral fringes; Finger I equal to
Finger II.
Hind limbs slender, short, TL 35% of SVL; upper surfaces of hind limbs smooth; posterior and ventral surfaces
of thighs coarsely areolate; heel and outer surface of tarsus lacking tubercles; two metatarsal tubercles, inner oval;
outer rounded, half size of inner; no supernumerary plantar tubercles; slightly areolate plantar surface; toes lacking
lateral fringes and basal webbing; toe tips rounded, lacking marginal grooves, about as large as those on fingers;
Toe III longer than Toe V (Fig. 4C).
Color in life. Overall dark brown dorsally and ventrally, with some light reddish areas on throat, chest and
upper venter. Iris bronze, turning darker towards the rounded pupil.
Color in preservative. Dark brown to brown dorsally and laterally, with a visible supratympanic stripe. Venter
brown to light brown with some darker spots.
Measurements of the holotype (in mm). SVL 23.3, HL 8.3, HW 8.6, TL 8.5, THL 9.2, FL 10.8, IOD 2.7, EW
2.3, IND 2.5, ED 3.2, E-N 2.7.
Vari a t i o n . Coloration pattern in some specimens (MUSM 3819 and 33116) includes a middorsal line that
extends onto back of thigh. Some specimens (e.g., MUSM 3815) have lighter areas on throat and venter. In darker
specimens, the supratympanic fold bordered by a dark stripe is barely visible (MUSM 9078 and 9079). The skin of
dorsum is usually shagreened, forming discontinuous dorsolateral ridges and occipital low folds.
Etymology. The specific name dumicola is a Latin adjective meaning "a dweller in thickets" in reference to the
habitat of forest patches bordering the tree line, inhabited by the species.
Distribution and ecology. This species is only known from Pampa del Cuy, Río Abiseo NP. Most specimens
were found in mosses on the ground of patches of elfin forest patches ranging from 0.01 to 5 ha in size. These elfin
forests grow on rocky substrates surrounded by wet montane grassland along a linear distance shorter than 3 km in
the Pampa del Cuy Valley (Figure 3B), from 3350 to 3550 m. The holotype was found in the grassland adjacent to
forest patches, during a rainy night. Phrynopus dumicola was the only species of frogs encountered in leaf-litter
plots in these elfin forest, while the surrounding grasslands contained other species of Pristimantis and
Gastrotheca. We found at least one P. d u m i c o l a specimen in every forest patch. We found some eggs (4.5 mm in
diameter) with adult specimens on 19 July 1987, suggesting parental clutch attendance. Most P. dumicola had
trombiculid mites under the skin, especially in the ventral area and limbs.
Phrynopus personatus new species
(Figures 6D, 9C, 12, 13)
Holotype. MUSM 33096, a gravid female (Figures 6D, 12) obtained near El Mirador (7°39'50"S, 77°26'55"W;
2890 m), ca. 20 km airline NE Pataz, Distrito Huicungo, Provincia Mariscal Cáceres, Departament (Región) of San
Martín, Peru, collected on 26 July 2000 by A. Catenazzi and R. von May.
Paratopotypes. MUSM 33166, an adult male collected with the holotype.
Paratypes. Five adult males and six adult females, all collected near the type locality, but at different elevations:
males MUSM 33121 (7°39'33"S, 77°28'07"W, 3260 m), MUSM 33109 (7°39'36"S, 77°27'46"W, 3180 m); females
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MUSM 33116 (7°39'36"S, 77°27'46"W, 3160 m), MUSM 33091 and 33099 (7°39'48"S, 77°28'26"W, 3110 m),
MUSM 33100 (7°39'43"S, 77°27'36"W, 3150 m), collected on 21–22 July 2000 by A. Catenazzi and R. von May;
female MUSM 3813 (7°39'45.92"S, 77°27'11.04"W, 3000 m), male AMNH 134153 and female MUSM 3811
(2930 m) collected on 23 July 1988 by L.O. Rodríguez; and males MUSM 3810 and 3812 (7°39´48” S, 77° 28´26”
W, 3100 m) collected on 26 July 1988 by L. O. Rodríguez.
FIGURE 12. Holotype of Phrynopus personatus sp. nov., female MUSM 33096 (SVL = 24.47mm), showing (A) head in
lateral view and (B, C) body in dorsal and ventral views. Specimen from El Mirador, 2890 m, Río Abiseo National Park, San
Martín, Peru. Photographs by M. Fleks.
Diagnosis. This species of Phrynopus has: (1) skin on dorsum shagreen with low scattered round tubercles on
dorsum and flanks, skin on flanks areolate; skin on throat, chest and belly finely areolate, ventral surface of thighs
coarsely areolate; discoidal fold absent, fine subgular and thoracic folds present; dorsolateral folds and an X shaped
medial ridge usually present; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal
and lateral views; (4) upper eyelid bordered by a dermal ridge and bearing some low rounded tubercles; width of
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upper eyelid narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers of vomers absent; (6)
males lacking vocal slits and nuptial pads; (7) Finger I slightly shorter than Finger II; tips of digits rounded,
terminal phalange knob-shaped (Fig. 9c); (8) fingers without lateral fringes; (9) ulnar and tarsal tubercles absent;
(10) heels lacking tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, about 1.5 times larger
than rounded outer metatarsal tubercle; supernumerary plantar tubercles absent; (12) toes without lateral fringes;
Toe V slightly longer than Toe III; toe tips rounded, about as large as those on fingers; (13) in life, dorsum dark
brown, reddish brown or olive green with irregular darker blotches, dark brown canthal and post orbital stripe
marking always a face mask; throat and ventral surfaces of forearms and thighs reddish brown to red-wine, ventral
surface of hands and feet lighter or dark brown with scattered creamy white flecks or blotches topping tubercles;
low venter and groin region dark brown, or black with sharp defined round white blotches, yellow in life,
sometimes extending onto the shank and also present behind the thigh; (14) SVL in females 23.4–28.2 mm (n = 7),
in males 17.5–21.1 mm (n = 7).
FIGURE 13. Live specimens of Phrynopus personatus sp. nov. (A, B) MUSM 33121, (C, D) MUSM 33100 and (E, F)
MUSM 33091 from near El Mirador, 2890 m, Río Abiseo National Park, San Martín, Peru. Photographs by A. Catenazzi.
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No other species of Phrynopus has such a combination of dorsal folds and masked dark face. Among
Phrynopus species lacking a tympanic annulus (present in P. auriculatus, P. montium, and P. peruanus) and the
dentigerous processes of vomers (character status for P. personatus in parentheses), P. capitalis is much larger,
lacks dorsolateral folds and has a different dorsal coloration pattern, possesses bulbous digital tips (rounded) and
wears lateral fringes in toes and fingers (absent). Phrynopus dumicola does not have a distinctive face mask, has
smaller nasal bones, a uniform dorsal coloration, and no light spots on groin whereas P. v a l q u i i has a weakly
defined canthus rostralis (sharp), much lighter coloration and bulbous digits (rounded in personatus). Among
species that exhibit bright coloration on groin (P. badius, P. bracki, P. dagmarae, P. heimorum, P. interstinctus, P.
nicoleae, P. paucari, P. peruanus, and P. vestigiatus), P. peruanus has a tympanic annulus (absent); P. bracki has
dentigerous processes of vomers (absent), and P. badius has smooth uniform brown ventral skin, scattered with
minute white dots and labial bar and very prominent subarticular at the base of fingers (absent in P. personatus); P.
paucari has a bright yellow venter. Phrynopus vestigiatus shares with P. personatus the presence of X ridges on the
scapular region, but has smooth dorsal skin, is smaller with single known female having SVL=18.8 mm (28.2 mm),
has minute dentigerous processes of vomers (absent), and a dark venter with small white flecks (light flecks
absent). Phrynopus dagmarae possesses light groin marks sharply bordered by black, but differs from P.
personatus by having the dentigerous processes of vomers and labial bars (both absent), different dorsal and ventral
coloration, and bulbous tip of fingers (rounded, not expanded). Phrynopus horstpauli has dorsal markings and large
nasal bones like P. personatus, but differs by its larger size, by lacking bright coloration on groin, and by having T-
shaped terminal phalanges (knobbed) and yellow-copper iris (upper iris golden with brown reticulations).
Phrynopus miroslawae is similar to P. personatus in having a supratympanic fold and broad dark canthal and
postorbital stripes delineating a face mask, and by bearing dorsolateral and occipital folds, but differs by lacking
bright coloration on groin, and by having bulbous digital tips (round), areolated ventral skin, and dorsal gray
coloration with dark blotches. Phrynopus nicoleae is similar to P. personatus in having first finger shorter than
second, and in bearing X-shaped dorsal marks and occipital and dorsolateral folds, but differs by its longer snout,
and by having labial bars and vomerine teeth. Among species of Phrynopus lacking tympanum and dentigerous
processes of vomers, P. c u r a t o r also has dorsal Y shaped ridges, and tubercles, but has a weakly areolate ventral
skin (aerolate), three tubercles on upper eyelid (dermal ridge with low tubercles), a conical tubercle on heel
(absent), longer limbs (TL/SVL= 49% in P. c u r a t o r, 33-35 % in P. personatus) and lacks bright coloration in the
groin (present in P. personatus). Species of the Pristimantis orestes group bearing reduced digital pads may
externally resemble Phrynopus personatus; those are P. atrabracus, P. chimu, P. cordovae, P mariaelenae, P.
melanogaster, P. pataikos, P. pinguis, P. seorsus, P. simonsii, P. stictoboubonus, P. stipa; all of which, except P.
simonsii, have either a visible tympanum, or the tympanic annulus is weakly defined or concealed beneath the skin
(in P. melanogaster, P. pataikos, P. stictoboubonus, P. seorsus and P. stipa). Furthermore, P. stipa, has a dark dorsal
skin, but differs from Phrynopus personatus by lacking medial dorsal ridges and dorsolateral folds, and bearing
prominent dentigerous processes of vomers (absent in P. personatus). Pristimantis simonsii also lacks the
dentigerous processes of vomers, but lacks dorsal folds and a distinctive face (both present in P. personatus), and
bright yellow or white spots on groin, on black ground (present). Pristimantis attenbouroughi differs by bearing
dentigerous processes of vomers (absent), by having the upper eyelid without dermal ridge (dermal ridge present,
bearing some low rounded tubercles), areolate ventral skin (finely areolate), heel with a small conical ulnar and
tarsal tubercles (absent), and a short cloacal sheath (absent).
Description of holotype. Head narrower than body, longer than width; HW 95% of HL; HW 37.8% of SVL;
interorbital area flat, no cranial crests; upper eyelid bearing several round supraorbital tubercles, EW 57% of IOD,;
snout semicircular in dorsal view, slightly truncate in profile; snout short, E-N 69% of eye length; canthus rostralis
sharp in lateral view, loreal region strongly sloping outward to lip, slightly concave; nostrils weakly protuberant,
directed dorsolaterally; lips not flared; tongue large, oval, posterior one-fourth free; choanae small, round, situated
well anterolaterally on palate, fairly separated; dentigerous processes of vomers absent; tympanic annulus and
tympanic membrane absent, supratympanic fold thick and prominent; several postrictal tubercles present,
extending onto the loreal area.
Skin of dorsum shagreened; prominent but incomplete dorsolateral ridges; flanks and upper surfaces of limbs
bearing scattered conical warts, lateral surfaces of limbs and paravertebral area bearing low small tubercles; an X-
shaped middorsal ridge formed by a series of closely packed warts starting postorbitally and joining the middorsal
line at the middle of the back, shorter and less prominent on posterior half; skin on throat shagreened, that of venter
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finely areolate; an outer row of tubercles present on forearms; palmar tubercle ovoid, thenar rounded barely
defined; subarticular tubercles low, simple; finger tips not bulbous; first finger shorter than second.
Tibia length 33% of SVL; showing an external row of tubercles on the heel and tarsal area, as an extension of
the warts on lateral surfaces of limbs; outer metatarsal tubercle flat, oval, barely defined, inner slightly smaller than
outer, rounded; supernumerary and subarticular tubercles not evident. Toes lacking lateral fringes and basal
webbing.
Color in life. The dorsal ground color was brown to reddish brown; venter, throat and inner forelimbs were
brown-red to red; bright to light rounded yellow spots on groin sharply bordered with black. Face and dorsal
surfaces as in preservative. Iris golden with fine dark reticulations on upper half, lower half dark brown.
TABLE 1. Measurements and proportions of two new species of Phrynopus. For measurements abbreviations see
Material and Methods. Mean and standard deviation are followed by range.
Color in preservative. The face is uniformly dark brown, with a faint line bordering the canthal stripe and the
tympanic fold. The tympanic and loreal regions underneath the faint line are dark brown, and form a pattern
resembling a mask. The upper lip is bordered by small cream blotches. A thin, pale cream middorsal stripe extends
from the internasal area to the vent. Above the cloacal opening, the stripe divides and extends across the thighs to
the lower edge of the knee, continuing on the posterior side of shanks to reach the heel, and forming an irregular
broad cream stripe (yellow in life) on the inner surface of the shank. Dorsal coloration is gray-brown above, with
dorsal folds, warts and ridges bordered with faint and dark brown, as on the face. Venter and concealed surfaces of
limbs, brown with dirty cream blotches. Rear surfaces of thigh and knees are dark brown.
P. dumicola sp. nov. P. personatus sp. nov.
♂ (n=6) ♀ (n=5) ♂ (n=7) ♀ (n=7)
SVL 20.1 + 0.5 24.1 + 0.6 19.2 + 0.5 25.3 + 0.6
(18.7–22.6) (22.3–25.3) (17.5–21.1) (23.4–28.2)
HL 6.9 + 0.2 8.0 + 0.4 6.8 + 0.2 8.9 + 0.3
(6.2–7.7) (7.2–9.4) (6.0–7.4) (7.7–10.0)
HW 7.5 + 0.2 8.6 + 0.4 7.0 + 0.2 9.3 + 0.3
(6.6–8.0) (7.9–10.1) (6.5–7.6) (8.1–10.4)
THL 7.2 + 0.1 8.8 + 0.4 7.8 + 0.2 9.3 + 0.2
(6.9–7.5) (8.1–9.5) (7.2–8.5) (8.7–10.2)
TL 6.9 + 0.1 8.2 + 0.2 6.7 + 0.2 8.4 + 0.1
(6.5–7.1) (7.6–8.9) (6.1–7.7) (8.2–9.2)
FL 8.5 + 0.2 10.0 + 0.2 8.5 + 0.3 10.0 + 0.3
(7.9–9.5) (9.5–10.8) (7.0–9.4) (8.3–11.3)
IOD 2.3 + 0.1 2.6 + 0.1 2.2 + 0.1 2.8 + 0.1
(2.1–2.7) (2.2–2.9) (1.9–2.5) (2.4–3.3)
EW 1.8 + 0.1 2.2 + 0.1 1.5 + 0.1 1.9 + 0.2
(1.6–2.1) (1.9–2.5) (1.3–1.9) (1.4–2.8)
E-N 1.5 + 0.1 1.9 + 0.2 1.6 + 0.2 1.8 + 0.1
(1.4–1.6) (1.3–2.7) (1.3–2.4) (1.4–2.4)
IND 2.0 + 0.1 2.6 + 0.2 2.0 + 0.2 2.6 + 0.1
(1.8–2.2) (2.2–3.1) (1.4–2.7) (2.2–3.1)
ED 2.5 + 0.2 3.1 + 0.2 2.4 + 0.2 3.0 + 0.3
(2.0–2.9) (2.6–3.8) (2.0–3.2) (2.3–3.9)
TL/SVL 34.9% 33.4%
HW/SVL 36.7% 36.6%
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Measurements (in mm) of the holotype. SVL 24.5, TL 8.1, FL 9.7, HW 9.3, HL 9.7, IOD 2.7, EW 1.6, IND
2.3, ED 2.3, E-N 1.6.
Vari a t i o n . Tibia length is 33% of SVL in females (n=7) and 35% in males (n=7). In some specimens, the
postympanic ridge does not continue over the eye. Some specimens bear small oblique ridges, formed by a row of
round tubercles (MUSM 3811 and MUSM 381, AMNH 134153), in the interorbital area. The dorsolateral folds are
not continuous in MUSM 33100 and completely absent in MUSM 33091. This last specimen has an additional
white spot (yellow in life) in the inner thigh, and two in the inner shank, as does MUSM 3811. In life, MUSM
9073, 90–74 and 9076, had very dark dorsal coloration and some ventral irregular spots. The thin pale cream mid-
dorsal line is persistent in all specimens but MUSM 33100, where the line is ill defined.
Etymology. The specific name personatus is a Latin adjective meaning masked and is used in reference to the
distinctive face mask of this species.
Distribution and ecology. Phrynopus personatus is known only from a narrow elevational range around the
type locality in the upper reaches of the continuous mountain forests of the Abiseo National Park. Fourteen
specimens were found in wet leaf litter of flat areas, all in the narrow elevational range from 2890 to 3110 m near
El Mirador (3000 m). The area is a continuous tropical montane rain forest about 15 m tall. Arboreal ferns are
common and tree trunks and ground surfaces are covered with moss, liverworts and lichens. Individuals of P.
personatus were notably inactive before and after capture. The reproductive stage of the holotype indicates that the
species may reproduce in the dry season. Males weighed 0.65– 0.95 g, females 1.90–3.05 g. Although no eggs
were found with the specimens, the large size of the well-developed eggs in the holotype strongly suggests direct
development. Some specimens had trombiculid mites under the skin (see cysts in Figures 13B, F).
Infection with Batrachochytrium dendrobatidis
All swabbed specimens were negative (Appendix 2), except for a specimen of P. dumicola (AMNH 134152)
collected on 19 July 1988 that had a low ZE = 9.56 zoospore equivalents.
Discussion
With the four species described herein, the genus Phrynopus now comprises 32 species, all endemic to Peru,
supporting the idea that many species await discovery in the eastern slopes of the Peruvian Andes, and that this
genus contains high regional diversity and endemism (Figure 1, Table 2). Our findings highlight this striking
diversity specifically for the region of Río Abiseo National Park, similarly to what has been reported for this genus
at Cordillera Carpish, Yanachaga-Chemillen NP, and other localities in central Peru (Hedges 1990; Lehr & Aguilar
2002, 2003; Chaparro et al. 2008; Trueb & Lehr 2008; Lehr et al. 2000, 2005, 2012, Lehr & Oróz 2012; Mamani &
Malqui 2014; Chávez et. al. 2015). The phylogenetic relationships and generic assignment of species of Phrynopus
of these two regions, and more broadly the biogeography of the genus in the Peruvian Andes, will not be fully
understood until molecular analyses can be conducted with a larger sample of species; the more recent analysis
included only eleven of twenty-eight nominal species (Padial et al. 2014).
The genus Phrynopus Peters, 1874, was first defined “by the lack of digital pads, simple digital tips, plantar
surfaces lacking conical or subconical supernumerary tubercles, having alary processes on the hyoid plate, and
having a “normal” skull architecture (i.e. not helmeted)” (Lynch 1975; Cannatella 1984). More recently, Phrynopus
sensu Lynch (1975) was found to be composed of several non-sister clades on the basis of molecular analyses, and
was split into different genera (Hedges et al. 2008). Although no synapomorphies are known for the genus,
Phrynopus was redefined as the clade containing most species of short-legged, terrestrial-breeding frogs having
head narrower than the body, digits lacking circumferential groves, terminal phalanges knob-shaped, and lacking
differentiated tympanic membrane and tympanic annulus in the high Andes of central and northern Peru. Most
species of Phrynopus lack cranial crests, and in most species the dentigerous processes of vomers are absent (but
present in P. anancites, P. auriculatus, P. bracki, P. dagmarae, P. interstinctus, P. kauneorum, P. kotosh, P.
lechriorhynchus, P. nicoleae, P. peruanus and P. vestigiatus). Finger I is usually smaller than Finger II (equal in
length in some species), Toe V slightly longer than Toe III, and digits can be narrow, rounded, or bulbous, with
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subarticular tubercles not projecting. Skin texture range from smooth to postulate dorsally, and smooth to areolate
ventrally, while SVL range from 14.5 mm in P. auriculatus to 54 mm in P. kauneorum.
The four species described herein are short-legged frogs lacking tympanum, columella and cavum
tympanicum, and only P. anancites wears dentigerous processes of vomers. We include in our comparisons a fifth
species, P. v a l q u i i , which we collected during our surveys and which occurs almost syntopically with P. anancites.
Examination of cleared and stained specimens and X-ray plates demonstrated relatively large nasal bones in P.
personatus, P. capitalis, and P. valquii, and small nasal bones in P. d u m ic ol a. The terminal phalanges show reduced
lateral processes in P. personatus, P. d u m i c o l a and P. v a l q u i i , while P. capitalis possesses moderate lateral
processes (Figure 9). In addition to having moderate lateral processes, P. capitalis also shares with species of the
Pristimantis orestes group the partial emargination of some digital tips. These similarities suggest that P. capitalis
could be related to species in the Pristimantis orestes group, but the absence of molecular data and morphological
synapomorphies for these genera prevents us from further exploring this hypothesis. Phrynopus dumicola differs
from P. valquii and P. capitalis by having reduced nasal bones and frontoparietals separated medially. Although the
characters presented here readily separate the new species from congeners, molecular information is currently
missing, preventing us from assessing the monophyletic relationships with other species of Phrynopus (Padial et al.
2014).
Species of Phrynopus occur in wet grasslands, elfin forests and rain forests from 2600 to 4490 m, between
latitudes 9°54´S (south of the Huancabamba depression) and 7°41´S (north of the Ene-Mantaro drainage; Figure 1,
Table 2). On the basis of available information, all species have small distribution ranges and, when co-occurring in
the same region, they segregate by elevation or by microhabitat type. A preliminary phylogeny including 11
species (Padial et al. 2014) suggests divergence between species inhabiting montane forests in the northern portion
of the distribution range, and species inhabiting the puna in central Peru. We lack sufficient sampling to examine
possible scenarios of diversification in the genus. Furthermore, several species inhabit fragmented elfin forests
(e.g., P. curator, P. dumicola, P. kotosh, P. lechriorhynchus, and P. oblivius).
The five species of Phrynopus of Río Abiseo NP occur along a linear distance shorter than 25 km, but are
separated spatially by microhabitat and elevation. Of these five species, only P. anancites and P. v a l q u i i are found
in proximity, share similar microhabitats in wet montane grasslands, and might occur sympatrically. Data from our
leaf-litter plots (which will be reported in detail in a different publication) indicate clumped distributions of most
species, which may be related to narrow microhabitat requirements and limited dispersal abilities. Phrynopus
capitalis inhabited a wet montane grassland at 3550 m, while P. valquii occurred in puna from 3550 to 3750 m and
wet montane grassland at 3850 m (similarly to P. anancites), P. d u m i c o l a was found in elfin forest between 3300
and 3550 m, and P. personatus in a narrow band (2930–3100 m) of continuous montane cloud forest well inside the
Park. These data suggest a fine altitudinal and habitat segregation within species of Phrynopus in the study area.
Small distribution ranges make species intrinsically more vulnerable to extinction, but at least two of the four
new species, P. dumicola and P. personatus, and P. v a l q u i i appear to have healthy populations within Río Abiseo
NP. According to the IUCN Red List criteria and categories (IUCN 2013), and considering that no anthropogenic
disturbance is currently threatening frog populations within the park, we propose to consider these two species for
the “Least Concern” (LC) category. A short survey within the park and surrounding areas (down to Pampa del Cuy)
in 2010 revealed the presence of Atelopus patazensis, Phrynopus valquii, P. dumicola and undescribed
Pristimantis. Furthermore, P. valquii has been described from specimens collected in 2014, after our last visit. The
other two species, P. anancites and P. capitalis, have not been collected within the park (although P. anancites was
found at the park’s boundary) and are known from few specimens; in absence of more detailed data regarding their
extent of occurrence, these species can provisionally be considered for the “Data Deficient” category of the Red
List.
The pathogenic fungus Batrachochytrium dendrobatidis has been reported from dying Atelopus patazensis
near Río Abiseo NP on 21 June 1999 (Venegas et al. 2008), which is the oldest case of chytridiomycosis causing
amphibian die-offs in Peru (Catenazzi & von May 2014). Of nine preserved specimens, all were negative, except
for P. dumicola (AMNH 134152) that had a low ZE = 9.56 zoospore equivalents. This specimen had been collected
on 19 July 1988, earlier than infected and dying Atelopus patazensis from nearby Pataz (Venegas et al. 2008).
Although this fungus can cause dramatic declines in species richness and abundance of stream-breeding frogs
(Catenazzi et al. 2011, 2014), it generally does not threaten terrestrial-breeding frogs in the Peruvian Andes
(Catenazzi & Ttito 2016, Catenazzi & von May 2014), and we did not observe dead specimens of Phrynopus and
other terrestrial-breeding frogs during our surveys in Río Abiseo NP.
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TABLE 2. Geographic and elevational ranges of species of Phrynopus.
Species Known distribution Elevation Habitat
P. anancites sp. nov. La Libertad: Ventanas 3820 m puna
P. auriculatus
1,2
Pasco: Oxapampa 2600 m montane forest
P. badius
3
Pasco: PN Yanachaga-Chemillén 2900 m montane forest
P. barthlenae
1
Huánuco: Maraypata 3425–3770 m puna
P. bracki
1
Pasco: Oxapampa 2600 m montane forest
P. bufoides
1,4
Pasco: Paucartambo 3850–4100 m puna
P. capitalis sp. nov. La Libertad: Pataz 3600 m puna
P. chaparroi
5
Junín: Comas 4205–4490 m puna
P. curator
2
Pasco: PN Yanachaga-Chemillén 3000 m montane forest/puna
P. daemon
6
Huánuco: Carpish 3140–3340 m montane forest
P. dagmarae
1,7
Huánuco: Chaglla (Carpish) 3020–3380 m montane forest
P. dumicola sp. nov. San Martín: PN Río Abiseo 3225–3550 m elfin forest/puna
P. heimorum
1
Huánuco: Conchamarca 3420 montane forest
P. horstpauli
1
Huánuco: Conchamarca 3010–3770 m montane forest
P. interstinctus
9
Huánuco: Umari (Carpish) 3100–3180 m montane forest
P. juninensis
1
Junín, Pasco (Cordillera Oriental)3420–3850 m puna
P. kauneorum
1,7
Huánuco: Chaglla (Carpish) 3020–3380 m montane forest
P. kotosh
1,10
Huánuco: Huancapallac 2950 m montane forest
P. lechriorhynchus
1
Huánuco: Acomayo 2740–2800 m montane forest
P. miroslawae
1
Pasco: Oxapampa 3363 m elfin forest
P. montium
1
Junín (several localities) 3000–4012 m puna
P. nicoleae
1
Pasco: Oxapampa 3589 m puna
P. oblivius
1,10
Junín: Maraynioc 3210–3220 m montane forest
P. paucari
1,4
Pasco: Paucartambo 3600 puna
P. personatus sp. nov. San Martín: PN Río Abiseo 2930–3100 m montane forest
P. peruanus
1
Junín: Maraynioc 3825 m puna
P. pesantesi
1,4
Pasco: Huachón 4280–4390 m puna
P. tautzorum
1
Huánuco: Maraypata 3770 m puna
P. thompsoni
1
La Libertad: Yamobamba 3290 m puna
P. tribulosus
1,2
Pasco: Oxapampa 2600 m montane forest
P. valquii
6
La Libertad: Parcoy 4025 – 4125 m puna
P. vestigiatus
9
Huánuco: Chinchao (Carpish) 3100 m montane forest
1
Duellman, W. E. and E. Lehr. 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Nature und Tier Verlag, Münster,
Germany, 382 pp.
2
Duellman, W. E., and S. B. Hedges. 2008. Two minute species of Phrynopus (Lissamphibia: Anura) from the Cordillera
Oriental in Peru. Zootaxa 1675: 59–66.
3
Lehr, E., J. Moravec, and J. C. Cusi. 2012. Two new species of Phrynopus (Anura, Strabomantidae) from high elevations in the
Yanachaga-Chemillén National park in Peru (Departamento de Pasco). ZooKeys 235: 51–71.
4
Lehr, E., M. Lundberg, and C. Aguilar. 2005. Three new species of Phrynopus from central Peru (Amphibia: Anura:
Leptodactylidae). Copeia 2005: 479–491.
5
Mamani, L., and S. Malqui. 2014. A new species of Phrynopus (Anura: Craugastoridae) from the central Peruvian Andes .
Zootaxa 3838: 207–214.
6
Chávez, G., R. Santa Cruz, D. Rodríguez, and E. Lehr. 2015. Two new species of frogs of the genus Phrynopus (Anura:
Terrarana: Craugastoridae) from the Peruvian Andes. Amphibian & Reptile Conservation 9(1, e105): 15–25.
7
Lehr, E., C. Aguilar, and G. Köhler. 2002. Two sympatric new species of Phrynopus (Anura: Leptodactylidae) from a cloud
forest in the Peruvian Andres. Journal of Herpetology 36: 208–216.
8
Lehr, E. 2001. A new species of Phrynopus (Anura: Leptodactylidae) from the eastern Andean slopes of central Peru.
Salamandra 37: 11–20.
9
Lehr, E., and A. Oróz. 2012. Two new species of Phrynopus (Anura: Strabomantidae) from the Cordillera de Carpish in central
Peru (Departamento de Huánuco). Zootaxa 3512: 53–63.
10
Lehr, E. 2007. New eleutherodactyline frogs (Leptodactylidae: Pristimantis, Phrynopus) from Peru. Bulletin of the Museum of
Comparative Zoology 159: 145–178.
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Chávez et al. (2015) suggested Phrynopus valquii to be Data Deficient, in absence of known localites within
the park. Here we confirm that P. valquii does indeed occur within Río Abiseo NP, because we found MUSM 9085
brooding a clutch of 20 eggs at the puna of Laplap (3850m), MUSM 3827 and AMNH 134155 at Paredones (3635
m), and MUSM 3824 near Lake Manchaqui (Figure 2). Therefore, similarly to P. d u m i c o l a and P. capitalis, we
propose the LC category for P. valquii. We recommend continuous monitoring of frog populations inside the park
to provide stronger support for future assessments of the conservation status of these species (Catenazzi & von
May 2014).
Acknowledgments
We thank M. Leo, M. Romo, E. Ortiz, R. von May, M. Fernandez, D. Aguilar and M. Silva for support during field
expeditions. Logistic support was provided by T. and I. Hurtado, P. Moore, the friendly people from Pataz, E.
Alayo, the park administration and rangers. LOR is especially grateful to D.W. Davidson for providing all
necessary support, working space and constructive comments on early version of the manuscript, C.W. Myers
(AMNH) , G.R. Zug (USNM), W.E. Duellman (KU), B. Clarke and C. McCarthy (BMNH) and G. Köhler (SMF)
kindly provided working space and facilities for specimen examination of collections at their care. We thank S.B.
Hedges for help with species comparisons; K. Tige for X-ray plates and G. Zug (USNM) and J. Icochea for clearing
and staining. We thank W.E. Duellman and L. Trueb for their hospitality during our visits to KU. LOR thanks C.W.
Myers for help with the initial descriptions and for extending many courtesies during her visits to the AMNH, L.S.
Ford for her help with defining localities, F. Chang (+) for drawing Figure 9 and M. T. Fuentes (CIMA-Cordillera
Azul), for Figure 2; D. Rödder for providing working space at the König Zoological Museum of Bonn, and M.
Flecks for assisting with holotype photographs.
The inventories conducted by the Asociación Peruana para la Conservación de la Naturaleza (APECO) in Río
Abiseo NP were funded by grants from the David and Lucille Packard Foundation and the PEW Charitable Fund,
through the University of Colorado at Boulder. Visits to AMNH were supported through a Collection Study Grant
and to KU with support from the University of Kansas. A visit to the National Museum of Natural History,
Smithsonian Institution was supported by the Biological Diversity Program in Latin America (BIOLAT). The
Dirección General Forestal y de Fauna of the Ministerio de Agricultura, Lima, provided the necessary permits for
scientific collections within Río Abiseo National Park.
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APPENDIX I. Specimens examined
Phrynopus bracki: PERU: P
ASCO
: 2.9 km N, 5.5 km E (airline) Oxapampa, Cordillera Yanachaga (2600 m), USNM 286918
(holotype), 286919, MUSM 4400 (paratype).
Phrynopus chaparroi: PERU: J
UNÍN
: Canchapalca, Distrito de Comas, Provincia Concepción, (11°44'45" S, 74°58'47"W):
MHNC 10983 (holotype, 4490 m.), MHNC 10980–10982 (4490 m.), MHNC 10984 and MHNC 10985 (4205 m.).
Phrynopus dagmarae: PERU: H
UÁNUCO
: Distrito Chaglla, Provincia Pachitea, Chaglla-Palteanan (09°51´27"S, 75°53´12"W):
SMF 80480–83; Maraypata (10°09´35"S, 7´6°06´05"W, 3370 m), SMF 80475 (paratype); Palma Pampa (09°53´12"S,
75°53´22"W): SMF 80476–79 and 80629–33 (paratypes); E slope Cordillera Carpish, Carretera Central, 2400 m: KU
196592.
Phrynopus heimorum: PERU: H
UÁNUCO
: Provincia Ambo, Conchamarca: SMF 80470, 80474 (09°59´44”S, 76°09´40”W,
3420m); 500 m east of Conchamarca (09°59´44”S, 76°09´40”W, 3420m): SMF 80471–80472.
Phrynopus horstpauli: PERU: H
UÁNUCO
: Provincia Ambo, 7 km east of Conchamarca, Huacamonte forest (09°5955”S,
76°09´43”W, 3070 m): SMF 80447–52, 80459–60; Huacamonte forest (09°59´41”S, 76°09´44”W, 3420 m): SMF 80466–
67; Ichocan, Jatunloma, 3100 m: KU 291399–291400, 311452: 10 km E of Conchamarca, 3420 m: KU 311453.
Phrynopus kauneorum: PERU: H
UÁNUCO
: Chaggla, SMF 80626–28, SMF 80484–85, AMNH 311451.
Phrynopus montium: PERU: J
UNÍN
: Tarma, 45 min. of Maranyioc (12000 ft = 3658 m), USNM 217416–17; J
UNÍN
: Prov. Yauli,
9.5 min. from La Oroya (13000 ft = 3962 m), AMNH 84795.
Phrynopus valquii: PERU: S
AN
M
ARTÍN
: Río Abiseo NP: MUSM 3824, MUSM 3821, 3823, KU 220918 and AMNH 134154–
55.
Pristimantis simonsii: PERU: C
AJAMARCA
: 23 km S Celendín, 3050, KU 181357- -181360; 33 km S Celendín, 3200 m, KU
181361--181391, 181830--181833, MUSM 1526--1527, 1546-1548; NE Encanada-, 23,5 km from Cajamarca, 3510 m,
MUSM 1160 (cleared and stained), 1161--1179; C
AJAMARCA
, 3500
M
, BM 1900.3.30.22-23, Lectotype "Paramo,
Cajamarca, 9000 feet".
APPENDIX 2. Museum specimens analyzed for the presence of Batrachochytrium dendrobatidis through skin swabbing
(November 2015) and quantitative PCR (see methods). ZE = zoospore equivalents
Specimen # Species Collection date Status ZE
AMNH 134158 P. capitalis 3 July 1989 Negative
AMNH 134149 P. dumicola 29 July 1988 Negative
AMNH 134150 P. dumicola 29 July 1988 Negative
AMNH 134152 P. dumicola 19 July 1988 Positive 9.57
KU 220917 P. dumicola 18 July 1987 Negative
AMNH 84795 P. montium 13 June 1964 Negative
AMNH 134153 P. personatus 23 July 1989 Negative
AMNH 134154 P. valquii 23 July 1987 Negative
AMNH 134155 P. valquii 28 July 1987 Negative
... We follow Lynch & Duellman (1997), Rodríguez & Catenazzi (2017) and for format of description, as well as character definitions given by Duellman & Lehr (2009). We follow Hedges, Duellman & Heinicke (2008) and Heinicke et al. (2017) as support of our phylogenetic analyses for family placement. ...
... We currently lack evidence of any synapomorphic phenotypic trait for Phrynopus that allows distinguishing species in this genus from Pristimantis and other Terrarana with similar body shape and anatomical structures. These phenotypic features (referred to Phrynopoid ecomorph by De la Riva, 2020) presumably converged in response to the conditions of high elevation environments (Padial, Grant & Frost, 2014;Rodríguez & Catenazzi, 2017). Therefore, based on the results of our phylogenetic analyses, overall body shape and combination of meristic traits, we assign the new species to the genus Phrynopus. ...
... Similar to most congeneric species, Phrynopus remotum inhabits humid puna grasslands in the Peruvian Andes (Lehr, Fritzsch & Müller, 2005;Chávez et al., 2015;Lehr & Rodriguez, 2017;Rodríguez & Catenazzi, 2017). Several Phrynopus species have been reported from two or more localities (Trueb & Lehr, 2008;von May, Lehr & Rabosky, 2018), but so far P. remotum is only known from the type locality; however, we are confident that future exploration in surrounding areas could detect the new species, and contribute to better documenting its distribution range. ...
Article
Full-text available
We describe a new, medium-sized species of terrestrial frog of the genus Phrynopus from a single locality in the central Andes of Peru (Departamento de Huánuco) at 3,730 meters of elevation. Phylogenetic analyses supported Phrynopus remotum sp. nov. as an independent lineage, sister to most of its congeners. The new species is morphologically distinguishable by the presence of small tubercles on upper eyelids and heels, an areolate venter, and the absence of dorsolateral folds or ridges. This species inhabits the highlands adjacent to the Marañón Dry valley. The only sympatric amphibian species recorded is the marsupial frog Gastrotheca peruana.
... Thus, the festae species Group is currently considered to contain the following 19 species [3,6,7,14] which are distributed in Panama, Colombia, Ecuador, Peru, and Bolivia ( Figure 1): R. acrolopha (Trueb, 1971) [10], R. arborescandens (Duellman and Schulte, 1992) [4], R. chavin, R. chullachaki Castillo-Urbina, Glaw, Aguilar-Puntriano, Vences and Köhler, 2021 [14], R. festae, R. lilyrodriguezae, R. lindae (Rivero and Castaño, 1990) [15], R. macrorhina, R. manu, R. multiverrucosa (Lehr, Pramuk and Lundberg, 2005) [16], R. nesiotes, R. nicefori (Cochran and Goin, 1970) [17], R. paraguas Grant and Bolívar-G., 2014 [18], R. rostrata, R ruizi (Grant, 2000) [19], R. tacana (Padial, Reichle, McDiarmid and De la Riva, 2006) [20], R. tenrec (Lynch and Renjifo, 1990) [21], R. truebae (Lynch and Renjifo, 1990) [21], and R. yanachaga. Herpetological surveys of the Río Abiseo National Park in northern Peru, Departamento San Martín, in the late 1990s led to the discovery of six new species of anurans: four Phrynopus [22], one Pristimantis [23], one Gastrotheca [24], and an unknown species of Rhinella. Recent herpetological surveys in northern Peru (Departamentos San Martín and Amazonas) by PJV and LAGA led to the discovery of several specimens of the same unknown species of Rhinella. ...
... Herein we describe this new species of Rhinella from northern Peru (Figures 1 and 2), which we tentatively assign to the festae species Group based on morphological characters. Herpetological surveys of the Río Abiseo National Park in northern Peru, Departamento San Martín, in the late 1990s led to the discovery of six new species of anurans: four Phrynopus [22], one Pristimantis [23], one Gastrotheca [24], and an unknown species of Rhinella. Recent herpetological surveys in northern Peru (Departamentos San Martín and Amazonas) by PJV and LAGA led to the discovery of several specimens of the same unknown species of Rhinella. ...
Article
Full-text available
We describe a new species of Rhinella from montane forests between 1788 and 2305 m a.s.l. in the Departamentos Amazonas and San Martín, Peru. We tentatively assign the new species to the Rhinella festae species Group based on morphological similarities with its other 19 members. It is characterised by large size (maximum SVL 91.6 mm in females), a pointed and protruding snout that is posteroventrally inclined, absence of a visible tympanic annulus and tympanic membrane, long parotoid glands in contact with upper eyelid, presence of a dorsolateral row of enlarged tubercles, outer dorsolateral tarsus surface with a subconical ridge of fused tubercles, and absence of subgular vocal sac and vocal slits in males. One specimen from Departamento Amazonas tested positive for Batrachochytrium dendrobatidis.
... 1500 m to above 4000 m), is one of the most biodiverse forests in Peru (Leo 1995;INRENA 2003). Recently, many new species have been described from this park, mainly amphibians (Lehr et al. 2013;Rodriguez & Catenazzi 2017), increasing the number of endemic species for Peru, highlighting the biodiversity value in this park and the Yungas (Leo 1995). In rodents, Thomasomys apeco and T. macrotis are known only from Abiseo National Park (Leo L. & Gardner 1993;Gardner & Romo 1993). ...
Article
Nephelomys albigularis is a sigmodontine rodent of the tribe Oryzomyini distributed in the Andean forests from central Ecuador to central Peru. Although several studies recognize this species as monotypic, significant morphological variation has been reported in Peruvian populations that were not properly assessed by direct comparisons with the type series from central Ecuador. We present a preliminary review of N. albigularis with an integrative approach and emphasis on Peruvian populations. We analyzed specimens using morphological and morphometric methods, complemented with phylogenetic analyses and species delimitation using sequence data from the cytochrome-b gene. Our results reveal that N. albigularis (sensu lato) comprises two taxa: N. albigularis s.s., from the montane forests in central and southern Ecuador and northwestern Peru, and Nephelomys sp. nov. from montane forest east of the Marañón River. These taxa are morphologically distinct and are separated by a genetic distance of 5.90 ± 1.01%. Nephelomys sp. nov. differs from N. albigularis s.s. by longer rump hairs, narrow hypothenar pads, faintly bicolor tail; absent interorbital ridges, low zygomatic plates, smoothly squared posterior margin of the hard palate or with a small median postpalatal process, smoothly edged ventral margin of the external auditory meatus, and slightly angular mandibular sigmoid notches. In this work, we present a diagnosis and description of the new species of Nephelomys and discuss the role of the Marañón River as a potential driver for speciation in the genus Nephelomys.
... Nevertheless, species diversity in this group constantly grows with the discovery of new species when remote regions are surveyed, museum specimens are examined, or cryptic species complexes are phylogenetically analyzed (e.g. Elmer and Cannatella [11]; Ortega-Andrade and Venegas [12]; Motta et al. [13]; Rodríguez and Catenazzi [14]; Páez and Ron [15]; Santa-Cruz et al. [16]). ...
Article
Full-text available
We describe two new species of terrestrial-breeding frogs from the Cordillera de Colán, in northeastern Peru. We used Parsimony and Maximum Likelihood approaches to infer a molecular phylogeny on a dataset composed of 75 terminals, including three terminals representing the new species, and 4202 bp of concatenated mtDNA and nuDNA fragments. Our phylogenetic analyses support the placement of the two new species in Lynchius and Oreobates, respectively. The new species of Lynchius occurs in two localities from 1,977 to 2,006 m a.s.l., and is characterized by having a dorsum covered by conical tubercles and a brown dorsal coloration lacking a pattern of blotches on the hidden surfaces of flanks and hindlimbs. The new species of Oreobates is only known from one location at 2608 m a.s.l. and is characterized by the absence of axillary and inguinal glands, and the presence of white or cream blotches on the dark brown hidden surfaces of the body.
... However, the herpetofauna of this region is far from being well surveyed and new species of anurans are continuously discovered, especially in remote localities (e.g. Duellman and Venegas 2005;Venegas and Barrio 2005;Lehr and Catenazzi 2011;Rivera-Correa et al. 2016;Rodriguez and Catenazzi 2017). ...
Article
Full-text available
We describe a new species of marsupial frog, genus Gastrotheca , using morphological characters and molecular data as lines of evidence. The new species was discovered in the páramo and the ecotone between páramo and humid montane forest of Cordillera de Colán, at elevations between 3136 and 3179 m a.s.l., in northeastern Peru. The new species is distinguished from all its congeners by the combination of the following characters: coarsely granular skin on dorsum, a green dorsal coloration without pattern, finger I shorter than finger II, turquoise iris, and a venter without blotches, flecks or dots. Furthermore, we include a detailed osteological description of the new Gastrotheca species based on Micro-CT scanning. Based on our phylogenetic analyses, the new species belongs to the Gastrotheca marsupiata species group, is sister to G. oresbios and closely related to G. psychrophila , G. spectabilis , G. stictopleura and one undescribed species. Additionally, we test for the presence of the fungal pathogen Batrachochytrium dendrobatidis ( Bd ). No Bd infection was detected for G. gemma sp. nov. specimens but Bd prevalence was detected among syntopic frogs.
... Its western part present high rainfall and low seasonality, as well as high anuran diversity and endemism rates (Duellman 1999), in which a number of complexes cryptic species have been recently recorded (e.g., Ferrão et al. 2016;Caminer et al. 2017;Rojas et al. 2018). A similar pattern is recorded in the Andes as well, where many new species have been discovered (mainly craugastorids and hylids) in the wet, high-altitude forests (e.g., River-Correa et al. 2015; De la Riva and Aparicio 2016;Navarrete et al. 2016;Rodriguez and Catenazzi 2017). The second region having recently described species in South America encompasses much of the mountains of the Atlantic Forest domain, in which many anuran species recently described have restricted ranges associated with mountains (e.g., Ribeiro et al. 2015;Roberto et al. 2017). ...
Chapter
Understanding the spatial distributions of phylogenetic diversity is an opportunity to support policy-makers and designing conservation strategies in megadiverse regions. Here, we mapped the spatial distribution of Faith’s phylogenetic diversity (PD) and phylogenetic endemism (PE) of amphibian species distributed across South America. Although we found areas of high species richness (SR) correlated with areas of high PD or PE, there are regions with much more PD/PE or much less PD/PE than expected given the SR. Using a phylogenetic approach, we found that the factors regulating amphibian biodiversity involve a complex interplay of evolutionary and biogeographical processes in different regions of South America. These results might help supporting conservation planning for this threatened vertebrate group.
... Its western part present high rainfall and low seasonality, as well as high anuran diversity and endemism rates (Duellman 1999), in which a number of complexes cryptic species have been recently recorded (e.g., Ferrão et al. 2016;Caminer et al. 2017;Rojas et al. 2018). A similar pattern is recorded in the Andes as well, where many new species have been discovered (mainly craugastorids and hylids) in the wet, high-altitude forests (e.g., River-Correa et al. 2015;De la Riva and Aparicio 2016;Navarrete et al. 2016;Rodriguez and Catenazzi 2017). The second region having recently described species in South America encompasses much of the mountains of the Atlantic Forest domain, in which many anuran species recently described have restricted ranges associated with mountains (e.g., Ribeiro et al. 2015;Roberto et al. 2017). ...
Chapter
Amphibians are especially diverse in the Neotropics and have also one of the highest rates of new species description among terrestrial vertebrates. The first systematic synthesis of South American anurans compiled a list of 1644 species, but there have been no update since the last 19 years. Here, we present a descriptive approach for temporal and spatial patterns of anuran species discoveries in South America, emphasizing trending changes in species description rates and number of researchers authoring a given species description. We recovered 2623 anuran species described in South America between 1758 and mid-2017 from 163 genera and 24 families. There is a high rate of species discovery across time, with at least 10 new descriptions per year in the period examined. Time span to reach multiples of 500 new species has dramatically decreased over time. For instance, it took more than two centuries for the description of 500 species since the first species (1750s), whereas it took about 10–12 years in order to add 500 new anuran species after 1990. Then, the curve of the cumulative anuran species description in South America is far from reaching an asymptote, yet it actually exhibits an exponential shape. Similar historical increase was recorded for the number of authors in papers over time, since descriptions are more collaborative in the last decades. Two major hotspots for new species discovery are depicted herein: (i) the Central and Northern Andes and the adjacent western Amazon (notedly in Ecuador, Peru, and Western Brazil) and (ii) the complex of Brazilian highlands encompassing the Atlantic and Brazilian plateau mountains. These trends are discussed according to singular historical events (including changes in research approaches) and possible explanations for the geographic pattern in species discovery.
... Its western part present high rainfall and low seasonality, as well as high anuran diversity and endemism rates (Duellman 1999), in which a number of complexes cryptic species have been recently recorded (e.g., Ferrão et al. 2016;Caminer et al. 2017;Rojas et al. 2018). A similar pattern is recorded in the Andes as well, where many new species have been discovered (mainly craugastorids and hylids) in the wet, high-altitude forests (e.g., River-Correa et al. 2015; De la Riva and Aparicio 2016;Navarrete et al. 2016;Rodriguez and Catenazzi 2017). The second region having recently described species in South America encompasses much of the mountains of the Atlantic Forest domain, in which many anuran species recently described have restricted ranges associated with mountains (e.g., Ribeiro et al. 2015;Roberto et al. 2017). ...
Book
This book analyzes different facets of anuran amphibian distribution in South America. We integrate alternative biological metrics employing cutting-edge methods to understand the dynamic processes underlying species distribution patterns. By using the modern biogeographic toolbox, we explore how richness gradients, phylogenetic diversity, functional diversity, and range size/endemism distribution of amphibians vary along the continent. Moreover, we present a robust proposal for priority areas for conservation of anurans in South America that maximizes representativeness of distinct biodiversity facets.
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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.
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