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A new terrestrial frog (Anura: Craugastoridae) from the montane cloud forests
of the southeastern Ecuadorian Andes

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DOI: 10.11646/zootaxa.4318.3.5

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 Zootaxa 4318 (3): 520–530 ISSN 1175-5326 (print edition)
http://www.mapress.com/j/zt/
Copyright © 2017 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4318.3.5
http://zoobank.org/urn:lsid:zoobank.org:pub:1215DB76-4A16-4C50-9D97-76F98B2A4517

A new terrestrial frog (Anura: Craugastoridae) from the montane cloud forests
of the southeastern Ecuadorian Andes

VERONICA L. URGILES1,2,4,5, VALENTINA POSSE1, BRUNO A. TIMBE1,

PEDRO X. ASTUDILLO3 & JUAN C. SÁNCHEZ-NIVICELA1,2
1
Laboratorio de Herpetología, Museo de Zoología de la Universidad del Azuay MZUA, Av. 24 de Mayo 7–77, P.O. box 01-01-981,
Cuenca, Ecuador.
2
Instituto Nacional de Biodiversidad del Ecuador, Pasaje Rumipamba 341 y Av. de los Shyris, Postal code 17–07–8976, Quito-Ecua-
dor.
3
Escuela de Biología, Ecología y Gestión, Universidad del Azuay, Av. 24 de Mayo 7–77, P.O. box 01.01.981, Cuenca, Ecuador.
4
Department of Biology, University of Central Florida, 4110 Libra Drive, Orlando, FL 32816, USA.
5
Corresponding author. E-mail: vurgiles@uazuay.edu.ec

Abstract

We describe a new species of Pristimantis from the Tinajillas-Rio Gualaceño Ecological Conservation Area, Morona San-
tiago province, an area of montane forest in the southeastern Andes of Ecuador. Pristimantis nimbus sp. nov. is similar to
P. altamazonicus, P. ardyae, P. bambu, P. bellator, P. caeruleonotus, P. cethospilus, P. churuwiai, P. croceoinguinis, P.
cryptomelas, P. diadematus, P. flavobracatus, P. gualacenio, P. nigrogriseus, P. ventrimarmoratus, and P. versicolor from
which it can be distinguished by the presence of flash marks on flanks, groin, and arm insertion composed of bright-yellow
oval spots and blotches surrounded by dark brown to black, forming a reticulated pattern, by having slightly enlarged toe
pads, a finely shagreen dorsum lacking dorsal folds, and males having nuptial pads but lacking vocal slits. Pristimantis
nimbus sp. nov. is only known from the type locality, at elevations between 2200–2400 masl. All individuals of the new
species were found at night, perching on leaves and branches between 80 cm to 150 cm above the ground. The analysis
of stomach contents rendered a total of 17 prey items of different unidentified species in five orders (Aranae, Coleoptera,
Diptera, Hemiptera, Lepidoptera). Other species found in the area are Pristimantis aff. altamnis, P. galdi, P. proserpens,
P. tinajillas, P. versicolor, Rhinella margaritifera, Noblella sp. and three unidentified species of Pristimantis.

Key words: Brachycephaloidea, montane forest, Morona Santiago, new species, Pristimantis, Terrarana

Introduction

The Ecuadorian Andes harbors ~ 30% of the amphibian diversity of the country (Ron et al. 2016), and the area is
particularly rich in frogs, especially in species in the genus Pristimantis (Lynch 1979; Guayasamín et al. 2004;
Bustamante & Mendelson 2008; Guayasamín & Arteaga 2013; Navarrete et al. 2016). New amphibian species are
constantly being discovered and described from the eastern Andes of Ecuador, particularly Pristimantis from
montane forests (Batallas & Brito 2014; Urgiles et al. 2014; Yanez et al. 2014; Hutter & Guayasamín 2015; Brito et
al. 2016; Navarrete et al. 2016). Several factors have been identified as potential explanations for the outstanding
diversity of this group in the Ecuadorian Andes, which include low dispersal abilities, the geographic complexity
of the Andes, and reproductive strategies independent of water bodies (Guayasamín & Arteaga 2013; Hedges et al.
2008; Wells 2007; Duellman & Lehr 2009; Gonzalez-Voyer et al. 2011).
Despite the high diversity of amphibians in this region, sampling efforts and geographic coverage are relatively
low, and even national parks remain poorly sampled (Navarrete et al. 2016). This is especially true for smaller
protected areas such as private lands or local-government protected lands, where amphibian surveys have been
rare. Within this context, in 2015, herpetologists from Universidad del Azuay conducted an expedition to the
montane forests of the Tinajillas—Rio Gualaceño Ecological Conservation Area, a small protected area

520 Accepted by J. Padial: 27 Jul. 2017; published: 8 Sept. 2017

administrated by the local government. These surveys resulted in the collection of eight specimens that according
to our morphological comparisons belong to an unknown species of Pristimantis that we name and describe herein.

Material and methods

The format for the diagnosis and holotype description is based on Lynch & Duellman (1997), while the
terminology for morphological traits follows Duellman & Lehr (2009). Specimens were collected under the
Ministerio del Ambiente de Ecuador research permit 03–2015-INVESTIGACIÓN-B-DPMS/MAE. Specimens
were euthanized using 2% benzocaine. Tissue samples of the liver were extracted and placed in cryovals with 95%
ethanol. Specimens were fixed in 10% formaldehyde and preserved in 70% ethanol, following McDiarmid (1994).
All the specimens as well as tissue samples were deposited in the Herpetology collection of Museo de Zoología de
la Universidad del Azuay (MZUA). Geographic coordinates and elevation were taken with a GPS. Descriptions of
the habitat where specimens were collected as well as coloration patterns in life are based on the author’s field
notes and photographs.

Morphological data

Sex and maturity were determined by gonadal inspection via flank incisions as well as through the observation of
secondary sexual traits (nuptial pads, vocal slits and vocal sac). All morphometric variables were measured three
times using a digital caliper to the nearest 0.1 mm. We present the average and the maximal and minimal values of
each morphometric character (Table 1). Abbreviations for measurements are as follows: eye to nostril distance
(EN), head length (HL), head width (HW), interorbital distance (IOD), internarinal distance (IND), snout vent
length (SVL), tibia length (TL), foot length (FL), tympanum diameter (TD), eye diameter (ED) and upper eyelid
width (EW).
We compared morphological characteristics and coloration patterns of the new species with other similar
Pristimantis species from southeastern Ecuador and northern Peru. We used original descriptions and
morphological notes available in Duellman & Lehr (2009) for P. bellator (Lehr et al. 2007), P. caeruleonotus (Lehr
et al. 2007), P. ceuthospilus (Duellman & Wild 1993), P. flavobracatus (Lehr et al. 2006) and P. versicolor (Lynch
1979). For comparisons with P. altamazonicus (Barbour & Dunn 1921) we used recent morphological descriptions
available in Ron et al. (2016) and Ortega-Andrade et al. (2017). Moreover, we studied the external morphology of
type material (holotypes and paratypes) using photographs available from Ron et al. (2016) for: Pristimantis
croceoinguinis (Lynch 1968), P. cryptomelas (Lynch 1979), P. diadematus (Jimenez de la Espada 1875), P.
ventrimarmoratus (Boulenger 1912) and P. versicolor (Lynch 1979). Finally, we studied directly type material of
Pristimantis ardyae (Reyes-Puig et al. 2013), P. bambu (Arteaga-Navarro & Guayasamin 2011), P. churuwiai
(Brito et al. 2017) and P. gualacenio (Urgiles et al. 2014) as well as additional specimens of P. altamazonicus
(Barbour & Dunn 1921) and P. nigrogriseus (Andersson 1945) available at Museo de Zoología de la Universidad
del Azuay Cuenca-Ecuador (MZUA), Instituto Nacional de Biodiversidad del Ecuador (INABIO), and Museo de
Zoología de la Pontificia Universidad Católica del Ecuador (PUCE).
We examined the stomach contents of five specimens of the new species. Contents were examined to the level
of family where possible, although very fragmented contents could only be identified to the level of order. The
identification of food items was carried out at the entomology laboratory of Universidad del Azuay, Ecuador.

Pristimantis nimbus new species

Figs. 2–5|

Holotype. MZUA.AN.1475, an adult male, collected on 19 September 2015 by Juan C. Sánchez, Veronica L.
Urgiles and Bruno A. Timbe at the Ecological Conservation Area Tinajillas-Río Gualaceño, (3°0’0.29’’S;
78°30’34.47’’W; 2399 masl), Provincia Morona Santiago, Ecuador (Fig. 1).

A NEW TERRESTRIAL FROG FROM ECUADOR Zootaxa 4318 (3) © 2017 Magnolia Press · 521
FIGURE 1. Map of Ecuador showing the distribution of Pristimantis nimbus sp. nov. at the Tinajillas-Río Gualaceño
Ecological Conservation Area, Morona Santiago, Ecuador.

Paratopotypes. MZUA.AN. 1462, 1465, 1466, 1468 (adult males) and MZUA.AN.1472, 1483 (subadult
males), collected with the holotype.
Paratypes. MZUA.AN.0705, a sub-adult male collected on 10 October 2013 by Veronica L. Urgiles and
Cristian Nieves at the Área Ecológica de Conservación Tinajillas Rio Gualaceño (3°1’47.20’’S; 78°35’3.43’’W;
2273 masl), Provincia Morona Santiago, Ecuador.
Diagnosis. Due to a lack of morphological synapomorphies for Pristimantis (Hedges et al. 2008; Padial et al.
2014), we tentatively assign the new species to this genus based on morphological similarity to other species in this
clade. Pristimantis nimbus (Figs. 2–4; Table 1) is a small species characterized by: (1) skin of dorsum finely
shagreen with scattered low and small tubercles, flanks areolate, being more noticeable towards the ventral region,
skin on venter finely areolate, lacking dorsolateral and discoidal folds; (2) tympanic membrane and annulus

522 · Zootaxa 4318 (3) © 2017 Magnolia Press URGILES ET AL.

differentiated, visible, rounded (42% of eye diameter), supratympanic fold present, one or two postrictal tubercles;
(3) short snout, subacuminate in dorsal view, rounded in profile, lacking tubercles on the tip, canthus rostralis
slightly concave; (4) upper eyelid bearing one or two small subconical tubercles and several scattered low
tubercles, cranial crests absent; (5) dentigerous process of vomer prominent, slightly oval with three to seven teeth,
rounded choana; (6) nuptial pads present, vocal sac and vocal slits absent; (7) finger I shorter than II, discs on all
fingers laterally but moderately expanded; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles low, one
diffuse antebrachial tubercle, palmar tubercles low and small; (10) heel with one subconical tubercle, shank lacking
tubercles, tarsal tubercles low and small; (11) toes bearing narrow lateral fringes; webbing absent; toe V longer
than III; discs of toes moderately expanded; (12) inner metatarsal tubercle ovoid three times bigger than outer one,
rounded; supernumerary plantar tubercles present, rounded, smaller than subarticular tubercles; (13) iris greenish-
cream with thin dark brown flecks and horizontal reddish copper stripe, dorsal coloration varies from copper-
brown with few dark brown and green marks to brown with greenish-brown; flanks are cream with dark brownish
flecks, ventral coloration is dusty brown with dark brown flecks; groin, thighs, posterior portion of flanks, and arm
insertion with bright yellow to orange oval-shaped flash marks surrounded by dark brown to black marks forming a
reticulated pattern; (14) SVL in adult males 23.9 mm ±2.9 (range: 22.4–28.6).

FIGURE 2. Adult male holotype of Pristimantis nimbus sp. nov. (MZUA.AN.1475, SVL 24.5 mm) and detail of flash marks
on the groin, flanks, hidden surfaces of thighs and arm insertion.

Comparison with similar species. Pristimantis nimbus is similar to other species of Pristimantis from north
eastern Ecuador and northern Peru which have distinctive flash marks on the groin and lack cranial crests:
Pristimantis altamazonicus, P. ardyae, P. bambu, P. bellator, P. caeruleonotus, P. cethospilus, P. churuwiai, P.
croceoinguinis, P. cryptomelas, P. diadematus, P. flavobracatus, P. gualacenio, P. nigrogriceus, P.
ventrimarmoratus and P. versicolor. However, Pristimantis altamazonicus differs from the new species by lacking
tubercles on tarsus and heel, and by having reddish to orange inguinal coloration with bold black bars and blotches.
Pristimantis ardyae differs from P. nimbus by lacking ulnar tubercles and by the amber with dark orange spots
dorsal coloration (brownish to olive green in P. nimbus) and finely granular skin. Pristimantis bambu lacks a
tympanic membrane and tubercles on tarsus and heel. Pristimantis bellator differs from the new species by having
a smooth dorsal skin with scattered tubercles, small tubercles on the upper eyelids (one or two subconical in P.
nimbus) and basal webbing on feet. Pristimantis caeruleonotus differs from the new species by having smooth
dorsal skin with scattered spicules, dorsolateral folds, a tubercle in the tip of the snout and black inguinal coloration
with white and pale blue spots. Pristimantis ceuthospilus differs from P. nimbus by lacking tubercles on the heel
and by having a sharper snout and large orange to yellow spots on the posterior surfaces of thighs, which are not

A NEW TERRESTRIAL FROG FROM ECUADOR Zootaxa 4318 (3) © 2017 Magnolia Press · 523
outlined by black or brown. Pristimantis gualacenio differs from the new species by lacking dentigerous process
and conspicuous conical tubercles on the dorsum, eyelids, tarsus, heel and ulna, while P. cryptomelas differs from
the new species by lacking nuptial pads and by presenting conspicuous dermal ridges on the head, occipital region
and anterior part of dorsum, and by having large conical tubercles on the tarsus and heel. Pristimantis versicolor
differs from P. nimbus by lacking tubercles on the heel and by presenting a brownish to black groin coloration with
red-pinkish flecks (dark brown with bright yellow spots in P. nimbus). Pristimantis churuwiai differs from the new
species by having dorsolateral folds, an internarial tubercle and a finely granular dorsal skin. Furthermore,
Pristimantis churuwiai is only known to occur at elevations between 1400 and 1800 masl. Pristimantis diadematus
differs from the new species by lacking tubercles on ulna, tarsus and upper-eyelid and by having scapular folds and
an inguinal coloration that varies from bluish-white, yellowish, pinkish or pale-green with dark brown diagonal
bars. Besides, Pristimantis diadematus only occurs in the lowland tropical rainforest at elevations below 1700
masl. Likewise, P. ventrimarmoratus and P. nigrogriseus have been only registered at elevations below 1800 masl,
and differ from the new species in that nuptial pads and tarsal tubercles are absent in P. nigrogriseus and the
tympanic membrane and nuptial pads are absent in P. ventrimarmoratus.

TABLE 1. Measurements (mm) of adult males of Pristimantis nimbus sp. nov. The mean and the standard deviation
(SD) values are shown for each morphological character.
MZUA MZUA MZUA MZUA MZUA Mean±SD
AN1475 AN1465 AN1462 AN1466 AN1468 (range)

Holotype Paratype Paratype Paratype Paratype

SVL 24.5 25.0 28.6 22.4 23.4 23.9±2.9
(22.4–8.6)
EN 4.3 4.1 5.1 3.8 4.1 4.2±0.5
(3.9–5.1)
HL 7.1 6.6 8.2 7.3 7.0 7.0±0.8
(6.6–8.2)
HW 9.4 8.9 11.4 8.2 8.7 9.0±1.4
(8.2–11.4)
IOD 2.5 2.4 3.3 2.5 2.5 2.5±0.5
(2.5–3.3)
IND 1.3 0.9 1.4 1.0 0.8 1.1±0.3
(0.8–1.4)
TL 13.0 13.1 16.2 12.1 12.7 12.9±1.9
(12.1–16.2)
FL 10.9 10.1 13.8 9.6 10.8 10.7±1.8
(9.6–13.8)
TD 7.3 7.5 9.4 6.4 6.8 7.1±1.4
(6.4–9.4)
ED 1.2 1.1 1.2 1.1 0.1 1.1±0.1
(1.0–1.1)
EW 2.7 2.9 3.6 2.4 2.7 2.7±0.5
(2.4–3.6)

Description of the holotype. Adult male (Figs. 2–4) with SVL 24.5 mm. The head is wider than long, head
length 75% of the head width. Snout subacuminate in dorsal view and rounded in profile (Fig. 4C–D), eye-nostril
length 11.71% of SVL. Canthus rostralis slightly concave in dorsal view, straight in profile view, nostrils laterally
oriented; interorbital space flat, narrower than the upper eyelid, interorbital distance 82% of the upper eyelid.
Cranial crests absent, upper eyelid with two subconical tubercles and five to eight low and small scattered
tubercles. Tympanic membrane differentiated from the surrounding skin, tympanic annulus 42% of eye diameter.
Two small rounded postrictal tubercles present. Small and round choanas not concealed by palatal shelf of maxilla.
Dentigerous process of vomers present (each with 3–7 teeth), ovoid in shape, situated posteromedially to the
choanas, tongue as wide as long, rounded, its anterior 30% attached to floor of mouth.

524 · Zootaxa 4318 (3) © 2017 Magnolia Press URGILES ET AL.

FIGURE 3. Dorsal (A) ventral (B) and lateral (C) views of the preserved male holotype of Pristimantis nimbus sp. nov.
(MZUA.AN.1475; SVL 24.5 mm).

Dorsum finely shagreen with small scattered low tubercles and the skin of venter finely areolate. Discoidal fold
absent (Fig. 3). Arms slim with respect to the body, fingers with discs, pads defined by circumferential grooves,
narrow lateral fringes on all fingers, palmar tubercle U-shaped, thenar tubercle ovoid, subarticular tubercles
rounded and prominent, supernumerary tubercles present (Fig. 4A). Nuptial pads present, vocal slits and vocal sac
absent. Hind limbs slim, tibia length 52% of SVL, heel with a small subconical tubercle, outer-edge of tarsus with
low and small tubercles; inner tarsal fold well defined. Toes bear narrowly lateral fringes, subarticular tubercles
rounded and prominent, inner metatarsal tubercle ovoid, three times longer that the round outer metatarsal tubercle.
Supernumerary plantar tubercles present, tip of toes with discs, toe V larger than III, reaching the subarticular distal
tubercle of toe IV (Fig. 4B).
Coloration of the holotype. In life (Fig. 2), the anterior region of dorsum copper-brown with tiny dark brown
spots; the posterior region dark brown. Head color on eyelid copper-brown with minute scattered black spots. Iris
greenish-cream with thin dark brown flecks and a horizontal reddish copper interorbital bar; a white sclera is
visible in the inferior and posterior surface of the eye. Supratympanic stripe dark brown, diffuse brown cantal stripe
and supralabial chevrons. A yellowish spot at the base of the humeral region. Thighs, groin and posterior region of
the flanks with bright-yellow to orange oval spots and blotches surrounded by dark brown to black, forming a
reticulated pattern. Venter salmon pink with dark brown flecks in the thoracic region; posterior surfaces of the
thighs and groin are dark brown with bright yellowish ovoid spots. Palmar and plantar surfaces red-brown with
dark brownish supernumerary tubercles.

A NEW TERRESTRIAL FROG FROM ECUADOR Zootaxa 4318 (3) © 2017 Magnolia Press · 525
FIGURE 4. Details of the hand (A), foot (B), and dorsal and lateral views of the head (C–D) of the adult male holotype of
Pristimantis nimbus sp. nov. (MZUA.AN.1475).

In preservative (Fig. 3), dorsum light brown with tiny dark brown spots. Upper eyelid, snout and occipital
region grey with small dark flecks. Flanks cream with narrowly dark brown chevrons, the groin dark brown with
cream oval spots. Venter pinkish-cream with small black and dark brown spots. Dorsal surfaces of digits pinkish-
cream with dark spots; dorsal surface of thighs dark brown with pale cream circular marks.
Variation. Morphometrics are presented in Table 1. On one specimen (MZUA.AN.1468), tubercles on the
dorsum are more conspicuous, while in a sub-adult (MZUA.AN.0705) the venter is notably areolate. In life, all
specimens have well differentiated low tarsal tubercles, although this condition is only partially visible in
preservative. Also, one of the paratypes (MZUA.AN.1472) presents two discontinuous copper-brown dorsolateral
stripes, and another paratype (MZUA.AN.0705) presents an olive green dorsal coloration with brown marks,
brighter yellowish humeral marks and dark-brown labial chevrons (Fig. 5).
Etymology. The specific epithet “nimbus” corresponds to the Latin masculine noun for cloudy or dark cloud,
and refers to the constant cloudy conditions of the type locality.
Distribution and natural history. Pristimantis nimbus is only known from the type locality, at elevations
between 2200–2400 masl. The area is part of the Tinajillas-Rio Gualaceño Ecological Conservation Area, a region
of montane cloud forests in the southeastern Andes of Ecuador (Fig. 1). The forest is dominated by tree species of
the genera Oreopanax, Weinmannia, Cinchona and Ocotea and the canopy reaches 30 m and is characterized by a
high dominance of epiphytes (Baez et al. 2013). All individuals of Pristimantus nimbus were found at night,
perching on leaves and branches between 80 cm to 150 cm above the ground. The analysis of stomach contents
rendered a total of 17 prey items of different unidentified species in five orders (Aranae, Coleoptera, Diptera,
Hemiptera, Lepidoptera). 30.9% of prey items could not be determined at the family level but belong to the
Coleoptera, Lepidoptera and Aranae. Among the items identified at the family level, most common preys were
species in the families Tipulidae (23.5%) and Membracidae (16.6%). Other families preyed upon were
Curculionidae (11.6%), Licosidae (5.8%), Tephritidae (5.8%) and Pentatomidae (5.8%). Other species of anurans
found in syntopy were Pristimantis cf. altamnis, P. galdi (Jimenez de la Espada, 1870), P. proserpens (Lynch

526 · Zootaxa 4318 (3) © 2017 Magnolia Press URGILES ET AL.

1979), P. tinajillas (Urgiles et al. 2014), P. versicolor (Jiménez de la Espada 1870), Rhinella margaritifera
(Laurenti 1768), Noblella sp., and three unidentified species of Pristimantis.
Remarks. The exposed tympanic membrane, expanded digital discs, presence of vomerine teeth, toe V much
longer than the III and the absence of a cranial crest are all features that indicate that Pristimantis nimbus fits the
definition of the Pristimantis unistrigatus species group (sensu Hedges et al. 2008). However, several studies (e.g.,
Hoyos et al. 2014; Padial et al. 2014) revealed rampant paraphyly in this group, which was dismantled by Padial et
al. (2014). Given the lack of evidence for the monophyly of the Pristimantis unistrigatus group, as well as
morphological evidence that would unambiguously place P. nimbus to any of the available monophyletic species
groups of Pristimantis, we leave the new species unassigned to group.

FIGURE 5. Color variation in specimens of Pristimantis nimbus sp. nov. A) MZUA.AN.1465 (SVL 25.0 mm), B)
MZUA.AN.1472 (SVL 18.5 mm), C) MZUA.AN.0705 (SVL 17.7 mm).

Acknowledgments

We gratefully acknowledge the field assistance of Cristian Maldonado, Saúl Benavidez, Patricio Estrella and
Cristian Nieves. The identification of stomach contents was performed by Sebastian Padrón. The local government
of Limon Indanza provided logistic support and facilities. This article benefited from extensive editing and advice
provided by Jose Padial, Anna Savage, David Siddons, Paul Szekely, Eduardo Toral and two anonymous
reviewers. Mario Yanez Muñoz provided access to herpetology collections at INABIO and valuable comments on
this article as well. Santiago Ron provided access to the herpetology collection at PUCE. We thank Jacinto Guillen
for his continuous support of our research. This study was funded by Universidad del Azuay (Fondos UDA 2015
[24, 2015]).

References

Andersson, L.G. (1945) Batrachians from East Ecuador, collected 1937, 1938 by Wm. Clarke-Macintyre and Rolf Blomberg.
Arkiv för Zoologi, 37A, 1–88.

A NEW TERRESTRIAL FROG FROM ECUADOR Zootaxa 4318 (3) © 2017 Magnolia Press · 527
Arteaga-Navarro, A.F. & Guayasamin, J.M. (2011) A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the
high Andes of southeastern Ecuador, discovered using morphological and molecular data. Zootaxa, 2876, 17–29.
Baez, S., Santiana, J. & Guevara, J. (2013) Bosque siempreverde montano del Sur de la Cordillera Oriental de los Andes. In:
Ministerio del Ambiente del Ecuador 2012. Sistema de Clasificación de los Ecosistemas del Ecuador Continental.
Ministerio del Ambiente del Ecuador, Quito, pp. 122–123.
Barbour, T. & Dunn, E.R. (1921) Herpetological novelties. Proceedings of the Biological Society of Washington, 34, 157–162.
Batallas, D. & Brito J. (2014) Nueva especie de rana del género Pristimantis del grupo lacrimosus (Amphibia: Craugastoridae)
del Parque Nacional Sangay, Ecuador. Papeís Avulsos de Zoologia, 54, 51–62.
https://doi.org/10.1590/0031-049.2014.54.05
Boulenger, G.A. (1912) Descriptions of new batrachians from the Andes of South America, preserved in the British Museum.
Annals and Magazine of Natural History, 8, 185–191.
https://doi.org/10.1080/00222931208693215.
Brito, J., Ojala-Barbour, R., Batallas, D. & Almendariz, A. (2016) A New Species of Pristimantis (Amphibia: Strabomantidae)
from the Cloud Forest of Sangay National Park, Ecuador. Journal of Herpetology, 50, 337–344.
https://doi.org/10.1670/13-103
Brito, J., Batallas, D. & Yanez, M.H. (2017) Ranas terrestres Pristimantis (Anura: Craugastoridae) de los bosques montanos del
Río Upano, Ecuador: Lista anotada, patrones de diversidad y descripción de cuatro especies nuevas. Neotropical
Biodiversity, 3, 125–126.
https://doi.org/10.1080/23766808.2017.1299529
Bustamante, M.R. & Mendelson, J.R. (2008) A new frog species (Strabomantidae: Pristimantis) from the High Andes of
Southeastern Ecuador. Zootaxa, 1820, 49–59.
Duellman, W.E. & Wild, E.R. (1993) Anuran amphibians from the Cordillera de Huancabamba, northern Peru: Systematics,
ecology, and biogeography. Occasional Papers of the Museum of Natural History, University of Kansas, 157, 1–53.
Duellman, W.E. & Lehr, E. (2009) Terrestrial-Breeding Frogs (Strabomantidae) in Peru. Natur und—Verlag,
Naturwissenschaft, Münster, 386 pp.
Gonzalez-Voyer, A., Padial, J.M., Castroviejo-Fisher, S., de la Riva, I. & Vilá, C. (2011) Correlates of species richness in the
largest neotropical amphibian radiation. Journal of Evolutionary Biology, 24, 931–942.
https://doi.org/10.1111/j.1420-9101.2011.02243.x
Guayasamín, J.M., Almeida-Reinoso, D. & Nogales-Sornosa, F. (2004) Two new species of frogs (Leptodactylidae:
Eleutherodactylus) from the high Andes of northern Ecuador. Herpetological Monographs, 18, 127–141.
https://doi.org/ 10.1655/0733-1347(2004)018[0127:TNSOFL]2.0.CO;2
Guayasamin, J.M. & Arteaga, A.F. (2013) A new species of the Pristimantis orestes group (Amphibia: Strabomantidae) from
the high Andes of Ecuador. Zootaxa, 3616 (4), 345–356.
https://doi.org/10.11646/zootaxa.3616.4.3.
Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular
phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1–182.
Hoyos, J.M., Mantilla, C., Galindo, D. & Salgar, L. (2014) Phylogenetic analysis within the Pristimantis unistrigatus (Anura,
Craugastoridae) group based on morphological characters. Caldasia, 36, 107–124.
https://doi.org/10.15446/caldasia.v36n1.43894
Hutter, C.R. & Guayasamin, J.M. (2015) Cryptic diversity concealed in the Andean cloud forests: two new species of rainfrogs
(Pristimantis) uncovered by molecular and bioacoustic data. Neotropical Biodiversity, 1, 35–59.
https://doi.org/10.1080/23766808.2015.1100376
Jimenez de la Espada, M. (1870) Fauna neotropicalis species quaedam nondum cognitae. Jornal de Sciências, Mathemáticas,
Physicas e Naturaes, 3, 57–65.
Jimenez de la Espada, M. (1875) Vertebrados del Viaje al Pacífico Verificado de 1862 a 1865 por una Comisión de Naturalistas
Enviada por el Gobierno Español. Batracios. M. Ginesta, Madrid, 208 pp.
Laurenti, J.N. (1768) Specimen medicum, exhibens synopsin reptilium emendatam cum experimentis circa venena et antidota
reptilium austracorum, quod authoritate et consensu. Joan. Thomae, Vienna, 217 pp
Lehr, E., Lundberg, M., Aguilar, C. & von May, R. (2006) New species of Eleutherodactylus (Anura: Leptodactylidae) from the
eastern Andes of Central Peru with comments on central Peruvian Eleutherodactylus. Herpetological Monographs, 20,
05–128.
https://doi.org/10.1655/0733-1347(2007)20[105:nsoeal]2.0.co;2
Lehr, E., Aguilar, C., Siu-Ting, K. & Jordán, J.C. (2007) Three new species of Pristimantis (Anura: Leptodactylidae) from the
Cordillera de Huancabamba in northern Peru. Herpetologica, 63, 519–536.
https://doi.org/10.1655/0018-0831(2007)63[519:tnsopa]2.0.co;2
Lehr, E., Catenazzi, A. & Rodríguez, D. (2009) A new species of Pristimantis (Anura: Strabomantidae) from the Amazonian
lowlands of northern Peru (Region Loreto and San Martín). Zootaxa, 1990, 30–40.
https://doi.org/10.5281/ZENODO.185535
Lynch, J.D. (1968) Two new frogs of the genus Eleutherodactylus from eastern Ecuador (Amphibia: Leptodactylidae). Journal
of Herpetology, 2, 129–135.
https://doi.org/10.2307/1563112

528 · Zootaxa 4318 (3) © 2017 Magnolia Press URGILES ET AL.

Lynch, J.D. (1979) Leptodactylid frogs of the genus Eleutherodactylus from Andes southern of Ecuador. Miscellanuos
Publications of Kansas University Natural History Museum, 66, 1–62.
Lynch, J.D. & Duellman, W.E. (1997) Frogs of the genus Eleutherodactylus (Leptodactylidae) in western Ecuador: systematics,
ecology, and biogeography. The University of Kansas Natural History Museum, 23, 1–236.
McDiarmid, R.W. (1994) Preparing Amphibians as Scientific Specimens. In: Heyer, W.R. (Eds.), Measuring and Monitoring
Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, pp. 289–297.
Navarrete, M., Venegas, P.J. & Ron, S.R. (2016) Two new species of frogs of the genus Pristimantis from Llanganates National
Park in Ecuador with comments on the regional diversity of Ecuadorian Pristimantis (Anura, Craugastoridae). Zookeys,
593, 139–162.
https://doi.org/10.3897/zookeys.593.8063
Ortega-Andrade, H.M., Rojas-Soto, O., Espinosa de los Monteros, A., Valencia, J.H., Read, M. & Ron, S.R. (2017)
Revalidation of Pristimantis brevicrusn(Anura, Craugastoridae) with taxonomic comments on a widespread Amazonian
direct-developing frog. Herpetological Journal, 26 87–103.
Padial, J.M., Grant, T. & Frost, D.R. (2014) Molecular systematics of terraranas (Anura: Brachycephaloidea) with an
assessment of the effects of alignment and optimality criteria. Zootaxa, 3825 (1), 1–32.
https://doi.org/10.11646/zootaxa.3825.1.1
Reyes-Puig, M.M., Reyes-Puig, J.P. & Yanez-Muñoz, M.H. (2013) Ranas terrestres del género Pristimantisb (Anura:
Craugastoridae) de la Reserva Ecológica Río Zúñag, Tungurahua, Ecuador: Lista anotada y descripción de una especie
nueva. Avances en Ciencias e Ingenierias, 5, B5–B13.
https://doi.org/10.18272/aci.v5i2.133
Ron, S.R., Guayasamin, J.M., Yanez-Muñoz, M.H., Merino-Viteri, A., Ortiz, D.A. & Nicolalde, D.A. (2016)
AmphibiaWebEcuador. Version 2016.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Available from:
http://zoologia.puce.edu.ec/Vertebrados/anfibios (accessed 4 July 2017)
Urgiles, V.L., Sánchez-Nivicela, J.C., Nieves, C. & Yanez-Muñoz, M.H. (2014) Ranas terrestres en los ecosistemas surandinos
de Ecuador I: Dos nuevas especies de Pristimantis (Anura: Craugastoridae) de la ladera oriental. Avances en Ciencias e
Ingenierias, 6, B51–B59.
https://doi.org/10.18272/aci.v6i1.159
Yanez-Muñoz, M.H., Bejarano-Muñoz, P., Brito, J. & Batallas, D. (2014) Ranas terrestres de los Andes Surorientales de
Ecuador II: Una nueva especie de Pristimantis verde espinosa de los bosques montanos del Parque Nacional Sangay
(Anura: Craugastoridae). Avances en Ciencias e Ingenierias, 6, B63–B77.
Wells, K. (2007) The ecology and behaviour of amphibians. The University of Chicago Press, Chicago, 1085 pp.
https://doi.org/10.7208/chicago/9780226893334.001.0001

APPENDIX 1. Specimens used for comparisons.

Pristimantis altamazonicus: ECUADOR-MORONA SANTIAGO: Central Hidroeléctrica Sopladora (2°36’15.58’’S;

78°26’2.84’’W; 949 masl) MZUA.AN.0069, MZUA.AN.0070. ECUADOR-MORONA SANTIAGO: La Virgen
(2°58’28.96’’S; 77°57’21.96’’W; 618 masl) QCAZ54419; QCAZ54438; Cerro Shaime (2°57’47.49’’S; 77°47’22.52’’W;
325 masl) QCAZ54507.
Pristimantis ardyae: ECUADOR-PASTAZA: Reserva Comunitaria Ankaku (1°16’34.11’’S; 77°54’04.59’’W; 768 masl)
QCAZ45956. ECUADOR-TUNGURAHUA: Reserva Ecológica Rio Zuñag (01° 22’31.3” S, 78°09’43.5” W, 2200 m)
QCAZ52498.
Pristimantis bambu: ECUADOR-CAÑAR: Rivera, Reserva Mazar, Parque Nacional Sangay (2°32’51.33’’S; 78°41’50.64’’W;
2934 masl) QCAZ46726, QCAZ46739.
Pristimantis cryptomelas: ECUADOR-LOJA: San Sebastian, Cajanuma (4°05’02.99’’S; 79°12’06.61’’W; 2454 masl)
QCAZ45581, QCAZ45592; Bosque Protector Huashapamba (3°39’39.06’’S; 79°16’30.70’’W; 2921 masl) QCAZ63279,
QCAZ63280, QCAZ63278. ECUADOR-ZAMORA CHINCHIPE: 5km E Loja (3°57’40.70’’S; 79°07’10.22’’W; 2541
masl) KU141992 (specimen revised via photographs), Parque Nacional Yacuri (4°45’43.29’’S; 79°24’50.46’’W; 3194
masl) QCAZ61181, QCAZ61191, QCAZ61187, QCAZ 61193, QCAZ61180; Reserva Tapichalaca (4°28’22.71’’S;
79°07’47.83’’W; 2044 masl) QCAZ45612, QCAZ45616-45620, QCAZ37146; QCAZ57148.
Pristimantis diadematus: ECUADOR-MORONA SANTIAGO: Central Hidroeléctrica Cardenillo (2°35’40.06’’S;
78°25’37.87’’W; 1270 masl), MZUA.AN.0254; San Carlos (2°36’15.58’’S; 78°26’2.84’’W; 1000 masl),
MZUA.AN.1218; MZUA.AN.1281; MZUA.AN.1282; MZUA.AN.1283. ECUADOR-NAPO: Río Aguarico
(0°11’50.04’’S; 77°28’58.74’’W; 1218 masl), KU123381 (specimen revised via photographs).
Pristimantis churuwiai: ECUADOR-MORONA SANTIAGO: Sardinayacu, Parque Nacional Sangay (2º04′45.3″S;
78º09′37.3″W, 1400 masl) DHMECN12242.
Pristimantis croceoinguinis: ECUADOR-SUCUMBIOS: Santa Cecilia (0°03’06.07’’S; 76°58’12.98’’W; 325 masl),
KU110789 (specimen revised via photographs).
Pristimantis nigrogriseus: ECUADOR-NAPO: 3-4 km carretera a Cuyuja-Papallacta (0°23’27.26’’S; 78°03’21.72’’W; 2770
masl) QCAZ01124; 10km Oyacachi (0°13’55.61’’S; 78°00’25.98’’W; 2788 masl) QCAZ07553, QCAZ_07655

A NEW TERRESTRIAL FROG FROM ECUADOR Zootaxa 4318 (3) © 2017 Magnolia Press · 529
 QCAZ_58902. ECUADOR-TUNGURAHUA: Rio Verde (1°24’24.56’’S; 78°17’27.97’’W; 1629 masl) QCAZ30808,
QCAZ30809, QCAZ30810, QCAZ30817. ECUADOR-MORONA SANTIAGO: Nueve de Octubre (2°13’08.04’’S;
78°00’26.57’’W; 1095 masl) QCAZ32272. ECUADOR-ZAMORA CHINCHIPE: Parque Nacional Podocarpus
(4°23’54.85’’S; 79°08’54.92’’W; 2166 masl) QCAZ_23816, QCAZ26951.
Pristimantis ventrimarmoratus: ECUADOR-MORONA SANTIAGO: Central Hidroeléctrica Sopladora, (2°35’11.65’’S;
78°27’57.13’’W; 1354 masl), MZUA.AN.1305, MZUA.AN.1310, MZUA.AN.1312, MZUA.AN.1316. ECUADOR-
TUNGURAHUA: El Topo, Río Pastaza, (1280 masl) BMNH 1947.2.15.74 (specimen revised via photographs).
Pristimantis versicolor: ECUADOR-LOJA: 15 km E from Loja city (3°57’35.98’’S; 79°06’35.61’’W; 2695 masl) KU119866
(specimen revised via photographs); Parque Nacional Podocarpus (4°23’54.85’’S; 79°08’54.92’’W; 2166 masl),
QCAZ45572, QCAZ45574, QCAZ63785, QCAZ63786; QCAZ62024, QCAZ62024. ECUADOR-ZAMORA
CHINCHIPE: Parque Nacional Colambo-Yacuri (4°45’43.29’’S; 79°24’50.46’’W; 3194 masl) QCAZ61567, QCAZ61570.

530 · Zootaxa 4318 (3) © 2017 Magnolia Press URGILES ET AL.

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