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A new Andean Toad of the genus Osornophryne (Amphibia: Anura: Bufonidae) from northwestern Ecuador, with taxonomic remarks on the genus

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We describe a new species of Andean toad of the genus Osornophryne from montane cloud forests of northwestern Ecuador between 2500–2750 m above sea level. The new species is characterized by moderately large size (snout-vent length 32.5–36.0 mm in adult females, 21.1–23.2 mm in adult males), dorsal skin highly rugose with scattered irregular warts, dorsolateral ridges of round or oblong glandular warts, venter with clusters of flattened pustular warts, angular rostrum shaped like a four-sided pyramid protruding in dorsal and lateral views but not forming a proboscis, smooth borders of transverse processes of pre-sacral vertebrae, a dark brown dorsum in life with ochre-brown warts, dorsolateral ridges maroon-brown, and abundant pale salmon spots on the venter. We suggest that it be clas-sified as Endangered under IUCN criteria, because of its small distributional, narrow al-titudinal range subjected to habitat loss and fragmentation. We discuss different aspects of taxonomy, natural history, and distribution of the different species of Osornophryne. In particular, we comment on the importance of morphological, chromatic, and molecular analyses that take into consideration ontogenic and dimorphic variation among species of Osornophryne. Abstract Describimos una nueva especie de sapo Andino del género Osornophryne de los bosques montano nublados del noroccidente de Ecuador entre 2500–2700 m sobre el nivel del mar. La nueva especie se caracteriza por su tamaño moderadamente largo (longitud hocico-cloaca 32.5–36.0 mm en hembras adultas, 21.1–23.2 mm en machos adultos), piel del dorso muy rugosa con verrugas irregularmente distribuidas, pliegues dorsolaterales compuestos de verrugas glandulares redondas u oblongas, vientre con agrupaciones de verrugas pustu-lares aplanadas, hocico angular con forma de una pirámide tetragonal proyectado en vistas dorsal y lateral pero sin formar una probóscide; bordes del proceso transverso lisos, dorso de color café oscuro en vida con pliegues dorsolaterales y verrugas café marrón y abun-dantes puntos salmón pálido en el vientre. Esta nueva especie es el único Osornophryne conocido que vive en las laderas occidentales de los Andes y recomendamos que sea clasifi-cado como En Peligro bajo los criterios de la UICN debido a su reducido rango de distribu-ción que está bajo fuerte presión debido a la pérdida de hábitat y fragmentación. También discutimos sobre diferentes aspectos de la taxonomía, historia natural y distribución de las diferentes especies de Osornophryne. En particular, comentamos sobre la importancia de detallados análisis morfológicos, cromáticos y moleculares que tomen en consideración la variación ontogénica y dimórfica para clarificar el complejo estado de la identidad de algunas especies de Osornophryne.
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ARTÍCULO/ARTICLE SECCIÓN/SECTION B
EN CIENCIAS E INGENIERÍAS
AVANCES
A new Andean Toad of the genus Osornophryne (Amphibia: Anura: Bufonidae) from
northwestern Ecuador, with taxonomic remarks on the genus
Diego F. Cisneros-Heredia1,2,3,, Andrew G. Gluesenkamp4,5
1Universidad San Francisco de Quito, Colegio de Ciencias Biológicas & Ambientales.
Diego de Robles y Vía Interoceánica, Quito, Ecuador.
2King’s College London, Department of Geography, London, UK.
3Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador.
4Texas Memorial Museum, University of Texas, Austin, Texas 78712, USA.
5700 Billie Brooks Drive, Driftwood, Texas, USA.
Autor principal/Corresponding author, e-mail: dcisneros@usfq.edu.ec
Editado por/Edited by: C. Zambrano, Ph.D.
Recibido/Received: 07/01/2010. Aceptado/Accepted: 09/01/2010.
Publicado en línea/Published on Web: 12/08/2010. Impreso/Printed: 12/08/2010.
Abstract
We describe a new species of Andean toad of the genus Osornophryne from montane cloud
forests of northwestern Ecuador between 2500–2750 m above sea level. The new species is
characterized by moderately large size (snout-vent length 32.5–36.0 mm in adult females,
21.1–23.2 mm in adult males), dorsal skin highly rugose with scattered irregular warts,
dorsolateral ridges of round or oblong glandular warts, venter with clusters of flattened
pustular warts, angular rostrum shaped like a four-sided pyramid protruding in dorsal and
lateral views but not forming a proboscis, smooth borders of transverse processes of pre-
sacral vertebrae, a dark brown dorsum in life with ochre-brown warts, dorsolateral ridges
maroon-brown, and abundant pale salmon spots on the venter. We suggest that it be clas-
sified as Endangered under IUCN criteria, because of its small distributional, narrow al-
titudinal range subjected to habitat loss and fragmentation. We discuss different aspects
of taxonomy, natural history, and distribution of the different species of Osornophryne.
In particular, we comment on the importance of morphological, chromatic, and molecular
analyses that take into consideration ontogenic and dimorphic variation among species of
Osornophryne.
Keywords. Anura; Conservation; Distribution; Ecuador; Osornophryne antisana;Os-
ornophryne bufoniformis;Osornophryne guacamayo;Osornophryne occidentalis new
species; Osornophryne sumacoensis;Osornophryne talipes; Taxonomy; Variation.
Abstract
Describimos una nueva especie de sapo Andino del género Osornophryne de los bosques
montano nublados del noroccidente de Ecuador entre 2500–2700 m sobre el nivel del mar.
La nueva especie se caracteriza por su tamaño moderadamente largo (longitud hocico-
cloaca 32.5–36.0 mm en hembras adultas, 21.1–23.2 mm en machos adultos), piel del dorso
muy rugosa con verrugas irregularmente distribuidas, pliegues dorsolaterales compuestos
de verrugas glandulares redondas u oblongas, vientre con agrupaciones de verrugas pustu-
lares aplanadas, hocico angular con forma de una pirámide tetragonal proyectado en vistas
dorsal y lateral pero sin formar una probóscide; bordes del proceso transverso lisos, dorso
de color café oscuro en vida con pliegues dorsolaterales y verrugas café marrón y abun-
dantes puntos salmón pálido en el vientre. Esta nueva especie es el único Osornophryne
conocido que vive en las laderas occidentales de los Andes y recomendamos que sea clasifi-
cado como En Peligro bajo los criteriosde la UICN debido a su reducido rango de distribu-
ción que está bajo fuerte presión debido a la pérdida de hábitat y fragmentación. También
discutimos sobre diferentes aspectos de la taxonomía, historia natural y distribución de las
diferentes especies de Osornophryne. En particular, comentamos sobre la importancia de
detallados análisis morfológicos, cromáticos y moleculares que tomen en consideración
la variación ontogénica y dimórfica para clarificar el complejo estado de la identidad de
algunas especies de Osornophryne.
Palabras Clave. Anura; Conservación; Distribución; Ecuador; Osornophryne antisana;
Osornophryne bufoniformis;Osornophryne guacamayo;Osornophryne occidentalis nueva
especie; Osornophryne sumacoensis;Osornophryne talipes; Taxonomía; Variación.
http://www.usfq.edu.ec/avances/aticulos/B64-2-3-2010
Avances, 2010, Vol. 2, No. 3, Pags. B64-B73
Cisneros-Heredia y Gluesenkamp Avances, 2010, Vol. 2, No. 3, Pags. B64-B73
Introduction
Toads of the bufonid genus Osornophryne Ruiz-Carran-
za and Hernández-Camacho are moderately diversified
along the Andes from central Colombia to central Ecua-
dor. They inhabit ecosystems with high humidity and
cool temperatures between 2100 and 3700 m above sea
level, i.e., montane cloud forest, upper montane forest,
timberline areas, and páramos. Eight species of Os-
ornophryne have been described and seven of them oc-
cur in Ecuador: Osornophryne antisana Hoogmoed, O.
bufoniformis(Peracca), O. cofanorumMueses-Cisneros,
Yánez-Muñoz & Guayasamin,O. guacamayo Hoogmo-
ed, O. puruanta Gluesenkamp & Guayasamin, O. suma-
coensis Gluesenkamp, and O. talipes Cannatella [1, 2,
3, 4, 5, 6, 7]. While most Ecuadorian Osornophryne
have small ranges and some are known only from their
type localities, O. bufoniformis has been reported from
a large area spanning the departments of Cauca, Huila,
Nariño, and Putumayo in Colombia and the provinces
of Carchi, Imbabura, Pichincha, Sucumbíos, and Tun-
gurahua in Ecuador [1, 2, 3, 4, 5, 6, 8, 9, 10]. Pop-
ulations assigned to O. bufoniformis in Colombia and
Ecuador show extensive phenotypic variation, and, al-
though some researchers consider it as a single highly
variable taxon [10] other herpetologists have suggested
that it is a species complex (T. Grant and D. F. Cisneros-
Heredia in [11]). Most Ecuadorian populations referred
to O. bufoniformis occur in the eastern Andean high-
lands and slopes over 2700 m, but a widely disjunct
population on the northwestern Andean slopes below
2750 m corresponds to an undescribed species of Os-
ornophryne, which we describe herein.
Methodology
Examined specimens were preserved in 70% ethanol af-
ter fixation in buffered 10% formalin. The following
measurements (in mm) were taken with digital calipers:
snout–vent length (SVL), greatest width of head (at ric-
tus; =HW), postocular head width (measured just be-
hind the eyes; =HWPO), head length (straight line dis-
tance from posterior margin of jaw to tip of rostrum;
=HL), interorbital distance (straight-line distance between
anterior margins of the orbits; =IOD); internarial dis-
tance (IND); eye-nostril distance (from anterior margin
of orbit to posterior margin of nostril; EN), horizon-
tal eye diameter (ED), greatest width of upper eyelid
(perpendicular to medial axis of skull, =UE), nostril-
rostrum distance (from anterior margin of nostril to tip
of rostrum; NR); hand length from base of thenar tuber-
cle to tip of third finger (HL3); hand length from base
of thenar tubercle to tip of fourth finger (HL4); tibia
length (TL), foot length (from base of inner metatarsal
tubercle to tip of fourth toe; FL4), foot length (from
base of inner metatarsal tubercle to tip of fifth toe; =
FL5). Sex of specimens was determined by noting the
presence of secondary sexual characters (nuptial pads)
and by direct observation of the gonads. Color pat-
Figure 1: Holotype of Osornophryne occidentalis (DHMECN
3520) in dorsal (A) and ventral (B) views, adult female, SVL =
36.0 mm.
tern in life was taken from field notes and color pho-
tographs. For the diagnosis, we followed the numbered
sequence proposed by Yánez-Muñoz et al. [12] (1) head
width/head length ratio; (2) snout form in dorsal and
lateral view, including the presence of papillae and pro-
boscis, and the form of the canthus rostralis and loreal
area; (3) dorsal skin texture; (4) ventral skin texture;
(5) flanks texture; (6) description of dorsal glandular
folds; (7) description of hind legs; (8) form and char-
acteristics of hands and feet; (9) dorsal and ventral col-
oration; (10) SVL range for adult females and males.
Classification of vegetation formations in Ecuador fol-
lows Sierra’s system [13].
Results
Osornophryne occidentalis sp. nov.
Atelopus bufoniformis Peracca, 1904 [1] in part; Peters
(1973:18–20) [2].
Osornophryne bufoniformis (Peracca, 1904) [1] in part;
Ruiz-Carranza and Hernández-Camacho (1976:
124–125) [3]; Cannatella (1986:618–622) [4]; Gluesen-
kamp (1995: 268–279) [6]; Quiguango (1997:195–209)
[14]; Toral et al. (2002:740) [15].
Holotype: DHMECN 3520, adult female (Fig. 1–3),
collected on February 2000, near Guarumos, approx-
imately 200 m from the Guarumos guard post on the
Nono–Tandayapa road, 11 km WNW of the town of
Nono (0002S, 7839W, 2550–2600 m), provincia de
Pichincha, República del Ecuador, by D.F. Cisneros-
Heredia and A. Cárdenas.
Paratypes: (4 specimens) ECUADOR: Provincia de Pich-
incha: adult male (DFCH-USFQ GU036) collected with
the holotype. Provincia de Imbabura: Two adult fe-
males (QCAZ 9318 and 10141) collected on the farm
of F. Rodríguez (0030N,7832W, 2500 m and 2750
m, respectively), W of El Rosario, Cordillera de Toisán,
by A. Quiguango et al. in September, 1994. An adult fe-
male (QCAZ 36894; Fig. 4) collected on the farm of F.
Rodríguez at 2580 m, by A. Gluesenkamp and J. Price
on 18 May 1996.
Avances, 2010, Vol. 2, No. 3, Pags. B64-B73 Cisneros-Heredia y Gluesenkamp
Figure 2: Head of the holotype of Osornophryne occidentalis
(DHMECN 3520) in dorsal (A) and lateral (B) views.
Referredmaterial: (10 juvenile or poorly-preservedspec-
imens) ECUADOR: Provincia de Imbabura: Three males
(QCAZ 9319–9321), collected on the farm of F. Ro-
dríguez at 2500 m by A. Quiguango et al. in September,
1994. A juvenile female (QCAZ 36895) collected on
the farm of F. Rodríguez at 2580 m on 18 May 1996
by A. Gluesenkamp and J. Price. A juvenile female
(KU 132126), collected near La Delicia (0022’N, 78
25’W,
2710 m), Cordillera de Intag [= Cordillera de Toisán].
Provincia de Pichincha: Two juvenile males (FHGO
1907 and 2981) and one female (FHGO 2984), collected
near Guantopungo, Parroquia de Yunguilla, buffer-zone
of the Maquipucuna Natural Reserve (0003’N,
7834’W, 2600 m), by K. Taylor on 05 January 1998
and 17 February 1999. A juvenile female (DFCH-USFQ
GU034) and a juvenile male (DFCH-USFQ GU035) col-
lected at the type locality in August 2000.
Diagnosis
(1) head wider than long (Table 1, Fig. 2); (2) snout
angular in dorsal and lateral views, shaped like four-
sided pyramid; tip of snout protracted into broad, con-
ical protuberance, projecting beyond upper jaw but not
forming proboscis; canthus rostralis distinct with ele-
vated glandular canthal ridge, straight in dorsal view,
Figure 3: Hand (A) and foot (B) of the holotype Osornophryne
occidentalis DHMECN 3520).
delimiting fairly rectangular, shallow, rugose, tubercu-
lar interorbital area (Fig. 2); (3) dorsal skin highly ru-
gose with scattered irregular warts, more dense poste-
riorly (Figs. 1, 2, 4); (4) ventral skin areolate with
clusters of flattened pustular warts; (5) prominent warts
on flanks; (6) dorsolateral ridges of round or oblong
glandular warts; (7) hind limbs short and stout, when
hindlimbs are flexed at right angles to sagittal plane,
heels do not touch, and remain widely separated; (8)
fingers and toes connected by thick web; fingers III and
IV clearly distinct, but just rounded tips of fingers I and
II protrude; toes I and II reduced, only rounded tips pro-
trude; Toe III slightly more distinct; Toe IV clearly dis-
tinct, well developed, long; toe V reduced but notice-
able, rounded tip protrudes at lateral side of Toe IV; (9)
dorsal surfaces dark brown in life with warts and dorso-
lateral ridges ochre-brown, venter maroon-brown with
abundant pale salmon spots; (10) moderately large body
size (32.5–36.0 mm SVL in adult females, n= 5; 21.1–
23.2 mm SVL in adult males, n= 5).
Comparisons
Osornophryne occidentalis is the only species of the
genus found in montane cloud forests of western Ecua-
dor. It differs from congeners by the following charac-
ters (characters of O. occidentalis in parentheses): O.
antisana has dorsum smooth to shagreen with warts and
paravertebral ridges and skin in occipital area relatively
smooth (dorsal skin rugose with warts and dorsolateral
ridges but without paravertebral ridges, skin of occipi-
tal area rugose), tip of snout protracted into fleshy pro-
boscis (tip of snout protracted into broad, conical pro-
tuberance, projecting beyond upper jaw but not form-
ing proboscis), and fourth finger and fifth toe shorter
and less pointed. Osornophryne bufoniformis has snout
fairly rounded, broader, less pointed in lateral view, with
tip barely pointed (snout angular in lateral view, shaped
like four-sided pyramid, with tip of snout protracted into
broad, conicalprotuberance, projectedbeyond upperjaw
but not forming proboscis), canthus rostralis more round-
ed, less elevated, and curved (canthus rostralis distinct,
Cisneros-Heredia y Gluesenkamp Avances, 2010, Vol. 2, No. 3, Pags. B64-B73
Females (n = 5) Males (n = 5)
Range (Mean ±SD); in mm Range (Mean ±SD); in mm
SVL 32.5-36.0 (34.5 ±1.5) 21.1-23.2 (22.3 ±0.8)
HW/HL 1.14-1.23 (1.19 ±0.05) 1.07-1.12 (1.09 ±0.02)
HW/SVL 0.32-0.37 (0.034 ±0.02) 0.35-0.37 (0.33 ±0.01)
HL/SVL 0.26-0.32 (0.29 ±0.03) 0.33-0.34 (1.09 ±0.02)
IOD/HW 0.27-0.38 (0.32 ±0.04) 0.34-0.38 (0.36 ±0.02)
IOD/HL 0.33-0.44 (0.38 ±0.04) 0.37-0.41 (0.39 ±0.01)
IOD/UE 1.00-1.46 (1.33 ±0.18) 1.26-1.42 (1.33 ±0.07)
IND/IOD 0.65-1.10 (0.78 ±0.18) 0.68-0.78 (0.73 ±0.04)
IND/NR 1.38-1.87 (1.62 ±0.20) 1.43-1.62 (1.53 ±0.09)
EN/HL 0.16-0.22 (0.19 ±0.02) 0.18-0.21 (0.19 ±0.01)
NR/HL 0.15-0.24 (0.18 ±0.04) 0.18-0.19 (0.18 ±0.01)
ED/HL 0.23-0.31 (0.27 ±0.03) 0.22-0.26 (0.24 ±0.01)
ED/IOD 0.57-0.81 (0.71 ±0.10) 0.55-0.67 (0.61 ±0.05)
TL/FL4 0.84-1.04 (0.94 ±0.09) 0.89-1.15 (0.98 ±0.11)
TL/HL 0.91-1.26 (1.03 ±0.14) 0.87-0.97 (0.91 ±0.04)
TL/SVL 0.29-0.32 (0.30 ±0.01) 0.28-0.32 (0.30 ±0.01)
Tabla 1: Variation of measurements (in mm) of adult Osornophryne occidentalis. See text for abbreviations.
Figure 4: Dorsal (A) and ventral (B) views of one paratype of
Osornophryne occidentalis (QCAZ 36894) in life, adult female,
SVL = 35.6 mm.
with elevated glandular canthal ridge, straight in dorsal
view), interorbital area poorly defined by low canthus
rostralis, fairly smooth (interorbital area fairly rectangu-
lar, rugose, tubercular), dorsum sepia or yellow-brown
to dark grayish-brown with or without warts and ridges
light brown to yellow-brown (dark brown with warts
and ridges ochre-brown), venter yellow-brown with yel-
low pustular warts to venter pinkish-red with tan pus-
tular warts, sometimes with dark brown blotches (ven-
ter maroon-brown with irregular clusters of pale salmon
pustules), fourth finger and fifth toe shorter less dis-
tinct and less pointed, see Hoogmoed (1987: Fig. 9).
Osornophryne guacamayo has longer limbs, thus when
hindlimbs are flexed at right angles to sagittal plane,
heels touch or remain close (heels do not touch and
remain widely separated), head with prominent bony
ridges in posterior part (head covered by warts), dor-
sal skin tubercular (highly rugose with abundant warts
and dorsolateral ridges), Finger III and Toe V elongated,
very distinct (Finger III distinct but not elongate, Toe
V reduced). Osornophryne percrassa has snout trun-
cate with low tip not projecting beyond anterior mar-
gin of jaw (snout angular in dorsal and lateral views,
shaped like four-sided pyramid, with tip of snout pro-
tracted into broad, conical protuberance, projected be-
yond upper jaw but not forming proboscis), dorsal skin
granular with warts but without dorsolateral ridges (dor-
sum rugose with warts and dorsolateral ridges), ven-
ter brown with bold white or yellow stripes or blotches
(venter maroon-brown with irregular clusters of pale
salmon pustules), and fourth finger and fifth toe shorter
less distinct, and less pointed. Osornophryne suma-
coensis has head rounded in dorsal view (angular in dor-
sal view), venter blue with black spots in adult females
(maroon-brown with clusters of small pale salmon pus-
tules), and transverse process of presacral vertebrae with
irregular borders (borders smooth). Osornophryne tal-
ipes has dorsal skin smooth to shagreen with warts and
paravertebral ridges (rugose with warts and dorsolat-
eral ridges but without paravertebral ridges), head that
is about as wide as long (head wider than long), tip
of snout protracted into highly acuminate fleshy pro-
boscis (tip of snout protracted into broad, conical pro-
tuberance, projected beyond upper jaw but not forming
proboscis), and Finger IV and Toe V shorter and less
pointed. Osornophryne puruanta, known only from fe-
males from the Puruanta and San Marcos lagoons in
eastern Andean Ecuador, differs in having larger females
Avances, 2010, Vol. 2, No. 3, Pags. B64-B73 Cisneros-Heredia y Gluesenkamp
(SVL 40.5–47.1 mm vs. 32.5-36.0 mm), dorsum rel-
atively smooth with numerous light-colored glandular
ridges (rugose with scattered irregular warts), andtip of
snout protracted into small proboscis in adult females
(fleshy and angular rostrum shaped like four-sided pyra-
mid). A new species from the Páramo del Ángel in
northern Andean Ecuador [12], differs from O. occiden-
talis by having dorsal skin granular with scattered large
pustules and tubercles and males with proboscis on tip
of snout.
Description of holotype
Adult female, SVL = 36.0 mm (Fig. 1), head slightly
wider than long, not as wide as adjacent part of body
(HW/HL = 1.15, HW/SVL = 0.37, HL/SVL = 0.32),
postocular width of head distinctly less than width at
corners of jaws (POHW/HW = 0.81), snout distinctly
angular in dorsal and lateral view, shaped like four-sided
pyramid (Fig. 2), tip of snout protracted into broad, con-
ical protuberance,projecting beyondupper jaw. (NR/EN
= 1.33, NR/HL = 0.21, EN/HL = 0.16, ED/HL = 0.25,
IOD/UE = 1.46). Nostrils closer to eye than to tip of
snout, posterior to margin of upper jaw, just below can-
thus rostralis in bulbous, slightly elevated, area; nos-
trils elongated, directed laterally and dorsally; canthus
rostralis distinct with elevated glandular canthal ridge,
straight in dorsal view, converging anteroventrally to
point above nares and sharply and curved anteroven-
trally in lateral view, forming raised point that extends
well past upper jaw in all views, delimiting fairly rect-
angular, shallow, rugose, tubercular platform in interor-
bital area. Occipital area rather flat, more rugose and
warty than interorbital area; upper eyelid rugose cov-
ered by small warts; loreal region with very low warts,
sloping steeply to flared upper lips; area between nos-
trils and base of protuberant tip distinctly concave; pupils
rounded; temporal region with numerous, heterogeneous
warts; tympanum absent (no vestige found after dissec-
tion); parotoid area with glands rounded, slightly tri-
angular covered by warts, externally bordered by low,
dorsolateral glandular ridges, sometimes discontinuous
as lines of warts, from posteromedial margin of orbits,
convergent but not in contact in scapular region.
Dorsal skin highly rugose with abundant warts round
and oblong; paravertebral ridges absent; dorsolateral
glandular ridges present; temporal area with warts low,
fairly uniform, and small; flanks with heterogenous, usu-
ally larger, warts on rugose skin; throat and chest dis-
tinctly areolate, almost warty especially to the sides;
venter distinctly areolate with disperse low formed by
clusters of flattened, pustular warts.
Limbs short, stout, covered with conical, rounded, and
flat warts; when hindlimbs are flexed at right angles to
sagittal plane, heels do not touch, and remain widely
separated (TL/HW = 0.80, TL/HL = 0.91, TL/FL4 =
0.84, TL/SVL = 0.29); palmar and plantar surfaces tu-
berculate; cloacal opening directed ventrally at end of
short cloacal tube.
Fingers with thick webbing, forming platform with tu-
bercular ventral surface; fingers III and IV clearly dis-
tinct (HaL4/HaL3 = 0.91), but just rounded tips of fin-
gers I and II protrude (Fig. 3); all fingers depressed with
rounded tips; dorsal surfaces of hands covered by warts,
wrist without transverse grooves; length of fingers III >
IV > II > I; toes connected by thick webbing, forming
platform with tubercular ventral surface; toes I and II re-
duced, only rounded tips protrude; Toe III slightly more
distinct; Toe IV clearly distinct, well developed, long;
Toe V reduced but noticeable, rounded tip protrudes at
lateral side of Toe IV (FL5/FL4 = 0.78; Fig. 3). Pha-
langeal formula of hand 2-2-3-3, phalangeal formula of
foot 2-2-3-4-3, tips of terminal phalanges slightly ex-
panded. Anterior and posterior margins of transverse
processes of presacral vertebrae smooth.
Coloration in life
Dorsum and dorsal surfaces of limbs dark brown, in-
cluding most dorsal warts; some warts, canthal ridge,
tip of snout, and dorsolateral ridges dark ochre-brown;
venter maroon-brown, with abundant pale salmon spots,
each composed of several pustular warts; ventral sur-
faces of limbs maroon-brown with small clusters of white
and pale salmon pustular warts; throat with fewer, small-
er pale salmon warts; palmar and plantar surfaces brown
with purplish-maroon tint and scattered pale pink tuber-
cles; iris black, heavily flecked with gold; palpebrum
translucent brown.
Coloration in preservative
Dorsum and dorsal surfaces of limbs dark brown; most
warts, canthal ridge, tip of snout, and dorsolateral ridges
ochre-brown; dorsal surfaces of hands and feet ochre-
brown with dark irregular spots; venter and ventral sur-
faces of limbs dark brown background with abundant
ochre-brown pustular warts; throat and chest with smaller
ochre-brown warts; palmar and plantar surfaces tan
brown with darker shadows; iris dark, palpebrum translu-
cent brown (Fig. 1).
Measurements
Holotype data is followed by measurements of adult fe-
males (F, n= 5) and adult males (M, n= 5) in parenthe-
sis (range, mean ±SD): SVL 36.0 mm; greatest head
width 13.3 mm (F: 10.9–13.3, 11.9 ±1.0; M: 7.8–8.5,
8.1 ±0.3); postocular head width 10.7 mm; head length
11.6 mm (F: 9.0–11.6, 10.0 ±1.2; M: 7.0–7.8, 7.5 ±
0.3); interorbital distance 5.1 mm (F: 3.0–5.1, 3.8 ±0.8;
M: 2.7–3.1, 2.9 ±0.1); internarial distance 3.3 mm (F:
2.5–3.3, 2.9 ±0.1; M: 2.0–2.2, 2.1 ±0.1); eye-nostril
distance 1.8 mm (F: 1.7–2.0, 1.9 ±0.1; M: 1.3–1.6, 1.4
±0.1); horizontal eye diameter 2.9 mm (F: 2.1–2.9, 2.7
±0.3; M: 1.7–1.9, 1.8 ±0.1); greatest width of upper
eyelid 3.5 mm (F: 2.4–3.5, 2.9 ±0.4; M: 1.9–2.4, 2.2
±0.2); nostril-rostrum distance 2.4 mm (F: 1.4–2.4, 1.8
±0.4; M: 1.3–1.5, 1.4 ±0.1); hand length to third fin-
ger 8.5 mm; hand length to fourth finger 7.7 mm; tibia
length 10.6 mm (F: 9.4–11.3, 10.3 ±0.8; M: 6.6–7.0,
Cisneros-Heredia y Gluesenkamp Avances, 2010, Vol. 2, No. 3, Pags. B64-B73
6.8 ±0.2); foot length to fourth finger 12.6 mm (F: 9.2–
12.6, 11.0 ±1.6; M: 5.9–7.9, 7.0 ±0.8); foot length to
fifth finger 9.8 mm.
Variation
Detailed information of body proportions is presented
in Table 1. Osornophryne occidentalis exhibits sexual
size dimorphism. Adult males are smaller than adult
females, and show smaller head width/head length and
head length/SVL ratios; these ratios do not overlap, how-
ever,our sample sizes are small and we will postpone a
more detailed study of sexual dimorphism (Tables 1).
Males have dorsal warts more densely packed posteri-
orly, ventral pustular warts bearing multiple coni, and
protuberance on the tip of the snout slightly broader
but never forms a proboscis. Gross morphological fea-
tures are relatively invariant. Dorsal skin is highly ru-
gose in all specimens, regardless of age or sex. Lips are
flared or slightly flared in adults, but not flared in juve-
niles. Dorsal coloration in all specimens is similar to
holotype, but ventral background coloration varies from
maroon-brown to black and the ventral pustular warts
vary from white to pale salmon to pale blue. Palmar and
plantar coloration varies from brown to reddish brown
to purplish-maroon. Tips of the digits vary from dark
brown to reddish brown. Males have a ventrally directed
cloacal tube, but some specimens have the cloaca open-
ing posteriorly at the upper level of the thighs rather
than ventrally at the end of a cloacal tube. The pres-
ence of ventrally directed cloacal tubes has been over-
looked in most species descriptions because they are
highly flexible and can retract depending on the position
of the individual, due to preservation artifacts or onto-
genic changes— juveniles seem to have shorter cloacal
tubes ([16], A.G. Gluesenkamp personal observation)..
Distribution and natural history
Osornophryne occidentalisis currently known from four
localities on the western slopes of the Cordillera Occi-
dental of northwestern Ecuador, between 2500 m and
2750 m (Fig. 5). Its presence in southern Colombia
is plausible, but no specimens are known even from
the northernmost Ecuadorian province of Carchi. The
two localities in the province of Imbabura are less than
20 km apart in the Cordillera de Toisán, a mountain
range part of the Cordillera Occidental of the Andes of
Ecuador. The two localities in the province of Pichincha
are 13 km apart in the Cordillera de Mindo, and 40 km
from the localities in Imbabura.
Osornophryne occidentalis inhabits primary and old-
secondary montane cloud forests, usually with dense
canopy and understory, abundant arboreal and terrestrial
epiphytes, and dense leaf litter. Specimens collected at
the type locality were found in secondary forest domi-
nated by small trees up to 10 m tall covered with abun-
dant epiphytes such as orchids, anthuriums, bromeli-
ads, and ferns. Similar conditions were observed at
Cordillera de Toisán, except that forest was primary and
Figure 5: Schematic map of Ecuador showing the distribution
of Osornophryne occidentalis (circles); from south to north:
near Guarumos, type locality; Guantopungo; and the northern-
most circle marks La Delicia and F. Rodríguez’s farm. Pre-
cise locations can be visualized in Google Earth by downloading
the supporting online material available at <http://www.cisneros-
heredia.org/osornophryne/occidentalis.kmz>
its canopy up to 25 m tall. Osornophryne occidentalis
has been collected under leaf litter or root mats, on moss-
covered floor, and in terrestrial and arboreal bromeliads
during day and early afternoon (J. D. Lynch field data,
D.F. Cis-neros-Heredia and A.G. Gluesenkamp personal
observation).
Osornophryne occidentalis may have clustering behav-
ior; adult female holotype and the topotypic paratype
were collected together on leaf litter and moss at the
base of an arboreal bromeliad (1 m above the floor).
Possible clustering behavior has also been observed in
O. antisana (several individuals collected under a log,
L. A. Coloma personal communication, A. G. Gluesen-
kamp personal observation), O. bufoniformis (several
individuals collected in and under terrestrial bromeli-
ads, M. H. Yánez-Muñoz personal communication, A.
G. Gluesenkamp personal observation), and in O. suma-
coensis (two males found together underneath a dead
bromeliad, A. G. Gluesenkamp personal observation).
Etymology
The specific epithet, a noun in opposition, of this new
species refers to its unique distribution in the montane
cloud forests on the western slope of Cordillera Occi-
dental of the Andes of Ecuador.
Suggested common name
Western Andean Plump Toad
Avances, 2010, Vol. 2, No. 3, Pags. B64-B73 Cisneros-Heredia y Gluesenkamp
Conservation status
We suggest that Osornophryne occidentalis be classi-
fied as Endangered by IUCN criteria, EN B1ab(i,ii,iii,iv),
2ab(i,ii,iii,iv). Although several specimens have been
collected at most known localities and habitat quality
varies from primary to secondary vegetation, the range
of this species is small (extent of occurrence = 555 km2;
Fig. 5) and confined to a rather narrow altitudinal band
(2500–2750 m a.s.l.). Populations in the Cordillera de
Toisán are within the buffer zone of the Cotacachi-Caya-
pas Ecological Reserve, but they are severely threatened
by habitat loss and fragmentation due to uncontrolled
expansion of the agricultural frontier and unsustainable
extraction of natural resources [17]. At least one pop-
ulation (La Delicia) may be already extinct due to ex-
treme habitat destruction (recent expeditions have failed
finding additional specimens). Various areas across the
Cordillera de Toisán are conceded for mining explo-
ration [17]. Direct and indirect environmental impacts
of mining operations may affect populations of O. occi-
dentalis (e.g., habitat transformation and fragmentation,
soil pollution). Populations in the Cordillera de Mindo
have enjoyed apparently better conservation conditions,
with many areas highly inaccessible, protected by sev-
eral private reserves, or part of two protected forests:
Mindo-Nambillo and Cuenca Alta del Río Guayllabam-
ba [17]. However, they are also impacted by habitat loss
and fragmentation. The area is crossed by the recently-
constructed oil pipeline “Oleoducto de Crudos Pesados”
that opened access for illegal colonization of areas that
were previously inaccessible [17].
Taxonomic remarks on some Osornophryne
Knowledge about the morphology, ecology, behavior,
breeding biology, and other evolutionary and biological
aspects of the natural history of Osornophryne is ex-
tremely scarce because few individuals have been ob-
served in the field or deposited in museums. However,
instead of being ecologically rare, most Osornophryne
seem to be difficult to find due to their habits (at least
some species are semi-fossorial) and few survey efforts
concentrated in their preferred habitats (M. H. Yánez-
Muñoz personal communication). Under the right eco-
logical circumstance it seems that some species of Os-
ornophryne may attain large densities; for example O.
cf. bufoniformis was reported as the most common spe-
cies at some localities in the Llanganates National Park
[18] and more than 60 specimens were rescued after
the flooding of 130 ha for the Salve-Faccha dam (M.H.
Yánez-Muñoz personal communication).
Due to insufficient knowledge on the morphological and
chromatic variation of Osornophryne, identification has
been troublesome. Osornophryne guacamayo has been
reported from four localities in Ecuador and two re-
cent localities in Colombia (i.e., Ecuador: Cordillera de
Los Guacamayos—type-locality, Sumaco volcano and
Figure 6: Hand (A) and foot (B) of textitOsornophryne guaca-
mayo(FHGO 1458).
Reventador volcanoes, and the Amazonian slopes of An-
tisana volcano; Colombia: Vereda Vichoy, Valle de Si-
bundoy; D. Lombeida personal communication, D.F. Cis-
neros-Heredia, personal observation, [5, 16, 10, 19]; Ap-
pendix 1). However, some specimens reported from
Colombia differ from Ecuadorian populations in the dor-
sal skin texture and body size and more than one taxa
may be involved (M. H. Yánez-Muñoz and D. F. Cisneros-
Heredia personal observation).
Osornophryne antisanaoccurs on the páramos and high-
lands on the eastern Andes and has been reported from
five localities in Ecuador and a recent locality from Co-
lombia (i.e., Ecuador: Antisana volcano—type local-
ity, Sumaco volcano, Cordillera de los Llanganates and
Vía Salcedo-Oriente, and Guandera; Colombia: Pasto;
[5, 6, 10, 20]). Specimens from Guandera and surround-
ings (i.e., La Angelina) are very similar to the holotype
of O. antisana and we agree that they seem to belong to
the species. All other records correspond to misidentifi-
cations or uncertain records.
Gluesenkamp [6] reported O. antisana from the Sumaco
volcano based on two adult males and a subadult female
(QCAZ 4573–75), but subsequent morphological and
molecular analysis showed that they were erroneously
identified and correspond to O. sumacoensis (A. Glue-
senkamp unpublished data). The irregular borders of
the transverse process in both sexes and blue venter of
females are the best diagnostic characters of O. suma-
coensis [6]. When first studied, the specimens from
Sumaco were sufficiently similar to Hoogmoed’s [5] de-
scription of O. antisana, but knowledge of ontogenic
change and sexual dimorphism in Osornophryne was
little known at that time; now it is recognized that ju-
veniles and males have generalized morphologies and
species specific morphologies are most apparent in adult
females ([16], D. F. Cisneros-Heredia and A. G. Glue-
senkamp personal observation). Gluesenkamp [6] also
reported Osornophryne antisana from Cordillera de los
Llanganates, Tungurahua, Ecuador; but a female from
Llanganates was identified as O. bufoniformis by Hoog-
Cisneros-Heredia y Gluesenkamp Avances, 2010, Vol. 2, No. 3, Pags. B64-B73
moed [5], and Ortíz and Morales [18] reported the pop-
ulation from Llanganates as O. bufoniformis. Further
studies are needed to determine the identity of the Os-
ornophryne from Llanganates.
When Hoogmoed [5] described Osornophryne antisana,
he considered it was closely related and rather alike O.
talipes. Mueses [10] reported O. antisana and O. talipes
for the first time from Colombia. He pointed out that his
specimen of O. antisana did not coincide with the origi-
nal description of the species [5] or with notes by Glue-
senkamp ([6], based on true O. antisana specimens), but
it was similar to the figure presented by Gluesenkamp
([6]: Fig. 1.D, based on misidentified O. sumacoen-
sis as mentioned above). Mueses [10] provided a re-
description of O. antisana based on his Colombian spec-
imen (ICN 12264). Yet, some characters used by Mue-
ses [10] were attributed to O. talipes by Hoogmoed [5].
In general, most diagnostic characters presented by Mue-
ses [10] and Hoogmoed [5] to differentiate O. antisana
from O. talipes are highly variable due to sexual dimor-
phism and ontogenic changes and are easily affected by
preservation (e.g., condition of the proboscis and cloa-
cal tube, head width vs. head length, and extension of
nuptial pads). However, three characters suggest that O.
antisana and O. talipes are different taxa: (1) body size:
O. talipes has larger SVL (adult males 23.8–24.3 mm)
than O. antisana (adult male 18.6 mm) and this differ-
ence is expected to be greater in females; (2) snout form:
O. talipes has a longer snout than O. antisana due to a
much longer loreal region; (3) dorsal skin texture and
coloration: O. talipes has a smooth dorsum with ele-
vated and distinctly-colored glandular ridges, whereas
O. antisana has a smooth to shagreen dorsal skin with
flat and mostly indistinct glandular ridges. Based on
descriptions provided by Mueses [10] and the examina-
tion of some Colombian specimens, we consider that
the Colombian specimen (ICN 12264) referred to O.
antisana does not belong to that species and may cor-
respond to an undescribed taxon. Definitive conclu-
sions about the identity of the phenetically similar pop-
ulations occurring in southeastern Colombia and north-
eastern Ecuador and their association with the names
antisana and talipes will require careful examination
of type-material and additional fresh topotypic mate-
rial to understand intraspefic variation. A molecular ap-
proach would likely be useful in elucidating relation-
ships among problematic populations. We emphasize
the importance of understanding individual, ontogenic
and sexual variation in Osornophryne. It is important to
remember species-specific morphologies are better ex-
pressed in adult females while juveniles and males have
generalized morphologies.
Discussion
Osornophryne is a bufonid genus diagnosed by the ab-
sence of the alary and posterolateral processes of the
hyoid plate, absence of the columella and tympanum,
quadratojugal reduced in size, pectoral girdle firmister-
nal with complete fusion of epicoracoid cartilages, six
presacral vertebrae, coccyx with greatly expanded lat-
eral processes fused to sacral diapophyses, phalangeal
formula variable, palmate hands and feet with extensive
webbing between digits, cloaca situated on a ventrally-
directed tube in both sexes in most species, amplexus
inguinal, large and unpigmented eggs terrestrially de-
posited [3, ?]. Osornophryne is the only bufonid genus
that lacks the alary process of the hyoid and the only
Neotropical genus with six presacral vertebrae [21]. It is
distinctive among Neotropical bufonids in having hands
and feet connected by thick webbing forming a fleshy
platform from which in some cases only the tips of the
fingers and toes protrude.
Osornophryne guacamayohas the most recognizable dis-
crete fingers and toes among the genus (Fig. 6), while
O. antisana and O. bufoniformis have hands and feet
where fingers and toes are hardly differentiated ([5]:
Fig. 7 and 9, D. F. Cisneros-Heredia and A. G. Glue-
senkamp personalobservation). Osornophryne occiden-
talis shows intermediate condition, where fingers (par-
ticularly the fifth) and toes are recognizable but still
forming a thick platform (Fig. 3). Ruiz-Carranza and
Hernández-Camacho [3] suggested the morphology of
hands and feet of O. bufoniformis and O. percrassa (i.e.,
extensively webbed digits forming a massive platform)
was an adaptation towards terrestrial habits. Hoogmoed
[5] suggested that the microhabitats reported for the types
of O. guacamayo and O. antisana refuted that hypoth-
esis and suggested the inverse tendency. The type se-
ries of O. guacamayo was collected on “moss in rock
crevices” and Hoogmoed [5] hypothesized that the spe-
cies was adapted for locomotion in saxicolous micro-
habitats. The microhabitat reported for the holotype of
O. antisana was an arboreal bromeliad ([5], based on
a letter written by the donor not the collector of the
specimen). Subsequent research has revealed that O.
guacamayo is more arboreal than terrestrial, although it
can occupy both microhabitats ([6, 16] D. F. Cisneros-
Heredia personal observation); and that O. antisana and
O. bufoniformis live in semi-fossorial and terrestrial con-
ditions, being common towards the center of terrestrial
bromeliads and beneath them (data for more than 50
specimens recently collected, M. H. Yánez-Muñoz per-
sonal communication). The report of O. antisana inside
an arboreal bromeliad is either an unusual event or an
error of the donor whom described the general situation
for different frogs collected with O. antisana rather than
its specific microhabitat. The occupation of both terres-
trial and arboreal habitats by O. occidentalis, a species
with discernible digits, seems to support the hypothesis
that more arboreal species have more discernible dig-
its while more terrestrial taxa have more compact hands
and feet.
Acknowledgments
For loan of specimens, provision of working space, and
Avances, 2010, Vol. 2, No. 3, Pags. B64-B73 Cisneros-Heredia y Gluesenkamp
kind hospitality, we thank L. A. Coloma (QCAZ), A.
Almendáriz (EPN), M. H. Yánez-Muñoz (MECN), R.
W. McDiarmid, W. R. Heyer, G. R. Zug (USNM), L.
Trueb, W. E. Duellman (KU), J. D. Lynch (ICN), and
J. M. Touzet, A. M. Velasco (FHGO). We are espe-
cially thankful to J.-M. Touzet who first identified O.
occidentalis as different and called DFCH’s attention to
it; to M. Hoogmoed who provided useful comments
and suggestions after examining one of the first col-
lected specimens; and to M. H. Yánez-Muñoz for his
extensive and constructive comments. We thank R. W.
McDiarmid, M. Hoogmoed, M. H. Yánez-Muñoz, J.
J. Mueses-Cisneros, L. A. Coloma, A. Quiguango, M.
Swartz, M. Bustamante, and J. M. Touzet for provid-
ing information, field data, literature, and useful dis-
cussions; N. Acosta, L. Coloma, D. Gluesenkamp, D.
Lombeida, J. Molineros, J. Price, A. Vallejo, V. Zak, A.
León-Reyes, T. Sugahara, K. Valarezo, and J. Robayo
for field companionship and support; and Sebastián Cruz
for the photographs of Osornophryne occidentalis and
O. guacamayo. DFCH is thankful to R. W. McDiarmid
and G. A. Wyngaard and to J. M. Guayasamin and E.
Bonaccorso for their hospitality during his visits to Wash-
ington, D.C., and Lawrence, KS, respectively. Research
by AGG was supported by the Dorothea Bennett Memo-
rial Graduate Fellowship Fund, a fellowship from the
International Studies/Student Travel Office of U. T. Aus-
tin, a Sigma Xi Research Fellowship, and a President’s
Undergraduate Fellowship from U.C. Davis. Research
by DFCH was supported by Ma. Elena and Laura Here-
dia, Universidad San Francisco de Quito, the 2002 Re-
search Training Program at the National Museum of
Natural History / Smithsonian Institution, the Smith-
sonian Women’s Committee, King’s College London,
and the Russell E. Train Education for Nature Program
of the Word Wildlife Fund–WWF. Research and export
permits were issued to AGG (several under QCAZ aus-
pices) and to DFCH (024-EXP-CIEN-FLO-DBAP-MA
under FHGO auspice) by the Ministry of Agriculture
and the Ministry of Environment of the Republic of Ecua-
dor.
Appendix 1: Material examined
Osornophryne antisana: ECUADOR: Vía Salcedo-Oriente, 3500-3600
m (QCAZ 1647-49, 9322); near La Angelina, ca. 3200 m (QCAZ
10047); Represa Salve Faccha, 3400-3500 m (DHMECN 0801-0831).
Osornophryne bufoniformis: ECUADOR: 15 km ESE El Carmelo
(USNM 193537); 1 km NW of Santa Bárbara (USNM 193538); Santa
Bárbara, 2650 m (KU 189945, QCAZ 11472). COLOMBIA: Páramo
Puracé, Laguna San Rafael (KU 144113-15, USNM 322775); Puracé,
3450 m (KU 145036-37); 23 km E Puracé, 3275 m (KU169134-35);
26 km E Puracé, 3180 m (KU 169136); 12 km E Pasto, 3050 m
(KU 169137, 169139-40). Osornophryne guacamayo: ECUADOR:
Cordillera de los Guacamayos, ca. 2200 m (QCAZ 1268-69, 2735,
3266, 3274, 3908, 7387, 8940, 9310-11, 9313-15, 13939-40); Vol-
cán Sumaco, 2500 m (QCAZ 4576-84); Volcán Reventador (QCAZ
9314, FHGO 1458-60); Sierra Azul (QCAZ 4121, 4889, 7402). Os-
ornophryne occidentalis: ECUADOR: near Guarumos (DHMECN
3520 - holotype, DFCH-USFQ GU036 - paratype, DFCH-USFQ
GU034, GU035); Cordillera de Toisán (QCAZ 10141, 9318, 36894 -
paratypes, 9319-21, 36895, KU 132126); near Guantopungo (FHGO
1907, 2981, 2984). Osornophryne percrassa: COLOMBIA: out of
Manizales, on old road to Nevado de Ruiz (USNM 153125); Páramo
de Herveo, E slope Cordillera Central (USNM 200467). Tolima, Ca-
jamarca, km 111-112 carretera Cajamarca-Calarcá (La Línea), 3040-
3080 m (ICN 10015-23); Herveo, 3100 m (ICN 01922, 01924-25,
02624-25). Osornophryne puruanta: ECUADOR: Laguna de Pu-
ruanta, Cordillera de Pimampiro, 3000 m (QCAZ 11471, 7684-85);
Laguna de San Marcos, 3400 m (QCAZ 13271, 13320, EPN 7081-
83). Osornophryne sumacoensis: ECUADOR: Volcán Sumaco, 2500
m (QCAZ 4570 - holotype, QCAZ 4571-72 - paratypes, 4573-75).
Osornophryne talipes: ECUADOR: Nudo de Mojanda, 3400 m (KU
131797 - holotype, KU 131798 - paratype). Osornophryne cf. tal-
ipes: COLOMBIA: Pasto, Hacienda La Marquesa, 3400-3450 m (ICN
12264); Parque Nacional Natural Puracé, 2880 m (ICN 07568, 07572);
carretera Pasto-Volcán Galeras, 3630 m (ICN 12252-53, 12255, 12262-
63). Osornophryne sp.: ECUADOR: Volcán Chiles, 3252 m (QCAZ
735, 9316-17). Estación Científica Los Encinos, 3600 m (DHMECN
2166-71). Km 51 vía Tulcán-Maldonado, 3252 m (QCAZ 9316). 23
km SW Tulcán (QCAZ 9316).
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... Each year, several species are described from montane forests of this biodiversity hotspot (e.g., Rojas-Runjaic et al., 2018;Guayasamin et al., 2019;Paez and Ron, 2019;Reyes-Puig et al., 2019b;San-ta-Cruz et al., 2019;Yanez-Muñoz et al., 2019;Acevedo et al., 2020;Ospina-Sarria et al., 2020;Lehr et al., 2021). Most described species from Ecuador belong to the hyper-diverse genus Pristimantis (Paez and Ron, 2019;Reyes-Puig et al., 2020a), but diversity in other anuran taxa has also increased considerably (e.g., Osornophryne, Hyloscirtus, Noblella, Centrolenidae; Mueses- Cisneros-Heredia and Gluesenkamp, 2010;Yánez-Muñoz et al., 2010a;Páez-Moscoso and Guayasamin, 2012;Almendáriz et al., 2014;Guayasamin et al., 2017aGuayasamin et al., , 2019Reyes-Puig et al., 2019c). ...
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