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We present new information on several species of centrolenid frogs from Ecuador and Peru that justify the placement of Centrolene fernandoi Duellman and Schulte as a junior synonym of Centrolenella audax Lynch and Duellman; Centrolenella puyoensis Flores & McDiarmid as a synonym of Centrolenella mariae Duellman & Toft; and Cochranella tangarana Du-ellman & Schulte as a synonym of Cochranella saxiscandens Duellman & Schulte.
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Notes oN the taxoNomy of some GlassfroGs from the aNdes
of Peru aNd ecuador (amPhibia: ceNtroleNidae)
dieGo f. cisNeros-heredia1,3
JuaN m. GuayasamiN2
We present new information on several species of centrolenid frogs from Ecuador and Peru that
justify the placement of Centrolene fernandoi Duellman and Schulte as a junior synonym of
Centrolenella audax Lynch and Duellman; Centrolenella puyoensis Flores & McDiarmid
as a synonym of Centrolenella mariae Duellman & Toft; and Cochranella tangarana Du-
ellman & Schulte as a synonym of Cochranella saxiscandens Duellman & Schulte.
K-W: Centrolene fernandoi; Centrolene audax; Nymphargus puyoensis; Nymphargus
mariae; Rulyrana saxiscandens; Rulyrana tangarana; Rulyrana spiculata; Synonymy.
Glassfrogs are conspicuous members of riverine
communities across Neotropical America and have
more than 140 described species (Frost, 2013). Twen-
ty-nine species of glassfrogs have been reported from
Peru (see below). Nymphargus ocellatus Boulenger was
the first centrolenid species to be described from Peru
(Boulenger, 1918), and no further glassfrogs were
reported from the country until Duellman’s (1976)
description of Nymphargus truebae Duellman and Ru-
lyrana spiculata Duellman, who also reported Nym-
phargus siren (Lynch & Duellman) and Hyalinoba-
trachium munozorum Lynch & Duellman. In 1979,
Nymphargus mariae Duellman & Toft was described
from the Serranía del Sira. Cannatella & Duellman
(1982) re-evaluated the Peruvian specimens assigned
to N.siren and regarded them as a different species:
Nymphargus phenax (Cannatella & Duellman). They
also described Nymphargus pluvialis (Cannatella &
Duellman), and provided the first Peruvian records
for Teratohyla midas (Lynch & Duellman) and Hya-
linobatrachium bergeri (Cannatella). Centrolene azulae
(Flores & McDiarmid) was described in subsequent
years from an isolated mountain range on the east-
ern Andes of Peru, while Centrolene hesperium (Cadle
& McDiarmid) and Cochranella euhystrix (Cadle &
McDiarmid) were described from the Pacific slopes
of northwestern Andean Peru (Flores & McDiar-
mid, 1989; Cadle & McDiarmid, 1990). Duellman
& Wild (1993) provided the first country record of
Centrolene buckleyi Boulenger. Duellman & Schulte
(1993) almost doubled the number of Peruvian cen-
trolenids with the description of eight species from
Volume 54(12):161-168, 2014
¹ Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Laboratorio de Zoología Terrestre,
calle Diego de Robles y Ave. Interoceánica, Campus Cumbayá, edif. Darwin, DW010-A, Casilla Postal 17-1200-841, Quito, Ecuador.
² Universidad Tecnológica Indoamérica, Centro de Investigación de la Biodiversidad y el Cambio Climático, Av. Machala y Sabanilla,
Quito, Ecuador.
³ King’s College London, Department of Geography, London, England, United Kingdom.
the eastern slopes of Cordillera Central and adjacent
ridges in the department of San Martín: Centrolene
fernandoi Duellman & Schulte, Centrolene lemnisca-
tum Duellman & Schulte, Centrolene muelleri Du-
ellman & Schulte, Nymphargus chancas (Duellman
& Schulte), Cochranella croceopodes Duellman &
Schulte, Rulyrana saxiscandens (Duellman & Schulte),
R. tangarana (Duellman & Schulte), and Hyalino-
batrachium lemur Duellman & Schulte. Nymphar-
gus mixomaculatus (Guayasamin, Lehr, Rodríguez
& Aguilar) was described from central Andean Peru
(Guayasamin et al., 2006). Torres-Gastello et al.
(2007) described Rulyrana erminea Torres-Gastello,
Suárez-Segovia & Cisneros-Heredia, and reported
the first records of Cochranella resplendens (Lynch &
Duellman) and Vitreorana oyampiensis Lescure from
Amazonian Peru. Cisneros-Heredia etal. (2008) de-
scribed Rulyrana mcdiarmidi from Ecuador and Peru,
and presented the first record of Nymphargus posadae
from Peru. Yánez-Muñoz et al. (2009) reported the
first Peruvian record of Hyalinobatrachium iaspidiense
Ayarzagüena from Amazonian Peru. Castroviejo-Fish-
er etal. (2009) described Hyalinobatrachium carlesvi-
lai Castroviejo-Fisher, Padial, Chaparro, Aguayo & de
la Riva from the Amazonian slopes of central Andean
Peru, assigned all previous records of H.munozorum
from Peru either to H. carlesvilai or to H. bergeri,
synonymized H. lemur with H. pellucidum Lynch
& Duellman, and extended the distribution of the
latter south to the department of Cusco, southern
Peru. Catenazzi etal. (2012) described Centrolene sa-
bini Catenazzi, von May, Lehr, Gagliardi-Urrutia &
Guayasamin from the Amazonian slopes of southeast-
ern Andean Peru. Catenazzi & Venegas (2012) pre-
sented photographs of Chimerella mariaelenae (Cisne-
ros-Heredia & McDiarmid) from Kampankis, in the
Amazonian slopes of northeastern Peru.
While developing our extensive reviews of Cen-
trolenidae (Cisneros-Heredia & McDiarmid, 2007;
Guayasamin et al., 2009), we cooperatively found
that there is no evidence to support specific recogni-
tion of several populations of Peruvian and Ecuador-
ian centrolenids currently hypothesised as different
species. Herein, we present our findings in an effort
to enhance the understanding on the diversity and
conservation of Neotropical amphibians.
Characters and terminology are standardized
following the definitions provided by Cisneros-Here-
dia & McDiarmid (2007). Taxonomy and systemat-
ics follow Guayasamin et al. (2009). The following
measurements (in millimetres) were taken with elec-
tronic digital callipers (0.05mm accuracy, rounded to
the nearest 0.1mm): snout-vent length, SVL; head
width, HW; head length, HL; horizontal eye diam-
eter, ED; inter-orbital distance, IOD; eye-nostril dis-
tance, EN; inter-narial distance, IN; width of disc on
the third finger, 3DW; tibia length, TL; foot length,
FL. Upper eyelid width was not measured because of
its limited utility due to preservation bias. We use the
notational device for webbing formulae of Savage &
Heyer (1967), as modified by Savage & Heyer (1997).
Sex and sexual maturity was determined by direct ex-
amination of the condition of gonads and develop-
ment of secondary sexual characters (vocal slits and
nuptial pads). We examined specimens (AppendixI)
deposited in the following collections: DHMECN
– División de Herpetología, Museo Ecuatoriano de
Ciencias Naturales, Quito; DFCH-USFQ – Universi-
dad San Francisco de Quito, Quito; QCAZ – Museo
de Zoología, Pontificia Universidad Católica del Ec-
uador, Quito; BMNH – Natural History Museum,
London; KU – The University of Kansas, Natural
History Museum, Lawrence; USNM – National Mu-
seum of Natural History, Washington, D.C.; MCZ
– Museum of Comparative Zoology, Harvard Univer-
Centrolene audax Lynch & Duellman, 1973
Centrolenella audax Lynch & Duellman, 1973.
Centrolene audax – Ruiz-Carranza & Lynch, 1991.
“Centrolene” audax – Guayasamin etal., 2009.
Centrolene fernandoi – Duellman & Schulte, 1993.
Holotype: KU 211770. Type locality: west slope
of Abra Tangarana, 7km (by road) northeast
of San Juan de Pacaysapa (06°12’S, 76°44’W,
1080m), Provincia Lamas, Departamento San
Martín, Perú. New synonymy.
Lynch & Duellman (1973) described Centrole-
nella audax for glassfrog populations diagnosed as
having small yellow spots on the dorsum, short and
distally-curved humeral spines in males, and extensive
webbing between outer fingers from the Amazonian
versant of the northern Andes. Centrolene audax is
currently known in Colombia and Ecuador from few
localities in Low Montane Evergreen Forest on the
Amazonian versant of the Andes, between 1350 and
            
1800 m (Mueses-Cisneros, 2005; Cisneros-Heredia
& McDiarmid, 2007; Yánez-Muñoz etal., 2010).
Centrolene fernandoi Duellman & Schulte
(1993) was described based on nine specimens col-
lected on the western slope of Abra Tangarana,
Amazonian versant of the Andes of Peru. Duellman
& Schulte (1993) compared C.fernandoi with Cen-
trolene audax and reported its high similarity, but dif-
ferentiated them by its snout form (bluntly rounded
in C. fernandoi, truncate in C. audax), dorsal skin
texture (with scattered small spicules in C.fernandoi,
without spicules in C.audax), dorsal fleck colouration
(bluish-white in C. fernandoi, golden in C. audax),
finger colouration (pale green in C. fernandoi, pale
yellow in C.audax), and iris background colouration
(silvery green in C.fernandoi, pale bronze in C.au-
dax). Centrolene fernandoi remains known only from
its type locality (Frost, 2011).
We examined six specimens of Centrolene fer-
nandoi (type-series) and 42 specimens of Centrolene
audax (including the type-series). We found that all
differences used to separate them are intraspecifically
variable within C.audax. Snout form of C.audax var-
ies between rounded to truncate in profile, presence of
spicules shows sexual and ontogenic variation (visible
in reproductive males, and absent in non-reproduc-
tive males and females), dorsal flecks vary from pale
yellow to golden yellow (furthermore, the photograph
of C.fernandoi in the original description shows pale
yellow spots), finger colouration varies from pale
green to bright yellow, and iris colouration varies
from pale bronze to silvery green or mustard with
thin black reticulation. Since no discrete differences
are evident and their populations have no obvious
biogeographic barriers, we place Centrolene fernandoi
Duellman & Schulte, 1993 as a junior synonym of
Centrolenella audax Lynch & Duellman, 1973. There-
fore, Centrolene audax inhabits Low Montane Ever-
green Forest on the Amazonian versant of the Andes
of southern Colombia, Ecuador, and northern Peru,
between 1080 and 1800 m (Duellman & Schulte,
1993; Mueses-Cisneros, 2005; Cisneros-Heredia &
McDiarmid, 2007; Yánez-Muñoz etal., 2010).
Nymphargus mariae (Duellman & Toft, 1979)
Centrolenella mariae Duellman & Toft, 1979.
Centrolenella puyoensis – Flores & McDiarmid, 1989.
Holotype: MCZ 91187, by original designation.
Type locality: “1.0 km W Puyo, Provincia de
Pastaza, Ecuador, between 1000-1050m eleva-
tion”. New synonymy.
Cochranella mariae – Ruiz-Carranza & Lynch, 1991.
Cisneros-Heredia & McDiarmid, 2007.
Cochranella puyoensis – Ruiz-Carranza & Lynch,
1991. Cisneros-Heredia & McDiarmid, 2006.
Centrolene mariae – Duellman & Schulte, 1993.
Centrolene puyoensis – Duellman & Schulte, 1993.
Centrolene puyoense – Stuart etal., 2008.
Nymphargus mariae – Guayasamin etal., 2009.
Nymphargus puyoensis – Guayasamin etal., 2009.
Centrolenella mariae Duellman & Toft was de-
scribed based on one female specimen collected at
Serranía de Sira, department of Huánuco, Peru (Du-
ellman & Toft, 1979). Flores & McDiarmid (1989)
described Centrolenella puyoensis and Centrolenella
azulae, hypothesising that, together with C.mariae,
they formed a monophyletic group (the C. mariae
species-group). Subsequent authors (Ruiz-Carranza
& Lynch, 1991, 1995; Duellman & Schulte, 1993)
followed this hypothesis, but Cisneros-Heredia &
FIGURE 1: Photographs of Centrolene audax: (A)KU 164500
from 2 km SSW of Río Reventador, Napo, Ecuador; (B) KU
211770, holotype of Centrolene fernandoi, from W slope of Abra
Tangarana, San Martín, Peru. Photos by W.E. Duellman.
      163
McDiarmid (2006, 2007) questioned the validity
of the C.mariae species-group, further pointing out
that although C.azulae is diagnosable, C.mariae and
C.puyoensis are very similar and probably conspecific
(Cisneros-Heredia & McDiarmid, 2007). Centrole-
nella mariae and C.puyoensis were placed in the genus
Nymphargus by Guayasamin et al. (2009) based on
morphological and molecular data, respectively. Flores
& McDiarmid (1989) separated N. puyoensis from
N.mariae by tympanum exposure (three-quarters in
N.puyoensis, one-half in N.mariae), hand and foot
webbing (slightly more extensive in N.mariae), ulnar
fold (present in N.puyoensis, absent in N.mariae), in-
tricate cloacal ornamentation (present in N.puyoensis,
FIGURE2: Photographs of Nymphargus mariae: (A)KU 174713, holotype of Centrolenella mariae, from Serranía de Sira, Huanuco, Peru;
(B)MCZ 91187, holotype of Centrolenella puyoensis, from 1kmW of Puyo, Pastaza, Ecuador. (C)DHMECN 4752, from Conambo,
Pastaza, Ecuador. Photo A by M. Bustamante; B by JMG; C by H.M. Ortega-Andrade.
            
absent in N.mariae) and differences in proportions
(greater eye-nostril/eye diameter and shank length/
snout-vent length in N.puyoensis).
We examined 16 specimens assignable to Nym-
phargus puyoensis from Ecuador and the holotype of
Nymphargus mariae and found no evidence to support
their differentiation. All characters used to diagnose
these two species can be attributed to subtle differenc-
es that fall inside the intraspecific variation of a single
species. Flores & McDiarmid (1989) proposed a po-
larization of characters that is rather subjective and bi-
ased due to their small sample size (one specimen for
each of their “species”). Variation of tympanum expo-
sure, webbing, and body proportions observed among
N.puyoensis and N.mariae is continuous and similar,
or even lower than the natural differences observed
within populations of other species of centrolenids.
The absence of ulnar folds and cloacal ornamenta-
tions in the type specimen of N.mariae could be at-
tributed to natural variation, but also due to preser-
vation artifacts (see Cisneros-Heredia & McDiarmid,
2006 for information on the variation of specimens of
N.puyoensis). Either way it has also been observed in
specimens of N.puyoensis.
In the absence of valid discriminating evidence
to support the hypothesis that Nymphargus puyoensis
and Nymphargus mariae are different lineages, we place
Centrolenella puyoensis Flores & McDiarmid, 1989 as
a synonym of Centrolenella mariae Duellman & Toft,
1979. Thus, Nymphargus mariae, as herein redefined,
inhabits Foothill Evergreen Forest and Lowland Ev-
ergreen Forest flooded by White-water Rivers on the
Amazonian versant of the Andes of Ecuador and Peru
(Cordillera del Sira), between 300 and 1550m (Flores
& McDiarmid, 1989; Cisneros-Heredia & McDiar-
mid, 2006, 2007; Yánez-Muñoz etal., 2010).
This synonym reflects the absence of evidence
to support the hypothesis that the population from
the Serranía del Sira in eastern Amazonian Peru (type-
locality of Nymphargus mariae) is different from those
of eastern Amazonian Ecuador. Although it might be
argued that the Serranía del Sira is a rather isolated
mountain range that likely contains several amphib-
ian endemics, our decision to place Nymphargus puy-
oensis in the synonymy of N.mariae is based on the
fact that there are no morphological traits that sup-
port the existence of two putative species, and that
potential biogeographic barriers cannot justify specif-
ic status without the corroboration of traits intrinsic
to the organisms. We encourage future researchers to
analyse other lines of evidence to evaluate the status of
these populations.
Rulyrana saxiscandens (Duellman & Schulte, 1993)
Cochranella saxiscandens Duellman & Schulte, 1993.
Cochranella tangarana Duellman & Schulte, 1993.
Type locality: “west slope of Abra Tangarana,
7km (by road) northeast of San Juan de Pacay-
sapa (06°12’S, 76°44’W), 1080 m), Provincia
Lamas, Departamento San Martín, Perú”. New
Rulyrana saxiscandens – Guayasamin etal., 2009.
Rulyrana tangarana – Guayasamin etal., 2009.
Duellman & Schulte (1993) described Co-
chranella saxiscandens and Cochranella tangarana
based on specimens collected at two nearby localities
of the Mayo River, Tarapoto region, department of
San Martín, Peru. Duellman & Schulte (1993) dif-
FIGURE3: Photographs of Rulyrana saxiscandens: (A)KU 211776, holotype of Cochranella tangarana, and (B)KU 211779, holotype of
Cochranella saxiscandens, both from Abra Tangarana, San Martín, Peru. Photos by W.E. Duellman.
      165
ferentiated these two species (now placed in the ge-
nus Rulyrana) by their snout form (bluntly round in
Rulyrana saxiscandens, truncate in Rulyrana tangara-
na), dorsal colouration in preservative (dark-grey to
black in R. saxiscandens, lavender in R. tangarana),
melanophores on the ventral surfaces of shanks and
tarsi (present in R.saxiscandens, absent in R. tanga-
rana); presence of spicules on dorsal surfaces (absent
in R. saxiscandens, present in R. tangarana); and in-
ner tarsal fold (absent in R.saxiscandens, present in
We examined 22 specimens of Rulyrana saxis-
candens and two of Rulyrana tangarana (including
all type-specimens) and found that all stated differ-
ences between them actually correspond to intraspe-
cific variation. Snout shape varies continuously from
round to truncate; tympanic annulus is conspicuous
at different degrees due to the supratympanic fold;
dorsal colouration in preservative varies continuously
from dark-purple, purplish-grey, dark-lavender, to
light-lavender (similar colour variation has been ob-
served in Rulyrana flavopunctata); melanophores are
always present on ventral surfaces although some-
times scarce; spicule presence and appearance varies
ontogenically and sexually (see Cisneros-Heredia &
McDiarmid, 2007); and inner tarsal fold is always
present but sometimes poorly noticeable.
In the absence of evidence to support the hy-
pothesis that two species are involved in the popu-
lations of the Mayo River, we place Cochranella tan-
garana Duellman & Schulte, 1993 as synonym of
Cochranella saxiscandens Duellman & Schulte, 1993.
Rulyrana saxiscandens remains very similar to
Rulyrana spiculata (Duellman, 1976), which is known
from forests on the Amazonian versant of the Andes
of central and southern Peru and eastern Bolivia,
between 1200 and 1700m (Frost, 2011; Rodríguez
et al., 2004). Duellman & Schulte (1993) reported
that Centronella saxiscandens (and Centrolenella tanga-
rana) were similar to Centrolenella spiculata, but dif-
fered due to snout form, tympanum and inner tarsal
fold appearance, coloration, and presence of spicules
on dorsal surfaces in Rulyrana tangarana. Evan Twom-
ey and associates are studying these species, and we
refer to them for a definitive conclusion.
With the present changes, the diversity of glass-
frogs of Peru currently includes 29 species: Centrolene
audax, C.azulae, C.buckleyi, C.hesperium, C.lemnis-
catum, C.muelleri, C.sabini, Chimerella mariaelenae,
Cochranella croceopodes, C. euhystrix, C. resplendens,
Hyalinobatrachium bergeri, H.carlesvilai, H. iaspidi-
ense, H.pellucidum, Rulyrana erminea, R.mcdiarmidi,
R. saxiscandens, R. spiculata, Vitreorana oyampiensis,
Nymphargus chancas, N. mariae, N. mixomaculatus,
N.ocellatus, N.phenax, N.pluvialis, N.posadae, Tera-
tohyla amelie, T.midas.
Hyalinobatrachium munozorum and Centrolene
condor are expected to occur in Peru. Hyalinobatra-
chium munozorum occurs in Ecuador and Bolivia
(Cisneros-Heredia & McDiarmid 2007, Castroviejo-
Fisher et al., 2011), and C. condor is known from
several localities in the Cordillera del Condor, just a
few kilometres from Peruvian territory (Cisneros-He-
redia & Morales-Mite, 2008; Almendáriz & Batallas,
Presentamos nueva información sobre algunas especies
de ranas centrolénidas de Ecuador y Perú que justifica
colocar a Centrolene fernandoi Duellman and Schul-
te como sinónimo junior de Centrolenella audax Lynch
and Duellman, Centrolenella puyoensis Flores &
McDiarmid como sinónimo de Centrolenella mariae
Duellman & Toft, y Cochranella tangarana Duellman
& Schulte como sinónimo de Cochranella saxiscandens
Duellman & Schulte.
P-C: Centrolene fernandoi; Centrolene
audax; Nymphargus puyoensis; Nymphargus mariae;
Rulyrana saxiscandens; Rulyrana tangarana; Rulyrana
spiculata; Sinonimia.
Funding for the development of diverse stages of
this study was obtained from the following: Russell E.
Train Education for Nature Program/WWF, Conser-
vation International, Smithsonian Women’s Commit-
tee, Research Training Program of the National Mu-
seum of Natural History – Smithsonian Institution,
María Elena Heredia, Laura Heredia, and Universidad
San Francisco de Quito to DFCH; and from the proj-
ect “Patrones de diversidad de los anfibios andinos del
Ecuador” of the Universidad Tecnológica Indoaméri-
ca to JMG. We are thankful to Linda Trueb (KU),
Mario Yánez-Muñoz (DHMECN), Roy W. McDiar-
mid, W. Ron Heyer and George R. Zug (USNM),
David Gower and Mark Wilkinson (BMNH), and
James Hanken and José Rosado (MCZ) for granting
access to specimens under their care. We are grate-
ful to Santiago Castroviejo, Marco Rada, and Evan
Twomey for discussions on the taxonomy of Peruvian
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Aceito em: 27/08/2013
Publicado em: 30/06/2014
      167
Examined specimens
Centrolene audax: ECUADOR: Napo: KU 146624 (holotype of Centrolenella audax): Salto de Agua; KU
155502-03: 7kmSW of Río Azuela; KU 164496: Azuela; KU 164497-504: 2kmSSW of Río Reventador;
USNM 286620-22: Cascada de San Rafael; KU 178018-27: Río Salado; USNM 286623-25, MCZ A97807-8:
14.6km (by road) NE of Río Salado; KU 190015: 43kmNE of Santa Rosa; KU 190016: 8.9kmNE Santa
Rosa; KU 143290, 143292: 16,5km NNE Santa Rosa; DHMECN 06788-89: Reserva Biológica Narupa.
PERU: San Martín: KU 211770 (holotype of Centrolene fernandoi), 211771-5: W slope Abra Tangarana.
Nymphargus mariae: ECUADOR: Napo: DFCH-USFQ D285: ca.45 kmE of Narupa. PASTAZA: MCZ
91187 (holotype of Centrolenella puyoensis): 1.0kmW Puyo; USNM 291298: Río Pucayacu. QCAZ 37932:
stream tributary of Río Lliquino; QCAZ 39293: near Villano; DHMECN 04752-53, 04756: Conambo; Orel-
lana: QCAZ 7104, 7499: Río Huataracu; Sucumbíos: DHMECN 06190: Río Verde. PERU: Huanuco: KU
174713 (holotype of Centrolenella mariae): Serranía de Sira.
Rulyrana saxiscandens: PERU: KU 211776 (holotype of Cochranella tangarana), 211777, 217299: W slope
of Abra Tangarana; KU 211779 (holotype of Cochranella saxiscandens), 211780-88, 211789-98, 211800-01:
Cataratas Ahaushiyacu; KU 211802-03: 15kmNE of Tarapoto.
            
... Centrolene fernandoi-Duellman and Schulte, 1993 [174]. Synonymy by Cisneros-Heredia and Guayasamin, 2014 [175]. Espadarana audax-Twomey, Delia, and Castroviejo-Fisher, 2014 [19]. ...
... The dominant frequency of a note measured at peak amplitude is 5426-6718 (mean = 6146, SD = 368) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 5254-6288 (mean = 5925, SD = 374) Hz and a higher limit of 6115-7063 (mean = 6608, SD = 297) Hz. [19,22,[174][175][176], this work). In Ecuador, the species is found within the Eastern Foothill Forest and Eastern Montane Forest ecoregions. ...
... Taxonomic Remarks: The morphological differences that separate Espadarana durrellorum and E. audax are subtle, but given the available genetic and morphological data, we consider them as valid species. "Centrolene" fernandoi was recently placed in the synonymy of E. audax [175]. The specific name callistomma is derived from the Greek kallistos-, meaning "most beautiful" and omma, meaning "eye", as a reference to the fantastic iris pattern in this species [9]. ...
Full-text available
Glassfrogs (family: Centrolenidae) represent a fantastic radiation (~150 described species) of Neotropical anurans that originated in South America and dispersed into Central America. In this study, we review the systematics of Ecuadorian glassfrogs, providing species accounts of all 60 species, including three new species described herein. For all Ecuadorian species, we provide new information on the evolution, morphology, biology, conservation, and distribution. We present a new molecular phylogeny for Centrolenidae and address cryptic diversity within the family. We employ a candidate species system and designate 24 putative new species that require further study to determine their species status. We find that, in some cases, currently recognized species lack justification; specifically, we place Centrolene gemmata and Centrolene scirtetes under the synonymy of Centrolene lynchi; C. guanacarum and C. bacata under the synonymy of Centrolene sanchezi; Cochranella phryxa under the synonymy of Cochranella resplendens; and Hyalinobatrachium ruedai under the synonymy of Hyalinobatrachium munozorum. We also find that diversification patterns are mostly congruent with allopatric speciation, facilitated by barriers to gene flow (e.g., valleys, mountains, linearity of the Andes), and that niche conservatism is a dominant feature in the family. Conservation threats are diverse, but habitat destruction and climate change are of particular concern. The most imperiled glassfrogs in Ecuador are Centrolene buckleyi, C. charapita, C. geckoidea, C. medemi, C. pipilata, Cochranella mache, Nymphargus balionotus, N. manduriacu, N. megacheirus, and N. sucre, all of which are considered Critically Endangered. Lastly, we identify priority areas for glassfrog conservation in Ecuador.
... Some species of glassfrogs (Centrolenidae) are notoriously difficult to identify morphologically due to the overall similarity in shapes and coloration , 2011aKok & Castroviejo-Fisher 2008). The use of molecular techniques, field observations, photographs, recordings of advertisement calls, tadpole morphology and other natural history information often are necessary to confirm identification of cryptic or previously unreported species of glassfrogs (Cisneros-Heredia and Guayasamin 2014, Twomey et al. 2014, Rada et al. 2019. Furthermore, the discipline of taxonomy increasingly incorporates contributions from a wide range of methodological approaches that study evolutionary relationships among species. ...
... This integrative framework has been termed integrative taxonomy and is proving extremely helpful for the accelerating rates of species discovery and description of amphibians common in tropical countries . Peru is no exception to the recent increase in the rate of frog species discovery (Catenazzi et al. 2012, Catenazzi and von May 2014, Cisneros-Heredia and Guayasamin 2014, Twomey et al. 2014. This is also evident for glassfrogs (Twomey et al. 2014;Cisneros-Heredia & Guayasamin 2014, Lujan et al. 2014, with several recent publications reporting new species, genetic and phenotypic variation, as well as range extensions. ...
... Peru is no exception to the recent increase in the rate of frog species discovery (Catenazzi et al. 2012, Catenazzi and von May 2014, Cisneros-Heredia and Guayasamin 2014, Twomey et al. 2014. This is also evident for glassfrogs (Twomey et al. 2014;Cisneros-Heredia & Guayasamin 2014, Lujan et al. 2014, with several recent publications reporting new species, genetic and phenotypic variation, as well as range extensions. Our study of new collected material revealed relevant data regarding species of Cochranella, Hyalinobatrachium and Rulyrana. ...
Full-text available
We used an integrative taxonomy approach to investigate the taxonomic identity of several populations of glassfrogs from Peru, which are notoriously challenging to identify due to their overall similarity in morphology and coloration. We relied on comparisons of morphology, bioacoustics, and partial fragments of 16S rRNA DNA sequences. We report for the first time the presence of Hyalinobatrachium mondolfii in Peru, being this the southernmost locality known for the species. Likewise, we update and extend the distribution ranges of Rulyrana spiculata and Cochranella nola in the Andes of Peru, provide a 16S sequence of a topotype of R. spiculata, and confirm its presence in Bolivia. For all three species, we increase the current knowledge on their geographic distribution and genetic and phenotypic variation.
... The range height of perching between 1.5 -2 m tend to provide favorable conditions to reduce predation by small terrestrial mammals on individuals or eggs. However, it does not represent a risk to the survival of tadpoles because of the impact when they hit the water (Hampton and Otto, 2014;Cisneros-Heredia and Guayasamin, 2014). ...
Full-text available
Objectives: to determine the population status of Centrolene savagei (Centrolenidae) in the Western and Central Andes of Colombia. Scope: promotion of the population study of the endemic glassfrogs species of Colombia. Methodology: C. savagei (Centrolenidae) has only been registered in the Central and Western Andes of Colombia. We conducted a visual encounter survey in eight different localities where C. savagei was previously reported: Alto Bonito, Florencia, La Pastora, Santa Rosa de Cabal, Filandia, Boquía, Alto el Otoño and Chicoral, with an effective sampling effort of 54 h per locality. Each captured individual was marked, sexed and the height above water level of the perch was registered. Furthermore, all detected clutches in each survey were also marked and height above water level was recorded. We use a standard capture-recapture model to estimate the size of the population in each study site assuming closed population conditions. Main Results: C. savagei was absent at La Pastora and Florencia. In all localities where C. savagei was registered, male individuals and clutches were usually found between 1.5-2.0 m, while female individuals remain most of the time above 6 m height, and only go down for reproduction. The population size of C. savagei in the study area of Alto del Otoño was 132 ± 13 ind, in Chicoral was 41 ± 8 ind and in Santa Rosa de Cabal was 37 ± 6 ind. In Filandia, Boquia and Alto Bonito, recaptures were not carried out, so the size of the population could not be estimated. Conclusions: The presence of C. savagei in 75% of the localities where it had previously been reported suggests that the biotic and abiotic conditions in these sites are still adequate to sustain populations of this endemic species.
Full-text available
We performed free sampling by visual encounter survey in order to document the biodiversity of amphibians and reptiles along an altitudinal transect that covered the Andean forest, foothills and lowlands of the cordillera Oriental, between the departments of Huila and Caqueta. In the transect, 50 species of amphibians and 42 species of reptiles were recorded, among which there are 8 new species for science; 6 new reptile records for the department of Caquetá and 4 new amphibian reports for the country (Hyloxalus italoi, H. maculosus, Pristimantis brevicrus and P. diadematus). Ameerega ingeri, Hyloscirtus torrenticola and Anolis ruizii are species in a category of threat; additionally, the introduced lizards Hemidactylus angulatus, H. frenatus and the transplanted species Gonatodes albogularis were registered. When descending, the biota found showed that around 900 m.a.s.l., the Andean herpetofauna and the Amazonian lowland herpetofauna began to be differentiated. Above this height, species considered to be Andean were observed. In amphibians a greater richness was observed between 400 and 700 m., while reptiles presented a greater number of species between 1000 and 1300 m. Comparisons with other studies of amphibians show a greater similarity between this trail and the fauna of Alto del Gabinete (both conducted in the department of Caquetá) and another group with Alto Sibundoy (Putumayo) along with the assemblage from Napo Province (Ecuador).
Full-text available
Peru is well known for amphibian diversity and endemism, yet there have been relatively few field studies of glassfrog (Centrolenidae) diversity in this country. Research in Colombia and Ecuador indicates that centrolenid diversity is higher in the northern Andes. However, part of this trend appears to be due to sampling effort. We conducted fieldwork through- out northern Peru, and based on phylogenetic analysis of DNA sequences, combined with bioacoustic and morphological analyses of new and available material we now recognize 33 species from the country (versus 30 species prior to this work). Field surveys led to the discovery of four remarkable species: Centrolene charapita new species is a large, orna- mented glassfrog that appears to be sister to Ce. geckoideum; Chimerella corleone new species represents the second- known member of the genus Chimerella; Cochranella guayasamini new species is the second-known member of the ge- nus with humeral spines; and Hyalinobatrachium anachoretus new species occurs in the cloud forest of the east-Andean versant in Peru. In addition to the new species described here, we provide new country records, new localities including range extensions of up to 875 km, information on diagnostic characters and phylogenetic relationships, call and larval de- scriptions, and observations on natural history for several Peruvian centrolenids. Our results also revealed several taxo- nomic problems concerning species of the genus Rulyrana, and we conclude that R. croceopodes and R. tangarana are junior synonyms of R. saxiscandens. By implication of our phylogenetic analyses, we recognize the following new com- binations: Espadarana audax new combination, Espadarana durrelorum new combination, and Espadarana fernandoi new combination.
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Presentamos nueva información que extiende la distribución latitudinal y altitudinal de cinco especies de ranas de cristal recientemente descritas y poco conocidas de la región oriental de Ecuador. Incluimos datos novedosos sobre su tamaño corporal e historia natural. Se discute información sobre la diversidad y biogeografía de ranas centrolenidas del oriente de Ecuador, encontrando que se encuentran asociadas con seis formaciones vegetales comprendidas entre las estribaciones orientales y la baja Amazonia. Identificamos tres importantes zonas de diversidad y endemismo en la región oriental de Ecuador asociadas con las cuencas hidrográficas de los ríos Napo, Pastaza y Santiago. Los ecosistemas de bosques Montano Bajos y Piemontanos concentran la mayor diversidad y endemismo para ranas centrolenidas, sin embargo 77% de ellas están amenazadas. Es trascendental juntar todos los esfuerzos posibles para investigar y conservar este substancial grupo de la fauna ecuatoriana.
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The Glassfrog Centrolenella puyoensis Flores & McDiarmid is a taxon known only from the female holotype, and recently placed in the genus Centrolene due to its supposed close relationship with Centrolene mariae (Duellman & Toft). Herein we report new material of puyoensis, including adult male specimens previously unknown. We propose the new combination Cochranella puyoensis (Flores & McDiarmid) n. comb., in recognition of the state of the humeral crista ventralis in males of this species, which lack a humeral spine. The hypothesis of relationships between three species, including puyoensis, proposed as the mariae species-group is questioned as it was based on phenetic rather than derived characters. We present new data that extend the distributional range of Cochranella puyoensis, and define its range along the Foothill Evergreen forests from 400 m to 1000 m above sea level in the provinces of Napo, Orellana, and Pastaza. New data presented herein also permit a re-evaluation of the conservation status of the species, previously classified under the IUCN category of Critically Endangered. We recommend that Cochranella puyoensis be reclassified as "Endangered": EN B1ab(i,ii,iii)+2ab(i,ii, iii); based on a better understanding of the presence of the species, its occupancy area, number of known localities, and habitat quality status.
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We describe a new species of glassfrog from the cloud forest of Manu National Park, southern Peru, at elevations of 2750–2800 m. The new species is similar in morphology to Centrolene lemniscatum, which occurs in northern Peru at elevations of 2000–2280 m. Both species have white labial stripes, humeral spines, and lack vomerine teeth. The new species differs from C. lemniscatum by its larger size, labial stripe extending into a distinct lateral stripe instead of a discontinuous lateral stripe, snout profile inclined anteroventrally instead of bluntly rounded, greater depression in the internarial area, and by having strongly protruding nostrils. Males of the new species emit long calls with 8–14 peaked notes, instead of a short tonal note in C. lemniscatum. Another morphologically similar species, C. buckleyi, has a short advertisement call composed of 1–5 notes, and is genetically distinct from the new species. This new Centrolene extends the known distribution of Centrolene to the south by 600 km, and is the southernmost species of this genus.
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We collected two specimens of the genus Hyalinobatrachium during fieldwork expeditions to the Departamento Pando-the northernmost region of Bolivia situated in the south-western Amazonian basin, within the zone of tall evergreen lowland rainforest. The specimens are deposited in the Coleccion Boliviana de Fauna, La Paz (CBF 6453) and in the National Museum, Prague (NMP6V 74059). Because species identification within Hyalinobatrachium based only on morphological characters is in many cases problematic (Kok & Castroviejo-Fisher 2008; Castroviejo-Fisher et al. 2009), we took advantage of published sequences of Hyalinobatrachium to identify our samples. Our results show that each specimen belongs to a different species (H. mondolfii and H. munozorum), none of them previously known to occur in Bolivia. The taxonomic implications of our discovery are briefly discussed.
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We collected two specimens of the genus Hyalinobatrachium during fieldwork expeditions to the Departamento Pando—the northernmost region of Bolivia situated in the south-western Amazonian basin, within the zone of tall evergreen lowland rainforest. The specimens are deposited in the Colección Boliviana de Fauna, La Paz (CBF 6453) and in the National Museum, Prague (NMP6V 74059). Because species identification within Hyalinobatrachium based only on morphological characters is in many cases problematic (Kok & Castroviejo-Fisher 2008; Castroviejo-Fisher et al. 2009), we took advantage of published sequences of Hyalinobatrachium to identify our samples. Our results show that each specimen belongs to a different species (H. mondolfii and H. munozorum), none of them previously known to occur in Bolivia. The taxonomic implications of our discovery are briefly discussed.
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The frog genus Phyllomedusa is represented in Costa Rica by six species. Analysis of variation in coloration, webbing, and measurements delineates features that distinguish the various forms. The characteristics of the flank pattern in the nominal species P. callidryas and P. helenae, utilized by previous authors to separate them, are shown to be subject to individual and geographic variation. The two forms represent two of many populations within a single species, P. callidryas. Reasons for not using the term subspecies for geographic segments of callidryas are presented. The diagnostic features and the geographic and ecologic distribution of the Costa Rican species, P. annae, P. calcarifer, P. callidryas, P. lemur, P. saltator and P. spurrelli, based on the entire species ranges, are discussed.