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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
ABSTRACT
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.
INTRODUCTION
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
http://dx.doi.org/10.1590/0031-1049.2014.54.12
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.
E-mail:dcisneros@usfq.edu.ec
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 etal. (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 etal. (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 etal. (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.
MATERIALS AND METHODS
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 (0.05mm accuracy, rounded to
the nearest 0.1mm): 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 (AppendixI)
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-
sity.
RESULTS AND CONCLUSIONS
Centrolene audax Lynch & Duellman, 1973
(Fig.1)
Centrolenella audax Lynch & Duellman, 1973.
Centrolene audax – Ruiz-Carranza & Lynch, 1991.
“Centrolene” audax – Guayasamin etal., 2009.
Centrolene fernandoi – Duellman & Schulte, 1993.
Holotype: KU 211770. Type locality: west slope
of Abra Tangarana, 7km (by road) northeast
of San Juan de Pacaysapa (06°12’S, 76°44’W,
1080m), 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
162
1800 m (Mueses-Cisneros, 2005; Cisneros-Heredia
& McDiarmid, 2007; Yánez-Muñoz etal., 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 etal., 2010).
Nymphargus mariae (Duellman & Toft, 1979)
(Fig.2)
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 1000-1050m 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 etal., 2008.
Nymphargus mariae – Guayasamin etal., 2009.
Nymphargus puyoensis – Guayasamin etal., 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,
FIGURE2: 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 1kmW 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.
164
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 1550m (Flores
& McDiarmid, 1989; Cisneros-Heredia & McDiar-
mid, 2006, 2007; Yánez-Muñoz etal., 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)
(Fig.3)
Cochranella saxiscandens Duellman & Schulte, 1993.
Cochranella tangarana Duellman & Schulte, 1993.
Type locality: “west slope of Abra Tangarana,
7km (by road) northeast of San Juan de Pacay-
sapa (06°12’S, 76°44’W), 1080 m), Provincia
Lamas, Departamento San Martín, Perú”. New
synonymy.
Rulyrana saxiscandens – Guayasamin etal., 2009.
Rulyrana tangarana – Guayasamin etal., 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-
FIGURE3: 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
R.tangarana).
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 1700m (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,
2012).
RESUMEN
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.
ACKNOWLEDGMENTS
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
centrolenids.
166
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Aceito em: 27/08/2013
Publicado em: 30/06/2014
167
APPENDIXI
Examined specimens
Centrolene audax: ECUADOR: Napo: KU 146624 (holotype of Centrolenella audax): Salto de Agua; KU
155502-03: 7kmSW of Río Azuela; KU 164496: Azuela; KU 164497-504: 2kmSSW of Río Reventador;
USNM 286620-22: Cascada de San Rafael; KU 178018-27: Río Salado; USNM 286623-25, MCZ A97807-8:
14.6km (by road) NE of Río Salado; KU 190015: 43kmNE of Santa Rosa; KU 190016: 8.9kmNE Santa
Rosa; KU 143290, 143292: 16,5km 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 kmE of Narupa. PASTAZA: MCZ
91187 (holotype of Centrolenella puyoensis): 1.0kmW 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: 15kmNE of Tarapoto.
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