Accepted by M. Vences: 29 Jul. 2009; published: 10 Sep. 2009
ISSN 1175-5326 (print edition)
1175-5334 (online edition)
Copyright © 2009 · Magnolia Press
Zootaxa 2227: 53–62 (2009)
Description of a new species of Teratohyla Taylor (Amphibia: Athesphatanura:
Centrolenidae) from north-western Ecuador
DIEGO F. CISNEROS-HEREDIA
, MARIO H. YÁNEZ-MUÑOZ
H. MAURICIO ORTEGA-ANDRADE
Universidad San Francisco de Quito, Colegio de Ciencias Biológicas & Ambientales, Calle Diego de Robles y Via. Interoceánica,
Campus Cumbayá, Edif. Charles Darwin, DW010, Quito, Ecuador. E-mail: email@example.com or
Museo Ecuatoriano de Ciencias Naturales, Sección Vertebrados, División de Herpetología, Quito, Ecuador
King's College London, Department of Geography, Strand, London, UK
Instituto de Ecología, A.C., Departamento de Biodiversidad y Ecología Animal, Veracruz, México
Corresponding author. E-mail: firstname.lastname@example.org or email@example.com
We describe a new species of Glassfrog of the family Centrolenidae. This new taxon, Terarohyla sornozai sp. nov., is
diagnosed by having a protruding snout in lateral profile, uniform green dorsal colouration in life, concealed prepollex,
extensive webbing between the outer fingers, fully webbed toes, and small body size. It inhabits the Non-Seasonal
Evergreen Foothill and Lowland forests in the provinces of Esmeraldas, Imbabura, and Pichincha, north-western
Key words: Centrolenidae, new species, taxonomy, Teratohyla sornozai sp. nov., Ecuador
Describimos una nueva especie de rana de cristal de la familia Centrolenidae. Este nuevo taxón, Teratohyla sornozai sp.
nov., es diagnosticado por tener el hocico protuberante en perfil lateral, la coloración dorsal verde uniforme en vida, el
prepolex oculto, membranas extensas entre los dedos externos de la mano, dedos del pie con membranas interdigitales
completas y tamaño corporal pequeño. Esta especie habita en los bosques Siempreverdes No-Estacionales Piemontanos
y de Tierras Bajas en las provincias de Esmeraldas, Imbabura y Pichincha en el noroccidente de Ecuador.
Palabras clave: Centrolenidae, nueva especie, taxonomía, Teratohyla sornozai sp. nov., Ecuador
The diversity of species of frogs of the family Centrolenidae is highest in Colombia and Ecuador, diminishing
northward and southward. The number of species in Ecuador has increased significantly since Lynch &
Duellman (1973) and Duellman & Burrowes (1989), with 10 species described as new since 2004; and
currently 32% (48 spp.) of the known anuran species of the family Centrolenidae (glassfrogs) have been
reported to inhabit in the Republic of Ecuador (see Cisneros-Heredia & McDiarmid, 2007 and subsequent
publications: Cisneros-Heredia et al., 2008; Cisneros-Heredia & Morales-Mite, 2008; Guayasamin et al.,
2008a; Yánez-Muñoz & Cisneros-Heredia, 2008; Yánez-Muñoz et al. 2009). Knowledge about the glassfrogs
from Ecuador is still fragmented and its species richness is expected to increase with the study of unidentified
CISNEROS-HEREDIA ET AL.54 · Zootaxa 2227 © 2009 Magnolia Press
museum specimens and the discovery of additional species with the exploration of previously uncollected
areas (Cisneros-Heredia & McDiarmid 2006a, b, 2007).
Despite been explored since the second half of the 19
century, western Ecuador holds several
undescribed or unreported amphibian species particularly on the Andean foothill forests and on the Cordillera
de la Costa. The foothill forests along the western base of the Andes of Ecuador hold a species-rich amphibian
fauna but have remained almost unstudied until recently. Various expeditions in the last years to the western
Andean foothills resulted in several new country records as well as in the discovery of new species of frogs
(e.g. Cisneros-Heredia & Yánez-Muñoz 2007; Yánez-Muñoz & Cisneros-Heredia 2008). Recent surveys
conducted in foothill forests in the provinces of Esmeraldas, Imbabura, and Pichincha on north-western
Ecuador by the Museo Ecuatoriano de Ciencias Naturales resulted in the collection of an undescribed species
of glassfrog which we are pleased to describe herein.
Materials and methods
Characters, terminology, measurements, and organization of the sections are standardised following
definitions provided by Cisneros-Heredia & McDiarmid (2007). Taxonomic arrangement follows
Guayasamin et al. (2009). The following measurements (in millimetres) were taken with electronic 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 diameter, ED; inter-orbital distance, IOD; eye-nostril distance, EN; inter-narial
distance, IN; width of disc on the third finger, TD, horizontal tympanum diameter; 3DW; tibia length, TL; foot
length, FL. For the webbing formulae, we use the notational device of Savage & Heyer (1967) as modified by
Savage & Heyer (1997). We use the standardised numeration system that codes the fingers in order from
innermost to outermost by Roman numerals from I to IV. Sex and sexual maturity was determined by direct
examination of the condition of gonads and development of secondary sexual characters (vocal slits and
nuptial pads). Specimens were fixed in 4% formalin and stored in 75% ethanol. We examined specimens
(Appendix I and those cited in Cisneros-Heredia & McDiarmid 2007) deposited in the following collections:
DHMECN—División de Herpetología, Museo Ecuatoriano de Ciencias Naturales, Quito; DFCH-USFQ—
Universidad San Francisco de Quito, Quito; QCAZ—Museo de Zoología, Pontificia Universidad Católica del
Ecuador, Quito; ICN—Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá; KU—
The University of Kansas, Natural History Museum, Lawrence; USNM—National Museum of Natural
History, Washington, D.C. Classifications of vegetation formations follow the definitions by Sierra (1999) and
the zoogeographic regions and endemic areas follow the definitions by Albuja et al. (1980) with modifications
by Cisneros-Heredia (2006, 2007).
Teratohyla sornozai sp. nov.
Holotype. DHMECN 3522, an adult female collected at a small stream tributary of Río Naranjal (00°21’05”
N, 78°55’01” W, 750 m elevation), cantón Cotacachi, provincia de Imbabura, República del Ecuador, 01 May
2006 by H. M. Ortega-Andrade.
Paratypes. DHMECN 2634, adult male, Reserva Biológica Canandé (00°18’22.5” N, 79°08’17.6” W,
550 m elevation), cantón Quinindé, provincia de Esmeraldas, República del Ecuador, 21 August 2004, M.
Yánez-Muñoz, P. Meza-Ramos, M. M. Reyes & A. Loaiza. DHMECN 4308, adult female, Bosque Protector
Mashpi (00°10’2.34” N, 78°52’2.32” W, 1100 m), provincia de Pichincha, República del Ecuador, May 2007,
M. H. Yánez-Muñoz y C. Castro.
Zootaxa 2227 © 2009 Magnolia Press · 55
A NEW TERATOHYLA (CENTROLENIDAE) FROM ECUADOR
FIGURE 1. Teratohyla sornozai sp. nov., adult female (paratype, DHMECN 4308) collected at Bosque Protector
Mashpi, province of Pichincha, Ecuador. Photo by H. M. Ortega-Andrade.
Referred material. DHMECN 3521, a juvenile female with the same data of the holotype.
Diagnosis. A glassfrog species (Figs. 1, 2, 3E) diagnosed from all other Centrolenidae by the combination
of the following characters: (1) vomerine teeth present; (2) snout truncate in dorsal view, protruding in profile;
nostrils slightly elevated producing an slight depression in the internarial area; loreal region concave; (3)
tympanic annulus evident, oriented dorsolaterally; very weak supratympanic fold above the tympanum; (4)
dorsal skin slightly shagreen with minute granulations; (5) ventral skin coarsely areolate; subcloacal tubercles
absent but subcloacal area granular; other cloacal ornamentation absent; (6) parietal peritoneum (condition
P3), visceral peritoneum translucent except for pericardium covered by iridophores (condition V1); (7) liver
tri-lobed (condition H0); (8) humeral crista dorsalis not forming a humeral spine in adult males; (9) webbing
basal between fingers I and II; webbing formula for outer fingers II 1
–3 III 2
–1IV (Fig. 2); (10) webbing on
feet I 1
–1 II 1
IV 1–1 V (Fig. 2); (11) no dermal folds or tubercles on hands, upper arm or feet;
ulnar and tarsal folds present (12) unpigmented nuptial pad Type I; concealed prepollex; (13) first finger
longer than second, (14) eye diameter larger than width of disc on finger III; (15) colour in life, uniform bright
green dorsum, green bones; (16) colour in preservative, dorsal surfaces deep grey-purple with few blurred
cream spots; (17) iris golden copper with dark fine reticulations; (18) melanophores present on fingers and
toes; (19) males call from upper side of leaves and ferns around smalls streams; call undescribed; (20, 21, 22)
fighting behaviour, egg clutches, and tadpoles unknown; (23) snout-vent length in adult females 20.4–21.3
mm (n = 2); adult male paratype 19.0 mm.
Comparisons. Teratohyla sornozai (Fig. 1) is the only Glassfrog with protruding snout in profile, uniform
green dorsal colouration, extensive webbing between outer fingers, and fully webbed toes. Extensive webbing
between the outer fingers, humeral crista ventralis not developed into a humeral spine and uniform dorsal
CISNEROS-HEREDIA ET AL.56 · Zootaxa 2227 © 2009 Magnolia Press
colouration also occurs in the following species: Rulyrana adiazeta (Ruiz-Carranza & Lynch), Teratohyla
amelie (Cisneros-Heredia & Meza-Ramos), Rulyrana orejuela (Duellman & Burrowes), T. spinosa (Taylor),
R. susatamai (Ruiz-Carranza & Lynch), and Cochranella xanthocheridia Ruiz-Carranza & Lynch. However,
all differ from T. sornozai as follows: Rurylana adiazeta and R. susatamai have rounded to truncate snouts in
lateral profile, less extensive webbing between the fingers and not fully webbed toes, abundant melanophores
on hands and feet, a different iris colouration (brown with dark brown reticulations), and they inhabit the
eastern slopes of Cordillera Oriental and Central of Colombia. Teratohyla amelie has a rounded snout in
lateral profile, iridophores covering the visceral peritonea, transparent parietal peritoneum, and inhabits the
central and southern lowlands and foothills of Amazonian Ecuador. Rulyrana orejuela has a truncate snout in
lateral profile, less webbing between the outer fingers and not fully webbed toes, abundant melanophores on
hands, feet, and ventral surfaces, different iris colouration (dark grey with yellow ring around pupil), and
larger SVL in adult males (27.3–28.3 mm). Teratohyla spinosa has a truncate snout in lateral profile, less
webbing between the outer fingers and not fully webbed toes, and adult males have a prepollical spine at the
base of the thumb. Cochranella xanthocheridia has a rounded to truncate snout, less webbing between the
outer fingers and not fully webbed toes, and it inhabits the northern portion of the Cordillera Occidental of
Colombia. Further, in preservative, T. sornozai shows a deep grey-purple colouration, while all other species
show a lavender colouration.
FIGURE 2. Ventral view of hand and foot of Teratohyla sornozai sp. nov. (adult female, holotype, DHMECN 3522).
Description of the holotype. Adult female, SVL = 21.3 mm (Fig. 20). Body slender. Head distinct,
slightly wider than long, and wider than body; HW/HL = 1.05, HW/SVL = 0.37, HL/SVL = 0.35. Snout short,
truncate in dorsal view, protruding in profile, EN/HL = 0.30; nostrils slightly elevated producing a slight
depression in the internarial area; canthus rostralis indistinct; concave loreal region; lips flared. Eyes large,
ED/HL = 0.35, directed anterolaterally at about 45° from midline, eyes can be seen when viewed from below,
Zootaxa 2227 © 2009 Magnolia Press · 57
A NEW TERATOHYLA (CENTROLENIDAE) FROM ECUADOR
interorbital area wider than eye diameter, IOD/ED = 1.54, EN/ED = 0.87, EN/IOD = 0.56. Tympanic annulus
evident, oriented dorsolaterally with dorsal inclination; weak supratympanic fold from behind the eye to the
insertion of the arm, separated from orbit by a longer distance than the tympanum diameter. Dentigerous
processes of vomers present, choanae moderately sized, rounded, near margin of mouth; tongue rounded,
slightly notched behind, not indented posteriorly.
FIGURE 3. (A): Teratohyla amelie; (B): Teratohyla midas; (C): Teratohyla pulverata; (D): Teratohyla spinosa; (E):
Teratohyla sornozai sp. nov. Photos by P. Meza-Ramos, A. Georges, M. H. Yánez-Muñoz, R. W. McDiarmid, and H. M.
CISNEROS-HEREDIA ET AL.58 · Zootaxa 2227 © 2009 Magnolia Press
Skin of dorsal surfaces of head, body, and limbs slightly shagreened with minute granulations. Ventral
surfaces coarsely areolate; cloacal opening directed posteriorly at upper level of thighs; no distinct cloacal
sheath, some small low tubercles around the cloacal opening; subcloacal tubercles absent, other cloacal
Upper arm thin, forearm moderately robust, breadth of upper arm about half that of forearm. Humeral
spine absent; ulnar fold present, tubercles absent. Relative lengths of fingers III > IV > I > II; webbing basal
between fingers I and II, webbing formula for other fingers I 1–3 III 2
–1 IV; bulla absent; finger discs slightly
wide, truncate; disc on third finger slightly larger than those on toes, and shorter than eye diameter, 3DW/ED
= 0.62; all subarticular tubercles rounded and elevated; indistinct supernumerary tubercles present in fingers;
palmar tubercle large, rounded, plane; thenar tubercle elliptic, rather indistinct.
Hind limbs slender; TL/SVL = 0.63, FL/SVL = 0.53. Tarsal fold present; inner metatarsal tubercle large
and elliptical, flat; outer metatarsal tubercle indistinct. Subarticular tubercles rounded and low; supernumerary
tubercles small, rather indistinct. Webbing on foot I 1
–1 II 1
IV 1–1 V; discs truncate on all toes,
Measurements of the holotype. Snout-vent length, 21.3 mm; head width, 7.9 mm; head length, 7.5 mm;
horizontal eye diameter, 2.6 mm; inter-orbital distance, 4.0 mm; eye-nostril distance, 2.3 mm; internarial
distance between the nostrils, 1.8 mm; width of disc on the third finger, 1.6 mm; tibia length, 13.3 mm; foot
length, 11.2 mm.
Colouration. In life (Fig. 1, 3E), dorsal surface uniform bright green, yellowish towards the flanks,
without light or dark flecks, spots, or ocelli. Upper eyelids yellowish-green or bright green; tips of digits and
interdigital webbing dull yellow to greenish-yellow. Throat and ventral surfaces of legs bluish-green. Parietal
peritoneum white (condition P3). Bones pale green. Iris golden copper to golden brown with dark thin
In preservative, dorsum deep grey-lavender with few blurred cream spots; dorsal surfaces of arms grey-
lavender, hind limbs cream-greenish or lavender; upper eyelid dark blue; lips cream. All ventral surfaces
cream. Parietal peritoneum white (condition P3); all visceral peritonea clear except for the white pericardium
TABLE 1. Variation of measurements (in mm) of adult Teratohyla sornozai sp. nov. See text for abbreviations.
Variation. Variation of measurements and body proportions is presented in Tables 1 and 2. Photographs
of specimens in life are shown in Figs. 1 and 3. Gross morphological and colouration features are fairly
Specimen DHMECN 3522 DHMECN 4308 DHMECN 2634
Holotype Paratype Paratype
Sex Female Female Male
SVL 21.3 20.4 19.0
HW 7.9 7.3 6.5
HL 7.5 7.2 6.7
ED 2.6 2.8 2.3
IOD 4.0 2.6 3.5
EN 2.3 2.5 2.4
IN 1.8 1.7 1.7
TD 0.9 0.9 0.8
3DW 1.6 1.6 1.2
TL 13.3 13.7 11.2
FL 11.2 13.3 8.8
Zootaxa 2227 © 2009 Magnolia Press · 59
A NEW TERATOHYLA (CENTROLENIDAE) FROM ECUADOR
invariant; male paratype, adult female paratype, and juvenile female are similar to female holotype. Hand and
foot webbing does not vary (Fig. 2). In preservative a few blurred cream irregular spots appear in some
specimens (DHMECN 3521–22), while other remain uniform lavender. These spaces are produced by the
absence of melanophores rather than by the presence of light pigmentation (iridophores or xanthophores).
Humeral spine and prepollical spine absent in both sexes. Adult male is smaller than adult females (SVL male
=19.0 mm; SVL females = 20.4–21.3 mm). Males have unpigmented nuptial pads Type–I.
TABLE 2. Variation of proportions of adult Teratohyla sornozai sp. nov. See text for abbreviations.
Etymology. The specific name of this Glassfrog is a noun in the genitive case and a patronym for our
friend Francisco (Pancho) Sornoza, Ecuadorian ornithologist and conservationist. He is president of
Fundación Jocotoco, an Ecuadorian NGO that manages the Reserva Biológica Canandé, and has actively
support the research activities of the Museo Ecuatoriano de Ciencias Naturales that resulted in the discovery
of this new species.
Distribution and natural history. Teratohyla sornozai is known from three localities in the provinces of
Esmeraldas, Imbabura, and Pichincha, northwestern Ecuador (Fig. 4). This species occur in Foothill
Evergreen Forests (DHMECN 3521, 3522, 4308) and Lowland Evergreen Forests (DHMECN 2634), with an
altitudinal range between 550 and 1100 m elevation. The female holotype and juvenile female were found
sitting on leaves of herbs and ferns up to 2 meters above a small stream, Naranjal River. No egg clutches or
tadpoles were seen in the area. At Naranjal, T. sornozai was found in sympatry with Espadarana prosoblepon
(Boettger), Pristimantis labiosus (Lynch, Ruiz-Carranza & Ardila-Robayo), and Hyloscirtus alytolylax
(Duellman). Although just females where found at Naranjal, males were calling in the surroundings but over 5
meters above the stream. At Canandé, the male was found calling from the upper surface of a leave and in
sympatry with E. prosoblepon, Sachatamia albomaculata (Taylor), S. ilex (Savage), Hyloscirtus palmeri
(Boulenger), Hypsiboas boans (Linnaeus), and H. picturatus (Boulenger). At Mashpi, the male was collected
Specimen DHMECN 3522 DHMECN 4308 DHMECN 2634
Holotype Paratype Paratype
Sex Female Female Male
HW/HL 1.01 1.05 0.97
HW/SVL 0.36 0.37 0.34
HL/SVL 0.35 0.35 0.35
EN/HL 0.34 0.30 0.35
EN/HW 0.34 0.29 0.36
EN/ED 0.88 0.87 1.02
EN/IOD 0.95 0.56 0.67
ED/HL 0.39 0.35 0.35
ED/HW 0.38 0.33 0.36
ED/3DW 1.79 1.63 1.92
IOD/ED 0.93 1.54 1.52
IOD/HW 0.36 0.51 0.54
IOD/HL 0.36 0.53 0.53
IN/IOD 0.66 0.44 0.49
TD/ED 0.32 0.33 0.35
TL/SVL 0.67 0.63 0.59
FL/SVL 0.65 0.53 0.46
TL/FL 1.03 1.19 1.27
CISNEROS-HEREDIA ET AL.60 · Zootaxa 2227 © 2009 Magnolia Press
in sympatry with Pristimantis labiosus (Lynch, Ruiz-Carranza & Ardila-Robayo), E. prosoblepon,
Hyloscirtus palmeri, and H. picturatus.
FIGURE 4. Schematic map showing the known localities of Teratohyla sornozai sp. nov. in Ecuador. Type-locality
marked by an empty dot.
Remarks. Teratohyla sornozai is similar to T. spinosa, sharing a similar body form and size, uniform
dorsal colouration, lobed liver, ventral parietal peritoneum covered by iridophores, and gastrointestinal and
hepatic peritonea without iridophores. We hypothesise that both are sister taxa based on their high
morphological similarity, but this remains to be tested. Teratohyla sornozai has not been found in sympatry
with T. spinosa. It is possible that both occur together particularly on lowland areas. In contrast with T.
spinosa that prefers low riverine vegetation along relatively quiet areas (Kubicki, 2007; pers. obs.); T.
sornozai apparently prefers higher strata along riverine vegetation (over 1.5 m) especially in the splash zone
Zootaxa 2227 © 2009 Magnolia Press · 61
A NEW TERATOHYLA (CENTROLENIDAE) FROM ECUADOR
We have not found egg clutches of Teratohyla sornozai but we have observed egg clutches of T. amelie, T.
midas, T. pulverata, and T. spinosa. Females of T. amelie, T. midas, and T. pulverata deposit their egg clutches
on the upper side of leaves, while T. spinosa may use both sides of leaves. Eggs are laid in a laminar array
(Altig and McDiarmid 2007) at different locations on the leave (centre or borders, showing intra-specific
variation). The egg clutches of T. amelie, T. midas and T. spinosa keep the original arrangement and location
where they were deposited, but the egg clutches of T. pulverata frequently absorb water and swell
considerably, forming a 3-dimensional clump that moves due to gravity to the tip of the leave and hangs
(Cisneros-Heredia & McDiarmid 2007: 28). Eggs of all four species have a greenish-tinted colouration
(Kubicki 2007; pers. obs.), which is apparently produced by the accumulation of biliverdin in the eggs, and
seems to be common to all members of this clade, probably a synapomorphy.
Teratohyla sornozai corresponds to the species mentioned as Cochranella sp. N11 by Yánez-Muñoz &
We thank J. D. Lynch (ICN), L. Coloma (QCAZ), W. E. Duellman and L. Trueb (KU), and R. W. McDiarmid,
G. W. Zug, and W. R. Heyer (USNM) for allowing access to specimens under their care; and to P. Meza-
Ramos (DHMECN), R. W. McDiarmid, and A. Georges (Applied Ecology Research Group, University of
Camberra) for allowing the use of their photographs. MHYM is grateful to F. Sornoza-Molina, N. Simpson
and R. Ridgely of the Fundación Jocotoco for their support to the surveys at Canande; to C. Castro, M. P.
Meza-R., M. Reyes-P., A. Loaiza, F. Timpe, and G. Velez for their field companionship and assistance at
Candande and Mashpi; and to M. Altamirano, P. Mena-Valenzuela, and C. Carrera of the Museo Ecuatoriano
de Ciencias Naturales for their constant support. HMOA thanks A. Ortiz T., C. Tobar, M. Arellano, R.
Cabrera, F. Narváez, and F. Armas for field companionship and help; to C. Ponce and M. Albarracín for
supporting his research between 2004 and 2006; and to M. Larrea, P. Tufiño and P. Moreno for their support at
the Monte Saíno and Río Naranjal reserves. DFCH express his gratitude to Ma. E. Heredia and L. Heredia for
their continuous support. This work was supported by the Fondo Ambiental del I. Municipio del Distrito
Metropolitano de Quito as part of the project “Monitoreo biológico: una herramienta para el manejo
adaptativo de las áreas protegidas y bosques protectores del Distrito Metropolitano de Quito, DMQ”;
Fundación Jocotoco; Fundación SIRUA; Museo Ecuatoriano de Ciencias Naturales; the Research Training
Program of the National Museum of Natural History, Smithsonian Institution; the Russel E. Train Education
for Nature Program, Word Wildlife Fund WWF; Conservation International; and the Iniciativa de Especies
Amenazadas managed by EcoCiencia, Conservation International, and the Royal Netherlands Embassy
(grants to DFCH and HMOA). Research and collecting permits (Nº 018 -IC-FAU-DNBAP/MA, 2003-2004,
and Nº 010) were provided by Ministerio del Ambiente del Ecuador granted to Museo Ecuatoriano de
Albuja, L., Ibarra, M., Urgilés, J. & Barriga, R. (1980) Estudio preliminar de los vertebrados ecuatorianos.
Departamento de Ciencias Biológicas, Escuela Politécnica Nacional, Quito, Ecuador.
Altig, R. & McDiarmid, R.W. (2007) Morphological diversity and evolution of egg and clutch structure in amphibians.
Herpetological Monographs, 21, 1–32.
Cisneros-Heredia, D.F. (2006) Distribution and ecology of the western Ecuador frog Leptodactylus labrosus (Amphibia:
Anura: Leptodactylidae). Zoological Research, 27 (3), 225–234.
Cisneros-Heredia, D.F. (2007) Distribution and natural history of the Ecuadorian snake Dipsas andiana (Boulenger,
1896) (Colubridae: Dipsadinae) with considerations on its conservation status. Russian Journal of Herpetology,
CISNEROS-HEREDIA ET AL.62 · Zootaxa 2227 © 2009 Magnolia Press
Cisneros-Heredia, D.F. & McDiarmid, R.W. (2007) Revision of the characters of Centrolenidae (Amphibia: Anura:
Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1–
Cisneros-Heredia, D.F. & Yánez-Muñoz, M.H. (2007) A new species of Glassfrog (Centrolenidae) from the southern
Andean foothills on the west Ecuadorian region. South American Journal of Herpetology, 2 (1), 1–10.
Cisneros-Heredia, D.F. & Morales-Mite, M.A. (2008) A new species of glassfrog from the elfin forests of the Cordillera
del Cóndor, southeastern Ecuador (Anura: Centrolenidae). Herpetozoa, 21 (1/2), 49–56.
Cisneros-Heredia, D.F., Venegas, P.J., Rada, M. & Schulte, R. (2008) A new species of glassfrog (Anura: Centrolenidae)
from the foothill Andean forests of Ecuador and Peru. Herpetologica, 64 (3), 341–353.
Duellman, W.E. & Burrowes, P.A. (1989) New species of frogs, Centrolenella, from the Pacific Versant of Ecuador and
Southern Colombia. Occasional Papers of the Museum of Natural History of the University of Kansas, 132, 1–14.
Guayasamin, J.M., Cisneros-Heredia, D.F. & Castroviejo-Fisher, S. (2008a) Taxonomic identity of Cochranella petersi
Goin, 1961 and Centrolenella ametarsia Flores, 1987. Zootaxa, 1815, 25–34.
Guayasamin, J.M., Castroviejo-Fisher, S., Trueb, L., Ayarzagüena, J., Rada, M. & Vilà, C. (2009) Phylogenetic
systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1–
Kubick, B. (2007) Ranas de vidrio de Costa Rica / Glass frogs of Costa Rica. Instituto Nacional de Biodiversidad
(INBio), Heredia, Costa Rica.
Lynch, J.D. & Duellman, W.E. (1973) A review of the Centrolenid frogs of Ecuador, with descriptions of new species.
Occasional Papers of the Museum of Natural history of the University of Kansas, 16, 1–66.
Savage, J.M. & Heyer, W.R. (1967) Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central
America. Beitrage zur Neotropischen Fauna, 5, 111–131.
Savage, J.M. & Heyer, W.R. (1997) Digital webbing formulae for anurans: a refinement. Herpetological Review, 28, 131
Sierra, R. (1999) Propuesta Preliminar de un Sistema de Clasificación de Vegetación para el Ecuador Continental.
Proyecto INEFAN/GEF-BIRF EcoCiencia, Quito, Ecuador.
Yánez-Muñoz, M.H. & Cisneros-Heredia, D.F. (2008) Notes on geographic distribution. Amphibia, Anura,
Centrolenidae: Cochranella orejuela, first country records from Ecuador. Check List, 4(1), 50–54.
Yánez-Muñoz, M, Pérez-Peña, P. & Cisneros-Heredia, D.F. (2009) New country records of Hyalinobatrachium
iaspidiense (Amphibia, Anura, Centrolenidae) from the Amazonian lowlands of Ecuador and Peru. Herpetology
Notes, 2, 49–52.
Appendix I. Examined material
H = holotype; P = paratype.
Cochranella xanthocheridia: ICN 27758 (H); ICN 27757 (P); ICN 10643–6 (P). Rulyrana adiazeta: ICN 17919 (H).
Rulyrana orejuela: KU 145081 (H); KU 145080 (P); DHMECN 04309; DHMECN 04551–2. Rulyrana susatamai:
ICN 18641 (H); ICN 15801 (P). Teratohyla amelie: DHMECN 3066 (H); DHMECN 3591. Teratohyla midas: ICN
23755 (P); DFCH-USFQ D102; QCAZ 22876; QCAZ 20001-2; USNM 286702–05; USNM 286707; USNM
288437; USNM 342716. Teratohyla pulverata: USNM 342214–21; USNM 219379–87; DHMECN 2612, 3194–5;
QCAZ 11367–8; DHMECN 2612; QCAZ 32224. Teratohyla sornozai: DHMECN 3522 (H); DHMECN 2634 (P);
DHMECN 3521. Teratohyla spinosa: USNM 288443; USNM 286741–44; USNM 219388–94.