La Marca et al.
40
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
hps://doi.org/10.5281/zenodo.21083985 / Anartia, 42 (junio 2026): 40-48
Updated geographic distribution and significant additional
museum records for the Horned Frog, Ceratophrys calcarata
Boulenger, 1890 (Anura: Ceratophryidae), in Venezuela
Distribución geográfica actualizada e importantes registros de museo adicionales
para el sapo de cuernos, Ceratophrys calcarata Boulenger, 1890
(Anura: Ceratophryidae), en Venezuela
Enrique La Marca1,2, Néstor Sánchez2,3, Iván José Mendoza Mendoza2,
Luis Fernando Navarrete2,4, Santos Bazó5, Luis Merlo6, Luis Alejandro Rodríguez2,
Karel Lemoine Nouel2 & Abraham Mijares-Urrutia7
1Escuela de Geografía, Facultad de Ciencias Forestales y Ambientales, Universidad de Los Andes, Mérida 5101, Venezuela.
orcid.org/0000-0002-6508-3375
2Centro de Conservación REVA (Rescate de Especies Venezolanas de Anfibios), Mérida, Venezuela.
3Departamento de Biología, Facultad de Ciencias, Universidad de Los Andes, Mérida, Venezuela.
4Serpentario. Instituto de Medicina Tropical, Universidad Central de Venezuela, Caracas.
5Serpentario.Com, Urb. Las Acacias, Valera, estado Trujillo, Venezuela.
6Fundatrópicos, Caracas, Venezuela.
7Conservation and Ecosystem Management Teaching Group, Technical and Further Education Institute, (TAFE NSW), Sydney, Australia.
Correspondence: enrique.lamarca@gmail.com
(Received: 15-02-2026 / Accepted: 24-04-2026 / On line: 30-06-2026)
ABSTRACT
e genus Ceratophrys is represented in Venezuela by two species: C. cornuta and C. calcarata. Despite being morphologi-
cally unique amphibians, most of the geographic distribution of C. calcarata in the north of the country has historically
depended on bibliographic mentions and sightings without voucher specimens that allowed scientic verication. e
present work formally reports rst museum voucher specimens for the states of Mérida, Trujillo, Yaracuy and Zulia, and
additional specimens that corroborate the presence of the species in Amazonas and Falcón states. Clarications on unsup-
ported historical records are included, and the importance of certain records is highlighted. ese new data will bring up
to date the geographic distribution map of the species within Venezuelan territory.
Keywords: Amphibia, Anura, Ceratophryidae, Ceratophrys calcarata, geographic distribution, Venezuela.
RESUMEN
El género Ceratophrys está representado en Venezuela por dos especies: C. cornuta y C. calcarata. A pesar de ser anbios
de morfología distintiva, la mayor parte de la distribución geográca de C. calcarata en el norte del país ha dependido
históricamente de menciones bibliográcas y avistamientos sin ejemplares testigo que permitan su vericación cientíca.
En el presente trabajo se reporta formalmente primeros registros de ejemplares de museo para los estados Mérida, Trujillo,
Yaracuy y Zulia, y ejemplares adicionales que corroboran la presencia de la especie en los estados Amazonas y Falcón. Se
incluye aclaratorias sobre registros históricos dudosos y se destaca la importancia de algunos registros. Estos nuevos datos
permiten actualizar el mapa de distribución geográca de la especie en territorio venezolano.
Palabras clave: Amphibia, Anura, Ceratophryidae, Ceratophrys calcarata, distribución geográca, Venezuela.
Ceratophrys calcarata in Venezuela
41
INTRODUCTION
e genus Ceratophrys Wied-Neuwied, 1824, com-
prises a group of robust anuran amphibians with a distinc-
tive morphology, popularly known as “horned frogs”, “sa-
pos de cachos” or “escuerzos”. Currently, eight species are
recognized, distributed throughout the South American
lowlands, covering Venezuela, Ecuador, Peru, Brazil, the
Guianas, Bolivia, Paraguay, and Argentina (Faivovich et
al. 2014, IUCN SSC Amphibian Specialist Group 2020,
2023, Frost 2026). In Venezuelan territory, the genus is
represented by only two species: Ceratophrys calcarata
Boulenger, 1890, and Ceratophrys cornuta (Linnaeus,
1758).
e oldest reference for the genus in Venezuela, as
Ceratophrys cornuta, is that of Ernst (1877). Lutz (1927)
indicated that Ceratophrys would be in this country but
hesitated to assign it to C. cornuta or C. calcarata; howev-
er, Rivero (1961: 85) stated that “it might well have been
an Eleutherodactylus cornutus maussi (= Strabomantis bi-
porcatus)”. Röhl (1942, 1949, 1956, 1959) continued the
use of the name C. cornuta without documenting speci-
mens. Nonetheless, Rohl’s color descriptions and illustra-
tions suggest that it most likely corresponds to Ceratoph-
rys calcarata. None of these early records of C. cornuta for
northern Venezuela have voucher specimens for verica-
tion and could be records of C. calcarata. In this regard,
Rivero (1961) had already shown skepticism about the
presence of C. cornuta in Venezuela and did not refer to
any specimen for the country. e rst sound record (al-
though only photographic) for C. cornuta in Venezuela,
used as a proof of its occurrence in this country, was nally
issued by Rojas-Runjaic et al. (2021). Dierences between
C. cornuta and C. calcarata were provided by Cochran &
Goin (1970); a necessary revision goes beyond the scope
of this paper.
Ginés (1959: 110) was the rst to give a precise locality
for a Ceratophrys in Venezuela (“Río El Palmar”, in Zulia
State), without identifying the species. We examined spec-
imens from this locality (see below), and they correspond
to C. calcarata. In that same work, Ginés mentioned that
Dr. Juan Rivero had informed him of having collected
specimens of C. calcarata “in the region of San Fernando
de Apure” (undoubtedly a lapsus for San Fernando de Ata-
bapo, as inferred from Rivero [1961: 84], who explicitly
indicates that it was in San Fernando de Atabapo where
he learned of a woman who had been bitten by a “sapo de
cachos” (horned toad) in Puerto Ayacucho. is lapsus
may be the source of Barrio-Amorós (1998) stating that
the species was present in San Fernando de Apure, without
documenting any specimens. To date, the presence of C.
calcarata (or any other Ceratophrys) in Apure State (and in
fact, in all the Venezuelan Llanos) remains undocumented.
Rivero (1961) did not provide veriable data (museum
records) regarding the existence of C. calcarata in northern
Venezuela, but for the rst time documented it for Vene-
zuelan territory with two specimens from what is current-
ly Amazonas State. Based on the presence of C. calcarata
there and in Santa Marta (Colombia), he extrapolated that
the species’ distribution range likely included the Llanos
and the Venezuelan Guayana, the Falcón region, and the
lower Maracaibo Lake basin, northeastern Colombia, and
probably the Guianas to the south and the savannas of
northern Brazil.
All subsequent mentions of the species (e. g., Rivero
1963, 1964, Gorham 1966, Lynch 1971, 1975, 1982, Mül-
ler 1973, Gorzula & Cerda 1979, Hoogmoed 1979, Péfaur
& de Pascual 1982, Mercadal de Barrio 1986, La Marca
1992, Yústiz 1996, Barrio-Amorós 1998, 2004, Vázquez
et al. 2011, IUCN 2020, Frost 2026) did not indicate
museum records. e only veriable museum records are
those of Rivero (1961), La Marca (1986), and Schalk et al.
(2014). It is imperative to document new localities of C.
calcarata to understand its total distribution range. In this
context, the primary objective of this article is to formally
report the rst museum voucher specimens for the states
of Mérida, Trujillo, and Yaracuy, and additional specimens
that corroborate the presence of the species in Amazonas
Falcón and Zulia States. is is a comprehensive review
of museum specimens in Venezuela; nonetheless we note
that there might be some more samples in U.S. and Euro-
pean museums, as deducted from some lists available on
the Internet.
NEW GEOGRAPHIC DISTRIBUTION RECORDS
OF CERATOPHRYS CALCARATA
FOR VENEZUELA
Amazonas State
CVULA (Colección de Vertebrados de la Universidad
de Los Andes, Mérida) 7509, from “Puerto Ayacucho,
Amazonas”, and MBUCV (Museo de Biología, Univer-
sidad Central de Venezuela) 886, from “Territorio Ama-
zonas, Amazonas”. ese records are important because
they corroborate the presence of C. calcarata in Amazo-
nas State, a controversial disjunct population. Records
assigned to this state in previous works include UPR-M
196 from “Territorio Amazonas” and UPR-M 197 from
“Pto. Ayacucho” (Rivero 1961), which were alternatively
considered as doubtful specimens of C. cornuta (Barrio-
Amorós, 1998) or a disjunct population of C. calcarata
(Barrio-Amorós et al. 2019). A more recent record (KU
La Marca et al.
42
-University of Kansas- 207528) was provided by Schalk et
al. (2014), also for Puerto Ayacucho.
A picture of a single live animal in Barrio-Amorós et al.
(2019), although without a museum number, is also from
“Puerto Ayacucho, Amazonas”. Another unvouchered re-
cord for the species in Barrio-Amorós (1998) deserves a
comment: “Puerto Ayacucho, Agualinda” probably refers
to a sector of the same name northeast of Puerto Ayacucho
city, while the record from “San Fernando de Atabapo” is
erroneous and most likely a misinterpretation of a com-
ment in Rivero (1961). In that work, the later author men-
tioned that in said locality he heard a report of a “sapo de
cachos” (“horned toad” in English, most likely a Ceratoph-
rys) from Puerto Ayacucho and explicitly noted that the
animal “is not known by the people of San Fernando de
Atabapo” (Rivero 1961: 85).
Falcón State
ere is only one documented museum record for a
locality in the state of Falcón, from the locality of Sanare
(EBRG -Estación Biológica de Rancho Grande, Mara-
cay- 1407, 1516), provided by Mijares-Urrutia & Arends
(1993). Here, we present the second museum record,
represented by specimen UMMZ (University of Michi-
gan Museum of Zoology) 2026, coming from “5 km by
road S of El Vínculo, Paraguaná Peninsula, Falcón State”
(Fig.1C). Other Ceratophrys calcarata from the Paraguaná
Peninsula, for which only photographs are available, are
shown for the rst time in Fig. 1(A, B, D) as examples of
the chromatic variability exhibited by the species in the
region. In addition to the locality of “Moruy, Paraguaná
Peninsula, Falcón municipality” (in Mijares & Arends
2000), also without voucher specimens, the individuals
illustrated here would be among the rst documented re-
cords (albeit only through photos) for the Paraguaná Pen-
insula.
Mérida State
CVULA 0131, from “El Vigía, near Guayabones, 22
km NE of El Vigía, Mérida State, Venezuela”. is consti-
tutes the rst record for the state and the southernmost
locality north of the Orinoco River; the presence of the
Figure 1. Ceratophrys calcarata specimens from Falcon state, Venezuela. (A, B) “Paraguaná”, (C) “Paraguaná, 5 km S El Vínculo”, (D)
“Yaracal, municipio Cacique Manaure”. Photos: A, B (Luis Merlo), C (Allan Markezich), D (Luis Fernando Navarrete and Karel Lem-
oine). Specimens not collected.
Ceratophrys calcarata in Venezuela
43
species in nearby Colombia (Acosta Galvis 2023) at simi-
lar latitudes and in equivalent ecosystems suggests that the
species could also be present on the lowlands close to the
Andean-lacustrine slopes of Táchira State.
Trujillo State
ULABG (Colección de anbios y reptiles, Laborato-
rio de Biogeografía, Universidad de Los Andes, Mérida)
7856-7857, from an articial lake on a farm “18 km W
Sabana de Mendoza, municipio Sucre, estado Trujillo,
Venezuela” (Fig. 2), geographic coordinates (WGS84)
9°25’47.73” N, 70°56’26.00” W. It represents the rst re-
cord for Trujillo State.
Yaracuy State
ULABG 7845, captured in “Aroa, SW San Felipe,
10°27.304” N, 68°54.169” W, 200 m s.n.m., estado Yar-
acuy, Venezuela”. It represents the rst record for Yaracuy
State. In the same locality, another specimen was observed
but not captured, buried on the grounds of a pastureland.
It exhibited a lime-green coloration on the anks and a
contrasting dark brown pattern on the back, somewhat
tiger-striped. Neither of these specimens was photo-
graphed alive, and for ULABG 7845, only the dry skull
is preserved.
Examination of the skull (Fig. 3) reveals that it belongs
to the family Ceratophryidae, genus Ceratophrys, as con-
rmed by the presence of non-pedicellate teeth on max-
illa and premaxilla and the architecture of the bony shield
(cf. Lynch 1982, Wild 1997). is identication is further
supported by the robustness and hyper-ossication of the
cranium, the presence of a massive postorbital arch, and
the dorsal orientation of the orbits –traits that clearly dis-
tinguish it from the sympatric bufonids described by Men-
doza (2011). Given its geographical origin, the specimen
is specically assigned to Ceratophrys calcarata, as it is the
only species of the genus identied to date in Venezuela
north of the Orinoco River.
Zulia State
MBUCV 508, from “El Laberinto, Río Palmar, Zu-
lia”; MBUCV 540, 6655-6663, from “Maracaibo, Zulia”;
MBLUZ (Museo de Biología de la Universidad del Zulia,
Maracaibo, Venezuela) 34, from “carretera vía Tulé, edo.
Zulia”; MBLUZ 36, from “Canchancha I, norte de Mara-
caibo, edo. Zulia, Venezuela”; MBLUZ 80, from “El Mo-
ján, edo. Zulia”; and MBLUZ 209, from “Agropecuaria
San Sebastián, municipio Machiques de Perijá, 9º52’3.50”
N, 72º23’58.80” O, 100 m s.n.m.”; represent new intra-
state records.
e rst museum record for Ceratophrys calcarata in
Zulia State consisted of the tadpoles (KU -University of
Kansas- 193492, 193498-193499) from ‘NW Maracaibo’
reported by La Marca (1986). is locality can now be
more accurately dened thanks to a post-metamorphic
specimen (ULABG 425), captured alongside these tad-
poles by the same collectors at a “pond NW of Maracaibo,
road near Don Bosco, Zulia State”. Infante-Rivero (2009
provided the second museum record (MBLUZ 105) giv-
ing “Cojoro” as the place of collection, which, according to
catalog data should specically be “La Guajira a 10 km de
Cojoro, municipio Páez, estado Zulia”. e closest Cera-
tophrys calcarata populations to this Venezuelan Guajira
record are those from the Peninsula of La Guajira in Co-
lombia (Acosta Galvis 2023, and Fig. 4 herein). Infante-
Rivero and Velozo (2015: 143) listed Ceratophrys calcara-
ta, without indicating voucher specimens, from a locality
Figure 2. Ceratophrys calcarata live adult (top), not collected,
and its habitat (bottom) at an articial lake in Sabana de Men-
doza, Trujillo state, Venezuela. Photos: Santos Bazó.
La Marca et al.
44
Figure 3. Dorsal (top) and ventral (bottom) views of cranium of Ceratophrys calcarata, museum specimen ULABG 7845, from Aroa,
Falcon state. Photos: Oscar Mendoza.
Ceratophrys calcarata in Venezuela
45
Figure 4. Individuals (not collected) of Ceratophrys calcarata from Peninsula de La Guajira in Colombia. Photos: Ricardo Pineda.
La Marca et al.
46
near Machiques de Perijá; one of them most probably cor-
respond to MBLUZ 209.
For the rst time we have a more precise view of the
geographic distribution of Ceratophrys calcarata in Ven-
ezuela, north of the Orinoco River (Fig. 5). More studies
are needed to obtain a more complete picture. Records
south of the Orinoco River (not shown in Fig. 5, but see
map 42 in La Marca 1992: 101) are still limited to the city
of Puerto Ayacucho and vicinities, in the Venezuelan Am-
azonas State.
ACKNOWLEDGMENTS
We are very grateful to Carmen Alicia Montilla, Miguel
David Montilla, José Danilo Ruz, Antonio José Fernán-
dez, F. Aroldo Rafael Valbuena, and Carlos Rosario for
their eld assistance and logistical support leading to the
capture of the voucher specimen for Ceratophrys calcarata
in Trujillo State. Hedelvy Guada and Leida Valero facili-
tated access to MBUCV and CVULA, respectively, for
the study of the Ceratophrys specimens deposited in those
collections. Gilson Rivas and Tito Barros provided info
on relevant Zulia records and museum specimens depos-
ited in MBLUZ. Gilson Rivas kindly brought to our at-
tention two pertinent papers citing Zulia records that we
were previously unaware of. We thank Alan Markezich
and Ricardo Javier Pineda Camargo for kindly providing
photographs of live Ceratophrys calcarata specimens used
in Fig. 1C and Fig. 4, respectively. Oscar Mendoza kindly
provided photographs of the skull of Ceratophrys calcara-
ta depicted in Fig. 3. e manuscript beneted from the
comments of two anonymous reviewers, to which we are
most thankful.
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Figure 5. Locality records of Ceratophrys calcarata north of the Orinoco River. Venezuelan records south of the Orinoco River (not
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Ceratophrys calcarata in Venezuela
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