La Marca
18
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.21083898 / Anartia, 42 (junio 2026): 18-39
http://zoobank.org/urn:lsid:zoobank.org:pub:FCF91A98-AC17-4E7D-A00E-75611D686270
A new Andean rocket frog (Amphibia: Anura: Leptodactylidae:
Leptodactylus) from urban spaces at Mérida, Venezuela
Una nueva rana andina (Amphibia: Anura: Leptodactylidae: Leptodactylus)
de espacios urbanos en Mérida, Venezuela
Enrique La Marca
Facultad de Ciencias Forestales y Ambientales (Escuela de Geografía), Universidad de los Andes, Mérida 5101, Venezuela;
and Fundación Biogeos para el estudio de la diversidad biológica, Mérida, Venezuela. orcid.org/0000-0002-6508-3375
Correspondence: enrique.lamarca@gmail.com
(Received: 23-01-2026 / Accepted: 10-06-2026 / On line: 30-06-2026)
ABSTRACT
A new species of Leptodactylus is described from the city of Mérida, in the Andean mountains of Venezuela. e new
taxon, a member of the Leptodactylus melanonotus species group, is distinguished from all congeners by a combination
of morphological and bioacoustic characters, including: moderate body size (adult males mean SVL 37.4 mm; female
44.5mm); a single pair of irregular, oen interrupted dorsolateral folds; slightly expanded (knob-shaped) toe tips; and a
ventral color pattern with pale mottling fading posteriorly. e new species is further characterized by an advertisement
call with an exceptionally high note repetition rate (~240 calls/min) and a high dominant frequency (2.4 kHz). We pro-
vide a detailed description of the holotype and variation within the type series, including data on coloration in life and
preservative and description of the larval stage. e species is restricted to internal valleys within the Cordillera de Mérida,
where it dwells in urban and peri-urban environments. We provide preliminary data on its ecology and conservation status.
Key words: Andes, Anura, conservation, Leptodactylus, natural history, taxonomy, Venezuela.
RESUMEN
Se describe una nueva especie de Leptodactylus de la ciudad de Mérida, en los Andes de Venezuela. El nuevo taxón, miem-
bro del grupo de especies Leptodactylus melanonotus, se distingue de todos sus congéneres por una combinación de carac-
teres morfológicos y bioacústicos, que incluyen: tamaño corporal moderado (longitud hocico-cloaca, LHC, promedio
en machos adultos de 37,4 mm; hembra de 44,5 mm); un único par de pliegues dorsolaterales irregulares y a menudo
interrumpidos; puntas de los dedos de los pies ligeramente expandidas (en forma de pomo o botón); y un patrón de colo-
ración ventral con un moteado pálido que se desvanece hacia la parte posterior. La nueva especie se caracteriza además por
un canto de advertencia con una tasa de repetición de notas excepcionalmente alta (~240 cantos/min) y una frecuencia
dominante elevada (2,4 kHz). Proporcionamos una descripción detallada del holotipo y de la variación dentro de la serie
tipo, incluyendo datos sobre la coloración en vida y en preservativo, y descripción de la etapa larvaria. La especie está res-
tringida a los valles internos de la Cordillera de Mérida, donde habita en entornos urbanos y periurbanos. Proveemos datos
preliminares sobre su ecología y estado de conservación.
Palabras clave: Andes, Anura, conservación, historia natural, Leptodactylus, taxonomía, Venezuela.
A new Andean Leptodactylus
19
INTRODUCTION
e biodiversity of the Venezuelan Andes is currently
in a prolic stage of discovery and inventory. In the rst
decade of the 21st century alone, numerous new species
were described from the Cordillera de Mérida, a moun-
tain range approximately 400 km long and 80 km wide
in northern South America. Most occurred in relatively
undisturbed natural habitats. However, a distinct and
yet undescribed taxon persists within the urban spaces of
Mérida, one of the largest and highest cities in the Ven-
ezuelan mountains. e genus Leptodactylus is a diverse
group of Neotropical frogs, currently organized into sev-
eral major species groups. e Leptodactylus melanonotus
group is characterized by having toes with lateral fringes
and, at most, a single pair of dorsolateral folds. In Ven-
ezuela, this group includes several recognized taxa, e. g.,
L. colombiensis, L. diedrus, L. leptodactyloides, L. magis-
tris, L. petersi, and L. sabanensis. Populations from the
Venezuelan Andes were oen referred to as a broad “Op-
erational Taxonomic Unit” (OTU); however, researchers
noted that this grouping might be too conservative and
likely contained multiple species. Previous literature has
occasionally mentioned the Andean populations under
various names, including L. podicipinus petersi, L. wag-
neri, and the nomen nudum “Leptodactylus meridensis”.
However, detailed morphological and bioacoustic analy-
ses now conrm that the populations inhabiting the inter-
nal valleys of the Cordillera de Mérida represent a distinct
evolutionary lineage. is new species is most similar to
L. colombiensis but can be readily distinguished by: 1. its
smaller size, distinct bioacoustic prole with a remarkably
fast call repetition rate and 2. specic coloration patterns,
i. e., greenish tinge in the groin. Herein we describe the
new species, provide a comprehensive diagnosis against
its congeners in the L. melanonotus group, and document
its natural history and conservation requirements within
the increasingly urbanized landscape of the Mérida ter-
race.
MATERIALS AND METHODS
Specimen collection and curation
All specimens examined in this study are deposited in
the Collection of Amphibians and Reptiles of the Labo-
ratorio de Biogeografía at the Universidad de Los Andes,
Mérida, Venezuela (ULABG). A total of 10 specimens
were collected for the species description; additional in-
dividuals were examined from existing holdings. Paratypes
consist exclusively of topotypes, being specimens collected
from the same locality as the holotype. Specimens consid-
ered to belong to the new species, but excluded from the
type series, are listed as additional specimens. e synony-
my lists provided for the new taxon follow the concept of
chresonymy (Smith & Smith 1972, La Marca 1994), i. e,
the usage of names in previous literature. is is intended
to clarify the taxonomic history of the populations from
the Mérida terrace, which have been historically assigned
to other taxa or mentioned as nomen nudum.
Morphometrics and character analysis
Measurements were taken with a digital caliper (He-
lios®, precision 0.01 mm). For the post metamorphic speci-
mens, the following morphometric variables were record-
ed: snout-vent length (SVL); head length (HL) from the
posterior corner of the mouth to the tip of the snout; head
width (HW) at the angle of the jaws; eye-to-naris distance
(EN) from the anterior corner of the eye to the center of
the naris; internarial distance (IN); eye length (EYE) from
the anterior to the posterior corner; horizontal diameter of
the tympanum (T); hand length (HAND) from the prox-
imal edge of the palmar tubercle to the tip of nger III;
tibia length (TL) from the outer edge of the exed knee
to the heel; and foot length (FOOT) from the proximal
edge of the outer metatarsal tubercle to the tip of toe IV.
Adult status was conrmed via gonad inspection. Mature
males were identied by the presence of open vocal slits in
the oor of the mouth, a pair of black keratinized thumb
spines, and enlarged testes. Mature females were identied
by the presence of pigmented eggs and thickened, convo-
luted oviducts. Terminology and descriptive methods fol-
low Heyer (1994) and La Marca et al. (2004).
Larval morphology
Tadpole developmental stages were determined fol-
lowing Gosner (1960), and morphological terminology
follows Mijares-Urrutia (1998). Morphometric data for
larvae included: body length (BL), tail length (TL), body
height (BH), eye-to-nostril distance (END), eye-to-snout
distance (ESD), eye diameter (ED), spiracle-to-snout dis-
tance (SSD), spiracle-to-dorsum distance (SDD), caudal
musculature height (CMH), dorsal n height (DFH),
ventral n height (VFH), anterior diastema width
(ADW), posterior diastema width (PDW), oral disc
width (ODW), body width (BW), inter-orbital distance
(IOD), and inter-nostril distance (IND).
Bioacoustics
Advertisement calls were recorded in the eld using a
Sony TCS-310 tape recorder with an integrated micro-
phone. Bioacoustic analyses were performed using Aviso-
Sonograph Pro (Windows), a PC-based spectrograph-
generating program. Spectrograms display frequency
La Marca
20
(kHz) on the vertical axis and time (s) on the horizontal
axis. Relative amplitude (dB) is represented by a grayscale
gradient, ranging from low (white) to high (black).\
DESCRIPTION OF SPECIES
Leptodactylus trinacria sp. nov.
(Figs. 1-2)
http://zoobank.org/urn:lsid:zoobank.org:act:49A4DFC5-FA66-48EF-A58A-
A88A12B5CC6F
Common names
Mérida’s Whistling Frog (La Marca 2016a, La Marca
2020), Sapito Silbador de Mérida (La Marca, 2015), Rana
Silbadora de Mérida (La Marca 2016b).
Chresonomy
Leptodactylus podicipinus petersi. Rivero, 1961: 47
(partim).
Leptodactylus wagneri. Heyer, 1970: 45 (partim); La
Marca, 1992: 63,107 (partim).
?Leptodactylus fuscus. Barrio-Amorós, 1998: 43
(partim).
Leptodactylus pallidirostris. Barrio-Amorós, 1998:
46 (partim).
Leptodactylus sp. 1. La Marca, 1999: 203; Barrio-
Amorós, 2004: 98,170 (partim).
Leptodactylus sp. García, Albornoz & La Marca,
2007: 65; La Marca, 2015: 217; La Marca et al.,
2015a, b: 10; La Marca, 2016a: 3, b: 7; La Marca,
2017a, b: 5; La Marca & Castellanos, 2018a, b:
17; La Marca, 2020a: [2]; La Marca, 2021: 31.
“Leptodactylus meridensis” (nomen nudum) – Karls-
dóttir, 2017a, b: 16; La Marca, 2022a, b: 7.
Holotype
ULABG 4873, adult male, with both vocal slits opened,
a pair of keratinized black thumb spines on each hand and
dark throat. Collected from the articial lake at Museo
de Ciencia y Tecnología of Mérida city (former “Central
Azucarero Los Andes”, CALA), Mérida state, Venezuela,
1320 m.asl, geographic cordinates WGS84: 8°33’44”N,
71°11’25”O (Fig. 1 A, B), collected by Enrique La Marca,
Diego Cadenas, Luis Felipe Esqueda, Enzo La Marca and
Francisco Nava, on 19 March 2002.
Paratypes (Topotypes)
ULABG 4871-4872, 4874-4879, adult males (vocal
slits opened, keratinized black spines, dark throat), col-
lected at the same locality of the holotype, but captured on
30 October 2001; ULABG 4880 (allotype, Fig. 1 C,D),
adult female (convoluted oviduct, large eggs up to 4.5
mm), same data as holotype.
Additional specimens
ULABG 1413, adult female, coming from Los Llani-
tos de Tabay, nr. Tabay, NE city of Mérida, estado Mérida,
Venezuela. 1475 m.asl, 8°37’14”N, 71°6’14”W, collected
on 28 marzo 1985 by Enrique La Marca.
ULABG 3061-3062, adult males from the Instituto
de Geografía y Conservación de Recursos Naturales
(IGCRN), Universidad de Los Andes (Mérida, Venezu-
ela), 8°37’20”N, 71°8’20”W collected on 5 November
1991 by Hans Peter Reinthaler.
ULABG 3733, adult female, coming from Ejido, estado
Mérida, Venezuela. 1100 m. asl., 8°33’30”N, 71°15’9”W,
collected on 27 February 1994 by María José Praderio and
Carlos Gottberg; ULABG 4286-4287, adult males, col-
lected by Carlos Gottberg on 12 March 1996.
ULABG 4481, 4482, lots of a single tadpole each, com-
ing from same locality as holotype and paratypes, collect-
ed on 16 September 2002 by María José Praderio.
Etymology
e specic epithet “trinacria” is a noun in apposition
derived from the Ancient Greek treis (meaning three;
Brown, 1956) and ákrai (meaning the extreme point of a
coast or territory; Rodríguez Adrados, 1980). Historically,
the term served as the classical Greek designation for the
island of Sicily, in reference to its triangular conguration.
Herein, the name alludes to the triangular dorsal blotch
located between the interorbital and scapular regions
(Fig.2), exhibited by several specimens of the new taxon,
including the holotype. While this character is not diag-
nostic for the species and may occur in other Leptodactylus
congeners, the name is formally dedicated to the author’s
father, Vincenzo La Marca Zirini, on the centenary of his
birth in Sicily (22/I/1926). is nomenclature thus estab-
lishes a symbolic link between the author’s ancestral Sicil-
ian heritage and this new Andean frog.
Diagnosis
Previous work recognized that there was a Venezuelan
Andes OTU” (Operational Taxonomic Unit) which pur-
portedly contained specimens from the Andean states of
Mérida, Táchira and Trujillo (Heyer 1994), as well as from
other western and central Venezuelan states from moun-
tains to lowlands. is Operational Taxonomic Unit was
deemed to represent a single species (“although this may
be too conservative a conclusion”; Heyer 1994:56). Aer
examination of specimens from the interior part of the An-
dean Cordillera de Mérida, we consider that a single spe-
A new Andean Leptodactylus
21
Figura 1. A, B: Dorsal and ventral views of preserved holotype (ULABG 4873) and C, D: allotype (ULABG 4880) of Leptodactylus
trinacria sp. nov. Circular section on le thigh of holotype shows place where a sample tissue was taken for Bd testing. Photos: Enrique
La Marca.
A
C
B
D
cies is involved, being well dierentiated from the already
described species. Nonetheless, future work analyzing au-
diospectrograms and DNA samples will be needed to fully
resolve the taxonomic identity of other populations previ-
ously included in Heyer’s (1994) Venezuela Andes OTU.
Leptodactylus trinacria sp. nov. is assigned to the L.
melanonotus species group due to the presence of lateral
toe fringes and a pair of dorsolateral folds (sometimes
interrupted). e new species is distinguished from its
congeners by the following combination of characters: (1)
moderate body size (mean SVL 37.4 mm in males, 44.5
mm in females); (2) a pair of irregular, frequently inter-
rupted, or inconspicuous dorsolateral folds; (3) slightly
expanded, knob-shaped toe tips; (4) a well-dened tym-
La Marca
22
panum, with a diameter approximately 2/3 of the eye di-
ameter; (5) ventral coloration with dense pale mottling on
the throat and chest, fading posteriorly toward the ingui-
nal region; (6)presence of a greenish tinge in the groin
and on the hidden surfaces of the thighs in life; and (7) an
advertisement call characterized by an exceptionally high
note repetition rate (~240 notes/min) and a dominant
frequency of 2.4 kHz.
Leptodactylus trinacria sp. nov. is a moderate-size spe-
cies associated with internal valleys within the Andean
Cordillera de Mérida, in western Venezuela. As all species
in L. melanonotus species group, L. trinacria sp. nov. spec-
imens have a single pair of dorsolateral folds. Dorsolateral
folds tend to have a dierent degree of expression (Fig. 3),
given by certain preserved conditions: poorly preserved
specimens do not exhibit the folds that are distinctive in
live specimens. Live specimens variably show the condi-
tions of absent (Fig. 3B, E, F), indistinct or fragmented
(Fig. 3A, C, D), and complete and moderate to long
(Fig.3G, H; 4); although the most frequent is indistinct
or fragmented (Carvahlo et al. 2022).
Comparisons with other species
Specimens of the new species exhibits an upper lip
stripe condition (following Heyer, 1994 and Carvahlo et
al. 2022) type “state B”, dened as extending from the pos-
terior corner of the eye towards the base of the arm (Fig. 4);
however, sometimes not easily discernible. is condition
is also exhibited by L. colombiensis and L. magistris. e
toe tips of the new species exhibit the “slightly expanded”
condition “E” of Carvahlo et al. (2022), also dened as
“knob-shaped swollen” (Fig. 5). e ventral color pattern
follows condition “A” of Carvahlo et al. (2022), where the
pale mottling is fading posteriorly (Fig. 6). e lower jaw
may be covered with pale spots in both males and females
(Fig. 6A, B). e ventral mottling concentrating towards
the anterior central parts and fading progressively towards
lower venter is also exhibited by Leptodactylus colombien-
sis and probably L. magistris (Carvahlo et al. 2022). e
groin can be greenish in some specimens, easily dierenti-
ating the species from others in the L. melanonotus group
that show a bright yellow coloration. e greenish tinge
may be also present on concealed anterior and posterior
parts of the thighs, and ventrolaterally at the posterior part
of ank (Fig. 7).
Venezuelan species of Leptodactylus with toe fringes
include members of the L. melanonotus and L. latrans
groups of species. Leptodactylus latrans have at least four
well-developed dorsolateral folds. All members of the L.
melanonotus group have a single pair of dorsolateral folds,
Figure 2. Dorsal view of a live Leptodactylus trinacria sp. nov. to show the triangular-shaped blotch between the interorbital and
scapular regions. Photo: Enrique La Marca.
A new Andean Leptodactylus
23
Figure 3. Live specimens, not collected, of Leptodactylus trinacria sp. nov. from the terrace of Mérida. Photos A-G: Enrique La Marca,
Photo H: James R. Dixon.
A
C
B
D
E
G
F
H
La Marca
24
istic of L. colombiensis and exhibits a signicantly higher
dominant frequency (2.4 kHz vs. 1.47–1.98 kHz). e
note repetition rate of the new species (~240 calls/min) is
nearly seven times faster than that of L. colombiensis (~36
calls/min) and remains one of the highest recorded for any
species within the L. melanonotus group (see Carvalho et
al. 2022). is accelerated repetition rate remains an ex-
ceptional diagnostic feature within the L. melanonotus
group.
Comparisons with other Leptodactylus species having
toe fringes (except for L. latrans and its closest relatives):
L. bolivianus Boulenger, 1898, is a larger species (SVL:
females to 88 mm, males to 94 mm) and the dorsolateral
folds are smooth, regular, and long. In the larger (SVL:
females to 81 mm, males to 63 mm) L. riveroi Heyer &
Pyburn, 1983), the entire upper lip and loreal region oen
have a broad and well-dened pale stripe. Leptodactylus
diedrus lacks dorsolateral folds and the toe tips are usu-
ally expanded into small discs. e ventral and posterior
thigh patterns do not merge but rather are in close con-
tact. Leptodactylus griseigularis (Henle, 1981) is a slightly
larger species (SVL: females 39-58 mm, males 35-51 mm)
with indistinct pale lip stripes, and presence of large black
thumb spines. Leptodactylus leptodactyloides specimens
commonly have a thigh pattern with distinct pale stripes,
and few specimens (10%) also have pale-spotted chins.
Almost all specimens have melanophores on the belly re-
sulting in a nely mottled pattern. Leptodactylus natalensis
Lutz, 1930 occurs along coastal Brazil from the State of
Rio Grande do Norte to the State of Rio de Janeiro; over
half of the specimens of this species have toe tips larger
than the narrow or just-swollen categories. e light pos-
terior stripe is just distinct in some (9%) specimens. Lep-
todactylus nesiotus Heyer, 1994 from Trinidad is a smaller
species (males 32-33 mm SVL) and has a broad pale stripe
on the entire upper lip or at least to under the eye. Lep-
todactylus magistris, from a relatively isolated mountain
range in Falcón State, Venezuela, shows keels on ngers
II and III, a pale white line on lips with well-dened bor-
ders and a weak tarsal fold. Leptodactylus pallidirostris
Lutz, 1930, from the Guianan shield region, is a smaller
species (females 30-43 mm SVL, males 28-37 mm SVL).
Belly bearing a ne mottling to dark blotches. Toe-tips ex-
panded or with small discs. Leptodactylus pascoensis, from
Amazonian anks of the Andes in central Peru, is a larger
species (SVL: females 52-67 mm, males 60-61 mm). Poste-
rior pale lip stripes absent. Toe tips swollen or just swollen.
Males with two large black thumb spines. Leptodactylus pe-
tersi (Steindachner, 1864), from greater Amazonia and the
Guiana shield region, including Venezuela, has a belly pat-
tern usually extensively mottled in an anastomotic pattern.
Figure 4. Live specimen of Leptodactylus trinacria sp. nov. ex-
hibiting a pale upper lip stripe extending from the posterior cor-
ner of the eye to base of arm. Photo by Christian Reymondin.
sometimes poorly developed. Other Venezuelan Lepto-
dactylus with lateral toe fringes are L. colombiensis Heyer,
1994; L. diedrus Heyer, 1994; L. leptodactyloides (Anders-
son, 1945); L. magistris Mijares-Urrutia, 1997; L. pascoen-
sis Heyer, 1994; L. riveroi Heyer & Pyburn, 1983; L. saba-
nensis Heyer, 1994; and L. validus Garman, 1888.
Leptodactylus trinacria sp. nov. most closely resembles
L. colombiensis, which also has dorsolateral folds that are
irregular and oen interrupted. Leptodactylus colombien-
sis is a large-size species (SVL: females 38-62 mm, males
to 36-56 mm) in which lip stripes, if present, extend only
from the mid-eye level posteriorly. Most L. colombiensis
individuals have some belly mottling (almost immacu-
late posterior belly in L. trinacria sp. nov.) and swollen
toe tips (toe tips little or no expanded). igh length in
L. trinacria sp. nov. is larger than that of L. colombiensis:
males of L. trinacria sp. nov. with a mean value of 48.8%
SVL (range 47%-51%), while L. colombiensis has a mean
value of 43 ±3% (range 37%-50%); the single female of
L. trinacria sp. nov. has a shank length of 49% SVL, while
that of females of L. colombiensis is 43 ± 3% SVL (range
37%-50%).
e advertisement call of new species has a higher dom-
inant frequency (2.4 kHz vs. <2.0 kHz) and a faster call-
ing rate (see below). e new species diers in bioacoustics
from Leptodactylus colombiensis by having a higher domi-
nant frequency (2.4 kHz vs. 1.47–1.98 kHz in L. colom-
biensis), lacks the low-frequency initial pulse character-
A new Andean Leptodactylus
25
Figure 5. A-F. Specimens of Leptodactylus trinacria sp. nov. to show the slightly expanded or knob-shaped swollen toe tips. Photos:
Enrique La Marca.
A B
Toe tips usually just swollen (62%), some with swollen or
just-expanded toe tips (34%). Posterior lip stripe usually
(44%) indistinct to not discernible (43%). Leptodactylus
podicipinus (Cope, 1862) occurs in Amazonia in Brasil,
Bolivia, Paraguay and northeastern Argentina. It oen has
distinct pale belly spots. Leptodactylus sabanensis, from La
Gran Sabana, Venezuela has the most common pale lip
stripe condition indiscernible, and, when discernible, they
extend from the posterior corner of eye. Leptodactylus
validus occurs in Guiana shield region, Trinidad, Tobago,
and Lesser Antilles. In those individuals with discernible
lip stripes, the stripes extend from the posterior corner of
C D
E F
La Marca
26
er to the tip of snout than to the anterior border of eye,
about 62% of the eye-to-nares distance. Horizontal length
of eye approximately 1.3 times the eye diameter and 1.1
times eye-to-nares distance. Horizontal length of eye
3times larger than eye-to-tympanum distance. Internarial
distance slightly shorter (0.9 times) than the eye-to-nares
distance. Tympanum rounded, distinct, with a conspicu-
ous tympanic annulus. Tympanum diameter 2.4 times
larger than eye-to-tympanum distance. Supra-tympanic
dermal fold distinct, covering the posterior part of eye to
arm insertion. e post commissural gland is about twice
as long as wide, single on le side and divided by a deep
trench on the right side. Vocal sac single, internal, no ex-
ternal modication visible. Vocal slits well-developed, on
posterior part of mouth, about 0.63 times shorter than
the tongue length. Tongue ¾ larger than wide. Tongue
sub-ovoid, about 30% adherent to oor of mouth, with
posterior border round and slightly notched. Choanae
sub-triangular, visible, not concealed by palatal shelf of
maxillary arch; widely separated between them (about 6
times the width of a choana). Vomerine teeth in two long
and almost straight series posterior to and almost entirely
between choanae; odontophore processes elevated, slight-
ly sub-triangular, approximately same width as length,
separated between them about 1/3 of its width. Maxilla
and pre-maxilla toothed; small teeth.
Dorsum smooth, bearing pale pointed tubercles of
various size; some tubercles faintly keratinized and more
abundant towards the posterior region, particularly over
the urostyle region. Dorsal folds absent. Skin of chest, ven-
ter and throat, smooth. Skin of venter, chest and throat,
smooth. Chest spines absent. Ventral disc fold distinct.
Arms not hypertrophied. Elbow and lateral posterior part
of forearm with minute keratinized spicules. Ventral, ex-
ternal and lateral surfaces of forearm with small, pointed
tubercles weakly keratinized. Ulnar ridge absent. enar
tubercle conspicuous, oval, and approximately twice larger
than wide. Palmar tubercle lobate, low-elevated and less
prominent than thenar one. Supernumerary tubercles ab-
sent. Sub-articular tubercles elevated and rounded. Finger
tips not expanded. Fingers without webbing. Keratin-
ized spicules on the internal lateral surfaces of ngers II
and III. Ridges and keels absent on lateral sides of ngers.
First nger slightly larger than second. Finger lengths
II=IV<I<III. First nger with two black nuptial conical
spines; distal spine broader than proximal one, both rela-
tively moderate in size and about 0.8 mm in length.
Cloacal opening well above the mid-line of thighs,
covered by a short and crenulate dermal fold with shallow
striae, giving a radiated appearance to the cloacal region.
Dorsal skin on thighs and tarsi with numerous low-kera-
Figura 6. Ventral color pattern of (A) female and (B) male of
Leptodactylus trinacria sp. nov. to show the pale ventral mot-
tling that is fading posteriorly. Note that the lower jaw is covered
with pale spots. Photos: Enrique La Marca.
A
B
the eye posteriorly. Leptodactylus wagneri (Peters, 1862), is
a larger species (males 39-61 mm SVL, females 52-82 mm
SVL) with long dorsolateral folds, and has moderately to
extensively mottled bellies. In L. trinacria sp. nov. all dis-
cernible lip stripes extend from the posterior corner of the
eye and the belly pattern is moderately mottled.
Description of holotype
Snout sub-ovoid in dorsal view; tip of snout rounded in
dorsal and lateral views. Head 0.1 times wider than long.
Inter-orbital space relatively smooth, without tubercles.
Interorbital distance 0.6 times larger than upper eyelid
width. Canthus rostralis straight, not well-dened. Loreal
region slightly concave, descending smoothly to lips. Lips
not ared. Nares elevated. Nares directed dorso-laterally
and tilted about 45 degrees back and upward. Nares clos-
A new Andean Leptodactylus
27
tinized small and pointed tubercles. ighs and shanks
ventrally smooth; tarsi and metatarsi bearing dorsal and
ventral small pointed tubercles scarcely or non-keratin-
ized. A row of weakly keratinized small pointed tubercles
from base of toe V to external border of the metatarsus
and tarsus. A relatively large postcommissural gland, 2.6
mm in length, on posterior edge of lips. Ventrolateral
gland bordering anterior 2/3 of belly. No other glands
evident. Tarsal fold conspicuous, extending 7/9 distance
of tarsus. Tarsal fold distinct, with numerous minute and
weakly keratinized spicules on and along the medial and
posterior portion of fold. Tarsal fold extending along tar-
sus, connecting with a membranous ap along the inter-
nal border of toe I. External metatarsal tubercle slightly
oval (1.25 times larger than wide), elevated and truncate
in lateral view. Internal metatarsal tubercle sub-spatulate
(narrower at proximal portion), 2.5 times larger than wide
in the middle, and about 2.5 times larger than the size of
external metatarsal tubercle. Tarsi and soles of feet with
several small, heterogeneous-size white tubercles, several
tan- tipped. Subarticular tubercles large, round to oval,
and round in lateral view. Toe-web formula: I1- ½ II1- ½
III ½ - ½ IV1- 1V. Toe tips slightly swollen. Toes bear-
ing membranous keels and well-developed lateral folds.
Tibio-tarsal articulation reaching to arm insertion when
leg adpressed forward.
Measurements of holotype (in mm)
Snout to vent length 36.2, thigh length 16.5, shank
length 17.8, head width 12.9, head length 12.0, tympa-
num diameter 2.7, eye diameter 3.4, eye to nostril distance
2.5, internarial distance 3.1, hand length 9.2, foot length
20.8.
Coloration of holotype in life
roat dark gray; venter “dirty” white with a yellow
tinge on the posterior end; groin with a yellowish-green
wash. Dorsum dark olive-brown, with cream-colored ar-
eas delimiting a dark, cross-shaped pattern. Head with an
inverted dark interorbital triangle bordered anteriorly by
a narrow cream band. Anterior concealed surfaces of the
thighs exhibit a faint reddish-orange tinge. Labial region
with alternating cream and dark brown bands. A distinct
cream-colored post-tympanic band extends from the low-
Figure 7. Female specimen of Leptodactylus trinacria sp. nov. exhibiting a greenish tinge in groin and on concealed anterior and poste-
rior parts of thighs, and ventrolaterally at the posterior part of ank. Same specimen as Figs. 5 A, B. Photo: Enrique La Marca.
La Marca
28
er posterior margin of the tympanum to the insertion of
the arm (most prominent on the right side of the head).
Coloration of holotype in ethanol 70%
Dorsum marbled; large dark brown blotches on paler
background, bordered by dark brown. Blotches more dis-
tinct posteriorly and at interorbital region. Irregular pale
interorbital band, black-bordered posteriorly. A pair of
narrow dark brown bands from the middle of upper eye-
lids converging diagonally to mid-posterior head, forming
an interrupted triangle. Upper lip with wide alternating
dark and pale brown bands, most posterior one separated
from tympanum by a very narrow dark band. Tympanum
pale brown; uppermost parts of tympanic annulus cream,
surrounded by dark brown. Tip of snout cream, dorsally
and laterally bordered by dark brown. Posterior articula-
tion of mouth has postcommissural gland cream colored.
Edge of chin dark with small irregular pale dots. roat
nearly uniformly brown; chest and belly cream, with irreg-
ular small dark brown blotches, especially on anterior part
of belly. Ventral part of forearms are dark brown, except to-
wards the anterior part near the arm insertion. Black (mel-
anized) thumb spines on right hand, dark brown on le
hand. Lower part of legs with irregular brown blotches on
a pale-cream background, resembling anterior belly pat-
tern. Palms brown. Inner metatarsal tubercle cream, bor-
dered by dark brown. Under parts of ngers pale brown,
with cream subarticular tubercles. Finger pads cream.
Palms dark brown, with pale supernumerary tubercles.
Toe pads cream to pale-brown. Metatarsal fold, toe fringes
and toe lateral aps tend to be cream to pale brown. Upper
parts of extremities, except on forearms, banded; banding
extends onto dorsal surfaces of ngers and toes. Supracloa-
cal fold cream. Posterior surfaces of thighs with irregular
pale and dark brown markings; no conspicuous pale or
cream bands on thighs. Dark brown band-like pattern
on posteroventral surfaces of thighs. Brown ventrolateral
glands. Cloacal opening surrounded by dark brown. Tes-
ticles cream.
Variation within the type series
Dorsolateral folds never form a continuous uninter-
rupted line. Folds, in type specimens, are absent in 20%,
are moderate in length in 30%, and long in 50%. Male
specimen ULABG 4876 have several characters that de-
part from the description of the holotype as follow: eye
diameter 2.5 larger that distance from eye to tympanum;
tongue 50% free from oor of mouth; choanae separated
among them by approximately three times the width of
a choana; dorsum of arm and forearm with few spicules;
arm with a single row of ve to six not keratinized spicules;
palmar tubercle slightly heart-shaped; thick lateral keel on
ngers II, III and IV, although less conspicuously on inter-
nal side of nger I; cloacal opening bordered by a striated
series of thin and shallow lines forming a radiated pattern;
tarsus and metatarsus smooth; foot web reaches the rst
tubercle on proximal side of rst toe, and the rst tubercle
on distal side of toe III; outer border of toe V with a cuta-
neous keel that extends from distal end of toe to the base
of the outer metatarsal tubercle. ULABG 4794 has odon-
tophore processes that are about twice wider than long
and a slightly conspicuous lateral ridge along the inner
side of the rst nger. In female specimen ULABG 4880
the choanae are separated between them by approximately
3 times the width of a choana. ick lateral ridges pres-
ent on all ngers. enar tubercle elongated. ick fold
from the base of the thenar tubercle, which connects with
a thick ridge that runs along the outside of the 4th nger.
Toe-web formula: I1- 1II1- ½ II1 - ½ IV1- 1V. It presents a
rounded protuberance on the back, at the level of the ante-
rior part of the sacrum (probably a parasite) embedded in
the skin (Fig. 1C), without reaching the muscles.
Morphometric variation of complete type series, including
holotype
Female 44.5 mm SVL, males 35.4-40.0 mm SVL
(x=37.4 ± 1.43); female head length 14.9 mm, male head
length 11.5-13.3 mm (x= 12.3 ± 0.6); female head width
16.2 mm, male head width 12.0-14.1 mm (x= 13.2 ± 0.6);
female tympanum diameter 3.3 mm, male tympanum di-
ameter 2.5-3.3 mm (x= 2.87 ± 0.25; female thigh length
21.8 mm, male thigh length 16.9-19.8 mm (x= 18.3 ±
1.06); female eye diameter 4.2 mm, male eye diameter
3.0-4.1 mm (x= 3.5 ± 0.3); eye-to-nostril distance in fe-
males 4.0, eye-to-nostril distance in males 2.5-3.8 mm
(x=3.3±0.4); inter-nostril distance in female 4.0, inter-
nostril distance in males 3.1-4.0 mm (x= 3.4 ± 0.3); female
hand length 11.0 mm, male hand length 8.3-10.3 mm
(x=9.24 ± 0.7); female foot length 23.3 mm, male foot
length 14.1-21.3 mm (x=20.1 ± 2.27). Measurements for
every individual of the type series is provided in Table 1,
and a summary of statistics for size and other selected mea-
surements is given in Table 2.
Color variation
Paratypes, in life, had dorsum olive brown, with ir-
regular dark brown blotches. A dark, incomplete inverted
triangular blotch, with apex directed backwards, usually
present and located between and behind eyes, always pre-
ceded by a cream inter-orbital narrow band. Lips barred
with dark and pale brown bands. Pale stripe on the poste-
rior lip region (cf. Heyer 1994: 30, Fig. 11) rather broad
A new Andean Leptodactylus
29
Table 1. Selected measurements (in mm) of the type series of Leptodactylus trinacria sp. nov. Le column indicates mu-
seum catalog number at the Collection of Amphibians and Reptiles of the Biogeography Lab at University of Los Andes in
Mérida, Venezuela. e holotype is indicated with an asterisk. All male specimens, except ULABG 4880 that is a female.
See methodology for abbreviations; additionally, “Rep. cond.” in the last column stands for the reproductive condition,
exhibited as indicated by the numbers 1 (both vocal slits opened), 2 (pair of keratinized black thumb spines), 3 (darkened
throat), 4 (gray throat), 5 (convoluted oviducts), 6 (only right vocal slit opened), 7 (gray throat with darkened borders
towards lips), 8 (corpora adiposa, “fat bodies”, present) and 9 (presence of mature eggs, 1.2 to 4.5 mm in diameter).
Museum Nr
ULABG SVL TL HW HL T EYE EN IN HAND FOOT Rep. cond.
4871 38.5 18.5 12.9 12.0 3.2 3.6 3.1 3.3 9.0 20.5 1,2,3,8
4872 37.8 19.3 13.3 13.0 2.9 3.0 2.5 3.1 9.7 20.6 1,2,3,8
4873 * 36.2 17.8 12.0 12.0 2.7 3.4 2.5 3.1 9.2 20.8 1,2,3
4874 35.8 16.9 12.8 12.0 2.8 3.5 3.3 3.1 8.4 14.1 2,3,6
4875 37.8 19.2 14.1 12.0 3.3 3.6 3.1 3.7 9.6 20.9 1,2,7
4876 35.4 16.9 13.0 12.0 2.5 3.5 3.8 4.0 8.9 20.9 1,2,3
4877 37.8 18.7 13.3 13.0 2.8 3.2 3.6 3.5 9.8 21.3 1,2,3
4878 37.5 17.5 13.3 13.0 2.9 4.1 3.3 3.5 8.3 20.8 1,2,3
4879 40.0 19.8 13.8 13.0 2.7 3.6 3.4 3.5 10.3 21.2 1,2,3,8
4880 44.5 21.8 16.2 15.0 3.3 4.2 4.0 4.0 11.0 23.3 5,9
Table 2. Summary of statistics for size and other selected measurements (in mm) of type series of Leptodactylus trinacria
sp. nov. Abbreviations as indicated in the methodology section.
Females (1) Males (9)
Value +/- SD Variation range Value ± SD Variation range
SVL 44.5 44.5 37.4 ± 1.43 35.4 – 40.0
TL 21.8 21.8 18.3 ± 1.06 16.9 – 19.8
HW 16.2 16.2 13.2 ± 0.6 12.0 – 14.1
HL 14.9 14.9 12.3 ± 0.6 11.5 – 13.3
T 3.3 3.3 2.87 ± 0.25 2.5 – 3.3
EYE 4.2 4.2 3.5 ± 0.3 3.0 – 4.1
EN 4.0 4.0 3.3 ± 0.4 2.5 – 3.8
IN 4.0 4.0 3.4 ± 0.3 3.1 – 4.0
HAND 11.0 11.0 9.24 ± 0.66 8.3 – 10.3
FOOT 23.3 23.3 20.1 ± 2.27 14.1 – 21.3
and extends from just past mid-eye in 100% of type
specimens, although it is not clearly dened on le side
of head of ULABG 4872. In some very dark specimens
from the Mérida terrace, not collected, the pale posterior
lip stripe was almost undistinguishable in 6 out of 10 ex-
amined specimens. White dots on the edge of the lower
lips. Iris golden to bronze, with brown reticulation, most
pronounced on the lower half. Olive-brown golden, with
irregular dark brown markings. Very dark throat in males.
Pale dots on a dark chin usually present. Groin with a
slight yellow tinge. Iris golden to bronze, with a brown
reticule more pronounced on inferior half (e. g., Fig. 3G).
Posterior extremities dorsally brown with dark brown
bands, especially noticeable on thighs; legs ventrally mar-
moreal. Posterior surface of thighs with a yellow-greenish
tinge sometimes bearing a yellow-green line. Although
males have usually a darker throat coloration, it is not a
useful character to distinguish between males and females
La Marca
30
of Leptodactylus trinacria sp. nov. Some adult males in-
deed have paler throats than adult females in some of the
additional specimens examined. In 80% of the type series
specimens there is a noticeable dark blotch, resembling
an incomplete inverted triangle between and posterior to
eyes, more noticeable when looking specimens submersed
in liquid. In darker specimens, this blotch tends to merge
with the background color. Posterior limbs barred and up-
per parts of thighs, shanks and tarsi barred with alternat-
ing pale- and dark-brown bars.
Tadpoles
Body elongated and oval in dorsal view, depressed
(wider than deep); widest at the beginning of 1/3 of body
length and in the last 1/3 length body. Snout broadly
rounded in dorsal view. Eyes directed dorsolaterally.
Nostrils located dorsolaterally and directed anterolater-
ally, slightly closer to tip-of-snout than to eye and about
32% distance from tip-of-snout to anterior border of eye.
Nostril-openings are rounded. Edges of nostrils with a
ring. Interorbital distance 22% of the internarial distance.
Chondrocranial elements not visible through skin of head.
Spiracle is below the middle line on le side of body, evi-
dent and forming a short tube almost completely attached
to skin of body. Spiracle opening located at about 56% of
body length from tip-of-snout. Cloacal tube short, me-
diomarginal and with median opening. Dorsal n arises
at body-tail junction. Dorsal n equal in depth to the ven-
tral n at mid length of tail. Length of caudal muscula-
ture equal to total tail-length. Caudal musculature strong,
with well-developed myomeres. e longitudinal tail axis
straight. Tip-of-tail rounded. Posterior legs intra- parallel
with respect to cloacal tube. Posterior legs without dermal
folds and toes distinct (Fig. 8A-C).
Coloration in preservative. Dorsum is brown. Upper
surface of anks same as dorsum; Venter pale brown with
numerous melanophores. Tail ns pale brown (numerous
minute melanophores when viewed in high detail). Tail
musculature cream, with skin bearing minute melano-
phores, same as ns. Legs brown; nger and toes with alter-
nated cream and browns bands. Lateral line system arising
on cheeks, continuing along the inferior border of eye and
along dorsolateral part of body (with a medial inexion to-
wards internal part of back. ere is a similar parallel line
below the later. Both lines continue over the tail as a series
of small white dots to middle part of tail (Fig. 8A-C).
Coloration in life. Pale brown with a greenish tinge,
with numerous little dark spots. Dark markings tend to
coalesce on top of dorsal n and along the medial lateral
part from the middle to posterior portion of tail; iris pale
gold (Fig. 8D).
Oral apparatus. Oral disc with keratinized parts, rela-
tively small, directed anteroventrally at tip-of-snout. Oral
apparatus medium-sized in relation to body width. Oral
disc trans-angular (i. e., without lateral inexions). A sin-
gle row of small and blunt alternating papillae, separated
by a wide rostral diastema on anterior margin of the oral
apparatus. Posterior border of the oral apparatus with pa-
pillae in a mixtured conguration. Beaks (“queratostoma”
of Mijares-Urrutia 1998) partially keratinized. Upper beak
broadly arched, bearing narrow lateral extensions; lower
beak broadly V-shape. Acute serrations on both beaks;
serrations on lower beak stronger than serrations on up-
per beak. Two upper and three lower rows of keratodonts
(“denticles”, tooth row formula 2/3); upper rows equal in
length to lower rows; upper rows complete; rst lower
with an inconspicuous medial gap; second and third lower
rows entire. Slightly larger keratodonts on second upper
row. Keratodonts on all rows well keratinized (Fig. 9).
Measurements of tadpoles (in mm)
e following data applies to tadpole ULABG 4881, in
stage 40 of Gosner:
BL= 1.7; TL= 10.5; BH= 5.0; END= 1.6; ESD= 2.6;
ED= 1.1; SSD= 6.9; SDD= 4.0; CMH= 2.2; DFH= 1.4;
FH= 1.2; ODW= 0.6; BW= 6.9; IOD= 2.3; IND= 1.7.
Data for metamorphosing froglet ULABG 4882, in
stage 44 of Gosner, as follows:
SVL= 9.1; TL= 2.5; IN=1.1; Tail bud= 2.7; END=
1.1; EYE= 0.7; IOD= 1.5.
Bioacoustics
Advertisement call of the new species (Fig. 10) is
characterized by short, impulsive notes exhibiting steep
upward frequency modulation. Spectrographic analy-
sis reveals that the primary energy of the call is concen-
trated between approximately 1.8 kHz and 3.0 kHz. e
dominant frequency is centered at 2.4 kHz, appearing as
a sharp, near-vertical trace on the spectrogram: a visual
result of its abrupt rise time and remarkably short dura-
tion (30–50 ms). e calling rate is exceptionally high,
with approximately 4 calls per second (240 calls/min).
e oscillogram (Fig. 10, bottom) displays pressure uc-
tuations with high-intensity peaks corresponding to each
impulsive note. When cross-referenced with the values
in previous work (Table 2, Carvalho et al. 2022), several
signicant details emerge. With a dominant frequency at
2,400 Hz, the new species places at the high end of the
range for L. melanonotus (2,132–3,141 Hz) and L. podici-
pinus (2,218–3,258 Hz) and it is notably higher than the
L. wagneri recorded in the table (1,572–1,898 Hz) and in
L. colombiensis values stated as 1,470–1,980 Hz (de Sá et
A new Andean Leptodactylus
31
Figure 8. Tadpole of Leptodactylus trinacria sp. nov. ULABG 4481. A: dorsal view, B: lateral view, C: ventral view, D: free-living larva,
coloration in life. Photos: Enrique La Marca and Michelle Castellanos.
AA
BB
CC
D
La Marca
32
Figure 9. Mouth parts of tadpole ULABG 4481. From a pencil drawing by Enrique La Marca, with right part mirrored from le part
due to slight deterioration of the later in the larval sample.
Figure 10. Audioespectrogram (above) and oscillogram (below) of an advertisement call of Leptodactylus trinacria sp. nov. from the
type locality, recorded synchronously with collection of the type series. No time nor temperature registered.
A new Andean Leptodactylus
33
al. 2014). Regarding the note repetition rate, the Mérida
rate of ~240 calls/minute is exceptionally high. Only L.
brevipes (150–330/min), L. nesiotus (264–288/min), and
L. natalensis (163–296/min) reach such speeds; most oth-
er species in the group, including L. wagneri (50–55/min)
and L. colombiensis (~36/min), are signicantly slower. As
per the note duration, the 30–50 ms in duration for the
Mérida frog is a perfect match for the “short-note” species
such as L. petersii call type 1 (31–50 ms), L. melanonotus
(28–48 ms), and L. sabanensis (29–45 ms).
Distribution
Populations of Leptodactylus trinacria sp. nov. have
been found at several places in internal valleys of the Cor-
dillera de Mérida close to the city of Mérida. Populations
are known in inter-Andean valleys between the towns of
Tabay and Ejido, along the mid valley of the Chama River,
all within the metropolitan area of Mérida city. All these
environments fall within the Life Zone of Premontane
moist forest in the Holdridge’s system (Ewel et al. 1976),
mainly dominated by seasonal semi deciduous forests,
mostly intervened by human activities (Fig. 11). All sur-
rounding environments are high elevation ecosystems
dominated by cloud forests and paramo environments or
lower ones dominated by semiarid vegetation and tropical
lowland humid forests.
ere is a geographic distribution report for a Lepto-
dactylus stated to come from “Amazonas: Río Pescado (=
Sabana Grande), 100 m” (Heyer 1994: 114) that is obvi-
ously erroneous. e specimen, ULABG 1413, actually
comes from the vicinities of Mérida city (see “Additional
specimens” herein), and represents a Leptodactylus trina-
cria sp. nov.. It was listed within the “Venezuelan Andes
OTU”, along with specimens from the states of Aragua,
Barinas, Distrito Federal, Falcón, Guárico, Miranda,
Táchira and Trujillo (Heyer 1994: 114). is grouping of
what could actually be dierent evolutionary units may ac-
count for the resulting composite nature inferred for this
taxonomic unit. For example, specimen UMMZ 216804,
coming from Cerro Socopó (sic: Socopo), 1,260 m, most
likely correspond to Leptodactylus magistris. Whereas, L.
trinacria sp. nov. has not been collected in sympatry with
any other member of the L. melanonotus species group.
Within the L. melanonotus group, L. colombiensis is the
geographically nearest species to L. trinacria sp. nov. Lep-
todactylus colombiensis is found in Colombia’s Caribbean
drainages (Sá et al. (2014). Its presence in Táchira, Ven-
ezuela, previously noted by Barrio-Amorós & Chacón-
Figure 11. Distribution (center, in green) of the Premontane moist forest (Holdridge’s Life Zone) at Mérida city and surroundings, the
region where all Leptodactylus trinacria sp. nov. specimens have been found.
La Marca
34
Ortiz (2001), remains to be conrmed following de Sá et
al. (2014).
Study area and natural history
Habitat and Ecology: e type locality (Fig. 12) is situ-
ated within the Premontane moist forest (Holdridge’s Life
Zone “bosque húmedo premontano” in Ewel et al. 1976).
e region is characterized by a two-seasonal rainfall pat-
tern: a primary dry season from late November to early
March and a shorter, moderate dry period (locally termed
“veranillo de San Juan”) between June and August. High-
precipitation peaks occur from early April to mid-May and
again from mid-September to mid-November. Average an-
nual temperatures range from 16.4°C to 17.8°C (García
et al. 2007). Specimens of the type series were collected
along the margins of an articial lagoon in the city of
Mérida. ey were found among grasses growing on or-
ganic matter, i. e., remnants of aquatic vegetation removed
during maintenance to prevent eutrophication. e site is
currently bordered by urban residential areas. Some speci-
mens were studied in swampy areas and within lentic pools
of slow-moving rivulets in forest remnants at the Urban-
ización Campo Claro, on the Mérida terrace, 8°33’42”N,
71°12’46”W (Fig. 13). At the latter site, a freshwater crab
(Pseudothelphusidae) was observed predating an uniden-
tied tadpole, suggesting a potential predator for the spe-
cies. At this locality, Leptodactylus trinacria sp. nov. was
found to be sympatric with Mannophryne collaris (Bou-
lenger, 1912).
Reproductive behavior
Specimens of Leptodactylus trinacria sp. nov. have
been heard calling in the early morning (approx. 06:00 h),
though calling may persist throughout the day during rainy
periods; no nocturnal activity has been observed. Court-
ship behavior include inguinal amplexus, as observed in
Figure 12. Type locality of Leptodactylus trinacria sp. nov., showing the articial lake at Museo de Ciencia y Tecnología of Mérida city
(within the former “Central Azucarero Los Andes”, CALA), Mérida state, Venezuela, 1,320 m asl. Photo by Enrique La Marca on 18
February 2013.
A new Andean Leptodactylus
35
mating pairs maintained in captivity at the Rescue of En-
dangered Venezuelan Amphibian (REVA) ex situ conser-
vation center in Mérida, Venezuela. e amplexus is ingui-
nal and the male stimulates the female on the sides of the
belly. During oviposition, the male on top of the female
constructs a foam nest by beating the eggs and jelly with
the hind limbs. ese nests are constructed on the surface
of lentic waters from where the tadpoles in early stages of
development emerge (Fig. 14).
Larval biology and parental care
e larvae are exotrophic. Tadpoles exhibit larval
schooling, moving together as a cohesive unit (Fig. 15), a
trait shared with other members of the genus Leptodacty-
lus. Furthermore, we observed larval attendance (parental
care), where an adult female closely guarded batches of
swimming tadpoles. is behavior has been documented
both in the eld and in captivity (La Marca 2017a, b).
Within the genus Leptodactylus, such parental care is oth-
erwise only known in the closely related L. latrans group
(de Sá et al. 2014).
Conservation status and legal framework
Leptodactylus trinacria sp. nov. represents the second
member of the genus in Venezuela requiring immediate
protection, alongside the non-Andean endemic L. magis-
tris. With the data at hand, we are condent to state that
Leptodactylus trinacria sp. nov. is an endangered species,
specially within the limits of Mérida city. Overall, the
populations of the new species have suered a continuous
decline in their original distribution area. e decline is
mainly due to the reduced availability of suitable habitats
because of human modication of their natural environ-
ment. is is an interesting case, joining to several oth-
ers, in which a species is threatened with extinction even
before being formally classied for science (La Marca
Figure 13. Microhabitat of Leptodactylus trinacria sp. nov. at forest remnant in Urbanización Campo Claro, close to Río Albarregas,
on the Mérida terrace. Photo: Enrique La Marca.
La Marca
36
2017ab). No specic legislative measures have been enact-
ed to protect the new species, nor is there ocial protec-
tion for its remaining habitats (La Marca 2016a, b). Before
its formal description, the species was tentatively listed (La
Marca 2015) in the Red Book of Venezuelan Fauna as En-
dangered (EN) B1ab(i,ii,iii). Based on more recent data,
we propose an updated IUCN category of Endangered
(EN) B1ab(iii,iv,v)+2ab(iii,iv,v).
Ex-situ initiatives and husbandry
In early 2014, an ex-situ conservation program was es-
tablished at the Chorros de Milla Zoological Park in Méri-
da (La Marca et al. 2015a, b). Founder individuals were
sourced from forest remnants within the Mérida urban
area. Following quarantine, adults were housed in large
terraria designed to simulate their natural environment,
featuring automated running water, native vegetation, leaf
litter, and rock shelters. Environmental parameters –in-
cluding photoperiod, misting, and seasonal rainfall– were
strictly regulated. Early reproductive eorts faced high
larval mortality due to developmental diseases. However,
aer rening nutritional protocols and micro-ecological
requirements, approximately two dozen froglets rst suc-
cessfully reached metamorphosis. Adults were maintained
on a diverse diet of invertebrates, including Drosophila
melanogaster, Acheta domestica, Galleria mellonella, Te n e-
brio molitor, Sitophilus spp., Armadillidium vulgare, and
Eisenia fetida. Prey items were dusted with calcium and
vitamin supplements, following established protocols al-
ready tested for Mannophryne collaris (La Marca 2016c,
d). Larvae were raised using a specialized formula devel-
oped at the REVA Conservation Center (La Marca &
Castellanos 2018a, b). Between 2014 and 2022, despite
sociopolitical instability (La Marca & Castellanos 2017)
and the COVID-19 pandemic (La Marca 2020b), the pro-
gram successfully released several F1 generation of dozens
Figure 14. Reproductive behavior in Leptodactylus trinacria sp. nov. A: Amplectant pair (note inguinal amplexus). B: Male on top
stimulates lateral sides of female. C: Female spawning aer stimulation. D: Tadpoles emerging from foam nest. Photos: Enrique La
Marca and Daniel Quihua. Archive of the Rescue of Endangered Venezuelan Amphibian (REVA) ex situ conservation center.
A
C
B
D
A new Andean Leptodactylus
37
Figure 15. Larval schooling of Leptodactylus trinacria sp. nov. in the wild. Photo: Enrique La Marca.
of captive-born specimens into the Mérida Botanical Gar-
den and a humid forest remnant in Campo Claro, near the
Albarregas River (La Marca 2017a, b).
Habitat fragmentation and destruction
e primary threat to Leptodactylus trinacria sp. nov. is
the near-total destruction of the seasonal premontane hu-
mid forests that once covered the Mérida terrace and other
places at the middle Chama River basin. is degradation is
largely driven by urban expansion and historical coee plan-
tations resulted in extensive deforestation. e species shares
this precarious situation with the sympatric Mannophryne
collaris; both serving as agship taxa for the vanishing eco-
systems of the Mérida terrace (La Marca, 2017a,b).
Emergent diseases
Analysis of museum specimens (ULABG 4871, 4873,
4875, 4878) tested negative for Batrachochytrium dendro-
batidis (Bd). However, a Leptodactylus specimen (ULA-
BG 4269) from nearby Chiguará tested positive (Lampo
et al. 2005, 2006). While the identity of the Chiguará
specimen remains to be fully conrmed, the geographic
proximity of this population suggests that the new taxon
is likely exposed to the pathogen. It is currently unclear
whether the relatively high ambient temperatures of its
habitat mitigate fungal pathogenicity or if the populations
possess a degree of resistance.
ACKNOWLEDGMENTS
e author is especially indebted to the Escuela de Geo-
grafía and the Instituto de Geografía y Conservación de
Recursos Naturales of the Facultad de Ciencias Forestales
y Ambientales, Universidad de Los Andes (ULA), Mérida,
Venezuela. Financial support for scientic research was
provided by the Consejo de Desarrollo Cientíco, Hu-
manístico y Tecnológico of the Universidad de Los Andes
(CDCHT-ULA). I am grateful to the British Library Na-
tional Sound Archive for generously providing the sono-
graph of the new species.
La Marca
38
I wish to thank Diego Cadenas, Rosemary Sánchez,
Carlos Gottberg, María José Praderio, Enzo La Marca,
Néstor Sánchez, Savanna Luna Sánchez Puente, Iván Men-
doza, Osmary Leal, Michelle Castellanos, Janina Puente,
Luis Felipe Esqueda, Argelia Rodríguez, Francisco Nava,
Daniel Quihua, and Hans-Peter Reinthaler for their in-
valuable assistance in the eld and laboratory. Amphibian
Ark is kindly acknowledged for providing nancial sup-
port to Enrique La Marca for the ex-situ captive breeding
program at the Rescue of Endangered Venezuelan Am-
phibian (REVA) ex situ conservation center in Mérida.
James (Jim) R. Dixon, Christian Reymondin and Mi-
chelle Castellanos generously provided Fig. 2G, Fig. 4, and
Fig. 8(A-C), respectively.
Collecting permit (“Licencia de caza con nes cientí-
cos”) was issued by the Ocina Nacional de Diversidad
Biológica, of the former Ministerio del Ambiente y de
los Recursos Naturales Renovables, of the República
Bolivariana de Venezuela, Ocio Nº 41-0296 of 6 May
2001. Institutional permits for the captive breeding fa-
cilities for Leptodactylus trinacria sp. nov. were issued
by the Ministerio del Poder Popular para el Ecosocial-
ismo (formerly Ministerio del Ambiente y de los Recur-
sos Naturales Renovables), under Providencia No. 0356
(17 October 2016); and the installation and operation
of these facilities were ocially registered under number
DGDB-0002.
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