Research Article |
Corresponding author: Andrés R. Acosta Galvis ( aacosta@humboldt.org.co ) Academic editor: Angelica Crottini
© 2018 Andrés R. Acosta Galvis, Rafael O. de Sá.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Galvis ARA, de Sá RO (2018) Leptodactylus validus Garman, 1888 in Colombia: its distribution and identification. ZooKeys 737: 113-123. https://doi.org/10.3897/zookeys.737.20442
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Leptodactylus validus is reported for the first time for Colombia, corresponding to the tenth species of the L. melanonotus species group occurring in the country. In collections, all L. validus specimens were identified as L. colombiensis. Morphological, coloration, and ecological characters are provided to differentiate the two species in Colombia. Furthermore, the distribution of L. validus is expanded based on the examination of specimens in both collections and literature records. In addition, the advertisement call of L. validus from Colombia is compared with those reported for other continental and insular populations; the calls are slightly more similar to those of insular populations.
Advertisement call, ecology, habitat, male nuptial spines, morphology
Neotropical frogs of the genus Leptodactylus represent a clade consisting of 75 currently recognized species; 32 % of the species are known to occur in Colombia (
The presence of Leptodactylus validus in Colombia is reported and the characteristics that differentiate it from L. colombiensis are discussed. Information on the call, characteristics of the habitat, and distribution of L. validus in Colombia are provided.
Specimens herein identified as L. validus are deposited in the amphibian collections of the Universidad Javeriana, Bogotá (
SVL snout-vent length,
HW head width,
HL head length from tip of snout to the posterior border of skull (i.e., posterior edge of prootic, noted through the skin),
IND internarinal distance,
IOD interorbital distance,
ED eye diameter,
END eye-nostril distance,
TD tympanum diameter,
UEW upper eyelid width,
ETS distance between the anterior edge of the eye to the tip of snout,
RW rostral width,
TL tibia length,
FL femur length,
FTL foot length,
HDL hand length (measurements are provided as a Suppl. material
Means are reported as +/- one standard error. Morphological terminology follows
Two recordings were analyzed, both are deposited (i.e., BSA call vouchers) at the Colección de Sonidos Ambientales del Instituto Alexander von Humbdolt. Call voucher BSA-15988 corresponds to a specimen recorded on September 4, 2008, Municipio San Martin, Departamento de Meta, Colombia, 460 mts altitude (3°39.3'N, 73°36.51'W). The specimen was calling inside a water pond, although the recorded individual escaped collection temperature was recorded. Other vouchers individuals of the species were collected at the time of recording and are deposited with numbers MUJ6168-69 (Universidad Javeriana). Call voucher BSA-15994 corresponds to a specimen recorded and collected on June 4, 2012, Municipio Puerto Gaitan, Departamento de Meta, Colombia, 204 mts altitude (4°19.44’N, 71°43.54’W). The specimen was calling in a temporal pond, air temperate was 24.9°, and specimen voucher is deposited with number
Leptodactylus validus Garman, 1888 is a small species within the L. melanonotus species group; in Colombian populations adult females have a snout-vent length (SVL) of 25.3–49.5 mm (x̄ = 32.3 mm, N = 43) whereas in adult males SVL ranges between 25.0–40.8 mm (x̄ = 30.1 mm, N = 26); snout is short and rounded in dorsal view, not spatulate; distinctive light color stripes on the upper lip, including under the eye; males lack chest spines; a light color stripe extends ventrally from the corner of the eye and running anterior to the tympanum to the jaw articulation; this white band also extends over an elongated gland running between the jaw articulation and the point of attachment of the arm to the body; in addition a supratympanic gland, overall light brownish in coloration and, in some specimens, with a darker brown ventral edge, extends from the posterior edge of the eye, over the tympanum, and curving ventrally to the insertion of the arm; dorsal folds absent; dorsolateral folds usually short and interrupted, but rarely absent, in life they have a reddish to light coffee coloration; lateral folds absent; coloration of dorsum is overall a light brownish, posterior thighs and shanks dorsally with distinct and transverse dark bands (Fig.
Distribution in Colombia. Leptodactylus validus occurs in north eastern South America; in Colombia the species is found in the western savannahs of Colombia, known as the “Llanos bajos” or “Llanos inundables” (Fig.
Advertisement Call. The advertisement call of Leptodactylus validus from populations in the western savannahs of Colombia consists of two notes. The first note consists of 1–3 pulses that are weak and increase gradually to the second note which consists of 2–6 pulses of wide amplitude that decrease gradually to lower amplitudes (Fig.
Habitat and ecology. Leptodactylus validus occurs in transitional ecotones between gallery forests and: a) savannahs, b) “Morichales” floodplains (i.e., communities dominated by Moriche palms, Mauritia flexuosa), and c) eolic savannahs. During the dry season (December–March) the species was found actively moving at night among the leaf litter in flooded areas close to streams and creeks within gallery forests; but males were not calling. Throughout the rainy season (May–June), L. validus was found in large numbers and calling in flooded areas and temporal ponds within savannahs and ecotonal environments. Examination of specimens collected in the rainy season show that they were sexually mature, e.g., larger body size, males with prepollical spines, and females with large oviductal eggs. Specimens collected in the dry season have smaller body size and were collected among the leaf litter at the edges of streams or permanent ponds within gallery forests (Fig.
Continental populations of Leptodactylus validus from Venezuela, Guyana, Surinam, and the north-central region of Roraima, Brazil, were identified as L. pallidirostris (Heyer, 1994). Furthermore, the author noted that a morphological similar species, L. validus, occurred in Trinidad and Tobago and across some of the oceanic islands of the Lesser Antilles (e.g., Sant Vicent, Bequia, Grenada). A study of Leptodactylus pallidirostris placed the species in the synonymy of L. validus (
Acoustic parameters were described and compared for South American mainland populations (i.e., Brazil and Venezuela) and from the continental islands of the Lesser Antilles (i.e., Trinidad and Tobago) (
Comparison of L. validus with L. colombiensis. Examinations found that in collections all specimens of L. validus had been misidentified as L. colombiensis based on external morphology. Interestingly, in the species accounts of the genus Leptodactylus (
Leptodactylus validus in the Colombian savannahs can be distinguished from L. colombiensis by its smaller adult body size [L. validus: females SVL = 25.3–49.5 mm (x̄ = 32.3 mm, N = 43) and males SVL = 25.0–40.8 mm (x̄ = 30.1 mm, N = 26) and L. colombiensis females SVL = 39.88–60.44 mm (x̄ = 50.59 mm, N = 37) and males 34.1–52.9 mm (x̄ = 40.65 mm, N = 21)]. Furthermore, the interorbital distance (IOD)/length of the tibia (TL) ratio consistently separates the two species; this ratio was previously found useful to differentiate species of Lithobates (Hillis and de Sá, 1988). The IOD/LT ratio in L. colombiensis ranges between 13.6–17.7% (N = 58) whereas in L. validus is always greater (17.9–27.6%; N = 86; Fig.
The advertisement calls of L. validus (as L. validus for insular populations and L. pallidirostris for mainland populations) have been previously analyzed (
Interestingly, distinct differences were found between the ecology and distribution of L. validus and L. colombiensis in Colombia. Leptodactylus validus has a strictly Cis-Andean distribution associated with the floodplains of the “Llanos Orientales”, that is savannahs with an altitudinal range from 35 m to 204 m and a precipitation of less than 2,800 mm (Fig.
The records for Leptodactylus validus reported here correspond to the western boundary of the species distribution (Fig.
Leptodactylus validus Garman, 1888 is reported to occur in the western savannahs of Colombia (= Llanos Orientales). The species has been commonly confused with L. colombiensis from which it can be distinguished by morphological, call, and ecological characteristics.
This research was done under Research Agreement # 15-276 between the Institute Alexander von Humboldt for Research of Biological Resources and the University of Richmond. RdS was supported by NSF-DEB award 1144692. The authors thank the different institutions, local communities, and projects that allow the collections of specimens along the Orinoco Basin, in particular to the Asociación de Becarios de Casanare (ABC). Collections in Meta and Vichada were done during the assessment of 'areas of high conservation value' under WWF-Fundación Omacha and Colombia Bio Program of COLCIENCIAS-IAvH (Cooperation Agreement Number FP44842-109-2016[IAVH 16-062]). We thank Dr. Julio Mario Hoyos, Universidad Javeriana, Bogotá for his assistance. We also thank Dr. Michael Jowers, Dr. Julio M. Hoyos, and an anonymous reviewer for their comments.
Measurements of Leptodactylus validus
Data type: specimens data