Catalogue of the Amphibians of Venezuela: Illustrated and Annotated Species List, Distribution, and Conservation 1,2César L
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Mannophryne vulcano, Male carrying tadpoles. El Ávila (Parque Nacional Guairarepano), Distrito Federal. Photo: Jose Vieira. We want to dedicate this work to some outstanding individuals who encouraged us, directly or indirectly, and are no longer with us. They were colleagues and close friends, and their friendship will remain for years to come. César Molina Rodríguez (1960–2015) Erik Arrieta Márquez (1978–2008) Jose Ayarzagüena Sanz (1952–2011) Saúl Gutiérrez Eljuri (1960–2012) Juan Rivero (1923–2014) Luis Scott (1948–2011) Marco Natera Mumaw (1972–2010) Official journal website: Amphibian & Reptile Conservation amphibian-reptile-conservation.org 13(1) [Special Section]: 1–198 (e180). Catalogue of the amphibians of Venezuela: Illustrated and annotated species list, distribution, and conservation 1,2César L. Barrio-Amorós, 3,4Fernando J. M. Rojas-Runjaic, and 5J. Celsa Señaris 1Fundación AndígenA, Apartado Postal 210, Mérida, VENEZUELA 2Current address: Doc Frog Expeditions, Uvita de Osa, COSTA RICA 3Fundación La Salle de Ciencias Naturales, Museo de Historia Natural La Salle, Apartado Postal 1930, Caracas 1010-A, VENEZUELA 4Current address: Pontifícia Universidade Católica do Río Grande do Sul (PUCRS), Laboratório de Sistemática de Vertebrados, Av. Ipiranga 6681, Porto Alegre, RS 90619–900, BRAZIL 5Instituto Venezolano de Investigaciones Científicas, Altos de Pipe, apartado 20632, Caracas 1020, VENEZUELA Abstract.—Presented is an annotated checklist of the amphibians of Venezuela, current as of December 2018. The last comprehensive list (Barrio-Amorós 2009c) included a total of 333 species, while the current catalogue lists 387 species (370 anurans, 10 caecilians, and seven salamanders), including 28 species not yet described or properly identified. Fifty species and four genera are added to the previous list, 25 species are deleted, and 47 experienced nomenclatural changes. Eleutherodactylus terraebolivaris Rivero, 1961 is synonymized with Hylodes incertus Lutz, 1927 as Pristimantis incertus. Oreophrynella dendronastes Lathrop and MacCulloch, 2007, is considered a junior synonym of O. macconnelli (Boulenger 1895). Centrolene Jiménez de la Espada, 1872, is a feminine genus, so all species in the genus are amended. Centrolenella pulidoi Rivero, 1968 is considered a junior synonym of Hyla benitezi Rivero, 1961, as Boana benitezi. Centrolenella estevesi Rivero, 1968 is considered a junior synonym of Hyla jahni Rivero, 1961, as Hyloscirtus jahni. Illustrated herein are 300 species (77.5% of the total). Lastly, the distributions for all species are revised, species that possibly occur within Venezuela are suggested, and comments are provided on nomenclature and conservation issues. Keywords. Biogeography, checklist, Anura, Urodela, Gymnophiona, management Resumen.—Se presenta una lista anotada de los anfibios de Venezuela, actualizada hasta diciembre de 2018. La última lista comprensiva (Barrio-Amorós 2019c) incluyó un total de 333 especies, mientras que la lista actual contiene 387 especies (370 anuros, 10 cecilias y siete salamandras), incluyendo 28 especies aun no descritas o identificadas propiamente. 50 especies y cuatro géneros se añaden a la lista previa, 25 especies se eliminan y 47 de ellas han experimentado cambios nomenclaturales. Eleutherodactylus terraebolivaris Rivero, 1961 se sinonimiza con Hylodes incertus Lutz, 1927, como Pristimantis incertus. Oreophrynella dendronastes Lathrop and MacCulloch, 2007, se considera sinónimo de O. macconnelli (Boulenger, 1895). Centrolene Jiménez de la Espada, 1872, es un género femenino, así que se emendan todos los nombres acorde. Centrolenella pulidoi Rivero, 1968 es considerado sinónimo de Hyla benitezi Rivero, 1961, como Boana benitezi. Centrolenella estevesi Rivero, 1968, se considera sinónimo de Hyla jahni Rivero, 1961, como Hyloscirtus jahni. Se presentan fotografías de 300 especies (77.5% del total). Por último, la distribución de todas las especies es revisada, se sugieren especies que podrían estar presentes en Venezuela y se presentan comentarios sobre nomenclatura y conservación. Palabras clave. Anura, biogeografía, Gymnophiona, lista, manejo, Urodela Citation: Barrio-Amorós CL, Rojas-Runjaic FJM, Señaris JC. 2019. Catalogue of the amphibians of Venezuela: Illustrated and annotated species list, distribution, and conservation. Amphibian & Reptile Conservation 13(1) [Special Section]: 1–198 (e180). Copyright: © 2019 Barrio-Amorós et al. This is an open access article distributed under the terms of the Creative Commons Attribution License [At- tribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org. Received: 27 August 2016; Accepted: 26 April 2018; Published: 14 July 2019 Introduction amphibian diversity, after Brazil (1,026 species; http:// br.herpeto.org/anfibios/), Colombia (813; https:// In consideration of the findings presented herein, www.batrachia.com), Peru (657; http://research.amnh. Venezuela (387 species) will be the seventh- org/vz/herpetology/amphibia/index.php//content/ most diverse country in the world with respect to search?taxon=&subtree=&subtree_id=&english_na Correspondence. 1,2 [email protected], 3,4 [email protected], 5 [email protected] Amphib. Reptile Conserv. 1 July 2019 | Volume 13 | Number 1 | e180 Catalogue of the amphibians of Venezuela Fig. 1. Number of amphibian species known in Venezuela since 1927. me=&author=&year=&country=167), Ecuador (594; many were deleted from the list due to synonymy or https://bioweb.bio/faunaweb/amphibiaweb/), Indonesia misidentification. The suprageneric systematics are still (457; http://research.amnh.org/vz/herpetology/amphibia/ in flux and subject to numerous, and sometimes shocking, index.php//content/search?taxon=&subtree=&subtree_ changes as well as errors (see Heinicke et al. 2009; Pyron id=&english_name=&author=&year=&country=166), and Wiens 2011; De Sá et al. 2012, 2014; Padial et al. and Mexico (392; Parra-Olea et al. 2014; https:// 2014; Duellman et al. 2016). For a relatively stable listing amphibiaweb.org/amphibian/newspecies.html). in the face of significant taxonomic changes, the reader is Several checklists for Venezuela, with varying advised to consult Frost (2019). This publication focuses levels of detail, have appeared since the 1950s (Fig. on concerns with taxonomic and nomenclatural issues 1), and clearly the number of known species has grown relevant to Venezuelan amphibians and not on major, with time and increased inventory activity, from 74 broad changes in species relationships or names. frog species in 1959 to 370 in 2019. The most data- intensive and continuously updated listing of the world’s Material and Methods amphibian species is currently Amphibian Species of the World 6.0 (http://research.amnh.org/vz/herpetology/ Several methods were used to clarify how the amphibia/index.php), an Online Reference whose book determination of the taxonomic status was made for some predecessor was published by Darrel Frost in the mid- taxa. The majority of species are listed by their generic 1980s (Frost 1985). Another important online source and specific names, under the Family level. Not used are is the AmphibiaWeb site (http://www.amphibiaweb. unranked taxa uniting families (Arboranae, Terraranae), org). Although both online databases are quite useful which can facilitate classification uses in some cases as references, they lack precision in their ability to (Hedges et al. 2008; Duellman et al. 2016). Rather, specifically identify local species, and should not be families are placed in alphabetical order. Where there are taken as the final taxonomic authority (Dubois 2017). It doubts regarding specific identity, three disclaimers are is also unwise to rely on a third major online database, used. The first is the placement of aff. (for affinis, from the IUCN Red List of Threatened Species (IUCN 2015), the Latin for closely related or akin, but not identical to) since entries often lag behind current research, and with between the generic and specific names. This is used few exceptions, corrections and updates are irregular. to identify taxa for which evidence indicates existing Ten years after the last published official checklist differences from a named, very similar taxon, but for (Barrio-Amorós 2009c), the panorama has changed which this difference has not yet been further evaluated. again in many ways. Several new species have been Thus, a species may be presented under a name, and a described, others reported as new for the country, and second entry may be made for the species name with aff. Amphib. Reptile Conserv. 2 July 2019 | Volume 13 | Number 1 | e180 Barrio-Amorós et al. inserted. A second disclaimer is cf. (for confer, from the Biogeography Latin for compare with), used in cases where there is insufficient evidence to connect the form to an existing Rivero (1961) was the first to present a zoogeography name, or to determine whether a close affinity exists. The of Venezuelan amphibians. His approach was insightful, last disclaimer is the addition of the term “sp.” to only the especially considering the level of knowledge at the time generic name, for forms with no current specific name