Systematics of the Neotropical Genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the Relevance of Non-molecular Evidence, and Species Accounts

A phylogeny of the species-rich clade of the Neotropical frog genus Leptodactylus sensu stricto is presented on the basis of a total evidence analysis of molecular (mitochondrial and nuclear markers) and non-molecular (adult and larval morphological and behavioral characters) sampled from > 80% of the 75 currently recognized species. Our results support the monophyly of Leptodactylus sensu stricto, with Hydrolaetare placed as its sister group. The reciprocal monophyly of Hydrolaetare and Leptodactylus sensu stricto does not require that we consider Hydrolaetare as either a subgenus or synonym of Leptodactylus sensu lato. We recognize Leptodactylus sensu stricto, Hydrolaetare, Adenomera, and Lithodytes as valid monophyletic genera. Our results generally support the traditionally recognized Leptodactylus species groups, with exceptions involving only a few species that are easily accommodated without proposing new groups or significantly altering contents. The four groups form a pectinate tree, with the Leptodactylus fuscus group diverging first, followed by the L. pentadactylus group, which is sister to the L. latrans and L. melanonotus groups. To evaluate the impact of non-molecular evidence on our results, we compared our total evidence results with results obtained from analyses using only molecular data. Although non-molecular evidence comprised only 3.5% of the total evidence matrix, it had a strong impact on our total evidence results. Only one species group was monophyletic in the molecular-only analysis, and support differed in 86% of the 54 Leptodactylus clades that are shared by the results of the two analyses. Even though no non-molecular evidence was included for Hydrolaetare, exclusion of that data partition resulted in that genus being nested within Leptodactylus, demonstrating that the inclusion of a small amount of non-molecular evidence for a subset of species can alter not only the placement of those species, but also species that were not scored for those data. The evolution of several natural history and reproductive traits is considered in the light of our phylogenic framework. Invasion of rocky outcrops, larval oophagy, and use of underground reproductive chambers are restricted to species of the Leptodactylus fuscus and L. pentadactylus groups. In contrast, larval schooling, larval attendance, and more complex parental care are restricted to the L. latrans and L. melanonotus groups. Construction of foam nests is plesiomorphic in Leptodactylus but their placement varies extensively (e.g., underground chambers, surface of waterbodies, natural or excavated basins). Information on species synonymy, etymology, adult and larval morphology, advertisement call, and geographic distribution is summarized in species accounts for the 30 species of the Leptodactylus fuscus group, 17 species of the L. pentadactylus group, eight species of the L. latrans group, and 17 species of the L. melanonotus group, as well as the three species that are currently unassigned to any species group.Se presenta una filogenia del género Leptodactylus, un ciado neotropical rico en especies, basada en análises combinados de datos moleculares (marcadores nuclear y mitocondriales) y no moleculares (caracteres de la morfología de adultos y larvas así como de comportamiento) se muestrearon > 80% de las 75 especies reconocidas. Los resultados apoyan la monofília de Leptodactylus sensu stricto, con Hydrolaetare como su grupo hermano. La monofília recíproca de Hydrolaetare y Leptodactylus no requiere considerar a Hydrolaetare como un subgénero o sinónimo de Leptodactylus sensu lato. Se reconocen Leptodactylus sensu stricto, Hydrolaetare, Adenomera y Lithodytes como géneros monofiléticos válidos. Los resultados en general resuelven los grupos tradicionalmente reconocidos de Leptodactylus, con excepciones de algunas especies que son reasignadas sin la necesidad de proponer nuevos grupos o alterar significativamente el contenido de los grupos tradicionales. Los cuatro grupos de especies forman una topología pectinada donde el grupo de L. fuscus tiene una posición basal, seguido por el grupo de L. pentadactylus que es el grupo hermano al clado formado por los grupo de L. latrans y L. melanonotus. Se estimó el impacto de los datos no moleculares en los resultados, comparándose los resultados de evidencia total con los de los análises de datos moleculares solamente. Los datos no moleculares representan un 3.5% de la matriz de evidencia total, pero estos datos tuvieron un impacto significativo en los resultados del análisis de evidencia total. En el análisis estrictamente molecular solamente un grupo de especies resultó monofilético, y el apoyo difirió en 86% de los 54 ciados de Leptodactylus compartidos entre los dos análises. A pesar que datos no moleculares no fueron incluidos para Hydrolaetare, la exclusión de evidencia no molecular resultó en el género estar dentro de Leptodactylus, demostrando que la inclusión de evidencia no molecular pequeña para un subgrupo de especies altera no solamente la posición topológica de esas especies, sino tambien de las especies para las cuales dichos datos no fueron codificados. La evolución de patrones de historia natural y reprodución se evalúan en el contexto filogenético. La invasión de afloramientos rocosos y la construción de cámaras de reprodución subterraneas está limitada a los grupos de Leptodactylus fuscus y L. pentadactylus, mientras que la oofagia larval está restringida al grupo de L. pentadactylus. Por otro lado, los cárdumenes larvales, la proteción del cárdumen, y otros comportamientos parentales complejos carecterizan al clado formado por los grupos de especies de L. latrans y L. melanonotus. Los resúmenes de especies incluyen información de sinonimias, etimología, morfología de adultos y larvas, cantos, y distribución geográfica para las 30 especies del grupo de Leptodactylus fuscus, 17 especies del grupo L. pentadactylus, ocho especies del grupo de L. latrans, 17 especies del grupo de L. melanonotus, así como para las tres especies que actualmente no se encuentran asociadas a ninguno de los grupos de especies.Taran Grant was supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico Proc. 307001/2011-3 and Fundação de Amparo à Pesquisa do Estado de São Paulo Proc. 2012/10000-5

On the presence of Rana ridibunda ridibunda Pallas in the Netherlands

Since long there has been a dispute whether in the Netherlands there occur one or two species of green frogs of the genus Rana. There has never been any doubt concerning the presence of Rana esculenta Linnaeus, a species widely distributed throughout the country (Van de Bund, 1964, 1968). The first to mention Rana ridibunda ridibunda Pallas from the Netherlands were Van Kampen & Heimans ( 1927), who observed that they saw a specimen of Rana esculenta var. ridibunda from Den Horn in the province of Groningen. Unfortunately they did not state where the specimen was deposited and no such specimen could be found in the collections of the Rijksmuseum van Natuurlijke Historie (RMNH) in Leiden or in the Instituut voor Taxonomische Zoölogie in Amsterdam. Since that time there have been several additional records of the species. Van Wijk (1946, 1947, 1951) recorded the species from the provinces of Groningen, Zuid-Holland and Limburg. However, these data are not very reliable as they were based on field-observations by untrained people, mainly persons who keep reptiles and amphibians as pets. The material on which their observations were based was not available for examination, so the identifications could not be checked. Van Wijk (1947, 1951) also stated that he had received specimens from Groningen which he identified as R. ridibunda. As the characters he used to distinguish between the green frogs were correct, his identification probably was correct too. Unfortunately, also this material is not available for examination. The next author dealing with this problem is Van de Bund (1964), who gives more details concerning the localities of the specimens previously recorded by Van Wijk (1946, 1947, 1951): Groningen (Groningen, Haren, Eelde), Zuid-Holland (Sassenheim, Schiedam), Limburg (Meerssen). However, Van de Bun

Rediscovery of a forgotten snake in an unexpected place and remarks on a small herpetological collection from southeastern Brazil

A collection of snakes and frogs collected in the area of Porto Real, Rio de Janeiro State, Brazil, was bought in December 1890 and remained unidentified until now. The collection comprises 100 snakes belonging to 18 species and 152 frogs belonging to 19 species. A comparison with the herpetofauna from nearby Serra do Japi in São Paulo is made. General remarks on variation and identification are made for a number of species. Among the snakes were two specimens of Cercophis aurata (Schlegel, 1837), a species described on the basis of one specimen from Suriname, and no new specimens having been recorded until now. The new individuals cause the known area of distribution to be greatly enlarged, but such a large distribution area occurs in several other snakes as well. Morphological data, drawings of details, habitus photographs and a detailed description of the species, based on the three available specimens, are provided

On a new species of toad from southern Morocco

A new species of toad, related with Bufo calamita Laurenti and B. viridis Laurenti, is described from the southern part of Morocco. Combining the data of the available specimens with literature records it was established that the new species probably ranges from the Sous valley southwards to Cape Bojador in the Spanish Sahara. The species seems to be restricted to semi-arid habitats. Tadpoles which probably belong to this species are described. Keys for the identification of adult toads and for the tadpoles are given

Resurrection of Hyla wavrini Parker (Amphibia: Anura: Hylidae), a gladiator frog from northern South America

During field work in Venezuela and Brazil a difference in call was noticed between specimens of large hylids of the Hyla boans-group. Based on this and on morphological differences, it was concluded that Hyla wavrini Parker, 1936, up till now considered a synonym of Hyla boans (Linnaeus, 1758), should be considered a valid species, of which Hyla miranda-ribeiri Melin, 1941 is a synonym. Durante trabajo de campo en Venezuela y el Brasil se nota una diferencia en el canto de especimenes de gran Hílidos del grupo Hyla boans. Basandose en este observación y en diferencias morfológicas, se concluye que Hyla wavrini, considerado até horita como sinónimo de Hyla boans debe ser considerado como especie valido, y que Hyla miranda-ribeiri es un sinónimo de el. Durante trabalhos de campo na Venezuela e Brasil, observou-se diferenças no canto entre espécimenes de grandes hilídeos do grupo Hyla boans. Baseado nisso e em diferenças morfológicas, conclui-se que Hyla wavrini, até o momento considerado sinónimo de Hyla boans, deve ser considerado uma espécie valida, da qual Hyla miranda-ribeiri é um sinónimo