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Taxonomic Status of Gomphobates Marmoratus Reinhardt and Lütken, 1862 “1861” and Eupemphix Fuscomaculatus Steindachner, 1864 (Amphibia, Anura, Leptodactylidae)

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Taxonomic Status of Gomphobates Marmoratus Reinhardt and Lütken, 1862 “1861” and Eupemphix Fuscomaculatus Steindachner, 1864 (Amphibia, Anura, Leptodactylidae) South American Journal of Herpetology, 1(3), 2006, 166-174 © 2006 Brazilian Society of Herpetology TAXONOMIC STATUS OF GOMPHOBATES MARMORATUS REINHARDT AND LÜTKEN, 1862 “1861” AND EUPEMPHIX FUSCOMACULATUS STEINDACHNER, 1864 (AMPHIBIA, ANURA, LEPTODACTYLIDAE) LUCIANA BARRETO NASCIMENTO1,3; BRUNO VERGUEIRO SILVA PIMENTA2; CARLOS ALBERTO GONÇALVES CRUZ2 AND ULISSES CARAMASCHI2 2 Museu de Ciências Naturais, Departamento de Ciências Biológicas e Programa de Pós-graduação em Zoologia de Vertebrados, PUC Minas, Av. Dom José Gaspar 290, 30535-610, Belo Horizonte, Minas Gerais, Brazil. 3 Departamento de Vertebrados, Museu Nacional/UFRJ, Quinta da Boa Vista, 20940-040, Rio de Janeiro, Rio de Janeiro, Brazil. 3 Corresponding author: [email protected] ABSTRACT: An analysis of specimens deposited in herpetological collections and identified as Physalaemus fuscomaculatus demonstrated that these differ from the types of this taxon, Eupemphix fuscomaculatus. Consequently, the taxonomic status of Gomphobates marmoratus and Eupemphix fuscomaculatus was revised based on the types and on data in the literature. Gomphobates marmoratus is associated to the genus Physalaemus, as Physalaemus marmoratus, and refers to specimens previously identified as Physalaemus fuscomaculatus. Additionally, Eupemphix fuscomaculatus is associated to the genus Pleurodema in the combination Pleurodema fuscomaculata. KEYWORDS: Amphibia, Anura, Taxonomy, Physalaemus, Pleurodema. INTRODUCTION Several specimens identified as Physalaemus fuscomaculatus collected in localities outside the Reinhardt and Lütken (1862 “1861”) described distribution range given by Cei (1990) are deposited Gomphobates marmoratus based on three syntypes in herpetological collections (Nascimento et al., collected in Lagoa Santa (19°37’S, 43°53’W), State of 2005). Analysis of this material demonstrated that Minas Gerais, Brazil. Currently, G. marmoratus is a such specimens differ from the taxon studied by Cei junior synonym of Physalaemus fuscomaculatus (1990), but agree perfectly with the types of Gom- (Steindachner, 1864) (Frost, 2006). phobates marmoratus Reinhardt and Lütken, 1862 Steindachner (1864) described Eupemphix fusco- “1861”. maculatus from one female from “Caiçará” [17°15’S, Based on the examination of the types, we propose 57°10’W, according to Cei (1990)], State of Mato Gros- the revalidation of G. marmoratus, under a new com- so, Brazil. Currently, E. fuscomaculatus is a junior bination, and designate a lectotype and paralectotypes. synonym of Physalaemus fuscomaculatus (Stein- We also assess the taxonomic status of dachner, 1863) (Frost, 2006). This taxon was better E. fuscomaculatus Steindachner, 1863 and propose the defined and illustrated by Cei (1990) based on the ho- allocation of this species in the genus Pleurodema lotype and on additional specimens from Rio Apa Tschudi, 1838. (22°30’S, 57°00’W), a tributary of Rio Paraguay. He considered this species “a relatively uncommon lepto- Historical background dactylid”, ranging from the type locality to the lower basin of Rio Paraguay, probably reaching the marginal Reinhardt and Lütken (1862 “1861”) erected the area of northern Argentinean provinces southwards. genus Gomphobates and described the species Physalaemus fuscomaculatus is presently allocated G. marmoratus from Lagoa Santa, State of Minas in the P. albifrons group (Nascimento et al., 2005) Gerais, Brazil. based on several morphological characters, such as Steindachner (1864) described Eupemphix fusco- dorsal and ventral color patterns, rounded snout, large maculatus from “Caiçará in Brasilien” and pointed the inguinal glands not associated with a dark ocellus, and presence of maxillary and vomerine teeth. He associ- shovel-like external and internal metatarsal tubercles ated Hiobates fuscomaculatus Fitz. Tschudi as a syn- with horned distal margins. onym of this species. Nascimento, L.B. et al. 167 Steindachner (1867) established part of Eupemphix Méhely (1904), after examining the type designat- nattereri Steindachner, 1863 as a junior synonym of ed by Spix, distinguished Paludicola albifrons (Spix) Gomphobates marmoratus, associated Eupemphix from P. fuscomaculata (Steindachner) by the pres- fuscomaculatus to this genus, under the new combi- ence of a second tarsal tubercle in the articulation be- nation Gomphobates fuscomaculatus, and indicated tween tibia and tarsus on the former, larger than the Iliobates fuscomaculatus (assigning it to “Fitz. Ts- distal tarsal tubercle, and by the absence of externally chudi”) as a synonym of the latter. visible lumbar glands. He also indicated that Cope (1869 “1868”) described Lystris brachyops P. fuscomaculata (Steindachner) presents no vomer- (= Pleurodema brachyops) and distinguished this ge- ine teeth. nus from Gomphobates by the presence of fontanela Miranda-Ribeiro (1926) included Eupemphix, En- and vomerine teeth, and from Eupemphix, by the pres- gystomops, Pleurodema, and Paludicola in the fam- ence of well-developed teeth on young and adults. ily Paludicolidae. Paludicola was characterized by the Eupemphix fuscomaculatus Steindachner was then presence of “more or less indistinct vomerine teeth”. included in the genus Lystris, under the combination He compared specimens of Paludicola albifrons Lystris fuscomaculatus (Steindachner). (Spix) and P. fuscomaculata (Steindachner) to the type Peters (1872) included Gomphobates marmora- of Eupemphix fuscomaculatus Steindachner and tus Reinhardt and Lütken, Liuperus marmoratus Bur- agreed with Méhely (1904) on the difference among meister, 1861, and part of Eupemphix nattereri Stein- these species. He also mentioned that dachner in the synonymy of Bufo albifrons Spix, 1824. P. fuscomaculata (Steindachner) presents “vomerine Boulenger (1882) allocated Pleurodema Tschudi, teeth indistinct, but present on a line anterior to the Cystignathus Duméril and Bibron, Leiuperus Duméril choanae”. He indicated the distribution of P. albifrons and Bibron, Pleurodema Günther, Gomphobates Re- (Spix) for the State of Bahia, Brazil, and that of inhardt and Lütken, Eupemphix Steindachner, and P. fuscomaculata (Steindachner) for the states of Mato Lystris Cope in the genus Paludicola Wagler, based Grosso, São Paulo, Paraná, and Rio Grande do Sul in on the presence of maxillary teeth. Consequently, the Brazil, and also in Argentina and Uruguay. new combinations Paludicola fuscomaculata (Stein- The first revision of the genus Paludicola was dachner) and Paludicola albifrons (Spix) were es- presented by Parker (1927), who distributed its spe- tablished for Lystris fuscomaculatus and Bufo albi- cies among the genera Physalaemus Fitzinger, 1826, frons, respectively. Boulenger cited the absence of Pseudopaludicola Miranda-Ribeiro, 1926, and Pleu- vomerine teeth in P. fuscomaculata (Steindachner) and rodema. Paludicola biligonigera (Cope), established its distribution range to Brazil, Uruguay, and P. fuscomaculata (Steindachner), and P. albifrons Buenos Aires (Argentina). Paludicola biligonigera (Spix) were included in the genus Physalaemus. He (Cope) was associated to Reinhardt’s specimens from also discussed the differences indicated by Mehély Lagoa Santa (State of Minas Gerais), and from the (1904) between Physalaemus albifrons (Spix) and States of Bahia and Pará, Brazil. P. fuscomaculatus (Steindachner). Parker (1927) re- Boulenger (1886) indicated the occurrence of Palu- moved Gomphobates marmoratus Reinhardt and dicola fuscomaculata (Steindachner) and Paludico- Lütken from the synonymy with P. albifrons (Spix) la albifrons (Spix) in the State of Rio Grande do Sul, and considered it a junior synonym of Brazil. Boulenger (1887) presented a diagnosis of Palu- P. fuscomaculatus (Steindachner), after requesting dicola albifrons from Porto Alegre, Rio Grande do the analysis of the type-specimens of P. albifrons Sul, Brazil. He indicated that Leiuperus marmoratus (Spix). Burmeister was not identical to Paludicola albifrons Cochran (1955) mentioned the absence of vomer- (Spix) as stated by Peters (1872), but was a junior syn- ine teeth in Physalaemus fuscomaculatus (Stein- onym of P. fuscomaculata (Steindachner). In reality, dachner), “although the bony projection on which they Leiuperus marmoratus D’Orbigny, 1847 (referred by would have arisen can be felt and in a few cases seen Burmeister, 1861), Leiuperus marmoratus Burmeis- between the choanae”. The description presented was ter, 1861, and Paludicola fuscomaculata Boulenger, based on specimens from the States of Minas Gerais, 1886, 1887 refers to the currently recognized Phys- Rio Grande do Sul, and São Paulo (Brazil), Argentina, alaemus biligonigerus (Cope, 1861). Bolivia, and Paraguay. 168 Taxonomic status of Gomphobates marmoratus Milstead (1960) characterized Physalaemus bili- He pointed out that the morphology of the former spe- gonigerus (Cope) and P. fuscomaculatus (Stein- cies is inaccurately determined, because Stein- dachner), species occurring in the State of Rio Grande dachner’s description of the type clearly indicated the do Sul, Brazil, and pointed that the first differs from all presence of maxillary and vomerine teeth. Physalae- congeneric species by the presence of vomerine teeth. mus fuscomaculatus (Steindachner) was character- He observed differences on snout-vent length between ized morphologically based on the holotype and on spec- specimens of P. fuscomaculatus (Steindachner) from imens collected at Rio Apa, a tributary of Rio Para- the States of São Paulo and Rio Grande do Sul, sug- guay, not far from the type-locality, adding
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