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Monophyly and Taxonomy of the Neotropical Seasonal Killifish Genus Leptolebias (Teleostei: Aplocheiloidei: Rivulidae), with the Description of a New Genus

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Monophyly and Taxonomy of the Neotropical Seasonal Killifish Genus Leptolebias (Teleostei: Aplocheiloidei: Rivulidae), with the Description of a New Genus Zoological Journal of the Linnean Society, 2008, 153, 147–160. With 11 figures Monophyly and taxonomy of the Neotropical seasonal killifish genus Leptolebias (Teleostei: Aplocheiloidei: Rivulidae), with the description of a new genus WILSON J. E. M. COSTA* Downloaded from https://academic.oup.com/zoolinnean/article/153/1/147/2606377 by guest on 23 November 2020 Laboratório de Ictiologia Geral e Aplicada, Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Caixa Postal 68049, CEP 21944-970, Rio de Janeiro, Brazil Received 30 March 2007; accepted for publication 4 July 2007 A phylogenetic analysis based on morphological characters indicates that Leptolebias Myers, 1952, a genus of small killifishes highly threatened with extinction, from Brazil, is paraphyletic. As a consequence, Leptolebias is restricted in this study to a well-supported clade that includes Leptolebias marmoratus (Ladiges, 1934), Leptolebias splendens (Myers, 1942), Leptolebias opalescens (Myers, 1942), and Leptolebias citrinipinnis (Costa, Lacerda & Tanizaki, 1988), from the coastal plains of Rio de Janeiro, and Leptolebias aureoguttatus (Cruz, 1974) (herein redescribed, and for which a lectotype is designated) and Leptolebias itanhaensis sp. nov., from the coastal plains of São Paulo and Paraná, in southern Brazil. Leptolebias is diagnosed by three synapomorphies: a caudal fin that is longer than deep, a single anterior supraorbital neuromast, and dark pigmentation that does not extend to the distal portion of the dorsal fin in males. A key is provided for the identification of species of Leptolebias. Three species formerly placed in Leptolebias, Leptolebias minimus (Myers, 1942), Leptolebias fractifasciatus (Costa, 1988), and Leptolebias cruzi (Costa, 1988), are transferred to Notholebias gen. nov., which is hypothesized to be the sister group to the clade comprising Leptolebias, Campellolebias Vaz-Ferreira & Sierra, 1974, and Cynopoecilus Regan, 1912. Notholebias gen nov. is diagnosed by two synapomorphies: a narrow basihyal and the presence of iridescent bars on the caudal fin in males; and three features interpreted as plesiomorphic, but not occurring in Leptolebias, Campellolebias,orCynopoecilus, the presence of dermosphenotic, well-developed contact organs on the pectoral fin in males, and an opercular region with red bars in males. ‘Leptolebias’ leitaoi, a species from Bahia, in north-eastern Brazil, is considered as having an uncertain phylogenetic position, as all known preserved material is presently lost, and the species may be extinct. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 147–160. ADDITIONAL KEYWORDS: annual fishes – Atlantic forest – biodiversity – Brazil – conservation – killifishes – systematics. INTRODUCTION Cynopoecilus and Campellolebias constitute an equally well-corroborated clade of internally insemi- The monophyly of the group comprising the south- nating killifishes (Costa, 1990a, 1998; Hrbek & eastern South American seasonal killifish genera Larson, 1999; Murphy et al., 1999), which have been Cynopoecilus Regan, 1912, Campellolebias Vaz- included in recent taxonomic revisions (Costa, 2002a, Ferreira & Sierra, 1974, and Leptolebias Myers, 1952 2006a). Leptolebias, a genus of small, externally was first proposed by Costa (1990a). This assemblage, inseminating species (maximum adult size ~20– named as the subtribe Cynopoecilina by Costa 30-mm standard length), is, however, still poorly (1990b), was subsequently tested and corroborated by diagnosed. more detailed phylogenetic analyses, based both on All species presently placed in Leptolebias are morphology (Costa, 1998) and molecular data (Hrbek endemic to the coastal plains of eastern, south- & Larson, 1999; Murphy, Thomerson & Collier, 1999). eastern, and southern Brazil (Costa, 2003). They inhabit seasonal pools of dark acidic water (pH 3.5– *Corresponding author. E-mail: [email protected] 6.0), which are often found in dense forests (Atlantic © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153, 147–160 147 148 W. J. E. M. COSTA forest) or sometimes in coastal open vegetation (formerly in Stanford University) [CAS (SU)]; (Restinga) (Costa, 1995a). These habitats are shallow Museu de Ciências e Tecnologia da Pontifícia Uni- (about 10–40-cm deep) and dry out seasonally, often versidade Católica, Porto Alegre (MCP); Museu twice a year, usually between February and March, Nacional, Universidade Federal do Rio de Janeiro, and between August and September. The greatest Rio de Janeiro (MNRJ); Museu de Zoologia, Univer- diversity of the genus is concentrated in south- sidade de São Paulo, São Paulo (MZUSP); Instituto eastern Brazil (Rio de Janeiro state), in which seven de Biologia, Universidade Federal do Rio de Janeiro, [Leptolebias citrinipinnis (Costa, Lacerda & Tanizaki, Rio de Janeiro (UFRJ). Comparative material 1988), Leptolebias cruzi (Costa, 1988), Leptolebias used in the phylogenetic analysis is listed in the fractifasciatus (Costa, 1988), Leptolebias marmoratus Appendix. The number of specimens are proceeded (Ladiges, 1934), Leptolebias minimus (Myers, 1942), by the catalog number. Downloaded from https://academic.oup.com/zoolinnean/article/153/1/147/2606377 by guest on 23 November 2020 Leptolebias opalescens (Myers, 1942), and Leptolebias Measurements and counts follow the procedures splendens (Myers, 1942)] of the nine valid species of described by Costa (1995a). Measurements are pre- the genus are endemic (Costa, 2003). One species, sented as percentages of the standard length (SL), Leptolebias leitaoi (Cruz & Peixoto, 1992), is endemic except for those related to head morphology, which to eastern Brazil (Mucuri River basin, Bahia state), are expressed as percentages of head length. Fin-ray and another species, Leptolebias aureoguttatus (Cruz, counts include all elements. Number of vertebrae, gill 1974), was first recorded from Paraná state, in the rakers, and caudal-fin rays were recorded only from southern Brazilian coastal plains (Myers, 1952), but cleared-and-stained (c&s) specimens. The compound was subsequently also recorded from adjacent areas caudal centrum was counted as a single element. of south-eastern Brazil, between the Itanhaém and Osteological preparations were made according to Ribeira de Iguape river basins, in São Paulo state the method decribed by Taylor & Van Dyke (1985). (Cruz, 1974; Costa, 1995a, 2003). Terminology for frontal squamation follows Hoede- Taxonomic studies involving Leptolebias are man (1958), and the terminology for the cephalic uncommon. Most species of Leptolebias are rare in neuromast series follows Costa (2001). nature, and are strongly threatened with extinction The phylogenetic analysis follows the cladistic or may already be extinct (Costa, 2002b; the present methodology, using morphological characters, study). Consequently, they are poorly represented in described in recent studies (Costa, 1998, 2006a). Ter- ichthyological collections. In addition, the hypothesis minal taxa were all species of Leptolebias listed by of monophyly of Leptolebias is weakly supported. Lep- Costa (2003), except L. opalescens and L. leitaoi (see tolebias was formerly defined by the colour pattern in the Discussion). In order to test the hypothesis of females, consisting of a body homogeneously light monophyly of Leptolebias, two species representing brown without dark markings (Myers, 1952; Costa, different lineages of each of the remaining genera of 1990a). However, this female colour pattern is not cynopoeciline killifishes were included as terminal unique to Leptolebias, as it also occurs in some taxa: Campellolebias chrysolineatus Costa, Lacerda & aplocheiloid taxa both in South America and in Africa. Brazil, 1989 and C. dorsimaculatus, Cynopoecilus Leptolebias was also tentatively diagnosed by some intimus Costa, 2002, and Cynopoecilus nigrovittatus osteological features (Costa, 1990a, 1998), but the Costa, 2002. Outgroups follow Costa (2006a): Nema- hypothesis of monophyly was not confirmed in subse- tolebias whitei (Myers, 1942), a basal member of quent studies (e.g. Costa, 1998; the present study). the clade hypothesized to be the sister group to On the other hand, molecular data suggest that Lep- Campellolebias + Cynopoecilus + Leptolebias (Costa, tolebias may be paraphyletic. In a study on phyloge- 1998, 2006b); Neofundulus paraguayensis (Eigen- netic relationships of the Rivulidae (Murphy et al., mann & Kennedy, 1903), a basal member of an 1999), including three nominal species of Leptolebias annual fish clade (Costa, 2005); and Kryptolebias bra- and one of Campellolebias among several other siliensis (Valenciennes, 1821), a basal species of the rivulids, both maximum parsimony and neighbour- most basal lineage of the Rivulidae (Costa, 2004). joining analyses supported a closer relationship Characters and character states are listed and coded between the clade comprising L. aureoguttatus, in the Supplementary Material, Appendix S1, and are L. citrinipinnis, and Campellolebias dorsimaculatus plotted in the data matrix of Table S1. Character and Costa, Lacerda & Brazil, 1989, than to L. minimus. character-state numbers are separated by a point, following the terminology used in the Supplementary Material, Appendix S1. Character states of multistate characters were treated as ordered when possible. MATERIAL AND METHODS The most parsimonious cladogram, using the BandB The material is deposited at the following institu- algorithm, and the bootstrap analysis
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