Neotropical Ichthyology, 3(1):1-60, 2005 Copyright © 2005 Sociedade Brasileira de Ictiologia Systematics of the subfamily Poeciliinae Bonaparte (Cyprinodontiformes: Poeciliidae), with an emphasis on the tribe Cnesterodontini Hubbs Paulo Henrique Franco Lucinda* and Roberto E. Reis** Osteological and soft anatomical features of representatives of poeciliine genera were studied to test the monophyly of the poeciliine tribes and to advance a hypothesis of relationships within the subfamily. The resultant hypothesis supports the proposal of a new classification for the subfamily Poeciliinae. Diagnoses are provided for suprageneric clades. The tribe Tomeurini is resurrected and the new tribes Brachyrhaphini and Priapichthyini as well as the supertribe Poeciliini are described. New usages of old tribe names are proposed based on the phylogenetic framework. Caracteres osteológicos e da anatomia mole de representantes dos gêneros de poeciliíneos foram estudados para se testar a monofilia das tribos de Poeciliinae e para propor uma hipótese de relações dentro da subfamília. A hipótese resultante suporta a proposição de uma nova classificação para a subfamília Poeciliinae. São fornecidas diagnoses para os clados supragenêricos. A tribo Tomeurini é ressuscitada e as novas tribos Brachyrhaphini e Priapichthyini bem como a supertribo Poeciliini são descritas. Novos usos para antigos nomes de tribos são propostos com base no arranjo filogenético. Key words: Alfarini, Brachyrhaphini, Gambusiini, Girardinini, Heterandriini, Priapellini, Priapichthyini, Poeciliini, Tomeurini. Introduction Nomenclatural and Taxonomic History This paper is resultant from a project that intended to Poeciliinae. The subfamily Poeciliinae is a cyprinodontiform perform the taxonomic revision of the tribe Cnesterodontini, group widely distributed throughout the Americas. Poeciliinae as well as to propose a phylogenetic hypothesis of rela- is the sister group of the Procatopodinae, a group composed tionships among its members. Testing the monophyly of of the South-American Fluviphylax Whitley and the African the Cnesterodontini was the first step attempting to achieve procatopodines. The clade Poeciliinae plus Procatopodinae our aims. Thus, the original project was broadened to em- is the sister group of the Aplocheilichthyinae (Costa, 1996; brace a phylogenetic hypothesis of relationships and the Ghedotti, 2000). These three subfamilies compose the family proposal of a provisional classificationPROOFS for the subfamily Poeciliidae. The Poeciliinae embraces approximately two hun- Poeciliinae. The resultant hypothesis included representa- dred twenty species currently allocated in approximately tives of all poeciliine genera, and all described twenty-eight genera (Lucinda, 2003). Theses fishes are char- Cnesterodontini species, as well as 24 new species revealed acterized by the uniquely derived possession of a by the taxonomic revisions of genera Cnesterodon Garman, gonopodium formed by the modified male anal-fin rays 3, 4, Phallotorynus Henn, and Phalloceros Eigenmann. and 5 (Parenti, 1981). Intrageneric relationships of Phalloptychus Eigenmann, The Poeciliinae includes well-known aquarium fishes Cnesterodon, Phalloceros, and Phallotorynus are provided such as the guppies, mosquito fishes, swordtails, platys, herein, but will be discussed in the aforementioned taxo- and mollies. Poeciliines are well known subjects of study for nomic revisions. ecologists, anatomists, embryologists, and other research- * Laboratório de Ictiologia Sistemática, Universidade Federal do Tocantins, Campus de Porto Nacional, rua 3, Quadra 17, s/n, Caixa Postal 136, 77500-000 Porto Nacional, TO, Brazil. e-mail:[email protected] ** Laboratório de Ictiologia, Pontifícia Universidade Católica do Rio Grande do Sul. Av. Ipiranga, 6681, Caixa Postal 1429, 90619-900 Porto Alegre, RS, Brazil. e-mail: [email protected] 1 2 Systematics of the subfamily Poeciliinae Bonaparte ers. Notwithstanding, this fish assemblage is only superfi- Table 2. Classification of Poeciliinae (Ghedotti, 2000) cially studied from the perspective of systematics. Intrageneric diversity and intergeneric relationships of the Subfamily Poeciliinae Bonaparte, 1831 Poeciliinae are poorly known, regardless of its huge distri- Tribe Poeciliini Bonaparte, 1831 bution, range, and notoriety. Similarly, phylogenetic hypoth- Poecilia, Xiphophorus, Phallichthys Tribe Alfarini Hubbs, 1924 eses for most genera are still wanting. Alfaro Taxonomic revisions and phylogenetic analyses have Tribe Cnesterodontini Hubbs, 1924 provided some insight into the relationships of smaller Cnesterodon, Phalloceros, Phalloptychus, Tomeurus, groups of the Poeciliinae (e.g., Rosen, 1967, 1979; Phallotorynus, Rauchenberger, 1989; Rosa & Costa, 1993; Meyer et al., Tribe Gambusiini Gill, 1893 1994; Mojica et al., 1997; Rodriguez, 1997; Poeser, 2003) or Gambusia, Belonesox, Brachyrhaphis have dealt with higher taxa (Rosen, 1964; Parenti, 1981; Costa, Tribe Girardinini Hubbs, 1924 1996, 1998; Ghedotti, 2000). The only comprehensive study Girardinus, Quintana, Carlhubbsia is the classic revision of “Poeciliidae” by Rosen & Bailey Tribe Heterandriini Hubbs, 1924 (1963), which did not deal with cladistic methodology. None- Heterandria, Priapichthys, Poeciliopsis, Neoheterandria theless, Rosen & Bailey (1963) is the basis for the current Tribe Xenodexini Hubbs, 1950 Xenodexia internal classification of Poeciliinae. Later, Parenti & Tribe Scolichthyini Rosen, 1967 Rauchenberger (1989) modified the classification of Rosen Scolichthys & Bailey (1963) in order to accommodate it into the taxo- Tribe Priapellini Ghedotti, 2000 nomic rank of subfamily proposed by Parenti (1981) (Table Priapella 1). Following Rosen & Bailey (1963) and Parenti & Rauchenberger (1989), Tomeurus Eigenmann alone is the Rafinesque (1810) created the “Ordini Cyprinidi”; this supertribe Tomeurini. The remaining genera form the group contained a species of Mugil Linnaeus and three spe- supertribe Poeciliini, which is subdivided in the tribes cies of Cyprinus Linnaeus. Later Wagner (1828) created the Poeciliini, Cnesterodontini, Gambusiini, Scolichthyini, family “Cyprinöidae” including the genera Lebias Goldfuss, Girardinini, Heterandriini, and Xenodexini. Later, Ghedotti Poecilia Bloch & Schneider, Fundulus Lacepède, Cyprinodon (2000) proposed another classification for the Poeciliinae Lacepède, and Mollienesia Lesueur. The word “Cyprinöidae” (Table 2) based in his phylogenetic study of the Poecilioidea is etymologically identical to “Cyprinidi”, for they share the despite the fact that only 12 genera were examined in his same stem. Thus, Wagner failed to coin a new name. Cuvier cladistic analysis. (1829) and Cuvier & Valenciennes (1846) followed Rafinesque The history of the subfamily began in 1801 with the de- (1810) employing the name “Cyprinöides” and “Cyprins”. scription of Poecilia vivipara Bloch & Schneider as new Bonaparte (1831) also accepted the “Ordini Cyprinidi” of genus and new species. Before establishment as a distinct Rafinesque (regarding it as a family), but divided it in three family, the history of the Poeciliidae is merged with that of groups: Cyprinini, Anableptini, and Poecilini [sic]. The other cyprinodontiform families and with the Cyprinidae. Poecilini was removed from the Cyprinidae by Bonaparte (1840) to form a separate family Poecilidae [sic] (including Table 1. Classification of Poeciliinae (Rosen & Bailey, 1963 Anableptini and Poecilini). Four years before Bonaparte, and Parenti & Rauchenberger, 1989).PROOFSAgassiz (1834) had already separated the Cyprinodonts and the Cyprinidae, removing Anableps Scopoli, Poecilia, Lebias, Subfamily Poeciliinae Bonaparte, 1831 Fundulus, Mollienesia, and Cyprinodon to a new family Supertribe Tomeurini Eigenmann, 1912 Tomeurus “Cyprinodontes”. Swainson (1838) placed Poeciliinae as a Supertribe Poeciliini Bonaparte, 1831 subfamily of Cobitidae. Gill (1857) adopted the name Tribe Poeciliini Garman, 1895 Cyprinodontes and later (Gill, 1865; 1894) employed Poecilia, Xiphophorus, Alfaro, Priapella Poeciliidae. Gill (1872) also used the name Cyprinodontidae Tribe Cnesterodontini Hubbs, 1924 as a subgroup of the Haplomi, which also included the Cnesterodon, Phalloceros, Phalloptychus, Phallotorynus. Amblyopsidae, Esocidae, and Umbridae. Günther (1866) cre- Tribe Scolichthyini Rosen, 1967 ated the genus Platypoecilus Günther and adopted Scolichthys Cyprinodontidae as family name, dividing it in Cyprinodontidae Tribe Gambusiini Gill, 1893 Limnophagae (=poeciliines) and Cyprinodontidae Carnivorae Gambusia, Belonesox, Brachyrhaphis Tribe Girardinini Hubbs, 1924 (=remaining Cyprinodontiformes). Girardinus, Quintana, Carlhubbsia. At the same time, new genera have been proposed: Tribe Heterandriini Hubbs, 1924 Mollienesia, Xiphophorus Heckel and Heterandria Heterandria, Poeciliopsis, Priapichthys, Neoheterandria, Agassiz. Poey (1854) added three new genera: Girardinus Phallichthys Poey, Limia Poey, and Gambusia Poey. Belonesox Kner Tribe Xenodexini Hubbs, 1950 was created in 1860. In the same year the genera Xenodexia Hemixiphophorus Bleeker and Pseudoxiphophorus Bleeker P. H. F. Lucinda & R. E. Reis 3 were erected. Soon after, De Filippi (1861) created Lebistes Hubbs, Parapoecilia Hubbs, and Xenophallus Hubbs, and De Filippi and Steindachner (1863) erected Poeciliodes recognized four subfamilies (Tomeurinae, Gambusiinae, Steindachner. Poeciliinae, and Poeciliopsinae) and nine tribes within the Garman (1895) adopted the family name Cyprinodontes