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Phylogeny of the Family Characidae (Teleostei: Characiformes): from Characters to Taxonomy

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Phylogeny of the Family Characidae (Teleostei: Characiformes): from Characters to Taxonomy Neotropical Ichthyology, 8(3):385-568, 2010 Copyright © 2010 Sociedade Brasileira de Ictiologia Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy Juan Marcos Mirande The family Characidae is the most diverse among Neotropical fishes. Systematics of this family are mainly based on pre- cladistic papers, and only recently a phylogenetic hypothesis for Characidae was proposed by the author. That phylogeny was based on 360 morphological characters studied for 160 species, including representatives of families related to Characidae. This paper is based on that phylogenetic analysis, with the analyzed characters described herein and documented, accompanied by comparisons of their definition and coding in previous papers. Synapomorphies of each node of the proposed phylogeny are listed, comparisons with previous classifications provided, and autapomorphies of the analyzed species listed. Taxonomic implications of the proposed classification and the position of the incertae sedis genera within Characidae are discussed. A discussion of the phylogenetic information of the characters used in the classical systematics of the Characidae is provided. La familia Characidae es la más diversa entre los peces neotropicales. La sistemática de esta familia está basada principalmente en trabajos pre-cladísticos, y sólo recientemente una hipótesis filogenética para Characidae fue propuesta por el autor. Esa filogenia estaba basada en 360 caracteres estudiados en 160 especies, incluyendo representantes de familias relacionadas a Characidae. Este trabajo está basado en ese análisis filogenético, y los caracteres analizados son aquí descriptos y documentados, comparando su definición y codificación con trabajos previos. Las sinapomorfías de cada nodo de la filogenia propuesta son listadas, y se incluyen comparaciones con clasificaciones previas; también se listan las autapomorfías de las especies analizadas. Las implicancias taxonómicas de la clasificación propuesta y la posición de los géneros incertae sedis dentro de Characidae son discutidas. También se presenta una discusión de la información filogenética de los caracteres usados en la sistemática clásica de Characidae. Key words: Ostariophysi, Osteology, Morphology, Classification, Systematics. Introduction Anostomidae, Characidae, Chilodontidae, Crenuchidae, Curimatidae, Ctenoluciidae, Cynodontidae, Erythrinidae, The order Characiformes includes more than 1,800 species, Gasteropelecidae, Hemiodontidae, Lebiasinidae, Parodontidae, of which the family Characidae is the most diverse, with Prochilodontidae, and Serrasalmidae) (Reis et al., 2003; approximately 1,200 species (Reis et al., 2003); indeed, the Calcagnotto et al., 2005), and one trans-Atlantic family Characidae is the fourth most diverse family of fishes, after the (Alestidae) (Zanata & Vari, 2005). The families Citharinidae Cyprinidae, Cichlidae and Gobiidae (Eschmeyer & Fricke, 2009). and Distichodontidae constitute the suborder Citharinoidei, Members of the Characidae occur from southern portions of considered as the sister group of the Characoidei, which the USA to northern Patagonia in Argentina, being especially includes all the remaining Characiformes (Vari, 1979; Fink & diverse in the Amazon, Orinoco, and La Plata River basins. Fink, 1981, 1996; Buckup, 1998; Calcagnotto et al., 2005). According to the currently accepted phylogenetic Among the Characoidei, the monophyly of a clade composed hypotheses, based on both morphological and molecular data, of the families Anostomidae, Chilodontidae, Curimatidae, and the Cypriniformes constitute the sister group of Prochilodontidae (Anostomoidea; Vari, 1983; Buckup, 1998), (Characiformes (Siluriformes + Gymnotiformes)) (Fink & Fink, and a clade formed by the Neotropical families Ctenoluciidae, 1981, 1996; Dimmick & Larson, 1996). According to Erythrinidae and Lebiasinidae and the African family classifications prior to Mirande (2009), the Characiformes Hepsetidae (Erythrinoidea; Vari, 1995; Buckup, 1998) had been consisted of three African families (Citharinidae, proposed. Relationships between these suprafamilial groups Distichodontidae and Hepsetidae) (Géry, 1977, Calcagnotto and the remaining Characiformes were unclear, and some et al., 2005), 14 Neotropical families (Acestrorhynchidae, hypotheses that conflicted in varying degrees were proposed CONICET-Fundación Miguel Lillo, Miguel Lillo 251, 4000 San Miguel de Tucumán, Argentina. [email protected] 385 386 Phylogeny of the family Characidae (Uj, 1990; Ortí & Meyer, 1997; Buckup, 1998; Calcagnotto Fink, 1985; Weitzman & Menezes, 1998), Cheirodontinae et al., 2005, Hubert et al., 2005). (Malabarba, 1998a) and Paracheirodon Géry (Weitzman & Most families of the Characiformes have evidences of Fink, 1983), Charax Scopoli (Lucena, 1987), Jupiaba Zanata monophyly (Weitzman, 1954; Roberts, 1973, 1974; Vari, 1979, (Zanata, 1997), Roestes Günther and Gilbertolus Eigenmann 1983, 1995; Buckup, 1998; Toledo-Piza, 2000; Zanata & Vari, (Lucena & Menezes, 1998), Spintherobolus Eigenmann 2005), whereas there are no consensus on the monophyly (Weitzman & Malabarba, 1999), Creagrutus Günther and and composition of the Characidae. Most currently Piabina Reinhardt (Vari & Harold, 1998, 2001), Deuterodon recognized subfamilial and generic groups in the Characidae Eigenmann (Lucena & Lucena, 2002), Cyanocharax are based on the pre-cladistic papers of Eigenmann (e. g. 1912, Malabarba & Weitzman (Malabarba & Weitzman, 2003), 1915, 1917, 1918, 1921, 1927) and Eigenmann & Myers (1929). Attonitus Vari & Ortega (Vari & Ortega, 2000), and Eigenmann (1917) was highly influential in terms of our Bryconadenos Weitzman, Menezes, Evers & Burns present concepts of relationships within the Characidae. (Weitzman et al., 2005)]. Most of these papers were focused Eigenmann defined 17 characters with discrete alternative on particular groups of the Characidae, without enough states, and used them in different combinations to diagnose exploration of their relationships with the remaining the genera in the Characidae, considering the most frequent Characidae. Malabarba (1998a) restricted the subfamily states as being primitive. The genus Astyanax Baird & Girard Cheirodontinae to a subset of the genera recognized in this has the combination of the most frequent states of all these subfamily by Géry (1977), leaving 33 genera as incertae characters, and it was consequently considered by Eigenmann sedis. Later, Reis (2003a) restricted the Tetragonopterinae (1917) as primitive within the family. Eigenmann, however, to its type genus Tetragonopterus Cuvier, leaving many recognized that the less frequent states could have genera as incertae sedis within the Characidae. Lima et al. independent origins in different species of the same genus, (2003) classified also several genera previously included in producing “polyphyletic” [sic] genera. Given the impossibility the subfamilies Bryconinae, Characinae, Cheirodontinae, to classify the Characidae in a branching scheme, Eigenmann and Paragoniatinae (Géry, 1977) as incertae sedis within the (1917) presented a radial pattern, identifying a “nucleus” of Characidae. The subfamilies and incertae sedis genera generalized morphology (represented by the genus Astyanax) recognized in the last revision of the Characidae are as follows and different lines of evolution diverging from it. (number of genera in each group in parentheses): incertae Eigenmann’s classification was followed by Greenwood sedis (88) (Lima et al., 2003), Agoniatinae (1) (Lima & Zanata, et al. (1966) and particularly by Géry (e. g. Géry, 1977). Géry 2003), Clupeacharacinae (1) (Lima, 2003a), Iguanodectinae also recognized the polyphyletic nature of this classification, (2) (Moreira, 2003), Bryconinae (3) (Lima, 2003b), and that this systematic scheme failed to reflect the phylogeny Serrasalminae (15) (Jégu, 2003), Aphyocharacinae (1) (Lima, (Géry, 1972). In this pre-cladistic systematic classification, 2003c), Characinae (12) (Lucena & Menezes, 2003), most genera of the Characidae, especially those with Stethaprioninae (4) (Reis, 2003a), Tetragonopterinae (1) “generalized” morphology, were included in the subfamily (Reis, 2003b), Rhoadsiinae (3) (Cardoso, 2003a), Tetragonopterinae. The remaining genera were distributed Cheirodontinae (15) (Malabarba, 2003), and across several subfamilies defined by the presence of Glandulocaudinae (19) (Weitzman, 2003). The subfamily somewhat arbitrarily chosen characters. Géry (1977), following Rhaphiodontinae (sensu Géry, 1977) was included into the the general classification of Eigenmann recognized the Cynodontidae (Lucena & Menezes, 1998; Toledo-Piza, 2003), subfamilies (number of genera in parentheses) Agoniatinae and all the genera assigned to the Paragoniatinae by Géry (1), Rhaphiodontinae (2), Characinae (14), Bryconinae (6), (1977) were included in the incertae sedis-group (Lima et Clupeacharacinae (1), Paragoniatinae (6), Aphyocharacinae (1), al., 2003). The Acestrorhynchidae was considered as a Glandulocaudinae (18), Stethaprioninae (3), Tetragonopterinae separate family (Lucena & Menezes, 1998, 2003). (49), Rhoadsiinae (2) and Cheirodontinae (13 genera sensu Malabarba & Weitzman (2003) described Cyanocharax stricto and 36 sensu lato). and proposed the monophyly of a group of genera (their clade The first genus of the Characidae explicitly diagnosed by A) including all members of Glandulocaudinae plus several shared presumably apomorphic features was Bramocharax incertae sedis genera.
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