Doradinae Bleeker, 1858

Doradini Bleeker, 1858: 39 (see also Bleeker, 1858: 48-52) [type genus Doras].

Included taxa. Anduzedoras, Centrochir, Centrodoras, Doraops, Doras, Hassar, Hemidoras, Leptodoras, Lithodoras, Megalodoras, Nemadoras, Orinocodoras, Ossancora, Oxydoras, Platydoras, Pterodoras, Rhinodoras, Rhynchodoras, Tenellus, and Trachydoras.

Diagnosis. The subfamily Doradinae is diagnosed by the following characters: tips of terminal diverticula of the gas bladder ditally divergent (#36: 0>1, present only in Doras zuanoni, Ossancora punctata, and Trachydoras, coded as missing data for the remaining species of Doradinae, which lack terminal diverticula), first free vertebrae eighth or nineth (#217: 1>3, modified in Doraops, Lithodoras, Megalodoras, Pterodoras, and also present in some non-Doradidae catfishes), serrations present on the posterior face of the dorsal-fin spine (#245: 0>1, reserved in some species of Leptodoras, present also in some non-Doradidae catfishes), transverse coracoid crest between adductor superficialis and arrector ventralis muscles (#274: 0>1, reserved in Doraops, Lithodoras, Oxydoras, Pterodoras, and fimbriate-barbel doradids [except Hemidoras morrisi, and Ossancora fimbriata], also present in Acanthodoras), eight branched rays in the ventral lobe of the caudal fin (#306: 0>1, also present in some non-Doradidae catfishes), hypurals 1 and 2 fused, parahypural distinct (#310: 2>1, reversed in Rhinodoras thomersoni, present also in Franciscodoras, Kalyptodoras, and Werthemeria).

Remarks. The present analysis did not recover Centrodoras as monophyletic, however, additional study including yet undescribed species of Centrodoras (Sabaj Pérez, pers. comm.) is necessary to establish the status of the genus.

Two major changes in the classification of Doradidae were made based on the present hypothesis of relationships among species of Doradinae: description of Tenellus as a new genus, and inclusion of the species previously placed in Opsodoras Eigenmann, 1925 within an expanded Hemidoras Bleeker, 1858. Hemidoras was defined by Eigenmann (1925), and subsequently followed by other authors, as comprising two species, H. stenopeltis and H. morrisi, both sharing dermal scutes between dorsal and adipose fins (#34). However, other characters analyzed herein indicate that Hemidoras morrisi is closely related to Opsodoras morei (now Hemidoras morei), a hypothesis corroborated by the similarity in the overall morphology of both species. Noteworthy to say, some specimens of Hemidoras morei (including the holotype, NMW 45433) have a single dermal plate between dorsal and adipose fins, a partial representation of the feature formerly used to distinguish Hemidoras from Opsodoras.

As discussed by Ferraris (2007: 164), the suprageneric names Centrochirinae, Hemidoradinae, and Lithodoradinae were proposed by Fowler (1951: 3) in a manner that made the names available when published, but the names were rendered unavailable in 1961 by Art. 13 of the Second Edition of the Code (ICZN, 1961). The names apparently remain unavailable, despite Art. 13.2.1 of the current version of the Code (ICZN, 1999), because they were treated as unavailable in Ferraris & de Pinna (1999: 8).

Within Doradinae, four major monophyletic groups were recovered in the present (Fig. 69). Clade 1: the Platydoras clade also includes Centrochir and is supported by two characters: a distinct dark longitudinal stripe on the middle of each caudal-fin lobe (#1: 0>1) (#1: 0>1, also present in Anadoras, Orinocodoras, Tenellus, and most Leptodoras), and the secondary chamber of the gas bladder formed by a single posterior chamber, thus without an inner septum (#38: 0>1, also present in Doras and Oxydoras, coded as missing data for Centrochir and Platydoras sp., which lack the secondary chamber). Clade 2: the Pterodoras clade (or largedoradids clade) also includes Centrodoras, Doraops, Lithodoras, and Megalodoras and is supported by two characters: large diverticula present on the main bladder and on the secondary chamber of the gas bladder (#41: 1>2, exclusive), 12 or more ribs (#221: 1>2, also present in Kalyptodoras, Oxydoras, and Wertheimeria). Clade 3: the Rhinodoras clade also includes Orinocodoras and Rhynchodoras and is supported by five characters: five or six pores on the mandibular canal (#113: 1>2, also present in Astrodoradinae, Centrochir, Franciscodoras, Kalyptodoras, Lithodoras, Platydoras, Pterodoras, and Wertheimeria, coded as missing data for Rhynchodoras), posterior limit of autopalatine approximately at vertical through the middle of orbit (#149: 0>1, exclusive), hyomandibular crest for Aw section of the adductor mandibulae muscle present (#156: 0>1, exclusive), posterior cleithral process triangular (#266: 0>1, exclusive), procurrent rays of the caudal fin plate-like (#301: 0>1, also present in Agamyxis, Francisodoras and Platydoras). Clade 4: the Doras / Oxydoras clade also includes Anduzedoras, Hassar, Hemidoras, Leptodoras, Nemadoras, Ossancora, Tenellus, and Trachydoras, and is supported by 18 characters, including among others: mesethmoid medially dilated in the shape of an arrow (#52: 0>2, exclusive), anteroventral keel of mesethmoid present (#55: 0>1, exclusive), premaxilla conical with a dorsal apex, depth distinctly greater than width (#125: 0>1, exclusive), ventral hypohyal quadrangular, length and width approximately equal (#172: 0>1, exclusive), conspicuous accessory lamellae on the medial face of the ceratobranchials (#179: 0>1, reversed in Trachydoras), conspicuous accessory lamellae on the medial face of the epibranchials (#180: 0>1, reversed in Trachydoras), ventral process of fourth basibranchial elongate (#188: 0>1, exclusive), pharyngobranchial tooth plate elongate, length about twice width (#199: 0>1, exclusive), anterior margin of pectoral girdle anteriorly elongate with truncate or acute tip (#249: 0>1, reversed in Ossancora, Trachydoras, and some species of Leptodoras), dorsoposterior border of coracoid extremely elevated dorsally (#271: 0>1, exclusive). The Doras / Oxydoras clade includes the Doras clade, which is herein recognized as composed by the fimbriate-barbel doradids, the most well supported species group of the family with 17 characters, three of them exclusive (see Appendix).