Accepted by M.R. De Carvalho: 11 Mar. 2013; published: 3 May 2013
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3646 (3): 265–276
www.mapress.com/ zootaxa/
Article
265
http://dx.doi.org/10.11646/zootaxa.3646.3.5
http://zoobank.org/urn:lsid:zoobank.org:pub:064C4B43-8D79-49ED-8247-91D74F93FAAC
Monophyly of the Agoniatinae (Characiformes: Characidae)
FERNANDO C.P. DAGOSTA
1,2
& ALÉSSIO DATOVO
1,2
1
Laboratório de Ictiologia de Ribeirão Preto, USP, FFCLRP, Departamento de Biologia, PPG Biologia Comparada, Av. Bandeirantes
3900, 14040-901 Ribeirão Preto, São Paulo, Brasil. E-mail: ferdagosta@yahoo.com.br
2
Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42.494, CEP 04218-970, São Paulo, SP, Brasil.
E-mail: adatovo@gmail.com
Abstract
The Characidae is the most diverse family of Neotropical fishes, currently encompassing more than one thousand valid spe-
cies. Some subgroups within this family still lack phylogenetic definitions, being diagnosed on the basis of combination of
characters, a common procedure in pre-cladistic studies. Agoniatinae, currently composed by two valid species, Agoniates
anchovia and A. halecinus, is one of them. In the present study the Agoniatinae is redefined using a phylogenetically oriented
comparative survey that included the two Agoniates species plus 114 species representing all the major clades of the Charac-
idae and their closest relatives. Six derived morphological characters are identified as synapomorphies for the Agoniatinae:
deep notch on the posterior region of the maxilla joining the ventral margin of the infraorbital 2; dentigerous portion of pre-
maxilla shorter than the ascending process of this bone; dentary canine preceded by tricuspid teeth; ventral margin of the
urohyal markedly convex; base of the uppermost ray of the ventral lobe of the caudal fin much expanded, being as deep as
the distal margin of hypural 2; and levator arcus palatini muscle with a posterodorsal bundle of fibers attaching to the dorsal
face of the sphenotic spine.
Key words: Agoniates, morphology, osteology, myology, systematics
Introduction
The Characidae includes 1027 valid species (Eschmeyer and Fong 2012) distributed throughout inland waters of large
portions of the Americas. The family contains several subunits whose phylogenetic relationships have been differently
interpreted recently (Reis et al. 2003; Mirande 2010; Oliveira et al. 2011). Noteworthy is the conflicting results of the
two most recent comprehensive studies dealing with phylogenetic relationships of the Characidae (Mirande 2010;
Oliveira et al. 2011) producing remarkably divergent classifications.
The subfamily Agoniatinae is represented by the single genus Agoniates Müller and Troschel, originally erected
to allocate A. halecinus Müller and Troschel from Guiana. A second species, A. anchovia Eigenmann, was described
by Eigenmann (1914) based on specimens from Río Beni, upper Amazon River basin, Bolivia, collected by J. D.
Haseman. A third species was described by Géry (1963) as A. ladigesi, but Zarske and Géry (1997) synonymized this
species with A. anchovia. Agoniates halecinus is widespread in the Amazon basin, while A. anchovia apparently
occurs primarily in the upper Amazon (Lima and Zanata 2003). The genus Agoniates was allocated into the group
Tetragonopterina by Günther (1864) and partially based on the conclusions of Géry (1963), Uj (1990) raised
Agoniatinae to family rank. Lima and Zanata (2003) defined Agoniates as being the sole member of the Agoniatinae, a
similar arrangement proposed by Mirande (2010) and Oliveira et al. (2011). In the present study, we follow these
recent classifications in assigning a subfamilial rank for the genus.
The first authors to make evolutionary inferences about the genus were Cuvier and Valenciennes (1849) who
commented that Agoniates was intermediate between “Hydrocyns” and “Tétragonoptères”. Regan (1911) suggested
the possible close relationship between Agoniates and Salminus Agassiz. Eigenmann (1912) commented about the
similarity between Agoniates and Hydrolycus Müller and Troschel and Howes (1976) admitted that some features
shared by agoniatines and cynodontines might represent derived specializations. Géry (1963), in a pre-cladistic context