“main” — 2011/5/11 — 19:20 — page 483 — #1 Anais da Academia Brasileira de Ciências (2011) 83(2): 483-511 (Annals of the Brazilian Academy of Sciences) Printed version ISSN 0001-3765 / Online version ISSN 1678-2690 www.scielo.br/aabc Taxonomic review and phylogenetic analysis of Enchodontoidei (Teleostei: Aulopiformes) HILDA M.A. SILVA and VALÉRIA GALLO Laboratório de Sistemática e Biogeografia, Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Rua São Francisco Xavier, 524, Maracanã, 20550-013 Rio de Janeiro, RJ, Brasil Manuscript received on September 24, 2010; accepted for publication on December 22, 2010 ABSTRACT Enchodontoidei are extinct marine teleost fishes with a long temporal range and a wide geographic distribution. As there has been no comprehensive phylogenetic study of this taxon, we performed a parsimony analysis using a data matrix with 87 characters, 31 terminal taxa for ingroup, and three taxa for outgroup. The analysis produced 93 equally parsimonious trees (L = 437 steps; CI = 0.24; RI = 0.49). The topology of the majority rule consensus tree was: (Sardinioides + Hemisaurida + (Nardorex + (Atolvorator + (Protostomias + Yabrudichthys) + (Apateopholis + (Serrilepis + (Halec + Phylactocephalus) + (Cimolichthys + (Prionolepis + ((Eurypholis + Saurorhamphus) + (Enchodus + (Paleolycus + Parenchodus))))))) + ((Ichthyotringa + Apateodus) + (Rharbichthys + (Trachinocephalus + ((Apuliadercetis + Brazilodercetis) + (Benthesikyme + (Cyranichthys + Robertichthys) + (Dercetis + Ophidercetis)) + (Caudadercetis + (Pelargorhynchus + (Nardodercetis + (Rhynchodercetis + (Dercetoides + Hastichthys)))))). The group Enchodontoidei is not monophyletic. Dercetidae form a clade supported by the presence of very reduced neural spines and possess a new composition. Enchodontidae are monophyletic by the presence of middorsal scutes, and Rharbichthys was excluded. Halecidae possess a new composition, with the exclusion of Hemisaurida. This taxon and Nardorex are Aulopiformes incertae sedis. Key words: Aulopiformes, Enchodontoidei, phylogeny, taxonomy. INTRODUCTION 1996, Fielitz 2004, Figueiredo and Gallo 2006, Gallo et Enchodontoidei are extinct marine teleosts generally al. 2006). with an elongate body and long and narrow rod-like The taxon was erected by Berg (1937) as a sub- maxilla included in the mouth gape (Nelson 1994). They order, which included only the family Enchodontidae. possess a long temporal range, extending from the Early According to this author, enchodontid fishes were sim- Cretaceous to the Early Eocene, and a wide geographic ilar to the members of the suborder Stomiatoidei also distribution in sedimentary deposits of South America created by him, but the enchodontids bear a median row (e.g., Bolivia and Brazil), Africa (e.g., Democratic of dorsal scutes and their vertebrae do not possess para- Republic of Congo, Egypt, and Morocco), Asia (e.g., pophyses. Goody (1969) accomplished a comprehensive Arabian Peninsula, India, Israel, Japan, and Lebanon), review of certain Late Cretaceous teleosteans, consider- Europe (e.g., Belgium, England, Germany, Holland, ing Enchodontoidei as part of the order Salmoniformes, Italy, and Sweden), and North America (Canada, together with three other suborders, Ichthyotringoidei, Mexico, and United States) (e.g., Goody 1969, Chalifa Cimolichthyoidei, and Halecoidei. Rosen (1973) erected the order Aulopiformes com- Correspondence to: Hilda Maria Andrade da Silva E-mail: [email protected] prising 15 living families, and the suborder Alepisau- An Acad Bras Cienc (2011) 83 (2) “main” — 2011/5/11 — 19:20 — page 484 — #2 484 HILDA M.A. SILVA and VALÉRIA GALLO roidei with 15 fossil genera, without dividing them sys- Enchodontidae. The extant aulopiforms were classified tematically (i.e., Ichthyotringa, Apateodus, Apateopho- in Synodontoidei (with four families), Chlorophthalmoi- lis, Cimolichthys, Dercetis, Rhynchodercetis, Pelargo- dei (with six families), and Giganturoidei (with two fam- rhynchus, Prionolepis, Enchodus, Palaeolycus, Eury- ilies). Additionally, four living families were placed in pholis, Saurorhamphus, Halec, Phylactocephalus, and the suborder Alepisauroidei. Hemisaurida), as well as the fossil genera incertae sedis In fact, the assemblage of extinct aulopiforms de- of the superfamily Synodontoidea, Sardinius and fined by Nelson (2006) corresponds to Enchodontoidei Volcichthys. sensu Nelson (1994). However, Nelson (2006) did not Nelson (1994) recognized the order Aulopiformes discuss his reasons for disregarding the name Encho- by Rosen (1973), as well as the suborders proposed by dontoidei and put its members in Alepisauroidei, Goody (1969) as superfamilies, putting them in a single Ichthyotringoidei, and Halecoidei. Moreover, Encho- suborder, Enchodontoidei, composed of four superfam- dontoidei were considered in the cladistic analysis of ilies: Enchodontoidea (Enchodus, Parenchodus, Palae- Dercetidae (Gallo et al. 2005), as well as in a prelimi- olycus, Eurypholis, and Saurorhamphus), Cimolichthy- nary approach by Silva and Gallo (2007). As there oidea (Cimolichthys, Prionolepis, Benthesikyme, Cyra- has not been a recent comprehensive phylogenetic study nichthys, Dercetis, Dercetoides, Pelargorhynchus, Rhyn- of Enchodontoidei, we review their classification his- chodercetis, and Stratodus), Halecoidea (Halec, Hemi- tory and provide a new cladistic analysis. saurida, and Phylactocephalus) and Ichthyotringoidea (Ichthyotringa and Apateodus). However, this classifi- SYSTEMATIC HISTORY OF ENCHODONTOIDEI cation was not developed in a phylogenetic framework. In this paper we use the general classification of Nel- Baldwin and Johnson (1996) accomplished a cla- son (1994), except for the family Enchodontidae (sensu distic analysis of Aulopiformes, including only extant Fielitz 2004). taxa. The authors maintained the monophyly of the taxon and added synapomorphies to those proposed by SUPERFAMILY ICHTHYOTRINGOIDEA Rosen (1973), which are mainly related to the morphol- According to Goody (1969), Ichthyotringoidea com- ogy of the dorsal portion of the gill arches. Their new prises two closely related families, Ichthyotringidae synapomorphies are from the intermuscular system, in- and Apateopholidae. The author considered mainly ternal soft anatomy, pigmentation pattern of larvae, and primitive features of the body and caudal skeleton, as morphology of the pelvic girdle. Most of these features well as a derived feature related to the rostral region. He are very difficult to assess in fossil specimens. stated that, despite the similarities shared by the taxa, Sato and Nakabo (2002) accomplished a phyloge- Apateopholidae should be the more advanced taxon. netic analysis of living Aulopiformes based on morpho- logical and molecular data. They divided it into the Family Ichthyotringidae. The family Ichthyotringidae suborders Synodontoidei, Chlorophthalmoidei, Alepi- (Table I) was created by Jordan (1905) to contain a sauroidei, and Giganturoidei. Moreover, the authors pro- single genus (i.e., Ichthyotringa). Goody (1969) po- posed a new family of Aulopiformes (i.e., Paraulopidae). sitioned the family in the suborder Ichthyotringoidei. Fielitz (2004) and Gallo et al. (2005) proposed Later, Nelson (1994) included the taxon in the suborder hypotheses of the phylogenetic relationships of some Enchodontoidei together with other fossil aulopiforms. fossil Aulopiformes (Enchodontoidea and Dercetidae, The generic epithet Ichthyotringa was created by Cope respectively). (1878) to replace the genus Rhinellus of Agassiz (1844), Nelson (2006) placed the extinct aulopiforms in which was pre-occupied. The genus Ichthyotringa in- three suborders: Ichthyotringoidei, comprising the fam- cludes the following species: Ichthyotringa furcata ilies Ichthyotringidae, Dercetidae, and Prionolepidae; (Agassiz, 1844), I. tenuirostris Cope, 1878, I. damoni Halecoidei, with a single family, Halecidae; and Ale- (Davis, 1887), I. ferox (Davis, 1887), I. delicata (Hay, pisauroidei, with two families, Cimolichthyidae and 1903), and I. africana (Arambourg, 1954). Forey et An Acad Bras Cienc (2011) 83 (2) “main” — 2011/5/11 — 19:20 — page 485 — #3 ENCHODONTOIDEI SYSTEMATICS 485 al. (2003), in the general list of fossil fishes from a great similarity regarding the structures of the skull Lebanon, placed Apateopholis in the family Ichthyo- and body, especially in the rostral region. Moreover, tringidae, but remarked that at least one species of general body squamation is lacking and two or three Apateopholis is often misinterpreted as a species of the rows of isolated scutes are present on the flanks. Re- closely related Ichthyotringa. Only more recently a new garding the family Prionolepididae, the author pointed ichthyotringoidei was reported to the El Doctor Forma- out some problems concerning its taxonomic place- tion in the Albian-Cenomanian of Mexico, I. mexicana ment. Unlike Woodward (1901), who assigned the genus Fielitz and González Rodríguez, 2008. Prionolepis to the Enchodontidae, Goody (1969) consid- Goody (1969) ranked Apateodus as an addendum ered it closely related to dercetids and cimolichthyids. (incertae sedis) to the Ichthyotringidae with a single Family Cimolichthyidae. The family (Table II) was species (A. striatus Woodward, 1901). Nelson (1994, erected by Goody (1969) to include the single genus 2006) placed Apateodus in Apateopholidae, but he Cimolichthys, which was designated by Leidy (1857) did not report the taxonomic status of Apateopholis. with