Taxonomic review and phylogenetic analysis of Enchodontoidei (Teleostei: Aulopiformes)

EARTH SCIENCES

Taxonomic review and phylogenetic analysis of Enchodontoidei (Teleostei: Aulopiformes)

Hilda M. A. Silva; 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

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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.

RESUMO

Os Enchodontoidei são peixes teleósteos marinhos extintos, com uma longa amplitude temporal e uma ampla distribuição geográfica. Tendo em vista que não há nenhuma proposta ampla para a filogenia deste táxon, foi realizada uma análise de parcimônia com base numa matriz de dados de 87 caracteres, 31 táxons terminais no grupo interno, e três táxons no grupo externo. Como resultado da análise, foram obtidas 93 árvores igualmente parcimoniosas (L = 437 passos; CI = 0,24; RI = 0,49). O consenso de maioria é representado pela seguinte topologia: (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 )))))). O grupo Enchodontoidei não é monofilético. Os Dercetidae formam um clado suportado pela presença de espinhos neurais muito reduzidos e possuem uma nova composição. Os Enchodontidae são um grupo monofilético, devido à presença de escudos no dorso, e Rharbichthys foi excluído do clado. Os Halecidae possuem uma nova composição, com a exclusão de Hemisaurida. Este táxon e Nardorex são Aulopiformes incertae sedis.

Palavras-chave: Aulopiformes, Enchodontoidei, filogenia, taxonomia.

INTRODUCTION

Enchodontoidei are extinct marine teleosts generally with an elongate body and long and narrow rod-like maxilla included in the mouth gape (Nelson 1994). They possess a long temporal range, extending from the Early Cretaceous to the Early Eocene, and a wide geographic distribution in sedimentary deposits of South America (e.g., Bolivia and Brazil), Africa (e.g., Democratic Republic of Congo, Egypt, and Morocco), Asia (e.g., Arabian Peninsula, India, Israel, Japan, and Lebanon), Europe (e.g., Belgium, England, Germany, Holland, Italy, and Sweden), and North America (Canada, Mexico, and United States) (e.g., Goody 1969, Chalifa

1996, Fielitz 2004, Figueiredo and Gallo 2006, Gallo et 1996, Fielitz 2004, Figueiredo and Gallo 2006, Gallo et al. 2006).

The taxon was erected by Berg (1937) as a suborder, which included only the family Enchodontidae. According to this author, enchodontid fishes were similar to the members of the suborder Stomiatoidei also created by him, but the enchodontids bear a median row of dorsal scutes and their vertebrae do not possess parapophyses. Goody (1969) accomplished a comprehensive review of certain Late Cretaceous teleosteans, considering Enchodontoidei as part of the order Salmoniformes, together with three other suborders, Ichthyotringoidei, Cimolichthyoidei, and Halecoidei

Rosen (1973) erected the order Aulopiformes comprising 15 living families, and the suborder Alepisauroidei with 15 fossil genera, without dividing them systematically (i.e., Ichthyotringa, Apateodus, Apateopholis, Cimolichthys, Dercetis, Rhynchodercetis, Pelargorhynchus, Prionolepis, Enchodus, Palaeolycus, Eurypholis, Saurorhamphus, Halec, Phylactocephalus, and Hemisaurida ), as well as the fossil genera incertae sedis of the superfamily Synodontoidea, Sardinius and Volcichthys.

Nelson (1994) recognized the order Aulopiformes by Rosen (1973), as well as the suborders proposed by Goody (1969) as superfamilies, putting them in a single suborder, Enchodontoidei, composed of four superfamilies: Enchodontoidea (Enchodus, Parenchodus, Palaeolycus, Eurypholis, and Saurorhamphus), Cimolichthyoidea (Cimolichthys, Prionolepis, Benthesikyme, Cyranichthys, Dercetis, Dercetoides, Pelargorhynchus, Rhynchodercetis, and Stratodus ), Halecoidea (Halec, Hemisaurida, and Phylactocephalus ) and Ichthyotringoidea (Ichthyotringa and Apateodus ). However, this classification was not developed in a phylogenetic framework.

Baldwin and Johnson (1996) accomplished a cladistic analysis of Aulopiformes, including only extant taxa. The authors maintained the monophyly of the taxon and added synapomorphies to those proposed by Rosen (1973), which are mainly related to the morphology of the dorsal portion of the gill arches. Their new synapomorphies are from the intermuscular system, internal soft anatomy, pigmentation pattern of larvae, and morphology of the pelvic girdle. Most of these features are very difficult to assess in fossil specimens.

Sato and Nakabo (2002) accomplished a phylogenetic analysis of living Aulopiformes based on morphological and molecular data. They divided it into the suborders Synodontoidei, Chlorophthalmoidei, Alepisauroidei, and Giganturoidei. Moreover, the authors proposed a new family of Aulopiformes (i.e., Paraulopidae).

Fielitz (2004) and Gallo et al. (2005) proposed hypotheses of the phylogenetic relationships of some fossil Aulopiformes (Enchodontoidea and Dercetidae, respectively).

Nelson (2006) placed the extinct aulopiforms in three suborders: Ichthyotringoidei, comprising the families Ichthyotringidae, Dercetidae, and Prionolepidae; Halecoidei, with a single family, Halecidae; and Alepisauroidei, with two families, Cimolichthyidae and Enchodontidae. The extant aulopiforms were classified in Synodontoidei (with four families), Chlorophthalmoidei (with six families), and Giganturoidei (with two families). Additionally, four living families were placed in the suborder Alepisauroidei.

In fact, the assemblage of extinct aulopiforms defined by Nelson (2006) corresponds to Enchodontoidei sensu Nelson (1994). However, Nelson (2006) did not discuss his reasons for disregarding the name Enchodontoidei and put its members in Alepisauroidei, Ichthyotringoidei, and Halecoidei. Moreover, Enchodontoidei were considered in the cladistic analysis of Dercetidae (Gallo et al. 2005), as well as in a preliminary approach by Silva and Gallo (2007). As there has not been a recent comprehensive phylogenetic study of Enchodontoidei, we review their classification history and provide a new cladistic analysis.

SYSTEMATIC HISTORY OF ENCHODONTOIDEI

In this paper we use the general classification of Nelson (1994), except for the family Enchodontidae (sensu Fielitz 2004).

SUPERFAMILY ICHTHYOTRINGOIDEA

According to Goody (1969), Ichthyotringoidea comprises two closely related families, Ichthyotringidae and Apateopholidae. The author considered mainly primitive features of the body and caudal skeleton, as well as a derived feature related to the rostral region. He stated that, despite the similarities shared by the taxa, Apateopholidae should be the more advanced taxon.

Family Ichthyotringidae. The family Ichthyotringidae (Table I) was created by Jordan (1905) to contain asingle genus (i. e., Ichthyotringa ). Goody (1969) positioned the family in the suborder Ichthyotringoidei. Later, Nelson (1994) included the taxon in the suborder Enchodontoidei together with other fossil aulopiforms. The generic epithet Ichthyotringa was created by Cope (1878) to replace the genus Rhinellus of Agassiz (1844), which was pre-occupied. The genus Ichthyotringa includes the following species: Ichthyotringa furcata (Agassiz, 1844), I. tenuirostris Cope, 1878, I. damoni (Davis, 1887), I. ferox (Davis, 1887), I. delicata (Hay, 1903), and I. africana (Arambourg, 1954). Forey et al. (2003), in the general list of fossil fishes from Lebanon, placed Apateopholis in the family Ichthyotringidae, but remarked that at least one species of Apateopholis is often misinterpreted as a species of theclosely related Ichthyotringa. Only more recently a new ichthyotringoidei was reported to the El Doctor Formation in the Albian-Cenomanian of Mexico, I. mexicana Fielitz and Gonzá lez Rodríguez, 2008.

Goody (1969) ranked Apateodus as an addendum (incertae sedis ) to the Ichthyotringidae with a single species (A. striatus Woodward, 1901). Nelson (1994, 2006) placed Apateodus in Apateopholidae, but hedid not report the taxonomic status of Apateopholis. In Frickhinger (1995), the latter was considered anichthyotringid and the former was not mentioned. Foreyet al. (2003) positioned Apateopholis in the family Ichthyotringidae. Taverne (2004) maintained Apateodus and two other genera of Cretaceous alepisauroids (Yabrudichthys and Rharbichthys ) as family incertaesedis. However, the same author (Taverne 2006c) suggested to exclude Apateodus from Ichthyotringidae. Fielitz and Gonzá lez Rodríguez (2008) accomplisheda cladistic analysis of Ichthyotringoidea and placed Apateodus tentatively among the species of Ichthyotringa. More recently, Fielitz and Shimada (2009) described a new species of Apateodus (A. busseni ) suggesting that the genus needs revision, but ranking it in Ichthyotringidae.

Family Apateopholidae. The family Apateopholidae (Table I) was erected by Goody (1969) to include only the genus Apateopholis, with two species [ A. laniatus (Davis, 1887); A. lanceolatus Woodward, 1901] . The genus Apateopholis was erected by Woodward (1891) to substitute Rhinellus by Davis (1887), which was posteriorly allocated in Belonostomus by Woodward (1888). Ten years after the creation of the genus, Woodward (1901) put Apateopholis in synonymy with Prionolepis. Goody (1969) revalidated the generic epithet Apateopholis with the single species Apateopholis laniatus.

SUPERFAMILY CIMOLICHTHYOIDEA

This superfamily includes three families (i.e., Cimolichthyidae, Dercetidae, and Prionolepididae). According to Goody (1969), Cimolichthyidae and Dercetidae show a great similarity regarding the structures of the skull and body, especially in the rostral region. Moreover, general body squamation is lacking and two or three rows of isolated scutes are present on the flanks. Regarding the family Prionolepididae, the author pointed out some problems concerning its taxonomic placement. UnlikeWoodward (1901), who assigned the genus Prionolepis to the Enchodontidae, Goody (1969) considered it closely related to dercetids and cimolichthyids.

Family Cimolichthyidae. The family (Table II) was erected by Goody (1969) to include the single genus Cimolichthys, which was designated by Leidy (1857) with the species C. levesiensis. Cope (1872), studying specimens from Niobrara (USA), recognized five species of Cimolichthys: C. nepaholica, C. sulcatus, C. semianceps, C. contracta, and C. merrillii. Later, Hay (1903) recognized only the species C. nepaholica, as other species were based mainly on isolated teeth and fragments of the jaws.

Family Dercetidae. Traditionally the creation of this family is attributed to Pictet (1850). However, he did not use the name Dercetidae, defining only a Dercetis group with D. tenuis, D. triqueter, and D. linguifer. As far as we know, the name Dercetidae was used for the first time by Woodward (1901).

The following taxa are regarded as valid in themost recent revisions of the Dercetidae (e. g., Taverne1987, 1991, 2005a, b, 2006b, Chalifa 1989a, Gallo etal. 2005, Blanco et al. 2008) and in general fossil fish lists (e. g., Frickhinger 1995, Forey et al. 2003) : Apuliadercetis tyleri Taverne, 2006a; Benthesikyme armatus (von der Marck, 1863); B. rostralis (Davis, 1887); B. gracilis (Signeux, 1954); Brazilodercetis longirostris Figueiredo and Gallo, 2006; Caudadercetis bannikovi Taverne, 2006b; Cyranichthys ornatissimus (Casier, 1965); Dercetis elongatus (Agassiz, 1837); D. triqueter Pictet, 1850; Dercetoides venator Chalifa, 1989a; Hastichthys gracilis (Chalifa, 1989a); Leccedercetis longirostris Taverne, 2008; Nardodercetis vandewallei Taverne, 2005a; Ophidercetis italiensis Taverne, 2005b; Pelargorhynchus dercetiformis von der Marck, 1858; Rhynchodercetis hakelensis (Pictet and Humbert, 1866); R. yovanovitchi Arambourg, 1943; R. gortanii (d'Erasmo, 1946); R. regio Blanco and Alvarado-Ortega, 2006; R. serpentinus (Hay, 1903); Robertichthys riograndensis Blanco-Piñ on and Alvarado-Ortega, 2005; and Scandiadercetis limhamnensis (Davis, 1890) (Table II).

Chalifa (1989a) stated that Dercetidae are a " relatively primitive" group, considering the presence of few apomorphies. She carried out a phylogenetic analysis of this taxon and recognized it as a monophyletic group, comprising the following clades: (Dercetis, (Pelargorhynchus, (Dercetoides, Rhynchodercetis))).

Taverne (1987, 1991) considered Dercetidae to be a clade of Stomiiformes. In reviewing the material studied by Chalifa (1989a) and discussing the relationships of genera, Taverne (1991) assumed a monophyletic condition for this family and used 33 characters to define it. He placed Benthesikyme as the most primitive genus, possessing most of (or all) these 33 " generalized conditions" . Taverne (1991) also " distributed" 40 " apomorphies" to genera and groups of genera in the family.

Gallo et al. (2005) accomplished a cladistic analysis of the family Dercetidae, using an outgroup composed of Enchodontoidei taxa. The authors obtained only one tree supporting the monophyly of Dercetidae, based on two synapomorphies: absence of a longitudinal opercular crest and reduced neural spine. They also verified that some of the superfamilies proposed by Nelson (1994) are not monophyletic. Taking into account the description of four new genera of Dercetidae from Nardò, Italy (Taverne 2005a, b, 2006a, b), as well as the new diagnoses of Dercetis and Benthesikyme (Taverne 2005b), Taverne (2006b) proposed a phylogenetic review of the Dercetidae, in which he provided a list of 43 plesiomorphies to this family. In the phylogenetic hypothesis furnished by Blanco et al. (2008), the monophyly of Dercetidae was confirmed and it was supported by the same synapomorphies shown in Gallo et al. (2005).

Dercetis was the first described genus of Dercetidae. It was created by Agassiz (1834) to accommodate the species D. scutatus. The taxon was briefly described on the basis of a single and almost complete specimen. The holotype was lost or destroyed, without being figured (Siegfried 1954, Goody 1969, Taverne 2005b). Pictet (1850) erected Dercetis triqueter and D. linguifer. Later, von der Marck (1863) identified a new genus within Dercetidae to include two new species, Leptotrachelus armatus and L. sagittatus. Pictet and Humbert (1866) synonymized D. triqueter and D. linguifer with Leptotrachelus triqueter. Siegfried (1966) redescribed the material by von der Marck (1863) and transferred the Leptotrachelus spp. for the genus Dercetis (D. armatus and D. sagittatus). Goody (1969) redescribed D. triqueter and claimed that D. linguifer was clearly D. triqueter.

Pelargorhynchus dercetiformis was erected by von der Marck (1858) based on several but poorly preserved specimens from Germany. There is no record of this genus in any other locality.

The genus Benthesikyme was created by White and Moy-Thomas (1940), including new species of Leptotrachelus described by several authors at the end of the nineteenth century and during the twentieth century. Taverne (2005b) furnished a comprehensive review of the genera Dercetis, Leptotrachelus, and Benthesikyme, in which L. sagittatus is probably a synonym of D. elongatus, L. virgulatus of D. triqueter, and L. longipinnis of Benthesikyme gracilis. The taxonomic status of D. reussi, D. latiscutatus, D. maximus, and L. serpentinus was also discussed but not in a conclusive way. Yet, Taverne (2005a, b) erected three new monotypic genera within the family Dercetidae, Ophidercetis (O. italiensis), Nardodercetis (N. vandewallei), and Scandiadercetis (S. limhamnensis). The latter had been proposed originally by Davis (1890) as Dercetis limhamnensis.

The genus Rhynchodercetis was erected by Arambourg (1943), comprising a single species, R. yovanovitchi, which is very abundant in deposits from the Lower Cenomanian of Morocco. Later, other Rhynchodercetis spp. were described: R. hakelensis (Pictet and Humbert, 1866); R. gortanii (d' Erasmo, 1946); and R. regio Blanco and Alvarado-Ortega, 2006. Chalifa (1989a) described Dercetoides venator and Rhynchodercetis gracilis. The latter was renamed as Hastichthys gracilis by Taverne (1991). Blanco-Piñ on and Alvarado-Ortega (2005) briefly described Robertichthys riograndensis, which is the second record of Dercetidae in the Turonian of Mexico. Blanco et al. (2008) provided a redescription of this taxon, as well as a discussion on its relationships.

Figueiredo and Gallo (2006) described Brazilodercetis longirostris, which is the first record of the family in South America.

Family Prionolepididae. The genus Prionolepis was created by Egerton (in Dixon 1850) with only one species, P. angustus. Later, Pictet and Humbert (1866) included one more species in the genus, P. cataphractus. Goody (1969) reviewed P. cataphractus and proposed the family Prionolepididae (Table II).

SUPERFAMILY ENCHODONTOIDEA

Family Enchodontidae. Previously to Woodward (1901), the genera assigned to Enchodontoidea were allocated in different families (Agassiz 1835, Pictet 1850, Cope 1872, 1874). The first attempt to classify the enchodontoids in a separate group was proposed by Woodward (1901), in which the author erected the family Enchodontidae and put it in the Isospondyli. He divided the family into two main groups based on the presence or the absence of a single tooth in palatine. In the first group, the author included the genera Enchodus, Palaeolycus, Eurypholis, and Saurorhamphus; the second consisted of the genera Halec, Cimolichthys, Prionolepis, Leptecodon, and Pantopholis. However, Woodward (1901) stated that the living families more closely related to the Enchodontidae were Alepisauridae and Odontostomidae, which possess the border of the upper jaw formed exclusively by the premaxilla, the maxilla being untoothed and excluded from the mouth gape.

After Woodward (1901), there was a long debate about the relationships of enchodontids and living fish families. Jordan (1905), Gregory (1933), and Arambourg (1954) agreed with the hypothesis of Woodward (1901) regarding the relationships of enchodontids and alepisaurids, but they positioned Enchodontidae in the suborder Iniomi. On the other hand, Regan (1911) and Romer (1945) rejected the hypothesis of Woodward (1901) and allocated the family into the Stomiatoidei, belonging to the suborder Isospondyli. Berg (1940) also rejected the enchodontids plus alepisaurids hypothesis and put enchodontids in the suborder Enchodontoidei into the Clupeiformes, as synonym of Isospondyli. The generic composition of Enchodontidae remained stable for some time, except for Halec, which was moved to the family Halecidae by Goody (1969), and for the inclusion of Rharbichthys by Arambourg (1954).

Goody (1969) accomplished a comprehensive review of the Enchodontidae, including the genera Enchodus and Palaeolycus. Also, he created the family Eurypholidae into Enchodontoidei to comprise the genera Eurypholis and Saurorhamphus.

Sorbini (1976) proposed a relationship between Rharbichthys and Cimolichthys. Taverne (1985) studied Rharbichthys and stated that it probably possessed close affinity with the halecids regarding the general proportions and the head shape.

Although the monophyly of Enchodontidae seems to be widely accepted, the previous diagnoses (e. g., Goody 1969, Rosen 1973, Chalifa 1989b) were not deduced from cladistic analyses.

Recently, Fielitz (2004) tested the monophyly of the family Enchodontidae, including living and extinct aulopiforms. The clade is supported by three synapomorphies: single dermopalatine tooth, dermopalatine bone with same length or shorter than the tooth, and interopercle absent. Alepisauridae appear in the analysis as the sister group of the clade formed by the extinct Aulopiformes. Enchodontidae were divided into four subfamilies: Rharbichthinae (with Rharbichthys), Palaeolycinae (with Palaeolycus), Eurypholinae (with Eurypholis and Saurorhamphus) and Enchodontinae (with Enchodus). The genus Parenchodus was put in synonymy with Enchodus.

The subfamily Rharbichthinae (Table III) is monotypic, being represented only by Rharbichthys ferox Arambourg, 1954. This species was considered by several authors as belonging to the Enchodontidae (e. g., Bertin and Arambourg 1958, Leonardi 1966, Goody, 1969), whereas Sorbini (1976) classified it in the family Cimolichthyidae. Yet, Taverne (1985) claimed that R. ferox is an alepisauroid.

The subfamily Palaeolycinae (Table III) is monotypic, comprising only Palaeolycus dreginensis described by von der Marck (1863). Later, it was reviewed by Siegfried (1954), who pointed out morphological similarities and putative relationships with the extant genus Odontostomus. This genus is in synonymy with Evermanella, which is in the family Evermannellidae of the suborder Alepisauroidei.

The subfamily Eurypholinae (Table III) was originally proposed as a family (Eurypholidae) by Goody (1969) to encompass the genus Eurypholis by Pictet (1850). This genus comprises only the type-species (E. boissieri Pictet, 1850) and another one initially proposed as Enchodus pulchellus by Woodward (1901), but laterredefined by Goody (1969) as Eurypholis pulchellus.

Nelson (1994) placed the family Eurypholidae inthe superfamily Enchodontoidea in the suborder Enchodontoidei.

Fielitz (2004) suggested the arrangement of the genera Eurypholis and Saurorhamphus in the subfamily Eurypholinae.

Saurorhamphus freyeri was originally described by Heckel (1850), and transferred to the genus Eurypholis by Woodward (1901). However, d'Erasmo (1912) claimed that Saurorhamphus was actually a distinct genus, closely related to Eurypholis. Another species, S. judeaensis, was described by Chalifa (1985).

The subfamily Enchodontinae (Table III) includes only the genus Enchodus Agassiz, 1835, containing, however, about 24 valid species, most of them erected on the basis of fragmentary material (isolated teeth or pieces of jaws), as follows: Enchodus brevis Chalifa, 1989b; E. bursauxi (Arambourg, 1952); E. dentex (Heckel, 1856); E. dirus (Leidy, 1857); E. faujasi Goody, 1968; E. ferox Leidy, 1855; E. gladiolus (Cope, 1872); E. gracilis (von der Marck, 1858); E. lewesiensis (Mantell, 1822); E. libycus (Quaas, 1902); E. longidens (Pictet, 1850); E. longipectoralis (Schaeffer, 1947); E. longipterygius (Raab and Chalifa, 1987); E. lycodon Kner, 1867; E. macropterus (von der Marck, 1863); E. major Davis, 1887; E. marchesettii (Kramberger, 1895); E. mecoanalis Forey, Yi, Patterson and Davies, 2003; E. oliverai Maury, 1930; E. petrosus Cope, 1874; E. shumardi Leidy, 1856; E. subaequilateralis Cope, 1886 (= E. elegans ); E. venator Arambourg, 1954; E. zinensis Chalifa, 1996 (e. g., Goody 1976, Chalifa1996, Forey et al. 2003, Fielitz 2004).

Parenchodus longipterygius was described by Raab and Chalifa (1987) as belonging to the family Enchodontidae. The authors suggested a relationship with the genus Enchodus, due to similarities in some structures of the head, the absence of scales, and the fusion of the elements of the caudal fin. Fielitz (2004) put the genus in synonymy with Enchodus.

SUPERFAMILY HALECOIDEA

Family Halecidae. The family Halecidae (Table IV) was originally proposed by Agassiz (1834) including forms similar to the clupeoids and salmonoids. The grouping and its name were used only by Pictet (1850) and Davis (1887), being disused later. The family was re-erected by Goody (1969) into the suborder Halecoidei containing three genera: Halec, Phylactocephalus, and Hemisaurida. Nelson (1994) put Halecidae in the superfamily Halecoidea in the suborder Enchodontoidei. Later, Nelson (2006) opted to use the suborder Halecoidei by Goody (1969).

Regarding the genus Halec, there are three valid species: H. sternbergi Agassiz, 1844 (type-species), H. eupterygius (Dixon, 1850), and H. haueri (Bassani, 1879).

The genus Phylactocephalus was erected by Davis (1887) and put in synonymy with Halec by Woodward (1901). Goody (1969) verified marked differences between the genera and separated them. He re-erected Phylactocephalus with a single species, P. microlepis Davis, 1887.

Kner (1867) created the genus Hemisaurida containing a single species, H. neocomiensis. Woodward (1901) and Romer (1966) suggested that this genus could belong to the family Myctophidae. Goody (1969) rejected this hypothesis based mainly on two features present in halecoids: maxilla partially excluded from the mouth gape and premaxilla without ascending and articular processes. Yet, Goody (1969) created the species H. hakelensis.

TAXA INCERTAE SEDIS

Family Nardorexidae. The monotypic family Nardorexidae (Table V) was proposed by Taverne (2004) with the species Nardorex zorzini. He placed the family in the suborder Alepisauroidei based on putative relationships with Enchodontoidei.

Family Serrilepidae. The taxon was proposed by Chalifa (1989c) with the single species Serrilepislongidens. Forey et al. (2003) added two new speciesto the genus, S. prymnostrigos and S. minor. According to these authors, among the aulopiforms, Serrilepis is more closely related to Halec, Hemisaurida, and Phylactocephalus and, therefore, it should be classified into the Halecidae. This relationship is based on two synapomorphies: fusion of dorsal hypohyal and anterior ceratohyal; fusion of first and second hypurals andthird and fourth hypurals. However, this latter synapomorphy is present in Atolvorator longipectoralis (Gallo and Coelho 2008), and we opted to use Chalifa'sclassification with Serrilepis in the family Serrilepidae (Table V).

Additionally, two other taxa, Yabrudichthys striatus Chalifa, 1989c, and Atolvorator longipectoralis Gallo and Coelho, 2008, are considered Enchodontoidei incertae sedis and Cimolichthyoidei incertae sedis, respectively (Table V).

MATERIALS AND METHODS

MATERIAL

The specimens of Enchodontoidei herein studied belong to several paleontological collections (see ^{Appendix I}). Extant aulopiform, stomiiform, and myctophiform fishes were used as comparative specimens represented by dry skeletons, alcohol-preserved, and cleared and stained specimens. They belong to the AO.UERJ, O.UERJ, and MZUSP (see ^{Appendix I}). Moreover, for the taxa of difficult access, as for instance, those deposited in Hebrew University of Jerusalem and Museo Civico di Storia Naturale di Verona, we selected information from available literature (e.g., Chalifa 1985, 1989 a, b, c, 1996, Raab and Chalifa 1987, Taverne 2005a, b, 2006a, b).

INSTITUTIONAL ABBREVIATIONS

AO. UERJ, Ichthyological Collection, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Riode Janeiro, Brazil; DGM, Divisão de Geologia e Mineralogia, Departamento Nacional da Produção Mineral, Rio de Janeiro, Brazil; FPH, Fundação Paleontológica Phoenix, Aracaju, Brazil; MN, Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; MNHN, Muséum National d'Histoire Naturelle, Paris, France; MZUSP, Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil; NHM, The Natural History Museum, London, England; O. UERJ, Ichthyological Collection, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil; Pz. UERJ, Paleozoological Collection, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Riode Janeiro, Brazil.

CLADISTIC METHODODOLOGY

A data matrix was built with 87 characters, unordered and unweighted, 31 terminal taxa for ingroup, and three taxa for outgroup. The parsimony analysis was carried out using the computer program PAUP* version 4.0b10 (Swofford 2001), with the heuristic algorithm HSearch. We tried to apply the exact branch-and-bound algorithm, but due to the length and complexity of the data matrix, we failed to obtain a result even running the analysis for a few days.

The character states that could not be verified mainly due to poor preservation were coded in the matrix as " ?" (missing data). All characters previously proposed in the literature have been reviewed: the character 54 was adapted from Goody (1969); the characters 1, 2, 21, 23, 26, 41, 63, 70, 71, 83, and 84 were from Chalifa (1989a); the characters 3, 10, 11, 16, 19, 24, 25, 29, 30, 33, 38, 42, 64, 65, 77, and 86 were from Taverne (1991); the characters 75, 76, and 81 were from Baldwin and Johnson (1996); the characters 5, 6, 7, 17, 31, 48, 49, 50, 52, 57, 62, 66, and 79 were from Fielitz (2004); the characters 4, 12, 14, 15, 18, 34, 39, 40, 43, 44, 45, 46, 47, 51, 56, 59, 60, 61, 67, 78, 85, and 87 were from Gallo et al. (2005). The new characters are 8, 9, 13, 20, 22, 27, 28, 32, 35, 36, 37, 53, 55, 58, 68, 69, 72, 73, 74, 80, and 82. The terminal taxa for the ingroup were: Ichthyotringidae (Ichthyotringa ), Apateopholidae (Apateodus, Apateopholis ), Cimolichthyidae (Cimolichthys ), Dercetidae (Apuliadercetis, Bentheskyme, Brazilodercetis, Caudadercetis, Cyranichthys, Dercetis, Dercetoides, Hastichthys, Nardodercetis, Ophidercetis, Pelargorhynchus, Robertichthys, Rhynchodercetis ), Prionolepididae (Prionolepis ), Enchodontidae (Rharbichthys, Palaeolycus, Eurypholis, Saurorhamphus, Enchodus, Parenchodus ), Halecidae (Halec, Hemisaurida, Phylactocephalus ), Serrilepidae (Serrilepis ), as well as the incertae sedis genera Nardorex, Yabrudichthys, and Atolvorator. Outgroup is based on Protostomias (Stomiiformes), Trachinocephalus (Aulopiformes), and Sardinioides (Myctophiformes).

^{Appendix II} includes the coded data matrix, which was built based on the list of characters presented in the Results. Only character states that resulted in apomorphies were illustrated. Although the strict consensus is the real consensus that shows all possible topologies, we opted to present the majority rule consensus (MRC) because, in general, it possesses a better resolution. The MRC is a form of consensus that preserves all clades present in the majority (i.e., in more than 50%) of the obtained set of equally parsimonious cladograms (Margush and McMorris 1981). The 50% rule ensures that all included clades are compatible (Sharkey and Leathers 2001). In spite of some criticism (e.g., Sharkey and Leathers 2001), several authors are using MRC as a method of weighting clades to solve ambiguous strict consensus trees (e.g., Swofford 1991, Candall and Fritzpatrick 1996, Titus and Larson 1996, Lutzoni 1997).

RESULTS

Eighty-seven characters were analyzed in this study (see ^{Appendix III}). The cladistic analysis produced 93 equally parsimonious trees, with a tree length of 437 steps, consistency index (CI) of 0.24, and retention index (RI) of 0.49. The majority rule consensus tree is shown in Figure 1. The majority rule consensus tree is represented by the following topology: (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 )))))).

DISCUSSION

Before discussing the cladistic analysis per se, we will furnish a brief comment on certain characters.

According to Chalifa (1989a) and Taverne (1991), the presence of a low head is a synapomorphy of Dercetidae. In this study, it is shared with two other genera outside the group, showing a homoplastic distribution.

In fossil aulopiform fishes, particularly in most of Cimolichthyoidei and Enchodontoidei (sensu Goody 1969), the snout length is equivalent to the diameter of the orbit (e.g., Cimolichthys and Eurypholis, respectively). All dercetids possess an elongate snout and the extreme condition is verified in Hastichthys, in which the snout length reaches more than 12 times the diameter of the orbit.

Fielitz (2004) considered the presence of vomerine teeth as a synapomorphy of the group formed by Cimolichthys and members of the family Enchodontidae. In the present study, this state of character has a homoplastic distribution, due to it is present in Nardorex and Prionolepis.

Gallo et al. (2005) interpreted the presence of amesethmoid with a bifid anterior extremity as an autapomorphy of Dercetis, but herein this condition was also verified in Nardorex and Sardinioides. Yet, Galloet al. (2005) pointed out a mesethmoid with a bifid posterior extremity in Dercetis. However, Taverne (2005b) described this bone with an acute posterior extremity, and we confirmed this feature in Dercetis elongatus (NHM P. 49793) and D. triqueter (MNHN SHA 523). Yet, Taverne (2006c) described Ichthyotringa africana as possessing a mesethmoid with acute posterior extremity, mainly based on specimens MNHN DTS 225-228. However, observing the specimen MNHN DTS 228, weverify a mesethmoid with bifid posterior extremity.

In Gallo et al. (2005), the autosphenotic spine posteriorly curved is a synapomorphy of the clade (Rhynchodercetis, Hastichthys ). However, in this study, the character shows a slightly wide distribution, being also present in Atolvorator and Trachinocephalus. Taverne (2006b) stated that the autosphenotic of Caudadercetis is hidden by the frontals. For this reason, we opted tocode it as missing data.

Accordig to Taverne (1991), Pelargorhynchus is characterized by four derived features, among them the loss of the supraoccipital crest. Gallo et al. (2005) considered the absence of the supraoccipital crest as an autapomorphy of Pelargorhynchus. In the present study, the character was differently interpreted, because its absence and/or loss were verified in other two taxa (Benthesikyme and Rharbichthys ).

Gallo et al. (2005) pointed out the presence of apterotic not projecting beyond the occiput in Dercetis based on available descriptions. However, we point edout another condition (projecting beyond the occiput), following the redescription of the genus furnished by Taverne (2005b), as well as by direct observation of specimens of Dercetis triqueter and D. elongatus.

The presence of an unroofed post-temporal fossa in Palaeolycus followed Goody (1969), in contrast with Fielitz (2004) who pointed out a roofed condition. We verify a roofed condition in Eurypholis and Saurorhamphus in disagreement with Fielitz (2004). Gallo et al. (2005) pointed out a partially roofed posttemporal fossa in the genus Dercetis. However, following the redescription of the genus furnished by Taverne (2005b), as well as the direct observation of specimens of Dercetis triqueter and D. elongatus, we indicate herein a roofed post-temporal fossa to Dercetis.

Rosen (1973) pointed out the absence of orbitosphenoid in enchodontids. Additionally, Taverne (1991) stated that its absence would be a primitive conditionof dercetids. Gallo et al. (2005) proposed the presenceof orbitosphenoid as an autapomorphy of Ichthyotringa. Generally, the common absence of an orbitosphenoid inthe specimens observed directly or indirectly is probablydue to its fragility, which impedes a good preservation.

Gallo et al. (2005) suggested the presence of the basisphenoid as an autapomorphy of Ichthyotringa, but other taxa (i. e., Apuliadercetis tyleri, Atolvorator longipectoralis and Robertichthys riograndensis ) bearing this bone were described after this study.

Taverne (1985) pointed out three conditions for the presence of teeth on the ectopterygoid: bone toothless or bearing some small conic teeth in a small portion of it as in Rharbichthys ferox, from the Cenomanian of Morocco; and well-developed teeth on the ectopterygoid similar to those found in the dentary of Rharbichthys cf. ferox from the Cenomanian-Turonian of Italy (see also Sorbini 1976). For this reason, the character was regarded as polymorphic.

In the cladistic analysis herein performed (Fig. 1), we demonstrate that the suborder Enchodontoidei is not a monophyletic group, as two genera of the outgroup belonging to another suborder and even to another order

(Trachinocephalus and Protostomias, respectively) went to the ingroup. Although Trachinocephalus belongs to the same order of the taxa herein studied (Aulopiformes), it is allocated into the suborder Synodontoidei, together with other extant members of Aulopiformes (Baldwin and Johnson 1996, Sato and Nakabo 2002). Arambourg (1954) and Taverne (1991) included Protostomias in the order Stomiifomes, based on generalized anatomical features, such as general shape of the body and a massive and tooth-bearing dentary, as well as the position of the median fins. Taverne (1992), in his comprehensive review of Protostomias, retained its placement in Stomiiformes.

Yet, the paraphyly of Enchodontoidei not allowed their taxonomic classification in the cladistic context.

The family Apateopholidae is not a monophyletic group, as Apateopholis is the sister-taxon of the clade including the family Enchodontidae (new usage) and the genera Cimolichthys, Prionolepis, Halec, Phylactocephalus, and Serrilepis. Apateodus, often placed with the Apateopholidae, is the sister-group of Ichthyotringa.

The monophyly of Dercetidae proposed by Gallo et al. (2005) and Blanco et al. (2008) was corroborated and supported by a single synapomorphy (very reduced neural spine; character 71) (Fig. 2), but the inclusion of new taxa changed the interrelationships of the family. The genus Apuliadercetis is the sister-group of Brazilodercetis and they are the basal clade of Dercetidae. Cyranichthys is the sister-taxon of Robertichthys, and Dercetis is the sister-taxon of Ophidercetis. These two clades plus Benthesikyme form a new clade, which is anintermediary group between the basal and crown groups. However, the relationships with in this intermediary group are uncertain. Caudadercetis appears as the most basal taxon in the major clade of Dercetidae (Caudadercetis, (Pelargorhynchus, (Nardodercetis, (Rhynchodercetis, (Hastichthys, Dercetoides )))). This clade is sustained by the unique presence of a convoluted suture marking the contact between second and third hypurals (character 83; Fig. 3B), although this characterin Pelargorhynchus was coded as missing data, as it scaudal skeleton is unknown. A pipe-shaped preopercle is an autapomorphy of Brazilodercetis (character 56; Fig. 4B).

Gallo et al. (2005) and Blanco et al. (2008) pointed out two synapomorphies of Dercetidae: the absence of a longitudinal opercular crest and a reduced neural spine. However, the absence of an opercular crest was also verified in other taxa. Yet, according to Gallo et al. (2005), Dercetoides is the sister-taxon of the clade formed by Rhynchodercetis and Hastichthys, but in Blanco et al. (2008) Dercetoides is the sister-taxon of Hastichthys, and both are sister-taxa of Rhynchodercetis. On the other hand, Taverne (2006b) stated that Dercetoides is the sister-taxon of Rhynchodercetis, and Hastichthys isthe sister-taxon of the clade (Caudadercetis, (Pelargorhynchus, (Rhynchodercetis, Dercetoides ))). Herein, Hastichthys appears as the sister-taxon of Dercetoides. Taverne (2006b) suggested Dercetis as the most basal dercetids, and Ophidercetis as the sister-taxon of Cyranichthys. However, we obtained different results: Dercetis forms a clade with Ophidercetis, and Cyranichthys is related to Robertichthys.

The family Enchodontidae is confirmed as monophyletic, as has already been proposed by Fielitz (2004), but herein it possesses a new composition. Parenchodus forms a clade with Palaeolycus, and Enchodus is the sister-group of this clade; the clade formed by (Eurypholis, Saurorhamphus ) is the basal sister-group. Rharbichthys was excluded from the Enchodontidae, being recognized in this analysis as the sister-group of the clade formed by the Dercetidae plus the genusTrachinocephalus. Fielitz (2004) proposed three synapomorphies of Enchodontidae: the presence of a single dermopalatine tooth, dermopalatine bone with same length or shorter than the tooth, and the absence of an interopercle. In this analysis, these features were not corroborated as synapomorphies. For instance, a single tooth on the dermopalatine is also present in Ophidercetis, a genus of Dercetidae (Taverne 2005b). The remaining synapomorphy of this family is the presence of middorsal scutes (character 87; Fig. 5). In addition, the clade (Eurypholis, Saurorhamphus ) is supported by two synapomorphies: quadrate-mandibular articulation hidden (character 52; Fig. 6) and the presence of a spine on posterior border of the opercle (character 61; Fig. 7).

Halecidae possess a new composition: Halec is the sister-group of Phylactocephalus; and Hemisaurida was excluded from the family, being considered Aulopiformes incertae sedis, like Nardorex.

The presence of a well-developed supraoccipital divided into two distinct regions is an autapomorphy of Nardorex (character 20; Fig. 8B). These two regions are separated by a slight transverse ridge: the anterior region is reduced and contacts the parietals, whereas the posterior one is large, contacts the epioccipitals and bears a high median crest (Fig. 8B).

ACKNOWLEDGMENTS

We are most grateful to R.C.T. Cassab (Departamento Nacional de Produção Mineral, Rio de Janeiro), D.D.R. Henriques (Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro), O.T. Oyakawa (Museu de Zoologia/Universidade de São Paulo, São Paulo), M. Richter (The Natural History Museum, London), H.R.S. Santos (Universidade do Estado do Rio de Janeiro, Rio de Janeiro), W. Souza-Lima (Fundação Paleontológica Phoenix, Aracaju), and M. Véran (Muséum National d'Histoire Naturelle, Paris), for access to specimens in their care. We thank S.A. Azevedo, M.J. Cavalcanti, F.J. de Figueiredo, and J. Alvarado Ortega for critical comments to the M.S. thesis of the H.M.A.S advised by V.G., on which this paper is based. We are particularly grateful to T. Moulton for his help with the English revision. This research was part of a Project supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 476708/2004-4) and the Fundação Carlos Chagas de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ E-26/171.215/2004). HMAS holds a fellowship from the FAPERJ and VG has research fellowship grants from the CNPq (Brazilian Federal Government), from the FAPERJ (Jovem Cientista do Nosso Estado), and from the PROCIÊNCIA (Rio de Janeiro State Government).

Manuscript received on September 24, 2010; accepted for publication on December 22, 2010

APPENDIX I

List of the analyzed material

RECENT MATERIAL

Synodontidae - Trachinocephalus myops: AO. UERJ17; AO. UERJ 123; AO. UERJ 127; AO. UERJ 128; O. UERJ 471; O. UERJ 1334; O. UERJ 2073.

Evermannellidae - Coccorella atlantica: MZUSP 78-193.

Paralepididae - Lestidium atlanticum: MZUSP 60327; Lestrolepis intermedia: MZUSP 80717.

FOSSIL MATERIAL

Enchodontidae - Enchodus sp. 1: Pz. UERJ 489; Pz. UERJ 490; Pz. UERJ 491; Enchodus sp. 2: Pz. UERJ 485; Pz. UERJ 492; Enchodus libycus: FPH-01-82; FPH-02-12; FPH-01 679; FPH-01 742; FPH-01 677; FPH-01 810; FPH-01 808; FPH-01 809; DGM 642-P; Enchodus oliveirai: DGM 643-P; Enchodus subaequilatelaris: DGM 644-P; MN 4329-V; Enchodus longipectoralis: DGM 501-P; Enchodus venator: MNHN DTS 156, MNHN DTS 157 d g; MNHN DTS 158 d g; MNHN DTS 160; MNHN DTS 148; MNHN DTS 138 d g; MNHN DTS 149; MNHN DTS 141 d g; MNHNDTS 159 d g; MNHN DTS 155; MNHN DTS 190 d g; MNHN DTS 143; Eurypholis boisseiri: Pz. UERJ 493; NHM P. 48; NHM P. 63323; Eurypholis longidens: NHM P. 48084; NHM P. 409507; Eurypholis pulchellus: NHM P. 1703; Eurypholis sp. : NHM P. 47317; Rharbichthys cf. ferox: Pz. UERJ 448 a b; Pz. UERJ 451; Pz. UERJ 452; Rharbichthys ferox: MNHN DTS 118 d g; MNHN DTS 162 d g; MNHN DTS 130 d g; MNHN DTS 81 d g; MNHN DTS 86 d g; MNHN DTS 164 d g; MNHNDTS97 d g; MNHNDTS119 g d; MNHN DTS 163 g d; MNHN DTS 165; MNHN DTS166; MNHN DTS 167; MNHN DTS 71; MNHN DTS161; MNHN DTS 72 d g; MNHN DTS 126 d g.

Dercetidae - Benthesikyme armatus: NHM P. 2109; Benthesikyme gracilis: NHM P. 48085; NHM P. 49539; NHM P. 48087; NHM P. 1902; NHM P. 47359; NHM P. 49538; NHM P. 9170; NHM P. 47360; NHM P. 4738; NHM P. 46553; NHM P. 48088; NHM P. 46533; NHM P. 48086; Benthesikyme rostralis: MNHN SHA 499; MNHN SHA 501; MNHN SHA 505; MNHN SHA 513; MNHN SHA 578; MNHN SHA 2840; MNHN SHA 2849; Benthesikyme sp.: NHM P. 4019; Brazilodercetis longirostris Pz.UERJ 447; Pz.UERJ 471; Pz.UERJ 472; Pz.UERJ 473; Pz.UERJ 474; Pz.UERJ 475; Pz.UERJ 476; Pz.UERJ 477; Pz.UERJ 478; Pz.UERJ 495.; Dercetis elongatus 4018; 49536; NHM P. 15513; NHM P. 43098; NHM P. 4134; NHM P. 4132-33; NHM P. 41198; NHM P. 49793; NHM P. 12895; NHM P. 43512; NHM P. 31075-82.; Dercetis sp. NHM P. 49740; NHM P. 48155; Dercetis triqueter NHM P. 4852; NHM P. 49541; NHM P. 49535; NHM P. 4007; NHM P. 46524; NHM P. 4963; NHM P. 49537; NHM P. 47362; NHM P. 47361, MNHN SHA 520; MNHN SHA 523; MNHN SHA 572; MNHN SHA 2192 d g; MNHN SHA 2317; MNHNSHA2444; MNHNSHA3107; Rhynchodercetis cf. gracilis NHM P. 63235 a/b; NHM P. 63332; NHM P. 63246 a/b; NHM P. 62677 a/b; Rhynchodercetis cf. yovanovitchi NHM P. 62678; NHM P. 63236 a/b; NHM P. 63599; NHM P. 63261; NHM P. 62690; NHM P. 63262; Rhynchodercetis gortanii NHM P. 10913; Rhynchodercetis hakelensis NHM P. 6001; NHM P. 4739; NHM P. 4683; NHM P. 4866; Rhynchodercetis serpentinus NHM P. 51939; Rhynchodercetis yovanovitchi MNHN DTS 8; MNHN DTS 9 d g; MNHN DTS 10 d g; MNHN DTS 14; MNHN DTS 15; MNHN DTS 20; MNHN DTS 21; MNHN DTS 22; MNHN DTS 23; MNHN DTS 123; MNHN DTS 47; MNHN DTS 175; MNHN DTS 174; MNHN DTS 186; MNHN DTS 40; MNHN DTS 302; MNHN DTS 303; MNHN DTS 304; MNHN DTS 306; MNHN DTS 307; MNHN DTS 308; MNHNDTS 309; MNHNDTS 261 d g; MNHNDTS 96 d g; MNHN DTS 43 d g; MNHN DTS 310; MNHN DTS 189; MNHN DTS 176 d g; MNHN DTS177 d g; MNHN DTS 2; MNHN DTS 7; MNHN DTS 41; MNHN DTS 262; MNHN DTS 6 d g; MNHN DTS 263 a b.

Apateopholidae - Apateodus striatus NHM P. 61919; NHM P. 49799; NHM P. 4090-1; NHM P. 49821; NHM P. 12899; NHM P. 12898; NHM P. 33309; NHM P. 49067; NHM P. 479224; NHM P. 10058; NHM P. 49070; Apateopholis laniatus NHM P. 4745; NHM P. 4026; NHM P. 4870; NHM P. 63263; NHM P. 4869.

Cimolichthyidae - Cimolichthys levesiensis NHM P. 4039; NHM P. 4026; NHM P. 1810a; NHM P. 1811; NHM P. 38113; NHM P. 5491.

Prionolepididae - Prionolepis cataphhractus NHM P. 4864; NHM P. 9966; NHM P. 9967; NHM P. 9968; NHM P. 9970; NHM P. 47516; NHM P. 4871; NHM P. 47332; NHM P. 4006; Prionolepis laniatus NHM P. 39234.

Ichthyotringidae - Ichthyotringa sp. NHM P. 6015; NHM P. 14204; NHM P. 9996; NHM P. 48089 a/b; NHM P. 1882; NHM P. 49544; NHM P. 48092; NHM P. 48155; Ichthyotringa africana MNHN DTS 225; MNHN DTS 226; MNHN DTS 227 g d; MNHN DTS228 d g.; Ichthyotringa damoni NHM P. 47367; NHM P. 4849; Ichthyotringa furcata NHM P. 49523; NHM P. 49525; NHM P. 48090; NHM P. 49545; NHM P. 48091; NHM P. 47363; NHM P. 48144; NHM P. 47364.

Halecidae - Halec sternbergi NHM P. 9004; NHM P. 9004; NHM P. 5732; Halec eupterygius NHM P. 11102; NHM P. 43388; NHM P. 32336; NHM P. 4289; NHM P. 43392; NHM P. 10920; NHM P. 32237; Halec rugosus NHM P. 13899; Phylactocephalus microlepis NHM P. 4757; NHM P. 105; NHM P. 47318; NHM P. 9151; Hemisaurida hakelensis NHM P. 48779; NHM P. 48780.

Atolvorator longipectoralis Pz. UERJ 486; Pz. UERJ 487; Pz. UERJ 496 a e b; Pz. UERJ 508 a e b; Pz. UERJ 509; Pz. UERJ 510.

Sardinioides minimus NHM P. 52513; NHM P. 52511; NHM P. 52512; NHM P. 52504 a.

Protostomias maroccanus MNHN DTS 18 d g; MNHNDTS 122 d g; MNHN DTS 172 d g; MNHN DTS 178; MNHN DTS 179 d g; MNHN DTS 184; MNHN DTS77; MNHN DTS 381.

Click to enlarge

Appendix II

APPENDIX III

List of characters used in the phylogenetic analysis of † Enchodontoidei.

1. Body length: slightly elongate [lesser than or equal to 1:10] (0); elongate [from 1:11 to 1:15] (1); very elongate [higher than 1:15] (2).

2. Head height: deep (0); low (1).

3. Snout length: short (0); long (1).

4. Dermal pattern on skull roof: smooth (0); only with tubercles (1); only with ridges (2); tubercles + ridges (3).

5. Vomerine teeth: absent (0); present (1).

6. Number of teeth on dermopalatine: two or more (0); none (1); single (2).

7. Dermopalatine length: twice or more times longer than its tooth (0); equal-sized or shorter than its tooth (1).

8. Antorbital: present (0); absent (1).

9. Nasal: present (0); absent (1).

10. Anterior extremity of mesethmoid: acute (0); bifid (1).

11. Posterior extremity of mesethmoid: bifid (0); acute (1); straight (2).

12. Autosphenotic spine: straight (0); posteriorly curved (1).

13. Suture between frontals: slightly sinuous (0); markedly sinuous (1).

14. Posterior border of frontal: behind the autosphenotic spine (0); at the level of the autosphenotic spine (1).

15. Shape of the post-orbital border: concave (0); convex (1).

16. Parietal length: long [length equal or larger than its height (0)]; short [length smaller than its height (1)].

17. Supraorbital sensory canal in the skull roof: covered (0); exposed (1).

18. Extension of the supraoccipital: not separating parietals (0); separating parietals (1).

19. Supraoccipital crest: present (0); absent (1).

20. Supraoccipital with two well-delimited regions: absent (0); present (1).

21. Extension of pterotic: not projecting backwards beyond the level of occiput; (0) projecting beyond the occiput (1).

22. Dilatator fossa: unroofed (0); roofed (1).

23. Exposition of the post-temporal fossa: roofed (0); unroofed (1).

24. Orbitosphenoid: present (0); absent (1).

25. Basisphenoid: present (0); absent (1).

26. Supraorbital: present (0); absent (1).

27. Lachrymal shape: subtriangular (0); suboval (1); trapezoidal (2); rod-shaped (3).

28. Position of the mandibular suspensorium: inclined (0); vertical (1).

29. Ectopterygoid: toothless (0); toothed (1).

30. Endopterygoid: toothless (0); toothed (1).

31. Placement of articular facet for the hyomandibula: posteroventral (0); ventral (1).

32. Number of articular facet for the hyomandibula: a continuous facet (0); two facets (1).

33. Premaxilla: toothed (0); toothless (1).

34. Posterior extension of the premaxilla: reaching the orbit (0); not reaching the orbit (1).

35. Dermal pattern on premaxilla: smooth (0), ornamented (1).

36. Fenestra in the premaxilla: absent (0); present (1).

37. Ascending process of the premaxilla: absent (0); present (1).

38. Maxilla: toothless (0); toothed (1).

39. Placement of the maxilla: over the premaxilla (0); behind the premaxilla (1).

40. Teeth on upper jaw: only straight (0); absent (1); curved + straight (2); only curved (3).

41. Supramaxilla: absent (0); present (1).

42. Mandible length: equal to the snout (0); shorter than the snout (1).

43. Teeth on mandible: only straight (0); only curved (1); curved + straight (2).

44. Teeth size on upper jaw: absent or with same height (0); with different height (1).

45. Teeth size on mandible: different height (0); equal height (1).

46. Rows of teeth on upper jaw: single (0); two or more (1).

47. Rows of teeth on mandible: single (0); two or more (1). 48. Anteroventral prongs on dentary: absent (0); present (1).

49. Mandibular sensory canal: enclosed by bone (0); partially open (1); open (2).

50. Mandibular dermal pattern: smooth (0), ornamented (1).

51. Flange on anguloarticular: present (0); absent (1).

52. Quadrate-mandibular articulation: exposed (0); hidden (1).

53. Articular facet for the quadrate: shallow (0); deep (1).

54. Retroarticular process: present (0), absent (1).

55. Ornamentation in the infraorbital bones: smooth (0), ornamented (1).

56. Preopercle shape: L-shaped (0); triangular (1); crescent-shaped (2); rod-shaped (3); pipe-shaped (4).

57. Preopercular dermal pattern: smooth (0); ornamented (1).

58. Posteroventral spine in the preopercle: absent or reduced (0), well-developed (1).

59. Dimension of the opercle: deeper than long (0); longer than deep (1).

60. Opercle crest: absent (0); (1) present.

61. Spine on posterior border of the opercle: absent (0); present (1).

62. Opercular and subopercular dermal pattern: smooth (0); ornamented (1).

63. Interopercle: present (0); absent (1).

64. Mesocoracoid: absent (0); present (1).

65. Scapula and coracoid: individualized (0); co-ossified (1).

66. Supraneurals: present (0); absent (1).

67. Total number of vertebrae: more than 50 (0); equal or minus than 50 (1).

68. Number of caudal vertebrae: more than 20 (0); equal or minus than 20 (1).

69. Ribs: extending to the pelvic fin origin (0); surpassing the pelvic fin origin (1).

70. Transverse processes: one pair (0); two pairs (1).

71. Neural spines: well-developed [their length surpassing the length of the vertebral centrum] (0); very reduced [their length equal or minus than half of the length of the vertebral centrum] (1).

73. Distribution of epipleurals: extending to more than a half of the body (0); up to half of the body (1).

74. Distribution of epineurals: extending to more than a half of the body (0); up to half of the body (1).

75. Position of the pectoral fin: high [last fin-ray placed at the level of the ventral border of the opercle or a little above (0)]; low [last fin-ray placed below the level of the ventral border of the opercle (1)].

76. Orientation of the pectoral fin base: vertical (0), horizontal (1), inclined (2).

77. Origin of the pelvic fin: anterior to the dorsal fin (0); opposite or posterior to the dorsal fin (1).

78. Dorsal fin length: short [lesser than 20 rays (0]; long [more than 20 rays] (1).

79. Shape of the first proximal pterygiophore of the dorsal fin: different from the remnants (0); all equal in shape (1).

80. Anal fin length: short [up to 15 rays (0)]; long [more than 15 rays (1)].

81. Anal fin edge: not serrated (0); serrated (1).

82. Fusion of hypurals: free (0); fused (1).

83. Contact between hypurals 2-3: free (0); with convoluted suture (1).

84. Body scales: present (0); absent (1). 85. Flank scutes: absent (0); triangular (1); cordiform (2); tripartite (3); retangular (4).

86. Number of rows of scutes on flanks: absent (0); single (1); two or more (2).

87. Middorsal scutes: absent (0); present (1).

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Appendix II

Appendix I

Appendix III

Publication Dates

• Publication in this collection
  03 June 2011
• Date of issue
  June 2011

History

• Accepted
  22 Dec 2010
• Received
  24 Sept 2010