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Interrelationships of Aulopiformes

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Interrelationships of Aulopiformes CHAPTER 14 Interrelationships of Aulopiformes CAROLE C. BALDWIN and G. DAVID JOHNSON Department of Vertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, D.C. 20560 I. Introduction relationships in which Rosen's (1973) aulopiforms and myctophiforms are united in the order Myctoph- In 1973, Rosen erected the order Aulopiformes for iformes. all non-ctenosquamate eurypterygians, that is, the Ini- In 1985, Rosen altered his concept of a monophy- omi of Gosline et al. (1966) minus the Myctophiformes letic Aulopiformes, noting that Aulopus shares several (Myctophidae and Neoscopelidae). Rosen's aulopi- derived features with ctenosquamates, most notably forms included 15 families (Alepisauridae, Anotopter- a median rostral cartilage. Hartel and Stiassny (1986) idae, Aulopidae, Bathysauridae, Bathypteroidae, considered a true median rostral cartilage a character of acanthomorphs and concluded that the morphol- Chlorophthalmidae, Evermannellidae, Giganturidae, ogy of the rostral cartilage is highly variable below Harpadontidae, Ipnopidae, Omosudidae, Paralepidi- that level. Nevertheless, Stiassny (1986) supported dae, Scopelarchidae, Scopelosauridae, and Synodon- Rosen's (1985) view of a paraphyletic Aulopiformes, tidae) and 17 fossil genera, a morphologically diverse proposing that Chlorophthalmus, Parasudis and Aulopus gro^u^p of benthic and pelagic fishes that range in habi- form the sister group of ctenosquamates based on an tat from estuaries to the abyss. elevated, reoriented cranial condyle on the maxilla Rosen (1973) diagnosed the Aulopiformes by the and concurrent exposure of a "maxillary saddle" for presence of. an elongate uncinate process on the reception of the palatine prong. G.D. Johnson (1992) second epibranchial (EB2) bridging the gap between discussed the shortcomings of Rosen's (1985) analysis a posterolaterally displaced second pharyngobran- and observed that neither Rosen nor Stiassny (1986) chial (PB2) and the third pharyngobranchial (PB3). mentioned the distinctive gill-arch configuration origi- He noted that paralepidid fishes lack this distinctive nally described by Rosen (1973) as unique to aulopi- configuration of EB2 and thus questioned their place- forms. He added an additional gill-arch character to ment in the order. Subsequently, R. K. Johnson Rosen's (1973) complex, the absence of a cartilaginous (1982) recognized that certain paralepidids (Paralepis, condyle on PB3 for articulation of EB2, and concluded and Notolepis) have an enlarged EB2 uncinate process that a suite of gill-arch modifications constitutes a but questioned Rosen's use of this feature to diag- complex specialization supporting the monophyly of nose aulopiforms because he believed the same Rosen's (1973) Aulopiformes. In addition, Johnson condition occurs in neoscopelids. Instead he sug- (1992) offered further evidence (absence of the fifth gested that the modification is a primitive iniome upper pharyngeal toothplate and associated third in- condition and that the small EB2 uncinate process ternal levator muscle) for the monophyly of Rosen's of myctophids is secondarily derived. R. K. Johnson (1973) Ctenosquamata, which include myctophids, (1982) resurrected a more traditional view of iniome neoscopelids and acanthomorphs, but not aulopi- Copyright © 1996 by Academic Press, Inc. 1NTERRELATIONSHIPS OF FISHES 355 All rights of reproduction in any form reserved. 356 CAROLE C. BALDWIN AND G. DAVID JOHNSON Gosline et al. (1966): Rosen (1973): Order Inioml Order Auloplformes, new name Myctophoidea Suborder Aulopoidei, new name Aulopidae Aulopidae Bathysauridae Bathysauridae Synodontidae Bathypteroidae Harpadontldae Ipnopidae Bathypteroidae Chlorophthalmidae Ipnopidae Notosudidae (=Scopelosauridae) Chlorophthalmidae Suborder Alepisauroidei Notosudidae (=Scopelosauridae) [15 fossil genera] Myctophidae Superfamily Synodontoidea, new usage Neoscopelidae [2 fossil general Alepisauroidea Synodontidae Paralepldidae Harpadontldae Omosudldae Giganturidae (? + Rosauridae) Alepisauridae Superfamily Alepisauroidea Anotopteridae Paralepldidae Evermannellidae Omosudldae Scopelarchidae Alepisauridae Anotopteridae Evermannellidae Scopelarchidae Sulak (1977): Benthic Myctophiformes: Aulopidae R. K. Johnson (1982): Aulopus (including Hime, Latropiscus) Myctophiformes: Synodontidae Aulopoids Subfamily Harpadontlnae Aulopidae Harpadon (incl. Peltharpadoril Myctopholds + Chlorophthalmoids Saurida Myctophoids Subfamily Bathysaurinae Myctophidae Bathysawus (incl. Macrtstium) Neoscopelidae Subfamily Synodontinae Chlorophthalmoids Syrwdus (incl. Xystodus) Notosudidae Trachinocephalus Scopelarchidae Chlorophthalmidae Chlorophthalmidae Subfamily Chlorophthalminae Ipnopidae Chlnrophthalmus Synodontoids + Alepisaurolds Parasudis Synodontoids Bathysauropsts (incl. Bathysaurops) Synodontidae Subfamily Ipnoplnae Harpadontldae Tribe Ipnopini Bathysauridae Ipnops (incl. Ipnoceps) Alepisaurolds Tribe Bathypteroinl Paralepldidae Bathypterois (incl. Benthosaurus) Anotopteridae Tribe Bathymicropini Evermannellidae Bathymlcrops Omosudldae Bathytyphlops (incl. Macristiella) Alepisauridae FIGURE 1 Four previously hypothesized classifications of aulopiform or myctophiform fishes. forms (see also Stiassny, this volume). Johnson et al. et al. (1966) recognized two "suborders": myctophoids (1996) argued that Aulopus is not closely related to (Aulopidae, Bathysauridae, Synodontidae, Harpa- ctenosquamates but is the cladistically primitive mem- dontldae, Bathypteroidae, Ipnopidae, Chlorophthal- ber of their Synodontoidei, a lineage that also includes midae, Notosudidae [= scopelosaurids of Marshall, Pseudotrichonotus, Synodus, Trachinocephalus, Saurida, 1966—see Paxton, 1972; Bertelsen et al, 1976], Myc- and Harpadon. Finally, Patterson and Johnson (1995) tophidae, and Neoscopelidae); and alepisaurolds (Par- provided corroborative evidence from the intermuscu- alepldidae, Omosudldae, Alepisauridae, Anotopteri- lar bones and ligaments for Rosen's (1973) Aulopl- dae, Evermannellidae, and Scopelarchidae). Rosen formes, in the extension of the epipleural series anteri- (1973) added synodontids and harpadontids (his syn- orly to the first or second vertebra. odontoids) and 17 fossil genera to the Alepisauroidei, Various schemes of relationships among iniomous described a new suborder, the Aulopoidei, for Aulopi- fishes have accompanied confusion about the recogni- dae, Bathysauridae, Bathypteroidae, Ipnopidae, tion of a monophyletic Auloplformes (Fig. 1). Gosline Chlorophthalmidae, and Notosudidae and, as noted, 14. Interrelationships of Aulopiformes 357 restricted the Myctophiformes to myctophids and notosudids and the evermannellid-scopelarchid neoscopelids. lineage, and bathysaurids and giganturids; and a Sulak (1977) examined aspects of the osteology of paraphyletic Paralepididae, with Paralepis forming the the benthic "myctophiforms" and envisioned them sister group of a monophyletic clade comprising Omo- forming two divergent lineages exhibiting progres- sudis and Alepisaurus. Patterson and Johnson (1995) sively greater differentiation from the basal aulopid noted that the paraphyly of the Paralepididae sug- body plan, an expanded Synodontidae that included gested by their data may be artificial, a result of the bathysaurids, synodontids, and harpadontids, and an greatly reduced number of intermuscular elements in expanded Chlorophthalmidae for chlorophthalmids Macroparalepis. (including Bathysauropsis) and ipnopids (including No other comprehensive studies of aulopiform rela- bathypteroids). tionships have been undertaken, and thus consider- To examine a previously proposed relationship be- able conflict about the evolutionary history of aulopi- tween the Evermannellidae and Scopelarchidae (e.g., form fishes existed when we initiated this study, the Marshall, 1955; Gosline et al., 1966), R. K. Johnson goal of which was to hypothesize a phylogeny of ex- (1982) studied the distribution of selected characters tant aulopiform genera based on cladistic analysis of among iniomes. He did not present a formal classifi- a wide range of morphological data. R. K. Johnsons's cation but described three perceived iniomous clades. (1982) cladistic analysis of iniome relationships used One comprised only aulopids, a second was equiva- commonality rather than outgroup comparison to as- lent to Rosen's (1973) alepisauroids minus scopelar- sess character polarity, and we thus found that many chids, and the third included myctophids, neoscopel- of his polarity decisions were reversed in our analysis. ids, chlorophthalmids, ipnopids, notosudids, and Patterson and Johnson's (1995) phylogeny is of limited scopelarchids. R. K. Johnson's (1982) phylogeny cor- value because it was constructed on the basis of a roborated Sulak's (1977) placement of bathysaurids in single complex. Despite their shortcomings these pub- the synodontid + harpadontid lineage, but he noted lications, as well as those of Rosen (1973), Sulak (1977) that only two clades resulting from his analysis, the and Okiyama (1984b), proved useful in this study, and myctophoids (Myctophidae, and Neoscopelidae) and we derived many informative characters from them. the alepisauroids (Paralepididae, Anotopteridae, Evermannellidae, Omosudidae, and Alepisauridae) were well supported. II. Methods Okiyama (1984b) examined R. K. Johnson's (1982) hypothesis in light of evidence from aulopiform lar- Osteological abbreviations are listed in Appendix vae. He did not produce an independent hypothesis 1, and a full list
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