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Phylogeny of the Trachichthyiformes (Teleostei: Percomorpha)

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Phylogeny of the Trachichthyiformes (Teleostei: Percomorpha) BULLETIN OF MARINE SCIENCE, 52(1): 114-136, 1993 PHYLOGENY OF THE TRACHICHTHYIFORMES (TELEOSTEI: PERCOMORPHA) Jon A. Moore ABSTRACT The osteology and soft anatomy of many genera of fossil and living fishes considered to be in the order Beryciformes were studied in a phylogenetic analysis ofacanthomorph fishes. The result of this analysis is that the beryciforms, as they are presently accepted, represent a non-monophyletic group. The "beryciform" suborders Polymixioidei and Dinopterygoidei are actually basal acanthomorph lineages. The remaining "beryciforms," generally those put in the suborders Berycoidei and Stephanoberycoidei, are basal percomorphs. Of those, the Holocentridae are more closely related to the Perciformes, Scorpaeniformes and Zeiformes, rather than to the other beryciforms, and no unequivocal features have been found to unite the Berycidae to the rest of the so called berycoid and stephanoberycoid beryciforms. The residual berycoid and stephanoberycoid beryciforms, minus the Holocentridae and Berycidae, are monophyletic based on a number of characters that are found in Cretaceous fossils and all or most of the recent taxa. The Cretaceous fossils arrange in a series of three sister groups outside the clade of recent taxa. This clade of recent taxa has been called the Trachichthyi- formes and is here recognized to consist of at least 13 families of fishes, which are: the Diretmidae, Anoplogastridae, Anomalopidae, Monocentridae, Trachichthyidae, Melam- phaidae, Gibberichthyidae, Stephanoberycidae, Hispidobcrycidae, Rondeletiidae, Barbour- isiidae, Megalomycteridae, and Cetomimidae. Characters that support the monophyly of each family are briefly described. "Re-analysis of Zehren's data using a methodology that allows character reversal would be worthwhile and could result in a new phylogenetic hy- pothesis for the Beryciformes" (Johnson and Rosenblatt, 1988: p. 86). The beryciforms are recognized as a morphologically diverse group of primitive spiny-finned fishes. They have been repeatedly mentioned as the ancestors or intermediate forms leading to the Perciformes (Starks, 1904; Jordan, 1905; Regan, 1911, 1929; Gregory, 1933; Patterson, 1964). Such statements would indicate that the beryciforms are possibly a paraphyletic group. Because of the many unique features found among beryciforms, the monophyly for most families within the group has been largely uncontested. The monophyly of the group as a whole, however, is still debated. Patterson (1964) divided the beryciforms into 3 sub- orders: the Polymixioidei (containing Polymixia and a few extinct genera); the Dinopterygoidei (containing a diverse group of extinct taxa); and the Berycoidei (containing many living beryciform families including berycids and holocentrids). Patterson contended that the stephanoberycoid fishes represented a separate order. Greenwood et al. (1966), however, added the suborder Stephanoberycoidei to the beryciforms. Two widely cited studies (Rosen, 1973; Zehren, 1979) have discussed in detail some features and phylogenetic relationships ofliving beryciform fishes. Rosen's discussion was part of a larger work, and the portions pertaining to the beryciforms described numerous interesting features and their distributions, but was in no way a quantitative analysis. Rosen concluded that beryciform fishes could be divided into five groups: polymixiids, berycids, holocentrids, trachichthyoids, and ste- phanoberycoids. He found it difficult, however, to relate the first three groups to the latter two. Zehren's (1979) study was, on the other hand, an explicitly quan- titative analysis, but contained many difficulties and assumptions which weakened 114 MOORE: TRACHICHTHYIFORM PHYLOGENY 115 the results. Both authors suggested, however, that the beryciforms might not be a monophyletic group for two reasons. Rosen (1973) suggested that the Holocen- tridae are more closely related to the perciform fishes. Zehren (1979) also could find little to relate the Holocentridae to the other beryciforms. In addition, Zehren adequately showed that the Polymixiidae are not related to the other beryciform fishes. This second conclusion has more recently been corroborated by Rosen (1985) and Stiassny (1986), who indicated that Polymixia is the most basal acan- thomorph lineage. Presumably the fossil polymixiids (e.g., Berycopsis and Hom- onotichthys; Patterson, 1964) are part of that lineage. Little has been said about the higher level relationships of the extinct dinop- terygoid fishes since Patterson's work (1964, 1967, 1968), other than Gayet's (1980, 1982) assertions that the families Apichthyidae and Aipichthyoididae are paracanthopterygians and the genus Pycnosteriodes is in the superfamily Holo- centroidea, with the extant Holocentridae and other Cretaceous holocentroids. Moore (1993) has found that the dinopterygoid fishes are more appropriately placed as one or more lineages at the base of the Acanthomorpha near the po- Iymixiids. This is supported by the presence of two series of intramuscular bones in dinopterygoids; the loss of one series of these bones is evidently a synapomorphy for the Paracanthopterygii + Acanthopterygii (Johnson and Patterson, 1993; Moore, 1993). The basal Percomorpha considered in this analysis include the Berycidae, Hol- ocentridae, all other "beryciforms," Lampriformes, percoid Perciformes, scor- paenoid Scorpaeniformes, Zeiformes, Cretaceous beryciforms usually placed in the Trachichthyidae (Patterson, 1964), and the Cretaceous fossil fishes referred to as "holocentroids" (Stewart, 1984). Based upon two characters of the pelvic anatomy, Stiassny and Moore (1992) have found that the Holocentridae are more closely related to the Perciformes, Scorpaeniformes and Zeiformes, rather than to the other beryciformes. Also, no unequivocal features have been found to unite the Berycidae to the rest of the so called beryciforms. A few equivocal features of the caudal skeleton and gill arches potentially align the berycids with the holocentrids and more derived perco- morphs, rather than the other trachichthyoid and stephanoberycoid beryciforms (sensu Rosen, 1973). Thus the fish order Beryciformes, as currently perceived, appears not to constitute a monophyletic group. The residual trachichthyoid and stephanoberycoid beryciforms, minus the Hol- ocentridae and Berycidae, are monophyletic, however, based on a number of characters that are found in all or most of the recent taxa. This group of recent taxa has been called the Trachichthyiformes (Moore, 1990, 1991; Stiassny and Moore, 1992) and is here recognized to consist of two suborders with at least 13 families of fishes. The suborder Trachichthyoidei contains the families Diret- midae, Anoplogastridae, Anomalopidae, Monocentridae, Trachichthyidae, and the suborder Stephanobercoidei contains the families Melamphaidae, Gibberich- thyidae, Stephanoberycidae, Hispidoberycidae, Rondeletiidae, Barbourisiidae, Megalomycteridae, and Cetomimidae. Other families of fishes formerly placed among the beryciforms include the Paradiretmidae, which have been shown to be juvenile pomacanthids (Allen et aI., 1976), and the Sorosichthyidae, which have been placed within the Trachich- I thyidae (Gomon and Kuiter, 1987; Moore, ms. ). Monophyly can be established , Moore, J. A. Sorosichlhys ananassa and the phylogeny of the family Trachichthyidae (Teleoslei: Percomorpha: Trachichlhyiformes). MS. 116 BULLETIN OF MARINE SCIENCE, VOL. 52, NO. I, 1993 for all 13 families and apomorphic characters for each family will be discussed later in this paper. METHODS AND MATERIALS Polarization of characters for the Trachichthyiformes, and similarly for other major acanthomorph clades, is a difficult problem due to the transformed nature of many of the lineages. For example, the nearest extant outgroups for the Trachichthyiformes consists of the Berycidae, the Perciformes and other derived percomorphs, the Holocentridae, the Atherinomorpha, and possibly the Lampriformes. In the very least, the last three groups have been characterized as highly divergent taxa with numerous autapomorphies (Olney, 1984; Stewart, 1984; Stiassny, 1990). Maddison et al. (1984) discussed the problems of making character polarity decisions based on distant or highly transformed outgroups. They concluded that more closely related outgroups could overturn polarity decisions. In the absence of extant outgroups closely related to the ingroup, fossil outgroups have proven to be important in polarity decisions (Gauthier et aI., 1988; Norell, 1988; Donoghue et aI., 1989). The results of a larger study of acanthomorphs (Moore, 1993) show that the Cretaceous fossil species usually assigned to the family Trachichthyidae (Patterson, 1964, 1967, 1968; Gayet, 1980, 1982; Grande and Chatterjee, 1987) form a sequential series of sister groups to the recent Trachichthyiformes. The most plesiomorphic fossil sister group includes the genus Acragaster and possibly Gnathaberyx. The next sister group consists of the genera Lissaberyx, Libanoberyx and Stichopteryx. The third fossil sister group consists of the genus Hop/apteryx. and maybe Antarctiberyx as well. The characters for the present study ofthe Trachichthyiformes were polarized using these three fossil outgroups and the holocentrids and berycids in an unresolved polychotomy with the most plesiomorphic fossil taxa. A data matrix was constructed and run on PAUP 3.0 (Phylogenetic Analysis Using Parsimony, version 3.0, Swofford, 1990) using the general heuristic search. The data matrix and constraints can
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