BULLETIN OF MARINE SCIENCE, 52(1): 114-136, 1993

PHYLOGENY OF THE (TELEOSTEI: )

Jon A. Moore

ABSTRACT The osteology and soft anatomy of many genera of fossil and living considered to be in the order 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 are more closely related to the , and , rather than to the other beryciforms, and no unequivocal features have been found to unite the 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 fossils and all or most of the recent taxa. The Cretaceous fossils arrange in a series of three sister groups outside the 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, , , Trachichthyidae, Melam- phaidae, Gibberichthyidae, , Hispidobcrycidae, Rondeletiidae, Barbour- isiidae, Megalomycteridae, and . 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 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 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 + (Johnson and Patterson, 1993; Moore, 1993). The basal Percomorpha considered in this analysis include the Berycidae, Hol- ocentridae, all other "beryciforms," , 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 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 , 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 , 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 be found in Table I. The fishes examined in this study were cleared and stained according to the methods described by Dingerkus and Uhler (1977) and Potthoff(1984). In addition, data collection was supplemented with intact and partially dissected alcohol specimens, a few dried skeletons, and with x-rays of alcohol specimens. Fossil specimens were examined when possible; however, a majority of the character information for fossil taxa was taken from the literature. For the most part, sub-adult and adult specimens were examined, although a few juvenile and larval specimens were also studied. This study was limited to mostly adult features, because until very recently only a few larvae and juveniles oftrachichthyiform taxa were known (Keene and Tighe, 1984). With the continued discoveries of larval and juvenile stages of a number of trachichthyiform taxa, a full study of features of these stages as well as ontogenetic transformations should be conducted. Dissections and observations were performed under a Wild M5 stereomicroscope and drawings were made with the aid of a camera-Iucida attachment. A full list of material examined can be found at the end of this paper in Appendix I. Institutional abbreviations are those of Levinton et al. (1985) except for the following: ANSP (VP) Paleontology Collection, Academy of Natural Sciences, Philadelphia; FMNH PFVertebrate Paleontology Collection, Field Museum of Natural History, Chicago; and YPM-PU Princeton Uni- versity Vertebrate Paleontology Collection, now housed at the Peabody Museum of Natural History, Yale University.

CHARACfERS Two Ocular Sclera Absent from Cartilaginous Ring Surrounding Eye. - In the Acanthomorpha, two ossified sclera are normally found on the anteriormost and posteriormost portions of the cartilaginous ring which circles the eyeball. The size and shape of these sclera are quite variable among taxa. Among the living trach- ichthyiforms, however, these ocular sclera are completely absent from all taxa studied, but found in all closely related acanthomorph lineages outside the trach- ichthyiforms. The condition in the fossil outgroups has not been determined, and until then it seems best to consider this character as a synapomorphy for the Trachichthyiformes. Neural Arch of First Vertebra Fused to Centrum. - In most percomorphs, the neural arch and spine of the first vertebra (NAY1) is autogenous (e.g., berycids, MOORE: TRACHICHTHYIFORM PHYLOGENY 117

Table I. Data matrix of character states for study taxa

Character'

I-S 6-10 II-IS 16-20 21-24 Berycidae 00000 00000 00000 00000 0000 Holocentridae 00000 00000 00000 00000 0000 Acrogaster ??OOO omo 00070 70000 0000 Lissoberyx ??OOO omo 00070 70000 0000 Hop/opteryx 70000 omo 00070 00000 0000 Anoplogasteridae 11111 11100 10000 00000 0000 Diretmidae 11111 11100 00000 00000 1000 Anomalopidae 11111 10011 00000 00000 0000 Monocentridae 11111 10011 20000 00000 0000 Trachichthyidae 11111 10011 20000 00000 0000 Melamphaidae 11100 00000 31110 01000 1000 Gibberichthyidae 11120 00000 31111 12000 0000 Hispidoberycidae 11120 00000 31111 12100 0000 Stephanoberycidae 11120 00000 31111 12100 0000 Rondeletiidae 11100 00000 31111 12010 0000 Barbourisiidae 11100 00000 31111 13011 1000 Ataxo/epis 11100 00000 31111 13011 1111 Cetomimidae 11100 00000 31111 13011 1111

I List of characters and character states: I. Ocular sclera~0 = present, J = absent. 2. Neural arch of first vertebra; 0 = autogenous, I = fused to centrum. 3. Number of supramaxillae; 0 = two, I = one. 4. Pallern of frontal ridges; 0 = various pallerns of converging or parallel ridges, I = modified X pallern, 2 = Y-shaped pallern. S. Bony arches over infraorbitals; 0 - absent, I - present. 6. Meseth- moid; 0 = large, I = small and confined to region between upper lateral ethmoids. 7. Larval head spines on parietals and frontals; o = absent, I = present. 8. Relation of second pharyngobranchial (PB2) to second epibranchial (EB2); 0 - in line and directly contacting, I = posterior PB2 underneath anterior EB2. 9. Founh pharyngobranchialtoothplate; 0 = present, I = absent. 10. Neural arch of second preural centrum; 0 = fused, I = unfused. II. Subocular shelf; 0 = continuous on two or more suborbitals or very wide-based on 10J, I = replaced by expanded orbital margin io 3. 2 = restricted to io 3 and narrow-based, tapered. and often hooked anteriorly. 3 = absent. 12. Orbitosphenoid; 0 = present. I = absent. 13. Cartilaginous neurocranium; 0 = absent, 1 = present. 14. Lower branchial tooth patches; o = presenl, I = losl. IS. Abdominal haemal arches; 0 = present, I = losl. 16. Exoccipital condyles; 0 = contact, I = widely separated. 17. Posterior placement of and insertion of first dorsal pterygiophore; 0 = somewhere anterior to fourth neural spine, I = betweeo founh and fifth neural spine, 2 = anterior to eighth or ninth neural spine, 3 = posterior to fourteenth neural spine. 18. Procurrent spines in caudal fin; 0 "'"fewer than 9 in either dorsal or ventral lobe, 1 = 9-11 in dorsal and ventral lobes. 19. Complete loss of fin spines; 0 = spines present, I = spines lost. 20. Increase in number of vertebrae; 0 = less than 38, I = 38 or more. 21. Basisphenoid; 0 = present, 1 = absent. 22. Haemal and neural spines on second pre ural centrum; 0 = single haernal spine and low neural crest, I = two spines on both haemal and neural arches. 23. Posttemporal and intercalar; 0 = forked and connected to cranium, intercalar present, I = splint-like and disconnected, intercalar abseot. 24. Pleural ribs; 0 = present, I - losl.

holocentrids, basal perciforms, scorpaeniforms; see Rosen, 1985). Hop/opteryx, the first outgroup for the trachichthyiforms, also has an autogenous NAYl. The condition in the other fossil outgroups is unknown since the first vertebra is usually covered by the bones of the skull. All living trachichthyiforms have the neural arch of the first vertebra fused to the underlying centrum. This feature is not unique to the trachichthyiforms (many derived paracanthopterygians, atherinomorphs, and some derived perciforms such as Echeneis, have this condition of the NAYI fused to the centrum [Rosen, 1985, figs. 33, 34, 38]). However, due to the absence of this feature in the first fossil outgroup, the holocentroid and berycid outgroups, and other basal percomorphs, the fusion of the NAYI to the underlying centrum in the living trachichthyiforms is presumed to be an independent derivation. Number of Supramaxillae. - The plesiomorphic condition for this feature in most primitive is the presence of two supramaxillae on the posterior end of the maxilla. The anterior supramaxilla is usually very small and the posterior supra- maxilla is relatively larger and often has a process over the dorsal portion of the anterior supramaxilla. Although reductions in the number of supramaxillae occur in numerous acanthomorph lineages, the loss of the anterior supramaxilla is a viable character for the Trachichthyiformes. The berycids, holocentrids, and the three fossil outgroups all possess two supramaxillae. All Trachichthyiformes (ex- cept some species of anomalopids and the melamphaid Scope/ogadus) have only 118 BULLETIN OF MARINE SCIENCE, VOL. 52, NO. I, 1993

pI

Figure L Dorsal view of the cranium of Slephanoberyx monae (FMNH 77987) exhibiting the Y-shaped frontal crests. Nasals were removed and are not shown. Abbreviations: ds, dermosphenotic; ex, extrascapular; fr, frontal; me, mesethmoid; pa, parietal; pt, pterotic; soc, supraoccipital.

the posterior supramaxilla. Johnson and Rosenblatt (1988) hypothesized that the two supramaxillae in some anomalopid genera are an independent derivation. The melamphaid genus Scopelogadus is unique among trachichthyiforms because both supramaxillae have been lost (Ebeling and Weed, 1963, 1973). Therefore, the most parsimonious distribution of these character states has the reduction to a single supramaxilla defining the trachichthyiforms, with an independent deri- vation of two supramaxilae in one clade within the Anomalopidae and the au- tapomorphous loss of both supramaxillae in Scopelogadus. Pattern of Cranial Ridges on Frontal. - It is possible that the patterns of cranial ridges among all trachichthyiforms are related. It is easy to imagine the transfor- mations that could produce the various patterns found in the entire group. It may also be that the patterns of cranial ridges in trachichthyiforms are somehow related to those of the fossil outgroups, berycids, and other percomorphs. However, until better comparative material oflarvae and juveniles oftrachichthyiforms becomes available to investigate possible ontogenetic changes, it is best to concentrate here on two very distinctive and putatively synapomorphous patterns. Trachichthyoids have a distinctive pattern of ridges on the frontal bone con- sisting of a modified X pattern (Zehren, 1979). The ridges all meet at a point over the eye and from there the two anteriorly directed ridges extend to the antero- medial and anterolateral corners of the frontal. From over the eye, three ridges are posteriorly directed towards the supraoccipital, parietal, and pterotic. This MOORE: TRACHICHTHYIFORM PHYLOGENY 119

105

Figure 2. Lateral view of right infraorbital series of Hoploslelhus occidenla/is (USNM 214196) dis- playing the complete bony arches on the first (io I), third, fourth and fifth (io 5) infraorbital bones. Scale bar equals 2 mm. pattern is illustrated in Patterson (1964, figs. 63, 65) for Monocentris and Hoplo- stethus. Zehren (1979) described and illustrated this feature for a number of trachichthyoid fishes (Anoplogaster, Monocentris, and Aulotrachichthys [his Para- trachichthys]); however, he never included it in his computational analysis of characters. Rosen (1973) also discussed this feature for the trachichthyoids in terms of a basic pattern of "membrane covered grooves and cavities." The X pattern is only found in the early juvenile stages of the Diretmidae. In all other trachichthyoid genera it is evident in the adults. I consider this modified X pattern of frontal ridges as a synapomorphy for the suborder Trachichthyoidei. Gibberichthyids, stephanoberycids, and hispidoberycids share a unique Y-shaped frontal crest pattern, as briefly discussed by Zehren (1979) and illustrated by Kotlyar (1980, 1990a). On the frontal bone of these fishes, the arms of the Y extend anteromedially to meet the counterpart from the other frontal and an- terolaterally to the lateral edge of the frontal and the posterolateral portion of the nasal (Fig. 1). The base of the Y extends posteriorly to the extra scapular and/or parietal. This Y-shaped pattern is not found in any ofthe other trachichthyiforms and is considered a synapomorphy for these three families. Complete Bony Arches over Infraorbitals. -Among acanthomorphs, the infraor- bitals (io) form an open channel for the sensory canal. The orbital edge of the infraorbital bones is rolled over to form a partial bony covering of the sensory canal. The rest of the sensory canal is covered by membrane and filled with mucus. This is the condition found in the fossil outgroups and the stephanoberycoids. In the Trachichthyoidei, however, complete bony "strut-like" arches (Rosen, 1973) cross over from the rolled-up orbital edge of the infraorbitals to the lower edge of the same bones. In most of the trachichthyoids these arches are very narrow bridges crossing over the anterior lachrymal (=io 1), in all cases io 3, very often io 4 and io 5, and occasionally io 2 as well (as in Anoplogaster). A typical example can be seen in the trachichthyid Hoplostethus (Fig. 2). Anomalopids have modified much of the ornamentation of the cranial bones and this includes the infraorbitals where the bony arches are often wide flat bridges that partially or completely cover some of the infraorbitals (Johnson and Rosenblatt, 1988, figs. 5, 8). Bony coverings of the infraorbital sensory canal can be found in numerous primitive 120 BULLETIN OF MARINE SCIENCE, VOL. 52, NO. I, 1993 teleosts (Nelson, 1969a), but not among the acanthomorphs. Possibly, to some extent, the bony arches in trachichthyiforms represent an atavistic rederivation of this bony covering. However, parsimony requires that the presence of complete bony arches over the lachrymal and the third infraorbital, and very often other infraorbitals as well, is considered independently derived and a synapomorphy for the Trachichthyoidei. Ethmoid Small and Confined to Region between Dorso-medial Portions of Lateral Ethmoids. -Zehren (1979) pointed out that among the trachichthyoids the eth- moid bone ("mesethmoid") is very small and confined to the area between the upper portions of the lateral ethmoids. Among the stephanoberycoids, as well as other acanthomorphs, including holocentrids, berycids, zeoids, perciforms and other percomorph groups, the ethmoid is larger and extends anterior to the dorsal portions of the lateral ethmoids (Stiassny, 1986, figs. 17-22 and note difference between Diretmus, fig. 21c, and all other examples). Although Patterson (1964) states that Hoplopteryx has a "very small" ethmoid, it is illustrated (fig. 47) as extending anteriorly relative to the lateral ethmoids, unlike the trachichthyoids. The condition is not determinable in other fossil outgroups at this time. Based on this distribution it appears that a small ethmoid confined to the area between the upper portions of the lateral ethmoids is an autapomorphy of the Trachichthy- oidei. Larval Head Spination.-Larval Anoplogastridae and Diretmidae share the un- usual feature of prominent spines projecting posteriorly from the parietals (Keene and Tighe, 1984, fig. 208 A, B). Johnson (1984) discussed the unique nature of parietal spines in larval percomorph fishes and stated that they are found elsewhere only among the scorpaeniforms. The presence of this spine defines the clade Anoplogastridae + Diretmidae. Relationship of Second Pharyngobranchial to Second Epibranchial. - In most per- comorphs examined, the posterior end of the second pharyngobranchial (PB2) is in line with and directly contacts the anterior tip of the second epibranchial (EB2). In diretmids and anoplogastrids, however, that direct contact is lost and instead the posterior end of the second pharyngobranchial extends underneath the anterior tip of the second epibranchial. Connective tissue apparently maintains the con- nection between the posterior PB2 and anterior EB2. This arrangement was not seen anywhere else and is taken to be another synapomorphy for these two families. Fourth Pharyngobranchial Toothplate Absent.-Rosen (1973) noted that among the Anomalopidae + Monocentridae + Trachichthyidae the fourth pharyngo- branchial (PB4) toothplate is absent and a large third pharyngobranchial (PB3) tooth plate is immovably fixed to the third pharyngobranchial. This is in contrast to the primitive acanthomorph condition ofa fourth pharyngobranchial toothplate present as a separate plate posterior to the third tooth plate, as found in diretmids, anoplogastrids, most stephanoberycoids (except Stephanoberyx), berycids, holo- centrids, and many other percomorphs and acanthomorphs. Stephanoberyx ap- pears to be a case where the fourth toothplate is simply lost, but since a separate fourth tooth plate can be found in all other stephanoberycoids, this absence is independent to that found in trachichthyoids. Rosen hypothesized that the absence of the fourth toothplate in the more derived trachichthyoids may have been due to a fusion of that toothplate to the third. Examination of juvenile Hoplostethus (20-30 mm, MCZ 70173) reveals no sep- arate fourth toothplate nor a suture anywhere within the large tooth plate on the MOORE: TRACHICHTHYIFORM PHYLOGENY 121

PB3. The posterior portion of this large tooth plate does, however, extend back below the ends of the third and fourth epibranchials and occupies the area where the separate fourth tooth plate is usually found in basal percomorphs. A suture is apparently evident in some smaller specimens of Hoplostethus (G.D. Johnson, pers. comm.). The absence of a separate fourth toothplate, most likely due to fusion to the posterior of the third toothplate, is unique within the trachichthyi- forms and is restricted to the Anomalopidae + Monocentridae + Trachichthyidae clade. Neural Arch of Second Preural Centrum Autogenous. -Zehren (1979, table 73) listed a number of outgroup taxa in which the neural arch of the second preural vertebra (PU2) is either fused to the centrum or autogenous. Zehren concluded that an autogenous second preural neural arch is primitive for beryciforms. This conclusion highlights one of the problems with Zehren's analysis. None of the outgroup taxa Zehren selected are acanthomorphs, except for four perciforms he examined. Maddison et al. (1984) discussed the problems associated with using distantly related outgroups. The neural arch of the second preural vertebra is fused to the centrum in paracanthopterygians, atherinomorphs, scorpaeniforms, zei- forms, polymixiids, holocentrids, berycids, stephanoberycoids, diretmids, ano- plogasterids, and perciforms. Thus, among acanthomorphs, an autogenous neural arch on the second preural centrum is unique to the Anomalopidae + Monocen- tridae + Trachichthyidae. Subocular Shelf. -In his discussion of the trachichthyoid beryciforms, Rosen (1973) mentioned a small "thumb-like" subocular shelf on the third infraorbital, a feature that is unusual among the basal percomorphs examined. The primitive condition for the subocular shelf in ctenosquamates is a continuous shelf on the second through fifth infraorbitals as found in myctophids (Paxton, 1972), poly- mixiids, and holocentrids. In some acanthomorph fishes, this shelf is somewhat reduced so that it is only found on the second and third or second through fourth infraorbitals. Berycids vary with the shelf on the second through fourth infraor- bitals (Centroberyx affinis), second and third (Beryx decadactylus), or third (B. splendens). The fossil outgroups to the trachichthyiforms also have variously reduced the shelf; Acrogaster has a continuous shelf on the second and third infraorbitals (Gayet, 1982), Lissoberyx has a "broad" shelf on the third infraorbital (Patterson, 1967), and Hoplopteryx has a large semicircular shelf on the third infraorbital (Patterson, 1964). For the fishes mentioned above with a subocular shelf restricted to the third infraorbital, that shelf is wide-based and originates from most of the length of the infraorbital bone. Within the trachichthyiforms, configuration of the subocular shelf is also quite variable. Anoplogastrids lack a true subocular shelf but have a distinct and unique thickening of the entire upper portion of the third infraorbital where the shelf would normally be found. Diretmids have a more plesiomorphic continuous shelf on the second and third infraorbitals. Anomalopids have a subocular shelf on the second and third infraorbitals that is uniquely discontinuous to accommodate the subocular light organ. A subocular shelf is entirely absent in the Stephanobery- coidei (a synapomorphy for that clade). Among the trachichthyiforms, only the monocentrids and trachichthyids have a subocular shelf restricted to the third infraorbital. It is narrow-based and originates from less than half the length of the infraorbital bone, extending slightly further under the eye, tapering, and often hooked anteriorly. This is considered a synapomorphy of the two families Mono- centridae + Trachichthyidae. 122 BULLETIN OF MARINE SCIENCE, VOL. 52, NO. I, 1993

Orbitosphenoid Absent. - The absence of an orbitosphenoid has been used by a number of authors to distinguish the stephanoberycoids from the other beryci- forms. An orbitosphenoid is present in the Trachichthyoidei, the fossil outgroups to the trachichthyiforms, the Holocentridae and the fossil "holocentroids" (Pat- terson, 1964; Stewart, 1984), the Berycidae, and other putative basal percomorphs such as Velifer and Lampris. Despite similar losses in other acanthomorph lineages (e.g., paracanthopterygians and atherinomorphs), the absence of the orbitosphe- noid corroborates the monophyly of the stephanoberycoids. Cranium Consisting Mostly of Cartilage and Connective Tissue with Thinly Ossified Bones. -Among all the stephanoberycoids, there is a tendency towards decreased ossification of the skeleton. The cranial bones are thinly ossified lamina over a neurocranium consisting largely of cartilage and connective tissue. It is very easy to see the large extent of cartilage and connective tissue in cleared and counter- stained specimens, although the connective tissue is also often apparent in alcohol specimens of melam phaids, for instance, as a white region underneath the frontals or inside the orbits. Loss of All Lower Branchial Tooth Patches. -In primitive teleosts, tooth patches that are not rudimentary gill rakers are associated with the lower branchial bones, particularly the basi- and hypo-branchials. Nelson (I969b, figs. 8-12) showed that "basibranchial" tooth patches are primitive for teleosts and illustrated numerous examples of tooth patches in trachichthyids, monocentrids, holocentrids, berycids, and perciforms. Zehren (1979) apparently had a more restricted definition for basibranchial tooth patches than did Nelson. For example, Zehren described the holocentrid Ostichthys as lacking basibranchial tooth patches, whereas Nelson illustrated tooth patches in that genus (Nelson, 1969b, fig. 10C). The patches Nelson illustrated are found on the hypobranchials, but do constitute tooth patches that are homologous with those found over the basibranchials in other holocen- trids. Therefore, many taxa that Zehren (1979) characterized as lacking basibran- chial tooth patches (e.g., Ostichthys, Plectrypops) possess lower branchial tooth patches. Tooth patches on the basi- and hypo-branchials are found in holocentrids, berycids, and trachichthyoids (except for some species of anomalopids). Lower branchial tooth patches are lacking in stephanoberycoids. Cetomimids are exceptional in having unusual median tooth patches, referred to as copular teeth (Paxton, 1989), which in most genera are fused into one median plate that extends over two or more basibranchials. These copular teeth are considered an autapomorphy of the Cetomimidae (Paxton, 1989). The absence of lower bran- chial tooth patches is considered here an autapomorphy of the Stephanobery- coidei, and the copular teeth of cetomimids are an independent derivation. Abdominal Haemal Arches Lost. - Rosen (1973) discussed in detail the presence of abdominal haemal arches of various morphologies in many percomorphs in- cluding Velifer, Lampris, all trachichthyoids, the fossil genus Hoplopteryx, all holocentrids, the berycid Centroberyx, zeiforms, and percoids. Abdominal haemal arches are also found in the fossil outgroup taxa Acrogaster (Gayet, 1980). Within stephanoberycoids, the Melamphaidae have abdominal haemal arches on the last three abdominal vertebrae, but the other stephanoberycoids lack them entirely. Exoccipital Facets Do Not Contact Each Other. - The exocciptal facets, which articulate with the first vertebra, suturally contact each other below the foramen magnum and above the basioccipital facet in most acanthomorphs. This is well illustrated in a number of taxa, such as Polymixia (Stiassny, 1986, fig. 12b), the fossil "holocentroid" Caproberyx (Patterson, 1967, fig. 68), Holocentrus (Rosen, MOORE: TRACHICHTHYIFORM PHYLOGENY 123

1985, fig. 29), various "basal" perciformes (Rosen, 1985, figs. 21-24, 26-29B), the trachichthyiform outgroup taxa Hoplopteryx (Patterson, 1964, figs. 48, 57), and the trachichthyid Hoplostethus (Starks, 1904, fig. 2). This contact is found in berycids, all trachichthyoids, and in the Melamphaidae, but is lacking in all non- melamphaid stephanoberycoids, where the exoccipital facets are widely separated and the foramen magnum is at the dorsal margin of the basioccipital facet. Loss of contact between the exoccipital condyles is taken to be an autapomorphy for that group. Posterior Placement of Dorsal Fin. - In most basal percomorphs the dorsal fin originates anterior to the midbody and often anterior to the anal fin. This is true of holocentrids, berycids, veliferids, percoids, scorpaenoids, caproids, zeoids, trachichthyoids, and melamphaids. In other stephanoberycoids, however, the dorsal fin originates posterior to the midbody and symmetrical to the anal fin. Among these fishes, only adult Gibberichthys does not have the dorsal and anal fins symmetrically arranged, but this genus has enlarged bases of its fin spines, most of which are autapomorphically fused to the underlying pterygiophores. This results in a wider spacing of the anterior pterygiophores and may obscure this symmetry. In larvae and juveniles of Gibberichthys (de Sylva and Eschmeyer, 1977), it can be seen that the median fins are posterior and symmetrical. This feature of posterior and symmetrical placement of median fins is a character for the stephanoberycoids exclusive of the Melamphaidae. Related to the posterior shift in the position of the dorsal fin is a change in the interdigitation of the first dorsal-fin pterygiophore relative to the vertebral neural spines. In the trachichthyoids, fossil outgroups to the trachichthyiforms, berycids, holocentrids, and "holocentroids," the first dorsal-fin pterygiophore inserts some- where anterior to the neural spine of the fourth vertebra. In melamphaids the first dorsal pterygiophore interdigitates between the fourth and the fifth neural spines. In Gibberichthys, Stephanoberyx, Hispidoberyx and Rondeletia the first dorsal pterygiophore interdigitates anterior to the eighth or ninth neural spine. In Barbourisia, megalomycterids, and cetomimids, the first dorsal pterygiophore interdigitates posterior to the 14th neural spine or even further back. When treated as an unordered transformation series, the various posterior shifts of the dorsal fin pterygitophore insertion described above define three nodes, namely the Steph-' anoberycoidei, the non-melamphaid stehanoberycoids, and the barbourisiids + megalomycterids + cetomimids.

9-11/9-11 Procurrent Spines. - Stephanoberycids and hispidoberycids share a dis- tinctive feature of the procurrent spines in the caudal fin. The number of procurrent spines for these two families (9 to II spines in both the dorsal and ventral lobes of the caudal fin) is higher than in any other acanthomorph fish.

Complete Loss of Fin Spines. - The cetomimoids (Rondeletiidae, Barbourisiidae, Megalomycteridae, and Cetomimidae) are characterized by the loss of dorsal-, anal- and pelvic-fin spines. Cetomimoids also lack the caudal procurrent spines and instead have procurrent soft rays, unlike any other trachichthyiforms, fossil outgroups, berycids or holocentrids. Fin spines have apparently been lost inde- pendently in some other percomorphs (i.e., monocentrids lack anal spines, di- retmids lack dorsal and anal spines, anoplogastrids lose dorsal, anal and pelvic spines). All these lineages, however, retain caudal procurrent spines. Complete absence of dorsal, anal, pelvic, and caudal procurrent spines is diagnostic for the cetomimoids. 124 BULLETIN OF MARINE SCIENCE, VOL. 52, NO.1, 1993

Increase in Vertebral Number. -Most trachichthyiforms and most other basal percomorphs have between 24-31 total vertebrae. Some specimens of stephanobe- rycids have as many as 33, and the hispidoberycids have 34. Barbourisia, me- galomycterids, and cetomimids have from 38 to 59 and in most cases more than 40 (Bertelsen and Marshall, 1984; Paxton, 1989). An increased number of ver- tebrae (38 or more) is recognized here as a synapomorphy for the Barbourisiidae + Megalomycteridae + Cetomimidae. Basisphenoid Absent. -Barbourisia, megalomycterids, and cetomimids share the loss of the basisphenoid. This bone is also lost in melamphaids and in diretmids (Zehren, 1979). The presence of the basisphenoid in other percomorphs (i.e., Velifer, berycids, holocentrids, percoids), fossil outgroups for the trachichthyi- forms, anomalopids + monocentrids + trachichthyids, Anoplogaster, rondele- tiids, gibberichthyids, stephanoberycids and hispidoberycids indicates that losses in diretmids, melamphaids, and Barbourisia + megalomycterids + cetomimids are most parsimoniously treated as the result of three independent losses within the trachichthyiforms. Two Spines on Neural and Haemal Arches of Second Preural Centrum. - The basal percomorphs, such as the holocentrids, berycids, veliferids, the fossil out- groups, and most trachichthyiforms have a low neural arch ("crest") and a single haemal spine on the second preural centrum. Megalomycterids and cetomimids share the feature of two spines on the neural and haemal arches of the second pre ural vertebra. These double spines are not considered pathologic since they are found on all specimens and are always on both arches (for examples see Rosen, 1973, figs. 54-57; Ebeling and Weed, 1973, fig. 5; Fedorov et aI., 1987, fig. 4). The presence of two spines on the neural (as opposed to a low neural crest) and haemal arches of the second preural centrum is diagnostic for the megalomycterids and cetomimids. Posttemporal Long, Unforked, and Splint-like. - In most teleosts, the posttemporal bone is forked with the upper limb contacting the epioccipital and the lower limb contacting the intercalar bone on the neurocranium. This is the condition for most of the basal percomorphs examined, including most trachichthyiforms. In the Megalomycteridae + Cetomimidae the posttemporal is completely free ofthe neurocranium and is reduced to a thin splint, apparently a remnant of the upper limb (Parr, 1929; fig. 10). The intercalar is also absent for these two families. These features are synapomorphies for the families Megalomycteridae and Ce- tomimidae. Loss of Pleural Ribs. -Paxton (1989) listed the loss of pleural ribs as a synapo- morphy for cetomimids, but pleural ribs are also lacking in megalomycterids. This loss is interpreted as a synapomorphy for the Megalomycteridae + Cetomim- idae.

FAMILy-LEVEL SYNAPOMORPHIES The following is a very brief discussion of apomorphies that help define the various families here placed within the Trachichthyiformes. Trachichthyidae. - The trachichthyids possess a posteriorly pointing spine on the posttemporal bone. The only other percomorph examined with a spine on the posttemporal is Centroberyx. The trachichthyid posttemporal spine has a different shape to the spine tip due to the passage of the lateral-line canal. Because of the number of intervening lineages, it is more parsimonious to presume that the two MOORE: TRACHICHTHYIFORM PHYLOGENY 125 posttemporal spines are not homologous and the condition found in trachichthyids is a synapomorphy for the family. Another feature of most trachichthyid species is a modal number of six pro- current spines in the upper caudal-fin lobe and either five or six spines in the lower caudal-fin lobe. The one exception to this is the species Sorosichthys an- anassa which has a modal number of four or five spines in both upper and lower lobes. However, Sorosichthys is a much modified trachichthyid (Moore, msY and probably represents a transformation from the plesiomorphic trachichthyid num- ber. Monocentridae. - The monocentrids exhibit many unique features, including bony armor developed from the scales and the possession of bacterial light organs on the dentary. Monocentrids also have a highly modified pelvic girdle with enlarged and suturally joined pelvic bones and a locking mechanism for keeping the en- larged pelvic spines erect. This girdle and sutural union is structurally very different from that described for the Holocentridae and "higher percomorphs" (Stiassny and Moore, 1992) and is here considered independently derived, most likely to support the erect pelvic spines as a defensive measure. The pelvic rays are reduced in size and number to two or three very small elements. The dorsal fin has a greatly exaggerated heteracanthy of the spines, which alternately point very far to the left and right, also presumably a defensive measure. Anomalopidae. -Johnson and Rosenblatt (1988) thoroughly described the sub- ocular light organ and associated complex of changes to the skull in anomalopids, such as the discontinuous subocular shelf. This seems more than sufficient to demonstrate the monophyly of this family. Diretmidae. -All diretmids lack a . Also, Zehren (1979) described the compound sphenotic of Diretmus as a fusion of the dermosphenotic to the un- derlying autosphenotic, and these fused bones are also found in Diretmoides and Diretmichthys (Kotlyar, 1990b). Diretmus and Diretmoides also share a sutural union of the three most ventral pectoral actinosts to each other, and the ven- tralmost actinost has an interdigitating suture with the adjacent coracoid (Fig. 3). The condition for Diretmichthys is not known. Anoplogastridae. - The anoplogastrids are easily characterized by a number of features including the unusual fangs developed on both the dentary and premaxilla in the adults; very steeply and posteriorly slanted neural spines on the vertebral column; an unusually minute, wedge-shaped urohyal; an interorbital space that is almost completely closed off by bone, as illustrated in Woods and Sonoda (1973); pedicellate-shaped scales (Woods and Sonoda, 1973); in place of a sub- ocular shelf there is a medial thickening of the entire third infraorbital, particularly at the orbital margin; and the second infraorbital is almost completely excluded from the orbital margin with only a small process of that bone extending to the orbit between the lachrymal and third infraorbital. Melamphaidae. - The genus Sio was not examined in this study, so all of the family-level characters listed below have been found in the other four melam phaid genera, Melamphaes, Scopeloberyx, Scopelogadus, and Poromitra. The para- sphenoid is a laterally widened flat bone, quite unlike the narrow shaft with ventrally directed lateral flanges mentioned or illustrated for other trachichthyi- forms and other acanthomorphs by some authors (Patterson, 1967; Gayet, 1980; Stewart, 1984; Johnson and Rosenblatt, 1988) or the simple narrow shaft found in most other fishes. Zehren (1979) described a unique ventrally directed process 126 BULLETIN OF MARINE SCIENCE, VOL. 52, NO.1, 1993

cl

cor· a 4,

Figure 3. Medial view of the right pectoral girdle of Direlmoides pauciradiatus (FMNH 67002). Fin rays have been removed to show the actinosts. Note the interdigitating sutures between actinosts 2- 4 and between aCIinost 4 (a 4) and the coracoid (cor). Other abbreviations: d, deithrum; sc, scapula.

on the fourth, or ventral most, pectoral actinost which is present in all specimens studied here. Zehren (1979) also pointed out that the scapula in melamphaids has two foramina; this condition is found elsewhere only in some genera of the Trach- ichthyidae and is presumably independently derived in that family. Another mod- ification is that a corner of the scapula extends ventrally for some distance; rather than sitting dorsal to the coracoid as in other trachichthyiforms (Zehren, 1979, figs. 26, 38, 46, 55), it is more posterior and even with the ventral edge of the coracoid (Fig. 4). Also, the anterior tips of both the and the coracoid are greatly enlarged. The four genera of melamphaids studied also have the parhypural fused to the first hypural (Zehren, 1979); this was not seen in any other percomorphs examined. The internarial process discussed by Ebeling (1962) is actually in part a unique dorsal projection of mesethmoid. Ebeling and Weed (1973) also mentioned that the lateral line consists of only one or two pores just above the which is quite reduced compared with most trachichthyoids and percomorph outgroups. Stephanoberycidae. -Stephanoberyx and Acanthochaenus share a fusion of in- fraorbitals 3 + 4. The condition is not known in Malacosarcus. Hispidoberycidae. - Hispidoberyx is characterized by the presence of a very long and unusually stout spine on the operculum, as shown in Kotlyar (1980). No MOORE: TRACHICHTHYIFORM PHYLOGENY 127

Figure 4. Lateral view of the pectoral girdle of Scopelogadus unispinis (FMNH 77986) exhibiting the ventrally directed process on the fourth actinost (a 4), two scapular foramina, extension of the ventral comer of the scapula (sc) to a point even with the ventral border of the coracoid (cor), and the greatly expanded anterior tips of both the cleithrum (cl) and coracoid. Scale bar equals 2 mm. other stephanoberycoid has such a prominent spine. Only some trachichthyids have anything approaching the size of the spine in Hispidoberyx. Hispidoberyx also has an extremely spinulose ornamentation to all of the dermal bones of the head (Kotlyar, pers. comm.), far exceeding that of any of the other stephanob- erycoids. Gibberichthyidae. -Gibberichthyids have apomorphic median fin spines in which the more anterior spines are immovably fused to the underlying pterygiophores, as described in de Sylva and Eschmeyer (1977). Gibberichthys has also a fusion of infraorbitals 2 + 3. Rondeletiidae. -Rondeletia has either 4 or 5 supraneurals before the dorsal fin. This is more than the usual 2 or 3 found in most other trachichthyiforms. Barbourisiidae. - Barbourisiids are very unusual in having 8 supraneurals anterior to the dorsal fin. This is more than any other acanthomorph examined. Barbourisia also has 7 infraorbitals, not counting the dermosphenotic. This is two more than is found in most other percomorph fishes. The only other taxon seen with as many infraorbitals is Cetostoma (Parr, 1929, fig. 10). The velvet-like spinulose skin of Barbourisia results from a very dense covering of tiny thumbtack-shaped scales extending over almost the entire head and body. Both the shape of the scales and their distribution appear to be particular to this genus and family. Megalomycteridae. - Myers and Freihofer (1966) described the unusually large nasal rosettes found in the megalomycterids. Megalomycterids and other mira- pinnatoids (i.e., Mirapinnidae and Eutaeniophoridae) have greatly reduced pha- ryngeal bones and dentition (Bertelsen and Marshall, 1956; Rosen, 1973). Atax- olepis almost completely lacks buccal or pharyngeal dentition or gill rakers with 128 BULLETIN OF MARINE SCIENCE, VOL. 52, NO. I, 1993 only vomerine teeth reported in a paratype of A. apus (Myers and Freihofer, 1966) and a few gill rakers on the ceratobranchials of A. henactis (Goodyear, 1970). Great reduction or complete loss of buccal and pharyngeal dentition may be a familial trait. Cetomimidae. - The apomorphic copular teeth over the basibranchials of cetom- imids are described more completely and illustrated for all genera in Paxton (1989). Another autapomorphous set of character states described by Paxton (1989) is the variously modi fed gill rakers found in all cetomimids. These mod- ifications range from individual teeth on the gill arches to club shaped gill rakers to toothy knobs to flat tooth plates, but taken as a whole these various modifi- cations are very different from the typical gill rakers found in Barbourisia, Ron- de/etfa, or more primitive stephanoberycoids. Finally, cetomimids have scales restricted to the anal fin lappets and the lateral line canals or, in the case of the two genera lacking a lateral line canal, the scales are in the skin underneath the papillate lateral line (Paxton, 1989). This particular distribution of scales is dis- tinctive for the cetomimids.

DISCUSSION One most parsimonious tree (length = 31, CI = 0.935, RI = 0.976) was com- puted by the PAUP heuristic search method (Fig. 5). From this tree it can be seen that the order Trachichthyiformes is composed of two subordinal clades, the Trachichthyoidei (containing the families Anolpogastridae, Diretmidae, Anom- alopidae, Monocentridae and Trachichthyidae) and the Stephanoberycoidei (con- taining the Melamphaidae, Stephanoberycidae, Hispidoberycidae, Gibberichthyi- dae, Rondeletiidae, Barbourisiidae, Megalomycteridae, Cetomimidae and probably the mirapinnid fishes also). The cetomimoids are a derived, monophyletic subset of the stephanoberycoids. In the past, the placement of the cetomimoids has been problematic. Originally they were placed in the Iniomi by Goode and Bean (1895) and maintained there by many other authors. Parr (1929) questioned this placement after studying the osteology of the Rondeletia and Cetostoma ("Cetomimus" regani). Although Parr retained the cetomimids within the Iniomi, he stated that Rondeletia is a "typical representative of the order Xenoberyces introduced by Regan 1911." This led some authors to state that the cetomimoids are a composite or polyphyletic group (Gosline, 197I). Other authors (Harry, 1952; Rofen, 1959) and this study have found that the cetomimoids are indeed a valid group and a subgroup of the stephanoberycoids (essentially Regan's Xenoberyces) with the family Cetomim- idae so highly transformed and reduced as to make them similar in some ways to various iniomous or pre-acanthomorph fishes (Rosen and Patterson, 1969; Rosen, 1973). Evidence that the cetomimoids do belong among the Acanthomorpha and within the Trachichthyiformes includes: Rondeletia and Barbourisia with a spina occipitalis, an acanthomorph synapomorphy (Stiassny, 1986). All cetomimoids lack urodermal and fulcral scales, the absence of which may also be acanthomorph features (Zehren, 1979). Cetomimoids usually have 8 or occasionally 9 branchio- stegals, a general reduction from the 9 or more found in most pre-acanthomorphs (Hubbs, 1920), but consistent with the number found in all of the trachichthyi- forms. Rondeletia and Barbourisia have a single row of intermuscular bones (usually called "epipleural ribs") extending from the first two centra and the region of the pleural rib heads; the epineural ribs are absent, which is the condition found in MOORE: TRACHICHTHYIFORM PHYLOGENY 129

GI III II II 'tJ II III CII III III 1lI Gl U 'tJ Gl Gl 'tJ 'tJ Gl -.::: 'tJ III >- Gl III 'tJ 1lI III U >. III Gl Gl >. -.::: 'tJ 'tJ >- CD ~ :E III ~ .D ;g U 'tJ a ~ CD 0 E >- E C 0 ~ .D 0 Gl D. C .!:! Gi ~ E E 0 iii 0 aI ;:, 0 'tJ Gi :c 0 E ~ CD 'tJ 0 iii E 0 E aI D. .D .D 0 aI C 0 a c til 0 C 'iii 1Il Gl CII .. :8 0 Ii Gl t= ::Ii 0( x (;j " a: III ::Ii 0

Figure S. One most parsimonious c1adogram resulting from the data matrix in Table 1. The characters are as numbered in the list of characters. The numbers in parentheses are the character state for multistate characters. the Paracanthopterygii + Atherinomorpha + Percomorpha and potentially this loss of the epineural ribs is a synapomorphy for that group (Johnson and Patterson, 1993; Moore, 1993). Rondeletia bieolor and Barbourisia rufa have high cranial ridges on the frontals with a large opening over the orbit and large mucus-filled cavities similar to those found in stephanoberycoids, trachichthyoids, berycids, and other basal perco- morphs. Rondeletia and Barbourisia have the teeth expanded somewhat antero-laterally at the dentary symphysis; a feature which unites two of the fossil outgroups with the trachichthyiforms. All the cetomimoids have the neural arch of the first ver- tebra fused to the underlying centrum and lack ocular sclera, two trachichthyiform characters. All of these characters indicate that the proper placement of the ce- tomimoids is within the trachichthyiforms, despite the various reductions and transformations which have confounded previous authors. In reviewing the literature regarding the Mirapinnidae (sensu Bertelsen and Marshall, 1984), no features are evident that would place these fishes anywhere other than as a sister group to the Megalomycteridae, and that clade as a sister group to the Cetomimidae, as suggested by Bertelsen and Marshall (1984). These authors also provide one feature, the particular distribution of red muscle on the body, to unite the cetomimids, megalomycterids, and mirapinnids. Paxton (1989) has given evidence for ruling out the mirapinnid and megalomycterid fishes as juveniles or sexual dimorphs of the cetomimids, as has been suggested by some authors (Gosline, 1971; Robins, 1974; de Sylva and Eschmeyer, 1977). Although no material from the Mirapinnidae (sensu Bertelsen and Marshall, 1984) has been examined in this study, it seems most likely from the literature that the Mira- 130 BULLETIN OF MARINE SCIENCE, VOL. 52, NO. I, 1993 pinnidae should be placed with the Megalomycteridae among the stephanobery- coid Trachichthyiformes.

ACKNOWLEDGMENTS

This paper is derived from some of the conclusions contained in my dissertation. I could not thank enough everyone who has helped me with that project, but I would like to make mention of a few individuals who have greatly facilitated this paper. B. Chernoff, N. Feinberg, K. Hartel, S. Jewett, D. Johnson. M. A. Rogers, M. Stiassny, and J. Williams have all kindly given access to the collections in their care. A. Seilacher has given valuable advice concerning the figures. W. Anderson Jr., C. Baldwin, T. Gill, D. Johnson, R. Mooi, C. Patterson, and M. Stiassny have provided numerous comments on this manuscript. Support for this research has come in part from a John Enders Fellowship and a grant from the Women's Seaman's Friend Society of Connecticut.

LITERATURE CITED

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DATEACCEPTED: June 9, 1992. ADDRESS: Division of Vertebrate Zoology, Peabody Museum of Natural History. Yale University, New Haven, Connecticut 0651 I. APPENDIX

Specimens are listed alphabetically by family. Cleared and stained, alcohol, x-ray specimens and dried skeletons are noted after the catalogue number as (c and s), (ale), (x-ray) or (skel), respectively. Uncatalogued specimens are noted as "uncat."

Acropomatidae Acropoma sp. USNM 287444 (c and s) wakiyae CAS-SU 23815 (c and s) Anomalopidae Anomalops katoptron CAS 55163 (e and s), AMNH 37949 (c and s), USNM uneat. (c and s), MCZ 56858 (alc) Kryptophaneron alfredi ANSP 136511 (c and s), ANSP 144633 (ale) Photoblepharon palpebratus SIO 74-58 (c and s), MCZ 56859 (ale) Photoblepharon steinitzi FMNH 95100 (c and s) Anoplogastridae Anoplogaster cornu/us MCZ 71611 (c and s), MCZ uncat. (e and s), YPM 2843 (c and s), FMNH 66620 (c and s), YPM 2845 (ale) Anoplogaster brachycera MCZ uncat. (c and s) Aphredoderidae Aphredoderus sayanus AMNH 27432 (c and s) Atherinidae Menidia menidia YPM uncat. (c and s) Telmatherina ladigesi YPM uncat. (c and s) Aulopodidae Aulopusjaponicus FMNH 65005 (c and s), AMNH 28635 (c and s) and 13024 (c and s), FMNH 89115 (ale) MOORE: TRACHICHTHYIFORM PHYLOGENY 133

Appendix. Continued

Barbourisiidae Barbourisia rufa AMS 1.18824-001 (e and s), YPM 1217 (holotype) (ale), USNM 197858 (ale) Bedotiidae Bedolia geayi AMNH 57452 (e and s) Rheoc/es alaolrensis AMNH 8800 I (e and s) and 88171 (e and s) Beryeidae Beryx dedaclylus MCZ 59527 (ale), FMNH 47972 (ale), USNM uneat. (x-ray) Beryx splendens FMNH 66446 (e and s), USNM 290352 (ale) Centroberyx affinis USNM 176984 (e and s), USNM 296453 (ale and x-ray) Anligonia capros USNM 289209 (e and s) Anligonia combalia USNM 188046 (e and s), USNM 188046 (x-ray) Anligonia rubescens USNM 182841 (x-ray) USNM 289207 (e and s), USNM uneat. (x-ray) Centrarehidae Microplerus salmoides YPM 1748 (skel) Centropomidae Cenlropomus unidecimalis YPM 8911 (e and s), USNM uneat. (x-ray) Cetomimidae Celomimus sp. YPM 8151 (ale and x-ray) CelOmimus kerdops YPM 3204 (holotype) (ale and x-ray) CeloslOma regani YPM 2132 (e and s), USNM 200531 (ale) Chlorophthalmidae Chlorophthalmus agassizi FMNH 67115 (e and s) oClodecimspinosus YPM 1617 (skel) Oiretmidae Direlmoides sp. YPM 9041 (ale), MCZ 70168 (ale) Direlmoides pauciradiatus FMNH 67002 (c and s), USNM 93454 (ale) Direlmus argenleus AMNH 49713 (e and s), MCZ 64749 (e and s), USNM 240760 (e and s), FMNH 47889 (e and s), YPM 2799 (ale) Gibberiehthyidae Gibberichlhys pumilus MCZ 44211 (e and s), FMNH 66805 (e and s), YPM 2838 (holotype) (ale), YPM 3232 (ale), USNM 205551 (ale), MCZ 44211 (ale) Hispidoberyeidae Hispidoberyx ambagiosus MMSU P.15416 (ho]otype) (x-ray), MMSU uneat. (x-ray) Holoeentridae Corniger spinosus ANSP 144596 (ale), USNM 83179 (ale and x-ray) Holocen/rus ascensionis AMNH 30240 (e and s), YPM 2320 (e and s), YPM 1722 (skel), USNM 109986 (skel) Holocenlrus rufus AMNH 27] 18 (e and s), JAM 87-1 (e and s), pers. coil. (ale) Myriprislis clarionensis USNM 67575 (e and s) Myriprislis jacobus AMNH 23380 (e and s), USNM 12686 (skel) Myriprislis murdjan USNM 207780 (e and s), USNM I] 1442 (skel) Myriprislis pralinius USNM 288353 (e and s) Neoniphon argenleus USNM 259911 (ale) Neoniphon opercularis USNM 154526 (e and s) Neoniphon sam mara USNM 218907 (e and s), pers. coil. (ale) OSlichlhys archiepiscopus USNM 204034 (x-ray) Oslichlhys della USNM 223716 (paratype) (ale and x-ray) OSlichlhys Irachypoma FMNH 65198 (e and s), USNM ]57990 (e and s), USNM 265723 (skel), 134 BULLETIN OF MARINE SCIENCE, YOLo 52, NO. I, 1993

Appendix. Continued

AMNH 34899 (ale) Plectrypops lima USNM 224721 (c and s), USNM 257406 (c and s), USNM 140895 (ale) Plectrypops retrospinus AMNH 34106 (c and s), USNM 79899 (alc and x-ray) Pristilepis oligolepis ANSP 86824 (alc), USNM 52746 (ale and x-ray) Sargocentron diadem a USNM 287735 (c and s) Sargocentron lepros USNM 233061 (paratype) (ale and x-ray) Sargocentron rubrum USNM 259103 (c and s) Sargocentron spiniferum MCZ uncal. (skel) Sargocentron suborbitalis FMNH 73918 (c and s) Sargocentron xantherythrus USNM 289208 (c and s) Lampridae Lampris guttatus AMNH 79642 (ske1) Macrurocyttidae hololepis AMNH 29463 (c and s) Megalomycteridae Ataxolepis apus USNM 221036 (e and s and ale), FMNH 71705 (ale) Melamphaidae Melamphaes eulepis FMNH 77982 (c and s) Melamphaes macrocephalus FMNH 77983 (c and s) Melamphaes microps YPM 8470 (c and s), SIO 54-100 (c and s) Melamphaes suborbita/is MCZ 52812 (ale) Poromitra sp. AMNH 28937 (c and s), YPM uncat. (c and s) Poromitra capito MCZ 76561 (c and s), YPM 8167 (ale) Poromitra crassiceps FMNH 86300 (c and s), YPM 2741 (ale) Poromitra cristiceps SIO 54-83 (c and s) Poromitra megalops YPM uncat. (c and s) Scopeloberyx sp. FMNH 77984 (c and s) Scopeloberyx opisthopterus MCZ 59920 (c and s), FMNH uncal. (c and s), YPM uncal. (c and s), YPM 2824 (ale) Scopelogadus beanii MCZ 53851 (ale) Scopelogadus mizolepis mizolepis MCZ 60850 (c and s), FMNH 77985 (c and s) Scopelogadus unispinis FMNH 77986 (c and s) Melanotaeniidae Melanotaenia sp. YPM uncal. (c and s) Melanotaenia nigrans AMNH 55067 (c and s) Monocentridae Cleidopus gloriamaris AMNH 38197 (c and s), MCZ 38581 (ale) Monocentrisjaponicus AMNH 27412 (c and s), FMNH 73560 (c and s), MCZ 25758 (ale), MCZ 47731 (ale) americanus YPM 5342 (c and s) Myctophidae Diaphus dumerili YPM 424 (c and s) Lampanyctus lineatus YPM 2298 (c and s) Osmeridae Osmerus mordax YPM uncat. (c and s) Ostracoberycidae dorygenys USNM 289214 (c and s), USNM 168442 (x-ray) Percopsidae Percopsis omiscomaycus YPM 8628 (c and s), FMNH 47972 (c and s) polymixiidae Polymixia lowei USNM 185284 (c and s), YPM 9054 (ale) Polymixia nobilis YPM 7548 (ale), USNM uncat. (x-ray) MOORE: TRACHICHTHYIFORM PHYLOGENY 135

Appendix. Continued

Rondeletiidae Rondeletia bieolor YPM 2103 (c and s), YPM 2105 (c and s), YPM 2108 (ale), USNM 203821 (ale) Rondeletia lorieata MCZ 43330 (c and s), USNM 206834 (ale) Dendroehirus zebra YPM uncal. (c and s) Eetreposebastes imus AMNH 27991 (c and s) Helieolenus daetylopterus USNM uncal. (x-ray) grandieornis YPM 257 (c and s) Scorpaena dispar USNM un cat. (x-ray) rosaceus USNM 106886 (x-ray) EpinephelusJulvus YPM 1695 (skel) Stephanoberycidae Acanthochaenus lutkeni MCZ 64948 (ale) Stephanoberyx monae AMNH 49679 (c and s), MCZ 41640 (c and s), USNM 208284 (c and s), FMNH 77987 (c and s), MCZ 41640 (ale), YPM 4250 (ale) Stomiidae Malacosteus niger YPM 2060 (c and s) Photoneetes intermedius YPM 2100 (c and s) Photostomias guernei YPM 2056 (c and s) Synodontidae Traehinoeephalus myops YPM 63 (c and s) Trachichthyidae Aulotraehichthys prosthemius USNM 179739 (ale) Aulotraehichthys sajademalensis FMNH 86460 (c and s and ale) Gephyroberyx darwini FMNH 64532 (c and s), 65227 (c and s), USNM 214210 (ale) Gephyroberyx philippinus AMNH 4970 I (c and s) Hoplostethus mediterraneus AMNH 26763 (c and s), FMNH 66788 (c and s), MCZ 70173 (c and s), MCZ 40338 (ale), USNM uncal. (x-ray) Hoplostethus meta/licus USNM 93344 (holotype), USNM 9339 (ale) (paratype) Hoplostethus occidentalis USNM 214196 (c and s) Optivus elongallls YPM 9869 (c and s), USNM 296454 (alc), YPM 9335 (ale) Paratrachichthys Jernandezianus FMNH 74351 (c and s) Paratrachichthys trailli YPM 9870 (c and s), YPM 9336 (ale), pers. coil. (ale) Sorosiehthys ananassa USNM uncal. (c and s) Trachichthys australus USNM 197649 (c and s and ale) Triacanthodidae Hollardia hollardia USNM 280328 (c and s) Triacanthodes anomalus USNM 93487 (c and s) Veliferidae Velifer aJricanus USNM 235676 (ale), USNM uncat. (x-ray) roseus YPM 385 (ale), USNM 186029 (x-ray) eonchifer USNM 303498 (c and s and ale) Jaber USNM 303497 (c and s and ale), USNM uncal. (x-ray) FOSSIL SPECIMENS Apipchthyidae Aipichthys minor FMNH PF 13392, FMNH PF 13396 Aipichthys vellfer FMNH PF 10440, FMNH PF 13794 Ctenothrissidae Ctenothrissa vexillifer ANSP (VP) 5801, ANSP (VP) 14523, FMNH PF 13377 136 BULLETIN OF MARINE SCIENCE, VOL. 52. NO. I. 1993

Appendix. Continued

Pycnosteroididae Pycnosteroides /evispinosus FMNH PF 10439 Percomorpha incerta sedis Acrosgaster ?heckeli ANSP (VP) 15486 Hop/apteryx sp. ANSP (VP) 14606, ANSP (VP) 14761, YPM-PU 11989, YPM-PU 11991, YPM-PU 11992, YPM-PU 12224 Hop/opteryx /ewesiensis ANSP (VP) 6042, ANSP (VP) 6045 Hop/opteryx simus ANSP (VP) 14762 Lissoberyx dayi FMNH PF 13523