BUI I 1 I IN 01 miBIOLOGICAL SOCIETY
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Editor: Richard v. SternbergReview Editor: Lynne R. ParentiReviewers: Gareth Nelson and Richard Winterbottom
Copies available as the supply lasts from:The Custodian of PublicationsBiological Society of WashingtonNational Museum of Natural HistorySmithsonian InstitutionWashington, D.C. 20560(Cost $75.00 for the 2-volume set, including postage and handling)
Cover illustration: Pholidichthys leucotaenia, dorsal gill-arch musculature(see Plate 169).
"This whole book is but a draught?nay, but the draft of a draft. "Herman Melville, Mobv Dick.
STUDY OF THE DORSAL GILL-ARCH MUSCULATUREOF TELEOSTOME FISHES, WITH SPECIALREFERENCE TO THE ACTINOPTERYGII
Victor G. Springer and G. David JohnsonKarolyn Darrow, Illustrator
Appendix: Phylogenetic Analysis of 147 Families of Acanthomorph FishesBased Primarily on Dorsal Gill-Arch Muscles and Skeleton
Victor G. Springer and Thomas M. Orrell
(VGS, GDJ, TMO), Division of Fishes MRC 159, Department of ZoologyNational Museum of Natural History, P.O. Box 37012Washington, D.C. 20013-7012, U.S.A.e-mail: Springer.Victor@nmnh.si.edu(KD) Department of Entomology MRC 105National Museum of Natural History, P.O. Box 37012Washington, D.C. 20013-7012, U.S.A.e-mail: Darrow.Karolyn@nmnh.si.edu
Abstract.?The dorsal gill-arch musculature (DGM), aspects of the asso-ciated skeleton, the transversus ventralis 4, and the semicircular ligament aredescribed for many species in over 200 families and over 300 genera ofteleostome fishes, and the DGM musculature of over 200 taxa is illustrated.A partially new system of DGM nomenclature is used. The transversus dor-
salis, is shown to be a much more complex system than has been generallyrecognized. DGM data are variously analyzed and shown to be of importancefor defining various currently recognized suprageneric pre-acanthomorph taxa.Among the many conclusions pertaining to acanthomorphs are: monophylyof Percopsiformes is hypothesized; Icosteidae, Menidae, and Centriscidae areprobably more closely related to pre-percomorph groups than to percomorphgroups; there is no basis for inclusion of Amarsipidae in the Stromateoidei;monophyly of the Polycentridae (Polycentriis, Polycentropsis, Afronandus,Monocirrhus) is corroborated based on a combination of gill-arch muscle andadditional characters. A new ordinal-group name, Anabantomorpha, is erectedto include the seven families with parasphenoid teeth: Nandidae, Badidae,Pristolepidae, Channidae, Anabantidae, Heleostomatidae, and Osphronemi-dae. Anabantoidei includes the last four of these families, which have supra-branchial organs. The superfamily name Labroidea is proposed to distinguishthe unequivocally monophyletic group of families, Labridae, Odacidae, Scar-idae, from other families included in the suborder Labroidei. The first syna-pomorphy for the gobioid family Odontobutidae is hypothesized based on theposition of the levator interims 2 relative to the obliquus dorsalis. Certaingill-arch skeletal characters previously unrecognized or inadequately evalu-ated are reported and discussed (e.g., epibranchial-ceratobranchial accessorycartilages; relationship of epibranchials 5 and 4; epibranchial 4 flange; esoph-ageal raphe).A separately authored appendix provides a cladistic analysis of 168 taxa in147 acanthomorph families based almost exclusively on DGM and gill-archskeletal characters. Among the many results implied by the study are: mono-phyly of Percopsiformes is corroborated. Smegmamorpha Johnson and Pat-terson (1993) are polyphyletic, and the name is rejected for nomenclaturalpurposes. Its constituents comprise two or three not closely related clades:Mugilomorpha + Atherinomorpha (= Percesoces Cope, 1875), Gasterosteo-morpha, and. possibly, Centrisciformes (a pre-percomorph group, comprisingonly Centriscidae). Gasterosteomorpha includes a monophyletic Hypoptych-idae {Hypoptychus + Aulichthys), Elassomatidae, Aulorhynchidae, monophy-letic Synbranchiformes (Synbranchidae + Mastacembelidae), and Gasteros-teidae, thus corroborating various hypotheses of Johnson and Patterson (1993)and Johnson and Springer (1997). Labroids are monophyletic only with in-clusion of Pholidichthyidae, but the group remains supported only by phar-yngognath characters. Blennioidei are monophyletic and their intra-relation-ships resolved. Their closest relatives are, stepwise: Gobiesocidae, Draconet-tidae (+ Callionymidae, which were not included in the analysis), Dactylop-teridae. A new ordinal-group name, Benfhomorpha, is proposed for this clade.Caproidae are polyphyletic; relationships of its two genera appear to be withtetraodontiforms on the one hand, and acanthuroids, on the other. As such,Johnson and Patterson's (1993) hypothesis that caproid relationships areamong percomorphs is corroborated. Sphyraenidae and Polynemidae form amonophyletic group (first proposed by Regan, 1912).
CONTENTS
First Author's Preface 1Introduction 2Methods 2Material 3Muscles and Skeletal Elements 3Muscle names 3Muscle types 4Origins and insertions 4Anatomical orientation 4Transverse muscles 4Acanthomorph accessory cartilages 4Epibranchials 5 and 4 5Abbreviations and Definitions for AnatomicalStructures 6Classification 16Gnathostomata 18Teleostomi 18Sarcopterygii 18CoelacanthomorphaCoelacanthidae 18Dipnoi 19Ceratodontidae 19Actinopterygii 21Pre-Acanthomorpha 21Cladistia 21Polypteridae 21Chondrostei 22Acipenseridae 22Polyodontidae 23Ginglymodi 24Lepisosteidae 24Halecomorphi 24Amiidae 24Osteoglossomorpha 25Hiodontidae 25Notopteridae 26Gymnarchidae 27Mormyridae 27Arapaimidae 28Pantodontidae 30Osteoglossidae 31Elopomorpha 32Megalopidae 32Elopidae 33Albulidae 34Notacanthidae 36Halosauridae 37Congridae 37Anguillidae 38Synaphobranchidae 39Clupeomorpha 39Denticipitidae 39Pristigasteridae 40Engraulidae 41Chirocentridae 42
Clupeidae 43Gonorynchiformes 44Chanidae 44Gonorynchidae 45Cypriniformes 46Cyprinidae 46Characiformes 48Characidae 48Distichodontidae 49Siluriformes 49Diplomystidae 49Gymnotiformes 51Gymnotidae 51Salmoniformes 51Salmonidae 51Retropinnidae 53Galaxiidae 54Osmeridae 56Argentiniformes 57Argentinidae 57Alepocephalidae 57Platytroctidae 58Esociformes 59Esocidae 59Umbridae 60Stomiiformes 62Diplophidae 62Sternoptychidae 63Gonostomatidae 63Ateleopodiformes 64Ateleopodidae 64Aulopiformes 65Aulopidae 65Synodontidae 66Chlorophthalmidae 66Myctophiformes 67Neoscopelidae 67Myctophidae 68Results?Pre-Acanthomorpha .... 69Tables 1-7 75-81Acanfhomorpha 74Additional material 74Lampridiformes 80Veliferidae 80Lampridae 81Polymixiiformes 82Polymixiidae 82Paracanthopterygii 83Percopsiformes 83Aphredoderidae 84Percopsidae 84Amblyopsidae 85Ophidiiformes 86Ophidiidae 86Bythitidae 88
Carapidae 88Gadiformes 89Ranicipitidae 89Batrachoidiformes 90Batrachoididae 90Lophiiformes 90Chaunacidae 90Acanthopterygii 91Stephanoberyciformes 91Melamphaidae 91Gibberichthyidae 92Stephanoberycidae 92Barbourisiidae 93Rondeletiidae 93Cetomimidae 94Icosteiformes 94Icosteidae 94Tables 8 & 9 98-107Zeiformes and Possible Relatives 107Zeiformes 108Oreosomatidae 108Parazenidae 108Zeniontidae 108Grammicolepidae 109Caproiformes 110Caproidae 110Tetraodontiformes IllTriacanthodidae IllMeniformes 112Menidae 112Beryciformes 113Trachichthyidae 113Berycidae 1 14Holocentridae 115Anomalopidae 116Percomorpha 117Smegmamorpha 117Gasterosteomorpha 117Centrisciformes 117Centriscidae 117Gasterosteiformes 118Gasterosteidae 118Hypoptychidae 120Aulorhynchidae 121Synbranchiformes 122Synbranchidae 122Mastacembelidae 123Elassomatiformes 124Elassomatidae 124Mugilomorpha 124Mugilidae 124Atherinomorpha 126Atheriniformes 126Atherinidae 126Bedotiidae 126Cyprinodontifomes 127
Aplocheilidae 127Cyprinodontidae 127Beloniformes 128Adrianichthyidae 128Belonidae 129Scomberesocidae 130Hemiramphidae 131Exocoetidae 131Perciformes 133Acropomatidae 133Percichthyidae 133Leptobramidae 134Latidae 135Centropomidae 135Centrarchidae 136Bathyclupeidae 137Symphysanodontidae .... 138Epigonidae 139Moronidae 140Serranidae 140Lutjanidae 141Haemulidae 142Inermiidae 142Apogonidae 143Priacanthidae 143Ostracoberycidae 144Cirrhitidae 144Pempheridae 145Glaucosomatidae 145Lactariidae 146Lateolabracidae 147Sciaenidae 147Polynemidae 148Sillaginidae 149Mullidae 150Centrogeniidae 151Ambassidae 151Caristiidae 152Bramidae 153Toxotidae 154Plesiopidae 155Percidae 156Cepolidae 157Callanthiidae 158Gerreidae 159Grammatidae 160Opistognathidae 160Pseudochromidae 161Leiognathidae 162Polycentridae 163Sphyraenidae 166Kurtidae 167Ammodytidae 168Trachinidae 168Uranoscopidae 169Cheimarrichthyidae 170Scorpaenoidei 170
Scorpaenidae 171Sebastidae 171Platycephalidae 172Champsodontidae 172Hexagrammidae 173Anoplopomatidae 175Rhamphocottidae 175Cottidae 176Normanichthyidae 177Carangoidei 177Nematistiidae 177Carangidae 178Rachycentridae 178Coryphaenidae 179Echeneidae 180Scombroidei 181Pomatomidae 181Scombrolabracidae 182Scombridae 182Sparoidei 183Nemipteridae 183Lethrinidae 184Centracanthidae 185Sparidae 185Girelloidei 187Girellidae 187Kuhliidae 188Terapontidae 188Labroidei 189Cichlidae 189Pomacentridae 191Embiotocidae 193Labroidea 194Labridae 196Pholidichthyoidei 199Pholidichthyidae 200Acanthuroidei 201Luvaridae 201Ephippidae 202Zanclidae 202Acanthuridae 203Anabantomorpha 203Nandidae 204Badidae 204Pristolepidae 205
Channidae 206Anabantidae 207Stromateoidei 207Amarsipidae 208Centrolophidae 209Zoarcoidei 209Bathymasteridae 209Zaproridae 210Pholidae 211Stichaeidae 211Notothenioidei 211Bovichtidae 211Pseudaphritidae 212Dactylopteroidei 213Dactylopteridae 213Malacanthidae 214Callionymoidei 215Draconettidae 215Callionymidae 216Gobiesocidae 218Blennioidei 218Tripterygiidae 219Blenniidae 220Dactyloscopidae 221Clinidae (Myxodinae) . . . 221Clinidae (Clininae) 222Labrisomidae 223Chaenopsidae 224Gobioidei 224Rhyacichthyidae 225Odontobutidae 226Xenisthmidae 228Eleotridae 228Microdesmidae 229Gobiidae 230Pleuronectiformes 233Psettodidae 233Tables 10 & 11 194, 225Acknowledgments 235Appendix: Phylogenetic Analysis of 147Families of Acanthomorph Fishes BasedPrimarily on Gill-arch Muscles andSkeleton 237Literature Cited 255Plates (separate volume)
First Author's Preface
This study was initiated about 1996 as the primaryeffort of the first author (VGS), who had long beeninterested in determining the interrelationships of theacanthomorph family Pholidichthyidae. Springer andFreihofer (1976) last discussed the problematic inter-relationships of the then monospecific family(Springer and Larson, 1996, described a second Phol-idichthys species). Subsequently, Stiassny and Jensen(1987), expanding on the work of Kaufman and Liem(1982), hypothesized the composition and interrela-tionships of an acanthomorph suborder Labroidei.Stiassny and Jensen based their hypothesis on a rel-atively broad selection of acanthomorph fishes andalmost exclusively on a limited number of gill-archcharacters. Referring only to Springer and Freihofer's(1976) study, they noted similarities of the gill-archskeleton of Pholidichthys to that of the labroids.Johnson (1993:9-10) discussed errors of oversightand commission in Stiassny and Jensen's (1987)study and noted, critically, that, other than charactersassociated with pharyngognathy, there was none thatcorroborated monophyly of the labroids. VGS re-ex-amined Pholidichthys in light of Stiassny and Jen-sen's and Johnson's studies, and noted problems withboth, although he agreed with Johnson's general crit-icism. As a result, VGS, with the assistance of GDJ,undertook to survey a wide variety of acanthomorphfishes to determine the distribution of the states ofthe muscle characters used by Stiassny and Jensen.VGS expanded the study to include a broad spectrumof non-acanthomorph fishes because of problems indetermining muscle homologies among the acantho-morphs.Darrow joined the project as full-time illustrator inthe fall of 1997 and remained on the project untilmid-2000, after which she continued on contract andthen as volunteer until late 2003, when all the gill-arch muscle illustrations were completed.VGS contracted with Tom Orrell to run PAUP pro-gram analyses of a limited number of acanthomorphtaxa beginning in 2002, but in 2003 involved him inthe preparation of the major cladistic analysis form-ing our co-authored Appendix to the present study.After essentially completing the actinopterygianpre-acanthomorph portion of the study, including an
analysis of the data, and much of the descriptive por-tion of the acanthomorphs, joint efforts with GDJceased in early 2002 at the sole instigation of VGS.VGS is responsible for selecting most of the taxaused in the study, preparing a large majority of thedissections, all the descriptions, supervision of prep-aration of all the illustrations, the partially new sys-tem of nomenclature used to designate the muscles,the interpretation and results of the non-acantho-morph portion of the study, most of the acantho-morph interrelationships, discussions that are not out-growths of the cladistic analyses, all of the choicesof taxa, characters, and character codes for the cla-distic analyses, and preparation of all preliminary andfinal drafts of the entire manuscript. /, VGS, there-fore, accept full responsibility for all errors, factualor otherwise, and eccentricities that this study em-bodies. On the other hand, I gladly share with myco-authors responsibility for any favorable aspects ofthe study. I especially want to thank them for theirinput: GDJ for his early encouragement and impor-tant suggestions for taxa to include, ready and com-prehensive knowledge of the existing classificationsof many groups of fishes and their defining charac-ters, and his often constructive challenges, his criticalreading of an early complete draft of the pre-acan-thomorph section of the actinopterygian portion ofthe study, early drafts of a large number of the acan-thomorph descriptions and discussions, commentingon a near final draft of the pre-Appendix portion ofthe manuscript, and very importantly, for bringingKarie Darrow to my attention; Karie Darrow for herdedication to the project, even after monetary com-pensation ceased, and the unstinting use of her greatillustrative talents, as well as for the numerous oc-casions on which she caught my descriptive mistakes;and to Tom Orrell for his insightful and knowledge-able handling of the PAUP program used in the phy-logenetic analysis, important suggestions for thepreparation and interpretation of the output, and pa-tience under stress of my importunities.In the pre-Appendix portion of the text that fol-lows, the editorial "we" and "our" are used to ac-cord with authorship, but their use is not intended toimply agreement by the second author.
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IntroductionThe present study has two main purposes. First, toprovide an annotated, descriptive atlas of the dorsalgill-arch muscles of the extant fishes (coelacanths,lungfishes, actinopterygians) included in the Super-class Teleostomi (= Grade Teleostomi, in part, ofNelson, 1994:65) or Osteichthyes (= branch 8 of Gilland Mooi, 2002:fig. 2.2) with special reference to theActinopterygii. Second, to determine how the char-acter states exhibited by these muscles accord withexisting morphologically based phylogenetic classi-fications of the fishes, and to a much lesser extent,molecular-based classifications.Because our study initially involved two ventralgill-arch structures (notably, the semicircular liga-ment and the transversus ventralis 4), we also sur-veyed them. We variously include treatment of otherventral gill-arch muscles and muscles that span boththe dorsal and ventral gill-arch elements. On the otherhand, as a result of the way the study developed, wedid not survey the protractor pectoralis, which is of-ten closely associated with the dorsal gill-arches. Weregret this omission, but the muscle was destroyedduring preparation of many of the gill-arches beforewe realized its potential importance (for an extensivesurvey of this muscle see Greenwood and Lauder1981).In our attempts to provide accurate illustrations ofthe muscles and their attachments, we also attemptedto apply the same attention to the dorsal gill-archskeletal elements. We frequently illustrate and dis-cuss skeletal structures that were previously unre-ported or erroneously described. On the other hand,we do not address all the skeletal characters that havebeen used in previous classifications, only those thatare obvious in the illustrations, have direct bearingon the position of muscle attachments, or that bychance came to our attention (e.g., whether infra-pharyngobranchial 2 is present or absent, and if pre-sent bears teeth or is edentate).In a study as broad as ours, keeping up with rel-evant literature was a problem. Although, we at-tempted to do this, we recognize the possibility thatreferences will have been missed. Another problemis that of references that became available after ourdiscussion and analyses were in an advanced stateand to adjust for them would have required continualrevision and delay of the study. As far as we areaware, we have included and adjusted for the litera-ture published through the end of 2002, and much ofthe literature of 2003.
MethodsDissections were made and studied primarily usinga Zeiss Operation Technoscope. Drawings were madeusing a Wild M-3 stereoscopic microscope with at-
tached camera lucida. As required, details werechecked using a high resolution Leitz stereoscopicdissecting microscope.In general, gill arches were removed from speci-mens using the following procedure. First, the hyoidarches including the branchiostegals were either sep-arated from the membranes connecting them to theopercular series and the interhyal linking them to thehyomandibula and retained attached to the gill-arch-
es, or, usually, the gill arches were released from thehyoid arches by separating the hypohyals from thebasibranchials. The basihyal was then freed by mak-ing a semicircular incision around the floor of themouth. Next, the gill filaments were stripped offwhile trying to assure that the external and posteriorlevators were not removed with them (levator exter-nus 1 and levator posterior, require the most care toavoid damage or removal). Tissue enclosing the gill-arch musculature laterally was then removed. Partic-ular care was necessary at this point as the levatorexternus 4, levator posterior, and protractor pecto-ralis in some forms (e.g., clupeomorphs, lampridi-forms) may be closely applied or imbedded in thetissue and would be damaged or removed with thetissue. A cut was then made across the pharyngealroof just anterior to the gill arches and pharyngo-branchial 1, if present, and the latter was releasedfrom its attachment to the skull. Muscles, ligaments,bones, nerves, and blood vessels attaching the gillarches to the skull were carefully scraped or cut free.A cut was then made through the pre-pectoral areato free the ventral attachment of the gill arches to thebody (cut passes through ventral aorta and, depend-ing on taxon, may slice through the urohyal). Fromthis cut, a posterodorsally arching cut was made oneach side of the specimen in the tissue containing thepharyngocleithrales and attaching the gill arches tothe surface of the cleithrum. The sphincter esophagiwas then severed and the retractor dorsales severedfrom their attachments to the vertebrae. Any attachedtissues (viscera, nerves, etc.) were cut and the gillarches released. The variety of taxa dissected fre-quently required considerable ad hoc modificationsto this general procedure.After removal, the gill arches were partiallycleaned by picking away the most obvious blood ves-
sels, nerves, viscera, and extraneous fatty and con-nective tissues. The gill arches were then stained.Early in the study we used a KOH-alizarin red-s so-lution to stain bone and an acetic acid-ethyl alcoholsolution of alcian blue to stain cartilage (Dingerkusand Uhler 1977). We found, however, that the KOH-alizarin solution was destructive of the muscles andwe substituted a non-destructive ETOH-alizarin so-lution (Springer and Johnson 2000) for it. Regardlessof which alizarin solution was used, the muscles usu-ally acquired a pink color or, in the case of alcian, a
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blue-green color, enabling one to distinguish themmore easily from one another or surrounding tissues.After staining, the muscles were further cleaned ofextraneous tissues and described.Descriptions and illustrations were often done instages. In the first stage, as much information as pos-sible was recorded without disturbing the musclesother than to truncate the levators, if they obscuredthe other muscles. The muscles were then drawn. Inthe next and subsequent stages, muscles were bisect-ed or removed in order to expose hidden muscles ormuscle attachments, and at each stage, the drawingwas modified, usually unilaterally, to show additionalinformation. Levator muscles and the retractor dor-salis are truncated, without mention in most of theillustrations.Some muscle attachments were inferred withoutdissection by referring to cleared and stained prepa-rations. In so far as they were not removed duringcleaning, ligaments and miscellaneous connective tis-sues are included in the illustrations and mentionedin the descriptive accounts, but the absence of suchstructures from the illustrations or descriptions doesnot necessarily imply that they are actually absent.We strove for clarity in the illustrations, at thesame time attempting to portray the muscles as close-ly as possible to their actual appearance?bilateralasymmetry and anomalies were included. Photo-graphs would have been more accurate, perhaps, butmuch less clear. The plates, with a few exceptionsnoted in the plate legends, are based on single spec-imens. The specimen illustrated may not be typicalof the taxon in every detail illustrated. For this rea-son, if the reader notes a difference between a char-acter as coded in the PAUP data matrix (Appendix.Table 12) and the representation of that character inthe illustration, the description of the taxon shouldbe read for explanation. All such conflicts, however,may not be explained, e.g.. proportional or presence-absence characters that are distorted resulting fromparallax or are obscured in the view illustrated.Finally, during the course of determining obscuredmuscles and skeletal elements, the muscles of manyof the specimens were of necessity greatly damagedor removed and the specimens will be of limited orno use to future investigators. This is particularly trueof specimens that were prepared early in the studyusing a KOH-alizarin solution to stain the bones, asthe solution badly macerates the muscles, and its res-idue continues to do so over time.
Material
Institutional abbreviations denoting specimens arethose proposed by Leviton et al. (1985) and Levitonand Gibbs (1988).Relevant study material is reported at the begin-
ning of each descriptive account. If an illustration isindicated, the first indicated specimen is usually theone illustrated. Occasionally, additional material ex-amined for only one or a few characters is cited inthe text. A list of material (see Acanthomorpha sec-tion), specifically for acanthomorphs, not otherwisementioned in the text, was examined for osteologicalinformation, and served as the source for informationin Table 8. In general, small specimens in the rangeof 50-150 mm SL were selected for dissection. De-pending on the taxon, these may have been adults orjuveniles (some taxa rarely attain a length of even 50mm as adults: specimens longer than 150 mm wereused for taxa with proportionally small heads, e.g.,eel-like forms). For many taxa, only one specimenwas available for study, but even for taxa where morespecimens were available, only one specimen mayhave been studied if the dissection was successful(i.e., little or no damage). We recognize this defi-ciency, but in a survey of the magnitude of the pre-sent one. we had to limit the depth of our examina-tions: any taxon could have been the basis for anindependent study, and we dissected about 500 spec-imens comprising 208 families and about 400 spe-cies.Muscles are highly variable in their expression,sometimes varying bilaterally in the same individual,between individuals of the same species, or ontoge-netically. Where our material and examinations per-mitted, and we considered the variation important, wediscuss variation.
Muscles and Skeletal ElementsThe muscles mentioned in the text and on the il-lustrations are listed with their definitions and the ab-breviations we use to represent them in the section
"Abbreviations and Definitions for AnatomicalStructures." The skeletal elements are similarly in-cluded, but only some are defined. A discussion ofepibranchial 5 (Eb5) is given below because the in-terpretation of the presence or absence of this ele-ment is important in deciding the attachment of ad-ductor 5, and we use the opportunity to note newphylogenetic inferences based on the relationship ofEb5 to Eb4, ceratobranchial 4 (Cb4), and Cb5 (seesection below. "Epibranchials 5 and 4").Muscle names.?There is a wealth of names avail-able for many gill-arch muscles, but relatively feware standardized. Winterbottom (1974b) valiantly andmost recently attempted a synonymy of teleost fishmusculature. We utilize many of the names he rec-ognized as senior synonyms and that are commonlyemployed in the literature (e.g., levator externus, orexternal levator). However, we also use a few nameshe treated as synonyms, and for many muscles wedevise our own names. For these last muscles, our
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intent is to have the name indicate the attachmentsof the muscle, e.g., musculus laminalis dentalis 5-ceratobranchialis 4 (M. UP5-Cb4). a muscle origi-nating on upper pharyngeal tooth plate 5 and insert-ing on ceratobranchial 4. Such names may be un-wieldy, but they are descriptive, and one rarely needsto refer to them other than by their abbreviations.There is no law of priority with regard to musclenames, and we find that despite Winterbottom's at-tempt to standardize muscle names, complete stan-dardization in the literature has not occurred, e.g.,names of ceratodontid muscles have not been stan-dardized. There is the additional problem, which wehave not solved, of assuring homology of usage. Ournomenclature is based variably on morphological to-pology and/or homology. If the reader is in doubt ofour usage from the context of our application or dis-cussion, it is probably safest to assume topology.Muscle types.?Aside from functional anatomicaltypes (e.g., contractors, extensors), we recognize twogeneral positional types of dorsal gill-arch muscles,those that are bilaterally paired, i.e., present on each
side, and those that are transverse, extending fromone side to the other. An interpretive problem devel-ops when the middle portion of a transverse muscleis lost, as often occurs in acanthomorphs (e.g., trans-versus pharyngobranchialis 2 of Pomacentridae,Plate 160; transversus epibranchialis 2 of Embioto-cidae, Plate 162.1); we have no evidence for thetransverse fusion of a bilaterally paired muscle).Origins and insertions.?Levators originate on thecranium, or in the case of the levator posterior, mayoriginate from the body musculature. Bilaterallypaired muscles attaching pharyngobranchial elementsto epibranchials and/or ceratobranchials are consid-ered to originate on the pharyngobranchial elements.Retractores dorsales are considered traditionally tooriginate on the vertebral column and insert on pha-ryngeal elements. Muscles attaching an epibranchialto another epibranchial {recti dorsales = RecD) onthe same side of the gill arches are considered (tra-ditionally) to originate on the more posterior epi-branchial: RecD4 originates on epibranchial 4 andinserts on epibranchial 3.Anatomical orientation.?Anterior and posteriorare defined by the dorsal mid-longitudinal axis of thefish that runs between the pharyngobranchials. Me-dial, or proximal, and distal, or lateral, are, in effect,positions relative to the pharyngobranchials. Anteriorand posterior refer to the positions relative to thehead and tail of a fish, but the medial angle of artic-ulation of epibranchials, and of the ceratobranchialswith the epibranchials, often imposes an almost lon-gitudinal orientation on them?thus, the proximaland distal ends of the epibranchials deceptively ap-pear to be the anterior and posterior ends. The de-
scriptions are based on the anatomical position, notthe deceptive appearances.The major axis of a pharyngeal element may beoriented almost perpendicularly, in which case its an-terior end may be described as dorsal and its dorsalsurface described as posterior.Transverse muscles.?Interpretation of the individ-ual muscles comprising the transversus dorsalis fre-quently involved subjectivity because of the natureor degree of continuity between the various compo-nents. Some components were unambiguously defin-able, but others were not (e.g., what degree of sep-aration of the components of transversus pharyngo-branchialis 3-epibranchialis 4 (TPb3-Eb4) shouldexist before recognizing the components as separatemuscles, TPb3 and TEb4: only complete disconti-nuity of the components; continuity, but by only afew muscle fibers, etc.?). For some taxa, where morethan one specimen was dissected, the componentsmight be continuous in one specimen and discontin-uous in another. The interpretations, therefore, con-tain a degree of subjectivity, which the illustrations
reflect.Acanthomorph accessory cartilages.?Rosen(1984:3, 25. fig. 25a) first noted the presence of anaccessory cartilage (AC) at the joint of an epibran-chial and ceratobranchial in the gill arches of anacanthomorph (the fourth arch of Acanthurus), inwhich he indicated that it was of unknown signifi-cance. Rosen and Patterson (1990:9, fig. 42a) nextcalled attention to an acanthomorph AC (in Lobotes,also in the fourth arch) and again indicated that itwas of unknown significance, neglecting to mentionRosen's earlier finding. We know of no other reportsof accessory cartilages at the Eb-Cb joints of acan-thomorphs, which is the only group in which theyoccur. Among acanthomorphs, Eb-Cb joint accessorycartilages are predominantly restricted to perciforms(Table 8). Although ACs may occur in any gill arch,they are most commonly associated with the fourtharch. In acanthomorphs having AC4, Ad5 usually at-taches to it.AC4 superficially resembles Eb5, which, as firstnoted by Baldwin and Johnson (1996), is restrictedto pre-acanthomorphs (here further restricted to pre-Ctenosquamata, see Table 6). Based on examinationof larvae of a few taxa (Osmeridae, Osmerus, 1 7 mmSL; Characidae, Corynopoma, >4 mm SL; Chloro-phthalmidae, Chlorophthalmus, 12-13 mm SL), Eb5is autogenous early in ontogeny (but may fuse on-togenetically with Eb4 and/or Cb4). In contrast, theacanthomorph AC4 is always associated with the dis-tal (usually posterodistal) end of Cb4, and appears tobud off Cb4 relatively late in ontogeny.In larval Morone (Moronidae) as large as 14 mmSL (all gill-arch elements with substantial ossificationand vertebral column fully differentiated and ossi-
NUMBER 1
1
fied), AC4 is absent and there is no evidence of acartilaginous projection from Cb4 from which AC4might form. In a 28 mm SL specimen of Morone,there is a projection extending from the cartilaginoustip of Cb4, which, we presume is the precursor ofthe autogenous AC4 present in the much larger spec-imens of Morone (100 mm SL) used for the muscledescriptions. Among four cleared and stained speci-mens of Ambassis sp., USNM 218805, AC4 on bothsides of two specimens, 30.7-35.4 mm SL, appear tobe in the process of budding off; one specimen, 39.0,has autogenous AC4s on both sides, and one speci-men, 45.6 mm SL, has an autogenous AC4 on oneside, and a bud on the other. In a specimen of Am-bassis buruensis, USNM 305331, 55 mm SL, a well-developed process extends posteriorly from the distalend of Cb4 with no evidence that budding off mightoccur.Although we did not investigate them, AC 1-3 andAC5 (the last known only in Lates, Latidae) probablyalso develop as buds off the distal ends of CM -3 andCb5, with which they are very closely associated (CTsurrounding the cartilaginous distal ends of the Cbsenvelops the associated AC).There is a problem in polarizing the character stateof AC4, but based on its appearance in Velifer, wearbitrarily treat its presence as an acanthomorph syn-apomorphy.Epibranchials 5 and 4.?The occurrence and in-terpretation of Eb5 has received considerable atten-tion (Nelson 1967d; Greenwood and Rosen 1971;Rosen 1974; Fink and Fink 1996; Johnson and Pat-terson 1996, 1997). It has not been established, how-ever, that the element, which is always cartilaginous,is an epibranchial (Nelson 1969a:520, reported Eb5was ossified on one side of one specimen of a gym-notid).Eb5, which is usually constrained as articulatingwith the distal end of Eb4 (e.g.. Nelson 1969a:520),is reported to be lacking in all ctenosquamates (Bald-win and Johnson 1996:372).Eb5 varies from being completely autogenous topartially fused with the distal end of Eb4, to puta-tively, completely fused with the distal end of Eb4,or infrequently, as we believe, with the distal end ofCb4 (only Albula). We find that in most groups withan autogenous Eb5, it is more closely associated withthe distal end of Cb4 than it is with Eb4, and it ap-pears to be fused with Cb4 in Albula based on com-parison with Eb5 and its association with Cb4 inPterothrissus, also Albulidae).Except for Osmerus (Osmeridae), it is problematicif Eb5 is always present in taxa in which the elementis interpreted as partially or completely fused to Eb4.The possible alternatives are that it has been com-pletely lost secondarily after fusion, or lost indepen-dently before fusion. Johnson and Patterson (1996:
275) observed that Eb5 is autogenous in O. mordaxlarvae, but becomes fused to Eb4 in later stages. Finkand Fink (1996:231) reported an autogenous Eb5 ina Gonorynchus larva, 18.5 mm SL, which is not pre-sent at later stages, but Johnson and Patterson (1997:597), who examined Fink and Fink's specimens, wereunable to find this cartilage. We also examined Finkand Fink's (1996:231) specimens (partially errone-ously cited; see Johnson and Patterson 1997:597, forcorrect citation) and confirm Johnson and Patterson'sfindings.In the plesiomorphic clupeomorph, Denticeps, theautogenous Eb5 attaches ventrally to the dorsodistalsurface of Cb4 and anteroventrally to the ventrodistalend of the rod-like Eb4. In clupeoids with an autog-enous Eb5, however, the element attaches ventrallyto the dorsodistal end of Cb4 and articulates closelydorsally and ventrally, but not in between, with thevertically expanded mostly bony distal end of Eb4(e.g., Dussumieria, Plate 29: B). Eb5 thus forms thecartilaginous distal border of a foramen throughwhich the posteriormost efferent artery passes (Nel-son 1969a:520); the proximal border of the foramenis mostly bony. Based on this landmark foramen.Nelson considered that Eb5 is present in clupeo-morphs in which there is no joint line dorsally be-tween it and the dorsodistal end of Eb4. but there isone at the ventrodistal end (foramen complete), orthe ventral portion of the foramen is open (foramenincomplete). Nelson did not specifically indicate thefusion of Eb5 with Eb4 in those clupeomorph taxain which the foramen is complete or the foramen iscompletely surrounded by cartilage with no joint linedorsally or ventrally. One can reasonably assume (asdid Rosen 1974), however, that based on the config-uration of the distal end of Eb4, that Eb5 is presentand that it has fused dorsally and ventrally with Eb4.There also appears to be inferential support for fu-sion of Eb5 and Eb4 in certain salmoniforms. In sal-monids, Eb5 is either autogenous (e.g., Oncorhyn-chus, Plate 36) or, apparently, fused dorsally withEb4 (e.g., Prosopium, foramen open ventrally; Ro-sen, 1974:fig. 9e). Johnson and Patterson (1997:596-597) appear to accept Eb5 as present in the ostario-physan Gonorynchus based on the ventrally open fo-ramen in the elongate posterior cartilaginous exten-sion of Eb4.Circumstantial evidence based on the attachmentsof adductor 5 (Ad5) supports the probable fusion ofEb5 with Eb4 in pre-acanthomorphs: Ad5, which al-ways attaches to Cb5 at one end, almost always at-taches to an autogenous Eb5 at the other end, or tothe distal end of Eb4 when Eb5 is putatively fusedwith Eb4 (Table 6).Probably extrapolating from the clupeomorph con-ditions in which Eb5 is fused dorsally to Eb4 and theforamen is open ventrally, Nelson (1967d:75, fig. Id;
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
also our Plate 7) considered that Eb5 might be pre-sent in the plesiomorphic osteoglossomorph Hiodon.In other osteoglossomorphs, the state of the foramenis variable and, with the exception of Pantodon,mimics the states of the clupeoids. Only Pantodon,among the osteoglossomorphs, has an autogenousEb5 (Plate 13B), but its articulations (one end withCb4 and the other with an accessory cartilage attach-ing to Cb5) are different from those that might give
rise to the putatively fused conditions seen in the oth-er osteoglossomorphs.Excluding Albula, we code three character statesfor Eb5 (Table 6): absent (0), autogenous (1), andputatively partially or completely fused with Eb4 (2).State 2 is interpreted based on the appearance of thedistal end of Eb4 (presence of a complete or incom-plete foramen). If the Eb5 character states are appliedto the pre-acanthomorph cladogram, it must be con-cluded parsimoniously that state 2 evolved indepen-dently in the Osteoglossomorpha and Clupeoidei;hence, there is no evidence for the existence of anindependent or fused Eb5 at the base of the Osteo-glossomorpha (the autogenous Eb5, and its articula-tions, in Pantodon is an autapomorphy). Eb5 autog-enous first appears as a basal synapomorphy in theElopocephala.Perhaps, ontogenetic studies of the osteoglosso-morphs will reveal, as in Osmerus, that an autoge-nous Eb5 is present and becomes fused with Eb4during development. If such is the case, however, thefusion of Eb5 to Eb4 would still have been achievedindependently by the Clupeocephala.There is another possible solution to the problem:the existing cladogram is incorrect. If Osteoglosso-morpha is replaced by Elopomorpha and placed asthe sister group of the Clupeomorpha. an autogenousEb5 becomes a synapomorphy of the newly com-posed Teleostei, and a fused Eb4-Eb5 becomes a syn-apomorphy of the Clupeomorpha + Osteoglosso-morpha. The reason for suggesting these changes isthat Arratia (1999), based on a limited number ofrecent taxa, but citing studies by other authors whoused additional characters, proposed that Elopiformesare the sister group of all other Teleostei and that theOsteoglossomorpha are close to the Clupeomorpha.The distribution of suprapharyngobranchial 1 (SPbl)also makes more sense parsimoniously if the Elopo-morpha exchange places with the Osteoglossomorphain the cladogram (see discussion of LI1 in pre-acan-thomorph Results section).This extended discussion bears on character statesfor the attachment of Ad5. To differentiate states inwhich Ad5 attaches to an autogenous Eb5 from thosein which it attaches to a putatively (partially or com-pletely) fused Eb5 with Eb4 or Eb5 with Cb4, weindicate the latter two skeletal conditions as "Eb4*"or "Cb4 :; . dicate these abbreviations, where
applicable, in the discussions and on the illustrationsof the Elopocephala, but not the Osteoglossomorpha.
Abbreviations and Definitions for AnatomicalStructuresNote: the following definitions occasionally con-tain important caveats on the interpretation of certainmuscles, e.g., ER, the esophageal raphe.
*?following a pre-acanthomorph Eb4 or Cb4 indi-cates putative fusion of Eb5 with Eb4 or Cb4. Seesection "Epibranchials 5 and 4" for discussion.AB?autogenous bone; tiny bone attached to tip ofEb4 levator process and dorsal end of Eb5; onlyin Cyprinidae.AC?accessory cartilage; mostly restricted to acan-thomorphs. Among pre-acanthomorphs havingACs (Polypterus, Atractosteus, Pantodon, Diplo-mystes, Galaxias), the ACs may occur in a varietyof locations and may be normal or adventitious inthe taxon in which they occur. Among acantho-morphs, ACs are common, but occur predomi-nantly at the joint between the distal ends of anEb-Cb pair, hence AC1, AC2, etc, with AC4 themost common in occurrence. Except for the rarelyoccurring AC5, acanthomorph ACs occurring atpositions other than at an Eb-Cb joint are not num-bered. See section "Acanthomorph accessory car-tilages" for discussion.Ad?adductor, adductores. Adl, Ad2, etc., adductorof 1st arch, 2nd arch, etc; plural. Ads, Adls, etc.;muscle attaching Eb to Cb of a single arch, exceptAd5, which attaches Cb5 variously to one or moreof the following: Cb4, AC4, Eb4, or Eb5. Ad4dorsal attachment is on dorsoposterior or ventralsurface of Eb4, often beginning anterior to OP onEb4 and usually extending laterally further thanOP. When present, one or more of first three Adsmay be completely obscured by, or fused with, anoverlying gill-filament muscle (GFM, q.v.) of samearch. In general. Ads are better developed thanGFMs. We occasionally had difficulty distinguish-ing Ads from GFMs in acanthomorphs, and sub-jectivity in deciding may have resulted in someerroneous decisions.Among, pre-acanthomorphs. Ads 1?3 occur onlyin Polyodontidae, Notacanthidae, Cyprinidae, andpossibly Anguillidae. They are variably presentamong acanthomoiphs, most commonly amongpercomorphs. It is unlikely that Adl?3 of pre-acanthomorphs and acanthomorphs are homo-logues. It is unlikely, furthermore, that the acan-thomorph Ads 1-3, which are absent in basal acan-thomorphs, are serial homologues of Ad4 and Ad5in acanthomorphs, which occur early in pre-acan-thomorph phylogeny.The identification of Ad5 in various non-perci-
NUMBER 11
form acanthomorphs in which ER is also presentcan be problematic. In some cases, it appears thatAd5 is absent, in others Ad5 ends dorsally at araphe with the ventral end of OP, and in others thatAd5 is present as the lateral portion of what oth-erwise appears to be OP. In pre-acanthomorphs,apparent fusion of Ad5 medially with OP ventro-laterally is common. Additional study of ER, Ad5,and OP is warranted. See also OP below and re-marks following Ad5 in description of Arapaima(Arapaimidae).Ad4'?adductor 4 primus, attaches to the anterodistalsurfaces of Eb4 and Cb4 (additional to Ad4); onlyin Notacanthidae.Bb?basibranchial, Bb3, Bb4, etc.; unpaired ventralgill-arch skeletal element.Cb?ceratobranchial; plural, Cbs; also Cbl, Cbls,
etc.; ventral gill-arch skeletal element.CPb
?
circumpharyngobranchialis; a sub-epithelialmuscle first described, but not named, by Anker(1978:261) for a cichlid muscle, and apparently notreported subsequently. CPb is present in variousperciform fishes and is often very well developed.
It appears to originate from the SO longitudinalmuscle layer, extending anteriorly, and surround-ing or only bordering, and attaching variously toPb2, Pb3, and UP4. In some perciforms (e.g., cich-lids, pomacentrids) TPb2a, may represent a dis-junct portion of CPb.CT?connective tissue.Eb?epibranchial; plural, Ebs; also, Ebl, Ebls, Eb2,
etc.; dorsal gill-arch skeletal element.Eb4* indicates putative fusion of Eb5 with Eb4; onlyin pre-acanthomorphs.Epibranchial flange?a dorsolateral or anterolateralextension of the dorsodistal bony edge of an epi-branchial such that it partly or completely
"shields" the cartilaginous distal end. Present onlyin some percomorphs and usually restricted to Eb4,although flanges may be present on other Ebs. Theflanges are much reduced in size in many taxa (anddifficult to show on our illustrations). A well-de-veloped Eb4 flange is present, e.g., in all membersof the Labroidei, Opistognathidae, Pseudochromi-dae, Grammatidae; variably developed in Plesiop-idae (well developed in Assessor and Paraple-siops; weakly developed in Trachinops and Belo-nepterygion; absent in Acanthoplesiops and Noto-graptus); well developed in all atherinomorphsexcept very weakly developed in belonids and ab-sent in scomberesocids. Also very weakly devel-oped in the mugilid, Agonostomus. In labroids, thecartilaginous distal end of Eb4 has been lost andthe cartilaginous end of Cb4 attaches by a tendonto the ventral surface of the flange.EO?epibranchial organ; plural, EOs; in pre-acatho-morphs usually formed, at least in part, by mod-
erate to extraordinary expansion of the cartilagi-nous distal end of Eb4; in acanthomorphs (onlystromateoids), EO involves an out pouching of theesophagus and distal end of Eb4 is not involved.Certain anabantoid families (e.g., Channidae. An-abantidae) have a suprabranchial organ which in-volves modification of the first epibranchial.ER?esophageal raphe; a fine line of connective tis-sue or myoseptum usually dividing OP transverse-ly at about mid-level or demarcating the ventralend of OP and separating it from Ad5 and/or SO.Very common in pre-acanthomorphs, but frequent-ly difficult to decide the constitution of the musclefibers ventral to ER: SO, OP, or Ad5. Relativelyuncommon in acanthomorphs, but when presentusually appears to separate OP ventrally from Ad5,resulting in OP attaching ventrally to Cb4 ratherthan Cb5 (its usual ventral attachment in acantho-morphs), and Ad5 attaching to Cb4 well medial todistal end of bone, rather than to the distal end.GC?gongyloid cartilage, first named by Di Dario(2002) and first described by him in print, but firstnoted by Nelson (1966a:157) in his Ph.D. disser-tation; present only in engrauloids, pristigasteroids,and Chanos (Chanidae). See discussions in Addi-tional remarks sections under Cetengraulis andChanos.GFM; GFM1, 2. 3?gill filament muscle. Here con-sidered to be essentially the same as Winterbot-tom's (1974b:260 and fig. 26c) interbranchialesabductores: "extrinsic [gill] filament muscles . . .connecting the bases of the oral filaments to thegill arch (cerato- or epibranchial [we would mod-ify this to cerato- and/or epibranchial]) . . . may[also] become intimately associated with the gillrakers . . ." They are often inconspicuous, fine, andstringy and are frequently destroyed when strip-ping gill filaments from the gill-arches. Those ofthe second and third arches in acanthomorphs mayextend dorsoanteriorly and attach to the posterioredge of the preceding arch or they may continuedorsomedially on the dorsal surfaces of Eb2 andEb3, that of the second arch sometimes meetingthe lateral end of TEb2. We report them only insome acanthomorphs, and only when they are con-spicuous or fused with an adductor (Ad). In sometaxa, they are questionably distinct from Ads(q.v.); decision on assignment as GFM or Ad issomewhat arbitrary. Additional study of the acan-thomorph Ads and GFMs is desirable.Winterbottom (1974b:259-260 and fig. 26) alsorecognized interbranchiales adductores, musclesattaching to the gill filaments of both hemibranchsof a single gill arch. We do not report on thesemuscles.Hb?hypobranchial; Hbl, etc.; plural, Hbs; ventralgill-arch skeletal element.
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
IAB?interarcual bone; a putatively ossified IAC,only in Synbranchidae and Carapidae.IAC?interarcual cartilage; usually an autogenousrod-like cartilage joining Ebl uncinate process toPb2; found only among acanthomorphs.IAC2?interarcual cartilage 2; autogenous cartilagejoining Eb2 and Pb2; only in Menidae.Interbranchiales abductores, adductores?see GFM.LCb
?
levator ceratobranchialis (i.e., LCb2, LCb4,LCb5) muscle originating on skull and insertingon a Cb; only in pre-acanthomorph Cyprinidae andacanthomorph Adrianichthyidae. See also remarksfollowing LCb5 in muscle description of Cyprini-dae for comment on homology. Not to be confusedwith LE5 of Dipnoi, which inserts on Cb5.LCb5A
?
levator ceratobranchialis 5 accessorius;muscle originating in supratemporal fossa of skullnear origin of LCb5, wrapping medially first, thenanteriorly around LCb5 and inserting in CT padattached to anterolateral surface of Cb5; only inCyprinidae. This muscle appears to be the same asHolstvoogd's (1965:216, and fig. 12b) M. troch-learis, which Winterbottom (1974:253) synony-mized with LP. It is also the same as Winterbot-tom's LP internus (internus and externus portionsnot labeled), as indicated in his fig. 22 (Cyprinus).Muscle re-named by us at suggestion of R. Win-terbottom, who correctly noted (in litt.) that tro-chelar most often refers to cranial nerve IV, whichsupplies the superior oblique muscle of the eye,hence, inappropriately applied to a gill arch leva-tor.lat?lateral.LE
?
levator externus or external levator; muscleoriginating on cranium and inserting on an Eb:LEI, LE2, etc.; plural LEs, LEls, etc. With rareexception (Carapidae), LEs originate on the skull,typically in a cluster or continuous line, usuallytogether with Lis; however, LE4 may be displacedposteriorly in some taxa, especially those with EOs(see also LP). LE3 always inserts on or close tothe Eb3 uncinate process in elopocephalans (ig-noring those taxa in which the uncinate process isabsent). According to Vanderwolle et al. (1998),
all carapid levators originate on the medial surfaceof the hyomandibula.LEI ', LE2', etc.
?
levator externus 1 primus, etc.; thesecond of two LEls, etc., arbitrarily designated,presumably the result of the division of an LE.Levator process (on Eb4)?a cartilaginously tippedprocess, lateral or posterior to the uncinate process(q.v.) on which LE4 and/or LP usually inserts.Presence or absence of the process may be onto-genetically associated: process isolated from distalcartilaginous end of Eb, or, in acanthomorphs, car-tilage lost during ontogeny by osseous exclusion.In the absence of a cartilage tip, the process is
arbitrarily considered absent.Johnson and Patterson (1996:272?275) discussthe confusion and phylogeny of uncinate and le-vator processes on Eb4 and note that an Eb4 un-cinate process "characterizes acanthomorphs,"and that "Loss of a separate levator process ap-pears to be a synapomorphy of Acanthopterygii,although it may occasionally occur secondarilywithin Percomorpha . . ." The presence of an Eb4levator process in percomorphs is more commonthan Johnson and Patterson implied. It is frequent-ly present in generally considered plesiomorphicpercomorphs (e.g., Acropomatidae, Percichthyi-dae, Moronidae, Scorpaenidae, Epigonidae, Apo-gonidae, Serranidae, Lutjanidae, Priacanthidae, La-teolabrachidae etc.) as well as in some specializedmembers (e.g., Opistognathidae, Trachinidae).LI
?
levator internus or internal levator; muscle orig-inating on cranium and inserting, variously, onPb2, Pb3, Pb4, UP4, or UP5 (LI1, LUP5, etc.; plu-
ral, Lis, LIls, etc.); LI1 inserts on Pb2 (and/or Pb3in some acanthomorphs; especially Blennioidei,which lack Pb2); LI2 normally inserts on Pb3; LI3,absent in ctenosquamates, variously inserts on Pb3,Pb4, UP4, UP5; LI4, only in Diplomystidae, in-serts on UP4. Some confusion may arise whencomparing our Lis with those mentioned in theliterature, as many authors number the LI based onthe Pb to which it attaches; hence our LI1 is oftenreferred to as LI2 in the literature, our LI2 as LI3,and our LI3 as LI4.LI la, LIlp
?
levator internus 1 anterioris, levator in-terims I posterioris; LI1 represented by two lon-gitudinally separated muscles, both inserting onPb2; only in Myctophidae.LI1'
?
levator internus J primus; the posterior divi-sion of LI 1 inserting on Pb3 anteriorly, often close-ly juxtaposed to LI1 insertion on Pb2; frequentlypresent in, but not limited to, Gobioidei.LI3 (part)?a separate, probably anomalous, basalportion of LI3; only in Heterotis (Osteoglosso-morpha).LI3' levator internus 3 primus; second of two LI3sinserting on Pb4; only in Searsia, (Platytroctidae).lig?ligament.LP?levator posterior (or levator posterioris); mostlyrestricted to acanthomorphs. but present in someclupeoids and ostariophysans. Muscle originatingvariously on skull or body musculature and usuallyinserting on Eb4 together with LE4. In some taxa,fused with LE4 or coalesced in a musculous andconnective tissue sheet with LE4 and/or PP. whichattaches along edges of gill arches 4 and 5, andthe individual muscles are not clearly separable.When LP is clearly distinguished, its origin is pos-terior or posteromedial to LE4 origin and usuallywell removed from it. In taxa with a single levator
NUMBER 11 9
muscle on Eb4 (including, however, LE4') thatoriginates with other LEs and/or Lis, there is noproblem identifying the muscle as LE4 (and LE4')because LP never clusters with the other LEs. Inpre-acanthomorph taxa that have the origin of thesingle levator on Eb4 well posterior to those of theother levators, one might be tempted to designatethe muscle LP, but examination of the muscles inrelated forms invariably indicates that only LE4 ispresent (i.e., it joins LE cluster). Additionally, LE4generally inclines anteriorly, whereas LP frequent-ly inclines medially or anteromedially. Only threeacanthomorph taxa appear to have lost LE4 andretained LP: Pholidichthys (Pholidichthyidae), Spi-nachia (Gasterosteidae), and, possibly, Echenei-dae.med?medial.mid?middle.M.
?
musculus: muscle.M. Ebl-Cbl
?
M. epibranchialis I-ceratobranchialis7; muscle joining dorsomedial end of Ebl withdorsoanterodistal end of Cbl (only in Calliony-midae).M. Ebl-IAC M. epibranchialis 1-cartilago inter-arcualis; muscle originating on Ebl and attachingto IAC (only in Adrianichthyidae).M. Eb4-F
?
M. epibranchialis 4 faucis; muscle orig-inating on Eb4 and meshing with SO in throat(Latin, faucis) region (only in Blenniidae).M. Intrb
?
M. intrabranchialis (pi. intrabranchiales);M. Intrb 1, Intrb 2 etc; muscle present in the CT(variously termed a diaphragm or septum) betweenthe hemibranchs of a single branchial arch, over-lain by the gill filaments, which must be scrapedaway to expose it. Known only for Chondrichthy-
es, in which they have been termed interbranchi-ales (Marion, 1905:905 & figs. 7. 8. 12; Daniel,1934:105 & fig. 108) or constrictor branchiales(Edgeworth, 1935:129), and Dipnoi, in which theyhave also been termed interbranchiales (Furbrin-ger, 1904:488) or constrictor branchiales (Edge-worth, 1935:129; Fox, 1965:490). Here renamedto avoid confusion with the "interbranchiales"(which include interbranchiales adductores andabductores), originally named by Winterbottom(1974b:259 & fig. 26) for small teleostean musclesattaching to the gill filaments (see also GFM).Edgeworth (1935:129) erroneously reported M.Intrbs in acipenserids (see Additional remarks sec-tion under Acipenser ruthenus).M. Pb2-Ebl
?
M. pharyngobranchialis 2-epibran-chialis 7; muscle originating on Pb2 and insertingon Ebl; only in mormyrids and some anguilli-forms.M. Pb2-Eb2 M. pharyngobranchialis 2-epibran-chialis 2; muscle originating on Pb2 and insertingon Eb2. According to Winterbottom's (1974b:253)
definition of obliquui dorsales, any muscle origi-nating on a Pb and inserting on an Eb could betermed an OD, but to do so might cause confusion.Our M. Pb2-Eb2 has been designated OD2 byEndo (2002:101) for gadiforms, in which he con-sidered its presence a specialization. In pre-acan-thomorphs, OD2 is present only in the osteoglos-somorphs (Hiodon and Heterotis), where it origi-nates on Pb3.In most acanthomorphs, the muscles we treat asobliquui dorsales originate entirely or primarily onPb3 and insert on Eb3 and/or Eb4. To distinguishthe acanthomorph OD2 (in Brotula, which also hasM. Pb2-Eb2), we elected to denominate the "OD"originating on Pb2 as a new muscle: M. Pb2-Eb2;likewise, we designate other "ODs" as M. Pbs-Ebs to avoid confusion. See also OD.M. Pb3-Cb5
?
M. pharyngobranchialis 3'-ceratobran-chialis 5; muscle originating on Pb3 and insertingon Cb5 (only in Sparidae and Centracanthidae).M. Pb3-Ebl
?
M. pharyngobranchialis 3-epibran-chialis 7; muscle originating on Pb3 and insertingon Ebl (only in Callionymidae).M. Pb3-Eb2 M. pharyngobranchialis 3-epibran-chialis 2; muscle originating on Pb3 and insertingon Eb2 (only in Pomatomidae).M. Pb3-Eb3-Eb4
?
M. pharyngobranchialis3-epi-branchialis 3-epibranchialis 4; muscle originatingon Pb3 and inserting on Eb3 and Eb4 (only ingobiid Gnatholepis).M. Pb3-Eb3 M. pharyngobranchialis 3-epibran-chialis 3; muscle originating on Pb3 and insertingon Eb3 (among pre-acanthomorphs, only in No-vumbra, Umbridae; among acanthomorphs, at leastin some gobioids).M. Pb3-Eb3-Eb2 M. pharyngobranchialis 3-epi-branchialis 3-epibranchialis 2; short muscle orig-inating on Pb3 and inserting on Eb3 and Eb2 (onlyin Gymnarchidae).M. Pb3-Eb4-Eb2-Cb3
?
M. pharyngobranchialis 3-epibranchialis 4-epibranchialis 2-ceratobranchial-is 3; muscle originating on Pb3 and inserting onEb4, Eb2, and Cb3 (only in Callionymus).M. Pb3p
?
M. pharyngobranchialis3 posterior, shortcone-like muscle attaching anteriorly to Pb3 andinserting, apparently without attaching, into a con-cavity at the anterior end of the first vertebra.Function problematic; found only in Hemiramphi-dae and Exocoetidae.M. Pb3-Pb4-Eb2
?
M. pharyngobranchialis 3-phar-yngobranchialis 4-epibranchialis 2?muscle orig-inating on Pb3 and Pb4 and inserting on Eb2; onlyin the osteoglossomorph Gymnarchidae.M. Pb3-UP4 M. pharyngobranchialis 3-laminalisdentalis 4; muscle attaching to Pb3 and UP4; onlyin Embiotocidae.M. Pb4-Eb2 M. pharyngobranchialis 4-epibran-
10 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
chialis 2; muscle originating on Pb4 and insertingon Eb2; only in notopteroids.M. SO-Pb2
?
M. sphinctoris esophagi-pharyngo-branchialis 2; muscle originating on each side asan anterior extension of the transverse layer of thesphincter oesophagi, becoming discrete anteriorlyas it extends along medial side of Pbs and insertson Pb2 and, variously, Eb4; noted in leiognathids,but probably more widely distributed.M. SO-Pb3 M. sphinctoris esophagi-pharyngo-branchialis 3; muscle originating on each side asan anterior extension of the dorsal SO longitudinalmuscle layer, becoming discrete anteriorly and in-serting on Pb3. More common than noted in thedescriptions or on the plates as it was identifiedlate in the study (e.g., Ditropichthys, Cetomimidae.Plate 74).M. SO-Pb4 M. sphinctoris esophagi-pharyngo-branchialis 4; dorsolateral extension of SO that at-taches to Pb4, only in Psenopsis, Centrolophidae.M. SPb2-Eb2
?
M. suprapharyngobranchialis 2-epi-branchialis 2; interrupted portion of LE2 originat-ing ventrolaterally on SPb2 and inserting on Eb2;only in Acipenseridae.M. SPb2L
?
M. suprapharyngobranchialis 2 lateral-is; interrupted portion of LE2 originating on skulland inserting on SPb2 dorsolaterally; only in Aci-penseridae.M. SPb2-LEl M. suprapharyngobranchialis 2-le-vator externus 1; probably a component of LE2originating from CT on medial surface of LEI andinserting anteriorly on mid-medial surface ofSPb2; only in Acipenseridae.M. SPb2Ma M. suprapharyngobranchialis 2 medi-alis anterioris; interrupted portion of LE2 origi-nating on skull and inserting on SPb2 anterome-dially; only in Acipenseridae.M. SPb2Mp^M. suprapharyngobranchialis 2 me-dialis posterioris; interrupted portion of LE2 orig-inating on skull and inserting on SPb2 postero-medially; only in Acipenseridae.M. TEb2-Pb2
?
M. transversus epibranchialis?
2
pharyngobranchialis 2. A part of TD arising fromventral surface of TEb2 and inserting on Pb2; nothomologous with TPb2, which arises dorsal or an-terior to TEb2; only in Cepolidae.M. UP4-Eb2 M. laminalis dentalis 4-epibranchialis2; muscle originating on UP4 and inserting onEb2; only in Albula.M. UP4-EM M. laminalis dentalis 4-epibranchialis4; muscle originating on UP4 and inserting onEb4; only in Congridae.M. UP4-Eb5-CM
?
M. laminalis dentalis 4-epibran-chialis5-ceratobra.nchia.lis 4 (not illustrated); mus-cle originating on UP4 and inserting on Eb5-Cb4joint; only in Megalopidae.M. UP5-CM M. laminalis dentalis 5-ceratobran-
chialis 4; muscle originating on UP5 and insertingon Cb4; very similar to M. UP5-Cb4-Eb5; only inAlbulidae and Gonostomatidae.M. UP5-Cb4-Eb5
?
M. laminalis dentalis 5-cerato-branchialis 4-epibranchialis 5\ muscle originatingon UP5 and inserting at inner angle of joint formedby Cb4 and Eb5; only in characoids, but very sim-ilar to M. UP5-Cb4.MPbl?mediopharyngobranchial; cartilage articulat-ing posteriorly with anterior or medial end of Eb 1
;
may comprise single, medial element; present onlyin Chanidae, Gonorhynchidae, and some Clupeo-idei.ObV3
?
obliquus ventralis 3 (not illustrated); ventralgill-arch muscle attaching to Hb3 and Cb3; mayalso insert on SCL.OD?(plural, ODs) obliquus dorsalis (obliquui dor-sales). We follow convention in describing thesemuscles as originating on Pbs and inserting onEbs. Gareth Nelson (commenting on a draft of theMS) noted that, functionally, it is more accurate toreverse the origin-insertion designations.ODs usually originate on Pb3, but origin mayinclude Pbl (only Diplomystes, Diplomystidae)and Pb4 (only pre-acanthomorphs) and Pb2 (onlyacanthomorphs). Designation derives from Eb onwhich muscle inserts. See M. Pb2-Eb2 for discus-sion of why this muscle is not considered to be anOD.OD2?origin on Pb3; only in osteoglossomorphs.OD3?insertion on Eb3 includes uncinate process,if present.OD3'?origin on Pb3 with, ventral to, or lateral toOD3 or OD3-4, becomes ventral to them pos-teriorly, and attaches on Eb3 dorsally ventral ormedial to uncinate process; except for Oncor-hynchus, present only in acanthomorphs.OD3-4?a complete or almost complete fusion ofOD3 and OD4, essentially restricted to acantho-morphs, in which origin is usually restricted toPb3 and insertions usually on Eb3 and Eb4 bonysurfaces supporting cartilaginous tips of unci-nate processes.OD4?in pre-acanthomorphs originates on Pb3,Pb3 and Pb4, or Pb4; in acanthomorphs origi-nates almost exclusively on Pb3; insertion onEb4 includes bony surface supporting cartilagetip of uncinate process, if present.OD4v?small ventral branch of OD4; only in Het-eropriacanthus (Priacanthidae).OD4'?originates on Pb4 in all pre-acanthomorphsexcept Megalops and Brycon, in which it orig-inates on Pb3. Among acanthomorphs, OD4' isonly present in percopsiforms, in which it orig-inates dorsal to OD4 or OD3-4 on Pb3, extendsposteriorly dorsal to them and inserts on the Eb4levator process.
NUMBER II I 1OP
?
obliquus posterioris; highly variable muscle,sometimes in as many as four parts, attaching dor-sally to the posterior surface of Eb4, usually me-dial to dorsal attachment of Ad4, but often almostcompletely overlapping Ad4 posteriorly. In mostpre-acanthomorphs, OP is usually interruptedtransversely at mid-length by ER, and in most ofthe few acanthomorphs that have it, ER usuallyseparates the ventral end of OP from Ad5 (see dis-cussion in Ad). OP medially is frequently insepa-rable from SO or ventrolaterally from Ad5. Insome pre-acanthomorph (e.g.. Amid) and mostacanthomorph taxa, OP is continuous, uninterrupt-ed by ER, and attaches ventrally to Cb5 near at-tachment of Ad5, although OP ventromedial edgemay join a restricted raphe with Ad5 posterodis-tally.Aerts (1982) reported that OP in cichlids com-prises three separate sections: medial, central (heretermed middle), and lateral, and that LE4 fuseswith the middle OP section to form a continuousmuscle extending from the origin of LE4 to theattachment of the OP middle section to Cb5. Thiscombined muscle has been called a "sling" byLauder and Liem ( 1983: 171 ) and Stiassny and Jen-sen (1987:284), and it also occurs in (but is notlimited to) labrids (broad sense), embiotocids, and,perhaps (our opinion) pomacentrids. and may in-clude participation by LP. Most acanthomorphsonly give evidence of having one OP section, mostprobably the middle section; we may have missedthe divisions early in our work and further studyis desirable.OP'
?
obliquus posterioris primus; slender muscle(possibly anomalous) originating on Eb4 levatorprocess and joining ER with OP; only in Albuli-dae.PP protractor pectoralis; muscle of pectoral girdle,occasionally illustrated and/or discussed, but onlywhen included in a CT sheet also containing, andusually not clearly distinguishable from, LE4 and/or LP. Greenwood and Lauder (1981) provide anextensive survey of this muscle in fishes.Pb?pharyngobranchial (commonly truncated spell-ing for "infrapharyngobranchial"; used for con-venience); Pbl, Pb2, etc.; dorsal gill-arch skeletalelement.PC pharyngoclavicularis, -es (or pharyngocleith-
ralis, -es), ventral gill-arch muscle, usually two oneach side, but only one present on each side in eelsand pre-halecostomes, but that of pre-halecostomesmay be divided. See also PCE, PCI below.PCa, PCp
?
pharyngoclavicularis anterioris, -poster-ioris; divisions of the single PC of the pre-hale-costome Polypterus.PCE, PCI pharyngoclavicularis externus, -interims(of Winterbottom, 1974b:267); ventral gill-arch
muscles attaching Cb5 to the cleithrum; both pres-ent on each side in most halecostome actinopter-ygians; one or both frequently illustrated in ourplates but usually not discussed (absence in illus-trations not intended to imply actual absence). PCIis illustrated or described in all acanthomorph taxain which the muscle attachment includes the distalend of Cb5 and/or joins a raphe with OP ventrally.PrO
?
protractor pharyngeus, anteriorly inclined le-vator-like muscle originating on ventral cranialsurface, extending posteriorly, and inserting ondorsal non-musculous esophageal connective tis-sue; present only in Neoceratodus (Dipnoi).RCb5E
?
retractor ceratobranchialis 5 externus;muscle originating on ventral basioccipital processand inserting dorsolaterally on Cb5; only in Cy-prinidae.RCb5I
?
retractor ceratobranchialis 5 interims; mus-cle originating as CT along dorsolateral surface of
vertical SO fold abutting basioccipital process andinserting by long tendon on CT pad attaching toCb5; only in Cyprinidae.RCb5T retractor ceratobranchialis 5 transversus;muscle originating medially from CT and SO, andinserting on dorsolateral margin of Cb5; only inCyprinidae. Appears to be the same as retractorpharyngeus superioris of Winterbottom, 1974b:258; fig. 22b). Only in Cyprinidae.RD retractor dorsalis; muscle usually originatingon anterior vertebrae and inserting variously onone or more of Pb3, Pb4, UP4. UP5, and Eb4.Origin and insertion usually not described by us.RDs may insert anteriorly or posteriorly on Pbs.and the difference is probably important. RDs maybe unpaired, branch only at beginning of insertion,comprise a bilateral pair (one RD on each side), abilateral pair and smaller unpaired median member(RD'), or vertical pair of muscles on each side. RDvaries from being incorporated almost entirelywithin the SO (ventral to the circular or transversemuscle layer) to being entirely external to SO (seealso SOD).RD' indicates either the unpaired median muscle be-tween the individual RDs of a bilateral pair or thedorsal muscle when RD consists of vertical pair ofmuscles on each side (see RD).RecCb rectus ceratobranchialis; short muscle con-necting distal ends of two successive Cbs; only inCallionymidae.RecCom
?
rectus communis, a ventral gill-arch mus-cle infrequently and only incidentally appearing inthe illustrations; not discussed in descriptions.RecD rectus dorsalis (plural recti dorsales); muscletypically joining epibranchial on one side with epi-branchial immediately anterior; RecD2, RecD3,RecD4, number derives from posterior epibranchi-
al, i.e., RecD2 originates on Eb2 and inserts on
12 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
RecCom
PCE
PCI
TV4
Fig. 1. Ventral view of gill-arches of Pempheris schomburgkii, USNM 318588 to show semicircular ligament (SCL). Both RecComsgreatly truncated; left-side PCE almost completely removed; basihyal removed. Ventrally elongate cartilaginous tip of posterior end ofBb3 joins SCL mid-posteriorly. Photograph extensively retouched.
Eb 1 ; RecD 1 , however, originates on Eb 1 and prob-ably inserts on skull or peters out in skin that roofsmouth; RecD5 (only Callionymus) origin includesCb5, insertion includes various skeletal elementsanteriorly. We have applied RecD to a variety ofproblematic muscles found in clearly unrelatedtaxa (e.g., Menidae, Callionymidae, Cyprinidae,Anguillidae). See also discussion following RecDsin Callionymus.RecV4
?
rectus ventralis 4 (Fig. 1); ventral gill-archmuscle attaching Cb4 to Hb3 and/or SCL.SCL?semicircular ligament (Fig. 1); anteriorly openU-shaped ligament attaching anteriorly to the ven-tromedial ends of Cb3 and Cb4 and, often, mid-posteriorly to ventral surface of cartilaginous pos-terior end of Bb3, which, in acanthomorphs, is of-
ten elongate and ventrally recurved. The ventralaorta divides into left and right branches, whichpass anteriorly on either side of the Bb3 attach-ment. ObV3 and RecV4 usually attach to SCL.The attachment of SCL to Bb3 is often obscuredin ventral view, and the connection is easily brokenduring dissection or in trying to determine if it isattached to Bb3. Several character states for theBb3 attachment to SCL were apparent in our dis-sections, but because of intermediates, we ana-lyzed SCL only for presence or absence. When weare certain of our observation, we report whenSCL is attached to Bb3. In the descriptions, whereSCL is merely described as present, we are uncer-tain as to whether it was free or attached to Bb3.Stiassny (1992:269-271) discussed and illustrated
NUMBER 11 13
SCL. Her statement that SCL is an "acanthomorphinnovation" is incorrect as SCL is present in sev-eral pre-acanthomorphs (Table 1 ; SCL of Novum-bra is especially similar to that of acanthomorphs).SL?standard length; all specimen lengths are SL,unless indicated otherwise.Sling?see OP.SO?sphincter esophagi; as generally recognized inthe literature.SOD?sphincter esophagi division; a narrow to broadband of SO transverse, or circular, muscle sepa-rated dorsally from the remainder of the transverselayer and passing dorsal to RDs. We imprecisely
restrict SOD to the condition in which RD extendsnoticeably anteriorly external to SOD before en-tering the transverse muscle layer (thus excludingthe Aulopiformes and Ateleopodiformes as havingSOD). SOD cannot be present if RD is absent, butSOD is not always present when RD is present.Although apparent in some of the acanthomorphillustrations, we may have failed to record the pres-ence of a fine, often inconspicuous mid-ventralbranch of SOD that separates the left and rightRDs.SPb
?
Suprapharyngobranchial; dorsal gill-arch skel-
etal element of endochondral origin articulatingwith the cranium and, normally, with Ebl (SPbl)or Eb2 (SPb2); present only in Latimeria and someactinopterans (i.e., Chondrostei, Ginglymodi, Ami-idae, Elopiformes, Albuliformes, and Platytrocti-dae).TD transversus dorsalis; transverse muscles attach-ing the gill-arch elements on one side with thoseon the other; not labeled as such on plates. Com-prises TDA and TDP and their components. TDAand TDP muscles may be continuous and on thesame level, or, most commonly in acanthomorphsTDA is somewhat dorsal to the level of TDP. TDAmay be broadly or narrowly continuous with TDPor completely separate. TDP muscles may be con-tinuous posteriorly with SO or SOD. Same namesfor component muscles of TDP or TDA reportedin different taxa do not necessarily imply homol-ogy; likewise, different names in different taxamay obscure homology.TDA
?
transversus dorsalis anterior; transverse mus-cles attaching to the anterior skeletal elements:Pb2, Ebl, Eb2 (e.g., TEb2; Pb3 attachments inacanthomorphs generally not reported in musclenames for TDA muscles); not labeled as such onplates. In acanthomorphs, the medial portion of aTDA muscle may be lost or replaced by tendinoustissue or a thick CT pad, resulting in a pair ofmuscles, each of which may attach secondarily toan additional skeletal element, e.g., interruptedTEb2, might attach to Pb3 as well as to Eb2.TDP transversus dorsalis posterior; transverse
muscles, attaching to the posterior skeletal ele-ments: Eb3, Eb4, Pb4, UP4 and/or to Pb3 in thearea joining these elements (e.g., TPb3-Eb3); notlabeled as such on plates.TD plexus?slender muscle branches joined to a me-dian CT sheet dorsal to TD; branches attach toEbl, Eb2, and Eb3; only in Acanthurus (Acan-thuridae).TEbl
?
transversus epibranchialis 1; essentially re-stricted to Labridae, Odacidae, Scaridae.TEbl-Eb2
?
transversus epibranchialis 1 -epibran-chialis 2; only in Pantodon (Osteoglossoidei).TEb2 transversus epibranchialis 2. Muscle may ap-pear to comprise two more-or-less fused segments,giving impression of twisting (see especially Go-biidae) as they pass between levators (usually LI1and LI2) to insert on Eb2.Borden ( 1999) differentiated a muscle he termedTD2 from another he termed OD2 in Naso (Acan-thuridae) on the basis that TD lacks a mid-line ra-phe and OD2 has one. Although usually present,the presence or absence of a raphe and its extentwhen present are highly variable, and we recognizea single muscle, TEb2, for Borden's TD2 andOD2. See also M. TEb2-Pb2.TEb2a
?
transversus epibranchialis 2 anterioris; inpre-acanthomorphs only in Maurolicus (Stomiifor-mes), variously in acanthomorphs (e.g., Pseuda-pocryptes, Gobiidae: labroids).TEb2p transversus epibranchialis 2 posterioris; inpre-acanthomorphs only in Maurolicus (Stomiifor-mes), variously in acanthomorphs (e.g., Pseuda-pocryptes, Gobiidae; Dicrolene, Ophidiidae).TEb2v transversus epibranchialis 2 ventralis; onlyin Diplophos (Stomiiformes).TEb2-Ebl transversus epibranchialis 2-epibran-chialis 7; only in Beiy.x (Berycidae), not to be con-fused with TEbl-Eb2.TEb3 transversus epibranchialis 3; present only inacanthomorphs. Except for the pre-acanthomorphengraulid genus Coilia, attachment of TD to Eb3alone or together with another skeletal element, isrestricted to Acanthomorphs, and is a synapomor-phy of the group.TEb3-Eb4 transversus epibranchialis 3?epibran-chialis 4.TEb4 transversus epibranchialis 4. May be discreteor continuous anteriorly and/or posteriorly withother muscles.TPbl-2-3-Ebl-2
?
transversus pharyngobranchialis1 ,2, 3'-epibranchialis 1,2; only in Diplomystes (Di-plomystidae).TPb2 transversus pharyngobranchialis 2. In pre-acanthomorphs and primitive acanthomorphs, thisis frequently a band-like muscle anterior to TEb2,if latter is present, and usually attaches to bothPb2s. In acanthomorphs, beginning with paracan-
14 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
thops and zeoids, TPb2 lies partly or entirely dor-
sal to TEb2 and fuses partly or entirely ventrallywith TEb2, and, except for its loss or the loss ofTEb2, rarely if ever has any other topographicalposition. TPb2 may also be pad-like or consist ofa bi-lateral muscle pair usually joined by CT; maycomprise a medially open semicircular ribbon ofmuscle on each side, which may join only TEb2antero- and posteromedially, or may attach ante-riorly only to IAC, Pb2, or Pb3. Occasionally, itmay be present only unilaterally and vestigially asa semicircular ribbon. Recognition of TPb2, whennot attached to Pb2 (e.g., Rhamphocottus, Moro-
ne), is based on the configuration of the muscle,which appears the same in other taxa in which itattaches to Pb2. For these reasons, we ignored thepresence of an attachment to Pb2 as the definingfactor in the identification of TPb2.TPb2 deserves more study than we were able todevote to it, and our designations of the variousstates of the muscle as TPb2 or some variety of it,possibly obfuscates its various homologies.In most acanthomorphs, our TPb2 is the samemuscle Anker ( 1978) designated as the "m. cranio-pharyngobranchialis 2," which designation hasbeen followed by most recent authors, and espe-cially Stiassny and Jensen (1987). The muscle thatthe latter authors treat as the transversus pharyn-gobranchialis 2, we nominate as TPb2a, as it ap-pears to have a different history from our TPb2.TPb2'
?
transversus pharyngobranchialis 2 primus; aseparate dorsoanterior portion of TPb2 (e.g., Ho-plostethus, Trachichthyidae) or separate muscle be-tween TPb2 and TPb3 (e.g., Tylosurus, Belonidae).TPb2-Pb2a
?
transversus pharyngobranchialis 2-phaiyngobranchialis 2 anterioris. Putative fusionof TPb2 and TPb2a (only in Labroides, Labridae,but see TPb2-Pb2a-Pb3).TPb2a transversus pharyngobranchialis anterioris(see also CPb and discussion of TPb2 above).Most prominently present in labroids, atherino-morphs, Pholidichthys.TPb2d transversus pharyngobranchialis dorsalis.Name applied to muscle of variable structure dor-sal to and continuous with TPb2 or TPb2v in afew acanthomorphs (e.g., Aphredoderus, Aphre-doderidae; Agonostomus, Mugilidae). In Agonos-tomus, it is flat, pad-like and attaches to IAC andis, perhaps, also represented by anterior portion ofTPb2 in moronids and mullids.TPb2p transversus pharyngobranchialis posterior-is; a posterior separation of TPb2.TPb2v transversus pharyngobranchialis ventralis;muscle joining ventral surfaces of Pb2s (e.g., Hem-iramphidae). The designation might be applied toTPb2a of Pholidichthys, but it appears that Pb2 hasrotated so that its anterior surface lies ventral; the
muscle is more similar to TPb2a of atherino-morphs than it is to TPb2v of atherinomorphs.TPb2-Pb2a-Pb3
?
Transversus pharyngobranchialis2-pharyngobranchialis 2 anterioris-pharyngo-branchialis 3. Compound muscle comprising a fu-sion of TPb2, TPb2a, and TPb3 (only in Sympho-dus, Labridae).TPb2-Pb3a
?
transversus pharyngobranchialis 2-pharyngobranchialis 3 anterioris; present only insome aulopiforms.TPb2-Pb3-Eb4
?
transversus pharyngobranchialis 2-pharyngobranchialis 3-epibranchialis 4; only inGymnarchus (Osteoglossomorpha).TPb2-Pb3-Pb4-Eb4
?
transversus pharyngobran-chialis 2-pharyngobranchialis 3-epibranchialis 4;only in Lovettia (Galaxiidae).TPb3
?
transversus pharyngobranchialis 3; common-ly present in a large variety of fishes.TPb3-Eb3
?
transversus phaiyngobranchialis 3-epi-branchialis 3; only and commonly present in acan-thomorphs (see also TEb3).TPb3-Eb3-EM transversus pharyngobranchialis 3-epibranchialis 3-epibranchialis 4; only in acantho-morphs (see also TEb3).TPb3-Eb4 transversus pharyngobranchialis 3-epi-branchialis 4.TPb3-Eb4-UP4-UP5 transversus pharyngobran-chialis 3-epibranchialis 4-laminalis dentalis 4-laminalis dentalis 5; only in Anguilla (Anguilli-dae).TPb3a
?
transversus pharyngobranchialis 3 anterior-is; only in pre-acanthomorphs.TPb3a-Eb2 transversus pharyngobranchialis 3 an-terioris-epibranchialis 2; only in a few pre-acan-thomorphs.TPb3p transversus phaiyngobranchialis 3 poster-ioris, only in pre-acanthomorphs.TPb3p-Pb4 transversus phaiyngobranchialis 3 pos-terioris-pharyngobranchialis 4; only in a few pre-acanthomorphs.TPb3-Pb4-Eb3 transversus pharyngobranchialis 3-pharyngobranchialis 4-epibranchialis 3; only inacanthomorphs.TPb3p-Pb4-Eb3-Eb4 transversus pharyngobran-chialis 3 posterioris-pharyngobranchialis 4-epi-branchialis 3-epibranchialis 4; only in veliferidsand some girellids.TPb3' transversus phaiyngobranchialis 3 primus;posteriormost of three TPb3s (after TPb3a,TPb3p); only in Aulopus (Aulopidae).TPb3-Pb4 transversus pharyngobranchialis 3-pharyngobranchialis 4; in various pre-acantho-morphs, but only in Bovichtus (Bovichtidae)among the acanthomorphs.TPb3-Pb4-Eb4 transversus pharyngobranchialis 3-pharyngobranchialis 4-epibranchialis 4; present
NUMBER 11 15
only in a few acanthomorphs, Psenopsis (Centro-lophidae), Tetracentrum (Ambassidae).TPb3p-Pb4-Eb4
?
transversus pharyngobfanchialis 3posterioris-phaiyngobranchialis 4-epibranchialis4; only in pre-acanthomorphs, e.g., Searsia (Pla-tytroctidae).TPb3-UP3-UP4 transversus phaiyngobranchialis3-laminaIis dentalis 3-laminalis dentalis 4\ only inthe eel Synaphobranchus (Synaphobranchidae).TPb3-UP4
?
transversus pharyngobranchialis 3-lam-inalis dentalis 4; present Xenocephalus (Uranos-copidae) and some gobioids.TPb3p-UP4?present only in Albula (Albulidae).TPb3-UP4-Eb4 transversus pharyngobranchialis3-laminalis dentalis 4-epibranchialis 4: presentonly in one species of Channa (Channidae).TPb4
?
transversus phaiyngobranchialis 4; presentin a variety of pre-acanthomorphs, but only inPseudaphritis (Pseudaphritidae) among the acatho-morphs.TPb4a
?
transversus phaiyngobranchialis 4 anterior-is; an almost completely separate anterior sectionof TPb4.TPb4-Eb3 transversus phaiyngobranchialis 4-epi-branchialis 3; only in Coilia (Clupeoidea) andNemipterus (Nemipteridae).TPb4-Eb4 transversus pharyngobranchialis 4-epi-branchialis 4\ present only in pre-acanthomorphs.TUP4 transversus lamina!is phaiyngobranchialis4; only in gobioids Pseudapocryptes and Ptereleo-tris.TUP4a transversus laminalis dentalis 4 anterioris;only in Diplomystes (Diplomystidae).TUP4p transversus laminalis dentalis 4 posterioris:only in Diplomystes (Diplomystidae).TUP5 transversus laminalis dentalis 5; muscle con-necting UP5s; only in Stomiiformes.TL?total length.TV4 transversus ventralis 4 (Fig. 1); ventral gill-arch muscle connecting Cb4 on one side to Cb4on the other; dorsally interrupted in some acantho-morphs (e.g., labroids) and attaching also to Cb5.TV5 transversus ventralis 5; ventral gill-arch mus-cle connecting Cb5 on one side to Cb5 on the oth-
er; occasionally illustrated, but not described.Uncinate process (pertaining to Ebsl-4; occasionallyrefers also to Pb2, which may have a distinct pro-cess that articulates directly or indirectly with Eblor Pb3)?defined for our purposes as a cartilage-tipped process. Hence, if the cartilage tip is absentthe process is considered absent; however, with re-gard to Ebl, Eb3, and Eb4, if the bony support ispresent but the cartilage tip is absent, the uncinateprocess is frequently described as "all bony," butnot differentiated from absent in the cladistic anal-ysis (Appendix).In acanthomorphs an uncinate process is fre-
quently present on Ebl, usually well medial to thelateral end of the bone, and in percomorphs it oftenarticulates with the lateral end of IAC. A cartilage-tipped uncinate process that articulates with Pb3may also be present on Eb2, but it appears to berestricted mainly to some pre-acanthomorphs.Among the Acanthomorpha, an Eb2 uncinate pro-cess has evolved independently in Hemiramphus(Hemiramphidae. in which the process does notarticulate with another skeletal element), and, var-iably, in Pholidichthys (Pholidichthyidae). inwhich it articulates with Pb2. Rosen and Patterson(1990:figs. 49c, 50c), probably erroneously, illus-trated a posteriorly directed Eb2 uncinate processin the acanthomorph genera Peprilus (Stromatei-dae) and Trichiurus (Trichiuridae). We have ex-amined a cleared and stained T. lepturus and findthat it has a bony process on Eb2. to which LE2probably inserts, the usual condition in acantho-morphs. Many cichlids have a separate, expandedcartilaginous process on the anteromedial edge ofEb2 which extends anteriorly ventral to Ebl, andwhich we do not consider to be an uncinate pro-cess.An uncinate process on Eb3 appears first in os-teoglossomorphs (Hiodon) followed by one onEb4 in elopiforms (Megalops). Tightly juxtaposeduncinate processes on Eb3 and Eb4 are first presentin some aulopiforms. but appear to be an acantho-morph synapomorphy, with occasional reversions.The acanthomorph Eb4 levator process is usu-
ally lateral or posterior to and well separated fromthe uncinate process. LE4 may insert directly onthe levator process or close to it. When only oneprocess, uncinate or levator, is present on Eb4,there is a problem in deciding which it is. If theprocess is tightly joined to the Eb3 uncinate pro-cess, there is usually little question that it is anuncinate process. If the process is well separatedfrom the Eb3 uncinate process, it is usually notpossible to decide, unless LE4 or LP inserts on it,in which case it almost certainly is the levator pro-cess (among acanthomorphs, LE4 inserts on theuncinate process only in a few lampridiforms).The uncinate process is absent in nemipterids,and "in Lethrinus and Gnathodentex [both Leth-rinidae] it . . . has become closely associated withthe levator process . . . and has lost a close asso-ciation with the uncinate process of the third epi-branchial" (Carpenter and Johnson, 2002:120).Early ontogenetic stages, therefore, may also offera solution to the problem of identification of a car-tilage-tipped process on Eb4 when only one suchprocess is present.UP?upper pharyngeal tooth plate, e.g., UP4, UP5.
16 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
o6 <p <*? \*?cr ><y <y &<r v? o* o?Mr!SarcopterygiiTeleostomi or Osteichthyes
I Gnathostomata
Fig. 2. Cladogram of major groups of extant Gnathostomatanfishes (derived essentially from Nelson, 1994:inside front cover)treated in the present study.
ClassificationThe classifications of fishes followed in our studyare given in Figs. 2-4. For the most part, these arecompilations of current hypotheses of relationshipsfrom which the fossil taxa have been removed. Thecladogram in Fig. 2 presents the entire higher clas-sification of taxa that include organisms generallycalled fishes. Fig. 3 illustrates the classification of theActinopterygii we examined, but provides detailedbranching only for the pre-Acanthomorpha. Theacanthomorph classification (Fig. 4) details the majorprimitive clades, but generalizes the terminal clade,Percomorpha; see also Footnote 1, page 18).The following discussion briefly examines the lit-erature or other information on which the classifica-tions are based.The classification of the recent Gnathostomata inFig. 2 derives essentially from Nelson (1994), and isgenerally accepted in some form (usually with moredetail) in most, if not all, current general classifica-tions of fishes (e.g., Long 1995:27, Maisey 1996:11,Gill and Mooi 2002:19).Pre-acanthomorph classification.?The five bas-almost clades of the Actinopterygii (Cladistia, Actin-opteri, Neopterygii, Halecostomi, and Teleostei) andtheir inter-relationships that we follow were first hy-pothesized by Patterson (1982:253; N.B., most recentauthors overlook this reference, in which Pattersonfirst included the Cladistia in the Actinopterygii, andplaced it as the basalmost clade). Patterson (1994),who did not cite his 1982 study, concluded that thefossil, anatomical (extant taxa), and molecular evi-dence supporting monophyly of the Actinopteri con-flicts, and that the Actinopteri must be considered tocomprise a polytomy of Ginglimodi, Halecomorphi,and Teleostei. The anatomical evidence (Patterson
1994:68) based on extant taxa, however, supportedhis earlier (1982) hypothesized relationships: (Ging-limodi (Halecomorphi, Teleostei)), which we employ.Gardiner et al. (1996) reported a similar conflict, withmorphological characters (including those of fossils)indicating that Ginglimodi and Halecomorphi areparaphyletic and molecular characters indicating thatthey are monophyletic.There are two main opposing classifications for thebasal clades of the Teleostei. Patterson and Rosen(1977) hypothesized the arrangement that we use inour cladogram. Arratia (1999) provided a radicallydifferent set of inter-relationships. Arratia based herhypothesis on 14 extant and 34 fossil taxa, using 196characters, of which only three (her character num-bers 165, 167, 190) can be considered as non-oste-ological. Character number 165, presence of adiposefin, appears only as a synapomorphy for two recentsalmonids; 167, coiling direction of intestine, appearsonly as a synapomorphy for two recent osteoglos-somorphs; and 190, presence of nasal sacs, supportsmonophyly of 12 of the 14 extant taxa and 12 of the14 fossil taxa she examined (excludes Amia and Lep-isosteus) and also supports monophyly of Esox andUmbra, the only two esociforms she examined. Themajor difference between Arratia's classification andthat in our Fig. 3, is that her Elops and Megalopsreplace the Osteoglossomorpha as the sister group of
all other teleosts, and the osteoglossomorphs aremade the sister group of the remaining teleosts. (Forpossible evidence of a closer relationship of Osteo-glossomorpha to Elopomorpha, see "bilaterallypaired Pb muscles" in Results section of pre-acan-thomorphs. For possible evidence for support of acloser relationship of Osteoglossomorpha to Clupeo-morpha see discussion of Eb5 and Eb4 in "Musclesand Skeletal Elements.")The classification of the Osteoglossomorpha is ex-tracted from Taverne (1998:figs. 21-22; timing pre-cluded accommodation of Hilton's (2003) rearrange-ment of the taxa in our Osteoglossiformes). The Elo-pomorpha is slightly modified from Forey et al.(1996:fig. 2) to make the Anguilliformes a polytomy.Forey et al. studied only the Anguillidae of the threeanguilliform families we include. There is little sup-port available for the sister group of the Elopiformes,comprising the Albuliformes, Notacanthiformes, andAnguilliformes. Nelson (1973:347) proposed An-guillomorpha for this group (noted as a new super-order only in the combined index of Greenwood et
al., 1973:520).The branchings of Clupeocephala into Otocephalaand Euteleostei, of Otocephala into Ostariophysi andClupeomorpha, of Euteleostei into Neognathi andProtacanthopterygii, of Neognathi into Neoteleosteiand Esociformes, and all the branches of the Prota-canthopterygii (Argentiformes, Salmoniformes, Os-
NUMBER 11 17
Ctenosquamata
Eurypterygii
Neoteleostei
Neognathi
Euteleostei
Myctophiformes{?Aulopiformes Acanthomorpha, * ~r-MyctophidaeLampanyctus -1NeoscopelusCliloroplitlialmiisSaurida
Ateleopoditbrmes
Stomiifomies
Esocifbrmes rCE
Argentiniformes
Prolacanthopterygii
Qupeocepliala
Elopocephala
Teleostei
Halecostomi
Neopterygii
Actinopteri
Actinopterygii
Osmeroidea
Osmeroidei
Salmoniformes GalaxioideaSalmonoidei
-cGvmnotifbrmes
Otophysi
Ostariophysi
Otocephala
r~ Siluriformes
Characiformes
Cypriniformes
Gonorvnchiforrnes
Clupeoidea
Qupeoidei
Clupeomorpha
Engrauloidea
Pristigasteroidea
Denticipitoidei
AulopusAteleopus
-Gonostoim
- MaiirolicusDiplopias
-Umbra
-Dallia
- No\w?ibraEsox
- Searsia
- Alepoceplialus
- Argentina
- Mailotus
- Hypomesus
- Lovettia
- Lepidogalaxias
- Galaxias
- Retropimm
- Coregonus
- Thymallus
- Oncorhynchus
- Gynvtotus
- Diplomystes
- Xenocliarax
- Brycon
-Zacco
- Opsariichthys
- Gonorynchits
- Otanos
- Qupea
- Dussumieria
- Oiirocentnts
- Coilia
- Cetengraidis
-Illislia
- Denticeps
3-
NeoscopelidaeChlorophthalmidaeSynodontidaeAulopidaeAteleopodidaeGonostomatidaeSternoptychidaeDiplophidae
Umbridae
EsocidaePlatytroctidaeAlepocephalidaeAreentinidae
^-Osmeridae
-Galaxiidae
3-
Anguillomorpha
Elopomorpha
I? SynaphobranclutsAngiulla
Retropinnidae
Salmonidae
GymnotidaeDiplomystidaeDistichodontidaeCharacidae
"H-Cyprinidae
GonorynchidaeQianidae
^-ClupeidaeChirocentridae
^-Engraulidae
PristigasteridaeDenticipitidaeSynaphobranchidae
Anguilliformes Lq?^Notacanthiformes i? AldrovandiacAlbuliformes
Elopiformes
Osteoglossoidei
Osteoglossiformes
Osteoglossomorpha Notopteroidei
Hiodonliformes r^Halecomorphi
Ginglymodi-l-Chondrostei
Cladistia-
NotacantlmsPterothrissusAlbidaElopsMegalopsScleropagesOsteoglossiimPantodonArapaimaHeterotisMormyrusPetrocepltahtsGymnarchusNotopterusHiodonAmiaAtractosteiisPolyodonAcipenserPolypterus
AnguillidaeCongridaeHalosauridaeNotacanthidae
~]-Albulidae
ElopidaeMegalopidae
^-Osteoglossidae
Panlodontidae
^-Arapaimidae
^]-MormyridaeGymnarchidaeNotopteridaeHiodontidaeAmiidaeLepisosteidaePolyodontidaeAcipenseridaePolypteridae
Fig. 3.study. Cladogram of the Actinopterygii (compiled from literature) with particular reference to pre-Acanthomorpha included in present
18 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
meroidei, Salmonoidei, Osmerioidea, Galaxioidea)are from Johnson and Patterson (1996:315-316). Thebranching of the Esociformes is taken from Nelson(1972:25) and Wilson and Veilleux (1982) and sup-ported by Johnson and Patterson (1996:314) and San-ford (2000:214, 218, 219). Lopez et al. (2000), basedon molecular evidence, however, arrived at a differ-ent set of relationships: (Umbra (Dallia (Esox, No-vumbra))).The branching of the Clupeomorpha. Greenwood(1968) hypothesized Denticipitoidei as the sistergroup of the Clupeoidei (= all other Clupeomorpha),and its position has persisted to the present. The un-resolved trichotomy of the Clupeoidei (Clupeoidea,Engrauloidea, Pristigasteroidea) was last hypothe-sized by Grande (1985). The branching of the Ostar-iophysi was hypothesized by Fink and Fink (1996:210-211).With the exception of the monofamilial Ateleo-podiformes, the branching of Neoteleostei to Acan-thomorpha was first proposed by Rosen (1973). Ro-sen included the ateleopodiforms as acanfhomorphs,but Olney et al. (1993:155) demonstrated they are notacanthomorphs and placed them as an unresolved tri-chotomy with Eurypterygii and Stomiiformes. Thestomiiform branches were hypothesized by Harold(1998:fig. 4). The aulopiform branches were hypoth-esized by Baldwin and Johnson (1996:359).Acanthomorpha classification.?The general ar-rangement of the higher acanthomorph groups is thatof Johnson and Patterson (1993:fig. 24; modifiedslightly in our Fig. 4). We arbitrarily recognize manygroups of families included in their Percomorpha asseparate branches (with unresolved interrelation-ships) of a polytomous Percomorpha bush, which in-cludes a polyphyletic Perciformes and polytomousSmegmamorpha as one of its several branches. Cur-
rently, there is no basis for separating Percomorphafrom Perciformes. 1
1 Johnson and Patterson (1993:591-592), in an attempt to con-serve the names Percomorpha and Perciformes, defined a mono-phyletic Percomorpha as a clade including, among other groups,the Perciformes. They illustrated their conclusions in their fig. 1 lb(polychotomous smegmamorphs as sister group of Perciformes).fig. 18 right (polychotomous Smegmamorpha as sister group of apolychotomy comprising Dactylopteriformes, Scorpaeniformes,Perciformes. Pleuronectiformes, Tetraodontiformes), and fig. 24(monophyletic Percomorpha comprising two clades, Smegmamor-pha and "Perciformes, etc." To summarize in their own words . . .we know ofno sound characters justifying a pre-perciform positionfor Scorpaeniformes, and this emphasizes the tenuity of the dis-tinction between our Smegmamorpha . . . and the Perciformes ....This raises the embarrassing or mortifying possibility that weshould wind up a volume on percomorph phylogeny by concludingthat the group does not exist. The alternative is to save the Per-comorpha by expanding it to include fishes that were originallyexcluded from it. the atherinomorphs. We believe that there is amonophyletic group comprising "perciforms and their immediaterelatives" and our smegmamorphs. That group can be character-
Gnathostomata(= Chondrichthyes + Telostomi)Telostomi(= Sarcopterygii + Actinopterygii)Remarks. Gill-arch levators are absent in Chon-drichthyes (Edgeworth, 1935:140, = his Elasmobran-
chii; Greenwood and Lauder, 1981:222), but are pres-ent in basal Sarcopterygii and Actinopterygii; there-fore, the presence of levators is a synapomorphy ofthe Teleostomi. Sarcopterygii(= Coelacanthidae + Dipnoi)COELACANTHIDAELatimeria chalumnae Smith.Remarks. Information on Latimeria is based hereprimarily on that reported by Millot and Anthony(1958), translated a bit freely from the originalFrench. Muscle-bone abbreviations and remarks areours.
Description.LEI originates as fibers on apex of the processuspost-oticus, near point of articulation of Pbl, and[continues] along posterior three-fourths of exteriorborder of Pb 1
.
Remarks. Nelson (1968b:fig. 5A) interpreted Mil-lot and Anthony's Pbl as a probable fusion of Pbland SPbl, but he (1969b:487 and fig. 1) later indi-cated it as "PHI" (= Pbl), noting that the Pbs ofLatimeria "are difficult to compare with those of oth-er fishes." Evidence possibly supporting the elementas either Pbl or Pbl+SPbl is that LEI attaches toPbl in the basalmost actinopterygian, Polypterus,and to SPbl in the relatively plesiomorphic Acipen-
ser, Chondrostei. Unfortunately, the basalmost extantdipnoan, Neoceratodus. lacks both Pbs and SPbs.Nelson (1968b:fig. 5a, 1969b:fig. 1) interpreted Mil-lot and Anthony's Pb2 as SPb2, their Eb2 as Pb2,and their Eb4+5 as Eb4.LE2, LE3, LE4 fuse at their cranial origin, whichoccupies the posterior surface of the processus post-oticus and extends to inferior border of the supratem-poral. LE2 insertion begins on posterior extremity ofPb2 [= Nelson's SPb2; see remarks following de-scription of LEI]. It soon divides and ends cappingthe posterior epiphysis of Cb2. LE3 and LE4 are unit-
ized by a series of apomorphies . . . and we propose to name itPercomorpha. Johnson and Patterson, could not offer support fora monophyletic "Perciformes, etc." and their Smegmamorphashould have been treated as just another "immediate relative" ofthe "Perciformes etc." polychotomy. Nelson (1994:253) failed tonote this discrepancy when he essentially reproduced Johnson andPatterson's fig. 18 right, as well as taking unexplained, althoughprobably correct, issue with the monophyly of the Smegmamorpha.
NUMBER 1
1
19///Jo j$> 4T Ac
*<& sP & aF c*f x? c# A?wPERCOMORPHAEUACANTHOPTERYGIIUNNAMED CLADEACANTHOPTERYGIIHOLACANTHOPTERYGII
EUACANTHOPTERYGIIACANTHOMORPHA
Fig. 4. Cladogram of major groups of Acanthomorpha (modified from Johnson & Patlerson ( 1993:fig. 24).
ed as a bulky fleshy body obliquely inclined poster-oventrally. Near their movable insertions, the fibersdivide and extend separately to posterior epiphysesof Cb3 and Cb4.LE5 is noticeable for its restricted mass and at-tachments. It originates as three weak tendons on ex-ternal surface of anocleithrum. The three tendonscontinue as brushlike muscles, fusing into a small,transversely flat muscle, which extends anteroven-trally toward the posterior end of Cb5, to which it isattached only by very poorly differentiated fibroustissue.Remarks. The origin of this muscle on a cleithralelement and its insertion on Cb5 suggest that it isprobably homologous with PP (which Millot and An-thony do not mention) in Dipnoi and Actinopterygii.
It does not appear to be homologous with LE5 ofNeoceratodus (Dipnoi), which originates on the cra-nium and inserts in the dorsal esophageal CT justposterior to the gill arches.Millot et al. (1978) expanded on the anatomy ofLatimeria. They (1978:fig. 7; plate 22) provided adrawing and photograph of a cross-section throughthe gill arches. They did not mention the presence ofany muscles, but clearly, there were no M. Intrbspresent in the illustrations. The absence of M. Intrbsin Latimeria is an indication that their absences in
Actinopterygii and Coelacanthidae represent inde-pendent losses (homoplasies).Ads absent.SO composition unrecorded.Remarks. Millot and Anthony (1958) do not men-tion which SO muscle layers are present, but Millot
et al. (1978:plate 4), without comment about the mus-cle layers, provided a cross section of the esopha-gous, which indicates only a single (transverse or cir-cular) muscle layer.RDs absent.
Dipnoi
Remarks. Only the generally considered least spe-cialized family, Ceratodontidae, is discussed below.CERATODONTIDAE
Neoceratodus forsteri (Krefft), AMS 1.40438001,200 mm TL. Plates 1.1, 1.2
Description.LEI mainly on dorsoposterolateralmost surface ofCbl with very minor tendinous attachment extendingfrom ventromedialmost edge of muscle to Ebl dor-soposterolaterally. See remarks following LE4.
20 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LE2 on posterolateral edge of Eb2-Cb2 joint. Seeremarks following LE4.LE3 originates with LE4 and M. Intrbr3 on cra-nium; inserts on posterolateral edge of Eb3-Cb3 joint.See remarks following LE4.LE4 originates with LE3 and M. Intrb3 on crani-um; inserts on posterolateral edge of Eb4-Cb4 joint.Remarks. LE1-4 are all relatively short and, ac-cording to Edgeworth (1935:129) and Fox (1965:490), who studied larvae, originate separately on theventral surface of the auditory capsule. Greenwoodand Lauder (1981.-fig. 2) illustrated LE3 and LE4originating from a common stalk. Fox (1965:fig. 8),based on a 34.5 mm larva (his largest specimen),illustrated the first three levators as originating sep-arately from the others, and the fourth as originatingwith a muscle he recognized as the 6th levator, whichwe believe is PP.Fiirbringer (1904:489) described the insertions ofthe four anteriormost levators as follows (translatedfrom German, substituting our muscle terminology):LEI on Cbl and posterior end of Ebl. LE2 withapproximately equal parts on Cb2 and posterior bor-der of Eb2. LE3 and LE4 insertions reduced to aminimum on [their respective] Cbs; insertions are al-most exclusively on posterior border of [their respec-tive] Ebs. Edgeworth (1935:129) described them asinserting "into the dorsal ends of the lower segmentsof the [branchial] bars," and Fox (1965:490), essen-tially agreed, "insert on the tops of ceratobranchiale1-4, respectively, laterally to the epibranchiale."LE5 originates on the ventrolateral surface of theauditory capsule posterior to the origin of LE4, and,superficially, appears to be an anterior continuationof the PP origin. LE5 has a distinct, moderately long,tendinous stem that inserts into the non-musculousesophageal tissue, where that tissue first constrictsposterior to the branchial arches, and at the dorsoan-teriormost edge of the musculous SO. At the LE5insertion. SO is continuous only around the ventralsurface of the esophageal tissue. LE5 can be consid-ered to be closely associated with Cb5 only by a fewfine, tendinous strands extending from the tendinousstem of the right-side LE5 to the PP in the vicinityof the posterior end of Cb5.Remarks. Fiirbringer (1904) and Edgeworth (1935)recognized only four LEs in Neoceratodus, whereasFox (1965) recognized six. We are uncertain as to theexact homologies of Fox's LE5 and LE6. Fox (1965:490) stated that, "Levators 5 and 6, which behindmerge ventrally with the coracobranchialis and pos-terior transversus musculature, may include a portionhomologous with the dilator laryngeus and the be-ginning of the cucullaris muscle [= our PP] . . ." Fox(1965:fig 7) indicated the presence of only LE5 inhis larval 27 mm specimen, in which it is quite long.Dorsally it closely approaches or attaches to the pos-
terior surface of the auditory capsule; anteriorly atmid-length, it closely approaches or attaches to theposterior (or distal) end of Cb5; posteroventrally it isfree, but clearly directed toward the clavicle or cleith-rum. In a 20 mm specimen. Edgeworth (1935:figs.43 and 43a) labels this muscle constrictor branchialis5 and shows it extending toward the clavicle and im-pinging on, or fusing with, coracobranchialis 5 pos-teriorly before attaching to the clavicle. In Fox's next,and last, stage larva, 34.5 mm, LE5 is no longer incontact with the cranium, having lost much of itsdorsal extent, but maintaining its mid-length attach-ment to the distal end of Cb5. It lies along (fuseswith?) the anterior or anterolateral surface of a verylarge LE6, which makes its first appearance andmore-or-less duplicates the relative length and posi-tion of LE5 in Fox's 27 mm larva. As Fox suggested,LE6 probably represents an early stage in the for-mation of the PP, but unless LE5 loses its identityand fuses with PP in specimens larger than our 200mm specimen, we do not believe it is part of PP.PP originates on ventrolateral surface of the au-ditory capsule beginning musculously as an apparent,narrow, posteriorly continuous extension of the LE5origin, but the origin becomes tendinous posteriorly;PP rapidly expands, becoming fan-like as it extendsventrally, attaching anteriorly along posterior (or lat-
eral) border of Cb5, and reaching ventrally to thesubarcualis rectus on Cb5 (Edgeworth, 1935:fig. 43b;Wiley, 1979:fig. 3c) and coracobranchialis 4 on Cb5(Edgeworth, 1935:fig. 43b); posteroventrally, it in-serts broadly on clavicle with and posterior to thecoracobranchiales. (See remarks following LE5.)M. Intrbl-4 origins are about same size as thoseof LEs. Intrbl and 2 originate on cranium near ori-gins of LEI and 2; Intrb3 originates together withcombined origins of LE3 and 4; Intrb4 originatesposterior to the previous. Ventral to their origins, thefibers of each M. Intrb gradually separate in a singleplane on the branchial septum, ultimately branchinginto about 8 filaments, each of which is separatedfrom an adjacent filament by a narrow space; eachmuscle arches posteriorly, then ventrally; the fila-ments re-unite anteroventrally and attenuate, finallycontinuing anteriorly as a long, fine tendon, whichinserts in CT near the anteroventral end of each mus-
cle's respective Cb.Remarks. M. Intrbs are known only in lungfish andelasmobranchs; thus constituting a possible synapo-morphy contra-indicating a sister-group relationshipbetween lungfish and sarcopterygians.PrO levator-like. oriented almost horizontally,originating on ventral surface of cranium a little me-dial to medial end of Eb4, extending posteromedially,and inserting on non-musculous esophageal tissueposterior to branchial arches.Remarks. We have not found this muscle men-
NUMBER 1
1
21
tioned in previous studies; it is tempting to interpretit as another form of RD, which otherwise makes itsfirst appearance in basal Neopterygii.SO consists only of transverse muscle layer.Ads absent.RDs absent (but see remarks following PrO).Additional remarks. Huxley (1876:27) illustratedthe gill arches of Neoceratodus. On page 37 he de-scribed them, recognizing anterior and posterior un-paired "mesobranchials" and another cartilage:
"close to the ventral end of the fifth arch [= Cb5],was a small nodule of cartilage, which is probably arudimentary sixth arch . . ." In our specimen, wefound the anterior mesobranchial, which lies mediallybetween the ventral ends of Cb2 and Cb3 on each
side. Huxley illustrated its position as extending an-teriorly from the ventral end of Cb2 to a point an-terior to the ventral end of Cbl. His mesobranchial1 is undoubtedly a basibranchial.On only the right side of our specimen there is anunpaired cartilage between the ventral ends of Cb3and Cb4. On the left side of our specimen, the ventralend of Cb5 appears to have become deeply notchedsub-terminally, but continuous posteriorly with theremainder of the Cb. A small autogenous plug ofcartilage fills the gap between the two continuousparts. It appears that ventral fragmentation of the Cbsmay be common in Neoceratodus.
ActinopterygiiPre-Acanthomorpha(Cladistia?Myctophiformes)CladistiaPOLYPTERIDAE
Polypterus ornatipinnis Boulenger, USNM 164514,170 mm TL. Plate 2
Description.LEI on dorsoposterior edge of Pbl and lateral por-tion of mostly cartilaginous Ebl (small ossificationcenter present in each Ebl).Remarks. The only actinopterygians in which a le-vator inserts in whole or in part on Pbl are Polyp-terus and some osteoglossiforms (Petrocephalus, Os-teoglossum, Scleropages).LE2 on dorsolateral surface of cartilaginous Eb2.LE3 on dorsal surface of cartilaginous Eb3.LE4 ventromedially on CT just lateral to SO, ven-trolaterally on cartilaginous distal end of Cb4 andgreatly reduced cartilage, which is here designatedEb4, and ventroanteriorly on distal end of Eb3 (seeadditional remarks at end of description).LP absent.LI 1?3 absent.
TD absent; SO joined dorsally to gill arches byCT, possibly resulting from loss or extreme reductionof Eb4. (TD is also absent in Dipnoi and probablycoelacanths.)OD3 and OD4 absent (OD first appears in Neop-terygii, Ginglymodi).OP absent, possibly concomitant with loss of Cb5.Ad 1-3 and Ad5 absent.Remarks. If it was present in early polypterid phy-logeny, Ad5, which normally attaches Cb5 to thefourth arch, was probably lost with the loss of Cb5(see also additional remarks).Ad4 is questionably represented by muscle joiningdistal end of Cb4 and reduced Eb4 with distal end ofEb3; muscle is adjacent to and questionably contin-uous with LE4.Remarks. Ad4 normally attaches Eb4 to Cb4 inother actinopterygians, but with the considerable re-duction of Eb4, Ad4 may have shifted its dorsal at-tachment to Eb3. The presence of Ads, which areabsent in Chondrichthyes and Sarcopterygii (Dipnoi+ Coelacanthidae), is a synapomorphy of the Actin-opterygii.RD absent.SO longitudinal muscle layer absent.Remarks. SO comprises a single (circular or trans-verse) muscle layer. SO extends anterodorsally onlyto a dorsally projected horizontal joining the distalends of Cb4s. As noted by Edgeworth (1935:167),SO in Polypterus "does not attach to any branchialbars," and as such, is unique among the Actinopter-ygii. Similar conditions pertaining to SO exist in theprimitive dipnoan, Neoceratodus, in which, however,SO begins relatively even more posteriorly.Additional remarks. SCL absent, TV4 absent.Prominent ligament, originating on parasphenoid, in-
serts on distal end of Cbl.Jollie (1984:fig. 17b) illustrated the gill arches ofPolypterus, purportedly based on Allis (1922:pl. 8,fig. 17b). Jollie, however, greatly increased the lengthof the ventral cartilaginous portion of Allis's Ebl,interpreted the cartilage as Ebl, and labeled the twobony arms extending dorsally from the cartilage as
"SPb" and "JPb." We believe Jollie's changes wereunwarranted.Polypterus is unusual in having only four gill arch-
es. Britz and Johnson (2003) discuss the two hypoth-eses concerning the homology of the missing arch(whether it is the fourth or fifth) and strongly supportthe argument that the missing arch comprises the fifthceratobranchials.Allis (1922:232 et seq.) reported that: the first gillarch comprises Pbl (which articulates with Ebl) Ebl,Cbl, and Hbl; the second arch comprises Pb2 witha fused, reduced Eb2, Cb2, and Hb2; the third archcomprises Pb3 with a fused, reduced Eb3, Cb3, andHb3; the fourth arch comprises only Cb4. We agree
22 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
with Allis on the composition of arch 1 and arch 4,considering that the vestigial Eb4 is variably absent.For arches 2 and 3, we recognize Allis's Pbs as Ebs;for arch 4, the vestigial Eb4 may be absent or easilyoverlooked, Allis gave no evidence for his assump-tion that fusion had occurred between an Eb and aPb.Allis (1922:233) mentioned that an autogenous bitof cartilage is interposed between the [first] epibran-chial and ceratobranchial of some specimens, but notothers. This cartilage was also reported by van Wijhe(1882:257 and pi. 5, fig. 7), who identified it as anepibranchial, but which we term an accessory carti-lage (AC). The AC (Plate 2A) is present in our spec-imen, and in an 85 mm SL cleared and stained spec-imen of Polypterus senegalus Cuvier (USNM229760), but not in a 132 mm SL specimen from thesame lot. The smaller of the two specimens also hasan AC on the third arch of one side. We note thataccessory cartilages are also present on the first andsecond arches of the lepisosteid Atractosteus (Plate5A).Neither Allis nor van Wijhe. noted another, small,autogenous cartilage attached to the distal end of Cb4(Plate 2B). which is present in P. omatipinnis andthe smaller, but not larger, specimen of P. senegalus.Although Britz and Johnson (2003:499) noted thepresence of this cartilage they refrained from identi-fying it. They cite literature in which the element hasbeen identified variously as Eb4 or an epipharyngo-branchial.Wiley (1979:161) discussed the homologies of thepharyngoclaviculares (PCs) in actinopterygians. Henoted that polypterids, chondrosteans, and gars,which compose the pre-halecostomes, have a singlePC and that halecostomes have PCI and PCE. Allis(1922:259) described PC in Polypterus as having asingle origin on the cleithrum, but dividing into twoparts with separate insertions, which he did not name,on Cb4, and which Wiley did not equate with thehalecostome PCI and PCE. One can infer from Wi-ley's discussion, however, that the PCs (or divisionsthereof) of the pre-halecostomes could be homolo-gous with the halecostome PCI. We assign PCp tothe more posterior insertion and PCa to the well-sep-arated more anterior insertion of PC in Polypterus.Wiley's hypothesis that [no portion of] the pre-hal-ecostome PC is homologous with PCE of halecos-tomes, is based on two considerations. First, Wiley(1979:161) credits Edgeworth (1911) with findingthat PCs are derived from the circular [our trans-verse] muscle layer of the oesophagus (among actin-opterygians, Edgeworth's work is based on the pre-halecostomes Polypterus, Acipenser, Lepisosteus, andthe halecostome Amia; Wiley accepted the presenceof PCI and PCE in Amia). Second, Wiley dissectedthe halecostome Hiodon, which has PCI and PCE,
and found that the muscle fibers of PCE are contin-uous with SO longitudinal fibers. He postulatedtherefrom, that the PCE of halecostomes could notbe homologous with the PC of pre-halecostomes, pu-tatively derived from the SO circular layer.We cannot find any indication that Edgeworth(1911), who studied the development of PCs, report-ed that they are derived from the SO circular musclelayer. In fact, Edgeworth (1911:232) concluded that
"the coraco-branchiales [= our pharyngoclavicula-
res] of Elasmobranchs, Teleostomi, and Dipnoi . . .are derived from the ventral end of one or more bran-chial myotomes, i.e., are of cranial origin." Edge-worth (1935:155) reaffirmed his earlier finding, stat-ing that in Dipnoi and Teleostomi (he modified hisearlier finding for elasmobranchs), "they are devel-oped from the outer ends of the Transversi ventralesjust after these have grown inwards from the ventralends of the branchial muscle-plates." Even so, Wi-ley's hypothesis that PCE of Hiodon (and other Te-leostei) is not homologous with PCE of Polypterusis probably viable. Pre-halecostomes lack the SO lon-gitudinal layer (the layer is a synapomorphy of hal-ecostomes, see Results section following pre-acan-thomorph section of this study), so that any musclederived from the halecostome SO longitudinal mus-cle layer cannot have a homologue in a pre-halecos-tome. ChondrosteiACIPENSERIDAE
Acipenser ruthenus Linnaeus, USNM 62372, ca. 260mm TL. Plate 3
Description.Remarks. Muscles are complex, difficult to char-acterize, generally not so discrete as we describe. Itis possible that considerable intraspecific variationoccurs; however, there was essential agreement be-tween the right and left sides of our specimen. Mar-inelli and Strenger (1973:fig. 24) present a lateralview of the gill-arch musculature of A. ruthenus thatagrees with our illustration (Plate 3B), insofar as themuscles they illustrate (they do not present a dorsalview of this musculature).LEI ventrally on Ebl, dorsoanteriorly on SPbland cranium; broad raphe on medial surface joinedby anterior end of M. SPb2-LEl.LE2 complex, comprising five parts:M. SPb2-Eb2 on SPb2 ventrolaterally and Eb2dorsally.M. SPb2L origin on cranium, insertion on SPb2dorsoanteriorly.M. SPb2Ma origin on cranium, insertion anteriorlyon mid-medial surface of SPb2.
NUMBER 11 23
M. SPb2Mp origin on cranium, insertion posteri-orly on mid-medial surface of SPb2 and raphe withanterormedial fibers of LE3.M. SPb2-LEl origin from CT on medial surfaceof LEI, insertion anteriorly on mid-medial surface ofSPb2 ventral to SPb2Ma.LE3 origin dorsally on cranium and SPb2 posteriorsurface; insertion on dorsal surface of Eb3; lateral-most fibers of left LE3 continuous with fibers ofRecD4 (aberrant?).LE4 on Eb4 lateral to and continuous with RecD4posterolateral^.LP absent.LI 1-3 absent.TD represented by TEb4, which is posteromediallycontinuous with SO.OD3 and OD4 absent.OP absent.RecD4 joins dorsolateral surfaces of Eb3 and Eb4.continuous with LE4 ventrally.Ad 1-3 absent (see remarks following Ad4).Ad4 on Eb4 dorsoposteriorly and Cb4 posteriorly,continuous ventromedially with SO.Remarks. Edgeworth (1935:131) reported thatAd 1-4 are present in A. sturio, but that only Ad4 ispresent in A. ruthenus, A. fulvescens, and Scaphir-hynchus.Ad5 dorsally on lateral process at distal end ofCb4, ventrally on Cb5 distal end.RD absent.SO comprises only transverse muscle layer.Remarks. Wiley (1976:30-32) reported that thereare two muscle layers lining the buccal cavity of Po-lyodon and Acipenser, an inner longitudinal layeroverlain by a circular layer, and that these layers areundifferentiated from the same muscle layers liningthe esophagus. We have examined both genera andfind there is no muscle dorsally in the area betweenthe gill arches. The SO in Acipenser and Polyodoncomprises a single muscle layer: circular (or trans-verse).Additional remarks. SCL absent. TV4 absent.Edgeworth (1935:129) stated that constrictor bran-chiales (also known as interbranchiales in elasmo-branchs (Daniel, 1934:105-106, 149) and lungfishes(Fiirbringer 1904:488) are present in acipenserids. He(1935:130) described these as "narrow muscle bandsexternal to the [epibranchials and ceratobranchials].In adult stages they lie in shallow grooves in the epi-and kerato-branchialia. In Acipenser their ventralends run into the outer ends of the Obliqui ventralies
i, ii and iii and Transversus ventralis iv. In Scaphir-hynchus they are overlapped by the outer end of thesemuscles." Edgeworth (1935:figs. 224b and 231) il-lustrated these putative muscles in a sagittal sectionof a 32 mm specimen of A. ruthenus and in a ventralview of the branchial muscles and arches of Sca-
phirhynchus platyrhynchus [= 5. platorynchus], sizenot given. We did not find the muscles in A. ruthenus(a second specimen, USNM 64607, 295 mm TL wasexamined for these muscles), and they were neitherreported nor illustrated by Marinelli and Strenger(1973) in their comprehensive anatomical study of A.ruthenus. We found only nerves and blood vesselscoursing along the grooves of the epi- and cerato-branchials (see also Marinellei and Strenger, 1973:fig. 240). We did not examine Scaphirhynchus.
POLYODONTIDAE
Polyodon spathula (Walbaum), USNM 101093, 217mm TL. Plate 4
Description.LEI on dorsal edge of Ebl near medial end andby tough CT on posterior edge of SPbl.LE2 on dorsal edge of Eb2 near medial end andby tough CT on posterior edge of SPb2.LE3 on Eb3 uncinate process, continuous withLE4 above insertion.LE4 on Eb4 dorsal edge somewhat distal to medialend, continuous with LE3 above insertion.LP absent.LI 1-3 absent.TD consists of TEb4, which is continuous and un-differentiated from SO.OD3 and 4 absent.OP absent.RecD4, small, anteriorly on dorsodistal edge ofEb3, posteriorly by tendon to Eb4 at insertion ofLE4.Ad 1?4, each dorsally, broadly on posterior surfaceof respective Eb, ventrally, narrowly on anterior sur-face of respective Cb just medial to inner angleformed by Eb-Cb joint.Ad5 dorsally on ventrodistalmost surface of Eb4,ventrally on dorsal surface of Cb5.Remarks. Edgeworth (1935:131) considered ourAd5 to be a 5th levator.RD absent.SO continuous with TEb4: comprises only trans-verse muscle layer (see remarks under SO in Acipen-
ser).Additional remarks. SCL absent. TV4 absent. Dan-forth (1913) described the musculature of Polyodon.Our findings are in essential agreement. He reportedthat the four levators arise from a continuous sheetand separate as they proceed toward their insertions.He worked on specimens much larger (ca. 1 m) thanthe one we describe, in which LE3 and LE4 arescarcely separate at their insertions. The differencenoted may be ontogenetic.
24 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONGinglymodiLEPISOSTEIDAE
Atractosteus tropicus Gill, USNM 120715, 415 mmTL. Plate 5
Description.LEI on dorsal surface of cartilaginous distal endof Ebl.LE2 on Eb2 (both removed when gill arches wereremoved; position in illustration is approximate).LE3 on dorsal surface of cartilaginous distal endof Eb3.Remarks. Short ligament (not illustrated) extendsposteriorly from base of LE3 and inserts on anteriormargin of cartilaginous distal end of Eb4.LE4 on dorsal surface of cartilaginous distal endof Eb4.LP absent.LI Ion dorsal bony surface of Pb2 and medial sur-face of SPbl.LI2 mostly on cartilage at ventral junction (notshown) of bony and cartilaginous portions of Pb3proximal to anteriormost Pb3 teeth.LI3 absent (Pb4 absent).TD comprises only TPb3, which is continuous an-teriorly but originates posteromedially from rapheswith SO and RDs and anastomoses ventrally with CTunderlying SO. Laterally, TPb3 attaches and sur-rounds Pb3 dorsolateral process (see also RD) and iscontinuous with anterior end of OD4.OD3 absent.OD4 originates on dorsal end of Pb3 process andinserts along anterodorsal edge of medial end of Eb4.OP absent or indistinguishable medially from Ad4and/or SO. Muscle laterally in this region joins Eb4and Cb4 medial to joint of these two elements.Ad 1-3 absent.Ad4 absent or indistinguishable medially from OPand/or SO (see OP and Ad5).Ad5 on posterodistal end of Cb4 and dorsodistalend of Cb5, inseparable medially from SO; some fi-bers on Cb4 questionably represent Ad4.Remarks. The muscles in this area are surroundedand penetrated with tough connective tissue, and thedelineation of the muscles is not as clear as they ap-pear in Plate 5.RD inserts anteriorly on raphe that joins SO, Pb3,and TPb3 in region at and ventral to posterior end ofTPb3.Remarks. Wiley (1976:32) stated that the RDs oflepisosteids share muscle fibers with the circularmuscle layer of the esophagous [SO]. We note thatSO does not have a longitudinal muscle fiber layerin Atractosteus and that RD fibers are not continuouswith the SO circular muscle fibers, although the fibers
of both RD and SO are permeated with (joined by)anastomosing connective tissue. (See discussion ofRD in Results section of pre-acanthomorphs.)SO attaches to Pb3 anteriorly and is free from Eb4,but dorsolaterally is inseparable from OP and/or Ad4on Eb4; comprises only transverse muscle layer.Additional remarks. SCL absent. TV4 absent. Wi-ley (1976:32), erroneously reported TV4 present inlepisosteids, and concluded that its presence was asynapomorphy of the Neopterygii. His fig. 14, how-ever, accurately shows TV4 is absent in lepisosteids.TV4 first appears in Amia and is a synapomorphy ofthe Halecostomi. Pb2 is well removed from contactwith Pb3. There are two cartilaginous SPbs, one eachon the uncinate processes of Ebl and Eb2. There isa pair of wedge-shaped ACs between the anterodistaland posterodistal ends of Ebl and Eb2 and their re-spective Cbs. See also references to Wiley (1976) inAdditional remarks under Amia calva.HalecomorphiAMIIDAEAmia calva Linnaeus, USNM 230909, 120 mm TL.Plate 6
Description.Remarks. Allis (1897) and Holstvoogd (1965) de-scribed and illustrated the dorsal gill-arch muscula-ture of A. calva; however, we find the illustrationsand descriptions wanting.LEI on dorsoposterior edge of Ebl just lateral toproximal cartilaginous end.LE2 on dorsomedial cartilaginous process (end) ofEb2, which articulates with Pb3 (end ventromediallyarticulates with Pb2).LE3 on posteriorly extending portion of Eb3 car-tilaginous medial end.LE4 on both cartilage and bone at distal end ofEb4.LP absent.LI1 on anterior cartilaginous tip of Pb2.LI2 on cartilaginous portion of Pb3 dorsal to un-coalesced tooth plates.LI3 absent (Pb4 absent).TD comprises only TPb3, which attaches to an-terolateral portion of Pb3, with some posteromedialfibers on right side continuous with right-side RD.OD3 absent.OD4 relatively short; origin on anteromedial car-tilaginous portion of Pb3, insertion on medial end ofEb4, posteromedially joining small raphe with dor-somedial end of OP.OP on Eb4 dorsomedially, on Cb5 distally, contin-uous with SO ventromedially, overlaps ventral end ofAd5; ER absent.Ad 1-3 absent.
NUMBER 11 25
Ad4 on dorsoposterior surface of Eb4 and dorsalsurface of Cb4.Ad5 on distal cartilaginous ends of Cb4 and Cb5.RD completely separate from SO; with separatedorsal and ventral attachments on Pb3 ventral toTPb3; few or no muscle fibers continuous with TPb3on left side, several fibers continuous with TPb3 pos-teriorly on right side. (See discussion of RD in Re-sults section of pre-acanthomorphs.)SO on Pb3 posteriorly; does not attach to Eb4;longitudinal muscle fibers sparsely distributed withinSO, roughly paralleling mucosal folds, extend ante-riorly to posterior margins of posteriormost upperpharyngeal tooth patches.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl positioned horizontally, in line with Ebl.We agree with Wiley's (1976:30-31) conclusions,based on Amia and lepisosteids, which have onlyOD4, that TD and OD muscles are apomorphic forNeopterygii; however, OD3 is a synapomorphy of theTeleostei. OsteoglossomorphaHIODONTIDAEHiodon alosoides (Rafinesque), USNM 350554, 2specimens, 102-130 mm.Plate 7A, B
Additional material. ? = Hiodon tergisus Lesueur.USNM 266581, 82.1 mm; USNM 342742. 154mm; USNM 350555, 163 mm.Plate 7C
Description.LEI finely, tendinously on cartilaginous medialend of Eb 1
.
LE2 on cartilaginous medial end of Eb2.LE3 on cartilaginous tip of Eb3 uncinate process.LE4 dorsodistally on Eb4.Remarks. LE4 origin is well separated posteriorlyfrom the linearly contiguous or clustered origins ofthe other levators. As such, LE4 in Hiodon could beinterpreted as LP (Winterbottom, 1974b:footnote p.252), which, in fishes with both LE4 and LP, alwaysoriginates posterior to the origins of the other leva-tors. In other osteoglossomorphs, except the highlyspecialized Heterotis, the origin of the levator insert-ing dorsodistally on Eb4 is linearly contiguous withthe other levators, and there is no levator insertingposterior to it. Greenwood and Lauder (1981:226)opined that LE4 in Hiodon has been displaced pos-teriorly because of "the large swimbladder extensionin the otic region." The interpretation of the poste-riormost levator as LE4 might be contraindicated bythe condition of the levators in one of the three spec-imens of H. tergisus. The disposition of the levatorsin two of the specimens of H. tergisus we examined
is similar to that of both specimens of H. alosoides(Plate 7A). In the third specimen of H. tergisus(USNM 342742), however, there is, bilaterally, aslender, additional levator tendinously inserting onEb4 well anterior to the typical posteriorly insertingLE4 (Plate 7C). The additional muscle is probablyanomalous. Our observations indicate that LP is pres-ent in pre-acanthomorphs only among some of thefishes belonging to the Otocephala (some clupeo-morphs and ostariophysans), but it is usually presentin acanthomorphs.LP absent (see remarks under LE4).LI1 on bony dorsal surface of Pb2.LI2 (see Remarks under LI3).LI3 (see Remarks).Remarks. There appears to be only one other LIbesides LI 1 . In the smaller specimen of H. alosoides,a few fibers of the second LI insert posteriorly onPb3. and the remainder insert on Pb4. In the largerspecimen, the second LI inserts extensively on bothPb3 (posteriorly) and Pb4 (anteriorly). In two of thethree specimens of H. tergisus. the second LI hasminor insertion posteriorly on Pb3 and the remainderon Pb4. In the third specimen, the second LI hascompletely separate insertions on Pb3 and Pb4, withthe more extensive insertion on Pb4. It is not possibleto decide if the second LI represents: LI2 insertingpartially on Pb4, fused LI2 and LI3, or LI3 with par-tial insertion on Pb3. Nelson (1969b: 18) indicated thepresence of only two Lis in osteoglossomorphs, butdid not specify their insertions. We find only one LIin Petrocephalus. Mormyrus, and Gymnarcluis (LI1),and only two in Arapaima (LI1, LI2). Notopterus,Heterotis. Osteoglossum, Scleropages, and Pantodon
all have three (LI 1-3).TD is broad, undifferentiated, consists of TPb3-Pb4-Eb4, continuous posteriorly with SO, fromwhich it is differentiated only by divergence of mus-cle fibers at posterior attachment to Eb4.OD2 absent. ? Present in two smaller specimens(Plate 7C) in which it is small, originates on Pb3 nearposterolateral origin of OD4, and inserts jointly withLE2 on Eb2 cartilaginous medial end.Remarks. Among pre-acanthomorphs, OD2 is oth-erwise known only in Heterotis, which, together withHiodon. also lacks TEb2. The bones joined by thebilateral pair of OD2 muscles are the same as thosejoined by TEb2 (or TEb2 complex) of other osteo-glossomorphs (Arapaima. Osteoglossum, Scleropa-ges, Pantodon). TEb2 in these fishes, however, is atransverse muscle connecting the two sides of the gillarches. It is possible that OD2 is a modified TEb2.OD3 origin on Pb3 lateral to and continuous withOD4 origin, insertion dorsally on Eb3 uncinate pro-cess just ventral to LE3 insertion.OD4 origin on Pb3 medial to and continuous withOD3 origin, insertion on anteromedialmost edge of
26 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Eb4 coincident with anterior attachments of Ad4 andOP.RecD4 origin on Eb4 dorsally anterolateral to OD4insertion; insertion split, dorsally on dorsoposteriorsurface and edge of Eb3 uncinate process and ven-trally on medial end of Eb3 (not illustrated), vari-ously fusing with LE3 basally.Remarks. Nelson (1967a:281; 1967c) used theterm obliquus inferior [of the dorsal gill-arch mus-culature] for a muscle joining two adjacent epibran-chials in eels. Winterbottom (1974b:259) noted thatobliquus inferior is a muscle of the eye and that theterm also had been applied to various other muscles.He, therefore, coined the name recti dorsales (sing.,rectus dorsalis) for longitudinal muscles joining ad-jacent epibranchials "to reflect their analogous po-
sition to that of the recti ventrales." RecDs occur ina diverse variety of fishes (e.g., osteoglossomorphs,anguilliforms, callionymids, etc.), and are probablyhomologous only within restricted groups of taxa.OP a narrow strap, originating on posteromedialmargin of Eb4 coincident with ventral portion ofOD4 insertion, ending posteroventrally at ER, whichis joined ventrally by Ad5 and SO; second, even slen-derer portion of OP may be present, originating sep-arately somewhat ventral to narrow portion, insertingtogether with narrow portion. ? Originating laterallywith Ad4 attachment; not distinguishable from SO.Remarks. Below ER, it is not possible to decide ifOP is continuous laterally with Ad5 or is replaced bya medially expanded Ad5.Adl-3 absent.Ad4 origin on Eb4 dorsomedially, coincident withOD4 insertion, insertion on dorsoposterior surface ofCb4.Ad5 on Eb4 dorsoposteriorly and on Cb5 laterally,joining ER medially with SO (see also remarks underAd5 in description of Arapaima).RD absent.SO longitudinal muscle layer thick dorsally and
ventrally, thin to almost absent laterally, extends an-teriorly only to horizontal at posterior margin ofTPb4-Eb4.Additional remarks. SCL absent. TV4 free fromCb5s. The distinct crossed anterior ends of Pb3s aredorsal to the crossed anterior ends of Pb2s. and thecomplex is completely encompassed in a ball of thickconnective tissue (not shown) that attaches to theventral surface of the cranium.
NOTOPTERIDAE
Notopterus notopterus Pallas, USNM 344674, ca.127 mm; USNM 191464, ca. 143 mm.Plate 8
Description.LEI on dorsal Ebl bony surface just lateral to car-tilaginous medial end; dorsoanteriorly, fibers on me-dial surface mesh with fibers on lateral surface ofLE2.LE2 dorsally on Eb2 cartilaginous and bony sur-faces just lateral to joint with Pb3; dorsoanteriorly,fibers on medial surface mesh with fibers on lateralsurface of LE3' and fibers on lateral surface meshwith fibers on medial surface of LEI.LE3 on tip of Eb3 uncinate process jointly with
"3" attachment of OD3-4.LE3' on dorsal surface of medial end of Eb3; dor-soanteriorly, fibers on lateral muscle surface meshwith fibers on medial muscle surface of LE2, andfibers on medial surface mesh with fibers on lateralsurface of LI3. See also "Additional remarks" be-low.LE4 on cartilaginous dorsoposterior end of Eb4.LP absent.LI1 on bony dorsal surface of Pb2 just lateral tomedial cartilaginous end.LI2 on dorsal surfaces of Pb3 and Pb4; insertionparalleling and distinct from LI3 insertion.LI3 on dorsal surface of cartilaginous Pb4, paral-leling and distinct from portion of LI2 insertion onPb4, also a few fibers dorsally on medial end of Eb3;dorsoanteriorly, fibers on lateral surface mesh withfibers on medial surface of LE2.TD broad, undifferentiated, consists of TPb3-Pb4-Eb4. In smaller specimen, TD overlies and is freefrom anterior end of SO. In larger specimen, TD iscontinuous posteriorly with SO, from which it is dif-ferentiated only by divergence of muscle fibers atposterior attachment to Eb4.OD3-4 origin on Pb3 bony portion adjacent to an-terior cartilaginous tip, posteriorly muscle divideslongitudinally with branch inserting on cartilaginoustip of Eb3 uncinate process and branch dorsally oncartilaginous distal end of Eb4 where it forms lateralhalf of raphe with dorsal end of Ad4 and is partiallycontinuous with LE4 insertion.OD4 origin beginning on dorsoposterior bony por-tion of Pb3 medial to LI2 insertion, continuing alongmost of length of cartilaginous Pb4, and inserting onEb4 dorsodistally and forming medial half of raphewith Ad4.RecD4 a sliver of muscle tendinously attached an-teriorly to dorsal tip of Eb3 uncinate process; passesposteriorly between Eb4 portion of OD3-4 and OD4,conforming with surface of OD4, and joins raphewith Ad4 along with and posterior to OD4.OP absent or fused indistinguishably with SO; ERpresent at about level of dorsalmost attachment ofAd5.Adl-3 absent.Ad4 on EB4 dorsoposteriorly, joined there by ra-
NUMBER 11 27
phe with OD3-4, OD4, and RecD4; on Cb4 dorsalsurface anterior to Eb4-Cb4 joint.Ad5 on posterior cartilaginous distal end of Eb4and posteromedial surface of Cb5, mostly undiffer-entiated from SO laterally.RD absent.SO longitudinal muscle fibers thickest dorsally, ex-tending anteriorly to anteromedial end of Pb3.Additional remarks. SCL absent. TV4 free fromCb5s. Holstvoogdt (1965:fig. 5) illustrated the leva-tors of the notopterid Xenomystus nigri. He foundonly six, four of which appear to be equivalent to ourLEI, LE2, LE4, and LI1. The equivalents of the othertwo levators, which he labels as L.III EX. andL.III.INT. are unclear, but possibly represent our LI2and LI3, which, if true, would indicate that Xeno-mystus appears to lack LE3 and LE3'.GYMNARCHIDAEGymnarchus niloticus Cuvier, USNM 319410. ca.315 mm TL. Plate 9
Description.Remarks. The dorsal gill-arch musculature andskeleton are considerably reduced and muscle fusionsare evident. Because of this, it is simpler to describemuch of the musculature in narrative form.The only levator that is clearly present is LI1,which is on Pb2 posteromedially. A large, long mus-
cle, or muscle complex, probably incorporating LEIand/or RecDl ("LEI complex" on Plate 9A), ex-tends well anteriorly from the anterior edge of Ebl.The dorsal surface of the muscle attaches along theventral surface of the cranium. A long muscle, pos-sibly comprising LE2 anteriorly, is incorporated dor-sally in the LEI complex. As this questionable LE2extends posteriorly, it attaches musculously to theposterior bony surface of Ebl, then tendinously tothe bony surface of Eb2 (incorporating a RecD2?),continues musculously (as RecD3?) posteriorly, andforms tendinous anterior and posterior attachments toEb3 and the anterior bony edge of Eb4. A short mus-cle (M. Pb3-Eb3-Eb2) connects the posteromedialbony dorsal surface of Eb2 with Pb3 and Eb3, wherethe latter two elements meet. Another muscle,RecD4, joins the dorsoposterior bony surface of Eb3with the dorsal bony surface of Eb4.TD comprises TPb2-Pb3-Eb4, originates anteriorly
at about level of Pb2s as sparse fibers on surface ofCT; anterolaterally, fibers appear to form part of me-dial edge of LEI muscle complex. TD becomessheet-like posteriorly beginning at about level ofPb3s, and is undifferentiated posteriorly from SO.OD3 absent.OD4 absent.OP absent; presence or absence of ER unclear.
Ad 1-3 absent.Ad4 on ventral surface of Eb4 and dorsal surfaceof Cb4.Ad5 dorsally attaches to tendon extending laterallyfrom Eb4, and ventrally to posterodistal end of Cb5;a unique, laterally extending strap of SO arises fromanterior surface of tendon and sandwiches Ad5 be-tween it and another SO strap arising from Cb5 areaventrally.RD absent.SO longitudinal muscle layer comprising sparselydistributed fiber bundles in "cottony" matrix, ex-tending anteriorly to anterior end of TD.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl and Pb4 are absent. Tiny AC present onlyon left side at posteromedial tip of Eb 1
.
MORMYRIDAEPetrocephalus tenuicauda (Steindachner), USNM118801. 92.3 mm. Plate 10
Additional material. ? = Monnyrus longirostris Pe-ters, USNM 261878, 225 mm.
Description.LEI on Ebl anterior and posterior edges (bridgingchannel that carries blood vessels and nerves) withsome fibers also on Pbl; fuses posteromedially withLE2.
?
Unclear if LE2 is represented among fusions.LE2 on Eb2 posteromedially, fuses laterally withLEI and medially with M. Pb3-Pb4-Eb2. which mayrepresent a modified LI2 and/or LI3. On left side, amuscle originates narrowly and separately on crani-um and inserts narrowly, tendinously on Eb2 amongfibers of broad, fused complex LE2. This muscle isalso labeled LE2 on Plate 10. ? Unclear if LE2 ispresent, but if so, is incorporated in RecD2.LE3 on all bony Eb3 uncinate process just lateralto RecD3 and OD3 attachments. ? Extremely longand variable in width along its length, changing al-most to thin tendon in places.LE4 absent.LP absent.LI1 on dorsomedial surface of Pb2.LI2 see LE2.LI3 see LE2.M. Pb3-Pb4-Eb2 originates on posterodorsal sur-face of Pb3 and dorsal surface of greatly reducedPb4, and inserts on Eb2 posteromedially; anteriorly,fuses with lateral portion of LE2. ? LE2 portionquestionably present.TD apparently undivided, consisting of TPb3-Pb4-Eb4, more-or-less continuous posteriorly with SO. ?Comprises TPb3 and questionably TPb4-TEb4. TPb3dense band of muscle fibers attached to Pb3 dor-soanteriorly, abruptly changing posteriorly to loose
28 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
strands of diagonal muscle fibers attaching to Pb4,and these continuing posteriorly as dense network offibers attaching to Eb4 posteromedially and continu-ing posteriorly as SO.OD3 tendinously on Pb2 (rather than Pb3 as inmost fishes), and tendinously on all bony Eb3 unci-nate process. On right side only, there is a possiblyanomalous strap of muscle extending anteriorly fromEb3 uncinate process that fuses anteriorly with LE1-LE2 complex; strap lies dorsal to M. Pb3-Pb4-Eb2.
?
Strap absent.OD4 absent.M. Pb2-Ebl narrowly on posteromedial edge ofEbl (ventral to anteriorly extending, fused LI2+3?and LE2) and broadly on anterodorsal surface of Pb2.
?
Originates on dorsoanterior surface of Pb3, withslender tendon on medial surface of muscle expand-ing anteriorly and joining CT covering Pb2-Pb3 joint.RecD2 broadly on dorsal surface of Eb2 ventral toLE2, and narrowly on posteromedial corner of Eblventral to LE2. ? Presence of LE2 questionable.RecD3 on all bony uncinate process of Eb3 anddorsal surface of medial cartilaginous end of Eb2. ?Asymmetrical; left side with cluster of small musclesoriginating on Pb3, Pb4, and Eb4 medially, and in-serting on Eb2 anteromedially; right side with twosmall muscles, one from Eb2 anteromedially to Eb4anteromedially, the other from Eb2 anteromedially toEb3 dorsomedially.OP questionably present.Remarks. ER, often denoting the ventral end of OP,when OP is present, is irregularly indicated on bothsides of SO at about the level of the distal end ofEb4. On the left side there is a narrow strap of musclethat originates on Eb4 and ends at the raphe. On theright side there is no strap. ? Many raphe-like CTintrusions in SO dorsolaterally; no defined strap-likemuscle; muscle fibers complexly oriented. Differenc-es from Petrocephalus possibly due to much largersize of specimen.Ad 1-3 absent.Ad4 dorsally on Eb4 dorsoanteriorly and ventrallyon Cb4.Ad5 dorsally on dorsoposterodistalmost cartilagi-nous tip of Eb4 and ventrally on Cb5; fuses mediallywith SO.RD absent.SO longitudinal fibers essentially restricted to dor-
sal arc of SO ventral to transverse fibers, few if anylongitudinal fibers ventrally. ? Fibers sparsely dis-tributed dorsally, few if any laterally or ventrally.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 is greatly reduced and offers little surfacefor attachment of a levator. This could explain theabsence of LI3, if present ancestrally.Bishai (1967:20-21) described and illustrated thedorsal gill-arch musculature of Mormyrus caschive
Linnaeus. Using terminology different from ours, hereported the presence of LEI -4, LI2 and 3 (on Pb2and 3, respectively, which would equal our LI1 andLI2), OD3, and a muscle that we would term OD3'(the two originating on separate parts of Pb3 and in-serting together on bony Eb3 uncinate process),RecD2 and 3, and Ad4. He did not mention fusionof any muscles, and his illustrations do not indicatethe presence of cartilage. It is possible that his LI3may insert partially on Pb4.Nelson (1969b: 18) reported on the dorsal gill-archmuscles of Mormyrus ovis, in which he recognizedonly LEI and 2, LI1, OD3, OD4, and RecD 2 and 3(no mention of Ads). Although Nelson mentionedBishai's study, he made no comment on the differ-ences between his findings and those of Bishai. Webelieve there is considerable room for differences ininterpretation of the muscles, as well as the possibil-ity that no two specimens of the same species (or twosides of a specimen) are more than generally similar.Our remarks undoubtedly also apply to our speci-mens of Petrocephalus and Mormyrus.ARAPAIMIDAE
Heterotis niloticus (Cuvier), USNM 303214, 237mm. Plate 11
Description.Remarks. Heterotis is unique among osteoglossi-forms we examined in having an epibranchial organ.The organ is formed by the complex coiling and ex-tensive development of what is probably the dorsalcartilaginous margin of Eb4, and, at least, includesthat margin. The thin muscle branches attaching var-iously to the surface of the cartilage are difficult tohomologize with reasonable certainty. Adding to thedifficulty are bilateral asymmetry of these muscles, asheet of muscles lining the internal surface of thecartilaginous coil, and an extensive network of in-vasive nerves supplying the organ internally and ex-ternally. We have not indicated nerves in other illus-trations, but in Plate 11 A, we show a large nerve(identification not determined), which superficiallyresembles a muscle. The nerve penetrates the dor-somedial side of the cartilaginous coil.LEI on cartilaginous and bony dorsomedial end ofEbl, fusing dorsally with anteroventral, longitudinalfibers of LE2. Short ligament (not illustrated) joinsdorsomedial surface of cartilaginous medial end ofEbl to bony dorsal surface of anterior end of Pb2.LE2 on dorsalmost prominence of cartilaginousmedial end of Eb2; muscle fusing ventrally with dor-
sal, longitudinal fibers of LEI. RecD2 parallels LE2ventrally, with ribbon of CT (not illustrated), whichconnects cartilaginous processes of Ebl and Eb2, ly-ing laterally and attaching to both muscles. Short lig-
NUMBER 11 29
ament (not illustrated) joins dorsomedial surface ofcartilaginous medial end of Eb2 to dorsal bony sur-face of anterior end of Pb3.LE3 tendinously on cartilaginous tip of Eb3 un-cinate process; muscle thin, weak, posterodorsally di-rected, applied medially to lateral surface of epibran-chial organ; origin not noted, but apparently not at-taching directly to cranium; muscle possibly absenton right side, or lost during dissection.LE4 on dorsal surface of epibranchial organ, con-forming with curvature of organ; possibly absent onright side or lost during dissection.LP absent.LI1 on Pb2 mid-dorsally.LI2 on bony Pb3 dorsal surface; right side withslender posterior branch inserting anteriorly on dorsalsurface of cartilaginous Pb4; branch absent on leftside.LI3 attached to fascia covering surface of cartilag-inous Pb4; anterior, elevated portion of muscle inter-rupted at attachment, continuing posteriorly as CTand then small portion of muscle applied to posteriorsurface of Pb4 (muscle uninterrupted on right side);CT at base of elevated portion gives rise to posteriorattachment of M. Pb4-Eb2. LI3 passes anteriorly lat-eral to LI2.TD possibly absent anteriorly (see OD2), compris-es TPb3-Pb4 (on posterior Pb3 segment and, possi-bly, also on Eb4), and is undifferentiated posteriorlyfrom SO.OD2 originates on dorsal surface of autogenouscartilaginous Pb3 anterior segment (Nelson, 1968a:268) anterior to OD3 origin, and inserts on cartilag-inous medial end of Eb2 dorsally. Medial fibers ofright-side OD2 overlap anterior end of left-side OD2and attach on dorsoanterior end of left-side Pb3 car-tilaginous segment. Anterior ends of Pb3 cartilagi-nous segments and OD2 origins are enveloped intough CT pad.Remarks. OD2 is known otherwise in pre-acantho-morphs only in some specimens of Hiodon tergisus.See remarks under OD2 in description of Hiodon.OD3 originates on dorsal surface of Pb3 anteriorsegment and inserts by long tendon on tip of elongatecartilaginous cap of Eb3 uncinate process. Right-sideorigin is ventral to OD2 origin; left-side origin begins
at posterior end of OD2 origin.OD4 absent on left side; questionably representedon right side by thin, elongate muscle originating onPb3 just anterior and medial to LI2 insertion and in-serting on posterior (= medial in view) cartilaginouswall of EO (Eb4).OP absent; ER absent.RecD2 on posterior surface of dorsal prominenceof medial cartilaginous end of Ebl and anterior sur-face of dorsal prominence of medial cartilaginous endof Eb2, parallels LE2 ventrally, with CT ribbon (not
illustrated) connecting cartilaginous processes of Ebland Eb2 passing laterally and attaching to both mus-
cles. Left-side RecD2 comprises a vertical pair ofmuscles, each member of which attaches tendinouslyto Eb2; right-side RecD2 has vertical series of threeseparate, but contiguous musculous attachments toEb2.RecD3 absent.RecD4 absent, but long, broad ligament arisingfrom Eb3 uncinate process and lateral edge of Pb4inserts on dorsal bony surface of Eb4.M. Pb4-Eb2 attaches to posterior surface of dor-somedial cartilaginous extension of medial end ofEb2 and inserts by flat tendon on Pb4 ventral to in-sertion of elevated portion of LI3 (q.v.).Ad 1-3 absent.Ad4 fan-like, attaching inner cartilaginous ventro-posterolateral surface of EO (Eb4) to bony medialsurface of posterior end of Cb4; dorsally, begins atlower internal rim of large foramen in anterior sur-face (lateral in view) of cartilaginous posterior endof Eb4 (Plate 1 IB) and expands anterodorsally end-ing as fine sheet of CT attaching to medial surfaceof dorsoposterior bony flange of Eb4. Large, fan-shaped gill-filament muscle (not illustrated) on pos-terolateral bony surface of Eb4 matches somewhatthe shape of Ad4 on internal surface. On left side, anapparently anomalous slip of Ad4, which attachesdorsal to remainder of muscle, is visible through theEb4 foramen, after removal of fine sheath of musclelining inner wall of epibranchial organ.Ad5 small, horizontal muscle joining posteroven-tral cartilaginous surface of EO (Eb4) to dorsopos-terior cartilaginous surface of Cb5.RD absent.SO longitudinal muscle fibers questionably absent,sparse if present.Additional remarks. SCL absent. TV4 free fromCb5s. Glottis present.
Arapaima gigas (Cuvier), USNM 177528, 2 speci-mens, ca. 135-139 mm.Plate 12
Description.LEI on Ebl dorsoposterior edge just lateral to me-dial cartilaginous end.LE2 absent.LE3 absent.LE4 on dorsoposterior edge of distal cartilaginousend of Eb4.LP absent.LI1 on Pb2 dorsoposteriorly.LI2 on dorsal surface of Pb3 posteromedially.LI3 absent.Remarks. M.Pb4-Eb2 conceivably represents LI3
30 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
that has shifted its origin to Eb2. (See also remarksunder LI3 in Hiodon).TD comprises two parts: TPb3-Eb2 and TPb3-Pb4-Eb4. TPb3-Eb2 a very thin sheet of muscle at-taching to dorsal surface of Pb3 anterior cartilaginoussegment (Nelson 1968a:267-268) and cartilaginousprocesses of Pb3 (posterior bony segment) and Eb2,where these processes join, and is anterodorsal toTPb3-Pb4-Eb4, which attaches to medial edges ofPb3, Pb4, and Eb4. TEb2 is dorsal to anterior attach-ments of OD3 and OD4. TPb3-Pb4-Eb4 is undiffer-entiated posteriorly from SO.OD3 on autogenous anterior Pb3 cartilaginous seg-ment and Eb3 uncinate process.OD4 anteriorly on Pb3 bony portion, posteriorlyon dorsomedial edge of Eb4, fusing posteriorly withRecD4.Remarks. Our interpretation of OD4 and RecD4(q.v.) may be in error and the muscle we indicate asRecD4 may be the Eb3 branch of an OD3?4, a mus-cle otherwise limited to Notopterus and Pantodonamong Osteoglossomorpha. Pantodon lacks a sepa-
rate OD4. Only Notopterus among the Osteoglosso-morpha has OD4, OD3-4, and RecD4, and Notop-terus lacks OD3, which is present in all other Osteo-glossomorpha except Pantodon and the highly spe-cialized Gymnarchus, which lacks OD completely.Our general observation among Neopterygii, is that,excluding Notopterus, OD4 or OD3-4 may be pres-
ent, but not both. Very few neopterygian taxa (Gym-narchus; the ostariophysan Chanos) lack an OD, alack we consider autapomorphic for each of thesetaxa.OP absent; ER absent.Remarks. There is considerable CT permeatingand obscuring the nature of the area that might bedefined as OP.RecD2 on cartilaginous posterior edge of Eb 1 me-dial end and bony posterior edge of Eb2 medial end;muscle strands continuous posteriorly with M. Pb4-Eb2.RecD3 absent.Remarks. Muscle reported to be present by Nelson(1969b: 18), but he may have considered it the sameas the muscle we designate M.Pb4-Eb2 (q.v.). We useRecD to indicate a muscle that connects two succes-sive epibranchials.RecD4 on posterior edge of cartilaginous tip ofEb3 uncinate process, fusing posteriorly with poste-
rior end of OD4. Possibly absent, see remarks underOD4.M. Pb4-Eb2 on Pb4 dorsally and posteromedialcartilaginous edge of Eb2, muscle strands continuousanteriorly with RecD2 (see also remarks under LI3).Ad 1-3 absent.Ad4 broadly on broad posterolateral surface of
Eb4 and narrowly on dorsodistal surface of Cb4 me-dial to Eb4-Cb4 joint (not visible in illustrations).Ad5 slender, from outer edge of dorsoposterior-most distal cartilaginous tip of Eb4 to posterior sur-face of cartilaginous distal end of Cb5.Remarks. Winterbottom (1974b:254-255) dis-cussed the problem of what to call a muscle attachingEb4 to Cb5 when there is only one such muscle. BothOP and Ad5 may exhibit these attachments (Ad5 mayattach Cb5 to Eb4, Eb4*, Eb5, or Cb4, but almostalways attaches to Eb5 when Eb5 is present (or toEb4* when this state replaces an autogenous Eb5),exceptions: cyprinids and Searsia, Argentiniformes).OP is usually attached well medial to the distal endof Eb4, whereas Ad5 attaches to the distal end (es-pecially when Eb4* is present). Winterbottom rec-ommends using Ad5 for the single muscle, and weconcur in the present case, especially as the appear-ance of the single muscle is more like an Ad5 thanan OP, and OP appears to be present in most otherosteoglossomorphs (see Hiodon alosoides).RD absent.SO longitudinal muscle fibers questionably absent,sparse if present.Additional remarks. SCL absent. TV4 free. SmallAC attached to dorsolateral tip of Cb5 (not reportedby Nelson, 1968a), found otherwise only in Panto-don, thus providing evidence supporting the cladecomprising these two genera and Heterotis, whichlacks the AC. Arapaima and Heterotis are the onlyosteoglossiforms we examined that have a glottis(opens dorsally into an air sac), at least one that isso close to the dorsal gill arches. The air sac is joinedbroadly to the SO, but was not illustrated.PANTODONTIDAEPantodon buchholzi + Peters, USNM 303224 (ca. 80mm), USNM 353993 (2:50.4-59.2 mm), USNM355626 (72 mm). Plate 13
Description (composite).LEI absent.LE2 on dorsoposterior edge of Eb2 somewhat lat-eral to medial end, meets attachment of TEbl-Eb2on Eb2.LE3 absent.LE4 on dorsoposterior edge of Eb4 somewhat me-dial to lateral end, meets attachment of OD3-4 onEb4.LP absent.LI1 primarily dorsally on Pb2, wrapping aroundcartilaginous medial end of Ebl with slight insertionon medialmost end of Ebl.LI2 on dorsolateral bony surface of Pb3, insertionposteriorly continuous with that of LI3 Eb4.LI3 anteriorly on posterolateral cartilaginous end
NUMBER 1
1
31
of Pb3, but mainly on dorsolateral surface of UP5(few fibers on cartilaginous anteromedial end of Eb4in one specimen), insertion anteriorly continuouswith that of LI2, posteriorly continuous with smallgroup of SO muscle fibers.TD comprises TEbl-Eb2 and TPb3-Eb4. TEbl-Eb2 divides briefly laterally with short branch attach-ing to posterior edge of medial end of Ebl, and longbranch attaching along posterior edge of medial halfof Eb2; continuous posteriorly by few crossing di-agonal muscle strands with TPb3-Eb4. TPb3-Eb4 at-taches to medial edges and surfaces of Pb3 and Eb4and is continuous posteriorly (undifferentiated) withSO.M. Pb2-Eb2 originates on ventroanterior surface ofPb2 (few weak muscle strands may attach on ventro-medial end of Ebl) and flares posteriorly as it insertsalong dorsoanterior edge of medial half of Eb2.OD3-4 origin on Pb3 dorsal surface, divides pos-teriorly with slender branch inserting on Eb3 mid-posterolaterally and much broader branch insertingon Eb4 mid-dorsolaterally together with insertion ofRecD4.Remarks. Muscle is similar to that of Notopterus,q.v.RecD4 originates ventrally on posteromedial edgeof Eb3 and inserts dorsally on Eb4 together with Eb4branch of OD3-4.OP questionably absent; however, possibly repre-sented by a thin, narrow band of muscle originatingon Eb4 near medial end of Ad4 and scarcely sepa-rable medially from OP, fans out ventrolaterally,passing anterior to Ad5, Eb5, and AC, and attachesto Cb5 at and anterior to ventral attachment of Ad5.ER absent.Adl-3 absent.Ad4 dorsally on posterolateral surface of Eb4, ven-trally on Cb4 dorsal surface at anterior angle formedby Eb4-Cb4 joint.Ad5 dorsally on posterolateral surface of Eb5, ven-trally on posterolateral surface of Cb5 together withlateral attachment of TV5.SO longitudinal muscle fibers questionably pres-
ent.SOD broad.RDs separate, on posterior ends of Pb3 and UP5,which is ventral to Pb3 posteriorly and Eb4 anteri-
orly.Remarks. RD is autapomorphic in Pantodon.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl and Pb4 absent (we presume that UP4 isalso absent and that the large toothplate ventral toPb3 and Eb4 is UP5). Eb5 and accessory cartilage(AC) present: Eb5 transverse, joined laterally andmore-or-less equally to posterodistal ends of Eb4 andCb4 and medially to dorsal end of AC; AC joinedventrally to dorsodistal end of Cb5. The orientation
of Eb5 is duplicated only in Diplomystes, Ostario-physi, which lacks AC. Among osteoglossomorphs,AC is found otherwise only in Arapaima, thus sup-porting the clade comprising these two genera andHeterotis, which lacks AC.OSTEOGLOSSIDAE
Osteoglossum bicirrhosum Cuvier, USNM 315447,210 mm; USNM 198123, 59.6 mm.Plate 14
Additional material. ? = Scleropages jardini (Sa-ville-Kent), USNM 217049, 215 mm.
Description.Remarks. Because of muscle fusions, our interpre-tations of LEI, LE2, RecD2, RecD4, and some otherproblematic muscles of Osteoglossum are, in part,subjective, and are described together. The smallerspecimen appeared to differ in no significant wayfrom the larger specimen.LEI is a thick ribbon of muscle originating on cra-nium inseparably from dorsal end of Pbl, and in-serting along most of the dorsal surface of elongate,cartilaginous Pbl and dorsomedial end of Ebl. Dor-somedial portion of LEI insertion on Ebl joining ra-phe with anterior end of RecD2, which originates onEb2 dorsomedially. RecD2 broadens considerablymedially, forming what we consider a separate mus-
cle, M. Pb2-Eb2, and attaching anteriorly to dorsalsurface of broad cartilaginous anterior end of Pb2 justventral to insertion of LI1 (anterior end of Pb2 spat-
ulate, becoming almost vertical medial to attachmentof M. Pb2-Eb2). At attachment on Eb2, RecD2 joinscomplex raphe with LE2 insertion, LI3 ventrolateralsurface, and anterior end of M. Pb4-Eb2 (describedbelow following OD4). LE2 and LI3 weakly (thinly)continuous dorsal to raphe on Eb2, with LI3 con-tinuing posteroventrally to its insertion. Anterior por-tion of LI3 insertion on Pb3 just lateral and parallelto LI2 insertion; posterior portion of insertion on car-tilaginous Pb4, fusing and attaching inextricably withorigin of M. Pb4-Eb2. Anterior end of RecD4 andposterior end of OD3 meet in raphe and attach onmedial surface of cartilaginous tip of Eb3 uncinateprocess. RecD4 attaches posteriorly to mid-dorsalcartilaginous edge of Eb4 anterior to insertion ofLE4.
?
Anterodorsal portion of LEI extends dorsal todorsal end of Pbl and originates separately on cra-nium. RecD2 not expanded medially?M. Pb2-Eb2absent?not attaching to Pb2. LE2 overlaps LI3 lat-
erally, but both muscles are clearly separate anteriorto their attachments to Eb2. RecD4 absent.LE3 on cartilaginous dorsal tip of Eb3 uncinateprocess.
32 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LE4 broadly along dorsodistal cartilaginous sur-face of Eb4.LP absent.LI1 on lateral surface of vertical expansion (Nel-son, 1968a:266) of cartilaginous anterior end of Pb2.
?
LI1 in deep excavation on lateral surface of ver-tical expansion of Pb2.LI2 on dorsal bony surface of Pb3; medially, in-sertion parallels and is inseparable from lateral at-tachment of TD; laterally, LI2 insertion parallels andis medial to anterior portion of LI3 insertion (the twomuscles are thin and "'plastered" together); posteri-
orly, insertion impinges on anterior attachment ofOD3, which is on posterior cartilaginous end of Pb3.
? Insertion does not impinge on OD3 anteriorly.LI3 (see initial paragraph under Descriptionabove).TD comprises two portions: short TEb2 and longTPb3-Pb4-Eb4 that is posteriorly scarcely separablefrom SO. TEb2 is broad, somewhat triangular, withpartial median raphe extending posteriorly from mus-
cle's anterior apex; muscle attaches to dorsomedialcartilaginous end of Eb2, where it is inextricablymeshed with complex attachments of LE2, LI3,RecD2, and M. Pb4-Eb2. TEb2 is dorsal to and dis-continuous from remainder of TD. ? An apparentlyanomalus strand of muscle is continuous from pos-terior margin of right side of TEb2 and posteromedialside of OD3.OD3 on dorsoanterior cartilaginous surface of Pb3(excluding small, separate anterior cartilaginous seg-ment of Pb3; see Additional remarks, below) and in-
serts on raphe with RecD4 on medial side of dorsalend of Eb3 uncinate process.OD4 anteriorly ventral to OD3, originates on an-teriormost bony surface of Pb3 medial to LI2 inser-tion, and inserts on dorsomedial cartilaginous surfaceof Eb4 just anteroventral to attachment of RecD4.M. Pb4-Eb2 on Pb4 dorsolaterally, extending an-terolaterally and inserting on dorsomedialmost carti-laginous edge of Eb2.OP probably represented by strap of muscle at-taching dorsally to posterior ventromedial surface ofEb4 ventral to Eb4 portion of TD, extending ven-trally to ER at about level of mid-distal end of Eb4.
? ER particularly well developed.Ad 1-3 absent.Ad4 broadly on posterior surface of cartilaginousdistal end of Eb4, ventrally, narrowly on dorsodistalbony surface of Cb4.Ad5 barely separable from SO medially, on pos-terodistal cartilaginous edge of Eb4 and posterodistalsurface of Cb5.RD absent.SO longitudinal muscle fibers in thin layer sur-rounding esophagus. ? Longitudinal fibers in thick
layer dorsally and ventrally, thinner to almost inter-rupted mid-laterally.Additional remarks. SCL absent. TV4 free fromCb5s. Pb3 with small, autogenous anterior segment;segment absent in Scleropages. Nelson (1968a:268)believed Pb3 cartilage segment among Teleostei tobe restricted to Arapaima and Heterotis, in both ofwhich it is relatively large. He considered its pres-ence an indication that the two genera are closelyrelated. The absence of the segment in Scleropagesand in some Osteoglossiformes with reduced gill-arch skeletons (e.g., Pantodon, Gymnarchus), mightbe secondary.Elopomorpha: ElopiformesMEGALOPIDAEMegalops cyprinoides (Broussonet), USNM 350458,133 mm. Plate 15
Additional material. ? = Megalops atlanticus Valen-ciennes, USNM 303317, 146 mm.
Description.Remarks. Holstvoogd (1965: especially figs. 3aand 3b) illustrated and briefly discussed the dorsalgill-arch muscles of M. cyprinoides. Although ourfindings generally agree with his, we find his illus-trations difficult to interpret, and he indicated thepresence of only two obliqui dorsales, apparentlymissing the muscle we identify as OD4.LEI on Ebl just lateral to cartilaginous tip of un-cinate process.LE2 on dorsoposterior edge of Eb2 just anterolat-eral to SPb2 and medial end of Eb2.LE3 on Eb3 uncinate process, which articulateswith Eb4 uncinate process.LE4 on dorsodistalmost end of Eb4.LP absent.LI1 on Pb2 anterodorsally and on SPbl anteriorsurface.LI2 absent.LI3 on Pb4 dorsolaterally.TD comprises TPb3a and TPb3p-Pb4-Eb4. TPb3ais on Pb3 at and anterior to uncinate process; musclenarrows anteriorly and joins CT between anteriorends of Pb3s; posteriorly, TPb3a overlies anterior-most end of TPb3p-Pb4-Eb4 and is continuous withthat muscle by slender muscle strand. ? TPb3a en-tirely anterior to TPb3p-Pb4-Eb4, continuous poste-riorly with remainder of TD by broad muscle strand.OD3 originates on Pb3 ventral to TPb3, partiallydivided longitudinally, inserts on Eb3 uncinate pro-cess.OD4 originates on Pb3 with and ventral to OD3,and inserts by long tendon anteriorly on Eb4 uncinateprocess lateral to OD4' insertion.
NUMBER 11 33
OD4' originates on Pb3 dorsoposteriorly and onPb4 dorsoanteriorly and inserts on Eb4 uncinate pro-cess medial to OD4 insertion.OP attaches dorsally on Eb4 posteromedial surfaceand joins ER ventrally, undifferentiated from SOventral to ER.M. UP4-Eb5-Cb4 (not illustrated) questionablydistinct muscle strap anterior to (overlain posteriorlyby) OP, with dorsoanterior fibers attaching to lateraledge of UP4 and posteroventrally tendinously attach-ing to junction of Eb5 and Cb4. The alternative toconsidering M. UP4-Eb5-Cb4 as a distinct muscle isthat it is a slightly differentiated SO portion.Ad 1-3 absent.Ad 4 broadly dorsally on posterodistal margin ofEb4, ventrally on Cb4 posteromedially (not visible inlateral view) medial to internal angle formed by Eb4-Cb4 joint.Ad5 complex: mainly attaching dorsally to Eb5posterior surface, but tendinously bound also to pos-terior end of Cb4; muscle fibers shift directions(featherlike) along mid-axis as muscle extends ven-trally, with lateral fibers attaching to Cb5 mediallyand continuing anteriorly along most of ventrallength of Cb5, then changing to broad membranoussheath medial to PCI and PCE (PCI not visible inlateral view, Plate 15C); medial fibers join raphe withTV5.RD absent.SO longitudinal muscle layer restricted to isolatedarea delimited posteriorly by horizontal between dis-tal ends of Eb4s and anteriorly by horizontal betweenmid-lengths of Pb4s.Additional remarks. SCL questionable, similar tothat of Elops (see Additional remarks under Elopssaurus for discussion of SCL). TV4 free from Cb5s.Large, tough CT pad covers dorsoanterior surfacesof Pbls and Pb2s. Eb5 attaches to posterodistal endof Cb4. Johnson and Patterson (1996:273) reportedthe presence of a small interarcual cartilage betweenthe uncinate processes of Eb3 and Eb4, in Megalopsand Elops. We did not find this cartilage in eitherspecimen of the two Megalops species we examined,nor in one cleared and double stained specimen eachof Megalops cyprinoides (USNM 173580) and M. at-lanticus (USNM 357435), but we reconfirmed itspresence in Johnson and Patterson's specimen of M.atlanticus (their specimen of M. cyprinoides was un-available). The presence of the structure is variableand probably of little use for establishing familial in-terrelationships. In Elops (q.v.), the putative rod-likeinterarcual cartilage is not autogenous, but is an ex-tension of the medial cartilage tip of Eb4. We didfind a small cartilage between the tips of the uncinateprocesses of Eb3 and Eb4 of the albulid Pterothrissus(Plate 18), but not in Albula, its sister group.
ELOPIDAEElops saurus Linnaeus, USNM 121694, 126 mm.Plate 16
Description.LEI on Ebl near dorsomedial end and ventral toarticulation with SPbl.LE2 on dorsomedial end of Eb2 and entire anteriorsurface of SPb2; LI2 and LI3 meet LE2 on SPb2.LE3 on dorsal tip of Eb3 uncinate process, joinedthere by OD3 insertion.LE4 attaches by long tendon to dorsodistalmostend of Eb4.LP absent.LI 1 on Pb2 dorsoanteriorly, attaching to entire me-dial surface of SPb2; muscle looping anteromediallyfrom posterior attachment to SPbl, forming twosheet-like layers with less extensive posterior layerincompletely overlapping surface of more extensiveanterior layer.Remarks. Unlike Megalops, there appears to be noseparate origin of LI1 on the braincase, its origin be-ing confined to SPbl.LI2 on Pb3 posterolaterally and SPb2 medial sur-face dorsally.LI3 on lateral edge of Pb4 at junction with UP4,anterior edge attached to SPb2.TD comprises TPb3 and TPb4-Eb4. TPb3 has twosections: smaller dorsoanterior section dorsally onPb3 dorsolateral process, overlies anterior end ofOD3; larger posterior section on Pb3 lateral edge,attaching posterolaterally along common line withLI2, overlies anterior end of OD4, and is posteriorlycontinuous by slender, diagonal muscle strand withSO.Remarks. Winterbottom (1974b:256 and fig. 18)stated that TD is absent in Elops because the trans-verse muscles are continuous with SO. We do notaccept that continuation with SO is sufficient basisfor rejecting the presence of a transversus muscle.Even if one rejects our TPb4-Eb4 as a transversus,TD is clearly indicated by TPb3 in Elops.OD3 anteriorly on posterior edge of Pb3 dorsolat-eral process ventral to TPb3 anterolateral attachment;posteriorly on dorsal tip of Eb3 uncinate process to-gether with LE3.OD4 anteriorly on posteromedial surface of Pb3and anteromedial surface of Pb4, posteriorly on an-terolateral surface of Eb4.OP dorsally on Eb4 posteriorly, ventromediallyjoins ER.Adl-3 absent.Ad4 dorsally broadly on posteriormost surface ofEb4, ventrally on posteriormost end of Cb4 lateral toEb5 ventrally.Ad5 dorsally on medial junction of Cb4 and Eb5,curving first ventromedially around posterior end of
34 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Cb5, then laterally, attaching along lateral surface ofCb5.RDs absent.SO longitudinal muscle layer restricted to isolatedarea delimited posteriorly by horizontal between dis-tal ends of Eb4s and anteriorly by horizontal at an-terior end of TPb4-Eb4.Additional remarks. TV4 free from Cb5s. Presenceof SCL questionable: Hb3 posteroventral flange isstraight anteriorly, curves medially posteriorly, and isattached tightly to posteroventral cartilaginous pro-cess of Bb3, thus forming with contralateral Hb3 ananteriorly open semicircle, the edge of which is linedwith CT. ObV3, on each side, attaches along the lat-eral edge of the straight portion of the Hb3 flangeand is musculously continuous medially with its re-spective RecV4, which attaches to the posterior edgeof the curved portion of the flange, but not to thecartilaginous tip of Bb3. There is no free ligamentousportion along the semicircle, but if some portion werefree, we would interpret the semicircular ligament asbeing present. {NB. There is no autogenous ball ofcartilage ventral to the posteroventral tip of Bb3, asis present in Albula.) Megalops (both species) is sim-ilar to Elops, but Hb3s do not curve medially pos-teriorly, the CT lining the semicircle is slightly loose,and RecV4 is attached to the posterior margin of theposteroventral cartilaginous process of Bb3.Nelson (1968b:fig. 6a) indicated the presence ofan interarcual cartilage between the uncinate processof Eb3 and the proximal end of Eb4. Johnson andPatterson (1996:273) claimed to have confirmed this.Our observations differ. The "interarcual cartilage"of our specimen, and Johnson and Patterson's, on re-examination, is a non-autogenous, elongate cartilag-inous extension of the dorsomedial cartilaginous headof Eb4. The extension attaches to the tip of the Eb3uncinate process. Conceivably, this cartilaginous ex-tension could bud off in large specimens, and suchshould be examined to determine if this occurs. Seealso Additional remarks under Megalopidae.Elopomorpha: AnguillomorphaALBULIDAE
Albula vulpes (Linnaeus)?, USNM 2475 11, 1 27 mmSL. Plate 17
Description.LEI broadly on Ebl, attaches to skull separatelyfrom other LEs, incorporates tendon dorsally.LE2 broadly on Eb2, incorporates tendon dorsally.LE3 on Eb3 uncinate process ventral to cartilagi-nous tip.LE4 on anterior surface of levator process of Eb4just ventral to cartilaginous edge.
LP absent.LI1 on Pb2 anteriorly, comprises two ventrallycontinuous sections; anterior section shorter than pos-terior section, attached all along medial surface ofcartilaginous SPbl.Remarks. See remarks under LI1 in Aldrovandia(Halosauridae) description.LI2 on dorsolateral bony portion of Pb3.LI3 on dorsolateral edges of UP4 and UP5, inser-tion continuous medially with attachment of M. UP4-Eb2.TD comprises two, more-or-less continuous sec-tions, TPb3a, which attaches along anterior marginof Pb3 dorsal process, and TPb3p-UP4, which attach-es laterally to medial edges of Pb3 and UP4, abuttingOD4 ventromedial margin; separable posterolaterallyfrom SO by slight change in orientation of musclefibers, but continuous posteriorly otherwise with SO.OD3 long, slender, originates on bony portion ofPb3 dorsal process and inserts on cartilaginous tip ofEb3 uncinate process; anomalous slip of muscle aris-es from origin of left-side OD3, passes through TPb3and inserts on cartilaginous tip of Eb3 uncinate pro-cess.OD4 large, vertically oriented; ventromediallyabutting TPb3; origin curving medially, attachingcontinuously from posterodorsal edge of Pb3, acrossmedial cartilaginous process of Eb3, over Pb4 to baseof Eb4; muscle twisting on itself dorsally (clockwiseon right side, counterclockwise on left), almost di-visible into two parts, inserting on dorsomedial (un-cinate) process of Eb4.Remarks. Wiley (1976:fig. 13c) believed this mus-cle to be a transversus dorsalis posterior and madeno mention of the muscle we treat as TPb3-UP4.Whatever the homology may be, we do not believeour OD4 is a homologue of a transversus dorsalis,which we consider to be a muscle joining the twosides of the gill arches. A vertically oriented OD4 isknown only in albulids.OP dorsally on posterior surface of Eb4 medial touncinate process, where it overlaps M. UP5-Cb4,ventrolaterally joining ER, below which Ad5 and SOappear to be confluent.OP' slender, originating on dorsoposterior bonysurface of Eb4 uncinate process (attachment coveredby OD4) and inserting tendinously at ER.Remarks. Although present on both sides of ourspecimen, verification in other specimens is needed.M. UP4-Eb2 origin on UP4 continuous with me-dial end of insertion of LI3; insertion on cartilaginouscap of Eb2 dorsomedial (uncinate?) process.Remarks. Similar muscle, M. Pb4-Eb2 present inAldrovandia affinis (Halosauridae).M. UP5-Cb4 dorsally with some fibers attachingto Eb4, but main portion of muscle continuing an-teriorly ventral to Eb4 and attaching to dorsal surface
NUMBER 1
1
35
of UP5; posteroventrally attaching to anterior surfaceof fingerlike (fused Eb5) posterior cartilaginous endof Cb4.Remarks. M. UP5-Cb4 is known elsewhere onlyin Pterothrissus.Ad 1-3 absent.Ad4 large, prominent, on posterior surface of Eb4levator process and posterior end of Cb4.Ad5 undifferentiated medially from SO, joineddorsomedially to ER; attaches dorsolaterally to car-tilaginous rod-like posterodistal end of Cb4 and ven-trally along lateral half of posterior surface of Cb5.Remarks. Albula lacks Eb5; however, the confor-mation of the cartilaginous distal end of Cb4 appearssimilar to the combined distal end of Cb4 and Eb5in Pterothrissus, presumably indicating fusion ofthese two elements in Albula. The possibility of suchfusion is indicated by our specimen of the anguillo-morph Notacanthus (Notacanthidae). in which Eb5 isautogenous on one side and, based on appearance,fused with Eb4 on the other.SO longitudinal layer appears to be restricted, atleast dorsally, to isolated area ventral to TPb3 (notpresent immediately posterior to TPb3).RD absent.Additional remarks. SCL attaches mid-dorsally tosmall, autogenous cartilaginous ball, which attachesin turn to ventral surface of cartilaginous tip of pos-terior end of Bb3 (anterior end of RecV4 attaches toSCL; ObV3 attaches medially only to Hb3 flangecontinuous with SCL). A relatively larger, somewhatcone-shaped autogenous cartilage is present in asmaller cleared-and-stained specimen examined (au-togenous cartilaginous element attached to ventro-posterior tip of Bb3 is known otherwise only in someacanthomorphs, e.g., the melamphaid Poromitra cap-ito, q.v., which lacks SCL). The autogenous cartilagewas not mentioned by Nelson (1969a. fig. 7a). TV4is free from Cb5s, but is attached to ventral surfaceof Bb4. Pbl is oriented horizontally?in line withEbl. SPb2 is absent; cartilaginous SPbl articulateswith medial end of Ebl (as opposed to ossified SPblin Elops and Megalops, both of which have cartilag-inous SPb2s).
Pterothrissus gissu Hilgendorf, FRSKU 22120, 132mm. Plate 18
Description.LEI on Ebl mid-dorsoposteriorly and on lateralsurface of SPbl dorsally ventral to origin of muscle.LE2 on anterior surface of Eb2 uncinate process,originates by long tendon (all other levators originatemusculously).LE3 on anterior surface of Eb3 uncinate process.
LE4 on dorsal cartilaginous tip of Eb4 levator pro-cess.LP absent.LI1 on dorsal surface of Pb2 and medial surfaceof SPbl.Remarks. See remarks under LI1 in Aldrovandia(Halosauridae) description.LI2 on posteromedial surface of Pb3 ventral toOD4.LI3 on dorsolateral edges of UP4 and UP5.TD comprises TPb3 and TPb4. TPb3 divided intoanterior and posterior sections by OD3, which orig-inates on Pb3 ventral to anterior section; anterior sec-tion attaches to anterolateralmost surface of Pb3 un-cinate process; posterior section attaches to uncinateprocess just ventral to anterior section and is contin-uous posteriorly by diagonal muscle strand withTPb4. TPb4 attaches to dorsomedial cartilaginousedge of Pb4 along line with OD4 origin.OD3 origin on dorsoanterior bony Pb3 surface be-low TPb3 anterior section; insertion on dorsoanter-iormost edge of tip of Eb3 uncinate process.OD4 large, vertically oriented; anteromedially,muscle originates continuously along bony dorso-medial margin of Pb3 and medial margin of Pb4 atjunction of cartilage and bony UP4; dorsolaterally,muscle divides into two sections: posterior sectioninserts on Eb4 uncinate process and small AC at tipof process, anterior section extends posteriorly pass-ing ventral to insertion of posterior section and in-
serts broadly on anterior surface of Eb4 levator pro-cess.Remarks. A vertically oriented OD4 is known onlyin albulids.OP dorsally on posteromedial surface of Eb4, ven-trally ending at ER.M. UP5-Cb4 dorsally with some fibers attachingto Eb4, but main portion of muscle continuing an-teriorly ventral to Eb4 and attaching to dorsal surfaceof UP5; ventrally attaching broadly to posterodistalend of Cb4 with minor attachment to Eb5 medialsurface.Remarks. M. UP5-Cb4 is known elsewhere onlyin Albula.Adl-3 absent.Ad4 large, prominent, dorsally on posterior surfaceof Eb4 levator process, ventrally on Cb4 dorsopos-terior surface anterior to inner angle formed by Eb4-Cb4 joint.Ad5 on Cb4 posterodistal end, wraps mediallyaround distal end of Cb5 and attaches to Cb5 pos-teroventral surface beginning at about mid-length,joined dorsomedially to ER and ventromedially byraphe with TV5.SO longitudinal muscle layer attaches anteriorly toPb3.RD absent.
36 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Additional remarks. SCL attached mid-dorsally toposteroventral cartilaginous tip of Bb3. TV4 freefrom Cb5s, but mid-dorsal surface attaches to Bb4ventral surface. Small, autogenous ball of cartilage(AC) present between dorsalmost cartilaginous tipsof Eb3 and Eb4 uncinate processes. Another AC atinner angle formed by Ebl-Cbl joints, supports gillraker at joint. Large, more-or-less vertically orientedEb5 attaches to posteromedialmost tip of Cb4. Eb5appears to be represented by non-autogenous process
at posteromedialmost end of Cb4 in Albula. Pbl car-tilaginous (ossified in Albula).NOTACANTHIDAE
Notacanthus chemnitzi Bloch, USNM 214342, ca.340 mm. Plate 19
Description.LEI on mid-posterior edge of Ebl just lateral touncinate process.Remarks. Slender ligament, originating on skull,inserts on medial end of Ebl; similar ligament insertson medial end of Pb2 in related Aldrovandia (Halo-sauridae).LE2 on Eb2 uncinate process.LE3 on cartilaginous tip of Eb3 uncinate process;muscle present and well developed only on left sideof illustrated specimen (but presence indicated in il-lustration based on USNM 44246, examined in situ,in which LE3 is present on both sides).LE 4 absent (both specimens).LP absent.LI1 on dorsal surface of Pb2.LI2 on dorsal surface of posterolateral cartilagi-nous process of Pb3.LI3 absent (Pb4 absent).TD comprises three continuous portions that aredemarcated only laterally: TEb2, TPb3, and TEb4.TEb2 attaches to anterior surface of medialmost endof Eb2, dividing laterally as it passes over anterolat-eral cartilaginous process of Pb3, to which a fewstrands of muscle also attach; TPb3 attaches to dor-solateral surface of Pb3; TEb4 attaches to postero-medial edge of Eb4 and is broadly continuous pos-teriorly with SO.OD3 absent.OD4 with split origin: on Pb3 dorsally ventral toTEb2, and on tip of Eb2 uncinate process; originsfuse posteriorly and insert on Eb4 dorsodistally.Remarks. Origin on Eb2 is unique.OP distinct, ribbon-like, originating dorsally onposteromedial surface of Eb4 and extending poster-oventrally, then curving anteriorly and joining con-tralateral OP in medial raphe, anterior point ofwhich joins medial raphe of TV5. As such, OP does
not attach ventrally to a gill-arch element. ER isabsent.Ad 1-3 present. Adl relatively large, inseparablefrom gill-filament muscle, which appears to overlieit dorsolaterally, but is continuous with it ventrome-dially. Ad2 and 3 well developed, with well-devel-oped gill-filament muscle portions dorsoanteriorly.Lateral portions of gill-filament muscles of Ad2 and3 were destroyed during dissection and their full ex-tent is unknown (see also remarks under Ad 1-3 inOncorhynchus (Salmoniformes).Ad4 relatively large, dorsally on long dorsoposter-ior edge of Eb4, ventrally on much of posterodorsalsurface of Cb4.Ad4', small (smaller than Ad3), vertical muscleband, dorsally on anterodistal surface of Eb4, ven-trally on bony anterolateral surface of Cb4; appar-ently not associated with gill-filament muscle.Remarks. Ad4' appears to be an autapomorphy andwas not observed elsewhere in this study. It differsfrom Ad4 in attaching to the anterior surface of Cb4,whereas Ad4 in pre-acanfhomorphs, except Poly-odon, attaches to the dorsoposterior surface of Cb4just medial to internal angle formed byEb4-Cb4 joint.In Polyodon, Ad4 attaches to posterior surface of Eb4and anterior surface of Cb4.Ad5 enveloped in tough CT (removed in Plate 19);on left side attaches dorsally to Eb5, which is at-tached to distalmost surfaces of Eb4 and Cb4; on
right side attaches dorsally to cartilaginous Eb4 pro-cess that has shape and position of separate Eb5 onleft side; relatively long and free as it wraps aroundCb5 and attaches well medially on Cb5.Remarks. In a cleared and stained specimen(USNM 214339), Eb5 is present on both sides and itis equally associated with the cartilaginous ends ofEb4 and Cb4. Cb5 bears simple gill rakers but noteeth.RD absent.SO longitudinal muscle layer present (not illus-trated), spongy, extends anteriorly beyond horizontalbetween medial ends of Ebls.Additional remarks. SCL absent. TV4 free fromCb5s, but attached to mid-anteroventral surface ofBb4. Pbl absent, apparently replaced functionally byelongate cartilaginous end of Ebl. Pharyngobranchialtoothplates absent. Bbl bears edentulous tooth platemid-dorsally, not reported by McDowell (1973:132)in Notacanthus. Tough, slender ligament on antero-lateralmost edge of anterior cartilaginous end of Ebl(originates on ventral margin of opercular bone Mc-Dowell, 1973:132). Slender ligament (not illustrated)originates on skull separately from clustered originsof LEs and Lis and inserts on Pb3 medial to Pb3origin of OD4.
NUMBER 11 37HALOSAURIDAEAldrovandia affinis (Giinther), USNM 319707, ca.425 mm TL. Plate 20
Description.LEI on base of Ebl uncinate process.LE 2 on anterior surface of Eb2 uncinate processalong and ventral to posterior cartilaginous tip of pro-cess, which also has separate cartilaginous dorsoan-terior tip (see M. Pb4-Eb2).LE3 on cartilaginous tip of Eb3 uncinate process
at attachment of Eb3 branch of OD3-4.LE4 on cartilaginous tip of Eb4 levator process.Remarks. Slender ligament, originating on skull,also inserts on levator process. Similar ligament alsopresent in closely related Notacanthus (Notacanthi-dae), which lacks LE4.LP absent.LI1 inserts over most of dorsoanterior surface ofPb2 and on cartilaginous tip of Ebl uncinate process.Remarks. Long, ribbon-like ligament, originatingon skull, inserts on medial tip of Pb2. Similar liga-ment, inserting, however, on medial end of Ebl, pres-ent in related Notacanthus (Notacanthidae).The unusual partial insertion of LI1 to include themedial end of Ebl may be the result of the loss ofSPbl. In some other elopomorphs (Elops, Megalops,Albula, and Pterothrissus), SPbl is present and at-taches on the dorsomedial end of Ebl (which pre-sumably divides to become the uncinate process), andLI1 expands ventrally to almost completely envelopSPbl in its insertion, which is otherwise on Pb2. Theinsertion on SPbl just misses including the edge ofEbl, so that it would be no great change for LI1 tohave its insertion extended (as in Aldrovandia) to in-clude the tip of the uncinate process if SPbl werelost. Compare LI1 in Aldrovandia with LI1 in Elops,Megalops, Albula, and Pterothrissus.LI2 on dorsolateral edge of Pb3.LI3 broad based, anteriorly on cartilaginous Pb4
at junction of cartilage with UP4, continuing poste-riorly onto edge of UP5, the anterior edge of whichlies under posterior end of Pb4.M. Pb4-UP5-Eb2 origin on posterolateral corner ofPb4 continuous anteriorly with insertion of LI3, pos-teriorly on dorsolateral surface of UP5 (not illustrat-ed); insertion on osseous dorsal edge of Eb2 uncinateprocess.TD comprising continuous TPb3-Pb4-Eb4 with ad-ditional, roughly trapezoidal TPb3 section arisingmid-dorsally from TPb4 area; anterior fibers of sep-arate section, attaching to and wrapping anteriorlyaround cartilage-tipped lateral process of Pb3; TDcompletely undifferentiated posteriorly from SO.OD3-4 origin broadly continuous on posterioredge of Pb3 and anterior end of Pb4, dividing pos-
teriorly with one section inserting on Eb3 uncinateprocess and other section inserting on Eb4 anterolat-
erally.OP dorsally on ventral surface and posteromedialedge of Eb4; ventrally on small Eb5 attached to Cb4and dorsodistal cartilaginous end of Cb4. ER absent.Ad 1-3 absent.Ad4 on Eb4 dorsoposteriorly and dorsoposteriorbony surface of Cb4.Ad5 dorsally on Cb4 distally and tiny Eb5, whicharticulates ventrally with Cb4 and is attached by lig-ament to Eb4 levator process; free ventrally for shortdistance posterior to attachment on mid-ventromedialsurface of Cb5.RD absent.SO longitudinal muscle layer (not illustrated) ex-tends anteriorly at least to horizontal between medialends of Ebls. SO fibers also attach to UP5, which isventral to, but separated by SO and OP muscle fibersfrom medial arm of Eb4.Additional remarks. SCL absent. TV4 free fromCb5s. Ligament connects posterior bony surface ofEb2 uncinate process to anterodistal cartilaginous endof Eb3; another connects posterior bony surface ofEb3 uncinate process to anterodistal cartilaginous endof Eb4; another attaches tip of Eb4 uncinate processto Eb5, which, unusually, attaches to Cb4 rather thanEb4, and continues dorsoposteriorly. Ribbon-like lig-ament inserts on cartilaginous tip of medial processof Pb2; its origin was not recorded, but a similarligament in Notacanthus attaches to the cranium. Pb2lacks tooth plate. CONGRIDAEConger cinereus Riippell, USNM 115969, 345 mm.Plate 21
Description.Remarks. Nelson (1967c) described and illustratedthe gill-arch musculature of Conger marginatus (cur-rently considered a junior synonym of C. cinereus).Insofar as Nelson's and our findings can be com-pared, there is much similarity, but some differencesmentioned below suggest further study is indicated.Although there is a question about the homology ofthe pharyngeal tooth plates, we use Nelson's (1966b)terminology for convenience.LEI tendinously on Ebl mid-dorsoposteriorly.LE2 tendinously on Eb2 mid-dorsoposteriorly.LE3 on cartilaginous tips of joined Eb3 and Eb4uncinate processes.LE4 finely, tendinously on Eb4 mid-dorsally.LP absent.LI1 on cartilaginous medial end of Pb2.LI2 on Pb3 laterally, UP3 dorsolateral^, and UP4dorsoanterolaterally.LI3 absent (Pb4 absent).
38 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONTD thin, sheet-like, overlies SO posteriorly, butscarcely separable from SO, comprises TPb3-Eb4, at-taching dorsally on Pb3 at and along line of attach-ment of OD4, and on Eb4 anteromedially.OD3 absent.OD4 origin on Pb3 medial to LI2 and lateral toline of attachment of TPb3-Eb4, insertion along en-tire length of Eb4 uncinate process.Remarks. The tips of the uncinate processes of Eb3and Eb4 are tightly bound together, and OD4 is pos-sibly minutely attached to the cartilaginous tip of Eb3uncinate process. In contrast to our findings. Nelson(1967c: 349) reported that the superior oblique (= ourOD) in C. marginata attaches Pb3 only to Eb3. If itis determined that the OD of C. cinereus lacks a
"true" connection to Eb3, the lack is autapomorphic.An OD attachment to Eb3 defines the Teleostei, andthere are exceedingly few other pre-acanthomorphtaxa that lack the attachment (i.e., Notacanthus, Gon-ostomo, Maurolicus).RecD2 on ventroanteriormost surface of Eb2 andmid-posterior edge of Ebl.RecD3 on ventroanteriormost surface of Eb3 andmid-posterior edge of Eb2.M. Pb2-Ebl on Pb2 dorsoposteriorly and Ebl me-dial cartilaginous tip.Remarks. Nelson (1967c:349 and fig. 2) termedthis muscle obliquus inferior accessorius.M. UP4-Eb4 on UP4 dorsoposterolaterally, poste-
riorly becoming broad, thin CT sheet, which attachesto ventrolateral margin of Eb4 (at angle of Eb4-Cb4joint) and extends medially, becoming incorporatedin SO.Remarks. Nelson (1967c:349) termed this muscle
"retractor dorsalis." Although appropriately namedfunctionally, the muscle does not originate on thevertebral column and cannot be considered even ho-moplastically as RD. Nelson (1966a: 123), however,reported that [exceptionally] among the several eelshe examined, which also included Conger and An-guilla, but not Synaphobranchus, "RD" in the [spe-cialized] Muraeninae, has become attached second-
arily to the vertebrae.OP strap of muscle attaching dorsally to Eb4 pos-teromedially, slightly distinguished by denser fibersfrom SO medially, ending ventrally at ER, which isjoined ventrally by Ad5; ventromedially partiallyoverlain by SO fibers.Ad 1-3 absent.Ad4 dorsally on Eb4 posteroventral surface andventrally on Cb4 posterodorsal surface.Ad5 joins distal end of Cb5 to bony sub-distal endof Cb4, dorsomedially joining ER.RD absent. See SO.SO thin longitudinal muscle layer surrounds eso-phagous and attaches to toothplates and Pb3.
Additional remarks. SCL absent. TV4 free fromCb5s. Uncinate process present on Eb4. Pbl absent.ANGUILLIDAEAnguilla rostrata (Lesueur), USNM 340815, 228mm, USNM 190998, 328 mm.Plate 22
Description.Remarks. All levators extend lateral to LI1, exceptLE4, which passes medial to LI1. LI1 extends dor-solateral^, all other levators extend anteriorly. Nel-son (1967c) described and illustrated the gill-archmusculature of Anguilla rostrata. There is much sim-ilarity between his and our findings, but a few dif-ferences exist.Although there is a question about the homologyof the pharyngeal tooth plates, we use Nelson's(1966b) terminology for convenience.LEI on dorsal surface of Ebl about half lengthdistally.LE2 on dorsal surface of Eb2 about half lengthdistally.LE3 finely tendinously on dorsoanterior surface ofbony Eb3 uncinate process, just ventral to OD3 in-sertion.LE4 very slender, on Eb4 mid-dorsally just lateralto bony uncinate process, which is tightly joined tobony Eb3 uncinate process.LP absent.LI1 broad, thin, on Pb2 dorsal surface; insertionjust medial to and paralleling M. Pb2-Ebl.LI2 on UP3 dorsolaterally.Remarks. Nelson (1967c:349) did not describe theinsertion of LI2 in either Anguilla or Conger, butreported that the muscles of Anguilla are "rather sim-ilar" to those of Conger. In Conger, LI2 inserts onPb3, UP3, and UP4.LI3 absent (Pb4 absent).TD comprises TPb3-UP3-UP4-Eb4, on medialmargin of Pb3, just barely on posteromedial edge ofUP3, on dorsomedial surface of UP4, continuing un-interrupted posteriorly as SO, and medial tip of Eb4.OD3 origin on dorsoanterior end of Pb3, continu-ous with OD4 origin, insertion on bony Eb3 uncinateprocess dorsal to LE3.Remarks. Nelson (1967c:fig. 4) indicated that OD4attaches to a cartilage tipped Eb4 levator process. Inboth our specimens, the tips of the levator processeson Eb3 and Eb4 are bony and although the two pro-cesses are tightly bound together by CT, OD3 attach-es only to Eb3.OD4 origin on Pb3 dorsoposteriorly, insertion onEb4 anteromedial edge.RecDl very long, on Ebl anterodistal tip, anteri-orly ends tendinously in CT of roof of mouth, par-
allels LEs.
NUMBER 11 39
Remarks. Ordinarily we would consider this mus-cle as LEI ', but its serial position, like that of RecD2and RecD3. supports Nelson's (1967c) assignment(our RecD is the same as Nelson's obliquus inferioraccessorius). With the questionable exception of theosteoglossiform Gymnarchus, we know of no othergenus in which RecDl occurs.RecD2 short, posteriorly on anterodistal end ofEb2, joined along anterior edge by well-developedportion of GFM, on Ebl posterolateral^ posterior toLEI insertion.RecD3, short, on anterodistal end of Eb3, joinedalong anterior edge by well-developed portion ofGFM; on Eb2 posterolateral^ posterior to LE2 in-sertion.M. Pb2-Ebl small, hidden by LI1, anteriorly onmedial tip of Ebl, posteriorly on lateral edge of Pb2,parallels lateral side of LI 1 insertion.OP dorsally on Eb4 dorsomedially, ventrally joinsER with SO and Ad5, not continuous below ER,which extends tendinously laterally and attaches toCb4 distally together with dorsal end of Ad5.Ad 1-3 absent.Ad4 dorsally on posterior edge of Eb4, ventrallyon Cb4 anterior to Eb4-Cb4 joint.Ad5 dorsally on ER with lateral tendinous attach-ment to Cb4 posterodistal end; ventrally on poster-odistal half of Cb5.RD absent. See SO below.SO longitudinal muscle layer surrounds esopha-gous, dorsally fibers attach to tooth plates and medialmargin of Pb3.Remarks. Nelson (1967c) considered this portionof the longitudinal muscle layer as RD.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl absent.SYNAPHOBRANCHIDAESynaphobranchus sp., USNM 316662, ca. 340 mm.Plate 23Remarks. Although there is a question about thehomology of the pharyngeal tooth plates, we use Nel-son's (1966b) terminology for convenience.
Description.LEI broadly on Ebl dorsally beginning at aboutmid-length of bone and extending laterally to smallposterodistal flange-like process.LE2 on small posterodistal Eb2 flange-like pro-cess.LE3 on posterior flange-like Eb3 process at OD3?4 insertion.LE4 on Eb4 dorsally slightly distal to mid-length.LP absent.LI1 on Pb2 anterolaterally.LI2 inserts posteriorly on Pb3 dorsoposteriorly,
UP3 dorsomedially, and on anterodorsal edge ofUP4, and inserts anteriorly on dorsoposterior edge ofEb2.LI3 absent (Pb4 absent).TD comprises undifferentiated layer of muscle,TPb3-UP3-UP4, attaching laterally to Pb3, UP3 dor-somedially, and on most of dorsal surface of UP4.OD3-4 origin on anterior end of Pb3, insertion onEb3 dorsally at and posterior to insertion of LE3 andon anteromedial edge of Eb4 beginning well lateralto joint with Pb3 and ending just proximal to pointopposite insertion of LE3.M. Pb2-Eb2 attached to ventral surfaces of Pb2and anterior end of Eb2.RecD2 on proximal two-thirds of anterior edge ofEb2 and on dorsomedial tip of Ebl.OP questionably present as strap of muscle attach-ing dorsally to posteromedial surface of UP4 andending ventrally at ER, which is at level of Cb4 andextends laterally as slender tendon to posterodistalend of Cb5; inseparable medially, and ventral to ER,from SO.Remarks. The shift of the dorsal attachment of OPfrom Eb4 to UP4 is unique among the taxa we ex-amined. Nelson (1967c) considered this muscle sec-tion as RD.Ad 1-3 absent.Ad4, very large, dorsally on most of Eb4 posteriorsurface, ventrally on Cb4 dorsal surface anterior tointernal angle formed by Eb4-Cb4 joint.Ad5 absent.RD absent. See SO.SO thin, longitudinal muscle layer extending an-teriorly to horizontal between anterior tips of Pb3s,surrounding esophagous, attaching to toothplates andPb3.Additional remarks. SCL absent. TV4 free fromCb5s. Pb2 reduced, cartilaginous. Pbl absent.ClupeomorphaDENTICIPITIDAE
Denticeps clupeoides Clausen, USNM 358795, 36.4mm, and BMNH uncataloged, 2 specimens. 30.0-30.6 mm. Plate 24
Description.Remarks. All levators except LI1 extend anteriorly,horizontally, at almost 180? angle and form a thin,posteriorly concave fan of overlapping muscles thatis closely applied to, and originates on, the convexsurface of the otic bulla. LI1 extends laterally atabout 90? angle from other levators, is medially con-cave, and conforms with the laterally convex surfaceof the otic bulla on which it originates.
40 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LEI on minute cartilaginous tip of poorly devel-oped Ebl uncinate process.LE2 on minute cartilaginous tip of poorly devel-oped Eb2 uncinate process.LE3 on minute cartilaginous tip of well-developedEb3 uncinate process.LE4 on cartilaginous dorsomedial end of Eb4 un-cinate process, there meeting OD4 insertion.LP absent.Remarks. Greenwood and Lauder (1981:226) wereuncertain that LP was absent in Denticeps because ofthe small size (ca. 50 mm SL) of their specimens.We agree that size was a problem for dissection andillustration, but are confident that LP is absent.LI1 on Pb2 posterolaterally (see also remarks atbeginning of description above).LI2 on Pb3 posterolaterally.LI3 on cartilaginous Pb4 (not illustrated) postero-laterally.TD comprises TEb2 and TPb3-Pb4-Eb4. TEb2 at-taches to Eb2 dorsomedially. TPb3-Pb4-Eb4 on Pb3and Pb4 at and along part of OD4 origin, becomingwider posteriorly and attaching on dorsal surface ofEb4 medial to uncinate process.OD3 origin on Pb3 dorsal surface ventral to TEb2,continuous with OD4 origin, and insertion on Eb3uncinate process.OD4 originates on Pb3 dorsoposterior surface ven-tral to TEb2, and on Pb4 dorsal surface at and medialto Pb4 portion of TPb3-Pb4-Eb4, and inserts on an-teromedialmost surface of Eb4 uncinate process.OP slender strap of muscle, dorsally on postero-medial edge of Eb4 uncinate process, overlappingmuch of Ad4 posterior surface, ending ventrally atER, which is dorsal termination of major portion ofAd5 (unless continuation is interpreted as OP fusedlaterally with AD5).Ad 1-3 absent.Ad4 dorsally on dorsoposterior edge of Eb4 un-cinate process continuing to distal end of Eb4, ven-trally on Cb4 dorsal edge anterior to internal angleformed by Eb4-Cb4 joint.Ad5 dorsally on posterior surface of Eb5 and atER ventral to OP, medially continuous with SO, ven-trally on Cb5 posterior surface.SO longitudinal muscle layer thickest dorsally andventrally; anterior extent undetermined.RDs absent.Additional remarks. SCL absent. TV4 free fromCb5s. Eb5 present (not previously reported, althoughpossibly implied by discussions in Johnson and Pat-terson, 1996). A single large UP present.
PRISTIGASTERIDAE
llisha africana (Bloch), USNM 357405, 120 mm.Plate 25
Description.LEI slender, weak, on dorsal bony edge of Ebluncinate process just distal to cartilaginous tip.LE2 slender, weak, on dorsal bony edge of Eb2uncinate process just distal to cartilaginous tip.LE3 on tip of all bony Eb3 uncinate process, atand anteromedial to LE3' insertion, larger and angleddorsoanteriorly about 10? lower than LE3'.LE3' on tip of all bony Eb3 uncinate process atand posterior to LE3 insertion, slenderer and angleddorsoanteriorly about 10? higher than LE3.LE4 broad, thin, on bony dorsoposterior edge ofEb4* coincident with posterior edge of OD4 inser-tion; questionably comprising two very thin sections,anterior and posterior, on right side, but not on left
side.LP broad, thin, on dorsodistal cartilaginous edgeof Eb4*, commencing at lateralmost edge of LE4 in-sertion, partially coincident with Ad4 dorsal attach-ment.LI1 on Pb2 dorsal surface medial to uncinate pro-cess.LI2 on Pb3 dorsoposteriorly.LI3 on Pb4 (not illustrated) dorsoposteriorly.TD comprises TPb3-Pb4-Eb4*, slightly partitionedlaterally at posterolateral origin of OD4 on Pb3 and
at attachment to medial end of Eb4*; on right sideonly, joining ER with OP ventrally; continuation pos-teriorly with SO marked by change in muscle fiberdirection.OD3 origin on Pb3 dorsal surface anterior to un-cinate process and on medial edge of uncinate pro-cess; insertion on dorsomedial edge of Eb3 bony un-cinate process.OD4 origin begins on Pb3 among TD fibers, pass-ing dorsal to most of Pb3 attachment of TD, andcontinuing on Pb4 dorsoanteriorly; insertion broadlydorsally on Eb4* anterior surface ventral to LE4 in-sertion.OP strap of muscle originating on Eb4* postero-medial to LE4 insertion, and extending ventrally toER; undifferentiated from SO ventral to ER.Ad 1-3 absent.Ad4 dorsoposteriorly on Eb4*, ventrally on Cb4
at anterior angle formed by Eb4*-Cb4 joint.Ad5 dorsally on distal end of Eb4*, ventrally onCb5 distal end, inseparable mid-medially from SO.RD absent.SO longitudinal muscle layer spongy, circumeso-phageal, thickest dorsally, extending dorsoanteriorlyto below anterior end of TD, but becoming extremelyattenuated anterior to horizontal between medial endsof Eb4*s.Additional remarks. SCL absent. TV4 free fromCb5s, attached dorsally to ventral surface of Bb3.Tiny MSPbl anterior to anterior end of each Ebl.GC attached ventrally to anterior ends of Pb2s, which
NUMBER 11 41
are slightly dorsal to anterior ends of Pb3s (see alsoAdditional remarks under Cetengraulis, and Chanos).ENGRAULIDAECetengraulis edentuhis (Cuvier), USNM 186377, 2specimens, 118-124 mm.Plates 26.1, 26.2
Description.Remarks. All levators except LEI pass medial toLI1. LE2-4 extend horizontally anteriorly at approx-imately 180? from insertions, LE4' extends dorsoan-teriorly about 15? from insertion, LI is more-or-less
vertical, and LI2 and LI3 are angled dorsoanteriorlyabout 45?.LEI thin, slender, on bony process at distal edgeof base of Ebl uncinate process; originates somewhatmore anteriorly on skull than other LEs.LE2 on tip of Eb2 uncinate process.LE3 tendinously on Eb3 uncinate process ventralto OD3 insertion.LE4 on medial edge of Eb4* levator process dorsalto insertion of OD4; originates as long tendon.LP slender, membranously attached along lateraledge of LE4, inserting with LE4 insertion; originatesas very long, slender tendon.LI1 broad, thin, convex medially (medial surfaceconforming with surface of cranium), insertingbroadly on dorsal surface of Pb2.LI2 on posterior end of Pb3 dorsal surface.LI3 inserting as long, slender tendon on medialedge of Pb4 dorsoanteriorly.TD comprises TPb3, TPb4, and TEb4*. TPb3 sep-arate muscle attaching to dorsolateral edge of Pb3beginning just posterior to base of uncinate process,continuous posteriorly by diagonal strand of muscleinserting on Pb4 anteriorly in one specimen and atmid-length on other. Anteriorly, TPb4 comprisesloose strands of fibers, which attach on each Pb4 dor-somedially, and is continuous posteriorly with slen-der TEb4*, which attaches to ventromedial ends ofEb4*s and is continuous posteriorly with SO+ EO.OD3 slender, originates on Pb3 uncinate processventromedial to dorsal tip and inserts tendinously onEb3 uncinate process just ventral to tip and dorsal toLE3.OD4 very weak, almost thread-like, originates byshort, fine tendon attached to tiny (almost pinpoint)bony area on dorsomedial edge of otherwise cartilag-inous Pb4; inserts on ventromedial edge of Eb4*.OP unclear if absent or included in complex sur-face muscles on EO.Ad 1-3 absent.Ad4 two widely separated dorsal attachments pos-teriorly on Eb4* levator process, uniting ventrally inslender attachment (not visible in lateral view) onCb4 at innermost angle formed by Eb4*-Cb4 joint.
Ad5 questionably included in complex surfacemuscles arising from EO and Cb5 and attaching an-terolaterally to Eb4*.Remarks. There is a posteroventral, rod-like car-tilaginous continuation of the dorsodistal end of Eb4*that articulates ventrally with the distal end of Cb5.Nelson (1967d:fig. 2j) indicated that this extensionrepresents a partially fused Eb5 in Engraulis and weagree.RDs absent.SO longitudinal muscle layer spongy, circumeso-phageal, thickest dorsally, becoming extremely atten-uated anterior to EO, and extending between UP4sto below anterior end of TPb4.Additional remarks. SCL absent. TV4 anterior toCb5s (condition relative to Bb complex not exam-ined). Elongate anterior cartilaginous tips of Ebl s notsegmented as in Coilia. UP4 edentate, extending dor-sally as thin plate. Short, cylindrical muscle-like lig-ament joining ventroposterior surface of Ebl unci-nate process to ventrolateral surface of Pb2 uncinateprocess. Another similar ligament joining ventropos-terior surface of Eb2 uncinate process to ventrolateralsurface of Pb3 uncinate process.The area between Pb2s and anterior ends of Pb3sis covered by a leathery CT sheet. The sheet attachesto the dorsalmost ends of the Pb2 uncinate processes,but the main part of the sheet lies well ventral to thedorsalmost ends, at about the level of the anteriorends of the Pb3s. The sheet apparently conforms withthe ventral surface of the cranium. Mid-ventrally, thesheet incorporates a small, seed-shaped cartilage,which Di Dario (2002:500) identifies as GC, an ele-ment he first named and described in a printed jour-nal (but see GC in Abbreviations and Definitions sec-tion): "[the engrauloid] Cetengraulis . . . has a typi-cal gongyloid cartilage in all aspects [our italics] ex-cept that it is markedly rod-shaped . . . this shape ishypothesized as convergent to that of the [clupeidsecond] mediopharyngobranchial," which Di Dariofound in three of the 20 clupeid genera he examined.Di Dario followed up by allowing that in the "ab-sence of simultaneous occurrence of the gongyloidcartilage and the second mediopharyngobranchial inany individual, the possibility of their homology can-not be conclusively discarded." We find that the pu-tative GC in our specimen of Cetengraulis (if present,it was destroyed during dissection in the other spec-imen) is quite dissimilar in shape, relative size, po-
sition, and in being incorporated in a CT sheet, fromthat of other clupeoid taxa (engrauloids, pristigaster-oids) as described and illustrated by Di Dario andthat we have examined. Because most (8 of 10) ofthe other engrauloids Di Dario (2002: table 1) ex-amined have a GC that is typical and the other twolack GC, we think that the element in question inCetengraulis is probably a modified (reduced) GC,
42 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
intermediate between character states of "full'velopment and complete absence. de-
Coilia neglecta Whitehead, USNM 357380, 2 spec-imens, 138-161 mm.Plate 27
Description.Remarks. All levators besides LI1 pass medial toLI1. LI1 is more-or-less vertical, following aroundthe convex surface of the prootic. LI2 and LI3 passdorsoanteriorly at about 60? and 45? angles and are
"laminated" against the dorsomedial surface of LI1.LE2-4 extend anteriorly at almost 180? from theirinsertions.LEI absent.LE2, very fine, narrowly on bony Eb2 uncinateprocess just distal to medial head of Eb2, which ar-ticulates with Pb2 uncinate process.LE3 on dorsoanterior surface of Eb3 uncinate pro-cess just ventral to cartilaginous tip.LE4 narrowly on dorsalmost edge of Eb4* justdorsal to OD4 insertion.LP absent.LI1 very broad, thin, on Pb2 dorsomedial margin,which is tightly bound and ventral to Pb3 slenderanterior process.LI2 thin, on Pb3 (not shown) dorsal surface pos-terolaterally.LI3 thin, on Pb4 dorsolaterally.TD comprises TPb3, TPb4-Eb3, TEb4*. TPb3,most distinct of the three, is broadly on Pb3 postero-laterally anteroventral to OD3 origin, and is posteri-orly dorsal to OD4 origin and continuous by loosediagonal strands of muscle with TPb4-Eb3. TPb4-Eb3 is on Pb4 dorsolaterally just medial to LI3 in-sertion and medial end of Eb3 (in smaller specimen,there is slight separation of Pb4 and Eb3 sections,hence these could be accorded separate names).TPb4-Eb3 is broadly continuous posteriorly withTEb4*, which is on Eb4* dorsalmost edge ventral toOD4 insertion.OD3 on bony dorsal surface of Pb3 uncinate pro-cess, extending a little anteriorly onto slender anteriorprocess of Pb3, and is anteriorly dorsal to TPb3 at-tachment; insertion on dorsomedial edge of Eb3 un-cinate process.OD4 origin mainly on Pb3 dorsoposteriorly, withfew strands on dorsoanteriormost surface of Pb4; in-sertion on dorsomedialmost edge of Eb4* ventral toLE4 insertion.OP consists of strands of muscle attaching dorsallyto Eb4* posteromedial surface, ventrally joining ER
at mid-level of cartilaginous distal end of Eb4*; ven-tral to ER, apparently comprises Ad5 and SO.Ad 1-3 absent.
Ad4 on Eb4* dorsoposteriorly lateral to OP, dor-sally with broad medial section separated by spacefrom slender lateral section (nerve passes throughspace), ventrally on Cb4 dorsal surface medial toEb4*-Cb4 joint.Ad5 continuous medially with SO, dorsolaterallyon distal cartilaginous end of Eb4*, ventrally on pos-terodistal end of Cb5, medially not separable fromSO.RD absent.SO longitudinal muscle layer thin, spongy, circu-mesophageal, evenly distributed, extending dorsoan-teriorly much attenuated at least to anterior end ofTPb3 (possibly continuing over CT anteriorly).Additional remarks. SCL absent. TV4 free fromCb5s, but slight attachment dorsally to ventral sur-face of cartilaginous Bb copula. There is an autoge-nous cartilage, (Plate 27A, MPbl) anterior to carti-laginous anterior tip of each Ebl (see additional re-marks under Dussumieria, Clupeidae). Ventral end ofPbl attaches to dorsal surface of this cartilage. GCabsent. CHIROCENTRIDAEChirocentrus dorab (Forsskal), USNM 283241, 195mm; USNM 283242, 144 mm.Plates 28.1, 28.2
Description.Remarks. All levators except LEI and the musclesheath termed LP, extend medial to LI1. LI1 is angleddorsoanteriorly at about 50-60?; the LP sheath com-prises a perpendicular pyramidal section overlying ormeshed with a horizontal section; the other levatorsare all angled horizontally at about 180?.LEI on tip of Ebl uncinate process.LE2 on tip of Eb2 uncinate process.LE3 on medial edge of distal tip of Eb3 uncinateprocess.LE4 on dorsomedialmost edge of Eb4* levatorprocess.LP thin, fanlike sheet of muscle fibers attachedalong much of lateral edge of LE4 and dorsomedialedge of Eb4*, dorsalmost fibers almost perpendicularto those of LE4.Remarks. Greenwood and Lauder (1981:226) be-lieved LP was absent in C. dorab "unless, atypically,it is closely associated with the 4th external levator
. .
." We find LP is typically closely associated withLE4 in clupeoids and characoids.LI1 posteromedially on Pb2 dorsal surface.LI2 absent.LI3 on lateral edge of Pb4 lateral to TPb4; appearsto have two slightly separate origins.TD comprises three parts: TPb3, TPb4, TEb4*.TPb3 on Pb3 uncinate process and Pb3 dorsolaterallyposterior to uncinate process, broadest centrally, dor-
NUMBER 1
1
43
sal to OD4 origin, continuous posteriorly by fewmuscle strands with TPb4 in smaller specimen, notcontinuous in larger specimen. TPb4 broad dorsally,narrow at attachment to Pb4 just medial to LI3 in-sertion, continuous by diagonal muscle strap withTEb4*. TEb4* on Eb4* medially ventral to OP, con-tinuous with SO posteriorly, but distinguishable fromSO by change in direction of muscle fibers.OD3 very small, slender, origin on Pb3 uncinateprocess just lateral to TPb3 attachment, insertion onposterior surface of Eb3 uncinate process ventral toLE3 insertion.OD4 massive, origin beginning on broad dorso-posterior bony surface of Pb3 and extending ontodorsoanterior surface of Pb4, insertion on anterome-dial surface of Eb4* levator process, joining raphewith OP.OP a muscle strap and filaments attaching poste-
riorly on dorsomedial surface of Eb4* levator processand medial arm of Eb4*. joins raphe dorsally withOD4, inserting ventromedially among SO fibers andventrolaterally on posterodistal surface of Eb4*.Ad 1-3 absent.Ad4 dorsomedially on posterior surface of Eb4*levator process, ventrally narrowly on Cb4 dorsalsurface just medial to inner angle formed by Eb4*-Cb4 joint.Ad5 dorsally, narrowly on Eb4* ventrodistally;
ventrally, broadly on posterior surface of Cb5; com-pletely separate from SO medially.RD absent.SO longitudinal muscle layer circumesophageal,evenly distributed, extending dorsoanteriorly as sheetto below TPb4. thence as sparse filaments to belowposterior end of TPb3.Additional remarks. SCL absent. TV4 free fromCb5s. ER absent (a chirocentrid apomorphy?).CLUPEIDAE
Dussumieria acuta Valenciennes, USNM 296827, 3specimens, 121?123 mm.Plate 29
Additional material.
?
= Clupea harengus Linnaeus,USNM 325930, 106 mm; USNM 349799, 112mm.
Description.LEI on cartilaginous tip of Ebl uncinate process.LE2 on cartilaginous tip of Eb2 uncinate process.LE3 on cartilaginous tip of Eb3 uncinate processtogether with LE3' insertion, angled dorsoanteriorlysimilarly to LE2, attaching to cranium near LE2 at-tachment.LE3' inserting on cartilaginous tip of Eb3 uncinateprocess together with LE3 insertion, almost horizon-
tal, extending anteriorly medial to LE1-3, attachingto cranium near cranial attachment of Pbl.LE4 narrowly inserted on dorsoanteriormost edgeof broad Eb4 levator process, continuous ventrolat-erally with abruptly thinner, sheet-like LP, which isalso distinguishable by abrupt change in inclinationof muscle fibers.
? On Eb4*; gradual change in mus-cle fiber inclination between LE4 and continuous,thin LP.LP thin, sheetlike, inserted broadly along dorsal-most edge of Eb4 levator process (lateral to insertionof LE4, but posterior given orientation of Eb4) andEb5, continuous anteroventrally with LE4, but distin-guishable by thinness and abrupt change in inclina-tion of muscle fibers.
? broadly on Eb4*, continuouswith LE4 along most of lateral edge of LE4, mainlydistinguishable from LE4 by abrupt thinness of mus-cle-fiber sheet.Remarks. Winterbottom (1974b:252) discussedLE4 in clupeids concluding that it was a "mootpoint" whether the sheet of thin muscle continuingposteriorly from LE4 could be interpreted solely asLE4 or as LP. If not as LP, he believed that LP couldhave evolved from a clupeid-like condition. Winter-bottom (1974b:fig. 24) provided a generalized illus-tration of the levators in Clupea harnengus and didnot differentiate an LP. Greenwood and Lauder(1981:215) disagreed with Winterbottom's interpre-tation of LE4 in clupeids. ". . . we would identify theusually thin, sheet-like but somewhat expanded mus-cle lying ventral to [the protractor pectoralis] as thelevator posterior muscle and not, as he does, a musclecomposed entirely of the expanded 4th levator exter-nus . . . even in Clupea harengus."' Greenwood andLauder believed that, "Winterbottom included theposterior levator, the 4th levator externus, and somenon-muscular tissue lying above and between thesemuscles, in the muscle he identified as the 4th levatorexternus." Our observations on Dussumieria acutasupport Greenwood and Lauder's general interpreta-tion of LE4 and LP in clupeids, but appear to supportWinterbottom's for C. harengus, in which the differ-entiation, other than a change in thickness, is unap-parent.Among pre-acanfhomorphs, LP (in any form) ispresent only in otocephalans, and the muscle cannotbe considered homologous with that of acantho-morphs.LI1 on Pb2 dorsal surface posteromedially. ? Onmedial margin of Pb2.LI2 on posterodorsal surface of Pb3, attaching toEb2 uncinate process as muscle extends anterodor-sally to origin; free from Pbl; joined by raphe withLI3 on Eb2 uncinate process (see LI3). ? At origin,attaches jointly to Pbl and cranium; completely freefrom Eb2 and LI3.LI3 on Pb4 dorsolaterally and dorsal edge of UP4
44 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(not illustrated), fans out broadly anterodorsally, mid-anteriorly forms partial raphe with LI2 at attachmentto Eb2 uncinate process. ? On dorsal surface of car-tilaginous Pb4 (UP4 absent) at and medial to TD at-tachment to Pb4.TD comprises TPb3-Pb4 and TEb4. TPb3-Pb4, ex-tensive, begins anteriorly at about mid-length ofPb3s, continues posteriorly along Pb3s and Pb4s me-dial to, and continuous by raphe with, OD3 and OD4origins, continuous posteriorly by diagonal musclestrands with TEb4. TEb4 on Eb4 dorsomedially, con-tinuous posteriorly with SO by diagonal musclestrands.
? TD comprises TPb3 and TPb4-Eb4*. TPb3
short, begins just posterior to Pb3 uncinate process(well posterior to OD3origin), overlaps OD4 origin,posteriorly continuous by sparse muscle fibers withTPb4-Eb4*. TPb4-Eb4* attaches to Pb4 posterolat-erally and ventromedial edge of Eb4* levator pro-cess, undifferentiated posteriorly from SO.OD3 short, origin on Pb3 uncinate process (whichjoins Eb2 uncinate process), insertion on Eb3 unci-nate process ventral to insertions of LE3 and LE3'.
? Long, origin on Pb3 anterior to uncinate process,fibers attach to Eb2 uncinate process as muscle pass-es posteriorly to insertion on Eb3 uncinate process.OD4 origin on lateral surface of Pb3, mediallyjoining raphe with TPb3-Pb4, faning out posteriorlyand inserting on dorsoanterior surface of Eb4 levatorprocess, obscures insertion of OD4' on Eb4. ? Originon posterolateralmost edge of Pb3, insertion on an-terodorsalmost medial edge of Eb4* levator process;raphe absent.OD4' origin at junction of Pb4 and UP4, extendsposteriorly mostly ventral to OD4, fans out posteri-orly and inserts along broad anterior surface of Eb4levator process medial to OD4 insertion. ? Origin onPb4 dorsoposteriorly, extends posteriorly (withoutfanning out) ventral to OD4 and inserts on antero-medial edge of Eb4* levator process just ventral toOD4; OD4 and OD4' more or less fused at Eb4*.OP (not labelled), questionable area of muscle,continuous with SO, originating on Eb4 posterome-dially ventral to TEb4 and joining ER posterolater-
ally. ? Same as Dussumieria, but substitute Eb4* andTEb4*; even less distinguishable in Clupea).Ad 1-3 absent.Ad4 dorsally on dorsoposterior edge of Eb4 leva-tor process and ventrally on Cb4, forming anteriorwall of "pocket" separating more-or-less vertical, an-terolateral continuation of SO-OP complex (see OPabove), which forms posterior wall of pocket (notillustrated).
? Substitute Eb4* levator process.Ad5 forms slender tendinous attachment dorsallyto large, cartilaginous Eb5; attaches musculously toCb5 di.stalm.ost end; undifferentiated medially from
SO ventral to ER. ? Musculously attached dorsallyto Eb4* distal end.SO with conspicuous posteromedially curving ERon each side, sheet of CT (not illustrated) arises fromentire length of ER and extends posteriorly (attach-ment not traced). Longitudinal muscle layer beginsas isolated dorsal patch at about horizontal throughposterior ends of ERs (longitudinal fibers absent pos-terior to patch) and extends anteriorly to below TEb4and attaches to mid-medial side of Pb4. ? ER veryreduced, present on only one side of one of two spec-imens examined.RD absent.Additional remarks. SCL absent. TV4 free fromCb5, but attached mid-dorsally to Bb3. Small, un-paired, roughly U-shaped, non-staining, questionablycartilaginous pad attached to anterior ends of Pb2sdorsally. In attachments and position, U-shaped padis faintly similar to GC of non-clupeid clupeoids.MPbl absent. Large Eb5 present, articulating dorsal-ly with dorsodistal end of Eb4, and ventrally withsmall ventrodistal end of Eb4 and broad distal endof Cb4.
?
U-shaped pad absent; small, cartilaginous MPblpresent just anterior to anterior tip of each Ebl(MPbl reported as absent in Clupea by Nelson,1967b:391, who, p. 392, implied that the unpairedMPbl- anterior to the tips of Ebl s in many clupeids,may have originated as segmentation of the cartilag-inous tips of Ebls). Eb5 absent or, by comparisonwith Dussumieria, fused with Eb4* dorsodistal end.Nelson (1967d:fig. 2b) illustrated Eb4* of Clupeaharengus and labeled the dorsally continuous, butventrally separate, posterior cartilaginous end of Eb4as Eb5. He (1967b:fig. 2d) illustrated the Eb5 portionof the distal end of Eb4 in C. harengus as fused ven-trally and separate dorsally and did not label it asseparate from the remainder of Eb4. On both sidesof both our specimens of Clupea, the ventral end isfused indistinguishably with the remainder of Eb4.GonorynchiformesCHANIDAEChanos chanos (Forsskal), USNM 173572, 140 mm,USNM 347538, 63.3 mm.Plate 30
Description.LEI on tip of Ebl uncinate process.LE2 on tip of Eb2 uncinate process.LE3 absent.LE4 originating as long tendon among other LEorigins, passing medial to all other LEs, except LP,
at about 180? angle, and inserting on tip of Eb4 un-cinate process in larger specimen (smaller specimenlacks distinct uncinate process, and LE4 inserts on
NUMBER 11 45
small, dorsally raised cartilaginous process of medialhead of Eb4).Remarks. In almost all other pre-acanfhomorphtaxa with LE4, it originates on the dorsal margin ofEb4 (no levator process present) or on or near theLE4 levator process (including those taxa in whichEb5 is fused with the dorsodistal end of Eb4). John-son and Patterson (1996:273) reported that Clianoslacks an uncinate process, quite possibly based onexamination of a small, early ontogenetic stage spec-imen (they did not list the size of the specimens theyexamined; see also illustrations of Chanos gill archesin Johnson and Patterson, 1996). There is a bonyridge separating the cartilaginous tip of the uncinateprocess from the cartilaginous proximal head of Eblin our larger specimen (obscured by LE4 insertion inPlate 30). The reasons we indicate the process as anuncinate process is that it appears to have developedas a separation of the medial head of Eb4 (Johnsonand Patterson, 1996:273), and a posterior (actuallylateral) levator process is present. The alternative,that two Eb4 levator processes are present is alsopossible, and would be uniquely synapomorphic forChanos.LP tiny, inserting on LE4 levator process well pos-terior to uncinate process; origin not recorded.Remarks. Greenwood and Lauder (1981:228)wrote that LP appeared to be absent in Chanos. Therelatively small size of the muscle may have causedthem to overlook it, or its presence may be variable.The fragile levator process was damaged on bothsides of our smaller specimen during dissection andit was not possible to determine if LP was present.LI1 on Pb2 bony surface dorsoposteriorly.LI2 on Pb3 bony surface dorsoposteriorly.LI3 on Pb4 dorsoanterolaterally.TD comprises TPb3 and TEb4. TPb3 a broad bandof muscle attaching to Pb3 bony dorsal surface, withslender, diagonal muscle strap extending from pos-terolateral end of left side and attaching to dorsalsurface of Pb4 (diagonal strap absent in smaller spec-imen). TPb3 well separated from and not continuouswith TEb4. TEb4 more extensive than TPb3, attach-ing to Eb4 long ventrolateral arm, continuous pos-teriorly with modified SO muscles.OD3 absent.OD4 absent.OP indistinguishable, if present, obscured by EO.See also Ad5.Ad 1-3 absent.Ad4 dorsally broadly on dorsoposterior edge oflong ventrolateral arm of Eb4, narrowing consider-ably ventrally and attaching to Cb4 just medial toinner angle formed by Eb4-Cb4 joint.Ad5 questionably represented by sheet of muscleattaching dorsally to large Eb5 cartilaginous plate andventrally to distal cartilaginous process at end of Cb5.
Also possibly represented by muscle strap (Plate30B, not labeled) extending from Eb5 dorsally to ra-phe (ER?. not labeled) on posterior surface of EOand undifferentiated from SO. Muscle sheet and/orstrap possibly including OP.SO longitudinal muscle layer circumesophageal instraight portion of esophagus (not illustrated) imme-diately posterior to EO; presence of SO longitudinalmuscle fibers in EO problematic; fibers absent ante-rior to EO.RD absent.Additional remarks. SCL absent. TV4 well anteriorto, and free from, anterior to tips of Cb5s. Pb2 andPb3 edentulous, UP4 and UP5 absent. Ebl with au-togenous cartilaginous segment of anterior tip(MPbl) present; see discussion in additional remarksunder Gonorynchus (Gonorynchidae).Large, unpaired, elongate-ovate cartilage, tenta-tively identified as GC, present overlying anterior tipsof Pb2s and Pb3s, surrounded by CT, and attachingmid-ventroanteriorly to medialmost ends of MPbls.In so attaching, Chanos's GC differs, perhaps, fromthat reported by Di Dario (2002), who first describedGC. He reported that GC occurs only in most en-grauloids and pristigasteroids among "remaining clu-peiforms [= our Clupeomorpha] and basal teleoce-phalans [= Recent Teleostei]," but did not mentionits attachments. We find that it attaches to the anteriorends of the Pb2s (our observation based on pristi-gasteroid Ilisha). Di Dario's comparative material in-cluded two specimens of Chanos, and it would ap-pear that if his specimens exhibited a GC-like ele-ment, he would have discussed it. His overlookingGC in Chanos is understandable. We failed to extractit in one of our two dissections of Chanos gill arches,and it appears that it has never been mentioned inany previously published study that treated the gillarches of the genus (but described by Nelson, 1966a:157, in his dissertation).Given the cladistic interrelationhips illustrated inFig. 3, GC in Chanos and GC in Clupeoidei are mostparsimoniously interpreted as homoplasies, althoughthe element may represent retention of a basal oto-cephalan character that has been lost independentlyseveral times. GONORYNCHIDAEGonorynchus moseleyi Jordan and Snyder, USNM354590, 231 mm. Plate 31
Additional material. Gonorynchus forsteri Ogilby,USNM 353921, 90.6 mm.
Description.Remarks. There is no substantive difference in themusculature of the two species.
46 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LEI on dorsoposterior edge of medialmost bonyportion of Ebl, which lacks uncinate process.LE2 on dorsoposterior surface of bony and carti-laginous medial end of Eb2, which lacks uncinateprocess.LE3 minute, on cartilaginous tip of Eb3 uncinateprocess.LE4 reduced, on dorsomedial edge of expandeddorsolateral cartilaginous margin of Eb4*. See alsoremarks following LP.LP absent.Remarks. Greenwood and Lauder (1981:228) re-ported that LP is present in Gonorynchus. It is pos-sible that they identified the muscle we believe to beLE4, of which they made no mention, as LP. Unlessa specimen is found that has both LE4 and LP, theidentification of the muscle in question may remainunresolved.LI Ion posteromedial surface of Pb2.LI2 on posterodorsal cartilaginous surface of Pb3.LI3 on posterolateral surface of cartilaginous Pb4.TD comprises TPb3, TPb4. and TEb4*. TPb3 at-taches over most of posterior half of surface of Pb3,well separated and discontinuous from TPb4. TPb4attaches to anterodorsal surface of Pb4 and is contin-uous ventroposteriorly with TEb4*. TEb4* verybroad, attaches to medial edge of surface of Eb4*medial arm dorsal to attachment of Ad4, and is con-tinuous posteriorly with greatly expanded SO as itforms EO.OD3 small, originates tendinously on bony dorsalsurface of Pb3 uncinate process at lateral edge ofTPb3 and inserts by slender tendon on dorsoanteriormargin of cartilaginous tip of Eb3 uncinate process.OD4 small, originates on anterolateral edge of Pb4and inserts by slender tendon on dorsoanteriormostcartilaginous edge of Eb4*.OP absent, perhaps highly modified by complex ofCT and nerves in area between epibranchial organlaterally and esophagus medially.Adl-3 absent.Ad4 on mid-posterior bony surface of Eb4* andbony dorsal surface of Cb4 medial to inner angle ofEb4*-Cb4 joint.Ad5 apparently absent.GFM? muscle with small portion on dorsomedialcartilaginous surface of Eb4* and long portion ondorsoposterior surface of AC (= ace of Johnson andPatterson, 1997:fig. 2b) joining Eb4* and Cb5. Inter-pretation highly questionable: are there gill filamentsattached to AC? If not, muscle may represent sepa-rate portion of SO.RDs absent.SO longitudinal muscle layer (not illustrated) cir-cumesophageal in esophagus leading into EO, con-tinuing dorsoanteriorly as sparse fibers to at least be-low TPb3 d probably further.
Additional remarks. SCL absent. TV4 tripartite, di-vided longitudinally, with each part attaching medi-ally to ventral surface of posterior, cartilaginous bas-ibranchial copula to which Cb3 and Cb4 attach me-dially. Triangular-like cartilaginous element attachesto anterior tip of each Ebl, which Johnson and Pat-terson (1997:596) consider a mediosuprapharyngo-branchial (MPbl). We note that the anteromedialmostpoint of each MPbl is minutely ossified where theelements of the two sides meet. Similar minute os-sifications occur posteromedially where each MPblmeets the anterior cartilaginous tip of its respectivePb2. A strong, slender ligament joins posterior edgeof cartilaginous tip of Eb3 uncinate process with an-terior bony edge of Eb4*. Pb2 and Pb3 edentate; UP4and UP5 absent. CypriniformesCYPRINIDAEZacco platypus, USNM 336890, 2 specimens, 84-101 mm; 3 cleared and counterstained, 28?82 mm.Plates 32.1, 32.2
Additional material. ? = Opsariichthys bidens Giin-ther, USNM 112443, 135 mm.
Description.Remarks. Many of the muscles described belowappear to be unique to cypriniforms. Our attempts tohomologize the names applied to these muscles, re-ferring to Takahashi (1925), Holtsvoogdt (1965), andWinterbottom (1974b), were only partly successfulbecause of unclear descriptions and illustrations. Weuse some of these authors' names for these muscles,but have introduced our own terminology for others.Homologies among many of the dorsal gill-arch mus-cles of otophysans remain to be elucidated.LEI on mid-posterodorsal surface of Ebl.LE2 on mid-posterodorsal surface of Eb2.LE3 on base of all bony Eb3 uncinate process.LE4 on cartilaginous tip of Eb4 levator processand tiny, horizontally oriented autogenous bone (AB,Plate 32.2) attached to posterior surface of cartilagi-nous tip of Eb4 levator process (AB also joined byCT to pad-like insertion of LCb5A; dorsal end of Eb5attached to AB).LP absent.Remarks. Winterbottom (1974b:252-253 and fig.22) treats our LCb5 as his LP externus and ourLCb5A as his LP internus. LCb5 inserts on Cb5 andLCb5A inserts on a CT pad attached to Cb5. Themuscle we define as LP inserts invariably on Eb4, orjoins LE4 in a combined insertion on Eb4. Further-more, with the exception of Chanos, in all pre-acan-thomorphs with LP, LP inserts on or together withLE4. In Chanos, LP inserts on Eb4 well posterior toLE4. We, therefore, infer that LP is absent in cypri-
NUMBER 1
1
47
nids. If one or the other of these two muscles, LCb5Aor LCb5, however, is derivative of the levator pos-terior, it would indicate that LP is a synapomorphyof the Otocephala.LI1 on posterior surface of Pb2 dorsolaterally, nearjoint with Eb2.LI2 on Pb3 mid-laterally and medial end of Eb3,which joins Pb3. ? On Pb3 mid- to posterolaterallyand Eb3 and Eb4 medial ends, which join Pb3 (in-sertion occupying same area as combined LI2 andLI2' of Z. platypus).Remarks. Takahashi (1925:41) reported that LI2has only one insertion in Z. platypus (and O. unci-
rostris); the muscle is apparently variable within thegenus; see also remarks under LI2'.LI2' on posterolateral cartilaginous edge of Pb3and anteromedial cartilaginous and bony edges ofEb4, spanning joint between Pb3 and Eb4. (see LI2).
? Not present; see LI2.Remarks. Pb4 is absent in both Z platypus and O.bidens, but is present in other cyprinids; e.g.. NewWorld Notropis hudsonius (USNM 315400, clearedand stained). Old World Abbotina (USNM uncat.,cleared and stained). With the exception of the cyp-rinid genus Pseudogobio, which has three Lis (= ourLI1, 2, 2'), Takahashi (1925:41-42) found only twoLis in cyprinoids, including Opsariichthys uncirostrisand Zacco platypus. He mentions, however, that thesecond LI of Cyprinus carpio has two "caudae" [or-igins?]. His descriptions (p. 42) of the three Lis incobitoids indicate very similar states to those wefound for the three Lis in Zacco platypus.LI3 absent.LI4 (see LCb4).LCb4 slender, originating on exoccipital and in-serting by long tendon on Cb4 among medial fibersof Ad4.Remarks. Holstvoogd (1965:fig. 12b) termed thismuscle levator PV interims in the cyprinid Leuciscus(and also for a muscle that we term LP attaching toEb4 in characiforms). We reserve LI for muscles in-serting on pharyngobranchial elements. LCb4 occursonly in cypriniforms, possibly only cyprinids. LCb4,however, may represent a different character state forLI4, which is found only in Diplomystes (Diplomys-tidae), in which it inserts on UP4.LCb5 massive, on dorsoposterior surface of hy-pertrophied Cb5, originating in subtemporal fossawith two levels of attachment: dorsoanteriorly mainlyor entirely on pterotic; ventroposteriorly mainly orentirely on exoccipital; found only in cypriniforms.See remarks following LCb5A.Remarks. Winterbottom (1974b:252-253) denotedthis muscle as his LP externus (see remarks followingLP above).LCb5A small, originating in supratemporal fossatogether with ventroposterior level of origin of LCb5
(q.v.). wrapping medially, then anteriorly aroundLCb5 and inserting in thick CT pad strongly attacheddorsally to anterolateral surface of Cb5 (pad alsojoined anteriorly by CT to tiny AB attached to tip ofEb4 levator process).Remarks. LCb5A appears to be the same as theinternus branch of the levator posterior of Winter-bottom (1974b:fig. 22b; indicated by the left lineleading from his label L.POST to the muscle; theright line leads to the externus branch, which equalsour LCb5).TD absent (unique among Halecostomi); possiblyreplaced by thick SO section extending anteriorlyventral to pharyngobranchials.OD3 on Pb3 bony dorsal surface anterolaterallyand tip of bony Eb3 uncinate process.OD4 on Pb3 bony surface posterolaterally andbony tip of Eb4 uncinate process.OP absent.RecD2 posteriorly on anteroventral surface of me-dial half of Eb2 and ventrolateral bony surface ofPb2, anteriorly on mid-ventromedial bony surface ofEbl.RecD3 posteriorly on anteroventral surface of me-dial half of Eb3 and ventrolateral surface of Pb3, an-teriorly on mid-ventromedial bony surface of Eb2.RecD4 posteriorly on anteromedial surface of Eb4,anteriorly on bony posteromedial surface of Eb3.Ad 1-4 each attaching to ventral surface of its re-spective Eb and dorsal surface of its respective Cb
at the internal angle formed by the two bones.Remarks. Among pre-acanthomorphs, only poly-odontids and cyprinids have Ad 1-3, and their attach-ments in the two groups are different and differentfrom those of acanthomorphs having Ad 1?3.Ad5 fan-like, with broad anterior end on dorso-posterior margin of Eb4 levator process and narrowposterior end on dorsolateral surface of Cb5.RCb5T originates medially from CT and SO. wellseparated from contralateral RCb5T, and inserts lat-erally on dorsolateral margin of Cb5.Remarks. Holstvoogd (1965:fig. 12a) called thismuscle transversus ventralis posterior. We think thename misleading as transverses ventrales are other-wise applied to ventral gill-arch muscles. Winterbot-tom (1974:258; fig. 22b) identified this muscle a theretractor pharyngeus superioris.RCb5E massive, originates on ventral basioccipitalprocess posterior to and partially dorsal to RCb5I,and inserts on Cb5 dorsolaterally.Remarks. Winterbottom (1974b:fig. 22) named thismuscle retractor pharyngeus inferioris.RCb5I relatively slender, originates as fine line ofCT along dorsolateral surface of a vertical SO foldmedially abutting a ventral basioccipital process andinserts by long tendon anteriorly on thick CT pad towhich LCb5A (q.v.) attaches (pad attaches to Cb5).
48 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Remarks. The vertical SO fold may be a separatemuscle. It is slightly disjunct anteriorly from a pos-teriorly ovoid area of SO muscles, which are anter-oventral to the ventral basioccipital process. A pair(one on each side) of muscle extensions continues ashort distance anteriorly from the ovoid area, andthese and the ovoid area muscles are covered by atough fascia. The fascia divides and continues ante-
riorly on each side forming a pad covering the sur-face of the anterodorsal ends of Pb2 and Pb3. An-teriorly from pair of muscle extensions, the fasciaforms long, slender tendons that attach to the medialsurface of Pb3. Winterbottom (1974b) did not men-tion RCb5I.SO longitudinal muscle layer absent at and poste-rior to horizontal joining posteriormost edges ofCb5s; very thick longitudinal layer begins belowtransverse layer anterior to horizontal and extendsventrally below gill arches well anterior to horizontal
at anterior margins of Ebls (roofs much of oral cav-ity); transverse muscle layer absent anterior to anter-iormost external extent visible in Plate 32.1A (ante-rior portion of longitudinal muscle layer not overlainby transverse muscle.RD absent.Additional remarks. SCL absent. TV4 divided (in-terrupted), attached medially to each Cb5. Slender,almost threadlike cartilaginous Eb5 (may be in 1 to3 pieces) attaches ventrally to dorsoposterior tip ofCb4 and dorsally to tiny autogenous bony element(AB) at posterodorsal tip of Eb4 levator process. Pbl,Pb4, UP4, and UP5 absent.
CharaciformesHowes (1976) treated cranial muscles of certaincharaciform fishes, essentially avoiding the dorsalgill-arch muscles except for mention of Kampf's( 1 96 1 ) brief treatment of of these muscles in Hydro-cyon forskali (= Hydrocynus forskalii, Alestiidae)and Winterbottom's (1974b) general study of fishmusculature, which included reference to Bryconguatemalensis (Characidae). In apparent justification,Howes (p. 219-220) stated, "The arrangement [ofthe branchial muscles] in the Cynodontini is basicallyas in Brycon, and a provisional survey of branchialarch myology in . . . various characoid families (pers.obs.) suggests relative uniformity throughout thegroup. However, some [unspecified] specializationshave been found in those taxa with epibranchial or-gans (Chilodus, Anodus)." There may be relativelymore variation among characiform gill-arch muscu-lature than Howes opined.Kampf (1961:436, figs. 29-20), using terminologydifferent from ours, reported that TD included at-tachment to Pb5, LE3 absent, and Adl-3 present,character states not present in the taxa we examined.
In the case of Pb5, we find that ventral strands of SOmuscle attach to the medial surfaces of UP4 and UP5.We arbitrarily did not treat the latter as part of TD,because these tooth plates extend posteriorly into theesophagous. Kampf's Adl-3 may equal our GFM1-3. But none of the three taxa we examined lackedLE3. CHARACIDAEBrycon guatemalensis Regan, USNM 114526, 99.0mm SL. Plate 33Additional material.
?
= Brycon melanopterus(Cope), USNM 307072, 78.4 mm SL.
Description.Winterbottom (1974b:fig. 20) presented a lateralview of the dorsal gill-arch muscles of B. guatema-lensis, but some muscles are obscured.LEI on dorsal edge of Ebl just lateral to cartilagetip of uncinate process; origin slightly dorsal to Pblarticulation with skull.
? On Ebl uncinate processjust ventrolateral to cartilaginous tip.LE2 on and just ventral to cartilaginous tip of Eb2uncinate process.
?
On Eb2 just lateral to uncinateprocess.LE3 on cartilaginous tip of Eb3 uncinate process.LE4 on dorsalmost tip of Eb4 levator process andtendinous base of LP.LP attached tendinously to dorsalmost tip of Eb4levator process.LI1 on dorsoposterior surface of Pb2.LI2 on dorsal surface of Pb3 posterolaterally.LI3 on dorsoposterior surface of cartilaginous Pb4.TD comprises TPb3a-Eb2, TPb3p, and TPb4-Eb4(see also discussion following Characiformes for pos-sibly excluded TD muscle). TPb3a-Eb2 with sparsemuscle strands attaching to Pb3 dorsoanteriorly, butmainly attaching to cartilaginous tip of Eb2 uncinateprocess laterally, posteriorly continuous by diagonalmuscle strap with TPb3p. TPb3p on Pb3 posterolat-erally, posteriorly continuous by diagonal muscleslips with TPb4-Eb4. TPb4-Eb4 broadly on Pb4 andnarrowly on Eb4 medially, broadly continuous pos-teriorly with SO.OD3 origin on Pb3 anteriorly ventral to TPb3a-Eb2, extends posteriorly dorsal to TPb3p, and insertson cartilaginous tip of Eb3 uncinate process medialto insertion of LE3.OD4 origin mostly on Pb4 anteriorly, slightly onPb3 at joint with Pb4, muscle fans out posteriorlyand inserts along most of lateral surface of broadlevator process of Eb4.OD4' long, slender; tendinous origin on Pb3 ven-tral to OD3 origin, joins OD4 insertion ventrally onanterolateral face of Eb4. ? OD4' absent.
NUMBER 1 1 49OP a strap of muscle originating on posteromedialsurface of Eb4, poorly separated from SO, endingventrally at ER; muscle fibers continuing ventrallyfrom ER include Ad5 and SO.M. UP5-Cb4-Eb5 (not illustrated), origin on lateralmargin of UP5, insertion at inner angle formed byCb4 and Eb5.Remarks. M. UP5-Cb4-Eb5 appears to be a vari-ation of Kampf's (1961:436, fig. 30, lower right) ob-liquus dorsalis inferior in Hydrocyon forskali (= Hy-drocynus forskallii (Cuvier)). Kampf described it,however, as connecting Pb5 (= our UP5?) with Cb5.Winterbottom (1974b) did not mention obliquus dor-salis inferior among his muscle synonymies. The al-bulids are the only other fishes we examined thathave a possibly equivalent muscle, which we indicateas M. UP5-Cb4.Ad 1-3 absent.Ad4 on Eb4 dorsoposteriorly and Cb4 dorsal sur-face medial to inner angle of Eb4-Cb4 joint.Ad5 on Eb5 mid-posteriorly and Cb5 posteriorly,with thin dorsolateral tendinous extension (not illus-trated) crossing Eb5 dorsally to distalmost bony endof Eb4; joins ER below OP laterally and SO medi-
ally, merges with SO medially and TV5 ventroanter-iorly.SO longitudinal muscle layer surrounds esophagus,thick dorsally, very thin elsewhere; fibers attachinganteriorly along medial edge of large UP4; fibers ex-tend anteriorly to below anterior end of TPb4-Eb4.
? Layer moderately evenly distributed around esoph-agus.RDs absent.Additional remarks. SCL absent. TV4 free fromCb5s. Slender ligament from cartilaginous tip of Eb4levator process to Eb5 dorsally. Eb5 ventrally on dis-talmost end of Cb4.DISTICHODONTIDAEXenocharax spilurus Gunther, USNM 227093, 125mm. Not illustrated
Description.LEI broadly on Ebl uncinate process beginning alittle lateral to cartilaginous tip, origin joins Pbl ar-ticulation with skull; short, band-like tendon joins or-igin with Pbl just ventrolateral to dorsal cartilage tip.LE2 on Eb2 just lateral to tip of uncinate process.LE3 on anteromedial edge of Eb3 uncinate pro-cess, there joining OD3 insertion.LE4 on dorsalmost edge of Eb4 levator process,joining raphes ventrolaterally with LP insertion andAd4 dorsally.LP on Eb4, joining raphes ventromedially withLE4 insertion and Ad4 dorsally.LI1 on dorsomedial surface of Pb2.
LI2 on dorsal surface of Pb3 posterolaterally.LI3 dorsoposterolaterally on Pb4.TD comprises TPb3a-Eb2, TPb3p, TPb4, andTEb4. TPb3a-Eb2 on Pb3 dorsally beginning at an-terior edge of bony surface and extending posteriorly,and on cartilaginous tip of Eb2 uncinate process dor-
sally, posteroventrally continuous by diagonal muscle
slip with TPb3p anteriorly. TPb3p on Pb3 postero-laterally, beginning ventral to TEb2, and just failingto meet LI2 insertion anteriorly; posteriorly contin-uous by diagonal muscle strand with TPb4. TPb4broadly on Pb4, sharply separated posteriorly fromTEb4. TEb4 on Eb4 anteriorly ventral to insertionsof OD3-4 and OD4'.OD3-4 origin on Pb3 anteriorly, divides posteri-orly with insertions on tip of Eb3 uncinate processmedial to LE3 insertion, and on Eb4 dorsolaterally.OD4' origin on Pb4 anteriorly joining raphe dor-soanteromedially with OD3?4 origin posteriorly; in-sertion on Eb4 ventral to OD3-4 insertion.OP a muscle strap originating on Eb4 posterome-dially, joining TEb4 ventrolaterally and irregular ERdorsally, which is joined ventrally by Ad5 (ER con-tinues short distance medially into SO). OP unclearlydifferentiated from SO medially.M. UP5-Cb4-Eb5 origin on lateral margin of UP5,insertion at inner angle formed by Cb4 and Eb5.(Also see remarks following description M. UP5-Cb4-Eb5 in Brycon).Ad 1-3 absent.Ad4 well developed on Eb4 dorsoposteriorly be-ginning medially ventral to uncinate process and ex-tending laterally to end of bone, joining raphes dor-sally with LE4 and LP insertions; ventrally very nar-rowly on Cb4 just medial to Eb4-Cb4 joint.Ad5 bulbous, broadly on Eb5 posteriorly and Cb5posteriorly, with thin dorsolateral tendinous exten-sion sheathing Eb5 dorsally and attaching to distal-most bony end of Eb4; muscle joins ER ventral toOP and merges medially with SO; ventrally joinsTV5 posterolaterally.SO longitudinal muscle layer thick dorsally, verythin or absent elsewhere, begins at about horizontalconnecting distal ends of Eb4s and extends anteriorlyto anterior end of TEb4.RDs absent.Additional remarks. SCL absent. TV4 free fromCb5s. Slender ligament joins cartilaginous tip of Eb4levator process to Eb5 dorsally, cartilage ventrally onposterodistalmost end of Cb4.
SiluriformesDIPLOMYSTIDAEDiplomystes chilensis (Molina)?, USNM 259097,119 mm. Plate 34
50 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Description.LEI on Ebl dorsoposteriorly.LE2 on Eb2 dorsoposteriorly.LE3 absent.LE4 absent, but see remarks under LI4.LP absent (see remarks under LI4).LI1 on posterior margin of small CT pad, whichimpinges on ventral cranial surface and incorporates:medial ends of Pbl-3 and Ebl; autogenous ball ofcartilage that articulates with medial ends of Ebl andPb2; and a few smaller autogenous cartilages; inser-tion splits OD3 origin.LI2 inserts mainly on dorsoanterior edge of largeUP4, with minor insertion on anterolateral cartilagi-nous edge of Pb4.Remarks. Insertion of LI2 on Pb4 (or UP4) is un-common among fishes we examined (Table 2).LI3 medially on lateral cartilaginous edge of Pb4,ventrolaterally on UP4 dorsoposteriorly, and Eb4 me-dial cartilaginous cap at Eb4-Pb4 joint.Remarks. Insertion including Eb4 is unique amongfishes we examined.LI4 originates on pterotic and inserts mainly onthe dorsoposterior surface of UP4, with some fibersinserting on cartilaginous joint formed by Pb4 withEb4.Remarks. Takahashi (1925:67), using different ter-minology, considered this muscle to represent RD insiluriforms, in which it originates variously on thepterotic or supraclavicle (? = posttemporal-supra-cleithrum) and inserts variously on the "posteriorpharyngobranchial" or Eb4 (Diplomystes was notamong his material). Takahashi did not distinguishPb4 from its toothplate, and we are uncertain of theexact location of the insertion in the taxa he listed.The origin of LI4 would seem to exclude interpre-tation of LI4 as RD, which usually originates on theanterior vertebrae, or possibly the basioccipital. Hol-stvoogd (1965:215 and fig. 10) termed this musclesimply "Levator IV" in the clariid siluriform Clar-ias. Winterbottom (1974b:253 and fig. 21) question-ably designated the muscle LP in Diplomystes.Among pre-acanthomorphs, LP occurs only amongOtocephala (clupeomorphs and ostariophysans).which might be evidence that the muscle we desig-nate LI4 is a modified LE4 that has shifted its inser-tion from Eb4 to UP4 (the slight partial insertion in-cluding the medial end of Eb4 providing evidence).LP, otherwise, invariably inserts on Eb4 together withLE4, which Diplomystes otherwise lacks.TD comprises four sections (a?d), of which thefirst is least differentiated. Anteriormost section, a,TPbl-Pb2-Pb3-Ebl-Eb2 (includes only anteriormosttip of Pb3) underlies pair of CT pads (see LI1) andis posteriorly continuous with next section, b, TPb3-Pb4, which joins medial bony edges of Pb3 and Pb4and overlies tanterior end of next section, c, TUP4a.
Laterally, TUP4a attaches to dorsomedial surface(ledge) of UP4, and is continuous posteriorly with d,TUP4p, which curves anteriorly and inserts alongdorsolateral surface of UP4; TUP4p is continuousposteriorly with SO.OD3 has a tripartite origin: dorsolateral origin (a)on Pbl and posterolateralmost margin of CT pad (seeLI1), including medial tip of Ebl, separated by in-sertion of LI1 from medial origin (b) on posterome-dial margin of CT pad; these two origins, separatedby LI1 insertion, meet and fuse posteriorly (obscuredby recumbent LI1); ventrolateral origin (c) a shortbranch on Eb2, fuses with ventral surface of dorso-lateral origin; complex inserts on anterior surface ofEb3 uncinate process.OD4 origin on dorsoposterior surface of Pb3, ven-tral to OD3, continuing onto dorsoanteromedial sur-face of Pb4; insertion on anterior surface of Eb4 un-cinate (levator?) process.Remarks. Arratia (1987:11, 41) reported that Di-plomystes lacks an uncinate process on Eb4, althoughmentioning that Eb4 "has a short lateral projectionwhich I do not consider an uncinate process." Shemade no mention of a levator process. We find theuncinate (or levator) process present and well devel-oped. The uncinate process on Eb3 is armlike, where-as that of Eb4 arises vertically from the horizontal asa broad, dorsally obtuse flange with a distinct carti-laginous cap.OP dorsally on posteromedial surface of Eb4, ven-trolaterally on medial end of Eb5, ventromediallyjoining raphe (ER?) with SO, continuing ventrallybelow raphe and attaching to bony medial surface ofCb5; ventrolaterally overlapping medial portion ofAd5.RDs absent (see LI4 above).SO longitudinal muscle band circumesophageal,thick dorsally and ventrally, thin laterally, extendinganteriorly to below anterior end of TPb3-Pb4.Ad 1-3 absent.Ad4 dorsally on posterodorsal surface of Eb4 un-cinate (levator?) process, ventrally on Cb4 dorsal sur-face.Ad5 dorsally on posteromedial surface of Cb4 andventral surface of Eb5; ventrally on dorsoposterioredge of Cb5, extending medially under OP.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 partially ossified dorsomedially. UP5 ab-sent. Eb5 on posterodistalmost end of Cb4, horizon-tally oriented, a position duplicated only in Pantodon(Osteoglossomorpha). Small, sesamoid bone (basi-hyal?) heavily enveloped in CT present between an-terior hypohyals (presumably, ventral hypohyals be-cause they articulate with the urohyal; Arratia, 1987:fig 27, indicates these are dorsal hypohyals, whereas,Azpelicueta, 1994:figs. 6b and 11. correctly indicatesthey are ventral hypohyals). de Pinna (E-mail, Mar-
NUMBER 11 51
Apr 1999) informs us that cartilaginously tipped Eb3and Eb4 uncinate processes occur only in Diplomys-tes among siluriforms, and that the basihyal and Bblare absent in all catfishes.GymnotiformesGYMNOTIDAEGymnotus carapo Linnaeus, USNM 260272, 320 mmTL. Plate 35
Description.Remarks. Hoz and Chardon (1984) reported on thegill-arch musculature of the gymnotid Sternopygusmacrurus, which is very similar to that of Gymnotuscarapo. We remark on the differences.LEI on Ebl dorsoposteriorly near distal end.LE2 posteriorly on Eb2 dorsoposterior edge justlateral to LI2 insertion, and anteriorly on dorsopos-terior edge of Ebl medial to LEI insertion.Remarks. Hoz and Chardon ( 1984) do not mentionan attachment to Ebl. Verification in other specimensis desirable.LE3 on bony Eb3 uncinate process just lateral toOD3 insertion.LE4 on levator process at dorsodistal end of Eb4.LP absent.LI1 broadly, tendinously on Pb2 dorsoanteriorlyand narrowly on posteromedial end of Ebl dorsally.Remarks. Hoz and Chardon (1984) indicate inser-tion only on Pb2.LI2 primarily on dorsoanteriormost surface of me-dial end of Eb3, barely continuing onto adjacent la-teralmost edge of Pb3, secondarily on dorsoposter-iormost edge of Eb2 immediately medial to LE2 in-sertion.Remarks. Identification of this muscle as LI2 isproblematic because of its primary insertions on Ebsrather than Pb3. Hoz and Chardon (1984) indicateinsertion is only on Eb3.LI3 on Pb4 and UP4 dorsolaterally.Remarks. Hoz and Chardon (1984) report insertionis only on Eb4. Their illustration (fig. 19b) is unclear,but the external position of the muscle appears sim-ilar to that of LI3 in G. carapo. We believe theyintended to report the insertion as Pb4, a lapsus sim-ilar to their listing (p. 44) of LE4 as a [second] LE3,but labeling the muscle LE4 on their fig. 19b.TD comprises TPb2-Pb3-Pb4 and TEb4. TPb2-Pb3-Pb4 attaches to medial edge of Pb2 on one sideand to CT on other, medial edge of Pb3 (or slightlymedial to edge) along OD4 origin, and Pb4 medially,and is narrowly continuous posteriorly with TEb4.TEb4 on posteromedial edge of Eb4, posteriorly con-tinuous with SO.Remarks. Hoz and Chardon (1984) divide TD into
two parts, one attaching to Pb2 and Pb3 and one at-taching to Pb4, with no mention of an attachment toEb4.OD3 relatively small, originating on Pb3 dorsoan-teriorly and Eb2 dorsomedialmost end, and insertingon medialmost end of bony Eb3 uncinate process.OD4 relatively large, originating on Pb3 and Pb4dorsally and inserting on Eb4 dorsolaterally.OP absent.Ad 1-3 absent.Ad4 dorsally broadly on ventrolateral surface ofEb4 and ventrally broadly on anterolateral surface ofCb4 medial to Eb4-Cb4 joint.Ad5 ventrally broadly on posterolateralmost sur-face of Cb5, just impinging on lateralmost end ofTV5, and dorsally narrowly on posterodistal surfaceof Eb5.RDs absent.SO longitudinal fibers circumesophageal, begin-ning posterior to horizontal connecting distal ends ofCb5s, forming very thick section dorsally in area be-tween distal ends of Eb4s. and essentially absent an-terior to horizontal through posterior end of TEb4.Additional remarks. SCL absent. TV4 continuousfrom one side to the other, forming median raphe andattaching tightly to dorsoanteriormost tips of Cb5s.UP5 absent. SalmoniformesSALMONIDAEOncorhynchus mykiss (Walbaum), USNM 351540,121 mm, USNM 333092, 91.8 mm.Plate 36
Description.LEI on tip of Ebl uncinate process.LE2 on tip of Eb2 uncinate process.LE3 on tip of Eb3 uncinate process just dorsal toOD3 insertion.LE4 on dorsolateralmost surface of Eb4.LP absent.LI1 broadly on Pb2 dorsally.LI2 on Pb3 dorsoposteriorly.LI3 insertion enveloping Pb4 dorsal and anteriorsurfaces and just attaching to dorsoanterior edge ofUP5.TD a broad sheet of muscle comprising TPb3a andTPb3-Pb4-Eb4 (Pb3 portion slightly differentiatedlaterally from remainder). TPb3a short, laterally onPb3 uncinate process, fusing medially with OD3 an-teroventral surface dorsal to OD3'. TPb3-Pb4-Eb4anteriorly on Pb3 ventral to OD3', forming raphewith OD4 origin, on Pb4 medial to LI3 insertion, andon medial end of Eb4, posteriorly continuous with,but noticeably demarcated from SO.OD3 anteriorly dorsal to anterolateral portion of
52 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
TPb3-Pb4-Eb4; dorsoanteriorly on Pb3 near anteriorend; joined to opposite OD3 by median raphe; an-other raphe lateral to median raphe setting off smallwedge-like anterior muscle section; anteroventral sur-face joined by medial end of TPb3a, possibly evi-denced by dorsolateral OD3 raphe; insertion on Eb3uncinate process ventral to LE3 insertion.OD3' completely ventral to OD3, origin on Pb3posteromedial to OD3 origin and anterior to TPb3-Pb4-Eb4; fuses with ventral surface of OD3 at aboutmid-length of OD3.OD4 origin on Pb3 dorsoposterior surface and Pb4dorsoanterior surface, forming raphe with TPb3-Pb4-Eb4 medially, inserting on dorsal surface of Eb4 me-dial to LE4 insertion, forming partial raphe with OPand/or Ad4 dorsally; some muscle strands continuouswith OP and Ad4.OP dorsally on Eb4, ventrally joining ER at aboutmid-level of Eb5, below which OP is undifferentiatedfrom SO (see also OD4). ER in smaller specimenbetter developed, extends medially from mid-level ofEb5 for width of OP and then turns relatively sharplyposteriorly for about same distance on SO.Adl-3 absent.Remarks. Dietz (1912:fig. 2; p. 19), illustrated andreported Ads 1?4 in Salmo salar Linnaeus as follows(our translation): "This system of weak rudiments ispresent on the anterior four gill arches as a border ofshort, obliquely placed fibers along the exterior of theepibranchials and the proximal greater part of the cer-tatobranchials. Only against the inside of the fourthgill arch is it well developed in form . . ." Although,we did not examine 5. salar, it is clear from his de-scription that Dietz identified the gill filament mus-cles on the first three arches as adductors.Ad4 dorsally on Eb4, ventrally on Cb4 medial toinner angle formed by Eb4-Cb4 joint (see also OD4).Ad5 dorsally on mid-posterior surface of Eb5. andventrally on Cb5 distal end.RD absent.SO longitudinal muscle layer absent posterior tohorizontal between distal ends of Cb5s, present asisolated, very thick band dorsally anterior to (andabove) horizontal connecting distal ends of Cb5s andcontinuing anteriorly only to about horizontal con-necting anterior ends of Pb4s.Additional remarks. SCL attached mid-dorsally tocartilaginous posteroventral tip of Bb3. TV4 freefrom Cb5s. UP4 absent.
Thymallus arcticus (Pallas), USNM 179764, 153mm. Plate 37Description.LEI on cartilage tip of Ebl uncinate process.LE2 on cartilage tip of Eb2 uncinate process dor-solateral^.
LE3 on cartilaginous tip of Eb3 uncinate process.LE4 on dorsalmost edge of distal end of Eb4, be-tween dorsal edge of OD4 insertion and dorsal at-tachment of Ad4.LP absent.LI1 on mid-dorsal bony surface of Pb2.LI2 relatively small; on left-side muscle almost in-separable from LI3 dorsal to LI2 insertion, which ison posterodorsal bony surface of Pb3; right-side LI2and LI3 completely separate, insertions as on leftside.LI3 on left side with two slightly separate inser-tions, anterior insertion on Pb4 dorsoanteriorly, pos-terior insertion on posterolateral edge of Pb4, con-tinuing posteroventrally and attaching tendinously todorsoposterolateral edge of UP5; on right side withtwo completely separate insertions (essentially twoseparate muscles), anterior muscle on dorsoanteriorsurface of Pb4, posterior muscle on dorsolateral edgeof UP5.Remarks. About one-fifth of left-side UP5 is ven-tral to Pb4 (remainder is ventral to Eb4), whereas onright side almost one-half is ventral to Pb4.TD comprises TPb3 and TPb4-Eb4. TPb3 on Pb3uncinate process joining raphe with anterior portionof OD3 and OD4 origins on right side and OD4 or-igin on left side; fibers changing from transverse tocrisscross about halfway along raphe, then continuingas transverse fibers of TPb4-Eb4. TPb4-Eb4 on dor-soposterior surface of Pb4 and dorsomedial surfaceof Eb4, continuous with SO posteriorly.OD3 strap-like, much smaller than OD4; left-sideOD3 origin on medial edge of cartilage tip of Pb3uncinate process ventral to OD4; right-side OD3 or-igin on dorsolateral surface of Pb3 uncinate processanterior to OD4 origin, but continuous posteriorlywith it; insertion on both sides identical, on medialedge of cartilaginous tip of Eb3 uncinate process.OD4 massive; left-side OD4 completely dorsal toOD3, origin begins on lateral edge of Pb3 uncinateprocess and continues posteriorly onto Pb4, joins ra-phe with TPb3; right-side OD4 origin in line andposterior to OD3 origin, joins raphe with TPb3; mus-cle on both sides identical, insert on dorsal surfaceof Eb4 medial to LE4 insertion.OP muscle strap beginning medially on posteriorsurface of Eb4 and extending laterally short distanceto dorsomedial edge of Ad4, barely separable medi-ally from SO, ventrally joining ER; ventral to ERindistinguishable from SO laterally and Ad5 medi-
ally.Adl-3 absent.Ad4 dorsally on posterolateral surface of Eb4 un-cinate process, ventrally on Cb4 dorsal surface justmedial to inner angle formed by Eb4-Cb4 joint.Ad5 dorsally on Eb5 mid-posterolateral surface,ventrally on Cb5 posterolaterally beginning at distal
NUMBER 11 53
tip and extending slightly medially, medially mostlyinseparable from OP.RD absent.SO longitudinal muscle layer absent posterior tohorizontal between distal ends of Cb5s, present asisolated, very thick band dorsally anterior to (andabove) horizontal connecting distal ends of Cb5s andcontinuing anteriorly only to about horizontal con-necting anterior ends of Pb4s.Additional remarks. SCL dorsomedially attachedto posteroventral cartilaginous end of Bb3. TV4 freefrom Cb5s. UP4 absent,adductors.
ginning lateral to OP attachment, ventrally on Cb4medial to inner angle formed by Eb4-Cb4 joint.Ad5 on mid-posterior surface of Eb5 dorsally, andCb5 distal end ventrally.RD absent.SO longitudinal muscle layer absent posterior tohorizontal between distal ends of Cb5s, present ven-trally and as isolated, very thick band dorsally ante-rior to (and above) horizontal connecting distal endsof Cb5s and continuing anteriorly only to about hor-izontal connecting anterior ends of Pb4s.Additional remarks. SCL present. TV4 complex,attached by tendons to Cb5.
Coregonus artedi Lesueur, USNM 1 1 7488. 2 speci-mens, 132-140 mm SL.Plate 38
Description.LEI, small, on tip of Ebl uncinate process.LE2 small, on tip of Eb2 uncinate process.LE3 on tip of Eb3 uncinate process dorsolaterally.LE4 tendinously on dorsalmost edge of distal endof Eb4.LP absent.LI1 on Pb2 dorsoposteriorly.LI2 on Pb3 dorsoposteriorly.LI3 on Pb4 dorsally.TD comprises three parts, a?c: a, TPb3a, raiseddorsally (dorsal to OD3 origin), attaching to anteriorbony surface of Pb3, including anterior bony edge ofPb3 uncinate process, posteriorly continuous (discon-tinuous in smaller specimen) by diagonal musclestrand with more ventral TPb3p (/?), which attachesto Pb3 bony surface dorsoposteriorly; TPb3p contin-uous posteriorly by broad diagonal muscle strap with
c, TPb4-Eb4, which attaches to dorsomedial edges ofPb4 and Eb4 and is broadly continuous by diagonalband of muscle with SO.OD3 origin divided; main portion on dorsoanteriorbony surface of Pb3 ventral to TPb3a; separate, ven-trolateral smaller portion (not illustrated) on posterioredge of Pb3 uncinate process; insertion on Eb3 un-cinate process dorsomedially.Remarks. Smaller specimen has undivided originon both sides.OD4 origin on Pb3 dorsoposteriorly and Pb4 dor-soanteriorly, insertion on dorsal edge of Eb4 well me-dial to LE4 insertion.OP dorsally on Eb4 medial to Ad4 dorsal attach-ment; ventrally joins ER at about level of dorsal at-tachment of Ad5 to Eb5, and is undifferentiated fromSO ventral to ER.Ad 1-3 absent.Ad4 dorsally, broadly on Eb4 posterolaterally be-
RETROPINNIDAERetropinno osmeroides Hector, USNM 304419, 106mm. Plate 39
Description.LEI finely, tendinously on cartilaginous tip of Ebluncinate process.LE2 finely, tendinously on cartilaginous tip of Eb2uncinate process.LE3 on tip of Eb3 uncinate process.LE4 on dorsodistal edge of Eb4*.LP absent.LI1 on Pb2 dorsoposteromedially ventral to ante-
rior end of Pb3.LI2 on Pb3 dorsoposteriorly (well separated fromLI3 insertion on Pb4).Remarks. Right-side LI2 probably anomalous, hasslender separate portion inserting on Pb4 anteriorly,and well separated from main LI2 and LI3 insertions.LI3 on Pb4 dorsoposteriorly.TD comprises TPb3a and TPb3p-Pb4-Eb4*.TPb3a lies dorsal to remainder of TD, comprisesloose strands of muscle connecting bony lateral edgesof Pb3 uncinate process dorsal to origin of OD3, con-nected posteriorly by fine muscle strand to TPb3p-Pb4-Eb4*. TPb3p-Pb4-Eb4* broad, compact, atta-ches to Pb3 dorsal surface dorsoposterolateral toOD4 origin, and to medial surfaces of Pb4 and Eb4*.OD3 origin on Pb3 dorsoanteriorly ventral toTPb3a. insertion on tip of Eb3 uncinate process ven-tromedial to LE3 insertion.OD4 origin on Pb3 posterodorsal bony surfaceventral to posterior margin of TPb3p portion ofTPb3p-Pb4-Eb4*, insertion mostly on Eb4 :i: dorso-lateral edge proximal to LE4 insertion.OD4' fine, originating on Pb4 dorsal surface com-pletely ventral to OD4; originating fibers passthrough TPb3p-Pb4-Eb4*; muscle fuses with OD4ventral surface well anterior to OD4 insertion.Remarks. Muscle is possibly anomalous; confir-mation of its presence in other specimens is desirable.OP comprises two or three muscle strands origi-
54 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
nating on posteromedial surface of Eb4* dorsal to SOattachment to Eb4*, joining ER dorsally, undiffer-entiated from SO ventral to ER.Ad 1-3 absent.Ad4 dorsally on posterior surface of Eb4*, ven-trally on Cb4 dorsal surface just medial to inner angleformed by Eb4*-Cb4 joint.Ad5 dorsally on posterodistal surface of cartilagi-nous Eb4* posterodistal hook-like process, ventrallyon posterodistal surface of Cb5, medially continuouswith SO or OP?.RD absent.SO longitudinal muscle layer absent or represented
at most by few sparsely distributed fibers dorsallyand ventrally posterior to horizontal through distalends of Cb5s, dorsally becoming thick, then attenu-ating, and extending anteriorly to anterior end of TD.Additional remarks. SCL absent. TV4 absent.GALAXIIDAEGalaxias auratus Johnston, USNM 344888, 85.6mm. Plate 40
Description.LEI on mid-posterior edge of Ebl, attaches to dor-
sal tip of Pbl at origin.LE2 on mid-posterior edge of Eb2.LE3 tiny, jointly with OD3 insertion on cartilagi-nous tip and lateral edge of Eb3 uncinate process.Remarks. LE3 reduced relative to LEI and LE2,similar to Lovettia.LE4 dorsodistally on Eb4* with posterodistalstrand extending ventrally and attaching to ventro-posterior surface of Eb4* distal end.LP absent.LI1 on Pb2 dorsal surface anteromedially.LI2 on Pb3 dorsal surface posteromedially.LI3 on Pb4 dorsal surface.TD undifferentiated sheet of muscle comprisingTPb3-Pb4-Eb4*, attaching to Pb3 medial to OD or-igins, Pb4 medial to LI3 insertion, and medial end ofEb4*.OD3 and OD4 are bilaterally asymmetrical. OD3originates dorsally on cartilaginous anterior end ofPb3 on both sides. Left-side origin well separatedanteriorly from OD4 origin on Pb3 bony dorsal sur-face, insertion on Eb3 cartilage-tipped uncinate pro-cess medial to and together with LE3; however,strand of muscle at anterior end of OD4 origin (wellseparated posteriorly from OD3 origin) also insertswith OD3. Right-side OD3 origin and insertion sim-ilar to left-side, but strand of muscle originates atposterior end of origin and inserts on Eb4* togetherwith OD4 insertion. Right-side OD3 origin, exceptfor strand, also well separated from OD4 origin.OD4 origin on both sides, except for strand on left
side, from multiple separate muscle strands on Pb3bony dorsal surface, combining to insert on Eb4*dorsodistal end medial to LE4 insertion.OP muscle band originating dorsally on dorsome-dial edge of Eb4*, questionably separate dorsome-dially from SO, joining ER dorsally at mid-posteriorlevel of Eb4* distal end; undifferentiated from SOventral to ER.Remarks. Configuration of OP, Ad5, SO, and ERsimilar to that of Lovettia.Ad 1-3 absent.Ad4 dorsally, broadly on posterior edge of Eb4*,ventrally on Cb4 medial to inner angle formed byEb4*-Cb4 joint.Ad5 broadly on posterodistal end of Cb5, dorsallynarrowly attaching to mid-posterodistal end of Eb4*,medially indistinguishable from SO.RDs absent.SO longitudinal muscle layer absent posterior tohorizontal between distal ends of Cb5s, present asisolated thick muscle band dorsally anterior to (andabove) horizontal connecting distal ends of Cb5s, andcontinuing anteriorly only to about horizontal con-necting medial ends of Eb4s*.Additional remarks. SCL absent. TV4 attached toCb5. Several tiny accessory cartilages on each sideof gill arches. Small cartilage just anterior to anteriorcartilaginous tip of Pb3 on both sides (possibly rep-resenting a segmentation of the anterior tips of Pb3).Anteriormost AC on left side between Ebl cartilag-inous medial end and anterior cartilaginous end ofPb2 just anterior to bony portion; anteriormost onright side between medialmost tip of Ebl and adja-cent cartilaginous edge of Pb2; posteriormost carti-lage on left side filamentous, parallels posteromedialedge of Pb2; posteriormost (not illustrated) on rightside, very tiny, between cartilaginous tips of Pb3-Eb3joint. Ebl and Eb2 lack uncinate processes.
Lepidogalaxias salamandroid.es, USNM 358461,43.7 mm; Murdoch University, 47.3 mm.Plate 41
Description.Remarks. The muscles are highly modified andconfusing. Our interpretations of TPb3, OD3, OD4,and OD4' are provisional.LEI on Ebl bony surface dorsodistally.LE2 on Eb2 at base and lateral to uncinate process.LE3 on dorsodistalmost end of Eb3 and ligament(not shown) extending from insertion to distal end ofCb4.LE4 on anterodistal surface of Eb4, muscle incross section V-shaped with apex directed medially.LP absent.LI1 on dorsoposterior surface of Pb2.
NUMBER 11 55
LI2 absent.LI3 unusually large, on Pb4 dorsally, insertionending posteriorly at margin of Pb4 with UP5? (notshown).Remarks. In the larger specimen, LI3 also appearsto be attached to UP5?, but damage during dissectionleaves this observation problematic.TD appears to comprise only broad TPb3, whichattaches on Pb3 dorsally, joining raphe anterolaterallywith OD3, extending posteriorly as complexly ori-ented fibers, interrupted by irregular, somewhat me-dian raphe, and joining another raphe on each sidemedial to LI3. OD4' appears to originate at latterraphe.OD3 passes lateral to LI3, origin on Pb3 anteriorend, joining raphe there with TPb3, insertion on Eb3dorsodistally.OD4 a muscle strap originating on Eb3 along ra-phe with OD3 and attaching to Eb4 dorsodistally be-tween insertion of LE4 and OD4'.Remarks. This muscle could be inteipreted as aRecD4, but other than its origin on Eb3, an apparentslight shift laterally from the highly reduced, tiny,rod-like, edentate Pb3, its insertion is fairly typicalof OD4, and the relationship with OD3 is reminiscentof the more-or-less in-line origins of OD3 and OD4in other pre-acanthomorphs.OD4' appears to originate at a raphe with TPb3and, on one side, RD well medial to LI3, and is in-terrupted by another raphe immediately medial to LI3before attaching to dorsal surface of Eb4 just medialto OD4. OD4' lies dorsal to unnamed band of muscleattaching anteriorly to posteromedial surface of Eb4and joining raphe posteriorly with RD. We were un-able to determine if OD4' attaches to Pb4 or UP5.OP dorsally on Eb4 posterior edge medial to Ad4dorsally, muscle ending ventrally at ER, which con-tinues laterally along dorsal end of Ad5, becomingtendinous and attaching to Eb4 posteroventrally andCb4 posteriorly (ER tendinous attachments to Eb4and Cb4 (obscured on Plate 41); OP undifferentiatedfrom SO medially.Ad 1-3 absent.Ad4 dorsally on Eb4 posterolaterally, ventrally onCb4 medial to Eb4-Cb4 joint.Ad5 on Cb5 posterodistally, joining ER dorsally,and forming tendinous attachment to Eb4 and Cb4.SO longitudinal fibers absent anterior to horizontalthrough posterior ends of Eb4s, unless represented byRDs; condition of specimen precluded determinationof posterior extent, unless indicated by posterior endsof RDs.SOD absent.RDs completely external to SO, broadly insertedon raphe with unnamed muscle band on one side andwith muscle band and OD4' on other side.Additional remarks. SCL absent. TV4 divided, free
from Cb5, medial end of each division attaching toventral surface of bony Bb4. Ebl and Eb3 lack un-cinate processes. UP4 questionably absent. Eb4 andCb4 appear to have unmodified distal cartilaginousends (see also Rosen, 1974:fig.l5B). with no trace ofEb5.
Lovettia sealei (Johnston), USNM 358617, 2 speci-mens, 50.2-54.2 mm.Plate 42
Description based primarily on smaller specimen.LEI on Ebl dorsoposterior edge somewhat medialto mid-length of bone.LE2 weakly developed (easily overlooked or de-stroyed), on Eb2 dorsoposterior edge somewhat me-dial to mdi-length of bone.LE3 greatly reduced, on finger-like dorsal exten-sion (modified, displaced uncinate process?) of car-tilaginous distal tip of Eb3.Remarks. LE3 reduced relative to LEI and LE2,similar to Galaxias LE2.LE4 on Eb4 :,: dorsodistally.LP absent.LI1 on Pb2 dorsomedially.LI2 on Pb3 dorsoposteriorly.LI3 on Pb4 dorsoposteriorly.TD not clearly partitioned, comprises TPb2-Pb3-Pb4-Eb4*, beginning as few muscle strands on Pb2,continuing posteriorly onto Pb3 and Pb4, and medialends of Eb4*, slightly differentiated posteriorly fromSO.OD3 origin on Pb3 dorsoanteriorly, insertion ondorsomedialmost edge of finger-like extension of car-tilaginous distal tip of Eb3.OD4 on Pb3 dorsoposteriorly and Pb4 dorsoanter-iorly, with slight separation of muscle between Pbs;insertion on dorsomedialmost cartilaginous edge ofdistal end of Eb4*.Remarks. Left-side OD4 of larger specimen has
split origins on Pb3.OP relatively small strap of muscle dorsally onposterior surface near medial end of Eb4*, joiningER ventrally, not distinguished from Ad5 or from SObelow ER.Remarks. Configuration of OP, Ad5, SO, and ERsimilar to that of Galaxias.Ad 1-3 absent.Ad4 on dorsoposterior surface of Eb4*, ventrallyon Cb4 anterior to inner angle formed by Eb4* andCb4.Ad5 on mid-distal end of Eb4* and posterodistalend of Cb5, not separable medially from SO and/orOP.SO, unable to determine by gross dissection if lon-gitudinal muscle layer is present.
56 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
RDs absent.Additional remarks. SCL absent. TV4 divided, freefrom Cb5, medial end of each side attached to ventralsurface of Bb3. Ebl, Eb2, and Eb3 uncinate pro-cesses absent. UP4 and UP5 absent.OSMERIDAEHvpomesus pretiosus (Girard), 2 specimens, USNM357404, 135 mm; USNM 70839, 172 mm.Plate 43
Additional material.104700, 148 mm. Mallotus villosus, USNM
Description.Remarks. The levators of the smaller Hvpomesusspecimen have relatively long tendinous origins,whereas the origins of the larger specimen (and thoseof Mallotus) are relatively short and mostly muscu-lous.LEI slender, on dorsodistal edge of Ebl uncinateprocess.LE2 slender, on dorsal tip of Eb2 uncinate process.LE3 on dorsal tip of Eb3 uncinate process.LE3', thin, applied closely to anterior surface ofLE3, insertion on Eb3 uncinate process just ventralto LE3 insertion.Remarks. The flat "anterior" surface of LE3 ispositioned parallel to the body midline, as is usual infishes, and the flat surface of LE3' is applied ("lam-inated") to the flat surface of LE3. The muscles orig-inate together and their distinction might escape ca-sual examination.LE4 originates in cluster with other LEs; insertson dorsalmost edge of Eb4* levator process.LE4' very slender, closely applied for almost entirelength to distal edge of LE4, origin not divergingmuch from that of LE4, insertion along edge of LE4.
?
Origin diverging slightly but noticeably from thatof LE4, but attached for most of length to distal edgeof LE4, becoming tendinous toward insertion.Remarks. Not considered LP because origin is withclustered LEs.LP absent.LI1 on dorsoposterior surface of Pb2.LI2 on dorsomedial surface of Pb3 posterior end,
at and lateral to attachment of TPb3p.LI3 on Pb4 dorsal surface at and lateral to TPb4-Eb4* attachment.TD comprises three parts: TPb3a-Eb2, TPb3p, andTPb4-Eb4* (described separately for Mallotus).TPb3a-Eb2 most distinct, dorsally situated relative toremainder of TD, with lateral attachments to joinedtips of Pb3 and Eb2 uncinate processes, and crossedlateral attachments to dorsoanterior Pb3 bony surfac-es dorsal to OD3 origins, completely separate fromTPb3p in larger specimen, continuous posteriorly by
diagonal fiber bundle with TPb3p in smaller speci-men. TPb3p on Pb3 dorsal surface posterior to un-cinate process and medial to LI2 insertion, continu-ous by diagonal muscle fibers with TPb4-Eb4*.TPb4-Eb4* on Pb4 dorsolateral^ medial to LI3, andon dorsomedial edge of Eb4, posteriorly almost in-separable posteriorly from SO.
?
Comprises TPb3a, TPb3p, and TPb4-Eb4*.TPb3a on Pb3 uncinate process laterally, ventrome-dially attached to CT between well-separated anteriorends of Pb3s (Pb3s of Hypomesus almost impingealong medial edges), no crossed lateral attachmentsto Pb3s dorsoanteriorly dorsal to OD3 origin, dor-sally situated relative to remainder of TD, continuousposteriorly with TPb3p posteriorly. TPb3p on Pb3laterally posterior to uncinate process, continuousposteriorly with TPb4-Eb4a*. TPb4-Eb4* on Pb4dorsolateral^ and Eb4* medially, posteriorly distin-guished from SO by change in muscle fiber direction.OD3 origin on dorsoanterior bony surface of Pb3ventral to TPb3a-Eb2, insertion on Eb3 uncinate pro-cess medial to LE3 and LE3' insertions. ? Originventral to TPb3a.OD4 origin on posteriormost end of Pb3 and dor-soanterior surface of Pb4, insertion on medial edgeof Eb4*. ? OD4 origin on Pb3 dorsoposteriorly andOD4' on Pb4 dorsoanteriorly, muscles joining justposterior to OD4' origin, but somewhat (artificially?)separable for entire length, with OD4 inserting onEb4* dorsoanteriorly and OD4' inserting on medialedge of Eb4* ventromedial to OD4.OD4' absent. ? Anteriorly on Pb4 dorsoanteriorlyand posteriorly on Eb4* medial edge; just posteriorto origin, joins OD4, but is somewhat separable (ar-tificially?) for entire length.OP strap of muscle dorsally on dorsomedial edgeof Eb4*, partly dorsal to attachment of TPb4-Eb4*to Eb4*; ventrally joins ER with SO; indistinguish-able ventral to ER.Adl-3 absent.Ad4 dorsally on dorsoposterior edge of Eb4*, ven-trally on Cb4 just medial to inner angle formed byEb4*-Cb4 joint.Ad5 on posterodistal end of Eb4* and posterodistalend of Cb5, weakly separable from SO (or OP?) me-dially. ? On posteriorly extending finger-like carti-laginous extension of Eb4* distal end and posterodis-tal end of Cb5.RDs absent.SO longitudinal fibers begin well posterior to hor-izontal through posterior ends of Cb5s and continueanteriorly to horizontal between anterior ends of Pb4s(posterior to TPb3p).Additional remarks. SCL attached posteromediallyto ventroposterior cartilaginous tip of Bb3?ligamentalmost circular with massive muscle attachments; ?SCL absent. TV4 free from Cb5s, divided at mid-
NUMBER 11 57
line, attaching dorsally to ventral surface of Bb series(specific element not determined). Smaller specimenwith two tiny ACs just anterior to anteroventral sur-face of Pbl on left side, one on right side; largerspecimen with one AC (not illustrated) on right sideonly. Johnson and Patterson (1996:277) discuss prob-lems concerning the identification of the upper pha-ryngeal tooth plate, or plates, as representing UP4 orUP5, but appear to favor identifying it as UP5.The posterior end of Eb4 has the foramen for thefifth efferent artery bordered completely by bone an-teriorly and by cartilage posteriorly in the largerspecimen of Hypomesus; the cartilaginous portion isinterrupted in the middle in the smaller specimen (seeRosen, 1974:fig. 16D). The posterior end of Eb4 inthe specimen of Mallotus is similar to that of thelarger specimen of Mallotus except that there is avery fine joint line (both right and left arches) be-tween the dorsomedial end of "Eb5" and the dor-solateral cartilaginous end of Eb4. The condition inour experience is exceptional. Eb5 usually appears tofuse to Eb4 dorsally first, and ventrally last (see com-ments on clupeomorphs by Rosen, 1974:280). Rosen(1974:284), however, states that the opposite is truefor osmeroids: fusion takes place ventrally first; how-ever, he provided no evidence for this. Based on hisillustrations of the osmeroid taxa (his fig. 16), allhave either a free Eb5 or a fused Eb5 that can beinterpreted alternatively as having had Eb5 fusedcompletely with Eb4 and subsequently variously andpartially lost, with no unequivocal indication whetherfusion took place dorsally or ventrally first.
ArgentiniformesARGENTINIDAEArgentina brucei Cohen and Atsaides, USNM238005, 99.5 mm. Plate 44
Description.LEI on Ebl at and just lateral to base of uncinateprocess.LE2 on dorsolateral edge of Eb2 uncinate process.LE3 on dorsolateral edge of Eb3 uncinate process.LE4 on dorsolateralmost tip of Eb4 levator processjust dorsal to insertion of OD4 anteriorly and dorsalattachment of Ad4 posteriorly.LP absent.LI1 on Pb2 mid-dorsally.LI2 on Pb3 dorsoposteriormost bony surface, be-ginning just lateral to OD4 origin.LI3 on Pb4 dorsoposteriorly.TD long, broad, essentially undifferentiated, com-prising TPb3-Pb4-Eb4, which attaches along dorso-medial surfaces of Pb3 and Pb4 and meeting inser-tions of OD3 and OD4, and with narrow attachment
to posteromedial edge of Eb4; posteriorly continuouswith SO.OD3 origin mid-dorsally on Pb3 just medial to LI2insertion, insertion on Eb3 uncinate process ven-troanterior to LE3 insertion.OD4 origin broadly on Pb3 beginning at posteriorend of OD3 origin, insertion on Eb4 levator processventroanterior to LE4 insertion.OP strap of muscle overlying SO dorsolaterally,dorsally on Eb4 posteromedial surface, ending ven-trally at ER (right side ER very weak), scarcely sep-arable ventrally from Ad5 and SO.Ad 1-3 absent.Ad4 dorsally on posterodorsal surface of Eb4, ven-trally on dorsal surface of Cb4 anterior to inner angleformed by Eb4-Cb4 joint.Ad5 dorsally on mid-distal surface of Eb5, ven-trally on posterodistal bonysurface of Cb5 ventrally,forming raphe with posterolateral end of TV5, me-dially continuous with SO.SO with strap of muscle arising laterally, extendingdorsally, then ventrally and attaching to anterior sur-face of Eb5 to form wall of pocket lined anteriorlyby Ad4; continuous ventrolaterally with Ad5 (or?OP). Longitudinal fibers commencing well posteriorto horizontal between distal ends of Cb5s and con-tinuing anteriorly, thinly to horizontal connecting me-dial ends of Eb3.RD absent.Additional remarks. SCL absent. TV4 free fromCb5s. Eb5 large, on dorsodistal end of Cb4, articu-lating ventroanteriorly with posterodistal end of Eb4,attached dorsally by long, slender ligament to Eb4levator process. Accessory cartilage on dorsodistalend of Cb5, surrounded by tubular CT which is at-tached to mid-posterior surface of Eb5.
ALEPOCEPHALIDAE
Alepocephalus tenebrosus Gilbert, USNM 215582,141 mm. Plate 45
Description.Remarks. LEI and Lis together surround and at-tach to dorsal end of very long, slender SPbl and tocranium; LI1 also attaches along dorsomedial surfaceof SPbl.LEI slender, on dorsolateral edge of Ebl uncinateprocess.LE2 slender, on dorsodistal edge of cartilaginousmedial end of Eb2.Remarks. Medial cartilaginous end of Eb2 expand-ed dorsally, and dorsally does not articulate with Pb3;Eb2 uncinate process absent, unless modified dorsalexpansion of medial end represents shift in positionof uncinate process.
58 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LE3 slender, on cartilaginous tip of Eb3 uncinateprocess.LE4 very slender, inserts finely, tendinously onmid-dorsal cartilaginous edge of distal end of Eb4.LP absent.LI1 broadly on bony dorsal surface of Pb2, dor-sally on SPbl beginning a little ventral to attachmentto skull and continuing to muscle origin.LI2 slender, on dorsal surface of bony and carti-laginous posterior end of Pb3.LI3 from single origin, spreads ventrally, becom-ing fan-like and inserting on Pb4 as group of sepa-rated muscle strands; on right side only, anteriormostmuscle strand is also attached to cartilaginous medialtip of Eb3.TD comprises short TPb3a, much longer TPb3p-Pb4, and short TEb4. TPb3a attaches to bony dorsalsurface of Pb3 anterior processes and is noticeablydistinct, although broadly continuous posteriorly withTPb3p-Pb4. TPb3p-Pb4 attaches along the dorso-medial margins of Pb3 and Pb4 and is broadly con-tinuous posteriorly with TEb4. although separatedfrom TEb4 by a posterolateral notch in the muscles
at the posterior end of Pb4 and anterior end of Eb4.TEb4 attaches along anteromedial margin of Eb4 andbecomes continuous posteriorly with SO by broadarea of crisscrossing muscle fibers.OD3 slender, origin on medial surface of Pb3 un-cinate process, with fibers extending anteriorly andjoining posterolateral edge of TPb3; insertion on dor-soanteriormost edge of Eb3 uncinate process.OD4 slender, origin on posteroventral edge of Pb3uncinate process ventral to OD3 origin; tendinous in-sertion on dorsoanterior cartilaginous edge of Eb4,continuous at insertion with OD4' insertion.OD4' slender, origin on anterodorsal surface ofPb4; insertion on Eb4 continuous with OD4 inser-tion.OP absent, but possibly represented by stringymuscle strands attaching to posteromedial surface ofEb4; strands become complexly meshed posterov-entrally with SO muscle strands (complexity not il-lustrated). ER absent, but possibly represented byarea where problematic OP muscle strands mesh withSO strands.Ad 1-3 absent.Ad4 on posterior surface of dorsodistal end of Eb4and dorsoposterior surface of Cb4 at internal angleformed by Eb4-Cb4 joint.Ad5 attaching cartilaginous Cb5 distal end poste-
riorly to ventral margin of large Eb5.SO longitudinal fibers loosely incorporated influffy brown matrix beginning well posterior to hor-izontal between posterior ends of Cb5s, and decreas-ing in density abruptly at horizontal between anteriorends of Eb4s; very thin matrix, possibly including
sparse muscle fibers, continuing into pharyngeal areaanterior to gill arches.RDs absent.Additional remarks. SCL absent. TV4 free fromCb5s. Cartilaginous end of Pbl ventromedial processarticulating with anterior end of Pb2, ventrolateralcartilaginous end articulating with anterior cartilagi-nous end of Ebl. Pbl and tips of Pb2 and Pb3 en-veloped in loose CT pad (not illustrated); pad at-tached posteriorly to long, slender ligament, whichjoins contralateral ligament at attachment to cranium.Large Eb5 present posteriorly, articulating ventral-ly with dorsal edge of posterior ends of Cb4 and Eb4;large AC articulating ventrally with Cb5 and Eb5 me-dially; right side only with two additional tiny ac-cessory cartilages laterally on Eb5. SPbl present.There is great similarity in the shapes and "lay-out" of the muscles of Alepocephalus and Searsia,especially when compared with their sister group, Ar-gentina. PLATYTROCHTIDAESearsia koefoedi Parr, USNM 206873, 117 mm.Plate 46
Description.Remarks. LEI and Lis surround and attach toSPbl ventral to origins on cranium (all levators orig-inate together on cranium).LEI slender, inserts by slender tendon on dorso-posterior edge of Ebl lateral to uncinate process.LE2 absent.LE3 slender, on cartilaginous tip of Eb3 uncinateprocess.LE4 very slender, finely, tendinously on mid-dor-
sal cartilaginous edge of Eb4.LP absent.LI 1 broadly on dorsomedial surface of bony por-tion of Pb2, dorsally on SPbl near insertion on skull;posteroventral fibers of left-side LI1 attach (anoma-lously) to lateral edge of Pb3.LI2 on posterodorsal surface of Pb3; right sidewith few fibers attaching to anteriormost surface ofPb4, dorsally difficult to separate from LI3'.LI3 on Pb4 posterolaterally; lightly attached to me-dial surface of SPb2.LI3' slender, but moderately broadly on Pb4 dor-soanteriorly lateral to origin of OD4'; right side LI3'both anterior and lateral to origin of OD4', dorsalfibers not clearly separable from LI2.TD comprises short TPb3a, which attaches to bonyPb3 anterior process, and much longer TPb3p-Pb4-Eb4, which attaches along dorsomedial edges of Pb3,Pb4, and Eb4, and is broadly continuous with SOposteriorly.OD3 origin on medial surface of Pb3 uncinate pro-cess; insertion on dorsomedialmost edge of Eb3 un-
NUMBER 1 1 59
cinate process: left side OD3 lies dorsal to and almostcompletely hides OD4; right-side hides all but anter-olateralmost OD4.OD4 origin with and ventral to OD3 origin on Pb3uncinate process; insertion on dorsomedial cartilagi-nous edge of Eb4, continuous posteriorly with pos-terior portion of OD4'.OD4' origin on anterodorsal surface of Pb4; leftside fuses with OD4 at about mid-length; right sidefuses with OD4 just anterior to insertion on Eb4.OP absent, but possibly represented by stringymuscle strands attaching to posteromedial surface ofEb4; strands become complexly meshed posteroven-trally with SO muscle strands (complexity not illus-trated). ER absent, but possibly represented by areawhere problematic OP muscle strands mesh with SOstrands.Ad 1-3 absent.Ad4 on dorsomedial surface of Eb4 and postero-dorsal surface of Cb4 at inner angle formed by Eb4-Cb4 joint.Ad5 narrowly on Eb4 ventrodistally and broadlyon Cb5 ventrodistally; right side with few fibers at-taching to AC.SO longitudinal fibers loosely incorporated influffy brown matrix beginning well posterior to hor-izontal between posterior ends of Cb5s, decreasing indensity and ending abruptly, or nearly so, at abouthorizontal between posterior thirds of Pb4s (matrix,
at least, continuing anteriorly into pharyngeal areaanterior to gill arches).RDs absent.Additional remarks. SCL present with dorsomedialCT pad attaching to Bb3 posteroventral tip. TV4 freefrom Cb5s. Pbl with cartilaginous end of ventro-medial process articulating with anterior tip of Pb2;ventrolateral process with two separated cartilaginoustips, articulating with anterior cartilaginous tip ofEbl. Pbl and tips of Pb2 and Pb3 enveloped in toughCT pad. Eb5 articulating laterally with Eb4 and me-dially with AC, which articulates ventrally with Cb5.Cartilaginous SPbl on dorsal cartilaginous tip ofEbl uncinate process, SPb2 on dorsal cartilaginoustip of Eb2 uncinate process. Johnson and Patterson(1996:275) reported that S. koefoidi has only SPbl.Their report was based on SIO 77-38 and 77-53(Johnson and Patterson, 1996:323), collected in theBanda and Sulu seas, whereas our specimen was col-lected in the northeastern Atlantic (type locality,however, is Bahamas). Matsui and Rosenblatt (1987:73) reported that, "the width of the frontals differsbetween S. koefoedi of the 3 ocean regions." It ap-pears likely that more than one species is involved.There is great similarity in the shapes and "lay-out" of the muscles of Alepocephalus and Searsia,especially when compared with their sister group, Ar-gentina.
EsociformesESOCIDAE
Eso.x niger Lesueur, USNM 355803, ca. 235 mm.Plate 47
Additional material. ? = Esox lucius Linnaeus,USNM68894, 87.5 mm.Remarks. Poor condition of E. lucius specimenprovided only limited amount of information, but nonoteworthy differences were observed in the musclesof the two species.
Description.LEI broadly, fibrotendinously on cartilaginous capof Ebl uncinate process.LE2 on tip of Eb2 uncinate process.LE3 on tip of Eb3 uncinate process.LE4 absent.LP absent.LI1 massive, broadly on Pb2 bony surface dorso-medially.LI2 broadly on Pb3 bony surface dorsoposteriorly.LI3 on Pb4 dorsomedial surface.TD comprises TPb3a and TPb3p-Pb4. TPb3a at-taches along lateral edge of anterior half of Pb3, isdorsal to origin of OD3. and posteriorly continuousby diagonal strand of muscle with TPb3p-Pb4.TPb3p-Pb4 attaches to Pb3 and Pb4 dorsal surfacesmedial to insertions of LI2 and LI3, joins dorsolateralraphe with OD4 origin, with which TPb3p-Pb4 isotherwise continuous, and is also continuous poste-
riorly with SO.OD3 origin on dorsal surface of Pb3 ventromedialto attachment of TPb3a, insertion on Eb3 uncinateprocess proximal to LE3 insertion.OD4 absent. See remarks following OD4'.OD4' originates dorsally from raphe with TPb3p-Pb4 and ventrally from Pb4 dorsal surface, and in-serts on Eb4 uncinate process.Remarks. OD4' of Esox is similar to OD4' of theputatively related Novumbra, particularly in that bothhave their main origins on Pb4 rather than Pb3. No-vumbra has OD3-4 fused or separate (OD3 andOD4) as well as OD4'.Johnson and Patterson (1996:275) argued that eso-coids may have an uncinate process, but not a levatorprocess. As far as we know, an Eb4 levator processnever articulates with the Eb3 uncinate process,whereas, the Eb4 uncinate process usually, if not al-ways, does, thus supporting the Johnson and Patter-son's interpretation of the process in Esox. The pres-ence of an Eb4 uncinate process, and its articulationwith the Eb3 uncinate process, in Esox is either anautapomorphy of Esox, or evidence that its closestrelationships are with the basal acanthomorphs, inwhich these states first occur.
60 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONOP indistinct, probably represented dorsally bymuscle attaching to posterior surface of Eb4 begin-ning ventral to uncinate process and extending me-dially, and ending ventrally at ER on Cb4, therejoined by Ad5 dorsally. This configuration OP, ER,and Ad5 also appears in Saurida (Synodontidae) andsome acanthomorphs, e.g., Polymixia.Ad 1-3 absent (condition of GFMs not deter-mined).Ad4 dorsally appears to attach along entire poste-rior surface of Eb4, although medial half probablyrepresents OP, which ventrally joins ER with Ad5.Ad5 on Cb5 distally and bony posterior surface ofCb4, where it joins ER dorsally with presumable OPventrally.RDs absent.SO longitudinal muscle layer originates well pos-terior to gill arches and extends anteriorly to hori-zontal at anterior ends of Pb4s, abruptly decreasingor vanishing anterior to horizontal.Additional remarks. SCL attached dorsomediallyto posteroventral cartilaginous end of Bb3. TV4 freefrom Cb5s (completely anterior to anterior tips ofCb5s). UP5 absent. Eb 5 absent.UMBRIDAENovumbra hubbsi, USNM 357403, 2 specimens,57.2-59.7 mm. Plate 48
Description.Remarks. In general the two dissected gill archeswere very similar, but because of small size and fra-gility, both our dissections involved damage. The de-scription and illustrations are mostly based on thelarger specimen, but the posterior view is based onthe smaller specimen.LEI on Ebl uncinate process at and just ventro-lateral to cartilaginous tip.LE2 on Eb2 uncinate process at and, variably, justventrolateral to cartilaginous tip.LE3 on tip Eb3 uncinate process dorsoanteriorly.LE4 weak, on Eb4 dorsoposteriorly near distalend, ventral fibers continuous with Ad4 dorsoposter-iorly.Remarks. Johnson and Patterson (1996:273) re-ported that Novumbra lacks LE4. The muscle is pres-ent in both of our specimens. Because LE4 is poorlydeveloped and specimens are small, the muscle couldhave been easily lost during dissection.LP absent.LI1 dorsomedially on bony surface of Pb2.LI2 on Pb3 dorsoposteriorly.LI3 on Pb4 dorsolaterally meeting junction of Pb4with UP4.TD complex. Distinct TPb3a anteriorly, attachingto Pb3 uncinate process anteriorly, continuous pos-
teriorly by crisscrossing muscle bands (both speci-mens) with TPb3p. TPb3p irregular, attaching to dor-soposteromedial surface of Pb3, continuous posteri-orly with TPb4 by continuations of crisscrossingmuscle bands, which meet raphe with OD4' posteri-orly. TPb4 attaching to Pb4, muscle continuous withSO posteriorly.OD3-4 comprises a fused single muscle in thelarger specimen (illustrated) and separate muscles(OD3 and OD4) in the smaller specimen. The fusedmuscle in the larger specimen can be artificially sep-arated, with the OD3 portion much more slender thanthe OD4 portion (relative sizes not apparent in illus-tration). The fused muscle divides slightly at its in-sertions on Eb3 and Eb4, which are similar to theinsertions of the two muscles in the smaller speci-men. In the smaller specimen, the two muscles areof about equal size and are described as folows:OD3 origin anteriorly on Pb3 dorsal surface ven-tral to TPb3a, insertion on medial edge of Eb3 un-cinate process.OD4 origin on Pb3 dorsal surface posterior to andcontinuous with OD3 origin, insertion on dorsome-dial all bony process of Eb4, fuses with insertion ofOD4'.OD4' origin dorsally from raphe with crisscrossingmuscle bands of TPb3p and mainly ventrally on dor-
sal surface of Pb4, insertion on dorsomedial bonyprocess of Eb4, fuses with insertion of 4 portion ofOD3-4.Remarks. OD4' of Novumbra is similar to OD4'of the putatively related Esox, particularly in thatboth have their main origins on Pb4 rather than Pb3.There is also a similarity with OD4' of Lepidoglaxias(Plate 41).OP on mid-dorsoposterior margin of Eb4, endingventrally at ER on Cb4 with Ad5.M. Pb3-Eb3 small, attached to dorsoposterior edgeof Pb3 uncinate process and dorsomedial edge of Eb3uncinate process near LE3 insertion.Ad 1-3 absent, but GFM2 and 3 unusual, superfi-cially like RecDs. GFM2 originates on dorsoanteriorsurface of Eb2 uncinate process and inserts on dor-soposterior edge of Eb 1 uncinate process, where it iscontinuous with LEI basally (not continuous basallywith LEI in smaller specimen). GFM3 (not illustrat-ed) slender muscle seen only on right side of smallerspecimen, originates on anterior margin of Eb3 andinserts on posterior surface of Eb2 uncinate process.Remarks. Similar GFM2 and 3 are known amongpre-acanthomorphs otherwise only in Dallia (Umbri-dae), which has a modified GFM1. GFM1 in Novum-bra was destroyed before it was realized that its char-acter state might have importance.Ad4 on Eb4 dorsoposteriorly and Cb4 dorsally justmedial to Eb4-Cb4 joint, just meeting and joining ER
at dorsodistalmost end of Ad5.
NUMBER 1 1 61
Ad5 on posterodistal surface of Cb4, joining ERwith ventral end of OP, and on dorsoposterior portionof Cb5. forming raphe ventrally with TV5.SO longitudinal muscle fibers begin well posteriorto horizontal between posterior ends of Cb5s and ex-tend anteriorly to below mid-posterior margin ofTPb3a.RDs absent.Additional remarks. SCL attached mid-dorsally toposteroventrally extending cartilaginous end of Bb3.TV4 free from Cb5s. Eb4 uncinate process bony.UP5 absent. Eb5 absent.
Dallia pectoralis Bean, USNM 11 1643, 107 mm.Not illustrated
Description.LEI on anterior surface of Ebl uncinate process.LE2 on anterior surface of Eb2 uncinate process.LE3 dorsoanteriorly on Eb3 uncinate process.LE4 mostly on Eb4 bony dorsolateral surface, ex-tending onto cartilaginous distal end.LP absent.LI1 on dorsomedial bony surface of Pb2.LI2 on Pb3 dorsally.LI3 on Pb4 dorsally.TD comprising TPb3 and TPb4. TPb3 shorter andwider of the two transverses, attaching to Pb3 dor-soanterolaterally; on left side attachment is dorsal toOD3 and OD4 origins; on right-side, posterolateraledge forms raphe with OD3 and OD4 origins andwith slender muscle (anomalous?) extending laterallyand inserting on Eb2 uncinate process; continuousbroadly posteriorly with anterior end of TPb4. TPb4on dorsal surface of Pb4 medial to LI3 insertion, con-tinuous posteriorly with SO, but noticeably distin-guished by change from horizontal (TPb4) to curvingmuscle fibers (SO).OD3 origin on Pb3 (see TPb3 above), insertion ondorsomedial edge of Eb3 uncinate process.OD4 origin on Pb3 (see TPb3 above), insertinglaterally on Eb4 dorsomedial edge, extending medi-ally along raphe with dorsal end of OP (left-side OD4posterolaterally and OP dorsally forming free strapof muscle?not attached ventrally; right-side raphealmost completely attached to dorsal edge of Eb4).OP dorsally joining raphe with OD4 (q.v.), ven-trally on posterodistal end of Cb5, forming raphe (notER, unless highly modified) posteroventrally withAd5, posterolaterally overlapping Ad4 and apparent-ly fusing with Ad4 (q.v.) ventroanteriorly on Cb5.Ad 1-3 absent, but GFM1-3 unusual; GFM2 andGFM3 RecD-like, similar condition observed only inNovumbra. GFM 1 originates by long, slender tendonattaching to anterior surface of cartilaginous medialend of Ebl, becoming musculous in area anterior to
uncinate process, and covering (attaching to) most ofdorsal surface of Ebl lateral to distal end of Ebl,there becoming slenderer and extending around Ebl-Cbl joint and continuing along dorsoanterior edge ofCbl. GFM2 inserts by short, fine tendon to posterioredge of Ebl ventral to uncinate process, muscle isover CT between Ebl and Eb2 and attaches to an-terior edge of Eb2. GFM3 similar to GFM2, insertionon Eb2 and origin on Eb2.Ad4 dorsally broadly on Eb4 beginning anterior tolateral end of OP and extending to end of bone; an-teroventrally on Cb4 dorsal surface medial to Eb4-Cb4 joint; posteroventrally attaching to cartilaginousfinger-like process extending medially from poster-odistal end of Cb4, there joining raphe with Ad5, andcontinuing ventrally and (unusually) attaching to Cb5anterior to OP, with which it appears to fuse.Ad5 very short, joins dorsodistal end of Cb5 toposterodistal end of Cb4, joins raphes with OP andAd4.SO beginning well posterior to horizontal throughposterior ends of Eb4s and extending anteriorly toposterior margin of TPb3.RDs absent.Additional remarks. SCL interrupted mid-posteri-orly by attachment to posteroventrally extending car-tilaginous end of Bb3. TV4 attached to Cb5s, exceptfor free short area at anterior end of muscle. UP5absent. Eb5 absent.
Umbra pygmaea (DeKay), USNM 343617, 61.1 mm.Plate 49
Description.LEI on Ebl uncinate process.LE2 on Eb2 uncinate process.LE3 on Eb3 uncinate process; right side LE3 withmuscle slip inserting on anteromedial edge of Eb4.LE4 on cartilaginous dorsodistalmost edge of LE4.LP absent.LI1 on Pb2 dorsoposteromedially.LI2 on Pb3 bony dorsal surface; posterior edgecontinuous with anterior edge of LI3, but slight break
at anteroventralmost insertion of LI3 on Pb4.LI3 on Pb4 dorsally (see also LI2).TD comprises single continuous sheet of muscle,TPb3-Pb4-Eb4, continuous posteriorly with SO.OD3 origin on Pb3 anterodorsal surface continu-ous with OD4 origin, insertion on medial edge of Eb3uncinate process ventral to OD4.OD4 origin on Pb3 anterodorsal surface immedi-ately at and posterior to OD3 origin, forming raphemedially with TPb3-Pb4-Eb4, insertion on dorso-medial edge of Eb4 at and just anterior to dorsal at-tachments of Ad4 and OP.OP separate band of muscle dorsally, attaching to
62 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
dorsomedial edge of Eb4 just medial to dorsal at-tachment of Ad4, just posterior to OD4 insertion, andpartially overlying (posterior to) SO attachment toposterior surface of Eb4; joins ER ventrally at aboutmid-level of cartilaginous Eb5, below which OP isinseparable from SO.Ad 1-3 absent (condition of GFMs not deter-mined).Ad4 dorsally on dorsoposterior edge of Eb4 ven-tral to LE4 insertion and lateral to OP attachment,ventrally on Cb4 at and medial to inner angle formedby Eb4-Cb4.Ad5 on distal surface of Eb5 and dorsodistalmostend of Cb5.SO longitudinal muscle fibers begin well posteriorto horizontal at posterior ends of Cb4s and extendanteriorly only to horizontal between mid-length ofPb4s.RD absent.Additional remarks. SCL attached mid-posteriorlyto ventral surface of cartilaginous posterior end ofBb3. TV4 questionably absent (rare condition), butan obliquus ventralis-like muscle attaches ventro-medial surface of each Cb4 to ventral surface of Bb4cartilage. UP5 absent. Eb5 present.StomiiformesDIPLOPHIDAE
Diplophos taenia Gunther, USNM 206614, 190 mm.Plate 50
Description.LEI very fine, on small bony process on lateralmargin of Ebl uncinate process.LE2 very fine, on small bony process ventrolateralto expanded cartilaginous dorsomedial end of Eb2.LE3 slender, on dorsomedial bony edge of Eb3uncinate process on and with OD3 insertion.LE4 originates in cluster with other LEs; insertstendinously on dorsal tip of Eb4 levator process.LP absent.LI1 on most of length of dorsomedial edge of Pb2.LI2 dorsally on posterolateral edge of Pb3.LI3 on dorsolateral edge of UP5, meeting TUP5laterally.Remarks. A slight portion of the posterior end ofUP5 slips under the medial end of Eb4. We followJohnson (1992) in recognizing the single tooth platein stomiiforms as UP5. However, its close associationwith Pb4 suggests that it could be UP4. Further in-vestigation of the homology of this tooth plate is de-sirable.TD comprises TEb2, TEb2v, TPb3, and TUP5.TEb2 broadest, on Eb2 dorsoanteriorly, attaching toexpanded cartilaginous medial end of Eb2 as musclepasses over it, continuous posteriorly by diagonal
strand of muscle with TPb3. TEb2v thin muscletightly attached to CT covering anterior ends of Pb3s,lies anteroventral to TEb2, with thin, slender pos-terolaterally extending straps that attach to antero-medial surface of expanded cartilaginous medial endof Eb2 (verification of presence in other specimensdesirable). TPb3 attaches dorsally on posterolateraledge of Pb3 with and medial to LI2 insertion, con-tinuous posteriorly by diagonal strand of muscle withTUP5. TUP5 attaches to UP5 anteromedial to LI3insertion, is continuous posteriorly by diagonal strandof muscle with SOD.OD3 origin ventral to TEb2 beginning on Pb3 dor-sally a short distance posterior to anterior cartilagi-nous tip of Pb3 and continuing uninterrupted poste-
riorly as OD4 origin to point a little posterior to be-ginning of cartilaginous posterior end of Pb3; OD3separates dorsolaterally from OD4 shortly posteriorto combined origins and inserts on Eb3 uncinate pro-cess anteriorly.OD4 origin on Pb3 continuous with OD3 origin,joins OD4' posterolaterally, and insert together onanterior surface of Eb4 levator process.OD4' originates on Pb4 anteriorly, beginning atedge joining Pb3, joins OD4 posteromedially, and to-gether insert on Eb4 levator process anteriorly.OP dorsally on Eb4 posteroventrally beginningnear medial end and extending laterally a short dis-tance, ventrally on Cb5 slightly posteromedial to dis-tal end; ER absent.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning later-
ally at Eb4 levator process and extending mediallyabout half distance to medial end of bone; ventrallynarrowly on Cb4 dorsoposteriorly medial to inner an-gle formed by Eb4-Cb4 joint; medial edge of muscletendinous.Ad5 dorsally on Eb4 dorsodistally, ventrally onEb5 (not illustrated) and posterodistally on Cb5, join-ing raphe with TV5.Remarks. Ad5 completely occludes the tiny Eb5from view. Fink and Weitzman (1982), using singlestained material, did not report the existence of Eb5in Diplophos taenia (their fig. 10 is based on a spec-imen from the same lot as ours). Baldwin and John-son (1996:372), using double-stained material, re-ported that Eb5 is present in Diplophos taenia.RDs separate by space equal to about diameter ofone RD, muscle inserts on dorsomedial edge of UP5
at junction with Pb4.SO longitudinal muscle fibers questionably pres-ent; requires histological examination.SOD present.Additional remarks. SCL absent. TV4 free fromCb5s. Identity of single UP questionable (see remarksfollowing LI3).
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1
63STERNOPTYCHIDAEMaurolicus muelleri (Gmelin), USNM 323033, 2specimens, 54.7?55.7 mm.Plate 51
Description.Remarks. The muscles are thin and tiny and it ispossible that alternative descriptions we provide inremarks following LE3, LE4, and OD4 may apply.LEI very fine, inserts by extremely fine tendon onmid-dorsoposterior edge of Ebl lateral to uncinateprocess.LE2 very fine, inserts by extremely fine tendon onmid-dorsoposterior edge of Eb2.LE3 on tip of Eb3 uncinate process.Remarks. Possibly comprising a bilateral pair ofclosely applied muscles (LE3, LE3'). based on whatappears to be a divided origin (attempts to separatemuscle into two natural parts were inconclusive).LE4 originates with cluster of other LEs; insertson tip of Eb4 levator process.Remarks. Possibly comprising a bilateral pair ofclosely applied muscles (LE4. LE4'). based on whatappears to be a divided origin (attempts to separatemuscle into two natural parts were inconclusive).LP absent.LI1 on Pb2 dorsomedial surface posterior to un-cinate process.LI2 on dorsoposterior surface of Pb3.LI3 on UP5 dorsally at junction with cartilaginousPb4.Remarks. Johnson ( 1992:fig. 3c) illustrated the up-per pharyngeal bones of Maurolicus and the inser-tions of LI 1-3 and RD. See also remarks regardingUP5 following LI3 in Diplophos, Diplophidae.TD comprises TEb2a, TEb2p, and TPb3-Pb4 (larg-er specimen) or TPb4 (smaller specimen). TEb2aoverlies bony anterior ends of Pb3 and attaches an-teriorly on dorsomedial surface of Eb2, separatedfrom TEb2p by depression. TEb2p on dorsoposteriorsurface of Eb2, extending slightly further distallythan TEb2a, overlies OD4 origins. TPb3-Pb4 slightlyposteroventral and unconnected to TEb2p, attaches todorsomedial edges of Pb3 and Pb4 and is indistin-guishable posteriorly from SOD; TPb4 is separatedfrom TEb2p by a relatively large gap in smaller spec-imen.Remarks. Separation of TEb2 into TEb2a andTEb2p is problematic.OD3 absent.Remarks. Absence of an OD attachment to Eb3,either as OD3 or OD3?4, is an uncommon speciali-zation within the Euteleostei, and occurs only insome Stomiiformes (Table 1) and ateleopodids. Itmay contribute to resolution of the Neoteleostei tri-chotomy (Fig. 3).OD4?4' origin ventral to TEb2p, on most of dor-
soposterior surface of Pb3 and all of dorsal surfaceon Pb4, insertion broadly on dorsolateral surface ofEb4 levator process.Remarks. There is a definite separation of the or-igins on Pb3 and Pb4 in the larger specimen, but littleor none in the smaller specimen. OD4', originatingon Pb4, is present in other stomiiforms we examined,and we infer that OD in Maurolicus represents a fu-sion of OD4 and OD4'.OP absent, ER absent.Ad 1-3 absent.Ad4 dorsally broadly on posterodorsal surface ofEb4 ventral to LE4, ventrally narrowly on Cb4 dorsalsurface just medial to inner angle formed by to Eb4-Cb4 joint.Ad5 dorsally on posterodistal end of Eb4, ventrallyon dorsodistal surface of Cb5, joining raphe withTV5 (not illustrated) ventroanteriorly in smaller spec-imen, but not in larger specimen.RDs separated by distance greater than one RD,on UP5 at junction with Pb4 (see remarks under LI3).SO longitudinal muscle fibers questionably pres-
ent, requires histological verification.SOD present, but broadly continuous with TPb3-Pb4 anteriorly.Additional remarks. SCL absent. TV4 free fromCb5s. Identity of single UP questionable.GONOSTOMATIDAEGonostoma elongatum Giinther, USNM 330309, 162mm. Plates 52.1, 52.2
Description.LEI very fine, inserts by long thread-like tendonto dorsal edge of Ebl lateral to base of uncinate pro-cess.LE2 absent.Remarks. Absence of LE2 rare in fishes.LE3 slender, inserts by very long tendon to carti-laginous tip of Eb3 uncinate process.LE4 relatively large, originates in cluster with oth-er LEs, inserts by tendon on dorsomedial cartilagi-nous end of Eb4 levator process.LP absent.LI1 massive, on almost entire medial edge of Pb2,lateral edge joins or is formed by tendon, which in-
serts on Ebl uncinate process.LI2 on posterolateral edge of Pb3.LI3 on lateral surface of UP5.Remarks. See also remarks regarding UP5 follow-ing LI3 in Diplophos, Diplophidae.TD comprises TEb2 anteriorly and well separatedTUP5 posteriorly. TEb2 attaches broadly on Eb2 an-teromedially, giving rise dorsoanteriorly on each sideto short muscle lamina that attaches to tendon in-serting on Ebl uncinate process, joins raphe with
64 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONOD4 origin beginning at tendon and extending entirelength of OD4 origin. TUP5 a small muscle strapattaching to lateralmost edge of UP5.OD3 absent (see remarks under OD3 in Mauroli-cus, Sternoptychidae).OD4 originating dorsally at raphe with TEb2 andventrally on Pb3 dorsal surface, inserting on anteriorsurface of Eb4 levator process.Remarks. Left-side OD4 appears to have two al-most continuous origins, and the muscle divides atabout mid-length with the ventral branch attachingseparately to Eb4 ventral to OD4'.OD4' originating on dorsal surface of Pb4 and in-serting on medial edge of Eb4 levator process.M. UP5-Cb4 of left side originating by long ten-don on medial edge of UP5 close to Pb4-Eb4 jointand inserting by long tendon on posterodistalmostcartilaginous surface of Cb4 close to tiny sesamoidbone (not illustrated) on medial side of Eb4-Cb4joint; muscle on right side originating as pair of slen-der tendons, one on medial edge of UP5 close to Pb4-Eb4 joint and the other on ventromedialmost bonyedge of Eb4, tendons uniting just before inserting ason left side.Remarks. The tiny sesamoid bone was not seen oneither side of two cleared and stained specimens, ca.130 mm (USNM 327091).OP dorsally on ventral surface of thick medial armof Eb4, ventrally at and medial to Ad5 insertion onCb5, interrupted by ER at about mid-length, butclearly continuous ventral to ER.Remarks. Other than Gonostoma, the dorsal at-tachment of OP is always on the dorsoposterior sur-face or posteroventral edge of Eb4.Ad 1-3 absent.Ad4 broadly on dorsoposterior edge of Eb4 levatorprocess, narrowing almost to point at tendinous in-sertion on posteromedial end of Cb4.Ad5 on Eb4 near dorsodistal end and Cb5 dorso-distal end; ligament runs along lateral edge of Ad5.RDs appressed medially, insert on posteroventro-medial surface of UP5.SO longitudinal fibers appear to begin at abouthorizontal joining Eb4 uncinate processes and endanteriorly at attachments to medial surfaces of UP5s.SOD a thin strap arising from SO and medial edgeof ER and extending dorsoanteriorly along dorsopos-terior surface of Eb4 from one side to other.Additional remarks. SCL absent. TV4 free fromCb5s. Identity of single UP questionable.AteleopodiformesATELEOPODIDAEAteleopus loppei (Roule), USNM 349721. ca. 350mm TL, Eastern Atlantic.Plate 53
Additional material. ? = Ateleopus sp., USNM359636, ca. 300 mm TL, Caribbean, only partialleft-side dorsal gill arches (information providedfor all muscles for which information is ascertain-able). ? = Ateleopodidae, USNM 361124, VityazCruise station 2560, W. Indian Ocean, small spec-imen, only incomplete left-side dorsal gill arches(information provided below for all muscles forwhich information is ascertainable).
Description.LEI on Ebl posterodorsal surface distally, com-prises two separate elements oriented longitudinallyon each side, right-side with additional element me-dial to paired elements. ? Comprises single elementon each side.LE2 on Eb2 posterodorsal surface distally. ?Same.LE3 slender, on connective tissue between Eb3 andEb4 distally. ? Same.LE4 on Eb4 dorsodistally. ? Same.LP absent. ? Same.LI1 on Pb2 dorsally. ? Same.LI2 on Pb3 bony surface dorsally, fuses dorsallywith LI3 ventral to origin. ? Same.LI3 left side: on cartilaginous posterior end of Pb3laterally, fuses dorsally with LI2 ventral to origin;right side: on bony and cartilaginous posterior end ofPb3 laterally, fuses dorsally with LI2 ventral to ori-gin. ? ? on Pb3 bony surface dorsoposteriorly andanterolateral surface of UP5, fuses dorsally with LI2ventral to origin.Remarks. Johnson (1992:11), who did not examineateleopodids, stated that LI3 inserts on Pb4 in allnon-neotelosts but among neoteleosts, shifts its in-sertion to UP5 in stomiiforms and aulopiforms; LI3and UP5 are absent in ctenosquamates.Olney et al. (1993:154), based on a specimen ofAteleopus japonicus Bleeker, reported that LI2 insertson Pb3 and LI3 inserts on UP5. Ateleopodids lackPb4, and so the insertion of LI3 would be expectedto occur on one or both, Pb3 and UP5. Whether theinsertion of LI3 on some portion of Pb3 in USNM349721 is typical or atypical for that species, doesnot bear on the problematic phylogenetic position ofateleopodids.TD comprises TEb2 and TPb3. TEb2 attaches toEb2 dorsoposterior margin and is posteriorly contin-uous with TPb3, which attaches to Pb3 dorsomedialmargin and is continuous posteriorly with SO. ?Same.OD3 absent. ? Same.OD4 origin on Pb3 dorsally near anteriormost tip,insertion on Eb4 dorsal bony surface just medial toLE4 insertion. ? Same.Remarks. Absence of Pb4 in ateleopodids presentsdifficulties in inferring synapomorphic states for the
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origin of OD4. See synapomorphies (34) (35) in Re-sults section.OP distinct dorsally as fan of muscle inserting onEb4 posteromedially, extending ventrally and fusingwith SO medially, becoming continuous ventrolater-ally with AD5 at posteriormost ventromedial edge ofCb4, where both attach to Cb4 by CT. ER absent.Ad 1-3 absent.Ad4 posteroventrally on Eb4 lateral to OP (not vis-ible in dorsal view), ventrally on dorsodistal cartilag-inous end of Cb4 just medial to Eb4-Cb4 joint.Ad5 dorsolaterally on posteromedialmost end ofCb4, ventrally on posterolateral end of Cb5, mediallyinseparable from OP (see OP above).RD broad, thick, dorosomedian sheet completelyincluded in SO, divides anteriorly at horizontal join-ing Eb4s at mid-length, attaches to UP5 and almostentire medial surface of Pb3.SO longitudinal fibers very sparse, restricted toventralmost area of SO.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl absent.
AulopiformesAULOPIDAEAulopus filamentosus (Bloch), USNM 313689. 167mm. Plate 54
Description.LEI on lateral edge of Ebl uncinate process.LE2 on anterior surface of lateral end of uncinateprocess at and lateral to insertion of TEb2.LE3 on cartilaginous tip of Eb3 uncinate processjust dorsal to OD3 insertion.LE4 origin clusters with other LEs; insertion ondorsal bony surface of Eb4 posterolaterally.LP absent.LI1 on Pb2 dorsally at and medial to base of un-cinate process.LI2 on posterodorsal surface of Pb3 at and lateralto TPb3' attachment and on Pb4 at and lateral toTPb4 attachment anteriorly.LI3 on dorsoanterior surface of UP5. anteriorly atand lateral to mid-point of attachment of TPb4 toPb4.Remarks. Nelson (1967a:286) stated mistakenlythat Aulopus has only two Lis. Johnson (1992) ar-gued persuasively that LI3 in Neoteleostei has shiftedits insertion from Pb4 to UP5. He illustrates his in-terpretation in his fig. 3b of Aulopus.TD comprises TPb2-Pb3a, TEb2, TPb3p, TPb3\TPb4, and TEb4. TPb2-Pb3a anteriorly on dorso-medial surface of Pb2 anterior to LI 1 . continuing ondorsomedial surface of anterior portion of Pb3 ante-
rior to OD3 origin, continuous posteriorly with TEb2
by diagonal muscle strand. TEb2 on most of anteriorsurface of Eb2 uncinate process, ending laterally atmedial edge of LE2 insertion, partially dorsal to OD3origin and anterolateral portion of TPb3p, continuousmid-posteriorly with TPb3p. TPb3p on Pb3 mid-dor-solaterally anterior to LI2 insertion, weakly continu-ous mid-posteriorly with TPb3'. TPb3' on Pb3 pos-terolaterally just medial to LI2 insertion, well-sepa-rated laterally, but broadly continuous mid-posteri-orly with. TPb4. TPb4 on Pb4 posterolaterally,abutting TEb4 mid-posteriorly. TEb4 attaching toEb4 posteromedially dorsal to SO and, on left sideonly, ventral to dorsomedial OP fibers, posteriorlyabutting SO.Remarks. The various divisions of TD are fairlywell differentiated and more numerous than in anyother taxon we examined.OD3 origin on Pb3 ventral to TEb2, insertion ondorsolateral surface of Eb3 uncinate process.OD4 origin on Pb3 ventral to TPb3p, insertion ondorsomedial edge of Eb4 uncinate process.LP absent.OP dorsally on dorsoposterior edge of Eb4 includ-ing uncinate process, joining raphe dorsally withOD4 insertion, ventrally joining ER at ventroposter-ior end of Eb5; not distinguishable from SO or Ad5below ER.Ad 1-3 absent.Ad4 dorsally on dorsoposterior edge of Eb4, ven-trally broadly on Cb4 dorsal surface medial to innerangle formed by Eb4-Cb4 joint.Ad5 dorsally on distal end of Eb5 and slightly ondistal end of Cb4, ventrally on Cb5 posterodistally,dorsomedially joining raphe with OP, medially notseparable from OP.RD not represented externally, comprises, single,broad, thick bundle of longitudinal fibers dorsally inesophagous between outer transverse SO muscle andinner mucosal layer; divides anteriorly at about levelof medial end of Eb3, inserts on medial edge of Pb4and UP5 and along medial edge of posterior half ofPb3.Remarks. Nelson (1967a:286) noted that pairedRDs in Aulopus are absent, and described the inser-tion of this muscle as follows: ". . . fibers anteriorlyattach mostly to the connective tissue of the pharyn-geal roof ..."SO longitudinal fibers questionably present; his-tological verification needed (see remarks followingRD under Saurida).Additional remarks. SCL present, attached anteri-orly to ventromedialmost end of Hb3, with ligamentarising dorsally from mid-posterior edge and attach-ing to ventroanterior surface of Bb3. TV4 free fromCb5s.
66 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONSYNODONTIDAESaurida gracilis (Quoy and Gairmard), USNM140822, 131 mm. Plate 55
Description.LEI on tip of Ebl uncinate process.LE2 on Eb2 just lateral to distal end of uncinateprocess and at and anterior to lateralmost attachmentof TEb2.LE3 on tip of Eb3 uncinate process.LE4 on Eb4 amid OD4 insertion; origin clusteredwith other LEs.LP absent.LI1 broadly on Pb2 anterior process just lateral toTPb2 attachment.LI2 on Pb3 posterolaterally.LI3 on posterolateral surface of UP5.Remarks. See remarks following LI3 in account ofAulopus filamentosus.TD comprises TPb2, TEb2, TEb3, and TEb4.TPb2 on Pb2 medial to LI1 insertion, dorsal to M.Pb2-Eb2 origin, continuous posteriorly by diagonalmuscle strap with TEb2. TEb2 on anterior bony sur-face of Eb2 uncinate process, continuous posteriorlyby diagonal muscle strap with TEb3. TEb3 on dor-soposterior surface of medial end of Eb3 ventral toOD3 insertion and OD4; posteriorly, TEb3 abuts fi-bers of TEb4. TEb4 slender muscle band on mid-dorsoposterior surface of Eb4, attachment ventral todorsal end of OP, posteriorly abuts fibers of SO.M. Pb2-Eb2 slender, longitudinal muscle, anteri-orly on Pb2 ventromedial to TPb2 attachment, pos-teriorly on anterior edge of medial end of Eb2 un-cinate process partially coincident with TEb2 attach-ment.OD3 origin broadly on Pb3 dorsomedially ventralto TPb2 and anterior to OD4 origin, extending pos-teriorly ventral to TEb2 and inserting on anterior andposterior surfaces of Eb3 uncinate process.OD4 origin on Pb3 posteromedial to posterior endof OD3 origin, insertion on anterior and posteriorsurfaces of dorsal crest of Eb4.OP broad, on Eb4 dorsoposterior edge medial todorsal attachment of Ad4 and ventral to TEb4, ven-trolaterally joins?ends at?ER at level of distal endof Eb4, fuses with SO ventromedially.Ad 1-3 absent.Ad4 dorsally on lateral dorsoposterior edge of Eb4,ventrally on Cb4 dorsal surface medial to inner angleformed by Eb4-Cb4 joint.Ad5 dorsally on posterodistal end of Cb4, ventrallyon posterodistal end of Cb5, ventromedially joins ra-phe with TV5, dorsally joins ER with OP.RD not represented external to SO, comprises bi-lateral pair of larger, longitudinal muscle bundlesventrally (RD) and very weakly differentiated small-
er pair dorsally (RD'); RD' fibers divide at posteriorend of Pb3s and each division extends anteriorly, in-serting along medial edge of most of posterior halfof Pb3; RD fibers attach to UP5.Remarks. Large longitudinal muscle bundle pres-ent on each side ventrally in esophagous betweentransverse muscle and mucosal layers (ventral musclebundle absent in Aulopus). There are no longitudinalmuscles in the lateral walls of the esophagous.Additional remarks. SCL absent. TV4 free fromCb5s. UP4 and Eb5 absent.CHLOROPHTHALMIDAEChlorophthalmus agassizi Bonaparte, USNM357798, 121 mm, USNM 159379, 2 specimens,119-125 mm. Plate 56
Description.LEI dorsolateral^ on bony edge of Ebl uncinateprocess just ventral to tip.LE2 on dorsolateralmost surface of distal end ofEb2 uncinate process.LE3 dorsoanteriorly on tip of Eb3 uncinate pro-cess.LE4 on dorsal tip of Eb4 levator process.LP absent.LI1 on bony anterior edge of Pb2 uncinate process.LI2 on dorsoposterior surface of Pb3.LI3 anterior third on Pb4 lateral surface, posteriortwo-thirds on dorsoanterior edge of UP5 at junctionwith Eb4.TD comprises TPb2-Pb3a, TEb2, TPb3p-Pb4, andTEb4. TPb2-Pb3a laterally on medial ends of proxi-mal Pb2 uncinate process, attaching ventromediallyto Pb3 dorsoanterior surface, continuous posteriorlyby diagonal muscle strand with TEb2. TEb2 veryslender, on dorsomedial surface of proximal end ofEb2 uncinate process, posteriorly continuous by di-agonal muscle strand with TPb3p-Pb4. TPb3p-Pb4on dorsomedial surfaces of Pb3 and Pb4, continuousposteriorly with TEb4. TEb4 on dorsomedial end ofEb4, continuous posteriorly with SO.OD3 origin on Pb3 just posteroventral to TEb2 and
at and dorsal to posterior portion of OD4 origin, lat-erally dorsal to OD4 for much of length of OD4;insertion on Eb3 uncinate process just ventral to LE3insertion.OD4 origin on Pb3 anteriorly ventral to TPb2-Pb3a and TEb2 and posteriorly on Pb4 anterodorsalsurface ventral to OD3, laterally ventral to OD3 formuch of length of OD4; insertion broadly on bonyanterior surface of Eb4 levator process.OP one or two straps of muscle attaching dorsallyto Eb4 dorsoposterior edge just medial to dorsal at-tachment of Ad4, and ventrally to Cb5 posterior sur-
NUMBER 1 1 67
face medial to Ad5 attachment, only slightly differ-entiated medially from SO. ER absent.Ad 1-3 absent.Ad4 broadly dorsally on Eb4 dorsoposterior edgelateral to OP attachment, ventrally on Cb4 dorsal sur-face just medial to inner angle formed by Eb4-Cb4joint.Ad5 dorsally on posterodistal ends of Cb4 andEb5, ventrally on posterodistal surface of Cb5.RD origin tendinous, muscle undivided (unpaired),completely ventral to SO transverse muscle layer, ex-tending broadly anteriorly and becoming continuouswith ventral surface muscle fibers of TPb2-Pb3a, at-taching to Pb3, Pb4, and Eb4.SO longitudinal muscle fibers questionably pre-sent; need histological verification.Additional remarks. SCL absent. TV4 free fromanterior ends of Cb5s.
MyctophiformesNEOSCOPELIDAENeoscopelus macrolepidotus Johnson. USNM358034, 127 mm; TCWC 7012.06, ca. 76 mm.Plate 57
Description.Remarks. The description is based on both speci-mens, but the illustration is based on the larger spec-imen because the smaller specimen was deeply dis-sected before it was decided to illustrate the taxon.The muscles in the larger specimen are more robustand folded than in the smaller specimen, and someelements (e.g., ER) and attachments are not readilyvisible without removing folds of overlying muscle,whereas in the smaller specimen the elements aremore obvious.LEI very weak, on mid-dorsoposterior edge ofEbl well lateral to uncinate process.LE2 on raised dorsoposterior edge of Eb2 some-what lateral to mid-length (uncinate process absent).LE3 on anterior edge of Eb3 uncinate process pos-terior to OD3 insertion, extends anteriorly medial toLI2.Remarks. In the smaller specimen, the uncinateprocess of Eb3 and levator process of Eb4 are tightlybound together. In the larger specimen, they are sep-arated as illustrated, but thin, strong tendinous con-nective tissue joins the processes and muscles thatthe tissue passes over.LE4 on Eb4 levator process just proximal to car-tilaginous tip, surrounded anteriorly by OD4 inser-tion, extends anteriorly medial to LI2.LP absent.LI1 on Pb2 medial to uncinate process and some-what ventral toTPb2a, passes medial to LI2.
LI2 on Pb3 dorsolaterally just anterior to joint withmedial end of Eb3.LI3 absent.Remarks. Johnson (1992) argued persuasively thatthe concomitant loss of LI3 and UP5 are a synapo-morphy of Ctenosquamates.TD complex, comprises TPb2a. TPb2p, TEb2,TPb3. and TEb4. TPb2a dorsally on Pb2 betweendorsoanteriorly projecting cartilaginous end and car-tilaginous tip of uncinate process, noticeably separatefrom TPb2p laterally and dorsally, but continuousposteroventrally with it. TPb2p on dorsal bony sur-face of Pb2 just medial to LI1 insertion, continuousposteriorly with TEb2. TEb2 on Eb2 dorsomedially,near articulation with Pb3, continuous posteroven-trally with TPb3. TPb3 extensive longitudinally, liesdorsal to Pb3s, muscle strands extending ventrallyfrom mid-anteroventral surface (ventral to TEb2) at-tach to medial surfaces of Pb3s. joins raphe laterallyon each side with origins of OD3 and OD4. poster-oventrally continuous with TEb4. TEb4 on Eb4 be-tween OD4 insertion and OP dorsal attachment.OD3 origin on Pb3 laterally ventral to TPb2p andTEb2, joins raphe anteriorly with TPb3, continuouswith OD4 origin (fibers appear separate dorsally butmesh ventrally), separating from OD4 posterolater-ally before inserting on Eb3 uncinate process anter-oventral to LE3 insertion.OD4 origin from dorsal raphe with TPb3 and dor-somedial surface of Pb3, insertion on Eb4 levatorprocess.OP dorsally on Eb4 uncinate process medial toAd4 attachment and ventral to OD4 insertion, inter-rupted by ER at mid-length (ER extends tendinouslylaterally and attaches to Cb4 distally together withdorsal end of Ad5), but continues below ER and joinsraphe with anteroventral portion of Ad5 and lateralend of TV5 before attaching to Cb5 medial to raphe.Ad 1-3 absent.Ad4 dorsally on Eb4 posterolateral edge lateral toOP, ventrally on Cb4 medial to inner angle formedby Eb4-Cb4 joint.Ad5 on Cb4 distal end at attachment of ER lateraltendinous extension, joins raphe with Ad4 anteroven-trally and TV5 medially.RD large muscle cluster consisting of dorsal, in-completely separated bilateral pair of bundles with
ventral, somewhat triangular, incompletely separatedbundle (RD') between them, inserting on posteriorsurface of Pb3 and Pb4 and posterior edge of UP4,and extending anteriorly between and attaching tomedial surfaces of Pb3s.SOD ventral to TEb4.SO longitudinal muscles restricted to dorsal andventral areas of SO.Additional remarks. SCL absent. TV4 free from
68 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Cb5s. UP5 and Eb5 absent. Eb4 uncinate process ab-
sent. MYCTOPHIDAELampanyctus macdonaldi (Goode and Bean), USNM303167, 118 mm.Plate 58A, B
Description.LEI absent, possibly represented by tough CT in-serting on Ebl uncinate process and attaching to skull(see remarks following LEI in Diaphus).LE2 on bony Eb2 process.LE3 on cartilaginous tip of Eb3 uncinate process.LE4 origin tendinous; insertion on cartilaginous tipof Eb4 levator process coincident with OD4 inser-tion. Originates with tendinous origins of other LEs.LP absent.LI la on Pb2 anterior to uncinate process (see re-marks under LIlp).LIlp on ventromedial surface of Pb2 uncinate pro-cess, questionably impinging on ventroposteriormostedge of LI la insertion.Remarks. LI la and LIlp are more accurately de-scribed, perhaps, as separated by the posterolateralportion of TPb2a, which lies dorsal to the insertionof LIlp, rather than by the ascending process (= ouruncinate process) of Pb2, as stated by Stiassny (1996:407). Stiassny possibly based her remark on John-son's (1992:fig. 7B) illustration of the condition inthe myctophid Diaphus mollis (but see remarks underLI la in our description of Diaphus mollis).LI2 on posterolateralmost dorsal surface of Pb3.LI3 absent..TD comprises TPb2a, TPb2p. TEb2. TPb3, andTEb4. TPb2a on dorsal bony surface of Pb2 anteriorto Pb2 uncinate process; muscle narrowly continuousposteromedianly with TPb2p. TPb2p on posteriorarm of Pb2 dorsally, continuous posteriorly by di-agonal muscle strand with TEb2. TEb2 on dorsal sur-face of medialmost end of Eb2 near articulation withPb3. TPb3, separated by large gap from TEb2, whichcontinues onto TEb4; muscle attached to Pb3 later-ally ventral to OD3 and OD4, continuous postero-medianly with TEb4. TEb4 on mid-posterior surfaceof Eb4 ventral to OD4 insertion and medial to OPdorsal attachment, continuous mid-posteriorly withSOD.Remarks. A median longitudinal raphe extendsfrom the posterior half of TPb2p to posterior marginof TEb2, and another extends from TPb3 to dorso-posterior margin of SOD.OD3 lies dorsal to mid-section of OD4; origin onPb3 dorsomedial edge dorsal to, and coincident with,origin of OD4 mid-section; insertion dorsoanteriorlyon Eb3 uncinate process ventral to LE3 insertion.OD4 origin extensive, beginning on Pb3 dorso-
medial edge dorsal to TPb3 and continuing posteri-orly ventral to OD3 origin, then dorsally to end ofPb3; insertion massive, on anterior surface of Eb4levator process.OP dorsally on Eb4 posterior surface medial toAd4 dorsal attachment, interrupted by ER ventrally,but continuing and attaching to Cb5 just anterior tocombined attachment of Ad5 and TV5 to Cb5.Adl-3 absent.Ad4 dorsally on posterolateral surface of Eb4; ven-trally, narrowly on Cb4 just medial to inner angleformed by Eb4-Cb4 joint.Ad5 broadly on Eb4 posterolaterally and Cb5 dor-sodistally, joining raphe with TV5 on Cb5.RD on each side comprising large RD ventrallyand small RD'. RD inserts completely separatelyfrom RD', attaching to posterior end of Pb4 and UP4and medial end of Eb4; RD' joins longitudinal SOmuscle layer at dorsoposterior end of Pb3 as layerdivides bilaterally (similar to Diaphus, Plate 58C).SOD present.SO longitudinal muscle layer restricted to dorsaland ventral areas; dorsal section divides bilaterally atposterior end of Pb3s, each division extending ante-
riorly between Pb3s and attaching to anteromedialsurface of its respective Pb3.Additional remarks. SCL absent. TV4 free fromCb5s. UP5 and Eb5 absent. Eb4 uncinate process ab-
sent.Jollie (1954) described and illustrated the gill-archmuscles of Lampanyctus leucopsarus (= Stenobra-chias leucopsarus Eigenmann and Eigenmann),which are very similar to those of L. macdonaldi.
Diaphus mollis Taning, USNM 300877 (Gulf ofMexico), 2 specimens, 50.5-54.4 mm; USNM274201, 54.2 mm (E. of New Zealand).Plate 58C
Description.Remarks. We found no differences worth notingamong the specimens of D. mollis from the two geo-graphic areas.LEI very fine, on dorsal tip of bony process justlateral to tip of Ebl uncinate process, where it isjoined by insertion of medial edge of LE2.Remarks. Stiassny (1996:412) stated that reductionof LEI to a thin slip (Neoscopelidae) or ligament(Myctophidae) is a synapomorphy of myctophiformfishes. We agree that a reduced LEI may be syna-pomorphic for myctophiforms, but we find that LEIin myctophids also may be present as a thin slip ofmuscle.LE2 on posterodorsal edge of Eb2 where bonewidens; muscle extends anteriorly and attaches to
NUMBER 1
1
69
bony process on Ebl lateral to uncinate process,thence continuing to origin on cranium.LE2' very fine, becoming even finer and tendinousbefore inserting among LE2 fibers (absent in Lam-panyctus).LE3 on cartilaginous tip of Eb3 uncinate process.LE4 on cartilaginous tip of Eb4 levator processcoincident with Od4 insertion.LI1 with single insertion equivalent to LI la ofLampanyctus macdonaldi.Remarks. Johnson (1992:fig. IB) illustrated LI1 ina specimen putatively identified as Diaphus mollis(AMNH 29449). In contrast to our finding for thespecies, his figure shows LI1 as having anterior andposterior sections with insertions equivalent to LI laand LIlp of Lampanyctus macdonaldi (q.v.). Thespecimen Johnson illustrated is actually identifiableas D. dumerilii (USNM 301132). AMNH 29449,however, is identifiable as D. mollis (from off Ber-muda). M. Stiassny examined a specimen from thesame AMNH lot and informed us (E-mail, 15 May2000) that LI1 has a single insertion. Aside from D.dumerilii and L. macdonaldi, we find LI1 also hastwo insertions in D. lucidus (USNM 269439), D. re-gard (USNM 269381), Hygophum hygomi (USNM206616), and Protomyctophum normani (USNM206631). Stiassny (1996:407) interpreted the subdi-vision of LI1 as a synapomorphy of the Myctophidae.The character state for LI1 in D. mollis is probablyautapomorphic.LI2 on posterolateralmost dorsal surface of Pb3.LI3 absent.TD comprises TPb2a, TPb2p. TEb2, TPb3, andTEb4. TPb2a on dorsal bony surface of Pb2 anteriorto Pb2 uncinate process, dorsal to anteriormost edgeof LI la insertion, and narrowly continuous postero-medianly with TPb2p. TPb2p on posterior arm ofPb2 dorsally, continuous posteromedially with TEb2.TEb2 on dorsal surface of medialmost end of Eb2near articulation with Pb3. TPb3, separated by largegap from TEb2, on Pb3 ventrolaterally, continuousposteromedianly with TEb4. TEb4 on mid-posteriorsurface of Eb4 ventral to OD4 insertion and medialto OP, with which in joins a raphe laterally, and iscontinuous mid-posteriorly with SOD.Remarks. A median longitudinal raphe extendsfrom TPb2a to posterior margin of TEb2, and anotherextends from TPb3 to dorsoposterior margin of SOD.TPb2a is much narrower relatively than it is in Lam-panyctus; it lacks the anterior portion without the me-dian raphe.OD3 lies dorsal to mid-section of OD4; origin onPb3 dorsomedial edge dorsal to, and coincident with,origin of OD4 mid-section; insertion dorsoanteriorlyon Eb3 uncinate process ventral to LE3 insertion.OD4 origin extensive, beginning on Pb3 dorso-medial edge, passing dorsal to TPb3 and continuing
posteriorly ventral to OD3 origin, then dorsally toend of Pb3; insertion massive, on anterior surface ofEb4 levator process; posteroventrally joining raphewith OP.OP a narrow strap; dorsally, narrowly on dorso-posterior Eb4 bony surface joining raphe laterallywith OD4 insertion and medially with TEb4 attach-ment; ventrally on distalmost tip of Cb5, not inter-rupted by ER, which appears to be absent.Ad 1-3 absent.Ad4 dorsally on posterolateral surface of Eb4, ven-trally, narrowly on Cb4 just medial to inner angleformed by Eb4-Cb4 joint.Ad5 narrowly on Eb4 posterodistally and Cb5 an-terodistally.RD comprising large RD ventrally and small RD'dorsally on each side. RD inserts completely sepa-rately from RD', attaching to posterior end of Pb4and UP4; RD' joins longitudinal SO muscle layer atdorsoposterior end of Pb3 as layer divides bilaterally.SOD present.SO longitudinal muscle fibers appear to be restrict-ed dorsally and ventrally in SO and originate at abouthorizontal between distal ends of Eb4s and extendanteriorly about to horizontal between distal ends ofEb3s.Additional remarks. SCL absent. TV4 free fromCb5s. See also additional remarks under Lampanyc-tus macdonaldi. UP5 and Eb5 absent. Eb4 uncinateprocess absent.
Results?Pre-AcanthomorphaMuch of the raw data on the muscles is presentedin Tables 1?6. The following discussion summarizesinformation for most of the muscles and assesses howthe information bears on the pre-acanthomorph clad-ogram (Fig. 4). Muscle synapomorphies are italicizedand numbered in parentheses, and summarized in Ta-ble 7. They are interpreted as such by parsimonybased on how the character states are distributedamong the taxa on the cladogram (for an example,see synapomorphy 9 below). We noted many cyprin-iform synapomorphies, but we include only those thatoccur homoplastically in other taxa (the two cyprinidgenera we examined are closely related). We usuallyignore autapomorphies of terminal taxa, but autapo-morphies that might be interpreted as synapomor-phies based on results of a total evidence analysis(not performed by us) are discussed. In some caseswe have noted skeletal synapomorphies; these are notnumbered nor indicated by italics.(1) Presence of Ads is a synapomorphy of Actin-opterygii.(2) Attachment ofpart of SO to a dorsal gill-archskeletal element is a synapomorphy of Actinopteri.
70 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Polypterus is the only taxon in which SO does notattach to a dorsal gill-arch element.(3) Insertion of LEI at dorsomedlalmost end ofEbl is synapomorphic for Osteoglossomorpha (nec-essarily ignoring the two osteoglossomorph taxa thatlack LEI or in which it is unrecognizable, i.e., fusedindistinguishably with another muscle). Among otherpre-acanthomorphs, the insertion point of LEI isquite variable, but with the exception of Elops, isalways lateral to the medialmost end of Ebl. LEI israrely absent, and only autapomorphically.(4) Insertion of LE2 at dorsomedialmost end ofEb2 is synapomorphic for Osteoglossomorpha. Pan-todon, which is well nested among the osteoglosso-morph clades on the basis of other evidence, is ex-ceptional. Among the other pre-acanthomorphs, theinsertion point of LE2 is quite variable, but with theexception of Elops, is always lateral to the medial-most end of Eb2. LE2 is rarely absent, and only au-tapomorphically, and the state is ignored in those taxathat lack it or in which it is unrecognizable, i.e., fusedindistinguishably with another muscle.(5) Doubling of LE3 is a synapomorphy of Os-meroidea. A doubling of LE3 (LE3, LE3') also oc-curs in some Clupeomorpha. LE3 is rarely absent,and only autapomorphically, except among Osteo-glossomorpha, in which three of the ten genera ex-amined lack the muscle or it is unrecognizable, i.e.,indistinguishably fused with another muscle. See also(6) for Salangidae.(6) Doubling ofLE4 (LE4, LE4') is a synapomor-phy of Osmeroidea. Too late for detailed inclusion,we examined the salangid Protosalanx chinensis (Os-beck), USNM 85840, and note that it also has LE3and LE4 doubled. This corroborates Johnson and Pat-terson's (1996:307) inclusion of "salangids" withOsmeridae.(7) LP present is a synapomorphy of Clupeoidei;see remarks following (8).(8) LP present is a synapomorphy of Characifor-mes. LP has been reported for several diverse groupsof pre-acanthomorph fishes, but we reserve the namefor a muscle that inserts on Eb4, with or slightly sep-arate from LE4, and originates "closely apposed tothe surface of the epaxial body muscles [of clu-peoids]" (Greenwood and Lauder, 1981:215), or onvarious cranial bones in all other groups that have it.Among pre-acanthomorphs, LP is restricted other-wise to Chanos, also a member of Otocephala. Al-though absent in Denticeps, currently recognized asthe sister group of all other clupeomorphs, the pres-ence of LP will probably be found to be synapo-morphic for Otocephala. LP next appears in Lampri-diformes, first clade of Acanthomorpha. Levator pos-terior has been applied to a variety of morphologi-cally similar but non-homologous muscles as thefollowing discussion amply demonstrates.
Greenwood and Lauder (1981:228) stated that allof the otophysan taxa they examined, which includedrepresentatives of all four major groups (Fig. 4), haveLP. We question the presence of LP in otophysansother than characiforms. The muscle Greenwood andLauder identify as LP in cypriniforms is based onWinterbottom's interpretation (1974:254-256), andcorresponds to our LCb5 (see Winterbottom's fig.22a), which inserts on Cb5; that of siluriforms cor-responds to our LI4 (see Winterbottom's fig. 21),which inserts on UP4. We are uncertain which gym-notiform muscle Greenwood and Lauder consideredto be LP unless it is the muscle we identify as LE4(inserts on Eb4), in which case they would considerLE4 as missing (see also Remarks following LI4 indescription of Diplomystes). If future study shouldindicate that LE4 is absent in gymnotiforms, then ab-sence of LE4 would appear to be a synapomorphy ofsiluriforms + gymnotiforms.(9) LI1 appears first in Ginglymodi and is a syn-apomorphy of Neopterygii. From its first appearance,LI1 is the only levator that is present in all actinop-terygian taxa we examined.Lauder and Wainwright (1992:457) indicate thatLis are plesiomorphic for Chondrichthyes + Actin-opterygii. We are unable to find the source on whichthey based their indication. Lis in Actinopterygii firstappear in Neopterygii, are synapomorphic for thatgroup, and cannot be plesiomorphic for Chondrich-thyes + Actinopterygii: chondrichthyans have no le-vators originating on the cranium.SPbl first appears in Chondrostei and is a syna-pomorphy of Actinopteri. LI1 is attached to SPblwhen LI1 first appears in Ginglymodi. Halecomorphaand Osteoglossomorpha lack SPbl, so loss of SPbl(and hence attachment of LI1 to it) is a synapomor-phy of Halecostomi. SPbl reappears in Elopomorphaand again LI1 is attached to it (condition present inboth elopiform families and both albuliform families,but absent in all Anguillomorpha). Reappearance ofSPbl is a synapomorphy of Elopomorpha, but wheth-er attachment of LI1 to SPbl is also a synapomorphyis equivocal, because we do not know whether LI1was attached to SPbl in the ancestor of those formsthat lack SPbl. The situation repeats. Loss of SPbl(and attachment of LI1 to it) is a synapomorphy ofClupeocephala, but SPbl, with LI attaching to it, re-appears in, and is a synapomorphy of. the clade Pla-tytroctidae + Alepocephalidae. The apparent conclu-sion to be drawn is that when both SPbl and LI1 arepresent, LI1 attaches to SPbl.(10) Division of LI1 (our Llla, Lllb) is a syna-pomorphy of Myctophidae. First reported by Stiassny(1996:407), our data corroborate her findings. Pres-ence of only LI 1 a in the myctophid D. mollis appearsto be autapomorphic (see also remarks under LI1 inaccount of D. mollis).
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(11) LI2 appears first in Neopterygii and is a syn-apomorphy of that clade. Aside from some special-ized osteoglossiforms, LI2 is present in all but three,distantly related, pre-acanthomorph taxa that we ex-amined.(12) Insertion of LI2 to include UP4 is a syna-pomorphy of Anguilliformes. LI2 inserts primitivelyon Pb3. and as a specialization may insert on otherskeletal elements.(13) LI3 is probably a synapomorphy ofTeleostei.LI3 appears first in Teleostei together with the firstappearance of Pb4, on which it inserts primitively,and is probably a synapomoiphy of that clade (but ifnot that clade, then of Osteoglossiformes and, inde-pendently, Elopocephala). The problem in recogniz-ing LI3 unequivocally as a synapomorphy of Teleos-tei is caused by the state of the Lis in Hiodon, whichis the sister group of all other Osteoglossomorpha (=Osteoglossiformes), which, in turn, is the sister groupof Elopocephala (= all other Teleostei). Hiodon hasonly two Lis, LI1 and a second, relatively large LI,which may simply be LI2 or, conceivably, a fusedLI2 and LI3. There are two possible scenarios: 1 ) thesecond LI represents only LI2, and LI3 has evolvedindependently in Osteoglossiformes and Elopocepha-la; 2) the second LI represents fused LI2 and LI3 oronly LI2, LI3 having been lost; the presence of LI3would, thus, represent a teleostean synapomorphy.We prefer the second scenario for reasons that follow.Osteoglossiformes, sister group of Hiodon, com-prises two clades: Notopteroidei and Osteoglossoidei.Three of the four genera of notopteroids are highlyspecialized with many muscle fusions or losses, andLI2 and LI3 (if the latter was present) are either lostor indistinguishably fused with other muscles. Thefourth genus, Notopterus and sister-group of the otherthree genera, has both LI2 and LI3. The character
state for LI3 at the base of Notopteroidei is "LI3present or ?". Four of the five genera of Osteoglos-soidei (including all three genera in the two basal-most clades) have LI2 and LI3 (the exception hasonly LI2). The character state for LI3 at the base ofOsteoglossoidei is "LI3 present." Combining thesetwo states, one concludes parsimoniously that thecharacter state of LI3 for Osteoglossiformes is "LI3present." The state for Hiodontiformes is, at its mostconservative, "LI3 absent"; therefore, the characterstate for Osteoglossomorpha would be "LI3 presentor absent." Given that "LI3 present" is the plesio-morphic state for Elopocephala, combining the statesfor the two teleostean clades results parsimoniouslyin "LI3 present" as the (synapomorphic) characterstate for Teleostei.(13a) Insertion of LI3 to include Pb3 is a syna-pomorphy of Osteoglossoidei. Among osteoglosso-morphs, LI3, when present, inserts variously on Pb3and Pb4, Pb4, or Pb3 and UP5. Pb4 is plesiomorphic.
therefore the insertion to include Pb3 is specialized.Insertion on UP5 is autapomorphic for Pantodon.(14) Absence of LI3 is a synapomorphy of An-guilliformes.(15) Shift of the insertion of LI3 from Pb4 to UPSis a synapomorphy of Neoteleostei. See remarks fol-lowing (16).
( 1 6) Loss of LI3 and UP5 is a synapomorphy ofCtenosquamata. Johnson (1992) first hypothesizedthese two synapomorphies (15 and 16) and our find-ings corroborate his hypotheses.TD is absent in Polypterus, possibly a result of lossor great reduction of Eb4. As such, it is not possibleto decide if presence of TEb4 is a synapomorphy ofChondrostei or Actinopterygii; we assume the former(see 18).(17) Absence of attachment ofTD to Eb4 is a syn-apomorphy of Albuliformes.(18) Presence of TEb4 is a synapomorphy ofChondrostei.(19) Presence of'TEb4 is a synapomorphy of Clu-peomorpha.(20) Presence of TEb4 is a synapomorphy of Eu-rypterygii.(21) Attachment of TD to include Pb2 is a syna-pomorphy of Gymnotiformes + Siluriformes.(22) Attachment of TD to include Pb2 is a syna-pomorphy of Eurypterygii.(23) Presence of TPb2a and TPb2b is a synapo-morphy of Myctophiformes.(24) Attachment of TD to Pb3 first occurs inNeopterygii, and is a synapomorphy of Neopterygii.(25) Attachment of TD to Pb4 first occurs in Te-leostei (where Pb4 first appears) and is a synapo-morphy of Teleostei.(26) Attachment of TD to Eb2 is a synapomorphyof Osteoglossoidei (Table 4).(27) Attachment of TD to Eb2 is a synapomorphyof the unnamed clade: (Characiformes (Siluriformes,Gymnotiformes). Attachment of TD to Eb2 is ple-siomorphic for Characiformes and present or absentin its sister group, Siluriformes (present) + Gymno-tiformes (absent).(28) Attachment of TD to Eb2 is a synapomorphyof Neoteleostei.(29) Presence of'TEb2 is a synapomorphy ofNeo-teleostei.(30) Presence of TEb2 is a synapomorphy of Os-teoglossidae.(31) OD4 first appears in Neopterygii, in which itattaches to Pb3, and is a synapomorphy of that clade.After its first appearance, some component of OD4(OD4' or OD4 as represented in the fused OD3-4)is rarely absent. See also (32-35).(32) OD3 first appears in Teleostei and is a syn-apomorphy of that clade. An OD3 component (OD3or OD3?4) is rarely absent.
72 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(33) Except for the osteoglossomorph Notopterus,where it occurs homoplasiously, some component ofOD4 (OD4, OD4 ', OD3^f) first attaches to Pb4 inElopocephala and is a synapomorphy of that clade(Table 4).(34) Loss of attachment of an OD4 component(OD4, OD4', or OD3^t) to Pb4 is a synapomorphyof Neognathi (Table 4). Absence of Pb4 in ateleo-podids makes it impossible to polarize the group withregard to the origin of OD4. However, the origin ofOD4 in ateleopodids is well anterior on Pb3 andwould appear to indicate that it is highly unlikely thatPb4 would have participated in the origin; we assumethis in our analysis of the synapomorphic states for(34) and (35).(35) Attachment of an OD4 component (OD4,OD4', or OD3-4) to Pb4 re-occurs in Stomiiform.esand is a synapomorphy of that clade (Table 4). Seecomments in (34).(36) A vertically oriented OD4 is a synapomorphyof Albuliformes.(37) OP first appears in Halecomorpha and isprobably a synapomorphy of Halecostomi.(38) Absence of OP is a synapomorphy of Gono-rynchiformes. See comments in (40).(39) Absence of OP is a synapomorphy of Cyprin-iformes. See comments in (34).(40) Absence of OP is a synapomorphy of Platy-trochtidae + Alepocephalidae (two representatives ofAlepocephaloidea). OP frequently appears to befused to SO and/or Ad5 and its presence is oftenquestionable.(41) ER first appears in Teleostei and is a syna-pomorphy of that clade. ER is often absent, and isnever present when OP is unquestionably absent.(42a) Presence of RecD4 is probably a synapo-morphy of Chondrostei. RecDs are relatively uncom-mon among pre-acanthomorphs. RecD4 is absent inPolypterus, conceivably as a result of the loss orgreat reduction of Eb4. As such, it is not possible todecide with certainty if presence of RecD4 beginningwith Chondrostei is a synapomorphy of Chondrostei.(42) Presence ofRecD4 is a synapomorphy ofOs-teoglossomorpha.(43) Presence of RecD4 is a synapomorphy ofCy-priniformes.(44) RecD2 first appears in Osteoglossiformesand is a synapomorphy of that clade.(45) Presence ofRecD2 is a synapomorphy ofCy-priniformes.(46) Presence ofRecDl is a synapomorphy ofAn-guilliformes.Bilaterally paired Pb muscles. The muscles of thisgroup (Table 5) have their origins on Pbs and/or UPsand their insertions on Ebs and/or Cbs. These pairedPb muscles are not homologous with TD componentmuscles, which, in pre-acanthomorphs, do not be-
come divided bilaterally by losing their median por-tions, as they may in acanthomorphs. Furthermore,the paired Pb muscles are usually aligned longitudi-nally and, within a species, often have origins on thesame Pbs and UPs as the TD muscles. Paired Pbmuscles, which first appear in the Osteoglossomor-pha, tend to be common only in Osteoglossomorphaand Anguillomorpha.Concerning specific bilaterally paired Pb muscles:(47) Presence of M. Pb4-Eb2 is a synapomorphyof Osteoglossoidei.(48) Presence of M. UP5-Cb4 is a synapomorphyof Albuliformes.(49) Presence M. UP5-Cb4-Eb5 is a synapomor-phy of Characiformes.(49a?) Ad 1-3 are present in only three unrelatedgroups of pre-acanthomorphs: Polyodontidae, Nota-canthidae, Cyprinidae. However, Edgeworth (1935:131) reported that Ad 1-4 are present in Acipensersturio (not examined by us), but not A. ruthenus, A.fulvescens, and Scaphirhynchus. It is possible, there-fore, that Ad1?3 are synapomorphic for Chondrostei.(50) Attachment ofAd5 to Eb5 (when Eb5 is pre-sent either as an autogenous element or when puta-tively fused with Eb4 or Cb4) is a synapomorphy ofTeleostei. This state is by far the most commonamong pre-acanthomorphs up to Neoteleostei. Ad5 isone of the most consistently present muscles and isalways attached at its ventralmost or posteriormostend to Cb5. Among the taxa we examined it is un-equivocally absent only in Polypterus and the eel,Synaphobranchus, both of which lack Cb5. Only theattachments to other skeletal elements will be dis-cussed further.The primitive state for Ad5 in Actinopterygii isconfused by the absence of the fifth arch in Polyp-terus, but it appears that the state is probably that ofAd5 attaching Cb5 to Cb4 (Table 6; Acipenser,Atractosteus, Amia). The primitive state appears au-tapomorphically above Halecomorpha in: Elopifor-mes, Anguilloida, Diplomystes, Lepidogalaxias, Eso-cidae, Ateleopus, Aulopiformes, and Neoscopelus.If attachment of Ad5 to Eb5 or to Eb4* (fusedEb5-Eb4) represents the same state (attachment toEb5), then:(51) Attachment ofAd5 to Eb4 (in the absence ofEb5 or Eb4*) is a synapomorphy of Stomiiformes.The state occurs otherwise, independently, only inMyctophidae, Lepidogalaxias, and in Polyodon.(52) Attachment of Ad5 to Eb4, when Eb5 is au-togenous, is a synapomorphy of Cypriniformes; it oc-curs otherwise, independently, only in Searsia.(53) Presence of RD is a synapomorphy of Neo-teleostei.Wiley (1976:31) observed that RDs in amiids andlepisosteids are derived from the outer circular mus-cle layer of SO, in contrast to their derivation from
NUMBER 11 73
the inner longitudinal layer of SO in other fishes.Thus, he inferred that the RDs of amiids and lepi-sosteids are not homologous with RDs in other fishes,but can be interpreted as homologous in lepisosteidsand amiids in having similar derivations. In opposi-tion to that inference, Wiley (p. 32) also describedseveral differences between the attachments of RDsin lepisosteids and amiids, and inferred that the stateof the RDs in each of these two groups is autapo-morphic, and thus their RDs are not homologous.Plotting the distribution of RD on the cladogram(Fig. 4), disregarding the differing character states ofRD in lepsosteids and amiids, indicates that somekind of RD is a synapomorphy of Neopterygii (butRDs lost is a synapomorphy in the next branch, Te-leostei); thus, Wiley was justified in his first infer-ence. Lacking knowledge of the particular character-istics of RD in the hypothetical neopterygian ances-tor, there are three possibilities: RD resembled thatof either extant lepisosteids or that of extant amiidsor differed from both. His second inference, there-fore, may be correct, but is unwarranted. The lack ofhomology at a higher phylogenetic level does notnecessarily preclude homology at a lower level.Another possibility is that the presence of RD maybe a synapomorphy of Ginglymodi + Halecomorpha,which if corroborated by overall parsimony, will re-quire rearrangement of these two clades in relationto Teleostei. (See also summary following this sec-tion.)RD exhibits three states in pre-acanthomorphs weexamined: RD completely dorsal to SO (SOD ab-sent): Amia, Atractosteus, and Lepidogalaxias (allpre-Neoteleostei); RD completely ventral to SO(SOD absent): Ateleopodiformes and Aulopiformes(both Neoteleostei); RD and SOD present: Pantodon(pre-Neoteleostei) and Stomiiformes and Myctophi-formes (Neoteleostei).Three aulopiform taxa {Gigantura chuni Brauer,USNM 221034; Omosudis lowei Gunther, USNM206792; Parasudis truculentus (Goode and Bean),USNM 159095), in addition to those listed in Fig. 4,were examined. All agree with those listed in Fig. 4in having RD completely ventral to SO. Although avery elongate RD was used to relate Gigantura toaulopiforms (i.e., synodontids, Rosen, 1983:440-441), the importance of RD being ventral to SO inpre-acanthomorphs has gone unnoticed, and offersadditional support for a relationship of Gigantura toaulopiforms, but not specifically to synodontids.Based on the intermusculars, Johnson and Patterson(1995:31) placed Gigantura with alepisauroids with-in aulopiforms, and Baldwin and Johnson (1996) pre-sented evidence that Gigantura and Bathysaurusform a sister group, which is the sister group of ale-pisauroids.Presence of a ventral RD in Ateleopodiformes,
which forms a polytomy with Stomiiformes and Eu-rypterygii might be evidence to relate Ateleopodi-formes with eurypterygian Aulopiformes. It is notpossible, however, to polarize the ventral and dorsalstates of RD. One character of RD, however, a singlebroad band of muscle that only divides at its attach-ment to the dorsal gill-arch skeleton, is unique toateleopodids and some aulopiforms. At least Aulopus,which Baldwin and Johnson (1996:359) included inthe basalmost aulopiform clade. and Chlorophthal-mus, which they included in one of the next two au-lopiform clades, share this state of RD with Ateleo-pus. Based on this evidence it is possible that Ate-leopodiformes and Aulopiformes form a sister group,possibly resolving the Neotelostean trichotomy. Suchan arrangement would require addressing conflictsprovided by synapomorphies (20 and 22), which sup-port an aulopiform and ctenosquamate clade.The presence of SOD basally in Acanthomorphaindicates that this state is plesiomorphic for Ctenos-quamata. An aulopiform-ateleopodiform resolutionof the neoteleostean trichotomy (Fig. 4), in which thenew clade is the sister group of Ctenosquamata,would indicate that presence of SOD is a synapo-morphy of Neoteleostei and its absence a synapo-morphy of Aulopiformes + Ateleopodiformes.(54) Longitudinal SO muscle layer first appears inHalecostomi and is a synapomorphy of that clade.We were unable to find a study on the compositionof the SO muscle fibers in fishes and doubt that oneexists. Indeed, the subject is rarely mentioned. Nev-ertheless, after discussion with several colleagues, webecame aware that there is a generally erroneous con-cept that SO in fishes comprises an outer transversemuscle layer and an inner longitudinal muscle layer(e.g., Wiley, 1976:30-32). We find, however, that al-though an outer transverse layer is always present, alongitudinal layer is not. In some taxa, we were un-able to determine on gross examination if a longitu-dinal muscle layer was present. Although we usuallydo not describe them, there are different distributionpatterns of the longitudinal muscle fibers around theesophagus (e.g., the fibers may be concentrated dor-sally and ventrally and absent laterally or they maybe distributed more-or-less evenly around the esoph-agus). We suggest that a detailed histological studyof the SO muscle layers is in order and would prob-ably reveal important phylogenetic information.(55) Presence of TV4 is a synapomorphy of Hal-ecostomi, and, within the pre-acanthomorph portionof that clade, TV4 is absent only in the galaxioidRetropinna. Note: attachment of TV4 to Hb4 is anautapomorphy of Halecomorpha.(55a) Attachment of TV4 to Cb4 is a synapomor-phy of Teleostei. Note: Absence of Hb4 is also a syn-apomorphy of Teleostei.(56) Attachment ofTV4 to Cb5 is a synapomorphy
74 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of Cypriniformes. Among pre-acanthomorphs, TV4 isprimitively unattached to Cb5s. TV4 is attached toCb5s in only three other pre-acanthomorph taxa, ineach of which it is autapomorphic.(57) SCL first appears unquestionably in Albuli-formes (= Albulidae) and is a synapomorphy of thatclade. Stiassny (1992:269) reported that the "semi-circular ligament system ... is an innovation of theacanthomorph fishes," by which she implied the firstappearance of SCL (and the way in which RecV4and ObV3 attach to it). She further stated (p. 270)that SCL is absent in pre-acanthomorphs. Stiassny iscorrect in indicating that SCL is primitively absentin pre-acanthomorphs. but it occurs in various pre-acanthomorph taxa. SCL is lacking in the most ad-vanced pre-acanthomorphs, myctophiforms and mostaulopiforms, but is present in the basalmost group ofacanthomorphs, Lampridiformes. Only in the sensethat its presence is homoplastic can it be consideredto be an acanthomorph innovation.(58?) Presence of SCL is a synapomorphy of Elo-pomorpha, if the condition of SCL in Elopiformes (seedescription in additional remarks in Elopidae) can beconsidered to indicate its presence. The differences incharacter states between Elopiformes and Albulifor-mes leaves unresolved which state is plesiomorphic.(59?) If SCL is considered to be absent in Elopi-formes, the condition described in additional remarksin Elopidae is a synapomorphy of Elopiformes.(60) SCL is a synapomorphy of the Esociformes.Summary (Table 7). The dorsal gill-arch muscu-lature (and TV4 and SCL) provides support formonophyly of Actinopterygii and several of its cur-rently recognized pre-acanthomorph clades. In gen-
eral, support exists mainly for clades that are alreadywell supported in the literature.The dorsal gill-arch muscles provide conflicting ev-idence regarding the interrelationships of the Gingly-modi, Halecomorpha, and Teleostei. The monophylyof the Halecostomi (Halecomorpha + Teleostei) issupported by three synapomorphies: (37), presence ofOP; (54), presence of SO longitudinal muscle layer;(55), presence of TV4. Alternatively, three synapo-morphies conflict with halecostome monophyly and,instead, support the monophyly of the Halecomorpha+ Ginglymodi: presence of RD, absence of attachmentof TD to Eb4, and absence of RecD4. Thus, based onour muscle evidence, each hypothesis is equally par-simonious. The first is congruent with the generallyaccepted hypothesis expressed in our cladogram (Fig.4). The second is congruent with the molecular evi-dence presented in Gardiner et al. (1996). We suggestthat the interrelationships of Ginglymodi and Hale-comorpha are worthy of additional study.Hilton (2003) hypothesized a different set of in-terrelationships for Osteoglossoidei than the one inour Fig. 4. His classification exchanges the position
of Osteoglossidae (his Osteoglossinae) with that ofArapaimidae (his Heterotinae). His classification dif-fers from ours only in aligning Pantodon as sistergroup of Osteoglossidae (his Osteoglossinae) insteadof Arapaimidae (his Heterotinae). Given Hilton's ar-rangement, the major change would be that synapo-morphy 13a would no longer be a synapomorphy ofOsteoglossoidei, but would, instead, become a syn-apomorphy of Osteoglossidae. The other osteoglos-somorph synapomorphies would remain the same.Although not supporting monophyly of the two oto-cephalan clades, Ostariophysi and Clupeomorpha, twocharacters, the gongyloid cartilage (Di Dario, 2002)and LP are suggestive of close relationship of theclades, i.e., supportive of Otocephala. Among pre-acanthomorphs, the gongyloid cartilage is present onlyin Chanos (Ostariophysi) and most of the genera ofPristigasteroidea and Engrauloidea, (Clupeomorpha;see Additional remarks under Chanos and Cetengrau-lis). Similarly, among pre-acanthomorphs, LP is pre-sent only in characiform genera (Ostariophysi), Chan-os (and possibly Gonorynchus, in which it was re-ported present by Greenwood and Lauder, 1981:228,although we did not find it), and all Clupeomorpha,except Coilia and Denticeps. It is also possible thatLCb5A in cyprinids represents a modified LP.There is evidence (not listed in Table 7) based onthe state of RD and SO that Aulopiformes and Ate-leopodiformes form a clade, but other evidence mayconflict. Acanthomorpha
Additional material of acanthomorph taxa notmentioned in descriptive accounts (used for supple-mentary data, e.g., Table 8).Cleared and stained. Acropomatidae: Acropoma
sp., USNM 287444 (2); Apogonops anomalus,USNM 287447; Doederleinia berycoides (Hilgen-dorf), USNM 290474. Ambassidae: Ambassis sp.,USNM 218805 (4 specimens); A. macleayi, USNM173817 (1). Arripidae: Arripis georgianus (Valen-ciennes), USNM 267149, 287442 Carangidae: Car-angoides crysos (Mitchill), USNM 167629; Seriola
sp., USNM 306575; Trachinotus falcatus (Linnaeus),USNM 280104. Chaetodontidae: Chaetodon trifas-ciatus Park, USNM 278739; C. melannotus Blochand Schneider, USNM266894; ForcipigerflavissimusJordan and McGregor, USNM 340962; Heniochusacuminatus (Linnaeus), USNM 147893. Coiidae:Coius sp., USNM 269799. Dinolestidae: Dinolesteslewini (Griffith and Smith), USNM 59932. Dinoper-CIDAe: Dinoperca petersi (Day), USNM 269543.Drepanidae: Drepane africana Osorio, USNM306264; D. longimanus (Bloch and Schneider),USNM 284472. Ephippidae: Chaetodipterus zonatus(Girard), USNM 220719, 220721; Ephippus orbis
NUMBER 11 75
Table 1.?Distribution of certain gill-arch muscles and SCL in genera of preacanthomorph fishes; homologies not implied. X = present;Y = present or absent; A = attached to Cb5; E = either OD3 or OD4 present, or only OD3-4 present; F = free from Cb5; L =longitudinal and transverse muscle layers present; T = only transverse muscle layer present: ? = presence or state questionable; * = stateof LI1.
Genera ? fNrgr^ro^t-^f a O- - <N rn en r<~, ^t ^t ^ Q Q Q Q^-_]JJjjJj_J_J_l_i_;_j_j_]_lOOOOOOOOOwc!ic::e:255ooHcnDiaphus X X X X X X X X X X X X 1. X FLampaiiyclits X X X X X X X X X X X X 1 X FNeoscopelus X X X X X X X X X X X 1 X FChlorophthulmus X X X X X X X X X X 9 FSaurida X X X X X X X X X X X X L FAulopus X X X X X X X X X X X X 9 F XAteleopus X X X X X X X X X 7 X 1 FGonostoma X X X X X X X X X X X I. X FMaurolicus X X X X 9 X X X X X 7 X FDiplophos X X X X X X X X X X X X 7 X FUmbra X X X X X X X X X X X L F XDallia X X X X X X X X X X L A XNovumbra X X X X X X X 1' !?: E X X X 1 F XEsox X X X X X X X X 7 X 1. F XSearsia X X X X X X X X X X 1 F XAlepocephalus X X X X X X X X X X 1 FArgentina X X X X X X X X X X X 1 FMallotus X X X X X X X X X X X X X 1 F"Hypomesus X X X X X X X X X X X X X 1 F XLovettia X X X X X X X X X X X ? FI.cpidogalaxias X X X X X X X X X X X X 9 FGalaxias X X X X X X X X X X X 1- AReiropinna X X X X X X X X X X 7 X X LCoregonus X X X X X X X X X X X 1 A XThymallus X X X X X X X X X X X 1 I' XOncorhynchus X X X X X X X X X X X X L F XGymnotus X X X X X X X X X 7 1 FDiplomysles X X X X X X X X X 9 1 1Xenocharax X X X X X X X X X X X X 1 !Brycon X X X X X X X X X X Y X X 1 FZacco X X X X X X X X X X X L AOpsariichthys X X X X X X X X X X X 1 AGonorynchus X X X X X X X X X ! FChanos X X X X X X X 1 FClupea X X X X X X X X X X X X 7 Y 1 FDussumieria X X X X X X X X X X X X " X L FChirocentrus X X X X X X X X X X 1 FCoilia X X X X X X X X X X 1 FCetengraulis X X X X X X X \ X X 7 1 FIlisha X X X X X X X X \ X X X X L FDenticeps X X X X X X X X X X X 1 FSynaphobranchus X X X X X X X 7 X X 1 FAnguilla X X X X X X X X X X X X 1 FConger X X X X X X X X X X X L FAldrovandia X X X X X X X X X 1 FNotacanthus X X X X X X X [ FPlerolhrissus X X X X X X X X X X X 1 F XAlbula X X X X X X X X X X X X I. F XElops X X X X X X X X X X X 1 F 9Megalops X X X X X X X X X X X L F 9Scleropages X X X X X X X X X X X X L FOsteoglossum X X X X X X X X X X X X X I. FPantodon X X X X X X 7 X X 9 X 1-Arapaima X X X X X X X X ? FHelerotis X X X X X X X X X 7 X 7 FMormyrus X 7 X X 9 ? X ? 7 7 7 1. FPetrocephalus X X X X 7 ? X 9 ? X X 1. FGymnarchus 7 7 X 7 7 7 X I, FNolopterus X X X X X X X X X X 7 X X 1. FHiodon X X X X X X 7 Y X X X X X I FAnna X X X X X X X X X L FAlractosteus X X X X X X X X TPolyodon X X X X X TAcipenser X X X X X TPolvpterus X X X X 1
76 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 2.?Insertion sites or states of LI2 and LI3 and state of RD, if present, ingenera of preacanthomorph fishes. X indicates insertion site or absence; ? = pres-ence or absence questionable. Data for LI2 and LI2' combined for Zacco. = RDdorsal to SO (SOD absent); 1 = RD ventral to SO (SOD absent); 2 = RD andSOD present. * = probably anomalous.
LI2 LI3
Genera j3 m "* 2 ? m "*< ? ? 5 5 i? uj RDDiaphus X X 2Lampanyctus X X 2Neoscopelus X X 2Chlorophthalmus X X X 1Saurida X X 1Aulopus X X X 1Ateleopus X X X 1Gonostoma X X 2Maurolicus X X 2Diplophos X X 2Umbra X XDallia X XNovumbra X XEsox X XSearsia X X XAlepocephalus X X X*Argentina X XMallotus X XHypomesus X XLovettia X XLepidogalaxias X XGalaxias X XRetropinna X XCoregonus X XThymallus X X XOncorhynchus X X XGymnotus X X X X XDiplomystes X X X X XXenocharax X XBrycon X XZacco X X X XOpsariichthys X X X XGonorynchus X XChanos X XClupea X XDussumieria X X X XChirocentrus X XCoilia X XCetengraulis X XIlisha X XDenticeps X XSynaphobranchus x X X XAnguilla X XConger X X X XAldrovandia X X XNotacanthus X XPterothrissus X X XAlbula X X XElops X XMegalops X XScteropages X X XOsteoglossum X X XPanlodon X X X 2Arapaima X XHeterolis X X XMormyrus ? ?Petrocephalus ? ?Gymnarchus X XNotopterus X X XHiodon X X ?Amia X XAtractosteus X XPolyodon X XAcipenser X XPolvplerus X X
NUMBER 11 77
Table 3.?TD muscle components in genera of preacanthomorph fishes. States within components not indicated. X = present; Uunique; ** = TD absent. Eb4* treated as Eb4. See also Table 4.
Genera
XIId3 x 25Id Id 'T ^A t * 4 ? t J -9x> x> x x r^B id(N (N rnrnmro ^?^"^t^j-JT1 <N X ? JD ?; en TfX X X X X X X X X X B? X UJ CUO-i?1 XX!d ?oi b* ? . a. cu b, b Wida.o-,D_Wi,_''' w u ? acd p. *> i cdp-iiii ! : - ~ ? ? ? ? . .V -T^ ,X X X X X X X XXXXXXXXXXXXXXXXXXXXXXXXX&-&&;WU-lpLipjpjtU uj a.CL.a.cuD.a.CL.cucucua,a.a.a.a-o.a-CL.cucL.cucuDHCucuDDDHHHS-S-S- H Hf-S-S-h-i-i-l-HHHHHHf-HI-Hf-f-f-l-Hf-S-Hf-HDiaphus X X X X XLampanyctus X X X X XNeoscopelus X X X X XChlorophthalmus X X X XSaurida X u X uAulopus X X X X u XAteleopus X XGonostoma X XMaurolicus u u XDiplophos X u X XUmbra XDallia X XNovumbra X X XEsox X XSearsia X XAlepocephalus X X XArgentina XMallotus X X XHypomesus X X XLovettia uLepidogalaxias XGalaxias XRetropinna X XCoregonus X X XThymallus X XOncorhynchus X XGymnotus X XDiplomysles u X II uXenocharax X X X XBrycon X X XZacco**Opsariichthys**Gonorynchus X X XChanos X XClupea X XDussumieria X XChirocentrus X X XCoilia X X uCetengraulis X X XIlisha XDenticeps X XSynaphobranchus uAnguilla uConger XAldrovandia X XNotacanthus X X XPterolhrissus X XAlbula X uElops X XMegalops X XScleropages X XOsteoglossum X XPantodon u XArapaima X uHeterotis XMormyrus X XPetrocephalus XGymnarchus uNotopterus XHiodon XAmia XAtractosteus XPolyodon XAcipenser XPolypterus**
78 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 4.?Skeletal elements to which TD attaches (Eb4 undifferentiated from EB4*;see also Table 3) and relation of OD4 (and/or OD4' or OD3-4; see also synapomorphies(30-40) in Results section) to Pb4 in genera of preacanthomorph fishes. A = Pb4 absent;P = Pb4 present, but TD not attached to it; X = attached to; ** = TD absent; 1 = origin
at least partly on Pb43; 2 = Pb4 present, but origin not on it.
Genera Skeletal elementsEbl Eb2 Eb3 Eb4 Pbl Pb2 Pb3 Pb4 UP3 UP4 UP5 OD4Diaphus X X X X p 2Lampanyclus X X X X p 2Neoscopelus X X X X P 2Chlorophthalmus X X X X X 1Saurida X X X X p 2Aulopus X X X X p 2Ateleopus X X A AGonostoma X p X 1Maurolicus X X X 1Diplophos X X p X 1Umbra X X X 2Dallia X X 2Novumbra X X 1Esox X X 1Searsia X X X 1Alepocephalus X X X 1Argentina X X 2Mallolus X X X 1Hypomesus X X X X 1Lovetlia X X X X 1Lepidogalaxias X p 2Galaxias X X X 2Retropinna X X X 1Coregonus X X X 1Thymallus X X X 1Oncorhynchus X X X 1Gymnotus X X X X 1Diplomystes X X X X X X 1Xenocharax X X X X 2Brycon X X X X 1Zacco** A AOpsariichthys** A AGonorynchus X X X 1Chanos X X p 2Clupea X X X 1Dussumieria X X X 1Chirocentrus X X X 1Coilia X X X X 1Celengraulis X X X 1Ilisha X X X 2Denticeps X X X X 1Synaphobranchus X A X X AAnguilla X X A X X AConger X X A AAldrovandia X X 1Notacanthus X X X A 2Pterothrissus X X 1Albula X p X 1EIops X X X 1Megalops X X X 1Scleropages X X X X 2Osteoglossum X X X X 2Panlodon X X X X A AArapaima X X X X 2Helerotis X X 2Mormyrus X X X 2Petrocephalus X X X 2Gymnarchus X X X A ANotoplerus X X X IHiodon X X X 2Amia X A AAtractosteus X A APolyodon X A AAcipenser X A APolvpterus** A A
NUMBER 1
1
79
Table 5.?Distribution of bilaterally paired pharyngobran-chial muscles in suprageneric groups of preacanthomorph fishes(all genera within each group are included). X = present; * =not illustrated.
? <n n m m U Wis\ (N ^- "n ^" <*Suprageneric Groups
o,5
J3uCN
a,s
w
0,
m
PL,7 4
a,54
a.
4 ?3a-5 w4a,D2
U2.D7.
(Jin
sAulopiformesChlorophthalmusSaurida XAulopusStomiiformesGonostoma XMaurolicusDiplophosEsociformesUmbraDalliaNovumbra XEsoxCharaciformesXenocharax X*Brycon X*ElopomorphaSynaphobranchus XAnguilla XConger X XAldrovandia XNotacanthusPterothrissus XAlbula X XElopsMegatopsOsteoglossomorphaScleropages XOsteoglossum X XPantodon XArapaima XHeterotis XMormyrus X XPetrocephalus X XGymnarchus XNotopterusHiodon
(Bloch), USNM 257868; Pfatax orbicularis (Forss-kal), USNM 268668. Epigonidae: Sphyraenops bair-dianus Poey, USNM 270279. Haemulidae: Haemu-lon sexfasciatus Gill, USNM 292785. Lobotidae:Lobotes pacificus Gilbert, USNM 82008. Monodac-TYLIDAE: Monodacytus argenteus (Linnaeus). USNM266897. Pempheridae: Parapriacanihus ransonnetiSteindachner, USNM 218867; Pempheris schwenkiiBleeker, USNM 269801. Percichthyidae: Bostockiaporosa Castelnau, USNM 218841; Gadopsis marmo-ratus Richardson, USNM 214836, 308109 (2). Po-macanthidae: Centropyge bicolor (Bloch), USNM56995; Pomacanthodes semicirculatus (Cuvier),USNM 273043. Scatophagidae: Scatophagies argus(Linnaeus), USNM 224393 (3), 259383; Selenetocamultifasciata (Richardson), USNM 173514, 245702(2). Scorpididae: Microcanthus strigatus (Cuvier).USNM 267047; Siganidae: Siganus vulpinus (Schle-
gel and Muller), USNM 325277. Trichiuridae: Tri-chiurus lepturus Linnaeus, USNM 272931.Muscle preparations in alcohol. Aplodactyudae:Crinodus lophodon (Gunther), USNM 227300. Car-angidae: Decapterus macrosoma Bleeker, USNM307977; Decapterus punctatus (Cuvier), USNM199037; Selene vomer (Linnaeus), USNM 338012.Centrarchidae: Acantharchus pomotis (Baird).USNM 237611; Ambloplites rupestris (Rafinesque),USNM 333731; Archoplites interruptus (Girard),USNM 39563; Centrarchus macropterus (Lacepede),USNM 243775; Lepomis auritus (Linnaeus), USNM243888; Pomoxis annularis Rafinesque, USNM129524. Chaetodonttdae: Chaetodon austriacusRuppell, USNM 267044. Chiasmodontidae: Chias-modon sp., USNM 1186139. Gempylidae: Neo-epinnula americana (Grey), USNM 366716; Pro-methichthys prometheus (Cuvier), USNM 289930.
80 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 6.?Character states for Eb5 and skeletal elements (otherthan Cb5) to which Ad5 attaches in genera of preacanthomorphfishes. = Eb5 absent; 1 = Eb5 autogenous; 2 = Eb5 putativelyfused to Eb4 (based on configuration of distal end of Eb4); 3 =Eb5 putatively fused to Cb4; X = major attachment; x = minorattachment. Eb5 character state 2 for Osteoglossomorpha reas-sessed in "Epibranchials 5 and 4" under section "Muscles andSkeletal Elements."
Genera Skeletal elements* ** 31 "1 "* "3t r )cur X> CO .o J3 X <JLbs w tu w u o <Diaphus XLampanyctus XNeoscopelus XChlorophthalmus 1 X XSaurida XAulopus 1 X XAteleopus XGonostoma XMaurolicus XDiplophos 1 X XUmbra 1 XDallia XNovumbra XEsox XSearsia 1 X XAlepocephalus 1 XArgentina 1 XMallolus 2 XHypomesus 2 XLovettia 2 XLepidogalaxias X XGalaxias 2 XRetropinna 2 XCoregonus 1 XThymallus 1 XOncorhynchus 1 XGymnotus 1 XDiplomystes 1 X XXenocharax 1 XBrycon 1 XZacco 1 XOpsariichthys 1 XGonorynchus 2 X XChanos 1 XClupea 2 XDussumieria 1 XChirocentrus 2 XCoilia 2 XCetengraulis 2 ?Ilisha 2 XDenticeps 1 XSynaphobranchus Ad 5 absentAnguilla XConger XAldrovandia 1 X XNotacanthus 1 or 2 X XPterothrissus 1 XAlbula 3 XElops 1 X XMegalops 1 X XScleropag.es 2 XOsteoglossum 2 XPantodon 1 XArapaima 2 XHeterotis 2 XMormyrus 2 XPetrocephalus 2 XGymnarchus 2 XNotoplerus 2 XHiodon 2 XAmia XAtractosteus XPolyodon XAcipenser XPolypterus Ad5 absent 1
Inermiidae: Inertnia vittata Poey, USNM 318643. Is-tiophoridae: Istiophorus sp., USNM 22999. Ky-phosidae: Kyphosus sectatrix (Linnaeus), USNM116955. Monodactylidae: Monodactylus argenteusLinnaeus, USNM 258894 (3), 266897. Notograpti-DAE: Notograptus guttatus Giinther, USNM 173798.Pomacanthidae: Centropyge bispinosus (Giinther),USNM 336476; C. vrolikii (Bleeker), USNM210295. Scatophagidae: Scatophagies argus (Lin-naeus), USNM 224393 Scombridae: Gasterochismamelampus Richardson, TMH D.1982; Euthynnus al-letteratus (Rafinesque). USNM 214658; Scombero-morus cavalla (Cuvier), USNM 289928; S. commer-soni (Lacepede), USNM 297407. Scorpididae: Scor-pis sp., USNM 339348. Siganidae: Siganus spinus(Linnaeus), USNM 233575. Trachipteridae: Tra-chipterus sp., USNM 346705.LampridiformesVELIFERIDAE
Velifer hypselopterus Bleeker, NSMT-P-59516, 30?S,168?E. 165 mm. Plate 59
Additional material. ? = Metavelifer multiradiatus(Regan), AMS 1.20605013, ca. 94 mm SL; USNM23953, cleared and stained.Remarks. AMS specimen dissection was faulty,but muscles, except for TEb2 and ODs, appear to beremarkably similar to V. hypselopterus.
Description.LEI on Ebl mid-dorsoposteriorly beginning lateralto base of uncinate process.LE2 on Eb2 mid-dorsoposteriorly.LE3 on anterior margin of cartilaginous tip of Eb3uncinate process, just dorsal to OD3.LE4 slender tendinous insertion on tip of Eb4 le-vator process.LP absent, possibly represented by discrete liga-mentous attachments, two on Eb4 levator process,one on Eb3 uncinate process, that fuse into broadsheet of fascia dorsally, which is also continuous withsheet of fascia representing PP.LI1 on Pb2 just ventroposterior to large dorsal,cartilage-tipped process.LI2 on Pb3 dorsolaterally anterior to medial endof Eb3.TD comprises completely separate TEb2 andbroad, more-or-less continuous TPb3-Pb4-Eb3-Eb4(attachments distinct laterally), which is posteriorlycontinuous with SOD. TEb2 consists of two, discon-tinuous parts. Left-side TEb2 originates as broad-dor-
sal and slender-ventral muscle straps (ventral strapnot visible in Plate 59) attached to thick CT pad en-veloping right-side Pb2 and Pb3 cartilaginous artic-
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Table 7.?Synapomorphies (as numbers), based on dorsal gill-arch muscles, TV4, and SCL, supporting Actinopterygian preacan-thomorph fish groups (see Results section for explanation).
Group Synapomorphy numberCtenosquamata 16Myc tophi formes 23Myctophidae 10Eurypterygii 20,22Neotelostei 15,28,29,53Stomiiformes 35,51Esociformes 60Neognathi 34Platytrochtidae +Alepocephalidae 40Osmeroidea 5,6Characiformes +Siluriformes +Gymnotiformes 27Gymnoti formes +Siluriformes 21Characiformes 8,49Cypriniformes 39,43,45,52,56Gonorynchiformes 38Clupeomorpha 19Clupeoidei 7Elopomorpha 58?Anguilliformes 12, 14,46Albuliformes 17,36,48,57Elopi formes 59?Elopocephala 33Teleostei 13,25,32,41,50,55aOsteoglossomorpha 3, 4, 42Osteoglossiformes 44Osteoglossoidei 13a, 26, 47Osteoglossidae 30Halecostomi 37,54,55Neopterygii 9, 11,24, 31, 42a?, 49a?Chondrostei 18Actinopteri 2Actinopterygii 1
uncinate process.
? Both muscles consist of loosestrands, but appear to be separate distal to their ori-gins.OP dorsally on posterior surface of Eb4 medial todorsal attachment of Ad4 and ventrally on dorsodistalsurface of Cb5 just posterior to posterior attachmentof Ad5.Ad 1-3 absent.Ad4 attaches dorsally to posterolateral surface ofEb4 and ventrally to Cb4 dorsally just anterior toinner angle of Eb4-Cb4 joint.Ad5 attaches anteriorly to Cb4 posterodistal endand AC4 posteriorly, and posteriorly to Cb5 distalend anterior to ventral attachment of OP.SOD present.RDs adjacent.Additional remarks. SCL attached mid-dorsally tocartilaginous ventroposterior tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. IAC absent. CTpad enveloping Pb2 and Pb3 processes gives rise toCT strap that attaches to cranium. AC4 and a muchsmaller AC3 present. Medial ends of Eb4 and Eb3about equal in size. ? AC3 and AC4 absent; medialends of Eb3 and Eb4 about equal in size.Olney et al. (1993) hypothesized that the Velifer-idae are the sister group of all other lampridiformsand that the Trachipteridae and Regalecidae are themost specialized. The Veliferidae lack ER as does theTrachipteridae (based on Trachipterus sp., USNM346705). LAMPRIDAE
Lampris guttatus (Briinnich), SIO82-70, not mea-sured (estimate >300 mm).Plate 60
ulating processes (pad mostly removed in Plate 59)and attaches to dorsomedial surface of left-side Eb2anterior to LE2 insertion. Right-side TEb2 originatesmedially from CT on left side and passes laterallybetween dorsal and ventral straps of left-side TEb2and inserts similarly on right-side Eb2. TPb3-Pb4-Eb3-Eb4 attaches to bony dorsoposterior surface ofPb3 and dorsomedial surfaces of Eb3 and Eb4 anddorsoanterior surface of Pb4. ? TEb2 damaged indissection, but could have been continuous across gillarches, with anterior edge attaching to CT pad en-veloping Pb2 and Pb3 processes.Remarks. The specialized medial overlapping ofTEb2 on each side was seen only in Lampris andVelifer, and supports close relationship of these twotaxa. The condition in Metavelifer should be verified.OD3 and OD4 originate together on dorsoanteriorbony surface of Pb3 and separate at about their mid-length before inserting broadly on anterior surface ofEb3 uncinate process and dorsomedial edge of Eb4
Description.LEI on anterior surface of posterior bony flangeof Ebl, cartilage-tipped uncinate process absent.LE2 on anterior surface of posterior bony flangeof Eb2 just anterolateral to LE2' insertion (see re-marks following LE2').LE2' on dorsal edge of posterior flange of Eb2,anterior surface appressed to posterior surface ofLE2, muscle fibers of LE2 and LE2' extend dorsally
at different angles.Remarks. Right-side LE2' cleanly separable fromLE2, but left-side pair did not separate cleanly. Ver-ification of distinctness in other, smaller specimensneeded.LE3 sheet-like, on tip of Eb3 uncinate process an-teriorly, muscle continuous dorsoposteriorly withpresumed LE4-LP complex.LE4 broad sheet, musculous distally, fascial prox-imally, inserting on Eb4 levator process, continuousventrolaterally in fascia with presumed LP fibers; fas-
82 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
cia continues ventrodistally around fourth arch andattaches to Cb5 distally.LP presumably represented by small strap of fibersarising from fascia at posterolateral edge of LE4 fi-bers (information from right side only; left side dam-aged); posterior fascia presumably continuous withPP.LI1 on Pb2 dorsoanterolaterally.LI2 on Pb3 dorsoanterolaterally.TD comprises TEb2 and TEb3-Eb4. TEb2 withdistinct right- and left-side muscle straps, each in-serting on respective Eb2 mid-dorsoanteriorly, ante-rior to LE2 insertion; straps originate medially fromventral surface of thick CT pad enveloping anteriorcartilaginous anterior ends of Pb2s and Pb3s and dor-sally from CT attaching to Pb3; left-side strap passesthrough right-side strap before attaching to CT; mus-cle extends laterally and attaches on Eb2 dorsoanter-iorly anterior to LE2 insertion. TEb3-Eb4 attaches todorsomedialmost surface of Eb3, dipping to level be-low that of attachment to dorsomedialmost surface ofEb4.Remarks. CT permeating TD obscures attach-ments; insertions should be verified on smaller spec-imen.OD3 origin on Pb2 dorsoanteriorly, more-or-lesscontinuous with OD4 origin, divides laterally withone branch, OD3, inserting on Eb3 uncinate process,and the other branch, OD3', inserting on Eb3 dor-soanterior surface.OD4 origin on Pb3 dorsoanteriorly; left-side in-serts on Eb4 levator process; right side inserts onboth Eb3 uncinate and Eb4 levator processes.OP dorsally broadly on Eb4 posteriorly ventral totip of uncinate process and extending laterally; ven-tral extent unclear.Ad 1-3 absent.Ad4 questionably absent (unusual if true), but ifpresent, much reduced, possibly represented by mus-cle fibers obscured from view by OP.Ad5 on Cb5 distally and Cb4 posterodistally atventral attachment of Ad4.SOD broad.RDs adjacent.Additional remarks. SCL present, attached mid-dorsally to posteroventral cartilaginous tip of Bb3.TV4 damaged, appears to have had a median septum,apparently free from Cb5s. IAC absent. Pb4 and UP4present. Eb4 uncinate process absent. Pb2 toothed.Tiny AC2 present on left side, absent on right side.PCI insertion restricted well medial to distal end ofCb5; muscles unusual in that right and left sides jointogether anteromedially.PolymixiiformesPOLYMIXIIDAEPolymixia lowei Gunther, USNM 159295, 89.3 mm,USNM 202153, 124 mm.
Plate 61
Description.LEI broadly on anterior surface of Ebl uncinateprocess just lateral to cartilage tip.LE2 on dorsoposterior edge of Eb2 just postero-lateral to lateral end of TEb2.LE3 on tip of Eb3 uncinate process just dorsal toinsertion of OD3-4 on Eb3.LE4 on Eb4 surrounding small Eb4 levator pro-cess, ventrolaterally continuous with broad PP fascia.LP slender, tendinously inserted among LE4 fibersbasally.LI1 broadly on Pb2 dorsal surface just anterome-dial to base of Pb2 uncinate process.LI2 on bony Pb3 dorsoposterolaterally meetingTPb3-Eb3 insertion anteriorly.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2with straps attaching to dorsal surface of cartilagi-nous and bony anterior end of Pb2 just anterior toLI1 insertion, lies dorsal to TEb2 anteriorly, and iscomplexly continuous ventrally with TEb2, and alsoposteriorly by diagonal muscle slip with TEb2. TEb2broad, with crossing straps extending mid-anteriorlyfrom attachments with TPb2 ventral surface and in-serting on Pb2 anteriorly, but muscle mainly insertingon Eb2 dorsoanteromedialmost surface, at mostreaching to opposite medial edge of LE2 insertion;muscle continuous posteriorly by crossing muscleslips with TPb3-Eb3. TPb3-Eb3 inserting on Pb3dorsoposteriorly, meeting anteromedial edge of LI2insertion, and on Eb3 dorsomedially.OD3?4 originating broadly on Pb3 bony dorsalsurface ventral to TEb2, inserting on Eb3 bony sur-face anteriorly beginning just ventral to tip of unci-nate process and on posterior bony surface of Eb4beginning just ventral to uncinate process.OP strap of muscle on Eb4 dorsoposteriorly ven-tral to LE4 insertion and medial to Ad4 on Eb4, ven-trally on Cb4 joining raphe (ER) with Ad5.M. SO-Pb3 (indicated, but not labeled on Plate 61as broad strap of longitudinal SO fibers lateral to eachRD) extends anteriorly, passing ventral to TPb3-Eb3and inserting on Pb3 dorsally anterior to RD inser-tion.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly, ventrally on Cb4medial to Eb4-Cb4 joint.Ad5 ventrally broadly on Cb5 dorsally beginning
at dorsodistal tip and extending medially; dorsallyjoining ER with OP ventrally at attachment of bothon Cb4. Tendinous tissue extends laterally from ERand is continuous with PP fascia, which attachesaround Eb4-Cb4 distally (incompletely indicated inPlate 61).SOD slender, continuous anteriorly by fine, diag-onal muscle slip with TPb3-Eb3 posteriorly.
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RDs slender, juxtaposed.Additional remarks. SCL attached mid-dorsally toventroposterior cartilaginous tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. IAC absent. Pb2toothed.
ParacanthopterygiiPercopsiformesPercopsiform monophyly and intrarelationships.?Rosen (1962) implied the monophyly of the Percop-sidae 4- Amblyopsidae + Aphredoderidae, whichsubsequently became known as the Percopsiformes(Greenwood et al., 1966:396). Later, Rosen (1985:42-45) was first to question the monophyly of thepercopsiform fishes, removing the Percopsidae, butaligning aphredoderids with amblyopsids on the basisof their thoracic anus and segmented premaxilla. Pat-terson and Rosen (1989:19?20) reaffirmed Rosen's(1982) findings, stating that they could find no char-acter synapomorphous for the Percopsiformes [of Ro-sen, 1962], They failed to note, however, that amongtheir many illustrations of paracanthopterygian dorsalgill-arch skeletons, the medial head of Eb4 of per-copsiforms is much broader than the combinedbreadth of the medial heads of any two other epi-branchials (first noted by Parenti (1993:181), who in-terpreted the character as a synapomorphy of atheri-nomorphs. percopsids, amblyopsids, and aphredod-
erids). Among the other paracanthops, the medialhead of Eb4 is comparatively narrow, and in only afew macrourid gadiforms, which are well removedfrom the percopsiforms in Patterson and Rosen'scladogram (1989:fig. 16), is the percopsiform con-dition approximated.An additional character, not noted previously, isthe presence of a levator process on the percopsiformEb4, which is present in some or all members of thethree families (e.g., the process is absent in Amblyop-
sis (Patterson and Rosen, 1989:figure 13E), but pres-ent in Chologaster?both Amblyopsisdae, thus, weconsider the process primitively present in percopsi-forms). The process is lacking in all other paracan-thops and present elsewhere among non-percomorphacanthomorphs only in Lampridae, Polymixia, andthe zeiform Xenolepadichthys.Murray and Wilson (1999) hypothesized a Percop-siformes comprising only percopsids and aphredod-erids. They placed the amblyopsids as a deeply em-bedded clade in the Anacanthini, which they placedas the sister group of Percopsiformes. They basedtheir classification on 47 osteological characters andincluded fossil taxa. They also overlooked the en-larged medial head of epibranchial 4 as a possiblecharacter, and simply disposed of the position of theanus and segmented premaxilla, which Rosen (1985)had used to unite aphredoderids and amblyopsids, by
stating, "The new placement of the amblyopsids re-quires viewing these characters as homoplasies [!]."Based on the three percopsiform taxa we examined(Aphredoderus sayanus, Aphredoderidae; Percopsisomiscomaycus, Percopsidae; Chologaster agassizi,Amblyopsidae) the group shares the following spe-cializations:1. OD4' (a branch of OD3-4) extending posteri-orly, passing dorsal to jointly articulating Eb3 andEb4 uncinate processes, and inserting on Eb4 levatorprocess. We have not found this muscle in any otheracanthomorph.2. OD origin includes Pb3 (mainly) and Pb2. ODonly originates on Pb3 in most other acanthomorphs;exceptions include: platycephalids, percids, caproids{Capros, but not Antigonia), elassomatids, gasteros-teids (only Apeltes), and hypoptychids.3. M. Pb2-Eb2 present. This is an uncommonmuscle with a spotty occurrence otherwise amongfishes (e.g., we found it in the distantly related par-acanthop family Ophidiidae, and the percoids: Mo-rone, Moronidae, and Sillago, Sillaginidae. Endo(2002:101, as OD2) reported that it is generally ab-sent in gadiforms, but present in lotines, gaidropsar-ines. phycines, most gadines, and the morid Lotella.He (2001:fig. 34) illustrated the musculature of Gair-dropsarus and and gadiform bregmaceratid, Breg-macerops).4. TEb2 absent. TEb2 is usually present in acan-thomorphs, but is notably absent in some other Par-acanthopterygii and in all Atherinomorpha (but pres-ent in Mugilomorpha, i.e., Agonostomus, Mugilidae,hypothesized sister group of the atherinomorphs(Stiassny, 1993; Johnson and Patterson, 1993).5. LE3 absent. Occurs variously among fishes, butis not common, except among Smegmamorpha andPolycentridae.6. Medial end of Eb4 much enlarged relative tomedial end of Eb3 (also common in many smegma-morphs, but not mugilomoph Mugilidae).7. Levator process present on Eb4. This processotherwise occurs variously among acanthomorphs,but uncommonly among pre-Percomorpha.8. An overall trapezoidal appearance, in dorsalview, of the combined transverses dorsales and ob-liqui dorsales. We have not seen such configurationin any other fishes.The combination of these specializations almostcertainly indicates the monophyly of the percopsi-forms. The presence of characters 4-6 commonlyamong Smegmamoipha, suggests that the percopsi-forms and smegmamorphs, may be more closely re-lated than current classifications indicate, a relation-ship most recently suggested by Parenti (1993). Theabsence of all three of these characters in the Mu-gilidae, would seem to preclude such relationship.Evidence from the gill-arch muscles is equivocal
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for resolving percopsiform intra-relationships. Ab-sence of an OD attachment to Eb3 is a specializationshared only by Chologaster and Percopsis. Con-versely, absence of a TPb3 attachment is a speciali-zation shared only by Chologaster and Aphredode-
rus. Non-musculature characters, however, such asthe advanced anus and segmented premaxilla sharedby Aphredoderus and Chologaster, strongly favor ahypothesis that they are the sister group of Percopsis.APHREDODERIDAEAphredoderus sayanus (Gilliams), USNM 238477,71.3 mm; USNM 238466, 69.0 mm.Plate 62
Description.LEI on and medial to small, sharp prominence atbase of Ebl uncinate process.LE2 on tip of expanded bony process on posterioredge of Eb2.LE3 absent.LE4 on Eb4 levator process dorsoanteriorly, join-ing raphe medially with OD4 insertion and anotherlaterally with LP insertion.LP insertion continuous with LE4 insertion ventro-laterally.LI1 on Pb2 anteriorly, ventromedial surface join-ing raphe with TPb2 just dorsal to LI1 insertion.LI2 broadly on Pb3 dorsolaterally, narrowly onPb4 anterolaterally and adjacent dorsoanterior edgeof UP4.TD comprises TPb2 (TPb2d + TPb2v), and TEb4.TPb2d dorsal to TPb2v, each sheetlike, united ante-
riorly; muscles together are pad-like and notchedmid-anteriorly, each with mid-longitudinal raphe,which join together mid-anteriorly. CT sheet overliesTPb2d raphe and is attached to it; TPb2d fibers formheart-shaped curve; TPb2d fibers curve broadly an-teriorly; muscles attach together on anteromedialmostsurface of Pb2 uncinate process, joining raphe therewith LI2, but continuing ventromedially to insert onanterolateralmost surface of Pb2 (insertion visibleonly in ventral view after removal of underlaying tis-sues). TEb4 triangular, apex anteriorly with fibersvariously joining SO longitudinal fibers and CT inmidline, posteriorly almost divisible transversely intotwo sections, anterior section maintains triangularity,posterior section more strap-like, two sections joininglaterally and attaching along most of posterior marginof Eb4 levator process, there joining raphe with OP.M. Pb2-Eb2 on Pb2 uncinate process and dorsalsurface mostly ventral to OD3-4, extending laterallyand inserting on most of dorsal surface of Eb2.OD unusually broad dorsoanteriorly, comprisesOD4' dorsally and OD3-4 ventrally; OD4' andOD3-4 originate inseparably, broadly on Pb3 andPb2 ventral to TPb2; OD4' extends posterolaterally
over joined Eb3 and Eb4 uncinate processes (but isnot attached to them) and inserts on anterior surfaceof Eb4 levator process, where it joins raphe with ven-troanterior edge of LE4; OD3-4 splits off ventrallyshortly posterior to joint origin with OD4' and insertson Eb3 uncinate process, with remaining portionpassing ventral to joined Eb3-Eb4 uncinate processesand inserting on Eb4 anterior surface medial to in-sertion of OD4'.Remarks. It is unusual for OD to attach to Pb2.OP dorsally on Eb4 posterior surface at and medialto levator process, joining raphe there with TEb4,ventrally on Cb5 dorsolaterally, ventrolaterally join-ing raphe with Ad5.Ad 1-3 absent.Ad4 on most of Eb4 posterior surface (medial halfof muscle anterior to OP) and distal half of Cb4 me-dial to Eb4-Cb4 joint.Ad5 dorsally on posterolateralmost two-fifths ofCb4 and ventrally on anterodistalmost end of Cb5,joining raphe medially with OP.SOD absent.RDs separate, adjacent.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Pb3 toothed. IAC absent.PERCOPSIDAEPercopsis omiscomaycus (Walbaum), USNM179711, 92.3 mm; USNM 334972, 72.1 mm.Plate 63
Description (based on larger specimen with remarkson smaller specimen).LEI on small bony prominence near base of Ebluncinate process.LE2 on expanded posterior edge of Eb2.LE3 absent.LE4 tendinously on dorsal tip of Eb4 levator pro-cess.LP on lateral surface of tendinous insertion of LE4and distalmost end of Eb4. Smaller specimen withLP inserting only on tendinous insertion of LE4.LI1 on dorsoanteriormost surface of Pb2.LI2 on Pb3 dorsolaterally.TD comprises TPb2 and TPb3-Eb4. TPb2 roughlyheart-shaped, notched anteriorly, with median raphe(from which CT sheet arises dorsally), lies dorsal toPb3 portion of TPb3-Eb4, but not continuous with it,comprises two scarcely separable parts (more obvi-ous in smaller, non-illustrated specimen) possiblyequivalent to TPb2d and TPb2v of Aphredoderus; an-terior part on dorsoanteriormost end of Pb2 and me-dial surface of uncinate process of Pb2; posterior partattaches to posterior surface of Pb2 uncinate processand forms raphe there with M. Pb2-Eb2. TPb3-Eb4broadly triangular, blunted apex anterior, long sideposterior, with irregular median raphe in anterior
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three-fourths, attaching by few muscle straps to Pb3posteromedially; long side slender laterally, attachingalong most of Eb4 dorsal arm, forming raphe later-ally with dorsal end of OP.Remarks. Left-side OP of large specimen compris-es two separate straps, lateral and medial. Right sidehas only lateral strap, with muscle area occupied bymedial strap on left side forming part of SO. In small-er specimen, OP is broad, single, and occupies jointarea of OP on left side of larger specimen.M. Pb2-Eb2 originates on Pb2 uncinate process atraphe with TPb2 posterior part and attaches alongmost of bony dorsal surface of Eb2.OD3 absent.OD unusually broad dorsoanteriorly. comprisesOD4' and OD4. OD4' originates broadly on Pb3bony dorsal surface ventral to TPb2, passes dorsal tojoined Eb3 and Eb4 uncinate processes, and insertson Eb4 levator process just medial to LE4 insertion.OD4 originates on Pb2 dorsally ventrolateral toOD4', joins OD4' just distal to origins, passes ven-tromedial to joined Eb3-Eb4 uncinate processes, andinserts on anterior surface of distal end of Eb4 ventralto OD4' insertion. In smaller specimen OD4 andOD4' are mostly fused, separating only just anteriorto their insertions on Eb4.OP dorsally on posteromedial surface of Eb4, join-ing raphe there with Eb4 portion of TPb3-Eb4, ven-trally forming raphe with Ad5 before both attach dor-sodistally to Cb5. No raphe with Ad5 in smaller spec-imen. See also remarks following TD.Ad 1-3 absent.Ad4 on distal half of ventral surface of Eb4, par-tially anterior to OP, and ventrally on Cb4 distal halfmedial to Eb4-Cb4 joint.Ad5 on posterodistal end of Cb4 and dorsodistalend of Cb5 forming raphe ventromedially with OPjust before attaching to Cb5. No raphe with OP insmaller specimen.SOD broad.RDs adjacent.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Tiny AC between right-side Pb2 and Ebl uncinate processes of larger spec-imen, possibly represents vestigial IAC, but IAC cod-ed as absent for this taxon.AMBLYOPSIDAE
Chologaster agassizii Putnam, USNM 232514, 65.4mm. Plate 64
Description.LEI on Ebl uncinate process about mid-dorsopos-teriorly; medial process absent.Remarks. Typically in acanfhomorphs the medial
Ebl process articulates with Pbl, and the uncinateprocess with Pb2 or IAC, which articulates with Pb2.LE2 beginning on about mid-dorsoposterior bonyedge of Eb2 and extending posteriorly onto CT be-tween Ebl and Eb2 (mostly on CT).LE3 absent.LE4 on Eb4 beginning at levator process anteriorlyand extending laterally along dorsoposterior edge ofOD4 insertion, and just medial to LP insertion.LP on Eb4 distally, anterior and slightly lateral tolateral edge of LE4 insertion.LI1 on Pb2 anterolaterally (at anteriormost junc-tion of Pb2 and Pb3).LI2 on Pb3 dorsolaterally.TD comprising TPb2d, TPb2v, and TEb4. TPb2da transverse strap, with median raphe ventrally at-tached to CT lining pharynx, laterally attached to Pb2just dorsoposterior to LI1 insertion and continuousand just anterior to TPb2v attachment. TPb2v a di-agonal strap on each side extending anterolaterallyfrom raphe with posterior end of OD4' on Pb3 onone side and attaching to Pb2 just posterior to TPb2don opposite side, some muscle strands passingthrough strands of contralateral TPb2v; asymmetri-cally continuous with TEb4. TEb4, roughly triangu-lar in shape, apex anterior, commencing along com-mon line with OD4' posteromedially, and extendingposterolaterally; muscle strands, somewhat inter-laced, becoming continuous with horizontal posteriorportion of muscle, and inserting on Eb4 dorsoposter-iorly; forming raphe with dorsal end of OP.Remarks. We have arbitrarily equated the two Pb2sections of Chologaster with TPb2d and TPb2v ofAphredoderus (see also Percopsis).M. Pb2-Eb2 originating on dorsoanterior edge ofPb2 (where it articulates with Ebl medial end) andinserting along most of bony dorsal surface of Eb2.OD3 absent.OD comprises OD4' and OD4, extremely broaddorsally and deep ventromedially, originating on Pb3dorsomedially. continuing onto Pb2 dorsolaterallyventral to TPb2v. Muscle joins raphe posteromediallywith posteromedial end of TPb2v of opposite sideand divides posterolaterally into OD4' and OD4, withOD4' passing posteriorly dorsal to joined Eb3 andEb4 uncinate processes and inserting on Eb4 levatorprocess just ventral (anterior in view) to LE4 inser-tion and lateral to TEb4 attachment, and OD4 passingventral to joined Eb3-Eb4 uncinate processes and in-serting on Eb4 dorsally medial to levator process.OP dorsally, broadly on posterior surface of Eb4,ventrally on dorsodistal end of Cb5 joining Ad5; me-dially continuous with SO.Ad 1-3 absent.Ad4 on most of dorsal surface of Eb4 anterior toOP attachment, and on distal half of surface of Cb4medial to Eb4-Cb4 joint.
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Ad5 dorsally on posterodistal fourth of Cb4, withslight tendinous extension to posterodistal end ofEb4; ventrally on distal end of Cb5, ventromediallyjoining raphe with OP.SOD present.RDs slightly separated.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 toothed. IAC absent.OphidiiformesOPHIDIIDAEDicrolene intronigra Goode and Bean, USNM362587, 2 specimens, 226-240 mm TL.Plate 65
Description.Remarks. Only one of the two sets of gill archeswas dissected to expose hidden muscle attachments,and the description of these is based mostly on thatspecimen. The remainder of the description is basedon the undissected specimen.LEI on anterior surface of Ebl uncinate processventrolateral to tip.LE2 on tip of prominent bony process on posteriormargin of Eb2, insertion continuous posteriorly asligament joining Eb2 with Eb3.LE3 on Eb3 uncinate process just ventrolateral totip, meeting insertion of OD3 portion of OD3-4.LE4 tendinously on dorsal edge of Eb4 lateral touncinate process and immediately dorsal to Ad4 at-tachment.LP absent.LI1 on dorsomedial edge of Pb2 joining raphe withTPb2p laterally and M. Pb2-Eb2 medially.LI2 on Pb3 dorsoposteromedially lateral to M. SO-Pb3 and RD; passes medial to OD3-4.TD complex, comprises TPb2, TPb2p, TPb3,TEb3, and, variably, TPb4 (absent in illustrated spec-imen). TPb2 very broad transversely, slender longi-tudinally, on bony edge lateral to cartilaginous edgeof dorsal margin of broad Pb2 dorsal process (entireedge is cartilaginous in other specimen), continuousbroadly posteriorly with TPb2p but overlappingTPb2p anterolaterally on right side (both sides non-illustrated specimen). TPb2p much shorter longitu-dinally than TPb2, passes dorsal to OD-3-4 origin,touching LI1, then extends ventrally medial to LI1and attaches on dorsomedial edge of Pb2; joins par-tial raphe with ventromedial surface of LI1; contin-uous by diagonal muscle strands with TPb3. TPb3 ondorsal surface Pb3 mid-laterally medial to lateral car-tilaginous process articulating with small AC (not il-lustrated) between process and medial end of Eb2(posteromedial cartilage-tipped process of Pb2 isventral to ACs and attached to them ventrally); con-tinuous posteriorly by diagonal muscle strands with
very broad TEb3. TEb3 dorsomedially on Eb3 bonysurface beginning slightly lateral to medial cartilag-inous end, continuous posteriorly by diagonal musclestrands with narrow TPb4 (TPb4 absent in illustratedspecimen, in which case continuous with SOD).TPb4 extends deeply laterally and attaches finely toPb4 dorsomedial surface; continuous posteriorly bydiagonal strands of muscle with SOD.M. Pb2-Eb2 on posterior edge of Pb2 dorsal pro-cess, just meeting lateral edge of TPb2 and the lateraledge of LI1 insertion, and on posterior surface ofraised anterior edge of Eb2 just anteromedial to LE2.OD3-4 origin on dorsoanterior surface of Pb3, di-vides posteriorly at mid-length or just before insert-ing narrowly on anterior surface of Eb3 uncinate pro-cess and medial edge of Eb4 uncinate process. Onleft side of illustrated specimen, an anomalous strapof OD3-4 attaches to Pb2 at the junction of LI1 andM. Pb2-Eb2.OP dorsally on Eb4 posterior surface below andmedial to uncinate process, ventrally joining raphe(ER) with Ad5 dorsolaterally where both attach tomedial end of CT pad enveloping posterolateral endof Cb4 (medial end of pad well separated from distalend of Cb4).M. SO-Pb3, longitudinal-fiber band originatingfrom SO in region of Eb4 and extending anteriorlyventral to TD components and inserting on Pb3 justventral to RD (q.v.).Ad 1-3 absent.Ad4 dorsally on dorsoposteriormost edge of Eb4beginning at and ventral to medial end of LE4 inser-tion and ending slightly medial to distal end of Eb4;ventrally on Cb4 dorsal surface medial to Eb4-Cb4joint.Ad5 dorsolaterally joining raphe (ER) with OP onfleshy CT pad enveloping distal end of Cb4, mediallyon posterior surface of lateral fourth of Cb5, joiningraphe there with TV5SOD present.RDs well separated, extending far anteriorly ven-tral to TD components and inserting on Pb3 just pos-terior to OD3-4 origin.Additional remarks. SCL absent. TV4 in two un-connected parts; anterior part with strong mid-lon-gitudinal raphe, originating on Cb4s ventral to pos-terior part and attaching mid-dorsoanteriorly to ven-tral surface of cartilaginous Bb4; ventral part contin-uous, originating on Cb4s dorsoposterior to anteriorpart, free from Cb5s. We have seen this type of TV4elsewhere only in the lophiiform Chaunax (Chauna-cidae), which differs in that a fine muscle strand joinsthe two portions and the two halves of the anteriorportion are separate at their attachment to the largecartilaginous Bb, which may be Bb4 or a complex.The dorsal tips of both Pbls are cartilaginous inone specimen and bony in the other. Pb4 and UP4
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present. IAC present. Eb4 levator process absent. Pb2toothed. Medial end of Eb3 and Eb4 about same size.There is a separate ball of cartilage between themedial end of Eb2 and the cartilage-tipped processmid-laterally on Pb3. Markle (1989:fig. 1A) illustrat-ed the dorsal gill-arch skeleton of a specimen heidentified as the neobythitine Dicrolene intronigra.The illustration does not accord with the skeleton ofour two specimens. For instance, he shows Eb2 ar-ticulating with Pb2 and does not indicate the presenceof an accessory cartilage either between this articu-lation or one attached to Pb3. Nor does he show acartilage-tipped process mid-laterally on Pb3. Patter-son and Rosen (1989:fig. 13g) illustrated the dorsalgill-arch skeleton of Monomitopus torvus Garman(another neobythitine), which is very similar to thatof our specimens of D. nigra. Consequently, one ofthe characters (contact between Eb2 and Pb2) Markleused to define a gadiform-batrachoidiform relation-ship apparently is invalid or needs modification.
Brotula multibarbata Temminck and Schlegel,USNM 340397, 166 mm, 214124, 133 mm.Not illustrated
Description.LEI on highest point of raised dorsoposterior bonyedge of Ebl, uncinate process lacks cartilaginous tip.LE2 on apex of prominent raised dorsoposteriorbony edge of Eb2.LE3 on dorsoanterior surface of Eb3 uncinate pro-cess lateral to cartilaginous tip.LE4 on bony dorsoposterior edge of Eb4 well lat-eral to uncinate process, fibers of posterior surfaceattaching on dorsal edge of CT covering Eb4 poste-
rior surface.Remarks. Dorsolateral fibers of OP and dorsome-dial fibers of Ad4 attach to same CT ventrally. Re-leasing CT from Eb4 results in strip of muscle ( =LE4-OP sling of Stiassny and Jensen, 1987) inter-rupted by band of CT.LP absent.LI1 on Pb2 dorsolaterally ventromedial to TPb2attachment to Pb2; larger than LI2.LI2 on dorsoposterolateral surface of Pb3 just me-dial to medial head of Eb3; muscle passes amidstOD3-4 dividing it into OD3-4 and OD4 (see OD3-4).TD comprises TPb2, TPb2p, TEb2. and TEb3.TPb2 a bean-shaped pad. covered with thin CT, at-tachment beginning anteriorly with CT of roof ofpharynx, continuing laterally and ending on anterior-most cartilaginous tip of Pb2, free edge continuingposterolaterally from this point and overlying all ODcomponents anteriorly; muscle attached ventrally todorsoanterior surface of Pb3 together with OD inser-
tions; broadly continuous posteriorly with TPb2p.TPb2p on Pb2 dorsolaterally at and medial to M.Pb2-Eb2 origin, posteriorly broadly continuous withTEb2. TEb2 on Eb2 dorsoanteriorly well medial toLE2 insertion, forming short raphe with ventral edgeof M. Pb2-Eb2 insertion and posterolateralmost edgeof OD2 insertion, broadly continuous posteriorlywith TEb3. TEb3 on dorsal surface of Eb3 medial tobase of uncinate process; broadly continuous poste-
riorly with SOD.OD comprises OD2, OD3-4, and OD3'. All ODcomponents originate together on Pb3 dorsoanterior-ly; components separate shortly distal to origin, withOD3-4 being variously split by penetration of LI2on its way to its insertion and recombining as theyinsert on Eb3 and Eb4: on anterior and posterior sur-faces, and medial edge of Eb3 just ventral to tip ofuncinate process, fusing ventrally with dorsal originof RecD3; on anterior surface and medial edge ofEb4 just ventral to tip of uncinate process, meetingOP dorsally. OD3' inserts on Eb3 dorsally ventral toOD3-4 insertion on Eb3. continuous there with ven-tral origin of RecD3.OD2 inserts on raised dorsoanterior edge of Eb2immediately ventral to M. Pb2-Eb2, there joining ra-phe with RecD3 insertion; posterolateralmost edge ofOD2 forming short raphe with TEb2.OP with two slightly separated attachments on Eb4posterior surface; dorsomedial attachment on unci-nate process, posteriorly overlapping ventrolateral at-tachment on Eb4 lateral to uncinate process, fusingtogether before ventrally joining raphe with dorso-lateral end of Ad5 (CT extension from raphe contin-ues onto Cb4 posterior surface and posteroventral fi-bers of Ad4 join raphe). Also see remarks under LE4.M. Pb2-Eb2 tendinously on anteriormost cartilag-inous tip of Pb2, to which TPb2 and Ebl uncinateprocess attach; broadly, musculously on Eb2 dor-soanterior edge.RecD3 with two separate origins, dorsal origin onEb3 uncinate process ventral to and fusing withOD3-4, ventral origin on dorsoanterior edge of Eb3,both parts fusing and inserting on Eb3 along insertionline of OD2.Ad 1-3 absent.Ad4 on Eb4 posterior surface lateral to OP, ven-trally on Cb4 dorsoposteriorly, attaching to CT ex-tending from ER. Also see remarks under LE4.Ad5 broadly on dorsodistal edge of Cb5, narrow-ing dorsally and joining raphe, which continues asCT on Cb4, with OP; attachment to Cb4 is well me-dial to distal end.SOD present.RDs present, separated.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Pbl with cartilaginous
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ends. Pb2 toothed. Medial end of Eb3 larger than thatof Eb4. IAC absent.BYTHITIDAECalomopteryx jeb Cohen, USNM 208341, 57.6 mm.Not illustrated
Description.LEI on Ebl mid-dorsoposteriorly, anterior (phar-yngobranchial supporting) process absent.Remarks. Anterior process considered absent (un-cinate present) because medial end of Ebl articulateswith Pb2.LE2 on Pb2 posteriorly a little lateral to mid-length, just lateral to lateral end of M.Pb2-Eb2.LE3 on Eb3 just lateral to cartilage tip of uncinateprocess.LE4 on Eb4 dorsoposteriorly between uncinateprocess and distal tip of Eb4.LP absent.LI1 broadly on Pb2 dorsoanterolaterally, ventro-medially joining raphe with TPb2 laterally, largerthan LI2.LI2 penetrating OD3-4, dividing it into almosttwo equal portions, and separating TPb3 posteriorlyfrom TEb3, while passing ventral to TEb3 on wayposteriorly to insertion on Pb3 dorsoposteriorly nearmedial end of Eb4.Remarks. Penetration of OD3-4 by LI2 is uncom-mon. Occurs also in related Ophidiidae and Carapi-dae, and unrelated Champodontidae and Odontobu-tidae (synapomorphic for all genera of latter).TD comprising TPb2, TPb3. and TEb3. TPb2broad, flat, thin, attaching to Pb2 beginning at ante-rior end and extending posteriorly to medial edge ofLI2, attached anteroventrally to CT of pharyngealroof, dorsally infiltrated with filmy CT, which coversTD, continuous posteriorly by diagonal musclestrands with TPb3. TPb3 slender, extends ventrolat-erally and slightly anteriorly anterior to LI2 (as latterextends ventroanteriorly from origin) and attaches toPb3 dorsally anterior to medial end of Eb3, contin-uous mid-dorsoposteriorly with TEb3. TEb3 broadmid-dorsally, extending narrowly laterally, passingdorsal to LI2 (as latter extends ventroanteriorly fromorigin) and attaches moderately broadly on Eb3 dor-somedially, broadly continuous mid-dorsoposteriorlywith SOD.M. Pb2-Eb2 medially broadly on Pb2 laterally be-ginning at anterior end and extending posteriorly toLI1 insertion, laterally on Eb2 dorsally, reaching me-dial edge of LE2 insertion.OD3-4 anteriorly, broadly on Pb3 dorsoantero-medially, almost completely divided longitudinallyby passage of LI2, posteriorly on Eb3 dorsoanteriorlybeginning immediately ventral to fine cartilage tip ofuncinate process, there meeting LE3 insertion ante-
roventrally, and on medial edge and posterior surfaceof Eb4 uncinate process.OP dorsally, broadly on Eb4 posteriorly beginningventral to uncinate process and extending medially,ventrally joining ER on Cb4 with Ad5 dorsally, dor-solateral^ slightly overlapping Ad4 medially, whichis also on Eb4 a little ventral to level of OP attach-ment.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly, beginning me-dially just anteroventral to lateral edge of OP andextending laterally to end of Eb4, ventrally on Cb4dorsally, beginning near lateral end and extendingmedially anterior to OP.Ad5 ventrally on Cb5 distally, joining ER dorsallywith OP ventrally on Cb4, beginning well medial todistal end of Cb4 and continuing medially.SOD present.RDs separated by distance less than one RD di-ameter, extending far anteriorly and inserting on Pb3sdorsally.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 absent, UP4 present. Pb2 toothed. Pbl ab-sent. IAC absent. Medial end of Eb3 larger than thatof Eb4. PCI begins attachment to Cb5 well medial todistal end of Cb5, attaches by moderately long ten-don to cleithrum. CARAPIDAE
Not examined.
Remarks. Vandewalle et al. (1998) briefly de-scribed and illustrated the gill-arch skeleton and mus-culature of the Carapidae. The publication was avail-able to us only as a poor quality photocopy. Never-theless, several important features were apparent.Some of the specializations they report (LP absent,LI2 penetrates OD, etc.), are shared with bythitidsand some ophidiids.All levators originate on inner surface of the hy-omandibula (we did not take note of the origins inour dissections of other ophidioids, which Vande-walle et al. noted would be of interest).LE1-LE4 present.LP absent.LI1 present (their LINT 2/3 or LINT3) insertingon Pb2 and Pb3 or Pb3, depending on genus.LI2 (their LINT4) inserting on "Pb4" (probablyUP4, presence of cartilaginous Pb4 not mentioned),penetrates OD3, which they recognize as OD3s andOD3p.IAC absent, but IAB (with cartilaginous ends)present, which Vandewalle et al. term an "interarcualelement." Ebl uncinate process apparently absent.TPb2, TEb2, and TEb3 present.ODl,2,3s,3p,4. Originating variously on Pb2. Pb2,
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Pb2 and 3, Pb4 [= UP4?], respectively, and insertingon Ebl, Eb2. Eb3, Eb3, Eb4 respectively.OP originating on Pb4 [= UP4?] or Eb4 dependingon genus, and inserting on Cb5.RecDs [their REDOs]. RecDl origin on Pb2, in-sertion on IAB; RecD2, origin on IAB, insertion onEb2; RecD3 origin on Eb2. insertion on Eb3; RecD4,origin on Eb3, insertion on Eb4.Ad 1-3 apparently absent.Ad4 apparently absent, but should be verified.Ad5 on distal end of Cb5 and on Eb4 (and in onegenus, also on Eb3).SOD apparently absent.RDs originate on second and third vertebrae, insert
at front of Pb3s.TV4 present, not attached to Cb5s.GadiformesEndo (2002) published a phylogenetic study of theof the Gadiformes. He discussed very little about thedorsal gill-arch musculature, but mentions (2000:82),following other authors, that TDA comprises onlyTEb2 in the gadiforms he examined. He (2002:fig.34) only illustrates (partially) the dorsal gill archmusculature of Gaidropsarus (Gadropsaridae) andBregmaceros (Bregmacerotidae). The musculature ofGaidropsarus is similar to that of Raniceps and Op-sanus (a batrachoidiform) in lacking LE3. but in-cludes M. Pb2-Eb2 (labelled as OD2), which neitherof the other two genera have. Endo (2002: 101 ) notedthat M. Pb2-Eb2 "is generally absent in most gadi-forms, but present in lotines. gaidropsarines. phyci-nes, gadines (except Gadiculus, Brosme) and themorid Lotella." His illustration of Bregmaceros alsoshows that it lacks LE3, as well as M. Pb2-Eb2 andLP. In Bregmaceros, he illustrates only the dorsoan-terior insertion of a muscle he identified as OP. whichattaches to Pb3 anteromedially. We suspect that thismay equate to our M. SO-Pb3.M. Pb2-Eb2 is commonly present in paracanthop-terygians.
RANICIPITIDAERaniceps raninus (Linnaeus), USNM 345222, 105mm S; USNM 307233, 105 mm.Plate 66
Description.LEI on Ebl dorsoanteriorly lateral to tip of unci-nate process.Remarks. Ligament extends medially from carti-laginous tip of uncinate process to dorsolateralmostcartilaginous edge of Pb2. Another ligament extendsposterolaterally from cartilaginous tip of uncinateprocess to anterior edge of Eb2 anterior to lateralmostend of TEb2, and is here joined by a third ligament
extending directly posterior from the posterior edgeof Ebl posterior to LEI insertion.LE2 on raised bony dorsal edge of Eb2 posteriorlyabout mid-laterally.LE3 absent.LE4 massive, on most of Eb4 bony surface dor-soposteriorly lateral to uncinate process.LP very small, easily removed or overlooked: join-ing LE4 insertion about mid-anteriorly.LI1 clasps edge of Pb3 between two anterior car-tilage-tipped processes (strut 1 and strut 2 of Markle,1989:fig. 2B), dorsally ventral to OD4 origin andventrally dorsal to Pb2.LI2 on Pb3 posterolaterally just anterior to artic-ulation with Eb4 and ventral to anteriorly extendingSO muscle strap, which passes ventral to Eb4 portionof TPb3-Eb3-Eb4.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2broad, bandana-like, attaching broadly mid-ventrallyto CT of pharyngeal roof, extending laterally and at-taching to Eb2 dorsally anterior to LE2 insertion,continuing broadly posteriorly as TPb3-Eb3-Eb4. In-dividual sections of TPb3-Eb3-Eb4 relatively distinctand well separated laterally, but obscured from viewby OD4, which lies dorsal to them: Pb3 sectionbroadly dorsally on Pb3 process articulating with Eb2medial end. separated by LI2 from Eb3 section,which attaches to dorsomedial bony surface of Eb3and is separated laterally, but adjacent posteriorly, toEb4 section, which attaches to ventromedial edge ofEb4 uncinate process (SO fibers attach to Eb4 dor-sally ventromedial to Eb4 section). Eb4 section con-tinuous mid-posteriorly with SOD.OD3-4 origin on Pb3 dorsomedially ventral toTEb2: insertion mainly on dorsomedial edge of Eb4uncinate process with slight tendinous connection touncinate process of Eb3.OP fused laterally with Ad4 and ventromediallywith SO, joined ventrolaterally to raphe (ER) withAd5: most clearly represented by strands on bonysurface of Eb4 uncinate process posteriorly and Cb5ventrally.Adl-3 absent.Ad4 fused medially with OP, ventrally joining ra-phe with Ad5 dorsolaterally; most clearly representedby broad attachment to Cb4 dorsally medial to Eb4-Cb4 joint.Ad5 dorsally on Cb4 well medial to distal end:ventrally on Cb5 dorsodistally; dorsolateral marginoutlined by raphe with Ad4 and OP, but free dorso-medially and medially.Remarks. It is unusual in acanthomorphs for theattachment of Ad5 to be so far removed mediallyfrom the distal end of Cb4. Condition also occurs inOpsanus (Batrachoididae), Lactarius (Lactariidae),cottoids, and Xenolepidichthys (Grammicolepidae).SOD broad.
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RDs separated by distance less than diameter ofone RD.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl tiny and cartilaginous. Pb4 and UP4 ab-
sent. IAC absent.
BatrachoidiformesBATRACHOIDIDAEOpsanus beta (Goode and Bean), USNM 301938,76.3 mm. Plate 67
Description.LEI on anterior surface of raised bony process onEbl.LE2 on anterior surface of raised bony process onEb2.LE3 absent.LE4 very broadly on dorsolateral surface of Eb4.LP absent.LI1 mainly on anterolateral edge of Pb3 with mi-nor attachment to anterior edge of Pb2 medially.LI2 on Pb3 dorsolaterally, immediately adjacent tomedial end of Eb3.TD comprises TEb2 and TEb3-Eb4. TEb2 broadwith mid-longitudinal raphe, which gives rise to CTsheets dorsally; attached mid-ventroanteriorly to CTof pharyngeal roof; muscle extends laterally and at-taches on Eb2 dorsally to point anterior to mid-pointof LE2 insertion. TEb3-Eb4 begins on dorsoantero-medialmost edge of Eb3 and continues around to pos-teromedial edge, with ventroposterior fibers attachingonto Eb4 dorsomedial surface and posteromedialedge.OD3-4 origin broadly on Pb3 dorsally with someventral muscle strands extending from origin ontoEb3 dorsomedially, slightly overlapping TEb3-Eb4on Eb3; minor insertion on medial edge of Eb3 un-cinate process, mainly on Eb4 uncinate process me-dially and posteromedially, joining raphe with OPdorsally.OP left side dorsally on Eb4 uncinate process pos-teriorly, joining raphe with OD3-4 on Eb4, right sidedorsally (anomalously?) on Eb3 and Eb4 uncinateprocesses posteriorly, ventrally on Cb5 dorsally me-dial to distal end, ventromedially not completely sep-arable from SO.Remarks. PCI (removed in Plate 67B) attachespartly to distalmost end of Cb5, but its tendinous dor-soanterior edge is joined by OP ventrolaterally and itmeshes with Ad5 ventrally just dorsal to distal endof Cb5.Ad 1-3 absent.Ad4 dorsally on ventral edge of distal half of Eb4medial to Eb4-Cb4 joint, its medialmost edge anter-
oventral to OP; ventrally dorsal edge of distal half ofCb4.Ad5 dorsally on Cb4 well medial to distal end,meeting Ad4 ventromedially, ventrally on Cb5 dis-tally, its posteromedial surface meshing with PCI justdorsal to Cb5.Remarks. It is unusual for the attachment of Ad5to be so far removed medially from the distal end ofCb4. Condition also occurs in Raniceps (Gadidae),Lactarius (Lactariidae), cottoids, and Xenolepidi-chthys (Grammicolepidae).SOD absent.RDs slightly separated.Additional remarks. SCL attached mid-dorsally toposteroventral end of Bb3. TV4 free from Cb5s. Pb4and UP4 absent. Pb2 toothed. IAC present, reduced.
LophiiformesCHAUNACIDAEChaunax pictus Lowe, USNM 187752, 73.3 mm.Plate 68
Description.Remarks. The anterior ends of the Pbs of Chaunaxappear to have been rotated dorsally and the gill arch-es are crowded close together.LEI slender, on dorsal edge of Ebl uncinate pro-cess just lateral to tip.LE2 very broad, on dorsoanterior edge of Eb2,meeting M. Pb2-Eb2 insertion posteriorly.LE3 absent.LE4 stout, on most of dorsal surface of Eb4 lateralto uncinate process, insertion meeting dorsal attach-ment of Ad4 on one side and insertion fibers inter-mingling with those of Ad4 on other side in what weconsider a "sling" (Stiassny and Jensen, 1987); mus-cle fibers of left side twist clockwise toward origin,those of right side twist counterclockwise.LP absent.LI1 mainly on dorsolateralmost edge of Pb3, butslight attachment to distinct spongy CT pad attachingto dorsal cartilaginous tip of Pb3 and, on right sideonly, a few fibers attach to dorsalmost tip of Pb2;joins raphe anteroventrally with dorsoposterior edgeof M. Pb3-Eb3.LI2 on dorsoposterior surface of Pb3 lateral to baseof dorsal process.TD comprises TEb2, TPb2, and TEb4; TEb2 high-ly modified, fails to attach to Eb2 (unique amongacanthomorphs we examined); muscle mostly ventralto TPb2, divided mid-longitudinally by broad raphe,which gives rise to thin CT sheets attaching to skull;anteromedially muscle on dorsoposterior surface ofPb2, attaching to spongy CT pad anterolateral to mid-longitudinal raphe, laterally with short converginganterolateral and posterolateral sections joining raphe
NUMBER 1 1 91
on dorsolateralmost edge of Pb2, which in turn isjoined by another raphe laterally with M. Pb2-Eb2medially. TPb2 a semicircular ribbon on each sidearising anteriorly and posteriorly from mid-longitu-dinal raphe of TEb2, also forming raphe mid-laterallywith TEb2. TEb4 on Eb4 posteriorly well medial touncinate process and well lateral to medial end ofEb4.M. Pb2-Eb2 originates anteriorly on Pb2 dorsola-teralmost edge, there joining raphe with TEb2 later-
ally, and inserts on Eb2 along anterior edge of broadLE2 insertion.OD3-4 originates broadly on Pb3 anteromedially,joining raphe dorsoposteriorly with ventrolateraledge of LI1; lateral fibers extend ventrally and inserton Eb3 dorsally well medial to uncinate process, re-maining fibers attenuate posteriorly insert by fine ten-don on Eb4 uncinate process just ventral to cartilagetip.OP dorsally on Eb4 posteriorly below and slightlymedial to uncinate process, ventrally on Cb5 dorso-posteriorly, ventromedially just meeting TV5.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 ventral to LE4 in-sertion, ventrally very broadly on Cb4 dorsal surfacemedial to Eb4-Cb4 joint.Ad5 absent.Remarks. The complete absence of Ad5 is uncom-mon in fishes. Even when not distinct, an indicationof its fusion with OP or another muscle is usuallydiscernible.SOD present.RDs adjacent.Additional remarks. SCL absent. TV4 in two dis-tinct parts, dorsal section a bilateral pair of musclesattaching separately to ventral surface of broad Bbcartilage plate; ventral section continuous, free fromCb5s, the two parts continuous by a fine musclestrand. A very similar TV4 is found only in Dicro-lene (Ophidiidae). q.v. Pb4 and UP4 absent. IAC ab-sent.
AcanthopterygiiStephanoberyciformesMELAMPHAIDAEPoromitra capito Goode and Bean, USNM 258325,96.1 mm. Plate 69
Additional material.
?
= Scopelogadus mizolepisbispinosus (Gilbert), USNM 356388, 81.7 mm.
Description.LEI on dorsoposterior edge of Ebl distal to un-cinate process.LE2 on raised mid-dorsoposterior edge of Eb2.
LE3 on Eb3 dorsally lateral to uncinate process.LE4 on posterodistal margin of Eb4 barely meet-ing LP anteriorly.LP on Eb4 at and just posterior to LE4 insertion.LI1 approximately mid-medially on dorsal surfaceof Pb2 anterior to uncinate process, smaller than LI2.
?
Laterally on dorsal surface of Pb2 anterior to un-cinate process.LI2 on Pb3 dorsoposteriorly medial to medial endof Eb3, larger than LI1.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2broadly on dorsal surface of anterior arm of Pb2 an-terior to uncinate process, posterormedianly dorsal to(appressed on) anterior end of TEb2, which it joinsalong mid-longitudinal raphe. TEb2 attaches on dor-soposterior edge of Eb2 medial to LE2 insertion.TPb2 and TEb2 are dorsal to and discontinuous fromTPb3-Eb3, which is very thin, broad, sheet-like, andventrally joins CT lining pharynx dorsally. Anteri-
orly, TPb3-Eb3 attaches broadly on Pb3 mediallyparalleling origin of OD3-4; posteriorly, TPb3-Eb3expands laterally and attaches to dorsomedial end ofEb3. Change in muscle fiber direction demarcatesseparation of TPb3-Eb3 from SO, with which it isposteriorly broadly continuous; @ Eb3 muscle por-tion continuing posteriorly as band of crisscrossingfibers, which change abruptly posteriorly to all trans-verse SO fibers.OD3-4 originates broadly along dorsolateral edgeof Pb3 and inserts dorsally on joined Eb3-Eb4 un-cinate processes.OP dorsally on Eb4 between Ad4 and SO attach-ments to Eb4, ventrolaterally attaches to dorsal edgeof gill raker contained in CT dorsal to Cb5, ventro-medially forms raphe (ER) with Ad5, scarcely sep-arable medially from SO. ? Well separable mediallyfrom SO.Ad 1-3 absent.Ad4 dorsoposteriorly on bony and cartilaginousdistal end of Eb4, attached by slender tendon ven-trally to dorsal surface of Cb4 slightly anterior toinner angle formed by Eb4-Cb4 joint. ? Attachedmusculously to Cb4.Ad5 narrowly, tendinously attached to distal endof Cb4; mid-dorsally more broadly attached to pos-teroventral surface of gill raker embedded in CT; dor-somedially joins raphe (ER) with to OP ventrome-dially; not separable medially from OP or SO; ven-trally on Cb5 dorsolaterally, ventromedially contin-uous with SO. ? Attached musculously to Cb4.SOD absent.RDs small, widely separate, inserting on Pb3 dor-soposteriorly; broad, short, thin band of longitudinalSO fibers extends anteriorly on each side and slightlyoverlaps respective RD insertion. ? Inserts on carti-laginous posteromedial end of Pb3 and medial endof Eb4.
92 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Additional remarks. SCL absent. TV4 free fromCb5s. Slender ligament (not illustrated) attaches tipof small prominence at cartilaginous medial end ofEb2 to Pb3 anterior to uncinate process; ? ligamentabsent. Autogenous rod-like cartilage attached toventroposterior cartilaginous end of Bb3; ? rod-likecartilage absent. Pb4 absent, UP4 present. Pbl withcartilage caps dorsally and ventrally. Pb2 toothed.Eb4 levator process absent. Medial end of Eb4 largerthan that of Eb3.Similar modified gill-raker noted in OP descrip-tion, otherwise noted in this study to occur only inScomber. GIBBERICHTHYIDAEGibberichthys pumilus Parr, USNM 214207, 79.5mm. Plate 70
Description.LEI, weak, on Ebl near tip of uncinate process.LE2 on posterodorsal edge of Eb2 somewhat prox-imal to mid-length.LE3 on dorsoanterior surface of Eb3 uncinate pro-cess just dorsal to OD3-4 insertion.LE4 on dorsodistal end of Eb4, joining LP inser-tion posteriorly, joining narrow raphe with Ad4 dor-
sally.LP slender, on Eb4 joining LE4 insertion antero-laterally.LI1 on Pb2 dorsoanteriorly ventral to anterior sec-tion of TPb2 (divides TPb2 into anterior and poste-
rior sections).LI2 on Pb3 mid-dorsoposteriorly, lateral to andmeeting lateral edge of TPb3 attachment.TD comprises TPb2 and TPb3. TPb2 on Pb2 an-terior process and basal area of uncinate process, di-vided into anterior and posterior sections by LI1; an-terior section partially overlies anterior end of pos-terior section; few muscle strands of anterior sectionattach to anterior end of right-side Pb3 and fewstrands of posterior section join OD3-4. TPb3 onPb3 posterolaterally, meeting medial edge of LI2 in-sertion, posteriorly continuous with SOD.OD3-4 anteriorly broadly on Pb3 anteromediallyventral to TPb2, divides at about posterior fourth oflength with anterior branch attaching to Eb3 uncinateprocess and posterior branch attaching to Eb4 unci-nate process.OP dorsally on Eb4 posteriorly beginning just ven-tral to tip of unciante process, ventrally joining raphe(ER) with Ad5 dorsally on Cb4.Ad 1-3 absent.Ad4 dorsally on dorsodistalmost cartilaginous endof Eb4 posteriorly, joining narrow raphe with LE4ventrally, ventrally broadly on Cb4 dorsodistally.Ad5 dorsolaterally on Cb4, there joining raphe
(ER) with OP ventrally, posteroventrally on dorso-distal end of Cb5, joining raphe with TV5.SOD present.RDs separate anteriorly, united posteriorly at ori-gin, inserting on Pb3 dorsal surface posteromedially.Additional remarks. SCL present, attached dorso-medially to cartilaginous posteroventral tip of Bb3.TV4 free from Cb5s. Pb4 and UP4 present. Pb2edentate. IAC absent. Eb4 levator process absent.PCI attaches to cleithrum by extremely long, fine ten-don. Medial end of Eb4 much larger than that of Eb3on one side, about same size on other.STEPHANOBERYCIDAEStephanoberyx monae Gill, USNM 304376, 104 mm.Plate 71
Description.LEI fine, on dorsal edge of Ebl distal to uncinateprocess.LE2 on dorsoposterior edge of Eb2 somewhat dis-tal to mid-length of Eb2.LE3 on dorsodistal edge of Eb3, inserts jointlywith OD3 insertion.LE4 on dorsodistalmost bony edge of Eb4, joinedposteroventrally by raphe with dorsolateral portion ofAd4 origin, some fibers continuing across raphe.LP slender, joining LE4 dorsoanterior to insertion,wrapping partway around LE4 and becoming slightlyseparate just ventral to origin.LI1 on Pb2 mid-dorsolateral edge ventral to ante-
rior section of TPb2.LI2 on Pb3 dorsoposteriorly somewhat lateral tomedial edge of bone; anteromedial half of insertionjoins posteromedial half of edge of TPb3 attachmentto Pb3 on one side, but is completely posterior toTPb3 insertion on the other side; posteromedial halfof insertion borders SO fibers extending dorsoanter-iorly along surface of Pb3.TD comprises TPb2, TPb3, and TEb2. TPb2 sep-arated laterally by LI1 insertion, anterior section at-taches to Pb2 anterior arm, posterior section attaches
at base of Pb2 uncinate process (on right side, fibersof posterior section continue short distance anteriorlyventral to anterior section before uniting with anteriorsection fibers). TEb2 attaches in CT enveloping dor-somedial end of Eb2-Pb2 joint. TPb3 passes laterallydorsal to OD4 origin, then extends deeply ventrallyand attaches to Pb3 dorsal surface medial surface an-terior to LI2 insertion on one side and meets antero-medial edge of LI2 insertion on the other side; ventrallayer (not illustrated) of TPb3 muscle fibers passesknifelike through OD4 and attaches to Pb3 medial toattachment of lateral TPb3 attachment.OD comprises OD3, OD3', OD4. OD3 and OD3'originate together on dorsoanterior surface of Pb3 be-ginning just posterior to cartilaginous anterior end.
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muscle forms two short divisions posteriorly; lateraldivision inserts with LE3 insertion on Eb3 dorsodis-tal edge; medial division ventral surface attaches toEb4 dorsal edge, but dorsal fibers continue withoutinterruption posteroventrally, becoming fused withposteromedial fibers of OP (we have observed asomewhat similar condition only in Hypoptychus,Hypoptychidae). OD3' branches off ventral surfaceof OD3 shortly posterior to origin and inserts on dor-sal edge of Eb3 ventral to OD3. OD4 originates ondorsal surface of Pb3 just posterior to OD3-OD3' or-igin and ventral to TPb3; on one side, muscle-fibersfrom ventral surface of TPb3 pass through OD4 an-teriorly before attaching to Pb3; OD4 muscle insertson medial edge of Eb4 bony process, which, in mostfishes, bears cartilaginous tip of uncinate process.OP originates on Eb4, fibers are continuous dor-sally with posteromedial division of OD3; muscle liesposterior to Ad4 ventrolaterally; inserts ventrally onCb4 posteriorly, joining small ER with dorsal end ofAd5.Ad 1-3 absent.Ad4 origin on Eb4 dorsolaterally, joined by raphewith LE4 insertion, with some fibers continuous be-tween the muscles.Ad5 originates dorsally from raphe (ER) joiningventral end of OP on Cb4, continuous anteriorly withventromedial OP fibers, inserting on Cb5 dorsally.SOD present.RDs originate together but separate before passinganteriorly ventral to SOD and TPb3; RD inserts onPb3 dorsoposteromedial surface of Pb3, parallelinglaterally adjacent SO longitudinal fibers, which alsoinsert on Pb3 surface beginning at posterior end ofOD4 origin.Additional remarks. SCL present, attached poster-omedianly by short ligament to posteroventral carti-laginous tip of Bb3. TV4 free from Cb5s. Pb4 re-duced. UP4 absent; tooth plate fused to Eb3 incor-rectly indicated as UP4 by Rosen (1973:472, fig. 91).Pb2 edentate. IAC absent. Eb4 levator process ab-
sent. Cartilage-tipped Eb3 and Eb4 uncinate process-es absent.
BARBOURISIIDAE
Barbourisia rufa Parr, AMNH 29772, ca. 205 mm.Plate 72
Description. Only left half of dorsal-gill arch musclesavailable for study.LEI on distal dorsal bony edge of base of Ebluncinate process.LE2 on dorsalmost bony edge of Eb2.LE3 on cartilaginous tip of Eb3 uncinate process.LE4 on Eb4 near dorsodistal end.LP on Eb4 at and posterolateral to base of LE4.
LI1 on Pb2 dorsally, anteriorly ventral to anteriorportion of TPb2.LI2 on Pb3 dorsally.TD comprises three parts: TPb2, TEb2, TPb3.TPb2 is divided into two portions; anterior portionon dorsal surface of Pb2 anteriorly, becoming atten-uated posterolaterally, broadly continuous posteriorlywith posterior portion, which attaches ventrolaterallyon Pb2 medial to LI1 insertion and is posteriorlybroadly continuous with TEb2 (see also remarks un-der SO). TEb2 attaches to Eb2 dorsally well medialto LE2 insertion and is completely separate fromTPb3. TPb3 attaches dorsoposteriorly on Pb3 at jointwith Eb3, and is continuously posteriorly with SOD.OD3-4 originates ventral to TPb2 and TEb2 onPb3 and inserts broadly on joined uncinate processesof Eb3 and Eb4.OP represented by broad muscle attaching to Eb4dorsoposteriorly, dorsolaterally partially continuouswith LE4, attaching to Cb4 at lateral end of ER, andjoining ER dorsally; broad Ad5 joins ER ventrally,possibly includes OP medially.Ad 1-3 absent.Ad4 on Eb4 dorsoposterolaterally and Cb4 dorsal-ly medial to internal angle of Eb4-Cb4 joint.Ad5 broad, joining ER dorsally and Cb5 ventro-posteriorly well medial to distal end; medial portionof muscle probably includes OP.SOD present, continuous posteriorly with SO.RDs separate, insert on dorsomedial surface ofPb3.SO branch of transverse muscle layer arises dor-somedially from SO, curves partly around medial arcof RD, and extends anteriorly, becoming continuouswith ventral surface of TPb3. Two distinct strap-likemuscles originate from longitudinal layer, one attach-es to Pb4 and the other to Pb3.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 toothed. Eb4 levatorprocess absent. IAC absent.
RONDELETIIDAE
Rondeletia loricata Abe and Hotta, USNM 206836,74.3 mm. Plate 73
Description.LEI on dorsolateral bony edge of Ebl uncinateprocess.LE2 on dorsalmost bony edge of Eb2.LE3 on tip of Eb3 uncinate process dorsal to in-sertion of Eb3 branch of OD3-4.LE4 near dorsolateral end of Eb4.LP threadlike, inserts by even finer tendon on Eb4just distal to LE4 insertion.LI 1 on anteromedial surface of Pb2 ventral to an-
94 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
terior half of TPb2 (splits TPb2 laterally into twoparts).LI2 on posterolateral surface of Pb3, mediallyabutting TPb3 attachment.TD comprises TPb2 and TPb3. TPb2 overliesOD3-4 and LI1 origins and attaches to dorsomedialsurface of Pb2 anterior to Pb2 uncinate process, isdivided mid-laterally by passage of LI1, and is con-tinuous posteriorly by a few muscle strands withTPb3. TPb3 attaches to dorsolateral surface of Pb3,posteriorly abutting medial edge of LI2 insertion, andis broadly continuous posteriorly with SOD.OD3?4 originates on Pb3 dorsoanteriorly ventralto TPb2, divides well posteriorly with short branchinserting on Eb3 uncinate process and another on Eb4uncinate process.OP dorsally on posterior surface of Eb4 uncinateprocess, in two sections; lateral section broader, ven-trally on Cb5 dorsally, joining ER with Ad5 dorsally;medial section slender, medially inseparable fromSO, ventrally on Cb5 dorsally posterior to Ad5 me-dially.Ad 1-3 absent.Ad4 dorsally on Eb4 distal cartilaginous end, ven-trally on Cb4 dorsoposterior bony surface.Ad5 on Cb4 posteriorly well medial to distal end,joining ER with OP lateral section ventrolaterally,ventrally on distal end of Cb5 dorsodistally.SOD present.RDs separate, cross (right side dorsal to left side)just anterior to origins (tendinous), insert on Pb3 pos-teromedially.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 edentate. IAC ab-sent, Eb4 levator process absent.CETOMIMIDAE
Ditropichthys storeri Goode and Bean, SIO 64-24-25, 77.7 mm. Plate 74
Description.LEI absent.Remarks. D. storeri is the only acanthomorph weexamined that lacks LEI. Another specimen shouldbe examined to verify that the absence is not an ar-tefact of a faulty dissection.LE2 on Eb2 mid-dorsally.LE3 absent.LE4 broadly, dorsally on Eb4 just dorsomedial toOD4 insertion on Eb4.LP filamentous, inserts by fine tendon on Eb4 atposterior margin of LE4 insertion.LI1 on Pb2 mid-dorsally.LI2 on Pb3 dorsal surface mid-posteriorly.TD comprises TPb2, TEb2, TEb3, and TEb4.TPb2 band-like, dorsoanteriorly on Pb2. small ven-
trolateral slip of fibers (not visible in illustration) at-taches to Pb3 lateral edge, muscle continuous pos-teriorly with TEb2. TEb2 string-like as it passes ontoand attaches dorsally on Eb2 just medial to LE2 in-sertion, continuous posteriorly by fine, spaced, di-agonal muscle threads with TEb3. TEb3 on Eb3 dor-somedially, forming raphe anteriorly with OD3 at-tachment, continuous posteriorly with TEb4. TEb4on Eb4 anteromedial to OD4 attachment, continuousposteriorly with SOD.OD3 and OD4 with joint origin on Pb3 dorsoan-teromedial surface; unified muscle divides at pointopposite mid medial edge of LI2. OD3 inserts on Eb3anteromedially, joining raphe with TEb3. OD4 in-serts on Eb4 dorsodistally at and anterior to LE4 in-sertion. OD4 lies mostly dorsal to OD3.OP dorsally on Eb4 posteriorly, anteriorly overlap-ping most of Ad4 and possibly fusing with Ad4 pos-teromedially; ventrally broadly on Cb5 posterodistal-iy-Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning medi-ally anterior to much of OP (and possibly fusing withit) and extending laterally to end of Eb4, ventrallyon Cb4 posterolaterally and CT joining distal ends ofCb4 and Cb5.Ad5 apparently absent.Remarks. It is unusual for Ad5 to be absent inacanthomorphs, and is possibly the result of the closeattachment of the posterolateral surface of Cb4 andthe anterolateral surface of Cb5, which obviates anyfunction Ad5 might serve.SOD slender, comprising loose threads of muscle.M. SO-Pb3 strap-like section of longitudinal mus-cle fibers, passes anteriorly from base of diverticulumof SO and ventral to SOD, TEb3, TEb4, and OD andattaches anterolaterally to Pb3 dorsal surface.RDs separate, vertically elliptical, insert on Pb3dorsoposteromedially.Additional remarks. SCL absent. TV4 complex,free from Cb5s. Pb4 tiny, UP4 large. Pb2 edentate.IAC absent. Eb3 and Eb4 uncinate processes absent.Eb4 levator process absent. Large, vertically oriented,bulb-like diverticulum on each side of SO just pos-terior to level of SOD; SO fibers attach to surface ofdiverticulum, which has two small glottis-like open-ings into esophagus.IcosteiformesICOSTEIDAEIcosteus aenigmaticus Lockington, LACM 35865-1,199 mm; HSU 81-305, 188 mm.Plate 75Description (based primarily on LACM specimen).LEI shortest levator, on bony dorsal edge of Ebluncinate process beginning just lateral to relativelylong cartilaginous medial end.
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LE2 on dorsoposterior edge of bony portion ofEb2 at about mid-length and well lateral to TEb2attachment.LE3 on dorsoanterior surface of Eb3 uncinate pro-cess, meeting OD3?4 insertion laterally.LE4 on bony dorsal edge of Eb4 well lateral touncinate process.LP on Eb4 beginning, variably, at lateral edge ofLE4 insertion, or slightly anteromedial to it, and ex-tending laterally to end of bony dorsal edge. Left-side LP anomalously doubled.LI1 on Pb2 dorsal bony surface just posterior toanteriormost cartilage tip and on anteromedial edgeof Pb3 adjacent to Pb2.LI2 on Pb3 dorsoposteriorly medial to medial endof Eb3.TD flat, entire dorsal surface attached to thickmuscular and CT layer that lines ventral surface ofskull, comprises TPb2, TEb2, and TEb3-Eb4. TPb2kidney-shaped, concave anteriorly, attached antero-laterally to dorsal surface of Pb2 just medial to me-dial end of IAC; anteroventral surface attached byCT (not illustrated) to dorsoanterior surface of Pb3.CT continuing and attaching to entire posterior Pblsurface; muscle fused ventromedially and posteriorlywith TEb2, which attaches to Eb2 bony surface dor-soposteriorly just lateral to medial end and well me-dial to LE2 insertion. TEb2 continuous posteriorlywith TEb3-Eb4, which attaches on Eb3 dorsomedi-
ally and Eb4 dorsomedial edge well medial to unci-nate process, there meeting SO fibers. TEb3-Eb4mid-posteriorly continuous by crossing musclestrands with SOD.OD3-4 origin on Pb3 dorsally posterior to anteriorcartilaginous tip, insertion on joined medial edges ofEb3 and Eb4 uncinate processes.OP bilaterally asymmetrical (undamaged side ofHSU 81-305 is like right side of LACM specimen,and is here considered to be the normal state). Right-side OP dorsally on Eb4 posteriorly, beginning lateralto uncinate process (and not clearly separate mediallyfrom SO fibers on Eb4) and extending laterally tobelow LE4-LP insertions, ventrally joins raphe (ER)with dorsal end of Ad5; tendon continues laterallyfrom raphe, is joined by Ad4 ventrally, and attachesto Cb4 posterodistally. Left-side OP dorsally likeright side, ventrally beginning laterally on lateral endof tendon attaching to posterolateral surface of Cb4and extending to lateral end of Cb5; muscle not dis-tinguishable from Ad5, if latter is present; Ad4 ven-trolaterally also joins lateral end of tendon.Remarks. The condition of OP on the right sideresembles the condition is some stephanoberyci-forms, e.g., Poromitra, or ophidiids, e.g., Dicrolene,particularly in that OP is either interrupted centrallyby ER or ends ventrally at ER, but in neither casedoes it attach to Cb5. For this reason, we record this
character as "OP wholly or partly on Cb4 and/orjoining ER at level of Cb4."Ad 1-3 absent, but fine GFM (not illustrated) onanterolateral edge of each arch.Ad4 dorsally on Eb4 posterolaterally, beginningnarrowly anteromedial to OP and extending to Eb4-Cb4 joint, ventrally on Cb4 dorsal surface medial toEb4-Cb4 joint, with some posterolateral fibers join-ing tendon at ventrolateral end of OP.Ad5 apparently, probably anomalously, absent onleft side, but apparently represented by musclestrands attaching ventrally to Cb5 and dorsally join-ing raphe with ventral end of OP well medial to distalends of Cb4 and Eb4. See remarks following OP.Remarks. Attachment to ER is somewhat similarto that of Dicrolene (Ophidiidae) and some stephan-oberycids.SOD broad.RDs short, joined at origin, slightly separated an-teriorly.Additional remarks. SCL present. TV4 free fromCb5s. Tiny AC at joint between Pbl and Ebl on oneside of LACM specimen, and on undamaged side ofHSU specimen. AC between Ebl and Cbl and an-other between Eb4 and Cb4 on right side of LACMspecimen; AC present between Eb4 and Cb4 on onlyone side of HSU specimen. Pb2 edentate; Pb3 bearsonly one or two fine teeth. Pb4 present. UP4 presenton only one side of each specimen, each representedby a single, fine tooth. Additionally, we examinedUSNM 49163 (ca. 200 mm), which had gill archespreserved only on one side, and in which only UP4had teeth (one) and LACM 32682-1 (1 14 mm) whichhad no pharyngobranchial teeth on either side.Interrelationships (with comments on Amarsipidaeand Stromateoidei). Aside from being assigned to theorder Perciformes in its own suborder, Icosteoidei(e.g., J. S. Nelson, 1994). or the catchall suborderPercoidei (Weitzman, 1998), the only group the Icos-teidae has been allied to is the perciform suborderStromateoidei. The basis for a stromateoid relation-ship has never been explained clearly, but appears tobe a general external similarity, sometimes referredto as the "stromateoid look" (Haedrich, 1967:44;Ahlstrom et al., 1976:290). Matarese et al. (1984:577) noted that the eggs, larvae, and early juvenilesof Icosteus superficially resemble those of stroma-teoid fishes, but added that more data were neededbefore a precise relationship could be determined. Weagree that there is a remarkable general similarity inthe appearance of Icosteus, particularly the young,and some stromateoids (compare Matarese et al.,1984:fig. 306 with Horn, 1984:fig. 333), but add thatit remains to be shown that the similarities are ho-mologs.Haedrich (1967), who reviewed the stromateoidscomprehensively (and included Icosteus among his
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non-stromateoid comparative material), did not in-clude Icosteus among possible close relatives of stro-mateoids. Nor did he mention Icosteus when he de-scribed Amarsipus (Haedrich, 1969), which he in-cluded among the stromateoids, even though it lacksthe main stromateoid synapomorphy: saccular out-growths posterior to the last gill arch.Haedrich (1969) justified the inclusion of Amar-sipus within the stromateoids by indicating a groupof characters that it shares with the other stroma-teoids, thus implicitly re-defining the Stromateoidei:perciform caudal skeleton, uniserial jaw teeth, ex-panded lacrimal, inflated and protruding top of thehead, extensive sub-dermal canal system, bonybridge over the anterior vertical canal of the ear,which Haedrich (1971) named the pons moultoni.Johnson and Fritzsche (1989:16), discredited thepons moultoni as a stromateoid synapomorphy be-cause it is widely distributed among acanthomorphs.Of the remaining characters, Icosteus clearly exhibitsonly the uniserial teeth and an inflated and protrudingtop of the head, both of which also occur variouslyamong perciforms, and a perciform caudal skeleton,which is too nebulous a character to constitute a basisfor relationships. The caudal fin of Amarsipus (Hae-drich, 1969:fig. 5) is relatively unspecialized, withautogenous: parhypural. hypurals 1?5, two epurals,and two pairs if uroneurals (the plesiomorphic actin-opterygian caudal fin differs in having three epurals).Also present are a short neural arch on PU2 and au-togenous hemal spines on PU2 and 3. The compo-sition of the fin, however, can vary considerably, de-pending on taxon, in almost every acanthomorphclade, but no percomorph has more than five hypur-
als.We are uncertain of the exact structure of the cau-dal fin of Icosteus as we only had radiographs of twospecimens to assess its composition. USNM 49163and 37327: parhypural and hypurals 1 and 2 autog-enous, hypurals 3, 4 and 5 (6 apparently not present)separate for most of their lengths, but all possiblyfused proximally to urostyle; autogenous [paired]uroneural, two epurals, long neural spine on PU2.USNM 49163: procurrent + caudal-fin rays (dorsal/ventral) 19/15, of which the central 9/9 are branched;autogenous hemal spines on PU2-6 (additionally,dorsal fin 54, of which anteriormost is a nubbin, lastray split to base; anal fin 38, of which anteriormostis a nubbin, last ray split to base; vertebrae, includingcaudal fin, 21+48 or 20+49). USNM 37327: pro-current + caudal-fin rays 17/16 (damaged), autoge-nous hemal spines on PU2-4 (additionally dorsal fin56 or 57 and anal fin 38, vertebrae 22 + 50). In anyevent, we find no basis for relating Icosteus withAmarsipus or with other stromateoids, in which thecaudal fin is often more specialized than that ofAmarsipus or Icosteus. We note with interest, how-
ever, the large number of autogenous hemal spines,3 and 5 in the two specimens of Icosteus we radio-graphed. It is most unusual for acanthomorphs tohave more than two autogenous hemal arches, Ste-phanoberyx monae (Stephanoberycidae), a notableexception, has 3.We have noted no specializations of the gill-archmuscles that might indicate a close relationship be-tween Icosteus and Amarsipus and/or the other stro-mateoids.Horn (1984) compiled meristics and other char-acters of stromateoid taxa, including Amarsipus, inan attempt to unravel their intra-subordinal relation-ships. Using Girellidae, Kyphosidae, and Scorpididae(last now generally included in the Kyphosidae),which he considered closely related to stromateoids,as outgroups. Horn provided a cladistic analysis ofthe stromateoids. for which he found only three char-acters that supported the mcnophyly of Amarsipus +stromateoids: cycloid scales (as opposed to ctenoid),lack of scales in the preopercular area (as opposed topresent), and presence of 6 hypurals (as opposed to5). While possibly valid indications of stromateoidrelations to the three outgroups, the first two char-acters are not particularly innovative and occur wide-ly among percomorphs (e.g., Icosteus lacks scales,except for imbedded prickles along the lateral line,and on the basis of scalation cannot be excluded fromrelationship with stromateoids). The hypural charac-ter is erroneous. The Girellidae and Kyphosidae (in-cluding scorpidinins) have an autogenous parhypuraland five autogenous hypurals, as does Amarsipus.Horn may have been misled by Haedrich's (1969:fig.5) recognition of 6 hypurals in Amarsipus, one ofwhich is actually the parhypural.Without further elongating this discussion, we. likeHaedrich, find no reason to ally Icosteus with thestromateoids. On the other hand, unlike Haedrich andmore recent followers, we find no reason to considerAmarsipus closely related to the stromateoids. We,thus, consider the Amarsipidae to be incertae sedisamong the percomorphs. (See also remarks followingdescription of Amarsipus, for a possible additionalcharacter of stromateoids that is not represented inAmarsipus.)Matarese et al. (1984:577) noted that the sequenceof fin formation and reduced number of pelvic-finrays in Icosteus are "blennioid" characters. All re-cently published accounts of the pelvic-fin formulaeof Icosteus variously state that it is 4, 5, or 1,4 (blen-nioids have 1,2?1,4), but we find that there is an im-bedded, greatly reduced spine closely applied to thebase of the first segmented ray and that the formulais 1,5 (pelvic fins are lost in the adult). We find noevidence of a relationship with the Blennioidei, oreven the so-called northern blennioids, Stichaeoidei,which formerly were believed to be closely related
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to the true blennioids (Springer, 1993:493). The pres-ence of IAC and Pb2 and, occasionally, UP4 in Icos-teus exclude it from inclusion among the Blennioidei,which possess none of these structures. The presenceof a basisphenoid in Icosteus (R. Fritzsche, in litt., 4Dec 2001, based on notes made by a former graduatestudent, K. Komori), two nostrils on each side of thehead, and IAC, exclude it from inclusion among theStichaeoidei, which lack a basisphenoid and IAC,and have a single nasal opening on each side of thehead.Of possible bearing on the relationships of theIcosteidae, are the presence of the accessory cartilag-es mentioned in additional remarks at the end of themuscle description of Icosteus. The distribution ofthese cartilages in acanfhomorph fishes, as far as wehave surveyed, is presented in Table 8. Icosteus hasthese cartilages variably at the joints between Ebland Cbl and between Eb4 and Cb4 (also, uniquelyin our experience, at the joint between Pbl and Ebl ).We have found these cartilages on the first or fourtharches in a variety of acanthomorphs (Table 8), buton both the first and fourth arches only in Centro-pomus (Centropomidae), Decapterus (Carangidae)and Selenotoca (Scatophagidae), besides Icosteus(Decapterus also has them on the second and thirdarches). For the most part, accessory cartilages arerestricted to percomorphs.Fin prickles. Another character that may havebearing on the interrelationships of Icosteus is thepresence of fine prickles on the lateral surfaces of therays of all fins (except for a reduced, imbedded pel-vic-fin spine and a vestigial anteriormost element inthe dorsal and anal fins, the fins comprise only seg-mented rays). Although a variety of fishes may havefin spines that are serrated or with prickles, the pres-ence of prickles on the segmented fin rays of all orsome fins is of relatively limited occurrence. Allen(2003) discusses the presence and ontogeny of theprickles in Icosteus. Among acanthomorphs. pricklesare also present in the following (1-14):1. Early stages of the three most specialized acan-thuroid families: Luvaridae, Zanclidae, and Acan-thuridae (Johnson and Washington, 1987:504); thus,are synapomorphic for these families.2. Adults of at least three tetraodontiform families(on dorsal, anal, and caudal fins), including the basalTriacanthodidae (Tydemania navigatoris Weber,USNM 30755 1 ; Parahollardia lineata) and the Bal-istidae {Sufflamen chrysopterus (Bloch and Schnei-der), USNM 224687, 353282; Balistes capriscus,USNM 240664) and Monacanthidae (Monacanthusciliatus (Mitchill), USNM 353975, 353293).3. Early juveniles (and, variously, adults) of theberyciform families Anoplogastridae, Diretmidae,Anomalopidae, and Trachichthyidae (Baldwin andJohnson, 1995; Konishi, 1999:figs. 3 & 5, provides
SEM photographs of the prickles in larval Anomal-ops and Hoplostethus), which together with the Mon-ocentridae have been hypothesized to form a mono-phyletic Trachichthyoidei (Moore, 1993), or togetherwith the Monocentridae, Holocentridae, and Beryci-dae, a monophletic Beryciformes (Johnson and Pat-terson, 1993).4. Monocentrus japonicus (Houttuyn) (Monocen-tridae; USNM 231938) has fine prickles on the seg-mented dorsal, anal, pectoral, and caudal fins (holo-centrids do not have the prickles).5. Segmented pelvic-fin rays of an unidentifiedzeiform (?Macrurocyttidae, USNM 367331).6. Copros aper (Linnaeus) (Caproidae, USNM289207), on segmented rays of all fins. Capros issometimes allied to the zeiforms (Tyler et al., 2003),but was excluded from that group by Johnson andPatterson (1993) and assigned to the Perciformes.(Not present in the other caproid genus, Antigonia).7. Four most specialized of the seven lampridi-form families (at least dorsal fin; Olney, 1984:379;Olney et al., 1993:160): Radiicephalidae, Lophotidae,Regalecidae (Regalecus), Trachipteridae (Trachipte-
rus, also at least caudal fin; USNM 175344; Zu,USNM 2721 1 1, also pectoral, pelvic, and caudal fins,and prickles on lateral line).8. Stephanoberycidae, all fins (prickles also on lat-eral line like Icosteus).9. Barbourisia (Barbourisidae), USNM 215469;
all fins like Icosteus, only segmented rays (ca. 42
vert.; 21-22 dorsal-fin rays, flexible body. No IAC).10. Chaunax (Chaunacidae, Lophiiformes), dorsaland caudal fins.11. Antennariidae. According to Pietsch and Gro-becker (1987:30 and table 1 ), spinules are present onthe body and fins of most taxa. We note that thespinules may occur on the interradial membranes aswell as the segmented rays (USNM 73115).12. Ogcocephalidae (Dibranchus atlanticus,USNM 158003, on base of caudal only; appears tobe continuation of body armature; variably presenton fins of Ogcocephalus). Not present in ceratioids.13. Centriscidae (all segmented fin-rays of Ma-crorhamphosus); however this state is not present inthe more specialized member of the family Aeoliscus(USNM 305976).14. Priacanthidae.To sum this up, fin prickles occur in at least somemembers of: Lampridiformes, Paracanthopterygii(Lophiiformes), Stephanoberyciformes, Zeiformes,Caproidae (possibly closely related to zeiforms), Be-ryciformes, Acanthuroidei, Tetraodontiformes, Cen-triscidae, and Priacanthidae.The nature and disposition of the prickles on thefin rays and the lateral line exhibit differences amongthe groups. A study of the variation might indicatevarious interrelationships among the groups. In gen-
98 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 8.?Distribution of accessory cartilages (ACs) at Eb-Cb joint of arches 1-4 in acanthomorph fishes. A few taxa that lack ACsand which are described in family accounts are included (all those not included lack ACs). * denotes taxa not mentioned in descriptiveaccounts. P = present; - = absent; PA = present or absent. Putatively closely related groups of families are boxed.
Taxa AC at joint of arch Taxa AC at joint of arch1 2 3 4 1 2 3 4Veliferidae PomacanthidaeVelifer hypsdopterus - P P Centropyge (three species)* - - PMetavelifer multiradiatus - - - Pomacanthodes semicirculatus - - PLampridae ChaetodontidaeLampris guttatus PA - - Chaetodon austriacus * - P -Trachipteridae Chaetodon melannolus * - P -Trachipterus sp. * - - - Chaetodon triafasciatus * P P -BerycidaeBeryx splendensCentroberyx affinisIcosteidaeIcosteus aenigmalicusMenidaeMene maculala PA PAP PSebastidaeSebasles prorigerScorpaenidaePontinus rathbuniNeomerinhe beanonimMoronidaeMorone americanaMorone mississippiensis
ChauliodontidaeChauliodus sloani *CentropomidaeCentropomus undecimalisLatidaeLates niloticus
PA
AmbassidaeAmbassis sp.*Ambassis buruensisTetracenlrum caudovittatusDinopercidaeDinoperca petersi*Epigonidae
PA
BelonidaeStrongylura timucu P - PA ?Tylosurus crocodilus PA - - 9ScomberesocidaeCololabis saira PA - - -ExocoetidaeExocoetus oblusiroslris PA - - -HemiramphidaeHemiramphusfar PA P - -
PA P
P
Epigonus pandionis - - PSphyraenops bairdianus * - - PHaemulidaeHaemulon (two species^ - - PPlectorhynchus pictus - - PPomadasys crocro - - PlnermiidaeInermia villata - - PLobotidaeLobotes pacificus . PCoiidaeCoius . PMullidaePseudupeneus maculatus PA -Mulloidesflavolineaus P -Parupeneus maculatus . -
Forcipigerflavissimus *Heniochus acuminatus *SymphysanodontidaeSymphysanodon berryiSymphysanodon ocloactinusSymphysanodon sp novAmmodytidaeAmmodytes dubiusNematistiidaeNemalistius pectoralis - - - PEcheneidaeEcheneis naucrates - - - PRemora remora* - - - PRachycentridaeRachycentron canadum - - - PCoryphaenidaeCoryphaena equiselis ? ? - PCoryphaena hippurus - - - -CarangidaeCarangoides crysos * - - - PDecaptenis macrosoma* P P P PDecapterus punctatus* PA PA P PSelar crumenophthalmus - - - PSeriola sp.* - - - PSelene vomer * - - - PScomberoides tol * - - - PCepolidaeAcanthocepola limbataCepola rubeseens
MonodacrylidaeMonodactylus argenteus *CichlidaeParatilapia polleni
1 1 other genera & speciesEmbiotocidaeAmphisticus argenteusCymatogaster aggregataEmbiotoca lateralisHysterocarpus traskii
PA
KuhliidaeKuhlia - PArripidaeArripis georgianus* P PTerapontidaeLeiopotherapon unicolor - PTerapon jarbua - -GirellidaeGirella tricuspidata - -KyphosidaeKyphosus seclatrox * - -ScorpididaeMicrocanthus strigatus* - -Scorpis sp. * - -
PA
P
NUMBER 11 99
Table 8.?Continued.
Taxa AC at joint of arch~ 2 3 4 Taxa AC at joint of arch~\ 2 3 4PercichthyidaeBostockia porosa *Gadopsis marmoratus *Macquaria colonorumSillaginidaeSillago sihama P -PempheridaeParapriacanthus - PPempheris - PGlaucosomatidaeGlaucosoma - PAcropomatidaeAcropoma sp.*Apogonops anomalus *Doederleinia berycoides*Synagrops bellaDinolestidaeDinolestes lewini *SphyraenidaeSphyraena barracuda
PriacanthidaeHeleropriacanlhus cruentalusDrepanidaeDrepane (two species)*EphippidaeChaetodipterus (two species)Ephippus orbis *Platax orbicularis *ScatophagidaeSelenotoca nmltifasciata *Scatophagus argus *SiganidaeSiganus (Lo) vulpinus *Siganus (S.) spinas*AcanthuridaeAcanthurus nigrofuscus
PA
PPA
PAPPP
GempylidaeNeoepinnula americana * - -Promethichthys prometheus * - -TrichiuridaeTrichiurus lepturus * - -ScombridaeEuthynnus alletleratus * P PScomber scombrus - PScomberomorus cavalla * P -Scomberomorus commersoni * P -Gasterochisma melampus * - -lstiophoridaeIsliophorus sp * P P
PA
P
PP
PA
Embiotocidae con'tPhanerodon alripes - - PPhanerodon furcatus PA PA -Rhachochilus vacca - - -Zalembius rosaceus - - PPomacentridaeAmphiprion melanopus P PA -Amphiprion allardi - - -13 other genera & 18 species - - -LabridaeAchoerodus virids PA P PBodianus mesothorax - - -Bodianus rufus P P -Cheilinus irilobatus - - -Cheilio inermis - - -Choerodon graphicus - PA -Choerodon cyanodus - - -Cleplicus parrae P P PCorisjulis P P PDecodon puellaris P PA -Halichoeres hortulanus PA - PAHalichoeres margaritaceus P P -Holog}mnosus doliatus P - -Labroides dimidialus P P PNololabrus celidotus - - -Polylepion cruenlum P - -Pseudodax moluccanus - P PPseudolabrus miles P P -Semicossyphus pulcher - - -Suezichthys aylingi - - -Symphodus roissali P - ?Tauloga onitis PA PA PATautogolabrus adspersus - - -OdacidaeOdax pullus - - -ScaridaeLeptoscarus vaigiensis - - -Nicholsina denticulata - - -Sparisoma aurofrenatum - - -
NotograptidaeNolograplus gutiatus *
AplodactylidaeCrinodus lophodon *CentrogeniidaeCentrogenys vaigiensisLeptobramidaeLeptobrama muelleri
eral, prickles on the segmented fin rays are mostcommon among deep-dwelling fishes and non-per-comorphs (acanthuroids, tetraodontiforms, and pria-canthids, not withstanding).ER, which is present in Icosteus, has a limited dis-
tribution among the other acanthomorphs (Table 9),and, with the possible exceptions of its presence inMenidae (Mene) and Centriscidae (Macrorampho-sus), ER occurs only among the more basal or non-percomorph acanthomorphs: Polymixiiformes (Poly-
100 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 9.?Distribution of certain gill-arch characters in acanthomorph fishes. Dash (-) = absent; P = present; PA = present or absent;N = no; Y = yes; na = not applicable; 2 1 = LI1 on Pb2 and IAC; 3 A = LI1 on Pb3 (Pb2 absent); 3P = LI1 on Pb3 (Pb2 not present);? = character state unknown or questionable.
Taxa
m
p UJ M r r> PJp.WWWa.p.0r.Pr 5pUr|0fl ^ in cA rA ? C 5JJJ HHS-HHHHHHHHh-f-HHHHHHHHHHHHOOOOOOSSS < Mh?W BVeliferidaeVelifer P 9 2 P P P P P N PMetavelifer P ? 2 P P P P P N P - -LamprididaeLampris 1' ? 2 P - P P P P P ? P ? -PolymixiidaePolymixia P P 2 P P P P P N P P -AphredoderidaeAphredoderus - P 2 - - - P P - - - - - - - - - - - - - - - - - - - - - - - - P - P - - - - N - - -PercopsidaePercopsis - P 2 - - - - P - - - - - - - - - P - - - - - - - - - - - - - P P - P - - - P N - - -AmblyopsidaeChologaster - P 2 - - - P P P P - P - - - P N - - -OphidiidaeDicrolene P - 2 - P - - P - - - - - - P - - - - - - - - - - - - - - - - - - P P - - - P N - P -Brotula P - 2 P P P P P P - - - P N - P -BythitidaeCalamopteryx P - 2 - P - - P P P - P - P N - P -RanicipitidaeRaniceps - P f P P P P N - P -BatrachoididaeOpsanus - - 2,3 P - P P N P P -ChaunacidaeChaunax - - 2,3 P - - P P - - - - - - - - - - - - - - - - - - - - - - - P - - P - - - - N - - -MelamphaidaePoromitra P P 2 P P P P N PScopelogadus P P 2 P P P P N PGibberichthyiidaeGibberichthys P P 2 P P I' P N P P -StephanoberycidaeStephanoberyx P P 2 P P P P P P P N P P -BarbourisiidaeBarbourisia P P 2 P P P P P N P P -RondeletiidaeRondeletia P P 2 P P P P N P P -CetomimidaeDitropichthys - P 2 P P - P P P P P N - - -IcosteidaeIcosteus P P 2,3 P P P P P N P P -OreosomatidaeAllocyttus - P 2,3 P P P P P P N P P .ParazenidaeParazen - P 3 P - P P P P P N P - -ZeniontidaeZenion - 9 2,3 P P - P - - - - - - - - - - - - - - - - - - - - - - P - - P - - - - - N P 9 -GrammicolepidaeXenolepidichthys P P 2,3 P P P P P - - - . - - - - N - - -CaproidaeCapros P - 2 P - P P P 1 23 P N P " -Antigonia P P 2 P P P P P N P -TriacanthodidaeParahollardia P P 2 P P P N P - -MenidaeMene P P 2 P P P P N P P .TrachichthyidaeHoplostethus P - 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N - P -BerycidaeBeryx P P 2 P - - P P P P P N - P .Centroberyx P P 2 P P P P P N - P -
NUMBER 1
1
Table 9.?Continued.
101
Taxa
m
v2- a a i i i i _? _L ? 2 .?> &? -? vCu -? JH XJ^3 X ^ ^ ^ J J XI C S ^ m ill c? W ?o.,bjo. uuao.a.o.ii.DD ul ? -isnn " c 5C i pj i i > i i i i i i _,. _L T X) jO JD ~- O n-tup.? wwwuoHOH<xcucucua.CLa.a.a.a.o.D-cua.cua.a.DDa.aggOQ--- -g O > U ct .nJJJ (-HH[-Hf-H[-H!-HHS-S-(-HHHHHHS-f-i-f-CJ00000222 < olh?)B WHolocentridaeHolocentrus P P 2 P - - P P P P P N - P -Sargocentron P P 2 P - - P P P P P N - P -AnomalopidaeAnomalops P P 2 P P P P N P P -Photobelpharon P P 2 P P P P P N P P -CentriscidaeMacroramphosus - P 2 P P P 123 P N P P -GasterosteidaeGasterosteus - P 2,3 P P P 123 P N P - -HypoptychidaeHypoplychus P - 2 P P P 1 23 P N P - -Aulichthys P P 2,3 P P P 123 P N P - -AulorhynchidaeAulorhynchus - P 2,3 P P P P 123 P N P - -SynbranchidaeSynbranchus - - ? P - - P - - - - - - - - P - - - - - - - - - - - - - - - - - P - - - 1 23 P N - - POphislernon PA - 7 P P P P 123 P N - - PMastacembelidaeMastacembelus - - 2,3 P P P 123 P N - - -ElassomatidaeElassoma P P 2,3 P P P 123 P N P - -MugilidaeAgonostomus P P 2 P P P P 23 - N P - PBedotiidaeBedotia P P 2 - - - P P P P P 123 - N P - PAthennidaeMenidia P P 2 - - - P P P P P 123 - N P - POdontesthes P P 2 - - - P P P P P 1 23 - N P - PAplocheilidaeRivulus P P 2 - - - - P P P P 123 - Y P - PCyprinodontidaeCyprinodon - P 3P - - - - P P P P 123 - na P - PAdrianichthyidaeXenopoecilus - P 2 - - - - P P P P 23 - N P - POryzias . P 2 - - - - P P P P 23 - N P - PBelonidaeTylosurus - P 2 - - - P P 7 P P P 23 P P P - PStrongylura - P 2 - - - - P ? P P P P 23 P P P - PScomberesocidaeCololabis - P 2 - - - - P P P P P 23 P Y P - -HemiramphidaeHemiramphus P P 2 . - - P P P P P P 23 - Y P - PExocoetidaeExocoetus P P 2 - - - P P P P P P 23 - Y P - PAcropomatidaeSynagrops P P 2 P P P P N P - -PercichthyidaeMacquaria P P 2 P P P P N P - -LeptobramidaeLeptobrama P P 2 P P N P - -LatidaeLates P P 2 P P PA P N P - -CentropomidaeCentropomus P P 2 P P P N P - -CentrarchidaeMicropterus P P 2,3 P P N P - -Enneacanthus P P 2,3 P 123 P N P - -BathyclupeidaeBalhyclupea P P 2 1 P P N - - -SymphysanodontidaeSymphysanodon P P 2 P - P N P - -
102
Table 9.?Continued.
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Taxa
x>UJ
JJJ Ht-Hf-S-HHI-HHI-HHHHHHHHHf-t-f-HS-UOOOOOSSS < m h M Hi UEpigonidae P P N P - -Epigonus I' P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - PA - - P - - - - P N P - -MoronidaeMorone 1' P 2 P - - - P - - - - - - - - - - - P - - - - - - - - - - - - - P P - - - P N P - -SerranidaeEpinephelus P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - 1' - - - - P N - - -Anthias P P 2 P P P 1' P N - - -LutjanidaePrislipomoides P P 2 P P P P - - - - P N P - -Hoplopagrus P P 2 P P P P P N P - -HaemulidaePomadasys P P 2' P P P P N PHaemulon P P 2' P P P P N PPleclorhinchus I' P 2' P - - - - - - - - - - - - - - - P - - - - - - - - - - P - - P - - - - - N - - PInermiidaeInermia P P 2 P - - - - - - - - - - - - - - - P - - - - - - - - - - P - - P - - - - - N - - PApogonidaeClossamia P P 2 P P P P P N P - PCheilodipterus P P 2 P P P P P N P - -PriacanthidaeHeteropriacanthus I' P 2 P P P NOstracoberycidaeOstracoberyx P P 2 P P - - P - - - - - - - - - - - - - - - - - - - - - - - - - P - - - - P N P - -CirrhitidaeParracirrhiles I5 P 2 P - - - P - - - - - - P - - - - - - - - - - - - - - - - - - P - - - - P N P - -Cirrhitus P P 2 P - - - P - - - - - - P - - - - - - - - - - - - - - - - - - P - - - - P N P - -PempheridaePempheris P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -Parapriacanlhus P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -GlaucosomatidaeGlaucosoma P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -LactariidaeLaclarius P lJ 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -LateolabracidaeLateolabrax P P 2' P - - - - - - - - - - - - - - - - - - - - - - - - - - P - - P - - - - P N P - -Sciaenidae - - -Cynoscion P P 2'.3 P P - - - - - P - - - - P N - - -PolynemidaePolydactylus I' P 2 P - - - P - - - - - - - - - - - P - - - - - - - - - - - - - I' - - - - P N - - -Filimanus P P 2,3 P - - - P - - - - - - - - P - - - - - - - - - - - - - P - - - - P N - - -SillaginidaeSillago P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - P P P - - P - - 23 P N I' - PMullidaePseudupeneus P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - P - - P - - - - P N - - -Parupeneus P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - P - - P - - - - P N - - -CentrogeniidaeCentrogenys P P 2,3 P - - - - - - - - - - P - - P - - - - - - - - - - - - - - - P - - - - P Y P - -AmbassidaeAmbassis P P 2,3 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - - N P - -Tetracenlrum P P 2,3 P - - - P - - - - - - - - - - - - P - - - - - - - - - - - - P - - - - - N P - PCaristiidaeCaristius P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -Bramidae - - -Brama - P 2' P - - - P - - - - - - - - - P - - - - - - - - - - - - - - - P - - - - P N P - -ToxotidaeToxotes P P 2 P - - - P - - - - - - - - - - - P - - - - - - - - - - - - - P - - - - P N P - PPlesiopidaeAssessor P P 2,3 P - - - P - - - - - - - P - - - - - - - - - - - - - .- .- - - I' - - - - P N P - PParaplesiops P P 2,3 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - I'
NUMBER II
Table 9.?Continued.
103
Taxa
m
? 03 M 0-0r> UJ0r WUJPJDr r O.0;.DDUJ tV ,3. IN rA rA ? C 3too,:- wwujWa.o-cua.o.a.cua.OHDHa.cua.a?a,a,a,a,cuDDa.aaQQCJ_;-_; -a O>U0^J3JJJ hhhhHHHHHHhhhhhHhhhhhhhHHUOOOOOSSS < tflholU WPercidaePerca P P 2,3 P P P P P N P - -Percina P P 2,3 P I' P P P - - 1 23 P N P - -CepolidaeAcanthocepola P P 2,3 P - - - - - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N 1' - -Cepola P P 2,3 P - - - - - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -CallanthiidaeCallanthias allporti P P 2 P - - - - - - - - - - - - - - - P - - - - - - - - - - P - - P - - - 1 23 P N P - -Callanthias austral P P 2 P - - - - - - - - - - - - - - P - - - - - - - - - - - P - - P - - - 1 23 P N P - -Grammalonolus P I' 2 P - - - - - - - - - - - - - - P - - - - - - - - - - - - - - P - - 1 23 P N - - -Gerreidae - - -Genes P P 2 P - - - - P - - - - - - - - - - - PA - - P - - - 1 23 P N P - PEucinostomus P P 2 P - - - - P - - - - - - - - - - - - - A - - P - - - 1 23 P N P - PGrammatidaeGramma P P 2,3 P - - - - - - - - - - - - - - P - - - - - - - - - - - - - - P - - - - P Y P - POpistognathidaeLonchopislhus P P 2,3 P - - - - - - - - - - - - P - - - - - - - - - - - - - - - - I' - - - - P Y P - POpistognalhus 1' P 2,3 P P P P Y P - PPseudochromidaeLabracinus I' P 2,3 P - - - - P - - - - - - P - - - - - - - - - - - - P - - - - P - - - 1 23 P Y P - PPseudochromis P P 2,3 P - - - - P - - - - - - P - - - - - - - - - - - - P - - - - P - - - 1 23 P Y P - PLeiognathidaeGazza P P 2 P - P - - - P P P Y P - PLeiognathus P P 2 P - 1' - - - P P P Y P - PSecutor P 1' 2 P - P P P P Y P - PPolycentridaeAfronandus PA P 2,3 P - - - - - - - - - - - - 1' - - - - - - - - - - - - - - - - P - - - - - N P - -Monocirrhus - P 2,3 P - - - - - - - - - - - - P - - - - - - - - - - - - - P - - P - - - - - N P - -Polycenlropsis PA P 2,3 P - - P - - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - - N P - -Polycentrus - P 2,3 P - - P P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - - N P - -SphyraenidaeSphyraena P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N - - -KurtidaeKurtus P P 2,3 P P - - P - - - - - - - - - - - - - - - - - - - - - - - - - P - - - - P N P - -AmmodytidaeAmmodyles P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -TrachinidaeTrachinus P P 2,3 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -UranoscopidaeKathetostoma - P 3A P - - - - - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - - N - - -Xenocephalus - P 2,3 P P - - - - - - - - - - - - - - - - - P - - - - - - - - - - P - - - - P N - - -CheimarrichthyidaeCheimarrichthys P P 2,3 P P P P N P - -ScorpaenidaePonlinus P P 2 P P P P P N P - -Neomerinthe P P 2,3 P P P P P N P - -SebastidaeSebastes P P 2 P P P P P N P - -PlatycephalidaePlatycephalus P P 2,3 P P P P N P - -Inegocia - P 2,3 P P P P N P - -ChampsodontidaeChampsodon P P 3 P P - - P P P N P - -HexagrammidaeHexagratnmos P P 2,3 P P P P N P - -AnoplopomatidaeAnoplopoma P P 2,3 P P P P P N P - -RhamphocottidaeRamphocottus - P 2,3 P P P P P N P - -
104
Table 9.?Continued.
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Taxa
m
rj *J n f n t
,-. CU J= JO .D XI ?
J J _1 H H H H f- -Hf-Hh-Hf-f-f-f-l-f-l-f-f-HHUOOOOOSSS < (*> H 00 W 2CottidaeMyoxocephalus P P 3 P P I' P P N P - -Cottus - P 3A P P P P P N P - -NematistiidaeNematistius P P 2 P P P P P N P - -CarangidaeScomberoides P P 2 P P P P P P N P - -Selar P P 2 P P P P P P N P - PRachycentridaeRachcentron P P 2 P P P P N P - .CoryphaenidaeCoryphaena P P 2 P P - P P P P N - - -EcheneidaeEcheneis naucrates P P 2,3 P P P P P N P . -Remora P P 2,3 P P P P N P . -PomatomidaePomatomus P P 2 P P P P P P N P . -ScombrolabracidaeScombrolabrax P P 2 P P P P P N - . .ScombridaeScomber P - 2 P P P P P N P - .NemipteridaeNemipterus P P 2 P P P P P P P P N P - -CentracanthidaeSpicara P P 2 P P P P P P P 123 P N P - PSparidaeAcanlhopagrus P P 2 P - - - P - - - - - - - - P - - - - - - - - P - P - - P - P - 1 23 P N P - PLagodon P P 2 P - - - P - - - - - - - - - - - P - - - - - - - - P - P - - P - P - 1 23 P N P - PSarpa P P 2 P - - - P - - - - - - - - - - - P - - - - - - - - - - P - - P - P - 1 23 P N P - PLethrinidaeLethrinus P P 2 P - - - P - - - - - - - - - P - - - - - - - - - - P - P - - P - - - 123 P N - - -Gymnocranius P P 2 P - - - P - - - - - - - - P - - - - - - - - P - P - - P - - - 123 P N - - PMonotaxis P P 2 P - - - P - - - - - - - - - - - P - - - - - - - - - - P - - P - - - 123 P N - - PGirellidaeGirella specimen A P P 2 P P P P P N P - -Girella specimen B P P 2 P - - - P - - - - - - - - - - - - - P - - - - - - - - - - - P - - - - P N P - -KuhliidaeKuhlia P P 21 P - - - P - - - - - - - - - - - P - - - - - - - - - - - - - P - - - - P N P - -TerapontidaeLeiopotherapon P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - P - - P - - - - P N P - -Terapon P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -CichlidaeCaquetaia P P 2,3 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 123 - Y P - PAstronotus P P 2,3 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PCichla P P 3 P P - - - P P - - - - - P - - - - - - - - - - - - - P - - - - P - - - 123 - Y - - PCichlasoma P P 2,3 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PCopadichromis P P 2,3 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 123 - Y P - PCrenicichla P P 2,3 P - - - P P - - - - - P - - - - - - - - - - - - - P - - - - P - - - 1 23 - Y - - PCyrtocara P P 2,3 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PSantanoperca P P 2 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PParatilapia P P 2,3 P - - - P P - - - - - P - - - - - - - - - - - - - P - - - - P - 23 - Y P - PPtychochromoides P P 2 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - 2 3 - Y P - PPtychochromis P P 2,3 P - - - P P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 23 - Y P - PPomacentridaeDischistodus P P 2 P - - - P P - - - - - P - - - - - - - - - - - - - P - P - - P - - - 1 23 - Y P - PAbudefduf PP PP 2 P - - - P P - - - - - P - - - - - - - - - - - - - - - P - - P - - - 1 23 - Y P - PChromis 2 P - - - P P - - - - - P - - - - - - - - - - - - - P - P - - P - - - 1 23 - Y P - PAcanthochromis P P 2 P - - - P P - - - - - P - - - - - - - - - - - - - P - P - - P - - - 1 23 - Y P - PAmblyglyphidodon P P 2 P - - - P P - - - - - P - - - - - - - - - - - - - P - P - - P - - - 1 23 - Y P - PAmphiprion P P 2 P - - - P P - - - - - P - - - - - - - - - - - - - P - P - - P - - - 1 23 - Y P - PChrysiptera P P 2 P - - - P P - - - - - P - - - - - - - - - - - - - P - P - - P - - - 123 - Y P - P
NUMBER 11
Table 9.?Continued.
105
Taxa
PL)js ^f n Tt n ^^ cu ?"= J= -o *> fS1? ?~? tUWWtuW rs ""> "* ?">
c w M or ^ > uJor uJurlujp-a.Pr PrDDwW -. rs A A ^ c gJJ-J HHHHHh-HHi-HHt-(-HHHHHf-(-!-l-HHHOOOOOOS2S < b! h ifl 3 EDascyllus P p 2 P - - - P P 1 23 - Y P - PLepidozygus P p 2 P - - - P p P 123 - Y P - PMecaenichlhys P p 2 P - - - P p P P P P 123 . Y P - PMicrospalhodon P p 2 P - - - p|p - - - - - P - - - - - - - - - - - - - P - P - - P - 123 - Y P - PPlectroglyphidodonT1 p 2 P - - - p p P P P 1 23 - Y P - PPomacenlrus p p 2 P - - - p p P P P P 1 23 - Y P - PStegastes p p 2 P - - - p p P P P 123 - Y 1' - PEmbiotocidaeAmphistichus p p 2,3 P P P P 123 P Y P - PEmbiotoca p p 2 P P P P 1 2 3 . Y P - PCymatogaster p p 2 P - P P P 1 2 3 - Y P - PHysterocarpus p p 2 P 1 23 - Y P - PPhanerodon p p 2 P P P 123 P Y P - PRhacochilus p p 2 P P P 123 - Y P - PZalembius p p 2 P 123 - Y P - PLabridaeAchoerodus p p 3P 123 - Y P _ PBodianus p p 3P PA P 123 . Y P - PCheilinus p p 3P P p P P P 1 23 - Y P - PCheilio p p 3P - - - - p p P P P 1 23 - Y P . PChoerodon p p 3" - - - - p p P P P 1 23 - Y P _ PCleplicus p p 3P P - - - p p P P P 1 23 - Y P - PCoris p p 3P P - - - p p P P 1 23 - Y P - PDecodon p p 3P P P 1 23 - Y P - PHalichoeres horl. p p 3P P - - - p p P P P 123 - Y P - PHalichoeres marga. p p 3P - - - - p p P P P P 1 23 - Y P - PHologymnosus p p 3 P P - - - p p P P P 1 23 - Y P - PLabroides p p 3 P P P P P P P - P 1 23 - Y P - PNotolabnis p p 3 P P - - p p - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PPolylepion p p 3' P - - p p - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PPseudodax p p 3 P 9 - - - p 7 P P P 123 - Y P - PPseudolabrus p p 3 P P - - - - - P - - - - - - - P - - - - - - - - - - P - - - - P - - - 123 - Y P - PSemicossyphus p p 3' - - - - - - P - - - - - - - P - - - - - - - - - - P - - - - P - - - 123 - Y P - PSuezichthys p p 3P - - - - p P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 123 - Y P - PSymphodus p p 3 P - - - - - - - P - - - - - - P - - - - - - - - - - P - - P - - P - - 1 23 - Y P - PTautoga p p 3 P - - - - p P P P P 123 - Y P - PTautogolabrus p p 3 P P - - - p P - - - - - - - - P - - - - - - - - - - P - - - - P - - - 1 23 - Y P - PXiphocheilus p p 3? - - - - p P P P P 123 - Y P - POdacidaeOdax pullus p p 3P P - - - p - - - - - - - - - P - - - - - - - - - - P - - P - - - - - 1 23 - Y P - PScaridaeLeptoscarus p p 3P - - - - p ? P P P 123 - Y P - PNicholsina p p 3P - - - - p 7 P P P 123 - Y P - PSparisoma p p 3P - - - - p 7 - - - - - - - - P - - - - - - - - - - P - - P - - - - - 123 - Y P - PPholidichthyidaePholidichthys p p 3P - - - - p - - - P P - - - - P - - - - - - - - - - P - - - - P - - - 123 - Y P - PLuvaridaeLuvarus p p 2 P - - P - - - - - - - P - - - - - - - - - - - - - - P - - - - - - - - P N P - -EphippidaeChaetodipterus p p 2 P - P P P P N P - PZanclidaeZanclus p p 2 P - P P P N P - -AcanthuridaeAcanthurus p p 2 P - - P - - - - - - - - - - - - - - - - - - - - - - P - - - - - - - - P N P - -NandidaeNandus p p 2 P - - - - - - - - - - - - P - - - - - - - - - - - - - - - - P - - - 123 - N P - PBadidaeBadis p p 2 P - - - - - - - - - - - - P - - - - - - - - - - - - - - - P - - - 123 - N P - PPristolepidaePristolepis p p 2 P - - P - - - - - - - - - - - - - - - - - - - - - - - - - - P - - - - - N 1' - -
106
Table 9.?Continued.
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Taxa
'3-X>uj2 * m t A "JnT J3 J2 ?> JO rH
?H' "5 <NCNCNmfnmrr?3-Tr-4'3-!?"A' rn''Et UJ M ^ m Cj CJj)a. J= J2JOJ2jDjDjOjDjOJ3jOjD&;e;-Om i i i C? W ra a/ a/> W^wuJPJo-a.a^Dr DwW ,~ <n rn rn <""> g 3en _ X!j=xijoxij2j2^xixj=j2j=j=xijDj=X!X5X)j3joj3?-&-J=S2S35 D- D- cu v> 9S-1 , ??JJJ HHHHH(-t-[-HHHHHr-HHf-HHHHHHHHU000005SS < 00 f- c/5 W UJChannidaeChanna asiatica P P 3P P - - - - - - - - - - - - - P - - - - - - - - - - P - - - - P - - - - - N P - PC. harcourtbutleri P P 3 P P - - - - - - - - - - - - - - - - - - - P - - - - P - - - - P - - - - - N P - PAnabantidaeCtenopoma P P 3 P - - - - P - - - - - - P - - - - - - - - - - - - - P - - - - P - - - 3 - N P - PSandelia P P 3 P - - - - P - - - - - - P - - - - - - - - - - - - - P - - - - P - - - 23 - N P - PAmarsipidaeAmarsipus P P 2 P - - - P - - - - - - - - - - P - - - - - - - - - - - P - - P - - - - P N P - -CentrolophidaePsenopsis P P 2 P - - - P - - - - - - - - - - - - P - - - - - - - - - - - - P - - - - - N I' - -BathymasteridaeBathymaster P P 2,3 P - - - - - - - - - - P - - - - - - - - - - - - - - - - - - P - - - - P N P - -ZaproridaeZaprora P P 2,3 P - - - P - - - - - - - P - - - - - - - - - - - - - P - - - - - - - - P N P - -StichaeidaeUlvaria P P 2,3 P - - - - - - - - - - - P - - - - - - - - - - - - - - P - - P - - - 1 23 P N P - -BovichtidaeBoviclus P P 2 P - - - - - - - - - - - - - - P - - - - - - - - - - - P - - P - - - 123 P N P - -PseudaphritidaePseudaphritis P P 2 P - - - - - - - - - - - - - - - - - - - - P - - - - - - - - P - - - - P N P - -DactylopteridaeDactyloptena P P 3 P P - - - P - - - - - - - P - - - - - - - - - - - - - - - - - P - - - - P N P - -MalacanthidaeCaulolalilus P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - P P - P PA N P - -Malacanthus P P 2 P - - - P - - - - - - - P - - - - - - - - - - - - P P PA P P N P - -DraconettidaeDraconetta P P 3 P P P - - > - - - - - - - - - - - - - - - - - - - - P - - - - P - - - 123 P N P - -CallionymidaeCallionymus - P 3 P P - - P P - - - - - - - - - - - - - - - - - - - - P - - P - - - - - - PA N P - -GobiesocidaeTrachelochismus - P 3A P P P P N - - -TripterygiidaeRuanoho P P 3A P P P 123 - Y P - -Lepidoblennius P P 3A P - - - - - - - - - - P - - - - - - - - - - - - - - - - - - P - - - 123 - Y P - -BlenniidaeParablennius gatto. P P 3A P - - - - - - - - - - P - - - - - - - - - - - - - - - - - - P - - - 123 P Y P - -Parablennius tasm. P P 3A P - - - - - - - - - - P - - - - - - - - - - - - - - - - - - P - - - 1 23 P Y P - -Scartella P P 3A P - - - - - - - - - - - - - P - - - - - - - - - - - - - - - P - - - 1 23 9 Y P - -Scartichthys P P 3A P P P 1 23 P Y P - -Istiblennius P P 3A P - - - - - - - - - - P - - - - - - - - - - - - - - - - - - P - - - 123 P Y P - -DactyloscopidaeDactylagnus - P 3A P P P P Y P - -ClinidaeGibbonsia P P 3A P P P P Y P - -Heterostichus P P 3A P P P P Y P - -Heteroclinus P P 3A P P P P Y P - -Springeratus P P 3 A P P P Y P - -Ophiclinus P P 3A P P P P Y P - -Clinus P P 3A P P P Y P - -LabrisomidaeCalliclinus P P 3A P P P Y P - -Labrisomus P P 3A P P P Y P - -ChaenopsidaeNeoclinus P P 3A P P P P Y P - -RhyacichthyidaeRhyacichthys P P 23 P P P 23 P N P - POdontobutidaeOdontobulis P P 23 P P - P P P P P - P - - - 123 P N P - PMicropercops P P 23 P 7 P P P P 123 P N P - PPercottus P P 23 P P P P P P 123 P N P - PXenisihmidaeXenisthmus P P 2 P P P 3 P N - - -
NUMBER I 1
Table 9.?Continued.
107
Taxa
?3-&? 2 222 ?
p W B D.Q- > UJOr WWWDr a.a.a.DDW UJ ?. <N cA rA <N c Srn J^J3?j3J3B^iiJlJ?XlJ3ilXlJ3XlJ3JlJ3j:jl5;S;JlSRAAA <" Q ? -1 , *ffl c -_ WWU)WQ.o.a.CLa.aHa.DHO.a?o.a.a.CL-CL.cu&.o.cuDDa.QDQQQ--_; -a o > U d jsJJJ HHHHF-S-HHHHHHHHl-HHHf-HHH!-HHUOOOOOS22 < wHtnUWEleotridaeEleotris P P 23 P P 1' P 1 23 P N P - POphiocara P P 23 P P - - - - - - - - - - - - - - P - - 123 P N P - PMicrodesmidaePtereleotris P P 23 P P P P 123 P N P - PNemateleotris P P 23 P P P P 1 2 3 P N P - PMicrodesmns P P 2 P P 1 2 3 P N P - -GobiidaeGlossogobius P P 2 P - - - - - - - - - - P - - - - - - - - - - - - - - P - P - - - - - 123 1' N P - PABollmannia P P 2 P - - - - - - - - - - - - - P - - - - - - - - - - - P - P - - - - - 123 P N P - -Padogobius P P 23 P - - - - - - - - - - - - - - - - - - P - - - - - - - - - - P - - - 123 P N I' - PPseudapocryptes P P 2 P P P - P - P 1 23 - Y P - -Gnatholepis P P 23 P - - - - - - - - - - P - - - - - - - - - - - - - - P - P - - - - - 123 P N P - PTrypauchen P P 23 P - - P - - - - - - - - - - - - - - - P - - - - - - P - P - - - - - 123 P N P - -PsettodidaePsettodes - P 2 P - - P P P P N P - -
mixiidae); Paracanthopterygii: Ophidiiformes (Ophi-diidae, Bythitidae), Gadiformes (Ranicipitidae),Batrachoidiformes (Batrachoididae); Stephanoberyci-formes (Stephanoberycidae, Gibberichthyidae, Ron-deletiidae, Barbourisidae, Melamphaidae); Zeiformes(Oreosomatidae): Beryciformes (Berycidae. Anom-alopidae, Holocentridae. Trachichthyidae).Among these groups, Icosteus, superficially, re-sembles the stephanoberyciform and paracanthopter-ygian families (but not Stephanoberycidae), withwhich it variously shares to the exclusion of the othergroups (data augmented from the literature): flexiblebody, a high number of precaudal (20-23 in Icosteus)and total vertebrae (67-72 in Icosteus), a high num-ber of dorsal-fin rays (52-58 in Icosteus), attaining alength of over a meter (ca. 2 m in Icosteus). It alsoshares with these families and some cetomimid ste-phanoberyciforms, imbedded lateral-line scales andweakly ossified bones. It further resembles the ce-tomimids in lacking pelvic fins (at least as an adult),having a highly distensible stomach (also present inlophiiforms), and dorsal and anal fins consisting onlyof segmented rays and placed far posteriorly. It dif-fers, at least, from all other basal acanthomoiphs inhaving only five, as opposed to six, hypurals, and,except for some ophidiiforms and gadiforms, in hav-ing an interarcual cartilage.The many similarities between Icosteus and someStephanoberyciformes, particularly Barbourisidae,mainly favors close relationship with that group;however, the presence of IAC tends to favor the Par-acanthopterygii. We assign Icosteidae to its own or-der, Icosteiformes, which we tentatively place nearthe Stephanoberyciformes, but, in any event, believe
that it occupies a pre-percomorph position among theacanthomoiphs.
Zeiformes and Possible Relatives
Nelson (1994:290-291) discussed the vacillatingassignments of the inter-relationships of the Capro-idae through 1993: whether they are zeiforms, per-ciforms, a sister-group of the zeoids and tetraodon-tiforms, or "in some position between the Stephan-oberyciformes and Beryciformes." Nelson rejectedJohnson and Patterson's (1993) treatment of caproidsas perciforms and retained them as the sister groupof the zeoids (Nelson, 1994:253).Tyler et al. (2003), conducted a phylogenetic studyof the zeiforms that also included a primitive tetrao-dontiform (Parahollardia), both genera of caproids,and seven other diverse outgroups. Relationship ofzeiforms with tetraodontiforms and caproids was sup-ported in three of their four analyses (data orderedwith and without most meristic characters and dataunordered with most meristic characters). The rela-tionship was not supported in the fourth analysis(data unordered without most meristic characters),which they considered, "the most rational and bestjustified."We elect to position the caproids together with thezeoids and tetraodontiforms in accordance with threeof the four cladistic analyses of Tyler et al. (2003).The presence of ER in an acanthomorph is usuallyan indication of a pre-percomorph condition. OnlyMene, the centriscid Macroramphosus, and the enig-matic Icosteus (Icosteidae) among those taxa usuallyincluded in the percomorphs have it, and we present
108 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
evidence that Icosteus (q.v.) is more appropriatelyplaced near the stephanoberyciforms. We also believethat Mene, which is generally unusual morphologi-
cally, is probably more appropriately placed amongthe pre-percomorphs, and have arbitrarily inserted itnear the zeiforms. Although we believe that the cen-triscids and relatives are also probably more closelyrelated to pre-percomorphs than to percomorphs, weplace them with the other groups Johnson and Pat-terson (1993) included in their Smegmamorpha to fa-
cilitate discussion of that group.
ZeiformesOREOSOMATIDAE
Allocyttus verrucosus (Gilchrist), USNM 329365,82.8 mm. Plate 76
Description.Remarks. The musculature appears to be skewedatypically, but the muscles and their attachments oth-erwise appear normal. The muscle fibers are gener-
ally stringy and weak, particularly the posterior mus-
cles, and are infiltrated with filmy tissue that wasimpossible to remove without causing considerabledamage to the muscles. An anomalous muscle strapon the right side attaches to Ebl mid-posteriorly,passes deeply ventrally, and attaches to Eb2 antero-medialmost edge.LEI very broadly on Ebl dorsoposteriorly. CTcovering levators laterally indents anteriorly and cov-ers much of Ebl posterior surface, becoming contin-uous with tendinous dorsal portion of LEI ventral toorigin.LE2 on Eb2 dorsoposteriorly, beginning just pos-terolateral to lateral end of TEb2.LE3 absent.LE4 dorsally on bony Eb4 process posteroventralto uncinate process, muscle joined posterolaterally byLP.LP very fine, continuous with LE4 ventrolaterallyslightly dorsal to combined insertion on Eb4.LI1 anterior surface attaches to Pb2 posteriorly innotch between two dorsally extending cartilage-tipped processes, continues ventrally and inserts onPb3 dorsally ventrolateral to dorsally extending car-tilage-tipped process.LI2 on Pb3 dorsolateral^ ventrolateral to medialend of Eb3.TD comprises TPb2, TEb2, TEb3, and TPb3. TPb2very thick pad with mid-anterior U-shaped notch ex-tending more than half distance across pad and shal-low mid-posterior notch; notches joined by mid-lon-gitudinal raphe, which gives rise to CT sheets dor-
sally; raphe attaches ventrally to CT of pharyngealroof; muscle attaches anteriorly to cartilage-tipped
Pb3 dorsal and Pb2 dorsomedial processes, fusesventrally with TEb2, and is continuous posteroven-trally by muscle strand with TEb3. TEb2 attaches onEb2 dorsally just anteromedial to LE2 insertion.TEb3 attaches to posterior margin of Eb3 medial touncinate process. A few, probably anomalous, musclefibers branch off right-side TEb3 and insert on Eb4dorsally. TPb3 is a very thin ribbon of muscle (notillustrated) that extends ventroanteriorly from TEb3,where TEb3 passes dorsal to dorsoposteromedialedge of Pb3. The ribbon extends deeply ventral toOD and attaches to Pb3 dorsally in a deep depressionwell posterior to the anterior end of Pb3.OD3, OD3' originate inseparably on Pb3 dorsallyventral to TEb2. OD3 inserts on Eb3 uncinate pro-cess dorsoanteriorly. OD3' branches off ventrallywell lateral to origin and inserts on Eb3 dorsally an-teroventral to uncinate process.OD4 absent.OP attaching dorsally on Eb4 posteriorly begin-ning medially ventral to uncinate process and ex-tending well laterally, meeting Ad4 dorsally. A fewmedialmost fibers (not visible in illustration) extendventrally to Cb5 anterior to Ad5; most fibers attachventrally on Cb4 joining ER with Ad5, medial todistal end of Cb4, and posterior to Ad4 (ER not inview in Plate 76B).Adl-3 absent.Ad4 dorsally on Eb4 mostly anterior to OP later-
ally, extending to near cartilaginous end of Eb4, ven-trally on Cb4 dorsally beginning short distance me-dial to Eb4-Cb5 joint and extending to joint.Ad5 dorsally on Cb4 well medial to distal end,ventrally on distal portion of Cb5 dorsally.SOD absent.RDs possibly anomalous, each divided longitudi-nally into two straps; right-side medial strap passesdorsal to left-side strap and lies appressed to right-side lateral strap; left-side medial strap passes me-dially ventral to right-side medial strap and lies ap-pressed to medial surface of right-side lateral strap.Additional remarks. SCL attached mid-dorsally tocartilaginous posterior tip of Bb3. TV4 free fromCb5s. Pb4 and UP4 absent. Pb2 toothed. IAC absent.Eb 1 uncinate process absent. Eb4 levator process ab-sent. PARAZENIDAEParazen pacificus Kamohara, USNM 364277, 73.0mm; USNM 187807, 93.9 mm.Not illustratedZENIONTIDAE
?
= Zenion hololepis (Goode and Bean), USNM187864, 68.5 mm.Not illustrated
NUMBER 11 109
Description.LEI on Ebl mid-dorsally (uncinate process ab-sent), attached anteriorly for entire muscular lengthto perpendicular section of pharyngeal roof CT, towhich Pbl also attaches for entire length.LE2 on Eb2 mid-dorsally.LE3 absent on one side, much reduced, but presenton other. ? Absent on both sides.LE4 broad based, on Eb4 mid-dorsoposteriorly.LP very fine, on Eb4 at ventrolateral edge of LE4insertion.
?
Specimen damaged, unable to determineif LP was present.LI1 on Pb3 anteriorly near base of dorsoanterior-most process; about twice size of LI2. ? Almost en-tirely on Pb3, but very fine CT attachment to Pb2present.LI2 on Pb3 dorsolaterally anterolateral to medialend of Eb3.TD comprises TPb2, TEb3, and TPb3. TPb2 athick pad with broad, deep V-shaped notch (open an-teriorly) leading to short mid-longitudinal raphe,which attaches dorsally to CT sheet covering musclesand ventrally to pharyngeal roof CT; muscle attachesto Pb2 and Pb3 dorsalmost surfaces posteriorly withtendinous continuation to medial edge of Pbl. TPb3a strap of muscle extending anterolaterally ventral toOD3-4 and attaching to Pb3 along medial edge ofLI2; muscle continuous mid-posteriorly with TEb3.TEb3 attaches broadly to Eb3 posterior edge, meetingOD3' posteriorly.
?
Comprises TPb2, TEb2, andTEb3. TPb2 notched anteriorly and posteriorly, fusedventrally with TEb2. which attaches on Eb2 dorsoan-teriorly ventral to LE2 insertion.OD comprises OD3-4 and OD3', muscles origi-nate on Pb3 dorsally, OD3' separating ventrally justlateral to origin; OD3-4 inserts on tip of Eb3 ante-
riorly, passing dorsal to tip and inserting on medialedge of Eb4; OD3' inserts on Eb3 dorsally ventral toOD3-4.OP dorsally broadly on Eb4 posteriorly beginningnear medial end and extending laterally to below lat-eral edge of LE4 insertion; ventrally. narrowly onCb5 beginning dorsally on most proximal surface oflong, rod-like cartilaginous distal end and extendingshort distance medially, meeting Ad5 medially andTV5 laterally. ? Possibly absent, but if present, at-taches ventrally to Cb4 (see Ad4).M. SO-Pb3 pair of SO longitudinal muscle straps,one on each side of each RD. extending well ante-riorly and inserting on medial edge of Pb3 ventral toOD3-4. ? Absent.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly, beginning me-dially at lateral edge of OP and extending laterally toEb4-Cb4 joint, ventrally broadly on Cb4 medial tojoint with Eb4. ? Two layered; anterior layer typical;posterior layer, separates dorsally from anterior layer
and fuses anteroventrally with anterior layer; poste-rior layer possibly represents modified OP.Ad5 triangular, apex dorsally even though muscleattaches broadly dorsally on posterior bony surfaceof Cb4 beginning slightly medial to cartilaginous dis-tal end, attaches ventrally along long cartilaginousdistal end of Cb5 and extends onto bony surface shortdistance anterior to OP.
?
Cb5 cartilaginous distalend normal (short rounded cap), hence, Ad5 muchless extensive ventrally.SOD absent.RDs separated by space more than 1.5X diameterof one RD.Additional remarks. SCL attached mid-dorsally tocartilaginous ventroposterior end of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 absent.Pb2 toothed. Eb4levator process absent.GRAMMICOLEPIDAEXenolepidichthys dalgleishi Gilchrist, USNM341954. 102 mm; USNM 320015, 100 mm.Plate 77
Description.LEI extremely broadly on Ebl dorsolaterally (un-cinate process absent).LE2 on dorsally expanded posterior margin ofEb2.LE3 on Eb3 lateral to tip of uncinate process.LE4 on Eb4 levator process dorsoanteriorly.LP tendinously joining LE4 slightly dorsal to car-tilage tip of Eb4 levator process.LI1 tendinously on Pb2, tendon continuing ontoPb3.LI2 on Pb3 laterally ventral to LI1.TD comprises TPb2, TEb2. and TEb3. TPb2 a bi-lateral pair of thick, rope-like muscles, separated byshallow notch mid-anteriorly and joined along theirmedial edges to tough, deeply depressed CT sheet(conforms dorsally with parasphenoid keel); CT sheetis tightly applied posteroventrally to TEb2 dorsally.TPb2 attaches tightly anterolaterally to cartilaginousdorsal tip of Pb2 and weakly to adjacent cartilaginousdorsal tip of Pb3. TEb2 divided medianly by narrowraphe (obscured from view in illustration); TEb2 lat-erally on Eb2 dorsally anterior to LE2 insertion.TPb2 and TEb2 not continuous posteriorly withTEb3. TEb3 on almost entire posterior edge of Eb3medial to uncinate process, also attached mid-ven-trally by CT to CT of pharyngeal roof.OD3, OD3' originate together on Pb3 dorsallyventral to TEb2 and divide immediately after exitingfrom under TEb2, with a slender dorsal branch(OD3) attaching narrowly to Eb3 uncinate processanteriorly and a slightly larger ventral branch (OD3')attaching on Eb3 dorsally ventral to uncinate process.OD4 absent.
110 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONOP on Eb4 dorsoposteriorly beginning ventral touncinate process and extending medially almost tomedial end, laterally overlapping Ad4 medially; ven-trally on Cb4 dorsally beginning well medial to distalend and extending a short distance medially, overlap-ping Ad4 medially and difficult to separate fromAd4. ER, to which OP usually attaches along withAd5 when OP attaches to Cb4, apparently absent.Ad 1-3 absent.Remarks. As SO longitudinal fibers extend anteriorto the gill arches, they spread laterally and anteriorlyand form a complex mesh over the roof of the oralchamber. From this mesh, a sheet of fibers (not illus-trated) extends ventrolaterally toward each of the firstthree arches and either attaches directly or throughconnective tissue to the ventral edge of the epibran-
chial, beginning about mid-laterally, and continuinglaterally and ventrally and attaching extensively tothe ceratobranchial dorsal surface medial to the innerangle of the Eb-Cb joint. The appearance in each caseis superficially like an Ad, but acanthomorph Adsl-3 are on the external surface of the associated Eb-Cbpair.Ad4 attaches dorsally along much of ventral edgeof Eb4 beginning medially anterior to OP and ex-tending laterally almost to distal end of Eb4; ven-trally on much of dorsal edge of Cb4 anterior to OP.Ad5 reduced or absent. On one side of each spec-imen, a short muscle (almost vestigial in smallerspecimen) joins Cb5 dorsodistally to Cb4 well ven-tral to its distal end.SOD absent.RDs adjacent.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 absent. Pb2 toothed IAC absent.Ebl uncinate process absent. Eb4 levator processpresent. CaproiformesCAPROIDAECapros aper (Linnaeus), USNM 289207, 2 speci-mens, 53.2-62.5 mm; USNM 327294, 64.8 mm.Plate 78
Description.LEI on dorsolateralmost bony edge of Ebl, welllateral to long uncinate process, which is attacheddorsoposteriorly by fine ligament to Pb2 anterolat-erally (no IAC). Thick, tendinous edge of thin CTsheet (not illustrated) covering LEs laterally, attachesalong entire posterolateral margin of LEI.LE2 on dorsolateralmost bony edge of Eb2, ven-troanteriorly meeting Ad2 dorsally and ventromedi-ally meeting TEb2 laterally.LE3 on Eb3 uncinate process anteriorly, ventro-medial edge meeting OD3 laterally.
LE4 on Eb4 dorsolateralmost bony edge well lat-eral to uncinate process, joining raphe posteroven-trally with Ad4 dorsally (unclearly or only partiallyjoining raphe with Ad4 in smaller specimen).LP absent.LI1 in depression in cartilaginous dorsalmost sur-face of Pb2 just posterior to anterior end.LI2 on Pb3 dorsoanterolaterally just ventral tooverlying anteromedial edge of Eb3, insertion ante-roventrally proximate to bony posterior surface ofPb2.TD comprises TEb2 and TEb3-Eb4. TEb2 withmid-longitudinal raphe, central dorsal area withtough CT pad (not illustrated) with raised convex lat-eral margins, which attach to expanded dorsoanteriorcartilaginous caps of Pb2s; pad attached mid-ven-trally to mid-longitudinal raphe; muscle laterally cov-ers much of dorsal surface of Eb2 and attaches dor-sally at ventromedial edge of LE2 and ventrally todorsomedial edge of Ad2, variously joining rapheswith each of these two muscles. TEb3-Eb4 well sep-arated posteriorly from TEb2, attaches broadly onposterolateral edge and surface of Eb3 (ventral toOD3 and OD3'), and posteromedial edge of Eb4;muscle slightly overlaps and is continuous ventrallyby fine muscle strands with SOD.OD3. OD3' origin on Pb2 posteromedially ventralto TEb2, continuing posteriorly broadly on Pb3 dor-somedially. OD3 inserting broadly on dorsoanterioredge of Eb3 uncinate process (there meeting ventro-medial edge of LE3). OD3' separating from OD3well ventrolateral to origin and inserting on dorsoan-terior surface of Eb3, meeting dorsomedial edge ofAd3.OD4 absent.OP on Eb4 dorsoposteriorly beginning mediallynear SOD and extending laterally to near uncinateprocess, dorsolateral^ meeting TEb3-Eb4 and dor-somedial end of Ad4, ventrally on Cb5 posteriorlyjust medial to distal end.M. SO-Pb3 (not illustrated), band of longitudinalSO fibers on each side extends anteriorly ventral toTD and inserts on Pb3 medially.Ad 1 broadly on anteroventral edge of Eb 1 and dor-soanteromedial surface of Cbl.Ad2 on dorsoanterolateralmost surface of Eb2,meeting LE2 ventrally and TEb2 laterally, and ondorsoanterior surface of Cb2.Ad3 on Eb3 anterolaterally ventral to uncinate pro-cess, meeting OD3' dorsomedially and on Cb3 dor-soanteriorly.Ad4 on Eb4 dorsoposteriorly beginning at lateralend of OP near uncinate process and extending lat-erally to end of Eb4; ventrally broadly on Cb4 lat-erally medial to Eb4-Cb4 joint, there meeting Ad5dorsoanterolaterally.Ad5 short, dorsally on Cb4 dorsodistally and ven-
NUMBER 11 111
trally on Cb5 dorsodistally. meeting OP ventrolater-
ally.SOD present.RDs separating slightly as they pass below SOD.Additional remarks. SCL attached mid-dorsally toelongate cartilaginous ventroposterior tip of Bb3.TV4 free from Cb5s. IAC absent. Pb4 absent, UP4present. Pb2 toothed. Eb4 levator process absent.
Antigonia rubescens (Gunther)?, USNM 365941,76.7 mm. Not illustrated
Additional material:
?
= Antigonia capros LoweUSNM 163521, 78.1 mm; ? = A. combatia Berryand Rathjen. USNM 188045, 91.8 mm.
Description.LEI finely, tendinously on dorsoposterior edge ofEbl just lateral to uncinate process.LE2 finely, tendinously on dorsoposterior edge ofEb2 much nearer lateral than medial end.LE3 finely, tendinously on dorsal tip of Eb3 un-cinate process.LE4 narrowly on dorsodistal end of Eb4, joinedslightly dorsal to insertion by LP.LP very fine, joined by long slender tendon to ven-trolateral edge of LE4 dorsal to joint insertion on Eb4dorsodistally.LI1 finely, tendinously on Pb2 posteroventrally.LI2 tendinously on Pb3 ventrolaterally just medialto cartilaginous edge articulating with medial end ofEb3.TD comprises TEb2 and TPb3-Eb3-Eb4. Verythick CT pad mid-dorsally extends anteriorly and at-taches to Pb2s dorsoanteriorly (all but obscuring theirdorsal cartilaginous ends); few small ACs irregularlydistributed dorsally on pad (all three species); padattaches mid-ventroanteriorly to CT of pharyngealroof. TEb2 continuous from one side to the otheronly around CT pad posteriorly; muscle otherwiseattaching medially to lateral surface of CT pad andextending laterally to, or almost to, dorsodistal endof bony surface of Eb2 (well lateral to LE2 insertion).TPb3-Eb3-Eb4 transversely continuous, free fromTEb2; on Pb3 dorsal surface posterolaterally ventralto OD3-OD3'. continuing onto Eb3 dorsomediallyand well along posterior edges of both Eb3 and Eb4;free from SOD.
? ?
TEb2 attaches to CT pad ven-trolaterally so that dorsolateral edge of pad overlapsTEb2.OD3, OD3' origin on Pb3 dorsoposteriorly, ex-tending laterally and branching ventrally (OD3') atabout level of medial end of Eb3; OD3' inserting onEb3 dorsally ventrolateral to uncinate process; OD3inserting on Eb3 uncinate process anteromedially.(Note: Eb3 and Eb4 uncinate processes closely bound
together. Superficially, OD3 also appears to insert ondorsomedialmost edge of Eb4 uncinate process, butprocesses can be readily separated and OD3 clearlyinserts only on Eb3.)OD4 absent.OP dorsally on Eb4 posteriorly beginning near me-dial end and extending laterally to below uncinateprocess, there meeting TPb3-Eb3-Eb4 ventrolateral-ly; ventrally on Cb5 posterolaterally, just meetingAd5 ventromedially.M. SO-Pb3, short band of longitudinal SO fiberson each side extends anteriorly ventral to TD andinserts on Pb3 medially.AD1?3 absent, reduced muscle strands (GFM1-3)on anterolateral surfaces of respective Eb and Cb.Ad4 dorsally on Eb4 posteriorly, beginning me-dially at lateral edge of OP and extending laterally todistal end of Eb4; ventrally on Cb4 dorsally extend-ing laterally to inner angle of Eb4-Cb4 joint and an-terior to Ad5 attachment.Ad5 relatively small, dorsally on cartilaginous dis-tal end of Cb4 posteriorly, beginning at Eb4-Cb4joint and extending ventrally to posterolateral surfaceof cartilaginous distal end of Cb5.SOD variously slender to moderately broad.RDs adjacent for much of length, narrowing, sep-arating, and becoming tendinous anteriorly and in-serting on Pb3 posteriorly.Additional remarks. SCL attached mid-posteriorlyto greatly elongated cartilaginous ventroposterior endof Bb3. TV4 free from Cb5s. IAC reduced, ball-likesuspended in ligamentous tissue (Tyler et al., 2003:table 1. indicate that IAC is absent in A. capros, butwe found it in all three species of Antigonia we ex-amined). Pb4 absent. UP4 present. Pb2 toothed. Eb4levator process absent.
TetraodontiformesTRIACANTHODIDAE
Parahollardia lineata (Longley), USNM 287252,119 mm. Plate 79
Description.LEI narrowly, tendinously on Ebl dorsopostero-laterally.LE2 narrowly, tendinously on Eb2 dorsopostero-laterally.LE3 narrowly, on tip of Eb3 uncinate process.LE4 musculously and tendinously on Eb4 dorso-lateral^.LP very fine, tendinously joining LE4 posteroven-trally just dorsal to LE4 insertion; origin also tendi-nous.Remarks. LP in tetraodontiforms (and many otherfishes) is much reduced, fragile, obscured by various
112 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
tissues, and easily destroyed when peeling away sur-rounding tissues. Because of this, and because he wasunaware of the existence of LP in fishes at that time,Winterbottom (1974a; in litt. 19 Mar 2001), in hisstudy of tetraodontiform musculature, did not reportthe presence of LP in any tetraodontiform. In addi-tion to Parahollardia, we found LP in Hollardia hol-lardi Poey (USNM 289328, Triacanthodidae) andTrixiphichthys weberi (Chaudhuri) (USNM 280329,Triacanthidae). Additionally, in Balistes vetula Lin-naeus (USNM 349662, Balistidae). we found a long,slender muscle extending dorsoposteriorly from a ra-phe with LE4 somewhat dorsal to the LE4 insertion(origin of muscle not determined). This muscle mayrepresent a modified LP, which usually joins LE4near the LE4 insertion. Investigation of LP in tetrao-dontiforms may provide information bearing on theinternal classification of the order.LI1 narrowly tendinously on dorsomedialmost tipof Pb2.LI2 by long, slender tendon on dorsal surface ofcartilaginous Pb3 process that is joined laterally bymedial end of Eb3.TD complex, comprising TPb2, TEb2, and TPb3-Eb3. TPb2 a pair of laterally curving, verticallyraised muscles, lying dorsal to and mostly free frombroad CT mid-section of TEb2; anteriorly, each TPb2muscle joins CT of pharyngeal roof, which gives risedorsally to thick CT shell that lies over TPb2s (andTEb2 portions underlying TPb2s) and forms cup at-taching to ventral process on skull; posteroventrally,each TPb2 joins CT, which attaches to TEb2; CT alsojoined by (forms tendinous mid-anterior portion of)TEb2 (mid-posteriorly, CT of TEb2 narrows consid-erably); muscle does not attach to Pb2. TEb2 mas-sive, attaching to entire bony dorsal surface of Eb2;anomalous dorsoanteromedial branch of right-sideTEb2 attaches finely, tendinously to anterior end ofright-side TPb2. TPb3-Eb3 on Pb3 dorsally medialto medial end of Eb2 (Eb3 portion with mid-longi-tudinal raphe joining TPb2 and TEb2 mid-ventrally),continuing onto Eb3 dorsolaterally ventral to OD3and meeting OD3 insertion on uncinate process, con-tinuous ventrally by diagonal strand of muscle withSOD.Remarks. Because of the large thick central CTsection of TEb2, we treated Pb3 as dorsally mostlynot covered by muscle. This area of the Pb3s formsa diarthrosis with a knob-like process on the ventralsurface of he skull, which is very similar to that ofpomacentrids, but not like that of labrids, cichlids, orembiotocids. The process has an irregular surface inParahollardia and pomacentrids. In the other threegroups, the process is divided into a bilateral pair ofbroad, smooth surfaces. Stiassny and Jensen (1987:282) characterized all the "labroids" as having a sim-ilar knob-like process, but we disagree.
OD3 origin on Pb3 dorsally ventral to TEb2, in-sertion massively on anterior surface of Eb3 uncinateprocess.OD4 absent.OP dorsally broadly on Eb4 posteriorly beginningon or near uncinate process and extending mediallyalmost to end of Eb4; ventrally broadly on Cb5 pos-teriorly, attachment tendinous laterally and joined onone side by Ad5 posterolateral^, but not on other.GFM1?3 large fan of fine filamentous muscle fi-bers attach on anterior surfaces of joined distal endsof respective Eb and Cb.Ad4 broadly dorsally on Eb4 posteriorly beginning
at or near uncinate process and extending laterallyalmost to end of Eb4; ventrally more restricted thandorsally, on Cb4 dorsally medial to Eb4-Cb4 joint.Ad5 dorsally on distal end of Cb4 posteriorly, ven-trally on Cb5 dorsodistally.SOD present, slender, entirely ventral to TPb3-Eb3.RDs slightly separated.Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Small, spherical IAC present. Pb4 absent,UP4 present. Pb2 toothed. Eb4 levator process ab-sent.Winterbottom (1974a) described and illustrated thebranchial arch musculature of several tetraodonti-form s. MeniformesMENIDAEMene maculata (Bloch and Schneider), USNM347107, 97.7 mm, USNM 102498, 87.8 mm.Plate 80
Description.LEI on broad anterior surface of Ebl lateral to tipof uncinate process.LE2 on mid-dorsoposterior edge of Eb2.LE3 tendinously on tip of Eb3 uncinate process,interrupted dorsally by CT and becoming musculusagain before attaching to skull.LE4 fan-like, joins LP ventrally on common ten-don, which inserts on Eb4 lateral to uncinate process.LE4 continuous dorsally with CT from which variousthin muscle straps continue dorsally and attach toskull.LP continuous ventrally on common tendon withLE4, dorsally joining extensive CT sheet, which dor-sally becomes musculus; dorsal extent of musculusLP not clearly separated from PP; CT sheet continuesventrally and attaches along outer curvature of 4thand 5th arches.LI1 on Pb2 tendinously just posterior to joint withIAC.LI2 on Pb3 dorsoposterolaterally.
NUMBER 11 113TD comprises TEb2 and TPb3-Eb3. TEb2 veryshort transversely, very long longitudinally, in twocontinuous sections: anterior section much the broad-
er, with mid-longitudinal raphe giving rise to CTsheets that attach to cranium; anteroventrally muscleattaches by CT to anteriormost ends of Pb2s, mid-ventrolaterally attaches to IAC2 dorsoposteriorly, andposterolaterally attaches to dorsoanteromedial surfaceof Eb2 anterior to medial edge of LE2 insertion; pos-terior section attaches to posteromedial edge of Eb2.TPb3-Eb3 begins anteriorly as diagonal strap of mus-cle on each side, which attaches to Pb3 dorsally ven-tral to posterior margin of TEb2; straps cross poste-
riorly just as they mesh with anterior end of broadEb3 portion of muscle, which attaches to medial edg-es of Eb3s.Remarks. The diagonal muscle straps appear to de-rive from crossing muscle strands present in manyacanthomorph TDs, and which connect the posteriorend of one TD muscle with the anterior end of an-other TD muscle or SOD. The straps in Mene are sodistinctive, that they possibly are not homologouswith TPb3-Eb3 in other acanthomorphs.OD3, OD3' large, columnar, originating on dor-soanteriormost end of Pb3 ventral to TEb2, dividingposteriorly as it exits from below TEb2, with ventralbranch, OD3', inserting on Eb3 dorsal surface ven-troanterior to insertion of OD3, which is on anteriorsurface of Eb3 uncinate process.OD4 absent.OP complex, distinctly three-parted dorsally onEb4, less so ventrally: medial part smallest, strap-like, beginning well medial to medial end of Eb4,curving ventrolaterally around OP middle part andinserting on tendinous raphe (ER, not illustrated) atabout level of Cb4; middle part originating on Eb4just ventral to tip of uncinate process, extending ven-trally, and joining ER immediately lateral to medialOP part; lateral OP part broadest, extends laterallyfrom middle part on Eb4 to cartilaginous distal endof Eb4 and onto AC4; sub-dorsally, medial edge oflateral OP part becomes tendinous and extends tonear distal end of Cb5; tendinous edge mid-posteri-orly joins ER and continues ventrally as lateral edgeof OP muscle attaching to Cb5. Ad5 joins tendinousOP edge ventroposteriorly.RecD2 slender, originating on Eb2 anteromedialedge and inserting on Ebl ventral surface just ventralto LEI, overlies CT to which gill rakers attach ven-trally.RecD3 well developed, originating on Eb3 anter-oventrolaterally and inserting on Eb2 just ventral toLE2, overlies CT to which till rakers attach ventrally.Ad 1-3 absent.Ad4 narrow, dorsally on ventrolateral surface ofEb4 fusing anteriorly with lateral OP part ventrally
(not as apparent in posterior view as presented inPlate 80C).Ad5 (see also OP) attaches ventrally on dorsodistaledge of Cb5, dorsally it attaches musculously andtendinously to posterodistal surfaces of Cb4 and Ad4.SOD absent.RDs proximate.Additional remarks. SCL weakly attached to ven-trally extending cartilaginous posterior tip of Bb3.TV4 free from Cb5s. Two interarcual cartilages: an-terior IAC joins Ebl uncinate process and tiny car-tilaginous process dorsally slightly posterior to an-terior bony tip of Pb2; posterior IAC (IAC2) joinscartilaginous process near posterior end of Pb2 (pro-cess is subtended by posterior extension of Pb2 tooth-plate; neither Pb2 process is visible in Plate 80C(IAC2 unknown for any other actinopterygian taxon).AC cartilage between distal ends of Eb3 and Cb3,another between distal ends of Eb4 and Cb4. Pb4absent, UP4 present. Pb2 toothed. Eb4 levator pro-cess absent.Leis (1994:140-142) compared characters (noneincluding the dorsal gill arches) of Lactariidae, Men-idae, and Carangidae and suggested the possibilitythat the first two families are the second and first
sister groups of the third. In the discussion followingGasteromorpha, below, we suggest reasons for con-sidering Menidae as being more closely related topre-percomorphs than to percomorphs (such as Lac-tariidae and Carangidae).
BeryciformesTRACHICHTHYIDAE
Hoplostethus mediterraneus Cuvier, USNM 307273,79.1 mm; USNM 361959, 98.0 mm.Plate 81
Description (based almost entirely on the smallerspecimen).Remarks. LE1-LE4 becoming long tendons dor-sally and coalescing into single origin on skull. LI1-LI2 musculously attached to skull.LEI on dorsal edge of Ebl well lateral to tip ofuncinate process.LE2 on posterodorsal edge of Eb2 at mid-lengthof Eb2.LE3 on cartilaginous tip of Eb3 uncinate process
at and dorsal to OD3-4 insertion on Eb3.LE4 on posterolateralmost surface of Eb4.LP absent.LI1 on medial surface of Pb2 uncinate process.LI2 on dorsal surface of Pb3 mid-laterally.TD comprises TPb2', TPb2, TEb2, and TPb3-Eb3.TPb2' thin, roughly triangular, sheetlike, dividedmid-longitudinally, closely applied to ventral surfaceof thin CT sheet, which attaches to ventral surface of
114 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
skull and envelops anterior ends of Pb2s. Mid-ven-troanteriorly, TPb2' muscle fibers mesh with semi-circular, pad-like TPb2, which attaches to Pb2 anter-odorsally and is anteroventrally continuous withTEb2. TEb2 inserts on Eb2 dorsal surface medial toLE2 insertion. Anteroventrally, TEb2 meshes withTPb3-Eb3. TPb3-Eb3 forms raphe dorsolaterallywith OD3-4 origin (forming "bun-like" mid-dorsalsection), sheet of fibers continue deep ventrolaterallyand attach broadly on dorsal surface of Pb3 medialto LI2 insertion, with small muscle strap attaching todorsomedial surface of Eb3. TPb3-Eb3 is continuousposteriorly by fine muscle strand with SOD.Remarks. TD anomalous anteriorly in larger spec-imen, with separate muscle below TPb2' that attachesto Pb2 on one side and Pb3 on other; TPb2' presentonly unilaterally.OD3-4 massive, originates anteriorly from Pb3anterior end (ventral to TEb2) and dorsoposteriorlyat raphe with TPb3-Eb3, inserts on lateralmost dor-soanterior surface of Eb3 uncinate process and later-almost dorsoposterior surface of Eb4, forms rapheposteroventrally with dorsal end of OP.OP originates dorsally from raphe with OD3-4 onEb4 (raphe occluded from view in Plate 8 IB), lat-erally continuous with Ad4, joins raphe (ER) ven-trally with Ad5.Ad 1-3 absent.Ad4 dorsally on posterolateral surface of Eb4, con-tinuous medially with OP, forms raphe posteroven-trally with Ad5 (raphe?as ER?continues mediallyat ventral end of OP), attaches along dorsal surfaceof Cb4 anteroventrally anterior to Eb4-Cb4 joint.Ad5 mid-ventrally broadly on Cb5 distally, ex-panded well ventrally beyond attachment to Cb5 withtendinous attachment to cleithrum; dorsolaterally onCb4 distally, expands medially and joins acutely an-gled (two-sided) raphe dorsally: with Ad4 laterallyand OP medially.SOD present.RDs adjacent, each inserts by short tendon on pos-terior end of Pb3 (in larger specimen tendinous in-sertion is elongate, extending well external to SO.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 absent. Pb3 toothed. IAC absent.Eb4 levator process absent.BERYCIDAEBeryx splendens Lowe, USNM 306134, 3 specimens,98.0-111 mm. Plate 82
Description.Remarks. LEI, 2, and 4 are tendinous (not illus-trated) along lateral edge of their insertions. LEI,may form a slender, separate muscle (section) origi-nating as a long tendon, becoming musculous ven-
trally and closely paralleling the remainder of LEI,and inserting as a long tendon at and lateral to theremainder of the LEI insertion. Because of the var-iability of LEI, we do not name the separate section.LEI origin tendinous, insertion on Ebl uncinateprocess dorsoanteriorly lateral to cartilaginous tip.LE2 on expanded bony edge of Eb2 dorsoanter-iorly.LE3 mostly on cartilaginous tip of Eb3 uncinateprocess, but few muscle fibers continue and insert ontightly abutting Eb4 uncinate process.LE4 on dorsodistalmost cartilaginous edge of Eb4.LP very slender, becoming tendinous basally andjoining lateral edge of LE4 insertion.LI1 insertion begins dorsally on medialmost edgeof Pb2 uncinate process just ventral to cartilaginoustip (joined there by dorsolateral edge of TPb2), andextends anteroventrally along posterior surface of un-cinate process. About same size as LI2.LI2 on Pb3 dorsolaterally near joint with Eb3 me-dialmost end.TD comprises TPb2, TEb2-Ebl. TEb2, TPb3-Eb3,and TEb4. TPb2 arises as broad strap from medianraphe on dorsoanterior surface of TEb2 and attachesto anterior surface of Pb2 uncinate process just ven-tral to tip of process; joined by LI1 anterior edgealong Pb2 uncinate process. TEb2-Ebl arises as nar-row strap from median raphe that is continuous withTEb2 median raphe, curves anterolaterally. becomingdorsal to TEb2, and attenuating as short, slender ten-don that inserts in CT surrounding tips of Ebl andPb2 uncinate processes; continuous posteriorly by di-agonal muscle strand with TPb3-Eb3. TEb2 broad,attaches to dorsal surface of proximal bony half ofEb2, with median raphe, which gives rise dorsally totough CT pad. TPb3-Eb3 on Pb3 bony dorsal surfacemedial to LI2 insertion and on Eb3 posteromedial-most surface, continuous dorsomedianly with TEb4,which is on anteromedial dorsal surface of Eb4.Remarks. Sasaki (1989:14) first reported TEb2-Ebl in Beryx, although he did not assign a name toit, nor illustrate it. We note that there is a remarkable,probably superficial, resemblance of TEb2-Ebl to theanterior branch of the TD plexus of Acanthurus(q.v.).OD3-4 originates broadly along most of length ofbony dorsal surface of Pb3, inserts broadly on ante-
rior surface of Eb3 uncinate process and medial edgeof Eb4 uncinate process.OP broad dorsally, on Eb4 posterior surface medialto dorsal attachment of Ad4, dorsolaterally joiningraphe with OD3-4, and ventrally joining ER withdorsal edge of Ad5 on Cb4, with a few medial mus-cle strands continuing onto Cb5.Remarks. Raphe with OD3-4 not present on eitherside of one specimen.Ad 1-3 absent.
NUMBER 1 1 115
Ad4 dorsally on Eb4 posterolateral surface andventrally on Cb4 anterior to Eb4-Cb4 joint.Ad5 on Cb4 posterodistally and Cb5 dorsodistally,forming raphe (ER) along dorsal edge with OP, andcontinuous ventrolaterally with SO.SOD present.RDs varying from well separated, to adjacent, tooverlapping, each becoming broad tendinous strapbefore inserting on Pb3.Additional remarks. SCL absent. TV4 free fromCb5s. Tiny AC 4 on dorsoposterior cartilaginous endof Cb4 (additional smaller AC4, dorsal to larger oneon one side of each of two specimens). UP4 present.Pb4 absent (see below). Eb4 levator process absent.Medial end of Eb4 much larger than medial end ofEb3. IAC absent. Pbl bony with cartilage ends. Pb2toothed.Rosen (1973:466, fig. 84) indicated the presenceof a reduced Pb4 in B. splendens. This element wasnot present in any of our three specimens (nor in theone of Centroberyx). Our specimens of Bery.x splen-dens were collected in the Gulf of Aden, off Somalia,whereas Rosen's specimen was collected in the Gulfof Mexico, off Mississippi. It is, therefore, quite pos-sible that there is variation in the presence of Pb4 inBery.x and that more than one species is involved; arevision of the genus is warranted. We did not at-tempt to verify Rosen's finding and for the purposesof our study, we have treated berycids as lackingPb4.
Centroberyx affinis (Gunther), USNM 176984, 93.5mm. Plate 83
Description.LEI originating as long tendon (other LEs mayhave tendinous component); muscle slender, on dor-solateral edge of medially projecting Ebl uncinateprocess.LE2 on tip of expanded bony edge of Eb2.LE3 on anterior surface of cartilaginous tip of Eb3uncinate process.LE4 on dorsodistalmost edge of Eb4.LP very fine, tendinously inserted at ventrodistal-most edge of LE4 insertion.LI1 on medial surface of Pb2 uncinate process.LI2 tendinously on Pb3 dorsolateral^ ; about samesize as LI1.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2roughly oblong with deep notch mid-anteriorly andextensive CT pad arising dorsally from broad mid-longitudinal raphe, which completely divides TPb2(and TEb2); muscle attachment begins anteriorly onanterior tip of Pb2 and extends posteriorly to pointon anterior edge of Pb2 uncinate process; muscle is
free anterolaterally and dorsal to anteromedial por-tion of TEb2, but is posteromedially and ventrallycontinuous with TEb2. TEb2 deeply notched mid-posteriorly at continuation of mid-longitudinal raphe;another, incomplete raphe mid-dorsally amidst TEb2,which extends laterally onto dorsal surface of Eb2 topoint just medial to base of spade-like process. TPb3-Eb3 free from TPb2 and TEb2, attaches laterally todorsal surface of Pb3 medial to LI2 insertion andattachment continues posteriorly, passing dorsal toanteromedial end of Eb4, onto posteromedialmostedge of Eb3. TPb3-Eb3 is dorsoposteriorly continu-ous by fine muscle strand with SOD.Remarks. The attachment of TPb3-Eb3 to Eb3 isimmediately adjacent to the abutting edges of Eb3and Eb4. We expect that other specimens might ex-hibit attachment to Eb4 as well. A well-developedTEb4 is present in the related Beryx splendens.OD3-4 massive, originates broadly along Pb3 ven-tral to TPb2 and TEb2, and inserts on anterior surfaceof Eb3 uncinate process and tip of Eb4 uncinate pro-cess, forming raphe posterodistally with dorsal endof OP on Eb4.OP broad dorsally, on Eb4 posterior surface medialto dorsal attachment of Ad4, dorsolaterally formingraphe with OD3-4, and ventrally joining ER withdorsal edge of Ad5 on Cb4, with a few medial mus-cle strands continuing onto Cb5.Ad 1-3 absent.Ad4 dorsally on posterodistal end of Eb4, attachedcontinuously on posterior edge of Eb4-Cb4 joint, andcontinuing medially short distance on Eb4.Ad5 dorsally on posterodistal surface of Cb4, ven-trally on distal end of Cb5.SOD very fine.RDs adjacent.Additional remarks. SCL absent. TV4 free fromCb5s. Tiny AC4 (not visible in dorsal view) on dor-soposterior edge of distal end of Cb4. UP4 present,Pb4 absent (see discussion in Additional remarks un-der Beryx). IAC absent. Medial end of Eb4 muchlarger than medial end of Eb3. Pb2 toothed.Eb4 levator process absent; however, it appears thenarrow cartilaginous distal end of Eb4, which con-nects expanded dorsal and ventral cartilaginous bulg-
es, could become ossified in larger specimens, thusisolating a levator process.
HOLOCENTRIDAE
Holocentrus adscencionis (Osbeck), USNM 345408,
1 1 1 mm. Plate 84
Additional material. Sargocentron diadema (Lace-pede), USNM 334012, 95.8 mm.
116 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Description.Remarks. The muscles of the two taxa are essen-tially the same.LEI broadly on anterolateral surface of expandedbony edge of Eb 1
.
LE2 on expanded dorsoposterior margin of Eb2.LE3 on joined cartilaginous tip of Eb3 uncinateprocess and cartilaginous edge of Eb4 uncinate pro-cess, also joins raphe with OD3-4 on these processes(joined processes completely enveloped in muscle,not visible externally).LE4 massive, dorsolaterally on Eb4 bony surface.LP on Eb4 at and lateral to LE4 insertion; insertioncontinuous ventrolaterally with CT sheet attaching tofourth and fifth arches.LI1 on Pb2 posteroventral to uncinate process.LI2 on Pb3 dorsoposterolaterally, just anterior tomedial end of Eb3, ventromedially joining raphe withlateral edge of TPb3.TD comprises TPb2, TEb2, TPb3, and TEb4. TPb2on Pb2 posteromedial surface, roughly circular andpad-like, dorsally with slight ventrolateral extensionon each side, notched anteriorly and giving rise toCT sheet dorsally from between notch, overlies andis broadly continuous ventrally with TEb2. TEb2dorsal to OD3-4 origin, attaching laterally on Eb2anterior to LE2 insertion. TPb3 broad, not connectedto TPb2 or TEb2, ventral to OD3-4 origin, on Pb3beginning on medial edge of base of Pb3 uncinateprocess, and extending posteriorly along ventrome-dial edge of LI2 insertion to position medial to me-dial end of Eb3, continuous by diagonal musclestrand with TEb4. TEb4 on dorsomedial surface ofEb4, continuous posteriorly by diagonal muscle bandwith SOD.OD3-4 origin on Pb3 dorsomedial edge ventral toTEb2, insertion broadly on anterior surface of Eb3uncinate process and posterior surface of Eb4 unci-nate process, insertion joining raphe with LE3, whichis on both processes (Eb3 uncinate process has well-developed rounded cartilaginous tip; Eb4 uncinateprocess is horizontal with narrow cartilaginous edge).OP dorsally on Eb4 posteriorly and medial to Ad4attachment, joining raphe with OD3-4 (on Eb4) pos-teroventrolaterally, ventrally joining ER with Ad5dorsally on Cb4.Adl-3 absent.Ad4 dorsally on Eb4 lateral to OP, ventrally onCb4 at and medial to Eb4-Cb4 joint.Ad5 ventrally extensively on dorsal surface ofCb5, dorsally on Cb4 well medial to distal end, therejoining raphe (ER) with OP ventrally.SOD present.RDs adjacent.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Eb4 levator process ab-sent. Pb2 toothed.
ANOMALOPIDAEAnomalops katoptron (Bleeker), SIO-75-466, notmeasured. Plate 85
Additional material.
?
= Photoblepharon palpebra-rum (Boddaert), USNM 264123, 71.2 mm.
Description.LEI origin by long tendon, insertion on dorsoan-terior surface of Eblwell lateral to tip of uncinateprocess.
? On Ebl immediately lateral to tip of un-cinate process.Remarks. All other levators originate musculously.Right-side uncinate process is incompletely separatedfrom medial end of Eb 1
.
LE2 on dorsoanterior surface of posterodorsalbony process of Eb2.LE3 on anterior surface of Eb3 uncinate process.LE4 on dorsodistalmost edge of Eb4, continuousat ventrolateralmost edge of insertion with weak mus-cle strands, representing LP, which is continuous pos-teriorly with CT sheet, which changes to muscle, rep-resenting PP; LE4 ventrolaterally joins raphe withAd4 dorsolaterally.LP represented by weak muscle strands attachingto ventrolateralmost edge of LE4 insertion and em-bodied in CT sheet also including PP.LI 1 on lateral surface of Pb2 uncinate process dor-soposteriorly, noticeably larger than LI2.
?
On dorsalsurface of Pb2 just ventral to tip of uncinate process,same size as LI2.LI2 on Pb3 dorsoposteriorly and slightly antero-ventral to dorsomedialmost edge of Eb3.TD comprises TPb2-Eb2 and TPb3-Eb3 (see alsoadditional remarks end of description). TPb2-Eb2 fi-bers complex anteriorly, attaching to medial surfacesof Pb2s anteriorly, muscle fibers "weave" into trans-verse muscle fibers posteriorly that insert dorsally onEb2, slightly lateral to medial end of Eb2 and slightlymedial to expanded dorsoposterior process. TPb3-Eb3 triangular, apex anteriorly between medial endsof OD3-4s, partially ventral to OD-3-4 laterally,ventrolaterally attaching to Pb3 and continuing ontodorsomedialmost surface of Eb3.
? Comprises TPb2, TEb2, and TPb3-Eb3-Eb4.TPb2 on Pb2 uncinate process, meeting LI1 insertion,continuous posteriorly by diagonal muscle strap withTEb2. TEb2 on Eb2 dorsoposteriorly well medial toLE2 insertion, continuous posteriorly by diagonalmuscle strap (passing ventral to OD3-4) with TPb3-Eb3-Eb4. TPb3-Eb3-Eb4 on Pb3 dorsally beginningslightly anterior to LI2 insertion and continuing alongmedial edge of insertion onto Eb3 dorsoposterome-dially and EB4 dorsomedially, continuous by diago-nal muscle strands with SOD.OD3-4 origin broadly on Pb3 medially ventral to
NUMBER 11 117
TPb2-Eb2 posteriorly, inserting on dorsomedial edgeof Eb3 uncinate process and dorsoposterior edge ofEb4 uncinate process, forming partial raphe with OPdorsolaterally.
?
On Pb3 medially ventral to TEb2.OP broad strap dorsally on posterior surface ofEb4 lateral to Ad4 dorsal attachment, forming partialraphe dorsomedially with OD3?4, narrowing slightlyventrally and forming raphe (ER) with Ad5 dorso-laterally, raphe continuous laterally between Ad5 andAd4.Adl-3 absent.Ad4 dorsally on posterolateral surface of Eb4 lat-eral to OP dorsally, dorsoposteriorly joining raphewith LE4 ventrally, and ventroposteriorly joining ERon Cb4 with Ad5 dorsally; anterior body of Ad4 ven-trally on Cb4 dorsal surface anterior to Eb4-Cb4joint.Ad5 ventrally broadly on Cb5 dorsodistal surface;dorsally, beginning posterodistally on Cb4, joiningraphe (ER) with Ad4 ventrally and OP ventrally.SOD present, broad.RDs separate, adjacent.Additional remarks. SCL present, attached dorso-medianly to posteroventral cartilaginous tip of Bb3.TV4 free from Cb5s. CT pad of TD anteriorly almostcompletely envelops Pbls. Pb2 uncinate processforms small, naked articulating facet (? facet absent).Pb4 and UP4 absent. IAC small. Eb4 levator processabsent.
PercomorphaSmegmamorpha
Johnson and Patterson (1993) hypothesized thisgroup to include Mugilomorpha. Atherinomorpha.Elassomatidae, Gasterosteiformes (comprising Gas-terosteidae, Aulorhynchidae, Hypoptychidae, Pegas-idae, Indostomidae, Solenostomidae, Syngnathidae,Centriscidae, Aulostomidae, Fistulariidae), Synbran-choidei, and Mastacembeloidei. The monophyly ofthe group, based primarily on a single synapomorphy(origin of first epineural at distal tip of a transverseparapophysis), has been controversial ever since.We do not believe the Smegmamorpha are mono-phyletic (a more formal denial is provided by Spring-er and Orrell in the Appendix), but, for the sake ofconvenience, we treat the members of the group thatwe examined together. We partition the Smegma-morpha into three groups, Gasterosteomorpha, Ath-erinomorpha, and Mugilomorpha. We believe the lasttwo groups are closely related, but not closely relatedto the Gasterosteomorpha. The Atherinomorpha havebeen strongly supported as a monophyletic group formany years and questionably allied to the Mugilo-morpha. We believe the Gasterosteomorpha are pos-sibly polyphyletic. We recognize four groups within
the Gasterosteomorpha: Centrisciformes, Gasterostei-formes, Synbranchiformes, and Elassomatiformes.Johnson and Springer (1997:176) implied thatthere is a close relationship between Elassomatidaeand Gasterosteidae. They did not follow this with aformal publication, and the matter remains unre-solved. GasterosteomorphaWe believe Centriscidae, particularly Macroram-phosus (Centriscops, Notopogon should also be ex-amined), are either a monophyletic group not closelyrelated to other gasterosteomorphs or are the sistergroup of the gasterosteomorphs. Macroramphosus isone of only three taxa (others: Icosteus, Icosteidae;Mene, Menidae) that are usually classified among thepercomorphs that possess ER and may have OP ven-trally attaching to Cb4, conditions otherwise knownto occur among pre-acanfhomorphs (commonly) andmore basal acanthomorphs (i.e., polymixiids, para-canthopterygians, zeiforms, stephanoberyciforms,beryciforms). All other of Macroramphosus 's puta-tively closely related gasterosteomorphs (i.e., syng-nathoids) are highly specialized and have lost manyskeletal elements; their muscles are also degenerate,but if OP is present, it attaches to Cb5.If Macroramphosus is validly removed from theGasterosteomorpha and placed in a more basal acan-thomorph position, the entire classification of theGasterosteomorpha needs to be re-evaluated, whichmight result in positioning the entire Gasterosteo-morphas among the pre-percomorphs.Insofar as we can determine, the term Gasterosteo-morpha was first used by Nakabo (2002:53, 56, 512),who recognized Johnson and Patterson's Smegma-morpha, and included under it four groups: Gaster-osteomorpha, Synbranchomorpha, Mugilomorpha,Atherinomorpha. The composition within each ofthese four groups mentioned only Japanese forms;therefore, it is uncertain how he would have treatedElassomatidae, although probably as a fifth group.We employ Gasterosteomorpha here to include Cen-trisciformes, Gasterosteiformes, Elassomatiformes,and Synbranchiformes. Although a reconstitutedSmegmamorpha (1993) might be used for the group,we find that name marginally offensive and its orig-inal composition misleading. The ICZN does notspecify that priority pertains to taxa above the family-group level. CentrisciformesCENTRISCIDAEMacroramphosus scolopax (Linnaeus), USNM366546 and 158813, both 111 mm; USNM366727, 110 mm, 177091, 126 mm.Plate 86
Hi BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Description.LEI on high, bony spike-like Ebl process dorso-medially (cartilage-tipped Ebl uncinate process ab-sent); strong, ribbon-like tendon on lateral edge ofmuscle, separating from muscle about halfway upfrom insertion and originating separately on skull;fine, thread-like tendon separates from posterior edgeof ribbon-like tendon and inserts in CT around dorsalend of Pb3 laterally.LE2 tendinously on mid-dorsal edge of Eb2 andcontinuing tendinously posteriorly and attaching toEb3 mid-ventroanteriorly; anteromedial edge of in-sertion joined by posterodistal edge of TEb2.LE3 absent.LE4 on Eb4 mid-dorsally lateral to uncinate pro-cess, insertion joined posteriorly by LP.LP tendinously on Eb4, joining LE4 insertion pos-teriorly.LI1 on Pb2 ventroposterior to dorsal process, andventrolateral to origin of OD3-4 on Pb2 [sic],LI2 on Pb3 dorsally medial to anterior portion ofjoint articulating with medial end of Eb3.TD comprises TEb2 and TPb3-Eb3. TEb2 rela-tively naiTOw, boomerang-like (anteriorly concave),with mid-longitudinal raphe continuing posteriorlyhalfway or completely across TPb3-Eb3; TEb2 atta-ches ventroanteriorly to complex CT pad and tissuescovering dorsal ends of Pb2, Pb3, and Eb3 (tissuesremoved in illustration); pad continuous anteriorlyand mid-ventrally with CT of pharyngeal roof (limp,but tough ligament inserts on Pb2 posteromediallyventral to cartilaginous Pb2 dorsal cap and immedi-ately medial to origin of OD3-4 portion inserting onPb2, and also deeply on Pb2 surrounding LI1 inser-tion medially, thus straddling Pb2 portion of OD3-4insertion; similar ligament inserts on Pb3 dorsopos-teriorly; see also Synbranchus, Synbranchidae, forsimilar ligaments); TEb2 attaches on Eb2 beginningat and medial to LE2 insertion and continuing dor-soanterolaterally almost to distal end of Eb2, thereoverlain anteriorly by GFM2; TEb2 continuous withEb3 portion of TPb3-Eb3 anteriorly by diagonal mus-cle strands. TPb3-Eb3 broadly triangular in dorsalview, variously appearing to consist of two meshingportions that extend laterally and attach to medialedge of Eb3 uncinate process and, separately, to Eb3dorsal surface medial to uncinate process; muscle ex-tends anterolaterally ventral to OD3-4-OD3' andTEb2 and attaches to Pb3 dorsomedial to joint withEb3, and is continuous posteroventrally and posteri-orly with SOD.OD3-4, OD3' with separate origins on Pb2 andPb3; origin on Pb2 posteromedially well ventral tolarge dorsal cartilage cap, straddled laterally and me-dially by limp ligament (described in TD above); fi-bers extend posteriorly from Pb2, meshing ventrallywith main body of muscle originating on Pb3 ventral
to TEb2; muscle divides posteriorly with ventral fi-bers (OD3') inserting on Eb3 anterior surface ventralto tip of uncinate process and dorsal fibers (OD3-4)inserting finely on medial edges of joined Eb3 andEb4 uncinate processes.OP dorsally on posterior surface of Eb4 beginninglateral to medial end and extending laterally a vary-ing distance, but no further than ventral to LE4 in-sertion, there variably separable or not from Ad4 dor-sally; divided at mid-level (level at Cb4) by smallraphe (ER) and attaching on Cb5 medial to Ad5, var-iably appearing undifferentiated from Ad5 ventral toER.GFM1 dorsally on and covering anterior surfaceof Ebl beginning a little ventral to spike-like processand extending anteroventrally most of length of Cbl.GFM2 beginning on Eb2 dorsomedial surface andcontinuing laterally on dorsoanterior surface anteriorto TEb2, fanning out anterolaterally from medial at-tachment to Eb2, thereby anteriorly covering TEb2laterally and Eb2-Cb2 joint, tapering ventrally, andextending anteroventrally along most of anterior sur-face of Cb2.GFM3 beginning on Eb3 dorsomedially and con-tinuing laterally, fanning out anterolaterally, coveringEb3-Cb3 joint, tapering ventrally, and extending an-teroventrally along distal fifth of Cb3; thin ribbon ofmuscle fibers splits off GFM posterodistally and at-taches to Eb3-Cb3 joint anteriorly (not visible in il-lustration).Ad4 dorsally on ventral and posterior surfaces ofEb4 lateral to OP, fusing with OP anterolaterally;ventrally relatively broadly on dorsolateral surface ofCb4 medial to Eb4-Cb4 joint.Ad5 relatively deep, dorsally on posterolateralmostsurfaces of Eb4 and Cb4, covering joint of these twobones, ventrally on Cb5 dorsolateral^ posterior toOP.SOD broad, continuous anteriorly with TEb3.RDs moderate, adjacent or slightly separated.Additional remarks. SCL attached mid-dorsally tobony posterior end of Bb3. TV4 free from Cb5s. Pblcartilaginous plate, Pb4 absent, UP4 present. Pb2toothed. IAC absent. Eb4 levator process absent.
GasterosteiformesGASTEROSTEIDAEGasterosteus aculeatus Linnaeus, USNM 344603,62.0 mm; USNM 58296, 64.6 mm.Plate 87
Additional material (information on these taxa in-cluded for the most part only in the data matrix.Table 12, used for the cladistic analyses). Apeltesquadracis (Mitchill), USNM 242691, 42.2 mm;Cuiea inconstans (Kirtland), USNM 69287 41.5
NUMBER 11 119
mm; Pungitius pungitius (Linnaeus), USNM77842, 49.8 mm; Spinachia spinachia (Linnaeus),USNM 23008, 138 mm, USNM 344839, 139 mm.
Description.LEI on Ebl surrounding bony or cartilage tippeduncinate process (see remarks), joining raphe ante-roventrally with Adl dorsally; fine ligament extendsposteriorly from posterolateral edge of raphe to an-terior edge of Eb2. See remarks following descriptionof LEI in Elassoma.Remarks. Primitively in acanthomorphs, the ante-rior or medial process of Ebl articulates with Pbland the uncinate process with IAC or Pb2 (articula-tion may be direct or ligamentous). On this basis weconsider the anterior process absent in Elassoma andgasterosteiforms, and the process articulating withPb2 to be the uncinate process. We interpret the oc-casional presence of a small cartilage tip on the ex-panded edge of Ebl in Gasterosteus, which is com-pletely surrounded by the insertion of LEI, as a fail-ure of the edge to completely ossify, as opposed totreating it as the tip of the "true" uncinate process.We know of no case where an uncinate process ispresent and the anterior process of Ebl articulateswith Pb2.LE2 on raised bony process on Eb2 posterolater-
ally, joining raphe ventromedially with lateral end ofTEb2 and another ventrolaterally with posterior por-tion of Ad2; fine ligament extends posteriorly fromlateral edge of latter raphe to anterior edge of Eb3.LE3 absent.LE4 broadly on Eb4 dorsoposteriorly, joining LPanteroventrally.Remarks. Uniquely among the gasterosteids ex-amined, Spinachia apparently lacks LE4. This inter-pretation is based on the fact that LE4 in the othergasterosteids angles dorsoanteriorly towards its ori-gin, whereas LP is almost vertical, as well as joiningAd5 dorsally (see LP).LP on most of Eb4 lateral to uncinate process (mayextend both lateral and medial to junction with LE4insertion), joining raphe with Ad5 dorsally, thusforming an LP-Ad5 sling; true in all gasterosteidsexamined.LI1 mostly on Pb3 dorsoanteromedially dorsal toand meeting origin of OD3-4, continuing, second-
arily, onto Pb2 dorsomedially.LI2 on Pb3 dorsoposteriorly opposite medial endof Eb3.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2broad medially, narrowing considerably laterally,with mid-longitudinal raphe giving rise dorsally toCT sheets and attaching ventroanteriorly betweenPb3s to CT of pharyngeal roof, extending laterallyand attaching on raised bony edge of Eb2 joiningraphe with medial edge of LE2 insertion (not ex-
tending laterally past medial edge of LE2 insertion);not continuous posteriorly with TPb3-Eb3-Eb4.TPb3-Eb3-Eb4 on Pb3 dorsolateral^ just medial toLI2 insertion, continuing posteriorly and attachingnarrowly on dorsomedialmost end of Eb3 (there join-ing raphe with medial end of anterior portion ofAd3), then continuing posteriorly, but well separatedlaterally, with attachment to Eb4 dorsally ventrome-dial to uncinate process; posteriorly continuousbroadly with SOD.OD3-4 origin on Pb3 broadly dorsomedially, an-teriorly ventral to and joining LI1 insertion, insertionon medial edges of Eb3 and Eb4 uncinate processes.OP dorsally beginning on Eb4 posteriorly ventralto uncinate process and extending broadly medially,ventrally on Cb5 beginning at ventrolateral edge ofAd5, medially unclearly separable from SO.Adl broad, consisting of a single portion (equiv-alent to posterior portion of Ad2 and Ad3) extendingfrom raphe with LEI ventroanteriorly across Ebl-Cbl joint to point on Cbl anterior bony surface justventral to joint, ventrolateral edge attaching to fila-ments. Present and similar in all gasterosteids ex-amined.Ad2 comprising two portions, posterior portion ex-tending from raphe with LE2 ventroanteriorly acrossEb2-Cb2 joint to point on Cb2 anterior bony surfacejust ventral to joint; anterior portion beginning onEb2 dorsomedially and extending laterally and pass-ing anterior to most of posterior portion, with lateraledge attaching to gill-rakers and/or gill filaments.Present and similar in all gasterosteids examined.Ad3 comprising two portions, posterior portion at-taching dorsally to anterolateral edge of Eb3 uncinateprocess, continuing on ligament joining Eb3 and Eb4uncinate processes and anterodistal end of Eb4; an-terior portion beginning on dorsomedial surface ofEb3, joining raphe with lateral end of TPb3-Eb3-Eb4and extending laterally and passing anterior to mostof posterior portion, with lateral edge attaching togill-rakers and/or gill filaments. Present and similarin all gasterosteids examined.Ad4 dorsally on Eb4 broadly ventrally, ventrallybroadly on Cb4 medial to Eb4-Cb4 joint, slightly fus-ing dorsoposteriorly with Ad5 dorsoposteriorly; com-pletely obscured from view posteriorly by Ad5.Ad5 dorsally beginning on Eb4-Cb4 joint and ex-tending broadly medially on Eb4 to edge of OP, join-ing raphe with LP ventroposteriorly. ventrally on Cb5dorsolaterally, completely overlapping Ad4 posteri-
orly.SOD broad continuous anteriorly with TPb3-Eb3-Eb4.SO lateral band of fibers extends anteriorly overmedial end of Eb4 and ventral to TPb3-Eb3 and in-
serts on Pb3 posteriorly.RDs adjacent.
120 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Additional remarks. SCL present, attached mid-dorsally to wedge of cartilage, which is attached toventral surface of cartilaginous posterior end of Bb3;Hb3s attach to wedge posterolaterally. TV4 free fromCb5s. Pbl, Pb4 and UP4 absent. Eb3 and Eb4 un-cinate processes present. Eb4 levator process absent.Anker (1974) described and illustrated the headskeleton and muscles of Gasterosteus aculeatus. Ourfindings are in essential agreement with his.HYPOPTYCHIDAE
Johnson and Patterson (1993) removed Aulichrhysfrom the Aulorhynchidae and included it in the Hy-poptychidae. We agree with that action.
Hypoptychus dybowskii Steindachner, USNM 51494,
ca. 85 mm; USNM 118009, 72.8 mm; UW 29655,3 specimens, 65?67 mm.Plate 88
Remarks. Damage occurred in every specimenduring release of the levators, and extreme difficultywas encountered in resolving the limits and attach-ments of the more posterior (non-levator) muscles.Additionally, there appeared to be complex variation.The description and Plate 88 are a composite basedon all the specimens. Every muscle described or il-lustrated was seen in at least two specimens.
Description.LEI extremely fine, posterodistally on Ebl, con-tinuing just onto CT (not illustrated) between Ebl anEb2.LE2 extremely fine, posterodistally on Eb2, con-tinuing just onto CT (not illustrated) between Eb2and Eb3.LE3 finest of all levators, usually lost during dis-section, on Eb3 lateral to uncinate process, continu-ing just onto CT (not illustrated) between Eb3 andEb4; expanding at origin, and joining, or nearly so,LE4 origin.LE4 relatively large, on Eb4 dorsolaterally, joiningraphe ventrolaterally with Ad5 dorsally; muscle mayappear incompletely divided into lateral section,which is continuous with Ad5, and medial section,which is not; LE4 expands at origin, probably join-ing, or nearly so, LE3 origin.Remarks. The LE3 and LE4 origins are posteriorto those of other levators. Usually, if not always, inpercomorphs, the origins of LE3 and LE4 join theorigins of the other levators, none of which ever joinsthe origin of LP. Our recognition of the levator onEb4 as LE4, rather than LP is based on this fact. Thealternative, that LE4 is lost and the origin of LE3 hasmoved posteriorly and joined that of LP, seems lessparsimonious.LP absent (see remarks following LE4).
LI1 on Pb2 dorsolaterally just anterior to articu-lation with Eb2.LI2 on Pb3 posterodistally just anterior to medialend of Eb3.TD comprises TEb2 and TPb3. TEb2 boomerang-like, attached mid-ventroanteriorly to CT of pharyn-geal roof, extending laterally and attaching to Eb2dorsally anterior to LE2; uniquely, ventrolaterallyfinely continuous with portion of OD3-4 origin onPb2; posteriorly meeting but not continuous withTPb3. TPb3 broad, attaching along much of Pb3 dor-
sal surface, continuous posteriorly with broad SOD.OD3-4 origin on Pb2 and Pb3 dorsally ventral toTEb2 (strands from origin on Pb2 attach on Eb2 insome specimens and join with TEb2); after passingposteriorly from under TEb2, muscle joins a branchof SO that passes posteriorly broadly over Eb4 (noraphe apparent) and continues to distal end of Cb5medial to OP. OD3-4 extends dorsally over Eb3 andEb4 uncinate processes (obscuring Eb4 uncinate pro-cess and, usually, Eb3 process in dorsal view), withslight insertion on Eb3 process and main insertion onEb4 process, and is continuous with OP medial sec-tion on Eb4 (raphe absent).OP strap-like, dorsally on Eb4 posterior to unci-nate process, dorsomedialmost fibers continuous withOD3-4 posteromedially; ventrally, attaching to Cb5dorsodistally near Ad5.Adl on anterodistal surface of Ebl and dorsoan-teriormost surface of Cbl.Ad2 on anterolateral bony surface of Eb2, meetingdistal end of TEb2, and dorsoanteriormost surface ofCb2.Ad3 on most of bony surface of Eb3 lateral touncinate process and on dorsoanteriormost surface ofCb3.Ad4 dorsally, broadly on Eb4 posteriorly betweenOP and Ad5; ventrally on Cb4, extent not deter-mined.Ad5 a small muscle strap, ventrally on Cb5 dor-soposterodistally; dorsally on Eb4 dorsoposterodis-tally, there joining raphe with LE4 ventrolaterally;small anterior portion of muscle (not visible in Plate88) attaches to posterodistal end of Cb4.SOD very broad.RDs slender, separated by space more than twiceone RD diameter.Additional remarks. SCL present, questionablyfree from Bb3, which has ventrally extending pos-terior end, usually indicative that SCL attaches to it.TV4 free from Cb5s. Pb4 present, Pbl and UP4 ab-
sent. Pb2 toothed. IAC absent. Ebl uncinate processpresent, anterior process absent (see remarks follow-ing description of LE 1 under Gasterosteus for expla-nation). Eb4 uncinate process present, levator processabsent.
NUMBER 1 1 121
Aulichthys japonicus Brevoort, USNM 347504, 136mm; USNM 59789, 129 mm; USNM 71104, 2specimens, 68.5?71.8 mm.Plate 89
Description.LEI on posterior edge of Ebl just medial to car-tilaginous distal end.LE2 on posterior edge of Eb2 just medial to car-tilaginous distal end.LE3 on tip of Eb3 uncinate process anteriorly.LE4 short, on Eb4 dorsoanteriorly near distal end.LP short, on Eb4 at and posterolateral to LE4 in-sertion.LI1 on Pb2 dorsoanteriorly and adjacent bony sur-face of Pb3 anterolaterally.LI2 on Pb3 dorsoposteriorly anterior to medial endof Eb3.TD comprises TEb2 and TPb3. TEb2 attaches ven-tromedially to CT of pharyngeal roof and laterally toEb2 dorsally medial to LE2 insertion, continuousmid-posteriorly with TPb3. TPb3 laterally passesventral to OD3-4 and attaches to Pb3 broadly ante-
rior to LI2 insertion; posteriorly muscle is broadlycontinuous with SOD; posterolaterally, anterior fibersof SO pass through and ventral to TPb3 and inserton Pb3 medially.Remarks. SO fibers originate diffusely medial toOP and extend dorsoanteriorly, passing among ante-
rior SOD fibers and (difficult to see) possibly throughposterior TPb3 fibers and attaching to Pb3.OD3-4 origin on Pb2 dorsally posterior to LI1 in-sertion and on Pb3 dorsally ventral to TEb2, insertionon Eb3 uncinate process anteriorly and medial edgeof Eb4 uncinate process.OP dorsally on Eb4 posteriorly, extending fromnear medial end to uncinate process, there apparentlyfusing completely with Ad4 medially; ventrally onCb5 dorsodistally, partially fusing anteriorly withAd5; medially fusing with SO.Ad 1-3, each beginning on anterior surface of re-spective Eb at about mid-length of Eb and extendinglaterally and spreading and attaching on anterior sur-face of Eb-Cb joint to include cartilaginous distal endof Cb; posterior surface of muscle takes twist at distalend and appears to pass posterior to itself.Ad4 attaches dorsally to Eb4 posteriorly lateral touncinate process and attaches ventrally to Eb4 dor-somedially, fusing medially with dorsal half of OP.Ad5 very reduced, ventrally on Cb5 distally, pos-teroventrally fusing with OP, dorsally on Cb4 poster-odistally.SOD very broad.RDs slightly separated.Additional remarks. SCL questionably free fromBb3. TV4 free from Cb5s. Pbl, Pb4 and UP4 absent.Pb2 toothed. IAC absent. Tiny AC present on dor-
sodistal end of Ebl, apparently not homologous withAC1 as it is completely removed from Cbl. Aulos-tomus has an elongate extension of the cartilaginousdistal end of Ebl, which, if budded off, would forma similar appearing AC (Aulostomus is unique amonggasterosteiforms in having IAC). Eb4 levator processabsent. Ebl uncinate process present, anterior processabsent (see remarks following description of LEI un-der Gasterosteus for explanation).AULORHYNCHIDAEAulorhynchus flavidus Gill, USNM 344688, 2 spec-imens, 143-144 mm SL; USNM 167915, 89.9 mmSL. Plate 90
Description.LEI on Ebl dorsoposterolaterally, insertion con-tinuing posteriorly on CT (not illustrated) joiningEbl to Eb2 anterodistally.LE2 on Eb2 dorsoposterolaterally, continuing pos-teriorly as CT (not illustrated) joining Eb3 anteriorly.LE3 absent.LE4 on Eb4 dorsolaterally.LP on Eb4 joining LE4 insertion posteriorly, join-ing raphe with Ad4 dorsally.LI1 massive, on Pb2 dorsoanteromedially and Pb3anterolateralmost edge, which abuts Pb2.LI2 on Pb3 dorsoposteromedially anterior to me-dial end of Eb3.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2flat, hood-like viewed dorsally; attaches finely later-ally on dorsoposterior edge of Eb2 at medial edge ofLE2 insertion, attaches ventrally along mid-longitu-dinal line to CT of pharyngeal roof; continuous pos-teriorly by diagonal muscle strand with TPb3-Eb3-Eb4. TPb3-Eb3-Eb4 anterolaterally on Pb3 dorsolat-erally between joints with medial ends of Eb3 andEb4. continuing posteriorly on medial ends of Eb3and Eb4 dorsally, continuous mid-posteriorly by di-agonal muscle strands with broad SOD.OD3-4. OD3' origin on almost entire dorsomedialmargin of Pb3, muscle separates into OD3?4 dorsallyand OD3' ventrally on exiting posteriorly from underTEb2; OD3-4 inserts on Eb3 uncinate process an-teriorly and Eb4 uncinate process medially; OD3' in-serts on Eb3 dorsally ventromedial to uncinate pro-cess. See also OP below.OP dorsally on Eb4 posteriorly beginning ventralto and between uncinate process and LP insertion andextending medially and becoming indistinguishablefrom SO (in one specimen slender group of musclefibers from OD3-4 extends posteroventrally medialto Eb4 uncinate process and joins OP fibers); ven-trally on Cb5 dorsodistally, apparently fusing withAd5 medially.Adl begins on dorsoanterior surface of Ebl an-
122 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
teromedial to LEI insertion and follows anterior edgeof Ebl laterally, attaching to anterior surfaces of Ebland Cbl where they join; short branch of muscle fi-bers separates anteriorly from remainder of muscle atdistal end of Ebl and attaches to distalmost gill rakerof dorsal arch (rakers do not follow Ebl margin, butextend directly anteriorly from distal end of Ebl).Remarks. Our description of the short branch ofAdl (and also that of Ad2 and Ad3) is problematic;Plate 90 presents another interpretation, with theshort branch of fibers separating posterolaterally fromthe remainder of the muscle.Ad2 begins on dorsomedialmost bony surface ofEb2 ventral to TEb2 and extends laterally on anterioredge of Eb2, covering anterior surfaces of Eb2 andCb2 where they join; short branch of muscle fibersseparates anteriorly from remainder of muscle at dis-tal end of Eb2 and extends anteriorly, diminishingshortly and becoming CT that attaches to Ebl pos-teriorly. See remarks following Adl.Ad3 like Ad2, but involving Eb3 and Cb3. Seeremarks following Ad 1
.
Ad4 dorsally on ventral surface of Eb4 posteriorlymostly lateral to OP, ventrally on Cb5 dorsal surfaceposteriorly medial to Eb4-Cb4 joint.Ad5 questionable small band of muscle joiningposterodistal end of Cb4 and anterodistal end of Cb5.fused indistinguishably with OP ventrolaterally.SOD very broad.RDs adjacent.Additional remarks. SCL attached mid-dorsally toventrally extending cartilaginous posterior tip of Bb3.TV4 free from Cb5s. Pbl and Pb4 absent, UP4 pres-ent. Pb2 toothed. IAC absent. Ebl uncinate processpresent, anterior process absent (see remarks follow-ing description of LEI under Gasterosteus for expla-nation). Eb4 levator process absent.
SynbranchiformesThe Synbranchiformes comprise the synbranchoidsand mastacembeloids. Britz (1996) last supportedrecognition of a monophyletic group consisting ofthese two suborders. The interrelationships of theSynbranchiformes are unclear.SYNBRANCHIDAESynbranchus marmoratus (Bloch), USNM 31 1207, 2specimens, 298-307 mm TL.Plate 91
Additional material.
?
= Ophisternon bengalensisMcClelland, USNM 135205, ca. 350 mm TL. Not
all data were available on this specimen; absenceof comment should not be interpreted as agreementwith description of S. marmoratus.
Description.Remarks. All of the levators extend anteriorly al-most horizontally parallel to mid-line axis of body.Their orientations in the illustrations have been mod-ified variously.
? Same.LEI on Ebl dorsally near distal end. Uncinate pro-cess absent. ? Same.LE2 divided, with separate anterior insertion onEb2 posterodistalmost edge and posterior insertion(LE2') on CT between Cbl and Eb3 or on CT butwith tendon extending posteriorly from insertion toCb3 somewhat ventral to joint with Eb3.
? Not di-vided.Remarks. In normal position LE2 and LE2' appearto be one muscle, but manipulating the muscles theyeasily separate basally and the separation can be con-tinued to (or almost to) their joint origin. The con-dition appears to be a derivation of the state of LE2in the related Mastacembelus, in which LE2 is notdivisible and inserts on Eb2 dorsoposterior edge andEb3 dorsoanterior edge.LE3 absent (both sides of both specimens). ?Highly reduced LE3 present on only one side; forcladistic analysis, we consider LE3 absent.LE4 on bony dorsodistal end of Eb4, divided atorigin.
?
Same.LP absent. ? Same.LI1 massive, inserting in CT attached to medialend of IAB, enveloping Pb2 (which may be vestigialor absent), and attached to or enveloping medial endof Eb2 and anterior end of Pb3 (none of the insertionpoints are visible in illustrations).
? Same.LI2 on Pb3 dorsoposterolaterally opposite medialend of Eb3.
?
Same.TD with almost complete mid-longitudinal raphe,comprises TEb2, TPb3-Eb3, and TEb4 (see addition-
al remarks for discussion of absence of TPb2). TEb2a pair of muscles, each angled anterolaterally and at-taching on respective Pb3 dorsoanteriormost end andEb2 dorsomedially, lies dorsal to OD3-4 origin andis continuous mid-posteriorly with TPb3-Eb3. TPb3-Eb3 (see also remarks following TD description) onPb3 dorsally along medial edge of LI2 insertion, pos-teriorly extending laterally and attaching on Eb3 dor-somedially, continuous narrowly mid-posteriorlywith TEb4. TEb4 more-or-less triangular, attachingalong posterior edge of Eb4 and ventral surface ofligament complex extending anteriorly to Pb3 andEb4 (exact nature of ligaments unclear), muscle con-tinuous posteriorly with SOD. ? Comprises TEb2(not split) and TPb3-Eb3-Eb4, shallow gap presentbetween Pb3 and Eb3, and Eb3 and Eb4 attachments.OD3?4 originates narrowly on joint formed by an-terolateral end of Pb3 and medial end of Eb2; insertsmassively dorsally on dorsolateral surface of Eb4(covering joint with Eb3 uncinate process and ante-
rior edge of Eb4 (lacks uncinate process), which may
NUMBER 1
1
123
be slightly modified as a bony process or only in-dented) and ventrally, less extensively on Eb3 unci-nate process, the tip of which may be bony or mi-nutely cartilaginous. ? Originates on Eb2 and adja-cent Pb3 and inserts similarly to S. marmoratus, ex-cept muscle does not cover Eb3 and Eb4 processes(Eb3 uncinate process with cartilaginous tip).OP dorsally on Eb4 ventrolaterally, ventrally onCb5 dorsoposteriorly, there overlapping Ad5 ventro-posteromedially, medially mostly inseparable fromSO.Adl (not illustrated) questionably interpreted as afan of muscle on Ebl-Cbl joint anteriorly; mainlyassociated with gill filaments.Ad2 on Eb2 mid-dorsoanteriorly, extending ven-trolaterally onto anterolateral surface of Cb2, ventraledge attached to gill filaments.Ad3 on Eb3 anterolaterally. extending onto Cb3anterolateral ly, ventral edge attached to gill filaments.Ad4 dorsally on Eb4 and ventrally on Cb4. medialto Eb4-Cb4 joint.Ad5 dorsally broadly on Cb4 ventrolaterally, ven-trally on Cb5 dorsally. extending medially anterior toOP.SOD broad, continuous anteriorly with TEb4.
?Present.RDs separated by space more than one RD diam-
eter.Additional remarks. SCL absent; ? Same. TV4free from Cb5s.
?
Same. Pbl and Pb4 absent. UP4present. Eb4 levator process absent.
?
Same. Smallbony flange at distal bony end of Eb4 dorsally ordorsoanteriorly shields cartilage. ? Same (see Rosenand Greenwood, 1976:fig. 35, where Eb4 flange isindicated but not labeled).Pb2 is present as an edentate bony fragment in S.marmoratus. Rosen and Greenwood (1976:fig. 36) il-lustrate a moderately well-developed, but edentate,rod-like Pb2 in the dorsal gill-arch skeleton of S.marmoratus, and (figs. 28-34) and as a small, eden-tate, plate-like bone in all seven species of Ophister-non; our specimen agrees.Rosen and Greenwood (1976:7) indicated the pres-ence of a tendon, "t-rab. tendon of M. retractoresarcum branchialium," (their rab = our RD), whichthey illustrated as inserting on UP4 posteriorly in var-ious synbranchids, but not Synbranchus. They did notdiscuss the tendon, which we believe is more accu-rately considered as a ligament). The ligament is in-tegral with RD ventrolaterally for the muscle's entirelength, and it remains after the muscle is digestedduring clearing and staining. Rosen and Greenwoodalso did not mention other non-muscle integrated lig-aments attaching variously to Pb3 (also attaches tocleithrum), Eb3, and Eb4 in synbranchids. On oneside of the specimen of Ophisternon, the ligamentsappear to unite in a complex mesh, and we are un-
certain if our dissections of Synbranchus did not de-stroy the integrity of the ligaments (hence, their ren-dition in Plate 91 should be verified).MASTACEMBELIDAEMastacembelus armatus (Lacepede), USNM 319465,275 mm; USNM 343569, 300 mm.Plate 92
Description.LEI on dorsoposterior edge of Ebl and dorsoan-terior edge of Eb2, meeting Ad2 anteriorly. Cartilage-tipped Ebl uncinate process absent.LE2 on dorsoposterior edge of Eb2 and dorsoan-terior edge of Eb3, meeting Ad3 anteriorly.LE3 absent.LE4 on Eb4 dorsally lateral to uncinate process.LP absent.LI1 massive, on joined anterior ends of Ebl, Pb2,and Pb3 dorsally, meeting anteriormost edge ofOD3-4 origin.LI2 on Pb3 dorsolaterally anterior to medial endof Eb3; muscle meets lateral edge of TPb3-Eb3.TD comprises TEb2 and TPb3-Eb3 (a few strandsof TD attach to posteromedial edge of Eb4 on oneside of one specimen). TEb2 strap-like with mid-lon-gitudinal raphe attaching dorsally to CT sheets (notshown) and ventrally to CT of pharyngeal roof; mus-cle attaches laterally on Eb2 dorsally, meeting pos-teromedial edge of LEI insertion and medialmostedge of Ad2, and is continuous mid-posteriorly bydiagonal muscle straps with TPb3-Eb3. TPb3-Eb3broadly, dorsally on Pb3 at medial edge of LI2 in-sertion, continuing posteriorly and attaching on pos-terior edge of Eb3 medial to uncinate process; con-tinuous posteriorly by diagonal muscle strands withSOD.OD3?4 origin on Pb3 dorsally ventral to TEb2,meeting posterior edge of LI1 insertion; muscle di-vides posteriorly with dorsal portion inserting on Eb4uncinate process and ventral portion inserting on Eb3uncinate process (including medial edges of joinedprocesses).OP dorsally on Eb4 posteriorly in area ventral touncinate process, laterally overlapping Ad4 medially,ventrally on Cb5 dorsolaterally meeting Ad5 ven-trally and PCI dorsally, medially not clearly separablefrom SO.Ad 1-3 problematically present (as GFM?). Oneach epibranchial dorsoanterolaterally, there is amore-or-less well-defined strip of muscle that be-comes weak and finer distally as it extends ventrallyalong the entire anterolateral edge of the associatedceratobranchial and attaches to the gill filaments ba-
sally.Ad4 dorsally on ventroposterior surface of lateralhalf of Eb4, medially overlapped by OP, ventrally on
124 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Cb4 dorsally medial to Eb4-Cb4 joint, there joiningAd5.Ad5 dorsally on posterolateral end of Cb4 con-tinuing short distance medially and joining Ad4 ven-trolaterally, ventrally on distal end of Cb5, there join-ing OP medially and PCI dorsoanteriorly.SOD present.SO muscle straps arise from SO in area lateral toSOD, extend anteriorly dorsal to Eb4 and attach var-iously to Pb4, Eb3, or Eb4.RDs separated by space about equal to diameter ofone RD.Additional remarks. SCL absent (Bb3 with ventro-posteriorly extending cartilaginous tip). TV4 freefrom Cb5s. Pbl absent. Pb2 toothed. Pb4 and UP4present. IAC absent. Eb4 levator process absent.
ElassomatiformesELASSOMATIDAEElassoma zonatum Jordan, USNM 173343, 30.5 mm;USNM 232536, 29.9 mm.Plate 93
Description.LEI on Ebl dorsolaterally, anterior edge of inser-tion bordering Adl dorsomedially.Remarks. Medial end of Ebl articulates with Pb2and we consider it to be the uncinate process. Weconsider the anterior Ebl process, which articulateswith Pbl, but never with Pb2,when present, is absentin Elassoma and the gasterosteiforms. See remarksfollowing description of LEI in Gasterosteus.LE2 on Eb2 dorsoposterolaterally, anterior edge ofinsertion bordering Ad2 dorsally, medial edge bor-dering TEb2 posterolaterally.LE3 on tip of Eb3 uncinate process.LE4 on Eb4 dorsolaterally posterior to Eb3 unci-nate process (Eb4 uncinate process absent).LP on Eb4 along posterior edge of LE4 insertion,posteriorly just dorsal to Ad4 dorsally.LI1 on anteromedial edge of Pb2 and, mainly, onadjacent anteromedial edge of Pb3 slightly anteriorto OD3-4 origin.LI2 on Pb3 dorsolaterally anteromedial to medialend of Eb3; medial edge of insertion joins lateraledge of TPb3-Eb3-Eb4.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2very wide, mid-anterior notch leading to mid-longi-tudinal raphe that continues across anterior half ofTPb3-Eb3-Eb4; attached mid-ventrally to CT of pha-ryngeal roof, giving rise dorsally to CT sheets; at-tached laterally to Eb2 dorsally at and anterior to LE2insertion, meeting LE2 insertion anteriorly and Ad2attachment medially; continuous posteriorly by finediagonal muscle filament with TPb3-Eb3-Eb4. TPb3-Eb3-Eb4 attaching to Pb3 dorsolaterally at medial
edge of LI2, continuing onto Eb3 dorsomedialmostsurface and Eb4 dorsomedially; continuous posteri-orly by diagonal muscle strand with SOD.OD3-4 origin broadly on Pb3 dorsoanteromediallyand adjacent edge of Pb2 ventral to TEb2; insertionon Eb3 uncinate process anteriorly and dorsal edgeof Eb4 adjacent to Eb3 uncinate process.OP dorsally on Eb4 ventroposteromedially, ven-trally on Cb5 posterolaterally.Remarks. Although we describe Adl and 3 as hav-ing one or two sections, we are uncertain in thosecases where only one section is present whether it isthe result of variation or damage in dissection. Fur-thermore, it is possibly more accurate to describe theAds as having one or two shapes.Adl variably with anterior and posterior sections;anterior section dorsally on anterolateral edge of Eb 1
,
posterior section dorsally meeting anterior edge ofLE 1 ; sections fuse laterally and attach to anterior sur-face of distal ends of Eb 1 and Cb 1
.
Ad2 with anterior and posterior sections; anteriorsection dorsally extending medially and meeting an-terodistal edge of TEb2; posterior section meetingventroanterior edge of LE2 insertion; sections fuselaterally and attach to anterior surface of distal endsof Eb2 and Cb2.Ad3 variably with anterior and posterior sections;anterior section on anterolateral edge of laterally andattach to anterolateral surface of Eb3 and Cb3; whenposterior section is absent, medial attachment is toEb3 ventroanterior to uncinate process.Ad4 dorsally on Eb4 posteriorly ventral to LP in-sertion and lateral to OP; ventrally on Cb4 dorsallymedial to Eb4-Cb4 joint.Ad5 dorsally on Cb4 posterolaterally, ventrally onCb5 dorsolaterally.SOD broad, continuous anteriorly with TPb3-Eb3-Eb4.RDs slightly separated.Additional remarks. SCL attached mid-dorsally toventral surface of cartilaginous posterior end of Bb3.TV4 free from Cb5s. Pbl and Pb4 absent, UP4 pres-
ent. Pb2 toothed. IAC absent. Eb4 levator processabsent. MugilomorpphaMUGILIDAEAgonostomus monticola (Bancroft), USNM 322336,90.4 mm, USNM 372452, 110 mm (gill archesnow cleared and stained).Plate 94
Description.LEI narrowly on cartilaginous tip of Ebl uncinateprocess.LE2 on tip of dorsally expanded bony posterior
NUMBER 1 1 125
edge of Eb2; ligament extends from posterior edgeof insertion to anterior edge of Eb2.LE3 on cartilaginous tip of Eb3 uncinate process.LE4 narrowly on posterior edge of mid-posteriorbony Eb4 process lateral to uncinate process.LP broadly on Eb4 dorsoposteriorly, beginning atLE4 insertion and extending well laterally.LI1 on Pb2 dorsoanterior process just ventral toarticulation with IAC.LI2 on Pb3 dorsolaterally just anterior to medialend of Eb3.Remarks. Stiassny (1990:6-7; 1993:200-202) firstproposed that the separation of the origin of LEIfrom those of the other LEs by the origins of LI1and LI2 is a synapomorphy of the Mugilidae andAtherinomorpha. The separation occurs in our spec-imens of Agonostomus, but it appears that it involvesonly LI2.TD comprises TPb2d, TPb2, TEb2, and TPb3.TPb2d. TPb2, and TEb2 originate from mid-longi-tudinal raphe which gives rise dorsally to flimsy CTsheets and is attached anteroventrally to CT of pha-ryngeal roof. TPb2d very broad, broadest anteriorly;attaches on dorsolateral half of IAC; central portionof muscle overlaps most of central portion of TEb2;laterally, beginning at medial surface of LI1 and con-tinuing posteriorly, TPb2d overlaps most of TPb2.TPb2 relatively thick, anterolaterally curving muscleon each side passing dorsal to TEb2 mid-laterally andattaching anteriorly to IAC posteromedially at jointwith Pb2. TEb2 extends laterally onto dorsoanteriorhalf of Eb2 well medial to LE2 insertion. TPb2d,TPb2. and TEb2 not connected posteriorly withTPb3, which is ventral to OD3-4; TPb3 on Pb3 dor-soposteriorly just medial to medial ends of Eb3 andEb4.Remarks. Attachment of TPb2 (including TPb2d)primarily to IAC occurs infrequently in acantho-morphs, but most extensively, and probably homo-plastically, only in Mullidae, Moronidae, and Epi-gonidae.OD3-4 origin on Pb3 dorsomedially ventral toTEb2, insertion on dorsoanterior surface of Eb3 un-cinate process and dorsomedial edge of Eb4 uncinateprocess.OP dorsally on Eb4 posteriorly beginning a littlemedial to uncinate process and extending laterally tobony process supporting LE4 insertion, overlappingLE4 posteromedially; ventrally on bony flange ex-tending posteriorly from Cb5 posterodistally, meetingAd5 posteriorly.Adl absent (small area of GFM spans Ebl-Cbljoint anteriorly).Ad2 begins medially at raphe with lateral end ofTEb2, extends along anterior surface of Eb2 andspreads across Eb2-Cb2 joint.Ad3 begins slightly lateral to anteromedial end of
Eb3, extends laterally along anterior surface of Eb3and spreads across Eb3-Cb3 joint.Ad4 on ventral surface of Eb4 beginning mediallyanterior to OP and extending laterally to near pos-terodistal end of bony edge; ventrally, attaches forequal extent along bony dorsal surface of Cb5 medialto Eb4-Cb4 joint.Ad5 dorsally beginning on cartilaginous distal endof Eb4 posteriorly and extending moderately wellmedially on bony surface, and on distal cartilaginoustip of Cb4 posteriorly; ventrally on distal end of Cb5dorsoanteriorly.Remarks. It is relatively uncommon in acantho-morphs for Ad5 to insert on the bony surface of Eb4.SOD absent in smaller specimen, but present, thinand moderately broad in larger specimen.RDs proximate, insert on Pb3 posteriorly.Remarks. Harrison and Howes (1991:114) statethat RD "in the majority of mugilids" they studied,which included A. monticola, inserts on Pb2. Theydid not mention which mugilids are exceptional orwhere RD inserts in them. They discussed the struc-ture and origin of RD in Agonostomus, but did notmention its insertion. They recognized that RD in-sertion on Pb2 is clearly a specialization in acantho-morphs.Harrison and Howes (1991:126) considered Ago-nostomus to be the sister group of all other mugilids,based on several characters including, among others,the slender structure of LI2 and the fact that each RDis a single muscle, as opposed to paired in most othermugilids. The insertion of RD on Pb3 in Agonosto-mus is additional evidence of the plesiomorphic po-sition of that genus within the Mugilidae.Additional remarks. SCL free from Bb3 (posteriorcartilaginous tip small, not extended posteroventral-ly). TV4 free from Cb5s. Pb4 absent, UP4 present.Pb2 toothed. Eb4 levator process absent.Both Rosen and Parenti (1981:fig. 3, AMNH11613) and Parenti (1993:fig. 5, USNM 73742) in-dicated that Pb4 is present in Agonostomus monti-cola. Of these, the AMNH specimen is missing (B.A.Brown, e-mail, 02/06/2003), but we have seen Par-entis specimens, which are small, but in our opinionlack Pb4, as do our two specimens. The absence ofPb4, a specialization, albeit moderately commonamong acanfhomorphs, is also true of atherinomorphsand might offer support for a relationship betweenthat group and mugilids (Stiassny, 1993); however,Harrison and Howes (1991) reported the presence ofPb4 in juvenile Mugil cephalus, three species of Liza,and Chelon labrosus, but made no mention of its
state in Agonostomus. We examined a few cleared-and stained juvenile specimens of specialized (havemodified Ebl) mugilids (USNM 315896, unidenti-fied) and Mugil cephalus (USNM 156159), and agree
126 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
that they have Pb4. The absence of Pb4 in Agonos-tomus is thus a specialization.A poorly developed bony Eb4 flange is presentdorsolaterally, but is easily overlooked as it onlyslightly overlaps the cartilaginous distal end of Eb4.It is much better developed in the larger specimenand is especially apparent when Eb4 is viewed fron-tally rather than dorsally.AtherinomorphaAtheriniformesATHERINIDAEMenidia peninsulae (Goode and Bean), USNM160465, 2 specimens, 88.0-95.6 mm.Plate 95A, B, D
Additional material. ? = Odontesthes regia (Hum-bolt), USNM 176485, 140 mm; USNM 127863,135 mm.
BEDOTIIDAE
?
= Bedotia sp., USNM 301513, 48.4 mm.Plate 95C
Description.LEI on Ebl at and just lateral to base of uncinateprocess.
?
On uncinate process dorsal to base.
?
Oncartilage tip of uncinate process laterally, extendingonto adjacent bony edge of process.LE2 on tip of expanded bony dorsal edge of Eb2,viewed laterally appears to be two longitudinallyfused muscles: anterior section broader, musculouslyon process; posterior section tendinous ventrolater-
ally, continuing posteriorly as ligament-like lateraledge of CT roofing pharynx and attaching to anterioredge of Eb3 (only CT edge shown in Plate 95.1 A).
?
No apparent longitudinal separation. ? Finely ontip of expanded bony dorsal edge of Eb2, no apparentlongitudinal separation.LE3 slender, on Eb3 uncinate process. ? Insertsby long, slender tendon on tip of uncinate process.LE4 on tip of bony process just lateral to Eb4 un-cinate process. ? Posteroventrally joins raphe withOP dorsally and except for fine attachment antero-laterally, is free from Eb4; forms sling (sensu Stiass-ny and Jensen, 1987).LP narrowly or, usually, broadly on Eb4 bony sur-face well lateral to LE4 insertion, variably joiningraphe ventrally with Ad5 or Ad5 and OP dorsally(forms sling).
?
On most of bony surface of Eb4lateral to LE4 insertion. ? Slightly separated laterallyfrom LE4 insertion; on one side, a muscle filamentcontinues dorsally with OP (here not considered asling).LI1 on Pb2 just medial to attachment of IAC to
Pb2. ? On bony surface of Pb2 well medial to IACattachment.LI2 on Pb3 posterolaterally and ventral to OD4and medial end of Eb3.TD comprises TPb2a, TPb2, TPb3, and TEb4.TPb2a attaching to broad, cartilaginous anterior endof Pb2 ventrally, completely ventral to and parallel-ing anterior portion of TPb2; muscle fibers mesh dor-sally with those of TPb2 just posterior to a horizontalanterior to the separation of the two muscles. TPb2roughly oblong, attaching to, and covering most of,broad cartilaginous anterior end of Pb2 dorsally, withirregular median raphe obscured by, and giving risedorsally to, muscle straps changing to CT (CT notillustrated), which attaches to skull, and ventrallyjoining to TPb2a and ventromedian CT. CT extendsposteriorly from muscle straps and covers Pb3 artic-ulating facets. TPb3 completely separated posteriorlyfrom TPb2 and TPb2a, passing posteroventral to Pb3dorsal articulating facets and attaching to Pb3 dor-soposteriorly medial to LI2 insertion and anterior toPb3 articulation with Eb4 medial end, continuous bydiagonal strand of muscle with TEb4. TEb4 on pos-teromedial surface of Eb4 near dorsolateral end ofOP.
? ?
TPb2 and TPb2a completely fused?no hor-izontal separation mid-anteriorly.OD3 absent.
? Present, see OD4.OD4 origin on Pb3 dorsomedially, beginning an-terior to flat dorsal articulating facet and ending onsurface of Pb3 just ventral to facet; insertion on dor-sal surface and medial edge of Eb4 uncinate process.? OD3-4 present, origin as in Menidia, but insertionincludes anterior surfaces of Eb3 and Eb4 uncinateprocesses.Remarks. Eb3 and Eb4 uncinate processes are ap-pressed and bound to each other by CT, such that inMenidia and Odontesthes it appears that the anterioredge of OD4 insertion includes the medialmost edgeof the Eb3 uncinate process. However, slicingthrough the separation of the two processes indicatesthat all of the insertion is on Eb4.OP dorsally broadly on posterior surface of Eb4.ventrally on posterodistal bony process of Cb5. ? OPventrally broadly joining raphe with OP, which ismostly released from Eb4.Ad 1?3 present, each attaches along most of anter-odistal edge and surface of respective Eb and to an-terodistalmost tip of respective Cb. ? Adl absent.Ad4 dorsally on ventrolateral edge of Eb4, theremeeting OD4 ventrally; ventrally on Cb4 dorsallyimmediately medial to Eb4-Cb4 joint; completely ob-scured from posterior view by Ad5 and OP.Ad5 dorsally on posterodistal surface of Eb4, ven-trally attaching to distal edge and dorsal surface ofCb5 posterodistal bony process.SOD absent.RDs proximate.
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Additional remarks. SCL attached mid-dorsally toposteroventral cartilaginous tip of Bb3. TV4 freefrom Cb5s, which are not ankylosed. Pbl and Pb4absent, UP4 present. Pb2 toothed. Eb4 levator pro-cess absent.
CyprinodontiformesAPLOCHEILIDAE
Rivulus marmoratus Poey, USNM 293487, 62.7 mm.Plate 96
Description.LEI on Ebl dorsodistally (no uncinate process).LE2 on Eb2 dorsodistally.LE3 on Eb3 slightly lateral to uncinate process,insertion continues posteriorly on ligament joiningEb3 and Eb4.LE4 massive, on dorsolateral half of Eb4.LP narrowly on Eb4 at and posterior to lateral endof LE4 insertion.LI1 on Pb2 dorsoanterolateralmost surface just me-dial to IAC attachment.LI2 on Pb3 posterolateral^ anterior to medial endof Eb3.TD comprises TPb2a, TPb2. TPb3. and TPb3p.TPb2a on ventroanterior surface of Pb2 and alongmedial margin of LI1 insertion, continuous with Pb2along mid-longitudinal raphe, which gives rise to CTpad attaching to skull. Ventral surface of raphe atta-ches to CT sheet covering Pb3 articulating facets.TPb2 anteromedially joins longitudinal raphe andmedially joins CT sheet covering articulating facets;laterally, TPb2 attaches to Pb2 dorsoanteromedialsurface, dorsomedial to LI1 insertion; ventromedial-ly, TPb2 joins CT junction with OD4 dorsoanteriorly,and posteromedially joins CT junction with TPb3 an-teriorly (CT junction attaches to Pb3). Median lon-gitudinal raphe weakly represented on semicircularTPb3, which is weakly attached mid-ventrally to CTbetween Pb3s, but is unconnected to TPb3p. Ante-roventrally, muscle fibers of TPb3 mesh with thoseof OD4. TPb3p finely, tendinously attached to pos-terior edge of Pb3 slightly medial to joint with Eb4;SO fibers join tendinous attachment.OD3 absent.OD4 originates anteriorly ventral to TPb2 on dor-soanterolateral surface of Pb3 dorsal articulating fac-
et (joining CT there with TPb2), and posteriorly ven-tral to TPb3 on CT covering facet, there joining withTPb3 anteriorly; inserts on broad mid-dorsal shelf onEb4.OP dorsally very broadly on Eb4 posterior surface,ventrally on posterodistal flange of Cb5, laterally es-sentially continuous with Ad5.Ad 1?3 very broadly on respective Eb, spanning
respective joint with Cb anteriorly and continuingwith CT supporting strip with gill rakers.Ad4 very broadly on ventrolateral surface of Eb4.and equally so on dorsal surface of Cb4, completelyobscured from view posteriorly by OP and Ad5 (onlyvisible in lateral view if obscuring CT and gill rakersare removed).Ad5 medially inseparable from OP, anterolaterallyattaching to distal ends of Eb4 and Cb4, ventropos-teriorly to distal end of Cb5.SOD absent.RDs proximate.Additional remarks. SCL band-like (as opposed tomore typical threadlike in other fishes), attached mid-dorsally to cartilaginous ventroposterior tip of Bb3.TV4 attached dorsally to Cb5s (continuous ventral-ly). Pbl and Pb4 absent, UP4 present. Eb3 and Eb4uncinate processes present. Eb4 levator process ab-sent. CYPRINODONTIDAECyprinodon variegatus Lacepede, USNM 107023, 3specimens, 48.1-49.0 mm.Plate 97
Description.Remarks. Assignments of LEI and LEI'. LE2 andLE2', and LE4 and LE4' are arbitrary designations.LEI on Ebl just lateral to articulation with IAC,joins raphe with Adl along anterior margin of inser-tion.LEI' on Ebl slightly lateral to LEI insertion.LE2 on posterodistal end of Eb2.LE2' laterally appressed to LE2 on posterodistalend of Eb2. Fine ligament extends from posterioredge of LE2 and LE2' to distal end of Eb3.LE3 absent; possibly represented by LE4'; see dis-cussion in Additional remarks section.LE4 on Eb4 dorsally lateral to dorsalmost point.Remarks. LE4 and LE4' appear to be one massivemuscle before mechanical separation, but separationpoint was same in all three specimens.LE4' on Eb4 at and lateral to LE4 insertion; veryfine ligament connects tip of Eb3 uncinate processwith LE4' insertion; see discussion in Additional re-marks section.LP on dorsodistalmost end of Eb4.LI1 on cartilaginous dorsoanteriormost tip of Pb3;in one specimen a few muscle filaments attach todorsoanteriormost tip of Pb2 on one side and to IACon the other.LI2 on Pb3 dorsolaterally.TD comprises TPb2a, TPb2, and TPb3-Eb4.TPb2a on anterior surface of dorsoanteriormost Pb2process, joined to TPb2 along mid-longitudinal ra-phe, which gives rise dorsally to CT pad coveringmuscles. TPb2 on posterolateral surface of Pb2 dor-
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soanteriormost process; TPb2 and TPb2a completely,although only narrowly separate from each other onlateral surface of process. TPb3-Eb4 attachment, ten-dinous, very fine, unclear, appearing variably to at-tach at Pb3-Eb4 joint, only to Pb3, or only to Eb4.OD3 absent.OD4 originates on medial surface of Pb3 begin-ning musculously anteriorly on lateral surface of dor-soanterior process becoming CT posteriorly and at-taching to groove ventral to articulating process; in-sertion splits distally with anterior branch on Eb4 un-cinate process and posterior branch on Eb4 dorsalsurface medial to LE4 insertion.OP dorsally broadly on Eb4 posterior surface, ven-trally on dorsodistal surface of Cb5; CT medial toOP covers Eb4-Pb3 joint.Adl with two continuous portions, dorsally on me-dial end of Ebl at anterior edge of LEI insertionventrolaterally, passing over Ebl-Cbl joint and at-taching to dorsoanterior surface of Cbl, continuousdorsally with ventral surface of crossing portion,which attaches on Ebl medially and to gill-raker andgill-filament strip laterally.Ad2 with two continuous portions, ventral portionattaching along most of dorsal surface of Eb2 passingover Eb2-Cb2 joint and attaching to Cb2; dorsalcrossing portion beginning at medialmost end of Eb2and extending laterally and attaching to gill-raker andgill-filament strip.Ad3 with two continuous portions, ventral portionon Eb3 anteromedial surface and Eb4 ventroanteriorsurface, extending ventrolaterally and attaching todorsoanterior surface of Cb3; dorsal portion attachingto most of medial arm of Eb3, extending laterally andoverlapping and joining dorsal surface of ventral por-tion and attaching to gill-raker and gill-filament strip.Ad4 dorsally on ventral surface, ventrally on dor-
sal surface of Cb4 medial to Eb4-Cb4 joint; obscuredin posterior view.Ad5 on posterodistal end of Cb4 and anterior sur-face of Cb5 well medial to distal end.Additional remarks. SCL attached mid-dorsally toventral surface of cartilaginous posterior end of Bb3.TV4 absent. Pbl, Pb4, and UP4 absent (or UP4 fusedto Pb3, according to Parenti 1981:417, but treated byus as absent. Unless it can be shown, perhaps onto-genetically, that UP4 fuses to Pb3 in some cyprino-dontids, it is equally parsimonious to treat UP4 asabsent as it is to treat it as fused to Pb3). Pb2 toothed.Eb4 levator process absent.
BeloniformesADRIANICHTHYIDAEXenopoecilus oophorus Kottelat, USNM 340431, 2specimens, 57.4?69.3 mm.Plate 98
Additional material. ? = Oryzias latipes (Temminckand Sclegel), 2 specimens, 29.0-29.4 mm.
Description.LEI on dorsal end of IAC, which is attached main-ly to dorsolateral end of Eb 1 and less so to dorsolat-eral end of Cbl (see also remarks following LCb2).
? IAC absent, LEI attached to lateral end of Ebldorsally.Remarks. IAC typically joins Ebl and Pb2 in acan-fhomorphs. This is also true of IAC in atheriniformssuch as Cyprinodon (Plate 97), which, similar to ad-rianichthyids, has lost the Ebl uncinate process. Itseems a relatively short transition to change, for ex-ample, from the cyprinodontid state to that of theadrianichthyids (compare Rosen and Parenti, 1981:figs. 10 and 1 1, who recognized the states of IAC inboth groups).LE2 on Ebl dorsodistally medial to LE2', verti-cally or slightly anteriorly directed toward origin.LE2' on Ebl dorsodistally lateral to LE2, dorso-posteriorly directed toward origin.LCb2 on dorsal end of dorsally autogenous carti-laginous end of Cb2. ? On dorsal end of Cb2.Remarks. The cartilaginous dorsoanteriormostends of Cb2, Cb3, and Cb4 are greatly expanded inadrianichthyids and vary from continuous to autog-enous, and that of Cb4 may consist of several sepa-rate pieces of cartilage. It is also possible that AC1(see LEI above) represents an autogenous piece ofCbl, but its position relative to the Ebl-Cbl jointdiffers from those of Cb2-4.LE3 absent (see remarks following LCb5).LE4 on bony dorsoposterior surface of Eb4.LP on dorsodistal end of Eb4 and autogenous car-tilaginous end of Cb4 (not visible in Plate 98), fusingventromedially with LCb5 laterally and with Ad5ventrolaterally. ? On Eb4 mid-dorsally and cartilag-inous distal end of Cb3 dorsally. See remarks follow-ing LCb2.LCb5, as it extends toward Cb5, attaches antero-laterally to posterolateral surface of distal end of Eb3(attachment released in Plate 98A, C), joins anter-oventromedialmost surface of LP on Eb4 and pos-terodistalmost surface of Cb4 just lateral to ventralattachment of Ad4, and has major portion of insertionon Cb5 dorsoanterodistalmost surface anterior to Ad5(origin was not recorded).Remarks. Aside from these two genera, we ob-served LCb5 otherwise only in the ostariophysancyprinids, in which the muscle inserts exclusively onCb5. The homology of LCb5 is problematic, and themuscle possibly represents a highly modified LE3.LI 1 on dorsoanterior tip of Pb2 posteriorly.LI2 broadly on Pb3 dorsoposterolaterally.TD comprises TPb2, TPb2a and TPb3. TPb2 hasmedian longitudinal raphe and attaches to Pb2 dor-
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soanterior process dorsal to TPb2a. TPb2a attachesto anteroventral edges of Pb2 dorsoanterior processand is continuous posterodorsally with TPb2. TPb3attaches to Pb3 in groove ventral to Pb3 articularsurface with Eb4.OD3 absent.OD4 originates on Pb3 dorsoanteromedial surfaceventral to TPb2 and inserts on bony Eb4 dorsal sur-face.OP dorsally on Eb4 posteroventral surface lateralto medial end and medial to Ad5 dorsomedially, ven-trally on Cb5 dorsodistally, only weakly separatedfrom Ad5 laterally.M. Ebl-IAC very fine, on Ebl dorsolaterally andlateral surface of IAC ventral to LEI insertion.Adl absent, but long GFM present as in secondarch.Ad2 broadly on Eb2 anterior surface and CT be-tween Eb2 and autogenous Cb2, and narrowly on,Cb2 dorsoanteriorly lateral to GFM dorsally.Ad3 very broadly on Eb3 anterior surface and CTbetween Eb2 and autogenous Cb3, and narrowly onCb3 dorsoanteriorly lateral to GFM.Ad4 dorsally on Eb4 ventral to OD4 insertion,ventrally on Cb4 dorsodistally; muscle broad dorsal-ly, very narrow ventrally.Ad5 dorsoposteriorly on Cb4 distally and Eb4 be-ginning lateral to OP and extending to distalmostend, there joining fine raphe with ventrolateralmostedge of LP, ventrally on Cb5 distally. ? Unclear ifAd5 joins LP.SOD absent.RDs well separated.Additional remarks. SCL attached dorsomedianlyto cartilaginous ventroposterior end of Bb3. TV4 freefrom Cb5s. A strap of SO extends anteriorly just lat-eral to RD and inserts in a deep bony pocket in Pb3ventrolateral to LI2. Pbl, Pb4, UP4, and Eb4 levatorprocess absent. Pb2 toothed.
BELONIDAE
Tylosurus crocodilus (Peron and Lesueur), USNM128454, ca. 345 mm; USNM 137545, not mea-sured, USNM 294990, ca. 170 mm.Plate 99
Additional material. ? Strongylura timucu (Wal-baum), USNM 203575, ca. 225 mm.
Description.LEI on dorsal edge of bony process projecting me-dially from enlarged lateral end of Eb 1
.
Remarks. Based on the configuration of the lateralend of the closely related scomberesocid, Cololabis(Plate 100), which has a cartilage-tipped uncinateprocess, we consider that the uncinate process of be-
lonids has lost the cartilage tip, hence, would not berecognized as an uncinate process.LE2 dorsally bifurcate, on posteriorly extendingbony process on Eb2.
? Not bifurcate dorsally.LE3 absent (see remarks following LE4).
?
On tipof all bony Eb3 uncinate process.LE4 (left side aberrant in illustrated specimen) onEb4 more-or-less mid-dorsally continuing muscu-lously anteriorly and then inserting on CT attachingEb4 to tip of all bony Eb3 uncinate process, insertionmeeting LP insertion posteromedial edge. ? On Eb3slightly lateral to bony uncinate process, continuingonto juxtaposed Eb4, meeting LP insertion ventro-medially.Remarks. LE4 in Tylosurus appears to have ex-tended its insertion a short distance anteriorly to in-clude the tip of bony Eb3 uncinate process. The typ-ical position of LP, at and lateral to the levator onEb4 lends support to identification of the muscle in-serting on Eb3 as LE4.LP on Eb4 dorsodistalmost bony surface at andlateral to LE4 insertion. ? Probably present, see
?in LE4.LI1 on Pb2 dorsoposteromedially.LI2 relatively massive, on Pb3 dorsoposterolater-
ally.TD complex, TDA comprising TPb2, TPb2',TPb3, and TDP comprising TEb4. TPb2 (see remarksfollowing this description) attaches to anterior end ofPb2 dorsoanterior process, wraps around process, andforms raphe with itself along mid-anteroventral sur-face of process; anteriorly. TPb2 extends ventrome-dially and joins a narrow raphe-like area of CT thatexpands posteriorly into a broad sheet that attachesto ventral surface of skull; dorsoposteriorly, TPb2forms a flat, shallow, thin layer (barely separate fromremainder of muscle), edged medially by CT, whichis continuous with broad CT sheet attaching to skull;TPb2 covers most of TPb2' and TPb3 (including dor-soanterior process of Pb3). and all three muscles joinalong median CT raphe-like area. TPb2' a thin, broadband passing dorsal to TPb3 and conforming with it;laterally, TPb2' attaches to Pb2 posterolaterally.TPb3 attaches to most of surface of Pb3 anteriorly,forming raphe posteriorly with OD4 on left side, andpartial raphe on right side. TEb4 disconnected fromremainder of SO, on Eb4 posteromedially, posteriorlycontinuous with SOD. Pb3's dorsoposteriorly arecompletely ventral to muscle, no articulating facetsare exposed.
? TPb2' almost completely fused with TPb2(TPb2' would not be recognized as such, lackingcomparison with Tylosurus); TPb3 posteriorly atta-ches by CT to dorsally exposed Pb3 articulating fac-
ets, with CT continuing around facets laterally andbecoming musculous OD3-4 anteriorly. TDP com-prises TPb3-Eb3-Eb4 attaches to Pb3 a little medial
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to medial end of LI2 continues posteriorly attachingto medial end of Eb3 and broadly along medial armof Eb4.Remarks. TPb2 in both species is possibly a fusionof TPb2 and TPb2a, which are separate in other ath-erinomorphs.M. Pb2-Eb2 on Pb2 dorsolateral^ beginning justposterior to dorsoanterior process and on Eb2 anter-omedially, laterally attaching to CT between Ebl andEb2. ? Muscle forks immediately anterior to its at-tachment to medial end of Eb2; arms of fork pass toeither side of LI1 and attach to separate areas, lateraland medial, on Pb2.OD3 absent. ? OD-3-4 origin by CT on Pb3 dor-
sal articulating fact, insertion on Eb3 just medial to
all bony uncinate process (very fine cartilage tip) andbroadly on Eb4 dorsally medial to LE4-LROD4 origin dorsoanteriorly joining raphe withTPb3 posteriorly, area below raphe attaching broadlyalong Pb3 medially; muscle in two incompletely sep-arated parts, lateral part passes medial to bony Eb3uncinate process and inserts on Eb4 dorsally medialto LE4-LP insertions, medial part inserts on Eb4 wellmedial to lateral part. Eb4 levator and uncinate pro-cesses absent (both taxa).Remarks. See similarity of insertions to those ofCyprinodon (Plate 97).OP dorsally on Eb4 posterolateral^, ventrally onCb5 posterolateral^, fusing dorsoanteriorly with Ad4posteroventrally.Adl absent.Ad2 dorsally with anterior branch (GFM2?) at-taching to dorsodistal end of Ebl and broader attach-ment to anterodistal end of Eb2, and ventrally at-taching to Cb2 dorsally.Ad3 like Ad2 but anterior branch (GFM3?) onEb2, remainder on Eb3 and Cb3; posterior branchextends medially almost to medial end of Eb3.Ad4 attaches to Eb4 ventrally anterior to OP, pos-teroventrally joins anterior surface of OP and attachesto Cb4 dorsoanteriorly.Ad5 apparently absent.SOD present.RDs well separated.Additional remarks. SCL attached mid-dorsally bylong tendon to posteroventral cartilaginous tip ofBb3. TV4 free from Cb5s. ? Dorsally attached nar-rowly to Cb5 anteriorly; ventrally free. Pbl, Pb4, andUP4 absent. Pb2 toothed. IAC absent.AC1 present between Ebl and Cblon both sidesof USNM 128454 and 294990, and latter has small,questionable AC4 on both sides associated with distalend of Eb4, which is well dorsal to distal end of Cb4(see Additional remarks in Cololabis on why this ACis not considered to be AC4); USNM 137545 lacksACs on all arches. Strongylura timucu has AC1 onboth sides, a tiny AC3 on one side, and a question-
able AC associated with Eb4 on one side. It also has.on both sides, a small disjunct cartilage associatedwith the distal (or dorsolateral) end of Cb2 and an-other with the distal end of Cb3. These two cartilagesappear to be fragments of the cartilaginous distalends of Cb2 and Cb3. The cartilages lie adjacent tothe anterior bony distal ends of these two elements,each of which bears a cartilage tip posteriorly (AC3is positioned between the cartilaginous ends of Eb3and Cb3).Bony flange on Eb4 distally, only slightly or not
at all extending over cartilaginous distal end of Eb4.hidden by LE4-LP insertions.SCOMBERESOCIDAECololabis saira (Brevoort), USNM 320999, 3 speci-mens, ca. 174?220 mm.Plate 100
Description.LEI beginning on cartilaginous tip of Ebl uncinateprocess and extending about half way along mediallyextending tendon attaching to medial end of Ebl(free from most of anterior arm of Ebl).LE2 on dorsally raised bony process on Eb2; lat-eral fibers continuous ventrally with anterior branchof Ad3.LE3 absent (see LE4).LE4 tendinously on Eb4 dorsally medial to distalend, tendinous insertion passes dorsoanteriorly and isapplied closely on cartilaginous tip of Eb3 uncinateprocess.LP on Eb4 dorsodistally, posterolateralmost fibersventrally variably continuous or not with OP dorsal-ly.Remarks. Although not as clearly forming an LP-OP sling (Stiassny and Jensen, 1987) as occurs inexocoetoids, the condition is, nevertheless a sling,which Stiassny and Jensen (1987:284) state is notpresent in scomberesocids; however, for this remark,they reference their fig. 2D, which is a belonid, whichfishes, as opposed to scomberesocids, do not have asling.LI1 on Pb2 dorsally near base of anterodorsallyextending Pb2 process.LI2 on Pb3 dorsoposteriorly.TD comprises TPb2a, TPb2, TPb3a, and TPb3p.TPb2a attaches along anterolateral edge of the longdorsoanterior Pb2 process and wraps anteriorlyaround process, becoming continuous with TPb2ventromedially where they join CT of pharyngealroof. Anterolaterally, TPb2 attaches to dorsal and me-dial surfaces of long dorsoanterior process of Pb2 andposterolaterally to dorsoposterior surface of Pb2, justextending onto anteromedialmost end of Eb2; alongits long ventromedial length, TPb2 joins CT of mid-line between two sides of gill arches, to which con-
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tralateral TPb2 also joins, and together join a sheetof CT lying dorsal to the flat, bony dorsoposteriorfacet on each Pb3; anteromedial extent of TPb2 com-pletely overlies TPb3a. Laterally, TPb3a attaches tomuch of dorsal surface of Pb3, including long dor-soanterior Pb3 process, and medially attaches (to-gether with TPb2) to CT of mid-line between twosides of gill arches; posteriorly, TPb3a continues asCT that attaches to and covers bony dorsoposteriorPb3 facet on each side. TPb3p short, joins Pb3s dor-soposteriorly, forming raphe there with SO, contin-uous posteriorly with SOD.M. Pb2-Eb2 anteriorly on ventroanterolateral sur-face of Pb2 dorsoanterior process, posteriorly on me-dial end of Eb2.OD3 absent.OD4 originating tendinously from CT coveringand attaching to dorsal surface of Pb3 medial to dor-soposterior facets, passing dorsal to flat, bony facetsand most of surfaces of Eb3 and Eb4 and attachingseparately to Eb4 dorsoposteriorly medial to LP andanteriorly near medial end. Eb4 uncinate process ab-sent.OP dorsally on most of lateral half of Eb4 poste-
riorly, overlapping Ad4 and Ad5 posteriorly and ex-cluding them completely in posterior view; postero-dorsal fibers variably continuous or not with LP pos-teroventrally; ventrally on dorsodistal end of Cb5. Asplay of fibers attaches to the posterolateral surfaceof Cb5, passes anterior to OP ventrally and appearsto join OP dorsoanteriorly in some specimens and/orAd5 posteromedially in others. Based on the attach-ment of OP to Cb5 in belonids. these fibers appearto be OP.Adl absent.Ad2 with dorsoanterior branch (= GFM2?) attach-ing to lateral edge of Ebl uncinate process, continu-ing posteriorly, spanning area between first and sec-ond arches and attaching (Ad2) to ventral surface ofEb2 and dorsolateral surface of Cb2 anterior to Eb2-Cb2 joint.Ad3 like Ad2 but on Eb3 and Cb3, and anteriorbranch (= GFM3?) attaching to raised bony processon Eb2 to which LE2 inserts.Ad4 dorsally on dorsoposterodistalmost surface ofEb4 ventral to OP, extending medially on Eb4 ante-
rior to OP, laterally joining distal ends and surfacesof Eb4 and Cb4 and fusing with dorsal end of Ad5on Cb4.Ad5 dorsally on Cb4 posterodistally, fusing therewith Ad4, extending ventrally and expanding as itattaches to posterodistal surface of Cb5.SOD present.RDs well separated.Additional remarks. SCL poorly defined; musclescrowded in region and attached to CT, which is at-tached mid-dorsally to very elongate posteroventral
cartilaginous tip of Bb3. TV4 ventrally free fromCb5s. but dorsally attaching narrowly, weakly to an-terior end of Cb5 (Cb5 a single bone). Pbl, Pb4, andUP4 absent. IAC absent. Two specimens have tinyAC1 bi-laterally, and one of these has very fine,somewhat rod-shaped cartilage between lateral endsof Eb4 and Cb4 on both sides; the third specimenlacks ACs and the cartilage between Eb4 and Cb4.It is doubtful that the cartilage between Eb4 and Cb4in Cololabis and belonids is homologous with AC4of other fishes. The cartilage in the former twogroups is close to the distal end of Eb4 and wellseparated from the distal end of Cb4. In belonids, thecartilage appears to have budded off Eb4, whereas,as far as is known, AC4 buds off the distal end ofCb4 in non-atherinomorphs.Cololabis is the only atherinomorph we examinedthat appears to lack an Eb4 flange.HEMIRAMPHIDAEHemiramphus far (Forsskal), USNM 294231, 2 spec-imens, 105-1 1 1 mm.Plate 101EXOCOETIDAE
Additional material. ? = Exocoetus obtusirostrisGiinther, USNM 198465, 92.8 mm; USNM295036, 161 mm; USNM 298987, 98.8 mm.
Description.Remarks. The muscles of Exocoetus appear to bemore specialized than those of Hemiramphus, al-though comparing muscles of the latter with those ofthe former, made it possible to identify most of them.We abbreviated descriptions of Ads, LEs, and slingof Exocoetus.LEI tendinously on bony and cartilaginous tip ofEbl uncinate process. ? Tendinously and muscu-lously on dorsalmost tip of expanded lateral end ofEbl, continuous posteroventrally with CT joiningEbl and Eb2 dorsodistally; uncinate process absent.LE2 fan-like or split dorsally, incompletely divis-ible into anterior and posterior sections, which fuse
at insertion on bony process near distal end of Eb2.
?
Similar but inserts on dorsodistal bony edge ofEb2.Remarks. Small, cartilaginously tipped uncinateprocess on Eb2 medial to bony process in both spec-imens (absent in Exocoetus). Among ctenosqua-mates, examined, a cartilage tipped Eb2 uncinate pro-cess is otherwise present only in some specimens ofPholidichthys (Pholidichthyidae).LE3 slender, inserts by long, slender tendon con-fluently with fine ligament joining Eb3 uncinate pro-cess with bony dorsolateral edge of Eb4 directly pos-terior (medial to Eb4 uncinate process); confluence
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is usually on bony edge of Eb4 but may be on liga-ment mid-way between Eb3 and Eb4; musculous por-tion united with anterior surface of LE4, but easilyand discretely separated. ? Musculous portion ofLE3 less easily separated from LE4.LE4 massive, mostly free from Eb4, but narrowlyattached medially to posterior surface of Eb4 unci-nate process at dorsolateral edge of OP: continuesventrally becoming tendinous, with OP joining ten-don dorsoanteriorly and Ad5 posteromedial surfacejoining tendon laterally, and together continuing ten-dinously and inserting on Cb5 dorsodistally (Ad5 at-tachment continues musculously on dorsomedialedge of Cb5); muscle variously, broadly forked ven-tral to origin.Remarks. Stiassny and Jensen (1987) first notedthe LE4-OP sling of exocoetoids and its similarity tothat of their labroids; however, the exocoetoid LPdoes not participate in the sling in the same way itdoes in some labroids.LP on dorsolateral surface of Eb4 and dorsopos-terior edge of Cb4, relatively distinct and separatefrom LE4. ? Muscle joins LE4 insertion laterally andwraps closely around LE4 posteromedially.LI1 on ventrolateral surface of elevated Pb2 an-terior process.LI2 on Pb3 dorsolaterally, posterior portion of in-sertion passing ventral to OD4 anteriorly. ? On Pb3posterolateral^.TD complex, comprising TPb2, TPb2a, TPb2v,TPb3, and TEb4. TPb2 comprises separate muscle oneach side, inserting broadly along ventral surface ofcranium (muscle is considerably truncated in Plate101 ); each side has broad fascia attachment on dorsalPb2 process dorsoanteriorly and muscle generallycovers TPb3, obscuring it dorsally. TPb2a completelyseparate from TPb2, attaches to medial edges of dor-sal Pb2 processes; muscle may represent a separationfrom TPb2, which wraps around Pb2 in various ath-erinomorphs (see Tylosurus, Plate 99). TPb2v super-ficially appears to be bilaterally paired muscle, butfibers fuse across ventral midline; anteriorly muscleattaches to skull, posteriorly attaches to ventroanter-ior edge of Pb2. TPb3 joins ventroanterior surfacesof Pb3 anterior processes (Pb3s are fused, but deeplycleft, ventromedially), has median longitudinal raphe.TEb4 arising on each side from lateral edge of CTsheet, passing dorsal to posterior surfaces of Pb3apophyses, and joining opposite TEb4; muscle insertson Eb4 uncinate process immediately posterior andessentially continuous anteriorly with OD4. SO mus-cle strap from each side joins edge of CT sheet pos-terior to TEb4 posterior edge.
? Composition same, muscles differently dis-posed; TPb3 and TEb4 are unpaired. TPb2 much lessextensive, attaches dorsolaterally on Pb2 dorsoanter-ior process, extends membranously around Pb2 and
Pb3 anterior processes encapsulating fatty and/orglandular mass of tissue; thins and fans out posteri-orly and overlaps TPb3, extends ventromedially andvery weakly attaches along midline between Pb3sand possibly dorsoposteriorly to cranium. TPb2a welldeveloped, triangular, apex attaching to dorsomedialsurface of Pb2 dorsoanterior processes, extends ven-tromedially toward midline and attaches to skull.TPb2v originates on ventrolateral surface of Pb2 dor-soanterior process and extends anterolaterally to an-teromedial end of Ebl, to which it is weakly attachedby CT; posteromedially weakly attached to skull.TPb3 a narrow band of undivided muscle with mid-longitudinal raphe, situated in area between posteriorends of bases of Pb3 dorsoanterior processes and Pb3posterodorsal articulating facets; lateral ends extendanteriorly as CT and attach to Pb3 dorsoanterior pro-cess. TEb4 interrupted at lateral edge of Pb3 dorso-posterior facet but continuous with CT across top offacets with opposite TEb4, muscle uninterrupted asit passes posterior to Pb3 posterodorsal articulatingfacets, attaches laterally to distal ends of Eb4s.OD3 absent.OD4 originates broadly on Pb3 dorsal surface lat-eral to anterolateral edge of Pb3 dorsoposterior facetand inserts on Eb4 uncinate process. ? Originates onPb3 posterodorsally anterior to articulating facet andmedial to LI2 insertion; inserts on Eb4 uncinate pro-cess.OP narrow muscle strap on posterolateral surfaceof Eb4, joining LE4 ventrally (see LE4 description,also remarks following LE4), extending ventrally andinserting on Cb5. ? Similar, but free portion dorso-laterally continuous with "sling."Adl absent.Ad2 on entire ventral edge of Eb2 and dorsoan-terior edge of Cb2; small IAC associated with dor-soanterior edge of muscle. ? Present, IAC similar.Remarks. Rosen and Parenti (1981 :fig. 12) showIAC in exocoetoids as extending between Ebl andEb2. The size of IAC in our specimens varied froma small sphere to a moderate-sized rod similar to thatin Rosen and Parenti (1981:fig. 12A).Ad3 bipartite, anterior branch on tip of Eb2 dorsalbony process and Cb3 dorsal surface where it joinsventral portion of posterior section, which is on mostof Eb3 anterior surface.
? Not bipartite (anterior por-tion absent).Ad4 on ventral edge of Eb4 and dorsoposterioredge of Cb4 medial to Eb4-Cb4 joint. ? Not clearlypresent; possibly fused in sling.Ad5 dorsally on Cb4 posterodistally, medial sur-face fusing with LP sling ventral tendinous end andattaching to Cb5 dorsodistally.SOD absent.RDs well separated.M. Pb3p, cone-shaped, narrowly joined to contra-
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lateral M. Pb3p anteroventrally at insertion on Pb3sposteroventral to Pb3 dorsal apophyses; cones areconfined dorsally, but are free from, conforming con-cave bony roof formed by posterior end of skull, andare restricted ventrally by RDs and viscera; dorsallyeach cone has thin, traversing ribbon of muscle.Additional remarks. SCL attached mid-dorsally tocartilaginous ventroposterior tip of Bb3. TV4 inter-rupted medianly and attached to ventrolateral edge ofCb5 keel. Pbl, Pb4, and UP4 absent. Eb3 and Eb4uncinate processes present. Eb4 levator process ab-sent. Tiny AC1 present or absent, AC2 present. ?AC1 present or absent, AC2 absent.
PerciformesACROPOMATIDAESynagrops bellus (Goode and Bean), USNM 359306,108 mm, USNM 186122, 124 mm.Plate 102Description.LEI with tendinous origin; insertion broadly onhigh Ebl uncinate process, beginning near tip andextending ventroanterolaterally almost to base.Remarks. Ligament originates on skull immediate-ly anterior to origin of LEI and inserts on Ebl an-terior to LEI insertion; ventrolaterally, ligament iscontinuous with low CT sheet that attaches along al-most entire dorsolateral edge of raised anterior mar-gin of Ebl.LE2 on raised posterior margin of Eb2 at aboutmid-length.LE3 on tip of Eb3 uncinate process.LE4 on tip of Eb4 levator process anteriorly.LP slender, on Eb4 at and lateral to LE4 insertion.LI1 tendinously on dorsoanterior Pb2 process pos-teriorly.LI2 on Pb3 posterolaterally at joint with Eb3, me-dial edge of insertion meeting lateral edge of TPb3-Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3; cov-ered dorsally by tough CT sheets, which attach alongentire posterior surface of Pbls, continuing mediallyalong anteriormost edge of Pb2s to mid-line of CTof pharyngeal roof, thence continuing posteriorly onmid-line raphe joining TPb2 and TEb2 posteriorly.TPb2 flat, thin, broadly V-shaped, with, narrow, an-terior mid-line notch deeply separating arms of V;muscle completely underlain by TEb2. TEb2 flat,very broad medially, narrowing considerably laterallyand attaching on Eb2 dorsally anterior to LE2 inser-tion; muscle continuous posteriorly by fine musclestrands with TPb3-Eb3. TPb3-Eb3 anteriorly broadlyon Pb3 dorsally ventral to OD3-4. posteriorly at-taching along medial edge of LI2 insertion and on
posteromedial cartilaginous end of Eb3 dorsally(passes dorsal to medialmost end of Eb4 without at-taching); muscle narrows posteriorly and is continu-ous by crossing muscle strands with SOD.OD3-4 origin on Pb3 dorsomedially ventral toTEb2, insertion mainly on Eb3 anterior surface be-ginning just ventral to tip of uncinate process andextending medially; lesser insertion on anterior edgeand surface of Eb4 just ventral to tip of uncinateprocess.OP dorsally on Eb4 posteriorly beginning on me-dialmost bony surface and extending laterally to orjust lateral to uncinate process, laterally overlappingAd4 dorsomedially; ventrally on Cb5 dorsoposterior-ly, beginning medially at junction of slender "horn"of Cb5 with expanded dentate portion and continuingalmost to distal tip of Cb5, there joining raphe withAd5 posteromedially.Ad 1-3 absent (GFM1 well developed; GFM2much reduced; GFM3 absent).Ad4 dorsally on Eb4 posteriorly, beginning me-dially anterior to OP and extending to posterodistal-most bony surface; ventrally on Cb4 dorsoposteriorlya short distance medial to distal end and extendinglaterally almost to distal end, there joining raphe withAd5 dorsomedially.Ad5 dorsally on Cb4 posteriorly beginning ventralto medial end of Ad4 and extending laterally almostto distal end of Cb4, there joining raphe with Ad4ventrally; ventrally on Cb5 dorsally anterior to OPattaching for distance paralleling attachment on Cb4.joining raphe with OP ventrolaterally.SOD present.RDs moderately slender, separated by distanceabout equal to about half diameter of one RD.Additional remarks. SCL weakly attached mid-dorsally to elongate, posteroventrally extending car-tilaginous tip of Bb3. TV4 free from Cb5s. Pb4 andUP4 present. IAC present. Tiny AC4 present; rela-tively well developed in larger specimen.Sasaki (1989:fig. 2A) partially illustrated the dor-sal gill-arch musculature of Acropoma japonicumGunther, which appears to differ from Synagrops bel-lus in that TPb2 is apparently broadly kidney-shapedrather than V-shaped.PERCICHTHYIDAEMacquaria colonorum (Gunther) USNM 59968. 125mm. Plate 103Description.Remarks. All LEs originate tendinously.LEI on anterior surface of Ebl uncinate processventrally.LE2 on dorsoanterior surface of Eb2 elevated bonyposterior ridge.
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LE3 on Eb3 uncinate process dorsoanteriorly withCT extending ventromedially from insertion joiningOD3?4 posterolaterally on Eb4 uncinate process.LE4 on bony edge of Eb4 levator process just me-dial to cartilage tip, ventrolateral edge joining raphewith OP.LP at lateral edge of LE4 insertion, extending ontocartilage tip of Eb4 levator process.LI1 on Pb2 dorsoanteriorly; anterior edge joiningraphe with TPb2 on Pb2 just medial to joint withIAC.LI2 on Pb3 dorsal surface at and lateral to TPb3-Eb3 attachment and medial to medial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2overlies TEb2, beanshaped, notched mid-anteriorly,attached to anterior cartilaginous tip of Pb2 near jointwith IAC, joining raphe with ventromedial edge ofLI1; muscle with mid-longitudinal raphe, anteriorend of which attaches to CT of pharyngeal roof andto Pb3 anterior end dorsally; raphe giving rise dor-sally to CT mass covering muscle, ventrally joiningTEb2, posteriorly continuing across middle of TEb2.TEb2 attaching laterally along medial half of Eb2dorsal surface and lateral to LE2 insertion, continu-ous posteriorly by slender, diagonal strap of musclewith anterior end of TPb3-Eb3. TPb3-Eb3 on Pb3medial to medial edge of LI2 insertion and dorsalsurface of medial end of Eb3, continuous posteriorlyby crossing muscle straps with SOD.OD3?4 on dorsoanterior surface of Pb3, insertionon anterior surface of Eb3 uncinate process and an-terior surface and medial edge of Eb4 uncinate pro-cess.OP broadly on most of Eb4 posterior surface join-ing raphe with ventrolateral edge of LE4 insertion,ventrally, narrowly on posterodistal surface of Cb5,medially not clearly separable from SO; bilaterallyasymmetrical, left side appearing to comprise twostraps of muscle, right side only one.Adl-3 absent.Ad4 dorsolaterally on Eb4 levator process poste-
rior surface lateral to OP, and medially on Eb4 ventralsurface anterior to OP, ventrally on dorsoposteriorsurface of Cb4 medial to Eb4-Cb4 joint and anteriorto Ad5.Ad5 on dorsal surface of Cb5 distally anterior toOP attachment and on Cb4 posterodistal surface pos-terior to Ad4 attachment, with dorsodistal portion ofmuscle continuous with tough CT (not illustrated) at-taching broadly to AC.SOD present.RDs separate, adjacent.Additional remarks. SCL attached mid-dorsally toposteroventral cartilaginous tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. Pb2 toothed. AC4present.
LEPTOBRAMIDAELeptobrama muelleri Steindachner, WAM P.556-001,195 mm. Plate 104
Description.LEI shortest levator; tendinously and musculouslyon Ebl beginning just lateral to tip of horizontallydirected uncinate process and continuing mediallyonto tip and on to IAC dorsolaterally. LEI and Ebltightly attached anteriorly to CT lining gill chamber.LE2 on tip of dorsally projecting peg-like processarising from Eb2 dorsoposteromedially; ventrolateralsurface continuous with CT extending anteriorlyaround, and attaching to, LEI and Pbl.LE3 narrowly joining OD3?4 just anterior to tipof Eb3 uncinate process and extending to CT cov-ering (joining) Eb3 and Eb4 uncinate processes, therejoining LE4 insertion.LE4 (see also LE3) beginning on CT envelopingEb3 and Eb4 uncinate processes and continuing shortdistance posteriorly to point just medial to tip of Eb4levator process, there joining LP ventromedially.LP on Eb4 beginning at ventrolateral edge of LE4insertion and extending to bony end of Eb4, therejoining CT attaching to PP; tough fascia extends me-dially from PP, attaches to lateral edge of OP andcovers OP and Ad5 posteriorly, also infiltrates SOlaterally.LI1 on Pb2 dorsoanteriorly.LI2 on Pb3 posterolaterally adjacent to anterome-dialmost edge of Eb3, muscle fibers posterolaterallyjust failing to meet anterolateral edge of TPb3-Eb3-Eb4 muscle fibers.TD comprises TPb2, TEb2, and TPb3-Eb3-Eb4.TPb2 comprising a pair of semicircular muscle bandsoverlying the mid-section of TEb2; bands join raphesmid-anteriorly and mid-posteriorly and attach antero-laterally to Pbl, anteroventrally to dorsomedial edgeof IAC and joint with Pb2, and also medial edge ofLI1 dorsal to insertion; anterior and posterior junc-tions of muscle bands attach ventrally to CT of pha-ryngeal roof; muscle bands surround central toughCT area forming TEb2 mid-section, which is dorsalto dorsomedial surfaces of Pb3s. TEb2 consisting ofanterodorsal broader section and posteroventral nar-row section joined medially by tough CT to contra-lateral muscle sections; muscle extends laterally ontoEb2 to position anterior to LE2 insertion. TPb3-Eb3-Eb4 free from TPb2-TEb2 posteriorly; beginning an-teriorly on Pb3 near ventromedial edge of LI2 inser-tion, continuing posteriorly a short distance and at-taching to posteromedialmost edge of Eb3 and to dor-somedial surface of Eb4; muscle continuousposteriorly by slender, diagonal muscle strand withSOD.OD3-4, OD3' originate together on Pb3 dorso-
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medially and branch shortly after exiting from underTEb2 into OD3-4 dorsally and OD3' ventrally.OD3-4 inserts on Eb3 bony surface ventral to tip ofuncinate process and on posteromedial edge of bonysupport of Eb4 uncinate process, joining raphe withOP dorsally beginning just medial to tip of levatorprocess and extending medially. OD3' inserts on Eb3dorsally ventral to uncinate process.OP dorsally on Eb4 posteriorly beginning at aboutmid-length of Eb4 and extending laterally to smallbony process just medial to tip of levator process,joining raphe for most of its dorsal extent with OD3-4; laterally, OP overlaps medial half of Ad4 on Eb4;lateral edge of OP tendinous, continuous as CT sheetgiving rise to PP; ventrally OP on Cb5 dorsoposter-iorly, ventrolaterally joining raphe with Ad5.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly, beginning wellmedial to lateral end of OP and extending laterallyalmost to distal end of bone (not including cartilage),ventrally on Cb4 dorsally, extending from near innerangle of Eb4-Cb4 joint for distance equal to dorsalattachment, meeting Ad5 dorsoanteriorly on Cb4.Ad5 dorsally on AC4 medially and Cb4 beginningnear posterodistal surface and extending medially arelatively short distance, meeting Ad4; ventromedi-ally joining raphe with OP ventrolaterally.SOD present.RDs slightly separated.Additional remarks. SCL present, attached mid-dorsally to tip of elongate, ventroanteriorly curvingcartilaginous posterior end of Bb3. TV4 free fromCb5s. Pb4 and UP4 present. Medial end of Eb4smaller than that of Eb3. Pb2 toothed. PCI begins atdistal end of Cb5 and extends broadly medially, doesjoin raphe with OP, but appears to join CT coveringAd5 anteriorly. LATIDAELates niloticus (Linnaeus), USNM 332869, 61.3 mm;USNM 332870, 123 mm, USNM 166851 (2):cleared and stained.Plate 105CENTROPOMIDAEAdditional material. ? = Centropomus undecimalis(Bloch), USNM 194201, 2:71.4-114 mm.Not illustratedRemarks. Mooi and Gill (1995:129-130) arguedthat the two synapomorphies Greenwood (1976) usedto recognize a single family, Centropomidae, com-prising two subfamilies, Centropominae and Latinae,are invalid. They, therefore, recognized the two sub-families as families, with unresolved interrelation-ships. We find very little difference between the twofamilies in their dorsal gill-arch musculature.
Description.LEI short, broadly on Ebl uncinate process ante-riorly just ventral to tip.LE2 on raised posterior rim of Eb2 posterior todistal end of TEb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 dorsoposteriorly well lateral to unci-nate process, joining raphe with Ad4 dorsomediallyand joined posteroventrolaterally by LP.LP on Eb4, extending medially from dorsodistal-most bony edge to, and joining, LE4 posteroventro-laterally; CT joins ventrolateral edge of LP with PP.LI1 on dorsoanteriormost surface of Pb2 immedi-ately medial to medial end of IAC and immediatelylateral to anterolateralmost edge of Pb3.LI2 on Pb3 dorsoposterolaterally just medial tomedial end of Eb3, meeting lateral edge of TPb3.TD comprises TPb2, TEb2, TPb3-Eb3. TPb2 flat,kidney-shaped, incurved anteriorly, divided by mid-longitudinal raphe that continues posteriorly acrossTEb3; raphe attaching ventroanteriorly to pharyngealroof CT, giving rise dorsally to thin, tough CT sheets,which also attach anterolaterally to dorsoanterior sur-face on each side of TPb2; TPb2 attached anterolat-erally to Pb2 dorsoanteriormost edge and anterioredge of LI1 slightly dorsal to insertion. TEb2 almostcompletely underlying TPb2, with fibers of bothmuscles meshing on each side of mid-longitudinalraphe. TEb2 extending laterally to medial edge ofraised anterior rim of Eb2, directly anterior to LE2insertion. TPb2 and TEb2 free from TPb3-Eb3.TPb3-Eb3 attaches to Pb3 posterolaterally, meetingmedial edge of LI2 insertion and attaching tendi-nously to posteromedialmost corner of Eb3. TPb3-Eb3 continuous posteriorly by diagonal musclestrands with SOD.
? TD comprises TPb2, TEb2, and TPb3. TPb2 V-shaped, arms broad, otherwise similar to Lates.OD3-4, 3' origin on Pb3 dorsoanteromedial edgeand surface, insertion on anterior surface of Eb3 un-cinate process and medial edge of Eb4 uncinate pro-cess. OD3' present on one side in smaller specimen,but not present in larger specimen; separates ventrallyfrom OD just ventral to origin and extends onto Eb3dorsally to position ventral to insertion of OD3-4 onEb3. ? OD3' absent.OP comprises two sections, a thinner medial sec-tion and thicker lateral section; dorsally, medial sec-tion begins on Eb4 posteriorly near bony medial endand extends laterally to medial edge of lateral section,which is on Eb4 posteriorly beginning just medial tobony rise of uncinate process and extending laterallyto just below LE4 insertion medially; lateral sectionjoins raphe with OD3-4 insertion on Eb4 uncinateprocess. Ventrally, medial section is on Cb5 well me-dial to distal end and overlaps lateral section poster-omedially; lateral section extends laterally almost to
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tip of Cb5, membranously overlapping Ad5 poster-oventrally. Left side of larger specimen is anomalousin that lateral section has a separate lateral branchsplitting off and joining Ad5 dorsoposteriorly. ? OPlateral section meets OD3-4 but does not join raphewith it.Ad 1-3 absent. Ad4 relatively broad dorsally, be-ginning on Eb4 dorsoposteriorly ventral to LE4 in-sertion and ventrolateral to medial edge of OP thicksection and continuing to Eb4-Cb4 joint; muscle at-taches on Cb4 dorsally beginning at Eb4-Cb4 jointand extending medially distance about equal to extentof dorsal attachment.Ad5 moderate, attaching dorsally on Cb4 poster-odistally near AC, and ventrally on Cb5 dorsodistallyanterior to OP thick section.SOD has mid-ventral branch that separates RDsfrom each other.RDs adjacent.Additional remarks. SCL questionably free fromcartilaginous posterior end of Bb3, which is not elon-gate, nor extends ventrally. Pb4 and UP4 present.Larger specimen has tiny AC4s and, uniquely, AC5s(latter attached to distal end of Cb5s) on both sides;smaller specimen only has AC4s. ? SCL attachedmid-dorsally by short, weak ligament to ventropos-teriorly extending cartilaginous tip of Bb3. TV4 freefrom Cb5s. ? Larger specimen has tiny AC Is onboth sides, well-developed AC4s on both sides, andtiny AC3 only on left side; smaller specimen onlyhas AC4s. that of left side represented by two smallcartilages. CENTRARCHIDAEMicropterus dolomieu Lacepede, USNM 332932, 2specimens, 71.0-78.4 mm; USNM 332991, 78.9mm. Plate 106
Description.LEI on Ebl anterolateral to tip of uncinate pro-cess.LE2 on expanded dorsoposterior edge of Eb2 mid-laterally.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 just medial to minute tip of levatorprocess.LP on Eb4 at and just anteromedial to ventrolateraledge of LE4 insertion.LI1 on dorsoanteriormost Pb2 surface, joins CTbinding Pb2 and Pb3.LI2 on Pb3 dorsolaterally just anteromedial to me-dial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 roughly heart-shaped with mid-anterior notchcontinuous with raphe, which widens as CT and con-tinues across TEb2; attaching anterolaterally to dor-
solateral surface of broad cartilaginous anterior endof Pb2 and tiny AC lying dorsal to anterior end ofPb2 (see also Additional remarks); attaching mid-ventrally to CT of pharyngeal roof; meshing poste-riorly with TEb2 anteriorly. TEb2 mostly posterior toTPb2, extending laterally and attaching to Eb2 dor-sally anterior to LE2 insertion; not continuous pos-teriorly with TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 anteri-orly ventral to TEb2. Attaching to Pb3 dorsally alongmedial edge of LI2 insertion, continuing posteriorlyand attaching to posteromedialmost edge of Eb3 anddorsomedial surface of Pb4, continuous posteriorlyby fine, diagonal muscle strand with SOD.OD3?4 origin broadly on Pb3 dorsomedially, only
slightly, anteriorly ventral to TEb2; insertion mas-sively on Eb3 uncinate process anteriorly and on Eb4dorsal edge beginning at uncinate process and ex-tending medially, with ventral fibers attaching sepa-rately to ventromedial edge of Eb4.OP dorsally broadly on Eb4 posteriorly, beginningbelow medial edge of LE4 and extending medially;ventrally broadly on Cb5 dorsoposteriorly; mediallydifficult to separate from SO.Adl-3 absent.Ad4 dorsally on Eb4 posteriorly beginning belowlevator process and extending medially; ventrally onCb4 medial to Eb4-Cb4 joint.Ad5 dorsally narrowly on Cb4 posterodistally,ventrally on Cb5 dorsally posterior to Eb4 and an-terior to OP.SOD present.RDs adjacent.Additional remarks. SCL attached mid-dorsally totip of posteroventrally curving cartilaginous end ofBb3. TV4 free from Cb5s. Pb4 and UP4 present.Tiny AC on dorsoanteriormost tip of Pb2, not foundin any other actinopterygians examined. In USNM348876, two cleared and counterstained specimens25.6-29.4 mm SL showed no evidence of the AC,but a third specimen, 36.8 mm SL, in the same lot,had the AC autogenous on one side and as a bud onthe other. A fourth cleared and counterstained spec-imen, USNM 348877 67.6 mm, had the AC autog-enous on both sides.
Enneacanthus gloriosus (Holbrook), USNM 243828,72.8 mm; USNM 90449, 57.4 mm.Plate 107
Description.LEI on Ebl anterolateral to tip of uncinate pro-cess.LE2 on expanded dorsoposterior edge of Eb2 mid-laterally.LE3 on tip of Eb3 uncinate process anteriorly.
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LE4 on Eb4 levator process anteriorly (uncinateprocess absent).LP beginning on Eb4 posterior to lateral edge ofLE4 insertion and continuing laterally.LI1 on Pb2 dorsoanteriorly, joins CT binding Pb2and Pb3, which lies ventral to Pb2.LI2 on Pb3 dorsally just medial to medial end ofEb3.TD comprises TEb2 and TPb3-Eb4. TEb2 verywide, interrupted mid-longitudinally by strip of CT,which gives rise dorsally to thick CT pad; attachesanteroventrally to Pb3 just posterior to insertion ofLI1 on Pb3, mid-ventrally amidst SO fibers and CTof pharyngeal roof, and laterally on Eb2 dorsally lat-eral to LE2 insertion, meeting medial end of Ad2;discontinuous posteriorly with TPb3-Eb4. TPb3-Eb4on Pb3 dorsoposterolaterally medial to medial end ofEb3, continuing posteriorly onto dorsomedial surfaceof Eb4; continuous posteriorly by diagonal musclefibers with SOD.Remarks. Of all the centrarchid taxa examined (seealso additional acanthomorph material section), E.gloriosus is the only one lacking TPb2.OD3-4 origin broadly on Pb3 dorsomedially ven-tral to TEb2, insertion massively on Eb3 uncinateprocess anteriorly and on Eb4 dorsal edge beginning
at articulation with Eb3 uncinate process and extend-ing medially, with ventral fibers attaching separatelyto ventromedial edge of Eb4.OP dorsally on posterior surface of Eb4 medial tolevator process, ventrally on Cb5 dorsolaterally, ex-tending medially anterior to SO on Cb5. ventrolateraledge of attachment finely tendinous.Ad 1-3 moderately developed, on anterior surfaceof distal ends of respective Eb and Cb.Ad4 dorsally on Eb4 posterolaterally, beginningbelow levator process, ventrally narrowly on Cb4dorsally medial to Eb4-Cb4 joint.Ad5 dorsally narrowly on Cb4 posterolaterally,ventrally narrowly on Cb5 dorsodistally.SOD slender.RDs adjacent.Additional remarks. SCL attached mid-ventrally toventral surface of posterior tip of Bb3. TV4 com-pletely free from Cb5s or with a few dorsoanteriormuscle strands attaching to anteriormost tips of Cb5sventrally. BATHYCLUPEIDAEBathyclupea argentea Goode and Bean, USNM372712, 167 mm; USNM 305668, cleared andstained. Plate 108
Description.LEI origin tendinous; insertion on Ebl anteriorlyjust ventrolateral to tip of uncinate process. Slender
ligament originates near LEI origin, expands broadlybasally and inserts on Ebl anteriorly beginning an-terior to ventrolateral edge of LEI insertion and ex-tending laterally to end of Ebl.LE2 on raised bony posterior edge at about mid-length of Eb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 beginning on tip of Eb4 levator process andextending short distance medially, joined at ventroan-terolateralmost edge by LP.LP small, short, on Eb4 at ventroanterolateralmostedge of LE4.LI 1 almost entirely on dorsoposteromedial half ofsurface of IAC; muscle dividing longitudinally intotwo sections (but appear superficially as single mus-cle) with separate but adjacent tendinous insertionsjoining CT extending medially and joining TPb2 at-tachment to dorsalmost surface of Pb2.Remarks. The division of LI1 into two sections isapparently unusual. A non-homologous condition isduplicated in some gobioids, in which LI1 complete-ly divides, with one part inserting on Pb2 and theother on Pb3. The totality of the two LI1 parts ofBathyclupea result in a muscle that is larger than LI2.LI2 from origin ventrally to level of OD3?4, mus-cle is essentially vertical and parallels LE2 closely;on reaching level of OD3-4, muscle curves sharplyanteromedially and becomes almost horizontal as itpasses ventral to OD3-4 to its origin on Pb3 imme-diately medial to anteromedial edge of Eb3, whichlies posteroventral to Eb2 medially.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2horizontally oblong, but with deep central anteriorinvagination; muscle completely dorsal to TEb2 andposteroventrally continuous with it; muscle attachesanterolaterally to Pb2 dorsoanteriorly at joint withmedial end of IAC, there joining tendinous continu-ation of LI1 insertion; dorsoposterior surface of mus-cle with unilaterally extending slip, which continuestendinously with CT sheets covering dorsal surfaceof TD. TEb2 narrow longitudinally and relativelywide horizontally, exposed mid-dorsoanteriorly ingap exposed by TPb2 invagination; muscle with mid-longitudinal raphe attaching dorsoanteriorly to CTsheets covering TD and attaching anteriorly and mid-ventrally to CT of pharyngeal roof; muscle attacheson Eb2 beginning a little medial to LE2 insertion andcontinuing dorsolaterally to point anterior and a littlelateral to LE2 insertion; continuous posteroventrallyby fine, diagonal muscle strand with TPb3-Eb3. TPb3originates on Pb3 beginning a little anterior to jointwith medial end of Eb3 continues posteriorly alongventromedial edge of LI2 insertion and attaches toposteromedial surface of Eb3.OD3?4 origin on Pb3 dorsoanteromedially; inser-tion on medial edge and anterior bony surface justventral to tip of uncinate process, and on medial edge
138 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of Eb4 beginning just ventral to tip of uncinate pro-cess.OP dorsally on Eb4 posteriorly beginning mediallyabout halfway to medial end of bone and extendinglaterally to point between uncinate process and me-dial end of LE4 insertion, failing to meet or overlapdorsomedial edge of Ad4; ventrally on Cb5 dorsallyposterior to Ad5 ventrally, beginning a short distancemedial to distal end of bone and extending laterallyto distal end, above which OP and Ad5 join a raphe.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly, beginning me-dially ventral to levator process and extending later-ally to lateral end of bone, ventrally on Cb4 dorsallybeginning laterally near Eb4-Cb4 joint and extendingmedially about one-fourth length of bone, joining ra-phe with Ad5 on Cb4.Ad5 dorsally on Cb4 dorsally beginning laterally
at lateral end of bony portion and extending mediallyfor distance parallel to that of Ad4, with which itforms raphe; ventrally on Cb5 dorsally anterior to OP,beginning laterally at lateral end of bony portion andextending medially a short distance; joining raphewith OP (q.v.)SOD present.RDs separated by distance equal to about half di-ameter of one RD.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Medial end of Eb3 largerthan medial end of Eb4.SYMPHYSANODONTIDAESymphysanodon berryi Anderson. USNM 370558,128 mm; ? = USNM 204088, paratype, 85.7 mm;USNM 208500, paratype, ca. 120 mm, cleared andstained.
Additional material. ? S. octoactinus Anderson,USNM 204085, ca. 80 mm; ? S. species, USNM371386, 86 mm.Remarks. Only the main differences exhibited by? ? are described. Plate 109
Description.Remarks. All levators and RDs are relatively slen-der.LEI origin a fine short tendon, insertion on broadEbl uncinate process anterolaterally just ventral tocartilage cap.LE2 on dorsalmost tip of raised posterior edge ofEb2, posterior or just posterolateral to lateral end ofTEb2; lateral fibers overlap medial fibers, such thatmuscle can be separated artificially into two partswith separate but juxtaposed insertions.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on bony dorsal edge of Eb4, slightly medial
to distal end, joining LP insertion medially, levatorprocess absent. ? ? Inserts just medial to cartilagecap of levator process.LP on dorsodistalmost bony edge of Eb4, joiningLE4 insertion laterally. CT extending ventrally frominsertion attaches to distal cartilaginous edges of Eb4,AC4, and Cb5, and is continuous with PP; no sepa-rate Eb4 levator process in any of the three speci-mens. ? ? Inserts on levator process and joins LE4insertion.LI1 on Pb2 dorsolaterally just ventral to cartilagecap joining IAC medially.LI2 origin a fine tendon, insertion on Pb3 dorso-posterolaterally, just medial to medial end of Eb3;medial edge of insertion joining lateral edge of TPb3-Eb3 on Pb3.TD flat, comprises TEb2, TPb2, and TPb3-Eb3.TPb2 and TEb2 with irregular mid-longitudinal rapheattaching dorsally to CT sheets (not illustrated),which are continuous anteriorly with CT of pharyn-geal roof. TPb2 dorsal to TEb2, relatively small,broadly V-shaped, arms open anteriorly, each armarising from mid-longitudinal raphe and extendinganteriorly beyond anterior margin of TEb2 and at-taching to broad, cartilaginous cap of Pb2 processthat articulates with IAC medially (right-side arm de-formed). TEb2 extending laterally and attaching toEb2 dorsally at or a little medial to LE2 insertion;posterolaterally, fine, unilateral muscle strand extendsdiagonally posterolaterally and becomes confluentwith TPb3-Eb3 dorsally. TPb3-Eb3 on Pb3 dorsallyalong medial margin of LI2 insertion, extends nar-rowly posteriorly, passing dorsal to Pb4 and attachesto Eb3 posteromedially; posteriorly, diagonal musclestrap joins SOD anteriorly. ? Mid-longitudinal rapheis straight; no muscle strands join TEb2 with TPb3-Eb3. ? TPb2 transverse, not V-shaped; TPb3-Eb3-Eb4 present, with attachment broadly to Eb4 dorsally.? TPb2 apparently absent; TEb2 broad medially, at-tenuating anteromedially with narrow extension pass-ing between dorsally naked Pb2 surfaces and attach-ing to CT of pharyngeal roof; TPb3-Eb3 present.OD3?4 origin broadly on Pb3 dorsomedially, in-sertion on Eb3 uncinate process anteriorly ventral tocartilage tip and on medial edge of Eb4 uncinate pro-cess just ventral to cartilage tip. ? Insertion continuesonto Eb4 uncinate process ventroanteriorly. ? OD3'present.OP in two sections, dorsally on Eb4 posteriorly:medial section begins medially about mid-way be-tween medial end of Eb4 and uncinate process andextends laterally to point a little medial to uncinateprocess, meeting medial end of lateral section, whichextends laterally to point ventral to LE4 insertion;ventrally, medial section attaches to Cb5 bony sur-face posteriorly beginning a little medial to distal endand extending medially for distance about equal to
NUMBER 1 1 139
its attachment on Eb4; lateral section becomes ten-dinous ventrally, joining Ad5 posteromedially anddistal end of Cb5. ?
?
Not divided into two sections,present as one broad section equal to the two.Ad 1-3 absent; GFMs moderately developed, par-
allel anterolateral edges of Ebs and Cbs.Ad4 broadly on Eb4 dorsoposteriorly, beginningmedially ventroanterior to OP lateral section and ex-tending laterally to Eb4-Cb4 joint, ventrally on Cb4dorsoposteriorly a short distance beginning at Eb4-Cb4 joint.Ad5 relatively small; dorsoanteriorly on AC4 pos-teriorly and Cb4 posterodistally; posteriorly on distalend of Cb5 anteriorly, posteriorly joining tendinousventral end of OP lateral section. ? Does not appearto attach to AC4. ? AC4 absent, attaches mainly onunmodified distal end of Cb4, with tendinous contin-uation to contact between Cb4 and Eb4. ? AC4 ab-sent, but distal end of Cb4 with cartilaginous poste-
rior extension to which Ad5 attaches.SOD present.RDs separated by distance equal to or greater thandiameter of one RD.Additional remarks. SCL attached mid-dorsally tolong, ventroposteriorly extending cartilaginous pos-terior end of Bb3. TV4 free from Cb5s. Pb4 and UP4present. AC4 present in all three specimens, but ab-sent in
?
and
?.We examined three additional specimens: ? = S.octoactinus Anderson, USNM 204085 (2), 69.8-81.4mm SL, paratypes; ? = S. species, USNM 371386,86.0 mm SL. ? is similar to S. berryi in lackingOD3', but differs in lacking AC4 or a finger-like pro-cess extending posteriorly from the cartilaginous dis-tal end of Cb4. ? Also differs from berryi in thatEb4 has a separate levator process and the distal endof the element is a cartilaginous knob, whereas inberryi and ? there is no levator process and the car-tilaginous distal end of the element is extended dor-
sally. It seems possible that the dorsal extension maybe an early ontogenetic stage, and that with growth,the cartilage will ossify in its mid-portion, resultingin a separate levator process and a knob-like distalend. EPIGONIDAEEpigonus pandionis (Goode and Bean), USNM159341, 95.2 mm, USNM 186123, 83.4 mm.Not illustrated
Description.LEI on tip of Ebl uncinate process anterolaterally.LE2 on dorsalmost tip of raised bony posterioredge of Eb2 at about mid-length of Eb2.LE3 on tip of Eb3 uncinate process.LE4 on Eb4 dorsally just medial to tip of levatorprocess, joining LP insertion posteriorly.
LP very slender, tendinously on Eb4 medial to tipof levator process, joining LE4 insertion anteriorly.LI1 on Pb2 dorsolaterally just posterior to tip ofdorsoanterior process, with tendinous attachment ofanteroventral edge of muscle to TPb2 where TPb2attaches on IAC medially.LI2 on Pb3 posterolaterally. medial edge of inser-tion meeting lateral edge of TPb3-Eb3 on Pb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2flat, relatively narrow, beginning slightly anterior toTEb2 mid-section and extending posteriorly dorsal toanterior half of mid-section (dorsal to all of TEb2mid-section in smaller specimen); muscle with broad,shallow notch mid-anteriorly, notch leading into mid-longitudinal raphe extending through TPb2 and mostof TEb2; ventrally raphe tightly joined to CT of pha-ryngeal roof; anterolaterally TPb2 attaches to IACdorsally just lateral to articulation with Pb2 dorsoan-terior process, thus forming anterior half of notchwith laterally extending TEb2; notch embraces me-dial edge of LI1, which has fine tendinous attachmentto TPb2 near joint with IAC. TEb2 flat, relativelynarrow, extending laterally and attaching on Eb2 dor-soanteriorly anterior or well lateral to LE2 insertion;TEb2 continuous mid-posteriorly by fine diagonalmuscle strand with TPb3-Eb3. TPb3-Eb3 broadly onPb3 dorsolaterally beginning posterior to joint withEb2, and extending posteriorly along medial marginof LI2 insertion, abruptly and narrowly expandingposterior to insertion and attaching to Eb3 postero-medially; muscle continuous mid-posteriorly bybroad crossing muscle strands with SOD.OD3-4 origin on Pb3 dorsomedially, beginning alittle posterior to anterior end; insertion variable: onleft side, apparently abnormally, only on Eb3 broadlyanteriorly beginning just ventral to tip of uncinateprocess; right side attachment on Eb3 similar, but in-sertion also on Eb4 beginning on medial edge justventral to tip of uncinate process and extending ashort distance anterolaterally. In smaller specimen,insertion on both sides is similar to that of right sideof larger specimen (attachments to both Eb3 andEb4).OD3' relatively small, present only in smallerspecimen; origin on Pb3 ventral to OD3-4; insertionon Eb3 dorsally ventral to, and well separated fromOD3-4.OP dorsally on Eb4 posteriorly beginning aboutmid-way between medial end and extending laterallya little lateral to uncinate process, ventrally on Cb5beginning moderately medial to distal end and ex-tending laterally beyond distal end onto Ad5 dorso-medially, there joining raphe with Ad5; muscle me-dially unclearly differentiated from SO laterally.Adl-3 absent. GFM1-3 weakly developed.Ad4 dorsally on Eb4 posteriorly beginning medi-ally below tip of levator process and extending lat-
140 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
erally to medial edge of cartilaginous distal end, therealso attaching minimally to AC4; ventrally on Cb4beginning medially at point ventrally opposite medialend of attachment on Eb4 (muscle ventromediallyoverlapped by OP) and extending laterally to medialangle of Eb4-Cb4 joint, joining raphe (not visible ex-ternally) with Ad5 anterolaterally.Ad5 very small, dorsoanteriorly on Cb4 posteri-orly a little medial to distal end, ventrally on Cb5beginning a short distance medial to distal end andextending to distal end, dorsally joining raphe withOP ventrolaterally, and another with Ad4 ventropos-teriorly.SOD present.RDs moderate, separated by space about half di-ameter of one RD (by space almost equal to one RDin smaller specimen).Additional remarks. SCL present, but only RV4attaches to it; appears to be fused with ventral aorta;possibly attached to elongate, ventrally curved car-tilaginous posterior end of Bb3 (removing aortawould have destroyed attachment, if present). TV4free from Cb5s. Pb4 present. UP4 present. Pbl most-ly bony. MORONIDAEMorone americana (Storer), USNM 61612, 99.9 mm.Plate 110
Additional material. ? = Morone mississippiensisJordan and Eigenmann, USNM 231447, >100mm.
Description.LEI tendinously on anterior bony surface of Ebluncinate process.LE2 on dorsoposterior surface of Eb2.LE3 on cartilaginous tip of Eb3 uncinate processdorsoanteriorly.LE4 on Eb4 levator process dorsodistally on oneside, just proximal to dorsal tip on the other. ? Justproximal to dorsal tip on both sides.LP origin tendinous, insertion at and ventrolateralto LE4 insertion. ? Ventroanterior to LE4 insertion.LI1 on dorsoanteriormost bony surface of Pb2.LI2 on Pb3 dorsoposterolaterally, at and lateral toTPb3-Pb4-Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 with mid-longitudinal raphe continuing poste-riorly across TEb2, raphe attaching ventrally to CTof pharyngeal roof and dorsally giving rise to CTsheet covering TD; anteriorly, muscle attaches onmost of dorsoanterior surface of IAC, posteriorlymuscle is dorsal to anterior end of TEb2, to which itis fused mid-posteroventrally (no attachment to Pb2).TEb2 with dorsolateral^ and ventrolaterally orientedmuscle strands fusing laterally but with slightly di-
vided attachments on dorsal surface of Eb2, anteriorattachment extending a little more laterally than pos-terior attachment, ending posteromedial to M. Pb2-Eb2 and anterior to LE2 insertion; muscle lying dor-
sal to, but posteriorly connected by CT and fine di-agonal muscle strands with, anterior end of TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 on Pb3 dorsoposterior bonysurface at and medial to LI2 insertion, on Pb4 dor-soanteriorly, and narrowly, tendinously on Eb3 pos-teromedialmost end, continuous posteriorly by diag-onal strands of muscle with SOD. ? TPb2 on onlymedial third of dorsoanterior surface of IAC.M. Pb2-Eb2 on anterolateralmost bony surface ofPb2 and dorsoanterior surface of Eb2 anterior toTEb2.Remarks. This muscle is uncommon in acantho-morphs, but occurs at least in the ophidiids Brotulaand Dicrolene, and all percopsiforms.OD3-4 anteriorly on dorsoposterior surface ofPb3, posteriorly on anterior surface of Eb3 uncinateprocess and anterior surface and medial edge of Eb4uncinate process.OP dorsally on most of Eb4 posterior surface me-dial to Ad4, ventrally, narrowly on Cb5 dorsal sur-face, joining small, strong raphe there with ventro-medial edge of Ad5; unusually well separated later-
ally from Ad4; a strap of SO muscle joins OP onCb5 but does not attach to Eb4 as it continues ante-
riorly and joins SO longitudinal muscle fibers ex-tending between Pb3s.Ad 1-3 absent.Ad4 dorsally on ventral surface of Eb4 lateral toOP ventrally on dorsoposterior surface of Cb4 medialto Eb4-Cb4 joint.Ad5 dorsally, tendinously and musculously onposterodistal surface of Cb4 and tendinously on mi-nute AC; ventrally on dorsodistal surface of Cb5.SOD present.RDs slightly separated.Additional remarks. SCL attached mid-dorsally tocartilaginous ventroposteriorly extending tip of Bb3.TV4 free from Cb5s. Tiny AC4 and AC2 present. ?AC4 relatively well developed; AC2 absent.SERRANIDAERemarks. Imamura and Yabe (2002) hypothesizeda reorganization and recomposition of the Scorpaen-iformes of previous authors. They recognized a scor-paenoid-serranoid sister group relationship within thePerciformes, but we have retained both groups sep-arately within our Perciformes pending further cor-roboration of their findings. See also remarks follow-ing Scorpaenoidei.Epinephelus merra Bloch, USNM 318074, 3 speci-mens, 68.6-92.5 mm.Plate 111
NUMBER li 141
Additional material.
?
= Anthias nicholsi, USNM151904, 110 mm (very similar to E. merra).
Description.LEI with tendinous origin, insertion broadly onbony anterior surface of Ebl uncinate process.LE2 on bony dorsoposterior edge of Eb2.LE3 on dorsoanterior edge of tip of Eb3 uncinateprocess.LE4 on dorsoposterior edge of Eb4 medial to tipof levator process.LP slender, on Eb4 at and lateral to lateral edge ofLE4 insertion. ? At and anterior to LE4 insertion.LI1 on Pb2 dorsomedially ventral to TPb2 attach-ment; slightly larger than LI2.
? On dorsal surfaceof Pb2 dorsoanteriorly just ventral to joint with IAC.LI2 on Pb3 dorsolaterally at anteriormedialmostend of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2heart-shaped, deep notch anteriorly leading to mid-longitudinal raphe, which gives rise to dorsally to CTsheets attaching to skull; on broad Pb2 dorsoanteriorprocess and ventroposterior end of Pbl (Pb2 and Pbltightly joined by CT at this point), continuous pos-teroventrolaterally with TEb2. TEb2 on Eb2 anteriorto LE2 insertion, continuous posteriorly by diagonalstrand of muscle with TPb3-Eb3. TPb3-Eb3 on Pb3dorsal surface medial to LI2 insertion and anterior toEb3 cartilaginous medial end, and on Eb3 bony sur-face dorsoposteromedially, continuous by diagonalstrands of muscle with SOD. ? TPb2 does not attachto Pbl.OD3?4 originates on Pb3 dorsomedially ventral toTEb2 and inserts on anterior surface of Eb3 uncinateprocess and anterior surface and medial edge of Eb4uncinate process.OP dorsally, broadly on Eb4 posteromedially be-ginning near levator process and extending medially;ventrally on Cb5 dorsolaterally posterior and coin-cident with Ad5, medially weakly separable fromSO.Ad 1-3 absent.Ad4 dorsally, broadly on Eb4 posterior surface lat-eral to OP; ventrally broadly on Cb4 dorsally medialto Eb4-Cb4 joint.Ad5 on posterodistal surface of Cb4 and dorsolat-eral surface of Cb5 anterior and coincident with OP.SOD with mid-longitudinal raphe, muscle fibersextend ventrally from raphe forming a short com-partment on each side that isolates each RD.RDs separate.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present.LUTJANIDAEPristipomoides aquilonaris (Goode and Bean),USNM 158472, 101 mm.
Plate 112
Additional material.
?
= Hoplopagrus guentherii,USNM 65544, 94.6 mm.
Description.Remarks. Hoplopagrus gill-arch muscles are verysimilar to those of Pristipomoides; however, they aremuch more massive relative to the size of the spec-imen.LEI finely, tendinously on dorsal edge of Ebl un-cinate process just lateral to process tip. ? Tendi-nously on and lateral to tip of uncinate process.LE2 on dorsal edge of raised posterior edge ofEb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 levator process dorsoanteriorly.LP at and ventroanterior to LE4 insertion.LI1 on Pb2 dorsal surface ventral to articulationwith IAC.LI2 on Pb3 dorsolaterally medial to anteromedialend of Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 is dorsal to TEb2, with median longitudinalraphe giving rise to CT sheets attaching to skull;muscle attaches anterolaterally to CT binding dor-soanteriormost cartilaginous ends of Pb2 and Pb3 anddorsomedial surface of IAC, attaches mid-ventrallyto CT of pharyngeal roof, fuses mid- and ventrolat-erally with TEb2, and is posteriorly free from TPb3-Pb4-Eb3. TEb2 twisting dorsoposteriorly after pass-ing laterally from under TEb2, attaching on Eb2 dor-soanteriorly anterior to LI2 insertion. TPb3-TPb4-Eb3 on Pb3 dorsolaterally medial to medial end ofEb3, continuing posterolaterally and passing dorsalto medial end of Eb4 and attaching to posteromedialedge of Eb3; muscle fibers delaminate from ventralsurface of Eb3 portion and attach to dorsal surfaceof Pb4; posteriorly continuous by fine muscle strandswith SOD.
?
Comprises TPb2, TEb2, and TEb3.TEb3 on medial end of Eb3 dorsoposteriorly. TEb2does not twist laterally.OD3-4 origin broadly on Pb3 dorsomedially ven-tral to TEb2, insertion mostly on anterior surface ofEb3 uncinate process, weakly on anterior surface ofEb4 uncinate process.OP dorsally on Eb4 posteriorly beginning near me-dial end and extending laterally almost to levator pro-cess; ventrally broadly on Cb5 dorsoposteriorly, pos-terior to Ad5.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning at le-vator process and extending laterally to end of bonysurface; ventrally broadly on Cb4 extending laterallyto Eb4-Cb4 joint.Ad5 ventrally broadly on Cb5 dorsally beginning
at distal end of Cb5, joining small raphe with OP
142 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ventroanterolaterally; dorsally on posterodistal end ofCb4.SOD present.RDs separated by space about equal to diameter ofone RD.Additional remarks. SCL attached mid-posteriorlyto very elongate cartilaginous ventroposterior tip ofBb3. TV4 free from Cb5s. Pb4 and UP4 present.HAEMULIDAEPomadasys crocro (Cuvier). USNM 338643, 2 spec-imens, ca. 68-78.1 mm.
Additional material. Haemulon scudderi Gill, USNM181299, 70.9 mm; Plectorhinchus pictus (Thun-berg), USNM 192546, 83.0 mm.Plate 113INERMIIDAE
*? = Inermia vittata Poey, USNM 318643, ca. 80mm SL. Not illustratedRemarks. We found no notable differences in themuscles among the three haemulid taxa. Inermiidmuscles are very similar to those of haemulids, towhich family they are closely related (Johnson,1980). Additional evidence for this relationship isthat the cartilage tip of the Ebl uncinate process isattached to the medial edge of a small, raised bonyflange, rather than to a distinct, rod-like arm. A some-what similar arrangement is present in centracanthidsand some sparids. e.g., Acanthopagrus. Aside fromthe fact that inermiid levators are less robust thenthose of haemulids, only the main differences are not-ed below. The inermiids and haemulids share in hav-ing the uncommon (for percomorphs) combination ofabsence of both SCL and SOD.
Description (based on Pomadasys).LEI origin slender, tendinous; insertion broad, onEbl beginning at joint of uncinate process with IACand extending laterally to point almost half distanceto distal end.LE2 on medial edge of raised posterior margin ofEb2.LE3 on Eb3 just anteroventral or just anterolateralto tip of uncinate process, meeting OD3?4 on Eb3.
? Inserts by long, fine tendon on tip of Eb3 uncinateprocess on one side, and by low, broad tendon onother.LE4 on Eb4 levator process dorsally, just posteriorto uncinate process, meeting but not joining LP me-dially. ? Inserts by long, fine tendon on tip of Eb4levator process on one side, and by low, broad tendonon other.LP broadly on Eb4 dorsally, extending mediallyfrom distal end to position just lateral to LE4 inser-
tion immediately anterior to tip of levator process,joining raphe with Ad4 dorsally.LI1 tendinously on Pb2 dorsoanteriorrmost tipposteriorly, extending laterally onto adjacent IACposteromedially (not visible in illustration) up toabout one-third length of IAC; tendinous attachmentto Pb2 continuous with CT joining Pb2 with anter-iormost tip of Pb3 dorsally (insertion here not con-sidered to include Pb3). ? Does not insert on IAC.LI2 tendinously on Pb3 dorsolaterally immediatelyanterior to anteromedialmost edge of Eb3, well sep-arated from TPb3-Eb3.TD comprises TEb2 and TPb3-Pb4-Eb3. TEb2 ro-bust, with broad, irregular central CT portion at-tached mid-ventroanteriorly to CT of pharyngeal roofand giving rise mid-dorsally to CT sheets attachingto skull; muscle extends laterally and attaches on Eb2dorsally anterior to LE2 insertion, also meeting me-dial end of GFM2 (not illustrated) as latter extendsdorsally on Eb2; muscle completely disjunct fromTPb3-Pb4-Eb3. TPb3-Pb4-Eb3 beginning on Pb3dorsolaterally medial to mid-medial end of Eb3 andcontinuing onto posteromedialmost corner of Eb3and dorsomedial edge of Pb4.OD3-4, OD3' with joint origin on Pb3 dorsopos-teromedially; relatively large OD3' splits off ven-troanteriorly shortly lateral to origin and inserts onEb3 dorsally ventrolateral to OD3-4 portion on Eb3;OD3-4 inserts massively on Eb3 uncinate processanteriorly and on Eb4 anteriorly just ventral to tip ofuncinate process.OP dorsally on Eb4 posteriorly beginning near me-dialmost bony end and extending laterally to medialedge of levator process, overlapping medial portionof Ad4 posteriorly; ventrally on Cb5 dorsoposteriorlybeginning medially near TV5 and extending laterallyalmost to distal end, but becoming membranous ven-trolaterally and overlapping Ad5 posteriorly; ventro-medially fusing with SO.Adl-3 absent. GFM1-3 (not illustrated) moderate-ly developed; GFM2 extending anteromedially, thendorsally on Eb2 to position anterior to distal end ofTEb2.Ad4 dorsally on Eb4 posteroventrally beginninganterior to OP near bony medial end and extendinglaterally to distalmost bony end; ventrally, moderate-ly broadly on Cb4 laterally, becoming fused poste-riorly with Ad5 anteromedially.Ad5 dorsally on AC4 posteriorly and Cb4 poster-odistally for short distance, there anteriorly joiningAd4 posteriorly; ventrally on Cb5 dorsoposterolater-
ally, mostly anterior to OP.SOD absent.RDs adjacent. ? Separated by distance equal todiameter of one RD.Additional remarks. SCL absent; cartilaginous pos-terior end of Bb3 not elongate. TV4 free from Cb5s.
NUMBER 11 143
Pb4 and UP4 present. Pbl with cartilaginous ends.Pb2 with teeth. AC4 present. Dorsoanterodistalmostbony margin of Eb4 and, variously, Eb2 and Eb3,with small flange in Haemulon and Plectorhinchus;flanges very reduced in Pomadasys, very well de-veloped in a cleared and stained specimen (USNM214488, 95 mm) of Parakuhlia macropthalmus (Oso-
rio). ? Flange clearly present, but so fine as to beeasily overlooked.APOGONIDAEGlossamia wichmanni (Weber), USNM 344886, 2specimens, 67.3?97.7 mm.Plate 114 (based on smaller specimen)
Additional material.
?
= Cheilodipterus macrodonLacepede, USNM 276623, 82.6 mm SL.Remarks. Differs from G. wichmanni mainly inhaving musculature much less robust.
Description.LEI on expanded dorsoposterior edge of Ebl lat-eral to tip of uncinate process.LE2 on expanded dorsoposterior edge of Eb2about mid-laterally.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on expanded dorsoposterior surface of Eb4lateral to tip of uncinate process.LP on Eb4 at and lateral to lateral edge of LE4.Remarks. Narrow, lateralmost cartilaginous edgeof expanded dorsoposterior surface of left-side Eb4narrowly continuous ventrally with knob-like carti-laginous distal end of Eb4. Cartilaginous lateral endcontinuous ventrally with knob-like distal end of Eb4on both sides in larger specimen and ?.LI1 on dorsoanterolateralmost surface of Pb2 justmedial to articulation with IAC.LI2 on Pb3 dorsoposterolaterally just anteromedialto medial end of Eb3.TD comprising TPb2, TEb2. and TPb3-Eb3, withmid-longitudinal raphe extending through all com-ponents and SOD, giving rise dorsally to CT sheetcovering surface of muscles, and attaching ventrallyto CT of pharyngeal roof between Pb3s. TPb2 verythin, flat, bilobed, arising dorsally from mid-longi-tudinal raphe in common with TEb2 posteriorly; an-teriorly joining raphe with TEb2 anterolaterally and
at that point weakly attaching to cartilaginous ante-
rior end of Pb2 just medial to LI1. TEb2 very broad,attaching on Eb2 dorsally anterior to LE2 insertion,continuous posteroventrally only by CT with TPb3-Eb3. TPb3-Eb3 on Pb3 dorsoposterolaterally begin-ning along medial edge of LI2 insertion and continu-ing posteriorly, crossing over medial end of Eb4 andattaching along posteromedial edge of Eb3; narrowlycontinuous mid-posteriorly with SOD.OD3-4 origin broadly on Pb3 dorsally ventral to
TEb2; insertion broadly on anterior surface of Eb3uncinate process and medial edge of Eb4 uncinateprocess.OP dorsally broadly on posterior surface of Eb4beginning ventral to LE4 and extending medially;ventrally on Cb5 dorsoposteriorly, joining small ra-phe ventrolaterally with Ad5.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 posteroventrolater-
ally, extending medially just under (anterior to) lat-eral edge of OP; ventrally broadly on Cb4 dorsallymedial to Eb4-Cb4 joint, meeting Ad5 attachment onCb4.Ad5 dorsally on Cb4 posterolaterally; ventrally onCb5 dorsally mostly anterior to OP, joining rapheposteroventrally with ventromedial edge of OP.SOD very fine.RDs separated by space less than half diameter ofone RD.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 toothed. Eb4 levatorprocess present on only one side of smaller specimen,absent on both sides of larger specimen. ? Eb4 le-vator process present on one side, absent on other.PRIACANTHIDAEHeteropriacanthus cruentatus (Lacepede), USNM319919, 63.3 mm; ? = USNM 141752, 88.0 mm.Plate 115
Description.LEI on dorsoposterior edge of Ebl well lateral touncinate process, but not extending onto distal quar-ter of surface.LE2 on mid-dorsoposterior edge of Eb2, insertioncontinuous posteriorly with ligament attaching Eb2to mid-anterior edge of Eb3.LE3 relatively small, on joined tips of Eb3 andEb4 uncinate processes.LE4 on dorsal edge of Eb4 levator process, inser-tion continuous ventrolaterally with ventromedialedge of CT sheet joining PP ventrally.LP finely tendinously on Eb4 at lateralmost edgeof LE4 insertion.LI1 on dorsomedialmost end of Pb2.LI2 on Pb3 dorsolaterally at articulation with me-dial end of Eb3.TD comprises TEb2 and TPb3-Eb4. TEb2 broad,wing-like, depressed centrally, with lateral margin ofdepression on each side represented by a surface ra-phe that gives rise to thick CT sheet attaching toskull; muscle attached antero- and ventromedianlybetween Pb3s to CT of pharyngeal roof; fibers ofanterior two-thirds of muscle curve posterolaterally,those of posterior third more-or-less transverse, fibersfuse laterally as muscle narrows and extends ontoEb2, attaching to Eb2 dorsally anterolateral to LE2
144 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
insertion; muscle is not connected posteriorly withTPb3-Eb4. TPb3-Eb4 on Pb3 dorsoposterolaterally,extending broadly posteriorly onto medial arm ofEb4 anteriorly, attached mid-anteroventrally to CT ofpharyngeal roof.OD3-4, OD4v origin broadly on Pb3 dorsomedi-ally ventral to TD, insertion on anterior surface ofEb3 uncinate process and medial edge of Eb4 unci-nate process, with short, separate branch (OD4v, notillustrated) attaching broadly dorsally on Eb4 ventro-medial to uncinate process.Remarks. OD4v probably autapomorphic.OP dorsally on Eb4 posteriorly, beginning on pos-terior surface of uncinate process and extending me-dially and becoming undifferentiated from SO, ven-trally on Cb5 dorsoposterolaterally, ventrolaterallyjoining raphe with Ad5 posteroventrally.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 beginning on ventralsurface of levator process and extending laterally onEb4 posteriorly, ventrally on Cb4 dorsolateral^ me-dial to Eb4-Cb4 joint.Ad5 dorsally on Eb4 posterodistally, including tinyAC4 attached to posterodistal end of Cb4 (present onright and left arches), ventrally on Cb5 dorsally an-terior to OP, joining raphe with OP ventrolaterally. ?AC absent.Remarks. Presence of AC4 is variable, and possi-bly ontogenetically correlated. In a cleared andstained specimen 58.9 mm (USNM 337685) it ispresent on one side but not the other.SOD absent.RDs separated by space less than one RD diameter.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. IAC reduced to tiny car-tilage adjacent to tip of Ebl uncinate process, at-tached by long, slender ligament to tip of Pb2 unci-nate process. OSTRACOBERYCIDAE
Ostracoberyx dorygenys Fowler, USNM 307282,80.0 mm. Plate 116
Description.LEI broadly on dorsoposterior edge of Ebl begin-ning on ventrolateral edge of long uncinate processand extending laterally; origin by long tendon.LE2 very broadly on expanded posterior surfaceof Eb2, ventromedial edge of insertion meeting TEb2posteriorly.LE3 on joined tips of Eb3 and Eb4 uncinate pro-cesses anteriorly, musculously on Eb3, tendinouslyon Eb4.LE4 broadly on Eb4 dorsolaterally, ventroposter-omedially joining raphe with Ad4 dorsomedially
(present on both sides and can be characterized as anLE4-Ad4 muscle sling).LP very slender, tendinously on dorsodistalmostedge of Eb4, joining ventrolateral edge of LE4 in-sertion.LI 1 on dorsalmost edge of Pb2, ventromedial edgeof insertion joining raphe with TPb2 anterolaterally;muscle about same size as LI2.LI2 on Pb3 dorsoposteriorly at medial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2a thick, roundish pad broadly continuous ventrallywith TEb2; attaching anteriorly to CT of pharyngealroof; attaching tendinously anterolaterally to dorsaledge of Pb2 and adjacent anterior tip of Pb3 (notillustrated); with broad mid-dorsal raphe-like areagiving rise to tough, filmy CT sheets covering TD.TEb2 extending laterally onto Eb2 anterior to LE2insertion, continuous mid-ventroposteriorly by diag-onal muscle strands with TPb3-Eb3. TPb3-Eb3 onPb3 dorsoposterolaterally near medial edge of LI2insertion, rising dorsally onto Eb3 beginning on pos-teromedialmost edge and extending a short distancealong posteriorly, continuous posteriorly with slenderSOD.OD3-4 origin on Pb3 dorsoanterolaterally ventralto TEb2, insertion broadly on Eb3 uncinate processanteriorly and medial edge of Eb4 uncinate process,joining narrow raphe posteroventrally with OP a littlemedial to Eb4 uncinate process.OP dorsally broadly on Eb4 posteriorly beginning
at uncinate process, extending medially, and becom-ing inseparable from SO, ventrally on Cb5 postero-laterally, joining short raphe with Ad5 ventrally.Ad 1-3 absent.Ad4 on Eb4 dorsoposterolaterally and Cb4 dor-solaterally medial to Eb4-Cb4 joint, joining raphewith Ad5 dorsoanteriorly on Cb4.Ad5 dorsally beginning tendinously on Cb4 pos-terodistally and extending medially on Cb4 dorsally,joining raphe with Ad4 ventrally; ventrally, narrowlyon Cb5 dorsodistally, joining raphe with OP ventral-ly-SOD moderately slender.RDs separated by space equal to about one RDdiameter.Additional remarks. SCL attached mid-posteriorlyto anterior surface of posteroventrally extending car-tilage tip of Bb3. TV4 free from Cb5s. Pb4 and UP4present. IAC present. Pb2 toothed. Eb4 levator pro-cess absent.
CIRRHITIDAE
Paracirrhites forsteri (Schneider), USNM 278070,91.3 mm; USNM 339098, 67.2 mm.Plate 117
NUMBER 11 145
Additional material.
?
= Cirrhitus pinnulatus (For-
ster), USNM 296746. 65.0 mm.
Description.LEI broadly on Ebl uncinate process anteriorlyjust lateral to cartilaginous tip; origin narrowly, ten-dinously united with posterior surface of epithelialsheet fronting gill arches dorsoanteriorly, separatingthem from pseudobranchs, also incorporates Pbls.Strong ligament inserts on Ebl anterior to LEI andextends to cranium.
? Ligament and epithelial sheetpossibly removed without notation (early dissection).LE2 on Eb2 at about mid-length dorsoposteriorly.LE3 finely on tip of Eb3 uncinate process anteri-orly, meeting insertion of OD3-4 on Eb3.LE4 on Eb4 dorsally beginning on tip of levatorprocess and extending medially a short distance,joined laterally or anterolaterally by LP insertion.LP relatively small, on Eb4 joining LE4 insertionlaterally or anterolaterally, insertion continuous ven-trolaterally with CT sheet (not illustrated) attachingto fourth and fifth arches and incorporating PP ven-trally.LI1 broadly on Pb2 dorsally beginning just pos-teroventral to dorsal tip; medial edge of muscle, dor-
sal to insertion, attaches to CT joining TPb2 anteri-orly to Pb2 and posteromedial edge of IAC.LI2 on Pb3 broadly posterolaterally opposite me-dial edge of Eb3, medial edge of insertion meetingTPb3 laterally.TD comprising TPb2, TEb2, and TPb3. TPb2,large, flat, circular, almost completely covering mid-section of TEb2, with shallow notch mid-anteriorlyand mid-posteriorly connected by mid-longitudinalraphe, which gives rise dorsally to thin sheets of CTand is joined mid-ventrally by TEb2; muscle attachedanterolaterally by CT to closely approximated Pbl-Ebl joint, IAC-Pb2 joint, and medial edge of LI1;attached mid-anteriorly and mid-ventroposteriorly toCT of pharyngeal roof. TEb2 medially ventral toTPb2, attaching along mid-longitudinal raphe ven-trally, extending laterally and attaching on Eb2 dor-sally a little anterolateral to LE2 insertion; posteriorlydiscontinuous from TPb3. TPb3 on Pb3 mostly ven-tral to OD3-4, attaching on Pb3 posterolaterallymeeting medial edge of LI2 insertion, continuous bydiagonal muscle strand with SOD. ? TPb2 on TEb2dorsally; a flat, semicircular ribbon, open anteriorly,originating anteriorly on each side at CT attachmentto closely approximated Pbl-Ebl joint, IAC-Pb2joint, and medial edge of LI1.OD3-4 origin broadly on Pb3 medially ventral toTEb2, insertion on anterior surfaces of Eb3 and Eb4uncinate processes.OP dorsally, broadly on Eb4 beginning at medialend of bony portion and extending laterally almostto levator process, there meeting dorsomedial end of
Ad4; ventrally on Cb5 posterolaterally, variably join-ing or not short raphe with ventroposterior end ofAd5 and more extensive one with PCI.M. SO-Pb3 originates from SO longitudinal mus-cle layer as two broad, anteriorly extending musclestraps, lateral strap inserts on Pb3 posteriorly, medialstrap (not visible in illustration) originates ventral orventromedial to RD and inserts on Pb3 posterome-dially; RD inserts on Pb3 between M. SO-Pb3 pair.Ad 1-3 absent, but very fine muscle strand ob-scured by bases of gill-filaments on each arch an-terolaterally.Ad4 dorsally on Eb4 ventrally, beginning belowlevator process and extending to Eb4-Cb4 joint, ven-trally, broadly on Cb4 dorsally medial to Eb4-Cb4joint.Ad5 dorsally, moderately broadly on posterolateralsurface of Cb4, ventrally on Cb5 posterolaterally,variably joining or not OP posterolaterally in a shortraphe.SOD present.RDs moderately separated or adjacent, insert bylong tendon on Pb3 posteriorly between insertions oflateral and medial M. SO-Pb3 pair, except on oneside of one specimen, in which RD tendon insertsinto CT of pharyngeal roof posterior to Pb3.Additional remarks. SCL attached mid-dorsally toposteroventrally elongated cartilaginous end of Bb3.TV4 free from Cb5s. Pb4, UP4, and IAC present.Pbls oriented perpendicular to distal ends of Ebls;
? Pbls oriented parallel to Ebls.PEMPHERIDAEPempheris schomburgkii Miiller and Troschel,USNM 318588, 95.5 mm; Pempheris schwenkiiBleeker, USNM 324224, 105 mm.Plate 118Additional material. ? = Parapriacanthus ranson-neti Steindachner, USNM 344281, 79.2 mm.GLAUCOSOMATIDAE
?
= Glaucosoma magnificum, WAM P. 14208-011,93.2 mm. Not illustrated
Remarks. We noted no substantive differencesamong the three genera.Description (based on Pempheris schomburgkii).LEI relatively slender, on dorsal edge of Ebl un-cinate process just lateral to cartilaginous tip; strong,fine ligament (not illustrated) extends dorsally to ven-tral surface of the skull from margin of Eb 1 anteriorto uncinate process; ligament is appressed to poste-rior surface of vertical CT pane lining pharyngealarea anterior to gill arches.
146 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LE2 on rounded high point of dorsoposterior edgeof Eb2 mid-laterally.LE3 on dorsal tip of Eb3 uncinate process.LE4 on dorsodistal edge of Eb4, joining LP ante-roventrally; levator process absent. ? LE4 and LP onlevator process. ? Levator process present.LP on Eb4 at and anterior to LE4 insertion. ? ?See LE4 above.LI1 has long, slender tendinous insertion on Pb2ventral to joint with IAC.LI2 short tendinous insertion on Pb3 dorsoposter-olateral surface at and just touching medial end ofEb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2,thick, pad-like, oblong, with irregular mid-longitu-dinal raphe giving rise to CT sheet covering muscleon anterior half; muscle attaches beginning anteriorlyat CT of pharyngeal roof, continuing laterally overand attaching to anterior end of Pb3, dorsomedialsurface of broad Pb2 uncinate process, and dorso-medial surface of IAC, remainder of muscle marginfree, but muscle fusing ventromedially with TEb2;amorphous CT connects mid-posteroventral surfaceto mid-anterior edge of TPb3-Eb3. TEb2 extendingout on dorsal surface of Eb2 to point anterolateral toLE2 insertion. TPb3-Eb3 on Pb3 posterolaterally.continuing on Eb3 posteromedial edge, continuousventrally by crossing strands of muscle with SOD,which is ventral to TPb3-Eb3. ? TPb2 flat, attach-ment on IAC extends to about mid-length of element.
? Attachment extends laterally only to dorsolateral-most edge of Pb2 (i.e., not on IAC).OD3-4 origin on Pb3 ventral to TPb2 and TEb2,inserting on anterior surface of Eb3 uncinate processand medial edge of Eb4 uncinate process.OP dorsally on Eb4 mid-dorsoposteriorly begin-ning at uncinate process and continuing mediallyabout half distance to medial end of Eb4, extendingventrally posterior to dorsomedial edge of Ad4 andposteroventral surface of Ad5 and inserting on pos-terodistal surface of Cb5.Ad 1-3 absent.Ad4 dorsally on ventral surface of Eb4 mostly lat-eral to OP, and ventrally, broadly on Cb4 dorsal sur-face medial to Eb4-Cb4 joint.Ad5 ventrally on dorsal surface of distal end ofCb5 anterior to OP insertion and broadly on Cb4 ven-troposterior surface and small AC4 posteriorly.SOD slender, ventral to TPb3-Eb3: moderatelywell developed, just ventroposterior to TPb3-Eb3 inP. schwenkii. ? Well-developed, well posterior toTPb3-Eb3, but continuous with it by slender, diago-nal muscle strap. ? Very fine, ventral to TPb3-Eb3.RDs adjacent.Additional remarks. SCL present, attached mid-dorsally by long, loose CT to tip of elongate, ventro-posteriorly extending cartilaginous posterior end of
Bb3. TV4 free from Cb5s. ? SCL apparently freefrom short, cartilaginous posterior end of Bb3(should be verified in additional specimens). Pb4 andUP4 present. Pb2 toothed.Johnson (1993:19), following Tominaga (1986),recommended recognizing the Glaucosomatidae as asubfamily of the Pempheridae and described addi-tional specializations shared by the two groups. Nel-son (1994:365-366), McKay (1997:5). and Mooi andGill (2002:23), without explanation, all chose not tofollow Johnson's recommendation. McKay (1997:5)proposed that the Pempheridae and Glaucosomatidaeare sister groups.To the specializations shared by pempherids andglaucosomatids previously reported in the literature,we add the ligament positioned anterior to LEI, ageneral similarity of the dorsal gill-arch musculature,and the shape of Eb4, which is posteriorly concave.
LACTARIIDAE
Lactarius lactarius (Bloch and Schneider), USNM343873, 119 mm. Plate 119
Description.LEI on Ebl uncinate process ventrolateral to tip;origin tendinous; long, broad-based ligament on Eblanterior to uncinate process (also in Glaucosoma?see additional remarks under Pempheridae).LE2 on dorsally expanded bony posterior marginof Eb2.LE3 on tip of Eb3 uncinate process medially.LE4 on Eb4 levator process beginning at ventrallypositioned cartilage tip and extending medially.LP on Eb4 at and anterior to anterior margin ofLE4 insertion, ventrolaterally joining CT sheet.LI1 very broad dorsally. on Pb2 dorsally, mediallyventral to TPb2.LI2 on Pb3 dorsoposterolaterally just anterior tomedial end of Eb3.TD comprises TPb2. TEb2, and TPb3-Eb3. TPb2heart-shaped, attaching to Pb2 dorsoanteriorly, withmedian longitudinal raphe posteriorly; raphe joinedventrally by TEb2 medially. TEb2 broad, attachingalong mid-longitudinal raphe ventrally between Pb3swith CT of pharyngeal roof, narrowing laterally andattaching along Eb2 dorsally anterior to LE2 inser-tion, continuous mid-posteriorly by CT (not illustrat-ed) with TPb3-Eb3 mid-anteriorly. TPb3-Eb3 on Pb3dorsoposterolaterally medial to medial end of Eb3and on posteromedial edge of Eb3, continuous pos-teriorly by diagonal muscle strands with SOD.OD3-4 origin on Pb3 dorsoanteromedially ventralto TEb2; insertion on anterior surface of Eb3 unci-nate process and medial edge of Eb4.OP dorsally on Eb4 posteriorly at and medial to
NUMBER 11 147
uncinate process; ventrally on Cb5 posterodistally,joining raphe ventrolaterally with Ad5.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 posteriorly beginningventrally at levator process and extending laterally toend of bone, ventrally broadly on Cb4 dorsally me-dial to Eb4-Cb4 joint, meeting Ad5 anterolaterally onCb4, but diverging from Ad5 medially.Ad5 dorsally on Cb4 bony surface entirely medialto cartilaginous end; ventrally on Cb5 dorsolaterallyjoining raphe posterolaterally with OP.SOD slender.RDs separated by space greater than one RD di-ameter.Additional remarks. SCL attached mid-dorsally totip of posteroventrally extending cartilaginous end ofBb3. TV4 free from Cb5s. Pb4 and UP4 present. IACpresent. LATEOLABRACIDAE2
Lateolabrax japonicus (Cuvier). USNM 344295, ?
= 102 mm; USNM 64631, 109 mm.Plate 120
Description.LEI on bony anterior surface of Ebl uncinate pro-cess.LE2 on dorsomedial edge of expanded bony edgeof Eb2.LE3 on joined tips of Eb3 and Eb4 uncinate pro-cesses, there joining raphe with OD3-4 insertion.LE4 on Eb4 dorsal surface lateral to uncinate pro-cess.LP on Eb4 at and lateral to LE4 insertion.LI1 on most of dorsal surface of Pb2; with fewanterolateral fibers on anteromedial surface of IACon one side and extensive attachment to medial sur-face of IAC on other.LI2 on Pb3 dorsal surface posterolaterally, begin-ning anteriorly a little anterior to articulation withmedial end of Eb3 and extending posteriorly to op-posite to medial end anterior to attachment of TPb3-Pb4-Eb3 to Eb3.TD comprises TEb2 and TPb3-Pb4-Eb3. TEb2with median longitudinal raphe, which gives rise dor-sally to CT sheets covering muscles and attachesmid-anteroventrally to CT of pharyngeal roof and toCT enveloping (attaching to) anterior ends of Pb3s;muscle attaching along Eb2 dorsally to area anteriorto LE2 insertion, continuing posteriorly by diagonal
2 The familial placement of Lateolabrax is problematic. It hasbeen variously assigned to Percoidei insertae sedis (Johnson, 1984;465) or Percichthyidae (Mochizuki, 1984:123). Springer andRaasch (1995:94, 104) assigned it to Lateolabracidae. which ap-pears to be the first published usage of a family name based onthe genus. The name appeared next in Orrell et al. (2002:628).
strand of muscle with TPb3-Pb4-Eb3. TPb3-Pb4-Eb3attaching laterally on dorsal surface of Pb3, dorsoan-terior edge of Pb4. and posteromedialmost surface ofEb3, continuing posteriorly by crossing strands ofmuscle with SOD.OD3-4 origin broadly on Pb3 dorsomedially ven-tral to TEb2, insertion on anterior surface of Eb3 un-cinate process, dorsal to OD3', and on medial edgeof Eb4 uncinate process.OD3' branches off OD3-4 anteroventrally justposterior to OD3-4's passing out posteriorly fromunder TEb2 and inserting on Eb3 dorsal surface ven-tral to OD3-4 insertion on uncinate process.OP dorsally on bony posterior surface of Eb4 me-dial to uncinate process, dorsolaterally overlappingAd4 dorsomedially, ventrally on Cb5 posterodistally,partially overlapping Ad5 ventromedially.Ad 1-3 absent.Ad4 dorsally on posterior surface of Eb4 lateral toOR ventrally on Cb4 anterior to Eb4-Cb4 joint.Ad5 dorsally on posterodistal surface of Cb4, ven-trally on dorsodistal surface of Cb5 anterior to OP.SOD present.RDs narrowly separated.Additional remarks. SCL attached mid-dorsally toventroposterior cartilaginous tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. Levator processpresent on posterior edge of right-side Eb4, not pres-ent on left side; present on both sides of ? and ofcleared and stained specimen, USNM 177447.SCIAENIDAECynoscion nebulosus (Cuvier), USNM 176237, 76.8mm. Plate 121
Additional material. Cynoscion arenarius Ginsburg,USNM 120047, 117 mm.
Remarks. We noted no significant differences be-tween the muscles of C. arenarius and C. nebulosus,except that SOD in the former is strap-like and nor-mal.
Description.LEI short, with tendinous origin, insertion broadlyon anterior surface of Ebl uncinate process.LE2 on expanded bony dorsoposterior surface ofEb2.LE3 on Eb3 uncinate process dorsally and dorsaledge of Eb4 uncinate process just medial to cartilagetip.LE4 on Eb4 dorsally beginning lateral to uncinateprocess and extending to cartilaginous distal edge.LP on LE4 at and just anterior to LE4 insertion.LI1 on dorsomedialmost surface of IAC, continu-ing onto adjacent surface of Pb2, and extending to
148 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONCT that binds anteriormost ends of Pb2 and Pb3;much larger than LI2.LI2 on dorsolateral surface of Pb3 just anterior toanteromedial end of Eb3.TD comprises TEb2 and TPb3-Eb3. TEb2 withmid-longitudinal raphe giving rise dorsally to filmyCT covering muscle, attached anteriorly and mid-ventrally to CT of pharyngeal roof and laterally ondorsal surface of Eb2 to point anterior to LE2 inser-tion, continuing posteroventrally by fine, diagonalmuscle strand with TPb3-Eb3. TPb3-Eb3 on Pb3 dor-solaterally posteromedial to LI2 insertion, and onEb3 dorsoposteromedial surface, passing freely overmedial end of Eb4 before attaching to Eb3, posteri-orly continuous by fine diagonal muscle strand withvery fine SOD.OD3-4 origin on Pb3 dorsoanteriorly ventral toTEb2, insertion on anterior surface of Eb3 uncinateprocess, joining raphe with anteroventral edge of LE3insertion, and on anterior surface of Eb4 uncinateprocess.OP dorsally on Eb4 posterior surface extendingmedially beginning ventral to uncinate process, ven-trally beginning on Cb5 posterior surface distally,there joining broad raphe with PCI and extendingmedially; distinctly separate from Ad4 and SO.Ad 1-3 absent.Ad4 dorsally on Eb4 posterior surface extendinglaterally beginning from point ventral to LE4 and LPinsertions, ventrally on Cb4 dorsally medial to Eb4-Cb4 joint.Ad5 on posterodistal end of Cb4 and dorsodistalend of Cb5, forming short raphe at that point withPCI and Ad4.SOD very fine, aberrant, passes dorsal to RD ofleft side and returns to SO between RDs. ? Strap-like, continuous from one side to the other.RDs well separated.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 toothed. Eb4 levatorprocess absent.Sasaki (1989) described and illustrated in detail thedorsal gill-arch musculature of Ctenosciaena gracil-icirrhus (Metzelaar), with which Cynoscion agrees.Sasaki did not detail the attachments of his transver-sus dorsalis posterior, but his illustration (his fig. 37)conveys the impression that it is TPb3-Eb3, similarto that of Cynoscion. Sasaki also described and illus-trated (his figs. 42, 44) differences shown by Pogon-ius, Aplodinodus, and Leiostomus, highly specializedwestern Atlantic and/or freshwater inhabitants. Thedifferences shown by these three genera mainly in-volve an interrupted TEb2: broad, musculously nakedPb3 dorsal facets; and an unusually well-developedAd5.The insertion of LI1 to include IAC is uncommon.It also occurs in haemulids and bathyclupeids, both
of which also share with Cynoscion the specializedstate of lacking SCL, and in bramids, kuhliids, andlateolabracids, which have SCL.POLYNEMIDAE
Polydactylus oligodon (Gunther), USNM 364370,128 mm. Plate 122
Additional material.
?
= Filimanus xanthonema (Va-lenciennes), USNM 278199, 99.1 mm.
Description.Remarks. LE1-4 insertions musculous and tendi-nous (tendinous portions not illustrated).LEI on Ebl just lateral to tip of uncinate process.LE2 on tip of bony process on proximal half ofposterior edge of Eb2.LE3 on anterior edge of tip of Eb3 uncinate pro-cess.LE4 on bony dorsal edge of Eb4 lateral to tip ofuncinate process.LP on Eb4 beginning at lateral edge of LE4.LI1 tendinously on Pb2 dorsally posteroventral toPb2 articulation with IAC with tendon continuingonto adjacent dorsoanterior end of Pb3. ? Musclefibers reach lateral edge of Pb3.LI2 on Pb3 dorsolaterally just medial to medialend of Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 flat, roughly kidney-shaped, in two sections;triangular small anterior section fitting into mid-an-terior notch of much larger posterior section; poste-rior section with mid-longitudinal raphe continuousposteriorly through TEb2; raphe giving rise dorsallyto CT sheets attaching to skull; raphe continuousmid-ventrally with CT of pharyngeal roof; muscleattaching anterolaterally to Pb2 just medial to dor-soanteriormost cartilaginous tip and joining ventrallyto TEb2 lateral to raphe. TEb2 anterior fibers ex-tending laterally and passing dorsoposterior to pos-terior fibers and both portions attaching dorsally ashort distance lateral to medial end of Eb2, posteri-ormost fibers continue finely tendinously dorsoanter-iorly over other TEb2 fibers and attach to IAC; mus-cle continuous posteriorly by diagonal strip of musclewith TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 on Pb3 dorso-laterally, partly joining posteromedial edge of LI2 in-sertion, continuing posteriorly medial to posterome-dial end of Eb3 and attaching tendinously to it, nextcontinuing onto Pb4 dorsolaterally medial to medialend of Eb4, then continuing posteriorly by diagonalmuscle strap with SOD. ? Anterior section of TPb2is continuous posteriorly with TEb2 ventral to pos-terior section, hence, is part of TEb2.OD3?4 robust, origin broadly on Pb3 dorsomedi-ally mostly ventral to TEb2, but partially posterior to
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TEb2; posteriorly on medial edge of Eb3 uncinateprocess and anterior surface and medial edge of Eb4uncinate process, joining raphe with OP dorsally me-dial to tip of Eb4 uncinate process (raphe longer onright side). ? No raphe with OP.OP dorsally beginning on Pb4 posterolaterally andextending laterally onto Eb4 posteriorly to pointslightly ventrolateral to uncinate process, there join-ing tendon on Eb4 with dorsomedial end of Ad4 (ten-don and associated CT sheets continue to cleithrum,with long tendinous attachment of PP to CT amongsheets ventral to EM?only tendon illustrated); ven-trally on Cb5 ventrolaterally posterior to Ad5, joiningraphe with Ad5 posterolateralmost edge; mid-medi-ally confluent with SO. ? No fibers on Pb4.Remarks. Fibers on Pb4 possibly more appropri-ately allocated to SO.Ad 1-3 absent, but frayed fan-like GFM on Ebl-Cbl joint anteriorly.Ad4 dorsally on Eb4 ventrally mostly lateral to OP,ventrally on Cb4 medial to Eb4-Cb4 joint, joiningraphe ventroposteriorly with Ad5.Ad5 dorsally on Cb4 posterolaterally, anteriorlyjoining raphe with Ad4.SOD present.RDs adjacent.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Pbl with cartilaginousends. Pb2 toothed. Eb4 levator process absent.SILLAGINIDAE
Sillago sihama (Forsskal), USNM 347113, 139 mm.Plate 123
Description.LEI narrowly on tip of small bony process ex-tending anteriorly from just below cartilage tip ofEbl uncinate process, muscle fanning out dorsally.LE2 narrowly on small bony process on Eb2 dor-soposteriorly, muscle expanding dorsally.LE3 on Eb3 uncinate process anteriorly, ventroan-teriorly joining OD3-4 on Eb3.LE4 on Eb4 dorsolaterally near distal end, joinedventrolaterally by LP.LP on dorsodistalmost surface of Eb4, joining LE4insertion ventrolaterally.LI1 thin, strap-like, on dorsoanterolateralmost sur-face of Pb2 and dorsomedial surface of IAC, justimpinging on fine tendinous origin of M. Pb2-Eb2,ventral half and dorsal fourth musculous, separatedby band of clear CT lying against LE2 medially (ar-eas of CT and adjacent LE2 about equal).LI2 on Pb3 dorsolaterally medial to articulationwith Eb3, medial edge of insertion meets anterolat-eral edge of TPb3-Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2roughly heart-shaped, notched mid-anteriorly, with
deep mid-longitudinal raphe, which attaches ventrallyto CT of pharyngeal roof, extending posteriorly fromnotch and continuing across ventrally lying TEb2broad portion, which attaches dorsally to TPb2 alongraphe; raphe thence continuing across dorsal TEb2narrow portion, becoming tendinous strand and con-tinuing across TPb3-Eb3 and anterior half of SOD;anterolaterally, TPb2 attaches to Pb2 dorsoanteriorlymedial to origin of M. Pb2-Eb2. TEb2 comprisingtwo incompletely separated parts: broad anterior part,which lies mostly ventral to TPb2, and narrow pos-terior part, which is external; anterior part attenuatinglaterally, passing ventral to M. Pb2-Eb2 and meshingwith it as they extend almost to distal end of Eb2bony surface, there meeting Ad2 posteromedially;posterior part narrows considerably laterally and at-taches to medial edge of bony process bearing LE2insertion (some fibers from anterior TEb2 part alsoattache to this edge). TPb3-Eb3 not continuous mus-culously with TEb2, almost completely occludedfrom view by overlying OD3 and OD4; musclecurves anterolaterally dividing into ventral branch,which attaches to Pb3 beginning at medial edge ofLI2 insertion and continuing posteriorly to bony edgebordering articulation with Pb4 and Eb4, and dorsalbranch, which passes dorsal to medial end of Eb4and attaches to posterior edge of medial half of Eb3.(See also M. Pb2-Eb2.)OD3, OD3', OD4, essentially separate muscles.OD4 robust, originating on Pb3 dorsomedially, over-lying much of posterior half of OD3, and insertingon medial edge of uncinate process and much of pos-terior surface of Eb4, meeting LE4 and LP insertions.OD3 robust, originating on Pb3 just ventrolateral toOD4 and inserting on most of bony anterior surfaceof Eb3 uncinate process, meeting LE3 insertion ven-trally and slightly separated dorsally from distal endof OD3'. OD3 and OD4 fibers mingle at tightlyjoined Eb3 and Eb4 uncinate processes. OD3' slen-der, originating on Pb3 ventral to OD3 and insertinga little more than halfway distally on Eb3 dorsopos-teriorly, there meeting Ad3.OP with two sections: medial section broadly dor-sally on Eb4 posteriorly beginning at medial end ofbony surface and extending laterally to near LP-LE4insertions, laterally overlapping dorsomedial half oflateral section; lateral section beginning on Eb4 pos-teroventrally (anterior to medial section), extendingdorsolaterally and attaining same attachment level asmedial section, and ending laterally at distal end ofbony surface. Medial section broadly ventrally on en-tire posterior surface of lateral arm (horn) of Cb5;lateral section narrowly distally on horn, meeting OPdistally, and both meeting Ad5 posteroventrally. OPcompletely occludes Ad4 in posterior view.M. Pb2-Eb2 relatively large, originating finely,tendinously on Pb2 dorsoanteriorly with and medial
150 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
to TPb2 insertion and extending posterolaterally ontoEb2 almost to distal end of dorsal bony surface, lyingdorsal to TEb2 as it extends onto Eb2 and meshingventrally with TEb2, meeting medial end of Ad2 pos-teriorly.Adl absent, weak GFM filaments on anterodistalsurfaces of Ebl and Cbl.Ad2 small, extending from dorsodistal surface ofEb2 to anterodistal surface of Cb2, meeting distal endof M. Pb2-Eb2.Ad3 extending laterally from anteromedialmostbony surface of Eb3 to anterodistal surface of Cb3,meeting anterodistal end of OD3'.Ad4 ventrally on Eb4 anterior to lateral OP sec-tion, ventrally broadly on Cb4 anterior to Eb4-Cb4joint, posteriorly excluded from view by OP.Ad5 bulky, dorsally relatively broadly on poster-odistal surface of Cb4, there meeting distal three-fourths of Ad4 attachment posteriorly, ventrally onCb5 dorsodistally, passing anterior to OP lateral sec-tion.SOD present.RDs adjacent; together forming cup-shaped de-pression dorsally.Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. Tiny AC2 presenton both sides (also on both sides of C&S specimen.USNM 269802). Eb4 levator process absent. Broadbony flange overlapping cartilaginous distal end ofEb4 anteriorly. MULLIDAEPseudupeneus maculatus (Bloch), USNM 267508, 2specimens, 94.0-116 mm.Plate 124
Additional material.
?
= Parupeneus multifasciatus(Quoy and Gaimard), USNM 267485. 2 speci-mens, 80.0-99.7 mm.
Description.Remarks. We observed no noteworthy differencesbetween any of the specimens.LEI on dorsoanterior surface of Ebl uncinate pro-cess.LE2 on dorsal tip of expanded bony posterior edgeof Eb2.LE3 tendinously on tightly bound tips of Eb3 andEb4 uncinate processes.LE4 on dorsoposterior edge of Eb4.LP on Eb4 at and lateral to lateral edge of LE4insertion.LI1 on Pb2 ventral to tip of dorsoanterior process.LI2 on Pb3 dorsoposteriorly just medial to medialend of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2consists of an anterior transverse strip of muscle at-
taching to each IAC dorsolaterally and continuingmid-posteriorly with TEb2 and slightly laterally withpair of slender, laterally convex ribbons of musclelying dorsal to TEb2; each ribbon with mid-lateralraphe and fusing anteriorly and posteriorly withTEb2 lateral to mid-longitudinal raphe, which ex-tends continuously from anterior end of TD almostto posterior edge of TPb3-Eb3; mid-longitudinal ra-phe attached ventroanteromedially to CT of pharyn-geal roof and giving rise dorsally to CT sheets thatcover muscle and attach to skull. TEb2 extensive,attaching on dorsal surface of Eb2 laterally anteriorto LE2 insertion, posteriorly continuous (or free, var-iable in specimens) by strands of muscle with TPb3-Eb3. TPb3-Eb3 beginning anteriorly on Pb3 dorsallyjust medial to medial end of Eb3 and continuing pos-teriorly and abruptly expanding greatly laterally andattaching on Eb3 dorsomedially; continuous by di-agonal strands of muscle with SOD.OD3-4, OD3' origin on Pb3 ventral to TEb2, di-viding ventrally (OD3') just before inserting on dor-soanterior surface of Eb3 uncinate process and me-dial edge of Eb4 uncinate process; OD3' insertion onEb3 dorsal surface just ventral to Eb3 portion ofOD3-4.OP dorsally broadly on Eb4 posterior surface ven-tromedial to insertions of LE4-LP, ventrally on dor-soposterior edge of Cb5.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 posterior surface lat-eral to OP, ventrally on Cb4 dorsal surface medial toEb4-Cb4 joint.Ad5 on dorsodistal surface of Cb5 and posterodis-tal end of Cb4 and Eb4.SOD present.RDs separate.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl and Pb4 absent, UP4 present. Eb4 levatorprocess absent; however, darkly blue-stained areanear posterior point of LP insertion seems to indicatethat a cartilage tip may have been present early inontogeny. AC2 (not illustrated) present on both sidesof one specimen but absent on both sides of otherspecimen; additionally, AC2 present on both sides oftwo specimens of Mulloides flavolineatus (USNM272965, cleared and stained).
? No ACs present ineither specimen.Kim (2002:35-36) briefly described and diagram-matically illustrated the dorsal gill-arch muscles ofUpeneus vittatus Forsskal, a member of the genus hehypothesized as the sister-group to all the other mul-lid genera. Although he describes TPb2 as includingIAC among its attachments, his illustration of TPb2does not exhibit the anterolateral extensions of themuscle that attach to IAC that are present in our spec-imens of Parupeneus and Pseudupeneus.
NUMBER 1 1 151CENTROGENIIDAECentrogenys vaigiensis (Quoy and Gaimard), USNM183016, 84.7 mm;
?
= USNM 327899, 93.5 mm.Plate 125
Description.Remarks. The musculature of Centrogenys is un-usually bulky and invested with tough CT, relative tothe small size of the specimens. Determinations ofextent and attachments of TD, OD, OP, and Ad4 &5 are particularly difficult and require considerabledestruction of muscles.LEI on Ebl uncinate process lateral to tip.LE2 on mid-dorsoposterior edge of Eb2 almosttouching TEb2 posteriorly.LE3 broadly dorsally on and lateral to tip of Eb3uncinate process and narrowly on dorsal tip of bonyEb4 uncinate process.LE4 massive, on Eb4 broadly dorsoposteriorly lat-eral to all bony uncinate process (see remarks follow-ing OD3-4) and anterior to levator process, joiningraphe posteroventromedially with OP dorsally. ? Ra-phe more extensive in larger specimen.LP on Eb4 dorsoposterior margin medial to minutecartilage tip of levator process, joining raphe withLE4 insertion posterolaterally; finely continuous ven-trolaterally with OP in
?, but ignored in coding char-acter for matrix (Table 12, Appendix).LI1 on dorsalmost edge of Pb2 and dorsoanteriorsurface of Pb3 posterior to anterior end (Pb2 eden-tate, much reduced and closely applied medially tolateral surface of Pb3).LI2 on Pb3 dorsoposteriorly medial to medial endof Eb3.TD comprises TPb2, TEb2, and TPb3 (see also
?).TPb2 bun-like muscle pad on each side continuouswith each other medially by tough, thick, broad fas-
cia, which gives rise to tough CT sheets attaching toskull; muscle dorsal to medial ends of bilaterally di-vided TEb2, attaches to dorsoanterior ends of Pb2and Pb3 and, between Pbs to CT of pharyngeal roof;muscle coverage mostly absent from dorsal Pb3 sur-face. TEb2 a muscle pair, each member becomingtendinous medially and dorsally joining ventral mid-line CT between Pbs, muscle extending onto Eb2dorsally well lateral to LE2 insertion. TPb3 on Pb3posterolaterally medial to medial end of Eb4, contin-uous posteriorly by diagonal muscle filament withSOD.
?
Comprises TPb2, TEb2, and TPb3-Pb4.OD3-4 massive, origin tendinous, on Pb3 antero-laterally ventral to TEb2, there joining fascia of TPb2and TEb2; insertion extensively on Eb3 dorsoanter-iorly and on medial edge of Eb4 bony uncinate pro-cess, continuing onto posterior surface of process.Remarks. In the described smaller specimen, theEb4 uncinate process bears an easily overlooked tinycartilage tip (absent in the larger specimen) ventral
to the more dorsal bony surface that is tightly joinedto the Eb3 uncinate process. The tip is visible onlyby severing the CT joining the two processes. It ap-pears that the cartilage tip is lost with growth, andits absence is considered typical for the taxon.OP complex, bulky, possibly representing a fusionof primitively mostly separate lateral and medial sec-tions; dorsally attached to most of broad ventral Eb4surface anterior to Ad4, dorsoposteriorly joining ra-phe with LE4 ventroposteromedially, ventrolaterallybecoming tendinous and joined laterally by Ad5 andventrally by PCI, ventrolaterally broadly on Cb5 pos-terolaterally, joining raphe with PCI anterolaterallyand TV5 anterolaterally. continuous medially withSO. ? Raphe with LE4 more extensive.Ad 1-3 absent.Ad4 broadly dorsally on ventral Eb4 surface pos-terior to OP, ventrally broadly on Cb4 parallelingAd5 attachment medial to Eb4-Cb4 joint.Ad5 dorsally broadly on posterolateral surface ofCb4 and AC, ventrally broadly on Cb5 posteriorly,joining raphe with PCI dorsoanteriorly and OP ten-don anteromedially.SOD present.RDs adjacent.Additional remarks. SCL free from Bb3. TV4 ven-trally continuous across ventral surface of Cb5s, dor-sally interrupted, attaching to anterolateral surface ofeach Cb5. Pb4 and UP4 present. Pb2 edentate. IACpresent. AC4 present.Cb5s are tightly bound medially along sinuousjoint, one side is larger than other and overlaps thesmaller side dorsoanteriorly. The bones are not fusedin specimens of the sizes we examined, but it seemslikely that they might be in much larger specimens(species variously reported to attain a length of 150or 200 mm). AMBASSIDAETetracentrum caudovittatus (Norman), USNM332862, 74.1 mm. Plate 126
Additional material. ? = Ambassis buruensis,USNM 305331, 55.0 mm.
Description.LEI on Ebl uncinate process dorsoanteriorly justlateral to cartilage tip. ? On uncinate process justanteroventral to tip.LE2 on dorsalmost edge of raised posterior surfaceof Eb2.LE3 on medial edge of cartilage tip of Eb3 unci-nate process. ? On tip of process dorsally.LE4 massive; on dorsolateral surface of Eb4 reach-ing to lateralmost bony edge, there narrowly joiningwith anterolateralmost edge of LP; posteriorly, nar-
152 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
rowly, musculously continuous with OP dorsolater-
ally, just medial to posteromedial edge of LP inser-tion. ? Not continuous with OP.LP little less robust than LE4; on Eb4 dorsally neardistal bony end, ventroanteriorly joining LE4, meet-ing OP dorsolaterally. ? Slender, on dorsodistalmostbony edge of Eb4, ventromedially joining LE4.LI1 passing between acute, laterally open angleformed by TPb2 with TEb2, and inserting by slendertendon on Pb2 dorsally beginning just ventral to me-dial end of broad cartilaginous cap, tendon joins CTextending onto adjacent bony surface of Pb3 just pos-terior to dorsoanteriormost cartilage tip.LI2 on Pb3 dorsoposterolaterally adjacent to artic-ulation with medial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 moderately robust, with shallow notch mid-an-teriorly leading to short, fine, mid-longitudinal sep-tum. Septum posteriorly joins broad, oblong, thick,tough CT area, representing non-muscular mid-por-tion of TEb2 covering flat, dorsal Pb3 surfaces andattaching mid-ventrally to CT of pharyngeal roof; CTsheets arise from margins of oblong CT area and cov-er TPb2 and TEb2. TPb2 entirely anterior to TEb2,but joining oblong CT area anteriorly at point justanterior to anteromedialmost musculous portion ofTEb2, there forming angle that embraces LI1 tendi-nous portion dorsal to insertion. TPb2 musculouslyon IAC beginning about mid-dorsally and extendingmedially onto Pb2 anteriorly a little medial to jointwith IAC (? begins on IAC dorsomedially close tojoint with Pb2); muscle is continuous across dorsaland anterior surfaces of Pb2s. CT area of TEb2 moreextensive than TPb2; musculous TEb2 portion ro-bust, but narrowing considerably as it extends ontoEb2, first dorsally, then anteriorly, reaching point an-terolateral to LE2 insertion and meeting medial endof GFM2. TEb2 not continuous posteriorly withTPb3-Pb4-Eb3. TPb3-Pb4-Eb3 beginning anteriorly,tendinously on posteromedial edge of Eb3 and ad-jacent surface of Pb3 and extending posteriorly onPb4 dorsally parallel to medial end of Eb4 (superfi-cially muscle appears to attach to medial end of Eb4).
? TPb3-Eb3 attaches to Pb3 (Pb4 absent) only onsurface immediately medial to medial corner of Eb3.M. SO-Pb3 (not visible in dorsal view) arises oneach side as anteriorly extending slender ribbon ofSO longitudinal fibers that inserts on Pb3 just ven-troposterior to raised dorsoposterior surface.OD3-4 robust, anteriorly ventral to TEb2 on trans-verse posterior edge of raised anterior half of Pb3,posteriorly, broadly on Eb3 uncinate process anteri-orly and Eb4 uncinate process posteriorly, joiningnarrow raphe with OP on Eb4 medial to tip of Eb4uncinate process.OP in two or 3 irregular and almost completelyseparate sections, attaching dorsally along most of
posterior bony surface of Eb4: medial section begin-ning at uncinate process and extending medially,middle section beginning at lateral end of medial sec-tion and extending to area ventral to LP, lateral sec-tion beginning at lateral end of medial section andextending laterally almost to end of Eb4; lateralmostsection narrowly, musculously continuous with LE4(q.v.); lateralmost section ventrally on Cb5 posteri-orly and dorsally, overlapping Ad5 posteriorly; othertwo sections more medially on Cb5. ? Not continu-ous with LE4, in two or three separate sections, la-teralmost section either absent or fused ventrally withAd5, q.v.Ad 1-3 absent. GFM1 fine on anterior surface ofEbl and Cbl at and near joint. GFM2 better devel-oped, extending onto Eb2 anteriorly to about mid-length and meeting lateral end of TEb2; GFM3 welldeveloped but limited to Eb3 dorsolaterally.Ad4 dorsally. narrowly on Eb4 posteroventrallynear angle with Cb4; ventrally, narrowly on Cb4 dor-sally near angle with Eb4; completely overlappedposteriorly by OP and not visible in posterior or lat-eral views (distal ends of Cb4 and Cb5 unusuallyclosely and tightly bound together).Ad5 dorsally on Cb4 posterolaterally, ventrally onCb5 dorsolaterally, including surface of dorsodistalbony flange; muscle extends medially on Cb5 a shortdistance anterior to OP. ? Ad5 appears to have fusedwith the lateralmost part of OP, or else has been lost,such that a single muscle representing one or both,OP or Ad5, extends from the dorsolateralmost end ofCb5 to Eb4.SOD absent.RDs massive, adjacent.Additional remarks. SCL attached to ventrally ex-tending cartilaginous posterior tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. (? Pb4 absent,UP4 present). Eb4 levator process absent. Very tinyAC3 present unilaterally (? no ACs present, but seediscussion of ACs in section "Abbreviations and Def-initions for Anatomical Structures" for Ambassis sp.with AC4). Bony flange of Eb4 projects laterally dor-
sal to cartilaginous distal end. Diameter of medialend of Eb3 greater than that of Eb4. ? Slightly lessthan diameter of Eb4; however, in two cleared andstained specimens, one each as Ambassis macleayi,USNM 173817, and Ambassis sp., USNM 218805,the medial end of Eb3 is slightly larger than that ofEb4. The medial end of Eb4 is treated as smaller thanthat of Eb3. Anterior end of Cb5 on one side overlapsanterior end on other side.
CARISTIIDAE
Caristius maderensis Maul, USNM 206895, 1 14 mm.Plate 127
NUMBER 11 153
Description.LEI on dorsal edge of Ebl just lateral to cartilagetip of well-developed uncinate process, which is wellremoved laterally from direct contact with anteriorend of Eb 1
.
LE2 on Eb2 mid-dorsoposteriorly.LE3 finely, tendinously on tip of Eb3 uncinate pro-cess medially.LE4 origin tendinous; left side, tendinously on lat-eral surface of tip of Eb4 uncinate process; right side,similar to left side, but separate strip of muscle fibersextends onto bony edge of Eb4 ventrolateral to tip ofuncinate process; ventrolateral edge of insertionmeets ventroanterior edge of LP and together theyare membranously continuous with Ad4 dorsally (seealso LP).LP origin tendinous; insertion joins ventrolateraledge of LE4 insertion and extends short distance lat-erally; LP and LE4 insertions join CT. which isjoined ventrally by dorsoposterior end of medial Ad4section, and the three muscles can be easily releasedas a cohesive group from Eb4. Broad CT sheet,which attaches to lateral edges of 4th and 5th arches,attaches medially to lateralmost edge of LP insertion.LI1 on bony surface of Pb2 dorsoanteriorly begin-ning just ventral to dorsalmost cartilaginous tip.LI2 on bony surface of Pb3 dorsoanteriorly justventral to cartilaginous process articulating with me-dial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2thick, roundish pad, depressed centrally, inseparableventrally from TEb2; with pair of longitudinal raphesthat give rise to CT sheets covering muscle and at-taching to skull; CT also arising from dorsoposteriorhalf of pad perimeter, beginning at mid-lateral rapheon each side, which is also joined ventrally by TEb2medially (TPb2 and TEb2 inseparable medial to ra-phe); weak, tendinous strand extends anteriorly frommid-lateral raphe and attaches to Pb2 dorsolaterally;weak, filmy CT attaches muscle mid-ventrally be-tween Pb3s to CT of pharyngeal roof. TEb2 extendslaterally and attaches on Eb2 dorsally well lateral toLE2 insertion; diagonal muscle strand extendingfrom posteroventral surface is continuous with Pb3portion of TPb3-Eb3. TPb3-Eb3 on Pb3 dorsolater-ally beginning medial to LI2 insertion and continuingposteriorly with laterally separate muscle strands at-taching to cartilaginous medial end of Eb3 dorsallyand posterior bony edge of Eb3; ventral diagonalmuscle strands continuous with SOD.Remarks. Vertical rotation of RDs has probably re-sulted in TPb3-Eb3 and SOD also becoming almostvertical (see also remarks following OD3-4).OD3-4 origin on Pb3 dorsoanteromedially ventralto TPb2 and TEb2; muscle divides posteriorly withanterior branch inserting broadly on anterior surfaceof Eb3 beginning on uncinate process and extending
laterally, posterior branch inserting equally broadlyon anterior surface of Eb4; most of Eb4 insertion ishidden from view.OP dorsally on Eb4 posteriorly beginning medialto cartilage tip of uncinate process and extending me-dially with several medialmost fibers extending wellanteriorly and inserting on Pb4 posteriorly; ventrallyon Cb5 dorsoposterolaterally, posterior to Ad5 at-tachment, just joining raphe with Ad5 posterodistally.Ad 1-3 absent.Ad4 with lateral and medial sections. Dorsally,both sections on posterior surface of Eb4; medial sec-tion beginning lateral to OP and continuing laterallyabout half distance to end of Eb4, dorsoposterome-dially continuous with CT joining LE4 and LP (pos-terior Ad4 fibers separate from anterior fibers, whichattach to Eb4 directly); lateral section on posterolat-eral portion of Eb4; ventrally both sections join Cb4dorsally.Ad5 dorsally relatively narrowly on Cb4 postero-lateralmost bony surface, ventrally relatively narrow-ly on Cb5 dorsolaterally.SOD present.RDs separated by distance less than diameter ofone RD; extend ventrally, vertically from origins,producing excavation of SO in region of anteriorlyextending SO longitudinal fibers, then turning ante-riorly and attaching to medial edge of UP4 anteriorly,and continuing onto most of medial surface of Pb3just dorsal to tooth plate.Additional remarks. SCL attached mid-dorsally totip of posteroventrally extending cartilaginous tip ofBb3. TV4 free from Cb5s. Pb4 and UP4 present.IAC absent. Eb4 levator process absent. Small ACpresent between Eb3 and Eb4 uncinate processes onright side. Pbl bony with cartilaginous ends. Pb2toothed.
BRAMIDAEBrama brama, USNM 240541, 82.2 mm.Plate 128
Description.LEI on raised bony dorsoposterior edge of Ebllateral to tip of uncinate process.LE2 on raised bony dorsoposterior edge of Eb2.LE3 absent.LE4 on cartilaginous tip of Eb4 levator process;ventroposteriorly tendinous, continuous with broadCT sheet, which includes PP, that attaches along arch-es 4 and 5 and ventral edge of Ad5.LP fine, fuses with anterolateral edge of LE4 in-sertion.LI1 on Pb2 dorsoanterolaterally, extending slightlyonto IAC at joint with Pb2.
154 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LI2 on Pb3 posterolaterally anterior to medial endof Eb3.TD comprises TPb2, TEb2, and TPb3-Eb4. TPb2concave, roundish, with raised lateral edges, attach-ing ventroanteriorly to CT of pharyngeal roof, withmid-longitudinal raphe giving rise to CT sheets cov-ering muscle; raphe mid-laterally with CT extensionattaching to dorsoanterior cartilaginous edge of Pb2;free ventrolaterally from, but broadly continuousventrally with, TEb2. TEb2 attaching on Eb2 dorsallyanterior to LE2 insertion. TPb3-Eb4 acutely trian-gular, with apex becoming tendinous anteriorly andjoining TPb2-TEb2 mid-ventroposteriorly, mainly at-taching broadly on Eb4 bony surface dorsally medialto uncinate process with few fibers delaminating fromventral surface and inserting on Pb3 dorsopostero-medially; mid-posteroventrally continuous withSOD, which it overlaps dorsally and partially ob-scures in dorsal view.OD3-4 origin broadly on Pb3 dorsally, insertionon Eb3 uncinate process anteriorly (and, on one side,on Eb3 dorsally just ventromedial to uncinate pro-cess) and medial edge of Eb4 uncinate process.OP dorsally on Eb4 posteriorly beginning on le-vator process extending medially to uncinate process,there joining raphe with OD3-4 on one side but notother, laterally overlapping Ad4, which extends shortdistance medially anterior to OP; ventrally becomingtendinous and fascia-like and closely applied to pos-teromedial surface of Ad5; medially separable fromSO, although fibers of both are closely adjacent.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning on le-vator process anteromedial to lateralmost edge of OPand extending to distal end. ventrally broadly on Cb4dorsally medial to Eb4-Cb4 joint.Ad5 dorsally narrowly on posterodistalmost sur-face of Cb4 (at edge of Eb4-Cb4 joint), ventrallymoderately broadly on dorsolateralmost surface ofCb5, posterior surface fused to tendinous fascia-likeventral extension of OP.SOD thin, dorsally ventral to TEb3-Eb4.RDs adjacent.Additional remarks. SCL attached weakly, if at all,mid-dorsally to elongate, posteroventrally extendingcartilaginous tip of Bb3. TV4 free from Cb5s. Pb4and UP4 present. Pb2 toothed.
TOXOTIDAEToxotes jaculatrix (Pallas), USNM 331444, 94.2 mm;289931, 86.9 mm.Not illustratedDescription.LEI on Ebl dorsoposteriorly a little lateral to tipof uncinate process; ventral half of lateral edge ofmuscle tendinous.
LE2 on Eb2 dorsoposteriorly about one-third dis-tance from medial end of Eb2 and posterior to lateralend of TEb2; anterior muscle surface tendinous nearinsertion.LE3 on Eb3 beginning at medial edge of tip ofuncinate process and extending medially a short dis-tance; ventral one-fourth of muscle lateral edge ten-dinous.LE4 on Eb4 medial to levator process, ventrolat-erally meeting LP, posteroventrolaterally joining nar-row raphe with OP dorsolaterally; lateral surface ofventral third of muscle edge tendinous.Remarks. The fine raphe with OP would hardlyseem to qualify as a specialized character state, LE4joins OP, but it is present on both sides of both spec-imens and as such is here considered to be a "sling"(sensus Stiassny and Jensen, 1987).LP largest levator, on Eb4 dorsally beginning atlateralmost bony edge and extending medially, meet-ing LE4 insertion medially and cupping LE4 ven-trally; LP insertion includes dorsal edge of Eb4 le-vator process, there meeting and joining raphe withmusculous portion of PP which continues ventrallyas CT and all but encapsulates Ad5.LI1 narrowly, tendinously on Pb2 dorsoanterola-terally; tendon continuous with CT binding anteriorends of Pb2 and Pb3 (see also description of TPb2);about same size as LI2.LI2 on Pb3 dorsolaterally, meeting anterolateralend of TPb3-Pb4-Eb3 on Pb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 almost entirely anterior to TEb2, posteriorlyoverlapping a little of anterior edge of TEb2; musclecomprising an oval-shaped section on each side,which is angled a little anterolaterally from shortmid-line raphe that widens and fills notch betweenovals anteriorly and is continuous anteriorly with CTof pharyngeal roof; raphe continues posteriorly andbecomes wide area of CT replacing central portionof TEb2 (leaving flat, dorsal Pb3 surfaces naked ofmuscle) and giving rise dorsally to CT sheets cov-ering TD and attaching mid-ventrally to CT of pha-ryngeal roof. Anteroventrally, TPb2 attaches tojoined dorsoanterior ends of Pb2 and Pb3 and dor-somedialmost surface of IAC, and is attached to an-terior edge of ventral tendinous portion of LI 1 . TEb2extends laterally and attaches on Eb2 dorsally ante-
rior to LE2 insertion, muscle is dorsal to, and dis-continuous with, TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 be-gins anteriorly on Pb3 dorsoposterolaterally at medialmargin of LI2 insertion and continues posteriorly at-taching onto posteromedial edge of Eb3, with strandsextending from muscle ventral surface and attachingto Pb4 dorsally, continuous posteriorly by fine, di-agonally crossing muscle strands with SOD anteri-
orly.
NUMBER 1 1 155OD3-4 origin on Pb3 dorsomedially, insertion onEb3 anteriorly beginning at tip of uncinate processand on medial edge of Eb4 beginning at tip of un-cinate process, joining raphe with dorsomedial edgeof OP on Eb4.OP dorsally on Eb4 posteriorly beginning just ven-tral to tip of uncinate process, there meeting OD3-4, and extending laterally to just below tip of levatorprocess, joining raphe dorsally with LE4, overlappingmost to almost all of Ad4 dorsally; narrowly, ven-trally on Cb5 posteriorly near distal end, muscle ven-troposteriorly tendinous, joining raphe with Ad5 pos-teromedially.Ad 1-3 absent.Ad4 broadly, dorsally on Eb4 ventrally beginningjust anterior to medial end of OP and extending lat-erally to posterodistalmost bony surface; broadly,ventrally on Eb4 dorsally beginning about one thirdfrom medial end of bone and extending laterally todistal end of bone at inner corner of Eb4-Cb4 joint,posterolaterally meeting Ad5 on Eb4.Ad5 dorsoanteriorly on Cb4 posterolaterally, pos-teroventrally on Cb5 dorsolaterally. anterior and pos-terior surfaces almost encapsulated by CT extendingventrally from PP.SOD present.Remarks. Johnson (1993:9) reported that Toxoteslacks SOD. but did not indicate the material on whichhe based his observations. He based it, however, ona set of gill arches that are from the same lot (USNM28993 1 ) as one of the specimens on which the pres-ent description is based. If so, the muscles are dam-aged in the area where SOD would be expected andit is not possible to decide whether SOD might havebeen present. Johnson (1993:9) also reported SODabsent in Scorpis, Kuhlia, Kyphosus, and Pholidi-chthys, but, similarly, did not cite his material. John-son based his observations on the following dissec-tions: USNM 218922 (Scorpis), 289925 (Kuhlia),and 366512 (Kyphosus), all of which are damaged inthe area where SOD might be expected to occur. Ofthese genera, and based on specimens dissected forthe present study, we have found SOD lacking onlyin Pholidichthys. Our observations on these threegenera are based on the following specimens: Scorpis
sp. (USNM 339348) and Kyphosus (USNM 366512),both not otherwise discussed in the present study, andspecimens cited in the description of Kuhlia (Kuhli-idae).RDs adjacent.Additional remarks. SCL attached mid-dorsally totip of ventrally extending cartilaginous posterior endof Bb3. TV4 free from Cb5s. Pbl bony with cartilageends. Pb4 and UP4 present. Very reduced Eb4 flangepresent.
PLESIOPIDAE
Assessor macneilli Whitley, USNM 269466, 47.7mm; USNM 274581, 51.7 mm.Plate 129
Additional material.
?
= Paraplesiops poweri Ogil-by, USNM 274579, 92.2 mm. ? = Acanthoclinusfuscus Jenyns, USNM 339246, not measured.
Description (based on smaller specimen of Assessorwith remarks based on larger specimen).LEI on dorsoposterior rim of Ebl beginning onuncinate process ventroanterolaterally and extendinga short distance laterally.LE2 on dorsalmost edge of raised, bony dorsopos-terior rim of Eb2, well lateral to TEb2 insertion.
?Inserts posterior to distal end of TEb2.LE3 on Eb3 immediately ventroanterior to carti-lage tip of uncinate process.LE4 finely, tendinously on Eb4 dorsally a little me-dial to dorsodistalmost end of bone, there meetingLP insertion (in smaller specimen, muscle insertsabove levator process, which is absent in larger spec-imen).
? ? Muscle massive, inserts broadly on Eb4dorsally beginning laterally just anterior to levatorprocess in
?
(process absent in
?), meeting OD3-4posterodistally on Eb4; fusing with LP ventroanter-iorly (or ventromedially, depending somewhat onviewing orientation) a little dorsal to insertion.LP narrowly on dorsodistalmost end of Eb4 bonysurface, meeting LE4 insertion ventroposteriorly.
?Muscle massive, ventroposteriorly joining raphe withAd5 dorsally (thus, forming an unusual LP-Ad5sling), fusing ventroanteriorly with LE4. ? Massivelike ?. but does not form LP-Ad5 sling.LI1 larger than LI2, ventrally becoming moderate-ly long, slender tendon passing ventral to TPb2 andinserting at junction between medial edge of Pb2 andadjacent impinging lateral edge of Pb3.
? ? Mus-culously and tendinously on Pb2 dorsoanteriorly pos-terior to anteriormost edge, tendinous portion con-tinuing medially as strong, slender tendon insertingon Pb3 dorsal facet mid-dorsoanteriorly.LI2 on Pb3 dorsoposterolaterally just medial tojoint with Eb3; medial edge of insertion separatedfrom lateral edge of TPb3-Eb3 in one specimen,meeting lateral edge of TPb3-Eb3 in other specimen.
?
Medial edge meets lateral edge of TPb3-Eb3.TD flat, comprises TPb2, TEb2, and TPb3-Eb3.TD with mid-longitudinal raphe, which gives risedorsally to filmy CT sheets covering TD. TPb2 ananterolaterally extending muscle on each side, in-serting on dorsoanterior surface of Pb2, almost com-pletely anterior to TEb2. TEb2 dorsoanteromediallyjoining TPb2 ventrally; main anterior portion contin-uous dorsally from one side to the other ventral toTPb2, lesser posterior portion interrupted medially
154 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LI2 on Pb3 posterolaterally anterior to medial endof Eb3.TD comprises TPb2, TEb2, and TPb3-Eb4. TPb2concave, roundish, with raised lateral edges, attach-ing ventroanteriorly to CT of pharyngeal roof, withmid-longitudinal raphe giving rise to CT sheets cov-ering muscle; raphe mid-laterally with CT extensionattaching to dorsoanterior cartilaginous edge of Pb2;free ventrolaterally from, but broadly continuousventrally with, TEb2. TEb2 attaching on Eb2 dorsallyanterior to LE2 insertion. TPb3-Eb4 acutely trian-gular, with apex becoming tendinous anteriorly andjoining TPb2-TEb2 mid-ventroposteriorly, mainly at-taching broadly on Eb4 bony surface dorsally medialto uncinate process with few fibers delaminating fromventral surface and inserting on Pb3 dorsopostero-medially; mid-posteroventrally continuous withSOD, which it overlaps dorsally and partially ob-scures in dorsal view.OD3-4 origin broadly on Pb3 dorsally, insertionon Eb3 uncinate process anteriorly (and, on one side,on Eb3 dorsally just ventromedial to uncinate pro-cess) and medial edge of Eb4 uncinate process.OP dorsally on Eb4 posteriorly beginning on le-vator process extending medially to uncinate process,there joining raphe with OD3-4 on one side but notother, laterally overlapping Ad4, which extends shortdistance medially anterior to OP; ventrally becomingtendinous and fascia-like and closely applied to pos-teromedial surface of Ad5; medially separable fromSO, although fibers of both are closely adjacent.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning on le-vator process anteromedial to lateralmost edge of OPand extending to distal end, ventrally broadly on Cb4dorsally medial to Eb4-Cb4 joint.Ad5 dorsally narrowly on posterodistalmost sur-face of Cb4 (at edge of Eb4-Cb4 joint), ventrallymoderately broadly on dorsolateralmost surface ofCb5, posterior surface fused to tendinous fascia-likeventral extension of OP.SOD thin, dorsally ventral to TEb3-Eb4.RDs adjacent.Additional remarks. SCL attached weakly, if at all,mid-dorsally to elongate, posteroventrally extendingcartilaginous tip of Bb3. TV4 free from Cb5s. Pb4and UP4 present. Pb2 toothed.
TOXOTIDAEToxotes jaculatrix (Pallas), USNM 331444, 94.2 mm;289931, 86.9 mm.Not illustratedDescription.LEI on Ebl dorsoposteriorly a little lateral to tipof uncinate process; ventral half of lateral edge ofmuscle tendinous.
LE2 on Eb2 dorsoposteriorly about one-third dis-tance from medial end of Eb2 and posterior to lateralend of TEb2; anterior muscle surface tendinous nearinsertion.LE3 on Eb3 beginning at medial edge of tip ofuncinate process and extending medially a short dis-tance; ventral one-fourth of muscle lateral edge ten-dinous.LE4 on Eb4 medial to levator process, ventrolat-erally meeting LP, posteroventrolaterally joining nar-row raphe with OP dorsolaterally; lateral surface ofventral third of muscle edge tendinous.Remarks. The fine raphe with OP would hardlyseem to qualify as a specialized character state, LE4joins OP, but it is present on both sides of both spec-imens and as such is here considered to be a "sling"(sensus Stiassny and Jensen, 1987).LP largest levator, on Eb4 dorsally beginning atlateralmost bony edge and extending medially, meet-ing LE4 insertion medially and cupping LE4 ven-trally; LP insertion includes dorsal edge of Eb4 le-vator process, there meeting and joining raphe withmusculous portion of PR which continues ventrallyas CT and all but encapsulates Ad5.LI1 narrowly, tendinously on Pb2 dorsoanterola-terally; tendon continuous with CT binding anteriorends of Pb2 and Pb3 (see also description of TPb2);about same size as LI2.LI2 on Pb3 dorsolaterally, meeting anterolateralend of TPb3-Pb4-Eb3 on Pb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 almost entirely anterior to TEb2, posteriorlyoverlapping a little of anterior edge of TEb2; musclecomprising an oval-shaped section on each side,which is angled a little anterolaterally from shortmid-line raphe that widens and fills notch betweenovals anteriorly and is continuous anteriorly with CTof pharyngeal roof; raphe continues posteriorly andbecomes wide area of CT replacing central portionof TEb2 (leaving flat, dorsal Pb3 surfaces naked ofmuscle) and giving rise dorsally to CT sheets cov-ering TD and attaching mid-ventrally to CT of pha-ryngeal roof. Anteroventrally, TPb2 attaches tojoined dorsoanterior ends of Pb2 and Pb3 and dor-somedialmost surface of IAC, and is attached to an-terior edge of ventral tendinous portion of LI1. TEb2extends laterally and attaches on Eb2 dorsally ante-
rior to LE2 insertion, muscle is dorsal to, and dis-continuous with, TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 be-gins anteriorly on Pb3 dorsoposterolaterally at medialmargin of LI2 insertion and continues posteriorly at-taching onto posteromedial edge of Eb3, with strandsextending from muscle ventral surface and attachingto Pb4 dorsally, continuous posteriorly by fine, di-agonally crossing muscle strands with SOD anteri-
orly.
NUMBER 1 1 155OD3-4 origin on Pb3 dorsomedially, insertion onEb3 anteriorly beginning at tip of uncinate processand on medial edge of Eb4 beginning at tip of un-cinate process, joining raphe with dorsomedial edgeof OP on Eb4.OP dorsally on Eb4 posteriorly beginning just ven-tral to tip of uncinate process, there meeting OD3?4, and extending laterally to just below tip of levatorprocess, joining raphe dorsally with LE4, overlappingmost to almost all of Ad4 dorsally; narrowly, ven-trally on Cb5 posteriorly near distal end, muscle ven-troposteriorly tendinous, joining raphe with Ad5 pos-teromedial^.Ad 1-3 absent.Ad4 broadly, dorsally on Eb4 ventrally beginningjust anterior to medial end of OP and extending lat-erally to posterodistalmost bony surface; broadly,ventrally on Eb4 dorsally beginning about one thirdfrom medial end of bone and extending laterally todistal end of bone at inner corner of Eb4-Cb4 joint,posterolateral^ meeting Ad5 on Eb4.Ad5 dorsoanteriorly on Cb4 posterolaterally, pos-teroventrally on Cb5 dorsolaterally. anterior and pos-terior surfaces almost encapsulated by CT extendingventrally from PP.SOD present.Remarks. Johnson (1993:9) reported that Toxoteslacks SOD. but did not indicate the material on whichhe based his observations. He based it, however, ona set of gill arches that are from the same lot (USNM28993 1 ) as one of the specimens on which the pres-ent description is based. If so, the muscles are dam-aged in the area where SOD would be expected andit is not possible to decide whether SOD might havebeen present. Johnson (1993:9) also reported SODabsent in Scorpis, Kuhlia, Kyphosus, and Pholidi-chthys, but, similarly, did not cite his material. John-son based his observations on the following dissec-tions: USNM 218922 (Scorpis), 289925 (Kuhlia),and 366512 (Kyphosus), all of which are damaged inthe area where SOD might be expected to occur. Ofthese genera, and based on specimens dissected forthe present study, we have found SOD lacking onlyin Pholidichthys. Our observations on these threegenera are based on the following specimens: Scorpis
sp. (USNM 339348) and Kyphosus (USNM 366512),both not otherwise discussed in the present study, andspecimens cited in the description of Kuhlia (Kuhli-idae).RDs adjacent.Additional remarks. SCL attached mid-dorsally totip of ventrally extending cartilaginous posterior endof Bb3. TV4 free from Cb5s. Pbl bony with cartilageends. Pb4 and UP4 present. Very reduced Eb4 flangepresent.
PLESIOPIDAE
Assessor macneilli Whitley, USNM 269466, 47.7mm; USNM 274581, 51.7 mm.Plate 129
Additional material.
?
= Paraplesiops poweri Ogil-by, USNM 274579, 92.2 mm. ? = Acanthoclinusfuscus Jenyns, USNM 339246, not measured.
Description (based on smaller specimen of Assessorwith remarks based on larger specimen).LEI on dorsoposterior rim of Ebl beginning onuncinate process ventroanterolaterally and extendinga short distance laterally.LE2 on dorsalmost edge of raised, bony dorsopos-terior rim of Eb2, well lateral to TEb2 insertion. ?Inserts posterior to distal end of TEb2.LE3 on Eb3 immediately ventroanterior to carti-lage tip of uncinate process.LE4 finely, tendinously on Eb4 dorsally a little me-dial to dorsodistalmost end of bone, there meetingLP insertion (in smaller specimen, muscle insertsabove levator process, which is absent in larger spec-imen).
?
? Muscle massive, inserts broadly on Eb4dorsally beginning laterally just anterior to levatorprocess in ? (process absent in
?), meeting OD3?
4
posterodistally on Eb4; fusing with LP ventroanter-iorly (or ventromedially, depending somewhat onviewing orientation) a little dorsal to insertion.LP narrowly on dorsodistalmost end of Eb4 bonysurface, meeting LE4 insertion ventroposteriorly. ?Muscle massive, ventroposteriorly joining raphe withAd5 dorsally (thus, forming an unusual LP-Ad5
sling), fusing ventroanteriorly with LE4. ? Massivelike
?, but does not form LP-Ad5 sling.LI1 larger than LI2, ventrally becoming moderate-ly long, slender tendon passing ventral to TPb2 andinserting at junction between medial edge of Pb2 andadjacent impinging lateral edge of Pb3. ? ? Mus-culously and tendinously on Pb2 dorsoanteriorly pos-terior to anteriormost edge, tendinous portion con-tinuing medially as strong, slender tendon insertingon Pb3 dorsal facet mid-dorsoanteriorly.LI2 on Pb3 dorsoposterolaterally just medial tojoint with Eb3; medial edge of insertion separatedfrom lateral edge of TPb3-Eb3 in one specimen,meeting lateral edge of TPb3-Eb3 in other specimen.
?
Medial edge meets lateral edge of TPb3-Eb3.TD flat, comprises TPb2, TEb2, and TPb3-Eb3.TD with mid-longitudinal raphe, which gives risedorsally to filmy CT sheets covering TD. TPb2 ananterolateral^ extending muscle on each side, in-serting on dorsoanterior surface of Pb2, almost com-pletely anterior to TEb2. TEb2 dorsoanteromediallyjoining TPb2 ventrally; main anterior portion contin-uous dorsally from one side to the other ventral toTPb2, lesser posterior portion interrupted medially
156 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and joining lateral margin of small, roundish CT padoverlying naked, fiat Pb3 facet posteriorly; muscleextends laterally, attaching to Eb2 dorsally medial toLE2 insertion; muscle slip (absent in larger speci-men) arises from TEb2 mid-anteriorly on right sideand extends medially as fine tendon, which inserts onPb2 near LI1 insertion. TPb3-Eb3 mid-anteromedi-ally continuous by CT with TEb2, attaches to Pb3dorsoposteriorly along LI2 insertion and has fine CTattachment to anteromedialmost edge of Eb3; musclecontinuous posteriorly by fine, diagonal musclestrands with SOD.Remarks. The flat Pb3 dorsal facet forms the dor-somedialmost surface of Pb3. In the larger specimenof Assessor, TEb2 is continuous only by CT acrossthe facets, the anterior one-third of each facet is ven-tral to TPb2; the posterior two-thirds are naked mus-culously. In the smaller specimen, only a small areaof the facet is naked.
?
Mid-dorsal CT area giving rise dorsally to toughCT sheets; most of Pb3 dorsal surface not coveredby muscle; muscle fibers of anterior portion of TPb2more-or-less transversely oriented, those of postero-lateral portion thinner, more longitudinally oriented,two portions meet at fine CT notch laterally nearpoint where ventroanterior edge of LI1 impinges onTPb2. TEb2 extends laterally to point anterior to LE2insertion.
? TPb2 and TEb2 musculously continuous acrossPb3s; TEb2 like Assessor.OD3-4 origin on Pb3 along and ventral to lateraledge of flat, dorsal Pb3 facet (ventral to roundish CTpad), insertion on Eb3 uncinate process dorsoanter-iorly and Eb4 uncinate process medially. ? ? Musclemassive.OP in two parts, medial part dorsally on Eb4 pos-teriorly beginning just ventral to uncinate process andextending short distance laterally, posteriorly over-lapping medial edge of lateral OP part, which extendslaterally on Eb4 to below medial edge of LE4-LPinsertions; ventrally two parts join raphe with ventro-posterior surface of Ad5 dorsal to attachment to Cb5,and attach to dorsal surface of small, posterodistallyprojecting bony Cb5 process that extends laterallypast medial margin of cartilage tip. OP of larger spec-imen comprises single part occupying same area astwo parts of smaller specimen. ? Lateral portionmuch thicker than medial portion, overlying medialportion posteriorly. ? Comprises single part.Ad 1-3 absent; fine GFMs present. ? GFMs mod-erately well developed, that on arch 2 extending dor-sally on Eb2 and just meeting distal end of TEb2.Ad4 relatively small, dorsally on Eb4 ventropos-teriorly lateral to lateral edge of OP, ventrally, nar-rowly on Cb4 medial to Eb4-Cb4 joint.Ad5 dorsally moderately narrowly on Cb4 pos-terolaterally; ventrally very narrowly on Cb5 dorso-
distally. ? Very well developed, joining raphe dor-sally with LP. ? Like Assessor.SOD relatively fine.RDs narrowly separate.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 absent, UP4 present. Pbl mostly bony.Pb2 toothed. IAC present.Although, TV4 is free from Cb5s in Paraplesiops,it consists of two layers, dorsal and ventral. The ven-tral layer is musculously continuous and unmodifiedas it passes across the Cb5s, whereas, the dorsal layerchanges to thick CT as it crosses the flattened ven-troanterior tips of the Cb5s. Bony Eb4 flange present.Medial end of Eb4 smaller than that of Eb3, also trueof Trachinops (USNM 269557), Plesiops (USNM264268), and Acanthoclinus (USNM 200546), allcleared and stained specimens, and ?, but medial endof Eb4 is larger than medial end of Eb3 in Paraple-siops.Mooi (1993:291) hypothesized the phylogeny ofthe Plesiopidae. He recognized Trachinops as com-prising the basalmost clade of the family, Assessoras comprising the next clade, and Paraplesiops andAcanthoclinus. separately, as members of the next,and final two, clades. We, therefore, consider that thestate of the medial end of Eb4 in Paraplesiops isapomorphic for that genus, as is the position of thelateral end of TEb2 reaching laterally anterior to LE2insertion, which is usually unspecialized for perco-morphs. On the other hand, the musculously uninter-rupted condition of TPb2 and TEb2 of Acanthocli-nus, which generally appears to be plesiomorphic inperciforms, is apomorphic for Acanthoclinus.
PERCIDAE
Perca flavescens (Mitchill), USNM 193135, 91.7mm. Plate 130
Additional material. ? = Percina caprodes (Rafin-esque), USNM 230758, 2:101-103 mm. ? = Per-cina sclera (Swain), USNM 161594, 91.5 mm.
Description.LEI on Ebl uncinate process just lateral to carti-lage tip. ? ? Uncinate process appears to havemoved medially and fused with cartilaginous medialend of Ebl; anterior arm of Ebl represented by slightcartilaginous bulge of ventromedial end in ?, whichdoes not join Pb2; bulge absent in ?; muscle insertson raised bony process on Ebl mid-dorsoposteriorly.LE2 on bony dorsoposterior surface of Eb2.LE3 on cartilaginous tip of Eb3 uncinate process.LE4 dorsoposteriorly on Eb4 just lateral to unci-nate process.LP on Eb4 at and lateral to LE4 insertion.
NUMBER II 157
LI1 mainly on Pb2 dorsoanteriorly, but few musclestrands appear to continue into CT covering Pb2 andanterior end of closely joined Pb3.
? ? Mainly onPb2, but some muscle strands clearly continue ontoPb3.LI2 on Pb3 dorsoposteriorly.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 thin, flat, laterally curving ribbon of muscle oneach side arising from TEb2 at anterior end of mid-longitudinal raphe and continuing to posterior end ofraphe; not attached to Pb2. TEb2 with mid-anteriorand mid-posterior notches joined by mid-longitudinalraphe giving rise dorsally to sheets of CT and con-necting ventrally to CT of pharyngeal roof; antero-lateral and posterolateral converging fibers fusing be-fore attaching on most of dorsal surface of Eb2.TPb3-Pb4-Eb3 on Pb3 dorsally just medial to LI2insertion, extending onto dorsomedial end of Eb3and. weakly, onto dorsal surface of Pb4, posteriorlycontinuous by diagonal slip of muscle with SOD. ?TPb2 thin, flat, laterally curving ribbon of muscle ononly one side of each of two specimens.
?
? TEb2scarcely notched posteriorly; TPb3-Pb4-Eb3 on Pb3dorsoposteriorly, Pb4 dorsally, and Eb3 dorsoposter-omedialmost end.OD3-4 origin broadly on Pb2 posterior to LI1 in-sertion and continuing broadly medially onto Pb3ventral to TEb2, inserts on anterior surface of Eb3uncinate process and anterior surface and medialedge of Eb4 uncinate process.OP dorsally on all of bony posterior surface of Eb4medial to LE4 insertion, ventrally on Cb5 bony sur-face beginning medial to distal end, overlapping pos-teriorly much of Ad5 posterior surface and joiningAd5 attachment to Cb5.M. Pb2-Eb2 absent. ? On one side of one speci-men, small strip of muscle extends from lateral edgeof Pb2 to anterior edge of Eb2 anteroventral to TEb2.
? On both sides, relatively broad strap of muscleoriginates on lateral edge of Pb2 ventral to OD3-4and attaches to anterior edge of Eb2 ventral to TEb2.Ad 1-3 absent. ? ? Adl on dorsoanterior surfaces
of, and covering joint between, Ebl and Cbl; Ad2beginning medially at raphe with lateral end of TEb2,extending laterally and spreading anterolaterally andattaching to anterior surfaces of Eb2-Cb2 at joint;Ad3 broad dorsomedially, on dorsolateral surface ofEb3, spreading anterolaterally and attaching to ante-
rior surfaces of Eb3-Cb3 joint.Remarks. GFMs (not illustrated) present in Percain positions similar to Ad 1-3, which attach to GFMsin
?
and
?, but relatively weakly developed, hence,difference in interpretation.Ad4 broadly dorsally on posterior surface of Eb4lateral to OP; ventrally, less broadly on Cb4 dorsalsurface anterior to Eb4-Cb4 joint, joining Ad5 at-tachment on Eb4.
Ad5 on Cb4 posterolateral surface and Cb5 dor-soposterior surface beginning lateral to OP attach-ment and extending medially anterior to OP attach-ment.SOD present.RDs slightly separated.Additional remarks. SCL attached mid-dorsally totip of cartilaginous ventroposterior end of Bb3. TV4free from Cb5s. Pb4 and UP4 present. IAC well de-veloped. LE4 levator process absent.
?
? Pbl absent.
? IAC a tiny sphere on both sides in larger specimen,absent in smaller specimen. ? IAC absent.
CEPOLIDAECepola rubescens Linnaeus, USNM 285452, ca. 260mm. Plate 131
Additional material. ? = Acanthocepola limbata(Valenciennes), NSMT P. 64733, 375 mm.
Description.Remarks. All levators and RDs relatively slender.LEI origin by short, fine tendon; insertion on dor-soanterior surface of Ebl just ventral to cartilagetipped process lateral to uncinate process, which ar-ticulates with IAC; dorsoanterior surface of LEI, andentire anterior surface of Pbl, attaches to posteriorsurface of CT arising from anterior edge of Ebl.LE2 on dorsoposteriormost edge of Eb2 posteriorto lateral end of TEb2.LE3 on tip of Eb3 uncinate process.LE4 on bony surface of Eb4 immediately medialto cartilage tip of levator process, posteriorly joiningLP insertion.
?
On levator process together with LP.LP beginning on tip of levator process and ex-tending medially, joining LE4 insertion posteriorly.
?
On levator process with LE4.LI1 divides into anterior and posterior branches asit passes over Pb2 articulation with IAC, anteriorbranch inserts on dorsalmost surface of Pb2, posteriorbranch, inserts on Pb2 dorsally well ventral to ante-
rior branch and continues onto adjacent lateral edgeof Pb3 anterior process. ? IAC absent on both sides;insertion divided on only one side; insertion on otherside passes posterior to Eb2-Pb2 joint and inserts likeposterior branch of other side.LI2 on Pb3 dorsoposterolaterally immediately me-dial to medial end of Eb3 anteriorly.TD comprises TEb2, M. TEb2-Pb2, and TPb3-Eb3. TEb2 with mid-longitudinal raphe giving rise toCT pad dorsally and attaching anteriorly to CT ofpharyngeal roof and laterally on Eb2 dorsally anteriorto LE2 insertion; muscle is joined ventromedially byM. TEb2-Pb2, and is continuous mid-posteroventral-ly by fine, diagonal muscle strap with TPb3-Eb3 dor-
158 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
soanteriorly (? TEb2 not continuous with TPb3-Eb3). M. TEb2-Pb2 broad, strap-like, originatingventrally from TEb2 posteriorly and extending ante-
riorly along mid-longitudinal raphe for about halflength of raphe (? muscle originates along mid-pos-terior half of raphe); muscle extends ventroanterola-terally dorsal to OD3-4 origin and inserts on Pb2dorsomedially just posterior to base of dorsoanteriorprocess. TPb3-Eb3 on Pb3 dorsally beginning ante-
rior to joint with medial end of Eb3 and extendingposteriorly and attaching to posteromedial corner ofEb3 on both sides, but on one side a fine, probablyanomalous, muscle strand extends to opposite sideand attaches to anteromedial edge of Eb4. (? No at-tachment to Eb4); muscle continuous posteriorly withSOD. ? TPb3-Eb3 joins raphe posterolateral^/ oneach side with SO muscle strap passing dorsoanter-iorly immediately lateral to OP on Eb4.Remarks. M. TEb2-Pb2 is not present in any otheracanthomorph we examined. It is difficult to equatethe muscle with TPb2, which arises dorsal and/or an-terior to TEb2. If present in Owstonia and/or Sphen-anthias, M. TEb2-Pb2 will either represent anothersynapomorphy of the Cepolidae or it will define amonophyletic group within the family.OD3-4 origin of Pb3 dorsally ventral to M.TEb2-Pb2, beginning a little posterior to anteriormost tipof Pb3; insertion on Eb3 anteriorly just ventral to tipof uncinate process and on bony medial edge of Eb4uncinate process.OP strap-like; dorsally on Eb4 posteriorly begin-ning medially ventral to tip of uncinate process andextending laterally about half distance between un-cinate and levator processes (? extends to or almostto levator process); ventrally on Cb5 beginning lat-erally on distal cartilaginous end medial to Ad5, andextending medially a short distance and meetingTV5.Ad 1-3 absent. GFM1-3 moderately developed.Ad4 on Eb4 dorsoposteriorly beginning mediallyjust medial to lateral edge of OP and extending lat-erally to distalmost bony edge, ventrally on Cb4 dor-sally beginning at Eb4-Cb4 joint and extending me-dially a short distance.Ad5 short, ventrally on Cb5 posterodistally, dor-sally on Cb4 posterodistally and AC4 posteriorly.
?does not join AC4.SOD slender.RDs separated by distance greater than diameter ofone RD. ? Separated by about one-half RD diameter.Additional remarks. SCL attached mid-dorsally toventrally extending posterior cartilaginous tip of Bb3.TV4 free from Cb5s. Pb4 and UP4 present. ? SCLattached mid-posteriorly to ventroanteriormost sur-face of ventrally extending posterior cartilaginous tipof Bb3. IAC small, ball-like; AC4 relatively small.
? IAC absent; AC4 relatively large.
The presence of two uncinate processes on Ebl isunique among the Actinopterygii (the tip of the la-teralmost process is attached by CT to Eb2 anteri-orly). Gill and Mooi (1993:331 and fig. 2) noted thattwo rod-like uncinate processes on Ebl supportmonophyly of the Cepolidae and Owstoniidae, whichthey recognized, as do we, as a single family, Ce-polidae. Nelson (1994:378), however, recognizedboth families in a superfamily Cepoloidea.
CALLANTHIIDAE
Callanthias allporti Gunther, USNM 350940, 129mm. Plate 132
Additional material. Callanthias australis Ogilby,AMS 1.18079-002, 87.9 mm. ? = Grammatonotuslaysanus Gilbert, BPBM 22757, 126 mm.
Description.Remarks. We noted no substantive differences inthe musculature between the two species of Callan-thias, or in the skeletal structure in both genera. Dif-ferences reported are for Grammatonotus.LEI on dorsal edge of Ebl immediately lateral tobroad uncinate process, continuing very slightly ontoCT joining Ebl and Eb2.LE2 on posterodorsal edge of Eb2 a little lateralto mid-length and continuing on CT between Eb2and Eb3; muscle is cleanly divisible into two sec-tions, one on bone and one on CT.LE3 on tip of Eb3 uncinate process.LE4 on tip of Eb4 levator process.LP on Eb4 levator process at and lateral to LE4insertion.LI1 on dorsal bony edge of Pb2 uncinate processjust posterior to cartilaginous tip.LI2 on Pb3 dorsolaterally at and anterior to medialend of Eb3.TD comprises TEb2, and TPb3-Pb4-Eb3. ThickCT pad attaches anterolaterally to anteriormost endof Pb2 and mid-anteroventrally to anterior end ofmid-longitudinal TEb2 raphe, which joins CT of pha-ryngeal roof. TEb2 attaches ventrally along mid-lon-gitudinal raphe to CT of pharyngeal roof; muscle ex-tends laterally attaching along most of Eb2 dorsalsurface; TEb2 not continuous with TPb3-Pb4-Eb3.TPb3-Pb4-Eb3 on Pb3 dorsal bony surface medial tomedial end of Eb3, continuing posteriorly onto pos-teromedialmost edge of Eb3 and adjacent dorsome-dial edge of Pb4, continuous posteriorly by diagonalmuscle straps with SOD. ? Attachment to medial endof Eb3 absent, hence TPb3-Pb4.OD3-4, OD3' origin on dorsoanterior surface ofPb3 ventral to TEb2, splitting shortly into dorsal(OD3) and ventral (OD4; not illustrated) sections.
NUMBER 11 159
Dorsal section completely overlies ventral section,hiding it in dorsal view; dorsal section splits laterallywith ventroanterior portion (OD3') inserting on Eb3dorsal surface well ventral to uncinate process anddorsal portion (OD3) inserting on anterior surface ofuncinate process. Ventral section inserts on Eb4 an-terior surface medial to levator process.
?
OD3' ab-sent; OD3-4 attachment on Eb4 only to tiny sectionof dorsal edge of Eb4 ventromedial to levator process(Eb4 section essentially absent).Remarks. The Eb3 uncinate process is tightlyjoined to Eb4, which has lost the uncinate process.That the Eb4 process posterior to this joining is nota displaced uncinate process is evidenced by the in-sertions of LE4 and LP on the process. These twomuscles rarely, if ever, insert on the Eb4 uncinateprocess.OP broadly on posterior surface of Eb4 medial tobase of levator process and broadly on posterodistalsurface of Cb5 posterior to Ad5.Adl-3 moderately developed, but gill-filaments at-tach to these muscles and they could be interpretedsimply as well-developed cord-like GFMs, present onEbs and extending only to dorsoanteriormost surfaceof respective Cbs; however, Ads 1-3 well developedin ?.Ad4 broadly on posterior surface of Eb4 beginning
at levator process and extending laterally, ventrallyon posterodorsal surface of Cb4 medial to Eb4-Cb4joint.Ad5 small, on posterodistal surface of Cb5, partlyanterior to OP ventral attachment, and on distalmostcartilaginous tip of Cb4 posteriorly.SOD present.RDs well separated.Additional remarks. SCL attached mid-dorsally totip of ventrally extending posterior cartilaginous endof Bb3. TV4 free from Cb5s. Pbl cartilaginous. Pb4and UP4 present IAC present. Eb4 uncinate processabsent. ? SCL questionably absent (should be veri-fied in another specimen); Pbl cartilaginous, veryslender, embedded in tough CT; one side in two seg-ments.
GERREIDAEGerres cinereus Eigenmann, USNM 331928, 90.4mm; USNM 337861, 2: 60.9-95.7 mm.Plate 133
Additional material.342106, 92.7 mm. Eitcinostomus sp., USNM
Description.LEI tendinously and musculously on anterior sur-face of Eb 1 uncinate process just ventral to cartilag-inous tip.
LE2 narrowly tendinously and musculously on tipof bony process on Eb2.LE3 tendinously and musculously on dorsomedialbony edge of Eb3 uncinate process.LE4 tendinously on mid-dorsoposterior surface ofEb4, fusing with medial end of LP insertion. ? In-sertion not fused with that of LP.LP massive, on much of dorsoposterior edge ofEb4, fusing with LE4 insertion, meeting, but not join-ing OP dorsally.
? Smaller than LE4, insertion en-tirely lateral to that of LE4.LI1 tendinously and musculously on bony poste-
rior surface of Pb2 dorsoanterior process ventral tocartilaginous tip, with tendon continuing ventrally toposterior surface of dorsoanteriormost Pb3 process.LI2 slender tendinously on Pb3 posterolaterallyventral to medial end of Eb3, which overlies Pb3.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2pair of muscles, each member joined to lateral edgeof thick CT pad covering Pb3 dorsal articulating fac-
ets and attaching on medial half of Eb2 dorsal surfacelateral to LE2 insertion. TPb3-Eb3-Eb4 on dorsopos-terolateral surface of Pb3, posteromedialmost edge ofEb3, and dorsomedialmost surface of Eb4.OD3-4, OD3'. OD3-4 origin on lateral edge ofPb3 dorsal articulating facet, insertion on medialedge and anterior surface of Eb3 uncinate processand medial edge and posterior surface of Eb4 unci-nate process; just posterior to origin, OD3' originateswith OD3-4, diverges well laterally and inserts ondorsal surface of Eb3 ventral to uncinate process,forming raphe with posteromedial end of Ad3; OD3'absent in 60.9 mm specimen and ?.OP massive, on much of posterior surface of Eb4and extending ventrally and attaching by tendons toposterior surface of Cb5 somewhat medial to distalend; Ad5 fibers joining OP tendons anteriorly; slen-der, separate OP strap attaches dorsally narrowly toEb4 medial to massive OP attachment and to Cb5medial to Ad5 attachment.Adl relatively small, restricted, spans anterior sur-face of Ebl-Cbl joint.Ad2 begins on Eb2 dorsoanteriorly adjacent to lat-eral end of TEb2 and expands laterally attaching overanterior surface of Eb2-Cb2 joint.Ad3 begins medially at raphe with lateral end ofOD3' and expands laterally attaching over anteriorsurface of Eb3-Cb3 joint. In smallest and largestspecimens and ? occupies most of dorsal and an-terolateral surfaces of Eb3 (area of attachment equalsthat of OD3' + Ad3 of 90.4 mm specimen).Ad4 on ventral surface of Eb4 and dorsal surfaceof Cb4 medial to Eb4-Cb4 joint; completely excludedfrom view posteriorly.Ad5 dorsally tendinously on distal end of Cb4, ten-don continues onto distal end of Eb4; muscle fusing
160 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(larger specimen) or not anteriorly with OP, ventrallyon Cb5 distal end.SOD present.RDs adjacent.Additional remarks. SCL attached mid-dorsally toventroposteriorly extending cartilaginous tip of Bb3.TV free from Cb5s. Rosen and Patterson (1990:13)reported that Pb4 is absent in gerreids. Although it isreduced in size, we find that it and UP4 are present.IAC present. Eb4 levator process absent. Well-devel-oped bony flange (occluded from view by LP in Plate133) at end of Eb4, completely overlies cartilaginousdistal end of Eb4. Medial end of Eb4 varies in di-ameter from about equal to a little larger than that ofEb3, but is smaller than that of Eb3 in
?
(two clearedand stained specimens of Eucinostomus, USNM397354 and 306708, also examined for this charac-ter). GRAMMATIDAEGramma loreto Poey, USNM 369710, 53.3 mm;199487, 54.4 mm; 319222, 50.9 mm; 360710, 52mm. Plate 134
Description.LEI on dorsoposterior edge of Ebl lateral to un-cinate process.LE2 on dorsoposterior surface of Eb2 at aboutmid-length.LE3 on Eb3 uncinate process just ventral to car-tilaginous tip.LE4 on bony edge of Eb4 levator process just me-dial to cartilaginous tip.LP on bony edge and cartilaginous tip of Eb4 le-vator process, joining LE4 insertion posteriorly.LI1 broad; finely tendinously on anterolateralmostedge of Pb3 adjacent to medial edge of Pb2.LI2 on Pb3 dorsoposteriorly along medial edge ofattachment of TPb3-Pb4 and near medial end of Eb3.TD comprises TEb2 and TPb3-Pb4. TEb2 a pairof muscles, each separated from (and continuouswith) counterpart by CT band equal to width of areaseparating Pb3s; each element of pair comprising an-terior and posterior portions, anterior portion attachesto dorsal surface of Eb2 anterior to LE2 insertion,posterior portion attaches to posterior surface andedge of Eb2 medial to LE2 insertion; free fromTPb3-Pb4. TPb3-Pb4 on Pb3 dorsal surface medialto LI2 insertion and on anterior edge of Pb4, contin-uous posteriorly by diagonal strands of muscle withSOD.OD3-4 origin on dorsal surface of Pb3 ventral toTEb2, insertion on dorsoanterior surface of Eb3 un-cinate process and medial edge of Eb4 uncinate pro-cess.OP dorsally on Eb4 posterior surface beginning
below LP and extending medially for variable dis-tance (unclear separation from SO on one side), ven-trally on Cb5 dorsodistally posterior to Ad5.Adl-3 absent.Ad4 dorsally on Eb4 beginning below LP insertionand continuing laterally to near distal end of Eb4,ventrally on Cb4 dorsally medial to Eb4-Cb4 joint.Ad5 on dorsoposterior end of Cb5 anterior to OPand on posterior surface of Eb4-Cb4 joint.SOD present.RDs separate.Additional remarks. SCL free from Bb3 (cartilag-inous posterior end not elongate). TV4 ventrally con-tinuous across Cb5s, but dorsally attaching to Cb5s(necessary to bisect muscle ventrally to expose at-tachment). Pb4 present. UP4 present. IAC present.OPISTOGNATHIDAE
Lonchopisthus higmani Mead, USNM 186058, 73.1mm; USNM 292182, 85.1 mm.Plate 135
Additional material.
?
= Opistognathus darwiniensisMacleay. USNM 174042, 60.5 mm.
Description.LEI on anteroventral surface of Ebl uncinate pro-cess.LE2 on dorsoposterior surface of Eb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 adjacent to tip of levator process an-teriorly. ? Broadly on Eb4 dorsoposteriorly (levatorprocess absent).LP at and posterior to LE4 insertion.LI1 on tendinous covering of anterior end of Pb2dorsolaterally, covering continues medially over ad-jacent anterior end of Pb3.Remarks. Articulation of Pbl with anteromedialtip of Ebl joins tiny CT pad that covers joined an-terior ends of Pb2 and Pb3. Pb2 edentate.
?
Condi-tion of preparation prevented determining if tiny CTpad was present.LI2 on Pb3 dorsoposterolaterally, lateral to attach-ment of TPb3-Eb3-Eb4 and just medial to joint withmedial end of Eb3.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2 apair of muscles connected by tough CT across flatdorsal Pb3 surfaces, CT continuous anteriorly be-tween Pb3s with CT of pharyngeal roof; muscle ex-tending laterally on Eb2 dorsal surface anterior toLE2 insertion. TPb3-Eb3-Eb4 on Pb3 dorsoposter-iorly well medial to LI2 insertion on right side and
at medial edge of insertion on left side, with tendi-nous attachments (not illustrated) to medialmost endsof Eb3 and Eb4, continuous mid-posteriorly by cross-ing muscle strands with fine SOD. ? Thick CT padover Pb3 dorsal surfaces; pad continuous with CT
NUMBER 1
1
161joining TEb2 muscular portions, partially separablefrom more ventral CT covering and attaching directlyto Pb3s.OD3-4 origin ventral to TEb2 on posterolateraledge of Pb3 flat dorsal surface, insertion broadly onanterior surface of Eb3 uncinate process and medialedge of Eb4 uncinate process, cartilage tip of whichis extremely small in smaller specimen and absent inlarger specimen. ? On anterior surface of Eb3 un-cinate process and dorsal edge of Eb4, which is at-tached to Eb3 uncinate process (no Eb4 uncinate pro-cess).OP dorsally on posterior surface of Eb4 beginning
at levator process and continuing well medial to un-cinate process, ventrally broadly on Cb5 beginninglaterally a little medial to distal end. ? Dorsally,broadly on posterior surface of lateral half of Eb4,posteriorly overlapping and almost completely ex-cluding Ad4 from posterior view.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning at le-vator process and extending laterally to just medialto distal end, ventrally on Cb4 dorsally medial toEb4-Cb4 joint, fusing there with anterior surface ofAd5.Ad5 dorsally on posterodistal end of Cb4. ventrallyon Cb5 dorsodistally.SOD present.RDs slightly separated.Additional remarks. SCL present. TV4 two lay-ered, ventral layer free from Cb5s; dorsal layer di-vided mid-longitudinally, each portion attached to re-spective Cb5 lateral surface. IAC present ? Eachdorsal portion attaches to ventral edge of respectiveCb5. Pb4 and UP4 present.PSEUDOCHROMIDAE
Labracinus cyclophthalmus (Miiller and Troschel),USNM 290900, 2 specimens, 78.3-82.4 mm;USNM 345601, 90.7 mm SL.Plate 136
Additional material. ? = Pseudochromis porphyreusLubbock and Goldman, USNM 290595. ? =Pseudochromis persicus Murray, USNM 147902.
Description.Remarks. The muscles of both species of Pseu-dochromis are essentially the same as those of La-bracinus.LEI on Ebl posteriorly just lateral to base of un-cinate process.LE2 on raised posterior margin of Eb2.LE3 on and lateral to Eb3 uncinate process ante-
riorly.LE4 massive, on Eb4 dorsolateral surface.LP narrowly on Eb4 dorsal surface at and posterior
to LE4 insertion, completely covering minute tip oflevator process.
?
Levator process very reduced onone side, absent on other. ? Levator process like La-bracinus.LI1 tendinously on fascia covering anterior end ofPb2 (which overlaps Pb3) and continuing onto dor-solateralmost edge of Pb3.LI2 on Pb3 dorsoposterolaterally just anterior tomedial end of Eb3.TD comprises TPb2a, TEb2, and TPb3-Eb3(TPb3-Eb3-Eb4 in largest specimen). TPb2a on an-terior surfaces of Pb2s, with mid-vertical raphe con-tinuing posteriorly and joining thick CT pad coveringmost of dorsal surfaces of Pb3s, which are only partlyroofed by muscle. TEb2 a pair of muscles joined me-dially by broad area of CT and joining mid-ventralsurface of CT pad overlying Pb3 articulating facets;muscle extending laterally and attaching to Eb2 dor-soanteriorly unusually well lateral to LE2 insertion;TEb2 not continuous posteriorly with TPb3-Eb3.TPb3-Eb3 on Pb3 dorsoposteriorly ventral and lateralto articulating facet and posteromedial end of Eb3(and finely, tendinously on Eb4 dorsoanteromediallyin largest specimen); continuous posteroventrally bydiagonal muscle strand with SOD.CPb originates posteriorly from longitudinal SOlayer, extends anteriorly between Pbs, with continu-ous major branch extending anterolaterally aroundPb2s and attaching to each UP4 anterolaterally; mi-nor branch extends posteromedially from continuousmajor branch, passes medial to Pb3 and attaches toUP4 medially.OD3-4 origin on Pb3 dorsoposteriorly ventral toTEb2, insertion broadly on anterior surface of Eb3uncinate process and medial edge of Eb4 uncinateprocess.OP dorsally on most of Eb4 posterior surface, ven-trally broadly on Cb5 dorsoposteriorly.Adl anterodistally on Ebl and Cbl.Ad2 medially joins raphe with distal end of TEb2on anterior edge of Eb2 and extends laterally attach-ing on anterior surfaces of Eb2 and Cb2 at Eb2-Cb2joint.Ad3 medially on Eb3 dorsally ventral to lateralportion of OD3-4, laterally attaching to anterior sur-faces of Eb3 and Cb3 at Eb3-Cb3 joint.Ad4 broadly on Eb4 ventrally almost entirely an-terior to OP, ventrally broadly on Cb4 medial to Eb4-Cb4 joint.Ad5 dorsally on Eb4-Cb4 joint posteriorly, con-tinuing, ventrally on Cb5 broadly anterodistally an-terior to OP.SOD slender.RDs adjacent.Additional remarks. SCL present. TV4 dorsal sec-tion attaches to Cb5s anteriorly, ventral section con-tinuous across anteroventral ends of Cb5s. Pb4 and
162 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONUP4 present. Pb2 toothed. Eb4 levator process withminute cartilage tip.Cb5 bears a tiny bit of cartilage at or near its distalend in Labracinus, but is usually completely ossifiedin Pseudochromis, although there is a "spot" of car-tilage on one Cb5 laterally well anterior to its distalend in ?. Labracinus has a tiny IAC, which is absentin ? and ?. Labracinus has a tiny, cartilaginous Pbl.
? Pbl has relatively long cartilaginous dorsal andventral ends separated by small central ossified area.? Pbl state unknown.LEIOGNATHIDAE(See additional remarks following description fordiscussion of valid family name for this taxon.)Gazza sp.. USNM 268914, 2 specimens, 68.6-69.2mm. Plate 137
Additional material. ? = Leiognathus equula Forss-kal, USNM 349512, 58.6 mm. ? = Secutor insi-diator (Bloch), USNM 329585, 70.9 mm.
Description.LEI broadly on Ebl dorsolateral^ and on anteriorand medial surfaces of large, dorsally expanded tri-angular process at lateral end of Ebl (lateral surfaceof process concave, receives gill filament bases; pro-cess apparently not an uncinate process as similarprocess is serially present on Eb2 and gill filamentsare not known to attach to Ebl uncinate process; car-tilaginously tipped uncinate process absent); musclewith lateral tendinous portion extending to origin. ?Like Gazza, but lateral edge of Ebl not expandednearly as much. ? Narrowly, tendinously on Eblposterolaterally medial to scarcely expanded lateralend of Eb 1
.
LE2 on medial edge of large, dorsally expandedtriangular process at lateral end of Eb2; ventrolater-ally joining raphe with posterolateral edge of TEb2;gill filaments attach to concave lateral edge of tri-angular process.
?
? Lateral end of Eb2 not ex-panded dorsally nearly as much as in Gazza.LE3 musculously and, variably, tendinously onbony portion of Eb3 uncinate process anteriorly (pro-cess has minute cartilage tip posteromedially. visibleonly after separation from Eb4 uncinate process).LE4 + LP fuse, well dorsal to Eb4, along tendi-nous raphe and form arms of a Y; raphe continuesalong anterior edge of musculous shank of Y, whichis presumably formed by lateral section of OP, andattaches between uncinate process and lateral end ofEb4 posteriorly; another tendinous raphe extendsalong lateral section of OP posteriorly and attachesto hook-like cartilaginous distal end of Cb4; lattertendinous raphe on Cb4 expands into CT sheet thatcontinues dorsoposteriorly as PP (not illustrated);
dorsoposteriorly, ventromedial portion of medial OPsection attaches to Cb5 posterolaterally. ? Tendinouscontinuation from fused LE4 + LP continues alonganterior edge of lateral OP section and attaches toposterodistalmost bony edge of Eb4, OP continuesventrally and meets Ad5 on Cb5; tendinous rapheabsent along lateral section of OP posteriorly. ? LE4+ LP fuse along strong tendon that divides beforeattaching between uncinate process and distal end ofEb4, tendon divides ventral to attachment, with lat-eral section of OP attaching to it.LI1 on Pb2 anterolaterally well ventral to cartilage-tipped dorsally extending process of Pb2.LI2 tendinously on Pb3 just medial to joint withposteromedial end of Eb3.TD comprises TEb2 and TEb3-Eb4. TEb2 com-prises a muscle on each side joined medially by ten-dinous CT to ventrolateral edge of a thick, cone-shaped CT pad (attenuated end ventral) that attachesmedially to a conforming dorsal shelf of Pb3 and sitsfreely in a conforming depression of dorsoanteriorsurface of OD3-3'; pads on either side joined by CTdorsally across dorsal edge of conforming Pb3 shelf;TEb2 attaches on Eb2 dorsally, joining LE2 ven-troanteriorly and extending well lateral to it; musclefree from TEb3-Eb4. TEb3-Eb4 on Eb3 posterioredge medial to base of uncinate process, continuingposteriorly and passing dorsal to medial end of Eb4but attaching ventrally to posteromedial edge of Eb4and, on left side only of one specimen, joining dor-somedial edge of OP medial section; muscle ven-troanteriorly continuous by diagonal muscle strand(obscured from view in illustration) with SOD. ? ?TD comprises TEb2 and TEb3; TEb3 attaches on me-dial edge of Eb3 uncinate process ventral to OD3insertion.OD3, OD3' massive, joint origin on lateral surfaceof conforming Pb3 dorsal shelf (see description ofTEb2), dividing laterally with small OD3 section in-serting on Eb3 uncinate process dorsoanteriorly, andlarge OD3' section inserting separately on remainderof Eb3 uncinate process anteriorly immediately ven-tral to OD3, insertion continues ventrally covering allof bony dorsolateral surface of Eb3 (superficially ap-pears as if OD3 and OD3' are entirely fused; sepa-ration most distinct in ?).OD4 absent.OP presumably in two sections, for lateral sectionsee LE4 above; medial section dorsally, broadly onEb4 posterolaterally, ventrally more broadly on Cb5posteriorly, joining posterior raphe of OP lateral sec-tion with Ad5 posteriorly; lateralmost portion of me-dial OP section joining tendinous raphe with lateralOP section on posterodistal end of Cb4 where rapheexpands into CT sheet which continues to PP (notillustrated). ? Posterior raphe of lateral OP section
NUMBER 11 163
absent; two OP sections join on Cb5; CT of PP at-taches to distal end of Cb5.M. SO-Pb2 strap of SO longitudinal fibers extend-ing anteriorly and inserting on Pb2 posteroventrally,passes posteriorly along Pb3 medially and then ven-tral to TEb3-Eb4 and. on left side only, divides, withone portion inserting on Eb4 at and medial to OPmedial section and the other becoming continuouswith SO and SOD anteriorly; on right side, Eb4 in-sertion is absent.Remarks. This muscle does not insert on Eb4 inLeiognathus (only one side is adequate for determi-nation) or on one side in Secutor, but does insert onEb4 on the other side.Ad 1-3 absent.Ad4 relatively small, scarcely if at all visible ex-ternally, on Eb4 ventrally and Cb4 dorsally medialto Eb4-Cb4 joint.Ad5 small, on dorsodistal end of Cb5 and cartilag-inous finger-like process at distal end of Cb4, joiningshort raphe there with ventromedial end of OP lateralsection and ventrolateral end of OP medial section.SOD slender.RDs well separate anteriorly, swelling posteriorlyand becoming proximate, each consisting of strap-like lateral section, RD', and much larger medial sec-tion, RD. RD' anteroventrally continuous (infiltrat-ed?) with CT of SO and inserting on UP4 and Pb4posteriorly; RD with minor insertion on posterome-dial edge of UP4 and major insertion on Pb3 poste-
riorly.Additional remarks. SCL weakly attached mid-an-teriorly to posteroventrally extending cartilaginousend of Bb3 (SCL easily separated from Bb3). TV4mostly continuous ventrally, but dorsoanterior fibersattach to anterolateral surfaces of Cb5s (hence similarto condition in to many other fishes that have an LE4-OP sling; e.g., labrids, cichlids, centrogenyids). Pb4and UP4 present. IAC absent. Eb4 with small, bonyflange dorsally at distal end. Dorsal end of Pbl bony;
. ? Dorsal end cartilaginous. Ebl uncinate processabsent. Eb3 uncinate process with minute cartilagi-nous tip.
?
Cartilage tip of uncinate process normal.? Tip of process bony on both sides. Eb4 uncinateprocess present.
?
Uncinate process on one side withminute cartilage tip, bony tip on other side. Eb4 le-vator process absent in all.Correct family name. Bleeker (1859:xxiii) coinedthe family name Equuloidei, in which he includedfishes currently recognized in the families Leiognath-idae and Menidae. Equuloidei was emended toEquulidae by Gill (1893:134), which Gill listed as asynonym of "Liognathidae Gill, 1892." We were un-able to find an 1892 or earlier publication of Gill'sthat refers to Liognathidae.Equuloidei Bleeker was based on the genus EquiilaCuvier, which is a junior synonym of Leiognathus
Lacepede; however, Equulidae has date seniority overLiognathidae Gill (first emended as Leiognathidae byJordan and Evermann, 1902:338). Hence, Equulidaehas seniority and, ostensibly, should replace Leiog-nathidae, but as far as we can tell has not been usedas a senior synonym since before 1899. Leiognathi-dae has been used as a valid senior synonym for thefamily for many years and we elect to continue touse it and recommend that Equulidae be suppressed.
POLYCENTRIDAE
Remarks. Until recently (Britz. 1997), Polycentrus,Polycentropsis, Monocirrhus and Afronandus wereincluded with Nandus in the family Nandidae. Britz(1997) demonstrated that a Nandidae that includedthese four genera was polyphyletic and removedthem. He hypothesized the monophyly of the firstthree genera based on egg morphology, but only
"tentatively" included Afronandus with them basedon a single shared character: presence of adhesivefilaments on the vegetal egg pole. Britz noted thatthis character also had been reported for some Pseu-dochromidae and Cichlidae, which he indicated werenot closely related to the Nandidae, but he did notassign either the first three or all four genera to aseparate family. Berra (2001:427), based on Britz(1997), apparently was first to define the Polycentri-dae as comprising only these four genera. Kullanderand Britz (2002:301) similarly recognized a Polycen-tridae, but again only "tentatively" included Afron-andus, based on Britz's (1997) character, which theyerroneously described as "presence of attachment fil-aments around the micropylar area of the egg."We find at least three synapomorphies, in additionto the vegetal-pole filaments, that support monophylyof the Polycentridae: loss of LE3, loss of sensorycanals on most lateral-line scales (including all ofthose on posterior half of body), and presence ofslender ligament attaching dorsal margin of Ebl an-terior to LEI to ventral surface of skull at or nearorigin of LEI.Notwithstanding its unilateral vestigial conditionin Monocirrhus and Afronandus, loss of LE3 is arelatively uncommon character state for percomorphs(various scorpaenoids, Psettodidae, many smegma-morphs, Bramidae, Uranoscopidae, dactyolscopidblennioids, callionymids, gobiesocids). Absence oflateral-line scale tubes, or restriction of the tubes toa limited area on the anteriormost portion of thebody, is also uncommon in perciform fishes, partic-ularly those inhabiting freshwaters, although Britz(pers. comm.) informs us that the condition is alsotrue of some deeply nested taxa within the Anaban-toidei. Among freshwater percomorphs, the reductionor absence of tubed lateral-line scales is generallyrestricted to mugilids, various atherinomorphs and
164 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
gasterosteomorphs, elassomatids, percichthyids, nan-nopercids, and specialized members of other groups(e.g., gobiids). The distribution of the Ebl ligamentin perciforms is less well known, as we may haveoverlooked or removed it in some taxa early in ourstudy (see Introduction, methods section); however,it is not present in anabantoids, ambassids, and Tox-
otes. We have noted the ligament at least in pem-pherids, cirrhitids, bathyclupeids, lacteriids, acro-pomatids.Another character, presence of a hook-like post-premaxillary process (reduced in Afronandus), is alsouncommon among acanthomorphs, and may repre-sent another polycentrid synapomorphy (see Liem,1970:figs. 13?16). A similar post-premaxillary pro-cess is also present in Nandus (Liem, 1970:fig. 12),but is reduced in the closely related Badis, possiblya reflection of the small size attained by this genus.Liem (1970), who examined the osteology andgill-arch musculature of all four polycentrid genera,included those genera in the Nandidae, along withNandus, which he also examined (Liem, 1970:9, 11).Liem (1970:56) reported that all five genera had onlythree external levators, but did not indicate whichlevator was missing. He suggested that the posteri-ormost levator might represent a fusion of the thirdand fourth. Liem was correct in asserting that Poly-centrus, Polycentropsis, Monocirrhus and Afronan-dus have only three LEs (he could easily have misseda vestigial LE3, if present, which we found unilat-erally in Afronandus and Polycentropsis). but he wasincorrect in implying the presence of only three LEsin Nandus, in which LE3 and LE4 are both well de-veloped. Commonly in acanthomorphs, includingNandus, LE3 and LE4 appear to be fused along theirentire length, except for their easily overlooked sep-arate insertions on the surfaces of Eb3 and Eb4. LE3and LE4 are easily separated and there is no inter-mingling of their fibers.Other specializations shared by the polycentridsare the absence of SOD and Pb4 and the presence ofmore than three anal-fin spines and only one epural.All these characters have a broad and varied distri-bution.In summary, the specializations shared by the fourpolycentrid genera, strongly imply their monophyly.
Polycentropsis abbreviata Boulenger, USNM302514, 34.2 mm.Not illustratedAdditional material. ? = Polycentrus schomburgkiiMiiller and Troschel, USNM 226071, 43.0 mm.
Description.Remarks. The arrangements of the muscles, exceptfor TD, are very similar in the two taxa.
LEI on Ebl posteriorly noticeably lateral to car-tilage tip of uncinate process. Slender ligament orig-inates on skull with LEI and inserts on Ebl anteriormargin anterior to LEI insertion.LE2 on Eb2 about mid-dorsoposteriorly.LE3 present only on left side; thread-like, less than10% width of LE4, inserting on tip of Eb3 uncinateprocess. ? Absent on both sides.LE4 broadly on Eb4 dorsoposteriorly beginningventral to OD3-4 insertion on posterior surface ofEb4 just ventral to tip of uncinate process, meetingLP insertion posteriorly; posteroventral edge of in-sertion meets OP dorsolateral^ and Ad4 dorsoanter-iorly; Eb4 levator process absent. CT sheet extendsfrom near LE4 and LP insertions to distal end of Eb4and becomes PP dorsally. ? Like Polycentropsis butbeginning lateral to OD3-4 insertion on posteriorsurface of uncinate process and extending laterally totip of levator process, meeting LP insertion posteri-orly; CT sheet begins at levator process.LP on Eb4 at and anterior to LE4 insertion.LI1 on lateral edge of dorsoanteriormost tip ofPb2, almost appearing to continue onto adjacent me-dialmost edge of IAC, and on dorsoanteromedialmostedge of adjacent Pb3. ? Like Polycentropsis, but in-sertion more extensive on Pb2, and, on one side, afew lateralmost muscle fibers continuing onto medi-almost surface of IAC.LI2 on Pb3 dorsoposteriorly immediately medialto medial end of Eb3.TD comprises TEb2 and TPb3-Eb3, and TEb4.TEb2 musculously continuous only narrowly poste-
riorly, with mid-anterior notch continuing posteriorlyas mid-longitudinal raphe, which gives rise dorsallyto CT sheets covering muscles and is continuous an-teriorly with CT of pharyngeal roof; ventroanteriorly,muscle attaches loosely to Pb3; laterally muscle at-taches on Eb2 dorsally anterior to LE2 insertion; pos-teriorly muscle is free from TPb3-Eb3. TPb3-Eb3 rel-atively narrow, begins anteriorly on Pb3 just anteriorto medial end of Eb3 and continues posteriorly onposteromedialmost edge of Eb3; muscle is free fromTEb4. TEb4 broader than TPb3-Eb3, attaches alongEb4 posteriorly between medial end and uncinateprocess.
? TD comprises TPb2, TEb2, TPb3-Eb3, andTEb4. TPb2 a semicircular ribbon of muscle on eachside, arising ventroanteromedially from TEb2 dorsal-ly and finely attached by CT to dorsoanteriormostsurface of Pb3 (not attached to Pb2); muscle curvesposterolaterally and joins medial band of CT cover-ing Pb3s posteriorly and joining the bi-lateral musclestraps of TEb2. CT joining muscle sections of TEb2notched deeply mid-anteriorly; anteriorly joining CTof pharyngeal roof and dorsally giving rise to CTsheets; muscle extends laterally from CT coveringPb3s and attaches on Eb2 dorsally well anterolateral
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to LE2 insertion; free from TPb3-Eb3. TPb3-Eb3ventral to OD3-4, beginning anteriorly a little ante-rior to LI2 insertion, continuing posteriorly along me-dial edge of LI2 insertion, and attaching finely toposteromedialmost corner of Eb3; posteriorly contin-uous by slender, diagonal muscle strand with TEb4.TEb4 slender, on Eb4 a little lateral to cartilaginousmedial end, meeting OP dorsomedially.OD3-4 on Pb3 dorsomedially ventral to TEb2, in-sertion on Eb3 anteriorly ventral to tip of uncinateprocess and on Eb4 posteriorly ventral to tip of un-cinate process.OP strap-like; dorsally on Eb4 posteriorly, begin-ning medially ventral to TEb4 attachment, continuinglaterally and meeting LE4 insertion posteriorly, andposteriorly overlapping Ad4 dorsomedially; ventrallyon Cb5 dorsoposteriorly, anteriorly meeting TV5posterolaterally, laterally posteriorly overlapping Ad5ventromedially. ? Dorsally on Eb4 posteroventrallybeginning medially ventral to OD3-4, continuing lat-erally almost to levator process (lateral to posteroven-tral edge of LE4 insertion), posteriorly overlappingAd4 dorsomedially; ventrally like Polycentropsis, butjoining well-developed raphe with Ad5 posterome-dially.Ad 1-3 absent.Ad4 dorsally on Eb4 beginning ventrally anteriorto lateral edge of OP and extending laterally almostto Eb4-Cb4 joint; ventrally on Cb4 dorsally just me-dial to Eb4-Cb4 joint.Ad5 strap-like, dorsoanteriorly on Cb4 beginningjust medial to distal cartilage tip, posteroventrally onCb5 beginning just medial to distal cartilage tip. ?Dorsoanteriorly meets Ad4 ventroposterolaterally onCb4; begins posteroventrally on distal tip of Cb5 andjoins raphe with OP ventrolaterally.SOD absent.RDs separated by space about one-half diameter ofone RD.
?
Space less than one-half diameter of oneRD.Additional remarks. SCL attached mid-dorsally toventral surface of cartilaginous posterior tip of Bb3.TV4 free from Cb5s. Pb2 toothed Pb4 absent; UP4present.
Afronandus sheljuzhkoi (Meinken), USNM 372183,65.9 mm. Plate 138
Description.LEI on Ebl dorsoposteriorly, well lateral to car-tilaginous tip of uncinate process; slender ligamentoriginates with LEI and inserts on Ebl anteriorly an-terior to LEI insertion.LE2 on Eb2 about mid-dorsoposteriorly, posteriorto posterolateralmost edge of TEb2.
LE3 absent on right side; on left side, a very finefilament inserting on tip of Eb3 uncinate process.LE4 on Eb4 posteriorly medial to lateral end, an-teromedially meeting OD3-4 and laterally meetingLP insertion medially.LP on Eb4 dorsally beginning at lateral edge ofLE4 insertion and extending laterally; CT sheet ex-tends from lateral edge of insertion and becomes PPdorsoposteriorly.LI1 on Pb2 dorsally just posterior to dorsoanter-iormost cartilage tip, inertion becoming tendinous asit joins adjacent anterior end of Pb3; muscle fibersabut medial IAC surface, but no fibers basally inserton IAC.LI2 narrowly on Pb3 dorsolaterally, medial edgemeeting TPb3-Eb3-Eb4 lateral edge.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2broad medial to much narrower extension onto Eb2dorsally, there attaching anterior to LE2 insertion;broadly, mid-anteriorly consisting of thick CT pad,which attaches anterolaterally to dorsoanteriormostends of Pb2s and is mid-anteriorly continuous withCT of pharyngeal roof; muscle fibers continuous nar-rowly and only posteriorly, but interrupted there byvery short, fine, mid-longitudinal raphe; muscle notcontinuous posteriorly with TPb3-Eb3-Eb4. TPb3-Eb3-Eb4 anteriorly ventral to OD3-4s, attaching toPb3 at medial edge of LI2 insertion, continuing pos-teriorly and attaching to posteromedialmost corner ofEb3, abruptly narrowing at that point and then ex-panding considerably and attaching to Eb4 postero-medially, there just reaching dorsomedialmost edgeof OP.OD3-4 origin on Pb3 dorsomedially ventral toTEb2, insertion on Eb3 anteriorly immediately ven-tral to tip of uncinate process and on Eb4 posteriorlyon and ventral to tip of uncinate process; meets LE4insertion ventroanteriorly.OP dorsally on Eb4 posteriorly beginning mediallynear lateral end of TPb3-Eb3-Eb4 and extending lat-erally to below LE4, overlapping medial half of Ad4posteriorly; ventrally on Cb5 dorsally beginning nearlateral end and extending medially and meeting pos-terolateral edge of TV5; anteriorly overlapping Ad5posteromedially.Adl-3 absent. GFMsl-3 (not illustrated) moder-ately well developed.Ad4 dorsally on Eb4 posteroventrally beginningmedially anterior to OP and extending laterally al-most to distal end of bony surface; ventrally on Cb4dorsally beginning medially anterior to OP (which ison Cb5) and extending laterally along Ad5 attach-ment dorsally to near lateral end of bony surface ofCb4.Ad5 dorsally on Cb4 posteriorly beginning wellmedially and extending laterally along ventral edgeof Ad4 almost to lateral end of bony surface of Cb4;
166 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ventrally on Cb5 posteriorly beginning near distalend of bony surface and extending medially; over-lapped posteriorly by OP.SOD absent.RDs almost adjacent.Additional remarks. SCL present. TV4 free fromCb5s. IAC articulates with cartilaginous medial endof Ebl uncinate process. Pb4 absent. UP4 present.Eb4 levator process absent on one side, greatly re-duced on other (treated as absent).
Monocirrhus polyacanthus Heckel, USNM 103840,78.1 mm: USNM 269969, cleared and stained,56.1 mm. Not illustrated
Description.LEI on Ebl well lateral to tip of uncinate process;strong, ribbon-like ligament originates with LEI andinserts on Ebl anteriorly anterior to LEI.LE2 on raised process on Eb2 posteriorly at aboutmid-length of element.LE3 absent.LE4 broadly on Eb4 dorsoposteriorly beginninglateral to uncinate process and extending laterallymuch of distance to distal end, laterally meeting slen-der LP insertion posteriorly.LP slender, at and anterior to LE4 insertion later-
ally.LI1 on Pb2 dorsally just posterior to anteriormosttip, insertion continuing on adjacent anterolateral-most margin of Pb3 and impinging on medialmostsurface of IAC, but no fibers basally inserting on it.LI2 on Pb3 laterally medial to medial end of Eb3.TD comprises TEb2 and TPb3-Eb3-Eb4. TEb2musculously continuous from attachment on Eb2dorsoanteriprly anterior to LE2 insertion on one sideacross Pbs to attachment to Eb2 on other side; areaover Pbs attached mid-ventroanteriorly to pharyngealroof CT and dorsally to CT sheets, with strongestattachment on each side to muscle surface just medialto extension of muscle onto Eb2; strong CT continuesanteriorly, attaching to ventroanterior edge of LI1,and tightly enveloping anterior ends of Pb2 and Pb3and medial surface of Pbl-Ebl joint; muscle not con-tinuous with TPb3-Eb3-Eb4. TPb3-Eb3-Eb4 originventral to OD3-4, beginning on Pb3 along medialedge of LI2 insertion, continuing onto posteromedi-almost corner of Eb3, followed posteriorly by con-siderable lateral extension, which attaches on Eb4mid-posteriorly.OD3-4, OD3' anteriorly on Pb3 dorsomediallyventral to TEb2; OD3?4 posteriorly on bony surfacesof Eb3 uncinate process anteriorly and Eb4 uncinateprocess posteriorly, almost covering tip of latter pro-cess; OD3' branches off OD3-4 ventroanteriorly at
about mid-length and attaches to Eb3 dorsally ventralto OD3-4 on Eb3.OP dorsally on Eb4 posteriorly beginning medially
at about mid-length of Eb4 and extending laterally tobelow mid-point of LE4 insertion, there joining Ad4dorsomedialmost edge; ventrally on Cb5 dorsally be-ginning at posterolateral edge of TV5 and extendinglaterally a short distance and joining Ad5 postero-medially; not clearly differentiated from SO medial-iy-Ad 1-3 absent.Ad4 on Eb4 dorsally beginning below about mid-point of LE4 insertion, where Ad4 and OP join, andextending laterally almost to lateral end of bony sur-face of Eb4; ventrally on Cb4 dorsally beginning me-dially at point ventral to its medial attachment on Eb4and extending laterally almost to posterodistalmostend of bony surface of Cb4, meeting Ad5 dorsoan-terolaterally.Ad5 strap-like, dorsoanteriorly on Cb4 dorsopos-teriorly well medial to lateral end of Cb4, there meet-ing Ad4 and extending medially a short distance;posteroventrally on Cb5 dorsally beginning near dis-tal end and extending a short distance medially, meet-ing OP ventroanteromedially.Remarks. It is unusual in a perciform for Ad5 toattach well medial to the distal end of Cb4.SOD absent.RDs separated by distance about diameter of oneRD.Additional remarks. SCL attached mid-dorsally totip of posteroventrally extending cartilaginous end ofBb3. TV4 free from Cb5s. Pb4 absent; UP4 present.IAC present. Eb4 levator process absent. Medial endof Eb4 larger than that of Eb3 in 78.1 mm specimen,about equal in 56.1 mm specimen.SPHYRAENIDAERemarks. Johnson (1986) included Sphyraenidaein the Scombroidei, but Orrell et al. (2003 and inpreparation) excluded it based on a molecular study.
Sphyraena barracuda (Edwards), USNM 331677,139 mm. Plate 139
Description.LEI on anterolateral surface of Ebl uncinate pro-cess.LE2 on raised bony posterior edge of Eb2.LE3 long tendinous origin, insertion on tip of Eb3uncinate process.LE4 long tendinous origin, insertion on dorsal sur-face of Eb4 just lateral to base of uncinate process.LP medial half of insertion at and anterior to in-sertion of LE4, posterior half is lateral to LE4.
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LI1 larger than LI2. broadly on Pb2 dorsoanteriorsurface beginning at and ventral to TPb2.LI2 on posterolateral edge of Pb3 anterior to me-dial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2dorsal to TEb2. V-shaped, arms anterior, joined pos-teriorly along mid-longitudinal raphe, which attachesventrally to CT of pharyngeal roof, each arm attachesto respective Pb2 dorsoanteriormost surface; musclefuses ventrally with TEb2 on either side of raphe.TEb2 broad, short laterally, attaching to dorsal sur-face of Eb2 well medial to LE2 insertion, posteriorlycontinuous by diagonal strap of muscle with TPb3-Eb3. TEb3-Pb3 on Pb3 dorsoposteriorly medial toLI2 insertion and on posteromedial edge of Eb3. freefrom SOD.OD3-4 originating on Pb3 anteromedially ventralto TEb2 and inserting on medial edges of Eb3 andEb4 uncinate processes.OP relatively slender, dorsally on posterior surfaceof Eb4 at and medial to uncinate process, ventrallyon posterodistal end of Cb5, joining raphe with Ad5ventrolaterally.Ad 1?3 absent.Ad4 relatively broad, on Eb4 posteriorly beginningventral to LE4 insertion and continuing laterally todistal end of bony portion of Eb4. broadly ventrallyon Cb4 dorsal surface medial to Eb4-Cb4 joint andmedial to broad Ad5 attachment to Cb4.Ad5 dorsally broadly on posterolateral bony sur-face of Cb4 just medial to distal end, fusing mediallywith SO: ventrally broadly on Cb5 laterally, fusinganteriorly with TV5.SOD present.RDs separated by space greater than width of oneRD. RD extends anteriorly between two longitudinalSO muscle straps: one originating from SO dorsallyin region ventral to SOD and extending anteriorlymedial to RD and inserting on Pb3 dorsally at andmedial to OD3-4 origin, and the other originating inarea anterior to OP and extending anteriorly lateralto RD, becoming dorsal to RD anteriorly and insert-ing on most of medial edge of Pb3 ventral to OD3^kAdditional remarks. SCL absent (note Bb3 hasposteroventrally extending cartilaginous tip which at-taches to the ventral aorta where this vessel dividesbilaterally and extends anteriorly). TV4 free fromCb5s. Pb4 absent, UP4 present. Pb2 toothed. IACpresent. Eb4 levator process absent. Medial end ofEb4 larger than that of Eb3. PCI originates by longtendon on cleithrum and attaches on Cb5 beginningwell medial to distal end of Cb5 and continuing me-dially. KURTIDAEKurtus sp., USNM 345060, 92.4 mm.Not illustrated
Description.LEI on dorsolateral edge of Ebl uncinate processbeginning just lateral to cartilage tip.LE2 on dorsal edge of expanded posterior marginof Eb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 dorsoposterolateral to tip of uncinateprocess, ventroposteriorly meeting LP insertion ven-troanteriorly.LP on Eb4 beginning at and extending posterior toLE4 insertion.LI1 inserting on Pb2 dorsoanteriormost processposteriorly, continuing posteroventrally and attachingto anteriormost edge of Pb3; joining posterior edgeof TPb2 attachment to Pb2; smaller than LI2.LI2 on Pb3 posterolaterally immediately adjacentto medial end of Eb3.TD comprises TPb2, TEb2, and TEb3. TPb2roughly V-shaped, arms broad, flat, open anteriorly,completely dorsal to TEb2, with raphe coursingthrough posterior apex and across TEb2 (thus com-pletely dividing both muscles) and attaching ventrallyto dorsomedial edge of each Pb3; muscle attachedmid-anterioiiy to CT of pharyngeal roof, attached an-terolaterally to dorsoanterior tip of Pb2, there joiningLI1 just dorsal to its insertion. TEb2 extending lat-erally and attaching to Eb2 dorsally anterior to LE2insertion; free posteriorly from TEb3 except for fineposterior CT continuations from dorsomedial edgesof Pb3s to mid-longitudinal raphe of TEb3. TEb3posteroventral to level of TEb2, attaching broadly toposteromedial margin of Eb3; raphe continuing pos-teriorly across SOD.OD3-4 origin on Pb3 dorsomedially ventral toTEb2; inserting on Eb3 broadly anteriorly ventral totip of uncinate process and on Eb4 somewhat lessbroadly ventral to tip of uncinate process.OP a relatively narrow strap, dorsally on Eb4 pos-teriorly beginning medial to uncinate process and ex-tending medially, ventrally on Cb5 dorsally posteriorto Ad5, beginning laterally near distal end of Cb5and extending medially.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 posteriorly beginning
at lateral edge of OP and extending laterally to endof bone, ventrally on Cb4 dorsoposteriorly beginninga little medial to lateral end and anterior to Ad5 at-tachment and extending medially.Ad5 dorsally on Cb4 posteriorly beginning neardistal end and extending medially, ventrally on Cb5dorsoposteriorly and extending medially anterior toOP.SOD very fine, medially with pair of fine ventralextensions from mid-dorsal raphe, each encirclingone RD.RDs separated by distance less than half diameterof one RD.
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Additional remarks. SCL interrupted medially byattachment to autogenous ball of cartilage attached toventral tip of posteroventrally extending cartilaginousposterior end of Bb3 (autogenous ball also present incleared and stained specimen of Kurtus indicus,USNM 305690). TV4 free from Cb5s. Pb4 absent.UP4 present. IAC present. Eb4 levator process ab-sent. Pbl bony with cartilage tips and, ventrolaterally,uniquely, has toothplate with fine teeth adjoiningsimilar toothplate on medial end of Ebl (consistentwith scale-eating behavior). Pb2 toothed. Medial endof Eb4 smaller than that of Eb3.
The next four families were included with nineothers in a suborder Trachinoidei by Nelson (1994:395 et seq.), who indicated that the composition ofthe suborder was probably polyphyletic.AMMODYTIDAEAmmodytes dubius Reinhardt, USNM 302478, 180mm. Plate 140Description.LEI on broad tip of Ebl uncinate process antero-laterally.LE2 on tip of raised mid-posterodorsal edge ofEb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 together with LP (laterally) tendinously ondorsal bony edge of Eb4 lateral to uncinate process.LP together with LE4 (medially) tendinously onbony distal dorsal edge of Eb4 at and lateral to LE4insertion.LI1 on bony dorsolateral edge of Pb2 just ventralto articulation with IAC.LI2 on Pb3 dorsoposteriorly at and lateral to me-dial edge of Pb3 portion of TPb3-Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2small, thin, V-shaped, dorsal to TEb2 anteriorly, eacharm attaching to cartilaginous tip of Pb2 anterior end,junction of arms divided by longitudinal raphe, whichcontinues across TEb2, attached ventrally to CT ofpharyngeal roof. TEb2 transversely broad, longitu-dinally narrow, attaching mid-ventrally to CT of pha-ryngeal roof, attaching laterally to Eb2 dorsally an-terior to LE2 insertion; free and well separated fromTPb3-Eb3. TPb3-Eb3 on Pb3 dorsoposteriorly at andmedial to LI2 insertion, continuing posteriorly andattaching finely to medial tip of Eb3, continuous mid-posteriorly by crossing strands of muscle with SOD.OD3-4 origin broadly on Pb3 dorsoposterolateraledge, anteriorly ventral to TEb2, but mostly posteriorto TEb2; insertion on Eb3 anteriorly ventral to tip ofuncinate process and on medial edge of Eb4 uncinateprocess.OP dorsally posteriorly on Eb4 beginning on un-
cinate process and extending laterally almost to be-ginning of cartilaginous distal end, ventrally on Cb5dorsolaterally, joining CT ventrolaterally with pos-terior end of Ad5, medially continuous with SO.Remarks. This description was completed muchearlier than that of Symphysanodon berryi (Symphy-sanodontidae, q.v.), in which the dorsal gill-arch mus-culature is otherwise remarkably similar. Re-exami-nation of our A. dubius specimen indicates the possi-bility that OP is divided into two parts, similar to S.berryi, but this should be verified in another specimen.Ad 1-3 absent.Ad4 broadly on Eb4 dorsoposteriorly lateral to OP,ventrally on Cb4 beginning at inner angle of Eb4-Cb4 joint and continuing a short distance medially.Ad5 posteriorly on distal end of Cb5 and anteriorlyon Cb4 posterolaterally and AC4 posteroventrally,joining tendinous raphe with OP ventromedially.SOD present, slender.RDs separated by distance greater than diameter ofone RD.Additional remarks. SCL attached mid-dorsally toventroposteriorly extending cartilaginous posteriorend of Bb3. TV4 free from Cb5s. Pb4 and UP4 pres-
ent. Pb2 toothed. IAC present. AC4 present (not yetbudded off in 1 1 1 mm, cleared and stained specimen,USNM 302247). Medial end of Eb3 larger than thatof Eb4.Kayser (1962) described and illustrated the skele-ton and musculature of A. tobianus Linnaeus. Thedorsal gill-arch muscles of A. tobianus differ mostnotably from those of A. dubius in that TPb2 is ab-sent, the origin of OD3-4 is completely ventral toTEb2, LI1 and LI2 are much more massive, and ACis apparently absent or has not budded off (thus Ad5attaches anteriorly only to Cb4). Kayser illustratedthe GFMs, which we did not.
TRACHINIDAE
Trachinus draco Linnaeus, USNM 349536, 2 speci-mens, 80.6-84.3 mm.Plate 141Description.LEI on Ebl anteriorly lateral to tip of uncinateprocess.LE2 on raised dorsoposterior bony edge of Eb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 dorsoposteriorly between uncinateand levator processes.LP on Eb4 posterior to LE4.LI1 on most of Pb2 bony surface dorsomediallywith muscle fibers separating dorsoposteriorly and in-serting on Pb3 bony surface slightly posteromedial toanterior end of Pb3.LI2 on Pb3 dorsally medial to medial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2
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flat, broad, bi-lateral pair of anterolaterally extendingmuscles originating posteriorly from mid-longitudinalraphe or somewhat heart-shaped ribbon of muscle di-vided mid-longitudinally; either type dorsally on mid-section of TEb2 and attaching anterolaterally to car-tilaginous dorsoanteriormost tip of Pb2; anterior fibersmay coalesce medially with those of TEb2; CT sheetsattaching to skull arise from mid-longitudinal raphe.TEb2 broad, extending laterally onto Eb2 dorsally an-terior to LE2 insertion, not continuous posteriorly withTPb3-Eb3. TPb3-Eb3 on Pb3 dorsolateral^ beginninganteriorly medial to LI2 insertion and continuing pos-teriorly and attaching to posteromedialmost edge ofEb3 (see Remarks following), continuous posteriorlyby diagonal muscle strand with SOD.Remarks. On right side only of larger specimen(illustrated), TPb3-Eb3 continues onto Eb4 dorso-medially. We consider this continuation to be anom-alous.OD3-4 anteriorly on Pb3 dorsomedially ventral toTEb2, posteriorly on anterior surface of Eb3 uncinateprocess and medial edge of Eb4 uncinate process.OP dorsally on Eb4 posteriorly, extending fromuncinate process about half distance to medial end ofEb4, ventrally on Cb5 dorsoposterolaterally.Ad 1-3 absent.Ad4 broadly dorsally on ventral surface of Eb4lateral to levator process, ventrally equally broadlyon Cb4 dorsally medial to Eb4-Cb4 joint.Ad5 dorsally on Cb4 posterolaterally (reachingdistal end), joining raphe with Ad4 ventral attach-ment; ventrally anterior to OP on Cb5 dorsally,reaching distal end.SOD present.RDs separated by space slightly less than diameterof one RD.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. IAC present. Eb4 levatorprocess present.URANOSCOPIDAE
Kathetostoma cubana Barbour, USNM 187953, 81mm.
Additional material. ? Xenocephalus egregius (Jor-dan and Thompson), USNM 186212, 89.1 mm.Plate 142
Description.LEI on anterior surface of broad bony flange lat-eral to tip of Ebl anterior process (uncinate processand Pbl absent). ? Uncinate process and tiny Pblpresent.LE2 short, bulky, on dorsoanterior surface of Eb2mid-laterally, medial edge of insertion joining raphewith TEb2 lateralmost edge.LE3 absent.
LE4 long, bulky, on Eb4 dorsoposterolaterally.LP very fine, at lateralmost edge of LE4 insertion;easily overlooked and lost during dissection.LI1 on dorsoanterolateralmost surface of Pb3 (Pb2absent); just dorsoanterior to insertion, attached tothick pharyngeal roof CT enveloping Ebl medial endand Pb3 dorsoanterior end (dorsoanterior surface ofOD3-4 origin also attached to CT medial to LI1; CTthins considerably medially).
?
On dorsoanterior-most surface of Pb3 and medial end of small, eden-tate Pb2; dorsomedialmost edge of insertion attachedto CT to which mid-anteromedial edge of TEb2 andanteromedialmost edge of OD3-4 origin also attach;Ebl medial end not enveloped by same CT; pharyn-geal CT not thickened.LI2 on Pb3 dorsoposteriorly opposite medial endsof Eb3 and Eb4.TD comprises TEb2 and TPb3-Eb3. TEb2 broad,flat, covering entire Pb area dorsally, with mid-lon-gitudinal raphe giving rise dorsally to CT sheets at-taching to skull, attaching mid-ventrally to CT ofpharyngeal roof between Pbs; muscle attaches onEb2 dorsomedially, joining raphe with medial edgeof LE2 insertion, and overlies, but is not continuouswith, anterior end of TPb3-Eb3. TPb3-Eb3 on Pb3dorsally medial to LI2 insertion and ventral to OD3-4, continuing on to Eb3 broadly dorsomedially.
?Comprises TEb2, TEb3, and TPb3-UP4. TEb2 broad-ly continuous posteriorly with TEb3, which attacheson Eb3 dorsally medial to base of uncinate process;TPb3-UP4 completely ventral and weakly attacheddorsally to TEb3; muscle attaches on Pb3 posterioredge and adjacent dorsomedialmost edge of UP4, andis continuous posteriorly by diagonal muscle strandwith SOD.OD3-4 origin on Pb3 dorsoanteriormost surfaceventral to TEb2, medial to LI1 insertion, and dorsalto LI2 insertion and TPb3-Eb3 anteriorly; dorsoan-teriormost surface of muscle strongly attached tothick CT of pharyngeal roof; insertion on medial edg-es of Eb3 and Eb4 uncinate processes. ? Origin onPb3 dorsomedially ventral to TEb2; anteromedial-most edge attaches to CT also joined by LI1 andTEb2 (see LI1 above); muscle is dorsal to LI2 inser-tion and TPb3-UP4 anterolaterally.OP in two sections: lateral section dorsally on Eb4posteriorly beginning below LE4 insertion mediallyand extending medially to below uncinate process,medial section dorsally, narrowly on Eb4 uncinateprocess, laterally folded over medial section posteri-orly (fibers of sections continuous in crotch of fold);both sections attaching together on Cb5 medially andjoin raphe with PCI dorsoanteriorly (PCI passes be-tween two sections to insert on Cb5); medial sectionincompletely separated medially from SO. ? Medialsection joins tendinous anterior extension of PCI ven-troanteriorly and lateral section joins dorsoanteriorly
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(also, Ad5 posteriorly joins lateral surface tendon);cartilaginous and adjacent bony area of Cb5 distalend not included in attachments.Ad 1-3 absent.Ad4 dorsally broadly on Eb4 ventrally below LE4insertion, ventrally on dorsolateral half of Eb4 medialto Eb4-Cb4 joint; medially, Ad4 attachment on Cb4is separate from Ad5 attachment on Cb4, but grad-ually the two muscles meet and distally are insepa-
rable. ? See Ad5.Ad5 dorsally broadly on Cb4 posterolaterally (seealso Ad4), ventrally on Cb5 distally. ? Musculousportion on Cb4 well posteromedial to distal end, butjoins CT attaching along posterolateral edge of Cb4;entire musculous portion joins Ad4 ventrally on Cb4;posterior surface joins lateral surface of tendinous an-terior end of PCI.SOD absent. ? Present.RDs exceptionally large, slightly separated. ?Moderately large, separated by space equal to abouthalf one RD diameter.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl, Pb2, and Pb4 absent, UP4 present. Medialend of Eb4 larger than medial end of Eb3. Ebl un-cinate process absent (see also Rosen and Patterson,1990, fig. 36b). Eb4 levator process absent.
? Pbl small, partly bony. Pb2 small, rod-like,edentate. Pb4 absent, UP4 present. IAC absent. Me-dial end of Eb4 not noticeably larger than that ofEb3. Ebl uncinate and anterior processes present.Eb4 levator process present, completely covered byLE4 insertion.Pietsch (1989) hypothesized Gnathagnus Gill (ju-nior synonym of Xenocephalus Kaup) and Pleuros-copus Barnard as the plesiomorphic sister group of aclade including Kathetostoma Giinther.CHEIMARRICHTHYIDAE
In a phylogenetic study, Imamura and Matsuura(2003) excluded Cheimarrichthys as a close relativeof the Pinguipedidae, in which some authors haveincluded it. They were, however, unable to proposea sister-group relationship for Cheimarrichthys andrecognized it, as have other authors, as the sole mem-ber of the Cheimarrichthyidae.
Cheimarrichthys fosteri Haast, USNM 362725,USNM 362725, 71.6 mm.Not illustrated(superfically resembles Callanthias, Plate 132)
Description.LEI on Ebl uncinate process anteriorly ventral tocartilage tip.LE2 broadly on Eb2 dorsoposteriorly, medial edgemeeting posterolateral end of TEb2.
LE3 broadly on Eb3 anteriorly, beginning at tip ofuncinate process and extending laterally.LE4 largest levator, on Eb4 dorsolaterally, reach-ing distal end of bony surface, posterolaterally join-ing LP insertion.LP on Eb4 at and lateral to LE4 insertion.LI1 on Pb2 dorsomedially and adjacent anteriorend of Pb3, there meeting OD3-4 anteromedially.LI2 on Pb3 posterolaterally, medial edge of pos-terior half of insertion meeting TPb3-Eb3.TD comprises TEb2 and TPb3-Eb3. TEb2 band-like with mid-longitudinal raphe, which attaches ven-trally to pharyngeal roof CT and anterior edge ofTPb3-Eb3, muscle extends laterally to point anteriorto LE2 insertion. TPb3-Eb3 band-like attaching onPb3 posterolaterally beginning anterior to LI2 inser-tion, continuing along medial edge of LI2 insertion,and attaching to posteromedialmost corner of Eb3;continuous posteriorly by diagonal muscle strap withSOD.OD3-4 broadly, anteriorly on Pb3 dorsomedially,meeting LI1 posteriorly; posteriorly on medial edgesof Eb3 and Eb4 uncinate processes.OP thick, strap-like; dorsally on Eb4 uncinate pro-cess posteriorly, there meeting OD3-4, and extend-ing laterally to below medial end of LE4 insertion,which it also meets, and, at that point, overlaps dor-somedial half of Ad4 posteriorly; ventrally on Cb5posteriorly beginning laterally at posteroventral endof Ad5 and extending medially for distance aboutequal to extent on Eb4; medially clearly differenti-ated from SO on one side, unclearly on the other.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly, beginning me-dially anteroventral to OP and extending laterally todistalmost end of bony surface; ventrally on Cb4 dor-sally beginning medially ventral to a perpendicularfrom dorsomedial origin on Eb4 and extending lat-erally to distalmost end of bony surface, meeting Ad5anteriorly for most of latter's length.Ad5 short, anteriorly on Cb4 posterodistal edge,meeting Ad4, posteriorly on Cb5 dorsally, beginningmedially a little anterior to OP ventrolaterally andending near distalmost end of bony surface.SOD present.RDs separated by space equal to about one RDdiameter.Additional remarks. SCL present, apparently freefrom ventrally curving cartilaginous posterior end ofBb3. TV4 free from Cb5s. Pb4 and UP4 present. Pblshort, cartilaginous, oriented medially (horizontally).Pb2 toothed. Eb4 levator process absent. IAC pres-
ent. ScorpaenoideiImamura and Yabe (2002) hypothesized a reorga-nization and recomposition of the Scorpaeniformes
NUMBER 1 I 171
of previous authors. They recognized a scorpaenoid-serranoid sister group, on the one hand, and a zoar-coid-cottoid sister group on the other. They did notimply that the two sister groups are closely related,but assigned both to the Perciformes. Furthermore,they excluded the Champsodontidae and Normani-chthyidae as possible members of either of the twosister groups.We retain the scorpaeniform composition of earlierstudies in our arrangement of the taxa that follow,but treat the group as a suborder. We do so only forconvenience and absent of intent to imply judgmenton Imamura and Yabe's study. In fact, see additionalremarks following the description of Hexagrammos
stelleri, Hexagrammidae.SCORPAENIDAEPontinus rathbuni Goode and Bean, USNM 190360,90.8 mm. Plate 143
Additional material.
?
= Neomerinthe beanorum(Evermann and Marsh), USNM 187913, 80.6 mm.SEBASTIDAE
?
= Sebastes proriger (Jordan and Gilbert), USNM1066601, 106 mm.Not illustrated
Description.Remarks. Ishida (1994) hypothesized the phylog-eny of the scorpaenoid fishes, in which he describedvarious aspects of the dorsal gill-arch musculature.He hypothesized that the Sebastidae are the sistergroup of all other scorpaenoids. We noted few tren-chant differences in the muscles of the three generaof scorpaenoids we examined, although some carti-laginous elements of the gill arches exhibit differ-ences. We illustrate Pontinus because our preparationwas much better than those of the other two genera.LEI origin tendinous, insertion on base of Ebl un-cinate process anteroventrally. ? Origin not ob-served.
?
Origin musculous.
?
? Insertion on Ebluncinate process anterolaterally.LE2 on Eb2 mid-dorsoposteriorly.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on medial bony edge of Eb4 levator process.LP at and lateral to LE4 insertion, just extendingonto tip of levator process.LI1 broadly on Pb2 dorsoanterior process begin-ning just posterior to dorsal tip, attaching to CT join-ing medial edge of process to adjacent surface of Pb3dorsoanterior process (medial edge of Pb2 process,where LI1 inserts, just overlaps lateral edge of Pb3process); ventral surface of muscle fans out posteri-orly toward insertion such that in dorsal view it ap-pears that there is a separate dorsoanterior insertion
overlying a ventroposterior insertion along medialedge of Pb2 process.
?
Insertion more restricted, butmuscle inserts on both Pb2 and Pb3.
? Insertion doesnot fan out ventrally, insertion area relatively abouthalf that of Pontinus.LI2 on Pb3 dorsoposterolaterally at lateral edge ofTPb3 and medial to medial end of Eb3.TD comprises TPb2, TEb2, TPb3-Eb3-Eb4. TPb2centrally oblong with mid-anterior and mid-posteriornotches joined by mid-longitudinal raphe, whichgives rise dorsally to CT sheets attaching to skull;attached anterolaterally to dorsoanteriormost surfaceof Pb2 (with weak CT strands attaching to dorsoan-teriormost tip of Pb3); attached anteriorly to CT ofpharyngeal roof between dorsoanterior ends of Pb2s;muscle lying dorsal to all but small posterior portionof TEb2, ventrally joining with TEb2 along mid-lon-gitudinal raphe and fusing with TEb2 posteromedi-ally on left side (less so on right side). TEb2 mostlyventral to posterior half of TPb2 and joining TPb2along mid-longitudinal raphe, muscle attaching onEb2 dorsally well medial to LE2 insertion; well sep-arated (possibly anomalous) from TPb3 posteriorly.TPb3-Eb3-Eb4 broadly dorsolaterally on Pb3, joiningmedial edge of LI2 insertion, fine, tendinous attach-ment to medial end of Eb3 on one side and poster-omedialmost bony edge of Eb3 on other, broadly onEb4 dorsoanterior surface well medial to uncinateprocess, continuous mid-posteriorly with slenderSOD. ? ? TPb2 unnotched posteriorly, completelydorsal to central part of TEb2; TEb2 continuous pos-teriorly by fine strands of muscle with TPb3-Eb3-Eb4; fine, tendinous attachment to medial end of Eb3in both taxa.OD3-4 origin on dorsoanterior surface of Pb3 ven-tral to TEb2, insertion on dorsoanterior surface ofEb3 uncinate process and medial edge of Eb4 unci-nate process.OP dorsally beginning on Eb4 uncinate processposteriorly and extending laterally to posteroventralsurface of levator process, there meeting Ad4; ven-trally broadly on Cb5 dorsoposteriorly, overlappingAd5 ventroposteriorly; indistinguishable from SOmedially.Ad 1-3 absent.Ad4 continuous sheet beginning dorsally on ven-tral surface of Eb4 levator process and continuinglaterally to Eb4-Cb4 joint, from there attaching dor-
sally along lateral third of Cb4, meeting Ad5 narrow-ly dorsally on Cb4 ventral to Eb4-Cb4 joint.Ad5 dorsally narrowly meeting Ad4 on Cb4 pos-terior surface medial to distal end, ventrally on Cb5dorsodistally anterior to OP attachment.SOD slender.RDs separated by space equal to about half di-ameter of one RD. ? Separated by space equal to
172 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
about diameter of one RD. ? Separated by spacemore than twice diameter of one RD.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. IAC attached by looseligament to Pb2. ? IAC attached directly to Pb2. ?IAC attached directly to Pb2; AC4 present.PLATYCEPHALIDAE
Platycephalus endrachtensis Quoy and Gaimard,USNM 173884, 113 mm.Plate 144
Additional material.
?
= Inegocia japonica (Tile-
sius), USNM 99761, 67.3 mm.
Description.LEI on Ebl dorsally anteroventral to uncinate pro-cess.LE2 on dorsally expanded posterior margin of Eb2anteriorly.LE3 on Eb3 uncinate process anteriorly. ? Absent.LE4 on Eb4 levator process dorsally just medialto cartilaginous edge.LP insertion fused anteroventrally with posterov-entral edge of LE4 insertion.LI1 on Pb2 dorsomedially, continuing mediallyalong anterolateralmost edge of Pb3 adjacent to Pb2;little, if any, larger than LI2.LI2 on Pb3 dorsoposteriorly, medial edge of at-tachment meeting lateral edge of TPb3-Eb4.TD comprises TEb2 and TPb3-Eb4. TEb2 broadwith mid-longitudinal raphe giving rise dorsally tofilmy CT sheets covering muscles; anterolateral andposterolateral muscle fibers coming together and at-taching to Eb2 dorsally anterior to LE2 insertion;muscle continuous posteriorly by fine diagonal mus-cle strand with TPb3-Eb4. TPb3-Eb4 on Pb3 dorsallymeeting medial edge of LI2 insertion; muscle fibersextend dorsomedially joining ventral surface of Eb4portion of muscle, which extends well laterally andinserts on Eb4 dorsally between medial end and sur-face ventral to uncinate process; on one side, an ap-parently anomalous muscle strand of Eb4 portion in-serts on dorsomedialmost bony surface of Eb3; mus-cle continuous posteriorly by fine diagonal musclestrand with SOD.
?
Comprises TEb2 and TPb3-Eb3;TEb3 on Eb3 mid-posteriorly.OD3-4 origin finely on Pb2 anteriorly, continuingbroadly on entire dorsomedial edge of Pb3; insertionbroadly on anteromedial edge of Eb3 uncinate pro-cess and posterior surface of Eb4 uncinate process.
?
On Pb2 and Pb3 on one side, but only on Pb3 onother.OP dorsally on Eb4, extending medially from ven-tral surface of levator process to posterior surface ofuncinate process, ventrally on Cb5 dorsoposteriorly,medially continuous with SO.
Ad 1-3 absent.Ad4 in two sections; anterior section broadly dor-sally on Eb4 ventrally lateral to OR ventrally broadlyon Cb4 dorsally anterior to posterior section; poste-rior section about half as broad as anterior section,fibers angled ventrolaterally (versus almost verticalfor anterior section), dorsally on ventral surface ofEb4 levator process, ventrally on Cb4 dorsally pos-terior to anterior section, joining raphe with Ad5 onCb4. ? Consists of anterior section only.Ad5 dorsally on posterolateral bony surface of Cb4well medial to distal end, joining raphe with Ad4;ventrally on bony surface of Cb5 dorsolaterally, join-ing raphe posteriorly with OP ventroanteriorly.SOD present.RDs separated by distance greater than diameter ofone RD.Additional remarks. SCL attached mid-dorsally byCT to tip of posterior cartilaginous end of Bb3. TV4free from Cb5s. Pb4 and UP4 present. Pbl mostlybony. IAC present (see discussion below). Medialend of Eb3 larger than that of Eb4. Pb2 toothed.
?Pb4, if present, greatly reduced. IAC absent.Platycephalus appears to be more plesiomorphicthan Inegocia in having IAC, LE3, and Pb4, andmore specialized, perhaps, in having TD attaching toEb4. Only Platycephalus is included in the cladisticanalysis.Imamura (1996) published a phylogenetic analysisof the Platycephalidae and related taxa. He describedthe dorsal gill-arch musculature (1996:172-173) ofplatycephalids in general, provided a generalized il-lustration (1996:fig. 43) of this musculature in theplatycephalid Sorsogona tuberculata Cuvier, and dis-cussed variation he found in other platycephalid taxa.In so far as they are comparable, our findings agreewith his.Imamura (1996:132-133) also illustrated and re-ported briefly on the gill-arch muscles of the Bem-bridae and Hoplichthyiidae, outgroup families mostclosely related to the Platycephalidae. He illustrates(but does not label) the most plesiomorphic of these,Bembridae, as having or lacking TPb2 (depending ongenus), which muscle he reports is lacking in Hopli-chthyiidae and Platycephalidae.CHAMPSODONTIDAEChampsodon atridorsalis Ochiai and Nakamura,USNM 297752, ca. 94 mm.Additional material. USNM 297752, ca. 82 mm; C.vorax Gunther USNM 122578, 126 mm, partial in
situ dissections to determine presence of LP, q.v.Not illustratedDescription.Remarks. Mooi and Johnson (1997) present a de-tailed morphological description of Champsodon vor-
NUMBER II 173
ax, which is representative of the genus. We agreewith most of their description of the dorsal gill-archmuscles, but expand on it and note a few differences.LEI broadly on Ebl dorsally well lateral to tip ofuncinate process, beginning near point of anteriorlydeflected (normally medially directed) arm of Ebl(see Mooi and Johnson, 1997:fig. 8b for dorsal gill-arch skeletal structure).LE2 on apex of prominent, raised Eb2 triangularprocess.LE3 slender, on tips of joined Eb3 and Eb4 unci-nate process anteriorly.LE4 largest levator, on Eb4 dorsoposteriorly.LP very fine, fragile, origin near dorsoanteriormostedge of PR insertion on Eb4 co-incident with ventro-lateralmost edge of LE4 insertion.Remarks. Mooi and Johnson (1997:152) reportedLP absent, which VGS verified (ibid., p. 174), basedon what now appears to have been a defective dis-section. LP is often fragile and destroyed in acantho-morphs unless special care is taken to ascertain itspresence early during dissection while the levatorsare still attached at both their origins and insertions.LI1 on Pb3 dorsally beginning just posterior to an-teriormost tip.LI2 on Eb3 posteriorly a little lateral to medialend, penetrates OD3?4 on way to insertion.Remarks. Champsodon is the only taxon we en-countered in which LI2 inserts exclusively on an Eb.In percomorphs, LI2 usually inserts on Pb3 at, oroccasionally ventral, to articulation of Pb3 with Eb3,and it is possible that we have overlooked the factthat some LI2 filaments might be associated with thedistal end of Eb3. Penetration of OD3-4 by LI2 oc-curs uncommonly in acanthomorphs (e.g., Brotula,Ophidiidae. and related Calomopteryx, Bythitidae;synapomorphic for gobioid family Odontobutidae).TD comprises a modified TPb2 and TEb2 (togeth-er with continuous mid-longitudinal raphe), andTEb4 (no raphe). TPb2 in three parts: anterior part abroad, transversely continuous strap attaching antero-laterally to tip of Ebl uncinate process and adjacentIAC dorsomedially, also weaker attachments to ad-jacent dorsal edges of anteriormost tips of Pb2 andPb3; posterior two parts (one on each side) each con-sisting of slender muscle slip arising from surface ofTEb2 lateral to mid-longitudinal raphe and curvinganterolaterally before finely, tendinously joiningshort raphe extending posteriorly from attachment ofanterior TPb2 part to Ebl; anteroventral edge of LI1also finely joined to same raphe, which marks shal-low constriction between anterior TPb2 part andTEb2; TPb2 otherwise broadly continuous posteri-orly with TEb2. TEb2 a broad, transversely contin-uous muscle strap narrowing laterally and twisting asit passes between LI1 and LI2 and attaches on TEb2dorsally anterior to LE2 insertion; TEb2 posteriorly
is free from and dorsal to anterior edge of TEb4.TEb4 attaches broadly on posterior edge of Eb4.Remarks. TPb2 is unusual but readily derivablefrom a common acanthomorph TPb2 muscle state inwhich TPb2 consists of a depressed, roundish paddorsal to and continuous partially or completely ven-trally with TEb2. Reduction of much of the pad, leav-ing only its lateral edges and anterior portion wouldresult in the condition found in Champsodon. Insome acanthomorphs (e.g., Psenopsis, Plate 177)only the anterolaterally curving portion of TPb2 ispresent. The attachment of TPb2 to the tip of Ebluncinate process appears to be concomitant with areduction in size and ventral displacement of IAC,which, unusual for an acanthomorph, articulates withthe bony surface of the Eb 1 uncinate process ventralto its cartilage tip. While not common among acan-thomorphs, TPb2 may attach well out on the dorsalsurface of IAC (e.g., Pseudupeneus, Plate 124).OD3-4 originates very broadly on Pb3 dorsome-dially and inserts on Eb3 anteriorly ventral to tip ofuncinate process and Eb4 posteriorly ventral to tip ofuncinate process; muscle penetrated by LI2 (see re-marks following LI2).OP dorsally on Eb4 posteroventrally (slightly ven-tral to level of SO attachment on Eb4), beginningmedially ventral to LE4 insertion and extending lat-erally and meeting Ad4 dorsomedially; curvingstrongly ventromedially and attaching on Cb5 nar-rowly posteriorly medial to posterior end of Cb5;mid-medially not clearly differentiated from SO.Remarks. Mooi and Johnson (1997:fig. 12a) do notdifferentiate OP and Ad4 from SO.Ad 1?3 absent.Ad4 dorsally on Eb4 ventrally beginning medially
at lateral edge of OP and extending laterally almostto distalmost end of bony Eb4 surface, attaching ven-trally on Cb4 dorsally, meeting entire edge of Ad5attachment on Cb4.Ad5 relatively long, dorsoanteriorly on Cb4 begin-ning medial to distal end and continuing medially onCb4 for about half length of Ad5, joining raphe withAd4 attachment on Cb4; posteroventrally on Cb5dorsally beginning at distal end and extending me-dially a short distance, meeting OP insertion.SOD absent.RDs separated by about diameter of one RD.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 toothed. Eb4 levatorprocess absent.HEXAGRAMMIDAE
Hexagrammos stelleri Telesius, USNM 130279, 2specimens, 119-123 mm.Plate 145
174 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Description.LEI on raised posterior surface of Ebldorsally lat-eral to tip of uncinate process.LE2 on Eb2 mid-dorsoposteriorly, meeting distalend of posterior branch of TEb2 anteriorly and, onright side only, meeting distal end of anterior branchposteriorly (fails anteriorly to meet TEb2 posteriorlyon left side).LE3 on Eb3 just anteroventral to tip uncinate pro-cess.LE4 on Eb4 dorsoposteriorly lateral to uncinateprocess, ventrolateral margin of insertion joined byLP.LP on Eb4 dorsolaterally, joining LE4 insertionventrolaterally and extending laterally to dorsodistal-most bony end of Eb4.LI1 extends ventrally between overlapping anteriorends of Pb2 (anteroventral) and Pb3 (posterodorsal),inserting almost entirely on Pb3 ventroanteriorly, in-cluding dorsoanteromedial edge of anterior end ofPb3 dorsal to dorsoanteriormost origin of OD3?4 onPb3; a few fibers insert on Pb2 dorsolateral surface.LI2 on Pb3 dorsoposteriorly just medial to medialend of Eb3; posterior half of medial edge of insertionmeets lateral edge of TPb3-Eb3 attachment on Pb3.TD comprises TEb2, TPb3-Eb3, and, questionably,TPb2 (see remarks following TD description). TEb2medially broad, notched mid-posteriorly, with irreg-ular mid-longitudinal raphe attaching dorsally tofilmy CT sheets and ventrally to CT of pharyngealroof; medial portion lies dorsal to, and extends wellanterior to, all Pbs except Pbl; dorsolateral marginon each side of central portion forms fine, flat muscleribbon, which unites anteromedially and posterome-dially with remainder of muscle; muscle narrows andextends laterally, dividing and attaching on Eb2 dor-sally anterior and posterior to LE2 insertion (see alsoLE2); few fine muscle strands join heart-shaped por-tion posteroventrally to mid-anterior edge of TPb3-Eb3 (strands not visible in dorsal view). TPb3-Eb3on Pb3 dorsoposteriorly joining posterior half of LI2insertion medially and continuing posteriorly ontoEb3 dorsomedially; posteromedially continuous bydiagonal muscle strands with SOD.Remarks. Shinohara (1994:50) illustrated the dor-sal gill-arch musculature of the hexagrammid Pleu-rogrammus azonus Jordan and Metz. The illustrationappears generalized, and he reported no variation inthe muscles either among hexagrammids or betweenhexagrammids and his comparative material. His re-port that TDA includes attachment to Eb4 is not trueof our specimens of H. stelleri (a species included inhis material), nor is his report that LI1 inserts [only]on Pb2. The extent to which unexplored variationprobably exists in his comparative material should beobvious from the variation in muscles often reportedin the present study.
He faintly indicates a fine circular perimeter ofmuscle fibers on a central differentiated plate-likemuscle section between the right and left "arms" ofwhat is clearly TEb2, but does not label or describeit. Imamura and Yabe (2002:fig. 14A) illustrate thedorsal gill-arch musculature of Hexagrammos lago-cephalus (Pallas), which, insofar as can be told, ap-pears mostly similar to ours of H. stelleri. It differsin having TPb2 well-developed and unambiguousand in indicating that the attachment of TEb2 to Eb2is only anterior to LE2 insertion, which may be anoversight.Based on Imamura and Yabe's illustration, we rec-ognize the presence of TPb2 in our specimen of H.
stelleri.OD3-4 originating broadly on posteroventral shelfof Pb3 and inserting on joined Eb3 and Eb4 uncinateprocesses medially; dorsoanteriorly, origin meets LI1insertion on Pb3.OP dorsally on Eb4 posteriorly beginning mediallyon uncinate process, meeting OD3-4 there in a raphe,and continuing laterally to point ventral to LE4 in-sertion; ventrally, broadly on Cb5 dorsally. meetingand narrowly joining ventrolateral end of Ad5 ondorsodistalmost end of Cb5.M. SO-Pb3 absent.Ad 1-3 absent; fine GFM on anterolateral surfacesof Ebl and Cbl, and anterolateral edges of Eb2 andCb2, and Eb3 and Cb3.Ad4 dorsally on Eb4 posteroventrally, beginningjust medial to lateral edge of OP and extending lat-erally to Eb4-Cb4 joint, ventrally on Cb4 broadly an-terior to Ad5.Ad5 dorsally on Cb4 beginning posterodistally andextending a short distance medially, ventrally, nar-rowly on Cb5 dorsally, mostly anterior to OP.SOD present.RDs adjacent or narrowly separated, inserting onPb3 posteriorly.Additional remarks. SCL present. TV4 free fromCb5s. IAC reduced to one or two small cartilagescontained in ligament attaching Ebl uncinate processto Pb2. Pb4 and UP4 present. Pbl cartilaginous. Ina C&S specimen of H. superciliosus (Pallas), USNM290478 (ca. 135 mm SL), Pbl is cartilaginous on oneside and ossified, with cartilaginous dorsal and ven-tral ends, on the other. Eb4 levator process absent.The attachment of TEb2 to Eb2 both anterior andposterior to LE2 insertion was otherwise encounteredonly in Imamura and Yabe's (2002:fig. 14B) illustra-tion of Pholis nemulosa. Another similarity of H.stelleri to Pholis is that both have cartilaginous Pbls.The ligament attaching Ebl uncinate process to Pb2in Pholis appears similar to that of H. stelleri, exceptthat the latter has a couple of small pieces of cartilagerepresenting IAC included in the ligament.
NUMBER 1 1 175We believe that the TEb2 attachment, and probablythe interarcual ligament and cartilaginous Pbl, sup-ports Imamura and Yabe's (2002) hypothesis of a sis-ter-group relationship between their cottoid and zoar-coid lines. ANOPLOPOMATIDAEAnoplopoma fimbria (Pallas), USNM 269910, 149mm. Not illustrated
Description.LEI on Ebl dorsoposteriorly a little medial to mid-length.LE2 on Eb2 dorsoposteriorly a little lateral to mid-length.LE3 on Eb3 just ventral to tip of uncinate process.LE4 on Eb4 dorsally somewhat medial to distalend, meeting LP insertion anteriorly.LP on Eb4 dorsally at bony posterior edge at me-dial end of thin cartilaginous extension of distal car-tilaginous tip of Eb4, meeting LE4 insertion poste-
riorly.LI1 on Pb2 broadly anterodorsally and extendingposteriorly on anterolateralmost edge of Pb3.LI2 on Pb3 dorsoposterolaterally medial to medialend of Eb3.TD comprises TPb2. TEb2. and TPb3-Eb3. TPb2not attached to Pb2. comprises two flat, semi-circularbands dorsal to TEb2 mid-section, which floors spacebetween bands; muscle confluent mid-laterally withTEb2; anteriorly and posteriorly joining mid-longi-tudinal raphe which gives rise dorsally to CT sheetscovering muscle and is attached mid-ventrally to CTof pharyngeal roof. TEb2 extends laterally and atta-ches on Eb2 anteriorly a little lateral to LE2 insertion,muscle not continuous posteriorly with TPb3-Eb3.TPb3-Eb3 on Pb3 dorsolateral^ ventral to OD3-4,beginning along medial margin of LI2 insertion andcontinuing posteriorly and joining medialmost edgeof Eb3, broadly continuous posteriorly with SOD.OD3-4 origin broadly on Pb3 dorsoanteriorly,posteriorly on Eb3 anteriorly beginning just ventralto tip of uncinate process, meeting LE3 insertion, andon Eb4 posteriorly beginning just ventral to tip ofuncinate process.OP stringy, infiltrated with amorphous CT, begin-ning dorsally on Eb4 ventral to LP insertion and ex-tending medially to attachment of OD3-4, broadlyoverlapping Ad4 medially; ventrally on Cb5 begin-ning dorsodistally posterior to Ad5 and extendingmedially a short distance.Adl muscle flat, beginning narrowly on anteriorsurface of lateral quarter of Ebl. fanning out laterallyand covering Ebl -CI joint; not as well-developedrelatively as Adl of other fishes, but better developed
than most GFMls; it and Ad2 and Ad3, possiblyshould be treated as GFMs.Ad2 similar to Adl, but muscle extends mediallyon Eb2 and meets distal end of TEb2.Ad3 similar to Adl, but muscle extends mediallyto medial end of bony surface.Ad4 dorsally broadly on Eb4 posteriorly, begin-ning medially near about medial margin of OP andextending laterally to axilla formed by posteriorly ex-tending (but medial in orientation) cartilaginous pro-cess on distal end of Cb5 and main portion of carti-laginous distal end; ventrally much more narrowly onCb4 medial to angle formed by Cb4 and Eb4.Ad5 relatively short, ventrally on Cb5 dorsodistal-ly anterior to OP, dorsally on Cb4 distally ventral toposteriorly extending cartilaginous process (seeAd4).SOD broad.RDs robust, barely separated from each other.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 and UP4 present. Pbl absent (anterior pro-cess absent). Pb2 toothed. IAC absent. Medial end ofEb4 smaller than that of Eb3. Eb4 levator processabsent. PCI begins a little medial to distal end of Cb5and extends well medially.There is a remarkable similarity of the gill-archmuscles of Anoplopoma and the zoarcoid Bathymas-ter. RHAMPHOCOTTIDAERhamphocottus richardsonii Giinther, USNM 49141,57 mm. Plate 146
Description.LEI on Ebl lateral to broad uncinate process.LE2 on Eb2 mid-dorsoposteriorly.LE3 absent. (See additional remarks.)LE4 on Eb4 dorsoposterolaterally medial to distalend, meeting LP insertion medially.LP at and lateral to LE4 insertion.LI 1 partially on Pb2 dorsoanteriorly, but mostly ondorsoanteriormost surface of Pb3, joining raphe me-dially with OD3-4 origin present).LI2 on Pb3 dorsolaterally just medial to medialend of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2a bilateral pair of laterally convex muscle straps, notattached to any skeletal element (muscle usually at-taches to Pb2 in acanthomorphs), arising broadly,seamlessly anteriorly and posteriorly from TEb2.TEb2 broadly oblong mid-dorsally with mid-longi-tudinal raphe giving rise dorsally to filmy CT; muscleattaching beginning mid-anteroventrally and continu-ing posteriorly, to CT of pharyngeal roof; attachinglaterally on Eb2 dorsoanteriorly anterior to LE2 in-sertion; overlapping anterior end of, and posteroven-
176 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
trally continuous by fine muscle strands with, TPb3-Eb3. TPb3-Eb3 on Pb3 dorsoposterolaterally medialto medial end of Eb3 and ventral to attachment ondorsomedial end of Eb3, broadly continuous poste-riorly with SOD.OD comprises OD3-4 and OD3'. OD3-4 originbroadly on Pb3 dorsomedially, branching ventrolat-erally almost immediately into short OD3' (not illus-trated), which inserts on medial end of Eb3 dorsally,and OD3?4, which inserts on Eb3 uncinate processanteriorly and Eb4 uncinate process medially.Remarks. OD3' usually attaches to Eb3 dorsallyventral to OD3-4 or OD3 attachment to anterior sur-face of Eb3 uncinate process.OP dorsally on Eb4 posteriorly beginning on pos-terior bony surface of uncinate process and extendinglaterally to below mid-insertion of LE4, ventrally onCb5 posterolateral^, joining raphe ventrolaterallywith Ad5, inseparable medially from SO.Adl-3 absent.Ad4 dorsally on Eb4 posteriorly beginning belowmedial end of insertion of LE4 and extending later-ally to Eb4-Cb4 joint, ventrally on Cb4 dorsally me-dial to Eb4-Cb4 joint.Ad5 dorsally on Cb4 well medial to distal end,ventrally on Cb5 dorsodistally.SOD very broad.RDs adjacent.Additional remarks. SCL attached mid-dorsally tocartilaginous posterior end of Bb3. TV4 free fromCb5s. Pb4 and UP4 absent. IAC absent. Eb4 levatorprocess absent.Yabe (1985) described the dorsal gill-arch mus-culature of cottoids and various related taxa. He hy-pothesized Rhamphocottus as the sister group of allother cottoids. Although his gill-arch muscle descrip-tions are generalized, we agree with him that there islimited variation in these muscles. Yabe reported,however, that the absence of LE3 is a synapomorphyof cottoids, although in some taxa a few strands ofLE4 attach to Eb3. We found LE3, varying from welldeveloped to moderately reduced in all three speciesof Myoxocephalus we examined, two of which werealso included in Yabe's material (see Myoxocephalus,Cottidae). The presence of LE3 in Myoxocephalus iseither autapomorphic for the genus among cottoids,or more widely distributed among cottoids than Yabenoted.Yabe and Uyeno (1996) repeated Yabe's (1985)cottoid synapomorphies in excluding Normanichthyscrockeri (Normanichthyidae) from among the cot-toids. COTTIDAE
Myoxocephalus niger (Bean), USNM 70823, 2 spec-imens, 83.8 mm (second specimen, 89.2 mm, ex-
amined only for presence of LE3, LE4, LP); M.jaok (Cuvier), USNM 127057, ca. 130 mm, andM. stelleri Tilesius, USNM 54232, 94.7 mm (bothexamined only for presence of LE3, LE4, LP).
?
= Cottus carolinae (Gill), USNM 163051, notmeasured. Not illustrated
Description.LEI dorsoposterolaterally on Ebl uncinate pro-cess.
?
On Ebl lateral to uncinate process.LE2 dorsoposteriorly on Eb2 at about mid-length.
? Near distal end of Eb2 dorsoposteriorly.LE3 well developed, dorsoposteriorly on Eb3slightly lateral to tip of uncinate process. ? Absent.(See also additional remarks under Ramphocottus(Ramphocottidae
.
)
LE4 broadly on posterodorsal surface of Eb4 lat-eral to uncinate process.LP on posterodorsal surface of Eb4 at and poste-rior to LE4 insertion.LI1 has split insertion: on dorsoanterolateralmostedge of Pb3 (along origin of OD3-4 anterolaterally)and on ventroanterior surface of Pb3 sandwiched be-tween Pb3 and dorsoanterior surface of Pb2.
?
Sim-ilar, but Pb2 is absent.LI2 on Pb3 dorsal surface ventral to OD3-4 andanterior to medial end of Eb3.TD comprises TEb2 and TEb3. TEb2 attached an-teroventrally to CT dorsal to LI1 and along ventralmidline to CT of pharyngeal roof; mid-longitudinalraphe dorsally giving rise to filmy CT sheets; attach-ing laterally on Eb2 dorsally anterior to medial endof LE2 insertion; faint indication dorsally on one sideof laterally curving muscle fibers overlying main por-tion of muscle laterally (possible vestigial remnant ofTPb2); continuous by fine strand of muscle posteri-orly with TEb3. TEb3 very narrow, finely, tendi-nously attaching to Eb3 about one-fourth length lat-eral to medial end; posteriorly continuous with SOD.
?
Mid-longitudinal raphe is continuous on anteriorportion of TEb3; TEb2 extends laterally to oppositeLE3 insertion, is broadly continuous with, and over-laps slightly, anterior end of TEb3; no indication ofTPb2 fibers; TEb3 attaches musculously to the me-dial end of Eb3.OD3-4 origin on dorsoanterior surface of Pb3, in-sertion on medial edges of Eb3 and Eb4 uncinateprocesses.OD3' origin on Pb3 dorsoanteriorly ventral toOD3-4, insertion on Eb3 anteromedial surface.
?Muscle not distinct, represented as muscle fibersoriginating on Pb3 immediately medial to OD3-4 in-sertion and inserting on medial end of Eb3 at andventral to TEb3 attachment. (See remarks followingOD3' in Rhamphocottus).OP broadly dorsally on posterior surface of Eb4
NUMBER 1 1 177
uncinate process and extending laterally to belowmid-insertion point of LE4, ventrally on Cb5 dorso-lateral^, joining raphe there with Ad5; only slightlyseparated dorsolaterally from Ad4. ? Distinctly sep-arated from Ad4.Ad 1-3 absent.Ad4 dorsally on Eb4 posterolaterally, ventrallyjoining raphe with Ad5 on Cb4 dorsolaterally.
?Ventrally broadly on Cb4 dorsally medial to Eb4-Cb4joint.Ad5 dorsally on Cb4 posterior surface beginningabout one-fourth length from lateral end and extending medially to point a little past mid-length of Cb4.ventrally on Cb5 dorsodistally. ? Dorsally on Cb4 alittle medial to distal end, ventrally on distal end ofCb5 dorsolaterally.SOD broad.RDs adjacent. ? RDs separated by space equal toabout half width of one RD.Additional notes. SCL attached mid-dorsally toventrally extending cartilaginous posterior end ofBb3. TV4 free from Cb5s. Pb4 and UP4 absent.NORMANICHTHYIDAENormanichthys crocked Clark.Not illustratedRemarks. Specimens unavailable. See Yabe andUyeno (1996) for anatomical description and illustra-tions. Information below extracted from their studyand modified to be consistent with present study. Fig-ures mentioned are theirs. Although they excludedNomanichthys as having close cottoid relationships,Yabe and Uyeno were unable to propose a reasonable
sister group for it. They assigned it as incertae sedisto Scorpaeniformes on the sole basis that it has asuborbital stay.LEI, LE2, LE3, LE4, LP present.LI1 on anterior margin of Pb2.LI2 on dorsolateral margin of Pb3.TEb2 present [TPb2 absent, based on their fig. 7c].An additional transversus of undescribed attachmentspresent posteriorly.OD3-4 originating on Pb3 dorsally, insertingbroadly on Eb4, some fibers inserting on "dorsal pro-cess" of Eb3.OP not described, but present in fig. 7c.Ad 1-4 present (fig. 8).Ad5 joins Cb5 and Eb4.SOD not described, but present in fig. 7c.RDs separate, inserting on dorsomedial margin ofPb3 (fig. 3c).Slender, muscular SO branch [modified CPb?] ex-tending anteriorly lateral to Pb4 [= UP4?] and Pb3and attaching to posterolateral corner of Pb2. [Seetheir fig. 3b]. This muscle differs from M. SO-Pb2in that the latter passes medial to UP4 and Pb3.
SCL present (see their fig. 7b, but not labeled).TV4 apparently undivided. Pbl and IAC absent. Pb2toothed. Pb4? UP4 present. Uncinate and levator pro-cesses not described. Their fig. 3c illustrates an allbony Eb3 uncinate process and no indication that Eb4has either an uncinate or levator process.CarangoideiAlthough Freihofer (1978) first noted a synapo-morphy uniting the four families of carangoid fishes,Smith-Vaniz (1984) first formally defended theirmonophyly. NEMATISTIIDAE
Nematistius pectoralis Gill, USNM 82203, 153 mm.Plate 147
Description.LEI on Ebl uncinate process just lateral to carti-laginous tip.LE2 on raised dorsoposterior edge of Eb2 aboutone-third length Eb2 laterally.LE3 on medial edge of cartilaginous tip of Eb3uncinate process.LE4 broadly on Eb4 posterior surface lateral touncinate process.LP on Eb4 posterior surface beginning a little an-terior to medial end of LE4 insertion, and extendinga little lateral to LE4 insertion. Right-side LP only(not illustrated) with slender strap of LP muscle pass-ing across posterior surface of LE4 and inserting onEb4 at mid-posterior point of LE4 insertion.LI1 insertion ventral to TPb2, on dorsoanterior-most bony and cartilaginous surface of anterior endof Pb2.LI2 on Pb3 dorsolaterally, extending posteriorly tomargin of medial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2kidney-shaped pad, deeply notched mid-anteriorlywith posterior end of notch continuing as laterallyoffset raphe (one side only; raphe gives rise dorsallyto CT covering muscles); attached anterolaterally tocartilaginous anteriormost end of Pb2 at junctionwith IAC, free edge of muscle from this point later-
ally giving rise to CT sheet covering muscles; ven-trally, broadly confluent with TEb2. TEb2 ventral toTPb2 and dorsal to OD3-4 origin, extending outabout one-third distance along dorsal surface of Eb2to point anterior to LE2. TPb3-Eb3 on Pb3 dorsalsurface along medial edge of LI2, and, passing dorsalto medial end of Eb4, inserts on dorsoposterior sur-face of medial end of Eb3; continuous dorsoposter-iorly by slender diagonal muscle strap with slenderSOD.OD3-4 anteriorly on dorsomedial surface of Pb3,posteriorly on anterior surface of Eb3 uncinate pro-
178 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
cess and anterior surface and medial edge of Eb4uncinate process.OP dorsally, broadly on Eb4 posterior surface justventral to LE4 and LP insertions, ventrally, broadlyon Cb5 bony posterior surface beginning medial tocartilaginous posterior tip, posteriorly overlappingAd5 attachment to Cb5. On right-side only. OP ven-trally becomes CT, which attaches to Ad5 posteriorsurface dorsal to Cb5 (not illustrated).Ad 1-3 absent.Ad4 dorsally on ventral surface of distal one-thirdof Eb4, partly overlapped posteriorly by lateral edgeof OP, ventrally on Cb4 medial to Eb4-Cb4 joint.Ad5 on distal fourth of dorsal surface of Cb5 an-terior to OP and dorsoposterior surface of Cb4 justventral to AC.SOD present.RDs well separated.Additional remarks. SCL attached mid-dorsally toposteroventral tip of Bb3. TV4 free from Cb5s. Pb2toothed. AC dorsoposteriorly on cartilaginous distaltip of Cb4. Pb4 and UP4 present. Eb4 levator processabsent. CARANGIDAE
Selar crumenophthalmus (Bloch), USNM 189251(not measured). Plate 148
Additional material.
?
= Scomberoides to! (Cuvier),USNM 76607, 136 mm.
Description.LEI on base of Ebl uncinate process anteriorly.
?On uncinate process dorsoanteriorly.LE2 on tip of expanded bony dorsoposterior mar-gin of Eb2.
?
On prominent bony process arisingfrom posterior margin of Eb2.LE3 finely, tendinously on tip of Eb3 uncinate pro-cess anteriorly.
?
Musculously on Eb3 uncinate pro-cess anteriorly, there meeting OD3-4.LE4 on Eb4 dorsally just lateral to uncinate pro-cess.
? On Eb4 dorsally between uncinate and le-vator processes.LP on Eb4 beginning at and anterior to LE4 in-sertion and extending ventrolaterally, completelycovering broad levator process, with posterior fiberscontinuous with Ad4 dorsoposteriorly.
? Levatorprocess relatively small, muscle not continuous withAd4.LI1 on Pb2 broadly dorsally, beginning anteriorlynear articulation with IAC.LI2 on Pb3 dorsolaterally opposite medial end ofEb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3-Eb4. TPb2 dorsal to TEb2, pad-like with shallow lat-eral folds, anterior half of muscle transversely con-
tinuous, with mid-longitudinal raphe, which expandsbroadly replacing the muscle medially with an areaof thick CT; CT sheets arise from lateral edge ofraphe and from thick CT and attach to skull; muscleattaches anterolaterally to Pb2 dorsoanterior processadjacent to IAC and mid-ventrally along raphe withCT of pharyngeal roof. TEb2 joins TPb2 ventrallyalong mid-longitudinal raphe, and attaches stronglyanteromedially to Pb3 dorsoanterolaterally posteriorto LI1 insertion, and laterally on Eb2 dorsoanterola-teral to LE2 insertion (strong attachment of TEb2 toPb3 is unusual in acanthomorphs). TPb2 and TEb2not joined musculously with TPb3-Pb4-Eb3-Eb4.TPb3-Pb4-Eb3-Eb4 broad, with mid-longitudinal ra-phe on anterior half; muscle attaches to Pb3 dorso-posteriorly medial to medial end of Eb3, continuesposteriorly and attaches along posteromedial edge ofEb3, dorsal surface of Pb4, and dorsomedial surfaceof Eb4.
? TEb2 destroyed before determining if itwas attached to Pb3. TD posteriorly appears to com-prise only TPb3-Eb3-Eb4; what appears to be insame position as Pb4 in other carangids, appears tobe an elongate cartilaginous tip of Pb3 to which UP4is attached dorsally.OD3-4, OD3'. OD3-4 origin on Pb3 dorsoposter-omedially, below TEb2 posteriorly; insertion on an-terior surfaces of Eb3 and Eb4 uncinate processes;OD3' (not illustrated) splits off ventrally from OD3-4 about half-way between origin and uncinate pro-cesses and inserts on Eb3 dorsally ventral to OD3^kOP dorsally on Eb4 uncinate process posteriorly,ventrally broadly tendinously on Cb5 posterolateral-ly, joining Ad5 posteroventrally.Ad 1-3 absent, but ropy GFMs present on antero-lateral surfaces of each Eb-Cb arch.Ad4 dorsally on Eb4 posterolateral^, mostly lat-eral to OP, continuous with LP; ventrally broadly onCb4 dorsoposteriorly medial to Eb4-Cb4 joint.Ad5 dorsally on medial surface of AC4 and pos-terodistal end of Cb4, ventrally on Cb5 dorsally, join-ing tendinous ventral end of OP. ? Dorsally joiningraphe with OP.SOD slender.RDs separated by space less than one RD diameter.Additional remarks. SCL attached mid-dorsally toventroposteriorly extending cartilaginous tip of Bb3.TV4 free from Cb5s. Pb4 and UP4 present. Pb2toothed. AC4 present.A moderately well-developed Eb4 flange is pres-ent. Flange is absent in
?, moderately well developedin Carangoides crysos, very well developed in Tra-chinotus falcatus, and weakly developed and scarcelynoticeable in Selene vomer and Decapterus macro-soma. RACHYCENTRIDAERachycentron canadum (Linnaeus), USNM 341455,
1 13 mm.
NUMBER 1 ] 179
Plate 149
Description.LEI broadly on Ebl uncinate process anteriorlyventral to cartilage tip.LE2 on dorsally expanded bony posterior edge ofEb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 narrowly tendinously on Eb4 just medial totip of levator process.LP broadly on dorsal surface of Eb4 anterior to tipof levator process and anteroventral to LE4 insertion.LI1 on Pb2 dorsoanteromedially.LI2 on Pb3 dorsally medial to broad medial endof Eb3.TD comprises TEb2 and TPb3-Pb4-Eb3 (on leftside only, a flat, thin, laterally curving, semicircularstrand of muscle dorsal to TEb2, arising anteriorly,represents vestigial TPb2). TEb2 very broad central-ly, ventrally continuous along mid-longitudinal raphewith CT of pharyngeal roof, narrowing laterally andjoining medial edge of LE2 insertion, continuing lat-erally and attaching to Eb2 dorsally anterior to LE2insertion, posteriorly continuous by diagonal strandof muscle with TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 dor-solateral^ attaching to dorsoposteromedial surface ofEb3, ventrolaterally attaching to dorsoposterolateralsurface of Pb3 and dorsal surface of Pb4, continuousby diagonal strand of muscle with SOD.OD3-4, OD3' originates broadly on Pb3 dorsally
ventral to TEb2, branches ventrally just lateral to or-igin; dorsal branch (OD3-4) inserts on bony anteriorsurfaces just ventral to cartilage tips of Eb3 and Eb4uncinate processes. OD3' branches off OD3?4 ven-trally shortly after emerging posteriorly from belowTEb2, and inserts on Eb3 dorsally ventroanterior toOD3-4 insertion on Eb3.OP dorsally on Eb4 posteroventrally beginningnear medial end and extending laterally to point me-dial to levator process, ventrally on Cb5 dorsolater-al^, joining raphe with ventroposterior end of Ad5,medially continuous with SO.Ad 1-3 absent.Ad4 broadly on Eb4 ventrally beginning mediallybelow levator process and extending laterally to bonyend of Eb4, ventrally broadly on Cb4 medial to Eb4-Cb4 joint, joining Ad5 attachment on Cb4.Ad5 dorsally on Cb4 posterolaterally and adjacentAC4, ventrally on Cb5 dorsolateral^ anterior to ORSOD present.RDs separated by space less than diameter of oneRD.Additional remarks. SCL questionably free fromBb3. TV4 free from Cb5s. Pb4 and UP4 present. IACpresent. Pb2 toothed.
CORYPHAENIDAECoryphaena equiselis Linnaeus, USNM 158126, 118mm.
Additional material.
?
= Coryphaena hippurus Lin-naeus, USNM 340988, 105 mm; damaged prepa-ration. Plate 150
Description.LEI originating tendinously and joining tendinousorigin and anterior margin of LE2; inserting verybroadly on Ebl dorsoposteriorly and bony dorsoan-terior surface of long uncinate process.LE2 on expanded posterior edge of Eb2; anterioredge of muscle tendinous, attaching to tip of Ebluncinate process as muscle extends anterodorsally;tendinous edge joining tendinous dorsal extension ofLEI ventral to origin. ? Not possible to tell if LE2attached to Ebl uncinate process.Remark. Ligament (Plate 150C, not labelled) at-taches tip of Ebl uncinate process posteriorly to an-terior edge of Eb2 near anterolateral end of TEb2.LE3 tendinously on tip of Eb3 uncinate process.LE4 on Eb4 just lateral to uncinate process.LP on Eb4 beginning at ventrolateral edge of LE4insertion, continuing laterally to anterior edge ofnotched cartilaginous distal end of Eb4, crossingnotch, and ending on AC4 medial to distal end ofEb4 and dorsal to medial end of Cb4. See also OP.LI1 on Pb2 dorsoanteriorly, beginning on anteriorprocess that joins medial end of IAC.LI2 on Pb3 dorsolaterally.TD comprises TPb2, TEb2, TEb3 (or possiblyTEb3-Eb4, see remarks). TPb2 circular, pad-like,centrally thin, thickened laterally, with mid-longitu-dinal raphe, completely covering and apparentlycompletely continuous with central portion of TEb2and almost completely continuous laterally withTEb2, tenuously attached by CT only to dorsal sur-face of Pb2 anteriormost tip (attachment easily bro-ken, not illustrated); circular area overlain with thin,very tightly applied CT sheet (could not be removedwithout damaging muscle). TEb2 inserting on mostof dorsoanterior surface of Eb2, narrowly, weaklycontinuous mid-posteriorly with mid-anterior end ofTEb3. TEb3 triangular, apex anteriorly continuouswith TEb2, inserts along Eb3 dorsal surface ventralto OD3-4 and posterior to OD3', posteriorly contin-uous by diagonal muscle strap with SOD. ? TPb2deeply undercut laterally and separated from TEb2;TEb2 well separated and unconnected with TEb3;TEb3 strap-like, not triangular.Remarks. Very weak CT fibers questionably arisefrom mid-lateral dorsal surface of TEb3 and narrowlyattach to medial edge of cartilaginous tip of Eb4 un-cinate process. The connection was destroyed on both
180 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sides inadvertently and was so tenuous that we de-cided against identifying TEb3 as TEb3-Eb4.
?
Con-dition precluded determination if Eb4 connection waspresent.There is no "substantive" attachment of TD toPb3. Weak, filmy CT attaches the ventral surface ofthe combined TPb2-medial TEb2 to a tough CT sheetcovering the dorsal surface of Pb3 (OD3-4 originateson dorsolateral edge of sheet).M. Pb3-Eb2 (not illustrated) small, possibly anom-alous muscle ventral to TEb2, present on only oneside; originating on Pb3 dorsoanteriorly, joining ra-phe there with OD-3-4, and inserting on dorsome-dialmost bony surface of Eb2. ? Not present.OD3-4, OD3' on Pb3 dorsolateral^ ventral toTEb2, branches ventrally just lateral to origin; dorsalbranch (OD3-4) inserts on bony dorsoanterior sur-face of Eb3 uncinate process and bony dorsomedialedge of Eb4 uncinate process; ventral branch (OD3')inserts along most of Eb3 dorsal surface ventral toOD3-4 insertion.OP on left side broadly on posteroventral surfaceof Eb4 beginning ventral to uncinate process and ex-tending medially, dorsolaterally joining LP along fineline of CT, muscle narrows as it extends ventrally andinserts on dorsodistal end of Cb5 together with Ad5insertion. Right side OP. possibly abnormal (illustratedreversed in rear view), differs in having two separatedportions dorsally, junction with LP along line of CTmore extensive, fusing dorsoposteriorly with Ad4 dor-
sal attachment to Eb4. ? Both sides with single OPsimilar to that of left side of C. equiselis.Ad 1-3 absent.Ad4 on Eb4 ventrolaterally, ventrally on Cb4 an-terior to Eb4-Cb4 joint, posterolaterally joining raphewith Ad5 near insertion on Cb4 (not visible in PI.150).Ad5 ventrally broadly on dorsolateral surface ofCb5 and dorsally on posterodistal surfaces of Cb4and AC4.SOD present.RDs separate.Additional remarks. SCL absent. TV4 free fromCb5s. IAC broadest laterally. Pb4 and UP4 present.Pb2 toothed. Eb4 levator process absent. AC4 presentattaching to posterodistal end of Eb4 medially anddorsoposterodistal end of Cb4 (autogenous on oneside and completely fused on other in
?). Small ACspresent on first and second arches, possibly derivedfrom segmentation of distal ends of Ebl and Eb2,rather than from distal ends of Cbs. ? No ACs onfirst and second arches.Johnson (1984:497) hypothesized that Coryphaen-idae, Rachycentridae, and Echeneidae form a mono-phyletic group with Echeneidae as the sister group tothe other two families. The results of a cladistic anal-ysis reported by O'Toole (2002:617) corroborated the
monophyly of the three families, but indicated Cor-yphaenidae as the sister group of the other two fam-ilies. ECHENEIDAE
Echeneis naucrates, USNM 202201, 2: 150. 175 mmSL; USNM 206662, 145 mm.
Additional material.
?
= Remora remora (Linnaeus),USNM 181890, 82.7 mm.Not illustrated
Description.Remarks.
?
Differs from Echeneis primarily incomposition of TD and relationship of L12 to OD3^kLEI broadly on bony anterior surface of Ebl un-cinate process.LE2 broadly, dorsally, on anterior surface of ex-panded posterior margin of Eb2 at about mid-length.LE3 variable: on tip of Eb3 uncinate process an-teriorly or on joined tips of Eb3 and Eb4 uncinateprocesses anteriorly.
? Same variation (each side dif-ferent).LE4 and/or LP. See following remarks.Remarks. Whether the relatively well-developedmuscle inserting broadly on the flat bony surface ofEb4 well lateral to the uncinate process representsonly LP or also includes LE4 is unclear (the musclewas destroyed during dissection of Remora). Themuscle originates well posterior to the origins of theother levators, which originate near each other or to-gether. This probably indicates that the muscle inquestion represents LP and that LE4 is absent, whichis rare among acanthomorph fishes (also probablytrue of Pholidichthys and Spinachia). The gill-archesof echeneids are greatly appressed against the ventralskull surface and the origin of LE4 (if the muscleincludes that levator) may have been separated pos-teriorly from those of the other levators with whichit is normally associated. Additionally, the muscle oneach side of each of the three specimens appears toincorporate two incompletely separated parts. Weconclude parsimoniously that LP is present, and thatLE4 is questionably absent.
? Information unavail-able.LI1 on dorsoanterior surfaces of Pb2 and adjacentPb3; insertion is ventral to tough CT extending an-teriorly from, and forming broad mid-section ofTEb2.LI2 largest levator, penetrating OD3?4 on wayfrom origin to insertion on Pb3 broadly posterolat-erally.
? Does not penetrate OD3-4, but passes ven-tral to it, as is usual in most acanfhomorphs.TD comprises TEb2, TEb3, and TUP4. TEb2 abroad muscle on each side joined by broad medianarea of tough CT covering large, musculously nakeddorsal Pb3 facets; muscle forms anterior two-thirds
NUMBER 11 181
of lateral CT margin and attaches laterally in twoseparate areas on Eb2: on medial edge of bony pro-cess supporting LE2 and dorsally anterior to LE2 in-sertion. TEb3 a slender muscle on each side extend-ing along posterior one-third of median area of toughCT covering Pb3 dorsal facets; muscle attaches onEb3 dorsally medial to bony support of uncinate pro-cess. TEb3 continuous posteriorly by slender, diag-onal muscle strand with broad, transversely muscu-lously uninterrupted TUP4, which attaches to UP4dorsally posterior to Pb4; posterolateral corner ofmuscle meets SO dorsolaterally and is broadly con-tinuous posteriorly with SOD.
? Comprises TEb2and TUP4-Eb4. TEb2 relatively slender, forms aboutone-fourth to one-third of lateral CT margin. TUP4-Eb4 transversely musculously continuous, attachingnarrowly anterolaterally on Eb4 dorsomedially andon medial edge of UP4.OD3-4 robust, origin broadly on lateral surface ofmusculously naked Pb3 dorsal facet, insertion onbony anterior surface and medial edge of Eb3 unci-nate process and bony medial edge of Eb4 uncinateprocess; muscle penetrated by LI2, separating mostof portion inserting on Eb3 from most of portion in-serting on Eb4; posterolaterally, muscle forms raphewith OP dorsally. ? Muscle relatively flat, not pen-etrated by LI2, inserting on medial bony edges ofEb3 and Eb4 uncinate processes.OP with two separate sections; dorsally medialsection on posterior bony surface of Eb4 uncinateprocess, lateral section on posterior surface of levatorprocess; two sections meet in raphe ventrally and at-tach on Cb5 posteromedially, posteromedial to me-dial end of Ad5; dorsally, OP sections overlap muchof Ad4 posteriorly.Ad 1-3 absent.Ad4 dorsally on Eb4 beginning posteroventrallyand posterorlaterally; medially mostly anterior to ORextending laterally to end of Eb4 bony surface; ven-trally on Cb4 anterior to both Ad5 and OP.Ad5 dorsally on AC4 and distal end of Cb4 pos-teriorly, ventrally on Cb5 beginning dorsodistally andextending medially anterior to OP.SOD present.RDs moderately slender, separated by distancegreater than one RD diameter.Additional remarks. SCL present, but highly mod-ified; apparently forming circle, the anterior portionof which is a sheet of tough ligamentous tissue. TV4free from Cb5s. Pb4 and UP4 present. AC4 present.IAC absent.
Scombroidei
Johnson (1986) included the Sphyraenidae in theScombroidei; however, Orrell et al. (2003:45 and inpreparation) concluded based on a molecular study
involving outgroups and representatives of most pu-tative scombroid genera that "there is no support fora close relationship between barracudas (Sphyraeni-dae) and the Scombroidei."POMATOMIDAEPomatomus saltatrix (Linnaeus), USNM 289926, 3specimens, 113-126 mm.Plate 151
Description.LEI on Ebl uncinate process dorsoanteriorly ven-tral to cartilage tip.LE2 on raised bony dorsoposterior edge of Eb2.Remarks. In a large articulated gill-arch skeletonof Pomatomus (USNM 016528, specimen size un-known; gill arches 200 mm from basihyal to 5th Cb),a distinct, bony, prong-like process arises from Eb2dorsoposteriorly.LE3 on tip of Eb3 uncinate process dorsoanterior-iy-LE4 on dorsal edge of Eb4 levator process lateralto uncinate process, posteroventromedially joiningraphe with OP dorsally.Remarks. The cartilaginous portion of the Eb4 le-vator process varies in and among the three speci-mens: small, round cartilage, two separated small car-tilages, or a single linear cartilage, all on the posterioredge of Eb4. The cartilaginous portion(s) is well lat-eral to the uncinate process and medial to the distalcartilaginous end of Eb4. These minute levator pro-cesses persist in large specimens, as indicated byremnants of their presence in a large skeleton (seeremarks following LE2).LP at and anterior to LE4 insertion.LI1 on Pb2 dorsoanteriorly beginning posterior toIAC attachment to Pb2.LI2 on Pb3 dorsolaterally well anterior to articu-lation with medial end of Eb3 and at and lateral toattachment of TPb3-Eb3 to Pb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2squarish, dorsal to TEb2, with mid-longitudinal ra-phe, continuing posteriorly across TEb2; muscle at-tached anteroventrolaterally to dorsal end of Pb2 andadjacent dorsomedialmost surface of IAC, fusingventroposterormedially with TEb2. TEb2 attachinglaterally on dorsal surface of Eb2 at point anterior toventromedial edge of LE2 insertion, posteriorly freeand well separated from TPb3-Eb3. TPb3-Eb3 onPb3 dorsoposteriorly beginning at and medial to LI2and continuing onto dorsomedialmost surface of Eb3,continuous posteriorly by crossing muscle straps withSOD.M. Pb3-Eb2 (not illustrated, easily overlooked.)small, hidden muscle originating on Pb3 at and an-terior to LI2 insertion and inserting on Eb2 ventralto LE2 insertion.
182 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONOD3?4, OD3' origin ventral to TPb2 on dorsoan-terior surface of Pb3, insertion on Eb3 uncinate pro-cess dorsoanteriorly and on medial edge and dor-soanterior surface of Eb4 medial to uncinate process.OD3' splits off ventromedial surface of OD3?
4
shortly posterior to origin and inserts on Eb3 dorsallyanteroventral to uncinate process.OP dorsally on medial half of posterior surface ofEb4, joining raphe with LP posteroventromedially,ventrally on Cb5 dorsolaterally posterior to Ad5.joining raphe with Ad5 posterolaterally.Ad 1-3 absent.Ad4, dorsally on posterolateral surface of Eb4,overlapped posteromedially by OP, ventrally on Cb4medial to Eb4-Cb4 joint.Ad5 ventrally on Cb5 dorsally mostly anterior toOP. dorsally on posterodistalmost surface of Cb4.SOD present.RDs adjacent or separated by space less than one-fourth diameter of one RD.Additional remarks. SCL attached mid-posteriorlyto ventroposterior cartilaginous distal end of Bb3.TV4 free from Cb5s. Pb4 and UP4 present. Pbl withdorsal and ventral cartilage ends. Pb2 toothed. PCIattaches to Cb5 beginning well medial to distal endand extends medially.SCOMBROLABRACIDAEScombrolabrax heterolepis Roule, USNM 187651, 2specimens, 98.5-104 mm.Plate 152
Description.Remarks. LEI origin tendinous, all other levatorsoriginate musculously.LEI on dorsoposterior edge of Ebl just lateral totip of uncinate process.LE2 on mid-dorsoposterior edge of Eb2.LE3 on dorsomedial edge of tip of Eb3 uncinateprocess.LE4 on dorsal edge of Eb4 between tips of unci-nate and levator processes.LP at and lateral to LE4 insertion, ventrolateraledge continuous with CT sheet attaching along edgesof fourth and fifth arches and containing PR whichimpinges on LP insertion.LI1 on bony surface of Pb2 just ventral to jointwith IAC.LI2 on bony Pb3 dorsal surface lateral to TPb3-Eb3 attachment and anterior to medialmost end ofEb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2completely overlying TEb2 medially, almost circular,notched anteriorly with mid-longitudinal raphe,which attaches dorsally to CT sheets; muscle attachedmid-anteriorly to CT of pharyngeal roof, attachmentcontinuing dorsolaterally to (and over) dorsoanterior
end of Pb2, at which point flat laterally convex rib-bon of muscle arises (on each side) and continues toposterior end of mid-longitudinal raphe; ventral sur-face fused with TEb2, but muscle partially replacedby CT on mid-portion of right side (TEb2 visiblethrough CT); TPb2 strongly attached to Pb2, overliesanterior end of Pb3 to which it is loosely attached.TEb2 extending laterally on dorsoanterior surface ofEb2 anterior to LE2 insertion, well separated from,but connected mid-posteriorly by CT (not illustrated)to TPb3-Eb3. TPb3-Eb3 anteriorly broadly on Pb3dorsal surface medial to LI2 insertion, posteriorlynarrowly on posteromedial surface of Eb3, continu-ous posteriorly by transverse muscle strands withSOD.OD3?4 originating on dorsoanteromedial edge ofPb3 ventral to TEb2 and inserting on joined anteriorsurface of Eb3 uncinate process and medial edge ofEb4 uncinate process.OP dorsally broadly on posterior surface of medialarm of Eb4 beginning just lateral to levator processand extending medially to end of bony surface, mid-medially inseparable from SO, ventrally broadly ondorsoposterior edge of Cb5 mostly posterior to Ad5.Ad 1-3 absent.Ad4 broadly dorsally on Eb4 dorsoposterior sur-face beginning below levator process and extendinglaterally to bony distal end, ventrally narrowly onCb4 medial to Eb4-Cb4 joint.Ad5 broadly on Cb5 dorsolaterally, mostly antero-medial to OP, very narrowly on posterodistalmost endof Cb4.SOD present.RDs slightly separated.Additional remarks. SCL absent. TV4 free fromCb5s. Pb4 and UP4 present. Pb2 toothed. PCI atta-ches on Cb5 beginning medial to distal end and con-tinues medially; origin is from cleithrum by long ten-don. SCOMBRIDAEScomber scombrus Linnaeus, USNM 203841, 143mm. Plate 153
Description.LEI very short, on tip of Ebl uncinate process,insertion tendinous.LE2 inserts by long tendon on dorsomedialmostedge of long bony Eb2 process.LE3 inserts by long tendon on dorsomedialmostedge of cartilaginous tip of Eb3 uncinate process,muscle becomes tendinous dorsally and then mus-culous again before attaching to skull.LE4 inserts by long tendon on posterodorsal edgeof Eb4 lateral to uncinate process.LP absent.
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LI1 tendinously on Pb2 dorsally slightly anteriorto joint with medial end of Eb2, joining raphe anter-omedially with anterolateral side of TPb2, where an-teromedial edge of TEb2 meets TPb2; about samesize as LI2.LI2 on Pb3 dorsoposterolaterally.TD comprises TPb2, TEb2, TPb3-Eb3. TPb2 thin,oval, attached dorsomid-longitudinally to thin, toughCT sheet, mid-anteroventrally to CT of pharyngealroof, and anteroventrolaterally to broad cartilaginousend of Pb2 just medial to articulation with IAC, con-tinuous ventrally with TEb2 (only narrow, arcing,free lateral edge distinguishes TPb2 from TEb2).TEb2 on Eb2 dorsally medial to LE2 insertion onlong bony process and just anterior to anterior at-tachment of GFM2. TPb3-Eb3 broadly on Pb3 bonysurface beginning anteriorly at about mid-length ofbone and well medial to lateral margin and LI2 in-sertion, continuous posteriorly by thin diagonal mus-cle strands with Eb3 portion, which is on postero-medial surface of Eb3.M. Pb2-Ebl (not visible in dorsal view) on Pb2anterolaterally ventral to joint with IAC, and on Eblposterior surface ventral to uncinate process.Remarks. M. Pb2-Ebl occurs otherwise only inpre-acanthomorphs.OD3-4, OD3' origin on most of dorsoanterome-dial surface of Pb3 ventral to TEb2, dividing poster-oventrally, with short ventral branch (OD3') insertingon dorsomedial surface of Eb3 and combined dorsalinsertion on medial surface of Eb3 uncinate processventral to cartilage tip and bony dorsomedial edge ofEb4 uncinate process.OP dorsally on Eb4 posteriorly beginning medially
at point between medial end and uncinate process andextending laterally well past uncinate process, to be-low LE4 insertion, ventrally on Cb5 joining tendon(raphe) with posterior edge of Ad5.Adl-3 absent. GFM1 and GFM2 present, super-ficially appear to be RecD2 and RecD3, but are as-sociated with gill filaments. Not interpreted as Adland Ad3, because they do not extend onto associatedCbs. GFM2 on anterior edge of Eb2 anterior to prom-inent bony process supporting LE2, narrowing topoint at attachment to ventrolateral edge of bony pro-cess supporting cartilage tip of Ebl uncinate process,lateral edge of muscle associated with gill filaments.GFM3 on dorsolateral edge of prominent Eb2 pro-cess and anterior edge of Eb3 mid-laterally, lateraledge of muscle associated with gill filaments.Ad4 on ventral surface of Eb4 dorsolateral^ anddorsodistal surface of Cb4 anterior to Eb4-Cb4 joint,ventroposteriorly fusing with Ad5 on Cb4.Ad5 moderately broadly on dorsodistal margin ofCb5 and narrowly on AC4 and posterodistalmost endof Cb4, fusing at about mid-anterior surface withAd4 and ventrally joining tendinous edge of OP.
SOD present.RDs separated by distance greater than twice di-ameter of one RD, each with small separate branch.Additional remarks. SCL present attached mid-dorsally to cartilaginous ventroposterior tip of Bb3.TV4 free from Cb5s. Pb4 absent, UP4 present. Eb4levator process absent. AC4 attached to posterodistalend of Eb4 and dorsodistal end of Cb4 (also presentin Rastralliger kanagurta (Cuvier), USNM 192526;not present in Scomberomorus cavalla (Cuvier),USNM 289928, which has a similarly positioned ACbetween Eb3 and Cb3). PCI attaches on Cb5 wellmedial to distal end and continues medially.Two enlarged, modified gill-raker patches are pres-ent on each side of SO (seen in posterior view, PI.153B); internally these patches support filamentousteeth. Similarly positioned gill-raker patch noted inpresent study only in Melamphaidae.SparoideiOrrell et al. (2002), based on a molecular phylo-genetic study, provided evidence that Lethrinidae arethe sister group of the Sparidae, within a weakly sup-ported monophyletic group comprising Sparidae,Centracanthidae, Lethrinidae, and Nemipteridae. Car-penter and Johnson (2002), in a morphological phy-logenetic study (not involving muscles), however, hy-pothesized Nemipteridae (Lethrinidae (polytomusSparidae-Centracanthidae)). In another molecularphylogenetic study, Orrell and Carpenter (2004)found that Sparidae are monophyletic only with in-clusion of Centracanthidae, which was not monophy-letic. In the same study, Sparoidei was not monophy-letic with inclusion of either Nemipteridae or Leth-rinidae, nor did the latter two families form a mono-phyletic group. NEMIPTERIDAENemipterus furcosus (Valenciennes), USNM 349457,2 specimens, 93.5-104 mm.Plate 154Description.LEI on tip of bony uncinate process at mid-lengthof Ebl. Cartilage tipped Ebl uncinate process absent.LE2 on raised posterior edge of Eb2; insertion pos-terorventrally continuous with ligament attachingEb2 to Eb3 anteriorly.Remarks. Imamura (2000:214) described LE2 innemipterids as inserting on both Eb2 and Eb3, a con-dition he found limited otherwise among perco-morphs to malacanthids. We believe the state in ma-lacanthids is different from that in nemipterids; seeremarks following description of LE2 in Malacanthi-dae. The nemipterid condition is more common thanImamura recognized, although we did not alwaysnote it in our descriptions.
184 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 levator process anteriorly.LP on Eb4 beginning at LE4 insertion posterolat-erally and extending laterally.LI1 on dorsoanteriormost surface of Pb2 beginningdorsally just ventral to cartilage tip.LI2 on Pb3 dorsolaterally just anterior to medialend of Eb3.TD comprises TPb2, TEb2, and TPb4-Eb3. TPb2a pair of muscles, each laterally convex, joined toeach other posteromedially and to TEb2 ventropos-teromedially; TPb2 and TEb2 attach to dorsalmost tipof Pb2 posteriorly. TEb2 attaches mid-anteroventral-ly to CT of pharyngeal roof, with mid-longitudinalraphe, which gives rise dorsally to CT sheets cover-ing muscles; TEb2 attaches laterally on Eb2 surfacelateral to LE2 insertion; muscle not continuous pos-teriorly with TPb4-Eb3. TPb4-Eb3 anteriorly ventralto TEb2, continuous posteriorly with longitudinal SOmuscle that extends anteriorly between Pb3s; TPb4-Eb3 dorsally slender, winglike, attaching to Eb3 pos-terior edge medial to uncinate process, ventromedi-ally attaching to Pb4 dorsally, continuous posteriorlywith SOD.Remarks. Attachment of TEb2 to Pb2 is uncom-mon; also present, homoplastically, in Amarsipus(Amarsipidae).CPb (not illustrated) moderately well-developed,extending along lateral surfaces of Pb2, Pb3 and UP4and attaching to lateral ends of Pb2 and UP4, withweak anterior branch extending medially betweenPb2 and Pb3 and posterior branch between Pb3 andUP4, anterior branch joining even weaker muscle at-taching to Pb2 posteromedially and, together withposterior branch, fading into sparse SO muscle fibersof pharyngeal roof. In the smaller specimen, CPbdoes not extend anteriorly past its attachment to Pb2;in the larger specimen, muscle continues anteriorlyfrom the attachment and begins to attenuate greatly
at about the mid-anterior Pb2 margin, then fades intoSO fibers medially.OD3. OD3', OD4 all well developed, originatingmassively and essentially together from a dorsome-dially raised bony flange on Pb3 and the Pb3 surfaceventrolateral to it, and separating almost immediatelyinto OD3 and OD4 dorsally and longitudinally, andOD3' ventrally from the other two. OD3 inserts onEb3 uncinate process anteriorly, OD3' on Eb3 dor-sally ventral to uncinate process, and OD4 on Eb4levator process (uncinate process absent).OP dorsally on Eb4 posteriorly medial to levatorprocess and ventrally on Cb5 posterolaterally, joiningsmall raphe at its ventrolateralmost edge with Ad5ventroposteriorly.Ad 1-3 absent.Ad4 dorsally on Eb4 posteriorly beginning anteriorto OP and extending laterally and attaching to Eb4
ventrally near joint with Cb4, ventrally broadly onCb4 dorsally medial to Eb4-Cb4 joint.Ad5 dorsally, moderately broadly on Cb4 postero-laterally, ventrally less broadly on Cb5 dorsolateralto OP.SOD broad.RDs separated by space less than half one RD di-ameter.Additional remarks. SCL very fine, easily broken,attached mid-dorsally to cartilaginous ventroposteriortip of Bb3. TV4 free from Cb5s. Pb4 and UP4 pres-
ent. IAC absent. Pb2 toothed.Eb4 uncinate process absent, as inferred from in-sertion of LE4 on levator process (in acanthomorphs,LE4 inserts on the uncinate only in a few basal taxa,
e.g., Lampris, Velifer. Also, indicated by apparent re-duction in size of the uncinate process and juxtapo-sition of it near the levator process in the closelyrelated Lethrinus (Lethrinidae, q.v.). Carpenter andJohnson (2003:120) report that the absence of theEb4 uncinate process is apomorphic for the Nemip-teridae among the sparoids.Cartilaginous, meniscus-like pad (not illustrated)present, tightly and closely attached and conformingwith dorsal surface of ventral cartilaginous end ofPbl. Pad is easily overlooked, but appears to act ascushion between skull and Pbl, which curves aroundskull and attaches dorsally to it.Carpenter and Johnson (2002) hypothesized themonophyly of the Sparoidea, in which they includednemipterids as the sister group to lethrinids, and thesetogether as the sister group of sparids and centracan-thids.
LETHRINIDAE
Lethrinus obsoletus (Forsskal), USNM 309317, 2specimens, 84.3-103 mm.Plate 155
Additional material. ? Lethrinus harak (Forsskal),USNM 259390, 85.5 mm. ? = Gymnocranius gri-seus (Temminck and Schlegel), 350957, 115 mm.? = Monotaxis grandoculis (Valenciennes),USNM 264135, not measured.
Description.LEI broad based, on Ebl just lateral to tip of un-cinate process; origin long, tendinous.LE2 on dorsally expanded posterior margin ofEb2.LE3 finely tendinously on tip of Eb3 uncinate pro-cess anteriorly.LE4 on tips of Eb4 levator and uncinate processes(see Additional remarks).LP finely, tendinously on Eb4 lateral to and notcoincident with LE4 insertion (see Additional re-
NUMBER 11 185
marks). ? ? LP insertion joins LE4 insertion later-
ally.LI1 on Pb2 dorsolaterally beginning ventral toTPb2 attachment to Pb2 and extending ventrally.LI2 on Pb3 dorsoanterolaterally just posterior toPb2 posterolaterally.TD comprises TPb2, TEb2, and TPb3-Eb4. TPb2thick, laterally convex muscle pair lying dorsal toTEb2 and joined posteriorly by CT, which extendsanteriorly as mid-longitudinal raphe of TEb2, whichgives rise dorsally to CT sheets covering muscles andattaching to skull; anteriorly each member of TPb2pair attaches to Pb2 dorsoanteriorly; posteromedially,along extension of raphe, and just anterior to its pos-terior end, each TPb2 member is joined ventrally byTEb2, which is transversely continuous anteriorly.Anterolaterally on each side, TEb2 joins TPb2 at at-tachment to Pb2; laterally TEb2 attaches on Eb2 dor-sally lateral to LE2 insertion; lateral end of TEb2divides into anterior and posterior branches, with me-dial end of fine Ad2 inserting into divide (not appar-ent on Plate 155A); TEb2 attached mid-ventroanter-iorly between Pb3s to CT of pharyngeal roof; musclenot continuous posteriorly with TPb3-Eb4. TPb3-Eb4ventral to origin of OD3-4 and OD3', begins on Pb3dorsolaterally just medial to posteromedialmost edgeof Eb3 and is dorsomedially continuous with Eb4portion of muscle anteroventrally; Eb4 portion ismostly separate from Pb3 section, and inserts on dor-somedial bony surface of Eb4; Pb3 portion is ventro-medially continuous with SOD, which is completelyobscured in dorsal view of gill arches. ? TD includesTPb3-Eb3-Eb4 instead of TPb3-Eb4; only fewstrands of muscle attach to posteromedial surface ofEb3. ? TD includes TPb3-Pb4-Eb3 instead of TPb3-Eb4; only few strands of muscle attach to dorsome-dial surface of Pb4.CPb very prominent externally, especially anteri-orly, where it is transversely continuous; completelyencircles Pb2 and Pb3 and attaches to posterolateraland posteromedial corners of UP4; medial fibers passinto SO longitudinal fibers. ? Muscle extends onlyfrom mid-lateral surface of UP4 on one side anteri-orly around to mid-lateral surface of UP4 on otherside.OD3-4, OD3' origin broadly on Pb3 dorsomedi-
ally; OD3-4 insertion on entire anterior surface ofEb3 uncinate process and entire medial edge of Eb4leading to levator and uncinate processes; OD3' split-ting off from OD3-4 almost immediately distal toorigin and inserting on Eb3 dorsal surface ventral toEb4 insertion of OD3-4.OP dorsally on most of bony posterior surface ofEb4 medial to levator process, ventrally broadly onCb5 posteriorly, ventrally joining raphe with PCI (seeAdditional remarks).Ad 1-3 relatively weak, on anterior surfaces of rel-
evant Eb and Cb; Ad2 extending medially onto dorsalsurface of Eb2 between split lateral end of TEb2.Ad4 dorsally broadly on Eb4 posteriorly beginningmedially on levator process and extending almost tolateral end of bone; ventrally broadly on Cb5 poste-
riorly.Ad5 dorsally relatively narrowly on Cb4 postero-laterally and ventrally on Cb5 dorsolaterally.SOD present, ventral to TPb3-Eb4, not visible indorsal view in illustrated specimen, but posterior toTPb3-Eb4 and visible in dorsal view in other speci-men and taxa.RDs adjacent.Additional remarks. SCL absent (but Bb3 has avery elongate posteroventrally extending cartilagi-nous end, normally present when SCL is present).TV4 free from Cb5s. IAC present. Pbl bony withcartilage tips. Pb2 toothed. Pb4 and UP4 present.Displacement of LP insertion from joining ormeeting LE4 insertion appears to be unique to Leth-rinus among acanthomorphs.In lethrinids, the uncinate processes on Eb3 andEb4 are not bound together as they are in most acan-thomorphs. The bony support of the Eb4 uncinateprocess is indistinguishable from that of the levatorprocess and its cartilaginous tip, when present, isgreatly reduced (vestigial) and barely separated fromthat of the levator process. In the nemipterids, theuncinate process has been entirely lost. The associ-ation of LE4 with the tip of Eb4 uncinate process isnot the same condition as occurs in Caristius (Car-istiidae) in which the uncinate process is well devel-oped and in its usual position (joining Eb3 uncinateprocess) and there is no Eb4 levator process.PCI inserts broadly on Cb5 reaching the cartilagetip of the element in Gymnocranius and Monotaxis,failing to extend even near the tip in both species ofLethrinus. OP joins raphe with PCI.CENTRACANTHIDAESpicara smarts (Linnaeus), USNM 269800, 2 speci-mens, 86.0?90.6 mm.Plate 156SPARIDAE
?
= Acanthopagrus bifasciatus (Forsskal), USNM191682, 71.8 mm.
Additional material. ? = Lagodon rhomboides (Lin-naeus), USNM 143843, not measured. ? = Sarpasalpa (Linnaeus), USNM 343618, 73.6 mm.Not illustratedRemarks. Carpenter and Johnson (2002) hypothe-sized that centracanthids and sparids form a mono-phyletic group "with placement of centracanthids un-resolved with respect to sparid genera [p. 114]"
186 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
among the Sparoidea, which also includes nemipter-ids and lethrinids.Except for CPb, muscles for all three sparid taxawe include were recorded only as present or absent,and all are present as in Spicara. although the finerdetails of the descriptions for Spicara may not applyto the sparid taxa.
Description.LEI on dorsoanterior surface of bony Ebl uncinateprocess.Remarks. The medial edge of the tip of the unci-nate process is minutely cartilaginous and easilyoverlooked.LE2 narrowly tendinously on raised posterior edgeof Eb2 anteriorly.LE3 finely tendinously on tip of Eb3 uncinate pro-cess anteriorly.LE4 finely tendinously on tip of Eb4 levator pro-cess anteriorly.LP on dorsal surface of Eb4 lateral to levator pro-cess.LI1 on Pb2 dorsally at base of cartilage-tipped dor-
sal process that articulates with IAC.LI2 on Pb3 dorsolaterally medial to medial end ofEb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 pair of flat, laterally curving muscles dorsal toTEb2; each member of pair attaching anteriorly toPb2 cartilage-tipped anterior process; pair joined pos-teromedially by raphe, which continues anteriorlyalong center of TEb2; dorsally, raphe gives rise toCT sheets covering muscles and attaching to skull;TPb2s join TEb2 at raphe, which is attached ventrallyto CT of pharyngeal roof between Pb3s; TPb2 andTEb2 not continuous posteriorly with TPb3-Pb4-Eb3.TEb2 broad, extending laterally and attaching to Eb2dorsally anterolateral to LE2 insertion, meeting me-dial end of Ad2, failing to cover dorsoposterior Pb3surfaces in illustrated specimen (probably unusual),but does so in other specimen and three sparid generaexamined. CPb. TPb3-Pb4-Eb3 on Pb3 dorsoposter-olaterally at medial end of Eb3, continuing posteri-orly on small Pb4 dorsally, and extending well lat-erally and attaching finely on posterior bony edge ofEb3 medial to uncinate process and posterior toOD3'; TPb3-Pb4-Eb3 continuous posteroventrally bycrossing strands of muscle with SOD.CPb (not illustrated) relatively weak musclestrands beginning on posterolateral corner of UP4,continuing anteriorly along Pb3 laterally and attach-ing to posterolateral end of autogenous tooth plate(Pb2'; see also remarks) joined to Pb2 posteriorly andabsent from Pb2' laterally, but beginning again onPb2' anterolaterally and continuing anteriorly aroundPb2 and across mid-line of arches to opposite side;muscle strands from posterior end of Pb2' also pass
medially between Pb2 and Pb3, with short branchattaching to Pb2 posteromedially and longer branchcontinuing posteriorly along medial side of Pb3 andattaching to posteromedial side of UP4, with strandsalso passing laterally between Pb3 and UP4 and join-ing strands passing anterolaterally to UP4 and Pb3.Muscle absent on Pb2 laterally, Pb2' mid-laterally,and UP4 posteriorly.Remarks. CPb present in ? and ?, in whichstrands may be continuous anteriorly along Pb2' lat-erally, but absent in ?.Pb2', which articulates closely with the posteriorend of Pb2, has been treated as an Eb2 tooth plate(e.g., Carpenter and Johnson, 2002: 120, character no.34; see also discussion by Johnson (1992:19, item 4))and is arbitrarily treated here as a part of Pb2.OD3-4, OD3' origin broadly on Pb3 dorsally, an-teriorly ventral to TEb2; muscle dividing ventroan-terolaterally well lateral to origin, with separate ven-tral insertion (OD3') on Eb3 dorsally ventroanteriorto uncinate process, meeting medial end of Ad3, and(OD3?4) on Eb3 uncinate process anteriorly and Eb4uncinate and levator processes medially.OP dorsally on Eb4 posteriorly medial to levatorprocess, ventrally on Cb5 dorsoposteriorly, laterallyjoining raphe with Ad5 medially.M. Pb3-Cb5 diagonal strap of muscle attaching toPb3 posteriorly and extending posteriorly medial tomedial end of Eb4, then ventrally and attaching ten-dinously to posterodistal surface of Cb5.Remarks. Muscle appears restricted to centracan-thids and sparids and provides additional evidence ofthe close relationship of these two groups.Adl?3, short, each on anterodistal surfaces of rel-evant Eb and associated Cb.Ad4 dorsally on Eb4 posteriorly beginning on le-vator process and extending laterally, ventrally onCb4 dorsally medial to Eb4-Cb4 joint.Ad5 anteriorly on posterodistal surface of Cb4 andposteriorly on Cb5 dorsally, joining raphe dorsome-dially with OP ventrolaterally.SOD present.RDs separated by space about one-half diameterone RD.Additional remarks. SCL present (also
?). ? ?Attached mid-dorsally to tip of ventroposteriorlycurving cartilaginous end of Bb3. TV4 free fromCb5s. Pb4 and UP4 present.Carpenter and Johnson (2002) described the pe-culiar relationship of Pbl to Ebl in centracanthids.The cartilaginous ventral end of Pbl tightly joins andconforms with the dorsomedial surface of the anteriorarm of Ebl, whereas the two elements are looselyarticulated in the sparids. lethrinids and nemipterids.Additionally, we found that Pbl of centracanthids ap-pears to have pivoted laterally from its articulationwith Ebl and that it is the ventrolateral surface of
NUMBER 1 1 187
the cartilaginous ventral end that is joining Ebl.Also, there is a thin wafer of fibrocartilage (not il-lustrated in Plate 156) that is tightly joined to thedorsal surface (or dorsomedial surface if Pbl wereupright) of the cartilaginous ventral end of Pbl. Thewafer rests against the ventral surface of the craniumand probably serves as a cushion.
Girelloidei
Girelloidei here coined as a subordinal taxon tofacilitate designating a group of families (Girellidae,Scorpididae, Microcanthidae, Kyphosidae, Kuhliidae,Terapontidae, Arripidae, Oplegnathidae, but exclud-ing those belonging to the Stromateoidei) that John-son and Fritsche (1989) hypothesized formed amonophyletic group. The group, including stroma-teoids, was based primarily on their possessing Freih-ofer's (1963) pattern 10 of the ramus lateralis acces-sorius (RLA). Johnson and Fritsche did not examineAmarsipus, sole member of the Amarsipidae, whichhas been included as a stromateoid since its originaldescription (Haedrich, 1969). Amarsipus lacks thestriking complex specialization that characterizes allother stromateoids (see both Amarsipus and Icosteus,Icosteidae, for discussions of the inter-relationshipsof Amarsipus), and if it is a stromateoid, is mostprobably the sister group of all other stromateoids.We examined Amarsipus for the presence of pattern10 of RLA. and found it absent, but are uncertainwhich of the other patterns it possesses. Amarsipus'srelationship with other stromateoids is, therefore, anopen question.We only examined the musculature of three of thefamilies of Girelloidei, and noted no informationbearing on its monophyly.GIRELLIDAE
Girella simplicidens Osburn and Nichols, USNM167579, 95.6 mm, USNM 321278, 78.0 mm.Not illustrated
Description.LEI on Ebl posteriorly at about mid-length andwell lateral to tip of horizontally directed uncinateprocess; tendon runs along lateral surface of muscle.LE2 on Eb2 posteriorly at about mid-length; slen-der tendon runs along ventral half of lateral edge ofmuscle.LE3 on tip of Eb3 uncinate process medially.LE4 on Eb4 posteriorly projecting levator processdorsally.LP on Eb4 dorsally a little lateral to LE4 insertion.Remarks. Among percomorphs, only Girella, Ra-chycentron (Rachycentridae), Spicara (Centracanthi-dae), lethrinids, and several atherinomorphs have LPinserting completely separate from LE4. In other per-
comorphs, the two muscles almost always insert to-gether and the insertions are often fused.LI1 on Pb2 dorsally just posterior to anteriormosttip; about same size as LI2.LI2 finely, tendinously on Pb3 dorsally immedi-ately medial to articulation with medial end of Eb3,near, if not bordering lateral edge of TPb3-Pb4-Eb3on Pb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3(TPb3-Pb4-Eb3-Eb4 in smaller specimen). TPb2 athick, laterally curving, semicircular muscle on eachside dorsal to TEb2 and ventral to thick CT pad;muscle arises posteriorly from posterolateral edge ofmoderately broad CT area, which narrows anteriorlyinto mid-longitudinal raphe as it extends acrossTEb2, gives rise to thick CT pad dorsally, and con-tinues anteriorly with CT of pharyngeal roof; anter-omedially muscle fades into TEb2; muscle impingeson Pb2 dorsoanteriorly, but is at most weakly, if at
all attached to Pb2. TEb2 extends laterally onto Eb2dorsally to point anterior to LE2 insertion, meetingmedial end of GFM2. TPb3-Pb4-Eb3 beginning onPb3 dorsolaterally a little anterior to articulation withEb3, extending posteriorly and attaching to posteriorcorner of medial end of Eb3 and, not visible exter-
nally, ventrally attaching on dorsal surface Pb4 (andon dorsal surface of medial end of Eb4 in smallerspecimen); muscle continuous posteroventrally byfine muscle strands (not visible externally) with SOD.OD3-4 anteriorly on Pb3 dorsomedially ventral toTEb2, posteriorly on Eb3 broadly anteriorly begin-ning just ventral to tip of uncinate process and onEb4 anteriorly beginning ventrolateral to uncinateprocess (muscle divides as it extends posterolateralfrom medial edge of Eb3 uncinate process).OP dorsally on Eb4 ventrally, beginning a littlelateral to medial end of Eb4 and extending laterallyand curving posteriorly as it follows posteriorly pro-jecting bony Eb4 shelf, but ending on shelf well an-terior (medial) to its posterior cartilage tip (levatorprocess); ventrally on Cb5 posteriorly, beginning me-dial to distal end and continuing medially a shortdistance past attachment of Ad5; lateral edge of mus-cle is tendinous, becoming fascia-like on attaching toCb5 and joining dorsomedial edge of Ad5; mediallymuscle is indistinguishable from SO. When gill arch-es are viewed posteriorly, only OP dorsal and ventralportions are visible, as Ad4 occludes mid-portionfrom view.Ad 1-3 absent; moderately well-developedGFMsl-3.Ad4 very broad, dorsally on Eb4 ventrally, begin-ning medially anterior to dorsolateral end of OP, ex-tending laterally posterolaterally around posteriorlyprojecting Eb4 shelf (supporting levator process),then laterally to end of bony Eb4 surface; ventrallyrelatively narrowly on Cb4 dorsally medial to Ad5.
188 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ad4, other than SO, is main muscle visible in pos-terior view of gill arches.Ad5 relatively small, anteriorly on posterodistalsurface of Cb4 and Eb4 at joint and lateral to OPlaterally, joining CT with OP ventrolaterally; poste-
riorly on Cb5 posterodistally.SOD present.RDs adjacent.Additional remarks. SCL weakly attached mid-dorsally to ventrally projecting cartilaginous posteriorend of Bb3. TV4 free from Cb5s. Pbl present, mostlybony. Pb2 toothed. Pb4 and UP4 present. IAC pres-ent. Medial end of Eb4 larger than medial end ofEb3. Eb4 flange absent.
KUHLIIDAE
Kuhlia mugil (Forster), USNM 1 14998, 2 specimens,87.0-94.8 mm. Plate 157
Description.LEI on bony surface of Ebl uncinate process be-ginning just lateral to joint with IAC.LE2 on dorsalmost edge of raised dorsoposteriormargin of Eb2.LE3 on Eb3 extending laterally from medial edgeof cartilage tip of uncinate process.LE4 on Eb4 bony surface just medial to cartilagetip of levator process.LP on Eb4 bony surface beginning just anterior tocartilage tip of levator process and extending to andjoining LE4 insertion anteriorly.LI1 insertion mainly on dorsoposteriormost edgeof Pb2 anterior process, with CT attachments to IACdorsoposteromedialmost surface (adjacent to Pb2 in-sertion) and anterolateralmost edge of TPb2.LI2 on Pb3 dorsoposteriorly immediately medialto anteromedialmost edge of Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb3.TPb2 divided; medially concave cord-like muscle oneach side dorsal to TEb2; attaches anteromedially todorsoanteriormost end of Pb2 with membranous con-tinuations onto IAC medially and adjacent LI1 inser-tion; anteromedially joining irregular mid-longitudi-nal raphe with TEb2 anteriorly, posteromedially fad-ing into TEb2; CT sheets arising from irregular mid-longitudinal raphe attach also on surface of TPb2.TEb2 flat medially, with irregular mid-longitudinalraphe, which is continuous ventroanteriorly with CTof pharyngeal roof; muscle thickening laterally andattaching broadly dorsally on Eb2 anteroventral toLE2; muscle discontinuous with TPb3-Pb4-Eb3.TPb3-Pb4-Eb3 attaching on Pb3 dorsolaterally ven-tral to OD3-4 and just medial to mid-medial edge ofEb3, continuing posteriorly on Eb3 posteromedialedge and ventrally on Pb4 dorsally (attachment on
Pb4 obscured in dorsal view), continuous by diagonalmuscle strand with SOD.OD3-4 origin broadly on Pb3 dorsoposteromedi-ally ventral to TEb2, insertion broadly on Eb3 dor-soanteriorly beginning just ventral to tip of uncinateprocess and narrowly on medial edge of Eb4 justventral to tip of uncinate process or on anterior sur-face just ventral to tip of uncinate process.OP dorsally on Eb4 posteriorly beginning at me-dial end of bony surface and extending laterally tobelow, or slightly lateral to, uncinate process, ven-trally joining tough, clear CT (not illustrated) sur-rounding Ad5, and attaching to Cb5 ventromediallycontinuous with SO.Ad 1-3 absent (GFMs moderately developed).Ad4 dorsally on Eb4 posteriorly beginning slightlyanteromedial to lateral edge of OP and extending lat-erally to below levator process, there becomingsharply less robust and extending somewhat ventrallyto end of bony surface; ventrally, narrowly on Cb4dorsally medial to Eb4-Cb4 joint.Ad5 dorsally on Cb4 posterodistally and AC4 ven-trally; ventrally on Cb5 posterolaterally. Muscle en-cased in tough, clear CT (removed in Plate 157),which is joined by OP ventrally.SOD present in both specimens.Johnson (1993:9) reported that SOD is absent inKuhlia (see also remarks following SOD in Toxotes(Toxotidae) description).RDs adjacent or very slightly separated.Additional remarks. SCL attached mid-dorsally toventroposteriorly extending cartilage tip of Bb3. TV4free from Cb5s. Pb4 and UP4 present. Pb2 toothed.AC4 present (also in cleared and stained specimen ofK. sandvicensis (Steindachner), USNM 28947 1
.
TERAPONTIDAE
Leiopotherapon unicolor (Gunther), USNM 173654,96.3 mm; USNM 173858, 120 mm, cleared andstained gill arches. Plate 158
Additional material. ? = Terapon jarbua (Forsskal),USNM 173657, 81.1 mm.
Description.LEI slender, on tip of Ebl uncinate process; mus-cle and Pbl attached anteriorly to pharyngeal roofCT. ? Not especially slender.LE2 finely, tendinously on Eb2 mid-dorsoposter-iorly. ? Insertion musculous, not fine.LE3 on Eb3 uncinate process dorsoanteriorly justventral to cartilage tip.LE4 on Eb4 posteriorly projecting levator process;ventroposterolateral and ventroposteromedial fiberscontinuous with Ad4. ? LE4 and Ad4 not continu-ous.
NUMBER 11 189LP on Eb4 beginning at ventro-anterolateral edgeof LE4 insertion and continuing laterally a short dis-tance; posterolateralmost fibers continuous with Ad4dorsally. ? Fibers not continuous with Ad4.LI1 on Pb2 dorsally just posterior to anteriormosttip. ? On Pb2 similarly, but, with additional attach-ment to posterior surface of Pb2-IAC joint.LI2 on Pb3 dorsolaterally just medial to medialend of articulation with Eb3.TD comprises TPb2, TEb2 and TPb3-Eb3. TPb2a thick, irregularly round, concave pad dorsal toTEb2, with mid-posterior notch extending anteriorlyas raphe, from which tough, filmy CT sheets ariseand also attach to pad dorsolaterally; muscle attachesanterolaterally to Pb2 and fuses ventrally with mid-medial area of TEb2. TEb2 attaches by CT mid-ven-trally to CT of pharyngeal roof, extends on Eb2 dor-sally to medial surface of tiny, low, diagonal bonystrut anterior to LE2 (medial end of GFM2 attachesto lateral surface of strut). TPb2 and TEb2 not con-tinuous with TPb3-Eb3. TPb3-Eb3 on Pb3 postero-laterally beginning just anterior to LI2, continuingposteriorly medial to LI2 insertion and extendingonto Eb3 posteromedially; muscle continuous poste-
riorly by diagonal muscle strand with SOD. ? Strutweakly developed, would have been overlookedwithout knowledge of occurrence in Leiopothera-pon?development possibly related to size of speci-men.OD3-4, OD3' originate together on Pb3 mediallyventral to TEb2 and TPb3-Eb3, extend laterally withshort OD3' branch separating anteroventrally and in-serting on Eb3 dorsally ventral to uncinate process,there meeting medial end of GFM3. Major portionof muscle continues laterally with anterior portion in-serting broadly on Eb3 uncinate process anteriorlyand posterior portion inserting on Eb4 uncinate pro-cess posteriorly, with fibers passing between uncinateprocesses and inserting on anterior surface of Eb4just ventral to tip of uncinate process. ? OD3' absent.OP dorsally on Eb4 posteriorly beginning mediallynear end of bony surface and extending laterally tojust medial to uncinate process; ventrally on Cb5 be-ginning laterally as broad CT raphe with mid-poste-
rior surface of Ad5 and continuing medially aboutsame extent as muscle occupies on Eb4.GFM1-3 each begin as fragile sparse muscle fanon anterodistal surfaces of respective Eb and Cb.GFM1 continues dorsoanteromedially a short dis-tance on Ebl. GFM2 continues dorsomedially be-coming dorsal and more compact and inserting onlateral surface of tiny, bony diagonal strut anterior toLE2 insertion (TEb2 inserts on medial surface of
strut). GFM3 follows path similar to GFM2, endingnear OD3'. ? OD3' absent.Ad4 dorsally begins on Eb4 dorsoposteriorly lat-eral to LP insertion, extends medially to uncinate
process, with few muscle strands continuous with LPposteroventrally and others joining raphe with LE4posteromedially; ventrally, muscle attaches on Cb4dorsally beginning just medial to medial end and ex-tends medially about a quarter length of Cb4. ? Mus-cle completely separated from LE4 and LP.Ad5 on Cb4 and dorsoposterodistally and ventrallyon Cb5 beginning dorsodistally and extending me-dially slightly less than distance occupied by Ad4;posterior surface joins broad CT raphe with OP ven-trally.SOD present.RDs adjacent.Additional remarks. SCL attached mid-dorsally toposteroventrally extending cartilaginous tip of Bb3.TV4 free from Cb5s. Pb4 and UP4 present. AC4 pre-sent on both sides, but present on only one side andin two pieces in cleared and stained specimen. Nei-ther AC4 nor any indication of a posterior cartilagi-nous extension of distal end of Cb5 present in
?.
LabroideiCICHLIDAECaquetaia kraussii (Steindachner), USNM 258004,62.8 mm. Plate 159
Additional material (lengths not recorded). Astrono-tus ocellatus (Agassiz), USNM 329642; Cichlaocellaris (Bloch and Schneider), USNM 226019;Cichlasoma bimaculatum (Linnaeus), USNM181457; Copadichromis jacksoni (lies), USNM261845; Crenicichla aha Eigenmann, USNM226024; Cyrtocara moorii Boulenger, USNM280311; Santanoperca leucosticta (Muller andTroschel), USNM 289647; Paratilapia polleni,USNM 344609; Ptychochromoid.es katria Reinthaland Stiassny, USNM 344607; Ptychochromis oli-gacanthus (Bleeker), USNM 344605.Remarks. Muscles of all the taxa are generally sim-ilar. The nature and relationships of LE4, LP, and OP,however, are particularly complex (see discussion inAdditional remarks), and variable among the taxa.The description of these three muscles is based al-most entirely on Caquetaia. Only a few variationspertaining to the other muscles are mentioned in thedescription.
Description.LEI on and lateral to Ebl uncinate process.LE2 on Eb2 mid-dorsoposteriorly.LE3 on Eb3 uncinate process ventral to tip and atOD3-4 attachment to Eb3.LE4 essentially free, inserting on Eb4 only at pointposterolaterally where LE4 joins with LP insertionmedially; continuous ventrally with central portion of
190 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTONOP (separation shown by a raphe), which inserts onCb5.LP on Eb4 laterally, insertion fusing with LE4 me-dially and partly joining raphe with putative lateralpart of OP dorsal ly.LI1 on Pb2 dorsoanteriorly and Pb3 ventroanter-iorly (sandwiched between Pb2 and Pb3).LI2 dorsolaterally on Pb3 lateral to OD3-4 originand medial to medial end of Eb3.TD comprises TPb2, TPb2a, TEb2 and TPb3-Eb4.TPb2 is deeply notched mid-anteriorly and almostcompletely divided mid-longitudinally; the divisionis an expansion of the mid-longitudinal raphe fre-quently present in the acanthomorph TD. The divi-sion also completely divides TEb2, but not TPb3-Eb4. TPb2 attaches to Pb2 dorsoanterolateral process(process not visible externally) and to tiny IAC; pos-teriorly, TPb2 joins CT sheet that covers Pb3 artic-ulating surfaces. TPb2a has mid-anterior raphe andattaches to anterior Pb2 surfaces ventral to TPb2; CTextends posteriorly from raphe and passes betweenPb2s and Pb3s; separation of TPb2 from TPb2a isindistinct (but separation may be distinct in othercichlids). TEb2 a pair of muscles, joined medially byCT sheet covering Pb3 articulating facets, and at-taching laterally on Eb2 to position anterior to LE2insertion. TPb2 and TEb2 well separated, not contin-uous posteriorly with TPb3-Eb4. TPb3-Eb4 attacheson Pb3 posterolaterally slightly ventral to attachmentto posteromedial end of Eb4 and continues slightlyanteromedially on Pb3. Posteriormost component insome other cichlids attaches only to Pb3 (see alsoTable 9; and discussion in Additional remarks).CPb comprises a pair of well-developed sub-epi-thelial muscles (Anker, 1978:261), each originatingposterolaterally on UP4, muscle divides anteriorlywith branch attaching to Pb2 laterally and branchpassing medially and attaching to Pb3 anteromedi-
ally.OD3-4 origin on Pb3 articulating facet anterolat-erally, insertion on medial edge of Eb3 uncinate pro-cess and on medial edge of, and enveloping Eb4 un-cinate process, joins raphe posteroventrally with a OPmedial portion.OP comprises four parts (apparently not all men-tioned or illustrated in the literature; see discussionin Additional remarks): OP1, OP2, OP3, and OP4.The first three parts attach ventrally near the distalend of Cb5 and the fourth part attaches ventrally,broadly on Cb5 dorsally medial to the distal end. OP1(appears to be the same as Aerts's (1982:233) parslateralis) is dorsally on Eb4 posteriorly just medial tothe membranous dorsal attachment of Ad5; dorso-posteromedially it joins a small raphe with LP ven-troposteriorly and is continuous ventromedially withOP2 and ventrolaterally with Ad5. OP2 (the same asAerts's (1982:233) pars centralis) is continuous with
LE4 ventrally, their junction indicated by a raphe.OP3 is on Eb4 dorsally, extending broadly mediallyfrom the uncinate process and joining a raphe withOD3?4 (dorsal attachment appears to be similar toAerts's (1982:233) pars medialis). OP4 is on most ofthe posteroventral edge of Eb4 (ventral attachmentappears to be similar to that of Aerts's (1982:233)pars medialis).Adl small, on Ebl anterolaterally and Cbl anter-odistally.Ad2 and Ad3 well developed, on anterolateral halfof respective Eb and anterodistally on respective Cb.Ad4 (not visible in illustration) dorsally, broadlyon Eb4 ventral surface, fusing posteriorly with OP4dorsoanteriorly, ventrally, narrowly on Cb4 dorsalsurface medial to Eb4-Cb4 joint.Ad5 dorsally, tendinously primarily on posterolat-eral end of Eb4, secondarily on posterodistal end ofCb4 dorsally; ventrally on posterodistal end of Cb5.SOD absent, but strap-like branch of SO arises lat-erally on each side and attaches to Pb3 mid-poster-oventrally, giving impression of an interrupted SOD.RDs adjacent.Additional remarks. SCL free from Bb3 (SCL ab-sent in Cichla and Crenicichla). TV4 with continuousventral portion across Cb5 and split dorsal portionattaching laterally to each Cb5 anteriorly (ventralportion absent in Cichla; also reported absent in var-ious African cichlids by Stiassny, 1992:265?267).Pb4 absent, UP4 present. IAC present, small.Anker (1978) termed our TPb2 as M. craniophar-yngobranchialis 2 and our TPb2a as M. transversuspharyngobranchialis 2. Both muscles appear to bederived from TPb2, and the second, which is not al-ways separate from the first in cichlids, should notcarry the main part of the name. TPb2a in acantho-morphs is a specialized part of TPb2 that is restrictedto "labroid" (sensus Stiassny and Jensen, 1987),pseudochromid, atherinomorph, and pholidichthyidfishes, and is questionably synapomorphic for thesefishes as group.Anker (1978:256-257) described the musculatureof the cichlid Haploehromis elegans Trewavas. Hetermed the posteriormost TD element the transversusepibranchialis 4, and described it as attaching to Pb3,Pb4 [actually UP4. Pb4 is absent], and Eb4. Stiassny(1981:96), who examined the dorsal gill-arch mus-culature in several cichlids (including some generawe also examined), followed Anker in recognizing aTEb4. She appears to have noted no variation in theattachment of this muscle, which she described asoriginating "from the caudal eminence formed at thejunction of Pb3 and UP4." In cichlids there is a tightassociation of UP4 dorsally with Pb3 at the joint ofPb3 with the medial end of Eb4, and the three ele-ments are bound by CT. The posteriormost TD mus-cle is attached to this complex dorsally, and as such
NUMBER 11 191
is removed from UP4. The posteriormost TD musclenever attaches to the bony portion of UP4, but mayattach to Eb4 and Pb3 or only to Pb3 (Table 9).Remarks. Aerts (1982) studied the development ofOP and LE4 in Haplochromis elegans Trewavas. Hereported that during ontogeny OP forms in three parts(lateral, central, and medial) and that LE4, which isinitially separate, combines with the dorsal fibers ofthe OP medial part. Claeys and Aerts (1984) dis-cussed further the ontogeny of LP, LE4, and OP inH. elegans (which they placed in Astatotilapia). Theyfound that during ontogeny a few fibers of LP attachdorsolaterally to Eb4, but most become attached tothe aponeurotic system of Eb4 dorsolaterally directlyopposite the insertion area of the lateral bundle ofOP and later join end to end with them to form acompound muscle. Thus, for H. elegans, and manyor most other cichlids, the "sling" comprises LE4,OP2, and LP, although we find that the contributionof LP to the sling is usually considerably more lim-ited than that of LE4. We further note that OP1 mayfuse almost completely with Ad5 (e.g., Ptychochrom-is). POMACENTRIDAE
Dischistodus fasciatus (Cuvier), USNM 328190,three specimens. 57.1-68.9 mm; USNM 179622,79.7 mm. Plate 160
Additional material. ? = Abudefduf sexfasciatus (La-cepede), USNM 221863, 68.4 mm; <D = Chromisamboinensis (Bleeker), USNM 338153, not mea-sured. Plate 161
Remarks. We also recorded limited data on thedorsal gill-arch musculature of several other species(data variably combined or assigned to individualspecies as warranted): Acanthochromis polyacanthus(Bleeker), USNM 275349, 309487; Amblyglyphido-don aureus (Cuvier), USNM 338213; Amphiprion al-lardi Klausewitz, MCZ 4489 (Stiassny and Jensen's(1987) specimen), USNM 275417; A. melanopusBleeker, USNM 338150; Chromis atrilobata Gill,USNM 321208, C. cyanea (Poey) (USNM 318909),C. iomelas Jordan and Seale (USNM 338155; C. ter-natensis (Bleeker), USNM 338158; C. viridis (Cu-
vier), USNM 338160; Chrysiptera taupou (Jordanand Seale), USNM 338076; Dascyllus reticulatus(Richardson), USNM 338175; Lepidozygus tapeino-soma (Bleeker), USNM 97140, 265252, 275893,348198; Mecaenichthys immaculatus (Ogilby),USNM 215191; Microspathodon chrysurus (Cuvier),USNM 194045; Plectroglyphidodon dickii (Lienard),USNM 338219; Pomacentrus vaiuli Jordan and Se-
ale, USNM 338226; Stegastes fasciolatus (Ogilby),USNM 338222.
Description.LEI broadly on Ebl uncinate process ventroanter-iorly.LE2 finely tendinously on bony tip of dorsally ex-panded posterior edge of Eb2.LE3 finely tendinously on tip of Eb3 uncinate pro-cess medially.LE4 on dorsal edge of Eb4 lateral to uncinate pro-cess, joining raphe with OP dorsal to level of Eb4;LE4 joined ventrolaterally by LP, which does not joinOP.
?
Lacks raphe with OP. ? Tendinously on dor-solateral edge of Eb4, does not join raphe ventrallywith OP. (See also Discussion following Additionalremarks.)LP finely tendinously on Eb4, joining LE4 pos-terolaterally. (See also discussion following Addi-tional remarks.)LI1 by slender tendon mainly on Pb2 dorsoanter-iorly just below broad cartilaginous tip of processarticulating with IAC, tendon extending secondarilyonto CT binding adjacent anteriormost end of Pb3with Pb2.LI2 on Pb3 dorsolaterally ventral to medial end ofEb3.TD comprises TPb2a. TPb2 (see Discussion ofTPb2 following Additional remarks), TEb2, andTPb3. TPb2a attached to anterior surfaces of Pb2s;attached mid-posteriorly to CT that passes betweenPb3s. TPb2 and TEb2 each comprising a pair of mus-cles (TPb2 dorsal to TEb2) joined medially by broadarea of CT. TPb2 attaching anteriorly to dorsoanteriortip of Pb2. TEb2 of each side attaches to ventrolateraledge of CT connecting TPb2s, muscle extends lat-erally and attaches on Eb2 dorsally anterior to LE2insertion, joining medial end of Ad2. TPb3 on dor-soposterolateral end of Pb3 opposite medial end ofEb4. ? TEb2 may just fail to reach medial edge ofLE2 or just reaches point anterior to LE2. ? Similarto Dischistodus, but has TPb3-Eb3 instead of TPb3;attachment to Eb3 is to posteromedialmost tip.Remarks. See discussion in remarks following de-scription of TD in Parahollardia (Triacanthodidae)for discussion of similarity of knob-like process at-taching Pb3s to ventral surface of skull in that taxonand pomacentrids, and difference from that of labrids,embiotocids, and cichlids.CPb fibers relative fine, closely bound and ob-scured by covering CT of pharyngeal area; originat-ing from longitudinal SO fibers passing posteriorlyalong medial surfaces of Pb2 and Pb3, branchingposteriorly and extending along posterior surfaces ofPb3 and UP4 and attaching to posterolateral cornerof UP4; another fine strap of muscle attaching to pos-terior surface of Pb2 and extending laterally around
192 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pb3 and UP4, and attaching to posterolateral cornerof Pb4.
?
CPb absent. Absent also in Lepidozygustapeinosoma.Remarks. This muscle is very difficult to find, andmay not be present in most genera. In large speci-mens of large species, such as Plectroglyphidodondickii, it is relatively easy to find.OD3-4, OD3' origin on lateral surface of Pb3 dor-sal articulating facet (facet partially ventral to TPb2),insertion on medial edges of Eb3 and Eb4 uncinateprocesses; OD3' questionably identified here as thinlayer of fibers originating on Pb3 laterally immedi-ately ventral to OD3-4 origin and inserting dorsallyon medial end of Eb3. On only one side of one offour specimens, insertion extended dorsolaterally onEb3 to point about halfway between medial end ofAd3 and base of uncinate process.Remarks. We also found OD3' in Acanthochromispolyacanthus, Amphiprion allardi, and Lepidozygustapeinosoma. It probably occurs in its shortened, spe-cialized state in most or all pomacentrids. The muscleis difficult to find as it is thin, completely overlainby OD3-4, and easily damaged when cutting throughOD3-4 to expose the insertions of LI2 and TPb3.OP dorsally joining raphe with LE4, dorsoanter-iorly attached to Eb4 posteriorly beginning medial touncinate process and extending laterally close to lat-eral end of Eb4, divided into lateral and medial sec-tions by long slender tendon, which extends fromnear dorsal end of muscle to dorsodistal end of Cb5;tendon joined laterally by ventromedial end of Ad5.? Not continuous dorsally with LE4, comprisingbroad lateral and slender medial sections, which fuseventrally and attach to Cb5 dorsoanterodistally, therealso joining Ad5.Ad 1-3 each begin on dorsoanterior surface of re-spective Eb and extend broadly onto anterior surfaceof joint with respective Cbl, then continue as slenderGFM along most of remaining Cb anterior edge.Ad4 dorsally broadly on ventral surface of Eb4lateral to uncinate process, ventrally on Cb4 dor-soanteriorly medial to Eb4-Cb4 joint.Ad5 dorsally broadly on posterolateral surface ofCb4, ventrally broadly on posterolateral surface ofCb5 extending dorsally and joining tendinous exten-sion at ventrolateral end of OP.SOD absent.RDs well separated.Additional remarks. SCL free from Bb3. TV4 intwo sections, ventral section relatively thin, contin-uous across fused Cb5s ventrally; dorsal sectionthick, interrupted, attaching to lateral surfaces of Cb5keel. Pb4 absent, UP4 present. Eb4 levator processabsent (present, at least, in Lepidozygus and Amphi-prion). See Table 8 for distribution of ACs in po-macentrids.Discussion. One of Stiassny and Jensen's (1987:
282) main synapomorphies of the Labroidei, is thatthe dorsal articulating Pb3 facets are completely na-ked and directly ("bone to bone") contact a ventrallyrounded neurocranial apophysis on the base of theskull. They contrasted this condition with that of sev-eral families (Gerreidae, etc.) in which the centralportion of TPb2 replaced by CT covering Pb3s dor-
sally. They (1987:276) reported that pomacentridslack cranio-pharyngobranchialis 2 (= our TPb2; theirtransversus pharyngobranchialis 2 = our TPb2a),presumably based on the alcohol preserved materialthey listed (* denotes taxa we also examined): Abu-defduf trochelii (Gill), A. saxatilis (Linnaeus); Am-phiprion allardi*; Chromis atrilobata*, C. cyanea*;Microspathodon chrysurus*; Neopomacentrus sin-densis; Pomacentrus otophorus ( = Stegastes otopho-rus (Poey)), Pomacentrus* moluccensis Bleeker; Ste-gastes* acapulcoensis (Fowler). All the pomacentridswe examined have TPb2 (and TPb2a) similar to thatillustrated in Plate 160), with the central portion com-prising a broad band of CT covering the Pb3 facets.We thought, perhaps, that Stiassny and Jensen mighthave considered the reduced pomacentrid TPb2 aspart of TEb2, but other than the levator sling, theydid not illustrate the pomacentrid muscles, citing il-lustrations in Stiassny's (1980) Ph.D. dissertation,which is not readily available, and Kauffman andLiem (1982:figs. 2A, 2B). Stiassny kindly sent us acopy of her illustration of the muscles in Pomacen-trus, and it does not include TPb2, nor do Kaufmanand Liem's illustrations.Among the pomacentrids they examined, Stiassnyand Jensen (1987:284 and fig. 8c) reported that LE4is not continuous with OP in Chromis (based on C.atrilobata and C. cyanea), Neopomacentrus (basedon N. sindensis (Day)) and Amphiprion (based on A.allardi). LE4 is continuous with OP in our specimenof C. cyanea, but not in our specimens of C. atrilo-bata and C. amboinensis. Considering the configu-ration of LE4 and LP in their illustration (the twomuscles insert together very narrowly), Stiassny andJensen appear to have based their description only onC. atrilobata.Stiassny and Jensen (1987:290) reported that LP
"never contributes to the muscle sling" in pomacen-trids. We examined Amphiprion melanopus and A.allardi (Stiassny and Jensen's specimen of A. allardiwas in too poor condition to determine the state ofLE4 and OP). In contrast to Stiassny and Jensen, wefound that not only LE4, but also LP is broadly con-tinuous with OP in both species of Amphiprion, andthat LP also contributes to the sling in Plectrogly-phidodon dickii and Stegastes fasciolatus.Among the four specimens of Lepidozygus tapei-nosoma we examined, a sling may be absent, LE4and LP may be continuous with OP, or only one orthe other may be continuous with OP, but only a few
NUMBER 11 193
muscle strands of LP are involved when LP is con-tinuous with OP.The pomacentrids have not been analyzed cladis-tically, and we are unable to hypothesize the plesiom-orphic state for the muscle sling, which is importantfor corroborating inclusion of the Pomacentridae inthe Labroidei of Kaufman and Liem (1982) andStiassny and Jensen (1987). In any event, the po-macentrid sling, when present, differs considerablyfrom the sling in the other labroids in being relativelysimple. K.L. Tang is preparing a phylogeny of thepomacentrid genera, which was not available at thetime we were preparing our study. We have arbitrari-ly selected Dischistodus, in which the sling consistsof LE4 + OP, the most common state, and Amphi-prion melanopus (not illustrated), in which the slingcomprises LE4 (not released from Eb4), LP, and ORfor inclusion in the cladistic analysis. The absence ofa sling in Chromis and Lepidozygous, appears to bespecialized in those two genera, because the other,more common character states are either shown bysome species of Chromis, or other specimens of themonotypic Lepidozygous.
EMBIOTOCIDAEAmphistichus argenteus Agassiz, USNM 132403, 3specimens, ca. 100+ mm, of which one is clearedand stained, and one damaged.Plates 162.1, 162.2
Additional material. ? = Embiotoca lateralis Agas-
siz, USNM 340889, 86.8 mm.Plate 162.1
Additional material. Cymatogaster aggregata Gib-bons, USNM 340888; Hysterocarpus traskii Gib-bons, USNM 61189; Phanerodon atripes (Jordanand Gilbert), USNM 337459; P. fiircatus Girard,USNM 126843 & 337461; Rhacochihis vacca (Gi-
rard), USNM 340884, Zalembius rosaceus (Jordanand Gilbert), USNM 337463. Most of the infor-mation on these taxa is found only on Table 9.
Description.LEI broad based, on dorsalmost edge and surfaceof Ebl near medial end of element (see Additionalremarks for comment about presence of Ebl uncinateprocess).LE2 on mid-dorsoposterior bony edge of Eb2.LE3 dorsoanteriorly on Eb3 uncinate process, fus-ing anteroventrally with OD3-4.LE4 complex, massive, essentially free from Eb4,fused posterolaterally with LP dorsal to Eb4, joiningraphe with presumable middle section of OP.LP laterally on Eb4 posterolaterally, fused withLE4 ventroanteriorly, joining raphe ventrally withpresumable middle section of OP. ? LP more distinct,
on levator process dorsally, continuous posteroven-trally by only a few muscle strands with OP.Remarks. There is a strong, bony process thatbears the cartilage tip of the levator process in
?.The same process is present in Amphisticus, whichlacks the cartilage (hence, uncinate process absent).LI1 on dorsomedial surface of Pb2 and dorsoan-terior surface of Pb3. ? On Pb2 dorsally.LI2 inserts by long tendon on Pb3 dorsopostero-laterally ventroanterior to medial end of Eb3.TD comprises TEb2 and TPb3-Eb4. TEb2 a pairof muscles joined medially by CT to ventrolateralsurface of CT pad covering otherwise naked dorsalPb3 articulating facets, extending laterally on dorsalsurface of Eb2 anterior to LE2 insertion, joining ra-phe with medial end of Ad2. TPb3-Eb4 on Pb3 dor-soposterolaterally, a little medial to LI2 insertion,continuing posteriorly and attaching to medialmostedge of Eb4, thence continuing along posteromedialsurface of Eb4 and joining raphe with dorsomedialedge of a branch of OP (possibly the same as eitherOP3 or OP4 of cichlids). TPb3-Eb4 posteromedially,continuous mid-posteriorly with SOD. ? TEb2 doesnot join raphe with Ad2.CPb absent, but, unusually, SO, which is coveredventrally by pharyngeal roof CT, spreads anterolat-erally (Plate 162.2) and lines the ventral surfaces ofthe epibranchials (see M. Pb3-UP4).Remarks. Muscle questionably distinct from loosemix of SO + CT fibers that spreads over pharyngealroof, and is possibly homologous with one or all ofthe muscles identified as CPb in other putative la-broids and pholidichthyids, callionymoids, centro-geniids, lethrinids. pseudochromids, nemipterids, andsparids. Muscle fibers do not attach to the (reduced)Pb2s or pass around them anteriorly, as they do inthe other taxa.M. Pb3-UP4 ventral to RD, on Pb3 ventroposter-iorly and UP4 ventroanteriorly (also present in ? andCymatogaster, but not other genera examined).OD3-4 originating on lateral surface of Pb3 dorsalarticulating facet, inserting massively on dorsomedi-almost surface of Eb3 and anterior surface of unci-nate process (there joining LE3 insertion anteriorly)and medial edge of bony Eb4 uncinate process.OP complex, presumably comprising two or threesections (only two indicated here): posterior sectionjoining raphes dorsally with LE4 and LP, ventrallyon Cb5 distally, joining tendinous raphe laterallywith Ad5; anterior section broad, on Eb4 posteriorly,fusing posteriorly with anterior section ventral toEb4.Ad 1-3, each broad, on anterolateral surfaces, cov-ering joint, of respective Eb and Cb.Ad4 thin muscle sheet dorsally on ventrolateraledge of Eb4 and ventrally narrowly on Cb4 dorso-
194 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
laterally, obscured from view by OP portion attach-ing broadly to Eb4 posteriorly.Ad5 dorsally on tiny AC4 (not visible in illustra-tion), narrowly on Cb4 distal ly, and broadly on Eb4posterodistally; ventrally on Cb5 distal end, joiningtendinous raphe with OP posterior section. ? Slight,conformational separation of Ad5 from OP alongraphe.SOD present. ? Absent. See also Additional re-marks.RDs slightly separate.Additional remarks. SCL attached mid-dorsally bylong tendon to posteroventral end of Bb3. TV4 com-plex, but dorsally attached to Cb5, ventrally contin-uous but interrupted by raphes, possibly comprisingseparate dorsal and ventral sets of muscles. Pb4 ab-sent, UP4 present. Pbl cartilaginous. Pb2 edentate.The uncinate process of Eb 1 appears to have movedmedially and essentially become confluent with themedial end of the anterior arm of Eb 1 in all embioto-cids, hence, a synapomorphy for the family. Evidencefor this assumption is that Pbl articulates with therounded ventral portion of the cartilaginous medialend of Ebl and a ligament connects the dorsal, blade-like cartilaginous medial edge of Ebl to Pb2.Tiny ACs present as follows (Table 8): Amphisti-cus argenteus, AC4 present on one side of each oftwo specimens (of which one cleared and stained;third specimen damaged), Ad5 attached to it in onespecimen; Phanerodon atripes, AC4, Ad5 not at-tached to it; P. furcatus, AC1 and AC2 present orabsent; Zalembius rosaceus, AC4, Ad5 attached to it.SOD in Amphistichus is distinct and readily rec-ognizable. In both species of Phanerodon, however,SOD arises from SO dorsolaterally as a short, fine
strip of muscle that fuses with the ventrolateral surfaceof TEb4, and is easily overlooked. Stiassny and Jensen
( 1 987), who examined Phanerodon, but not Amphis-tichus, reported that all labroids, in which they in-cluded embiotocids, lack SOD. Presence of SOD isgenerally plesiomorphic for percomorphs and possiblyindicative that Amphistichus is one of the most ple-siomorphic genera in the family. The state of SOD inPhanerodon appears to be more specialized.Liem (1986) illustrated and described the gill-archmusculature of Embiotoca lateralis. Our observationsare in general agreement with his; however, Liem de-scribed LI1 as inserting only on Pb2, and LP as com-pletely separate from LE4. We examined Liem's ma-terial (MCZ 58890) and find that LI1 and LP accordwith the description of our specimen.
Labroidea
In order to facilitate discussion of the unquestion-ably monophyletic group comprising the Labridae,
Table 1 0.?Distribution of certain characters in genera of labro-idean fishes. Dash (-) = absent; P = present; C = continuous; D
= divided: F = fused; S = separate.
*x>uJia,HGenera en 04XIto gLabridaeAchoerodus - - - C FBodianus - P PA C FCheilinus - P PA C FCheilio - - A c FChoerodon P - A C FClepticus - P P C FCoris P - P c SDecodon - P A c FHalichoeres - - PA C SHologymnosus - - P C SLabroides P - P c SNotolabrus - - P c SPolylepion P P P c SPseudodax - 9 ? C FPseudolabrus - - P C SSemicossyphus - P A C sSuezichlhys - - A C sSymphodus - P A c FTautoga - P A C FTautoglabrus - P P C FXiphocheilus - P A C FOdacidaeOdax pullus - P P C FScaridaeLeptoscams - P A D FNicholsina - P A D FSparisoma - P A D F
Odacidae, and Scaridae, and distinguish them fromother families (Embiotocidae, Pomacentridae, Cich-lidae) included in the putatively monophyletic sub-order Labroidei, we use the superfamily name La-broidea. The Labridae are generally considered theleast specialized of the three families and, for themost part, we restrict our description and discussionto them.Additional material. We examined the gill-archmuscles of several taxa of Labroidea other than thelabrids described in the detailed accounts (data onthese taxa are included in Tables 8-10): Labridae:Bodianus mesothorax (Bloch & Schneider), USNM217854; Bodianus rufus (Linnaeus), USNM 320990;Cheilinus trilobatus Lacepede, USNM 224020 (Plate167B); Cheilio biennis (Forsskal), USNM 114811,332257; Choerodon graphicus (De Vis), USNM218548 (Plate 167A); Choerodon cyanodus (Rich-ardson), USNM 328226; Clepticus parrae (Bloch &Schneider), USNM 318548 (Plate 167C); Decodonpuellaris (Poey), USNM 185260; Halichoeres mar-garitaceus (Valenciennes), USNM 334551; Halicho-eres hortulanus (Lacepede), USNM 336931; Holo-gymnosus doliatus (Lacepede), USNM 218484; No-tolabrus celidotus (Bloch & Schneider), USNM
NUMBER 11 195
339231; Polylepion omentum Gomon, USNM215466; Pseudolabrus miles, USNM 339193; Pseu-dodax mohtccanus (Valenciennes), USNM 295262;Semicossyphus pulcher (Ayres), USNM 338987; Sue-zichthys aylingi Russell, USNM 339187; Symphodusroissali (Risso), USNM 198879; Tautoga onitis (Lin-naeus), USNM 118352. 163713; Tautogolabrus ad-spersus (Walbaum), USNM 1 18349; Xiphocheilus ty-pus Bleeker, USNM 260876; Odacidae: Odax pullus(Forster), USNM 339188; Scaridae: Sparisoma au-rofrenatum (Valenciennes), USNM 319033; Leptos-carus vaigiensis (Quoy & Gaimard), USNM 330153;Nicholsina denticulata (Evermann & Radcliffe),USNM 202270.Common and other characters. Many aspects of themusculature of the taxa are similar (e.g., all haveLE1-3. LI1, LI2. TPb3-Eb4, Adl-3; all but Bodianushave CPb). They differ mainly in the composition ofthe anterior TD muscles (Tables 9 and 10), whetherthe RDs are fused ventrally (underlain by a strap ofCT) at least to a point just posterior to their insertionon Pb3s, or are separate, whether TPb3 is continuousfrom one side of the gill arches to other (or whetherit is interrupted at the mid-line), and whether a sling(Stiassny and Jensen, 1987:283-284, fig. 8) is present(absent only in Labroides).An interrupted TPb3-Eb4 is a scarid synapomor-phy (hence, the problematic Pseudodax does not be-long in the scarid clade and possibly merits a family-group name equivalent to Labridae, Odacidae, andScaridae).One character, LEI insertion includes area medialto the Ebl uncinate process (usually entirely medial),is a synapomorphy of the Labroidea among all acan-thomorphs (we exclude the peculiar state in Colola-bis (Scomberesocidae, Plate 100) in which LEI ex-tends medially along a tendon from the tip of theuncinate process, but is restricted, nevertheless, to anarea well medial to the medial end of Ebl). Only inthe highly specialized labrid, Labroides, is the inser-tion on and lateral to the uncinate process, a condi-tion we consider specialized within the Labroidea. Aproblem concerning this character is provided by taxalacking an Ebl uncinate process. In most, if not all,of the problematic taxa, except possibly the labroidfamily Embiotocidae, the insertion is well removedlaterally from the medial end of Ebl and probablyindicates that the muscle insertion has not moved me-dially. The insertion in the embiotocids, which lackan uncinate process, is close to the medial end ofEbl, and is possibly indicative of a close relationshipbetween Labroidea and Embiotocidae.Choerodon (Plate 167A) is unique in that LEIcomprises two separate muscles (LEI, LEI'), onewhich inserts entirely medial to the uncinate process,and the other, which inserts just ventrolateral to thetip of the uncinate process. Such a condition could
develop from the typical labroidean condition by lossof the central portion of the typical LEI. Loss of themedial portion would result in a condition similar tothat of Labroides.LE2'. We consider the distinction between LE2'and LE2 to be problematic. LE2' only occurs in thepresence of LE2, and is possibly an artefact of dis-section. Ebl and Eb2 are closely juxtaposed in la-brids. In most labrids, a tendon extends along LE2from about mid-length to insertion on a bony processon the posterior margin of Eb2. At this point the ten-don continues as a ligament attaching to a process onthe anteroventromedial margin of Eb3. In many la-brids. it appears that the LE2 comprises two separateelements, one (LE2) inserting musculously and theother tendinously on Eb2. In attempting to decidewhether one or two muscles are involved, it is nec-essary to force apart Eb2 and Eb3. In doing this, thequestionable muscle sometimes remains entirely onEb2, sometimes it appears to divide, but remain onEb2, sometimes it appears to completely separate andbe associated only with Eb3. In Tautoga and Pseu-dodax. LE2 is clearly restricted to Eb2. with no in-dication of possible division. In Choerodon, Halicho-eres, and Notolabrus, LE2' appears to be present. Inthe other genera, the situation is questionable.Transversus dorsalis. The names we apply to theanterior TD muscles should be considered highly ten-tative. As much as possible, we identify these mus-cles with those commonly found among acantho-morphs.One example of how a pair of muscles can varyis exemplified by Coris, Choerodon, and Xiphochei-lus. The lateral attachments of TPb2 and TPb2a inCoris (Plate 166) are to the anterolateral tip of Pb2,the most common attachments of these two musclesin labrids. In Choerodon (Plate 167A), TPb2 appearsto have shifted its attachment to the tip of the Eb3uncinate process, and the anterior portion of TPb2aappears to have an anterior branch that also attachesto the process. In Xiphocheilus (not illustrated), it ap-pears that the condition in Choerodon has been car-ried one step further, and the TPb2a portion has beenlost. We have arbitrarily indicated the muscles inChoerodon and Xiphocheilus as TPb2 and TPb2a inTable 9 and on Plate 167A. A study of the dorsalgill-arch musculature of labroideans will probablyuncover informative characters for intra- and inter-familial relationships.Transversus epibranchialis 1. Unaware, perhaps,that some labrids lacked TEbl, Stiassny (1980:248-249) proposed that the presence of TEbl is a syna-pomorphy of the Labridae. A cladistic analysis of theLabridae will be required in order to determinewhether the absence of TEbl in Labridae is a sec-ondary loss. Pending such an analysis, we treat thelack of TEbl in labrids as the plesiomorphic state.
196 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pharyngobranchial 1. Our identification of Pbl inlabrids, the only labroideans that have an element thatmight be interpreted as Pbl, is problematic. All la-brids we examined, except Labroides, have such anelement, which is always cartilaginous. Pbl is un-ambiguously absent in Labroides, which is clearly aspecialized state for an acanthomorph. In the otherlabrids, there is a range of variation in morphologyof the cartilage at the anteromedial end of Ebl: anunremarkable cartilaginous end joining a separate,curved or straight rod-like cartilage (e.g., Achoero-dus, Plate 163.1 A; Polylepion, Plate 165A), whichmight be interpreted as Pbl; a greatly expanded, me-dially flattened cartilage cap giving rise anteriorly toa continuous, posteriorly directed, curved cartilagerod (e.g., Coris, Halichoeres), which might be inter-preted either as Pb 1 fused to the end of Eb 1 or as ade novo shape of the anteromedial end of Ebl; agreatly reduced autogenous filament of cartilage,which may be present on one side and absent on theother, and which might also be interpreted as Pbl.In the two specimens of Coris (USNM 337450),the cartilaginous medial end of Eb 1 is basally cuboidand medially plate-like with the anterior marginforming a worm-like, dorsally extending process. Ev-idence that the process originates as part of the plate-like portion is indicated by the conditions in USNM92292. On the left side of this specimen the plate-like portion is almost completely demarcated fromthe remainder of the cartilaginous end of Ebl by aline of unstained tissue, as precedes the budding offof cartilaginous ACs. On the right side, there are twoisolated non-staining areas in the same relative po-sition as the unstained line on the left side. The un-stained areas appear to mark the budding off of theputative Pbl. The situation in labrids is unusual,however, in that developmentally, Pbl appears toform separately from the medial end of Ebl in theearliest ontogenetic stages of perciforms (e.g., Pott-hoff et al., 1987, pomacentrid), if not all acantho-morphs.Like the presence or absence of TEb 1 , the state ofthe medial end of Ebl may offer clues to the intra-familial relationships of the labrids.Sling. The composition of the sling is also com-plex, involving LP, LE4, OP, and Ad5 in apparentlydifferent ways. Some of the variation in the sling canbe gleaned from the illustrations.LABRIDAERemarks. Yamaoka (1978) describes seven origintypes for LP, which he associates with feeding types.We did not investigate LP origins.Achoerodus viridis (Steindachner), AMS 1.7019-006,88.9 mm; USNM 218488, 56.8 mm.Plates 163.1, 163.2
Description.LEI on Ebl medially beginning laterally ventralto tip of uncinate process and extending to dorso-medial end of Ebl.LE2 on dorsoposterior edge of Eb2 at mid-length.LE3 tendinously and musculously on all bony Eb3and cartilage-tipped Eb4 uncinate processes; muscletwists on itself as it descends from origin.Remarks. A cartilage tipped Eb3 uncinate processis absent in all labrids. The tip of the Eb4 uncinateprocess (usually not visible as it is often obscured byLE3 insertion) may be cartilaginous or all bony vary-ing with the taxon and possibly with growth (begin-ning as cartilage and becoming bony).LE4 complex, essentially free from Eb4 (exceptfor "point" attachment to Eb4, where LE4 and LPfuse), fused with LP along broad, ventrally extendingtendon that attaches to dorsoposterior edge of distalend of Cb5, presumably incorporating middle sectionof OP ventral to level of Eb4.LP complex, massive, on dorsal and ventral sur-faces of Eb4 (bone appears tilted up so that surfacescould be interpreted as anterior and posterior), ex-tends ventrally fusing with presumed lateral portionof OP.LI1 on Pb3 articulating surface ventroanterolater-
ally.LI2 on Pb3 posterolaterally, ventral to Eb3 medi-
ally.TD comprises TPb2-Pb2a and TPb3-Eb4. TPb2-Pb2a originates posteriorly from anteriorly extendingfibers from transverse SO muscle layer passing be-tween naked dorsal Pb3 articulating facets andspreading anteriorly and attaching on dorsoanteriorsurfaces of Pb2s. TPb3-Eb4 attaches to posterolater-almost surface of Pb3 and ventroposterolmedialmostedge of Eb4.Remarks. Homology of anterior TD muscles of la-brids, odacids, and scarids with those of other acan-fhomorphs is problematic.CPb extends from posterolateral corner of Pb3 an-teriorly around Pb2 and Pb3 ventral to TPb2-Pb2a toposterolateral corner of opposite Pb3.OD3-4 origin on lateral surface of dorsal Pb3 ar-ticulating facet, insertion on medial edge of bony Eb3uncinate process and mid-anterior edge of Eb3 ven-tral to process, and on medial edge of cartilage-tippedEb4 uncinate process.OP complex, possibly consisting of three sections;questionable middle section participating in sling asmuscle ventral to raphe joining LE4 and LP; lateralsection fusing with LP ventrally and Ad5 posteriorly;medial section in two parts: dorsal part on posteriorsurface of Eb4 ventral to OD3?4, becoming tendi-nous ventrally and joining broad tendon extendingfrom OP middle section shortly ventral to broad ten-don's giving rise to long posteriorly extending liga-
NUMBER 11 197
ment; ventral part musculously on posterodistal sur-face of Cb5, fused with ventral end of lateral OPsection and/or posterior surface of Ad5.Remarks. The OP sections of labrids are possiblyhomologous with the three OP sections identified inembiotocids and three of the four sections identifiedin cichlids, but await confirmation from cladistic andembryological studies.Adl spans anterior surface of Ebl-Cbl joint.Remarks. Slender GFM1 extends ventrolaterallyacross Adl surface from dorsomedial origin of Adl;others may have been present on arches 2 and 3, butwere not noted during dissection. GFMs were notspecifically checked for during dissection of labrids.Ad2 with two portions joined to raphe; dorsal por-tion extends from dorsal surface of Eb2 just anteriorto LE2 insertion and attaches to posterior surface ofEbl ventral to uncinate process; ventral portionspreads ventroanteriorly from raphe and attaches toCb2 anterior surface just ventral to Eb2-Cb2 joint.Ad3 dorsally on almost entire anterior edge of Eb3ventroanterior to uncinate process, tendinously at-tached (not illustrated) to Eb2 at and continuous withLE2 insertion, extending ventrolaterally across anter-odistal surface of Cb3 and attaching just ventral tocartilaginous distal end.Ad4 on Eb4 ventrolaterally and on Cb4 dorsallyanterior to Eb4-Cb4 joint.Ad5 most recognizable dorsolaterally, attachingdorsally broadly, tendinously to Eb4 and Cb4 pos-terodistally, and ventrally to Cb5 dorsodistally, me-dial surface fused with OP complex.SOD absent.RDs fused, dividing just before attaching to Pb3s.Additional remarks. SCL attached mid-dorsally byCT to posterior surface of posteroventrally extendingcartilaginous tip of Bb3. TV4 mostly split, attachedto Cb5s laterally except for thin, continuous musclestrap passing across Cb5s ventrally. Pb4 and UP4absent. Pbl cartilaginous. Eb4 levator process absent.See Table 8 for distribution of ACs in labrids.
Symphodus roissali (Risso), USNM 198879, 2 spec-imens, 50.0-55.8 mm.Plates 164.1, 164.2
Description (see also remarks following various mus-cle descriptions under Achoerodus).LEI broadly on Ebl beginning at base of uncinateprocess and extending medially to medialmost bonyedge.LE2 narrowly, tendinously on mid-dorsoposterioredge of Eb2.LE3 tendinously on tip of Eb4 [sic] uncinate pro-cess (Eb3 and Eb4 tightly joined, but muscle hasshifted its usual insertion from Eb3 to Eb4).
Remarks. Eb4 uncinate process with minute car-tilage tip at bottom of minute, shallow bony depres-sion, as if bone was about to grow over tip.LE4 essentially free from Eb4, joins LP dorsal tolevel of Eb4, ventrally continuous with presumedmiddle section of OP (raphe not illustrated).LP massive, on Eb4 dorsolaterally, posteroventral-ly joining raphe (not illustrated) with presumed lat-eral section of OP, which continues ventrally as broadtendon that attaches to Ad5 and inserts on Cb5.LI1 on anterolateral surface of Pb3 just posteriorto dorsoanterior tip of Pb2.LI2 tendinously on posterolateral surface of Pb3.TD comprises TEbl, TPb2-Pb2a-Pb3, and TPb3-Eb4. TEbl on mid-anterior edge of Ebls with finecrossing muscle filaments mid-posteriorly attachingto cartilaginous Pbls and medial edge of Ebls. TPb2-Pb2a-Pb3 originating posteriorly as longitudinal fi-bers extending anteriorly from transverse SO layer,passing between Pb3s, dividing and passing over andattaching to Pb2s and around each Pb3 and attachingto it; dorsal divisions continuous ventrally with un-interrupted portion, which attaches to Pb2s laterally.TPb3-Eb4 attaches to posterior surface of Pb3 dorsalarticulating facets and to Eb4s posteromedially.CPb semicircular band passing anteriorly around,but mostly separated by epithelial tissue from, Pb2and Pb3 ventral to uninterrupted portion of TPb2-Pb2a-Pb3, attaching to posterolateral corner of Pb3,and continuing posteromedially and meshing com-plexly with SO.M. Pb2-Eb2 on Pb2 ventrolateral edge, passing be-tween TPb2-Pb2a-Pb3 and CPb and attaching tosmall bony process on mid-anterior edge of Eb2.Remarks. This muscle is possibly a separate ex-tension of CPb. In many other species of labrids, CPbhas a fine tendinous attachment to Eb2 ventromedi-ally and Pb2 ventrolaterally as it extends posteriorly,or muscle fibers are continuous with CPb.OD3 absent.OD4 on lateral surface of Pb3 articulating facetand Eb4 uncinate process (see remarks followingLE3).OP presumably in three sections: lateral sectionjoining raphe dorsally with LP and continuing ven-trally as broad tendon, which joins Cb5 dorsoposter-iorly, and is joined by Ad5 anteriorly; tendon contin-ues dorsally. joining OP middle section; medial sec-tion on Eb4 posteriorly and Cb5 dorsally medial tobroad tendon, medially continuous with SO.Adl broadly on Ebl and cartilaginous Pbl dorsallyand Cbl anteriorly well ventral to distal end.Ad2 on Eb2 dorsomedially and Cb2 anteriorly wellventral to distal end.Ad3 on Eb3 dorsomedially ventral to OD4 andCb3 anteriorly well ventral to distal end.
198 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ad4 dorsally on ventrolateral surface of Eb4, ven-trally on Cb4 dorsolaterally medial to Eb4-Cb4 joint.Ad5 ventrally on dorsoanterodistal surface of Cb5,dorsally broadly on posterodistal ends of Eb4 andCb4, medially joining tendon from OP middle sec-tion.SOD absent.RDs fused, separating just before to attaching toPb3s.Additional remarks. SCL present (cartilaginousposterior end of Bb3 recurved ventrally). TV4 com-pletely divided medially, attaching to Cb5 anterolat-erally on each side. Pb4 and UP4 absent. Pbl carti-laginous. Eb4 levator process absent. See Table 8 fordistribution of ACs in labrids.
Polylepion cruentum Gomon, USNM 215466. 82.7mm. Plate 165
Additional material.
?
= Coris julis (Linnaeus),USNM 337450, 2 specimens: 135-164 mm;USNM 92292, 105 mm.Plate 166
Description.LEI on dorsomedialmost edge of Ebl medial tocartilage-tipped uncinate process. ? Broadly on Eblbeginning near dorsoposteromedialmost bony surfaceand extending to base of uncinate process.LE2 on bony process formed by mid-dorsoposter-ioredge of Eb2, joining LE2' insertion. LE2' (Seediscussion of LE2' in Labroidea section) tendinouslyon bony process at mid-dorsoposterior edge of Eb2together with LE2, ligament continues posteriorlyfrom insertion and attaches to bony process on ven-troanteromedialmost edge of Eb3.
? On bony processat ventroanteromedialmost edge of Eb3, ligament at-taches process to Eb2 process supporting LE2.LE3 tendinously on tip of all bony Eb3 uncinateprocess.
?
On tips of tightly joined bony Eb3 andcartilage-tipped Eb4 uncinate processes.LE4 massive, essentially free from Eb4, fusingwith LP dorsal to level of Eb4 and presumable mid-dle portion of OP, which continues ventromedially aslong tendon and attaches to Cb5, and fuses laterallywith Ad5.LP massive, partly on dorsoanterior surface ofEb4, but mainly on posterolateral surface, where it isjoined by LE4; joins raphe ventrally with presumablelateral section of OP.LI1 on Pb3 dorsoanterolaterally, anteroventral todorsal articulating facet.LI2 on Pb3 dorsolaterally anterior to joint withEb3 medial end.TD complex, comprising TEbl, TEb2, TPb2,TPb2a, and TPb3-Eb4. TEbl on dorsoanteromedial
surfaces of Ebls, anterolaterally joining raphe withAdl, joined posteriorly by complex muscle strandsto CPb; some strands continue posteriorly as bilateralpair of longitudinal SO bands that pass posteriorlybetween Pb3s and give rise dorsally to two bilateralpairs of small, slender muscles: each member of an-terior pair, TPb2, attaches to anterior tip of Pb2 to-gether with TPb2a; each member of posterior pair,TEb2, attaches to Eb2 dorsal surface near medial at-tachment of Ad2. TPb2a attaches broadly to medialedge of slender anterior Pb2 process. TPb3-Eb4 at-taches to posterior surface of Pb3 articulating facetsand to posteromedial surface of Eb4. ? TEbl andassociated longitudinal strands of SO absent. TEb2joins raphe laterally with Ad2.CPb with broad anterolateral extension attachingto anterior edge of Eb2 ventral to Ad2; muscle oth-erwise semicircular, with median anterior raphe,passing posterolaterally around Pb2 and Pb3 on eachside, thinning and attaching to posterolateral cornerof Pb3, thence spreading posteriorly and continuingas SO. ? No median raphe; does not continue pos-teriorly beyond attachment to Pb3.OD3?4 origin on anterolateral edge of Pb3 dorsalarticulating facet, continuing posteriorly on Pb3 ven-tral to facet, insertion on medial edges of bony Eb3and Eb4 uncinate processes. ? Eb4 uncinate processwith cartilage tip.OP presumably comprising three sections; lateralsection dorsally joining raphe with LP ventrally andmedially joining long tendon that joins middle OPsection dorsally, ventrally fusing indistinguishablywith Ad5 posteriorly; medial section on distal end ofCb4, becoming tendinous dorsally on one side (re-maining musculous on other) and attaching to Eb4posteriorly.
?
Medial OP section absent.Adl broadly on Ebl dorsoanteriorly beginning atraphe with TEbl (which is absent in
?), extendingventrolaterally medial to Ebl-Cbl joint, and attach-ing to Cbl anteriorly ventral to joint.Ad2 broadly on most of bony length of Eb2, ex-tending ventrolaterally just medial to Eb2-Cb2 jointand attaching to Cb2 anteriorly just ventral to joint.Ad3 broadly on most of bony length of Eb3, ex-tending ventrolaterally just medial to Eb3-Cb3 jointand attaching to Cb3 anteriorly just ventral to joint.Ad4 broadly on Eb4 ventrolateral edge and Cb4dorsally medial to Eb4-Cb4 joint.Ad5 ventrally on Cb5 dorsolaterally, dorsallybroadly on Eb4 and Cb4 laterally, fused laterally withOP.SOD absent.RDs fused ventrally just posterior to attaching toPb3s.
?
RDs separate well posterior to Pb3s.Additional remarks. SCL attached mid-dorsally toventrally extending cartilaginous posterior end ofBb3. TV4 completely divided, attaching to Cb5 an-
NUMBER 1 1 199
terolaterally. Pb4 and UP4 absent. Eb4 levator pro-cess absent.
?
SCL present; cartilaginous posteriorend of Bb3 extends ventrally; thin band of TV4 con-tinuous across Cb5 ventral to anterolateral attachedportions. See Table 8 for distribution of ACs in la-brids.
Labroides dimidiatus (Valenciennes), ? = USNM309376, 70.7 mm;
?
= USNM 205283, ca. 65 mmSL;
?
= USNM 363249, 80.3 mm SL; ? =USNM 369939. 69.5 mm.Plate 168
Description (unless noted otherwise, description ap-plies to all four specimens).LEI originates tendinously, inserts on bony dor-soanterior surface of Ebl uncinate process.LE2 on tip of expanded mid-posterior edge of Eb2,probably joining LE2' ventrally (see remarks follow-ing LE2').LE2' (See discussion of LE2' in Labroidea sec-tion) tendinously on bony process ventral to medialend of Eb3 and questionably on mid-posterior edgeof Eb2 at or with LE2 insertion.Remarks. Eb2 is tightly joined to Eb3 posterior toLE2 insertion, and it is difficult to decide whetherLE2' actually joins LE2. Eb2 and Eb3 are artificiallyseparated in Plate 168A.LE3 tendinously on tip of all bony Eb3 uncinateprocess.LE4 ribbon-like, on Eb4 dorsolaterally, varying asfollows: ? inserting musculously at and anterior toLP on one side and at and posterior to LP on theother;
?
fusing with LP (forming a Y) and insertingmusculously together (both sides);
? inserting ten-dinously at and posterior to LP (both sides); ? ex-tending tendinously across and joining ventromedialsurface of LP and inserting on bony distal end of Eb4dorsally together with ventroposteromedial surface ofLP.LP ribbon-like, on Eb4 dorsolaterally, variable (seeLE4), insertion continuous posteroventrally with CTto which PP (not illustrated) also joins. ? Musclefibers are continuous ventromedially with OP on leftside but not on right side.LI1 slender, on Pb3 dorsoanteriorly near articula-tion with Pb2 dorsal process.LI2 on Pb3 dorsolaterally ventral to dorsal articu-lating facet.TD comprises TPb2-Pb2a. TPb3, TEb2, and TPb3-Eb4. Anterior three muscles originate posteriorlyfrom transverse SO fibers that pass anteriorly be-tween Pb3s and divide on exiting from between Pb3s.TPb2-Pb2a fuses ventrally with uninterrupted trans-verse muscle portion (dorsal to CPb) and attaches toPb2s anterolaterally. TEb2 extends laterally posterior
to ascending process of Pb2 and attaches on Eb2 dor-soanteriorly. meeting anteromedial margin of Ad2.TPb3 attaches to Pb3s anterolaterally dorsal to TPb2-Pb2a and along medial edge of Eb2. TPb3-Eb4 at-taches medially to posterior surfaces of Pb3 articu-lating facets and to posteromedial half of Eb4.CPb, beginning posteriorly, attaches on posterolat-eral corner of Pb3, extends anterolateral to Pb3 at-taching minutely to Eb2s ventroanteriorly, and con-tinues anteromedially ventral to TPb2-Pb2a to op-posite side.Remarks. It was unclear if the Pb2a portion ofTPb2-Pb2a was present in
?.OD3 origin on lateral surface of Pb3 articulatingfacet, insertion on anterior surface of all bony Eb3uncinate process.OD4 origin on lateral surface of Pb3 articulatingfacet posterior to OD3, insertion beginning on dorsalsurface of cartilage-tipped Eb4 uncinate process andextending laterally.OP on Eb4 ventrolaterally and Cb5 dorsomedial toAd5.Ad 1-3 similar, muscle on most of dorsoanteriorsurface of respective Eb, extending anteriorly overEb-Cb joint and attaching along anterior surface ofCb.Ad4 (occluded in posterior view) on ventrolateraledge of Eb4 and dorsolateral edge of Cb4 medial toEb4-Cb4 joint.Ad5 dorsally on cartilaginous distal end of Cb4continuing as CT with which LP and PP are alsocontinuous; ventrally on Cb5 dorsolaterally.SOD absent.RDs well separated.Additional remarks. SCL present. Except for thin,ventral, continuous section of fibers, TV4 attached toCb5 anterolaterally. Pbl, Pb4, and UP4 absent. SeeTable 8 for distribution of ACs in labrids.Labroides. Labroides is the most distinctive of thelabroidean genera we examined. It alone lacks a slingand has the LEI insertion on and lateral to the Ebluncinate process. It also lacks TEbl and has separateRDs. Although, all these characters are plesiomorph-ic for acanthomorphs, we believe all, except possiblythe absence of TEbl, will prove to be apomorphic atvarious levels within the Labroidea. The continuousportion of LP with OP on one side of one of fourspecimens, is probably anomalous and, if our inter-pretation is correct, not homologous with the LE4-OP sling of other labrids.
PholidichthyoideiThe interrelationships of the Pholidichthyidae re-main unresolved. The family shares specializationswith a diverse group of fishes. In recent times, it hasbeen questionably or provisionally associated with
200 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
the blennioids (Springer and Freihofer, 1976:40), thelabroids (Stiassny and Jensen, 1987), and the trachi-noids (Nelson, 1994:397).PHOLIDICHTHYIDAE
Pholidichthys leucotaenia Bleeker, CAS 32408, onevery large (illustrated) and one 230 mm SL;USNM 289924 and uncataloged, two juveniles.Plate 169
Additional material. ? = Pholidichthys anguisSpringer and Larson, USNM 337860 and NTMSI 3529-001, both juveniles.
Description.LEI inserts by slender tendon on Ebl anterior tobase of uncinate process; ligament (not shown) joinsposterior edge of Ebl to anterior edge of Eb2.LE2 inserts on tip of bony Eb2 process postero-lateral to uncinate process; ligament (not shown)joins process to anterior edge of Eb3. ? Process ab-sent.Remarks. A low, cartilage-tipped Eb2 uncinateprocess occurs in some specimens of P. leucotaenia,but not others. In some specimens the cartilaginousmedialmost end of Eb2 is more anteriorly directedand overrides the dorsal surface of Pb2 (see Springerand Freihofer, 1976:fig. 7, left side of fish), but whenarticulating with Pb3 is attached to bony surface.Among ctenosquamates, a cartilage-tipped uncinateprocess on Eb2 occurs elsewhere only in Hemiram-phus, (Hemiramphidae), in which it does not articu-late with another skeletal element.LE3 inserts on Eb3 uncinate process.Remarks. Eb3 uncinate process is closely boundby CT to anterior edge of Eb4, which lacks an un-cinate process. Because of the tight joint, it is pos-sible to interpret LE3 as inserting on both Eb3 andEb4, although the attachment seems to favor Eb3.Except for the Lampridae and Veliferidae, LE4 neverinserts on the Eb4 uncinate process in acantho-morphs.LE4 absent.Remarks. Pholidichthys, the gasterosteid Spina-chia, and the echeneids are apparently the only acan-thomorphs that lack LE4.LP broadly on most of lateral half of dorsoposter-ior surface of Eb4, insertion posteriorly parallelingand closely approximating OP attachment on Eb4;muscle extends posteromedially towards its origin onthe basioccipital, which is well removed from theother levators, which originate together on the pro-
otic.LI1 narrowly on anterolateral edge of Pb3 just pos-terior to anteriormost tip of Pb3.LI2 on posterolateral edge of Pb3 somewhat me-dial to medial end of Eb3.
TD comprises TPb2, TPb2a, TEb2, and TPb3-Eb4.CT covers entire dorsal surface of anterior TD com-ponents. TPb2 in two essentially separate parts, pos-sibly the result of LIl's passing between parts; an-terior part bilobed with mid-longitudinal raphe sep-arating lobes and attaching dorsally to CT cover;muscle attaches to anterolateral cartilaginous tip ofedentate Pb2; muscle forms shallow, posteriorlyopening pocket into which anterior fourth of Pb2 in-
serts. Posterior portion of TPb2 arises medially frombroad CT band extending across dorsally naked Pb3articulating facets; muscle extends anterolaterally andinserts on dorsolateral edge of Pb2. TEb2 very re-duced, may be present bi- or unilaterally or totallyabsent (absent in the two juveniles and in ?); musclearises from CT at posterolateral end of posterior sec-tion of TPb2 extends short distance and attaches toposteromedialmost edge of Eb2. TPb2a with mid-longitudinal raphe, smaller than and mostly ventro-posterior to TPb2, attaches to ventroanterolateraledge of Pb2. covering most of Pb2 ventral surfaceextending posterior to TPb2 pocket (N.B., althoughmuscle is posterior to TPb2, it appears that Pb2 hasrotated dorsoposteriorly in effecting its more lateralattachment with Eb2; without rotation, the musclewould be on the anterior surface of Pb2, similar tothat of afherinomorphs and the labrid Achoerodus).TPb3-Eb4 attaches to dorsomedian surface of Pb3,continuing laterally past attachment to Pb3 and at-taching on expanded posteromedial surface of Eb4.OD3-4 originates broadly from dorsoposterior sur-face of Pb3 and inserts on both Eb3 and Eb4 at jointformed with Eb3 uncinate process, and along Eb4surface posteromedial to this joint.OP origin on posterior surface of Eb4 along linejust below insertion line of LP (q.v.), insertion ondistal end of Cb5 dorsally.CPb complex, apparently derived from SO longi-tudinal fibers with which it is continuous posteriorly;mainly comprises anterior cylindrical portion (no an-teromedian raphe) that curves around anterior andlateral margins of both Pb3s. Cylindrical portion fi-bers almost vanish at posterolateral corner of eachPb3, but, on each side, a few fibers curve mediallyand become confluent with broad, strong group offibers that attach posteromedially to Pb3. The latterfibers curve medially, passing anteriorly between thePb3s and ventral to dorsal fibers that are continuousposteriorly with the SO longitudinal fibers. The an-teriorly passing fibers branch right and left anteriorlyand continue on their respective side as slips of mus-cle ventral to the ventroanterior surface of Pb3. Themuscle slips become confluent with the anterior innermargin of the cylindrical portion of CPb.Ad 1-3 present, each attaches along anterolateralsurface of its respective Eb and anterodistal surfaceof its respective Cb.
NUMBER 1
1
201
Ad4 (not illustrated) on ventroanterior surface ofEb4 and dorsal surface of Cb4 just medial to innerangle formed by Eb4-Cb4 joint; not visible in dorsalview, obscured by OP in posterior view.Ad5 vertically elongate, on posterodistal ends ofCb4 and Cb5; in small specimens of both speciesAd5 not clearly separable from OP and a few fibersof Ad5 attach to distal end of Eb4.SOD absent.RDs moderately separated, vertically short, eachconsisting of a partially round (in vertical cross sec-tion) anterior section that is continuous posteriorlywith a horizontally longer, obliquely posteriorly di-rected section; band-like tendon incorporated as ven-tral surface of each RD, which inserts on medial sur-face of Pb3.Additional remarks. SCL fine, free from posteriorend of Bb3, which is not elongate. TV4 bipartite,attaches anteriorly on each side of median ventralkeel of Cb5. Pb4 and UP4 absent. Pb2 edentate. IACabsent. Eb4 levator process absent.
AcanthuroideiLUVARIDAELuvarus imperialis Rafinesque, MCZ 55003, 156mm. Plate 170
Description.LEI short, on cartilaginous tip of Ebl uncinateprocess.LE2 dorsally on posteromedial edge of Eb2.LE3 tendinously on joined cartilaginous tips ofEb3 and Eb4 uncinate processes, attachment more onEb3 than Eb4; additionally, a long slender ligamentattaches cartilaginous tip of Eb3 uncinate process tocranium.LE4 mostly destroyed, along with LP, in dissec-tion; remainder on Eb4 dorsoposterior edge distal touncinate process, connected by raphe with dorsolat-eral edge of OP, such that release of raphe from Eb4produces a "sling"; attached to, but easily separatedfrom, medial surface of broad membrane (not illus-trated) that passes lateral to other levators and ex-tends posteriorly and includes LP and PP (see fol-lowing remarks).Remarks. Winterbottom (1993a:29) described LE4and LP in acanthuroids:
"In all the acanthuroids examined, the fourth levator externusnot only inserts on the dorsal rim of epibranchial 4, but contin-ues posterior to this into the dorsomedial wall of the branchialchamber (Fig. 11). In siganids and Luvarus, a levator posteriorarises from the region of the prootic/pterotic junction and joinsthe posterior fibers of levator externus IV in the wall of thebranchial chamber (Fig. 11). Posterolaterally, the protractor pec-toralis arises from the ventrolateral tip of the pterotic and passesventrally to the posterior wall of the branchial chamber and the
anterolateral face of the cleithrum. All of these muscles arerepresented by thin sheets of muscle fibers. In Zanclus, the pro-tractor pectoralis is joined medially by a fan-shaped sheet offibers from the ventrolateral prootic (= levator posterior?) ....In acanthurids (Fig. 12). the fibers to the medial wall of thebranchial chamber are separated by connective tissue from thoseof the protractor pectoralis. Occasionally, some fibers are pre-sent in this connective tissue in a position where one mightexpect a levator posterior to be (e.g., Acanthurus triostegus),but this is not the case in the majority of the acanthurids dis-sected."LP destroyed during dissection. See remarks fol-lowing LE4 above.LI Ion Pb2 dorsoanteriorly.LI2 on Pb3 cartilaginous posterior end dorsoanter-iorly, opposite anteromedial end of Eb3.TD comprises TEb2, TPb3, and TEb4. TEb2, inthree parts: one in middle and one on each side linkedby thick CT pad; middle part small, triangular; lateralparts strap-like, medially joining lateral edges of CTpad and laterally extending onto Eb2 dorsally lateralto LE2 insertion. TPb3 joins medial margins of Pb3sventral to OD3s. TEb4 a slender strap joining Eb4sjust ventromedial to uncinate processes.Remarks. Winterbottom (1993a:30) stated that Lu-varus is unique among acanthuroids in lacking TD4(= our TEb4), and possessing TD5. TEb4 in Luvarusis very similar to TEb4 in Acanthurus. We are un-certain what he meant by TD5, but this is possiblythe unusually anterior SOD we recognize in Luvarus.Although a fine lateralmost SOD muscle filamentjoins OP medially, SOD can be traced to Cb5 as itsventromedial fibers fuse with OP; however, in Acan-thurus, we also find a thin normally situated SODthat can be traced to Cb5.OD3 origin broadly on bony dorsal surface of Pb3ventral to TEb2, insertion on Eb3 somewhat medialto distal end continuing well up on uncinate process.OD4 absent.OP comprises two more-or-less distinct sections,variably almost completely separated from each otherand from SOD; medial section dorsally on Pb4 pos-teriorly, lateral section on Eb4 dorsoposteriorly inarea ventral to uncinate process, joining raphe withLE4 insertion (see OP description and remarks); bothOP sections on Cb5 near posterior end.Remarks. It is unusual for OP or SO to attach toPb4. It is equally possible to interpret the medial OPsection as SO fibers that have shifted from Eb4 toPb4. SO attaches to Pb4 in Polydactylus but not inFilimanus (both Polynemidae).Ad 1-3 absent.Ad4 on Eb4 posterolaterally and Cb4 narrowlyposteriorly anterior to Ad5 attachment.Ad5 very short, transverse, on Cb4 dorsoposter-iorly and Cb5 dorsoposterodistally.RDs separated by narrow space.SOD dorsally anterior to, and free from, TEb4.
202 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Remarks. SOD in other fishes is posterior to allTD components. See also remarks following TDabove.Additional remarks. SCL attached mid-dorsally toposteroventrally extending cartilaginous end of Bb3.TV4 free from Cb5s. Eb4 levator process absent. Pb4and UP4 present. EPHIPPIDAEChaetodipterus faber (Broussonet), USNM 289421,74.5 mm; USNM 159268, 85.4 mm; USNM118501, 61.3 mm. Plate 171
Description.LEI broadly on Ebl beginning lateral to uncinateprocess and extending almost to distal end.LE2 broadly on Eb2 posterolaterally.LE3 on CT joining Eb3 and Eb4 uncinate pro-cesses anteriorly (more closely associated with Eb3than Eb4).LE4 on Eb4 dorsally lateral to uncinate process,joining raphe with dorsal end of Ad4; Eb4 levatorprocess absent.LP finely tendinously on Eb4 at and lateral to LE4.LI1 on Pb2 dorsoanteriorly, about same size asLI2.LI2 on Pb3 dorsally opposite medial end of Eb3.TD comprises TEb2 and TEb3-Eb4. TEb2 variablycontinuous musculously transversely or broadly in-terrupted anteriorly by thick CT mid-section; at-tached dorsoanteriorly to Pb2 dorsoposteriorly bytough CT, which is continuous mid-anteroventrallywith CT of pharyngeal roof; dorsmedial surfacetightly covered by fascia, which gives rise to CTsheets attaching to skull; muscle extending laterallyand reaching almost to dorsodistal bony end of Eb2;not continuous posteriorly with TEb3-Eb4. TEb3-Eb4 on posterior edge of Eb3 medial to uncinate pro-cess and medial edge of Eb4 ventral to tip of uncinateprocess, posteriorly continuous with SOD.OD3-OD3' anteriorly on Pb3 dorsolaterally ventralto TEb2, posteriorly broadly on anterior surface ofEb3 between uncinate process and distal end, sepa-rated by shallow notch from OD3' on dorsal surfaceof Eb3 immediately ventral to OD3.OD4 absent.OP dorsally on Eb4 posteriorly beginning ventralto uncinate process and extending medially, ventrallyon Cb5 dorsoposterolaterally.Ad 1-3 absent, but moderately well-developed fan-like GFM1 present (not illustrated), but GFM2 andGFM3 weakly developed.Ad4 broadly dorsally on Eb4 posteriorly beginningventral to uncinate process and extending laterally,ventrally broadly on Eb4 dorsally medial to Eb4-Cb4joint.
Ad5 dorsally on Eb4 posterodistally (but not at-tached to AC4), ventrally on dorsoanterolateral sur-face of Cb5 anterior to OP.SOD present.RDs adjacent.Additional remarks. SCL attached mid-dorsally totip of ventroposteriorly extending cartilaginous endof Bb3. TV4 free from Cb5s. Pb4 and UP4 present.Pb2 toothed. Eb4 levator process absent. Cartilage tipon Ebl uncinate process reduced and variably absent.IAC present or absent, reduced when present, not at-taching or directly impinging on Ebl uncinate pro-cess. AC4 well-developed. Very fine Eb4 flange pres-
ent. Medial end of Eb3 larger than that of Eb4.ZANCLIDAEZanclus cornutus (Linnaeus), USNM 350564, 159mm, USNM 365537, 157 mm.Plate 172
Description.LEI slender tendinously on dorsoposterodistalmostedge of Ebl.LE2 slender tendinously on dorsoposterodistalmostbony edge of Eb2, joining posterodistalmost edge ofTEb2, ligament from insertion extends posteriorlyand attaches to Eb3 anterolaterally.LE3 slender tendinously on tip of Eb3 uncinateprocess.LE4 and LP not separable ( 1 57 mm specimen doesnot exhibit diffuse separation shown by muscle fibersof illustrated specimen, which is similar on bothsides), muscle sheet-like becoming fascia ventrallyand attaching along lateral edges of fourth and fiftharches; PP joins fascia posteriorly.LI1 on Pb2 dorsoanteriorly just lateral to dorsal-most cartilaginous tip.LI2 slender tendinously on Pb3 dorsoanterolater-ally medial to medial end of Eb3.TD comprises TEb2 and TEb3-Eb4. TEb2 vari-able, either widely interrupted mid-dorsally by thickCT pad or continuous from one side to other; whencontinuous, almost inseparably attached mid-dorsallyto thick CT pad (which, in either type TEb2, attachestightly ventrally to Pb2 and Pb3 and is tightly at-tached dorsally to thin mid-ventral process of paras-phenoid), with small muscle remnant attaching mid-posteriorly on CT pad; muscle of both types extendslaterally and attaches along most of dorsal surface ofEb2. TEb3-Eb4 of two continuous sections, anteriorsection triangular, with mid-longitudinal raphe, apexanterior, attaches to CT between Pb3s; posterior sec-tion, broad attaches mainly to dorsomedial edge ofEb4 near articulation with Eb3 uncinate process andsecondarily tendinously to medial edge of Eb3 justventral to tip of uncinate process, continuous poste-riorly by diagonal muscle strands with SOD.
NUMBER 11 203OD3 origin mostly ventral to TEb2, but smallbranch (not illustrated) extends anteroventrally andattaches on dorsolateral surface of Pb2, main portionattaches to CT pad attached to dorsolateral surface ofPb3, insertion on dorsolateral surface of Eb3.OD4 absent.OP slender, dorsally on Eb4 posteriorly joiningTEb3-Eb4 ventrolaterally, ventrally on Cb5 just pos-teromedial to Ad5.Ad 1-3 absent, but weak GFMs present on anteriorsurfaces of relevant Eb-Cb joints of first three archesand, unusual for fishes, on anterolateral surface ofEb4.Ad4 dorsally on Eb4 posterolaterally beginningventral to margin articulating with Eb3 uncinate pro-cess and extending laterally to near end of bony sur-face, ventrally, narrowly on Cb4 dorsally medial toEb4-Cb4 joint.Ad5 dorsally narrowly on Cb4 posterodistally,ventrally narrowly on Cb5 dorsodistally anterior toOP.SOD present.RDs relatively small, slightly separated.Additional remarks. SCL attached mid-dorsally toautogenous cartilage, which is attached to ventropos-terior cartilaginous end of Bb3. TV4 free from Cb5s.Pb4 and UP4 present. Pb2 toothed.ACANTHURIDAEAcanthurus nigrofuscus (Forsskal), USNM 338195,3 specimens, 101-104 mm.Plate 173
Description.LEI finely tendinously and musculously (tendonborders muscle anteriorly) on dorsoposterodistalmostedge of Eb 1
.
LE2 finely tendinously and musculously (tendonborders muscle anteriorly) on dorsoposteriorly ex-panded surface Eb2 laterally.LE3 finely tendinously and musculously (tendonborders muscle anteriorly) on tip of Eb3 and Eb4uncinate processes, mainly on Eb3.LE4 and LP not separable (if both are present),muscles sheet-like, becoming broad CT sheet ven-trally and attaching along lateral edges of fourth andfifth arches; various tendinous extensions of the CTsheet attach to other skeletal elements; fine tendonextends from LE4+7LP ventroanteriorly and insertson Eb4 lateral to uncinate process (possibly indicat-ing LE4 insertion); CT sheet joined posteriorly byPP. See also remarks following LE4 in Luvarus.LI1 on Pb2 dorsoanteriorly ventral to dorsalmosttip of Pb2.LI2 on Pb3.TD complex comprising TEb2, TEb4, and TDplexus. TD plexus comprises three slender muscles
(easily removed during dissection) originating fromdorsoposterolateral edge of CT pad; anterior muscleextends anterolaterally passing dorsal to TEb2 andinserts on posterodistalmost cartilaginous edge ofEbl at LEI insertion; middle muscle extends laterallyand slightly anteriorly and inserts finely, tendinouslyon posterior edge of Eb2 just posterior to LE2 inser-tion; posterior muscle extends laterally and slightlyposteriorly, and inserts on Eb4 uncinate process.TEb2 broad, originates on CT pad just anterior to TDplexus, fans out laterally, and attaches on most ofbony dorsal surface of Eb2. TEb4 uninterrupted, in-
serts on Eb4 posteriorly just medial to uncinate pro-cess, finely continuous posteriorly with SOD.OD3 originates on lateral edge of Pb3 and insertson most of bony dorsal surface of Eb3.OD4 absent.OP dorsally mostly on Eb4 posteroventrally. butmeeting TEb4 dorsolaterally; ventrally on Cb5 dor-
sally.GFM1-3 fine splay of muscle fibers on anteriorsurface of Eb-Cb joints (weakest on Eb3-Cb3), at-taching laterally to gill filaments.Ad4 dorsally on Eb4 posteriorly beginning ventralto uncinate process and extending laterally, ventrallynarrowly tendinously on Cb4 posterolaterally.Ad5 on Cb4 posterodistal end and AC posterolat-
erally, joining raphe dorsoanteriorly with OP ventro-laterally; debatable if Ad5 joins CT attaching AC toCb4.SOD slender.RDs slender, proximate posteriorly, separated bydistance less than one RD diameter anteriorly.Additional remarks. SCL attached mid-dorsally toautogenous cartilage, which is attached to elongateposteroventral cartilaginous end of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. Pb2 toothed. IACabsent. Relatively large AC4 attached to dorsodistal-most surface of Cb4 and posterodistalmost surface ofEb4. IAC absent. Eb4 levator process absent.Anabantomorpha
Britz (2003) briefly recapped the classificatory his-tory of the suborder Anabantoidei, in which he rec-ognized three families (Anabantidae, Helostomatidae,Osphronemidae), which have well-developed supra-branchial organs. He stated that the closest relativesof the group appear to be the Channidae, which havea much lesser-developed suprabranchial organ, andthat the four families, together with Badidae, Nandus(family not indicated, but Nandidae is available), andPristolepis (family not indicated, but Pristolepidae isavailable) form a monophyletic group based on thesynapomorphic presence of parasphenoid teeth. Torecognize the monophyly of this group of families,we erect a new ordinal-group name Anabantomor-
204 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
pha, retaining Anabantoidei for the four familieswith suprabranchial organs (note: we did not examineany heleostomid or osphronemid gill arches).NANDIDAENandus nebulosus (Gray), USNM 328105, 49.3 mm.Not illustrated
Description.LEI on Ebl dorsoposteriorly a little lateral to ar-ticulation with IAC (no uncinate process, IAC verylong, articulates directly with bone of Ebl; like Pris-tolepis, Plate 174).LE2 on bony prominence on Eb2 mid-dorsopos-teriorly.LE3 finely, tendinously on tip of Eb3 uncinate pro-cess.Remarks. Liem (1970:56-57) stated (and illustrat-ed) that only three LEs are present in the nandids heexamined (included N. nandus and N. nebulosus).There is a possibility that he either overlooked thefine LE3 we observed, or that there is variation in itsoccurrence.LE4 largest levator, on bony surface of Eb4 justanterior to cartilage tip of levator process.LP on Eb4 posterolaterally beginning medially atlateral edge of LE4 insertion and extending laterally.LI1 on dorsoanteriormost cartilaginous tip of Pb2
at joint with IAC.LI2 on Pb3 dorsolaterally with main portion of in-sertion continuing onto UP4 dorsally ventral to me-dial end of Eb3.TD comprises TEb2, TPb3-Eb3-Eb4. TEb2, welldeveloped, uninterrupted (musculous medially), withmid-longitudinal raphe; muscle extending laterallyand attaching on Eb2 dorsally anterior to LE2 inser-tion (dorsal surfaces of Pb3s covered by muscle).TPb3-Eb3-Eb4 on Pb3 beginning anteromedial tomedial end of Eb3, continuing posteriorly on carti-laginous posteromedial end of Eb3, and then ontoanterior and posterior bony edges of Eb4 medial tobony support of uncinate process.OD3-4 originating on Pb3 dorsomedially and in-serting on Eb3 anterior surface ventral to tip of un-cinate process and on Eb4 dorsally just lateral to tipof uncinate process.OP strap-like; dorsally on Eb4 ventroposteriorlybeginning about mid-way between medial end andlevator process and extending laterally almost to le-vator process, posteriorly overlapping dorsomedialedge of Ad4; ventrally on Cb5 dorsally, beginningimmediately medial to cartilaginous posterior tip andextending a short distance medially and meeting pos-terolateral edge of TV5.Ad 1-3 weakly developed, spanning respective Eb-Cb joint anteriorly posterior to gill rakers; easilyoverlooked.
Ad4 dorsally on Eb4 ventroposteriorly, beginninga little ventromedial to levator process (and anteriorto OP laterally) and extending laterally most of bonylength of Eb4; ventrally on Cb4 dorsally beginninga little medial to Eb4-Cb4 joint and extending me-dially a short distance.Ad5 anteriorly on posterodistalmost surface ofCb4; posteriorly on posterodistal surface of Cb5,meeting OP.SOD absent.RDs well separated at origins, gradually converg-ing and becoming adjacent at insertions on Pb3s me-dially and UP4s dorsomedially.Additional remarks. SCL free from posterior endof Bb3. TV4 free from Cb5s. Pb4 absent. UP4 pres-ent. IAC very long. Pbl present, bony, with dorsaland ventral cartilaginous caps. Eb4 with small bonyflange dorsoanterodistally. Pb2 toothed.See Interrelationships section following additionalremarks in description of Badis kyar (Badidae) forcomments on other characters of anabantomorphs.BADIDAEBadis kyar Kullander and Britz, USNM 343545, 39.1mm. Not illustrated
Description.LEI on Ebl mid-dorsoposteriorly.LE2 insertion begins on Eb2 mid-dorsoposteriorlyand continues onto Eb3 mid-dorsoanteriorly.LE3 on Eb3 just ventral to tip of uncinate process.LE4 on Eb4 dorsally posterior to all bony uncinateprocess, meeting LP insertion anteriorly. Insertion isdorsal to minute cartilage tip of levator process onone side, cartilage tip absent on other side.LP on Eb4 at and posterior to LE4 insertion, fiberscontinuous posteriorly with Ad4 dorsoposterolater-
ally.LI1 on dorsoanteriormost cartilaginous tip of Pb2
at joint with IAC.LI2 on Pb3.TD comprises an undivided TEb2 (which is onlyweakly developed as it passes over the Pb3s dorsally)and attaches on Eb2 anterior to LE2 insertion, andTPb3-Eb4. Most of the dorsal surface of Pb3s notcovered by muscle.OD3-4 origin on Pb3 dorsomedially ventral toTEb2; insertion on Eb3 dorsoanteriorly just ventralto tip of uncinate process and on Eb4 dorsoposter-iorly just ventral to tip of uncinate process; separate(anomalous?), small posterior section originates onPb3 dorsomedially immediately posterior to other(usual) OD-3-4 origin, extends laterally, and curvesa little as it extends to its attachment on Eb4 justmedial to dorsomedial edge of OP.OP present, strap-like, dorsally on Eb4 ventrally
NUMBER 11 205
ventral to LP insertion, ventrally on Cb5 dorsallynear distal end, there meeting Ad5 posteriorly.Ad 1-3 very well developed, probably relativelylarger than in any other acanthomorph taxon we ex-amined, broadly spanning respective Eb-Cb joint.Ad4 dorsally on Eb4 ventrally beginning anteriorto OP laterally and extending to Eb4-Cb4 joint, con-tinuous dorsoposterolaterally with LP posteroventral-ly; ventrally on Cb4 extending medially from Eb4-Cb4 joint.Ad5 attaches distal end of Cb5 to distal end ofCb4.SOD absent.RDs relatively massive, juxtaposed.Additional remarks. SCL apparently free fromBb3. TV4 free from Cb5s. IAC similar to that ofNandus in relative length and in joining only thebone of Ebl (uncinate process absent in both taxa).Pbl is entirely cartilaginous (mostly bony in Nan-dus). Pb4 absent. UP4 present. Eb4 with small bonyflange dorsoanterodistally. Pbl cartilaginous. Pb2toothed.Interrelationships. Kullander and Britz (2002:300)hypothesized a monophyletic group comprising Bad-idae and Nandidae (latter including only Nandus)based primarily on the shared presence of a deeplybifurcated hemal spine on vertebra PU2, which oc-curs in no other teleost [we found a deeply bifurcatedHPU2 in two of three radiographed specimens ofScatophagus argus, USNM 259383, but not in fivecleared and stained specimens, USNM 197528,224393]. Kullander and Britz offered as additionalevidence for monophyly, the presence of 7 + 7 prin-cipal caudal-fin rays, compared with 8 + 8 in Poly-centridae and Pristolepidae, various members ofwhich have been considered by authors as related tothe nandids.In addition to those characters reported by Kullan-der and Britz, we find that our specimens of Badidaeand Nandidae are unusual in having the IAC articu-lating directly with the bony surface of Eb 1 , whichlacks a cartilage-tipped Ebl uncinate process. A sim-ilar condition is found in the probably closely relatedPristolepis (Pristolepidae). A somewhat similar con-dition also exists in the synbranchiform Synbranchi-dae, in which there is an IAB (purportedly an ossifiedIAC; Rosen and Greenwood, 1976). The synbranchidIAB articulates directly with the bony surface of Eb 1
,
which lacks a cartilage tipped uncinate process. Thecondition in certain atherinomorphs (aplocheilids, cy-prinodontids), in which an Ebl uncinate process ap-pears to be absent and IAC articulates with the distal,cartilaginous end of Ebl, is also dissimilar to the con-dition in Badidae and Nandidae.Nandids and badids share in having another spe-cialization, presence of a bony flange, or spur, over-lapping cartilaginous distal end of Eb4. Although
limited to and common among percomorphs, the Eb4flange occurs otherwise in anabantomorphs only inanabantids, osphronemids (Luciocephalus, Britz,1995:fig. 4; Macropodus, Betta, Britz, 2001:figs. 2b,
c; flanges not labelled in any of Britz's figures, butpresence clearly indicated), and probably heleosto-matids (not examined by us). The flange is not pres-ent in pristolepids and the channids we examined.PRISTOLEPIDAE
Pristolepis fasciata (Bleeker), USNM 103105, 68mm. Plate 174
Description.LEI slender, tendinously on Ebl dorsoposteriorlya little lateral to articulation with IAC (no uncinateprocess; IAC articulates directly with bone, as it doesin Nandus and Badis), left side with muscle fibersextending along entire medial edge of incorporatedtendon; muscle attaching only to distal end of slendertendon on right side.LE2 slender, mostly tendinously on slight bonyprominence on dorsoposterior edge of Eb2.LE3 finely tendinously on tip of Eb3 uncinate pro-cess, tendon partly joining OD3-4 (muscle also pres-ent on both sides of a partially dissected specimen).Remarks. Liem (1970:56-57) stated that only threeLEs are present in the anabantomorphs he examined(included P. fasciata). There is a possibility that heeither overlooked the fine LE3 we observed, or thatthere is variation in its occurrence.LE4 largest levator, tendinously on dorsodistal-most bony edge of Eb4 levator process.LP tendinously at and anterior to LE4 insertion,extending onto dorsodistalmost end of levator pro-cess.LI1 on dorsoanteriormost cartilaginous tip of Pb2
at joint with IAC.LI2 tendinously on dorsoposterolateral surface ofPb3 near junction with Pb4 and medial end of medialend of Eb3.TD comprises TEb2 and TEb4. TEb2 comprises amuscle pair, each member of which is joined to lat-eral edge of broad, very thick CT pad, which attachesventroanteromedially to Pb2; posteroventral half ofpad is free, overlies and is continuous anteroventro-laterally with bilateral pair of thinner CT pads, eachof which covers dorsal bony surface of its respectivePb3 and is attached to dorsomedial edge of Pb3; ven-tral pad is joined laterally by OD3-4 origin; muscleextends laterally and attaches on Eb2 dorsally ante-
rior, or slightly anterolateral, to LE2 insertion. TEb4attaches on Eb4 posteromedially.OD3-4 origin on lateral edge of CT pad coveringPb3 dorsally and on dorsoanterior bony surface ofPb3, insertion on medial edge and anterior surface of
206 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Eb3 uncinate process and medial edge and anteriorsurface of Eb4 uncinate process.OP dorsally on posterior surface of Eb4, extendinglaterally from uncinate to levator process, ventrallyon Cb5, ventrolaterally penetrated by raphe, mediallycontinuous with SO.GFM1-3, that on first arch very weakly developed,
all three just reaching to anterodistalmost end of re-spective Cb.Ad4 dorsally on posterolateral surface of Eb4, ven-trally on dorsal Cb4 surface anterior to Eb4-Cb4joint.Ad5 dorsally tendinously on posterolateral surfaceof Eb4 and ventrally on dorsodistalmost end of Cb5,joining raphe anteroventrolaterally with OP.SOD absent.RDs separate, insert on Pb3 and UP4 posteriorly.Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Pb4 absent. UP4 present. Pbl bony withcartilaginous ends. Pb2 toothed.See Interrelationships section following additionalremarks in description of Badis kyar (Badidae) forcomments about the relationships of Nandidae, Bad-idae, Pristolepidae, Channidae and Anbantoidei.CHANNIDAE
Channel asiatica (Linnaeus), USNM 191304, 117mm; USNM 192925. 98.0 mm.Plate 175
Additional material. Channa harcourtbutleri (Annan-dale), USNM 191465, 90.4 mm.
Description.LEI on posterior edge of broadly expanded Ebl alittle lateral to mid-length (uncinate process absent).LE2 slender, on tip of bony process at distal endof Eb2. .Remarks. Insertion of LE2 at the distal end of Eb2is uncommon in acanthomorphs. It also occurs insome gobioids and atherinomorphs as a specializedstate within these groups. It also appears to be a spe-cialized state within anabantomorphs. That TEb2fails to reach the LE2 insertion in Channa is probablythe result of the extreme lateral shift in position ofthe LE2 insertion, not of a reduction in the lateralextent of TEb2.LE3 slender, on tip of Eb3 uncinate process.LE4 slender, tendinously on Eb4 dorsally at aboutmid-anterior point of LP insertion, closely appliedposteriorly to anterior surface of LP and difficult todistinguish from LP.LP massive, broadly dorsally on most of distalmosthalf of Eb4, joining extensive raphe posteriorly withOP dorsally.Remarks. The angle of direction from insertion toorigin is the same as that of LE4. In most acantho-
morphs the LP angle diverges noticeably from thatof LE4. LP normally inclines posteriorly or poster-omedially from LE4.LI1 on Pb3 dorsoanteromedially (note: Pb2 welldeveloped; positioned ventral to anterior portion ofEb2, relatively well-separated from Pb3); insertionheavily enveloped in CT involving medial end ofEb2 and ventral surface of thick CT area joiningTEb2 with contralateral TEb2.LI2 on Pb3 dorsoposteriorly.TD comprises TEb2 and TPb3-Eb4. TEb2 a pairof muscles joined by broad area of tough, thick, lay-ered CT, which is notched anteriorly as it passesaround the parasphenoid tooth patch, and gives riseto CT sheet attaching to skull, muscle fibers essen-tially cover Pb3 dorsal surface, but are separatedfrom it by thick CT layer; muscle attaching to a littlemore than medial half of posterior edge of Eb2, fail-ing to reach the insertion of LE2 (see remarks fol-lowing LE2), not continuous posteriorly with TPb3-Eb4. TPb3-Eb4 narrowly on Pb3 dorsoposteromedi-
ally, continuing more broadly posteriorly and attach-ing on Eb4 posteromedially.CPb (not illustrated) a slender extension of the SOlongitudinal muscle layer jointly encircling Pb3 andUP4, with an even more slender muscle filamentpassing between Pb3 and UP4 and connecting bothsides of the encircling portion of the muscle.OD3-4 origin dorsally from margin of large cen-trally, located CT sheet, ventrally from Pb3 dorsallyjust lateral to dorsomedial surface; minor insertion onEb3 uncinate process anteriorly, main insertion onEb4 beginning on anterior surface just posterior toEb3 uncinate process, and continuing laterally onnarrow dorsal surface to medial edge of LP.OP dorsally variably broadly on Eb4 posteriorly,meeting OD3?4 and LP; always most distinct ven-trolaterally with fibers attaching to tendinous sheetcoursing along ventrolateral surface of muscle andconforming with Ad5 dorsally; ventrally OP meetsPCI but does not join raphe with it; medial edge ofOP weakly to well separated from SO.Adl absent.Ad2 bulky muscle dorsally on Eb2-Cb2 joint an-teriorly, extending short distance ventrally on Cb2.Ad3 dorsally on much of anterior edge of Eb3, butceasing a little medial to distal end, there a short,separate portion present dorsally, both portions at-taching ventrally to anterior surface of Cb3 slightlyventral to Eb3-Cb3 joint.Ad4 dorsally on most of Eb4 ventrally, continuinglaterally to end of inner angle of Eb4-Cb4 joint, ven-trally on most of Cb4 dorsally (not visible external-ly).Ad5 dorsally on posterodistal surfaces of Eb4 andCb4, ventrally on Cb5 posterodistally; with fine ra-phe-like inclusion on surface (both sides) only in
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larger specimen (illustrated); joins raphe with PCInear distal end of Cb5.
?
Dorsally only on Cb4 dis-tally.SOD absent.RDs separated by space less than half one RD di-ameter.Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Pbl bony with ventral end cartilaginous.Pb2 finely toothed. Pb4 absent, UP4 present. Eb4 un-cinate and levator processes absent. IAC absent.ANABANTIDAESandelia bainsii Castelnau, USNM 363314. 83.7mm. Not illustrated
Additional material. Ctenopoma kingsleyae Giinther.USNM 288037. 54.1 mm. The muscles appear tobe essentially the same as those of Sandelia.
Description.LEI slender, short, on posterior edge of broadlyexpanded Ebl mid-dorsoposteriorly.Remarks. Right-side Ebl has a fine, cartilagetipped uncinate process joining a very slender IAC,which becomes finely ligamentous medially. It is un-clear what the ligament attaches to; it was brokenduring dissection, but was probably attached to thetip of long dorsal process of Pb2. Ebl uncinate pro-cess and IAC are absent on left side.LE2 slender, short, on raised bony prominence onEb2 mid-posteriorly.LE3 on tip of Eb3 uncinate process anteriorly.LE4 robust, on Eb4 dorsolaterally. fused poster-oventrally with LP, but the two muscles have slightlyseparate origins.Remarks. The bony dorsal surface of Eb4 extendslaterally as a shelf (flange), the ventral surface ofwhich attaches to the somewhat elongate dorsal car-tilaginous distal end of Cb4. On one side, just medialto the attachment of Cb4 to Eb4, is an alcian-bluestained area, which probably represents a vestige ofa former (present early in ontogeny?) distal cartilag-inous end of Eb4. The area is absent on the otherside. Eb4 and the joint with Cb4 are closely remi-niscent of the structures in cichlids, except that cich-lids retain the cartilaginous distal end of Eb4. Alsoin cichlids, LE4 fuses posteroventrally with LP.LP fusing ventroanteriorly with LE4 and insertingtogether on Eb4 dorsolaterally.LI1 inserts by slender tendon on Pb3 dorsoanter-olaterally.LI2 inserts by short tendinous strap on Pb3 an-terolaterally, a little medial to articulation with me-dial end of Eb3.TD comprises only TPb2 and TPb3. TPb2 joinslateral margin of CT pad covering otherwise naked
Pb3 dorsal facet; muscle is small, sheet-like, and al-most vertical, extending laterally and attaching toPb2 along medial edge and surface of long, dorso-medially extending Pb2 uncinate process. TPb3 at-taches broadly to posteromedial margins of Pb3s.CPb extends as a broad continuation of SO lon-gitudinal muscle between pharyngobranchials, divid-ing and becoming string-like anteriorly and extendingaround periphery of Pbs on each side, but with shortmedial branch attaching to posterolateral corner ofPb2, and posteromedially re-joining SO.OD3-4 relatively large, most prominent muscle;anteriorly, broadly on Pb3 dorsomedially, posteriorlyon Eb3 uncinate process dorsoanteriorly and on allbony Eb4 uncinate process dorsally.OP dorsally on Eb4 ventrally beginning mediallywell medial to LE4-LP insertions and extending lat-erally to about mid-way below insertions, ventrallyon Cb5 dorsoposterolaterally, meeting but not joiningraphe with PCI and joining raphe with Ad5 ventro-medially.Adl-3 absent. GFM1 a thin muscle extendingalong posterior edge of Ebl from near LEI insertionto Ebl-Cbl joint. GFM2 with small attachment toEbl posteriorly well lateral to LEI insertion, withmain portion of muscle on anterolateral edge of Eb2,reaching to Eb2-Cb2 joint. GFM3 well developed, onEb3 dorsally with small attachment to Eb2 posteri-orly. These muscles do not span the anterior surfacesof the Eb-Cb joint.Ad4 dorsally, broadly on Eb4 ventrally beginningmedially anterior to OP and ventral to LE4-LP in-sertions and extending laterally to near distal end;ventrally on Cb4 dorsally, narrowly on distalmostbony surface.Ad5 dorsally on Eb4 ventrally at distalmost sur-face, ventrally on Cb5 dorsodistally, joining raphewith OP ventromedially.SOD absent.RDs adjacent.Additional remarks. SCL present. TV4 free fromCb5s. Pb4 absent. UP4 present. Pbl bony with car-tilage ends. Pb2 finely toothed; teeth absent in Cten-opoma. Eb4 levator process absent. Medial end ofEb4 much larger than medial end of Eb3.See Interrelationships section following additionalremarks in description of Badis kyar (Badidae) forcomments about the relationships of Nandidae, Bad-idae, Pristolepidae, Channidae and Anbantoidei.
Stromateoidei
Ever since Haedrich's (1969) original descriptionof Amarsipus and consideration of it as an aberrantstromateoid, Amarsipus has been considered to be astromateoid. Although there is a general external sim-ilarity of Amarsipus to the "true" stromateoids (taxa
208 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
sharing the specialization of having a toothed sac-cular outgrowth of the gut posterior to fourth gillarch), a close relationship between the two groupshas not been hypothesized cladistically. Amarsipuslacks the saccular outgrowth, and, therefore, thatcharacter cannot be used to relate it to the stroma-teoids. The other characters used by Haedrich (1967)to define the Stromateoidei (for list, see interrelation-ships discussion of the Icosteidae, which has alsobeen considered to be closely related to the stroma-teoids), are either generalized or inexplicit, with thepossible exception of their possessing "an extensivesubdermal canal system," a character that, to ourknowledge, has not been surveyed.Freihofer (1973:table 1, p. 138) reported that stro-mateoids have the highly specialized pattern 10 ofthe orbito-pectoral branch of the ramus lateralis ac-cessorius (RLA-OP). Only in pattern 10 does theRLA-OP pass posteriorly beneath the skin parallelingthe pterotic canal, then pass between the dorsal endof the preopercular canal and posterior end of thepterotic, continue beneath the skin overlying the le-vator opercularis muscle, and then extend mediallyaround the joint between the post-temporal and su-pracleithrum onto the medial side of the pectoral gir-dle. We checked the posterior extension of RLA-OPin the stromateoid Peprilus burti (USNM uncata-loged, field no. U60-7), beginning in the region nearthe dorsal end of the preopercular canal and corrob-orate Freihofer's observations. We also dissected aspecimen of Amarsipus (SIO 61-547), beginning inthe region near the dorsal end of the preopercle todetermine the position of its RLA-OP. RLA-OP inAmarsipus appears to parallel the pterotic, but it doesnot pass below the surface of the skin nor does itapproach the post-temporal-supracleithrum joint.Rather, it passes medial to the thick mass of opercularmuscles and extends posteriorly well ventral to thepost-temporal-supracleithrum joint as it enters themedial side of the pectoral girdle. Inasmuch as wewere unable to completely trace RLA-OP from itsorigin, we cannot say which, if any, of Freihofer'sRLA-OP patterns pertains to Amarsipus, but it is cer-tainly not pattern 10 as he described it. This findingdoes not necessarily provide evidence that Amarsipusis not closely related to the stromateoids (see alsoInter-relationships discussion following descriptionof Icosteus, Icosteidae), but it does suggest that morestudy is necessary before it can be "concluded" thatAmarsipus and stromateoids are closely related.Johnson and Fritsche (1989) included the stroma-teoids together with a group of families that we havesegregated in the Girelloidei. They based their groupprimarily on the fact that all shared Freihofer's RLApattern 10. Johnson and Fritsche did not examineAmarsipus. If Amarsipus is a stromateoid (sistergroup to all other stromateiods), more evidence will
be necessary in order to relate the stromateoids to thegirelloids.Our inclusion of the monotypic Amarsipidae in theStromateoidei is provisional.AMARSIPIDAEAmarsipus carlsbergi Haedrich, SIO-75-116, 44.0mm. Plate 176
Description.LEI on dorsal edge of Ebl at and just lateral totip of uncinate process.LE2 on raised dorsoposterior edge of Eb2.LE3 very slender, on dorsoanterior edge of Eb3uncinate process.LE4 on Eb4 just lateral to cartilage tip of uncinateprocess (no levator process).LP on Eb4 at distal edge of LE4 insertion.LI1 on anteromedial surface of Pb2 medial to IACattachment.LI2 on Pb3 dorsolaterally.TD comprises TPb2, TEb2, and TPb3-Pb4. TPb2thin, heart-shaped (notched anteriorly) with mid-lon-gitudinal raphe, attached anterolaterally to anterior tipof Pb2, broadly continuous ventrally with TEb2.TEb2 with (unusual) slight but definite tendinous at-tachment to anterior tip of Pb2 at junction of LI1insertion and with anterolaterally and posterolaterallyextending fibers joining at about medial end of Eb2and attaching on dorsal surface of Eb2 anterior toLE2 insertion; posteriorly free from TPb3-Pb4. Fi-bers from TPb3-Pb4 extending anteriorly ventral toTEb2 and attaching to Pb3s anteromedially, posteri-orly attaching to dorsoposterior surface of Pb3 anddorsal surface of Pb4, continuous posteriorly withSOD.OD3-4, OD3' origin on Pb3 dorsolaterally ventralto TEb2, inserting on dorsoanterior surface of Eb3uncinate process and dorsoanterior surface and dor-somedial edge of Eb4, forming ventrolateral branch,OD3', slightly distal to mid-length, and inserting onEb3 dorsoanterior surface just ventral to OD3-4 in-sertion.OP dorsally on Eb4 posterior surface at and medialto uncinate process, ventrally on posterodistal surfaceof Cb5, posterior to Ad5 attachment.Ad 1-3 absent.Ad4 dorsally on ventral surface of Eb4 lateral touncinate process, ventrally on Cb4 dorsally anteriorto Eb4-Cb4 joint.Ad5 ventrally on Cb5 dorsodistal surface, dorsallyon Cb4 distal end.SOD present.RDs separate.Additional remarks. SCL appears to be free fromBb3 (needs verification in another specimen). TV4
NUMBER 1 1 209
free from Cb5s. See Icosteidae for discussion ofAmarsipus interrelationships. Pb4 and UP4 present.Pb2 toothed. CENTROLOPHIDAEPsenopsis sp., USNM 304398, two specimens, 80.5-90.0 mm. Plate 177
Description.LEI on dorsal edge of Ebl uncinate process justlateral to cartilage tip.LE2 on mid-dorsoposterior edge of Eb2.LE3 on tip of Eb3 uncinate process.LE4 on dorsoanterior surface of Eb4 levator pro-cess.LP on Eb4 levator process just anterior to LE4insertion.LI1 tendinously on dorsoanterolateralmost surfaceof Pb2 just medial to joint with IAC.LI2 on Pb3 dorsoposteriorly and dorsoanterolateraledge of Pb4, meeting posterolateral edge of TPb3-Pb4-Eb3.TD comprises TPb2, TEb2, and TPb3-Pb4-Eb4.TPb2 an almost circular ribbon of muscle interruptedanteriorly where it ventrally joins CT of pharyngealroof and dorsally joins CT sheet covering muscle andattaching to skull; muscle attaching anteroventrola-terally to joined cartilaginous dorsoanterior ends ofPb2 and Pb3, posteriorly fusing with TEb2 and pos-terior continuation of pharyngeal roof CT. whichtightly covers musculously naked Pb3 dorsal facets(TEb2 muscle fibers do not extend across central areabounded by TPb2, but in larger specimen three iso-lated islands of muscle are present posterior to TPb2mid-anteriorly). TEb2 thickest anteriorly, meetingthin posterior fibers and "squeezing" as the muscleextends onto Eb2 dorsally, thick portion attaching onEb2 dorsoanteriorly at point anterolateral to LE2 in-sertion, thin portion attaching to Eb2 dorsally at andanterior to LE2 insertion; muscle not continuous pos-teriorly with TPb3-Pb4-Eb3. TPb3-Pb4-Eb3 with finestrands of muscle attaching to Pb3 along medial edgeof LI2 insertion, to bony posteromedial edge of Eb3,and to dorsal surface of Pb4 where fibers mesh withthose of anterior end of M. SO-Pb4.M. SO-Pb4 originates from SO fibers associatedwith lateral surface of EO and extends dorsoanter-iorly and attaches to dorsal surface of Pb4 togetherwith TPb3-Pb4-Eb3.OD3-4 originating on Pb3 dorsoanteriorly ventralto TEb2, inserting on dorsoanterior surface of Eb3uncinate process and anteromedialmost surface ofEb4 uncinate process.OP apparently absent. Muscle fibers attaching toposterior surface of Eb4 levator process mesh later-ally with Ad4 and ventrally with EO and M. SO-Pb4
fibers; a few fibers may extend to Cb5; situation un-resolvable in specimens.Ad 1-3 absent.Ad4 dorsally on posterodistal surface of Eb4, ven-trally attaching by CT to dorsodistalmost end of Cb4;EO attaches to posterodistalmost surface of Cb4 andseparates Ad4 and Ad5 attachments.Ad5 dorsally on ventrodistalmost end of Cb4, ven-trally on Cb5 distally (see also Ad4).SOD absent.RD posteriorly laterally compressed, slightly sep-arated from counterpart, extends well anteriorly andattaches to anteromedial surface of Pb3.Additional remarks. SCL attached mid-posteriorlyto ventroposterior cartilaginous tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. IAC present.
ZoarcoideiImamura and Yabe (2002) hypothesized that theZoarcoidei and Cottoidei comprise a monophyleticgroup (unnamed), which they include in the Perci-formes. We found some evidence that might corrob-orate their hypothesis (see Additional remarks fol-lowing description of Hexagrammos stelleri, Hexa-grammidae, included under Scorpaeniformes).BATHYMASTERIDAEBathymaster signatus Cope, USNM 339381, 101mm; USNM 339378, 107 mm.Plate 178
Description.LEI on medial half of Ebl dorsally.Remarks. Anterior arm of Ebl interpreted as ab-sent because the medial end of Ebl articulates withPb2.LE2 on raised dorsoposterior edge of Eb2.LE3 on dorsoanterior edge of tip of Eb3 uncinateprocess.LE4 on dorsoposterior surface of Eb4 a little morethan half distance to distal end, beginning above la-teralmost edge of OP.LP at and posterior to posterior edge of LE4 in-sertion.LI1 in smaller specimen on Pb2 dorsoanteriorlyand Pb3 anterolateralmost edge; in larger specimenon dorsolateral surface of Pb2 and anterolateral edgeof Pb3.LI2 on Pb3 at and lateral to TPb3 attachment.TD comprises TEb2, TPb2, and TPb3-Eb3. TPb2a laterally curving semicircular ribbon of muscle oneach side arising anteriorly on each side from mid-anterior end of TEb2 and fusing at mid-posterior end;muscle not attached to Pb2. TEb2 lies dorsal to an-terior ends of Pb2 and Pb3 but is free from them; itsoblong dorsomedial section is notched mid-anteriorly
210 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and mid-posteriorly, and divided mid-Iongitudinallyby a raphe that attaches ventrally to CT of pharyngealroof and dorsally to CT sheet that attaches to skull;laterally, TEb2 attaches along more than half lengthof Eb2 dorsal surface; posteriorly, TEb2 is free from,and dorsal to level of, TPb3-Eb3. TPb3-Eb3 broadlyon Pb3 dorsal surface beginning along medial edgeof LI2 insertion and continuing posteriorly and at-taching to the very medialmost edge of Eb3, contin-uous posteriorly by broad, diagonal muscle strap withSOD.OD3-4 origin on dorsomedial surface of Pb3 ven-tral to TEb2, insertion on anterior surface of Eb3 un-cinate process, and medial edge of Eb4 uncinate pro-cess.OP dorsally on posterior surface of Eb4, beginningslightly lateral to tip of uncinate process and extend-ing well medially, ventrally on dorsolateral surfaceof Cb5, overlapping much of Ad5 posteriorly.Ad 1-3 absent (short cord-like GFM on each Ebdorsolaterally, extending onto anterodistalmost endof respective Cb).Ad4 broadly on ventral surface of Eb4 and dorsalsurface of Cb4 medial to Eb4-Cb4 joint, beginningmedially a little anterior to OP.Ad5 broadly on both dorsal surface of Cb5 andposterolateral surface of Cb4.SOD present.RDs separated by narrow space less than one-halfdiameter of one RD.Additional remarks. SCL attached mid-dorsally tocartilaginous ventroposterior tip of Bb3. TV4 freefrom Cb5s. Pb4 and UP4 present. Pbl, IAC, and Eb4levator process absent. Pb2 toothed. Medial end ofEb3 larger than that of Eb4.ZAPRORIDAEZaprora silenus Jordan, USNM 306369, 1 1 3 mm.Not illustrated
Description.LEI tendinously on mid-dorsoposterior edge ofEbl; uncinate process absent.LE2 narrowly tendinously on Eb2 nearer medialend than distal end, ligamentous connection fromposterior base of insertion to anterior margin of Eb3.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 posterolaterally, but medial to distalend, insertion overlapped slightly by anteromedialedge of LP insertion.LP on Eb4 beginning slightly anterior to lateraledge of LE4 insertion and extending to cartilaginousdistal end Eb4.LI1 on joined dorsoanteriormost tips of Pb2 andPb3, slightly larger than LI2.LI2 on Pb3 dorsally medial to medial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2
a bilateral pair of broadly ribbon-like semicircularmuscles arising from TEb2 mid-dorsoposteriorly,curving laterally, and joining mid-longitudinal raphe,forming cup-shaped depression floored by TEb2 me-dially and some CT. TPb2 attaches anteroventrolater-ally to anterodorsalmost tip of Pb2; mid-longitudinalraphe attaches ventrally to CT of pharyngeal roof anddorsally gives rise to CT sheets covering muscles.TEb2 extends laterally from TPb2 and attaches toEb2 dorsally well anterolateral to LE2 insertion,meeting anteromedial end of GFM2. TPb3-Eb3 be-gins anterior as small, almost separate branch attach-ing to Pb3 anterior to LI2 insertion (same on bothsides) and continues posteriorly as much broader por-tion, which attaches to posteromedial edge of Eb3,posteriorly continuous by diagonal muscle strandswith SOD.OD3 originates on Pb3 dorsoanteromedially andinserts on Eb3 just ventral to tip of uncinate process;on one side only, a few muscles fibers attach weaklyto dorsomedialmost edge of Eb4 uncinate process.OD4 (see OD3). We consider OD4 absent, but ver-ification in other specimens needed.OP dorsally on Eb4 dorsoposteriorly. beginningmedial to uncinate process and extending laterally tobelow LE4 insertion, ventrally on Cb5 posterolater-
ally, overlapping Ad5 posteromedially.Ad 1-3 absent.GFM 1 weakly fan-shaped.GFM2 well developed, dorsomedially meetingTEb2 distally, extending laterally, weakly spanningEb2-Cb2 joint and extending narrowly ventrallyalong Cb2 margin.GFM3 similar to GFM2 (however, does not meetOD3).Ad4 dorsally on Eb4 posterolaterally beginningbelow LE4 insertion and extending laterally to distalend of Eb4, ventrally on Cb4 dorsally anterior toAd5.Ad5 ventrally on Cb5 dorsodistally, medial edgeanterior to OP, dorsally on Cb4 dorsodistally.SOD very broad, similar to that of Bathymaster(Plate 178), except diagonal muscle strap passes toopposite side, and muscle slip passes through RD onone side, similar to Nemipterus (Plate 154).RDs crossed and joined at origin and only brieflyexposed before separating and passing ventral toSOD, but extending well anteriorly before attachingto Pb3 dorsoanteriorly, meeting OD3 attachment pos-teriorly.Additional remarks. SCL free from posterior endof Bb3, which is not elongate. TV4 free from Cb5s.Pbl absent. Pb2 toothed. Pb4 and UP4 present. IACabsent Eb4 levator process absent. Medial end of Eb3larger than that of Eb4. PCI attaches laterally wellmedial to distal end of Cb5. Ebl anterior process
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absent (interpreted as such based on attachment ofmedial end of Ebl to anterior tip of Pb2).PHOLIDAENot examined.Imamura and Yabe (2002:fig. 14B, 118) illustratethe dorsal gill-arch musculature of Pholis nebulosa(Temminck and Schlegel), but only describe Ad 1?3and a circular TPb2 (only indicated as part of TDA).LEI -4, LP, LI 1-2, TEb2 attaching both anterior andposterior to LE2, OD (but not whether it attaches toboth Eb3 and Eb4), TDP with attachment at least toEb3, OP with dorsal attachment to Eb4 medial toLE4. A mid-longitudinal raphe divides TPb2 andTEb2. They also illustrate a reduced, cartilaginousPbl and a cartilage-tipped Ebl uncinate process,which are the only incidences of the presence of thesetwo elements in the zoarcoids we examined. Theyillustrate a ligament attaching to the cartilaginous tipof the Eb 1 uncinate process and extending medially,presumably, to Pb2. Pholis appears to be less spe-cialized than the stichaeid Vivaria.STICHAEIDAEVivaria subbifurcata (Storer), USNM 364344, 95.4mm. Plate 179
Description.LEI on Ebl mid-dorsoposteriorly (anterior processand Pbl absent; uncinate process present).Remarks. Anterior process assumed absent be-cause medial end of Ebl joins dorsal end of Pb2.LE2 on Eb2 mid-dorsoposteriorly.LE3 on anterior edge of tip of Eb3 uncinate pro-cess, meeting anteroventrally with OD3-4.LE4 on Eb4 bony surface dorsoposterolaterally.LP on posterolateralmost bony edge of Eb4, con-tiguous with posterior edge of LE4 insertion.LI1 on dorsal surface of Pb2 and anteriormostedge of Pb3, overlying, but unattached to, medial endof Ebl uncinate process.LI2 on Pb3 dorsolaterally, joining raphe mediallywith TPb3-Eb3.TD comprises TEb2, TPb3-Eb3, and vestigial rem-nant of TPb2, present only unilaterally as fine, lat-erally curving, semicircular muscle strand arisingdorsoanteriorly from, and dorsoposteriorly re-enter-ing, TEb2 (not connected to Pb2). TEb2 broad, withmid-longitudinal raphe, which gives rise dorsally toCT sheets covering muscles, attached mid-anteroven-trally to CT of pharyngeal roof, attaching on Eb2dorsally anterior to LE2 insertion, joining raphe withmedial end of Ad2, continuous posteriorly by diag-onal strand of muscle with TPb3-Eb3. TPb3-Eb3 onPb3 dorsally joining raphe with medial edge of LI2
insertion and on posterior corner of medial end ofEb3, continuous posteriorly by diagonal musclestrand with SOD.OD3-4, OD3' origin on Pb3 dorsoanteromediallyventral to TEb2, insertion on Eb3 uncinate processdorsoanteriorly and Eb4 uncinate process ventrome-dial to tip; OD3' splits off ventrally from OD3-4 justposterior to origin and inserts on Eb3 dorsoanteriorlyventral to uncinate process, joins raphe with medialend of Ad3.OP on posterior surface of Eb4 at and little lateralto uncinate process, ventrally on Cb5 posterolaterallyposterior to Ad5, medially partially continuous withSO.Adl weak, on anterodistal bony surface of Ebl anddorsoanterior end of Cbl just ventral to Ebl-Cbljoint.Ad2 on Ebl dorsolaterally. joining raphe with lat-
eral end of TEb2, and on anterior surface of Cb2 justventral to Eb2-Cb2 joint.Ad3 on dorsoanterolateral surface of Eb3, joiningraphe with lateral end of OD3', and on anterior sur-face of Cb3 just ventral to Eb3-Cb3 joint.Ad4 dorsally on Eb4 posteriorly ventral to LP, ven-trally on Cb4 dorsolaterally medial to Eb4-Cb4 joint.Ad5 on posterolateral surface of Cb4 (extendingonto cartilage tip on one side, but not the other) anddorsolateral surface of Cb5 anterior to OP.SOD very broad.RDs adjacent.Additional remarks. SCL very fine (damage pre-cluded determining whether it is attached to Bb3,posterior end of which is slightly curved ventrally,usually an indication of such attachment). TV4 freefrom Cb5s. Pb4 and UP4 present. Pbl and IAC ab-sent. Pb2 toothed.
NotothenioideiBOVICHTIDAE
Bovichtus veneris (Sauvage), USNM 200412, 107mm. Plate 180
Description.LEI on raised posterior edge of Ebl anteriorly, atabout mid-length of Eb 1 . No uncinate process.LE2 on raised posterior edge of Eb2 anteriorly, atabout mid-length of Eb2.LE3 on tip of Eb3 uncinate process anteriorly.LE4 broadly on Eb4 dorsoposteriorly beginninglateral to uncinate process.LP on Eb4 joining LE4 insertion posterolaterally.LI 1 mainly on Pb2 dorsoposterior to dorsoanteriorprocess with tendinous attachment ventromedially toanteriormost tip of Pb3.
212 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LI2 on Pb3 dorsally, medial edge of insertionmeeting lateral edge of TPb3-Pb4.TD comprising TEb2 and TPb3-Pb4. TEb2 broadcentrally, narrowing laterally and attaching on Eb2dorsally anterior to LE2 insertion, meeting medialend of Ad2; with mid-longitudinal raphe giving riseto CT sheets covering TD; anterior edge of muscle,between short tendinous raphe on each side of mid-longitudinal raphe, attaching tightly to CT of pha-ryngeal roof; attaching by filmy CT along mid-lon-gitudinal raphe ventrally between Pb3s to CT to pha-ryngeal roof; not continuous posteriorly with TPb3-Pb4. TPb3-Pb4 beginning on Pb3 dorsally alongmedial-edge of LI2 insertion and continuing poste-
riorly onto medial edge of Pb4 ventral to joint withmedial end of Eb4 ventrally, continuous posteriorlyby diagonal muscle strap with SOD.OD3-4, OD3' origin broadly on Pb3 dorsomedi-ally ventral to TEb2, insertion on Eb3 uncinate pro-cess anteriorly continuing onto Eb4 uncinate processposteromedially, there joining raphe with OP dorso-lateral^; muscle branches (OD3') ventroanteriorlyjust before insertion on Eb3 uncinate process and in-
serts on Eb3 dorsally well ventromedial to uncinateprocess, meeting lateral end of Ad3.OP dorsally on Eb4 posteriorly beginning on un-cinate process (joining raphe with OD3-4) and ex-tending medially to medial end, fibers of medial halfpassing posteroventrolaterally over those of lateralhalf and together attaching broadly on Cb5 postero-laterally; ventrodistally tendinously joining toughfascia (not illustrated) covering Ad5 surface; musclemedially not clearly separated from SO.Ad 1-3 moderately developed. Each beginning onEb dorsoanterolaterally and extending laterally thenventrally over Cb dorsoanteriorly; Ad2 medial endjoining raphe with TEb2; Ad3 medial end just meet-ing lateral end of OD3'.Ad4 broadly dorsally on Eb4 ventrally, dorsome-dial half overlapped posteriorly by OP, ventrallybroadly on Cb4 dorsally medial to Eb4-Cb4 joint.Ad5 posteriorly covered by tough fascia, dorsallyon Cb4 posterodistally with tendinous connection ex-tending to Eb4 posterodistally (surface of Eb4-Cb4joint covered by tough fascia (not illustrated), whichis continuous with fascia covering Ad5, ventrally onCb5 mostly anterior to OP.SOD present.RDs separated by distance less than half one RDdiameter.Additional remarks. SCL present. TV4 free fromCb5s. Pbl absent. Pb2 toothed. IAC absent. Pb4 pres-ent. UP4 present. Eb4 levator process absent.PSEDUAPHRITIDAEPseudaphritis urvillii (Valenciennes),344898, 97.2 mm. USNM
Not illustrated
Description.LEI on anterior surface of raised bony process onEbl mid-dorsoposteriorly (uncinate process absent).LE2 on anterior surface of raised bony process onEb2 mid-dorsoposteriorly.LE3 on tip of Eb3 uncinate process anteriorly.LE4 on Eb4 dorsoposteriorly lateral to uncinateprocess.LP on Eb4 beginning at posterior edge of LE4 in-sertion and extending well laterally.LI1 on Pb2 dorsoanteriorly.LI1' on Pb3 dorsoanterolaterally just ventral to an-terolateralmost edge of OD3-4.Remarks. Presence of LI1 and LI1' very similar tothese two muscles in some gobioids.LI2 on Pb3 dorsoposteriorly medial to medial endof Eb3.TD comprises TEb2, TPb4a, and TPb4. TEb2 withmid-longitudinal raphe, attached anteroventrally toCT extending anterior to gill arches, raphe giving riseto broad CT sheets covering TD; muscle extendinglaterally onto Eb2 anterior to LE2 insertion, contin-uous at posterior end of raphe with TPb4a. TPb4aslender, extending laterally, passing through OD3?
4
and attaching ventrally to Pb4 at and lateral to an-terolateral attachment of TPb4, continuous postero-laterally by fine, diagonal muscle filament with TPb4.TPb4 relatively broad, on Pb4 medialmost edge atjoint with Eb4, questionably with extremely fine ten-dinous attachment to posteromedialmost edge of Eb3adjacent to Eb4; continuous posteriorly by fine,crossing diagonal muscle fibers with SOD.Remarks. This is the only taxon we encounteredin which a TD muscle passes through OD3?4. Ver-ification in another specimen is desirable. OD3-4 insome forms (e.g., Odontobutidae) is split by LI2.OD3-4 origin on Pb3 broadly dorsally ventral toTEb2, anteriorly just medial to LI1' insertion, pene-trated by TPb4a just after OD3-4 passes posteriorlyfrom under TEb2, insertion beginning on Eb3 unci-nate process anterodorsally and continuing mediallyto medial edge of Eb4 uncinate process. (See remarksfollowing TD.)OP dorsally on Eb4 posteroventrally beginning be-low uncinate process and extending laterally to pointbelow LP insertion, ventrally on Cb5 dorsopostero-laterally, posterior to Ad5 medially.Ad 1-3 absent, but weak GFI on first arch antero-laterally; on arches 2 and 3, GFI begins on Eb dor-soanterior surface and passes to anterior edges of Eband Cb; GFI best developed on Eb3.Ad4 dorsally broadly on Eb4 ventrally, beginninganterior to OP laterally and extending laterally toEb4-Cb4 joint, ventrally on Cb4 broadly.Ad5 dorsally on Cb4 posterolaterally, ventrally on
NUMBER 11 213Cb5 beginning dorsoposterodistally and extendingmedially short distance anterior to OP.SOD present.RDs separated by space equal to less than half di-ameter of one RD.Additional remarks. SCL attached tendinouslymid-dorsoposteriorly to posteroventral cartilaginousend of Bb3. TV4 free from Cb5. IAC absent. Pblabsent. Pb2 toothed. Pb4 and UP4 present. Eb4 le-vator process absent.Pseudaphritis is sometimes included in the Bovi-chtidae, but recent studies (Balushkin, 1992; Eastmanand Clarke, 1998) do not support this inclusion.
DactylopteroideiImamura (2000) excluded the Dactylopteridaefrom the Scorpaeniformes based on several charac-ters and included them with the Malacanthidae in anexpanded family Dactylopteridae. He hypothesizedthe latter action based on eight putative synapomor-phies. His first appears to be the most significant,nasals of dactylopterids are fused in both larvae andadults and the nasals of malacanthids [and holocen-trids] are fused in larvae, but separate later. He basedthe malacanthid information on Johnson (1984:491et seq.). Imamura, thus, equated fusion of the nasalbones with subsequent separation, as found in ma-lacanthids, with fusion of nasal bones without sub-sequent separation, as found in dactylopterids. Webelieve that this consideration needs further study;however, for convenience, we maintain Imamura'sassociation of the two families by including them ina single suborder.Imamura's synapomorphies 2-6, and 8 (absence ofsupramaxillary, absence of prevomerine teeth, ab-sence of palatine teeth, presence of six branchiostegalrays, absence of toothed plate on Eb2, presence of aTDA circular element) have a wide and varied dis-tribution and we believe should be part of a largercharacter-based test of interrelationships. He reportedthat his seventh synapomorphy, adductor mandibulaesection 2 subdivided, is "a rare synapomorphy"based on its absence in members of 18 perciformfamilies (he reported 16 families, but we recognizetwo additional included among his material) and "75species of Scorpaeniformes . . . listed by Imamura(1996)." We are unable to comment on the seventhsynapomorphy, but note there are a great many morefamilies that should be evaluated for it.
DACTYLOPTERIDAE
Dactyloptena macracantha (Bleeker), USNM224473, 92.7 mm. Plate 181
Description.LEI broadly on Ebl mid-dorsally.LE2 on Eb2 mid-dorsally.LE3 on Eb3 uncinate process anteriorly just ven-tral to OD3-4 insertion, which is on cartilage tip ofprocess anteriorly.Remarks. This is the only instance we know of inwhich LE3 inserts on Eb3 ventral to OD3-4 insertionon Eb3. The condition should be verified in anotherspecimen.LE4 on Eb4 dorsoposterolaterally.LP on Eb4 anterolateral to LE4 insertion.LI1 On Pb3 dorsoanterolaterally.LI2 on Pb3 dorsoposterolaterally just medial tomedial end of Eb3.TD comprises TPb2, TEb2, and TPb3-Eb3. TPb2flat, laterally curving, semicircular ribbon of muscleon each side dorsal to TEb2; joined to contralateralTPb2 anteromedially by broad wedge of CT whichnarrows to raphe posteriorly; muscle arising fromTEb2 anteromedially, becoming fine tendon poster-omedially attaching to raphe and contralateral TPb2;attached anterolaterally by CT to dorsoanterior endof Pb2; mid-longitudinal CT areas attached ventrallyto CT of pharyngeal roof, giving rise dorsally tofilmy CT sheets covering muscles. TEb2 separatedfrom counterpart by same CT areas as TPb2; extend-ing laterally and fanning out more-or-less verticallyand attaching to Eb2 anterolaterally lateral to LE2origin, meeting and attaching partially ventrally tobroad Ad2 medially, connected posteroventrally byvery fine filaments of muscle to TEb3. TP3-Eb3 fine-ly tendinously on dorsomedial surface of Eb3 and onPb3 dorsoposterolaterally just posterior to LI2; pos-teriorly joined by fine crossing strands of muscle withSOD.OD3-4 origin broadly dorsomedially on Pb3 ven-tral to TEb2, insertion on Eb3 uncinate process dor-soanteriorly just dorsal to LE3 insertion and on me-dial edge of Eb4 uncinate process.OP broadly dorsally on Eb4 ventrally beginningventral to LE4 insertion and extending medially,broadly ventrally on Cb5 dorsoposteriorly medial todistal end, medially inseparable from SO.Ad 1-3 unusually well developed.Adl extensively on Ebl beginning on uncinateprocess laterally and extending laterally, becomingfan-like and attaching to anterior surface of Eb 1 -Cb 1joint.Ad2 on Eb2 anterolaterally, overlying lateralmostend of TEb2 and extending laterally to anterior sur-face of Eb2-Cb2 joint.Remarks. It is unusual for Ad2 to overlie the lat-
eral end of TEb2.Ad3 beginning on dorsal surface of Eb3 medial touncinate process and on Eb3 uncinate process lateral
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to LE3 insertion, and extending laterally and fanningout over anterior surface of Eb3-Cb3 joint.Ad4 dorsally broadly on Eb4 ventrally mostly lat-eral to OP, ventrally broadly on Cb4 medial to Eb4-Cb4 joint.Ad5 on posterolateral end of Eb4 and dorsolateralend of Cb5, medially anterior to OP ventrally.SOD present.RDs separated by space less than diameter of oneRD. Right-side RD incompletely divided longitudi-nally, joint cross section noticeably larger than thatof left side RD.Remarks. In Imamura's (2000:fig. 9B) illustrationof the RDs of Dactyloptena macracantha, the di-ameter of the left-side RD is noticeably larger thanthat of the right side. Asymmetry of RD size is com-mon in fishes, although usually not differentially sogreat as in Dactyloptena.Additional remarks. SCL attached mid-dorsally bytendon to posteroventrally extending cartilaginous tipof Bb3. TV4 free from Cb5s. Pbl cartilaginous. UP4present, Pb4 absent, but present in Dactlopterus vol-itans (Linnaeus), based on USNM 261386, clearedand. IAC absent. Pb2 toothed. Eb4 levator processabsent.Remarks. Imamura (2000) described some aspectsof the dorsal-gill arch musculature of dactylopteridsand presented a diagrammatic illustration of the mus-culature of D. macracantha.Imamura listed several perciform groups, besidesdactylopterids, in which TPb2 is circular (he was un-certain the muscle was homologous with TPb2) andindicated that the condition is a percoid [percomorph]apomorphy. The muscle state is much more widelydistributed than Imamura recognized; it also occurs,
e.g., in Chaunacidae (Paracanthopterygii), Bathymas-teridae (Zoarcoidei), and Centrolophidae (Stromateo-idei), among several other groups. Because of thevarious states of TPb2 present in a wide variety ofacanthomorphs, the derivation of the circular TPb2.including the loss of its attachment to Pb2, cannot bedecided readily.MALACANTHIDAE
Caulolatilus affinis Gill, USNM 211424, ca. 98 andca. 100 mm. Plate 182
Additional material. ? = Malacanthus brevirostrisGuichenot, USNM 334628, 150 mm.
Description.LEI on Ebl uncinate process at or just lateral tojoint with IAC.LE2 on bony prominence rising from Eb2 mid-dorsoposteriorly and continuing posteriorly onto mid-anterior edge of Eb3; tendon extends length of mus-
cle surface laterally, attaching to bony prominenceand continuing posteriorly and attaching to anterioredge of Eb3.Remarks. Imamura (2000:214) reported that the in-sertion of LE2 on Eb2 and Eb3 is restricted to ma-lacanthids and nemipterids among the perciforms heexamined. We find that in our specimens of Nemip-terus there is a ligament attaching the posterior edgeof Eb2 at the posteroventral margin of the LE2 in-sertion to the anterior edge of Eb3. We do not equatethese two conditions. In malacanthids, LE2 splits itsinsertion between the closely adjacent margins ofEb2 and Eb3. such that if the bones are forced apart,LE2 splits longitudinally, with one part remainingwith Eb2 and the other with Eb3. In nemipterids, theentire muscular insertion of LE2 in on Eb2.LE3 on Eb3 uncinate process anteriorly.LE4 on Eb4 beginning on levator process dorsallyand extending laterally and meeting LP insertion an-teriorly. ? Like Caulolatilus, but insertion broadly,ventroanteriorly meeting OD4.LP on Eb4 at and joining LE4 insertion posteri-orly.LI1 on Pb2 dorsally just posterolateral to joint withIAC.LI2 on Pb3 dorsolaterally medial to medial end ofEb3, ventromedial edge meeting TPb3-Eb3 laterally.TD comprises TPb2, TEb2, and TPb3-Eb3. Thick,cup-shaped CT pad arises mid-dorsally from mid-longitudinal raphe extending across TPb2 and TEb2;pad continuous anteriorly with pharyngeal roof CT.TPb2 a bilateral pair of roughly semicircular musclesdorsal to TEb2 medially; each member of pair at-taching anteromedially to CT covering dorsoanter-iormost end of Pb2 process articulating with IAC andcontinuous ventrally with CT of pharyngeal roof;posteromedially each member fades into TEb2 andmid-longitudinal raphe. TEb2 attaches to Eb2 alongdiagonal beginning posteriorly at medial edge of LE2insertion and extending anterolaterally to point nearlateral edge of CT (not illustrated) attaching Eb2 toEbl (lateral to LE2 insertion). TPb2-TEb2 not con-tinuous with TPb3-Eb3. TPb3-Eb3 laterally ventralto OD3?4, with mid-longitudinal raphe; muscle intwo layers: dorsal and ventral; both layers attach to-gether laterally on Pb3 dorsolaterally at medial edgeof LI2 insertion; fibers of dorsal layer with slightchange in orientation between portions attaching toPb3 and Eb3 (change not obvious in smaller speci-men); dorsal layer continues posteriorly attaching toposteromedial margin of Eb3; ventral layer extendsposteriorly beyond mid-posterior margin of dorsallayer (ventral layer much less discrete in smallerspecimen and absent in
?); muscle not continuousposteriorly with SOD (SOD absent, probably anom-alously, in smaller specimen). ? TPb2-TEb2 contin-uous posteriorly with SOD; TPb3-Eb3 muscle strap
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attaching dorsally to Eb3 along posteromedial edge(as in Caulolatilus), but ventrally attaching to Pb3dorsoposteriorly posterior to Pb3-Eb3 joint.CPb fine muscle thread, easily overlooked, sepa-rated by thin epithelial tissue from Pbs, extendingposteriorly from Pb2 anterolaterally to UP4 laterally,with short, fine branch extending medially anterior toPb3 and another medially anterior to UP4; medialbranches absent in smaller specimen. ? Slightly bet-ter developed, but beginning as continuation from an-teriorly extending SO longitudinal fibers medial toPbs, extending anterolaterally between Pb2 and Pb3(no extension anterior to Pb2), then posteriorly alonglateral surfaces of Pb3 and UP4, with medial branchextending anterior to UP4.Remarks. Of those taxa possessing CPb (Table 9),it is most poorly developed in malacanthids, andprobably vestigial; considerable variation in its man-ifestation is probably to be expected.OD3. OD4 originate as slender CT pad attachingalong Pb3 dorsomedial edge; posterior half of originis ventroposterior to TPb2 and TEb2 in larger spec-imen, but completely ventral to TPb2 and TEb2 insmaller specimen. Muscle divides into OD3 and OD4well before inserting on Eb3 dorsoanteriorly just ven-tral to tip of uncinate process and on Eb4 dorsopos-teriorly, beginning about midway between medialend and levator process and extending laterally tomedial edge of cartilage tip of levator process; tinycartilaginous tip of Eb4 uncinate process sandwichedbetween, partially overlain by, and obscured fromview by OD3 and OD4 muscle insertions. ? OD3and OD4 separate almost immediately posterior toinsertion; OD4 passes dorsal to all bony Eb4 uncinateprocess; OD3' present (anomalously?) on one side,separates ventrally from joint origin of OD3 and OD4and inserts on Eb3 dorsally well ventral to lateral endof OD3.OP dorsally on Eb4 posteriorly beginning near me-dial end and extending laterally to below levator pro-cess, overlapping medial fibers of Ad4 posteriorly;ventrally, broadly on Cb5 beginning well mediallyalong PCI attachment and extending laterally to justmedial to distal end.Ad 1-3 absent, but GFMs 1-3 (not illustrated) welldeveloped; on anterior margins of Ebl and Cbl, butbeginning on dorsoposterolateral surfaces of Eb2 andEb3 and attenuating distally and attaching along an-terior surface of respective Cb.Ad4 dorsally on Eb4 posteroventrally, beginningmedially below levator process and anterior to dor-solateralmost OP fibers and extending laterally al-most to distal end; ventrally on Cb4 dorsoposteriorlybeginning medially a short distance from lateral endand joining Ad5 attachment and extending laterallyto Eb4-Cb4 joint medially.Ad5 beginning dorsally on posteriorly extending
cartilaginous process at distal end of Cb4 and con-tinuing a short distance on Cb4 medially, meetingAd4 ventrally; ventrally on Cb5 dorsodistally extend-ing a short distance medially anterior to lateralmostOP fibers.SOD present, well posterior to TPb3-Eb3, moder-ately wide (absent, probably anomalously, in smallerspecimen). ? SOD broad, joined mid-anteriorly toTPb3-Eb3.RDs adjacent; separated by distance half diameterof one RD in smaller specimen and ?.Additional remarks. SCL attached mid-dorsally toelongate ventroposteriorly extending cartilaginousposterior end of Bb3 (only ObV3 attaches to it). TV4free from Cb5s. Pb2 toothed. Pb4 and UP4 present.
CallionymoideiDRACONETTIDAEDraconetta oregona Briggs and Berry, USNM159234, 110 mm. Plate 183
Description.LEI on raised mid-dorsoposterior bony edge ofEbl (no uncinate process).LE2 on mid-dorsoposterior bony edge of Eb2.LE3 tendinously on dorsal edge of Eb3 lateral touncinate process.LE4 massive, dorsally on expanded posterior sur-face of Eb4 well lateral to joint with Eb3 uncinateprocess.LP on dorsal surface of Eb4 between lateral edgeof LE4 insertion and distal end of Eb4.LI1 on dorsoanteriormost surface of Pb3, whichoverlies all of dorsal surface of Pb2 except for an-teriormost cartilaginous end.LI2 on Pb3 posterolaterally medial to medial endof Eb3.TD broad, continuous, comprising TEb2, TEb3,and, very doubtfully, TPb2. Questionable TPb2 athin, slender posterolaterally convex strip of musclebordering TEb2 anteriorly, widening laterally as itextends dorsal to TEb2 and inserts on Eb2 amongTEb2 fibers as they pass laterally onto Eb2; not at-tached to Pb2, see remarks). TEb2 with mid-longi-tudinal raphe continuing posteriorly on all but pos-terior section of TEb3; raphe giving rise to filmy CTcovering TD; TEb2 attaching mid-anteriorly to CT ofpharyngeal roof and amid sparse, anteriorly extend-ing longitudinal muscle fibers, attaching anteroven-trolaterally to dorsoanterior ends of Pb2 and Pb3 (un-usual for TEb2 to attach to Pb2); as muscle extendslaterally, anterior fibers twist posteroventrally overposterior fibers and extend onto Eb2 dorsally to at-tach anterior to LE2 insertion, joining raphe with me-dial end of Ad2; posterior fibers attach to posterior
216 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
edge of Eb2 medial to LE2 insertion; continuous pos-teriorly with TEb3. TEb3 on Eb3 dorsoposterome-dially ventral to OD3-4 insertion, continuous poster-oventrally by fine muscle filament with SOD.Remarks. Most probably the questionable TPb2 ispart of TEb2 and is treated as such for cladistic pur-poses. Attention is drawn to it here because TPb2 ispresent in the Callionymidae, which is the sistergroup to Draconettidae.CPb comprising subcutaneous filaments of muscledifficult to free from skin, simple strip passing aroundPb2 on each side and continuing down lateral andmedial margins of Pb3, then meshing with matrix ofsubcutaneous muscle extending posteriorly from Pb3and becoming SO.Remarks. The presence of CPb is evidence forclose relationship with Callionymidae, which has afar more complex development of dorsal gill-archmusculature that obscures the long-accepted sister-group relationship between the Callionymidae andDraconettidae.OD3-4 origin on Pb3 dorsomedially ventral toTEb2, insertion on anterior surface of Eb3 uncinateprocess and posterior surface of Eb4 where Eb4 ar-ticulates with Eb3 uncinate process.OP on most of posterior surface of Eb4, fused in-distinguishably ventrolaterally with Ad5, broadly onCb5 dorsally continuous with Ad5, distinguishablemedially from SO mainly by abrupt change fromthickness of OP to very thin SO.Adl attaching on anterior surface of Ebl just an-teromedial to medial edge of LEI and extending lat-
erally, fanning out distally, and attaching across an-terior surfaces of distal ends of Eb 1 and Cb 1
.
Ad2 with anterior and posterior sections, anteriorsection on Eb2 dorsally meeting lateral end of TEb2,posterior section dorsally meeting anterior edge ofLE2 insertion, sections fuse laterally, fan out and at-tach across anterior surfaces of distal ends of Eb2and Cb2.Ad3 with anterior and posterior sections, anteriorsection on Eb3 dorsally, posterior section dorsallymeeting anterior edge of LE3 insertion, sections fuselaterally, fan out and attach across anterior surfacesof distal ends of Eb3 and Cb3.Ad4 (obscured in posterior view) dorsally broadlyon Eb4 ventral surface anterior to OP, ventrallybroadly on Cb4 dorsal surface medial to Eb4-Cb4joint.Ad5 completely fused medially with OP (includingattachment to Cb5), attaching anteriorly to poster-odistal surfaces of Eb4 and Cb4.SOD very slender.RDs adjacent.Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Tiny cartilaginous Pbl attached to medial
tip of Eb2. Pb4 and UP4 absent. Eb4 cartilage-tippeduncinate and levator processes absent.
CALLIONYMIDAECallionymus lyra Linnaeus, USNM 197584, 3 spec-imens, 86.3-128 mm.Plates 184.1, 184.2
Additional material. ? = Callionymus filamentosusValenciennes, USNM 232253, 68.7 mm.
Description.Remarks. Kayser (1962) described and illustratedthe gill-arch musculature and skeleton of C. lyra. Al-though we generally agree with his findings, his de-scription is brief and does not do justice to the ex-treme complexity exhibited by these muscles in Cal-lionymus. Additionally we report a muscle (CPb) hedid not include. Winterbottom (1974b:fig. 23) pro-vided a redrawn version of one of Kayser's figures.LEI on Ebl dorsodistally, joining tough CT rapheventromedially with distal end of M. Pb3-Ebl.LE2 on dorsodistal end of Eb2 meeting RecD3dorsoposteromedially and M. Pb3-Eb4-Eb2-Cb3 dis-tally.LE3 absent (see LE4).LE4 massive, inserts on CT binding distal ends ofEb3 and Eb4 and is joined there by posterior end ofRecD4 and anterior end of Ad5; posteroventrallyjoins LP anteroventrally (see LP for variation in ?).Remarks. Either LE3 and LE4 are fused or LE3has been completely lost; we arbitrarily opt for thelatter interpretation because the general configurationexhibited by LP and the questionable muscle is likethat of LP and LE4 of many fishes; Kayser (1962)assumed that LE3 and LE4 are fused.LP joins LE4 insertion posteriorly on Eb4 andjoins Ad5 dorsally. ? LP and LE4 joined tendinouslyand only lateralmost edge of LE4 portion attached toEb4 near distal end; muscles together forming
"sling" (Stiassny and Jensen, 1987:284), with tendonfrom LP extending ventrally and attaching to Cb5anterior to OP; Ad5 absent.LI1 on Pb3 broadly dorsoanterolaterally at anddorsal to M. Pb3-Ebl origin.LI2 on Pb3 dorsoposteriorly medial to M. Pb3-Eb4-Eb2-Cb3 origin on Pb3.TD comprises TEb2, TPb2, and TEb4. TEb2 thin,bilateral muscle pair, continuous membranously dor-
sal to Pb3s and from mid-longitudinal pharyngealroof muscle fibers, attaching sheet-like along pos-terolateral surface of Eb2. TPb2 a bilateral pair ofsemicircular ribbons attaching to CT surrounding an-terior edges of Pb3 and Pb2 (Pb2 almost entirely ven-tral to Pb3), fusing posteromedially with TEb2 andTEb4. TEb4 comprising anterior and posterior por-
NUMBER 11 217
tions; anterior portion with mid-longitudinal raphe,posterior portion uninterrupted, both portions fusingtogether laterally and inserting on Eb4 dorsomediallyventral to OD4; joined by CT mid-ventrally to SODmid-dorsally when SOD is present.CPb originates as SO longitudinal fibers extendinganteriorly dorsal to pharyngeal roof tissue, dividingwell posterior to gill arches with branch on each sideextending around Pb2 and Pb3 and attaching antero-lateral^ to Eb2 and Pb2 ventrally, where the twobones meet, and to Pb3 posterolaterally.OD3 absent.OD4 origin on Pb3 dorsally ventral to TEb2, in-sertion on bony Eb4 uncinate process dorsoanteriorly.M. Ebl-Cbl attaches dorsomedial end of Ebl todorsoanterodistal end of Cbl; muscle joins CT sur-rounding distal end of Cbl, which is also joined byanterior end of RecD2.Remarks. The skeletal elements to which M. Ebl-Cbl attaches are the same as those that define Adlin other acanthomorphs. but the position of the Adlattachment on Ebl is closer to the distal end of thebone and spans the Ebl-Cbl joint, very unlike M.Ebl-Cbl.M. Pb3-Ebl origin on Pb3 anterolateral^ at andventral to lateral edge of LI1 insertion, insertion onposterodistal edge of Ebl, there joining raphe withLEI insertion and raphe of RecD2 with RecD3.M. Pb3-Eb4-Eb2-Cb3 origin beginning on Pb3dorsoposterolaterally and continuing onto Eb4 dor-somedially along attachment of TEb4, insertion alongposterior edge of LE2 insertion on Eb2 and with ra-phe joining RecD3 and RecD4 to dorsodistal end ofCb3.OP dorsally on posterior surface of bony Eb4 un-cinate process, ventrally on Cb5 posterolateral sur-face, joining small raphe with Ad5 ventrolaterally atdorsodistal end of Cb5.Ad 1-3 absent.Ad4 dorsally on Eb4 posterolaterally (mostly over-lapped posteriorly by OP and Ad5), ventrally on Cb4dorsally medial to Eb4-Cb4 joint.Ad5 posteriorly on Cb5 dorsodistally, dorsoanter-olaterally attaching joining distal ends of Eb4 andEb3 and to which LE4 inserts, dorsoanteriorly joiningraphe with LP insertion, ventroanteriorly fusing in-distinguishably with RecD5 (see also Remarks pref-acing RecD2). ? Ad5 absent.Remarks. Kayser (1962:413; fig. 35) treated themuscle we identify as Ad5 as a fusion of two muscles
"Museums pharyngo-arcualis I + II." He illustratedthe muscle as being divided by a raphe into right andleft halves. Based on his illustration of Ammodytesin the same paper (1962:fig. 15), he considered thefused muscle to comprise the muscles we designateas Ad5 and OP. Although we agree that Ad5 is in-volved in a fusion, we find that the composition is
Ad5 + RecD5, and the direction of the fusion is an-terior-posterior. OP is distinct. We were only able toresolve the composition of the fusion in C. lyra be-cause Ad5 is completely absent in C. filamentosus,but RecD5 is distinct and very similar to RecD2?4.RecDs. The next four muscles are designated as
recti dorsales. a name coined by Winterbottom(1974b:259) to apply to muscles that interconnectepibranchials of successive arches. He included sim-ilar muscles in Callionymus lyra, although the con-nections do not agree entirely with his definition, aremore complex, and are possibly synapomorphic forthe family. In the two larger specimens in USNM197584, the RecDs appear to be individual muscles.In the smallest specimen RecD3 and 4 each appearsto comprise two muscles (Plate 184.2D), a RecD andan underlying muscle, RecCb, attaching the distalends of successive Cbs. In all three specimens RecD5is fused posteriorly with Ad5 and appears to includea fused RecCb anteroventrally. In the specimen of C.filamentosus, we only confidently observed RecCbsbetween arches 2 and 3 and 3 and 4. The RecDs ofCallionymus do not appear to be homologous withthose of the only other acanthomorph, Mene, inwhich they are present.RecD2 anteriorly on distal end of Cbl, there join-ing raphe with M. Ebl-Cbl, posteriorly joining raphewith and anterior end of RecD3 on distal ends of Ebland Cb2.RecD3 anteriorly joining raphe with posterior endof RecD2 on Ebl and Cb2 and posteriorly joiningraphe with anterior end of RecD4 on distal ends ofEb2 and Cb3.RecD4 anteriorly joining raphe with posterior endof RecD3 on distal ends of Eb2 and Cb3, and pos-teriorly joining raphe with anterior end of RecD5 ondistal ends of Eb3, Eb4, and Cb4.RecD5 anteriorly on distal ends of Eb3, Eb4, andCb4, posteriorly fusing with Ad5 on Cb5 anterodis-tally. ? Posteriorly on anterodistal end of Cb5; Ad5absent.SOD present in only one of three specimens (113mm SL); very thin, slender, completely ventral toTEb4, with median raphe attaching to ventral surfaceof TEb4. ? Absent.RDs separated by distance less than half diameterof one RD.Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Pb4 and UP4 absent. Pb2 toothed. IACabsent. Cartilage tipped Eb4 uncinate and levatorprocesses absent.Kayser (1962:412) designated the muscles we callM. Ebl-Cbl, M. Pb3-Ebl, and M. Pb3-Eb4-Eb2-Cb3as M. obliquus inferior 1-3. Despite their differentattachments, Kayser considered them to be serial ho-mologs and different from OD4, which he also rec-ognized in C. lyra. Kayser did not report the presence
218 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of OD3 in C. lyra, and we note that it is absent.Winterbottom (1974b:253-254) considered a varietyof bilaterally paired transverse muscles joining Pbsto Ebs as ODs. He included Kayser's obliquui infer-iores among them. The muscle in Callionymus thatwould be interpreted as OD3 does not include Eb3among its attachments and we do not consider it tobe homologous with OD3 in other actinopterygians.The presence and attachments of OD3 and OD4 inactinopterygians are so consistent that we reserve theterms OD3 and OD4 for them.Usually, PCI is attached to Cb5 musculously or byshort tendon, but in Callionymus and Trachelochis-mus (Gobiesocidae) it is attached by a long, slendertendon. These two families, together with the Dra-conettidae, were hypothesized by Gosline (1970) toform a natural group.GOBIESOCIDAETrachelochismus sp., USNM 339196, 57.1 mm.Plate 185
Description.LEI on Ebl dorsally at about mid-length; uncinateprocess absent.LE2 on Eb2 posteriorly near distal end.LE3 absent.LE4 on Eb4 dorsodistally.LP on Eb4 at and posterior to LE4 insertion.LI1 on Pb3 dorsoanteriorly.LI2 on Pb3 dorsally lateral to TPb3-Eb4 attach-ment.TD comprises TEbl, TEb2, and TPb3-Eb4. TEbla slender filament on medial tip of Ebl, continuousposteriorly with TEb2. TEb2 broad, flat mid-dorsally,with median longitudinal raphe, muscle narrowinglaterally, attaching to Eb2 dorsomedially, and poste-
riorly continuous with Eb4 portion of TPb3-Eb4.TPb3-Eb4 delaminating (bilaterally) from postero-ventral surface of TEb2, extending laterally, and hav-ing two separate attachments to Pb3, one curving an-teriorly, passing ventral to OD3-4 origin, and attach-ing to dorsal surface of Pb3, the other extending lat-erally and attaching to Pb3 near medial ends of Eb3and Eb4; Eb4 portion slender, on medial end of Eb4,continuous posteriorly with SOD.Remarks. Although undoubtedly homoplastic,TEbl is known otherwise only in certain labrid taxa.OD3-4 originates on Pb3 and, unusually, on me-dial end of Eb2 anteriorly ventral to TEb2, overlyinglateral attachments of TPb3-Eb4 to Pb3, and insertson dorsoanterior surface of proximal half of Eb3 andposterior edge of proximal quarter of Eb4.OP, if present, inseparable from SO.Adl on mid-anterior surface of Ebl and dorsoan-terior surface of Cbl just ventral to Ebl-Cbl joint.Ad2 on much of distal half of Eb2 anterior surface.
extending anterolaterally and attaching to posterodis-talmost end of Ebl (unusual), anterior surface of Cb2just ventral to Eb2-Cb2 joint, and dorsolateral edgeof CT joining Ebl and Eb2.Ad3 on most of medial half of Eb3 anterior sur-face, extending anterolaterally and attaching to pos-terodistalmost edge of Eb2 (unusual), at and just ven-tral to Eb3-Cb3 joint, and dorsolateral edge of CTjoining Eb3 and Cb3.Ad4 dorsally on Eb4 posteriorly beginning atabout mid-length of Eb4 and extending laterally al-most to end of bone, ventrally on Eb4 for about dis-tance equal to dorsal attachment.Ad5 on anterolateral surface of Cb5, extendingonto posterodistal half of surface of Cb4 and con-tinuing onto posterodistal end of Eb4.SOD present, continuous anteriorly with TPb3-Eb4.RDs separated by space greater than twice diam-eter of one RD.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl. Pb2, Pb4, and UP4 absent. Ebl, Eb3, andEb4 uncinate processes absent. Eb4 levator processabsent. Medial end of Eb3 much larger than that ofEb4. PCI attaches by long slender tendon to distalend of Cb5.
Blennioidei
Remarks. Springer (1993) defined the SuborderBlennioidei and its included six families: Blenniidae,Chaenopsidae, Clinidae, Dactyloscopidae, Labrisom-idae (monophyly not hypothesized), and Tripterygi-idae. He was unable to resolve the intra-relationshipsof the families, but hypothesized that the Triptery-giidae are the sister group of the other five families.Hastings and Springer (1994) hypothesized the intra-relationships of the Chaenopsidae, placing Neoclinusas sister group of the other genera.Although all of the gill-arch specializations thatcharacterize blennioids are found among other acan-thomorph taxa, the number and combination of thesespecializations are distinctive. Specializations in-clude: TPb2 absent (except a vestige in Labrisomi-dae); Pb3 musculously naked dorsoanteriorly; attach-ment of TV4 to Cb5; absence of cartilage tipped Ebl,Eb3, and Eb4 uncinate processes and Eb4 levatorprocess; PCI broadly on Cb5, extending to distal endand joining raphe with OP ventrally (in all familiesexcept Dactyloscopidae, in which the two musclesjust fail to meet). All blennioids lack IAC, Pb2, Pb4,and UP4. LI1 attaches to Pb3, which is also a spe-cialized state; however, Pb2 is absent.The most restricted of these specializations are themusculously naked dorsoanterior surface of Pb3 andthe combination of the attachment of PCI to includethe distal end of Cb5 and the muscle's joining a raphe
NUMBER I I 219
with OP ventrally (we consider failure of PCI to joina raphe with OP in the Dactyloscopidae to be deriv-ative for blennioids). The combined PCI-OP state oc-curs in centrogeniids, sciaenids, nemipterids. pingui-pedids, mastacembelids, lethrinids (Monotaxis andGymnocranius, but PCI just failing to reach distalend of Cb5 in Lethrinus, see description, additionalremarks).
TRIPTERYGIIDAERuanoho decemdigitatus (Clarke), USNM 92.0 mm;USNM 339242, 73.6 mm.Plate 186
Description.LEI broadly on expanded bony dorsoposterioredge of Ebl (no uncinate process).LE2 broadly on expanded bony dorsoposterioredge of Eb2.LE3 anteriorly On dorsoposterior edge of Eb3 allbony uncinate process, joining raphe with dorsolat-eral edge of OD3-4 on Eb3.LE4 large, on Eb4 dorsolaterally, extending to dis-tal end of bone.LP on Eb4 posterior to LE4 insertion.LI1 on Pb3 beginning on anterolateralmost edgeand continuing posteriorly onto dorsoanterior surfaceventral to TEb2 (Pb2 absent).LI2 on Pb3 dorsoposterolaterally medial to medialend of Eb3, aligned along lateral edge of TPb3.TD comprises TEb2 and TPb3. TEb2 a broad bandinterrupted by mid-longitudinal raphe, which givesrise dorsally to filmy CT and is attached ventrally toCT of pharyngeal roof; attaches on Eb2 dorsally wellmedial to LE2, continuous posteriorly by diagonalstrand of muscle with TPb3. TPb3 on bony surfaceof Pb3 dorsolaterally along medial edge of LI2 in-sertion, extending posteriorly opposite to medial endof Eb4.OD3-4 origin on Pb3 dorsally ventral to TEb2,insertion on Eb3 dorsoanterior surface medial to LE3(joining ventromedial edge of LE3) and on mid-dor-sal surface of Eb4.OP broadly dorsally on Eb4 coincident with ven-troposterior edge of LP insertion; broadly ventrallyon Cb5 almost coincident with PCI attachment; me-dially incompletely separable from SO.Adl on Ebl bony surface anterodistally and Cblanteriorly just medial to distal end of bony surface.Ad2 broadly on most of Eb2 bony surface dor-soanteriorly lateral to TEb2, narrowly on anterodis-talmost bony surface of Cb2.Ad3 broadly on Eb3 bony surface dorsoanteriorlyventral to OD3-4, narrowly on Cb3 anterodistalmostbony surface.Ad4 broadly on Eb4 dorsolaterally anterior to OP,
extending very broadly onto Cb4 dorsally medial toEb4-Cb4 joint.Ad5 on Cb4 posterolaterally (extending mediallyanterior to OP) and on Cb5 anterior to OP attach-ment, beginning at distal end and extending medially,joining raphe with PCI.SOD absent.RDs well separated in larger specimen, much lessso in smaller specimen.Additional remarks. SCL free from Bb3. TV4 ven-trally free across Cb5s, but with divided dorsal at-tachment to lateral surface of anterior end of eachCb5. IAC, Pbl, Pb2, Pb4, and UP4 absent. Cartilagetipped uncinate process on Ebl, Eb3, and Eb4 absent.Eb4 levator process absent.
Lepidoblennius marmoratus (Macleay), USNM201625, 82.1 mm; USNM 201626, 90.7 mm.Plate 187
Description.LEI broadly on expanded bony dorsoposterioredge of Eb 1
.
LE2 broadly on expanded bony dorsoposterioredge of Eb2.LE3 anteriorly on dorsoposterior edge of Eb3 allbony uncinate process.LE4 massive, on entire dorsolateral surface of Eb4beginning medially at posterolateral edge of OD3-4.LP massive, fused with LE4 posteroventrally, ex-tending posteroventrally ventral to Eb4 and joiningraphe with dorsal end of Ad5; tendon from rapheextends to Cb5; muscle fuses with OP medially;some posteromedial fibers continuous ventrally withOP.LI1 on dorsoanterolateralmost edge of Pb3 (Pb2absent).LI2 on Pb3 dorsoposterolaterally, paralleling me-dial ends of Eb3 and Eb4.TD comprises TEb2 and TPb3. TEb2 broadly in-terrupted medially, becoming thin, fascia-like and at-taching to broad, flat Pb3 dorsal surfaces; fascia alsocontinuous with TPb3 anteriorly; attaching laterallyon Eb2 dorsally well medial to LE2, continuous pos-teriorly by diagonal muscle strand with TPb3. TPb3on Pb3 dorsolaterally at medial edge of LI2 insertion,ending posteriorly medial to medial end of Eb4.OD3?4 origin on Pb3 dorsally ventral to TEb2,insertion on Eb3 dorsoanterior surface medial to LE3insertion (joining ventromedial edge of LE3) and onmid-dorsal surface of Eb4.OP dorsally, broadly on Eb4 coincident with ven-troposterior edge of LP insertion; ventrally broadlyon Cb5 joining raphe ventrolaterally with PCI attach-ment to Cb5, medially incompletely separated fromSO.
220 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Adl on Ebl bony surface anterodistally and Cblbony surface medial to distal end.Ad2 broadly on most of Eb2 bony surface dor-soanteriorly lateral to TEb2, narrowly on dorsoanter-iormost bony surface of Cb2.Ad3 broadly on most of Eb3 bony surface dor-soanteriorly, narrowly on Cb3 bony anterodistalmostsurface.Ad4 broadly on Eb4 dorsolaterally anterior to OP,very broadly on Cb4 medial to Eb4-Cb4 joint. (Notvisible in illustration.)Ad5 on Cb4 posterolaterally (extending mediallyanterior to OP), on Cb5 anterolateral^ anterior to OP,some fibers continuous with OP.SOD absent, but on one side of only one specimen,a slender filament of SO muscle extends dorsome-dially, passes dorsal to the RD on that side and re-turns to SO between the two RDs.RDs separated by space less than half diameter ofone RD.Additional remarks. SCL present. TV4 dorsal fi-bers attach to lateral surface of anterior end of Cb5on each side, ventral fibers continuous across Cb5s.IAC, Pbl, Pb2, Pb4, and UP4 absent. Cartilagetipped uncinate process on Ebl, Eb3, and Eb4 absent.Eb4 levator process absent.BLENNIIDAEParablennius gattorugine (Linnaeus),276284, 94.4 mm. Plate 188 USNM
Additional material. ? Parablennius tasmanianus(Richardson), USNM 276284, 81.3 mm; ? Scar-tella cristata (Linnaeus), USNM 208445, 105 mm;? Scartichthys gigas (Steindachner), USNM227556, 77.8 mm; ? Istiblennius edentulus(Schneider and Forster), USNM 334144, 108 mm.
Description.Remarks. Although the disposition of the musclesin all the taxa is generally similar, information on ?
? ? ? should be considered limited to the specificfeatures described.LEI on tip of all bony Ebl uncinate process andtendon attaching posterior end of insertion to Eb2. ?Like P. gattorugine.
? ? ? Restricted to bony pro-cess and surface of Eb 1
.
LE2 enveloping pointed, vertical bony mid-dorsalEb2 process and on tendon attaching posterior end ofinsertion to Eb3. ? Like P. gattorugine. ? ? ? Re-stricted to bony process and surface of Eb2.LE3 on raised edge of bony Eb3 uncinate processand on CT tendon attaching Eb3 to Eb4. ? Like P.gattorugine.
? ? ? Restricted to bony process andsurface of Eb3.LE4 on Eb4 dorsally near lateral end of bony sur-
face, joining raphe ventroanteriorly with OD4. ?Does not join raphe.LP joining raphe with lateral edge of LE4 insertionand extending laterally to end of bony portion of Eb4.LI1 on lateral surface of dorsal Pb3 articulatingprocess near joint with Eb2. ? ? ? ? Like P. gat-torugine.LI2 on Pb3 dorsolaterally medial to medial end ofEb3. ? ? ? ? Like P. gattorugine.TD comprises TEb2 and TPb3. TEb2 musclechanges to CT medially as muscle passes dorsal toessentially naked Pb3 dorsal facets, attaches on Eb2anterior surface ventral to pointed mid-dorsal bonyprocess, there joining raphe with dorsal edge of Ad2;continuous by fine strand of muscle posteriorly withTPb3. TPb3 on Pb3 dorsally medial to LI2 insertionand medial end of Eb4, abutting but not continuousposteriorly with SOD. ? Comprises TEb2 and TPb3;anomalous strap of TEb2 attaches to Eb4 dorsome-dially on one side. ? Comprises TEb2 and TPb3-Eb4; TEb2 uninterrupted; TPb3-Eb4 attaches to Pb3dorsolaterally ventral to attachment on Eb4 medially.? Comprises TEb2 and TPb3-Eb4; TEb2 broadlycontinuous across Pb3s, notched anteriorly, with me-dian longitudinal raphe; TPb3-Eb4 attached to Pb3anterior to medial end of Eb4 and to Eb4 dorsome-dially. ? TD like P. gattorugine, but TEb2 is contin-uous posteriorly across Pb3s and attaches on dorso-posterior surface of Eb2.OD3?4 origin on lateral edge of Pb3 articulatingfacet, insertion very finely on dorsal edge of bonyEb3 uncinate process, joining raphe there with LE3insertion, and massively on Eb4 dorsally joining ra-phe there with ventromedial edge of LE4 insertion.
? ?
? Like P. gattorugine. ? Insertion only on Eb4(OD4).Remarks. When present in any of the taxa, the in-sertion on Eb3 consists at most of a fine musclestrand. We think it probable that the presence or ab-sence of the attachment may vary within a taxon.OP dorsally broadly on Eb4 posteriorly, overlap-ping Ad4 posteromedially; ventrally on Cb5 poster-odistally, joining raphe ventrolaterally with PCI. ? ?? ? Like P. gattorugine.M. Eb4-F, dorsally thin, sheet-like, on anterodistalbony edge of Eb4 ventrally anterior to Ad4, extend-ing ventromedially, becoming more string-like andmeshing into SO. Also present, at least, in ? ? ?.Adl dorsally on anterolateral half of bony surfaceof Ebl, ventrally on lateral third of bony surface ofCb 1 . ? ? ? ? Present in all taxa.Ad2 dorsally on anterolateral bony surface of Eb2joining raphe with distal end of TEb2 ventrally, ven-trally on anterodistal bony surface of Cb2. ? ? ?Present in all taxa. ? Extending from dorsomedial-most bony surface of Eb2 to anterodistal bony sur-face of Cb2, joining raphe posteriorly with TEb2.
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1
221
Ad3 dorsally on anterolateral half of Eb3 bony sur-face, ventrally on dorsodistal bony surface of Cb3.
?
? ?
? Present in all taxa.Ad4 dorsally on ventral surface of lateral half ofEb4 anterior to OD and posterior to M. Eb4-F, ven-trally on Cb4 dorsally medial to Eb4-Cb4 joint andanterior to Ad5. ? ? ? ? Present in all taxa.Ad5 dorsally on Cb4 posterolaterally, ventrally ondorsolateral half of Cb5. ? ? ? ? Present in all taxa.SOD present. ? ? ? Present; questionably absentin ? (needs verification in another specimen).RDs adjacent. ? ? ? ? Like P. gattorugine.Additional remarks. SCL present. TV4 in two sec-tions: broad, uninterrupted ventroanterior section andnarrow, interrupted slightly dorsoposterior section at-taching to anterolateral surface of Cb5. ? ? ? ?Like P. gattorugine. IAC, Pbl, Pb2, Pb4, and UP4absent in all taxa. Ebl, Eb3, and Eb4 cartilage-tippeduncinate processes absent.DACTYLOSCOPIDAE
Dactylagnus mundus Gill, USNM 205741, 109 mm;USNM 205742, 2 specimens, 92.2-92.7 mm.Plate 189
Description.LEI broadly on Ebl mid-dorsally.LE2 broad based, beginning on dorsoposterome-dial edge of Eb2 and continuing laterally onto CTjoining Eb2 and Eb3, about half insertion on Eb2,half on CT.LE3 absent.LE4 broadly on most of bony anterior edge of Eb4lateral to OD3-4.LP on Eb4 at and paralleling entire posterior edgeof LE4 insertion.LI1 broadly on ventral surface of anterior arm ofPb3 (Pb2 absent).LI2 on Pb3 dorsolaterally medial to medial end ofEb3.TD comprises TEb2 and TPb3-Eb4. TEb2 broadwith mid-longitudinal raphe attaching mid-ventrallyto CT of pharyngeal roof and giving rise to filmy CTsheets attaching to skull; muscle not overlying dorsalarticulating surfaces of anterior Pb3 arms, extendingdorsolaterally to point on Eb2 ranging from oppositemedial edge of LE2 insertion to mid-point of inser-tion (less extensive than any other blennioids excepttripterygiids); posteriorly dorsal to, and not continu-ous with, TPb3-Eb4. TPb3-Eb4 narrowly on Pb3 dor-soposterolaterally, continuing posteriorly and attach-ing to much of posterior edge of Eb4, posteriorlydorsal to, and not continuous with, SOD.OD3-4 origin on Pb3 dorsolaterally ventral toTEb2, insertion on Eb3 dorsally in area near and dor-
sal to all bony uncinate process and on most of Eb4
medial and dorsal to all bony uncinate process, par-alleling TPb3-Eb4 attachment to Eb4.OP dorsally on Eb4 beginning a lttle medial todistal end and extending medially aboult half lengthof bone, medially indistinguishable from SO, ven-trally broadly on Cb5 beginning near distal end andextending medially joining raphe ventrally with at-tachment of PCI on Cb5 distally, laterally overlap-ping much of slender Ad5.Ad 1-3 absent.Ad4 hidden in posterior view, distinguishable ven-trally by broad, separate insertion on Cb4 anterior toattachment of Ad5, dorsally on Eb4 ventrally, appar-ently fusing with OP posteriorly.Ad5 slender, on dorsoposterior surfaces of distalends of Cb4 and Cb5, medial edge posteriorly over-lapped by OP laterally.SOD present.RDs separated by space less than diameter of oneRD.Additional remarks. SCL free from Bb3. TV4 in-terrupted, attaching to lateral surface of each Cb5,few, if any, muscle strands continuous across Cb5s
ventrally. IAC, Pbl, Pb2, Pb4, and UP4 absent. Eb4levator process absent.CLINIDAE (MYXODINAE)Gibbonsia evides (Jordan and Gilbert), USNM152005, 2 specimens, 116-126 mm.Plate 190
Additional material. ? = Heterostichus rostratus,USNM 132367, 208 mm.Remarks. Gibbonsia and Heterostichus are mem-bers of the oviparous, putatively least specializedclinid tribe Myxodini.
Description.LEI broadly on dorsoposterior bony expansion ofEbl.LE2 broadly, anteriorly on dorsoposterior bony ex-pansion of Eb2.LE3 on Eb3 dorsoanteriorly lateral to tip of allbony uncinate process.LE4 broadly on dorsodistal bony surface of Eb4.
?
Joins raphe ventrally with OP lateral section dor-
sally.LP at and posterior to LE4 insertion.LI1 on lateral edge and much of ventral surface ofPb3 anterior process.LI2 on Pb3 dorsoposterolaterally at medial end ofEb3.TD comprises TEb2 and TPb3-Eb4. TEb2 sinu-soidal, with mid-longitudinal raphe anteriorly, whichgives rise to filmy CT covering muscles dorsally; at-taching anteriorly by tough fascia to Pb3s and pha-ryngeal roof (most of dorsal surface of Pb3 anterior
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
processes not covered by muscle); attaching overmost of Eb2 dorsal surface, meeting anterior edge ofLE2 insertion; overlapping, but not continuous with.TPb3-Eb4 mid-anteriorly. TPb3-Eb4 broadly dorsallybeginning anterior to LI2 insertion, continuing me-dial to insertion and onto dorsomedial Eb4 surface,not continuous with SOD.OD3-4 origin broadly on lateral edge of Pb3 pos-terolaterally ventral to TEb2, insertion on medialedges of all bony Eb3 and Eb4 uncinate processes.
? Insertion continues dorsoposteriorly across Eb4and joins raphe with OP medial section dorsally.OP dorsally beginning on Eb4 uncinate processposterior surface, there meeting OD3?4, and extend-ing laterally about half distance to distal end of Eb4,there meeting Ad5 dorsomedially; ventrally broadlyon Cb5 posterolaterally; medially inseparable fromSO. ? OP more or less divisible into lateral and me-dial sections.Ad 1-3 absent, moderately developed GFMs 1-3present.
?
GFMs even more weakly developed thanin Gibbonsia.Ad4 dorsally narrowly on Eb4 ventrolaterally, con-tinuing on Cb4 dorsolaterally medial to Eb4-Cb4joint, fused broadly posteriorly with Ad5. (Not visi-ble in illustrations.) ? Not fused with Ad5.Ad5 dorsally on Eb4 posterolaterally, just attach-ing to posterodistalmost end of Cb4, ventrally on Cb5dorsoposterodistally, there meeting PCI; anterior sur-face fuses with Ad4. ? Not fused with Ad4.SOD broad.RDs separated by distance less than one-fourth di-ameter one RD.Additional remarks. SCL free from Bb3 (? at-tached mid-dorsally to posteroventral cartilaginoustip of Bb3). TV4 in two sections, dorsal portion di-vided and attached broadly to anterolateral surface ofCb5 on each side; ventral portion continuous acrossanterior ends of Cb5s. Pbl cartilaginous; IAC, Pb2,Pb4 and UP4 absent. Ebl, Eb3, and Eb4 cartilage-tipped uncinate processes absent. Eb4 levator processabsent. CLINIDAE (CLININAE)Heteroclinus perspicillatus (Valenciennes), USNM201518, 110 mm SL.Plate 191A, BAdditional material.
?
= Clinus acuminates (Blochand Schneider), USNM 199579, 74.3 mm SL(Plate 19 1C); ? = Ophiclinus gracilis (Waite),USNM 218794, 99.4 mm SL; ? = Springeratusxanthosoma (Bleeker), USNM 204628, 68.9 mmSL.Description.Remarks. In general, only minor differences areexhibited by the four taxa.
LEI on expanded bony dorsoposterior edge of Ebl(no uncinate process).LE2 very broad, on expanded bony dorsoposterioredge of Eb2.LE3 very broad, on and lateral to all bony Eb3uncinate process.LE4 very broad but with much narrower insertionon Eb4 bony surface lateral to bony uncinate process,joining short raphe ventromedially with dorsomedialend of lateral section of OP.
?
? Does not join OD3-4. ? Just barely joins raphe with OP at Eb4 (see LP).
? LE4 joins OD3-4 laterally, raphe with OP is justventral to Eb4. ? Raphe with OP just ventral to Eb4,involves most of posterior edge of insertion.LP on Eb4 at and slightly posterior to LE4 inser-tion, ventroposterolaterally joining raphe with dor-solateral end of OP lateral section. ? Insertion an-teriorly joins almost entire ventral edge of LE4 in-sertion, such that only medial edge of LE4 insertionjoins OP, whereas entire posterior LP insertion joinsraphe with OP.LI 1 on lateral edge and much of ventral surface ofPb3 anterior process (Pb2 absent).LI2 on Pb3 dorsoposterolaterally medial to medialend of Eb3.TD comprises TEb2 and TPb3-Eb4. TEb2 withbroad anteromedial CT portion overlying surfaces ofPb3s anteriorly; muscular portion with mid-longitu-dinal raphe, which gives rise to CT sheets dorsally;muscular portion connected mid-ventrally betweenPb3s to CT of pharyngeal roof, attaches laterally todorsal surface of Eb2 at point opposite lateral end ofLE2 insertion, joining fine raphe with dorsomedialend of GFM2, not connected posteriorly with TPb3-Eb4. TPb3-Eb4 on Pb3 dorsoposteriorly medial toLI2 insertion and on Eb4 medial end dorsoposterior-ly, joined by diagonal muscle strand with SOD. ?TEb2 relatively broad, covers all but anteriormostPb3 surfaces.OD3?4 origin on Pb3 dorsoposterolaterally ventralto TEb2, insertion on dorsoposterior surfaces of allbony Eb3 and Eb4 uncinate processes, joining rapheposteromedially with dorsomedial end of medial sec-tion of OP.OP with posteriorly distinct lateral and medial sec-tions; lateral section joining raphe dorsally with LE4and LP insertions on Eb4, ventrally joining commonraphe with ventroposterior end of Ad5, ventroposter-olateral end of medial OP section, and dorsoanteriorend of PCI; medial section dorsally on bony Eb4 un-cinate process posteriorly, joining raphe mediallywith OD3-4, ventrally broadly on posterior surfaceof Cb5 impinging broadly on PCI attachment andventrolaterally joining raphe with PCI, medially con-tinuous with SO (continuation obscured in Plate191 A). ? ? Two sections appear to be fused.Adl-3 absent (GFM1-3 weakly developed).
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Ad4 (not visible in illustrations) dorsally on Eb4anterolateral^ ventral to LE4 insertion, ventrallycontinuing broadly onto distal one-third of Cb4 me-dial to Eb4-Cb4 joint.Ad5 dorsally broadly on posterolateral edge ofCb4, passing mostly anterior to OP lateral portion,ventrally broadly on Cb5 dorsolaterally, joining com-mon raphe posteroventrally with PCI dorsoanteriorlyand OP lateral section ventrolaterally. ? Appears tobe fused medially with OP, but should be verified inanother specimen.SOD present. ? ? Absent. ? Present.RDs separated by narrow space. ? Separated byspace greater than diameter of one RD.Additional remarks. SCL unusually thick, appar-ently continuous mid-dorsally with thick walls ofaorta, which appears to be attached to ventroposteriorsurface of Bb3 (? ? ? SCL present). TV4 ventrallycontinuous between Cb4s, dorsally interrupted andattached to anterolateral surface of each Cb5. Pblcartilaginous; IAC, Pb2. Pb4 and UP4 absent. Car-tilage tipped Ebl. Eb3. and Eb4 uncinate processesabsent. Eb4 levator process absent.LABRISOMIDAE
Calliclinus geniguttatus (Valenciennes), USNM269371, 91.3 mm. Plate 192
Additional material. ? = Labrisomus philippi (Stein-dachner), USNM 128206, 103 mm.
Description.LEI on expanded bony dorsoposterior margin ofEbl.LE2 broadly on bony dorsoposterior edge of Eb2.ventroanterolateral edge joining raphe with TEb2posterodistally.LE3 on expanded bony dorsal edge of Eb3 dor-soanterodistally.LE4 on most of bony dorsal surface of lateral halfof Eb4, medial fibers joining raphe ventrally withpresumed OP portion of fused Ad5 + OP + SO,ventrolaterally meets LP ventromedially. ? Joins ra-phe with medial section of OP dorsally.LP on Eb4 meeting LE4 ventrolaterally; ventro-medially joining CT with presumed Ad5 portion ofAd5 + OP + SO. ? Joins LE4 insertion posteriorly,not continuous with any other muscle.LI1 on ventroanterolateralmost surface of Pb3.LI2 on Pb3 dorsoposterolaterally just medial tomedial end of Eb3.TD comprises TEb2, TPb3-Eb4, and, vestigialTPb2. TPb2 (see discussion under Additional re-marks) a fine, flat, semicircular ribbon of muscle oneach side dorsal to TEb2; muscle originates antero-laterally on dorsoposterolateral edge of thick CT area
forming anteromedial area of TD and fuses withTEb2 anteromedially near posteromedial edge of CT.TEb2 anteriorly originates on posterior margin of CTarea and is divided by mid-longitudinal raphe (CTarea continuous anteriorly and ventrally with CT ofpharyngeal roof, attaches also to medial end of Eb 1
,
does not overlie Pb3, and gives rise dorsally to CTsheets covering TD); laterally, muscle attaches onEb2 dorsally. reaching slightly anterolateral to LE2insertion, almost to distal end of bony surface; mus-cle not continuous posteriorly with TPb3-Eb4. TPb3-Eb4 attaches mid-ventrally to CT of pharyngeal roof,laterally, broadly on Pb3 dorsally beginning anteriorto LI2 insertion and continuing posteriorly, formingraphe with median edge of LI2 insertion, and con-tinuing onto dorsodistal and posterodistal surfaces ofEb4. ? Lacks TPb2.OD3-4 origin on Pb3 posterolaterally ventral toTEb2. insertion on dorsal edge of Eb3 that joins Eb4and massively on Eb4 dorsally, with narrow sectionof fibers continuous posteroventrally with presumedOP portion of Ad5 + OP+ SO.OP fused indistinguishably with Ad5 medially andSO medially on right side, left-side Ad5 separate;ventrally, muscle complex joins raphe with PCI onCb5. ? OP in two sections: medial section dorsallyjoining fibers of OD3-4 on Eb4 uncinate process;lateral section dorsolaterally on Eb4 ventral to LPinsertion, dorsomedially continuous with LE4.Ad 1-3 absent (GFMs moderately developed).Ad4 dorsally on Eb4 ventrolaterally. ventrally ondorsal surface of lateral quarter of Cb4, medial toEb4-Cb4 joint, not visible in posterior view.Ad5 fused indistinguishably with OP on one side,distinct on other: on Cb4 posterolaterally and Cb5dorsodistally, partly anterior to OR ? Distinct.SOD absent.RDs separated by space about one-half RD diam-
eter.Additional remarks. SCL attached mid-dorsopos-teriorly by CT to ventroposterior end of Bb3 (carti-lage tip not elongated). TV4 attached dorsally to an-terolateral surfaces of Cb5s, continuous ventrallyacross Cb5s. Pbl cartilaginous; IAC, Pb2, Pb4 andUP4 absent. Cartilage-tipped uncinate processes onEbl, Eb3, and Eb4 absent. Eb4 levator process ab-sent.TPb2 is usually defined by its attachment to Pb2,which is absent in all blennioids. The muscle in Cal-liclinus is similar in shape and position to TPb2found in some other acanthomorphs (e.g., Dactylop-tena, Dactylopteridae; Callionymus, Callionymidae)and is undoubtedly vestigial, similar to its occurrencein Rachycentron (Plate 149) which has TPb2 weaklyrepresented, present only unilaterally, and possiblyanomalously.Additional specimens of Calliclinus should be ex-
224 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
amined to verify presence of a vestigial TPb2. Pres-ence of the muscle in our specimen may be anoma-lous. A specimen of the labrisomid genus Auchen-ionchus, which VGS considers a possibly plesiomor-phic labrisomid, was not available for dissection andshould also be examined. The presence of TPb2 inCalliclinus, probably has little bearing on the inter-relationships of the blennioid families, although itspresence in another labrisomid genus might indicateclose relationship with Calliclinus.CHAENOPSIDAE
Neoclinus blanchardi Girard, SIO 85-14, 129 mm.Plate 193
Description.LEI broadly on anterior surface of all bony unci-nate process of Ebl.LE2 broadly on dorsoanterior surface of expandedbony flange of Eb2.LE3 broadly dorsolaterally on Eb3 bony surface,joining Eb3 portion of OD3?4.LE4 broadly on dorsodistal bony surface of Eb4,just encroaching on cartilaginous distal end.LP on Eb4 posterior to LE4, fusing anteroventrallywith LE4 insertion.LI1 on lateral edge of Pb3 anterior process (Pb2absent).LI2 on Pb3 dorsoposterolaterally, just anterior tomedial end of Eb3.TD comprises TEb2 and TPb3-Eb4. TEb2 broadmid-dorsally, attenuated laterally, notched mid-ante -riorly and mid-posteriorly, with mid-longitudinal ra-phe, which gives rise to CT sheets that attach to skull;broad mid-dorsal portion attached mid-anteroventral-ly to CT of pharyngeal roof and ventrolaterally toPb3 dorsally; attenuated portion on Eb2 dorsallyreaching point anteroventral to LE2 insertion: muscleoverlapping anterior end of, and unattached to, TPb3-Eb4. TPb3-Eb4 broadly on Pb3 dorsal surface ante-rior to LI2 insertion, continuing posteriorly to pointmedial to insertion and onto Eb4 dorsomedially, con-tinuous posteriorly with SOD.OD3-4 origin broadly on Pb3 dorsally ventral toTEb2, insertion on Eb3 dorsomedially and dorso-medial edge of bony Eb4 uncinate process.OP broadly dorsally on posterior surface of Eb4extending from bony uncinate process laterally topoint below LP insertion, ventrally on posterolateralbony surface of Cb5 dorsally, almost or partly meet-ing PCI, which attaches along much of posterolateralsurface of Cb5 ventral to OP attachment and com-pletely envelops distal end of Cb5.Ad 1-3 absent.Ad4 beginning on ventral surface of Eb4 anteriorto OP dorsally and extending to Eb4-Cb4 joint, and
attaching to dorsal surface of distal third of Cb4 me-dial to joint.Ad5 on dorsal edge of Cb5 distally and much ofposterolateral surface of Cb5; attachment includesposterodistalmost surfaces of Eb4 and Cb4.SOD slender.RDs unequal, well separated.Additional remarks. SCL present, free from Bb3cartilaginous posteroventral end (which has small,separate cartilage attached to tip). TV4 in two sec-tions, dorsal portion divided and attached to antero-lateral surface of anterior end of Cb5 on each side;ventral portion continuous across anterior ends ofCb5s. Pbl cartilaginous; IAC, Pb2, Pb4 and UP4 ab-sent. Cartilage-tipped uncinate processes absent onEbl, Eb3, and Eb4. Eb4 levator process absent.
Gobioidei
Remarks. Miller (1973), corroborated by Springer(1983), hypothesized the Rhyacichthyidae as the sis-ter group of all other gobioids. Hoese and Gill(1993b) hypothesized Odontobutidae as the next go-bioid clade, hence sister group to all remaining go-bioids, but lacked a synapomorphy for the group. Inadditional remarks following the description ofOdontobutis, we hypothesize a synapomorphy for theOdontobutidae. Recognition of family level taxaamong the remaining gobioids is in a state of flux;our usage follows some generally recognized groups.To the gobioid synapomorphies listed by Winter-bottom (1993b) and Shibukawa et al. (2001), we addthe absence of an Eb4 uncinate process, or, if a bonyprocess is present, it lacks a cartilage tip. This char-
acter, however, occurs variously among other acan-thomorphs. We have summarized a few other gill-arch character states for gobioids in Table 1 1
.
Wang et al. (2001) based a molecular phylogenyof the gobioids on 32 genera belonging to six sub-families in four different families. Our and their ge-neric taxonomic coverages overlap very little, butone of their findings is, perhaps, supported by one ofours. Their strict consensus tree (their fig. 3) indicatesa monophyletic grouping of their Sicydiinae (Stipho-don, Sicyopterus), Oxudercinae (Periophthalmus, Bo-leophthalmus), and some of their Gobionellinae gen-era (Rhinogobius, Oligolepis, Stenogobius).Among the genera we examined, Pseudapocryptes(Oxudercinae) and Sicydium (Sicydiinae) are the onlygobioids having the muscle, M. Pb3-Eb3. However,a similar muscle, M. Pb3-Eb3-Eb4, also occurs inGnatholepis (the only member of the Gobionellinaewe examined). These three genera also have a thickpad covering Pb3 dorsally. The muscle and pad per-haps offer support for Wang et al.'s (2001) findingsand indicate a potentially new monophyletic groupwithin the gobioids.
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Table 1 1 .?Distribution of certain characters in selected families and genera of gobioid fishes. P = present; absent.
CharactersTaxa LI1 on LI1
split LI2on Pb4 UP4 TEb3 TEb4 TPb3 TPb3-Eb4 TPb3-Pb4 TPb3-UP4 TUP4RhyacichfhyidaeRhyacichthys Pb2 & Pb3 no Pb3 & Pb4 P P - - - P - - -OdontobutidaeOdontobutis Pb2 & Pb3 no Pb3 P P P P P - - - -Micropercops Pb2 & Pb3 no Pb3 (& Pb4 ?) P P ? - P - - - -Percottus Pb2 & Pb3 no Pb3 P P P - P - - - -EleotridaeEleotris Pb2 & Pb3 yes Pb3 & Pb4 P P - - - - P - -Ophiocara Pb2 & Pb3 yes Pb3 & Pb4 P P - - - - P - -MicrodesmidaeMicrodesmus Pb2 no Pb3 - - - - - - - - -Ptereleotris Pb2 & Pb3 no Pb3 & UP4 - P - - P - - - PNemateteotris Pb2 & Pb3 yes? UP4 P P - - - - P - -XenisthmidaeXenisthmus Pb2 no Pb3 - - P - - - - - -GobiidaeGlossogobius Pb2 & Pb3 yes Pb3 & UP4 - - - - P - - - -Bollmannia Pb2 & Pb3 yes Pb3 & UP4 - P - - - P - - -Oxuderces * Pb2 & Pb3 yes Pb3 - P** ? ? ? ? ? ? ?Pseudapocryptes Pb2 & Pb3 yes Pb3 - P - - P - - - PGnatholepis Pb2 & Pb3 yes Pb3 - P - - P - - - PPadogobius Pb2 & Pb3 no Pb3 - P - - - - - P -Sicydium * Pb2 & Pb3 no Pb3 - P - P - - - - -Trypauchen Pb2 & Pb3 no Pb3 - P - P - - - P -
*Genus treated only incidentally in descriptive accounts. *Apparently fused with UP3.
Another gobioid specialization, also present in awide variety of percomorphs, is the Eb4 flange, abony distal extension of Eb4 that overlaps the carti-laginous distal end of the element. Among the taxawe examined, its presence is variable among generaand even among individuals of the same species. Ina cleared and stained specimen of Rhyacichthys(AMS 48695), the flange is well developed anteriorlyon one side and absent on the other. In the illustratedspecimen, the flange is weakly trilobed (anterior-mid-dle-posterior) on one side and weakly bilobed (an-terior-posterior) on the other, with the posteriormorelobe slightly better developed. In the odontobutids, itis only present anteriorly and varies from reduced tomoderately developed. It is moderately developed an-teriorly in eleotrids, weakly bilobed {Ptereleotris, Ne-mateleotris) or absent {Microdesmus) in microdes-mids, well developed in {Oxuderces) or absent {Pseu-dapocryptes) among oxudercines, and absent in thegenerally reduced skeletons of xenithmids and cer-dalids. Among gobiids, it is well developed in Gnath-olepis and Padagobius, well developed or absent inGlossogobius, and absent in Bollmannia. and Try-pauchen.
RHYACICHTHYIDAE
Rhyacichthys aspro (Valenciennes), USNM 247300,175 mm; QM 1.31044, 165 mm.Plate 194
Description.LEI on dorsoposterior edge of Ebl just lateral tobase of uncinate process.LE2 on dorsoposterolateralmost edge of Eb2, in-sertion impinges on TEb2 attachment.LE3 on anterior surface of Eb3 just lateral to tipof uncinate process, insertion impinging medially onEb3 insertion of OD3-4, and laterally on Ad3 at-tachment.LE4 on dorsolateral end of Eb4, just meeting LPinsertion.LP on posterior edge of lateral end of Eb4.LI1 on Pb2 uncinate process ventromedially andadjacent dorsoanterior edge of Pb3; posteriorly, in-sertion forms raphe ventromedially with anterolateraledge of OD3-4.LI2 on Pb3 posterolaterally (at attachment ofTPb3-Pb4 on Pb3) and on Pb4 dorsally.Remarks. The insertion of LI2 on Pb4 occurs only
226 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
in gobioids among acanthomorphs (not all gobioidshave Pb4).TD comprises TEb2 and TPb3-Pb4. Anteriorly,TEb2 attaches broadly to CT of pharyngeal roof;muscle with mid-longitudinal raphe; anterolateralhalf of muscle crosses posterior half laterally andjoins it, and together they attach broadly on Eb2 dor-
sally, almost reaching its lateralmost end. TPb3-Pb4attaches broadly dorsolaterally on Pb3 and Pb4, in-cluding along entire medial line of LI2 insertion.OD3?4 origin broadly on dorsoanterior bony sur-face of Pb3, insertion on joined uncinate processesof Eb3 and Eb4 (Eb4 process is all bony).OP dorsally on posterior surface of Eb4, ventrallyon posterior surface of Cb5.Adl absent; GFM1 exceptionally well developed,on anterodistal surfaces of Ebl and Cbl, distal edgeattaching to gill filaments.Ad2 well developed, comprising two layers, ante-
rior layer (GFM) on distal half of Eb2 and dorsoan-terior surface of Cb2, fibers running diagonally (ven-trodistally), fusing with posterior layer (Ad) on an-terodistal end of Eb2; fibers of posterior layer ori-ented almost vertically; distal edge of anterior layerattaching to gill filaments.Ad3 comprising two layers similar to Ad2, excepton Eb3 and Cb3.Ad4 dorsally on dorsoposterior surface of Eb4. andventrally on Cb4 medial to inner angle formed byEb4-Cb4 joint.Ad5 on posterodistalmost end of Cb4 and distal-most end of Cb5.SOD broad.RD well separated from contralateral element, in-serts on Pb3 and, mostly, UP4.Additional remarks. SCL free from Bb3 (cartilag-inous posterior end not elongate or curved ventrally).TV4 free from Cb5s. Threadlike, cartilaginous Pblpresent as two or three linear segments, imbedded inCT and muscle of pharyngeal roof. Pb2 toothed. IACpresent. ODONTOBUTIDAEOdontobutis obscura (Temminck and Schlegel),USNM 264892, 71.2 mm; USNM 264893, 2 spec-imens, 62.3?71.4 mm.Plate 195
Additional material.
?
= Micropercops swinhonis(Gunther), USNM 336883, 56.6 mm; ? = Percot-tus glenii Dybowsky, USNM 105188, 63.9 mm.See also additional remarks for information onNeodontobutis aurarmus (Vidthayanon).
Description.Remarks. We were unable to observe clearly theattachments of the posterior parts of the transversus
dorsalis and the insertion of LI2 in Micropercops.These attachments are much obscured by overlyingmuscles. In the case of LI2, the insertion joins a ra-phe with TD dorsal to an area where several crowdedskeletal elements occur, all or only some of whichmight participate in the insertion. We indicate the at-tachments that we feel confident of, but further studyis needed.LEI broadly, dorsally at and lateral to Ebl unci-nate process, reaching close to distal end of Ebl.LE2 on Eb2 dorsoposteriorly, bordering TEb2 pos-terodistal margin.LE3 on bony Eb3 uncinate process, borderingOD3 insertion posteriorly.LE4 dorsodistally on bony surface of Eb4, inser-tion joined posteriorly with LP.LP dorsodistally on Eb4, insertion joined anteri-orly with LE4.LI1 dorsally on joined anteriormost ends of Pb2and Pb3, about same size as LI2.LI2 on Pb3 dorsolaterally, passes between OD3and OD3' on one side and OD3 and OD4 on theother on way ventrally from origin. ? Passes betweenOD3 and OD4 (OD3' absent); insertion joins raphewith TD medially and includes Pb3, at least (alsopossibly UP4 and Eb3; see remarks following de-scription). ? Passes through OD4 (OD3' absent) andinserts on Pb3 near joint with Pb4.TD comprises TEb2, TPb3, TEb3, TEb4, withmid-longitudinal raphe coursing along TEb2, TEb3,and dorsoanterior half of TEb4. TEb2 broadly at-tached ventroanteriorly with CT of pharyngeal roof,anterior section twists as it extends laterally and joinsremainder of muscle, which attaches on most of Eb2dorsal surface; fascia attaches anterior surface ofmuscle to anterolateral end of Pb2 and IAC; musclecontinuous posteriorly with TPb3. TPb3 extends an-terolaterally and attaches to Pb3 ventral to TEb2 andOD4 and is continuous posteriorly with TEb3. TEb3attaches to dorsomedial end of Eb3 and is posteriorlycontinuous with TEb4. TEb4 attaches to dorsomedialend of Eb4 and is weakly continuous mid-posteriorlywith SOD. ? TD comprises TEb2 and at least TPb3,and attachments to Eb3 and UP4 may also be present(see remarks under description); TEb4 absent. ?TEb4 absent.OD3, OD3' origin on Pb3 laterally (sandwichedbetween Pb3 and Pb2) and extensively on Eb2 ven-tromedial surface, divides into dorsal (OD3) and ven-tral (OD3') portions distally; OD3 inserts on anteriorsurface of well-developed bony Eb3 uncinate processand OD3' inserts on anterodistal surface of Eb3 atmedialmost point of attachment of Ad3. ? OD3 or-igin mainly on Pb3 dorsoposteriorly, with minor at-tachment to Pb2 dorsoposterolaterally; insertion onEb3 dorsally at anterior edge of LE3 insertion; OD3'absent. ? OD3 origin on Pb3 anterolaterally and Eb3
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posteromedialmost end, insertion on Eb3 uncinateprocess; OD3' absent.OD4 origin on Pb3 dorsoanteriorly ventral toTEb2, and slightly on dorsomedial end of Eb2 where
it meets Pb3; divides into dorsal and ventral halvesposterior to origin, insertion anteromedially on Eb4dorsal surface. ? Insertion on Eb4 dorsoanterome-dially. continuing onto ligament joining Eb3 at me-dial edge of LE3 insertion to Eb4 mid-posteriorly. ?Origin on Pb3 with OD3, divides into dorsal and ven-tral portions by passage of LI2, portions recombineposterior to LI2 and insert on anterior surface of bonyEb4 uncinate process, meeting OD3 on Eb3 uncinateprocess.OP dorsally on Eb4 ventroposterior margin pos-terior to main portion of Ad4, ventrally on poster-odistal surface of Cb5 posterior to Ad5 attachment,medially inseparable from SO. ? ? Distinct from SOmedially.Adl dorsally on anterolateral surface of Ebl, ven-trally on anterolateral end of Cbl. ? Dorsally nearend of anterolateral surface of Ebl. ? On Ebl an-terior to LEI insertion and on dorsoanteriormost sur-face of Cbl.Ad2 short, on anterolateral ends of Eb2 and Cb2.
?
On lateral third of Eb2 and anterodistal end ofCb2. ? Meets ventrolateral edge of TEb2 and anter-odistal end of Cb2.Ad3 short, on anterolateral ends of Eb3 and Cb3.
?
Extremely well developed, extends entire length ofEb3, but only on Cb3 anterolateralmost surface. ?On most of Eb3 dorsal surface and anterolateral sur-face of Cb3.Ad4 dorsally on distal half of Eb4 ventral surface,medially just anterior to dorsal attachment of OPventrally on Cb4 dorsoposterior margin just medialto inner angle of Eb4-Cb4 joint.Ad5 dorsally on ventroposterior surface of Cb4,ventrally on posterodistalmost surface of Cb5 just an-terior to ventral attachment of OP.SOD present, continuous mid-anteriorly withTEb4.RD juxtaposed to contralateral RD, inserts on Pb3posterior margin and UP4 dorsally.Additional remarks. SCL free from Bb3 (cartilag-inous posterior end not elongated or curved ventrally.TV4 free from Cb5s. Pbl present, cartilaginous. Pb4and UP4 present. Pb2 toothed. Medial end of Eb4larger than that of Eb3.Some data were recorded for a paratype of Odon-tobutis aurannus (recently assigned to the genusNeodontobutis, by Chen et al., 2002:233), USNM325486, female, 37. 1 mm, which has ambiguous af-finities (Shibukawa et al., 2001:231): LI1 is on Pb2and Pb3. LI2 divides OD4 into a large posterior sec-tion and a fine anterior section comprising a fewmuscle strands contiguous with OD3. TD comprises
TEb2 and, at least, TPb3 and TEb3 (we were unableto verify whether Pb4 is present, and whether thereis an attachment to UP4). There is no TD attachmentto Eb4. M. Pb2-Eb2, not present in the other gobioidswe examined, is on Pb2, ventrolateral to the dor-soanterior cartilaginous tip, and Eb2 dorsoanterola-terally, there becoming continuous with TEb2 antero-laterally. OD3 originates ventrolateral to OD4 on Pb3laterally and adjacent posteromedial edge of Eb3 andinserts anteriorly on bony support of uncinate pro-cess. OD4 originates on Pb3 dorsomedially and in-serts on Eb4 bony uncinate process medial edge andanterior surface, with a few muscle strands on medialedge of Eb3 uncinate process. Pbl is absent. SCL isfree from Bb3. TV4 is free from Cb5s.Hoese and Gill (1993:434) were unable to hypoth-esize a synapomorphy for the Odontobutidae, al-though they suggested, based on Micropercops, thepossibility that a modification of the procurrent cau-dal-fin cartilages might provide a synapomorphy.Watson (in Berra. 2001:460) indicated that "the mor-phology of procurrent cartilages of Odontobutis andPercottus is unremarkable," thus implying that thispotential specialization is not useful.Akihito et al. (2000) provided results from a mo-lecular based unrooted phylogenetic study of 28 go-bioid taxa representing most of the gobioid familiescurrently recognized based on morphologicalgrounds. They found that the taxa grouped into eightgroups: six clusters comprising two to eight taxa anda separate phyletic line each for Odontobutis andXenisthmus. Micropercops swinhonis, the only otherodontobutid included in their study, fell into a mor-phologically highly diverse group of six genera. Theynoted the morphological differences, shown by Mi-cropercops compared with the other genera in itsgroup, and the similarity of its morphology to that ofOdontobutis and to that of their Rhyacichthys-Pro-togobius cluster. They indicated that more study wasdesirable to reconcile the conflicts, but concluded,nevertheless that their data did not support the com-position (monophyly) of Hoese and Gill's (1993)Odontobutidae.We find that the passage of LI2 through OD isunique to odontobutids, including TV. aurarmus,among gobioids, and appears to satisfy the need fora synapomorphy that defines the Odontobutidae. It ispossible the character is more widespread than re-corded here, and it would be useful to determine ifit is present in some other plesiomorphic gobioidgenera of unresolved relationships, e.g., TerateleotrisShibukawa et al., Protogobius Watson and Polla-bauer.Because of the variation in the position of LI2within OD (whether separating the Eb3 from the Eb4components, or splitting only the Eb4 component), itis possible that the position of LI2 vis-a-vis the OD
228 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
components is the chance result of individual ontog-eny. A similar interposition of LI2 in OD3?4 alsooccurs, homoplastically in various ophidiids, bythi-tids, and champsodontids, at least.
XENISTHMIDAE
Xenisthmus sp., USNM 247389, 2 specimens, 29.2-33.3 mm. Plate 196
Description.Remarks. The gill arches are small and greatly de-pressed (almost fiat) and the muscles are thin, un-stained, and essentially transparent, making them dif-ficult to interpret. The relationship of OP and SO areuncertain. Most of the description is based on thelarger of the specimens.LEI on Ebl at and just lateral to uncinate process.LE2 absent.Remarks. Muscle in position of LE2 is weakly,musculously attached to posterolateral end of Eb2,with musculous end continuing as fine tendon andinserting on distal end of Eb3. During cleaning, mus-cle invariably releases from Eb2, but maintains con-nection to Eb3.LE3 present (see remarks under LE2).LE4 dorsoanteriorly on distal end of Eb4.LP just posterior to LE4 insertion (on Eb4-Cb4joint on one side of larger specimen).LI1 on Pb2 dorsolaterally.LI2 on Pb3 dorsoposterolaterally.TD comprises TEb2 and TEb3. TEb2 on mid-dor-soposterior edge of Eb2, continuous mid-posteriorlyby diagonal muscle strap with TEb3, which is onmid-posterior edge of Eb3. continuous posteriorly bydiagonal muscle strap with SOD.OD3 absent.OD4 origin broadly on Pb2 dorsoposteriorly andPb3 dorsoanteriorly, insertion on Eb4 mid-anteriorlyand on ligament joining Eb3 uncinate process withEb4.OP muscle fibers extending from most of posteriorsurface of Eb4 and attaching to Cb5, difficult to sep-arate medially from SO (see remarks following de-scription, above).Remarks. Long, slender muscle strap (adventi-tious?) originates from SO and inserts on Eb4-Cb4joint posteriorly on one side of larger specimen.Ad 1-2 absent.Ad3 long, attaching along most of anterior surfaceof Eb3, but just reaching and attaching to Cb3 an-terordistalmost surface.Ad4 reduced, dorsally on Eb4 posterolaterally,ventrally on Cb4 dorsally anterior to inner angle ofEb4-Cb4 joint; dorsally just visible posteriorly lateralto OP.
Ad5 dorsally on Cb4 distally and Cb5 posterodis-tally.SOD present, broad.RD widely separated from contralateral RD, insertsby long tendon to posterior end of Pb3.Additional remarks. SCL absent. TV4 free fromCb5s. Pbl vestigial, minute and cartilaginous whenpresent. Pb2 edentate. Pb4 absent; UP4 present, butreduced and entirely ventral to Eb4. IAC present.Springer ( 1983:18-21; fig. 11) described and illus-trated the osteology of Xenisthmus clarus (Jordan andSeale). The skeletal structure of the dorsal gill archesof the specimens in the present study is similar tothat which Springer illustrated, except that the con-stricted medial end of the interarcual cartilage of X.clarus is present as a tiny autogenous cartilage. Seealso miscellaneous remarks under Microdesmus.ELEOTRIDAE
Eleotris melanosoma Bleeker, USNM 321251, 3specimens, 59.4-86.7 mm.Plate 197
Additional material. ? = Ophiocara porocephala(Valenciennes), USNM 342613, 59.8 mm.
Description.Remarks. Only conspicuous differences noted forO. porocephala.LEI broadly on Ebl dorsal surface lateral to un-cinate process.LE2 broadly on Eb2 dorsoposteriorly, anterioredge of insertion along posterior edge of TEb2.LE3 on Eb3, variably just lateral to tip of uncinateprocess or on and lateral to process.LE4 tendinously on distal end of Eb4 dorsoanter-iorly, joining LP insertion.LP on dorsodistalmost surface of Eb4, anterome-dially joining LE4 insertion.LI 1 anteriorly on dorsolateral surface of Pb2 ven-tral to LIT insertion.LI1' laterally on dorsoanterior edge of Pb3 dorsalto LI1 insertion.LI2 on posterolateral dorsal surface of Pb3 andadjacent medialmost tip of Eb3, continuing uninter-rupted onto lateral edge of Pb4.
? On Pb3 dorsolat-erally and Pb4 laterally.TD comprises TEb2 and TPb3-TPb4. TEb2 withmid-longitudinal raphe attaching ventrally to CT ofpharyngeal roof, giving rise dorsally to filmy CTcovering gill arches; flat ribbon of fibers arises frommuscle anterolaterally, twists and extends laterallyand inserts among fibers of remainder of TEb2,which extends onto Eb2 almost to dorsolateralmostend; muscle continuous mid-posteriorly with TPb3-Pb4. TPb3-Pb4 with mid-longitudinal raphe in Pb4portion; muscle on Pb3 dorsoposterolaterally at me-
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dial edge of LI2 insertion and on anterior margin ofPb4; muscle discontinuous (two specimens) or con-tinuous by fine, diagonal muscle strand with SOD.OD3 and OD4 with broad, continuous origin ondorsomedial surface of Pb3; muscle divides longitu-dinally into OD3 and OD4 after passing posteriorlyfrom under TEb2; OD3 insertion on anterior surfaceof Eb3 uncinate process and short ligament connect-ing Eb3 and Eb4 uncinate processes. OD4 insertionon same ligament and Eb4 bony uncinate process. ?OD3-4, indivisible.OP with separate overlapping anterior and poste-
rior layers, which may partially fuse in overlap area;anterior layer angled ventromedially from attachmentto Eb4, attaches to Cb5 posteromedially; posteriorlayer angled ventrolaterally from attachment to Eb4posteromedial surface, attaches ventrally on Cb5 pos-terolateral surface.Adl dorsally on most of anterolateral surface ofEbl, ventrally on Cbl anterodistalmost bony surface.Ad2 dorsally on distal half of ventrolateral surfaceof Eb2; ventrally on anterior surface of distal end ofCb2.Ad3 dorsally on most of dorsoanterior surface ofEb3, ventrally on anterodistalmost end of Cb3.Ad4 dorsally on ventrolateral half of surface ofEb4, medially anterior to OP dorsal attachments; ven-trally on dorsolateral surface of Cb4 anterior to innerangle formed by Eb4-Cb4 joint; separation from in-ner angle varying from little to noticeable.Ad5 dorsally on posterolateralmost end of Cb4,ventrally on dorsolateral end of Cb5, meeting ventralattachment of posterior OP layer.SOD present.RD separated from counterpart by space less thanhalf width of RD, tendinously attached to posteriorend of Pb3.Additional remarks. SCL present. TV4 free fromCb5s. Pbl reduced, cartilaginous. Pb2 toothed. Pb4and UP4 present. IAC present.MICRODESMIDAE
Ptereleotris microlepis (Bleeker), USNM 257075,78.0 mm. Plate 198
Additional material. ? Nemateleotris magnifiedFowler, USNM 214103, 48.4 mm SL.
Description.Remarks. The muscles of the two taxa are verysimilar in general appearance.LEI on Ebl from lateral edge of uncinate processto dorsodistal tip of Ebl. ? On Ebl lateral to unci-nate process.LE2 on Eb2 dorsodistally.
LE3 finely, tendinously on Eb3 lateral to uncinateprocess.LE4 finely, tendinously on dorsodistal end of Eb4.LP at and posterolateral to LE4 insertion.LI1 broadly, dorsoanteriorly on Pb2 and Pb3. ?Same, but easily divisible (artificially?) into two mus-cles, one on each of the skeletal elements.LI2 very broadly on Pb3 posterolaterally and UP4medially (Pb4 absent), easily divisible (artificially?)into two muscles, one on each of the skeletal ele-ments. ? On UP4 dorsally lateral to small Pb4, mus-cle not obviously divisible.Remarks. ? Absence of insertion on Pb3 and Pb4noteworthy.TD comprises TEb2, TPb3, and TUP4. TEb2deeply notched anteriorly, notch dividing broad an-terior portion of muscle into right and left halves,each of which originates on CT pad conforming withdorsomedial surface of Pb3; posterior portion of mus-cle broad, transversely continuous with mid-longitu-dinal raphe continuing posteriorly from anteriornotch; both sections extend laterally, with posteriorsection attaching to posterior edge of Eb2 and dorsalsection twisting and overlapping posterior sectionand attaching to dorsal surface of Eb2; TEb2 endingposteriorly as CT, which is continuous mid-poster-oventrally with mid-longitudinal raphe of TPb3.TPb3 attaches to Pb3 laterally, anteriorly meetingmedial edge of LI2 insertion on Pb3 (unclear if someTPb3 fibers also attach to UP4), mid-posteriorly con-tinuous with raphe of TUP4. TUP4 somewhat asym-metrical, attaching to UP4 posteromedially, continu-ous mid-posteriorly with median raphe of SOD. ?Comprises TEb2 and TPb3-Pb4. TEb2 similar toPtereleotris, but "dorsal" section not clearly over-lapping posterior section (therefore, not dorsal).TPb3-Pb4 dorsolateral^ on Pb3 just medial to car-tilage tip of process articulating with medial end ofEb3 and on medialmost edge of small Pb4.OD3-4 broadly on Pb3 dorsomedially ventral toTEb2, insertion on Eb3 uncinate process and Eb4mid-dorsolaterally (no uncinate process). ? OD ori-gin narrowly anteriorly on Pb2 and broadly posteri-orly on Pb3, divides into OD3 and OD4 shortly afterpassing posteriorly from under TD; OD3 inserts onEb3 uncinate process and OD4 inserts on Eb4 mid-dorsolaterally (no uncinate process).OP on Eb4 dorsoposteriorly; ventrally, narrowlyon Cb5 anterodistalmost surface.Ad 1-3 broadly on anterior surface of respectiveEb, and less extensively on associated Cb; Adl over-lain anteriorly by broad bases of two dorsalmost gillrakers.Ad4 almost completely overlapped posteriorly byOP; dorsally, broadly on Eb4 ventral surface; ven-trally, narrowly on Cb4 anterior to internal angle ofEb4-Cb4 joint.
230 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ad5 dorsally on Cb4 posterodistal end, ventrallyon Cb5 distally.SOD present.RD separated from counterpart by interspace equalto about half width of RD, inserts on Pb3 posteriorly.Additional remarks. SCL present, free from Bb3(cartilaginous posterior elongate, not extended ven-trally). TV4 free from Cb5s. IAC present. Pbl ques-tionably present, but possibly the separate rod-likecartilage at the tip of Eb 1 anterior process representsa segmentation near the base of the cartilaginous endof the anterior process, which is often unusually longin gobies. Pb4 absent, UP4 present. ? Ebl with elon-gate cartilaginous tip, no separate Pbl, Pb4 and UP4present. Pb2 toothed.
Microdesmus longipinnis (Weymouth), USNM199614, 162 mm. Plate 199
Description.LEI on Ebl dorsally lateral to uncinate process.LE2 on Eb2 dorsodistally.LE3 slender, on Eb3 just dorsolateral to bony un-cinate process.LE4 very fine, inserts by short, fine tendon on Eb4dorsodistally at medial edge of LP insertion.LP much broader than LE4, on distalmost end ofEb4.LI1 on dorsomedial surface of Pb2.Remarks. Pb2 posterior end abuts Pb3 anteriorend. Except for xenisthmids (Springer, 1983:fig. 11,Xenisthmus; Gill, 1993:fig. 6, Paraxenisthmus) schin-dleriids (Johnson and Brothers, 1993:fig. 8), and Mi-crodesmus, the anterior portion of the gobioid Pb3lies dorsal to Pb2, and LI1 (or LI1 + LI1') insertson both Pb2 and Pb3. LI1 is also only on Pb2 inXenisthmus, which does not have LI1', which is pos-sibly present in the least specialized xenisthmid, Par-axenisthmus (condition in Schindleria also un-known). Although the alignment of Pb2 and Pb3 inthese families may be attributable to developmentaltruncation of the head (the species are small and/orhave small heads), Pb2 and Pb3 retain their dorsal-ventral relationship in the gobiid genus Eviota (le-vators not studied), one of a few gobioid genera thatinclude some of the smallest fish species.LI2 on Pb3 dorsoposterolaterally.TD comprises only TEb2, which attaches broadlyalong Eb2 posteriorly and is mid-posteriorly contin-uous by a fine muscle strand with SOD.OD3-4 arises broadly along medial edges of Pb2and Pb3 and inserts dorsoanteriorly on bony Eb3 un-cinate process and on Eb4 mid-dorsally, joining ra-phe with OP; posteriorly partly continuous with dor-
sal attachment of OP.
OP dorsally with two separate attachments on Eb4;lateral attachment joining raphe with OD3-4, medialattachment posteromedially on Eb4; ventrally on Cb5dorsodistally at and posterior to ventral attachment ofAd5.Ad 1-3 on respective Eb near medial end, mostlyfree from shaft, and on respective Cb anteriorly neardistal end.Ad4 on Eb4 ventrodistally and Cb4 dorsally me-dial to Eb4-Cb4 joint.Ad5 dorsally on Cb4 dorsodistally, ventrally onCb5 dorsodistally.SOD unusually broad.RD separate slightly from counterpart, inserting onPb3 medially and posteriorly.Remarks. The RDs enter the esophagus well pos-terior to gill arches.Additional remarks. SCL present. TV4 free fromCb5s. Pbl absent Pb2 minutely toothed. Pb4 andUP4 absent. IAC absent.GOBIIDAEPseudapocryptes elongatus (Cuvier), USNM 341281,137 mm. Plate 200
Description.LEI on Ebl dorsolateral surface beginning at baseof uncinate process and extending to end of bone.LE2 on dorsolateral one-third of surface of Eb2.LE3 in two sections dorsal to insertion; inserts bytendon bridging dorsomedial edge of uncinate pro-cess and dorsolateral surface of Eb3.LE4 slender, tendinously attached to Eb4 dorso-distally just anterior to lateral edge of LP insertion.LP broad based, insertion covering almost all ofbony Eb4 surface distal to uncinate process.LI1 on Pb2 dorsally ventral to dorsomedial carti-lage-tipped process.LI1' completely separate from LI1, on Pb3 dor-sally near anteromedial end.Remarks. LI1' also occurs in Gnatholepis, Boll-mannia, Glossogobius, Eleotris, and Ophiocara.LI2 on Pb3 dorsoposteromedially ventral to OD3?4 origin (Pb4 absent; UP4 present).TD complex, all elements either completely sepa-rated medially or joined by broad CT area, comprisesTEb2a, TEb2p, TPb3, TUP4, and TD median fibersthat overlie the broad CT area. The CT area appearsto be an expansion of the normal median raphe thatoccurs in the TD of other gobies. TD median fibersoriginate as complex web of fine fibers mid-dorsallyon CT area between TEb2p and give rise to a slenderlongitudinal muscle on each side that inserts on theCT pad on which TEb2a originates. TEb2a origindorsally on spongy CT pad dorsal to Pb3 anteriorly,insertion on Eb2 dorsally just proximal to medial
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edge of LE2 insertion, a few fibers attaching to ten-dinous insertion of TEb2p, completely separate fromcontralateral TEb2a. TEb2p origin as multiple slen-der muscle straps on mid-dorsal CT section, insertiontendinous on Eb2 ventromedial to TEb2a insertion.TPb3 origin on mid-dorsal CT area, insertion on Pb3dorsoposterolaterally just medial to medial end ofEb3. TUP4 origin on mid-dorsal CT area, insertionon medial edge of UP4.Remarks. TEb2a & p appear to be a specializedseparation into two parts of the typical gobioid TEb2,which comprises two incompletely separate sectionslaterally.M. Pb3-Eb3 origin on Pb3 dorsoposteriorly, inser-tion on Eb3 dorsoposterodistally.OD3 and OD4 originate broadly from mixed fibersattaching to Pb3 dorsomedial edge ventral to TEb2p.but are distinct muscles for almost all of their lengths.OD3 is divided mid-longitudinally dorsally, but fibersof two sections mesh ventrally, muscle inserts ten-dinously to medial edge of Eb3 uncinate process.OD4 inserts on dorsoanteriormost edge of Eb4 justanterior to bony Eb4 uncinate process.OP dorsally on Eb4 ventrally posterior to lateralportion of Ad4, ventrally, finely, tendinously on edgeof Cb5 at attachment of Ad5.Adl dorsally on almost entire anterior surface ofEbl medial to uncinate process, ventrally on Cblbroadly anteriorly medial to cartilaginous distal end.Ad2 dorsally on bony anterior edge of Eb2 later-
ally, ventrally on anterior surface of distal bony andcartilaginous end of Cb2 medial to Eb2-Cb2 joint.Ad3 dorsally on bony anterior edge of Eb3 anter-oventral to uncinate process, ventrally on anteriorsurface of cartilaginous distal end of Cb3 ventral toEb3-Cb3 joint.Ad4 in two separate sections. Anterior sectionoriginates on Eb4 dorsally anterior to posterior sec-tion and inserts finely on Cb4 dorsal edge just medialto Eb4-Cb4 joint; posterior section originates on Eb4dorsally posterior to origin of anterior section andinserts narrowly on Cb4 near lateral end of insertionof anterior section.Ad5 on cartilaginous Cb4 posterodistal end anddistally on broad, distal cartilaginous edge Cb5.SOD absent.RDs separated by narrow space anterior to origins,each divides anteriorly into lateral section consistingof loose fibers that insert on posterior edge of UP4,and median robust section that inserts along medialedge of Pb3.Additional remarks. SCL free from Bb3 (cartilag-inous posterior end not elongate or extended ven-trally). TV4 in two parts: posterior part, narrow, con-tinuous, and free across ventral surfaces of Cb5s;broad anterior part on each side attaching medianly
to ventral surface of cartilaginous anterior ends ofCb5s. Pbl absent. Pb2 toothed. IAC absent.
Glossogobius aureus Akihito and Meguro, USNM241833, 2 specimens, 81.4-117 mm.Plate 201Additional material,dan, 71.5 mm.
Additional material. G
nestrini), 70.2 mm.
Bollmannia chlamvdes Jor-
Plate 202Padogobius nigricans (Ca-
Description.LEI mid-dorsally on Ebl just lateral to uncinateprocess.
?
On distalmost bony surface of Ebl.LE2 mid-dorsally on Eb2 just lateral to bony pro-cess on posterior edge. ? Broadly on Eb2 beginning
at distalmost end and extending medially.LE3 on Eb3 just lateral to tip of uncinate process.
?
On lateral edge of tip of uncinate process and ex-tending a short distance laterally.LE4 on dorsodistalmost bony end of Eb4.LP at and posterior to insertion of LE4. ? Fusingwith LE4 laterally just dorsal to LE4 insertion.LI1 on Pb2 dorsoanteriorly. ? On Pb2 and Pb3dorsoanteriorly.LI1' on Pb3 dorsoanteriorly just posterior to LI1.
? Absent.LI2 in larger specimen on Pb3 dorsal surface pos-terolaterally and UP4 anterolaterally, in smaller spec-imen also on tiny Pb4 (Pb4 probably anomalous, notpresent in larger specimen or ? and ?). ? On Pb3dorsoposterolaterally with few strands on Eb3 dor-somedialmost surface.TD comprises TEb2 and TPb3. TEb2 broad withmedian longitudinal raphe on posterior half in largerspecimen (complete in smaller specimen; apparentpair of raphes anteriorly in larger specimen are arti-facts), raphe continues posteriorly across TPb3 andSOD; muscle twists and divides laterally with pos-terior portion of muscle passing anteroventral to an-terior portion and attaching to hook-like process (notillustrated) on mid-anterior edge of Eb2; anterior por-tion attaches to Eb2 anterior surface just posterior tohooklike process; posteriorly broadly continuouswith TPb3. TPb3 originates from posterior half ofmedian raphe, passes laterally ventral to OD3 andOD4 and attaches to Pb3 dorsally medial to LI2 in-sertion. TPb3 narrowly continuous mid-posteriorlywith SOD at median raphe.
? TD comprises TEb2 and TPb3-Eb4, mid-longi-tudinal raphe like Glossogobius. TEb2 similar toGlossogobius, but no hooklike process on Eb2.TPb3-Eb4 on Pb3 dorsoposterolaterally immediatelyanterior to medial end of Eb4, continuing onto medialend of Eb4 dorsally.
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<D TD comprises TEb2 and TPb3-UP4; median ra-phe restricted to posterior portion of TEb2, continu-ing across TEb3-UP4, and only anterior half of SOD;TEb2 twist not apparent, extending laterally to, andjoining medial edge of LE2 insertion and joining ra-phe with Ad2 dorsally (no hook-like process on Eb2);continuous broadly posteriorly with TPb3-UP4.TPb3-UP4 on Pb3 dorsoposteriorly medial to medialend of Eb4, continuing onto UP4 dorsally just pos-teroventral to medial end of Eb4, broadly continuousposteriorly with SOD.OD3 and OD4 originate inseparably on dorsome-dial surface of Pb3 ventral to TEb2; muscle dividesinto OD3 and OD4 as it passes out from under TEb2,with OD3 inserting on dorsomedial edge of Eb3 un-cinate process and OD4 inserting on dorsomedialedge of bony Eb4 uncinate process. ? OD3-4 pres-
ent, not separable into two muscles.OD4 (see OD3 above).OP dorsally broadly on Eb4 posterior surface ex-tending medially almost to medial end, ventrally onposterolateral surface of Cb5 posterior to Ad5 attach-ment.Adl dorsally on most of bony length of Ebl ven-trally, ventrally on anterodistalmost surface of Cbl(muscle does not pass anterior to cartilaginous distalend).Ad2 dorsally on ventral bony surface of Eb2 be-ginning at mid-anterior hooklike process (anteroven-tral to LE2 insertion) and extending laterally to endof bone, ventrally on anterodistal surface of Cb2(muscle does not pass anterior to cartilaginous distalend). ? Dorsally begins at mid-length of Eb2 (nohooklike process). ? Dorsally on ventral surface ofEb2 well medial to LE2 insertion, forming raphe dor-sally with anterior edge of Eb2 portion of TEb2.Ad3 dorsally on most of bony surface of Eb3, ven-trally on anterodistal surface of Cb3 (muscle does notpass anterior to cartilaginous end); section of fibersarises dorsoposterolaterally from main muscle massand attaches to anterior surface of uncinate processventral to OD3 (in frontal view, Ad3 appears to betwo muscles, more-or-less horizontal anterior portionoverlapping posterior vertical portion). ? Verticalsection smaller, inconspicuous, inclined dorsomedi-
ally, only on posterolateral surface of uncinate pro-cess.Ad4 dorsally broadly on Eb4 ventrally anterior toOP, ventrally broadly on Cb4 dorsal surface medialto Eb4-Cb4 joint.Ad5 dorsally beginning on posterodistal end ofCb4 and extending medially short distance, ventrallyto anterodistal end of Cb5 and extending short dis-tance medially anterior to OP attachment.RDs slightly separated. ? Adjacent.SOD slender, connected at mid-dorsal raphe withTPb3. ? SOD broad.
Additional remarks. SCL free from Bb3. TV4 freefrom Cb5s. Pbl absent. Pb2 toothed. IAC present.
Trypauchen vagina (Bloch and Schneider), USNM339608, 149 mm. Plate 203
Description.LEI on Ebl dorsolateralmost bony surface.LE2 on Eb2 dorsolateralmost bony surface, meetsdorsal attachment of Ad2 and distal edge of TEb2.LE3 on Eb3 just lateral to tip of uncinate process.LE4 on Eb4 dorsally near distal end, joins LP in-sertion laterally.LP on dorsodistal end of Eb4 at and lateral to LE4insertion.LI1 on Pb2 dorsoanteriorly and adjacent anteriorend of Pb3, not divisible.LI2 inserts by broad tendon posterolaterally onPb3 dorsal surface.TD comprises TEb2, TPb3-UP4, and TEb4. TEb2attached broadly anteroventrally and ventrally frommid-longitudinal raphe to CT of pharyngeal roof (ra-phe continues across TPb3-UP4); on left side only,thin, medially concave slip of muscle arises frommain muscle mass anterolaterally and re-enters pos-terolaterally; thin, flat muscle slips extend postero-laterally and laterally from mid-longitudinal rapheand join as lateral arm of TEb2 attaches to Eb2 dorsalsurface; broadly continuous posteriorly with TPb3-UP4. TPb3-UP4 attaches to Pb3 posteromedially andUP4 dorsally, and is continuous posteriorly withTEb4. TEb4 attaches to Eb4 mid-posteriorly and liesdorsal to SOD.OD3 and OD4 superficially completely fused, butactually almost completely separate except for fineline of fusion ventrally; muscles originate broadlyalong Pb3 dorsal surface; OD3 inserts along dor-soanterior surface of Eb3 uncinate process and an-terior portion of tight ligament connecting Eb3 andbony Eb4 uncinate processes; OD4 inserts on liga-ment and dorsomedial edge of Eb4 uncinate process.OP dorsally on dorsoposterior surface of Eb4, ven-trally on Cb5 posterodistally, meeting Ad5 ventralattachment; posteriorly, broadly overlaps Ad4 dorsalattachment on ventral surface of Eb4.Adl on most of length of anterior edge of Ebl andon anterolateral surface of Cbl.Ad2 broadly on Eb2, forming raphes with ventro-lateral edge of TEb2 and ventromedial edge of LEI,ventrally on anterolateral surface of Cb2.Ad3 dorsally on most of anterolateral edge of Eb3,ventrally on anterolateral surface of Cb3.Ad4 dorsally broadly on Eb4 ventral surface,mostly occluded in posterior view by posteriorly
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overlying OP, ventrally on Cb4 dorsolateral surfacemedial to inner angle formed by Eb4-Cb4 joint.Ad5 dorsally on posterodistal end of Cb4, ventrallyon Cb5. meeting OP ventral attachment.RDs contiguous, inserting on Pb3 and UP4 pos-teriorly: right side RD with anomalous fibers branch-ing off ventral surface posterior to level of SOD andmeshing with SO fibers.SOD present.Additional remarks. SCL present (condition pre-cluded observing if it was free or attached to Bb3.TV4 free from Cb5s. Pbl and Pb4 absent, UP4 pres-
ent. Pb2 toothed. IAC present, reduced.
Gnatholepis cauerensis (Bleeker), USNM 327750, 2specimens, 42.4-45.6 mm.Plate 204
Description.LEI dorsolaterally on Ebl beginning at base ofuncinate process.LE2 on Eb2 dorsally near distal end.LE3 on dorsal edge of Eb3 between tip of uncinateprocess and distal end of Eb3.LE4 on Eb4 dorsally near distal end.LP massive, broadly on Eb4 dorsally, lateralmostend at and posterior to LE4 insertion, inserted pos-terolaterally on dorsal end of ligament joining Eb4and Cb5.LI1 on Pb2 anterolaterally ventral to TEb2 anter-iormost section.LI1' on Pb3 anterolateralmost end ventral to TEb2anteriormost section; insertion separated by spacefrom LI1 insertion; LI1 and LIT appearing as onemuscle after extending out from under TEb2.LI2 on Pb3 dorsomedial surface near cartilaginousprocess that articulates with medial cartilage tip ofEb4.TD comprises TEb2 and TPb3. TEb2 comprisesthree, almost completely separated sections; anteriortwo sections originate on and are broadly interruptedmedially by CT pad over dorsal surface of Pb2 andanterior surface of Pb3; interrupted sections divide asthey extend laterally, posterior section twisting andbecoming ventral to anterior section before attachingon posteroventral edge of Eb2 well medial to distalend; anterior section attaches on dorsal surface ofEb2 just medial to LE2 insertion; uninterrupted pos-teriormost section of TEb2 with median longitudinalraphe, muscle continuous mid-posteriorly with CTsheet covering dorsoposteriormost surfaces of Pb3s;as posteriormost section of TEb2 extends laterally, itjoins posterior interrupted TEb2 section. TPb3 orig-inates from all margins except anteriormost of CTsheet covering Pb3 dorsoposterior surfaces and atta-ches on Pb3 dorsoposterolaterally near process that
articulates with Eb4; TPb3 is narrowly continuousmid-posteriorly with SOD.M. Pb3-Eb3-Eb4, slender muscle originating most-ly on anterior surface of Pb3 process that articulateswith Eb4 and dorsally on medial cartilaginous tip ofEb4, extends laterally at right angles to long axis ofgill arches and inserts on posterior surface of Eb3ventral to tip of uncinate process.OD3 originates ventral to TEb2 on mid-lateraledge of anterior Pb3 articulating surface and at andventral to OD4 origin, and inserts on Eb3 uncinateprocess.OD4 originates ventral to TEb3 on mid-lateraledge of anterior Pb3 articulating surface and at anddorsal to OD3 origin, and inserts anteriorly on mid-dorsal flat, bony Eb4 uncinate process.OP broadly dorsally on Eb4 posteriorly and ven-trally on broad ligament joining LP and Eb4 to Cb5;completely obscures Ad4 from posterior view.Adl on Ebl bony anterior surface, extending frommedialmost end to medial edge of LEI, there cross-ing ventrally and attaching on anterior surface of dis-tal end of Cb 1
.
Ad2 on bony process on anterior edge of Eb2 andon anterior surface of cartilaginous end of Cb2.Ad3 on ventral edge of distal bony half of Eb3 andanterior surface of cartilaginous end of Cb3 (not vis-ible in illustration).Ad4 dorsally on ventral edge of Eb3, ventrally nar-rowly on Cb4 dorsally medial to Eb4-Cb4 joint; com-pletely occluded from external view.Ad5 short, dorsally on Cb4 posterodistally andCb5 posterodistalmost end.SOD slender.RDs paired on each side, ventral member of pairslightly separated from counterpart, inserts on pos-terior margin of Pb3; dorsal member of pair well sep-arated from counterpart, inserts mostly on posteriorend of UP4 with few medial fibers attaching to Pb3together with lateralmost fibers of ventral member.Additional remarks. SCL free from Bb3 (cartilag-inous posterior end not elongate or extending ven-trally). TV4 free from Cb5s. Pbl absent. Pb2 toothed.IAC present.
PleuronectiformesPSETTODIDAE
Psettodes erumei (Bloch and Schneider), USNM366443, 106 mm, eyes on left side.Plates 205.1, 205.2
Description.LEI on Ebl dorsally at base of uncinate process.LE2 on expanded dorsal edge of Eb2 at about mid-length.LE3 absent.
234 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LE4 on posteroventral surface of Eb4 uncinateprocess.LP slender, fragile filament on Eb4 at lateral edgeof LE4 insertion.LI1 on Pb2 dorsoanteriorly; anteromedially joiningmoderately long raphe with TPb2 laterally.LI2 broadly on lateral edge of Pb3 posteriorly.TD comprises TPb2, TEb2, and TEb4. TPb2 at-taches musculously dorsoanteriorly to Pb2 dorsoan-teriorly, and tendinously anterolaterally to Pbl ven-trolaterally and Ebl dorsoanterolaterally (attachmentto Pbl and Ebl unknown in other fishes); right-sideportion of TPb2 folds over left side posteriorly, andmedial edge is interrupted by raphe that gives rise toCT sheet that attaches to cranium; posterolaterally,on each side, TPb2 joins another raphe with TEb2,and muscle strands extend posteromedially from ra-phe of each side and become continuous with TEb4posteromedially. TEb2 attaches laterally along mostof anterior edge of Eb2 medial to raised dorsal Eb2margin and LE2 insertion. TEb4 attaches mid-ven-trally by CT among longitudinal fibers of long, strap-like muscle extension of SO (extension attaches toPb3 anteromedially); TEb4 ventrally continuous bymuscle filaments with SOD anteriorly.OD3-4 origin on dorsoanterior surface of Pb3; in-
sertion on Eb3 dorsoanteriorly and, beginning on me-dial edge of Eb4 uncinate process and continuing pastEb3 uncinate process, attaching broadly on Eb4 an-teriorly ventral to uncinate process.OP in medial and lateral sections. Medial sectiondorsally on Eb4 posteromedial to uncinate process;lateral section dorsally on Eb4 posteriorly ventral toLE4 insertion; medial section attaches ventrally tohook-like process on Cb5 mid-posteriorly (not visiblein illustrations) and to adjacent arm of Cb5, overlap-ping posteriorly and continuous with lateral OP sec-tion, which is on Cb5 dorsolaterally between attach-ments of TV5 and Ad5.Ad 1-3 absent.Ad4 broadly dorsally on Eb4 lateral to LE4 inser-tion, ventrally, not so broadly on Cb4 slightly medialto Eb4-Cb4 joint.Ad5 anteriorly on posterodistal ends of Eb4 andCb4, posteriorly on Cb5 anterodistally.SOD present.RDs adjacent, insert on posteromedial surface ofPb3 and anteromedial edge of UP4.Additional remarks. SCL attached mid-dorsally toelongate cartilaginous posteroventral tip of Bb3. TV4free from Cb5s. Pb4 absent, UP4 present. Pb2toothed. IAC present. Eb4 levator process absent.
NUMBER 11 235
Acknowledgments
Our study has benefitted from the generous assis-tance of many colleagues who provided material forstudy, suggestions for taxa to be included, curatorialassistance, photographs, information, and discussionof various aspects of the study. We could not havecompleted this study without their input: I. Akagawa(University of Miyazaki, Japan), specimens; G. R.Allen (WAM), specimens; M. E. Anderson (SAIAB-RUSI), specimens; C. Baldwin (USNM), discussion;D. R. Bellwood (James Cook University, Townsville,Australia), information on Scaridae; R. Britz(USNM) specimens, discussions, suggestions, partic-ularly on polycentrids, anabantomorphs. polypterids,and Dipnoi; B. A. Brown (AMNH): G. Burgess(UFMNH), specimens; K. Carpenter (ODU), speci-mens, discussion of sparoids; D. Catania (CAS),specimens; J. Clayton (USNM), specimens, curatorialassistance; B. B. Collette (NOAA, Washington,D.C.), specimens, discussion of beloniforms, scom-broids; M. C. C. de Pinna (MZUSO). discussion ofcatfishes; J. Finan (USNM), curatorial assistance;S. V. Fink (Michigan State University) and W L.Fink (UMMZ), information on cyprinids; T. H. Fraser(Port Charlotte, FL), discussion of centropomids,apogonids; R. Fritzsche (HSU), specimens, informa-tion on Icosteidae; A. C. Gill (BMNH), discussion ofpseudochromids, plesiopids. gobioids. percopsiforms;K. Hoshino (USNM), information on Psettodes; S.Jewett (USNM), curatorial assistance; L. W. Knapp(USNM). information on platycephalids; H. K. Lar-son (NTM), specimens; K. Matsuura (NSMT), spec-imens; M. McGrouther (AMS), specimens; R. M.McDowall (Christchurch, New Zealand), specimens;J. McEachran (TCWC). specimens; R. Mooi (MPM),information on plesiopids, pempherids; G. Moore(WAM). specimens; T. Munroe (NOAA. Washington,D.C.), specimens, labrid larvae; E. O. Murdy (NSF,Washington, D.C.), information on gobioids; K. Mur-phy (USNM), radiography, paperwork, dedicatedgeneral curatorial assistance; D. Nelson (UMMZ),specimens; G. J. Nelson (University of Melbourne,Australia), discussions on gill arches, gill-arch mus-cles; J. Nelson (UAMZ), specimens; J. Nielsen(ZMUC), specimens, discussions on ophidiids andbyhthids; D. Nolf (IRSNB), discussions; J. W Orr
(NOAA, Seattle. WA), information on gasterostei-forms; T. M. Orrell (USNM), computer maven; L.Palmer (USNM), curatorial assistance; L. R. Parenti(USNM), atherinomorph. mugiliomorph discussions,suggestions, specimens, and encouragement; J. Pax-ton (AMS), specimens, prodding; T. Pietsch (UW).specimens; J. E. Randall (BPBM), specimens; S. Ra-redon (USNM), radiography, photography, curatorialassistance; R. Robins (UF), specimens; R. H. Rosen-blatt (SIO), specimens; K. Sasaki (BSKU), speci-mens; J. Seigel, specimens; D. G. Smith (USNM),discussion about eels, nomenclatural discussions, cu-
ratorial assistance; S. Smith (USNM), curatorial andcomputer program assistance; W. F Smith-Vaniz(U.S. Geological Survey, Gainesville, FL), referenc-
es, information on opistognathids, carangids, cepol-ids; C. Spencer (formerly Alexandria. Virginia), art-work early in study; R. v. Sternberg (USNM), dis-cussions on epigonids, bathyclupeids, acropomatids,character coding; M. J. Stiassny (AMNH), speci-mens, information on labroids; K. Sulak (U.S. Geo-logical Survey, Gainesville, FL), specimens, infor-mation on ateleopodids; A. Suzumoto (BPBM), spec-imens; K. Tighe (USNM), information on eels; J. C.Tyler (USNM), information and discussions aboutzeiforms, tetraodontiforms, caproids; B. Urbain(UW), specimens; R. P. Vari (USNM), information onterapontids; S. H. Weitzman (USNM), information oncharaciforms; M. Westneat (FMNH), information onlabrids; J. T. Williams, ever helpful computer maven,photography, suggestions for character coding; R.Winterbottom (ROM), general discussions and rec-ommendations about gill-arch muscles, charactercoding.R. Winterbottom and G. Nelson each reviewed acomplete draft of the manuscript and performed yeo-man's work, noting many discrepancies and offeringvaluable suggestions for its improvement. We are in-debted to them for their sage advice, no less for theimportant ground work their publications have laidfor the study of fish musculature.Our special thanks to R. Von Sternberg, for en-couragement, enlightening discussions on cladisticpractice, advice on manuscript preparation, and forhis editing the MS for publication.
This study is dedicated to Dr. Herbert R. and Mrs. Evelyn Axelrod inrecognition of their over 40 years of loyal friendship with VGS and in grat-itude for their generous support of his research, which made this study pos-
sible, and for their support of systematic ichthyology generally.
236 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
APPENDIX
NUMBER 11 237
Phylogenetic analysis of the families of acanthomorph fishes based ondorsal gill-arch muscles and skeleton
Victor G. Springer and Thomas M. Orrell
"The unraveling ofphyletic lines within the Perciformes is made difficult bythe sheer number of species and genera. One is faced with the choice ofmining a narrow vein for nuggets of knowledge which He isolated, or engag-ing in a strip mining operation which reveals broad patterns at the expenseof ignoring contradictory details." Richard H. Rosenblatt (1984).
Introduction
This section comprises a phylogenetic analysis ofthe Acanthomorpha based almost exclusively on gill-arch muscle and skeletal characters examined and de-scribed in the Springer and Johnson (henceforth.S&J) portion of this study. The analysis includes 168individual taxonomic groups (Table 12) comprising147 families out of a total of approximately 250acanthomorph families and 2400 genera (increasedarbitrarily based on Nelson's 1994:253. figures). Ofthese. 15 families in the analysis are represented bymultiple taxa (e.g., all four genera of Polycentridae;three different genera of Labridae), and 132 familiesare represented individually. The individually repre-sented families include either all the taxa S&J ex-amined for the family (see the pertinent family ac-counts), which might be several genera and species.or a single genus and all its examined species, whichmight only be one (e.g., Cichlidae. Ptychochromis).In the latter case, the generic name follows the familyname on the cladograms (Figs. 5 and 6). The reasonsfor selecting a single taxon, as opposed to all thatwere examined, varied. Usually, the single taxon waschosen because a published study or colleague indi-cated that it was the basal member, or among themore basal members, of the family group that S&Jexamined. For example, of the six gobioid familiestreated in the descriptive accounts, only Rhyaci-chthyidae are entered in the matrix, based on: Miller(1973), Springer (1983), and Hoese and Gill (1993).For Opistognathidae, Lonchopisthus, was suggestedby W.F. Smith-Vaniz (pers. comm.). In other caseswe selected taxa that might include a relatively un-specialized taxon in a group for which a basal mem-ber has not been hypothesized (e.g., Labridae, Am-bassidae).Fifty-six characters (1-36, 36a, 37-55), compris-ing 137 character states, were used in preparing Table12. Characters l-36a refer to gill-arch muscles, 37-55 are osteological, mostly gill-arch skeletal charac-ters examined during the descriptive part of the study.Character 36a was added after all other charactershad been coded and rather than re-number characters37-55, we chose to interpolate the additional char-acter as 36a.
S&J described character 36 for many, but not all,taxa. The character was observed in the remainingtaxa and coded directly into the matrix. Similarly,S&J did not describe character 43 in any taxa. Be-cause it pertains to the dorsal gill arches, and wasnoticed during the study, the character was examinedfor all taxa for the Appendix and coded directly intothe matrix.Character states were coded based on the states inSynagrops (Acropomatidae), a relatively unspecializedpercomorph, because it was initially intended to ana-lyze only percomorphs. Non-percomorphs were addedand rather than re-code based on a non-percomorphwe continued to code based on Synagrops.Character 53 is the single synapomorphy Johnsonand Patterson (1993) used to hypothesize their Smeg-mamorpha. We did not verify the character states forcharacter 53 in our study, but infer them from John-son and Patterson (1993). We added this character inorder to enable a stronger test of the monophyly ofthe Smegmamorpha.Among acanthomorphs, character 55 is a uniquesynapomorphy of the Anabantomorpha and character54 is confined to the Anabantoidei. We added thesetwo characters in an attempt to restrict the anaban-tomorph taxa included in the analysis.
Parsimony AnalysisA PAUP NEXUS file was developed from thecharacters described below. The file contained PAUPcommands and a matrix of 168 taxa by 56 characters,or 9408 cells. Multi-state characters were assignedsingle symbols using the PAUP "equate" function.Although character states were coded based on 5w-agrops (Acropomatidae). characters were not orderedand were given equal weight for the parsimony anal-ysis because of the large number and diversity of taxainvolved. Each of the 56 characters was informativeunder parsimony. States were unknown or inappli-cable for several characters. Both states are treatedas "missing" by the parsimony algorithm, becauseunder the algorithim there is no way to differentiatebetween them (Swofford 2003). Characters withmissing data do not affect the placement of taxa onthe tree. Trees were mid-point rooted. This method
238 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of tree drawing roots at the midpoint of the longestpath linking any pair of taxa. Functionally, this pro-duces a tree that is not explicitly rooted. However,the use of this option in PAUP* forces MINF re-optimization of the tree (the only optimization meth-od which is not dependant on root position).Parsimony analyses were conducted using PAUP*version 4.0bl0 (Swofford 2003) and were run on anAlpha 64 bit processor running RedHat Linux ver-sion 7.1 Alpha. The analysis was run with the fol-lowing parameters: starting trees obtained via step-wise addition; swapping algorithm tree bisection-re-connection; no topological restraints; all trees un-rooted; character-state optimization acceleratedtransformations (MINF optimization was used todraw trees); MulTrees option in effect; starting seed
= 524970930; heuristic search of 1000 random ad-dition replicates with 10 trees held at each step duringstepwise addition; no more than 100 trees of score(length) greater than or equal to 602 saved in eachreplicate; number of rearrangements (per addition-se-quence replicate) limited to 1,000,000.000.Because of the large size of the matrix and limitedprocessing power, it was necessary to restrict theanalysis in order to complete it. For example, we at-tempted to run an analysis of 1 unrestricted randomaddition replicates. After 240 hours (10 days), thesearch had not proceeded past the first random ad-dition replicate. The search tried >2.04 X 109 rear-rangements and had saved 180,550 equally parsi-monious trees of 598 steps (of which it still had179,005 remaining to be swapped). We chose to runan analysis in which multi-state characters for anytaxon were interpreted as uncertain. This allowedPAUP to select the variable state that minimized treelength. A heuristic search consisting of 1000 randomaddition sequences was performed. In order to in-crease the chance of improving the tree score foundby stepwise addition, 10 trees were held at each step.Only 1000 trees were held for each step greater than602 steps. This value was determined by conductingnumerous single random addition sequence searches.In all of these searches, trees of at least 602 stepswere generated. Therefore, 602 was set as an upperlimit to restrict searching of sub-optimal trees, whichresulted in decreasing the overall time required tocomplete all 1000 replicates. In addition, the maxi-mum number of branch swapping rearrangementswas limited to 100.000.000 per repetition.We recognize the inherent problems of placing toogreat a reliability on parsimony analyses of large datamatrices, especially when there are relatively fewcharacters per taxon and considerable homoplasy. Webelieve it worthwhile, however, to have the results ofan analysis that is restricted essentially to charactersderived from a single character-rich complex, dorsalgill-arch muscles and skeleton, that has received rel-
atively limited attention in the past (Johnson 1993:27). In spite of the large amount of homoplasy in-volved, our analysis clearly represents stronger testsof the monophyly of the Smegmamorpha than John-son and Patterson's (1993; henceforth, J&P) hypoth-esis based essentially on a single character (includedin our analysis), mode of articulation of the anterior-most epineural. It may not be, however, an adequatetest of Stiassny and Jensen's (1987) hypothesizedmonophyly of the Labroidei also based essentially ongill-arch skeletal and muscle characters (included inour analysis, but comprising only a few of the char-acters we address). Additionally, there is heuristicvalue in the clades we have found, which will pro-vide bases for others to test or derive support.
Characters (Numbers in Brackets) and CharacterStates (Numbers in Parentheses)
[1] LEI inserts completely medial to lateral fourthof surface of Ebl (0); insertion includes all orpart of lateral fourth of surface (1); LEI absent(2).[2] LEI origin groups with other LE origins (0);separate from other LE origins (1); not appli-cable (-).[3] LE3 present (0); vestigial or absent (1).[4] LP and LE4 insert together on Eb4 (0); inser-tions well separated on Eb4 (1); LE4 and/or LPsheet-like, usually continuous with PP. withbroad attachment continuing from Eb4 and Cb5along length of Cb4 (2); LP absent (3); LE4absent.[5] LP does not insert by long, slender tendon (0);inserts by long, slender tendon (1); not appli-cable (-).[6] LI1 only on Pb2 (0); on Pb2 and Pb3 (1); onlyon Pb3 (2); on Pb2 and IAC (3); not applicable(-)[7] LI1 slightly, if at all, larger than LI2 (0); muchlarger than LI2 (1).[8] LI2 does not penetrate OD (0); penetrates OD(1).[9] TD muscles essentially completely cover Pb3sdorsally (0); TD muscle broadly replaced or in-terrupted medially by CT dorsal to Pb3s, Pb3susually with fiat, dorsal articulating facets,which do not form diarthrosis with process onventral surface of cranium (1); TD musclesbroadly replaced medially by CT dorsal to mod-erate-sized Pb3 dorsal articulating facets, facetsjoin diarthrosis with irregular-surfaced knob-like process on ventral surface of cranium (2);TD muscle broadly replaced medially by CTdorsal to relatively large Pb3 dorsal articulatingfacets, facets form diarthrosis with smooth, con-
NUMBER 1
1
239
forming, usually weakly bilobed ventral surfaceof process on ventral surface of cranium (3).[10] TEb2 right and left sides do not overlap medi-ally (0); overlap medially ( 1 ); TEb2 absent (2).[11] TEb2 extends laterally well past medial edge ofLE2 insertion (0); does not extend laterallymuch, if any, past medial edge of LE2 insertion
( 1 ): inapplicable (-).[12] TPb2 insertion not on IAC (0); insertion in-cludes IAC (1); TPb2 vestigial or absent (2).[13] TPb2a absent (0); present ( 1 ).[14] TD attachments include Eb3 (0); exclude Eb3(1).[15] TD not on Eb4 (0): on Eb4 ( 1 ).[16] TDP attaches to Pb3 (0), does not attach to Pb3(1).[17] TD not attached to Pb4 when Pb4 is present (0);attached to Pb4 ( 1 ); Pb4 absent (2).[18] SOD present (0); absent (1).[19] CPb absent (0); present ( 1 ).[20] OD origin only on Pb3 (0); on Pb2 and Pb3 (1).[21] OD3' absent (0); present (1).[22] OD insertion includes Eb3 (0); excludes Eb3(1).[23] OD on Eb4 (0): not on Eb4 (1).[24] OD4' absent (0); present (1).[25] OP posteroventrally mostly to entirely on Cb5(0); wholly or partly on Cb4 and/or joining ERat level of Cb4 (1); OP absent (2).[26] Neither OP nor Ad4 joins either LE4 or LPwhen both LE4 and LP are present (0); OP joinsonly LE4, which is not released from Eb4; Ad4not involved (1); OP joins only LE4, which ismostly released from Eb4; Ad4 not involved(2); OP joins only LP; Ad4 not involved (3);OP joins both LP and LE4, which is not re-leased from Eb4; Ad4 not involved (4); OPjoins both LP and LE4, which is released fromEb4; Ad4 not involved (5); Ad4 joins only LE4;OP not involved (6); Ad4 joins only LP; OPnot involved (7); Ad4 joins both LP and LE4;OP not involved (8); Inapplicable (-).M. Pb2-Eb2 absent (0); present ( 1 ).M. Pb3p absent (0); present (1).GFM 1 moderately developed or absent, not fan-like (0); enlarged, fan-like (1).Adl absent (0); present (1).Ad2 and Ad3 absent (0); present (1).Ad5 dorsally or anteriorly: attaches only to dis-tal end of Cb4 and/or AC4 (0); attaches wellmedial to distal end of Cb4 and/or to ER wellmedial to distal end of Cb4 (1); attaches onlyor more than incidentally to Eb4 (2); Ad5 ab-sent (3).[33] ER absent (0); present ( 1 ).[34] Ad5 joins neither LP nor LE4 (0); joins LE4and/or LP (1); inapplicable (-).
[35]
[36]
[27][28][29]
[30][31][32]
[36a[37]
[38]
[39][40]
[41]
[42][43]
[44]
[45][46][47][48][49][50][51]
[52][53]
[54]
[55]
TV4 free from Cb5s (0); attached to Cb5s (1);TV4 absent (2).PCI attachment on Cb5 begins well medial todistal end and extends medially (0); attachmenton Cb5 begins at distal end and extends medi-
ally, muscle does not join raphe with OP ven-trally (1); attachment on Cb5 begins at distalend and extends medially, muscle joins raphewith OP ventrally (2).
| M. Pb3-Cb5 absent (0); present (1).Pbl bony and cartilaginous, articulates with an-terior Ebl process (0); Pbl all cartilaginous, ar-ticulates with anterior Ebl process (1); Pbl ab-sent, anterior Ebl process present (2); Pbl ab-sent, Ebl anterior process absent (3).Pb2 with teeth (0); Pb2 edentate (1); Pb2 absent(2).UP4 present (0); absent (1).Cartilage-tipped Ebl uncinate process present(0); absent ( 1 ).Cartilage-tipped Eb3 uncinate process present(0); absent (1).Eb4 bony flange absent (0); present ( 1 ).Medial end of Eb4 equal to or smaller than me-dial end of Eb3 (0); larger than medial end ofEb3 (1). Character not discussed by S&J; ob-servations coded directly in matrix.Cartilage-tipped Eb4 uncinate process present(0); absent (1).Levator process present on Eb4 (0), absent (1).AC1 absent (0); present (1).AC2 absent (0); present (1).AC3 absent (0): present (1).AC4 present (0); absent (1).IAC (or IAB) present (0); IAC absent (1).Cb5s not fused together (0); fused but suturepresent (1); fused but suture absent (2).SCL present (0); absent (1).Anteriormost epineural rib does not articulatewith distal end of transverse process (0); artic-ulates with distal end of transverse process (1).Ebl not modified as suprabranchial organ (0);modified as suprabranchial organ (1).Parasphenoid teeth absent (0); present (1).
Results
The results are summarized in two cladograms,one, a strict consensus tree (SCT; Figure 5) derivedfrom the 1823 most parsimonious (retained) trees,each with 595 steps, found during the PAUP analysis,and the other, a 50 percent majority-rule tree (MRT;Figure 6). In the SCT, there are 77 non-terminalnodes (those present in all 1823 trees retained) andin the MRT there are 145 non-terminal nodes.The large number of taxa surveyed made it im-practical to show character and nodal support directly
240 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 12.?Character-state matrix for selected acanthomorph taxa examined for present study. Family name followd by genus indicatescharacters coded only for genus; otherwise coding refers to all taxa examined. Characters 1-52 are those examined during present study.Assignment of states for character 53 are inferred from Johnson and Patterson (1993:table 2, character 3); 54 is based on Nelson (1994:432, compiled): and 55 is based on R. Britz (pers. comm.) and literature. Other character states are assigned based on Synagrops(Acropomatidae), as a single outgroup. Key: A = (01); B = (02): C = (03); D = (06); E = (12); - = (not applicable); ? = (missing).
Characters000 000 000 111 111 111 122 222 222 223 333 333 333 344 444 444 445 555 55TAXA 123 456 789 012 345 678 901 234 567 890 123 456 6783 . . 901 234 567 890 123 45Acropomatidae 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 00Percichthyidae 000 000 000 000 000 000 000 000 010 000 000 000 000 000 000 000 000 000 00Pempheridae 000 000 000 001 000 000 000 000 000 000 000 000 000 000 000 A00 000 000 00Glaucosomatidae 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 100 000 000 00Ammodytidae 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 100 000 000 00Terapontidae, Terapon 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 000 00Terapontidae, Leiopolherapon 000 000 000 000 000 000 001 000 080 000 000 000 000 000 000 000 000 000 00Girellidae 000 100 000 000 00A 010 000 000 000 000 000 000 000 000 010 000 010 000 00Kuhliidae 000 003 000 001 000 010 000 000 000 000 000 000 000 000 010 000 000 000 00Epigonidae 000 000 000 001 000 000 00A 000 000 000 000 000 000 000 000 000 000 000 00Lactariidae 000 000 100 000 000 000 000 000 000 000 000 000 000 000 000 000 010 000 00Bathyclupeidae 000 003 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 010 00Ostracoberycidae 000 000 000 000 000 000 000 000 060 000 000 000 000 000 000 100 010 000 00Sebastidae 000 000 000 010 001 000 000 000 000 000 000 000 000 000 000 000 000 000 00Scorpaenidae 000 00A 000 010 001 000 000 000 000 000 000 000 000 000 000 000 010 000 00Platycephalidae, Platycephalus 000 001 000 002 011 000 010 000 000 000 010 000 000 000 000 000 010 000 00Lutjanidae 000 000 000 001 000 AA0 000 000 000 000 000 000 000 000 000 000 010 000 00Centrarchidae, Micropterus 000 000 000 000 000 010 000 000 000 000 000 000 000 000 000 000 010 000 00Moronidae 000 000 000 001 000 010 000 000 001 000 000 000 000 000 000 00A 000 000 00Latidae 000 000 100 000 000 000 000 000 060 000 000 000 000 000 000 000 000 000 00Centropomidae 000 000 100 000 010 000 000 000 060 010 000 000 000 000 000 1A0 A00 000 00Caristiidae 000 000 000 000 000 000 000 000 080 000 000 000 000 000 000 100 Oil 000 00Apogonidae, Glossamia 000 000 000 000 000 000 000 000 000 000 000 000 000 000 100 A00 010 000 00Kurtidae 000 001 000 000 000 120 000 000 000 000 000 000 000 000 000 100 010 000 00Cirrhitidae, Paracirrhites 000 000 000 000 010 000 000 000 000 000 000 002 000 000 010 000 010 000 00Percidae, Perca 000 001 000 000 000 010 010 000 000 000 000 000 000 000 010 100 010 000 00Percidae, Percina 000 001 000 000 000 010 010 000 001 001 100 000 030 000 010 100 01A 000 00Cepolidae, Cepola 000 001 000 002 000 000 000 000 000 000 000 000 000 000 000 000 000 000 00Serranidae 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 010 00Symphysanodontidae 000 000 000 01B 000 000 00A 000 000 000 000 000 000 000 000 A00 0A0 000 00Nematistiidae 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 100 000 000 00Malacanthidae 000 000 000 000 000 00A 100 000 000 000 000 000 000 000 000 000 010 000 00Carangidae, Selar 000 000 000 000 001 010 001 000 070 000 000 000 000 000 100 000 000 000 00Priacanthidae 000 000 000 002 011 001 000 000 000 000 000 000 000 000 000 000 0A0 010 00Centracanthidae 000 100 000 000 000 010 101 000 000 001 100 000 100 010 100 000 010 000 00Haemulidae 000 003 A00 002 000 011 001 000 070 000 000 000 000 000 100 000 000 010 00Inermiidae 000 000 000 002 000 011 001 000 070 000 000 000 000 000 100 000 000 010 00Nemipteridae 000 000 000 000 000 110 101 000 000 000 000 002 000 010 001 000 Oil 000 00Lateolabracidae 000 003 100 002 000 010 001 000 000 000 000 000 000 000 000 000 010 000 00Callanfhiidae 000 000 000 002 0A0 010 00A 000 000 001 100 000 010 000 001 000 010 0A0 00Bramidae 001 003 000 000 011 000 000 000 000 000 000 000 000 000 000 000 010 000 00Icosteidae 000 001 000 010 001 100 000 000 100 000 011 000 000 A00 000 1A0 0A0 000 00Centrolophidae 000 000 001 000 000 011 000 000 700 000 000 000 000 000 000 000 010 000 00Amarsipidae 000 000 000 000 010 010 001 000 000 000 000 000 000 000 000 100 010 000 00Mullidae 000 000 000 001 000 020 001 000 000 000 020 000 020 000 000 10A 010 010 00Sciaenidae, Cynoscion 000 003 100 002 000 000 000 000 000 000 000 002 000 000 000 100 010 010 00Rhyacichthyidae 100 001 000 002 010 010 000 000 000 010 100 000 010 000 111 100 010 000 00Cichlidae, Ptychochromis 000 001 103 001 111 021 100 000 050 001 120 010 000 000 110 100 010 100 00Pomacentridae, Dischistodus 000 001 102 000 110 021 101 000 010 001 100 010 000 000 100 100 010 200 00Pomacentridae, Amphiprion 000 001 102 000 110 021 101 000 040 001 100 010 000 000 100 0AA 010 200 00Labridae, Achoerodus 000 002 003 2-0 111 021 100 000 050 001 120 010 011 101 110 111 111 200 00Labridae, Choerodon 000 002 003 2-0 111 021 100 000 050 001 120 010 011 101 110 10A Oil 200 00Labridae, Coris 000 002 003 010 111 021 100 000 050 001 120 010 011 101 110 111 111 200 00Embiotocidae, Amphisticus 000 001 003 002 011 020 000 000 050 001 120 010 001 000 100 100 0A1 200 00
NUMBER 1
1
Table 12.?Continued.
241
Characters000 000 000 111 111 111 122 222 222 223 333 333 333 344 444 444 445 555 55TAXA 123 456 789 012 345 678 901 234 567 890 123 456 678 901 234 567 890 123 45
Embiotocidae, Embiotoca 000 000 003 002 011 021 000 000 050 001 120 010 001 000 100 000 001 200 00Pholidichthyidae 000 402 101 010 111 021 100 000 0-0 001 100 010 001 100 111 100 Oil 200 00Lethrinidae 000 110 000 000 0AA 0A0 101 000 000 001 100 00B 000 000 A01 000 010 010 00Sparidae 000 000 000 000 000 010 A01 000 000 001 100 000 100 000 100 000 010 000 00Draconettidae 000 002 000 002 000 120 100 000 000 001 120 000 010 110 001 100 Oil 000 00Champsodontidae 000 002 010 002 011 101 000 000 000 000 000 000 020 000 000 100 010 000 00Uranoscopidae, Xenocephalus 001 001 000 012 000 020 000 000 000 000 000 000 011 000 000 000 Oil 010 00Bovichtidae 000 001 000 002 010 010 001 000 000 001 100 000 030 000 000 100 Oil 000 00Pseudaphritidae 000 000 000 002 010 110 000 000 000 000 000 000 030 000 000 100 Oil 000 00Cheimarrichthyidae 000 001 000 002 000 000 000 000 000 000 000 000 010 000 000 100 010 000 00Trachinidae 000 001 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 000 00Plesiopidae, Assessor 000 001 10A 010 000 020 000 000 000 000 000 000 000 000 100 A00 010 000 00Grammatidae 000 001 101 002 010 010 000 000 000 000 000 010 000 000 100 000 010 000 00Opistognathidae, Lonchopisthus 000 001 001 002 001 000 000 000 000 000 000 010 001 000 10A 000 010 000 00Pseudochromidae 000 001 101 002 10A 000 000 000 000 001 120 010 0E0 000 100 000 01A 000 00Centrogeniidae 000 001 001 000 010 000 000 000 010 000 000 012 001 000 001 000 000 000 00Ambassidae, Ambassis 000 001 001 001 000 021 000 000 000 000 000 000 000 000 000 100 0A0 000 00Ambassidae, Telracentrum 000 001 001 001 000 011 000 000 010 000 000 000 000 000 100 100 A10 000 00Bathymasteridae 000 001 100 000 000 020 000 000 000 000 000 000 030 000 000 100 Oil 000 00Anoplopomatidae 000 001 100 000 000 000 000 000 000 001 100 000 030 000 000 100 Oil 000 00Stichaeidae 000 001 100 002 000 000 001 000 000 001 100 000 030 000 000 100 Oil 000 00Hexagrammidae 000 001 000 000 000 000 000 000 000 000 000 000 010 000 000 100 010 000 00Gerreidae 000 001 001 002 001 000 00A 000 000 001 100 000 000 000 1A0 100 010 000 00Polynemidae 000 00A 000 010 000 010 000 000 000 000 000 000 000 000 010 100 010 010 00Toxotidae 000 000 001 001 000 010 000 000 010 000 000 000 000 000 100 000 010 000 00Polycentridae, Afronandus 001 001 100 002 001 021 000 000 000 000 000 000 000 000 000 100 010 000 00Polycentridae, Monocirrhus 001 001 100 002 001 021 001 000 000 000 010 000 000 000 0A0 100 010 000 00Polycentridae, Polycentropsis 001 001 000 002 001 021 000 000 000 000 000 000 000 000 000 100 010 000 00Polycentridae, Polycentnis 001 001 100 000 001 021 000 000 000 000 000 000 000 000 000 000 010 000 00Nandidae 001 000 000 002 001 021 000 000 000 001 100 000 000 010 100 000 010 000 01Badidae 001 000 001 002 011 021 000 000 070 001 100 000 010 010 100 A00 010 000 01Pristolepidae 000 000 001 002 011 121 000 000 000 000 020 001 000 010 010 000 010 000 01Channidae 000 002 000 012 011 021 100 000 030 000 1B0 001 000 010 111 100 Oil 000 11Anabantidae, Sandelia 000 002 001 2-0 010 021 100 000 000 000 020 001 000 0A0 111 100 01A 000 11Caproidae, Amigonia 000 010 000 002 001 020 001 010 000 000 000 000 000 000 000 100 01A 000 00Caproidae, Capros 100 3-0 000 012 001 120 001 010 060 001 100 000 000 000 000 100 Oil 000 00Sillaginidae 000 003 000 000 000 000 001 000 001 000 100 000 000 000 100 101 010 000 00Dactylopteridae 000 002 100 000 000 020 000 000 000 001 100 000 010 000 000 100 Oil 000 00Mugilidae, Agonostomus 010 000 000 Oil 010 02A 000 000 000 000 120 000 000 000 100 100 010 001 00Bedotiidae 010 100 001 2-0 111 021 000 000 020 000 120 000 000 000 110 100 010 001 00Atherinidae 010 100 001 2-0 111 021 000 100 030 001 120 000 000 000 110 100 010 001 00Adrianichthyidae, Xenopoecihis 111 100 001 2-0 110 021 000 100 000 000 120 000 020 110 111 100 010 001 00Cyprinodontidae 111 102 001 2-0 111 021 000 100 000 001 100 020 020 110 110 100 010 001 00Aplocheilidae 110 000 001 2-0 110 021 000 100 000 001 120 010 020 010 110 100 010 001 00Hemiramphidae 010 100 001 2-0 111 021 000 100 020 100 100 010 020 100 110 1A1 010 201 00Exocoetidae 110 000 001 2-0 111 021 000 100 020 100 100 010 020 110 110 1A0 010 201 00Belonidae, Strongylura 110 000 001 2-0 ?11 020 000 000 001 000 13- 010 020 111 111 110 A?l 201 00Belonidae, Tylosurus 111 000 000 2-0 ?11 020 000 100 001 000 13- 010 020 111 111 1A0 0?1 201 00Scomberesocidae 011 000 001 2-0 110 020 000 100 0C1 000 100 010 020 100 Oil 1A0 Oil 201 00Elassomatidae 100 001 100 002 001 020 000 000 000 001 100 000 030 000 001 100 Oil 001 00Gasterosteidae, Apeltes 001 001 000 012 001 020 000 000 000 001 120 100 030 100 010 100 Oil 001 00Gasterosteidae, Culea 001 001 100 012 001 020 000 000 000 001 120 100 030 100 000 100 Oil 001 00Gasterosteidae, Gasterosleus 001 001 100 012 001 020 000 000 000 001 120 100 030 100 000 100 Oil 001 00Gasterosteidae, Pungitius 001 001 100 012 001 020 000 000 000 001 120 100 030 100 010 100 Oil 001 00Gasterosteidae, Spinachia 001 001 100 012 001 020 000 000 000 001 120 100 030 100 010 100 Oil 001 00Aulorhynchidae 101 001 100 012 001 020 001 000 070 001 100 000 030 000 000 100 Oil 001 00Hypoptychidae, Aulichthys 100 001 000 012 010 020 010 000 000 001 100 000 030 100 000 100 Oil 001 00Hypoptychidae, Hypoptychus 100 3-0 000 002 010 000 010 000 060 001 120 100 030 100 000 100 Oil 001 00
242
Table 12.?Continued.
BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Characters000 000 000 111 111 111 122 222 222 223 333 333 333 344 444 444 445 555 55TAXA 123 456 789 012 345 678 901 234 567 890 123 456 678 901 234 567 890 123 45
Synbranchidae 101 3-- 100 012 001 020 000 000 0-0 001 100 000 021 010 111 100 010 Oil 00Mastacembelidae 001 3-1 100 012 000 000 000 000 0-0 001 100 002 030 010 010 100 Oil Oil 00Centriscidae 001 000 000 002 000 020 Oil 000 100 010 021 000 010 010 000 100 Oil 001 00Leptobramidae 000 000 000 001 001 000 001 000 000 000 000 001 000 000 000 000 000 000 00Tripterygiidae, Ruanoho 100 002 000 012 010 021 000 000 000 001 100 Oil 022 111 001 100 Oil 000 00Tripterygiidae, LepidoblenniusDactyloscopidae 100 002001 002 001 012 010001 0A2 011 021 000 000 030 001020 000 000 000 000 100 112 022 111 Oil000 Oil 022 111 Oil 100 Oil 000 00100 Oil 000 00Chaenopsidae 000 002 001 002 011 020 000 000 000 000 000 012 012 111 001 100 Oil 000 00Labrisomidae, Calliclinus 000 002 001 000 011 021 000 000 010 000 020 012 012 111 001 100 Oil 000 00Climdae, Clininae A00 002 001 002 011 020 000 000 040 000 000 012 0E2 111 0A1 100 Oil 000 00Clinidae, Myxodinae 100 002 001 002 011 020 000 000 0A0 000 020 012 012 111 Oil 100 Oil 000 00Blenniidae 100 002 00A 002 01A 020 000 A00 000 001 100 012 022 111 Oil 100 Oil 000 00Berycidae 000 010 000 010 00A 020 000 000 100 000 001 000 000 000 010 100 001 010 00Trachichthyidae 000 3-0 000 010 000 020 000 000 1-0 000 001 000 000 100 010 100 Oil 010 00Grammicolepidae 100 011 100 000 000 121 001 010 100 000 010 000 000 110 010 000 Oil 010 00Oreosomatidae 101 001 000 010 000 121 001 010 100 000 Oil 000 000 110 010 100 Oil 000 00Triacanthodidae 000 010 002 000 000 020 000 010 000 010 000 000 000 000 000 100 010 000 00Menidae 000 200 000 012 000 021 001 010 100 000 001 000 000 000 000 100 100 000 00Leiognathidae 100 0-0 001 002 00A 100 001 010 050 000 000 010 000 010 100 100 Oil 000 00Ephippidae, Chaeiodipterus 100 010 000 002 001 100 001 010 060 000 000 000 000 0A0 100 100 00A 000 00Luvaridae 000 200 000 002 011 000 000 010 010 000 000 000 000 000 000 100 010 000 00Zanclidae 100 200 100 002 001 100 010 010 000 000 000 000 000 000 001 100 Oil 000 00Acanthuridae 100 2?0 000 002 011 100 000 010 0?0 010 000 000 000 010 000 100 001 000 00Holocentridae 000 000 100 000 011 000 000 000 100 000 Oil 000 000 000 010 100 Oil 010 00Aphredoderidae 001 000 000 2-0 011 101 010 001 001 000 000 000 000 000 010 000 Oil 010 00Percopsidae 001 000 000 2-0 011 000 010 101 001 000 000 000 000 000 010 000 Oil 010 00Amblyopsidae 101 000 000 2-0 011 100 010 101 001 000 000 000 030 000 010 000 Oil 010 00Stephanoberycidae 000 000 000 010 010 000 001 000 160 000 Oil 000 001 101 001 100 Oil 000 00Gibberichthyidae 000 000 000 2-0 010 000 000 000 160 000 Oil 000 001 000 0A0 100 Oil 000 00Rondeletiidae 000 000 000 2-0 010 000 000 000 100 000 Oil 000 001 000 000 100 Oil 000 00Cetomimidae 2-1 000 000 010 001 100 000 000 000 000 030 -00 001 001 Oil 100 Oil 010 00Barbourisiidae 000 000 000 010 010 000 000 000 110 000 Oil 000 000 000 010 100 Oil 000 00Melamphaidae 000 000 000 010 000 021 000 000 100 000 001 000 000 000 010 100 Oil 010 00Lampridae 000 2-0 000 102 001 100 Oil 000 700 000 00? 000 000 010 001 00A Oil 000 00Veliferidae, Velifer 000 3-0 000 102 001 010 000 000 0-0 000 000 000 000 000 Oil 000 101 000 00Polymixiidae 000 010 100 010 000 000 000 000 100 000 Oil 000 000 000 000 000 Oil 000 00Gobiesocidae 001 002 000 012 011 020 000 000 000 001 120 001 022 111 001 100 Oil 010 00Rhamphocottidae 001 001 000 000 000 020 001 000 000 000 010 000 000 100 000 100 Oil 000 00Ranicipitidae 001 002 100 002 001 020 000 000 100 000 Oil 000 010 100 010 100 Oil 010 00Batrachoididae 001 3-1 000 002 001 121 000 000 0-0 000 010 000 000 100 000 100 010 000 00Ophidiidae, Brotula 000 3-0 110 010 000 100 001 000 161 000 Oil 000 000 010 000 100 Oil 010 00Ophidiidae, Dicrolene 000 3-0 000 2-0 000 0A0 000 000 1-1 000 Oil 000 000 000 000 100 010 010 00Bythitidae, Calamopleryx 000 3-0 110 2-0 000 020 000 000 1-1 000 Oil 000 030 000 000 100 Oil 010 00Chaunacidae 001 3-2 000 010 011 120 000 000 -Dl 000 030 -00 000 100 000 100 Oil 010 00Scombridae, Scomber 000 3-0 000 010 000 020 001 000 0-0 000 000 000 000 000 010 100 000 000 00Sphyraenidae 000 000 100 010 000 020 000 000 000 000 000 000 000 000 010 100 010 010 00Pomatomidae 000 000 000 Oil 000 000 001 000 010 000 000 000 000 000 000 000 010 000 00Scombrolabracidae 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 010 00Rachycentridae 000 100 100 002 000 010 001 000 000 000 000 000 000 000 000 000 000 000 00Coryphaenidae 000 100 000 000 000 100 001 000 000 000 000 001 000 000 000 100 000 010 00Zaproridae 000 001 100 000 000 000 000 010 000 000 000 000 030 000 000 100 Oil 000 00Psettodidae 001 000 000 010 Oil 120 000 000 000 000 000 000 000 000 000 100 010 000 00Zeniontidae 001 ??1 100 000 000 121 001 000 ??0 000 00? 000 000 110 010 100 Oil 000 00Parazenidae 001 002 100 2-0 000 021 001 000 000 000 000 000 000 110 010 100 Oil 000 00Anomalopidae, Anomalops 000 000 100 010 000 020 000 000 160 000 001 000 000 100 010 100 010 000 00Anomalopidae, Photoblepharon 000 000 000 010 001 020 000 000 160 000 001 000 000 100 010 100 010 000 00
NUMBER 1
1
243
on the subsequent phylogenetic trees. We chose torepresent these values for the MRT in Table 13, rath-er than to enlarge the tree and split it across multiplepages. In the MRT (Fig. 6) we designated four majorclades with letters (A, B, C, D) and then labeled allinternal nodes with numbers in boxes (starting fromthe top left hand side of the tree and sequentiallyworking towards the terminal nodes whenever pos-
sible). In Table 13, we chose only to show those char-acters that support a node unambiguously and in-cluded both uncontradicted synapomorphic and ho-moplastic characters. We have given the character
state for each supporting character, but have notshown a direction of change, as all data were rununordered.The cladograms include a mix of interrelationshipsthat have been suggested or formally hypothesizedby other workers (usually based on few. if any, ofthe characters used in our analysis) and some thathave never been proposed. Among the latter aremany we suspect are unreasonable, but some of theseprobably deserve serious consideration. We believethat the SCT obscures much of the phylogenetic sig-
nal; therefore, the following discussion is basedmainly on the MRT. We restrict most of our discus-sion to those clades we believe are most informativeor corroborated by other studies. The interrelation-ships implied by the PAUP analysis had no influenceon S&J's suggested interrelationships.Bolded letters in brackets [ ] refer to the four majornodes in the MRC (Fig. 6); numbers following thebolded letters refer to small boxed numbers of non-terminal nodes included in the major node, e.g., [A13].Lampridiformes (Veliferidae, Lampridae), Acan-thuroidei, Caproidae, Leiognathidae [B 40]. TheMRC retrieved the two families of lampridiforms asa sister group [B 41], which is sister to a clade [B42] comprising all the acanthuroids (except Luvari-dae) we examined, Leiognathidae, and Capros, oneof the two genera of Caproidae. Luvaridae are in-cluded in a larger clade [B 32] containing the taxamentioned, but they are well removed stepwise fromthem. Both Tyler et al. (1989) and Winterbottom(1993a), hypothesized Luvaridae as a member of theAcanthuroidei in well-defended phylogenetic analy-
ses, and we believe their affinities are with the acan-thuroids. The other genus of caproid, Antigonia, wasretrieved as sister group to the Triacanthodidae (Par-ahollardia) in a clade [D 120] well removed fromthat containing Capros.S&J discussed Tyler et al.'s (2003) study in whichthe latter's preferred cladistic analysis retrieved bothcaproid genera, the tetraodontiform genus Parahol-lardia and their other outgroups in a polytomy withthe zeiforms. If acanthuroids, among which Caproswas retrieved in our analysis, and tetraodontiforms,
with which Antigonia was retrieved, are perciforms,J&P's claim that caproids are perciforms (or at leastpercomorphs) is corroborated, although caproidmonophyly remains problematic.A caproid-tetraodontiform relationship has alsobeen found in two recent molecular studies, Chen et
al. (2003) and Miya et al. (2003). Chen et al. (2003),who included only Capros, presented the results ofthree different molecular analyses and a simultaneousanalysis (their fig. 5) of the data on which the threeseparate analyses were based. The simultaneous anal-ysis shows Capros as the sister group of the tetrao-dontiform Mola. One of the other three analyses hasa clade with Capros, Drepane (an acanthuroid rela-tive), and Mola. The other two analyses link Caproswith a pleuronectiform or syngnathiform. Miya et al.(2003), who included only Antigonia, retrieved it assister to the tetraodontifom genera Stephanolepis(Monacanthidae) and Sufflamen (Balistidae), a cladedeeply nested among their percomorphs.Considering all the studies together, the evidencehints at the possibility that there is a monophyleticgroup comprising caproids, acanthuroids, and tetrao-dontiforms. A close relationship between the last twogroups was discussed and rejected by Rosen (1984),but we strongly believe deserves further study.The interrelationships of the Leiognathidae havenever been seriously investigated. They have beenplaced near the Menidae in classifications beginningwith Bleeker (1859:xxiii), who included them withthe fossil genus Acanthonemus Agassiz. an acanthu-
roid, (J.C. Tyler, pers. comm.), as the only membersof his family Equulidae (= Leiognathidae, see dis-cussion of proper family name in S&J). S&J hypoth-esized that the Menidae, which along with the Leiog-nathidae, have long been considered to be perci-forms, are probably a pre-percomorph group, and ourcladistic analysis corroborates that by retrievingMenidae well nested in a clade [D 130] containingberyciforms and stephanoberyciforms. We believe,however, that there is a good possibility that theleiognathids' closest relationhips, as evidenced byour analysis, are with the acanthuroids.Polymixiiformes. The MRT retrieved the Poly-mixiidae as part of a clade [D 137] that also includesa beryciform (Holocentrus) and two of the Paracan-thopterygian groups, a gadiform (Raniceps) and theophidiiforms. The sister group of this clade [D 134]contains all but one of the stephanoberyciforms, Ce-tomimidae [B 36], included in our study.J&P (1993:fig. 24) hypothesized (Lampridiformes(Polymixiidae (Paracanthopterygii))) as the three bas-almost groups of the Acanthomorpha. Wiley et al.,(2000:fig. 6), in a combined molecular-morphologicalstudy, hypothesized the same two basalmost acantho-morph groups, but retrieved the Paracanthopterygiias polyphyletic: ((paracanthop percopsiforms (para-
244 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
? Grammatidae
? Centrogeniidae
? Opistognathidae, Lonchopisthus
? Gerreidae
? Pseudochromidae
^
? Embiotocidae, Amphisticus
? Embiotocidae, Embiotoca
? Cichlidae, PtychochromisCPomacentridae, DischistodusPomacentridae, AmphiprionPholidichthyidaeLabridae, ChoerodonLabridae. AchoerodusLabridae, Con's
? PristolepidaeCNandidaeBadidaeCChannidaeAnabantidae, SandeliaMugilidae, Agonostomus
? Bedotiidae
? AtherinidaeK Adrianichthyidae, XenopoecilusCyprinodontidaeApiocheilidaeHemiramphidaeExocoetidaeScomberesocidaeBelonidae, StrongyluraBelonidae, TylosurusAcropomatidaePercichthyidaePempheridaeGlaucosomatidaeAmmodytidaeTerapontidae. TeraponTerapontidae, LeiopotheraponGirellidaeKuhliidaeEpigonidaeLactariidaeBathyclupeidaePlatycephalidae, PlatycephalusLutjanidae
? Centrarchidae, MicropterusMoronidae
? Apogonidae, GlossamiaKurtidaeCirrhitidae, ParacirrhitesCepolidae, CepolaSerranidaeNematistiidae
? MalacanthidaePriacanthidaeLateolabracidaeBramidae
? AmarsipidaeMullidae
? Sciaenidae, CynoscionChampsodontidae
? Uranoscopidae, XenocephalusPseudaphritidae
? Cheimarrichthyidae
? TrachinidaePlesiopidae, AssessorHexagrammidaePolynemidaeCaproidae, Antigonia
- Caproidae, Capros
- CentriscidaeTriacanthodidae
- Menidae
- LuvaridaeZanclidaeAcanthuridae
? CetomimidaeRhamphocottidae
- Chaunacidae
- Scombridae, ScomberSphyraenidae
- Scombrolabracidae
- Rachycentridae
- Psettodidae
-c
-c
?c
?
c
-E
m^
^
HZ
^
-C
LeptobramidaeCoryphaenidaeLeiognathidaeEphippidae, ChaetodipterusLampridaeVeliferidae, VeliferAnomalopidae, AnomalopsAnomalopidae, PhotoblepharonCarangidae, SelarHaemulidaeInermiidaeBerycidaeTrachichthyidaeMelamphaidaeAphredoderidaePercopsidaeAmblyopsidaeOstracoberycidaeCaristiidaeLatidaeCentropomidaeSymphysanodontidaePomatomidaeSebastidaeScorpaenidaeSillaginidaeRhyacichthyidaePercidae, PercaPercidae, PercinaToxotidaeCentrolophidaeAmbassidae, AmbassisAmbassidae, TetracentrumParazenidaeZeniontidaeGrammicolepidaeOreosomatidaeNemipteridaeCallanthiidaeLethrinidaeCentracanthidaeSparidaePolycentridae, AfronandusPolycentridae, MonocirrhusPolycentridae, PolycentropsisPolycentridae, PolycentrusBatrachoididaeIcosteidaeBarbourisiidaeStephanoberycidaeGibberichthyidaeRondeletiidaePolymixiidaeHolocentridaeRanicipitidaeOphidiidae, BrotulaOphidiidae, DicroleneBythitidae, CalamopteryxBathymasteridaeZaproridaeAnoplopomatidaeBovichtidaeStichaeidaeDactylopteridaeDraconettidaeGobiesocidaeTripterygiidae, RuanohoTripterygiidae, Lepidoblennius
? Blenniidaei? Dactyloscopidae
? Chaenopsidae
? Labrisomidae, Calliclinus
? Clinidae, Clininae
? Clinidae, MyxodinaeCHypoptychidae, AulichthysHypoptychidae, Hypoptychus
? ElassomatidaeAulorhynchidae
Le
SynbranchidaeMastacembelidae(? Gasterosteidae, Apeltes
? Gasterosteidae, Culea
? Gasterosteidae, Gasterosteus
? Gasterosteidae, Pungitius
? Gasterosteidae, Spinachia
NUMBER 1 I 245
canthop gadiforms + zeiforms) + (mix of groups, ofwhich most deeply nested are some paracanthops)).Miya et al. (2003:fig. 2), in a molecular study, hy-pothesized a somewhat similar arrangement, with Po-lymixiidae as sister group of a group comprising per-copsiforms. gadiforms and zeiforms. Chen et al.(2003, fig. 6C), in another molecular study, summa-rized their series of three molecular studies (seeSmegmamorpha, below) and included Polymixiifor-mes in the Paracanthopterygii.The evidence presently is mixed as to whether Po-lymixiidae are the sister group of all other acanthop-terygians (except lampridiforms) or should be in-cluded among a group comprising some of the par-acanthopterygians (which could be the sister groupof all other acanthomorphs).Paracanthopterygii: Percopsiformes, Lophiifor-mes (chaunacidae), ophidiiformes. gadiformes(ranicipitidae), Batrachodiformes (batrachoidi-dae). The MRT retrieved the Paracanthopterygii aspolyphyletic, its five components comprising parts offour separate monophyletic groups.
1 ) [B 37] Percopsiformes, Chaunacidae. Thefreshwater percopsiforms were retrieved as a re-solved monophyletic clade [B 38] (Aphredoderidae(Percopsidae + Amblyopsidae)) that is the unlikely
sister group of the moderately deep-dwelling, marineChaunacidae. S&J hypothesized monophyly of per-copsiforms based on eight characters (for discussionof the recent vacillating history of the group seeS&J"s account of Percopsiformes).While the resolution of the Percopsiformes we re-port will probably undergo rearrangement, we feelconfident that the monophyly of the three familieswill be further corroborated. Molecular studies (Wi-ley et al. 2000; Miya et al. 2003) have included onlytwo percopsiforms, aphredoderids and percopsids, intheir analyses and these indicate that the two familiesform a sister group; however, it is the phylogeneticposition of Amblyopsidae that has been the source ofmost disagreement.2) Ophidiiformes [D 140] are monophyletic, butOphidiidae are paraphyletic without inclusion of By-thitidae. J. Nielsen (pers. comm.) informs us thatthere is increasing support for combining these twofamilies.3) [D 139] Ranicipitidae (sister group of all othergadiforms; Yabe 1985) seems unlikely to be closelyrelated to the beryciform Holocentridae with whichit is paired in [D 139], but close relationship of Ran-
icipitidae with Ophidiiformes, as indicated by node[D 137] is probable.4) [C 63] Batrachoididae are well nested withinthe perciform family Polycentridae [C 60]. We be-lieve a close relationship between these two groupsto be improbable.Markle (1989) hypothesized the batrachoidids assister group of the gadiforms, of which Ranicipitidaeare the basal member. Patterson and Rosen (1989)hypothesized batrachoidids and lophiiforms as sistergroup to the gadiforms, and Wiley et al. (2000:fig.6), found batrachoidids to be the sister group of by-thitids. Miya et al. (2003:fig. 2) found gadiforms tobe the sister group of zeiforms. Thus, the interrela-tionships of all these groups remains problematic, butin all studies, except ours, the evidence indicates thatbatrachoidiforms are related to a pre-perciformgroup.Stephanoberyciformes. The MRT retrieved thefive families in two separate and well removedclades. Four of the families were retrieved in a re-solved clade [D 134], the fifth family. Cetomimidae,was retrieved as part of a clade [B 36] containing avariety of taxa. Paxton et al. (2001) suggested thatRondeletiidae and Gibberichthyidae are sister taxa.but the two are separated by Stephanoberycidae inthe MRT.Zeiformes [C 56]. In the MRT zeiforms comprisea monophyletic group that is the sister group of theRhamphocottidae. We know of no other evidence fora zeiform-rhamphocottid relationship.Although we examined few zeiform taxa, theirstepwise arrangement in the MRT partly differs fromtheir relative positions in the recent classification hy-pothesized by Tyler et al. (2003). Of the taxa in theirclassification, Oreosomatidae are the least specializedzeiform family that we examined. The MRT, how-ever, retrieved Parazenidae and Zeniontidae as thefirst two branches of the clade. The position of Par-azenidae and Zeniontidae in our analysis agrees withJ&P's (1993:596) assertion that these families are themost plesiomorphic zeiforms. The MRT positions arealso supported by Miya et al.'s (2003) molecularstudy based on complete mitochondrial DNA se-quences, in which they reported essentially the samearrangement as in our MRT: (Parazenidae (Zenion-tidae (Oreosomatidae + Zeidae))).Holocentridae [D 139]. The position of the Hol-ocentridae, whether its affiliations are with the be-ryciforms or the percomorphs. has been the subject
Fig. 5. Strict consensus of 1.823 equally parsimonious trees (EPT) generated from a restricted parsimony analysis of 147 families ofacanthomorph fishes (see Appendix for analysis specifics). Each EPT has 595 steps, consistency index 0.14. homoplasy index 0.86, andre-scaled consistency index 0.09. Star character indicates continuation of tree which is split into two columns. Tree is mid-point rooted.
246 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
-Gj
GrammatidaeOpistognathidae, LonchopisthusGerreidaePseudochromidaeEmbiotocidae, EmbiotocaEmbiotocidae, AmphisticusCichlidae, PtychochromisPomacentridae, DischistodusPomacentridae, AmphiprionPholidichthyidaeLabridae, ChoerodonLabridae, AchoerodusLabridae, CorisNandidaeBadidaePristolepidaeChannidaeAnabantidae, SandeliaMugilidae, AgonostomusBedotiidaeAtherinidaeAdrianichthyidae, XenopoecilusCyprinodontidaeApiocheilidaeHemiramphidaeExocoetidaeScomberesocidaeBelonidae, StrongyluraBelonidae, TylosurusCentrogeniidaeBramidaePlatycephalidae, PlatycephalusPriacanthidaeLuvaridaeChampsodontidaePseudaphritidaePsettodidaeCetomimidaeChaunacidaeAphredoderidaePercopsidaeAmblyopsidaeLampridaeVeliferidae, VeliferZanclidaeCaproidae, CaprosAcanthuridaeLeiognathidaeEphippidae, ChaetodipterusCepolidae, CepolaTrachinidaeCheimarrichthyidaeHexagrammidaeSillaginidaeRhyacichthyidaePercidae, PercaPercidae, PercinaKurtidaeUranoscopidae, XenocephalusRhamphocottidaeParazenidaeZeniontidaeGrammicolepidaeOreosomatidaePolycentridae. PolycentrusPolycentridae, AfronandusPolycentridae, MonocirrhusPolycentridae, PolycenlropsisBatrachoididaePlesiopidae, AssessorBathymasteridaeZaproridaeAnoplopomatidaeBovichtidaeStichaeidaeDactylopteridaeDraconettidaeGobiesocidaeTripterygiidae, RuanohoTripterygiidae, LepidoblenniusBlenniidaeDactyloscopidaeClinidae, ClininaeClinidae, MyxodinaeChaenopsidaeLabrisomidae, CalliclinusHypoptychidae, AulichthysHypoptychidae, HypoptychusElassomatidaeAulorhynchidaeSynbranchidaeMastacembelidaeGasterosteidae, CuleaGasterosteidae, GasterosteusGasterosteidae, ApeltesGasterosteidae, PungiliusGasterosteidae, Spinachia
Terapontidae, TeraponLactariidaeApogonidae, GlossamiaCirrhitidae, ParacirrhitesMalacanthidaeSerranidaeScombrolabracidaeBathyclupeidaeSciaenidae, CynoscionSebastidaeScorpaenidaeSymphysanodontidaePomatomidaeAcropomatidaePercichthyidaeTerapontidae, LeiopotheraponLeptobramidaeCoryphaenidaeGlaucosomatidaeAmmodytidaeNematistiidaeOstracoberycidaeCaristiidaeLatidaeCentropomidaePempheridaeEpigonidaeKuhliidaeMoronidaeLutjanidaeToxotidaeCentrolophidaeAmbassidae, AmbassisAmbassidae, TetracentrumGirellidaeCentrarchidae. MicropterusLateolabracidaeRachycentridaeCarangidae, SelarHaemulidaeInermiidaeNemipteridaeCentracanthidaeSparidaeCallanthiidaeLethrinidaeAmarsipidaeCaproidae, AntigoniaTriacanthodidaeMullidaeScombridae, ScomberPolynemidaeSphyraenidaeMelamphaidaeBerycidaeTrachichthyidaeAnomalopidae, AnomalopsAnomalopidae, PhotoblepharonCentriscidaeMenidaeIcosteidaeBarbourisiidaeRondeletiidaeStephanoberycidaeGibberichthyidaePolymixiidaeHolocentridaeRanicipitidaeOphidiidae. BrotulaOphidiidae. DicroleneBythitidae, Calamopteryx
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of much current debate. Baldwin and Johnson (1995)discussed the evidence, which they found inconclu-sive. The MRT retrieved the Holocentridae as sistergroup of the gadiform Ranicipitidae among a largergroup of clades [D 133] comprising ophidiiforms.stephanoberyciforms, and Polymixia. Holocentridspossess ER. which is restricted to pre-perciforms, andthe putative percomorph families Centriscidae, Men-idae, and Icosteidae, which we be believe are pre-perciforms and which our MRC also retrieved amonga clade [D 123] that includes many of those fishes.Smegmamorpha. The MRT retrieved the taxa J&P(1993) included in the Smegmamorpha (Synbranchi-formes, Elassomatidae, Gasterosteiformes, Mugili-dae, Atherinomorpha) as a polyphyletic assemblageof three clades, none of which appears as closely re-lated to another: Clade [A 18], Mugilidae + Atheri-nomorpha; Clade [C 81], Gasterosteiformes (lessCentriscidae), Elassomatidae, and Synbranchiformes;Clade [D 130], Centriscidae ( + Menidae). Clearly,J&P's putative smegmamorph synapomorphy (char-acter 53) was swamped by other characters.Clade [A 18] appears as the final branch in a clade[A 13] that includes the Anabantomorpha as its basalmembers. Clade [C 81] appears as the seemingly un-likely sister group of a clade [C 70] comprising theDactylopteridae, Gobiesociformes (Draconettidae,Gobeisocidae), and Blennioidei. Clade [D 130] wasretrieved as the sister group of Menidae among aclade [D 125] of pre-perciforms that also includes theIcosteidae.The monophyly of the Smegmamorpha was firstrejected by Nelson (1994:252-253). who placed Mu-gilomorpha at the base of the Acanthopterygii, fol-lowed by a branch, Atherinomorpha. which was sep-arated by several branches from a polyphyletic Gas-terosteiformes + Synbranchiformes. Wiley et al.(2000:figs. 6 and 8), in a combined morphologicaland molecular cladistic study, failed to retrieve amonophyletic Smegmamorpha. Neither Chen et al.(2003), in their three molecular analyses (see follow-ing), nor Miya et al. (2003), using mitogenomic data,recovered a monophyletic Smegmamorpha. Otherthan J&P's (1993) minimally supported hypothesis,no one has shown support for a monophyletic Smeg-mamorpha. Considering our results together withthose of the other studies mentioned, there is no basisfor recognizing the taxon. Older names are availablefor each of its originally hypothesized componentsand we suggest that Smegmamorpha be permanently
rejected for nomenclatural purposes. Discussion of itspurported three clades follows.Clade [A 18]. A sister-group relationship of theMugilidae (= Mugilomorpha) and Atherinomorphahas been suggested by many authors over the years,but the most recent morphological studies (Stiassny1993; Parenti, in press) consider the relationshipproblematic. Molecular studies are also problematic,but show a tendency of support of the relationship.Miya et al. (2003), in a mitogenomic study including100 species representing all but one (Batrachoidifor-mes) of the higher teleosotean orders, retrieved aclade (((Mugilidae) + (Blenniidae + Gobiesocidae))+ (Atherinomorpha)), which hints strongly at a mu-gilomorph-atherinomorph relationship. Chen et al.(2003), in a study of acanthomorphs, reported onthree analyses: (1) 12S and 16S mtDNA from 97taxa; (2) 28S rDNA from 74 taxa; (3) nuclear pro-tein-coding gene, rhodopsin from 86 taxa. In (1), ath-erinomorphs were polyphyletic. but one of the cladeswith atherinomorphs has ((Mugilidae + Atherino-morpha) + (Blennioidei + Gobiesocidae)). In (2),atherinomorphs are also polyphyletic, but one cladehas ((Mugilidae + Atherinidae) + (Serranidae)). In(3), atherinomorphs are monophyletic, and the cladeis (Mugilidae + Atherinomorpha). In a combinedanalysis of the three analyses, atherinomorphs arepolyphyletic, but the clade containing all atherino-morphs and the mugilid is ((Poecilliidae + Mugili-dae) + (Bedotiidae + Belonidae)).Including our study, there appears to be growingsupport for a monophyletic Mugilomorpha-Afheri-nomorpha sister group. To formally acknowledge thissister-group, and simplify future discussion of it, weresurrect Cope's (1871:480) Percesoces. in which heincluded only Mugilidae and Atherinidae, as its or-dinal-group name.The branchings of the Atherinomorpha in Clade[A 18] have been the subject of many studies sinceRosen ( 1 964) first proposed the group. Parenti (Inpress) presents a considered review of the relevantpost- 1964 literature and summarizes current hypoth-eses of atherinomorph inter- and intra-relationships.The intra-relationships of the atherinomorphs as re-trieved by our MRT generally reflect those proposedin the literature, with the major difference being thatthe positions of Adrianichthyidae and Aplocheilidaein our tree are exchanged.Clade [C 81] supports J&P's (1993:578-579) hy-pothesis that an Aulorhynchidae including Aulorhyn-
Fig. 6. 50% majority rule consensus of 1823 equally parsimonious trees (EPT) generated from a restricted parsimony analysis of 147families of acanthomorph (see Appendix for analysis specifics). EPT statistics are as in Fig. 5. Four major clades are lettered A, B, C,and D. All other internal nodes are numbered in boxes. Character support for all nodes and frequency values for internal nodes are foundin Table 13. Star character indicates continuation of tree, which is split into two columns. Tree is mid-pointed rooted.
248 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 13.?Support values for all nodes of majority rule consensus tree (MRT; Fig. 6). Taxonomic names indicate terminal nodes. A,B, C, D refer to four major internal clades. Numbered nodes refer to clades within A, B, C, D. Bold and underlined characters areuncontradicted synapomorphies. All other characters are homoplastic. Character state follows colon and each character is separated by acomma. Frequency (F) corresponds to nodes represented in at least 50% of all 1823 equally parsimonious trees. Symbols: - = no supportor support ambiguous; n/a = not applicable.
Node Supporting Characters F Node Supporting Characters FA 42:1 100 Cheimarrichthyidae 12:2 n/aB 15:1 96 Cichlidae, Ptychochromis 12:1,51:1 n/aC 45:1 96 Cirrhitidae, Paracirrhites 14:1,36:2,43:1 n/aD 6:0 79 Clinidae, Clininae 26:4 n/aAcanthuridae 14:1,29:1 n/a Clinidae, Myxodinae ? n/aAcropomatidae ? n/a Coryphaenidae 4:1,16:1,45:1,52:1 n/aAdrianichthyidae, Xenopoecilus 15:0,44:1 n/a Cyprinodontidae 6:2,32:0,35:2 n/aAmarsipidae 14:1 n/a Dactylopteridae 12:0 n/aAmbassidae, Ambassis 17:2 n/a Dactyloscopidae 3:1,36:1 n/aAmbassidae, Telracenlrum 26:1,42:1 n/a Draconettidae 16:1, 19:1 n/aAmblyopsidae 1:1,37:3 n/a Elassomatidae 44:1 n/aAmmodytidae ? n/a Embiotocidae, Amphisticus 18:0 n/aAnabantidae, Sandelia 10:2, 15:0,31:0 n/a Embiotocidae, Embiotoca 49:0 n/aAnomalopidae, Anomalops 7:1 n/a Ephippidae, Chaelodipterus ? n/aAnomalopidae, Photoblepharon 15:1 n/a Epigonidae ? n/aAnoplopomatidae ? n/a Exocoetidae ? n/aAphredoderidae 18:1 n/a Gasterosteidae, Apelles 7:0 n/aAplocheilidae 15:0 n/a Gasterosteidae, Culea ? n/aApogonidae, Glossamia 42:1 n/a Gasterosteidae, Gasterosteus ? n/aAtherinidae 26:3 n/a Gasterosteidae, Pungitius ? n/aAulorhynchidae 21:1,26:7 n/a Gasterosteidae, Spinachia ? n/aBadidae 26:7,37:1 n/a Gerreidae 35:0,45:1 n/aBarbourisiidae 26:1,43:1 n/a Gibberichthyidae ? n/aBathyclupeidae ? n/a Girellidae 4:1,43:1 n/aBathymasteridae ? n/a Glaucosomatidae ? n/aBatrachoididae 4:3, 16:1,39:1 n/a Gobiesocidae 3:1,15:1,52:1 n/aBedotiidae 26:2 n/a Gratnmatidae 7:1,17:1 n/aBelonidae, Strongylura 22:0 n/a Grammicolepidae 3:0,5:1,45:0,52:1 n/aBelonidae, Tylosurus 9:0 n/a Haemulidae 6:3 n/aBerycidae 49:0 n/a Hemiramphidae 1:0,4:1,40:0,47:1 n/aBlenniidae ? n/a Hexagrammidae ? n/aBovichtidae 7:0, 14:1, 17:1 n/a Holocentridae 14:1 n/aBramidae 3:1,6:3 n/a Hypoptychidae, Aulichthys 11:1 n/aBythitidae, Calamopteryx 17:2,37:3 n/a Hypoptychidae, Hypoptychus 4:3,6:0,17:0,26:6,32:2,34:1 n/aCallanthiidae . 12:2, 19:0,37:1 n/a Icosteidae 6:1, 15:1, 16:1 n/aCaproidae, Anligonia 12:2,15:1 n/a Inermiidae ? n/aCaproidae, Capros 4:3,11:1,17:2,30:1,31:1 n/a Kuhliidae 6:3,43:1 n/aCarangidae, Selar 12:0, 15:1 n/a Kurtidae 16:1 n/aCaristiidae 26:8,50:1 n/a Labridae, Achoerodus ? n/aCentracanthidae 4:1,40:1 n/a Labridae, Choerodon ? n/aCentrarchidae, Micropterus ? n/a Labridae, Coris ? n/aCentriscidae 3:1, 11:0,20:1,29:1,32:2, Labrisomidae, Calliclinus 12:0, 18:1,26:1 n/a37:1,40:1,50:1,53:1 n/a Lactariidae 7:1 n/aCentrogeniidae 26:1,36:2,38:1,44:1,49:0 n/a Lampridae 20:1,21:1,40:1 n/aCentrolophidae 12:0 n/a Lateolabracidae 6:3 n/aCentropomidae 14:1,29:1 n/a Latidae 45:0 n/aCepolidae, Cepola 49:0 n/a Leiognathidae 9:1,26:5,35:1 n/aCetomimidae !i2, 14:0,38:1,41:1,44:1 n/a Leptobramidae 12:1,15:1 n/aChaenopsidae 6:2,8:1, 18:1,37:2 n/a Lethrinidae 4:1,5:1 n/aChampsodontidae ? n/a Lutjanidae ? n/aChannidae 9:0, 26:3 n/a Luvaridae 4:2,23:1,26:1 n/aChaunacidae 4:3,6:2, 17:2,39:1 n/a Malacanthidae 19:1 n/a
NUMBER I 1 249
Table 13.?Continued.
Node Supporting Characters F Node Supporting Characters FMastacembelidae 15:0, 17:0,36:2 n/a Terapontidae, Terapon ? n/aMelamphaidae 18:1 n/a Terapontidae, Leiopolherapon j 26:8 n/aMenidae 4:2, 18:1,23:1,48:1,49:0 n/a Toxotidae 26:1,42:1 n/aMoronidae 27:1 n/a Trachichthyidae 4:3,39:1 n/aMugilidae, Agonostomus 9:0, 15:0,43:0 n/a Trachinidae ? n/aMullidae 12:1,32:2,37:2 n/a Triacanthodidae 9:2,21:0,29:1 n/aNandidae 9:0, 14:0 n/a Tripterygiidae, Lepidobtennius 26:3,34:1 n/aNematistiidae 43:1 n/a Tripterygiidae, Ruanoho ? n/aNemipteridae 16:1,36:2,40:1,50:1 n/a Uranoscopidae, Xenocephalus 11:1, 12:2,37:1,38:1,45:0,Ophidiidae, Brotula 16:1,21:1,40:1 n/a 52:1 n/aOphidiidae, Dicrolene 7:0, 50:0 n/a Veliferidae, Velifer 4:3, 16:0, 17:1,43:1,48:1,49:0 n/aOpistognathidae, Lonchopisthus 38:1 n/a Zanclidae 7:1,20:1 n/aOreosomatidae 7:0, 11:1,33:1 n/a Zaproridae 23:1 n/aOstracoberycidae ? n/a Zeniontidae ? n/aParazenidae 6:2, 10:2 n/a 1 14:0,15:1 100Pempheridae ? n/a 2 30:1,31:1 100Percichthyidae 26:1 n/a 3 32:2 100Percidae, Perca 31:0 n/a 4 14:1, 17:2, 18:1 100Percidae, Percina 27:1,30:1,37:3 n/a 5 9i3, 26:5, 38:1, 50:1, 51:2 100Percopsidae 16:0 n/a 6 45:1 99Pholidichthyidae 4j4, 9:1,32:0,44:1 n/a 7 12:0,13:1,19:1 100Platycephalidae, Platycephalus 20:1,32:1 n/a 8 38:0, 50:0 99Plesiopidae, Assessor 11:1,42:1 n/a 9 9:2, 15:0,21:1,32:0 100Polycentridae, Afronandus ? n/a 10 6:2, 11:1,39:1 100Polycentridae, Monocirrhus 21:1 n/a 11 37:1,41:1 100Polycentridae, Polycentropsis ? n/a 12 46:1,48:1 100Polycentridae, Polycentnis 45:0 n/a 13 35:0,40:1,55:1 100Polymixiidae 5:1,45:0 n/a 14 3:1,32:0 100Polynemidae 17:1 n/a 15 30:0,43:1 99Pomacentridae, Amphiprion 45:0 n/a 16 11:1,45:1 99Pomacentridae, Dischistodus ? n/a 17 6:2,19:1,44:1,54:1 100Pomatomidae 12:1,26:1 n/a 18 1A, 40:0, 53:1, 55:0 100Priacanthidae 18:1,52:1 n/a 19 4:1, 10:2, 13:1 100Pristolepidae 16:1,31:0,42:0 n/a 20 22:1 100Psettodidae 17:2,50:0 n/a 21 1:1,37:2,40:1 100Pseudaphritidae 15:0,17:1,37:3 n/a 22 3:1 98Pseudochromidae 7:1, 13:1,37:1,2 n/a 23 4:0,35:1 98Rachycentridae 4:1 n/a 24 32:0,51:2 100Ranicipitidae 3:1,6:2, 12:2, 17:2,37:1,39:1 n/a 25 26:2,28:1 100Rhamphocottidae 32:1 n/a 26 18:0,27:1,44:1,50:1 100Rhyacichthyidae 1:1, 12:2, 14:1,29:1,37:1,44:1 n/a 27 32:3,41:1 100Rondeletiidae ? n/a 28 ? 100Sciaenidae, Cynoscion 7:1, 12:2,36:2,45:1 n/a 29 9:0,35:0 100Scomberesocidae 1:0,15:0,40:0,42:0 n/a 30 ? 86Scombridae, Scomber 4:3,49:0 n/a 31 6:0 91Scombrolabracidae ? n/a 32 45:1 97Scorpaenidae ? n/a 33 16:1 97Sebastidae 49:0 n/a 34 50:1 96Serranidae ? n/a 35 3:1, 11:1, 12:0 100Sillaginidae 6:3,21:1,27:1,47:1 n/a 36 52:1 98Sparidae ? n/a 37 27:1 97Sphyraenidae 7:1 n/a 38 10:2,20:1,24:1,45:0 100Stephanoberycidae 21:1,39:1,41:1,44:1 n/a 39 22:1 100Stichaeidae ? n/a 40 4:2, 14:0 99Symphysanodontidae ? n/a 41 10:1, 45:0 100Synbranchidae 37:2,38:1,42:1,44:1,50:0 n/a 42 1:1,23:1 99
250 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 13.?Continued.
Node Supporting Characters F Node Supporting Characters F43 26:6 98 92 11:1 10044 40:1 97 93 15:1 10045 4:0,42:1 100 94 ? 9446 14:0 81 95 49:0 7847 ? 68 96 21:1 9648 37:1 100 97 36:1 10049 31:1 100 98 45:1 9950 17:1,43:1 100 99 26:6 10051 20:1 100 100 49:1 10052 17:2 97 101 7:1 10053 ? 75 102 12:1 9354 50:1 90 103 17:1 9455 21:1,39:1 100 104 49:1 9556 7:1,18:1,40:1,43:1 100 105 9:1 10057 16:1 100 106 18:1 10058 1:1,23:1,32:1 100 107 6:1,45:1 10059 7:1 58 108 17:1 7760 15:1,18:1 100 109 21:1 9261 12:2 83 110 12:2 9662 ? 81 111 49:0 8863 7:0 81 112 26:7,42:1 10064 ? 80 113 18:1,52:1 10065 37:3,50:1 100 114 19:1 10066 17:0 100 115 30:1,31:1 10067 30:1,31:1 100 116 36a:l,42:l 10068 12:2 100 117 ? 9869 21:1 100 118 45:1 9670 17:2 100 119 17:2 9671 6:2,37:1 100 120 5:1,23:1 9972 7:0,39:1,40:1,44:1 100 121 ? 8973 11:1, 14:1,36:1,37:2,38:2, 122 11:1,43:1 9941:1 100 123 21:0 9874 1:1,35:1 100 124 ? 9975 9:1,36:2,43:1 100 125 25:1,33:1 9876 11:0 100 126 50:1 10077 30:0,31:0 100 127 ? 9678 ? 91 128 ? 9179 ? 89 129 26:6,39:1 10080 43:0 92 130 12:2,21:1 9981 1:1,53:1 91 131 43:0 9682 7:0, 14:1,20:1,39:1 100 132 17:0,32:1 10083 15:1 100 133 50:1 10084 3:1, 11:1 100 134 14:1 10085 ? 98 135 38:1 10086 4:3,40:1,52:1 100 136 26:6 9987 32:2,34:1,39:1 100 137 7:1 10088 ? 97 138 52:1 10089 ? 99 139 11:0, 15:1,43:1 10090 52:1 100 140 4:3,27:1 10091 6:3 100 141 10:2 99
chits and Aulichthys is polyphyletic, and that Hypop-tychus and Aulichthys form a sister group, Hypop-tychidae. In retrieving Elassomatidae as sister groupto a series of branches including Aulorhynchidae,Synbranchiformes, and Gasterosteidae, the arrange-ment in the MRT is congruent with Johnson and
Springer's (1997:176, abstract) first suggestion of aclose relationship between Elassomatidae and Gas-terosteidae. The structure of the clade, and its re-trieved sister group, are worthy of more study. Re-trieval of Synbranchidae and Mastacembelidae as amonophyletic group and their inclusion as closely re-
NUMBER 1 1 251
lated to gasterosteiforms in Clade 2 variously sup-ports the findings of J&P and other authors.Clade [D 130]. Centriscidae (includes Macroram-phosidae) have usually been treated as members ofthe Gasterosteiformes, among which they are consid-ered most closely related to the Aulostomidae andFistulariidae (Nelson 1994:302), which were nottreated by S&J. Among other characters, centriscids,aulostomids, and fistulariids share in having the an-terior vertebrae elongate. Chen et al. (2003) foundmolecular support for a close relationship of the threefamilies.We examined the gill-arch musculature and skel-eton of two specimens of Aulostomus chinensis (Lin-naeus), USNM 111979 (147 mm) and 327565 (221mm) and find that it is very reduced and generallyuninformative. Among other things, Eb4 is absent(also absent in Fistularia, J&P:table 1 ) and the pos-terior musculature is incompletely differentiated fromSO. A small muscle, probably Ad5, attaches to thedistal ends of Cb4 and Cb5; there is no indication ofOP or ER, which does not occur in the absence ofOP. A short, IAC (also present in Fistularia, J&P:table 1) attaches to Ebl uncinate process, but doesnot join the well-developed Pb2, which lacks artic-ulating processes. Nelson (1969a:fig. 17A) illustratedthe gill-arch skeleton of A. chinensis.S&J treated Centriscidae, Menidae, and Icosteidaeas more probably related to pre-percomorph groups,because the three families possess ER and have OPattaching to or at the level of Cb4. Retrieval of thethree families among a group of pre-perciforms byour MRT supports S&J's opinion. If the centriscids,aulostomids, and fistulariids are closely related, andtheir linkage with the other gasterosteiforms is cor-roborated, there are two main possibilities: the pre-perciform characters of centriscids are reversals (ornew acquisitions), or centriscids represent the basalgasterosteiform branch, in which case, the gasteros-teiforms are probably pre-perciforms.Anabantomorpha [A 13 to 17]. Britz (2003:427)proposed, but incompletely resolved the interrelation-ships of these fishes. The MRT retrieved the anaban-tomorphs as the basal series of steps leading to thePercesoces, hence the anabantomorphs are indicatedas polyphyletic without inclusion of Percesoces.Character 55 (parasphenoid teeth) was inadequate todefine the anabantomorphs, but character 54 (Eblsmodified as suprabranchial organs) did define the An-abantoidei [A 17]. We believe the anabantomorphs(and anabantoids) are monophyletic, and that the in-trarelationships of the families in our analysis areprobably ((Nandidae + Badidae) + (Pristolepidae(Channidae + Anabantidae))). We have no sugges-tion as to their nearest relatives, but do not excludethe Percesoces as a possiblility.Apparently, ours is the first cladistic analysis to
indicate a close relationship between anabantomorphsand Percesoces. Without suggesting a direct relation-ship, Gosline (1968), in a survey of perciform fishes,uniquely segregated these two groups (as Anabanto-idei, Mugiloidei) as the only "Protopercoid" Sub-orders (our emphasis) of his "Percoidei and Deriv-ative Suborders." In part following Regan (1912),Gosline's Mugiloidei included the Polynemidae, Mu-gilidae, Sphyraenidae, Atherinidae and Phallostethi-dae, but excluded the "cyprinodontoids" (= our Cy-prinodontiformes) and "exocoetoids" (= our Belon-iformes), which he placed among the perciforms, andhis Anabantoidei excluded Pristolepis and Nandus,which he also placed among the perciforms.Gosline's main character for associating his Mu-giloidei and Anabantoidei, was a plesiomorphic con-dition, separation of pelvices from cleithra. He rec-ognized that there were problems with the characterand that reversals to the plesiomorphic state hadprobably occurred. Although Gosline did not resolvethe classification so neatly as does our MRT, we be-lieve there was more than luck involved in his per-ception.Labroidei, Pholidichthyidae [A 5]. The MRT im-plies that the Labroidei are paraphyletic without in-clusion of Pholidichthyidae, thus essentially corrob-orating Stiassny and Jensen (1987), who most re-cently hypothesized a monophyletic Labroidei (Em-biotocidae, Pomacentridae, Cichlidae, Labroidea)based on gill-arch morphology. They lacked materialof Pholidichthys, but noted (their p. 294) several os-teological similarities shared by that genus and la-broids, and indicated that a study of Pholidichthysmusculature would be important, particularly as towhether it included a "sling" (a joining of LE4 andOP). It does not?it lacks LE4, but even if S&J areincorrect in identifying the single levator on Eb4 asLP, rather than LE4, there is no muscular continua-tion of the levator with OP.Johnson (1993:8-10) noted problems with some ofStiassny and Jensen's characters and hoped that "thehypothesis of a monophyletic Labroidei does not be-come dogma" in the "absence of corroborative evi-dence independent of the pharyngeal apparatus." Hebelieved that "labroid monophyly will be an impor-tant hypothesis to test with molecular data."Using single-copy nuclear DNA, Streelman andKarl (1997), who lacked material of Pholidichthys,tested the monophyly of Stiassny and Jensen's La-broidei. They used Sebastes (Sebastidae) as outgroupin their analysis and included a cottid, percid, po-macanthid, and acanthurid together with three ormore taxa in each of the labroid families. Their re-sults (their fig. 2) indicate a polyphyletic Labroidei:((Embiotocidae + Pomacentridae) + (Cottidae (Per-cidae (Pomacanthidae (Acanthuridae (Labridae 4-Cichlidae)))))).
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Given that the monophyly of the Labroidei, in-cluding Pholidichthys, still rests only on morphologyof gill-arch muscles and skeleton, our addition ofmore characters from the same anatomical complexmay only be adding correlated characters. Polyphylyof the group has been hypothesized by only one mo-lecular study, which included relatively few non-la-broids and did not include Pholidichthys. Molecularstudies involving labroids, Pholidichthys, and non-labroids additional to those used by Streelman andKarl (1997) are still needed, as are morphologicalstudies based on more than gill-arch anatomy.POLYCENTRIDAE [C 60]. Both the MRT and SCTrecovered Polycentridae and Batrachoididae as amonophyletic clade. As discussed under Batrachoid-iformes, we doubt a close relationship between thetwo groups. S&J provide several non-gill arch spe-cializations in support of polycentrid monophyly,which we believe will continue to be corroborated byfuture studies.NOTOTHENOIDEI (BOVICHTIDAE, PSEUDAPHRITIDAE)[C 69 and B 34]. The MRT retrieved the essentiallysouthern hemispheric restricted notothenioids aspolyphyletic. Pseudaphritidae were retrieved as abranch of a polytomy including an apparently het-erogeneous assemblage of perciform and pre-perco-morph taxa. Bovichtidae were retrieved as the sistergroup of the zoarcoid family Stichaeidae. The sistergroup is the fifth step in a multi-stepwise clade [C64] whose first four steps are Plesiopidae (perciform),Bathymasteridae (zoarcoid), Zaproridae (zoarcoid),Anoplopomatidae (traditionally allied with scorpae-noids, but Quast, 1965, questioned the relationship,and the matter has not been resolved).The monophyly of the notothenioids has long beenaccepted (Balushkin 1992:90-91, references the mainpublications). Balushkin (1992, 2000), based on mor-phology, hypothesized the Pseudaphritidae and Bov-ichtidae as the first and second branches of the no-tothenioid clade. Near et al. (2004), using completegene sequences of mtDNA and 16S rRNA, foundBovichtidae and Pseudaphritidae as the first and sec-ond branches.The position of Bovichtidae as closely related tozoarcoids appears to offer the first cladistic supportfor this relationship, which has been generally sug-gested (Anderson 1984:581, 2003:309; Nelson 1994:388). Molecular studies (Chen et al. 2003), however,have not found a close relationship between the twogroups.Dactylopteridae, Callionymoidei, Gobiesocoid-
ei, Blennioidei. The MRT retrieved this group as acompletely resolved monophyletic clade [C 71].Some of the implied relationships are new, but webelieve that the group is worth serious consideration.For this reason we assign a new ordinal-group name,Benthomorpha, to it. Perhaps, the most questionable
member is the Dactylopteridae, which Imamura(2000) hypothesized as including Malacanthidae. OurMRT retrieved the latter family as part of a poly-chotomous clade [D?see polytomy] well removedfrom the Benthomorpha. One of the main characters,a ring-like TPb2, that Imamura used to relate the twofamilies is present in a wide variety of acantho-morphs (e.g., Chaunacidae, Percidae, Kuhliidae,Bathymasteridae). We did not use this character statein our analysis because the many different shapeswith intermediates that TPb2 assumes makes it dif-ficult to code.Callionymoidei are universally recognized asmonophyletic and comprise the Draconettidae andhighly specialized Callionymidae, which we did notinclude in our analysis. The callionymoids are gen-erally considered to be closely related to the Gobie-socoidei (Gosline, 1970; Allen, 1984:629; Nelson,1994:409; Leis and Carson-Ewart, 2000:131). Nelson(1994:409-410) summarized the literature, citing dis-senting authors, and mentioned that there was no os-teological study that demonstrated a cladistic rela-tionship between the two groups. Springer (1993) hy-pothesized the monophyly of the Blennioidei [C 74]and removed it from close relationship to the "Sti-chaeoidei" (= Zoarcoidei of current authors). John-son (1993:10) and Mooi and Gill (1995:130) reportedadditional synapomophies for the Blennioidei, and itsmonophyly is currently strongly entrenched. A mor-phologically based close relationship between theGobiesocoidei and Blennioidei has not been hypoth-esized, although Rosen and Patterson (1990) notedconsiderable similarity in the dorsal gill-arch skeletonof both groups. Chen et al.'s (2003:fig. 5) molecular-based study, however, retrieved a monophyletic clade((Blenniidae + Tripterygiidae) + (Gobiesocidae)) intheir simultaneous analysis; the first two families rep-resented by one taxon each, and the third by two taxa.Support for all of Clade [C 71], except possibly forinclusion of Dactylopteridae, appears to be accumu-lating.In spite of many years study of blennioids, Spring-er has not suggested a sister group for them. Springer(1993) did, however hypothesize the monophyly ofeach of the blennioid families except the Labrisom-idae, proposing the Tripterygiidae as sister group of
all other blennioids. Springer also did not hypothe-size the interrelationships of the other families. Al-though not resolving a monophyletic Tripterygiidae,Clade [C 73] does accord with Springer's proposedposition of the family (which is monophyletic basedon other characters). No one has hypothesized amonophyletic Labrisomidae, and Springer (1993:493) suggested that there was possibly a continuumof taxa joining Labrisomidae and Chaenopsidae, in-cluding Neoclinus, which he "arbitrarily" includedin the Labrisomidae. Stepien et al. (1993), using
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rDNA and allozyme data and Tripterygiidae andBlenniidae as outgroups, analyzed the interrelation-ships of three families of blennioids, Clinidae [My-xodinae], Labrisomidae, Chaenopsidae. Their mostparsimonious results suggested (Chaenopsidae +(Labrisomidae + Chaenopsidae)), but the allozymedata suggested that Labrisomidae are paraphyletic. Ina morphological study using an array of blennioidtaxa as outgroups, Hastings and Springer (1994) hy-pothesized a monophyletic Chaenopsidae in whichNeoclinus was retrieved as the sister-group of all oth-er chaenopsids. They did not hypothesize a sistergroup for their Chaenopsidae, and left open the pos-
sibility that its composition might change in a morecomprehensive analysis. Considering the trend, it ap-pears reasonable to synonymize Labrisomidae Hubbs(1952:56) with Chaenopsidae Gill (1865:141).Sparoidei, Callanthiidae [D 114]. Johnson(1980:20), based on anatomy, first proposed Sparo-idei to comprise Sparidae, Centracanthidae, Lethrin-idae, Nemipteridae. Carpenter and Johnson (2002), ina morphological phylogenetic study, corroboratedthis composition. Orrell et al. (2002), in a molecularstudy, found support for monophyly in only one oftheir two analyses: weighted cytochrome b nucleotideanalysis. Their equally weighted nucleotide analysis,has ((Nemipteridae + Lateolabracidae + Moronidae)+ (other sparoids)). Orrell and Carpenter (2004), inanother molecular study, using 16S rRNA and cyto-chrome b retrieved only a polyphyletic Sparoidei.Neither of the molecular studies included Callanthi-idae among its outgroups, but almost all the familiesused in the outgroups were included in our analysis.SlLLAGINIDAE (PERCIDAE + RHYACICHTHYIDAE) [C49]. These three families were retrieved as a mono-phyletic clade in a polytomy [C] with a variety ofother fishes. In spite of considerable effort on the partof many workers, the interrelationships of the Go-bioidei have not been convincingly hypothesized.Winterbottom (1993b), made the most complete mor-phological search for a gobioid sister group. He de-scribed 23 putative gobioid apomorphies and foundthat the scorpaenoid Hoplichthyidae, among all thefamilies he studied, exhibited the greatest number, 11,of these apomorphies, but he was unsatisfied that aclose relationship existed between the two groups.Miya et al. (2003 :fig. 2) retrieved the gobioids as thesister group of the Dactylopteridae, which, as notedabove, we retrieved as sister group to a callionymoid,gobiesocoid, blennioid clade [C 72].Perhaps of interest, is the appearance of Sillagin-idae as a close relative of Percidae. McKay (1985:1?2) discussed the classificatory history of the sillagin-ids and quoted "W. Schwarzhans pers. comm." ashaving informed him that based on otoliths, "I havelittle doubt that [the percid] Aspro (= Zingel) reallyis the closest relative to the sillaginids." McKay went
on to write. "It appears that the family Sillaginidaeis related to the Sciaenidae, Percidae, and to a lesserextent the Haemulidae." Although our analysis didnot retrieve haemulids or sciaenids in a clade nearsillaginids, we believe that sillaginids are probablymost closely related to sciaenids. If Zingel is not sis-ter-group of all other percids, any similarity to thesillaginids is undoubtedly convergent.Miscellaneous families. Menidae [D 130] werediscussed, in part, under Smegmamorpha. AlthoughS&J's suggestion that Menidae are a pre-perciformfamily is supported by the MRT, Chen et al. (2003)found Menidae to be enmeshed in a perciform cladeof carangoids in two of their three analyses and intheir simultaneous analysis. In the third analysis(their fig. 3), Menidae is in a clade with a sphyraenid,a polynemid, and a bothid, and this clade is sister toa clade with more pleuronectiforms, a latid, and sev-eral carangoids.Sphyraenidae, Polynemidae [D 124]. These twofamilies were retrieved by our MRT as a sister group.Sphyraenidae and Polynemidae were originally as-sociated closely by Regan (1912:846-847; fullyquoted in Gosline, 1962:210, who essentially agreedwith Regan's assessment). Johnson (1993:7-8) dis-cussed the problematic classificatory history of thetwo families beginning with Gosline (1968), and con-cluded that the evidence favored a Polynemidae-Sciaendae sister group. Johnnson (1986:fig. 1) hy-pothesized that the Sphyraenidae are part of a scom-broid clade: (Scombrolabracidae (Pomatomidae(Sphyraenidae (Scombridae))). In our MRT, theSphyraenidae-Polynemidae clade is only one step re-moved from Scombridae in the clade [D 123] com-prising these three families, but which includes noother scombroids. Orrell et al. (2003), in a molecularstudy, found no support for inclusion of Sphyraeni-dae in the Scombroidei. The resolution of sphyraenidand polynemid interrelationships remains problem-
atic.Icosteidae [D 132] were discussed, in part, underSmegmamorpha. S&J presented both gill arch andnon-gill arch evidence that the family is probablymost closely related to stephanoberyciforms. TheMRT retrieved the Icosteidae as sister group of a pairof clades, one [D 134], comprising four of the fivestephanoberyciform families included in our study,and the other [D 137], comprising a mix of Holo-centridae, Ranicipitidae, and Ophidiiforms, thus lend-ing support to S&J's conclusion.Carangoidei. Our analysis included four of thefive carangoid families: Nematistiidae, Coryphaeni-dae, Rachycentridae, Carangidae (Echeneidae, not in-cluded). These were retrieved in three well separatedclades [D 97, 98, 110]. Only Rachycentridae andCarangidae appear as closely related.Haemulidae, Inermiidae. These two families were
254 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
retrieved as a sister group [D 1 13], thus corroboratingJohnson's (1980:46-48) hypothesis.Girelloidei. Johnson and Fritzsche (1989:15) con-curred with Freihofer's (1963) hypothesis that the Ra-mus Lateralis Accsessorius nerve pattern 10 charac-terized a natural assemblage of fishes. They consid-ered RLA pattern 10 as the synapomorphy unitingGirellidae, Scorpididae, Kyphosidae, Microcanthidae,Kuhliidae, Arripidae, Oplegnathidae, Terapontidae,and Stromateoidei, but did not provide a name forthe group. S&J excluded the stromateoids and ap-plied the ordinal-group name Girelloidei to the re-maining families. Of the Girelloidei, only Girellidae,Kuhliidae, and Terapontidae are included in our anal-ysis, but we include the stromateoid Centrolophidaeand its putative relative, Amarsipidae, here for pur-poses of discussion. The MRT retrieved each of thesefamilies either in well separated clades or well sep-arated within a highly branched clade, and the well-defined Terapontidae (Vari, 1978) was retrieved intwo separate clades. The results probably indicatethat gill-arch muscle and skeletal morphology as con-sidered here are either not useful in defining girelloidintra-relationships or that the Girelloidei are poly-phyletic.Grammatidae, Opistognathidae, Gerreidae,Pseudochromidae. These four families form the fourbasal stepwise clades of a series of clades [A 4] ter-minating in (Labroidei) + (Anabantoidei, Perceso-ces). The Psudochromidae and Opistognathidae havelong been associated, but usually together with othercurrently recognized families. Jordan and Snyder(1902:492) appear to be the first to have proposedthat the two are "very closely related."
Mok et al. (1990:38) first hypothesized a mono-phyly clade of three of the families, indicating (Pseu-dochromidae (Grammatidae + Opistognathidae)).Mooi and Gill (1995:121), however, disputed one ofMok et al.'s characters, nature of the M. epaxialisassociation with the dorsal-fin pterygiophores, andfurther commented, "Our continuing studies on thephylogenetic positions of Grammatidae, Opistognath-idae and other pseudochromoid families have failedto provide corroborating evidence for a sister-grouprelationship between the Grammatidae and Opistog-nathidae." This statement does not appear to rejectthe possibility that Mok et al.'s three families areclosely related.Mok et al. (1990:38) hypothesized that the Ple-siopidae (including the Acanthoclinidae) are the sis-ter group of the pseudochromoid families. Our anal-ysis has the Plesiopidae in a clade [C 64] well re-moved from the Pseudochromidae.In conclusion, a morphological cladistic analysisbased on a single complex character set strongly cor-roborates some previously hypothesized phylogeniesbased on different characters, partially supports oth-ers, has no bearing on yet others, suggests possiblerelationships that are worth further research, and im-plies relationships in some cases that seemingly makeno sense. The confusion is partly due to the taxa wechose to include and the incomplete spectrum of taxathat we had available to include, as well as to therestricted character complex. Although taxon-richcharacter-poor phylogenetic analyses currently caninvolve extended computer time and result in consid-erable homoplasy, we believe even more expandedstudies than ours will contribute to refinement of theore from Rosenblatt's mining operation.
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