136 



R.D. MOOI AND AC. GILL 



EPAX 



Fig. 19 Epaxial musculature in the gadid Urophycis regia (MPM 

 31175, 133.0 mm SL). Individual slips of muscle extend from the 

 main epaxialis body to insert on the dorsal-fin pterygiophore 

 shafts under the depressores dorsales. Abbreviations and other 

 methods of presentation as in Figs 1, 3. Scale bars = 5 mm. 



tigation of epaxial muscle variation elucidates is the need to 

 shrug off the straitjacket of present classifications when 

 investigating phylogeny of higher taxa. This is particularly 

 true when the taxa are already recognized as non- 

 monophyletic, undefined, or poorly defined (e.g., Percoidei, 

 Perciformes, Paracanthopterygii), but have in essence been 

 reified over time. It is necessary to look beyond the tradi- 

 tional taxonomic boundaries, not only when dealing with 

 undefined groups such as the percoids, but also when investi- 

 gating apparently well-defined or well-established taxa such 

 as the scorpaenoids and trachinoids. Epaxial muscle inser- 

 tions to dorsal-fin pterygiophores provide one character 

 complex that illustrates the potential and novel relationships 

 that such an approach can suggest. These possible relation- 

 ships await rejection or corroboration from similar studies of 

 additional characters. 



Acknowledgements. Specimens were kindly made available by: 

 Mary Anne Rogers, Kevin Swagel, Mark Westneat (FMNH), Tony 

 Harold, Marty Rouse, Rick Winterbottom (ROM), Susan Jewett, 

 Lisa Palmer, Dave Johnson (USNM), Norma Feinberg, Melanie 

 Stiassny (AMNH), Mark McGrouther, Sally Reader, Tom Trnski 

 (AMS), Oliver Crimmen, Anne-Marie Hodges (BMNH). Earlier 

 drafts of this manuscript were reviewed by the late Humphry 

 Greenwood, Dave Johnson, Jeff Leis, Nigel Merrett, Colin Patter- 

 son, Darrell Siebert and Vic Springer; their comments were greatly 

 appreciated. This material is based upon work supported by Smithso- 

 nian postdoctoral fellowships (RDM, ACG), a Lerner-Gray 

 Research Fellowship at the American Museum of Natural History 

 (ACG), and the National Science Foundation under Grant No. 

 DEB-9317695 (RDM). 



REFERENCES 



Allen, G.R. & Swainston, R. 1988. The marine fishes of north-western Australia. 

 201 pp. Western Australian Museum, Perth. 



Anderson, M.E. 1984. On the anatomy and phylogeny of the Zoarcoidei 

 (Teleostei: Perciformes). 254 pp. Ph.D. Thesis, College of William and 

 Mary, Williamsburg, Virginia. 



Baldwin, C.C. 1990. Morphology of the larvae of American Anthiinae (Teleo- 

 stei: Serranidae), with comments on relationships within the subfamily. 

 Copeia 1990(4): 913-955. 



& Johnson, G.D. 1993. Phylogeny of the Epinephelinae (Teleostei: 



Serranidae). Bulletin of Marine Science 52(1): 240-283. 



Balushkin, A.V. & Voskoboynikova, O.S. 1990. A new family, Bathylutichthy- 

 idae (Cottoidei, Scorpaeniformes), for the deepwater fish Bathylutichthys 

 taranetzi gen. et sp. nov. from South Georgia Island (Antarctica). Journal of 



Ichthyology 30: 185-191 [originally published in Russian in Voprosy ikhti- 



ologii 30: 185-191 (1989)]. 

 Chapleau, F. 1993. Pleuronectiform relationships: a cladistic reassessment. 



Bulletin of Marine Science 52(1): 516-540. 

 Day, F. 1875. The fishes of India; being a natural history of fishes known to 



inhabit the seas and fresh waters of India, Burma, and Ceylon. Part 1 . 168 p. 



William Dawson & Sons, Ltd., London. 

 Fowler, H.W. & Bean, B.A. 1922. Fishes from Formosa and the Philippine 



Islands. Proceedings of the United States National Museum 62(2448): 1-73. 

 Fraser, T.H. 1968. Comparative osteology of the Atlantic snooks (Pisces. 



Centropomus) . Copeia 1968(3): 433-460. 

 Gill, A.C. & Mooi, R.D. 1993. Monophyly of the Grammatidae and of the 



Notograptidae, with evidence for their phylogenetic positions among perci- 



forms. Bulletin of Marine Science 52(1): 327-350. 

 Gosline, W.A. 1966. The limits of the fish family Serranidae, with notes on 



other lower percoids. Proceedings of the California Academy of Sciences, 4th 



Series 33(6): 91-112. 

 1968. The suborders of perciform fishes. Proceedings of the United Stales 



National Museum 124(3647): 1-78. 



1970. A reinterpretation of the teleostean fish order Gobiesociformes. 



Proceedings of the California Academy of Sciences, 4th Series 38(19): 

 363-382. 



Greenwood, P.H. 1976. A review of the family Centropomidae (Pisces. 

 Perciformes). Bulletin of the British Museum (Natural History), Zoology 

 series 29(1): 1-81. 



Ishida, M. 1994. Phylogeny of the suborder scorpaenoidei (Pisces: Scorpaeni- 

 formes). Bulletin of Nansei National Fisheries Research Institute 27: 1-112. 



Johnson, G.D. 1983. Niphon spinosus: a primitive epinepheline serranid. with 

 comments on the monophyly and intrarelationships of the Serranidae. 

 Copeia 1983(3): 777-787. 



1984. Percoidei: development and relationships. In Moser, H.G., Rich- 

 ards, W.J., Cohen, D.M., Fahay, M.P., Kendall Jr., A.W. & Richardson. 

 S.L. (eds), Ontogeny and systematics of fishes. American Society of Ichthy- 

 ologists and Herpelologists Special Publication 1: 464-498. 



1986. Scombroid phylogeny: an alternative hypothesis. Bulletin of Marine 



Science 39(1): 1-41. 



1988. Niphon spinosus, a primitive epinepheline serranid: corroborative 



evidence from the larvae. Japanese Journal of Ichthyology 35(1): 7-18. 



1993. Percomorph phylogeny: progress and problems. Bulletin of Marine 



Science 52(1): 3-28. 



& Anderson, Jr., W.D. (eds) 1993. Proceedings of the symposium on 



phylogeny of Percomorpha, June 15-17, 1990, held in Charleston, South 

 Carolina at the 70th annual meetings of the American Society of Ichthyolo- 

 gists and Herpetologists. Bulletin of Marine Science 52: 1-626. 



& Patterson, C. 1993. Percomorph phylogeny: a survey of acanthomorphs 



and a new proposal. Bulletin of Marine Science 52(1): 554-626. 



Jordan, D.S. 1923. A classification of fishes including families and genera as far 

 as known. Stanford University Publications, University Series, Biological 

 Sciences 3(2): 77-243. 



Kaufman, L. & Liem, K.F. 1982. Fishes of the suborder Labroidei (Pisces: 

 Perciformes): phylogeny, ecology, and evolutionary significance. Breviora, 

 Museum of Comparative Zoology 472: 1-19. 



Kendall, Jr., A.W. 1984. Serranidae: development and relationships. In 

 Moser, H.G., Richards, W.J., Cohen, D.M., Fahay, M.P., Kendall Jr.. 

 A.W. & Richardson, S.L. (eds), Ontogeny and systematics of fishes. Ameri- 

 can Society of Ichthyologists and Herpetologists Special Publication 1: 

 499-510. 



Kido, K. 1988. Phylogeny of the family Liparididae, with the taxonomy of the 

 species found around Japan. Memoirs of the Faculty of Fisheries, Hokkaido 

 University 35(2): 125-256. 



Leis, J.M. 1984. Tetraodontiformes: relationships. In Moser, H.G., Richards, 

 W.J., Cohen, D.M., Fahay, M.P., Kendall Jr., A.W. & Richardson. S.L. 

 (eds), Ontogeny and systematics of fishes. American Society of Ichthyologists 

 and Herpetologists Special Publication 1: 459-463. 



Leviton, A.E., Gibbs, Jr., R.H., Heal, E. & Dawson, C.E. 1985. Standards in 

 herpetology and ichthyology. Part I: Standard symbolic codes for institu- 

 tional resource collections in herpetology and ichthyology. Copeia 1985(3): 

 802-832. 



Mandrytsa, S.A. 1991. The structure of the seismosensory system of the 

 members of the family Pataecidae (Pisces, Acanthopterygii). USSR Acad- 

 emy of Sciences, Proceedings of the Zoological Institute, Leningrad 235: 

 29-36. (In Russian with English summary.) 



Mok, H.-K., Chang, H.-J. & Lee, C.-Y. 1990. Phylogenetic interrelationship of 

 the perciform Acanthoclinidae, Grammidae, Plesiopidae, Pseudochromidae 

 and Opistognathidae. Bulletin of the Institute of Zoology. Academia Sinica 

 29(1): 29-39. 



