Polyphyly of the mail-cheeked fishes (Teleostei: Scorpaeniformes): Evidence from mitochondrial and nuclear sequence data

Molecular Phylogenetics and Evolution

Volumn 32, Issue 2, 2004, Pages 627-646

  Smith, Wm Leo ^a,b   Wheeler, Ward C ^a  

^a DIVISION OF INVERTEBRATE ZOOLOGY   (United States)

^b Och Spine at New York Presbyterian Hospitals   (United States)

Author keywords

Anoplopomatoidei; Cottoidei; Hexagrammoidei; Platycephaloidei; Scorpaeniformes; Scorpaenoidei; Trichodontidae; Zaniolepidoidei

Indexed keywords

MITOCHONDRIAL DNA;

ANIMAL; ARTICLE; CELL NUCLEUS; CLASSIFICATION; DNA SEQUENCE; FISH; GENETICS; PHYLOGENY;

ANIMALS; CELL NUCLEUS; DNA, MITOCHONDRIAL; FISHES; PHYLOGENY; SEQUENCE ANALYSIS, DNA;

ACANTHOMORPHA; ANOPLOPOMATOIDEI; ATHERINIFORMES; BLENNIOIDEI; COTTOIDEA; COTTOIDEI; GASTEROSTEOIDEI; GRAMMATIDAE; HEXAGRAMMOIDEI; NOTOTHENIOIDEI; PERCIDAE; PISCES; PLATYCEPHALOIDEI; SCORPAENIFORMES; SCORPAENOIDEI; TELEOSTEI; TRICHODONTIDAE; ZOARCOIDEI;

EID: 3042541169     PISSN: 10557903     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.ympev.2004.02.006     Document Type: Article

References (118)

1. Allen G.R., Midgely S.H., Allen M. Field Guide to the Freshwater Fishes of Australia. 2002;Western Australian Museum, Perth
2. Andriashev A.P. A new fish of the lumpfish family (Pisces, Liparidae) found at a depth of more than 7 kilometers. Trudy Inst. Okeanol. 12:1955;340-344. (in Russian)
3. Baldwin C., Johnson G.D. Phylogeny of the Epinephelinae (Teleostei: Serranidae). Bull. Mar. Sci. 52:1993;240-280
4. Balushkin A.V. Morphology, classification, and evolution of notothenioid fishes of the Southern Ocean (Notothenioidei, Perciformes). J. Ichthyol. 40:2000;S74-S109
5. Berg L.S. Freshwater Fishes of the U.S.S.R. and Adjacent Countries. fourth ed. 1948;Israel Program for Scientific Translations Ltd, Jerusalem
6. Bolin R.L. The evolution of the marine Cottidae of California with a discussion of the genus as a systematic category. Stanford Ichthyol. Bull. 3:1947;1-135
7. Bowne P.S. Systematics and morphology of Gasterosteiformes. Bell M.A., Foster S.A. Evolutionary Biology of the Threespine Stickleback. 1994;28-60 Oxford University Press, Oxford
8. Breder C.M. Jr., Rosen D.E. Modes of Reproduction in Fishes. 1966;The Natural History Press, New York
9. Bremer K. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution. 42:1988;795-803
10. Bremer K. Branch support and tree stability. Cladistics. 10:1995;295-304
11. Burnett J.W. Aquatic adversaries: stonefish. Cutis. 62:1998;269-270
12. Busby M.S. Guide to the identification of larval and early juvenile poachers (Scorpaeniformes: Agonidae) from the northeastern Pacific Ocean and Bering Sea. NOAA Tech. Rep. 137:1998;1-88
13. Chen W.J., Bonillo C., Lecointre G. Repeatability of clades as a criterion of reliability: a case study for molecular phylogeny of Acanthomorpha (Teleostei) with larger number of taxa. Mol. Phylogenet. Evol. 26:2003;262-288
14. Colgan D.J., McLauchlan A., Wilson G.D.F., Livingston S.P., Edgecombe G.D., Macaranas J., Cassis G., Gray M.R. Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Aust. J. Zool. 46:1998;419-437
15. Cuvier, G., 1829. Le Règne Animal, distribuéd'après son organisation, pour servir de base àl'histoire naturelle des animaux et d'introduction àl'anatomie comparée, second ed. Règne Animal v. 2
16. Davies P.L., Ewart K.V., Fletcher G.L. The diversity and distribution of fish antifreeze proteins: new insights into their origins. Hochachka P.W., Mommsen T.P. Biochemistry and Molecular Biology of Fishes. vol. 2:1993;279-291 Elsevier Science, Amsterdam
17. Eakin R.R. Biology of the antarctic seas. X. Osteology and relationships of the fishes of the Antarctic family Harpagiferidae (Pisces, Notothenioidei). Antarctic Res. Ser. 31:1981;81-147
18. Elmerot C., Amason U., Gojobori T., Janke A. The mitochondrial genome of the pufferfish, Fugu rubripes, and ordinal teleostean relationships. Gene. 295:2002;163-172
19. Eschmeyer W.N. Catalog of Fishes: Volume 3 - Genera of Fishes, Species and Genera in a Classification, Literature Cited, Appendices. 1998;California Academy of Sciences, San Francisco
20. Eschmeyer W.N., Hureau J.C. Sebastes mouchezi, a senior synonym of Helicolenus tristanensis, with comments on Sebastes capensis and zoogeographical considerations. Copeia. 1971:1971;576-579
21. Farris J.S. The retention index and the rescaled consistency index. Cladistics. 5:1989;417-419
22. Feller A.E., Hedges S.B. Molecular evidence for the early history of living amphibians. Mol. Phylogenet. Evol. 9:1998;509-516
23. Freihofer W.C. Patterns of the ramus lateralis accessorius and their systematic significance in teleostean fishes. Stanford Ichthyol. Bull. 8:1963;80-189
24. Freihofer W.C. Cranial nerves of a percoid fish, Polycentrus schomburgkii (family Nandidae), a contribution to the morphology and classification of the order Perciformes. Occ. Pap. Cal. Acad. Sci. 128:1978;1-78
25. Gill T. On the classification of the mail-cheeked fishes. Proc. U.S. Nat. Mus. 11:1888;567-592
26. Gill A.C. Comments on an intercalary path for the glossopharyngral (cranial IX) nerve as a synapomorphy of the Paracanthopterygii and on the phylogenetic position of the Gobiesocidae (Teleostei: Acanthomorpha). Copeia. 1996:1996;1022-1029
27. Gill A.C., Mooi R.D. Monophyly of the Grammatidae and of the Notograptidae, with evidence for their phylogenetic position among perciforms. Bull. Mar. Sci. 52:1993;327-350
28. Giribet G., Ribera C. A review of arthropod phylogeny: new data based on ribosomal DNA sequences and direct character optimization. Cladistics. 16:2000;204-231
29. Goloboff, P.A., 1998. NONA ver. 3.0. Program and documentation. Fundación e Instituto Miguel Lillo, Tucumán, Argentina
30. Goloboff P.A. Analyzing large data sets in reasonable times: solutions for composite optima. Cladistics. 15:1999;415-428
31. Gosline W.A. The suborders of perciform fishes. Proc. U.S. Nat. Mus. 124:1968;1-78
32. Gosline W.N. Functional Morphology and Classification of Teleostean Fishes. 1971;University of Hawaii Press, Honolulu
33. Greenwood P.H., Rosen D.E., Weitzman S.H., Myers G.S. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bull. Am. Mus. Nat. Hist. 131:1966;341-455
34. Hall T.A. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl. Acids. Symp. Ser. 41:1999;95-98
35. Hallacher L.E. The comparative morphology of extrinsic gasbladder musculature in the scorpionfish genus Sebastes (Pisces: Scorpaenidae). Proc. Cal. Acad. Sci. 40:1974;59-86
36. Halstead B.W. Poisonous and Venomous Marine Animals of the World. 1988;U.S. Government Printing Office, Washington, DC
37. Hillis D.M., Dixon M.T. Ribosomal DNA: molecular evolution and phylogenetic inference. Quart. Rev. Biol. 66:1991;411-453
38. Imamura H. Phylogeny of the family Platycephalidae and related taxa. Species Div. 1:1996;123-233
39. Imamura H. An alternative hypothesis on the phylogenetic position of the family Dactylopteridae (Pisces: Teleostei), with a proposed new classification. Ichthyol. Res. 47:2000;203-222
40. Imamura H., Shinohara G. Phylogenetic studies of the order Scorpaeniformes (Pisces: Acanthopterygii): progress, present condition and problems. Jpn. J. Ichthyol. 44:1997;77-95. (In Japanese with English summary)
41. Imamura H., Shinohara G. Scorpaeniform fish phylogeny: an overview. Bull. Nat. Sci. Mus., Ser. A. 24:1998;185-212
42. Imamura H., Yabe M. Demise of the Scorpaeniformes (Actinopterygii: Percomorpha): an alternative phylogenetic hypothesis. Bull. Fish. Sci., Hokkaido Univ. 53:2002;107-128
43. Ishida M. Phylogeny of the suborder Scorpaenoidei (Pisces: Scorpaeniformes). Bull. Nansei Nat. Fish. Res. Inst. 27:1994;1-112
44. Jackson K.L., Nelson J.S. Ambophthalmos, a new genus for Neophrynichthys angustus and Neophrynichthys magnicirrus, and the systematic interrelationships of the fathead sculpins (Cottoidei, Psychrolutidae). Can. J. Zool. 76:1998;1344-1357
45. Johns G.C., Avise J.C. Tests for ancient species flocks based on molecular phylogenetic appraisals of Sebastes rockfishes and other marine fishes. Evolution. 52:1998;1135-1146
46. Johnson G.D. Niphon spinosus: a primitive epinepheline serranid, with comments on the monophyly and intrarelationships of the Serranidae. Copeia. 1983:1983;777-787
47. Johnson G.D. Percoidei: development and relationships. Moser H.G., Richards W.J., Cohen D.M., Fahay M.P., Kendall A.W. Jr., Richardson S.L. Ontogeny and Systematics of Fishes. American Society of Ichthyologists and Herpetologists Special Publication 1. 1984;464-498 Allen Press, Lawrence
48. Johnson G.D. Percomorph phylogeny: progress and problems. Bull. Mar. Sci. 52:1993;3-28
49. Johnson G.D., Patterson C. Percomorph phylogeny: a survey of acanthomorphs and a new proposal. Bull. Mar. Sci. 52:1993;554-626
50. Jungersen, H.F.E., 1910. Ichthyological contributions. II. The structure of the Aulostomidæ, Sygnathidæ and Solenostomidæ. D. Kgl. Danske Vidensk Selsk. Skrifter 8 (7), 267-364
51. Kai Y., Kouji N., Nakabo T. Molecular phylogenetic perspective on speciation in the genus Sebastes (Scorpaenidae) from the Northwest Pacific and the position of Sebastes within the subfamily Sebastinae. Ichthyol. Res. 50:2003;239-244
52. Kanayama T. Taxonomy and phylogeny of the family Agonidae (Pisces: Scorpaeniformes). Mem. Fac. Fish., Hokkaido Univ. 38:1991;1-199
53. Keenan C.P. Phylogeny of Australian species of flatheads (Teleostei, Platycephalidae) as determined by allozyme electrophoresis. J. Fish Biol. 39:1991;237-249
54. Kendall A.W., Vinter B. Development of hexagrammids (Pisces: Scorpaeniformes) in the Northeastern Pacific Ocean. NOAA Tech. Rep. 2:1984;1-44
55. Kido K. Phylogeny of the family Liparididae with the taxonomy of the species found around Japan. Mem. Fac. Fish., Hokkaido Univ. 35:1988;125-256
56. Kizer K.W., McKinney H.E., Auerbach P.S. Scorpaenidae envenomation: a five-year Poison Center experience. JAMA. 253:1985;807-810
57. Kluge A.G., Farris J.S. Quantitative phyletics and the evolution of anurans. Syst. Zool. 18:1969;1-32
58. Kocher T.D., Thomas W.K., Meyer A., Edwards S.V., Paabo S., Villablanca X.F., Wilson A.C. Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proc. Natl. Acad. Sci. USA. 86:1989;6196-6200
59. Kochzius M., Söller R., Khalaf M.A., Blohm D. Molecular phylogeny of the lionfish genera Dendrochirus and Pterois (Scorpaenidae, Pteroinae) based on mitochondrial DNA sequences. Mol. Phylogenet. Evol. 28:2003;396-403
60. Kontula T., Kirilchik S.V., Väinölä R. Endemic diversification of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing. Mol. Phylogenet. Evol. 27:2003;143-155
61. Koya Y., Matsubara T. Ultrastructural observations on the inner ovarian epithelia of kichiji rockfish Sebastlobus macrochir with special reference to the production of gelatinous material surrounding the eggs. Bull. Hokkaido Nat. Fish. Res. Inst. 59:1995;1-17
62. Le H.L.V., Lecointre G., Perasso R. A 28S rRNA-based phylogeny of the gnathostomes: first steps in the analysis of conflict and congruence with morphologically based cladograms. Mol. Phylogenet. Evol. 2:1993;31-51
63. Leipertz S.L. The rockhead poacher, Bothragonus swani (Teleostei: Agonidae): Selected osteology, with comments on phylogeny. Copeia. 1988:1988;64-71
64. Leis J.M., Rennis D.S. Scorpaenidae (scorpionfishes, stonefishes). Leis J.M., Carson-Ewart B.M. The Larvae of Indo-Pacific Coastal Fishes: An Identification Guide to Marine Fish Larvae. 2000;226-235 Brill, Leiden
65. Leis J.M., Trnski T. Larval development of the Indo-Pacific perciform fish, Centrogenys vaigiensis (Pisces: Centrogeniidae). Ichthyol. Res. 46:1999;413-418
66. Love M.S., Yoklavich M.M., Thorsteinson L. The Rockfishes of the Northeast Pacific. 2001;University of California Press, Berkeley
67. Lovejoy N. Reinterpreting recapitulation: systematics of needlefishes and their allies (Teleostei: Beloniformes). Evolution. 54:2000;1349-1362
68. Mandrytza, S.A., 2001. The seismosensory system and classification of scorpaenoid fishes. Izdatelstvo Permskogo Universiteta. (in Russian with English summary)
69. Matarese A.C., Kendall A.W. Jr., Blood D.M., Vinter B.M. Laboratory guide to early life history stages of northeast Pacific fishes. NOAA Tech. Rep. 80:1989;1-652
70. Matsubara K. Studies of the scorpaenoid fishes of Japan: anatomy, phylogeny, and taxonomy (I). Trans. Sigenkagaku Kenkyusho. 1:1943;1-170
71. Matsubara K., Ochiai A. A revision of the Japanese fishes of the family Platycephalidae (the flatheads). Mem. Col. Agri., Kyoto Univ. 68:1955;1-109
72. Miya M., Kawaguichi A., Nishida M. Mitogenomic exploration of higher teleostean phylogenies: a case study for moderate-scale evolutionary genomics with 38 newly determined complete mitochondrial DNA sequences. Mol. Biol. Evol. 18:2001;1993-2009
73. Miya M., Takeshima H., Endo H., Ishiguro N.B., Inoue J.G., Mukai T., Satoh T.P., Yamaguchi M., Akira K. Major patterns of higher teleostean phylogenies: a new perspective based on 100 complete mitochondrial DNA sequences. Mol. Phylogenet. Evol. 26:2003;121-138
74. Mooi R.D., Gill A.C. Association of epaxial musculature with dorsal-fin pterygiophores in acanthomorph fishes, and its phylogenetic significance. Bull. Nat. Hist. Mus., London (Zool.). 61:1995;121-137
75. Mooi R.D., Johnson G.D. Dismantling the Trachinoidei: evidence of a scorpaenoid relationship for the Champsodontidae. Ichthyol. Res. 44:1997;143-176
76. Moser H.G. Reproduction and development of Sebastodes paucispinis and comparison with other rockfishes off Southern California. Copeia. 1967:1967;773-797
77. Moyer J.T., Zaiser M.J. Social organization and spawning behavior of the pteroine fish Dendrochirus zebra at Miyake-jima, Japan. Jpn. J. Ichthyol. 28:1981;52-69
78. Munehara H., Koya Y., Hayakawa Y., Takani K. Extracellular environments for the initiation of external fertilization and micropylar plug formation in a cottid species, Hemitripterus villosus (Pallas) (Scorpaeniformes) with internal insemination. J. Exp. Mar. Biol. Ecol. 211:1997;279-289
79. Musick J.A., Harbin M.M., Berkeley S.A., Burgess G.H., Eklund A.M., Lindley L., Gilmore R.G., Golden J.T., Ha D.S., Huntsman G.R., McGovern J.C., Parker S.J., Poss S.G., Sala E., Schmidt T.W., Sedberry G.R., Weeks H., Wright S.G. Marine, estuarine, and diadromous fish stocks at risk of extinction in North America (exclusive of pacific salmonids). Fisheries. 25:2000;6-29
80. Nazarkin M.V., Voskoboinikova O.S. New fossil genus and species of Trichodontidae and the position of this family in the order Perciformes. J. Ichthyol. 40:2000;687-703
81. Nelson J.S. Fishes of the World. third ed. 1994;Wiley, New York
82. Nixon K.C. The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics. 15:1999;407-414
83. Nixon, K.C., 2000. WinClada (beta) ver. 0.9.9. Published by the author, Ithaca, NY
84. Norman J.R. On the affinities of the Chilean fish, Normanichthys crockeri Clark. Copeia. 1938:1938;29-32
85. O'Bannon, B.K. (Ed.), 2001. Fisheries of the United States 2000. NOAA, NMFS, Silver Spring
86. Okiyama M. An Atlas of the Early Stage Fishes in Japan. 1988;Tokai University Press, Kanagawa
87. Palumbi S.R. Nucleic acids II: the polymerase chain reaction. Hillis D.M., Moritz C., Mable B.K. Molecular Systematics. second ed. 1996;205-247 Sinauer, Sunderland
88. Parenti L.R. Relationships of atherinomorph fishes (Teleostei). Bull. Mar. Sci. 52:1993;170-196
89. Pietsch T.W. Evolutionary relationships of the sea moths (Teleostei: Pegasidae) with a classification of gasterosteiform families. Copeia. 1978:1978;517-529
90. Quast J.C. Osteological characteristics and affinities of the hexagrammid fishes, with a synopsis. Proc. Cal. Acad. Sci. 31:1965;563-600
91. Regan C.T. XIV. - The osteology and classification of the teleostean fishes of the order Scleroparei. Ann. Mag. Nat. Hist. 11:1913;169-184
92. Richards W.J., Jones D.L. Preliminary classification of the gurnards (Triglidae: Scorpaeniformes). Mar. Freshwater Res. 53:2002;275-282
93. Richardson S.L. Current knowledge of larvae of sculpins (Pisces: Cottidae and allies) in northeast Pacific genera with notes on intergeneric relationships. Fish. Bull. 79:1981;103-121
94. Rocha-Olivares A., Rosenblatt R.H., Vetter R.D. Molecular evolution, systematics, and zoogeography of the rockfish subgenus Sebastomus (Sebastes, Scorpaenidae) based on mitochondrial cytochrome b and control region sequences. Mol. Phylogenet. Evol. 11:1999;441-458
95. Rocha-Olivares A., Kimbrell C.A., Eitner B.J., Vetter R.D. Evolution of a mitochondrial cytochrome b gene sequence in the species-rich genus Sebastes (Teleostei, Scorpaenidae) and its utility in testing the monophyly of the subgenus Sebastomus. Mol. Phylogenet. Evol. 11:1999;426-440
96. Rosen D.E. An essay on euteleostean classification. Am. Mus. Novitates. 2827:1985;1-57
97. Rosen D.E., Patterson C. On Müller's and Cuvier's concepts of pharyngognath and labyrinth fishes and the classification of percomorph fishes, with an atlas of percomprph dorsal gill arches. Am. Mus. Novitates. 2893:1990;1-57
98. Shinohara G. Comparative morphology and phylogeny of the suborder Hexagrammoidei and related taxa. Mem. Fac. Fish., Hokkaido Univ. 41:1994;1-97
99. Sideleva, V.G., 1994. Speciation of endemic Cottoidei in Lake Baikal. In: Martens, K., Goddeeris, B., Coulter, G. (Eds.), Speciation in Ancient Lakes. Arch. Hydrobiol. Beih. Ergebn. Limnol., Schweizerbart, Stuttgart, vol. 44, pp. 441-450
100. Slaughter D., Fletcher G.L., Ananthanarayanan V.S., Hew C.L. Antifreeze protein genes from the sea raven, Hemitripterus americanus: further evidence for diversity among fish polypeptide antifreezes. J. Biol. Chem. 256:1981;2022-2026
101. Starks E.C. The primary shoulder girdle of the bony fishes. Stanford Univ. Publ. Univ. Ser.: Biol. Sci. 6:1930;147-239
102. Stiassny M.L.J. The limits and relationships of the acanthomorph teleosts. J. Zool., London. 1:1986;411-460
103. Stiassny, M.L.J., Wiley, E.O., Johnson, G.D., de Carvalho, M.R., in press. Gnathostome fishes. In: Donaghue, M.J., Cracraft, J. (Eds.), Assembling the Tree of Life. Oxford University Press, New York
104. Streelman J.T., Karl S.A. Reconstructing labroid evolution with single-copy nuclear DNA. Proc. Roy. Soc. Lond. B. 264:1997;1011-1020
105. Taranets A.Y. K. klassifikatsii i proiskhotzhdenif bychov semeystva Cottidae. Izvestiya Akad. Nauk SSSR. Otd. Biol. 3:1941;427-447. (Translated by Wilmovsky, N.J. and E. Lanz, Published as Inst. Fish., Univ. British Columbia, Mus.Contrib. 425: 421-428)
106. Titus, T.A., 1992. A phylogenetic analysis of the Desmognathinae (Caudata: Plethodontidae): Evolutionary patterns inferred from mitochondrial DNA sequences. Unpubl. Ph.D. Diss, Univ. of Kansas, Lawrence
107. Velez J.A., Watson W., Sandknop E.M., Arntz W., Wolff M. Larval and osteological development of the mote sculpin (Normanichthys crockeri) (Pisces: Normanichthyidae) from the Independencia Bight, Pisco, Peru. J. Plank. Res. 25:2003;279-290
108. Washington B.B., Eschmeyer W.N., Howe K.M. Scorpaeniformes: relationships. Moser H.G., Richards W.J., Cohen D.M., Fahay M.P., Kendall A.W. Jr., Richardson S.L. Ontogeny and Systematics of Fishes. American Society of Ichthyologists and Herpetologists Special Publication 1. 1984;438-447 Allen Press, Lawrence
109. Weitzman S.H. The osteology of Brycon meeki, a generalized characid fish, with an osteological definition of the family. Stanford Ichthyol. Bull. 8:1962;3-77
110. Wheeler W.C. Optimization alignment: the end of multiple sequence alignment in phylogenetics. Cladistics. 12:1996;1-9
111. Wheeler W.C. Iterative pass optimization of sequence data. Cladistics. 19:2003;254-260
112. Wheeler W.C. Implied alignment: a synapomorphy-based multiple-sequence alignment method and its use in cladogram search. Cladistics. 19:2003;261-268
113. Wheeler W.C., Gladstein D., DeLaet J. POY Version 3.0 and Command Summary. 2002;American Museum of Natural History, New York
114. Wiley E.O., Johnson G.D., Dimmick W.W. The interrelationships of acanthomorph fishes: a total evidence approach using molecular and morphological data. Biochem. Syst. Ecol. 28:2000;319-350
115. Winterbottom R. A descriptive synonymy of the striated muscles of the Teleostei. Proc. Acad. Nat. Sci. Philadelphia. 125:1974;225-317
116. Yabe M. Osteological review of the family Icelidae Berg, 1940, (Pisces; Scorpaeniformes), with comment on the validity of this family. Bull. Fac. Fish., Hokkaido Univ. 32:1981;293-315
117. Yabe M. Comparative osteology and myology of the superfamily Cottoidea (Pisces: Scorpaeniformes) and its phylogenetic classification. Mem. Fac. Fish., Hokkaido Univ. 32:1985;1-130
118. Yabe M., Uyeno T. Anatomical description of Normanichthys crockeri (Scorpaeniformes, Incertae sedis: Family Normanichthyidae). Bull. Mar. Sci. 58:1996;494-510