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Phylogeny and Zoogeography of Glyptocephalines (Pisces: Pleuronectidae)

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Phylogeny and Zoogeography of Glyptocephalines (Pisces: Pleuronectidae) AN ABSTRACT OF THE THESIS OF Tai-sheng Chiufor the degree of _Doctor of Philosophyin Fisheries presented on July 10, 1987 Title: Phylogeny and Zoogeographyof Glyptocephalines (Pisces: Pleuronectidae) Abstract approved: Redacted for Privacy Douglas F. Markle Hypotheses of phylogenetic and historicalzoogeographic relationships of glyptocephalines (Glyptocephaluscynoglossus, G. zachirus, G. stelleri, Microstoinuskitt, H. pacificus, H. achne, Embassichthys bathybius, and Tanakius kitaharae)were constructed based on comparative anatomy andexternal morphology. The phylogenetic approachwas cladistic and character polaritywas determined by using out-groupcomparison.One nominal in-group, Platichthys bicoloratus, isshom to be a derived out-grouptaxon. Two other out-group taxa usedare Parophrys vetulus and Atheresthes stoinias. Monophyly of glyptocephalinesis suggested by the resultsof minimum distance clustering ofshape, as well as asymmetricaljaw dentition and mi-serial arrangementof jaw teeth; loss of ural neural arch; median numberof rays on fifth hypuralgreater than two; and median number of parhypural rays greater thanthree. Two monophyletic groups are identifiedwithin glyptocephalines: CladeN, Ernbassichthys and Microstomus;and Clade G, Tanakiusand Glyptocephalus. Each dade includes a primitive monotypic genus and a derived genus with three species. The most recent speciation event in each dade separated an Atlantic species from a western Pacific species. One trans-Pacific dispersal event is hypothesized for the ancestor of Clade G and one recent trans-Arctic dispersal is hypothesized for both clades. The trans-Pacific dispersal probably occurred during the Miocene. The trans-Arctic dispersal probably occurred during the Pliocene. The western North Pacific or North Pacific is the most likely the center of origin of glyptocephalines, since this area possesses the most primitive taxa. Some ontogenetic data are discussed to corroborate glyptocephaline monophyly and phylogenetic relationships. Relative timing of ontogenetic events and ontogenetic shape changes corroborate generic groupings. A more thorough study of ontogenetic biornetry may shed light on the robustness of glyptocephaline phylogeny. In addition, the specialized giant larvae of eastern North Pacific glyptocephalines deserves further study of ecologic and ontogenetic interactions. Phylogeny and Zoogeography of Glyptocephalines (Pisces: Pleuronectidae) by Tai-sheng Chiu A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Completed July 10, 1987 Commencement June 1988 APPROVED: Redacted for Privacy Associate Professor of Fisheries and Wildlife in charge of major Redacted for Privacy Head of Department of Fisheries and Wildlife Redacted for Privacy Dean of Gr School Date thesis is presented July 10, 1987 Typed by Tai-sheng Chiu for Tai-sheng Chiu ACKNOWLEDGMENTS I would like to thank the members of my graduate committee, Dr. Arthur Boucot, Dr. Bruce Coblentz, Dr. Alvin Smith, and Dr. David Thomas, for their comments and criticisms on the manuscript. I am truly appreciative of my major professor Dr. Douglas i4arkle for his assistance and encouragement, and his extraordinary patience during the weekly discussions of the last six months and his extraordinarily rapid review during the last days. I am grateful to the following persons and their institutions for loans and gifts of material used in this study: Mr. J.R. Dunn, Northwest and Alaska Fisheries Center; Mr. K.E. Hartel, Museum of Comparative Zoology, Harvard University; Mrs. G. Haynes, Ministry of Agriculture, Fisheries and Food, ngland; Mr. S.L. Jewett and Mr. L.P. Norrod, National Museum of Natural History, Smithsonian Institution; Mr. S.L. Leipertz, University of Washington; Drs. D.E. McAllister and B.W. Coad, National Museum of Canada; Dr. H.G. Moser, Southwest Fisheries Center, National Marine Fisheries Service; Dr. J. Nielsen, Zoologisk Museum, University of Copenhagen; Drs. K. Sulak and L.V. Guelpen, Atlantic Reference Centre; Dr. J.A. Seigel, Natural History Museum of Los Angeles County; Dr. A. Wheeler, British Museum (Natural History); Dr. K. Amaoka, Hokkaido University; Dr. D. Kitagawa, Iwate Prefectural Fisheries Experimental Station; Dr. K. Matsuura, National Science Museum; and Dr. T. Nakabo, Kyoto University. I would also like to thank the many scholars who provided much specimen information. I am grateful for the two-year financial support from the National Science Council and National Taiwan University, Taiwan, Republic of China. Finally, I would like to recognize and laud my wife, Yuh-chen Chern, for her oriental virtue and endurance. She took care of everything in Taiwan while I was abroad. TABLE OF CONTENTS Page INTRODUCTION 1 1. Phylogenetic history 1 2. Zoogeographic considerations 7 3. Objectives 8 MATERIALS AND METHODS 9 1. Materials 9 2. Methods 9 A. Comparative anatomy 9 B. Biometry 10 C. Phylogenetic and zoogeographic cladograrn 18 3. Terminology 18 COMPARATIVE ANATOMY 22 1.Jaw apparatus 22 2. Gill arches 34 3.Secondary body cavity and intestinal tract 60 4.Vertical fins 67 5.Pectoral girdle 71 6. Pelvic girdle 85 7.Caudal fin 86 8.Vertebral column 98 BIOMETRY 112 1. Meristic character 112 2. Morphometric character 116 A. Range of events 116 B. Ontogenetic sequence 133 C. Inter-specific comparison 162 PHYLOGENETIC CLADOGRAM 192 1. Coding of characters 192 2. Phylogenetic cladogram 197 3. Corroboration 210 ZOOGEOGRPLPHIC CLADOGRAM 219 1. Distribution 219 2. Zoogeographic cladograin 225 3. Zoogeographic scenario 229 BIBLIOGRAPHY 266 APPENDIX 276 LIST OF FIGURES Figure Page 1. Relationship of glyptocephalines based on interpretation of Norman (1934). 5 2. Relationship of glyptocephalines based on interpretation of Richardson (1981). 5 3. Relationships of glyptocephalines proposed by Sakamoto (1984, Fig. 51). The names follow Norman (1934). The names in parenthesis are generic names proposed by Sakamoto (1984). P. is composed of Pleuronectus and other taxa. A. is composed of Atheresthes and other taxa. Similarity scales has been omitted. 5 4. The set of morphological landmarks used to quantify general shape. 14 5. Lateral view of jaw apparatus of: A) G. cynoglossus, ARC 8601037, 100 mm; B) G. zachirus, OS uncatalogued, 79 mm; and C) G. stelleri, OS 11272, 172 mm. Left denotes right side upper and lower jaws and right denotes left side upper and lower jaws). 24 6. Lateral view of jaw apparatus of: A) N. kitt, OS 1805, 162 mm; B) N. pacificus, OS 3181, 117 mm; and C) N. achne, OS 11270, 167 mm. Left denotes right side upper and lower jaws and right denotes left side upper and lower jaws). 26 7. Lateral view of jaw apparatus of: A) E. bathybius, OS 7493, 197 mm; B) P. kitaharae, UW 21194, 160 mm; and C) P. bicoloratus, OS 11271 032, 111 mm. Left denotes right side upper and lower jaws and right denotes left side upper and lower jaws). 28 8. Lateral view of jaw apparatus of: A) P. vetulus, OS 5908, 59 mm; B) A. stomias, OS 3906, 98 mm. Left denotes right side upper and lower jaws and right denotes left side upper and lower jaws). 30 9. Three schematic types of tooth. 30 10. Right dorsal gill arches and pharyngobranchial teeth of G. cynoglossus, ARC 8601037, 100 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 35 List of Figures (Con't) Figure Page 11. Right dorsal gill arches and pharyngobranchial teeth of 0. zachirus, OS uncatalogued, 79 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 37 12. Right dorsal gill arches and pharyngobranchial teeth of 0. stelleri, OS 11272, 172 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 39 13. Right dorsal gill arches and pharyngobranchial teeth of N. kitt, OS 1805, 162 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 41 14. Right dorsal gill arches and pharyngobranchial teeth of N. pacificus, OS 3181, 117 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 43 15. Right dorsal gill arches and pharyngobranchial teeth of M. achne, OS 11270, 167 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 45 16. Right dorsal gill arches and pharyngobranchial teeth of E. bathybius, OS 7493, 197 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 47 17. Right dorsal gill arches and pharyngobranchial teeth of T. kitaharae, TJW 21194, 160 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 49 18. Right dorsal gill arches and pharyngobranchial teeth of P. bicoloratus, OS 11271, 111 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 51 19. Right dorsal gill arches and pharyngobranchial teeth of P. vetulus, 05 5908, 59 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 53 List of Figures (Con't) Figure Page 20. Right dorsal gill arches and pharyngobranchial teeth of A. stomias, OS 3906, 98 mm. Right denotes dorsal view of right side gill arch and left denotes ventral view of pharyngobranchial tooth plate. 55 21. Intestinal tract: A) G. cynoglossus, B) G. zachirus, C) G. stelleri, D) M. kitt. 61 22. Intestinal tract: A) N. pacificus, B) N. achne, C) E. bathybius, D) T. kitaharae. 63 23. Intestinal tract: A) P. bicoloratus, B) P. vetulus, C) A. stomias. 65 24. A) Structure of the first 5 elements of the dorsal fin. B) Structure of the anal fin elements between the first pterygiophore and first haemal spine. C) Structure of the right pelvic fin. 69 25. Lateral view of pectoral girdle of: A) G. cynoglossus, ARC 8601037, 100 mm; B) G. zachirus, OS uncatalogued, 79 mm; and C) G.
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