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+Atractosteus Atrox (Leidy) Figures 48B, 50A, 52A, 53D, 55D, 61, 62 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Abstract The relationships of gars to other actinopterygian. fishes, the number of recognizable species of gars, the relationships of these species, and the biogeography of these species were investigated. The phylogenetic method of Hennig (1966) was used to analyze current hypotheses concerning the relationships among gars and of gars to other actinopterygian groups. Hennig's method is discussed and several points taken up in detail. Croizat's (1958) method of biogeographic analysis was used to describe the major features of gar biogeography. Gars were found to comprise a monophyletic group. Gars are the sister-group of the Halecostomi (Amiidae plus Teleostei). These three taxa comprise another monophyletic group, the Neopterygii. The Neopterygii is the sister-group of the Chondrostei. These conclusions corroborate certain previous hypotheses and refute others. Sixteen species of gars are recognized. They are split - equally among two genera, Lepisosteus and Atractosteus. The genus Lepisosteus includes an undescribed fossil species from the Cretaceous of Montana and the following seven species in approximate phylogenetic order: L. cuneatus (Eocene, North America); L. platostomus (Recent, North , America); L. indicus (Cretaceous, India); L. osseus (Recent, North America); L. finbratus(Eocene and Oligocene, Europe); Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Acknowledgements To my colleagues and friends at the American Museum of Natural History I express my thanks for many hours of discussion about gars and systematic theory. Special thanks go to Dr. Donn E. Rosen whose help and encouragement throughout the project are gratefully acknowledged. Thanks also to Dr. Gareth Nelson who spent many hours discussing problems of fish phylogeny and systematics with me and who taught me many techniques used during the project. Dr. Bobb Schaeffer gave me valuable advice and guidance on the fossils. Mr. Walter Sorensen helped and guided me in fossil preparation. The section concerning ancestor recognition grew out of a dialogue with my colleague George Englemann; we expect to publish our thoughts on this subject in an expanded joint paper. Ms. Louise LoPresti typed the rough draft, Ms. Lynne Judge and Ms. Joan Mey typed the final draft. Their skills are appreciated. My thanks to Dr. Colin Patterson (British Museum, Natural History) and M. Daniel Gouget (Museum National d'Histoire Naturelle) for their help during my visit abroad. Thanks also to Dr. Darrell D. Hall (Sam Houston State University) and his students for assisting with the field work in Texas. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table of Contents Introduction ...................................... 1 Materials and Methods .............................. 13 Systematic methodology ......................... 18 Biogeographic method ........................... 37 Material examined .............................. 40 Anatomical abbreviations ....................... 46 Relationships of gars to other actinopterygian fishes 50 The skull ..................................... 51 The hyoid and visceral arches ...................75 Post cranial skeleton .......................... 90 Summary hypotheses of actinopterygian relationships ..................................97 Systematics accounts ............................... 100 Division Ginglymodi ............................ 100 Family Lepisosteidae ........................... 104 Genus Lepisosteus .............................. 106 Genus Atractosteus ............................. 136 Phylogenetic Relationships among gars ............... 165 Monophyly of the genera ........................ 166 Relationships among Lepisosteus gars ............ 172 Relationships among Atractosteus gars ........... 176 A classification of gars ........................... 182 Gar biogeography .................................. 186 Summary of the study ............................... 192 Literature cited ........................ 196 Tables ............................................ 248 Figures ........................................... 252 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. List of Figures Figure Page 1. Four hypotheses of the relationships between chondrosteans (C), gars (a), amiids (A), teleosts (T), and semionotids (S). a., after Nelson, 1969a; b. after Jessen, 1972; c. after Patterson, 1973; d. after Rayner, 1948 ................................ 252 2. The skull of Lepisosteus oculatus (LACM 33916-3). Upper, dorsal view; lower, lateral view ....... 253 3. Lateral view of the snout of five teleostome fishes. a. Lepisosteus; b. Amia; C. Elops; d. Eustenopteron; e. Pteronfinius. (a, b after Jollie, 1969; c after Forey, 1972; d, e after Stensio, 19)47) .................... 254 4. Dorsal view of the posterior skull of four act inopterygians. A, Pteronisculus; B, Lepisosteus; C, Amia; D. pops: A after Neilsen, 1942; B, C a ter Patterson, 1973; D after Forey, 1972 ................... 255 5. Pterygoid bones of Atractosteus tristoechus (USNM 111309). a., dorsal view; b., lateral view. Dermopalatine omitted ................... 256 6. Medial view of the post-orbital region of Atract- osteus tropicus (AMNH 27939). Suspeni3Far- ossifications unstippled, other bones stippled ....................................... 257 7 Lower jaw of Atractosteus spatula (AMNH uncat.). Upper, lateral view; lower, medial view ..... 258 ossifications of Lepisosteus 8. Neurocranial ) oculatus (LACM 33914-2). Top, posterior view; middle, ventral view; bottom, lateral view .... 259 9. Diagramatic lateral view of the hyoid apparatus of a gar. Bone stippled ....................... 260 10. -Tongue" of Lepisosteus oseeus (AMNH 509). Dorsal view .................................... 261 11. Lateral view of the oasibranchial copula of four actinopterygians. a. Lepisosteus platostomus; b. Amia calva; C. Polyodon spatula; d. Pol- ypterus sp. Cartilage stippled, hypobranchial articulatory surfaces black, pone unstippled. Redrawn after Nelson, 1969a ................. 262 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure Page 12. Ventral view of left dorsal gill arch elements of (A.) "Gogo" palaeoniscid, (B) Atractosteus tropicus, and (C) Amia calva. Suprapharyngo- branchials not shown in B or C. (A. after Gardiner, 1973) ............................. 263 13. Dorsal view of dorsal gill arch elements of (A.) Amia calva; (B.) Lepisosteus oculatus; (C) Albula vulpes,. Semidiagramatic, bone stippled, dorsal aorta unstippled ........... 264 14. Ventral views of the posterior gill arches of Amia calva (left) and Atractosteus spatula T7IihIT7--Semidiagramatic, bone and cartilage stippled, tendons white ..................... 265 15. Pectoral girdle of Lepisosteus osseus, medial view, endoskeletal mesocorocoid stippled ............ 266 16. Vertebrae of Atractosteus africanus (MHNP N.29, length of ventral ridge, 13mm ) ............... 267 17. Alternate hypothesis of the relationships of chondrosteans (C), gars (G), amiids (A), and teleosts (T). (a.) Least refuted hypothesis; (b.) alternate hypothesis with corroborating characters. Synapomorphies (blacl; rectangles) connecting taxa are: (1) a dermal intercalar; (2) a posterior myodome; (3) a supramaxilla; (4) a post-temporal process; (5) loss of in- ternares commissure; (6) an endochondrali rostral; (7) maxilla with internal articula- tory head; (8) a single supratemporal on each side of the midline; (9) circumorbital ring incomplete; (10) an interopercular; (11) un- cinate process on third infrapharyngobranchial; (12) a rectus communis muscle; (13) medial neural spines in caudal region; (14) basipter- ygoid process entirely composed of parasphenoid; (15) a postnares commissure between the supra- and infraorbital canals; (16) no clavicle; (17) uncinate processes on first and second infrapharyngobranchials; (18) infrapharyngobranchials laterally supported; (19) differentiated dorsal gill arch musculature (i.e. presence of OD, TD, etc.); (20) four basibranchial copulae; (21) course of spinal nerves; (22) general similarities in pectoral girdle anatomy ........................ 268 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure Page 18. Alternate hypotheses of the phylogenetic position of the semionotiform genus Lepidotes. (a.) Least rejected hypothesis, (b.) alternate hypo- thesis with corroborating characters. Synapomorphies (black rectangles) connecting taxa are: (1) medial neural spines in caudal region; (2) a posterior myodome; (3) a supra- maxilla; (4) an interopercular; (5) a maxilla with internal articulatory head; (6) loss of opisthotic; (7)
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