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Smithsonian Institution Scholarly Press smithsonian contributions to zoology • number 627 Smithsonian Institution Scholarly Press TheA Chronology Therian Skull of MiddleA Missouri Lexicon with Plains EmphasisVillage on the OdontocetesSites J. G. Mead and R. E. Fordyce By Craig M. Johnson with contributions by Stanley A. Ahler, Herbert Haas, and Georges Bonani SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of “diffusing knowledge” was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.” This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions in History and Technology Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Museum Conservation Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology In these series, the Institution publishes small papers and full-scale monographs that report on the research and collections of its various museums and bureaus. The Smithsonian Contributions Series are distributed via mailing lists to libraries, universities, and similar institu- tions throughout the world. Manuscripts submitted for series publication are received by the Smithsonian Institution Scholarly Press from authors with direct affilia- tion with the various Smithsonian museums or bureaus and are subject to peer review and review for compliance with manuscript preparation guidelines. General requirements for manuscript preparation are on the inside back cover of printed volumes. For detailed submissions require- ments and to review the “Manuscript Preparation and Style Guide for Authors,” visit the Submissions page at www.scholarlypress.si.edu. smithsonian contributions to zoology • number 627 The Therian Skull A Lexicon with Emphasis on the Odontocetes J. G. Mead and R. E. Fordyce WASHINGTON D.C. 2009 ABSTRACT Mead, J. G., and R. E. Fordyce. The Therian Skull: A Lexicon with Emphasis on the Odontoce- tes. Smithsonian Contributions to Zoology, number 627, 248 pages, frontispiece, 2 diagrams, 32 figures, 9 tables, 2009. — Cetaceans form one of the most unique groups in the evolutionary history of mammals. They have returned to the sea and modified their tail as an efficient means of locomotion. As they adapted to the limited visibility in the aquatic environment, the odontocetes developed a system of echolocation that resulted in extensive modifications to the skull bones. This made descriptive comparisons very difficult and early anatomists unwittingly composed new terms for anatomical structures that had already been named in other taxa. This made anatomical comparisons, based on the literature, extremely tenuous. This lexicon is an attempt to remedy that situation in that it provides headwords and defini- tions for all the terms that have been used in describing the mammal skull and notes the synony- mous terms. The lexicon includes the human nomenclature (Nomina Anatomica), the veterinary nomenclature (Nomina Anatomica Veterinaria) and the nomenclature that is used in descriptive comparative anatomy. The lexicon covers not only extant but extinct mammalian groups and is extensively indexed. Cover: Detail from Frontispiece, illustration of a dolphin skull (cf. Delphinus sp.) presented by Gesner (1560). Published by Smithsonian Institution Scholarly Press P.O. Box 37012, MRC 957 Washington, D.C. 20013-7012 www.scholarlypress.si.edu Library of Congress Cataloging-in-Publication Data Mead, James G. The therian skull : a lexicon with emphasis on the odontocetes / J. G. Mead and R. E. Fordyce. p. cm. — (Smithsonian contributions to zoology, ISSN 0081-0282 ; no. 627) Includes bibliographical references and index. 1. Skull—Dictionaries. 2. Mammals—Anatomy. 3. Cetacea—Anatomy. 4. Anatomy, Comparative. I. Fordyce, R. Ewan. II. Title. QL822M43 2009 571.3'195—dc22 2007045891 ∞ The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48–1992. Dedication The authors dedicate this work to the memory of Dr. John E. Heyning (6 January 1957–17 February 2007), who was a colleague in anatomy, a fellow museum curator, and a valued friend. FRONTISPIECE. Dolphin skull (cf. Delphinus spp.) with the caption “Delphini Calvaria è libro Bellonij” presented by Gesner (1560:162). Contents LIST OF ILLUSTRATIONS vii LIST OF TABLES ix INTRODUCTION 1 History 1 Evolutionary Changes 2 MATERIALS AND METHODS 3 Protocol for Terms in the Lexicon 3 Priority 4 Embryological Terms 4 Orientation Terms and Terms that Imply Size, Shape, and Proportion 4 Eponymous Terms 5 Usage of Terms 5 Absence of Landmarks 5 Illustrations 5 Separate Treatment of Bones Associated with the Ear 5 Craniometric Landmarks 5 Specimens Used 5 OSTEOLOGICAL RELATIONSHIPS OF THE SKULL BONES 6 Types of Features 6 Topographic Relationships of the Bones 6 Cranial Nerves 6 GUIDE TO LEXICON ENTRIES 10 LEXICON 11 Basioccipital Bone 11 Ethmoid Bone 13 Exoccipital Bone 17 Frontal Bone 24 Hyoid Apparatus 32 Interparietal Bone 33 Jugal Bone 35 Lacrimal Bone 38 Mandible 42 v i • smithsonian contributions to zoolog Y Maxilla 48 Nasal Bone 68 Palatine Bone 70 Parietal Bone 76 Premaxilla 79 Pterygoid Bone 82 Septomaxilla 90 Sphenoid Bone 90 Squamosal Bone 102 Supraoccipital Bone 108 Tympanoperiotic Complex 111 Periotic Bone 111 Tympanic Bulla 133 Auditory Ossicles 149 Malleus 149 Incus 151 Stapes 151 Vomer 151 MULTI-ELEMENT COMPLEXES: STRUCTURES SHARED BY ADJACENT BONES 154 Topographic Features 154 The Rostrum 154 Antorbital Region 154 Facial Region 154 The Palate and Pterygopalatine Fossa 154 The Cranium 154 Vertex 155 Occipital Area 155 Zygomatic Arch and Temporal Fossa 155 Orbit and Orbital Fissure 155 Basicranium 155 Tympanoperiotic 155 Cranial Hiatus 155 Pterygoid Sinus Complex 156 Venous Sinus System 156 Mandible and Mandibular Foramen 156 Transmission of Vessels, Nerves, and Passages 156 Infraorbital Complex 156 Nasal Passages 156 Nares 156 Nasal Cavity 157 Basipharyngeal Canal 157 Paranasal Sinuses 157 Arterial Circulation and Basicranial Foramina 157 FIGURES 159 ACKNOWLEDGMENTS 201 APPENDIX A. Historical TerminologY of the Cranial Nerves 203 APPENDIX B. Skull Measurements for Delphinids 205 REFERENCES 207 INDEX 217 Illustrations Cover. Detail from Frontispiece. Frontispiece. Dolphin skull (cf. Delphinus spp.) presented by Gesner (1560). iv Diagram 1. Schematic cross section of the rostrum and antorbital fossa. 50 Diagram 2. Schematic dorsal view around the antorbital notch in Odontoceti. 51 Figure 1. Intact skull of adult specimen, dorsal view. 160 Figure 2. Intact skull of adult specimen, ventral view. 161 Figure 3. Intact skull of adult specimen, lateral view. 162 Figure 4. Intact skull of adult specimen, anterior view. 162 Figure 5. Intact skull of adult specimen, posterior view. 163 Figure 6a–f. Intact neonatal skull. 164 Figure 6g–i. Disarticulated neonatal skull (g) and anterodorsal views of young (h) and adult (i) specimens. 165 Figure 7. Disarticulated neonatal skull, basioccipital. 166 Figure 8. Disarticulated neonatal skull, ethmoid. 167 Figure 9. Disarticulated neonatal skull, exoccipital. 168 Figure 10. Disarticulated neonatal skull, frontal. 169 Figure 11. Disarticulated neonatal skull, hyoid. 170 Figure 12. Disarticulated neonatal skull, interparietal. 171 Figure 13. Disarticulated neonatal skull, jugal. 172 Figure 14. Disarticulated neonatal skull, lacrimal. 173 Figure 15. Disarticulated neonatal skull, mandible. 174 Figure 16. Disarticulated neonatal skull, maxilla. 175 Figure 17. Disarticulated neonatal skull, nasal. 176 Figure 18. Disarticulated neonatal skull, palatine. 177 Figure 19. Disarticulated neonatal skull, parietal. 178 Figure 20. Disarticulated neonatal skull, premaxilla. 179 Figure 21. Disarticulated neonatal skull, pterygoid. 180 Figure 22. Disarticulated neonatal skull, sphenoid. 181 Figure 23. Disarticulated neonatal skull, squamosal. 182 Figure 24. Disarticulated neonatal skull, supraoccipital. 183 v i i i • smithsonian contributions to zoolog Y Figure 25a–c. Disarticulated neonatal skull, tympanoperiotic. 184 Figure 25d,e. Disarticulated neonatal skull, tympanic, right. 185 Figure 25f–k. Adult skull, tympanic, right. 186 Figure 25l–q. Adult skull, periotic, right. 187 Figure 25r,s. Adult skull, tympanic and malleus, left. 188 Figure 25t–v. Adult skull, periotic, left. 189 Figure 25w,x. Adult skull, sectioned tympanoperiotic, right. 190 Figure 25y. Disarticulated neonatal skull, tympanoperiotic, left. 191 Figure 26a–f. Skull of juvenile specimen, malleus, right. 192 Figure 26g–l. Skull of juvenile specimen, incus, right. 193 Figure 26m–r. Skull of juvenile specimen, stapes, right. 194 Figure 27. Disarticulated neonatal skull, vomer. 195 Figure 28. Disarticulated neonatal skull, basicranium, right, dorsal view. 196 Figure 29. Disarticulated neonatal skull, bulla, right, ventrolateral view. 197 Figure 30. Skull of adult specimen, basicranium, right, ventrolateral view. 198 Figure 31. Skull