HARVARD UNIVERSITY Library of the Museum of Comparative Zoology ins of , mricayf UUontowasJ 'AM '? /*??/ VOLUME 99, NUMBER 335 DECEMBER 12, 1990 HARVARD UNIVERSITY Review of the Bullia Group (Gastropoda: Nassariidae) with comments on its Evolution, Biogeography, and Phylogeny by Warren D. Allmon Paleontological Research Institution 1259 Trumansburg Road Ithaca, New Yoik, 14850 U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION Officers President Harry A. Leftingwell Vice-President J. Thomas Dutro. Jr. Secretary Henry W. Theisen Treasurer James C. Showacre Assistant Treasurer Roger J. Howley Director Peter R. Hoover Legal Counsel Henry W. Theisen Trustees Bruce M. Bell (to 6/30/93) Edward B. Picou, Jr. (to 6/30/92) Carlton E. Brett (to 6/30/92) James C. Showacre (to 6/30/93) J. Thomas Dutro, Jr. (to 6/30/93) James E. Sorauf (to 6/30/91) Harry A. Leffingwell (to 6/30/93) John Steinmetz (to 6/30/91) Robert M. Linsley (to 6/30/92) Henry W. Theisen (to 6/30/92) Cathryn Newton (to 6/30/91) Raymond Van Houtte (to 6/30/91) Samuel T. Pees (to 6/30/92) William P. S. Ventress (to 6/30/93) A. D. Warren, Jr. (to 6/30/91) BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA Peter R. Hoover Editor Reviewers for this issue David R. Lindberg Elizabeth Nesbitt A list of titles in both series, and available numbers and volumes may be had on request. Volumes 1-23 of Bulletins of American Paleontology have been reprinted by Kraus Reprint Corporation, Route 100, Millwood, New York 10546 USA. Volume I of Palaeontographica Americana has been reprinted by Johnson Reprint Corporation. 1 1 1 Fifth Ave., New York, NY 10003 USA. Subscriptions to Bulletins of American Paleontology may be started at any time, by volume or year. Current price is US $30.00 per volume. Numbers of Palaeontographica Americana are priced individually, and arc invoiced separately on request. for additional information, write or cal Paleontological Research Institution 1259 Trumansburg Road Ithaca, NY 14850 USA (607) 273-6623 The Paleontological Research Institution acknowledges with special thanks the contributions of the following individuals and institutions PATRONS ($1000 or more at the discretion of the contributor) James E. Allen (1967) Robert C. Hoerle (1974-1977) American Oil Company (1976) Richard I. Johnson (1967, 1986) Atlantic Richfield Company (1978) J. M. McDonald Foundation (1972,1978) Christina L. Balk (1970. 1982, 1983) Mobil Oil Corporation (1977 to date) Hans M. Bolli(1984) Samuel T. Pees (1981) RuthG. Browne (1986) Richard E. Petit (1983) Mr. & Mrs. Kenneth E. Caster (1967) Robert A. Pohowsky (1982) Chevron Oil Company (1978, 1982) Texaco, Inc. (1978, 1982, 1987) Exxon Company (1977 to date) Union Oil of Californl\ (1982 to date) Lois S. Fogelsanger (1966) United States Steel Foundation (1976) Gulf Oil Corporation (1978) Charles G. Ventress (1983 to date) Merrill W. Haas (1975) Christine C. Wakeley (1976-1984) Norman E. Weisbord(1983) (continued overleaf) LIFE MEMBERS ($200) R. Tucker Abbott William F. Klose, II James E. Allen JiRj KrIz Elizabeth A. Balcells-Baldwin Ralph L. Langenheim, Jr. Christina L. Balk Harry A. Leffinowell Bruce M. Bell Egbert G. Leigh, Jr. Robert A. Black Gerard A. Lenhard Richard S. Boardman Louie N. Marincovich, Jr. Hans Bolli Donald R. Moore David John Bottjer Shuji Niko Ruth G. Browne Sakae O'Hara J. David Bukry Samuel T. Pees Sybil B. Burger Richard E. Petit Lyle D. Campbell Edward B. Picou, Jr. John L. Carter Robert A. Pohowsky Anneliese S. Caster John Pojeta, Jr. Kenneth E. Caster John K. Pope John E. DuPont Anthony Reso J. Thomas Dutro, Jr. Arthur W. Rocker J. Mark Erickson Walter E. Sage, III Richard J. Erickson John B. Saunders Lois S. Fogelsanger Judith Schiebout A. Eugene Fritsche Miriam W. Schriner Christopher L. Garvie Edward S. Slagle Ernest H. Gilmour David H. Stansbery Merrill W. Haas Jorge P. Valdes Anita G. Harris Charles G. Ventress Steven M. Herrjck WiLLL\M P. S. Ventress Robert C. Hoerle Emily H. Vokes F. D. Holland, Jr. Harold E. Vokes Frederick H. C. Hotchkiss Christine C. Wakeley David Jablonski Thomas R. Waller Richard I. Johnson Albert D. Warren. Jr. David B. Jones Gary D. Webster Peter Jung Norman E. Weisbord Tomoki Kase Ralph H. Willoughby David Garrett Kerr Armour C. Winslow Cecil H. Kindle Victor A. Zullo Membership dues, subscriptions, and contributions are all important sources of funding, and allow the Paleontological Research Institution to continue its existing programs and services. The P.R.I, publishes two series of respected paleontological monographs, Bullclins ofAmerican Paleonlology and Falaeonlofiraphica Americana, that give authors a relatively inexpensive outlet for the publication of significant longer manuscripts. In addition, it reprints rare but important older works from the pa- leontological literature. The P.R.I, headquarters in Ithaca, New York, houses a collection of inver- tebrate type and figured specimens, among the five largest in North America; an extensive collection of well-documented and curated fossil specimens that can form the basis for significant future pa- leonlologic research; and a comprehensive paleontological research library. The Paleontological Research Institution is a non-profit, non-private corporation, and contributions may be U.S. income lax deductible. For more information on P.R.I, programs, memberships, or subscriptions to P.R.I, publications, call or write; Peter R. Hoover Director Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850 U.S.A. 607-273-6623 -fJipicrican toicqy DECEMBER 12, 1990 OLUME 99, NUMBER 335 Review of the Bullia Group (Gastropoda: Nassariidae) with comments on its Evolution, Biogeography, and Phylogeny by Warren D. AUmon Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. Library of Congress Card Number: 90-63452 The publication costs of this volume were in large part defrayed by a generous grant of $15,000 from the EXXON Corporation. The Paleontological Research Institution gratefully acknowledges this rec- ognition and support of its publications program. Printed in the United States of America Allen Press, Inc. Uwrcnce, KS 66044 U.S.A. CONTENTS Page Abstract 5 Acknowledgements 5 Introduction Study Justification 6 Systematic Zoology and Paleontology; Introductory Remarks 8 Institutional Abbreviations 9 Familial Classification 9 The Bullia Group and Melanopsids 12 Recent Representatives Ecology 12 Previous Taxonomic Work 13 Systematic Characters in Recent Species 13 Systematic Summary 19 Fossil Representatives Miocene of the Middle Atlantic Coastal Plain Introduction 29 Previous Work 30 Present Study Stratigraphic Context 31 Methods 32 Results 37 Discussion 39 Form of the Shell Apex 46 Bulliopsis from New Jersey 50 Systematic Summary 50 Cretaceous of the southeastern U. S 52 Paleogene of the Gulf Coastal Plain 53 Cenozoic of the West Coast of North America 63 Cenozoic of the Caribbean, Central and northern South America 71 Cenozoic of southern South America and Antarctica 79 Cenozoic of Europe Genera of Nuttall and Cooper 82 Problematic Continental Forms 83 Neogene of Africa 86 Neogene of the Indian Subcontinent 89 Phylogeny, Biogeography, and Evolution Phylogenetic Analysis and Paleontology 90 Completeness of the Fossil Record 90 Characters and Character Analysis 97 Polarity 99 Ancestor-Descendant Series 1 00 Phylogenetic Analysis of the Bullia Group Method 100 Outgroup Comparison 102 Morphological Analysis 103 Paleobiogeographic Framework 109 Discussion and Conclusions 1 1 Systematic Summary 115 Appendix: Paleobiogeographic History of the Atlantic 116 References Cited 119 Plates 140 Index 155 . LIST OF ILLUSTRATIONS Text-figure Page 1 Proposed phylogenetic tree for the evolution of the family Nassariidae 7 2. Sketches of species of Buccinidae s. I. 10 3. Radular dentitions of living species in the Bullia group 15 4. Opercular outlines of living sp>ecies of the Bullia group 16 5. Generalized stratigraphic classification for Miocene and early Pliocene sediments of the Middle Atlantic coastal plain 31 6. Map of the Chesapeake Bay region 33 7. Map of the Middle Atlantic coastal plain 35 8. Alternative scenarios for the evolution of Miocene species of Bidliopsis 36 9. Morphometric variables recorded for specimens of Bulliopsis 37 10. Scatterplot of scores on the first two factor axes for all specimens of Bulliopsis collected from the Chesapeake Bay region for this study 40 1 1. Scatterplot of scores on the second and third factor axes for all specimens of Bulliopsis collected from the Chesapeake Bay region fiar this study 41 12. Scatterplot of scores on the first two discriminant axes for all specimens of Bulliopsis collected from the Chesapeake Bay region for this study 42 1 3. Proportional representation of the four Bulliopsis morphotypes at each of the four stratigraphic levels sampled in the Chesapeake Bay region 43 14. The four "select groups" of specimens chosen to represent the four Bulliopsis morphotypes 45 1 5. Representative samples of Bulliopsis from four stratigraphic levels in the upper Miocene of Maryland and Virginia 47 16. Scatterplot of scores on the first two factor axes for specimens of Bulliopsis Integra and Bulliopsis quadrala from Windmill Point. St. Mary's Co., Maryland 48 17. Plots of mean values of the two calculated shouldering indices for all specimens of Bulliopsis quadrala from the four stratigraphic horizons sampled in the Chesapeake Bay region 49 1 8. Generalized correlation chart for Paleogene sediments of the Gulf coastal plain 54 1 9. Summary phylogenetic tree for all species of Bullia (Bulliopsis) 56 20. Map of southern Africa, showing localities where fossil specimens of Bullia have been found 89 21. Approximate stratigraphic ranges of the species of the Bullia group that have a known fossil record foldout inside back cover 22. Histogram showing the number of species of the Bullia group from the Upper Cretaceous through the Recent 91 23. Histogram showing the stratigraphic and geographic distribution of published fossil faunas examined in this study 92 24.