SV Jurassic Rhynchonellids Internal Structures and Taxonomic Revisions 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, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 73 Jurassic Rhynchonellids: Internal Structures and Taxonomic Revisions Shi, Xiao-ying and Richard E. Grant SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1993 ABSTRACT Shi, Xiao-ying and Richard E. Grant. Jurassic Rhynchonellids: Internal Structures and Taxonomic Revisions. Smithsonian Contributions to Paleobiology, number 73, 190 pages, 83 figures, 18 plates, 1993.—Jurassic brachiopods of the order Rhynchonellida are classified according to modem concepts and techniques, with special attention to internal structures. They are grouped into 6 families and 16 subfamilies of which three are new: the Acanthorhynchiinae, the Cryptorhynchiinae, and the Piarorhynchiinae. Subfamilies emended or revised are the Acanthothyridinae Schuchert (1913) raised to family rank, Davanirhynchinae Ovtsharenko (1983), Dzhangirhynchinae Ovtsharenko (1983), Erymnariinae Cooper (1959), Indorhynchiinae Ovtsharenko (1975), Septocrurellinae Ager, Childs, and Pearson (1972), and Striirhynchiinae Kamyshan (1968). New genera are Aalenirhynchia (type-species Rhynchonella subdecorata Davidson, 1853), Bradfordirhynchia (type-species Cryptorhynchia bradfordensis Buckman, 1918), and Sharpirhynchia (type-species Kallirhynchia sharpi Muir-Wood, 1938). A new subgenus is Burmirhynchia (Hopkinsirhynchia) (type-species Burmirhynchia hopkinsi Da­ vidson, 1854). The only new species is Pycnoria depressa. Eleven genera are revised, and many are transferred among the subfamilies; lectotypes are designated where needed. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The trilobite Phacops rana Green. Library of Congress Cataloging-in-Publication Data Shi, Xiao-ying Jurassic Rhynchonellids: internal structures and taxonomic revisions / Shi, Xiao-ying and Richard E. Grant. p. cm. — (Smithsonian contributions to paleobiology ; no. 73) Includes bibliographical references (p. ) and index. 1. Rhynchonellida, Fossil. 2. Paleontology-Jurassic. I. Grant, Richard E. II. Title. III. Series. QE701.S56 no. 73 [QE797.R5] 560 s-dc20 [564'.8] 93-23896 ® 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—1984. Contents Page Introduction 1 Previous Work 2 Acknowledgments 3 Internal Structures 3 Terminology 4 Crura 4 Septalium 14 Classification 17 Rhynchonellid Hierarchical Synopsis 17 Characteristics of Rhynchonellid Suprageneric Taxa 19 Superfamily RHYNCHONELLACEA Gray, 1848 19 Family WELLERELLIDAE Likharev, 1956 19 Subfamily CIRPINAE Ager, 1965 19 Family BASILIOLIDAE Cooper, 1959 19 Subfamily DAVANIRHYNCHIINAE Ovtsharenko, 1983 19 Subfamily DZHANGIRHYNCHIINAE Ovtsharenko, 1983 20 Subfamily LACUNOSELLINAE Smirnova, 1936 20 Family RHYNCHONELLIDAE Gray, 1848 20 Subfamily RHYNCHONELLINAE Gray, 1848 20 Subfamily PlARORHYNCHllNAE, new subfamily 21 Subfamily IVANOVIELLINAE Makridin, 1964 21 Subfamily TETRARHYNCHIINAE Ager, 1965 22 Subfamily INDORHYNCHUNAE Ovtsharenko, 1975 22 Subfamily CYCLOTHYRIDINAE Makridin, 1955 23 Subfamily STRIIRHYNCHIINAE Kamyshan, 1968 23 Family ACANTHOTHYRIDIDAE Schuchert, 1913, new status 23 Subfamily ACANTHOTHYRIDINAE Schuchert, 1913 24 Subfamily ACANTHORHYNCHIINAE, new subfamily 24 Family ERYMNARIIDAE Cooper, 1959 24 Subfamily ERYMNARIINAE Cooper, 1959 24 Subfamily CRYPTORHYNCHIINAE, new subfamily 24 Subfamily SEPTOCRURELLINAE Ager, Childs, and Pearson, 1972 25 Family SEPTIRHYNCHIIDAE Muir-Wood and Cooper, 1951 25 Systematic Descriptions of Selected Rhynchonellid Genera and Species 25 Prionorhynchia Buckman, 1918 25 Bradfordirhynchia, new genus 26 Ptyctorhynchia Buckman, 1918 32 Costirhynchia Buckman, 1918 36 Rhynchonelloidea Buckman, 1918 36 Lotharingella Laurin, 1984 41 Rhynchonelloidella Muir-Wood, 1936 43 Sharpirhynchia, new genus 49 Thurmannella Leidhold, 1921 53 Aalenirhynchia, new genus 57 Burmirhynchia Buckman, 1918 60 Hopkinsirhynchia, new subgenus 63 in 1V SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Cymatorhynchia Buckman, 1918 71 Daghanirhynchia Muir-Wood, 1935 73 Goniorhynchia Buckman, 1918 75 Isjuminella Makridin, 1955 78 Kallirhynchia Buckman, 1918 80 Kutchirhynchia Buckman, 1918 86 Lacunaerhynchia Almeras, 1966 88 Obsoletirhynchia Shi, 1992 90 Rhactorhynchia Buckman, 1918 98 Somalirhynchia Weir, 1925 104 Sphenorhynchia Buckman, 1918 105 Moquellina Jin, Sun, and Ye, 1979 106 Tanggularella Shi, 1990 107 Bicepsirhynchia Shi, 1990 113 Flabellirhynchia Buckman, 1918 115 Globirhynchia Buckman, 1918 116 Granulirhynchia Buckman, 1918 121 Pararhactorhynchia Shi, 1990 123 Pycnoria Cooper, 1989 125 Pycnoria depressa, new species 127 Septaliphoria Leidhold, 1921 132 Acanthothiris d'Orbigny, 1850 136 Aethirhynchia Shi, 1990 138 Cryptorhynchia Buckman, 1918 139 References 143 Plates 151 Index 188 Jurassic Rhynchonellids: Internal Structures and Taxonomic Revisions Shi, Xiao-ying and Richard E. Grant Introduction rhynchonellids, we realized that most of the genera are ordinary looking forms, they are similar in external morphology and not The order Rhynchonellida, one of the major groups among easy to distinguish. It is almost impossible to put some species the Jurassic brachiopods, presents challenges of long standing. into their relevant genera confidently without studying the Although it has been more than two hundred years since internal structures. In the "Treatise" (Williams and Rowell, Linnaeus (1767) recorded the first species of Jurassic rhyn­ 1965) a tentative classification was suggested for the Mesozoic chonellid, and more than 160 Jurassic rhynchonellid genera and Cenozoic Rhynchonellida and many genera were presented have been named, the classification has not been established with some interior information; however, a lot of uncertainty solidly and many genera are still not properly studied in the remained and many genera that were not sufficiently known terms of internal structures. Jurassic rhynchonellids are internally had to be grouped solely on their external appear­ important not only in stratigraphical and paleontological ances. Because of insufficient study of the internal structures, studies, or for their abundance and diversity in the marine much confusion in the taxonomy has arisen, and homonyms sediments, but also for their widespread distribution and rapid and synonyms are not rare in the literature. Comprehensive evolution and their ecological adaptation to different environ­ study of Jurassic rhynchonellids reveals that species or generic ments. names vary from country to country, or even from area to area. The basic systematics of the Jurassic rhynchonellids were These problems in taxonomy, in turn, produce new difficulties laid out by Buckman (1918). As a result of the limitation in and confusion in the studies of biostratigraphy, paleobiogeo- techniques, equipment, and knowledge at that time, Buckman's graphy, and evolution. Therefore, as the basis of paleontology system is far from satisfactory in the view of present and its related subjects, taxonomic study should still be knowledge. Most of his genera were established without emphasized and the study of interiors, which is of great knowledge of internal characters, and many of them were not importance in the classification, should receive especially close defined clearly on any consistent feature, so that much attention. confusion has resulted. Buckman did, of course, contribute a Here we attempt to revise the Jurassic rhynchonellid genera, great deal to the systematics, and his monograph is still used as