MITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY NUMBER 76 33 m£$vy Brachiopods Near the Permian-Triassic Boundary in South China 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 76 Brachiopods Near the Permian-Triassic Boundary in South China Guirong Xu and Richard E. Grant SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1994 ABSTRACT Xu, Guirong, and Richard E. Grant. Brachiopods Near the Permian-Triassic Boundary in South China. Smithsonian Contributions to Paleobiology, number 76, 68 pages, 54 figures, 7 tables, 1994.—Sixty-eight genera and 164 species in the Changxingian Stage and 12 genera and 20 species in the lower Griesbachian Stage are recorded on the basis of brachiopod fossils collected from 32 sections in South China and from review of the Chinese literature. Of these, 24 genera and 34 species are described here, including three new genera ( Fanichonetes, Prelissorhynchia, and Rectambitus) and 24 new species ( Acosarina strophiria, Enteletes asymmatrosis, Peltichia schizoloides, Derbyia pannuciella, Perigeyerella altilosina, Choneti- nella cursothonia, C. volitanliopsis, Fanichonetes campigia, Cathaysia spiriferoides, Uncin- unellina multicostifera, Prelissorhynchia triplicatioid, Cyrolexis antearcus, Cyrolexis beccojec- tus, Cartorhium xikouensis, C. twifurcifer, Callispirina rotundella, Araxathyris subpentagulata, A. beipeiensis, Spirigerella discsella, S. ovaloides, Squamulariaformilla, Hustedia orbicostata, Rostranteris ptychiventria, and Notothyris bifoldes). The Cathaysia chonetoides-Chonetinella substrophomenoides assemblage zone and the Cathaysia sinuata-Waagenites barusiensis assemblage zone represent respectively faunas of the lower Changxingian and the upper Changxingian in clastic lithofacies; whereas the Peltichia zigzag-Prelissorhynchia triplicatioid assemblage zone and the Spirigerella discusella- Acosarina minuta assemblage zone represent faunas in limestone lithofacies. The Crurithyris pusilla-Lingula subcircularis assemblage zone and Permian-type brachiopods are present in the lower Griesbachian. The Changxingian brachiopod fauna can be correlated with the Dorashamian fauna of Armenia; the brachiopod faunas of the Ali Bashi Formation, North-West Iran; unit 7 of the Hambast Formation, Central Iran; and the upper part of the Bellerophon Formation of the Southern Alps. The genera Cathaysia, Peltichia, and Prelissorhynchia are especially characteristic of the Cathaysia Tethyan Subprovince. In contrast, the WestTethyan Subprovince is characterized by the genera Costiferina, Ombonia, Comelicania, and many other species. Four brachiopod ecofacies are recognized in the Changxingian of South China: (1) antibiohermal dwellers; (2) calcareous substratum dwellers; (3) biohermal dwellers; and (4) ubiquitous substrate dwellers. In the lower Griesbachian, the brachiopod fauna of Lingula and Crurithyris spreads across the entire Tethys and is called the Circum-Pangaea brachiopod fauna. Massive extinction of brachiopod faunas occurred at the close of the Changxingian, with only a few Permian-types surviving into the early Griesbachian, and they completely vanished after the early Griesbachian except for harbingers of Mesozoic brachiopods. 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 Xu, Guirong Brachiopods near the Permian-Triassic boundary in South China / Guirong Xu and Richard E. Grant, p. cm. — (Smithsonian contributions to paleobiology ; no. 76) Includes bibliographical references. 1. Brachiopoda, Fossil—China. 2. Paleontology—Triassic. 3. Paleontology—Permian. I. Grant, Richard E. II. Title. III. Series. QE701.S56 no. 76 [QE796] 560 s-dc20 [564'.8'0951] 93-11614 ® 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 Acknowledgments.3 Zonation and Correlation.3 Relationship with Other Biozones.6 Correlations within Tethys.8 Brachiopod Provinces and Biofacies.12 Biofacies.12 Changxingian Brachiopod Biofacies.12 Brachiopod Biofacies of the Lower Griesbachian.13 Extinction and Evolution.13 Conclusions.16 Systematic Listing.17 Order Orth ida Schuchert and Cooper, 1932 . 18 Superfamily Rhipidomelloidea Schuchert, 1913.18 Family SCHIZOPHORIIDAE Schuchert and LeVene, 1929 . 18 Genus Acosarina Cooper and Grant, 1969 . 18 Acosarina strophiria sp. nov.18 Superfamily Enteletoidea Waagen, 1884 . 19 Family Enteletidae Waagen, 1884 . 19 Genus Enteletes Fischer de Waldheim, 1825 .19 Enteletes asymmatrosis sp. nov.19 Genus Peltichia Jing and Liao, 1981.20 Peltichia schizoloides sp. nov.21 Order Strophomenida Opik, 1934 . 24 Suborder Orthotetidina Waagen, 1884 . 24 Superfamily Derbyioidea Stehli, 1954 . 24 Family Derbyidae Stehli, 1954 . 24 Subfamily Derbyinae Stehli, 1954 . 24 Genus Derbyia Waagen, 1884 . 24 Derbyia pannuciella sp. nov.24 Family Meekellidae Stehli, 1954 . 24 Genus Meekella White and St. John, 1867 . 24 Meekella langdaiensis Liao, 1980 . 24 Superfamily Orthotetoidea Waagen, 1884 . 26 Family ORTHOTET1DAE Waagen, 1884 . 26 Genus Perigeyerella Wang, 1955 . 26 Perigeyerella altilosina sp. nov.26 Suborder Chonetidina Muir-Wood, 1955 . 27 Superfamily CHONETOIDEA Bronn, 1862 . 27 Family Rugosochonetidae Muir-Wood, 1962 . 27 Subfamily CHONETINELUNAE Muir-Wood, 1962 . 27 Genus Chonetinella Ramsbottom, 1952 . 27 C hone tine l la cursothornia sp. nov.27 Chonetinella volitanliopsis (Xu, 1987). 28 Subfamily RUGOSOCHONETINAE Muir-Wood, 1962 . 29 Fanichonetes gen. nov.29 iii IV SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Fanichonetes campigia sp. nov.29 Suborder Productidina Waagen, 1883 . 29 Superfamily PRODUCTOIDEA Gray, 1840 . 29 Family Chonetellidae Likharev, 1960 . 29 Genus Cathaysia Ching, 1966. 29 Cathaysia sinuata Chan, 1979 . 31 Cathaysia spiriferoides sp. nov.32 Cathaysia obicularis Liao, 1980 . 34 Superfamily RlCHTHOFENlOlDEA Waagen, 1885 . 34 Richthofeniid, genus and species undetermined.34 Order Rhynchonellida Kuhn, 1949 . 35 Superfamily Wellerelloidea Xu and Liu, 1983 . 35 Family Wellerellidae Likharev, 1956 . 35 Genus Uncinunellina Grabau, 1932 . 35 Uncinunellina multicostifera sp. nov.35 Family PONTISHDAE Cooper and Grant, 1976 . 36 Subfamily LlSSORHYNCHUNAE Xu and Liu, 1983 . 36 Prelissorhynchia gen. nov.36 Prelissorhynchia pseudoutah (Huang, 1933), new combination .... 38 Prelissorhynchia triplicatioid sp. nov.38 Superfamily Stenoscismatoidea Oehlert, 1887 (1883). 39 Family Atriboniidae Grant, 1965 . 39 Subfamily Atriboniinae Grant, 1965 . 39 Genus Cyrolexis Grant, 1965 . 39 Cyrolexis ante arcus sp. nov.39 Cyrolexis beccojectus sp. nov.39 Order Spiriferida Waagen, 1883 . 41 Suborder Spiriferidina Waagen, 1883 . 41 Superfamily Spiriferoidea King, 1846 . 41 Family Spiriferidae King, 1846 . 41 Subfamily Neospiriferinae Waterhouse, 1968 . 41 Genus Cartorhium Cooper and Grant, 1976 . 41 Cartorhium xikouensis sp. nov.41 Cartorhium twifurcifer sp. nov.42 Superfamily CYRTIOIDEA Frederiks, 1924 . 43 Family Ambocoeliidae George, 1931.43 Genus Crurithyris George, 1931.43 Crurithyris pusilla Chan, 1979 . 43 Suborder Spiriferinidina Cooper and Grant, 1976 . 45 Superfamily Spiriferinoidea Davidson, 1884 . 45 Family Paraspiriferinidae Cooper and Grant, 1976 . 45 Genus Paraspiriferina Reed, 1944 . 45 Paraspiriferina alpha (Huang, 1933). 45 Genus Callispirina Cooper and Muir-Wood, 1951.47 Callispirina'l rotundella sp. nov.47 Suborder ATHYRIDIDINA Boucot, Johnson, and Staton, 1964 . 48 Superfamily Athyrididoidea McCoy, 1844 . 48 Family Athyrididae McCoy, 1844 . 48 Subfamily SPIRIGERELL1NAE Grunt, 1965 . 48 Genus Araxathyris Grunt, 1965 . 48 Araxathyris beipeiensis sp. nov.48 Araxathyris subpentangulata sp. nov.49 Rectambitus gen. nov.50 Rectambitus bisulcatus (Liao, 1980), new combination.50 NUMBER 76 V Genus Spirigerella Waagen, 1883 . 51 Spirigerella discusella sp. nov.51 Spirigerella guizhouensis (Liao, 1980), new combination.52 Spirigerella ovaloides sp. nov.54 Spirigerella shuizhutangensis (Chan, 1979), new combination.56 Genus Tongzithyris Ching, Liao, and Fang, 1974 . 56 Tongzithyris sichuanensis Xu, 1987 . 57 Superfamily Reticularioidea Waagen, 1883 . 58 Family ELYTHIDAE Frederiks, 1924 . 58 Genus Squamularia Gemmellaro, 1899 . 58 Squamularia formilla sp. nov.58 Suborder RETZIIDINA Boucot, Johnson, and Staton, 1964 . 60 Superfamily Retzioidea Waagen, 1883 . 60 Family Retziidae Waagen, 1883 . 60 Genus Hustedia Hall and Clarke, 1893 . 60 Hustedia orbicostata sp. nov.60 Order Terebratulida Waagen, 1883 . 61 Superfamily Cryptonelloidea Thomson, 1926 . 61 Family NOTOTHYRIDIDAE Likharev, 1960 . 61 Genus Rostranteris Gemmellaro, 1899 . 61 Rostranteris ptychiventria sp. nov.61 Genus Notothyris Waagen, 1882 . 63 Notothyris bifoldes sp. nov.63 Literature Cited.65 Brachiopods Near the Permian-Triassic Boundary in South China Guirong Xu and Richard E. Grant Introduction The study of brachiopods near the Permian-Triassic bound¬ ary provides fossil evidence useful in establishing and correlating the stratigraphic boundary stratotype of the Per¬ mian-Triassic. Such a study also enables one to reconstruct the environment where brachiopods lived and adapted during the passage from the Paleozoic to the Mesozoic and to elucidate the evolutionary relationship between Paleozoic and Mesozoic brachiopods. Former works that systematically described brachiopods from the Late Permian of South China are by Chao (1927, 1928, 1929) and Huang (1932, 1933). Investigations of coal deposits of the Late Permian and research on the Permian- Triassic boundary stratotype in South China in the last decade has provided continued impetus to this line of endeavor. Papers by Chan (in Hou et al., 1979), Liao (1979, 1980a, 1981), Liao and Meng (1986), and Xu (in Yang et al., 1987) reveal that brachiopods are abundant in rocks of Late Permian age in South China and that several Permian-type brachiopods extend into the lowermost Triassic. This paper deals with the Changxingian Stage (formerly spelled Changhsingian) and the lower Griesbachian Stage brachiopods in South China, based upon materials collected from 32 sites whose locations are shown in Figure 1. (Hereafter, all collection spots will be referred to by their spot number and will not include the reference to Figure 1.) The study area is situated mainly in the Yangtze platform, which was essentially surrounded by “old lands” (i.e., emergent at the time) during the Late Permian and Early Triassic. North of the area is the North China platform, which was completely above sea level and, at that time, was the source of erosional Guirong Xu, Paleontology Laboratory, China University of Geo¬ sciences, Wuhan, Hubei Province, 430074, China. Richard E. Grant, Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. sediments. The Cathaysia old landmass extended north¬ eastward from the North China platform. The Kangdian old landmass was to the west, and the Yun-Kai old land was located south of the platform. Therefore, during the Late Permian and Early Triassic, this area was a barrier sea basin. Another old landmass, called the Jiangnan old landmass, was located in the middle of the basin. The sedimentary characters to a certain extent relate to that very complicated landform (Figure 3). The Changxingian Stage with its abundantly preserved brachiopods has several sedimentary types in the Yangtze basin, largely of three types: limestone, silicalite (i.e., particu¬ late chert), or arenite. The Changxing Formation, type section located at Meishan, Changxing County, Zhejiang Province (spot 26), consists of gray to dark medium-bedded pure limestone, bituminous limestone and limy dolomite, and intercalated cherty limestone. The Changxing Formation is most widely distributed in Zhejiang, Sichuan, and North Guizhou, generally in the platform facies of the basin. Chert sediments are represented in the Dalong Formation, whose type locality is at Matan, Geshan County, Guangxi Province (near spot 15), where it is composed of gray or black thin-bedded silicalite, yellowish brown siliceous tuffite, and intercalated siliceous or tufaceous arenite. Distribution of the Dalong Formation follows closely the distribution of volcaniclastic rocks extending along a narrow belt in South Guizhou, Guangxi, and Hubei Provinces. On the east side of the Kangdian old landmass, the west side of Cathaysia old landmass, and the north side of the Yun-Kai old landmass are mostly arenites of the Xuanwei Formation (type locality is at Xuanwei, Yunnan Province) and the Yanshi Formation (type locality is at the Yanshi section, Longyan County, Fujian Province) (spot 31). The former consists of intercalated terrigenous sediments and the latter is marine arenite containing dark gray, yellowish green, and grayish yellow calcareous siltstone and silty mudstone, interbedded 1 2 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Figure 1.—Distribution of the Permian-Triassic boundary sections of South China. 1. Kucaoping section, Xixiang County, Shaanxi Province 2. Mingxuexia section, Guangyuan County, Sichuan Province 3. Xindianzi section, Guangyuan County, Sichuan Province 4. Shangsi section, Guangyuan County, Sichuan Province 5. Xinjiacao section, Guangan County, Sichuan Province 6. Huayingchan section, Linshui County. Sichuan Province 7. Yanjingxi section, Hechuan County, Sichuan Province 8. Liangfengya section, Zhingqing County, Sichuan Province 9. Banzhuyuan section, Nantong County, Sichuan Province 10. Liuchang section, Qingzhen County, Guizhou Province 11. Yingpanpo section, Guiyang County, Guizhou Province 12. Xiaochehe section, Guiyang County, Guizhou Province 13. Huopu section, Pan County, Guizhou Province 14. Longtoujiang section, Yishan County, Guanxi Province 15. Paoshui section, Laibin County, Guanxi Province 16. Penglaitan section, Laibin County, Guangxi Province 17. Shatian section, Huangshi County, Hubei Province 18. Guanyinshan section, Fuqi County, Hubei Province 19. Baimu section, Yiahun County, Jiangxi Province 20. Jueguangsi section, Laiyang County, Hunan Province 21. Matian section, Yongxi County, Hunan Province 22. Xiaoyuanchong section, Jiahe County, Hunan Province 23. Meitian section, Yizhang County. Hunan Province 24. Majiashan section, Chao County, Anhui Province 25. Yueshan section, Huaining County, Anhui Provonce 26. Meishan section, Changxing County, Zhejiang Province 27. Huangzhishan section, Wuxing County, Zhejiang Province 28. Minfa section, Guangfeng County, Jiangxi Province 29. Xijia section, Shangroa County, Jiangxi Province 30. Yading section, Shangping County, Fujian Province 31. Yanshi section, Longyan County, Fujian Province 32. Xikou section, Zhenan County, Shaanxi Province with fine sandstone and lenses of limestone. Lower Griesbachian strata, which bear poor brachiopod fossils, are petrographically more uniform than are the underlying Changxingian rocks. They can be divided into two kinds of rocks: limestone and arenite. The lower Daye and the lower Qinglong formations consist mainly of limestone and calcareous mudstone. They occur on the platform facies belt of the basin. The clastic-type strata are dispersed along the margins of old land provinces, such as the Xikou Formation, which is located on the west of Cathaysia, the Kayitou Formation, and the lower Feixianguan Formation, which occurs to the east of the Kangdian old landmass. The Yinkeng Formation is currently becoming known internationally be¬ cause its reference section is situated at the same location as the type section of the Changxingian Stage (spot 26). It contains many fossils indicative of the Griesbachian Stage. It overlies the Changxing Formation; therefore, it is a prime candidate for the Permian-Triassic boundary stratotype. It consists of grayish yellow-green calcareous mudstone intercalated with a few thin-and medium-bedded limestone stringers (Table 1). Brachiopods from the 32 sections that are recorded in the literature of South China consist of about 68 genera and 164 species in the Changxingian and about 12 genera and 20 species that extend into the lower Griesbachian (Table 6). Permian-type brachiopods are completely absent above the Griesbachian. Twenty-four genera and 34 species are described in this paper, including 3 new genera and 24 new species. NUMBER 76 3 Table 1.—Summary of formations and general lithology near the Permian-Triassic boundary in South China. Stratigraphic Units Zhejiang and Jiangsu Provinces Hubei, Hunan and Guangxi Provinces Sichuan Province Fujian Province West Guizhou and Yunnan Provinces Series Stage O c Yinkeng or lower Lower Daye Fm.: Lower Lower Xikou Kayitou Fm.: 0 limestone calcareous and grey mudy stone, > O _l 6 silt mudstone limestone mudstone c c Changxing Fm: Dalong Fm.: Changxing Yanshi Fm.: Upper Xuan- Cu 0 Fm.: wei Fm.: E 0) limestone and chert silicalite inter- calcareous k_ 0 c limestone calated lime- limestone siltstone siltstone Ol X stone and mud- intercalated and intercalated k— O) r~ stone siliceous mudstone coal beds 0 0 limestone Q. _C and CL — ) O calcareous mudstone Acknowledgments Guirong Xu was a senior postdoctoral fellow at the Smithsonian Institution and received support from the Institu¬ tion’s Office of Fellowships and Grants, 1985-86. He expresses special thanks to the Smithsonian Institution and his supervisor and coauthor Richard E. Grant. The Department of Paleobiology, National Museum of Natural History (NMNH), Smithsonian Institution, provided space, equipment, and its huge collections that gave him an opportunity to compare specimens collected from South China to the collections from other Tethyan areas. Xu was especially pleased to have the opportunity to visit the classic Permian area of West Texas, guided by B.R. Wardlaw of the United States Geological Survey (USGS), R.E. Grant (NMNH), and D.M. Rohr and S. Rudine of Sul Ross State University. G. Arthur Cooper and Norman D. Newell contributed through their useful discussions of stratigraphy and brachio- pods. Rex A. Doescher provided us the use of his database at the Smithsonian International Brachiopod Information Center, which helped to make the manuscript consistent. The typescript was reviewed for scientific content by Yang Zunyi, China University of Geosciences, Stanley S. Beus, Northern Arizona University, and Bruce R. Wardlaw, United States Geological Survey, whom we thank for substantial insights. The collections of brachiopods described in this paper from 32 sections of South China were contributed by teachers and students of the Wuhan College of Geology who worked in the field every summer from 1980 to 1984. Data on stratigraphic sections were published by Z.Y. Yang et al. (1987). Other fossils referred to in this paper include ammonoids identified by F.Q. Yang, conodonts by M.H. Ding, and fusulinids by R.G. Gu. We appreciate their help greatly. Illustrated specimens are deposited in the collections of the former United States National Museum (USNM), now in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Zonation and Correlation Several schemes of brachiopod zonations near the Permian- Triassic boundary in South China have been advanced during the past two decades (Table 2). Summing up the brachiopod sequences of the 32 sections as well as information taken from other works, we put forward the brachiopod zonation presented in the left column in Table 2. Brachiopod faunas are closely related to lithofacies in the Changxingian, so brachiopod assemblage zones are linked to the several lithofacies of that stage. 1. Cathaysia chonetoides ( ChdiO)-C honetinella substro- phomenoides (Huang) assemblage zone (abbreviation C- C): The representative brachiopod fauna in the clastic lithofacies of the lower Changxingian is not highly diverse, being composed mainly of Chonetoidea, such as Waagenites, Chonetinella , and Fanichonetes, new genus; Chonetellidae, such as Cathaysia; and Meekellidae, such as Orthothetina and Perigeyerella. The following species play important roles in this assemblage zone: Cathaysia chonetoides (Chao), C. 4 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Table 2.—The course of zonation change of brachiopods near the Permian-Triassicboundary in South China. Stratigraphic Units this paper Hou et al. 1980 Liao, 1980 Yang, et al. clastic limestone lithofacies Series Stage 1987 lithofacies Lower Triassic Griesbachian Crurithyris speciosa- Lingula sub- circularis assemblage zone Crurithyris pusilla— Lingula subcircularis assemblage zone c (0 E L_ 8 x > 0 COMPARISON. The new species closely resembles A. FIGURE 38.—Serial sections of Araxathyris subpentangulata, new species subpentangulata in its anterior commissure and sulcus shape; showing the internal features. 50 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY convexity somewhat posterior to midlength; beak short, incurved; foramen covered by beak; sulcus from umbonal region narrow, extending over */2 length of valve, widening rapidly. Brachial valve somewhat less strongly convex; beak slightly incurved; sulcus narrow from umbonal region to near anterior margin, no fold, only flattened crest at anterior margin. Pedicle valve interior with small teeth, short thin dental plates fused to wall, only anterior edge free, strongly divergent near floor then curved; muscle area deeply impressed, separated by very low median ridge. Brachial valve interior with thin plate, perforated near posterior margin, at middle part becoming a shallow furrow, short, disappearing when hinge plate flattened anteriorly; hinge socket shallow, inner-socket ridge proportionately high; cardinal process short, comb-like at posterior, bilobed at anterior; crural bases seemingly forming sides of hinge plate; axis of spiralia laterally pointed, including 5-6 volutions per spiralium. Holotype.— USNM 456168. Measurements (mm).— Pedicle valve Brachial valve Curved Curved Thick- USNM Length length Length length Width ness 456165 26.6 35.7 24.2 28.0 25.4 15.4 456166 22.9 28.9 20.0 21.7 18.5 13.1 456167 20.1 26.0 19.0 20.2 17.8 12.9 456168 13.5 16.7 11.9 12.8 12.7 7.8 Stratigraphic Occurrence and Locality. —Long- dongchuan Formation: Xikou section, Zhenan County, Shaanxi Province. Diagnosis. —Smooth, subpentagonal outline, with sulci on both valves, anterior commissure usually weakly parasulcate. COMPARISON.— Araxathyris subpentangulata is character¬ ized by its narrow sulci on both valves and a weakly parasulcate commissure. In these respects it resembles A. quadrilobata (Abich, 1878:53, 54, pi. 7: fig. 6, pi. 9: figs. 8, 9; Grunt, 1965:243, 244, pi. 43: figs. 1, 2); but it differs in its smaller maximum size, slightly elongated outline, and sulcus wider at the anterior margin of the pedicle valve. It differs from A. zhijinensis Liao (1980a:267, pi. 8: figs. 52, 53) in its subpentagonal outline and parasulcate commissure; the latter species has a triangular shape and an uniplicate anterior commissure. Etymology.— From the Latin prefix, sub, meaning some¬ what; latinized from the Greek, penta. meaning five; and from the Latin, angulata, meaning having angles; hence, somewhat pentagonal. Rectambitus gen. nov. Small to medium size, biconvex, outline subpentagonal to subelliplical slightly elongate or transverse; anterior commis¬ sure rectiinarginate with emarginate anterior; narrow and shallow sulcus at each valve, brachial sulcus moderately or weakly developed; growth laminae irregularly spaced, most frequent near anterior margin; pedicle beak somewhat incur¬ ved, arching over brachial beak. Pedicle valve interior with strong knob-like teeth; dental plates thin, bow-like, converging together to form spondylium, attached at floor in apical cavity, and supported by low and broad median septum at anterior edge of spondylium; muscle scars narrow, elongate in spondylial floor. Brachial valve interior with undivided hinge plate, thin and flat; hinge socket shallow, inner-socket ridges relatively high; 2 comb-like processes interpreted as cardinal processes, separated widely; crural bases butting between outer and inner hinge plates; athyridid spiralia. Diagnosis.—S mooth Athyridoidea with rectimarginate commissure and emarginate anterior edge; narrow sulcus in both valves; internally, dental plates convergent, forming spondylium. Comparison. —The new genus is comparable to Araxathy¬ ris in having a dorsal sulcus, thin dental plates, and undivided hinge plate. The obvious distinctions are that the new genus has a rectimarginate commissure and a spondylium. It resembles Leptathyris Siehl (1962:212) in its bisulcate and rectimarginate commissure, but that Devonian genus has deeply depressed hinge plates in the dorsal valve and dental plates in the ventral valve, and both plates are fused completely with the shell wall (see A.J. Boucot et al. in Moore, 1965:H633, fig. 538,4). Type Species.— Araxathyris bisulcata Liao (1980a:268, pi. 9: figs. 14-17). Etymology. —From the Latin, rectus, meaning proper, and ambitus, meaning curvature (of margin). The gender is masculine. Rectambitus bisulcatus (Liao, 1980), new combination Figures 38(19-30), 39 Araxathyris bisulcata Liao, 1980a:268, pi. 9: figs. 14-17. DIAGNOSIS.— Average size for genus; moderately biconvex; greatest width somewhat anterior to midlength, maximum thickness near both umbonal regions; bisulcate, narrow and shallow, pedicle sulcus slightly wider than brachial sulcus; anterior commissure rectimarginate, rounded edge emarginate in middle. Pedicle valve spondylium supported by low and broad median septum extending about 2 mm at anterior edge of spondylium. Brachial valve with bilobate, comb-like hinge processes, diverging widely. Measurements (mm).— Pedicle valve Brachial valve Curved Curved Thick- USNM Length length Length length Width ness 456170 17.9 21.2 14.1 16.0 16.6 8.8 456171 15.5 16.0 14.4 16.0 16.7 9.0 456172 15.3 18.9 16.1 13.5 8.8 456173 13.7 17.9 12.4 15.9 12.6 8.4 NUMBER 76 51 FIGURE 39.— Serial sections of Rectambitiis bisulcata (Liao) showing internal features. Stratigraphic Occurrences and Localities.— Longtan Formation: Qiaozishan section, Anshun County. Guizhou Province. Longdongchuan Formation: Xikou section, Zhenan County, Shaanxi Province. Discussion.— Liao's (1980a:268) brief description of Araxathyris bisulcata, based on a single specimen from Qiaozishan section, Anshun County, Guizhou Province, em¬ phasized its rectimarginate anterior commissure and narrow sulcus. Examination of our specimens from Xikou section, Zhenan County, Shaanxi Province, shows them to be externally identical to Liao's specimen. Liao did not illustrate the internal features of his specimen, but because he placed it in the genus Araxathyris. we must assume that the specimen has internal features compatable with those of that genus. Rectambitus and Araxathyris differ in that Araxathyris has a spondylium; however, this character easily may be overlooked, and otherwise the two genera share many similar internal features. In addition, the localities for Liao's specimen and our specimens are near the west margin of the South China Sea, and they also share a common age. Based upon all of this, we consider them to be the same species. Genus Spirigerella Waagen, 1883 Diagnosis.— Smooth Athyrididae with strongly incurved pedicle beak, fairly sharp brachial beak, uniplicate or parasul- cate anterior commissure; pedicle valve interior with bow-like dental plates, with most parts merged into thickened wall; brachial valve interior with flat, undivided hinge plate, posterior part perforated, cardinal process usually well devel¬ oped or with inner-socket ridges, position of crura usually near sides of hinge plates. Discussion.— Some species of smooth Athyrididae that occur in the Upper Permian of South China have been assigned incorrectly to the genus Araxathyris. For example, A. shuizhu- tangensis Chan (in Hou et al„ 1979:99, pi. 8: figs. 7, 8, 10, 15, 16) and A. guizhouensis Liao (1980a:268, pi. 9: figs. 1-4) should be attributed to Spirigerella. Every species of Araxathy¬ ris has a distinct median groove in the dorsal valve. Examination of a large collection of Araxathyris compiled by Grant from Kuh-e-Ali Bashi, Iran (inventory of USNM), proves that this character is one of the most consistent features of the genus. Araxathyris guizhouensis Liao (= Araxathyris shuizhutangensis Chan) has a trace of a median groove that can be seen only dimly; this character essentially differs from the obvious median groove that is deeply scored and somewhat widened forward. Grant’s comparison of Composita with the genus Spirigerella (Grant, 1976:204, 205) pointed out several features that belong especially to Spirigerella: strongly incurved beak with foramen absent or only very small; greatest width farther forward; thickened wall submerging most of dental plates; and crura normally extending from the sides of the hinge plate. In his original description, Waagen (1883:451) said that the genus Spirigerella has an “exceedingly large” cardinal process. The features mentioned by Grant (1976) and Waagen (1883) are visible in the species described below from South China (Figures 42-44). Grant (1976:204) suggested that the species under the name Athyris timorensis (Rothpletz) by Huang (1933) might belong to the genus Spirigerella. This work proves that analysis to be true. Based on its strongly incurved beak, no foramen, no median groove on the fold, thickened wall submerging dental plates in the pedicle valve, and flattened hinge plate in the brachial valve, we place Athyris timorensis in the genus Spirigerella. Araxathyris normally has a roof-like hinge plate (see Grunt, 1965:241). Type Species.— Spirigerella derbyi Waagen (1883:450- 455, pi. 35: figs. 4-7,9-13). Spirigerella discusella sp. nov. Figures 40, 43(36-47) Average size for genus; moderately biconvex; outline nearly circular, an oblate spheroid with greatest width at midlength, maximum thickness somewhat posterior to midvalves; anterior commissure rectimarginate or weakly incurved with emargi- nate or straight edge; growth laminae irregularly spaced. Pedicle valve evenly and moderately convex in both lateral and anterior profile; beak short and incurved; sulcus narrow and shallow, originating from umbonal region. Brachial valve evenly and slightly inflated in both lateral and anterior profile. 52 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Figure 40. —Serial sections of Spirigerella disciisella. new species, showing the internal features. umbonal region roundly arched; no fold; brachial beak strongly incurved, attenuated, and sharply pointed. Pedicle valve with relatively strong teeth; dental plates thin, converging at apical cavity. Brachial valve with undivided hinge plate; hinge sockets shallow, inner-socket ridges high; crural bases butting between outer and inner hinge plates. Holotype.— USNM 456175. Measurements (mm).— Pedicle valve Brachial valve Curved Cun’ed Thick USNM Length length Length length Width ness 456174 12.4 16.6 10.8 12.6 11.9 8.0 456175 14.4 18.2 13.2 14.5 13.1 6.7 Stratigraphic Occurrences and Localities.— Changxing Formation: Beipci section, Zhongqing City, Si¬ chuan Province; Huangzhishan section, Wuxing County, Zhejiang Province. Diagnosis. —Oblate spheroid in outline; no fold on brachial valve; anterior commissure rcctimarginate or slightly up- curved. Comparison.— Spirigerella diseusella, new species, is similar to S. shuizhutangensis (Chan in Hou et al., 1979:99, pi. 8: figs. 7, 8, 10, 15, 16) in its subcircular outline and strongly incurved and attenuated brachial beak; it differs from S. shuizhutangensis in its slightly up-curved anterior commissure and lack of fold. Etymology.— From the Latin, discus, meaning a circular plate, plus -ella, diminutive. Spirigerella guizhouensis (Liao, 1980), new combination Figures 41(1-51), 42 Athyris timorensis (Rothpletz).—Huang, 1933:69-71, pi. 10: figs. 13-19.— Wang et al., 1964:612, pi. 120: figs. 13-16. Araxathyris guizhouensis Liao, 1980a:268, pi. 9: figs. 1-4. Medium size, strongly biconvex; outline subcircular, either slightly elongate or slightly transverse, greatest width normally anterior to midlength; anterior commissure parasulcate; growth lines fine, closely spaced, visible only in well-preserved specimens, growth laminae irregularly spaced, most frequent near anterior margin. Pedicle valve strongly and evenly convex; beak short, pointed or slightly blunt, strongly incurved, usually embracing the brachial beak; sulcus shallow, beginning near midlength and widening rapidly, rounded floor forming relatively wide tongue-like edge, bounded by short, rounded lateral plicae at each side. Brachial valve somewhat less strongly convex, greatest inflation in umbonal region; fold gently arched, parasulcus on each side, shallowly rounded. Pedicle valve interior with hinge teeth small; dental plates thin, bow-like, free entirely, extending about or l /4 length of valve, converging at floor. Brachial valve interior with small cardinal process, slightly elevated over the posterior edge of hinge plate; hinge plate flat, undivided; dental socket shallow; inner-socket ridges rather high, contacting at sides of hinge plate at approximately right angles; fulcral plates hanging under sockets, low and short; crural bases butting in hinge plate, between inner and outer hinge plates, continuing toward anterior after hinge plate disappears, and connecting with FIGURE 41 (opposite page).— Spirigerella guizhouensis (Liao): 1-3, 9, 10, dorsal, ventral, anterior, lateral, and posterior views (xl), USNM 456177; 4, 5, 11-13, dorsal, ventral, lateral, posterior, and anterior views (xl), USNM 456179a; 16-20, idem (xl), USNM 456176; 40-42,51, dorsal, ventral, lateral, and anterior views (x2) and 46, anterior view (xl), USNM 456179b; 43-45, dorsal, ventral, and lateral views (xl) and 47-50, dorsal, ventral, posterior, and anterior views (x2), USNM 456180; all from section 27. 6-8, 14, 15, dorsal, ventral, lateral, posterior, and anterior views (xl), USNM 456178, from section 4. 23-27, dorsal, ventral, lateral, posterior, and anterior views (xl), USNM 456181a; 21, 22, anterior views (xl), USNM 456181b,c; 29, 30, ventral and lateral views (xl); 33-35, 38, 39, dorsal, ventral, lateral, posterior, and anterior views (x2), USNM 456182; all from section 6. 28, 36, 37, ventral, dorsal, and lateral views (x2) and 31, 32, dorsal and lateral views (xl), USNM 456218, from section 8. NUMBER 76 53 54 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 42. —Serial sections of Spirigerella guizhouensis (Liao) showing internal features. spiralia; muscle scars narrow, separated by median ridge. Measurements (mm).— Pedicle valve Brachial valve Curved Curved Thick- length to USNM Length length Length length Width ness thickness 456176 16.9 26.2 14.8 19.5 15.8 12.4 1.36 456177 14.8 22.0 13.2 15.2 13.5 9.6 1.54 456178 14.9 19.8 13.7 16.8 14.7 10.3 1.44 Stratigraphic Occurrences and Localities.— Changxing Formation: Huayingshan section, Linshui County; Shangsi section, Guangyuan County; Beipei section, Zhongq- ing City, Sichuan Province; Huangzhishan section, Wuxing County, Zhejiang Province. Diagnosis.— Subcircular in outline; parasulcate anterior commissure; beak strongly incurved; internally, dental plates relatively long, converging at floor; hinge plate flat. Comparison.— Spirigerella grandis (Davidson) and S. obesa were identified by Huang (1933), based on material from Huayingshan (Hanyuanhsien) section, Sichuan Province, and Zunyi County (Tsunyihsien), Guizhou Province, respectively. Spirigerella guizhouensis differs from them in that S. grandis and S. obesa are elongate in outline, their sulcus and fold are almost obsolete, and they have strongly swollen umbonal regions (Huang, 1933:75-78, pi. 10: figs. 27-29, pi. 11: figs. 1 . 2 ). Discussion.— These specimens are identified here to be the same species as specimens from Guizhou Province that were assigned to Athyris timorensis (Rothpletz) by Huang (1933) and later assigned to Araxathyris guizhouensis by Liao (1980a), It is reasonable to discriminate specimens of South China from Timor’s specimens, as Huang (1933:71) implied when he described Athyris timorensis, pointing out that “it is to be noted that our shells do differ from Broili’s in some small but definite characters. Firstly, our shells are decidedly smaller in size; secondly, the lateral depressions on the brachial valve are better developed in ours than in the Timor specimens...As stated in the “Discussion” of the genus Spirigerella, this work assigns Athyris timorensis (Rothpletz) to the genus Spirigerella. Spirigerella ovaloides sp. nov. Figures 43(1-15), 44 Small for genus, strongly biconvex; outline subovate; hinge short and curved, greatest width near midlength; commissure strongly parasulcate; growth lines fine and closely spaced, growth laminae irregularly distributed, most frequent near anterior margin. Pedicle valve strongly convex; beak pointed and incurved over brachial beak; sulcus shallow, beginning somewhat anterior to midlength, rounded tongue-like at anterior edge, bounded by low, narrow and short plica, with median groove in sulcus, its trace extending to umbonal region in many specimens. Brachial valve somewhat equally strongly convex; beak incurved; fold roundly arched from midlength to anterior margin, parasulci beside fold shallow and narrow, visible only near anterior margin. Pedicle valve interior with stout teeth; short thin dental plates fused to wall, anterior edge free only for about V 2 their height; muscle area deeply impressed. Brachial valve interior with thin hinge plate, posterior part somewhat thickened and perforated by pit at posterior edge, becoming median furrow about midlength; hinge sockets shallow, inner-socket ridges forming sides of hinge plate, extending anteriorly, connected with crural bases. Syntypes.— USNM 456183-456185. Measurements (mm).— Pedicle valve Brachial valve Curved Curved USNM Length length Length length Width Thickness 456183 12.6 18.0 11.2 12.1 10.1 9.5 456184 11.8 16.0 10.5 11.9 10.3 8.9 456185a 11.2 16.1 10.0 11.8 9.9 8.8 456185b 11.2 16.0 10.2 12.5 10.7 8.3 456185c 9.7 15.5 8.4 12.0 8.3 6.7 Stratigraphic Occurrence and Locality. —Changxing Fonnation: Yutangjiao section, Nantong County, Sichuan Province. NUMBER 76 55 FIGURE 43 (opposite column).— Spirigerella. 1-15, Spirigerella ovaloides, new species: 1-5, dorsal, ventral, lateral, posterior, and anterior views (xl), USNM 456183; 6-10, idem (xl), USNM 456184; 11-15, idem (xl), USNM 456185; all from section 9, all syntypes. 16-35, Spirigerella shuizhutangen- sis (Chan): 16-20, dorsal, ventral, lateral, posterior, and anterior views (xl), USNM 456186; 21-25, idem (xl). USNM 456187; 26-30, idem (xl), USNM 456188; and 31-35, idem (xl), USNM 456189; all from section 15. 36-47, Spirigerella discusella. new species: 36-40, dorsal, ventral, lateral, posterior, and anterior views (xl), from section 27, USNM 456174, paratype; 41, 43, 45-47, dorsal, ventral, posterior, anterior, and lateral views (x2) and 42, 44, dorsal and ventral views (xl), from section 8, USNM 456175, holotype. 56 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Diagnosis. —Small, strongly biconvex, strongly parasulcate with narrow and short plica each side of sulcus at anterior edge of pedicle valve, and rounded tongue-like sulcus edge. Comparison. — Spirigerella ovaloides is similar to S. derbyi Waagen (1883:453-455, pi. 35: figs. 4-7, 9-13, pi. 37: figs. 11-13; see also Grant, 1976:233, pi. 65: figs. 35-49) in its elongated elliptical, globular, strongly incurved beak, and tongue-like sulcus edge, but the new species has short plicae beside the sulcus, short parasulci beside the fold, and a median groove in the sulcus. The specimen from Qidi Village, Xizang (Tibet), identified by Diener (1897:63, pi. 11: fig. 14) as S. derbyi has a proportionately wide sulcus and small shell; therefore, it rather resembles the new species except for the differences mentioned above. The new species is most nearly comparable to S. obesa Huang (1933:77, 78, pi. 11: figs. 1, 2) in its size being small, elongate, and obese, but S. obesa has neither a fold nor a sulcus. It differs from S. pentagonalis Chao (1927:95-97, pi. 1: figs, la-d) in its obvious fold and short parasulci lateral to the fold, and its tongue-like sulcus edge. Etymology. —From the Latin, ovalis, meaning oval, and -oides, meaning similar. Spirigerella shuizhutangensis (Chan, 1979), new combination Figures 43(16-35). 45 Araxathyris sliuizluitangensis Chan in Hou et al., 1979:99, pi. 8: figs. 7, 8, 10, 15, 16. Medium size, moderately to strongly biconvex; outline subtriangular to subcircular, length nearly equal to width or slightly elongate; hinge short and curved, widest anterior to midlength; anterior commissure uniplicate to parasulcate, some specimens only weakly uniplicate; growth laminae irregularly spaced. Pedicle valve with maximum convexity in posterior part, flattened from midvalve forward; sulcus very shallow to deep, flattened or rounded floor, narrow furrow from umbonal region, widened from midvalve forward; beak small, incurved. Brachial valve with even convexity; fold prominent only near anterior margin; brachial beak incurved, attenuated, and pointed. Pedicle valve interior with short dental plates, almost converged at floor; teeth stout. Brachial valve interior with shallow hinge sockets, inner-socket ridges high; hinge plate flat, undivided. Measurements (mm).— Pedicle valve Brachial valve Curved Curved Thick USNM Length length Length length Width ness 456186 14.4 18.0 12.5 15.8 13.0 10.0 456187 12.6 16.5 11.4 14.2 11.8 8.7 456188 11.7 16.4 10.7 13.1 11.6 8.5 456189 10.5? 13.0 10.3 11.7 10.2 6.5 FIGURE 45. —Serial sections of Spirigerella shuizhutangensis (Chan) showing the interior features. Stratigraphic Occurrences and Localities. —Paoshui Formation: Paoshui section, Laibin County, Guangxi Province. Changxing Formation: Huangzhishan section, Wuxing County, Zhejiang Province. Diagnosis. —Subcircular-shape Spirigerella with attenu¬ ated and pointed brachial beak. Comparison. — Spirigerella shuizhutangensis (Chan) is similar to S. guizhouensis in its subcircular outline, shallow sulcus that widens rapidly from midvalve, and convergent dental plates, but the former usually has the anterior part of the pedicle valve more flattened, a wider anterior edge of sulcus, and an attenuated and pointed brachial beak. It differs from S. ovaloides. new species, in its medium-size shell, subcircular outline, flattened anterior part of pedicle valve, and attenuated and pointed brachial beak. Genus Tongzithyris Ching, Liao, and Fang, 1974 Tongzithyris Ching, Liao, and Fang, 1974:313.—Liao, 1980a:268. Large size, strongly or moderately inflated; outline subpen¬ tagonal to subcircular; anterior commissure para-episulcate (not typical episulcate because of no median fold in pedicle sulcus); growth laminae irregularly spaced, most frequent near anterior margin; shell thickened at posterior part. Pedicle valve strongly convex, lateral slopes moderately steep or gently inclined; beak short, slightly incurved, foramen covered by beak; delthyrium covered by a pair of plates analogous to deltidial plates; sulcus beginning near beak, bounded by 2 gently arched plications, anterior edge tongue- shape. Brachial valve somewhat less strongly inflated; beak incurved; fold gently arched with median groove, lateral sulci originating somewhat posterior to midvalve. Pedicle valve interior with strong teeth; dental plates proportionately thin, bow-like, convergent at floor, forming pseudospondylium by filling with callus. Brachial valve interior with bilobate or comb-like cardinal processes; hinge plate undivided, flat; inner-socket ridges high; muscle scars narrow and elongate, separated by median ridge; spiralia pointed laterally. Comparison. — Tongzithyris resembles certain large species of Araxathyris, such as, A. protea (Abich) from Dzhulfian NUMBER 76 57 strata at Kuh-e-Ali Bashi, Northeast Iran, and at Dzhulfa, Azerbaijan (Smithsonian Accessions 284193 and 346823), in external features, but the former has more pronounced lateral sulci flanking the fold and a pseudospondylium in the pedicle valve. Discussion. —Ching et al. (1974) did not point out that this genus has a pseudospondylium in the pedicle valve when they established Tongzithyris as a genus. Liao (1980a:268) rede- scribed the genus in more detail: “Ventral valve interior with big teeth; dental plates well-developed, bowlike, extending at both sides, converged at floor; median ridge low.” After examining specimens of T. anshunensis Liao, which has a filled callus between convergent dental plates (Liao, 1980:269, fig. 2), we conclude that a pseudospondylium exists in Tongzithy¬ ris. Furthermore, a specimen of Tongzithyris episulcata from Tongzi County, Guizhou Province (Smithsonian Accession 339345), which is an inner core, verifies that a pseudospondy¬ lium is clearly visible in the umbonal area (Figure 47(5-8)). TYPE Species. — Tongzithyris episulcata Ching, Liao, and Fang (1974:313, pi. 165: figs. 10-13). Tongzithyris sichuanensis Xu, 1987 Figures 46,47(1-4) Tongzithyris sichuanensis Xu in Yang et al., 1987:233, 234, pi. 8: figs. 13, 14. Diagnosis. —Large size, moderately convex; subpentagonal in outline; anterior commissure para-episulcate; pedicle sulcus beginning near beak, roundly shallow with median grove at floor, anterior edge tongue-like, turn-up to dorsal, lateral furrows pronounced near anterior margin; brachial fold starting near umbonal region, dominated at anterior to midvalve with narrow and shallow groove, flanks of fold steeply tilted, lateral FIGURE 46.—A section near the beak of Tongzithyris sichuanensis Xu showing the pseudospondylium in the pedicle valve and the median ridge in the brachial valve. FIGURE 47.— Tongzithyris. 1-4, Tongzithyris sichuanensis Xu: dorsal, ventral, lateral, and anterior views (xl), from section 6, USNM 456190, holotype. 5-9, Tongzithyris episulcata Ching, Liao, and Fang: dorsal, ventral, posterior, lateral, and anterior views (xl), from Tongzi County, Guizhou Province, USNM 481645. (Reduced to 9772% for publication.) 58 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY plications gently arched, forming asymmetrical lateral sulci between fold and lateral plications. Measurements (mm).— Pedicle valve Brachial valve USNM Length Cun’ed length Length Curved length Width Thick¬ ness 456190 38.0 57.5 34.0 41.8 40.0 28.5 Stratigraphic Occurrence and Locality. —Changxing Formation: Huayingshan section, Linshui County, Sichuan Province. Comparison. —This species is characterized by its roundly shallow sulcus with median groove at floor, gently arched fold with shallow median groove that is flanked by asymmetrical lateral sulci. These features are similar to T. episulcata, but the latter has a more inflated and regular-shape shell, median groove that begins at the beak, and a deeply V-shape sulcus. It resembles T. anshunensis Liao in outline and features of the fold and sulcus, but it differs in that the latter has no lateral plications nor a median groove on either fold. Superfamily Reticularioidea Waagen, 1883 Family Elythidae Frederiks, 1924 Genus Squamularia Gemmellaro, 1899 Squamularia formilla sp. nov. Figure 48(1-24.26, 27) Small, outline subcircular, length and width about equal; widest posterior to midlength of brachial valve; biconvex with pedicle valve more inflated; commissure rectimarginate, neither fold nor sulcus, or, if present, only mace of sulcus on pedicle valve; growth lamellae regularly spaced, more closely spaced on brachial valve than on pedicle valve, numbering about 12-13 and 7-9 per 5 mm, respectively, in juvenile individuals and about 8-10 and 6-8 per 5 mm, respectively, in mature individuals; each edge of lamellae bearing 2 rows of fine pustules, upper row is spine-base scars, in well-preserved specimens fine spines visible, arranged radially with backward pointing spines attaching on each lamella, rarely preserved double-barrelled spine-scars. Pedicle valve strongly convex with maximum convexity near umbonal area in lateral profile and evenly arched in anterior profile; beak thick, blunt, strongly incurved; interarea small, poorly defined; delthyrium proportionately large and open. Brachial valve slightly and evenly convex; beak short, suberect; notothyrium broad, shallow. Pedicle valve interior with small teeth, short dental ridges; adminicula and septa absent; delthyrial ridges narrow, fur¬ rowed distally, hook-like in cross section view. Brachial valve interior with shallow hinge sockets. Syntypes. —USNM 456191-456195. Measurements (mm).— Pedicle Brachial valve valve Maximum Hinge USNM length length width width Thickness 456191 10.1 9.0 10.6 6.5 6.1 456194 11.3 11.0? 7.0? 456192 18.0 18.5 456195 19.3 16.7 19.5 9.8 11.3 Stratigraphic Occurrences and Localities. —Longtan Formation: Shatian section, Daye County, Hubei Province. Changxing Formation: Huayingshan section, Linshui County, Sichuan Province; Huangzhishan section, Wuxing County, Zhejiang Province. Diagnosis. —Small-size Squamularia with two rows of spine scars at edge of each growth lamella, neither fold nor sulcus, and proportionately wide delthyrium. Comparison. —Squamularia formilla, new species, is simi¬ lar to Squamularia elegantuloides Grabau (1931:198-200, pi. 16: figs, la-f, 2) in its wide delthyrium and its rectimarginate commissure with neither fold nor sulcus; however, the Mongolian species is larger, more transverse, and has more prominent concentric lamellae with very fine pustules. Squamularia elegantula (Waagen, 1883:545, 546, pi. 44: fig. 1) has a large shell and a narrow sulcus, so it is readily distinguishable from the new species. Squamularia formilla differs from S. waageni (Loczy) (see Chao, 1929:93-95, pi. 11: figs. 7-11; Grabau, 1931:204-206, pi. 16: figs. 5a-d) in that the latter has a medium-size shell, an elongately ovate outline, and a subelevated pedicle beak. Even though Chao’s illustrated specimens present a shallow sulcus and Grabau’s specimen has no sulcus, they seem to resemble the new species when characters of the hinge area and the delthyrium are considered. Squamularia formilla resembles S. asiatica Chao (1929:91— 93, pi. 11: figs. 12-14), which was regarded as the type species when Likharev (1934) established the genus Neophricadothy- ris, but the spiralia of S. asiatica have yet to be found. The new species differs from S. asiatica in having a wide delthyrium, small hinge area, and a relatively less convex brachial valve. As Liao (1980a:265, 266) pointed out, S. asiatica is a Carbonifer¬ ous species, but some specimens collected from the Upper Permian were identified as that species. A specimen from the Longtan Formation in Zunyi County, Guizhou Province, identified by Huang (1933:40,41, pi. 6: figs. 7,8), turns out not FIGURE 48.—1-24, 26, 27, Squamularia formilla, new species: 1-5, dorsal, ventral, lateral, posterior, and anterior views (x2), 8-10, dorsal, ventral, and lateral views (xl). USNM 456191, from section 27; 6, 7, ventral and lateral views (x2), 11, 12, ventral and lateral views (xl), and 23, 26. micro- omamentation (xl0.4, x20.8, respectively), USNM 456194, from section 6; 13-15, 19, 20, dorsal, ventral, lateral, posterior, and anterior views (x2) and 17, 18, dorsal and lateral views (xl), USNM 456195, from section 17; 16, 22, dorsal and ventral views (x2) and 24, 27, micro-ornamentation (xl0.4, x20.8, respectively), USNM 456193; 21, ventral view (x2), USNM 456192; all from section 6, all syntypes. 25, 28, Neophricadothyris sp.: micro-ornamentation (xl0.4), USNM 481647, 481646, respectively. NUMBER 76 59 _ * 4 ^ ^ % 60 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 49.—Serial sections of Hustedia orbicostata, new species, showing the internal features. to be that species. The new species differs from Huang’s specimen in that it has no pseudodeltidium. Etymology. —From the Latin, forma, pretty, and -ilia, diminutive. Suborder Retziidina Boucot, Johnson, and Staton, 1964 Superfamily Retzioidea Waagen, 1883 Family Retziidae Waagen, 1883 Genus Hustedia Hall and Clarke, 1893 Hustedia orbicostata sp. nov. Figures 49,50(1-23) Medium size for genus, moderately biconvex; outline elongately subovate; commissure strongly serrate; neither fold nor sulcus present; costae strong, rounded, numbering 10-12 on pedicle valve, 9-11 on brachial valve. Pedicle valve moderately convex, greatest convexity near umbo, slowly curved to anterior in lateral profile; anterior profile flattened or slightly arched with inclined sides; beak short, suberect; foramen round, mesothyridid; delthyrium relatively large, covered by deltidial plates. Brachial valve slightly more strongly convex with maximum convexity at posterior; beak blunt, slightly incurved over hinge. Pedicle valve interior with stout teeth, tooth ridges very short; pedicle collar not well preserved. Brachial valve interior with median septum high but short, only in umbonal area; hinge plate greatly thickened with proportionately big hole at posterior part and ligulate process projecting anteriorly; hinge sockets narrow with high inner-socket ridges; crura rod-like. short; spiralia unknown. SYNTYPES.— USNM 456196-456198. Measurements (mm).— Pedicle valve Brachial valve Pedicle USNM Length Curved length Curved Length length Width Thick¬ ness valve costae 456196 9.8 12.7 7.8 8.1 7.5 6.1 10 456197 9.9 12.9 8.4 8.2 8.0 6.1 10 Stratigraphic Occurrences and Localities.— Changxing Formation: Yutangjiao section, Nantong County; Beipei section, Zhogqing City, Sichuan Province. Lower Yinkeng Formation: Huangzhishan section, Wuxing County, Zhejiang Province. Diagnosis. —Medium-size Hustedia with rounded costae, no fold and sulcus, but with high median septum in brachial valve. Comparison. —Two species of the genus, Hustedia indica (Waagen) and H. remota (Eichwald), have been reported in Upper Permian rocks of South China. Hustedia orbicostata, new species, most nearly resembles the specimen from Guiyang County, Guizhou Province that was identified as H. indica by Huang (1933:78, 79, pi. 11: figs. 3, 3a-c; see also Wang et al., 1964:639, 640, pi. 126: fig. 21). It differs from Huang’s H. indica in that the new species has more than ten costae and lacks a sulcus and fold. Eumetria indica Waagen (1884:492-494, pi. 35: figs. 1, 2) has a globular outline, width nearly equal to length, and plications fewer than ten, thus, it is distinguished easily from the new species. According to Huang’s description, H. remota has plications numbering 12 on the pedicle valve and 14 on the brachial valve and has a remarkable character in that “the interplical spaces are NUMBER 76 61 FIGURE 50.— Hustedia orbicostata, new species: 1-5, dorsal, ventral, lateral, posterior, and anterior views (x2) and 6-10, idem (xl), USNM 456197, from section 9; 11-15, dorsal, ventral, lateral, posterior, and anterior views (x2) and 19, 20, 22, dorsal, ventral, and lateral views (xl), USNM 456196, both from section 27: 16-18, dorsal, ventral, and lateral views, (x2), USNM 456198, from section 8; 21. ventral view (xl), USNM 456199; 23, idem (xl), USNM 456200, both from section 27; all syntypes. (Reduced to 97'/2% for publication.) distinctly flat-bottomed and appear wider than the plicae” (1933:81). Obviously, Huang's specimens collected from Dading County (Tatinghsien), Guizhou Province, differ from the new species. Grabau (1931:125-128, pi. 7: fig. 8a-e; 1934:103, 104, pi. 7: fig. 3) emphasized the “sub-rotund” outline of H. remoia , and the figure shown by Huang (1933, pi. 11: fig. 5) is virtually circular in outline. Another specimen of H. remota. described by Grabau (1934:103, 104. pi. 7: fig. 3), is “nearly as wide as long.” Thus, the new species is readily distinguishable from //. remoia. Hustedia orbicosiaia. new species, is similar to H. spicata Cooper and Grant (1976b:2799. 2800, pi. 741: figs. 12-40) in both nonelevated median costa on the brachial valve and high median septum, but it differs in that the new species has rounded costae and neither fold nor sulcus. It somewhat resembles H. cepoidea Cooper and Grant (1976b:2767-2769, pi. 732: Figs. 56-85) in its outline and costae numbers, but the latter has angular costae and a low median septum. Hustedia decoUatensis Cooper and Grant (1976b:2778, 2779, pi. 735: figs. 54-68) is readily distinguishable from the new species in that the former has a low median ridge rather than a septum in the brachial valve even though they both are similar in their numbers of rounded costae and in having neither fold nor sulcus. Etymology. —From the Latin, orbis, meaning circle, and cosiata, meaning ribbed (rounded costae). Order Terebratulida Waagen, 1883 Superfamily Cryptonelloidea Thomson, 1926 Family Notothyrididae Likharev, 1960 Genus Rostranteris Gemmellaro, 1899 Rostranteris ptychiventria sp. nov. Figures 51,52(1-16) Small, usual size for genus; unequally biconvex; elongate- ovate to subtriangular in outline, widest anterior to midvalve or near anterior margin; sides broadly rounded to slightly angular; anterior commissure antiplicate; surface of brachial valve with posterior */2 smooth but anterior ‘/2 plicated, plicae on pedicle valve beginning from umbonal area, interspaces somewhat narrower than plicae; growth laminae well developed near anterior margin. Pedicle valve strongly convex in lateral profile with umbonal area fairly strongly curved with maximum convexity near midvalve, anterior profile strongly domed, crest serrate and sides sloping steeply or moderately; 2 strongest median costae forming fold, each side having 1 or 2 moderate costae. Brachial valve gently inflated, with steep anterior slope in lateral profile, anterior profile flattened or slightly depressed at median region, forming steep slope at each side; sulcus observable near anterior margin, bounded by 2 strong plicae, median plica in sulcus sometimes weak, sometimes strong, but always slightly lower than lateral plicae; each side having 1-2 plicae. Interior of pedicle valve without dental plates but with strong dental ridges, teeth stout. Brachial valve interior with small bilobate cardinal process; hinge sockets slit-like with strong inner-socket ridges; hinge plate undivided, flat, perforated at posterior edge; crural bases rod-like, butting in both sides of hinge plate and touching inner-socket ridges; loop centronelli- form, long, extending about 3.5 mm, with high median plate that extends beyond conjunction of main hands. HOLOTYPE.— USNM 456201. Measurements (mm).— Pedicle valve Brachial valve Curved Curved Thick- USNM Length length Length length Width ness 456201 10.1? 17.1 8.2 10.2 12.5 8.5 456203 10.2 14.9? 7.5 9.5 8.1 8.0 456202 11.2? 14.2 9.8 11.0 9.7 7.2 Stratigraphic Occurrence and Locality. —Long- dongchuan Formation: Xikou section, Zhenan County, Shaanxi Province. Diagnosis. —Small, antiplicate commissure; anterior l /2 plicated on brachial valve, plicae beginning from umbonal area on pedicle valve. Comparison. —This new species is mainly characterized by its long plicae on the pedicle valve, which distinguishes it from SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 51.—Serial sections of Rostranleris ptychiventria, new species, showing the internal features. NUMBER 76 63 FIGURE 52.— Roslranleris ptychiventria. new species: 1-4, dorsal, ventral, lateral, and anterior views (x2): 5-7, dorsal, ventral, and anterior views (xl), LSNM 4^6201, holotype; 8-11, dorsal, ventral, lateral, and anterior views (x2t. LSNM 456203, paratype: 12, 13, dorsal and ventral views (xl), LSNM 456203, paratype: 14-16, dorsal, ventral, and anterior views (xl), LSNM 456202. paratype: all from section 32. most other species of the genus Rostranteris. Rostranteris adrianensis Gemmellaro (1899:105, 106, pi. 25: figs. 35-39; see also Stehli, 1962:100, pi. 20: group A) has five weak plicae, three on the brachial valve and two on the pedicle valve, which set out near the frontal margin. This arrangement forms a striking contrast to the new species, which has strong plicae, more than 4 on each valve. Rostranteris pulchrum Gemmellaro (1899:106, pi. 25: figs. 58-62) is similar to the new species in its size and outline, but it is readily distinguishable in that the former has rather less thickness and has weak and short plicae. Rostranteris gibbosum Gemmellaro (1899:110. pi. 25: figs. 40, 41) has a relatively big, swollen shell and a conspicuous pedicle sulcus (USNM Accession 180508), although it is comparable to the new species in having long plicae on the pedicle valve. Rostranteris ptychiventria, new species, closely resembles R inflatum Gemmellaro (1899:109, pi. 25: figs. 42-45, pi. 30: fig. 41; Stehli. 1962:100, 101, pi. 20: group B) in its appearance, including shell size, outline, and profile, but it differs in that the latter has an obvious pedicle sulcus and relatively short plications originating near midvalve. It is externally compilable to Mongolina subdieneri Grabau (1931:103-105, pi. 8: figs. 5a-f), which has short plicae and whose interior features are not known. Discussion. —Several species of the genus have the pedicle valve sulcus, for example, R. sinuatum Gemmellaro (1899: 111, pi. 25: figs. 52-57; USNM Accessions 180508, 183389. and 2252.39) has a typical sulcus in the pedicle valve and a rather broad interspace between the two main plications. In the new species, however, the interspace between the two main plications on the pedicle valve is narrower than the plications, so it is unreasonable to call it a “sulcus” in this case. Etymology. —From the Greek, ptychos, meaning fold, and from the Latin, venter , meaning front (as in ventral). Genus Notothyris Waagen, 1882 Notothyris bifoldes sp. nov. Figures 53, 54(1-4) Small, moderately biconvex, elongate ovate in outline, greatest width near midlength; sides smoothly rounded, anterior margin subtruncated; anterior commissure rectimargi- nate with sharp angular indentation; surface smooth except for anterior '/3 where marked by 8-9 plications; 5 somewhat strong plications forming low fold with slightly depressed crest, brachial valve of some specimens having 3 median plications somewhat lower than 2 bound-lateral plications; 2 somewhat strong plications bounding low fold on pedicle valve, those having 2 slightly weakened plications between them; 1 or 2 lateral plications at each side; growth lines weak, occasionally raised. Pedicle valve strongly and evenly convex in lateral profile, with rounded bow-like curvature; anterior profile evenly rounded with steep sides; beak erect and slightly incurved; foramen circular, mesothyridid; delthyrium appearing to be covered by deltidial plates. Brachial valve somewhat less strongly convex; beak pointed, incurved. Pedicle valve interior with strong pedicle collar in delthyrial cavity; no dental plates, teeth stout. Brachial valve interior with shallow hinge sockets, fulcral plates strong but short, inner- socket ridges low; hinge plates fused together, crural bases forming at sides of hinge plates; posterior foramen seemingly expressed as narrow tube; muscle scars narrow oval-shape at umbonal region. Holotype. —USNM 456205. Measurements (mm).— Pedicle valve Brachial valve Curved Curved Thick- USNM Length length Length length Width ness 456204 10.2 13.6 8.0 8.6 7.8 6.8 456205 10.4 13.3 9.0 9.5 8.4 7.1 Stratigraphic Occurrences and Localities.— Changxing Formation: Huangzhishan section, Wuxing County, Zhejiang Province. Paoshui Formation: Paoshui section, Laibing County. Guangxi Province. Diagnosis. —Small, elongately ovate; anterior commissure rectimarginate with sharply indented margin; 7-8 plications; weak folds on both valves. Comparison. —Multiple plications at anterior '/3 and 2 weak folds on both valves distinguish this from other species of the genus. Notothyris triplax Grant (1976:253, 254, pi. 46: figs. 64 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 53.—Serial sections of Notothyris bifoldes, new species, showing the internal features. FIGURE 54 .—Notothyris bifoldes, new species: 1-4, dorsal, ventral, lateral, and anterior views (x 1.5), from section 15, USNM 456205, holotype. 1-8) is similar to the new species in outline, but it differs in having 3 plications on the dorsal valve and 4 on the ventral valve. As Grant pointed out, N. praelecta Reed (not Water- house and Piyasin, 1970) has more than 5 plicae on the dorsal valve (Grant, 1976:253), and Reed (1944:167) described it as “bearing 6 subangular longitudinal plications.” Thus, the number of plicae of N. praelecta seems to resemble the new species; however, N. bifoldes is smaller than N. praelecta, and N. praelecta has 2 median plicae that are much stronger than others on the pedicle valve. Notothyris bifoldes, new species, strongly resembles N. revocata Reed in respect to number and nature of median plications, which initially caused us to identify it as the same species, but considering the uncertain nature of the Salt Range species’ internal structures and obvious external differences, we describe this new species as differing from N. revocata in its small size, elongately ovate outline, no sulcus on the brachial valve, and fewer lateral plications. Discussion,— The interior structures of the new species basically coincide with genera Notothyris Waagen and Ros- tranteris Gemmellaro, but we have not ascertained what type of loop it contains. The cardinalia in several genera of the family Notothyrididae are almost the same, but the loops are different. The external dimension seems to reflect the shape of the loop among families within the Upper Permian. For example, Notothyris has numerous plications and an anterior commissure that is rectimarginate to faintly sulcate (Stehli in Treatise, 1965:H758; Grant, 1976:253); Rostranteris has an intraplicate anterior commissure (Stehli in Treatise, 1965:H759); and Timorina has 2, 3, or more median plications on the pedicle valve that are raised into a slight fold (Stehli in Treatise, 1965:H760). Accordingly, we hereby place the new species in the genus Notothyris, although it may be that N. praelecta is attributable to Timorina Stehli, 1961. 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