Polymerid Tribolites from the Cambrian of Northwestern Hunan, China, Two-Volume Set

POLYMERID TRILOBITES FROM THE CAMBRIAN OF NORTHWESTERN HUNAN, CHINA Volume 1 Corynexochida, Lichida, and Asaphida PENG...

Author: S. Peng | L.E. Babcock | H. Lin

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POLYMERID TRILOBITES FROM THE CAMBRIAN OF NORTHWESTERN HUNAN, CHINA Volume 1 Corynexochida, Lichida, and Asaphida

PENG Shanchi Loren E. BABCOCK LIN Huanling

i ~ Science Press

Beijing

Responsible Editresses: HU Xiaochun and LIN Caihua

POLYMERID

TRILOBITES

FROM THE CAMBRIAN

HUNAN, CHINA

Volume 1 Corynexochida, Lichida, and Asaphida PENG Shanchi, Loren E. BABCOCK, and LIN Huanling

Copyright © 2004 by Science Press Published by Science Press http://www, sciencep, com 16 Donghuangchenggen North Street Beijing 100717 P. R. China

Printed in Beijing, 2004 Reprinted in Beijing, 2006 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. ISBN 7-03-014905-X/Q • 1538

OF NORTHWESTERN

LIST OF AUTHORS PENG Shanchi Nanjing Institute of Geology and Palaeontology Chinese Academy of Sciences 39 East Beijing Road Nanjing 210008, China

Loren E. BABCOCK Department of Geological Sciences The Ohio State University 275 Mendenhall Laboratory 125 South Oval Mall Columbus, Ohio 43210, USA

LIN Huanling Nanjing Institute of Geology and Palaeontology Chinese Academy of Sciences 39 East Beijing Road Nanjing 210008, China

This research was funded by National Natural Science Foundation of China (49070077, 40072003, 40023002, 40332018) Chinese Academy of Sciences (KZCX2-SW- 129) Ministry of Technology and Science of China (2001 DEB20056, G2000077700) National Geographic Society (No. 5819-96; 7151-01) State Key Laboratory of Paleobiology and Stratigraphy (No. 933114) State Education Commission of China The Ohio State University US National Science Foundation (EAR 9405990, 9526709, 0073089, 0106883, EAR OPP-0346829)

FOREWORD China is richly endowed with Cambrian strata yielding some of the best-preserved fossils known anywhere in the world. The trilobites are most important element in the fauna, and of scientific relevance for several reasons. First, they are of regional importance in the precise correlation of strata, and are of primary use in characterising mappable formations. Second, some of the species are widespread internationally, and these permit the placing of Chinese stratigraphical schemes within the global chronostratigraphy. Thirdly, the variety, relationships and endemicity of the faunas contributes to the broad questions of how evolution proceeded in the Cambrian, and is relevant to debates about whether special conditions applied at the early stage of the Phanerozoic radiation. The polymerid trilobite faunas of northwestern Hunan are remarkable for their diversity and excellence of preservation. Many of them belong to genera confined to China or to its palaeogeographic neighbours. However, some of these important taxa have remained imperfectly known or undescribed. This monograph makes good this omission. Thus in several cases pygidia or free cheeks are assigned where they had not been known previously, thus providing a much fuller picture of morphology in assessing relationships. This is particularly welcome where the species concerned is the type of its genus. In addition a number of new genera are added to the fauna. The paper also documents the endemic radiation of the specialised and interesting Dameselloidea, a family showing some of the most specialised pygidia in the Trilobita. The systematics of these trilobites is fully discussed, and the whole work is illustrated by photographs of the highest quality. This work should remain the standard account for the foreseeable future. The detailed work on the trilobites is placed in the wider context of biostratigraphy and correlation, which should be of interest to all those concerned with Cambrian geology. A refinement of the biostratigraphic zones based on the ranges of trilobite species will have implications for those currently seeking to increase the precision in international correlation of subdivisions within the Cambrian. Careful new work, of which this paper is an excellent example, is a more valuable contribution than almost anything else in this endeavour.

Professor Richard A. Fortey F R S Department of Palaeontology The Natural History Museum and Department of Zoology Oxford University

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CONTENTS FOREWORD ...................................................................................................................................... i INTRODUCTION .............................................................................................................................. 1 ACKNOWLEDGMENTS

........................................................................................................... 2

R E P O S I T O R I E S .......................................................................................................................... 2

PREVIOUS WORK ........................................................................................................................... 3 GEOLOGIC SETTING ..................................................................................................................... 4 PALEOGEOGRAPHY

................................................................................................................ 4

S T R A T I G R A P H Y ....................................................................................................................... 4 M E A S U R E D S E C T I O N S ......................................................................................................... 10

CORRELATION ............................................................................................................................. 4 0 N O R T H C H I N A A N D N O R T H E A S T C H I N A P L A T F O R M S ............................................... 40 T A R I M A N D N O R T H W E S T C H I N A ...................................................................................... 4 0 W E S T E R N Z H E J I A N G ............................................................................................................ 42 A U S T R A L I A ............................................................................................................................. 42 B A L T I C A .................................................................................................................................. 42 K A Z A K H S T A N ........................................................................................................................ 43 L A U R E N T I A ............................................................................................................................. 43 S I B E R I A .................................................................................................................................... 43

SYSTEMATIC PALEONTOLOGY .............................................................................................. 4 4 C l a s s TRILOBITA W a l c h , 1771 ................................................................................................... 4 4 O r d e r CORYNEXOCHIDA K o b a y a s h i , 1935 ................................................................................. 44 S u b o r d e r CORYNEXOCHINA K o b a y a s h i , 1935 ............................................................................ 4 4 F a m i l y CORYNEXOCHIDAE A n g e l i n , 1854 ................................................................................. 44 G e n u s CORYNEXOCHUS A n g e l i n , 1854 ...................................................................................... 44

Corynexochus xiangxiensis sp. n o v . . ................................................................................. 45 G e n u s CORYNEXOCHINA L e r m o n t o v a , 1940 .............................................................................. 46

Corynexochina sinensis sp. nov.- ....................................................................................... 47 in Z h o u et al., 1977 ............................................................................ 48 Chatiania chatianensis Y a n g in Z h o u et al., 1977 ............................................................ 49 Chatiania expansa ( Y u a n a n d Yin, 1998) ......................................................................... 52 Chatiania sp. cf. C. chatianensis Y a n g in Z h o u et al., 1977 ............................................. 53 Chatiania convexa sp. n o v . . ............................................................................................... 53

G e n u s CHATIANtA Y a n g

F a m i l y DOLICHOMETOPIDAE W a l c o t t , 1916 .............................................................................. 54 G e n u s AMPHOTON L o r e n z , 1906 ................................................................................................ 54

Amphoton deois ( W a l c o t t , 1905) ....................................................................................... 56 Amphoton alceste ( W a l c o t t , 1905) ..................................................................................... 57 Amphoton sp. cf. A. typicum ( K o b a y a s h i , 1942a) .............................................................. 59 G e n u s FUCHOUIA R e s s e r a n d E n d o in K o b a y a s h i , 1935 ........................................................... 60 Fuchouia chiai Lu, 1957 ................................................................................................... 62 Fuchouia kuruktagensis Z h a n g , 1981 ................................................................................ 64 •

iii

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Fuchouia Fuchouia Fuchouia Fuchouia

oratolimba Yang in Z h o u et al., 1977 ............................................................... 65 bulba sp. nov.. .................................................................................................... 67 sixinensis sp. nov.. ............................................................................................. 68 sp. indet.. ............................................................................................................ 69

Family DORYPYGIDAE Kobayshi, 1935 ...................................................................................... 70 Genus DORYPYGE Dames, 1883 ................................................................................................. 70

Dorypyge richthofeni Dames, 1883 ................................................................................... 71 Dorypyge perconvexalis Yang in Zhou et al., 1977 .......................................................... 73 Dorypyge bisulcata sp. nov.. .............................................................................................. 75 Dorypyge huaqiaoensis sp. nov.. ....................................................................................... 76 Dorypyge globosa sp. nov.- ................................................................................................ 77 Dorypyge? sp.. ................................................................................................................... 78 Suborder LEIOSTEGIINA Bradley, 1925 ...................................................................................... 79 Superfamily LEIOSTEGIOIDEA Bradley, 1925 ............................................................................ 79 Family LEIOSTEGIIDAE Bradley, 1925 ....................................................................................... 79 Subfamily LEIOSTEGIINAE Bradley, 1925 .................................................................................. 79 Genus CHUANGIA Walcott, 1911 ............................................................................................... 79

Chuangia subquadrangulata Sun, 1935 ............................................................................ 79 Chuangia austriaca Yang in Zhou et al., 1977 ................................................................. 80 Genus MEROPALLA Opik, 1967 .................................................................................................. 81

Meropalla bella Yuan and Yin, 1998 ................................................................................ 82 Meropalla gibbera sp. nov.. ............................................................................................... 82 Subfamily CHELIDONOCEPHALINAE Wittke, 1984 ..................................................................... 83 Genus GEMINICLAVULA gen. n o v . . ............................................................................................ 83

Geminiclavula wangcunica gen. et sp. n o v . . ..................................................................... 84 Subfamily CHIAWANGELLINAE Chu, 1959 ................................................................................. 84 Genus CHIAWANGELLA Chu, 1959 ............................................................................................ 85

Chiawangella hunanensis sp. nov.- .................................................................................... 86 Subfamily ORDOSnNAE Lu, 1954 ............................................................................................... 89 Genus WANSHANIA R o n g and Yang in Zhou et al., 1977 .......................................................... 89 Wanshania wanshanensis Rong and Yang in Z h o u et al., 1977 ........................................ 90 Subfamily PAGODIINAE Kobayashi, 1935 ................................................................................. 93 PAGODIINAE gen. et sp. indet. •................................................................................................... 93 Genus PROCHUANGIA Kobayashi, 1935 ..................................................................................... 93

Prochuangia granulosa Lu, 1956 ...................................................................................... 94 Prochuangia linicispinata Peng, 1992 ............................................................................... 95 Prochuangia sp. cf. P. leiocephala Peng, Geyer, and Hamdi, 1999 ................................. 95 Order LICHIDA Moore, 1959 ...................................................................................................... 96 Superfamily DAMESELLOIDEA Kobayashi, 1935 ....................................................................... 96 Family DAMESELLIDAE Kobayashi, 1935 .................................................................................. 96 Subfamily DAMESELLINAE Kobayashi, 1935 ............................................................................. 96 Genus DAMESELLA Walcott, 1905 ............................................................................................. 96

Damesella hunanensis sp. nov.- ......................................................................................... 96 Damesella? sp.. .................................................................................................................. 98 Genus BLACKwELDERIA Walcott, 1906 ..................................................................................... 98

Blackwelderia youshuica sp. nov.. ..................................................................................... 99 Blackwelderia? sp.. .......................................................................................................... 100 Genus PARABLACKwELDERIA Kobayashi, 1942b .................................................................... 101 Parablackwelderia jimaensis (Yang in Lu et al., 1974a) ................................................ 104 •

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Parablackwelderia Parablackwelderia Parablackwelderia Parablackwelderia

laterilobata ( Y a n g in Z h o u et al., 1977) .......................................... 108 sp. cf. P. huabeiensis ( Z h a n g in Qiu et al., 1983) ............................ 109 yangi sp. nov. •.................................................................................. 110 s p . . ....................................................................................................

111

G e n u s PROTAITZEHOIA Y a n g 1978 in Yin and Li, 1978 ......................................................... 112

Protaitzehoia Protaitzehoia Protaitzehoia Protaitzehoia Protaitzehoia

yuepingensis Y a n g in Yin and Li, 1978 .................................................... granifera Y a n g in Yin and Li, 1978 .......................................................... subquadrata P e n g , 1987 ............................................................................ spinifera sp. n o v . . ......................................................................................

114 115

117 119

sp.. .............................................................................................................

120

S u b f a m i l y DORYPYGELLINAE K o b a y a s h i , 1935 ...................................................................... 121 G e n u s TEINISTION M o n k e , 1903 ..............................................................................................

121

Teinistion posterocostum ( Y a n g in Z h o u et al., 1977) .................................................... 123 G e n u s TAIHANGSHANIA Z h a n g and W a n g , 1985 ..................................................................... 125

Taihangshania wangcunensis sp. nov.. ............................................................................ 126 S u b f a m i l y DREPANURINAE Hup6, 1953 ...................................................................................

128

G e n u s DREPANURA B e r g e r o n , 1899 .........................................................................................

128

Drepanura ? crassispina sp. n o v . . ................................................................................... 129 G e n u s PALAEADOTES Opik, 1967 ............................................................................................

130

Palaeadotes hunanensis ( Y a n g in Z h o u et al., 1977) ...................................................... Palaeadotes bella (Qiu in Qiu et al., 1983) ..................................................................... G e n u s PARADAMESELLA Y a n g in Z h o u et al., 1977 ................................................................ Paradamesella typica Y a n g in Z h o u et al., 1977 ............................................................ Paradamesella peculiaris Z h o u in Z h o u et al., 1977 ...................................................... Paradamesella nobilis L u and Lin, 1989 .........................................................................

132 136 137 139 142 145

S u p e r f a m i l y ODONTOPLEUROIDEA K o b a y a s h i , 1935 ............................................................... 145 F a m i l y EOACIDASPIDAE P o l e t a e v a , 1957 .................................................................................

145

G e n u s PARAACIDASPIS P o l e t a e v a , 1960 ..................................................................................

146

Paraacidaspis hunanica E g o r o v a in P o l e t a e v a , 1960 ..................................................... 147 Paraacidaspis sp.. ............................................................................................................ 149 O r d e r ASAPHINA Salter, 1864 ...................................................................................................

150

S u p e r f a m i l y ANOMOCARAROIDEA P o u l s e n , 1927 ....................................................................

150

F a m i l y ANOMOCARIDAE P o u l s e n , 1927 ...................................................................................

150

G e n u s AFGHANOCARE Wolfart, 1974 ......................................................................................

150

Afghanocare truncatum (Peng, 1987) .............................................................................. 150 G e n u s GLYPHASPELLUS Ivshin, 1953 .......................................................................................

152

Glyphaspellus? sinensis sp. nov.. ..................................................................................... 152 G e n u s PARACOOSIA K o b a y a s h i , 1936 ......................................................................................

153

Paracoosia sp. cf. P. kingi W i t t k e , 1984 ......................................................................... 154 Paracoosia huayuanensis sp. nov.- .................................................................................. 156 G e n u s PAIBIANOMOCARE gen. n o v . . ........................................................................................

157

Paibianomocare paibiense gen. et sp. n o v . . .................................................................... 158 Paibianomocare lineatum gen. et sp. nov.- ...................................................................... 159 F a m i l y PTEROCEPHALIIDAE K o b a y a s h i , 1935 .........................................................................

160

S u b f a m i l y CILIINAE P e n g , 1992 ...............................................................................................

160

G e n u s YANGWEIZHOUIA Y u a n and Yin, 1998 .........................................................................

161

Yangweizhouia carinata Y u a n and Yin, 1998 ................................................................. 161 S u p e r f a m i l y ASAPHOIDEA Salter, 1864 ....................................................................................

162

F a m i l y CERATOPYGIDAE L i n n a r s s o n , 1869 .............................................................................

162 °

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Subfamily CERATOPYGINAE Linnarsson, 1869 ........................................................................ 162 Genus PROCERATOPYGE Wallerius, 1895 ................................................................................ 162 S u b g e n u s PROCERATOPYGE (PROCERATOPYGE) Wallerius, 1895 ............................................ 163

Proceratopyge Proceratopyge Proceratopyge Proceratopyge

(Proceratopyge) fenghwangensis Hsiang in E g o r o v a et al., 1963 .......... 163 (Proceratopyge)fuyangensis Lu and Lin in Peng, 1987 ........................ 164 (Proceratopyge) truncata Y a n g in Zhou et al., 1977 ............................. 165 sp. indet. •................................................................................................. 166

Subfamily IWAYASPIDINAE Kobayashi, 1962 .......................................................................... 166 Genus PSEUDOYUEPINGIA Chien, 1961 .................................................................................... 166

Pseudoyuepingia laochatianensis Yang in Z h o u et al., 1977 .......................................... 167 Superfamily TRINUCLEOIDEA H a w l e and Corda, 1847 ............................................................ 169 F a m i l y ALSATASPIDIDAE Turner, 1940 .................................................................................... 169 Subfamily HAPALOPLEURINAE Harrington and Leanza, 1957(nom. trans, ex. Hapalopleuridae Harrington and Leanza, 1957) ............................................................................................. 169 Genus GAOLOUPINGIA Yuan and Yin, 1998 ............................................................................ 170

Gaoloupingia gaoloupingensis Yuan and Yin, 1998 ....................................................... Genus AJRIKINA K r a s k o v in B o r o v i k o v and Kraskov, 1963 ................................................... Ajrikina wannanensis (Qiu in Qiu et al., 1983) ............................................................... Ajrikina hunanensis (Peng, 1987) ....................................................................................

170

171 172

173 Genus TORIFERA Wolfart, 1974 ............................................................................................... 174

Torifera taoyuanensis (Peng, 1987) ................................................................................. Torifera tuma (Yang in Z h o u et al., 1977) ...................................................................... Torifera abrupta sp. nov.. ................................................................................................ Torifera? paraconvexa (Yang in Yin and Li, 1978) ........................................................

175 176 178

179 Family LIOSTRACn~rIDAE R a y m o n d , 1937 ................................................................................ 180 Genus LIOSTRACINA M o n k e , 1903 .......................................................................................... 180

Liostracina bella Lin and Zhou, 1983 ............................................................................. 180 Superfamily UNCERTAIN .......................................................................................................... 181 Family MONKASPIDIDAE Kobayashi, 1935 .............................................................................. 181 Genus MONKASPIS Kobayashi, 1935 ........................................................................................ 181

Monkaspis quadrata Y a n g in Z h o u et al., 1977 .............................................................. 183 G e n u s METOPOTROPIS 0 p i k , 1967 ........................................................................................... 184

Metopotropis sinensis sp. nov.. ........................................................................................ 184 REFERENCES ............................................................................................................................... 186 INDEX OF SPECIES AND GENERA ......................................................................................... 201 P L A T E S .......................................................................................................................................... 207

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LIST OF TEXT-FIGURES Text-figure 1. Text-figure 2. Text-figure 3. Text-figure 4. Text-figure 5.

Text-figure 6. Text-figure 7.

Text-figure 8. Text-figure 9. Text-figure 10. Text-figure Text-figure Text-figure Text-figure Text-figure Text-figure Text-figure Text-figure Text-figure

11. 12. 13. 14. 15. 16. 17. 18. 19.

Map showing distribution of rocks assigned to the Jiangnan Slope Belt facies (Cambrian) in Hunan and Guizhou provinces, China Table showing the correlation of polymerid and agnostoid biozones recognized within the Huaqiao Formation of northwestern Hunan, China Geological map of northwestern Hunan, China, showing locations of measured stratigraphic sections near Paibi and Wangcun Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Paibi section near Paibi, Huayuan, northwestern Hunan Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Paibi-2 section near Paibi, Huayuan, northwestern Hunan Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Wangcun section, northwestern Hunan Correlation of zonal sequence for the studied interval of Huaqiao Formation in northwestern Hunan in South China with zonal schemes for the North and Northeast China Platform, Western Zhejiang in East China, and Tarim in Northwest China, and the zonal schemes of other continents Chatiania chatianensis Yang in Zhou et al., 1977. Reconstruction of cephalon and pygidium, and lectotype Reconstruction of cranidium and pygidium of Chiawangella hunanensis sp. nov. Type material of Wanshania wanshanensis Rong and Yang in Zhou et al., 1977 and specimens referred to the species Reconstructions of some species attributed to Parablackwelderia Kobayashi, 1942 Type material of Paradamesops jimaensis Yang in Lu et al., 1974a Parablackwelderia sp. Holotype cranidium of Protaitzehoia yuepingensis Yang in Yin and Li, 1978 Holotype cranidium of Protaitzehoia granifera Yang in Yin and Li, 1978 Holotype cranidium of Protaitzehoia subquadrata Peng, 1987 Type material of Teinistion posterocostum (Yang in Zhou et al., 1977) Reconstruction of cranidium and pygidium of Taihangshania wangcunensis sp. nov. Type material of Bergeronites hunanensis Yang in Zhou et al., 1977, transferred to Palaeadotes Opik, 1967 Reconstruction of dorsal exoskeleton of Palaeadotes hunanensis (Yang in Zhou et al., 1977) Type material of Paradamesella typica Yang in Zhou et al., 1977 Reconstruction of dorsal exoskeleton of Paradamesella typica Yang in Zhou et al., 1977 Holotype of Paradamesella peculiaris Zhou in Zhou et al., 1977 and specimens referred to the species Reconstruction of cephalon and pygidium of Paraacidaspis hunanica Jegorova in oo

Text-figure 20. Text-figure 21. Text-figure 22. Text-figure 23. Text-figure 24.

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Poletaeva, 1960 Text-figure 25. Lectotype of Coosia asiatica Mansuy, the type species of Paracoosia Kobayashi, 1938 Text-figure 26. Reconstruction of cephalon and pygidium of Paracoosia sp. cf. P. kingi Witteke, 1984 Text-figure 27. Holotype cranidium and associated paratype pygidium of Proceratopyge (Proceratopyge) truncata Yang in Zhou et al., 1977 Text-figure 28. Lectotype of Pseudoyuepingia laochatianensis Yang in Zhou et al., 1977 Text-figure 29. Reconstruction of cephalon of Torifera tuma (Yang in Zhou et al., 1977) Text-figure 30. Reconstruction of cephalon and pygidium of Liostracina bella Lin and Zhou, 1983 Text-figure 31. Lectotype of Monkaspis quadrata Yang in Zhou et al., 1977

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INTRODUCTION The Wulingshan Mountains region of northwestern Hunan and eastern Guizhou provinces, China, contains some of the most complete and fossiliferous strata known in the upper part of the Cambrian. Easy access to good exposures has helped make this area the subject of numerous systematic, biostratigraphic, lithostratigraphic, chemostratigraphic, and sequence-stratigraphic studies. A west-to-east cross-section through eastern Guizhou and northwestern Hunan reveals a paleogeographic profile representing a shelf-to-basin transect through carbonate shelf paleoenvironments of the Yangtze (or South China) Platform, the adjacent carbonate-dominated Jiangnan Slope Belt, and the more distal Jiangnan Basin (e.g., Gao and Duan, 1985; Lu and Qian, 1986; Liu et al., 1990; Pu and Ye, 1991; Peng and Babcock, 2001). In recent years, dark carbonates of the Huaqiao Formation, which represent the Jiangnan Slope Belt, have been particularly well studied as part of a larger effort to develop series and stage subdivisions within the Cambrian System. The Huaqiao Formation contains the most diverse trilobite fauna known from anywhere. More than 260 valid species occur through a stratigraphic interval of approximately 400 meters. Agnostoid trilobite species, 76 in number, were documented by Peng and Robison (2000), and many of them have intercontinental distributions and considerable utility for the correlation of strata globally. Agnostoid trilobites occurring in a section through the Huaqiao Formation of northwestern Hunan provide the key indicators for the base of the Paibian Stage and the Furongian Series, the first internationally ratified subdivisions within the Cambrian. The purpose of this two-volume set is to document the polymerid (or non-agnostoid) trilobites from the Huaqiao Formation of northwestern Hunan. A total of 196 species, some in open nomenclature, are reported, including a large number of new genera and new species. Some additional individual sclerites remain undetermined, however so the true diversity of polymerid taxa in the Huaqiao Formation is slightly greater than that reported here based on good, identifiable material. The polymerids have paleogeographic distributions ranging from regional to intercontinental. Most of the genera and some of the species are useful as biostratigraphic zonal indicators within Gondwana, and a few have utility intercontinentally. The work reported here continues and builds on earlier studies of the trilobite faunas and stratigraphy of northwestern Hunan by Peng (1984, 1987, 1990a, b, 1992), Peng and Robison (2000), Peng and Babcock (2001), and Peng et al. (1995,2001 a, 2001 b, 2001 c, 2001 d, 2001 e, 2001 f, 2004). Volume 1 of this set contains general geologic information concerning the Huaqiao Formation of northwestern Hunan, lithostratigraphic data, biostratigraphic zonation, detailed descriptions of three stratigraphic sections, and the systematics of corynexochid, lichid, and asaphid trilobites. Volume 2 contains systematic treatments of ptychopariids, eodiscids, trilobites of uncertain affinity, and some undetermined sclerites. In both volumes, considerable effort was made to reillustrate type material of many previously described taxa. In some older literature, illustrations were not always reproduced well, so the new illustrations should help to clarify taxonomic concepts.

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ACKNOWLEDGMENTS This work is the outgrowth of nearly two decades of field and laboratory work, and we are grateful to the numerous colleagues and institutions that have provided support for this effort over the years. Among them, Chen Yongan, G. Geyer, A. R. Palmer, R. A. Robison, J. H. Shergold, and Zhang Wentang (W. T. Chang), have been particularly helpful in a variety of aspects of the study. Han Yaojun, Luo Kunli, Li Jun, Li Yue, M. N. Rees, and Wang Huayu aided in collecting fossils. G. A. Wasserman helped to prepare and photograph specimens, and J. St. John helped to locate some references. Ren Yugao helped to draft figures, and K. Polak helped to prepare the manuscript for publication. Research was supported by grants to S. Peng from the National Natural Science Foundation of China (49070077, 40072003, 40023002, 40332018), the Chinese Academy of Sciences (KZCX2-SW-129), the Ministry of Technology and Science of China (2001 DEB20056, G2000077700), the State Key Laboratory of Paleobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (No. 933114), the National Geographical Society (No. 5819-96; 7151-01), the National Scholarship Council for International Studies, State Education Commission of China; and by grants to L. E. Babcock from the US National Science Foundation (EAR 9405990, 9526709, 0073089, 0106883, EAR OPP-0346829) and The Ohio State University (Seed Grant). REPOSITORIES All the specimens mentioned and described in the text, and illustrated on the text-figures and plates are reposited in the following institutions. Acronyms used to identify the repositories precede the repository names. CUGB EM (E.N.S.M) GMC GPIBo HIT MB NIGP USNM XTR

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China University of Geology (Beijing), Beijing, China l~cole des Mines, Departement des Sciences de la Terre, Universit6 de Lyon, France Geological Museum of China, Beijing, China Institut fiir Pal~iontologie, Universit~it Bonn, Germany Nanjing Institute of Geology and Mineral Resources, the Chinese Academy of Geological Sciences, Nanjing, China Museum ftir Naturikund of Humboldt-Universit~it, Berlin, Germany Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA Xinjiang Regional Surveying Team, Bureau of Geology and Mineral Resources, Ortimqi, China

PREVIOUS WORK Geologic study of the Cambrian stratigraphy in northwestern Hunan and eastern Guizhou began in the early Twentieth Century. Most of the major Cambrian lithologic units in eastern Guizhou were named during regional studies conducted by Ting (1930) and others during the 1940s (Tian, 1940; Lu, 1942, 1945; Liu, 1945; Yin et al., 1945; Wang, 1947). In western Hunan (Tian, 1940; Liu, 1945; Lu, 1956) and central Hunan (Wang and Pien, 1949; Liu, 1945), Cambrian lithologic units were investigated primarily in the search for mercury ore. During the 1960s and 1970s, regional geologic mapping and surveying conducted through Hunan and Guizhou resulted in the documentation of numerous Cambrian sections, and provided the basis for regional stratigraphic and paleogeographic syntheses. The work also provided important information about Cambrian faunas, and established the basis for comprehensive work on the biostratigraphy of the HunanGuizhou region (e.g., Qian, 1961; Egorova et al., 1963; Lin et al., 1966; Lu et al., 1974a; Yin and Li, 1978; Yang, 1978; Zhou et al., 1979, 1980; Peng and Tan, 1979; Peng, 1984, 1987, 1990a, 1990b, 1992; Dong 1990, 1991; Peng and Robison, 2000; Peng and Babcock, 2001; Peng et al., 2001 a, 200 lb, 2001c, 2001d, 2001e, 2001f, 2004; Dong and Bergstr6m, 2001). Biostratigraphic zonation through Hunan and Guizhou forms a major part of the zonal scheme applied generally through the South China tectonic block (Peng, 2000; Peng and Babcock, 2001 a; Peng et aL, 2004).

-3.

GEOLOGIC SETTING PALEOGEOGRAPHY The paleogeographic setting of northwestern Hunan and eastern Guizhou was summarized by Peng (1992) and Peng and Babcock (2001a). Major regions, defined by sedimentary facies and fauna, are the Yangtze (or South China) Platform, the Jiangnan Slope Belt, and the Jiangnan Basin (Text-figure 1). Together, these regions comprise a carbonate platform-to-basin transition. The Yangtze Platform was located tropically, and received sediments ranging from dolostones, phosphorites, and evaporites to siliciclastics such as black shales, calcareous and carbonaceous shales, siltstones, and marls. Strata representing the Jiangnan Slope Belt include calcareous shales, siltstones, and black shales. The Huaqiao Formation, which is dominated by dark carbonates, was deposited in the Jiangnan Slope Belt. Strata representing the Jiangnan Basin are dominated by dark-gray or black, thinly laminated carbonates. STRATIGRAPHY

Lithostratigraphy Strata representing the upper part of the Cambrian System in northwestern Hunan (Wulingian to lower Hunanian series of South China terminology; Peng and Babcock, 2001) were originally subdivided into four formations (Anonymous, 1974). In ascending order, the units were the Aoxi, Huaqiao, Chefu, and Bitiao formations. Peng and Robison (2000) recognized that differences between the Huaqiao, Chefu, and B itiao formations were biostratigraphically based, rather than lithostratigraphically based, and expanded the definition of the Huaqiao Formation to include intervals formerly referred to the Chefu and B itiao formations. The Aoxi Formation is a heterogeneous unit consisting mostly of dark-gray to yellowish gray thinly laminated to medium-bedded dolostone, with interbeds of black shale near the top. The Huaqiao Formation consists mostly of alternating thin beds of dark limestone and argillaceous limestone with interbeds of dolomitic limestone and intercalated light-colored, medium- to thickbedded carbonate breccia. Carbonate breccia beds are more numerous in the upper part of the Huaqiao Formation at Paibi. Only one such bed occurs at the Wangcun section. The Huaqiao Formation is inferred to represent sedimentation in a slope environment (part of the Jiangnan Slope Belt), and the distribution of breccia beds suggests that the Paibi sections represent deposition higher on the slope apron than does the Wangcun section.

Biostratigraphy Cambrian polymerid and agnostoid trilobites have markedly different paleogeographic distributions. Most polymerid genera are restricted to shelf and slope deposits of individual paleocontinents, whereas many agnostoid genera and species are cosmopolitan in open-marine deposits (Robison, 1976; Peng and Robison, 2000). As a result of the differences in distribution patterns, Robison (1976)proposed that separate sets of zones be developed for polymerid and -4-

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agnostoid trilobites, and we continue to advocate that practice. Peng and Robison (2000) documented thoroughly the agnostoid zonation of the Huaqiao Formation. Here, we provide a significantly revised zonation of polymerid trilobites through the stratigraphic interval represented by the Huaqiao Formation. The correlation and relationship between the polymerid and the agnostoid biostratigraphy is shown on Text-figure 2.

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Text-figure 2. Table showing the correlation of polymerid and agnostoid biozones recognized within the Huaqiao Formation of northwestern Hunan, China. Chronostratigraphic names in boldface a r e applicable globally, and those in plain type are applicable to South China. Biozones based on polymerid trilobites used here are identified by an eponymous species and characterized by a number of other species. Eponymous species are not necessarily confined to the zones that they represent, and the species do not necessarily have the same stratigraphic ranges outside of northwestern Hunan. Each of the zones defined here is identified by the first appearance of a single species selected for its relatively short stratigraphic range and abundance. The species or genera also have been selected for their relatively wide geographic distributions within Gondwana, wherever practical. The base of each zone is defined by the lowest stratigraphic occurrence of its eponymous species. In most, but not all cases, characterizing species are absent from zones above. The top of each zone is defined by the base of the next overlying zone. In ascending order, six zones are named for the species Dorypyge richthofeni, Pianaspis sinensis, Wanshania wanshanensis, Liostracina bella, Chuangia subquadrangulata, and Shengia quadrata.

.6-

1. Dorypyge richthofeni Zone. The lowest observed occurrences of Dorypyge richthofeni in the Huaqiao Formation are at 3.2 m above the base of the formation in the Paibi section and 33.0 m above the base of the formation in the Wangcun section. Because underlying strata of the Huaqiao and Aoxi formations are poorly fossiliferous, these occurrences may be younger than the first appearances of D. richthofeni elsewhere. In the Wangcun section, a species that characterizes the D. richthofeni Zone, Sudanomocarina sp. cf. S. changi, occurs at 1.2 m above the base of the Huaqiao Formation, so it is inferred that the D. richthofeni Zone extends downward to at least that level. D. richthofeni extends slightly into the lowermost part of the overlying Pianaspis sinensis Zone. Species that characterize the lower part of the D. richthofeni Zone include Huayuanaspis spp., Sudanomocarina sp. cf. S. changi and Maotunia spp. Species that characterize the upper part of the D. richthofeni Zone include Wangcunia wangcunensis, Fuchouia sixinensis, Fuchouia kuruktagensis, Paranomocarella parallela, and Amphoton deois. Paranomocarella fortis ranges from the upper part of the D. richthofeni Zone through much of the overlying Pianaspis sinensis Zone. The base of the D. richthofeni Zone is close to the base of the Ptychagnostus atavus Zone of agnostoid trilobite zonation. 2. Pianaspis sinensis Zone. The lowest observed occurrences of Pianaspis sinensis in the Huaqiao Formation are at 112.6 m above the base of the formation in the Paibi section and 82.1 m above the base of the formation in the Wangcun section. P. sinensis extends through approximately the lower half of the zone. Species that characterize the lower part of the P. sinensis Zone include Lisania yuanjiangensis, Parapianaspis hunanensis, Fuchouia bulba, and Amphoton alceste. Damesellids including Palaeadotes, Blackwelderia, Parablackwelderia, and Paradamesella make their first appearances in the middle to upper part of this zone. Species that characterize and range through much of the P. sinensis Zone include Fuchouia chiai, Fuchouia bulba, and Dorypyge bisulcata. Parablackwelderia jimaensis and Fuchouia oratolimbata range from the upper part of the P. sinensis Zone through much of the Wanshania wanshanensis Zone. Huzhuia paratypica, Palaeadotes hunanensis, Paradamesella peculiaris, and Baojingia subquadrata range from the P. sinensis Zone through the lower part of the Liostracina bella Zone. The base of the Pianaspis sinensis Zone is just below the base of the Goniagnostus nathorsti Zone of agnostoid trilobite zonation. 3. Wanshania wanshanensis Zone. The lowest observed occurrences of Wanshania wanshanensis in the Huaqiao Formation are at 240.5 m above the base of the formation in the Paibi section and 170.0 m above the base of the formation in the Wangcun section. W. wanshanensis extends into about the lower third of the overlying Liostracina bella Zone, although it is relatively uncommon above the W. wanshanensis Zone. Species that characterize the lower part of the W. wanshanensis Zone include Paibiella paibiensis and Dorypyge perconvexalis. Species that characterize the upper part of the W. wanshanensis Zone include Chatiania convexa, Neoglaphyraspis nitida, Damesella hunanensis, and Fenghuangella spp. Damesellids are both abundant and diverse in this zone; they comprise as much as 20% of the polymerid trilobite fossils. Neoanomocarella asiatica, Teinistion posterocostum, Luyanhaoaspis decorosa, Ajrikina hunanensis, Protaitzehoia subquadrata, Monkaspis quadrata, Protaitzehoia granifera, and Buttsia globosa are among the species that range from the W. wanshanensis Zone into the overlying Liostracina bella Zone. Torifera is common in this zone although it ranges into the lower part of the overlying zone. Paraacidaspis hunanica and Proceratopyge (Proceratopyge) fuyangensis range from the W. wanshanensis into the Chuangia subquadrangulata Zone. °7

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The base of the W. wanshanensis Zone is just below the base of the Proagnostus bulbus Zone.

4. Liostracina bella Zone. The lowest observed occurrences of Liostracina bella in the Huaqiao Formation are at 310.4 m above the base of the formation in the Paibi section and 218.3 m above the base of the formation in the Wangcun section. L. bella has not been observed to extend into the overlying Chuangia subquadrangulata Zone; its last observed occurrence is just below the base of the overlying zone. Species that characterize L. bella Zone include Liostracina bella, Oculishumardia hunania, Chatiania chatianensi, Taihanshania wangcunensis, Paradamesella typica, and Rhyssometopus zhongguoensis. Species that characterize the lower part of the L. bella Zone include Paracoosia huayuanensis, Paibianomocare paibiensis, and Luyanhaoaspis inflata. Damesellids, which range upward from the underlying W. wanshanensis Zone, are present in the L. bella zone, but they are much less numerous than they are in the W. wanshanensis Zone. Species that characterize the upper part of the L. bella Zone include Pseudoyuepingia laochatianensis, Fenghuangella liostracinala, and Meteoraspis sinensis. Placosema bigranulosum ranges from the upper part of the L. bella Zone into the lower part of the overlying Chuangia subquadrangulata Zone, and Shengia trapezia ranges from the top of the L. bella Zone through the overlying C. subquadrangulata Zone. The base of the L. bella Zone is just above the base of the Linguagnostus reconditus Zone of agnostoid trilobite zonation.

5. Chuangia subquadrangulata Zone. The lowest observed occurrence of C. subquadrangulata in the Huaqiao Formation is at 370.6 m above the base of the formation in the Paibi; strata above the L. bella Zone in the Wancun section have not been studied as part of this work. In the Paibi-2 section, C. subquadrangulata is not present, but the first appearance of Chuangia austriaca is used as a proxy for the base of the C. subquadrangulata Zone. C. austriaca first appears at 4.3 m above the base of the Huaqiao Formation in the Paibi-2 section. C. subquadrangulata has not been observed to extend into the overlying Shengia quadrata Zone. Species or subspecies that characterize the C. subquadrangulata Zone include Olenus punctatus, Proceratopyge truncata, Fenghuangella laochatiensis crassa, Prochuangia granulosa, and Stigmatoa yangziensis. The base of the C. subquadrangulata Zone is immediately above the base of the Glyptagnostus reticulatus Zone (just above the base of the Paibian Stage and Furongian Series). 6. Shengia quadrata Zone. The lowest observed occurrence of Shengia quadrata in the Huaqiao Formation is at 72.0 m above the base of the formation in the Paibi-2 section. The eponymous species has not been observed in the Paibi section, and strata above the L. bella Zone in the Wangcun section have not been studied as part of this work However, Peng (1992) discussed S. quadrata and other co-occurring trilobites from northwestern Hunan.

Chronostratigraphy In recent years, a chronostratigraphic scheme for the Cambrian comprising four series subdivisions has been undergoing development. This scheme, which is intended to be applicable globally, has evolved in concert with a developing chronostratigraphic subdivision of the rocks of South China (Peng et al., 1998, 1999, 2000; Peng, 2000, 2003; Peng and Babcock, 2001). The four-part system introduced for South China, which uses widely recognizable biostratigraphic tie-points, represents an improvement over the more traditional three-part subdivision of the Cambrian of the North China Platform, which was based largely on lithostratigraphic units. The scheme used in North China has been applied in South China, but correlations based on it can be

.8.

imprecise (Peng, 2003). Three boundaries relevant to Cambrian chronostratigraphy globally have been ratified: the lower and the upper boundaries of the system, and the base of the uppermost series. The base of the Cambrian System is marked by the first appearance datum (FAD) of the trace fossil Tricophycus pedum (Brasier et al., 1994; Landing, 1994; Gehling et al., 2001). The base of the Ordovician System, which also corresponds to the top of the Cambrian System, is marked by the FAD of the conodont Iapetognathus fluctivagus (Brasier et al., 1994; Cooper et al., 2001). The conterminous base of the uppermost Cambrian series (Furongian) and the lowermost stage of the series (Paibian), which corresponds to the base of the Hunanian Series and Waergangian Stage as used in South China, is defined by the FAD of the agnostoid trilobite Glyptagnostus reticulatus (Peng et al., in press). The Cambrian System of South China comprises four series, including one pre-trilobite series succeeded by three trilobite-dominated series (Peng et al., 1999; Peng and Babcock, 2001). The series are subdivided into a total of nine stages. The base of each Cambrian stage as used in South China is defined at a level coinciding with the FAD of a fossil distributed widely in South China or distributed globally. The chronostatigraphic framework of South China, and the biostratigraphic tie-points currently used to identify boundaries (Peng and Babcock, 2001), is listed below in descending order. The tie-point used to identify the base of each series is the same as that used to identify the base of the lowermost stage of the series. IV, Hunanian Series 9, Taoyuanian Stage (identified by FAD of lrvingella angustilimbata) 8, Waergangian Stage (identified by FAD of Glyptagnostus reticulatus) III, Wulingian Series 7, Youshuian Stage (identified by FAD of Linguagnostus reconditus) 6, Wangcunian Stage (identified by FAD of Ptychagnostus punctuosus) 5, Taijiangian Stage (identified by FAD of Oryctocephalus indicus) II, Qiandongian Series 4, Duyunian Stage (identified by FAD of Arthricocephalus chauveaui) 3, Nangaoan Stage (identified by FAD of trilobites [Tsunyidiscus, Sinodiscus]) I, Diandongian Series 2, Meishucunian Stage (identified by FAD of Paragloborilus subglobosus) 1, Jinningian Stage (identified by FAD of Trichophycus pedum) The Diandongian Series is a pre-trilobite series, and the Qiandongian, Wulingian, and Hunanian series are based on trilobite-bearing strata. Sections and criteria used for definition of the series and their stages were reviewed by Peng and Babcock (2001). The Diandongian Series and its stages (Jinningian and Meishucunian) are based on sections in eastern Yunnan Province. The Qiandongian Series, the stages of the Qiandongian Series (Nangaoan and Duyunian), and the Taijiangian Stage of the Wulingian Series, are based on sections in eastern Guizhou Province. The other stages of the Wulingian and Hunanian series are based on sections in northwestern Hunan Province. The Wangcunian and Youshuian stages of the Wulingian Series are based on strata in the Wangcun section, from which many of the specimens reported in this work were collected. The lower boundary of the Wangcunian Stage in the Wangcun section is 56.7 m above the base of the Huaqiao Formation, and the lower boundary of the Youshuian Stage in the Wangcun section is 210.5 m above the base of the Huaqiao Formation (Peng et al., 200 l e). The FAD of the agnostoid trilobite Ptychagnostus punctuosus, which marks the base of the Wangcunian Stage, occurs slightly below the FAD of the polymerid trilobite Pianaspis sinensis (eponymic species of the P. sinensis Zone). The FAD of Linguagnostus reconditus, which marks the base of the Youshuian Stage, occurs slightly below the FAD of the polymerid trilobite Liostracina bella (eponymic species of the .9-

L. bella Zone). The conterminous lower boundaries of the Hunanian Series and the Waergangian Stage, and the lower boundary of the Taoyuanian Stage, are based on strata in the section at Waergang, Taoyuan County, northwestern Hunan Province. The base of the Hunanian Series and the base of the Waergangian Stage were defined at the level of the FAD of the agnostoid trilobite Glyptagnostus reticulatus. However, the equivalent position for global chronostratigraphic purposes (conterminous base of the Furongian Series and Paibian Stage) was defined based on the FAD of G. reticulatus in the Paibi section, northwestern Hunan. That position in the Paibi section is 369.06 m above the base of the Huaqiao Formation (Peng et al., 2001a, 2004b, in press). The FAD of G. reticulatus is slightly above the FAD of the polymerid trilobite Chuangia subquadrangulata (eponymic species of the C. subquadrangulata Zone). Just like the section at Paibi, sections in the Wangcun area have great potential for defining or characterizing Cambrian stages. Peng et al. (2004a) reviewed two widely recognizable datum points, the FADs of the cosmopolitan agnostoid trilobites Ptychgnostus punctuosus and Lejopyge laevigata, in the Wangcun section and the Wangcun North section. Rich polymerid trilobite faunas in these sections enhance the importance of these sections for global chronostratigraphic correlation. MEASURED SECTIONS Specimens forming the basis of this work were collected from three measured sections in the Huaqiao Formation of northwestem Hunan. The sections lie on opposite limbs of a large undulating syncline, the Liexi-Zhuitun Syncline, which strikes to the north-northeast (Text-figure 3). The Liexi-Zhuitun syncline exposes mostly Cambrian and Ordovician formations at the surface. Two measured sections (Paibi and Paibi-2), on the southwestern flank of the syncline, are located near the village of Paibi, Huayuan County, Hunan (Text-figure 1). The third measured section, on the northeastem flank of the syncline, is about 4 km southeast of Wangcun, along the Wangcun to Luoyixi road, along the north side of the Fengtan Reservoir on the Youshui River (Text-figure 1). The Paibi-1 and Wangcun sections represent long, essentially uninterrupted intervals through the upper Aoxi and Huaqiao formations. The Paibi-2 section is a much shorter interval through the Huaqiao Formation. Dark to medium gray carbonate beds constitute the background sedimentation pattern in the Huaqiao Formation in both the Paibi and Wangcun areas. Allochthonous carbonate debris beds occur in the Paibi and Paibi-2 sections but they do not appear to interrupt the stratigraphic appearance of trilobite species (Peng et al., 2004). The Wangcun section is almost completely lacking in carbonate debris beds. The sections near Paibi have a considerably greater diversity of polymerid trilobite species than does the section near Wangcun. The diversity of agnostoid trilobite species is nearly the same in both areas. Measured sections near Paibi and Wangcun, Hunan, China, are described below and accompanied by columnar stratigraphic sections showing the ranges of polymerid trilobite species (Text-figures 4-6). In the following descriptions, numbered beds are indicated at the far left of each bed description. Bed 1 represents the stratigraphically lowest position, and beds are numbered successively in ascending order. Within each bed description collection numbers are indicated. Numbers preceded by a P or W refer to levels in meters above the base of the Huaqiao Formation in the Paibi and the Wangcun sections, respectively. Numbers preceded by a P[3 refers to level in meters above an arbitrary zero point (at the base of the massive breccia in the basal part of Bed 58) in the Paibi-2 section. Numbers within parentheses are altemate field collection numbers. Numbers preceeded by HP, Gs, and Gr refer to collections of trilobites, and numbers preceded by PC refer to collections of conodonts.

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Paibi Section (Text-figure 4) This well exposed section is located near the village of Sixincun. It was orignally measured by Chen Yongan and his colleagues of the 405 Geological Team, Jishou, northwestern Hunan, in 1981. The measured section is on the north side of the Jishou-Huayuan highway (part of the Chinese National Highway 319), about 35 km north of Jishou and 35 km south of Huayuan. Beginning just west of Sixincun, the section extends eastward approximately 1.7 km to the foot of a small hill north of Sixincun, about 1 km nortwest of Paibi. The measured interval is composed almost completely of carbonates. Exposures reveal the uppermost strata of the Aoxi Formation, which consist of dolostone intercalated with black shale, overlain by limestones (lime mudstones, wackestones, packstones, and some intercalated carbonate debris beds) of the the Huaqiao Formation. The Huaqiao Formation includes strata representing the Dorypyge richthofeni through the Chuangia subquadrangulata biozones (equivalent to the upper part of the Ptychagnostus atavus through the lower part of the Glyptagnostus reticulatus biozones in terms of agnostoid trilobite zonation). The section contains the Global Standard Stratotype-section and Point for the base of Furongian Series, which is the uppermost series of the Cambrian System, and the base of the Paibian Stage, which is the lowermost stage of the series (Peng et al., 2004). .11.

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breccia bed argillaceous limestone or dolomite and interbedded shale black shale

Huaqiao Formation of the Paibi section near Paibi, Huayuan, northwestern Hunan; see Peng and of the same section. Their agnostoid zonation is adopted in this figure. -13.

Huaqiao Formation 41. Medium gray to dark-gray, thin-bedded packstone intercalated with medium gray, thin bedded grainstone containing small siliceous concretions (with diameters of 0.5-1.0 cm). 385.50-388.5 m 40. Medium gray, thick-bedded grainstone, with a layer of thick-bedded limestone breccia at base and top. 381-385.5 m P383.5: Chuangia subquadrangu/ata, Placosema bigranulosum sp. nov.; agnostoid

Glyptagnostus reticu/atus. 39. Medium gray to dark-gray, thin-bedded packstone and grainstone containing small siliceous concretions (with diameters of 0.5-1.0 cm). 378.2-381 m P379: Placosema bigranulosum sp. nov. P378.3: agnostoids Peratagnostus obsoletus, Pseudagnostusjosepha. P378.25 (HP31)" polymerids Afghanocare truncatum, Baikadamaspis paibiensis sp. nov.,

Fenghuangel/a laochatianensis crassa, Paraacidaspis hunanica, Proceratopyge (Proceratopyge) fenghwangensis, Proceratopyge (Proceratopyge) truncata, Prochuangia granulosa, Prochuangia cf. P. leiocepha/a, Prochuangia linicispinata, Pseudomapania cylindrica, Shengia wannanensis, Stigmatoa yangziensis; agnostoid Glyptagnostus reticu/atus. 38. Dark-gray, thin-bedded packstone intercalated with several layers of dark-gray, striped grainstone and a single layer of limestone breccia. This bed is divided into 4 subbeds (total thickness 7.44 m). 38d. Dark-gray, thin-bedded packstone, intercalated with a single layer of medium gray grainstone. 376.32-378.2 m P376.4 ( HP30d ): polymerids Paibianomocare paibiensis gen. et sp. nov., P. ( Pro-

ceratopyge) fenghwangensis. P376.22 (Gr6): polymerids Acmarhachis typicalis, Chuangia subquadrangulata, P.

(Proceratopyge) fenghwangensis; agnostoids Glyptagnostus reticulatus, Peratagnostus obsoletus; conodont Furnishinafurnishi. 38c. Dark-gray, thin-bedded packstone, intercalated with two layers of dark, thin-bedded grainstone and a single layer of medium gray, thin-bedded limestone breccia (0.25 m thick). 372.62-376.32 m P375.9" agnostoid Glyptagnostus reticulatus. P375.22 (Gr5, PC59): agnostoids Agnostus inexpectans, Glyptagnostus reticulatus,

Peratagnostus obsoletus, Pseudagnostus josepha; conodonts Westergaardodina bicuspidata, Muellerodus sp. cf. M. pomeranensis. P375.15: polymerids Chuangia subquadrangulata, Placosema bigranulosum, Prochuangia cf. P. leiocephala. P375.0: polymerids Placosema bigranulosum, P. (Proceratopyge) fenghwangensis; Placosema bigranulosum sp. nov.; agnostoids Peratagnostus obsoletus, Pseudagnostus josepha. P374.9 (HP30c): polymerids Afghanocare truncatum, Chuangia subquadrangulata, Dorypyge perconvexalis, Huzhuia curvata sp. nov., Placosema bigranulosum, Proceratopyge (Proceratopyge) truncata, Prochuangia leiocephala, Prochuangia cf. P. leiocephala; agnostoids Glyptagnostus reticulatus, Agnostus inexpectans, Peratagnostus obsoletus, Pseudagnostusjosepha. P374.76 (PC56): conodont Furnishina sp. cf. F. a/am. P373.50 (PC43): eonodonts Huayuanodontus tricornis, Furnishina furnishi, F. sp. cf. F. a/am. P373.22 (Gr4): polymerid Baikadamaspis paibiensis; agnostoid Glyptagnostus .14.

reticulates. P373.0: agnostoid Peratagnostus obsoletus. P372.84 (PC35): conodonts Muellerodus pomeranensis, Phakelodus tenuis. P372.65 (PC32): conodonts Furnishina furnishi, Phakelodus tenuis. P372.62 (Gr3, PC31): polymerid Proceratopyge (Proceratopyge)fenghwangensis; agnostoid Glyptagnostus reticulatus; conodonts Furnishina furnishi, F. quadrata. P372.4: agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha. 38b. Dark-gray, thin-bedded packstone, intercalated with two layers of medium gray, thin-bedded grainstone. 371.42-372.62 m P372.12 (Gr2, PC23): polymerid Proceratopyge (Proceratopyge)fenghwangensis;

agnostoids Glyptagnostus reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha; eonodonts Westergaardodina bicuspidata, Furnishina kranzae, F. furnishi, F. quadrata, F. sp. P371.42 (Grl, PC16): polymerids Glyptagnostus reticulatus, Proceratopyge (Proceratopyge) fenghwangensis; conodont Furnishina sp. cf. F. quadrata. P371.2: polymerid P. (Proceratopyge) fenghwangensis. P370.9: agnostoids Agnostus inexpectans; Glyptagnostus reticulatus, Peratagnostus obsoletus. 38a. Medium gray, thick-bedded limestone breccia (0.66 m thick), no trilobites collected. 370.76-371.42 m 37. Dark-gray, thin-bedded packstone, intercalated several beds of light-gray, striped grainstone and two layers of medium gray, thin-bedded limestone breccia. This bed is divided into 5 subbeds (total thickness 8.76 m). 37e. Dark-gray, thin-bedded packstone, containing conodonts and rare inarticulate brachiopods, no trilobites collected. 369.53-370.76 m P370.76 (PC15): conodonts Furnishina kranzae, F. cf. F. bicarinata. P370.6: polymerid Chuangia subquadrangulata; agnostoids Agnostus inexpectans,

Peratagnostus obsoletus, Pseudagnostus josepha. P370.48(PCll): conodont F. quadrata; agnostoids Glyptagnostus reticulatus,

Peratagnostus obsoletus. P370.4: agnostoids Agnostus inexpectans, Peratagnostus obsoletus, Pseudagnostus josepha. P370.3: agnostoids Agnostus inexpectans, Glyptagnostus reticulatus. P370.0: agnostoid Peratagnostus obsoletus. P369.3: agnostoid Glyptagnostus reticulatus. 37d. Dark-gray, thin-bedded packstone interbedded with medium gray, thin-bedded grainstone, with a layer of thin-bedded limestone breccia near the top of this bed. 367.76-369.53 m P369.06 (Gr8): polymerids Baikadamaspis paibiensis, Shengia wannanensis; agnostoids Acmarhachis typicalis, Glyptagnostus reticulatus, Idolagnostus agrestis?,

Peratagnostus obsoletus. P368.8: polymerid: P. ( Proceratopyge ) fenghwangensis; agnostoids Agnostus inex-

pectans, Peratagnostus obsoletus, Pseudagnostus josepha. P368.49 (Gs7): agnostoids Aspidagnostus lunulosus, Glyptagnostus stolidotus, Peratagnostus obsoletus, Pseudagnostus josepha. P368.20 (PC 10): eonodonts Gapparodus bisulcatus, Phakelodus tenuis. P368.0(Gs6): polymerids Agnostardis amplinatis, Fenghuangella liostracinala, Teinistion posterocostum; Palaeadotes sp., P. (Proceratopyge) fenghwangensis; • 15"

agnostoids

Glyptagnostus stolidotus, Lisogoragnostus sp., Peratagnostus obsoletus, Pseudagnostus josepha. P367.76 (Gs5): polymerids Fenghuangella liostracinala, Liostracina bella, Peratagnostus obsoletus, Proceratopyge fenghwangensis; agnostoids Acmarhachis typicalis, Agnostardis amplinatis, Agnostus inexpectans, Glyptagnostus stolidotus, Pseudagnostus josepha. 37c. Dark-gray, thin-bedded packstone, interbedded with light-gray, thin-bedded grainstone. 366.36-367.76 m P367.50 (HP30a): polymerid P. (Proceratopyge) fenghwangensis; agnostoids

Aspidagnostus lunulosus, Glyptagnostus stolidotus, Peratagnostus obsoletus, Pseudagnostus josepha. P367.30 (PC19)" eonodonts Westergaardodina elegans, Furnishina furnishi, F. sp. cf. F. polonica. P367.26 (Gs4): polymerids Chatiania chatianensis, Pseudoyuepingia laochatianensis; agnostoids Agnostardis amplinatis, Glyptagnostus stolidotus, Peratagnostus obsoletus, Pseudagnostus josepha. P367.0: polymerid P. ( Proceratopyge ) fenghwangensis; agnostoid Agnostus inexpectans. P366.86 (Gs3): agnostoids Glyptagnostus stolidotus, Pseudagnostus josepha. P366.56 (Gs2): agnostoids Agnostardis amplinatis, Glyptagnostus stolidotus. P366.40 (PC23): conodonts Westergaardodina elegans, W. sp.,Furnishina quadrata, F. furnishi, F. kranzae, F. sp. cf. F. alata, F. sp., Phakelodus tenuis, P. elongatus, Paibiconus proacuatus. P366.36(Gs 1)" agnostoids Glyptagnostus stolidotus , Pseudagnostus josepha. 37b. Dark-gray, thin-bedded packstone, with a layer of trilobite-bearing grainstone of 5 cm thick at the base, and a layer of thick bedded limestone breccia (0.6 cm thick) in the middle part. 364.2-366.36 m P366.08(PC27): eonodonts Westergaardodina elegans, Gapparodus bisulcatus, Phakelodus tenuis, P. elongatus, Furnishina sp. P364.9: agnostoids Agnostus inexpectans, Glyptagnostus stolidotus. P364.5" agnostoids Glyptagnostus stolidotus, Pseudagnostus josepha. P364.30(PC37): polymerids Paradamesella typica, Paradistazeris hunanensis?; agnostoid Glyptagnostus stolidotus; eonodonts Westergaardodina matsushitai, W. grandidens, Furnishina quadrata, F. furnishi, F. sp. cf. F. alata, Phakelodus tenuis,

P. elongates. P364.20 (Gs0): polymerids

Paradamesella typica, Protaizehoia yuepingensis, Pseudoyuepingia laochatianensis, Teinistion posterocostum; agnostoids Glyptagnostus stolidotus, Lisogoragnostus sp., Peratagnostus obsoletus, Pseudagnostus josepha.

37a. Darkgray, thin-bedded packstone. 362.0--364.2 m P363.9: agnostoids Glyptagnostus stolidotus, Agnostus inexpectans. P363.7: agnostoids Agnostardis amplinatis, Glyptagnostus stolidotus, Pseudagnostus

josepha; P363.5 (HP30, GS- 1): polymerids Fenghuangella liostracinala, Luyanhaoaspis decorosa, Metopotropis sinensis sp. nov., Neoanomocarella asiatica, Placosema bigranulosum sp. nov., Pseudoyuepingia laochatianensis, Pseudagnostus prolongus, Rhyssometopus zhongguoensis, Teinistion posterocostum; agnostoids

Agnostogonus incognitos, Acmar~hachis apicula, Agnostardis amplinatis, Ammagnostus histus, Aspidagn'ostus lunulosus, Glyptagnostus stolidotus, Peratagnostus .16-

obsoletus, Pseudagnostus josepha. P362.45 (HP29g): polyrnerids Chatiania chatianensis, Paradistazeris hunanensis?, Pseudoyuepingia laochatianensis; agnostoids Agnostardis amplinatis, Agnostardis

typicalis, Glyptagnostus stolidotus, Peratagnostus obsoletus. P362.4: agnostoids Agnostardis amplinatis, Pseudagnostus koerferi. 36. Dark-gray, thin-to medium-bedded packstone, interbedded with 10 layers of medium gray, medium- to thick-bedded limestone breccia (single layer ranging up to 90 cm thick); one such layer (about 4 cm thick) occurs at base and top of this bed. 345.60-362.0 m P361.6(H29c): polymerid Paradamesella typica; agnostoids Ammagnostus histus,

Glyptagnostus stolidotus, Peratagnostus obsoletus. P361.5 (H29b, PC46): polymerids Chatiania chatianensis, Liostracina bella, Pseudoyuepingia laochatianensis, Rhyssometopus zhongguoensis, Shengia trapezia, Teinistion posterocostum; agnostoids Agnostogonus incognitus, Glyptagnostus stolidotus, Pseudagnostus koerferi; eonodonts Westergaardodina tetragonia, W. elegans, W. sp., Muellerodus pomeranensis, Phakelodus tenuis, Gapparodus bisulcatus, Prosagittodontus dunderbergiae, Furnishina quadrata, F. kranzae, F. bigeminata, F. furnishi, F. sp. cf. F. alata, F. sp. P361.12 (PC50): conodonts Phakelodus tenuis, P. elongates. P357.10 (PC67): eonodonts Furnishina furnishi, Huayuanodontus tricornis, Paibicornus proarcuatus, Phakelodus tenuis, P. elongates. P356.5 (HP29a): polymerids Chatiania chatianensis, Proceratopyge sp., Rhyssometopus zhongguoensis; agnostoid Peratagnostus obsoletus. P353.7(HP29-1): polymerids Ajrikina hunanensis, Chatiania chatianensis, Fenghuangella liostracinala, Palaeadotes hunanensis, Paradamesella typica, Pseudoyuepingia laochatianensis; agnostoids Agnostogonus incognitus, Peratagnostus obsoletus. P353.64: polymerids Palaeadotes hunanensis, Pseudoyuepingia laochatianensis, Teinistion posterocostum; agnostoid Peratagnostus obsoletus. P351.15 (PC74): eonodonts Westergaardodina elegans, W. sp., Furnishina furnishi, F. sp. cf. F. alata, Huayuanodontus tricornis, Phakelodus tenuis, P. elongates. P348.0(HP29): polymerids Fenghuangella laochatianensis, Liostracina bella, Neoanonocarella asiatica, Palaeadotes hunanensis, Paradamesella typica, Protaitzehoia granifera, Protaizizehoia sp., Rhyssometopus zhongguoensis, Teinistion posterocostum; agnostoid Ammagnostus sinensis. P346.7 (HP28i): polymerids Chatiania chatianensis, Fenghuangella laochatianensis, Liostracina bella, Luyanhaoaspis decorosa, Protaitzehoia granifera. 35. Dark-gray, thin- to medium-bedded, laminated packstone with limestone concretions (5-10 cm by 2-5 cm in size), no trilobites collected. 344.6-345.6 m 34. Dark-gray, thin-bedded packstone, intercalated with medium gray, thin-bedded, laminated packstone and dolomitic grainstone, and five layers of thin- to medium-bedded limestone breccia (0.2-0.5 m thick), varying in thickness laterally. 330.3-344.6 m P344.6 (HP28h): polymerids Neoanomocarella asiatica, Palaeadotes hunanensis, Para-

damesella typica, Pseudoyuepingia laochatianensis, Rhyssometopus zhongguoensis; agnostoid Pseudagnostus koerferi. P341.8 (HP28f): polymerids Adelogonus sp., Baikadamaspis sp., Chatiania cf. C. chatianensis, Iniotoma porosus sp. nov., Liostracina bella, Luyanhaoaspis inflata sp. nov., Palaeadotes hunanensis, Parablackwelderia jimaensis, Protaitzehoia spinifera sp. nov.; agnostoid Ammagnostus histus. P341.7" polymerids Chatiania chatianensis, Monkaspis quadrata, Liostracina bella; .17.

agnostoid Pseudagnostus koerferi. P337.5 (HP28e): polymerids Chatiania chatianensis, Chatiania sp. cf. C. chatianensis, Eymekops? sp. 2, Fenghuangella laochatianensis, Liostracina bella, Protaitzehoia yuepingensis, Paracoosia huayuanensis sp. nov., Prodamesella punctata, Protaitzehoia spinifera sp. nov. P332.0: agnostoids Ammagnostus? cryptus, ?Ammagnostus sinensis, Hadragnostus mo-

destus, Pseudagnostus koerferi. P331.8 (HP28d): polymerids Adelogonus hunanensis sp. nov., Baojingia subquadrata, Chatiania chatianensis, Damesella sp., Dorypyge perconvexalis?, Fenghuangella laochatianensis, Gaoloupingia gaoloupingensis, Huayuanella paibiensis gen. et sp. nov., Huzhuia paratypica, Liostracina bella, Luyanhaoaspis decorosa, Neoano-

mocarella asiatica, Palaeadotes hunanensis, Paraacidaspis hunanica, Paradamesella peculiaris, Protaitzehoia subquadrata, Taihangshania wangcunensis sp. nov., Torifera tuma; agnostoids Hadragnostus modestus, Kormagnostus minutus, Pseudagnostus

l¢oerferi. 33. Thick-bedded limestone breccia (1.30 m thick), no trilobites collected. 329.0-330.3 m 32. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, laminated packstone and four layers of light-gray, thin-bedded, dolomitic graintone, and a single layer of gray, thin-bedded limestone breccia (0.2 m thick). 322.0-329.0 m P327.4: polymerids Palaeadotes hunanensis, P. (Proceratopyge) fuyangensis, Protai-

tzehoia subquadrata. P326.9 (HP28b-1, HP28c): polymerids Luyanhaoaspis inflata sp. nov., Oculishumardia

hunania, Palaeadotes hunanensis, P. (Proceratopyge) fuyangensis, Protaitzehoia subquadrata, Teinistion posterocostum; agnostoid Pseudagnostus koerferi. P325.7 (HP28b): polymerids Chatiania chatianensis, Luyanhaoaspis decorosa, Paradamesella typica, Protaitzehoia subquadrata, Wanshania wanshanensis; agnostoid Pseudagnostus koerferi. 31. Thick-bedded breccia (1.0 m thick), no trilobites collected. 321.0-322.0 m 30. Dark-gray, thin-bedded packstone, intercalated with gray, thin-to thick-bedded, laminated packstone beds and laminated dolomitic grainstone beds, and eight layers of thin-bedded limestone breccia (0.15-1.102 m thick), most of which vary in thickness laterally. 298.1-321.0 m P319.8(HP28a-1):polymerids Chatiania sp. cf. C. chatianensis, Buttsia globosa, Eymekops? sp. 1, Huzhuia paratypica, Lobocephalina sinensis sp. nov., Liostracina

bella, Neoanomocarella asiatica, Palaeadotes hunanensis, Paracoosia huayuanensis sp. nov., Paradamesella peculiaris, Protaitzehoia subquadrata, Pseudomapania cylindrical, Taihangshania wangcunensis, Teinistion posterocostum; agnostoids Agnostogonus incognitus, Linguagnostus reconditus, Pseudagnostus koerferi. P319.6 (HP28a-0): polymerids Afghanocare truncatum, Baikadamaspis linearis sp. nov., Buttsia globosa, Chatiania chatianensis, FenghuangeUa laochatianensis, Huzhuia paratypica, Luyanhaoaspis decorosa, Monkaspis quadrata, Neoanomocarella asiatica, Palaeadotes hunanensis, Paraacidaspis hunanica, Paracoosia huayuanensis sp. nov., Paradamesella peculiaris, Paradistazeris hubeiensis, P. (Proceratopyge) fuyangensis, Proceratopyge sp. indet., Prodamesella punctata, Prodamesella sp. cf. P. biserrata, Protaitzehoia granifera, Teinistion posterocostum, Undetermined librigena 1; agnostoids Ammagnostus hunanensis, Aspidagnostus laevis, Pseudagnostus koerferi. P317.4 (HP28a): polymerids Chatiania chatianensis, Chiawangella hunanensis sp. nov., Liostracina bella, Meteoraspis sp. cf. M. orientalis, Onchonotellus curvitensus sp. nov., Paradamesella peculiaris, Paracoosia huayuanensis sp. nov., Protaitzehoia subquadrata, P. (Proceratopyge)fuyangensis, Teinistion posterocostum, Torifera tuma; .18.

agnostoids Nahanna gnostus ng anasanicus, Pse uda gno stus koe rfe ri. P316.1(HP28-4): polymerids Baikadamaspis linearis sp. nov., Huzhuia paratypica; agnostoid Agnostogonus incognitus. P316.0 (HP28-3): polymerids Liostracina bella, Paraacidaspis hunanica, Palaeadotes hunanensis, Teinistion posterocostum, Wanshania wanshanensis. P315.8: polymerid Wanshania wanshanensis. P311.7" polymerids Chatiania chatianensis. P310.4 (HP28-2): polymerids Liostracina bella, Wanshania wanshanensis; agnostoids Hadragnostus modestus, Kormagnostus minutus, Proagnostus bulbus sinensis. P308.0(HP28-1): polymerids Baikadamaspis linearis sp. nov., Chatiania expansa; Fenghuangella laochatianensis, Palaeadotes hunanensis, Taihangshania wangcunensis sp. nov.; agnostoids Hadragnostus modestus, Kormagnostus minutus, Linguagnostus reconditus, Nahannagnostus nganasanicus, Proagnostus bulbus bulbus, Proagnostus bulbus sinensis. P307.4 (HP28): polymerids Baojingia subquadrata, Buttsia globosa, Darnesella? sp., Dorypyge perconvexalis, Palaeadotes hunanensis, Protaitzehoia granifera, Wanshania wanshanensis; agnostoids Ammagnostus? cryptus, Kormagnostus minutus. P305.4 (HP27i): polymerid Wanshania wanshanensis. P301.9(HP27h):polymerids Baojingia subquadrata, Buttsia globosa, Parablackwelderia jimaensis, Luyanhaoaspis decorosa, Neoanomocarella asiatica, Palaeadotes hunanensis, Paradistazeris hunanensis, Wanshania wanshanensis, Yangweizhouia carinata, Undetermined librigena 1. P301.0 (HP27g): polymerids Baikadamaspis linearis sp. nov., Baikadamaspis sp. cf. B. granulosa, Baojingia subquadrata, Fuchouia oratolimba, Palaeadotes hunanensis, Huzhuia paratypica; agnostoid Linguagnostus reconditus. P300.4 (HP27f): polymerids Baojingia subquadrata, Palaeadotes hunanensis, Wanshania wanshanensis, P. (Proceratopyge) fuyangensis. P298.54: polymerids Huayuanella gen. nov. and sp. indet., Luyanhaoaspis decorosa, L. sp. cf. L. inflata sp. nov., Monkaspis quadrata, Neoglaphyraspis nitida, Parablackwelderia jimaensis, Protaitzehoia subquadrata, Wanshania wanshanensis; agnostoids Hadragnostus modestus, Kormagnostus minutus. P298.4 (HP27d/HP27e): polymerids Baikadamaspis linearis sp. nov., Baojingia subquadrata, Buttsia globosa, Iniotoma porosus sp. nov., Neoglaphyraspis nitida, Luyanhaoaspis decorosa, Meropalla bella, Monkaspis quadrata, Neoanomocarella asiatica, Palaeadotes hunanensis, Paradamesella peculiaris, Paradistazeris hunanensis, P. (Proceratopyge) fuyangensis, Protaitzehoia subquadrata, Qiandongaspis sinensis. 29. Dark-gray, thin- to medium-bedded, laminated, dolomitic packstone bearing limestone concretions, intercalated with three layers of light-gray, medium-bedded grainstone. 293.5-298.1 m P296.4: agnostoid Hypagnostus parvifrons. P295.13: polymerids Palaeadotes hunanensis, Parablackwelderia jimaensis, Paracoosia cf. P. kingi, Wanshania wanshanensis. 28. Medium to dark-gray, thin-bedded packstone, intercalated with numerous dark-gray, thin-bedded, laminated argillaceous limestone beds, and two intercalated layers of gray, medium-bedded limestone breccia. 284.8-293.5 m P293.2: polymerids Parablackwelderia jimaensis, Palaeadotes hunanensis, Wanshania wanshanensis; agnostoids Goniagnostus fumicola, Hadragnostus modestus, Kormagnostus minutus, Proagnostus bulbus bulbus. -19-

P293.0(HP27-8):polymerids Fenghuangella coniforma, Wanshania wanshanensis; agnostoids Goniagnostusfumicola, Linguagnostus reconditus. P292.5" agnostoid Proagnostus bulbus bulbus. P290.5: polymerids Damesella hunanensis, Dorypyge? sp., Luyanhaoaspis inflata sp. nov., Palaeadotes hunanensis, Parablackwelderia jimaensis, Wanshania wanshan-

ensis. P289.4: agnostoids Hadragnostus modestus, Kormagnostus minutus. P288.0: agnostoid Lisogoragnostus mictus. P287.1 (HP27c): polymerids Palaeadotes hunanensis, Wanshania wanshanensis; agnostoid Lejopyge sinensis. P286.3 (HP27-7): polymerids Adelogonus hunanensis sp. nov., Fuchouia oratolimba, P. (Proceratopyge) fuyangensis, Wanshania wanshanensis; agnostoids Ammagnostus hunanensis, Goniagnostusfumicola, Hypagnostus brevifrons, Valenagnostus imitans. P285.0(HP27-6):polymerids Fuchouia oratolimba, Parablackwelderia jimaensis; agnostoids Glaberagnostus altaicus, Goniagnostus fumicola, Lejopyge sinensis, Linguagnostus paibiensis, Linguagnostus stenus. 27. Base and top with a single layer of thick-bedded limestone breccia (1.9 m and 1.2 m thick respectively), interbedded with dark-gray, thin-bedded, trilobite-bearing packstone layers. 279.6-284.8 m P283.75: polymerids Buttsia globosa, Dorypyge perconvexalis, Fuchouia oratolimba, Parablackwelderia jimaensis, Wanshania wanshanensis. P283.67: polymerids Adelogonus hunanensis, Parablackwelderia jimaensis. P283.47: polymerids Dorypyge perconvexalis (?), Fuchouia oratolimba, Parablackwelderia jimaensis, Wanshania wanshanensis; agnostoid Aspidagnostus laevis. P283.0(HP27b): polymerid Fuchouia oratolimba; agnostoids Aspidagnostus laevis, Hypagnostus brevifrons, Hypagnostus parvifrons. P282.8" agnostoids Acmarhachis typicalis, Linguagnostus stenus. P282.75"polymerids Dorypyge perconvexalis(?), Luyanhaoaspis decorosa, Madarocephalus orientalis sp. nov., Palaeadotes hunanensis, Parablackwelderia jimaensis, Protaitzehoia granifera, Wanshania wanshanensis; agnostoid Proagnostus bulbus bulbus. P282.6(HP27a): polymerids Ajrikina hunanensis, Fuchouia oratolimba, Monkaspis quadrata, Palaeadotes hunanensis, Paradamesella peculiaris, Teinistion posterocostum, Torifera tuma, Wanshania wanshanensis, Undetermined pygidium 1; agnostoids Hadragnostus modestus, Hypagnostus brevifrons, Hypagnostus parvifrons, Lisogoragnostus rnictus, Quadrahornagnostus tienshifuensis, Valenagnostus imitans. 26. Dark-gray, thin-bedded, laminated, dolomitic packstone with dark to medium gray, intercalated with light-gray, thin-bedded grainstone, and a layer of medium gray, medium-bedded breccia (0.4 m thick). 260.0-279.6 m P279.7: polymerids Fuchouia oratolimba, Palaeadotes hunanensis, Parablackwelderia jimaensis; agnstoids Acmarhachis typicalis, Glaberagnostus bituberculatus. P279.55 (HP27-5): polymerids Fuchouia oratolimba, Huzhuia latilimbata sp. nov.; agnostoids Glaberagnostus altaicus, Hypagnostus parvifrons, Lejopyge laevigata, Valenagnostus imitans. P279.0 (HP27-4): polymerids Fuchouia oratolimba, Wanshania wanshanensis; agnostold Lejopyge laevigata. P278.3" agnotoid Hypagnostus parvifrons. P278.1" polymerids Fuchouia oratolimba, Parablackwelderia jimaensis, Wanshania wanshanensis; agnostoids Goniagnostus spiniger, Hypagnostus parvifrons, • 20.

Valenagnostus imitans. P278.0: polymerids Fuchouia oratolimba, Fenghuangella coniforma, Wanshania wanshanensis; agnostoids Goniagnostus spiniger, Hypagnostus brevifrons, Hypagnostus

parvifrons, Proagnostus bulbus bulbus, Valenagnostus imitans. P277.6 (HP27-3a): polymerids Fuchouia oratolimba, Wanshania wanshanensis; agnostoids Hypagnostus brevifrons, Lejopyge laevigata, Lisogoragnostus mictus. P277.0 (HP27-3): polymerids Ajrikina hunanensis, Baojingia subquadrata, Damesella

hunanensis, Dorypyge perconvexalis, Fuchouia oratolimba, Luyanhaoaspis decorosa, Neoglaphyraspis nitida, Paranomocarella? obvia sp. nov., Glyphaspellus? sinensis, Madarocephalus orientalis sp. nov., Paibianomocare paibiense gen. et sp. nov., Parablackwelderia jimaensis, Prodamesella punctata, Protaitzehoia subquadrata, Wanshania wanshanensis, Torifera tuma, Undetermined hypostome 2; agnostoids Acmarhachis typicalis, Ammagnostus laiwuensis, Clavagnostus trispinus, Glaberagnostus altaicus, Goniagnostus spiniger, Hypagnostus parvifrons, Linguagnostus stenus, Lisogoragnostus mictu.s, Tomagnostella sulcifera, Valenagnostus imitans, agnostoid gen. and sp. indet. P276.2 (HP27-2): polymerids Dorypyge perconvexalis, Neoglaphyraspis nitida; agnostoids Ammagnostus laiwuensis, Goniagnostus spiniger, Hypagnostus brevifrons. P275.1 (HP27-1): polymerids Palaeadotes hunanensis, Paradamesella peculiaris, Parablackwelderia jimaensis, Wanshania wanshanensis; agnostoid Ammagnostus

laiwuensis. P273.8 (HP27-0): polymerids

Baojingia subquadrata, Dorypyge perconvexalis, Fuchouia oratolimba, Luyanhaoaspis decorosa, Madarocephalus orientalis sp. nov., Paibianomocare paibiense gen. et sp. nov., Parablackwelderia jimaensis, Paranomocarella? obvia sp. nov., Wanshania wanshanensis, Undetermined pygidium 2; agnostoids Acmarhachis typicalis, Clavagnostus trispinus, Hypagnostus parvifrons, Proagnostus bulbus bulbus, Valenagnostus imitans. P273.66: polymerids Baojingia subquadrata, Fuchouia oratolimba, Madarocephalus orientalis sp. nov., Parablackwelderia jimaensis, Wanshania wanshanensis; agnostoid Valenagnostus imitans. P273.6: polymerid Fuchouia oratolimba. P273.52: polymerids Dorypyge perconvexalis, Fuchouia oratolimba, Wanshania wanshanensis; agnostoid Ammagnostus laiwuensis. P273.46:polymerids Fuchouia oratolimba, Madarocephalus orientalis sp. nov., Teinistion posterocostum, Wanshania wanshanensis. P269.0 (HP27): polymerids Baojingia jiudiantangensis, Blackwelderia ? sp., Fuchouia oratolimba, Paibianomocare paibiense gen. et sp. nov., Paranomocarella? obvia sp. nov., Protaitzehoia sp., Qiandongaspis sinensis, Wanshania wanshanensis, Undetermined pygidium 2; agnostoids Acmarhachis typicalis, Ammagnostus hunanensis, Hypagnostus parvifrons, Lejopyge laevigata, Valenagnostus imitans. P268.3 (HP26b): polymerids Fuchouia oratolimba, Madarocephalus orientalis sp. nov., Qiandongaspis sinensis, Parablackwelderia jimaensis, Prodamesella sp. cf. P. biserrata, Schmalenseeia fusilis sp. nov., Torifera tuma; agnostoids Ammagnostus laiwuensis, Goniagnostus spiniger, Hadragnostus modestus, Hypagnostus parvifrons, Lejopyge laevigata, Tomagnostella sulcifera. P265.1 (HP26-3): agnostois Hypagnostus parvifrons, Valenagnostus imitans. P264.2 (HP26-2): polymerids Baojingia subquadrata, Wanshania wanshanensis. P263.85: polymerid Wanshania wanshanensis. P262.4: agnostoid Tomagnostella sulcifera. -21-

P261.5 (HP26-1): polymerids

Dorypyge perconvexalis, Fuchouia oratolimba, Wanshania wanshanensis, Qiandongaspis sinensis; Undetermined pygidium 1, agnostoids Ammagnostus laiwuensis, ?Clavagnostus repandus, Clavagnostus trispinus, Oidalagnostus changi, Hypagnostus parvifrons. P261.35 (HP26-0): polymerids Baojingia quadrata, Dorypyge perconvexalis, Madarocephalus orientalis sp. nov., Neoanomocarella asiatica, Paraacidaspis hunanica ?, Paracoosia cf. P. kingi, Fuchouia oratolimba, Parablackwelderia jimaensis, Qiandongaspis sinensis; agnostoids Goniagnostus spiniger, Hadragnostus modestus, Clavagnostus trispinus, Oidalagnostus changi, Proagnostus bulbus bulbus, Tomagnostella sulcifera. P261.0(HP26 ):polymerids Baojingia youshuiensis, Fuchouia oratolimba, Qiandongaspis sinensis, Torifera tuma, Wanshania wanshanensis; agnostoids Helepagetia bitruncula. P260.1" agnostoids Goniagnostus spiniger, Lisogoragnostus hybus. P260 (HP25b): polymerids Baojingia subquadrata, Fuchouia oratolimba, Parablackwelderia jimaensis, Paracoosia cf. P. kingi, Qiandongaspis sinensis, Torifera tuma; agnostoid Oidalagnostus changi.

25. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, laminated packstone and 5 layers of medium gray, medium-bedded limestone breccia (each about 0.4 m thick). 223.5-260.0 m P259.85: polymerids Madarocephalus orientalis sp. nov., Parablackwelderia jimaensis; agnostoids Clavagnostus trispinus, Hypagnostus parvifrons, Lisogoragnostus hybus,

Proagnostus bulbus bulbus, Tomagnostella sulcifera. P256.0: Hypagnostus parvifrons. P251.0: polymerid Baojingia subquadrata; agnostoids Hypagnostus parvifrons,

Lisogoragnostus hybus. P250.0: polymerid Baojingia subquadrata. P249.0 (HP25a): polymerids Baojingia subquadrata, Baojingia youshuiensis, Dorypyge

perconvexalis, Fuchouia oratolimba, Huzhuia paratypica, Madarocephalus orientalis sp. nov., Paibiella paibiensis gen. et sp. nov., Paradamesella nobilis, Prodamesella punctata; agnostoids Ammagnostus laiwuensis, Diplagnostus sp., Hadragnostus modestus, Linguagnostus kjerulfi, Lisogoragnostus hybus, Oidalagnostus changi. P246.5 (HP25-1): polymerid Pianaspis sinensis; agnostoids Lejopyge laevigata, Linguagnostus kjerulfi. P240.5 (HP25): polymerids Madarocephalus orientalis sp. nov., Paibiella paibiensis gen. et sp. nov., Parablackwelderia laterilobata, Schmalenseeia sinensis, Wanshania wanshanensis; agnostoids Ammagnostus laiwuensis, Clavagnostus repandus, Diplagnostus planicauda, Hypagnostus parvifrons, Lejopyge laevigata, Linguagnostus kjerulfi. P240.0: agnostoids Oidalagnostus trispinifer, Valenagnostus imitans. P237.25 (HP24): polymerids Baojingia subquadrata; Fuchouia oratolimba (?), Parablackwelderia jimaensis; agnostoids Hypagnostus brevifrons, Hypagnostus parvifrons, Lejopyge laevigata, Lisogoragnostus hybus, Valenagnostus imitans. P230.5 (HP23b): polymerid Baojingia subquadrata; agnostoid Lejopyge laevigata. P225.5(HP23a):polymerid Baojingia subquadrata; agnostoid Diplagnostus planicauda. P223.7 (HP23-1):polyrnerids Baojingia paralala, Huzhuia paratypica; agnostoids Ammagnostus laiwuensis, Linguagnostus kjerulfi, Lisogoragnostus hybus, Valenagnostus imitans. • 22-

24. Thick-bedded limestone breccia (1.0m thick) in upper part, light-gray, medium-bedded grainstone in lower part, no trilobites found. 222.2-223.5 m 23. Dark-gray, thin-bedded, laminated packstone, intercalated with dark-gray, thin-bedded, dolomitic limestone and a layer of light-gray, medium-bedded grainstone, no trilobites found. 218.0-222.2 m 22. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, dolomitic limestone and several layers of light-gray, medium-bedded grainstone, with two layers of medium-bedded limestone breccia occurring in the lower and the middle part respectively. 199.0-218.0 m P216.0 (HP23): polymerids Baojingia paralala, Paradamesella peculiaris; agnostoid

Ammagnostus laiwuensis. P212.0: agnostoid Hypagnostus brevifrons. P209.5 ( HP22b ): polymerid Fuchouia oratolimba; agnostoids Clavagnostus repandus, Diplagnostus sp., Lejopyge laevigata, Ptychagnostus aculeatus. P208.9 (HP22-2): polymerid Fuchouia oratolimba; agnostoids Lejopyge laevigata,

Ptychagnostus aculeatus. P207.0: agnostoids Ammagnostus laiwuensis, Phalagnostus? sp. P204.5 ( HP22-1 ): agnostoid Clavagnostus trispinus. P204.4: agnostoid Lejopyge laevigata. P204.0 ( HP22a ): polymerids Baojingia tungrenensis, Baojingia paralala, Lisania paratungjenensis, Qiandongaspis sinensis, Prodamesella sp. cf. P. biserrata; agnostoids Ammagnostus laiwuensis, Hypagnostus brevifrons. P200.7(HP22):polymerids Baojingia latilimbata, Huayuania subcalva, Lisania

paratungjenensis, Parablackwelderia laterilobata, Paranomocarella fortis, Prodamesella sp. cf. P. biserrata; agnostoid Hypagnostus brevifrons. P199.9 (HP21e): polymerids Fuchouia chiai?, Parablackwelderia jimaensis, Palaeadotes hunanensis; agnostoids Lejopyge laevigata, Ptychagnostus aculeatus. 21. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, dolomitic limestone; with a layer of light-gray, medium-bedded grainstone, and a layer of medium gray, thin-bedded, limestone breccia. 189.0-199.0 m P190.7 (HP21d): polymerids Baojingia tungrenensis, Parablackwelderia cf. P. huabeiensis, Pianaspis sinensis; agnostoid Linguagnostus kjerulfi. P190.0: agnostoid Diplagnostus planicauda. P 189.4: agnostoid Ammagnostus wangcunensis. 20. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, dolomitic packstone with two layers of light-gray, thin-bedded grainstone, no trilobites found. 186.0-189.0 m 19. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, dolomitic packstone with 4 layers of medium gray, thin- to medium-bedded limestone breccia (single layer 0.15-0.4 cm thick, among which the layer at base is 0.4 m thick ) in the lower part and 5 layers of light-gray, thin- to medium-bedded grainstone in the upper part. 180.3-186.0 m P184.0(HP21c ): polymerids Baojingia latilimbata, Fuchouia chiai, Huzhuia paratypica, Parablackwelderia cf. P. huabeiensis, Pianaspis attenuata; agnostoids

Clavagnostus repandus, Hypagnostus brevifrons. P180.3 ( HP21b ): polymerids Dorypyge bisulcata sp. nov., Lisania paibiensis sp. nov.; agnostoids Glaberagnostus bituberculatus, Lejopyge laevigata, Linguagnostus

kjerulfi. 18. Dark-gray, thin-bedded packstone, intercalated with dark-gray, thin-bedded, laminated, • 23

•

dolomitic packstone; with 3 layers of medium gray, thin- to medium-bedded limestone breccia (single layer 0.2-0.3 m thick) in the lower part, and 7 layers of light-gray, thinbedded grainstone in the upper part. 171.0-180.3 m P178.3 (HP21a): polymerids Baojingia latilimbata, Huzhuia paraS'pica, Lisania paibiensis sp. nov., Pianaspis sinensis. P174 (HP21-2): polymerids Dor3,pyge bisulcata sp. nov., Lisania paibiensis sp. nov.; agnostoid Lejopyge laevigata. P171.5(HP21-1/HP20e):polymerids Fuchouia chiai, Huzhuia parat3'pica, Lisania paibiensis sp. nov., Parablacl,q+'elderia laterilobata, Pianaspis attenuata, Sudanomocarina sp." agnostoids Goniagnostus spiniger, ?Hypagnostus parvifrons. P171.0 (HP21): polymerids Huzhuia paras'pica, Lisania paibiensis sp. nov., Parablackwelderia laterilobata, Pianaspis sinensis, Sudanomocarina? huananensis sp. nov.; agnostoids Ammagnostus laiwuensis, Clavagnostus repandus, Goniagnostus

spiniger, ?Linguagnostus kjerulfi, Lisogoragnostus hvbus. 17.

Dark-gray, thin-bedded packstone, intercalated with medium gray dolomitic limestone. 161.0-171.0 m P 168.5 (HP20d): polymerid Pianaspis attenuata. P167.0(HP20c): polymerids Dor3'pyge bisulcata sp. nov., Fuchouia chiai, Qiandongaspis xiangxiensis sp. nov." agnostoids Goniagnostus spiniger, Hypagnostus brevi-

frons, Linguagnostus kjerulfi, Lisogoragnostus hvbus. P164.2 (HP20a): polymerids Dor3"pyge bisulcata sp. nov., Fuchouia chiai, Huzhuia paratypica, Lisania paibiensis sp. nov., Sudanomocarina? huananensis sp. nov.; agnostoid Linguagnostus kjerulfi. P163.2:agnosoids Clavagnostus trispinus, Diplagnostus planicauda, Hypagnostus

brevifrons. 16. Dark-gray, thin-bedded laminated packstone, interbedded with dark-gray laminated argillaceous dolostone, with 3 layers of light-gray, medium-bedded grainstone (2 layers in the lower part and 1 near the top) and a layer of limestone breccia in mid-upper part. 141.5-161.0 m P 160.5 (HP20-5): polymerid Lisania yuanjiangensis. P156.0 (HP20-4): polymerids Dor3'pyge bisulcata sp. nov., Dor3'pyge globosa sp. nov., Huzhuia paratypica, Lisania yuanjiangensis, Parablackwelderia sp. P149.5 (HP20-3): polymerids Huzhuia parat3'pica, Qiandongaspis convexa sp. nov., Qiandongaspis xiangxiensis sp. nov.; agnostoid Hypagnostus brevifrons. P147.0 (HP20-2): agnostoids Hypagnostus brevifrons, Hypagnostus parvifrons. P145.5 (HP20- la): polymerids Fuchouia bulba sp. nov., Lisania yuanjiangensis; agnostoids Hypagnostus parvifrons, Lejopyge laevigata. 15. Dark-gray, thin-bedded laminated packstone, interbedded with dark-gray argillacerous dolostone, and intercalated with 4 layers of thin- to medium-bedded grainstone. 133.0-141.5 m P141.0 (HP20-1): polymerid Pianaspis sinensis: agnostoids H3pagnostus parvifrons,

Lejopyge calva. P138.8: (HP20): polymerids

Huayuanaspis granulosa?, Lisania yuanjiangensis, Pianaspis sinensis. P 138.2" agnostoid Hypagnostus parvifrons. P138.0: agnostoid Hypagnostus brevifrons. P137.8" polymerids Fuchouia bulba sp. nov., Pianaspis sinensis. P136.7 (HP19g): polymerids Lisania yuanjiangensis, Pianaspis sinensis, Pagodiinae gen. et sp. indet.; agnostoid Hypagnostus brevifrons. P136.3 (HP19f): polymerids Amphoton alceste, Fuchouia bulba sp. nov., Lisania • 24-

yuanjiangensis, Paranomocarella similaris sp. nov., Pianaspis sinensis; agnostoids Hypagnostus brevifrons, Lejopyge laevigata, Tomagnostella exsculpta. P 135.7 (HP 19e): polymerids Lisania yuanjiangensis, Pianaspis sinensis, Paranomocarella similaris sp. nov.; agnostoid Pseudophalacroma ovale. P135.0(HP19c):polymerids Amphoton alceste, Pianaspis sinensis, Qiandongaspis sinensis; agnostoid Hypagnostus brevifrons. P 134.2 (HP 19b): polymerids Amphoton alceste, Fuchouia chiai, Lisania yuanjiangensis, Paranomocarella similaris sp. nov., Pianaspis sinensis, Sudanomocarina? huananensis sp. nov.; agnostoids Hypagnostus brevifrons, Pseudophalacroma ovale. 14. Dark-gray, thin-to medium-bedded laminated, argillaceous packstone bearing limestone concretions, and dark-gray laminated packstone, intercalated with 3 layers of light-gray, thin-bedded grainstone in mid-upper part, and two layers of thin-bedded limestone breccia. 126.0-133.0 m P131.7 (HP19a): polymerids Amphoton sp. cf. A. typicum, Fuchouia bulba sp. nov., Lisania yuanjiangensis, Paranomocarella fortis, Paranomocarella similaris sp. nov.,

Pianaspis sinensis; agnostoids Hypagnostus brevifrons, Tomagnostella exsculpta. P130.95"polymerids Dorypyge bisulcata sp. nov., Pianaspis sinensis; agnostoids

Hypagnostus brevifrons, Pseudophalacroma ovale. P130.5 (HP19b): polymerids Fuchouia chiai, Lisania yuanjiangensis, Paranomocarella fortis, Paranomocarella similaris sp. nov., Prodamesella tumidula sp. nov., Qiandongaspis xiangxiensis sp. nov., Sudanomocarina? huananensis sp. nov." agnostoids

Ammagnostus laiwuensis, Hypagnostus brevifrons, Tomagnostella exsculpta. P130.25: polymerids Fuchouia bulba sp. nov., Paranomocarella similaris sp. nov.; agnostoids Ammagnostus laiwuensis, Hypagnostus brevifrons, Pseudophalacroma ovale, Tomagnostella exsculpta. P130.0: agnostoids Ammagnostus laiwuensis, Hypagnostus brevifrons, Hypagnostus parvifrons, Pseudophalacroma ovale, Tomagnostella exsculpta. P126.9 (HP19): polymerids Lisania yuanjiangensis, Paranomocarella similaris sp. nov.; agnostoids Ammagnostus laiwuensis, Hypagnostus brevifrons, Hypagnostus parvifrons, Pseudophalacroma ovale. 13. Dark-gray, thin-bedded packstone, interbedded with dark-gray, thin-bedded, argillaceous dolostone, intercalated with light-gray, medium-bedded grainstone. 110.0-126.0 m P126.0(HP18a):polymerids Amphoton sp. cf. A. typicum, Baojingia tungrenensis, Dorypyge richthofeni, Huayuanaspis? sp., Pianaspis sinensis; agnostoids

Diplagnostus planicauda, Pseudophalacroma ovale, Tomagnostella exsculpta. Amphoton sp. cf. A. typicum, Dorypyge richthofeni, Fuchouia chiai, Lisania paratungjenensis; agnostoids Hypagnostus brevifrons, Hypagnostus parvifrons, Pseudophalacroma ovale. P123.6(HP17a):polymerids Amphoton sp. cf. A. ~'picum, Changqingia laevis, Dorypyge richthofeni, Qiandongaspis convexa sp. nov., Qiandongaspis xiangxiensis sp. nov., Parapianaspis hunanensis gen. et sp. nov., Paranomocarella fortis, Pianaspis sinensis, Prodamesella tumidula sp. nov.; agnostoids Ammagnostus laiwuensis, Diplagnostus planicauda, Goniagnostus nathorsti, Hypagnostus parvifrons, Pseudophalacroma ovale, Tomagnostella exsculpta. P 121.48: polymerids Dorypyge richthofeni, Lisania yuanjiangensis. P122.4 (HP17): polymerids Amphoton cf. A. typicum, Paranomocarella fortis, Pianaspis sinensis; agnostoids Hypagnostus brevifrons, Pseudophalacroma dubium, Pseudophalacroma ovale. P122 (HP16e): polymerids Dorypyge richthofeni, Pianaspis sinensis; agnostoids

P125.0(HP18):polymerids

• 25.

Ammagnostus laiwuensis, Hypagnostus brevifrons, Hypagnostus parvifrons, Pseudophalacroma ovale. Pll4(HP15e):polymerids ?Dorypyge richthofeni, Lisania sp. cf. L. agonius, Parapianaspis hunanensis. P112.6 (HP15d): polyrnerids Amphoton sp. cf. A. typicum, Dorypyge richthofeni, Paranomocarella parallela, Pianaspis sinensis, Menocephalites? sp. 2; agnostoids Diplagnostus planicauda, Hypagnostus brevifrons. P 112.4 (HP 15c): polyrnerid Changqingia laevis; agnostoid Ammagnostus laiwuensis. 12. Upper part: dark-gray, thin- to medium-bedded, laminated, dolomitic limestone, intercalated with dark-gray, thin-bedded packstone bearing limestone concretions; lower part: dark-gray, thin-bedded packstone, interbedded with dark-gray, thin- to mediumbedded, argillaceous dolostone. 93.0-110.0 m P 108.5: polyrnerid Zhujia hunanensis; agnostoid Tomagnostella exsculpta. P108.0 (HP15c): polyrnerids Amphoton alceste, Amphoton sp. cf. A. typicum, Dorypyge richthofeni, Paranomocarella fortis, Ptychopariidae gen. and sp. indet., Prodamesella tumidula sp. nov., Qiandongaspis sinensis, Zhujia hunanensis; agnostoids Ammagnostus laiwuensis, Diplagnostus planicauda, Hypagnostus parvifrons, Pseudophalacroma dubium, Pseudophalacroma ovale. P105.0: agnostoids Hypagnostus parvifrons, Pseudophalacroma ovale, Ptychagnostus punctuosus. P102.5 (HP15b): polymerid Amphoton sp. cf.A. typicum; agnostoids Pseudophalacroma dubium, Pseudophalacroma ovale, Ptychagnostus atavus. P 102.0: agnostoid Diplagnostus planicauda. P98.0 (HP15a): agnostoids Hypagnostus brevifrons, Pseudophalacroma dubium, Ptychagnostus atavus, Ptychagnostus leptus. P97.3: agnostoids Diplagnostus planicauda, Ptychagnostus leptus. P95.2 (HP 15-1c): agnostoids Ptychagnostus atavus, Ptychagnostus punctuosus. P95.0 (HP15-1b): agnostoids Diplagnostus planicauda, Hypagnostus brevifrons, Hypagnostus parvifrons, Ptychagnostus atavus, Ptychagnostus cuyanus, Ptychagnostus punctuosus, Utagnostus songae. P94.5 (HP 15-1a): agnostoid Ptychagnostus atavus. P93.74 (HP15): agnostoids Ptychagnostus atavus, Ptychagnostus punctuosus. P93.2 (HP14c): agnostoids Hypagnostus brevifrons, Ptychagnostus atavus, Ptychagnostus punctuosus. 11. Upper part: dark-gray, thin- to medium-bedded laminated, argillaceous, dolomitic limestone, interbedded with dark-gray, thin-bedded packstone. 74.0-93.0 m P92.7 (HP 14b): agnostoid Ptychagnostus atavus. P90.0: agnostoids Diplagnostus planicauda, Hypagnostus parvifrons, Ptychagnostus atavus. P85.5" agnostoid Ptychagnostus cuyanus. P84.9: agnostoid Ptychagnostus cuyanus. P84.3(HP14a):polymerid Fuchouia sixinensis sp. nov.; agnostoids Ptychagnostus atavus, Ptychagnostus cuyanus. P82.8: agnostoids Hypagnostus parvifrons, Ptychagnostus cuyanus. P82.5 (HP14): polymerids Dorypyge richthofeni, Fissanomocarella paibiensis, Fuchouia sixinensis sp. nov., Paranomocarella sp.; agnostoids Ammagnostus laiwuensis, Ptychagnostus cuyanus. P82.1 (HP 13a): polymerid Paranomocarella fortis; agnostoid Hypagnostus parvifrons. P82.0: agnostoid Ptychagnostus cuyanus. • 26"

P81.0: polymerids Dorypyge huaqiaoensis sp. nov., Changqingia intermedia; agnostoid

Hypagnostus parvifrons. P80.5 agnostoids Hypagnostus parvifrons, Ptychagnostus atavus. P79.0 agnostoid Hypagnostus parvifrons, Ptychagnostus cuyanus. P78.0 (HP 13): agnostoids Baltagnostus? ambonus, Ptychagnostus atavus. 10. Dark-gray, thin- to medium-bedded laminated, argillaceous dolostone, intercalated with dark-gray, thin-bedded packstone. 47.1-74.0 m P71.4 (HP12a): polyrnerids Amphoton deois, Fuchouia sixinensis sp. nov." agnostoid

Ptychagnostus atavus. P68.8 (HP12): polyrnerids Dorypyge richthofeni, Fuchouia kuruktagensis, Menocephalites hunanensis sp. nov. P64.5 (HP11): polyrnerids Amphoton deois, Dorypyge richthofeni, Fuchouia sixinensis sp. nov., Grandioculus obscurus sp. nov., Menocephalites hunanensis sp. nov.; agnostoids Ptychagnostus atavus, Ptychagnostus cuyanus. P57.0 (HP10d): agnostoid Ptychagnostus gibbus. P51.0: agnostoid Diplagnostus planicauda. P48.5 (HP10a): polymerids Amphoton deois, Dorypyge richthofeni, Sudanomocarina traynorae; agnostoid Ptychagnostus cuyanus. 9. Dark-gray, thin- to medium-bedded laminated, dolomitic limestone, intercalated dark-gray, thin-bedded packstone. 43.5-47.1 m P45.6 (HP9a): polymerids Amphoton deois, Crepicephalina? sp., Dorypyge richthofeni, Huayuania quadrilateralis, Huayuanaspis granulosa gen. et sp. nov., Huayuanaspis perpauca gen. et sp. nov., Paranomocarella parallela. P45.1 (HP9): polymerid Dorypyge richthofeni. 8. Dark-gray, thin-bedded laminated, dolomitic limestone, intercalated dark-gray, thin-bedded packstone. 41.0--43.5 m P43.0: agnostoids Diplagnostus planicauda, Ptychagnostus cuyanus. P42.5 (HP8a): polymerids Amphoton deois, Dorypyge richthofeni, Huayuania quadrilateralis, Mapania jiangnanensis sp. nov., Paranomocarella parallela; agnostoids

Hypagnostus parvifrons, Ptychagnostus atavus. 7. Upper part and lower part: dark-gray, thin-bedded packstone, intercalated with thin-bedded argillaceous, dolomitic limestone; middle part: medium-bedded, thin-bedded packstone and grainstone. 28.5-41.0 m P38.5 (HP8): polyrnerids Dorypyge richthofeni, Huayuanaspis granulosa gen. et sp. nov., Paranomocarella parallela, Proashaphiscus (Honanaspis)? sp. cf. P. (H.) parvus, Sudanomocarina sp. cf. S. changi; agnostoid Ptychagnostus atavus. P37.0 (HP7c): polymerids Menocephalites cf. M. pauperata, Grandioculus truncatus sp. nov., Huayuanaspis granulosa gen. et sp. nov., Parayujinia constricta gen. et sp. nov., Sudanomocarina sp. cf. S. changi. P36.0: agnostoid Diplagnostus planicauda. P35.5 (HP7b): polymerids Dorypyge richthofeni, Grandioculus truncatus sp. nov., Huayuanaspis granulosa gen. et sp. nov., Parayujinia constricta gen. et sp. nov., Proashaphiscus (Honanaspis)? sp. cf. P. (H.) parvus, Sudanomocarina sp. cf. S. changi; agnostoid Ptychagnostus affinis. P34.3" polymerid Proasaphiscus? sp. cf. P. butes. P29.0 (HP7a): polymerids Huayuanaspis granulosa gen. et sp. nov., Sudanomocarina sp. cf. S. changi; agnostoid Ptychagnostus affinis. 6. Upper part: dark-gray, thin- to medium-bedded laminated packstone; lower part: dark-gray, thin-bedded packstone intercalated grainstone. 15.2-28.5 m • 27-

P19.8 (HP7): polymerids Dorypyge richthofeni, Huayuanaspis granulosa gen. et sp. nov.;

agnostoid Ptychagnostus affinis. 5. Dark-gray, thin-bedded packstone, intercalated dark-gray laminated, argillaceous dolostone and light-gray grainstone. 12.4-15.2 P12.5 (HP6a): polymerids Do~pyge richthofeni, Kingstonia euryaxis sp. nov., Parayujinia constricta gen. et sp. nov., Sudanomocarina sp. cf. S. changi; agnostoid Ptychagnostus atavus. 4. Dark-gray, thin- to medium-bedded, laminated, argillaceous dolostone intercalated dark-gray packstone. 6.80-12.4 P7.25(HP6-1):polymerids Dorypyge richthofeni, Kingstonia eurvaxis sp. nov., Maotunia sp. cf. M. blackwelderi; agnostoids Agnostus babcocki, P~chagnostus atavus. 3. Grayish dark, thin- to medium-bedded, argillaceous dolostone, with a layer of thin-bedded dolostone with wavy lamination at the base. 2.8-6.8 P3.2 (HP6): polymerids Corynecxochus xiangxiensis sp. nov., Dorypyge richthofeni, Kingstonia euryaxis sp. nov., Sudanomocarina sp. cf. S. changi; agnostoid Ptychagnostus atavus. 2. Grayish dark, thin- to medium-bedded, laminated dolostone, interbedded dark-gray, thin-bedded packstone and black shale. 0-2.8 P0.3: agnostoids Agnostus babcocki, Hypagnostus parvifrons, Ptychagnostus atavus, Ptychagnostus gibbus; P0.05 (HP5): fragments of trilobites. Aoxi Formation 1. Light-gray, thin- to medium-bedded, laminated dolostone, intercalated with black shale in upper part. - 15.5-0

m

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Paibi-2 Section (Text-figure 5) This section is located on the east side of the rough highway connecting the Sixincun and Banlizhai villages, about 0.8 km north of Sixincun and 0.3 km south of Banlizhai. It exposes the upper Huaqiao Formation through the Lower Ordovician. Only the upper portion of the Huaqiao Formation is described here.

Huaqiao Formation 67. Medium gray, medium-bedded, dolomitic sandy grainstone. 75.0-76.0m 66. Medium gray, thick- to very thick-bedded limestone breccia, intercalated with light-gray grainstone; thickness of breccia varies greatly. 69.5-75.0 m P[3 72 (HP36): polymerids Metopotropis sinensis sp. nov., Prochuangia granulosa, Paraacidaspis hunanica, P. (Proceratopyge) truncata, Shengjia quadrata, Undetermined librigena 2, Undetermined librigena 3; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha. P[3 71.2 (HP35b): polymerid Shengia trapezia; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha. P[3 70.7" polymerids Meteoraspis cf. M. orientalis, Paraacidaspis hunanica, Shengia trapezia. P[3 70.3 (HP35a): polymerids Prochuangia granulosa, Paraacidaspis hunanica, Shengia trapezia. 65. Dark-gray, thin-bedded grainstone bearing richly siliceous small siliceous concretions, interbedded dark-gray, thin- and very thin-bedded, laminated packstone, containing small siliceous concretions. 64.8-69.5 m • 28.

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Text-figure 5. Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Paibi-2 section near Paibi, Huayuan, northwestern Hunan. PI3 69.5 (HP35): agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratag-

nostus obsoletus, Pseudagnostus josepha. P[3 65.8 (HP34b): polymerids P. (Proceratopyge) truncata, Shengia trapezia; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha. 64. Medium gray, medium- to very thick-bedded breccia (single layer 0.49-1.8 m thick), intercalated with dark-gray, very thin- to thin-bedded, laminated packstone and medium gray grainstone bearing humerous small siliceous concretions. 62.3-64.8 m PI3 64.3 (HP34a): polymerid P. (Proceratopyge) truncata; agnostoids Glyptagnostus

reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha. 63. Medium gray, thin-bedded grainstone beating abundant, small siliceous concretions, intercalated with dark-gray very thin- to thin-bedded, laminated packstone, and 4 layers of medium gray, thin- to medium-bedded limestone breccia (single layer 0.2-0.4 m). 26.6-62.3 m PI360.3 (HP33d): polymerids Baikadamaspis sp. cf. B. granulosa, Paraacidaspis hunanica, P. sp., Prochuangia granulosa, P. (Proceratopyge) truncata, Shengia • 29-

trapezia, Stigmatoa yangziensis, Sulcareclava sagitta gen. et sp. nov.; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Pseudagnostus josepha. PI3 60.15: polymerid P. (Proceratopyge) truncata. PI3 56.5 (HP33c): polymerids Olenus punctatus sp. nov., Paraacidaspis hunanica, Prochuangia granulosa, P. (Proceratopyge) truncata, Stigmatoa yangziensis; agnostoids Aspidagnostus lunulosus, A gnostus inexpectans, Pseudagnostus josepha. PI3 55.5: polymerids Paraacidaspis hunanica, Prochuangia granulosa. PI3 36 (HP33b): polymerids Baikadamaspis sinensis, Olenus austriacus, Paraacidaspis hunanica, Prochuangia granulosa, Stigmatoa yangziensis, Shengia trapezia. PI3 35.4(HP33a):polymerids Baikadamaspis sinensis, Olenus punctatus sp. nov., Paraacidaspis hunanica, P. (Proceratopyge) truncata, Shengia trapezia. • PI3 27.2(HP33): polymerids Chuangia austriaca, Olenus austriacus, Paraacidaspis hunanica, P. (Proceratopyge) truncata, Prochuangia granulosa; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Pseudagnostus josepha. 62. Medium gray, medium thick-bedded breccia. 26.0-26.6 m 61. Medium gray, thin-bedded grainstone beating small siliceous concretions. 23.4-26.0 m PI3 23.5 (HP32a): polymerids Baikadamaspis paibiensis sp. nov.(?), Chuangia austriaca,

Paraacidaspis hunanica, Prochuangia granulosa, Shengia wannanensis, Stigmatoa yangziensis; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Pseudagnostus josepha. 60. Medium gray, medium thick-to thick-bedded grainstone, intercalated with 3 layers of thick-bedded limestone breccia (single layer 0.6-1.1 m thick). 15.6-23.4 m PI3 22.4 (HP32): polymerids Baikadamaspis paibiensis sp. nov., Chuangia austriaca,

Olenus austriacus, Prochuangia granulosa, Shengia trapezia, Shengia wannanensis, Shengia sp.; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus. 59. Dark-gray, thin-bedded packstone, interbedded with 2 layers of medium gray, thinbedded limestone breccia (each layer 0.3 m thick). 9.0-15.6 m 58. Dark-gray, thin-bedded packstone and grainstone, interbedded with one layer of thick limestone breccia in upper part (1.9 m thick) and two layers of thick limestone breccia in lower part (0.8 and 1.2 m thick respectively). 0-9.0 m PI3 7.0 (HP31-4): polymerid Proceratopyge fenghwangensis; agnostoids Agnostus

inexpectans, Glyptagnostus reticulatus. PI3 5.4 (HP31-3): agnostoids Agnostus inexpectans, Glyptagnostus reticulatus. PI35.1 (HP31-2):polymerids Baikadamaspis paibiensis sp. nov., Fenghuangella liostracinala, Huzhuia curvata sp. nov., Paraacidaspis hunanica, Paibianomocare lineatum gen. et sp. nov., Shengia wannanensis. PI3 4.3 (HP3 l a): polymerids Chuangia austriaca, Paibianomocare paibiense gen. et sp. nov., Prochuangia linicispinata, Shengia wannanensis; agnostoids Agnostus inex-

pectans, Glyptagnostus reticulatus. PI] 3.3 (HP(2)-3): agnostoids Agnostus inexpectans, Glyptagnostus reticulatus. 57. Dark, thin-bedded packstone, intercalated with light to medium gray, thin-bedded -3-0 m grainstone and a layer of medium gray limestone brecia (0.4 m thick). PI]-0.3" polymerid Olenus austriacus; agnostoids Agnostus inexpectans, Glyptagnostus

reticulatus, Peratagnostus obsoletus. PI3-1.5: polymerid Olenus austriacus; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Idolagnostus sp. PI3-1.6: polymerids Olenus austriacus, Prochuangia linicispinata; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Idolagnostus sp. • 30"

PI3-1.65: polymerids Shengia wannanensis, Proceratopyge fenghwangensis; agnostoids

Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Idolagnostus sp. PI3-1.72: agnostoids Glyptagnostus reticulatus; polymerid Proceratopyge fenghwangensis. PI3-1.8: polymerids Baikadamaspis paibiensis sp. nov., Olenus austriacus, Proceratopyge fenghwangensis; agnostoids Agnostus inexpectans, Glyptagnostus reticulatus, Peratagnostus obsoletus, Idolagnostus sp. PI3-2.75: polymerids Baikadamaspis paibiensis sp. nov., Chiawangella hunanensis sp. nov., Chatiania chatianensis, Dorypyge perconvexalis, Fenghuangella liostracinala, Liostracina bella, Oculishumardia hunania, Palaeadotes hunanensis, Paradamesella typica, Paradistazeris sp., Protaitzheoia granifera(?), Rhyssometopus zhongguoensis, Teinistion posterocostum; agnostoids Agnostus amplinatis, Agnostus inexpectans, Ammagnostus sinensis, Glyptagnostus stolidotus, Lisogoragnostus sp., Nahanagnostus nganasanicus, Peratagnostus obsoletus.

Wangcun Section (Text-figure 6) This section is located about 4 km southeast of Wangcun, an historic town in southernmost Yongshun County, northwestern Hunan. The section was measured along a roadcut of the highway constructed on the north bank of the Youshui River ; the highway connects Wangcun with the Luoyixi (now renamed as Mendonghe) railway station in Guzhang County. The section is situated on the southeast limb of the Liexi-Zhuitun Syncline. It exposes carbonate formations including the middle Cambrian Aoxi Formation through the Lower Ordovician Nantsinkwan Formation. However, only the uppermost Aoxi Formation and the Ptychagnostus gibbus Zone through lower Glyptagnostus reticulatus Zone of the Huaqiao Formation are described here.

Huaqiao Formation 34. Dark-gray, thin-bedded packstone bearing small siliceous concretions. 259.0-307.0 m W288.95: agnostoid Glyptagnostus reticulatus. 33. Dark-gray, thin-bedded, packstone, interbedded with dark-gray, thin-bedded laminated packstone, both containing rarely small siliceous concretions. 230.0-259.0 m W254.1: polymerids Chatiania chatianensis, FenghuangeUa fusilis sp. nov., Liostracina bella, Lobocephalina sinensis sp. nov., Oculishumardia hunania, Neoglaphyraspis

nitida, Neoanomocarella asiatica, Paradamesella typica, Pseudoyuepingia laochatianensis, Teinistion posterocostum; agnostoids Agnostogonus incognitus, Ammagnostus? cryptus, Ammagnostus hunanensis, Glyptagnostus stolidotus, Idolagnostus agrestis?, Pseudagnostus koerferi, Tomagnostella sulcifera. W251.15: polymerids Fenghuangella liostracinala, Oculishumardia hunania, Neoanomocarella asiatica, Paradamesella typica, Taihangshania wangcunensis sp. nov.; agnostoids Agnostogonus incognitus, Ammagnostus? cryptus, Aspidagnostus lunulosus, Glyptagnostus stolidotus, Hypagnostus brevifrons, Nahannagnostus nganasanicus, Peratagnostus obsoletus, Tomagnostella sulcifera. W243.6: polymerids Fenghuangella liostracinala, Oculishumardia hunania, Liostracina bella, Neoanomocarella asiatica, Teinistion posterocostum, Paradamesella typica; agnostoid Nahannagnostus nganasanicus. 32. Dark-gray, thin- to medium packstone, intercalated with 2 layers of light-gray grainstone bearing trilobites in middle part. 227.0--230.0 m W229.9: polymerids; Neoanomocarella asiatica, Palaeadotes hunanensis, P. (Procera-

topyge )fuyangensis, Pseudomapania cylindrica, Rhyssometopus zhongguoensis?, -31-

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Formation of the Wangcun section, northwestern Hunan; see Peng and Robison (2000) for the agnostoid Their agnostoid zonation is adopted in this figure. • 33-

Teinistion posterocostum. W228.3: polymerids Ajrikina hunanensis, Fenghuangella coniforma, Fenghuangella laochatianensis, Lobocephalina sinensis sp. nov., Meropalla bella, Oculishumardia

hunania, Neoanomocarella asiatica, Palaeadotes hunanensis, P. (Proceratopyge) fuyangensis, Prodamesella punctata, Pseudomapania cylindrica, Teinistion posterocostum, Torifera abrupta sp. nov.?, Townleyella rara, Undetermined librigena 2; agnostoids Hypagnostus brevifrons, Kormagnostus minutus, Peratagnostus obsoletus, Tomagnostella sulcifera. 31. Dark-gray, thin-bedded, argillaceous packstone, interbedded with light to medium gray, thin-bedded grainstone, most of which bear trilobites. 209-227.0 m W227.0: polymerids Afghanocare truncatum, Buttsia globosa, Chatiania expansa, Chiawangella hunanensis sp. nov., Fenghuangella laochatianensis, Iniotoma laevis sp. nov., Oculishumardia hunania, Liostracina bella, Neoanomocarella asiatica, Paibianomocare paibiense gen. et sp. nov., Palaeadotes hunanensis, Paracoosia huayuanensis sp. nov., Paradamesella typica, P. (Proceratopyge) fuyangensis, Protaitzehoia granifera, Sulcareclava sagitta gen. et sp. nov., Taihangshania wangcunensis sp. nov., Teinistion posterocostum, Torifera abrupta sp. nov., Yangweizhouia carinata; agnostoids Ammagnostus hunanensis, Ammagnostus sinensis,

Ammagnostus wangcunensis, Hadragnostus modestus, Kormagnostus minutus, Proagnostus bulbus sinensis, Tomagnostella sulcifera. W225.0: polymerids Buttsia globosa, Chatiania expansa, Fenghuangella laochatianensis, Liostracina bella, Meropalla bella, Neoglaphyraspis nitida, Neoanomocarella asiatica, Palaeadotes bella, Parablackwelderia jimaensis, P. (Proceratopyge) fuyangensis, Protaitzehoia spinifera sp. nov., Pseudomapania cylindrica, Taihangshania wangcunensis sp. nov., Teinistion posterocostum; agnostoids Ammagnostus wangcunensis, Kormagnostus minutus, Proagnostus bulbus sinensis. W223.2: polymerids Buttsia globosa, Monkaspis quadrata?, Neoanomocarella asiatica, Townleyella rara; agnostoid Kormagnostus minutus. W221.5" polymerids Buttsia globosa, Fenghuangella laochatianensis, Liostracina beUa, Neoanomocarella asiatica, Neoglaphyraspis nitida, Palaeadotes hunanensis, Palaeadotes bella, Protaitzehoia spinifera sp. nov., Teinistion posterocostum; agnostoids Ammagnostus wangcunensis, Hadragnostus modestus, Kormagnostus minutus, Proagnostus bulbus sinensis, Tomagnostella sulcifera. W219.7: polymerids Ajrikina hunanensis, Buttsia globosa, Drepanura? crassispina sp. nov., Fenghuangella laochatianensis, Lobocephalina sinensis sp. nov., Neoglaphyraspis nitida, Palaeadotes bella, P. hunanensis, Paradamesella typica, Parablackwelderia jimaensis, Pseudomapania cylindrica, Teinistion posterocostum, Undetermined larva; agnostoids Acmarhachis typicalis, Ammagnostus hunanensis, Ammagnostus wangcunensis, Aspidagnostus rugosus, Kormagnostus minutus, Linguagnostus reconditus, Nahannagnostus nganasanicus, Proagnostus bulbus sinensis, Tomagnostella sulcifera. W218.3: polymerids Ajrikina hunanensis, Drepanura? crassispina sp. nov., Fenghuangella laochatianensis, Liostracina bella, Monkaspis quadrata, Neoanomocarella asiatica, Parablackwelderia jimaensis, Paradistazeris rotundus sp. nov., P. (Proceratopyge) fuyangensis, Teinistion posterocostum; agnostoids Ammagnostus wangcunensis, Kormagnostus minutus, Linguagnostus reconditus, Proagnostus bulbus sinensis. W216.5:polymerids Ajrikina hunanensis, Baojingia subquadrata, Buttsia globosa, Fenghuangella coniforma, Gaoloupingia gaoloupingensis, Neoanomocarella asiatica, • 34-

Palaeadotes bella, Paradamesella typica, Paradistazeris rotundus sp. nov., P. (Proceratopyge) fuyangensis, Pseudomapania cylindrica, Teinistion posterocostum, Torifira taoyuanensis, Wanshania wanshanensis; agnostoids Ammagnostus wangcunensis, Korrnagnostus minutus, Linguagnostus reconditus, Nahannagnostus nganasanicus, Peratagnostus obsoletus, Proagnostus bulbus sinensis. W215.1: polymerids Buttsia globosa, Monkaspis quadrata, Neoanomocarella asiatica, Palaeadotes bella, Paradistazeris hunanensis, Paradistazeris rotundus sp. nov., Qiandongaspis sinensis, Teinistion posterocostum; agnostoids Hypagnostus brevifrons, Kormagnostus minutus, Proagnostus bulbus sinensis. W213.0: polymerid Chatiania expansa. W212.45" polyrnerids Baojingia subquadrata, Fenghuangella laochatianensis crassa subsp, nov., Meropalla gibbera sp. nov., Monkaspis quadrata, Neoanomocarella asiatica, Palaeadotes hunanensis, Paracoosia sp. cf. P. kingi, Paradistazeris rotundus sp. nov., Protaitzehoia subquadrata; agnostoids Ammagnostus wangcunensis, Hadragnostus modestus, Kormagnostus minutus, Peratagnostus obsoletus, Proagnostus bulbus sinensis. W211.7: polymerids Baojingia subquadrata, Damesella hunanensis sp. nov., Fenghuangella laochatianensis crassa subsp, nov., Monkaspis quadrata, Neoglaphyraspis nitida, Palaeadotes hunanensis, Parablackwelderia jimaensis, Paradistazeris hubeiensis, P. (Proceratopyge) fuyangensis, Protaitzehoia spinifera sp. nov., Pseudomapania cylindrica, Torifira taoyuanensis, Torifera tuma, Undetermined librigena 4; agnostoids Ammagnostus wangcunensis, Hadragnostus modestus, Kormagnostus minutus, Valenagnostus imitans. W210.5: polymerids Aethia rectangular, Ajrikina hunanensis, Baojingia subquadrata, Fenghuangella laochatianensis, Monkaspis quadrata, Neoglaphyraspis nitida, Neoanomocarella asiatica, Palaeadotes hunanensis, Parablackwelderia jimaensis, P. (Proceratopyge) fuyangensis, Protaitzehoia subquadrata, Pseudomapania cylindrica, Torifira taoyuanensis; agnostoids Acmarhachis typicalis, Ammagnostus wangcunensis, Hadragnostus modestus, Hypagnostus brevifrons, Kormagnostus minutus, Linguagnostus reconditus, Nahannagnostus nganasanicus, Proagnostus bulbus sinensis, Tomagnostella sulcifera. 30. Dark-gray, medium- to thick-bedded (thickness of 0.5-1 m in single bed), laminated packstone, intercalated with light to medium gray, thin-bedded grainstone. 191.0-209.0 m W207.5: polymerids Palaeadotes hunanensis, Paradistazeris hubeiensis, Wanshania wanshanensis; agnostoids Goniagnostus fumicola, Hadragnostus modestus, Hypag-

nostus brevifrons, Kormagnostus minutus, Oidalagnostus trispinifer. W203.0: agnostoid Goniagnostus fumicola. W201.0: agnostoid Hypagnostus brevifrons. W200.7: polymerids Protaitzehoia subquadrata, P. hubeiensis. W200.3:polymerids Ajrikina wannanensis, Baojingia subquadrata, Palaeadotes hunanensis, Paradistazeris hubeiensis, Protaitzehoia spinifera sp. nov., Torifera taoyuanensis; agnostoids Goniagnostus fumicola, Hadragnostus modestus, Hypag-

nostus brevifrons, Kormagnostus minutus, Proagnostus bulbus sinensis. W199.2: polymerids Ajrikina wannanensis, Baojingia subquadrata, Fenghuangella

coniforma, Limbishumardia wangcunensis, Neoglaphyraspis nitida, Neoanomocarella asiatica, Parablackwelderia jimaensis, Paracoosia sp. cf. P. kingi, Paradistazeris hubeiensis, Palaeadotes hunanensis, Paradamesella peculiaris, Protaitzehoia subquadrata, Pseudomapania cylindrica, Pseudoyuepingia laochatianensis, Qiandongaspis sinensis, Torifira taoyuanensis; agnostoids Glaberagnostus altaicus, .35.

Goniagnostus fumicola, Hadragnostus modestus, Hypagnostus brevifrons, Kormagnostus minutus, Proagnostus bulbus bulbus, Valenagnostus imitans. W198.45: polymerids Fuchouia oratolimba, Palaeadotes hunanensis, Paradamesella peculiaris; agnostoids Goniagnostus fumicola, Oidalagnostus trispinifer, Hypagnostus brevifrons, Lejopyge sinensis, Valenagnostus imitans. W197.6: agnostoids Acmarhachis typicalis, Glaberagnostus altaicus, Goniagnostus fumicola, Hypagnostus brevifrons, Nahannagnostus nganasanicus, Valenagnostus imitans. W197.5: polymerids Ajrikina wannanensis, Buttsia globosa, Fenghuangella coniforma, Fuchouia oratolimba, Palaeadotes hunanensis, Protaitzehoia yuepingensis, Qiandongaspis sinensis, Wanshania wanshanensis; agnostoids Acmarhachis typicalis, Ammagnostus wangcunensis, Glaberagnostus altaicus, Goniagnostus fumicola, ?Hadragnostus modestus, Hypagnostus brevifrons, Lejopyge sinensis, Linguagnostus stenus, Oidalagnostus trispinifer, Proagnostus bulbus bulbus, Valenagnostus imitans. W196.3:polymerids Ajrikina wannanensis, Baojingia quadrata, Fenghuangella coniforma, Neoanomocarella asiatica, Palaeadotes hunanensis, Parablackwelderia jimaensis, Paracoosia cf. P. kingi, Paradamesella peculiaris, ProdameseUa punctata, Protaitzehoia yuepingensis, Pseudomapania cylindrica, Pseudoyuepingia laochatianensis, Qiandongaspis sp., Torifira taoyuanensis; agnostoids Glaberagnostus altaicus, Goniagnostus fumicola, Lejopyge sinensis, Linguagnostus paibiensis, Oidalagnostus trispinifer, Oidalagnostus changi. W195.7: polymerids Paracoosia cf. P. kingi, Paradamesella peculiaris, Wanshania wanshanensis; agnostoids Goniagnostus fumicola, Lejopyge laevigata, Lejopyge sinensis, Oidalagnostus trispinifer, Proagnostus bulbus bulbus. W195.3"polymerids Buttsia globosa, Palaeadotes hunanensis; agnostoids Paradamesella peculiaris, Oidalagnostus trispinifer, Proagnostus bulbus bulbus. W 194.7: agnostoids Goniagnostus fumicola, Oidalagnostus trispinifer. W 194.6: polymerid Palaeadotes hunanensis; agnostoid Proagnostus bulbus bulbus. W 194.1: agnostoid Hypagnostus brevifrons. W 193.6: polymerids Palaeadotes hunanensis, Parablackwelderia jimaensis, Qiandongaspis sinensis; agnostoid Oidalagnostus trispinifer. 29. Dark-gray, thin- to medium-bedded, laminated packstone, intercalated with light- to medium-gray, thin-bedded, striped grainstone. 173.9-191.0 m W 189.0: polymerids Baojingia quadrata, Neoanomocarella incilis sp. nov., Palaeadotes hunanensis, Paradamesella peculiaris, Parablackwelderia jimaensis, Torifera tuma. W 188.8: polyrnerids Fenghuangella coniforma, Fuchouia oratolimba, Neoanomocarella incilis sp. nov., Parablackwelderia jimaensis, Paradamesella peculiaris, Torifera tuma. W187.8:polyrnerids Ajrikina wannanensis, Madarocephalus orientalis sp. nov., Neoanomocarella incilis sp. nov., Palaeadotes hunanensis, Parablackwelderia jimaensis, Paradamesella peculiaris, Torifera tuma; agnostoids Ammagnostus laiwuensis, Glaberagnostus bituberculatus, Goniagnostus spiniger, Hypagnostus brevifrons, Lejopyge laevigata. 28. Grayish black, thin-bedded, laminated packstone, intercalated frequently with light-gray, thin-bedded grainstone (thickness ranging between 4 and 10 cm). 139.3-173.9 m W173.8: polymerids Baojingia quadrata, Torifera tuma; agnostoids Ammagnostus laiwuensis, Hypagnostus parvifrons, Linguagnostus kjerulfi, Proagnostus bulbus bulbus. • 36.

W171.9: polymerids Fuchouia oratolimba, Paradamesella peculiaris, Prodamesella sp. cf. P. biserrata, Torifera tuma; agnostoids Ammagnostus laiwuensis, Goniagnostus

spiniger, Hypagnostus parvifrons, Lejopyge multifora, Proagnostus bulbus bulbus. W170.0: polymerids Palaeadotes hunanensis, Wanshania wanshanensis; agnostoids

Clavagnostus repandus, Goniagnostus spiniger, Hypagnostus parvifrons, Linguagnostus kjerulfi, Oidalagnostus changi. W 169.1" agnostoid Linguagnostus kjerulfi. W 167.9: agnostoids Goniagnostus spiniger, Lejopyge multifora. W158.7" agnostoids Glaberagnostus bituberculatus, Lejopyge laevigata. W 151.6: agnostoid Lejopyge laevigata. W146.2: polymerids Baojingia paralala, Fuchouia sp. indet., Huzhuia latilimbata sp. nov., Huzhuia paratypica, Lisania paibiensis sp. nov., Lisania? sp., Madarocephalus orientalis sp. nov., Mapania jiangnanensis sp. nov., Prodamesella sp. cf. P. biserrata, Sudanomocarina? huananensis sp. nov.; agnostoids Ammagnostus laiwuensis, Glaberagnostus bituberculatus, Linguagnostus kjerulfi, Lejopyge laevigata, Lisogoragnostus hybus. 27. Light-gray, thin-bedded grainstone (thickness ranging between 4 and 15 cm). 138.8-139.3 m W 139.2: polymerids Lisania paibiensis sp. nov., Lisania wangcunensis sp. nov. W 139.2" polymerid Lisania paratungjenensis. W138.9: polymerids Fuchouia bulba sp. nov., Lisania paratungjenensis" agnostoid

Lejopyge laevigata. 26. Grayish black, thin-bedded, laminated packstone bearing limestone concretions. 132.9-138.8 m 25. Light-gray grainstone, interbedded with dark-gray, thin-bedded, laminated packstone. 132.0-132.9 m W132.5: polymerids Blackwelderia youshuica sp. nov., Huzhuia paratypica, Lisania paibiensis sp. nov., Parablackwelderia laterilobata, Schmalenseeia sinensis; agnostoids

Ammagnostus laiwuensis, Clavagnostus trispinus, Hypagnostus brevifrons, Lejopyge laevigata. 24. Grayish black, medium- to thick-bedded packstone bearing limestone concretions, interbedded with light-gray, thin-bedded grainstone. 122.00-132.0 m W128.9" polymerid Lisania yuanjiangensis; agnostoids Ammagnostus laiwuensis,

Lejopyge laevigata. W 125.0: agnostoid Lejopyge laevigata. 23. Light-gray, thick-bedded grainstone, interbedded with medium-bedded, laminated packstone. 118.6-122.0 m W121.6: polymerids Baojingia latilimbata, Fuchouia bulba sp. nov., Lisania yuanjiangensis, Pianaspis sinensis; agnostoid Lejopyge laevigata. 22. Grayish black, medium- to thick-bedded, argillaceous packstone bearing limestone concretions, interbedded with numerous light-gray, thin-bedded grainstone beds. 90.5-118.6 m W lll.3"polymerids Dorypyge bisulcata sp. nov., Fuchouia sp. indet., Lisania yuanjiangensis, Palaeadotes hunanensis, Pianaspis sinensis; agnostoids Hypagnostus

brevifrons, Lejopyge laevigata. W105.0: polymerids Fuchouia bulba sp. nov., Lisania yuanjiangensis, Pianaspis sinensis; agnostoids Goniagnostus nathorsti, Hypagnostus brevifrons, Pseudo-

phalacroma ovale. W101.0"polymerids

Fuchouia bulba sp. nov., Pianaspis sinensis; agnostoids Goniagnostus nathorsti, Hypagnostus parvifrons, Pseudophalacroma ovale, Tomagnostella exsculpta. • 37"

W99.3: polymerids

Fuchouia bulba sp. nov., Pianaspis sinensis; agnostoids Hypagnostus parvifrons, Lejopyge calva, Pseudophalacroma ovale, Tomagnostella exsculpta. W98.0: polymerids Fuchouia bulba sp. nov., Lisania yuanjiangensis, Pianaspis sinensis; agnostoids Hypagnostus parvifrons, Tomagnostella exsculpta. W95.8" agnostoid Pseudophalacroma ovale. 21. Light-gray, thin-bedded, fine-grained grainstone, extending laterally into lenses. 90-90.5 m 20. Grayish black, medium- to thick-bedded, laminated packstone bearing limestone concretions, interbedded with few light-gray, thin-bedded grainstone beds. 82.3-90.0 m W88.1: polymerids Fuchouia kuruktagensis, Parapianaspis hunanensis gen. et sp. nov.; agnostoids Diplagnostus planicauda, Pseudophalacroma ovale, Tomagnostella

exsculpta. 19. Grayish black, laminated, dolomitic packstone, interbedded with light-gray, thin-bedded grainstone beds, some of which extend laterally into lenses. 80.5-82.3 m W82.2: Polymerids Amphoton alceste, Dorypyge richthofeni, Fuchouia kuruktagensis" agnostoids Hypagnostus parvifrons, Pseudophalacroma lundgreni, Pseudophalac-

roma ovale, Tomagnostella exsculpta. W82.1: polymerids Amphoton cf. A. typicum, Dorypyge richthofeni, Fuchouia bulba, Fuchouia kuruktagensis, Paranomocarella similaris, Pianaspis sinensis; agnostoids

Ammagnostus laiwuensis, Diplagnostus planicauda, Goniagnostus nathorsti, Hypagnostus parvifrons, Pseudophalacroma dubium, Pseudophalacroma lundgreni, Pseudophalacroma ovale, Pseudophalacroma scanense, Tomagnostella exsculpta, Utagnostus trispinulus. 18. Grayish black, medium- to thick-bedded, laminated packstone bearing limestone concretions and light-gray, thin-bedded grainstone. 72.6-80.5 m W77.8" polymerids Dorypyge richthofeni, Fuchouia kuruktagensis; agnostoids Bal-

tagnostus? ambonus, Goniagnostus scarabaeus, Hypagnostus parvifrons, Pseudophalacroma dubium, Ptychagnostus punctuosus, Tomagnostella exsculpta. W75.7: agnostoid Pseudophalacroma dubium. 17. Grayish black, medium- to thick-bedded, laminated packstone bearing limestone concretions, intercalated with few light-gray, thin-bedded grainstone beds, with a 15 cm thick layer of loosely cemented black shale appearing at 72.6 m, and a 10 cm thick grainstone bearing rich trilobites at 56.5 m. 52.1-72.6 m W70.2: agnostoid Ptychagnostus leptus. W69.5: agnostoid Ptychagnostus leptus. W68.5: agnostoid Ptychagnostus leptus. W64.7: polymerids Dorypyge richthofeni, Fuchouia sixinensis sp. nov., Menocephalites? sp. 1; agnostoids Ptychagnostus atavus, Ptychagnostus punctuosus. W56.7: polymerids Dorypyge richthofeni, Fuchouia kuruktagensis, Geminiclavula wangcunica gen. et sp. nov., Paranomocarella fortis, Prodamesella tumidula sp. nov., Szeaspis huananensis sp. nov., Wangcunia wangcunensis; agnostoids Ammagnostus

laiwuensis, Baltagnostus? ambonus, Doryagnostus incertus, Glaberagnostus? cicer, Ptychagnostus cuyanus, Ptychagnostus punctuosus, Tomagnostella exsculpta. 16. Medium gray, thin-bedded grainstone with few laminations, interbedded with dark-gray, thin-bedded packstone. 51.3-52.1 m 15. Dark-gray, thin-bedded packstone bearing limestone concretions. 50.2-51.3 m 14. Dark-gray, thin-bedded grainstone with few laminations, intercalated with dark-gray, thin-bedded laminated packstone. 49.5-50.3 m • 38.

13. Dark-gray, thin-bedded packstone beating limestone concretions. 12. Medium gray, thin-bedded packstone with few laminations. 11. Grayish black, thin- to medium-bedded packstone bearing limestone concretions.

48.5-49.5 m 48.0-48.5 m 46.7-48.0 m

10. Grayish black, medium-bedded, laminated packstone bearing limestone concretions, intercalated with light-gray, thin-bedded grainstone. 36.4-46.7 m W42.0: polymerid Fuchouia sixinensis sp. nov" agnostoids Ptychagnostus atavus,

Ptychagnostus cuyanus, P~chagnostus gibbus. W36.6: polymerids Crepicephalina pergranosa, Sudanomocarina sp. cf. S. changi; agnostoid Ptychagnostus atavus. 9. Dark-gray, thin-bedded grainstone with few laminations, intercalated with dark-gray, thin-bedded, laminated packstone. 33.4-36.4 m W33.6: agnostoid Ptychagnostus atavus. W33.0: polymerids Dorypyge richthofeni, Sudanomocarina sp. cf. S. changi; agnostoids

Hypagnostus parvifrons, Ptychagnostus atavus. 8. Grayish black, thin-to medium-bedded, laminated packstone bearing limestone concretions, intercalated with few light-gray, thin-bedded grainstone beds. 22.5-33.4 m 7. Dark-gray, thin-bedded grainstone, intercalated with dark-gray, thin-bedded, laminated packstone. 21.6-22.5 m 6. Grayish black, medium- to thick bedded, laminated packstone bearing rare limestone concretions. 1.2-21.6 m W3.0: polymerids Corynexochina sinensis sp. nov., Maotunia sp., Parayujinia constricta gen. et sp. nov., Xilingxia ichangensis; agnostoid P~chagnostus atavus. W1.2: polymerids Crepicephalina cf. C. quadrata, Maotunia sp. cf. M. blackwelderi, Sudanomocarina sp. cf. S. changi; agnostoids Po,chagnostus atavus, P~chagnostus

affinis, Ptychagnostus atavus. 5. Black, laminated, argillaceous, dolomitic packstone and thin-bedded packstone. -0.8-1.2m W0: polymerids Corynexochina sinensis sp. nov., Maotunia sp. cf. M. semiplectra, Maotunia rectangularis sp. nov., Sudanomocarina sp. cf. S. changi; agnostoid Peronop-

sis sp. 4. Black, thin-bedded grainstone.

-1.1--0.8 m

Aoxi Formation

3. Black shale. -2.0--1.1 m 2. Grayish black, thin-bedded, dolomitic packstone, interbedded with black shale. -5.0--2.0m 1. Black shale, interbedded with a few layers of dolostone in upper part; grayish, laminated dolostone, bearing thin, belt-like layers of chert in lower part, with a layer of 30-cm-thick black chert with oolites at base. -18.5- -5.0 m

• 39"

CORRELATION Agnostoid and polymerid trilobites permit biostratigraphic correlation of the studied interval of the Huaqiao Formation in South China with slope or shelf facies of many other regions of the Cambrian world. Text-fig. 7 summarizes the correlation of biostratigraphic zones recognized here from South China with those of other regions within China (North China and Northeast China, western Zhejiang, and Tarim) and with those of other paleocontinents (Australia, Kazakhstan, Siberia, Laurentia, and Baltica). Intercontinental correlation of the Huaqiao Formation is based on the use of cosmopolitan agnostoid trilobites (see Peng and Robison, 2000). Polymerid trilobites provide additional biostratigraphic information that is summarized in this section. NORTH CHINA AND NORTHEAST CHINA PLATFORMS

The Dorypyge richthofeni Zone contains Amphoton deois, D. richthofeni, Grandioculus, Mapania, Menocephalites, Moutunia, Sudanomocarina, Szeaspis, all of which are known from the upper Crepicephalina and Amphoton zones of the North China and Northeast China platforms. Lisania is characteristic of the lower to middle part of the Pianaspis sinensis Zone of South China and is characteristic of the Taitzuia-Poshania Zone of North China and Northeast China. Parablackwelderia and some other damesellids appear first in the upper part of the Pianaspis sinensis Zone, whereas Parablackwelderia [=Damesops] is restricted to the Damesella-Yabeia Zone of North China and Northeast China. Prodamesella has a similar stratigraphic distribution. The Wanshania wanshanensis Zone contains Monkaspis and Teinistion, both of which are known from the Blackwelderia Zone of North China and Northeast China. The Wanshania wanshanensis Zone of South China is probably equivalent to the Blackwelderia Zone of North and Northeast China. Moreover, the base of the overlying zone as recognized in South China, the Liostracina bella Zone, is evidently correlative with the Drepanura Zone of North and Northeast China because the guide fossils Chaiwangella, Taihangshania, and Liostracina first appear in the basal parts of both the L. bella and the Drepanura zones. Guo and Zhang (1992) recognized the ProchuangiaParacoosia Zone as the basal zone of the Changshanian Stage. This zone is probably correlative with the upper part of the L. bella Zone, although in Hunan Paracoosia occurs in the lower part of the zone, whereas Prochuangia occurs in the overlying Chuangia subquadrangulata Zone. The Chuangia Zone of North and Northeast China is characterized by abundant specimens of Chuangia spp. (including Chuangia subquadrangulata) and Prochuangia; the zone is probably correlative with the Chuangia subquadrangulata Zone and the lowermost Shengia quadrata Zone of northwestern Hunan. TARIM AND NORTHWEST CHINA

The Buttsia-"Cycloagnostus" Zone of Tarim and Northwest China seems to be correlative with the Wanshania wanshanesis Zone of South China. Guide fossils shared between the two regions are Buttsia globosa, Wanshania, Torifera [ =Cyclolorenzella ], Parablackwelderia [ = Paradamesops ] • 40.

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jimaensis and Protaizehoia (see Lin et al., 1990). The Liostracina Zone of Tarim correlates with the Liostracina bella Zone of South China. Guide fossils shared between the two regions are Ajrikina [=Jiangnania], Liostracina, Teinistion [-Do©'pygella] , and Meteoraspis. WESTERN ZHEJIANG Agnostoid biostratigraphy (Peng and Robison, 2000; Lu and Lin, 1989) for northwestern Hunan and western Zhejiang suggests a precise correlation of the studied interval of the Huaqiao Formation with the Huayanshi and the basal Siyanshan formations. Although Ptychagnostus punctuosus and Goniagnostus nathorsti have not been recorded yet from western Zhejiang, Pseudophalacroma ovale, which ranges from the P. punctuosus through the G. nathorsti zones, suggests a correlation of these two zones of western Hunan to the Pseudophalacroma ovale Zone of western Zhejiang. Lejopyge armata appears prior to L. laevigata in sections in both northwestern Hunan and western Zhejiang, suggesting that the base of the Lejopyge armata Zone of western Zhejiang falls within the upper part of the Goniagnostus nathorsti Zone of northwestern Hunan. Proagnostus bulbus and Lejopyge sinensis are known from both sites, and indicate the base of the Lejopyge sinensis Zone of western Zhejiang is slightly younger than that of Proagnostus bulbus Zone of northwestern Hunan. Polymerids known also from western Zhejiang include Fuzhouia,

Pianaspis sinensis, Baojingia, Buttsia globosa, Paradamesella, Palaadotes, Torifera, Teinistion, Liostracina bella, Proceratopyge fuyanensis, and P. fenghwangensis. They all share similar occurrences there as in Paibi and Wangcun, northwestern Hunan. AUSTRALIA The Erediaspis Zone and Acmarharchis quasivespa Zone of Queensland, Australia (Opik, 1967) are correlative with the Wangshania wangshanensis Zone and the lower and middle part of Liostracina bella Zone of South China. Guide fossils shared between the two regions include damesellids (i.e., Parablackwelderia [=Merigaspis and Blackwelderia (in part)] and Palaeadotes), and Rhyssometopus. Taihangshania [=Teinistion? amydium] and Liostracina suggest a direct correlation between the upper part of the Liostracina bella Zone of South China and the Glyptagnostus stolidotus Zone of Queensland. BALTICA Correlation of the Huaqiao Formation with the equivalent interval in Baltica is principally on the basis of agnostoid biostratigraphy. The base of the Po,chagnostus punctuosus Zone and the base of the Lejopyge laevigata Zone in both Sweden and northwestern Hunan are directly correlative. Liguagnostus reconditus ranges through the upper two-thirds of the Agnostus pisiformis Zone in Sweden (Ahlberg and Ahlgren, 1996; Ahlberg, 2003). This suggests that the base of the L. reconditus Zone falls within the lower part of the A. pisiformis Zone. Glyptagnostus reticulatus first appears near the base of the Homagnostus obesus Zone (Olenus gibbosus Subzone) in Sweden (Ahlberg, 2003; Westerg~.rd, 1947), and this suggests that this level is correlative with the base of the Furongian Series (Paibian Stage). Olenus from the basal part of the Paibian Stage supports the correlation. Besides Olenus, several additional polymerids such as Dorypyge, Schmalenseeia, Palaeodotes, Paradamesella, and Proceratopyge conifrons are also known from Sweden (Moberg, 1903; Westerg~rd, 1922, 1947, 1948.) or elsewhere in Baltica, but their occurrences in Baltica are not always consistent with those in northwestern Hunan.

-42-

KAZAKHSTAN Agnostoids from Kazakhstan permit rather precise correlation between South China and Kazakhstan (Peng and Robison, 2000; Eragliev, 1980). In addition to the agnostoids, both regions also share many polymerids. Parablackwelderia [=Meringaspis and Blackwelderia], Wanshania [Peichiashania], Neoanomocarella, Palaeodotes, Paradamesella [=Palaeodotes (in part)], Prodamesella, and Ajrikina from the upper Pianaspis sinensis Zone through the lower Liostracina bella Zone of South China are all known from the Lejopyge laevigata and Kormagnostus simplex zones of Kazakhstan (Ergaliev, 1980); whereas Fenghuangella liostracinala [-Cyclolorenzella coniformis], Glaphyraspis, Tienistion [=Blackwelderia sp.], Baikadamaspis, Prochuangia, and Olenus [=Eugonocare cf. E. tesselatum] from the upper Liostracina bella Zone and the Chuangia subquadrangulata Zone of South China are known from the Glyptagnostus stolidotus Zone and the G. reticulatus Zone of Kazakhstan. LAURENTIA In Laurentia, the base of the Bolaspidella Zone coincides with the base of the Ptychagnostus atavus Zone (Rowell et al., 1982; Ludvigsen and Westrop, 1985). That horizon correlates to the base of the P. atavus Zone of northwestern Hunan, and the position is slightly lower than the base of the Dorypyge richthofeni Zone. The base of the Aphelaspis Zone of Laurentia correlates closely with the base of the Glyptagnostus reticulatus Zone (Palmer, 1962, 1998, 1999); that horizon is slightly lower than the base of the Chuangia subquadrangulata Zone of northwestern Hunan. Pratt (1992, fig. 20 ) correlated the Cedaria and Crepicephalina zones of the Great Basin with four successive Cedaria zones, the Cedaria minor, C. selwyni, C. prolifica, and C. brevifrons zones, of the Mackenzie Mountains, northwest Canada. The first appearance of Proagnostus bulbus at the base of the Cedaria minor Zone suggests that the base of the zone correlates with the base of the Proagnostus bulbus Zone of western Hunan. Correlation of the base of the Crepicephalius is tentative, but the range of Nahannagnostus nganasanicus from the base of the C. selwyni Zone through the C. brevifrons Zone in the Mackenzie Mountains suggests that the upper part of the Cedaria Zone of the Great Basin (=C. selwyni Zone) correlates with the upper part of the P. bulbus Zone through the Glyptagnostus stolidotus Zone of northwestern Hunan. If so, the base of the Crepicephalus Zone may correspond closely with the base of the Linguagnostus reconditus zone. SIBERIA Correlation of the Tomagnostus fissus-Paradoxides hicksi, Anopolenus henrici, and Anomocarioides limbataeformis zones of the Siberian Platform with the Dorypyge richthofeni Zone and the lower part of the Pianaspis sinensis Zone of the South China Platform is based on the presence of Pseudanomocarella horrida and Corynexochus perforatus in the T. fissus-P, hicksi Zone; Ptychagnostus atavus, P. punctuosus, Goniagnostus nathorsti, Linguagnostus kjerulfi, and Doryagnostus incertus in the A. henrici Zone; and Pianaspis attenda in the A. limbataeformis Zone (see Egorova et al., 1982, fig. 2). These correlations are supported by the occurrence of some related taxa in the two regions. For example, Pseudanomocarella horrida and Corynexochus perforatus of Siberia are similar to Sudanomocarina cf. S. changi and Corynexochus xiangxiensis from the basal part of Dorypyge richthofeni Zone of northwestern Hunan. A variety of taxa known from the upper D. richthofeni and the lower Pianaspis sinensis zones of northwestern Hunan are also present in Siberia.

• 43"

SYSTEMATIC PALEONTOLOGY Class TRILOBITA Walch, 1771 Order

CORYNEXOCHIDA

Suborder

Kobayashi, 1935

CORYNEXOCHINA

Kobayashi, 1935

Family CORYNEXOCHIDAEAngelin, 1854 Genus CORYNEXOCHUSAngelin, 1854

Corynexochus Angelin, 1854, p. 59; Zittel, 1885, p. 602; Gr6nwall, 1902, p. 136; Lindstr6m, 1901, p. 22; Walcott, 1916b, p. 309-312; Lake, 1934, p. 180-184; Resser, 1936, p. 22, 23; Whitehouse, 1939, p. 233,234; Lermontova, 1940, p. 144" WestergS.rd, 1948, p. 9-11" Ivshin, 1953, p. 71-75" Hup6, 1953a, p. 167; Moore, 1959, p. 0227" Chernysheva, 1960, p. 77; Egorova, Ivshin, Pokrovskaya, Repina, Rozova, Romenenko, and Sivov, 1960, p. 191; Suvorova, 1964, p. 198-202; Opik, 1967, p. 177, 178; Palmer, 1968, p. 42; Egorova and Savitsky, 1969, p. 166, 167; Bognibova, Kontev, Mikhailova, Poletaeva, Romanenko, Romanenko, Semashko, Tomashnoliskaya, Fedjanina, and Chernysheva, 1971, p. 127; Zhou, Liu, Meng, and Sun, 1977, p. 136; Zhu, Lin, and Zhang, 1979, p. 88; Shergold, 1982, p. 47; Zhou, Li, and Qu, 1982, p. 234; Egorova, Pegel, and Chernysheva, 1982, p. 79; Lisogor, Rozov, and Rozova, 1988, p. 68; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 63; Peng, 1992, p. 34; Pratt, 1992, p. 44, 45. Karlia Walcott, 1889, p. 444; 1916a, p. 223. Type species. Corynexochus spinulosus Angelin, 1854 (p. 59, pl. 33, figs. 9, 9a; non fig. 11), from the Paradoxides forchhammeri Zone, Andrarum Limestone, Andrarum, Sweden; by original designation. Other species. At least sixteen species, including the type species, and two questionably assigned species (Suvorova, 1964, p. 199; Babcock, 1994, p. 92, fig. 10) are included in this genus. The following species should be included in this genus: Corynexochus solitus Egorova in Bognibova, 1965 (p. 69, pl. 1, figs. 7, 8 ) from the Middle Cambrian of Siberia; Corynexochus terminalis Poletaeva in Poletaeva and Romanenko (1970a, p. 78, pl. 11, figs. 10-16; 1970b, p. 223, pl. 11, figs. 10-16), from the upper part of the middle Cambrian to the lower part of the upper Cambrian (now Furongian Series) of Altai; Corynexochus donggouesis Lin and Zhang in Zhu et al. (1979, p. 88, pl. 37, figs. 6-7, ?8) from the Nidangshan Group (Middle Cambrian) at Donggou, Huangzhong, Gansu; Corynexochus pulcher Zhou in Zhou et al. (1982, pl. 60, figs. 3, 4) from the upper part of the Shuangyingshan Group (upper Cambrian; now Furongian Series) at Shuangyuingshan, Subei, Gansu; and Corynexochus xiangxiensis sp. nov. Species left in open nomenclature, and which are questionablly assigned to Corynexochus, are •

44

•

Corynexochus aff. C. perforatus Lermontova (Repina et al., 1975, p. 130, pl. 15, fig. 8) from the Dignaceps Zone, Ryis-cai, Madyigen, Turkestan; Corynexochus aft. C. solitus Egorova in Bognibova (Egorova et al., 1982, p. 80, pl. 53, figs. 8, 11, 12) from the Tomagnostusfissus Zone of Siberia; Corynexochus sp. sensu Ivshin (1953, p. 76, 77, pl. 6, fig. 2) from the AnomocareGlaberagnostus [=Phoidagnostus] biturberculatus Zone of Boschekuli, Kazakhstan, Corynexochus sp. sensu Repina et al. (1975, p. 130, p|. 15, fig. 9) from the Sdzuyella-Aegunaspis Zone, Suliuktyi, Turkestan; and Corynexochus sp. (Zhou et al., 1996, pl. 7, figs. 3, 4) from the top part of the Heicigou Group (middle Cambrian) at Honggedacun, Tianzhu, Gansu. Corynexochus chinensis Lin and Zhang in Zhu et al., (1979, p. 88, pl. 37, fig. 5) from the upper Cambrian (now Furongian Series) at Chuancigou, Qilian, Qinghai, is probably a junior synonym of C. termilaris. Corynexochus cf. C. chinensis (Xiang and Zhang, 1985, p. 100, pl. 27, fig. 1) from the Glyptagnostus reticulatus Zone at Guozigou, Huocheng, Xinjiang, is regarded here as a junior synonym of C. plumula Whitehouse. C. hunanensis Duan, Yang, and Shi, 1999 (p. 164, 165, figs. 7G, 7H, 15), from the Huaqiao Formation at Haichongkou, Fenghuang (Furongian), northwestern Hunan, is based on juvenile specimens. The juvenile specimens are poorly preserved and associated with adults of C. plumula. Thus, it is likely that C. hunanensis is a junior synonym of C. plumula. Corynexochus eosimple Yang in Yang et al., 1991 (p. 124, pl. 9, figs. 3-5; 1993, p. 160, 161, pl. 9, figs. 3-5) and Corynexochus elongatus Yang in Yang et al. (1991, p. 125, pl. 9, figs. 6-8; 1993, p. 161, pl. 9, figs. 6-8), both from the "Mufushania"-Holocephalina Zone of the Fengjiamiao Formation (lower middle Cambrian) at Xiouzigou, Xichuan, Henan, are based on poorly preserved material, and their specific validity remains uncertain. Except for C. plumula and C. pulcher, both of which are Furongian (late Cambrian) in age, and C. terminalis, which ranges from the middle Cambrian to the upper Cambrian, all species assigned to Corynexochus are middle Cambrian in age (pre-Youshuiian in South China stage terms). Remarks. The concept of this genus has been discussed by various authors (see Shergold, 1982, p. 47). The diagnosis given by Suvorova (1964, p. 198, 199) is followed here. Corynexochus xiangxiensis sp. nov.

Plate 1, figures 1-10, ? 11, ? 12 2001b Corynexochussp.; Peng, Babcock, and Lin, p. 101, pl. l, figs. 1, 2. Etymology. From Chinese, Xiangxi, brief name for northwestern Hunan. Holotype. Cranidium (P1. 1, figs. 5-7; NIGP 137285) from collection P3.2. Other material. More than 10 sclerites including cranidia, hypostomes, and pygidia (NIGP 137282-137284, 137286-137288) in collection P3.2. Diagnosis. Corynexochus with elongate, anteriorly expanded glabella. Four faint lateral glabellar furrows; S 1, $2 nearly transverse; $3, $4 inwardly and forwardly directed. Preglabellar area absent. Palpebral lobe long, reaching to position of S1. Posterior area of fixigena short, with nearly transverse, abaxially broadening posterior margin. Pygidium nearly effaced on axis and pleural •

45

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areas, anterior border deep, lateral and posterior border furrows faint.

Description. Cranidium trapezoidal in outline, moderately convex; width twice length; anterior margin moderately arched forward. Glabella long, length excluding occipital ring about twice width at base; 4 pairs of faint lateral glabellar furrows; S 1, $2 long, nearly transverse; $3 shorter than S 1 and $2, inwardly and forwardly directed; $4 shortest, inwardly and forwardly directed, extending beyond position of anterolateral pit. Transverse occipital furrow deeply incised. Occipital ring tumid, posterior margin strongly curved rearward; occipital node faint, located posteriorly. Palpebral lobe long, gently curved, well defined by palpebral furrow, located between S 1 and $4, length (exs.) about half that of glabella. Fixigena moderately convex; preocular area short, narrow; palpebral area gently convex, width equal to two-thirds of glabellar width at occipital furrow; posterior area short, posterior border furrow deeply incised but shallowing abaxially; posterior border convex, widening abaxially. Posterior margin nearly transverse. Anterior branch of facial suture short, convergent forward; posterior branch of facial suture long, strongly divergent rearward. Hypostome fused to rostral plate, slightly longer than wide with small, inflated middle body, wide and shallow border furrows, convex posterior border, and forwardly narrowing lateral border. Rostral plate long, convex, moderately arched forward, defined distally by straight, diagonally directed connective sutures. Pygidium subtriangular in outline, width two-thirds length, moderately convex; axis and pleurae mostly effaced; lateral and posterior border furrows shallow. Axis obscurely divided into 5 tings and a tiny, subtriangular terminal piece, reaching almost to posterior border furrow. Pleural field with 3 obscure pleural furrows; only the first interpleural furrow, which is fine and threadlike, present. Remarks. Corynexochus xiangxiensis, which is from the base of the Ptychagnostus atavus Zone of the Huaqiao Formation near Paibi, Hunan, is one of the earliest known species assigned to this genus. It is characterized by having a long glabella with four pairs of lateral furrows and a poorly furrowed pygidium, and by lacking a preglabellar area. Among the species assigned to Corynexochus, the new species is most closely similar to C. pussilus (Illing, 1916, p. 431, pl. 37, figs. 1, 2) from the Stockingford Shale of Nuneaton, England. It differs from C. pussilus, however, in having a diagonal posterior branch of the facial suture, a wider posterior border of the cranidium and, apparently, weaker lateral glabellar furrows. C. perforatus Lermontova (1940, p. 144, pl. 44, figs. 4, 4a) and C.filix Suvorova (1964, p. 203, pl. 24, figs. 9, 10; pl. 25, figs. 1-15, text-figs. 58, 59) are also similar to C. xiangxiensis, but they are distinguished by having a more expanded glabella. The former is further distinguished by having more distinctly furrowed pleural areas in the pygidia. C. filix also has wider palpebral lobes than does C. xiangxiensis. C. terminalis (Poletaeva and Romanenko, 1970, pl. 11, figs. 10-16) is similar to C. xiangxiensis in terms of cranidial outline and effacement, but it differs in having a proportionally shorter glabella, and smaller palpebral lobes. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Genus CORYNEXOCHINALermontova, 1940

Corynexochina Lermontova, 1940, p. 144; 1951, p. 23" Ivshin, 1953, p. 77, 78; Hupr, 1953a, p. 167" • 46

•

Moore, 1959, p. 0227; Suvorova and Chernysheva, 1960, p. 78; Repina, Petrunina, and Khairullina, 1975, p. 131.

Type species. Corynexochina weberi Lermontova, 1940 (p. 144, pl. 44, figs. 5, 5a), from the Middle Cambrian, southern Fergana, Uzbekistan; by original designation. Other species. Corynexochina asiatica Ivshin, 1953 (p. 78, pl. 6, fig. 5) from the AnomocareGlaberagnostus [=Phoidagnostus] bituberculatus Zone, Boschekuli, Kazakhstan; Corynexochina sinensis sp. nov. Diagnosis. Genus of Corynexochidae that is largely effaced. Glabella strongly expanded forward; occipital ring uniformly long (sag., exs.) or narrowing abaxially. Palpebral lobe medium to small sized, located near or slightly anterior to glabellar midlength. Pygidium strongly effaced, with axis merging with pleurae.

Remarks. As discussed by Lermontova (1951, p. 23) and Ivshin (1953, p. 78), Corynexochina is closely similar to Corynexochus in the structure of the cranidium and pygidium. However, Corynexochina differs from Corynexochus by being quite effaced on the dorsal surface. Blandiaspis Qian in Qiu et al., 1983 (p. 164, 165) (type species, B. anhuiensis), is closely comparable to Corynexochina and probably is a junior synonym of it Blandiaspis is similar to Corynexochus in overall morphology and effacement. It is also similar in size. Blandiaspis is monospecific. The type species includes only a single cranidium (the holotype) and a single pygidium. Both specimens are poorly preserved. The limited, poorly preserved material of B. anhuiensis is insufficient to determine its validity.

Corynexochina sinensis sp. nov. Plate 1, figures 13-16; Plate 2, figures 1-7 2001b Corynexochinasp. Peng, Babcock, and Lin, p. 101, pl. 1, figs. 5, 6.

Etymology. From Latin, sinae, China. Holotype. Cranidium (P1. 1, fig. 13, NIGP 137289) from collection W3. Other material. One cranidium, one hypostome, and one pygidium (NIGP 137290-137294) in collections W0 and W3. Diagnosis. Corynexochina with short glabella; glabellar length slightly less than maximum glabellar width; palpebral lobe small; occipital ring of uniform length. Pygidium with incomplete axial furrows; posterior part of axis merging with pleurae.

Description. Cranidium trapezoidal in outline, moderately convex. Glabella mostly effaced, gently convex, short, wide; width at base 0.56- 0.58 of the length; glabella expanding forward or slightly constricted near midlength, evenly rounded anteriorly. Furrows on axis effaced except for occipital furrow, which is obscurely defined and gently curved rearward. Axial furrow faint, beating • 47

•

anterolateral pit anteriorly. Occipital ring of uniform length. Palpebral lobe short, lying about midlength of cranidium. Palpebral furrow faint on external surface, wide and shallow on steinkern. Palpebral field of fixigena narrow. Facial suture possibly proparian. Posterior branch of facial suture diagonal, enclosing a triangular posterolateral projection. Anterior branch of facial suture short, convergent forward; posterior branch of facial suture long, strongly divergent rearward. Posterior border furrow wide, deep, with abaxial end curved strongly forward. Posterior border widened abaxially, possibly with short, broad genal spine. Posterior margin nearly transverse. Hypostome somewhat longer than wide; middle body ovate in outline, fused to rostral plate, inflated, separated from the rostral plate by wide, shallow anterior border furrow; lateral border narrow; widening posteriorly into crescentic posterior border; posterior border convex. Rostral plate convex, moderately arched forward, with straight, nonfunctional connective suture distally. Pygidium subtriangular in outline, moderately convex, width two-thirds length; axis and pleurae mostly effaced, lateral and posterior border furrows shallow. Axis divided into 5 obscure tings and tiny, subtriangular terminal piece, reaching almost to posterior border furrow. Pleural field with 3 obscure pleural furrows; only the first interpleural furrow, which is fine, threadlike, present. Remarks. Lermontova (1951, pl. 2, figs. 5-9) refigured C. webei, the type species of Corynexochina, together with additional material from the type locality. New material reported here as the new species C. sinensis represents the first record of Corynexochina in China. The new species is similar in general to the type species, but differs in some important ways, notably in having a proportionally shorter and wider glabella that is less expanded anteriorly, and in having an occipital ring that is of even width, rather than narrowing abaxially. It also differs in having a smaller palpebral lobe, and a wider but shorter pygidium. The pygidial axis of the new species is shorter and more convex anteriorly than is the axis of C. webei. C. asiatica from Kazakhstan is closely comparable to the new species in having a less expanded glabella. However, the glabella of that species is proportionately much longer than wide. C. asiatica can also be distinguished by having more clearly defined axial and occipital furrows. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the unnamed Zone and the Dorypyge richthofeni Zone (equivalent to the upmost of Ptychagnostus gibbus Zone and the basal of Ptychagnostus atavus Zone).

Genus CHATIANIAYang in Zhou et al., 1977 Chatiania Yang in Zhou et al., 1977, p. 36; Yin and Li, 1978, p. 448; Yang, 1978, p. 35, 36; Luo, 1982, p. 2; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 63. Parachatiania Yuan and Yin, 1998, p. 150. non Chatiania (Eochatiania) Yuan and Yin, 1998, p. 151. Type species. Chatiania chatianensis Yang in Zhou et al. (1977, p. 136, pl. 43, figs. 14, 15) from the Liostracina-Chatiania Zone of the Huaqiao Formation, Chatian, Fenghuang, northwestern Hunan, China; by original designation. Emended diagnosis. Glabella large, rectangular, moderately to strongly convex, expanding forward gently, with median notch anteriorly. Lateral glabellar furrows effaced or obscure. Preglabellar field • 48

•

absent. Anterior border narrow, length uniform or narrowing abaxially; anterior border furrow, palpebral furrow, and occipital furrow variably defined. Occipital node positioned anteriorly or centrally. Palpebral lobe moderately large, situated opposite glabellar midlength. Fixigena with small preocular field; palpebral area narrow to moderately wide; with subtriangular to bladelike posterolateral projection. Librigena with wide genal field and short, broad-based genal spine. Pygidium with lateral and posterior border furrows shallow to obscure; pleural furrows variably expressed, interpleural furrows faint. Remarks. Chatiania is a small-sized corynexochid trilobite characterized by the presence of a clearly defined anterior border. So far as known, this genus is limited to the Jiangnan Slope Belt of South China (eastern Guizhou and northwestern Hunan, southern Anhui, and western Zhejiang). Parachatiania is here considered to be a junior synonym of Chatiania. The genus was erected by Yuan and Yin with Parachatiania expansa Yuan and Yin (1998, p. 150, 151, pl. 3, figs. 1, 3, 4; non fig. 3) as the type species. The genetic diagnosis and all the stated differences between the type species and Chatiania chatianensis, the type species of Chatiania, seem to be of no more than species-level significance. Characters used to differentiate P. expansa (and, by implication, Parachatiania) include a deeper anterior border furrow, a more strongly vaulted glabella, a narrower palpebral area of the fixigena, and a longer palpebral lobe. Yuan and Yin (1998) erected a subgenus Chatiania (Eochatiania), with Chatiania (Eochatiania) laevigata as the type species. The subgenus is monospecific, and the type species is known from only the holotype. The holotype of C. (E.) laevigata is a cranidium having a length of less than 2 mm (Yuan and Yin, 1998, p. 151, pl. 3, figs. 5a, b). It is difficult to determine any difference between the holotype of C. (E.) laevigata and cranidia of similar-sized juveniles of Luyanhaoaspis decorosa (Peng et al., 1995) [formerly Luaspis decorosa Peng et al., 1995, pl. 6, fig. 12] except for the more anteriorly located palpebral lobes in C. (E.) laevigata. Because the holotype of C. (E.) laevigata is from the same formation and the same paleogeographic region as L. decorosa, and because it occurs within the stratigraphic range of L. decorosa, it is probably conspecific with L. decorosa. Here we consider C. (E.) laevigata to be a probable junior synonym of L. decorosa, and we consider Chatiania (Eochatiania) to be a possible invalid subgenus. Chatiania chatianensis Yang in Zhou et al., 1977

Plate 2, figures 8, 9, 13, 14; Plate 3, figures 1-11; Text-figure 8 1977 1978 1982 1983

Chatiania chatianensis Yang in Zhou et al., p. 136, pl. 43, figs. 14-15. Chatiania chatianensis Yang; Yang, p. 36, pl. 5, figs. 5-7. Chatiania chatianensis Yang; Liu, p. 304, pl. 215, figs. 9, 12 (refigured). Chatiania chatianensis Yang; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, p. 64, pl. 21, figs. 8, 9. 1987 Chatiania chatianensis Yang; Peng, p. 92, pl. 5, fig. 18. 1990 Chatiania chatianensis Yang; Dong, p. 79, pl. 3, fig. 13. 1991 Chatiania chatianensis Yang; Dong, p. 457, pl. 3, fig. 10 (refigured). 2001c Chatiania chatianensis Yang; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 2, fig. 21. 2001d Chatiania chatianensis Yang; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.7. Lectotype. Cranidium (Zhou, Liu, Meng, and Sun, 1977, pl. 43, fig. 14; same as Yang, 1978, pl. 5, fig. 5; CUGB 0108106) from the Liostracina bella Zone (as defined here) of the Huaqiao Formation, Chatian, Fenghuang, northwestern Hunan; designated herein.

.49.

N e w material. More than 30 cranidia and pygidia (illustrated specimens NIGP 137295-137297;

137300-137308) in collections P311.7, P317.4, P319.6, P325.7, P331.8, P337.5, P341.7, P346.7, P353.7, P356.5, P361.5, P362.4, P387.36, PI3-2.75, and W254.1. E m e n d e d diagnosis. Chatiania with short, flat anterior border, uniform in width. Glabella gently

convex, expanding forward gently. Anterior border furrows, axial furrow, palpebral furrow, and occipital furrow well incised. Palpebral area greater than one-third of glabellar width at glabellar midlength. Occipital node central. Pygidium with well-incised pleural furrows and faint border furrows. Axis poorly segmented behind first one or two tings. Lateral and posterior borders of pygidium sloping away from pleural field.

B

C Text-figure 8. Chatiania chatianensis Yang in Zhou et al., 1977. A, B, reconstruction of cephalon and pygidium based on specimens NIGP 137303, 137304 and 137297 (see P1.3, figs- 5 and 6 and P1.2, figs. 13, 14); C, lectotype cranidium (original of Zhou et al., 1977, pl. 43, fig. 14 and Yang, 1978, pl. 5, fig. 5; CUGB 0108106; selected herein).

• 50"

Description. Cranidium subtriangular in outline, slightly wider than long. Glabellar large, expanding forward gently to anterior border furrow, with median notch anteriorly; lateral glabellar furrows effaced. Axial furrow deeply incised, shallowing near eye ridge. Occipital furrow deep, transverse, ending sharply at sides; occipital ring with posterior margin gently curved posteriorly. Anterior border fiat, narrow, uniform in width. Palpebral lobe moderately long, with distinct palpebral furrow; palpebral lobe close to axial furrow anteriorly, distant from axial furrow posteriorly. Palpebral area of fixigena is more than half glabellar width in small cranidium (P1. 2, fig. 3), and about one-third glabellar width in large cranidium (P1. 2, fig. 6). Anterior branch of facial suture convergent forward, enclosing a small preocular field; posterior branch strongly divergent outward and rearward to the level of posterior border furrow, then turning sharply posteriorly and slightly outward, enclosing a blade-like posterolateral projection; posterior border defined by incised posterior border furrow, widening adaxially. Librigena with wide, gently convex genal field, border furrow shallow, broad; genal spine short, broadly based. Pygidium semielliptical to semicircular in outline. Axis with first two tings weakly defined, others fused. Pleural area with incised anterior border furrow and pleural furrows, interpleural furrows obscure. Lateral and posterior border furrows effaced on dorsal surface; lateral and posterior borders continuing slope of pleural field. On exfoliated surface, pleural furrows are shallow, wide; interpleural furrows and posterior and lateral border furrows faint, defining moderately wide borders. Remarks. A cranidium from the syntypic material of Yang (1978) is here designated as the lectotype for Chatiania chatianensis, the type species of Chatiania. The type material of Yang (1978) includes two cranidia and one pygidium. One of these cranidia and the pygidium (CUGB 0108106 and 0108105) were illustrated by Zhou et al. (1977). When Yang (1978) subsequently published his material, the two specimens illustrated by Zhou et al. (1977) were both designated as holotypes in the the text and the figure caption (Yang, 1978, p. 36, 78). From these two specimens, the cranidium is selected here as the lectotype. The rich new material of Chatiania chatianensis from the Huaqiao Formation resembles the type material described by Yang (in Zhou et al., 1977) from the same area in northwestern Hunan, and allows for an emended diagnosis of the species. This species is characterized principally by a relatively large, forwardly expanding glabella of low convexity; a narrow, flat, anterior border; a moderately large palpebral lobe; and poorly defined lateral and posterior borders on the pygidium. The new material shows ontogenetic variation in the shape of glabella and the width of the palpebral area. The glabella is well expanded anteriorly in small cranidia (P1. 2, figs. 3, 4) but becomes less expanded and nearly parallel-sided in large cranidia (P1.2, fig. 6). Occurrence. The species is known from northwestern Hunan at Chatian in Fenghuang, Wa'ergang in Taoyuan, Paibi in Huayuan, and Wangcun in Yongshun. Its observed stratigraphic range is from the Linguagnostus reconditus Zone to the G. reticulatus Zone. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). In South China, it is known also from the equivalent interval (Huayansi Formation or Tuanshan Formation) in Guichi and Jingxian, southern Anhui, and in Fuyang, westem Zhejiang.

.51.

Chatiania expansa (Yuan and Yin, 1998)

Plate 4, figures 1-14 1998

Parachatiania expansa Yuan and Yin (in part), p. 150, 151, pl. 3, figs. 1, 3, 4; non fig. 3 [?=Townleyella rara (Yuan and Yin)]. 2001b Chatiania expansa (Yuan and Yin); Peng, Babcock, and Lin, p. 104, pl. 11, figs. 3, 4. Holotype. Cranidium (Yuan and Yin, 1998, pl. 3, fig. 1; NIGP 127917) from the middle part of the

Huaqiao Formation (previously referred to the Chefu Formation). New material. More than 20 cranidia and pygidia (illustrated specimens NIGP 137313-137322) in

collections P308, W213, W225, and W227. Diagnosis. Chatiania with thick palpebral lobe; posterior end of posterior lobe close to axial furrow,

defined by shallow palpebral furrow; palpebral area narrow (tr.), less than one-fourth glabellar width; anterior border moderately wide, fiat. Pygidium with short, wide axis reaching to posterior border furrow; lateral border and posterior border wide, defined by shallow border furrows and change of slope in pleural field. Description. Glabella large, gently convex, subrectangular in outline, parallel-sided or gently

tapering forward, broadly or obtusely rounded anteriorly with faint indications of F1 and F2, and a weak median notch in the front. Axial furrow shallow, weakly defined; occipital furrow rather deep, transverse, becoming shallow at sides; occipital ring prominent, with posterior margin gently curved posteriorly, bearing a weak node lying anteriorly and close to occipital furrow. Anterior border fiat, rather wide, uniform in width. Palpebral lobe thick, moderately long, defined weakly by shallow palpebral furrow, with both the anterior and the posterior ends close to axial furrow. Palpebral area of fixigena fiat, about the same width as glabella in small cranidium (P1. 4, fig. 1) and one-fifth the width of glabella in large cranidium (P1. 4, fig. 14). Anterior branch of facial suture short, converging forward and enclosing a small preocular field; posterior branch diverging strongly outward and rearward to enclose a blade-like posterolateral projection; posterior border furrow shallow, posterior border gently widening adaxially. Pygidium semielliptical in outline. Axis with first two tings weakly or faintly defined, others fused. Pleural area gently convex, with deep anterior border furrow and pleural furrows, and faint interpleural furrows; anterior band and posterior band are about equally wide (exs.). Lateral and posterior border furrows shallow; lateral and posterior borders flat, moderately wide. Remarks. Chatiania expansa is closely similar to C. chatianensis, the type species of Chatiania, in general aspects of both the cranidium and pygidium. C. expansa differs, however, in having

shallow or faint furrows on the cranidium, rather than thin and incised furrows; in having a wider palpebral lobe whose posterior end is located close to the axial furrow; in having well-defined, but not deeply incised, pleural furrows on the pygidium; and in having well-defined borders. An ontogenetic series for this species shows a pattern similar to that known from the type species, C. chatianensis (cf. P1. 3, figs. 1-6). In early ontogenetic stages, the glabella of C. expansa was expanded forward. By the late holaspid period, however, the glabella became nearly parallel-sided. Also through ontogeny, the palpebral area of the fixigena changed from being large • 52.

and wide in earlier stages to small and narrow in later stages. Two pygidia assigned to C. expansa by Yuan and Yin (1998, pl. 3, figs. 2, 3) evidently belong to different species. One pygidium (Yuan and Yin, 1998, pl. 3, fig. 2) seems to be conspecific with pygidia that we assign here to the species. The other pygidium (Yuan and Yin, 1998, pl. 3, fig. 3), however, does not seem to agree with pygidia assigned to either C. expansa or C. chatianensis. The pygidium bears clearly defined interpleural furrows, but they are faint in C. expansa; the posterior band of the pleura is not gently curved as in C. expansa, but runs nearly transversely from the axial furrow and then curves strongly outwards and rearwards as it reaches the narrow and convex posterior border. In both C. expansa and C. chatianensis, the pleural bands are not connected with the lateral and posterior borders, and the border furrows are continuous, rather than interrupted by the bands. The pygidium illustrated by Yang and Yin (1998, pl. 3, fig. 3) appears to belong to Townleyella rara (Yuan and Yin, 1998, pl. 2, fig. 13). As noted by Yuan and Yin (1998, pl. 2, fig. 3), another pygidium from the same field collection (WY23F1; now in the NIGP collection) may belong to Townleyella rara. Occurrence. The species occurs in the northwestern Hunan-eastern Guizhou area, including Paibi, Huayuan; Wangcun, Yongshun; Yangweizhou, Wanshan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Chatiania sp. cf. C. chatianensis Yang in Zhou et al., 1977

Plate 3, figures 12-16 2001b Chatiania cf. C. chatianensis Yang; Peng, Babcock, and Lin, p. 105, pl. 13, figs. 3, 4. Material. Five cranidia and one pygidium(illustrated specimens NIGP

137309-137312) in

collections P319.8, P337.5, and P341.8. Remarks. Material compared to C. chatianensis is similar to specimens referred without question to

that species in the shape of glabella, the shape of the palpebral area, and the appearance of the cranidial and pygidial furrows. It differs from unquestionable C. chatianensis in having a convex, abaxially narrowing anterior border in the cranidium; a short, wide pygidial axis; and broad lateral and posterior borders in the pygidium. When exfoliated, the pygidium of C. chatianensis bears borders comparable to that of the pygidium referred to C. sp. cf. C. chatianensis. However, its pleural and interpleural furrows are faint or shallow rather than incised (P1.2, fig. 8). Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Chatiania convexa sp. nov.

Plate 5, figures 1-9 2001b

Chantiania sp.; Peng, Babcock, and Lin, p. 103, pl. 8, fig. 3.

.53"

Etymology. From Latin, convexus, convex, in reference to the convex glabella. Holotype. Cranidium (P1.5, figs. 5-7, NIGP 137325), from collection P273.66. Other material. Three cranidia (NIGP 137323, 137324, 137326) from collections P269, P278.1, and P282.6 are figured as paratypes; six additional cranidia from collections P273.8, P296.4, P298.54, and P341.7. Diagnosis. Chatiania with strongly convex glabella, subquadrate or barrel-like in outline; palpebral lobe moderately long, lying close to axial furrow; palpebral furrow well defined; palpebral area of fixigena narrow. Description. Cranidium subquadrate in outline with gentle forward curve at anterior margin. Glabella large, tumid, strongly convex sagittally, slightly expanded anteriorly or at midlength; except for a faint S 1, glabellar furrows are effaced completely. Axial and occipital furrows shallow but clearly defined; occipital ring with posterior margin gently curved rearward, beating a tiny and weak node anteriorly. Anterior border narrow, fiat or weakly convex, uniform in width. Palpebral lobe medium-sized, placed opposite midlength of glabella and close to axial furrow; palpebral furrow shallow; palpebral area of fixigena narrow, convex. Anterior branch of facial suture short, extending forward and slightly outward, enclosing a short and narrow preocular field; posterior branch directed diagonally; posterolateral projection subtriangular, bearing wide and shallow border furrow and narrow (exs.) posterior border. Remarks. The new species is referred to Chatiania because it resembles closely both C. chatianensis and C. expansa. The shape, proportion and effacement of glabella, the nature of the anterior border, the position, shape and size of the palpebral lobe are all greatly reminiscent of those species. However, it can be easily differentiated from both previously known species by the greater convexity of the glabella, a palpebral lobe that is much closer to the axial furrow, and a narrower palpebral area of the fixigena. The large, effaced glabella, which has high convexity and narrow palpebral areas, are reminiscent of those characters in Luyanhaoaspis decorosa (Peng et al., 1995, pl. 6, figs. 1-9). L. decorosa differs in having a wider (sag.) anterior border, a smaller palpebral lobe that is defined by an obscure palpebral furrow, and a narrower (tr.) but longer (exs.) posterolateral projection. In L. decorosa, the axial furrow is much shallower than that in the new species, and the occipital furrow is completely effaced. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the Linguagnostus reconditus Zone). Family DOLICHOMETOPIDAEWalcott, 1916 Genus AMPHOTON Lorenz, 1906

Amphoton Lorenz, 1906, p. 89-91; Kobayashi, 1935, p. 137, 138; 1942a, p. 162; 1942d, p. 471,472; Hup6, 1953a, p. 165; Lu, 1957, p. 263" Poulsen in Moore, 1959, p. 0222; Suvorova and Chernysheva, 1960, p. 74; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 112; Zhou, Liu, Meng, and • 54-

Sun, 1977, p. 134; Yin and Li, 1978, p. 447; Opik, 1982, p. 57; Liu, 1982, p. 302; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 60; Zhang and Wang, 1985, p. 341; Zhang and Jell, 1987, p. 62, 63; Guo, Zan, and Luo, 1996, p. 95, 96; Jago and McNeil, 1997, p. 87; Jago and Brown, 2001, p. 11. Amphoton (Amphotonella) Kobayashi, 1942a, p. 164, 165; 1942d, p. 472; Poulsen in Moore, 1959, p. 0222. Amphoton (Sunia) Kobayashi, 1942a, p. 165; 1942d, p. 472; Poulsen in Moore, 1959, p. 0222; Opik, 1982, p. 66, 67. non Amphoton (Fuchouia) Resser and Endo; Kobayashi, 1942a, p. 166, 167; Poulsen in Moore, 1959, p. 0222. Sunia Kobayashi; Lu, 1957, p. 263, 264; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 115; Opik, 1982, p. 66, 67; Xiang and Zhang, 1985, p. 102. Amphotonella Kobayashi; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 114; Nan, 1980, p. 486; Opik, 1982, p. 57, 58; Guo, Zan, and Luo, 1996, p. 97. Amphoton (Paramphoton) Yang in Zhou et al., 1977, p. 135; Yang, 1978, p. 34; Yin and Li, 1978, p. 447; Liu, 1982, p. 302. non Amphoton (Pseudamphoton) Chu and Zhang in Zhu et al., 1979, p. 86. Eurodeois Opik, 1982, p. 58.

Type species. Amphoton steinmanni Lorenz, 1906 (p. 89, pl. 4, figs. 15-17) from Laiwu, Shandong, China [subjective junior synonym of Dolichometopus deois Walcott, 1905]; by original designation. Remarks. Kobayashi (1942a) divided Amphoton into four subgenera: A (Amphoton), A. (Amphotonella), A. (Sunia), and A. (Fuchouia). Such a classification was accepted by Poulsen (in Moore, 1959), but Lu et al. (1965), followed by Opik (1982), elevated the subgenera to separate genera. Opik (1982) erected Eurodeois with Dolichometopus deois (Walcott, 1905) as the type species. However, D. deois is commonly considered to be synonymous with A. steinmanni, the type species of Amphoton. Zhang and Jell (1987, p. 62, 63) discussed Amphoton at length and suppressed Amphoton (Sunia) Kobayashi, and A. (Amphotonella) Kobayashi as junior synonyms of Amphoton. They also synonymized Eurodeois. We agree with Zhang and Jell (1987) in regarding Amphoton and Fuchouia to be separate, valid genera. These two genera are principally differentiated from each other in the size of the palpebral lobes and thus the size of the cranidial posterolateral projections, the number of thoracic segments, and the nature of the borders and interpleural furrows in the pygidium. As suggested by Opik (1961) and Zhang and Jell (1987), Dolichometopus may eventually prove to be a senior synonym of Amphoton, but because the characters of Dolichometopus are insufficiently known, we continue to retain Dolichometopus and Amphoton as separate genera. New material of Amphoton from the Huaqiao Formation, northwestern Hunan, shows that the subgenus Amphoton (Paramphoton) Yang (in Zhou et al., 1977) should also be considered a junior subjective synonym of Amphoton. Yang (1978, p. 34) differentiated the subgenus primarily on the presence of an occipital spine. However, as noted by Zhang and Jell (1987, p. 62), Amphoton also bears an occipital spine. Other major distinguishing features (Yang, 1978) include the presence of a pair of fossulae in front of the cranidial axial furrows and the absence of an anterior curvature in the posterior margin of the pygidium. Fossulae are also known in Amphoton (see Opik, 1982 for descriptions of species of Sunia). The sinuous posterior margin that Yang regarded as a diagnostic character of Amphoton may be based on incorrect illustrations of Kobayashi (1942a, pl. 1, fig. 10) and Poulsen (in Moore, 1959, fig. 163.5). In any case, if the character exists, it is probably an example of intraspecific variation. Guo et al. (1996) proposed to revive Amphotonella as a separate genus. This proposal is •

55.

rejected here because it was mainly based on the consideration that some of the occipital spine-bearing forms (i. e., their Amphotonella) have relatively higher stratigraphic occurrences than forms of Amphoton, which are presumed to lack an occipital spine. As discussed by Zhang and Jell (1987, p. 62), examination shows that the type of the type species of Amphoton bears spines throughout its meraspid and holaspid periods. The "non occipital spine-beating" Amphoton is due to preservation, and the genus is found with an occipital spine throughout its stratigraphic range. The stratigraphic occurrence has no value for a genetic assignment. Amphoton deois (Walcott, 1905)

Plate 6, figures 10 - 12 1905 1906 1906 1913 1916b 1924 1935 1937 1937 1937 1937 1937 1942a 1942 1942 1957 1965 1977 1982 1987 ?1996 1996 non 2001b

2001b

Dolichometopus deois Walcott, p. 94. Bathyuriscus asiaticus Lorenz, p. 87-89, pl. 5, figs. 1-5. Amphoton steinmanni Lorenz, p. 89-91, pl. 4, figs. 15-17. Dolichometopus deois Walcott, p. 216, 217, pl. 21, figs. 13, 13a- d; pl. 22, figs. 1, l a-h, 2, 2a, b. Dolichometopus? deois Walcott; Walcott, p. 365-367, pl. 54, figs. 1, 1a-m, 2. Dolichometopus deois Walcott; Sun, p. 81, 82, pl. 5, fig. 9. Amphoton deois Walcott; Kobayashi, p. 138, 139, pl. 22, fig. 12. Amphoton deois Walcott; Resser and Endo, p. 205,206, pl. 38, figs. 1, 9; pl. 58, fig. 23. Amphoton parallela Resser and Endo, p. 206, 207, pl. 38, figs. 2-8, 10-13; pl. 39, figs. 19, 20. Amphoton alia Resser and Endo, p. 207, pl. 38, figs. 14-18. Amphoton divergens Resser and Endo, p. 208, pl. 48, figs. 31, 32, non fig. 33. Dorypyge manchuriensis Resser and Endo (in part), p. 208-209, pl. 31, fig. 3 only. Amphoton deois Walcott; Kobayashi, p.175, 176, pl. 1, fig. 10; pl. 3, fig. 6. Amphoton blackwelderi Resser, p. 5. Amphoton kaipingense Resser, p. 5. Amphoton deois Walcott; Lu, p. 263, pl. 140, fig. 5. Amphoton deois Walcott; Lu, Zhang, Zhu, Qian, and Xiang, p. 113, pl. 17, fig. 16. Amphoton deois Walcott; Schrank, p. 148, pl. 2, figs. 6, 7" pl. 3, fig. 1. Eurodeoisdeois (Walcott); Opik, p. 58, pl. 20, fig. 5, text-fig. 28. Amphotondeois Walcott; Zhang and Jell (in part), p. 63-65, pl. 17, figs. 2-3, 5-14; pls. 18-20, 23" pl. 22, figs. 1-7; pl. 122, ?fig. 5; non pl. 17, figs. 4, 5 [=Amphoton alceste (Walcott)]. Amphoton deois Walcott; Guo, Zan, and Luo, 1996, p. 96, 97, pl. 38, figs. 1-8; pl. 39, figs. 1-15. Amphotonellajingxianensis Guo and Luo in Guo et al., 1996, p~ 98, pl. 38, figs. 9, 10; pl. 40, figs. 6-13. Amphoton deois Walcott; Peng, Babcock, and Lin, p. 102, pl. 3, figs. 4, 5 [=Amphoton sp. cf. A. typicum (Kobayashi, 1942a), herein]. Amphoton sp., Peng, Babcock, and Lin, p. 101, pl. 1, fig. 17.

Lectotype. Incomplete cranidium (Walcott, 1913, pl. 22, fig. 1, 1'; also Walcott, 1916b, pl. 54, fig. If; Resser and Endo, 1937, pl. 38, fig. 9; Zhang and Jell, 1987, pl. 2, 3, USNM 58249) from south of Yanzhuang, Shandong, China; by subsequent designation (Zhang and Jell, 1987). New material. Six pygidia (illustrated specimens NIGP 137341-137343) in collections P42.5, • 56"

P45.6, and P48.5. Remarks. Zhang and Jell (1978) discussed the specific concept at length and refigured numerous specimens previously illustrated by Walcott (1913), Resser and Endo (1937), and Lorenz (1906); together these specimens illustrate well the morphological variation within the species. Some of these specimens were previously referred to different species or even different genera. Except for a few specimens that are reassigned here, we concur with Zhang and Jell's (1987) new assignments for other specimens. Three pygidia from northwestern Hunan are assigned here to the species as they fall within the variation range of the species. They are almost identical with some of the pygidia figured by Zhang and Jell (1987, pl. 23, figs. 10, 13) and by Guo et al. (1996, pl. 39, figs. 4, 6, 12-15) from Liaoning. This is the first recognition of the species in South China. Previously this common species was known from the lower Changhia Formation in North China and from Australia. The key features that warrant placing the Hunan specimens into the species include the pygidial outline, the markedly wide and flat borders, the segmentation of the axis, and the nature of the pleural field. Like some of Walcott's specimens, the surface in one of the present pygidia is also minutely punctate. Occurrence. This species is known from the Amphoton Zone of the Changhia Formation in Shandong and Liaoning, North China, and from the Ptychagnostus punctuosus Zone in Australia. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Amphoton alceste (Walcott, 1905) Plate 6, figures 13, 14; Plate 7, figures 1-6 1905 1905 1913 1913

Dolichometopus alceste Walcott, p. 94. Dolichometopus dirce Walcott (in part), p. 96 [cranidium and librigena, non pygidium]. Dolichometopus alceste Walcott, p. 215, pl. 22, figs. 3, 3a, b. Dolichometopus dirce Walcott (in part), p. 218, 219, pl. 22, figs. 5, 5a, non 5b [=Lisania agonius (Walcott, 1905)]. 1942 Amphoton alceste (Walcott); Resser, p. 5. 1942a Amphoton (Amphotonella) alceste (Walcott)" Kobayashi, p. 164, pl. 3, figs. 4, 5. 1965 Amphotonella alceste (Walcott)" Lu, Zhang, Zhu, Qian, and Xiang, p. 114, pl. 17, figs. 17-19. 1977 Amphotom robustum Zhou in Zhou et al., p. 134, 135, pl. 43, figs. 2, 3. 1977 Amphoton (Paramphoton) xiangxiensis Yang in Zhou et al., p. 135, pl. 43, figs. 4, 5. 1978 Amphoton (Paramphoton) xiangxiensis Yang; Yin and Li, p. 448, pl. 159, figs. 13, 14. 1978 Amphoton (Paramphoton) xiangxiensis Yang; Yang, p. 34, pl. 4, figs. 4, 5. 1980 Amphotonella alceste Walcott; Nan, p. 487, pl. 200, fig. 26. 1982 Amphotom robustum Zhou; Liu, p. 302, pl. 214, figs. 7, 8. 1982 Amphoton (Paramphoton) xiangxiensis Yang; Liu, 1982, p. 302, pl. 215, figs. 1, 2. ?1985 Amphoton sp.; Xiang and Zhang, p. 103, pl. 26, figs. 1, 2. 1987 Amphoton alceste (Walcott); Zhang and Jell, p. 65, 66, pl. 21, figs. 1-5; pl. 22, figs. 8-11. 1987 Amphoton deois Walcott; Zhang and Jell, p. 63-65, pl. 17, figs. 4, 5 only. 1987 Anomocarellid indet.; Zhang and Jell, 1987, p. 263,300, pl. 39, fig. 4. • 57-

1996 1996

Amphotonella alceste (Walcott); Guo, Zan, and Luo, p. 97, pl. 41, figs. 1-6, 7a, 8-11, 13, ?12; pl. 42, figs. 1-8. Amphotonella dirce (Walcott); Guo, Zan, and Luo, p. 97, 98, pl. 41, 7b.

Holotype. Partly exfoliated cranidium (Walcott, 1913, pl. 22, fig. 3; Zhang and Jell, 1987, pl. 21, figs. 1, 2, USNM 58249)from south of Yanzhuang, Shandong, China. New material. Six cranidia and one librigena (illustrated specimens NIGP 137345-137348) in collections P108, P134.2, P135, P136.3, and W82.1. Remarks. Amphoton alceste is characterized by having a librigena bearing a genal spine, and a parallel-sided, highly convex glabella with long and well defined S1 furrows extending rearward and inwards. This species has a surface bearing fine, scattered punctae (Walcott, 1905, p. 94; 1913, p. 215) that seem to be present not only externally, but also on exfoliated surfaces of the cephalon (Zhang and Jell, 1987, pl. 22, figs. 1, 2, 5). New cranidia from northwestern Hunan resemble those from North China in glabellar shape and convexity, and in the configuration of the S1 furrows. The associated librigena bears a moderately convex genal field, a genal spine, and fine punctae on an exfoliated surface. The new specimens have palpebral areas that are similar in width to those of the type material and the material of Guo et al. (1996) from Liaoning. In all these specimens, the palpebral areas are narrower than those of A. deois. Amphoton (Paramphoton) xiangxiensis Yang (in Zhou et al., 1977), which is based on a cranidium and a pygidium from the same formation and the same region as the new material of A. alceste, is regarded as junior synonym of A. alceste. The cranidium has a highly convex, parallel-sided glabella with long and obliquely directed S 1 furrows. The holotype cranidium of A. xiangxiensis appears to be conspecific although the glabellar furrows look to be somewhat more shallow. The pygidium is identical with that of A. alceste in bearing axial tubercles on each ring. The new cranidia show slight differences from the material of North China. They bear fine, densely spaced granules on the surface of the test, commonly on the L1 lobe and on the occipital ring. So far there is no specimen from North China reported to have such ornamentation, but since granules on the Hunan material are variably developed, they are considered to be the result of intraspecific variation. Zhang and Jell (1987) synonymized the species Dolichometopus dirce Walcott, 1905, which was based on an incomplete cranidium, a librigena, and a pygidium from the Changhia Formation at Zhangxia, Shandong (Walcott, 1913, pl. 22, figs. 5, 5a, 5b), with A. deois, referring only the holotype cranidium to D. dirce. Zhang and Jell (1987) reassigned the pygidium to Lisania spinosa Resser and Endo and considered the librigena to belong to an anomocarellid or manchuriellid. However, the refigured librigena (Zhang and Jell, 1978, pl. 39, fig. 4) shows that it is almost identical with those assigned to A. alceste by Guo et al. (1996, pl. 42, fig. 7); it is also associated with cranidia of A. alceste. This suggests reassignment of the librigena to A. alceste. The "steep posterior portion of the facial suture" emphasized by Zhang and Jell (1987) to exclude the librigena from Amphoton seems to be the result of the orientation of the specimen when observed. As discussed above, we prefer to assign the cranidium and the librigena to A. alceste, and the pygidium, following Zhang and Jell (1987), to Lisania agonius (Walcott) [=Aojia luna Resser and Endo]. Occurrence. The species is known from the Amphoton Zone at Zhangxia and Yanzhuang, Shandong, and the Peishania-Liopaishania Zone in southern Liaoning, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it • 58.

occurs with trilobites indicative of the top part of the Dorypyge richthofeni Zone to the lower part of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone).

Amphoton sp. cf. A. typicum (Kobayashi, 1942a) Plate 5, figures 10-16; Plate 6, figures 1-9; Plate 16, figure 7b cf. 1942a Amphoton (Sunia) typica Kobayashi, p. 176-177, pl. 1, figures 1-9; pl. 3, figs. 1-3. cf. 1957 Sunia typica Kobayashi; Lu, pl. 140, figs. 6, 7. cf. 1965 Sunia typica Kobayashi; Lu, Zhang, Zhu, Qian, and Xiang, p. 115-116, pl. 18, fig. 9. 1978 Amphoton (Paramphoton) parallela (Resser and Endo); Yang, p. 34, pl. 5, figs. la, b. non 1937 Amphoton parallela Resser and Endo, p. 206, pl. 38, figs. 2-8, 10-13; pl. 39, figs. 19, 20 [=Amphoton derios (Walcott)]. ?1978 Amphoton (Paramphoton) sp.; Yang, p. 34, pl. 5, figs. 2a, b, 3. 2001b Amphoton deois (Walcott); Peng, Babcock, and Lin, p. 102, pl. 3, figs. 4, 5.

Material. Nearly 30 cranidia, librigenae, pygidia (illustrated specimens NIGP 137327-137340, 137453) in collections P102.5, P108, P112.6, P122.4, P123.6, P125, P126, P131.7, and W82.1. Description. Cranidium moderately convex, length four-fifths width. Glabella long, with length (excluding occipital ring) more than twice basal width, constricted at about mid-length and then expanded forwardly, rounded anteriorly with faint sagittal keel on exfoliated surface; three to four pairs of lateral furrows present: S 1 deep and straight, rearward and inward; $2 shallow and short, parallel to S 1; $3 and $4 short and faint, exfoliated specimen showing that they run nearly inwards; occipital furrow deep, nearly transverse medially, outward- and forward-directed distally; occipital ring markedly narrowing abaxially with long spine sagittally. Palpebral lobe long, well defined by palpebral furrow, with anterior end near $4, and posterior end opposite midlength of L1. Anterior border short, gently convex, separated from glabella by wide and relatively deep border furrow. Palpebral area of fixigena moderately convex, 0.28-0.45 as wide as glabella; posterior area of fixigena short (exs.) and long (tr.), with deep and wide posterior border furrow. Anterior branch of facial suture short, slightly divergent forward, posterior branch long, running outward and slightly rearward. Glabella and, sometimes, fixigena ornamented fine, dense granules. Librigena with stout spine and flat and moderately wide lateral border being upturned anteriorly. Most pygidia are largely exfoliated. One pygidium (P1. 6, fig. 5) shows a testaceous fight pleural area. Pygidium width twice length, with wide facet sloping steeply forward. Axis short and wide, occupying 0.75-0.80 of pygidial length, with three tings and a large, semicircular terminal piece extending as a stout postaxial ridge onto posterior margin. Pleural area with four segments separated by thin and shallow interpleural furrows; pleura with ridge-like anterior and posterior bands and very wide pleural furrow. The posterior band is less convex than the anterior band. Lateral and posterior borders broad and fiat, almost of uniform width. Structure of exfoliated surfaced almost identical with that of the test surface. Pygidial surface in some specimens with densely spaced granules. Remarks. Specimens in the large collection from northwestern Hunan show a close similarity to the specimens described under the name Amphoton (Sunia) typica by Kobayashi (1942a). The cranidium is characterized by having a long glabella that is constricted at cranidial midlength and expanded anteriorly, genal spine-bearing librigenae, and a long occipital spine. The specimens are • 59-

almost identical in all respects with A. typicum except for having more distinct $2 furrows and a less stout occipital spine. Because the type material of A. typicum is preserved in shale and is more or less flattened, whereas the specimens from Hunan are in limestone, the appearance of $2 on the present material, and the higher convexity of the glabella as well, may be caused by preservation rather than morphological differences. In the pygidium, the present material differs in having a more transverse outline, a shorter and wider axis, and much wider borders. The present specimens also closely resemble A. deois, but the librigena in the latter bears no genal spine, and the glabellar furrows are shallower. The present pygidium seems indistinguishable from that of A. deois. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper Dorypyge richthofeni Zone to the Pianaspis sinensis Zone (equivalent to the Ptychagnostus punctuosus Zone and the Goniagnostus nathorsti Zone). Genus FUCHOUIA Resser and Endo in Kobayashi, 1935 Fuchouia Resser and Endo in Kobayashi, 1935, p. 136; Resser and Endo, 1937, p. 225; Hupr, 1953a, p. 165; 1953b, p. 184; Lu, 1957, p. 264; Egorova, Xiang, Li, Nan, and Guo, 1963, p. 30; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 116; Zhou, Liu, Meng, and Sun, 1977, p. 135; Yin and Li, 1978, p. 446; Yang, 1978, p. 33; Zhang, 1981, p. 157; t3pik, 1982, p. 26, 27; Liu, 1982, p. 302; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han and Wei, 1983, p. 61; Zhang and Jell, 1987, p. 66, 67; Lu and ]_,in, 1989, p. 124, 125" Jell and Hughes, 1997, p. 98; Jago and Brown, 2001, p. 11. Amphoton (Fuchouia) Resser and Endo, 1937, p. 227; Kobayashi, 1942a, p. 166, 167; Poulsen in Moore, 1959, p. 0222. Fuchouia (Parafuchouia) Lu and Chien in Lu et al., 1974b, p. 103; Yin and Li, 1978, p. 446; Zhu, Lin, and Zhang, 1979, p. 87; Lu and Qian, 1983b, p. 33. Borovikovia Kraskov, 1977, p. 59. Fuchouia (Pseudofuchouia) Zhang, 1981, p. 158. Parafuchouia Lu and Chien; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 62. Type species. Bathyuriscus manchuriensis Walcott, 1911 (p. 97, pl. 16, fig. 4) from the Changhia Formation, Changxingdao Island, southem Liaoning, China; by original designation. Other species. Apart from the type species, the following species have been assigned to Fuchouia: Fuchouia chai Lu, 1957 (p. 264, pl. 140, figs. 11, 12) from northwestern Hunan and eastern Guizhou; Fuchouia spinosa Chang, 1959 (Zhang, p. 210, pl. 4, figs. 12-16) from the Changhsia Formation, Boshan (Poshan), Shandong, which was considered to be a possible variant population of F. manchuriensis by Zhang and Jell (1987, p. 68); Amphoton bensoni Opik, 1961, p. 136-141, p. 11, text-fig. 47, from Queensland, Australia; Amphoton limbatum Zhou in Lu et al., 1974b (p. 102, 103, pl. 40, figs. 10, 11) from the Jialao Formation, near Nangao, Kaili, Guizhou; Amphoton promptum Zhou in Lu et al., 1974b (p. 102, 103, pl. 40, fig. 12) is from the same formation and locality as A. limbatum Zhou, and may be synonymous with the latter; Fuchouia delicata Lu, Zhang and Zhu (1963, p. 24, pl. 1, fig. 3), locality unknown; Fuchouia (Parafuchouia) elongata Lu and Qian (in Lu et al., 1974b, p. 103, pl. 40, figs. 13, 14) from the Jiaolao Formation, near Xingren, Kaili, Guizhou; Fuchouia oratolimba Yang in Zhou et al., 1977 (p. 135, 136, pl. 43, fig. 8, ?fig. 9) from the middle part of the Huaqiao Formation, northwestern Hunan; Fuchouia angusta Yang, • 60

•

1978 (p. 33, 34, pl. 4, fig. 8), a junior synonym of F. oratolimba, also from the middle part of the Huaqiao Formation, northwestern Hunan; Fuchouia kuruktagensis Zhang, 1981 (p. 157, 158, pl. 60, fig. 5) from the Mohershan Formation, near Kuruktag, Xinjiang; Fuchouia (Pseudofuchouia) mohershanensis Zhang, 1981 (p. 158, pl. 60, figs. 6, 7), a junior synonym of F. oratolimba from the Mohershan Formation, near Kuruktag; Fuchouia formosa Liu, 1982 (p. 302, 303, pl. 215, fig. 10), a junior synonym of F. oratolimba from the Huaqiao Formation, near Yongshun, northwestern Hunan. Fuchouiafecunda Opik (1982, p. 27-39, pl. 6-8; pl. 9, fig. 1; pl. 10, figs. 3, 4; pl. 11-13; text-figs. 8a-17), from Queensland and the Northern Territory, Australia; Fuchouia aft. F. fecunda Opik (1982, p. 39, pl. 3, fig. 6) from Queensland, Australia; Fuchouia labda Opik (1982, p. 39, 40, pl. 2, figs. 5, 6; pl. 3, fig. 1; pl. 31, fig. 1; text-fig. 18), Fuchouia atoba Opik (1982, p. 40, 41, pl. 3, figs. 7-9; pl. 9, figs. 3a-c; pl. 10, figs. 1, 2a, b), from Queensland and the Northern Territory, Australia; Fuchouia sp. aft. F. atoba Opik (1982, p. 41, pl. 3, fig. 2), from Queensland and the Northern Territory, Australia; and Fuchouia morstonensis Opik (1982, p. 42, 43, pl. 3, figs. 2, 4, 5, 3?) from Queensland, Australia; Fuchouia (Parafuchouia) transversa Lu and Qian (1983b, p. 34, 35), a junior synonym of Fuchouia (Parafuchouia) elongata Lu and Qian in Lu et al., 1974b, from eastern Guizhou; Fuchouia tetrasolena Ju in Qiu et al., 1983 (p. 62, pl. 21, figs. 3-5) from the Yangliugang Formation (Middle Cambrian), Tonglu, Zhejiang; Fuchouia antiqua Rosov (in Lisogor et aL, 1988, p. 69, 70, pl. 6, figs. 1, 2), from Maliy Karatau, Kazakhstan; and Fuchouia sp. (Lisogor et al., 1988, p. 70, pl. 6, fig. 5) from Maliy Karatau, Kazakhstan. Fuchouia aft. F. manchuriensis (Walcott) described also from Maliy Karatau, Kazakhstan (Lisogor et al., 1988, p. 68, 69, pl. 6, fig. 4) probably belongs to Shengia because of the presence of long eye ridges. Two species described from the Xijiadian Formation (Middle Cambrian), Xichuan, Henan, Fuchouia sp. cf. F. oratolimba Yang (in Yang et al., 1991, p. 131, pl. 10, figs. 11-13; 1993, p. 168, pl. 10, figs. 11-13) from the Aspidagnostus primitivus-A, orientalis Zone-Linguagnostus kjerulfi-Diplagnostus qinlingensis Zone and Fuchouia (Parafuchouia) quadratoglabella Yang (in Yang et al., 1991, p. 131, 132, pl. 11, figs. 2-7; 1993, p. 168, 169, pl. 11, figs. 2-7 ) from the Ptychagnostus gibbus-Doryagnostus incertus Zone, are of uncertain assignment within Fuchouia. Two additional Fuchouia species, described as Fuchouia (Parafuchouia) sp. 1 and F. (P.) sp. 2, from the Xijiadian Formation are, respectively, a meraspid cranidium and indeterminable fragments (Yang in Yang et al., 1991, pl. 11, figs. 8-11; 1999, pl. 11, figs. 8-11). They are associated with F. (P.) quadratoglabella and may be conspecific with it. The species described from southern Kazakhstan under the name of Borovikovia juvenilia Kraskov (1977, p. 59, 60, pl. 13, figs. 8-11, text-fig. 8) is a junior synonym of Fuchouia oratolimba Yang (in Zhou et al., 1977). An indeterminate Fuchouia was described from the Kurgiakh Formation (middle Cambrian), Zanskar, northwestern Himalaya, India (Whittington, 1986, pl. 19, fig. 5; pl. 20, figs. 2, 3). This work adds two new species, F. bullba sp. nov. and F. sixinensis sp. nov., from northwestern Hunan. Remarks. The genetic concept of Fuchouia and its differences from Amphoton have been discussed by Opik (1982) and Zhang and Jell (1987). Their concept is followed here. We concur with Zhang and Jell (1987) in regarding the subgenus Fuchouia (Parafuchouia) as a junior synonym of Fuchouia. The type species, Fuchouia (Parafuchouia) elongata Lu and Chien, is characterized by the presence of a preglabellar field, a long palpebral lobe and a short posterior area on the fixigena, which together are considered to be the major distinctive features for the subgenus. However, as noted by Zhang and Jell (1987, p. 67), the lectotype of Fuchouia manchuriensis, the type species of Fuchouia, and other material assigned to the genus also bear a short preglabellar field, a long palpebral lobe, and a short posterior area on the fixigena. They concluded that there is no basis for separation of Fuchouia (Parafuchouia). Opik (1982) sorted Fuchouia into three species groups according to the length of the palpebral

• 61 •

lobes. Species belonging to Group B and Group C have long palpebral lobes and narrow posterior areas of the fixigenae, and are comparable in those features to F. (Parafuchouia). It is obvious that 0pik's (1982) concept of Fuchouia embraces already Fuchouia (Parafuchouia). Some Australian species assigned to Fuchouia, like F. elongata and F. fecunda are closely similar to Amphoton because they have long palpebral lobes. Based on such similarities, Lu and Lin (1989, p. 125, 241) suggested either reviving Fuchouia (Parafuchouia) as a subgenus of Amphoton [=Sunia] or treating it as a separate genus. This suggestion is rejected here. We prefer to retain these species in Fuchouia because the species are distinguishable from Amphoton in some key respects. The distinct preglabellar field, the divergent anterior branches of the facial suture, the pleural fields with diagonal pleural furrows and differently shaped pleural bands, and the absence of pygidial borders all suggest that these species belong in Fuchouia. Fuchouia (Pseudofuchouia) is considered to be a junior synonym of Fuchouia. As discussed by Zhang (1981, p. 158), F. (Pseudofuchouia) differs from Fuchouia only in the presence of a preglabellar field, which is also a feature of Fuchouia. Borovikovia Kraskov (1977) was erected on the basis of poor material. The holotype of the type species, Borovikovia juvenilia, is an incomplete cranidium with three incomplete thoracic segments attached (Kraskov, 1977, pl. 13, fig. 8). In comparison with other specimens of the species, this cranidium is small in size and, as indicated by the specific name, belongs to an immature individual. The strongly expanded glabella is comparable to that of meraspid Fuchouia. The associated paratype pygidium and fragmental paratype cranidia (ibid., pl. 13, figs. 9-11; text-fig. 8) show that those sclerites are certainly conspecific with Fuchouia oratolimba Yang in Zhou et al., 1977 (p. 135, pl. 43, figs. 5, 6, 8). B. juvenilia is therefore regarded as a junior synonym of Fuchouia oratolimba, and Borovikovia is regarded as a junior synonym of Fuchouia. Both Opik (1982) and Zhang and Jell (1987) noted that no type specimen had been designated for the type species, Fuchouia manchuriensis, and they designated and refigured a cranidium from Walcott's collection as the lectotype (Zhang and Jell, 1987, p. 67, p. 331, pl. 24, fig. 1, upper fight; also pl. 122, fig. 10, USNM 57587 ). A pygidium on the same slab with that cranidium was also mistakenly designated as the lectotype of F. manchuriensis by the same authors in the same publication (Zhang and Jell, p. 295, pl. 24, fig. 1, lower left). Fuchouia chiai Lu, 1957

Plate 7, figures 7, 8 1957 1962 non 1963 non 1964

1965 1977 1978 1981 non 1982

Fuchouia chiai Lu, p. 264, pl. 140, figs. 11, 12. Fuchouia chiai Lu; Lu, Zhu, and Qian, p. 31, pl. 4, figs. 6, 9. Fuchouia chiai Lu; Egorova, Xiang, Li, Nan, and Guo, p. 31, 32, pl. 6, figs. 3-5 [=Fuchouia oratolimba Yang in Zhou et al., 1977], ?6. Fuchouia chiai Lu; Lu and Qian, p. 30, pl. 4, figs. 4, 5 [=Fuchouia oratolimba Yang in Zhou et al., 1977]. Fuchouia chiai Lu (in part); Lu, Zhang, Zhu, Qian, and Xiang, p. 117, 118, pl. 18, figs. 21, 22 non 18-20 [=Fuchouia oratolimba Yang in Zhou et al., 1977]. Fuchouia chiai Lu; Zhou, Liu, Meng, and Sun, p. 135, pl. 43, figs. 6, 7. Fuchouia chiai Lu; Yin and Li, p. 446, pl. 157, fig. 7; Liu, 1982, p. 302, pl. 214, figs. 6, 11. Fuchouia delicata Lu; Zhang, p. 157, pl. 61, figs. 16, 17. Fuchouia chiai Lu; Yang, p. 303, 304, pl. 2, figs. 12-20 [?=Fuchouia sp. cf. F. spinosa

Chang, 1959 as interpreted herein]. non 1983b Fuchouia chiai Lu; Lu and Qian, p. 29-33, pl. 4, figs. 1-9 [=Fuchouia kuruktagensis Chang,

1981]. • 62-

non 1989

Fuchouia chiai Lu; Lu and Lin, p. 126, 127, pl. 17, fig. 9 [?=Fuchouia kuruktagensis Chang,

1981 as interpreted herein]. non 1991

Fuchouia chiai Lu; Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p. 131, pl. 11, figs. 1, 2 [=Fuchouia sp. indet.]. non 1991 Fuchouia chiai Lu; Lin, p. 131, pl. 3, figs. 3, 4 [=Fuchouia kuruktagensis Zhang, 1981]. non 1993 Fuchouia chiai Lu; Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p. 168, pl. 11, figs. 1, 2 [=Fuchouia sp. indet.]. non 1995 Fuchouia chiai Lu; Zhang in Zhang et al., p. 22, 23, pl. 5, figs. 7-9 [?=Fuchouia manchuriensis (Walcott)]. 2001b Fuchouia chiai Lu; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 3. 2001e Fuchouia chiai Lu; Peng, Babcock, Lin, Chen, and Zhu, p. 159, figs. 10.22-10.24. Lectotype. The cranidium from the original material of Lu (1957, pl. 140, fig. 1) is here designated as lectotype. This specimen was refigured at least twice (Lu et al., 1962, pl. 4, fig. 6; Lu et al., 1965, pl. 18, fig. 21). A "lectotype" was also designated by Zhang in Zhang et al. (1995, p. 22) for this

species. This "lectotype" is rejected here because it was not selected from the syntypes of Lu, 1957, but from a specimen described subsequently by Egorova et al. (1963, pl. 6, fig. 3; refigured also in Lu et al., 1965, pl. 18, fig. 18). For this reason, it loses its status of lectotype (International Commission on Zoological Nomenclature, 1999, p. 82, Article 74.2). The specimen previously selected as a "lectotype" could serve as a neotype, but this is unnecessary because the syntypic suite of Lu (1957) still exists. N e w material. One cephalon and five cranidia (illustrated specimens NIGP 137349, 137350) in

collections P125, P130.5, P134.2, P164.2, P167, P171.5, P184,? P199.9. Emended diagnosis. Fuchouia with an obtusely angled anterior margin, a moderately long glabella

that is gently expanded anteriorly; palpebral lobe short, opposite $2 furrow; fiat anterior border strongly upturned; preglabellar field absent; librigena with moderately wide border, upturned anteriorly; genal spine slender and short. Remarks. The type material of Fuchouia chiai includes a cranidium and a pygidium from the

Huaqiao Formation of the northwestern Hunan-eastern Guizhou border area (Lu, 1957, pl. 140, figs. 1, 2). No holotype was originally designated for the species when it was erected by Lu. The cranidium has a steeply upturned anterior border that has an angular anterior margin, has a glabella that is slightly expanded anteriorly, and lacks a preglabellar field. Some specimens assigned subsequently to the species by Egorova et al. (1963, pl. 6, figs. 3-6) are not conspecific with the type material because they lack these key features. The material of Egorova et al. (1963) includes three incomplete exoskeletons in one collection and a pygidium in another collection. All the exoskeletons are early holaspid, and are rather small in size. Among these specimens, the larger specimens show parallel-sided glabellas, and one exoskeleton (Egorova et al., 1963, fig. 3), which is the only specimen with an anterior border, shows a narrow and convex anterior border and a preglabellar field. Cranidial features of these exoskeletons suggest that they are not conspecific with F. chai but with F. oratolimba Yang in Zhou et al., 1977 (see P1. 10, figs. 1-3 herein), and should be referred to that species. The concept of Fuchouia chiai had long been confused because these exoskeletons were subsequently refigured together with Lu's (1957) collection under the name E. chiai in the important atlas of Chinese trilobites (complied by Lu et al., 1965). More unfortunately, such confusion has been enhanced since one of the exoskeletons in the material of Egorova et al., 1963 (pl. 6, fig. 3) was designated as lectotype of Fuchouia chiai by Zhang et al. • 63

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(1995, p. 22). Because the "lectotype" of Zhang et al. (1995) is neither from among the syntypes of Lu, 1957 nor conspecific with Fuchouia chiai, according to ICZN, Article 74.2 (4th edition, 2000), it should not be considered as a lectotype. Occurrence. This species was previously reported from the Huaqiao Formation of northwestern Hunan and eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone and the Lejopyge laevigata Zone). Fuchouia kuruktagensis Zhang, 1981

Plate 8, figures 1-15; Plate 9, figures 1-8 ?1942a Amphoton (Fuchouia) manchuriensis (Walcott); Kobayashi, p. 178, 179, pl. 1, figs. 12-19; pl. 3, 7-9 (reconstruction). 1981 Fuchouiakuruktagensis Zhang, 1981, p. 157, 158, pl. 60, fig. 5. 1982 Fuchouiaoratolimba Yang; Liu, p. 302, pl. 215, figs. 3, 4. 1983b Fuchouia chiai Lu; Lu and Qian, p. 29-33, pl. 4, figs. 1-9. ?1989 Fuchouiachiai Lu; Lu and Lin, p. 126, 127, pl. 17, fig. 9. ?1991 Fuchouiacf. F. oratolimba Yang; Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p 131, pl. 10, figs. 11-13. ?1993 Fuchouiacf. F. oratolimba Yang; Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p 168, pl. 10, figs. 11-13. 1991 Fuchouia chiai Lu; Lin, p. 131, pl. 3, figs. 3, 4. 2001b Fuchouia oratolimba Yang; Peng, Babcock, Lin, p. 101, pl. 2, figs. 10, 11. Holotype. An early holaspid exoskeleton (Zhang, 1981, pl. 60, fig. 5; XTR-107), from the Mohershan Formation, Kuruktag, Xinjiang, China. New material. More than 40 sclerites, including an exoskeleton, cranidia in ontogenetic series, librigena, hypostomes, and pygidia (illustrated specimens NIGP 137353-137372) in collections P64.5, P68.8, W56.7, W77.8, W82.1, W82.2, and W88.1. Remarks. The cranidium of F. kuruktagensis is about 10 mm long, and shares all observed characters, including the glabellar outline, the nature of the lateral glabellar furrows, and the presence of a tiny intergenal spine at the distal end of the posterior margin, with cranidia of similar size in our collection. The pygidia from Hunan are quite similar to the holotype pygidium in the shape, segmentation, and the presence of clearly defined interpleural furrows on the first three or four segments of the pleural regions. The new material from Hunan includes an ontogenetic series, providing additional features for the species, especially in the characters of the late holaspid stages. These include a narrow preglabellar field, the presence of a short occipital spine, and the rectangular-shaped, more effaced glabella. The first recorded hypostomes added hypostomal morphology to the species concept. The new material is also similar to Fuchouia spinosa Zhang (1959, p. 235, pl. 4, figs. 12-16; text-figs. 26, 27) from North China, which is characterized by having a rectangular glabella, tiny intergenal spines, and a short occipital spine in the cranidium. However, F. spinosa differs in having an upturned anterior cranidial border and pygidial border spines, and in lacking a preglabellar field. According to Zhang (1959), F. spinosa has only two pairs of lateral glabellar furrows, but in

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our collection of F. kuruktagensis, some specimens show faint $3 and $4 furrows. This may be another means of distinguishing F. kuruktagensis. So far, there is only one adult cranidium known for F. spinosa and such a determination must wait until additional cranidial material of F. spinosa is reported. Because the preglabellar field is variably present in F. kuruktagensis, as are the marginal spines on the pygidium of F. spinosa, these two species may eventually prove to be synonymous. Pending the acquisition of more information about the North China species, we assign the present material to F. kuruktagensis. As noted by Zhang (1959, p. 235), some cranidia described as Amphoton (Fuchouia) manchuriensis by Kobayashi (1942a, pl. 1, figs. 12-19) contain intergenal spines. The largest cranidium in Kobayashi's material (1942a, pl. 1, fig. 15) is probably an early holaspid. It seems to have a short preglabellar field, and both intergenal and occipital spines. The associated pygidia resemble closely the pygidia assigned now to F. kuruktagensis in having clearly defined interpleural furrows and lacking border spines. Kobayashi's (1942a) material is probably conspecific with specimens in our collections, and until more is known, it is questionably referred to the present species. Occurrence. This species was originally reported from the Mohershan Formation of Kuruktag, Xinjiang, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone to the lower part of the Pianaspis sinensis Zone (equivalent to the Ptychagnostus atavus Zone through the lower part of the Goniagnostus nathorsti Zone). Fuchouia oratolimba Yang in Zhou et al., 1977

Plate 10, figures 1-11" Plate 11, figures 1-5 1963 1964 1965

Fuchouia chiai Lu; Egorova, Xiang, Li et al., p. 31, 32, pl. 6, figs. 3-6. Fuchouia chiai Lu; Lu and Qian, 1964, p. 30, pl. 4, figs. 4, 5. Fuchouia chiai Lu (in part); Lu, Zhang, Zhu, Qian, and Xiang, p. 117, 118, pl. 18, figs.

1977

Fuchouia oratolimba Yang in Zhou et al. (in part), p. 135, 136, pl. 43, fig. 8, non fig. 9 [?=Fuchouia bulba sp. nov.]. Borovikovia juvenilia Kraskov, p. 59, 60, pl. 13, figs. 8-11, text-fig. 8. Fuchouia angusta Yang; Yin and Li, p. 446, pl. 159, fig. 3. Fuchouia oratolimba Yang (in part); Yang, p. 33, 34, pl. 4, figs. 5, 6, non fig. 7 [?=Fuchouia bulba sp. nov.]. Fuchouia angusta Yang, p. 33, 34, pl. 4, fig. 8. Fuchouia (Pseudofuchouia) mohershanensis Zhang, p. 158, pl. 60, figs. 6, 7. Fuchouia oratolimba Yang; Liu, p. 302, pl. 215, figs. 3, 4 [=Fuchouia kuruktagensis Zhang,

18-20 only.

1977 1978 1978 1978 1981 non 1982

1981]. 1982 Fuchouia formosa Liu, p. 302, 303, pl. 215, fig. 10. 1983 Fuchouia dingxiangensis Zhang in Qiu et al., p. 61, pl. 21, figs. 6, 7. 1989 Fuchouia oratolimba Yang; Lu and Lin, p. 125, 126, pl. 17, figs. 1-8. 2001b Fuchouia angusta Yang; Peng, Babcock, and Lin, p. 103, 104, pl. 7, figs. 5, 6; pl. 10, figs. 5, 11. Holotype. Early holaspid cranidium (Yang in Zhou et al., 1977, pl. 43, fig. 8; refigured in Yang, •

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1978, pl. 4, fig. 6, and in Liu, 1982, pl. 215, fig. 3), from the Parablackwelderia [=Paradamesops] jimaensis-Torifera [=Cyclolorenzella] tuma Zone, at Jiudiantang, Xinhuang, northwestern Hunan. New material. More than 80 specimens, including cranidia, pygidia, librigenae, and thoracic segments (illustrated specimens NIGP 137378-137393) in collections P209, P209.5, P237.25(?), P260.08, P261, P261.35, P261.5, P269, P273.46, P273.52, P273.6, P273.66, P273.8, P277, P277.6, P278.04, P278.1, P279, P279.7, P282.95, P283, P283.47, P283.75, P301, W171.9, W188.8, W197.5, and W 198.45. Emended diagnosis. Fuchouia with parallel-sided, elongated glabella bearing 4 pairs of lateral furrows; anterior border narrow, arched gently forward or rearward; preglabellar field present, wider than anterior border; palpebral lobe placed anterior to cranidial midlength; occipital node positioned near posterior margin of occipital ring. Thorax has 8 segments with broad pleural furrows and thin pleural bands. Pygidium with long, thin, clearly segmented axis, deeply furrowed pleural field, and narrow border. Description. Glabella rectangular in outline, length twice width, obtusely rounded anteriorly, well defined laterally by slightly sinuous axial furrows, with 4 pairs of lateral furrows; S1 longest, deep and straight, diagonally oriented, nearly isolating L 1 from glabella; $2 deeply to weakly impressed; $3 faint, pit-like, not extending to axial furrow; $4 short, transversely oriented; occipital furrow shallow and broad, deep at sides; occipital ring prominent, beating small node lying posteriorly; anterior border narrow and elevated, arched gently forward to rearward, well defined by border furrow; preglabellar field fiat, depressed in front of glabella; palpebral lobe short, located just anterior of the cranidial midlength, with posterior end opposite the middle of L2 or $2. Anterior branch of facial suture short, diverging forward at angle of 10°-30 ° to sagittal line; posterior branch moderately sigmoidal, strongly diverging rearward, enclosing short and long, blade-like posterolateral projection of fixigena. Librigena with narrow, ridge-like border, broad genal field with densely anastomosing ridges, and slender genal spine. Thorax of 8 segments with axis tapering gently toward posterior; medial nodes observed on axial tings 7 and 8 (P1. 10, figs. 1, 7); but partly exfoliated specimen suggests presence of medial nodes on more anterior rings; each ring bears pair of pits laterally; pleural field width twice width of axis, with each pleuron bearing tiny spine at posterolateral comer; thin anterior band widening abaxially, wide pleural furrow branches into two thin furrows adaxially. Pygidium with narrow borders; axis with 7-8 tings and semicircular terminal piece; pleural field bears incised interpleural furrows and broad pleural furrows; pleura with anterior band widening abaxially; pleural furrow diagonal, narrowing abaxially. Remarks. Ontogenetic series show that the glabella changes in shape from somewhat expanded anteriorly in small cranidia to parallel-sided in large holaspid specimens. The axial furrow becomes sinuous in the cranidia of large holaspids. The anterior margin changes from anteriorly curved to transverse or posteriorly curved. The S 1 furrow becomes bifurcate, the position of the $3 furrow increases from axial furrow, and the $4 furrow extends obliquely to transversely. The anterior branch of the facial suture becomes less divergent forward in larger specimens. The preglabellar field is present in all known stages and its proportional length remains nearly unchanged through ontogeny. Type material of F. oratolimba Yang (in Zhou et al., 1977, pl. 43, figs. 8, 9) includes the holotype cranidium from Jiudiantang, Xinhuang, northwestern Hunan, and a paratype pygidium from a different horizon in Huaqiao, Baojing, northwestern Hunan, some 150 km north of • 66

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Jiudiantang. An additional incomplete exoskeleton, originally included as a paratype, was first illustrated by Yang in a later publication (Yang, 1978, pl. 4, fig. 5). The specimen comes from the same collection as the holotype but has a much larger cranidium than the holotype. Our material, complete with ontogenetic information, shows that the holotype is not a large holaspid cranidium. Instead, it notably represents an early holaspid. The paratype pygidium is not conspecific with the two other specimens in the type suite. Fuchouia angusta is based on a single pygidium (Yang, 1982, pl. 4, fig. 8) that is identical in all respects to pygidia known from the complete specimens of F. oratolimba. It leaves little doubt that this species should be considered as a junior synonym of F. oratolimba. Borovikovia juvenilia (Kraskov, 1977) from the Kendyiktas Range, southern Kazakhstan, is indistinguishable from F. oratolimba in every respect, and the two species are here considered to be synonymous. Both F. oratolimba and B. juvenilia species were described in 1977. F. oratolimba, which was published in February, 1977 (Yang in Zhou et al., 1977), has seniority over B. juvenilia, which was published in May, 1977 (Kraskov, 1977). F. oratolimba is a distinctive species of Fuchouia, characterized by a parallel-sided glabella with four pairs of lateral furrows, a narrow and convex anterior border, a preglabellar field, and a multisegmented pygidium with clearly incised interpleural furrows. By these features it can be easily differentiated from all other species of Fuchouia. Occurrence. This species was first reported from the Parablackwelderia [=Paradamesops] jimaensis-Torifera [=Cyclolorenzella] tuma Zone, at Jiudiantang, Xinhuang, northwestern Hunan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Pianaspis sinensis Zone to the Wanshania wanshanensis Zone (equivalent to the Lejopyge laevigata Zone through the Proagnostus bulbus Zone). Fuchouia bulba sp. nov.

Plate 11, figures 6-14; Plate 12, figures 1-12; Plate 13, figures 1-13 ?1977 ?1978 ?1982 2001b

Fuchouiaoratolimba Yang in Zhou et al. (in part), p. 135, 136, pl. 43, fig. 9 only. Fuchouiaoratolimba Yang (in part); Yang, p. 33, 34, pl. 4, fig. 7 only. Fuchouiaoratolimba Yang (in part); Liu, p. 302, pl. 215, fig. 4 only. Fuchuoia sp. 2; Peng, Babcock, and Lin, p. 102, pl. 4, fig. 15.

Etymology. From Latin bulba, bulb; referring to the glabellar shape. Holotype. A testaceous cranidium (P1. 12, figs. 7, 8; NIGP 137409) from collection P131.7. Other material. Numerous exoskeletons, cranidia, librigenae, hypostomes, and pygidia in collections P130.5, P131.7, P136.3, P137.8, P145.5, W82.1, W98, W99.3, W101, W105, and W138.9, among which one exoskeleton, 17 cranidia, four librigenae, two hypostomes, and eight pygidia (NIGP 137394-137324) are illustrated herein. Diagnosis. Fuchouia with gently arched anterior cranidial margin; bulb-like glabella, well rounded anteriorly; four pairs of lateral furrows with S1 bifurcated, variably defined; $3 obsolete; preglabellar field absent; librigena with genal spine that is wide proximally; pygidium with

.67"

interpleural furrows effaced except for first one or two.

Description. Anterior border convex, narrow to moderately wide; preglabellar field absent. Glabella slightly expanded forward to the position of $2, more greatly expanded at the position of middle of L4, rounded anteriorly; 4 pairs of lateral furrows, but commonly only S 1, $2, and $4 present on testaceous surface; S1 moderately long, deep and bifurcated; $2 and $4 short, weak; occipital furrow shallow medially, deep at sides; occipital ring narrowing abaxially with tiny node close to posterior margin. Palpebral lobe short, extending between positions of the middle of L2 and $3. Anterior branch of facial suture diverging forward to border furrow, then turning inward to cross anterior border diagonally; posterior branch sigmoidal; posterolateral projection much wider than long. Librigena with moderately wide lateral border defined by wide shallow border furrow; genal field wide, gently convex, smooth, bearing genal spine that is wide at proximal end. Hypostome with obtusely angled anterior margin (hypostomal suture); median body large, with ovate anterior lobe and crescentic, gently convex posterior lobe; anterior wing narrow (tr.), steeply sloping outward; border furrow broad and shallow; lateral border ridge-like, obtusely angled, turning near distal end of middle furrow. Thorax with eight segments, axis slightly narrower than pleural region; pleura pointed outward, with shallow pleural furrow, anterior band convex, expanding distally; posterior band wide and weak.

Pygidium with weakly defined border; axis with four clearly defined rings and one obscure ring, and semicircular terminal piece; first one or two interpleural furrows obscure, other effaced; pleural furrows wide and shallow, extending diagonally, width nearly uniform.

Remarks. The new species is most similar to Fuchuoia chiai, from which it differs in having a more expanded glabella anteriorly, a convex rather than upturned anterior border, an anterior margin that is gently curved rather than angled forward, and a stronger, broad-based genal spine on the librigena. The pygidium is similar to that of F. kuruktagensis in the pattern of segmentation and the outline, but the pygidium of F. kuruktagensis differs in having clearly defined interpleural furrows. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone and the lower part of the Lejopyge laevigata Zone). Fuchouia sixinensis sp. nov. Plate 9, figures 9-14 2001b Fuchouiasp. 1; Peng, Babcock, and Lin, p. 102, pl. 4, fig. 12.

Etymology. From Chinese, Sixincun, in reference to Sixin Village, located just to the south of the Paibi section; the Paibi section yielded most of the type specimens of the new species. Holotype. Incomplete cranidium, NIGP 137376 (P1.9, fig. 12) from collection collection P82.5. Other material. Eight cranidia in collections P64.5, P71.4, P82.5, P84.3, W42, and W64.7, among which four specimens are figured as paratypes (NIGP 137373-137375; 137377).

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Diagnosis. Fuchouia with rectangular, relatively short glabella, with four pairs of narrow lateral furrows; palpebral lobe at about cranidial midlength; anterior border thin and elevated, curved slightly forward; preglabellar field wide (sag.); occipital ring with stout spine. Description. Anteror border thin, elevated; preglabellar field flat and slightly depressed medially; glabella length about 1.5 times width, anterior obtusely rounded, S1 deeply incised, diagonally directed; $2 narrow, with abaxial end incised, isolated from axial furrow; $3 faint, directed inwardly and forwardly, separated from axial furrow; $4 weak, pit-like, lying close to axial furrow; palpebral lobe situated at midlength of cranidium, posterior end opposite distal end of S 1; occipital furrow deep at side; occipital ring with stout spine; anterior branch of facial suture extending forward and outward at angles of 15°-20 ° to the sagittal line; posterior branch gently sigmoidal, strongly divergent rearward. Remarks. The presence of a preglabellar field, the thin, elevated anterior border, and the unexpanded glabella are characters most reminiscent of Fuchouia oratolimba. The latter, however, can be differentiated from the new species by the longer proportion of its glabella, deeper and wider lateral furrows, a more anteriorly placed palpebral lobe, and by the absence of an occipital spine. The new species resembles F. manchuriensis (Walcott, 1911; pl. 16, fig. 4; Zhang and Jell, 1987, pl. 24, figs. 1-5, 7-11; pl. 122, fig. 10), the type species of the genus from North China, in having a relatively short, rectangular-shaped glabella. The position and size of the palpebral lobe and the course of the facial suture also seem identical. However, the type species differs partly in the absence of a preglabellar field. In the new species, the preglabellar field is present. Also, the new species possesses an occipital spine; F. manchuriensis lacks an occipital spine. It may be further differentiated by lacking $3 and $4 furrows. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and W angcun sections, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone and the lower part of the Ptychagnostus punctuosus Zone). Fuchouia sp. indet. Plate 7, figures 9-11

Material. Two cranidia (NIGP 137351, 137352), in collections W111.3 and W146.2. Remarks. Material includes two fragmental cranidia with a more or less inflated, forwardly expanded glabella. The S1 furrow is deeply impressed and bifurcated, the $2 furrow is weakly indicated, and the $4 furrow is deeply pitted. They are closely comparable to the holotype cranidium of Fuchouia quadratoglabella Yang (Yang et al., 1991, pl. 11, fig. 3; 1993, pl. 11, fig. 3) from the Xijiadian Formation, Xichuan, Henan, but differ in having a shorter palpebral lobe and a narrower palpebral field of the fixigena. Because the holotype cranidium and the paratypes of F. quadratoglabella are strongly deformed or distorted, an extensive comparison is not possible. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone and the lower part of the Lejopyge laevigata Zone).

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Family DORYPYGIDAEKobayshi, 1935 Genus DORYPYGEDames, 1883

Dorypyge Dames, 1883, p. 23, 24; Zittel, 1885, p. 595; Walcott, 1886, p. 221,222; 1889, p. 443; 1905, p. 28; 1913, p. 107; Matthew, 1897, p. 186; Toll, 1899, p. 35; Gr6nwall, 1902, p. 126; Sun, 1924, p. 29; Resser and Endo, 1937, p. 208; Lermontova, 1940, p. 141; Resser, 1942, p. 15-17; Westergg.rd, 1948, p. 7; Egorova, Lomovitskaya, Poletaeva, and Sivov, 1955, p. 123; Hup6, 1955, p. 91; Lu, 1957, p. 264; Poulsen in Moore, 1959, p. O217; Kraskov, Lazarenko, Ogienko, and Chernysheva, 1960, p. 218; Chernysheva, 1960, p. 78; Kobayashi, 1960b, p. 347; Lu, Zhu, and Qian, 1963, p. 62; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 96, 97; KhairuUina, 1970, p. 18; Kushan, 1973, p. 134, 135; Repina, Petunina, and Khairullina, 1975, p. 142; Nan, 1976, p. 333, 334; 1980, p. 486; Zhou, Liu, Meng, and Sun, 1977, p. 129; Schrank, 1977, p. 145; Yang, 1978, p. 31; Zhu, Lin, and Zhang, 1979, p. 85; Zhang, 1981, p. 154, 155; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 56, 57; Han, 1984, p. 15; Zhang and Wang, 1985, p. 339; Zhang and Jell, 1987, p. 56-58; Yang, Yu, Li, Su, He, Shang, Zhang, Zhu, Li, and Yan, 1991, p. 127; 1993, p. 163; Zhu, 1992, p. 338; Guo, Zan, and Luo, 1996, p. 92; Luo, 2001, p. 377. Dorypigaspis Khairullina, 1973, p. 56, 57. Olenoides (Dorypyge) Lorenz, 1906, p. 81. Dorypyge (Jiuquania) Li in Zhang, 1981, p. 155. Type species. Dorypyge richthofeni Dames (1883, p. 24, 25) from the Changhia Formation in Liaoning; by original designation. Other species. Species described prior to 1987 were listed and reassigned by Zhang and Jell (1987, p. 57, 58). To this list, the following species may be added: Dorypyge forta Repina in Repina et al., 1975 (p. 143, 144, pl. 21, figs. 8-12) from the Dignaspis Bed (Mayan Stage), Turkestan; Dorypyge zujiensis Ju in Qiu et al., 1983 (p. 57, pl. 19, figs. 1-3) from the Yangliugang Formation, Zhuji, Zhejiang; Dorypyge yantouensis Zhang and Wang (1985, p. 340, pl. 105, figs. 14, 15) from the Changhia Formation, Fanshi, Shanxi; Dorypyge subovata Zhang (1981, p. 155, 156, pl. 59, fig. 6) from the Mohershan Formation, Kuruktag, Xinjiang, Dorypyge kunshanensis Qian and Zhou (1984, p. 175, 176, pl. 2, figs. 5, 6, ?7) from Kunshan, Jiangsu, and three new species described herein, Dorypyge bisulcata sp. nov., Dorypyge huaqiaqoensis sp. nov. and Dorypyge globosa sp. nov. An Australian species erected by t3pik (1967, p. 174, pl. 2, fig. 8; text-fig. 55) under the name of Olenoides tranans from Queensland, Australia, should be reassigned to Dorypyge. Several species assigned to Dorypyge are possible junior synonyms of D. richthofeni. These include Dorypyge fuzhouwanensis Han, 1984 (p. 15-17, pl. 1, figs. 1-6; text-fig. 1) from shales in the lower part of the Changhia Formation, Fuxian, southern Liaoning; Dorypyge yuhangensis Ju (in Qiu et al., 1983, p. 57, pl. 19, figs. 5, 6) from the Yangliugang Formation, Yuhang, Zhejiang; three species from the Changhia Formation, North China, Dorypyge lata Zhang and Wang (1985, p. 339, pl. 105, fig. 11), Dorypyge zhaogeichuangensis Zhang and Wang(1985, p. 339, pl. 105, fig. 12), and Dorypyge angusta Zhang and Wang (p. 339, 340, pl. 106, fig. 13); and Dorypyge qinlingensis Yang in Yang et al. (1991, p. 127, 128, pl. 10, figs. 1, 2; 1993, p. 163, 164), from the Xijiadian Formation (Middle Cambrian), Xichuan, Henan. Both Dorypyge (Jiuquania) multiformis Li in Zhou et al. (1982, p. 232, pl. 59, figs. 10-13) from the Germogou Formation, Jiuquan, Gansu, and D. (Jiuquania) xinjiangensis Zhang (1981, p. 155, pl. 59, fig. 7) from the Mohershan Formation, • 70.

Kuruktag, Xingjiang, are junior synonyms of D. perconvexalis Yang.

Remarks. The genetic concept of Zhang and Jell (1987, p. 56) is followed here. Complete exoskeletons of Dorypyge fuzhouwanensis and D. richthofeni, the type species, were recently illustrated by Han (1984, pl. 1, figs. 1-3; text-fig. 1) and Guo et aL (1996, pl. 36, fig. 1). The thorax in each species is of seven segments, and has pleurae bearing rather deep and incised pleural furrows and narrow bands. Dorypigaspis Khairullina, 1973 (p. 56, 57) is a possible junior synonym of Dorypyge. Both the genus and the type species, Dorypigaspis bifida Khairullina, are monotypic, based only on a single incomplete, largely exfoliated pygidium from the middle Cambrian (Mayan age) of the Turkestan range, Uzbekistan (Khairullina,1973; pl. 4, figs. 8, 9). The pygidium is indistinguishable from that of Dorypyge. Khairullina (1973) stated that her new genus can be differentiated from Dorypyge because in the latter, the marginal pygidial spines equal the number of axial tings, but in Dorypigaspis, the spines have twice the number of axial tings. Dorypigaspis, as described, has five pairs of marginal spines on the pygidium, with the first four pairs being branched. The fifth spine, the posterolateral, is the longest and the thickest. According to Khairullina (1973) the posterolateral spine is the eighth pair of spines. However, Khairullina apparently had not described the last pair of short spines lying between the posterolateral spines. The number of spines for Dorypigaspis as originally reported seems doubtful, because spines are not shown on the illustrations of the holotype pygidium. Our material from northwestern Hunan suggests the holotype pygidium belongs to Dorypyge, and that the number of spines may have been misinterpreted because of unsatisfactory preservation of the pygidium. In Dorypyge, the first four pairs of pygidial spines are commonly located at the posterolateral corners of the anterior bands (or fibs, if the interpleural furrows are obliterated), immediately anterior to pleural furrows (see P1. 15, figs. 10, 11; P1. 16, figs. 1, 6, 11). It is possible that the anterolateral comers of the ribs were misinterpreted as one of the branches of bifurcated spines for the pygidium of Dorypigaspis bifida (Khairullina, 1973, pl. 4, figs. 8, 9). The holotype pygidium needs to be restudied to clarify morphological details. For comparison, the pygidium of Dorypyge is variable in the number of marginal spines and axial tings. Dorypyge richthofeni has five to six pairs of marginal spines, and four to five axial tings (see Schrank, 1977, pl. 1, fig. 6; pl. 2, figs. 1-4) whereas D. khademi pallilosa (Kushan, 1973, pl. 27, figs. 2, 7, 8) has as many as seven pairs of marginal spines. This variation indicates that differences in spine numbers should not be accorded more than species-level significance in dorypygiids. Khairullina (1973) also differentiated her new genus from Dorypyge in the lack of both interpleural furrows and granulose ornamentation. These differences, however, also are of no more than species-level significance. As the axial and pleural characters of Dorypigaspis are so similar to those of Dorypyge, we tentatively consider Dorypigaspis to be synonymous with Dorypyge, regardless of the true number of marginal spines on the pygidium. The subgenus D. (Jiuquania) Li (in Zhang, 1981, p. 155) is apparently synonymous with Dorypyge. It differs from Dorypyge sensu stricto only in having deeper interpleural furrows on the pygidium, and this should be regarded as a specific rather than subgeneric criterion.

Dorypyge richthofeni Dames, 1883 Plate 14, figures 1-14; Plate 15, figures 1-11; Plate 16, figures 1-7a, 8-11; Plate 19, figure 5 1883 1906 1913 1924

Dorypyge richthofeniDames, p. 24-27, pl. 1, figs. 1-6. Olenoides (Dorypyge) richthofeni Dames; Lorenz, p. 81-87, pl. 4, figs. 1-5. Dorypyge richthofeni Dames; Walcott, p. 108, 109, pl. 8, figs. 1, 1a-f. Dorypyge richthofeni Dames; Sun, p. 29, 30, pl. 2, figs. 3a-d. .71.

1937 1937 1938 1942 1942 1942 1942 1942 1955

Dorypyge richthofeni Dames; Kobayashi, p. 434, pl. 17, figs. 13a, b. Dorypyge damesi Dames; Resser and Endo, p. 209, 210, pl. 31, figs. 14-18. Dorypyge laiwuensis Kobayashi, p. 887, fig. 1. Dorypyge richthofeni Dames; Resser, p. 16. Dorypyge lorenzi Resser, p. 18. Dorypyge suni Resser, p. 19. Dorypyge shantungensis Resser, p. 19. Dorypyge chihliensis Resser, p. 19. Dorypyge richthofeni Dames; Egorova, Lomovitskaya, Poletaeva, and Sivov, p. 123, pl. 13, figs. 11a, b. 1960b Dorypyge richthofeni Dames; Kobayashi, p. 347, 348. 1962 Dorypyge richthofeni Dames; Lu, Zhu, and Qian, p. 30, pl. 5, fig. 10; pl. 6, fig. 6. non 1963 Dorypyge richthofeni Dames; Egorova, Xiang, Li, Nan, and Guo, p. 33, 34, pl. 6, fig. 7 [=Dorypyge perconvexalis Yang in Zhou et al., 1977]. 1965 Dorypyge richthofeni Dames; Lu, Zhang, Zhu, Qian, and Xiang, p. 97, 98, pl. 14, figs. 10, 11. 1965 Dorypyge richthofeni chihliensis Resser; Lu, Zhang, Zhu, Qian, and Xiang, p. 98, 99, pl. 14, figs. 12, 13. 1965 Dorypyge richthofeni damesi Resser; Lu, Zhang, Zhu, Qian, and Xiang, p. 98, 99, pl. 14, figs. 12, 13. 1977 Dorypyge richthofeni Dames; Schrank, p. 145-147, pl. 1, figs. 1-6; pl. 2, figs. 1-5. 1977 Dorypyge richthofeni Dames; Zhou, Liu, Meng, and Sun, p. 129, pl. 42, fig. 8. 1978 Dorypyge cf. D. richthofeni Dames; Yang, p. 77, pl. 4, fig. 2. 1980 Dorypyge richthofeni Dames; Nan, p. 486, pl. 200, figs. 24, 25. 1981 Dorypyge richthofeni damesi Resser and Endo; Zhang, p. 155, pl. 59, fig. 5. 1982 Dorypyge richthofeni Dames; Liu, p. 299, pl. 214, fig. 14. 1985 Dorypyge richthofeni Dames; Zhang and Wang, p. 338, 339, pl. 105, figs. 6-8. 1985 Dorypyge lata Zhang and Wang, p. 339, pl. 105, fig. 11. 1985 Dorypyge zhaogeichuangensis Zhang and Wang, p. 339, pl. 105, fig. 12. 1985 Dorypyge angusta Zhang and Wang, p. 339, 340, pl. 106, fig. 13. 1987 Dorypyge richthofeni Dames; Zhang and Jell, p. 58, pl. 12, figs. 4-7; pl. 13, figs. 1-10; pl. 15, fig. 7. 1987 Dorypyge damesi Dames; Zhang and Jell, p. 59, 60, pl. 14, figs. 13-16. 1992 Dorypyge richthofeni Dames; Zhu, p. 338, pl. 116, fig. 8. 1996 Dorypyge richthofeni Dames; Guo, Zan, and Luo, p. 93, 94, pl. 36, figs. 1-13. 2001 Dorypyge richthofeni Dames; Luo, p. 337, p. 2, figs. 1-15. 2001b Dorypyge richthofeni Dames; Peng, Babcock, and Lin, p. 101, 102, pl. 2, figs. 15, 16, pl. 3, figs. 14, 15. 2001 e Dorypyge richthofeni Dames; Peng, Babcock, Lin, Chen, and Zhu, p. 159, figs. 10.13, 10.14. Lectotype. Cranidium (Dames, 1883, pl. 1, fig. 1; also Kobayashi, 1937, pl. 17, fig. 13a; Schrank, 1977, pl. 1, fig. 1; MB.T 3794) from Dames's (1883) type material from Liaoning; designated by Kobayashi (1937). Material. Numerous sclerites, including cranidia, hypostomes, and pygidia (illustrated specimens NIGP 137425-137456) are in collections P3.2, P7.25, P12.5, P18.2, P35.5, P38.5, P42.5, P45.1, P45.6, P48.5, P64.5, P81, P82.5, P108, P121.48, P112.6, P114(?), P122, P. 123.6, P125, P126, W33, W56.7, W64.7, W77.8, W82.1, and W82.2. • 72-

Remarks. Specimens from the Huaqiao Formation in northwestern Hunan are quite similar to the type material from southern Liaoning, North China (Dames, 1883). The key points of comparison are six pairs of marginal spines on the pygidium, four to five axial tings and a terminal piece on the pygidial axis, and coarse to fine, widely-spaced granules on the cranidium. The species has been redescribed several times by Walcott (1913), Kobayashi (1937), Schrank (1977), and Zhang and Jell (1987). Variability in the characteristic features of the original material includes the surface granulation, the width of the preocular area of the fixigena, the segmentation of the pygidial axis, and the size of the fourth pygidial border spines. Our specimens from northwestern Hunan fall well within the apparent range of morphologic variability in the species. They also add information about ontogenetic variation in the cranidium and pygidium of D. richthofeni. Resser (1942) erected numerous new species using Dames's (1883) original material, although he did not figure any of the specimens. Zhang and Jell (1987) reviewed this material and synonymized all of Resser's species as junior synonyms of D. richthofeni. They also noted that several other species described from North China, including D. damesi, D. laevis, D. kidoi, and D. leei were probably junior synonyms of D. richthofeni. We concur with their assignments and suggestions. Dorypyge laiwuensis Kobayashi, 1938 was erected using some of Lorenz's original material of D. richthofeni (Lorenz, 1906, pl. 4, figs. 1-5; also Kobayashi, 1938); it was regarded by Zhang and Jell (1987, p. 57) as a valid species. However, after examination of Dames's type material at the Naturkund Museum in Berlin, Germany, by one of us (SP), and after examination of photographs of Lorenz's figured and unfigured specimens, provided by G. Geyer, we conclude that Lorenz's original assignment was correct, D. laiwuensis should be regarded as a junior synonym of D. richthofeni. Occurrence. The type material occurs in oolitic limestone and interbedded limestone and shale in the Amphoton Zone of the Changhia Formation, Liaoning, China. The species has a wide distribution through China, where it occurs in middle Cambrian strata. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan. Dorypyge richthofeni is the eponymic species of the D. richthofeni Zone. The species occurs through the entire biozone and into the lower part of the overlying Pianaspis sinensis Zone (equivalent to the Ptychagnostus atavus Zone through the lower part of the Goniagnostus nathorsti Zone). Dorypyge perconvexalis Yang in Zhou et al., 1977 Plate 17, figures 1-14; Plate 19, figures 1-3 1963 1977 1978 1981 1982 1982 2001b 200 lc

Dorypyge richthofeni Dames; Egorova, Xiang, Li, Nan, and Guo, p. 33, 34, pl. 6, fig. 7. Dorypyge perconvexalis Yang in Zhou et al., p. 129, pl. 42, figs. 9, 10. Dorypyge perconvexalis Yang, p. 31, 32, pl. 6, figs. 3, 4. Dorypyge (Jiuquania) xinjiangensis Zhang, p. 155, pl. 59, fig. 7. Dorypyge (Jiuquania) multiformis Li in Zhou et al., p. 232, pl. 59, figs. 10-13. Dorypyge perconvexalis Yang; Liu, p. 299, pl. 213, figs. 2, 6. Dorypyge perconvexalis Yang; Peng, Babcock, and Lin, p. 103, pl. 7, figs. 12, 13. Dorypyge perconvexalis Yang; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 3, fig. 9.

Lectotype. Cranidium (Yang in Zhou et al., 1977, pl. 6, fig. 3; CUGB 0318105) from the Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan, China; designated subsequently as holotype of the species by Yang (1978). • 73.

New material. More than 20 sclerites, including more than 10 cranidia, two librigena, one hypostome, and six pygidia (illustrated specimens INGP 133527, 137457-137466; 137474, 137475) are in collections P249, P261.3, P261.35, P261.5, P273.52, P273.8, P276.2, P277, P282.75(?), P283.47(?), P283.75, P307.4, P331.8(?), and P[3-2.75. Emended diagnosis. Dorypyge with cranidium obtusely angled anteriorly; glabella highly convex, subrectangular in outline, expanded gently forward. Pygidium with wide and relatively short axis; both pleural and interpleural furrows incised; with five pairs of marginal spines developed as outgrowths of the pleurae; last pair of marginal spines long and stout; lateral borders absent. Remarks. The combination of the presence of deep and clearly defined pleural and interpleural furrows, and long and stout posterolateral spine on the pygidium, characterize this species. Other distinctive features include the angled anterior cranidial margin, the highly convex glabella, and the relatively narrow pleural field. Lateral borders and border furrows are absent on the pygidium, so in this species, the marginal spines are developed as extensions of the pleurae. The type material of D. perconvexalis from the Huaqiao Formation of northwestern Hunan is unsatisfactorily preserved and illustrated (Yang in Zhou et al., 1977; also in Yang, 1978). New material from the same region helps to clarify the species concept. It shows that two species referred to the subgenus Dorypyge (Jiuquania) by Zhang (1981), D. (Jiuquania) xinjiangensis and D. (Jiuquania) multiformis, both from Northwest China, are identical in all respects with D. perconvexatis. They should be suppressed as junior synonyms of D. perconvexalis. As discussed previously, the subgenus Dorypyge (Jiuquania) should be suppressed as a junior synonym of Dorypyge. The well furrowed pygidium of D. perconvexalis is most reminiscent of that of D. khademi papillosa Kushan (1973, p. 141-144, pl. 27, figs. 5-11, text-fig. 11) from the Mila Formation of the Alborz Mountains, Iran, which also possesses incised interpleural furrows, and a similarly shaped axis with four tings and a terminal piece. D. khademi papillosa, however, has two more pairs of tiny border spines on the posterior margin between the large posterolateral spines of the pygidium. The Iranian subspecies also differs from D. perconvexalis in having a cranidium with a nearly parallel-sided, less convex glabella that bears distinct lateral furrows, a rounded rather than angled anterior margin, posteriorly located palpebral lobes, and a wide (tr.) anterior area with a well impressed anterior border furrow. D. perconvexalis is also similar closely to Olenoides tranans Opik (1967, p. 174, pl. 2, fig. 8; text-fig. 55) from the Erediaspis eretes Zone in the Mungerebar Limestone, Queensland, Australia. O. tranans should be considered a Dorypyge species because its pygidium bears a long and stout posterolateral spine. No such spine is known in Olenoides, but it is one of the diagnostic characters of Dorypyge. The Australian species, however, can be differentiated from D. perconvexalis by having a slender, less tapered axis, shallow instead of incised furrows on the pleural region; and by lacking granulation. D. perconvexalis is the youngest known species of Dorypyge. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Lejopyge laevigata Zone through the Glyptagnostus stolidotus Zone).

• 74-

Dorypyge bisulcata sp. nov. Plate 18, figures 1-6 2001b Dorypygebispinosa Walcott; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 5.

Etymology. From Latin, prefix bi, two, plus sulcatus, furrowed, referring to the distinctive interpleural and pleural furrows on pygidium. Holotype. A pygidium (P1. 18, fig. 3, NIGP 137468) in collection P167. Other material. Ten sclerites including cranidia and pygidia in collections P130.95, P156, P164.2, P174, and W111.3, among which one cranidium, one pygidium, and two fragments are figured as paratypes (NIGP 137467, 137469-137471). Diagnosis. Dorypyge with glabella expanding strongly forward on posterior two-thirds, and semicircular on anterior one-third; pygidium with wide pleural field beating incised interpleural and pleural furrows, and five border spines with posterior two spines being much longer and thicker than others, posterior two spines diverging widely outward. Description. Glabella moderately convex, expanded strongly forward on posterior two-thirds, semicircular in outline in anterior one-third; S1 short, slightly impressed. Occipital furrow broad and shallow, deep at sides. Axial furrow well defined with antennal pit besides anterolateral comer of glabella. Anterior border ridge-like and clearly defined medially, becoming wide and poorly defined by broad and shallow border furrow laterally. Small palpebral lobe located at the level of glabellar midlength; eye ridge oriented diagonally, clearly defined in front of palpebral lobe, merging with fixigena adaxially. Anterior branch of facial suture slightly convergent forward, enclosing steeply downsloping preocular area; posterior branch sigmoidal, enclosing a subtriangular posterolateral projection. Pygidium semicircular in outline, slightly longer than wide. Axis long, cylindrical, with four tings and large, subrectangular terminal piece; terminal piece obtusely rounded posteriorly; first three ring furrows relatively deep, last ring furrow weakly defined. Pleural field wider than axis, well elevated above border; with four pleural segments with wide and deep pleural furrows and thin but clearly defined interpleural furrows; anterior band inflated, widening abaxially. Borders relatively narrow, separated from pleural field by faint border furrows and change of slope; five pairs of border spines almost evenly spaced: the first three pairs long and thin, becoming progressively shorter rearward, nearly evenly spaced; posterior two pairs much longer and stronger, diverging widely outward. Ornamentation consisting of dense, medium-sized granules on glabella, closely spaced, medium-sized, scattered fine granules on fixigenae and on pygidial surface. Remarks. One pygidium assigned here as Dorypyge bisulcata sp. nov. was previously referred to D. bispinosa (Peng et al., 2001b) because it bears two pairs of long posterolateral spines that are almost identical with those of D. bispinosa. More careful comparison reveals that those specimens represent a new species of Dorypyge. So far only one pygidium is known for D. spinosa. It was from a locality southwest of Yanzhuang, Xintai district, Shandong, North China, and was illustrated -75.

several times (Walcott, 1913, p. 107, pl. 8, fig. 3; Lu et al., 1965, pl. 15, fig. 1; Zhang and Jell, 1987, pl. 14, fig. 12). The pygidium of the new species differs from the holotype pygidium of D. bispinosa in having a semicircular rather than subtrapezoidal outline, a relatively longer and slender, more segmented axis, a pleural field that is much wider than the axis, and a more widely spaced posterior pair of spines. In D. bispinosa, the first pleura is not transverse but backward-curved distally, the interpleural and pleural furrows are poorly rather than clearly defined, the anterior bands of pleurae are not inflated as in the new species, and the posterior bands are ridgelike and much thinner than those of the new species. Although the pygidium of D. bispinosa has a somewhat larger size than the pygidium of the new species, it has less segmentation than the new species. This indicates that they belong to different species. In other dorypygids (e.g., D. richthofeni), large pygidia commonly bear more segments. Besides D. bisulcata, the only other Dorypyge species that bears deeply defined pleural and interpleural furrows is D. perconvexalis Yang in Zhou et al. (1977). The new species differs from D. perconvexalis in having two pairs of long and stout posterolateral spines on the pygidium, rather than one pair of spines, and a rounded rather than angular anterior margin on the cranidium. Furthermore, the lateral spines in D. perconvexalis are derived from pleurae rather than from the border. The new species is similar to D. subovata Zhang (1981, p. 155, 156, pl. 59, fig. 6) from the Mohershan Formation, Kuruktag, Xinjiang. D. subovata is based on a single, mostly exfoliated cranidium and differs only in having a glabella that is more convex and more elliptical in outline, and probably in having more deeply impressed fossulae on the axial furrows. Because of the close similarity in the cranidium, the new species eventually may prove to be synonymous with D. subovata, but until the pygidium and more cranidial features (such as surface ornamentation) are known for the species from Xinjiang, we preferred to assign the Hunan specimens to a separate, new species. The holotype pygidium of Olenoides tranans Opik (1967, p. 174, 175, pl. 2, fig. 8) from Queensland, Australia, which properly belongs to Dorypyge, shows some similarity to the pygidium of the new species. The Australian specimens, however, have less well defined pleural and interpleural furrows, less well defined borders, and a relatively longer but narrower outline.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone and the Lejopyge laevigata Zone). Dorypyge huaqiaoensis sp. nov. Plate 19, figures 6, 7

Etymology. From Chinese, Huaqiao, in reference to Huaqiao Formation, from which the type material was collected. Huaqiao is a village in Baojing County, northwestern Hunan, after which the well-known Huaqiao Formation is named. Holotype. Cranidium (P1. 19, fig. 7; NIGP 137479) in collection P81. Other material. One cranidium and one pygidium (NIGP 137478a, b) in collection P81. Diagnosis. Dorypyge with cranidial and pygidial surfaces lacking granulose ornamentation" Glabella wide and relatively short, parallel-sided, truncate anteriorly, with anterolateral comers • 76.

deeply notched inward by prominent fossulae. Pygidium with a thick and short axis with three tings and long terminal piece.

Description. Cranidium with narrow, convex anterior border being broadly arched forward. Glabella less than two times as long as wide, nearly parallel-sided, obtusely rounded anteriorly with anterolateral comers markedly notched by fossulae that are deeply impressed. Occipital furrow shallow mesially, deepening abaxially; occipital ring wide (sag.), occupying about one-fifth total glabellar length. Pygidium with an axis that has four tings and a long terminal piece. Pleural furrow faintly impressed, interpleural furrow effaced. No surface granulation on both cranidium and pygidium.

Remarks. This species is represented by a few incomplete sclerites. The non-granulation surface and the parallel-sided, anteriorly truncate glabella warrant an assignment of new species of Dorypyge. D. laevis (Walcott, 1913, p. 109, figs. 2, 2', 2a; also Zhang and Jell, 1987, pl. 13, figs. 11-13; pl. 15, fig. 13) from the Changhia Formation near Dongyu, southern Shanxi has a nearly smooth surface, but its glabella is distinctly rounded at anterolateral comer and seems not truncate but very rounded anteriorly. The incompleteness of both species prevents a further comparison. In cranidial morphology, the new species resemble mostly D. pergranosa Resser and Endo (1937, p. 210, pl. 31, figs. 6-13; also Zhang and Jell, 1987, p. 59, pl. 14, figs. 1-11; pl. 15, figs. 1-5, 8, 9, 12) from the Changhia Formation, Changxingdao Island, southern Liaoning, but differs in having a glabella that is less rounded anteriorly, and in lacking granules on surface.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Dorypyge globosa sp. nov. Plate 18, figures 7-9 2001b Dorypygesp., Peng, Babcock, and Lin, p. 102, pl. 5, fig. 4.

Etymology. From Latin, globosus, globose, referring to the distinctive shape of the glabella. Holotype. Cranidium (P1. 18, figs. 7, 8; NIGP 137472) in collection P156. Other material. A pygidium (NIGP 137473), also in collection P156. Diagnosis. Dorypyge with rounded, highly convex glabella, and broad occipital furrow; pygidium with interpleural furrows effaced; border clearly defined, with five pairs of spines, fifth pair weak, directed posteriorly.

Description. Glabella convex, subspherical in outline, slightly longer than wide, lateral furrows effaced. Axial furrows deeply impressed anteriorly. Occipital furrow shallow; occipital ring as wide as occipital furrow sagittally, narrowing abaxially, with a large median spine or node. Fixigena • 77.

moderately convex, anterior area sloping strongly downward, beating broad and shallow border furrow. Palpebral lobe unknown, but probably located at about midlength of cranidium. Partially preserved eye ridge weakly defined, merging with fixigenae proximally. Posterior border furrow broad and deep, posterior border evenly wide (exs.). Surface with dense granules on glabella and occipital ring, and scattered granules on fixigenae. Pygidium semicircular in outline with a long, cylindrical axis extending to border furrow. Pleural field moderately convex; interpleural furrows effaced, bearing shallow and wide pleural furrows; border narrow, bearing five pairs of marginal spines, with first three pairs short and slender; fourth pair of spines strong, long; fifth pair of spines long, strong, directed posteriorly. Surface with fine, dense granules.

Remarks. This species is represented by a single incomplete but distinguishable cranidium characterized by having a strongly convex, globose glabella and a broad occipital furrow. The glabella is reminiscent of Dorypyge subovata Zhang (1981, pl. 59, fig. 6) from Kuruktag, Xinjiang, but the granulation in D. subovata is finer and less densely spaced, the occipital and the posterior border furrows in D. subovata are much narrower and shallower than they are in D. globosa. The pygidium referred to D. globosa is distinctive in having a broad pleural field, narrow borders, in lacking interpleural furrows, and in having a relatively weak fifth border spine. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Dorypyge? sp. Plate 19, figure 4

Material. One pygidium, NIGP 137476, from P 290.5. Remarks. A single pygidium is tentatively placed in the genus Dorypyge as it bears a postaxial ridge that is not observed in any species of Dorypyge. The pygidium resembles some other pygidia of Dorypyge in having five pairs of marginal spines, six axial tings and a short terminal piece, weak interpleural furrows, strong pleural furrows, and coarse granules. A pygidium referred to Dorypyge dames (Resser and Endo, 1937, pl. 31, figs. 17, 18; also Zhang and Jell, 1987, pl. 14, figs. 15, 16) is most comparable in having an elongate subtriangular outline with a narrow pleural field. However, the pygidium of D. dames has narrower borders and thinner border spines. Two pygidia assigned as Damesellidae genus and species undetermined B from the Mila Formation, Alborz Montains, northern Iran (Wittke, 1984, pl. 11, figs. 11, 12) may be congeneric or even conspecific with the new pygidium. The two pygidia compare well with the new specimen, differing only in having a wider outline and a proportionally wider and shorter axis. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone).

• 78"

Suborder LEIOSTEGILNABradley, 1925 Superfamily LEIOSTEGIOIDEABradley, 1925 Family LEIOSTEGIIDAEBradley, 1925 Subfamily LEIOSTEGIINAEBradley, 1925 Genus CHUANGIAWalcott, 1911

Type species. Ptychoparia? batia Walcott (1905, p. 75) from the Changshan Formation, Shandong, China; by original designation.

Remarks. The genetic concept of Zhang and Jell (1987, p. 199) is followed here. Chuangia subquadrangulata Sun, 1935 Plate 20, figures 15, 16 1935 Chuangia subquadrangulata Sun, p. 24, pl. 1, fig. 12. 1965 Chuangia subquadrangulata Sun; Lu, Zhang, Zhu, Qian, and Xiang, p. 369, pl. 68, figs. 17, 18. 1980 Chuangia subquadrangulata Sun; Nan, p. 502, pl. 205, fig. 11. non 1983 Chuangia subquadrangulata Sun; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, p. 171, pl. 56, fig. 11 [= Chuangia batia Walcott]. 1985 Chuangia subquadrangulata Sun; Zhang and Wang, p. 453, pl. 134, figs. 5, 6. 1994 Chuangia (Chuangia) subquadrangulata Sun; Qian, p. 107, 108, pl. 24, figs. 3-5, ?6.

Holotype. By monotypy; cranidium (Sun, 1935, pl. 1, fig. 12) from the Changshan Formation at Dawenkou, Shandong, China.

New material. One cranidium and one pygidium (NIGP 137491, 197492) and fragments are in collections P370.6, P 374.9, P375.15, and P383.5.

Remarks. New material from the Huaqiao Formation, northwestern Hunan, is similar to the type material from Shandong, North China, in most respects including the possession of a subrectangular glabella with three pairs of short, shallow lateral glabellar furrows. However, the glabella in the new material is not straight-sided like that in the type material; instead, it is somewhat constricted in the center. Two pygidia from the Chuangia Zone, Changshan Formation at Qingshuihe, Inner Mongolia, were referred or questionably referred to this species (Qian, 1994). These pygidia are different in having an axis that is shorter and thicker, and more tapered, than that of the pygidium in the new material from Hunan.

Occurrence. The holotype is from the Changshan Formation at Dawenkou, Shandong, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is the eponymic species of the Chuangia subquadrangulata Zone (equivalent to the upper • 79.

Glyptagnostus stolidotus Zone and the Glyptagnostus reticulatus Zone). Chuangia austriaca Yang in Zhou et al., 1977

Plate 20, figures 1-14 1977 1978 1982 2001b

Chuangia austriaca Yang in Zhou et al., p. 191, pl. 56, figs. 8, 9. Chuangia austriaca Yang, p. 55, pl. 10, figs. I-5. Chuangia austriaca Yang; Liu, p. 317, 318, pl. 200, figs. 20, 23. Chuangia austriaca Yang; Peng, Babcock, and Lin, p. 106, pl. 16, figs. 4-6.

Lectotype. Partly exfoliated cranidium (Zhou et al., 1977, pl. 56, fig. 8; refigured by Yang, 1978, pl. 10, fig. 1; CUGB 0242002) from the Chuangia-Prochuangia Zone in the middle part of the Huaqiao Formation at Tingziguan, Fenghuang, northwestern Hunan; designated subsequently as holotype of the species by Yang (1978). New material. More than 20 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 137481-137490) are in collections PI34.3, P~22.4, PI323.5, and P1327.2. Emended diagnosis. Chuangia with anterior border flexed into sharply angled ridge and arched moderately forward; fixigena relatively narrow, with palpebral area sloping strongly towards axial furrow; librigena with genal spine extending from position in front of arterolateral comer; anterior branch of facial suture forwardly convergent, pygidium with moderately convex pleural field, weakly defined border furrow, and rather wide border. Description. Anterior border arched gently forward and flexed into sharply angled ridge, with anterior slope inclined forward and posterior slope inclined rearward. Glabella moderately convex, tapering forward onto border furrow, obtusely rounded to truncate anteriorly, with three pairs of faint lateral furrows; S1 bifurcated; occipital ring crescentic in outline with small, medial node. Palpebral area of fixigena about one-third length of glabella, gently convex and sloping strongly inward; palpebral lobe relatively small, with midpoint opposite posterior one-third of glabellar length; eye ridge weak, oriented diagonally. Anterior branch of facial suture converging forward gently to anterior border furrow, curving inward to anterior cranidial margin; posterior branch strongly divergent rearward, turning sharply rearward and slightly outward after crossing posterior cranidial border furrow. Librigena with broad genal field, gently upturned lateral border defined by broad and shallow border furrow, and genal spine extending from position in front of anterolateral comer. Pygidium subtriangular in outline, width twice length. Axis long and narrow, tapering slightly rearward with five tings defined by shallow or obscure ring furrows and semicircular terminal piece, beyond posterior border furrow, thick postaxial ridge extending to posterior margin. Each axial ring with a pair of widely spaced nodes. Pleural field gently convex, faintly segmented with 2 or 3 fibs. Lateral and posterior borders moderately wide and of uniform width, slightly convex, defined by shallow border furrows. Ornamentation consists of dense punctae. Remarks. The lectotype cranidium of Chuangia austriaca is partly exfoliated and its occipital ring is partly broken, but paratype cranidia, although exfoliated, have complete occipital tings. The • 80.

syntype pygidia are mostly testaceous (Yang, 1978, pl. 10, figs. 2-5). These specimens show both testaceous and exfoliated features, providing a base for a better understanding of the species concept. The new material from the Huaqiao Formation at Paibi is from the same formation and the same geographic region as the type material. It agrees in all observed details with the type material and is confidently assigned to this species. C. austriaca is most similar to C. tolli Resser and Endo (1937, pl. 54, figs. 16-25) from the Chuangia Zone at Jinjiachengzi, southern Liaoning. The type material of C. tolli was refigured by Zhang and Jell (1987, pl. 93, figs. 6-10; pl. 94, figs. 1-3). Except for the glabella that is more tapered and slightly constricted in the anterior one-third in C. tolli, it is almost indistinguishable from C. austriaca in cranidial features. In the pygidium, C. tolli differs in having a more rounded outline, a thicker and shorter, less segmented axis, and in lacking borders. C. austriaca is also comparable with C. wulingensis Yang in Zhou et al., 1977 (pl. 56, figs. 11, 12). These two species are from the same area, but C. wulingensis was also reported from Taoyuan, northwestern Hunan (Peng, 1992, figs. 42I-K, M-O). C. wulingensis has a thinner and more effaced exoskeleton than does C. austriaca. Cranidial furrows of C. wulingensis are much shallower than are those of C. austriaca, and the eye ridges are less defined. C. wulingensis is also distinguished by having a thinner anterior border, and a narrower fixigena.

Occurrence. The holotype is from the Chuangia-Prochuangia Zone in the middle part of the Huaqiao Formation at Tingziguan, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs with trilobites indicative of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). Genus MEROPALLA13pik, 1967

Meropalla Opik, 1967, p. 269; Yuan and Yin, 1998, p. 138. Type species. Meropalla quadrans Opik, 1967 (p. 269, 270, pl. 1; pl. 18, fig. 7; pl. 19, figs. 1-3) from the Erediaspis eretes Zone and the Acmarharchis (=Cyclagnostus) quasivespa Zone, Queensland, Australia; by original designation. Other species. Meropalla auriculata Opik, 1967 (p. 271, 272, pl. 1; pl. 19, figs. 4, 5) from the Erediaspis eretes and the Acmarharchis [=Cyclagnostus] quasivespa zones, Queensland, Australia; Meropalla bella Yuan and Yin (1998, p. 138-140, pl. 1, figs. 1-6) from the Hadragnostus modestus [=Formosagnostus formosus]-Blackwelderia Zone, Huaqiao Formation at Jimachong, Wanshan, eastern Guizhou. Remarks. The genetic concept of Opik (1967, p. 269) is followed here. Opik placed Meropalla in the family Leiostegiidae but left its subfamilial status uncertain. We consider this genus to be referable to the subfamily Leiostegiinae because the general morphology of the cranidium and pygidium are close to such leiostegiinean genera as Leiostegium, Paraszechuanella, and Chuangia. The anterior border shares a flexed appearance with that of Chuangia, and the small posteriorly located palpebral lobe is similar to that of Leiostegium.

.81.

Meropalla bella Yuan and Yin, 1998 Plate 21, figures 1-11 1998 Meropallabella Yuan and Yin, p. 138-140, pl. 1, figs. 1-6.

Holotype. Cranidium (Yuan and Yin, 1998, pl. 1, fig. 1, NIGP 127884) from the Hadragnostus modestus [=Formosagnostus formosus]-Blackwelderia Zone in the middle part of the Huaqiao Formation at Jimachong, Yuping, eastern Guizhou.

New material. Two cranidia and two pygidia (NIGP 137493-137496) are in collections W225, W228.3, and P298.4.

Emended diagnosis. Meropalla with a parallel-sided, anteriorly truncated glabella; strongly depressed triangular area on inner palpebral area and postocular field of fixigena; and moderately wide (tr.) anterior border.

Remarks. New material from the Huaqiao Formation near Wangcun, northwestern Hunan, is identical in all respects with the type material from the same region. Yuan and Yin (1998) reported that the lateral furrows of the glabella are completely effaced on testaceous surfaces in the type material. In the new material, three pairs of lateral furrows are observed. They are weakly impressed, with the S 1 being bifurcated and the $2 and $3 being forwardly and inwardly directed (P1. 21, figs. 6-9). The key features that characterize the species include the strongly convex (tr. and sag.), quadratic-shaped glabella, the small, posteriorly placed palpebral lobes, the deep axial furrows, the posteriorly placed occipital node, and the wide (tr.) anterior border that occupies more than half of the cranidial width, with its distal ends opposite the midpoint of the fixigena. This species is readily differentiated from Meropalla quadrans Opik, the type species, in having a parallel-sided instead tapered glabella, a more strongly elevated palpebral area, and a more distinctly depressed triangular area on the posterior part of fixigena, which is more strongly inclining inward.

Occurrence. The holotype is from the Hadragnostus modestus-Blackwelderia Zone in the middle part of the Huaqiao Formation at Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone).

Meropalla gibbera sp. nov. Plate 22, figures 1-6 2001b Meropallasp., Peng, Babcock, and Lin, p. 104, pl. 11, figs. 1, 2.

Etymology. From Latin gibber, hunch-backed or hump-backed, referring to the extremely convex, hunchback-like glabella. • 82.

Holotype. Cranidium (P1.22, figs. 1-6, NIGP 137497) in collection W212.45. Diagnosis. Meropalla with strongly convex, subrectangular or elongate-elliptical glabella that is strongly convex in posterior half, obtusely rounded anteriorly; anterior border short (tr.); deeply depressed and steeply inclined triangular area on posterior part of fixigena.

Remarks. The new species is represented by a single cranidium. It is most similar to Meropalla bella Yuan and Yin, which occurs in the same section with the new species but has a slightly higher stratigraphic occurrence. Morphologically, M. bella has a proportionally shorter but wider, and anteriorly truncate glabella that is less convex in the posterior part. In M. bella, the anterior branches of the facial sutures are less convergent forward, and the anterior border is much wider (tr.) than that of M. gibbera.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the Linguagnostus reconditus Zone). Subfamily

CHELIDONOCEPHALINAE Wittke,

1984

Remarks. Wittke (1984, p. 111, 112) erected the subfamily Chelidonocephalinae and classified questionably it within the Ordosiidae. Subsequently, Peng and Geyer (1999, p. 30, 31) discussed the subfamily at great length. In emending the subfamilial concept, they included Adelogonus Opik, 1967; Chelidonocephalus King, 1937; Derikaspis Dean, 1982; Iranoleesia King, 1937; and Iranochresterius Peng and Geyer, 1999 in the subfamily. The concept of Peng and Geyer (1999) is followed here, except for transferring Adelogonus into Proashaphiscidae. The new genus Geminiclavula, described below, is tentatively referred to the family. Genus GEMINICLAVULAgen. nov.

Etymology. From Latin geminus, double, combined with clavula, small bar, referring to the paired bar-like swellings on the frontal area of cranidium.

Type species. Geminiclavula wangcunica gen. et sp. nov. from the Huaqiao Formation, near Wangcun, Yongshun, northwestern Hunan.

Diagnosis. Chelidonocephalid? genus characterized by a pair of transverse swellings on preocular areas of fixigena; anterior border furrow comprising transverse medial portion and paired gently forward-convex outer portions; anterior border wide (sag.) and upturned; glabella mostly effaced, front truncate; occipital ring large, with tiny, posteriorly located occipital node; palpebral lobes large and thick, located posteriorly; anterior branch of facial suture strongly divergent; posterior branch of facial suture enclosing narrow (exs.), transverse posterolateral projection.

Remarks. Only a single cranidium is available for the type species G. wangcunica, but the distinct morphology makes it clear that new generic and specific names are warranted to embrace the specimen. The cranidium of the new genus resembles in part the cranidia of Wanshania and Baojingia, but differs in having larger, thicker, and more closely spaced palpebral lobes, having a •

83.

proportionally shorter glabella, having a longer (sag.) occipital ring, and having more divergent anterior branches of the facial sutures. The transverse swellings on the preocular areas of the fixigenae are not present in either Wanshania or Baojingia. In fact, this structure has not been reported previously from any liostegiid. The relatively wide anterior area, divergent anterior branches of the facial suture and narrow (tr.) palpebral areas of the fixigenae are reminiscent of pterocephaliids. However, pterocephaliids are readily distinguished by lacking transverse swellings on the anterior area; by facial sutures that are more smoothly curved on the anterior branches; and having smaller, more anteriorly located palpebral lobes.

Geminiclavula wangcunica gen. et sp. nov. Plate 22, figures 8, 9 2001e Leiostegiid gen. et sp. nov., Peng, Babcock, Lin, Chen, and Zhu, p. 159, fig. 10.25.

Etymology. Named for the town of Wangcun, Hunan Province. Holotype. Testaceous cranidium (P1.22, figs. 8, 9, NIGP 137499) in collection W56.7. Diagnosis. Same as for genus. Description. Description based only on holotype. Cranidium length equal to width, length of holotype about 4.25 mm. Anterior border wide (sag., tr.), upturned gently anteriorly, defined posteriorly by sinuous anterior border furrow consisting of transverse, shallow middle portion and with outer end deflected forward as deep, posterolaterally curving furrow that is convex forward and defined anteriorly by pair of bar-like swellings on preocular areas of fixigenae. Glabella subrectangular, tapering forward gently, truncate anteriorly, with only S1 visible as obscure impression. Occipital furrow weakly impressed, ill-defined; occipital ring of uniform width, slightly wider (tr.) than basal width of glabella, beating tiny, posteriorly located node. Palpebral lobe long (exs.) and thick, occupying three-fourths glabellar length, with posterior end close to dorsal furrow and opposite occipital furrow; eye ridge of low convexity, as thick as palpebral lobe, directed diagonally and poorly defined. Palpebral area of fixigena narrow, obliquely directed, of low convexity in front. Anterior branch of facial suture nearly straight, diverging forward at angle of about 110°; posterior branch transverse, enclosing short and narrow (exs.) posterolateral projection.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus punctuosus Zone). Subfamily CHIAWANGELLINAEChu, 1959

Remarks. This subfamily is monogeneric. Zhu (1959, p. 185) erected this subfamily and classified it in the Damesellidae. The assignment has been followed by most authors (Kobayashi, 1960b; Lu et aL, 1965; Qiu et al., 1983; Zhang and Jell, 1987). However, Zhu's assignment was based on two damesellid cranidia that she mistakenly associated with the pygidia of Chiawangella pustulosa Zhu. C. pustulosa was designated as the type species of Chiawangella but, according to Zhang and Jell (1987, p. 217), is in fact a junior synonym of Chiawangella pacifica (Walcott). The concept of • 84.

Chiawangella is based on pygidial characters because both C. pacifica and C. pustulosa were erected with pygidia as holotypes. These pygidia, characterized by having an inverted trapezoidal outline, narrow pleural fields, and a pair of macropleural spines on the first segment, show similarities in morphology to both damesellids and leiostegiids. Based on the new cranidial material of Chiawangella hunanensis sp. nov., which shows such leiostegiid characters as an effaced glabella, this subfamily is reassigned to the Leositegiidae. Damesellid normally have more strongly furrowed glabellas. Pygidial character of Chiawangella also support such a reassignment. So far as known, pygidia of leiostediids lack spines or bear only one pair of large anterolateral spines derived from the posterior band of the first pleural segment and the anterior band of the succeeding pleura. The pygidium of Chiawangella, which bears only two pairs of spines and is similar to leiostegiids in the structure of the anterolateral spine, may be more comparable with leiostediids than damesellids because the pygidium in damesellids is commonly multispinose. Also, the anterolateral spines in the pygidia of damesellids are derived from the first pleural segment (i. e., from the anterior band, or the anterior border, and the posterior band of the first segment). The cranidium of C. hunanensis sp. nov. has a forwardly deflected posterior border that bears two intergenal spines. The spines on the pygidium and on the cranidium serve as the principal means of characterizing the subfamily Chiawangellinae. Genus CHIAWANGELLA Chu, 1959

Chiawangella Chu (Zhu), 1959, p. 71; Kobayashi, 1960b, p. 353, 354; Lu, Qian, and Zhu, 1963, p. 111; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 402; Nan, 1980, p. 505; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 185; Zhang and Jell, 1987, p. 217.

Type species. Chiawangella pustulosa Chu, 1959 (Zhu, p. 71, 72, pl. 5, figs. 6, 7; non figs. 4, 5) from the Drepanura premesnili Zone, Kushan Formation at Jiawang, northern Jiangsu, and on Changxingdao Island, southern Liaoning; by original designation. C. pustulosa is a junior synonym of Albertella pacifica Walcott, 1911 (p. 79, pl. 14, fig. 6). Other species. Chiawangella hunanensis sp. nov. described herein. Tricrepicephalus asiaticus Yuan and Yin (1998, p. 144, pl. 6, fig. 5) from the Hadragnostus modestus [=Formosagnostus formosus]-Blackwelderia Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou may also belong to Chiawangella. Chiawangella karakolica Goncharova in Goncharova et al. (1972, p. 223, 224, pl. 53, fig. 19), from the lower upper Cambrian of Kirghizia (Kyrgyzstan), is based on a single damesellid cranidium belonging not to Chiawangella but probably to Protaizehoia Yang in Yin and Li, 1978. For this reason C. karakolica is here excluded from Chiawangella.

Emended diagnosis. Cranidium with posterior border deflected forward medially, with double intergenal spines; glabella trapezoidal, weakly furrowed, with truncate front and straight flanks; palpebral lobe located anteriorly, moderately elevated above the anterior half of glabella, pygidium inverted trapezoidal in outline, with narrow pleurae, large axis, broad (sag.) posterior border and two pairs of marginal spines, with the anterolateral spine stout, long, and the posterior spine slender, directed rearward.

Remarks. The concept of the genus is emended here, based largely on new material from the • 85-

Huaqiao Formation of northwestern Hunan. Zhu (1959) stated that the genus has three pairs of marginal spines on the pygidium, but new, well preserved material indicates that Chiawangella bears only two pairs of marginal spines. Examination on Zhu's (1959) material at the Museum of the Nanjing Institute of Geology and Palaeontology confirms that pygidia in the type suite have only two pairs of spines. The type species, Albertella pacifica Walcott, was based on a single pygidium from Changxingdao Island, southern Liaoning. The pygidium is distinctively spinose and quite reminiscent of that of the zacanthoidid genus Albertella. However, as noted by Kobayashi (1935), Resser and Endo (1937), and Zhu (1959), it is not closely related to Albertella. Zhu (1959) erected Chiawangella based on two A. pacifica-like pygidia from the Kushan Formation in the Jiawang Coal Field, northern Jiangsu. She designated one of the pygidia as the holotype of Chiawangella pustulosa, the type species of the genus, and transferred A. pacifica to Chiawangella. Zhang and Jell (1987) suppressed C. pustulosa as a junior synonym of C. pacifica with reference only to the pygidium. Cranidia originally assigned by Zhu (1959) to C. pustulosa are indistinguishable from those of Taihangshania, a damesellid genus erected by Zhang and Wang (1985) on the basis of cranidia from central Shanxi. Taihangshania is characterized by having a bacculae-bearing glabella that tapers more rapidly in the anterior half than in the posterior half, and by having a posteriorly located palpebral lobe. Examination of Zhu's (1959) material at the Museum of the Nanjing Institute of Geology and Palaeontology reveals that cranidia that Zhu (1959) assigned to C. pustulosa agree well Taihangshania in all essential respects and belong to that genus. These cranidia also may be conspecific with the pygidia described as Drepanura transversa (Zhu, 1959, pl. 3, figs. 2, 3). The reason for this tentative reassignment is that Zhu's (1959) pygidia of D. transversa are closely comparable in morphology to pygidia assigned to Taihangshania by Zhang and Wang (1985, pl. 136, figs. 13-16). As described below, Taihangshania also occurs in the Huaqiao Formation of northwestern Hunan (P1. 37, figs. 7-15; P1. 38, figs. 1-6 herein). New material of Taihangshania supports the assignment made by Zhang and Wang (1985) for Taihangshania. A leiostegiid cranidium from the Huaqiao Formation, northwestern Hunan, occurs in association with pygidia of Chiawangella hunanensis sp. nov., and is interpreted as belonging to the same species because the spines on both the cranidium and pygidium are consistent with that assignment.

Chiawangella hunanensis sp. nov. Plate 23, figures 1-14; Text-figure 9 2001 c Chiawangella sp., Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 3, figs. 5-7. 2001b Chiawangella sp. cf. C. pacifica (Walcott); Peng, Babcock, and Lin, p. 105, pl. 14, figs. 1-4.

Etymology. Named for Hunan Province. Holotype. Testaceous cranidium (P1.23, figs. 1-5; NIGP 137500) in collection W227. Other material. Five pygidia (NIGP 133524, 133525, 137501-137503) in collections W227, P317.4, and PI3-2.75. Diagnosis. Chiawangella with trapezoidal, anteriorly truncated glabella; preglabellar field absent; palpebral lobes located anteriorly; posterior border deflected forward medially and distally, bearing two intergenal spines. Pygidium longer than wide, with long anterolateral spine extending from •

86.

pleura, broad at base; posterior border wide (sag.); posterior border spines on pygidium relatively closely spaced; posterior margin arched forward.

•

.

. . . .

Text-figure 9. Reconstruction of cranidium and pygidium of Chiawangella hunanensis sp. nov. Cranidium based on specimen NIGP 137500 (see P1.23, figs. 1-5); pygidium based on specimens NIGP 137501, 137502, 133524, 133525 (see P1.23, figs. 6-13). Description. Cranidium length four-fifths width, convex, with two pairs of intergenal spines. Anterior border narrow, gently upturned; preglabellar field absent. Glabella moderately convex (sag. tr.), tapering forward evenly, width at basal lobe about twice that at frontal margin, with nearly straight sides, truncated anteriorly; with three pairs of weak lateral glabellar furrows; S1 long, slightly curving inward and rearward; $2 faint, slightly shorter than and nearly parallel to S1; $3 slightly shorter than $2, obscure. Occipital furrow deep, transverse, curving slightly forward • 87.

distally; occipital ring gently convex (sag.), of nearly uniform width, with weak medial node. Palpebral lobe short (exs.), elevated slightly above glabella, located in anterior one-fifth of cranidium; eye ridge short, thick, almost transverse, obscure; preocular area of fixigena small, inwardly and forwardly inclined; palpebral area narrow (tr., exs.), steeply inclined inward; postocular area subtriangular in outline, moderately convex, defined posteriorly by deep, slightly curved, almost diagonally directed posterior border furrow. Anterior branch of facial suture short, converging forward; posterior branch extending downward initially, then proceeding gently inward and rearward, and finally outward and slightly rearward to meet posterior margin in front of distal intergenal spine. Posterior border furrow narrow, gently convex, with posterior margin extending outward from end of axial furrow, deflected forward at midlength, bearing thick, short intergenal spine at inflection point, and another intergenal spine distally. Pygidium trapezoidal or irregular-hexangular in outline except for anterolateral spines, length 1.2-2 times width, with anteriorlateral margin at angle of about 35 ° to sagittal line, and long, straight lateral margins 15°-20 ° to sagittal line. Axis large, tapering gently rearward, occupying about three-fourths to four-fifths of pygidial length (sag.), rounded or acutely rounded posteriorly with narrow (sag.), bar-like articulating half-ring, four tings defined by straight, shallow ring furrows, and a semicircular or subtriangular terminal piece. Anterior border with narrow, transverse, depressed inner portion and oblique, bar-like outer portion that is about twice as long (tr.) as inner portion; small node or projection on distal end of anterior border present. Pleural field narrow, about half to one-fourth of axial width, with long anterolateral spine, broad at base; posterior border flat, about two-fifths as long as axis, posterior margin arched forward medially; bearing pair of fork-like border spines. Surface of cranidium and pygidial axis covered with densely spaced fine granules. No granules on surface of borders and spines.

Remarks. The cranidium and the paratype pygidia are associated here because they occur in association in a single collection (bed) in the Wangcun section, and because of strong morphological similarities: spinosity, granulosity, and size. The cranidium is rather different from the cranidia that were associated by Zhu (1959, pl. 5, figs. 3, 4) with the pygidia of Chiawangella pustulosa Chu 1959 [=C. pacifica (Walcott, 1913)], the type species of the genus. As discussed above, those cranidia should be referred to Taihangshania. In cranidial characteristics, C. hunanensis is reminiscent of leiostegiids. Differences are in the anterior location of the palpebral lobe, and the intergenal spines on the posterior border. The spinose pygidium is also similar to leiostegiids such as Prochuangia (cf. Prochuangia linicispinata Peng, 1992, fig. 41B-I; P1. 25, figs. 7, 13; P1.26, figs. 5, 6, 8, 10, herein). Other similarities to the leiostegiid pygidium include the short, gently rearward-tapered axis, the deflected anterior border, and the elevated anterolateral spine. The new species C. hunanensis is closely similar to C. pacifica in general respects, but differs in having an elongated outline with a much greater length than width. In the type species, the pygidium is roughly as wide as long. The new species can be further differentiated in having a narrower (tr.) pleural field, a wider (sag.) posterior border with a narrower (tr.) posterior margin, and thus more closely spaced posterior border spines, and greater arching of the posterior margin. Tricrepicephalus asiaticus Yuan and Yin (1998, p. 144, pl. 6, fig. 5) from the Huaqiao Formation at Jimachong, Yuping, eastern Guizhou, is based on a single, fragmental pygidium with a pair of long and stout anterolateral spines. The holotype pygidium may actually belong to Chiawaugella. However, the border of the holotype pygidium is covered by matrix. Further preparation is needed to determine whether it belongs to Chiawangella. T. asiaticus is distinguished from species confidently assigned to Chiawangella by having a shorter, subconical (rather than subcylindrical) axis with narrow (sag., exs.), bar-like tings. • 88"

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2 and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Subfamily ORDOSIINAE Lu, 1954 We follow the original classification of Lu (1954, p. 421) in recognizing this group of trilobites at subfamily rank and placing the subfamily in the family Leiostegiidae. Lu (1954) erected the subfamily Ordosiinae to accommodate one new genus, Ordosia. The type species of Ordosia, O. fimbricauda Lu (1954, pl. 1, figs. 12-18), is clearly a leiostegiid. Lochman-Balk (in Moore, 1959, p. 313-315) suppressed the subfamily Ordosiinae as a junior synonym of Leiostegiinae and classified the genus within the subfamily Leiostegiinae. Later, Zhang (1963, Zhang in Lu et al., 1965) elevated the subfamily to family rank and expanded the genetic concept, emphasizing the structure of the anterior cranidial border, which is defined by border furrows that are confluent medially with the preglabellar furrow and convex-forward at the sides. In some ordosiids, the anterior border bears a wide plectrum. Zhang (1963) referred seven genera to the family, although they may not all be closely related phylogenetically. Zhang and Jell (1987, p. 109) restricted the familial concept slightly by including only four of the previously assigned genera; they also added two genera to the family including the new genus Paralevisia. That assignment seems problematic, as Zhang and Jell (1987) indicated, because their new genus Paralevisia does not completely fit the familial concept. The type species of Paralevisia is.P. globosa. It has a subtriangular cranidium with an ovate, strongly convex glabella (Zhang and Jell, 1987, pl. 43, figs. 3, 4), suggesting a solenoparioid, rather than an ordosioid, affinity. Yuan and Yin (2001, p. 344, 345) transferred Protaizehoia Yang from the Damesellidae to the Ordosiidae, but such an assignment was based on the finding of a pygidium of the genus Protaizehoia. However, the pygidial material of Yuan and Yin (2001) is too poorly preserved to show the morphology of the pygidial spines. Peng et al. (2003b) maintained the original familial status of Protaizehoia because both cephalic and pygidial characters seem most consistent with the inclusion of Protaizehoia among the dameselloids rather than among the ordosioids. Key characters include a glabella with deeply incised lateral furrows, an evenly arched or slightly yoked anterior border furrow, an anterior cranidial border lacking a plectrum, and a pygidium that is spinose in meraspids but reduced in spinosity in holaspids. A thorax has not been recorded previously for ordosioid taxa, but a new specimen of Wanshania from northwestern Hunan (P1. 19, figs. 8-10) shows nine thoracic segments. It is uncertain if other ordosioids had a similar number of segments. Complete specimens of Chuangia and Chosenia (Madaoyuites), both of the subfamily Leiostegiinae (Qiu et al., 1983, pl. 56, fig. 1; Zhang and Wang, 1985, pl. 134, fig. 6; Peng, 1990a, pl. 8, figs. 10, 11), have thoraxes with 10 segments. Genus WANSHANIA Rong and Yang in Zhou et al., 1977 Wanshania Rong and Yang in Zhou et al., 1977, p. 164; Yin and Li, 1978, p. 487; Yang, 1978, p. 40, 41; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 112. Type species. Wanshania wanshanensis Rong and Yang in Zhou et al., 1977 (p. 164, pl. 49, figs. 21, 22), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [-Torifera] tuma Zone; Huaqiao Formation, Dongpo and Jiudiantang, Xinhuang, northwestern Hunan, and • 89"

Jimachong of Yuping and Wanshan of Tongren, eastern Guizhou; by monotypy. Emended diagnosis. Ordosiidae with largely effaced, parallel-side glabella, occipital ring bearing

short spine and centrally located node. Palpebral lobe relatively long (exs.), crescentic, situated posterior of cranidial midlength; anterior border narrow (exs.), with strong plectrum; fixigena wide (tr.) with blade-like posterolateral projection; anterior branches of facial suture curving outwardly; librigena moderately wide (tr.), with border furrow confluent at genal comer and continuing onto genal spine. Thorax with nine segments bearing pleural spines. Pygidium large, semicircular to semielliptical, with long and multisegmented axis and broad (tr.) pleural fields. Remarks. The diagnosis of Rong and Yang (in Zhou et al., 1977, p. 164) is emended here on the

basis of new material, especially on the basis of first recorded exoskeleton and a cephalon with librigenae. Wanshania was classified in the Ordosiidae by Yang (1978) and Zhang and Jell (1987, p. 109) because all of the cephalic and pygidial characteristics are closely similar to those of the genera usually included in this family (Ordosia, Poshania, and Taitzuia; see Lu et al., 1965). Ordosia is similar to Wanshania in glabella shape, fixigenal width, and curvature of the anterior branch of the facial suture, but differs primarily in having a concave anterior border that is defined by a similarly concave border furrow, lacking a plectrum, and having a triangular rather than blade-like posterolateral projection on the fixigena. Lateral glabellar furrows in Ordosia are clearly defined rather than completely effaced as in Wanshania. Poshania, based on Poshania poshanensis Chang (Zhang, 1957, pl. 1, fig. 3.4; 1959, pl. 2, figs. 4-10), has an anterior cranidial border and ocular ridges that are similar to those of Wanshania, but differs in having a forwardly tapering, proportionally shorter glabella, straight rather than curved anterior branches on the facial suture, and a pygidium with clearly defined interpleural furrows that define wire-like anterior bands. Taitzuia is differentiated from Wanshania by its wide (sag., exs.), tumid anterior cranidial border and its subovate glabella. Wanshania wanshanensis Rong and Yang in Zhou et al., 1977

Plate 19, figures 8-10; Plate 24, figures 1-13; Text-figure 10 1977 1978 1978 1978

Wanshania Wanshania Wanshania Wanshania

wanshanensis Rong and Yang in Zhou et al., 164, pl. 49, figs. 21, 22. wanshanensis Rong and Yang; Yang, p. 41, pl. 7, figs. 1, 2. dongpoensis Rong and Yang ; Yang, p. 42, pl. 7, figs. 3-5. wanshanensis Rong and Yang; Yin and Li, p. 487, 488, pl. 163, fig. 15; pl. 165,

figs. 14, 15. 1978 Wanshania dongpoensis Rong and Yang; Yin and Li, p. 488, pl. 165, figs. 9, 10. 1978 Wanshania wanshanensis Rong and Yang; Liu, p. 311, pl. 210, figs. 6, 7. 1980 Peichiashania bispina Ergaliev, p. 146, 147, pl. 3, fig. 18; pl. 5, fig. 16; pl. 6, figs. 1, 2. 1983 Wanshania angustata Qian in Qiu et al., p. 112, pl. 37, fig. 7. Wanshania fenshuiensis Ju in Qiu et al., p. 112, pl. 37, fig. 8. 1983 1987 Wanshania constricta Peng, p. 96, pl. 7, fig. 5. 2001 b Wanshania wanshanensis Rong and Yang; Peng, Babcock, and Lin, p. 103, 105, pl. 6, fig. 9; pl. 8, figs. 4, 5; pl. 14, fig. 5. Lectotype. Cranidium (Rong and Yang in Zhou et al., 1977, pl. 49, fig. 21; also in Yang, 1978, pl. 7, fig. 1, CUGB 1301010) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Dongpo, Xinhuang, northwestern Hunan; designated subsequently as the • 90"

holotype of the species by Yang (1978).

Text-figure 10. A-F, Type material of Wanshania wanshanensis Rong and Yang in Zhou et a1.,1977 and other specimens referred to the species. A, B, testaceous cranidium, lectotype, in dorsal and anterior views; and C, partly exfoliated pygidium, CUGB 1301010, 1301001 x 3.2, z 3.2, × 4(original of Zhou et al., 1977, pl. 49, figs. 21, 22; also Yang, 1978, pl. 7, figs. 1, 2), the cranidium was subsequently

designated as holotype of the species; D, testaceous cranidium, CUGB 1301014, x 4, designated as holotype of Wanshania dongpoensis Rong and Yang in Yang, 1978 (original of Yang, 1978, pl. 7, fig. 3); E, partly exfoliated cranidium, CUGB 1104102, × 3, paratype of Wanshania dongpoensis Rong and Yang in Yang, 1978 (original of Yang, 1978, pl. 7, fig. 5); F, mostly exfoliated pygidium, CUGB 1301011, x 5, paratype of Wanshania dongpoensis Rong and Yang in Yang, 1978 (original of Yang, 1978, pl. 7, fig. 4).

Other species. As indicated in the synonymy list, all other species assigned previously to Wanshania are now regarded as junior synonyms of the type species. Peichiashania bispina Ergaliev, 1980 from southern Kazakhstan is also regarded as a junior synonym of Wanshania

.91"

wanshanensis. New material. More than 200 sclerites, including an exoskeleton, a cephalon, cranidia, librigenae, and pygidia in P240.5, P261, P261.5, P264.2, P269, P273.46, P273.52, P273.66, P273.8, P273.85, P275.1, P277, P277.6, P278.01, P278.1, P279, P282.6, P282.75, P283.47, P283.75, P286.3, P287.1, P290.5, P293, P293.21, P295.13, P296.54, P298.4, P300.4, P301.9, P305.4, P307.4, P310.4, P315.8, P316.0, P325.7, W170, W195.7, W207.5, and W216.5 (illustrated specimens NIGP 137480, 137454137517b). Description. Cranidium rectangular in outline, with transverse or forwardly ached anterior margin, length 0.7-0.8 of width between palpebral lobes. Anterior border with wide (tr.) plectrum of variable width (tr.), extending rearward onto preglabellar furrow. Glabella cylindrical, mostly effaced; S1 weakly bifurcated to fully effaced; $2 and $3 faint to fully effaced. Occipital furrow weakly impressed, deep laterally but not connected with axial furrows; occipital ring slightly narrowing abaxially, bearing tiny medial node subcentrally and tiny spine on posterior margin. Palpebral lobe crescentic, of median size, clearly defined by shallow palpebral furrow, opposite midlength of glabella (including occipital ring); eye ridge extends obliquely rearward. Fixigena variable in width, palpebral area nearly or equal to glabellar width. Anterior branch of facial suture rounded, curving outward; posterior branch strongly divergent rearward, enclosing blade-like posterolateral projection. Librigena moderately wide, with gently convex genal field, moderately wide lateral and posterior borders, and slender genal spine that is posterolaterally directed. Lateral and posterior border furrows confluent at genal comer and extending onto proximal part of genal spine. Thorax with nine segments. Axis narrow, about two-fifths width of pleural region. Pleura transverse, with small posterolateral spine; anterior and posterior bands narrow, separated by wide and relatively deep pleural furrow. Pygidium semicircular to subelliptical in outline, length 0.5-0.6 width. Axis narrow, long, with seven to eight tings and short crescentic terminal piece, tapering gently rearward nearly to posterior border furrow, with narrow and short postaxial ridge. Pleural field wide, moderately convex, with seven to eight ribs separated by shallow and clearly defined pleural furrows, and small terminal area. Anterior border wide abaxially, with facet about half as wide (tr.) as pleural region; lateral and posterior borders fiat, moderately wide, and of uniform width. Remarks. Zhang and Jell (1987, p. 109) suppressed Wanshania dongpoensis, from the Huaqiao Formation of northwestern Hunan, and Peichiashania bispina from the Lejopyge laevigata Zone and the Kormagnostus simplex Zone of Malyi Karatau, Kazakhstan, as junior synonyms of Wanshania wanshanensis, the type species of Wanshania. The type material of both W. wanshangensis and W. dongpoensis is refigured here as text-fig. 10. It shows that W. dongpoensis differs only in the width (tr., sig.) of the anterior border. The anterior border is quite variable in width in the new, abundant material from the Huaqiao Formation of northwestern Hunan. Occurrence. The type material is from the Paradamesops[=Parablackwelderias]jimaensisCyclolorezella [=Torifera] tuma Zone of Dongpo, Xinhuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is the eponymic species of the Wanshania wanshanensis Zone, although it ranges upward into the lower part of the overlying Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the basal part of the Linguagnostus reconditus Zone).

• 92-

Subfamily PAGODIINAEKobayashi, 1935 PAGODIINAEgen. et sp. indet. Plate 22, figure 7 2001b Menocephalites?sp.; Peng, Babcock, and Lin, p. 102, pl. 4, fig. 13.

Material. Incomplete cranidium, NIGP 137498, in collection P136.7. Description. Anterior border upturned, defined posteriorly by shallow anterrior border furrow; preglabellar field narrow, gently depressed. Glabella trapezoidal, length slightly greater than width at base, obtusely rounded anteriorly; S 1 broad, weakly impressed, inward and rearward-directed; other lateral glabellar furrows effaced; occipital ring subtriangular, with stout occipital spine; occipital furrow deep, arched rearward. Palpebral lobe relatively small, reniform, located near cranidial midlength; eye ridge faint, diagonally directed; palpebral area narrow, length about one-third of basal width of glabella. Anterior branch of facial suture converging slightly. Cranidium except for anterior border covered with coarse, widely separated granules. Remarks. A single cranidium left in open nomenclature, bears greatest resemblance to both Prochuangia and Menocephalites. The specimen is granulose, similar to some species of Prochuangia, but the presence of a preglabellar field suggests an affinity with Menocephalites. The stout occipital spine and the upturned anterior border also weigh against an assignment of the specimen to Prochuangia. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone). Genus PROCHUANGIAKobayashi, 1935

Type species. Prochungia mansuyi Kobayashi, 1935 [=Chuangia nais Walcott sensu Mansuy, 1915, p. 20-22, pl. 2, fig. 14a-g; non Walcott, 1911, p. 84, 85; sensu Kobayashi, 1935, p. 186, 187; pl. 8, fig. 8; pl. 10, figs. 1-7; pl. 8, fig. 8 ] from the Prochungia Zone, Saisho-li, South Korea, and lower upper Cambrian (Furongian, at least in part), northeast Vietnam; by original designation. Other species. See Shergold (1980, p. 66), Peng (1992, p. 73), Qian (1994, p. 126, 127), and Shergold et al. (2000, p. 614-616). To these lists, the following three species from the Alborz Mountains, Iran, should be added: Prochuangia sp. aft. P. linicispinata Peng (Peng, Geyer, and Hamdi, 1999, p. 48-50, Fig. 25.1-25.16), Prochuangia leiocephala Peng, Geyer, and Hamdi, (1999, p. 50-53, Fig. 26.1-26.20; 27.1-27.9), and Prochuangia pachycephala Peng, Geyer, and Hamdi, (1999, p. 53-56, Fig. 28.1-28.18). An additional form, described here as Prochuangia sp. cf. P. leiocephala, should also be included in Prochuangia. Remarks. The genetic concept and classification of Shergold (in Shergold et al., 1976, p. 277, 278; •

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2000, p. 614-616; Shergold, 1980, p. 66-69) is followed here.

Prochuangia granulosa Lu, 1956 Plate 25, figures 1-13 1956 1963 1964 1965

Prochuangia granulosa Lu, p. 369, pl. 1, fig. 5. Prochuangia granulosa Lu; Egorova, Xiang, Li, Nan, and Guo, p. 51, 52, pl. 12, figs. 1-5. Prochuangia granulosa Lu; Lu and Qian, p. 33, 34, pl. 8, figs. 7, 8. Prochuangia granulosa Lu; Lu, Zhang, Zhu, Qian, and Xiang, p. 414, 415, pl. 78, figs. 22, 23. 1975 Kaolishania? cf. quadriceps (Dames) (in part), Schrank, p. 598, pl. 5, fig. 3 (only). 1977 Prochuangia granulosa Lu; Zhou, Liu, Meng, and Sun, p. 202, pl. 60, figs. 5, 6. 1978 Prochuangia granulosa Lu; Yin and Li, p. 514, pl. 174, figs. 1, 2. non 1978 Prochuangia granulosa Lu; Yang, p. 81, pl. 11, figs. 12, 13 [=Prochuangia linicispinata Peng, 1992]. 1981 Prochuangia granulosa Lu; Zhang, p. 178, 179, pl. 65, figs. 5, 6. 1983 Prochuangia granulosa Lu; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, p. 188, pl. 63, figs. 1, 2. ?1984 Prochuangia granulosa Lu; Wittke, p. 139, pl. 8, figs. 13, 14. 1992 Prochuangia granulosa Lu; Peng, p. 75, fig. 41J-M. 1999 Prochuangia granulosa Lu; Peng, Geyer, and Hamdi, p. 47, 48, fig. 24. 200 lb Prochuangiagranulosa Lu; Peng, Babcock, and Lin, p. 106, pl. 16, figs. 10, 11. 2001d Prochuangiagranulosa Lu; Peng, Babcock, Lin, and Chen, p. 142, fig. 10.10. Holotype. By monotypy; incomplete pygidium (Lu, 1956, pl. 1, fig. 5; NIGP 8646) from the upper Cambrian (Furongian) at Longtiancong, Wanshan, eastern Guizhou. New material. More than 30 sclerites including cranidia, broken librigenae, and pygidia (illustrated specimens NIGP 137518-137528) are in collections P378.25, PI322.4, PI323.5, PI327.2, P[336, P[355.5, P1356.5, PI360.3, P1370.3 and P1372. Description. Anterior cranidial margin transverse or arched gently forward; anterior border deflected, with posterior slope nearly vertical and anterior slope inclined moderately forward. Librigena with wide (tr.) genal field, shallow lateral border furrow, and prominent eye socle. Omamentation consists of coarse, densely spaced granulation. Remarks. New material from the middle part of the Huaqiao Formation in northwestern Hunan resembles in all respects the holotype and other materials from the same region. Key characters of the species as shown by Lu (1956) and Egorova et al. (1963) are the coarse and densely spaced granulation on the cranidium, the pygidial axis with six tings and a terminal piece, and the fine granulation on the pygidium. Occurrence. The holotype is from the upper Cambrian (Furongian Series) at Longtiancong, Wanshan, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it is associated with trilobites indicative of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). • 94

•

Prochuangia linicispinata Peng, 1992 Plate 26, figures 7-11 1978 1992

Prochuangiagranulosa Lu; Yang, p. 81, pl. 11, figs. 12, 13. Prochuangialinicispinata Peng, p. 75, fig. 41B-I. 2001c Prochuangiasp.; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 6, fig. 12. 200 ld Prochuangialinicispinata Peng; Peng, Babcock, Lin, and Chen, p. 142, figs. 10.8, 10.9. Holotype. Cranidium (Peng, 1992, fig. 41B, NIGP 95196) from the Glyptagnostus reticulatusChuangia wulingensis Zone at Wa'ergang, Taoyuan, northwestern Hunan. New material. More than 10 sclerites (illustrated specimens NIGP 133577, 137539-137542) in collections P374.9, P375, P375.15, P378.25, PI3-1.6, and PI34.3.

Remarks. New material from the Huaqiao Formation, northwestern Hunan, resembles the type material from the same region. Key characters that warrant assignment of the new material to this species include scattered granules on the cephalon and a slightly tapered glabella that is constricted laterally.

Occurrence. The holotype is from the Glyptagnostus reticulatus-Chuangia wulingensis Zone at Wa'ergang, Taoyuan, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone and the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). Prochuangia sp. cf. P. leiocephala Peng, Geyer, and Hamdi, 1999 Plate 25, figures 14-18; Plate 26, figures 1-6 cf. 1999 Prochuangialeiocephala Peng, Geyer, and Hamdi, p. 50-53, figs. 26, 27.

Material. More than 30 cranidia and pygidia (illustrated specimens NIGP 137529-137538) are in collections P374.9, P375.15, and P378.25.

Remarks. The new specimens, left in open nomenclature, represent a smooth to punctuate Prochuangia species. Most cranidia and pygidia in the collections from the Huaqiao Formation are rather densely punctate, both on the test and on exfoliated surfaces. This species has a relatively wider (sag., exs.) anterior border that is flat and deflected. The species also has a strongly tapered glabella in holaspids: the anterior part of the glabella is transverse, and extends slightly onto the anterior border. These features are mostly reminiscent of P. leiocephala from the Alborz Mountains, Iran (Peng et al., 1999, figs. 26, 27), which also is a smooth to punctate species of Prochuangia. Other characters comparable to those of the Iranian species include the gently oblique orientation of the eye ridge and the course of the facial sutures, with the anterior branch being subparallel in large holaspids. However, the associated pygidia from Hunan are quite distinct from those of Iranian P. leiocephala. In the Iranian material, the pygidial • 95

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axis is more tapered rearward, the posterior margin between the spines is gently curved rather than deeply bowed, and the borders are much wider. As a result, the pleural fields of the Iranian specimens are much smaller than those of the Hunan specimens. Furthermore, in the Iranian material, the first pleural furrow in the pygidium is strongly effaced rather than deeply incised as it is in the Chinese material. The new material from Hunan also resembles Prochuangia depressa (Qian, 1994, pl. 28, figs. 7-10; pl. 29, figs. 1-3; pl. 30, figs. 1-6; text-fig. 42) from the Prochuangia mansuyi Subzone (of the Chuangia Zone) of the Changshan Formation in southern Liaoning. P. depressa differs, however, in having a transverse rather than forward-arched anterior border furrow, a more tapered glabella, and a proportionally wider pygidium with more strongly based posterolateral spines.

Occurrence. The holotype is from the Glyptagnostus reticulatus-Chuangia wulingensis Zone at Wa'ergang, Taoyuan, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone and the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). Order

LICHIDA

Moore, 1959

Superfamily DAMESELLOIDEAKobayashi, 1935 Family DAMESELLIDAEKobayashi, 1935 Subfamily DAMESELLINAEKobayashi, 1935 Genus DAMESELLAWalcott, 1905

Type species. Damesella blackwelderi Walcott, 1905 [junior synonym of Damesella paronai (Airaghi, 1902)] from the Changhia Formation, southwest of Yangzhuang, central Shandong, China; by original designation. Remarks. The genetic concept of Walcott (1905, p. 34, 35; 1913, pl. 23, 24) is followed here. Damesella hunanensis sp. nov. Plate 27, figures 1-12 1978

Blackwelderia granosa Endo; Yang, p. 81, pl. 11, fig. 6 (misspelled as Blackwelderia granulosa). 2001b Damesellasp., Peng, Babcock, and Lin, p. 103, pl. 8, fig. 2. 2001b Blackwelderiagranosa Endo; Peng, Babcock, and Lin, p. 104, pl. 10, fig. 2 (misspelled as Blackwelderia granulosa). Etymology. From Chinese, Hunan, in reference to Hunan Province, where the type material was collected. Holotype. Cranidium (P1.27, fig. 5, NIGP 137546) in collection P277. Other material. Six cranidia and five pygidia (illustrated speciemens NIGP 137543-137545, • 96

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137547-137551) in collections P277, P294.51 and W211.7. Diagnosis. Damesella having sinuous cranidial margin, convex glabella with deeply incised S 1 and $2 lateral furrows, occipital furrow bowed rearward, and rounded at front; fixigena two-thirds as wide as glabella at S1; anterior cranidial border narrow (sag.), convex, defined by wavy border furrow; eye ridges clear, strong. Pygidium subtriangular, with axis and pleural region subequal in width; seven pairs of marginal spines, with the anterior and fifth pairs longer than others. Cranidium and pygidium covered with coarse granules. Description. Cranidium width about twice length, moderately convex. Glabella wide, tapering forward, broadly rounded to acutely rounded at front; S1 deep, posteromedially directed, nearly isolating small lateral lobe; $2 short and deep, nearly transverse; $3 short and shallow, anteromedially directed. Occipital furrow arched posteriorly; occipital ring convex (tr., sag.), of nearly uniform width; fixigena width three-fifths as wide as glabella; palpebral lobe short (exs.), weakly defined and posteriorly directed, opposite L2; eye ridge stout and straight; posterior border furrow deep; and posterior border a narrow ridge. Pygidium width three-fourths length. Axis long, consisting of four to five tings and short semicircular terminal piece, tapering posteriorly, extending to posterior border. Pleural field containing five deep pleural furrows and four shallow interpleural furrows. Marginal spines consist of seven long pairs. Cranidial surface with densely spaced granules of bimodal size; pygidium including spines covered with fine, densely spaced granules. Remarks. One pygidium (P1. 27, figs. 10-12) had been previously assigned to Blackwelderia granosa because of its general resemblance to the pygidia assigned to that species (Endo, 1937, p. 323, pl. 63, figs. 7-12). However, associated cranidia suggest that the pygidium and cranidia instead represent a new species of Damesella. The cranidium of the new species differs from that of B. granosa in having a ridge-like anterior border rather than a thin, upturned anterior border and a narrow rather than broad anterior border furrow. The new species also differs in having a less tapered axis in the pygidium. This new species is closely similar to Damesella dongvanensis Mansuy 1915 (p. 13-17, pl. 2, figs. 7a-7n) from the middle Cambrian of Drng-van, northeast Vietnam, but D. dongvanensis differs in having a much more tapered, less convex glabella, and a narrower palpebral area of the fixigena. D. hunanensis sp. nov. is also similar to Damesella huoshanensis Zhang and Wang, 1985 (p. 461, pl. 137, figs. 10, 11) from the Changhia Formation at Sanyanyao, Huoshan, Shanxi, North China. The new species differs from D. huoshanensis in having a more distinctly furrowed glabella, relatively wider (tr.) fixigenae, and stronger eye ridges. Damesella huoshanensis is possibly synonymous with both Menocephalites abderus (Walcott, 1905, p. 88; 1913, p. 173, pl. 16, fig. 3; refigured in Zhang and Jell, 1987, pl. 102, figs. 6, 7) from the Changhai Formation of Shandong, North China, and Damesella bella Resser and Endo (1937, p. 199, pl. 45, figs. 1, 2, 7, 8; refigured in Zhang and Jell, 1987, pl. 99, fig. 10; pl. 98, fig. 3) from the Changhai Formation of Liaoning, Northeast China. Menocephalites abderus is more likely to be a species of Damesella than Menocephalites. If these three species concepts represent a single conspecific form, then Menocephalites abderus, based on a single cranidium, has priority. No pygidium was assigned to either D. huoshanensis or M. abderus, but a pygidium with seven pairs of marginal spines was assigned to D. bella (Resser and Endo, 1937, pl. 45, figs. 7, 8). A pygidium that is morphologically similar to that of D. bella (Walcott, 1913, pl. 18) was present in association with the holotype •

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cranidium of M. abderus in collection C19 of Blackwelder (1907, p. 33). The pygidium was named by Walcott as Damesella brevicaudata (Walcott, 1913, p. 129, pl. 9, fig. 9; refigured in Zhang and Jell, 1987, pl. 98, figs. 1, 2). It is possible that D. bella and D. brevicaudata, based on pygidia, and D. abderus, based on a cranidium, all belong to the same species. Zhang and Jell (1987) however followed Walcott in recognizing sclerites as representative of separate species. Material of Damesella hunanensis sp. nov. from the Huaqiao Formation of northwestern Hunan suggests that the cranidium and pygidium associated by Resser and Endo (1937) for Damesella bella is reasonable, and that the cranidium-based M. abderus and the pygidium-based D. brevicaudata are conspecific. Furthermore, available material indicates that M. abderus is referable to Damesella. This conclusion is supported particularly by the material of D. brevicaudata, most of which is from a single collection from the Changhia Formation of Liaoning, Northeast China (Guo et al., 1996, pl. 66, figs. 1-3a, 4-14). Both the cranidia and the pygidia assigned to D. brevicaudata in the material of Guo et al. (1996) are identical in almost all respects with those that Resser and Endo (1937) assigned to D. bella. The cranidia are also indistinguishable from the holotype cranidium of Menocephalites abderus. The cranidium-based Menocephalites abderus and the pygidium-based Damesella brevicaudata are here regarded as belonging to a single species, and D. brevicaudata is regarded as a junior synonym of M. abderus. M. abderus is a damesellid species and should be transferred to Damesella. D. bella, and D. huoshanensis are tentatively regarded as junior synonyms of M. abderus. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the upper part of the Proagnostus bulbus Zone and the lower part of the Linguagnostus reconditus Zone). Damesella? sp.

Plate 57, figure 8 Material. Single fragmental pygidium, NIGP 137842, from P331.8. Remarks. The single known pygidium is incompletely preserved. Observed features, particularly macropleural spines on the first segment and at least six pairs of short pleural spines on the successive segments, indicate that the species is a damesellid. The pleural region bears rather broad, moderately deep pleural furrows that become pit-like depressions distally. The interpleural furrows are almost completely effaced. Among damesellids, distally depened pleural furrows are commonly observed in Damesella (see Zhang and Jell, 1987, pl. 96, figs. 5-7, 9, 10; pl. 97, figs. 4, 10; pl. 98, figs. 1-3). For this reason, this fragmental pygidium is tentatively referred to that genus. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the upper part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone).

Genus BLACKwELDERIAWalcott, 1906 Type species. Calymmene? [misprint for Calymene?] sinensis Bergeron, 1899 (p. 500-503, pl. 13, fig. 1, text-figs. 1, 2) from unknown formation and locality in North China (possibly from the • 98"

Kushan Formation, Dawenkou, central Shandong, China); by original designation. Remarks. The concept of the genus Blackwelderia has been discussed at length by several authors (Walcott, 1913; Kobayashi, 1942b; Lu et al., 1965; Opik, 1967). Walcott's original concept is followed here. The type species, Blackwelderia sinensis, and all other taxa described by Bergeron (1899), except for Arthricocephalus chauveaui, are from a single slab of "Bianfushi (Bat-stone)", a name used locally for the B. sinensis and Drepanura premesnili-rich mudstone (see remarks under genus Drepanura). Monke (1903) and Walcott (1913) noted that the cranidium and librigena described as Calymene? sinensis, and the associated pygidia described as Olenoides leblanci belong to the same species, and that O. leblanci should be suppressed. Bergeron's (1899) syntypes of both Calvmene? sinensis and Olenoides leblanci are refigured on P1.45. One of the syntypic cranidia of Calymene? sinensis (P1.45, fig. 1B) is here chosen as the lectotype of the species. Blackwelderia voushuica sp. nov. Plate 28, figures 1-7 200 lb Blackwelderia sp., Peng, Babcock, and Lin, p. 103, pl. 6, fig. 10. Etymology. From Chinese, named for the Youshui River, along which occurs the Wangcun section, from which the holotype was collected. Holotype. Testaceous, broken cranidium and its counterpart (P1. 28, figs. 3-5, NIGP 137553) in collection W 132.5. Other material. One small cranidium and one large, fragmental cranidium (NIGP 137552, 137554), also in collection W 132.5. Diagnosis. Blackwelderia with broad subconical glabella, bacculae not completely isolated; palpebral lobes well-defined, reniform, lying at about cranidial midlength; anterior border flat and wide, gently upturned; eye ridges well-defined, strongly oblique. Description. Preglabellar field absent; anterior border moderately wide (sag.), gently upturned. Glabella tapering moderately forward, truncate anteriorly in small holapsids, but obtusely rounded in large holapsids; with long, deep S 1, slightly sinuous laterally; $2 short or shallow pit; L 1 with a subovate baccula defined by weak furrow subparallel to S 1. Occipital furrow shallow, deep at side; occipital ring crescentic with median node and transverse ridge medially. Eye ridge weak to well defined, oblique rearward at about 70 ° angle to sagittal line; palpebral lobe reniform, located opposite L2, defined by outwardly arched palpebral furrow. Anterior branch of facial suture straight initially, curving across anterior border furrow and anterior border; posterior branch straight, strongly divergent rearward at about a 60 ° angle to sagittal line, enclosing triangular posterolateral projection of fixigena with length about 0.5 of width. Posterior border furrow moderately incised, posterior border narrow and of uniform width (exs.). Remarks. The new species is most similar to Blackwelderia longispina Resser and Endo (1937, p. 186, pl. 50, figs. 8-11; Zhang and Jell, 1987, pl. 103, figs. 8-10) from the Kushan Formation near •

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Dapingzhuang (Tapingchuang), Changxingdao Island, Liaoning. The nature of the anterior region, the course of the facial suture, and the shape of the posterior area of the fixigena in both species are almost identical. However, the new species differs in having glabellar bacculae, and clearly rather than obscurely defined eye ridges. The S 1 and $2 furrows are isolated from the axial furrows in B. youshuica, but are connected with the axial furrows in B. longispina. The new species is morphologically similar to some species assigned to Parablackwelderia. The strongly tapered glabella with nearly straight sides and rounded front, the deeply incised and strongly oblique S 1 and $2 furrows that are isolated from the axial furrows, and the clearly defined eye ridge, are all features typical of Parablackwelderia. The new species, however, differs from Parablackwelderia in having normal rather than stalk-like palpebral lobes. Stratigraphically, the new species occurs just below the earliest Parablackwelderia. Morphological similarity between B. youshuica and Parablackwelderia, and the stratigraphic ranges of the two taxa, suggests a close phylogenetic relationship between Blackwelderia and Parablackwelderia. Blackwelderia youshuica or a closely related species probably gave rise to the Parablackwelderia stock.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is associated with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Blackwelderia? sp. Plate 45, figure 2

Material. Incomplete pygidium (NIGP 137709) in collection P269. Remarks. Single granulose pygidium left in open nomenclature, has a subcylindrical axis with five tings and a terminal piece, deep pleural furrows and faint interpleural furrows, and seven pairs of border spines. The fifth pair of border spines seems stouter and longer than the others. The pygidium is most similar to the pygidium of Blackwelderia sinensis (P1. 45, fig. l b', b"), which differs only in having border spines of even length, and in lacking granulose sculpture. In having a long, stout spine, the undetermined pygidium is similar to a pygidium of B. chiawangensis Chu (Zhu, 1959, pl. 5, fig. 1) that was referred to B. granosa Endo by Yang (1978, pl. 11, fig. 6, misspelled as B. granulosa). However, in both of those species, the sixth pair of spines, rather than the fifth pair, is long and stout. Among the species from the Paibi and the Wangcun sections, the new pygidium bears some similarity with pygidia of Damesella hunanensis sp. nov. (P1. 27, figs. 7-12) and Taihangshania wangcunensis sp. nov. (P1. 37, fig 14; P1. 38, figs. 1-6). The pygidium of D. hunanensis sp. nov. has narrower pleural fields and coarser and less dense granules, whereas T. wangcunensis sp. nov. has wider (exs.) pleural bands on the first segment, more oblique pleural furrows, and a shorter, more slender fifth pair of border spines. This general morphology of the single known pygidium is consistent with Blackwelderia, but until an associated cranidium is discovered, that assignment remains uncertain. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone).

"100.

Genus PARABLACKWELDERIAKobayashi, 1942b Parablackwelderia Kobayashi, 1942b, p. 210; 1960b, p. 352; Hupf, 1955, p. 166; Lochman-Balk in Harrington et al., 1959, p. O318; Lu, Qian, and Zhu, 1963, p. 110; Luo, 1974, p. 653; Nan, 1976, p. 339. Damesops Chu (Zhu), 1959, p. 69, 70; Kobayashi, 1960b, p. 352, 353; Lu, Qian, and Zhu, 1963, p. 110; Lu, Zhang, Zhu, Qian, Xiang, 1965, p. 390; Nan, 1980, p. 504; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 177, 178; Guo, Zan, and Luo, 1996, p. 125; Zhang and Jell, 1987, p. 216; Jell and Hughes, 1997, p. 100. Meringaspis Opik, 1967, p. 323; Ergaliev, 1980, p. 149; Lisogor, Rozov, and Rozova, 1988, p. 71. Paradamesops Yang in Lu et al., 1974a, p. 86; Zhou, Liu, Meng, and Sun, 1977, p. 198; Yin and Li, 1978, p. 511; Yang, 1978, p. 58; Zhang, 1981, p. 177; Liu, 1982, p. 321; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 179; Peng, 1987, p. 104; Kim, 1987, p. 58; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, 1991, p. 166; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, 1993, p. 215; Guo, Zan, and Luo, 1996, p. 127. ?Guancenshania Zhang and Wang, 1986, p. 667,668. Type species. Blackwelderia spectabilis Resser and Endo, 1937 (p. 188, pl. 52; also Zhang and Jell, 1987, p. 214, pl. 102, figs. 8, 9; pl. 103, figs. 3-5) from the Drepanura Zone, near Haishang, Changxindao Island, southern Liaoning, China; by original designation. Lectotype of B. spectabilis, selected here, a cranidium from Resser and Endo's syntypes (1937, pl. 52, insert figure, second cranidium from fight; also Zhang and Jell, 1987, pl. 103, fig. 3, uppermost cranidium; USNM 86767d). Emended diagnosis. Damesellidae with glabella broadly conical, rounded or acutely rounded in front; palpebral lobe small, stalk-like, lying anterior to midlength of glabella (excluding occipital ring); preglabellar field narrow (sag., exs.) or absent; posterior branch of facial suture straight, strongly oblique or diagonally directed; posterolateral projection of fixigena long (exs.), triangular. Pygidium with slender axis and six pairs of lateral border spines; border spines of unequal length, sixth pair long and stout; pygidial margin between the sixth pair of spines rounded, angulate, or rounded with additional pair of short spines. Other species. Blackwelderia chiawangensis Chu, 1959 (Zhu, p. 67, pl. 5, fig. 1) [=Blackwelderia disticha Zhang in Qiu et al., 1983, p. 175, pl. 57, figs. 9, 10] from the Drepanura premesnili Zone of Kushan Formation, Jiawang, northern Jiangsu; Damesops convexus Chu, 1959 (Zhu, p. 70, pl. 5, figs. 2, 3) [=Damesops sponosus Guo and Luo in Guo et al., 1996. p. 125, 126, pl. 68, figs. 7-9], from the Blackwelderia paronai Zone of Kushan Formation, Jiawang, northern Jiangsu; Meringaspis meringaspis Opik, 1967 (p. 323, 326, pl. 44, figs. 6* (fight-center rather than below), 7-9; pl. 45, figs. 1, 2; text-figs. 119-121) from the Glyptagnostus stolidotus Zone in the O'Hara Shale, Queensland, Australia; Paradamesopsjimaensis Yang in Lu et al., 1974a (p. 86, pl. 1, fig. 15 only) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Jiudiantang, Xinhuang, northwestern Hunan; Paradamesops laterilobatus Yang in Zhou et al., 1977 (p. 198, 199, pl. 59, figs. 1, 2) from the Paramphoton Zone and Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan; Meringaspis huabeiensis Zhang in Qiu et al., 1983 (p. 181, pl. 59, figs. 5a, 5b) from the Changhia Formation, Jiawang, northern Jiangsu; • 101.

Paradamesops triangularis Guo and Luo in Guo et al., 1996 (p. 127, pl. 68, figs. 10, 14) from the Blackwelderia-Damesops [=Parablackwelderia] Zone of the Kushan Formation, Jinxian, Liaoning; Damesops sheridanorum Jell and Hughes (1997, p. 100, 101, pl. 32, figs. 1-7, 9) from the Kurgiakh Formation, Zanskar Valley, Ladakh, India; and Parablackwelderia sp. cf. P. huabeiensis (Zhang), described herein. Meringaspis sp. B (Opik, 1967, p. 326, pl. 46, figs. 1, 3) is possibly a junior synonym of Parablackwelderia meringaspis. Some pygidia described under the names of Blackwelderia alastor Walcott (Mansuy, 1915, p. 12, fig. 12; 1916, p. 21, 22, pl. 2, fig. 5) and Blackwelderia biloba (Kobayashi, 1942c, p. 43, pl. 2, fig. 4) probably belong to Parablackwelderia. Two cranidia described under the names Menocephalus belenus Walcott (1905, p. 62) and Dorypygella alastor Walcott (1905, p. 31) also probably belong to Parablackwelderia. Walcott (1913) transferred these two species to Lisania, whereas Zhang and Jell (1987) questionably transferred both species to Damesops. A cranidium described as Damesops angustus (Guo and Luo in Guo et al., 1996, p. 126, pl. 10, fig. 13) probably represents a species of Parablackwelderia. Remarks. Jell and Hughes (1997) synonymized Damesops Chu, 1959 with Meringaspis Opik, 1967, and Paradamesops Yang in Lu et al., 1974a. All three genera are here considered to be junior synonyms of Parablackwelderia Kobayashi, 1942b. This genus was referred to the subfamily Drepanuriinae (Hupr, 1955; Lochman-Balk in Moore, 1959; Kobayashi, 1960b), but we prefer to follow Opik (1967) in classifying it with the B lackwelderiinae. Kobayashi (1942b, p. 210) erected Parablackwelderia with Blackwelderia spectabilis (Resser and Endo, 1937, p. 188, pl. 52) as the type species. Kobayashi (1942b, p. 210) diagnosed the genus using its small, anteriorly located eyes. According to Kobayashi (1942b), the genus is also differentiated from Blackwelderia by its broad librigena, and its truncate-conical glabella. A more detailed diagnosis for this genus was provided by Lochman-Balk in Moore (1959), but that diagnosis was chiefly based on Kobayashi's (1942b) redescription of the type species of Parablackwelderia. By including Damesops, Meringaspis, and Paradamesops in Parablackwelderia the diagnosis is emended here to embrace a wider genetic concept. Zhang and Jell (1987, p. 214, pl. 102, figs. 8, 9; pl. 103, figs. 3-5) refigured the type material of B. spectabilis and interpreted the genetic characters noted by Kobayashi (1942b) for Parablackwelderia to be of only species-level value. Zhang and Jell (1987) regarded Parablackwelderia as a junior synonym of Blackwelderia. Here, Parablackwelderia is retained as a separate genus because it differs from Blackwelderia in several important respects. Most importantly, Parablackwelderia has a long eye stalk that extends outward and upward to support the small, anteriorly placed eye lying opposite the glabellar frontal lobe (see Zhang and Jell, 1987, pl. 103, figs. 3, 4; Text-fig. 11 herein). This feature has not been observed in any species of Blackwelderia. The glabella of Parablackwelderia is proportionally larger than that of Blackwelderia, and is usually broad-conical in shape with nearly straight sides and a rounded front. In Blackwelderia, the palpebral lobe is located near the cranidial midlength, and the glabella is relatively smaller in proportion (Bergeron, 1899, p. 500-503, pl. 13, figs. 1, 5; also Zhang and Jell, 1987, pl. 100, figs. 7-10; pl. 101, figs. 1-6; pl. 104, fig. 9), with the sides more or less constricted, and the front truncate or evenly rounded. Parablackwelderia may be further differentiated from Blackwelderia by having a straight, obliquely directed S 1, a shorter anterior cranidial border, and a shorter preglabellar field. The pygidium of Parablackwelderia also seems to be distinctive from Blackwelderia. Pygidia of species assigned to Blackwelderia and Parablackwelderia differ in minor ways. Parablackwelderia usually has a more slender and more segmented axis; it also bears pygidial border spines of varying size, among which the first and particularly the sixth pairs are stouter and much longer than the others. In Blackwelderia, the pygidial axis is relatively wide and less segmented; the pygidial border spines are normally subequal in length and width. Some species • 102•

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and Endo, 1937), the type species of Purubluckwelderiu from the Drepunuru Zone of southern Liaoning, China; based on specimens figured by Resser and Endo (1937, pl. 52); also by Zhang and Jell (1987, pl. 102, fig. 9, USNM 86767b; pl. 103, figs. 3 , 4, USNM 86767d-f, g); C, D, Purubluckwelderiu meringuspis (dpik, ~

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jimaensis (Yang in Lu et al., 1974a), based on specimens NIGP 13757 1-137573, and 137581 (see PI. 30, figs. 5-7, 15).

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1967) from the Glyptugno.~tusstolidotus Zone of Queensland, Australia, based on specimens figured by opik (1967, pl. 45, figs. 1, 2); and E, F, Purubluckwelderiu • 103•

currently assigned to Blackwelderia that have a stout sixth pair of pygidial spines (e.g.B. chiawangensis, Zhu, 1959, pl. 5, fig. 1) should perhaps not be referred to Blackwelderia, but to Parablackwelderia instead. The genus Paradamesops is apparently a junior synonym of Parablackwelderia. Type material of Paradamesops jimaensis, the type species from northwestern Hunan, has no distinct differences from Parablackwelderia spectabilis, the type species of Parablackwelderia, except for lacking the posterior (seventh) pair of marginal spines on the posterior margin. Such a difference is probably of only specific value. Meringaspis Opik, 1967 from Queensland, Australia, is also a junior synonym of Parablackwelderia. The type species M. meringaspis apparently bears an abaxially elevated eye stalk, which is characteristic of both Parablackwelderia and Paradamesops. It resembles Parablackwelderia spectabilis and Paradamesops jimaensis in all essential aspects except for having a pointed or triangular posterior portion of the pygidium, as interpreted by 0pik (1967, p. 325, text-fig. 121), and having somewhat more posteriorly located palpebral lobes. However, new material of Parablackwelderia from Hunan indicates that the location of the palpebral lobe is somewhat variable, although it is always anterior to the distal end of the S 1 furrow (which is near the midlength of the glabella, excluding the occipital ring). Other material of Meringaspis (e.g., Meringaspis sp. A sensu 0pik, 1967) tends to confirm that the palpebral lobe is located variably in that genus, and that it falls well within the variation of Parablackwelderia. Upon further examination of photographs of the type material of M. meringaspis, the posterior portion of one pygidium (0pik, 1967, pl. 45, fig. 2), which is the best preserved pygidium in Opik's collection of Meringaspis, is broken, and its shape cannot be positively identified. More material is needed to confirm whether the posterior pygidial border is pointed. Even if the pygidium has a pointed posterior border, the distinction should be interpreted as being of only specific, rather than genetic, significance. Guancenshania Zhang and Wang, 1986 is tentatively considered to be a junior synonym of Parablackwelderia. Guancenshania is most comparable to Meringaspis, as both genera have palpebral lobes that are placed opposite L2. Differences between Guancenshania and Meringaspis are minimal, and they are most likely synonymous. Damesops is also regarded as a junior synonym of Parablackwelderia. The type species of Damesops is D. convexus Chu (Zhu, 1959, p. 69, 70; pl. 5, figs. 2, 3). The holotype of the type species is an incomplete cranidium having the palpebral lobes broken away. The subcentrally located palpebral lobes are reminiscent of Meringaspis, and the pygidium agrees well with that of Parablackwelderia. Morphologically the holotype cranidium is closely comparable to both Paradamesops and Meringaspis in having a broadly conical glabella, double eye ridges and by lacking a preglabellar field. Upon examination of the type material of the type species, there appears to an incomplete stalk-like palpebral lobe on the holotype cranidium, suggesting that it is closely comparable with Parablackwelderia. We concur with Jell and Hughes (1997) in synonymizing Damesops Chu, 1959 with Meringaspis Opik, 1967 and Paradamesops Yang in Lu et al., 1974a. As a result, Damesops also should be suppressed as a junior synonym of Parablackwelderia.

Parablackwelderia jimaensis (Yang in Lu et al., 1974a) Plate 29, figures 1-13; Plate 30, figures 1-15; Plate 57, figures 6, 7" Text-figures 11E, F, 12A-D 1963 1963 • 104.

Blackwelderia sp. (in part), Egorova, Xiang, Li, Nan, and Guo, p. 40, pl. 8, fig. 3 only; non fig. 4 [= Proitaizehoia granifera Yang, 1978]. Genus and species indeterminate, Egorova, Xiang, Li, Nan, and Guo, p. 43, pl. 8, figs. 7, 8.

1965

Blackwelderia sp. (in part), Lu, Zhang, Zhu, Qian, and Xiang, p. 383, 384, pl. 72, fig. 3 only; non fig. 4 [= Proitaizehoia granifera Yang, 1978 ].

1965

Genus and species indeterminate, Lu, Zhang, Zhu, Qian, and Xiang, p. 648, 649, pl. 133, figs. 20, 21. 1967 Meringaspis sp. A, Opik, p. 326, pl. 46, figs. 2-5. 1974a Paradamesops jimaensis Yang in Lu et al. (in part), p. 86, 87, pl. 1, figs. 15a, b; pl. 2, figs. 1la, b; non pl. 1, figs. 16a, b [=Parablackwelderia yangi sp. nov.]. 1978 Paradamesops jimaensis Yang (in part); Yin and Li, p. 511, pl. 169, fig. 10 non fig. 11 [=Parablackwelderia yangi sp. nov.]. 1978 Paradamesops jimaensis Yang (in part); Yang, p. 58, pl. 13, figs. la, b; non fig. 2 [=Parablackwelderia yangi sp. nov.]. 1980 Meringaspis karatauensis Ergaliev, p. 149, 150, pl. 3, fig. 15; pl. 6, figs. 9-12. 1981 Paradamesops jimaemsis Yang; Zhang, p. 177, pl. 66, figs. 4, 5. 1981 Paradamesops karuktagensis Zhang, p. 178, pl. 65, fig. 3. 1983 Paradamesops depressus Qian in Qiu et al., p. 180, pl. 61, fig. 1. 1983 Paradamesopsflatilimbatus Qiu in Qiu et al., p. 180, pl. 60, fig. 7. 1983 Paradamesops trigonatus Qian in Qiu et al., p. 180, pl. 60, figs. 9, 10. 1983 Paradamesops latilimbatus Qian in Qiu et al., p. 180, pl. 59, figs. 3, 4. 1983 Paradamesops lirellatus Qiu in Qiu et al., p. 180, 181, pl. 60, fig. 8. 1983 Paradamesops reticulatus Qiu in Qiu et al., p. 181, pl. 60, fig. 11. 1987 Paradamesops karatauensis (Ergaliev); Peng, p. 104, pl. 8, figs. 9-11; pl. 9, fig. 1. 1987 Paradamesops rieni (Kim) (in part), p. 58, pl. 21, fig. 3, non fig. 11. ?1987 Paradamesops whangjuensis (Kim), p. 58, 59, pl. 21, figs. 2, 4, 10. 1988 Meringaspis karatauensis Ergaliev; Lisogor, Rozov, and Rozova, p. 71, pl. 6, figs. 6, 7. 1991 Paradamesops qinglingensis Yang in Yang et al., p. 166, pl. 21, figs. 3, 4. 1993 Paradamesops qinglingensis Yang in Yang et al., p. 215, 216, pl. 21, figs. 3, 4. 2001 b Meringaspis jimaensis (Yang); Peng, Babcock, and Lin, p. 103, pl. 7, figs. 2-4. 2001 d Meringaspis karatauensis Ergaliev; Peng, Babcock, Lin, and Chen, p. 141, figs. 9.10, 9.11. Holotype. Testaceous cranidium (Lu et al., 1974a, pl. 1, fig. 15; refigured in Yin and Li, 1978, pl. 69, fig. 11, Yang, 1978, pl. 13, fig. 1; CUGB 1211301) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [= Torifera] tuma Zone of the Huaqiao Formation, Jimachong,

Yeping, eastern Guizhou. New material. More than 80 sclerites including cranidia, pygidia, and a hypostome (illustrated

specimens NIGP 137558-137581, 137841), in collections P199.9, P230.09, P237.25, P259.85, P260, P261.35, P269, P273.66, P273.8, P275.1, P277, P278.1, P279.7, P282.75, P283.47, P283.67, P283.75, P290.5, P293.21, P295.13, P298.54, P341.8, W 187.8, W 188.8, W 196.3, W 199.2, W210.5, W211.7, W218.3, W219.7, and W225. Description of new sclerites. Librigena with narrow, moderately convex genal field; having long,

flat genal spine, broad at base; lateral border fiat, wide (tr.) at genal angle, narrowing evenly forward; posterior border short (tr.) and moderately wide (exs.). Hypostome length equal to width at front. Anterior margin arched gently anteriorly; posterior margin broadly curved with broad but shallow indentation sagittally, merged with posterior margin of median body. Median body inverted trapezium in outline, moderately convex; median furrow wide and deep, diagonally directed, with inner end close to sagittal line; posterior lobe short (sag.) crescentic in outline. Lateral border narrow, triangular, developed only posterior of the midlength •

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of the hypostome, inclining strongly inward; posterolateral border wing-shaped, as wide as posterior lobe of median body (exs., tr.), with length being equivalent to about half the width.

D

E

Text-figure 12. Type material of Paradamesops jimaensis Yang in Lu et al., 1974a, the holotype cranidium and the paratype pygidium are now regarded as separate species of Parablackwelderia. A-D, Parablackwelderia jimaensis(Yang in Lu et al., 1974a), holotype cranidium (original of Lu et al., 1974a, pl. 1, figs. 15a, b; CUGB 1211301) in dorsal, anterolateral, and oblique-anterior views, and partial enlargement showing surface features, from the Huaqiao Formation at Jimachong, Yeping, eastern Guizhou; A-C, × 2.6, D, × 8; E, F, Parablackwelderia yangi sp. nov. (original of Lu et al., 1974a, pl. 1, fig. 16; CUGB 1430201), holotype, designated herein, partial enlargement and dorsal view, showing the surface granulation, x 5.5, x 9.5, from the Huaqiao Formation at Jiudiantang, Xinhuang, western Hunan. Remarks. Yang's (1978) type material of the species is refigured here (Text-fig. 12). Prior to being published by Yang, it was figured twice (Lu et al., 1974a; Yin and Li, 1978) as Paradamesops jimaensis, the type species of Paradamesops Yang. Paradamesops is now regarded as a junior synonym of Parablackwelderia. The type material includes a single cranidium that was assigned as the holotype, and a single pygidium. They are quite different in terms of the pattern of surface prosopon. The holotype cranidium, collected from Jimachong, Yuping, eastern Guizhou, has extremely fine and closely spaced granules that can be seen only under high magnification, but the paratype pygidium is strongly granulated. In addition to the fine granules the holotype bears also dense anastomosing ridges on the fixigenae. The paratype pygidium, however, bears notably coarser and scattered granules. The large new material of Parablackwelderia jimaensis from the Huaqiao Formation at Paibi and Wangcun, northwestern Hunan, includes cranidia and pygidia in association, and shows that the pygidium of this species bears no ornamentation. It either lacks surface prosopon or has extremely fine granules and anastomosing ridges similar to that observed •

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on the cranidium, suggesting that the paratype pygidium should be excluded from Parablackwelderia jimaensis. The paratype pygidium appears to represent a new granulated species of Parablackwelderia (Parablackwelderia yangi sp. nov.). P. jimaensis is characterized by a moderately tapered glabella with an obtusely rounded front and faintly defined bacculae, anteriorly located palpebral lobes, a relatively narrow (tr.) anterior cranidial area and strongly convergent anterior branches of the facial suture. The cranidia in the new material are identical in all major respects with the holotype cranidium, but show some morphological variation in the nature of the glabellar front and the width of the palpebral area of the fixigena. The associated pygidia are also variable in outline and in location of the posterolateral spines. The shape varies from transverse-subelliptical in outline with widely spaced posterolateral spines to subtriangular in outline with length subequal to width and with closely spaced posterolateral spines. Morphological variation in the new material enables synonymization of many species assigned previously to Paradamesops as junior synonyms of Parablackwelderia jimaensis. The new material from Hunan includes librigenae, hypostomes and juvenile cranidia, none of which were known previously for the species. The librigena has a narrow (tr.), gently convex, strongly anastomosing genal field with an anterior-inner portion elevated adaxially to fit the palpebral stalk, and with a flat lateral border widening backward to form a broad genal angle from which extends a stout genal spine. The border bears a shallow furrow posteriorly; it is parallel to the lateral margin of the librigena, and extends to the proximal part of the genal spine. The hypostome (P1. 30, fig. 11) bears wide, wing-like lateral borders having a posterior portion that projects outward and gently forward. The middle body has a subtriangular anterior lobe with an obtusely pointed posterior end; a middle furrow that is deeply incised laterally, shallowing medially, and a crescentic, convex posterior lobe. The hypostome is much different in morphology from that assigned by Opik (1967, pl. 44, fig. 9) to Meringaspis meringaspis, although the hypostome also has broad posterolateral borders. 0pik's (1967) hypostome from Queensland is closely comparable to those assigned here to Paradamesella. Juvenile cranidia have long, gently forward-tapering glabellas with deeply incised S1, $2, and $3 furrows; strong eye ridges; wide (sag.), gently upturned anterior borders, and forwardly divergent anterior branches of the facial suture. All following nine species are considered to be junior synonyms of Paradamesops jimaensis: Meringaspis sp. A (Opik, 1967, p. 326, pl. 46, figs. 2-5) from Queensland, Australia; Meringaspis karatauensis Ergaliev, 1980 (p. 149, 150, pl. 3, fig. 15; pl. 6, figs. 9-12) from Malyi Karatau, Kazakhstan; Paradamesops kuruktagensis Zhang, 1981 (p. 178, pl. 65, fig. 3) from the middle Cambrian Mohershan Formation of Kuruktag, Xinjiang; Paradamesops depressus Qian in Qiu et al., 1983 (p. 180, pl. 61, fig. 1) from Jingxian, southern Anhui; Paradamesops trigonatus Qian in Qiu et al., 1983 (p. 180, pl. 60, figs. 9, 10), from Qingyang, southern Anhui; Paradamesops reticulatus Qiu in Qiu et al., 1983 (p. 181, pl. 60, fig. 11), also from Jingxian, southern Anhui; Paradamesops flatilimbatus Qiu in Qiu et al., 1983 (p. 180, pl. 60, fig. 7) from Guichi, southern Anhui; Paradamesops lirellatus Qiu in Qiu et al., 1983 (p. 180, 181, pl. 60, fig. 8), from Guichi, southern Anhui; and Paradamesops latilimbatus Qian in Qiu et al., 1983 (p. 180, pl. 59, figs. 3, 4) from Guichi, southern Anhui.

Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Jimachong, Yeping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Pianaspis sinensis Zone though the lower part of the Liostracina bella Zone (equivalent to the Lejopyge laevigata Zone through the lower part of the Linguagnostus reconditus Zone).

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Parablackwelderia laterilobata (Yang in Zhou et al., 1977)

Plate 31, figures 1-5 1977 1978 1978 1980 ?1987 ?1987 2001b

Paradamesops laterilobatus Yang in Zhou et al., p. 198, pl. 59, figs. 1, 2. Paradamesops laterilobatus Yang; Yin and Li, p. 512, pl. 170, figs. 5, 6. Paradamesops laterilobatus Yang, p. 59, pl. 13, figs. 3-5. Paradamesops laterilobatus Yang; Liu, p. 321, pl. 221, figs. 7, 15, 20. Hwangjuella chonjuriensis Kim, p. 40, 41, pl. 22, figs. 1, 10, 11, ?2-4, ?9. Paradamesops sp., Kim, pl. 22, figs. 5, 7, 8, 14, ?2 (figured. only). Meringaspis laterilobatus (Yang), Peng, Babcock, and Lin, p. 102, pl. 6, figs. 3, 4.

Lectotype. Cranidium (Yang in Zhou et al., 1977, pl. 59, fig. 1; refigured in Yin and Li, 1978, pl. 170, fig. 5, Yang, 1978, pl. 13, fig. 3, Liu, 1982, pl. 221, fig. 15; CUGB 0110001) from the upper part of the Lisania tungjenensis Zone, Huaqiao Formation, at Huaqiao, Baojing, northwestern Hunan; designated subsequently as holotype of the species by Yang (1978). New material. Eight cranidia, and one pygidium (illustrated specimens NIGP 137582-137585) in collections P 171.5, P200.7, P240.5, and W 132.5. Emended diagnosis. Parablackwelderia with strong, transverse eye ridge having length greater than glabellar width at $2; bacculae large and prominent, separated or nearly separated from L1 lobes; pygidium with pleural furrows shallow or gently impressed; pygidial axis occupying two-thirds of pygidial length; posterior margin rounded. Remarks. New material from the Huaqiao Formation of northwestern Hunan agrees in all observed details with the type material from the same region. This species is most closely related to P. jimaensis, differing pricipally in having a pair of well defined bacculae. Furthermore, it can be differentiated by having a wider (tr.) anterior border, a wider (tr.) eye ridge, and more widely spaced palpebral lobes. Key features of this species include a cranidium having large, prominent bacculae, a broad (tr.) anterior cranidial area and palpebral area of the fixigena, a pygidium having a relatively less segmented axis, and a relatively effaced pleural field with weak ribs in the posterior portion. Hwangjuella chonjuriensis Kim, 1987 and Paradamesops sp. (Kim, 1987) from North Korea no doubt belong to Parablackwelderia. Based on Kim's (1987) poor illustrations, both species are probably synonymous with P. laterilobata. Until more is known about these two Korean species, they are questionably assigned to P. laterilobata. Occurrence. The holotype is from the upper part of the Lisania tungjenensis Zone, Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Pianaspis sinensis Zone though the lowermost part of the Wanshania wanshanensis Zone (equivalent to the Lejopyge laevigata Zone).

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Parablackwelderia sp. cf. P. huabeiensis (Zhang in Qiu et al., 1983)

Plate 31, figures 8-12 cf. 1983 Paradamesops huabeiensis Zhang in Qiu et al., p. 181, pl. 59, figs. 5a, 5b. 2001b Meringaspis sp., Peng, Babcock, and Lin, p. 102, pl. 6, figs. 1, 2. Material. Three cranidia and one pygidium (illustrated specimens NIGP 137588-137590) in collections P 184 and P190.7. Description. Cranidium trapezoidal, length half width. Glabella broadly conical, obtusely rounded anteriorly, with only two pairs of lateral furrows; S 1 straight and long, deeply incised and strongly oblique rearward, defining triangular L 1; $2 short, nearly transverse, with outer end deepened into pit; L1 with weak oblique furrow paralleling S1 and defining subtriangular baccula. Occipital furrow wide (sag., exs.), moderately impressed, deep laterally; occipital ring narrow, gently curving forward laterally, with transverse ridge medially. Preglabellar field absent; anterior border narrow (sag.) and upturned. Eye ridge double, belt-like, gently oblique rearward. Palpebral lobe slender stalk, extending outward and upward with base just in front of $2. Anterior branch of facial suture gently convergent forward and across anterior border furrow, then turning strongly inward to cut anterior border inward and forward; posterior branch nearly straight, diagonally directed, enclosing broad (exs.) triangular posterolateral projection of fixigena. Posterior border furrow transverse, shallow and wide (exs.), posterior border a narrow ridge. Pygidium wider than long, with slender, slightly rearward tapering axis beating at least six tings. Borders flat and relatively wide, defined by moderately impressed border furrows, bearing six pairs of border spines. Glabella covered with fine and dense granules; fixigena with fine, anastomosing caecal ridges; pygidium with fine, moderately dense granules. Remarks. Paradamesops huabeiensis is based on an incomplete cranidium from the Changhia Formation in the Jiawang region, northern Jiangsu; its exact locality is unknown. Although poorly preserved, the cranidium shows some distinct features that differentiates it from other species of Parablackwelderia. These include oblique eye ridges, a flat and upturned anterior border, and no glabellar bacculae. P. huabeiensis is closely similar to Parablackwelderia merigaspis (Opik) from Queensland, Australia, but P. meringaspis is distinguished by having a narrower palpebral area of the fixigena, a more oblique eye ridge, and a shorter (exs.) but wider (tr.) posterolateral projection of the fixigena. New cranidia from Hunan are left in open nomenclature because the holotype of Paradamesops huabeiensis, with which they are compared, is poorly preserved and poorly illustrated. Cranidia from the Huaqiao Formation, northwestern Hunan, are about twice as large as the holotype cranidium. The Hunan material is similar to the holotype in having a similar brow/d-conical glabella, and a similar narrow and upturned anterior border. Both the new cranidia and the holotype cranidium have obliquely directed eye ridges. However, the eye ridge in the holotype is less oblique than the new cranidia and it seems to be more posteriorly located, so that the holotype differs in having a more posteriorly located palpebral lobe (or the base of palpebral lobe). The inner end of the eye ridge in the holotype is situated far posterior of the glabellar front, but is more anteriorly located in the new cranidia. (i.e., it is connected with the anterior portion of the frontal lobe). The baccula seems to be absent from the holotype. It is uncertain if the differences mentioned above • 109.

have specific value or just represent intraspecific variation. It is also uncertain whether the holotype bears the thin and long palpebral stalks as seen in the present cranidia. Until more is known about Paradamesops huabeiensis, and a closer comparison is possible, the new material is assigned only with question to that species. The Hunan specimens also resemble Parablackwelderia spectabilis, the type species. The glabellar shape, the direction of the lateral glabellar furrows, the shape of the preglabellar field, the course of facial sutures, and the presence of palpebral stalks are all comparable. In cranidial respects, differences bel~ween the Hunan specimens and P. spectabilis are minimal. The new specimens have more tapered and less effaced glabellas, stronger eye ridges, less thick palpebral stalks, and more divergent posterior branches of the facial suture. In pygidial respects, the new specimens differ in having a complete rather than spinose posterior margin between the posterolateral spines. Pygidia assigned to this species have wide and flat lateral borders, which are clearly defined by border furrows. Such a character distinguishes the present species from those species with rounded posterior margins in Parablackwelderia, in which the lateral borders are much narrower, and the posterolateral spines are not derived from border but from pleurae.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the Pianaspis sinensis Zone though (equivalent to the Lejopyge laevigata Zone). Parablackwelderia yangi sp. nov. Text-figure 12E, F 1974a Paradamesopsjimaemsis Yang in Lu et al. (in part), p. 86, 87, pl. 1, fig. 16 only. 1978 Paradamesopsjimaemsis Yang (in part); Yin and Li, 1978, p. 511, pl. 169, fig. 10 only. 1978 Paradamesopsjimaemsis Yang (in part); Yang, p. 58, pl. 13, fig. 2 only.

Etymology. The name honors Yang Jialu from China University of Geology (Wuhan). Holotype. Testaceous pygidium (Yang in Lu et al., 1974a, pl. 1, fig. 16; refigured in Yin and Li, 1978, pl. 169, fig. 10; Yang, 1978, pl. 13, fig. 2; CUGB 1430201). Diagnosis. Parablackwelderia with rounded posterior pygidial margin; axis with eight tings and short terminal piece; lateral and posterior border narrow; surface covered with granules. Remarks. New Parablackwelderia material from Hunan demonstrates that the pygidium assigned to Parablackwelderia jimaensis as a paratype (Yang in Lu et al., 1974a) is not conspecific with the holotype cranidium. The pygidium is now designated as the holotype of a new species, Parablackwelderia yangi. The coarse and scattered granulation present on the holotype differentiate this species from most others assigned to Parablackwelderia that have a complete posterior margin. P. sp. cf. P. huabeiensis has a similar granulation. However, P. sp. cf. P. huabeiensis is differentiated by having a proportionally wider outline, broader lateral and posterior borders, and a clearly defined, less segmented pleural field bearing narrower pleural furrows. Occurrence. From the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] • 110.

tuma Zone, Huaqiao Formation, Jiudiantang, Xinhuang, western Hunan. Parablackwelderia sp. Plate 31, figures 6, 7; Text-figure 13 2001b

Meringaspis sp. 1, Peng, Babcock, and Lin, p. 102, pl. 5, figs. 1, 2.

Material. One incomplete cranidium and one incomplete pygidium (NIGP 137586, 137587) in

collection P 156.

Text-figure 13. Parablackwelderia sp. A, composite photo from the incomplete cranidium figured on P1.31, fig. 7, the right side is a mirror image of the left side; B, partial enlargement of right posterolateral glabella, showing granules and baccula, × 8.

Description. Cranidium with anterior margin arched forward slightly. Glabella large, tapered

moderately forward, obtusely rounded anteriorly with baccula expanded outward and not completely isolated from glabella; S1 furrow deeply pitted anteriorly, shallowing adaxially. Occipital ring with a centrally located node and pair of weak, transverse depressions at sides; occipital furrow clearly defined. Eye ridge weak, slightly oblique rearward. Facial suture and posterolateral projection unknown. Pygidium subtriangular in outline, with broad-based, closely spaced posterolateral spines. Pleural field about as wide as, or slightly narrower than, axis, with interpleural furrows nearly effaced, shallow but clearly impressed pleural furrows that define five fibs. Lateral and posterior borders wide and flat, bearing tiny border spines, of which only the third and the fourth spines on fight lateral border are preserved. Surface of glabella, occipital ring, fixigenae, pygidial border and posterolateral spines covered with dense, fine granules; surface of anterior border and pygidial pleural field smooth. Remarks. This species is known from only an incomplete cranidium and a broken pygidium. So far, it is the oldest-known species of Parablackwelderia from the northwestern Hunan-eastern Guizhou

region. Morphologically, the cranidium of this species is characterized by having a glabella with long and curved S 1 furrows, small bacculae that expand at the bases of the L1 lobes, a narrow and strongly upturned anterior border, weak and nearly transverse eye ridges, and dense fine granules. The pygidium has a wide and flat lateral border, tiny lateral border spines, and stout posterolateral border spines. • 111.

The present species is most similar to Parablackwelderia sp. cf. P. huabeiensis described above, but differs in having a curved rather than straight S 1, a weaker eye ridge, a narrower (sag., exs.) anterior border, broader lateral pygidial border, and more broadly based lateral border spines. In P. cf. P. huabeiensis, the baccula is poorly developed, and less expanded outward as that in the present cranidium; the occipital furrow is broader and deepened at the sides, and the occipital ring is narrower (sag.), beating no additional transverse impressions. An incomplete cranidium assigned to Menocephalius belenus by Walcott (1905, p. 62) from the Changhia Formation in Zhangxia, Shandong, is also comparable to P. sp. It was later referred to Lisania by Walcott (1913, p. 166, pl. 15, fig. 16), and more recently was transferred questionably to Damesops by Zhang and Jell (1987, p. 217, pl. 102, fig. 10). The specimen is here questionably transferred to Parablackwelderia. It differs from the present cranidium in having a more effaced, non-granulose glabella, and probably a raised, rather than flat and upturned anterior cranidial border. The present cranidium and pygidium may represent an unnamed species of Parablackwelderia, which is oldest in age among the parablackweldiids in the northwestern Hunan and eastern Guizhou region. However, as both are unsatisfactorily preserved, open nomenclature is used until better material is available.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Genus

PROTAITZEHOIA

Yang 1978 in Yin and Li, 1978

Protaitzehoia Yang in Yin and Li, 1978, p. 512; Yang, 1978, p. 59, 60; Peng, 1987, p. 100; Yuan and Yin, 2001, p. 344, 345" Peng, Babcock, Lin, and Lin, 2003b, p. 473-477. ? Shanchengziella Lu and Qian, 1983a, p. 245,252, 253" Qian, 1994, p. 56. Type species. Protaitzehoia yuepingensis Yang in Yin and Li, 1978, p. 512, pl. 170, fig. 3, from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [-Torifera] tuma Zone, Huaqiao Formation in eastern Guizhou; by original designation. Emended diagnosis. Cephalon transversely subelliptical with obtuse genal angle and genal spines forward of poserolateral comer. Glabella subrectangular with sides convergent, subparallel, or constricted in anterior half, obtusely rounded with weak sagittal notch anteriorly; four pairs of lateral furrows with incised, weakly bifurcated S 1. Palpebral lobe relatively small, located at or slightly posterior to cranidial midlength; eye ridge thick, weakly to clearly defined. Anterior border moderately thick, upturned and widened laterally; preglabellar field absent. Anterior branch of facial suture straight, gently to strong convergent forward; posterior branch straight and diverging strongly rearward. Pygidium semicircular or transverse-subrectangular in outline, axis variably conical, with four to eight tings and a short terminal piece; with five to six pairs of short border spines, or with one to three pairs of reduced spines anteriorly, or without spines; pleural field with relatively flat inner portion and down-sloping, concave outer portion; pleura with wide anterior band and linear posterior band. Surface variably granulose. Other species. Damesella quadrata Resser and Endo, 1937 (p. 203, pl. 49, fig. 32), from the Kushan Formation, near Dapingzhuang, Changxingdao Island, southern Liaoning, Northeast China; • 112.

Protaitzehoia granifera Yang in Yin and Li, 1978 (p. 512, 513, pl. 170, fig. 3), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Jimachong, Yeping, eastem Guizhou; Protaitzehoia subquadrata Peng, 1987 (p. 101, pl. 9, fig. 7), from the Formosagnostus formosus [=Hadragnostus modestus]-Distazeris [=Paradistazeris] Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestemn Hunan; Zhalongtania valleculata Zhou in Zhou et al., 1996 (p. 41, pl. 4, figs. 13-15) from beds equivalent to the Proagnostus bulbus Zone, Nidanshan, Hualong, Qinghai, Northwest China; Protaitzehoia spinifera sp. nov., and Protaitzehoia sp. (described below). Remarks. Yuan and Yin (2001) recently revised the genetic diagnosis by adding the pygidial characters. This genetic diagnosis is emended here to emphasize the presence or absence of pygidial spines in the genus. When the genus was erected, only cranidia were available. It was classified within the Damesellidae by Yang (1978) because it has close similarity in cranidial characters to genera assigned to that family. This assignment was widely accepted by most subsequent authors (Yin and Li, 1978, p. 512; Peng, 1987, p. 100; Zhang and Jell, 1987, p. 208). Yuan and Yin (2001, p. 344), however, recently transferred the genus to the Ordosiidae. Such a transfer was based mainly on the absence of pygidial spines in Protaitzehoia granifera from the Huaqiao Formation, eastern Guizhou. Such a transfer was rejected by Peng et al. (2003), as their material from northwestern Hunan does bear reduced pygidial spines. Peng et al. (2003) demonstrated that some other species of Protaitzehoia bear either reduced or "normal" pygidial spines. Protaitzehoia spinifera bears pleural spines through much of the meraspid and holaspid periods (Peng et al., 2003), and those spines are closely comparable in nature with those of other damesellids. Besides the presence of pygidial spines, the pygidium of Protaitzehoia is unlike that of ordosiids in the subdivision of pleurae. In Protaitzehoia, the pleura is divided into a wide (exs.) anterior band, and a linear posterior one, whereas ordosiids, such as Ordosia, Taitzuia, and Poshania, have reversed pleurae with a linear anterior band and a wide (exs.) posterior band (see Lu et al., 1965, pl. 38, figs. 9, 19; pl. 39, figs. 8, 11, 16). The cranidial morphology of Protaitzehoia is more consistent with that of damesellids than ordosiids. A complete cephalon present in new material from the Huaqiao Formation of northwestern Hunan (P1. 33, fig. 11) displays a clear dameselloid aspect including a well furrowed glabella, an upturned anterior cranidial border, and genal spines in front of the posterolateral comer. Meanwhile, several early meraspid cranidia (P1. 33, figs. 1-3) are reminiscent of the damesellid genus Parablackwelderia. Key differences in the cranidium between the ordosiids (Ordosia, Taizuia, Poshania, and Parataizeia) and damesellids are that the ordosiids have effaced rather than well-furrowed glabellas, swollen anterior cranidial borders, and a tripartite cranidial border furrow. Protaizehoia shares none of the diagnositic features with ordosiids. Shanchengziella Lu and Qian is probably a junior synonym of Protaitzehoia. Shanchengziella is monotypic, and its type species, Shanchengziella elongata is known only from the holotype. The holotype cranidium (Lu and Qian, 1983a, pl. 2, fig. 2) of S. elongata is relatively small in size, and characterized by having a forward-tapering glabella, an upturned anterior border, a thick eye ridge, small palpebral lobes lying at the cranidial midlength, and by lacking a preglabellar field. It is identical in almost all respects with cranidia of similar size in the ontogenetic series of Protaitzehoia subquadrata (P1. 33, fig. 6), suggesting that the genus may be based on a juvenile of Protaitzehoia. Lu and Qian (1983a) and Qian (1994) considered Shanchengziella to be comparable with Paokania, a lower Cambrian genus characterized by possessing transglabellar furrows, and classified them within the same family. We consider such an association unlikely. Qian's (1994) subsequent assignment of two Paokania-like pygidia to S. elongata also seems problematic. Until more is known for Shanchengziella, it is tentatively placed in synonymy with Protaitzehoia. • 113.

Stratigraphically, Shanchengziella occurs in the upper Cambrian Chuangia Zone of the Changshan Formation in Bengxi, Liaoning. It is slightly younger in age than Protaitzehoia, which ranges upward through the Proagnostus bulbus Zone into the overlying Linguagnostus reconditus Zone, and probably into the succeeding Glyptagnostus stolidotus Zone. Zhalangtania Zhou, which has Zhalangtania valleculata Zhou (in Zhou et al., 1996, p. 42, pl. 4, figs. 13-15) as its type species, bears close resemblance to Protaitzehoia. However, this genus differs from Protaitzehoia in having a largely effaced glabella with faint lateral furrows, obscure eye ridges, and a narrow (tr., sag.), flat and slightly upturned anterior border.

Protaitzehoia yuepingensis Yang in Yin and Li, 1978 Plate 32, figures 8-13; Text-figure 14 1978 1978 2001 2001b 2003b

Protaitzehoiayuepingensis Yang in Yin and Li, p. 512, pl. 170, fig. 4. Protaitzehoiayuepingensis Yang; Yang. p. 60, pl. 13, fig. 7. Protaitzehoianitida Yuan and Yin (in part), p. 347. 348, pl. 1, fig. 2 only. Protaitzehoiayuepingensis Yang; Peng, Babcock. Lin, p. 104, pl. 9, figs. 7, 8. Protaitzehoiayuepingensis Yang; Peng. Babcock. Lin, and Lin, p. 479, pl. 1, figs. 4-6.

Holotype. By monotypy; mostly exfoliated cranidium (Yang in Yin and Li, 1978, pl. 170, fig. 4; refigured by Yang, 1978, pl. 13, fig. 7; Text-fig. 14 herein; CUGB 1211304), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou. New material. Three cranidia and two pygidia (illustrated specimens NIGP137596-137599) in collections W 196.3, W 197.55, and P337.5.

Text-figure 14. Holotype cranidium of Protaitzehoia yuepingensis Yang in Yin and Li, 1978 (pl. 170, fig. 4; also Yang, 1978, pl. 13, fig. 7, CUGB 1211304). A-C, in anterior, anterolateral, and dorsal views, all × 6; D, E, partial enlargement of the anterolateral and posterolateral glabella. The enlarged areas are indicated by rectangles on figure C, both × 12. •

114.

Emended diagnosis. Protaitzehoia with anterior cranidial border narrow and raised; glabella short, with length equal to or slightly greater than maximum width at L1; sides of glabella straight or slightly concave in anterior half. Eye ridge and eye well defined; palpebral area of fixigena narrow. Pygidium semicircular, with wide axis and obscure postaxial ridge. Surfaces covered with fine, dense, granules of variable size. Remarks. This species was based on a single cranidium that was first illustrated by Yin and Li (March, 1978) and later by Yang (August, 1978). The concept of the species has been unclear as the holotype cranidium, from the Huaqiao Formation of eastern Guizhou, was never satisfactorily illustrated previously. The holotype is refigured here (Text-fig. 14) to help clarify the concept of the species and the genus. Key characters of the species include a glabella that is relatively short and wide; moderately tapered, narrow (tr.) palpebral and preocular areas of the fixigena; a thick, well raised eye ridge; a thick, gently curved palpebral lobe; and a narrow (sag.) and strongly raised anterior border. The holotype cranidium bears fine granules that are uniform in size. They are most obvious on the fixigenae. Cranidia in the new material are about the same size as the holotype, and agree in all essential respects with the holotype cranidium, which comes from the same general region. One cranidium shows that the granulation varies somewhat in size (P1. 32, figs. 8, 9) in the species. The specimen bears granules that are slightly coarser than those observed on the holotype and on another cranidium in the new material. Pygidia associated with the cranidium from Hunan are the first recorded for the species (P1.32, figs. 11, 13). The pygidium has a semicircular outline, and a conical axis with five tings and a short, ill-defined terminal piece, and a complete posterior margin. The pleural field is weakly furrowed and comprises a flat inner portion and a concave outer portion. The pleural and interpleural furrows are transverse on the inner portion, deflected sharply rearward and then merging as broad and shallow impressions that are posterolaterally directed on the outer portion and not connected to the lateral and posterior margins. Granulation on the pygidium seems to be variable. One pygidium (P1. 32, fig. 13) bears coarser, unevenly sized and widely spaced granules, whereas fine granules seem to be more common for the species. The variably granulose ornamentation also occurs in other species of Protaitzehoia. In P. granifera, granulation seems to vary with stratigraphic occurrence, becoming weaker in specimens having higher stratigraphic occurrences. Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the Linguagnostus reconditus Zone). Protaitzehoia granifera Yang in Yin and Li, 1978

Plate 32, figures 1-7; Text-figure 15 1963 1965 1978

Blackwelderia sp., Egorova, Xiang, Li, Nan, and Guo (in part), p. 40, pl. 8, fig. 4 only; non fig. 3 [-- Parablackwelderia jimaensis (Yang in Lu et al., 1974a)]. Blackwelderia sp., Lu, Zhang, Zhu, Qian, and Xiang (in part), p. 383, 384, pl. 72, fig. 4 only; non fig. 3 [= Parablackwelderia jimaensis (Yang in Lu et al., 1974a) ]. Protaitzehoia granifera Yang in Yin and Li, p. 512, 513, pl. 170, fig. 3. • 115.

1978 1987 1987 2001 2001

2001 2003b

Protaitzehoiagranifera Yang; Yang, 1978, p. 60, pl. 13, fig. 8. Protaitzehoiagranifera Yang; Peng, p. 100, 101, pl. 9, fig. 4. Protaitzehoiasp. Peng, p. 100, pl. 9, fig. 8. Protaitzehoialatilimbata Yuan and Yin, p. 346, 347, pl. 1, figs. 5-8. Protaitzehoianitida Yuan and Yin (in part), p. 347, 348, pl. 1, figs. 4, 9, 15, ?10, ?16 [probably belongs to Baikadamaspis]; non fig. 2 [= Protaitzehoia yuepingensis Yang in Yin and Li, 1978], non fig. 11 [= Protaitzehoia subquadrata Peng, 1987]. Protaitzehoiagranifera Yang; Yuan and Yin, p. 348, pl. 1, figs. 12-14. Protaitzehoia granifera Yang; Peng, Babcock, Lin, and Lin, p. 479, pl. 1, figs. 1-3, 7, 8, 10-13, ?9.

Holotype. By monotypy; cranidium (Yang in Yin and Li, 1978, pl. 170, fig. 3; refigured in Yang, 1978, pl. 13, fig. 8; Text-fig. 15 herein, CUGB 1207208) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [= Torifera] tuma Zone of the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou. New material. About 12 sclerites, including cranidia and pygidia (illustrated specimens NIGP 137591-137595), in collections P282.75, P307.4, P319.6, P346.7, P348, and W227. Emended diagnosis. Protaitzehoia with anterior cranidial border that is moderately wide, gently convex and deflected; palpebral area of fixigena more than half as wide as glabella at L 1; eye ridge weak. Pygidium with slender axis and postaxial ridge reaching posterior margin. Surface with granules of bimodal size.

C

Text-figure 15. Holotype cranidium of Protaitzehoia granifera Yang in Yin and Li, 1978 (pl. 170, fig. 3; also Yang, 1978, pl. 13, fig. 8, CUGB 1207207) in anterolateral and dorsal views (A, B), x 8, x 8, and partial enlargement of glabella showing two sets of granules (C), x 12.

Description. Cranidium with anterior border moderately wide (sag.), gently convex, gently deflected; glabella constricted in anterior one-third, with sides concave and front transverse or • 116.

weakly notched. Eye ridge weak; palpebral area wider than half of glabellar width at L 1. Pygidial axis slender with seven to eight tings and small terminal piece with postaxial ridge reaching to posterior margin; each ring with pair of nodes, forming two longitudinal rows of nodes on axis. Pleural field with flat inner portion and concave outer portion; pleural furrows relatively short, sigmoidal, clearly defined; interpleural furrows long, shallow, abaxially broadened and rearward curving; anterior pleural band wide (exs.) and moderately convex, posterior band short, linear, horizontally directed. Pleural field with scattered granules and strong wrinkles.

Remarks. This species was originally based on a single cranidium (Yang in Yin and Li, 1978). The holotype cranidium, which is refigured here to clarify the species concept, is characterized by having a relatively rather narrow (tr.) glabella that is strikingly constricted in its anterior one-third, weak eye ridges, relatively wide fixigenae, a relatively wide (sag.) and moderately upturned anterior border, and strongly convergent anterior branches of the facial suture. Ornamentation of the cranidium consists of two sets of granules, fine, closely spaced granules and coarse scattered granules. These features serve to distinguish P. granifera from P. yuepingensis. The new material from the Huaqiao Formation, northwestern Hunan, resembles the holotype cranidium in all observed respects except for having weaker granulate ornamentation. However, such a minor difference should be of no more than intraspecific variation because similar weak granulation is also known from additional material collected from the type locality of P. granifera (Yuan and Yin, 2001, pl. 1, figs. 5, 6; Peng et al., 2003b, pl. 1, figs. 10-13). The pygidium of this species is comparable to that of P. yuepingensis in general respects but differs in having a more slender, less rearward-taped, and more segmented axis, more deeply furrowed pleural fields, and a concave rather than convex posterior margin. Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the Linguagnostus reconditus Zone). Protaitzehoia subquadrata Peng, 1987 Plate 33, figures 1-11; Plate 34, figures 1-7; Text-figure 16 1987 1987 2001 2001 2001 2001 2001b 2001d 2003b

Protaitzehoia subquadrata Peng, p. 101, pl. 9, fig. 7. Protaitzehoia intermedia Peng, p. 101,102, pl. 9, figs. 5, 6. Protaitzehoia quadrata jimachongensis Yuan and Yin, p. 345,346, pl. 1, fig. 3, ?fig. 1. Protaitzehoia nitida Yuan and Yin (in part), p. 347, 348, pl. 1, fig. 11 only. Protaitzehoia subquadrata Peng; Yuan and Yin, p. 349, pl. 1, fig. 17. Protaitzehoia intermedia Peng; Yuan and Yin, p. 349, pl. 1, fig. 18. Protaitzehoia subquadrata Peng; Peng, Babcock, and Lin, p. 104, pl. 12, figs. 3, 4. Protaitzehoia subquadrata Peng; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.9. Protaitzehoia subquadrata Peng; Peng, Babcock, Lin, and Lin, p. 475,479, pl. 1, figs. 14, 15; text-fig. 2A-H.

Holotype. Cranidium (Peng, 1987, pl. 9, fig. 7; Text-fig. 16 herein; NIGP 74548) from the Proagnostus bulbus Zone of the Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan. • 117-

New material. More than 50 sclerites including, cranidia, librigenae, pygidia, and one cephalon (illustrated specimens NIGP 137600-137614), from P277, P298.4, P298.54, P317.4, P319.8, P325.7, P326.9, P327.4, P331.8, W199.2, W200.7, W210.5, and W212.45.

Text-figure 16. Holotype cranidium of Protaitzehoia subquadrata Peng, 1987 (pl. 9, fig. 7; NIGP 74548) in anterolateral and dorsal views, × 5.5.

Emended diagnosis. Protaitzehoia with anterior cranidial border narrow, notably upturned; glabella relatively wide (tr.), palpebral area of fixigena subequal to or less than half as wide as glabella at L 1; eye ridge clearly defined. Pygidium with narrow and relatively short axis with postaxial ridge reaching to paradoublural line; anterior one or two segments with spinelike, rearward projection. Surface covered with granules of bimodal size. Remarks. New material from the Huaqiao Formation, northwestern Hunan, is closely similar to the type material, which is also from northwestern Hunan. Characters that warrant assignment of the new material to this species include a relatively wide cranidium; a wide, short glabella that is weakly constricted in the middle; a clearly defined eye ridge, a strongly upward curving anterior border, relatively narrow palpebral areas of the fixigenae, and subparallel or gently convergent anterior branches of the facial sutures. P. subquadrata is closely similar to P. yuepinensis, the type species of the genus, in having a thick eye ridge, relatively less convergent anterior branches of the facial sutures, a relatively broad glabella, and a relatively narrow fixigena. However, the anterior border in P. yuepingensis is gently raised and deflected rather than strongly curved upward. It is further differentiated in lacking the coarse granules. P. subquadrata is similar to P. granifera in having granulose ornamentation, but differs in having a wider glabella, distinct rather than obscure eye ridges, and in having less convergent anterior branches of the facial sutures. 118. •

Small cranidia (P1. 33, figs. 1, 2) in the new material probably represent early meraspid stages. They are characterized by having a strongly tapered glabella, a transverse eye ridge, an upward-deflected anterior border, and relatively broad (exs.) posterolateral projections of the fixigena. These features are reminiscent of some species assigned here to Parablackwelderia, suggesting that Protaitzehoia and Parablackwelderia share a close affinity, and should be classified in the same family. This species is characterized by a subtrapezoidal pygidium with subparallel lateral margins and a tiny spine on the first one or two pleural segments. The pygidium of P. subquadrata most closely resembles that of P. granifera. It is distinct from P. granifera, however, in having a shorter, less segmented, more strongly tapering axis; having a wrinkled pleural field; and having more prominent spinelike projections from the anterolateral margin. The pygidium varies through ontogeny in the shape of axis, the pleural fields, and the posterior margin. The axis becomes narrower through ontogeny. The inner portion of the pleural field becomes progressively more wrinkled through ontogeny. The posterior margin varies through ontogeny from entire and rounded to sinuous with an anterior curvature medially.

Occurrence. The holotype is from the Proagnostus bulbus Zone of the Huaqiao Formation at Wa'ergang, Taoyuan, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the Linguagnostus reconditus Zone). Protaitzehoia spinifera sp. nov. Plate 34, figures 8-17 2001b Protaitzehoia sp., Peng, Babcock, and Lin, p. 104, pl. 12, figs. 5, 6. 2003b Protaitzehoia sp., Peng, Babcock, Lin, and Lin, p. 475, text-fig. 2I-L.

Etymology. From Latin spinifer, spine-beating, in reference to the presence of pygidial spines. Holotype. Pygidium (P1.34, fig. 16, NIGP 137621) in collection W221.5. Paratypes. Four cranidia, and three pygidia (NIGP 137615-137620) in collections P337.5, P341.8, W200.3, W211.7, W221.5, and W225. Diagnosis. Protaitzehoia with coarse, subequally-sized granules on cranidium and pygidium; glabella weakly furrowed. Pygidium with relatively wide, subconical axis and five pairs of pleural spines. Description. Anterior border strongly upturned, preglabellar field absent. Glabella gently tapering forward, evenly rounded anteriorly, straight along sides or slightly constricted adjacent to $3, with indentation at anterior margin; S 1 incised, gently oblique rearward; $2 shallow and short, directed inward or parallel to S1; $3 faint and isolated from axial furrow; $4 faint. Fixigena one-third to two-fifths of glabellar width at S 1; palpebral eye ridge wide, gently raised. Anterior branches of facial suture convergent forward at an angle of 70 ° to the sagittal line; posterior branches almost transverse. • 119.

Pygidium subrectagular; length twice width; with straight, slightly expanding lateral margin and five pairs of short, pleural spines. Axis conical, occupying about three-fourths of pygidial length, with four tings and subtriangular terminal piece. Pleural field with flat inner portion and outward-sloping outer portion, bearing four pleurae; pleura transverse initially, deflected strongly rearward distally; with convex, anterior band that widens abaxially, and deep pleural furrows at anterior three segments, and short (tr.), linear posterior band defined by faint interpleural furrow. Surface covered with relatively coarse granules of subequal size. Each axial ring in pygidium with pair of relatively large granules, forming two longitudinal lines of granules on axis. Remarks. The cranidium has strong and coarse granules, a glabella with shallow lateral furrows and a slightly oblique S 1. The pygidium has distinct marginal spines. These features serve to distinguish this new species from all other species of Promitzehoia. A single pygidium, described as a new species of Stephanocare, S. fenghangensis (Zhou in Zhou et al., 1977, p. 196, pl. 58, fig. 7) from the Huaqiao Formation near Fenghuang, northwestern Hunan, may also represent a species of Protaitzehoia. It is from the same formation and region as the new species, and is morphologically similar to the meraspid pygidium of P. spinifera. However, its shorter, less segmented axis and dense granulation serve to differentiate S. fenghuagensis from the new species. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the Linguagnostus reconditus Zone). Protaitzehoia sp. Plate 32, figure 14 2001c

?Protaitzehoiasp., Peny, Babcock, Lin, Chen, and Zhu, p. 166, pl. 3, fig. 11.

Material. One incomplete pygidium, NIGP 133529, in collection PI3-2.75. Remarks. A single, incomplete pygidium is characterized by having a long, conical axis beating two longitudinal rows of coarse nodes, a flat pleural region with a narrow pleural field, faint border furrows, and wide lateral and posterior borders. The pleural field has a wide, convex anterior band that is directed outward and slightly rearward within the border furrow and strongly deflected rearward beyond the border furrow. The posterior band is short (tr.) and ridge-like, merging into the border abaxially (beyond the border furrow). Scattered granules are present on the pleural field. In having granulose ornamentation, this pygidium is similar to the pygidium of Protaitzehoia subquadrata (see P1. 34, figs. 3-6), but it differs in lacking wrinkles on the pleural field, and in having an anterior pleural band that is more convex and more strongly deflected rearward. This pygidium occurs in the upper part of the Liostracina bellla Zone and therefore is the youngest species of Protaitzehoia. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone (equivalent to the Glyptagnostus stolidotus Zone). • 120-

Subfamily DORYPYGELLINAEKobayashi, 1935 Genus TEINISTIONMonke, 1903 Teinistion Monke, 1903, p. 117; Walcott, 1913, p. 109, 110; Sun, 1924, p. 31; Kobayashi, 1931, p. 176; 1935, p. 254, 255; 1955, p. 92, 93; 1960b, p. 352; Endo, 1937, p. 337; 1944, p. 78; Hup6, 1955, p. 166; Howell and Moore in Moore, 1959, p. 248; Zhu, 1959, p. 61, 99; Lu, Qian, and Zhu, 1963, p. 112; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 408; 13pik, 1967, p. 333, 334; Zhou, Liu, Meng, and Sun, 1977, p. 200; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 186; Zhang and Wang, 1985, p. 463; Peng, 1987, p. 109; Zhang and Jell, 1987, p. 218; Lu and Lin, 1989, p. 140; Zhu and Wittke, 1989, p. 221; Zhu, 1992, p. 355, 356; Guo, Zan, and Luo, 1996, p. 127. Dorypygella Walcott, 1905, p. 29; Kobayashi, 1935, p. 255; 1941, p. 29; 1960b, p. 352; Hup6, 1955, p. 166; Lu, 1957, p. 274; Zhu, 1959, p. 62, 101; Lu, Qian and Zhu, 1963, p. 111; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 405; Zhou, Liu, Meng, and Sun, 1977, p. 200; Yin and Li, 1978, p. 513; Yang, 1978, p. 63; Liu, 1982, p. 321; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 185; Lin, Lin, and Zhou, 1983, p. 406; Qian and Zhou, 1984, p. 176; Zhang and Wang, 1985, p. 463; Zhu, 1992, p. 355. Metashantungia Chang (Zhang), 1957, p. 31; Zhu, 1959, p. 64; Lu, Qian, and Zhu, 1963, p. 112; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 411,412; Nan, 1976, p. 339, 340; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983 p. 187; Zhang and Jell, 1987, p. 220. Histiomona Opik, 1967, p. 335, 336. Jiawangaspis Zhang in Qiu et al., 1983, p. 187, 188. Type species. Teinistion lansi Monke (1903, p. 117) from the Kushan Formation, Yan-tzy-yai, Shandong. Other species. Besides the type species, Opik (1967, p. 333) included two species, T. subconicum Sun (1924, p. 31, pl. 2, fig. 4) and T. tuncatum Endo (1937, p. 337, pl. 64, fig. 7; pl. 65, figs. 1-3), in Teinistion, and excluded T. obtusum Endo, 1937 and T. sulcatum Endo, 1937 from the genus. To this list the following species should be added: Dorypygella typicalis Walcott (1905, p. 29; 1913, p. 111-113, pl. 9, figs. 2, 2a-c) from the Blackwelderia Zone, Kushan Formation, near Yanzhuang, Xintai, Shandong; Metashantungia brevica Chang (Zhang, 1957, p. 31, pl. 1, fig. 6) from the Drepanura Zone, Kushan Formation, near Yanzhuang, Xintai, Shandong; Teinistion tangshihlingensis Chu (Zhu, 1959, p. 62, pl. 2, figs. 24, 25) from the Blackwelderia paronai Zone, Kushan Formation, Dangshi, Yantai, Liaoning; Teinistion liaoningensis Chu (Zhu, 1959, p. 62, pl. 2, figs. 26, 27), from the Blackwelderia paronai Zone, Kushan Formation, Dangshi, Yantai, Liaoning; Teinistion? amydium Opik (1967, p. 334, 335, pl. 44, figs. 1, 2, text-fig. 126) from the Glyptagnostus stolidotus Zone, Georgina Limestone, northwestern Queensland, Australia; Histiomona oculosa Opik (1967, p. 336, 337, pl. 44, figs. 3, 4, text-fig. 127), also from the Glyptagnostus stolidotus Zone, Georgina Limestone, northwestern Queensland, Australia; Dorypygella posterocosta Yang in Zhou et al. (1977, p. 200, pl. 59, figs. 15, 16) from the Huaqiao Formation, Laochatian, Fenghuang, northwestern Hunan, and Jimachong, Yuping, eastern Guizhou; Teinistion handanensis Zhang and Wang from the Kushan Formation, southern Hepei; Teinistion bellum Guo and Luo (in Guo et al., 1996, p. 128, 129, pl. 64, figs. 11, 12), from the Damesella-Yabeia Zone, Changhia Formation, southern Liaoning; Jiawangaspis dananzhuangensis Zhang (in Qiu et al., 1983, p. 188, pl. 62, fig. 2) from the Kushan Formation, Jiawang, southern • 121.

Jiangsu and Teinistion wangcunense sp. nov. The following species, which were assigned originally either to Teinistion or Dorypygella, are invalid or problematic: Teinistion yangi Chu, 1959 (junior synonym of Teinistion lansi Monke, 1903); Teinistion hongshuizhuangense Zhang and Wang, 1985 (junior synonym of Teinistion tangshihlingense Chu); and Teinistion hunanense Peng, 1987 (junior synonym of Teinistion posterocostum Yang). Dorypygella alastor Walcott, 1905 does not belong to Teinistion; it was transferred to Blackweldera by Walcott (1913) and transferred tentatively to Parablackwelderia [=Damesops] by Zhang and Jell (1987). Dor3'pygella langa Kobayashi 1942c (p. 42, pl. 2, fig. 5) is a pygidium that appears to not belong to Teinistion. Teinistion lansiforme Endo, 1944 assigned questionably to Teinistion by Kobayashi, 1955 is an unrecognizable species. Both Teinistion nanhaitouense Guo and Luo (in Guo et al., 1996, p. 128, pl. 9, fig. 14; pl. 67, fig. 13) and Teinistion granulosum Guo and Luo (in Guo et al., 1996, p. 128, pl. 67, fig. 12), from the same locality in southern Liaoning, are characterized by having a triangular, pointed glabella, and are probably synonymous with T. sulcatum Endo (1937, p. 338, 339). T. sulcatum was questionable transferred to Dorypygella by Kobayashi (1955, p. 93) and was excluded from Teinistion by Opik (1967, p. 333) because of its distinct glabellar morphology.

Remarks. Walcott (1913) and Zhang and Jell (1987) discussed the concept of the genus, and considered Teinistion and Dorypygella to be synonymous. The genetic concept of both Walcott (1913) and Zhang and Jell (1987) is followed here. Besides Doo'pygella, new material from Hunan suggests that Histiomona Opik, 1961 and Jiawangaspis Zhang in Qiu et al., 1983 should also be considered junior synonyms of Teinistion. Zhang and Jell (1987, p. 218) had tentatively considered Histiomona to be synonymous with Teinistion. Histiomona includes only one species, H. oculosa, which is based on two cranidia (Opik, 1967, pl. 44, figs. 3, 4) from the Glyptagnostus stolidotus Zone, Queensland, Australia. The description and illustration of H. oculosa indicate no significant differences from specimens assigned here to Teinistion. H. oculosa has a wider and less upturned anterior cranidial border than that usually present in species assigned to Teinistion; however, these characters are considered to be of only specific value. Width and degree of upturning of the anterior border are variable even within a single species, as demonstrated by Monke (1903), and as also indicated in our new material. The stratigraphic occurrence of Histiomona falls within the range of Teinistion, and this is consistent with synonymizing the two taxa. Jiawangaspis Zhang is monotypic. The genus and its type species, J. dananzhuangensis, are best considered to be Teinistion. The holotype and only known specimen of J. dananzhuangensis is a small (3 mm), incomplete cranidium from the Kushan Formation, Jiawang, northern Jiangsu (Qiu et al., 1983, pl. 62, fig. 2). The holotype cranidium has steep inwardly inclined palpebral areas, which are characteristic of Teinistion. Although the palpebral lobes are broken, they were apparently in a relatively posterior position. Like Histiomona, Jiawangaspis has a wide and gently upturned anterior border. Features of Jiawangaspis are consistent with the morphology of juveniles of Teinistion (see Monke, 1903, P1.4, figs. 7, 6; P1. 35, figs. 1-3, herein), and the holotype of J. danazhuangensis occurs in association with holaspid Teinistion in the Kushan Formation of Jiawang, where the holotype was collected (Zhu, 1959). Metashantungia Chang (Zhang, 1957) is apparently a synonym of Teinistion. Both taxa were originally reported from the Drepanura Zone in Shandong. The type species of Metashantungia is Metashantungia brevica Chang, which is based on an incomplete cranidium referred to Shangtungia spinifera Walcott, 1905 (Walcott, 1913, pl. 14, fig. 6e; also Zhang and Jell, 1987, pl. 108, fig. 3, left). The cranidium conforms in all respects to that of Teinistion lansi (Monke, 1903) except for angulation of the anterior margin and a transverse, rather than forwardly concave, border furrow. As demonstrated by a large collection described subsequently for Teinistion and • 122.

Dorypygella (see synonymy above) and by a the new material assigned here to Teinistion, such differences fall well within the morphological variety of Teinistion. Moreover, the pygidium that is preserved in ventral view is associated with the holotype cranidium of M. brevica (Zhang and Jell, 1987, pl. 108, fig. 3, fight). It is indistinguishable from the ventrally preserved pygidia of Teinistion lansi figured by Monke (1903, pl. 4, fig. 16) and Lu et al. (1965, pl. 77, fig. 15). This evidence supports suppression of Metashantungia as a junior synonym of Teinistion. Teinistion posterocostum (Yang in Zhou et al., 1977)

Plate 35, figures 1-12; Plate 36, figures 1-12; Plate 37, figures 1-6; Text-figure 17 1977 1978 1978 1982 1983 1987 1989 2001c 2001b

Dorypygella posterocosta Yang in Zhou et al., p. 200, pl. 59, figs. 15, 16. Dorypygella posterocosta Yang; Yin and Li, p. 513, pl. 171, figs. 1, 2. Dorypygella posterocosta Yang; Yang, p. 63, pl. 11, figs. 7-11. Dorypygella posterocosta Yang; Liu, p. 321, pl. 222, figs. 4, 9. 18. Dorypygella posterocosta Yang; Lin, Lin, and Zhou, p. 406, pl. 3, fig. 11. Teinistion hunanensis Peng, p. 109, 110, pl. 12, figs. 4-6, 12. Teinistion sp. Lu and Lin, p. 254, pl. 22, fig. 8. Teinistion posterocosta (Yang); Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 3, figs. 1-4. Teinistion posterocosta (Yang); Peng, Babcock, Lin, p. 104, pl. 10, figs. 12, 13.

Lectotype. Incomplete, mostly exfoliated cranidium (Yang in Zhou et al., 1977, pl. 59, fig. 15; refigured by Yang, 1978, pl. 11, fig. 9, Liu, 1982, pl. 222, fig. 4; CUGB 0321501; Text-fig. 16A-C herein), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Laochatian, Fenghuang, northwestern Hunan; designated subsequently as the holotype of the species by Yang (1978). New material. More than 100 sclerites, including cranidia, pygidia, and one hypostome (illustrated specimens NIGP 137622-137644), in collections P273.46, P282.6, P316.1, P317, P317.2, P317.4, P319.6, P319.8, P326.9, P348, P353.64, P361.5, P363.5, P13-2.75, W215.1, W216.5, W218.3, W219.7, W221.5, W225, W227, W228.3, W229.9, W243.6, W254.1, and P[3-2.75. Remarks. New material from the Huaqiao Formation, northwestern Hunan, resembles the type material from northwestern Hunan and eastern Guizhou. Key characters of this species include an angulate anterior cranidial margin; a glabella that is moderately tapered forward and rounded to truncate anteriorly; large and thick palpebral lobes that are semicircular in shape and strongly elevated; a pair of weakly defined bacculae that are separated from the L1 lobes; wide eye ridges that are oblique and curve posteriorly from the frontal lobe of the glabella; steeply inclined palpebral and postocular areas of the fixigena; an occipital furrow that curves forward at the sides; gently divergent or subparallel anterior branches of the facial suture; a subtriangular pygidium that possesses a pair of long anterolateral spines derived from first pleura, and six pairs of moderately long, flat border spines; a long conical axis with a prominent postaxial ridge reaching to the posterior border; and deep, straight pleural furrows. The surface in this species is covered with fine granules. Examination of the type material of Yang (1978) shows that the holotype is a mostly exfoliated cranidium. Paratypes include a testaceous cranidium and two exfoliated associated pygidia that are covered with fine and dense granules, just like the surfaces in new collections from Hunan.

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Text-figure 17. Type material of Teinistion posterocostum (Yang in Zhou et al., 1977), all × 10, from the Huaqiao Formation at Laochatian, Fenghuang, western Hunan. A-C, holotype cranidium, original of Yang in Zhou et al., 1977 (pl. 59, fig. 15; also Yang, 1978, pl. 11, fig. 9, CUGB 0321501), in dorsal, obliquely anterior, and anterolateral views; D, two paratype pygidia in association, original of Yang in Zhou et al., 1977 (pl. 59, fig. 16; also Yang, 1978, pl. 11, fig. 10; CUGB 0324202); E, paratype cranidium, original of Yang, 1978 (pl. 11, fig. 8, CUGB 0324201). A large number of specimens in new collections show considerable variation in the species. The glabella tapers moderately to strongly. The glabellar front varies from acutely rounded to evenly rounded, and to firmly truncate. The anterior cranidial border is upturned variably, or upturned initially and then deflected gently forward. The pygidium varies in shape from subtriangular to transverse-subrhomboidal with an anterolateral spine that is directed posterolaterally. The smallest known cranidium (P1.35, fig. 1) shows that the palpebral lobe is gently curved rather then semicircular as in adults; and that the eye ridge is faint, rather than distinct. The anterior border is long (sag.), slightly upturned, and defined posteriorly by a preglabellar furrow and a pair of weak, anteriorly-arched border furrows at the sides. The anterior branches of the facial sutures are moderately divergent rather than subparallel. An associated hypostome is tentatively assigned to the species. Like the hypostomes in other damesellids, it has wide (tr.) lateral borders that are moderately inclined inward. The hypostome has an angulate anterior margin and an evenly rounded posterior margin, a long and convex median body that is acutely rounded anteriorly with a short (sag.), crescentic posterior lobe, and a wide (sag., exs.) and flat posterior border. • 124.

The trivial name T. posterocosta has been modified to T. posterocostum in order to agree with the neuter gender of the genetic name.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2 and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone through the Glyptagnostus stolidotus Zone). Genus TAIHANGSHANIAZhang and Wang, 1985

Taihangshania Zhang and Wang, 1985, p. 464, 465. Type species. Taihangshania shanxiensis Zhang and Wang (1985, p. 465, pl. 139, figs. 9-16), Kushan Formation, central Shanxi Province, China; by original designation. Other species. Drepanura transversa Chu (Zhu, 1959, p. 63, pl. 3, figs. 2, 3; also pl. 5, figs. 4, 5, cranidia referred previously to Chiawangella pustulosa Chu, 1959 [=Chiawangella pacifica (Walcott, 1911) ]), from the Drepanura premisnili Zone, Kushan Formation, near Jiawang, northern Jiangsu; Dorypygella hsihsienensis Chu (Zhu, 1959, p. 62, 63, pl. 2, fig. 28; pl. 3, fig. 1) from the Kushan Formation, Xixian, Shanxi; Teinistion? amvdium Opik (1967, p. 334, 335, pl. 44, figs. 1, 2; text-fig. 126) from the Glyptagnostus stolidotus Zone, Georgina Limestone, northwestern Queensland, Australia; and Taihangshania wangcunensis sp. nov. Emended diagnosis. Dorypgellinae with glabella that is bullet-shaped, acutely rounded anteriorly, and with pit-like lateral furrows; bacculae present; palpebral lobe large, strongly elevated and located posteriorly; palpebral area of fixigena narrow (tr.), inclined strongly adaxially; preglabellar field present or absent; facial suture with anterior branches convergent forward and posterior branches straight, diagonally directed; posterolateral projection of fixigena subtriangular or triangular in shape. Pygidium transversely subtriangular, axis tapered, occupying one-fifth of pygidial length; borders obscure; with seven pairs of pygidial spines; anterolateral pair of spines stout and long, derived from first segment; posterior six pairs of border spines short and uneven in length. Remarks. Taihangshania is closely similar in both cranial and pygidial aspects to Teinistion but differs in having a glabella that is tumid posteriorly and strongly tapered in the anterior half; having lateral glabellar furrows that are located on the slopes of the glabella and impressed deeply inward, giving a pit-like appearance; having oblique or diagonally direrted posterior branches of the facial suture rather than transversely directed posterior branches; and in having narrow (tr.) and triangular-shaped rather than long (tr.), blade-like posterolateral projections on the fixigenae. In pygidial characters, Taihangshania differs from Teinistion in having stronger anterolateral spines, and in lacking a clearly defined border and border furrows. Drepanura transversa Chu is here transferred to Taihangshania. This species was based on two pygidia from a single collection (Ls 56) in the Drepanura premesnili Zone of the Kushan Formation at Jiawang Coal Field, northern Jiangsu (Zhu, 1959, pl. 3, figs. 2, 3). Morphologically, these two pygidia are considerably different from those of Drepanura premesnili, the type species of Drepanura (cf. Zhang and Jell, 1987, pl. 105, figs. 1, 2; pl. 107, figs. 1-4) but closely similar to those assigned here to Taihangshania wangcunensis sp. nov., from the Huaqiao Formation, • 125-

northwestern Hunan. Examination of the collection Ls 56 (see Zhu, 1959, p. 52) at the Nanjing Institute of Geology and Palaeontology reveals that the pygidia are in association with cranidia of Taihangshania, which were originally assigned to Chiawangella pacifica (Walcott) [=Chiawangella pustulosa Chu] by Zhu (1959, pl. 5, figs. 4, 5). As discussed above under the genus Chiawangella Chu, the pygidium-based Chiawangella has a unique leiostegiid-like cranidium bearing two pairs of intergenal spines. A single cranidium referred to Shangtungia (Metashandongia) brevica Chang from the Kushan Formation of Liaoning (Zhu, 1959, pl. 3, fig. 6) resembles the type species Taihangshania shanxiensis in most respects except for lacking bacculae. More material is needed to clarify whether the cranidia are referable to Taihangshania. Taihangshania resembles Oculens Poletaeva and Romanenko, 1970a, b, which is known only from northern Gornyi Altay, Russia. The large palpebral lobes, short (tr.) posterolateral projections, bullet-shaped glabella, and the nature of the anterior area of the cranidium are similar in both genera. The Russian genus is also of similar age to Taihangshania. Oculens, however, is regarded as a richardsonellid (Remopleurididae), and can be differentiated by the placement of its palpebral lobes nearly against the axial furrow, and by the forwardly divergent anterior branches of the facial suture.

Taihangshania wangcunensis sp. nov. Plate 37, figures 7-15; Plate 38, figures 1-6; Text-figure 18 1977

Bergeronitesaustriacus Yang, Zhou, Liu, Meng, and Sun (in part), p. 199, pl. 59, fig. 4; non fig. 3 [= Paradamesella peculiaris Zhou in Zhou et al., 1977]. 1978 Bergeronites austriacus Yang, Yang (in part), p. 62, pl. 11, figs. 3, 4; non figs. 1, 2 [= Paradamesella peculiaris Zhou in Zhou et al., 1977]. 1982 Bergeronites austriacus Yang, Liu (in part), p. 321, pl. 222, fig. 3; non figs. 1, 2 [= Paradamesella peculiaris Zhou in Zhou et al., 1977]. 2001b Blackwelderia sp., Peng, Babcock, and Lin, p. 105, pl. 13, figs. 5, 6. Etymology. From Chinese, named for Wangcun, an historic town in Yongshun County, Hunan Province, and from where the type material was obtained. Holotype. Testaceous, incomplete cranidium (P1.37, fig. 9, NIGP 137647) in collection P319.8. Paratypes. Five cranidia, and two broken pygidia (NIGP 137645, 137646, 137648-137652), in collections P308, P319.8, P331.8, W225, W227, and W251.15. Diagnosis. Taihangshania with a glabellar length about twice width, and with small triangular posterolateral projection; preglabellar field absent in holaspids; anterior branch of facial suture slightly convergent forward, posterior branches of facial suture diagonally directed. Pygidium with markedly furrowed pleural field, and narrow, poorly defined borders. Description. Cranidium rectangular except for palpebral lobe and posterolateral projection, longer than wide. Glabella tapered moderately forward between L 1 and $2, then tapered strongly to form acutely rounded front; baccula narrow and long, separated from L1. Lateral glabellar furrows consist of three pairs, deeply impressed; S1 longest and widest, directed rearward and slightly • 126-

inward; $2 pit-like, isolated from axial furrow; $3 located on anterolateral comer of glabella, just behind inner edge of eye ridge, directed forward and inward. Occipital furrow shallow and transverse medially, deeply impressed and deflected forward at sides; occipital ring large, much longer than L1 and wider (tr.) than glabellar base, narrowing abaxially; occipital node small and weak, lying close to occipital furrow. Eye ridge wide, diagonally directed, becoming faint anteriorly. Palpebral lobe reniform, thick, lying posterior to glabellar midlength, elevated slightly higher than summit of glabella, with posterior part hanging over anterior part of posterolateral projection; palpebral furrow incised, semicircular. Preocular, palpebral, and postocular areas of fixigena about half as wide as glabella, with the palpebral and postocular areas inclined steeply to axial furrows. Anterior branch of facial suture initially convergent at angle about 15°-20 ° to the sagittal line, then strongly curving adaxially after crossing anterior border furrow to meet anterior margin about one-fourth of width of anterior border at each side; posterior branch short and straight, diagonally directed, enclosing a narrow triangular posterolateral projection. Posterior border furrow shallow and wide; posterior border of uniform width, with inner portion of posterior margin denticulated.

/

Text-figure 18. Reconstruction of cranidium and pygidium of Taihangshania wangcunensis sp. nov. Cranidium is based mainly on the holotype NIGP 137647 and the paratype cranidia NIGP 137648, 137650 (see P1. 37, figs. 9, 11, 12); pygidium is based on the paratype pygidia NIGP 137651, 137652 (see P1.37, fig. 14; PI. 38, figs. 1, 3).

• 127.

Pygidium subtriangular except pleural spines, length half width. Axis tapered gently rearward, constricted slightly at sides, beating four tings defined by moderately deep ring furrows and a fifth ring defined by faint ring furrow; six pairs of pleural furrows and seven pairs border spines; anterolateral spine long and stout, other spines short and directed posteriorly, with length decreasing adaxially except for the fifth pair. Border furrow absent. Surfaces of the posteromedial part of glabella, occipital ring, pygidial axis and pleural area with dense or moderately dense granules of uniform size. Remarks. This new species includes some specimens previously assigned to Blackwelderia or Bergeronites. The convergent facial suture, excavated anterior area of the cranidium, and the spinebearing posterior border of the pygidium suggests the morphology is comparable to that of some specimens assigned to Blackwelderia sinensis (Bergeron) by Walcott (1913, pl. 9, figs. 5a, 5b), but the large, strongly curved palpebral lobe that is highly elevated and hangs over the posterior area of the fixigena, the steeply inclined palpebral and posterior areas of the fixigenae, and the thick eye ridges, suggest that specimens referred here to the new species are not conspecific with B. sinensis. Two pygidia assigned originally to Bergeronites austriacus Yang in Zhou et al., 1977 (pl. 59, fig. 4; Yang, 1978, p. 62, pl. 11, figs. 3, 4) are referred here to Taihangshania wangcunensis sp. nov. The type specimens of B. austriacus includes two cranidia and two pygidia housed in the Museum of the China University of Geology (Beijing). Examination of the museum collection by one of us (SP) indicates that only the holotype cranidium and one paratype pygidium remain. The remaining paratype pygidium is incomplete (P1. 37, figs. 5, 6 herein), but its observed characters make it indistinguishable from the pygidium of the new species. In having a bullet-shaped glabella, convergent anterior branches of the facial suture, narrow fixigenae, and granulate ornamentation on part of the surface of the cranidium, T. wangcunensis is most similar to Taihangshania shanxiensis, the type species of the genus. However, T. shanxiensis is differentiated by having a proportionally shorter glabella, a wider preglabellar area, a short and less impressed S1, a relatively shorter (sag.) occipital ring with a centrally rather than anteriorly located median node, and a wider (tr.) posterior area of the fixigena. The pygidium assigned here to the new species differs from that of T. shanxiensis by having poorly defined borders and rounded rather than fiat pygidial spines. Teinistion? amydium Opik (1967, pl. 44, figs. 1, 2), which is based on two fragmentary cranidia from the Glyptagnostus stolidotus Zone, Georgina Limestone, northwestern Queensland, Australia, may represent immature individuals. This interpretation is based on their close similarity, both in morphology and in size, to juvenile cranidia of the new species. The Australian species differs in having wider palpebral areas of the fixigena. Poor preservation of the Australian specimens limits further comparison. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and W angcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone and the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Subfamily DREPANURINAE Hup6, 1953 Genus DREPANURABergeron, 1899 Type species. Drepanura premesnili Bergeron (1899, p. 406, pl. 13, fig. 8) from the Kushan Formation of North China, locality unknown; by original designation. • 128.

Remarks. Cooper et al. (1996, p. 380) synonymized Drepanura, Bergeronites, and Palaeadotes. All species reassigned by Zhang and Jell (1987, p. 220, 221) to those genera were referred by Cooper et al. (1996) to Bergeronites Sun in Kuo (Guo,1965). This synonymization apparently ignores the priority held by Drepanura Bergeron, 1899, and is not supported here. Morphologically, Drepanura is characterized by having a strongly tapering glabella; a small, anteriorly located palpebral lobe that is close to the anterior lobe of the glabella, strongly oblique posterior branches of the facial sutures; a broad (exs., tr.), subtriangular posterolateral projection; and a pygidium having a short axis and flat pleural field. As discussed further under Palaeadotes, these distinctive features warrant genetic rank for Drepanura, and warrant retaining Drepanura, Bergeronites and Palaeadotes as separate genera (following Zhang and Jell, 1987). Zhang and Jell (1987) designated "the pygidium figured by Bergeron (1899, pl. 13, fig. 8)" as the 'holotype' of Drepanura premesnili. However, the original illustration (Bergeron, 1899, pl. 13; E.N.S.M. 9000) has six pygidia or more of D. premesnili, of which three are numbered as 8. A unique name-bearing specimen should be chosen from among those pygidia, and the pygidium on the upper left of Bergeron's (1899) pl. 13 (P1.45, fig. 1A, herein) is here designated as lectotype. This is probably the specimen on which Bergeron's line-drawing of D. premesnili (Bergeron, 1899, p. 509, text-fig. 8) is based. Bergeron's (1899) plate is a rock slab illustrated at natural size, purchased from Beijing, and said to be from the hills north of Beijing. As noted by Monke (1903, p. 100), the locality is questionable. So far, there are no reports of Drepanura from the northern Beijing area, although there are upper Cambrian outcrops known in this area. Bergeron's specimen is probably from Dawenkou, Tai'an, Shandong, where the Drepanura premesnili and Blackwelderia-rich mudstone, the famous "Bat-stone", has been quarried for commercial purposes for more than 100 years. Besides the type specimens of D. premesnili, this slab also contains the type material of Blackwelderia sinensis (Bergeron), the type species of Blackwelderia, Stephanocare(?) sinensis (Bergeron), and Pseudagnostus douvillei (Bergeron). The original slab is probably missing or in the personal collection of Premisnil's descendents (Abel Prieur, Universit6 Claude Bernard, Lyon, France, personal communication). The 'type specimen' housed in the Museum of Lyon University is only a gypsum cast from that slab (P1. 45, fig. 1). Even so, the concept of Drepanura is understandable from the plastotype. As noted by Zhang and Jell (1987), the genetic concept is sufficiently understood because of the local abundance of the type species in North China. As noted by Zhang and Jell (1987, p. 220, 221), the type species is the only species belonging with certainty to Drepanura. A number of species have been classified in Drepanura, but they should be transferred to other damesellid genera. A new species, D.? crassispira is referred with question to Drepanura.

Drepanura ? crassispina sp. nov. Plate 45, figures 5-7 2001b Undeterminedpygidium; Peng, Babcock, and Lin, p. 105, pl. 14, fig. 14.

Etymology. Combination of two Latin words, crass, stout, fat, and spina, spine, referring to the stout and tumid anterolateral spines.

Holotype. Incomplete, testaceous pygidium in collection W218.3 (P1.45, fig. 7, NIGP 137712). Paratype. One pygidium (NIGP 137711) in collection W219.7. • 129.

Diagnosis. Damesellid with pygidium having large and tumid first pleural segment beating wide pleural furrows and rearward directed pleural spines; axis moderately long, poorly subdivided, rounded posteriorly, without postaxial ridge; border furrow obscure, lateral and posterior borders moderately wide, with at least four pairs of denticulate border spines.

Description. Pygidium transverse, length about twice width except for spines; anterior margins transverse, with large triangular facets anterolaterally; fulcrum located about two-thirds of pleural width from axial furrow. Axis about three-fourths of sagittal length, tapering gently rearward, rounded posteriorly, with a linear articulating half-ring, three poorly defined tings, and semicircular terminal piece reaching poorly defined border furrow. First pleural segment markedly large, beating wide, slightly curved pleural furrow directed almost diagonally and extending onto the base of stout, rearward-directed, rapidly tapering anterolateral spine; second pleura ondicated only by anterior band. Lateral and posterior borders poorly defined, occupying about one-fourth sagittal length, smooth, slightly convex, beating four pairs of small, denticulate spines. Posterior margin (excluding the border spines) bowed rearward slightly.

Remarks. In general respects, especially the large anterolateral spines and the poorly divided, flattened pleural area behind the macropleura, the pygidium of D.? crassispina is closely comparable to the pygidium of Drepanura premesnili, the type species of Drepanura. However, some notable differences prevent us from confidently referring them to Drepanura. In D. premesnili, the pygidium has no facets, and probably no pleural furrows on the first segment. Its axis is proportionally shorter than that of the new species, occupying only about half of the pygidial sagittal length, and is posteriorly obtusely pointed rather than rounded. Its first pleura is proportionally much smaller than that of the new species, and lacks a pleural furrow. Moreover, D. premesnili has a subtriangular rather than subrectangular pygidial outline. The pygidium of Neimonggolaspis pata,us Zhang and Liu, 1991 (p. 96, pl. 1, figs. 10-18) has a similarly strong first pleural segment and similar directed anterolateral spines, but differs in having a postaxial ridge on the axis and in lacking border spines between the anterolateral spines. Drepanura is rare in South China although it is abundant in North China. The report of Mansuy (1915, 1916) on its occurrence near the border between Yunnan and Vietnam are questionable.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is associated with trilobites indicative of the uppermost part of the Wanshania wanshanensis Zone and the lowermost part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Genus

PALAEADOTES

Opik, 1967

Palaeadotes (3pik, 1967, p. 339, 340: Ergaliev, 1980, p. 152: Zhang and Jell, 1987, p. 220; Lisogor, Rozov, and Rozova, 1988, p. 72; Zhang, 1996, p. 71, 72; Shergold, Feist, and Vizcaino, 2000, p. 620. Drepanura (Spinopanura) Kushan, 1973, p. 144. Bergeronites Sun in Kuo (Guo), 1965: Zhou, Liu, Meng, and Sun, 1977, p. 199; Yang, 1978, p. 61; Lu and Zhu, 1980, p. 20, 21; Zhang, 1981, p. 178: Liu, 1982, p. 321; Luo, 1982, p. 6; Lin, Lin, and Zhou, 1983, p. 406; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 181; Feist and Courtessole, 1984, p. 178; Lu and Lin, 1989, p. 139: Wang, Mills, Webby, and • 130.

Shergold, 1989, p. 115, 116; Duan, Yang, and Shi, 1999, p. 161. non Bergeronites Sun in Kuo (Guo), 1965. p. 631,637: Nan, 1976, p. 339; Guo and Duan, 1978, p. 453; Zhang and Wang, 1985, p. 461: Zhang and Jell, 1987, p. 222; Zhang and Liu, 1991, p. 97; Zhang, Xiang, Liu, and Meng, 1995. p. 77; Cooper, Jago, and Begg, 1996, p. 380; Guo, Zan, and Luo, 1996, p. 129. non Bergeronites (Bergeronites) Sun in Kuo; Qian and Zhou, 1984, p. 177. non Bergeronites (Falkopingia) Qian and Zhou, 1984, p. 177 [junior synonym of Paradamesella Yang in Zhou et al., 1977]. Bergeronites (Palaeadotes) Opik; Qian and Zhou. 1984, p. 177: Peng, 1987, p. 105, 106; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li. and Yan, 1991. p. 167: Yang, Yu, Liu. Su, He, Shang, Zhang, Zhu, Li, and Yan, 1993, p. 216. Type species. Palaeadotes dissidens Opik (1967, p. 341), from the Glyptagnostus stolidotus Zone, Georgina Basin, western Queensland, Australia; by original designation. Other species. Species of Palaeadotes were listed by Qian and Zhou (1984), Zhang and Jell (1987), and recently by Shergold et al. (2000). Among the species previously attributed to Palaeadotes, Bergeronites austriacus Yang and Palaeadotes acutisulcata Ergaliev are now referred to Paradamesella. Palaeadotes cf. P. italops Opik sensu Ergaliev (1980, pl. 7, fig. 4) also belongs to Paradamesella, and is probably conspecific with Paradamesella acutisulcatus (Ergaliev). Drepanura binodosa Egorova (1984, p. 24-26, pl. 5, figs. 8, 9) was reassigned to Palaeadotes by Gogin and Pegel (1997), and is now transferred to Paradamesella (see discussion under Paradamesella). The pygidium assigned to P. binodosa (Egorova) by Gogin and Pegel (1997, pl. 29, fig. 6) also belongs in Paradamesella. A number of species are now considered to be junior synonyms of Palaeadotes hunanensis (Yang in Zhou et al., 1977). Palaeadotes taoyuanensis Peng, 1987 is now regarded as a junior synonym of Palaeadotes bella (Qiu in Qiu et al., 1983). Bergeronites kaipinensis Kuo (Guo, 1965, pl. 1, fig. 8) should be reassigned back to Bergeronites, but a cranidium referred to B. kaipinensis Kuo by Zhou et al. (1977, pl. 59, fig. 10) does belong to Palaeadotes. Blackwelderia speciosa Mansuy, 1915 (pl. 2, fig. 5), transferred to Palaeadotes by Shergold et al. (2000), should also be excluded from Palaeadotes. The fragmental pygidium of Bergeronites sp. indet. (Duan, Yang, and Shi, 1999, p. 161, Fig. 7F) belongs to an undetermined species of Palaeadotes, but the fragmental cranidium (ibid. 7E) of the species belongs to Paradamesella. Zhang (1996, p. 72) noted that Bergeronites angustus (Zhang in Qiu et al., 1983) is a junior homonym of Palaeadotes angusta Ergaliev, 1980 (nora. correct., pro P. angustus). However, Ergaliev's species is now regarded as a junior synonym of Palaeadotes hunanensis (see discussion under the species P. hunanensis), and, because of this synonymy, P. angusta (Zhang, 1983) is revived as a valid species. So far as known, Palaeadotes includes 18 species" Drepanura ingens Poletaeva, 1960; Blackwelderia speciosa Lazarenko, 1966; Palaeadotes dissidens (~pik, 1967; Palaeadotes italops Opik, 1967; Drepanura (Spinopanura) erbeni Kushan, 1973; Bergeronites hunanensis Yang in Zhou et al., 1977; Bergeronites langyashanensis Lu and Zhu, 1980; Bergeronites angustus Zhang in Qian et al., 1983; Bergeronites bellus Qiu in Qiu et al., 1983; Bergeronites guichiensis Qiu in Qiu et al., 1983; Bergeronites transversus Qiu in Qiu et al., 1983; Bergeronites wannanensis Qiu in Qiu et al., 1983; Bergeronites (Palaeadotes) obtus Qian and Zhou, 1984; Bergeronites (Palaeadotes) punctatus Yang in Yang et al., 1991; Palaeadotes cf. P. dissidens Opik, 1967; Palaeadotes cf. P. italops Opik sensu Cooper et al., 1996; Bergeronites sp. sensu Yang, 1978; and Bergeronites sp. sensu Lu and Zhu, 1980. •

131.

Remarks. As indicated under Drepanura, Cooper et al. (1996) synonymized Drepanura, Bergeronites, and Palaeadotes, and included all species assigned to these genera by Zhang and Jell (1987, p. 220, 221) in Bergeronites Sun in Kuo, 1965, even though Drepanura Bergeron, 1899 holds priority over Bergeronites and Palaeadotes. We prefer to follow Zhang and Jell (1987) and Zhang (1996) in maintaining these three genera as separate genera. Palaeadotes commonly has been considered either as a subgenus of Bergeronites or as a junior synonym, but significant differences between B. (Palaeadotes) and B. (Bergeronites) support separate genetic status for Palaeadotes. In Palaeadotes, the glabella is tapered forward rather than drum-shaped as in Bergeronites, and its S 1 is bifurcate rather than non-bifurcate. Palaeadotes possesses a pair of posterolateral accessory lobules (i.e., the preoccipital lobes of Zhang, 1996) on L1, and a pair of weakly defined bacculae, whereas Bergeronites lacks these features. Palaeadotes has larger eye lobes than Bergeronites. These two genera are also different in pygidial morphology. Palaeadotes possesses much wider (tr.) pleural fields with clearly defined but incomplete interpleural and pleural furrows. The interpleural furrows in Palaeadotes become largely effaced adaxially (within the paradoublural line) and the pleural furrows become largely effaced beyond paradoublural line. The interpleural and pleural furrows are largely effaced or eliminated in Bergeronites. One pygidium assigned to B. kettleri has deep and long pleural furrows (Zhang and Jell, 1987, pl. 108, fig. 8) and may not belong to the species. On the ventral side, Palaeadotes possesses a much wider (sag., exs.) doublure than that of Bergeronites. The concept of Drepanura has been discussed by Opik (1967, p. 338, 339) and Zhang and Jell (1987, p. 220). Zhang and Jell (1987) stated that a morphological gradation exists from Drepanura through Palaeadotes to Bergeronites. It is uncertain that such a gradation exists, at least between Drepanura and Palaeadotes, or between Palaeadotes and Bergeronites, because there are no clear morphological intermediates between either pair of genera. Drepanura appears to be more closely related to Bergeronites than to Palaeadotes because they share similar pygidial and hypostomal morphology (Kobayashi, 1941). The stratigraphic distribution of these genera also weighs against a Drepanura to Palaeadotes to Bergeronites gradation because Palaeadotes is the oldest form, occurring first in the uppermost Goniagnostus nathorsti Zone (Peng and Robison, 2000). Drepanura and Bergeronites are younger, co-occurring in the Drepanura premesnili Zone, which is roughly equivalent to the Glyptagnostus stolidotus Zone of South China. Hypostomes of Palaeadotes, which were figured by Kushan (1973, pl. 30, figs. 1-3), Peng (1987, pl. 11, figs. 4, 5), and Shergold et al. (2000, pl. 5, figs. 7, 8), have gently angulated anterior margins, large, wing-shaped lateral borders, and posterior margins that are strongly arched forward. They are considerably different from those assigned to Palaeadotes by Opik (1967, pl. 50, fig. 3; pl. 51, figs. 1, 2, middle part of fig. 4; text-figs. 129, 133), which possibly belong to Paradamesella. As noted by Zhang (1996), Palaeadotes has a widespread geographic distribution, and it is usually present in slope facies. In China, Palaeadotes occurs only in South (Hunan, Guizhou, Anhui, Jiangsu, Zhejiang, Yunnan, Henan) and Northwest China (Xinjiang). It is also known from Siberia, Iran, Australia, Kazakhstan, and southern France. In contrast, Drepanura and Bergeronites have more limited distributions, occurring mostly in North and Northeast China (i.e., the North China Platform). Drepanura, however, also occurs in South China. Palaeadotes hunanensis (Yang in Zhou et al., 1977) Plate 38, figures 7-13; Plate 39, figures 1-13; Plate 40, figures 1-5; Plate 45, figures 3, 4; Plate 57, figures 9-13; Text-figures 19, 20 1963 • 132.

Drepanurasp., Egorova, Xiang, Li, Nan, and Guo, p. 39, pl. 8, fig. 2.

1965 1977 1977 1978 1980

Drepanura sp., Lu, Zhang, Zhu, Qian, and Xiang, p. 400, pl. 75, figs. 11, 12. Bergeronites hunanensis Yang in Zhou et al., p. 199, pl. 59, figs. 6-8. Bergeronites dissidens (Opik); Zhou, Liu, Meng, and Sun, p. 199, pl. 59, fig. 5. Bergeronites hunanensis Yang; Yang (in part), p. 61, 62, pl. 10, figs. 6, 8-10, non fig. 7 [= Palaeadotes bella Qiu in Qiu et al., 1983]. Palaeadotes acutisulcata Ergaliev (in part) (nom. correct., pro P. acutisulcatus), p. 152, 153,

pl. 7, fig. 7 only. 1980 1980 1981 1982 1982 1983 1983 1983 1983 1983 1987 1987 1987 ?1987 1988 1988 1989 1991 2001b

Palaeadotes angusta Ergaliev (nom. correct., pro P. angustus), p. 154, 155, pl. 7, figs. 2, 3, 6. Bergeronites sp., Lu and Zhu, p. 22, pl. 4, fig. 8. Bergeronites cf. B. hunanensis Yang; Zhang, p. 178, pl. 65, fig. 4. Bergeronites hunanensis Yang; Liu, p. 321, pl. 222, figs. 10, 11. Bergeronites dissidens ((3pik); Liu, p. 199, pl. 222, fig. 6; pl. 223, figs. 2, ?3. Bergeronites hunanensis minocollus Qian in Qiu et al., p. 182, 183, pl. 61, figs. 2, 3. Bergeronites wannanensis Qiu in Qiu et al., p. 183, pl. 61, figs. 4, [?]5. Bergeronites jiangsuensis Lin H. and Zhou in Lin T. et al., p. 406, pl. 3, figs. l a, b. Bergeronites kunshanensis Lin T. in Lin T. et al., p. 406, pl. 3, fig. 2. Bergeronites yushanensis Lin T. in Lin T. et al., p. 406, pl. 3, figs. 3, 4. Bergeronites (Palaeadotes) hunanensis (Yang); Peng, p. 106, pl. 11, figs. 1-5, text-fig. 14. Bergeronites (Palaeadotes) wulingensis Peng, p. 108, pl. 11, figs. 6, 7. Bergeronites (Palaeadotes) changdeensis Peng, p. 107, 108, pl. 11, figs. 8-10. Damesella sp., Kim, pl. 21, fig 7. Palaeadotes acutisulcata Ergaliev; Lisogor, Rozov, and Rozova (nom. correct., pro P. acutisulcatus), p.72, pl. 7, fig. 4. Bergeronites jiangsuensis Lin and Zhou; Zhu, Lin, and Zhang, p. 79, pl. 9, figs. 3a, b. Bergeronites hunanensis Yang; Lu and Lin, p. 140, pl. 22, figs. 6, 7. Bergeronites (Palaeadotes) hunanensis Yang; Lin, p. 375, pl. 1, fig. 1. Palaeadotes hunanensis (Yang); Peng, Babcock, and Lin, p. 103, 104, pl. 9, figs. 1-3; pl. 12,

figs. 1, 9. 2001 d Palaeadotes hunanensis (Yang); Peng, Babcock, Lin, and Chen, p. 141, fig. 9.13. 2001d Palaeadotes wulingensis Peng; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.14. Lectotype. Broken, almost completely exfoliated cranidium (Yang in Zhou et al., 1977, pl. 59, fig. 6;

also in Yang, 1978, pl. 10, fig. 6; Liu, 1982, pl. 22, fig. 10; Text-fig. 19A herein) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Dongpo, Xinhuang, northwestern Hunan; designated subsequently as holotype of the species by Yang (1978). New material. More than 160 sclerites, including one exoskeleton, cranidia, hypostomes, librigenae,

and pygidia (illustrated specimens 137653-137672, 137710, 137834-137845) in collections P199.9, P275.1, P279.7, P282.6, P282.75, P287.1, P290.5, P293.21, P295.13, P298.4, P298.54, P300.4, P301, P301.9, P307.4, P308, P316.1, P319.6, P319.8, P326.9, P327.4, P331.8, P341.8, P344.6, P348, P353.64, P353.7, PI3-2.75, Wl13, W170, W187.8, W189, W193.6, W194.6, W195.3, W 196.3, W 197.55, W 198.45, W 199.2, W200.3, W207.5, W210.5, W211.7, W212.45, W219.7, W221.5, W227, and W229.9. Remarks. Reinvestigation of the type material of Palaeadotes hunanensis (Yang) from the Huaqiao

Formation, northwestern Hunan, and study of rich new material of this species from the same •

133

•

formation and same region as the type material show that a number of species assigned originally to either Bergeronites or Bergeronites (Palaeadotes) from the Jiangnan Slope Belt (Peng, 1990a) in northwestern Hunan, northern Jiangsu, and southern Anhui are synonymous with Palaeadotes hunanensis. Some specimens described from Xinjiang and Kazakhstan are also referable to this species.

Text-figure 19. Type material of Bergeronites hunanensis Yang in Zhou et al., 1977. A, D-F are here transferred to Palaeadotes, and B, C is referred to Palaeadotes taoyuanensis Peng, 1987. A, lectotype cranidium from the Huaqiao Formation at Dongpo, Xinhuang, western Hunan; original of Zhou et al., 1977, pl. 59, fig. 7, also Yang, 1978, pl. 10, fig. 6, CUGB 1301003, x 2.4; D-F, pygidia, original of Yang, 1978, pl. 10, figs. 8-10, CUGB 131001, 1301002, 1211101, x 2.4, x 3.9, x 2.5. D, E, from the Huaqiao Formation at Dongpo, Xinhuang, western Hunan; F, from the Huaqiao Formation at Jimachong, Yuping, eastern Guizhou.

Palaeadotes hunanensis is distinguished by a short, wide, and relatively convex, tapered • 134.

glabella; a relatively narrow (tr.) posterior cranidial border that is shorter than the basal glabellar width; a pair of convergent anterior branches of the facial suture; and a transversely subrectangular pygidium with a wide axis bearing three clearly defined tings. It has a pair of large anterolateral spines that are relatively short (about one-third longer than the pygidial sagittal length), nearly triangular distally; and six pairs of shorter, flat, triangular, sawtooth-like spines between the anterolateral spines. Morphological variation in the pygidium includes the pygidial outline, which is transversely rectangular to transversely subtriangular. Also, variation exists in the shape and the width of the axis, and in the shape of the anterolateral spines, which vary from nearly straight to gently curved. The only complete specimen known of Palaeadotes (P1. 40, figs. 1-4) is probably an early holaspid exoskeleton of P. hunanensis. It has 12 thoracic segments, each of which bears pleural spines. The third and the seventh pairs of pleural spines are macropleural spines. The librigena has a moderately convex and relatively narrow genal field, a wide lateral border that narrows forward and becomes strongly upturned laterally, and a genal spine that extends from a position forward of the posterolateral comer. The preoccipital lobe on L1 of the glabella is clearly defined in an early holaspid stage of ontogeny, but it becomes faint during the late holaspid period.

Text-figure 20. Reconstruction of the dorsal exoskeleton of Palaeadotes hunanensis (Yang in Zhou et al., 1977), based on an early holaspid exoskeleton NIGP 137671 (see P1.40, figs. 1-4). The holotype cranidium of Palaeadotes acutisulcata Ergaliev, 1980 from Malyi Karatau, Kazakhstan, apparently belongs to Paradamesella rather than to Palaeadotes, but the pygidium that • 135.

Ergaliev (1980) assigned to that species is identical in all respects to Palaeadotes hunanensis, to which it is here referred. According to Ergaliev (1980), the pygidium is associated with the holotype cranidium of Palaeadotes angusta Ergaliev, 1980 (in collection 1351-2), a species that is indistinguishable from P. hunanensis in both cranidial and pygidial aspects, and is regarded as a junior synonym of P. hunanensis. The Kazakhatanian Palaeadotes angusta Ergaliev, 1980 and the Chinese Palaeadotes angusta (Zhang in Qian et al., 1983) are homonymous, but because the Kazakhstanian species is suppressed, the Chinese species name becomes valid. Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Dongpo, Xinhuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2, and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Pianaspis bella Zone through the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone through the Glyptagnostus reticulatus Zone). Palaeadotes bella (Qiu in Qiu et al., 1983)

Plate 41, figures 1-11 1978 Bergeronites hunanensis Yang; Yang (in part), p. 61, 62, pl. 10, fig. 7 only. 1983 Bergeronites bellus Qiu in Qiu et al., p. 182, pl. 60, figs. 3, 4. 1987 Bergeronites (Palaeadotes) taoyuanensis Peng, p. 107, pl. 12, figs. 10, 11. Holotype. Pygidium (Qiu et al., 1983, pl. 60, fig. 4" HIT 0390) from the Yangliugang Formation, Huamiao, Guichi, southern Anhui. New material. More than 20 sclerites, including cranidia and pygidia (illustrated specimens NIGP 137673-137680), in collections W215.1, W216.5, W219.7, W221.5, and W225. Remarks. New material from the Huaqiao Formation, northwestern Hunan, closely resembles the type material from southern Anhui, which is distinguished by having only five pairs of border spines between the anterolateral spines. Additionl key characters of the species include a glabella that is relatively short, wide, moderately convex, and moderately tapered; a posterior border that is wider (tr.) than the basal width of the glabella; a relatively wide palpebral area of the fixigena; and anterior branches of the facial suture that are slightly diverging forward or parallel-sided. The pygidium is characterized by having a subtriangular outline; a relatively long axis bearing four clearly defined tings; a and pair of anterolateral spines that are relatively long and posteriorly curved. Occurrence. The holotype is from the Yangliugang Formation of Huamiao, Guichi, southern Anhui, China. New material is from dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone and the lower part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone).

• 136.

Genus PARADAMESELLAYang in Zhou et al., 1977 Paradamesella Yang in Zhou et al., 1977, p. 197; Yin and Li, 1978, p. 510; Yang, 1978, p. 53; Liu, 1982, p. 320; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 178; Peng, 1987, p. 102; Lu and Lin, 1989, p. 138; Zhang, 1996, p. 70; Pegel, 2000, p. 1011, 1015, 1016. Bergeronites (Falkopingia) Qian and Zhou, 1984, p. 177. Type species. Paradamesella typica Yang in Zhou et al., 1977 (p. 197, pl. 58, figs. 9-12) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Laochatian, Fenghuang and Jiudiantang, Xinhuang, northwestern Hunan Province; Liangweizhou, Yuping, eastern Guizhou, China; by original designation. Emended diagnosis. Cranidium width 2.5 to 3 times length. Preglabellar field absent. Anterior border narrow (sag., exs.), strongly upturned; glabella subrectangular to subtrapezoidal in outline, truncate anteriorly with four pairs of lateral furrows; S 1 deeply incised with outer end obscurely bifurcate; $2 deep and short; $3 pit-like and isolated from axial furrow; $4 short and moderately incised; L1 swollen, with weak accessory lobule posterolaterally (i.e., preoccipital lobe of Zhang, 1996); bacculae present or absent. Palpebral lobe relatively small, well defined, located near cranidial midlength; eye ridge raised, with inner end connecting to frontal lobe of glabella. Anterior branch of facial suture converging forward to subparallel. Librigena with broad genal field and genal spine forward of posterolateral comer. Thorax with 12 segments; macropleurae on segments 3 and 7. Pygidium relatively large; axis variably wide, extending close to border furrow or posterior margin, with five to seven tings and a relatively swollen terminal piece; with flattened pleural field; first segment bearing long marginal pleural spines; seven to ten pairs of additional border spines. Surface covered with granules. Other species. Drepanura eremita Westerg~d (1947, p. 12-14, pl. 13, fig. 10, non fig. 9 [=Palaeadotes dissidens Opik]) from the Agnostus pisiformis Zone, Djupadalen, Falkrping, V~istergrtland, Sweden; Damesella(?) lata Romanenko in Romanenko and Romanenko (1967, p. 83, 84, pl. 3, figs. 1, 2; Romanenko, 1977, pl. 24, figs. 22a, b) from the lower part of the upper Cambrian, Bolshaya Isha River Valley, Gornyi Altai, Russia; Paradamesella peculiaris Zhou in Zhou et al. (1977, p. 198, 199, pl. 58, fig. 15), from the Huaqiao Formation, Chatian, Fenghuang, northwestern Hunan; Palaeadotes acuticulcatus Ergaliev (1980, p. 152-154, pl. 7, fig. 5, non fig. 7), from the Kormagnostus simplex Zone, Malyi Karatau, Kazakhstan; Drepanura binodosa Egorova (1984, p. 24-26, pl. 5, figs. 8, 9) from the upper Cambrian near the Aldan River, Siberian Platform; Paradamesella nobilis Lu and Lin (1989, p. 138, pl. 22, figs. 1-4) from the Lejopyge laevigata Zone, Dachen, Jiangshan, western Zhejiang; Paradamesella sp. (Pegel, 2000, Fig. 12.13; Figs. 13.17, 13.20) from the upper Cambrian Eyra Formation near the Kotoy River and the Ogon'or Formation near the Khos-Nelege River, Siberian Platform. Remarks. The genetic concept of Yang in Zhou et al. (1977, p. 197; Yang, 1978, p. 55, 56) is revised based on the material from South China, including specimens showing the thorax (Qiu et al., 1983, pl. 58, fig. 11; Lu and Lin, 1989, pl. 22, fig. 1). Paradamesella is similar in morphology to Palaeadotes Opik, and is easily confused with it because they share a glabella of similar shape with similarly impressed lateral furrows, similarly located palpebral lobes, and similar facial sutures. Such similarity has led to assignment of some specimens belonging to Paradamesella to • 137•

Palaeadotes Opik (i. e., those of Bergeronites austriacus Yang in Zhou et al., 1977 and Yang, 1978, Palaeadotes acutisulcata Ergaliev 1980, cranidium only and Palaeadotes sp. cf. P. italops Opik in Ergaliev, 1980). However, Paradamesella differs from Palaeadotes in some essential respects. It has a much wider fixigena, particularly in the anterior area, where the width is nearly as wide as the anterior margin of the glabella. The fixigena in Palaeadotes is much narrower, being only one-third to one-half of the width of the anterior margin of the glabella. The basal segment of the glabella in Paradamesella bears tumid L1 lobes at the sides and a notably depressed central area, whereas it is evenly convex (tr., sag.) in Palaeadotes. The S1 furrow in Paradamesella is either widened or obscurely bifurcate at the abaxial end, but extends inward for a short distance and then bifurcates strongly at the adaxial end in Palaeadotes. In the pygidium, the pleural and interpleural furrows are complete and of even depth in Paradamesella, but they are incomplete or partly effaced in Palaeadotes; the borders are narrow and well defined in Paradamesella, but wide and obscurely defined in Palaeadotes. Moreover, Paradamesella is differentiated by having a smaller palpebral lobe, a more oblique and more raised eye ridge, and a librigena that has a much wider genal field and a more forwardly located genal spine. Qian and Zhou (1984) erected a subgenus, Bergeronites (Falkopingia), with Drepanura eremite (Westerghrd, 1947, p. 12-14, pl. 3, figs. 9-11) from Sweden as the type species. Previously, Opik (1967) had assigned this species to Palaeadotes, mainly on the basis of cranidial features. The Swedish species includes a cranidium, a fragment of thoracic segment and a holotype pygidium. The specimens are from different localities (Westerg~d, 1947). Zhang (1996) correctly noted that the holotype pygidium of D. eremita belongs neither to Drepanura nor to Palaeadotes but to Paradamesella, and the paratype cranidium belongs to Palaeadotes. Therefore Bergeronites (Falkopingia) should be suppressed as a junior synonym of Paradamesella. Paradamesella has been assigned to the Damesellinae (Yang, 1978; Peng, 1987; Lu and Lin, 1989; Zhang, 1996), but is here classified in the Drepanurinae based on its close similarity in cranidial and pygidial morphology and thoracic segmentation to Bergeronites and Palaeadotes. Both of these genera have been traditionally classified in the subfamily Drepanurinae. In the thorax, members of all three of these genera bear macropleurae on the third and seventh segments. So far, 17 species have been assigned to Paradamesella, but of them, only six species are considered to be valid. Paradamesella euo'pterophora Zhang (in Qiu et al., 1983, p. 179, pl. 58, fig. 10) from southern Anhui and Paradamesella depressa Peng (1987, pl. 10, figs. 1-7) from the Glyptagnostus stolidotus Zone, Wa'ergang, Taoyuan, northwestern Hunan, are regarded as junior synonyms of P. typica, the type species. Paradamesella septemispinosa Yang (in Zhou et al., 1977, p. 198, pl. 58, fig. 14), Paradamesella novemospinosa Yang (in Zhou et al., 1977, p. 199, pl. 58, fig. 16), Paradamesella decemospinosa Yang (in Zhou et al., 1977, p. 199, pl. 58, fig. 13), Paradamesella paratypica Yang (1978, p. 56, 57, pl. 7, fig. 9), Paradamesella sp. indet. (Yang, 1978, p. 57, 58, fig. 13), Paradamesella trapezoidalis Peng (1987, p. 103, pl. 10, figs. 8-10), and Paradamesella subquadrata Peng (1987, p. 104, pl. 10, fig. 11) are all regarded as junior synonyms of P. peculiaris Zhou in Zhou et al., 1977. Palaeadotes sp. cf. P. italops Opik (Ergaliev, 1980, pl. 7, fig. 4) from Malyi Karatau, Kazahkstan, is a juvenile cranidium of Paradamesella. It is associated with Paradamesella acuticulcata (Ergaliev), and it is possibly conspecific with that species. A specimen referred to Paradamesella sp. cf. P. paratypica Yang (Lu and Lin, 1989, p. 139, pl. 22, fig. 5) is a fragmental pygidium, and may be conspecific with P. typica Yang. Paradamesella is widely distributed in South China (Hunan, Guizhou, Anhui, Jiangsu, Zhejiang); it occurs also in Sweden, Kazakhstan, and Russia (Siberia and Altai).

•

138

•

Paradamesella ~pica Yang in Zhou et al., 1977

Plate 42, figures 1-10; Plate 43, figures 1-7; Text-figures 21, 22 1977 1978 1978 1982 1983 1983

Paradamesella typica Yang in Zhou et al., p. 197, pl. 58, figs. 9-12. Paradamesella typica Yang; Yin and Li, p. 511, pl. 170, figs. 1, 2. Paradamesella typica Yang, p. 56, pl. 12, figs. 1-8. Paradamesella typica Yang; Liu, p. 320, pl. 221, figs. 11, 12, 14, 16. Paradamesella eurypterophora Zhang in Qiu et al., p. 179, pl. 58, fig. 10. Paradamesella decemospinosa Yang; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, p. 179, pl. 58, fig. 1. 1983 Paradamesella typica Yang; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, p. 179, pl. 59, fig. 2. 1987 Paradamesella depressa Peng, p. 102, 103, pl. 10, figs. 1-7. ?1989 Paradamesella cf. P. paratypica Yang; Lu and Lin, p. 139, 253, pl. 22, fig. 5. 1996 Paradamesella typica Yang; Zhang, p. 73, pl. 1, fig. 8. ?1999 Paradamesella cf. P. typica Yang; Duan, Yang, and Shi, p. 163, fig. 121. 2001b Paradamesella typica Yang; Peng, Babcock, and Lin, p. 104, pl. 12, figs. 7, 8. 2001c Paradamesella typica Yang; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 3, fig. 8. 2001 d Paradamesella typica Yang; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.15. Lectotype. Pygidium (Yang in Zhou et al., 1977, pl. 58, fig. 11; refigured in Yin and Li, 1978, pl. 170, fig. 1; Yang, 1978, pl. 12, fig. 6; Text-fig. 22A herein, CUGB 0321202), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Jiudiantang, Xinhuang, northwestern Hunan; designated subsequently as the holotype of the species by Yang (1978). New material. More than 50 sclerites, including cranidia, hypostomes, and pygidia (illustrated specimens NIGP 137681-137694), in collections W216.5, W219.7, W227, W242.6, W251.15, W251.5, W254.1, P325.7, P344.6, P348, P353.7, P361.6, P364.3, and PI3-2.75. Description. Yang (1978, p. 56) described the species in detail. His description in Chinese, with supplemental information from the new material, is translated as follows: Glabella subrectangular, tapering gently forward, forming an obscure crest sagittally with four pairs of lateral furrows: S 1 long, deeply incised, inward and rearward directed, with its external end bifurcated; $2 short, deeply incised, subparallel to S 1; $3 pitted, shallow and isolated from axial furrow; $4 short, moderately deep, inward and forward directed; L1 lobe large with its posterolateral portion defined by an obscure accessory furrow and forming a transversely elliptical, swollen accessory lobule; bacculae large and ill-defined. Occipital furrow transverse to gently sinuous, shallow medially, markedly deepened laterally; occipital ring widest sagittally, narrowing abaxially with a prominent, centrally placed median node. Preglabellar field absent; anterior border narrow (sag., exs.) but wide, strongly upturned. Fixigena wide (tr.), more or less flat. Palpebral lobe small semicircular, lying at about cranidial midlength, around the level of external end of S 1. Eye ridge of uniform width, oblique rearward, well defined and notably raised with its inner end connecting with frontal lobe of glabella just in front of $4 furrow. Anterior branches of facial suture divergent forward at about 25°-30 ° to sagittal line; posterior branches running outward for a distance as wide as palpebral area, then gently curving rearward to meet cranidial posterior margin. • 139-

Posterior border furrow, transverse, moderately deep, with its inner portion widened into a transversely triangular impression lying just behind baccula; posterior border transverse with its distal one-fifth to one-sixth reflected gently forward (reflected beyond fulcral joint). Hypostome subquadrate with a forward-arched anterior margin, outward-arched lateral margin, and transverse posterior margin with distal ends deflected posterolaterally to form broad notch on posterior border. Median body with subovate anterior lobe defined laterally by pair of short, deeply incised, rearward and slightly inward furrows, and separated from crescentic posterior lobe by shallow, posteriorly arched furrow. Anterior wing small, subtriangular. Lateral border broad, bearing deep impression centrally.

Text-figure 21. Type material of Paradamesella rypica Yang in Zhou et al., 1977. A, B, incomplete, somewhat crushed cranidium, CUGB 1435002, × 5, × 3, original of Zhou et al., 1977 (pl. 58, fig. 9; also Yang, 1978, pl. 12, fig. 1), B is a composite photo from A with the left half being a mirror image of the fight half; C, thorax, CUGB 0321201, x 6, original of original of Zhou et al., 1977 (pl. 58, fig. 10; also Yang, 1978, pl. 12, fig. 4), note the 3rd and the 7th segments are macropleurae; D, external mold of pygidium, reversed from the negative image, CUGB 0321202, × 6, designated subsequently as holotype (= lectotype) of this species by Yang (1978, p. 56, 82), original of Zhou et al., 1977 (pl. 58, fig. 11; also Yang, 1978, pl. 12, fig. 6). Thorax of 12 segments, with flat pleural field nearly twice width of axis, and macropleurae on third and seventh segments. Interpleural furrow clearly defined, transverse to slightly oblique; pleural furrow deep, with proximal end at anterior margin of pleura, running obliquely across pleura. Pleura bearing slender, posterolaterally directed spine and short, hooklike accessory spine in front of pleural spine. Pygidium semicircular to subtriangular in outline. Axis slender, occupying about one-fifth pygidial width at anterior margin, with 5-6 tings and long, variably swollen terminal piece being • 140.

poorly defined laterally and posteriorly. Pleural field flat and broad with posterior portion slightly curved downward. Pleura with clearly defined, straight, diagonally directed pleural furrow, and linear ridge lying posteriorly on posterior band and defined by weak interpleural furrow. First pleura beating long pleural spine and a short, hook-like accessory spine; border furrow shallow; border narrow, beating 9-10 pairs of moderately long and sharp spines. Surface with fine, densely spaced granules.

Text-figure 22. Reconstruction of dorsal exoskeleton of Paradamesella opica Yang in Zhou et al., 1977. Cephalon and pygidium are based on specimens NIGP 137687, 133526 (see P1.42, fig. 8; pl. 43, fig. 3); and the thorax is based on a specimen figured by Yang (1978, pl. 12, fig. 4; herewith, Text-fig. 19C). R e m a r k s . New material from the Huaqiao Formation, northwestern Hunan, is identical with the

type material from northwestern Hunan in almost all respects. Key cephalic characters that warrant assignment of the new material to this species include a gently tapered glabella, and a wide proximal portion of the posterior border on the cephalon. Key pygidial characters include a wide" pleural field, a slender axis, and thin spines on the pygidium. An ontogenetic series (P1. 42, figs. 1-9) shows that the glabella is cylindrical, that the $3 lobe connects with the axial furrow in the early meraspid stage, and that the anterior border is progressively shortened and increases in degree of upturning during ontogeny. The eye ridge is • 141.

horizontal and becomes progressively oblique through the holaspid period. The palpebral lobe, which is located anteriorly early in the holaspid period, shifts rearward to the midlength of the cranidium. Associated hypostomes in the new collections are almost identical to those from Australia assigned to Palaeadotes by 0pik (1967, pl. 50, fig. 3; pl. 51, figs. 1, 2, middle part of 4; text-figs. 129, 133). The Australian hypostomes, however, probably do not belong to Palaeadotes, but to Paradamesella. Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone of the Huaqiao Formation, Jiudiantang, Xinhuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2, and Wangcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone through the Liostracina bella Zone (equivalent to the upper part of the Linguagnostus reconditus Zone through the basal Glyptagnostus reticulatus Zone). Paradamesella peculiaris Zhou in Zhou et al., 1977

Plate 43, figure 8; Plate 44, figures 1-14; Text-figure 23A-G 1977 1977 1977 1977 1977

Paradamesella peculiaris Zhou in Zhou et al., p. 197, pl. 58, fig. 15. Paradamesella septemispinosa Yang in Zhou et al., p. 198, pl. 58, fig. 14. Paradamesella novemospinosa Yang in Zhou et al., p. 198, pl. 58, fig. 16. Paradamesella decemospinosa Yang in Zhou et al., p. 198, pl. 58, fig. 13. Bergeronites austriacus Yang in Zhou et al. (in part), p. 198, pl. 59, fig. 3, non fig. 4 [= Taihangshania wangcunensis sp. nov.]. 1978 Paradamesella paratypica Yang, 1978, p. 56, 57, pl. 12, fig. 9. 1978 Paradamesella novemospinosa Yang; Yang, p. 57, pl. 12, fig. 10. 1978 Paradamesella decemospinosa Yang; Yang, p. 57, pl. 12, fig. 11. 1978 Paradamesella septemispinosa Yang; Yang, p. 57, pl. 12, fig. 12. 1978 Paradamesella sp. indet., Yang, p. 57, 58, pl. 12, fig. 13. 1978 Bergeronites austriacus Yang (in part), p. 62, pl. 11, figs. 1, 2, non figs. 3, 4 [- Taihangshania wangcunensis sp. nov.]. 1982 Paradamesella decemospinosa Yang; Yang, p. 306, pl. 2, fig. 25. 1982 Paradamesella septemispinosa Yang; Liu, p. 320, pl. 221, fig. 8. 1982 Paradamesella novemospinosa Yang; Liu, p. 320, pl. 221, fig. 18. 1982 Paradamesella decemospinosa Yang; Liu, p. 320, 321, pl. 221, fig. 17. 1982 Bergeronites austriacus Yang (in part); Liu, p. 321, pl. 222, figs. 1, 2, non fig. 3 [= Taihangshania wangcunensis sp. nov.]. 1982 Paradamesella peculiaris Zhou; Liu, p. 320, pl. 221, fig. 13. 1983 Paradamesella novemospinosa Yang; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, p. 179, pl. 58, fig. 11. 1987 Paradamesella trapezoidalis Peng, p. 103, pl. 10, figs. 8-10. 1987 Paradamesella subquadrata Peng, p. 104, pl. 10, fig. 11. 1999 Paradamesella cf. P. decemospinosa Yang; Duan, Yang, and Shi, p. 164, fig. 6I. ?1999 Bergeronites sp. indet. (in part), Duan, Yang, and Shi, p. 161, fig. 7E only. 2001b Paradamesella paratypica Yang; Peng, Babcock, and Lin, p. 103, pl. 8, figs. 7, 8. 2001 d Paradamesella subquadrata Peng; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.12. • 142•

Text-figure 23. A-G, Holotype of Paradamesella peculiaris Zhou in Zhou et al., 1977 and specimens referred to the species. All specimens are from the collection of Yang (1978). All the specimens except for G were described originally by Yang (in Zhou et al., 1977; Yang, 1978) as types of species referred to either Paradamesella or Bergeronites. A, incomplete pygidium, CUGB 0329107, × 3.5, originally described as holotype of Paradamesella septemispinosa Yang (in Zhou et aL, 1977, p. 198, pl. 58, fig. 14; also Yang, 1978, pl. 12, fig. 12); B, incomplete, broken pygidium, CUGB 1301007, × 6, originally described as holotype of Paradamesella novemospinosa Yang (in Zhou et al., 1977, p. 198, pl. 58, fig. 16; also Yang, 1978, pl. 12, fig. 10); C, D, incomplete pygidium in dorsal and posterolateral views, CUGB 1301032, × 4, originally described as holotype of Paradamesella decemospinosa Yang (in Zhou et al., 1977, p. 198, pl. 58, fig. 13; also Yang, 1978, pl. 12, fig. 11); E, incomplete pygidium, CUGB 1301006, × 2.8, originally described as holotype of Paradamesella paratypica Yang (1978, p. 56, 57, pl. 12, fig. 9); F, broken, crushed, incomplete cranidium, CUGB 0329101, × 5, originally described as holotype of Bergeronites austriacus Yang in Zhou et al., 1977 (p. 198, pl. 59, fig. 3; also Yang, 1978, pl. 11, fig. 2); G, undescribed broken cranidium in association with CUGB 0329107(A), × 5. • 143"

Holotype. By monotypy; pygidium (Zhou in Zhou et al., 1977, pl. 58, fig. 15) from the Huaqiao Formation (lower part of upper Cambrian), Chatian, Fenghuang, northwestern Hunan. New material. More than 30 sclerites, including cranidia, hypostomes, pygidia, and fixigenae (illustrated specimens NIGP 137695-137708), in collections W187.8, W188.8, W189, W195.3, W195.7, W196.3, W198.45, W199.2, P216, P275.1, P277, P282.6, P298.4, P317.4, P319.6, P319.8, P325.7, and P331.8. Emended diagnosis. Paradamesella with glabella strongly tapered forward; proximal part of posterior cranidial border shorter than width of glabella at L 1. Pygidium with relatively wide axis, width of axis nearly width of pleural region; axis with five tings and markedly swollen terminal piece; border nearly flat, bearing eight to ten wide, triangular border spines. Remarks. Five species from northwestern Hunan, described by Zhou et al. (1977) in the same publication, are probably synonyms. They are Paradamesella peculiaris, P. septemispinosa, P. novemospinosa, P. decemospinosa, and P. paratypica; all based on single specimens, and all from the same zone and the same area of northwestern Hunan; the last three species are even from a single collection (single bed). These species were erected mostly on the number of border spines. New material from the Huaqiao Formation of northwestern Hunan shows a spectrum of characters indicating that these five species are gradational. P. peculiaris is chosen as the senior synonym because it is the first of these species to appear in print (Zhou et al., 1977, p. 197). The emended diagnosis embraces the morphological variation exemplified in these five species. In addition, examination of Yang's material by one of us (SP) reveals that, except for the number of pygidial spines, these species have no significant differences in morphology. A cranidium was not described previously for any of the species described by Zhou et al. (1977), but cranidia described under the name Bergeronites austriacus Yang (1978, pl. 11, figs. 1, 2) and Paradamesella sp. indet. (Yang, 1978, pl. 12, fig. 13) apparently belong to the species. These cranidia occur in the same collection (HHd.1) as P. novemospinosa, P. decemospinosa, and P. paratypica. B. austriacus Yang (1978, p. 62, pl. 11, figs. 1-4) is an invalid species, and a junior synonym of Paradamesella peculiaris. The species is based on two cranidia and two pygidia, all of which are poorly preserved. A paratype cranidium and a paratype pygidium were figured in advance of Yang's (1978) paper by Zhou et al. (1977, pl. 59, figs. 3, 4). Yang (1978, pl. 11, fig. 4) added one more pygidium to the species. The holotype cranidium of B. austriacus (Zhou et al., pl. 59, fig. 3; Yang, 1978, pl. 11, fig. 3) belongs to Paradamesella, whereas the pygidia (Yang, 1978, pl. 11, figs. 3, 4) belong to Taihangshania wangcunense sp. nov. The holotype is a badly crushed cranidium; Text-fig. 23F, herein) that has its glabella and fight fixigena rotated clockwise, giving the eye ridge a strongly oblique appearance. However, the paratype cranidium and an associated unillustrated cranidium in Yang's (1978) type material (Text-fig. 23G) show that these cranidia, characterized by having a notably tapered glabella, are conspecific with the cranidia assigned here to P. peculiaris. P. peculiaris is most similar to P. lata (Romanenko and Romanenko, 1967) from the lower upper Cambrian of Altai, and it may be a junior synonym of P. lata. Pending further investigation, we regard both species names as valid, and refer the present material to P. peculiaris. P. peculiaris also resembles P. typica, the type species of Paradamesella. However, P. peculiaris can be differentiated by having a more tapered glabella, and a posterior border that has a relatively shorter proximal portion (between the inner end and the fulcral joint). The length of the proximal part is about two-thirds of the basal glabellar width in P. peculiaris, but is as wide as, or even greater than, the basal glabellar width in P. ~pica.

.144.

Occurrence. The holotype is from the Huaqiao Formation (lower part of the upper Cambrian), Chatian, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Pianaspis sinensis Zone through the lower part of the Liostracina bella Zone (equivalent to the Lejopyge laevigata Zone through the Linguagnostus reconditus Zone). Paradamesella nobilis Lu and Lin, 1989

Plate 28, figures 8-10 1989 Paradamesella nobilis Lu and Lin, p. 138, 252, pl. 22, figs. 1-4. 2001b Paradamesella typica Yang; Peng, Babcock, and Lin, p. 102, pl. 6, fig. 5. Holotype. Incomplete thoracopygon (Lu and Lin, 1989, pl. 22, fig. 1; NIGP 66432) from the middle part of the Lejopyge armata Zone, Yangliugang Formation, Dachen, Jiangshan, western Zhejiang. New material. Three pygidia (NIGP 137555-137557) in collection P249. Remarks. New material from the Huaqiao Formation, northwestern Hunan, resembles the type material from Jiangshan, western Zhejiang, in key respects including the pygidial outline; the shape, proportion, and segmentation of the axis; and the slender, sharp spines on the pygidial margin. The weakly furrowed pleural field also falls within the morphological variation of the species. The only difference between the Hunan material and the Zhejiang material is that the new material from Hunan bears nine pairs of spines whereas the type material has only eight pairs of spines. Such a difference, however, is considered to represent intraspecific variation rather than a difference of species-level significance. Occurrence. The holotype is from the middle part of the Lejopyge armata Zone, Yangliugang Formation, Dachen, Jiangshan, western Zhejiang, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the Lejopyge laevigata Zone).

Superfamily ODONTOPLEUROIDEAKobayashi, 1935 Family EOACIDASPIDAEPoletaeva, 1957 Thomas and Holloway (1988) regarded the family Eoacidaspidae Poletaeva as a junior synonym of Lichakephalidae Tripp, 1957. This proposal was later rejected by Zhang (1990) and Shergold et al. (2000). We agree with those authors in maintaining separate family names. As noted by Shergold et al. (2000), the Eoacidaspidae embraces the morphologically similar and phylogenetically related genera Usoviana Poletaeva, 1977b, Belovia Poletaeva, 1956, Paraacidaspis Poletaeva, 1960, Eoacidaspis Poletaeva, 1956, and Archikainella Liu, 1982, all of which are time-successive, ranging from the Lejopyge laevigata Zone to the late Furongian.

• 145.

Genus PARAACIDASPIS Poletaeva, 1960

Paraacidaspis Poletaeva, 1957, p. 162 (nomen nudum); 1960, p. 68-70; Egorova, Xiang, Li, Nan, and Guo, 1963, p. 52, 53; Lu and Qian, 1964, p. 35; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 634; Luo, 1974, p. 693; 1983, p. 17; Zhou, Liu, Meng, and Sun, 1977, p. 264, 265; Yang, 1978, p. 71; Liu, 1982, p. 342; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 262; Zhang, 1990, p. 174; Shergold, Feist, and Vizcaino, 2000, p. 621,622. Type species. Paraacidaspis hunanica Egorova in Poletaeva, 1960 (p. 68, pl. 3, fig. 1; refigured with additional specimens by Egorova et al., 1963, p. 53, 54, pl. 12, figs. 6-8) from the ChuangiaProchuangia Zone, Huaqiao Formation, Tingziguan, Chatain, Fenghuang, northwestern Hunan, China; by original designation. Other species. Paraacidaspis ultima Shergold, Feist, and Vizcaino, 2000 (p. 622-626, pl. 6, figs. 1-17, text-fig. 4) from the Val d'Homs Formation, Montagne Noire, France; Paraacidaspis sp., described below. Paraacidaspis latilimbus Yang (1978, p. 72, pl. 13, fig. 15; text-fig. 9c), from the same zone in the same area as P. hunanica, is considered a junior synonym of P. hunanica. As noted by Zhang (1990, p. 176), Paraacidaspis sibirica Poletaeva (1960, p. 70, 71. pl. 3, fig. 2), from the lower part of the upper Cambrian of western Siberia, is indeterminate. Paraacidaspis sp. of Yang (1978, p. 72, pl. 13, fig. 16; text-fig. 9d), from the same zone and locality as P. hunanica, and Paraacidaspis sp. of Ergaliev (1980, pl. 14, fig. 21), from post-Chuangia-Prochuangia Zone-equivalent strata in southern Kazakhstan, are also regarded as junior synonyms of P. hunanica. Paraacidaspis triangularis Luo (1983, pl. 6, fig. 13) from the Baoshan Formation, Luishui, Shidian, western Yunnan, and Paraacidaspis sp. of Apollonov et al. (1984, p. 8, pl. 6, figs. 13, 14) do not belong to Paraacidaspis, but to Archikainella Liu, 1982. Diagnosis. The emended diagnosis of Zhang (1990, p. 174) is here followed. Remarks. The genetic concept of Zhang (1990, p. 174) is followed here. The genetic concept, validity, and taxonomic position of Paraacidaspis have been discussed by Zhang (1990) and Shergold et al. (2000) at some length. Bruton (1983, p. 898) considered the genus to be invalid, because the type species, P. hunanica Jegorova in Poletaeva, 1960 is a nomen nudum. He further regarded Paraacidaspis as a junior subjective synonym of Eoacidaspis Poletaeva, 1956. Jell and Adrain (2003, p. 335) affirmed a valid status for Paraacidaspis, but regarded P. sibirica Poletaeva, 1960 as the type species of Paraacidaspis. Shergold et al. (2000) regarded the date of validation of Paraacidaspis as 1963, when the type species was formally described by Egorova et al. (1963, p. 53, 54). However, according to the ICZN (1999), Articles 11, 13.1.1, the year 1960 should be considered the date of validation for the genus. The genetic diagnosis provided and the type species nominated and illustrated in that year (Poletaeva, 1960, p. 68, 69, pl. 3, fig. 1) met the requirements for a new taxon, and made the genetic name available. This date is also the date of validation for the type species, although the species was not formally described at that time. A statement in the discussion used to differentiate Paraacidaspis sibirica from P. hunanica clarified the characters of the type species (Poletaeva, 1960, p. 71). We concur with Zhang (1990) and Shergold et al. (2000) in regarding Paraacidaspis as a valid genus belonging in the family Eoacidaspidae. • 146-

Pygidia of Paraacidaspis were first illustrated by Shergold et al. (2000, pl. 6, figs. 11-17) for the French species Paraacidaspis ultima. The pygidium of P. ultima is characterized by having a relatively short, thin, and clearly segmented axis, wide and poorly defined borders, and pleurae with narrow anterior bands and wide posterior bands. These pygidia are comparable in overall morphology to the pygidia assigned to P. hunanica (Peng et al., 200 lb, pl. 15, fig. 11; P1. 46, figs. 7-12, 14 herein). Yang (1978, pl. 13, fig. 14) referred a fragmental pygidium to P. hunanica, but it apparently does not belong to Paraacidaspis as it has a wide, long, effaced axis and a largely effaced pleural field. That specimen is too fragmental to be interpreted, and is regarded here as an undetermined polymerid pygidium. Paraacidaspis hunanica Egorova in Poletaeva, 1960

Plate 46, figures 1-16; Text-figure 24 1960 1963

Paraacidaspis hunanica Egorova in Poletaeva, p. 68-70, pl. 3, fig. 1 (figured only). Paraacidaspis hunanica Egorova; Egorova, Xiang, Li, Nan, and Guo, p. 53, 54, pl. 12, figs.

6-8. 1964 1965

Paraacidaspis hunanica Egorova; Lu and Qian, p. 35, pl. 7, fig. 4. Paraacidaspis hunanica Egorova; Lu, Zhang, Zhu, Qian, and Xiang, p. 635, pl. 130, figs. 15,

16. Paraacidaspis hunanica Egorova; Zhou, Liu, Meng, and Sun (in part), p. 265, pl. 81, figs. 1-3, non fig. 4. 1977 Paraacidaspis latilimbus Yang in Zhou et al., p. 265, pl. 80, fig. 7. 1977b Paraacidaspis hunanica Egorova; Poletaeva, pl. 1, fig. 13 (not described). 1978 Paraacidaspis hunanica Egorova; Yang (in part), p. 71, 72, pl. 13, fig. 13, text-fig. 9a; non fig. 14, non text-fig. 9b. 1978 Paraacidaspis latilimbus Yang; Yang, p. 72, pl. 13, fig. 15; text-fig. 9c. 1978 Paraacidaspis sp.; Yang, p. 72, pl. 13, fig. 16, text-fig. 9d. 1980 Paraacidaspis hunanica Egorova; Ergaliev, p. 187, pl. 16, fig. 12 (not described). 1980 Paraacidaspis sp.; Ergaliev, p. 186, pl. 14, fig. 21 (not described). 1982 Paraacidaspis hunanica Egorova; Liu, p. 342, pl. 239, fig. 8. 1982 Paraacidaspis latilimbus Yang; Liu, p. 342, pl. 239, fig. 9. 1983 Paraacidaspis hunanica Egorova; Qiu in Qiu et al., p. 252, pl. 88, fig. 8. 1990 Paraacidaspis hunanica Egorova; Zhang, p. 174-177, pl. 1, figs. 5-9. 1996 Paraacidaspis hunanica Egorova; Zhou, Cao, Hu, and Zhao, p. 46, pl. 7, figs. 11, 12. 2001b Paraacidaspis? sp., Peng, Babcock, and Lin, p. 103, pl. 7, fig. 7. 2001b Paraacidaspis hunanica Egorova; Peng, Babcock, and Lin, p. 105, pl. 13, fig. 13; pl. 15, figs.

1977

9-11. Holotype. By monotypy; cranidium (Poletaeva, 1960, pl. 3, fig. 1; MGC 2604) from the Huaqiao

Formation, Tingziguan, Chatian, northwestern Hunan. The holotype and other specimens in the type suite of P. hunanica were refigured subsequently by Egorova, Xiang, Li, Nan, and Guo, 1963, pl. 12, figs. 6-8; and the holotype was refigured by Lu, Zhang, Zhu, Qian, and Xiang, 1965, pl. 130, fig. 15; Zhou, Liu, Meng, and Sun, 1977, pl. 81, fig. 2; Poletaeva, 1977b, pl. 1, fig. 13; and Zhang, 1990, pl. 1, fig. 6. New material. More than 30 sclerites, including cranidia, librigenae, and pygidia (illustrated

specimens 137713-137726) in collections ?P261.35, P316.0, P319.6, P331.8, P378.25, P[35.1, • 147•

PI323.5, PI327.2, PI356.5, PI360.3, PI370.7, and P[372. Remarks. A description of Paraacidaspis hunanica, translated from the original text of Egorova et al. (1963) was provided by Zhang (1990, p. 175, 176). Paraacidaspis hunanica varies somewhat in morphology. It shows variation in the pro-

portional length and the effacement of the glabella, the sinuousity of the axial furrows in the cranidium, the divergence of anterior branches of the facial suture, and in the degree of curvature of the posterior margin of the pygidium. An ontogenetic series shows that the glabella is more rounded anteriorly, and less tapered in the anterior half, in immature cranidia, but becomes truncated anteriorly and strongly tapered in the anterior half in large holaspids.

,

.#-

Text-figure 24. Reconstruction of cephalon and pygidium of Paraacidaspis hunanica Jegorova in Poletaeva, 1960. Cephalon based on specimens NIGP 137714, 137717, 137718 (see P1.46, figs. 2, 5, 6); pygidium based on specimens NIGP 137720, 137721 (see P1.46, figs. 8, 9). The smallest cranidium is about 3.4 mm, and shows a slightly wider palpebral area, a baccula that is poorly defined anteriorly, and a glabellar front that is separated from the paradoublural line. Fragmental librigenae attributed to P. hunanica are illustrated here (P1. 46, figs. 6, 13). They show the course of the facial suture, the wide lateral border, the distinct paradoublural line that continues the paradoublural line on the cranidium, the acutely pointed genal angle, and a wrinkled surface. Librigenae of this species are quite different from those of P. ultima from the Montagne Noire, southern France (Shergold et al., 2000, pl. 6, fig. 9). P. ultima has a wider genal field, and has a genal spine that is narrow at the base, and lacks a paradoublural line. Such great differences suggest that the librigenae from either northwestern Hunan or southern France do not belong to Paraacidaspis. Pygidia of P. hunanica are nearly indistinguishable from those of P. ultima. Specimens of the Chinese species are preserved in limestone. They are less effaced, and their axes appear to be • 148•

slightly more slender than the French specimens, which are preserved in shale. One pygidium (P1.46, fig. 10) is questionably assigned to P. hunanica. In general, it resembles other pygidia referred to this species, but the pleurae are less posteriorly diverted; it also has an entire posteriorly bowed rear margin, rather than an anteriorly bowed margin. Paraacidaspis latilimbus Yang, 1978; Paraacidaspis sp. sensu Yang, 1978, and P. sp. sensu Ergaliev, 1980 are regarded as synonyms of P. hunanica because they all fall within the range of morphological variation of P. hunanica. P. latilimbus is based on a single cranidium (Yang, 1978, pl. 13, fig. 15) from the Huaqiao Formation at Huaqiao, Baojing, northwestern Hunan. Zhang (1990, p. 177) considered it as valid, but noted that the species agrees with the type species in all respects except for having a slightly shorter glabella. Our new material reveals that this difference is no more than intraspecific variation within P. hunanica. Paraacidaspis sp. sensu Yang (1978, pl. 13, fig. 16) was based on a single cranidium from the same zone and stratigraphic section as P. hunanica. It is a rather fragmental specimen, but apparently identical in all observed respects with the type species. Paraacidaspis sp. sensu Ergaliev (1980, pl. 14, fig. 21) was based on a single cranidium from the equivalent of post-Paibian Stage strata in southern Kazakhstan. It is apparently a juvenile cranidium, and it closely resembles cranidia of similar size in the new ontogenetic series of P. hunanica. In Kazakhstan, P. hunanica ranges up through the Neoagnostus quadratiforms Zone (Ergaliev, 1980). This represents the youngest known occurrence of the type species.

Occurrence. The holotype is from the Huaqiao Formation, Tingziguan, Chatian, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it is questionably associated with trilobites indicative of the Wanshania wanshanensis Zone and definitely associated with trilobites indicative of Liostracina bella Zone through the Shengia quadrata Zone (equivalent to the uppermost of the Lejopyge laevigata Zone through the Linguagnostus reconditus Zone). Paraacidaspis sp. Plate 46, figure 17

Material. One pygidium, INGP 137727, in collection P[360.3. Remarks. A single, incomplete pygidium is similar to pygidia assigned to Paraacidaspis hunanica or to P. ultima in features such as the shape and length of the axis, the nature of the border, and segmentation of the pleural field with narrow, outward-expanding anterior bands and wide posterior bands. Granulation of the specimen, however, suggests that it differs from all described species. It is too fragmentary for meaningful description at the present time. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi-2 section, Hunan, where it is associated with trilobites indicative of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone).

• 149.

Order Superfamily Family Genus

ASAPHINA

Salter, 1864

ANOMOCARAROIDEA ANOMOCARIDAE AFGHANOCARE

Poulsen, 1927

Poulsen, 1927

Wolfart, 1974

Afghanocare Wolfart, 1974, p. 101, 102; Fortey, 1994, p. 35" Yuan and Yin, 1998, p. 164; Jell and Adrain, 2003, p. 356. Paranomocare Lee and Yin (Li and Yin), 1973, p. 28; Zhou, Liu, Meng, and Sun, 1977, p. 175" Yin and Li, 1978, p. 495" Jell and Adrain, 2003, p. 420. Type species. Afghanocare angustegenatum Wolfart, 1974 (p. 103-106, pl. 13, figs. 7-11; pl. 14, figs. 1-8; pl. 15, figs. 1-3, text-fig. 12) from the upper Cambrian, Surkh, Bum, central Afghanistan; by original designation. Remarks. As noted by Fortey (1994), Afghanocare is similar to Changshanocephalus Sun, 1935 from North China, and may eventually prove to be synonymous with it. The type species of Changshanocephalus is C. reedi Sun (1935, pl. 1, fig. 25; also Lu et al., 1965, pl. 29, fig. 8), which is based on a single, poorly illustrated cranidium. With that illustration, the concept of the genus became muddled. Several species have been referred to Changshanocephalus (Kobayashi, 1960b, 1962; Lu et al., 1965; Schrank, 1975; Qian, 1994), but apparently not all of them are congeneric. For example, specimens attributed by Qian (1994) to C. conica (Endo, 1944) and C. subconica Qian are characterized by having short anterior areas and short, anteriorly located palpebral lobes; whereas the anterior area of C. majus (Dames, 1883), as revived by Schrank (1975, p. 598, pl. 5, figs. 6-9; pl. 6, figs. 1-7), is broad and concave, and the palpebral lobe is long and located posteriorly. Features shown by C. majus seem more conformable to Sun's (1935) original diagnosis of Changshanocephalus. The species is closely comparable and probably congeneric with the type species of Afghanocare and the material referred here to Afghanocare. Pending further investigation of the type species of Changshanocephalus, Afghanocare should be regarded as a valid genus. Paranomocare Lee and Yin (Li and Yin, 1973, from eastern Guizhou and western Hunan, is regarded as a junior synonym of Afghanocare. The valid date of publication should be 1973, rather than 1978 as indicated by Jell and Adrain (2003). The genetic names and its type species P. guizhouensis (Li and Yin 1973, p. 28, 29, pl. 1, figs. 1, 2) were first published with adequate descriptions and illustrations, in a widely distributed journal in 1973. Except for the type species, Li and Yin described five additional species, among which four were refigured by Yin and Li (1978). Although the type species shows clear furrows, the genus bears no significant differences from Afghanocare in general respects of the cranidium and pygidium. By suppressing Paranomocare, its type species Paranomocare guizhouensis becomes a senior synonym of Afghanocare guizhouense Yuan and Yin (1998, p. 164, pl. 5, fig. 17; pl. 6, figs. 1-4). Afghanocare truncatum (Peng, 1987) Plate 47, figures 1-15; Plate 51, figures 13-15 1987 Hunanaspis? truncatus Peng, 1987, p. 98, 99, pl. 7, fig. 4, text-fig. 12. 1998 Afghanocare guizhouense Yuan and Yin (in part), p. 164, 165, pl. 6, figs. 1-4, non pl. 5, fig. • 150.

17.

Holotype. By monotypy; cranidium (Peng, 1997, pl. 7, fig. 4; NIGP 74519), from the Liostracina bella-Ammagnostus sinensis [=Proagnostus bulbus] Zone, Huaqiao Formation, Taoyuan, northwestern Hunan. New material. More than 10 sclerites, including cranidia and pygidia (illustrated specimens NIGP 137728-137735, 137778), in collections P319.6, P374.9, P378.25, and W227. Remarks. This species, based on a single cranidium from the Huaqiao Formation in northwestern Hunan, is transferred to Afghanocare. Originally it was assigned to Hunanaspis because of its similarity to H. gracilis Zhou (in Zhou et al., 1977, p. 176, 177, pl. 62, fig. 6), the type species of Hunanaspis, which is also based on a single cranidium from the Huaqiao Fornation in western Hunan. The assignment was not satisfactory because Hunanaspis is characterized by having an anterior area of the fixigena that bears a gently convex anterior field, a clearly defined anterior border and a pitted anterior border furrow. With a pitted anterior border furrow, Hunanaspis seems more comparable to some eulomoids (e.g., Stigmatoa Opik, 1963) than to anomocaroids. H. truncatus lacks a row of pits in the anterior border furrow, and its anterior border is not convex as in Hunanaspis, but is flat and upturned. New material from the Huaqiao Formation at Paibi and Wangcun, western Hunan, include several cranidia and pygidia. The cranidia are indistinguishable from the holotype cranidium of Afghanocare truncatum, suggesting an assignment to this species. Afghanocare guizhouensis sensu Yuan and Yin (1998, pl. 6, figs. 1-4) is here suppressed as a junior subjective synonym of A. truncatum because both have identical morphologies, are of similar age, and are from the same paleogeographical region. Both A. guizhouensis and Hunanspis ? trancatus are from the traditional "early Late Cambrian", and the discovery of new material extends the stratigraphic range of the species upwards into the Glyptagnostus reticulatus Zone (base of the Furongian Series). Afghanocare truncatum differs from Afghanocare angustegenatum, the type species, by having a longer preglabellar area, a truncate rather than rounded glabellar front, and more effaced pleural regions in the pygidium. Afghanocare hubeiense (Zhu and Sun in Zhou et al., 1977, p. 176, pl. 52, figs. 1, 2) from western Hubei is similar in glabellar shape, concavity of the anterior area, and divergence of the anterior branches of the facial suture, but it can be differentiated from A. truncatum in having a slightly shorter preglabellar area and more pairs of incised pleural furrows in the pygidium. A. hubeiense is from the traditional "Middle Cambrian" Guanzhuling Formation and is older than A. truncatum. A. truncatum can be easily differentiated from Afghanocare lategenatum Wolfart, 1974 because A. lategenatum has a subconical, rather than anteriorly truncated, glabella. Afghanocare lategenatum is probably conspecific with Paranomocare songtaoense Lee and Yin, 1973 (Li and Yin, p. 29, pl. 1, figs. 7, 8) from eastern Guizhou and Paranomocare bagushangense Zhu and Sun (in Zhou et al., 1977, p. 175, 176, pl. 52, figs. 3, 4) from western Hubei, both of which are also characterized by having a subconical glabella. If so, P. songtaoense has priority over the other two species names. Occurrence. The holotype is from the Liostracina bella-Proagnostus bulbus Zone, Huaqiao Formation, Taoyuan, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone through the lower part of the Chuangia subquadrangulata Zone (equivalent to the Linguagnostus reconditus Zone and the Glyptagnostus reticulatus Zone). • 151•

Genus GLYPHASPELLUS Ivshin, 1953

Type species. Glyphaspellus primus Ivshin, 1953 (p. 144-146, pl. 10, figs. 1-5) from the Anomocare-Phoidagnostus [=Glaberagnostus] bituberculatus Zone, Boshekuly, Kazakhstan; by original designation.

Glyphaspellus? sinensis sp. nov. Plate 5 l, figures 11a, 12 2001b

Undetermined cranidium, Peng, Babcock, and Lin, p. 104, pl. 9, fig. 14.

Etymology. From Latin, Sinae, China, referring to China. Holotype. By monotypy; cranidium in collection (P1.51, figs. 11, 12; NIGP 137777), P277. Diagnosis. Glyphaspellus? with preglabellar area about half length of glabella excluding occipital ring; glabella constricted slightly at sides, truncated anteriorly, with four pairs of lateral furrows, S 1 long, weakly bifurcated; palpebral lobe located posteriorly, with anterior and posterior ends close to axial furrow; occipital furrow gently sinuous, deep at sides.

Description. Cranidium subrectangular; width at anterior margin about three-fourths cranidial length. Glabella trapezoidal, with sides gently constricted inward, truncate anteriorly; with four pairs of lateral furrows: S1 long and deep, directed diagonally; $2 shallow, directed inward and slightly rearward; $3 shallow, directed slightly forward; $4 faint, directed inward and slightly forward; occipital furrow transverse, deep at sides; occipital ring narrow (sag.), half as long as L1 lobe, bearing tiny, medially placed node. Preglabellar area wide (sag., tr.) with moderately convex anterior border, narrowing gently abaxially; anterior border furrow shallow anteriorly, gently forward-arched posteriorly. Palpebral lobe gently arcuate, moderately elevated, lying close to axial furrow, with anterior end opposite midpoint of L4, posterior end opposite midpoint of L1; palpebral furrow shallow; palpebral area narrow (tr.), gently convex, about one-third as wide as glabella. Anterior branch of facial suture diverging forward at angle of 90 ° to posterior transverse furrow on anterior area of fixigena, then curving gradually inward to enclose a rounded anterolateral comer of cranidium. Posterior branch and posterolateral projection unknown.

Remarks. According to Ivshin (1953, p. 142), Glyphaspellus is characterized by its distinctive structure in the anterior field of the fixigena: it bears a transverse, auxiliary furrow that divides the anterior field into a flat or slightly concave posterior part and a raised, forwardly inclined anterior part. The anterior border is flat and forwardly deflected, defined clearly by the anterior border furrow. The holotype cranidium of G. ? sinensis has such a morphology in the anterior area of the fixigena. It is questionably attributed to Glyphaspellus, however, because the anterior end of the palpebral is so close to the axial furrow that an eye ridge is almost absent, and the anterior branch of the facial suture extends nearly from the axial furrow. These features, as well as the laterally constricted glabella, serve to differentiate this species from all other species assigned to Glyphaspellus. In addition, the glabella of G. ? sinensis has constricted flanks and bears four pairs of lateral furrows, with S 1 being notably long. • 152•

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone).

Genus

PARACOOSIA

Kobayashi, 1936

Paracoosia Kobayashi, 1936, p. 170, 172; 1944, p. 136; 1960a, p. 248; Hupr, 1953b, p. 212; 1955, p. 189; Howell in Moore, 1959, p. 0288; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 175; Chernysheva, 1960, p. 94; Kraskov, Lararenko, Oginko, and Chernysheva, 1960, p. 232; Opik, 1967, p. 225; Luo, 1974, p. 638; Egorova, Pegel, and Chernysheva, 1982, p. 95; Wittke, 1984, p. 121,122; Fortey, 1994, p. 35; Gogin and Pegel, 1997, p. 120, 121; Jell and Adrain, 2003, p. 418. Type species. Coosia asiatica Mansuy, 1916 (p. 40-42, pl. 7, figs. 6a-h; Text-fig. 25 herein), from the middle Cambrian (horizon of Annamitia sinifera), Penkai, northwestern Vietnam; by original designation. Other species. Coosia deprati Mansuy, 1915 (p. 27, 28, pl. 3, figs. 4a-g) from the middle Cambrian of Haut-Tonkin, Vietnam; Coosia sukhanica Chernysheva in Kraskov et al., 1960 (p. 230-232, pl. 52, figs. 1, 2) from the lower part of the upper Cambrian, middle reaches of the Olenek River, Siberia, Russia; Paracoosia pulchra Chernysheva (1960, p. 232, 233, pl. 53, figs. 7, 8) from the upper part of the Mayan Stage, middle reaches of the Aldan River, Siberia, Russia; Paracoosia aspis Opik, 1967 (p. 225, pl. 31, figs. 3a, b) from the Glyptagnsotus stolidotus Zone, O'Hara Shale, Queensland, Australia; Paracoosia mirzzadi Wolfart, 1974 (p. 110, 111, pl. 16, figs. 7, 8; pl. 17, figs. 1-3) from the lower upper Cambrian, central Afghanistan; Paracoosia kingi Wittke, 1984 (p. 16, 17, pl. 2, figs. 11-16) from the Drepanura-Torifera-Eokaolishania Zone and the ProchuangiaParacoosia Zone, Mira Formation, northern Iran; Paracoosia cf. P. mirzzadi Wolfart, 1974 (Fortey, 1994, p. 35-37, figs. 5A-C) from the upper Cambrian of Oman; Coosia suchovi Egorova, 1984 (p. 21, pl. 5, fig. 1) from Siberia, Russia; Paracoosia ditigenta Gogin and Pegel, 1997 (p. 121, 122, pl. 27, figs. 3, 4, 6) from the Maspakites Zone, Usti-Maiskaya Formation, Aldan River, Siberia, Russia; Paracoosia cf. P. kingi Wittke, 1984, from the Huaqiao Formation of western Hunan, and Paracoosia huayuanensis sp. nov. from the Huaqiao Formation of western Hunan. Remarks. Kobayashi (1936) erected Paracoosia to embrace two species, Coosia asiatica Mansuy, 1916 and Pterocephalus asiaticus Walcott, 1905. In so doing, the two species became homonyms, and Pterocephalus asiaticus holds priority over Coosia asiatica. Kobayashi (1936) renamed C. asiatica as Paracoosia mansuyi and selected it as the type species of Paracoosia. Subsequent to Kobayashi's (1936) work, the name P. mansuyi has been used by most authors except Opik (1967) as a replacement for Coosia asiatica and as the name of the type species of Paracoosia. Opik (1967, p. 226) noted there are significant differences between Coosia asiatica Mansuy and Pterocephalus asiaticus Walcott, including a baccula-bearing glabella and a pygidium with denticulate spines in P. asiaticus, and concluded that C. asiatica Mansuy and P. asiaticus represent separate genera. Following reexamination of Walcott' s (1905, 1906, 1913) collections from China, Zhang and Jell (1987, p. 192, pl. 84, fig. 1; pl. 85, figs. 7, 8) reillustrated the type material of Pterocephalus asiaticus, and concluded that P. asiaticus should be transferred to Monkaspis. That assignment is followed here, as is Opik's (1967) genetic concept. Thus, Paracoosia mansuyi should be abandoned as a junior objective synonym of Coosia asiatica Mansuy, and the valid name of the type species of Paracoosia should be Paracoosia asiatica (Mansuy). • 153-

Text-figure 25. Lectotype of Coosia asiatica Mansuy, the type species of Paracoosia Kobayashi, 1938; original of Mansuy, 1916, pl. 7, fig. 6c, x 3.1. The slab is in the collection of the Department of Earth Sciences, Universit6 Claude Bernard, Lyon, France. A single name-bearing specimen had not been selected previously for Coosia asiatica. As illustrated, Mansuy's (1916, pl. 7, figs. 6a-h) syntypes of Coosia asiatica include four incomplete exoskeletons, two cranidia, and a pygidium showing doublure. One of us (SP) attempted to examine the type material at several institutions in France, but found that only one exoskeleton (Mansuy, 1916, pl. 7, fig. 6c; Text-fig. 25 herein) is accessible in the collection of the Department of Earth Sciences, Universit6 Claude Bernard. The others seem to be missing. The remaining specimen is the most complete exoskeleton among Mansuy's (1916) syntypes, and is here selected as the lectotype of Paracoosia asiatica (Mansuy). Paracoosia sp. cf. P. kingi Wittke, 1984

Plate 48, figures 1, 2; Plate 50, figures 1-15; Text-figure 26 1963 1965 1978 cf. 1984 ?1991

Coosia sp.; Egorova, Xiang, Li, Nan, and Guo, p. 49, pl. 11, fig. 15. Coosia sp.; Lu, Zhang, Zhu, Qian, and Xiang, p. 362, pl. 66, fig. 28. Metanomocare? sp.; Yang, p.78, pl. 5, figs. 12-14. Paracoosia kingi Witteke, p. 123, pl. 2, figs. 11-16. Coosia sp. 1; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 141, 142, pl. 13,

figs. 12-14. • 154.

?1991 ?1993 ?1993 2001 b

Coosia sp. 2; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 142, pl. 13, fig. 15. Coosia sp. 1; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 141, 183, pl. 13, figs. 12-14. Coosia sp. 2; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 183, 184, pl. 13, fig. 15. Pterocephalid gen. et sp. nov. 1, Peng, Bobcock, and Lin, p. 102, 103, pl. 5, fig. 6; pl. 7, figs. 10, 11; pl. 9, figs. 5, 6.

Holotype of Paracoosia kingi. Cranidium (Wittke, 1984, pl. 2, fig. 11; GPIBo/Wi. 72) from the Ablaz Mountains, northern Iran. Material. More than 20 sclerites, including cranidia, librigenae, hypostomes, and pygidia (illustrated specimens NIGP 127736, 137759-137771), in collections P260, P261.35, P295.13, W195.7, W196.3, W199.2, and W212.45.

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Text-figure 26. Reconstruction of cephalon and pygidium of Paracoosia sp. cf. P. kingi Wittke, 1984. Cephalon based on specimens NIGP 137763, 137764 (see P1. 50, figs. 5, 6)" pygidium based on specimen NIGP 137769 (see P1.50, fig. 13).

Remarks. Material from the Huaqiao Formation of western Hunan is closely comparable with that from the Ablaz Mountains of Iran. The large pygidia in the new collections have long, incised, and oblique pleural furrows, and thin and long axes with slightly concave flanks and nine to ten tings. In these respects, the material is almost identical to Paracoosia kingi. Associated cranidia also conform in general respects to the holotype cranidium from Iran, particularly in such characters as the broad conical glabella, the relative wide palpebral area, and the long, moderately defined eye •

155.

ridge. However, the Hunan specimens show some differences from the holotype. They have a proportionally longer glabella that occupies more than half of the cranidial length, weak lateral glabellar furrows, and a somewhat differently shaped anterior branch of the facial suture. In the Hunan material, the anterior branch extends forward and outward until it nears the anterior margin, where it then curves sharply inward. In Paracoosia kingi, the glabella is effaced, and is less than half of the cranidial length; the anterior branch of the facial suture is gently and smoothly curved. A small, possible meraspid, pygidium in the Hunan material (P1.48, figs. 1, 2) is different from the small pygidia assigned to P. kingi by Wittke (1984, pl. 2, figs. 15, 16). The Iranian pygidia have a short and wide axis, and are probably not conspecific with P. kingi. Examination of material assigned as Coosia sp. by Egorova et al. (1963) and later assigned to ? Metanomocare sp. by Yang (1978) show that they are conspecific with Paracoosia sp. cf. P. kingi. Both species have almost the same occurrence in the Huaqiao Formation with the new material, and they are from the same paleogeographic region in western Hunan (?Metanomocare sp. and Coosia sp.) and eastern Guizhou (?Metanomocare sp.). The strongly deformed material assigned as Coosia sp. 1 and Coosia sp. 2, both figured by Yang et al. (1991, 1993), belongs to Paracoosia and also may be conspecific with P. sp. cf. P. kingi.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone (equivalent to the lower part of the Proagnostus bulbus Zone through the lower part of the Linguagnostus reconditus Zone). Paracoosia huayuanensis sp. nov. Plate 48, figures 3-16 2001b Pterocephalid gen. et sp. nov. 2, Peng, Babcock, and Lin, p. 104, pl. 10, figs. 9, 10.

Etymology. From Huayuan County, from where most known material of the species was collected. Holotype. Cranidium (P1.48, fig. 11, NIGP 137743) in collection P319.8. Other material. Five cranidia and four pygidia (NIGP 137737-137742, 137744, 137745), in collections P317.4, P319.6, P319.8, P337.5, and W227. Diagnosis. Paracoosia with subtrapezoidal or elongate-subconical glabella less than half cranidial length, and bearing weak carina; lateral glabellar furrows consist of three weak pairs; palpebral area about half as wide as glabella between palpebral lobes. Pygidium with axis occupying two-thirds of pygidial length; pleural region rather effaced, relatively wide, and weakly segmented; pleural furrows short. Description. Anterior area of fixigena long, gently concave and upturned slightly anteriorly; some specimens show short, obscure anterior border; cranidium longer than wide. Glabella gently convex, tapering forward gently, obtusely rounded anteriorly, occupying half or less of cranidial length, with weak carina and three pairs of lateral furrows; S 1 weakly impressed, $2 and $3 faint; occipital furrow distinct, bowed gently rearward or transverse; occipital ring of uniform width, flat in profile, bearing weak median node located forward of midlength of occipital ring; palpebral long, • 156.

semicircular, lying opposite midpoint of glabella, with posterior end slightly anterior of occipital furrow and anterior end at three-fourths glabellar length; eye ridge short, weak, diverging posteriorly about 60 ° to sagittal line. Anterior branch of the facial suture diverging strongly forward from anterior end of palpebral lobe, curving inward evenly, enclosing anterior area of fixigena by smooth curvature; posterior branches transverse, enclosing narrow belt-like posterolateral projection; posterior border furrow weak; posterior border a linear ridge. Pygidium semicircular, length two-thirds width, with transverse anterior margin and rounded anterolateral comers; facet sloping slightly forward and outward. Axial subcylindrical, relatively wide, tapering gently rearward, occupying about two-thirds pygidial length; ring furrows shallow on anterior two or three segments, becoming progressively weak or obscure rearward. Pleural field convex, largely effaced, with two to four fibs, first rib well defined, and others weak; borders about as wide as pleural field, flat, uniform in width. Cranidial surface smooth; pygidial surface with fine terrace lines on axis, pleural fields, and borders.

Remarks. The new species most closely resembles Paracoosia kingi but differs in having a less tapered, less acutely rounded glabella, a relatively wider palpebral area that is about half the glabellar width, a weaker eye ridge, and an effaced pygidium with a shorter and wider axis and shorter pleural furrows. Paracoosia asiatica (Mansuy), the type species of Paracoosia, can be easily differentiated from P. huaynanensis by its broad-conical glabella and its less effaced pygidium with longer pleural furrows. Paracoosia diligenta Gogin and Pegel, 1997 is also closely comparable to the new species fron Hunan, but the Siberian species has a more rounded glabella anteriorly, an obliterate palpebral furrow, a smaller palpebral lobe, and a much wider pygidium. Paracoosia mizzadi Wolfart has a subquadrate cranidium with a more tapered glabella and a much wider palpebral area. These characters serve to easily differentiate P. mizzadi from the new species. The effaced pygidium of the new species is different from all others species of Paracoosia. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Genus PAIBIANOMOCAREgen. nov.

Etymology. From Paibi, a village in Huayuan County, western Hunan, combined with the generic name Anomocare. Type species. Paibianomocare paibiense gen. et sp. nov. Diagnosis. Anomocarid with relatively short preglabellar field; glabella relatively large, tapered forward, weakly furrowed, occupying three-fourths cranidial length; anterior margin of cranidium gently rounded, anterior border upturned, poorly defined; palpebral lobe medium to rather long in size, opposite midpoint of glabella; palpebral area narrow; pygidium subovate with relatively wide and short axis slightly longer than half pygidial length; pleural furrows of two to three pairs, weakly defined; borders wide, gently concave. Discussion. This genus is characterized by having a relatively large glabella with weakly defined lateral furrows, a relatively short preglabellar field, a pygidium with a relatively wide and short axis, • 157.

and rather effaced pleural regions. The cranidium is generally of anomocarid aspect, but the proportion of the preglabellar field and the glabella bear similarity with aphelaspids. The pygidium, however, shows a clear relationship of this species to anomocarids. The new genus resembles most closely Glyphaspellus Ivshin, 1953 from the middle Cambrian of Kazakhstan, and it resembles especially its type species Glyphaspellus primus Ivshin (1953, p. 144-146, pl. 10, figs. 1-5). The new genus differs from Glyphaspellus in having a more tapered, anteriorly rounded rather than truncate glabella, more widely divergent anterior branches of the facial suture, and in lacking a tropidium on the anterior area of the cranidium. In the Kazakhstan species, the anterior border is clearly defined, the palpebral lobes are longer, defined by deep and wide palpebral furrows, and the preglabellar field is not depressed as it is in the new genus. Afghanocare Wolfart, 1974 from central Afghanistan has a similarly depressed anterior area of the cranidium, but the Afghanistan genus has a much longer preglabellar field, and a proportionally shorter glabella. Guizhouanomocare Lee (in Yin and Li, 1978), from the middle Cambrian of central Guizhou, resembles the new genus in the nature and the proportion of the anterior area, the relatively short pygidial axis, and the narrow, posteriorly effaced pleural region of the pygidium, but differs in having large palpebral lobes that are more widely spaced. Yingziaspis Zhang, 1999 (type species Y. erdaogouensis from Liaoning, northeast China), is also comparable to Paibianomocare. Y. erdaogouensis is based on three cranidia, among which the holotype cranidium (Zhang, 1999, pl. 1, fig. 17) is rather distorted, which prevents a precise comparison. The paratype cranidia, however, are closely similar, differing only in the presence of a frontal spine on the anterior cranidial border, and in the less converging anterior branches of the facial suture. Protapatokephalus Ergaliev, 1980 from southern Kazakhstan, which is based on a single cranidium (Ergaliev, 1980, pl. 17, fig. 14), has a palpebral lobe of similar shape and position, and has a glabella of similar size, but the Kazakhstan genus can be differentiated by having less divergent anterior branches of the facial suture, and, therefore, a narrower (tr.) anterior area that bears a less curved anterior border furrow, a flat rather than depressed preglabellar area, and a gently convex rather than upturned anterior border.

Paibianomocare paibiense gen. et sp. nov. Plate 49, figures 1-14 1998 Palemansuyia prima Yuan and Yin (in part), p. 168, pl. 6, fig. 7 only. 2001b Proasaphiscid? gen. et sp. nov., Peng, Babcock, and Lin, p. 103, pl. 8, figs. 14, 15. 2001b Ptychoparioid gen. et sp. nov., Peng, Babcock, and Lin, p. 105, pl. 15, fig. 2.

Holotype. Cranidium (P1 49, fig. 9; NIGP 137754) in collection P277. Other material. Twelve cranidia, and three pygidia (illustrated specimens NIGP 137746-137753, 137755-137758) in collections P260, P269, P273.8, P277, P376.4, P134.3, and W227.

Diagnosis. Paibianomocare lacking paradoublural line on anterior area of fixigena; palpebral lobe moderately large, lying posterior of cranidial midpoint; anterior border furrow shallow, obscure; anterior border upturned but flexed slightly downward medially.

Description. Cranidium subquadrate with anterior margin arched gently forward. Glabella relatively large, gently convex, trapezoidal or broadly conical in outline, with three pairs of faint lateral furrows, obtusely rounded anteriorly, occupying three-fourths to four-fifths cranidial length; sides •

158.

straight, slightly concave or slightly convex outward; S 1 diagonally, obscurely bifurcated; $2 and $3 faintly impressed; occipital furrow shallow, widening somewhat at sides; occipital ring widest sagittally, narrowing abaxially, bearing tiny, centrally located median node. Palpebral lobe gently curved, situated opposite midpoint of glabella, defined by shallow palpebral furrow; palpebral area narrow, inclined steeply to axial furrow, width less than one-fourth that of glabella between palpebral lobes. Preglabellar area relatively short, sloping gently forward, preglabellar field gently depressed; anterior border furrow shallow, obscurely defined; anterior border upturned gently but flexed slightly downward and forward medially to form narrow (sag., exs.) and wide, slightly convex belt-like area. Anterior branch of facial suture diverging forward at angle of 800-90 ° onto anterior border furrow, then turning inward gently after crossing anterior border furrow to cut anterior border obliquely and meet anterior margin at point opposite lateral margin of occipital ring; posterior branch diverging strongly rearward about 160 °, enclosing wide (tr.), blade-like or subtriangular posterolateral projection; posterior border wider (exs.) than fixigenal field on posterolateral projection, gently convex, defined by deeply incised posterior border furrow. Pygidium subovate in outline, slightly longer than wide; axis relatively short and wide, obtusely rounded posteriorly, occupying about two-thirds of pygidial length; articulating half-ring narrow (sag.) and long (tr.), bar-like, defined by linear, shallow articulating furrow; axis with three to four faint tings and possible long terminal piece; pleural field narrow, gently convex, bearing two to three fibs; borders gently concave, width greater than pleural field. Cranidial surface nearly smooth, lacking ornamentation; pygidial surface with terrace lines on borders.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2, and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone through the Chuangia subquadrangulata Zone (equivalent to the Proagnostus bulbus Zone through the Glyptagnostus reticulatus Zone). Paibianomocare lineatum gen. et sp. nov. Plate 51, figure 5

Etymology. From Latin, lineatus, line-bearing, referring to the paradoublural line-bearing cranidium. Holotype. One cranidium, NIGP 137774, in collection P135.1. Diagnosis. Paibianomocare having paradoublural line on anterior area of fixigena; relatively large palpebral lobe lying slightly posterior to midpoint of glabella; anterior border furrow obscure; anterior border thin, upturned. Description. Glabella subcylindrical in outline, gently tapering forward, with nearly straight sides and subangular front, occupying about five-sixths cranidial length; S 1 faint, $2 short and obscure; occipital furrow transverse, moderately deep; occipital ring widest sagittally, narrowing abaxially. Palpebral lobe rather long (sag.), wide, defined by rather deep palpebral furrow, palpebral area nearly half as wide as glabella between palpebral lobes, inclined steeply to axial furrow. Paradoublural line shallow, broadly arched forward, lying immediately before preglabellar furrow, dividing anterior area of fixigena into a flat posterior portion and an anterior portion that is slightly convex immediately in front of paradoublural line, becoming forward-sloping and continuing as shallow, poorly defined anterior border furrow and upturned anterior border. Anterior branch of •

159

•

facial suture diverging forward at angle of about 110 ° onto anterior border furrow, then turning forward and inward after crossing anterior border furrow to cut anterior border obliquely and meet anterior margin at point opposite to the anterolateral comer of glabella; posterior branch diverging strongly rearward at 160 °, enclosing narrow (exs.), blade-like posterolateral projection; posterior border rather convex, wider than fixigenal field on posterolateral projection; posterior border furrow clearly defined. Thorax and pygidium unknown.

Remarks. At first glance, the new species is quite similar to Glyphaspellus primus. The ridge-like convex region, defined posteriorly by a transverse furrow, on the anterior field of the fixigena is easily confused with the tropidium of Glyphaspellus primus. However, varied the structure on the anterior area of the fixigena in the new species seems to be a paradoublural line rather than a tropidium. The new species is referred to Paibianomocare because it is more closely comparable in morphology to P. paibiense gen. et sp. nov. than it is to any species of Glyphaspellus. The new species has a proportionally larger glabella, lacks a clearly defined anterior border, has an anterior border that is upwardly flexed and defined by a shallow, poorly defined anterior border furrow. P. lineatum can be differentiated easily from P. paibiense principally by having a transverse furrow anteriorly and a longer palpebral lobe. P. lineatum also seems to have a proportionally longer and less tapered glabella.

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi-2 section, Hunan, where it is associated with trilobites indicative of lower part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). Family PTEROCEPHALIIDAE Kobayashi, 1935 Subfamily CILIINAE Peng, 1992 Peng (1992, p. 63) erected this subfamily to accommodate Cilia Peng, 1992, Proapatokephaloides Ergaliev, 1980, and Mecophrys Shergold, 1982. Yangweizhouia Yuan and Yin, 1998, which was originally classified within the Dokimocephalidae, also belongs to the subfamily. This subfamily includes pterocephaliid trilobites characterized by an elongate cranidium, a long frontal area of the cranidium with a plectrum-bearing anterior border, a forward-tapered, rounded to obtusely rounded glabella, rather long, posteriorly located palpebral lobes, narrow palpebral areas, and a laterally narrowing occipital ring that bears a median node. The anterior branch of the facial suture diverges rather strongly forward initially, and then meets the anterior cranidial margin at or near the sagittal line. The pygidium, known only in Mecophrys and Yangweizhouia, is relatively small, transverse or semicircular, with obtusely rounded anterolateral comers and a short axis beating two or three tings and reaching nearly to the posterior border furrow. With the inclusion of Yangweizhouia in the Ciliinae, the observed range of the subfamily is extended somewhat downward. Now the subfamily has a total range from the Linguagnostus reconditus Zone to the Lotagnostus punctatus Zone, and has an age of latest Wulingian (South China terminology) to middle Furongian.

• 160.

Genus

YANGWEIZHOUIA

Yuan and Yin, 1998

Type species. Yangweizhouia carinata Yuan and Yin, 1998 (p. 156, 157, pl. 4, figs. 1-6; pl. 5, figs. 15, 16), from the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou; by original designation. Remarks. This genus is monospecific, and the genetic concept of Yuan and Yin (1998, p. 155, 156) is followed here. So far the genus is known only from the western Hunan-eastem Guizhou region. It is most similar to Cilia in the nature of the frontal area, the course of the facial sutures, and the shape and size of the palpebral lobes, but it differs in having a proportionally larger glabella with an acutely rather then obtusely rounded front, and a more effaced occipital furrow. In glabellar shape, and the course of the facial sutures, Yangweizhouia is similar to Mecophrys Shergold, 1982, but Mecophrys has a much wider (sag.) preglabellar field, a larger palpebral lobe, and a relatively wider (tr.) palpebral area. Yuan and Yin (1998) compared Yangweizhouia with Chalfontia Shergold, 1982 and Prismenaspis Henderson, 1976, both of which are based on species attributed to Prismenaspis by Henderson (1976). Both taxa can be differentiated from Yangweizhouia by having deep and broad axial and occipital furrows, a shorter preglabellar field, a strongly inward-sloping palpebral area, and a subcentrally located, well-defined palpebral lobe. These differences are sufficiently strong to suggest that Chalfontia and Prismenaspis are not related to Yangweizhouia but belong to a different lineage. Compared to Yangweizhouia, Abharella Wittke from the Mila Formation in the Alborz Mountains, northem Iran (Wittke, 1984) and the Val d'Homs Formation in the Montagne Noire, southern France (Shergold et al., 2000) has a similar wide (sag.), plectrum-bearing anterior border, similarly shaped glabella, and similarly located palpebral lobes. In addition, the occurrence of Yangweizhouia in western Hunan falls well within the stratigraphic range of Abharella in Iran, as documented by Wittke (1984). Abharella, however, is readily differentiated by having a large palpebral area of the fixigena, a longer preglabellar field, and a relatively longer palpebral lobe.

Yangweizhouia carinata Yuan and Yin, 1998 Plate 51, figures 6-10 1998 Yangweizhouiacarinata Yuan and Yin, p. 156, 157, pl. 4, figs. 1-6; pl. 5, figs. 15, 16.

Holotype. Cranidium (Yuan and Yin, 1998, pl. 5, fig. 15; NIGP 127937) from the Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou.

New material. Two cranidia, NIGP 137775, 137776, in collections W227 and P301.9. Remarks. The type material (Yuan and Yin, 1998, pl. 4, figs. 1-6; pl. 5, figs. 15, 16) from eastern Guizhou shows that the glabellar shape is variable. In some specimens, including the holotype, the glabella is unevenly tapered, with the anterior half being more strongly tapered than the posterior half, leading to a pyriform shape; it also has a-more acutely rounded glabellar front. In other specimens, however, the glabella is evenly tapered forward and the glabellar front is broadly 161. •

rounded. Additionally, the fine, curved ridge connecting the anterolateral comer of the glabella and the 13 point of the facial suture on the preocular field of the fixigena is variable; and the glabellar and occipital furrows are variably effaced. A pair of baccula-like raised structures are consistently developed on the posterior part of the palpebral area of the fixigenae. The new material from western Hunan shows an evenly tapered, largely effaced, anteriorly rounded glabella and faint curved ridges on the preocular fields. The specimens are similar to others from the same paleogeographic region in eastern Guizhou. The key characters that warrant assignment of specimens to this species include posteriorly located palpebral lobes; narrow palpebral areas showing ovate convexity posteriorly; divergent anterior branches of the facial suture that enclose a wide (sag., exs.) frontal area; fine, curved ridges on the preocular areas; a convex anterior border with a short plectrum; and a long (tr.) blade-like posterolateral projection.

Occurrence. The holotype is from the Hadragnostus modestus-Blackwelderia Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone). Superfamily ASAPHOIDEASalter, 1864 Family

CERATOPYGIDAE

Linnarsson, 1869

Subfamily CERATOPYGINAELinnarsson, 1869 Genus PROCERATOPYGEWallerius, 1895

Type species. Proceratopyge conifrons Wallerius, 1895 (p. 56, 57, pl. 1, figs. 6a, b; Westerggtrd, 1948, p. 5, 6, pl. 1, figs. 7-16 ), from the upper part of the Lejopyge laevigata Zone, Gudhem, Falbygden area, V~isterg6tland, Sweden; by original designation.

Other species. Species were listed by Rushton (1983, p. 131, 132), Lee and Choi (1995, p. 28-30), and Yuan and Yin (1999, p. 173-175). To these lists, several species are added from Australia, northern Germany, and southern France: Proceratopyge ocella Webby, Wang, and Mills, 1988 (p. 926, pl. 85, figs. 1-10), Proceratopyge sp. Webby, Wang, and Mills, 1988 (p. 926, pl. 85, fig. 11); Proceratopyge sellinensis Buchholz, 2000 (p. 751-753, pl. 7, figs. 3-7; text-fig. 11) and Proceratopyge (Proceratopyge) sp. indet. (Shergold et al., 2000, p. 619, pl. 2, figs. 16-20). Proceratopyge elonga Bao and Jago, 2000 (p. 896, pl. 2, figs. 8-13), described from the uppermost upper Cambrian of southwestern Tasmania, does not belong to Proceratopyge but to Diceratopyge Troedsson, 1937. Remarks. Shergold et al. (2000, p. 619) discussed the status of the genus and summarized differing views on its subgeneric division. Those authors followed Lu and Lin (1989) and Peng (1992) in recognizing three subgenera for the genus: Proceratopyge (Proceratopyge), P. (Sinoproceratopyge), and P. (Lopnorites). Peng's (1992) subgeneric concept of P. (Proceratopyge) is followed here.

• 162.

Subgenus PROCERATOPYGE(PROCERATOPYGE) Wallerius, 1895 Proceratopyge (Proceratopyge) fenghwangensis Hsiang in Egorova et al., 1963

Plate 53, figures 5, 9" Plate 54, figures 1-4 Proceratopyge fenghwangensis Hsiang in Egorova et al., p. 47, 48, pl. 10, figs. 1-10. Proceratopyge fenghwangensis Hsiang; Lu, Zhang, Zhu, Qian, and Xiang, p. 548, 549, pl. 114, figs. 10-13. 1977 Proceratopyge fenghwangensis Hsiang; Zhou, Liu, Meng, and Sun, p. 232, 233, pl. 70, figs. 10-18. 1978 Proceratopyge fenghwangensis Hsiang; Yin and Li, p. 179, pl. 70, figs. 6, 7. 1978 Proceratopyge fenghwangensis Hsiang; Yang, p. 65, 66. text-figs. 7a-j, 8a-d. 1982 Proceratopyge fenghwangensis Hsiang (in part); Liu, p. 333, pl. 228, fig. 4, pl. 229, fig. 1; non pl. 228, fig. 14 [?=P. (P.) truncata Yang, 1978]. ?1982 Proceratopygefenghwangensis ellipsoides; Liu, p. 333, pl. 228, fig. 6. 1989 Proceratopyge fenghwangensis Hsiang; Lu and Lin, p. 146, pl. 23, fig. 8. 1999 Proceratopyge (Lopnorites) orthogonialis Yang in Duan et al. (in part), p. 149, fig. 5Aa: ?fig. 4A, non fig. 4E. 2001 c Proceratopyge fenghwangensis Hsiang; Peng. Babcock, Lin, Chen, and Zhu, p. 166, pl. 4, fig. 17; pl. 5, figs. 12, 13.

1963 1965

Lectotype. Pygidium (Egorova et al., 1963, pl. 10, fig. 6; CMG 2651), from the middle Huaqiao Formation (lower and middle upper Cambrian as recorded by Xiang in Egorova et al., 1963) from Huanghexiang, Fenghuang, western Hunan; designated herein. New material. More than 40 sclerites, including cranidia, librigenae, and pygidia (illustrated specimens NIGP 133563, 137797, 137798, 137083, 137084) in collections P317.2, P367, P367.76, P368, P368.8, P372.12, P371.2, P373.62, P374.9, P375, P376.4, P378.25, PI3-1.72, PI3-1.8, PI3-1.65, PI37. Remarks. No single name-bearing specimen was assigned previously for this species. Xiang (in Egorova et al., 1963, p. 47, 108) assigned two specimens as the "holotype" for this species (a cranidium, Egorova et al., 1963, pl. 10, fig. 1; and a pygidium, Egorova et al., 1963, pl. 10, fig. 6). The pygidium is here chosen as the lectotype of the species because it has more distinct characters than the cranidium. By the assignment of the lectotype, this species is characterized by having a rather wide pygidium; furrowed pleural areas; and posterior margin with shallow a bow. The ontogenetic material of Yang (1978) shows that the eye ridge is relatively long and that the lateral glabellar furrows are fairly distinct in the fully developed cranidia. Occurrence. The lectotype is from the middle Huaqiao Formation, Huanghexiang, Fenghuang, western Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone and the Chuangia subquadrangulata Zone (equivalent to the Linguagnostus reconditus Zone through the Glyptagnostus reticulatus Zone).

• 163-

Proceratopyge (Proceratopyge)fuyangensis Lu and Lin in Peng, 1987 Plate 52, figures 1-14; Plate 53, figures 1-4 1977 1987

Proceratopyge conifrons Wallerius; Yang, pl. 7, p. 80, figs. 22, 23. Proceratopyge (Proceratopyge)fuyangensis Lu and Lin in Peng, p. 114, 115, pl. 8, figs.

1989 1999

Proceratopyge (Proceratopyge)fuyangensisLu and Lin; Lu and Lin, p. 148, pl. 24, figs. 4, 5. Proceratopyge (Proceratopyge) conifrons sinensis Yuan and Yin, p. 175, 176, pl. 1, figs. 18,

?2000

Proceratopyge (Proceratopyge) sp. undet., Shergold, Feist, and Vizcaino, p. 619, 620, pl. 2,

7-10.

19. figs. 16-20. 2001b Proceratopyge conifrons Wallerius, Peng, Babcock, and Lin, p. 104, pl. 12, fig. 2.

Holotype. Cranidium (Peng, 1987, pl. 13, fig. 10; NIGP 66475), from the Lejopyge sinensis Zone, Huayansi Formation, Yaogongbu, Fuyang, western Zhejiang.

New material. More than 30 sclerites, including cranidia and pygidia (illustrated specimens NIGP 137779-137792; 137794-137796) in collections P286.3, P298.4, P300.4, P317.4, P319.6, P319.6, P3226.9, P327.4, P371.5, W210.5, W211.7, W216.5, W218.3, W225, W227, W228.3, and W229.9.

Remarks. This species is characterized by having a broadly conical, rather effaced glabella, and a transverse pygidium with a short and somewhat less segmented axis; the first pleural segment is large, and the pleural spines are slender. A plectrum is not obvious on the type material, but the new material includes a large collection of cranidia from the middle part of the Huaqiao Formation (upper Proagnostus bulbus Zone) showing that the anterior border in this species does bear a plectrum that is variably defined and variable in length (sag.). The $2 and $3 furrows are faint in specimens from lower stratigraphic occurrences, clarified in later forms. This species is most similar to P. (P.) conifrons from the Lejopyge laevigata Zone of Sweden, but differs in having a relatively larger and more posteriorly located palpebral lobe; a transverse, rather than oblique, posterior branch of the facial suture that encloses a blade-like, rather than triangular, posterolateral projection; and a relatively broad glabella. This species has a total observed stratigraphic range though the Proagnostus bulbus Zone into the lower Linguagnostus reconditus Zone, and is younger in age than P. (P.) conifrons. The wide pygidium of this species helps to differentiate it from all other species of P. (Proceratopyge).

Occurrence. The holotype is from the Lejopyge sinensis Zone, Huayansi Formation, Yaogongbu, Fuyang, western Zhejiang, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone through the lower part of the Linguagnostus reconditus Zone).

• 164.

Proceratopyge (Proceratopyge) truncata Yang in Zhou et al., 1977

Plate 53, figures 6-8: Plate 54, figures 5-10; Text-figure 27 1977 1978 ?1987 ?1991 2001 b

Proceratopyge truncatum Yang in Zhou et al., p. 233,234, pl. 70, figs. 18, 19. Proceratopyge truncatum Yang; Yang, p. 67, pl. 7, figs. 19, 20. Proceratopyge gordonensis Jago; Yang, p. 69, 70, pl. 3, figs. 5-17. Proceratopyge sp. cf P. gordonensis Jago; Jell, Hughes, and Brown, p. 467, fig. 6. Proceratopyge fenghwangensis Hsiang in Egorova et al.; Peng, Babcock, and Lin, p. 106, pl.

16, fig. 14. Lectotype. Cranidium (Zhou et al., 1977, pl. 70, fig. 18; also Yang, 1978, pl. 7, fig. 19a, CUGB 0104802-a; refigured Yang, 1978; Text-fig. 27A, left herein) from the Chuangia-Prochuangia Zone (Glyptagnostus reticulatus Zone), upper Huaqiao Formation, Huaqiao, Baojing, western Hunan;

designated subsequently as the holotype of the species by Yang (1978). New material. More than 50 sclerites, including cranidia and pygidia in (illustrated specimens

NIGP 137799-137801; 137805-137809) collections P374.9, P378.25, P[327.2, P[335.4, P[356.5, P]360.3, P[360.15, P[364.3, P[365.8, and P[3 72.

A

B

Text-figure 27. A, B, Lectotype cranidium and associated paratype pygidium of Proceratopyge (Proceratopyge) truncata Yang in Zhou et al., 1977, CUGB 0104802-a, 0104802-b, x 2.7, original of Zhou et al., 1977 (pl. 70, fig. 18; also Yang, 1978, pl. 7, fig. 19)" B is the paratype pygidium in dorsal view. Remarks. The lectotype cranidium and the associated paratype pygidium are refigured as Text-fig.

27A. The lectotype cranidium is mostly exfoliated, and both sides of the anterior area of the fixigena, as well as the left posterolateral projection, are incompletely exposed. Additionally, the posterior end of the palpebral lobe is broken. Key characters of the species include an elongate, anteriorly truncate glabella with deep S 1, and weak $2 and $3 furrows; a palpebral lobe that lies close to the axial furrow; a short, strongly oblique eye ridge; a relatively long (sag.) but narrow (tr.) anterior area of the fixigena; a gently diverging anterior branch of the facial suture with the posterior end being close to the axial furrow; a pygidium having a length greater than the width of the anterior margin of the pleural region; and a posterior margin that is rather deeply bowed. The • 165-

new material from the Glyptagnostus reticulatus Zone of the Huaqiao Formation is from the same stratigraphic horizon in the same syncline of western Hunan as is the holotype. The new material agrees in all characters with P. (Proceratopyge) truncata Yang. Proceratopyge gordonensis Jago from the Singing Creek Formation, southwestern Tasmania, is similar to P. (Proceratopyge) truncata. The new material from Hunan shows that the Tasmanian species resembles P. (P.) truncata in all its essential characters. The Tasmanian species is also similar in age, and we tentatively regard it as a junior synonym of P. (P.) truncata.

Occurrence. The lectotype is from the Chuangia-Prochuangia Zone, Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it is associated with trilobites indicative of the Chuangia subquadrangulata Zone through the base of the Shengia quadrata Zone (equivalent to the Glyptagnostus reticulatus Zone).

Proceratopyge sp. indet. Plate 52, figure 15

Material. One incomplete cranidium, NIGP 137793, in collection P319.6. Remarks. A broken cranidium is similar to P. (Proceratopyge)fuyangensis Lu and Lin in Peng, 1987, and it occurs within the observed stratigraphic range of P. (P.)fuyangensis. It differs in having better defined $2 and $3 furrows, a longer preglabellar field, a more clearly defined anterior border, a less well defined plectrum, and more divergent anterior branches of the facial suture. Incomplete preservation of the specimen, however, precludes a firm taxonomic assignment for this cranidium.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Subfamily IWAYASPIDINAE Kobayashi, 1962 Genus PSEUDOYUEPINGIA Chien, 1961

Pseudoyuepingia Chien (Qian), 1961, p. 106, 126; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 506; Zhou, Liu, Meng, and Sun, 1977, p. 215, 216; Yin and Li, 1978, p. 534; Lu and Lin, 1980, p. 127, 128; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 207; Jago, 1987, p. 226, 227; Webby, Wang, and Mills, 1988, p. 920-922; Lu and Lin, 1989, p. 155; Pratt, 1992, p. 46, 47; Duan, Yang, and Shi, 1999, p. 145. lwayaspis Kobayashi, 1962, p. 122; Lazarenko, 1968, p. 184; Palmer, 1968, p. 53. Aplotaspis Henderson, 1976, p.339, 340; Shergold, 1982, p. 52; Pegel, 2000, p. 1017. Aplotaspis (Cataplotaspis) Yuan and Yin, 1999, p. 176.

Type species. Pseudoyuepingia modesta Chien, 1961 (Qian, p. 106, pl. 3, figs. 5-7), from the upper Cambrian Sandu Formation, Sandu, southeastern Guizhou; by original designation.

• 166.

Remarks. The genetic concept of Qian (1961) is followed here. Pseudoyuepingia is a non-spinose ceratopygoid genus, resembling Proceratopyge in overall respects but lacking paired anterolateral pleural spines in the pygidium. We follow Lu and Lin (1980) in synonimizing Iwayaspis Kobayashi, 1962 as a junior synonym of Pseudoyuepingia. Pratt (1992) further regarded lwayaspis asaphoides, the type species of Iwayaspis, and P. venusta Jago (1987) as synonyms of the type species of Pseudoyuepingia. Aplotaspis Henderson, 1976 and its subgenus Aplotaspis (Cataplotaspis) Yuan and Yin, 1999 are also suppressed, as the both type species of Aplotaspis and A. (Cataplotaspis) are quite similar to Pseudoyuepingia modesta. Yuan and Yin (1999) listed a number of characters by which Aplotaspis (Cataplotaspis) could be distinguished from Pseudoyuepingia and transferred Pseudoyuepingia laochatianensis to that subgenus. However, the differences are all minor, and none of them is regarded here as of more than species-level significance. In addition, the type species of A. (Cataplotaspis) is obviously a junior synonym of Pseudoyuepingia laochatianensis (see discussion under the latter species). Pseudoyuepingia laochatianensis Yang in Zhou et al., 1977 Plate 55, figures 1-18; Text-figure 28

Proceratopyge conifrons Wallerius; Egorova, Xiang, Li, Nan, and Guo, p. 46, 47, pl. 10, figs. 11,12. ?1965 Proceratopyge conifrons Wallerius; Lu, Zhang, Zhu, Qian, and Xiang, p. 547, pl. 114, figs. 7, 8. ?1968 lwayaspis cf. L asaphoides Kobayashi; Palmer, 1968, p. 53, pl. 10, figs. 9, 10. 1977 Pseudoyuepingia laochatianensis Yang in Zhou et al., p. 216, pl. 63, figs. 14, 15. ?1977 Proceratopyge conifrons Wallerius; Zhou, Liu, Meng, and Sun, p. 232, pl. 70, figs. 5, 6. 1978 Pseudoyuepingia laochatianensis Yang; Yang, p. 69, 70, pl. 3, figs. 5-17. 1982 Pseudoyuepingia laochatianensis Yang; Liu, p. 329, pl. 225, figs. 8-10. 1983 Pseudoyuepingia laochatianensis koutianwuensis Qiu in Qiu et al., 1983, p. 208, pl. 69, fig. 5. 1983 Pseudoyuepingia aspinosa Qian in Qiu et al., 1983, p. 207, 208, pl. 69, fig. 4. 1989 Proceratopyge (Proceratopyge) conifrons Wallerius; Lu and Lin (in part), p. 147, pl. 23, figs. 12, 13 only. ?1989 Proceratopyge (Proceratopyge) zhejiangensis Lu and Lin, p. 147, pl. 24, figs. 1-3. 1999 Aplotaspis (Cataplotaspis) quadrata Yuan and Yin, p. 177-179, pl. 1, figs. 7-17. 1999 Aplotaspis (Cataplotaspis) spiculata Yuan and Yin, p. 179, pl. 2, figs. 15-17. 1999 Aplotaspis (Cataplotaspis) laochatianensis (Yang); Yuan and Yin, p. 180, pl. 2, figs. 1-12. 1999 Aplotaspis (Cataplotaspis) laochatianensis nobilis Yuan and Yin, p. 180, 181, pl. 1, figs. 1-6; pl. 2, figs. 13, 14. 2001b Pseudoyuepingia laochatianensis Yang; Peng, Babcock, and Lin, p. 105, pl. 14, figs. 6, 7, 16. ?1963

Lectotype. Incomplete, immature exoskeleton (Yang in Zhou et al., 1977, pl. 63, fig. 14; also Yang, 1978, pl. 3, fig. 9, CUGB 0321314; Text-fig. 28, herein) from the middle part of Huaqiao Formation, western Hunan; designated subsequently as the holotype of the species by Yang (1978). New material. More than 50 sclerites, including an ontogenetic series of cranidia and pygidia (illustrated specimens NIGP 137810-137827) in collections P344.6, P353.64, P353.7, P361.5, P362.35, W 196.3, W 199.2, and W254.1.

• 167.

Remarks. This species was originally described (Yang, 1978) on the basis of several cranidia, pygidia, and exoskeletons; two specimens were published earlier than the others (Yang in Zhou et al., 1977). Previous to this report, all figured material of Pseudoyuepingia laochatianensis was from the same section at Tingziguan, near Fenghuang, western Hunan. Yang's (in Zhou et al., 1977)

figured specimens include an early holaspid exoskeleton and a well developed but incomplete exoskeleton lacking the cephalon. The immature exoskeleton was subsequently selected as the 'holotype' for the species (Yang, 1978, pl. 3, fig. 9; Text-fig. 28, herein), and it shows some morphological differences from larger holaspids. This specimen must be regarded as the lectotype of the species. The cephalon of the lectotype has a subcylindrical glabella with faint lateral furrows, a large median node between the S 1 furrows, and a rounded front. The lectotype has a pygidium with faint border furrows, a gently concave posterior border, and obscurely furrowed axial and pleural regions. Large exoskeletons and cranidia in Yang's (1978) original suite show that the glabella is subconical, has an acutely rounded front, has clearly defined S1 furrows, and has a rather well furrowed pygidium. The present material from the Huaqiao Formation of western Hunan is assigned to P. laochatianensis because both immature and mature specimens conform in all characters to Yang's (1978) material of the species.

Text-figure 28. The lectotype of Pseudoyuepingia laochatianensis Yang in Zhou et al., 1977, an early holaspid exoskeleton lacking librigenae, CUGB 0321314, x 11, refigured and designated subsequently as holotype of the species by Yang (1978), original of Zhou et al., 1977 (pl. 63, fig. 14; also Yang, 1978, pl. 3, fig. 9). The new material from Hunan reveals some additional ontogenetic variation in the species. It shows that during ontogeny the furrows on both the cranidium and pygidium change progressively from obscure to more distinct, the cranidial outline becomes progressively elongated, and the pygidial outline becomes proportionately shorter and transverse. The smallest cranidium (P1. 55, fig. 1) in the new material, which probably belongs to an early meraspid, is characterized by having a transverse-subtriangular outline, a poorly defined glabella, low convexity, and no lateral furrows. The smallest pygidium (P1. 55, fig. 9) is semicircular or inverted-trapezoidal in outline, and bears • 168.

almost no furrows in the pleural region. A cranidium (P1. 55, fig. 8) in the new material shows a glabella with four pairs of rather distinct lateral furrows. P. laochatianensis differs from P. modestus Chien (Qian, 1961), the type species of Pseudoyuepingia, in having a plectrum-bearing cranidium, a more tapered glabella, a large, more posteriorly located palpebral lobe, a transverse rather than oblique posterior branch of the facial suture, and a pygidium with a shorter, postaxial ridge-beating axis. Aplotaspis (Cataplotaspis) quadrata Yuan and Yin (1999), the type species of A. (Cataplotaspis), is considered to be a junior synonym of P. laochatianensis because it appears to be indistinguishable from P. laochatianensis. The holotype cranidium of A. (C.) quadrata was said (Yuan and Yin, 1999) to be characterized by having a glabella with a subquadrate outline, but this is probably the result of longitudinal compression. Other material assigned to A. (C.) quadrata by Yuan and Yin (1999) is essentially indistinguishable from P. laochatianensis, except for somewhat weaker furrowing of the pleural fields of the pygidium. This feature is considered to be within the range of intraspecific variation.

Occurrence. The lectotype is from the middle part of the Huaqiao Formation, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus reconditus Zone through the basal Glyptagnostus reticulatus Zone). Superfamily

TRINUCLEOIDEA

Family

Hawle and Corda, 1847

ALSATASPIDIDAE Turner,

1940

Subfamily HAPALOPLEURINAEHarrington and Leanza, 1957 (nora. trans, ex. Hapalopleuridae Harrington and Leanza, 1957)

Remarks. The taxonomic rank of the hapalopleurine trilobites is somewhat in flux, with authors regarding the group as either of the family Hapalopleuridae (Harrington and Leanza, 1957; Ludvigsen et al., 1989; Peng, 1992) or of subfamily rank within the family Orometopidae (i.e., Hapalopleurinae; Fortey and Shergold, 1984; Fortey and Chatterton, 1988; Babcock and Smith, 2003). Peng (1992) regarded the Egorovaiidae as a junior synonym of the Hapalopleuridae, and Fortey (1997) considered the Hapalopleuridae, Myindidae, and Egorovaiidae all to be junior synonyms of the Orometopidae. Jell and Adrain (2003) regarded the Orometopidae and the Hapalopleuridae to be junior synonyms of the Alsataspididae. Here, the Gaoloupingiinae Yuan and Yin, 1998 is regarded as a junior synonym of the Hapalopleuridae. Pending further comprehensive study of the hapalopleurine and related trilobites, Cambrian and Ordovician genera formerly assigned to the Hapalopleuridae are referred to the subfamily Hapalopleurinae (family Alsataspididae). The definition of the Hapalopleurinae followed here is that of Harrington in Moore (1959), as emended by Babcock and Smith (2003). The subfamily includes, at a minimum, the Cambrian genera Ajrikina Kraskov in Borovikov and Kraskov, 1963, Torifera Wolfart, 1974, and Gaoloupingia Yuan and Yin, 1998, and the Early Ordovician genera Hapalopleura Harrington and Leanza, 1957, Araiopleura Harrington and Leanza, 1957, Rhadinopleura Harrington and Leanza, 1957, Palquiella Sufirez-Soruco, 1975, and Guanacunopleura Babcock and Smith, 2003.

• 169.

Genus GAOLOUPINGIAYuan and Yin, 1998 1998 GaoloupingiaYuan and Yin, p. 161, 162.

Type species. Gaoloupingia gaoloupingensis Yuan and Yin, 1998 (p. 162, 163, pl. 5, figs. 4-10) from the Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone to the Glyptagnostus stolidotus Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou; by original designation. Other species. Gaoloupingia triangularis Yuan and Yin, 1998 (p. 163, pl. 5, fig. 11), Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone to the Glyptagnostus stolidotus Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou.

Emended diagnosis. Alsataspididae with subparallel-sided, anteriorly rounded glabella bearing four pairs of deeply notched furrows; frontal area with depressed frontal field and thick, weakly defined anterior border; eye ridges transverse, strongly raised; palpebral lobes located subcentrally, lying opposite $3 furrow; surface granulose. Remarks. Yuan and Yin (1998) erected this genus with Gaoloupingia gaoloupingensis Yuan and Yin, 1998 as the type species. The genus is characterized in part by having a wide and upturned cranidial border; distinct, transverse eye ridges; and small palpebral lobes that are located anteriorly. On the basis of this genus, Yuan and Yin (1998) erected a new subfamily, Gaoloupingiinae, and referred it to the Shumardiidae. Yuan and Yin's (1998) classification is rejected here because the presence of strong eye ridge, small eyes, depression of the frontal field, and the presence of an anterior border are inconsistent with the morphology of the Shumardiidae. Commonly the shumardiids are blind, lack eye ridges and an anterior border, and have a convex rather than concave frontal area. Only two known shumardiid genera have eyes (Oculishumardia Peng et al., 2003a, Akoldinioidia Zhou and Zhang, 1984 and Limbishumardia gen. nov. in Volume 2 of this two-part work), and only two species (Shumardia goniolata Lu and S. lata Lu) have eye ridges (Lu, 1975). The convex, anteriorly rounded glabella with short, inwardly notched glabellar furrows, the strong eye ridges, and the small palpebral lobes lying far from the glabella all support an assignment of Gaoloupingia to the Hapalopleurinae.

Gaoloupingia gaoloupingensis Yuan and Yin, 1998 Plate 2, figures 10-12 1998 2001b

Gaoloupingia gaoloupingensis Yuan and Yin, p. 162, 163, pl. 5, figs. 4-10. Gaoloupingia gaoloupingensis Yuan and Yin; Peng, Babcock, and Lin, p. 105, pl. 13, fig. 7.

Holotype. Incomplete cranidium (Yuan and Yin, 1998, pl. 5, fig. 10; NIGP 127959) from the Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone to Glyptagnostus stolidotus Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou. New material. Two cranidia, NIGP 137298, 137299, in collections P331.8 and W216.5. • 170.

Remarks. New material from the Huaqiao Formation of western Hunan is identical in all respects to the type material from the same biogeographic region in eastern Guizhou. Besides the type species of the genus, Yuan and Yin (1998) described a second species, G. triangularis, based on a single cranidium collected from the same locality as G. gaoloupingensis, but in a bed that is slightly lower than that containing the type species. Yuan and Yin (1998) differentiated G. triangularis from the type species by a narrow, triangular cranidium with a strongly arched anterior margin, a flatter frontal area, narrower fixigenae, shorter eye ridges, and more strongly convergent anterior branches of the facial suture. Examination of Yuan and Yin's (1998) type material of both G. gaoloupingensis and G. triangularis by one of us (SP) shows that among the differential features Yuan and Yin listed, the "flatter frontal area" is hard to recognize as a distinct feature on the holotype cranidium of G. triangularis. Other features, such as the shorter eye ridges, the narrower fixigenae, and the more convergent anterior branches of the facial suture, are taphonomic in origin because the left side of the cranidium has been damaged; the fight side of the cranidium has not been completely exposed. The shape of the anterior cranidial margin in the holotype of G. triangularis seems to be the only feature that distinguishes G. triangularis from the type species. However, a cranidium in the new material (P1. 2, fig. 10) from Hunan shows curvature of the anterior margin intermediate between that of G. triangularis and G. gaoloupingensis, suggesting that this distinctive feature falls within the range of variation of G. gaoloupingensis. In short, the validity of G. triangularis remains problematic, and further investigation may prove that it is synonymous with the type species. However, so little is known about Gaoloupingia that, until more material is available for both G. gaoloupingensis and G. triangularis, these two species are provisionally considered to be valid. Occurrence. The holotype is from the Hadragnostus modestus-Blackwelderia Zone to the Glyptagnostus stolidotus Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone and the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Genus AJRIKINA Kraskov in Borovikov and Kraskov, 1963

Ajrikina Kraskov in Borovikov and Kraskov, 1963, p. 279; Romanenko in Goncharova et al., 1972, p. 214; Ergaliev, 1980, p. 140. Huamiaocephalus Qiu in Qiu et al., 1983, p. 202, 203. Jiangnania Lin and Zhou in Lin et al., 1983, p. 407; Peng, 1987, p. 111; Lu and Lin, 1989, p. 141, 255; Duan, Yang, and Shi, 1999, p. 150. Type species. Ajrikina bulakeensis Kraskov in Borovikov and Kraskov, 1963 (p. 279, 280, pl. l, figs. 21-23), from the lower part of the Bulak-Airyk Formation, Kendyktas Range, southern Kazakhstan; by original designation. Other species. Ajrikina lusoria Romanenko, 1977 (p. 173, pl. 24, figs. 4, 8) from the lower upper Cambrian [equivalent to the latest Wulingian], Bolishaya Isha River Basin, Gorlyi Altai; Ajrikina elongata Ergaliev, 1980 (p. 140, 141, pl. 5, fig. 7) from the Lejopyge laevigata Zone and the Kormagnostus simplex Zone, Malyi Karatau, Kazakhstan; Huamiaocephalus wannanensis Qiu in Qiu et al., 1983 (p. 203, pl. 67, fig. 8) from the middle part of the upper Cambrian [late Wulingian], • 171.

Huamiao, Guichi, southern Anhui; Jiangnania miranda Lin and Zhou in Lin et al., 1983 (p. 407, pl. 3, fig. 5), from the Tuanshan Formation, Kunshan, southern Jiangsu; Jiangnania hunanensis Peng (1987, p. 12, figs. 2, 3; text-fig. 15) from the Agnostascus (Paragnostascus) sinensis [=Proagnostus bulbus] Zone, Wa'ergang, Taoyuan, northwestern Hunan; Jiangnania granulosa Peng, from the Agnostascus (Paragnostascus) sinensis [=Proagnostus bulbus] Zone, Wa'ergang, Taoyuan, northwestern Hunan; Ajrikina sp. (Romanenko, 1977, not described, pl. 24, figs. 6, 7) from the upper middle Cambrian (middle Wulingian-equivalent, Bolishaya Isha River Basin, Gorlyi Altai; and specimens referred to Ajrikina mera by Romanenko (1977, pl. 24, figs. 3, 5), from the lower upper Cambrian (upper Wulingian-equivalent), Bolishaya Isha River Basin, Gorlyi Altai. Ajrikina mera Romanenko in Goncharova et al. (1972, p. 214, 215, pl. 53, figs. 3-5, text-fig. 53) from the lower upper Cambrian [latest Wulingian-equivalent], Gorlyi Altai, western Siberia, should be excluded from the genus. The species apparently belongs to Torifera.

Remarks. Huamiaocephalus is here considered to be a junior synonym of Ajrikina. Huamiaocephalus includes only one species, Huamiaocephalus wannanensis Qiu, which was based on a single cranidium. The holotype cranidium of H. wannanensis is subtriangular in outline, has a small, subconical glabella, has a raised anterior portion of the frontal area, has a long, nearly transverse eye, and has a transverse, convex, belt-like posterior part of the fixigena. Except for a semicircular cranidial outline, all these features are shared by Ajrikina bulakeensis, the type species of Ajrikina. Such a minor distinction should be of no more than species-level significance. Jiangnania was published four months after the publication of Huamiaocephalus. The type species, J. miranda, is based on a single specimen. The holotype is a fragmentary cranidium having damage to the glabella, the fight palpebral lobe, and the fight posterior areas of the fixigena. All the observed features of the cranidium show close similarity to the holotypes of both A. bulakeensis (Borovikov and Kraskov, 1963, pl. 1, figs. 22, 22a) and H. wannanensis (Qiu et al., 1983, pl. 67, fig. 8). The holotype of J. miranda differs only in having a forwardly oblique eye ridge, and thus a more anteriorly located palpebral lobe. Such a difference is insufficient to serve as a genus-level character, and Jiangnania should be placed in synonymy with Huamiaocephalus. Huamiaocephalus, in turn, is considered to be a junior synonym of Ajrikina. In following Lin et al. (1983) and Peng (1987), Lu and Lin (1989) regarded Jiangnania as a valid genus, but failed to compare the genus with Huamiaocephalus. Lu and Lin (1989, p. 141, 255) considered Jiangnania to be distinguishable from Ajrikina. However, the differences they listed are either not true (absence of bacculae and transverse depressions and elevations on the fixigenae of Ajrikina) or they are not of genetic significance (more oblique eye ridges and more anteriorly located palpebral lobes in Jiangnania). Based on large collections, Peng et al. (2001b) suppressed Jiangnania as a junior synonym of Ajrikina, and transferred J. hunanensis Peng, 1987 to Ajrikina. Ajrikina wannanensis (Qiu in Qiu et al., 1983) Plate 56, figures 1, 2 1983 Huamiaocephaluswannanensis Qiu in Qiu et al., p. 203, pl. 67, fig. 8.

Holotype. By monotypy; incomplete cranidium (Qiu in Qiu et al., 1983, pl. 67, fig. 8, HIT 0425), from the middle part of the upper Cambrian (upper Wulingian-equivalent), Huamiao, Guichi, southern Anhui. New material. Five cranidial fragments (illustrated specimens NIGP 137830, 137831) in collections • 172.

W187.6, W196.3, W197.5, W199.2, and W200.3. Remarks. The new material is poorly preserved, but all observed features seem to be identical with those of Huamiaocephalus wannanensis Qiu (in Qiu et al., 1983), a species that is based on a poorly preserved cranidium that is now regarded as belonging to Ajrikina. The holotype cranidum is characterized by having a subtriangular cranidial outline and a transverse eye ridge. A. wannanensis is most similar to Ajrikina lusoria Romanenko, 1977, but differs in having a wide (sag.) preglabellar field, a wider (exs.) and more convex transverse swelling on the posterior part of the fixigena, and a proportionally longer (sag.) cranidium. Occurrence. The holotype is from the middle part of the upper Cambrian, Huamiao, Guichi, southern Anhui, China. New material is from dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone (equivalent to the Linguagnostus reconditus Zone). Ajrikina hunanensis (Peng, 1987) Plate 56, figures 3-17; Plate 57, figures 1-5 1987 1989 1989 2001b

Jiangnaniahunanensis Peng, p. 111, 112, pl. 12, figs. 2, 3; text-fig. 15. Jiangnaniafuyangensis Lu and Lin, p. 141,142, 255; pl. 22, fig. 9. Jiangnaniafengzuensis Lu and Lin, p. 142, 256; pl. 22, fig. 10. Ajrikina hunanensis (Peng); Peng, Babcock, and Lin, p. 103, pl. 7, figs. 14, 15; p. 104, pl. 11, figs. 15-17. 200 ld Ajrikina hunanensis (Peng); Peng, Babcock, Lin and Chen, p. 141, fig. 9.8.

Holotype. Cranidium (Peng, 1987, pl. 12, fig. 2; NIGP 74572), from the middle part of the upper Agnostascus (Paragnostascus) sinensis [=Proagnostus bulbus] Zone (upper Wulingian), Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan. New material. More than 20 cranidia and four pygidia (illustrated specimens NIGP 137830-137840) in collections P277, P282.3, P353.7, W210.5, W216.5, W218.3, W219.7, and W228.3. Description. Cranidium subsemicircular; length half width. Glabella small, tapering gently forward, obtusely rounded anteriorly, occupying one-half to three-fifths of cranidial length; glabella with maximum convexity in front of occipital furrow, with poorly defined node at highest point. Glabella with three pairs of short lateral glabellar furrows; S 1 deeply incised, directed diagonally, almost isolating L1 from glabella; $2 pit-like, impressed inwardly; $3 short, faintly defined. Occipital furrow transverse, deeply incised, deepest at sides; occipital ring narrow, bearing short stout spine posteriorly. Baccula small, weakly defined, same length as L1. Eye ridge thin, long, clearly defined, outwardly and forwardly directed; palpebral lobes small, located anteriorly, widely spaced. Anterior border strongly convex, widest sagittally, narrowing abaxially, gently swollen medially; anterior field of fixigena flat, depressed; posterior area of fixigena large, with wide (exs.) transverse depression widening abaxially. Lateral border narrow, strongly convex, gently curved anteriorly, flexed inward and rearward posteriorly. Posterior border furrow transverse, deep, widening slightly abaxially; posterior border narrow, linear. • 173-

Pygidium transverse, triangular, width three to four times length. Axis narrow, nearly reaching posterior border furrow, with six tings and transverse bar-like terminal piece. Pleural field flat, with posterior portion gently convex; pleura transverse, with linear anterior and posterior bands, moderately deep, broad pleural furrow; interpleural furrow deeply incised. Lateral and posterior borders linear, defined by moderately deep border furrows. Posterior part of pleural field bears terrace lines. Remarks. The new material from northwestern Hunan is identical in all observed features with the type material of Jiangnania hunanensis from Wa'ergang, Taoyuan, and is referred to the species. Jiangnania is regarded as a junior synonym of Ajrikina. A large collection from northwestern Hunan adds cranidial features and the pygidial morphology to the species concept. The new material shows that the anterior portion of the anterior border is folded beneath the posterior portion, and that the occipital ring bears a spine on the posterior margin sagittally. The new pygidium, which is the first recorded for the species and the genus, shows a distinctive outline and pleural structure. Ajrikina hunanensis is comparable to A. miranda (Lin and Zhou; in Lin et al., 1983, pl. 3, fig. 5), but A. miranda has a wider (sag.) and less well defined anterior border. A. miranda is based on a single fragmental cranidium that has a broken glabella. Poor preservation prevents further comparison. Ajrikina elongata Ergaliev, 1980 is also similar to A. hunanensis. The Kazakhstan species, however, differs in having a more narrow (tr.) and convex anterior cranidial border with an acutely curved anterior margin, less well defined bacculae, and probably in lacking an occipital spine. Occurrence. The holotype is from the Proagnostus bulbus Zone, Huaqiao Formation, We'ergang, Taoyuan, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone and the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Genus

TORIFERA

Wolfart, 1974

Torifera Wolfart, 1974, p. 96, 97; Wittke, 1984, p. 17, 18; Fortey, 1994, p. 38; Jell and Hughes, 1997, p. 101. Xiangia Peng, 1987, p. 112, 113. Type species. Torifera triangularis Wolfart (1974, p. 97-101, pl. 12, figs. 1-9; pl. 13, figs. 1-6; text-fig. 11) from the upper Cambrian, about 1 km west of Surkh Bum, central Afghanistan; by original designation. Other species. Ajrikina mera Romenenko in Goncharova et al. (1972, p. 214, 215, pl. 53, figs. 3-5; text-fig. 53) from the lower upper Cambrian, Gorlyi Altai, western Siberia; Cyclolorenzella tuma Yang in Zhou et al. (1977, p. 170, pl. 50, figs. 17, 18) from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Laochatian, Fenghuang, northwestern Hunan, China; Torifera youyangensis Zhu in Zhang et al. (1980, p. 383, pl. 132, figs. 10, 11) from the Dashuijing Formation (basal upper Cambrian), Youyang, southeastern Sichuan; Xiangia taoyuanensis Peng (1987, p. 113, pl. 13, figs. 1-6; text-fig. 16) from the Huaqiao Formation, Taoyuan, northwestern Hunan; Torifera cf. T. triangularis Wolfart (Fortey, • 174.

1994, p. 38, Fig. 6B-F) from Fauna A, Andam Formation, Qarn Mahata Humaid, Oman; and Torifera abrupta sp. nov. Species questionably transferred here to Torifera are: Cvclolorenzella paraconvexa Yang (in Yin and Li, 1978, p. 488, 489, pl. 165, fig. 17; Yang, 1978, p. 42, 43, pl. 7, fig. 7) from the Parablackwelderia jimaensis-Torifera tuma Zone, Huaqiao Formation, Liangweizhou, Yuping, eastern Guizhou; Cyclolorenzella? sp. (Whittington, 1986, pl. 18, figs. 2, 3) from the upper middle Cambrian, Zanskar, Ladakh, India; Agraulos tonkinensis Mansuy ( 1915, pl. 2, figs. 13a-c) from the middle Cambrian, Vietnam; and Cyclolorenzella ventaiensis (Chu) sensu Zhou et al. (1977, p. 170, pl. 50, fig. 19).

Remarks. As remarked by Fortey (1994), the generic concept of Wolfart (1974) is clear; that concept is followed here. Xiangia is considered to be synonymous with Torifera. Peng (1987) differentiated Xiangia (type species X. taoyuanensis) from Torifera by the absence of an anterior border, shorter palpebral lobes, and the presence of a preglabellar boss defined by paired exsagittal furrows. New material from western Hunan reveals that X. taovuanensis also bears an anterior border. The presence or absence of a preglabellar boss is likely related to sexual dimorphism (see Fortey and Hughes, 1998). T. taoyuanensis has relatively small palpebral lobes, but these are considered to be of species-level significance. Torifera taoyuanensis (Peng, 1987) Plate 58. figures l - l l

Xiangia taovuanensis Peng (in part), p. 113. pl. 13. figs. 1-4, 6: text-fig. 16: mm 5 [=Ajriki,a hunanensis Peng, 1978]. 2001b Torifera paraconvexa (Yang): Peng, Babcock. and Lin (in part), p. 104, pl. 11, fig. 14. mm figs. 12, 13 [=Torifera abrupta sp. nov.]. 2001d Ajrikina hunanensis (Peng): Peng, Babcock, Lin, and Chen, p. 141. fig. 9.6. 1987

Holotype. Cranidium from the Formosagnostus formosus [=Hadragnostus modestlts]-Distazeria [Paradistazeria] Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan (Peng, 1987, pl. 13, figs. la, b; NIGP 74583). New material. More than 20 sclerites including cranidia and pygidia (illustrated specimens NIGP 137846-137853) are in collections W196.3, W199.2, W200.3, W210.5, W211.7, and W 216.5. Remarks. All observed characters of the new specimens agree with those expressed in Xiangia taoyuanensis from the Huaqiao Formation at Wa'ergang, Taoyuan, northwestern Hunan. Careful preparation of the new material revealed a tiny, short (tr.), ridge-like anterior cephalic border. The presence of this feature in the type species of Xiangia indicates that the genus should be suppressed as a junior synonym of Torifera. T. taoyuanensis resembles Torifera? paraconvexa (Yang) (see also Pl. 60, fig. 11 herein) from the same paleogeographic region, but T.? paracon~'exa is differentiated by having narrower axial furrows, more posteriorly located eye ridges, and by lacking bacculae. Occurrence. The holotype is from the Formosagnostus formostts [=Hadragnostus modesttts]Distazeria [Paradistazeria] Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Wangcun •

175.

section, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the upper part of the Proagnostus bulbus Zone to the lower part of the Linguagnostus reconditus Zone). Torifera tuma (Yang in Zhou et al., 1977)

Plate 59, figures 1-15; Text-figure 29A-C 1977 1978 1978 1982 1983 2001b

Cyclolorenzella tuma Yang in Zhou, Liu, Meng, and Sun, p. 170, pl. 50, figs. 17, 18. Cyclolorenzella tuma Yang; Yin and Li, p. 489, pl. 165, fig. 16. Cyclolorenzella tuma Yang; Yang, p. 43, pl. 7, figs. 9-11. Cyclolorenzella tuma Yang; Liu, p. 312, pl. 219, figs. 13, 15. Cyclolorenzella caijiapingensis Yang; Lin, Lin, and Zhou, p. 404, pl. 2, fig. 6. Torifera tuma (Yang); Peng, Babcock, and Lin, p. 103, pl. 7, fig. 9.

Lectotype. Cranidium (Zhou et al., 1977, pl. 50, fig. 18, CUGB 0329201; Text-fig. 29C, herein) from Laochatian, Fenghuang, western Hunan; designated subsequently as the holotype of the species by Yang (1978). New material. More than 50 sclerites, including cephala, cranidia, and pygidia (illustrated specimens NIGP 137854-137861) in collections P260, P261, P268.3, P277, P282.6, P317.4, P331.8, W171.9, W173.8, W187.8, W188.8, and W211.7. Emended diagnosis. Torifera with conical glabella; long, linear anterior border; palpebral lobe small, ovate, and located anteriorly; eye ridge wide, poorly defined; librigena narrow (tr.) with linear lateral border, short, widely-based genal spine, lacking border furrow. Description. Cephalon semicircular, except for genal spines. Glabella subtriangular, slightly longer than wide, obtusely pointed anteriorly, with incised, gently oblique S1 furrow, almost isolating small L1 lobe; $2 short, faint; baccula obscure or obliterated; occipital ring wider (tr.) than glabellar base, markedly narrowing laterally, extending rearward into stout, wide-based occipital spine, divided by pair of shallow, inward and rearward-directed furrows into a large, triangulate median part and two small ridge-like distal parts. Preglabellar field and fixigena moderately convex; preglabellar boss weak; anterior border linear, moderately long(tr.), well defined by linear border furrow. Eye ridge transverse, poorly defined, widening slightly abaxially; palpebral lobe small, semicircular, gently elevated. Anterior branch of the facial suture converging forward, slightly convex, meeting anterior cranidial margin at distance to sagittal line equal to basal glabellar width, posterior branch curved, cutting posterior border as rearward and inward curve; posterior area of fixigena broad (tr., exs.) with wide (exs.) and deep posterior border furrow, that ends abruptly before meeting lateral cranidial margin, and abaxially widened posterior border. Librigena narrow (tr.), slightly convex, with raised linear lateral border defined only by change of convexity on genal area, without border furrow; genal spine short, wide at base. Surface of cephalon covered with coarse granules; anterior part of preocular field and base of genal spine have few terrace lines. Remarks. This species was originally referred to Cyclolorenzella (Zhou et al., 1977; Yang, 1978), a genus erected by Kobayashi (1960, p. 389, 390) with Lorenzella quadrata Kobayashi (1935, p. 210, • 176.

C Text-figure 29. Reconstruction of cephalon of Torifera tuma (Yang in Zhou et al., 1977). A, B, reconstruction of cephalon based mostly on specimen NIGP 137856 (see P1. 59, figs. 3-9); C, lectotype of Cyclolorenzella tuma Yang in Zhou et al., 1977, CUGB 0329201, × 15, designated subsequently as the holotype of the species by Yang (1978), original of Zhou et al., 1977 (pl. 50, fig. 18; also Yang, pl. 7, fig. 11). pl. 12, figs. 2-5; pl. 13, figs. 2, 3) as the type species. As noted by Zhang and Jell (1987, p. 131, 132), C y c l o l o r e n z e l l a lacks an anterior border. Zhang and Jell also stated that C y c l o l o r e n z e l l a has • 177.

transverse eye ridges and small, anteriorly located palpebral lobes. These characters, however, are based mainly on specimens referred by Yang (1978) to Cyclolorenzella, which are from western Hunan, South China, rather than being based on specimens from South Korea and North China (Kobayashi, 1935; Resser and Endo, 1937; Lu, 1957; Zhu, 1959). Actually, as demonstrated by Kobayashi (1935), C. quadrata, the type species of Cyclolorenzella, is characterized by having relatively large, quite posteriorly located palpebral lobes, and oblique eye ridges. All species from North China including the type species lack an occipital spine. Among 17 species previously assigned to Cyclolorenzella (Zhang and Jell, 1987, p. 132), three species, C. tuma, C. paraconvexa, C. caijiapingensis, are from western Hunan. C. tuma and C. paraconvexa (refigured in P1. 60, fig. 11, herein) are transferred here to Torifera. C. caijiapingensis has a forward-outward directed eye ridge, a small, anteriorly located palpebral lobe, and a conical glabella. These characters suggest that it is a synonym of Fenghuangella coniforma (see remarks under F. coniforma). New material from western Hunan (Wangcun and Paibi) agrees in all respects with the type material of Torifera tuma from elsewhere in western and northwestern Hunan (Baojing and Fenghuang), and is attributed to the species. T. tuma is distinguished from all other species of Torifera in having narrow fixigenae, weakly defined eye ridges, small palpebral lobes, and coarse granulation.

Occurrence. The holotype is from Laochatian, Fenghuang, western Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the lower part of the Proagnostus bulbus Zone through the lower part of the Linguagostus reconditus Zone). Torifera abrupta sp. nov. Plate 60, figures 1-10 ?1987 Liostracina taoyuanensis Peng (in part), p. 110, 111, pl. 12, fig. 9 only. 2001 b Torifera paraconvexa (Yang); Peng, Babcock, and Lin (in part), p. 104, pl. 11, figs. 12, 13, non fig. 14 [=Torifera taoyuanensis (Peng, 1987)].

Etymology. From Latin, abruptus, abrupt or steep, referring to the steep anterior part of the preglabellar field. Holotype. Cranidium (P1.60, figs. 4, 5, NIGP 137865) in collection W227. Other material. Nine cranidia and one pygidium, of which four cranidia and one pygidium are illustrated as paratypes (NIGP 137862-137864, 137866-137868), in collections W223.2?, W277. A pygidium (NIGP 74580), assigned originally to Lioastracina taovuanensis Peng, from Wa'ergang, Taoyuan, northwestern Hunan, is reassigned to the new species. Diagnosis. Torifera with glabella truncate anteriorly, preglabellar field with anterior portion turned abruptly downward; occipital ring bearing spine; anterior border short (tr., sag.), anteriorly angulate; triangular posterolateral projections present. Description. Cranidium semicircular. Small anterior border defined by shallow and narrow border • 178.

furrow, angulate anteriorly. Glabella tapered forward moderately, acutely rounded or obtusely truncate anteriorly, defined by broad and deep axial and preglabellar furrows, occupying about two-thirds of cranidial length; S 1 short, deeply incised, strongly oblique rearward, almost isolating L1 from glabella; $2 short, well impressed, lying slightly anterior to midlength of glabella, $3 faint or absent; baccula prominent, reniform in outline, separated from glabella. Occipital furrow transverse, deep distally, shallow medially; occipital ring triangular, extending rearward into a short spine bearing weak keel sagittally. Palpebral lobe small, opposite midlength of cranidium; eye ridge linear, long, transverse or slightly oblique rearward. Preocular field broad, divided into two parts by abrupt change of slope and an obscure ridge that is broadly W-shaped in anterior view; posterior part with preglabellar boss defined laterally by pair of shallow, longitudinally subparallel furrows and gently convex posterior portion of preocular field; anterior part turned downward abruptly to form sharp and crescentic, anteriorly arched, vertical surface. Anterior branch of facial suture curving gently inward to cut anterior border obliquely and meet anterior margin sagittally; posterior branch diverging rearward moderately, enclosing triangular posterolateral projection. Posterior border furrow and posterior border widening abaxially. Surface with fine, moderately spaced granules of similar size. Pygidium transverse, width greater than twice length. Axis conical, extending rearward to posterior margin, with crescentic articulating half ring, four tings and tiny terminal piece. Articulating furrow and anterior two ring furrows with paired, transversely elongated pits. Pleural field undivided with dense, raised terrace lines. Lateral border broad and shallow, lateral border linear.

Remarks. Torifera abrupta sp. nov. is similar to Torifera taoyuanensis (Peng), which has a slightly lower occurrence in the Wangcun section, Hunan. Both species have broad and deep axial and preglabellar furrows, prominent bacculae, and a short (tr., sag.) anterior border. T. abrupta differs from T. taoyuanensis, however, in having a more strongly tapering glabella, a divided preocular field, an angulate margin of the anterior border, and a pygidium that has a pointed anterolateral comer. In T. taoyuanensis, the preocular field curves continuously downward and forward without an edge-forming slope change, the anterior margin of the cranidial anterior border is smoothly curved, and the pygidium is not pointed but obliquely truncated distally. Torifera triangularis, the type species of Torifera, differs from T. abrupta sp. nov. in having a wider (sag., tr.) anterior border, a broad (sag., exs.) anterior border furrow, a triangular occipital ring that is wide at the base, and a less transverse pygidial outline. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is associated with trilobites indicative of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Torifera? paraconvexa (Yang in Yin and Li, 1978) Plate 60, figure 11 1978 Cyclolorenzella paraconvexa Yang; Yin and Li, p. 488,489, pl. 165, fig. 17. 1978 Cyclolorenzella paraconvexa Yang; Yang, p. 42, 43, pl. 7, fig. 7.

Holotype. Incomplete cranidium (Yang, 1978, pl. 7, fig. 7; P1.60, fig. 11 herein, CUGB 100400), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Liangweizhou, Yuping, eastern Guizhou. • 179.

Remarks. This species is represented only by the holotype cranidium. The holotype is refigured here to show its morphological resemblance to specimens assigned here to Torifera from western Hunan. Although originally described (Yang, 1978) in Cyclolorenzella, the specimen shows greater morphologic resemblance to Torifera, and is transferred to that genus with question. The figure (P1.

60, fig. 11) is a photograph of a replica from the latex cast of the holotype; the whereabouts of the original specimen is unknown. Family

LIOSTRACINIDAE

Raymond, 1937

Genus LIOSTRACINAMonke, 1903 Type species. Liostracina kraisei Monke, 1903 (p. 114-117, pl. 3, figs. 10-17) from the Drepanura

Zone, Kushan Formation, near Yanzhuang, Shandong; by original designation. Remarks. The concept of the genus has been discussed by a number of authors (Walcott, 1913;

Resser and Endo, 1937; Raymond, 1937; Endo, 1944; Kobayashi, 1962; Opik, 1967; Zhang and Jell, 1987). Opik (1967, p. 352, 253) expanded the genetic concept slightly by adding two new species, and that concept is followed here. Liostracina bella Lin and Zhou, 1983

Plate 61, figures 1-14; Plate 62, figures 1-13" Text-figure 30 1963

Liostracina krausei Monke; Egorova, Xiang, Li, Nan, and Guo (in part), p. 42, pl. 7, fig. 5, non fig. 6 [=Fenghuangella liostracinala Yang in Zhou et al., 1977]. 1977 Liostracina krausei Monke; Zhou et al., p. 210. pl. 61, fig. 18. 1982 Liostracina krausei Monke; Liu, p. 324, pl. 223, fig. 15. 1983 Liostracina bella Lin and Zhou in Lin, Lin, and Zhou, p. 407,408, pl. 3, figs. 7-10. 1987 Liostracina bella Lin and Zhou; Peng, p. 110, pl. 12, fig. 7. 1987 Liostracina taoyuanensis Peng, p. 110, 111, pl. 12, figs. 8, 9. 2001 c Liostracina bella Lin and Zhou; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 3, fig. 12. 2001b Liostracina bella Lin and Zhou" Peng, Babcock, and Lin, p. 105, pl. 13, figs. 1, 2. 200 ld Liostracina bella; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.3. Holotype. An incomplete cranidium (Lin et al., p. 407, 408, pl. 3, fig. 7, NIGP 67134), from the

Tuanshan Formation, Kunshan, southern Jiangsu. New material. More than 50 sclerites, including cranidia, librigenae, and pygidia (illustrated

specimens NIGP 137869-137889), in collections P308, P310.4, P316.1, P317.4, P319.8, P331.8, P337.5, P341.7, P341.8, P346.7, P361.5, PI3-2.75, W218.3, W221.5, W225, W227, W243.6, and W254.1. Remarks. The type material of Liostracina bella includes three poorly preserved cranidia and a

librigena from drill cores at Kunshan, southern Jiangsu. The holotype cranidium is incomplete, without an occipital ring, posterolateral projections, and palpebral lobes. It is characterized by having a curved ridge immediately behind the anterior border furrow, a strongly ananstomose anterior field of the fixigena, a small, subcylindrical glabella bearing three pairs of short lateral furrows, and a pair of large, clearly defined bacculae. The type material shows that the eye ridge is 180. •

variably directed and that the front of the glabella varies in appearance from acutely rounded to obtusely rounded.

~t_

r -

T

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-

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Text-figure 30. Reconstruction of cephalon and pygidium of Liostracina bella Lin and Zhou, 1983. Cephalon based on specimens NIGP 137877, 137878 and 137883, (see PI. 61, figs. 12, 13; P1.62, fig. 7); pygidium based on specimen NIGP 137886 (see PI. 62, fig. 10). The cranidia in the new material are almost identical with cranidia in the type material, but the associated librigena differs from a librigena in the type material. The paratype librigena has a rather smooth genal field, a nearly completely effaced paradoublural line, shallow border furrows, and a less upturned lateral border. These distinctions suggest that the specific assignment for the paratype librigena is questionable, although this librigena is found in association with the holotype cranidium. The librigena could belong to Liostracina. Occurrence. The holotype is from the Tuanshan Formation, Kunshan, southern Jiangsu, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2, and Wangcun sections, Hunan, where it is the eponymic species of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone through the lower part of the Glyptagnostus reticulatus Zone).

Superfamily UNCERTAIN Family MONKASPIDIDAE Kobayashi, 1935 Genus MONKASPIS Kobayashi, 1935 Monkaspis Kobayashi, 1935, p. 289, 300; 1960a, p. 237, 268; Howell in Moore, 1959, p. 0288; Lu, 181. •

Zhang, Zhu, Qian, and Xiang, 1965, p. 164; Zhou, Liu, Meng, and Sun, 1977, p. 149; Yang, 1978, p.37; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 92; Zhang and Wang, 1985, p. 368; Peng, 1987, p. 95; Zhang and Jell, 1987, p. 191, 192; Xiang in Zhang et al., 1995, p. 79; Guo, Zan, and Luo, 1996, p. 90. Liaoningaspis Chu (Zhu), 1959, p. 74, 75, 118, 119; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p.167, 168; Lu, Zhu, Qian, Lin, Zhou, and Yuan, 1974a, p. 95, 96; Lu, Zhang, Qian, Zhu, Lin, Zhou, Qian, Zhang, and Wu, 1974b, p. 106; Zhou, Liu, Meng, and Sun, 1977, p. 150; Li, 1978, p. 229; Nan, 1980, p. 490; Luo, 1982, p. 2; Zhang and Wang, 1985, p. 336. Kushanopyge Chu (Zhu), 1959, p. 77, 122; Kobayashi, 1960a, p. 269. Paraliaoningaspis Chu (Zhu) in Lu et al., 1965, p. 168; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 92.

Type species. Anomocare? daulis Walcott, 1905 (p. 50, 51; 1913, p. 189, 190, pl. 18, figs. 7, 7a), from the Kushan Formation, Zhangxia, Shandong; by original designation. Emended diagnosis. Cranidium with rounded or obtusely rounded anterior margin; frontal area wide (sag., exs.), gently depressed, with short anterior border and shallow anterior border furrow; paradoublural line coinciding with or lying posterior to preglabellar furrow; glabella parallel-sided to gently tapered forward, rounded to obtusely rounded anteriorly with bacculae and faint lateral furrows; palpebral lobe located posteriorly. Thorax with 12 segments, axis much narrower than pleural region. Pygidium with two to eight pairs of pleural spines; axis with four to six rings and long terminal piece; postaxial ridge variably present, if present, wide; pleural furrow incised, reaching base of spine; doublure wide, with inner margin at or lying slightly anterior to end of axis.

Remarks. Kobayashi (1960a) remarked that Kushanopyge may be not a separate genus from Monkaspis. Zhang and Jell (1987, p. 190, 191) synonymized Kushanopyge Chu (Zhu), 1959, and Paraliaoningaspis Zhu in Lu et al., 1965 as junior synonyms of Monkaspis. Kobayashi (1960a, p. 268) and Xiang (in Zhang et al., 1995, p. 79) noted the close similarity of Liaoningaspis Chu (Zhu), 1959 with Monkaspis, and considered the genus to be possibly congeneric with Monkaspis or to be a possible subgenus of it. Liaoningaspis is here regarded as a junior synonym of Monkaspis. As noted by Zhang and Jell (1987), Liaoningaspis differs only in the size of the baccula, the degree of arc of the palpebral lobes, and the shape and segmentation of the pygidium. However, the first two differences are minor, and they vary even within individual species of Monkaspis (e.g., Monkaspis quadrata Yang from western Hunan). The pygidium of Liaoningaspis taitzehoensis, the type species of Liaoningaspis, differs from that of Monkaspis serrata Mong (Zhou et al., 1977, pl. 46, figs. 22-25; Zhang and Jell, 1987, pl. 86, figs. 4-6) only in having fewer pairs of pleural spines (two pairs rather than seven to eight pairs), and an entire posterior margin between the spines. In these respects, Liaoningaspis is comparable with Kushanopyge except that Kushanopyge has two or three more pairs of marginal spines. In Monkaspis, as in some other genera of trilobites (e.g., various aphelaspidids, damesellids, cheirurids), the number of marginal spines on the pygidium is variable, and is commonly regarded as a character of species-level significance. In pygidial characteristics, Kushanopyge shows features intermediate between those of Liaoningaspis and Monkaspis. By suppressing Kushanopyge and Liaoningaspis as junior synonyms of Monkaspis, Monkaspis includes a series of species having a variable number of pleural spines in the pygidium.

• 182.

Monkaspis quadrata Yang in Zhou et al., 1977 Plate 63, figures 1-13; Text-figure 31 1977 1978 1982 200 lb

Monkaspis quadratus Yang in Zhou et al., p. 150, pl. 46, figs. 17, 18. Monkaspis quadratus Yang; Yang, p. 37, pl. 5, figs. 9-11. Monkaspis quadratus Yang; Liu, p. 306, pl. 216, figs. 15, 16. Monkaspis quadrata Yang; Peng, Babcock, and Lin, p. 104, pl. 10, figs. 3-6.

Lectotype. Cranidium (Zhou et al., 1977, pl. 46, fig. 17; also Yang, 1978, pl. 5, fig. 9; CUGB 231005; Text-fig. 31 herein), from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Laochatian, Fenghuang, western Hunan; designated subsequently as the holotype of the species by Yang (1978). New material More than 15 sclerites, including cranidia, and pygidia (illustrated specimens NIGP 137890-137900), in collections P282.6, P298.4, P298.54, P319.6, P341.7, W210.5, W211.7, W212.45, W215, W218.3, and W223.2 (?). Remarks. Monkaspis quadrata from the Huaqiao Formation, western Hunan and eastern Guizhou, differs from M. daulis (Walcott, 1905), the type species of Monkaspis, in having a glabella that is less tapered forward and less rounded anteriorly, and a more strongly curved anterior cranidial margin. It is different from M. hunanensis Peng, from the Huaqiao Formation in northwestern Hunan, which has a proportionally longer, more strongly tapering glabella that bears more constricted sides, and an acutely rather than obtusely rounded front. M. hunanensis can be further differentiated by having a cranidium with a more rounded anterior margin, a more rearward-oblique eye ridge, and a subtriangular rather than blade-like posterolateral projection.

Text-figure 31. Lectotype of Monkaspis quadrata Yang in Zhou et aL, 1977, a cracked cranidium, designated subsequently as holotype of the species by Yang (1978, p. 37), CUGB 0231005, x 4.6, original of Zhou et al., 1977 (pl. 46, fig. 17; also Yang, 1978, pl. 5, fig. 9). New material of M. quadrata from western Hunan resembles the type material of the species. Key characters that warrant placement of the new material into this species include a short subquadrate glabella; a flat, slightly rounded anterior margin of the cranidium; and pygidial pleural 183•

furrows that extend nearly to the tips of the pleural spines. Juvenile cranidia in the new material show that in early ontogeny, the glabella is parallel-sided rather than tapered, and it is proportionally longer than in large holaspids.

Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, Laochatian, Fenghuang, western Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is associated with trilobites indicative of the upper part of the Wanshania wanshanensis Zone through the Liostracina bella Zone (equivalent to the upper part of the Proagnostus bulbus Zone through the lower part of the Linguagnostus reconditus Zone). Genus METOPOTROPIS Opik, 1967

Type species. Metopotropis travesi Opik, 1967, from the Glyptagnostus stolidotus Zone, Georgina Limestone, Queensland, Australia; by original designation.

Other species. Metopotropis sinensis sp. nov. In addition, Sigmocheillus orientalis Lazarenko and Nikforov (1968, p. 44-46, pl. 10, figs. 20, 21) from the Faciura-Garbiella Zone, northwestern Siberian Platform questionably belong to the genus. Remarks. Metopotropis is here assigned to the Monkaspididae. When Opik (1967, p. 224) erected the genus, he regarded it as an anomocaroidean and classified it as subfamily uncertain within the family Austitamidae. However, the absence of a defined anterior border, the depressed and anteriorly upturned frontal area, and the presence of well-defined bacculae, are inconsistent with classification within the superfamily Anomocaroidea. These characters are more consistent with classification in the family Monkaspididae. This interpretation is reinforced by the gently tapered to cylindrical glabella, which is comparable to that of immature Monkaspis (e.g., M. quadrata; P1.63, figs. 1, 2).

Metopotropis sinensis sp. nov. Plate 51, figures 1-4 2001b Monkaspid gen. et sp. nov., Peng, Babcock, and Lin, p. 105, pl. 15, fig. 3.

Etymology. From Latin, Sinae, China, referring to the discovery of the genus in China. Holotype. Cranidium (Plate 51, figs. 1, 2, NIGP 137772) in collection P363.5. Other material. One fragmentary cranidium, NIGP 137773, in collection P319.8. Diagnosis. Metopotropis with glabella that is gently tapered, acutely rounded anteriorly, with weak lateral furrows; anterior branch of facial suture diverging at 35o-50 ° to sagittal line; palpebral lobe markedly arcuate and oblique.

Description. Cranidium subquadrate, length equal to width at anterior margin. Glabella sub• 184.

rectangular, moderately convex, parallel-sided behind glabellar midpoint, tapering slightly forward anteriorly to midpoint, rounded anteriorly, occupying about two-thirds of total glabellar length, bearing three pairs of shallow lateral glabellar furrows. Occipital furrow deep, gently curved rearward; occipital ring with even width. Bacculae ovate, distinctive. Anterior area wide (sig., tr.), gently upturned anteriorly, with broad and shallow anterior border furrow, gently wavy transverse ridge close to preglabellar furrow medially; preglabellar field behind transverse ridge depressed. Palpebral lobe long, notably arcuate and oblique, located posteriorly, with posterior end slightly anterior to occipital furrow and anterior end near anterolateral comer of glabella; palpebral area narrow, fiat, half width of glabella. Anterior branch of facial suture diverging forward about 350-50 ° to sagittal line, then curving sharply after crossing anterior border furrow, meeting anterior margin, at a point oppsite the anterior end of palpebral lobe; posterior branch and posterior area of fixigena unknown. Surface of anterior area of fixigena with dense wrinkles.

Remarks. In having a tropidium-bearing anterior area, a long and gently tapered glabella, and prominent, elongated bacculae, the new species is similar to Metopotropis travesi 0pik (1967, p. 224, 225, pl. 16, figs. 9a, b-10; text-fig. 77), the type species of Metopotropis, from Queensland, Australia. However, the Australian species has strongly diverging anterior branches of the facial suture, which are almost transverse before reaching the cranidial margin; a nearly completely effaced glabella that is obtusely rather than acutely rounded anteriorly; and a longer, less oblique and less arcuate palpebral lobe. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is associated with trilobites indicative of the upper part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone to the Glyptagnostus stolidotus Zone).

• 185.

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• 200.

INDEX

OF SPECIES

AND GENERA

A a b r u p t a , T o r i f e r a .............................................................................................................................. 178 alceste, A m p h o t o n ... ........................................................................................................................... 57 A f g h a n o c a r e ..................................................................................................................................... 150 t r u n c a t u m ................................................................................................................................. 150 A j r i k i n a ............................................................................................................................................ 171 h u n a n e n s i s ................................................................................................................................ 173 w a n n a n e n s i s ............................................................................................................................. 172 A m p h o t o n ........................................................................................................................................... 54 alceste ......................................................................................................................................... 57

cf. t y p i c u m .................................................................................................................................. 59 deois ........................................................................................................................................... 56 a u s t r i a c a , C h u a n g i a ........................................................................................................................... 80

B bisulcata, D o r y p y g e ........................................................................................................................... 75 B l a c k w e l d e r i a .................................................................................................................................... 98 y o u s h u i c a ................................................................................................................................... 99 B l a c k w e l d e r i a ? sp.. .......................................................................................................................... 100 bella, L i o s t r a c i n a ............................................................................................................................. 180 bella, M e r o p a l l a ................................................................................................................................. 82 bella, P a l a e a d o t e s ............................................................................................................................ 136 bulba, F u c h o u i a ................................................................................................................................. 67 C carinata, Y a n g w e i z h o u i a .................................................................................................................. 161 C h a t i a n i a ............................................................................................................................................ 48

cf. c h a t i a n e n s i s .......................................................................................................................... 53 c h a t i a n e n s i s ................................................................................................................................ 49 c o n v e x a ....................................................................................................................................... 53 e x p a n s a ...................................................................................................................................... 52

cf. c h a t i a n e n s i s , C h a t i a n i a ................................................................................................................ 53 cf. h u a b e i e n s i s , P a r a b l a c k w e l d e r i a ................................................................................................. 109 cf. kingi, P a r a c o o s i a

154

cf. l e i o c e p h a l a , P r o c h u a n g i a ............................................................................................................. 95 cf. t y p i c u m , A m p h o t o n ....................................................................................................................... 59 c h a t i a n e n s i s , C h a t i a n i a ...................................................................................................................... 49 chiai, F u c h o u i a .................................................................................................................................. 62 C h i a w a n g e l l a ..................................................................................................................................... 85 • 201 •

h u n a n e n s i s ..................................................................................................................................

86

C h u a n g i a ............................................................................................................................................

79

a u s t r i a c a ....................................................................................................................................

80

s u b q u a d r a n g u l a t a ......................................................................................................................

79

c o n v e x a , C h a t i a n i a ............................................................................................................................

53

C o r y n e x o c h i n a ...................................................................................................................................

46

sinensis .......................................................................................................................................

47

C o r y n e x o c h u s .....................................................................................................................................

44

xiangxiensis .................................................................................................................

45

c r a s s i s p i n a , D r e p a n u r a ? .................................................................................................................. 129

D D a m e s e l l a ..........................................................................................................................................

96

h u n a n e n s i s ..................................................................................................................................

96

D a m e s e l l a ? s p . . .................................................................................................................................. 9 8 deois, A m p h o t o n .................................................................................................................................

56

D o r y p y g e ............................................................................................................................................

70

b i s u l c a t a .....................................................................................................................................

75

g l o b o s a .......................................................................................................................................

77

h u a q i a o e n s i s ..............................................................................................................................

76

p e r c o n v e x a l i s .............................................................................................................................

73

r i c h t h o f eni ..................................................................................................................................

71

D o r y p y g e ? s p . . ................................................................................................................................... 7 8 D r e p a n u r a ........................................................................................................................................ 128 D r e p a n u r a ? c r a s s i s p i n a ................................................................................................................... 129

E e x p a n s a , C h a t i a n i a ............................................................................................................................

52

F f e n g h w a n g e n s i s , P r o c e r a t o p y g e ( P r o c e r a t o p y g e ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163 F u c h o u i a ............................................................................................................................................

60

bulba ..........................................................................................................................................

67

chiai ............................................................................................................................................

62

k u r u k t a g e n s i s .............................................................................................................................

64

o r a t o l i m b a ..................................................................................................................................

65

s & i n e n s i s ....................................................................................................................................

68

sp. i n d e t . •.................................................................................................................................... 6 9

f u y a n g e n s i s , P r o c e r a t o p y g e ( P r o c e r a t o p y g e ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164 G g a o l o u p i n g e n s i s , G a o l o u p i n g i a ....................................................................................................... 170 G a o l o u p i n g i a ................................................................................................................................... 170 g a o l o u p i n g e n s i s ....................................................................................................................... 170 G e m i n i c l a v u l a ....................................................................................................................................

83

w a n g c u n i c a ................................................................................................................................

84

g i b b e r a , M e r o p a l l a ............................................................................................................................

82

g l o b o s a , D o r y p y g e .............................................................................................................................

77

G l y p h a s p e l l u s ? sinensis ................................................................................................................... 152 • 202 •

granifera, Protaitzehoia ................................................................................................................... 115 granulosa, Prochuangia .................................................................................................................... 9 4

H huaqiaoensis, Dorypyge ..................................................................................................................... 76 huayuanensis, Paracoosia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5 6 hunanensis, Ajrikina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 7 3 hunanensis, Chiawangella ................................................................................................................. 8 6 hunanensis, Damesella ........................................................................................................ 9 6 hunanensis, Palaeadotes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3 2 hunanica, Paraacidaspis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 6

J jimaensis, Parablackwelderia .......................................................................................................... 101

K kuruktagensis, Fuchouia .................................................................................................................... 6 4 L laochatianensis, Pseudoyuepingia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 7 laterilobata, Parablackwelderia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 8 lineatum, Paibianomocare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5 9 linicispinata, Prochuangia ................................................................................................................. 9 5 Liostracina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 8 0 bella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 8 0

M Meropalla ........................................................................................................................................... 81 bella ............................................................................................................................................ 8 2 gibbera ....................................................................................................................................... 8 2 Metopotropis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 8 4 sinensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 8 4 Monkaspis ........................................................................................................................................ 181 quadrata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 8 3

N nobilis, Paradamesella

.................................................................................................................... 145

0 oratolimba, Fuchouia ........................................................................................................................ 6 5

P Pagodiinae

gen. et sp. indet.. .............................................................................................................. 93

Paibianomocare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lineatum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . paibiense . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . paibiense, Paibianomocare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palaeadotes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

157 159 158 158 130 136

• 203.

hunanensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraacidaspis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hunanica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

132

sp.. ............................................................................................................................................

149

Parablackwelderia ........................................................................................................................... c f . huabeiensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . jimaensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . laterilobata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . yangi ........................................................................................................................ sp.. ............................................................................................................................................

146 147

101 109 104

108 110 111

paraconvexa, Torifera? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 7 9 Paracoosia ....................................................................................................................................... 153 huayuanensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5 6 c f . kingi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154 Paradamesella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3 7 nobilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 5 peculiaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 2 typica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3 9 peculiaris, Paradamesella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 2 perconvexalis, Dorypyge .................................................................................................................... 7 3 posterocostum, Teinistion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 3 Proceratopyge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 2 Proceratopyge (Proceratopyge) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 3 fenghwangensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 3 fuyangensis ............................................................................................................................... 164 truncata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 5 Proceratopyge s p . i n d e t . •. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 6 6 Prochuangia ....................................................................................................................................... 9 3 c f . leiocephala ............................................................................................................................ 9 5 granulose .................................................................................................................................... 9 4 linicispinata ................................................................................................................................ 9 5 Protaitzehoia .................................................................................................................................... 112 granifera .................................................................................................................................. 115 spinifera ................................................................................................................................... 119 subquadrata ............................................................................................................................. 117 yuepingensis ............................................................................................................................. 114 sp.. ............................................................................................................................................ 120 Pseudoyuepingia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . laochatianensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

166 167

Q quadrata, Monkaspis

........................................................................................................................

183

R richthofeni, Dorypyge ........................................................................................................................ 71

S sinensis, Corynexochina .................................................................................................................... 4 7 sinensis, Glyphaspellus? .................................................................................................................. 1 5 2 • 204.

sinensis, M e t o p o t r o p i s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

184

sixinensis, Fuchouia ........................................................................................................................... 6 8 s p . , Blackwelderia? .......................................................................................................................... 1O0 s p . , Damesella? .................................................................................................................................. 9 8 s p . , Dorypyge? ................................................................................................................................... 7 8 s p . , Paraacidaspis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

149

s p . , P a r a b l a c k w e l d e r i a .................................................................................................................... 111 s p . , Protaitzehoia ............................................................................................................................. 1 2 0 sp. i n d e t . , Fuchouia ............................................................................................................................ 6 9 sp. i n d e t . , P a g o d i i n a e

g e n . i n d e t . •. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

sp. i n d e t . , Proceratopyge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

93

166

spinifera, Protaitzehoia .................................................................................................................... 119 subquadrangulata, Chuangia ............................................................................................................ 7 9 subquadrata, Protaitzehoia .............................................................................................................. 117 T Taihangshania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

125

wangcunensis ...........................................................................................................................

126

taoyuanensis, Torifira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175 Teinistion .......................................................................................................................................... 121 posterocostum ..........................................................................................................................

123

Torifira ............................................................................................................................................. 1 7 4 abrupta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

178

taoyuanensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

175

tuma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Torifera? p a r a c o n v e x a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

179

176

truncata, P r o c e r a t o p y g e (Proceratopyge) .......................................................................................

165

truncatum, A f g h a n o c a r e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

150

tuma, Torifira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176 typica, P a r a d a m e s e l l a ...................................................................................................................... 1 3 9

W wangcunensis, Taihangshania ......................................................................................................... 1 2 6 wangcunica, Geminiclavula ............................................................................................................... 8 4 Wanshania .......................................................................................................................................... 8 9 w a n s h a n e n s i s ............................................................................................................................. 9 0 wannanensis, Ajrikina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

172

wanshanensis, Wanshania ................................................................................................................. 9 0

X xiangxiensis, C o r y n e x o c h u s ............................................................................................................... 4 5 Y yangi, Parablackwelderia ................................................................................................................ 110 Yangweizhouia ................................................................................................................................. 161 carinata .................................................................................................................................... 161 yuepingensis, Protaitzehoia ............................................................................................................. 114 youshuica, Blackwelderia .................................................................................................................. 9 9

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PLATES All illustrated specimens were coated with black ink and then magnesium oxide before being photographed. All specimens are preserved in limestone. Specimens having a horizon number with the prefix P are from the Paibi section; those with the prefix PI3 are from the Paibi-2 section; and those with the prefix W are from the Wangcun section. Unless otherwise stated, all specimens are deposited in the collections of the Nanjing Institute of Geology and Palaeontology, the Chinese Academy of Sciences (NIGP).

• 207.

Plate 1

Figures 1-10, ? 11, ? 12. Corynexochusxiangxiensis sp. nov.; all from P3.2. 1. Juvenile cranidium, NIGP 137282, × 20. 2, 3. Slightly exfoliated cranidium in dorsal and anterolateral views, NIGP 137283, × 12. 4. Hypostome, NIGP 137284, × 20. 5-7. Mostly exfoliated cranidium in anterolateral, dorsal, anterolateral, and anterior views, holotype, NIGP 137285, × 12. 8, 9. Testaceous pygidium, in dorsal and posterolateral views, NIGP 137286, × 17. 10. Broken testaceous pygidium, NIGP 137287, × 18. 11, 12. Small, testaceous cranidium in dorsal and anterolateral views, NIGP 137288, × 20. From Dorypyge richthofeni Zone. Figures 13-16. Corynexochina sinensis sp. nov.; all from W3. 13, 14. Incomplete, mostly exfoliated cranidium in dorsal and anterolateral views, holotype, NIGP 137289, × 12. 15. Hypostome, NIGP 137290, × 25. 16. Incomplete, mostly exfoliated pygidium, NIGP 137291, × 15. From Dorypyge richthofeni Zone.

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Figures 1-7. Corynexochina sinensis sp. nov. 1, 2. Small, nearly complete cranidium in dorsal and anterolateral views, NIGP 137292, x 22, W0. 3, 4. Possibly exfoliated cranidium, in dorsal and anterolateral views, NIGP 137293, × 12, W3. 5-7. Complete testaceous cranidium in dorsal, obliquely anterior, and anterolateral views, NIGP 137294, × 15, W0. Figures 1, 2, 5-7, unnamed zone; Figures 3, 4, Doo'pyge richthofeni Zone. Figures 8, 9, 13, 14. Chatiania chatianensis Yang in Zhou et al., 1977 8. Small, testaceous cranidium, NIGP 137295, × 22, P337.5. 9. Slightly exfoliated cranidium, NIGP 137296, × 10, P337.5. 13, 14. Testaceous librigena in anterolateral and dorsal views, NIGP 137297, × 10, P337.5. All from Liostracina bella Zone. Figures 10-12. Gaoloupingia gaoloupingensis Yuan and Yin, 1998 10, 12. Incomplete cranidium and the latex cast from its external mold, NIGP 137298, × 20, P331.8. 11. Testaceous cranidium, NIGP 137299, × 25, W216.5. Figures 10, 12 from Liostracina bella Zone; Figure 11 from upper most Wanshania wanshan-

ensis Zone.

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Plate 3

Figures 1-11. Chatiania chatianensis Yang in Zhou et al., 1977 1. Small, incomplete, mostly exfoliated cranidium, NIGP 137300, x 20, W254.1. 2. Small, testaceous cranidium, NIGP 137301, x 18, P311.7. 3. Testaceous cranidium, NIGP 137302, x 10, P311.7. 4. Small, testaceous cranidium, NIGP 133519, x 15, P[~-2.75. 5. Incomplete, testaceous cranidium, NIGP 137303, x 12, W254.1. 6. Testaceous cranidium, NIGP 137304, x 9, P331.8. 7. Small, testaceous pygidium, NIGP 137305, x 20, P317.4. 8, 9. Testaceous pygidium in dorsal and posterolateral views, NIGP 137306, x 10, P254.1. 10. Mostly exfoliated pygidium, NIGP 137307, P331.8, x 8. 11. Testaceous pygidium in dorsal and posterolateral views, NIGP 137308, x 10, P311.7. All from Liostracina bella Zone. Figures 12-16. Chatiania sp. cf. C. chatianensis Yang in Zhou et al., 1977 12, 13. Testaceous cranidium in dorsal and anterolateral views, NIGP 137309, x 10, P337.5. 14. Testaceous cranidium, NIGP 137310, x 12, P319.8. 15. Testaceous cranidium, NIGP 137311, x 12, P319.8. 16. Testaceous pygidium, NIGP 137312, x 8, P341.8. All from Liostracina bella Zone.

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Plate 4

Figures 1-14. Chatiania expansa (Yuan and Yin, 1998) 1. Testaceous, meraspid cranidium, NIGP 137313, × 20, W225. 2. Testaceous, possibly meraspid, cranidium, NIGP 137314, × 20, W225. 3. Nearly complete, testaceous cranidium, NIGP 137315, × 18, W213. 4. Nearly complete, testaceous cranidium, NIGP 137316, × 12, W227. 5, 6. Testaceous cranidium in anterolateral and dorsal views, NIGP 137317, × 15, W225. 7, 8. Testaceous cranidium in dorsal and anterolateral views, NIGP 137318, × 15, W225. 9, 10. Testaceous cranidium in dorsal and anterolateral views, NIGP 137319, × 15, W227. 11. Testaceous cranidium, NIGP 137320, × 9.6, P308. 12, 13. Testaceous pygidium in dorsal and posterolateral views, NIGP 137321, × 15, W225. 14. Testaceous cranidium, NIGP 137322, × 10, W225. Figures 1, 2, 4-10, 12-14 from Liostracina bella Zone; Figures 3, 11 from upper Wanshania wanshanensis Zone.

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Plate 5

Figures 1-9. Chatiania convexa sp. nov. 1. Nearly complete, testaceous cranidium, NIGP 137323, × 8.5, P278.1. 2-4. Mostly exfoliated cranidium in dorsal, lateral, and frontal views, NIGP 137324, × 11, P282.6. 5-7. Testaceous cranidium in dorsal, anterolateral, and obliquely anterior views, holotype, NIGP 137325, × 9.4, P273.66. 8, 9. Testaceous cranidium in dorsal and anterior views, NIGP 137326, × 12, P269. All from Wanshania wanshanensis Zone. Figures 10-16. Amphoton sp. cf. A. typicum (Kobayashi, 1942a) 10. Incomplete, slightly exfoliated cranidium, NIGP 137327, × 4, P125. 11. Slightly exfoliated cranidium, NIGP 137328, × 5, P123.6. 12. Partly exfoliated cranidium, NIGP 137329, × 3, P122.4. 13. Incomplete cranidium, NIGP 137330, × 4, P122.4. 14. Partly exfoliated librigena, NIGP 137331, × 4, P122.4. 15, 16. Mostly exfoliated pygidium in lateral and dorsal views, NIGP 137332, × 3, P126. All from Pianaspsis sinensis Zone.

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Plate 6

Figures 1-9. Amphoton sp. cf. A. typicum (Kobayashi, 1942a) 1. Incomplete, exfoliated cranidium, NIGP 137333, x 4, W82.1. 2. Partly exfoliated cranidium, NIGP 137334, x 4, P 102.5. 3. Incomplete, mostly exfoliated cranidium, latex cast, NIGP 137335, x 3, P108. 4. Mostly exfoliated cranidium, NIGP 137336, x 2, P126. 5, 6. Partly exfoliated pygidium in dorsal and anterolateral views, NIGP 137337, × 6, P122.4. 7. Mostly exfoliated pygidium, NIGP 137338, x 6, P123.6. 8. Mostly exfoliated, incomplete pygidium, NIGP 137339, x 5, P122.4. 9. Mostly exfoliated, incomplete pygidium, NIGP 137340, x 5, P131.7. Figures 1, 4-9 from Pianaspsis sinensis Zone; Figures 2, 3 from Dorypyge richthofeni Zone. Figures 10-12. Amphoton deois (Walcott, 1905) 10. Testaceous pygidium, NIGP 137341, x 3, P42.5. 11. Incomplete, testaceous pygidium, NIGP 137342, x 5, P42.5. 12. Exfoliated pygidium, NIGP 137343, x 5, P42.5. From Dorypyge richthofeni Zone. Figures 13, 14. Amphoton alceste (Walcott, 1905) 13, 14. Incomplete, testaceous cranidium in lateral and dorsal views, NIGP 137344, x 4, P134.2 From Pianaspsis sinensis Zone.

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Figures 1-6. Amphoton alceste (Walcott, 1905) 1, 2. Incomplete, testaceous cranidium in lateral and dorsal views, NIGP 137345, × 4, P135. 3, 4. Incomplete, testaceous cranidium in lateral and dorsal views, NIGP 137346, × 3, P136.3. 5. Exfoliated librigena, NIGP 137347, × 4, P 115.6. 6. Incomplete, exfoliated cranidium, NIGP 137348, × 2.25, P108. Figures 1-5 from Pianaspsis sinensis Zone; Figure 6 from Dorypyge richthofeni Zone. Figures 7, 8. Fuchouia chiai Lu, 1957; all from P171.5. 7. Mostly exfoliated cephalon with two segments attached, latex cast, NIGP 137349, × 3, showing slender genal spine. 8. Incomplete exfoliated cranidium, NIGP 137350, × 4. From Pianaspsis sinensis Zone. Figures 9-11. Fuchouia sp. indet. 9. Incomplete cranidium, NIGP 137351, × 2.5, W 111.3. 10, 11. Broken cranidium in dorsal and anterolateral views, NIGP 137352, × 3, W146.2. From Pianaspsis sinensis Zone.

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Plate 8

Figures 1-15. Fuchouia kuruktagensis Zhang, 1981 1, 2. Meraspid cranidium in dorsal and anterolateral views, NIGP 137353, x 20, W88.1. 3. Small testaceous cranidium, NIGP 137354, x 8, W56.7. 4. Incomplete, small testaceous cephalon, NIGP 137355, x 8, W56.7. 5. Small exfoliated cranidium, NIGP 137356, x 10, W88.1. 6. Librigena, NIGP 137357, x 5, W88.1. 7. Exfoliated cranidium, NIGP 137358, x 8, W88.1. 8. Partly exfoliated cranidium, NIGP 137359, x 10, W56.7. 9, 10. Hypostome in lateral and dorsal views, NIGP 137360, x 10, W88.1. 11. Testaceous cranidium, NIGP 137361, x 10, W56.7. 12. Partly exfoliated cranidium, NIGP 137362, x 8, W88.1. 13. Incomplete, crashed exoskeleton, NIGP 137363, x 2, W56.7. 14, 15. Slightly exfoliated cranidium in anterolateral and dorsal views, NIGP 137364, x 6, W88.1. All from base of Pianaspsis sinensis Zone.

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Figures 1-8. Fuchouia kuruktagensis Zhang, 1981 1. Partly exfoliated cranidium, NIGP 137365, × 6, W77.8. 2. Partly exfoliated cranidium, NIGP 137366, × 6, P64.5. 3. Partly exfoliated hypostome, NIGP 137367, × 7.5, W82.1. 4. Testaceous pygidium, NIGP 137368, × 8, W77.8. 5. Testaceous pygidium, NIGP 137369, × 6, W56.7. 6. Testaceous pygidium, NIGP 137370, × 6, W56.7. 7. Incomplete, slightly exfoliated pygidium, NIGP 137371, × 6, P.68.8. 8. Pygidium, NIGP 137372, × 4, W56.7. All from Dorypyge richthofeni Zone. Figures 9-14. Fuchouia sixinensis sp. nov. 9. Small, partly exfoliated cranidium, NIGP 137373, x 10, W42. 10. Small, testaceous cranidium, NIGP 137374, x 5, P82.5. 11. Testaceous cranidium, NIGP 137375, x 8, P64.5. 12. Partly exfoliated cranidium, holotype, NIGP 137376, x 6, P82.5. 13, 14. Partly exfoliated cranidium in dorsal and anterolateral views, NIGP 137377, x 6, W64.7. Figures 9, 11, 13, 14 from Dorypyge richthofeni Zone; Figures 10, 12 from Pianaspsis sinensis Zone.

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Plate 10 Figures 1-11. Fuchouia oratolimba Yang in Zhou et al., 1977 1. Small, testaceous cephalon with six thoracic segments attached, NIGP 137378, × 5, P269. 2. Small, incomplete, testaceous cranidium, NIGP 137379, × 4, P261. 3. Incomplete testaceous exoskeleton, latex cast, NIGP 137380, × 5, P301. 4. Testaceous cranidium, NIGP 137381, × 3, P261.35. 5. Incomplete, cranidium, latex cast, NIGP 137382, × 4, P269. 6. Incomplete, testaceous cranidium, NIGP 137383, × 4, P261. 7. Incomplete, testaceous thorax, NIGP 137384, × 2.25, P273.8. 8. Incomplete, testaceous cranidium, NIGP 137385, × 4, W 198.45. 9. Incomplete, possibly exfoliated cranidium, NIGP 137386, × 3, W 198.45. 10. Incomplete, testaceous exoskeleton, NIGP 137387, × 2.5, W 198.45. 11. Incomplete, testaceous pygidium, latex cast, NIGP 137388, × 3, P269. All from Wanshania wanshanensis Zone.

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Plate 11

Figures 1-5. Fuchouia oratolimba Yang in Zhou et al., 1977 1. Librigena, NIGP 137389, x 3, P279. 2. Testaceous cranidium, NIGP 137390, x 3, P301. 3. Testaceous pygidium with three thoracic attached, latex cast, NIGP 137391, x 3, P277. 4. Three cranidia, NIGP 137392, x 3, P273.52. 5. Testaceous exoskeleton with broken cranidium, NIGP 137393, × 2.25, P273.8. All from Wanshania wanshanensis Zone. Figures 6-14. Fuchouia bulba sp. nov.; unless otherwise stated, all from P131.7. 6. Testaceous meraspid cranidium, NIGP 137394, x 20. 7. Testaceous meraspid cranidium, NIGP 137395, x 20, P 130.5. 8. Testaceous meraspid cranidium, NIGP 137396, x 20, P130.5. 9. Testaceous cranidium, NIGP 137397, x 20. 10. Testaceous cranidium, NIGP 137398, x 15, P 130.5. 11. Testaceous cranidium, NIGP 137399, x 10, W82.1 12. Testaceous cranidium, NIGP 137400, x 10. 13. Testaceous cranidium, NIGP 137401, x 12, W98. 14. Testaceous cranidium, NIGP 137402, x 15, P 130.5 All from Pianaspsis sinensis Zone.

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Figures 1-12. Fuchouia bulba sp. nov.; unless otherwise stated, all from P131.7. 1. Testaceous meraspid cranidium, NIGP 137403, × 8. 2. Testaceous cranidium, NIGP 137404, × 6. 3. Testaceous cranidium, NIGP 137405, × 8. 4. Testaceous librigena, NIGP 137406, × 5. 5. Testaceous cranidium, NIGP 137407, x 6, W98. 6. Exfoliated cranidium, NIGP 137408, × 4, W98. 7, 8. Testaceous cranidium in lateral and dorsal views, holotype, NIGP 137409, × 3. 9. Partly exfoliated hypostome, NIGP 137410, × 5, W98. 10. Incomplete, testaceous librigena, NIGP 137411, × 3, W98. 11, 12. Testaceous exoskeleton, NIGP 137412, × 4, W99.3 (corrected from P99.3 in Peng et al., 2001, p. 102). All from Pianaspsis sinensis Zone.

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Plate 13 Figures 1-13. Fuchouia bulba sp. nov.; unless otherwise stated, all from P 131.7. 1, 2. Two testaceous cranidium in dorsal and lateral views, NIGP 137413, × 5. 3. Testaceous cranidium and hypostome, NIGP 137414, × 5.5. 4. Testaceous librigena, NIGP 137415, × 3, P 130.95. 5. Testaceous pygidium, NIGP 137416, × 10. 6. Testaceous pygidium, NIGP 137417, × 6. 7. Testaceous pygidium, NIGP 137418, × 4, W98. 8. Partly exfoliated librigena, NIGP 137419, × 5. 9. Testaceous pygidium, NIGP 137420, × 6. P131.5. 10. Testaceous pygidium, NIGP 137421, × 10. 11. Broken, partly exfoliated pygidium, NIGP 137422, × 6. 12. Partly exfoliated pygidium, NIGP 137423, × 5. 13. Partly exfoliated pygidium, NIGP 137424, × 7.5, P 130. All from Pianaspsis sinensis Zone.

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Plate 14 Figures 1-14. Dorypyge richthofeni Dames, 1883 1. Small testaceous cranidium, NIGP 137425, × 12, W56.7. 2. Small testaceous cranidium, NIGP 137426, × 10, W56.7. 3. Partly exfoliated cranidium, NIGP 137427, × 7, P82.5. 4. Small testaceous cranidium, NIGP 137428, × 6, P42.5. 5. Testaceous cranidium, NIGP 137429, × 4, P45.6. 6. Incomplete, testaceous cranidium, NIGP 137430, × 4, W56.7. 7, 8. Exfoliated cranidium in anterolateral and dorsal views, NIGP 137431, × 3, W56.7. 9, 10. Testaceous cranidium in lateral and dorsal views, NIGP 137432, × 4, P82.5. 11. Partly exfoliated hypostome, NIGP 137433, × 5, W56.7. 12, 13. Testaceous cranidium in lateral and dorsal views, NIGP 137434, × 4, P35.5. 14. Exfoliated, crushed cranidium, NIGP 137435, × 2, W56.7. All from Dorypyge richthofeni Zone.

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Figures 1-11. Dorypyge richthofeni Dames, 1883 1. Incomplete, partly exfoliated cranidium, NIGP 137436, × 2, W56.7. 2. Partly exfoliated hypostome, NIGP 137437, × 3, W82.2. 3. Small, testaceous pygidium, NIGP 137438, × 8, P3.2. 4. Small, testaceous pygidium, NIGP 137439, × 5, P42.5. 5. Small, testaceous pygidium, NIGP 137440, × 5, P45.6. 6. Partly exfoliated pygidium, NIGP 137441, × 4, P42.5. 7. Partly exfoliated pygidium, NIGP 137442, × 5, P42.5. 8. Partly exfoliated pygidium, NIGP 137443, × 4, P42.5. 9. Exfoliated pygidium, NIGP 137444, × 2, W56.7. 10. Testaceous pygidium, NIGP 137445, × 2, W56.7. 11. Exfoliated pygidium with partial thoracic segments, NIGP 137446, × 1.5, W56.7. All from Dorypyge richthofeni Zone.

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Figures 1-7a, 8-11. Dorypyge richthofeni Dames, 1883; unless otherwise stated, all from W82.1. 1. Nearly complete, testaceous pygidium, NIGP 137447, × 2.5, W56.7. 2. Testaceous cranidium, NIGP 137448, × 5, P42.5. 3, 4. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 1374498, × 4. 5. Testaceous pygidium, NIGP 137450, × 1.6. 6. Partly exfoliated pygidium, NIGP 137451, × 3. 7a. Partly exfoliated pygidium, NIGP 137452, in association with Amphoton sp. cf. A. typicum (Kobayashi, 1942a), × 3. 8. Testaceous, incomplete pygidium, NIGP 137454, × 3. 9, 10. Partly exfoliated hypostome in obliquely lateral and dorsal views, NIGP 137455, × 7.5. 11. Testaceous, incomplete pygidium, NIGP 137456, × 3. All from Dorypyge richthofeni Zone. Figure 7b. Amphoton sp. cf. A. typicum (Kobayashi, 1942a) 7b. mostly exfoliated pygidium, NIGP 137453, × 3. From Dorypyge richthofeni Zone.

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Plate 17 Figures 1-14. Dorypyge perconvexalis Yang in Zhou et al., 1977; unless otherwise stated, all from P277. 1. Small, incomplete, testaceous cranidium, NIGP 137457, × 5. 2, 4. Testaceous cranidium in dorsal and lateral views, NIGP 137458, × 3. 3. Incomplete, testaceous cranidium, NIGP 137459, × 2. 5. Incomplete testaceous librigena, latex cast, NIGP 137460, × 5. 6. Same librigena as in fig. 5, × 5. 7-9. Slightly exfoliated, broken cranidium in dorsal, anterior, and lateral views, NIGP 137461, x2. 10. Testaceous hypostome, NIGP 137462, × 3. 11. Testaceous pygidium, latex cast, NIGP 137463, × 1.5. 12. Testaceous pygidium, NIGP 137464, × 5, P273.8. 13. Incomplete testaceous pygidium, NIGP 137465, × 6, P273.8 14. Incomplete testaceous pygidium, NIGP 137466, × 5. All from Wanshania wanshanensis Zone.

• 240 •

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e

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Plate 18

Figures 1-6. Dorypyge bisulcata sp. nov. 1, 2. Nearly complete, testaceous cranidium in anterolateral and dorsal views, NIGP 137467, × 4, P156. 3. Nearly complete, testaceous pygidium, holotype, NIGP 137468, × 4, P 167. 4. Broken, testaceous pygidium, NIGP 137469, × 6, P 167. 5. Incomplete, testaceous pygidium, NIGP 137470, × 4.5, P130.95. 6. Broken pygidium, NIGP 137471, × 4, P 130.95. All from Pianaspsis sinensis Zone. Figures 7-9. Dorypyge globosa sp. nov. 7, 8. Partly exfoliated cranidium in dorsal and lateral views, holotype, NIGP 137472, × 4, P156. 9. Incomplete, testaceous pygidium, NIGP 137473, × 3.2, P 156. From Pianaspsis sinensis Zone.

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Plate 19

Figures 1-3. Dorypyge perconvexalis Yang in Zhou et al., 1977 1. Incomplete librigena, NIGP 137474, × 10, P374.9. 2. Fragmental cranidium, NIGP 137475, × 7, P273.8. 3. Incomplete testaceous pygidium, NIGP 133527, × 9, PI3-2.75. Figure 1 from Chuangia subquadrangulata Zone; Figure 2 from Wanshania wanshanensis Zone; Figure 3 from Liostracina bella Zone. Figure 4. Dorypyge? sp. 4. Testaceous pygidium, NIGP 137476, × 4.5, P290.5. From Wanshania wanshanensis Zone. Figure 5. Dorypyge richthofeni Dames, 1883 5. Incomplete, testaceous cranidium, NIGP 137477, × 2.25, P 108. From Dorypyge richthofeni Zone. Figures 6, 7. Dorypyge huaqiaoensis sp. nov. 6. Partly exfoliated cranidium and pygidium in association, NIGP 137478a, NIGP 137478b, × 4.5, P81. 7. Partly exfoliated cranidium, holotype, NIGP 137479, × 4.5, P81. From Dorypyge richthofeni Zone. Figures 8-10. Wanshania wanshanensis Rong and Yang in Zhou et al., 1977 8-10. Testaceous exoskeleton with librigenae missing, partial enlargement of thorax, and dorsal and anterolateral views, NIGP 137480, × 9, × 4.5, × 4.5, P273.8. From Wanshania wanshanensis Zone.

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Figures 1-14. Chuangia austriaca Yang in Zhou et al., 1977 1. Partly exfoliated cranidium, NIGP 137481, × 6, P1327.2. 2, 3. Incomplete, exfoliated cranidium in dorsal and anterolateral views, NIGP 137482, × 6, × 6, PI327.2. 4. Incomplete, mostly exfoliated cranidium, NIGP 137483, × 10, PI322.4. 5. Incomplete, mostly exfoliated cranidium, NIGP 137484, × 5, PI323.5. 6. Incomplete, partly exfoliated cranidium, NIGP 137485, × 5, PI322.4. 7, 8. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 137486, × 6, PI322.4. 9. Partly exfoliated librigena, NIGP 137487, × 3, PI322.4. 10. Mostly exfoliated pygidium, NIGP 137488, × 4, PI323.5. 11. Mostly exfoliated pygidium, NIGP 137489, × 8, P1323.5. 12, 14. Testaceous pygidium, partially enlargement and dorsal view, × 15, × 7.5, PI323.5. 13. Mostly exfoliated pygidium, NIGP 137490, × 5, PI323.5. All from Chuangia subquadrangulata Zone. Figures 15, 16. Chuangia subquadrangulata Sun, 1935 15. Testaceous cranidium, NIGP 137491, × 4.5, P370.6. 16. Testaceous pygidium, NIGP 137492, × 7.5, P374.9. From Chuangia subquadrangulata Zone.

• 246 •

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11 10 13

14

15

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Figures 1-11. Meropalla bella Yuan and Yin, 1998 1-5. Partly exfoliated cranidium in dorsal, anterolateral, posterodorsal, posterior, and lateral views, NIGP 137493, all x 15, W225. 6-9. Almost fully exfoliated cranidium in anterior, lateral, dorsal, and posterior views, NIGP 137494, all x 6, W228.3. 10. Exfoliated pygidium, NIGP 137495, x 8, P298.4. 11. Broken testaceous pygidium, NIGP 137496, x 5, P298.4. Figures 1-9 from the base of Liostracina bella Zone; Figures 10, 11 from the upper part of Wanshania wanshanensis Zone.

• 248.

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8

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Figures 1-6. Meropalla gibbera sp. nov. 1-6. Partly exfoliated cranidium in anterolateral, dorsal, anterolateral, oblique-posterior, oblique-lateral, and anterolateral views, holotype, NIGP 137497, × 8, W212.45. From the upper part of Wanshania wanshanensis Zone. Figure 7. Pagodiinae gen. et sp. indet. 7. Incomplete, testaceous cranidium, NIGP 137498, × 10, P 136.7. From Pianaspsis sinensis Zone. Figures 8, 9. Geminiclavula wangcunica gen. et sp. nov. 8, 9. Cranidium in dorsal and anterolateral views, holotype, NIGP 137499, × 10, W56.7. From Dorypyge richthofeni Zone.

• 250.

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8

9

Plate 23 Figures 1-14. Chiawangella hunanensis sp. nov. 1-5. Broken, testaceous cranidium in anterior, anterolateral, posterolateral, and dorsal views and partial enlargement, holotype, NIGP 137500, × 10, × 10, × 10, × 10, × 15, W227. 6, 7. Broken, testaceous pygidium in anterolateral and dorsal views. The anterolateral view shows the anterior and lateral borders and a small node on turning point of the borders, NIGP 137501, × 15, × 15, W227. 8-10. Broken, testaceous pygidium in dorsal, anterolateral and anterior views, latex cast, NIGP 137502, all × 12.5, W227. 11. Broken, testaceous pygidium, NIGP 133524, × 10, PI3-2.75. 12, 13. Broken, testaceous pygidium in dorsal and posterolateral views. Note the small node at the junction of anterior and posterior borders, NIGP 133525, x 5, x 5, P[3-2.75. 14. Broken, testaceous pygidium, NIGP 137503, x 15, W227. All from Liostracina bella Zone.

• 252 •

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12

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Figures 1-13. Wanshania wanshanensis Rong and Yang in Zhou et al., 1977 1. Small, incomplete, exfoliated cranidium, NIGP 137504, × 12, P240.5. 2. Nearly complete, testaceous cranidium, NIGP 137505, × 6, P277. 3. Nearly complete, testaceous cranidium, NIGP 137506, × 5, P277. 4. Nearly complete, testaceous cephalon, NIGP 137507, × 4, P325.7. 5. Two associated, partly exfoliated cranidia, NIGP 137508, NIGP 137509, × 3, P273.8. 6. Testaceous librigena, NIGP 137510, × 4, P273.8. 7. Testaceous cranidium, NIGP 137511, × 3, P315.8. 8. Exfoliated cranidium, latex cast, NIGP 137512, × 4, P261. 9. Incomplete, partly exfoliated cranidium, NIGP 137513, × 2.5, W207.5. 10. Slightly exfoliated pygidium, NIGP 137514, × 4.5, P264.2. 11. Testaceous pygidium, NIGP 137515, × 2.25, P273.8. 12. Partly exfoliated pygidium, NIGP 137516, × 3.5, P277. 13. Two associated, partly exfoliated pygidia, NIGP 137517a, NIGP 137517b, × 4, P307.4. Figures 1-3, 5, 6, 8-13 from Wanshania wanshanensis Zone; Figures 4, 7 from Liostracina bella Zone.

• 254-

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Plate 25

Figures 1-13. Prochuangia granulosa Lu, 1956; unless otherwise stated, all from PI372. 1. Testaceous cranidium, NIGP 137518, × 2.25, PI356.5. 2. Small, incomplete, testaceous pygidium, NIGP 137519, x 15, P378.25. 3. Nearly complete, testaceous cranidium, NIGP 137520, × 10. 4. Nearly complete, testaceous cranidium, NIGP 137521, × 10, PI360.3. 5. Incomplete, exfoliated librigena, NIGP 137522, × 10. 6. Nearly complete, partly exfoliated cranidium, NIGP 137523, × 5. 7. Nearly complete, slightly exfoliated pygidium, NIGP 137524, x 3.5. 8, 9. Testaceous cranidium in anterolateral and dorsal views, NIGP 137525, × 5. 10. Nearly complete, testaceous cranidium, NIGP 137526, × 10, PI322.4. 11, 12. Testaceous hypostome in dorsal and lateral views, NIGP 137527, × 6. 13. Nearly complete, slightly exfoliated pygidium, NIGP 137528, x 6. Figures 1, 2, 4, 10 from Chuangia subquadrangulata Zone; Figures 3, 5-9, 11-13 from base of Shengia quadrata Zone. Figures 14-18. Prochuangia sp. cf. P. leiocephala Peng, Geyer, and Hamdi, 1999 14. Small, exfoliated cranidium, NIGP 137529, × 12, P374.9. 15. Small, partly exfoliated cranidium, NIGP 137530, × 7.5, P375.15. 16. Small, partly exfoliated pygidium, NIGP 137531, × 15, P375.15. 17. Small, testaceous pygidium, NIGP 137532, × 7.5, P374.9. 18. Partly exfoliated pygidium, NIGP 137533, × 4, P375.15. All from Chuangia subquadrangulata Zone.

• 256.

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e

10

11

12

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15

16

17

18

Plate 26

Figures 1-6. Prochuangia sp. cf. P. leiocephala Peng, Geyer, and Hamdi, 1999 1. Nearly complete, partly exfoliated cranidium, NIGP 137534, x 5, P378.25. 2, 3. Nearly complete, partly exfoliated cranidium in anterolateral and dorsal views, NIGP 137535, x 5, P378.25. 4. Incomplete, partly exfoliated cranidium, NIGP 137536, x 3.7, P378.25. Note that a few granules are present on posterior fields of the fixigenae. 5. Incomplete, exfoliated pygidium, NIGP 137537, x 8.7, P374.9. 6. Incomplete, partly exfoliated cranidium, NIGP 137538, x 5, P378.25. All from Chuangia subquadrangulata Zone. Figures 7-11. Prochuangia linicispinata Peng, 1992 7. Small cranidium, NIGP 137539, x 13.2, P378.25. Note that a few granules are present on posterior fields of fixigenae. 8. Small testaceous pygiium, NIGP 137540, x 18, P378.25. 9. Incomplete testaceous cranidium, NIGP 133577, × 12, P13-1.6. 10. Testaceous pygidium, NIGP 137541, x 9, P378.25. 11. Partly exfoliated cranidium, NIGP 137542, x 12.5, P378.25. All from Chuangia subquadrangulata Zone.

• 258.

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8

10

11

Plate 27

Figures 1-12. Damesella hunanensis sp. nov. 1. Nearly complete testaceous cranidium, NIGP 137543, × 6, P277. 2. Nearly complete testaceous cranidium, NIGP 137544, × 6, P277. 3, 4. Testaceous cranidium in dorsal and anterolateral vievs, NIGP 137545, × 6, P277. 5. Nearly complete testaceous cranidium, holotype, NIGP 137546, × 6, P277. 6. Fragmental testaceous cranidium, NIGP 137547, × 7.5, P294.5. 7. Incomplete, exfoliated pygidium, NIGP 137548, × 5, W211.7. 8. Partly exfoliated pygidium, NIGP 137549, × 5, W211.7. 9, 10. Broken, testaceous pygidium, and latex cast from its external mold, NIGP 137550, × 4, P298.4. 11. Partial enlargement of fig. 9 to show fine, densely spaced granules and coarse scattered granules on pygidial surface, × 12.5. 12. Testaceous, malformed pygidium having one more medial spine than other pygidia of the species, NIGP 137551, × 3, P294.5. All from Wanshania wanshanensis Zone.

• 260.

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10

11

12

9

Plate 28

Figures 1-7. Blackwelderia youshuica sp. nov. 1, 2. Testaceous cranidium in dorsal and anterior views, NIGP 137552, × 11, W132.5. 3-5. Broken, testaceous cranidium in dorsal, anterolateral and dorsal views, holotype. Fig. 5 is a latex cast from the external mold of figs. 3, 4. NIGP 137553, × 3, W132.5. 6. Composite photo of fragmental, testaceous cranidium with the fight part of palpebral area and the posterolateral projection from a latex cast, NIGP 137554, × 3, W132.5. 7. Partial enlargement of fig. 6, × 9. All from Pianaspsis sinensis Zone. Figures 8-10. Paradamesella nobilis Lu and Lin, 1989 8. Incomplete, testaceous pygidium, NIGP 137555, × 12, P249. 9. Nearly complete, testaceous pygidium, NIGP 137556, × 9, P249. 10. Incomplete, testaceous pygidium, NIGP 137557, × 9.5, P249. All from the base of Wanshania wanshanensis Zone.

• 262 •

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9

10

Plate 29

Figures 1-13. Parablackwelderiajimaensis (Yang in Lu et al., 1974a) 1. Incomplete, testaceous librigena, showing anastomosing caecal ridges, NIGP 137558, x 3, W196.3. 2. Incomplete, testaceous cranidium, NIGP 137559, x 4, W 199.2. 3. Small testaceous cranidium, NIGP 137560, x 5, P199.9. 4. Testaceous cranidium, NIGP 137561, x 3, W188.8. 5, 6. Testaceous cranidium with posterior portion of glabella damaged, dorsal and anterolateral views, NIGP 137562, x 6, W187.8. 7, 8. Incomplete testaceous cranidium, dorsal view and partial enlargement of glabella and occipital ring to show the granular ornamentation, NIGP 137563, x 3, x 6, W187.8. 9. Incomplete, testaceous pygidium, NIGP 137564, x 3, W187.8. 10. Partly exfoliated pygidium, NIGP 137565, x 5, W211.7. 11, 12. Testaceous pygidium, latex cast, dorsal view and partial enlargement of left pleural field, NIGP 137566, x 3, x 5, P269. 13. Testaceous pygidium, latex cast, NIGP 137567, x 3, W188.8. Figures 1, 2, 4-13 from Wanshania wanshanensis Zone; Figure 3 from upper part of Pianaspsis sinensis Zone.

• 264 •

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11

12

13

Plate 30

Figures 1-15. Parablackwelderiajimaensis (Yang in Lu et al., 1974a) 1. Meraspid cranidium, latex cast, NIGP 137568, × 15, P277. 2. Meraspid cranidium, NIGP 137569, × 15, P277. 3. Incomplete, testaceous cranidium, NIGP 137570, × 3, P260. 4. Partially enlargement of fig. 6, showing granules on posterior portion and occipital ring, × 5, P277. 5. Testaceous librigena, NIGP 137571, × 4, P277. 6. Testaceous cranidium, NIGP 137572, × 3, P277. 7. Testaceous librigena, NIGP 137573, × 3, P277. 8. Incomplete, testaceous librigena, NIGP 137574, × 3, P277. 9. Incomplete, testaceous cranidium, NIGP 137575, × 4, P261.35. 10. Testaceous pygidium, NIGP 137576, × 4, P277. 11. Hypostoma, NIGP 137577, × 2.25, P298.54. 12. Incomplete, testaceous pygidium, NIGP 137578, × 4, P275.1. 13. Incomplete, testaceous pygidium, showing anastomosing caecal ridges on pleural field, NIGP 137579, × 2, P277. 14. Incomplete, testaceous pygidium, showing anastomosing caecal ridges on pleural field, NIGP 137580, × 5, P273.8. 15. Incomplete, testaceous pygidium, NIGP 137581, × 2, P277. All from Wanshania wanshanensis Zone.

• 266-

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10 8

12

11

14

15

Plate 31

Figures 1-5. Parablackwelderia laterilobata (Yang in Zhou et al., 1977) 1, 2. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 137582, × 4, P240.5. 3. Testaceous pygidium, latex cast, NIGP 137583, × 5, P200.7. 4. Small, testaceous cranidium, NIGP 137584, × 5, P171.5. 5. Incomplete, testaceous cranidium, NIGP 137585, × 4, W 132.5. From Pianaspsis sinensis Zone. Figures 6, 7. Parablackwelderia sp. 6. Testaceous pygidium with axis damaged, NIGP 137586, x 3, P 156. 7. Incomplete, testaceous cranidium, NIGP 137587, x 4, P 156. From lower part of Pianaspsis sinensis Zone. Figures 8-12. Parablackwelderia sp. cf. P. huabeiensis (Zhang in Qiu et al., 1983) 8. Partly damaged and partially exfoliated cranidium, NIGP 137588, × 3, P184. 9, 10. Incomplete, partially exfoliated pygidim, dorsal view and enlargement of fight posterolateral part, NIGP 137589, × 3, × 6, P 190.7. 11, 12. Incomplete, partially exfoliated cranidium in dorsal and anterolateral views, NIGP 137590, × 2.5, P184. From upper part of Pianaspsis sinensis Zone.

• 268•

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10

11

12

Plate 32

Figures 1-7. Protaitzehoia granifera Yang in Yin and Li, 1978 1. Incomplete, partly exfoliated cranidium, NIGP 137591, × 5, P346.7. 2. Incomplete, partly exfoliated cranidium, NIGP 137592, × 5, P346.7. 3, 4. Incomplete, partly exfoliated cranidium, NIGP 137593, × 3.5, P348. Fig. 4 is a composite photograph of fig. 3 with left part being a mirror image of fight side. 5. Incomplete, testaceous pygidium with injured border on left side, NIGP 137594, × 5, P348. 6, 7. Testaceous pygidium in dorsal and posterior views, NIGP 137595, × 5, P307.4. Figures 1-5 from Liostracina bella Zone; Figures 6, 7 from the top part of Wanshania wanshanensis Zone. Figures 8-13. Protaitzehoia yuepingensis Yang in Yin and Li, 1978 8, 9. Incomplete, testaceous cranidium, and partial enlargement of glabella showing fine granules, NIGP 137596, × 7, × 18, W 197.5. 10. Incomplete, mostly exfoliated cranidium, NIGP 137597, × 7, W196.3. 11, 12. Partly exfoliated pygidium and partial enlargement of left pleural field showing fine granules, NIGP 137598, x 10, × 20, W 197.5. 13. Testaceous pygidium, NIGP 137599, × 4, P337.5. Figures 8-12 from upper part of Wanshania wanshanensis Zone; Figure 13 from lower part of Liostracina bella Zone. Figure 14. Protaitzehoia sp. 14. Incomplete pygidium, latex cast, NIGP 133529, × 15, PI3-2.75. From Liostracina bella Zone.

• 270.

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10

11

13

12

14

Plate 33

Figures 1-11. Protaitzehoia subquadrata Peng, 1987 1. Testaceous juvenile cranidium, NIGP 137600, × 15, P277. 2. Incomplete, testaceous juvenile cranidium, NIGP 137601, × 15, P277. 3. Incomplete, testaceous juvenile cranidium, NIGP 137602, × 12, P277. 4. Incomplete, testaceous juvenile cranidium, NIGP 137603, × 12, P298.4. 5. Incomplete, testaceous juvenile cranidium, NIGP 137604, × 10, P331.8. 6. Testaceous juvenile cranidium, NIGP 137605, x 10, P277. 7. Testaceous cranidium, NIGP 137606, × 8, W 199.2. 8, 9. Testaceous cranidium in anterolateral and dorsal views, NIGP 137607, × 6, W 199.2. 10. Testaceous librigena, NIGP 137608, × 7.5, P319.8. 11. Broken, partly exfoliated, cephalon, × 3.5, NIGP 137609, × 8, P325.7. Figures 1-4 from Wanshania wanshanensis Zone; Figures 5, 10, 11 from Liostracina bella Zone.

• 272.

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11

Plate 34

Figures 1-7. Protaitzehoia subquadrata Peng, 1987 1. Testaceous juvenile pygidium, NIGP 137610, x 8, P325.7. 2. Testaceous pygidium, NIGP 137611, × 7, W200.7. 3. Broken testaceous pygidium, NIGP 137612, × 5, P326.9. 4. Testaceous pygidium, NIGP 137613, × 6, P317.4. 5-7. Testaceous pygidium in posterior and dorsal views, and partial enlargement showing fine and coarse granules and wrinkles on axis and pleural field, NIGP 137614, × 5.5, × 5.5, × 15, P319.8. Figures 1, 3-7 from Liostracina bella Zone; Figure 2 from upper part of Wanshania wanshanensis Zone. Figures 8-17. Protaitzehoia spinifera sp. nov. 8. Testaceous juvenile cranidium, NIGP 137615, × 6, P341.8. 9. Incomplete, testaceous cranidium, NIGP 137616, × 7, W200.3. 10. Incomplete, testaceous cranidium, NIGP 137617, × 8, W221.5. 11, 12. Incomplete, testaceous cranidium, and partial enlargement, NIGP 137618, × 5, × 10 W225. 13. Testaceous juvenile pygidium, NIGP 137619, × 20, P337.5. 14. Exfoliated juvenile pygidium, NIGP 137629, × 10, W211.7. 15-17. Testaceous pygidium in dorsal and posterolateral views and partial enlargement, holotype, NIGP 137621, × 5.5, × 5.5, × 11, W221.5. Figures 8, 10-13, 15-17 from Liostracina bella Zone; Figures 9, 14 from upper part of Wanshania wanshanensis Zone.

• 274•

Plate 34

.

r

.

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10 d

11

12 13 15

14

16

17

Plate 35

Figures 1-12. Teinistion posterocostum (Yang in Zhou et al., 1977) 1. Incomplete, testaceous juvenile cranidium, NIGP 137622, × 15, W225. 2, 3. Incomplete, testaceous juvenile cranidium in dorsal and anterolateral views, NIGP 137623, ×12, W221.5. 4. Broken cranidium, NIGP 137624, × 8, P319.6. 5. Incomplete, testaceous cranidium, NIGP 133521, × 15, PI3-2.75. 6. Incomplete, partly exfoliated cranidium, latex cast, NIGP 137625, × 7, W227. 7. Nearly complete, testaceous cranidium, NIGP 137626, × 12, P319.6. 8. Nearly complete, testaceous cranidium, NIGP 137627, × 8, P317.4. 9. Nearly complete, testaceous cranidium, NIGP 137628, x 10, P317.4. 10-12. Nearly complete, slightly exfoliated cranidium in anterior, dorsal, and anterolateral views, NIGP 137629, × 5.5, W227. All from Liostracina bella Zone.

• 276.

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11

12

Plate 36

Figures 1-12. Teinistion posterocostum (Yang in Zhou et al., 1977) 1. Incomplete, possibly exfoliated cranidium, NIGP 133520, × 10, PI3-2.75. 2. Broken, testaceous cranidium, NIGP 137630, × 10, W218.3. 3. Hypostome, NIGP 137631, × 6, P317.4. 4. Incomplete, testaceous juvenile pygidium, NIGP 137632, × 12, W215.1. 5. Nearly complete, testaceous pygidium, NIGP 137633, × 9, W227. 6. Nearly complete, testaceous pygidium, NIGP 137634, × 8, W225. 7. Testaceous juvenile pygidium, NIGP 133522, × 15, PI3-2.75. 8. Nearly complete, testaceous juvenile pygidium, NIGP 133523, × 15, PI3-2.75. 9. Testaceous pygidium, NIGP 137635, × 10, P319.8. 10. Broken, partly exfoliated pygidium, NIGP 137636, × 9, W219.7. 11. Slightly exfoliated pygidium, NIGP 137637, × 8, W254.1. 12. Pygidium in ventral view, showing doublure, NIGP 137638, × 7, W228.3. All from Liostracina bella Zone.

• 278-

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11

12

Plate 37

Figures 1-6. Teinistion posterocostum (Yang in Zhou et al., 1977) 1. Testaceous pygidium, NIGP 137639, × 8, W225. 2. Testaceous pygidium, NIGP 137640, × 5.2, W221.5. 3. Testaceous pygidium, NIGP 137641, × 5, W225. 4. Testaceous juvenile pygidium, NIGP 137642, × 6, W254.1. 5. Testaceous pygidium, NIGP 137643, x 5, W225. 6. Testaceous pygidium, NIGP 137644, × 4.5, P319.6. All from Liostracina bella Zone. Figures 7-15. Taihangshania wangcunensis sp. nov. 7. Incomplete, testaceous juvenile cranidium, NIGP 137645, × 15, W251.15. 8. Partial, testaceous juvenile cranidium, NIGP 137646, × 15, W225. 9, 15. Incomplete, testaceous cranidium and partial enlargement of posterior portion showing granulate ornamentation and the course of posterior branch of facial suture. Arrow indicates the anterior end of the posterior branch, holotype, NIGP 137647, × 5, × 12, P319.8 (corrected, previously listed as W227 in Peng, Babcock, and Lin, 2001 b). 10. Testaceous, partial cranidium, NIGP 137648, × 7, W227. 11. Incomplete, testaceous cranidium, NIGP 137649, × 5, W227. 12, 13. Testaceous, partial cranidium in dorsal and anterior views, showing elevated palpebral lobe and steeply inclined palpebral area of fixigena, NIGP 137650, × 5, W227. 14. Incomplete, slightly exfoliated pygidium, NIGP 137651, × 5, W227. All from Liostracina bella Zone.

• 280.

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13

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11

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Plate 38

Figures 1-6. Taihangshania wangcunensis sp. nov. 1, 2. Broken slightly exfoliated pygidium, dorsal view and partial enlargement, NIGP 137652, × 3, × 6, W227. 3. Composite photograph from fig. 1 with fight part reversed from the left part, x 4.5. 4. Partial enlargement of left posterior border of cranidium (PI. 37, fig. 11), showing tooth-like spines, × 10. 5, 6. Partly exfoliated pygidium, posterolateral and dorsal views, CUGB 0324302, × 6, Laochatian, Fenghuang, western Hunan (originally a paratype of Bergeronites austriacus Yang, 1978, pl. 11, figs. 4a, b). Figures 1-4 from Liostracina bella Zone. Figures 7-13. Palaeadotes hunanensis (Yang in Zhou et al., 1977) 7. Incomplete, slightly exfoliated cranidium, latex cast, NIGP 137653, × 5, W 193.6. 8. Nearly complete, testaceous cranidium, NIGP 137654, × 5, W 187.8. 9, 10. Nearly complete, slightly exfoliated cranidium in anterolateral and dorsal views, NIGP 137655, × 3, W193.6. 11. Nearly complete, testaceous pygidium, NIGP 137656, × 2, W 196.3. 12. Nearly complete, slightly exfoliated pygidium, NIGP 1376577, × 2.5, P287.1. 13. Nearly complete, slightly exfoliated pygidium, latex cast, NIGP 137658, x 3, W212.45. All from Wanshania wanshanensis Zone.

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10 11

12

13

Plate 39

Figures 1-13. Palaeadotes hunanensis (Yang in Zhou et al., 1977) 1. Testaceous librigena, NIGP 137659, × 11, W 187.8. 2. Nearly complete, slightly exfoliated pygidium, NIGP 137660, × 3, W 198.45. 3. Partly exfoliated hypostome, NIGP 137661, × 3, P298.54. 4. Slightly exfoliated hypostome, NIGP 137662, × 5, W227. 5. Slightly exfoliated hypostome, NIGP 137663, × 4, P298.4. 6. Testaceous pygidium, NIGP 137664, × 2.5, P287.1. 7. Nearly complete, testaceous pygidium, NIGP 137665, × 2.5, P287.1. 8. Testaceous pygidium, NIGP 137666, × 1.5, P348. 9. Partial, testaceous pygidium, NIGP 137667, × 2.5, W 199.2. 10. Testaceous pygidium in ventral view, showing doublure, NIGP 137668, × 4, P341.8. 11. Nearly complete, slightly exfoliated pygidium with partial exoskeleton, NIGP 137669, × 1.5, P348. 12, 13. Nearly complete, testaceous pygidium and partial enlargement to show flange on left anterior margin, NIGP 137670, × 3, × 9, W221.5. Figures 1-3, 5-7, 9, 10 from Wanshania wanshanensis Zone; Figures 4, 8, 11-13 from Liostracina bella Zone.

• 284.

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13

10

11

12

Plate 40 Figures 1-5. Palaeadotes hunanensis (Yang in Zhou et al., 1977) 1-4. Complete, testaceous exoskeleton in dorsal, lateral, dorsal, and anterolateral views, × 6, × 5, × 5, × 6, NIGP 137671, × 11, P308. 5. Nearly complete, testaceous pygidium, latex cast, NIGP 137672, × 3, W 111.3. Figures 1-4 from upper part of Wanshania wanshanensis Zone; Figure 5 from upper part of Pianaspsis sinensis Zone.

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Plate 41

Figures 1-11. Palaeadotes bella (Qiu in Qiu et al., 1983) 1. Incomplete, testaceous cranidium, NIGP 137673, × 3, W215.1. 2. Incomplete, slightly exfoliated cranidium, NIGP 137674, × 2, W215.1. 3. Partial, slightly exfoliated cranidium, NIGP 137675, × 3, W219.7. 4, 5. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 137676, × 4, W216.5. 6, 7. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 137677, × 3, W219.7. 8. Pygidial doublure, NIGP 137678, × 6, W225. 9. Slightly exfoliated pygidium, NIGP 137679, × 3, W219.7. 10, 11. Incomplete, partly exfoliated pygidium, and enlarged posterior border, NIGP 137680, × 3, × 4.5, W221.5. Figures 3-11 from base of Liostracina bella Zone; Figures 1, 2 from upper part of Wanshania wanshanensis Zone.

• 288.

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11

Plate 42

Figures 1-10. Paradamesella typica Yang in Zhou et al., 1977 1. Small juvenile cranidium, NIGP 137681, × 16, W216.5. 2. Small, incomplete, juvenile cranidium, NIGP 137682, × 16, W216.5. 3. Small, incomplete juvenile cranidium, NIGP 137683, × 15, W227. 4. Small, juvenile, testaceous cranidium, latex cast, NIGP 137684, × 11, W254.1. 5. Incomplete testaceous juvenile cranidium and associated hypostome, NIGP 137685, × 6, W216.5. 6. Incomplete juvenile cranidium, NIGP 137686, × 6, W242.6. 7. Incomplete, testaceous cranidium, associated with Proagnostus bulbus Butts, 1926, × 4, W219.7. 8, 9. Testaceous cranidium in dorsal and anterolateral views, NIGP 137687, × 5, W254.1. 10. Incomplete, testaceous pygidium, NIGP 137688, × 6, W219.7. Figures 1, 2, 5 from upper part of Wanshania wanshanensis Zone; Figures 3, 4, 6-10 from Liostracina bella Zone.

• 290 •

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9

10

Plate 43

Figures 1-7. Paradamesella typica Yang in Zhou et al., 1977 1. Testaceous cranidium, latex cast, NIGP 137689, × 4.5, P325.7 2. Testaceous hypostome, NIGP 137690, × 6, W216.5. 3. Testaceous pygidium, NIGP 133526, × 5, PI3-2.75. 4. Testaceous pygidium, NIGP 137691, × 2, P348. 5. Testaceous pygidium, NIGP 137692, × 6, W216.5. 6. Incomplete, testaceous pygidium, NIGP 137693, × 4, W251.15. 7. Incomplete, testaceous pygidium, NIGP 137694, × 4, W254.1. Figures 1, 3, 4, 6, 7 from Liostracina bella Zone; Figures 2, 5 from upper part of Wanshania wanshanensis Zone. Figure 8. Parademesella peculiaris Zhou in Zhou et al., 1977 8. Incomplete, testaceous pygidium, NIGP 137695, × 3, P277. From Wanshania wanshanensis Zone.

• 292-

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7

8

Plate 44

Figures 1-14. Parademesella peculiaris Zhou in Zhou et al., 1977 1. Small juvenile cranidium, latex cast, NIGP 137696, x 10, W 199.2. 2. Incomplete, testaceous cranidium, NIGP 137697, x 6, W 198.45. 3. Broken, testaceous cranidium, NIGP 137698, x 5, P331.8. 4. Testaceous librigena, note that the posterior branch of the facial suture curves inward after acrossing posterior border furrow, NIGP 137699, x 8, W 188.8. 5, 6. Incomplete testaceous cranidium in dorsal and anterolateral views, NIGP 137700, x 4, x 4, P331.8. 7. Incomplete, testaceous librigena, showing the anterior branch of the facial suture, NIGP 137701, x 6, W 198.45. 8. Testaceous hypostoma, NIGP 137702, x 8, W 198.45. 9. Broken, testaceous pygidium, NIGP 137703, x 6, W 187.8. 10. Incomplete, testaceous pygidium, NIGP 137704, x 5, P216. 11. Incomplete, testaceous hypostoma, NIGP 137705, x 4, P298.4. 12. Broken, testaceous pygidium, NIGP 137706, x 6, P282.6. 13. Incomplete, testaceous pygidium, NIGP 137707, x 6, P216. 14. Broken, testaceous pygidium, NIGP 137708, x 3, W 189. Figures 1, 2, 4, 7-9, 11, 12, 14 from Wanshania wanshanensis Zone; Figures 3, 5, 6 from Liostracina bella Zone; Figures 10, 13 from upper part of Pianaspis sinensis Zone.

• 294.

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Figure 1. Gypsum cast of rock slab containing syntypes of Drepanura premesnili Bergeron, 1899, Blackwelderia sinensis (Bergeron, 1899), and Shantungia sinensis (Bergeron, 1899), and holotype of Pseudagnostus douvillei Bergeron, 1899; original of Bergeron, 1899, pl. 13, which is said to to be from north of Beijing but is probably from Dawenkou, Shandong, China. This plastotype is housed in the Department of Earth Sciences, Universit6 Claude Bernard, Lyon, France. Natural size (scale bar = 2 cm). A, a-a". Drepanura premesnili Bergeron, 1899. A, lectotype pygidium, a. pygidia in dorsal view; a', pygidia in ventral view; a", broken cranidia. B, b-b". Blackwelderia sinensis (Bergeron, 1899). B. cranidium, lectotype, selected herein; b, librigena; b', pygidia in dorsal view; b", pygidia in ventral view. C, c. Pseudagnostus douvillei (Bergeron, 1899). C, broken cephalon?, holotype in dorsal view; c, cephalon, possibly belonging to the species. D. Shantungia sinensis (Bergeron, 1899) [=Shantungia spinidera Walcott, 1905]. D. Incomplete pygidium. Figure 2. Blackwelderia? sp. 2. In complete, testaceous pygidium, dorsal view, NIGP 137709, × 5, P269. From lower part of Wanshania wanshanensis Zone. Figures 3, 4. Palaeadotes hunanensis (Yang in Zhou et al., 1977) 3, 4. Incomplete, slightly exfoliated cranidium, and its composite photograph with left side being a mirror image of the fight side, NIGP 137710, × 2, x 1.4, W219.7. From Liostracina bella Zone. Figures 5-7. Drepanura? crassispina sp. nov. 5, 6. Slightly exfoliated pygidium NIGP 13771 l, x 4, W219.7 7. Slightly exfoliated pygidium NIGP 137712, × 10, holotype, W218.3. From Liostracina bella Zone.

• 296 •

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Figures 1-16. Paraacidaspis hunanica Egorova in Poletaeva, 1960 1. Small, nearly complete, testaceous cranidium, NIGP 137713, × 10, PI35.1. 2. Testaceous cranidium, NIGP 137714, × 8, P378.25. 3. Mostly exfoliated cranidium, NIGP 137715, × 6, P[327.2. 4. Broken, testaceous cephalon, NIGP 137716, × 4.5, P316.0. 5. Crushed, testaceous cephalon, NIGP 137717, × 2, P378.25. 6. Testaceous librigena, NIGP 137718, × 9, P331.8. 7. Testaceous pygidium, NIGP 137719, × 4.5, PI356.5. 8. Testaceous pygidium, latex cast, NIGP 137720, × 2, P331.8. 9. Partly exfoliated pygidium, latex cast, showing broad doublure with wrinkled surface, NIGP 137721, × 2.5, PI327.2. ? 10. Partly exfoliated pygidium, NIGP 137722, × 7, P261.35. 11. Broken, testaceous pygidium, showing broad doublure with terrace lines, NIGP 137723, × 3, P319.6. 12. Incomplete, testaceous pygidium, latex cast, NIGP 137724, × 3, PI372. 13. Incomplete, testaceous librigena, showing acutely pointed genal angle, NIGP 137725, × 4, PB56.5. 14. Broken, testaceous pygidium, NIGP 137726, × 6, PI360.3. 15. Partial enlargement of fig. 2, showing wrinkle ornamentation, × 12. 16. Partial enlargement of fig. 11, showing wrinkle ornamentation, × 6. Figures 1-3, 5, 7, 9, 13-15 from Chuangia subquadrangulata Zone; Figure 12 from base of Shengia quadrata Zone; Figures 4, 6, 8, 11, 16 from Liostracina bella Zone; Figure 10 from Wanshania wanshanensis Zone. Figure 17. Paraacidaspis sp. 17. Incomplete, testaceous pygidium, latex cast, NIGP 137727, × 7, PI360.3. From Chuangia subquadrangulata Zone.

• 298•

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Figures 1-15. Afghanocare truncatum (Peng, 1987) 1, 2. Incomplete, mostly exfoliated cranidium in dorsal and oblique anterior views, NIGP 137728, × 10, W227. 3-5. Incomplete, slightly exfoliated cranidium in anterolateral, oblique anterior, and dorsal views, NIGP 137729, × 6, W227. 6. Incomplete, mostly exfoliated cranidium, NIGP 137730, × 5, W227. 7. Broken, slightly exfoliated cranidium, NIGP 137731, × 4.5, P378.25. 8-10. Complete, mostly exfoliated pygidium in dorsal, posterolateral, and posterior views, NIGP 137732, × 2, W227. 11. Incomplete, partly exfoliated pygidium, NIGP 137733, × 4, P374.9. 12-14. Nearly complete, testaceous pygidium in posterior, dorsal, and posterolateral views, NIGP 137734, × 4, P319.6. 15. Incomplete, slightly exfoliated pygidium, NIGP 137735, × 4.5, P378.25. Figures 1-6, 8-10, 12-14 from Liostracina bella Zone; Figures 7, 11, 15 from Chuangia subquadrangulata Zone.

• 300.

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13

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Plate 48

Figures 1, 2. Paracoosia sp. cf. P. kingi Witteke, 1984 1, 2. Small testaceous pygidium in dorsal and posterior views, NIGP 137736, x 8, P261.35. From Wanshania wanshanensis Zone. Figures 3-16. Paracoosia huayuanensis sp. nov. 3. Small, incomplete, partly exfoliated cranidium, NIGP 137737, × 4, P319.6, 4, 5. Small, fragmental, testaceous cranidium in dorsal and oblique-lateral views, NIGP 137738, × 3, P319.8. 6. Small incomplete, testaceous cranidium, NIGP 137739, × 3, P337.5. 7, 8. Testaceous cranidium in dorsal and oblique-anterior views, NIGP 137740, × 3, P317.4. 9. Small, partly exfoliated pygidium, NIGP 137741, × 7, P319.6. 10. Fragmental, partly exfoliated cranidium, NIGP 137742, × 2, W227. 11. Incomplete, partly exfoliated cranidium, holotype, NIGP 137743, × 2, P319.8. 12-15. Incomplete, broken pygidium showing part of doublure, in dorsal (12) and posterior (13) views, × 2; 14 is the enlarged anterior part of axis of the pygidium, × 4; 15 is the enlarged anterolateral comer of the pygidium, × 4, NIGP 137744, P319.8. 16. Incomplete, mostly exfoliated pygidium, NIGP 137745, × 2, P319.8. All from Liostracina bella Zone.

• 302.

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Figures 1-14. Paibianomocare paibiense gen. et sp. nov. 1. Incomplete, slightly exfoliated cranidium, NIGP 137746, × 10, P260. 2. Incomplete, partly exfoliated cranidium, NIGP 137747, × 7.5, PI34.3. 3. Fragmental, testaceous cranidium, NIGP 137748, × 10, W227. 4. Incomplete, testaceous cranidium, NIGP 137749, × 8, P376.4. 5. Incomplete, testaceous cranidium, NIGP 137750, × 8, P277. 6. Incomplete, testaceous cranidium, NIGP 137751, × 6, P277. 7, 10. Nearly complete, testaceous cranidium in dorsal and lateral views, NIGP 137752, × 6, P277. 8. Incomplete, slightly exfoliated cranidium, latex cast, NIGP 137753, × 5, P269. 9. Incomplete, testaceous cranidium, holotype, NIGP 137754, × 6, P277. 11. Incomplete, testaceous cranidium, NIGP 137755, × 6, P277. 12. Incomplete, testaceous pygidium, latex cast, NIGP 137756, × 5, P269. 13. Incomplete, testaceous pygidium, latex cast, NIGP 137757, × 6, P277. 14. Incomplete, testaceous pygidium, latex cast, NIGP 137758, × 7.5, P273.8. Figures 1, 5-14 from Wanshania wanshanensis Zone; Figures 2, 3 from Liostracina bella Zone, Figure 4 from Chuangia subquadrangulata Zone.

• 304.

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Figures 1-15. Paracoosia sp. cf. P. kingi Witteke, 1984 1. Small incomplete, exfoliated cranidium, NIGP 137759, × 10, P261.35. 2. Small incomplete, partly exfoliated cranidium, NIGP 137760, × 5, P261.35. 3. Fragmental, exfoliated cranidium, NIGP 137761, × 4.5, P295.13. 4. Incomplete, testaceous cranidium, NIGP 137762, × 2, P260. 5. Complete, slightly exfoliated librigena, NIGP 137763, × 1.5, P261.35. 6. Incomplete, slightly exfoliated cranidium, NIGP 137764, × 1.5, P261.35. 7. Incomplete, testaceous cranidium, NIGP 137765, × 4, W 195.7. 8, 9. Hypostome in posterolateral and dorsal views, NIGP 137766, × 7, P261.35. 10. Incomplete, partly exfoliated cranidium, NIGP 137767, × 1.5, P261.35. 11. Fragmental, testaceous pygidium, NIGP 137768, × 3, W 196.3. 12, 13. Slightly broken, slightly exfoliated pygidium in posterior and dorsal views, latex cast, NIGP 137769, × 1.5, P261.35. 14. Nearly complete, slightly exfoliated pygidium, latex cast, NIGP 137770, × 1.5, W 196.3. 15. Broken, slightly exfoliated pygidium, NIGP 137771, × 2, W212.45. All from Wanshania wanshanensis Zone.

• 306 •

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Figures 1-4. Metopotropis sinensis sp. nov. 1, 2. Nearly complete, partly exfoliated cranidium in dorsal and anterolateral views, holotype, NIGP 137772, × 10, P363.5. 3, 4. Fragmentary, testaceous cranidium in dorsal and anterolateral views, NIGP 137773, × 10, P319.8. From Liostracina bella Zone. Figure 5. Paibianomocare lineatum gen. et sp. nov. 5. Incomplete, testaceous cranidium, holotype, NIGP 137774, × 10, PI35.1. From Chuangia subquadrangulata Zone. Figures 6-10. Yangweizhouia carinata Yuan and Yin, 1998 6, 7. Incomplete, slightly exfoliated cranidium, NIGP 137775, × 9, P301.9. 8-10. Nearly complete, testaceous cranidium in anterior, anterolateral, and dorsal views, NIGP 137776, × 11, W227. Figures 6, 7 from upper part of Wanshania wanshanensis Zone; Figures 8-10 from base of Liostracina bella Zone. Figures 11, 12. Glyphaspellus? sinensis sp. nov. 11,12. Incomplete, slightly exfoliated cranidium (a) in association with Prodamesella punctata Ergaliev, 1980 (b), dorsal and anterolateral views, holotype, NIGP 137777, × 12, P277. From base of Liostracina bella Zone. Figures 13-15. Afghanocare truncatum (Peng, 1987) 13-15. Nearly complete, testaceous cranidium in dorsal, anterior, and anterolateral views, NIGP 137778, × 7.5, P319.6. From base of Liostracina bella Zone.

• 308.

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Plate 52

Figures 1-14. Proceratopyge (Proceratopyge)fuyangensis Lu and Lin in Peng, 1987 1. Small, incomplete, testaceous cranidium, NIGP 137779, × 12, P319.6. 2. Incomplete, testaceous cranidium, NIGP 137780, × 12, W216.5. 3. Incomplete, testaceous cranidium, NIGP 137781, × 8, W210.5. 4. Incomplete, slightly crushed, testaceous cranidium, NIGP 137782, × 12, P319.6. 5. Incomplete testaceous cranidium, NIGP 137783, × 8, P298.4. 6. Incomplete testaceous cranidium, NIGP 137784, × 8, W216.5. 7. Incomplete, partly exfoliated cranidium, NIGP 137785, × 8, W210.5. 8. Incomplete, partly exfoliated cranidium, NIGP 137786, × 8, W228.3 9. Nearly complete, slightly exfoliated cranidium, NIGP 137787, × 7, W210.5. 10. Crushed, testaceous pygidium with four thoracic segments attached, NIGP 137788, × 8, P317.4. 11. Incomplete, slightly exfoliated pygidium, NIGP 137789, × 10, P317.4. 12. Pygidium with spines broken, NIGP 137790, × 7.5, P317.4. 13. Testaceous pygidium, NIGP 137791, × 8, P317.4. 14. Incomplete, mostly exfoliated pygidium, NIGP 137792, × 12, W227. Figures 1, 4, 8, 10-14 from base of Liostracina bella Zone; Figures 2, 35-79 from upper Wanshania wanshanensis Zone. Figure 15. Proceratopyge sp. indet. 15. Incomplete cranidium, NIGP 137793, × 6, P319.6. From base of Liostracina bella Zone.

• 310.

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Plate 53

Figures 1-4. Proceratopyge (Proceratopyge)fuyangensis Lu and Lin in Peng, 1987 1. Incomplete, testaceous cranidium, NIGP 137794, × 5.5, W228.3. 2. Broken, testaceous cranidium, NIGP 137795, × 7, P298.4. 3, 4. Incomplete, testaceous exoskeleton in anterolateral and dorsal views, NIGP 137796, × 6, P327.4. Figures 1, 3, 4 from Liostracina bella Zone; Figure 2 from upper Wanshania wanshanensis Zone. Figures 5, 9. Proceratopyge (Proceratopyge)fenghwangensis Hsiang in Egorova et al., 1963 5. Incomplete, mostly exfoliated cranidium, NIGP 137797, × 7.5, P317.4. 9. Librigena, NIGP 137798, × 8, P317.4. From base of Liostracina bella Zone. Figures 6-8. Proceratopyge (Proceratopyge) truncata Yang in Zhou et al., 1977 6. Incomplete, partly exfoliated cranidium, NIGP 137799, × 7.5, PI360.15. 7. Slightly exfoliated cranidium, NIGP 137800, × 4, P374.9. 8. Nearly complete, slightly exfoliated cranidium, NIGP 137801, × 4, P378.25. All from Chuangia subquadrangulata Zone.

• 312.

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Figures 1-4. Proceratopyge (Proceratopyge) fenghwangensis Hsiang in Egorova et al., 1963 1. Nearly complete, testaceous pygidium, NIGP 137802, × 5, P374.9. 2. Testaceous pygidium, NIGP 137803, × 4, P376.4. 3. Testaceous pygidium, NIGP 133563, × 6, PI3-1.72. 4. Mostly exfoliated, broken pygidium showing doublure, NIGP 137804, × 7, P1372. Figures 1, 2 from Chuangia subquadrangulata Zone; Figure 3 from Liostracina bella Zone; Figure 4 from the base of Shengia quadrata Zone. Figures 5-10. Proceratopyge (Proceratopyge) truncata Yang in Zhou et al., 1977 5. Pygidium with nine thoracic segments attached, NIGP 137805, × 4, PI3 65.8. 6. Nearly complete, slightly exfoliated exoskeleton, latex cast, NIGP 137806, ×4, PI3 27.2. 7. Two broken, partly exfoliated pygidia, NIGP 137807, × 3, PI3 60.15. 8, 9. Incomplete, mostly exfoliated librigena in lateral and dorsal views, NIGP 137808, × 5, P378.25. 10. Mostly exfoliated librigena in lateral and dorsal views, NIGP 137809, × 5, P378.25. All from Chuangia subquadrangulata Zone.

• 314.

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Plate 55

Figures 1-18. Pseudoyuepingia laochatianensis Yang in Zhou et al., 1977 1. Small testaceous cranidium, NIGP 137810, × 12, W 196.3. 2. Small testaceous cranidium, NIGP 137811, × 12, W 199.2, 3. Small, testaceous cranidium, NIGP 137812, × 10, W 199.2. 4. Small, incomplete, testaceous cranidium, NIGP 137813, × 20, P353.7. 5. Small, nearly complete, testaceous cranidium, NIGP 137814, × 8, P353.7. 6. Testaceous cranidium, NIGP 137815, × 10, P353.64. 7. Nearly complete, testaceous cranidium, NIGP 137816, × 2, P353.7. 8. Fragmental, slightly exfoliated cranidium, NIGP 137817, × 10, W 199.2. 9. Small, testaceous pygidium, NIGP 137818, × 20, P353.7. 10, 15b. Small, testaceous pygidium, NIGP 137819, × 21, P353.7. 11. Small, testaceous pygidium, NIGP 137820, × 9, W 199.2. 12. Small, testaceous pygidium, NIGP 137821, × 10, W199.2. 13. Incomplete, testaceous pygidium, NIGP 137822, × 8, P353.7. 14. Nearly complete, mostly exfoliated pygidium, NIGP 137823, × 7, W254.1. 15a. Partly exfoliated pygidium, NIGP 137824, × 6, P353.7. 16. Partly exfoliated pygidium, NIGP 137825, × 4, P344.6. 17. Partly exfoliated pygidium, NIGP 137826, × 4, P353.7. 18. Slightly exfoliated pygidium, NIGP 137827, × 4.5, P361.5. Figures 1-3, 8, 11, 12 from upper part of Wanshania wanshanensis Zone; Figures 4-7, 9, 10, 13-18 from Liostracina bella Zone.

• 316.

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Plate 56

Figures 1, 2. Ajrikina wannanensis (Qiu in Qiu et al., 1983) 1. Incomplete, slightly exfoliated cranidium, latex cast, NIGP 137828, × 8, W 197.5. 2. Broken, slightly exfoliated cranidium, NIGP 137829, × 11, W 187.8. From Wanshania wanshanensis Zone. Figures 3-17. Ajrikina hunanensis (Peng, 1987) 3. Incomplete, testaceous cranidium, NIGP 137830, × 15, W216.5. 4, 5, 11. Incomplete, testaceous cranidium in dorsal and anterolateral views, and partial enlargement, NIGP 137831, × 9, × 9, × 13.5, W210.5. 6. Broken, incomplete, testaceous cranidium, NIGP 137832, × 8, W216.5. 7-10. Slightly exfoliated cranidium, partially enlarged, and in dorsal, oblique-anterior, and anterolateral views, NIGP 137833, × 13, × 10, × 10, × 10, W216.5. 12-14. Nearly complete, slightly exfoliated cranidium in dorsal, oblique-anterior, and anterolateral views, NIGP 137834, × 10, × 8, × 10, P277. 15. Small, nearly complete, testaceous pygidium, NIGP 137835, × 25, W216.5. 16. Nearly complete, testaceous pygidium, NIGP 137836, × 20, W216.5. 17. Incomplete, testaceous pygidium, NIGP 137837, × 20, W216.5. All from Wanshania wanshanensis Zone.

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Figures 1-5. Ajrikina hunanensis (Peng, 1987) 1-3. Incomplete, testaceous cranidium in dorsal, anterolateral, and oblique-anterior views, NIGP 137838; note that anterior portion of anterior border is folded beneath the posterior portion, x 10, W216.5. 4. Nearly complete, testaceous cranidium, latex cast, showing a node on the summit of L1 and an occipital spine, NIGP 137839, x 10, W216.5. 5. Partly exfoliated, incomplete cranidium, showing left palpebral lobe, NIGP 137840, × 12, W219.76.5. From upper Wanshania wanshanensis Zone. Figures 6, 7. Parablackwelderia jimaensis (Yang in Lu et al., 1974a) 6, 7. Nearly complete, partly exfoliated hypostome in dorsal and lateral views NIGP 137841, × 4, P341.8. From Liostracina bella Zone. Figure 8. Damesella? sp. 8. Fragmental pygidium, dorsal view, NIGP 137842, x 3, P331.8. From Liostracina bella Zone. Figures 9-13. Palaeadotes hunanensis (Yang in Zhou et al., 1977) 9. Incomplete, testaceous pygidium, NIGP 137843, × 16, PI3-2.75. 10-12. Hypostome in posterolateral, anterolateral, and dorsal views, NIGP 137844, × 4.2, P331.9. 13. Nearly complete, testaceous hypostome, latex cast, NIGP 137845, × 8, P341.8. From Liostracina bella Zone.

• 320 •

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Figures 1-11. Torifera taoyuanensis (Peng, 1987). 1. Small, testaceous cranidium, NIGP 137846, × 12, W 199.2. 2, 3. Incomplete, slightly exfoliated cranidium, in anterolateral and dorsal views, NIGP 137847, × 8, W199.2. 4. Incomplete testaceous cranidium, NIGP 137848, × 12, W216.5. 5, 6. Incomplete, partly exfoliated cranidium in dorsal and anterolateral views, NIGP 137849, × 12, W196.3. 7. Nearly complete, partly exfoliated cranidium, NIGP 137850, × 10, W211.7. 8, 9. Nearly complete, partly exfoliated cranidium in anterolateral and dorsal views, NIGP 137851, × 10, × 10, W211.7. 10. Testaceous pygidium, NIGP 137852, × 20, W199.2. 11. Testaceous pygidium, NIGP 137853, × 20, W210.5. All from Wanshania wanshanensis Zone.

• 322 •

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Figures 1-15. Torifera tuma (Yang in Zhou et al., 1977) 1. Small, testaceous cranidium, NIGP 137854, × 20, P317.4. 2. Incomplete, testaceous cephalon, showing linear anterior and fight lateral border, and fight facial suture, NIGP 137855, × 15, P268.3. 3-9. Nearly complete, testaceous cephalon, NIGP 137856. 3-5, latex cast, in fight anterolateral, dorsal, and left anterolateral views, all × 12; 6-9, in fight anterolateral, dorsal, oblique anterior, and anterior views, × 12, × 15, × 15, × 12, W 187.8. 10. Incomplete, testaceous cephalon, showing linear anterior and lateral borders on fight side, fight facial suture, and terrace lines on posterior part of librigena, NIGP 137857, × 12, P261. 11. Incomplete, testaceous cranidium, NIGP 137858, × 15, P261. 12. Nearly complete cephalon with librigena slightly displaced, NIGP 137859, × 10, W211.7. 13, 14. Fragment of partly exfoliated cranidium in dorsal and anterolateral views, showing linear anterior border, NIGP 137860, x 12, × 12, W 173.8. 15. Slightly exfoliated cranidium, latex cast, NIGP 137861, × 12, W171.9. Figure 1 from the basal part of Liostracina bella Zone; Figures 2-15 from Wanshania wanshanensis Zone.

• 324.

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Plate 60

Figures 1-10. Torifera abrupta sp. nov. 1. Small, testaceous, incomplete cranidium, NIGP 137862, × 12, W227. 2. Nearly complete, mostly exfoliated cranidium, NIGP 137863, × 11, W227. 3. Incomplete, testaceous cranidium in anterior view, NIGP 137864, × 10, W227. 4, 5. Nearly complete, testaceous cranidium in dorsal and anterior views, holotype, NIGP 137865, × 12, W227. 6. Slightly exfoliated cranidium, NIGP 137866, × 12, W227. 7, 8. Exfoliated cranidium in dorsal and anterolateral views, NIGP 137867, × 12, W227. 9. Incomplete, testaceous pygidium, NIGP 137868, × 20, W227. 10. Testaceous pygidium, NIGP74580, × 15, WT8b, Huaqiao Formation, Wa'ergang, Taoyuan (originally assigned to Liostracina taoyuanensis by Peng, 1987, pl. 12, fig. 9). From Liostracina bella Zone. Figure 11. Torifera? paraconvexa (Yang in Yin and Li, 1978) 11. Incomplete, testaceous cranidium, CUGBP 1004001, x 12, holotype, Huaqiao Formation, Liangweizhou, Yuping, eastern Guizhou (holotype of Cyclolorenzella paraconvexa Yang, 1978, pl. 7, fig. 7).

• 326 •

Plate 60

10

11

Plate 61

Figures 1-14. Liostracina bella Lin and Zhou, 1983 1. Nearly complete, slightly exfoliated cranidium, NIGP 137869, × 12.5, W227. 2. Nearly complete, testaceous cranidium, NIGP 137870, × 12, W225. 3. Incomplete, testaceous cranidium, NIGP 137871, × 13, W227. 4. Nearly complete, testaceous cranidium, NIGP 137872, × 13, W254.1. 5, 6. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 137873, both × 12, W218.3. 7. Incomplete, partly exfoliated cranidium, NIGP 137874, × 10, W254.1. 8, 9. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 137875, both × 9, W227. 10. Incomplete, testaceous librigena, NIGP 137876, × 12, W218.3. 11, 12. Nearly complete, testaceous cranidium in anterolateral and dorsal views, NIGP 137877, × 11, W227. 13, 14. Complete, testaceous librigena in dorsal and anterior views, NIGP 137878, × 13, W227. All from Liostracina bella Zone.

• 328.

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11

/

13

12

14

Plate 62

Figures 1-13. Liostracina bella Lin and Zhou, 1983 1. Incomplete, testaceous cranidium, NIGP 137879, x 10, W225. 2. Incomplete, exfoliated cranidium, NIGP 137880, x 10, P317.4. 3. Enlargement of the cranidium in P1.61, figs. 8, 9, x 16, W227. 4. Nearly complete, testaceous cranidium, NIGP 137881, x 10, W225. 5. Nearly complete, testaceous cranidium, NIGP 137882, 9, W227. 6, 7. Nearly complete librigena in anterior and dorsal views, NIGP 137883, x 12, W227. 8. Incomplete, testaceous cranidium, NIGP 137884, x 10, W218.3. 9. Incomplete, slightly exfoliated librigena, NIGP 137885, x 8, W254.1. 10. Incomplete pygidium, latex cast, NIGP 137886, x 17.5, P346.7. 11. Incomplete pygidium, latex cast, NIGP 137887, x 14, W227. 12. Incomplete, partly exfoliated cranidium, NIGP 137888, x 8, W227. 13. Complete, testaceous librigena in ventral view, NIGP 137889, x 10, W225. All from Liostracina bella Zone.

• 330 •

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10

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13

Plate 63 Figures 1-13. Monkaspis quadrata Yang in Zhou et al., 1977 1. Small incomplete, slightly exfoliated cranidium, NIGP 137890, × 6, P282.6. 2. Small incomplete, testaceous cranidium, NIGP 137891, × 8, P298.4. 3. Small incomplete, testaceous cranidium, NIGP 137892, × 8, W218.3. 4. Small incomplete, testaceous cranidium, NIGP 137893, × 7, W212.45. 5. Nearly complete, partly exfoliated cranidium, NIGP 137894, × 6, W210.5. 6. Incomplete, slightly exfoliated cranidium, NIGP 137895, × 4.5, P298.54. 7, 8. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 137896, × 2, P319.6. 9. Small, testaceous pygidium, NIGP 137897, × 12, W210.5. 10. Nearly complete, testaceous pygidium, NIGP 137898, × 4.5, P298.54. 11. Incomplete, testaceous pygidium, NIGP 137899, × 10, W211.7. 12, 13. Incomplete, testaceous pygidium in dorsal and posterolateral views, NIGP 137900, × 5, P298.4. All from Wanshania wanshanensis Zone.

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POLYMERID TRILOBITES FROM THE CAMBRIAN OF NORTHWESTERN HUNAN, CHINA Volume 2

Ptychopariida, Eodiscida, and Undetermined Forms

PENGShanchi

Loren E. BABCOCK LIN Huanling

~

SciencePress Beijing

Responsible Editresses: HU Xiaochun and LIN Caihua

POLYMERID TRILOBITES HUNAN, CHINA

Volume 2

FROM THE CAMBRIAN

Ptychopariida, Eodiscida, and Undetermined Forms

PENG Shanchi, Loren E. BABCOCK, and LIN Huanling

Copyright © 2004 by Science Press Published by Science Press http://www, sciencep, corn 16 Donghuangchenggen North Street Beijing 100717 P. R. China

Printed in Beijing, 2004 Reprinted in Beijing, 2006 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. ISBN 7-03-014905-X/Q • 1538

OF NORTHWESTERN

LIST OF AUTHORS PENG Shanchi Nanjing Institute of Geology and Palaeontology Chinese Academy of Sciences 39 East Beijing Road Nanjing 210008, China

Loren E. BABCOCK Department of Geological Sciences The Ohio State University 275 Mendenhall Laboratory 125 South Oval Mall Columbus, Ohio 43210, USA

LIN Huanling Nanjing Institute of Geology and Palaeontology Chinese Academy of Sciences 39 East Beijing Road Nanjing 210008, China

This research was funded by National Natural Science Foundation of China (49070077, 40072003, 40023002, 40332018) Chinese Academy of Sciences (KZCX2-SW-129) Ministry of Technology and Science of China (2001 DEB20056, G2000077700) National Geographic Society (No. 5819-96; 7151-0 l) State Key Laboratory of Paleobiology and Stratigraphy (No. 933114) State Education Commission of China The Ohio State University US National Science Foundation (EAR 9405990, 9526709, 0073089, 0106883, EAR OPP-0346829)

CONTENTS I N T R O D U C T I O N .............................................................................................................................. 1 A C K N O W L E D G M E N T S ........................................................................................................... 1 R E P O S I T O R I E S .......................................................................................................................... 1 S Y S T E M A T I C P A L E O N T O L O G Y ................................................................................................ 7 Order PTYCHOPARIIDA Swinnerton, 1915 .................................................................................... 7 Suborder PTYCHOPARIiNA Richter, 1933 ..................................................................................... 7 Superfamily ~YCHOPARIOtDAE Matthew, 1887 ......................................................................... 7 ?Family PTYCHOPARIIDAE Matthew, 1887 .................................................................................. 7 Genus TOWNLEYELLA Opik, ! 967 ............................................................................................... 7

Townleyella rata Yuan and Yin, 1998 ................................................................................ 7 Genus X~L~GXL~ Lu in Z h a n g et al., 1980 .................................................................................. 8

Xilingxia ichangensis (Chang, 1964) ................................................................................... 9 Ptychopariid gen. et sp. indet.. ........................................................................................... 10 Supeffamily AS~HISCOIDEA R a y m o n d , 1924 .......................................................................... 11 F a m i l y ANOMOCARELLIDAE Hup~, 1953 ................................................................................... 11 Genus FISSANOMOCARELLA Peng, Lin, and Chen, 1995 ........................................................... 11

Fissanomocarella paibiensis Peng, Lin, and Chen, 1995 .................................................. 11 Genus HUAYUAMA Peng, Lin, and Chen, 1995 ......................................................................... ! ! Huayuania subcalva Peng, Lin, and Chen, ! 995 ............................................................... Huayuania quadrilateralis (Resser and Endo, 1937) ........................................................ Genus PARANOMOCARELLA Yang in Z h o u et al., 1977 ............................................................. Paranomocarella parallela Yang in Zhou et al., 1977 ..................................................... Paranomocarella fortis Peng, Lin, and Chen, 1995 .......................................................... Paranomocarella similaris sp. nov.. .................................................................................. Paranomocarella sp.. ......................................................................................................... Paranomocarella? obvia sp. nov.- .....................................................................................

12 12 14 15

17 18 19

19 Genus SZEASPIS Chang, 1959 .................................................................................................... 20

Szeaspis huananensis sp. n o v . . .......................................................................................... 21 Family PROASAPHISCIDAE Chang, 1963 .................................................................................... 22 Genus PROASAPHISCUS Resser and E n d o in Kobayashi, 1935 .................................................. 22

Proasaphiscus? sp. cf. P. butes (Walcott, 1905) ............................................................... 23 Subgenus PROASAPHISCUS (HONANASPIS) Chang, 1959 ........................................................... 23

Proasaphiscus (Honanaspis)? sp. cf. P. (H.) parvus Kuo in Lu et al., 1965 .................... 24 Genus ADELOGONUS Opik, 1967 ............................................................................................... 25 Adelogonus hunanensis sp. n o v . . ....................................................................................... 26 Adelogonus? sp.. ................................................................................................................ 27 Genus EYMEKOPS Resser and Endo, 1937 ................................................................................. 28

Eymekops? sp. 1 ................................................................................................................. 28 Eymekops? sp. 2 ................................................................................................................. 29 Genus GRANDIOCULUS C o s s m a n n , 1908 ................................................................................... 29 °i

°

Grandioculus obscurus sp. nov.- ........................................................................................ 31 Grandioculus truncatus sp. n o v . . ....................................................................................... 32 G e n u s HUAYUANASPIS gen. n o v . . .............................................................................................. 33

Huayuanaspis granulosa gen. et sp. n o v . - ......................................................................... 34 Huayuanaspis perpauca gen. et sp. n o v . - .......................................................................... 35 Huayuanaspis? sp. •............................................................................................................ 36 G e n u s INIOTOMA O p i k , 1967 ..................................................................................................... 37

Iniotoma laevis sp. n o v . . .................................................................................................... 37 Iniotoma porosus sp. n o v . . ................................................................................................. 38 G e n u s MAOTUNIA Z h a n g a n d Jell, 1987 .................................................................................... 39

Maotunia Maotunia Maotunia Maotunia

M. blackwelderi ( R e s s e r a n d E n d o , 1937) .............................................. 39 M. semiplectra Z h a n g a n d Jell, 1987 ...................................................... 41 rectangularis sp. n o v . . ....................................................................................... 41 sp. cf.

sp. cf.

s p . . ..................................................................................................................... 43

in Jell a n d R o b i s o n , 1978 ......................................................... 43 Sudanomocarina traynorae Z h a n g a n d Jell, 1987 ............................................................. 4 4 Sudanomocarina sp. cf. S. changi Jell in Jell a n d R o b i s o n , 1978 ..................................... 4 4 Sudanomocarina? huananensis sp. nov.- ........................................................................... 45 Sudanomocarina sp.. .......................................................................................................... 47 G e n u s ZHUJIA Ju in Q i u et al., 1983 .......................................................................................... 47 Zhujia hunanensis P e n g , Lin, a n d C h e n , 1995 .................................................................. 48 G e n u s SUDANOMOCARINA Jell

G e n u s PAIBIELLA gen. nov.- ....................................................................................................... 48

Paibiella paibiensis gen. et sp. n o v . . ................................................................................. 49 G e n u s PARAYUJINIA gen. n o v . . .................................................................................................. 50

Parayujinia constricta gen. et sp. n o v . . ............................................................................. 50 F a m i l y CATILLICEPHALIDAE R a y m o n d , 1938 ............................................................................ 52 S u b f a m i l y CATILLICEPHALINAE R a y m o n d , 1938 ...................................................................... 52 G e n u s BUTI'SIA W i l s o n , 1951 .................................................................................................... 52

Buttsia globosa L u a n d L i n in P e n g , 1987 ......................................................................... 53 G e n u s MADAROCEPHALUS R e s s e r , 1 9 3 8 a .................................................................................. 55

Madarocephalus orientalis sp. nov.. .................................................................................. in Z h a n g et al., 1 9 8 0 a .................................................................. Paradistazeris hubeiensis Z h u in Z h a n g et al., 1 9 8 0 a ....................................................... Paradistazeris hunanensis ( P e n g , 1987) ............................................................................ Paradistazeris rotundus sp. n o v . . ...................................................................................... Paradistazeris sp.. ..............................................................................................................

G e n u s PARADISTAZERIS Z h u

56 57 58 58 59 60

S u b f a m i l y ONCHONOTININAE Lu, 1965 ..................................................................................... 61 G e n u s HUZHUIA Chu, 1965 ........................................................................................................ 62

Huzhuia paratypica Y a n g , 1978 ........................................................................................ Huzhuia curvata sp. n o v . . .................................................................................................. Huzhuia latilimbata sp. n o v . . ............................................................................................. G e n u s ONCHONOTELLUS L e r m o n t o v a in I v s h i n , 1956 .............................................................. Onchonotellus curvitensus sp. n o v . . ..................................................................................

63 64 65 66 67

G e n u s PLACOSEMA O p i k , 1967 .................................................................................................. 68

Placosema bigranulosum sp. n o v . . .................................................................................... 68 F a m i l y CREPICEPHALIDAE K o b a y a s h i , 1935 .............................................................................. 69 G e n u s CREPICEPHALINA R e s s e r a n d E n d o

in K o b a y a s h i , 1935 ................................................. 69

Crepicephalina pergranosa R e s s e r a n d E n d o , 1937 ......................................................... 69 Crepicephalina sp. cf. C. quadrata R e s s e r a n d E n d o , 1937 .............................................. 70 • ii°

Crepicephalina? sp.. .......................................................................................................... 71 Genus METEORASPIS Resser, 1935 ............................................................................................. 71

Meteoraspis sp. cf. M. orientalis Yuan and Yin, 1998 ...................................................... 72 Family DICERATOCEPHALIDAE Lu, 1954 ................................................................................... 73 Genus FENGHUANGELLA Y a n g in Z h o u et al., 1977 .................................................................. 73 Fenghuangella laochatianensis laochatianensis Yang in Z h o u et al., 1977 ..................... 74

Fenghuangella laochatianensis crassa subsp, nov.. .......................................................... 76 Fenghuangella coniforma Yang in Zhou et al., 1977 ........................................................ 77 Fenghuangella liostracinala Yang in Zhou et al., 1977 .................................................... 79 Fenghuangella fusilis sp. n o v . . .......................................................................................... 80 Family DOKIMOCEPHALIDAE Kobayashi, 1935 ......................................................................... 81 Subfamily CONOKEPHALININAE Hup6, 1955 ............................................................................. 81 Genus LOBOCEPHALINA Rfi~i6ka, 1940 ..................................................................................... 81

Lobocephalina sinensis sp. n o v . . ....................................................................................... 82 Family EULOMIDAE Kobayashi, 1955 ........................................................................................ 83 Subfamily EULOMINAE Kobayashi, 1955 ................................................................................... 83 Genus STIGMATOA Opik, 1963 .................................................................................................. 83

Stigmatoa yangziensis Yang in Z h o u et al., 1977 .............................................................. 84 Family KINGSTONIIDAE Kobayashi, 1933b ................................................................................ 85 Genus KINGSTONIA Walcott, 1924 ............................................................................................. 85

Kingstonia euryaxis sp. nov.. ............................................................................................. 85 Family LISANIIDAE Chang, 1963 ............................................................................................... 87 Genus LISANIA Walcott, 1911 .................................................................................................... 87

Lisania paratungjenensis Yang in Zhou et al., 1977 ......................................................... 90 Lisania yuanjiangensis (Yang in Z h o u et al., 1977) .......................................................... 92 Lisania sp. cf. L. agonius (Walcott, 1905) ......................................................................... 94 Lisania paibiensis sp. nov.- ................................................................................................ 95 Lisania wangcunensis sp. nov.. .......................................................................................... 97 Lisania? sp.- ....................................................................................................................... 97 Genus BAOJINGIA Y a n g in Z h o u et al., 1977 ............................................................................. 98 Baojingia youshuiensis Yang in Zhou et al., 1977 ............................................................ 99 Baojingia jiudiantangensis (Yang in Zhou et al., 1977) ................................................. 101 Baojingia latilimbata (Peng, 1987) .................................................................................. 102 Baojingia paralala (Yang in Z h o u et al., 1977) .............................................................. 103 Baojingia quadrata (Yang in Z h o u et al., 1977) ............................................................. 104 Baojingia subquadrata (Yang, 1978) .............................................................................. 106 Baojingia tungjenensis (Nan in E g o r o v a et al., 1963) ..................................................... 108 Genus NEOANOMOCARELLA Hsiang in E g o r o v a et al., 1963 .................................................. 109 Neoanomocarella asiatica Hsiang in E g o r o v a et al., 1963 ............................................. 1 1 0 Neoanomocarella incilis sp. nov.. .................................................................................... 112 Genus QIANDONGASPIS Yuan and Yin, 1998 ........................................................................... 113

Qiandongaspis Qiandongaspis Qiandongaspis Qiandongaspis

sinensis (Peng, 1987) .............................................................................. 114 convexa sp. nov.. ..................................................................................... 115 xiangxiensis sp. nov.. .............................................................................. 116 sp.. ........................................................................................................... 117

Genus SHENGIA Hsiang in E g o r o v a et al., 1963 ...................................................................... 1 1 7

Shengia quadrata Hsiang in E g o r o v a et al., 1963 ........................................................... 1 1 8 Shengia trapezia Peng, 1992 ............................................................................................ 119 Shengia wannanensis Qiu in Qiu et al., 1983 .................................................................. 120 • iii.

Shengia

sp.. ................................. .....................................................................................

121

F a m i l y LONCHOCEPHALIDAE HupG, 1953 ................................................................................

121

G e n u s AETHIA Qian and Z h o u , 1984 .......................................................................................

121

Aethia rectangula

Q i a n and Z h o u , 1984 ..........................................................................

122

G e n u s NEOGLAPHYRASPIS Y u a n and Yin, 1998 ...................................................................... 122

Neoglaphyraspis nitida

Y u a n and Yin, 1998 ................................................................... 123

G e n u s PRODAMESELLA C h a n g , 1959 .......................................................................................

Prodamesella punctata E r g a l i e v , 1980 ............................................................................ Prodamesella sp. cf. P. biserrata Jell in Jell and R o b i s o n , 1978 .................................... Prodamesella tumidula sp. nov.. ...................................................................................... F a m i l y MAPANIIDAE C h a n g , 1963 ........................................................................................... G e n u s MAPANIA R e s s e r and E n d o

Mapania jiangnanensis

in K o b a y a s h i ,

124 125 126 127 128

1935 ........................................................... 128

sp. nov. •.....................................................................................

129

F a m i l y ONCHONOTOPSIDAE Shaw, 1966 .................................................................................

130

G e n u s LUYANHAOASPIS P e n g , B a b c o c k , and Lin, 2001 b ........................................................ 130

Luyanhaoaspis decorosa (Peng, Lin, and C h e n , 1995) ................................................... Luyanhaoaspis inflata sp. n o v . . ....................................................................................... Luyanhaoaspis sp. cf. L. inflata sp. n o v . . ........................................................................

131 132 132

G e n u s HUAYUANELLA gen. n o v . . .............................................................................................

133

Huayuanella paibiensis gen. et sp. nov. •......................................................................... Huayuanella zhiqiangi gen. et sp. nov.. ...........................................................................

134

F a m i l y PAPYRIASPIDIDAE W h i t e h o u s e , 1939 ..........................................................................

135

G e n u s PIANASPIS Saito and S a k a k u r a , 1936 ............................................................................

135

Pianaspis attenuata ( L e r m o n t o v a and C h e r n y s h e v a in C h e r n y s h e v a , 1950) .................. Pianaspis sinensis ( Y a n g in Z h o u et al., 1977) ............................................................... G e n u s WANGCUNIA P e n g , Lin, and C h e n , 1995 ......................................................................

134

137 139 140

Wangcunia wangcunensis Peng, Lin, and Chen, 1995 .................................................... 140 G e n u s PARAPIANASPIS gen. nov.- ............................................................................................. 141 Parapianaspis hunanensis

gen. et sp. n o v . . ..................................................................... 142

F a m i l y RHYSSOMETOr'IDAE Opik, 1967 ...................................................................................

143

G e n u s RHYSSOMETOPUS Opik, 1967 .......................................................................................

143

Rhyssometopus zhongguoensis Z h o u in Z h o u et al.,

1977 .............................................. 143

F a m i l y SOLENOPLEURIDAE A n g e l i n , 1854 ...............................................................................

144

in Qiu et al., 1983 .............................................................. Changqingia intermedia ( W a l c o t t , 1906) ........................................................................ Changqingia laevis P e n g , Lin, and C h e n , 1995 ...............................................................

144

G e n u s MENOCEPHALITES K o b a y a s h i , 1935 .............................................................................

145

G e n u s CHANGQINGIA L u and Z h u

Menocephalites sp. cf. M. pauperata ( W a l c o t t , 1906) .................................................... Menocephalites hunanensis sp. nov.. ............................................................................... Menocephalites? sp. 1 ...................................................................................................... Menocephalites? sp. 2 ...................................................................................................... F a m i l y SHUMARDIIDAE L a k e , 1907 .........................................................................................

144 145 146 147 148 148 149

G e n u s OCULISHUMARDIA P e n g , B a b c o c k , H u g h e s , and Lin, 2003 ......................................... 149

Oculishumardia hunania

Peng, B a b c o c k , H u g h e s , and Lin, 2003 .................................. 149

G e n u s LIMBISHuMARDIA gen. nov.. .........................................................................................

Limbishumardia wangcunensis

PTYCHOPARIOIDAE FAMILY UNCERTAIN ..................................................................................

152

G e n u s PSEUDOMAPANIA Y u a n and Yin, 1998 .........................................................................

152

Pseudomapania cylindrica •

iv

°

150

gen. et sp. nov. •............................................................. 151

Y u a n and Yin, 1998 ............................................................. 152

S u b o r d e r OLENINA B u r m e i s t e r , 1843 ....................................................................................... 153 S u p e r f a m i l y OLENOIDEA B u r m e i s t e r , 1843 ............................................................................. 153 F a m i l y OLENIDAE B u r m e i s t e r , 1843 ........................................................................................ 153 S u b f a m i l y O L E N I N A E B u r m e i s t e r , 1843 ................................................................................... 153 G e n u s OLENUS D a l m a n , 1827 .................................................................................................. 153

Olenus austriacus Y a n g in Z h o u et al., 1977 .................................................................. 153 Olenus punctatus sp. n o v . . ............................................................................................... 155 S u b f a m i l y PELTURINAE C o r d a , 1847 ....................................................................................... 156 G e n u s SULCARECLAVA gen. nov.. ............................................................................................ 156

Sulcareclava sagitta gen. et sp. nov.. ............................................................................... 157 S u b o r d e r HARPINA W h i t t i n g t o n , 1959 ..................................................................................... 158 ? F a m i l y HARPIDIDAE W h i t t i n g t o n , 1950 ................................................................................. 158 G e n u s B A I K A D A M A S P I S E r g a l i e v , 1980 ................................................................................... 158

Baikadamaspis sinensis ( Y a n g in Z h o u et al., 1977) ....................................................... 159 Baikadamaspis sp. cf. B. granulosa ( Y u a n a n d Yin, 1998) ............................................. 161 Baikadamaspis linearis sp. n o v . - ..................................................................................... 161 Baikadamaspis paibiensis sp. n o v . . ................................................................................. 163 Baikadamaspis sp.. ........................................................................................................... 165 TRILOBITA ORDER UNCERTAIN .................................................................................................

165

F a m i l y BURLINGIIDAE W a l c o t t , 1908 ...................................................................................... 165 G e n u s S C H M A L E N S E E I A M o b e r g , 1903 .................................................................................... 165

Schmalenseeia sinensis Y a n g in Y i n and Li, 1978 .......................................................... 167 Schmalenseeia fusilis sp. nov.- ......................................................................................... 170 UNDETERMI]~ JED T R I L O B I T E S C L E R I T E S . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

171

F a m i l y UNDETERMINED ........................................................................................................... 171 G e n u s U n d e t e r m i n e d ............................................................................................................... 171 U n d e t e r m i n e d larva ......................................................................................................... 171 U n d e t e r m i n e d p y g i d i u m 1 ............................................................................................... 172 U n d e t e r m i n e d p y g i d i u m 2 ............................................................................................... 172 U n d e t e r m i n e d l i b r i g e n a 1 ................................................................................................ 173 U n d e t e r m i n e d l i b r i g e n a 2 ................................................................................................ 173 U n d e t e r m i n e d l i b r i g e n a 3 ................................................................................................ 173 U n d e t e r m i n e d l i b r i g e n a 4 ................................................................................................ 174 U n d e t e r m i n e d h y p o s t o m e 1 ............................................................................................. 174 U n d e t e r m i n e d h y p o s t o m e 2 ............................................................................................. 175 O r d e r EODISCIDA K o b a y a s h i , 1939 .......................................................................................... 175 S u b o r d e r EODISCINA K o b a y a s h i , 1939 .................................................................................... 175 S u p e r f a m i l y E O D I S C O I D E A R a y m o n d , 1913 ............................................................................ 175 F a m i l y EODISCIDAE R a y m o n d , 1913 ....................................................................................... 175 G e n u s HELEPAGETIA Jell, 1975 ................................................................................................ 175

Helepagetia bitruncula Jell, 1975 .................................................................................... 176 REFERENCES ............................................................................................................................... 177 INDEX OF SPECIES AND GENERA ......................................................................................... 191 PLATES .......................................................................................................................................... 199

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13. 14. 15. 16. 17.

Text-figure 18. Text-figure 19. Text-figure 20. Text-figure 21. Text-figure 22. Text-figure 23. Text-figure 24.

Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Paibi section near Paibi, Huayuan, northwestern Hunan Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Wangcun section, northwestern Hunan Zonation and observed stratigraphic distribution of non-agnostoid species in the Huaqiao Formation of the Paibi-2 section near Paibi, Huayuan, northwestern Hunan Lectotype of Paranomocarella parallela Yang in Zhou et al., 1977 and the invalidly designated holotype of the species Reconstruction of cephalon and pygidium of Parayujinia constricta gen. et sp. nov. Buttsia globosa Lu and Lin in Peng, 1987, described originally as the holotype of Buttsia (Waergangia) spectabilis Lin, 1991, the type species of Buttsia (Waergangia) Lin, 1991 Type material of Fenghuangella laochatianensis laochatianensis Yang in Zhou et al., 1977 Type material of Fenghuangella coniforma Yang in Zhou et al., 1977 Holotype of Fenghuangella liostracinala Yang in Zhou et al., 1977 Holotype of Stigmatoa yangziensis Yang in Zhou et al., 1977 Lisania paratungjenensis Yang in Zhou et al., 1977 Type material of Lisania yuanjiangensis (Yang in Zhou et al., 1977) and Lisania hespera (Yang in Zhou et al., 1977) Type material of Baijingia zhengwanensis (Yang in Yang et al., 1991) Lectolype of Baojingia youshuiensis Yang in Zhou et al., 1977 Holotype of Baojingia jiudiantangensis (Yang in Zhou et al., 1977) Holotype of Baojingia paralala (Yang in Zhou et al., 1977) Type material of Baojingia quadrata (Yang in Zhou et al., 1977), and the holotype of Eoshengia paragenalata Yang in Zhou et al., 1977, which is now reassigned to B. quadrata The type material of Baojingia subquadrata (Yang, 1978) and the type material of Eoshengia spinosa Yang, 1978, which are now reassigned to E. subquadrata Reconstruction of cephalon and pygidium of Neoanomocarella asiatica Hsiang in Egorova et al., 1963 Holotype of Olenus austriacus Yang in Zhou et al., 1977 Holotype of Baikadamaspis sinensis (Yang in Zhou et al., 1977) Reconstruction of cephalon and pygidium of Baikadamaspis linearis sp. nov. Reconstruction of cephalon and pygidium of Baikadamaspis paibiensis sp. nov. Reconstruction of dorsal exoskeleton of Schmalenseeia sinensis Yang, 1978

• vii.

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INTRODUCTION This is the second and final volume documenting the polymerid trilobites from the Huaqiao Formation of northwestern Hunan, China. This volume includes the systematics of ptychopariids, eodiscids, trilobites of uncertain affinity, and some undetermined sclerites. Most geologic information concerning the Huaqiao Formation of northwestern Hunan, including lithostratigraphic data, biostratigraphic zonation, and detailed descriptions of three stratigraphic sections, is contained in Volume 1 of this two-part set. For convenience, however, stratigraphic range charts for the polymerids identified from the Paibi, Paibi-2, and Wangcun sections are repeated in this volume. In the following descriptions of fossil material, the collection numbers preceded by a P or W refer to levels in meters above the base of the Huaqiao Formation in the Paibi and the Wangcun sections, respectively. Numbers preceded by a P~ refers to level in meters above an arbitrary zero point (at the base of the massive breccia in the basal part of Bed 58) in the Paibi-2 section. ACKNOWLEDGMENTS This work is the outgrowth of nearly two decades of field and laboratory work, and we are grateful to the numerous colleagues and institutions that have provided support for this effort over the years. Among them, Chen Yongen, G. Geyer, A. R. Palmer, R. A. Robison, J. H. Shergold, and Zhang Wentang (W. T. Chang), have been particularly helpful in a variety of aspects of the study. Han Yaojun, Luo Kunli, Li Jun, Li Yue, M. N. Rees, and Wang Huayu aided in collecting fossils. G. A. Wasserman helped to prepare and photograph specimens, and J. St. John helped to locate some references. Ren Yugao helped to draft figures, and K. Polak helped to prepare the manuscript for publication. Research was supported by grants to S. Peng from the National Natural Science Foundation of China (49070077, 40072003, 40023002, 40332018), the Chinese Academy of Sciences (KZCX2-SW-129), the Ministry of Technology and Science of China (2001 DEB20056, G2000077700), the State Laboratory of Paleobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, the Chinese Academy of Sciences (No. 933114), the National Geographic Society (No. 5819-96; 7151-01), the National Scholarship Council for International Studies, State Education Commission of China; and by grants to L. E. Babcock from the National Science Foundation (EAR 9405990, 9526709, 0073089, 0106883, EAR OPP-0346829) and The Ohio State University (Seed Grant). REPOSITORIES Illustrated specimens are reposited in the following institutions. Acronyms used to identify the repositories precede the repository names. CUGB GMC HIT

China University of Geology (Beijing), Beijing, China Geological Museum of China, Beijing, China Nanjing Institute of Geology and Mineral Resources, the Chinese Academy of Geological .1.

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IGSK NIGP UQF USNM YIGM XTR

.6.

Sciences, Nanjing, China Institute of Geological Sciences, Kazakhstan Academy of Sciences, Almaty, Kazakhstan Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China Department of Geology and Mineralogy, University of Queensland, St. Lucia, Queensland, Australia United States National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA Yichang Institute of Geology and Mineral Resources, the Chinese Academy of Geological Sciences, Yichang, Hubei, China Xinjiang Regional Surveying Team, Bureau of Geology and Mineral Resources, Ortimqi, China

SYSTEMATIC PALEONTOLOGY Order PTYCHOPARIIDA Swinnerton, 1915 Suborder PTYCHOPARIINA Richter, 1933 Superfamily PTYCHOPARIOIDAE Matthew, 1887 ?Family

PTYCHOPARIIDAE

Matthew, 1887

Genus TOWNLEYELLAOpik, 1967

Type species. Townleyella townleyi Opik, 1967 (p. 385, pl. 8, figs. 4, 5; text-fig. 147), from the Glyptagnostus stolidotus Zone, O'Hara Shale, western Queensland, Australia; by original designation. Remarks. The suprageneric placement of this genus is problematic. When it was erected, Opik (1967) classified it as of uncertain superfamilial assignment within the Ptychopariina. This placement was based on the presence of bacculae in Townleyella townleyi. The bacculae prevented Opik (1967) from making a conclusive assignment. Opik (1967) noted the similarity of Townleyella to Calymenidius, although Calymenidius lacks a preglabellar field. Yuan and Yin (1998) referred some specimens from eastern Guizhou to Townlevella, and placed the genus in the family Paracedariidae Hfipe, 1955. However, the facial sutures, which are apparently not of the cerdariiform type, make an assignment to the Paracedariidae unsatisfactory. Because bacculae are variably presented in the Chinese material of Townleyella, we follow Jell and Adrain (2003) in assigning the genus with question to the Ptychopariidae. Townleyella rara Yuan and Yin, 1998 Plate 45, figures 10-12 1998 Townleyella rara Yuan and Yin, p. 147, 148, pl. 2, figs. 11, 12, 14, 15, ?13. ?1998 Cathayanella granulosa Yuan and Yin (in part), p. 149, pl. 2, fig. 10 only. 1998 Townleyella similis Yuan and Yin, p. 147, 148, pl. 2, figs. 16, 17.

Holotype. Cranidium with a broken glabella (Yuan and Yin, 1998, pl. 2, fig. 11, NIGP 127910), from the Glyptagnostus stolidotus Zone, Jimachong, Yuping, eastern Guizhou. New material. An early meraspid cranidium and two holaspid cranidia (illustrated specimens NIGP 138366, 138367), in collections W223.2 and W228.3. Remarks. The holaspid cranidia from Wangcun, northwestern Hunan, agree in all respects with the •

7

•

type material of Townleyella rara from eastern Guizhou. Key characters that warrant placement of the new material into this species include a moderately tapered, anteriorly rounded glabella with deeply incised S1 furrows, pit-like $2 furrows, faint $3 furrows, and a lack of bacculae; a narrow, convex, gently forwardly arched anterior border; a broad (tr.), moderately convex fixigena, nearly transverse eye ridges, converging anterior branches of the facial suture, and fine, densely spaced granules. In comparison to the holotype cranidium, the new material lacks a plectrum on the anterior border, which Yuan and Yin (1998) used as a character to differentiate T. rara from T. townleyi, the type species of Townleyella, which lacks a plectrum. However, examination of the type material of T. rara reveals that the plectrum is variably developed and is even obliterated on some paratypes, suggesting that it is a weak choice for a differential criterion. Townleyella similis is probably synonymous with T. rara. The paratype cranidium of T. similis (Yuan and Yin, 1978, pl. 2, fig. 17) appears to be identical in all respects with cranidia of T. rara. The holotype cranidium of T. similis, which occurs in a collection with T. rara, has a slightly less tapered glabella than those cranidia of T. rara. This difference is probably related to ontogenetic variation: the holotype of T. similis is much smaller in size than the paratype. In other respects, the holotype cranidium is indistinguishable from T. rara. Occurrence. The holotype is from the Glyptagnostus stolidotus Zone, Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it is occurs with species indicative of the lower part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone). Genus

XILINGXIA

Lu in Zhang et al., 1980

Type species. Yuehsienszella ichangensis Chang (Zhang, 1964, p. 25, pl. 1, figs. 3, 4; also Zhang et al., 1980a, p. 305, 306, pl. 101, figs. 8-10) from the upper part of the Tianheban Formation (upper lower Cambrian), at Shilongdong near Shipai, Yichang, western Hubei, central China; by original designation. Remarks. The genetic concept of Lu (in Zhang et al., 1980a) is followed here. The genus includes small ptychoparids having a sagittally depressed preglabellar field, subcentrally located palpebral lobes, faint lateral glabellar furrows, and subparallel anterior branches of the facial suture. As discussed by Lu (in Zhang et al., 1980a, p. 305), Parapoulsenia Rasetti, 1957, and Poulsenia Resser, 1936, both from Laurentia are closely comparable to Xilingxia. Parapoulsenia is distinguished by having deeper glabellar lateral furrows, more anteriorly located palpebral lobes, shorter posterior areas of the fixigenae (tr.), and converging anterior branches of the facial suture, whereas Poulsenia is distinguished by having narrower fixigenae, a more strongly tapered glabella, and converging anterior branches of the facial suture. In South China, Xilingxia is most comparable with Yuehsienszella, but Yuehsienszella differs in having a relatively shorter, more strongly tapered glabella, a wider (sag.) preglabellar field without a sagittal depression, and a wider axis in the thorax and pygidium. Yuan and Zhao (1999, p. 121) regarded Xilingxia as a junior synonym of Binodaspis Lermontova, 1951. Their proposal is rejected here as Binodaspis with its type species B. spinosa from the Lower Cambrian Protolenus Zone of eastern Siberia differs greatly from Xilingxia. The type material of B. spinosa includes two cranidia and one librigena (Lermontova, 1951b, pl. 19, figs. 5, 5a, 5b), none of which was designed as holotype. However, the cranidium figured as fig. 5 appears to be receipted as base of understanding the type species through the refigration in "Osnovy Paleontologii" by Chernysheva (1960, pl. 3, fig. 8). It differs from Xilingxia by having a notably .8-

longer preglabellar field that bears no sagittal depression, a proportionally shorter glabella that is much more tapered forward, a narrower palpebral field of the fixigena, and an oblique rather than transverse eye ridge. The librigena shows that B. spinosa differed in having a wider lateral border. The other cranidium in the syntypes of B. spinosa (Lermontova, 195 lb, pl. 19, fig. 5b) is from a different locality and seems not conspecific with the two other specimens. This cranidium has a wide (sag., exs.), transverse anterior border and a conical glabella with clearly defined S1 and $2 furrows, and a narrow fixigena. These features indicate that it differs from Xilingxia. The concept of Binodaspis seems obscure, and needs to be clarified. Specimens that were subsequently assigned to Binodaspis from eastern and western Sayan and the Siberian Platform (Egorova et al., 1960; 1987; Suvorova, 1960; Chernysheva, 1961; Egorova and Savitsky, 1969; Repina, 1972; Ogienko et al., 1974; Pegel, 1984) are variable in morphology. A few species, for example, B. paula Suvorova, 1960 and B. rara Chemysheva, 1961, are more or less comparable to Xilingxia. Besides the type species, at least seven species have been assigned to Xilingxia (Zhu, 1987; Xiang and Zhou, 1987; Zhang et al., 1980a). Among these species, X. convexa Xiang and Zhou (1987, p. 7, text-fig. 2-1) is apparently invalid (nomen nudum). X. brevica Zhang and Yao (in Zhang et al., 1980a, p. 102, figs. 1-4) and X. tenuis Zhang and Yao (fig. 5) are conspecific. The holotype cranidia of both X. brevica and X. tenuis, and most of the paratypes of X. brevica, are from the same collection (F20) at Yangjiaba, near Chengkou, Sichuan Province; there is no significant difference in morphology between cranidia of the two species. Even the assignment of these species to Xilingxia is problematical because their wide and short glabellas, and their incised lateral glabellar furrows, prevent definite assignment to the genus as conceived by Lu in Zhang et al. (1980a). For the same reason, the assignment of X. elongata Zhu, 1987 is also problematic. A species described as Chondragraulos (Antagmopleura) intercedens Chemysheva (1977, pl. 13, fig. 15) from Irgigkhemsk Formation (Middle Cambrian, Amgan Stage), Tuva, Russia, is closely comparable to X. ichangensis and may belong to Xilingxia. Xilingxia ichangensis (Chang, 1964) Plate 1, figures 15, 16 1957 Yuehsienszella szechuangensis (Sun); Zhang et al., p. 145 (in part, list only). 1964 Yuehsienszella ichangensis Chang; Zhang, p. 25, pl. 1, figs. 3, 4. 1980a Xilingxia ichangensis (Chang); Zhang, Lu, Zhu, Qian, Lin, Zhou, Zhang, and Yuan, p. 305, 306, pl. 101, figs. 8-10. 1984 Xilingxia ichangensis (Chang); Sun, p. 354, pl. 129, fig. 5. 1987 Xilingxia ichangensis (Chang); Xiang and Zhou, p. 7 (list only). Holotype. Complete exoskeleton and its counterpart (Zhang, 1964, pl. 1, figs. 3, 4, NIGP 15247; refigured by Zhang et al., 1980a, pl. 101, figs. 8, 9), from the Tianheban Formation, near Shilongdong, Yichang, western Hubei. New material. A cranidium, NIGP 137906, in collection W3. Remarks. Only one incomplete cranidium is present in new collections from northwestern Hunan. The cranidium is small, 2.6 mm long, and is characterized by having a glabella that is moderately long, rather convex, gently tapering forward, and rounded anteriorly, with three pairs of short, oblique lateral furrows, and a prominent occipital node that is located anteriorly. The preglabellar field is as about as wide as the anterior border (sag.), and bears a preglabellar median furrowlike .9.

depression sagittally. The palpebral lobes, although mostly broken, are located subcentrally, and the eye ridges are transverse and well defined. The palpebral area is wide and gently convex. The facial sutures have subparallel anterior branches and subdiagonally directed posterior branches. Xilingxia ichangensis is a rather small ptychoparid, characterized by having a glabella that is relatively more tapered in the posterior half and subparallel anterior to the glabellar midlength; the preglabellar field bears a broad, clearly depressed median preglabellar furrow sagittally and is nearly as wide (sa.) as the anterior border; the lateral glabellar furrows are short, and the eye ridges are transverse. The new cranidium, which was collected from the base of the Huaqiao Formation, northwestern Hunan, is identical in almost all respects to previously reported specimens of Xilingxia ichangensis (Chang), except that it has a more distinct occipital node. This subtle difference is regarded as an example of intraspecific variation. The type material of X. ichangensis from the Tianheban Formation, western Hubei, is of late early Cambrian age. X. ichangensis also has been reported from the upper part of the Megapalaeolenus Zone in the Tianheban Formation near Shipai, Yichang (Xiang and Zhou, 1987, p. 7). Discovery of this species in northwestern Hunan extends its range upward into the basal part of the Ptychagnostus atavus Zone.

Occurrence. The holotype is from the Tianheban Formation, near Shilongdong, Yichang, western Hubei, China. New material is from dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Ptychopariid gen. et sp. indet. Plate 1, figures 12-14

Material. A single incomplete, partly exfoliated cranidium (NIGP 137905) in collection P108. Remarks. This cranidium has a subquadrate, gently tapered, rather convex glabella that is defined by broad and deep axial and preglabellar furrows; a transverse front; a flat, forward-sloping preglabellar field; and an upturned anterior border that is slightly shorter (sag.) than the preglabellar field. The fixigena is narrow, and the palpebral lobe appears to be rather large and defined by a deep palpebral furrow. Two pairs of lateral glabellar furrow are visible: F1 is weak, straight, rather long, and diagonally directed; F2 is long, faint, and subparallel to F1. Overall, and especially in the glabellar shape, the size and the location of the palpebral lobes, and the shape of the fixigena, this illustrated specimen is mostly reminiscent of eulomaiids, especially the earlier eulomatids such as Protoeuloma Sdzuy, 1958, Kytina Rozova, 1963, and Euloma (Archaeuloma) Lee in Yin and Li, 1978, all from the upper Cambrian. These eulomaiids are characterized by having weak lateral glabellar furrows. However, these trilobites are different in the structure of the anterior cranidial area, which comprises a convex anterior border, a pitted anterior border furrow, and a preocular field that is commonly wider (sag.) than the anterior border.

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Dorypyge richthofeni Zone (equivalent to the lower part of the Ptychagnostus punctuosus Zone).

"10.

Superfamily

ASAPHISCOIDEA Raymond,

1924

Family ANOMOCARELLIDAEHup6, 1953 Genus FISSANOMOCARELLAPeng, Lin, and Chen, 1995 Type species. Fissanomocarella paibiensis Peng, Lin, and Chen, 1995 (p. 293, pl. 2, figs. 1-11; pl. 3, figs. 1-3) from the Ptychagnostus atavus Zone, Huaqiao Formation, Paibi, northwestern Hunan; by original designation. Remarks. This monospecific anomocarellid genus is characterized by a spinose pygidium. It was erected based on specimens from the Paibi section, northwestern Hunan, and most of the specimens are illustrated here. Peng et al. (1995, p. 292, 293) discussed the genetic concept, and that concept is followed here. Fissanomocarella paibiensis Peng, Lin, and Chen, 1995 Plate 2, figures 1-13 1995 Fissanomocarella paibiensis Peng, Lin, and Chen, p. 293, pl. 2, figs. 5-11; pl. 3, figs. 1-3. 2001b Fissanomocarella paibiensis Peng, Lin, and Chen; Peng, Babcock, and Lin, p. 101, pl. 2, figs. 7,8. Holotype. Cranidium (Peng et al., 1995, pl. 2, fig. 8, NIGP 118825; P1. 2, fig. 3 herein) from collection P82.5 (=HP14); upper part of the Ptychagnostus atavus Zone, Huaqiao Formation, Paibi, Huayuan, northwestern Hunan. Material. More than 20 sclerites including cranidia and pygidia (illustrated specimens 118822-118824, 118827-118830, 137907-137909) in collection P82.5. Diagnosis. See Peng et al. (1995, p. 292). Remarks. This species was fully described by Peng et al. (1995, p. 293). Additional specimens collected from the same locality and same bed as the holotype add no further information. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Genus HUAYUANIAPeng, Lin, and Chen, 1995 Type species. Huayuania subcalva Peng, Lin, and Chen, 1995 (p. 296, pl. 4, figs. 5-14; pl. 5, figs. 1-6) from the Lejopyge laevigata Zone, Huaqiao Formation, Paibi, northwestern Hunan; by original designation.

.11.

Other species. Proasaphiscus quadrilateralis Resser and Endo (1937, p. 263, 264, pl. 48, figs. 23-28), from the Changhia Formation, Shichangtun, Liaoyang, Liaoning. Remarks. The genetic concept of Peng et al. (1995) is followed here. Based on information from new pygidia, P. quadrilateralis Resser and Endo is transferred to the genus. Two cranidia from the Liopeishania Zone, Changhia Formation, near Hancheng, Shaanxi, which were referred to Liopeishania lubrica Chang (Zhang, 1957) by Luo (2001, pl. 3, figs. 15, 16) possibly belong to Huayuania. The pygidia referred to Peishania marginata (Resser and Endo) by Guo et al. (1996, pl. 59, figs. 4, 6; pl. 63, fig. 7) also may belong to Huayuania. The pygidia were not associated with cranidia assigned to P. marginata by Guo et al. (1967) and differ from the paratype pygidium of P. marginata (Endo and Resser, 1937, p. 249, pl. 46, 17; also Zhang and Jell, 1987, pl. 58, fig. 4) in having clearly defined, wide, and flat borders. Huayuania subcalva Peng, Lin, and Chen, 1995 Plate 3, figures 1-16; Plate 4, figures 1-8 1995 Huayuania subcalva Peng, Lin, and Chen, p. 296, pl. 4, figs. 5-14; pl. 5, figs. 1-6. 2001b Huayuania subcalva Peng, Lin, and Chen; Peng, Babcock, and Lin, p. 102, pl. 5, figs. 12, 13.

Holotype. Cranidium (Peng et al., 1995, pl. 4, fig. 7, NIGP 118848; P1. 3, figs. 9-11 herein) from collection P200.7 (=HP22); Lejopyge laevigata Zone, Huaqiao Formation, Paibi, Huayuan, northwestern Hunan. New material. More than 100 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 118846-118849, 118852, 118853, 118856, 118857, 118859, 118860, 118863, 124311, 137910-137920) in collection P200.7. Diagnosis. Peng et al. (1995, p. 295) provided a diagnosis for the species; new material does not warrant emendation of that information. Remarks. An ontogenetic series of this species was collected from the same locality and bed as the type material. It shows that the most distinctive changes during ontogeny are an increase in the degree of taper of the glabella and a decrease in the width of the fixigena. The pygidium becomes proportionately longer by a slight amount late in the holaspid period. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Pianaspis sinensis Zone (equivalent to the middle of Lejopyge laevigata Zone). Huayuania quadrilateralis (Resser and Endo, 1937) Plate 5, figures 1-14 1913 1937 1937 • 12.

Lisania cf. bura Walcott, p. 258, pl. 15, fig. 14. Proasaphiscua quadrilateralis Resser and Endo, p. 263,264, pl. 48, figs. 23-28. Aojia? vilis Resser and Endo, p. 178, pl. 32, figs. 9, ?10.

1960b Lisania quadrilateralis (Resser and Endo); Kobayashi, p. 370. 1965 Lisania quadrilateralis (Resser and Endo); Lu, Zhang, Zhu, Qian, and Xiang, p. 265, pl. 45, figs. 13, 14. 1965 Aojia? vilis Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 271, pl. 46, figs. 15, ?16. 1987 Lisania quadrilateralis (Resser and Endo); Zhang and Jell, p. 135, 136, pl. 51, figs. 11, 12; pl. 52, figs. 6, 7; pl. 54, fig. 2. 1987 Peishania sp. b; Zhang and Jell, p. 187, pl. 78, figs. 10, 11. 2001b Anomocarella sp.; Peng, Babcock, and Lin, p. 101, pl. 2, figs. 3, 4.

Lectotype. Cranidium (Resser and Endo, 1937, pl. 48, fig. 23, USNM 86880a; refigured by Zhang and Jell, 1987, pl. 52, fig. 7), from Changhia Formation, near Liaoyang, Liaoning; designated by Zhang and Jell (1987, p. 135, 305). New material. More than 30 sclerites, including cranidia and pygidia (illustrated specimens NIGP 137928-137940), in collections P42.5, P45.6, and P123.6. Emended diagnosis. Huayuania with cranidial length subequal to width between palpebral lobes; glabella tumid, notably convex sagittally, broadly rounded anteriorly; palpebral lobe subcentrally located, 0.26-0.3 of glabellar length. Anterior border long, weakly convex, and upturned; posterolateral projection transversely triangular. Pygidium rather effaced with small pleural field, and wide, flat borders. Description. Cranidium subqudrate in outline, about as long as the width between palpebral lobes. Glabella two-thirds as wide as long, slightly tapered forward or parallel-sided, broadly rounded anteriorly, notably convex in lateral profile; lateral glabellar furrows completely effaced or presented as obscure impressions; exfoliated surface with four pairs of lateral furrows: S 1 bifurcate, forming a subtriangular impression; $2 short, weakly impressed; $3 and $4 obscure; occipital furrow shallow, transverse; occipital ring narrowing abaxially with an obscure median node and rearward arched posterior margin. Anterior border very weakly convex, gently upturned, anterior border furrow well impressed; preglabellar field very short. Preglabellar furrow almost confluent sagittally with anterior border furrow on some specimens. Palpebral lobe gently curved, relatively small in size, defined by faint palpebral furrow; anterior branches of facial suture divergent at about 30 ° to sagittal line; posterior branches straight, outward and rearward directed; posterolateral projection triangular, less than half of basal width of glabella. Pygidium semicircular, mostly effaced, length more than half of width. Axis subconical, with straight lateral margins, rounded posteriorly; with four to five weak tings and semicircular terminal piece, reaching or extending beyond posterior border furrow. Pleural field narrower than axis, gently convex; pleural furrow weakly impressed, nearly straight, slightly oblique rearward; pleurae obscure or effaced. Lateral and posterior borders wide, flat, almost uniform in width. Remarks. Kobayashi (1960b) transferred Proasaphiscus quadrilateralis to Lisania, and such an assignment was followed by some subsequent authors (Lu et al., 1965; Zhang and Jell, 1987). Zhang and Jell (1987) opined that some pygidia with effaced furrows and wide borders that were associated with the cranidia of P. quadrilateralis are not consistent with an assignment to Lisania and excluded the specimens from the species. New material from the Huaqiao Formation in northwestern Hunan confirms that the pygidia that Resser and Endo (1937) originally assigned to P. quadrilateralis are associated correctly, and further suggests that this species is better referred to Huayuania. Based on the new material, the pygidia that were excluded from the species by Zhang •

13.

and Jell (1987) are reassigned back to the species. Besides the cranidium and pygidium, the librigena figured by Resser and Endo (1937, pl. 48, fig. 25; also Zhang and Jell, 1987, pl. 52, fig. 6) for P. quadrilateralis is also consistent with that of Huayuania. This species differs from Huayuania subcalva, the type species of Huayuania, in having a less tapered and less convex glabella, smaller palpebral lobes that are rather closely spaced, and wider (exs.) posterolateral projections. The first appearance of H. quadrilateralis in stratigraphy precedes that of H. subcalva. Huayuania quadrilateralis is similar to Sudanomocarina changi, the type species of Sudanomocarina Jell (in Jell and Robison, 1978, pl. 4, figs. 9-16), but S. changi can be differentiated by having a longer cranidial outline, a lower convexity, longer, and more posteriorly located palpebral lobes that are relatively closely spaced. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone and Pianaspis sinensis Zone (equivalent to the Ptychagnostus atavus Zone through P. punctuosus Zone). Genus PARANOMOCARELLAYang in Zhou et al., 1977 Paranomocarella Yang in Zhou et al., 1977, p. 183; Yin and Li, 1978, p. 505; Yang, 1978, p. 53; Liu, 1982, p. 314; Lin, Lin, and Zhou, 1983, p. 405; Yang and Liu in Yang et al., 1991, p. 161; 1993, p. 208; Peng, Lin, and Chen, 1995, p. 283, 295,296. Type species. Paranomocarella parallela Yang in Zhou et al., 1977 (in part, p. 183, pl. 54, fig. 1, non fig. 2) from the Lisania tungjenensis Zone, Huaqiao Formation, Huaqiao, Baojing, western Hunan and Fengmuping, Tongren, eastern Guizhou. Other species. Among the species listed by Peng et al. (1995, p. 294), Paranomocarella parapolita Yang in Yin and Li, 1978 should be excluded. To this list should be added Jiangsuia? cylindrica Qian and Zhou (1984, p. 179, pl. 2, figs. 3, 4) from the "Paotaishan" Formation, near Kunshan, Jiangsu; Paranomocarella fortis Peng et al. (1995, p. 283,294, 295, pl. 1, fig. 12a; pl. 3, figs. 4-13; pl. 4, figs. 1-4) from the Ptychagnostus atavus Zone to the Lejopyge laevigata Zone, Huaqiao Formation, Paibi and Wangcun, northwestern Hunan; Paranomocarella similaris sp. nov., and Paranomocarella yangi sp. nov. (Yang in Zhou et al., 1977, pl. 54, fig. 2, as reassigned herein). Remarks. Designation of a lectotype for Paranomocarella parallela, the type species of Paranomocarella, is a confusing issue. No holotype was designated when the species was erected. Syntypes of the type species include a cranidium and a pygidium (Yang in Zhou et al., 1977, pl. 54, figs. 1, 2). According to Yang (1978), the2¢ are from the same locality and the same formation (the Huaqiao Formation), but from different biozones (see Yang, 1978). The cranidium is from the Lisania tungjenensis Zone, whereas the pygidium is from the Dorypyge cf. richthofeni Zone, which is older in age. Yang (1978) designated a "holotype" cranidium (Yang, 1978, pl. 9, fig. 2) for P. parallela, but it is an invalid designation because it was not selected from the syntypes (see International Commission on Zoological Nomenclature, 1999, Art. 74.2). The syntype cranidium is here designated as lectotype of the type species. Yang (1978) designated this lectotype cranidium as the "holotype" of P. parapolita Yang (1978, p. 54, pl. 9, figs. 4-6). The selection of the lectotype for the type species makes P. parapolita a junior objective synonym of P. parallela because they are based on the same cranidium. Furthermore, the cranidium figured by Yang as the "holotype" of .14.

P. parallela (1978, pl. 9, fig. 2; also Yin and Li, 1978, pl. 169, fig. 17 and herein Text-fig. 4C, D) does not seem to be conspecific with P. parallela, but, as discussed below, it may belong to P. fortis Peng et al., 1995. The syntype pygidium of P. parallela may represent a new species of Paranomocarella. Paranomocarella resembles Anomocarella Walcott, 1905 but differs from it primarily in having more posteriorly placed palpebral lobes, narrower palpebral areas, lower convexity in the cranidium, a proportionately smaller glabella, and a well-furrowed pygidium with a well defined paradoublural line. The librigena of Paranomocarella (Peng et al., 1995, pl. 3, figs. 6, 9, 12) has a short genal spine with a broad base; in Anomocarella the librigena is long and narrow at the base of the genal spine. Most species of Paranomocarella have three to four pairs of faint lateral glabellar furrows, of which the S1 furrow is bifurcated. Some species show a median cafina on glabella. In Anomocarella, the lateral glabellar furrows are commonly absent or, if present, they are almost obliterated. Paranomocarella parallela Yang in Zhou et al., 1977

Plate 7, figures 10-16; Text-figure 4 A, B 1977

Paranomocarella parallela Yang in Zhou et al. (in part), p. 183, pl. 54, fig. 1, non fig. 2 [=Paranomocarella yangi sp. nov.]. non 1978 Paranomocarella parallela Yang; Yin and Li (in part), p. 505, 506, pl. 169, fig. 16 [=Paranomocarella yangi sp. nov.], fig. 17 [=Paranomocarellafortis Peng et al., 1995]. 1978 Paranomocarella parapolita Yang in Yin and Li (in part), p. 506, pl. 169, fig. 15, non fig. 14 [=Paranomocare bagushangensis Zhu and Sun in Zhou et al., 1977]. 1978 Paranomocarella parapolita Yang; Yang (in part), p. 54, pl. 9, figs. 4, 5, non fig. 6 [=Paranomocare bagushangensis Zhu and Sun in Zhou et al., 1977]. non 1978 Paranomocarella parallela Yang, p. 53, 54; pl. 9, fig. 2 [=Paranomocarellafortis Peng et al., 1995], fig. 3 [=Paranomocarella yangi sp. nov.] 1982 Paranomocarella parallela Yang; Liu (in part), p. 314, pl. 220, fig. 13, non fig. 19 [=Paranomocarella yangi sp. nov.]. non 1983 Paranomocarella parallela Yang; Lin, Lin, and Zhou, p. 405, pl. 2, fig. 13 [=Paranomocarella fort& Peng, Lin, and Chen, 1995]. 1983 Paranomocarella parapolita Yang; Lin, Lin, and Zhou, p. 405, pl. 2, figs. 14, 15. 2001b Paranomocarella parallela Yang; Peng, Babcock, and Lin, p. 101, pl. 2, figs. 12, 13. ?2001 b Paranomocarella fortis Peng, Lin, and Chen; Peng, Babcock, and Lin, p. 102, pl. 2, fig. 14. Lectotype. Designated herein; cranidium (Zhou et al., 1977, pl. 54, fig. 1, CUGB 0112702; Text-fig. 4A, B herein), from the Lisania tungjenensis Zone, Huaqiao, Baojing, northwestern Hunan. This cranidium was refigured as Paranomocarella parapolita Yang by Yin and Li (1978), and subsequently designated as holotype of Paranomocarella parapolita Yang by Yang (1978). New material. Two cranidia and seven pygidia in collections P38.5, P45.2, P45.6, P82.5, and P112.6. Emended diagnosis. Paranomocarella with anterior border that is flat or slightly concave, wider than preglabellar field; glabella proportionately wide and short, parallel sided, lacking median carina; lateral glabellar furrows mostly effaced. Pygidium subtriangular, width less than twice width; doublure relatively narrow.

.15-

Description. Glabella large, gently convex, parallel-sided, well rounded anteriorly, with weak or faint S 1 furrow. Occipital furrow shallow; occipital ring slightly wider than basal glabellar width, bearing weak median node. Anterior border moderately long to long, moderately concave, defined posteriorly by shallow to distinct anterior border furrow that is divided sagitally into two parts by wide, poorly defined plectrum. Palpebral area about one-third as wide as glabella. Anterior branch of facial suture diverging forward about 90 ° . Pygidium semicircular, length about three-fifths width, borders gently concave. Axis reaching to paradoublural line, with short, postaxial ridge extending slightly beyond paradoublural line; anterior part poorly divided into 5-6 tings, posterior part obscurely segmented. Pleural field moderately convex, with faint pleural furrows and weak interpleural furrows; both pleural and interpleural furrows long and rather straight, extending onto or beyond paradoublural line. Paradoublural line shallow but clearly defined. Remarks. As discussed above under the genus Paranomocarella, the "holotype" designated by Yang (1978) for Paranomocarella parallela, the type species of Paranomocarella, is invalid, and P. parapolita should be suppressed as a junior objective synonym of P. parallela. The "holotype" designated by Yang (1978) for P. parallela belongs not to that species but probably to P. fortis Peng et al. (1995).

Text-figure 4. A, B, Lectotype of Paranomocarella parallela Yang in Zhou et al., 1977, selected herein from syntypes of this species; an incomplete cranidium in dorsal and obliquely anterior views, CUGB 0112702, both × 2.3 (original of Zhou et al., 1977, pl. 54, fig. 1; also Yang, 1978, pl. 9, fig. 4); C, D, Paranomocarella fortis Peng, an incomplete cranidium in dorsal and anterolateral views, CUGB 0114203, × 4, original of Yang, 1978, pl. 9, fig. 2. This cranidium was invalidly designed as the holotype of Paranomocarella parallela by Yang (1978, p. 53, 80). -16.

The syntype cranidium (which is designated here as the lectotype) and the syntype pygidium of P. parallela are from different stratigraphic horizons: the cranidium is from the Lisania tungjenensis Zone, whereas the pygidium is from the older Dorypyge cf. richthofeni Zone. Yang (in Yin and Li, 1978), subsequent to his original description of P. parallela (Yang in Zhou et al., 1977), noted that the cranidium and pygidium belong to different species. New collections confirm that the pygidium is not conspecific with the holotype cranidium, and that it represents a previously undescribed species of Paranomocarella. That species, Paranomocarella yangi sp. nov., is characterized by a short axis and distinctive pleural furrows that increase in depth and width beyond the paradoublural lines. The pygidium associated subsequently with the holotype of P. parallela by Yang (in Yin and Li, 1978, pl. 169, fig. 14) is not from Huaqiao, but from Wanshan, Yuping, eastern Guizhou. This pygidium is not a Paranomocarella because it lacks a paradoublural line and bears rather weak pleural furrows. Instead, it agrees well with the pygidium of Paranomocare bagushangensis Zhu and Sun (in Zhou et al., 1977, pl. 52, fig. 4) from the Guangzhuling Formation (Middle Cambrian), at Bagushang, Xianfeng, southwestern Hubei, and should be referred to that species. New material from the Huaqiao Formation agrees with the holotype of the species in all essential characters except for having a slightly narrower (sag., exs.) anterior cranidial border. The anterior branches of the facial suture seem to be more widely diverging in the new material than in the holotype, but examination of the holotype shows that this is not the case, because the left preocular area of the holotype is not completely exposed. The new material shows slight variation in the pygidial outline and the length of the pygidial axis. Stratigraphically lower specimens have outlines that are proportionately wider, and have axes that are relatively shorter, than those in stratigraphically higher layers. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone to the Pianaspis sinensis Zone (equivalent to the Ptychagnostus atavus Zone and the Ptychagnostus punctuosus Zone). Paranomocarella fortis Peng, Lin, and Chen, 1995 Plate 6, figures 1-16; Plate 7, figure 1; Text-figure 4 C, D 1978 Paranomocarella parallela Yang in Zhou et al. (in part); Yin and Li, p. 505, 506, pl. 169, fig. 17 only. 1978 Paranomocarella parallela Yang; Yang (in part), p. 53, 54, pl. 9, fig. 2 only. 1983 Paranomocarella parallela Yang; Lin, Lin, and Zhou, p. 405, pl 2, fig. 13. 1995 Paranomocarella fortis Peng, Lin, and Chen (in part), p. 283, 294, 295, pl. 1, fig. 12a; pl. 3, figs. 4-9, 11-13; pl. 4, figs. 1-4, non pl. 3, fig. 10 [=Paranomocarella similaris sp. nov.]. 2001b Paranomocarella fortis Peng, Lin, and Chen; Peng, Babcock, and Lin (in part), p. 102, pl. 3, fig. 13, non figs. 11, 12 [=Paranomocarella similaris sp. nov.]. non 2001b Paranomocarella fort& Peng, Lin, and Chen; Peng, Babcock, and Lin, p. 102, pl. 2, fig. 14 [=Paranomocarella parallela Yang]. Holotype. Cranidium (Peng et al., 1995, pl. 3, figs. 4a, b, NIGP 118832; P1. 6, figs. 7, 8 herein) from collection P122.4. New material. More than 30 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 118833, 118835, 118837, 118840, 118842, 118844, 118845, 137941-137948)in -17-

collections P82.1, P108, P123.6, P130.5, P131.7, P200.7, and W56.7 Remarks. New material of Paranomocarella from the Huaqiao Formation at Paibi, northwestern Hunan, shows that two cranidia and one pygidium previously included in P. fortis (Peng et al., 1995, 2001b) do not belong to P. fortis. A previously illustrated cranidium (Peng et al., 1995, pl. 3, fig. 10) and pygidium (Peng et al., 2001b, pl. 3, fig. 12) are here reassigned to P. similaris sp. nov., and another described pygidium (Peng et al., 2001b), which occurs in association with other sclerites of P. parallela, is reassigned to P. parallela. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone to the Pianaspis sinensis Zone (equivalent to the Ptychagnostus atavus Zone to the Lejopyge laevigata Zone). Paranomocarella similaris sp. nov. Plate 7, figures 2-9 1995 Paranomocarella fortis Peng, Lin, and Chen (in part), p. 283,294, 295, pl. 3, fig. 10 only. 2001b Paranomocarella fortis Peng, Lin, and Chen; Peng, Babcock, and Lin (in part), p. 102, pl. 3, figs. 11, 12 only. Etymology. From Latin, similaris, similar, referring to its similarity to Paranomocarella fortis. Holotype. Cranidium (P1.7, fig. 3, NIGP 137950) from collection P130.5. Other material. Five cranidia and eight pygidia (illustrated specimens NIGP 118838, 137949137955) in collections P126.9, P130.25, P130.5, P131.7, P134.2, P135.7, P136.3 and W82.1. Diagnosis. Paranomocarella with anterior border that is convex and narrow; glabella proportionately short, tapered, beating median carina and lateral furrows. Pygidium subelliptical, length greater than half of width; doublure moderately wide. Description. Cranidium subquadrate, length approximately same as width of anterior area. Anterior border short (sag., exs.), moderately convex, defined posteriorly by distinct anterior border furrow that is divided sagitally into two parts by short, poorly defined plectrum. Preglabellar area 1.5-2 times as long as anterior border. Glabella occupying three-fourths to two-thirds of cranidial length, tapering gently forward, rounded or acutely rounded anteriorly, with weakly defined carina; with three pairs of lateral glabellar furrows; S1 clearly impressed, bifurcated, $2 and $3 weakly impressed. Occipital furrow well impressed; gently curved rearward; occipital ring gently convex (sag., tr.), narrowing gently abaxially, beating tiny median node. Palpebral area about half as wide as glabella. Anterior branch of facial suture diverging forward about 90 °. Anterior area beating fine, dense radiating caecae. Pygidium semicircular, length about two-thirds width, pleural field moderately convex, borders flat or gently concave. Axis long, thin, occupying three-fourths of pygidial length and reaching to paradoublural line, extending as short, faint postaxial ridge; anterior part poorly divided into 6-7 tings, posterior part obscurely segmented. Pleural field with well impressed interpleural furrows and faint pleural furrows; pleural furrows long, nearly straight, extending onto borders or "18-

nearly to pygidial margin; interpleural furrows shorter than pleural furrows, extending onto or slightly beyond paradoublural line. Paradoublural line shallow but clearly defined; doublure evenly wide, occupying about one-fourth of pygidial length (sag.).

Remarks. This new species is similar to Paranomocarella fortis Peng et al., 1995 in having a distinct anterior cranidial border and plectrum, strongly diverging anterior branches of the facial suture, and a relatively long preglabellar field. However, it differs in having a tapered, well furrowed, and carina-bearing glabella, and a pygidium that is proportionally longer, with more rounded anterolateral comers. In P. fortis, the glabella is parallel-sided, largely effaced, evenly convex, without a median carina; the pygidium is wider than long, with less rounded anterolateral comers and a relatively wide doublure that is slightly widening forward, rather than narrowing, as in the new species. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone). Paranomocarella sp. Plate 45, figures 15, 16

Material. Two pygidia (NIGP 138370, 138371) in collection P82.5. Remarks. Two similar pygidia from the same bed in the Huaqiao Formation at Paibi, Huayuan, Hunan, are referred to Paranomocarella, although the species assignment is uncertain. The specimens are similar to those assigned here to Paranomocarella parallel, differing only in having narrower lateral and posterior borders that are flat, rather than concave, and by lacking the paradoublural line. These two pygidia are from the same collection as P. parallela in the Paibi section. More material is needed to clarify whether they are conspecific with that species. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Paranomocarella? obvia sp. nov. Plate 8, figures 14-16 2001b Anomocarellid gen. et sp. indet.; Peng, Babcock, and Lin, p. 103, pl. 9, fig. 4.

Etymology. From Latin, obvius, obvious, referring to the clearly defined interpleural furrows and paradoublural furrows on the pygidium. Holotype. Pygidium (P1. 8, fig. 15, NIGP 137977) from collection P277. Other material. Two pygidia (NIGP 137976, 137978), in collections P269 and P277. .19.

Diagnosis. Paranomocarella? with pygidium length two-thirds width; axis relatively wide, occupying three-fourths sagittal length of pygidium, with anterior four tings well defined; pleural and interpleural furrows clearly impressed and deepened beyond paradoublural line; pleura with narrow anterior band and wide posterior band separated by diagonally directed pleural furrow; paradoublural line deeply impressed. Description. Pygidium subtriangular, width three-fourths length. Axis wide, relatively short, bearing five tings and short, semicircular terminal piece, reaching nearly to paradoublural line, extending as short, thin, and moderately defined postaxial ridge. Pleural field as wide as axis, slightly convex, defined clearly by moderately deep paradoublural line; pleural and interpleural furrows long, well impressed, extending nearly to pygidial margin, dividing each pleura into narrow anterior band and much wider posterior band; furrows deepened distally, so that they outline ridge-like thin band on distal part of borders. Borders moderately wide, occupying about one-fifth of pygidial length (sag.), widening slightly abaxially. Remarks. Two well furrowed pygidia from the Huaqiao Formation show close similarity to those of Paranomocarella in having an axis that is moderately segmented with four well defined tings, one weakly defined ring, and a short terminal piece; the axis reaches to or nearly to the paradoublural line; the paradoublural line is clearly defined, and there is a relatively wide doublure. The gently curved pleural furrows that extend beyond the paradoublural line and the narrow borders are consistent with assignment to Paranomocarella. The pygidia, however, are distinguished from known Paranomocarella species by being proportionately longer, having a comparatively wider axis, having more deeply impressed interpleural furrows, and having a deeper paradoublural line. The overall morphology suggests that the new pygidia represent a new species of Paranomocarella, but until a cranidium is discovered for this species, the specimens are left in open nomenclature, and only questionably referred to this genus. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone). Genus SZEASPIS Chang, 1959

Szeaspis Chang (Zhang), 1959, p. 207, 208, 229, 230; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 306; Zhou, Liu, Meng, and Sun, 1977, p. 180; Jell and Hughes, 1997, p. 57; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 150; Zhang and Jell, 1987, p. 153; Yang et al., 1991, p. 159; 1993, p. 206. Spitiaspis Kobayashi, 1967, p. 488; Jell and Adrain, 2003, p. 447. Type species. Szeaspis reticulatus Chang, 1959 (Zhang, p. 207, 208, 230, pl. 3, figs. 11-16) [=Anomocare conjunctiva Reed, 1910] from the Fuchouia-Luia Subzone, Changhia Formation, Boshan, Shandong; by original designation. Other species. Anomocare conjunctiva Reed, 1910 [=Proasaphiscus centronatus Resser and Endo, 1937, p. 250, 251, pl. 37, figs. 17-20; = Szeaspis reticulatus Chang, 1959 =S. reticulatus var. brevicus Chang, 1959, p. 208, pl. 3, figs. 7-10], from the middle Cambrian at Changnu campground, Spiti, India; Manchuriella pustulosa Resser and Endo (1937, in part, p. 244, 245, pl. • 20.

36, fig. 13, non figs. 14, 15) from the Amphoton Zone of the Changhia Formation, Xibianling, Taizihe district, Liaoning; Asaphiscus iddingsi Walcott (1911, p. 99-101, pl. 16, fig. 3) from the Crepicephalina Zone, Changxingdao, eastern Liaoning; Szeaspis mopoensis Mong in Zhou et al., 1977 (p. 180, pl. 53, fig. 9) from the Changhia Formation at Mopo, Gongxian, Henan; and Szeaspis huananensis sp. nov. Zhang and Jell (1987) questionably referred Conokephalina maia Walcott (1913, pl. 13, fig. 13) to the Szeaspis. Assignment of a cranidium from C. D. Walcott's collection to Szeaspis sp. (Zhang and Jell, 1987, p. 155, 156, pl. 61, fig. 4) also seems questionable. Judging from the short preglabellar field and crescentic occipital ring of the specimen, it seems more likely to be Manchuriella than Szeaspis.

Remarks. The genetic concept of Zhang (1959, p. 208, 229) is followed here. However, Szeaspis is classified in the Paranomocarellidae on the basis of the long preglabellar field and the relatively large pygidium. Jell and Hughes (1997) suppressed Spitiaspis Kobayashi, 1967 as a junior synonym of Szeaspis. Zhang and Jell (1987) considered Szeaspis reticulatus, the type species of Szeaspis, to be a junior synonym of S. centronatus (Resser and Endo), and Jell and Hughes (1997) suppressed both species as junior synonyms of S. conjunctiva (Reed).

Szeaspis huananensis sp. nov. Plate 8, figures 1-13 2001b Paranomocarella?sp. 1; Peng, Babcock, and Lin, p. 101, pl. 2, fig. 5. 2001b Paranomocarella?sp. 2; Peng, Babcock, and Lin, p. 102, pl. 3, figs. 7, 8. 2001e Paranomocarella?sp., Peng, Babcock, Lin, Chen, and Zhu, p. 159, fig. 10.19-10.21.

Etymology. From Chinese, Huanan, South China, in reference to this, the first record of a Szeaspis species in South China.

Holotype. Cranidium with broken anterior border (P1. 8, fig. 8, NIGP 137970) from collection W56.7.

Other material. More than 30 sclerites, including an ontogenetic series of cranidia and pygidia, and one hypostome (illustrated specimens NIGP 137963-137969, 137971-137975), in collection W56.7.

Diagnosis. Szeaspis with long preglabellar field; glabella with three pairs of shallow lateral furrows; palpebral lobe strongly arcuate, located posteriorly, defined by shallow palpebral furrow; anterior branch of facial suture strongly diverging. Pygidium transverse, with axis that is slender, elongate-conical, and weakly segmented; doublure wide.

Description. Cranidium subquadrate, length subequal to width or slightly greater than width. Anterior border slightly convex, with sagittal length about half of preglabellar field, narrowing abaxially, without plectrum on posterior margin; anterior border furrow clearly defined, arched forward gently. Glabella subconical, gently convex, tapering moderately forward, rounded anteriorly, occupying 0.58-0.62 cranidial length; with three pairs of lateral furrows; S1 and $2 moderately long, weakly impressed, directed inward and slightly rearward; $3 short, obscure, • 21 •

directed inward. Occipital furrow shallow, equal in depth or slightly deepened at sides, curved slightly rearward; occipital ring of uniform width, bearing weak, subcentrally located node. Eye ridge moderately long, directed rearward slightly obliquely; palpebral lobe moderately long, arcuate, located posteriorly, defined by shallow palpebral furrow; palpebral area wide, with width slightly more than half that of glabella between palpebral lobes, gently inclined to axial furrow. Anterior branch of facial suture diverging abaxially, enclosing narrow (exs.) posterolateral projection. Hypostorne with subovate middle body that is divided by faint middle furrow into large, subcircular anterior lobe and short, crescentic posterior lobe, slightly sinuous anteriorly and obtusely rounded posteriorly. Lateral and posterior borders narrow, ridge-like, defined by deep lateral furrow and shallow posterior furrow that is even more shallow medially. Anterior wing subtriangular, strongly sloping downward. Pygidium transverse, width twice length, with rounded anterolateral comer. Axis slender, elongate-conical, with five tings, semicircular terminal piece, and short and obscure postaxial ridge. Pleural furrows broad and well impressed, interpleural furrows almost obliterated. Paradoublural line well impressed; doublure occupying about one-third of pygidial length, narrowing slightly posteriorly. Pleural field gently convex inside of paradoublural line, gently concave outside of doublural line; borders narrow, poorly defined.

Remarks. With the exceptions of one species from Spiti, India, all other species previously assigned to Szeaspis are from the North China Platform. The new species from the Huaqiao Formation of northwestern Hunan is differentiated from all other species in the genus by the combination of having a long preglabellar field; diverging anterior branches of the facial suture; a rather strongly transverse pygidium with a slender, rather wide axis; and a distinct paradoublural line. An incomplete ontogenetic series shows that the glabella changes from subcylindrical to subconical, and that the palpebral lobe changes from gently curved to strongly arcuate. Little change other than size occurs in the pygidium through ontogeny.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus punctuosus Zone). Family

PROASAPHISCIDAE

Chang, 1963

Remarks. Zhang and Jell (1987, p. 141, 142) discussed the familial concept and included 14 genera in the family. This concept is expanded to embrace two new genera, Huayuanaspis and Paibiella, each of which bears an occipital spine. Adelogonus Opik is now classified within the Proasaphiscidae. When Opik (1967) erected this genus, he placed it in the superfamily Ptychoparioidea, but its familial position remained unassigned. Material from eastern Guizhou (Yuan and Yin,1998) and northwestern Hunan (herein) shows that Adelogonus has a close affinity with proasaphiscoids. Genus

PROASAPHISCUS

Resser and Endo in Kobayashi, 1935

Type species. Proasaphiscus yabei Resser and Endo in Kobayashi, 1935 (p. 287, pl. 24, fig. 16) from the Hsuchuang Formation, Tangshishan, near Yantai, Liaoning; by original designation.

Remarks. The genetic concept of Zhang and Jell (1987, p. 142) is followed here. •

22.

Proasaphiscus? sp. cf. P. butes (Walcott, 1905) Plate 14, figure 11 cf. cf. cf. cf.

1905 1913 1937 1965

Anomocare? butes Walcott, p. 49, 50. Anomocare butes Walcott; Walcott, p. 199, 200, pl. 19, figs. 7, 7a-d. Proasaphiscus affiuens Resser and Endo, p. 265, pl. 48, figs. 1-13. Proasaphiscus affiuens Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 285, pl. 49, figs. 12-16. cf. 1965 Proasaphiscus butes Walcott; Lu, Zhang, Zhu, Qian, and Xiang, p. 285, 286, pl. 49, figs. 17, 18. cf. 1987 Proasaphiscus butes Walcott; Zhang and Jell, p.142-144, pl. 39, fig. 12, pl. 55, fig. 6, pl. 57, figs. 1-13; pl. 58, figs. 1-4. Lectotype of Proasaphiscus butes. A cranidium (Walcott, 1913, pl. 19, fig. 7, USNM 58168; refigured by Zhang and Jell, 1987, pl. 57, figs. 11, 12), from the Bailiella-Lioparia Zone, Hsuchuang Formation, Xintai, Shandong; designated by Zhang and Jell (1987, p. 143, 308). Material. Single cranidium, NIGP 138029, in collection P34.3. Remarks. A broken, mostly exfoliated cranidium from the Huaqiao Formation, northwestern Hunan, is assigned questionably to Proasaphiscus. The cranidium has a wide, subrectangular glabella with nearly straight sides that taper forward, and an obtusely rounded front; four pairs of faint lateral glabellar furrows, with S1 being bifurcated; and a faint carina. The specimen is most closely comparable with the cranidium of Proasaphiscus butes (Walcott, 1913, pl. 19, fig. 7; Zhang and Jell, 1987, pl. 57, figs. 11, 12). The new cranidium is identical with the holotype of Proasaphiscus butes in all observed respects except for the preglabellar field, which is somewhat shorter. Because the single available specimen is broken, it can be only questionably assigned to Proasaphiscus. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Subgenus PROASAPHISCUS(HONANASPIS)Chang, 1959

Proasaphiscus (Honanaspis) Chang (Zhang), 1959, p. 204, 205, 226, 227; Zhang and Jell, 1987, p. 150; Zhang in Zhang et al., 1995, p. 60; Guo, Zan, and Luo, 1996, p. 77. Honanaspis Chang; Zhang in Lu et al., 1965, p. 294, 295; Zhou, Liu, Meng, and Sun, 1977, p. 178; Yin and Li, 1978, p. 498; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 140, 141; Zhang and Wang, 1985, p. 418. Type species. Proasaphiscus (Honanaspis) honanensis Chang (Zhang, 1959, p. 205, 206, 227, 228, pl. 3, figs. 1-4; text-fig. 24), from the Bailiella Zone, Hsuchuang Formation, Laofenkou, Songbiao, near Dengfeng, western Henan; by original designation.

• 23-

Remarks. Zhang (in Lu et al., 1965) elevated Honanaspis from subgeneric status to genetic status, and this assignment has been followed by a number of authors. However, according to Zhang (1959) it differs from Proasaphiscus only in having slightly shorter palpebral lobes, a thorax with 11 rather than 13 segments, shorter pleural spines, and a smaller pygidium. Most of these distinctions, however, have only specific significance. We follow Zhang (1959) in regarding Honanaspis as a subgenus of Proasaphiscus. Proasaphiscus (Honanaspis)? sp. cf. P. (H.) parvus Kuo in Lu et al., 1965 Plate 10, figure 2 non 1959 Proasaphiscus(Honanaspis) honanensis Chang (Zhang), p. 205, pl. 3, figs. 1-4; text-fig. 24. cf. 1965 Honanaspishonanensis Chang; Zhang in Lu et al., p. 295, pl. 52, fig. 14. non 1959 Proasaphiscus (Honanaspis) honanensis Chang; Zhou, Liu, Meng, and Sun, p. 178, 179, pl. 53, figs. 1, 2. cf. 1965 Honanspisparvus Kuo in Lu et al., p. 296, pl. 53, fig. 3. cf. 1977 Honanspisparvus Kuo; Zhou, Liu, Meng, and Sun, p. 179, pl. 52, fig. 13. cf. 1995 Honanspisparvus Kuo; Zhang and Meng, p. 86. Holotype ofP. (Honanaspis) parvus. By monotypy; cranidium (Lu et al., 1965, pl. 53, fig. 3, GMC, catalog number unknown), from the Bailiella Zone, Hsuchuang Formation, Gongxian, Henan. Material. One incomplete cranidium, NIGP 137987, in collection P33.5. Remarks. Zhang and Meng (in Zhang et al., 1995, p. 86) suppressed Proasaphiscus (Honanaspis) parvus as a junior synonym of P. (Honanaspis) honanensis, the type species of Proasaphiscus (Honanaspis). This view is rejected here because P. (H.) parvus differs in having strongly furrowed glabella. As figured by Zhang (1959, pl. 3, figs. 1-4), the glabella of the type specimens of the type species is notably effaced. The poorly preserved cranidium from the Huaqiao Formation, northwestern Hunan resembles Proasaphiscus (Honanaspis) parvus. This specimen is characterized by a gently convex, rectangular glabella with an obtusely rounded front; four pairs of deeply impressed lateral glabellar furrows; a narrow and upturned, forwardly convex anterior border defined by a shallow and broad anterior border; a narrow preglabellar field; a narrow (tr.) palpebral area; and diverging, outwardly convex anterior branches of the facial suture. Among these characters, the lateral glabellar furrows are most distinctive. The cranidium of Honanaspis parvus Kuo in Lu et al. (1965, pl. 53, fig. 3), has a subrectangular glabella with four distinct pairs of lateral glabellar furrows, and a narrow palpebral area. However, the previously illustrated cranidium differs from the new specimen left in open nomenclature in having a transverse anterior border defined by a more distinct border furrow, and a glabella that is slightly inwardly concave laterally. Inclusion of P. (H.) parvus in Proasaphiscus (Honanaspis) seems problematic because glabellar furrows in all other species assigned so far to the subgenus are strongly effaced. To accommodate this species and the new crandium left in open nomenclature, the concept of the subgenus needs to be expanded to include forms having strong lateral glabellar furrows. Alternatively, the two forms could be placed in a new proashaphiscid genus. Pending further study, we refer both forms to Proasaphiscus (Honanaspis). Pseudanomocarina? eldachia from Altay-Sayan (Chernysheva in Bognibova et al., 1971, p. 157, pl. 16, figs. 13, 14) is broadly comparable to the cranidium described here. P.? eldachia is • 24.

characterized by a rather well furrowed glabella with a similar pattern of lateral furrows, a narrow palpebral field, an upturned anterior border, and both palpebral lobes and facial sutures that are similar to those of the new cranidium. P. ? eldachia differs only in having a more rounded glabellar front, less distinct lateral glabellar furrows, and three rather than four pairs of lateral glabellar furrows. The cranidia from Altai-Sayan and northwestern Hunan may be closely related, possibly even congeneric.

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Genus ADELOGONUS Opik, 1967

Adelogonus Opik, 1967, p. 203,204. Sinocoosella Yuan and Yin, 1998, p. 143, 144. Plesiocilia Yuan and Yin, 1998, p. 154. Type species. Adelogonus solus Opik, 1967 (p. 204, pl. 8, fig. 3, text-fig. 70), from the Glyptagnostus stolidotus Zone, O'Hara Shale, northwestern Queensland Australia; by original designation. Other species. Sinocoosela typica Yuan and Yin, 1998 (p. 144, pl. 1, figs. 14-16) [=Plesiocilia minuta Yuan and Yin, 1998 (in part), p. 154, 155, pl. 3, figs. 11, 12, non 13, 14] from the Glyptagnostus stolidotus Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou; Adelogonus hunanensis sp. nov.; and Adelgonus sp. (herein). Emended diagnosis. Proasaphiscidae with semicircular cephalon, border strongly upturned, and border furrows shallow and broad. Preocular ridges present or absent; if present, transverse. Glabella tapering broadly forward, with concave sides in anterior half, rounded in front; lateral furrows variably effaced; S 1 bifurcated. Occipital furrow becoming shallow distally, not connected to axial furrow. Preglabellar area about 0.3 of cranidial length. Palpebral area narrow (0.25-0.3 of glabellar width at the level of palpebral lobes); located subcentrally; palpebral lobe narrow, of medium size (about 0.3 of the glabellar length excluding occipital ring). Pygidium semicircular, with outer portion of pleural field broadly concave; borders narrow, strongly upturned. Axis extending to near paradoublural line; with three tings and semicircular terminal piece and wide postaxial ridge. Surface densely granulose or smooth. Remarks. The diagnosis of t3pik (1967, p. 203) is emended here on the basis of new and better preserved material from eastern Guizhou (Yuan and Yin, 1998) and northwestern Hunan. Sinocoosella and Plesiocilia are regarded as junior synonyms of Adelogonus. Both Sinocoosella and Plesiocilia are monospecific and are apparently synonymous. The type species of Sinocoosella, Sinocoosella typica (Yuan and Yin, 1998, p. 144, in part, pl. 1, figs. 15, 16, non 14), and the type species of Plesiocilia, Plesiocilia minuta (Yuan and Yin, 1998, p. 154, 155, pl. 3, figs. 11, 12, ?13, non 14), are from the same collection (WY23F1) in the same section. The holotype and paratype cranidia of P. minuta are small in size. Morphologically, they are identical in all respects with those of S. typica and are clearly immature individuals of that species. Furthermore, Sinocoosella is apparently a junior synonym of Adelogonus. The Chinese and the Australian genera • 25.

are of comparable age, as both occur in the Glyptagnostus stolidotus Zone. There seems to be no significant distinction in morphology between these two taxa, although illustrated specimens of Sinocoosella seem to have deeper glabellar furrows, and seem to lack the paired, shallow convex furrows (i.e., the frontal limits of the cephalic pleural lobes of Opik, 1967, p. 203; text-fig. 70) on the preocular area observed in Adelogonus. These differences are considered to be of no more than species-level significance.

Adelogonus hunanensis sp. nov. Plate 9, figures 1-7 2001b Adelogonuscf. A. typica (Yuan and Yin); Peng, Babcock, and Lin, p. 105, pl. 13, fig. 1.

Etymology. From Hunan Province. Holotype. Cranidium (P1.22, figs. 1-3, 7, NIGP 137979) from collection P331.8. Paratypes. Two pygidia (NIGP 137980, 137981), in collections P286.3 and P331.8. Diagnosis. Adelogonus with glabella having faint lateral furrows; preglabellar field wider (sag.) than anterior border; frontal field bears pair of transverse, slightly convex ridges at extreme anterior end and pair of ovate nodes in front of anterolateral comers of glabella. Pygidium with subconical axis, reaching paradoublural line, with three tings and terminal piece; pleural field outside paradoublural line broadly concave; borders narrow, strongly upturned; pleural furrows distinct proximally, interpleural furrows almost obliterated. Surface covered with dense granules.

Description. Anterior border narrow, convex and upturned; anterior border furrow shallow; preglabellar field gently depressed, wider (sag.) than anterior border. Glabella moderately convex, strongly tapered, slightly concave in anterior half, obtusely rounded anteriorly, beating obscure carina. Axial furrow deep, gently sinuous. Lateral glabellar furrows consist of four pairs; S 1 broad, gently impressed, with abaxial end deepened into shallow cavity; $2, $3, $4 short and gently curved, $2 forward of level of anterior end of palpebral lobe. Occipital furrow shallow and wide, becoming narrower and more shallow distally; occipital ring of uniform width. Fixigena narrow, with palpebral area about one-fourth of glabellar width between palpebral lobes. Preocular field with paired transverse convex ridge at anterior limit. The transverse convex ridge is defined posteriorly by thin, incised furrow that fails to extend into preglabellar field adaxially. Paired subovate node present between proximal end of transverse ridge on preocular field and anterolateral comer of glabella, defined abaxially by short furrow extending exsagittally from anterior end of axial furrow to proximal end of furrow-defined transverse ridge in front of preocular field. Palpebral lobe thin, about one-quarter as long as glabella, defined by moderately deep palpebral furrow. Anterior branch of facial suture arched outward, posterior branch unknown. Posterior border furrow shallow, broad; posterior border consists of linear ridge. Posterolateral projection subtriangular. Pygidium semicircular, width about twice length, with linear flange anteriorly. Axis tapered gently rearward, extending almost to paradoublural line, consisting of bar-like articulating half-ring, three tings defined by shallow, thin ring furrows, a semicircular terminal piece that continues as a wide postaxial ridge and merges with pleural field before reaching posterior border. Pleural area • 26.

with inner part gently convex and outer part broadly concave. Pleural furrows distinct proximally, becoming shallow or faint distally; borders narrow, strongly upturned, defined by shallow or obscure border furrows. Surface covered with fine, densely spaced granules.

Remarks. Adelogonus hunanensis sp. nov. is quite similar to Adelogonus solus Opik (1967, p. 204, pl. 8, fig. 3, text-fig. 70), the type species of Adelogonus, from northwestern Queensland, but differs from A. solus in having a glabella with more sinuous sides and a more transverse front, a wider (sag.) preglabellar field, and a longer (exs.) preocular area that bears a transverse prefixigenal ridge instead of a shallow convex furrow. The new species is also distinct from the type species of the genus in having a pair of ovate nodes between the preglabellar field and the preocular area. Adelogonus typica (Yuan and Yin, 1998, pl. 1, figs. 15, 16; non 14 [=Plesiocilia minuta Yuan and Yin, 1998, pl. 3, figs. 11, 12, non figs. 13, 14]), which is from the same region as the new species, is moderately similar to A. hunanensis. A. typica is distinguished by having an anteriorly rounded glabella with deeper lateral furrows, of which the S1 furrow is bifurcated rather than shallowly impressed, and the $2 furrow is weakly rather than obscurely defined; and by lacking both the paired transverse ridges and the paired ovate nodes on the frontal field. Pygidia previously assigned to A. typica (Yuan and Yin, 1998, pl. 1, fig. 14) and its junior synonym Plesiocilia minuta (Yuan and Yin, 1998, pl. 3, fig. 14) do not appear to be conspecific, nor does either one seem to be conspecific with pygidia assigned here to A. hunanensis. New material of Adelogonus indicates that the pygidia of A. typica and P. minuta illustrated by Yuan and Yin (1998) are probably misassociated. The pygidium assigned to A. typica (Yuan and Yin, 1998, pl. 1, fig. 14) may be conspecific with pygidia that Yang and Yin (1998) assigned to Poriplethopeltis sinensis Yuan and Yin (1998, pl. 5, figs. 12, 13). However, the pygidium assigned to Plesiocilia minuta (Yuan and Yin, 1998, pl. 3, fig. 14) is less than 1 mm in length and too small for a meaningful evaluation at present. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the middle part of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the upper part of the Proagnostus bulbus Zone to the Linguagnostus reconditus Zone). Adelogonus? sp. Plate 31, figures 15, 16

Material. One pygidium (NIGP 138210) in collection P341.8. Remarks. When originally described (Opik, 1967), Adelogonus was known only from the holotype cranidium of A. solus. Solenocoosella typica, which is transferred here to Adelogonus, was described based on two cranidia and an associated pygidium (Yuan and Yin, 1998). The paratype pygidium of A. typica is similar to the pygidium that is illustrated here, but left in open nomenclature. The new pygidium is broken along the anterior margin, and on the fight and posterior sides. The anterior margin is strongly deflected backward. The axis is convex, wide, nearly cylindrical, and formed of at least one distinct axial ring, plus a large subcircular terminal piece and a rather broad postaxial ridge. Weak furrows divide the relatively narrow pleural fields into four segments. A posterior arch seems to have been present. The new pygidium differs from the pygidium of A. typica in having slightly wider pleural areas, and a larger terminal piece on the • 27"

axis. Both the paratype pygidium of A. typica and the new pygidium are certainly congeneric, but both pygidia are notably different from that assigned here to Adelogonus hunanensis sp. nov. (see P1. 9, figs. 4, 5). For this reason, the new pygidium and the paratype pygidium of A. typica are regarded as questionably belonging to Adelogonus.

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the upper part of the Linguagnostus reconditus Zone). Genus EYMEKOPS Resser and Endo, 1937

Type species. Anomocarella hermias Walcott, 1911 (p. 92, pl. 15, fig. 10) from the Crepicephalina Zone, Changhia Formation, Changxingdao Island, Liaoning; by original designation. Remarks. Zhang and Jell (1987, p. 162-165) rediagnosed the genus and reviewed the species previously assigned to the genus. The type species and three other species assigned to Eymekops by Resser and Endo (1937) were considered to be synonymous, and most of the other species were either excluded from the genus or were questionably included in it. Zhang and Jell's (1987) genetic concept is followed here. Eymekops? sp. 1 Plate 10, figures 11-14

Material. One cranidium (NIGP 137994) in collection P319.6. Description. Anterior border narrow, convex, gently arched forward; preglabellar field wide, gently depressed. Cranidium subquadrate, slightly longer than wide. Glabella subrectangular in outline, slightly tapered forward, obtusely rounded anteriorly, with 4 pairs of lateral furrows; S 1 trifurcated, with posterior branch curving rearward to meet occipital furrow, defining ovate lobe distally on L 1; median branch of S 1 longest, straight, oblique rearward; anterior branch of S 1 oblique forward; $2, $3, and $4 short and faint. Palpebral lobe large, moderately curved, with anterior and posterior ends opposite occipital and $4 furrows respectively; palpebral area wider than half glabellar width. Anterior branch of facial suture gently curved; posterior branch enclosing narrow (exs.), transverse posterolateral projection. Surface covered with fine, densely spaced granules. Remarks. This cranidium is questionably assigned to Eymekops because it resembles Eymekops hermias, the type species of Eymekops (see Zhang and Jell, 1987, p. 163, pl. 66, figs. 3-12; pl. 67, figs. 1-8, 10, 11), in most essential respects. Shared characteristics include a large palpebral lobe; wide, short, and gently oblique eye ridges; a subrectangular glabella; a convex, forwardly arched anterior border; and four pairs of lateral glabellar furrows, with S 1 being branched. However, the new cranidium differs from E. hermias in having a much wider (exs.) preglabellar field and in lacking a plectrum on the posterior margin of the anterior border. Also, the rearward-curved posterior branch of S 1, which isolates the distal part of L l, is distinctive. • 28"

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone). Eymekops? sp. 2 Plate 1O, figure 15

Material. One pygidium (NIGP 137995) in collection P337.5. Description. Pygidium transverse-subrectangular in outline, width twice length, beating one pair of rather thin lateral spines, and one pair of posterior spines that are broad at base. Anterior margin transverse, lateral margin straight, parallel to sagittal line; anterolateral comer of pygidium rounded-rectangular, anterolateral facet not evident. Axis unknown, but probably occupying half pygidial length. Pleural region with abaxial part gently convex, sloping gently outward and rearward to posterior margin, without border furrows, divided into about four segments; first pleural furrow deeply incised and transverse initially, deflected rearward and gently outward; other pleural furrows deeply incised and oblique initially, obliterated beyond inner part of pleural region; all interpleural furrows obscure initially, becoming deep and rather broad on outer part of pleural region, nearly confluent with pleural furrows. Surface covered with fine, densely spaced granules. Remarks. This spinose, non-bordered pygidium bears some similarity to the pygidium of Eymekops (see Zhang and Jell, 1987, pl. 66, figs. 3, 6-8, 10, 11; pl. 67, figs. 5, 8 for E. hermias, the type species, and Chemysheva, 1961, pl. 25, figs. 12, 13 for Kolpura oculeata, a possible species of Eymekops). The broad-based posterior spines, the wavy posterior margin, the relatively short axis, and the deeply impressed pleural or interpleural furrows are more or less comparable. The pygidium is distinguished by having a rectangular rather than rounded anterolateral comer, and two rather than three pairs of spines. For those reasons and because of the absence of an associated cranidium it is tentatively referred to Eymekops. It is uncertain whether this pygidium can be associated with the cranidium assigned here as Eymekops? sp. 1 (pl. 10, figs. 11-14), which is from a different but stratigraphically close collection (i.e., from the level of 17.9 m, below), and is smaller in size. If they do belong to a single species, it may represent an undescribed new species of Eymekops or even a new genus related closely to Eymekops. Stratigraphically, Eymekops is known from the Crepicephalus Zone in the North China Platform, having much lower occurrences than both Eymekops? sp. 1 and E. ? sp. 2. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Genus GRANDIOCULUSCossmann, 1908

Megalophthalmus Lorenz, 1906, p. 76 (preoccupied); Kobayashi, 1935 p. 87; 1937, p. 433, 434; Jell and Adrain, 2003, p. 404. non Megalophthalmus Leach, 1830 (not seen, fide Zhang and Jell, 1987, p. 166). non Megalophthalmus Gray, 1832 (not seen, fide Zhang and Jell, 1987, p. 166). Grandioculus Cossmann, 1908, p. 68; Howell in Moore, 1959, p. 290; Kobayashi, 1962, p. 70, 71;Lu, • 29"

Zhang, Zhu, Qian, and Xiang, 1965, p. 337; Zhang and Jell, 1987, p. 166; Jell and Adrain, 2003, p. 380. Honania Lee in Lu et al., 1965, p. 338, 339; Zhou, Liu, Meng, and Sun, 1977, p. 184; Schrank, 1977, p. 152; Zhang, Lin, Wu, and Yuan, 1980, p. 82; Zhou, Li, and Qu, 1982, p. 252, 253; Zhang and Wang, 1985, p. 439; Zhang, Xiang, Liu, and Meng, 1995, p. 61, 62; Jell and Adrain, 2003, p. 385. Type species. Liostracus megalurus Dames (1883, p. 20, 21, pl. l, figs. 7, 8) from the Changhia Formation, Saimaji, eastern Liaoning; designated by Kobayashi (1935, p. 87). Other species. Zhang and Jell (1987) restricted the genus Grandioculus to the type species, Anomocare bigsbyi Walcott (1906, p. 581, 582; 1913, p. 198, pl. 21, figs. 3, 3a, b), and Anomocarella? dingxiangensis Chang (in Lu et al., 1965, p. 327, pl. 60, figs. 20-24). Both G. bigsbyi and G. dingxiangensis are from the Amphoton Zone in the Changhia Formation of Shanxi. To this list, the following species should be added: Grandioculus sp. (Zhang and Jell, 1987, p. 168, pl. 68, fig. 4) from the Changhia Formation at Gaojiapu, north of Xintai, Shandong; Psilaspis suni Resser (1942, p. 45) [=Anomocare flava (Walcott) sensu Sun, 1924, p. 80, 81, pl. 5, figs. 8a-d] from the middle Cambrian near Zhaogezhuang, western Hebei; Honania lata Lee in Lu et al., (1965, p. 383, pl. 63, figs. 3-5), possibly from the Changhia Formation, Dengfeng and Gongxian, northwestern Henan; Honania yinjiensis Mong in Zhou et al., 1977 (p. 185, pl. 54, figs. 12, 13) from the Changhia Formation near Wuyang, central Henan; Honania pansa Zhang and Yuan (in Zhang et al., 1980b, p. 82, pl. 10, fig. 4) from the Changhia Formation near Ruicheng, southern Shanxi; Honania panda Zhang and Yuan (in Zhang et al., 1980b, p. 82, 83, pl. 10, fig. 5) in the same collection as Honania pansa Zhang and Yuan; Honania symmetrica Zhou in Zhou et al., 1982 (p. 253, pl. 63, figs. 9, 10) from the uppermost Hsuchuang Formation near Alxa Zuoqi (Bayan Hot), Inner Mongolia; Honania zhongyangensis Zhang and Wang, 1985 (p. 439, 440, pl. 129, fig. 12) from the Hsuchuang Formation, Cangwan, Zhongyang, western Shanxi; Honania huoshanensis Zhang and Wang, 1985 (p. 440, pl. 129, fig. 13) from the Hsuchuang Formation, Guojiajie, Hongdong, southern Shanxi; Grandioculus truncatus sp. nov.; and G. obscurus sp. nov. Anomocarella irma Walcott (1906, p. 584; 1913, p. 202, 203, pl. 19, figs. 8, 8a) from the upper part of the Changhia Formation near Yantou, Wutai, Shanxi, was transferred questionably to Grandioculus by Zhang and Jell (1987). Honania? yinjiensis Mong in Zhou et al. (1977, p. 186, pl. 54, fig. 11) is a proasaphiscioid but not a species of Grandioculus. The species is based on a single cranidium characterized by having an elongate, parallel-sided, anteriorly truncate glabella; and a wide (sag., exs.) anterior area and narrow (tr.) posterolateral projections of the fixiganae. Remarks. Grandioculus was erected to replace the preoccupied name Megalophthalmus Lorenz (1906, p. 76). Names of that spelling were erected twice before Lorenz (1906) proposed the genetic name again. Zhang and Jell (1987) rediagnosed the genus Grandioculus and considered Honania Lee in Lu et al., 1965 to be a junior synonym of Grandioculus. Subsequently, Zhang (in Zhang et al., 1995, p. 61, 62) revived Honania without comment. In comparing the lectotype of Honania lata (Lu et al., 1965, pl. 63, fig. 3; lectotype designated by Zhang in Zhang et al., 1995, p. 61, 62), the type species of Honania, with the lectotype of Grandioculus megalurus Cossmann, 1908, p. 68 (lectotype designated by Schrank, 1977, pl. 5, fig. 1), the type species of Grandioculus, there seem to be no significant differences in essential characteristics. The lectotype of G. megalurus does differ slightly from that of H. lata in having outwardly convex rather than outwardly slightly concave • 30"

axial furrows and a moderately rounded rather than broadly rounded glabellar front, but these distinguishing features appear to be of only species-level value at most. The pygidia of Grandioculus and Honania are indistinguishable. Here, Honania continues to be suppressed as a junior synonym of Grandioculus (following Zhang and Jell, 1987). All species assigned originally to Honania are here transferred to Grandioculus and listed above. After the erection of Honania, most species assigned to it have been based on single cranidia, most of which are neither preserved nor illustrated satisfactorily, and the specific concepts of these species are more or less obscure. Some of them eventually may be shown to be synonymous. For instance, H. panda and H. pansa are likely to be synonyms. H. zhongyangensis is likewise probably a junior synonym of H. lata. More material is needed to clarify the status of these species. Grandioculus obscurus sp. nov.

Plate 10, figure 1 2001b Anomocarellid gen. et sp. nov., Peng, Babcock, and Lin, p. 101, pl. 2, fig. 9. Etymology. From Latin, obscurus, obscure, referring to the obscure lateral glabellar furrow. Holotype. By monotypy; cranidium (P1. 10, fig. 1, NIGP 137986) from collection P64.5. Diagnosis. Grandioculus with an anteriorly rounded, laterally constricted glabella having obscure lateral furrows; anterior border upturned moderately; preglabellar field short, depressed. Description. Cranidium subquadrate, slightly longer than wide, with rounded anterior margin. Anterior border flat, upturned, defined posteriorly by shallow anterior border furrow. Preglabellar field short (sag.), length about one-third that of anterior border, depressed. Glabella tapering forward gently, sides concave, front rounded, length about five-sixths of cranidium; lateral glabellar furrows nearly effaced. Occipital furrow transverse, wide, moderately impressed, shallowing distally; occipital ring prominent, widest medially, constricted abaxially, with obtusely angular posterior margin distally, probably beating median node at posterior. Eye ridge wide, directed diagonally. Palpebral lobe moderately large, located slightly posterior to midlength of cranidium; palpebral area of fixigena probably half of glabellar width at its midlength. Anterior branch of facial suture diverging gently forward; posterior branch diverging strongly rearward. Posterolateral projection triangular, with deeply incised posterior border furrow and narrow (exs.) convex posterior border; both posterior border furrow and border gently widening abaxially. Surface covered with fine, densely space punctae. Remarks. A single cranidium is considered to represent a new species of Grandioculus. The new species, G. obscurus, which is known only from a cranidium, is similar to G. latus (Lee in Lu et al., 1965, p. 338, pl. 63, figs. 3, 5) except for having a proportionally longer glabella and effaced lateral glabellar furrows. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone).

.31"

Grandioculus truncatus sp. nov.

Plate 11, figures 1-15 2001b Honania sp. cf. H. lata Lee; Peng, Babcock, and Lin, p. 101, pl. 1, fig. 9. 2001b Solenopleurid gen. et sp. nov. (in part), Peng, Babcock, and Lin, p. 101, pl. 1, fig. 11 only. Etymology. From Latin, truncatus, truncated, referring to the truncate-tapering glabella. Holotype. Cranidium (P1. 11, figs. 4, 11, 12, NIGP 137999) from collection P37. Other material. More than 20 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 137996-137998, 138000-138006) in collections P35.5 and P37. Diagnosis. Grandioculus with an anteriorly truncated glabella; broad occipital furrow; wide eye ridge; and narrow palpebral area, which is about one-fourth of glabellar width at L2. Description. Cranidium subquadrate, slightly wider than long. Anterior border gently convex, moderately elevated, anterior border furrow shallow, gently arched forward. Glabella trapezoidal, tapering forward gently; with straight sides and truncate front; lateral glabellar furrows consist of four pairs; S1 long, bifurcated, with transverse anterior branch and diagonally directed posterior branch; $2 moderately long, transverse or slightly oblique rearward; $3 almost as long as $2, slightly oblique forward; $4 short, incised, lying slightly forward of $3, directed inward and forward. Occipital furrow wide and deep, transverse medially or gently sinuous, medial portion arched forward slightly. With distal ends deflecting gently forward; occipital ring wide sagittally, gently narrowing abaxially, beating posteriorly located median node. Preglabellar field shorter (sag.) than anterior border, gently impressed. In large cranidium, anterior border is thin and strongly upturned. Eye ridge thick, short, subparallel to anterior border furrow; palpebral lobe of moderate size, crescentic, slightly oblique, extending from the level of midpoint of L1 to $2. Palpebral area narrow, gently convex, with width about one-fourth that of glabella at L2. Anterior branch of facial suture diverging forward ; posterior branch diverging strongly rearward. Posterolateral projection transverse, triangular; posterior border furrow widening abaxially. Librigena bearing a short, thin, genal spine that is broad at base. Genal field wide, with radiating, anastomosing ridges; lateral border gently convex, and elevated or progressively upturned forward, defined by broad and shallow lateral border furrow. Posterior border narrow (tr.) and wide (exs.). Pygidium semicircular, width about 1.5 times length; with markedly convex pleural field, and subequally convex axis. Axis long, gently tapered rearward, consisting of five tings, with anterior four tings defined clearly, last ring weakly defined, and a crescentic terminal piece extending as a postaxial ridge on posterior border. Pleural field wider than axis, beating 4-5 pleurae. Pleural furrow broad, well impressed, separating narrow anterior and posterior bands of uniform width. Lateral and posterior border moderately wide, horizontal or slightly downsloping. Cranidial, librigenal, and pygidial surfaces covered with fine, densely spaced granules. Remarks. The new species is most similar to Honania lata, but differs mainly in glabellar shape. In H. lata, the glabella is rounded anteriorly and laterally, and is moderately constricted. In addition, H. lata has a narrower (sag., exs.) occipital furrow, and relatively wider palpebral areas. • 32.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Genus HUAYUANASPIS gen. nov.

Etymology. From Huayuan County in northwestern Hunan, plus Latin aspis, shield, carapace. Type species. Huayuanaspis granulosa gen. et sp. nov.; designated herein. Other species. Huayuanaspis perpauca gen. et sp. nov.; Huayuanaspis? sp. Diagnosis. Proasaphiscidae with tapering glabella, truncate anteriorly, beating occipital spine; lateral glabellar furrows consist of four pairs, S 1 bifurcated; anterior border flat, wider (sag.) than preglabellar field; fixigena moderately wide; palpebral lobe of moderate size, located immediately behind midlength of cranidium; librigena with wide genal field and short, stout genal spine. Remarks. The new genus, Huayuanaspis, is classified within the Proasaphiscidae because its morphology resembles in almost all respects that of genera assigned to that family previously. However, inclusion of the new genus in the Proasaphiscidae requires slight expansion of the concept of the family because genera previously referred to the family all lack an occipital spine (Zhang, 1963). Among the proasaphiscids, some species of Proasaphiscus and Honania that have similarly shaped glabellas and similar patterns of lateral glabellar furrows are most comparable to species included here within Huayuanaspis. For example, Proasaphiscus termieri (Mansuy, 1916, p. 26, 27, pl. 4, fig. 1) and Grandioculus truncata sp. nov. (pl. 11, figs. 1-15) agree with the new genus in having a truncate-tapering glabella, four pairs of lateral furrows, of which S 1 is bifurcated and with a long, diagonally directed posterior branch. Except for having an occipital spine, the new genus differs from Proasaphiscus in having a wide (sag.) cranidial frontal area, smaller palpebral lobes, and a relatively broad cranidial outline. Honania is distinguished from Huayuanaspis by having a shorter frontal area, lacking an occipital spine, having a strongly upturned anterior border, and having more anteriorly located palpebral lobes. Zhaishania Qiu in Qiu et al. (1983, p.144) bears some resemblance to the new genus. Except for lacking an occipital spine, Zhaishania is differentiated from Huayuanaspis by having less diverging anterior branches of the facial suture that meet the anterior cranidial margin more close to the sagittal line, less curved palpebral lobes, and shallower palpebral furrows. Asteromajia Nan and Chang, 1982 (p. 33) and Jiangsuaspis Lin in Qiu et al., 1983 (p. 149) are also similar to Huayuanaspis. These two genera are monospecific, and characterized by having coarsely granulate cranidia with tapered glabellas. They may be synonymous, although it is not certain whether Asteromajia lacks an occipital spine. Both genera can be differentiated from Huayuanaspis by having a more or less rounded glabellar front and a more convex anterior border. In these two genera, the eye ridges are almost obliterated and the ocular ridges are almost completely effaced, so the palpebral areas appear to merge with the palpebral lobes. Jiangsuaspis is further differentiated by the absence of an occipital spine. Yuan et al. (1999, p. 23; 2002, p. 209) suppressed Asteromajia as a junior synonym of Temnoura Resser and Endo in Kobayashi, 1935. Their proposal is rejected here. Yuan et al. (1999, pl. 2, figs. 10, 11; 2002, pl. 62, figs. 1, 2) figured two complete exoskeletons as Temnoura mesembrina Yuan and Zhao, which bear a Temnoura-like pygidium but have a cranidium differing • 33.

greatly from that of Asteromajia. In T. mesembrina, the cranidium has a non-granulate surface, the glabellar flanks are straight rather than inward-constricted, the anterior cranidial border is flat rather than notably convex, the glabellar furrows are strongly impressed rather than largely effaced, and the posterior area of the fixigena is proportionally much wider (tr.) than that of Asteromajia. Until more material is available for Temnoura and Asteromajia, they are regarded as separate genera. The presence of an occipital spine, the glabellar shape, and the nature of the frontal area in Huayuanaspis are reminiscent of Idahoia Walcott, 1924 or, according to Ludvigsen and Westrop (1983, p. 23), its senior synonym Saratogia Walcott, 1916. However, the North American genus differs in having a much narrower fixigena and a variably effaced glabella; the S 1 furrow, if present, is not bifurcated in Idahoia or Saratogia.

Huayuanaspis granulosa gen. et sp. nov. Plate 12, figures l- 12 2001b Solenopleurid gen. et sp. nov. (in part), Peng, Babcock, and Lin, p. 101, pl. l, fig. 10 only. 2001b Undetermined cranidium, Peng, Babcock, and Lin, p. 102, pl. 4, fig. 11.

Etymology. From Latin, granulosus, granulate, referring to the granulate ornamentation. Holotype. Cranidium (P1. 12, figs. 2-4, NIGP 138008) from collection P37. Paratypes. Five cranidia and three librigenae (illustrated specimens NIGP 138007, 138009-138016) in collections P19.8, P29, P35.5, P37, P38.5, P45.6 and ?P138.8.

Diagnosis. Huayuanaspis with distinct lateral glabellar furrows; cephalic surface covered with both fine, densely spaced granules and coarse, scattered granules, or with only coarse, closely spaced granules on cranidial surface.

Description. Cranidium subquadrate with width between palpebral lobes subequal to length, excluding occipital spine. Anterior border flat or gently convex, horizontal or slightly upturned, defined posteriorly by slope change and shallow anterior border furrow that is gently arched forward. Frontal area occupying 0.25-0.28 cranidial length excluding occipital spine, with anterior border wider than preglabellar field. Glabella evenly tapering forward, truncate to obtusely rounded anteriorly, with four pairs of lateral furrows, defined by deep axial and preglabellar furrows; S 1 bifurcated; anterior branch of S 1 shallow, transverse or inward and slightly forwardly directed; posterior branch of S 1 long, straight or somewhat sinuous, well impressed, extending inward and rearward from axial furrow to nearly meet occipital furrow, defining subtriangular L1; $2 short, clearly defined, slightly curved, inwardy and slightly rearward directed, lying forward of midpoint of glabella excluding occipital ring; $3 shorter than $2, inwardly and forwardly directed, lying about midway between $2 and preglabellar furrow; $4 weak, shorter than $3, lying about midway between $3 and preglabellar furrow. Occipital furrow wide and deep, transverse medially, deflected forward distally; occipital ring widest sagittally, narrowing abaxially, extending into moderately long spine that is slightly elevated toward rear. Eye ridge convex and clearly defined, extending subparallel to anterior border furrow from anterior end of palpebral lobe into axial furrow at level of $3. Palpebral lobe of moderate size, crescentic in shape, defined by deep, strongly arcuate palpebral furrow, lying immediately behind cranidial midlength. Palpebral area moderately convex, • 34.

about half as wide as glabellar width at L2, with inner part inclined steeply to axial furrow. Anterior branch of facial suture diverging forward gently, becoming straight until meeting anterior border furrow, then turning rather sharply, converging forward to cut anterior border and meet cephalic anterior margin at point opposite distal end of occipital furrow; posterior branch nearly transverse, enclosing blade-like posterolateral projection. Librigena wide, with moderately wide lateral and posterior borders; lateral and posterior border furrows broad and shallow, converging at genal angle and extending into base of genal spine; genal spine horn-like with distal end curving inward. Doublure narrow and convex, cut anteriorly by straight, obliquely directed connective suture. Cranidial and librigenal surfaces with fine, densely spaced granules and coarse scattered granules, or with only coarse, closely spaced granulation.

Remarks. Huayuanaspis granulosa sp. nov., the type species of the genus, is represented by the most abundant and best preserved specimens assigned to any of the three species embraced by Huayuanaspis. H. granulosa is readily differentiated from the other two species of Huayuanaspis by the presence of coarser, more densely spaced granules on the cephalic surface. The granules are fine and sparsely distributed over the exoskeleton of H. perpauca sp. nov., and they are absent from specimens assigned to Huayuanaspis? sp. H. granulosa also differs from H. perpauca in having a proportionally larger glabeUa with a more rounded front, and it differs from Huayuanaspis? sp. in having more clearly defined lateral glabellar furrows.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone to the Pianaspis sinensis Zone (equivalent to the Ptychagnostus atavus Zone through the lowermost part of the Lejopyge laevigata Zone). Huayuanaspis perpauca gen. et sp. nov. Plate 13, figures 1-10

Etymology. From Latin, perpaucus, extremely few, referring to having few granules on the fixigena.

Holotype. Mostly exfoliated cranidium (P1.13, figs. 4, 5, NIGP 138018) from collection P45.6. Paratypes. Six cranidia (NIGP 138017, 138019-138023) in collection P45.6. Diagnosis. Huayuanaspis with lateral glabellar furrows weakly impressed, anterior area of fixigena with anastomosing lines; fixigena with few, widely scattered, fine granules.

Description. Cranidium subquadrate, width between palpebral lobes subequal to length excluding occipital spine. Anterior border gently convex, slightly upturned and slightly elevated above preglabellar field; preglabellar field shorter (sag.) than anterior border. Glabella trapezoidal, tapering gently forward, truncate anteriorly, occupying three-fourths total cranidial length, defined by broad and moderately deep axial and preglabellar furrows; with four pairs of lateral furrows; S 1 weak, bifurcate, with long, posterior branch directed strongly obliquely rearward and anterior branch short, directed forward and inward; $2-$4 short, faint, directed forward and inward. -35.

Occipital furrow transverse, moderately wide, deflected slightly forward distally; occipital ring widest medially, narrowing abaxially, probably extending rearward into long, stout spine. Eye ridge weak, directed diagonally. Palpebral lobe moderately large, crescentic, defined by deep, strongly arcuate palpebral furrow, lying immediately behind cranidial midlength; palpebral area about half as wide as glabellar width at L2, gently convex, steeply inclined to axial furrow. Anterior branch of facial suture diverging forward gently, converging forward after crossing anterior border furrow to cut anterior border and meet cephalic anterior margin at point opposite inner end of eye ridge; posterior branch straight, diverging strongly rearward; posterolateral projection triangular in shape, short (exs.), relatively narrow (tr.), beating wide and moderately deep posterior border furrow.

Remarks. This new species is quite similar in morphology to the type species, Huayuanaspis granulosa, but differs in having a rather effaced glabella, anastomosing lines in the frontal field of the cranidium, and almost no granulation other than small granules on the posterior area of the fixigenae, or occasionally on the preocular areas. In a small, mostly exfoliated cranidium, scattered granules are present on the glabella. The anterior border in the new species is gently convex, and the palpebral area and posterior area of the fixigena is inclined rather steeply to the axial furrow. Exfoliated surfaces commonly show dense punctae. The occipital spine is broken in all known specimens but its presence is clearly indicated by a strong spine base on the posterior margin of the occipital ring.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Huayuanaspis? sp. Plate 13, figure 11 2001b Undetermined cranidium, Peng, Babcock, and Lin, p. 102, pl. 4, fig. 14.

Material. A single, mostly exfoliated cranidium (NIGP 138024) in collection P126. Remarks. A single broken cranidium left in nomenclature has a truncate-tapering glabella with four pairs of lateral glabellar furrows. The S 1 furrow is bifurcate, and the $3 and $4 furrows are closely spaced near the anterior comer of the glabella. Most of the surface is smooth, but the posterior part of the palpebral area bears fine, densely spaced granules. The outer part of the frontal field shows obscure anastomosing lines, and the anterior part of the anterior border has punctae. The single known cranidium is similar to Huayuanaspis perpauca gen. et sp. nov., but differs in having a less tapered glabella, a thinner exoskeleton, a narrower palpebral area, and fainter and more oblique eye ridges. The morphology of this specimen suggests that its most likely affinity is with Huayuanaspis, and it probably represents an undescfibed species of the genus. However, because the occipital ring is mostly broken away, and the presence of an occipital spine is uncertain, assignment of the specimen to Huayuanaspis is uncertain.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the lower part of the Goniagnostus nathorsti Zone). • 36.

Genus INIOTOMA Opik, 1967 Type species. Iniotoma iniotoma t3pik, 1967 (p. 231,232, pl. 11, figs. 1-3, text-fig. 81) from the Mungerebar Limestone, northwestern Queensland; by original designation. Other species. Two new species, Iniotoma laevis sp. nov. and Iniotoma porosus sp. nov. Remarks. The genetic diagnosis provided by Opik (1967) is emended slightly to include species having the occipital furrows either ending distally at the lobules or extending to the axial furrows. Iniotoma laevis sp. nov. Plate 9, figures 8-11 2001b Adelogonus? sp.; Peng, Babcock, and Lin, p. 105, pl. 14, fig. 11. Etymology. From Latin, laevis, smooth, referring to the smooth cranidial surface. Holotype. Cranidium (P1.9, figs. 8-11, NIGP 137983) from collection W277. Paratype. A small cranidium (NIGP 137982) in collection W277. Diagnosis. Iniotoma with glabella markedly inflated, tapering strongly forward, rounded anteriorly. Surface smooth. Remarks. Iniotoma laevis sp. nov. closely resembles Iniotoma laevis, the type species of Iniotoma. The narrow, strongly arched anterior border, the glabellar shape, and the course of the facial sutures are comparable between the two species. However, in the new species, the glabella is more strongly tapered forward and more convex, resulting in an inflated appearance; the palpebral area is much narrower; and the ends of the occipital furrow are connected to the axial furrows rather than being closed by lobules. The cranidium of L laevis sp. nov. superficially resembles cranidia referred to Blountia montanensis (Rasetti, 1965, pl. 9, figs. 13, 20) from the Crepicephalus Zone of Tennessee, USA. The specimens from Tennessee, however, can be easily distinguished by the absence of an occipital furrow. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone).

• 37.

Iniotoma porosus sp. nov.

Plate 14, figures 12-16 Etymology. From Latin, porosus, rich in pores, referring to the punctate cranidial and pygidial surfaces. Holotype. Pygidium (P1. 14, figs. 13, 16, NIGP 138030) from collection P298.4. Paratype. An incomplete cranidium (NIGP 138031) in collection P341.8. Diagnosis. Iniotoma with cranidial and pygidial surfaces densely punctuate; glabella with three pairs of weakly impressed lateral furrows; occipital ring with bluntly angled posterior margin. Pygidium with axis having weak ring furrows. Description. Cranidium subquadrate in outline. Glabella tapered forward gently, with straight sides, evenly rounded in front; S 1, $2, and $3 lateral furrows faintly impressed. Occipital furrows broadly bowed rearward; occipital ring subtriagular, narrowing abaxially, elevated posteriorly, with posterior margin angled bluntly. Palpebral lobe wide, curved gently, located subcentrally, defined by incised palpebral furrow; palpebral field one-fourth as wide as glabella; eye ridge obscure, strongly oblique. Posterior branch of facial suture directed diagonally; anterior branch unknown. Pygidium subsemicircular, width twice length. Axis occupies about three-fourths pygidial length, with linear articulating half-ring; weakly segmented, with possibly four tings and poorly defined terminal piece. Pleural field with inner part gently convex, outer part broadly impressed; pleural furrows shallow; interpleural furrows obliterated. Surfaces mostly covered with fine, densely spaced punctae. Remarks. A punctate cranidium and a punctate pygidium from the Huaqiao Formation of northwestern Hunan are inferred to be conspecific based on the appearance of the surface prosopon although the specimens are from separate, but stratigraphically close, collections. The frontal area of the cranidium is partly broken, but other observed cranidial features are similar to those of Iniotoma iniotoma, the type species of Iniotoma. These features include a glabella that tapers forward gently and that is raised above the level of the fixigenae; bifurcate S 1 furrows; a narrow anterior border and a relatively gently sloping preglabellar field; a narrow fixigena; a thin and slightly curved palpebral lobe; a triangular occipital ring; and a narrow (tr.), triangular posterolateral projection. Except for beating punctate prosopon, the new species can be differentiated from the type species by its more transverse glabellar front, the posteriorly more strongly angled occipital ring, and the less effaced glabellar furrows. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone).

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Genus

MAOTUNIA

Zhang and Jell, 1987

Type species. Maotunia semiplectra Zhang and Jell, 1987 (p. 157, pl. 82, figs. 1-4; pl. 83, fig. 1; pl. 84, fig. 1), from the Bailiella-Lioparia Zone, Hsuchuang Formation, near Maotun, Fuzhouwan, Liaoning; by original designation. Other species. Mapania? liaotungensis Resser and Endo, 1937 (p. 252-253, pl. 34, figs. 23, 24) from the Changhia Formation at Jinjiachengzi, Fuzhouwan, Liaoning; Anomocarella distincta Resser and Endo, 1937 (in part, p. 168, pl. 34, figs. 7, 9, non pl. 32, fig. 21) from Crepicephalina Zone, Changhia Formation, Changxingdao Island, Liaoning; Anomocarella blackwelderi Resser and Endo, 1937 (p. 170, 171, pl. 33, fig. 10; pl. 34, figs. 3a-d, 4) from Amphoton Zone, Changhia Formation, Changxingdao Island, Liaoning; Mapaniidae gen. nov. and sp. nov. (sensu Opik, 1967, p. 304, pl. 23, fig. 2) from the middle-upper Cambrian Passage Zone in northwestem Queensland. Four additional species Maotunia sp. cf. M. blackwelderi (Resser and Endo, 1937), Maotunia sp. cf. M. semiplectra Zhang and Jell, 1987, M. rectangularis sp. nov., and Maotunia sp., all from the basal part of the Huaqiao Formation, northwestem Hunan, are described herein. Diagnosis. See Zhang and Jell (1987, p. 156). Remarks. The genetic concept of Zhang and Jell (1987) is followed here. Zhang and Jell (1987) classified Maotunia provisionally within the Proasaphiscidae. Morphologically it is similar to proasaphiscids (e.g., Mapania) in overall features, especially in the shape and arrangement of the lateral glabellar furrows, and the presence of a plectrum. Zhang and Jell (1987) differentiated Maotunia from Mapania in the shape of the cranidial outline, the degree of development of the plectrum, and the proportions of the preglabellar field and the glabella. However, it is not clear that the two genera are actually distinguishable on the basis of glabellar proportion. Mapania seems to have an important role in trilobite phylogeny. Because of a close morphologic similarity between Maotunia and Mapania, Zhang and Jell (1987, p. 156, 188-190) viewed Maotunia as ancestral to both Mapania and the Anomocarellidae during the late Taijiangian and early Wangcunian [=early and middle Changhian age in North China]. Mapania jiangnanensis sp. nov., from the Ptychagnostus atavus Zone (late Taijiangian) in northwestem Hunan, shows morphologic characters intermediate between Maotunia and Mapania, and provides further support for the inferred phylogenetic relationship between these taxa. An alternative interpretation of the relationship between Mapania and Maotunia was provided by Opik (1961, p. 166), who drew attention to the similarity in cranidial morphology between Mapania and Proceratopyge. Proceratopyge and the ceratopygid lineage probably were derived from Mapania (Zhang and Jell, 1987, p. 187, 188; Yuan and Yin, 1999, p. 169), or from the Anomocarellidae (Fortey and Chatterton, 1988), and Maotunia may have given rise to members of the Ceratopygidae through the mapaniids. Maotunia sp. cf. M. blackwelderi (Resser and Endo, 1937) Plate 10, figures 3, 4 cf. 1913 cf. 1937

Anomocarella chinensis Walcott (in part), p. 200, 201, pl. 20, fig. 4a only. Anomocarella blackwelderi Resser and Endo, p. 170, 171, pl. 33, fig. 10; pl. 34, figs. 10-19. • 39-

cf. 1965 cf. 1987

Anomocarella (?) blackwelderi Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 326, 327, pl. 60, figs. 15-19. Moutunia blackwelderi (Resser and Endo); Zhang and Jell, p. 160, pl. 76, figs. 8-13; pl. 70, fig. 2.

Lectotype of Maotunia blackwelderi. Exoskeleton (Walcott, 1913, pl. 20, fig. 4a, USNM 58210; refigured by Reser and Endo, 1937, pl. 33, fig. 10 and by Zhang and Jell, 1987, pl. 77, fig. 2), from the Amphoton Zone, Changhia Formation, Changxingdao Island, Liaoning; designated by Zhang and Jell, 1987 (p. 160). Material. Two cranidia (NIGP 137988, 137989) in collections P7.25 and W 1.2. Description. Cranidium subquadrate, arched forward anteriorly, length subequal to width between palpebral lobes. Anterior border gently convex, arched forward; plectrum short, separating deep border furrow into two parts, not extending onto glabellar front; preglabellar field narrow, slightly convex, subequal in length with anterior border. Glabella subrectangular, straight-sided, or tapering forward slightly, obtusely rounded anteriorly; lateral furrows largely effaced; S 1 isolated from axial furrow, bifurcate, with posterior branch apostrophe-like, and anterior branch transverse; other furrows obscure. Occipital furrow shallow, transverse; occipital ring narrowing abaxially, with prominent node close to posterior margin. Palpebral lobe moderately long, strongly arcuate; eye ridge weak, directed diagonally. Palpebral area nearly half as wide as glabella, gently convex; palpebral furrow shallow medially, deepened slightly at anterior and posterior. Anterior branch of facial suture diverging strongly forward at angle of about 60 ° to sagittal line; posterior branch nearly transverse, slightly convex forward, enclosing blade-like posterolateral projection; posterior border straight. Remarks. Zhang and Jell (1987) illustrated morphologic variation in Maotunia blackwelderi, a species that is based on specimens preserved in shale. The holotype (Walcott, 1913, pl. 20, fig. 4a; Endo and Resser, 1937, pl. 33, fig. 10; Zhang and Jell, 1987, pl. 77, fig. 2) is an internal mold of the exoskeleton; it shows a truncate-conical glabella. Other material indicates that the glabella varies from parallel-sided to gently tapered forward and the glabellar front is commonly obtusely rounded. The holotype also shows the morphology of the doublure. The cephalic doublure bears a connective suture that extends inward and rearward and defines a rather wide (tr., exs.) rostral plate that probably bears a rearward-bowed posterior margin. This species is identical to Maotunia blackwelderi in most important respects, especially the relatively large, arcuate palpebral lobes, the subrectangular glabella with an obtusely rounded front, the transverse occipital furrow, the short preglabellar field, the nature of the fixigenae, and the course of the facial sutures. It can be differentiated from M. blackwelderi by the more strongly curved anterior border, and the location of a point of the facial suture, which is present at a greater distance from the sagittal line than it is in M. blackwelderi. The new cranidia from northwestern Hunan may represent an undescribed species of Maotunia, but until more material becomes available, the specimens are left in open nomenclature. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone).

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Maotunia sp. cf. M. semiplectra Zhang and Jell, 1987 Plate 18, figure 10 cf. 1987 Maotunia semiplectra Zhang and Jell, p. 157, pl. 82, figs. 1-4; pl. 83, fig. 1; pl. 84, fig. 1.

Holotype of Maotunia semiplectra. Exoskeleton (Zhang and Jell, 1987, pl. 82, figs. 1, 2, NIGP 76411), Crepicephalina Zone, Changhia Formation, Maotun, Fuxian, Liaoning. Material. One cranidium (NIGP 138070) in collection W0. Remarks. The single cranidium left in open nomenclature has a subconical glabella with rounded front and straight lateral margins; a weakly convex, poorly defined, forwardly arched anterior border; and narrow (exs.) and long (tr.) posterolateral projections. The exfoliated surface shows that the anterior border is narrow (sag., exs.) and distinctly convex with a wide and short, obscurely defined subtriangular area behind; this was regarded as a primitive plectrum by Zhang and Jell (1987). The specimen is closely comparable to the exfoliated cranidia referred to Maotunia semiplectra, the type species of Maotunia (Zhang and Jell, 1987, pl. 82, figs. 1-3, pl. 83, fig. 1), differing only in having an arched rather than transverse anterior border. The new cranidium is also comparable to Sudanomocarina? huananensis sp. nov. (P1.4, figs. 9-17), but S.? hunanensis has a more clearly defined anterior border and a rearward-bowed rather than transverse occipital furrow. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the unnamed zone (probably equivalent to the Ptychagnostus gibbus Zone). Maotunia rectangularis sp. nov. Plate 15, figures 1-16; Plate 16, figures 1-12

Etymology. From Latin, rectangularis, rectangular, referring to the rectangular shape of glabella. Holotype. Cranidium, mostly exfoliated (P1. 16, figs. 2-4, NIGP 138042) from collection W0. Paratypes. Eighteen cranidia, three librigena, and 10 pygidia (illustrated specimens NIGP 138032138041, 138043-138049), all in collection W0. Diagnosis. Maotunia species with proportionally short, rectangular glabella; with 4 pairs of weak lateral furrows; preglabellar field short; palpebral lobe of medium size; anterior branch of facial suture diverging strongly; pygidium with axis occupying about two-thirds of pygidial length. Description. Cranidium subquadrate, length subequal to width of anterior area of fixigena. Anterior border rather long, fiat, moderately deflected, beating obscure, short plectrum; preglabellar field short to nearly obliterate. Glabella rectangular, parallel-sided, obtusely rounded anteriorly, with weak carina; lateral furrows consist of four weak pairs; S 1 long, bifurcate; $2 as long as S 1, oblique -41-

rearward, slightly curved, lying anterior to glabellar midlength (excluding occipital ring); $3 short, isolated from axial furrow; $4 slightly longer than $3, oblique forward. Occipital furrow incised, transverse; occipital ring slightly narrowing abaxially, with median node. Palpebral area gently convex, length about one-third glabellar width; palpebral lobe of medium size, gently arcuate, extending between midpoint of L 1 and $2; eye ridge weak. Anterior branch of facial suture straight between palpebral lobe and anterior border furrow, diverging diagonally forward, turning sharply inward after crossing anterior border furrow and cutting anterior border as gentle curve; posterior branch gently sinuous, cutting posterior margin close to genal angle, enclosing blade-like posterolateral projection. Lateral border furrow shallow; posterior border furrow moderately deep; posterior border as wide (tr.) as glabella, uniform in width (exs.); lateral and posterior border furrows confluent at genal angle and extending rearward into the base of genal spine as a broad and shallow furrow. Librigena moderately wide, with moderately convex genal field, moderately wide lateral border, and genal spine with broad base. Pygidium elliptical, width nearly twice length. Axis occupying 0.75-0.67 sagittal length; with 5-6 tings, short terminal piece, and postaxial ridge that is distinct, extending beyond paradoublural line but not to posterior margin. Pleural field with inner part gently convex, outer part downsloping strongly; pleural furrows distinct, interpleural furrows weak; pleural and interpleural furrows curved gently; anterior and posterior bands narrow, nearly equal in width. Border narrow, poorly defined. Cephalon and pygidium with thin shell; external surface smooth.

Remarks. In having a short preglabellar field and poorly developed plectrum, Maotunia rectangularis sp. nov. resembles Maotunia semiplectra Zhang and Jell, the type species of Maotunia. The latter differs, however, in having a tapered glabella, a relatively narrower (sag.) anterior border, less diverging anterior branches of the facial suture and a pygidium with longer axis. A cranidium attributed to Maotunia blackwelderi (Resser and Endo, 1937, pl. 34, fig. 18; also Zhang and Jell, 1987, pl. 76, fig. 13) shows similarity to M. rectangularis in the pattern of the lateral glabellar furrows and the configuration of the anterior branches of the facial suture. Other specimens referred to this species show similarly wide and deflected anterior borders. M. blackwelderi, however, can be easily differentiated from the new species by having longer and more arcuate palpebral lobes, a wider (sag.) preglabellar field, and a tapered rather than parallelsided glabella. The new species is also similar to Mapania jiangnanensis sp. nov., which has a somewhat higher occurrence than Maotunia rectangularis in the Paibi-Wangcun area. M. rectangularis, however, differs from M. jiangnanensis in having a rearward-bowed anterior border, a proportionally longer glabella with deeply incised rather than weak lateral furrows, and a long and more clearly defined plectrum. In Mapania jiangnanensis, the palpebral areas of the fixigenae are wider (tr.), the palpebral lobes are longer, and the anterior branch of the facial suture between the anterior end of the palpebral lobe and the anterior border furrow is not straight but outwardly arched.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the unnamed Zone (probably equivalent to the Ptychagnostus gibbus Zone).

-42.

Maotunia sp. Plate 18, figure 13 Material. One incomplete pygidium (NIGP 138072) in collection W3. Remarks. This single pygidium from the Huaqiao Formation, northwestern Hunan, is almost indistinguishable from the pygidium referred to Ptychoparia kochibei Walcott, 1911 by Resser and Endo (1937, pl. 32, fig. 9) from the Crepicephalina Zone, Changhia Formation, Changxingdao Island, Liaoning, Northeast China. Resser and Endo's (1937) pygidium was subsequently transferred to Maotunia distincta (Resser and Endo) by Zhang and Jell (1987, pl. 83, fig. 3). The new pygidium may be conspecific with that pygidium. Until more material belonging to this species becomes is available, we prefer to leave the new pygidium in open nomenclature. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Genus

SUDANOMOCARINA

Jell in Jell and Robison, 1978

Sudanomocarina Jell in Jell and Robison, 1978; p. 14; 15; Zhang and Jell, 1987, p. 168. Type species. Sudanomocarina changi Jell in Jell and Robison, 1978 (p. 15, 16, pl. 4, figs. 9-16) from the Peronopsis opimus Zone, Thomtonia area, northwestern Queensland, Australia; by original designation. Other species. See Jell in Jell and Robison, 1978, p. 14. To this list, Sudanomocarina traynorae Zhang and Jell, 1987 (p. 168, 169, pl. 77, figs. 6a, b), from the Crepicephalina Zone, Changhia Formation, Changxingdao Island, eastern Liaoning, and three additional species, Sudanomocarina sp. cf. S. changi Jell, Sudanomocarina? huananensis sp. nov., and Sudanomocarina sp., described below from the Huaqiao Formation, northwestern Hunan should be added. Remarks. Jell (in Jell and Robison, 1978, p. 16) erected Sudanomocarina to replace Pseudanomocarina Chemysheva in Chemysheva et al., 1956 by transferring the holotype cranidium of Pseudanomocarina plana Chemysheva, the type species of Pseudanomocarina, to Chondranomocare Chemysheva in Chemysheva et al., 1956, and by transferring all other species of Pseudanomocarina to Sudanomocarina. Recently, Jell and Adrian (2003) argued that Chondranomocare, Pseudanomocarina, and Sudanomocarina, are all valid genera. This proposal is tentatively accepted here, although some species assigned to these genera seem to have morphologies that are intermediate among the genetic concepts, and ultimately, the genera might eventually prove to be synonymous. Both Pseudanomocarina and Chondranomocare were erected simultaneously, with Pseudanomocarina having been described three pages earlier than Chondranomocare. If these two genera are synonymous, then Chondranomocare rather than Pseudanomocarina should be suppressed. For the moment, the concept of Pseudanomocarina can be restricted to its holotype cranidium and probably the paratype librigena of P. plana. Specimens subsequently referred to P. plana (Chemysheva, • 43 •

1961, pl. 22, figs. 1-10) and the species subsequently assigned to the genus by various authors (see Jell in Robison, 1978, p. 14) should be transferred to Sudanomocarina. Sudanomocarina, as defined by Jell (in Jell and Robison, 1978), may be differentiated from Pseudanomocarina by having a shorter and smaller anterior area; a shorter, somewhat convex rather than flat anterior border; a less tapered, anteriorly less rounded glabella; and probably longer palpebral lobes. Palmer and Gatehouse (1972) argued that a concave, rather than flat or convex, anterior border characterizes Chondranomocare.

Sudanomocarina traynorae Zhang and Jell, 1987 Plate 19, figure 16 1987 Sudanomocarinatraynorae Zhang and Jell, p. 168, 169, pl. 77, figs. 6a, b.

Holotype. By monotypy; cranidium (Zhang and Jell, 1987, pl. 77, figs. 6a, b, USNM 258612), from the Crepicephalina Zone, Changhia Formation, Changxindao Island, Liaoning. Material. Single cranidium (NIGP 138086) in collection P48.5. Remarks. A cranidium from Paibi, Hunan, agrees in almost all respects with the holotype cranidium from the Crepicephalina Zone in the Changhia Formation, Changxingdao Island, Liaoning. Key characters include a subrectangular, rather convex glabella with a nearly transverse front, defined by a well impressed occipital furrow; a short, weakly convex anterior border; moderately arcuate palpebral lobes located slightly posterior to the midlength of the glabella; a narrow palpebral area (about one-fourth as wide as the glabella); and a narrow (tr.), subtriangular posterolateral projection of the fixigena. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Sudanomocarina sp. cf. S. changi Jell in Jell and Robison, 1978 Plate 17, figures 1-17; Plate 18, figures 1-9; Plate 78, figures 6-8 cf. 1961 Pseudanomocarinaplana Chernysheva (in part), p. 188-191, pl. 22, figs. 3-8, ?figs. 2, 9, 10. cf. 1971 Pseudanomocarina plana Chernysheva; Bognibova, Koptev, Mikhailova, Poletaeva, Romanenko, Semashko, Tomashpoliskaya, Fedjanina, and Chernysheva, 1971, p. 153, 154, pl. 16, figs. 9, 15. cf. 1978 Sudanomocarinachangi Jell in Jell and Robison, p. 15, 16, pl. 4, figs. 9-16. 2001b Sudanomocarina sp., Peng, Babcock, and Lin (in part), p. 101, pl. 1, fig. 7, non figs. 8, 13 [=Parayujinia constricta gen. et sp. nov.].

Holotype ofSudanomocarina changi. Cranidium (Jell and Robison, 1978, p. 4, fig. 14, UQF 69375), from the Peronopsis opimus Zone, near Thorntonia, Northwestern Queensland, Australia. Material. More than 20 sclerites, including cranidia, librigenae, pygidia (illustrated specimens •

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NIGP 138050-138069, 138682) in collections P3.2, P12.5, P29, P33.5, P37, P38.5, W0, W1.2, W33, and W36.6.

Description. Anterior border slightly convex, upturned, much wider than preglabellar field, narrowing abaxially; anterior border furrow weak. Glabella subrectangular, gently convex, tapering forward slightly, broadly or obtusely rounded anteriorly with rounded anterolateral comers; lateral furrows consist of 3-4 faint to almost completely effaced pairs; S 1, $2, and $3 rather long, directed inward and slightly rearward; $4 short, directed inward and forward. Occipital furrow shallow, slightly sinuous; occipital ring crescentic, with small, median node; posterior margin of occipital ring arched rearward. Palpebral lobe about 0.45 glabellar length (excluding occipital ring), gently arcuate and slanted, located posterior to cranidial midlength, defined by weak palpebral furrow; palpebral area crescentic, gently convex, about one-fourth width of glabella. Anterior branch of facial suture diverging forward 250-30 ° to sagittal line, deflecting inward and forward after crossing anterior border furrow; posterior branch gently sigmoidal, enclosing narrow (exs.), wide, and long (tr.) blade-like posterolateral projection. Librigena with wide, gently convex genal field, moderately wide border and border furrow and genal spine that is broad at base; border furrow shallow but distinct, not extending to genal angle. Pygidium subelliptical; lateral margin angled rather obtusely; posterior margin broadly rounded; axis long, reaching close to posterior border furrow, with 4-5 tings. Pleural field gently convex; distal part downsloping slightly; pleural furrows gently curved, clearly impressed; interpleural furrows obscure. Posterior border furrow obscure. Doublure moderately wide. Remarks. Cranidia of this species left in open nomenclature from the Huaqiao Formation of northwestern Hunan bear close similarity to both cranidia of Sudanomocarina changi from Australia (Jell in Jell and Robison, 1978), and most cranidia assigned to Pseudanomocarina plana (other than the holotype) from the Pseudanomocarina Zone (Amgan Stage) of Siberia (Chernysheva, 1961). Minor differences are present in both the cranidia and pygidia. The cranidium from Hunan has weakly sinuous rather than straight axial furrows, a wider anterior border, and probably greater convexity than the specimens from Australia or Siberia. The pygidium has a more slender axis, a relatively wider pleural area, and angled rather rounded lateral margins as compared to the Australian and Siberian specimens. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the unnamed zone and the Dorypyge richthofeni Zone (probably equivalent of the Ptychagnostus gibbus Zone to the lower part of the Ptychagnostus atavus Zone). Sudanomocarina? huananensis sp. nov. Plate 4, figures 9-17 2001b Sudanomocarinasp." Peng, Babcock, and Lin, p. 103, pl. 6, fig. 8.

Etymology. From Chinese, Huanan, South China, referring to its record from South China. Holotype. Incomplete cranidium (P1.4, fig. 16; NIGP 137926) from collection P204.

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Other material. Four cranidia and two pygidia in collections P130.5, P134.2, P 164.2, P171 and W146.2. Diagnosis. Sudanomocarina? with glabella that is subconical, largely effaced; fixigenae narrow; anterior border furrow arched forward; plectrum clearly defined, not extending to preglabellar furrow; eye ridge wide, short; palpebral lobes rather small, located near glabellar midlength. Pygidium with wide axis; borders moderately wide. Description. Anterior border gently convex, slightly longer (sag.) than preglabellar field, narrowing rapidly abaxially; preglabellar field slightly depressed, sloping gently forward; preocular field gently convex, sloping forward and outward. Glabella large, moderately convex, tapered gently forward, rounded to obtusely rounded anteriorly, occupying about four-fifths of cranidial length; lateral furrows mostly effaced. Occipital furrow well defined, shallowing at sides; occipital ring crescentic, posterior margin arched rearward, beating tiny medial node. Palpebral area gently convex, about one-fourth width of glabella; palpebral lobe moderately arcuate, about one-third as long as glabella, located near axial furrow opposite midlength of glabella, defined by deep palpebral furrow; eye ridge short and wide. Anterior branch of facial suture diverging forward at 350-40 ° to sagittal line, turning sharply inward and forward, cutting anterior border almost straight and meeting anterior margin opposite anterolateral comer of glabella; posterior branch extending outward and slightly rearward, deflected rearward about two-thirds of distance to posterior margin, enclosing broad, blade-shaped posterolateral projection. Pygidium semicircular, length about three-fifths width. Axis rather wide, with 4-5 faint tings, plus semicircular terminal piece extending to posterior border furrow, and short, wide postaxial ridge extending to posterior margin; pleural field gently convex, rather effaced, with weakly defined pleural furrows and obscure interpleural furrows; border furrows shallow; borders flat, of almost uniform width. Remarks. Except for the structure of the anterior area of the cranidium, and probably also the shorter palpebral lobes, this new species is essentially indistinguishable from the holotype of Pseudanomocarina plana (Chernysheva in Chernysheva et al., 1956, p. 31, fig. 7), the type species of Pseudanomocarina, from the Amgan Formation of Siberia. It differs from all other species assigned to Sudanomocarina in lacking a distinct plectrum. In this respect, Sudanomocarina? huananensis sp. nov. is comparable to species assigned to Maotunia, especially M. distincta (Resser and Endo, 1937, pl. 37 figs. 7, 9 (cranidium only); also Zhang and Jell, 1987, pl. 82, figs. 5-14; pl. 83, figs. 2-12). M. distincta, however, differs in having a proportionally larger glabella with a more broadly rounded front, shorter and more anteriorly placed palpebral lobes, and shorter eye ridges. Maotunia semiplectra is also comparable to S. ? huananensis, but can be readily distinguished by the transverse rather than forward-arched anterior border furrow, longer palpebral lobes, wider fixigenae, and narrower border furrows. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Goniagnostus nathorsti Zone and the Lejopyge laevigata Zone).

• 46

•

Sudanomocarina sp. Plate 19, figure 15 Material. One cranidium (NIGP 138085), in collection P 171.5. Remarks. The single known cranidium of this species, which may represent a new species, is left in open nomenclature. It is most similar to Sudanomocarina changi. The cranidium of this undetermined species is different from that of S. changi in having a proportionally larger glabella and a shorter preglabellar field. The undetermined cranidium is also similar to Sudanomocarina? huananensis sp. nov., but can be differentiated from that species by the less tapered and more broadly rounded anterior glabella, the shorter preglabellar field, the shorter anterior border, and a nearly transverse rather than rearwardly arched occipital furrow. Stratigraphically, S. sp. occurs slightly higher than S. ? huananensis. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Genus ZHUJIA Ju in Qiu et al., 1983 Zhujia Ju in Qiu et al., 1983, p. 160; Peng, Lin, and Chen, 1995, p. 285. Type species. Zhujia lubrica Ju in Qiu et al. (1983, p. 160, 161, pl. 50, figs. 6, 7) from the Yangliugang Formation, Zhuji, westem Zhejiang; by original designation. Emended diagnosis. Proasaphiscidae with cranidium and pygidium rather effaced; glabella tapered forward moderately, rounded anteriorly, with weak carina and four pairs of shallow lateral furrows; preglabellar area moderately wide; palpebral lobe relatively small, located subcentrally. Pygidium semielliptical, wider than long; axis with 5-6 tings, reaching to posterior border furrow. Exfoliated surface with clearly defined axial, occipital, preglabellar, and posterior border furrows. Remarks. Peng et al. (1995) transferred this genus from the family Tsinaniidae, to which the genus was first referred (Ju in Qiu et al., 1983), to the family Proasaphiscidae, on the basis of the morphology of the exfoliated cranidium and pygidium. Ju (in Qiu et al., 1983, p. 160) discussed the differences between Zhujia and Liopeishania Chang (Zhang, 1963, p. 473, 486). Although Zhujia and Liopeishania are similar and probably closely related, Zhujia is sufficiently different to warrant status as a separate genus. Liopeishania was erected with Psillaspis (?) convexus Endo, 1937 as the type species, and the genetic concept was based on a single cranidium (Endo, 1937, p. 350, pl. 59, fig. 1, non 3, 4; Zhang, 1963, pl. 2, figs. 15, 16). The other three specimens assigned to the species by Endo were later excluded from the species by Zhang (1963). This cranidium serves to demonstrate that the species is strongly convex, both transversely and sagitally; it has a strongly convex glabella (especially in the posterior half), and the glabella is conical and elongate-ovate in outline. The glabella lacks both a carina and lateral furrows. The anterior branches of the facial suture in Liopeishania are subparallel or slightly diverging forward, and the occipital furrow is distinct on testaceous cranidia. In comparison, the •

47

•

occipital furrow is mostly effaced in testaceous specimens of Zhujia. Peng et al. (1995) outlined the distinction between Zhujia and Lioparella Kobayashi, 1937. Lioparella differs from Zhujia in having a larger palpebral lobe, a glabella that is truncated anteriorly, and a spinose pygidium.

Zhujia hunanensis Peng, Lin, and Chen, 1995 Plate 14, figures 1-10 1995 Zhujia hunanensis Peng, Lin, and Chen, p. 297, 298, pl. 5, figs. 7-10. 2001b Zhujia hunanensis Peng, Lin, and Chen; Peng, Babcock, and Lin, p. 102, pl. 3, fig. 6.

Holotype. Partly exfoliated cranidium (Peng et al., 1995, pl. 5, figs. 8a, b; pl. 14, figs. 6, 7 herein; NIGP 118863), from the Ptychangnsostus punctuosus Zone, Huaqiao Formation, Paibi, northwestern Hunan.

New material. About 20 sclerites, including cranidia and pygidia (illustrated specimens NIGP 118862, 118864, 118865, 138025-138028), in collections P108, P108.5, and P115.6.

Remarks. Yuan et al. (2000, p. 141) transferred Zhujia tiunanensis to Liopeishania. Because of differences between Liopeishania and Zhujia discussed under Zhujia, that opinion is rejected, and the species is retained in Zhujia. New topotypic material from northwestern Hunan is identical with the type series. As recorded previously (Peng et al., 1995, p. 298), this species is confined to the Ptychagnostus punctuosus Zone (Peng and Robison, 2000). A reference to specimens from the Lejopyge laevigata Zone provided in the plate explanation of Zhujia hunanensis (Peng et al., 1995, p. 298) was in error.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Dorypyge richthofeni Zone to the lower part of the Pianaspis sinensis Zone (equivalent to the Ptychagnostus punctuosus Zone). Genus PAIBIELLA gen. nov.

Etymology. From Paibi Village, plus Latin suffix -ella, small, referring to the site where the genus was first discovered, and the small size of the fossil.

Type species. Paibiella paibiensis gen. et sp. nov. Diagnosis. Proasaphiscidae with glabella that is elongate-subrectangular, sides constricted gently; with four pairs of lateral furrows; S 1 bifurcate; $3 isolated from axial furrow; occipital ring with moderately large occipital spine; preglabellar area occupying less than one-fifth of cranidial length (excluding occipital spine), with anterior border subequal in width to preglabellar field; anterior branch of facial suture converging gently; posterior branch enclosing short (tr.) posterior area of fixigena. Surface covered with dense granules.

Remarks. Paibiella gen. nov. and Huayuanaspis are the only genera referred to the family • 48

•

Proasaphiscidae that have an occipital spine. Huayuanaspis has an anteriorly truncated glabella with four pairs of lateral furrows, bifurcate S 1 furrows, a gently arched anterior border furrow, a moderately long and strongly oblique eye ridge, and an occipital spine. Paibiella differs from Huayuanaspis principally in having a proportionally longer glabella with constricted sides, a shorter (sag.) preglabellar area, and forward-converging anterior branches of the facial suture. In Huayuanaspis, the glabella has straight sides, the preglabellar area occupies more than one-fourth of the cranidial length, and the anterior branches of the facial suture diverge forward. Excluding the occipital spine, the new genus bears some similarity to Proasaphiscina Lin and Wu (in Zhang et al., 1980b, p. 76). Proasaphiscina (type species Proasaphiscina quadrata) is characterized by having a subrectangular glabella with four pairs of lateral furrows, a short preglabellar area, and a gently arched anterior border furrow (Zhang et al., 1980b, p. 76, 77, pl. 9, figs. 1-3). However, it can be distinguished from Paibiella by its larger palpebral lobe, divergent rather than convergent anterior branches of the facial suture, and more strongly diverging posterior branches.

Paibiella paibiensis gen. et sp. nov. Plate 9, figures 12-14 2001b Crepicephalid? gen. et sp. nov.; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 7.

Etymology. From Paibi Village. Holotype. Cranidium (P1.9, figs. 13, 14, NIGP 137985) from collection P240.5. Other material. A meraspid cranidium (NIGP 137984) in collection P249. Diagnosis. As for the genus. Description. Anterior border subequal in width with preglabellar field, gently convex, arched gently forward, defined posteriorly by distinct border furrow. Glabella subrectangular, defined by deep axial and preglabellar furrows, with sides gently constricted, occupying about 0.82 of cranidial length; lateral glabellar furrows consist of four pairs; S 1 bifurcate with posterior branch extending rearward for about one-third of its length, then deflecting inward and rearward; anterior branch transverse, weakly impressed; $2 and $3 weak, long, directed inwardly and rearwardly; $4 shorter but deeper than $3, directed inwardly and forwardly, S1, $2, $4 connected to axial furrow; $3 isolated from axial furrow. Occipital furrow transverse, shallow medially, deeper at sides; occipital ring widest sagittally, narrowing abaxially, with long occipital spine extending rearward and strongly upward. Preglabellar area less than one-fifth of cranidial length. Eye ridge distinct, strongly oblique, subparallel to anterior border furrow. Palpebral lobe broken, but probably of moderate size, located subcentrally, posterior end opposite anterior one-third of L1, anterior end opposite $3. Anterior branch of facial suture converging gently forward, straight until meeting anterior border furrow, then tuming rather sharply, converging forward and deflected inward slightly to anterior border furrow, then continuing inward and forward to cut anterior border obliquely; posterior branch diverging about 40 ° to sagittal line. Posterior area of fixigena about two-thirds as wide as occipital ring, with deep border furrow and abaxially widening posterior border. • 49.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the upper part of the Lejopyge laevigata Zone). Genus PARAYUJINIA gen. nov.

Etymology. From Greek prefix, para-, near (in the sense of a comparison); plus genetic name Yujinia, referring to its resemblance to the genus. Type species. Parayujinia coustricta gen. et sp. nov. Diagnosis. Proasaphiscidae with glabella that is large, tapered, constricted laterally, with four pairs of moderately impressed lateral furrows and deep occipital furrow; preglabellar field short; anterior cranidial border upturned steeply; palpebral lobe located subcentrally, close to axial furrow; palpebral area narrow. Librigena with genal spine that is broad at base; lateral border steeply upturned. Remarks. Parayujinia gen. nov. is characterized by having a proportionally large glabella that is furrowed, constricted laterally, and broadly rounded anteriorly. It also has a nearly effaced preglabellar field, strongly upturned anterior and lateral cephalic borders, and palpebral lobes that are situated close to the axial furrows. In having a furrowed glabella and upturned anterior border, the new genus bears similarity to Yujinia Zhang and Yuan (in Zhang et al., 1980b), but Yujinia differs in having much wider fixigenae, more widely spaced palpebral lobes, and a longer preglabellar field. Yujinia may be further differentiated by having an unconstricted glabella with less rounded anterolateral comers, and less well impressed lateral and occipital furrows. The new genus, with its furrowed glabella beating bifurcated S1, is reminiscent of some species previously referred to Honania, such as Honania lata Lee (in Lu et al., 1965, pl. 63, figs. 3-5) and H. pansa Zhang and Yuan (in Zhang et al., 1980b, pl. 10, fig. 4). Honania was suppressed as junior synonym of Grandioculus Cossmann, 1908 by Zhang and Jell (1987, p. 166). Both G. lata and G. pansa are essentially identical to Parayujinia constricta gen. et sp. nov. in the shape and the convexity of the glabella, the nature of the lateral and occipital furrows, the shape of the occipital ring, and the nature of the anterior border. However, those two species differ from P. constricta in having wider fixigenae, longer eye ridges, shorter palpebral lobes, and subtriangular rather than blade-shaped posterolateral projections. These differences, as well as the shape of the glabella, which is proportionately shorter and unconstricted laterally, serve also to distinguish Liostracus megalurus Dames, the type species of Grandioculus (see Schrank, 1977, p. 152, 153, pl. 4, figs. 7-9; pl. 5, figs. 1-4), from the new genus. The new genus is referred to the Proasaphiscidae because it bears resemblance to Proasaphiscus Chang, 1963 and related taxa in overall features. However, the new genus differs from other proasaphiscids in having a proportionally larger glabella, a shorter preglabellar field, and an upturned anterior cranidial border. Parayujinia constricta gen. et sp. nov. Plate 19, figures 1-14; Text-figure 5

Etymology. From Latin, constrictus, constrict, referring to the constricted nature of the axial furrow. • 50.

Holotype. Cranidium (P1. 19, fig. 3; NIGP 138075) from collection P37. Other material. More than 20 sclerites, including cranidia, librigenae, and pygidia (illustrated specimens 138073, 138074, 138076-138084) in collections P12.5, P33.5, P37 and W3. Diagnosis. As for the genus.

' " -~ • -

~:~. " "" "i' .

"';'. . . . ' :

.

. . . . ":'~.'.::,,

Text-figure 5. Reconstruction of cephalon and pygidium of Parayujinia constricta gen. et sp. nov. Cephalon based mostly on specimens NIGP 138073 and 138075 (see P1. 19, figs. 1, 3); pygidium based on specimen NIGP 138081 (see P1. 19, fig. 10).

Description. Cephalon semicircular; anterior and lateral borders upturned; with short genal spine, broad at base. Anterior area short, with extremely short preglabellar field and strongly upturned anterior border; anterior border furrow shallow, identified by change in slope of anterior and preglabellar field; preocular area flat, subtriangular, sloping slightly forward and outward, with anterolateral margin opposite outer limit of palpebral lobe. Glabella large, occupying about 0.85 of cranidial length, tapering gently forward, constricted gently at $2, broadly rounded anteriorly, with rounded anterolateral comers; lateral glabellar furrows consist of four distinct pairs; S1 long, bifurcate, posterior branch directed diagonally; $2 curved forward gently, directed inward and slightly rearward, located near midlength of glabella (excluding occipital ring); $3 shallow, short, horizontal, isolated from axial furrow; $4 equal in length to $3, directed inward and forward, located immediately anterior to eye ridge; occipital furrow deeply impressed, slightly sinuous, arched forward slightly near middle, curving forward laterally; occipital ring prominent, widest sagittally, narrowing abaxially, posterior margin bowed, with median node. Palpebral area narrow, gently convex, about one-fourth as wide as glabella; palpebral lobe curved gently, slanting gently, about one-third as long as glabella, lying slightly posterior to cranidial midpoint, defined by shallow palpebral furrow that becomes incised deeply on exfoliated surface. Anterior branch of facial suture diverges forward initially at 30°-40 ° to sagittal line, then curves inward evenly to .51.

define rounded anterolateral comer of cranidium; posterior branch gently sigmoidal, enclosing narrow, blade-like posterolateral projection; posterior border furrow well impressed. Librigena with wide, gently convex genal field, lateral border moderately wide, becoming progressively upturned forwardly; lateral and posterior border furrows merge without extending to genal angle. Pygidium subpentagonal with obtusely angled lateral margin. Axis conical, occupying 0.70-0.80 of pygidial length, with 3-4 tings and short, wide postaxial ridge; anterior two tings clearly defined, following one or two tings weakly defined. Pleural field with 4-5 pleurae; pleural furrows deep; interpleural furrows faint; borders narrow, weakly defined.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Family

CATILLICEPHALIDAE Raymond,

1938

Remarks. Fortey and Chatterton (1988, p. 210) regarded the Catillicephalidae as a probable polyphyletic taxon. The group includes taxa beating either a rostral plate or a median suture ventrally. According to Fortey and Chatterton (1988), the median suture indicates a monophyletic origin of the order Asaphina. The median suture-beating catillicephalids such as Acheilus Raymond, 1924 (non Clark 1924; non Theodenisia Clark, 1948; see Fortey, 1983, p. 186 for nomenclatural discussion) are regarded as resembling the Isocolidae more than the Catillicephalus. The presence of a rostral plate in Catillicephalus suggests that this genus has a close affinity with the Ptychopariidae. The earlier catillicephalids, those of middle and early late Cambrian age, probably have a similar ventral structure. Here, the median suture-beating Acheilus and its allies are regarded as asaphinids, and excluded from the Catillicephalidae. Recently, Fortey (in Whittington et al., 1997, p. 302) classified the Catillicephalidae as an undetermined suborder within the Ptychopariida. That proposal is not followed here. Pending the discovery of further information about the ventral structure of other catillicephalids, we prefer to retain the Catillicephalidae sensu stricto within the Ptychopariina. Subfamily CATILLICEPHALINAERaymond, 1938 Genus BUTI'SIA Wilson, 1951

Buttsia Wilson, 1951, p. 626-628; Rasetti, 1954, pl. 606; Lazarenko, 1966, p. 69, 70; Khramova, 1977, p. 66; Lu and Lin, 1989, p. 136, 137, 251. Buttsia (Waergangia) Lin, 1991, p. 371-375. Type species. Buttsia drabensis Wilson, 1951 (p. 627, 628, pl. 89, figs. 19-24) from the Conococheague Limestone and the Ore Hill Member of the Gatesburg Formation, central Appalachian Mountains, USA; by original designation. Other species. Buttsia pinga Lararenko (1966, p. 70, 71, pl. 8, figs. 9-20) from the Agnostus pisiformis Zone and the Homagnostus fecundus Zone, northwestern Siberian Platform; Bunsia parvula Khramova (1977, p. 66, 67, pl. 15, figs. 1, 2) from the Chukuk Formation (basal upper Cambrian), northern Siberian Platform; Bunsia sp. (Romanenko, 1977, p. 183, pl. 25, fig. 18) from "52"

the upper part of the middle Cambrian and the basal part of the upper Cambrian, northeastern Altai; Buttsia inexpectata Fortey (1980, p. 37-39, pl. 3, figs. 25-28) from the Olenidsletta Member (middle Arenig), Spitzbergen, Norway; Buttsia globosa Lu and Lin in Peng (1987, p. 94, 95, pl. 6, figs. 1-5) from the Huayanssu Formation, western Zhejiang, and the Huaqiao Formation, northwestern Hunan.

Emended diagnosis. Catillicephalidae having cranidium that is subsemicircular, convex; glabella inflated, expanding forward, with faint lateral furrows; occipital ring with or without median node and occipital spine; preglabellar field short or absent; palpebral lobe of medium to large size, located medially or posteriorly; fixigena downsloping strongly. Thorax with nine segments, having short axial spines. Pygidium with large, convex, broadly-conical axis reaching close to posterior margin; pleural region narrow (tr.), poorly segmented, with or without lateral pleural spines. Remarks. Characters that serve to distinguish Buttsia from Catillicephala include the presence of a distinct anterior border, the presence of lateral glabellar furrows, larger palpebral lobes, and a pygidial axis that does not reach to the posterior margin. Rasetti (1954) noted that the posterior branch of the facial suture meets the posterior margin of the cranidium at the genal angel in Catillicephala, but at a distance from the genal angel in Buttsia. The course of the posterior branch of the facial suture of Buttsia is well exemplified by the cephalon of Buttsia pinga from the northwestern part of the Siberian Platform (Lazarenko, 1966, pl. 8, fig. 17), which bears a somewhat displaced librigena with a genal spine at the genal angle. Lazarenko (1966) and Lin (1991) illustrated the exoskeletons of B. pinga and B. globosa respectively, and each bears a thorax of nine segments. Each thoracic segment bears an axial spine. Buttsia has some similarity to specimens assigned to Paradistazeria [=Distazeris (Paradistazeria) Zhu in Zhang et al., 1980a and Distazeris sensu Peng, 1987]. However, the cranidia in these Chinese specimens can be differentiated by having a considerably shorter (tr.) anterior border that is usually transverse rather than arched forward, converging anterior branches of the facial suture, and a pygidium with well furrowed pleurae. Lu and Lin (1989, p. 137, 251) indicated that Distazeris (Paradistazeria) can also be distinguished from Buttsia by a proportionally longer glabella and more pairs of lateral glabellar furrows, but variation observed in large collections of this genus obscures such distinctions. Buttsia has a relatively wide geographic distribution, being known from Laurentia (Wilson, 1951; Rasetti, 1954; Stitt, 1977), Siberia (Lazarenko, 1966; Khramova, 1977; Romanenko, 1977; Gogin, 1990), Baltica (Fortey, 1980), and South China (Peng, 1987; Lu and Lin, 1989). Stratigraphically, it ranges from the Proagnostus bulbus Zone to the Glyptagnostus reticulatus Zone in South China. It also occurs in the lower part of the equivalent interval in Altai, and the middle part of the equivalent interval in Siberia. It occurs in the younger Elvinia Zone in North America in even younger strata (middle Arenig age) in Baltica. Geographically progressively younger occurrences suggest that members of the genus migrated from South China or Altai via Siberia to Laurentia, and then on to Baltica. Buttsia globosa Lu and Lin in Peng, 1987 Plate 20, figures 1-14; Plate 21, figures 1-3; Text-figure 6 1987 1989 1991

Buttsia globosa Lu and Lin in Peng, p. 94, 95, pl. 6, figs. 1-5. Buttsia globosa Lu and Lin, p. 137, 138, pl. 21, figs. 7-10. Buttsia (Waergangia) spectabilis Lin, p. 372, pl. 2, figs. 4, 5, 7. .53"

1991 1999 2001b 2001 d

Buttsia(Waergangia) globosa (Lu and Lin), p. 372, pl. 2, fig. 6. Distazerissp. cf. D. hunannensis Peng; Duan, Yang, and Shi, p. 161, fig. 12F. Buttsiaglobosa Lu and Lin; Peng, Babcock, and Lin, p. 104, pl. 9, fig. 12; pl. 11, figs. 5, 6. Buttsiaglobosa Lu and Lin; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.1.

Holotype. Well preserved cranidium (Peng, 1987, pl. 8, figs. 4a, b; Lu and Lin, 1989, pl. 21, 9a, b, NIGP 66430), from the Lejopyge sinensis Zone, Duibian, Jiangshan, western Zhejiang. New material. More than 30 cranidia and one pygidium (illustrated specimens NIGP 138087138097) in collections P282.75, P298.4, P301.9, P307.4, P319.6, P319.8, W195.3, W197.5, W215.1, W216.5, W219.7, W221.5, W223.2, W225, and W227. Remarks. New material from the Huaqiao Formation, northwestern Hunan, resembles the type material from western Zhejiang and northwestern Hunan. Key characters that warrant assignment of the new material to this species include a wide, inflated glabella with two pairs of long, faint, curved lateral furrows; a long and thin, moderately convex, notably forward-arched anterior border; an occipital ring with a small median node lying close to the occipital furrow and a short occipital spine; a strongly downsloping fixigena; a prominent, tumid, broadly conical, pygidial axis; and a pleural region having only the anterior part segmented. Well-preserved pygidia from western Zhejiang (Lu and Lin, 1989, pl. 21, figs. 7, 8) show that each of first three pleural segments bears a pair of lateral spines.

Text-figure 6. A-D. Buttsia globosa Lu and Lin in Peng, 1987, described originally as the holotype of Buttsia (Waergangia) spectabilis Lin, 1991, the type species of the subgenus Buttsia (Waergangia) Lin, 1991, NIGP T180, all x 6 (original of Lin, pl. 2, figs. 4, 5). A, internal mold of exoskeleton; note the displaced cephalic doublure (indicated with triangle); B-D, latex cast from external mold of the same exoskeleton, dorsal, lateral, and anterolateral views. In having an occipital spine, a long and curved S 1 furrow, and a well defined anterior border, Buttsia globosa is different from B. drabensis, the type species of Buttsia, but closely similar to B. pinga Lazarenko. B. pinga, however, is differentiated from B. globosa by having a less inflated • 54.

glabella with sides that are less bowed outward, a transverse rather than curved $2, a shorter pygidial axis; and in lacking an occipital node and lateral pleural spines on the pygidium. Buttsia (Waergangia) spectabilis Lin (1991; Text-fig. 6 herein) is apparently a junior synonym of B. globosa. Lin's (1991) specimens were collected from the same horizon and section that yielded Peng's (1987, pl. 6, figs. 1-3) material of B. globosa, and are assigned to two species: B. (W.) spectabilis and B. (W.) globosa. A large collection of B. globosa from northwestern Hunan (Peng, 1987) and western Zhejiang (Lu and Lin, 1989) show that Lin's (1991) specimens fall well within the range of morphological variation of that species, and that B. spectabilis and B. globosa are synonymous.

Occurrence. The holotype is from the Lejopyge sinensis Zone, Duibian, Jiangshan, western Zhejiang, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the upper part of the Proagnostus bulbus Zone and the lower part of the Linguagnostus reconditus Zone). Genus MADAROCEPHALUSResser, 1938a

Type species. Madarocephalus laetus Resser, 1938 (p. 87, pl. 10, figs. 51, 52, ?53) from the Nolichucky Formation, Alabama, USA; by original designation. Other species. Except for the type species and Madarocephalus orientalis sp. nov., this genus contains one recognized species, M. scolus Robison (1988, fig. 16.8-10) from the Holm Dal Formation, North Greenland; and one species left in open nomenclature, cf. M. laetus Resser (Pratt, 1992, pl. 26, fig. 23) from the Mackenzie Mountains, northwestern Canada. Madarocephalus minor Rasetti was suppressed as a junior synonym of M. laetus (Pratt, 1992). Remarks. Madarocephalus is a distinctive catillicephalid, differing from all other genera of the Catillicephalidae by having a long, highly convex, forward-expanding glabella with straight or inwardly bowed sides, and palpebral lobes that are located rather far forward. Robison (1988) emended the genetic concept of Resser (1938a) and Rasetti (1946, 1954) to include a species from Greenland that bears an anterior border. That concept is followed here. A pygidium has not been associated with confidence to cranidia of Madarocephalus. A pygidium that was associated with the syntype cranidia of M. laetus by Resser (1938, pl. 10, fig. 53) was assigned only questionably to that species by Pratt (1992). Previously, Madarocephalus was known only from Laurentia (Alabama, Quebec Tennessee, Greenland, and Mackenzie Mountains) (Resser, 1938a, Rasetti, 1946, 1965; Robison, 1988; Pratt, 1992). Collections from northwestern Hunan, South China, are the first record of the genus outside of North America-Greenland. The Hunan material includes a complete cephalon, not just a cranidium, and this is the first such specimen recorded for Madarocephalus. The cephalon of M. orientalis sp. nov. bears narrow (tr.), borderless librigenae, broadly rounded genal angles, a small inflated eye, and gonatoparian facial sutures. Madarocephalus resembles Catillicephala in these respects.

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Madarocephalus orientalis sp. nov. Plate 23, figures 1-17 2001b Madarocephalus sp. cf. M. laetus Resser; Peng, Babcock, and Lin, p. 103, pl. 6, figs. 6, 7; pl. 7, fig. 1.

Etymology. From Latin, orientalis, eastern, referring to its occurrence in the Far East. Holotype. Cephalon (P1.23, figs. 4-7; NIGP 138121) from collection P240.5. Other materal. Ten cranidia (NIGP 138119, 128120, 138122-138127) in collections P240.5, P249, P268.3, P277, P273.8, W146.2, and W187.8. Unfigured material includes several cranidia in collections P259.8, P261.35, P273.46, P273.66, and P282.75. Diagnosis. Madarocephalus with broadly or obtusely rounded anterior cranidial margin and without anterior border; glabella (except occipital ring) with bifurcated S 1 furrow, length about two times the basal width; prominent palpebral lobe situated at the level of anterior third of glabella. Description. Cranidium trapezoidal or semielliptical, highly convex, slightly wider than long (except occipital spine). Anterior margin arched gently or rounded obtusely. Glabella moderately convex, expanding forward; length (excluding occipital ring) about twice basal width, occupying entire cranidial length. Axial furrow slightly concave outward, deeply impressed posteriorly, becoming shallow anteriorly. Lateral glabellar furrows consist of 2-3 pairs, obscure on external surface, faint on exfoliated surface; S1 long, bifurcate, with posterior branch strongly oblique rearward, anterior branch directed inwardly; $2 and $3, if visible, short and transverse. Occipital furrow deeply incised, curved gently rearward; occipital ring moderately long, with stout occipital spine. Palpebral lobe a short bar, defined by deep palpebral furrow, deflected gently, located near level of anterior third of glabella (except occipital ring); posterior end just anterior of midpoint of glabella; eye inflated, teardrop-shaped, anterior bluntly pointed, posterior end rounded; eye ridge short and nearly effaced. Anterior branch of facial suture short, converging forward; posterior branch meets posterior margin at genal angle, acutely rounded. Fixigena moderately convex, downsloping strongly abaxially with maximum width at posterior margin; posterior margin narrower than basal glabellar width. Posterior border furrow deeply incised, terminating before reaching facial suture; posterior border narrow medially, widening rapidly distally, and extending strongly rearward. Librigena narrow, long, slightly convex, smooth; border and border furrow absent. Remarks. Madarocephalus orientalis sp. nov. is most similar to M. laetus, the type species of the genus. M. orientalis is distinguished from M. laetus by having a proportionally longer glabella with sides that are bowed inward rather than straight, more prominent palpebral lobes, and a less well arched anterior margin of the cranidium. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Pianaspis sinensis Zone to the upper part of the Wanshania wanshanensis Zone (equivalent to the Lejopyge laevigata Zone • 56"

through the upper part of the Proagnostus bulbus Zone). Genus PARADISTAZERISZhu in Zhang et al., 1980a

Distazeris (Paradistazeris) Zhu in Zhang et al., 1980a, p. 379, 380; Sun, 1984, p. 363. Type species. Distazeris (Paradistazeris) sichuanensis Zhu in Zhang et al., 1980a (p. 380, pl. 132, fig. 12) from the Dashuijing Formation, Youyang, southeastern Sichuan, China; by original designation. Other species. Distazeris (Paradistazeris) hubeiensis Zhu in Zhang et al. (1980a, p. 380, pl. 132, figs. 13-15) from the lower part of the Haozituo Group (upper Cambrian), Xianfeng, southwestern Hubei; Distazeris hunanensis Peng (1987, p. 93, 94, pl. 6, figs. 7-9) from the lower part of the Proagnostus bulbus Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan; Distazeris dongtingensis Peng (1987, p. 94, pl. 6, fig. 6) from the lower part of the Proagnostus bulbus Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan; Distazeris sp. cf. D. hunanensis Peng (Duan, Yang, and Shi, 1999, p. 161, fig. 12F, from the Huaqiao Formation, Fenghuang, northwestern Hunan. Remarks. Paradistazeris was erected as subgenus of Distazeris to accommodate two Distazeris-like species from South China (Zhu in Zhang et al., 1980a). The subgeneric assignment was based largely on the genetic concept of Rasetti (1954) and the reconstruction of Rasetti (1954, fig. l d; also in Moore, 1959, fig. 210.2) for Distazeris acuta Raymond, the type species of Distazeris. The reconstruction shows a forward-expanding or pyriform glabella and a preglabellar area lacking a preglabellar field but having an anterior border. According to Pratt (1992, p. 92), the reconstruction is misleading in the shape of the glabellar outline and the addition of an anterior border. Pratt (1992) rediagnosed the genus as a catillicephalid with a rectangular-shaped glabella beating two pairs of faint lateral furrows, a short occipital spine, and a short, downsloping preglabellar area lacking an anterior border. Pratt's (1992) concept differs substantially from that of Rasetti (1954). Based on the revised concept, Pratt (1992) transferred Ucebia simplex Raymond, 1937 and Distazeris rasettii Shaw, 1962 [=Cephalocoelia rotunda Rasetti, 1946, pl. 68, figs. 18-23], both of which were previously referred to Distazeris by Rasetti (1954), to Pemphigaspis. Pratt (1992) restricted Distazeris to its type species, and excluded all other species, including four species from China, from the genus. D. (Paradistazeris) sichuanensis Zhu and D. (Paradistazeris) hubeiensis Zhu were reassigned to Agelagma Opik, 1967, whereas Distazeris hunanensis Peng and Distazeris dongtingensis Peng were regarded as resembling Pemphigaspis Hall, 1863 in cranidial morphology. Pratt's (1992) reassignment of the Chinese species is rejected here. The four Chinese species, all of which are all from the Jiangnan Slope Belt (Peng, 1990) and adjacent areas on the eastern margin of the Yangtze Platform, seem to be congeneric. They do not fall within the concept of Agelagma, which is characterized by the presence of a wide (sag.) preglabellar field and the absence of an occipital spine. They also do not fall within the concept of Pemphigaspis, which is characterized by lacking a clearly defined anterior border and in having a pygidium with distinct kidney bean-shaped pleural fields and a slender axis (see Palmer, 1951). The genetic concept of Distazeris remains unclear. Pratt's (1992) concept of the genus was probably based more on specimens from the Mackenzie Mountains of Canada than on Raymond's (1937) holotype of D. acuta, the type species from Vermont. Specimens from the Mackenzie Mountains differ from the Vermont material in having a less convex glabella, and the glabella is forward-tapering rather than parallel-sided. Until more information is known about the holotype, • 57-

we prefer to regard Paradistazeris as a separate genus, and refer D. (Paradistazeris) sichuanensis, D. (Paradistazeris) hubeiensis, Distazeris hunanensis, and Distazeris dongtingensis to the genus. Paradistazeris differs from Distazeris in having large, arcuate palpebral lobes and in having a prominent anterior border. In cephalic morphology, Paradistazeris is similar to Pemphigaspis, but it differs in having an anterior border, and in having smaller and shorter genal spines. Paradistazeris hubeiensis Zhu in Zhang et al., 1980a

Plate 21, figures 7-18; Plate 32, figures 8, 9 1980a 1984 1987 2001b

Distazeris (Paradistazeris) hubeiensis Zhu in Zhang et al., p. 380, pl. 132, figs. 13-15. Distazeris(Paradistazeris) hubeiensis Zhu; Sun, p. 363, pl. 138, figs. 8-11. D&tazerishubeiensis Zhu; Peng, p. 92, 93, pl. 6, figs. 10, 11. Distazeris hubeiensis Zhu; Peng, Babcock, and Lin, p. 104, pl. 10, fig. 1.

Holotype. Incomplete cranidium (Zhu in Zhang et al., 1980a, pl. 132, fig. 13; NIGP 38152) from the lower part of the Haozituo Group (lower Furongian), Xianfeng, southwestern Hubei. New material. About 20 sclerites, including cranidia, librigena, and pygidia (illustrated specimens 138101-138109, 138216) in collection P 319.6, W199.2, W200.3, W201, W207.5, and W211.7. Remarks. New material of Distazeris hubeiensis from the Huaqiao Formation of northwestern Hunan resembles material in the type series from the same region. The cranidium of the species has a narrow, straight-sided glabella with four distinct pairs of lateral glabellar furrows; the S 1 furrows are obscurely bifurcated, the $2, $3, and $4 furrows are isolated from the axial furrows; the fixigena is narrow and has anastomosing lines; and a long median spine is present on the occipital ring. The librigena has a wide, convex lateral border bearing terrace lines; a shallow border furrow is present; the genal field is relatively flat and has anastomosing lines; a short genal spine having a broad base is present. The eye is elongate-kidney bean-shaped, of holochroal type, with round, densely spaced lenses arranged in linear fashion. The associated pygidium is similar to that of Distazeris hunanensis (Peng, 1987, pl. 6, figs. 7-9) in having a cylindrical axis that extends to a position in front of the posterior border, and a well furrowed pleural field with evenly divided pleural bands. The pygidium of D. hubeiensis differs only in having an axis that is proportionally narrower and longer than that of D. hunanensis. Occurrence. The holotype is from the lower part of the Haozituo Group, Xianfeng, southwestern Hubei, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone and the lower part of the Linguagnostus reconditus Zone). Paradistazeris hunanensis (Peng, 1987)

Plate 21, figures 4-6 1987 Distazeris hunanesis Peng, p. 93, 94, pl. 6, figs. 7-9.

• 58.

Holotype. Cranidium (Peng, 1987, pl. 6, fig. 7; NIGP 74511), from the Proagnostus bulbus Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan. New material. Seven cranidia (illustrated specimens NIGP 138098-138100) in collections P298.4, P301.9, P362.45(?), P364.5(?), and W215.1. Remarks. Additional material from the Huaqiao Formation, northwestern Hunan, agrees in all respects with the type material, which was collected from the same region. The cranidium of D. hunanesis is characterized by a subrounded, somewhat tumid glabella; the S 1, $2, and $4 furrows are prominent but the $3 furrow is almost completely effaced. The S 1 furrow is the longest of the lateral glabellar furrows. The S1 furrow has a transverse outer part that occupies about half the distance between the axial furrow and the sagittal line; it has a clearly bifurcated inner part in which the posterior branch is well impressed and directed rearwardly and slightly inwardly, and in which the anterior branch is weakly impressed and directed inwardly and slightly forwardly. The occipital ring bears a moderately long, stout median spine. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the upper part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone through the Glyptagnostus stolidotus Zone). Paradistazeris rotundus sp. nov. Plate 22, figures 1-15

Etymology. From Latin rotundus, nearly rounded, referring to the shape of glabella. Holotype. Partly exfoliated cranidium (P1. 22, figs. 9, 14, 15; NIGP 138115) from collection W216.5. Other material. More than 20 sclerites, including cranidia, and pygidia (illustrated specimens 138110-138114, 139116-138118) in collections W212.45, W215.1, W216.5, and W218.3, among which seven cranidia and two pygidia are illustrated as paratypes. Diagnosis. Paradistazeris with glabella that is nearly round, largely effaced, bearing only S1 and $2 furrows, S 1 and $2 weak to moderately distinct; palpebral lobe large, with anterior end opposite or slightly posterior to glabellar front; preocular field short. Pygidium with axis reaching to posterior border furrow. Description. Cranidium strongly convex, semicircular, length slightly more than half of maximum width. Anterior border linear, transverse in dorsal view and gently arched upward in anterior view; preglabellar field absent. Glabella nearly round, moderately inflated, length approximately maximum width at glabellar midlength; defined by narrow but deep axial and preglabellar furrows. Lateral glabellar furrows consist of two pairs; S1 long, weak to moderately impressed, directed inwardly and somewhat rearwardly, weakly bifurcated; $2 weak, transverse or directed inwardly and slightly forwardly; occipital furrow relatively shallow, curved rearward gently; occipital ring crescentic, widest sagittally, narrowing abaxially, bearing median node located slightly forward of •

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center, and short, stout median spine directed slightly upward. Preocular area of fixigena small, downsloping strongly. Palpebral area narrow, one-fourth as wide as glabella, moderately convex, downsloping strongly abaxially; palpebral ridge long, gently arcuate, defined by incised palpebral furrow, with anterior end slightly posterior to glabellar front, posterior end opposite S 1. Anterior branch of facial suture short, converging forward and directed strongly downward; posterior branch of facial suture straight, diverging rearward and downward, enclosing triangular posterolateral projection. Posterior border furrow wide and deep; posterior border narrow, of nearly uniform width. Pygidium semicircular, width about twice length. Axis strongly convex, with 3 tings and long, semielliptical terminal piece, reaching posterior border furrow. Pleural region gently convex, with four distinct pleural segments and small, obscurely segmented terminal area; each pleura beating ridge-like bands and deep and wide pleural furrow; pleurae defined by deep interpleural furrows that are somewhat narrower than pleural furrows. Lateral and posterior borders linear, separated from pleural region by wide, shallow border furrow. Posterior bands of pleurae connect with lateral margin, and anterior bands extending only to border furrow. Surface with fine, closely spaced granules on palpebral and posterior areas of fixigena, occipital ring, and sometimes posterior region of glabella; fine anastomosing ridges on palpebral and posterior areas of fixigena, and anterior and lateral regions of glabella. Remarks. Paradistazeris rotundus sp. nov. is characterized by having a proportionally large, rounded glabella whose anterior end nearly hangs over the preglabellar area; lateral glabellar furrows that are largely effaced; long, thin palpebral lobes defined well by incised palpebral furrows; strongly reduced preocular areas of the fixigenae; and a long pygidial axis that reaches to the border furrow. These features serve to distinguish the new species from all other species of Paradistazeris. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lowermost part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Paradistazeris sp.

Plate 68, figures 12, 13 2001c Distazeris sp., Peng, Babcock, Lin, Chen, and Zhu, p. 165, pl. 2, figs. 15, 16. Material. Meraspid and adult cranidia (NIGP 133513, 133514) in collection P[3-2.75. Remarks. Two cranidia collected from the uppermost part of the Glyptagnostus stolidotus Zone are the youngest Paradistazeris known from northwestern Hunan. Morphologically the two available cranidia are similar to Paradistazeris hunanensis and P. rotundus. However, P. hunanensis has a clearly defined anterior border, and P. rotundus has an anterior deflection in the area of the posterolateral comer; neither of these characters is present in the two specimens from the G. stolidotus Zone. Until more material is available, the two cranidia are left in open nomenclature. The large cranidium is badly broken, but shows a rounded glabella with three pairs of lateral glabellar furrows, all of which are isolated from the axial furrow. The S1 furrow is weakly bifurcated with a deep, rearward and slightly inwardly directed posterior branch, and a faint, short • 60

•

anterior branch. The $2 and $3 furrows are pit-like and faint. The anterior border is poorly separated from the preocular field of the fixigena. The small cranidium assigned to this species seems to be of an immature specimen. It has an elliptical glabella, and all of the lateral glabellar furrows are almost completely effaced. A short (sag., exs.) occipital ring extends rearward into a short spine, and a narrow fixigena with an angular posterolateral comer is present. The anterior border of the specimen appears is poorly defined. The palpebral lobe is small and located anteriorly.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs with trilobites indicative of the upper part of the Liostracina bella Zone (equivalent to the Glyptagnostus stolidotus Zone). Subfamily ONCHONOTININAE Lu, 1965

Remarks. Lu (in Lu et al., 1965, p. 218) erected the family Onchonotinidae to accommodate Onchonotina Lu in Lu and Qian, 1964 and Onchonotellus Lermontova in Ivshin, 1956 (p. 26-28). Peng (1992, p. 65) synonymized Onchonotina and Guotangia Zhou (1981, p. 243) with Onchonotellus. Onchonotellus was previously classified within the Solenopleuridae Angelin by Ivshin (1956) and Chernysheva (1960), but transferred to the Catillicephalidae by 0pik (1967, p. 206, 207). Opik (1967) described an Onchonotellus-like genus, Placosema, from Queensland, Australia, and erected a new family, Placosematidae, to accommodate the genus. The Placosematidae is here considered to be synonymous with the Onchonotinidae. This group of trilobites is classified here as a subfamily within the Catillicephalidae. Study of well preserved material of Huzhuia Chu from northwestern Hunan shows that it bears close similarity to both Onchonotellus and Placosema, and suggests that these genera constitute a group of catillicephalids with a moderately sized, ovate or subrectangular glabella, a wide (tr.) fixigena, converging anterior branches of the facial suture, and small, anteriorly or subcentrally located palpebral lobes. Morphologically, this group of trilobites differs from other such catillicephalid genera as Catillicephala, Buttsia, Distazeris, Pemphigaspis, and Madarocephalus. This group of trilobites has large, tumid, outward- or forward-expanded glabellas, narrow fixigenae, and rather posteriorly located, relatively large palpebral lobes. The subfamily Onchonotininae (including the concept of the Placosematidae as a junior synonym) is revived here to accommodate Onchonotellus, Placosema, and Huzhuia. Pratt (1992, p. 84) reassigned Placosema caelatum, the type species of Placosema, to Genevievella Lochman, 1936. This conclusion was evidently based largely on Genevievella simon Pratt (1992, pl. 32, figs. 12-21; text-fig. 32a), which differs from all other species of Genevievella in having a short preglabellar field, or lacking the preglabellar field altogether. This characteristic means that G. simon bears some similarity to Placosema. Genevievella differs from Placosema in having a forwardly tapering glabella, a narrower fixigena that is inclined inward, and a preglabellar field. Despite the similarities, and pending further study, it is probably best to consider Placosema and Genevievella as separate genera. Opik (1967) tentatively included Albansia Howell, 1937 in the Placosematidae. Here, Albansia is regarded as a probable catillicephalid, rather than as a representative of the Onchonotininae. Shergold (1980, p. 62) tentatively synonymized Seletella Ivshin (1962, p. 255-257) with Onchonotellus. Here, Seletella is regarded as a genus of the Catillicephalidae, and thus transfer to the Onchonotininae does not seem to be warranted.

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Genus HUZHUIAChu, 1965 Type species. Huzhuia typica Chu (Zhu, 1965, p. 141, 142, pl. 2, fig. 9), from the middle Cambrian at Xiaoshiwan, Huzhu, Qinghai; by original designation. Diagnosis. Onchotiniinae with cranidium subtrapezoidal to subsemicircular; glabella moderately convex, cylindrical to subcylindrical; lateral glabellar furrows largely effaced; S1, if present, is strongly oblique rearward; palpebral lobe located anteriorly; eye ridge weak; preglabellar field as wide as or shorter (sag.) than anterior border; fixigena somewhat convex with wide (tr., exs.) posterolateral projection. Other species. Solenoparia hsinkiangense Troedsson (1937, p. 66, 67, pl. 4, fig. 15) from the upper Cambrian, western Kuruktag, Xinjiang; Huzhuia paratypica Yang (1978, p. 40, pl. 6, fig. 5) from the Parablackwelderia jimaensis-Torifera tuma Assemblage-zone, Tingziguan, Fenghuang, northwestern Hunan; Huzhuia lajishanensis Chu and Zhang in Zhu, Lin, and Zhang (1979, p. 97, 98, pl. 39, figs. 9-11) from the Nidanshan Group, Lajishan, Huangzhong, Qinghai; Onchonotina tenuis Zhang (1981, p. 170, pl. 63, figs. 5, 6) from the Torsuqtagh Formation, Kuruktag, eastern Tianshan, Xinjiang; Huzhuia borohoroensis Xiang and Zhang (1985, p. 115, 116, pl. 37, figs. 5-9, 11 [=Huzhuia cf. H. longa Chu in Xiang and Zhang, 1985, p. 116, pl. 37, fig. 10]) from the Goniognostus nathorsti-Ptychagnostus punctuosus Zone at Guozigou, Hocheng, western Xinjiang; Huzhuia curvata sp. nov.; and Huzhuia latilimbata sp. nov. Onchonotellus kuruktagensis (Zhang) sensu Xiang and Zhang (1985, p. 117, pl. 39, figs. 2-4; non Zhang, 1981, p. 169, 170, pl. 63, figs. 2-4) and Onchonotellus jingheensis Xiang and Zhang (1985, p. 117, pl. 39, figs. 5-7), both from the Glyptagnostus reticulatus Zone at Keguqin, Jinghe, western Xinjiang, are tentatively transferred to Huzhuia. Huzhuia sp. (Zhang in Zhang et al., 1995, p. 29, pl. 6) should be excluded from the genus. The species is based on an indeterminate fragmental cranidium. However, the wide (sag.) preglabellar field and the strongly diverging anterior branches of the facial suture indicate that the specimen does not belong to Huzhuia. Remarks. The genetic concept of Zhu (1965, p. 147) is emended slightly to include species assigned subsequent to Zhu's (1965) work, including those species erected here. Just like Onchonotellus, Huzhuia was classified within the family Solenopleuridae when Zhu (1965) erected the genus, but this classification scheme no longer seems tenable. Huzhuia is relatively small in size, and closely comparable to Solenopleura subcincta Lermontova (1951, pl. 5, figs. 4, 5), the type species of Onchonotellus (see Opik, 1967, p. 206, 207 for discussion of validity), in the general respects; it also seems to be similar to other Onchonotellus species such as O. perlatus Invshin (1962, p. 121-124, text-fig. 33; Ergaliev, 1980, pl. 13, fig. 12), O. kuruktagensis Zhang (1981, pl. 63, figs. 2-4), O. globosa (Zhou et al., 1977, pl. 49, figs. 5, 6), and O. sp. cf. O. kuruktagensis Zhang (Peng, 1992, p. 65, 66, figs. 37, 39D-I). These species are all characterized by having short preglabellar fields. However, the more convex cranidium, the ovate glabella, the more convex fixigenae, the strongly converging anterior branches of the facial suture, and the relatively shorter (tr.) anterior border distinguish species of Ochonotellus from species of Huzhuia. Onchonotellus kuruktagensis (Zhang) sensu Xiang and Zhang (1985) and O. jingheensis Xiang and Zhang (1985) show characters that are transitional between Onchonotellus and Huzhuia. Stratigraphically these two species occur between the middle Cambrian Huzhuia and the Furongian .62-

(upper Cambrian, post-Irvingella-Zone) Onchonotellus. We tentatively refer these species to Huzhuia because they are morphologically more similar to that genus. Onchonotopsis? kobluki Pratt (1992, p. 74, 75, pl. 27, figs. 1-6) from the Rabbitkettle Formation, southern Mackenzie Mountains, Canada, bears close similarity to Huzhuia in having a parallel-sided, anteriorly rounded glabella, anteriorly situated palpebral lobes, and a short preglabellar field. It is tentatively considered to be a Laurentian representative of Huzhuia. Huzhuia longa Chu (Zhu, 1965, p. 142, pl. 2, figs. 10, 11) is evidently a junior synonym of Huzhuia typica. All specimens described for both species by Zhu (1965) from a single collection made near Qinghai, China, have a relatively low glabellar convexity and lack lateral glabellar furrows. Huzhuia jiuquanensis Zhou in Zhou et al. (1982, p. 240, pl. 61, figs. 14, 15), from the Germogou Formation (middle Cambrian) near Jiuquan, Gansu, has a relatively large, subquadrate cranidium with a gently convex, distinctly tapered, completely effaced glabella, a narrow (tr., exs.) fixigena, and relatively large palpebral lobes. These characters suggest that it does not conform to Huzhuia, and should be excluded from the genus. It may be referable to or closely related to Aphelaspis.

Huzhuia paratypica Yang, 1978 Plate 24, figures 1-17; Plate 25, figures 1-3; Plate 72, figure 2b 1963 1963 1965 1965 1978 1991

Solenoparia? sp., Egorova, Xiang, Li, Nan, and Guo, p. 37, pl. 6, fig. 15. Gen. et sp. indet., Egorova, Xiang, Li, Nan, and Guo, p. 37, pl. 11, fig. 18. Solenoparia? sp., Lu, Zhang, Zhu, Qian, and Xiang, p. 650, pl. 36, fig. 3. Gen. et sp. indet., Lu, Zhang, Zhu, Qian, and Xiang, p. 650, pl. 134, fig. 5. Huzhuia paratypica Yang, 1978, p. 40, pl. 6, fig. 5. Huzhuia paratypica Yang; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 145, pl. 15, fig. 6. 1993 Huzhuia paratypica Yang; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 187, pl. 15, fig. 6. 2001b Huzhuia longa Chu; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 10. Holotype. Cranidium (Yang, 1978, pl. 6, fig. 5, CUGB 0329210) from the Huaqiao Formation, Tingziguan, Fenghuang, northwestern Hunan. New material. More than 60 cranidia, and one librigena (illustrated specimens NIGP 138128138145, 138632) in collections P149.5, P156, P164.2, P171, P171.5, P178.3, P184, P223.7, P301, P319.6, P319.8, P331.8, W132.5, and W146.2. Emended diagnosis. Huzhuia with straight anterior border furrow on cranidium; glabella parallel-sided, front rounded, with three pairs of faint lateral furrows; S 1 strongly oblique rearward; palpebral lobe located anteriorly, opposite anterolateral comer of glabella; anterior branch of facial suture diverging slightly to converging slightly forward. Description. Cranidium subtrapezoidal, moderately convex, length slightly more than half maximum glabellar width at posterior border. Anterior border narrow, moderately convex, narrowing abaxially, defined posteriorly by deep, transverse anterior border furrow. Preglabellar field short (sag.), length subequal to anterior border, sloping forward moderately. Glabella •

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rectangular, convex, rounded to obtusely rounded anteriorly, occupying 75-80 percent of cranidial length; with three pairs of short lateral glabellar furrows; S 1 weakly incised, directed diagonally; $2 and $3 pitlike, faint. Occipital furrow deep, gently curved rearward; occipital ring convex, with weak but large median node. Eye ridge weak, transverse. Palpebral area about two-thirds as wide as glabella; posterior area large, convex, with distal part sloping downward; beating deeply incised posterior border furrow and narrow, convex posterior border; palpebral lobe small, located anteriorly, opposite anterolateral comer of glabella; palpebral furrow short, shallow. Anterior branches of facial suture short, converging forward gently; posterior branches curved outward, diverging rearward at angle of about 100°-110 °, enclosing long (exs.) subtriangular posterolateral projection. Librigena with relatively narrow, gently convex genal field and narrow, convex lateral border of uniform width.

Remarks. This species from the Huaqiao Formation, northwestern Hunan, resembles Huzhuia longa Chu, the type species of Huzhui, but differs in having a more convex glabella with clearly defined, strongly oblique S 1 furrows, and a more anteriorly located palpebral lobe. An ontogenetic series of the species shows that the glabella is expanded forward slightly and with an obtusely rounded front in the late meraspid or early holaspid stage. The glabella becomes parallel-sided in the early to mid-holaspid stage, and remains so through to the late holaspid stage. Occurrence. The holotype is from the Huaqiao Formation, Tingziguan, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Pianaspis sinensis Zone to the lower part of the Liostracina bella Zone (equivalent to the lower part of the Lejopyge laevigata Zone through the Linguagnostus reconditus Zone). Huzhuia cura,ata sp. nov. Plate 25, figures 7-15 2001b

Huzhuiasp.; Peng, Babcock, and Lin, p. 106, pl. 16, fig. 2.

Etymology. From Latin, curvatus, curved, referring to the forward-curved anterior border and border furrow. Holo~pe. Cranidium with broken occipital ring (P1. 25, figs. 8, 9, NIGP 138149) from collection P13 5.1. Other material. Four cranidia (NIGP 138148, 138150-138153) in collections P374.9 and P]3 5.1. Diagnosis. Huzhuia with anterior border of cranidium arched forward considerably; glabella tapered forward gently, obtusely rounded anteriorly, with transverse or gently concave apex; palpebral lobe situated immediately anterior to midpoint of glabella. Description. Cranidium subtrapezoidal, moderately convex, length two-thirds width. Anterior border narrow and wide (tr.), moderately convex, arched forward strongly, of almost uniform width; anterior border furrow distinct. Glabella long, occupying 85-90 percent of cranidial length, convex, • 64

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tapering forward gently, front obtusely rounded with apex straight or curved slightly rearward; with three pairs of short, nearly evenly-spaced lateral glabellar furrows; S1 long, straight, strongly oblique rearward, weakly to deeply incised; $2 short, parallel to S 1, weakly to deeply impressed, lying near glabellar midlength (excluding occipital ring); $3 pitlike, faint; occipital furrow deeply incised, gently curved rearward; occipital ring arched upward, nearly as wide (sag.) as L1 (exs.) medially, markedly narrower at sides. Palpebral lobe small, slightly deflected outward, located immediately forward of cranidial midpoint, opposite $2 or L3, defined by shallow palpebral furrow; eye ridge thin, faint, extending inward and forward from anterior end of palpebral lobe and meeting axial furrow about midpoint of frontal lobe of glabella. Preglabellar field slightly shorter (sag.) than anterior border, depressed; fixigena moderately convex, with preocular and palpebral areas about half as wide as glabella, and postocular area as wide as glabella; posterior border furrow deeply incised, widening abaxially, defining narrow, moderately convex posterior border. Anterior branches of facial suture subparallel; posterior branches nearly straight, diverging rearward at angle of about 60 ° . Remarks. All previously described species of Huzhuia have an anterior border that is transverse or curved forward gently and that is relatively shorter in transverse width due to convergence of the anterior branches of the facial suture. The strongly arched and relatively longer (sag.) anterior border distinguishes H. curvata sp. nov. from all other species of Huzhuia. Two cranidia described as Peishania sp. from the northern submontane belt, Turkestan (Repina et al., 1975, p. 159, pl. 25, figs. 1-3) are closely comparable to the new species. However, the Turkestani species is differentiated by having a highly effaced glabella that lacks lateral furrows, and has a more posteriorly located palpebral lobe. Minor intraspecific variation present in H. curvata. Variation exists in the curvature of the glabellar front, the position of the palpebral lobe, the proportional width (tr.) of the fixigenae, the relative prominence of the eye ridge, and the degree of effacement of the lateral glabellar furrows. Among the three species of Huzhuia that are present in the Huaqiao Formation of northwestern Hunan, Huzhuia curvata has the highest stratigraphic occurrence. In comparison with Huzhuia paratypica, which is an older species, evolutionary development of H. curvata involved rearward movement of the palpebral lobes, arching of the anterior border, change in the shape of the anterior branches of the facial suture (from converging forward in H. paratypica to subparallel in H. curvata), and a diminishing degree of divergence of the posterior branches of the facial suture. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs with trilobites indicative of the lower part of the Chuangia subquadrangulata Zone (equivalent to the lower part of the Glyptagnostus reticulatus Zone). Huzhuia latilimbata sp. nov. Plate 25, figures 4-6 Etymology. From Latin, latus, plus limbatus, wide border, referring to the wide preglabellar field. Holotype. Incomplete cranidium (P1.25, fig. 6, NIGP 138147) from collection P279.55. Paratype. Cranidium (NIGP 138146) in collection W 146.2.

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Diagnosis. Huzhuia with anterior border furrow arched forward, rather obscure; preglabellar field wide (sag.) and concave; eye ridge horizontal, faint; palpebral lobe opposite $3; anterior branches of facial suture parallel. Description. Cranidium about three-fifths as long as wide, with narrow, ridge-like anterior border that is gently curved forward, defined posteriorly by shallow, obscure anterior border furrow. Preglabellar field at least two times as long as anterior cranidial border. Glabella subovate, rounded to obtusely rounded anteriorly, with lateral furrows obscurely or weakly impressed inward; occipital furrow deeply incised, forwardly deflected distally; occipital ring moderately convex upward, bearing a prominent median node that is placed slightly anterior to the midpoint of occipital ring. Palpebral lobe small, located much posterior to preglabellar furrow, opposite L3; palpebral areas about two-thirds as wide as glabella. Anterior branches of facial suture subparallel; posterior branches gently curved, diverging rearward at angle of about 90 °- 120 °. Remarks. Only two reasonably well preserved cranidia are present in the collections from northwestern Hunan. This new species differs from all other species of Huzhuia except H. curvata in having an anterior border that is strongly arched forward and relatively wide (tr.); a poorly defined, anteriorly curved border furrow; and a relatively wide (sag., tr.) preglabellar field. H. curvata sp. nov. differs from H. latilimbata in the absence of a preglabellar field and the more posterior location of the palpebral lobe. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Pianaspis sinensis Zone to the lower part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Lejopyge laevigata Zone through the Proagnostus bulbus Zone). Genus ONCHONOTELLUS Lermontova in Ivshin, 1956

Onchonotellus Lermontova, 1951, p. 21; Ivshin, 1956, p. 26-28; 1962, p. 117; Chernysheva, 1960, p. 119, 120; Lazarenko, 1966, p. 65, 66; Opik, 1967, p. 206, 207; Rozova, 1968, p. 66; 1985, p. 117; Shergold, 1980, p. 61, 62; Peng, 1992, p. 65; Westrop, 1995, p. 36; Duan, Yang, and Shi, 1999, p. 159. ?Seletella Ivshin, 1962, p. 255-257. Onchonotina Lu in Lu and Qian, 1964, p. 34; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 218, 219; Lu, Zhu, Qian, Lin, Zhou, and Yuan, 1974, p. 87; Zhou, Liu, Meng, and Sun, 1977, p. 162; Liu, 1982, p. 311; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 110; Zhang, 1981, p. 169; Lu and Lin, 1989, p. 135. Guotangia Zhou, 1981, p. 243, 244. Onchonotina (Guotangia) Zhou; Lu and Lin, 1984, p. 97; Peng, 1984, p. 339. Type species. Solenopleura subcincta Lermontova, 1951 (p. 22-25, pl. 5, figs. 4, 5, 5a) from the Tortkuduk Effusive Suite, central Kazakhstan (Ivshin, 1962, p. 117), designated by Lermontova (in Ivshin, 1956, p. 26 ; see t3pik, 1967, p. 206, 207 for discussion of validity). Other species. Species described by 1980 were listed by Shergold (1980, p. 61, 62). To the list, the following species should be added: Onchonotina vigilans Lu in Lu and Qian, 1964 (p. 34, pl. 7, fig. 9) from the Xiyangshan Formation (upper Cambrian), Changshan, western Zhejiang; Onchonotina • 66

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globosa Zhou in Zhou et al., 1977 (p. 162, pl. 49, figs. 5, 6) from the upper Cambrian, Qidong, central Hunan; Onchonotina longiceps Liu in Zhou et al., 1977 (p. 162, pl. 49, figs. 7, 8) from the upper Cambrian, Huangshi, Taoyuan, northwestern Hunan; Guotangia guotangensis Zhou, 1981 (p. 244, pl. 1, figs. 8, 9) from the Guotan Formation (Lower Ordovician: Tremadocian), Sandu and Pu'an, eastern Guizhou; Onchonotina kuruktagensis Zhang, 1981 (p. 169, 170, pl. 63, figs. 2-4) from the Torsuqtagh Formation (upper Cambrian), Kuruktag, eastern Tianshan, Xinjiang; Onchonotellus jingheensis Xiang and Zhang, 1985 (p. 117, pl. 39, figs. 5-7) from the Glyptagnostus reticulatus Zone, Jiangjugou Formation, Jinghe, northern Tienshan, Xinjiang; Onchonotellus sp. cf. O. kuruktagensis (Zhang) sensu Peng (1992, p. 65-67, figs. 37, 39: D-I) from the Rhaptagnostus ciliensis-Onchonotellus cf. O. kuruktagensis Zone, Huaqiao Formation, Cili and Taoyuan, northwestern Hunan; Onchonotellus sp. (Peng, 1992, p. 67, fig. 39: B, C) from the Lotagnostus (Lotagnostus) punctatus-Hedinaspis regalis Zone, Shejiangwang, Cili, northwestern Hunan; Onchonotellus sp. indet. (Westrop, 1995, p. 36, pl. 15, figs. 12, 13) from the Noelaspis bilobata Zone, Rabbikettle Formation, Mountain River region, northern Mackenzie Mountains, Canada; and Onchonotellus curvitensus sp. nov. Onchonotina? taoyuanensis Zhou in Zhou et al., 1977 (p. 162, 163, pl. 49, fig. 9) from the upper Cambrian of Huangshi, Taoyuan, northwestern Hunan, is a junior synonym of O. longiceps Liu in Zhou et al., 1977. Remarks. Opik (1967) and Shergold (1980) discussed the taxonomic classification and nomenclatural problems associated with Onchonotellus. Peng (1992) suppressed Onchonotina Lu in Lu and Qian, 1964 and Guotangia Zhou, 1981, as junior synonyms of Onchonotellus. The genetic concept of Peng (1992) is followed here. Onchonotellus curvitensus sp. nov. Plate 26, figures 8-11 Etymology. From Latin, curvi, neck, combined with tensus, elongated, referring to the elongate occipital ring. Holotype. Cranidium (P1.26, figs. 8-10, NIGP 138159) from collection P317.4. Diagnosis. Onchonotellus with anterior border weakly defined; preglabellar field slightly shorter than anterior border; glabella relatively small, with elongate occipital ring occupying about onefourth of glabellar length; surface with scattered granules. Description. Cranidium subtrapezoidal, moderately convex, width two-thirds length. Anterior border wider than preglabellar field, gently convex; anterior border furrow shallow with medial portion weak, slightly deepened distally. Glabella ovate, convex, rounded anteriorly; lateral glabellar furrows consist of two faint pairs; S1 curved, extending inward and strongly rearward from axial furrow at about posterior one-third of glabellar length (excluding occipital ring), becoming rearward-directed distally, defining a narrow (tr.), lenticular L1; $2 short, lying slightly anterior to midlength of glabella, directed inwardly. Occipital ring long, tongue-like, moderately convex transversely and flat sagittally, with posterior part raised gently, occupying more than one-fourth of glabellar length; occipital furrow narrow but clearly defined, curving forward distally. Eye ridge long and faint, slanting rearward; palpebral lobe located subcentrally, moderate in size; palpebral area of fixigena about half glabellar width; anterior branch of facial suture converging -67-

forward; posterior branch of facial suture directed diagonally, enclosing subtriangular posterolateral projection that slopes downward strongly abaxially. Posterior border distinct; border furrow curving forward abaxially; posterior border moderately convex, widening distally. Surface covered with small scattered granules.

Remarks. In having a narrow preglabellar field, Onchonotellus curvitensus sp. nov. is most similar to Onchonotellus sp. cf. O. kuruktagensis from Cili and Taoyuan, northwestern Hunan. O. sp. cf. O. kuruktagensis is distinguished from O. curvitensus in having a strongly convex anterior cranidial border, a proportionally shorter (sag.) occipital ring, and densely spaced fine granules. By having a long, distinct occipital ring and by having a preglabellar field, the new species can be differentiated from all other previously described species of Onchonotellus. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Genus

PLACOSEMA

Opik, 1967

Type species. Placosema caelatum Opik, 1967 (p. 381,382, pl. 20, figs. 2-4) from the Erediaspis eretes and the Cyclagnostus [=Acmarhachis] quasivespa zones, Mungerbar Limestone, northwestern Queensland, Australia; by original designation. Other species. Placosema addnatum Opik, 1967 (p. 382, pl. 19, fig. 6; pl. 20, fig. 1), from the Cyclagnostus [=Acmarhachis] quasivespa and Glyptagnostus stolidotus zones, Georgina Limestone and Mungerbar Limestone, northwestern Queensland, and Placosema bigranulosum sp. nov. Remarks. The genetic concept of Opik (1967, p. 380) is followed here. Placosema bigranulosum sp. nov. Plate 26, figures 1-7 1996 Placosema sp.; Zhou, Cao, Hu, and Zhao, p. 44, pl. 3, fig. 13. 1996 Pagodia sp.; Cooper, Jago, and Begg (in part), p. 379, fig. 7I, J, non K, P-R. 2001 b Placosema sp.; Peng, Babcock, and Lin, p. 105, pl. 15, fig. 1.

Etymology. From Latin, bi, plus granulosus, doubly granular, referring to the bimodal size of granules present on the cranidium. Holotype. Cranidium (P1.26, figs. 3, 4, NIGP 138156) from collection P374.9. Other material. Three cranidia in collections P374.9 and P375. Additional material, including about 10 cranidia, are in collections P 375, P375.15, and P383.5. Diagnosis. Placosema with glabella that is bulbous; covered with large, widely spaced granules. • 68

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Description. Cranidium trapezoidal, width about twice length, convex. Anterior border narrow (sag.), convex; anterior border furrow deep, transverse. Glabella tumid, ovate, extending to anterior border, defined by broad and deep axial furrows; lateral glabellar furrows consist of three faint pairs; occipital furrow transverse, forward curving laterally, moderately well defined; occipital ring crescentic, posterior margin bowed rearward, with medial node. Eye ridge thin, distinct or weak, directed diagonally; palpebral lobe small, located subcentrally, defined by deeply incised palpebral furrow. Preocular area downsloping forward strongly; palpebral area half as wide as glabella. Anterior branch of facial suture converging forward; posterior branch of facial suture diverging rearward, enclosing triangular posterolateral projections. Posterior border furrow broad, deep; posterior border narrow. Surface of cranidium with granules of bimodal size; fine, densely spaced granules cover entire cranidium; coarse granules occur in regular pattern, with four pairs of coarse granules on glabella including one pair of coarse granules on occipital ring, and six pairs of coarse granules on fixigena, including one pair on the distal end of posterior border. Remarks. Placosema bigranulosum sp. nov. is distinguished from the two Australian species previously assigned to Placosema by the distinct, bimodal pattern of granulation. That granulation consists of fine granules coveting the cranidium together with three coarse granules on each side of the glabella, and six coarse granules on each fixigena. This ornamentation pattern, and the ovate glabella, suggest that P. bigranulosum has affinities with two specimens from North Victoria Land, Antarctica, which were referred to Pagodia by Cooper et al. (1996, fig. 7: I, J). The Antarctic specimens may be conspecific with those from Hunan. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the upper part of the Liostracina bella Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the lower part of the Glyptagnostus stolidotus Zone through the G. reticulatus Zone). Family

CREPICEPHALIDAE

Kobayashi, 1935

Genus CREPICEPHALINA Resser and Endo in Kobayashi, 1935

Type species. Crepicephalus convexus Walcott, 1911 (p. 79, 80, pl. 14, figs. 11, 1 l a) from the Crepicephalina Zone, Changhia Formation, Changxingdao Island, Liaoning, China; by original designation. Remarks. Zhang and Jell (1987) rediagnosed the genus and referred eight species to it. Their genetic concept is followed here. Crepicephalina pergranosa Resser and Endo, 1937 Plate 68, figure 11 1937 Crepicephalina pergranosa Resser and Endo, p. 196, pl. 37, fig. 23; pl. 45, figs. 30, 31; pl. 46, figs. 18-21. 1937 Crepicephalina mukdensis Resser and Endo, p. 197, pl. 33, fig. 3; pl. 45, figs. 32-34. 1937 Manchuriella mukdensis Resser and Endo, p. 245, pl. 36, fig. 22. 1937 Proasaphiscus kimurai Resser and Endo, p. 263, pl. 49, fig. 1. • 69

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1937 Proasaphiscus? mantouensis Resser and Endo (in part), p. 266, pl. 46, fig. 28, non fig. 29. 1937 Proasaphiscus? willis Resser and Endo, p. 267, pl. 46, fig. 27. 1965 Crepicephalina mukdensis Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 160, pl. 26, figs. 3-5. 1965 Crepicephalina pergranosa Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 160, pl. 26, figs. 6-9. 1965 Proasaphiscus willis Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 289, pl. 59, fig. 18. 1965 Proasaphiscus? kimurai Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 291, pl. 51, fig. 6. 1965 Proasaphiscus? mantouensis Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang (in part), p. 291, pl. 51, fig. 9, non fig. 10. 1965 Manchuriella mukdensis Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 300, pl. 53, fig. 18. 1987 Crepicephalina pergranosa Resser and Endo; Zhang and Jell, p. 83, 84, pl. 36, figs. 5-9; pl. 37, figs. 1-11; pl. 38, figs. 1, 2. 1996 Crepicephalina pergranosa Resser and Endo; Guo, Zan, and Luo, p. 100, 101, pl. 45, figs. 1-6, 8,9.

Holotype. Cranidium (Resser and Endo, 1937, pl. 37, fig. 23, USNM 86784; refigured by Zhang and Jell, 1987, pl. 38, fig. 1) from the Crepicephalina Zone, Changhia Formation, Changxingdao Island, southern Liaoning. Material. Mostly exfoliated cranidium (NIGP 138600) in collection W33.6. Remarks. Resser and Endo's (1937) type material was refigured by Zhang and Jell (1987, pl. 36, figs. 5-9; pl. 37, figs. 1-11; p. 38, figs. 1, 2), who suppressed four species referred to Crepicephalina, Manchuiella, or Proasaphiscus by Resser and Endo (1937) as its junior synonyms. The holotype is a well preserved cranidium showing densely spaced, fine granules on the surface. The new cranidium resembles the type material in all respects, including the presence of granules on testaceous specimens. This is the first record of C. pergranosa from South China; previously, it was known only from North China. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the lower part of the Dorypyge richthofenis Zone (equivalent to the Ptychagnostus atavus Zone). Crepicephalina sp. cf. C. quadrata Resser and Endo, 1937 Plate 18, figures 11, 12 cf. 1937 Crepicephalinaquadrata Resser and Endo, p. 197, 198, pl. 46, figs. 22, 23. cf. 1965 Crepicephalinaquadrata Resser and Endo; Lu, Zhang, Zhu, Qian, and Xiang, p. 160, pl. 26, figs. 10, 11. cf. 1987 Crepicephalinaquadrata Resser and Endo; Zhang and Jell, p. 84, pl. 37, figs. 12, 13. cf. 1996 Crepicephalinaquadrata Resser and Endo; Guo, Zan, and Luo, p. 101, pl. 45, fig. 11.

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Holotype of Crepicephalina quadrata. Cranidium (Resser and Endo, 1937, pl. 46, fig. 23, USNM 86789a; refigured by Zhang and Jell, 1987, pl. 37, fig. 12) from the Crepicephalina Zone, Changhia Formation, near Lujiafangshan, Liaoyang, Liaoning.

Material. One cranidium (NIGP 138071) in collection W 1.2. Remarks. Except for having slightly shorter, and more arcuate palpebral lobes, the illustrated cranidium appears to be identical in all aspects with C. quadrata Resser and Endo (1937) from the Crepicephalina Zone of Changhia Formation in Liaoyang, Liaoning. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Crepicephalina? sp. Plate 1, figures 9-11

Material. One crushed cranidium (NIGP 137904) in collection P45.6. Remarks. A single cranidium is tentatively referred to Crepicephalina because it resembles some other cranidia of Crepicephalina described from the North China Platform (Zhang and Jell, 1987). The subtrapezoidal glabella with four pairs of shallow lateral furrows and an obtusely rounded front; the narrow preglabellar field; the convex anterior border; the narrow (exs.) posterior areas of fixigenae; and the scattered granules on the fixigenae, preglabellar field and anterior border are all consistent with Crepicephalina. However, the new cranidium differs from species of Crepicephalina in having proportionally shorter palpebral lobes. The species is tentatively referred in Crepicephalina also because it lacks an associated pygidium that can be used for comparison. In Crepicephalina, the pygidium is characterized by beating a pair of posterolateral spines.

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Genus METEORASPIS Resser, 1935

Meteoraspis Resser, 1935, p. 40, 41; Lochman, 1938b, p. 472; Shaw, 1952, p. 475; Palmer, 1954, p. 753; Deland and Shaw, 1956, p. 557; Lochman-Balk in Moore, 1959, p. 250; Rasetti, 1961, p. 116; Lochman and Hu, 1961, p. 141, Rasetti, 1965, p. 54; Yuan and Yin, 1998, p. 145. Coleopachys Raymond (in part), 1937, p. 1120. Greylockia Raymond, 1937, p. 1108.

Type species. Ptychoparia? metra Walcott (1890, p. 273, pl. 21, fig. 7) from the upper Cambrian at Potatotop, Burnet, and Packsaddle Mountain, Llano, Texas; by original designation. Remarks. The genetic diagnosis of Resser (1935, p. 40, 41) was based on a single, poorly preserved .71-

cranidium that was referred questionably to Ptychoparia by Walcott (1890). Shaw (1952, p. 475) synonymized Greylockia Raymond (1937) and Coleopachys Raymond (1937) as junior synonyms of Meteoraspis. Palmer (1954, p. 753) expanded the genetic concept by emending the genetic diagnosis. Palmer's (1954) concept is followed here.

Meteoraspis sp. cf. M. orientalis Yuan and Yin, 1998 Plate 1, figures 1-8 cf. 1996 cf. 1998 200 lb

Meteoraspis sp.; Zhou, Cao, Zhao, and Hu, p. 44, pl. 3, figs. 15, 16. Meteoraspis orientalis Yuan and Yin, p. 145, pl. 6, figs. 6a, b. Meteoraspis sp., Peng, Babcock, and Lin, p. 104, pl. 10, fig. 16.

Holotype of Meteoraspis orientalis. Cranidium (Yuan and Yin, 1998, pl. 6, fig. 6, NIGP 127970) from the Glyptagnostus reticulatus Zone, Huaqiao Formation, Jimachong, Wanshan, eastern Guizhou. Material. Single incomplete cephalon, single incomplete cranidium, and single librigena (NIGP 137901-137903) in collections P317.4, and PI3 70.7. Description. Glabella gently tapering forward, moderately convex, rounded to obtusely rounded anteriorly, with three pairs of faintly depressed lateral glabellar furrows; S1 bifurcate with long, rearward and inward directed posterior branch and short forward and inward directed anterior branch; $2 transverse, located in anterior one-third of glabellar length; $3 obscure. Occipital ring furrow broad and deep; occipital ring moderately convex (tr., sag.), evenly wide medially, narrowing distally, with a large, poorly defined median node adjacent to occipital furrow. Preglabellar field nearly as wide (sag.) as anterior border, anterior border furrow beating three pits. Eye ridge faint; palpebral lobe crescentic, moderately long, lying about cranidial midlength; palpebral area of fixigena narrow, gently convex; posterolateral projection triangular, downsloping strongly abaxially, posterior border furrow deeply incised, widening abaxially. Anterior branch of facial suture diverging slightly forward, curving gently inward after crossing border furrow; posterior branch diverging rearward to meet posterior border furrow, then extending subparallel to rear. Librigena with moderately wide, gently convex genal field defined by broad and shallow border furrows; lateral border wide (tr.), long; posterior border wide (tr.), short (tr.); genal spine short, broad at base. External surface and internal mold covered with granules varying from coarse to bimodal (fine, dense, and coarse, scattered). Remarks. This species from western Hunan is compared to M. orientalis (Yuan and Yin, 1998, pl. 6, figs. 6a, b) from eastern Guizhou. The new material and the holotype of the M. orientalis occur in the same paleogeographic region. In general respects including the convexity of the cranidium, the presence of three pits on the anterior border furrow, the size and position of the palpebral lobes, and the presence of granulate ornamentation, this taxon is identical with the holotype. Differences between the Hunan and Guizhou specimens are apparent, however: M. orientalis has a shorter, broader, more strongly tapered glabella; a shorter (sag.) preglabellar field; a narrower (tr.) palpebral area; and a much wider (tr.) posterolateral projection. Both M. orientalis from Guizhou and the specimens from Hunan are in small collection, and •

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more specimens are needed to clarify whether the differences that have been observed represent intraspecific variation or are of specific significance.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs with trilobites indicative of the lower part of the Liostracina bella Zone to the upper part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus stolidotus Zone through the G. reticulatus Zone). Family DICERATOCEPHALIDAELu, 1954

Remarks. Zhang and Jell (1987, p. 131) discussed the concept of Diceratocephalidae Lu, and suppressed Aulacodigmatidae Opik, 1967 as a junior synonym. Except for Diceratocephalus Lu, Zhang and Jell (1978) referred five genera, Cyclolorenzella Kobayashi (1960b), Aulacodigma Opik (1967), Torifera Wolfart (1974), Fenghuangella Yang (1978), and Xiangia Peng (1987) to the family. The concept of Zhang and Jell (1978) is followed here, except that Torifera and Xiangia is excluded from the family. Xiangia is considered to be synonymous with Torifera. Genus FENGHUANGELLAYang in Zhou et al., 1977

Fenghuangella Yang in Zhou et al., 1977, p. 169, 170; Yin and Li, 1978, p. 489; Yang, 1978, p. 44; 1982, Liu, 1982, p. 312; Luo, 1982, p. 4; Lin, Lin, and Zhou, 1983, p. 403; Qian and Zhou, 1984, p. 178; Peng, 1987, p. 97. Cyclolorezellina Ergaliev, 1980, p. 137. Type species. Fenghuangella laochatianensis Yang in Zhou et al. (1977, p. 170, pl. 50, fig. 20) from the Huaqiao Formation, Laochatian of Fenghuang and Jiudiantang of Xinhuang, western Hunan, and Jimachong of Yuping, eastern Guizhou; designated subsequently by Yang (1978, p. 44). Other species. Fenghuangella coniforma Yang in Zhou et al. (1977, p. 170, 171, pl. 50, fig. 2 l) and Fenghuangella liostracinala Yang in Zhou et al. (1977, p. 171, pl. 50, figs. 22, 23), both from the Huaqiao Formation in Fenghuang and Xinhuang, western Hunan, and Yuping, eastern Guizhou; Fenghuangella sp. (Luo, 1982, p. 4, pl. 1, fig. 11) and Cyclolorenzella sp. 1 (Luo, 1982, p. 3, pl. 1, figs. 4-6), both from the Hetaoping Formation, near Baoshan, western Yunnan; Fenghuangella fusilis sp. nov., and Fenghuangella laochatianensis crassa subsp, nov. Fenghuangella modesta (Zhou, Liu, Meng, and Sun, 1977, p. 171, pl. 50, figs. 24, 25), F. magnispina (Lin, Lin, and Zhou, 1983, p. 403, 404, pl. 2, figs. 3-5), F. striate (Qian and Zhou, 1984, p. 178, 179, pl. 2, figs. 8, 9), and F. lata (Qian and Zhou, 1984, p. 178, pl. 2, fig. 10) are suppressed as junior synonyms of F. laochatianensis, whereas F. subtriangularis (Lin, Lin, and Zhou, p. 403, pl. 2, figs. 1, 2) is a junior synonym of F. coniforma. Two species described under the names Cyclolorenzella caijiapingensis Yang (Yin and Li, 1978, p. 489, pl. 165, fig. 19) from western Hunan and Cyclolorenzellina coniformis Ergaliev from Kazakhstan, (Ergaliev,1980, p. 138, pl. 8, fig. 14) are regarded as junior synonyms of F. coniforma and F. liostracinala respectively. Remarks. The concept of Fenghuangella and the differences between Fenghuangella and Cyclolorenzella Kobayashi (1960b, p. 389, 390) were reviewed by Peng (1987), and the genetic • 73"

concepts are followed here. Cyclolorenzella differs from Fenghuangella in having a truncateconical glabella, a large posteriorly located palpebral lobe, a posteriorly oblique eye ridge, and a narrow (tr., exs.) posterior area of the fixigena. Fenghuangella is comparable to Torifera Wolfart (1974, p. 97, 98), but Torifera has an anterior border on the cephalon. Cyclolorenzellina and its type species Cyclolorenzellina coniformis Ergaliev (1980, p. 138, pl. 8, fig. 14) from Malyi Karatau, Kazakhstan, were suppressed by Peng (1987, p. 97) as junior synonyms of Fenghuangella and Fenghuangella liostracinala Yang in Zhou et al. (1977, p. 171, pl. 50, figs. 22, 23), respectively. Although the genetic concept for Fenghuangella is clear enough, some species referred to the genus are basically indeterminate, as most of them are illustrated with poor photographs. Type specimens of most described species will need to be refigured in order to clarify the various species concepts. Fenghuangella laochatianensis laochatianensis Yang in Zhou et al., 1977 Plate 27, figures 1-15; Text-figure 7 1977 1977 1978 1978 1982 1982 1983 1984 1984 1987 1991

Fenghuangella laochatianensis Yang in Zhou et al., p. 170, pl. 50, fig. 20. Fenghuangella modesta Zhou in Zhou et al., p. 171, pl. 50, figs. 24, 25. Fenghuangella laochatianensis Yang; Yin and Li, p. 489, pl. 165, fig. 20. Fenghuangella laochatianensis Yang; Yang, p. 44, pl. 7, figs. 12, 13. Fenghuangella laochatianensis Yang; Liu, p. 312, pl. 219, fig. 14. Fenghuangella modesta Zhou; Liu, p. 312, pl. 219, figs. 16, 19. Fenghuangella magnispina Lin in Lin et al., p. 403,404, pl. 2, figs. 3-5. Fenghuangella striate Qian and Zhou, p. 178, 179, pl. 2, figs. 8, 9. Fenghuangella lata Qian and Zhou, p. 178, pl. 2, fig. 10. Fenghuangella laochatianensis Yang; Peng, p. 97, pl. 7, figs. 6-9. Fenghuangella cf. F. modesta Zhou; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 147, pl. 15, fig. 12. 1993 Fenghuangella sp. cf. F. modesta Zhou; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 189, pl. 15, fig. 12. ?1999 Fenghuangella laochatianensis Yang; Duan, Yang, and Shi, p. 158, fig. 13c. 2001b Fenghuangella laochatianensis Yang; Peng, Babcock, and Lin, p. 104, pl. 10, fig. 4. 2001d Fenghuangella magnispina Lin; Peng, Babcock, Lin and Chen, p. 141, fig. 9.4. Lectotype. Testaceous cranidium (Zhou et al., 1977, pl. 50, fig. 20, CUGB 0324304; Text-fig. 7 herein) from the Huaqiao Formation, Laochatian, Fenghuang, western Hunan. The lectotype was assigned as the holotype by Yang (1978) and subsequently split and damaged. New material. More than 50 sclerites, including cephala, cranidia, and pygidia (illustrated specimens NIGP 138160-138168), in collections P308, P319.6, P331.8, P337.5, P. 348, W210.5, W218.3, W221.5, W227, and W228.3. Emended diagnosis. Fenghuangella with semicicular to transverse glabella, moderately elongate, tapered unevenly forward, gently in posterior two-thirds, tapered strongly in anterior one-third; preglabellar area beating paired short, oblique, closely spaced furrows posteriorly; fixigena broad (tr.), narrow (tr.) librigena with short genal spine; surface partly granulated.

• 74.

Text-figure 7. A-F, Fenghuangella laochatianensis laochatianensis Yang in Zhou et al., 1977. A, B, cranidium designed subsequently as holotype of the species in Yang, 1978 (p. 44), anterolateral and dorsal views, CUGB 0324304, both x 16, (original of Zhou et al., 1977, pl. 50, fig. 20; also Yang, 1978, pl. 7, fig. 12, and damaged after photograph was taken in 1977), rephotographed in 2002; C, replica from a latex cast of the holotype made in 1980, x 16; D-F, paratype cranidium, CUGB 1206202, x 16, x 16, x 28, (original of Yang, 1978, pl. 7, fig. 3), D, replica from the latex cast of the specimen; E, composite photo with the left half being a mirror image of the fight side; F, enlargement of posterolateral part of left fixigena, showing granulate ornamentation. Remarks. This is the type species of F e n g h u a n g e l l a , and the type material is from the Huaqiao

Formation at Laochatian, Fenghuang. The type species of this small trilobite had been illustrated previously four times, but always with small photographs. For this reason, the concept of the species has remained obscure. Here, the holotype has been refigured with a large photograph (Text-fig. 7). The holotype cranidium was damaged since it was first illustrated. However, the •

75

•

specimen clearly shows a subconical glabella with flanks curved slightly outward, and a short but deep S 1 furrow that is directed rearward and inward. The fixigena is broad, downward-curving, and has undivided preglabellar and preocular areas, together with a weak eye ridge. The occipital ring is wider (tr.) than the base of the glabella and bears a short but wide occipital spine. Most of the fixigena is smooth, but fine granules are present on the inner parts of the postocular areas. Similar ornamentation is also present on the paratype cranidium. Reinvestigation of the type material and study of large collections of the species from Paibi and Wangcun make it possible to emend the specific diagnosis and to synonymize some species described either from Fenghuang, western Hunan (Zhou in Zhou et al., 1977), or from Kunshan, southern Jiangsu (Lin et al., 1983; Qian and Zhou, 1984). Both of these localities are located within the Jiangnan Slope Belt of South China.

Occurrence. The lectotype is from the Huaqiao Formation at Laochatian, Fenghuang, western Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone through the upper part of the G. stolidotus Zone). Fenghuangella laochatianensis crassa subsp, nov. Plate 27, figure 16; Plate 28, figures l-15 2001b Fenghuangella sp.; Peng, Babcock, and Lin, p. 105, pl. 15, figs. 5, 6.

Etymology. From Latin, crassus, thick, dense, coarse-grained, referring to the dense, coarse-grained ornamentation of granules. Holotype. Testaceous cranidium (P1.28, fig. 3, NIGP 138172) from collection P378.25. Paratypes. One cephalon, seven cranidia and one pygidium (NIGP 138170, 138171, 138173138179) in collections P378.25, W211.7, W211.45 and W228.3. Diagnosis. Fengwangella laochatianensis with palpebral lobes and eye relatively small and weak; surface granulation relatively coarse, dense. Description. Cephalon semicircular, length about half width, with obtusely angular genal angle. Cranidium transverse-subtriangular. Anterior area convex, sloping forward anteriorly, beating pair of incomplete preglabellar furrows. Glabella conical, angular, or obtusely angular anteriorly, with three pairs of short, inwardly directed lateral glabellar furrows. Occipital furrow deep; occipital ring subtriangular, with stout spine medially. Axial furrow wide, deep. Eye ridge transverse, faint to effaced. Posterior area of fixigena large, moderately convex, outer part sloping steeply downward. Surface with relatively coarse granules; spacing of granules dense in inner part of fixigena, becoming less dense abaxially and forwardly. Remarks. Some material from Hunan is recognized as a new subspecies of Fenghuangella laochatianensis because it differs from all other specimens referred to the species in having coarser and denser granulation, smaller and weaker palpebral lobes and eyes, and probably in lacking genal •

76.

spines. The new subspecies is the youngest Fenghuangella with a stratigraphical occurrence from the middle of the Linguagnostus reconditus Zone to the middle of the Glyptagnostus reticulatus Zone.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the Linguagnostus reconditus Zone through the G. reticulatus Zone). Fenghuangella coniforma Yang in Zhou et al., 1977 Plate 29, figures 1-17; Text-figure 8 1977 Fenghuangella coniforma Yang in Zhou et al., p. 170, 171, pl. 50, fig. 21. 1978 Fenghuangella coniforma Yang; Yin and Li, p. 490, pl. 164, fig. 12. 1978 Cyclolorenzella caijiapingensis Yang; Yin and Li, p. 489, pl. 165, fig. 19. 1978 Fenghuangella coniforma Yang; Yang, p. 45, pl. 7, figs. 15, 16. 1978 Cyclolorenzella caijiapingensis Yang; Yang, p. 43, pl. 7, fig. 8. 1982 Fenghuangella coniforma Yang; Liu, p. 312, pl. 219, fig. 20. 1983 Fenghuangella subtriangularis Lin and Zhou in Lin et al., p. 403, pl. 2, figs. 1, 2. 1987 Fenghuangella magnispina Lin; Peng, p. 98, pl. 7, figs. 10, 11. 1987 Fenghuangella coniforma Yang; Peng, p. 97, 98, pl. 7, fig. 12. 1988 Fenghuangella subtriangularis Lin and Zhou; Zhu, Lin, and Zhang, p. 79, pl. 9, fig. 4. 2001b Fenghuangella coniforma Yang; Peng, Babcock, and Lin, p. 104, pl. 9, figs. 9, 10. non 2001b Fenghuangella coniforma Yang; Peng, Babcock, and Lin, p. 105, pl. 14, figs. 8, 9 [=Fenghuangella fusilis sp. nov.].

Holotype. By monotypy; exfoliated cranidium (Zhou et al., 1977, pl. 50, fig. 21, CUGB 0231006; refigured by Yang, 1978, pl. 7, fig. 16; Text-fig. 8A, B, herein) from the Huaqiao Formation, Laochatian, Fenghuang, western Hunan. New material. More than 50 sclerites, including cranidia, and pygidia (illustrated specimens NIGP 138180-138194), in collections P278, P293, W188.8, W196.3, W197.55, W199.2, W216.5. Remarks. The type series includes two specimens, both of which are nearly completely exfoliated. Only one cranidium, which is from western Hunan, was figured in the original publication of the species (Yang in Zhou et al., 1977, pl. 50, fig. 21). Subsequently, Yang (1978) designated the second cranidium from eastern Guizhou as the holotype and referred to the first published cranidium as the paratype of the species. Considering that the species became valid in 1977, the first-published cranidium should be regarded as the holotype of the species. New material from the Huaqiao Formation of western Hunan resembles the type material from the same region. This species has a subtriangular cranidium; strong eye ridges that project anterolaterally or run horizontally; a small, poorly defined palpebral lobe located on the anterolateral area of the cranidium; a conical glabella; a narrow preglabellar area; relatively coarse granules on the glabella and occipital ring, the inner part of the fixigena, and the basal part of the preglabellar area; a genal spine on the librigena; and terrace lines on the anterior part of the preocular area. • 77-

Text-figure 8. Fenghuangella coniforma Yang in Zhou et ai., 1977. A, B, holotype cranidium in dorsal and anterolateral views, mostly exfoliated, CUGB 0231006, both x 8, (original of Zhou et al., 1977, pl. 50, fig. 16; also Yang, 1978, pl. 7, fig. 16): C, D, mostly exfoliated cranidium that was invalidly designed as holotype of the species in Yang, 1978 (p. 45, 79), CUGB 1206107, x 14, (original of Yang, 1978, pl. 7, fig. 15); E, F, a slightly exfoliated cranidium, CUGB 0328201, x 13. This specimen was designed as the holotype of Cyclolorenzella caijiapingensis Yang in Yin and Li, 1978 (pl. 165, fig. 19; figured also in Yang, 1978. pl. 7, fig. 8). In having paired furrows on the preglabellar area, this species resembles F. laochatianensis. F. coniforma differs from F. liostracinala in having paired furrows on the pregrabellar area rather than a single furrow (an incomplete median preglabellar furrow). Yang (in Zhou et al., 1977) differentiated F. coniforma from F. laochatianensis by the shape of glabella and by the absence of an occipital spine, but examination of the type material reveals that the occipital spine is broken out in the two specimens assigned to F. coniforma (see Text-fig. 8, herein). Thus, the glabellar shape becomes the only difference between these two species. New material demonstrates that F. coniforma may be distinguished by having a proportionately narrower and longer cranidium, a relatively narrower (tr.) palpebral area of the fixigena, more widely spaced preglabellar furrows, • 78"

and a thicker exoskeleton on the cranidium. The meraspid cranidium referred here to F. coniforma is not readily distinguishable from holaspides of F. laochatianensis. Occurrence. The holotype is from the Huaqiao Formation, Laochatian, Fenghuang, western Hunan China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone through the lower part of the Linguagnostus reconditus Zone). Fenghuangella liostracinala Yang in Zhou et al., 1977

Plate 30, figures 9-15" Text-figure 9 Liostracina krausei Monke; Egorova, Xiang, Li, Nan, and Guo (in part), p. 42, 43, pl. 8, fig. 6 only. 1977 Fenghuangella liostracinala Yang in Zhou et al., p. 171, pl. 50, figs. 22, 23. 1978 Fenghuangella liostracinala Yang; Yin and Li, p. 490, pl. 164, fig. 13. 1978 Fenghuangella paraconiforma Yang; Yin and Li, p. 490, pl. 165, fig. 128. 1978 Fenghuangella liostracinala Yang; Yang, p. 44, 45, pl. 7, fig. 14. 1978 Fenghuangella paraconiforma Yang; Yang, p. 46, pl. 7, figs. 17, 18. 1980 Cyclolorenzellina coniformis Ergaliev, p. 138, pl. 8, fig. 14. 1982 Fenghuangella liostracinala Yang; Liu, p. 312, pl. 219, fig. 30. 2001c Fenghuangella liostracinala Yang; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 2, figs. 17, 18. 2001b Fenghuangella liostracinala Yang; Peng, Babcock, and Lin, p. 105, pl. 14, fig. 10.

1963

Lectotype. Partly exfoliated cranidium (Zhou et al., 1977, pl. 50, fig. 22, CUGB 0324203; refigured by Yang, 1978, pl. 7, fig. 14 and by Yun and Li, 1978, pl. 164, fig. 13; Text-fig. 9 herein) from the Huaqiao Formation, Laochatian, Fenghuang, western Hunan; assigned by Yang (1978). New material. One incomplete exoskeleton and more than 20 cranidia (illustrated specimens NIGP 138199-138202) in collections P353.7, P363.5, P367.76, P368.0, W243.6, W251.15, P13-2.75, and P]55.1. Remarks. New material from the Huaqiao Formation, western Hunan, resembles material described or figured previously from westem Hunan (Zhou et al., 1977; Yang, 1978; Peng et al., 2001b, 2001c). This species has just one preglabellar furrow. Other key characters include a semicircular, basically transverse, cranidium; a gently tapered, anteriorly sharply-pointed glabella, faint eye ridges that project anterolaterally; wide fixigenae with few granules on the posterolateral areas including the posterior border. New material shows that the thorax consists of 8 or possibly even 9 segments. It is densely granulated with a markedly wide axis, transverse, ridge-like pleural bands with anterior bands that the slightly narrower (exs.) than the posterior ones, and moderately deep pleural furrows. Occurrence. The lectotype is from the Huaqiao Formation, Laochatian, Fenghuang, western Hunan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2, and • 79"

Wangcun sections, Hunan, where it occurs with trilobites indicative of the upper part of the Liostracina bella Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the upper part of the Linguagnostus reconditus Zone through the upper part of the Glyptagnostus reticulatus Zone).

Text-figure 9. Lectotype of Fenghuangella liostracinala Yang in Zhou et al., 1977, partly exfoliated cranidium in dorsal and anterolateral views, CUGB 0324203, × 20 (original of Zhou et al., 1977, pl. 50, fig. 22; also Yang, 1978, pl. 7, fig. 14).

Fenghuangella fusilis sp. nov. Plate 30, figures 1-8

2001b Fenghuangella coniforma Yang; Peng, Babcock, and Lin, p. 105, pl. 14, figs. 8, 9. Etymology. From Latin, fusilis, fused, melted, referring to the fusion of the glabellar anterior lobe and the preglabellar area. Holotype. Testaceous cranidium (P1.30, figs. 4-7, NIGP 138198) from collection W254.1, from the Huaqiao Formation, Wangcun, western Hunan. Other material. Three testaceous cranidia (P1. 30, figs. 1-3; NIGP 138195-138197), all in collection W254.1. Diagnosis. Fenghuangella with a conical glabella that is fused with preglabellar area anteriorly, and single, pit-like preglabellar furrow; surface smooth. Description. Cranidium subtriangular, length three-fifths width (except occipital spine), obtusely angled anteriorly. Glabella conical, pointed and fused with preglabellar area anteriorly, with two pairs of lateral furrows: S 1 short, directed diagonally; $2 short, impressed inwardly; occipital ring furrow shallow or faint medially, deeper laterally; occipital ring wider (tr.) than glabellar basal lobe, with short spine sagittally. Eye ridge effaced almost completely, transverse or slightly oblique forward; palpebral lobe small, located at level of fused area of glabella and preglabellar area. Anterior branch of the facial suture curving inward and forward to meet anterior margin sagittally; posterior branch curving outward and rearward from palpebral lobe. Posterior border furrow transverse, merging into fixigena about half way from axial furrow to genal comer. • 80-

Remarks. This species is distinguished from all other species of Fenghuangella by having the glabella and preglabellar area fused, and by the absence of granules. It differs further from F. coniforma in having a single median preglabellar furrow, and from F. laochatianensis and F. liostracinala by having a conical glabella, and a less transverse cranidial outline. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the lower part of the Glyptagnostus reticulatus Zone). Family

DOKIMOCEPHALIDAE

Kobayashi, 1935

Subfamily CONOKEPHALININAEHup6, 1955

Remarks. Opik (1967, p. 247, 248) revived Lobocephalina Rfi2i6ka, 1940, which was placed in synonymy with Conokephalina Brcgger, 1878 by Snajdr (1958), and classified with Conokephalina and Wuhuia Kobayashi, 1935 within the Conokephalinidae. These three genera have large palpebral lobes lying close to the axial furrows. Conokephalina is distinguished by a broad, slightly tapering glabella (BrCgger, 1878, pl. 3, figs. 5, 5a; also Kobayashi, 1962, pl. 9, fig. 18); Lobocephalina is characterized by a pyriform glabella; and Wuhuia is characterized by a rather tapered glabella with straight sides. As noted by Opik (1967), the Treatise illustration of Conokephalina (Lochman-Balk in Moore, 1959, fig. 176.1a, b) led to misconceptions about the concept of Conokephalina. The Wuhuiinae was erected to accommodate Wuhuia and related species of the AustralianAsiatic Dokimocephalidae (Shergold, 1980), including Lorrettina Shergold, 1972, Protemnites Henderson, 1976, and Chalfontia Shergold, 1982. Saimachia Kobayashi and Prismenaspis Henderson, 1976 were also included in the Wuhuiinae, but Saimachia was regarded by Zhang and Jell (1987, p. 141) as belonging to the Proasaphiscidae, and Prismenaspis was regarded by Shergold (1982, p. 43) as a junior synonym of Protemnites. Wuhuia, Lorrettina, Protemnites, and Chalfontia are closely similar to Lobocephalina. These genera occur in Australia and South China, and are probably related phylogenetically. We prefer to regard Conokephalininae and Wuhuiinae as synonymous, and to transfer Lorrettina, Protemnites and Chalfontia to the Conokephalininae. Genus

LOBOCEPHALINA

Rfi2i6ka, 1940

Type species. Conokephalites emmrichi Barrande, 1846 (Barrande, 1852, p. 428, pl. 11, figs. 2-6; revised by Snajdr, 1958) from the Jince Beds, Jince, Bohemia; by original designation. Other species. Lobocephalina pyriceps Opik (1967, p. 248-250, pl. 30, figs. 1-3, text-fig. 85) from the Glyptagnostus stolidotus Zone of the O'Hara Shale, northwestern Queensland, Australia, and Lobocephalina sinensis sp. nov. Remarks. Opik (1967, p. 248) discussed the genus at length. His genetic concept is followed here.

-81"

Lobocephalina sinensis sp. nov. Plate 31, figures 1-10

200 lb Cern~taspissp. cf. C. abundans Opik, 1967, Peng, Babcock, and Lin, p. 104, pl. 11, fig. 7. Etymology. From Greek, Sina, China. Holotype. Cranidium (P1.31, figs. 2, 3, 7, NIGP 138204) from collection P319.6. Other material. Three cranidia and one pygidium (NIGP 124312, 138203, 138205, 138206) in collections P319.8, W219.7, W228.3 and W254.1 are figured paratypes; other fragments are in collections W219.7 and W228.3. Diagnosis. Lobocephalina with anterior cranidial border that is wide (sag.), posteriorly depressed, with short plectrum extending rearward onto glabella, and narrow preglabellar field. Glabella pyriform with lateral furrows almost obliterated. Surface densely punctuate. Description. Cranidium longer than wide. Anterior border wide, gently convex anteriorly and concave posterior, beating short poorly defined plectrum on posterior margin; anterior border furrow shallow, transverse or gently arched forward, interrupted sagittally by plectrum; preglabellar field narrow. Glabella pyriform, gently convex with posterior half subparallel-sided and anterior half rather tapered, obtusely rounded anteriorly; lateral glabellar furrows faint on testaceous surface, shallow on internal mold; S1 bifurcated with posterior branch directed diagonally and anterior branch directed inward and forward; $2 nearly transverse; $3 slightly oblique forward; occipital furrow transverse, moderately deep medially, becoming narrower, shallower and forwardly deflected distally; occipital ring narrowed slightly distally, with poorly defined median node. Fixigena with narrow (tr.) palpebral area and crescentic, triangular, preocular areas. Palpebral lobe large, flat, gently curved, lying close to axial furrow, opposite midpoint of glabella, anterior and posterior ends almost touching the axial furrow; eye ridge short, obscure; palpebral furrow slender, shallow. Anterior branch of facial suture diverging forward strongly about 40 ° to sagittal line, posterior branch straight, diverging rearward strongly, enclosing transverse triangular posterolateral projection that almost equals posterior area of fixigena. Posterior border wider (exs.) than posterior field of fixigena, widening slightly abaxially; posterior border a linear ridge. Pygidium semicircular, width about twice length, with transverse anterior margin, and facet sloping outward and forward. Axis subcylindrical, tapering gently rearward, occupying three-fourths pygidial length, extending rearward as postaxial ridge to posterior border furrow. Pleural field moderately convex, with four ribs defined by shallow pleural furrows, border moderately wide, rather flat, defined poorly by shallow and broad border furrows. Cranidial and pygidial surface with fine punctae, variable in density. Remarks. Based on the presence of a plectrum, the frontal area of the new species is interpreted as having a wide anterior border that is depressed posteriorly and is slightly convex anteriorly, having a short preglabellar field, and a paired subtriangular preocular areas. The new species resembles Lobocephalina pyriceps Opik from Queensland, Australia. Both species are similar in glabellar shape, the presence of a plectrum, and in the closely spaced palpebral lobes, which have their • 82.

anterior and posterior ends almost touching the axial furrows. The new species, however, differs in the structure of the frontal area. The frontal area has a wider (sag.) anterior border that is convex anteriorly and depressed posteriorly, rather than being simply convex, and it has a narrower prglabellar field. L. pyriceps is also differentiated by having a granulose surface prosopon. L. pyriceps and L. sinensis sp. nov. resemble L. emmrichi (Barrande) in essential characters such as glabellar shape, the large and closely spaced palpebral lobes, the course of the facial sutures, the shape of the posterior area of the fixigena, and the number of lateral glabellar furrows. However, as noted by Opik (1967, p. 249) for L. pyriceps, L. emmrichi (Barrande), the type species of Lobocephalina, is distinguished by a less tumid glabella posteriorly, more widely spaced palpebral lobes with anterior and posterior ends widely separated from the axial furrows, and a more distinct anterior border furrow. These distinctions also serve to differentiate the new species from the type species. It should be noted that the plectrum that is present in the Chinese and Australian species is absent in the type species. As a result of these differences, Opik ( 1967, p. 249 ) suggested that L. pyriceps might represent a new conokephalinoid genus, or even an early form of the Richardsonellidae. If so, the Chinese species would be congeneric with L. pyriceps. Until more is known of the Australo-Asiatic Lobocephalina, we follow Opik's (1967) practice in referring the present material to that genus.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs with trilobites indicative of the Liostracina bella Zone (equivalent to the lowermost part of the Glyptagnostus reticulatus Zone). Family

EULOMIDAE

Subfamily

Kobayashi, 1955

EULOMINAE

Genus

Kobayashi, 1955

STIGMATOA

Opik, 1963

Stigmatoa Opik, 1963, p. 87-89; Shergold, Cooper, Mackinnon, and Yochelson, 1976, p. 265; Henderson, 1976, p. 352, 353; Zhou, Liu, Meng, and Sun, 1977, p. 141; Romanenko, 1977, p. 179; Yang, 1978, p. 36; Shergold, 1982, p. 31; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 78, 79; Wittke, 1984, p. 143; Rickards, Bailie, and Jago, 1990, p. 209; Guo and Zhang, 1992, p. 243; Peng, 1992, p. 40; Lee and Choi, 1995, p. 38; Shergold, Feist, and Vizcaino, 2000, p. 661,662. Kerbinella Jegorova (Egorova), 1984, p. 26, 27. Type species. Stigmatoa diloma ()pik (1963, p. 89, 90, pl. 4, fig. 2), from the Erixanium sentum Zone, Georgina Basin, western Queensland, Australia; by original designation. Remarks. Kerbinella Jegorova, 1984 from the upper Cambrian of Siberia is regarded as a junior synonym of Stigmatoa Opik, 1963. The illustration of the holotype cranidium of Kerbinella conica Jegorova (Egorova, 1984, pl. 5, fig. 3), the type species of Kerbinella, seems indistinguishable in general respects from cranidia of Stigmatoa described from Australia, China, Siberia, and Iran. The genetic diagnosis and the description of the type species confirm the synonymy, as they agree with those of Opik (1963) for Stigmatoa and its type species S. diloma. o°

• 83"

Stigmatoa yangziensis Yang in Zhou et al., 1977 Plate 32, figures 1-7; Text-figure 10

Stigmatoa yangziensis Yang in Zhou et al., p. 141,142, pl. 12. Stigmatoa yangziensis Yang; Yang, p. 36, 37, pl. 5, figs. 8a, 8b. Stigmatoa yangziensis Yang; Liu, p. 305, pl. 216, fig. 7. Stigmatoa yangziensis Yang; Peng, p. 40, fig. 18: A-C. Stigmatoa yangziensis Yang; Peng, Babcock, and Lin, p. 105, pl. 15, fig. 12.

1977 1978 1982 1992 2001b

Holotype. By monotypy; cranidium (Zhou et al., 1977, pl. 44, fig. 12, CUGB 0242002; refigured by Yang, 1978, pl. 5, figs. 8a, 8b and by Liu, 1982, pl. 216, fig. 7; Text-fig. 10 herein) from the Chuangia-Prochuangia Zone, Huaqiao Formation, Tingziguan, Fenghuang, northwestern Hunan. New material. Ten cranidia (illustrated specimens NIGP 138211-138215) from collections PI323.5, PI336.5, and PI360.3.

A

Text-figure 10. Holotype of Stigmatoa yangziensis Yang in Zhou et al., 1977, partly exfoliated cranidium in anterolateral and dorsal views, CUGB 0242002. × 3 (original of Zhou et al., 1977, pl. 44, fig. 12; also Yang, 1978, pl. 5, fig. 8).

Remarks. The specimens from Paibi, northwestern Hunan, resemble in all essential respects the holotype of S. yangziensis from Fenghuang, northwestern Hunan (Text-fig. 10). Minor morphological variation in the sagittal width of the anterior border, the effacement of $3, and the degree of divergence of the anterior branches of the facial suture, is present in the new material. Occurrence. The holotype is from the Huaqiao Formation at Tingziguan, Fenghuang, northwestern Hunan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs with trilobites indicative of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone).

• 84"

Family KINGSTONIIDAE Kobayashi, 1933b Genus KINGSTONIA Walcott, 1924

Type species. Kingstonia apion Walcott, 1924 (p. 58, pl. 14, fig. 2, line drawing), from the Maryville Formation, Tennessee, USA; by original designation. Remarks. The concept of Kingstonia has been discussed by various authors (Walcott, 1924, p. 24; 1925, p. 103; Resser, 1936, p. 24, 25; Tasch, 1951, p. 295; 1952, p. 859; Shaw, 1952, p. 471,472; Chernysheva, 1953, p. 65; Palmer, 1954, p. 724, 725; 1962b, p. 29; Rozova, 1960, p. 67, 70; Kobayashi, 1933a, p. 277, 278; 1933b, p. 143; 1962, p. 68, 69; Palmer and Peel, 1981, p. 24, 25; Pratt, 1992, p. 67). Ucebia Walcott, 1924 is commonly regarded as synonymous with Kingstonia or a subgenus of Kingstonia. Pratt (1992) rediagnosed the genus without grouping it into two subgenera, K. (Kingstonia), and K. (Ucebia). Pratt's (1992) concept is followed here, but new material described as Kingstonia euryaxis sp. nov., which represents the oldest species of the genus, expands the genetic concept slightly to include species having an anteriorly expanded glabella, a slightly convex pygidial axis, and proparian facial sutures. Kingstonia was known previously only from Laurentia. According to Pratt (1992), all Laurentian species are from the upper Marjuman (traditional middle Cambrian). Four species assigned to Kingstonia from the upper Cambrian of Kazakstan and the Sayan-Altai Mountains were transferred to Parakoldinia Rozova by Shergold and Sdzuy (1984). Seven species were described from Northeast China and South Korea (Kobayashi, 1933a, 1933b, 1935, 1958; Endo, 1937), all of which are middle to latest late Cambrian (Furongian) in age. Pratt (1992, p. 67) regarded these species as not likely to be congeneric with Kingstonia. About the same time, Peng (1992, p. 104) transferred two of Endo's (1937) species, Kingstonia kuantungensis and K. perconvexa, to Parakoldinia and transferred questionably two of Kobayashi's (1933a) species, K. paichiaensis and K. parallela, to that genus. Peng (1992) regarded three other species as of uncertain assignment. Shergold and Sdzuy (1984, p. 99-101) noted that many homeomorphs exist among effaced, strongly convex trilobites that were described under a variety of genetic names referred to different families such as Kingstoniidae, Illaenuridae, and Plethopeldidae. Kingstonia is strongly similar to Koldinia and Parakoldinia (with its possible synonyms, Wanwanaspis Kobayashi, 1966 and Wanwanoglobus Kobayashi, 1966)in morphology, but the latter genera are obviously different in having palpebral lobes that lie well behind the midpoint of the glabella. Well preserved material of Parakoldinia from Turkey (Shergold and Sdzuy, 1984), and northwestem Hunan (Peng, 1992) show that Parakoldinia has a median glabellar node. The median node distinguishes Parakoldinia from Kingstonia, and suggests classification (with Symphysurina) within the family Nileiidae Angelin (Peng, 1992, p. 104). Koldinia and Parakoldinia have a much younger stratigraphic occurrence than Kingstonia.

Kingstonia euryaxis sp. nov. Plate 33, figures 1-14 2001b Kingstoniasp., Peng, Babcock, and Lin, p. l01, pl. l, figs. 3, 4.

Etymology. From Greek, eury, broad; axis, axial lobe, referring to the broad pygidial axis. Holotype. Cranidium (P1. 33, figs. 4-6, NIGP 138220) from collection P6. • 85"

Paratypes. Six cranidia and five pygidia (NIGP 138217-138219, 139121-138228) in collections P3.2, and P7.25; other fragment are in collections P7.5 and P 12.5. Diagnosis. Kingstonia with cranidium lacking anterior border; cranidium semicircular, with narrow (tr.) posterolateral projection beating clearly defined border furrow and border; axial furrows absent or poorly impressed, defining gently expanded glabella. Pygidium semicircular, weakly convex, with large, broadly conical axis; pleural furrows absent to poorly impressed. Description. Cranidium length three-fifths width, strongly convex (tr., sag.), with broadly rounded anterior margin, lacking anterior border. Glabella variably distinctive, faintly defined to defined only by posterior margin of occipital ring; glabella, if obvious, occupies full length of cranidium, subparallel-sided posteriorly, gently expanded in front of its midlength; occipital ring narrow (sag.), defined anteriorly by faint occipital furrow, twice as wide as posterolateral margin of cranidium. Palpebral lobe small, bar-like, curved downwardly, lying about the midlength of glabella. Fixigena narrow (tr.), subtriangular in outline, with clear posterior border that widens abaxially; lateral border wide (tr.), weakly defined. Anterior branch of facial suture subparallel to gently converging forward; posterior branch diverging diagonally rearward, diverging less after across crossing lateral border furrow to cut lateral border obliquely and meet lateral margin of cephalon anterior of genal angle. Pygidium semicircular, gently convex, length half width, with rounded anterolateral comer. Axis wide (tr.) and long, strongly to completely effaced; defined by straight, faint axial furrows; strongly tapered rearward, obtusely pointed posteriorly, continuing as short postaxial ridge to posterior margin. Pleural field half as wide (tr.) as axis on anterior margin, smooth or weakly to faintly segmented, with 4 pairs of interpleural furrows. Anterior border furrow weakly impressed; lateral and posterior border furrows absent or obscure. Remarks. According to Pratt (1992, p. 68), Kingstonia bealesoides, from the Cedaria minor Zone of the Rabbitkettle Formation in the Mackenzie Mountains, Canada, is the only species of Kingstonia that lacks an anterior cranidial border. In the absence of an anterior border, and in the general shape of the cranidial outline and the cranidial convexity, K. euryaxis sp. nov. is closely comparable with the Canadian species, but K. bealesoides differs in lacking a posterior border and a posterior border furrow. The pygidium of K. bealesoides is unknown, but the pygidium is known for most North American species. Where known, pygidia of Kingstonia have a relatively narrow, moderately convex, subcylindrical axes that are obtusely rounded posteriorly. Although the pleural segmentation of K. euryaxis is reminiscent of the exfoliated pygidia of some North American species (e.g., K. walcotti Resser, 1938a, pl. 12, fig. 4; K. rotundatai Resser, 1938a, pl. 12, fig. 10; K. clevelandensis Resser, 1938a, pl. 12, fig. 15), the broad conical axis differentiates the new species from all Laurentian species. K. euryaspis is probably the first valid record of a Kingstonia species in China. As noted above, formerly recorded species of Kingstonia from the upper Cambrian of China belong to Parakoldinia. Stratigraphically, the new species occurs in the basal P~chagnostus atavus Zone, and is much older than K. bealesoides, which Pratt (1992, p. 68) regarded as the oldest species of Kingstonia. In general features, the morphology of K. euryaxis is consistent with an assignment to Kingstonia. This species, the oldest known for the genus, possesses some features not known from the North American species. The features include an anteriorly expanded glabella and a weakly convex, broadly conical pygidial axis. The posterior branch of the facial suture in K. euryaxis meets the lateral margin of the cranidium in advance of the genal angle, and this suggests a proparian facial suture. In the North American species, the glabella, where visible on the exfoliated surface, is • 86"

subparallel-sided to gently tapered forward (Resser, 1942, pl. 7, fig. 35; pl. 8, fig. 44), the pygidial axis is commonly subcylidrical, and the facial suture is opisthoparian. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicatives of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone).

Family

LISANIIDAE

Chang, 1963

Genus LISANIAWalcott, 1911 Lisania Walcott, 1911, p. 82, 83" 1913, p. 163, 164; 1916, p. 403, 404; Resser, 1942, p. 28" Hup6, 1955, p. 181; Lu, 1957, p. 271; Lochman-Balk in Moore, 1959, p. O312; Kobayashi, 1960b, p. 368-370; Lu, Zhang, Zhu, Qian, and Xiang, 1963, p. 262; Zhou, Liu, Meng, and Sun, 1977, p. 171; Yin and Li, 1978, p. 491; Yang, 1978, p. 45; Nan, 1980, p. 496; Zhang, 1981, p. 170; Liu, 1982, p. 312; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 128; Lu and Qian, 1983, p. 60; Zhang and Wang, 1985, p. 408; Zhang and Jell, 1987, p. 134; Zhu, 1992, p. 345; Zhang in Zhang et al., 1995, p. 68; Guo, Zan, and Luo, 1996, p. 116. Aojia Resser and Endo in Kobayashi, 1935, p. 89; Resser and Endo, 1937, p. 172, 173; Resser, 1942, p. 6; Endo, 1944, p. 6; Hup6, 1955, p. 135; Lu, 1957, p. 271" Lochman-Balk in Moore, 1959, p. O311; Kraskov, Lazarenko, Ogienko, and Chernysheva, 1960, p. 235; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 266, 267; Zhou, Liu, Meng, and Sun, 1977, p. 172; Zhang and Wang, 1985, p. 410; Yin and Li, 1978, p. 490; Yang, 1978, p. 46; Liu, 1982, p. 313; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, 1991, p. 149; 1993, p. 192. ?Paraojia Sun and Zhou in Zhou et al., 1977, p. 173. Paraaojia Rozova in Lisogor et al., 1988, p. 73, 74. Metalishania Ju in Qiu et al., 1983, p. 129, 130. Type species. Anomocarella? bura Walcott (1905, p. 56; 1911, p. 82, pl. 15, fig. 2) from the Changhia Formation, Zhangxia, Shandong; by original designation. Other species. Arionellus agonius Walcott (1905, p. 58; 1913, p. 164, pl. 15, figs. 17, 17a) from the Amphoton Zone, Changhia Formation, Yangzhuang, Xintai, Shandong; Lisania rectangularis Sun (1924, p. 55, pl. 4, figs. 2a, b) from the Changhia Formation, Zhaogezhuang, northeastern Hebei; Aojia spinosa Resser and Endo in Kobayashi (1935, p. 89, pl. 24, figs. 3, 4) from the Changhia Formation, near Yantai (now Dengta), central Liaoning; A. fracta Resser and Endo (1937, p. 175, pl. 47, fig. 5) from the Changhia Formation, near Liaoyang, central Liaoning; A. quadrata Resser and Endo (1937, p. 177, pl. 34, fig. 5) from the Changhia Formation, Changxingdao Island, southern Liaoning; Proasaphiscus quadrilateralis Resser and Endo, 1937 (p. 263, 264 in part, pl. 48, figs. 23-25 non 26-28) from the Changhia Formation, near Liaoyang, central Liaoning; Proasaphiscus humilis Resser and Endo (1937, p. 264, pl. 49, fig. 2) from the Changhia Formation, Changxingdao Island, southern Liaoning; Lisania paratungjenensis Yang in Zhou et al. (1977, p. 171, pl. 51, fig. 2) [=Lisania placida Yang in Yin and Li, 1978, p. 491, pl. 165, figs. 5, 6], from the Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan and from Fengmuping, Tongren, eastern Guizhou; Aojia hespera Yang in Zhou et al. (1977, p. 172, pl. 51, figs. 4-6) from the middle part of the Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan [=Aojia puteata Qiu in Qiu et al., 1983, p. 129, pl. 41, figs. 10-12]; Lisania subcylindrica Zhang, 1981 (p. 170, pl. 64, fig. 1); Aojia yuanjiangensis Yang in Zhou et al. (1977, p. 172, pl. 51, figs. 8, 9) from the Huaqiao Formation, • 87.

Huaqiao, Baojing and Niutouping, Fenghuang, northwestern Hunan [=Metalisania acutifrons Ju in Qiu et al., 1983, p. 130, pl. 41, fig. 13; ? =Metalisania acutifrons Ju in Qiu et al., 1983, p. 130, pl. 41, fig. 14; =Paraaojia compacta Rozova in Lisogor et al., 1988, p. 74, 75, pl. 4, fig. 7]; Lisania subcylindrica Zhang (1981, p. 170, pl. 64, fig. 1) from the Mohershan Formation, Kuruktag, eastern Tienshan, Xinjiang; Metalisania subquadrata Ju in Qiu et al. (1983, p. 130, pl. 42, figs. 1, 2) from the upper part of the Yangliugang Formation, Fuyang, Zhejiang; Metalisania zhujiensis Ju in Qiu et al., 1983 (p. 131, pl. 42, figs. 3-5) from the upper part of the Yangliugang Formation, Zhuji, western Zhejiang; Lisania handanensis Zhang and Wang (1985, p. 408, pl. 123, fig. 4) from the Changhia Formation, near Wuan, southern Hebei; Lisania lubrica Zhang and Wang (1985, p. 409, pl. 123, fig. 5) from the Changhia Formation, near Wutai, northern Shanxi; Lisania lingqiuensis Zhang and Wang (1985, p. 410, pl. 123, fig. 3) from the Changhia Formation, near Lingqiu, northern Shanxi [=Aojia elongata Zhang and Wang, 1985, p. 411, pl. 123, fig. 12]; Aojia wanyuanensis Yang and Liu (in Yang et al., 1991/1993, p. 149/192, pl. 17, figs. 6, 7), from the Baguamiao Formation, near Wanyuan, southeastern Sichuan. In following Zhang and Jell (1987), Aojia crassa Resser and Endo, 1937 is tentatively assigned to Lisania. Remarks. We follow Kobayashi (1960b, p. 368) and Zhang and Jell (1987, p. 134) in regarding Aojia as a junior synonym of Lisania. We further consider Metalisania Ju in Qiu et al., 1983 and Paraaojia Rozova in Lisogor et al., 1988 to be additional junior synonyms of Lisania. Paraojia Sun and Zhou in Zhou et al., 1977 is possibly a junior synonym of Lisania. An emended diagnosis for Lisania was given by Zhang and Jell (1987), and that genetic concept is followed here. Kobayashi (1960b, p. 369) listed 27 species from the North China Platform that were placed either in Aojia or Lisania. In addition, he added one additional species to the genus, Lisania convergens, and transferred Proasaphiscus quadrilateralis to Lisania. Most of the species listed by Kobayashi (1960), however, were subsequently placed in synonymy or excluded from Lisania by Lu et al. (1965) and Zhang and Jell (1987). These species include: Arionellus alala Walcott, 1905, Aojia yentaiensis Resser and Endo, 1937, A. luna Resser and Endo, 1937, A. reflexa Endo, 1944, and Lisania convergens Kobayashi, 1960b, which were regarded as junior synonyms of Lisania agonius (Walcott, 1905; Zhang and Jell, 1987); Aojia punctata Resser and Endo, 1937, which was regarded as a junior synonym of Lisania quadrata (Resser and Endo, 1937; see Zhang and Jell, 1987); Arionellus ajax Walcott, 1905, which was excluded and used as the type species of their new genus Ajacicrepida by Zhang and Jell (1987); Ptychoparia tellus Walcott, 1905, A. longispina Resser and Endo, 1937, and A. peiciliensis Resser and Endo 1937 were transferred to the genus Pla~lisania, with A. peiciliensis being suppressed as a junior synonym of A. longispina (Zhang and Jell, 1987); Lisania? hsuchiachuangensis Sun, 1924 is transferred to Redlichaspis (=Lisanella; Lu et al., 1965, p. 274); Anomocarella biston Walcott, 1905 and Ptychoparia undata Walcott, 1906, which were transferred to Redlichaspis Kobayashi, 1935 (Zhang and Jell, 1987); Menocephalus belenus Walcott, 1905, which was tentatively transferred to Damesops Chu 1959 by Zhang and Jell (1987). Damesops is probably a junior synonym of Parabalckwelderia Kobayashi, 1942; Aojia reflexa Endo, 1944, which was considered to be an invalid species (Kobayashi, 1960b, p. 370); Aojia tumida Resser and Endo, 1937 is regarded as an immature form belonging to Metanomocarella, whereas Aojia divergens Endo, 1944 is also an immature trilobite according to Lu et al. (1965, p. 269, 337); A. ? carina Resser and Endo, 1937, A. angustata Endo, 1944, and A. depressa Endo, 1944 were • 88"

not considered to belong to Aojia by Lu et al. (1965), although they included questionably these species in Aojia; Lisania sp. indet, b (Walcott, 1913) was considered to be a representative of Chuangia (Zhang and Jell, 1987, p. 203,204); Ju (in Qiu et al., 1983) transferred Aojia aigawaensis Endo, 1937 to his new genus Metalisania. However, we consider Metalisania to be a junior synonym of Lisania, and A. aigawaensis may be better classified within Platylisania Zhang and Jell, 1987. Based on newly discovered pygidia, Proasaphiscus quadrilateralis can be only questionably referred to Lisania. After 1960 some 21 additional species from South China and North China have been assigned to either Lisania or Aojia. These species are: Lisania tungjenensis Nan in Egorova et al., 1963 Lisania paralala Yang in Zhou et al., 1977 Lisania paratungjenensis Yang in Zhou et al., 1977 Lisania placida Yang in Yin and Li, 1978 Lisania guizhouensis Lee and Yin in Yin and Li, 1978 Lisania punctata Lu and Chien in Yin and Li, 1978 Lisania subcylindrica Zhang, 1981 Lisania handanensis Zhang and Wang, 1985 Lisania taihangshanensis Zhang and Wang, 1985 Lisania gengzhenensis Zhang and Wang, 1985 Lisania lubrica Zhang and Wang, 1985 Lisania wutaiensis Zhang and Wang, 1985 Lisania wuanensis Zhang and Wang, 1985 Lisania lingqiuensis Zhang and Wang, 1985 Lisania sp. Zhang in Zhang et al., 1995 Aojia hespera Yang in Zhou et al., 1977 Aojia yuanjiangensis Yang in Zhou et al., 1977 Aojia puteata Qiu in Qiu et al., 1983 Aojia robusta Zhang and Wang, 1985 Aojia elongata Zhang and Wang, 1985 Aojia yantouensis Zhang and Wang, 1985 Aojia wanyuanensis Yang and Liu in Yang et al., 1991 (reprinted 1993) Among species in this list, 13 species are based only on a single cranidium, and others are based on a few specimens. In most cases, new species were unsatisfactorally illustrated, and their validity needs to be reevaluated. One of us (SP) examined the material from the northwestern Hunan and eastern Guizhou region (i.e., that of Egorova et al., 1963 and Yang, 1978). The examination reveals that Lisania tungjenensis Nan (Egorova et al., p. 34, pl. 6, figs. 8-12) and L. paralala Yang (in Zhou et al., 1977, p. 171, pl. 51, fig. 1) are best placed within Baojingia. Lisania paratungjenensis Yang (in Zhou et al., 1977, p. 171, pl. 51, fig. 2) and L. placida Yang (in Yin and Li, 1778, p. 491, pl. 165, figs. 5, 6), both from the Huaqiao Formation in the Guizhou - Hunan border area, are synonymous. These two species are identical in nearly all respects, differing only in the absence of a short occipital spine in L. placida. New material shows that such a distinction in Lisania is of no more than an intraspecific variation. For the same reason, Aojia elongata Zhang and Wang (1985, p. 411, pl. 123, fig. 12) is regarded as a junior synonym of Lisania lingqiuensis Zhang and Wang (1985, p. 410, pl. 123, fig. 3). According to Zhang and Wang (1985), both species occur in the same formation and at same locality in northwestern Shanxi. Lisania guizhouensis Lee and Yin (in Yin and Li, 1978, p. 491, pl. 165, fig. 1) from northern Guizhou and L. wuanensis Zhang and Wang (1985, p. 410, pl. 123, fig. 2) from southern Hebei • 89•

belong probably to Qiandongaspis Yuan and Yin, 1998. Both species differ from Lisania in having relatively longer, more posteriorly placed palpebral lobes, well-defined palpebral furrows, and narrower (tr.), strongly inward inclined palpebral areas. These features seem to be most similar to those of Qiandongaspis. The presence of a short preglabellar field in Lisania punctata Lu and Chien (in Yin and Li, 1978, p. 492, pl. 165, fig. 8) from eastern Guizhou, and in L. taihangshanensis Zhang and Wang (1985, p. 408, pl. 123, fig. 1) from northern Shanxi suggests that these two species may belong to Redlichaspis Kobayashi, 1935. Aojia puteata Qiu (in Qiu et al., 1983, p. 129, pl. 41, figs. 10-12) from northern Jiangsu resembles A. hespera Yang (in Zhou et al., 1977, p. 172, pl. 51, figs. 4-6) from northwestern Hunan in both cranidial and pygidial respects, and it may be a junior synonym of the latter. Its more effaced palpebral furrows are apparently due to the testaceous state of the holotype cranidium. Aojia robusta Zhang and Wang (1985, p. 410, 411, pl. 123, fig. 13) has a short glabella that is tapered forward moderately and obtusely rounded anteriorly; a strongly effaced occipital furrow, and an occipital ring with a stout median spine. This species is almost certainly a junior synonym of Paralisanella tenuilabrosa Qiu (in Qiu et al., 1983, p. 132, 133, pl. 43, figs. 2, 3). Similarly, A. yantouensis Zhang and Wang (1985, p. 411, pl. 123, figs. 10, 11 ) is probably also a representative of Paralisanella Ju (in Qiu et al., 1983). It closely resembles Paralisanella longispinosa Qiu (in Qiu et al., 1983, p. 132, 133, pl. 43, figs. 2, 3). Lisania gengzhenensis Zhang and Wang (1985, p. 408, 409, pl. 123, fig. 5) and L. wutaiensis Zhang and Wang (1985, p. 409, pl. 123, fig. 9), both from the Changhia Formation at the same locality in northwestern Shanxi, appear to be indistinguishable from each other in morphology. They are characterized by a shorter, notably tapered glabella with a truncate front and a fairly broad anterior border furrow. These features are nothing like those found in Lisania, but suggest that these species may constitute an undescribed genus that is related to Paralisania Qiu in Qiu et al., 1983. Two cranidia from northwestern Hunan were assigned to Aojia? peichiliensis Resser and Endo by Zhou et al. (1977, pl. 51, figs. 8, 9). They differ apparently from Platylisania longispina [=Aojia? peichiliensis] of the North China Platform (see Zhang and Jell, 1987, p. 140) in having a relatively elongate glabella, and are probably referable to Lisania hespera (Yang). Lisania paratungjenensis Yang in Zhou et al., 1977

Plate 37, figures 5-10; Text-figure 11 1977 1978 1978 1978 1982

Lisania paratungjenensis Yang in Zhou et al., p. 171, pl. 51, fig. 2. Lisania placida Yang in Yin and Li, p. 491, pl. 165, figs. 5, 6. Lisania paratungjenensis Yang, p. 46, pl. 6, fig. 8. Lisania placida Yang, p. 46, pl. 6, figs. 14, 15. Lisania paratungjenensis Yang; Liu, p. 312, pl. 219, fig. 17.

Holotype. By monotypy; an exfoliated cranidium (Yang in Zhou et al., 1977, pl. 51, fig. 2, CUGB 0112201; refigured by Yang, 1978, pl. 6, figs. 6a, 6b and by Liu, 1982, pl. 219, fig. 17; Text-fig. 11 A, B herein) from the Lisania paratungjenensis Zone, Huaqiao Formation, at Huaqiao, Baojing, northwestem Hunan. New material. Two cranidia, plus an incomplete cephalon, and one pygidium that are tentatively assigned to this species, NIGP 138276-138279, in collections P125, P200.7, P204, P207, and W138.9. • 90

•

Remarks. Lisania paratungjenensis and L. placida are considered to be synonymous. L. placida was differentiated from L. paratungjenensis only in the absence of an occipital spine (Yang in Yang and Lin, 1978). Large collections of Lisania huayuanensis sp. nov. show that an occipital spine is variably expressed and simply represents one form of intraspecific variation. Both L. paratungjenensis and L. placida co-occur in the Huaqiao Formation at Huaqiao, Baojing, northwestern Hunan. According to Yang (1978), figured specimens of both species are from collection Hbh 12, whereas the type specimens of L. placida are in both collection Hbh 12 and in collection Hbh 13.

Text-figure 11. Lisania paratungjenensis Yang in Zhou et al., 1977, all from the Huaqiao Formation at Huaqiao, Baojing, northwestern Hunan. A, B, holotype, exfoliated cranidium in dorsal and anterolateral views, CUGB 0112201, × 6 (original of Zhou et al., 1977, pl. 51, fig. 2; also Yang, 1978, pl. 6, fig. 8); C-F, exfoliated cranidium in dorsal and anterolateral views and pygidium in dorsal and posterolateral views, CUGB 0113201, 0113202, × 5.2, × 5.2, × 8, × 8 (original of Yin and Li, 1978, pl. 165, figs. 5, 6; also Yang, 1978, pl. 6, figs. 14, 15). The cranidium and the pygidium are the syntypes of Lisania

placida Yang in Yin and Li, 1978, and were subsequently designed as the holotype and paratype of the species respectively by Yang 1978 (p. 45, 79). "91-

Cranidia from the Huaqiao Formation at Paibi, Huayuan, Hunan, conform in all observed respects to the holotypes of both L. paratungjenensis and its junior synonym, L. placida, and are thus referred to L. paratungjenensis. A pygidium lacks anterolateral spines and conforms to Yang's (1978, p. 47) description of the pygidium of L. placida. Morphologically it agrees with the pygidium that Yang (1978, pl. 6, fig. 15) assigned to L. placida in all respects except for the rather closely spaced fulcra, and these seem to represent a form of intraspecific variation. However, both of the anterolateral comers of Yang's (1978) pygidium of L. placida, which is the only figured pygidium for the species, are damaged and the status of the anterolateral spines remains uncertain for L. paratungjenensis (see Text-fig. 11 E, F). More material is needed to clarify if the pygidium illustrated here is conspecific with the one assigned by Yang (1978) to the species. Occurrence. The holotype is from the Huaqiao Formation at Huaqiao, Baojing, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicatives of the Pianaspis sinensis Zone (equivalent to the lower part of the Goniagnostus nathorsti Zone to the Lejopyge laevigata Zone). Lisania yuanjiangensis (Yang in Zhou et al., 1977)

Plate 35, figures 12-16; Plate 36, figures 1-18; Plate 37, figures 1-3" Text-figure 12 A-C, ?F, G 1977 ?1977 ?1978 1978 ?1978 1982 1983 ?1983 1988 2001b

Aojia yuanjiangensis Yang in Zhou et al., p. 172, pl. 51, figs. 6, 7. Aojia hespera Yang in Zhou et al. (in part), p. 172, pl. 51, fig. 5 only. Aojia hespera Yang (in part); Yin and Li, p. 490, 491, pl. 165, fig. 4 only. Aojia yuanjiangensis Yang, p. 47, pl. 6, figs. 10, 11. Aojia hespera Yang (in part), p. 47, pl. 6, fig. 13 only. Aojia yuanjiangensis Yang; Liu, p. 313, pl. 219, figs. 9, 22. Metalisania acutifrons Ju in Qiu et al., p. 130, pl. 41, fig. 13. Metalisania acutifrons Ju in Qiu et al., p. 130, pl. 41, fig. 14. Paraaojia compacta Rozova in Lisogor et al., p. 74, 75, pl. 4, fig. 7. Lisania yuanjiangensis (Yang); Peng, Babcock, and Lin, p. 102, pl. 4, figs. 7, 8; non fig. 3 [=Baojingia tungjenensis (Nan in Egorova et al., 1963)].

Holotype. Exfoliated cranidium (Yang in Zhou et al., 1977, pl. 51, fig. 6, CUGB 0113102; refigured by Yang, 1978, pl. 6, figs. 10a, b and by Liu, 1982, pl. 219, fig. 9; Text-fig. 12 A, B herein), from the Paramphoton [=Amphoton] Zone, Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan. New material. More than 100 sclerites including cranidia, librigena, and pygidia (illustrated specimens NIGP 138254-138274), in collections P121.48, P126.9, P130.5, P131.7, P134.2, P135.3, P135.7, P136.3, P136.7, P138.8, P145.5, P156, P160, W98, W105, W111.3, W121.6, and W128.9. Remarks. New material from Paibi and Wangcun, northwestern Hunan, resembles closely the type material, which comes from the same region, in both cranidial and pygidial respects. Key characters that warrant placing the new material in this species include a yoke-shaped anterior border; an elongate subrectangular glabella with parallel or slightly constricted sides and a rounded front; relatively wide palpebral areas; relatively short, notably slanted eye ridges; converging anterior branches of facial suture; and a transverse pygidium with a stout axis beating four segments and a • 92.

pair of short anterolateral spines. The paratype pygidium of Lisania hespera (Yang in Zhou et al., 1977, pl. 51, fig. 5) is reassigned here to L. yuanjiangensis, as it seems indistinguishable from the new pygidia from Hunan, which are associated with the cranidia of L. yuanjiangensis. According to Yang (1978), L. yuanjiangensis and L. hespera co-occur in collection HBh 13 from the Huaqiao section in Baojing, northwestern Hunan, and this supports the decision to reassign the paratype.

Text-figure 12. A-C, ?F, G Lisania yuanjiangensis (Yang in Zhou et al., 1977). A, B, partly exfoliated cranidium in dorsal and anterolateral views and C, exfoliated pygidium, CUGB 0113102 and 0113103, × 4.5, × 4.5, × 6, designated subsequently as the holotype and paratype of the species respectively by Yang, 1978 (p. 47, 79; original of Zhou et al., 1977, pl. 51, figs. 6, 7: also Yang, 1978, pl. 6, figs. 10, 11); ?F, G, testaceous pygidium, CUGB 0113502, × 6, originally referred to Lisania [=Aojia] hespera Yang in Zhou et al., 1977 (pl. 51, fig. 5; also Yang, 1978, pl. 6, fig. 13); D, E, Lisania hespera (Yang in Zhou et al., 1977). Testaceous cranidium in dorsal and anterolateral views, CUGB 011350, both x 5, designated subsequently as the holotype of this species by Yang (1978) (original of Zhou et al., 1977, pl. 51, fig. 4: also Yang, 1978, pl. 6, fig. 12). New material adds some previously unknown information about L. yuanjiangensis.

The •

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posterior branches of the facial suture diverges strongly rearward from the posterior end of the palpebral lobe until meeting the posterior border furrow and then defecting rearward and slightly outward to cross the posterior border furrow and border and cut the posterior margin at an angle of about 80 ° to the sagittal line. The librigena has a narrow and convex genal field, a moderately high eye socle, and a stout genal spine with a broad base. The new material also shows some morphological variation in the species. The most evident variation involves the curvature in the anterior border and border furrow, in the width of the palpebral area, in the proportional length of the glabella, and in the nature of the glabellar sides.

Occurrence. The holotype is from the Huaqiao Formation in northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicatives of the lower part of the Pianaspis sinensis Zone (equivalent to the upper part of the Pt3,chagnostus punctuosus Zone to the Lejopyge laevigata Zone). Lisania sp. cf. L. agonius (Walcott, 1905) Plate 37, figures 11-13

Arionellus agonius Walcott, p. 58. Arionellus alala Walcott, p. 59. Lisania agonius (Walcott); Walcott, p. 164, pl. 15, figs. 17, 17a. Lisania alala (Walcott); Walcott, p. 165, 166, pl. 15, figs. 19, 19a-d. Aojia yentaiensis Resser and Endo, p. 175, 176, pl. 46, figs. 24-26. Aojia luna Resser and Endo (in part), p. 176, pl. 46, figs. 33, 35 non fig. 34. Aojia luna Endo, p. 64, 65, pl. 2, figs. 6-9. Lisania agonius (Walcott); Kobayashi, p. 369. Lisania alata (Walcott); Kobayashi, p. 369. Lisania convergens Kobayashi, p. 369. Lisaniaagonius (Walcott); Lu, Zhang Zhu, Qian, and Xiang, p. 263, pl. 45, figs. 3, 4. Lisaniaalata (Walcott); Lu, Zhang Zhu, Qian, and Xiang, p. 264, pl. 45, figs. 9-11. Aojia (?) luna Resser and Endo; Lu, Zhang Zhu, Qian, and Xiang, p. 269, 270, pl. 46, figs. 7, 8. cf. 1965 Aojia (?) yantaiensis Resser and Endo; Lu, Zhang Zhu, Qian, and Xiang, p. 271, pl. 47, figs.l, 2. cf. 1987 Lisaniaagonius (Walcott); Zhang and Jell, p. 134, 135, pl. 51, figs. 13-16; pl. 52, figs. 1-5; pl. 53, fig. 8; pl. 55, fig. 2. cf. 1996 Lisaniaagonius (Walcott); Guo, Zan, and Luo, p. 116, 117, pl. 56, figs. 1a-c, 2, 3, 5, 6, 8.

cf. 1905 cf. 1905 cf. 1913 cf. 1913 cf. 1937 cf. 1937 cf. 1944 cf. 1960b cf. 1960b cf. 1960b cf. 1965 cf. 1965 cf. 1965

Lectotype ofLisania agonius. Cranidium, USNM 58048 (Walcott, 1913, pl. 15, fig. 17; refigured by Zhang and Jell, 1987, pl. 51, fig. 13), from the Amphoton Zone, Changhia Formation,Yangzhuang, Shandong; designated by Zhang and Jell, 1987 (p. 135). Material. Four cranidia and one pygidium (illustrated specimens NIGP 138280-138282) in collections P 108, P 114, and P 115.6. Remarks. Specimens from the Huaqiao Formation, northwestern Hunan, that are compared to Lisania agonius are characterized by a glabella that is tapered forward gently; by a broadly rounded •

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anteriorly and beating four pairs of faintly impressed lateral furrows and a weak carina; by a pair of fossulae on the axial furrows; by a narrow palpebral area that is about one-fourth as wide as the glabella; and by a gently convex, moderately wide (sag.) anterior border. The anterior branches of the facial suture in this species extend forward in subparallel fashion, becoming directed inwardly and forwardly after crossing the anterior border furrow and meeting the anterior margin at the points opposite the fossulae. The posterior branches diverge rearward diagonally until reaching the level of the posterior one-third of the posterior field of the fixigena, then turn rearward and slightly outward to meet the posterior margin with the o3 point beyond the outer limit of the palpebral lobe. The associated pygidium is characterized by an axis that bears only two tings and an inverted trapezoidal terminal piece, and by short paired anterolateral spines. This species resembles closely Lisania agonius from the North China Platform, especially the lectotype of L. agonius (Zhang and Jell, 1987, pl. 51, fig. 13). The shape of the glabella, the width and shape of the fixigena, the presence of a carina, the proportional length and convexity of the anterior cranidial border, and the shape of occipital ring are similar to those features in L. agonius. However, some features in L. agonius are different from those in the new material from Hunan. These characters include the presence of fossulae, the nature of posterior branch, which is deflected at its midway point in the present species but, as shown by the librigena and the cranidium (Zhang and Jell, 1987, pl. 51, figs. 15, 16), is straight and directed diagonally in L. agonius. The posterolateral projection in L. agonius is triangular rather than subrectangular. Only one pygidium is known for L. agonius (Zhang and Jell, 1987, pl. 52, fig. 16), and it differs in having a conical axis and in lacking posterolateral spines. It is uncertain if the new specimens represent a new species. Because the species is represented by a small collection that is unsatisfactorily preserved, open nomenclature is used.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicatives of the upper part of Dorypyge richthofeni Zone to the lower part of Pianaspis sinensis Zone (equivalent to the Pt3,chagnostus punctuosus Zone). Lisania paibiensis sp. nov. Plate 34, figures 1-16; Plate 35, figures 1-11 2001b Lisaniatungjenensis Nan; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 14.

Etymology. After Paibi Villlage. Holotype. Cranidium (P1.35, fig. 1, NIGP 138243) from collection P180.3. Other material. More than 80 sclerites including cranidia, hypostomes, librigena, and pygidia, illustrated paratypes (NIGP 138229-138242, 138244-138253) in collections P164.2, P171, P171.5, P174, P178.3, P180.3, W132.5, W139.2, and W146.2. Diagnosis. Lisania with glabella truncated anteriorly. Eye ridge moderately long and slanting gently rearward. Anterior border crescentic; anterior border furrow evenly and gently curved. Posterolateral projection narrow, with distal end opposite the outer margin of palpebral lobe. Anterior branch of facial suture subparallel to slight diverging forward; posterior branch deflected midway, with the posterior section directing inward then outward. •

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Description. Cranidium subquadrate in outline, length subequal to width, with evenly curved anterior margin. Anterior border crescentic, gently convex, occupying about 0.15 of sagittal length of cranidium; anterior border furrow wide and moderately defined, evenly and gently curved forward. Glabella rectangular, length about twice width, truncate anteriorly, with sides slightly concave or straight. Lateral glabellar furrows consist of four faint pairs; S 1 bifurcate with posterior branch sinuous, directed diagonally with outer end lying slightly posterior to midpoint of axial furrow; $2 curved gently, moderately long, extending inward from axial furrow at the level of anterior one-third of glabellar length, curving gently inward and rearward; $3 short, curved gently, isolated from axial furrow; $4 shortest, directed inward and forward, isolated from axial furrow, opposite eye ridge. Occipital furrow weak, bowed gently rearward; occipital ring crescentic, longest sagittally, narrowing rapidly abaxially, with median node lying close to occipital furrow, and with or without medial occipital spine. Paired lateral bulges of glabella prominent, located immediately behind S1. Palpebral lobe moderately large, located subcentrally, somewhat slanted, defined by shallow palpebral furrow. Palpebral area of fixigena half as wide as glabella, gently convex; preocular field wider than long, sloping downward and outward; posterior area slightly wider than half of glabella, with posterolateral projection narrow, curving strongly downward with distal end slightly beyond or within outer limit of palpebral lobe. Anterior branch of facial suture diverging slightly forward or parallel forward to anterior border furrow, then turning abruptly inward, crossing anterior border obliquely with ct point opposite axial furrow; posterior branch bracket-like, with short anterior section directed diagonally and long posterior section directed inwardly and rearwardly with c0 point lying well within the outer limit of palpebral lobe. Librigena with genal field narrower than lateral border, slightly convex, widening rearward, defined by shallow border furrow. Genal angle with moderately long, flat spine being broad at base. Hypostome subrectangular, with ovate middle body and deeply incised middle furrow that is not connected medially. Anterior margin gently curved forward; anterior wing triangular. Lateral border ridge-like, defined by deep and broad lateral border furrow that continues rearward as posterior border furrow. Posterior border narrow sagittally; posterior margin transverse. Pygidium transverse-subelliptical. Axis thick, gently tapered rearward to posterior border furrow, slightly narrower than pleural region, with four well defined tings, a poorly defined fifth ring, and short crescentic terminal piece. Pleural field rather convex, with two to three fibs, defined by deep and broad pleural furrows. Interpleural furrows faint on first one or two segments. Border furrows distinct; borders flat, rather wide, of uniform width; posterolateral spine tiny, directed posteriorly. Remarks. The new species is similar to Lisania paratungjenensis, but differs principally in having a proportionally narrower and longer glabella that is truncate rather than obtusely rounded anteriorly, a more convex anterior border, a deeper and broader anterior border furrow, a wider palpebral area, a narrower posterolateral projection, a deflected posterior branch of the facial suture, and less convexity. In Lisania paratungjenensis, the occipital node is centrally rather than anteriorly placed. According to Yang (1978), the pygidium of L. paratungjenensis [=L. placida] lacks anterolateral spines and its axis bears four segments. If true, then L. paibiensis sp. nov. also can be distinguished by the presence of posterolateral spines and by a more segmented axis. Lisania paibiensis is also similar to L. yuanjiangensis, but the latter is readily differentiated by having a glabella that is broadly rounded anteriorly, having a yoke-shaped rather than crescentshaped anterior border, having a sinuous anterior border furrow, having a shorter and more slanting eye ridge, having a wider but shorter posterolateral projection, and by having a relatively wide genal field on the librigena. In L. yuanjiangensis, the anterior branch of the facial suture is more convergent forward and the posterior branch has a shorter posterior section. The pygidium of the new species is almost indistinguishable from that of L. yuanjiangensis except for the axis, which •

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has one fewer segment. The presence of an occipital spine was commonly used previously as a character of species-level significance in Lisania, but a large collection of the new species shows that the occipital spine is variably developed. As such, the presence or absence of an occipital spine is here regarded as an example of intraspecific variation, and it is possibly a dimorphic feature.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicatives of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Lisania wangcunensis sp. nov. Plate 37, figure 14

Holotype. Cranidium (P1.37, fig. 14, NIGP 138283) from collection W 139.2. Diagnosis. Lisania with narrow, bar-like anterior cranidial border, defined by shallow and shallow border furrow; glabella rectangular, largely effaced with broadly rounded front; palpebral area width more than half that of glabella. Anterior branch of facial suture diverging slightly forward. Posterior branch enclosing rectangular posterior area of fixigena. Description. Cranidium subquadrate, length slightly shorter than width between palpebral lobes. Anterior border bar-like, moderately convex, of nearly uniform width (sag. exs.), curved forward slightly. Preglabellar field absent. Glabella subrectangular, with parallel sides and obtusely rounded front, defined by shallow axial and preglabellar furrows; lateral glabellar furrows effaced. Occipital furrow shallow; occipital ring crescentic, probably beating median spine. Eye ridge faint. Palpebral lobe crescentic, located at midlength of cranidium; palpebral furrow shallow. Palpebral area about glabellar width. Anterior branch of facial suture diverging forward slightly; posterior branch of facial suture enclosing rectangular posterior area of fixigena. Surface smooth. Remarks. This single cranidium is considered to represent a new species of Lisania. It possesses a narrow, bar-like anterior border that differentiates it from all other species assigned previously to either Lisania or Aojia except for L. bura (Walcott), the type species of Lisania. The anterior border, however, is more curved in the type species of Lisania. Moreover, the type species is also differentiated by having a narrower fixigena, and a broader anterior border furrow. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicatives of the Pianaspis sinensis Zone (equivalent to the lower part of the Lejopyge laevigata Zone). Lisania? sp. Plate 37, figure 4

Material. One pygidium (NIGP 1388275) in collection W 146.2. Remarks. This pygidium is tentatively assigned to Lisania because it has features suggesting either • 97"

Lisania or Eoshengia. It is characterized by having a slender axis that bears 6 tings, with the last one being poorly defined; a small, crescentic terminal piece; and a pair of tiny anterolateral spines. The pleural ribs are of uniform width, and defined by deep, trough-like pleural furrows. Such pleural furrows were not known previously in Lisania, and are more similar to the furrows of Eoshenjia. The illustrated pygidium differs from reported species of Eoshenjia in having anterolateral spines. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicatives of the upper part of the Pianaspis sinensis Zone (equivalent to the lower part of the Lejopyge laevigata Zone). Genus BAOJINGIA Yang in Zhou et al., 1977

Baojingia Yang in Zhou et al., 1977, p. 173, 174; Yang, 1978, p. 48; Lu and Lin, 1989, p. 136. Eoshengia Yang in Zhou et al., 1977, p. 174; Yang, 1978, p. 49; Yin and Li, 1978, p. 492, 493; 1981, p. 171. Xichuania Yan in Yang et al., 1991, p. 154; 1993, p. 199. Quandraspis Yang in Yang et al., 1991, p. 156; 1993, p. 201,202. Type species. Baojingia youshuiensis Yang in Zhou et al., 1977 (p. 174, pl. 51, figs. 10, 11), from the Lisania tungjenensis Zone of the Huaqiao Formation at Huaqiao, Baojing, northwestern Hunan; by original designation. Remarks. Eoshengia (type species, with E. subquadrata Yang, 1978) was considered to be possibly synonymous with Shengia by Shergold et al. (2000, p. 613) but here it is regarded as a junior synonym of Baojingia. Both Baojingia and Eoshengia were erected by Yang (in Zhou et al., 1977), but Baojingia appears earlier in terms of page position in the same volume. Baojingia is monospecific, whereas Eoshengia was erected to embrace three species, and a type species was not designated. This situation lasted until Yang (1978) selected E. subquadrata as the type species. Morphologically Baojingia and Eoshengia are extremely similar to each other, the only difference being that Baojingia has deeply impressed lateral glabellar furrows. The furrows are weak to nearly completely effaced in Eoshengia. Such a difference in the effacement of lateral glabellar furrows is considered to be of only species-level significance. Baojingia is apparently related to Shengia Hsiang (in Egorova et al., 1963). Superficially, Baojingia and Eoshengia resemble Shengia, but close comparision reveals that both genera are different in the structure of the anterior cranidial border and the anterior cranidial border furrow. Baojingia has a W-shaped anterior border furrow; an anterior border that is relatively wide (sag.) and flat; narrowing unevenly abaxially; and an anterior cranidial margin that is sinuous or yoked-shaped. In Shengia, both the anterior border furrow and the anterior cranidial margin are evenly curved, and the anterior cranidial border is narrower and quite convex. Xichuania Yan (in Yang et al., 1991; reprinted 1993) is apparently a junior synonym of Baojingia. The type species of Xichuania, X. xiuzigouensis Yan, is almost identical to Baojingia youshuiensis Yang, the type species of Baojingia. Quandraspis Yang (in Yang et al., 1991; reprinted 1993) is also regarded as a junior synonym of Baojingia (Text-fig. 13). All of the illustrated specimens of Q. zhengwanensis Yang (in Yang et al., 1991, pl. 16, fig. 14; pl. 18, figs. 9-11; reprinted 1993), the type species of Quandraspis Yang, are strongly compressed longitudinally, resulting in a transverse appearance for those cranidia, and a subquadrate outline for • 98

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their glabella. Restored images of the holotype and of one of the paratypes of Q. zhengwanensis (Text-fig.13) show that its morphology agrees well with those of Baojingia.

A

C

D

Text-figure 13. Baojingia zhengwanensis (Yang in Yang et al., 1991) [=Quandraspis zhengwanensis Yang in Yang et al., 1991, the type species of Quandraspis]. A, D, deformed and computer-aided restored views of holotype cranidium, HXX 100348, both x 5 (original of Yang et al., 1991, pl. 18, fig. 9; reprinted 1993); B, C, deformed and computer-aided restored views of paratype cranidium, HXX100337a, both x 5 (original of Yang et al., 1991, pl. 18, fig. 10; reprinted 1993). Baojingia youshuiensis Yang in Zhou et al., 1977

Plate 38, figures 1-9; Text-figure 14 1977 1978 1982 1991 1993 2001b

Baojingia youshuiensis Yang in Zhou et al., p. 174, pl. 5 l, figs. 10, 11. Baojingia youshuiensis Yang; Yang, p. 48, 49, pl. 6, figs. 16-18. Baojingia youshuiensis Yang; Liu, p. 313, pl. 219, figs. 10, 25. Xichuania xiuzigouensis Yan in Yang et al., p. 154, 155, pl. 18, figs. 1, 2, ?3. Xichuania xiuzigouensis Yan in Yang et al., p. 199, 200, pl. 18, figs. 1, 2, ?3. Eoshengia youshuiensis (Yang); Peng, Babcock, and Lin, p. 103, pl. 6, figs. 12, 13.

Lectotype. Exfoliated cranidium (Zhou et al., 1977, pl. 51, fig. 10, CUGB 0112605; refigured by Yang 1978, pl. 6, fig. 16 and by Liu, 1982, pl. 219, fig. 19; Text-fig. 14 herein), from the Lisania tungjenensis Zone of the Huaqiao Formation, Huaqiao, Baojing, northwestern Hunan. This

cranidium was subsequently designated as holotype by Yang (1978). New material. More than 10 sclerites including including cranidia, librigenae, and pygidia •

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(illustrated specimens NIGP 138285-138291 ) in P249 and P261. Description. Cranidium width greater than length. Anterior border wide medially, and narrow

laterally, defined posteriorly by moderately deep, weakly wavy border furrow. Glabella straight-sided, tapering forward moderately, moderately convex, obtusely rounded anteriorly. Lateral glabellar furrows deep; S1 directed posterolaterally, weakly to strongly bifurcated; $2 directed posterolaterally; $3 transverse, isolated from axial furrow; $4 directed anterolaterally. Medial carina present on glabella. Occipital ring narrow at sides, wide medially, with small medial node located either slightly anterior to center of ring or anteriorly close to occipital furrow. Occipital furrow sinuous, wide and deep, with anterior margin curved forward medially. Fixigena moderately wide. Eye ridge distinct; palpebral lobe crescentic, moderately long, lying about cranidial midlength, defined by deep and wide ocular furrow. Anterior branch of facial suture converging gently forward; posterior branch transverse initially, curving rearward sharply distally. Librigena with genal spine; genal field moderately wide and moderately convex; lateral and posterior borders moderately wide and upturned; border furrows deep. Pygidium semicircular. Axis long and wide, with 5-6 tings and terminal piece, tapering gently onto posterior border furrow. Pleural field as wide as or slightly wider than axis, convex; pleural furrows groove-like; interpleural furrow effaced. Border flat, of uniform width. Surface covered with fine to moderately coarse, closely spaced granules on cranidium, librigena, and pygidium. Librigena with radiating and anastomosing ridges on genal field and terrace line on lateral border.

Text-figure 14. Baojingia youshuiensis Yang in Zhou et al., 1977. A, B, exfoliated cranidium in dorsal and anterolateral views, CUGB 0112605, × 6.5, designed subsequently as the holotype of the species (original of Zhou et al., 1977, pl. 51, fig. 10; also Yang, 1978, pl. 6, fig. 16). Remarks. The holotype (Text-fig. 14) and the paratype cranidia (Yang, 1978, pl. 6, fig. 18) of this •

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species are completely exfoliated. The specimens show four pairs of deeply impressed lateral glabellar furrows, with the S1 furrow being weakly sinuous. A weak carina, and non-granulate surface is also expressed in the type series. The occipital ring is damaged in the holotype, but present in the paratype cranidium. It is triangular in shape, and the posterior margin is angled obtusely. New material from the Huaqiao Formation at Paibi, Huayuan, Hunan, agrees with the type material in general respects but it also shows some morphological variation within the species. The testaceous surface of the species in the new material bears densely spaced granules, the S1 furrow is variable from weakly sinuous to strongly bifurcated, the carina is subtle to distinct, and the occipital ring varies in sagittal length. The new material adds information about the shape and segmentation of the pygidium. Occurrence. The holotype is from the Lisania tungjenensis Zone of the Huaqiao Formation, Huaqiao, Baojing County, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the upper part of the Lejopyge laevigata Zone through the basal part of the Proagnostus bulbus Zone). Baojingia jiudiantangensis (Yang in Zhou et al., 1977) Plate 39, figures 8, 9; Text-figure 15 1977 1977 1978 1978 1980 1982 1982 1983 ?1983

Lisania tungjenensis Nan; Zhou, Liu, Meng and Sun, p. 171, 172, pl. 51, fig. 3. Eoshengia jiudiantangensis Yang in Zhou et al., p. 174, pl. 51, fig. 16. Eoshengia jiudiantangensis Yang; Yang, p. 59, 60, pl. 8, fig. 3. Eoshengia jiudiantangensis Yang; Yin and Li, p. 493, pl. 166, fig. 6. Lisaniella sp. cf. L. elongata Chang; Ergaliev, pl. 6, fig. 4 (not described). Lisania tungjenensis Nan; Liu, p. 312, 313, pl. 219, fig. 26. Eoshengia jiudiantangensis Yang; Liu, p. 313, pl. 219, fig. 27. Lisania sp.; Lin, Lin, and Zhou, p. 402, 403, pl. 1, fig. 8. Eoshengia sp.; Lin, Lin, and Zhou, p. 403, pl. 1, figs. 14, ? 13.

Holotype. By monotypy; testaceous cranidium (Zhou et al., 1977, pl. 51, fig. 16, CUGB 1430206; refigured by Yang, 1978, pl. 8, fig. 3 and Liu, 1982, pl. 219, fig. 27; Text-fig. 15 herein) from the Huaqiao Formation at Jiudiantang, Xinhuang County, northwestern Hunan. New material. Single cranidium (NIGP 138299) in collection P269. Remarks. The cranidium conforms in all observable details with the holotype cranidium from the Huaqiao Formation of northwestern Hunan. Key features of this species are a forwardly arched anterior border furrow, slightly curved palpebral lobes located near the cranidial midlength, diagonally directed posterior branches of the facial suture enclosing narrow (tr.) posterolateral projections of the fixigena, and a lack of surface granulation. Occurrence. The holotype is from the Huaqiao Formation at Jiudiantang, Xinhuang County, northwestern Hunan, China. New material is from dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Proagnostus bulbus • 101.

Zone).

Text-figure 15. A, B, holotype of Baojingia jiudiantangensis (Yang in Zhou et al., 1977), slightly exfoliated cranidium in dorsal and anterolateral views, CUGB 1430206, x 2.5 (original of Zhou et al., 1977, pl. 51, fig. 16; also Yang, 1978, pl. 8, fig. 3). Baojingia latilimbata (Peng, 1987)

Plate 39, figures 1-7 1987 1987 1980 1991 1993

Eoshengia latilimbata Peng, p. 96, 97, pl. 8, fig. 7. Eoshengia curvata Peng, p. 97, pl. 8, fig. 8. Taitzuia? sp.; Ergaliev, pl. 6, fig. 3. Eoshengia shorteglabella Yang in Yang et al., p. 150, 151, pl. 16, fig. 11. Eoshengia shorteglabella Yang in Yang et al., p. 194, 195, pl. 16, fig. 11.

Holotype. Cranidium (Peng, 1987, pl. 8, fig. 7, NIGP 74536) from the Formosagnostus formosus [=Hadragnostus modestus]-Distazeris [=Paradistazeris] Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan. New material. Three cranidia and one pygidium (NIGP 138295-138298) in collections P178.3, P184.8, P200.7, and W121.6. Remarks. The holotype cranidium of Baojingia latilimbata from the Huaqiao Formation in Taoyuan, northwestern Hunan, is characterized by having a smooth surface, a parallel-sided, weakly carinate glabella beating three pairs of weak lateral glabellar furrows, and a wide (sag.) anterior border with sinuous anterior margin and a pair of small, weakly defined tubercles near the rear and close to the cranidial border furrow. The tubercles, however, are expressed variably in the new material. E. curvata is considered to be a junior synonym of E. latilimbata. It is represented by a broken cranidium, which is from the same bed and same collection as the holotype, and was differentiated from E. latilimbata only by its variable features, the narrower (tr.) anterior border and the less obliquely directed eye ridges (Peng, 1987, p. 97). One cranidium in the new material (P1. 39, fig. 1) is tectonically distorted and flattened. By using computer-graphic restoration technique (Hughes and Jell, 1992), a restored image of this cranidium (Pl. 39, fig. 2) has been achieved, showing that it is almost indistinguishable from the holotype cranidium of Baojingia latilimbata: key features, such as the W-shaped anterior cranidial •

102.

border furrow, the wide (sag.) anterior border beating tubercles posteriorly, and the presence of a weak carina are all identical. This technique suggests that Eoshengia shorteglabella Yang (in Yang et al., 1991), which is based on a distorted cranidium from the area of eastern Qinling-Dabashan Mountains, also should be synonymized.

Occurrence. The holotype is from the Huaqiao Formation, Formosagnostus formosus [=Hadragnostus modestus]-Distazeris [=Paradistazeris] Zone, Wa'ergang, Taoyuan, northwestem Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Baojingia paralala (Yang in Zhou et al., 1977) Plate 40, figures 1-18; Text-figure 16

Lisania paralala Yang in Zhou et al., p. 171, pl. 51, fig. 1. Lisania paralala Yang, p. 45, 46, pl. 6, figs. 9, 11. Lisania paralala Yang; Yin and Li, p. 491, pl. 165, fig. 13. Lisania subcylindrica Zhang, p. 170, pl. 64, fig. 1. Lisania paralala Yang; Liu, p. 312, pl. 219, fig. 18. Eoshengia subquadrata Yang; Peng, Babcock, and Lin, p. 102, pl. 5, figs. 9, 11.

1977 1978 1978 1981 1982 2001b

Holotype. By monotypy; exfoliated cranidium (Zhou et al., 1977, pl. 51, fig. 1, CUGB 0112104; refigured by Yang, 1978, pl. 6, fig. 9 and Liu, 1982, pl. 219, fig. 18; Text-fig. 16 herein), from the Lisania tungjenensis Zone, Huaqiao Formation, at Huaqiao, Baojing, northwestern Hunan.

.¢

t

i!

Text-figure 16. A, B, holotype of Baojingia paralala (Yang in Zhou et al., 1977), exfoliated cranidium in dorsal, anterior, and anterolateral views, CUGB 0112104, x 4 (original of Zhou et al., 1977, pl. 51, fig.l" also Yang, 1978, pl. 5, fig. 9).

New material. More than 40 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 138305-138317, 138417) in collections P204, P216, P223.7, and W146.2. Emended diagnosis. Baojingia having a glabella with parallel-sided or tapered, effaced or faintly furrowed glabella; transverse or gently yoked anterior border furrows; wide and upturned anterior • 103"

border; narrow (tr.), strongly inward-sloped palpebral area and wide (exs.) posterolateral projection of fixigena; and long and broadly-based librigenal spine.

Remarks. The holotype is almost completely exfoliated, the palpebral lobes are broken, and the facial sutures are unexposed. Rich new material from the same region in northwestern Hunan adds information on the palpebral lobes, the facial suture, and on the librigena and pygidium. It also shows variation in cranidial morphology. A cranidium and a pygidium in the new material were assigned previously as Eoshengia subquadrata (Peng, Babcock, and Lin, 2001b), and are now considered to be conspecific with Lisania paralala Yang (in Zhou et al., 1977). The latter is now regarded as a species belonging to Baojingia because of its similarity to effaced species of Baojingia, which were commonly assigned to Eoshengia previously. Key features of this species are the wide (sag.), upturned anterior border, the relatively narrow (tr.) palpebral areas of the fixigenae, the transverse to gently forwardly arched cranidial border furrow, the diverging anterior branches of the facial suture, the deflected posterior branch of the facial suture, the long occipital ring, the narrow (tr.) librigena with border furrows extending onto the genal spine, and the narrow (tr.) pygidial pleural fields. Occurrence. The holotype is from the Lisania tungjenensis Zone, Huaqiao Formation, at Huaqiao, Baojing County, northwestern Hunan, China. New material is from dark-gray limestone of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata Zone). Baojingia quadrata (Yang in Zhou et al., 1977) Plate 38, figures 10-13; Text-figure 17 1977 1977 1978 1978 1978 1978 1980 1981 ?1981 1981 1982 1982 ?1991 1991 ?1993 1993

Eoshengia quadrata Yang in Zhou et al., p. 174, pl. 51, figs. 14, 15. Eoshengia paragenalata Yang in Zhou et al., p. 174, pl. 51, figs. 17, 18, ? 19. Eoshengia quadrata Yang; Yang, p. 50, pl. 8, figs. 4, 5. Eoshengia paragenalata Yang; Yang, p. 51, pl. 8, figs. 9, 10, ?11. Eoshengia quadrata Yang; Yin and Li, p. 493, pl. 165, figs. 11, 12. Eoshengia paragenalata Yang; Yin and Li, p. 493, pl. 166, figs. 14, ?15. Lisanella elongataformis Ergaliev, p. 147, pl. 3, fig. 17. Eoshengia quadrata Yang; Zhang, p. 171, pl. 63, figs. 10, 11. Eoshengia tenuis Zhang, p. 172, pl. 63, figs. 12-14. Eoshengia xinjiangensis Zhang, p. 172, pl. 63, figs. 16-19. Eoshengia quadrata Yang; Liu, p. 313, pl. 219, figs. 11, 28. Eoshengia paragenalata Yang; Liu, p. 313, pl. 220, figs. 1, 2. Eoshengia paratenuis Yang in Yang et al., p. 149, pl. 16, figs. 5-8. Eoshengia rigida Yan in Yang et al., p. 151, pl. 16, figs. 12, 13. Eoshengia paratenuis Yang in Yang et al., p. 192, 193, pl. 16, figs. 5-8. Eoshengia rigida Yan in Yang et al., p. 195, pl. 16, figs. 12, 13.

Lectotype. Exfoliated cranidium (Zhou et al., 1977, pl. 51, fig. 14, CUGB 0329103; refigured by Yang, 1978, pl. 8, fig. 4 and Liu, 1982, pl. 219, fig. 11; Text-fig. 17 herein), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, at Laochatian, Fenghuang, northwestern Hunan. This cranidium was subsequently • 104.

designated as the holotype of the species by Yang (1978). N e w material. Four cranidia and one pygidium (illustrated specimens NIGP 138292-138294) in

collections P261.35, W 173.8, W 189, W 196.3. Remarks. New material from the Huaqiao Formation, northwestern Hunan, resembles closely the

type material, which comes from the same region. Key features of this species are a proportionally long and narrow, subparallel-sided glabella, and a relatively narrow (sag.) anterior border.

Text-figure 17. A-F, Baojingia quadrata (Yang in Zhou et al., 1977). A, B, exfoliated cranidium in dorsal and anterolateral views, originally a syntype of Eoshengia paragenalata Yang in Zhou et al., 1977 (pl. 51, fig. 17) and subsequently designed as the holotype of E. paragenalata by Yang (1978, p. 51, pl. 8, fig. 9), CUGB 0329101, × 3; C, D, slightly exfoliated cranidium from syntypes of Baojingia quadrata, designed subsequently as the holotype of this species by Yang, 1978 (p. 50, 80), in dorsal and

anterolateral views, CUGB 0329103, × 3.5 (original of Zhou et al., 1977, pl. 51, fig. 14; also Yang, 1978, pl. 8, fig. 4); E, F, mostly exfoliated pygidium from syntypes of B. quadrata, in dorsal and obliquely anterior views, CUGB 0329109, × 3 (original of Zhou et al., 1977, pl. 51, fig. 15; also Yang, 1978, pl. 8, fig. 5). •

105

•

Following Yang (in Zhou et al., 1977; Yang, 1978), this species is included in Eoshengia (now Baojingia). Baojingia quadrata seems to be morphologically similar to Wanshania wanshanensis, and may be congeneric with that species. It agrees almost in all the cranidial and pygidial respects except for having a transverse, strongly curved anterior border furrow, and thus a longer glabella.

Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, at Laochatian, Fenghuang County, northwestern Hunan China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone).

Baojingia subquadrata (Yang, 1978) Plate 41, figures 1-12; Plate 42, figures 1-14; Text-figure 18 1978 1978 1978 1978 1999

Eoshengia subquadrata Yang in Yin and Li, p. 493, pl. 166, figs. 12, 13. Eoshengia spinosa Yang in Yin and Li, p. 493, pl. 166, figs. 16, 17. Eoshengia subquadrata Yang; Yang, p. 49, pl. 8, figs. 1, 2. Eoshengia spinosa Yang; Yang, p. 50, 51, pl. 8, figs. 6-8. Eoshengia cf. E. spinosa Yang; Duan, Yang, and Shi, p. 157, figs. 9A-D. 2001b Eoshengiaspinosa Yang; Peng, Babcock, and Lin, p. 104, pl. 9, fig. 11. non 2001b Eoshengiasubquadrata Yang, Peng, Babcock, and Lin p. 102, pl. 5, figs. 9, 11 [=Baojingia paralala (Yang in Zhou et al., 1977)]. 2001b Eoshengiasp.; Peng, Babcock, and Lin, p. 104, pl. 10, figs. 14, 15. Holotype. Exfoliated cranidium (Yang, 1978, pl. 8, fig. 1, CUGB 114103; Text-fig. 18E herein), from the Paradamesops [=Parablackwelderia] jimaensis-Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, at Laochatian, Fenghuang, northwestern Hunan.

New material. More than 50 sclerites including an incomplete exoskeleton, cranidia, librigenae, and pygidia (illustrated specimens NIGP 138318-138336) in collections P225.5, P230.5, P249, P250, P251, P260. P264.2, P273.66, P273.8, P277, P298.4, P301, P301.9, P307.7, P331.8, W199.2, W200.3, W210.45, W210.5, W211.7, W212.95, and W216.5.

Remarks. Eoshengia subquadrata is now transferred to Baojingia. Originally it was designated as the type species of Eoshengia. The holotype of E. subquadrata is a holaspid cranidium 10 mm in length. It is the only cranidium known for the species. It was described as lacking occipital spines. However, the left half of the occipital ring was damaged (Text-fig. 18A), and it is hard to say if the spine as originally absent on the holotype. E. subquadrata Yang (1978, pl. 8, figs. 1, 2) is similar in morphology to E. spinosa Yang (1978, pl. 8, figs. 6-8). Even the surface ornamentation of both species is identical. According to Yang (1978, p. 51), E. spinosa differs in having a more convex glabella, an occipital furrow connected with the dorsal furrow laterally, an undulating anterior cranidial furrow, and in the presence of an occipital spine. However, examination of the holotype shows that the first three differences are invalid, and the last one is uncertain. Yang (1978) differentiated the pygidium of E. spinosa from that of E. subquadrata by the relatively greater length, the more segmented axis, and the absence of anterior curvature (sag.) on the posterior margin. •

106.

New material from the Huaqiao Formation, northwestern Hunan, shows that a range of character states exist in E. spinosa, and the range is sufficiently wide as to warrant accomodating E. spinosa within E. subquadrata. Some specimens lack an occipital spine (P1. 42, fig. 3) and others possess it; also some specimens are strongly granulated on the external surface and other specimens are only partially granulated (P1. 42, figs. 7, 8). The type specimens of both species seem to be identical in all respects other than these. The two species occur in the same horizons and in the same localities in northwestern Hunan.

Text-figure 18. A-G, Baojingia subquadrata (Yang, 1978). A-C, holotype cranidium in dorsal view and paratype pygidium in dorsal and anterolateral views, CUGB 1104103, 1104101, all x 4 (original of Eoshengia subquadrata Yang, 1978, pl. 8, figs. 1, 2); D-G, cranidium in anterolateral and dorsal views and pygidium in dorsal and posterolateral views, CUGB 1207206, 1206201, x 3.2, x 2.5; originally assigned respectively as holotype and paratype of Eoshengia spinosa Yang, 1978 (p. 50, 80).

Occurrence.

The holotype

is

from

the

Paradamesops

[=Parablackwelderia] jimaensis• 107.

Cyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, at Laochatian, Fenghuang County, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the lower part of the Pianaspis sinensis Zone to the lower part of the Liostracina bella Zone (equivalent to the upper part of the Lejopyge laevigata Zone through the Linguagnostus reconditus Zone). Baojingia tungjenensis (Nan in Egorova et al., 1963)

Plate 39, figures 10-14 Lisania tungjenensis Nan in Egorova et al., p. 34, 35, pl. 6, figs. 8-10, 12, ?11. Lisania tungjenensis Nan; Lu, Zhang, Zhu, Qian, and Xiang, p. 265,266, pl. 45, figs. 16-18. Lisania tungjenensis Nan; Zhou, Liu, Meng and Sun, p. 171, 172, pl. 51, fig. 3 [=Baojingia jiudiantangensis (Yang in Zhou et al., 1977)]. 1978 Lisania tungjenensis Nan; Yin and Li, p. 492, pl. 165, fig. 2. non 1982 Lisania tungjenensis Nan; Liu, p. 312, 313, pl. 219, fig. 26 [=Baojingia jiudiantangensis (Yang in Zhou et al., 1977)]. 1991 Lisania tungjenensis Nan; Lin, p. 376, pl. 3, figs. 1, 2. Quandraspis shengwanensis Yang in Yang et al., p. 156, pl. 16, fig. 14; pl. 18, figs. 9-11. 1991 1993 Quandraspis shengwanensis Yang in Yang et al., p. 202, pl. 16, fig. 14; pl. 18, figs. 9-11. 2001b Lisania yuanjiangensis (Yang); Peng, Babcock, and Lin, p. 102, pl. 4, fig. 3. non 2001b Lisania tungjenensis Nan; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 14 [=Lisania paibiensis sp. nov.].

1963 1965 non 1977

Lectotype. Cranidium (Egorova et al., 1963, pl. 6, fig. 8; GMC IV2631) from the Huaqiao Formation near Huangbaizhen, Tongren, eastern Guizhou; designated herein. New material. Three cranidia and one pygidium (NIGP 138300-138304) in collections P 126, 190.7, and P2 04. Remarks. Two specimens, a cranidium and a pygidium in the type collection (Egorova et al., 1963, pl. 6, figs. 8, 12) were designated as holotypes. The syntype cranidium is selected herein as the lectotype of the species. The species was originally assigned to Lisania, but its morphology suggests a classification with Baojingia. B. tungrenensis is a weakly furrowed species of Baojingia. As diagnosed by Nan (in Lu et al., 1965), the species is characterized by the presence of three pairs of weakly impressed lateral glabellar furrows, a subtriangular occipital ring bearing both a node and a spine, relatively wide fixigenae, and a conical pygidial axis with five tings. The present material resembles closely the type material in most cranidial and pygidial respects. The number of lateral glabellar furrows counted by Nan may be erroneous; all species of Baojingia have four pairs of glabellar furrows. The type material has been illustrated at least three times, but always was poorly illustrated, showing no details of such furrows. However, previous illustrations show that there is a pair of rounded lobes isolated from the anterolateral comers of the L1 furrows of the glabella by an x-shaped S1. Such auxiliary lobes are variably presented in the present material. The present material has a relatively narrow (sag.) occipital ring, and the shape of occipital ring is also variable. The only feature that differentiates the type cranidia from the new material is the greater sagittal convexity of the types. This difference may reflect population-level differences, or may be the • 108•

result of taphonomic influences. Occurrence. The lectotype is from the Huaqiao Formation near Huangbaizhen, Tongren, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it is occurs in association with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the lower part of the Goniagnostus nathorsti Zone through the Lejopyge laevigata Zone).

Genus NEOANOMOCARELLAHsiang in Egorova et al., 1963 Neoanomocarella Hsiang in Egorova et al., 1963, p. 55; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 335; Zhou, Liu, Meng, and Sun, 1977, p. 183; Yin and Li, 1978, p. 504, 505; Yang, 1978, p. 52; Liu, 1982, p. 314; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 157; Peng, 1987, p. 99; Duan, Yang, and Shi, 1999, p. 155. Type species. Neoanomocarella asiatica Hsiang in Egorova et al., 1963 (p. 55, pl. 12, figs. 9-11) from the Huaqiao Formation, Fenghuang, northwestern Hunan (in association with Liostracina bella); by original designation. Other species. Neoanomocarella incilis sp. nov. All previously described species from Hunan (Yang in Zhou et al., 1977; Qiu in Qiu et al., 1983; Peng, 1987) are regarded as junior synonyms of N. asiatica. Emended diagnosis. Lisaniidae with convex anterior cranidial border beating transverse depression medially and plectrum posteriorly; glabella largely effaced, tapering forward gently or subrectangular in outline, with occipital node located anteriorly; palpebral lobe moderately small, located subcentrally. Librigena with narrow (tr.) genal field; lateral border wide, posterior border rather wide, with border furrow-like depression continuing the depression on anterior border of cranidium and extending onto genal spine, genal spine broad at base. Pygidium transverse, axis with four segments, extending nearly to border furrow; lateral and posterior borders wide, upturned; pleural field weakly segmented. Surface smooth or finely granulated. Remarks. This genus is here regarded as a plectrum-bearing lisaniid, and is rediagnosed based on new material from northwestern Hunan, which is the region from which it was first described. The genus was erected on the basis of cranidia alone. Because of similarity with the cranidium of Anomocarella, Xiang (in Egorova et al., 1963) classified this genus within the Anomocarellidae Hup6, 1953. Such an assignment was accepted by later authors (Lu et al., 1965; Yang in Zhou et al., 1977; Yin and Li, 1978; Yang, 1978; Qiu et al., 1983; Peng, 1987; Duan et al., 1999). New information concerning the pygidium, librigena, and cranidium suggests that it is more appropriate to refer Neoanomolocarella to the Lisaniidae. The transverse pygidium, which has an axis occupying most of the pygidial length (sag.), has a strong lisaniid aspect, and overall features of the cephalon are closely comparable with such lisaniid genera such as Lisania, Baojingia, and Shengia. The shape of glabella, the pattern of the lateral glabellar furrows, the subcentrally located palpebral lobes, the trend of the eye ridge (subparallel to the anterior border), the deflected posterior branches of the facial suture, and the wide fixigenae resemble those features in Baojingia and Shengia. The presence of a plectrum is similar to Baojingia, which bears a plectrum-like extension defined by a W-shaped anterior border furrow. In spite of the similarities to other lisaniids, Neoanomocarella • 109"

remains a distinctive genus by virtue of a convex, upturned anterior cranidial border, relatively small palpebral lobes, short posterolateral projections of the fixigenae, and effaced pleural fields of the pygidium. Neoanomocarella asiatica Hsiang in Egorova et al., 1963

Plate 43, figures 1-13; Plate 44, figures 1-18; Text-figure 19 1963 1965

Neoanomocarella asiatica Hsiang in Egorova et al., p. 55, 56, pl. 12, figs. 9-12. Neoanomocarella asiatica Hsiang; Lu, Zhang, Zhu, Qian, and Xiang, p. 335,336, pl. 62, figs.

1977 1977 1977 1978 1978 1978 1978 1978 1980 1982 1982 1983

Neoanomocarella asiatica Hsiang; Zhou, Liu, Meng, and Sun, p. 183, pl. 54, figs. 3, 4.

20-22. Neoanomocarella hunanensis Yang in Zhou et al., p. 183, pl. 54, fig. 5. Neoanomocarella quadrata Rong and Yang in Zhou et al., p. 183, pl. 54, figs. 6, 7. Neoanomocarella asiatica Hsiang; Yin and Li, p. 505, pl. 169, figs. 18, 19. Neoanomocarella quadrata Rong and Yang; Yin and Li, p. 505, pl. 169, fig. 12. Neoanomocarella asiatica Hsiang; Yang, p. 52, pl. 8, figs. 15-17. Neoanomocarella hunanensis Yang; Yang, p. 52, 53, pl. 8, fig. 18. Neoanomocarella quadrata Rong and Yang; Yang, p. 53, pl. 9, fig. 1. Neoanomocarella Neoanomocarella Neoanomocarella Neoanomocarella

asiatica Hsiang; Ergaliev, p. 183, pl. 5, figs. 17, 18. hunanensis Yang; Liu, p. 314, pl. 220, fig. 8. quadrata Rong and Yang; Liu, p. 314, pl. 220, figs. 6, 16. asiatica Hsiang; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and

Wei, p. 157, pl. 50, fig. 12. 1983 1983 1987 1987 ?1999

Neoanomocarella brevis Qiu in Qiu et al., p. 158, pl. 50, fig. 11. Neoanomocarella elongaa Qiu in Qiu et al., p. 158, pl. 51, figs. 7, 8. Neoanomocarella asiatica Hsiang; Peng, p. 99, pl. 7, figs. 13, 14; pl. 8, figs. 1-3.

200 lb

Neoanomocarella intermedia Peng, p. 99, 100, pl. 8, figs. 4-6. Neoanomocarella asiatica Hsiang; Duan, Yang, and Shi, p. 155, fig. 4H. Neoanomocarella asiatica Hsiang; Peng, Babcock, and Lin, p. 103, pl. 7, figs. 10, 11.

2001 d

Neoanomocarella asiatica Hsiang; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.2.

Holotype. Cranidium (Egorova et al., 1963, pl. 12, fig. 9, GMC IV2612) from the Huaqiao

Formation, near Chatian, Fenghuang, western Hunan. N e w material. More than 100 sclerites including cranidia, librigenae, and pygidia (illustrated

specimens NIGP 138337-138361), in collections P261.35, P298.4, P301.9, P319.6, P319.8, P331.8, P344.6, P348, P363.5, W 196.3, W 199.2, W210.5, W212.45, W215.1, W216.5, W218.3, W221.5, W223, W225, W227, W228.3; W229.9, W243.6, W251.15, and W254.1. Description. Cranidium subquadrate; length subequal to width. Anterior border with medial

transverse depression, upturned slightly at front; plectrum present, weak to distinct. Glabella elongate, tapered forward, truncate to slightly concave anteriorly. Lateral glabellar furrows consist of four pairs; furrows nearly effaced on testaceous surface, weak on exfoliated surface; S1 bifurcated; $2 straight, directed inward and slightly rearward; $3 short, straight; $4 short, directed inward and forward. Occipital furrow well impressed, curved gently rearward, with pair of short furrows that almost isolate pair of small anterolateral lobes; occipital ring wide medially, narrowing towards sides, with small median node located anteriorly. Palpebral lobe moderately long; eye ridge • 110.

nearly effaced. Anterior branch of facial suture diverges forward about 90 ° to anterior border furrow, turns inward and forward sharply toward anterior margin; posterior branch diverges strongly rearward, then deflects rearward sharply and gently inward to posterior margin. Librigena with narrow (tr.), outward-sloping genal field defined by well-impressed, gently curved border furrow; lateral border broad with convex inner portion and upturned outer portion; a border furrow-like depression present, connecting with the depression on anterior cranidial border and extending far onto genal spine; genal spine broad at base.

Text-figure 19. Reconstruction of cephalon and pygidium of Neoanomocarella asiatica Hsiang in Egorova et al., 1963. Cephalon based mostly on specimens NIGP 138347, 138350 and 138351 (see P1. 44, figs. 1, 4, 5); pygidium based mostly on specimen NIGP 138356 (see P1.44, fig. 11). Pygidium subelliptical, transverse. Axis subcylindrical, slightly tapered, convex; with four tings and short terminal piece. Pleural field with 3-4 interpleural furrows, pleural furrow present on first 1-2 pleurae, weak. Border moderately narrow, convex. Dorsal surface usually smooth; pygidium may have unevenly distributed fine granules. Terrace lines on lateral and posterior borders of pygidium, subparallel to margin. R e m a r k s . New material from the Huaqiao Formation at Paibi and Wangcun agrees with the type

and other specimens, which are from the same formation in the same paleogeographic area (Egorova et al., 1963; Yang in Zhou et al., 1977; Yin and Li, 1978; Peng, 1987). Key characters of the new material that indicates assignment to this species include the transverse depression on the anterior border, the presence of a plectrum, the obscure lateral glabellar furrows on both external and exfoliated surfaces, the shallow occipital furrow, and the short (tr.) posterolateral projection. •

111.

The large collection shows wide variation in cranidial morphology, and enables us to synonymize Neoanomocarella hunanensis Yang in Zhou et al., 1977, N. quadrata Rong and Yang in Zhou, 1977, and N. intermedia Peng, 1987, all from the same area as N. asiatica. Stated differences between these species and N. asiatica (for example, the proportional length or strength of the plectrum, the relatively broader glabella, the continuity of the anterior border furrow, and the strength of the transverse depression on the anterior border) fall well within the range of variation of N. asiatica. N. brevis and N. elongata, both from the Yangliugang Formation of southern Anhui (Qiu in Qiu et al., 1983), are also synonymized. Stated distinctions of these two species are apparently based on distorted features. The present collection shows a morphologic distinction between the external and exfoliated surfaces of the eye ridges. The eye ridge is faint on testaceous cranidia, but clearly defined on exfoliated surfaces. Occurrence. The holotype is from the Huaqiao Formation, near Chatian, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone through the upper part of the Liostracina bella Zone (equivalent to the lower part of the Proagnostus bulbus Zone through the lowermost part of the Glyptagnostus reticulatus Zone). Neoanomocarella incilis sp. nov. Plate 45, figures 1-9 Etymology. From Latin, incile, furrow, referring to the incised or clearly defined lateral glabellar and occipital furrows on the exfoliated surface. Holotype. Cranidium (P1.45, figs. 2-4, NIGP 138363) from collection W188.8. Other material. Three cranidia in collections W187.8 and W188.8 are illustrated as paratypes (NIGP 138362, 138364, 138365); a fragmental cranidium in collection W 189. Diagnosis. Neoanomocarella with long, subcylindrical glabella bearing weak lateral furrows, S1 bifurcated; exfoliated surface with clearly defined lateral furrows; cranidial surface densely granulate. Description. Cranidium subquadrate, length subequal to width. Glabella subcylindrical, twice as long as wide, truncate anteriorly, with sides expanded slightly and front slightly concave. Anterior border with medial transverse depression, developed variably; plectrum depressed and poorly defined. Lateral glabellar furrows consists of four pairs, weak on external surface but incised on exfoliated surfaced: S1 bifurcated; $2 straight, diverted inward and slightly rearwardly directed, located opposite midlength of glabella (excluding occipital ring). Occipital furrow well impressed, curved gently rearward, with pair of short furrows that almost isolate pair of small anterolateral lobes from the occipital ring. Palpebral lobe extending from level of anterior one-third of L1 to middle of L2; eye ridge obscure or clearly defined on exfoliated surface. Anterior branch of facial suture diverges forward about 90 ° to anterior border furrow, turns inward and forward sharply in gentle arch to anterior margin; posterior branches diverges strongly rearward, then deflects • 112.

rearward sharply and gently inward to posterior margin. Cranidial surface with densely spaced, fine granules. Thorax and pygidium unknown.

Remarks. Neoanomocarella incilis sp. nov. differs from N. asiatica, the type species of the genus, in the shape of glabella (subcylindrical rather than tapered forward), the presence of granulate prosopon, the better impressed occipital furrow, the weak rather than obscure eye ridge, and the incised rather than obscure lateral glabellar furrows on exfoliated surfaces.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone). Genus QIANDONGASPIS Yuan and Yin, 1998

Qiandongaspis Yuan and Yin, 1998, p. 145, 146. Type species. Qiandongaspis qiandongensis Yuan and Yin, 1998 (p. 146, 147, pl. 2, figs. 1-8), a junior synonym of Rhyssometopus sinensis Peng, 1987 (p. 100, pl. 8, fig. 12, text-fig. 13), from the Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone of the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou; by original designation. Other species. Qiandongaspis laevis sp. nov., Q. convexa sp. nov., Q. xiangxiensis sp. nov., and Q. sp., all from the Huaqiao Formation of northwestern Hunan, described below. Two other species assigned previously to Lisania are tentatively transferred to Qiandongaspis. They are Lisania guizhouensis Lee and Yin in Yin and Li, 1978 (p. 491, pl. 165, fig. 1) from the Pingjing Formation, Wuchuan, northern Guizhou, and Lisania taihangshanensis Zhang and Wang, 1985 (p. 410, pl. 123, fig. 2) from the Hsuchuang Formation, near Wuan, southern Hebei.

Remarks. Qiandongaspis qiandongensis, the type species of Qiandongaspis, is a junior synonym of Rhyssometopus sinensis Peng, 1987. The monospecific genus Qiandongaspis was erected by Yuan and Yin (1998, p. 146, 147) on the basis of material from the Huaqiao Formation (formerly Chefu Formation) of eastern Guizhou. Yuan and Yin (1998) considered the genus and the type species to be closely similar to Rhyssometopus Opik, 1967 and referred the taxa to the Rhyssometopidae, but they apparently overlooked the description of R. sinensis Peng (1987). R. sinensis is from a nearly equivalent horizon of the same formation in the same general region as Q. qiandongensis. Type material of both R. sinensis and Q. qiandongensis is reposited in the Museum of the Nanjing Institute of Geology and Palaeontology. Examination of the materials shows that there is no morphological difference between the species; thus Q. qiandongensis is suppressed as a junior synonym of R. sinensis. The genetic concept of Qiandongensis (Yuan and Yin, 1998) is followed here, and R. sinensis is transferred to Qiandongaspis. Qiandongaspis is characterized by having a normal, marginal genal spine on the librigenae, having a pair of fossulae, having bifurcated S1 lateral glabellar furrows, and by lacking lateral glabellar bulges. Rhyssometopus differs from Qiandongaspis in having a transmarginal genal spine that interrupts the lateral and posterior border furrows, lacking a pair of fossulae in the axial furrows, having a pair of lateral bulges on the glabella, and having non-bifurcated S1 lateral • 113.

glabellar furrows. The pygidium of Qiandongaspis, if correctly associated (Yuan and Yin, 1998, pl. 2, figs. 3, 5), differs from that of Rhyssometopus in having a subconical rather than cylindrical, proportionally shorter axis with flat rather than upturned borders. Overall in morphology Qiandongaspis is more similar to lisaniids, especially Lisania and Redlichaspis Kobayashi, 1935 [=Lisanella Chang, 1963], than it is to species of the Rhyssometopidae. Thus, Qiandongaspis is here classified in Lisaniidae. Lisania is differentiated from Qiandongaspis by having shorter, weakly defined palpebral lobes and more effaced lateral glabellar furrows. Redlichaspis (sensu Zhang and Jell, 1987, p. 137, 138), another lisaniid, differs from Qiandongaspis in having a proportionally wider, subquadrate cranidium with more slanting palpebral lobes, wider fixigenae, and an occipital spine. Two new species of Qiandongaspis, Q. convexa sp. nov. and Q. xiangxiensis sp. nov., are described below. Besides Q. sinensis, Q. laevis, and Q. xiangxiensis, this genus probably includes two more species that were originally referred to Lisania: L. guizhouensis Lee and Yin in Yin and Li, 1978 and L. taihangshanensis Zhang and Wang, 1985. The relatively long, well-defined, posteriorly located palpebral lobes, and the narrow palpebral area, suggest that these two species are more likely to have an affinity with Qiandongaspis.

Qiandongaspis sinensis (Peng, 1987) Plate 46, figures 1-13; Plate 47, figures 1-4 1987 Rhyssometopus sinensis Peng, p. 100, pl. 8, fig. 12, text-fig. 13. 1998 Qiandongaspis qiandongensis Yuan and Yin, p. 146, 147, pl. 2, figs. 1-8. 2001b Rhyssometopus sp. cf. R. sinensis Peng, Babcock, and Lin, p. 102, pl. 3, fig. 3. 2001b Rhyssometopus sp., Peng, Babcock, and Lin, p. 102, pl. 5, fig. 8. 2001b Rhyssometopid gen. et sp. nov., Peng, Babcock, and Lin, p. 103, pl. 7, fig. 16. 2001d Rhyssometopus sinensis Peng; Peng, Babcock, Lin, and Chen, p. 141, fig. 9.5.

Holotype. By monotypy; partly exfoliated cranidium (Peng, 1987, pl. 8, fig. 12, NIGP 74541) from the Formosagnostus formosus [=Hadragnostus modestus]-Distazeris [=Paradistazeris] Zone, Huaqiao Formation, at Wa'ergang, Taoyuan, northwestern Hunan.

New material. More than 30 cranidia (illustrated specimens NIGP 138372-138385) in collections P108, P135, P204, P260, P261.5, P268.3, P269, P298.4, W196.3, W197.6, and W215.1.

Remarks. Comparison of the holotypes of both Rhyssometopus sinensis and Qiandongaspis qiandongensis show that they are identical in all details. As discussed above under the generic concept, R. sinensis is now transferred to Qiandongaspis and is regarded as a senior synonym of Q. qiandongensis, the type species of Qiandongaspis. New material from the Huaqiao Formation of northwestern Hunan shows a wide morphological variation in the shape of glabella in Q. sinensis. The glabella varies from tapered slightly forward or parallel-sided with a truncate front, sinuous with a weak concavity medially, to obtusely rounded. Variation in the effacement of lateral glabellar furrows is also present. The preglabellar area varies in sagittal length, and the anterior border varies in the degree of forward-curvature. A weak carina is present on some specimens; the carina is more clearly defined on exfoliated cranidia.

Occurrence. The holotype is from the Formosagnostus formosus [=Hadragnostus modestus]• 114.

Distazeris [=Paradistazeris] Zone, Huaqiao Formation, Wa'ergang, northwestern Hunan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Dorypyge richthofeni Zone to the upper part of the Wanshania wanshanensis Zone (equivalent to the Ptychagnostus punctuosus Zone through the lower part of the Linguagnostus reconditus Zone). Qiandongaspis convexa sp. nov. Plate 47, figures 10-17 2001b Lisaniaagonius (Walcott); Peng, Babcock, and Lin, p. 102, pl. 4, figs. 2, 4.

Etymology. From Latin convexus, smooth, referring to the gently convex anterior border. Holotype. Cranidium (P1.47, figs. 10, 15, NIGP 138390) from collection P 149.5. Other material. More than 20 cranidia (illustrated paratypes NIGP 138391-138396) in collections P123.6 and P149.5. Diagnosis. Qiandongaspis with gently tapered, anteriorly truncate, faintly furrowed glabella, and transverse occipital furrow. Anterior border gently convex, moderately wide. External surface smooth. Description. Anterior border crescentic, gently convex, rather wide (tr., sag.); anterior border furrow curved evenly forward, rather wide, well impressed. Glabella subrectangular, tapered gently forward, length about twice width, truncate anteriorly, with lateral glabe~lar furrows largely effaced; S1, or sometimes S land $2, faintly impressed; S 1 bifurcate with long, straight, diagonally directed posterior branch and short, transverse anterior branch; $2 moderately long, very gently curved, directed inward and slightly rearward. Occipital furrow shallow and rather wide, transverse, with distal ends curving slightly forward; occipital ring longest sagittally, narrowing laterally, beating tiny median node located subcentrally. Palpebral lobe moderately large, located posteriorly, with posterior end opposite middle of L1 lobe and anterior end opposite $2 furrow; palpebral furrow shallow; palpebral area narrow (tr.), gently convex, with inner portion strongly inclined to axial furrow. Anterior branch of facial suture diverging forward at about 50 ° to anterior border furrow, then turning sharply inward and forward to cross anterior border at about 90 ° with ot point opposite anterolateral comer of glabella; posterior branch diverging strongly about 120 °, enclosing triangular posterolateral projections. Surface smooth. Remarks. Qiandongensis convexa closely resembles Q. sinensis in general respects, but differs in having a convex rather than upturned anterior border, relatively shorter palpebral lobes, and a transverse rather than rearward-bowed occipital furrow, and in lacking granulate ornamentation. In addition, the new species can be differentiated by the course of the anterior branch of the facial suture. In the new species both the ~,-13and the 13-c~sections of the anterior branch are straight, but in Q. sinensis, the ~/-13-otsections are continuously curved. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it 115•

occurs in association with trilobites of the Pianaspis sinensis Zone (equivalent to the lowermost part of the Goniagnostus nathorsti Zone through the resembles Lejopyge laevigata Zone).

Qiandongaspis xiangxiensis sp. nov. Plate 47, figures 5-9

2001b Lisaniabura (Walcott); Peng, Babcock, and Lin, p. 102, pl. 4, fig. 1. Etymology. From Chinese, xiangxi, northwestern Hunan. Holotype. Cranidium (P1.47, figs. 8, 9, NIGP 138389) from collection P167. Paratypes. Three cranidia (NIGP 138386-138388) in collections P123.6, P130.5, and P149.5. Diagnosis. Qiandongaspis with subrectangular glabella that bears notch anteriorly. Anterior border moderately narrow (sag., exs.) and gently convex; preglabellar field absent; palpebral lobe moderately arcuate; palpebral area about one-fourth of glabellar width. Surface finely granulated.

Description. Anterior border bar-like, curved forward gently, defined posteriorly by deeply incised anterior border furrow. Glabella subrectangular, tapered forward gently, length about twice width, obtusely rounded anteriorly; frontal lobe beating faint median notch anteriorly; with four pairs of shallow lateral glabellar furrows; S 1 longest, bifurcated; $2 long, slightly oblique rearward; $3 long, slightly oblique forward; $4 short, spaced closely to $3, directed forwardly and inwardly. Occipital furrow shallow, transverse, wide medially, narrowing abaxially; occipital ring crescentic, with large median node centrally. Palpebral area gently convex, inclined to axial furrow, about one-fourth as wide as glabella; palpebral lobe arcuate, close to axial furrow, defined by moderately deep palpebral furrow, located posterior to cranidial midlength, with anterior end opposite midpoint of L1 and posterior end opposite midpoint of L3. Preocular area of fixigena small. Anterior branch of facial suture short, diverging slightly forward about 30 ° with ot point opposite anterolateral comer of glabella; posterior branch of facial suture diverging strongly about 120 °, enclosing blade-like posterolateral projection. Surface covered with fine, dense granules.

Remarks. One cranidium (P1.47, fig. 7) referred previously to Lisania bura is reassigned here to the new species. Additional material shows that this species is most comparable with Qiandongaspis in general morphology, especially in the more closely spaced, relatively long palpebral lobes, the clearly defined lateral glabellar furrows, the short (tr.) eye ridges, and the surface ornamentation of fine, dense granules. Q. xiangxiensis sp. nov. differs from Q. sinensis (Peng) in having a shorter (sag.) anterior border, a relatively shorter but more curved palpebral lobe, a relatively wider palpebral area, and an anteriorly incised glabella. The new species also resembles Q. convexa sp. nov., but Q. convexa is differentiated by a wider, gently convex anterior border, a more tapered glabella, a wider (tr.) palpebral area, and by the absence of surface granules.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites of the lower part of Pianaspis sinensis Zone (equivalent to the lowermost part of the Goniagnostus nathorsti Zone through the Lejopyge laevigata Zone). • 116.

Qiandongaspis sp. Plate 37, figure 15

Material. A single, mostly exfoliated cranidium (NIGP 138284) in collection W 196.3. Remarks. This cranidium has a subquadrate outline, a broad glabella that is strongly convex (sag., tr.) and slightly expanded laterally, a thick upturned border, a rather wide (tr.) preocular field of the fixigena that is enclosed by a forwardly directed anterior branch of the facial suture, and, as shown by the remainder of the test on the anterolateral comer, a granulate surface. The illustrated specimen is most similar to Qiandongaspis sinensis, but it is too incomplete to assign it confidently to species. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites of the upper part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Proagnostus bulbus Zone). Genus SHENGIA Hsiang in Egorova et al., 1963

Type species. Shengia quadrata Hsiang in Egorova et al. (1963, p. 57, 58, pl. 13, figs. 1-11), from the Glyptagnostus reticulatus Zone, Huaqiao Formation, northwestern Hunan; by original designation. Other species. See species listed by Shergold et al. (2000, p. 613). Among species in Shergold's (2000) list, Shengia genalata Lu (in Lu et al., 1963, p. 28, pl. 3, fig. 1) should be indicated as being from the Torsuqtagh Formation (upper Cambrian) of Kuruktag, rather than Xibei, Xinjiang, China. Shengia intermedia (Resser and Endo, 1937) and Shengia intermedia Lin and Zhang in Zhu et al. (1979) both of which are in Shergold's (2000) list of Shengia species, are homonymous. Shengia sp. cf. S. spinosa Yang (Shergold et al., 2000, p. 613) from the Val d'Homs Formation of southern France may not belong to the genus but to Lisania. To replace the junior homonym S. intermedia Lin and Zhang (in Zhu et al., 1979), the new species name Shengia shergoldi is proposed here. Remarks. The concept of Xiang in Egorova et al. (1963) is followed here. This genus is known only from South China and Northwest China (Hunan, Guizhou, Anhui, Liaoning, Qinghai, Xinjiang, Shaanxi). Shergold et al. (2000, p. 613) considered Eoshengia Yang in Zhou et al., 1977 and Shengia to be synonyms, but close comparision reveals that Eoshengia is more likely an effaced Baojingia, and is here regarded as a junior synonym of that genus. Shengia cf. spinosa Yang reported from southern France (Shergold et al., 2000) possibly belongs to Lisania. The species that was from southern Kazakhstan referred to Lisanella by Ergaliev (1980) but regarded as a representive of Shengia by Shergold et al. (2000) more likely belongs to Baojingia judging from its morphology and stratigraphic occurrence (Lejopyga laevigata Zone and Kormagnostus simplex Zone).

• 117.

Shengia quadrata Hsiang in Egorova et al., 1963

Plate 48, figures 1-10 1963

Shengia quadrata Hsiang in Egorova et al. (in part), p. 57, pl. 13, figs. 1 (cranidia only), 2-6, 11; non fig. 1 (pygidium only) [=Proceratopyge fenghwangensis Hsiang], non figs. 7-10 [=?Shengia trapezia Peng]. 1965 Shengia quadrata Hsiang; Lu, Zhang, Zhu, Qian, and Xiang (in part), p. 275, pl. 47, figs. 12, 14, 15 (cranidia only); non fig. 13 [=Shengia trapezia Peng], non fig. 15 (pygidium only) [=Proceratopyge fenghwangensis Hsiang]. 1977 Shengia quadrata Hsiang; Zhou, Liu, Meng, and Sun (in part), p. 175, pl. 51, figs. 22-24; non fig. 21 [=Shengia trapezia Peng]. 1978 Shengia quadrata Hsiang; Yang, p. 80, pl. 8, fig. 12. 1983 Shengia mina Qian in Qiu et al., 1983, p. 134, pl. 43, fig. 4. 1983 Shengia convexa Qiu in Qiu et al., 1983, p. 133, 134, pl. 43, figs. 6-8. 1992 Shengia quadrata Hsiang; Peng (in part), p. 76, figs. 40A-E, L, non figs. 40M, N, O [=Shengia trapezia Peng]. 2001 b Shengia quadrata Hsiang; Peng, Babcock, and Lin, p. 106, pl. 16, figs. 12, 13. Holotype. Pygidium (Egorova et al., 1963, pl. 13, fig. 5; GMC IV2610), from the upper part of the Glyptagnostus reticulatus Zone, Huaqiao Formation, near Tingziguan, Fenghuang, western Hunan. New material. More than 100 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 138397, 138406) in collection PI372. Emended diagnosis. Shengia with smooth dorsal surface. Pygidium with axis ending in front of posterior margin; pleural field narrow, with shallow pleural furrows; defined by wide border furrow; pygidial border flat and broad, moderately upturned abaxially. Remarks. Pygidia of Shengia quadrata, the type species of Shengia, in the type series include two morphologic forms. One form has wide borders, a relatively short axis, and narrow (tr.) pleural fields (Egorova et al., 1963, pl. 13, figs. 5, 6, 11). The other form has narrow borders (Egorova et al., 1983, figs. 8-10) with a relatively long axis that reaches almost to the posterior margin and has wide pleural fields. All the pygidia have a smooth surface. According to Egorova et al. (1963, p. 8), these specimens are from two different intervals of the Huaqiao Formation at Tingziguan, Fenghuang, northwestern Hunan. The narrow-bordered pygidia are from the lower interval, called the "the topmost of the lower part of upper Cambrian" (outcrop numbers 417 and 418), whereas the wide-bordered pygidia are from the "the middle part of upper Cambrian" (outcrop number 424). The holotype pygidium is a wide-bordered pygidium. Shengia pygidia in the new collections from the Huaqiao Formation at Paibi and Wangcun, northwestern Hunan, include both wide- and narrow-bordered specimens. Based on their associated cranidia, they likely represent two taxa. The wide-bordered pygidia agree well with the holotype pygidium of S. quadrata in every respect and the associated cranidia confirm the assignment, whereas the cranidia associated with narrow-bordered pygidia show that these specimens belong to S. trapezia. Previously no pygidium was known for S. trapezia. The finding of a pygidium for S. trapezia indicates that some smooth-surfaced pygidia with narrow borders and long axes referred previously to S. quadrata do not belong to S. quadrata but belong to S. trapezia. •

118.

The occurrence of Shengia in western and northwestern Hunan show that S. quadrata is restricted to a relatively high interval of Glyptagnostus reticulatus Zone, whereas S. trapezia ranges throughout this zone.

Occurrence. The holotype is from the upper part of the Glyptagnostus reticulatus Zone, Huaqiao Formation, near Tingziguan, Fenghuang, northwestern Hunan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it is the eponymous species of the Shengia quadrata Zone (equivalent to the Glyptagnostus reticulatus Zone). Shengia trapezia Peng, 1992 Plate 49, figures 1-10; Plate 78, ?figure 10 ?1963 ?1965

Shengia quadrata Hsiang in Egorova et al. (in part), p. 57, pl. 13, figs. 7-10 only. Shengia quadrata Hsiang; Lu, Zhang, Zhu, Qian, and Xiang (in part), p. 275, pl. 47, fig. 12 only. ?1978 Shengia quadrata Hsiang; Zhou, Liu, Meng, and Sun (in part), p. 175, pl. 51, fig. 23 only. 1992 Shengia quadrata Hsiang; Peng (in part), p. 76, figs. 40M, N, O only. 1992 Shengia trapezia Peng, p. 76, 77, fig. 40F-J. non 200 l b Shengia trapezia Peng; Peng, Babcock, and Lin, p. 105, pl. 15, figs. 7, 8. [=Shengia wannanensis Qiu in Qiu et al., 1983]. Holotype. Cranidium (Peng, 1992, fig. 40F-J; NIGP 95180) from the Glyptagnostus reticulatusChuangia wulingensis Zone, Huaqiao Formation, at Wa'ergang, Taoyuan, northwestern Hunan. New material. More than 20 sclerites including an incomplete exoskeleton, cranidia, librigenae, and pygidia (illustrated specimens NIGP 138047-138416, 138684) in collections P360.15, P1322.4, P1335.4, Pl336, P1360.3, P[365.8, PJ370.7, and P[371.2. Remarks. New material from the Huaqiao Formation, northwestern Hunan, includes the first recorded thorax, librigena, and pygidia of Shengia trapezia. The cranidium resembles the type material from the Cili-Taoyuan area of northwestern Hunan. Key features that warrant placing the new material into this species include a tapered, anteriorly truncate glabella, a transverse anterior border furrow, a long blade-like posterolateral projection, and a smooth external surface. The librigena has a broad genal field that bears fine radiating ridges on the exfoliated surface, and a wide lateral border bearing terrace ridges. The lateral and posterior border furrows are confluent at the genal angle, and extend onto the genal spine. The librigena seems indistinguishable from that of S. quadrata (see Peng, 1992, fig. 40A; text-fig. 43B). Thoracic segments in S. trapezia have a relatively narrow inner part and a wide outer part that bears strongly sloping facets. It is similar to the thorax of Baojingia subquadrata (Yang, 1978) ( P1. 41, fig. 12 herein). The pygidium differs from that of S. quadrata in having a longer, more segmented axis, a wider pleural field, and narrower borders. A tiny projection is present on each anterolateral comer of small pygidia (P1.49, figs. 5, 7) indicating a close relationship between Shengia and Lisania. Specimens previously assigned to the species by Peng et al. (2001b) include a cranidium and a pygidium. Both specimens have fine granules, are here reassigned to Shengia wannanensis Qiu.

• 119•

Occurrence. The holotype is from the Glyptagnostus reticulatus-Chuangia wulingensis Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan, China. New material is from darkgray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Liostracina bella Zone through the upper part of the Chuangia subquadrangulata Zone (equivalent to the lower part of the Glyptagnostus stolidotus Zone through the Glyptagnostus reticulatus Zone). Shengia wannanensis Qiu in Qiu et al., 1983 Plate 50, figures 1-16 1983 Shengia wannanensis Qiu in Qiu et al., 1983, p. 134, pl. 37, figs. 15, 16. 2001c Shengia sp., Peng, Babcock, Lin, Chen and Zhu, p. 166, pl. 4, fig. 11; pl. 5, ?fig. 15. 2001b Shengia trapezia Peng; Peng, Babcock, and Lin, p. 105, pl. 15, figs. 7, 8.

Holotype. Cranidium (Qiu et al., 1983, pl. 37, fig. 15, HIT 0249) from the lower part of the Furongian, near Beigong, southern Anhui. New material. More than 50 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 138418-138428) in collections P378.25, P13-1.60, P134.3, P135.1, and P[322.4. Emended diagnosis. Shengia with fine, closely spaced but unevenly distributed granules on cephalon and pygidium. Pygidium slightly wider than long, with relatively narrow borders, axis long and wide, reaching almost to posterior margin; pleural field with ribs defined by broad, deep pleural furrows. Description. Glabella parallel-sided or tapered gently forward, sides being gently convex. Occipital furrow wide medially. Lateral glabellar furrows consist of 4 pairs of faint impressions or areas lacking granules. Anterior branch of facial suture diverging gently forward; posterior branch diverges strongly rearward, deflected rearward at about two-thirds of the way to posterior margin, enclosing subrectangular posterolateral projections. Pygidium length about two-thirds width. Axis wide and long, almost as wide as pleural field, with 5 tings and semicircular terminal piece, reaching almost to posterior margin. Pleural field with four ribs and deep and rather broad pleural furrows. Border narrow, gently convex, defined by shallow border furrows, beating 2-3 terrace lines. Surface with fine, closely spaced granules, distributed unevenly on external surface. Smooth or poorly granulated areas include preocular area, anterior border, inner part of palpebral and posterior area of fixigena, anterior part of librigena, lateral parts of axis, anterior and outer parts of pleural field. Remarks. Shengia wannanensis Qiu was considered to be a junior synonym of Shengia quadrata Hsiang by Peng (1992), but new material from the Huaqiao Formation of northwestern Hunan suggests that S. wannanensis should be recognized as valid. The granulate prosopon on the new material agrees well with the description of Qiu (in Qiu et al., 1983) for S. wannanensis. Such prosopon has not been observed on other shengiids and serves a diagnostic character for this species. Other features of the new cranidia and pygidia that are consistent with S. wannanensis include the gently curved anterior border furrow on the cranidium; a long axis, and relatively narrow borders • 120•

on the pygidium. Aside from the surface prosopon, this species differs from S. quadrata in having a pygidium with a longer axis and narrower borders, and it differs from S. trapezia in having a less tapered glabella and a proportionally narrower pygidium. This species has a morphology intermediate between Baojingia Yang (in Zhou et al., 1977), especially between those species previously assigned to Eoshengia Yang (in Zhou et al., 1977) and Shengia Hsiang (in Egorova et al., 1963). The cranidium with convex anterior border, evenly curved anterior border furrow, and relatively narrow fixigena are similar to Shengia, but the granulate prosopon and the pygidial morphology are similar to Baojingia.

Occurrence. The holotype is from the middle part of the upper Cambrian, near Beigong, southern Anhui, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs in association with trilobites indicative of the lower part of the Liostracina bella Zone through the lower part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus stolidotus Zone through the Glyptagnostus reticulatus Zone). Shengia sp. Plate 78, figure 11

Material. Pygidium (NIGP 138685) in collection PI322.4. Remarks. An incomplete, partly exfoliated pygidium is characterized by having narrow, slightly convex lateral and posterior borders, and a pleural field with shallow pleural furrows. It is most similar to that of Shengia wannanensis Qiu (in Qiu et al., 1983, pl. 37, fig. 16; P1.50, figs. 8, 10-16 herein), but differs in having more effaced pleural fields and narrower borders. The less furrowed pleural field differentiates it also from all other species referred to Shengia. Occurrence. The pygidium is from dark-gray limestone of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). Family LONCHOCEPHALIDAEHup6, 1953 Genus

AETHIA

Qian and Zhou, 1984

Aethia Qian and Zhou, 1984, p. 179, 180. Type species. Aethia rectangula Qian and Zhou, 1984 (p. 180, pl. 2, fig. 2) from the "Paotaishan" Formatiom, Ma'anshan, Kunshan, southern Jiangsu; by original designation. Remarks. This is a small trilobite characterized by a cylindrical glabella, lenticular anterior cranidial border, faint eye ridges, and wide trapezoidal fixigenae. It resembles Prodamesella, but Prodamesella differs in having a tapered glabella, shorter anterior border, and distinct, well-raised eye ridges.

• 121.

Aethia rectangula Qian and Zhou, 1984 Plate 56, figures 15, 16 1984 Aethiarectangula Qian and Zhou, p. 181, pl. 2, fig. 2.

Holotype. Cranidium (Qian and Zhou, 1984, pl. 2, fig. 2, NIGP 69161) from the "Paotaishan Formation" (middle Wulingian), Kunshan, southern Jiangsu, China. New material. One cranidum (NIGP 138477) in collection W210.5. Description. Cranidium trapeziform, length two-thirds width. Anterior border lenticular, length (sag.) nearly equal that of occipital ring, narrowing evenly abaxially; anterior border furrow clearly defined, arched rearward medially, deflected rearward at sides. Glabella cylindrical, constricted slightly at $2, truncate anteriorly, with two pairs of shallow, notched lateral furrows. Occipital furrow incised, transverse; occipital ring large and tumid, wider than L1, with a faint median node, occupying one-fourth of glabellar length, gently narrowing abaxially. Eye ridge long (tr.), nearly effaced, directed nearly transversely; palpebral lobe small, located anteriorly, opposite to midlength of anterior lobe of glabella. Fixigena large, gently convex, trapezoidal, with maximum width at level of posterior border furrow, narrowing forward, width at level of anterior border furrow about two-thirds of that of anterior glabellar lobe; posterior border rather wide, of uniform width, defined by deep posterior border furrow. Anterior branch of facial suture converging forward diagonally; posterior branch diverging rearward at about 60 ° , cutting lateral border obliquely with posterior half. Occurrence. The holotype is from the middle Cambrian "Paotaishan Formation", Kunshan, southern Jiangsu, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Linguagnostus reconditus Zone). Genus NEOGLAPHYRASPIS Yuan and Yin, 1998

Glaphyraspis (Neoglaphyraspis) Yuan and Yin, 1998, p. 141, 142. Type species. Glaphyraspis (Neoglaphyraspis) nitida Yuan and Yin, 1998 (p. 142, pl. 1, figs. 7-12) from the Huaqiao Formation, Jimachong, Yuping, eastern Guizhou; by original designation. Remarks. This form was originally referred to Glaphyraspis Resser, 1937 as a subgenus (Yuan and Yin, 1998). Here, Neoglaphyraspis is regarded as a separate genus that is closely related to Glaphyraspis. The type species of Glaphyraspis, Liostracus parvus Walcott, 1899 (p. 463, 464, pl. 45, fig. 6), from northwestern Wyoming, was based on fragmental cranidia, but Walcott's (1899) line drawing seems correct in showing subcentrally located palpebral lobes and thin and oblique eye ridges. North American Glaphyraspis and its junior synonym Raaschella Lochman, 1938a were reported by various authors (e.g., Lochman, 1938a; Lochman and Duncan, 1944; Show, 1956; •

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Lochman and Hu, 1960, 1962; Rasetti, 1961, 1965; Palmer, 1954, 1962a, b, 1965; Hu and Tan, 1971; Robison, 1988; Pratt, 1992). Palmer (1965, p. 50, 51) clarified the genetic concept and recognized three groups of Glaphyraspis species that appear successively in stratigraphy. Pratt (1992) considered two of Palmer's (1962) Glaphyraspis species groups, one represented by the type species, the other represented by G. ornata (Lochman) and G. occidentalis to be indistinguishable and suppressed two the latter species as junior synonyms of the type species. As reported by Yuan and Yin (1998, p. 141, 142), Neoglaphyraspis differs from Glaphyraspis in many important respects, among which the anteriorly located palpebral lobes; the wide, transverse, depressed preglabellar field; the presence of general spines on the librigena, and the short axis on the pygidium are significant. Neoglaphyraspis resembles Glaphyraspis in most glabellar characters including the glabellar shape, the deeply incised lateral furrows, and the median indentation in the front of glabella; however, its lateral glabellar furrows are quite different from those of Glaphyraspis. The lateral glabellar furrows are all straight and subparallel in Neoglaphyraspis, but are directed differently, with the S1 furrow commonly curving rearward distally, in Glaphyraspis. Neoglaphyraspis resembles Rimouskia Resser, 1938, but Rimouskia can be differentiated by the absence of a clear preglabellar field, and a rounded glabellar front. The close relationship between Chinese Neoglaphyraspis and North American Glaphyraspis is also apparent. In the cranidium, the transverse eye ridges and the rather anteriorly located palpebral lobes, which characterize Neoglaphyraspis are also present on small cranidia of Glaphyraspis parva (Pratt, 1992, pl. 26, fig. 16). In the pygidium, the posterior bands of the pleurae are extended onto the lateral margin in both genera, although distal nodes are absent from each of the posterior bands in Neoglaphyraspis.

Neoglaphyraspis nitida Yuan and Yin, 1998 Plate 51, figures 1-17 1998 Glaphyraspis(Neoglaphyraspis) nitida Yuan and Yin, p. 142, pl. 1, figs. 7-12. 1998 Glaphyraspis(Neoglaphyraspis) trapezoidalis Yuan and Yin, p. 143, pl. 1, fig. 13. 2001b Neoglaphyraspisnitida Yuan and Yin; Peng, Babcock, and Lin, p. 103, pl. 8, fig. 11.

Holotype. Cranidium (Yuan and Yin, 1998, pl. 1, fig. 8, NIGP 127891) from the Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou.

New material. More than 50 sclerites including cranidia, pygidia, librigena (illustrated specimens NIGP 138429-138440) in collections P276.2, P277, P298.4, W199.2, W210.5, W211.7, W219.7, W221.5, W225, and W254.1.

Remarks. Neoglaphyraspis trapezoidalis is based on a single cranidium that comes from a single collection (WY23F1) from which most specimens assigned to the type species of the genus were derived (Yuan and Yin, 1998, pl. 1, figs. 7-12). Morphologically, the differences that Yuan and Yin (1998) used to distinguish G. trapezoidalis from the type species fall well within the range of variation of N. nitida, suggesting that G. trapezoidalis should be suppressed as a junior synonym of

N. nitida. This species, described first from eastern Guizhou, is common in northwestern Hunan. Large collections show that the shape of the glabella, including the median indentation in the front, the curvature of the anterior border, and the width of preglabellar field, are variable. • 123.

Yuan and Yin (1998) stated that the pygidial axis of N. nitida bears five to six tings. This interpretation was probably based on their small pygidium (Yuan and Yin, 1998, pl. 1, fig. 11), which is apparently a transitory pygidium with two unreleased thoracic segments. New material shows that the pygidial axis consists of three tings in the holaspid stage; each of the axial tings bears a large median node. The pygidium also shows a tiny terminal piece and a bar-like articulating half ring. Ring furrows are weak or faint. Occurrence. The holotype is from the Formosagnostus formosus [=Hadragnostus modestus] Blackwelderia Zone, Huaqiao Formation, Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun section, Hunan, where it occurs in association with trilobites indicative of the Wanshania wanshanensis Zone to the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone through the lowermost part of the Glyptagnostus reticulatus Zone).

Genus PRODAMESELLAChang, 1959 Prodamesella Chang (Zhang), 1957, p. 20 (nomen nudum); 1959, p. 196; Kobayashi, 1960b, p. 352; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 397; Jell and Robison, 1978, p. 16, 17; Peng, 1987, p. 104; Zhang and Jell, 1987, p. 212; Ergaliev, 1980, p. 151; Lisogor, Rozov, and Rozova, 1988, p. 71; Guo, Zan, and Luo, 1996, p. 122; Peng, Babcock, and Lin, 2001a, p. 411-413. Danjiangella Yang in Yang et al., 1991, p. 133; 1993, p. 171. Prodamesella (Prodamesella) Chang; Yuan and Yin, 2000, p. 252, 253. Prodamesella (Metaprodamesella) Yuan and Yin, 2000, p. 255. Prodamesella (Neoprodamesella) Yuan and Yin, 2000, p. 257, 258. Type species. Prodamesella convexa Chang (1959, p. 196, 197, pl. l, fig. 13) from the Changhia Formation, Boshan, Shandong; by original designation. Other species. Prodamesella biserrata Jell in Jell and Robison (1978, p. 17, pl. 1, fig. 9; pl. 4, figs. 1-6), from the middle Cambrian of northwestern Queensland, Australia; P. punctata Ergaliev (1980, p. 152, pl. 6, fig. 8), from the Kormagnostus simplex Zone, Malyi Karatau, Kazakhstan; and Danjiangella tenuispinosa Yang in Yang et al. (1991, p. 133, pl. 11, figs. 11, ?12; also 1993, p. 171, 172, pl. 11, figs. 11, ?12; text-fig. 49, cranidia only, ?pygidium) from the Pseudophalacroma scanense [=Triplagnostus gibbus posterus]-Doo'agnostus incertus Zone, Xijiadian Formation, Xichuan, southwestern Henan. Emended diagnosis. Small trilobite with trapezoidal cranidium. Glabella subrectangular, parallel sided or tapering gently forward, truncate to obtusely rounded in front, with posterior 3 pairs of lateral furrows deep and short; anterior border flat or ridge-like, with or without plectrum; preglabellar field narrow (sag.) or absent. Palpebral lobe small or tubercle-like, located anteriorly, eye ridge moderately distinct. Anterior branch of facial suture short, converging forward; posterior branch long, enclosing wide (tr., exs.) posterior area of fixigena. Thorax with convex axis and deeply furrowed pleurae. Pygidium transversely lanceolate; with short, weakly segmented axis. Surface punctate or granulate. Remarks. Jell (in Jell and Robison, 1978) discussed the genetic concept by excluding two species that Zhang (1959) tentatively referred to Prodamesella and restricting the genus to Prodamesella • 124.

convexa, the type species, and P. biserrata Jell in Jell and Robison. Based on rich material of Prodamesella from northwestern Hunan, Peng et al. (2001a, p. 412) rediagnosed the genus, and suppressed P. subtriangularis Peng, 1987 and most species erected by Yuan and Yin (2000) as junior synonyms of P. punctata. They also added information about the thoracic and pygidial features of Prodamesella by describing an incomplete thoracopygon of P. punctata. Although the holotype of Prodamesella convexa is incomplete, rich material from Australia, Kazakhstan, and South China (Jell and Robison, 1978; Ergaliev, 1980; Peng, 1987; Lisogor et al., 1988; Yuan and Yin, 2000; Peng et al., 2001 a) provides a strong basis for understanding the genetic concept. Danjiangella Yang in Yang et al., 1991 from southwestern Henan is here regarded as a junior synonym of Prodamesella. This genus is monotypic. The holotype of its type species, D. tenuispinosa (Yang et al., 1981, pl. 11, fig. 11), which is an incomplete cranidium with no anterior area preserved, shows that the genus has no significant difference from Prodamesella. The paratype

pygidium seems to be incorrectly associated as it bears border spines. Prodamesella punctata Ergaliev, 1980

Plate 52, figures 1-15; Plate 53, figures 1-14 1980 1987 2000 2000 2000

Prodamesella punctata Ergaliev, p. 152, pl. 6, fig. 8. Prodamesella subtriangulata Peng, p. 105, pl. 8, fig. 13. Prodamesella (Prodamesella) cylindrica Yuan and Yin, p. 254, 255, pl. 1, figs. 4-7, 9, 13, non fig. 8. Prodamesella (Metaprodamesella) subtriangulata Peng; Yuan and Yin (in part), p. 255, 256, pl. 2, figs. 1-3, 5, 7-9, 11-16, non figs. 4, 6, 10. Prodamesella (Metaprodamesella) subtriangulata prisca Yuan and Yin, p. 256, pl. 1, figs.

10-12. Prodamesella (Metaprodamesella) granulosa Yuan and Yin (in part), p. 256, 257, pl. 1, figs. 14-18, non 19. 2000 Prodamesella (Metaprodamesella) sp., Yuan and Yin, pl. 1, fig. 20. 2000 Prodamesella (Neoprodamesella) spinosa Yuan and Yin (in part), p. 258, 259, pl. 2, figs. 19, 21, 23, 24; non 18, 20, 22. 2001a Prodamesella punctata Ergaliev; Peng, Babcock, and Lin, p. 413, 414, pl. 1, figs. 1-14. 2001 b Prodamesella punctata Ergaliev; Peng, Babcock, and Lin, p. 105, pl. 13, figs. 10-12.

2000

Holotype. Cranidium (Ergaliev, 1980, pl. 6, fig. 8, IGSK 1950/107) from the Kormagnostus simplex

Zone, Malyi Karatau, Kazakhstan. New material. More than 50 sclerites including cranidia, pygidia, and librigena (illustrated

specimens NIGP 132807-132819, 132821, 138441-138449) in collections P249, P277, P319.6, P337.5, W196.3, W211.7, and W228.3. Remarks. Material from northwestern Hunan, some of which was illustrated previously by Peng et al. (2001a), is identical in all respects with the type series from Kazakhstan. Key characters that

warrant assignment of the Hunan material to this species include a tapered glabella with concave sides and a truncate front; a narrow (tr., sag.), convex anterior border bearing a broad plectrum; small anteriorly located palpebral lobes; wide (tr., exs.), and moderately convex posterior areas of the fixigenae; diagonal, slightly convex posterior branches of the facial suture, and densely spaced punctae on the surface. • 125.

Material illustrated here includes an ontogenetic series showing that in the meraspid stage the glabella is cylindrical in shape, bears transglabellar S 1 to $3 furrows and a large frontal lobe that is bilobed and sinuous anteriorly. Also, the fixigena is relatively narrow, and the occipital ring is proportionally large. In meraspid stages, the species has a transverse, narrow (sag.) pygidium that is transversely lanceolate, with a short, weakly segmented axis, and narrow borders. P. punctata is almost identical to P. convexa Chang, the type species of Prodamesella, in glabellar features, but P. convexa can be differentiated by having a thin, wider (sag.), and upturned anterior border, and more posteriorly located palpebral lobes. P. biserrata is distiguished by having a glabella that is concave anteriorly, subcentrally located palpebral lobes, and relatively narrow (tr.) posterior areas of the fixigenae. Occurrence. The holotype is from the Kormagnostus simplex Zone, Malyi Karatau, Kazakhstan. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the upper part of the Lejopyge laevigata Zone to the lower part of the Linguagnostus reconditus Zone). Prodamesella sp. cf. P. biserrata Jell in Jell and Robison, 1978 Plate 54, figures 5-16 cf. 1978 Prodamesella biserrata Jell in Jell and Robison, p. 17, pl. 1, fig. 9 (left), pl. 4, figs. 1-6. cf. 2001 b Prodamesella biserrata Jell; Peng, Babcock, and Lin, p. 102, pl. 5, fig. 15. Holotype of Prodamesella biserrata. Cranidium (Jell and Robison, 1978, pl. 4, figs. 2a, b, UQF 69369), from the Peronopsis opimus Zone, near Thomtonia, northwestern Queensland, Australia. Material. More than 30 cranidia (illustrated specimens NIGP 138454-138463) in collections P200.7, P204, P207, P268.3, P319.6, W 146.2, and W 171.9. Remarks. Specimens of Prodamesella from northwestern Hunan closely resemble P. biserrata from northwestern Queensland, Australia, in most key respects. The glabella that is tapered to the $3 furrow, slightly expanded in the frontal lobe, and bears an anteromedial indentation that has a bilobed appearance. Other features include a thin, wide anterior cranidial border with an obscure plecturm, a raised circular area on the occipital ring, a tiny node surrounded by four pits on the occipital ring, and relatively narrow (tr.) fixigenae. The Hunan specimens differ from the Australian P. biserrata in having a palpebral lobe that is located more anteriorly, opposite the base of the frontal lobe of the glabella (L4) rather than opposite the L3 lobe as in P. biserrata; and an external surface that is granulate rather than punctate. However, punctate ornamentation is present on exfoliated surfaces of some specimens from Hunan (P1.54, fig. 7). Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the Pianaspis sinensis Zone to the lower part of the Liostracina bella Zone (equivalent to the Lejopyge laevigata Zone through the lower part of the Linguagnostus reconditus Zone).

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Prodamesella tumidula sp. nov. Plate 54, figures 1-4 2001b Prodamesellabiserrata Jell; Peng, Babcock, and Lin, p. 102, pl. 4, fig. 10. 2001e Prodamesellabiserrata Jell; Peng, Babcock, Lin, Chen, and Zhu, p. 159, fig. 10.15.

Etymology. From Latin, tumidulus, rather or fairly tumid, referring to the proportionally large, somewhat tumid glabella. Holotype. Cranidium (P1. 54, fig. 3, NIGP 138452), which is about 2.2 mm long, from collection 123.6. Other material. Three cranidia, including an immature cranidium (NIGP 138450, 138451, 138453), in collections P123.6 and W56.7; a fragmental cranidium from P108. Diagnosis. Prodamesella with glabella parallel-sided or with straight sides tapered slightly forward, somewhat tumid, and somewhat concave anteriorly; anterior border a narrow (sag., exs.) ridge, lacking plectrum; palpebral lobe at level of or anterior to $3. Description. Cranidium trapezoidal, length two-thirds width; anterior margin curved forward gently. Anterior border a narrow ridge lacking plectrum, defined posteriorly by narrow, deeply incised border furrow. Glabella subrectangular, relatively large, tapered forward gently, with straight sides and sinuous front; with three pairs of short, deep lateral furrows that are subequally spaced. Occipital furrow deep, transverse; occipital ring crescentic, slightly wider (tr.) than basal glabellar lobe, beating tiny, posteriorly located median node. Eye ridge weak. Palpebral lobe located anteriorly, opposite frontal lobe of glabella; palpebral area narrower (tr.) than frontal lobe. Posterior area of fixigena with maximum width equal to or somewhat narrower than basal glabellar width. Remarks. Two cranidia (P1. 54, figs. 2, 4) that were assigned previously to Prodamesella biserrata Jell (Peng et al., 2001b, e), are now reassigned to a new species based on additional material that shows consistent differences from P. biserrata. P. tumidula sp. nov. differs from P. biserrata in having a proportionally large glabella that is less tapered or even parallel-sided, a narrower anterior border that lacks a plectrum, and a surface that lacks punctate ornamentation. In P. biserrata, the occipital ring bears a raised circular area with a tiny occipital node that is surrounded by four pits. In the new species, the occipital ring lacks the raised circular area and the pits. An obscure median node is present in P. tumidula. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Dorypyge richthofeni Zone to the lower part of the Pianaspis sinensis Zone (equivalent to the Ptychagnostus punctuosus Zone through the Goniagnostus nathorsti Zone).

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Family MAPANIIDAEChang, 1963 °o

Remarks. Opik (1967, p. 296, 297) rediagnosed the family Mapaniidae and added two genera, Quitacetra Opik, 1967 and Quitalai Opik, 1967 but Zhang and Jell (1987) used a restricted concept of the family and excluded Opik's (1967) genera from Australia from the family. Zhang and Jell (1987) retained Mapanopsis Chang 1963, along with Mapania Resser and Endo, 1937, in the Mapaniidae. Mapanopsis is characterized by having a subquadrate outline, diverging anterior branches of the facial suture, and an incomplete plectrum; it is quite similar to the Australian genera that Opik (1967) classified in the Mapaniidae. Maotunia Zhang and Jell, 1987 is also similar to Mapanopsis, although it was classified tentatively in the Proasaphiscidae by Zhang and Jell, 1987. Zhang and Jell (1987) suggested that Maotunia is ancestral to the mapaniids. It is clear that Maotunia, Mapania, Mapanopsis, Quitalai, and Quitacetra constitute a group of phylogenetically related taxa. Here, all of these genera are included in the Mapaniidae. Yuan and Yin (1998) erected Pseudomapania and placed it in the Mapaniidae. The genus is small in size, and characterized by small, subcentrally located palpebral lobes that are close to the axial furrow; a glabella with short, inwardly impressed lateral furrows; and narrow fixigenae. All these features suggest that its ancestry is not with the mapaniid lineage, and therefore it should be excluded from the Mapaniidae. The family Plectriferidae Opik (1967, p. 273) may be synonymous with the Mapaniidae. As diagnosed, the presence of a pair of pygidial spines seems to be the only character differentiating it from the Mapaniidae. The presence or absence of pygidial spines alone is not necessarily a firm family-level characteristic. Genus

MAPANIA

Resser and Endo in Kobayashi, 1935

Mapania Resser and Endo in Kobayashi, 1935, p. 228, 229; Resser and Endo, 1937, p. 250, 251; Lu, 1957, p. 266; Howell in Moore, 1959, p. 0288; Chernysheva, 1960, p. 93; Opik, 1961, p.164-166; 1967, p. 296, 297; Lu, Qian, and Zhu, 1963, p. 101; Lu, Zhang, Zhu, Qian, and Xiang, 1965, p. 339, 340; Nan, 1980, p. 500; Zhang and Jell, 1987, p. 188; Guo, Zan, and Luo, 1996, p. 89.

Type species. Mapania striata Resser and Endo in Kobayashi, 1935 (p. 229) [=Ptychoparia typus (Dames); see Walcott, 1913, p. 134, pl. 12, figs. 14, 14a-c] from the Amphoton Zone, Changhia Formation, Changxingdao Island, eastern Liaoning; by original disgnation.

Other species. Mapania beihoensis Kobayashi, 1935 (p. 229, pl. 20, figs. 8-10) from the Mapania Zone, Doten, South Korea; Mapania synopsis Opik, 1961 (p. 168-170, pl. 13, figs. 1-3; text-fig. 56) from the Proampyx agra Zone, middle part of the Devoncourt Limestone, Northern Territory, Australia; and Mapania jiangnanensis sp. nov. Mapania? dicella t3pik, 1961 (p. 171, 172, pl. 13, figs. 5, 7; text-fig. 56), and Mapania sp. cf. M. faceta Opik (1961, p. 170, pl. 13, fig. 6; text-fig. 56) are referred questionably to Mapania. For M. dicella, the genetic assignment is uncertain because the lateral glabellar furrows are absent. A cranidium is not known for M. sp. cf. M. faceta, and this fact renders its genetic assignment uncertain. Mapania? liaoningensis Resser and Endo (1937, p. 252, 253, pl. 43, figs. 23, 24) and Mapania angusta (Whitehouse) (1939, p. 224, pl. 223, fig. 21; also Opik, 1961, pl. 14, figs. 1-4, text-fig. 55) were transferred to Maotunia by Zhang and Jell (1987, p. 156, 157); Mapania faceta ()pik (1961, p. 170, pl. 13, fig. 4; text-fig. 56) also may belong to Maotunia. •

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Remarks. Zhang and Jell (1987) refigured Walcott's (1913) material of Ptychoparia typus (Dames), which was renamed M. striata by Resser and Endo (in Kobayashi, 1935), the type species of Mapania. Zhang and Jell (1987) selected a syntypic cranidium (Walcott, 1913, pl. 12, fig. 14a) as the lectotype of M. striata. The lectotype is flattened and somewhat deformed, but well preserved specimens, either in limestone or in shale, illustrated by Zhang and Jell (1987) demonstrate the cephalic features of the type species, and provide a basis for understanding the genetic concept. Zhang and Jell's (1987) additional material shows that Opik's (1961, text-fig. 53) reconstruction of the type species is not accurate in the shape of the glabella (gently tapered forward and rounded or obtusely rounded anteriorly rather than conical and truncate anteriorly), the course of the anterior branch of the facial suture (converging rather than diverging), and the number of lateral glabellar furrows (four pairs rather than three pairs, omitting the $3 furrows). Mapania jiangnanensis sp. nov. Plate 10, figures 5-10 2000b Mapaniasp., Peng, Babcock, and Lin, p. 101, pl. 1, fig. 16.

Etymology. From Chinese, jiangnan, south of the Yangtze River, South China, referring to the first record of a mapaniid from South China. Holotype. Broken cranidium (P1. 10, figs. 7-9, NIGP 137992) from collection P42.5. Other specimens. A cranidium with its counterpart, and a librigena in collection P42.5, and a cranidium possibly belonging to the species in collection W146.2 (NIGP 137990-137993). Diagnosis. Maotunia with a subrectangular glabella having incised lateral glabellar furrows, and an anterior cranidial border that bows rearward. Description. Cranidium subquadrate, length equal to width between palpebral lobes. Anterior border convex, bowed rearward, with thick plectrum that extends onto anterior lobe of glabella and separates frontal field of fixigenae into pair of subtriangular, wide and flat anterior fields. Glabella subrectangular, defined laterally by straight axial furrows, tapering forward slightly, rounded anteriorly; with four pairs of distinct lateral glabellar furrows; S1 bifurcate with long, curved posterior branch and shallow, transverse anterior branch; $2 long, arched anteriorly, lying near midpoint of cranidium; $3 short, isolated from axial furrow; $4 weak, isolated from but close to axial furrow, lying just behind inner end of eye ridge, directed forwardly and inwardly; occipital furrow transverse, shallow medially, deep laterally; occipital ring widest sagittally, narrowing abaxially, bearing tiny, weak, medial node. Palpebral area gently convex, width half glabellar width. Palpebral lobe rather large, crescentic, rather oblique, extending between midpoint of L1 and $3; palpebral furrow shallow. Eye ridge rather wide, directed diagonally. Anterior branch of facial suture diverging forward initially about 45 ° to sagittal line, becoming subparallel anteriorly; posterior branch nearly transverse, enclosing blade-like posterolateral projection; posterior border furrow shallow, posterior border wide. Librigena with moderately wide genal field and genal spine that is broad at base. Lateral and posterior borders moderately wide, somewhat elevated above outer part of genal field; bearing shallow furrow formed by concavity of border and parallel to lateral and posterior margins. Lateral • 129-

and posterior border furrows moderately deep, confluent at genal angle and extending into the base of genal spine as a broad, shallow furrow. Surface smooth except for frontal field of fixigena and genal field of librigena, which bear dense, radiating, anastomosing ridges.

Remarks. Mapania jiangnanensis sp. nov. seems to be an intermediate form between Mapania and Maotunia or between Mapania and Mapanopsis. The distinct lateral glabellar furrows and complete plectrum are characters typical of Mapania, and the slightly tapered glabella and strongly divergent anterior branches of the facial suture are typical of Maotunia and Mapanopsis. The new species is provisionally referred to Mapania because it seems most similar to the genetic concept of that genus. In the Paibi section, the new species appears higher in the section than does Maotunia rectangularis sp. nov. and Maotunia sp. cf. M. blackwelderi. The species occurs in association with Amphoton dios (in the lower Ptychagnostus atavus Zone). The intermediate features and the stratigraphic occurrence suggest that, phylogenetically, it represents the midpoint between

Maotunia and its descendent Mapania striata. One exfoliated cranidium referred to Mapania? liaodongensis (Resser and Endo, 1937, pl. 34, fig. 25, above; Zhang and Jell, 1987, pl. 84, fig. 6) has rather distinct lateral glabellar furrows and a glabella with a more rounded front. It is similar to the new species in having distinctive anastomosing ridges on the anterior field of the cranidium, and in having well defined eye ridges. However, it differs from the new species in lacking a plectrum and in having a more tapered glabella with less incised lateral glabellar furrows. M. ? liaodongensis was transferred to Maotunia by Zhang and Jell (1987). Zhang and Jell (1987, p. 156) considered the cranidium described as Mapaniidae gen. et sp. nov. (Opik, 1967, pl. 23, fig. 2) from the middle-upper Cambrian Passage Beds of Queensland, Australia, to belong to Maotunia. The cranidium is rather poorly preserved, but the overall features resemble those of the new species except for the proportionally longer glabella with less distinct lateral furrows, and the transverse rather than rearward-bowed anterior border.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the Dorypyge richthofeni Zone to the Pianaspis sinensis Zone (equivalent to the P~chagnostus atavus Zone through the Lejopyge laevigata Zone). Family ONCHONOTOPSIDAE Shaw, 1966

Remarks. The concept of Onchonotopsidae was rediagnosed and discussed at length by Robison (1988, p. 82). That concept is expanded slightly to include the Chinese genus Luyanhouaspis Peng, Babcock, and Lin, which bears a relatively wide, flat or slightly concave anterior border, and an effaced occipital furrow. Genus LUYANHAOASPISPeng, Babcock, and Lin, 2001 b

Luaspis Peng, Lin, and Chen, 1995, p. 286 (preoccupied). non Luaspis Hup6, 1953, p. 158. Luyanhaoaspis Peng, Babcock, and Lin, 2001, p. 100. Type species. Luaspis decorosa Peng, Lin, and Chen, 1995 (p. 286, pl. 6, figs. 1-9), the from •

130-

Huaqiao Formation, Paibi, Huayuan, northwestern Hunan; by original designation.

Other species. Chatiania (Eochatiania) laeviugata Yuan and Yin (1998, p. 152, pl. 3, figs. 5, 6), Luyanhaoaspis inflata sp. nov. and Luyanhaoaspis sp. cf. L. inflata sp. nov. Remarks. Peng et al. (2001b) proposed Luyanhaoaspis as a replacement name for Luaspis Peng et al. (1995) because Luaspis is preoccupied by an ellipisocephaloid trilobite (Hupr, 1953). The original genetic concept of Peng et al. (1995) is followed here. Peng et al. (1995) discussed the similarity of Luyanhaoaspis to Laurentian and Australian genera belonging to the Blountiinae but left open the familial classification of the genus. Luyanhaoaspis differs from blountioids in lacking a preglabellar field. A new species, Luyanhaoaspis inflata, bears some similarity to such genera as Matania Rasetti, 1946, Onchonotopus Rasetti, 1946, and Cryptoderaspis Rasetti, 1946, all from western Quebec, Canada. All these genera are placed within the family Onchonotopsidae on the basis of having a highly inflated glabella. Some features observed in immature individuals of Luyanhaoaspis are reminiscent of onchonotopsoids. A small cranidium of L. inflata (P1. 56, figs. 1, 2), which is probably in an early holaspid stage, bears an occipital furrow that is as clearly defined as in onchonotopsoids. A meraspid or early holaspid cranidium of L. decorosa (Peng et al., 1995, pl. 6, fig. 5) shows a convex, almost evenly wide, anterior border that is compatible with the morphology of onchonotopsoids. On the strength of these characters, Luyanhaoaspis is now classified tentatively within the Onchonotopsidae. Luyanhaoaspis decorosa (Peng, Lin, and Chen, 1995) Plate 55, figures 1-15 1995 Luaspis decorosa Peng, Lin, and Chen, p. 298, 299, pl. 6, figs. 1-9. 2001b Luayanhaoaspis decorosa (Peng, Lin, and Chen); Peng, Babcock, and Lin, p. 103, pl. 8, figs. 9,10.

Holotype. Cranidium (Peng, Lin, and Chen, 1995, pl. 6, figs. 2a-c, NIGP 118867; P1.55, figs. 1-3 herein), from the Lejopyge laevigata Zone, Huaqiao Formation, Paibi, Huayuan, northwestern Hunan. New material. About 20 sclerites including cranidia and pygidia (illustrated specimens NIGP 118868, 118872, 138464-138472) in collections P273.8, P277, P282.75, P290.5, P298.4, P298.54, P301.9, P319.6, P325.7, P331, P346.7, and P363.5. Remarks. New material from the Huaqiao Formation, northwestern Hunan, is closely similar to the type material from the same locality. This species has a cranidium with a bulbous glabella, a wide anterior border that is horizontal to slightly downsloping forward, small palpebral lobes that are very close to the axial furrows, and a small median node on the occipital ring. The pygidium is mostly effaced. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where occurs in association with trilobites indicative of the Wanshania wanshanensis Zone to the upper part of the Liostracina bella Zone (equivalent to the Proagnostus bulbus Zone through the lower • 131.

part of the Glyptagnostus stolidotus Zone).

Luyanhaoaspis inflata sp. nov. Plate 56, figures 1-9 2001b Luyanhaoaspissp. nov. Peng, Babcock, and Lin, p. 105, pl. 13, figs. 15, 16.

Etymology. From Latin, inflatus, inflated, referring to the highly inflated glabella. Holotype. Broken cranidium (P1.56, figs. 1, 4, 5, NIGP 138473) from collection P326.9. Other material. One small, testaceous cranidium and one fragmental cranidium (NIGP 138474, 138475) in collection P290.5, P326.5, and P341.8.

Diagnosis. Luyanhaoaspis with glabella that is highly inflated, subrectangular to elliptical; anterior border slightly concave; anterior border furrow obscure.

Description. Cranidium smooth, longer than wide. Anterior border incompletely known, moderately wide (sag.), slightly concave. Glabella subrectangular to elliptical, acutely rounded anteriorly, width two-thirds length, lacking lateral furrows; occipital furrow nearly effaced. Palpebral lobe close to axial furrow, length about one-fourth that of glabella. Posterior area of fixigena downsloping strongly, beating narrow border that is widened slightly abaxially and shallow border furrow. Anterior branch of facial suture diverges gently forward; posterior branch diverges strongly rearward for short distance, then curves inward gradually to cut posterior border exsagittally.

Remarks. Luyanhaoaspis inflata sp. nov. is differentiated from Luyanhaoaspis decorosa, the type species of the genus, by the shape and convexity of the glabella, and by the shape of the anterior border. A cranidium figured as Onchonotopsididae gen. et sp. indet. (Zhou et al., 1996, pl. 5, fig. 15) from Nidanshan, Hualong, northeastern Qinghai, China, which is now regarded as a new species belonging to Huayuanella gen. nov., resembles the small cranidium of L. inflata illustrated here. It differs in having a relatively narrower, more clearly defined anterior border furrow, and an occipital node that is located subcentrally. The glabellas of that specimen are somewhat more rounded than is the glabella of the small L. inflata cranidium.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Luyanhaoaspis sp. cf. L. inflata sp. nov. Plate 64, figures 11-15

Material. One cranidia (NIGP 138558) in collection P298.54. • 132.

Description. Cranidium subtriangular, width slightly greater than length. Anterior border narrow, gently convex, curved forward gently; anterior border furrow narrow but clearly defined. Preglabellar field narrow. Glabella subelliptical, width five-sixths length, rounded anteriorly and posteriorly, convex, defined by narrow, shallow axial and preglabellar furrows. Occipital furrow faint. Palpebral lobe small, close to axial lobe, located immediately anterior of cranidial midlength. Posterior area of fixigena triangular, sloping downward strongly, with maximm width at posterior margin, maximam width equal to one-third of basal glabellar width; with narrow border and shallow border furrow.

Remarks. One cranidia left in open nomenclature most closely resembles Luyanhaoaspis inflata sp. nov., described above, but differs in having a less convex, subquadrate rather than subovate glabella, and a much narrower, flat rather than upturned (sag. exs.) anterior border. Luyanhaoaspis decorosa, the type species, is differentiated by having a wide (sag., exs.) anterior border, a proportionally longer glabella, and more posteriorly located palpebral lobes. This specimen may represents an unnamed species of Luyanhaoaspis, but not enough material is known at present to ascertain that possibility.

Occurrence. From dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Linguagnostus reconditus Zone). Genus HUAYUANELLAgen. nov.

Etymology. From Huayuan County, Hunan, combined with the Latin suffix, -ella, small. Type species. Huayuanella paibiensis gen. et sp. nov. Other species. Peishania parallela Endo, 1944 (p. 66, 67, pl. 6, figs. 12-17) from the Changhia Formation, near Shuanmiaozi (Sobyosi), Liaoyang, Liaoning; Huayuanella zhiqiangi gen. et sp. nov.

Diagnosis. Onchonotopsidae with narrow preglabellar field. Glabella subquadrate, moderately convex; lateral furrows effaced; occipital ring nearly uniform in width (sag., exs.); anterior cranidial border strongly convex, arched moderately forward; palpebral lobes small, located subcentrally; anterior branch of facial suture diverging forward; posterior branch deflected rearward distally, subparallel to sagittal line, enclosing wide (sag.), short (exs.), subrectangular posterior area of fixigena.

Remarks. The new genus from northwestern Hunan most closely resembles Matania Rasetti in such features as the shape of the anterior border; the convexity of the glabella; the size, shape, and position of the palpebral lobes; and the width of the palpebral area. A subquadrate glabella is present in Matania (Robison, 1988). However, Matania is distinguished by the absence of a preglabellar field, the course of the facial suture (which is subparallel anterior to the palpebral lobes and directed diagonially behind the palpebral lobe), and the distally narrowing occipital ring.

• 133"

Huayuanella paibiensis gen. et sp. nov. Plate 56, figures 1O- 13

Etymology. From Paibi Village, Hunan. Holotype. Testaceous cranidium (Plate 56, figs. 10-13, NIGP 138476) from collection P331.8. Diagnosis. As for the genus. Description. Cranidium subquadrate, slightly longer than wide. Glabella subquadrate, moderately convex, anteriorly obtusely rounded, length (excluding occipital ring ) about 1.2 glabellar width; lateral glabellar furrows effaced; occipital ring nearly uniform in width (sag., exs.), slightly wider (tr.), than glabella at base, with tiny, faint, median node; occipital furrow shallow, transverse. Anterior border strongly convex, markedly arched forward, wider (sag.) than preglabellar field, occupying about 0.1 of total cranidial length; preglabellar field narrow. Palpebral lobe small, crescentic, close to axial furrow; palpebral furrow faint; palpebral area about one-fourth as wide as glabella; eye ridge faint, directed diagonally. Anterior branch of facial suture diverging forward, about 40 ° to sagittal line; posterior branch diverging rearward strongly, then deflecting rearward sharply to meet posterior margin subvertically. Posterior area of fixigena about half as wide as occipital ring, beating shallow and wide posterior border furrow. Posterior border wide (exs.), slightly convex. Remarks. A cranidium and a pygidium figured by Lu et al. (1965, pl. 62, figs. 9, 10) as H. parallela (Endo, 1944) are probably from the syntypic series of Endo (1944, pl. 6, fig. 13, and 16 respectively). The figured specimens of Lu et al. (1965), although illustrated poorly, provide a basis for understanding the species concept, and that species resembles Huayuanella paibiensis sp. nov. in having a proportionally shorter glabella. A lectotype was not designated previously for H. parallela, so a cranidium figured by Endo (1942, pl. 6, fig. 13) is here chosen as the lectotype of the species. H. paibiensis differs from H. parallela in having a proportionally shorter glabella with a less rounded front, and an occipital ring that is less narrowed at the sides. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the Linguagnostus reconditus Zone). Huayuanella zhiqiangi gen. et sp. nov. Plate 56, figure 14 1996 Onchonotopsididae gen. et sp. nov.; Zhou, Cao, Hu, and Zhao, p. 45, pl. 5, fig. 15.

Etymology. Named after Zhou Zhiqiang, who collected and illustrated the holotype of the species. Holotype. By monotypy; A cranidium illustrated originally by Zhou et al. (1996, pl. 5, fig. 15) and • 134.

herein (P1. 56, fig. 14, NIGP 132820), from the Nidanshan Group, Nidanshan, Hualong, Qinghai, Northwest China.

Remarks. A cranidium illustrated by Zhou et al. (1996) as a representative of a new species of an undescribed genus within the Onchonotopsidae is here described as Huayuanella zhiqiangi. It differs from H. paibiensis, the type species of Huayuanella, in having greater convexity, a glabella that is subelliptical in outline and is more rounded anteriorly, an anterior border that is flatter and more clearly defined posteriorly, and in having an occipital furrow that is more extensively bowed to the rear. Occurrence. The holotype is from the uppermost part of the Nidanshan Group (probably Wangcunian-equivalent), Nidanshan, Hualong, Qinghai, Northwest China. Family PAPYRIASPIDIDAEWhitehouse, 1939 Genus PIANASPIS Saito and Sakakura, 1936

Pianaspis Saito and Sakakura, 1936, p. 114-116; Chernysheva, 1970, p. 123, 124; Jago, 1974, p. 141, 142; Lu and Zhu, 1980, p. 15; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 101; Xiang and Zhang, 1985, p. 118; Lu and Lin, 1989, p. 132, 133; Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, 1991, p. 142; 1993, p. 184; Yuan, Zhao, Li, and Huang, 2002, p. 207, 208. Prohedinia Lermontova and Chernysheva in Chernysheva, 1950, p. 68, 69; Egorova, Lermontova, Poletaeva, and Sivov, 1955, p. 127; Howle in Moore, 1959, p. 0269; Balashova, Ivshin, and Chernysheva in Chernysheva, 1960, p. 109; Egorova, Ivshin, Pokrovskaya, Poletaeva, Repina, Rozova, Romanenko, Sivov, Tomashpoliskaya, Chernysheva, 1960, p. 230; Kobayashi, 1962, p. 57; Egorova, Xiang, Li, Nan, and Guo, 1963, p. 35, 36; Lu, Zhang, Zhi, Qian, and Xiang, 1965, p. 189; Soloviev, 1966, p.15, 16; Palmer, 1968, p. 69; Zhou, Liu, Meng, and Sun, 1977, p. 153; Yin and Li, 1978; p. 480, 481; Zhang, 1981, p. 167, 168; Zhang in Zhang, Xiang, Liu, and Meng, 1995, p. 27, 28. Pinaspis Saito and Sakakura; Kim, 1987, p. 35 (misspelled generic name). Tosotychia Opik, 1961, p. 160. Pianaspis (Pianaspis) Saito and Sakakura; Yuan, Zhao, Li, and Huang, 2002, p. 208. Pianaspis (Prohedinia) Lermontova and Chernysheva; Yuan, Zhao, Li, and Huang, 2002, p. 208. Type species. Pianaspis kodairai Saito and Sakakura, 1936 (p. 114, pl. 8, figs. 1-3) from the Ssukkhol Formation, south of Pyongyang, North Korea; by original designation. Remarks. This genus has been discussed by a number authors (e.g., Chernysheva, 1970; Lu and Zhu, 1980; Jago, 1974; Lu and Lin, 1989; Zhang in Zhang et al., 1995). We follow Chernysheva (1970, p. 122-124) in regarding Prohedinia and Toso~chia as junior synonyms of Pianaspis. Suppression of Tosotychia as a junior synonym of Prohedinia has been accepted by most authors, but the placement of Prohedinia in synonymy with Pianaspis has not received universal acceptance. Jago (1974) and Lu and Lin (1989) accepted the synonymy with reservations, whereas Zhang in Zhang et al. (1995) rejected it. According to Zhang (in Zhang et al., 1995, p. 27), Prohedinia is not synonymous with Pianaspis, but a valid genus that differs from Prohedinia in having a broader cranidium, a very narrow anterior border, a prominent median plectrum, a depressed preglabellar field, a long eye 135. •

ridge, and more arcuate palpebral lobes. In addition, Zhang (in Zhang et al., 1995, p.27) noted that the palpebral lobe of Pianaspis seems smaller than that of Prohedinia. Comparison of Pianaspis hodairai (Saito and Sakakura, 1936), the type species of Pianaspis, with Prohedinia attenuata, the type species of Prohedinia (Chernysheva, 1950), and the variation expressed in Siberian (Soloviev, 1966) and Chinese material of Prohedinia tends to support inclusion of Prohedinia as a junior synonym of Pianaspis. In Prohedinia attenuata, the anterior border is not narrower but wider than that of Pianaspis kodairai; the plectrum seems to be absent; and the eye ridge, which is variable in P. attenuata, seems indistinguishable in length from that of P. kodairai. New material of Pianaspis sinensis from Hunan includes an ontogenetic series. Compared to Pianaspis kodairai, some specimens assigned here to P. sinensis have narrower rather than broader cranidial proportions (P1. 57, figs. 11, 12; P1. 58, figs. 1, 4; P1. 61, figs. 12-15). The specimens show that the anterior border varies in length through ontogeny. It is narrower in earlier stages and becomes wider in later stages. The observed anterior border width of P. kodairai falls within the range of variation of the anterior border width of P. sinensis. The plectrum is extremely variable in Prohedinia. It may be absent, as in Prohedinia siberica Soloviev (1966, pl. 1, figs. 1-4; pl. 2, figs. 1-6) and P. attenuata Lermontova and Chernysheva sensu Soloviev (1966, pl. 2, figs. 9-11), or it may be present, as in our material of P. sinensis. In the latter case, it varies from small and weak to large and prominent. In addition, some specimens assigned to P. attenuata from Uzberkistan (Khairullina, 1973), Siberia (Egorova et al., 1982), and Kazakhstan (Lisogor et al., 1988) clearly lack a plectrum. We do not see much difference in the nature of the preglabellar field and the length of eye ridges between Prohedinia and Pianaspis, and some specimens in our collection (P1.57, fig. 9; P1. 61, figs. 12, 13) even seem to have shorter eye ridges than P. kodairai. Most species of Prohedinia have a palpebral lobe that is longer than that of Pianaspis kodairai, but P. sors (Opik, 1961, pl. 15, figs. 1-7; text-fig. 50) seems indistinguishable from P. kodairai in the size of the palpebral lobe. Thus, the more arcuate palpebral lobe becomes the only character that serves to distinguish it from Pianaspis. This character, however, should be only of specific significance. Zhang (in Zhang et al., 1995, p. 27, 28) listed the species and subgenera of Prohedinia that were known to him. He excluded Prohedinia brevifrons Palmer (1968) and Prohedinia (?) sp. sensu Xiang and Zhang (1985, p. 115, pl. 36, fig. 8) from the genus, and referred Prohedinia (?) sp. sensu Zhou et al. (1977, p. 154, pl. 47, fig. 7) to Prohedinia attenuata. We agree with these assignments. Zhang (in Zhang et al., 1995) also synonymized all other species with P. attenuata. However, among these species, at least Tosotychia sors Opik, 1961 and Prohedinia sinensis Yang in Zhou et al., 1977 should be considered species of Pianaspis. Prohedinia sp. 1 sensu Yang, 1978 (p. 39, pl. 6, fig. 7) belongs to Wangcunia wangcunensis. Zhang (in Zhang et al., 1995) provided a list of Pianaspis species. To the list the following records of Pianaspis should be added: Prohedinia attenuata Lermontova and Chernysheva; Egorova, Pokrovvskaya, Poletaeva, Repina, Rozova, Romanenko, Sivov, Tomashpoliskaya, Fedyanninag and Chernysheva, 1960, p. 231, pl. Cm-26, fig. 9. Prohedinia sibirica Soloviev, 1966, p. 17-23, pl. 1, figs. 1-4; pl. 2, figs. 1-8; text-fig. 4a. Prohedinia attenuata Lermontova and Chernysheva; Soloviev, 1966, p. 23, 24, pl. 2, figs. 9-11; text-fig. 4b. Pianaspis attenuata (Lermontova and Chernysheva); Khairullina, 1973, p. 79, pl. 11, figs. 1-4, 6, 8, 11. Pianaspis recta Khairullina, 1973, p. 81, pl. 11, fig. 7. Pianaspis sp. 1, Khairullina, 1973, p. 112, pl. 11, figs. 5, 9. Pianaspis sp., Repina, Petrunina, and Khairullina, 1975, p. 168, 169, pl. 27, fig. 9. • 136.

Pianaspis attenuata (Lermontova and Chernysheva); Egorova et al., 1982, p. 109, pl. 20, figs. 2, 3;

pl. 23, figs. 6, 7; pl. 25, fig. 5; pl. 27, fig. 18; pl. 43, fig. 12; pl. 45, fig. 1; pl. 45, fig. 7. Pianaspis attenuata (Lermontova and Chernysheva); Lisogor, Rozov, and Rozova, 1988, p. 73, fig. 8. Pianaspis sinensis (Yang); Lu and Lin, 1989, p. 133, 134, pl. 20, figs. 1-5. Pianaspis attenuata (Lermontova and Chernysheva); Yang et al., 1991, p. 142, pl. 14, figs. 1-3a (reprinted 1993, p. 184). Pianaspis sinensis (Yang); Yang et al., 1991, p. 142, 143, pl. 14, figs. 4-8 (reprinted 1993, p. 184). Pianaspis dictyodroma Yang and Liu in Yang et al., 1991, p. 143, 185, pl. 14, figs. 9, 10 (reprinted 1993, p. 184). Pianaspis sp. 1, Yang and Liu in Yang et al., 1991, p. 143, 144, 186, pl. 14, fig. 11 (reprinted 1993, p. 185). Pianaspis sp. 2, Yang and Liu in Yang et al., 1991, p. 144, pl. 14, figs. 12, 13 (reprinted 1993, p. 186). Pianaspis sp. 3, Yang and Liu in Yang et al., 1991, p. 144, pl. 15, fig. 1 (reprinted 1993, p. 186). Prohedinia attenuata Lermontova and Chernysheva; Zhang in Zhang et al., 1995 (in part), pl. 4, fig. 9; pl. 10, figs. 2, 3, non 10. Here, Prohedinia sibirica is considered to be a valid species of Pianaspis, characterized by having a narrower (sag.) preglabellar field, an oblique eye ridge, and a thorax that contains as many as 29 segments. Pianaspis recta Khairullina is based on a single distorted cranidium. This species and Pianaspis sp. sensu Khairullina seem to be synonymous with P. attenuata. All specimens assigned to P. dictyodroma, and P. spp. 1 - 3 by Yang (in Yang et al., 1991; reprinted 1993) are also synonymous with Pianaspis attenuata. They are from the same locality and probably the same horizon as the material of P. attenuata from Xichuan, Henan, China. Pianaspis is a widespread genus, occurring in Korea, northern and southeastern Siberia, Sayan-Altai, Uzbekistan, Kazakhstan, Australia (Queensland, Tasmania), and China (Qinghai, Hunan, Guizhou, Xinjiang, Zhejiang, Henan). Except for P. kodairai, the type species, Pianaspis contains only four species: P. attenuata (Lermontova and Chernysheva, 1950), P. sors (Opik, 1961), P. sibirica (Soloviev, 1966), and P. sinensis (Yang in Zhou et al., 1977). So far as known, P. kodairai is the oldest species (probably in the Ptychagnostus gibbus Zone) and then P. sinensis, P. attenuata, P. sibirica, and P. sors probably appear in succession. The lowest occurrence of each of the last four species is reported repectively in the P. punctuosus Zone, the Anomocarioides limbataeformis Zone (=post-punctuosus and pre-armata Zone) (Egorova et al., 1982), the L. armata Zone, and the basal Lejopyge laevigata Zone. Prohedinia sp. sensu Yang, 1978 (p. 39, pl. 6, fig. 7) is probably a species of Pianaspis, and it occurs in association with Dorypyge richthofeni in the basal Ptychagnostus atavus Zone. Pianaspis attenuata (Lermontova and Chemysheva in Chernysheva, 1950)

Plate 58, figure 10 1950 Prohedinia attenuata Lermontova and Chernysheva in Chernysheva, p. 69-72, pl. 1, figs. 9-12. 1955 Prohedinia slida Poletaeva in Egorova et al., p. 127, 128, pl. 14, figs. 5a, b. 1960 Prohedinia attenuata Lermontova and Chernysheva; Egorova, Ivshin, Pokrovskaya, Poletaeva, Repina, Rozova, Romanenko, Sivov, Tomashpoliskaya, Chernysheva, p. 231, pl. Cm-24, fig. 9. non 1963 Prohedinia attenuata Lermontova and Chernysheva; Egorova, Xiang, Li, Nan, and Guo, p. 36, 37, pl. 6, figs. 13, 14 [=Pianaspis sinensis (Yang in Zhou et al., 1977)]. • 137.

non 1965

1965 1966

Prohedinia attenuata Lermontova and Chernysheva, Lu, Zhang, Zhu, Qian, and Xiang, p. 189, pl. 32, fig. 5 [=Pianaspis sinensis (Yang in Zhou et al., 1977)]. Prohedinia attenuata Lermontova and Chernysheva; Zhu, p. 141, pl. 2, fig. 8. Prohedinia attenuata Lermontova and Chernysheva; Soloviev, p. 23, 24, pl. 2, figs. 9-11;

1973

Pianaspis attenuata (Lermontova and Chernysheva); Khairullina, p. 79, pl. 11, figs. 1-4, 6, 8,

text-fig. 4b. 11. 1973 Pianaspis recta Khairullina, p. 81, pl. 11, fig. 7. 1973 Pianaspis sp. 1, Khairullina, p. 112, pl. 11, figs. 5, 9. 1974 Pianaspis (?) loveni Jago, p. 143-146, pl. 1, figs. 1-12, text-fig. 2. 1975 Pianaspis sp., Repina, Petmnina, and Khairullina, p. 168, 169, pl. 27, fig. 9. 1977 Pianaspis sp., Zhou, Liu, Meng, and Sun, p. 154, pl. 47, fig. 7. 1982 Pianaspis attenuata (Lermontova and Chernysheva); Egorova et al., p. 109, pl. 20, figs. 2, 3; pl. 23, figs. 6, 7; pl. 25, fig. 5; pl. 27, fig. 18; pl. 43, fig. 12; pl. 45, fig. 1; pl. 46, fig. 7. 1985 Prohedinia? sp.; Xiang and Zhang, p.115, pl. 36, fig. 8. 1988 Pianaspis attenuata (Lermontova and Chernysheva); Lisogor, Rozov, and Rozova, p. 73, fig. 8. 1991 Pianaspis attenuata (Lermontova and Chernysheva); Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p. 142, pl. 14, figs. 1-3a. 1991 Pianaspis sinensis (Yang); Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p. 142, 143, pl. 14, figs. 4, 5, 7, 8. 1991 Pianaspis dictyodrama Yang in Yang et al., p. 143, pl. 14, figs. 9, 10. ?1991 Pianaspis sp. 1, Yang and Liu in Yang et al., p. 143, 144, pl. 14, fig. 11. 1991 Pianaspis sp. 2, Yang and Liu in Yang et al., p. 143, 144, pl. 14, figs. 12, 13. 1991 Pianaspis sp. 3, Yang and Liu in Yang et al., p. 144, pl. 15, fig. 1. 1993 Pianaspis attenuata (Lermontova and Chernysheva); Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p. 184, pl. 14, figs. 1-3a. 1993 Pianaspis sinensis (Yang); Yang, Yu, Liu, Su, He, Sheng, Zhang, Zhu, Li, and Yan, p. 184, pl. 14, figs. 4, 5, 7, 8. 1993 Pianaspis dictyodrama Yang in Yang et al., p. 184, 185, pl. 14, figs. 9, 10. ?1993 Pianaspis sp. 1, Yang and Liu in Yang et al., p. 185, 186, pl. 14, fig. 11. 1993 Pianaspis sp. 2, Yang and Liu in Yang et al., p. 186, pl. 14, figs. 12, 13. 1993 Pianaspis sp. 3, Yang and Liu in Yang et al., p. 186, pl. 15, fig. 1. 1995 Prohedinia attenuata Lermontova and Chernysheva; Zhang in Zhang et al., 1995 (in part), pl. 4, fig. 9; pl. 10, figs. 2, 3, non 10. Holotype. Cranidium (Chernysheva, 1950, pl. 1, fig. 9; repository unknown) from the Paradoxides f o r c h h a m m e r i Zone (upper Middle Cambrian), eastern Siberia. N e w material. A single cranidium (NIGP 138501) in collection P 184 and two cranidial fragments in

collection P 168. Remarks. A cranidium referred to P. attenuata that has a straight anterior border, wide (tr.)

palpebral areas, nearly transverse eye ridges, a wide (sag.) preglabellar field, and outwardly bowed anterior branches of the facial suture. Morphologically, the specimen falls well within the range of variation of P. attenuata, and it is especially similar to the holotype and paratype cranidia (Chernysheva, 1950, pl. 1, figs. 9, 10) and some other specimens assigned to P. attenuata from Siberia (cf. Egorova et al., 1982, pl. 23, fig. 7; pl. 43, fig. 12). • 138.

Occurrence. The holotype is from the Paradoxides forchhammeri Zone (upper Middle Cambrian), eastern Siberia. New material is from dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the Pianaspis sinensis Zone (equivalent to the Lejopyge laevigata). Pianaspis sinensis (Yang in Zhou et al., 1977)

Plate 57, figures 1-15; Plate 58, figures 1-9; Plate 61, figures 11-15 1963

Prohedinia attenuata Lermontova and Chernysheva, Egorova, Xiang, Li, Nan, and Guo, p.

36, 37, pl. 6, figs. 13, 14. Gen. et sp. indet., Egorova, Xiang, Li, Nan, and Guo, p. 38, pl. 6, fig. 16. Prohedinia attenuata Lermontova and Chernysheva; Lu, Zhang, Zhu, Qian, and Xiang, p. 189, pl. 32, fig. 5. 1965 Inouyella sp., Lu, Zhang, Zhu, Qian, and Xiang, p. 231, pl. 40, fig. 10. 1977 Prohedinia sinensis Yang in Zhou et al., p. 154, pl. 47, fig. 6. 1978 Prohedima attenuata Lermontova and Chernysheva; Yin and Li, p. 481, pl. 163, fig. 4. 1978 Prohedinta sinensis Yang; Yin and Li, p. 481, pl. 163, fig. 14. 1978 Prohedinia sinensis Yang; Yang, p. 39, pl. 6, fig. 6. 1981 Prohedima sinensis quadrata Zhang T., p. 168, pl. 62, figs. 1a, b. 1982 Prohedinta sinensis Yang; Liu, p. 310, pl. 217, figs. 5, 6. 1983 Prohedima sinensis quadrata Zhang T.; Zhang Q. in Qiu et al., p. 101, pl. 34, figs. 7, 8. 1989 Pianaspis sinensis (Yang); Lu and Lin, 1989, p. 133, 134, pl. 20, figs. 1-5. non 1991 Pianaspis sinensis (Yang); Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 142, 143, pl. 14, figs. 4, 5, 7, 8. non 1993 Pianaspis sinensis (Yang); Yang, Yu, Liu, Su, He, Shang, Zhang, Zhu, Li, and Yan, p. 184, pl. 14, figs. 4, 5, 7, 8. 2001b Pianaspis sinensis (Yang); Peng, Babcock, and Lin, p. 102, pl. 4, fig. 9. 1963 1965

Holotype. By monotype; incomplete cranidium (Yang in Zhou et al., 1977, pl. 47, fig. 6, CUGB 0508303), from the Paramphoton [=Amphoton] Zone, Huaqiao Formation, Fenghuang, north-

western Hunan. New material. More than 30 sclerites including cranidia, librigena, hypostomes, and pygidia

(illustrated specimens NIGP 138478-138550, 138527-138531) in collections P112.6, P122, P122.4, P123.6, P126, P130.95, P131.7, P134.2, P135, P135.7, P136.3, P136.7, P137.8, P138.8, W82.1, W98, W99.3, W101, W105, W113, and W121.6. Remarks. Zhang (in Zhang et al., 1995, p. 27, 28) placed P. sinensis in synonymy with Prohedinia attenuata (together with some other species assigned to either Prohedinia or Pianaspis). Here, however, P. sinensis is regarded as a valid species. The holotype of P. sinensis is an incomplete

cranidium showing a rather wide, flat, upturned anterior border, an oblique eye ridge, and a prominent, rather large plectrum. Excellent new material from the same area as the holotype cranidium of P. sinensis shows that characters exposed in the holotype are consistent and distinctive. Our material, and other additional material assigned previously to P. sinensis or to P. sinensis quadrata (Zhang, 1981; Qiu et al., 1983), demonstrate that P. sinensis is further differentiated from P. attenuata in having an arched rather than straight anterior border furrow, a relatively wide (sag.) preglabellar area, and a narrower (tr.) palpebral area. • 139-

The first reported ontogenetic series for Pianaspis (herein) shows some notable changes in morphology during ontogeny. The preglabellar field is absent in early meraspid stages and becomes wide (sag.) in succeeding stages. The palpebral area decreases in width progressively during ontogeny. The glabella becomes more tapered, the anterior border becomes more forward-arched, and the S 1 furrow becomes bifurcate in holaspid stages. The front of the glabella varies in shape from transverse to obtusely rounded, and then finally to rounded. Most specimens show an increase in size of the plectrum during ontogeny, but a few cranidia show a weak plectrum.

Occurrence. The holotype is from the Huaqiao Formation, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it is the eponymic species of the Pianaspis sinensis Zone (equivalent to the Ptychagnostus punctuosus Zone through the lower part of the Lejopyge laevigata Zone). Genus

WANGCUNIA

Peng, Lin, and Chen, 1995

Type species. Wangcunia wangcunensis Peng, Lin, and Chert, 1995 (p. 278, 279; 290, 291, pl. 1, figs. 1-11, 12b, 13) from the Huaqiao Formation, Wangcun, northwestern Hunan.

Remarks. This genus was tentatively referred to the family Papyriaspididae Whitehouse, 1939 when it was erected (Peng et al., 1995). Because it is almost identical with other papyriaspidids in cephalic morphology, and bears a mutisegmented thoracopygon, it is now referred with confidence to that family. The genetic concept of Peng et al. (1995) is followed here. Wangcunia wangcunensis Peng, Lin, and Chen, 1995 Plate 59, figures 1-14 1978 Prohedinia sp., Yang, p. 39, pl. 6, figs. 7a, b. 1995 Wangcunia wangcunensis Peng, Lin, and Chen, p. 290, 291, pl. l, figs. 1-1 l, 12b, 13. 2001b Wangcunia wangcunensis Peng, Lin, and Chen; Peng, Babcock, and Lin, p. 102, pl. 3, figs. 9, 10. 2001 e Wangcunia wangcunensis Peng, Lin, and Chen; Peng, Babcock, Lin, Chen, and Zhu, p. 159, figs. 10.16-10.18.

Holotype. Crushed incomplete cranidium (Peng et al., 1995, pl. 1, fig. 2, NIGP 118805, from the basal Ptychagnostus punctuosus Zone, Huaqiao Formation, Wangcun, northwestern Hunan. New material. Several cranidia and pygidia collected from the same bed and same locality as the type material, collection W56.7 (illustrated specimens including part of type material: NIGP 118804, 118806-118808, 118810-118813, 138817, 138502-138505). Remarks. The species concept of Peng et al. (1995) is followed here. Prohedinia sp. (Yang, 1978), based on a single cranidium from the Dorypyge sp. cf. D. richthofeni [=Dorypyge richthofeni] Zone at Fengmuping, Fenghuang, northwestern Hunan, is considered synonymous with W. wangcunensis. Both species are identical in cranidial features. They are also from the same paleogeographic region. The type and new material of W. wangcunensis occur in association with Dorypyge richthofeni in the Wangcun section. 140•

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus punctuosus Zone). Genus PARAPIANASPIS gen. nov.

Etymology. Latin prefix, para, next to, plus genetic name Pianaspis, referring to its close similarity to that genus. Diagnosis. Papyriaspididae with glabella having parallel sides, anterior obtusely rounded, with weak lateral furrows; anterior branch of facial suture diverging strongly; anterior border flat and wide (sag.); preglabellar field somewhat wider than anterior border; palpebral lobe large, located posteriorly, with posterior end slightly anterior to occipital furrow. Pygidium subrounded, with relatively large, multisegmented axis and wide borders.

Remarks. The new genus is similar to Pianaspis in having a nearly straight anterior border furrow interrupted by a plectrum, a glabella occupying about two-thirds of cranidial length, a long eye ridge, and an arcuate palpebral lobe. However, Parapianaspis gen. nov. is differentiated from Pianaspis by the subrectangular rather than tapered glabella; the more effaced lateral glabellar furrows; the strongly diverging rather than parallel-sided or converging anterior branches of the facial suture; the narrower preglabellar field; the larger, more posteriorly located palpebral lobes; the flat rather than convex anterior border; the evenly wide (sag., exs.) rather than laterally narrowed occipital ring; and the subrounded rather than transverse pygidium. Parapianaspis can be differentiated from Pianaspis throughout its post-protaspid ontogeny. Small cranidia of the new genus (P1. 61, figs. 1-6, 9, 10) have an anteriorly expanded glabella, large, posteriorly located palpebral lobes, a wide anterior border and strongly diverging anterior branches of the facial suture. However, in similar ontogenetic stages (P1.57, figs. 3-6), the glabella of Pianaspis is parallel-sided or tapered anteriorly, the palpebral lobe is medium in size, and is located subcentrally, the anterior border is narrow, ridge-like, and the facial suture is diverging slightly forward or parallel-sided. The rectangular glabella and the strongly diverging anterior branch of the facial suture are reminiscent of Hedinaspis Troedsson (1937, 1951) and Asiocephalus Palmer (1968). Both genera, however, differ in having smaller palpebral lobes that are located in advance of the glabellar midlength. Hedinaspis has small palpebral lobes throughout its post-meraspid ontogenetic stages (Taylor, 1976). Parapianaspis shows some similarity to Wangcunia Peng et al. (1995). Both genera are characterized by having a multisegmented pygidium, but Wangcunia differs in having a proportionally large glabella with an acutely rounded front, converging anterior branches of the facial suture, a smaller palpebral lobe that is located subcentrally, and a pygidium with a wider (tr.) pleural field and much narrower borders. Rhodonaspis Whitehouse (type species, R. longula Whitehouse, 1939, p. 220) from the Glyptagnostus stolidotus Zone of Queensland, Australia, was questionably assigned to the Papyriaspididae (Poulsen in Moore, 1959). It is similar to the new genus in cranidial features, especially the glabellar outline, the course of the facial suture, and the size and shape of palpebral lobes. It differs, however, in having transverse eye ridges, deeper lateral glabellar furrows, a convex preglabellar field, a convex rather than flat anterior border, and a spinose pygidium, and in lacking a plectrum. These features suggest that it is an olenoid rather than papyriaspidid (0pik, 1961, 1963). Papyriaspis Whitehouse (1939, p. 217) is easily differentiated from Parapianaspis by its • 141.

tapered glabella and much more converging anterior branches of the facial suture.

Parapianaspis hunanensis gen. et sp. nov. Plate 60, figures l- 13; Plate 61, figures 1-10 2001b

Papyriaspidid? gen. et sp. nov., Peng, Babcock, and Lin; pl. 4, figs. 5, 6.

Etymology. From Hunan Province, South China. Diagnosis. As for the genus. Holotype. External mold of nearly complete cranidium (P1. 61, figs. l, 2, NIGP 138519) from collection W88.1.

Other material. Eleven cranidia, two librigenae, and seven pygidia (NIGP 1 3 8 5 0 6 - 138531) in collections P114, P123.6, and W88.1.

Diagnosis. As for the genus. Description. Cranidium subquadrate, length slightly shorter than width between palpebral lobes. Anterior border flat, of even width (sag.) medially with sides narrowing abaxially; plectrum weak. Glabella subrectangular, with weak carina, obtusely rounded anteriorly, occupying three-fourths cranidial length; sides somewhat sinuous, parallel to sagittal line; with four pairs of weak lateral glabellar furrows, all isolated from axial furrow; S1 curved, narrowing adaxially, extending inward and rearward for rather long distance; $2 short and wide or pit-like; $3 pit-like, closer to sagittal line than other furrows; $4 pitlike, short, S 1, $2 and $4 almost evenly spaced, $3 closer to $4 than to $2. Occipital ring furrow transverse or gently bowed rearward, deepened at sides, isolated from axial furrows; occipital ring of uniform width (sag.), about as wide (sag.) as L1, beating tiny node lying subcentrally. Preglabellar field flat, as wide (sag.) as or slightly wider than anterior border. Eye ridge thick, gently convex, about 60 ° to sagittal line; palpebral lobe arcuate with anterior end opposite $2 or $3, and posterior end extending to the midpoint of L1. Palpebral area flat, slightly narrower than glabella. Anterior branch of facial suture diverging about 60" until meeting anterior border furrow, then turning inward and forward to cross anterior border as gentle curve and meeting anterior cranidial margins on each side of sagittal line at point opposite axial furrow; posterior branch almost tranverse, enclosing narrow, blade-shaped posterolateral projection. Posterior border furrow weakly impressed, posterior border of linear-like ridge. Pygidium subrounded, relatively large in size, length nearly same as width to much longer than width. Axis long, slightly tapered rearward, consisting of 6-7 tings defined by shallow ring furrows, and a small rounded-triangular terminal piece reaching to border furrow. Lateral and posterior borders broad and flat, well defined; anterior border narrow and shallow. Pleural field narrower than axis, divided into 6-7 pleurae beating shallow, broad pleural furrows, defined by weak interpleural furrows. Posterior margin complete or curved anteriorly. Frontal field of cranidium with weak, radiating, anastomosing ridges; surface of pygidial axis finely wrinkled.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, • 142-

where it occurs in association with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus punctuosus Zone). Family RHYSSOMETOPIDAE Opik, 1967 Genus RHYSSOMETOPUS {3pik, 1967

Type species. Rhyssometopus (Rhyssometopus) rhyssometopus Opik, 1967 (p. 274-278, pl. 25, figs. 1-4; pl. 33, fig. 4; pl. 46, fig. 3; text-figs. 93-95) from the Acmarharchis quasivespa Zone, Mungerebar Limestone, northwestern Queensland, Australia; by original designation. Remarks. The diagnosis of Opik (1967, p. 273,274) is followed here. Rhyssometopus zhongguoensis Zhou in Zhou et al., 1977 Plate 62, figures 1-17 1977

Rhyssometopus (Rhyssometopus) zhongguoensis Zhou in Zhou et al., p. 209, pl. 61, figs. 19, 20. 1977 Rhyssometopus (Rhyssometopus) elongatus Zhou in Zhou et al., p. 209, pl. 61, fig. 21. 1982 Rhyssometopus (Rhyssometopus) zhongguoensis Zhou; Liu, p. 324, pl. 222, figs. 15, 16. 1982 Rhyssometopus (Rhyssometopus) elongatus Zhou; Liu, p. 324, pl. 223, fig. 10. 2001c Rhyssometopus (Rhyssometopus) zhongguoensis Zhou; Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 2, figs. 19, 20. 2001b Rhyssometopuszhongguoensis Zhou" Peng, Babcock, and Lin, p. 105, pl. 13, figs. 8, 9. Holotype. Partly exfoliated cranidium (Zhou et al., 1977, pl. 61, fig. 20; YIGM 70141) from the Huaqiao Formation, Chatian, Fenghuang, northwestern Hunan. New material. More than 50 sclerites, including cranidia, librigena, and pygidia (illustrated specimens NIGP 138532-138545) in collections, P344.6, P348, P356.5, P361.5, P363.5, and P[3- 2.75. Remarks. Two species, R. zhongguoensis and R. elongatus, described by Zhou (in Zhou et al., 1977) are from the Huaqiao Formation at two different but adjacent localities in Fenghuang, northwestern Hunan. According to Zhou (in Zhou et al., 1977), R. elongatus, which is represented by a single cranidium, differs from R. zhongguoensis by having a more elongate and convex glabella, a narrower fixigena, a wider (sag.) preglabellar field, and less distinct lateral bulges on the glabella. However, new material from the same formation in the same paleogeographic region as Zhou's (in Zhou et al., 1977) specimens shows that these distinctions fall within the range of variation of R. zhongguoensis. The new material agrees in all cranidial respects with the type material, and adds information concerning the librigena and pygidium. Among the species described under the subgenus Rhyssometopus (Rhyssometopus) by Opik (1967), R. zhongguoensis is most similar to R. rugiceps Opik, a species from the Acmarharchis quasivespa Zone of northwestern Queensland. R. zhongguoensis differs in having shorter and more arcuate palpebral lobes, more oblique distal parts of the paradoublural line on the anterior field of •

143

•

the cranidium (i.e, the frontal wrinkle, see Opik, 1967, text-fig. 93 for terminology), and a more segmented pygidial axis that bears four tings rather than three tings. Occurrence. The holotype is from the Huaqiao Formation, Chatian, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Liostracina bella Zone (equivalent to the upper part of the Linguagnostus reconditus Zone and the Glyptagnostus stolidotus Zone).

Family SOLENOPLEURIDAE Angelin, 1854 Genus CHANGQINGIA Lu and Zhu in Qiu et al., 1983 Changqingia Lu and Zhu in Qiu et al., 1983, p. 104; Zan, 1992, p. 253; Peng, Lin, and Chen, 1995, p. 280; Guo, Zan, and Luo, 1996, p. 105. Austrosinia Zhang and Jell, 1987, p. 91, 92; Zhang, Xiang, Liu, and Meng, 1995, p. 67. Type species. Changqingia shandongensis Lu and Zhu in Qiu et al. (1983, p. 105, pl. 36, figs. 9, 10) from the Changhia Formation, near Gushan, Tai'an, Shandong; by original designation. Remarks. The genetic concept of Lu and Zhu (in Qiu et al., 1983, p. 104) is followed here. We follow Zan (1992, p. 252) and Peng et al. (1995, p. 280) in suppressing Austrosinia Zhang and Jell as a junior synonym of Changqingia. Solenoparia jiangsuensis Lin and Zhou in Lin et al. (1983, p. 405, pl. 2, figs. 11, 12) from the Yangliugang Formation in a borehole at Kunshan, southern Jiangsu belongs to Changqingia. Changqingia intermedia (Walcott, 1906)

Plate 63, figures 6, 7 1906 1913 1913 1937 1942 1965

Ptychoparia (Liostracus) intermedia Walcott, p. 592. Solenopleura intermedia Walcott, p. 169, pl. 17, figs. 16, 16a. Shumardia sp. indet. Walcott, p. 241, pl. 7, fig. 9. Solenoparia hemicycla Resser and Endo, p. 290, pl. 47, fig. 23. Solenoparia intermedia (Walcott, 1906); Resser, p. 51. Solenoparia intermedia (Walcott, 1906); Lu, Zhang, Zhu, Qian, and Xiang, p. 201, pl. 34, figs. 28, 29. 1987 Austrosinia intermedia (Walcott, 1906); Zhang and Jell, p. 94, 95, pl. 39, figs. 5-8; pl. 40, fig. 9. 1995 Austrosinia intermedia (Walcott, 1906); Zhang, Xiang, Liu, and Meng, p. 68, pl. 30, figs. 6-8; pl. 40, fig. 9. Holotype. Nearly complete cranidium (Walcott, 1913, pl. 17, fig. 16, USNM 58072) from the Changhia Formation, near Chaomidian, north of Zhangxia, Shandong. New material. Two cranidia (NIGP 138547, 138548) in collection P81.

-144-

Remarks. This species was reassigned to Changqingia by Peng et al. (1995). A cranidium from the Huaqiao Formation of the Paibi section, northwestern Hunan, closely resembles the type material form Shandong, North China. Key features that warrant assignment of these specimens to C. intermedia include a cephalon with a narrow, conical glabella; narrow palpebral areas; small palpebral lobes; and fine granules together with a few scattered coarser granules, coveting the cranidial surface. Occurrence. The holotype is from oolitic limestone and shale in the Amphoton Zone of the Changhia Formation, Shandong, North China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Changqingia laevis Peng, Lin, and Chen, 1995 Plate 63, figures 1-5 1995 Changqingia laevis Peng, Lin, and Chen, p. 280, 281, pl. 2, figs. 1-4. 2001b Changqingia laevis Peng, Lin, and Chen" Peng, Babcock, and Lin, p. 102, pl. 3, fig. 16.

Holotype. Nearly complete exfoliated cranidium (Peng et al., 1995, pl. 2, fig. 1, NIGP 118818; P1. 63, fig. 3, herein) from the Goniagnostus nathorsti Zone, Huaqiao Formation, Paibi, northwestern Hunan. New material. Single cranidium (NIGP 138546) in collection P112.42 is added to the type material (Peng et al., 1995), which is in collection P123.6 (NIGP 118818-118820). Remarks. Changqingia laevis is characterized by having a smooth cranidial surface. It is similar to Changqingia puteata (Resser and Endo; see Zhang and Jell, 1987, pl. 34, figs. 9-12; pl. 122, figs. 8, 9; Zan, 1992, pl. 1, figs. 1, 2; Guo et al., 1996, pl. 47, fig. 9), but differs in having a more acute glabCia, narrower palpebral areas, and more clearly defined glabellar furrows. C. puteata is also differentiated in having a pitted cranidial surface. The new material includes one incomplete, testaceous cranidium from the same locality as the type series of C. laevis. The smooth cranidial surface suggests that it is conspecific with C. laevis. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the uppermost part of the Dorypyge richthofeni Zone and the basal part of the Pianaspis sinensis Zone (equivalent to the Ptychagnostus punctuosus Zone and the lowermost part of the Goniagnostus nathorsti Zone). Genus MENOCEPHALITES Kobayashi, 1935

Type species. Solenopleura acantha Walcott, 1905 (p. 88, 89; 1913, p. 173, 174, pl. 16, figs. 4, 4a, b) from the Poshania-Taitzuia Zone, Changhia Formation, Zhangxia, Shandong; by original designation. Other species. Zhang and Jell (1987, p. 102) reassigned the following species to Menocephalites" • 145.

Solenopleura acantha Walcott, 1905; Menocephalus agave Walcott, 1905; Solenopleura acidalia Walcott, 1905; Menocephalus adrastia Walcott, 1905; Solenopleura pauperata Walcott, 1906; Damesella blackwelderi var. minor Sun, 1924; Solenopleura triangularis Resser and Endo, 1937; and Menocephalites jiawangensis Zhang and Jell, 1987. Remarks. Zhang and Jell (1987) rediagnosed Menocephalites and suppressed Parataitzuia Chang, 1963 as a junior synonym of the genus. According to Zhang and Jell (1987), Menocephalites is characterized by a conical and globular glabella, and by a discontinuous anterior border furrow with the proximal ends of two branches not joining the preglabellar furrow. The genetic concept of Zhang and Jell (1987) is followed here. Menocephalites was known previously from the Amphoton and the Taitzuia-Poshania zones of the Changhia Formation on the North China Platform. Material from the basal Huaqiao Formation described herein from northwestern Hunan is its first record in South China. In northwestern Hunan the genus occurs prior to the appearance of Amphoton, which extends the range of the genus downward into the Ptychagnostus atavus Zone (equivalent of the Crepicephalina Zone of the North China Platform).

Menocephalites sp. cf. M. pauperata (Walcott, 1906) Plate 63, figures 8-11 cf. 1906 cf. 1913 cf. 1937

Solenopleura pauperata Walcott, p. 593,594. Solenopleura pauperata Walcott; p. 169, pl. 17, fig. 18. Solenopleura triangularis Resser and Endo (in part), p. 289, pl. 47, figs. 18-22, non pl. 34,

cf. 1942 cf. 1965 cf. 1987 2001b

Solenopleura pauperata Walcott; Resser, p. 52. Trachoparia pauperata (Walcott); Lu, Zhang, Zhu, Qian, and Xiang, p. 210, pl. 37, figs. 1, 2. Menocephalitespauperata (Walcott); Zhang and Jell, p. 105, pl. 43, figs. 1, 7-11. Changqingiachalcon (Walcott)" Peng, Babcock, and Lin, p. 101, pl. 1, fig. 12.

figs. 1, 2.

Holotype of Menocephalites pauperata. A cranidium (Walcott, 1913, pl. 17, fig. 18, USNM 58074) from the Amphoton Zone, Changhia Formation, near Dongyu, Shanxi, North China. Material. Three cranidia (NIGP 138549-138551) in collection P37. Remarks. One cranidium (P1. 63, figs. 10, 11) was assigned previously to Changqingia but the strongly convex glabella and the shape of the anterior border furrow suggest that it belongs to

Menocephalites. New cranidia agree in almost all respects, including surface ornamentation, with specimens of Menocephalites pauperata from the North China Platform (Walcott, 1913, pl. 17, fig. 18; also Zhang and Jell, 1987, pl. 43, fig. 8). The only significant difference between specimens from Hunan and those from North China is that the Hunan specimens have a less tapered glabella.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone).

• 146.

Menocephalites hunanensis sp. nov. Plate 64, figures 1-4

2001b Menocephalitessp. 1, Peng, Babcock, and Lin, p. 101, pl. 2, figs. 1, 2. Etymology. After Hunan Province. Holotype. Cranidium (P1.64, figs. 2, 3, NIGP 138553) from collection P68.8. Other material. One small cranidium and one pygidium (138552, 138554) in collections P64.5 and P68.8. Diagnosis. Menocephalites with glabella that tapers forward gently, and is obtusely rounded at the front; with weak lateral furrows; anterior border wide and upturned; surface with fine, densely spaced granules and coarse, scattered granules. Description. Cranidium subquadrate, length about width between palpebral lobes. Anterior border moderately wide, moderately upturned; anterior border furrow moderately deep, gently arched forward. Glabella convex, subrectangular, tapering gently forward, obtusely rounded anteriorly, almost reaching anterior border furrow; with 3 pairs of weak lateral glabellar furrows; S 1 long, diagonally directed, almost isolating subtriangular L1; $2 short, pitlike, located slightly anterior to midpoint between occipital and preglabellar furrows; $3 faint, pitlike, located at the level of inner end of eye ridge; occipital furrow transverse, deeply incised, with distal ends gently deflected forward; occipital ring long (sag.), narrowing distally, with posterior margin strongly bowed rearward. Palpebral lobe of moderate size, located subcentrally; eye ridge wide, weak, strongly oblique. Palpebral area about one third as wide as glabella. Anterior branches of facial suture subparallel; posterior branches directed diagonally on small cranidium, directed outward and gently rearward on large cranidium; posterior border furrow deep, widening abaxially. Pygidium subpentagonal, wider than long. Axis strongly convex, tapering gently rearward, with posterior end reaching posterior border furrow, beating probably 4-5 tings and terminal piece. Pleural field moderately convex; with curved, deep, incised pleural furrows and slightly shallower and narrower interpleural furrows; four pleural segments with ridge-like anterior bands and wide, convex posterior bands; border moderately wide, separated from pleural field by change of slope. Surfaces of cephalon and pygidium with fine, densely spaced granules and scattered coarse granules. Remarks. This species is assigned to Menocephalites because its anterior border is not confluent with the preglabellar furrow, and because it has a general similarity to other species assigned to the genus. Menocephalites hunanensis sp. nov. resembles Menocephalites adrastia (Walcott, 1906), which is based on a single fragmentary cranidium (Walcott, 1913, pl. 16, figs. 5, 5a; see also Lu et al., 1965, pl. 40, fig. 17; Zhang and Jell, 1987, pl. 43, fig. 6) and Menocephalites acidalia (Walcott, 1905), which is based on a fragmentary cranidium and a librigena (Walcott, 1913, pl. 16, figs. 8, 8a; see also Lu et al., 1965, pl. 38, figs. 11, 12). All of these species have almost identical granulation, and have similarly effaced lateral furrows. However, the shape of glabella, which is globular in M. adrastia, and subtrapezoidal in M. acidalia, and the more oblique eye ridges, differentiate both M. • 147.

adrastia and M. acidalia from the new species. One pygidium referred to M. acidalia by Zhang and Jell (1987, pl. 42, fig. 7) from Walcott's (1906) material is similar to that of the new species in such characters as the long axis, the curved pleural and interpleural furrows, the shallow border furrows, and the surface ornamentation. The pygidium is distinguished from M. hunanensis, however, by having a proportionally narrower and more effaced pleural field, with all the furrows except for the first one being faint, by having relatively narrower borders, and by having a more tapered axis.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Menocephalites? sp. 1 Plate 64, figures 5, 6

2001b Menocephalitessp. 2, Peng, Babcock, and Lin, p. 101, pl. 3, fig. 1. Material. One cranidium (NIGP 138555) in collection W64.7. Remarks. This cranidium is assigned questionably to Menocephalites because of its narrow, ridge-like anterior border, the shape of the anterior border furrow (which is continuous medially), and the presence of four pairs of lateral glabellar furrows. This species is similar to Menocephalites hunanensis sp. nov. but, in addition to the above-mentioned characters, it differs in having a more arched anterior border furrow, and converging rather than parallel anterior branches of the facial suture. The coarse, scattered granules on the cranidial surface are also reminiscent of the holotype of Solenoparia bigranosa Endo (1937, pl. 59, fig. 5; Zhang, 1963, pl. 1, figs. 12, 13). Zhang (1963) designated S. bigranosa as the type species of Trachoparia. T. bigranosa, however, differs in having a proportionally smaller, tapered glabella and a transverse anterior border furrow.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lowermost part of the Dorypyge richthofeni Zone (equivalent to the Ptychagnostus atavus Zone). Menocephalites? sp. 2 Plate 64, figures 7, 8 2001b

Menocephalites?sp. 1, Peng, Babcock, and Lin, p. 101, pl. 3, fig. 2.

Material. One cranidium (NIGP 138556) in collection P112.6. Remarks. The illustrated cranidium has a broad, gently convex anterior border, defined posteriorly by a discontinuous border furrow that is not confluent with the preglabellar furrow, and a subrectangular, moderately convex glabella with three pairs of faint lateral furrows. Fine granules are present on the posterior part of the glabella, on the fixigenae, and on the occipital ring. The specimen is questionably referred to Menocephalites because of the almost invisible granulation. • 148.

Otherwise it is identical in general features with other species assigned to Menocephalites.

Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lowermost part of the Pianaspis sinensis Zone (equivalent to the Ptychagnostus punctuosus Zone). Family SHUMARDIIDAE Lake, 1907

Remarks. Peng et al. (2003, p. 203) expanded the definition of the family slightly to include genera beating facial sutures and palpebral lobes, or having as few as four thoracic segments. In addition, Peng et al. (2003) referred Oculishumardia to this family. This expanded concept of the Shumardiidae is followed here. Here, one new genus, Limbishumardia, is added to the family. Yuan and Yin (1998) referred their genus Gaoluopingia to the Shumardiidae without comment. We agree that Gaoluopingia is similar to Zacompsus. Here, both genera are classified with the Alsataspididae (subfamily Hapalopleurinae) (see Volume 1 of this two-part set). Genus OCULISHUMARDIAPeng, Babcock, Hughes, and Lin, 2003

Type species. Oculishumardia hunania (Peng, Babcock, Hughes, and Lin, 2003, p. 204, text-fig. 5A-I), from the Linguagnostus reconditus and Glyptagnostus stolidotus zones, Huaqiao Formation, Wangcun and Paibi, northwestern Hunan; by original designation. Emended diagnosis. Shumardiidae having palpebral lobes; glabella with small, poorly defined anterolateral lobes, pair of bacculae, and weak lateral furrows; preglabellar furrow faint; preglabellar field long, sloping forward. Remarks. Oculishumardia is a shumardioid that retains palpebral lobes throughout holaspid ontogeny. Most other shumardioids are blind. Aside from Oculishumardia, exceptions are Limbishumardia gen. nov. and Akoldinioidia dydimacantha from the middle upper Cambrian of northwestern Hunan (Peng, 1992, figs. 20I-K; 21A, B). Limbishumardia has eyes throughout ontogeny, but Akoldinioidia loses its eyes in later holaspid stages. Except for the presence of eyes, O. hunania shows an overall similarity to previously known Shumardiidae. Oculishumardia hunania Peng, Babcock, Hughes, and Lin, 2003 Plate 31, figure 6b; Plate 65, figures 8-15" Plate 66, figures 1-9 ?1967 Larva gen. indet, et sp. indet. (in part), Opik, p. 387, 388, pl. 36, figs. 9, 12 only. Shumardioid gen. et sp. nov., Peng, Babcock, Lin, Chen, and Zhu, p. 165, pl. 2, figs. 13, 14. 2001 2001 b Shumardioid gen. et sp. nov., Peng, Babcock, and Lin, p. 105, pl. 14, figs. 12, 13. Oculishumardia hunania Peng, Babcock, Hughes, and Lin, p. 204, 205, text-fig. 5A-I. 2003

Holotype. Cranidium (Peng et al., 2003, text-fig. 5B, C, NIGP 132983; P1. 66, figs. 1, 2 herein) from the Glyptagnostus stolidotus Zone, Huaqiao Formation, Wangcun, northwestern Hunan. New material. More 10 cranidia (illustrated specimens NIGP 132882-132886, 133511, 133512, • 149.

138562-138564) in collections P326.9, P13-2.75, W277, W228.3, W243.6, W251.15, and W254.1.

Diagnosis. Same as for the genus. Description. Cranidium semicircular or subellepitical in outline, length 0.65-0.8 times width. Anterior border and anterior border furrow present or absent; if present, border furrow is shallow, and border is narrow. Glabella moderately convex, at least two-thirds cephalic length, parallel-sided, acutely rounded anteriorly, with small, rounded baccula adjacent to L1, and with frontal lobe slightly expanded, defined by broad, deep axial furrows; with 3 pairs of short lateral furrows; S1 directed rearward and inward; $2 transverse, located near midlength of glabella excluding occipital ring; $3 directed forward and inward, delimiting a small anterolateral lobe that is best observed on exfoliated specimens; frontal lobe expanded slightly. Occipital ring slightly longer (sag.) than L1, narrowing abaxially, bearing a tiny, centrally located occipital node; occipital furrow distinct, transverse. Preglabellar furrow shallow or poorly defined. Palpebral lobe of medium size, defined by faint palpebral furrow, located slightly posterior to midlength of cranidium. Fixigena moderately wide; palpebral area gently convex, width slightly narrower than glabella; posterior area spatuliform, beating distinct, transverse border furrow that shallows abaxially and does not meet facial suture; posterior border long, widening outward. Facial suture opisthoparian; anterior branch converging forward; posterior branch directed obliquely to nearly transversely. Remarks. This genus was discussed at length by Peng et al. (2003). Occurrence. The holotype is from the Glyptagnostus stolidotus Zone, Huaqiao Formation, Paibi and Wangcun, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi, Paibi-2, and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone and the Glyptagnostus stolidotus Zone). Genus LIMBISHUMARDIAgen. nov.

Etymology. From Latin, limb, border, combined with genetic name Shumardia, referring to the Shumardia-like, border-beating taxon. Type species. Limbishumardia wangcunensis gen. et sp. nov. Diagnosis. Shumardiidae with anterior cranidial border; glabella bluntly pointed anteriorly, with anterolateral lobes isolated from L3 and weakly defined bacculae; preglabellar field sloping strongly forward; palpebral lobe a thick ridge, located posteriorly. Remarks. Limbishumardia gen. nov. is classified within the Shumardiidae because the glabella is morphologically similar to the glabellas of Shumardia (Conophrys) acutifrons Liu (in Zhou et al., 1977, p. 147, pl. 46, figs. 4, 5; Peng, 1992, pl. 5, figs. 12-15; pl. 6, figs. 1-3) from the Lower Ordovician (Tremadocian) of northwestern Hunan, and Acanthopleura meridionalis Henderson (1983, fig. 3A, C-F) from the Lower Ordovician (Arenigian) of northeastern Queensland, Australia. These two shumardioids are characterized by having isolated anterolateral lobes and bluntly pointed fronts on the glabellas. S. (C.) acutifrons has short, notched lateral glabellar furrows that are almost • 150.

identical in morphology to the lateral glabellar furrows of Limbishumardia. As recorded previously (Peng et al., 2003), an anterior border and palpebral lobes are present in Oculishumardia; and facial sutures are present in Shumardia sp. cf. S. exophthalma Ross (Ludvigsen and Westrop, 1985, p. 300), S. (Conophrys) salopiensis (Callaway; see Fortey and Owens, 1991, p. 458), and Akoldinioidia didymocantha Peng (1992, figs. 20I-K; 21A, B). These features, although unusual among shumardioids, serve to reinforce the interpretation that Limbishumardia should be classified in the family Shumardiidae. The new genus represents the second in this family that was sighted throughout ontogeny. As is the case with Oculishumardia, the presence of functional eyes suggests a close affinity between ptychopariids and shumardioids. Limbishumardia, which ranges from the Proagnostus bulbus Zone, extends downward the stratigraphic range of the Shumardiidae.

Limbishumardia wangcunensis gen. et sp. nov. Plate 65, figures 1-7 2001b Conocoryphidgen. et sp. nov., Peng, Babcock, and Lin, p. 103, pl. 8, figs. 12, 13.

Etymology. From Wangcun, site of the section yielding the holotype of the species. Holotype. Cranidium (P1.65, figs. 1-5, NIGP 138559) from collection W 199.2. Other material. Two incomplete cranidia (138560, 138561) in collection W 199.2. Diagnosis. As for the genus. Description. Cranidium trapezoidal, slightly wider than long, with obtusely rounded anterior margin. Anterior border wide (sag.), gently convex; frontal field more than twice as wide as anterior border, slightly convex, sloping forward steeply; with preglabellar field strongly depressed posteriorly. Glabella conical, strongly convex, bluntly pointed anteriorly; with three pairs of lateral glabellar furrows; S1 and $2 deeply notched inward; $3 directed forward and slightly inward, isolating anterolateral lobe from L3; occipital furrow, transverse, shallow medially, deep laterally; occipital ring transversely subtriangular, with pair of shallow furrows laterally, parallel to posterior margin of occipital ring and confluent with distal end of occipital furrow; bacculae reniform, weak, isolated from glabella. Palpebral and posterior areas of fixigena rising rapidly outward, inclining strongly toward axial furrow; palpebral lobe large, wide, elevated almost as high as crest of glabella; palpebral lobe, together with raised part of posterior portion of preocular area, forms curved "wall" that extends outward and rearward from in front of preglabellar furrow to posterior border furrow; with anterior end close to axial furrow, opposite anterior end of anterolateral lobe, with posterior end opposite L1. Eye ridge short, linear, weak, extending almost transversely and downward across fixigena from anterior end of palpebral lobe to meet axial furrow just in front of anterolateral lobe. Anterior branch of facial suture converging gently forward, curving strongly inward after crossing border furrow, meeting anterior margin at point opposite anterolateral lobe; posterior branch probably short, directed diagonally, enclosing narrow (tr.) triangular posterolateral projection. Posterior border narrow (exs.) transverse, curving slightly inward distally; posterior border furrow deep, widening slightly abaxially. • 151"

Surface of frontal area, palpebral lobe, glabella, and occipital ring covered with coarse, densely spaced granules; palpebral area with scattered coarse granules.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicative of the Wanshania wanshanensis Zone (equivalent to the Proagnostus bulbus Zone). PTYCHOPARIOIDAE FAMILY UNCERTAIN Genus PSEUDOMAPANIA Yuan and Yin, 1998

Type species. Pseudomapania cylindrica Yuan and Yin, 1998 (in part, p. 158, pl. 4, figs. 12, 13, 15-17, non fig. 14) from the Glyptagnostus stolidotus Zone, Huaqiao Formation, at Jimachong, Yuping, eastern Guizhou; by original designation.

Remarks. Pseudomapania was classified within the Mapaniidae by Yuan and Yin (1998) when they erected the genus. It is a tiny trilobite, distinguished by a cranidium with a rectangular glabella; small palpebral lobes lying close to the axial furrow; a thick, convex anterior border beating a plectrum and diverging anterior branches of the facial suture. The lateral glabellar furrows are all connected with the axial furrows; the S 1 furrows are oblique and the $2 to $4 furrows are notched. Except for having a distinct plectrum, the genus bears little resemblance to Mapania. This is especially true of the shapes of the lateral glabellar furrows. Also unlike Pseudomapania, Mapania is a relatively large sized trilobite. Zhang and Jell (1987, p. 188) excluded from the Mapaniidae Australian genera, other than Mapania, that were included previously in the family by Opik (1961, 1967). Here, we also exclude Pseudomapania from the family. The most distinctive features of the Mapaniidae are the lateral glabellar furrows. The furrows resemble those of ceratopygiids, with a pair of apostrophe-like S 1 furrows that are unconnected with the axial furrows, and with $2 and $3 commonly unconnected from the axial furrows. A general similarity between Pseudomapania and Anomocarella suggests that Psuedomapania should be classified within the Anomocarellidae. However, this possibility remains uncertain because the clearly defined lateral glabellar furrows, the small palpebral lobes, and the narrow borders on the pygidium are inconsistent with referral to the Anomocarellidae.

Pseudomapania cylindrica Yuan and Yin, 1998 Plate 67, figures 1-18; Plate 68, figures 1-9 1998

Pseudomapania cylindrica Yuan and Yin (in part), p. 158, pl. 4, figs. 12, 13, 15-17, non fig.

1998

Pseudomapania trancata Yuan and Yin (in part), p. 158, 159, pl. 4, figs. 7-9, non figs. 10,

14. 11. ?1998 2001b

Cathayanella granulosa Yuan and Yin (in part), p. 149, 150, pl. 2, fig. 10 only. Pseudomapania cylindrica Yuan and Yin; Peng, Babcock, and Lin, p. 105, pl. 13, fig. 14.

Holotype. Cranidium (Yuan and Yin, 1998, pl. 4, fig. 16, NIGP 127948) from the Glyptagnostus stolidotus Zone, Jimachong, Yuping, eastern Guizhou. • 152-

New material. More than 100 sclerites including cranidia, librigenae, and pygidia (illustrated specimens NIGP 138575-138598) in collections P319.8, P378.25, W196.3, W199.2, W210.5, W211.7, W216.5, W219.7, W225, W228.3, and W299.2. Remarks. Large collections from the Huaqiao Formation in both the Paibi and the Wangcun sections of northwestern Hunan agree in all respects with Pseudomapania cylindrica from the same paleogeographic region in eastern Guizhou. Specimens show notable variation in glabellar shape and proportion, the course of the facial sutures, effacement of the glabellar furrows, and the curvature of the anterior cranidial margin. All the specimens bear granules on the surface; the granules are of variable size. Terrace lines are present on the anterior cranidial border and on the posterior part of the pygidium. The new material demonstrates that the differences that Yuan and Yin (1998) used to distinguish Pseudomapania trancata from P. cylindrica, the type species of the genus, fall within the range of variation of a single species, and this suggests that these two taxa are synonyms. Two pygidia and one librigena assigned to either P. cylindrica or its junior synonym P. trancata by Yuan and Yin (1998), may not belong to that species. The pygidia more likely belong to Fenghuangella. It is uncertain as to what genus the librigena belongs. Occurrence. The holotype is from the Glyptagnostus stolidotus Zone, Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the upper part of the Proagnostus bulbus Zone to the Glyptagnostus reticulatus Zone). Suborder

OLENINA

Superfamily Family

OLENOIDEA

OLENIDAE

Subfamily Genus

Burmeister, 1843

Burmeister, 1843

OLENINAE OLENUS

Burmeister, 1843

Burmeister, 1843

Dalman, 1827

Type species. Entomostracites gibbosus Wahlenberg, 1821, from the Olenus Zone, upper Cambrian of Sweden; by subsequent designation (Salter, 1864, p. 3). Other species. See Henningsmoen (1957) and the supplement of Rushton (1983). To these lists, a new species, Olenus punctatus sp. nov., is added here. Remarks. Henningsmoen (1957, p. 96-100) and Rushton (1983, p. 121-124) discussed the genetic concept, and that concept is followed here. Olenus austriacus Yang in Zhou et al., 1977 Plate 69, figures 1-8; Text-figure 20 ?1922

Olenus transversus? Westerg~d (in part), p. 126, pl. 3, figs. 16, 17 only. • 153"

1972 1977 1978 1982 1983 1992 2001c

sp. 1, Taylor and Rushton, p. 20, pl. 4. Yang in Zhou et al., p. 152, pl. 47, fig. 5. Yang; Yang, p. 38, 39, pl. 6, figs. 1, 2. Yang; Liu, p. 307, pl. 216, fig. 9. Yang; Rushton, p. 125, pl. 16, figs. 7, 8, 11, 12; text-fig. 5a. Olenus austriacus Yang; Peng, p. 54, 55, fig. 22G-I. Olenus austriacus Yang; Peng, Babcock, Lin, Chen, and Zhu, pl. 5, figs. 10, 11, pl. 6, figs. 3, 5, 10, 15. 2001d Olenus austriacus Yang; Peng, Babcock, Lin, and Chen, p. 142, fig. 10.7. Olenus Olenus Olenus Olenus Olenus

austriacus austriacus austriacus austriacus

Holotype. By monotypy; incomplete, partly exfoliated exoskeleton (Zhou et al., 1977, pl. 47, fig. 5,

CUGB 0318105; refigured by Yang, 1978, pl. 6, fig. 1 and by Liu, 1982, pl. 216, fig. 9), from the C h u a n g i a - P r o c h u a n g i a Zone, Huaqiao Formation at Tingziguan, Fenghuang, western Hunan. material. Numerous sclerites including exoskeletons, cranidia, librigenae, and pygidia (illustrated specimens NIGP 133568, 133569, 133576, 133580, 138603-138605)in collections PI3-1.8, PI3-1.6, PI3-1.5, P[3-0.3, PI322.4, PI327.2, and PI336.

New

Diagnosis. See Rushton (1983, p. 126).

Rushton (1983) described the first complete exoskeleton of this species (from the Nuneaton district, central England), and emended the species diagnosis. Rushton (1983) also discussed features that differentiate O. austriacus from such species as O. ogilvis Opik, 1963 and O. transversus Westerg~d, 1922.

Remarks.

Text-figure 20. Holotype of Olenus austriacus Yang in Zhou et al., 1977. Incomplete exoskeleton, CUGB 0318105, × 4 (original of Zhou et al., pl. 47, fig. 5" also Yang, 1978, pl. 6, fig. 1).

• 154-

New material from the Paibi section is identical in all respects with the type material from Fenghuang (Yang in Zhou et al., 1977; Yang, 1978) and the material from the Cili-Taoyaun area, northwestern Hunan (Peng, 1992) and from central England. Key characters of this species include a rectangular glabella that is longer than wide, parallel-sided, and bearing three pairs of lateral furrows; a wide (sag.) preglabellar field; long palpebral lobes; forwardly diverging anterior branches of the facial suture; and sinuous posterior branches of the facial suture. An articulated specimen from northwestern Hunan is identical to those from central England in having a thorax of 13 segments, and a pygidium that is about half as long as wide, bearing a rather narrow axis, and having narrow or linear borders. In East Asia, Olenus has been reported from South Korea and from South China (Zhejiang, Guizhou, and Hunan). Olenus asiaticus Kobayashi (1944, p. 229-232, pl. l a, b; also Lee and Choi, 1994, p. 129-132, pl. 2, figs. 13-15) from the Glyptagnostus reticulatus Zone of the Machari Formation, South Korea, is distinguished from O. austriacus mainly by its shorter, forwardly tapered glabella, a thorax that bears widely spaced fulcra, and a proportionally longer, subtringular pygidium. O. sinensis Lu, from the much higher Lotagnostus punctatus Zone of the Siyangshan Formation, western Zhejiang (Lu in Lu and Qian, 1964; also Lu and Lin, 1989, pl. 19, figs. 1, 2), has a broad (sag.) preglabellar field similar to O. austriacus, but differs in having converging rather than diverging anterior branches of the facial suture, shorter palpebral lobes, and wider (exs.) posterior areas of the fixigenae. O. guizhouensis Lu and Chien (in Yin and Li, 1978, p. 475,476, pl. 163, fig. 3; also Lu and Qian, 1983, p. 45, 46, pl. 6, figs. 3-5) from the upper Cambrian of southeastern Guizhou has diverging anterior branches of the facial suture similar to O. austriacus, but is readily differentiated by having a shorter glabella, shorter palpebral lobes, and a shorter preglabellar field. The thorax of O. guizhouensis shows some resemblance to that of O. austriacus but differs in having 15 rather than 13 segments, and having more widely spaced fulcra so that its pleural field is much wider than the axis.

Occurrence. The holotype is from the Chuangia-Prochuangia Zone, Huaqiao Formation at Tingziguan, Fenghuang, northwestern Hunan, China. In England, it is from the Olenus gibbosus Subzone of the Olenus-Homagnostus obesus Zone. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs in association with trilobites indicative of the Liostracina bella Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone). Olenus punctatus sp. nov. Plate 68, figures 14a, 15; Plate 70, figures 1-13 2001c Olenus austriacus Yang; Peng, Babcock, Lin, Chen, and Zhu, p.166, pl. 4, figs. 12-14. 2001b Olenus austriacus Yang; Peng, Babcock, and Lin, p.105, pl. 15, fig. 13.

Etymology. From Latin, punctatus, punctate, referring to its punctate surface ornamentation. Holotype. Exoskeleton (P1. 70, fig. 7, NIGP 138610) from collection PI335.4. Other material. More than 20 sclerites including exoskeletons, cranidia, librigena, and pygidia (illustrated specimens NIGP 133544, 133545, 138601, 138602, 138606-138613)in collections P13-1.80, P1335.4, P1336, and P1356.5. • 155-

Diagnosis. Olenus with glabella tapered gently; preglabellar field long; palpebral lobes long; eye ridge transverse; anterior branch of facial suture diverging strongly; thorax with 15 segments; segments with short pleural spines. Pygidium short, transverse, length less than half width. Surface covered with dense punctae. Remarks. Olenus punctatus sp. nov. has punctate ornamentation that differentiates it from all other species of Olenus. The new species is most similar to Olenus austriacus Yang (in Zhou et al., 1977, p. 152, pl. 47, fig. 5), with which it co-occurs in the upper Glyptagnostus reticulatus Zone of the Huaqiao Formation in northwestern Hunan. Aside from having punctae, the new species differs in having a more tapered glabella, more strongly diverging anterior branches of the facial suture, a thorax that bears 15 rather than 13 segments, and a proportionally shorter and wider pygidium with a broader axis. The presence of a tapered glabella and long preglabellar field in O. punctatus invites comparison with O. asiaticus Kobayashi, 1944 (p. 229, 230, text-fig, l a, b; see also Kobayashi, 1962, pl. 9, figs. 16a, b; Lee and Choi, 1994, pl. 2, figs. 13, 14) and O. sinensis Lu (in Lu and Qain, 1964, p. 35, pl. 7, fig. 6; also Lu and Lin, 1989, pl. 19, figs. 1, 2). O. asiaticus can be differentiated from O. punctatus, however, by its lack of surface punctae; and by the presence of a proportionally shorter cranidium, shorter palpebral lobes, and parallel-sided or converging rather than diverging anterior branches of the facial suture. O. sinensis can be differentiated from O. punctatus by its lack of surface punctae; and by the presence of converging anterior branches of the facial suture, and by the number of thoracic segments (13 rather than 15 segments). O. guizhouensis Lu and Chien (in Yin and Li, 1978, p. 475, 476, pl. 163, fig. 3; also Lu and Qian, 1983, pl. 6, figs. 3-5) has the same number of thoracic segments as O. punctatus, but it differs in having shorter palpebral lobes, a concave anterior border, divergent-straight rather than divergent-sinuous posterior branches of the facial suture, and surface sculpture consisting of fine, densely anastomosing ridges. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Liostracina bella Zone and the Chuangia subrectangulata Zone (equivalent to the Glyptagnostus stolidotus Zone and the Glyptagnostus reticulatus Zone). Subfamily PELTURINAECorda, 1847 Genus SULCARECLAVAgen. nov.

Etymology. From Latin, sulcare, furrowed, and clava, club; in reference to the furrowed, clubshaped glabella in large holaspides. Type species. Sulcareclava sagitta gen. et sp. nov. Diagnosis. Olenidae with cranidium subtrapezoidal; anterior margin expanded anteromedially; glabella narrower than posterior area of fixigena, slightly concave along sides, expanded anteriorly in large holaspides, with anterolateral comers rather truncate, with medial depression at front; lateral glabellar furrows consist of 4 distinct pairs; S 1 transcurrent, directed medially and slightly rearward; $2 directed medially and slightly rearward; $3 elongate-pitlike, directed medially, separated from axial furrow; $4 elongate-pitlike, directed anteromedially; occipital furrow transcurrent with posterior border furrow; palpebral lobe small, located in anterior one-third of cranidium; eye ridge weak, directed slightly anteromedially; fixigena narrow in anterior part, wide in posterior part. •

156.

Remarks. Sulcareclava is a monospecific genus.The cranidium of Sulcareclava gen. nov. is similar to cranidia of both Peltura Milne-Edwards, 1840 and Protopeltura Br6gger, 1882. Sulcareclava differs in having a more expanded anterior glabella with more convex lateral sides, four rather than three lateral glabellar furrows, a more posteriorly located palpebral lobe, a wider fixigena, and divergent rather than convergent anterior branches of the facial suture. Protopeltura can be further differentiated by having a wider preglabellar field.

Sulcareclava sagitta gen. et sp. nov. Plate 3 l, figures 11-14

Etymology. From Latin, sagittus, arrow; in reference to the arrow-shaped anterior part of the glabella.

Holotype. Cranidium (P1. 31, fig. 14, NIGP 138209) from collection P1360.3, Chuangia subquadrangulata Zone, Huaqiao Formation, Paibi, Huayuan, northwestern Hunan, China. Other material. Two cranidia (NIGP 138208, 138210) in collections W277 and P1360.3. Diagnosis. As for the genus. Description. Olenidae with cranidium subtrapezoidal, moderately convex. Anterior margin expanded anteromedially; preglabellar area small (sag., tr.), depressed, limited to arcuate area in front of glabella. Glabella long, extending to anterior boder in holaspids, narrower than posterior area of fixigena, slightly concave along sides, expanded anteriorly in large holaspides, with anterolateral comers rather truncate, with medial depression at front; lateral glabellar furrows consist of 4 distinct pairs; S1 transcurrent, directed medially and slightly rearward; $2 directed medially and slightly rearward; $3 elongate-pitlike, directed medially, separated from axial furrow; $4 elongate-pitlike, directed anteromedially, barely separated from axial furrow. Occipital ring moderately wide medially, narrowing at sides, with small medial node; occipital furrow well impressed, nearly straight, transcurrent with posterior border furrow. Palpebral lobe small, located opposite L3; eye ridge weak, directed slightly anteromedially; fixigena narrow in anterior part, wide in posterior part. Posterior border narrow close to occipital ring, width expanding slightly distally; palpebral furrow well impressed, nearly straight. Surface covered with fine granules. Thorax and pygidium unknown.

Remarks. Sulcareclava sagitta gen. et sp. nov. is represented by three cranidia, the smallest one representing a probable meraspid, and the other two representing successive holaspid stages. Through the available ontogenetic sequence, the glabella increases slightly in length, the frontal area of the glabella expands in size, the lateral glabellar furrows appear and then become better impressed, the anterior margin becomes bowed forward, and the anterior part of the fixigena becomes narrower.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi-2 and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the Liostracina bella Zone through the upper part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus stolidodus to the Glyptagnostus reticulatus Zone). • 157.

Suborder HARPINAWhittington, 1959 ?Family Genus

HARPIDIDAE

Whittington, 1950

BAIKADAMASPIS

Ergaliev, 1980

Baikadamaspis Ergaliev, 1980, p. 145; Peng, 1987, p. 114; Lisogor, Rozov, and Rozova, 1988, p. 73. Pterocephalops Li & Zhang in Zhu et al., 1979, p. 91, 92. non Pterocephalops Rasetti, 1944, p. 254. Cathayanella Yuan and Yin, 1998, p. 148, 149. Pterocephalopsinus Jell & Adrain, 2003, p. 436. Type species. Baikadamaspis proprius Ergaliev, 1980 (p. 145, 146, pl. 10, figs. 7-10) from the Glyptagnostus reticulatus Zone and Homagnostus longiformis Zone, Malyi Karatau, Kazakhstan; by original designation.

Other species. Loganopeltoides sinensis Yang (in Zhou et al., 1977, p. 244, pl. 73, fig. 18; also Yang, 1978, pl. 13, figs. 11, 12) from the Chuangia-Prochuangia Zone, Huaqiao Formation at Tingziguan, Fenghuang, northwestern Hunan; Pterocephalops granulus Li & Zhang (in Zhu et al., 1979, p. 92, pl. 38, fig. 2) from the upper Cambrian, Chuanlagou, Qilian, eastern Qinghai; Baikadamaspis sp. (Peng, 1987, p. 114, pl. 13, fig. 11) from the Liostracina bella-Proagnostus bulbus Zone, Huaqiao Formation, Wa'ergang, Taoyuan, northwestern Hunan; Cathayanella granulosa Yuan and Yin, 1998 (in part, p. 149, 150, pl. 2, fig. 9, non fig. 10) from the Huaqiao Formation, eastern Guizhou. Four additional species, all from the Hauqiao Formation of northwestern Hunan, are Baikadamaspis sp. cf. B. granulosa (Yuan and Yin), Baikadamaspis linearis sp. nov., Baikadamaspis paibiensis sp. nov., and Baikadamaspis sp. Two possible representatives of Baikadamaspis are a fragmental cranidium reported as Guizhoucephalina? sp. indet. (Shergold, 1980, p. 66, pl. 20, fig. 8) from the Wentsuia iota-Rhaptagnostus apsis Assemblage-Zone of the Chatsworth Limestone, western Queensland, Australia; and a pygidium reported as Eugonocare? sp. indet. (Shergold, 1980, p. 61, pl. 35, fig. 7) from the Wentsuia iota-Rhaptagnostus apsis Assemblage-Zone of western Queensland, Australia. Emended diagnosis. Cephalon semicircular; cranidium with broad (sag. exs.) frontal area, transverse ridge or paradoublural line present or absent; glabella subrectangular, medium size, moderately convex, with 2-4 pairs of deeply incised lateral furrows; glabellar carina present or absent; occipital ring with median node; palpebral lobe small or tubercle-like, located subcentrally; eye ridge distinct, transverse or slightly oblique rearward; anterior branch of facial suture diverging forward gently to strongly, posterior branch diverging forward slightly to rearward slightly, enclosing broad (sag., exs.) posterior area of fixigena; fixigena with posterolateral projection subrectangular or subrhomboidal, extending posteriorly in some species; librigena with relatively small, narrow genal field. Pygidium transverse-elliptical or lenticular, with convex axis bearing 5 segments, extending nearly to posterior border furrow. Surface granulate.

Remarks. Ergaliev's (1980) genetic concept is emended on the basis of large collections from China that show all cranidial and pygidial features except for those of the hypostome. Baikadamaspis is assigned tentatively to the family Harpididae based on its general similarity to Entomaspis Ulrich in Bridge, 1931, Loganopeltoides Rasetti, 1945, and Fissocephalus •

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Lermontova, 1951, both in cephalic and pygidial morphology (see '~'ang in Zhou et al., 1977; Jell and Adrain, 2003). Assignment of the genus to the Paracedaiidae (Ergaliev, 1980; Yuan and Yin, 1998) or to the Eulomiidae (Jell and Adrain, 2003) is rejected here. The presence of a rostral plate defined by subparallel connective sutures renders the superageneric classification of the genus inconclusive. Baikadamaspis may have a close affinity with Loganopeltiodes. Rasetti (1945) originally interpreted Loganopeltoides as having a proparian facial suture, but later (Rosetti, 1948) interpreted the facial suture to be opisthoparian. Well preserved material shows that Baikadamaspis has opisthoparian sutures, similar to those of Loganopeltoides. Except for the widely separated anterior and posterior branches of the facial suture, the rearward direction of the posterior branch of the facial suture, and the larger genal field of the librigena, Baikadamaspis resembles Loganopeltoides in most features. Baikadamaspis and Loganopeltoides are quite similar in cephalic shape and convexity, the shape of the eye ridge, the shape and segmentation of the glabella, the size and position of the palpebral lobes, and the presence of bacculae. Baikadamaspis is slightly older than Loganopeltoides (which occurs in Furongian strata), and it may be ancestral to it. Baikadamaspis is similar to Sailoma Schrank, 1975 and is possibly synonymous with it. However, so far Sailoma is only known from fragmental cranidia from Northeast China and Iran (Schrank, 1975, pl. 8, figs. 5, 6; Peng, Geyer, and Hamadi, 1999, fig. 32). Pending further information about Sailoma, the material from Hunan is retained as Baikadamaspis. Cathayanella Yuan and Yin, 1998 is regarded as a junior synonym of Baikadamaspis. Large collections of Baikadamaspis from northwestern Hunan show that the differential characters of Cathayanella listed by Yuan and Yin (1998) are of relatively little significance. The characters are regarded as being of intrageneric or even intraspecific significance. Yuan and Yin (1998) compared their Cathayanella with Kathrynia Westrop, 1986, which is based on K. limbata Westrop (1986, p. 80, 81, pl. 40, figs. 1-3; text-fig. 38) from upper Sunwaptan strata of the southern Canadian Rocky Mountains. The Canadian genus, however, can be easily distinguished from Baikadamaspis by its posteriorly located palpebral lobes, its broader (sag.) frontal area, and the anterior branches of the facial suture, which meet the anterior cranidial margin near the sagittal line.

Baikadamaspis sinensis (Yang in Zhou et al., 1977) Plate 74, figures 6-12; Text-figure 21 1977 1978 1982 1996

Loganopeltoides sinensis Yang in Zhou et al., p. 244, pl. 73, fig. 18. Loganopeltoides sinensis Yang; Yang, p. 70, 71, pl. 13, figs. 11, 12. Loganopeltoides sinensis Yang; Liu, p. 893, pl. 230, fig. 18. Loganopeltoides sinensis Yang; Zhou, Cao, Hu, and Zhao, p. 46, pl. 7, figs. 9, 10.

Holotype. By monotypy; incomplete cranidium (Zhou et al., 1977, pl. 73, fig. 18, CUGB 514034; refigured by Yang, 1978, pl. 13, figs. 11, 12; Text-fig. 21 herein), from the Chuangia-Prochuangia Zone, Huaqiao Formation, Tingziguan, Fenghuang, western Hunan. New material. One cranidium and one incomplete external mold of cranidium (NIGP 138656, 138657) in collections PD35.4 and P]336. Remarks. This species was assigned originally to Loganopeltoides by Yang (in Zhou et al., 1977), and it certainly bears great morphologic similarity to Laurentian species assigned to •

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Loganopeltoides (Rasetti, 1945, 1948). Similarities to Loganopeltoides include the glabellar shape, the short (tr.) eye ridge that is slightly oblique to the sagittal line, the small palpebral lobes, and the ornamentation (fine radiating ridges and closely species granules) on the genal area. However, examination of the type specimens of Baikadamaspis sinensis indicates that the species has an outward- and rearward-directed posterior section of the facial suture, a feature that is more typical of Baikadamaspis, particularly that of Baikadamaspis linearis sp. nov. Loganopeltoides by contrast, has a forward-outward directed anterior section of facial suture. In addition, the small paratype cranidium of B. sinensis (P1. 74, figs. 8, 9) is closely comparable in morphology with that of Baikadamaspis.

Text-figure 21. Holotype of Baikadamaspis sinensis (Yang in Zhou et al., 1977). Incomplete cranidium, CUGB 0308001, × 5 (original of Zhou et al., pl. 73, fig. 18; also Yang, 1978, pl. 13, fig. 12). This species was originally known from only two incomplete cranidia that were collected from the Huaqiao Formation of northwestern Hunan (pl. 74, figs. 8, 9; Text-fig. 21 herein). New material from the Paibi section is also fragmentary, but bears a complete anterior area, a feature that was incompletely known from the type material. The new specimens show that the anterior area of this species is about 1.5 times as long as the glabella with a very narrow, ill-defined anterior border.

Occurrence. The holotype is from the Chuangia-Prochuangia Zone, Huaqiao Formation, • 160-

Tingziguan, Fenghuang, northwestern Hunan, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi-2 sections, Hunan, where it occurs in association with trilobites indicative of the lower part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus reticulatus Zone).

Baikadamaspis sp. cf. B. granulosa (Yuan and Yin, 1998) Plate 74, figures 1-3 cf. 1998 Cathayanella granulosa Yuan and Yin (in part), p. 149, 150, pl. 2, fig. 9, non fig. 10 [? = Pseudomapania cylindrica Yuan and Yin, 1998].

Holotype of Baikadamaspis granulosa. Cranidium (Yuan and Yin, 1998, pl. 2, fig. 9, NIGP 127948), from the Formosagnostus formosus [=Hadragnostus modestus]-Blackwelderia Zone, Jimachong, Yuping, eastern Guizhou.

Material. Two cranidia (NIGP 138653, 138654) in collections P301 and P[3 60.3. Remarks. The material from the Huaqiao Formation, northwestern Hunan, resembles the holotype cranidium from the same paleogeographic region in eastern Guizhou in most respects, but differs in having a glabella that is more tapered in the anterior one-third rather than evenly tapered forward, and a transverse preglabellar ridge that is more close to the anterior border furrow.

Occurrence. The holotype is from the Formosagnostus formosus-Blackwelderia Zone, Jimachong, Yuping, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the Chuangia subquadrangulata Zone (equivalent to the lower part of the Linguagnostus reconditus Zone through the Glyptagnostus reticulatus Zone). Baikadamaspis linearis sp. nov. Plate 53, figure 10b, Plate 71, figures 1-16; Plate 72, figures 1-12; Plate 78, figure 12; Text-figure 22 2000b

Baikadamaspissp. 1; Peng, Babcock, and Lin, p. 104, pl. 10, figs. 7, 8.

Etymology. From Latin, linearis, line-beating, referring to the paradoublural lines on the frontal area and genal field of the cephalon.

Holotype. Cranidium (P1.72, figs. 3-5, NIGP 138633) from collection P319.6. Other material. More than 100 sclerites including cranidia, librigena, and pygidia in collections P298.4, P301, P309, P316.1, P319.6, among which 14 cranidia, five librigena, and six pygidia are selected as paratypes (NIGP 138614-138631, 138634-138652, 138686). Diagnosis. Baikadamaspis with broad frontal area that is slightly convex; frontal area sloping •

161

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forward posteriorly, and upturned anteriorly; beating distinct paradoublural line a short distance from glabella; anterior cranidial border poorly defined by broad (sag.) border furrow; glabella with four pairs of lateral furrows; S1 and $2 deeply incised; pair of tiny, obscure bacculae present; glabellar carina present; palpebral lobe tubercle-like; posterior branch of facial suture transverse or directed outward and forward before crossing paradoublural line; posterolateral projection of the fixigena prolonged rearward; pygidium lenticular, width twice length; with axis gently tapered rearward, pleural field obscurely segmented posteriorly; lateral border furrows wide.

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.

Text-figure 22. Reconstruction of cephalon and pygidium of Baikadamaspis linearis sp. nov. A, B, dorsal view of cephalon and pygidium based mainly on specimens NIGP 138624, 138627, and 138638 (see P1.71, figs. 12, 15 for cephalon and P1.72, fig. 10 for pygidium); C, ventral view of cephalon based on specimens NIGP 138620, 138640 (see PI. 71, fig. 8; P1.72, fig. 12). Description. Cranidium semicircular, moderately convex, length two-thirds width, anterior margin arched gently forward, genal angles rather elongated rearward; preglabellar area occupying about one-third of cranidial length. Anterior border strongly upturned, poorly defined by broad, concave border furrow; glabella subrectangular, moderately convex, posterior half nearly parallel-sided, anterior half tapering gently forward; glabellar front commonly obtusely rounded, truncate, rounded, or acutely rounded; glabellar carina weak; lateral glabellar furrows consist of 4 pairs; S 1 incised, directed inward and rearward, sometimes weakly bifurcated; $2 deep and short, isolated from axial furrow, transverse or subparallel to S 1; $3 shorter and shallower than $2, transverse, isolated from axial furrow; $4 faint, as short as $3, extending inward and forward from axial furrow immediately in front of eye ridge; occipital furrow transverse, deflected slightly forward, deep at sides; occipital ring narrow (sag.), gently convex (sag., exs.), beating prominent median node; bacculae small, sometimes obscure, raised slightly above fixigenae. Paradoublural line prominent, slightly sinuous, extending rearward across genal field, with posterior ends merged into genal angles before meeting posterior border furrow. Palpebral lobe small, tubercle-like, with distinct palpebral furrow, opposite $2; palpebral field about half glabellar width at level of $2; eye ridge raised, curved slightly, oblique slightly rearward. Anterior branch of facial suture with outward-convex curvature; posterior branch extending outward or outward and slightly forward to lateral border furrow, then turning rearward as a smooth curvature to cross lateral border furrow and border. Posterior border furrow deep, rather wide, transverse, with distal end deflected slightly rearward, becoming shallow before •

162-

meeting paradoublural line; posterior border narrow, transverse ridge adaxially, widened and deflecting rearward abaxially. Librigena with small genal field, border furrow broad; border poorly defined; genal spine flat, broad at base. Hypostome with ovate median body beating subrounded anterior lobe, crescentic posterior lobe, and anteriorly curved posterior margin. Posterior border absent, lateral border flat, widening rearward. Pygidium lenticular, width two or more times length, with rounded anterolateral comer. Axis tapered slightly rearward, occupying 0.8 of pygidial length, with thin, curved, bar-like articulating half-ring, 3 clearly defined tings, 1-2 poorly defined tings and thin, crescentic terminal piece. Pleural field moderately convex, with first two pleurae beating clearly defined pleural furrows, defined by weak interpleural furrows, posterior pleurae obscurely segmented. Borders narrow, defined by rather wide, shallow border furrows. Cephalic doublure moderately wide, extending rearward to genal spine, with inner part turned strongly dorsally; rostral plate subrectangular, defined laterally by straight connective suture that is subparallel to sagittal line. Pygidial doublure as wide as cephalic doublure. Weak concentric terrace lines developed on cephalic and pygidial doublure. Surface of cephalon and pygidium covered with dense granules.

Remarks. Baikakmaspis linearis sp. nov. resembles B. proprius in most important aspects of the cranidium and pygidium. These include the shape of glabella, the shape and direction of the S 1 and $2 furrows, the shape and the proportional length of the frontal area, the shape of the eye ridge, the course of the facial suture, the shape, proportion, and segmentation of the pygidial axis, and the segmentation of the pleural region. However, the new species is readily differentiated from B. proprius by its more curved border furrow, its smaller palpebral lobe, its narrower (tr.) palpebral area, its oblique eye ridge, its rearward elongated posterolateral projuction, its wider pygidial outline, its more effaced interpleural furrows, and by the presence of paradoublural lines on the frontal area. According to the description of Ergaliev (1980), the type species, B. proprius, has only 2 to 3 pairs of lateral glabellar furrows. Baikadamaspis granulosa (Yuan and Yin) differs in having a truncate-conical glabella, a more clearly defined anterior cranidial border, a transverse rather than oblique eye ridge, and a posterolateral projection that is not prolonged rearward. B. granulosa lacks the paradoublural line but has a transverse ridge on the frontal area.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Liostracina bella Zone (equivalent to the lower part of the Linguagnostus reconditus Zone). Baikadamaspis paibiensis sp. nov. Plate 73, figures 1-16; Plate 78, ?figures 13, 14; Text-figure 23 2001c Baikadamaspis sp., Peng, Babcock, Lin, Chen, and Zhu, p. 166, pl. 4, fig. 15. 2001b Baikadamaspis sp. nov. 2, Peng, Babcock, and Lin, p. 105, pl. 16, figs. 1, 7-9.

Etymology. From Paibi Village, where the holotype was collected. Holotype. A cephalon (Plate 73, figs. 7-9, NIGP 138646) in collections P378.25, P135.1. •

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Other material. More than 50 sclerites including a cephalon, cranidia, librigena, and pygidia (illustrated paratypes NIGP 138641-138645, 138647-138652, 138687) in collections P378.25, PI3-2.75, PI3-1.8, P135.1, PI322.4 and ?PI323.5.

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Text-figure 23. Reconstruction of cephalon and pygidium of Baikadamaspispaibiensis sp. nov., based on the specimen NIGP 138646 (see P1, 73, figs. 7-9 for cephalon and fig. 14 for pygidium).

Remarks. Baikadamaspis paibiensis sp. nov. from the Glyptagnostus reticulatus Zone in the Paibi-2 section has a higher stratigraphic occurrence than the other three species assigned to Baikadamaspis from the Paibi section. Morphologically, the new species is characterized by having a proportionally larger, moderately inflated glabella that is obtusely rounded anteriorly, three pairs of lateral glabellar furrows, an anteriorly sloping frontal field that lacks a paradoublural line or a transverse ridge, a thick anterior border that is arched strongly forward, palpebral lobe that is elongate-elliptical in outline, a straight eye ridge that is transverse, or oblique rearward slightly, a relatively short (tr., exs.) posterolateral projection that is somewhat extended rearward, and a pygidium that bears a subconical axis, a segmented pleural field, and poorly defined borders. The new species differs from both B. proprius and B. linearis sp. nov. The new species differs from B. paibiensis in having a shorter (tr.) posterolateral projection, a well-defined anterior cranidial border, and a more tapered pygidial axis. It differs from B. granulosa (Yuan and Yin) in having a more arched anterior border, a shorter (exs.) posterior area of the fixigena, longer palpebral lobes, and in lacking a transverse ridge on the frontal field. A hypostome that is from a slightly higher level than most of other specimens assigned to the new species is questionably referred to the species. Except for having slightly a narrow (tr.) posterolateral border, it seems indistinguishable from those assigned to Baikadamaspis linearis sp. nov. (P1. 72, fig. 9; P1.78, fig. 12). More material of hypostome is needed to clarify the assignment.

.164.

Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi and Paibi-2 sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Liostracina bella Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the Glyptagnostus stolidotus Zone through the G. reticulatus Zone). Baikadamaspis sp. Plate 74, figures 4, 5

Material. One cranidium, NIGP 138655, in collection P341.8. Remarks. The illustrated cranidium has a glabella with four pairs of lateral furrows, among which the S 1 furrow is long and strongly oblique rearward; a paradoublural line, which lies immediately in front of preglabellar furrow, is straight and transverse, an anterior border that is well defined; and a short (tr.) transverse eye ridge. The posterolateral projection of the cranidium is not significantly elongated rearward, and the anterior branches of the facial suture are gently divergent. The cranidium is similar to B. granulosa (Yuan and Yin), but B. granulosa has a poorly defined paradoublural line that is not extended to the posterolateral projection, and has a shorter S 1 furrow. This cranidium probably represents an undescribed species that has a close affinity with B. granulosa. However, limited material renders any decision about the systematics of this trilobite difficult. Occurrence. Dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the Liostracina bella Zone (equivalent to the upper part of the Linguagnostus reconditus Zone). TRILOBITA ORDER UNCERTAIN Family BURLINGIIDAEWalcott, 1908

Remarks. This family includes only two genera: Burlingia Walcott, 1908 and Schmalenseeia Moberg, 1903. Whittington (1994) discussed the burlingiids at length and provided a revised diagnosis for the family. As discussed by Whittington (1981), the origin and relationship of the burlingiids remains problematical. Genus SCHMALENSEEIAMoberg, 1903

Schmalenseeia Moberg, 1903, p. 93; Westergkd, 1922, p. 119; 1929, p. 8 (footnote); 1948, p. 3; Hupr, 1955, p. 198; Poulsen in Moore, p. 0293; Lazarenko in Kraskov, Lazarenko, Ogienko and Chernysheva, 1960, p. 253; Chernysheva, 1960, p. 130; Jago, 1972, p. 232, 233; Taylor and Rushton, 1972, p. 18 (on pl. 4); Rushton, 1978, p. 273; Yin and Li, 1978, p. 506; Yang, 1978, p. 63; Whittington, 1981, p. 595; Qiu, Lu, Zhu, Bi, Lin, Zhou, Zhang, Qian, Ju, Han, and Wei, 1983, p. 158, 159; Xiang and Zhang, 1985, p. 120; Buchholz, 1991, p. 107; 2000, p. 726; Whittington, 1994, p. 12; Ahlberg and Ahlgren, 1996, p. 131; Jell and Hughes, 1997, p. 100; Duan, Yang, and Shi, 1999, p. 163; Pegel, 2000, p. 1011, 1016; Yuan, Zhao, Li, and Huang, 2002, p. 132; Ebbestad and Budd, 2003, p. 1181.

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Type species. Schmalenseeia amphionura Moberg, 1903 (in part, p. 96-101, pl. 4, figs. 1-8, 10) from the Agnostus pisiformis Zone of Sweden; by monotypy. Other species. Schmalenseeia acutangula Westerggtrd, 1948, in part, p. 4, pl. 1, figs. 2-4, non figs. 5, 6 [= Burlingia jagoi (Whittington, 1994)], from the Tomagnostus fissus and Ptychagnostus atavus zones of the Paradoxides paradoxissimus beds, Gils6vshammar, southeast coast of Scania and J~imatland, Sweden; Schmalenseeia spinulosa Lazarenko (in Kraskov et al., 1960, p. 253, 254, pl. 53, fig. 18) from the Agnostus pisiformis Zone of the middle reaches of the Olenko River, North Siberian Platform; Schmalenseeia gostinensis Jago (1972, p. 233-236, pl. 44, figs. 19-22) from the Lejopyge laevigata Zone of Australian usage (Shergold, 1995; Lejopyge laevigata III Zone of Opik, 1961; or the middle Cambrian/upper Cambrian Passage Zone of Opik, 1967) at St. Valentine Peak, northwestern Tasmania; Schmalenseeia sp. cf. S. spinulosa Lazarenko (Rushton, 1978, p. 274, pl. 25, fig. 16) from the the Agnostus pisiformis Zone in the Outwoods Shale, Merevale No. 3 Borehole, Nuneaton, England; Schmalenseeia longa Ju in Qiu et al., 1983 (p. 159, pl. 51, figs. 9, 10), from the upper part of the Yangliugang Formation (middle Cambrian), Tonglu, northwestern Zhejiang; Schmalenseeia rara Zhao, Yuan, and Ahlberg in Yuan et al., 2002 (p. 132, 133, pl. 38, fig. 1), from the lower part of the Kaili Formation (lower middle Cambrian), Balang, Taijing, eastern Guizhou; Schmalenseeia athrotryphe Ebbestad and Budd, 2003 (p. 1182, pl. 3, figs. 1-5) from the lower part of the Lejopyge laevigata Zone, Paradoxides forchhammeri Stage in the Alum Shale, Ringsaker district, Norway; Schmalenseeia sp. indet. (sensu Whittington, 1981, pl. 1, figs. 11, 14; original of Moberg, 1903, pl. 4, fig. 9) which is probably an incomplete thoracopygon from the Agnostus pisiformis Zone of Sweden; Schmalenseeia sp. (sensu Xiang and Zhang, 1985, p. 120, pl. 37, fig. 1), which is based on a single thoracopygon from the Agnostascus orientalis [=Proagnostus bulbus] Zone, Keguqin, Jinghe, western Xinjiang; and the new species Schmalenseeia fusilis. Schmalenseeia transversa Ju in Qiu et al., 1983 (p. 159, pl. 51, fig. 11) is apparently a junior synonym of Schmalenseeia longa Ju. Both species are from the same horizon and locality, and are somewhat distorted. The relatively long preglabellar median ridge of S. transversa, which was the only difference discussed by Ju (in Qiu et al., 1983) was evidently caused by distortion due to compaction. Schmalenseeia jagoi Whittington, 1994 (p. 14, 15, pl. 4, figs. 7-10; also Westerg~rd, 1948, pl. 1, figs. 5, 6), from the Tomagnostus fissus and Ptychagnostus atavus zones, Paradoxides paradoxissimus beds (lower middle Cambrian), Brantevik, southeastern coast of Scania, Sweden was transferred to Burlingia by Ebbesta and Budd (2003). Emended diagnosis. Burlingiidae with exoskeleton functionally articulated or with cranidium, thoracic segments, and pygidium fused; glabella beating moderately deep to well-defined SO and S1-$3 furrows; median preglabellar ridge absent or variably present; posterior cephalic margin transverse or arched posteriorly; thoracopygon either articulated or fused; thorax with 7-17 segments; pygidium with 1-8 pairs of fused pleurae, posterior pair of pleurae and terminal area directed posteriorly. Remarks. Jago (1972) and Whittington (1981, 1994) discussed the genetic concept of Schmalenseeia extensively. Further discussion and cladistic analyses of burlingiids, including Burlingia and Schmalenseeia, were published by Ebbestad and Budd (2003). The type species of Schmalenseeia, S. amphionura from the Agnostus pisiformis Zone of Sweden, was the only species when Moberg (1903) erected the genus. Subsequently WestergS.rd (1948) described another species, S. acutangula, which is also from Sweden, but is much older than the type species (early middle Cambrian; equivalent to the early Wulingian Epoch in South China terminology). Jago (1972) • 166.

showed that the original material of S. acutangula included two separate forms, and both were sufficiently different from S. amphionura to be removed from Schmalenseeia. As Jago (1972) noted, both Form 1 and Form 2 lack preglabellar median ridges. Also, Form 2 of S. acutangula has a proportionally large glabella with shallow lateral glabellar furrows that are isolated from the axial furrow. In all these respects, these two forms are much more comparable to Burlingia, which also has an age equivalent to the early Wulingian. Whittington (1994) rejected Jago's (1972) view of excluding S. acutangula from Schmalenseeia, but concurred with him in separating the original material of S. acutangula. The original species name was restricted to Jago's (1972) Form 1 and, Schmalenseeia jagoi was erected to embrace Jago's (1972) Form 2. The genetic diagnosis provided by Whittington (1994) stresses the segmentation of the thoracopygon, in addition to the rather deeply impressed occipital and lateral glabellar furrows. Based mostly on the Swedish material, Schmalenseeia was diagnosed to have 7 or 8 segments in the thorax, and 7 or 8 fused segments in the pygidium. This diagnosis is emended here based on new material from western Hunan, which includes an unusually large exoskeleton of S. sinensis Yang, and a new species, Schmalenseeia fusilis, which has fused tagmata. The large exoskeleton of S. sinensis is about 11 mm, which is about 1.5 times the size of any previously described exoskeleton of Schmalenseeia (length ranges to 2.8 in S. amphionura and to 7.7 mm in S. spinulosa). It shows a multisegmented thorax with 17 segments, and a small, narrow pygidium, beating only one pleural segment and a long and narrow terminal area. S. fusilis sp. nov. is based on an ontogenetic series of exoskeletons, 1.6 to 7.8 mm long, all having fused tagmata. The largest exoskeleton has 13 segments in the thorax. The fused tagmata is an autapomorphy. Schmalenseeia has a wide paleogeographic distribution, suggesting that it was a pelagic form (Jago, 1972; Whittington, 1994). Representatives of the genus occur in the Wulingian, or equivalent strata of Sweden, Norway, England, Newfoundland, Siberian, Tasmania, India, Germany, and China (Hunan, Guizhou, Zhejiang, and Xinjiang). In China, all recorded occurrences of Schmalenseeia are from slope facies.

Schmalenseeia sinensis Yang in Yin and Li, 1978 Plate 75, figures 1-8; Text-figure 24 1978 1978 1999 2001b

Schmalenseeia sinensis Yang in Yin and Li, p. 506, pl. 169, fig. 9. Schmalenseeia sinensis Yang; Yang, p. 63, 64, pl. 13, fig. 10; text-fig. 6. Schmalenseeia sinensis Yang; Duan, Yang, and Shi, p. 163, figs. 12G, H. Schmalenseeia amphionura Moberg; Peng, Babcock, and Lin, p. 103, pl. 6, fig. 11.

Holotype. By monotypy; exoskeleton (Yin and Li, 1978, pl. 169, fig. 9, CUGB 0514034; refigured by Yang, 1978, pl. 13, fig. 10), from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, at Fengmuping, Tongren, eastern Guizhou. New material. An incomplete exoskeleton (with counterpart) and an incomplete cranidium (NIGP 138660, 138661) in collections P240.5 and W 132.5. Emended diagnosis. Schmalenseeia with conical glabella and deeply impressed SO and S1-$3 lateral furrows; L1, L2, and L3 each with medial node, L4 with two nodes sagittally; posterior cranidial margin forward-curved; posterior area rapidly widened abaxially, beating tiny genal spine posterolaterally; median preglabellar ridge extended into frontal lobe of glabella; paired boss • 167.

located on anterior ends of palpebral lobes; thorax with 17 segments ending in points. Pygidium with large semicircular axis, with one pair of pleurae directed rearward. Description. Exoskeleton subovate, width 0.75 length, maximum width at midlength of exoskeleton.

Cranidium semielliptical in outline, length half width; posterior margin arched strongly rearward; glabella conical, gently convex, obtusely pointed anteriorly, with occipital furrow deeply incised at side, interrupted medially by a large node or spine-base; S 1 and $2 furrows probably transglabellar, notably deep side; $3 moderately deep, transglabellar, deep at side, defining subtrangular frontal lobe; L1-L3 each beating median node, large in L2, L3; frontal lobe (L4) beating two small nodes, one close to $3 and the other close to anterior margin of frontal lobe; fixigena flat, with anterior area occupying about two-thirds of total cranidial length, beating strong preglabellar median ridge that extends slightly into anterior part of frontal lobe and linear ridge at each side; anterior border and anterior border furrow absent; posterior area occupied mostly by large posterolateral projection, widening sharply abaxially with straight lateral margin and short genal spine curving slightly rearward; both the anterior and the posterior margin of posterolateral projection raised as linear ridges; palpebral lobe long, narrow (tr.) and steeply inclined, with its anterior end opposite midpoint of L1 and posterior end opposite $3; palpebral area flat, narrow (tr.), separated from palpebral lobe by clearly defined palpebral furrow; eye ridge short, beating prominent node; anterior branch of facial suture straight, at angle of 30 ° to sagittal line; posterior branches nearly parallel to anterior branch of facial suture.

Text-figure 24. Reconstruction of dorsal exoskeleton of Schmalenseeia sinensis Yang, 1978, based on specimens NIGP 138660 and 138661 (see PI. 75, figs. 1-8).

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Thorax with 17 segments. Axis gently convex, occupying about one-quarter of thoracic width; axial tings of uniform length with linear half-tings; tings of anterior 10 segments each beating transverse elliptical convexity on posterolateral part; axial tings, 6, 10, 14?, bearing median node; pleural region flat; pleurae with straight edges and sharply pointed tips, lateral margins of pleurae becoming increasingly laterally oblique rearward until last 3-4 segments, which become medially oblique. Pygidium subrectangular in outline; lateral margins parallel to axial line, posterior margin arched forward; pleural region with two strongly concave pleurae lacking pleural furrows; second pygidial pleura with upturned posterior margin that looks like postaxial ridge; terminal area absent. Remarks. This species was erected on the basis of a single exoskeleton preserved in limestone. It is about 6.0 mm long and lacks the librigenae. Examination of the holotype by one of us (SP) shows that the cranidium has a narrow, conical glabella beating a node interrupting the SO furrow; and nodes on each of the L2, L3, and frontal lobes; a median preglabellar ridge; and a rearward-arched posterior margin of the pygidium. The thorax bears 15 segments, among which the sixth segment has an obscurely defined node, and a tiny pygidium beating probably only one pair of pleurae that is fork-like in shape. A reconstruction (Yang, 1978, text-fig. 6) of the holotype is inaccurate, both in morphology and magnification, as it shows only 14 segments in the thorax and a length of only about 5 ram, rather than 6 mm. Comparison with the new material from Hunan indicates that the holotype is a juvenile in the meraspid period. A new specimen (P1. 75, figs. 4, 6) shows that S. sinensis could reach lengths of at least 11 mm (cranidial length of at least 3-4 mm). Because some of the exoskeleton is crumpled and shifted to the left, the original size may have been larger than the measured length. Whittington (1981, 1994) noted that all species resembling Schmalenseeia amphionara known up to that time have a median preglabellar ridge, a paired boss on the fixigena beside the frontal glabellar lobe, and well-defined glabellar furrows are based on minor distinctions. These species include S. spinulosa from Siberia (Lazarenko in Kraskov et al., 1960), S. gostinensis from Tasmania (Jago, 1972), S. cf. S. spinulosa from England (Rushton, 1978), and S. sinensis and S. longa [=S. transversa] from South China (Yang, 1978; Ju in Qiu et al., 1983). Except for S. sinensis (herein), additional material has not been reported for any of these species since the time of Whittington (1994), so further comment is not warranted on most of them. New material of S. sinensis reinforces the distinction between S. amphionara and S. sinensis. S. sinensis differs from all other Schmalenseeia species in having 17 thoracic segments in the holaspid stage. The presence of the articulating half-tings in each of the last twelve segments indicates that these segments are articulated rather than fused. Another articulating half-ring separating the posterior margin of the 17th segment is the boundary separating the thorax from the pygidium. In addition to the segmentation, S. sinensis differs from S. amphionara in having a more anteriorly located occipital node; node-beating L1, L2, L3, and L4 lobes; a strongly curved, rather than transverse, posterior cranidial margin; more widely spaced anterior and posterior branches of the facial suture; a narrower (exs.) posterior area of the fixigena; and thoracic pleurae with more oblique lateral margins and more pointed tips. S. sinensis is most similar to S. spinulosa. S. spinulosa is based on a single exoskeleton that is 7.7 mm long. According to Lazarenko (in Kraskov et al., 1960, p. 253) and Rushton (1978, p. 274), S. spinulosa has a forward-curved posterior cranidial margin; spine- or large node-beating L1, L2, L3, and L4 lobes; distinct occipital and lateral glabellar furrows; a trace that continues from the median preglabellar ridge in front of L4; and a proportionally wide (sag.) preglabellar area. However, both the anterior and the posterior branches of the facial suture in S. spinulosa seem to be more widely diverging than those of S. sinensis, and the posterior area of the fixigena is probably wider (exs.) proximally and less rapidly widening abaxially. S. spinulosa further differs from S. •

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sinensis in having a proportionally smaller frontal glabellar lobe that bears one large, centrally located node, rather than two as present in S. sinensis. S. spinulosa has thoracic segments with less oblique lateral margins and less pointed tips than those in S. sinensis. Differences between the Chinese and the Siberian species seem to be minor, and the glabellar morphology of the Siberian species seems to be within the range of variation of the Chinese material of S. sinensis. More material is needed to evaluate whether S. sinensis should be regarded as a separate species from S. spinulosa. S. sinensis also resembles S. athrotryphe from the lower Lejopyge laevigata Zone of Norway. The two species are nearly indistinguishable in cranidial morphology. The Norwegian species has 18 to 19 thoracic segments, which is similar in number to S. sinensis. However, the Norwegian species has a relatively narrower (tr.) pleural region, less pointed tips in the pleural segments, and a larger pygidium containing nine segments. S. sinensis resembles S. gostinensis from Tasmania, but S. gostinensis can be differentiated by having a less curved posterior cranidial margin, more closely spaced anterior and posterior branches of the facial suture, and probably the pattern of nodes on the glabella. The Tasmanian species has nine segments in the thorax, and a larger, more segmented pygidium. Occurrence. The holotype is from the Paradamesops [=Parablackwelderia] jimaensisCyclolorenzella [=Torifera] tuma Zone, Huaqiao Formation, at Fengmuping, Tongren, eastern Guizhou, China. New material is from dark-gray limestones of the Huaqiao Formation in the Paibi and Wangcun sections, Hunan, where it occurs in association with trilobites indicative of the upper part of the Pianaspis sinensis Zone to the lowermost part of the Wanshania wanshanensis Zone (equivalent to the Lejopyge laevigata Zone). Schmalenseeia fusilis sp. nov. Plate 76, figures 1-12; Plate 77, figures 1-11 2001 Schmalenseeia sp.; Peng, Babcock, and Lin, p. 103, pl. 8, fig. 1.

Etymology. From Latin fusilis, melted, fused, referring the fully fused exoskeleton. Holotype. Mostly exfoliated exoskeleton lacking librigenae (P1.77, figs. 3-6, NIGP 138673) from collection P268.3. Other material. More that 20 exoskeletons in various stages of ontogenetic development (illustrated specimens NIGP 138662-138672, 138674, 138675), all in collection P268.3. Diagnosis. Schmalenseeia having all tagmata fused. Description. Exoskeleton elongate-elliptical, length twice width, gently convex. Cephalon semicircular, length about two-thirds width. Anterior border a thin ridge (P1. 76, fig. 9), defined posteriorly by thin and shallow anterior border furrow. Preglabellar field long, subequal in length (exs.) with preocular field, rather flat. Glabella conical, acutely rounded anteriorly; with 3 pairs of lateral glabellar furrows that are shallow, broad, transverse, moderately long, evenly spaced. Occipital furrow broad, deep, developed only at sides, interrupted by convex area that bears a median node; occipital ring narrow. Eye ridge short, weak, beating large, poorly-defined node • 170.

medially. Palpebral lobe large, located posteriorly, with anterior end opposite L1, posterior end opposite $3. Both anterior and posterior branches of facial suture diverging forward, subparallel to each other. Thorax and pygidium undivided. Axis about half as wide as pleural area; with 11-13 tings defined by transverse, broad ring furrows that are deepened at sides. Pleurae with thin ridge-like anterior and posterior bands and broad, shallow pleural furrows.

Remarks. Schmalenseeia fusilis sp. nov. is distinctive among species of Schmalenseeia by virtue of complete fusion of the tagmata. Occurrence. Dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Proagnostus bulbus Zone). UNDETERMINED TRILOBITE SCLERITES Family UNDETERMINED Genus Undetermined Undetermined larva Plate 66, figures 10-19

Material. Seven meraspid exoskeletons and three cranidia (NIGP 138565-138574) in collection W219.7. Remarks. Specimens represented in the ontogenetic sequence of an undetermined trilobite range from a late degree 1 protaspis to a degree 1 meraspis. Protaspid specimens are separated axially but the pleural lobes remain fused, and there is no articulating joint between the cephalon and pygidium. Meraspid degree 1 specimens show a thoracic segment and articulations separating the segment from the cephalon and pygidium. The earliest protaspid is subcircular in outline with a distinct, convex axis that extends nearly to the anterior margin and is furrowed only in the protopygidium; boundaries between pleural areas are marked by faint furrows. Through the observed part of the ontogenetic sequence, the exoskeleton becomes ellipsoidal, pits marking the positions of the lateral glabellar lobes appear, the palpebral areas take shape, the interpleural furrow appears in the thoracic segment, and pleural and interpleural furrows appear in the pygidium. Although the palpebral area begins to become apparent on the fixigena, the trilobite is apparently still eyeless in the earliest part of the meraspid period. The meraspid cranidium has a subcylindrical glabella that is parallel-sided anteriorly to S1, gently constricted at L1, and truncate anteriorly. The glabella bears four pairs of lateral furrows, with S 1 being broad and short, and deeply impressed inward from the axial furrow; $2, $3, and $4 are pitted and progressively shallowing forward. The occipital furrow is deep and transverse. The occipital ring has a rearward curving posterior margin, and possibly bears a median node anteriorly. The anterior border is narrow (sag.) and gently upturned; the anterior border furrow is shallow and transverse. The palpebral area of the fixigena is the same width as the glabella, and the posterior area is two to three times the width of the glabella. A preglabellar field is absent. The palpebral ridge is linear, long and transverse, with the inner end being shorter (sag.) and opposite the anterior lobe of the glabella; the palpebral lobe is gently curved, moderately large, and located anteriorly with the posterior end opposite the $2 furrow. Anterior branches of the facial suture are gently • 171-

convergent to parallel-sided, whereas the posterior branches are nearly diagonal, and each encloses a large, triangular posterolateral projection.

Occurrence. The material is from dark-gray limestones of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone to the lower part of the Chuangia subquadrangulata Zone (equivalent to the upper part of the Proagnostus bulbus Zone to the Glyptagnostus reticulatus Zone). Undetermined pygidium 1 Plate 45, figures 13, 14

Material. Two pygidia (NIGP 138368, 138369) in collections P261.5 and P282.6. Remarks. Two pygidia, tentatively regarded as representing the same species, are left in open nomenclature. Both specimens are semielliptical in outline and convex. Five distinct tings in the axis are present; the tings are separated by deep ring furrows. In one specimen (P1.45, fig. 14) the posterior axis is broken, but the other specimen (P1. 45, fig. 13) show that the posterior axis includes a short terminal piece and a distinctive, narrow postaxial ridge that extends to the posterior margin. The pleural fields consist of about four pleurae separated by interpleural furrows that are moderately impressed and shallowing posteriorly. Pleural furrows are weak. The lateral and posterior borders are moderately wide, becoming widest posteromedially, and slightly upturned. In both pygidia, width exceeds length; however, one specimen (P1. 45, fig. 13) is somewhat more elongate than the other specimen (P1.45, fig. 14). Differences between the specimens are presumed to represent intraspecific variation. Occurrence. The material is from dark-gray limestones of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the middle part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Proagnostus bulbus Zone). Undetermined pygidium 2 Plate 68 figure 10; Plate 74, figures 13-17

Material. Three pygidia (NIGP 138599, 138658, 138659) in collections P237.8 and P269. Remarks. Three broken pygidia, all evidently belonging to the same species, are represented among the new material from northwestern Hunan. The pygidia are broadly subelliptical in outline, much wider than long, flattened in the pleural areas, and convex axially. The axis is short, tapering backward rapidly, and contains four tings separated by distinct ring furrows and a short terminal piece. A distinct postaxial ridge extends afrom the terminal piee of the axis about midway onto the posterior border area. Four pleurae are present in each pleural field; interpleural furrows are moderately impressed. Plerual furrows are present in only the first two segments. Pleurae extend to the margin, and a distinct border is absent. The surface is covered with coarse granules. Occurrence. The material is from dark-gray limestones of the Huaqiao Formation in the Paibi • 172.

section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the upper part of the Lejopyge laevigata Zone to the lower part of the Proagnostus bulbus Zone). Undetermined librigena 1 Plate 78, figure 1

Material. One librigena, NIGP 138676, in collection P319.6. Remarks. A single librigena with an upturned lateral border that is obscurely defined by a slope change in the genal field rather than by a border furrow. The genal field is gently convex, having a smooth surface. This librigena is relatively large in size. This librigena is associated with a number of polymerids (see description of Paibi section in Volume 1 of this two-part set), but from the morphology and the size, it can not be assigned to any of them. Occurrence. The librigena is from dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Liostracina bella Zone (equivalent to the upper part of the Lejopyge laevigata Zoneto the lower part of the Linguagnostus reconditus Zone). Undetermined librigena 2 Plate 78, figures 2, 3

Material. Two librigenae, NIGP 138677, 138678, in collection P1372 and W228.3. Remarks. Two undetermined librigenae are characterized by having a wide lateral border that is flat and slightly concave anteriorly, a relatively narrow (tr.) genal field that is gently convex, and a moderately wide (exs.) posterior border. The course of the facial suture shows that the trilobite has a large, moderately curved, posteriorly located palpebral lobe and a short (exs., tr.) posterolateral projection. The librigenae may belong to Lobocephalina sinensis sp. nov. Cranidia of L. sinensis and the two undetermined librigenae occur in the Liostracina bella Zone but they are present in separate collections. Until more definitive evidence of a relationship between these sclerites becomes available, the librigenae are left unassigned. Occurrence. The librigenae are from dark-gray limestones of the Huaqiao Formation in the Wangcun and Paibi-2 sections, Hunan, where they occur in association with trilobites indicative of the Liostracina bella Zone to the lowermost part of the Shengia quadrata Zone (equivalent of the Linguagnostus reconditus Zone to the Glyptagnostus reticulatus Zone). Undetermined librigena 3 Plate 78, figure 4

Material. One librigena, NIGP 138679, in collection PI372.

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Remarks. A single undetermined librigena from the Huaqiao Formation is characterized by having a rounded genal angle, a broad genal field, and a moderately wide lateral border. The sclerite resembles the librigenae assigned to Monkaspis taizehoensis (Chu) by Zhu (1959, pl. 6, figs. 5, 6) from the Kushan Formation of Liaoning, but differs in having a narrower, more clearly defined border furrow, and, therefore, a wider genal field. The sclerite is also similar to librigenae assigned to Amphoton dios (Walcott) by Walcott (1913, pl. 21, fig. 13b; pl. 22, fig. 1a) and Zhang and Jell (1987, pl. 17, figs. 3, 8). The librigenae of A. dios reported by Walcott (1913) and Zhang and Jell (1987), however, are much older than the new specimen reported here. A. dios occurs in the Dorypyge richthofeni Zone in northwestern Hunan, whereas the new librigena is from the Shengia quadrata Zone. Occurrence. The librigena is from dark-gray limestone of the Huaqiao Formation in the Paibi-2 section, Hunan, where it occurs in association with trilobites indicative of the lowermost part of the Shengia quadrata Zone (equivalent to the upper part of the Glyptagnostus reticulatus Zone). Undetermined librigena 4 Plate 78, figure 5

Material. One librigena, NIGP 138680, in collection W211.7. Remarks. A single undetermined librigena is relatively small in size, has poorly defined borders, a broad genal field, and an angular genal angle without a genal spine. The course of the facial suture shows that the trilobite has a small palpebral lobe that is probably located anteriorly, and a relatively large posterior area of the fixigena. The librigena is associated with a variety of polymerids (see description of the Wangcun section in Volume 1 of this two-part set), but cannot be assigned to any of the co-occurring taxa at the present time. Occurrence. The librigena is from dark-gray limestone of the Huaqiao Formation in the Wangcun section, Hunan, where it occurs in association with trilobites indicative of the upper part of the Wanshania wanshanensis Zone (equivalent to the lowermost part of the Linguagnostus reconditus Zone). Undetermined hypostome 1 Plate 78, figure 9

Material. One hypostome, NIGP 138683, in collection P319.8. Remarks. A single hypostome has a central body with a short anterior; a long, inverted, subtrapezoidal posterior lobe that bears a pair of depressed areas posteriorly; and a long, deeply incised middle furrow. Scattered granules are present on the central body. The lateral borders are flat, upturned steeply, and have fine caecal ridges. This librigena is associated with various polymerid taxa (see description of the Paibi section in Volume 1 of this two-part set) but cannot be assigned to species at the present time. Occurrence. The hypostome is from dark-gray limestone of the Huaqiao Formation in the Paibi • 174.

section, Hunan, where it occurs in association with trilobites indicative of the lower part of the

Liostracina bella Zone (equivalent to the the lower part of the Linguagnostus reconditus Zone). Undetermined hypostome 2 Plate 78, figures 15, 16

Material. One hypostome, NIGP 138688 in collection P277. Remarks. This hypostome is characterized by having a forwardly curved anterior margin, a relatively wide anterior border, and a narrow posterior border. The posterior margin is straight, and angular at the posterolateral comers. The central body has a large anterior lobe, a small, crescentic posterior lobe, and a short middle furrow. The lateral border is upturned, widest (tr.) medially, narrowing forward and rearward. This hypostome differs considerably from the hypostome figured here as Undetermined hypostome 1, but is similar to hypostomes assigned to Baikadamaspis (P1.78, figs. 12-14).

Occurrence. The hypostome is from dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the the lower part of the Proagnostus bulbus Zone). Order

EODISCIDA

Suborder

Kobayashi, 1939

EODISCINA

Kobayashi, 1939

Superfamily EODISCOIDEA Raymond, 1913 Family EODISCIDAE Raymond, 1913 Genus HELEPAGETIAJell, 1975 1975 HelepagetiaJell, 1975, p. 82; Jell in Whittington et al., 1997, p. 384.

Type species. Helepagetia bitruncula Jell, 1975 (p. 82-84; pl. 29, figs. 1-11); by original designation.

Diagnosis. The genetic diagnosis of Jell (1975, p. 82) is followed here. Remarks. Helepagetia is a monospecific genus. As discussed by Jell (1975), Helepagetia is a pagetiid lacking facial sutures. Aside from the lack of facial sutures, this genus resembles Opsidiscus. Not all species assigned to Opsidiscus, however, lack facial sutures: O. longispinus Babcock, 1994 from North Greenland apparently lacks sutures. Only the type species of Helepagetia, H. bitruncula, has been placed in Helepagetia. This report represents the first record of Helepagetia outside of Australia.

•

175

•

Helepagetia bitruncula Jell, 1975 Plate 64, figures 9, 10 1975 Helepagetiabitruncula Jell, 1975, p. 82-84, pl. 29, figs. 1-11.

Material. One cranidium (NIGP 138557) in collection P261. Description. Cephalon semielliptical, convex, slightly indented anteromedially. Preglabellar field short, depressed. Glabella narrow, tapering anteriorly; S1 and $2 short, $3 forming shallow transglabellar furrow; occipital furrow shallow medially, deeper at sides; cranidial spine elongate; small median node behind occipital ring and near anterior of spine. Axial furrow deep, of uniform width. Palpebral lobe extends around anterior of eye gap; eye ridge and palpebral furrow absent. Anterior and lateral border convex, upturned; border furrow moderately deep anteriorly, shallowing laterally, with few poorly impressed scrobicules in lateral border furrow. Surface covered with fine, dense pustules and few larger granules. Remarks. A single, well-preserved cephalon from northwestern Hunan appears to be identical to specimens reported previously from Queensland and Tasmania, Australia (Jell, 1975). On the new specimen, the cephalic spine is broken, but enough remains to suggest that it is quite long, perhaps longer than the rest of the cephalon. H. bitruncula seems to be closest morphologically to Opsidiscus longispinus Babcock, 1994. The main diagnostic characters than can be used to distinguish O. longispina from H. bitruncula are: the presence of distinct eye ridges; the presence of eyes; and the presence of a glabella that is swollen just behind the anterior glabellar lobe. Occurrence. In Australia, H. bitruncula ranges from the P~chagnostus punctuosus Zone to the Lejopyge laevigata Zone. New material is from the dark-gray limestone of the Huaqiao Formation in the Paibi section, Hunan, where it occurs in association with trilobites indicative of the lower part of the Wanshania wanshanensis Zone (equivalent to the lower part of the Proagnostus bulbus Zone).

• 176-

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• 190.

INDEX OF SPECIES AND GENERA

A Adelogonus .........................................................................................................................................25 hunanensis ..................................................................................................................................2 6 Adelogonus? s p . . ................................................................................................................................ 27 Aethia ............................................................................................................................................... 121 rectangular ............................................................................................................................... 122 asiatica, Neoanomocarella .............................................................................................................. 110 attenuate, Pianaspis ......................................................................................................................... 137 austriacus, Olenus ............................................................................................................................ 153 B

Baikadamaspis ................................................................................................................................. 158 cf. granulose ............................................................................................................................ 161

linearis ..................................................................................................................................... 161 paibiensis ................................................................................................................................. 163 sinensis ..................................................................................................................................... 159 s p . . ............................................................................................................................................ 165

Baojingia ............................................................................................................................................9 8 jiudiantangensis ....................................................................................................................... 101 latilimbata ................................................................................................................................ 102 paralala .................................................................................................................................... 103 quadrata ................................................................................................................................... 104 subquadrata ............................................................................................................................. 106 tungjenensis .............................................................................................................................. 108 youshuiensis ...............................................................................................................................9 9 bigranulosum, Placosema ..................................................................................................................68 bitruncula, Helepagetia ................................................................................................................... 176 Buttsia ................................................................................................................................................5 2 globosa .......................................................................................................................................53 C cf. agonies, Lisania ............................................................................................................................9 4 cf. biserrata, Prodamesella .............................................................................................................. 126 cf. blackwelderi, Maotunia ................................................................................................................ 39 cf. butes, Proasaphiscus? ...................................................................................................................23 cf. changi, Sudanomocarina .............................................................................................................. 44 cf. granulosa, Baikadamaspis .......................................................................................................... 161 cf. inflata, Luyanhaoaspis ................................................................................................................ 132 cf. orientalis, Meteoraspis ................................................................................................................. 7 2

• 191-

cf. p a r v u s , P r o a s h a p h i s c u s ( H o n a n a s p i s ) ? ....................................................................................... 24 cf. p a u p e r a t a , M e n o c e p h a l i t e s ......................................................................................................... 146 cf. q u a d r a t a , C r e p i c e p h a l i n a ............................................................................................................. 70 cf. s e m i p l e c t r a , M a o t u n i a .................................................................................................................. 41 C h a n g q i n g i a ..................................................................................................................................... 144 i n t e r m e d i a ................................................................................................................................ 144 laevis ........................................................................................................................................ 145 c o n i f o r m a , F e n g h u a n g e l l a ................................................................................................................. 77 c o n s t r i c t a , P a r a y u j i n i a ...................................................................................................................... 50 c o n v e x a , Q i a n d o n g a s p i s .................................................................................................................. 115 c r a s s a , F e n g h u a n g e l l a l a o c h a t i a n e n s i s ............................................................................................. 76 C r e p ic e p h a l ina ................................................................................................................................... 69 cf. q u a d r a t a ................................................................................................................................ 70 p e r g r a n o s a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 C r e p i c e p h a l i n a ? sp. •.......................................................................................................................... 71 c u r v a t a , H u z h u i a ................................................................................................................................ 4 4 c u r v i t e n s u s , O n c h o n o t e l l u s ................................................................................................................ 67 c y l i n d r i c a , P s e u d o m a p a n i a .............................................................................................................. 152

D d e c o r o s a , L u y a n h a o a s p i s ................................................................................................................. 131 E

e u r y a x i s , K i n g s t o n i a .......................................................................................................................... 85 E y m e k o p s ? .......................................................................................................................................... 28 sp. 1 ............................................................................................................................................ 28 sp. 2 ............................................................................................................................................ 29

F e n g h u a n g e l l a ................................................................................................................................... 73 f u s i l i s .......................................................................................................................................... 80 c o n i f o r m a ................................................................................................................................... 77 c r a s s a , l a o c h a t i a n e n s i s .............................................................................................................. 76 l a o c h a t i a n e n s i s .......................................................................................................................... 74 l i o s t r a c i n a l a ............................................................................................................................... 79 F i s s a n o m o c a r e l l a ............................................................................................................................... 11 p a i b i e n s i s ................................................................................................................................... 11 f o r t i s , P a r a n o m o c a r e l l a ..................................................................................................................... 17 f u s i l i s , S c h m a l e n s e e i a ...................................................................................................................... 170 f u s i l i s , F e n g h u a n g e l l a ........................................................................................................................ 80 G g l o b o s a , B u t t s i a .................................................................................................................................. 53 G r a n d i o c u l u s ...................................................................................................................................... 29 o b s c u r u s ..................................................................................................................................... 31 t r u n c a t u s .................................................................................................................................... 32 g r a n u l o s a , H u a y u a n a s p i s .................................................................................................................. 34 I-I • 192

•

Helepagetia ...................................................................................................................................... 175 bitruncula ................................................................................................................................. 1 7 6 huananensis, S u d a n o m o c a r i n a ? ........................................................................................................ 4 5 huananensis, Szeaspis ........................................................................................................................ 21 H u a y u a n a s p i s ..................................................................................................................................... 3 3 granulosa ................................................................................................................................... 3 4 p e rpauca .................................................................................................................................... 35 H u a y u a n a s p i s ? sp. •............................................................................................................................ 3 6 H u a y u a n e l l a ..................................................................................................................................... 133 paibiensis ................................................................................................................................. 1 3 4 zhiqiangi ................................................................................................................................... 1 3 4 H u a y u a n i a .......................................................................................................................................... 11 subcalva ..................................................................................................................................... 12 quadrilateralis ............................................................................................................................ 12 hubeiensis, Paradistazeris .................................................................................................... 5 8 hunania, Oculishumardia ................................................................................................................ 149 hunanensis, A d e l o g o n u s ..................................................................................................................... 2 6 hunanensis, M e n o c e p h a l i t e s ............................................................................................................ 147 hunanensis, Paradistazeris ................................................................................................................ 5 8 hunanensis, Parapianaspis .............................................................................................................. 1 4 2 hunanensis, Zhujia ............................................................................................................................. 4 8 H u z h u i a .............................................................................................................................................. 6 2 paratypica .................................................................................................................................. 6 3 curvata ....................................................................................................................................... 6 4 latilimbata .................................................................................................................................. 6 5 hypostome

1, U n d e t e r m i n e d

............................................................................................................ 1 7 4

hypostome

2, U n d e t e r m i n e d

............................................................................................................ 175

ichangensis, Xilingxia .......................................................................................................................... 9 incilis, N e o a n o m o c a r e l l a ................................................................................................................. 112 inflata, L u y a n h a o a s p i s ..................................................................................................................... 1 3 2 Iniotoma ............................................................................................................................................. 3 7 laevis .......................................................................................................................................... 3 7 p o r o s u s ....................................................................................................................................... 3 8 intermedia, Changqingia ................................................................................................................. 144

J j i a n g n a n e n s i s , M a p a n i a ................................................................................................................... 1 2 9 jiudiantangensis, Baojingia ............................................................................................................. 101 K

Kingstonia .......................................................................................................................................... 85 euryaxis ...................................................................................................................................... 85

laevis, Changqingia ......................................................................................................................... 145 laevis, Iniotoma .................................................................................................................................. 3 7 laochatianensis, F e n g h u a n g e l l a ........................................................................................................ 7 4 • 193"

latilimbata, B a o j i n g i a ...................................................................................................................... latilimbata, H u z h u i a ..........................................................................................................................

102 65

l a r v a , U n d e t e r m i n e d .........................................................................................................................

171

l i b r i g e n a 1, U n d e t e r m i n e d ...............................................................................................................

173

l i b r i g e n a 2, U n d e t e r m i n e d

...............................................................................................................

173

l i b r i g e n a 3, U n d e t e r m i n e d

...............................................................................................................

173

l i b r i g e n a 4, U n d e t e r m i n e d ...............................................................................................................

174

L i m b i s h u m a r d i a ...............................................................................................................................

150

w a n g c u n e n s i s ...........................................................................................................................

151

linearis, B a i k a d a m a s p i s ...................................................................................................................

161

liostracinala, F e n g h u a n g e l l a .............................................................................................................

79

L i s a n i a ................................................................................................................................................

87

cf. a g o n i e s ..................................................................................................................................

94

p a i b i e n s i s ...................................................................................................................................

95

p a r a t u n g j e n e n s i s ........................................................................................................................

90

w a n g c u n e n s i s .............................................................................................................................

97

y u a n j i a n g ensis ............................................................................................................................

92

L i s a n i a ? s p . . .......................................................................................................................................

97

L o b o c e p h a l i n a ....................................................................................................................................

81

sinensis ....................................................................................................................................... L u y a n h a o a s p i s .................................................................................................................................

82 130

cf. inflata ..................................................................................................................................

132

d e c o r o s a ...................................................................................................................................

131

inflata .......................................................................................................................................

132

M

M a d a r o c e p h a l u s ................................................................................................................................

55

orientalis ....................................................................................................................................

56

M a o t u n i a ............................................................................................................................................

39

cf. b l a c k w e l d e r i ..........................................................................................................................

39

cf. s e m i p l e c t r a ............................................................................................................................

41

r e c t a n g u l a r i s ..............................................................................................................................

41

s p . . ..............................................................................................................................................

M a p a n i a ...........................................................................................................................................

43 128

j i a n g n a n e n s i s ...........................................................................................................................

129

M e n o c e p h a l i t e s ................................................................................................................................

145

cf. p a u p e r a t a ............................................................................................................................

146

h u n a n e n s i s ................................................................................................................................

147

M e n o c e p h a l i t e s ? sp. 1 ......................................................................................................................

148

M e n o c e p h a l i t e s ? sp. 2 ......................................................................................................................

148

M e t e o r a s p i s ........................................................................................................................................

71

cf. orientalis ...............................................................................................................................

72

N

N e o a n o m o c a r e l l a .............................................................................................................................

109

asiatica ..................................................................................................................................... incilis ........................................................................................................................................

110 112

N e o g l a p h y r a s p i s ..............................................................................................................................

122

nitida ........................................................................................................................................

123

• 194•

nitida, N e o g l a p h y r a s p i s ...................................................................................................................123 0 o b s c u r u s , G r a n d i o c u l u s .....................................................................................................................

31

o b v i a , P a r a n o m o c a r e l l a ? ...................................................................................................................

19

O c u l i s h u m a r d i a ................................................................................................................................149 h u n a n i a ....................................................................................................................................149 O l e n u s ..............................................................................................................................................153 a u s t r i a c u s .................................................................................................................................

153

p u n c t a t u s ..................................................................................................................................

155

O n c h o n o t e l l u s ....................................................................................................................................

66

c u r v i t e n s u s .................................................................................................................................

67

o r i e n t a l i s , M a d a r o c e p h a l u s ...............................................................................................................

56

P P a i b i e l l a .............................................................................................................................................

48

p a i b i e n s i s ...................................................................................................................................

49

p a i b i e n s i s , B a i k a d a m a s p i s ...............................................................................................................163 p a i b i e n s i s , F i s s a n o m o c a r e l l a ............................................................................................................

11

p a i b i e n s i s , H u a y u a n e l l a ...................................................................................................................134 p a i b i e n s i s , L i s a n i a .............................................................................................................................

95

p a i b i e n s i s , P a i b i e l l a ...........................................................................................................................

49

P a r a d i s t a z e r i s ....................................................................................................................................

57

h u b e i e n s i s ...................................................................................................................................

58

h u n a n e n s i s ..................................................................................................................................

58

r o t u n d u s .....................................................................................................................................

59

s p . . ..............................................................................................................................................6 0

p a r a l a l a , B a o j i n g i a ..........................................................................................................................103 p a r a l l e l a , P a r a n o m o c a r e l l a ...............................................................................................................

15

P a r a n o m o c a r e l l a ...............................................................................................................................

14

f o r t i s ...........................................................................................................................................

17

p a r a l l e l a .....................................................................................................................................

15

s i m i l a r i s ......................................................................................................................................

18

s p . . ..............................................................................................................................................19

P a r a n o m o c a r e l l a ? o b v i a .................................................................................................................... P a r a p i a n a s p i s ..................................................................................................................................

19 141

h u n a n e n s i s ................................................................................................................................142 p a r a t u n g j e n e n s i s , L i s a n i a ..................................................................................................................

90

p a r a t y p i c a , H u z h u i a ...........................................................................................................................

63

P a r a y u j i n i a ........................................................................................................................................

50

c o n s t r i c t a ....................................................................................................................................

50

p e r g r a n o s a , C r e p i c e p h a l i n a ..............................................................................................................

69

p e r p a u c a , H u a y u a n a s p i s .................................................................................................................... P i a n a s p i s ..........................................................................................................................................

35 135

a t t e n u a t e ...................................................................................................................................137 s i n e n s i s .....................................................................................................................................139 P l a c o s e m a ..........................................................................................................................................

68

b i g r a n u l o s u m .............................................................................................................................

68

p o r o s u s , I n i o t o m a ..............................................................................................................................

38

• 195"

Proasaphiscus .................................................................................................................................... 2 2 Proasaphiscus? cf. butes .................................................................................................................... 2 3 P r o a s a p h i s c u s (Honanaspis) ............................................................................................................. 2 3 cf. p a r v u s .................................................................................................................................... 2 4

P r o d a m e s e l l a ...................................................................................................................................

124

cf. biserrata ..............................................................................................................................

126

p u n c t a t a ....................................................................................................................................

125

tumidula ................................................................................................................................... 127 P s e u d o m a p a n i a ................................................................................................................................ 152 cylindrica ................................................................................................................................. 152 Ptychopariidae

g e n . a n d sp. i n d e t . •....................................................................................................

10

punctata, P r o d a m e s e l l a ................................................................................................................... 1 2 5 punctatus, Olenus ............................................................................................................................. 1 5 5 pygidium

1, U n d e t e r m i n e d

..............................................................................................................

172

pygidium

2, U n d e t e r m i n e d

..............................................................................................................

172

Q Qiandongaspis ................................................................................................................................. 113 convexa ..................................................................................................................................... 115 sinensis ..................................................................................................................................... 114 xiangxiensis .............................................................................................................................. 116 s p . . ............................................................................................................................................

117

quadrata, Baojingia .........................................................................................................................

104

quadrata, Shengia ............................................................................................................................ 118 quadrilateralis, H u a y u a n i a ................................................................................................................ 12

R rara, Townleyella ................................................................................................................................. 7 rectangula, Aethia ............................................................................................................................

122

rectangularis, M a o t u n i a .................................................................................................................... 41 R h y s s o m e t o p u s .................................................................................................................................

143

zhongguoensis ..........................................................................................................................

143

rotundus, Paradistazeris .................................................................................................................... 5 9

saginata, Sulcareclava .....................................................................................................................

157

Schmalenseeia ..................................................................................................................................

165

sinensis .....................................................................................................................................

167

fusilis ........................................................................................................................................ 170 Shengia ............................................................................................................................................. 117 quadrata ................................................................................................................................... 118 trapezia .................................................................................................................................... 119 w a n n a n e n s i s .............................................................................................................................

120

s p . . ............................................................................................................................................

121

similaris, P a r a n o m o c a r e l l a ............................................................................................................... 18 sinensis, B a i k a d a m a s p i s ..................................................................................................................

159

sinensis, Lobocephalina ..................................................................................................................... 8 2 sinensis, Pianaspis ...........................................................................................................................

139

sinensis, Qiandongaspis ................................................................................................................... 114 • 196.

s i n e n s i s , S c h m a l e n s e e i a ................................................................................................................... 167

sp., A d e l o g o n u s ? ................................................................................................................................

27

sp., B a i k a d a m a s p i s ........................................................................................................................... 165 sp., C r e p i c e p h a l i n a ? ..........................................................................................................................

71

sp., H u a y u a n a s p i s ? ............................................................................................................................

36

sp., L i s a n i a ? .......................................................................................................................................

97

sp., M a o t u n i a .....................................................................................................................................

43

sp., P a r a d i s t a z e r i s ..............................................................................................................................

60

sp., P a r a n o m o c a r e l l a .........................................................................................................................

19

sp., Q i a n d o n g a s p i s ...........................................................................................................................

117

sp., S h e n g i a ......................................................................................................................................

121

sp., S u d a n o m o c a r i n a ..........................................................................................................................

47

sp. 1, E y m e k o p s ? ................................................................................................................................

28

sp. 1, M e n o c e p h a l i t e s ? ..................................................................................................................... 148 sp. 2, E y m e k o p s ? ................................................................................................................................

29

sp. 2, M e n o c e p h a l i t e s ? ..................................................................................................................... 148 S t i g m a t o a ...........................................................................................................................................

83

y a n g z i e n s i s .................................................................................................................................

84

s u b c a l v a , H u a y u a n i a ..........................................................................................................................

12

s u b q u a d r a t a , B a o j i n g i a .................................................................................................................... 106 S u d a n o m o c a r i n a ................................................................................................................................

43

cf. c h a n g i ....................................................................................................................................

44

t r a y n o r a e ....................................................................................................................................

44

sp.. .............................................................................................................................................. 47 S u d a n o m o c a r i n a ? h u a n a n e n s i s ......................................................................................................... 45 S u l c a r e c l a v a ..................................................................................................................................... 156 s a g i t t a ....................................................................................................................................... 157 S z e a s p i s ..............................................................................................................................................

20

h u a n a n e n s i s ................................................................................................................................

21

T T o w n l e y e l l a ..........................................................................................................................................

7

r a r a ...............................................................................................................................................

7

t r a p e z i a , S h e n g i a .............................................................................................................................. t r a y n o r a e , S u d a n o m o c a r i n a .............................................................................................................. t r u n c a t u s , G r a n d i o c u l u s .................................................................................................................... t u m i d u l a , P r o d a m e s e l l a ...................................................................................................................

119 44

32 127

t u n g j e n e n s i s , B a o j i n g i a .................................................................................................................... 108

U U n d e t e r m i n e d t a x a ........................................................................................................................... 171 h y p o s t o m e 1 ............................................................................................................................. 174 h y p o s t o m e 2 ............................................................................................................................. 175 l a r v a .......................................................................................................................................... 171 l i b r i g e n a 1 ................................................................................................................................ 173 l i b r i g e n a 2 ................................................................................................................................ 173 l i b r i g e n a 3 ................................................................................................................................ 173 l i b r i g e n a 4 ................................................................................................................................ 174 p y g i d i u m 1 ............................................................................................................................... 172 • 197 •

p y g i d i u m 2 ...............................................................................................................................

172

W

wangcunensis, Limbishumardia ....................................................................................................... wangcunensis, Lisania ....................................................................................................................... wangcunensis, Wangcunia ............................................................................................................... wannanensis, Shengia ...................................................................................................................... Wangcunia ....................................................................................................................................... wangcunensis ...........................................................................................................................

151 97

140 120 140 140

X xiangxiensis, Qiandongaspis ............................................................................................................ 116 Xilingxia ............................................................................................................................................... 8 ichangensis ................................................................................................................................... 9 Y yangziensis, Stigmatoa ....................................................................................................................... yuanjiangensis, Lisania ...................................................................................................................... youshuiensis, Baojingia .....................................................................................................................

84 92 99

Z

zhiqiangi, Huayuanella .................................................................................................................... zhongguoensis, Rhyssometopus ....................................................................................................... Zhujia ................................................................................................................................................. hunanensis ..................................................................................................................................

• 198.

134 143 47 48

PLATES All illustrated specimens were coated with black ink and then magnesium oxide before being photographed. All specimens are preserved in limestone. Specimens having a horizon number with the prefix P are from the Paibi section; those with the prefix PI3 are from the Paibi-2 section; and those with the prefix W are from the Wangcun section. Unless otherwise stated, all specimens are deposited in the collections of the Nanjing Institute of Geology and Paleontology, the Chinese Academy of Sciences (NIGP).

•

199

•

Plate 1

Figures 1-8. Meteoraspis sp. cf. M. orientalis Yuan and Yin, 1998 1-5. Testaceous cephalon with fight librigena missing, in dorsal, anterolateral, oblique anterior, lateral, and anterior views, NIGP 137901, x 12.5, P317.4. 6. Testaceous librigena, NIGP 137902, x 10, P1370.7. 7, 8. Incomplete, partly exfoliated cranidium (8) and latex cast from its external mold (7), NIGP 137903, × 9, PI3 70.7. Figures 1-5 from Liostracina bella Zone; Figures 6-8 from Chuangia subquadrangulata Zone. Figures 9-11. Crepicephalina? sp. 9-11. Crushed, testaceous cranidium in anterolateral, dorsal, and anterior views, NIGP 137904, × 6, P45.6. From Dorypyge richthofeni Zone. Figures 12-14. Undetermined Ptychopariid. 12-14. Incomplete, partly exfoliated cranidium in dorsal, obliquely anterior and anterolateral views, NIGP 137905, all × 10, P 108. From the top part of Dorypyge richthofeni Zone. Figures 15, 16. Xilingxia ichangensis (Chang, 1964) 15, 16. Incomplete cranidium in anterolateral and dorsal views, NIGP 137906, × 12, W3. From the basal part of Dorypyge richthofeni Zone.

• 200-

Plate 1

't 4 •

it~ ~

/

,

11

/ f

. ~.~,,

12

10

% tf

15

it

' ~ "~.e*

11

13

14

16

Plate 2

Figures 1-13. Fissanomocarella paibiensis Peng, Lin, and Chen, 1995 1. Nearly complete, mostly exfoliated cranidium, NIGP 118823, × 5, P82.5. 2. Nearly complete, partly exfoliated cranidium, NIGP 137907, × 3, P82.5. 3. Nearly complete, slightly exfoliated cranidium, holotype, NIGP 118825, × 4, P82.5. 4. Incomplete, slightly exfoliated cranidium, NIGP 118824, × 3, P82.5. 5, 6. Nearly complete, slightly exfoliated cranidium in dorsal and anterolateral views, NIGP 118829, × 5, P82.5. 7. Incomplete, slightly exfoliated cranidium, NIGP 118822, × 5, P82.5. 8. Nearly complete, slightly exfoliated pygidium, NIGP 118828, × 6, P82.5. 9. Complete, partly exfoliated pygidium, NIGP 118827, × 2, P82.5. 10. Incomplete pygidium in ventral view, showing doublure, NIGP 118830, × 2, P82.5. 11. Broken, partly exfoliated cranidium, NIGP 137908, × 3, P82.5. 12. Nearly complete, partly exfoliated pygidium, NIGP 118830, × 3, P82.5. 13. Complete, partly exfoliated pygidium, NIGP 137909, × 3, P82.5. All from Dorypyge richthofeni Zone.

• 202.

Plate 2

o

o.

r y .....

11

10

12

13

Plate 3

Figures 1-16. Huayuania subcalva Peng, Lin, and Chen, 1995 1. Small testaceous cranidium, NIGP 137910, × 20, P200.7. 2. Small slightly exfoliated cranidium, NIGP 137911, × 25, P200.7. 3. Small slightly exfoliated cranidium, NIGP 137912, × 12, P200.7. 4. Small slightly exfoliated cranidium, NIGP 137913, × 10, P200.7. 5. Small slightly exfoliated cranidium, NIGP 137914, × 10, P200.7. 6. Largely exfoliated cranidium, NIGP 118847, × 10, P200.7. 7. Incomplete, testaceous cranidium, NIGP 118846, × 8, P200.7. 8. Testaceous librigena, NIGP 118852, × 5, P200.7. 9-11. Partly exfoliated cranidium in dorsal, anterolateral, and obliquely anterior views, holotype, NIGP 118848, × 5, P200.7. 12. Exfoliated and partly exfoliated pygidia in association NIGP 137915, x 5, P200.7. 13. Librigena in ventral view, NIGP 137916, × 8, P200.7. 14. Testaceous pygidium, NIGP 118856, × 10, P200.7. 15. Largely exfoliated pygidium, NIGP 137917, × 5, P200.7. 16. Partly exfoliated pygidium, NIGP 118860, × 10, P200.7. All from Pianaspsis sinensis Zone.

• 204-

Plate 3

10

~r

i'

.

S

11 12 13

14

15

16

Plate 4

Figures 1-8. Huayuania subcalva Peng, Lin, and Chen, 1995 1. Testaceous librigena, NIGP 118853, × 10, P200.7. 2. Testaceous cranidium, NIGP 137918, x 12, P200.7. 3. Largely exfoliated cranidium, NIGP 118849, × 8, P200.7. 4. Testaceous librigena, NIGP 137919, × 10, P200.7. 5. Incomplete, testaceous pygidium, NIGP 137920, × 10, P200.7. 6. Exfoliated pygidium, showing doublure, NIGP 118859, x 6, P200.7. 7. Largely exfoliated pygidium, NIGP 118857, × 5, P200.7. 8. Exfoliated pygidium, NIGP 124311, × 4.5, P108. Figures 1-7 from Pianaspsis sinensis Zone; Figure 8 from the top part of Dorypyge richthofeni Zone. Figures 9-17. Sudanomocarina? huananensis sp. nov. 9, 11. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 137921, × 15, W146.2. 10. Nearly complete, testaceous cranidium, NIGP 137922, × 15, P164.2. 12, 13. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 137923, × 12, P134.2. 14. Nearly complete, testaceous cranidium, NIGP 137924, × 10, P171. 15. Partly exfoliated pygidium, NIGP 137925, x 7, P164.2. 16. Incomplete, testaceous cranidium, holotype, NIGP 137926, × 10, P204. 17. Partly exfoliated pygidium, NIGP 137927, × 10, P130.5. All from from Pianaspsis sinensis Zone.

• 206 •

Plate 4

h

:'.,. . . . .

,i r

10

12

11

14

13

i~i~¸ i~ ¸

J.a

r

15

16

17

Plate 5

Figures 1-14. Huayuania quadrilateralis (Resser and Endo, 1937) 1. Nearly complete, testaceous cranidium, NIGP 137928, × 2, P123.6. 2. Small, nearly complete, partly exfoliated cranidium, NIGP 137929, × 10, P45.6. 3. Incomplete, largely exfoliated cranidium, NIGP 137930, x 6, P42.5. 4, 5. Nearly complete, testaceous cranidium in dorsal and lateral views, NIGP 137931, x 6, P42.5. 6. Nearly complete, testaceous cranidium, NIGP 137932, × 5, P45.6. 7. Nearly complete, testaceous cranidium, NIGP 137933, x 5, P45.6. 8. Incomplete, testaceous cranidium, NIGP 137934, x 4, P45.6. 9. Incomplete, slightly exfoliated cranidium, NIGP 137935, × 4, P42.5. 10. Partly exfoliated pygidium, NIGP 137936, x 6, P42.5. 11. Testaceous pygidium, NIGP 137937, x 8, P48.5. 12. Incomplete mostly exfoliated pygidium, NIGP 137938, x 5, P42.5. 13. Slightly exfoliated pygidium, NIGP 137939, × 6, P42.5. 14. Broken, testaceous pygidium, NIGP 137940, × 4, P45.6. Figure 1 from basal part of Pianaspsis sinensis Zone; Figures 2-14 from Dorypyge richthofeni Zone.

• 208.

Plate 5

10

12

13

11

14

Plate

6

Figures 1-16. Paranomocarellafortis Peng, Lin, and Chen, 1995 1. Small, incomplete, testaceous cranidium, NIGP 118833, × 5, P122.4. 2. Incomplete, testaceous cranidium, NIGP 137941, × 6, W56.7. 3. Largely exfoliated cranidium, NIGP 118835, × 4, P122.4. 4. Incomplete, testaceous cranidium, NIGP 137942, × 2, P122.4. 5. Incomplete, testaceous cranidium, NIGP 137943, × 6, P82.1. 6. Incomplete, partly exfoliated cranidium, NIGP 137944, × 4.5, P130.5. 7, 8. Nearly complete, testaceous cranidium, holotype, NIGP 118832, × 3, P122.4. 9. Incomplete, partly exfoliated cranidium, NIGP 137945, × 3, P122.4. 10. Partly exfoliated librigena, latex cast, NIGP 118837, × 3.6, P122.4 11. Small, testaceous pygidium, latex cast, NIGP 137946, × 5, P122.4. 12. Partly exfoliated pygidium, latex cast, NIGP 118842, × 1.5, P82.5. 13. Partly exfoliated pygidium, NIGP 137947, × 3, P 108. 14. Largely exfoliated pygidium, NIGP 137948, × 3, P 131.7 15. Partly exfoliated pygidium, NIGP 118845, × 2, P 122.4. 16. Testaceous pygidium, latex cast, NIGP 118844, × 2, P 122.4. Figures 1, 3, 4, 6-11, 13-16 from Pianaspsis sinensis Zone; Figures 2, 5, 12 from the upper part of Dorypyge richthofeni Zone.

• 210

•

Plate 6 I

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Figure 1. Paranomocarella fortis Peng, Lin, and Chen, 1995 1. Librigena in ventral view, showing diagonally directed connective suture, latex cast, NIGP 118840, x 3, P82.5. From Dorypyge richthofeni Zone. Figures 2-9. Paranomocarella similaris sp. nov. 2. Incomplete, slightly exfoliated cranidium, NIGP 137949, x 3, P 136.3. 3. Nearly complete, slightly exfoliated cranidium, holotype, NIGP 137950, x 3, P130.5. 4. Incomplete, testaceous cranidium, NIGP 137951, x 5, P 134.2. 5. Incomplete, partly exfoliated cranidium, NIGP 118838, x 3, P 126.9. 6. Nearly complete, testaceous pygidium, NIGP 137952, x 4, P130.5. 7. Testaceous pygidium, NIGP 137953, x 2, P130.25. 8. Partly exfoliated pygidium, NIGP 137954, x 5, P 131.7. 9. Slightly crushed, testaceous pygidium, NIGP 137955, x 5, P134.2. All from Pianaspsis sinensis Zone. Figures 10-16. Paranomocarella parallela Yang in Zhou et al., 1977 10. Incomplete, slightly exfoliated cranidium, NIGP 137956, x 4, P82.5. 11. Nearly complete, partly exfoliated cranidium, NIGP 137957, x 6, P68.8. 12. Testaceous pygidium, NIGP 137958, x 5, P82.5. 13. Testaceous pygidium, NIGP 137959, x 6, P82.5. 14. Partly exfoliated pygidium, NIGP 137960, × 8, P82.5. 15. Partly exfoliated pygidium, NIGP 137961, x 2, P112.6. 16. Partly exfoliated pygidium, NIGP 137962, x 4, P38.5. Figures 1-14, 16 from Dorypyge richthofeni Zone; Figure 15 from the base of Pianaspsis sinensis Zone.

• 212

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Figures 1-13. Szeaspis huananensis sp. nov. 1. Testaceous, juvenile cranidium, NIGP 137963, × 12, W56.7. 2. Incomplete, testaceous, juvenile cranidium, NIGP 137964, × 15, W56.7. 3. Nearly complete, testaceous, juvenile cranidium, NIGP 137965, x 12, W56.7. 4. Nearly complete, testaceous, juvenile cranidium, NIGP 137966, × 12, W56.7. 5. Nearly complete, slightly exfoliated cranidium, latex cast, NIGP 137967, × 4, W56.7. 6. Incomplete, testaceous cranidium, NIGP 137968, × 6, W56.7. 7. Nearly complete, largely exfoliated cranidium, latex cast, NIGP 137969, × 4, W56.7. 8. Nearly complete, partly exfoliated cranidium, holotype, NIGP 137970, × 4, W56.7. 9. Incomplete, testaceous hypostome, NIGP 137971, × 8, W56.7. 10. Testaceous, juvenile pygidium, NIGP 137972, × 10, W56.7. 11. Testaceous pygidium, NIGP 137973, × 4, W56.7. 12. Exfoliated pygidium, NIGP 137974, x 8, W56.7. 13. Testaceous pygidium, NIGP 137975, × 8, W56.7. All from the base of Pianaspsis sinensis Zone. Figures 14-16. Paranomocarella? obvia sp. nov. 14. Incomplete, testaceous pygidium, NIGP 137976, x 3, P277 15. Largely exfoliated pygidium, holotype, NIGP 137977, × 5, P277. 16. Incomplete, slightly exfoliated pygidium, NIGP 137978, × 5, P269. All from Wanshania wanshanensis Zone.

• 214-

Plate 8

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Plate 9

Figures 1-7. Adelogonus hunanensis sp. nov. 1-3, 7. Testaceous cranidium in anterior, dorsal, anterolateral views, and partial enlargement of basal glabellar lobe, holotype, NIGP 137979, x 2, × 2, x 2, x 6, P331.8. 4. Most exfoliated pygidium, NIGP 137980, x 2.25, P331.8 5, 6. Testaceous pygidium in dorsal view and partial enlargement of left anterior pleural region, NIGP 137981, x 6, x 18, P286.3. Figures 1--4, 7 from Liostracina bella Zone; Figures 5, 6 from the upper part of Wanshania wanshanensis Zone. Figures 8-11. Iniotoma laevis sp. nov. 8. Small cranidium, NIGP 137982, x 16, W277. 9-11. Incomplete, mostly exfoliated cranidium, holotype, 9, 10, latex cast in anterolateral and dorsal views, NIGP 137983, x 6, W227. All from Liostracina bella Zone. Figures 12-14. Paibiella paibiensis gen. et sp. nov. 12. Small, incomplete cranidium, NIGP 137984, x 20, P249. 13, 14. Incomplete testaceous cranidium in dorsal and anterolateral views, holotype, NIGP 137985 x 12, P240.5. All from Wanshania wanshanensis Zone.

• 216.

Plate 9

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Figure 1. Grandioculus obscurus sp. nov. 1. Incomplete, testaceous cranidium, holotype, NIGP 137986, x 6, P64.5. From Dorypyge richthofeni Zone. Figure 2. Proasaphiscus (Honanaspis)? sp. cf. P. (H.) parvus Kuo in Lu et al., 1965 2. Incomplete, testaceous cranidium, NIGP 137987, x 14, P35.5. From Dorypyge richthofeni Zone. Figures 3, 4. Maotunia sp. cf. M. blackwelderi (Resser and Endo, 1937) 3. Incomplete, partly exfoliated cranidium, NIGP 137988, × 7.5, P7.25. 4. Incomplete, partly exfoliated cranidium, NIGP 137989, × 11, W I.2. All from Dorypyge richthofeni Zone. Figures 5-10. Mapania jiangnanensis sp. nov. 5. Incomplete, probably testaceous cranidium in dorsal and anterolateral views, NIGP 137990, × 11, W146.2. 6. Incomplete, testaceous cranidium, NIGP 137991, x 6, P42.5. 7-9. Anterior part of the incomplete cranidium of fig. 8, latex cast, holotype; 9, cranidium, composite photo from fig. 7 and 8, NIGP 137992, all x 6, P42.5. 10. Testaceous librigena, NIGP 137993, x 6, P42.5. Figure 5 from Pianaspsis sinensis Zone; Figures 6-10 from Dorypyge richthofeni Zone. Figures 11-14. Eymekops? sp. 1 1 1 - 1 4 . Nearly complete, testaceous cranidium in lateral, obliquely anterior, anterolateral and dorsal views, NIGP 137994, × 14, P319.6. From Liostracina bella Zone. Figure 15. Eymekops? sp. 2 15. Broken, testaceous pygidium, NIGP 137995, x 5, P337.5. From Liostracina bella Zone.

• 218-

Plate 10

11 12

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Plate 11

Figures 1-15. Grandioculus truncatus sp. nov. 1. Incomplete, mostly exfoliated cranidium, NIGP 137996, x 12, P35.5. 2. Incomplete, testaceous cranidium, NIGP 137997, x 12, P35.5. 3. Testaceous librigena, NIGP 137998, x 8, P35.5. 4, 11, 12. Nearly complete testaceous cranidium in dorsal and anterolateral, and partial enlargement of fight frontal area to show fine granulation of cranidial surface, holotype, NIGP 137999, × 12, x 10, x 18, P37. 5. Partly exfoliated librigena, NIGP 138000, x 8, P35.5. 6. Incomplete, largely exfoliated cranidium, NIGP 138001, x 8, P37. 7. Mostly exfoliated pygidium, NIGP 138002, x 6, P35.5. 8. Broken, incomplete, exfoliated pygidium, NIGP 138003, x 6, P35.5. 9, 14. Broken, testaceous pygidium in dorsal view and partial enlargement of left pleural field and axis to show granulation of pygidial surface, NIGP 138004, x 12.5, P35.5. 10, 15. Exfoliated pygidium in dorsal and posterolateral views, NIGP 138005, x 6, P35.5. 13. Exfoliated pygidium, NIGP 138006, x 5, P35.5. All from Dorypyge richthofeni Zone.

• 220-

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Figures 1-12. Huayuanaspis granulosa gen. et sp. nov. 1. Testaceous librigena, NIGP 138007, x 8, P37. 2-4. Nearly complete, testaceous cranidium in dorsal, anterior, and anterolateral views, holotype, NIGP 138008, x 6, P37. 5. Incomplete, testaceous cranidium, NIGP 138009, x 8, P37. 6. Incomplete, mostly exfoliated cranidium, NIGP 138010, x 8, P37. 7. Incomplete, testaceous cranidium, NIGP 138011, x 4, P37. 8. Incomplete, testaceous cranidium, latex cast, NIGP 138012, x 5, P38.5. 9. Partly exfoliated librigena, NIGP 138013, x 4, P37. 10. Incomplete, testaceous cranidium, NIGP 138014, x 4, P45.6. 11. Incomplete, testaceous cranidium, NIGP 138015, x 4, P 19.8. 12. Crushed, testaceous librigena, showing anterior doublure, NIGP 138016, x 3.5, P35.5. All from Dorypyge richthofeni Zone.

• 222•

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Plate 13 Figures 1-10. Huayuanaspis perpauca gen. et sp. nov. 1-3. Testaceous cranidium in dorsal, anterolateral, and anterior views, NIGP 138017, × 8, x 8, × 7, P45.6. 4, 5. Mostly exfoliated cranidium in dorsal and anterolateral views, holotype, latex cast, NIGP 138018, x 5.2, P45.6. 6. Incomplete, testaceous cranidium, NIGP 138019, x 8, P45.6. 7. Crushed, incomplete, testaceous cranidium, NIGP 138020, × 4, P45.6. 8. Incomplete, testaceous cranidium, NIGP 138021, x 4, P45.6. 9. Partly exfoliated cranidium, NIGP 138022, × 3.8, P45.6. 10. Broken, partly exfoliated cranidium, NIGP 138023, × 4.8, P45.6. All from Dorypyge richthofeni Zone. Figure 11. Huayuanaspis? sp. 11. Incomplete, testaceous cranidium, NIGP 138024, x 4, P126. From Pianaspsis sinensis Zone.

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Figures 1-10. Zhujia hunanensis Peng, Lin, and Chen, 1995 1. Exfoliated cranidium, NIGP 138025, × 6, P108. 2. Exfoliated cranidium, latex cast, NIGP 118865, x 8, P 108. 3. Incomplete, partly exfoliated cranidium, NIGP 118864, × 4, P 108. 4, 5. Most exfoliated cranidium in dorsal and anterior views, NIGP 118862, x 4, P 108. 6, 7. Partly exfoliated cranidium in lateral and dorsal views, holotype, NIGP 118863, × 4, P108. 8. Mostly exfoliated pygidium, NIGP 138026, × 4, P 108. 9. Mostly exfoliated pygidium, NIGP 138027, x 5, P108. 10. Hypostome, NIGP 138028, × 4, P 108. All from Dorypyge richthofeni Zone. Figure 11. Proasaphiscus? sp. cf. P. butes (Walcott, 1905) 11. Incomplete cranidium, NIGP 138029, × 8, P34.3. From Dorypyge richthofeni Zone. Figures 12-16. Iniotoma porosus sp. nov. 12, 14, 15. Incomplete, testaceous cranidium in dorsal, lateral and partial enlargement of left posterior parts of fixigena and glabella, NIGP 138030, × 9, × 9, × 18, P341.8. 13, 16. Incomplete, testaceous pygidium in dorsa and partial enlargement of left anterior part of pleural region, holotype, NIGP 138031, x 9, x 18, P298.4. Figures 12, 14, 15 from Liostracina bella Zone; Figures 13, 16 from Wanshania wanshanensis Zone.

• 226.

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Figures 1-16. Maotunia rectangularis sp. nov. 1, 2. Small testaceous cranidium and latex cast from its external mold, NIGP 138032, x 11.5, W0. 3. Small, mostly exfoliated cranidium, NIGP 138033, x 12, W0. 4, 5. Small, mostly exfoliated cranidium in dorsal and anterolateral views, NIGP 138034, x 8, W0. 6, 9. Small, mostly exfoliated cranidium in dorsal and anterior views, NIGP 138035, x 8, W0. 7, 8. Slightly exfoliated cranidium in dorsal and anterolateral views, NIGP 138036, both x 6.8, W0. 10. Broken, mostly exfoliated cranidium, NIGP 138037, x 8.8, W0. 11. Partly exfoliated librigena, NIGP 138038, x 7, W0. 12-14. Mostly exfoliated cranidium in anterolateral, dorsal, and oblique anterior views, NIGP 138039, x 7, W0. 15, 16. Incomplete, testaceous cranidium in anterolateral and dorsal views, latex cast, NIGP 138040, x 5.7, W0. All from the unnamed zone.

• 228.

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Figures 1-12. Maotunia rectangularis sp. nov. 1. Incomplete, testaceous cranidium, NIGP 138041, × 4.6, W0. 2-4. Nearly complete, mostly exfoliated cranidium in dorsal, anterolateral, and oblique anterior views, holotype, NIGP 138042, x 4, W0. 5. Small, exfoliated pygidium, latex cast, NIGP 138043, × 9, W0. 6, 7. Partly exfoliated pygidium in dorsal and posterior views, NIGP 138044, × 8, W0. 8. Small, testaceous pygidium, NIGP 138045, × 10, W0. 9. Small, exfoliated pygidium, NIGP 138046, x 10, W0. 10. Mostly exfoliated pygidium, NIGP 138047, × 6, W0. 11. Exfoliated pygidium showing doublure, latex cast, NIGP 138048, × 8.2, W0. 12. Mostly exfoliated pygidium, NIGP 138049, × 4.5, W0. All from the unnamed zone.

• 230-

Plate 16

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Figures 1-17. Sudanomocarina sp. cf. S. changi Jell in Jell and Robison, 1978 1, 2. Testaceous, juvenile cranidium, NIGP 138050, x 13, W0. 3. Incomplete, exfoliated, juvenile cranidium, NIGP 138051, × 8, W 1.2. 4. Incomplete, mostly exfoliated cranidium, latex cast, NIGP 138052, × 7.5, W0. 5. Mostly exfoliated cranidium, NIGP 138053, × 10, W0. 6. Incomplete, testaceous cranidium, NIGP 138054, × 8, W 1.2. 7. Incomplete, partly exfoliated cranidium, NIGP 138055, × 8, W 1.2. 8, 9. Incomplete, mostly exfoliated cranidium, NIGP 138056, x 7, × 7, W0. 10-13. Nearly complete, slightly exfoliated cranidium in anterolateral, dorsal, oblique anterior, and lateral views, NIGP 138057, × 7, W0. 14. Broken, partly exfoliated, pygidium, NIGP 138058, × 10, W1.2. 15. Broken, testaceous pygidium, NIGP 138059, × 14, W0. 16. Testaceous pygidium, NIGP 138060, × 20, P 12.5. 17. Incomplete, partly exfoliated pygidium, latex cast, NIGP 138061, × 7, W0. Figures 1, 2, 4, 5, 8-13, 15, 17 from the unnamed zone; Figures 3, 6, 7, 14, 16 from Dorypyge richthofeni Zone.

• 232-

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Figures 1-9. Sudanomocarina sp. cf. S. changi Jell in Jell and Robison, 1978 1. Incomplete, partly exfoliated cranidium, NIGP 138062, x 8, P33.5. 2. Complete, testaceous cranidium, NIGP 138063, x 6, P33.5. 3. Mostly exfoliated cranidium, NIGP 138064, x 5, P37. 4. Mostly exfoliated librigena, showing portion of doublure and connective suture, NIGP 138065, x 6, P33.5. 5, 7. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 138066, x 8, x 8, P29. 6. Complete, testaceous librigena, NIGP 138067, x 5, P33.5. 8. Nearly complete, testaceous cranidium, NIGP 138068, x 6, P33.5. 9. Partly exfoliated librigena, NIGP 138069, x 6, P33.5. All from Dorypyge richthofeni Zone. Figure 10. Maotunia sp. cf. M. semiplectra Zhang and Jell, 1987 10. Nearly complete, mosrly exfoliated cranidium, NIGP 138070, x 7.5, W0. From the unnamed Zone. Figures 11, 12. Crepicephalina sp. cf. C. quadrata Resser and Endo, 1937 11, 12. Nearly complete, exfoliated cranidium in dorsal and anterolateral views, NIGP 138071, x 6, W1.2. From Dorypyge richthofeni Zone. Figure 13. Maotunia sp. 13. Incomplete pygidium, NIGP 138072, x 6, W3. From Dorypyge richthofeni Zone.

• 234-

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Figures 1-14. Parayujinia constricta gen. et sp. nov. 1. Partly exfoliated librigena, NIGP 138073, × 8, P35.5. 2. Incomplete, partly exfoliated cranidium, NIGP 138074, × 12, P35.5. 3. Incomplete, partly exfoliated cranidium, holotype, NIGP 138075, × 6, P37. 4. Mostly exfoliated librigena, NIGP 138076, × 8, P35.5. 5, 6. Nearly complete, mostly exfoliated cranidium in anterolateral and dorsal views, NIGP 138077, × 8, P35.5. 7. Nearly complete, mostly exfoliated cranidium, NIGP 138078, × 10, P37. 8. Nearly complete, mostly exfoliated pygidium, latex cast, NIGP 138079, × 12, P35.5 9. Incomplete, exfoliated pygidium, NIGP 138080, × 12, P37. 10. Nearly complete, partly exfoliated pygidium, latex cast, NIGP 138081, × 8, P35.5. 11. Incomplete, partly exfoliated pygidium, NIGP 138082, × 13, P12.5. 12. Complete, testaceous pygidium, NIGP 138083, × 5, P35.5. 13, 14. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 138084, × 12, W3. All from Dorypyge richthofeni Zone. Figure 15. Sudanomocarina sp. 15. Incomplete, testaceous cranidium, NIGP 138085, × 10, P 171.5. From Pianaspsis sinensis Zone. Figure 16. Sudanomocarina traynorae Zhang and Jell, 1987 16. Nearly complete, mostly exfoliated cranidium, NIGP 138086, × 5, P48.5. From Dorypyge richthofeni Zone.

• 236.

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Figures 1-14. Buttsia globosa Lu and Lin in Peng, 1987 1, 2. Exfoliated cranidium in dorsal and anterolateral views, NIGP 138087, × 18, W216.5. 3-5. Partly exfoliated cranidium in dorsal, obliquely anterior, and anterolateral views, NIGP 138088, × 15, W223.2. 6. Small, broken pygidium, NIGP 138089, × 22, P282.75. 7. Partly exfoliated cranidium, NIGP 138090, × 14, W221.5. 8, 9. Incomplete, mostly exfoliated cranidium in dorsal and anterolateral views, NIGP 138091, × 10, W219.7. 10, 11. Nearly complete, exfoliated cranidium in dorsal and anterior views, NIGP 138092, × 8, W 215.1. 12, 13. Incomplete, mostly exfoliated cranidium in dorsal and anterolateral views, NIGP 138093, × 8, W197.5. 14. Incomplete pygidium, NIGP 138094, × 15, W216.5. Figures 1, 2, 6, 10-14 from Wanshania wanshanensis Zone; Figures 3-5, 7-9 from Liostracina bella Zone.

• 238.

Plate 20

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Plate 21

Figures 1-3. Buttsia globosa Lu and Lin in Peng, 1987 1. Partly exfoliated cranidium, NIGP 138095, × 8, P307.4. 2. Mostly exfoliated cranidium, NIGP 138096, × 10, W219.7. 3. Incomplete, partly exfoliated cranidium, NIGP 138097, × 12, W227. Figure 1 from Wanshania wanshanensis Zone; Figures 2, 3 from Liostracina bella Zone. Figures 4-6. Paradistazeris hunanensis (Peng, 1987) 4. Cranidium, NIGP 138098, × 10, P298.4. 5. Incomplete, partly exfoliated cranidium, NIGP 138099, × 10, P298.4. 6. Incomplete, mosttly exfoliated cranidium, NIGP 138100, × 10, W215.1. Figure 4, 5 from Liostracina bella Zone; Figure 6 from Wanshania wanshanensis Zone. Figures 7-18. Paradistazeris hubeiensis Zhu in Zhang et al., 1980a 7. Small cranidium, NIGP 138101, × 15, W211.7. 8. Small, partly exfoliated cranidium, NIGP 138102, × 15, W211.7. 9,10. Partly exfoliated cranidium in dorsal and anterolateral views, NIGP 138103, × 9, W199.2. 11, 12. Posterior portion of eye, enlarged from the eye, showing regular placed lenses, × 48, and incomplete librigena with the eye, NIGP 138104, × 12, W 199.2. 13. Incomplete, mostly exfoliated cranidium, NIGP 138105, × 9, W 199.2. 14, 15. Mostly exfoliated cranidium in dorsal and anterolateral views, NIGP 138106, × 10, W199.2. 16. Incomplete librigena with the incomplete eye, showing thick, convex lateral border with terrace lines, NIGP 138107, × 12, W 199.2. 17. Partly exfoliated, incomplete, pygidium, latex cast, NIGP 138108, × 9, W 199.2. 18. Incomplete pygidium, latex cast, NIGP 138109, × 9, W 199.2. All from Wanshania wanshanensis Zone.

• 240 •

Plate 21

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Plate 22 Figures 1-15. Paradistazeris rotundus sp. nov. 1. Small incomplete, partly exfoliated cranidium, NIGP 138110, x 15, W216.5. 2-4. Small, testaceous cranidium in dorsal, anterolateral, and obliquely anterior views, NIGP 138111, x 15, W216.5. 5. Partly exfoliated cranidium, NIGP 138112, x 10, W215.1. 6. Incomplete, partly exfoliated cranidium, NIGP 138113, x 10, W216.5. 7, 8. Slightly broken, testaceous cranidium in dorsal and anterolateral views, NIGP 138114, x 12, W215.1. 9, 14, 15. Partly exfoliated cranidium in dorsal, obliquely anterior, and anterolateral views, holotype, NIGP 138115, × 8, W216.5. 10. Incomplete, mostly exfoliated pygidium, NIGP 138116, × 15, W216.5. 11, 12. Testaceous cranidium in dorsal and anterolateral views, NIGP 138117, × 9, W216.5. 13. Testaceous pygidium, latex cast, NIGP 138118, x 15, W216.5. All from the top part of Wanshania wanshanensis Zone.

• 242.

Plate 22

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11

13

14

12

15

Plate 23

Figures 1-17. Madarocephalus orientalis sp. nov. 1, 2. Small, incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 138119, × 20, W187.8. 3. Small, testaceous cranidium, NIGP 138120, × 25, P277. 4-7. Testaceous cephalon in dorsal, lateral, anterolateral, and anterior views, holotype, NIGP 138121, × 25, P240.5. 8, 9. Testaceous cranidium in dorsal and anterolateral views, showing weak lateral glabellar furrows, NIGP 138122, × 20, P249. 10. Testaceous cranidium, NIGP 138123, × 20, W 146.2. 11-13. Testaceous cranidium in oblique anterior, dorsal, and lateral views, NIGP 138124, × 25, P268.3. 14. Partly exfoliated cranidium, NIGP 138125, × 25, P268.3. 15, 16. Mostly exfoliated cranidium in lateral and dorsal views, NIGP 138126, × 25, P268.3. 17. Testaceous cranidium, NIGP 138127, × 25, P273.8. Figures 1-9, 11-17 from Wanshania wanshanensis Zone; Figure 10 from Pianaspsis sinensis Zone.

• 244.

Plate 23

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13 15

14

16

17

Plate 24

Figures 1-17. Huzhuia paratypica Yang, 1978 1. Small, incomplete, testaceous cranidium, NIGP 138128, × 20, W 146.2. 2. Small, testaceous cranidium, NIGP 138129, × 15, W 146.2. 3. Small, testaceous cranidium, NIGP 138130, × 15, W 146.2. 4. Small, incomplete, partly exfoliated cranidium, NIGP 138131, × 15, W 146.2. 5, 6. Slightly broken, testaceous cranidium in dorsal and lateral views, NIGP 138132, × 15, P184. 7. Slightly broken, slightly exfoliated cranidium, NIGP 138133, × 15, P223.7. 8. Testaceous cranidium, showing fine and dense granules, NIGP 138134, × 15, × 10, P319.8. 9. Librigena, NIGP 138135, × 12, P331.8. 10. Testaceous cranidium, NIGP 138136, × 15, P 171. 11. Slightly broken, testaceous cranidium, NIGP 138137, × 15, P 171.5. 12. Incomplete testaceous cranidium, NIGP 138138, × 15, W 146.2. 13, 14. Slightly exfoliated cranidium in anterolateral and dorsal views, NIGP 138139, × 12, P331.8. 15. Slightly broken, testaceous cranidium, NIGP 138140, × 12, W 146.2. 16. Incomplete testacerous cranidium, NIGP 138141, × 12, P331.8. 17. Broken, testaceous cranidium, NIGP 138142, × 11, W 146.2. Figures 1-7, 10-12, 15, 17 from Pianaspsis sinensis Zone; Figures 8, 9, 13, 14, 16 from Liostracina bella Zone.

• 246 •

Plate 24

10

12

15

11

13

16

14

17

Plate 25

Figures 1-3. Huzhuia paratypica Yang, 1978 1. Slightly broken, testaceous cranidium, NIGP 138143, × 12, P 149.5. 2. Testaceous cranidium, NIGP 138144, × 11, W 146.2. 3. Incomplete, testaceous cranidium, NIGP 138145, × 12, P331.8. Figures 1, 2 from Pianaspsis sinensis Zone; Figure 3 from Liostracina bella Zone. Figures 4-6. Huzhuia latilimbata sp. nov. 4, 5. Testaceous cranidium in dorsal and anterolateral views, NIGP 138146, × 11, W 146.2. 6. Incomplete mostly exfoliated cranidium, holotype, NIGP 138147, × 12, P279.55. Figures 4, 5 from Pianaspsis sinensis Zone; Figure 6 from Wanshania wanshanensis Zone. Figures 7-15. Huzhuia curvata sp. nov. 7. Testaceous cranidium, NIGP 138148, × 20, PI35.1. 8, 9. Testaceous cranidium in dorsal and anterolateral views, holotype, NIGP 138149, × 15,

r'135. . 10, 11. Partly exfoliated cranidium, NIGP 138150, × 15, × 15, P374.9. 12, 13. Testaceous cranidium in anterior and dorsal views, NIGP 138151, × 18, PI35.1. 14. Slightly broken, testaceous cranidium, NIGP 138152, × 15, P374.9. 15. Slightly broken, testaceous cranidium, NIGP 138153, × 15, PJ35.1. Figures 7-9, 12, 13, 15 from Chuangia subquadrangulata Zone; Figures 10, 11, 14 from Wanshania wanshanensis Zone.

• 248 •

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11 12

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Plate 26

Figures 1-7. Placosema bigranulosum sp. nov. 1. Incomplete, testaceous cranidium, NIGP 138154, × 8, P374.9. 2. Nearly complete, testaceous cranidium, NIGP 138155, × 9, P374.9. 3, 4. Testaceous cranidium in lateral and dorsal views, holotype, NIGP 138156, × 7, × 9, P374.9. 5, 6. Partly exfoliated cranidium, in anterior, anterolateral and anterolateral views, NIGP 138157, × 8, P374.9. 7. Testaceous cranidium, NIGP 138158, x 15, P374.9. All from Chuangia subquadrangulata Zone. Figures 8-11. Onchonotellus curvitensus sp. nov. 8-11. Testaceous cranidium in dorsal, anterolateral, lateral, and oblique anterior views, holotype, NIGP 138159, × 7, P317.4. From Liostracina bella Zone.

• 250.

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10

11

Plate 27

Figures 1-15. Fenghuangella laochatianensis laochatianensis Yang in Zhou et al., 1977 1. Testaceous cranidium, NIGP 138160, × 20, P308. 2. Nearly complete, testaceous cranidium, NIGP 138161, × 17, W228.3. 3. Nearly complete, testaceous cranidium, NIGP 138162, × 19, W227. 4. Testaceous cranidium, NIGP 138163, × 15, P337.5. 5. Slightly exfoliated cranidium, NIGP 138164, × 12, W210.5. 6-10. Exfoliated cephalon, NIGP 138165, P331.8.6, in dorsal view, x 16; 7, in lateral view, showing librigena and facial suture, × 20; 8, partial enlargement showing left eye that is subovate in center, and hook-shaped palpebral lobe surrounding the eye on its anterior and left side, × 35; 9, 10, in anterior and anterolateral views, × 12. 11-13. Testaceous cranidium in dorsal, anterior, and anterolateral views, NIGP 138166, × 12, W210.5. 14. Mostly exfoliated cranidium, NIGP 138167, × 12, P331.8. 15. Incomplete pygidium, NIGP 138168, × 10, W227. Figures 1, 5, 11-13 from the top part of Wanshania wanshanensis Zone; Figures 2-4, 6-10, 14, 15 from Liostracina bella Zone. Figure 16. Fenghuangella laochatianensis crassa subsp, nov. 16. Incomplete pygidium, NIGP 138169, × 20, W228.3. From Liostracina bella Zone.

• 252 •

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12

10 13

11

14

15

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Plate 28

Figures 1-15. Fenghuangella laochatianensis crassa subsp, nov. 1. Nearly complete, testaceous cranidium, NIGP 138170, × 20, W218.3. 2. Small, nearly complete, testaceous cranidium, NIGP 138171, × 30, P378.25. 3. Nearly complete, testaceous cranidium, holotype, NIGP 138172, × 20, P378.25. 4. Nearly complete, testaceous cranidium, NIGP 138173, × 20, P378.25. 5-7, 15. Nearly complete cephalon in dorsal, anterior, anterolateral, and lateral vies, NIGP 138174, × 15, P135.1. Note the weak palpebral lobe, facial suture, and probable absence of genal spine. 8. Small incomplete cranidium, latex cast, NIGP 138175, × 25, P378.25. 9. Nearly complete, testaceous cranidium, NIGP 138176, × 20, P378.25. 10, 11. Testaceous cranidium in dorsal and anterolateral views, NIGP 138177, × 19, × 19, W211.7. 12, 13. Testaceous cranidium in anterolateral and dorsal views, NIGP 138178, × 20, W227. 14. Testaceous cranidium, NIGP 138179, × 12, W228.3. Figures 1-7, 12-15 from Liostracina bella Zone; Figures 8, 9 from Chuangia subquadrangulata Zone; Figures 10, 11 from Wanshania wanshanensis Zone.

• 254.

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Plate 29 Figures 1-17. Fenghuangella coniforma Yang in Zhou et al., 1977 1. Broken, testaceous cranidium, NIGP 138180, x 20, W 199.2. 2. Incomplete testaceous cranidium, latex cast, NIGP 138181, x 15, W 188.8. 3. Two cranidia in association, NIGP 138182, x 15, W 199.2. 4, 5. Nearly complete testaceous cranidium in dorsal and anterolateral views, NIGP 138183, × 15, W199.2. 6, 7. Partly exfoliated cranidium in dorsal and anterolateral views, NIGP 138184, × 15, W199.2. 8. Partly exfoliated cranidium, NIGP 138185, × 15, P278. 9. Partly exfoliated cranidium, NIGP 138186, x 15, W 188.8. 10. Partly exfoliated cranidium, NIGP 138187, x 16, W219.7. 11. Nearly complete, testaceous cranidium, NIGP 138188, x 16, W219.7. 12. Incomplete, testaceous cranidium, NIGP 138189, x 12, W 188.8. 13. Incomplete, testaceous cranidium, NIGP 138190, × 12, W 196.3. 14. Incomplete, testaceous cranidium, latex cast, NIGP 138191, x 15, W 196.3. 15. Incomplete, testaceous pygidium, NIGP 138192, x 18, W 199.2. 16. Incomplete, testaceous pygidium, latex cast, NIGP 138193, x 18, W 199.2. 17. Testaceous testaceous pygidium, NIGP 138194, x 14.5, W 197.5. All from Wanshania wanshanensis Zone.

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Figures 1-8. Fenghuangella fusilis sp. nov. 1, 8. Testaceous cranidium in dorsal and anterolateral views, NIGP 138195, × 28, W254.1. 2. Testaceous cranidium with occipital spine broken out, NIGP 138196, × 25, W254.1. 3. Nearly complete, testaceous cranidium, NIGP 138197, × 18, W254.1. 4-7. Testaceous cranidium, holotype, NIGP 138198, × 15, W254.1.4, 5, stereopair in dorsai view; 6, 7, in anterolateral and anterior views. All from Liostracina bella Zone. Figures 9-15. Fenghuangella liostracinala Yang in Zhou et al., 1977 9, 10. Incomplete, testaceous exoskeleton with broken occipital spine, latex cast in dorsal and lateral views, NIGP 138199, x 19, W251.15. 11, 15. Testaceous cranidium with broken occipital spine in dorsal view and in partial enlargement, NIGP 138200, x 18, × 36, P13-2.75. 12, 13. Testaceous cranidium with broken glabella and occipital spine, in anterolateral and dorsal views, NIGP 138201, × 15, W251.15. 14. Testaceous cranidium with broken occipital spine, NIGP 138202, x 22, PI3-2.75. All from Liostracina bella Zone.

• 258 •

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11

13 12

10

14

15

Plate 31

Figures 1-10. Lobocephalina sinensis sp. nov. 1. Mostly exfoliated cranidium, NIGP 138203, × 16, W219.7. 2, 3, 7. Testaceous cranidium in dorsal, anterolateral views and partial enlargement of posterior glabella showing fine punctae, holotype, NIGP 138204, × 10, × 10, × 25, P319.8. 4. Incomplete, largely exfoliated cranidium, dorsal view, NIGP 124312, × 11, W228.3. 5, 6a, 10. Fragment of testaceous cranidium in dorsal and obliquely anterolateral views, and partial enlargement of anterior border showing fine punctae, NIGP 138205, x 10, × 10, x 25, W254.1. 8, 9. Testaceous pygidium, partial enlargement of left pleural region and dorsal view, NIGP 138206, × 27.5, × 11, W254.1. All from Liostracina bella Zone. Figure 6b. Oculishumardia hunania Peng, Babcock, Hughes, and Lin, 2003 6b. The same cranidium as on P1.66, fig. 9, × 10, W254.1. From Liostracina bella Zone. Figures 11-14. Sulcareclava sagitta gen. et sp. nov. 11, 12. Meraspid cranidium in anterolateral and dorsal views, NIGP 138207, x 16, W227. 13. Small cranidium, NIGP 138208, × 15, PI360.3. 14. Cranidium, dorsal view, holotype, NIGP 138209, × 10, PI360.3. Figures 11, 12 from Liostracina bella Zone; Figures 13, 14 from Chuangia subquadrangulata Zone. Figures 15, 16. Adelogonus? sp. 15, 16. Incomplete, partly exfoliated pygidium, NIGP 138210, in dorsal and lateral views, × 4, P341.8 From Liostracina bella Zone.

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10

11

13

12

14

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Plate 32

Figures 1-7. Stigmatoa yangziensis Yang in Zhou et al., 1977 1. Incomplete, exfoliated cranidium, NIGP 138211, × 5, PI323.5. 2, 3. Incomplete, partly exfoliated cranidium, in anterolateral and dorsal views, NIGP 138212, × 4, P378.25. 4, 5. Nearly complete, slightly exfoliated cranidium in anterolateral and dorsal views, NIGP 138213, × 3, PI356.5. 6. Incomplete, mostly exfoliated cranidium, NIGP 138214, × 2.5, PI360.3. 7. Incomplete, mostly exfoliated cranidium, NIGP 138215, × 3, P1356.5. All from Chuangia subquadrangulata Zone. Figures 8, 9. Paradistazeris hubeiensis Zhu in Zhang et al., 1980 8, 9. Very slightly exfoliated pygidium in dorsal and posterolateral views, NIGP 138216, × 10, W201. From Wanshania wanshanensis Zone.

• 262 •

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7

8

9

Plate 33 Figures 1-14. Kingstonia euryaxis sp. nov. 1. Small, incomplete, testaceous cranidium, NIGP 138217, × 18, P3.2. 2. Small, incomplete, testaceous cranidium, NIGP 138218, × 18, P3.2. 3. Small, testaceous, crushed cranidium, NIGP 138219, × 20, P3.2. 4-6. Testaceous cranidium in anterior, anterolateral, and dorsal views, holotype, NIGP 138220, × 12.5, P6. 7. Testaceous cranidium, NIGP 138221, × 12, P3.2 8. Incomplete, testaceous cranidium, NIGP 138222, × 12, P3.2. 9. Small, testaceous pygidium, NIGP 138223, × 16, P7.25. 10. Testaceous, slightly crushed cranidium, NIGP 138224, × 12, P3.2. 11. Testaceous pygidium, NIGP 138225, × 15, P3.2. 12. Small, testaceous pygidium, NIGP 138226, × 17, P7.25. 13. Testaceous pygidium, NIGP 138227, × 15, P3.2. 14. Testaceous pygidium, NIGP 138228, × 12. P3.2. All from the basal part of Dorypyge richthofeni Zone.

• 264.

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Figures 1-16. Lisania paibiensis sp. nov. 1. Small, testaceous cranidium, NIGP 138229, × 20, W 132.5. 2. Small, testaceous cranidium, NIGP 138230, × 20, W 132.5. 3. Two small, testaceous cranidia in association, NIGP 138231, × 12, W 132.5 4. Small, testaceous, nearly complete cranidium, NIGP 138232, × 15, W 132.5. 5. Small, testaceous, nearly complete cranidium, NIGP 138233, × 12, W 146.2. 6. Nearly complete, testaceous cranidium, NIGP 138234, × 11, W 132.5. 7. Broken, testaceous cephalon, NIGP 138235, × 10, P 180.3. 8. Small, testaceous cranidium, NIGP 138236, × 8, P164.2. 9. Partly exfoliated cranidium, NIGP 138237, × 8, W 132.5. 10. Testaceous, nearly complete cranidium, NIGP 138238, × 8, W 132.5. 11. Mostly exfoliated cranidium, NIGP 138239, × 8, P 164.2. 1 2 - 1 4 . Testaceous cranidium in anterolateral, dorsal, and lateral views, NIGP 138240, × 7, P164.2. 15. Testaceous, incomplete cranidium, NIGP 138241, × 11, W 132.5. 16. Testaceous, incomplete cranidium, NIGP 138242, × 8, P180.3. All from Pianaspsis sinensis Zone.

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11 12

13

15

16

14

Plate 35

Figures 1-11. Lisania paibiensis sp. nov. 1. Testaceous, nearly complete cranidium, holotype, NIGP 138243, × 5, P180.3. 2. Slightly exfoliated cranidium, NIGP 138244, × 4, P 180.3. 3. Mostly exfoliated cranidium, NIGP 138245, × 4, P164.2. 4. Small, incomplete, testaceous cranidium, NIGP 138246, × 7, P 180.3. 5. Partly exfoliated hypostome, NIGP 138247, × 15, W 132.5. 6. Small, testaceous cranidium and pygidium in association, NIGP 138248, × 8, W 132.5. 7. Testaceous pygidium, NIGP 138249, × 8, W 132.5. 8. Testaceous, broken pygidium, NIGP 138250, × 7, W 132.5. 9. Slightly exfoliated hypostome, NIGP 138251, × 6, P 171. 10. Librigena in ventral view, showing doublure, NIGP 138252, × 6, W 132.5. 11. Testaceous pygidium, NIGP 138253, × 12, W139.2. All from Pianaspsis sinensis Zone. Figures 12-16. Lisania yuanjiangensis (Yang in Zhou et al., 1977) 12. Partly exfoliated cranidium and broken pygidium in association, NIGP 138254, x 6, P126.9. 13, 14. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 138255, × 5, W111.3. 15. Exfoliated pygidium, NIGP 138256, × 6, W 111.3. 16. Exfoliated, incomplete pygidium, NIGP 138257, × 6, P130.5. All from the lower part of Pianaspsis sinensis Zone.

• 268•

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14

Plate 36 Figures 1-18. Lisania yuanjiangensis (Yang in Zhou et al., 1977) 1. Small, testaceous cranidium, NIGP 138258, × 15, P130.5. 2. Nearly complete, partly exfoliated cranidium, NIGP 138259, × 4, P156. 3. Nearly complete, testaceous cranidium, NIGP 138260, × 10, P 138.8. 4, 5. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 138261, × 6, × 6, P130.5. 6, 7. Nearly complete, mostly exfoliated cranidium in dorsal and lateral views, NIGP 138262, × 4, × 4, P131.7. 8. Testaceous librigena, NIGP 138263, × 5, P134.2. 9. Nearly complete, testaceous cranidium, NIGP 138264, × 7, P134.2. 10. Slightly exfoliated librigena, NIGP 138265, × 5, P 134.2. 11. Nearly complete, exfoliated cranidium, NIGP 138266, × 5, W 111.3. 12, 15. Nearly complete, testaceous cranidium in dorsal and lateral views, NIGP 138267, x 5.5, × 5.5, P 130.5. 13. Testaceous pygidium, NIGP 138268, × 6, P136.3. 14, 17. Testaceous pygidium, NIGP 138269, × 5, P 136.3. 16. Testaceous pygidium, NIGP 138270, × 6, P 131.7. 18. Partly exfoliated pygidium, NIGP 138271, × 7, P 134.2. All from the lower part of Pianaspsis sinensis Zone.

• 270-

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11 13

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15 14

16

17

18

Plate 37

Figures 1-3. Lisania yuanjiangensis (Yang in Zhou et al., 1977) 1. Two cranidia in association, NIGP 138272, x 7, P138.8. 2. Nearly complete, testaceous cranidium, NIGP 138273, x 4.5, P 121.48. 3. Testaceous pygidium, NIGP 138274, x 8, P136.7. All from the lower part of Pianaspsis sinensis Zone. Figure 4. Lisania? sp. 4. Nearly complete, testaceous pygidium, NIGP 138275, x 14, W 146.2. From Pianaspsis sinensis Zone. Figures 5-10. Lisania paratungjenensis Yang in Zhou et al., 1977 5, 6. Testaceous pygidium in dorsal and posterolateral views, NIGP 138276, × 6.5, P204. 7, 9. Incomplete exfoliated cephalon, NIGP 138277, × 5, P200.7. Fig. 9 is a composite photograph of the same cranidium with the left side being a mirror image of the fight side. 8. Incomplete, testaceous cranidium, NIGP 138278, × 8, P125. 10. Nearly complete, testaceous cranidium, NIGP 138279, × 8, P204. All from the upper part of Pianaspsis sinensis Zone. Figures 11-13. Lisania sp. cf. L. agonius (Walcott, 1905) 11. Incomplete, testaceous cranidium, NIGP 138280, × 7, P108. 12. Testaceous pygidium, NIGP 138281, × 6, P108. 13. Incomplete, slightly exfoliated cranidium, NIGP 138282, × 4, P108. All from the upper part of Dorypyge richthofeni Zone. Figure 14. Lisania wangcunensis sp. nov. 14. Nearly complete, testaceous cranidium, holotype, NIGP 138283, × 8, W 139.2. From the upper part of Pianaspsis sinensis Zone. Figure 15. Qiandongaspis sp. 15. Broken, mostly exfoliated cranidium, NIGP 138284, x 4, W 196.3. From Wanshania wanshanensis Zone.

• 272.

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Plate 38 Figures 1-9. Baojingia youshuiensis Yang in Zhou et al., 1977; all from P249. 1. Incomplete, partly exfoliated cranidium, NIGP 138285, x 8. 2, 3, 6. Testaceous cranidium in dorsal and anterolateral views and enlargement of left palpebral area, NIGP 138286, x 4, x 4, x 8. 4. Exfoliated librigena, NIGP 138287, x 4. 5. Testaceous cranidium, NIGP 138288, x 8. 7. Partly exfoliated pygidium, NIGP 138289, x 3. 8. Partly exfoliated pygidium, NIGP 138290, x 5. 9. Broken, testaceous cranidium, NIGP 138291, x 4.5. From the basal part of Wanshania wanshanensis Zone. Figures 10-13. Baojingia quadrata (Yang in Zhou et al., 1977) 10, 11. Broken, partly exfoliated cranidium in do rsal and anterolateral views, NIGP 138292, × 4.5, W173.8. 12. Exfoliated pygidium, NIGP 138293, x 6, W 196.3. 13. Testaceous cranidium, NIGP 138294, x 4, P261.35. From the lower part of Wanshania wanshanensis Zone.

• 274-

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10

11

12

13

Plate 39

Figures 1-7. Baojingia latilimbata (Peng, 1987) 1, 2. Partly exfoliated cranidium and its restored image, white strain ellipse in the lower left of the restored image has its long axis in the direction of compression, NIGP 138295, × 5, W121.6. 3, 4. Testaceous pygidium in dorsal and anterolateral views, NIGP 138296, × 4.5, W121.6. 5, 6. Exfoliated cranidium in lateral and dorsal views, NIGP 138297, x 4, x 4, P 178.3. 7. Partly exfoliated cranidium, NIGP 138298, x 4, P200.7. All from Pianaspsis sinensis Zone. Figures 8, 9. Baojingia jiudiantangensis (Yang in Zhou et al., 1977) 8, 9. Testaceous cranidium and partial enlargement to show the course of posterior branch of facial suture, NIGP 138299, x 4, × 6, P269. From Wanshania wanshanensis Zone. Figures 10-14. Baojingia tungjenensis (Nan in Egorova et al., 1963) 10. Testaceous cranidium, NIGP 138300, × 6, P204. 11. Testaceous cranidium, NIGP 138301, × 3, P204. 12. Partly exfoliated cranidium, NIGP 138302, × 8, P126. 13. Testaceous cranidium, NIGP 138303, × 5, P190.7. 14. Testaceous pygidium, NIGP 138304, × 8, P204. All from Pianaspsis sinensis Zone.

• 276.

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12

13

14

Plate 40 Figures 1-18. Baojingia paralala (Yang in Zhou et al., 1977) 1. Testaceous cranidium, NIGP 138305, × 6, W 146.2. 2. Exfoliated cranidium, NIGP 138306, × 4, P216. 3, 4. Exfoliated cranidium in dorsal and anterior views, NIGP 138307, × 5, P216. 5, 6. Partly exfoliated cranidium in anterolateral and dorsal views, NIGP 138308, × 4, P223.7. 7. Exfoliated cranidium, NIGP 138309, × 4, P216. 8. Testaceous 1: brigena, NIGP 138311, × 8, W146.2. 9. Testaceous pygidium, NIGP 138310, × 6, P223.7. 10. Testaceous pygidium, NIGP 138312, × 6, P223.7. 11, 12. Testaceous librigena in dorsal and anterior views, NIGP 138313, × 5, P223.7. 13. Testaceous librigena, NIGP 138314, × 4, P204. 14. Testaceous pygidium, NIGP 138315, × 6, P223.7. 15. Testaceous pygidium, NIGP 138316, × 6, P223.7. 16, 17. Small testaceous pygidium in anterolateral and dorsal views, NIGP 138317, × 11, W146.2. 18. Testaceous librigena, NIGP 138417, × 4, P223.7. All from the upper part of Pianaspsis sinensis Zone.

• 278.

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Plate 41 Figures 1-12. Baojingia subquadrata (Yang, 1978) 1. Testaceous cranidium, NIGP 138318, x 8, P301. 2. Partly exfoliated cranidium, NIGP 138319, x 7, W210.5. 3. Broken, testaceous cranidium, NIGP 138320, x 5, P301. 4. Exfoliated cranidium, NIGP 138321, x 4, P301.9. 5. Partly exfoliated cranidium, NIGP 138322, x 6, P277. 6. Partly exfoliated cranidium, NIGP 138323, x 8, W211.7. 7. Testaceous librigena, NIGP 138324, x 6, W216.5. 8-10. Broken, testaceous cranidium in dorsal and anterolateral views, and partially enlargement, NIGP 138325, x 4, x 4, x 8, W 199.2. 11, 12. Exfoliated cephalon with two thoracic segments attached, dorsal, anterior views, NIGP 138326, x 4, W212.45. All from Wanshania wanshanensis Zone.

• 280-

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Figures 1-14. Baojingia subquadrata (Yang, 1978) 1, 2. Testaceous cranidium in anterolateral and dorsal views, showing malformed left posterior area of fixigena, NIGP 138327, x 4, P227. 3. Broken, partly exfoliated cranidium, showing the absence of occipital spine, NIGP 138328, x 4, W210.5. 4. Partly exfoliated librigena, NIGP 138329, x 5, W210.5. 5, 8. Partly exfoliated cranidium and partial enlargement, NIGP 138330, x 3, x 4.5, P331.8. 6. Testaceous librigena, NIGP 138331, x 6, P301. 7. Testaceous cranidium, NIGP 138332, x 8, W212.45. 9. Testaceous librigena, NIGP 138333, x 8, W216.5. 10. Partly exfoliated pygidium, NIGP 138334, x 5, P301. 11, 12. Partly exfoliated pygidium in dorsal and posterolateral views, NIGP 138335, x 4, P298.4. 13, 14. Partly exfoliated pygidium in dorsal and posterolateral views, NIGP 138336, x 6, W216.5. Figures 1, 2 from the upper part of Pianaspsis sinensis Zone; Figures 3-14 from Wanshania wanshanensis Zone.

• 282-

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Figures 1-13. Neoanomocarella asiatica Hsiang in Egorova et al., 1963 1. Small, testaceous cranidium, NIGP 138337, x 10, W212.45. 2. Small, slightly exfoliated cranidium, NIGP 138338, x 12, W201.5. 3. Small, incomplete, testaceous cranidium, NIGP 138339, x 9, W218.3. 4. Incomplete, partly exfoliated cranidium, NIGP 138340, x 5, W221.5. 5, 8. Incomplete, mostly exfoliated cranidium in anterolateral and dorsal views, NIGP 138341, x 4.5, W227. 6. Mostly exfoliated cranidium, NIGP 138342, x 4, W221.5. 7. Nearly complete, testaceous cranidium, NIGP 138343, x 4.5, W221.5. 9. Nearly complete, testaceous cranidium, NIGP 138344, x 4, W221.5. 10, 13. Nearly complete, partly exfoliated cranidium in lateral and dorsal views, NIGP 138345, x 3, P261.35. 11, 12. Incomplete, partly exfoliated cranidium in anterolateral and dorsal views, NIGP 138346, x 3, W221.5. All from Liostracina bella Zone.

• 284.

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11

12

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Plate 44

Figures 1-18. Neoanomocarella asiatica Hsiang in Egorova et al., 1963 1. Incomplete, testaceous librigena, NIGP 138347, × 3, W227. 2. Nearly complete, mostly exfoliated cranidium, NIGP 138348, × 4, W 199.2. 3. Nearly complete, mostly exfoliated cranidium, NIGP 138349, × 4, W 199.2. 4. Incomplete, testaceous librigena, NIGP 138350, × 3, W221.5. 5, 9. Mostly exfoliated cranidium in dorsal and anterolateral views, NIGP 138351, × 2.5, W221. 6. Partly exfoliated cranidium, NIGP 138352, × 2.5, W215.1. 7. Partly exfoliated cranidium, NIGP 138353, × 4, P261.35. 8. Nearly complete, partly exfoliated cranidium, NIGP 138354, × 2.5, P298.4. 10. Small, incomplete, testaceous pygidium, NIGP 138355, × 15, W225. 11. Small, complete, testaceous pygidium, NIGP 138356, × 10, W221.5. 12, 13. Incomplete, testaceous pygidium in posterolateral and dorsal views, NIGP 138357, × 7, × 7, W221.5. 14. Testaceous pygidium, NIGP 138358, × 5, P331.8. 15. Testaceous pygidium, NIGP 138359, × 6, P298.4. 16, 17. Incomplete, partly exfoliated pygidium in dorsal and posterolateral views, latex cast, NIGP 138360, × 4.5, × 4.5, W227. 18. Nearly complete, partly exfoliated pygidium, NIGP 138361, × 3, W225. Figures 1, 4, 5, 9-14, 16-18 from Liostracina bella Zone; Figures 2, 3, 6-8, 15 from Wanshania wanshanensis Zone.

• 286.

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18

Plate 45

Figures 1-9. Neoanomocarella incilis sp. nov. 1. Small, testaceous cranidium, NIGP 138362, x 7, W 188.8. 2--4. Mostly exfoliated cranidium in anterolateral view, partial enlargement (occipital ring), and dorsal view, NIGP 138363, × 5, × 15, × 5, holotype, W 188.8. 5, 6. Mostly exfoliated cranidium and the latex cast from its external mold, NIGP 138364, × 5, W188.8. 7-9. Nearly complete, testaceous cranidium in anterolateral and dorsal and views and in partial enlargement (posterior portion of glabella), NIGP 138365, x 8, x 8, x 20, W 187.8. All from the lower part of Wanshania wanshanensis Zone. Figures 10-12. Townleyella rara Yuan and Yin, 1998 10. Juvenile cranidium, NIGP 138366, x 30, W228.3. 11, 12. Broken cranidium in anterolateral and dorsal views, NIGP 138367, x 20, W228.3. From Liostracina bella Zone. Figures 13, 14. Undetermined pygidium 1 13. Testaceous pygidium, NIGP 138368, × 8, P261.5. 14. Slightly exfoliated pygidium, NIGP 138369, x 12, P282.6. From Wanshania wanshanensis Zone. Figures 15, 16. Paranomocarella sp. 15. Slightly exfoliated pygidium, NIGP 138370, x 5, P82.5 16. Testaceous pygidium, latex cast, NIGP 138371, x 1.5, P82.5 From Dorypyge richthofeni Zone.

• 288.

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10

11

13

12

14

15

16

Plate 46

Figures 1-13. Qiandongaspis sinensis (Peng, 1987) 1. Incomplete, testaceous cranidium, NIGP 138372, × 10, W215.1. 2. Nearly complete, partly exfoliated cranidium, NIGP 138373, × 6, P298.4. 3. Nearly complete, testaceous cranidium, NIGP 138374, × 18, P260. 4. Testaceous cranidium, NIGP 138375, × 10, P261.5. 5. Broken, mostly exfoliated cranidium, NIGP 138376, × 6, P298.4. 6, 7. Incomplete, exfoliated cranidium in dorsal and lateral views, NIGP 138377, x 7, P135. 8, 9. Nearly complete, testaceous cranidium in anterolateral and dorsal views, NIGP 138378, × 6, W196.3. 10. Incomplete, testaceous cranidium, latex cast, NIGP 138379, × 6, W 197.55. 11. Incomplete, testaceous cranidium, NIGP 138380, × 8, P268.3. 12. Mostly exfoliated cranidium, NIGP 138381, × 7, P 108. 13. Broken, mostly exfoliated cranidium, NIGP 138382, × 8, P108. Figures 1-11 from Wanshania wanshanensis Zone; Figures 12, 13 from Dorypyge richthofeni Zone.

• 290 •

Plate 46

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Figures 1-4. Qiandongaspis sinensis (Peng, 1987) 1, 2. Nearly complete, slightly exfoliated cranidium in dorsal and lateral views, NIGP 138383, × 5, P204. 3. Testaceous cranidium, NIGP 138384, × 10, P269. 4. Incomplete, Testaceous cranidium, NIGP 138385, x 8, P268.3. All from Wanshania wanshanensis Zone. Figures 5-9. Qiandongaspis xiangxiensis sp. nov. 5. Small, incomplete, testaceous cranidium, NIGP 138386, x 12, P 149.5. 6. Incomplete, testaceous cranidium, NIGP 138387, × 12, P 130.5. 7. Nearly commplete, testaceous cranidium, NIGP 138388, × 10, P 123.6. 8, 9. Incomplete, testaceous cranidium in anterolateral and dorsal views, holotype, NIGP 138389, × 6, P167. All from lower part of Pianaspsis sinensis Zone. Figures 10-17. Qiandongaspis convexa sp. nov. 10, 15. Incomplete, testaceous cranidium in anterolateral and dorsal views, holotype, NIGP 138390, × 10, × 10, P123.6. 11. Small, fragmental, testaceous cranidium, NIGP 138391, × 12, P 149.5. 12. Small, incomplete, testaceous cranidium, NIGP 138392, × 12, P123.6. 13. Small, nearly complete, testaceous cranidium, NIGP 138393, x 12, P123.6 14. Small, nearly complete, testaceous cranidium, NIGP 138394, × 12, P123.6 16. Testaceous cranidium, NIGP 138395, x 10, P 123.6. 17. Incomplete, testaceous cranidium, NIGP 138396, × 8, P 123.6. All from the lower part of Pianaspsis sinensis Zone.

• 292-

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Figures 1-10. Shengia quadrata Hsiang in Egorova et al., 1963 1. Small testaceous cranidium, NIGP 138397, x 10, PJ372. 2. Nearly complete, partly exfoliated cranidium, NIGP 138398, x 4, P~72. 3. Three partly or mostly exfoliated cranidial and one exfoliated librigena in association, NIGP 138399, x 3.5, PJ372. 4. Early meraspid, incomplete, testaceous pygidium, latex cast, NIGP 138400, x 25, PJ372. 5. Mostly exfoliated cranidium, NIGP 138401, x 4, P[372. 6. Incomplete, testaceous pygidium, latex cast, NIGP 138402, x 11, PJ372. 7. Nearly complete, testaceous pygidium, NIGP 138403, x 4, P~72. 8. Incomplete, partly exfoliated pygidium, latex cast, NIGP 138404, x 6, P[372. 9. Nearly complete, slightly exfoliated pygidium, NIGP 138405, x 8, P~72. 10. Broken, partly exfoliated pygidium, NIGP 138406, x 3, P[372. All from the basal part of Shengia quadrata Zone.

• 294.

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9

10

Plate 49

Figures 1-10. Shengia trapezia Peng, 1992 1. Incomplete testaceous cranidium, NIGP 138407, × 2.25, P[360.3. 2. Nearly complete, crushed testaceous cranidium with four thoracic segments, latex cast, NIGP 138408, × 3.5, PI335.4. 3. Mostly exfoliated librigena, NIGP 138409, × 4, PI322.4. 4. Early meraspid, incomplete, testaceous pygidium, latex cast, NIGP 138410, × 4, PI371.2 5. Small, slightly exfoliated cranidium, NIGP 138411, × 12, P1322.4. 6. Incomplete, mostly exfoliated cranidium, NIGP 138412, × 3.5, PI335.4. 7. Small, incomplete, mostly exfoliated pygidium, NIGP 138413, × 7, PI370.3. 8. Incomplete, partly exfoliated cranidium, NIGP 138414, × 6, PI371.2. 9. Incomplete, partly exfoliated pygidium, latex cast, NIGP 138415, × 4, PI335.4. 10. Nearly complete, partly exfoliated pygidium, NIGP 138416, × 4.5, PI322.4. All from ChuangiasubquadrangulataZone.

• 296 •

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9

10

Plate 50

Figures 1-16. Shengia wannanensis Qiu in Qiu et al., 1983 1. Small slightly exfoliated cranidium, NIGP 138418, × 12, P378.25 2, 9. Nearly complete, slightly exfoliated cranidium in dorsal and anterolateral views, NIGP 138419, × 2.8, × 2.8, P378.25. 3. Incomplete, mostly exfoliated librigena with posterior portion broken, NIGP 138420, x 4, P378.25. 4, 5. Incomplete, partly exfoliated cranidium, dorsal view and partially enlargement, showing granulate ornamentation, NIGP 138421, x 4.5, x 12, P[323.5. 6. Incomplete partly exfoliated librigena with anterior portion broken, NIGP 138422, x 4, P378.25. 7. Incomplete, mostly exfoliated cranidium, NIGP 138423, x 5.5, P[35.1. 8. Small, testaceous pygidium, latex cast, NIGP 138424, x 6.5, P387.25. 10, 15. Partly exfoliated pygidium in partial enlargement showing granulate ornamentation and dorsal view, NIGP 138425, x 10, × 4.2, P387.25. 11-13. Partly exfoliated pygidium in dorsal, obliquely posterior, and posterolateral views, NIGP 138426, × 3, P387.25. 14. Incomplete, testaceous pygidium, NIGP 138427, x 8, P]322.4. 16. Incomplete, partly exfoliated pygidium, NIGP 138428, x 4, P378.25. All from Chuangia subquadrangulata Zone.

• 298-

Plate 50

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Figures 1-17. Neoglaphyraspis nitida Yuan and Yin, 1998 1. Testaceous cranidium, latex cast, NIGP 138429, x 15, P277. 2. Incomplete, testaceous cranidium, NIGP 138430, x 15, W210.5. 3, 4. Incomplete, testaceous cranidium in anterolateral and dorsal views, NIGP 138431, x 15, W199.2. 5. Nearly complete, testaceous, cranidium, NIGP 138432, × 15, P277. 6, 7. Testaceous cranidium in anterolateral and dorsal views, NIGP 138433, x 15, P277. 8. Broken, slighly exfoliated cranidium, NIGP 138434, x 15, W219.7. 9. Broken, testaceous, cranidium, NIGP 138435, x 15, W225. 10, 11. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 138436, × 10, P298.4. 12, 13. Testaceous librigena in lateral and dorsal views, NIGP 138437, × 11, W254.1. 14. Broken, testaceous pygidium, NIGP 138438, x 15, W210.5. 15. Broken, testaceous pygidium, NIGP 138439, x 16, W211.7. 16, 17. Testaceous pygidium in dorsal and posterolateral views, NIGP 138440, x 15, W227. Figures 1-7, 10, 11, 14, 15 from Wanshania wanshanensis Zone; Figures 8, 9, 12, 13, 16, 17 from Liostracina bella Zone.

• 300 •

Plate 51

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Figures 1-15. Prodamesella punctata Ergaliev, 1980 1. Small, testaceous cranidium, NIGP 132807, × 20, W211.7. 2. Small, incomplete, testaceous cranidium, NIGP 132808, × 20, P249. 3. Small, broken, testaceous cranidium, NIGP 138441, × 20, W211.7. 4. Small, testaceous cranidium, NIGP 138442, x 20, W211.7. 5. Small, broken, testaceous cranidium, NIGP 138443, × 18, W228.3. 6, 7. Small, testaceous cranidium in dorsal and anterior views, NIGP 138444, × 16, W228.3. 8. Small, partly exfoliated cranidium, NIGP 138445, × 20, P337.5. 9. Small, incomplete, testaceous cranidium, NIGP 132809, x 20, P277. 10. Small, broken, testaceous cranidium, NIGP 132810, × 15, P319.6. 11. Broken, partly exfoliated cranidium, NIGP 138446, × 13, W228.3. 12, 15. Nearly complete, mostly exfoliated cranidium in dorsal and anterolateral views, NIGP 138447, × 12, W196.3. 13, 14. Somewhat enrolled thorax beating 8 thoracic segments and pygidium, NIGP 138448, × 10, P319.6. Figures 1, 5-8, 10-15 from Wanshania wanshanensis Zone; Figures 2-4, 9 from Liostracina bella Zone.

• 302-

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10 11

13

14

12

15

Plate 53

Figures 1-14. Prodamesella punctata Ergaliev, 1980 1, 2. Testaceous cranidium in dorsal and anterolateral views, NIGP 138449, x 15, P319.6. 3. Testaceous cranidium, NIGP 132812, x 12, P319.6. 4. Incomplete, testaceous cranidium, NIGP 132811, x 12, P277. 5. Testaceous cranidium, NIGP 132813, x 12, P319.6. 6. Nearly complete, testaceous cranidium, NIGP 132817, x 10, P277. 7. Testaceous librigena, NIGP 132816, x 15, P319.6. 8. Mostly exfoliated cranidium, NIGP 132814, x 12, P319.6. 9, 10a. Mostly exfoliated cranidium in dorsal and anterior views, NIGP 132815, in association with Baikadamaspsis linearis sp. nov. x 10, P319.6. 11. Testaceous pygidium, NIGP 132821, x 15, P319.6. 12a. Incomplete, testaceous cranidium, NIGP 132818, in associated with Aspidagnostus laevis Palmer (12b, original of Peng and Robison, 2000, fig. 30.5), latex cast, x 10, P319.6. 13, 14. Incomplete, testaceous thorax and pygydium in posterolateral and dorsal views, NIGP 132819, x 18, P319.6. Figures 1-3, 5, 7-14 Liostracina bella Zone; Figures 4, 6 from Wanshania wanshanensis Zone. Figure 10b. Baikadamaspis linearis sp. nov. 10b. Cranidium in anterolateral view, x 10, P319.6 (same as P1.71, fig. 5).

• 304 •

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10

12

11

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14

Plate 54

Figures 1-4. Prodamesella tumidula sp. nov. 1. Small, testaceous cranidium, NIGP 138450, × 16, W56.7. 2. Partly exfoliated cranidium, NIGP 138451, × 16, P 123.6. 3. Partly exfoliated cranidium, holotype, NIGP 138452, × 16, P123.6. 4. Incomplete, testaceous cranidium, NIGP 138453, x 16, P 123.6. Figure 1 from Dorypyge richthofeni Zone; Figures 2--4 from Pianaspsis sinensis Zone. Figures 5-16. Prodamesella sp. cf. P. biserrata Jell in Jell and Robison, 1978 5. Small, testaceous cranidium, NIGP 138454, x 20, P204. 6. Testaceous cranidium, NIGP 138455, × 20, P204 (collection level corrected; originally reported as P146.2 in Peng et al., 2001, pl. 5, fig. 15). 7. Incomplete, exfoliated cranidium, NIGP 138456, × 20, P200.7. 8. Incomplete, exfoliated cranidium, NIGP 138457, x 16, W146.2. 9, 10. Incomplete, slightly exfoliated cranidium in dorsal and anterolateral views, NIGP 138458, × 17, W146.2. 11. Incomplete, testaceous cranidium, NIGP 138459, × 16, P319.6. 12. Incomplete, testaceous cranidium, NIGP 138460, × 16, W 146.2. 13. Incomplete, testaceous cranidium, NIGP 138461, × 16, P319.6. 14. Incomplete, slightly exfoliated cranidium, NIGP 138462, × 15, W 146.2. 15, 16. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 138463, × 9, W171.9. Figures 5-10, 12, 14 from Pianaspsis sinensis Zone; Figures 11, 13 from Liostracina bella Zone; Figures 15, 16 from Wanshania wanshanensis Zone.

• 306 •

Plate 54

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Plate 55

Figures 1-15. Luyanhaoaspis decorosa (Peng, Lin, and Chen, 1995) 1-3. Testaceous cranidium in dorsal, oblique anterior, and lateral views, holotype, NIGP 118867, × 4, P277 (HP27-3). 4. Small, slightly broken, testaceous cranidium, NIGP 138464, × 11, P277. 5. Nearly complete, testaceous cranidium, NIGP 118872, x 4, P277. 6. Testaceous cranidium, NIGP 138465, × 8, P277. 7. Testaceous cranidium, NIGP 118868, × 8, P277. 8. Incomplete, testaceous pygidium, NIGP 138466, × 3, P277. 9. Testaceous pygidium, NIGP 138467, × 6, P363.5 10. Testaceous pygidium, NIGP 138468, × 6, P298.4. 11. Partly exfoliated pygidium, NIGP 138469, × 6, P325.7. 12. Testaceous pygidium, NIGP 138470, × 4, P301.9. 13, 14. Partly exfoliated pygidium, NIGP 138471, × 2.5, P298.4. 15. Testaceous pygidium, NIGP 138472, x 4, P346.7. Figures 1-8, 10, 12-14 from Wanshania wanshanensis Zone; Figures 9, 11, 15 from Liostracina bella Zone.

• 308.

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15

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Plate 56

Figures 1-9. Luyanhaoaspis inflata sp. nov. 1, 4, 5. Broken, testaceous cranidium in lateral, dorsal, and anterior views, NIGP 138473, × 4, P326.9. 2, 3. Small, testaceous cranidium in lateral and dorsal views, NIGP 138474, × 12, P341.8. 6-9. Broken cranidium in anterolateral, dorsal, anterior, and lateral views, NIGP 138475, × 9, P290.5. Figures 1-5 from Liostracina bella Zone; Figures 6-9 from the upper part of Wanshania wanshanensis Zone. Figures 10-13. Huayuanella paibiensis gen. et sp. nov. 10-13. Testaceous cranidium in anterolateral, obliquely anterior, lateral, and dorsal, views, holotype, NIGP 138476, × 5.7, P331.8. From Liostracina bella Zone. Figure 14. Huayuanella zhiqiangi gen. et sp. nov. 14. Incomplete, testaceous cranidium, holotype (refigured from Zhou et al., 1996, pl. 5, fig. 15), NIGP 132820, × 9. From the uppermost part of Nidanshan Group, Nidanshan, Hualong, Qinghai (in collection Lw 6). Figures 15, 16. Aethia rectangula Qian and Zhou, 1984 15, 16. Testaceous cranidium in dorsal and anterolateral views, NIGP 138477, × 18, W210.5. From the upper part of Wanshania wanshanensis Zone.

• 310.

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11

12 13

14

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Plate 57

Figures 1-15. Pianaspis sinensis (Yang in Zhou et al., 1977) 1. Meraspid cranidium, NIGP 138478, x 25, P 135.7. 2. Meraspid cranidium, NIGP 138479, x 25, P135.7. 3. Meraspid cranidium, NIGP 138480, x 25, P 135.7. 4. Meraspid cranidium, NIGP 138481, × 20, P 135.7. 5. Small testaceous cranidium, NIGP 138482, x 12, P135.7. 6. Small testaceous cranidium, NIGP 138483, x 12, P 135.7. 7. Testaceous cranidium, NIGP 138484, x 8, P135.7. 8. Nearly complete, testaceous cranidium, NIGP 138485, x 7.5, P130.95. 9. Incomplete, partly exfoliated cranidium, latex cast, NIGP 138486, x 8, P123.6. 10. Incomplete, partly exfoliated cranidium, NIGP 138487, × 5, P 123.6. 11. Incomplete testaceous pygidium, NIGP 138488, x 4, P135.7. 12. Partly broken and exfoliated pygidium, NIGP 138489, x 10, W82.1. 13. Testaceous pygidium, NIGP 138490, x 7, P123.6. 14. Testaceous pygidium, NIGP 138491, x 8. P136.3. 15. Incomplete hypostome, NIGP 138492, x 8. P 138.8. All from the lower part of Pianaspsis sinensis Zone.

• 312•

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Figures 1-9. Pianaspis sinensis (Yang in Zhou et al., 1977) 1, 2. Two testaceous cranidia in association, dorsal and anterolateral views, NIGP 138493, NIGP 138494, x 4, W82.1. 3. Small, testaceous librigena, NIGP 138495, x 12, P 123.6. 4. Nearly complete, partly exfoliated cranidium, NIGP 138496, x 3, P122.4. 5. Nearly complete, partly exfoliated cranidium, NIGP 138497, x 5, P122.4. 6. Testaceous librigena, NIGP 138498, x 5, P136.3. 7, 8. Testaceous cranidium in dorsal and anterolateral views, NIGP 138499, x 4, x 4, W99.3. 9. Testaceous librigena, NIGP 138500, x 5, W82.1. All from Pianaspsis sinensis Zone. Figure 10. Pianaspis attenuata (Lermontova and Chemysheva in Chemysheva, 1950) 10. Testaceous cranidium, NIGP 138501, x 5. P 184. From the lower part of Pianaspsis sinensis Zone.

• 314.

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8

10

Plate 59

Figures 1-14. Wangcunia wangcunensis Peng, Lin, and Chen, 1995 1. Crushed, nearly complete, testaceous exoskeleton, NIGP 118807, × 5, W56.7. 2. Incomplete, testaceous cranidium, NIGP 118808, × 8, W56.7. 3. Nearly complete, testaceous cranidium, NIGP 138502, × 8, W56.7. 4. Crushed, nearly complete, testaceous cranidium, NIGP 118804, × 6, W56.7. 5. Incomplete librigena, latex cast of fig. 8, NIGP 118806, x 8, W56.7. 6. Hypostome, NIGP 118811, × 8, W56.7. 7. Hypostome, NIGP 118810, × 8, W56.7. 8. Incomplete librigena, the same as fig. 5. 9. Incomplete, testaceous pygidium, NIGP 138503, × 2, W56.7. 10. Testaceous pygidium, NIGP 118812, × 6, W56.7. 11. Testaceous pygidium, NIGP 118817, × 5, W56.7. 12. Testaceous pygidium, NIGP 138504, × 3, W56.7. 13. Testaceous pygidium, NIGP 138505, × 2.5, W56.7. 14. Testaceous pygidium, NIGP 118813, x 2.5, W56.7. All from Dorypyge richthofeni Zone.

• 316.

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Figures 1-13. Parapianaspis hunanensis gen. et sp. nov. 1. Incomplete, testaceous, meraspid cranidium, NIGP 138506, x 12, P 123.6. 2. Nearly complete, testaceous, meraspid cranidium, NIGP 138507, x 12, P114. 3. Incomplete, testaceous, meraspid cranidium, NIGP 138508, x 12, P 123.6. 4. Nearly complete, testaceous, meraspid cranidium, NIGP 138509, x 12.5, P123.6. 5, 6. Nearly complete, testaceous, meraspid cranidium and latex cast, NIGP 138510, x 12.5, P123.6. 7. Incomplete, testaceous librigena, NIGP 138511, x 5, W88.1. 8. Testaceous librigena, NIGP 138512, x 5, W88.1. 9. Broken, testaceous cranidium, NIGP 138513, x 10, W88.1. 10. Incomplete, testaceous cranidium, latex cast, NIGP 138514, x 10, W88.1. 11. Broken, testaceous pygidium, NIGP 138515, x 6, W88.1. 12. Testaceous cranidium, latex cast, NIGP 138516, x 6, W88.1. 13. Two cranidia in association, NIGP 138517, NIGP 138518, x 6, W88.1. All from the lower part of Pianaspsis sinensis Zone.

• 318-

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10

12

13

Plate 61

Figures 1-10. Parapianaspis hunanensis gen. et sp. nov. 1, 2. Nearly complete, testaceous cranidium in dorsal and anterolateral views, latex cast, holotype, NIGP 138519, × 4, W88.1. 3. Nearly complete, testaceous cranidium, latex cast, NIGP 138520, × 2.5, W88.1. 4. Small, crushed, testaceous pygidium NIGP 138521, x 10, W88.1. 5. Incomplete testaceous pygidium, NIGP 138522, × 4, P123.6. 6. Incomplete testaceous pygidium, latex cast, NIGP 138523, × 6, W88.1. 7. Fragmentary pygidium, latex cast, NIGP 138524, × 6, W88.1. 8. Testaceous pygidium, latex cast, NIGP 138525, × 4, P123.6. 9. Slightly broken, testaceous pygidium, NIGP 138526, × 4, P123.6. 10. The same pygidium as fig. 8 with partial pleural region removed to show doublure, × 4. All from the lower part of Pianaspsis sinensis Zone. Figures 11-15. Pianaspis sinensis (Yang in Zhou et al., 1977) 11. Nearly complete, testaceous cranidium, NIGP 138527, × 4, P 138.8. 12. Nearly complete, testaceous cranidium, NIGP 138528, × 6, W82.1. 13. Broken, incomplete, testaceous cranidium, NIGP 138529, × 6, W82.1. 14. Inomplete, testaceous cranidium, NIGP 138530, × 6, W82.1. 15. Partly exfoliated, nearly complete cranidium, NIGP 138531, × 5, W87.6. All from the lower part of Pianaspsis sinensis Zone.

• 320 •

Plate 61

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Figures 1-17. Rhyssometopus zhongguoensis Zhou in Zhou et al., 1977 1. Incomplete, testaceous cranidium, NIGP 138532, × 7.5, P361.5. 2. Mostly exfoliated, incomplete cranidium, NIGP 138533, × 4, P344.6. 3. Partly exfoliated, incomplete cranidium, NIGP 138534, × 5, P348. 4. Mostly exfoliated, incomplete cranidium, NIGP 138535, × 5, P348. 5. Partly exfoliated, incomplete librigena, NIGP 138536, × 5, P348. 6-8. Mostly exfoliated, incomplete cranidium in oblique anterior, dorsal, and anterolateral views, NIGP 138537, × 4, P344.6. 9. Partly exfoliated, incomplete librigena, NIGP 138538, × 5, P348. 10. Mostly exfoliated, incomplete librigena, NIGP 138539, × 5, P348. 11, 12. Slightly exfoliated, nearly complete cranidium in oblique anterior, and dorsal views, NIGP 138540, × 4, P348. 13. Testaceous pygidium, NIGP 138541, × 5, P348. 14. Testaceous pygidium, NIGP 138542, × 5, P348. 15. Testaceous librigena, NIGP 138543, × 5, P356.5. 16. Testaceous pygidium, NIGP 138544, × 5, P348. 17. Partly exfoliated, incomplete pygidium, NIGP 138545, × 5, P348. All from the upper part of Liostracina bella Zone.

• 322 •

Plate 62

10

12

15

16

14

17

Plate 63 Figures 1-5. Changqingia laevis Peng, Lin, and Chen, 1995 1. Exfoliated cranidium, NIGP 118819, × 8, P 123.6. 2. Nearly complete, testaceous cranidium, NIGP 138546, × 7.5, P112.42. 3. Nearly complete, mostly exfoliated cranidium, holotype, NIGP 118818, × 8, P 123.6. 4, 5. Nearly complete, exfoliated cranidium in dorsal and anterior views, NIGP 118820, × 6, × 6, P123.6. Figures 1, 3, 4 from the lower part of Pianaspsis sinensis Zone; Figure 2 from the uppermost part of Dorypyge richthofeni Zone. Figures 6, 7. Changqingia intermedia (Walcott, 1906) 6. Incomplete, testaceous cranidium, NIGP 138547, x 7.5, P81. 7. Fragmentary, testaceous cranidium, NIGP 138548, × 7.5, P81. All from the upper part of Dorypyge richthofeni Zone. Figures 8-11. Menocephalites sp. cf. M. pauperata (Walcott, 1906) 8. Broken, testaceous cranidium, NIGP 138549, x 8, P37. 9. Incomplete, testaceous cranidium, NIGP 138550, x 8, P37. 10, 11. Nearly complete, testaceous cranidium in dorsal and lateral views, NIGP 138551, × 10, P37. All from the lower part of Dorypyge richthofeni Zone.

• 324.

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11

Plate 64

Figures 1--4. Menocephalites hunanensis sp. nov. 1. Small, testaceous cranidium, NIGP 138552, × 20, P64.5. 2, 3. Testaceous cranidium in anterolateral and dorsal views, holotype, NIGP 138553, × 15, P68.8. 4. Broken, testaceous pygidium, NIGP 138554, × 10, P68.8. All from Dorypyge richthofeni Zone. Figures 5, 6. Menocephalites? sp. 1 5, 6. Testaceous cranidium in dorsal and anterolateral views, NIGP 138555, × 18, W64.7. From the upper part of Dorypyge richthofeni Zone. Figures 7, 8. Menocephalites? sp. 2 7, 8. Partly exfoliated cranidium in anterolateral and dorsal views, NIGP 138556, × 8, P112.6. From the base of Pianaspsis sinensis Zone. Figures 9, 10. Helepagetia bitruncula Jell, 1975 9, 10. Cephalon in dorsal and anterolateral views, NIGP 138557, × 28, P261. From Wanshania wanshanensis Zone. Figures 11-15. Luyanhaoaspis sp. cf. L. inflata sp. nov. 11-15. Broken cranidium in dorsal, anterolateral, lateral, anterior and posterior views, NIGP 138558, × 7.5, P298.54. From the upper part of Wanshania wanshanensis Zone.

• 326 •

Plate 64

11

14 12

10

13

15

Plate 65

Figures 1-7. Limbishumardia wangcunensis gen et sp. nov. 1-5. Incomplete testaceous cranidium in anterolateral, dorsal, dorsal, obliquely anterior, obliquely posterior views, holotype, 3 is latex cast, NIGP 138559, × 16, W199.2. 6. Incomplete cranidium, NIGP 138560, × 16, W 199.2. 7. Incomplete cranidium, NIGP 138561, × 16, W 199.2. From Wanshania wanshanensis Zone. Figures 8-15. Oculishumardia hunania Peng, Babcock, Hughes, and Lin, 2003 8. Testaceous cranidium, NIGIP 132985, × 20, P326.9. 9. Incomplete, testaceous cranidium NIGIP 132982, × 27, W254.1. 10-12. Testaceous cranidium in dorsal, anterolateral and anterior views, NIGP 138562, × 30, PI3-2.75. 13, 14. Exfoliated cranidium showing anterolateral lobes, NIGIP 132984, × 20, W243.6. 15. Incomplete, testaceous cranidium, NIGP 133511, × 20, P[3-2.75. All from Liostracina bella Zone.

• 328 •

Plate 65

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Figures 1-9. Oculishumardia hunania Peng, Babcock, Hughes, and Lin, 2003 1, 2. Testaceous cranidium in anterolateral and dorsal views, holotype, NIGP 132983, x 20, x 20, W227. 3. Incomplete, testaceous cranidium, NIGP 133512, x 20, PI3-2.75. 4-6. Mostly exfoliated cranidium in dorsal, anterolateral and anterior views, NIGP 132986, all x 20, W243.6. 7, 8. Testaceous cranidium in anterolateral and dorsal views, NIGP 138563, x 25, x 25, W251.15. 9. Testaceous cranidium, the same as figured on P1.31, fig. 6b, NIGP 138564, x 25, W254.1. All from Liostracina bella Zone. Figures 10-19. Undetermined larva 10. Complete exoskeleton, NIGP 138565, x 25, W219.7. 11. Complete exoskeleton, NIGP 138566, x 25, W219.7. 12. Complete exoskeleton, NIGP 138567, x 25, W219.7. 13. Complete exoskeleton, NIGP 138568, x 25, W219.7. 14. Complete exoskeleton, NIGP 138569, x 25, W219.7. 15. Complete exoskeleton, NIGP 138570, x 25, W219.7. 16. Complete exoskeleton, NIGP 138571, x 25, W219.7. 17. Cranidium, NIGP 138572, x 25, W219.7. 18. Cranidium, NIGP 138573, x 25, W219.7. 19. Cranidium, NIGP 138574, x 25, W219.7. All from the basal part of Liostracina bella Zone.

• 330 •

Plate 66

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Figures 1-18. Pseudomapania cylindrica Yuan and Yin, 1998 1. Incomplete, testaceous cranidium, NIGP 138575, × 16, W228.3. 2. Nearly complete, slightly exfoliated cranidium, NIGP 138576, × 15, W210.5. 3. Incomplete, testaceous cranidium, NIGP 138577, × 16, W211.7. 4. Nearly complete, testaceous librigena, NIGP 138578, × 12, W211.7. 5. Nearly complete testaceous cranidium, NIGP 138579, × 15, W210.5. 6, 7. Nearly complete, testaceous cranidium in dorsal and anterolateral views, NIGP 138580, × 16, W211.7. 8, 9. Incomplete, testaceous cranidium in dorsal and anterolateral views, NIGP 138581, × 15, W199.2. 10. Incomplete, testaceous cranidium, NIGP 138582, × 15, W229.9. 11. Nearly complete, testaceous cranidium, NIGP 138583, × 11, W 196.3. 12. Incomplete, testaceous cranidium, NIGP 138584, × 15, W210.5. 13. Testaceous pygidium, NIGP 138585, × 20, W225. 14. Testaceous pygidium, NIGP 138586, × 20, W211.7. 15. Testaceous pygidium, NIGP 138587, × 20, P378.25. 16. Testaceous pygidium, NIGP 138588, × 15, P319.8. 17. Incomplete, testaceous pygidium, NIGP 138589, × 15, P319.8. 18. Mostly exfoliated pygidium, NIGP 138590, × 15, P319.8. Figures 1, 10, 13, 15 from the basal part of Liostracina bella Zone; Figures 2-9, 11, 12, 14, 16-18 from the upper part of Wanshania wanshanensis Zone.

• 332.

Plate 67

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Figures 1-9. Pseudomapania cylindrica Yuan and Yin, 1998 1. Incomplete, testaceous cranidium, NIGP 138591, x 18, W216.5. 2. Nearly complete, testaceous cranidium, NIGP 138592, x 16, W211.7. 3, 4. Testaceous cranidium in anterolateral and dorsal views, NIGP 138593, x 18, W219.7. 5. Nearly complete testaceous cranidium, NIGP 138594, x 20, W225. 6. Broken, testaceous, NIGP 138595, x 16, W211.7. 7. Testaceous pygidium, NIGP 138596, x 20, W225. 8. Mostly exfoliated pygidium, NIGP 138597, x 18, W 199.2. 9. Testaceous pygidium, NIGP 138598, x 20, W225. Figures 1, 2, 6, 8 from Wanshania wanshanensis Zone; Figures 3-5, 7, 9 from Liostracina bella Zone. Figure 10. Undetermined pygidium 2 10. Testaceous pygidium, latex cast, NIGP 138599, x 6, P269. From Wanshania wanshanensis Zone. Figure 11. Crepicephalina pergranosa Resser and Endo, 1937 11. Incomplete, mostly exfoliated cranidium, NIGP 138600, x 8, W33.6. From Dorypyge richthofeni Zone. Figures 12, 13. Paradistazeris sp. 12. Juvenile cranidium, NIGP 133514, x 16, PI3-2.75. 13. Incomplete, cranidium, NIGP 133513, x 13, P[3-2.75. From Liostracina bella Zone. Figures 14a, 15. Olenus punctatus sp. nov. 14a. Librigena in association with librigena of Proceratopyge (Proceratopyge)fenghwangensis Hsiang in Ergorova et al., 1963 (14b), dorsal view, showing anterior doublure, NIGP 138601, x 2, P[335.4. 15. Incomplete exoskeleton, latex cast, NIGP 138602, x 2, P[335.4. From Chuangia subquadrangulata Zone.

• 334 •

Plate 68

10

12

13

11

14

15

Plate 69

Figures 1-8. Olenus austriacus Yang in Zhou et al., 1977 1, 2. Partly exfoliated cranidium, NIGP 138603, × 3, × 3, PI336. 3. Testaceous exoskeleton, latex cast, NIGP 138604, × 2, PI327.2. 4. Largely exfoliated librigena, NIGP 138605, × 3, PI3 22.4. 5. Incomplete, testaceous cranidium, NIGP 133568, × 6, PI3-1.6. 6. Small, incomplete, testaceous cranidium, NIGP 133580, x 10, P13-1.6. 7. Small, nearly complete, testaceous cranidium, NIGP 133569, x 10, PI3-0.3. 8. Small, broken, testaceous cranidium, NIGP 133576, x 18, PI3-1.6. Figures 1-4 from Chuangia subquadrangulata Zone; Figures 5-8 from Liostracina bella Zone.

• 336 •

Plate 69

7

8

Plate 70

Figures 1-13. Olenus punctatus sp. nov. 1, 10. Slightly exfoliated, incomplete librigena in dorsal view and partial enlargement showing punctate prosopon, latex cast, NIGP 138606, x 3, x 7.5, PI335.4. 2. Testaceous, incomplete cranidium, NIGP 133545, x 11, PI3-1.80. 3. Testaceous, incomplete cranidium, NIGP 133544, x 7, PI3-1.80. 4. Largely exfoliated librigena, NIGP 138607, x 3, PI335.4. 5. Incomplete partly exfoliated cranidium, NIGP 138608, x 3, P[336. 6. Incomplete largely exfoliated cranidium, NIGP 138609, x 3, P[336. 7. Nearly complete, testaceous exoskeleton, holotype, NIGP 138610, x 3, PI335.4. 8, 9. Incomplete, partly exfoliated cephalon with 8 thoracic segments attached, dorsal view and partial enlargement showing punctate prosopon, NIGP 138611, x 5.5, PI336. 11. Largely exfoliated pygidium, NIGP 138612, x 3, x 7.5, PI335.4. 12, 13. Testaceous pygidium in dorsal view and partial enlargement showing punctate prosopon, NIGP 138613, x 3, x 18, PI335.4. Figures 1, 4-13 from Chuangia subquadrangulata Zone; Figures 2, 3 from Liostracina bella Zone.

• 338.

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Figures 1-16. Baikadamaspis linearis sp. nov. 1. Small, testaceous cranidium, NIGP 138614, × 15, P319.6. 2. Small, testaceous cranidium, NIGP 138615, x 15, P319.6. 3, 4. Small testaceous cranidium in dorsal and anterolateral views, NIGP 138616, x 8, P319.6. 5. Small, testaceous cranidium, NIGP 138617, × 10, P319.6. 6. Incomplete testaceous cranidium, latex cast, NIGP 138618, × 7, P319.6. 7. Testaceous cranidium, NIGP 138619, x 7, P319.6. 8. Incomplete cephalic doublure with rostral plate, NIGP 138620, x 12, P319.6. 9. Testaceous cranidium, NIGP 138621, x 8, P301. 10. Incomplete cephalic doublure, NIGP 138622, x 12, P319.6. 11. Incomplete, testaceous cranidium, NIGP 138623, x 5, P319.6. 12. Testaceous cranidium, NIGP 138624, × 5, P319.6. 13. Testaceous librigena (anterior portion) and cephalic doublure (posterior portion, showing the paradoublural line coinciding with inner margin of doublure), NIGP 138625, x 6.5, P319.6. 14. Testaceous pygidium, NIGP 138626, x 12, P319.6. 15. Testaceous pygidium, NIGP 138627, x 12, P319.6. 16. Partly exfoliated pygidium, showing pygidial doublure, NIGP 138628, x 15, P319.6. All from Liostracina bella Zone.

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Plate 72 Figures 1-12. Baikadamaspis linearis sp. nov. 1. Two testaceous cranidia in association, NIGP 138629, NIGP 138630, x 5, P319.6. 2a. Incomplete, partly exfoliated cranidium, NIGP 138631, x 7, P301. 3-5. Testaceous cranidium in anterolateral, dorsal and anterior views, holotype, NIGP 138633, x 6, P319.6. 6. Incomplete partly exfoliated cranidium, latex cast, NIGP 138634, x 7, P301. 7. Incomplete, testaceous librigena, latex cast, NIGP 138635, x 6, P319.6. 8. Incomplete testaceous pygidium, NIGP 138636, x 13, P319.6. 9. Hypostome, NIGP 138637, x 8, P319.6. 10. Testaceous pygidium, NIGP 138638, x 12, P319.6. 11. Testaceous pygid.ium, NIGP 138639, x 11.5, P319.6. 12. External mold of doublure, showing straight connective suture, NIGP 138640, x 6.5, P319.6. Figures 1, 3-5, 7-12 from Liostracina bella Zone; Figures 2, 6 from Wanshania wanshanensis Zone. Figure 2b. Huzhuia paratypica Yang, 1978 2b. Incomplete cranidium, NIGP 138632, x 7, P301.

• 342 •

Plate 72

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Figures 1-16. Baikadamaspispaibiensis sp. nov. 1. Small, incomplete testaceous cranidia, NIGP 138641, × 10, P378.25. 2. Incomplete, testaceous cranidium, NIGP 138642, × 12, PI35.1. 3. Partly exfoliated cranidium NIGP 138643, × 12, PI35.1. 4. Testaceous cranidium, NIGP 138644, × 12, PI35.1. 5, 6. Testaceous cranidium in dorsal and lateral views, NIGP 138645, × 12, PI35.1. 7-9. Nearly complete cephalon in dorsal, oblique anterior, and anterolateral views, holotype, NIGP 138646, × 10, P135.1. 10. Testaceous librigena with partial posterolateral projection attached, showing course of facial suture, NIGP 138647, × 8, PI35.1. 11, 12. Incomplete, testaceous cranidium in lateral and dorsal views, showing palpebral lobe, eye, and anterior branch of facial suture, NIGP 138648, × 5, PI322.4. 13. Testaceous pygidium, NIGP 138649, × 12, PI35.1. 14. Testaceous pygidium, NIGP 138650, × 12, PI35.1. 15. Testaceous pygidium, NIGP 138651, × 12, PI35.1. 16. Testaceous pygidium, NIGP 138652, × 12, PI35.1. All from the basal part of Chuangia subquadrangulata Zone.

• 344.

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Figures 1-3. Baikadamaspis sp. cf. B. granulosa (Yuan and Yin, 1998) 1, 2. Testaceous cranidium in anterolateral and dorsal views, NIGP 138653, × 12, x 12, P301. 3. Testaceous cranidium, NIGP 138654, x 8, PI360.3. Figures 1, 2 from Wanshania wanshanensis Zone; Figure 3 from Chuangia subquadrangulata Zone. Figures 4, 5. Baikadamaspis sp. 4, 5. Testaceous cranidium, NIGP 138655, × 7, P341.8. From Liostracina bella Zone. Figures 6-12. Baikadamaspis sinensis (Yang in Zhou et al., 1977) 6, 10-12. Incomplete cranidium in partial enlargement, showing surface prosopon and in dorsal (10), anterior (11), and anterolateral (12) views, latex cast, NIGP 138656, × 6, x 4, x 4, x 4, PI335.4. 7. Incomplete, testaceous cranidium, in dorsal view, latex cast, NIGP 138657, x 3.5, PI336. 8, 9. Small, incomplete, testaceous cranidium in dorsal, anterior and anterolateral views, CUGB 308003, x 8, from Chuangia-Prochuangia Zone, Huaqiao Formation, at Tingziguan, Fenghuang, western Hunan (original of Yang, 1978, pl. 13, fig. 11). Figures 6, 7, 10-12 from Chuangia subquadrangulata Zone. Figures 13-17. Undetermined Pygidium 2 13. Broken pygidium, latex cast, NIGP 138658, × 8.3, P273.8. 14-17. Broken pygidium. 14, latex cast from external mold, dorsal view; 15-17, in dorsal, anterolateral, and anterior views, NIGP 138659, × 8.5, P273.8. From Wanshania wanshanensis Zone.

• 346 •

Plate 74

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Figures 1-8. Schmalenseeia sinensis Yang in Yin and Li, 1978 1, 3. Incomplete, crumpled exoskeleton in dorsal and anterior views NIGP 138660, × 7 , P240.5. 2. Posterior portion of the same exoskeleton as in figures 1 and 3, latex cast from incomplete external mold, x 7. 4. Composite exoskeleton, photo from the fig. 1 and fig. 2, x 11. The numbers 10, 14, and 17 indicate the 10th, the 14th and the 17th thoracic segment respectively. 5. Incomplete cranidium, NIGP 138661, x 12, W 132.5. 6. Partial enlargement of the posterior part of fig. 2, showing the last four thoracic segments and pygidium. The numbers 14 and 17 indicate the 14th and the 17th thoracic segment respectively; pll and p12 are the first and the second pleural segments of the pygidium; lateral margins of the pygidium are indicated by white triangulars. 7, 8. Lateral view of the same incomplete exoskeleton as in figs. 1 and 2, x 9.4. The number 5 indicates the fifth thoracic segment. Figures 1-4, 6-8 from the base of Wanshania wanshanensis Zone; Figure 5 from Pianaspsis sinensis Zone.

• 348 •

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Figures 1-12. Schmalenseeia fusilis sp. nov., all from P268.3. 1. Small testaceous exoskeleton lacking librigenae, NIGP 138662, × 20. 2. Small incomplete, testaceous exoskeleton lacking librigenae, latex cast, NIGP 138663, x 20. 3. Small testaceous exoskeleton lacking librigenae, NIGP 138664, × 20. 4. Small testaceous exoskeleton lacking librigenae, NIGP 138665, × 20. 5. Small testaceous exoskeleton lacking librigenae, NIGP 138666, × 20. 6. Small testaceous exoskeleton lacking librigenae, NIGP 138667, × 20. 7. Testaceous exoskeleton lacking librigenae, NIGP 138668, x 20. 8. Testaceous exoskeleton lacking librigenae, note the linear cranidial border, NIGP 138669, x18. 9. Incomplete, testaceous exoskeleton lacking librigenae, the linear anterior cranidial border is indicated by triangular mark, NIGP 138670, x 18. 10, 11. Testaceous exoskeleton lacking librigenae in dorsal and lateral views, NIGP 138671, both x 15. 12. Incomplete, testaceous exoskeleton lacking librigenae; note the linear anterior cranidial border at the fight margin of the cranidium and the linear lateral border of pygothorax, NIGP 138672, x 15. All from the lower part of Wanshania wanshanensis Zone.

• 350.

Plate 76

10

11

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Plate 77

Figures 1-11. Schmalenseeia fusilis sp. nov., all from P268.3. 1, 2. The same exoskeleton as in P1. 76, fig. 12, in anterolateral and lateral views; note the linear anterior cranidial border and thoracopygidial lateral border indicated by triangle, × 15. 3-6. Mostly exfoliated exoskeleton lacking librigenae in dorsal, posterior, anterolateral, and lateral views, holotype; note the linear posterolateral border of pleural region of the thoracopygon is indicated by triangle, NIGP 138673, × 15. 7. Incomplete, testaceous exoskeleton lacking librigenae, NIGP 138674, x 15. 8-11. The largest-known, broken exoskeleton lacking librigena (length about 8 mm). 8, latex cast from the external mold of the anterior portion of the exoskeleton; 9, 10, posterior portion of the exoskeleton in dorsal and posterior views; 11, composite photo from 8 and 9, testaceous exoskeleton lacking librigenae, NIGP 138675, × 9. All from the lower part of Wanshania wanshanensis Zone.

• 352.

Plate 77

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Plate 78

Figure 1. Undetermined librigena 1 1. Incomplete, partly exfoliated librigena, NIGP 138676, × 2, P319.6. From Liostracina bella Zone. Figures 2, 3. Undetermined librigena 2 2. Testaceous librigena, NIGP 138677, × 4.5, W228.3. 3. Mostly exfoliated librigena, latex cast, NIGP 138678, × 6, P1372. Figure 2 from Liostracina bella Zone; Figure 3 from the base of Shengia quadrata Zone. Figure 4. Undetermined librigena 3 4. Slightly exfoliated librigena, latex cast, NIGP 138679, x 6, P~72. From the base of Shengia quadrata Zone. Figure 5. Undetermined librigena 4 5. Nearly complete librigenna, NIGP 138680, × 10, W211.7. From the top part of Wanshania wanshanensis Zone. Figures 6-8. Sudanomocarina sp. cf. S. changi Jell in Jell and Robison, 1978 6, 7. Nearly complete librigena in dorsal and anterolateral views, NIGP 138681, x 8, W0. 8. Incomplete, exfoliated librigena, NIGP 138682, x 8, W0. From the unnamed Zone. Figure 9. Undetermined hypostome 1 9. Nearly complete, testaceous hypostome, NIGP 138683, x 4, P319.6. From the basal part of Liostracina bella Zone. Figure 10. ?Shengia trapezia Peng, 1992 10. Nearly complete, partly exfoliated hypostome, NIGP 138684, × 8.5, PI322.4. From Chuangia subquadrangulata Zone. Figure 11. Shengia sp. 11. Incomplete, partly exfoliated pygidium, NIGP 138685, × 4, PI322.4. From Chuangia subquadrangulata Zone. Figure 12. Baikadamaspis linearis sp. nov. 12. Nearly complete, partly exfoliated hypostome, NIGP 138686, × 12, P301. From Wanshania wanshanensis Zone. Figures 13, 14. Baikadamaspis paibiensis sp. nov. ? 13, 14. Nearly complete, mostly exfoliated hypostome, NIGP 138687, × 4.5, PI323.5. From Chuangia subquadrangulata Zone. Figures 15, 16. Undetermined hypostome 2 15, 16. Slightly exfoliated hypostome in dorsal and lateral views, NIGP 138688, × 7, P277. From Wanshania wanshanensis Zone. • 354 •

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