Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229 – 249 www.elsevier.com/locate/palaeo Early Cambrian metazoan fossil record of South China: Generic diversity and radiation patterns Guoxiang Li a,⁎, Michael Steiner b , Xuejian Zhu a , Aihua Yang c , Haifeng Wang a , Bernd D. Erdtmann b a State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China b Technische Universtät Berlin, Sekr. ACK14, Ackerstrasse 71-76, D-13355 Berlin, Germany c Department of Earth Sciences, Nanjing University, Nanjing 210008, China Accepted 5 March 2007 Abstract South China with its rich fossil record is an important region for studying Early Cambrian metazoan diversity patterns. Compilation of a database of metazoan genera from the traditional Lower Cambrian (Cambrian Series 1–2) of South China allows a quantitative analysis of biodiversity changes. The dataset shows that about 876 genus names have been reported from the Lower Cambrian of South China. Of these only about 582 genera are now considered valid after taxonomic scrutiny. Plotting generic diversity shows a two-part, first-order trend: 1, diversity increase through the most of the Meishucunian and Qiongzhusian stages (South China usage), but punctuated by a modest decline during the upper Meishucunian; followed by 2, diversity decline in the Canglangpuan and Longwangmiaoan stages (South China usage). About 155 genera of small shelly fossils (SSFs) occur in Meishucunian rocks. The Meishucunian fauna is dominated by helcionellids, orthothecimorph hyoliths and other enigmatic fossils and is very consistent with the pretrilobitic ‘Tommotian Fauna’. Diversity increases in the Qiongzhusian stage, reaching 304 genera, the peak value known for the Cambrian. The Qiongzhusian fauna is characterized mainly by the occurrence of dominant pan-arthropods and crown-group brachiopods, representing an initial phase of the Cambrian Evolutionary Fauna. A substantial decline in generic diversity through the Canglangpuan (ca 153 genera) and Longwangmiaoan (ca 69 genera) stages is due partly to a eustatic regression, and it approximates the global pattern, reflecting an extinction event at the end of the Lower Cambrian. The observed regional biodiversity through this interval has both evolutionary and taphonomic influences: the decline in SSF diversity beginning in the upper Meishucunian was not only related to the decimation of SSFs, but also to a reduction in phosphogenesis. An apparent peak in diversity during the Qiongzhusian was somewhat exaggerated by the exceptional Chengjiang Biota. © 2007 Elsevier B.V. All rights reserved. Keywords: Generic diversity; Diversification pattern; Metazoa; Early Cambrian; South China ⁎ Corresponding author. E-mail address: gxli@nigpas.ac.cn (G. Li). 0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2007.03.017 230 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 1. Introduction On the basis of taxonomic diversity, quantitative analysis of the metazoan fossil record is one of the most significant approaches for improving our understanding of general diversity patterns. In past decades, several quantitative evaluations of metazoan diversity during the traditional Early Cambrian have been made mainly through analyzing global generic diversity (e.g., Sepkoski, 1992; Zhuravlev and Wood, 1996; Zhuravlev, 2001). Previous diversity analyses show that the early– mid Early Cambrian witnessed a striking increase in the diversity and abundance of metazoan fossils (Sepkoski, 1992; Zhuravlev, 2001). Subsequent to the early–mid Early Cambrian radiation event, metazoan fossils suffered a strong decline in diversity during the late Early Cambrian. This substantial decline reflects an extinction event, which was related to the Hawke Bay Regression (Palmer and James, 1979; Zhuravlev and Wood, 1996) and represents the first major extinction of the Phanerozoic (Zhuravlev, 2001). But previous statistical results may be rendered less conclusive by imprecision and uncertainties in international correlations of Cambrian strata (Geyer and Shergold, 2000; Babcock et al., 2005), taxonomic oversplitting (Bengtson, 1992; herein), taphonomic biases (Porter, 2004), and perhaps other factors. Ideally global diversity patterns should be synthesized from internally consistent regional data. Detailed quantitative study of the regional fossil records should be a prerequisite to a well integrated evaluation of the global diversity and radiation patterns of Early Cambrian metazoans since regional data are usually less influenced by uncertainties in stratigraphic correlation. The relation between biodiversity and taphonomic factors can also be better understood at the regional scale. Lower Cambrian sedimentary successions ranging from shallow to deep marine environments are well developed and widely exposed in South China, and contain highly diverse and abundant metazoan fossils (Zhu et al., 2003). South China is one of the most important regions for investigating the origin and early diversification of metazoans. Early Cambrian metazoan fossils from this region have been intensively studied and systematically described in hundreds of papers (or books), revealing that they represent significant material for studying Early Cambrian metazoan diversity and radiation patterns. Although previous achievements in taxonomy, biological affinities and stratigraphic correlation provide substantial data for assembling a database, which could stimulate quantitative improvement in our understanding of diversity changes and radiation patterns of early metazoans, most previous analyses have used a qualitative approach. General patterns of Early Cambrian metazoan diversity (especially at genus or species level) in South China largely remain an open question. The purpose of this paper is to quantitatively analyze generic diversity of Early Cambrian metazoan fossils from South China. Using database compilations of metazoan genera, we attempt to clarify our understanding of the general patterns of radiation and diversity, and to increase our understanding of relative influences of general radiation patterns and taphonomic factors upon diversity. 2. Data compilation 2.1. Chronological framework A quantitative analysis of generic diversity requires a practical bio- or chrono-stratigraphic framework so that genus occurrences of early metazoan fossils can be logically ordered with respect to time intervals in order to compile effectively a database of genus occurrences from published studies. The traditional Lower Cambrian stratigraphy in South China has been intensively studied during past decades, and new bio-, litho-, and chemostratigraphic data (e.g., Qian, 1999; Zhu et al., 2001; Zhang, 2003; Steiner et al., this volume) add to and revise the information developed over several decades of work. These studies enable compilation of a database of metazoan fossils and allow quantitative analysis of changes in taxonomic diversity, which is accepted here as simple diversity: a result of alpha and beta diversity (Zhuravlev and Naimark, 2005). Historically, the Lower Cambrian of South China has been subdivided into four stages (Table 1), in ascending Table 1 Six stratigraphic intervals utilized for the database compilation of genus occurrences in South China, and tentative correlations with the Siberian Platform Lower Cambrian South China Siberian Platform Longwangmiaoan Canglangpuan Qiongzhusian Upper Meishucunian Mid Meishucunian Lower Meishucunian Toyonian Botomian Atdabanian Tommotian Nemakit–Daldynian The lower Meishucunian covers the Anabarites trisulcatus–Protohertzina anabarica assemblage zone, the mid Meishucunian covers the Siphogonuchites triangularis–Purella squamulosa and the Watsonella crosbyi assemblage zones, and the upper Meishucunian covers the barren zone and Sinosachites flabelliformis–Tannuolina zhangwentangi assemblage zone. G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 order, the Meishucunian, Qiongzhusian (Chiungchussian), Canglangpuan (Tsanglungpuan), and Longwangmiaoan (Lungwangmiaoan) (Luo et al., 1994; Zhang, 2003). It needs to be noted that recently the conventional Lower Cambrian has been divided into two series, i.e. the 1st (subtrilobitic interval) and 2nd Series of the Cambrian System (Ogg, 2005), and each series comprising two stages (Peng, 1999; Peng and Babcock, 2001; Babcock et al., 2005; Peng et al., 2006). The new framework is not completely ready for implementation in the lower half of the Cambrian because international stratigraphic tie points have not been fully resolved (Zhu et al., 2001; Steiner et al., this volume). Until a globally applicable chronostratigraphic subdivision is agreed upon, it is best to use one of the regional schemes developed for South China for work of the type reported here. Furthermore, because the occurrences of Cambrian metazoan genera have usually been reported relative to the traditional four-stage biostratigraphically based scale, we prefer to continue to follow this usage here. At the moment, this is the most practical methodology for developing a database of generic occurrences from published references. The lower half of the Cambrian is bracketed in time between ca. 542 Ma and ca. 510 Ma (Ogg, 2005), but there is some uncertainty about those dates and about the duration of each traditional stage interval of South China. The dating within the Lower Cambrian is up to now largely uncertain especially in South China. Except for a SHRIMP zircon age of 538.2 ± 1.5 Ma from the lower Zhongyicun Member (lowest Meishucunian stage) in eastern Yunnan (Jenkins et al., 2002), there are indeed few well constrained numerical age data for the Lower Cambrian in South China. Therefore, the duration of each of the four conventional stages of the Lower Cambrian has to be inferred from extrapolation and international stratigraphic correlation, notwithstanding the considerable uncertainties regarding Lower Cambrian correlation due to the strong provincialism of the small skeletal fossils and trilobites and to the clear facies dependence of archaeocyaths (Xiao et al., 2005a). Strata in the lower half of the Cambrian of South China are commonly correlated with the Siberian Platform (Table 1). Based on bio- and chemo-stratigraphic evidence, the Meishucunian has been roughly correlated with the Nemakit–Daldynian and the Tommotian (Brasier et al., 1990; Zhang et al., 1997; Qian et al., 2001; Zhu et al., 2001), the Qiongzhusian with the Atdabanian (Zhu et al., 2001; Steiner et al., this volume), the Canglangpuan with the Botomian, and the Longwangmiaoan with the Toyonian (Luo et al., 1994). However, divergent interpretations have been published, e.g., the upper Meishucunian has been correlated with the upper Atdabanian 231 (Zhuravlev and Riding, 2001), or Botomian (Landing, 1994). Clearly, the time duration of the Meishucunian, as traditionally defined, is extremely long, roughly equal to the length of the 1st Series in the model stratigraphic chart (Babcock et al., 2005; Ogg, 2005). Even if the pretrilobitic Meishucunian is correlated with the Nemakit– Daldynian and the Tommotian, its duration may span from 10–15 Myrs (Bowring and Erwin, 1998) to 22 Myrs (ca. 542–520 Ma) (Landing et al., 1998; Babcock et al., 2001). Hence, the duration of the other three stages (Qiongzhusian, Canglangpuan and Longwangmiaoan) together may be roughly equivalent to, or even shorter than, the Meishucunian. The pre-trilobitic Meishucunian is biostratigraphically subdivided into four SSF assemblage zones (Qian et al., 2001; Steiner et al., this volume). Considering that its duration is much longer than each of the other three stages, a general statistical account of the Meishucunian metazoan taxa will easily obscure some significant features of the biotic diversity. During compilation of the database, the Meishucunian was therefore subdivided into three intervals based on SSF biostratigraphic zonation (Qian et al., 2001; Steiner et al., this volume): (1) the lower Meishucunian (Anabarites trisulcatus–Protohertzina anabarica assemblage zone), (2) the middle Meishucunian [Siphogonuchites triangularis–Purella squamulosa and Watsonella crosbyi (=Heraultipegma yunnanensis) assemblage zones], (3) the upper Meishucunian [the barren zone (e.g., the lower Shiyantou Formation in east Yunnan) and Sinosachites flabelliformis–Tannuolina zhangwentangi Assemblage Zone]. The middle Meishucunian covers two SSF biozones because biotratigraphic occurrences of many SSF genera in literature were commonly labeled as the former Siphogonuchites triangularis–Paragloborilus subglobosus Assemblage Zone, although later it was further subdivided into two zones (the Siphogonuchites triangularis–Purella squamulosa and the Watsonella crosbyi assemblage zones)(see Steiner et al., this volume). Definition and correlation of the Qiongzhusian, Canglangpuan and Longwangmiaoan stages is mainly based on trilobite biostratigraphic zonation (Luo et al., 1994; Zhang, 2003). Added to the three Meishucunian intervals, altogether six chronostratigraphic bins are utilized in the database compilation of genus occurrences, though they are not intended to represent equal time intervals. 2.2. Faunal data Metazoan fossils are found widely in the Lower Cambrian strata of South China and are described in published literature, thus providing significant material 232 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 for analyzing diversity changes. Metazoans of varying taxonomic levels, from genus to order, generally exhibit congruent diversity patterns through the Cambrian (Sepkoski, 1992), but generic diversity is most often used for studying radiation or extinction patterns (e.g., Bengtson, 1992; Zhuravlev and Wood, 1996; Zhuravlev, 2001; Porter, 2004) because genus-level diversity is closest to species diversity. In addition, it is apparent that many synonyms among the metazoan species reported from the Lower Cambrian, especially for the small shelly fossils described from South China. A full taxonomic revision of all such species would be a difficult task at present. Hence, our compilation incorporates all published genus occurrences of metazoan fossils (including poriferans) from South China. The dataset includes valid metazoan genera, as well as their synonyms (except for trilobites) and invalid taxonomic names. Altogether, about 876 genera have been described from the Lower Cambrian of South China (see Appendix A), but there exist many problems concerning generic identifications. After referring to previous taxonomic revisions in the literature and our preliminary taxonomic scrutiny, only about 582 generic names are considered valid. The other names include respectively junior synonyms (189 generic names), problematic erections (72 generic names), inappropriate assignments (28 generic names) or even pseudofossils (5 generic names). A multiple treatment of disarticulated sclerites of one organism is one of the major reasons for the proliferation of taxonomic names (Bengtson, 1992), while incorrect morphological erection is another reason (Hou et al., 2001). In accordance with the six chronostratigraphic bins of the Lower Cambrian utilized in the data compilations, about 745 generic occurrences were used in analyses of diversity patterns after elimination of some inappropriate names. Although this dataset represents the most complete list of genus occurrences in the Lower Cambrian of South China at present, there still exist some limitations. The assessment of synonymies could not wholly exclude a subjective view, and the stratigraphic occurrences of some genera are somewhat imprecise. New discoveries and insights are being made continuously, and so the results are subject to future modification. For convenience in assessing generic occurrences and factoring in future taxonomic changes, the database of generic occurrences is provided here in Appendix A. Presently known chronostratigraphic ranges of genera are indicated, and instances of apparent taxonomic oversplitting are marked. Genera are listed alphabetically within higher taxa, although the assignments of many genera to higher taxa are uncertain. 3. Results: patterns in generic diversity In South China, the earliest records of metazoans derive from the early Ediacaran Doushantuo Biota. The Doushantuo Biota includes abundant phosphatized embryo fossils (Xiao et al., 1998; Chen, 2004) and some body microfossils, such as a secondary phosphatized cnidarian (Xiao et al., 2000) and a sponge (Li et al., 1998). Some macrofossils from the Miaohe Biota of the upper Doushantuo Formation were described as metazoans, but most of them are now considered to be colonial prokaryotes or multicellular algae, and only about two genera possibly resemble animal remains (cnidarian or sponge) (Xiao et al., 2002). Apparently there are no body fossils of complex bilaterians in the Doushantuo Formation, though some of the phosphatized embryos may show affinities with bilaterians (Xiao et al., 1998; Chen, 2004). In the Gaojiashan Biota of the upper Ediacaran Dengying Formation, there occur several tubular fossils, such as Cloudina, Sinotubulites, Conotubus and Chenella (Hua et al., 2000, 2005), which represent some of the earliest mineralized exoskeletons. Most of the tubular fossils have debatable zoological affinities. Cloudina, for example, may be assignable to cnidarians or sponges (Conway Morris, 1993; Budd and Jensen, 2003), or perhaps to annelids (Chen and Sun, 2001; Hua et al., 2005). Except for a fossil with a quilted body plan from the Yangtze Gorges region (Xiao et al., 2005b), typical Ediacaran-grade fossils are scarce in South China. Likewise, Ediacaran-type metazoan genera have rarely been discovered from Lower Cambrian deposits, and this is consistent with the global record. Compared with the limited diversity of metazoan fossils in the Ediacaran, the Lower Cambrian is marked by an extraordinary metazoan diversification. Bilaterian metazoans rapidly diversified into the habitats vacated by the demise of the Ediacaran fauna. Among the 34 extant animal phyla (Brusca and Brusca, 2002), about 19 (Fig. 1), including Chordata (Shu et al., 1999; Chen et al., 1999; Shu, 2003), are known from the fossil record up to the Qiongzhusian stage in South China. In addition, many extinct clades made their first appearance in the Lower Cambrian. The explosive radiation occurred mainly during Meishucunian and Qiongzhusian, while biodiversity declined through the Canglangpuan and Longwangmiaoan. 3.1. Major trend Plotting generic diversity in South China reveals a striking change of a sigmoidal pattern through the Early G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 233 Fig. 1. The sequential occurrence of the early metazoan fossils during the Ediacaran and early Cambrian in South China. Cambrian (Fig. 2A). First, generic diversity increases through the Meishucunian–Qiongzhusian interval, but is punctuated by a modest decline during the late Meishucunian. Second, the most dramatic increase in generic diversity occurs during the Qiongzhusian, when metazoans attained their highest generic diversity not only of the Early Cambrian but also of the whole Cambrian in South China (Rong et al., in press). Third, diversity decreases significantly through the Canglangpuan–Longwangmiaoan interval. 3.1.1. The early–mid Meishucunian SSF diversification The increase in generic diversity through the early–mid Meishucunian is exemplified by the abrupt occurrence of diverse small skeletal fossils (i.e., small shelly fossils), and this has been taken as the first stage of the “Cambrian Explosion” (Qian, 1999; Chen, 2004). The dataset shows that about 19 SSF genera occur in the lower Meishucunian deposits (i.e., in the Anabarites trisulcatus–Protohertzina anabarica assemblage zone). SSFs of this interval mainly include cnidarians (6 genera), hyoliths (4), mollusks (1), protoconodonts (1), enigmatic tubular fossils (2), enigmatic sclerites (2) and spheroidal fossils (3). All 19 genera originated anew and extended to the mid Meishucunian. Edicacaran taxa are not known to have ranged upward into this interval. The highest SSF generic diversity occurs in the mid Meishucunian (i.e., the 2nd and 3rd SSF assemblage zones) (Steiner et al., this volume). Among 140 known genera in this interval, about 121 are new occurrences, representing a major diversification of SSFs. SSFs of the mid Meishcunian record include fossils assigned to the Porifera, Cnidaria, Annelida(?), Mollusca, Chaetognatha (protocondonts), and to many extinct clades, e.g., hyolithes, stem-group brachiopods, tommotiids, coeloscleritophorans, cambroclaves, coleoloids. Various enigmatic tubular fossils and sclerite fossils are known from this interval. The dominant components are hyoliths 234 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 mainly include poriferans (4 genera), cnidarians (5), hyoliths (10), mollusks (12), coeloscleritophorans (6), protoconodonts (1), enigmatic tubular fossils (6), enigmatic sclerites (12) and spheroidal fossils (2). 3.1.3. The Qiongzhusian major diversification Strata of the Qiongzhusian Stage record a dramatic increase in biodiversity. The metazoan diversity attains a Cambrian peak at various taxonomic levels, representing the major phase of the “Cambrian Explosion”. The skeletal fossils (Qian, 1999) and both biomineralized and non-biomineralized organism of the Chengjiang biota (Hou et al., 2004; Chen, 2004) show that there are about 19 extant metazoan phyla (of which ca. 14 phyla with their first appearances; Fig. 1), and more than 120 families occur in the Qiongzhusian deposits (Rong et al., in press). The dataset shows that about 304 metazoan genera (Fig. 2A) occur in the Qiongzhusian Stage, among which 266 genera have their first appearances. Metazoan fossils of this interval mainly include arthropods (ca. 113 genera) (Fig. 5), poriferans (51, including 18 archaeocyathid genera), hyoliths (22), phosphatic microbrachiopods (13), priapulids and other worms (16), vetulicolians and chordates (12). The presence of abundant arthropods, crown-group brachiopods (Li and Holmer, 2004) and enchinoderms (Shu et al., 2002) imply that the Cambrian Evolutionary Fauna (sensu stricto; Sepkoski, 1992) began from the Qiongzhusian age. Fig. 2. Showing patterns of metazoan generic diversity through the Early Cambrian. (A) Diversity curve for South China. (B) Global diversity curve (redrawn and compiled from Zhuravlev, 2001, Fig. 8.1A). NEMAK = Nemakit–Daldynian. (about 19 genera), mollusks and mollusk-like fossils (about 55 genera), and enigmatic sclerite fossils (more than 20 genera). 3.1.2. The late Meishucunian decline A modest decline in generic diversity occurred in the late Meishucunian, and it looks like a temporary interruption of the general radiation pattern exemplified through the Meishucunian–Qiongzhusian interval on the diversity trajectory (Fig. 2A). This diversity decline is most evident among SSFs (especially mollusks and mollusk-like fossils), and reflects the beginning of the decimation of SSFs. The dataset shows that about 58 SSF genera occur in the upper Meishucunian deposits (i.e., the Sinosachites flabelliformis–Tannuolina zhangwentangi assemblage zone), of which 15 genera made their first appearance in this interval. SSFs of this interval 3.1.4. The late Early Cambrian decline The generic diversity of the metazoan fossils exhibits a prolonged substantial decline through the Canglangpuan–Longwangmiaoan interval in South China. Low generic diversity was maintained through the late half of the Cambrian (the traditional Middle and Late Cambrian) (Rong et al., in press). The dataset shows that about 153 genera occur in Canglangpuan deposits, of which 112 have their first appearances during this interval. Canglangpuan faunas mainly include arthropods (ca. 107 genera), poriferans (23, including 19 archaeocyathid genera), hyoliths (7), brachiopods (5), mollusks (5), priapulids and other worms (3), ventulicolians (1), cnidarians (2), chancelloriids (1). The dominance of arthropods (mainly trilobites) is quite apparent. Compared with the Qiongzhusian faunal assemblage, most metazoan clades (except for in trilobites and archaeocyaths) exhibit a very distinct decline in generic diversity. Trilobites show a dramatic increase in generic diversity (from 33 genera in the Qiongzhusian to about 91 genera in the Canglangpuan). Archaeocyaths have a generic diversity of 19 in the Canglangpuan, which is nearly equal with that of the Qiongzhusian (18 genera). G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 The decline in generic diversity is most dramatic from the upper Canglangpuan to the Longwangmiaoan. All metazoan clades either exhibit a distinct decline in generic diversity or have no fossil record in Longwangmiaoan rocks. The dataset shows that about 69 genera occur in the Longwangmiaoan deposits, of which 20 make their first appearances in this interval. Trilobite fossils are the predominant components of the Longwangmiaoan assemblages: among the total record of 69 genera, 55 are trilobites (Fig. 8). Longwangmiaon faunas mainly include arthropods (55 genera), cnidarians (3), hyoliths (2), brachiopods (3), mollusks (3), and echinoderms (1). 3.2. Origination rate and extinction rate Both origination and extinction rates play an important role in the diversity patterns. If the two rates are equal, then diversity will be in equilibrium. A diversity increase usually results from an origination rate being larger than the extinction rate. Based on the dataset, we attempted to derive curves of origination and extinction rates at the generic level (Fig. 3A, B) although it is understood that the subdivision of the Early Cambrian into six chronostratigraphic bins is not timeaveraged. Origination rate was calculated by newly 235 occurring genera/total genera per biostratigraphic interval, while extinction rate was calculated by extinct (or disappeared) genera/total genera per biostratigraphic interval. At the genus level, the origination rate attained its first peak in the early–mid Meishucunian, and this peak mainly reflects a rapid diversification of SSFs. The peak is then followed by a dramatic decline in the late Meishucunian. The second peak in origination rate occurs in the Qiongzhusian, which received a major boost due to the exceptional Chengjiang Biota. Origination rate through the Canglangpuan and Longwangmiaoan interval is also fairly high (above 0.7) but still less than that of the Qiongzhusian (Fig. 3A). The first peak in extinction rate occurs at the end of the mid-Meishucunian (Fig. 3B), and it is followed by the lowest extinction rate of the Early Cambrian in the late Meishucunian. Extinction rates attain their highest values through the late Qiongzhusian and Canglangpuan. The high extinction (disappearance) rate at the end of the Qiongzhusian is apparently biased by the fact that many exceptionally preserved fossils in the Chengjiang Biota scarcely occur in the overlying rocks of the Canglangpuan stage, implying that the disappearances have a taphonomic component. The high extinction rate indicated in the Canglangpuan largely reflects an extinction event at the end of the Canglangpuan. The early–mid Meishucunian increase in generic diversity is inferred to be due to high origination and low extinction rates. That the late Meishucunian diversity decline shows both low extinction and origination rates indicates that this decline may not be a true extinction but a consequence of environmental and taphonomic factors. Generic diversity in the Qiongzhusian is characterized by both high origination and extinction (disappearance) rates, and these rates are strongly influenced by the appearance of many non-biomineralized fossils that most occur in the Chengjiang Lagerstätten. 3.3. Faunal succession through the Early Cambrian interval Fig. 3. Origination rate (A) and extinction rate (B) of the Early Cambrian metazoans in South China. Compared with the Ediacaran fossil record, the Early Cambrian is characterized by the rapid diversification of various bilaterians (Budd, 2003). Although the Early Cambrian biodiversification seems quite abrupt or explosive in relation to the enormity of geologic history, first appearances of clades in the Early Cambrian interval tend to be sequential (Fig. 1). For instance, hyoliths, mollusk-like maikhanellids, and chaetognaths (protoconodonts) first appear in the early Meishucunian Age, coeloscleritophorans and tommotiids in the middle and 236 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 Fig. 4. The composition of the Meishucunian faunas in South China. Abbreviations for Figs. 4–8: AT = Pan-arthropods, BR = Brachiopods, CD = Cnidarians, CG = Chaetognaths, CS = Coeloscleritophorans, DT = Deuterostomians, EC = Echinoderms, HT = Hyoliths and hyolith-like fossils, MC = Mollusks and cap-shaped fossils, OC = other fossils from the Chengjiang biota, OM = other Meishucunian skeletal fossils, PP = Priapulids and other worms, PR = Poriferans, SB = Stem group brachiopods, TM = Tommotiids, TR = Trilobites, OT = Others. late Meishucunian, while crown group brachiopods, arthropod body fossils and lobopods first appear in the early Qiongzhusian Age. The arthropod record is somewhat complicated by arthropod trace fossils (Zhu, 1997) and possible stem group arthropod embryos (Steiner et al., 2004), both of which first occur in the Meishucunian Age. Different taxa within single phylum also have sequential first appearances. For example, helcionellids (monoplacophorans) first appear in the early Meishucunian, rostroconchs (if Watsonella being a rostroconch) first appear in the middle Meishucunian, and if the taxa with coiling shells such as Aldanella and Pelagiella belong to helcionellids, then the first occurrence of gastropods may possibly be in the latest Cambrian (Gubanov and Peel, 2000). Due to the asynchronous diversifications of different clades and due to taphonomic factors, there seems to be a rapid succession of distinct faunas through the Lower Cambrian. The fossil record of South China shows that the clades occurring in the Meishucunian include the extant Porifera, Cnidaria, Annelida (?), Mollusca, Chaetognatha, and include many extinct taxa, such as hyoliths, Fig. 5. The composition of the Qiongzhusian faunas in South China. Fig. 6. The composition of the Qiongzhusian Chengjiang Biota in eastern Yunnan. stem-group brachiopods, tommotiids, coeloscleritophorans, cambroclaves, etc. About 155 metazoan fossil genera have been described from Meishucunian rocks of South China, and the Meishucunian Fauna is dominated by helcionellids, hyoliths, problematic cap-shaped and tubular fossils (Fig. 4). Most of the Meishucunian skeletal fossils are problematic in terms of biological affinities, and infrequently extend upwards to the Qiongzhusian. The Meishucunian fauna is distinctively different from the Qiongzhusian to Furongian faunal assemblages, and is consistent with the “Tommotian Fauna” (Sepkoski, 1992). The occurrence of abundant skeletal fossils in the Meishucunian reflects the beginning of rapid diversification of metazoans. After the late Meishucunian, many groups of SSFs – the major components of the “Tommotian Fauna” – either disappeared or became subdominant. The fossil record of the Qiongzhusian demonstrates that the clades having their first appearances in the interval include 14 extant phyla and an extinct phylum, Vetulicolia (Shu et al., 2001). Considering that the Qiongzhusian may only represent an interval of a few million years (Babcock et al., 2001), it becomes evident that the rate of metazoan diversification during this interval is very rapid. Among 304 metazoan genera described from the Qiongzhusian rocks of South China, Fig. 7. The composition of the Canglangpuan faunas in South China. G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 Fig. 8. The composition of the Longwangmiaoan faunas in South China. the dominant fossils are pan-arthropods (ca 127 genera) (Fig. 5). A separate analysis of the Chengjiang Biota also shows a similar result: among 162 metazoan genera, about 81 genera belong within pan-arthropods (Fig. 6). The Qiongzhusian fauna is mainly characterized by the occurrence and dominance of pan-arthropods and by the decline in the SSF diversity. The obvious decline of SSF diversity reflects the decimation of the enigmatic small shelly fossils which flourished in the Meishucunian. The dominance of pan-arthropods and the occurrence of crown-group brachiopods (Li and Holmer, 2004) and echinoderms (Shu et al., 2002) demonstrate that the Qiongzhusian fauna represents an initial phase of the Cambrian Evolutionary Fauna (sensu strict; Sepkoski, 1992). The faunal assemblages of the Canglangpuan and Longwangmiaoan ages are still typical of the Cambrian Evolutionary Fauna (sensu stricto) with regard to the predominance of trilobites although the generic diversity shows a dramatic decline through the interval. The Canglangpuan fauna mainly consists of trilobites, bradoriids, archaeocyaths, brachiopods, and hyoliths (Fig. 7), and more than half of the genera are trilobites. The depauperate Longwangmiaoan fauna comprise mainly trilobites (about 80% of genera are trilobites) (Fig. 8). There is no record of archaeocyaths in the Longwangmiaoan fauna since they became extinct at the end of the Canglangpuan in South China (Yang et al., 2006). 4. Discussion 4.1. Analyses of biodiversity variations The observed variation in generic diversity is an integrated result of radiation, extinction and taphonomic influences. Generic diversity increase through the Meishucunian and Qiongzhusian is consistent with the major episode of the Cambrian radiation. The early–mid 237 Meishucunian diversity increase is mainly due to the diversification of SSFs. About the same time, phosphatebearing deposits were deposited widely across the Yangtze Platform, e.g., in the Zhongyicun and Dahai members in eastern Yunnan, the Maidiping Formation in southwestern Sichuan, and the Kuanchuanpu Formation in southern Shaanxi and northern Sichuan. Phosphatebearing strata favord the preservation of SSFs: many originally calcareous SSFs were preserved primarily through secondary phosphatization (Bengtson, 1992; Porter, 2004). The modest late Meishucunian decline in generic diversity not only resulted from the disappearance of SSFs, but was also strongly influenced by taphonomic and environmental factors. In South China, sedimentary strata of this interval are usually composed of dark siliciclastic rocks (black shale or siltstone, e.g., the Shiyantou Formation in eastern Yunnan, the Guojiaba Formation in southern Shaanxi). These dark rocks are not only poor in fossils but also are apparently unfavorable for SSF collection using the acid maceration method. Furthermore, the apparent reduction of phosphate deposits beginning from the late Meishucunian also contributes to the apparent decline in SSF diversity. The Qiongzhusian diversity peak shows that this interval represents the major phase of the “Cambrian Explosion”. Although exceptional preserved fossils of the Chengjiang Biota partly contribute to the high diversity, the generic diversity increase is mainly related to the radiation of arthropods, priapulids and other animals. That SSFs suffered an obvious decline in the Qiongzhusian may be partly related to a significant reduction in phosphogenesis. Fossil evidence from the Chengjiang Lagerstätten indicates that a modern-style marine ecosystem had already been well established by the Qiongzhusian, both in the benthic and pelagic realms (Hu, 2005; Vannier and Chen, 2005). The fossil record also shows that, along with the rapid biodiversification, metazoans progressively exploited more ecological space: most Meishucunian faunas are from shallow sedimentary environments, but during the Qiongzhusian, there was a clear differentiation between shallow and deep water habitats. There was also a strong linkage between the occurrences of skeletal fossils and sedimentary environments (Li et al., 2004; Hu, 2005). The Canglangpuan decline in generic diversity is mainly due to the following two factors. First, lacking major deposits of exceptional preservation, there are only a few genera of non-biomineralized fossils known from Canglangpuan deposits (e.g., Zhang and Hua, 2005). Second, a dramatic decrease in the generic diversity of SSFs (only ca. 13 genera) resulted from 238 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 severe decimation of SSFs and reduction of phosphate deposits. The dramatic generic diversity decline from the late Canglangpuan to the Longwangmiaoan implies that there was an extinct event in the late Canglangpuan. The most distinct disappearance is that of archaeochyaths (Yang et al., 2006). The low generic diversity of the Longwangmiaoan is not only due to the late Canglangpuan extinction but is also related to a eustatic fall recorded in Longwangmiao strata. Sea level falls reduced habitat space on shallow shelves. Sedimentary sequences of this interval are mainly composed of coarse-grained siliciclastic rocks (especially in eastern Yunnan), which are not only unfavorable for nonbiomineralized tissue preservation but also impede the collection of small skeletal fossils by acid maceration. 4.2. Comparison with global patterns The sigmoidal pattern of generic diversity in South China (Fig. 2A) through the Early Cambrian is roughly consistent with global diversity pattern (Fig. 2B) (Zhuravlev, 2001). The stepwise increase in generic diversity through the early and mid-Meishucunian is comparable with the global diversity increase through the Nemakit–Daldynian to early Tommotian interval. Both increases resulted largely from the diversification of SSFs. A secondary peak in generic diversity in the middle Meishucunian is somewhat comparable with the early Tommotian peak on the global diversity curve. In contrast to the Siberian Platform and Mongolia (Zhuravlev and Wood, 1996), there is no archaeocyath record from Meishucunian rocks in South China. The mid Meishucunian peak did not receive loading from archaeocyath fossils. On the Yangtze Platform, archaeocyaths first occur in the Qiongzhusian rocks (Yang et al., 2006). The generic diversity decline through the late Meishucunian to earliest Qiongzhusian interval in South China seems to be comparable with the diversity decrease through the late Tommotian and early Atdabanian interval. The decline in South China is largely due to the widespread deposition of black shales (anoxic environments) and to the beginning of decimaltion of SSFs. Phosphate deposition was also reduced. However, comparison of this decline with the mid-Botomian extinction (Zhuravlev and Wood, 1996) is not well supported by biostratigraphic correlations or faunal components. The major diversity peak of the late Qiongzhusian, when metazoans attained their highest generic diversity of the Cambrian, compares with the diversity peak of the early Botomian. The diversity increase through the Qiongzhusian could be compared with the diversity increase through the late Atdabanian and early Botomian. But unlike the global diversity increase, which got at least 24% loading from archaeocyath genera (Zhuravlev, 2001), the South China increase in generic diversity was partly related to the soft-bodied preservation of the Chengjiang fauna. In South China, archaeocyaths comprise only ca. 6% of total generic diversity, and are not the principal group contributing to the diversity pattern. The distinctive diversity decline through the Canglangpuan–Longwangmiaoan interval is more or less consistent with the global diversity decline through the late Botomian and Toyonian. The Canglangpuan decline in generic diversity is due partly to the soft-bodied fossils of the Chengjiang Biota scarcely occurring in the Canglangpuan rocks and due to a dramatic decline in the generic diversity of the Meishucunian SSFs (only ca. 13 genera). In contrast, trilobites show a rapid increase in generic diversity from 33 during the Qiongzhusian to 91 genera during the Canglangpuan. The decline in the generic diversity is the most dramatic from the Canglangpuan (ca. 153 genera) to the Longwangmiaoan (ca. 69 genera), and all metazoan clades either exhibit a distinct decline in generic diversity or even have no fossil record in Longwangmiaoan rocks. 5. Conclusion Quantitative analysis of Early Cambrian metazoan generic diversity in South China shows the following patterns: an early–mid Meishucunian increase, a modest late Meishucunian decline, a dramatic Qiongzhusian increase, and a prolonged Canglangpuan–Longwangmiaoan decline. The major Cambrian bioradiation mainly occurred through the Meishucunian–Qiongzhusian interval. Metazoan fossils attained their highest generic diversity (ca. 304 genera) of the Cambrian during the Qiongzhusian. The prolonged diversity decline through the Canglangpuan– Longwangmiaon interval was due partly to sea level fall in South China and to decimation of SSFs. The Meishucunian fauna is fairly consistent with the pretrilobitic “Tommotian Fauna” (Sepkoski, 1992). The abrupt occurrence of abundant SSFs in the Meishucunian could be taken as the first phase of the “Cambrian Explosion” (Qian, 1999). The Qiongzhusian fauna is mostly characterized by the occurrence and dominance of pan-arthropods, and by a decline in SSF diversity. Dominance of the pan-arthropods and the occurrence of crown-group brachiopods and echinoderms indicate that the Qiongzhusian fauna represents an initial phase of the Cambrian Evolutionary Fauna (sensu stricto; Sepkoski, 1992). The generic diversity pattern through the Early Cambrian in South China is relatively similar to the G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 global pattern. The secondary peak of generic diversity in the mid Meishucunian of South China is approximately close to the early Tommotian peak in the global pattern. The diversity decline through the late Meishucunian and early Qiongzhusian may be compared with the decline through the late Tommotian and early Atdabanian. Most striking is the major increase in diversity in the Qiongzhusian which may correspond to that in the early Botomian. The diversity decline through the Canglangpuan and Longwangmiaoan is more or less consistent with the global pattern through the late Botomian and Toyonian. Acknowledgements We acknowledge financial support from Chinese Academy of Sciences (KZCX03-SW-141), the National Natural Science Foundation of China (Nos.40572006, 40232020), the Ministry of Science and Technology of China (No.2006CB806401), and the State Key Laboratory of Palaeobiology and Stratigraphy (No.063102). Constructive comments by L. Babcock, G. Shields, M.Y. Zhu and Y. Qian have greatly improved the manuscript. Critical reviews by A. Gubanov and J. Vannier significantly improved this paper. G. X. Li thanks the Max–Planck–Gesellschaft (Munich, Germany) for supporting his work at the Technical University Berlin. Appendix A. Genera of the metazoan fossils reported from the Lower Cambrian of South China This listing is based on references with the published genera of Early Cambrian metazoan fossils from South China. The grouping of some genera into higher taxa is still a subject of active research, although tentatively, most of them are alphabetically listed within higher taxonomic categories herein. Genera occurring in the Chengjiang Biota are preceded by a superscript “c”. Generic names marked by asterisks (⁎) are considered to be junior synonyms of the genera shown in the appressed parentheses, but the generic names of trilobites deemed invalid are not included. Names denoted by rhombs (#) repesenting inappropriate identifications or incorrect assignments, and those marked by question marks (?) representing problematic erections. The generic ranges occurring in the Lower Cambrian of South China are shown in the square brackets. The data of the Chengjiang Biota were compiled by M. Steiner, archaeocyaths by A.H. Yang, trilobites by X.J. Zhu, and the others by G.X. Li. Abbreviations: “E Meis” — early Meishucunian, “M Meis” — middle Meishucunian, “L Meis” — late 239 Meishucunian, “Qiong” — Qiongzhusian, “Cangl” — Canglangpuan, “Longw” — Longwangmiaoan. PORIFERANS Demosponges c Allantospongia Rigby et Hou [Qiong] c Choia Walcott [Qiong] c Choiaella Rigby et Hou [Qiong] c Hazelia Walcott [Qiong] c Leptomitella Rigby [Qiong] c Leptomitus Walcott [Qiong] c Paraleptomitella Chen, Hou et Lu [Qiong] Sanshapentella Mehl et Erdtmann [Qiong] Hexactinellids Asteractinella Hinde [M Meis] ?Bashanites Ding et Li [Qiong] Calcihexatina Sdzuy [M Meis–Qiong] c Crumillospongia Rigby [Qiong] Diagoniella Rauff [Qiong] Gabelia Rigby et Murphy [Qiong] c Halichondrites Dawson [Qiong] Hexatractiella Mehl [Cangl] Hunanospongia Qian et Ding [M Meis–Qiong] c Hyalosinica Mehl et Reitner [Qiong] Lantianospongia Xiao et al. [Qiong] ?Licroites Qin et Li [Qiong] Metaxyspongia Wu et al. [Cangl] Protospongia Salter [M Meis–Qiong] c Quadrolaminiella Chen, Hou et Li [Qiong] Ratcliffespongia Rigby [Cangl] c Saetaspongia Mehl et Reitner [Qiong] Sanshadictya Mehl et Reitner [Qiong] Solactiniella Mehl et Reitner [Qiong] c Takakkawia Walcott [Qiong] c Triticispongia Mehl et Reitner [Qiong] ?Xixiangites Ding et Li [Qiong] Calcareans ⁎Actinoites Duan (=Eiffelia Walcott) Eiffelia Walcott [L Meis–Qiong] ⁎Lenastella Missarzhevsky (=Eiffelia Walcott) ⁎Niphadus Duan (=Eiffelia Walcott) Archaeocyathans Afiacyathus Voronin [Qiong–Cangl] #Agastrocyathus Debrenne (being Zunyicyathus Debrenne, Kruse et Zhang) [Cang] #Ajacicyathus Bedford et Bedford (being Nochoroicyathus Zhuravleva) #Anaptychocyathus Debrenne (being Erismacoscinus Debrenne) Archaeocyathus Billings [Cangl] #Archaeofungia Taylor (being Afiacyathus Voronin or Sibirecyathus Vologdin) Archaeopharetra Bedford et Bedford [Qiong] #Cambrocyathellus Zhuravleva (being Graphoscyphia Debrenne et Zhuravleva or Changicyathus Debrenne et Zhuravlev) Changicyathus Debrenne et Zhuravlev [Qiong] Chengkoucyathus Yuan [Qiong–Cangl] Chouberticyathus Debrenne [Qiong] Clathricoscinus Zhuravleva [Cangl] Conannulofungia Yuan [Cangl] #Coscinocyathus Bornemann (being Erismacoscinus Debrenne) 240 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 Dailycyathus Debrenne [Cangl] Densocyathus Vologdin [Cangl] Dictyocyathus Bornemann [Qiong–Cangl] Erismacoscinus Debrenne [Qiong–Cangl] Erugatocyathus Debrenne [Cangl] Graphoscyphia Debrenne et Zhuravleva [Qiong] Ichnusocyathus Debrenne [Cangl] Inessocyathus Debrenne [Cangl] Khirgisocyathus Voronin [Qiong] Leptosocyathus Vologdin [Qiong] Metacyathellus Debrenne et Zhuravlev [Cangl] Nochoroicyathus Zhuravleva [Qiong] #Okulitchicyathus Zhuravleva (being Graphoscyphia Debrenne et Zhuravleva) Pilodicoscinus Debrenne et Jang [Qiong] #Porocoscinus Debrenne (being Chengkoucyathus Yuan) Protopharetra Bornemann [Qiong–Cangl] #Rasetticyathus Debrenne (being Dailycyathus Debrenne) ⁎Retecyathus Vologdin (=Archaeocyathus Billings) Robustocyathellus Konyushkov [Qiong] #Rotundocyathus Vologdin (being Dailycyathus Debrenne or Robustocyathellus Konyushkov) Rudanulus Debrenne [Cangl] ⁎Sanxiacyathus Yuan et Zhang (=Archaeocyathus Billings) Sibirecyathus Vologdin [Cangl] Spirillicyathus Bedford et Bedford [Cangl] Stillicidocyathus Ting [Cangl] Thalamocyathus Gordon [Qiong] Taylorcyathus Vologdin [Qiong] Taylorfungia Perejon [Qiong] #Tumuliolynthus Zhuravleva (none archaeocyaths, being Acanthocassis He et Xie) [M Meis] Yhecyathus Belyaeva et Kesin [Qiong] Zunyicyathus Debrenne, Kruse et Zhang [Cangl] Problematic sponges Blastulospongia Pickett et Jell [Qiong] c Sinoflabrum Zhang et Babcock [Cangl] CNIDARIANS ⁎Archaeotuba Hou et al. (=Cambrorhytium Conway Morris et Robinson) Byronia Matthew [Qiong–Longw] c Cambrohydra Hu [Qiong] c Cambrorhytium Conway Morris et Robinson [Qiong–Cangl] Cambrovitus Mao et al. [Longw] ⁎Dentachites He et Lin(=Punctatus He) c Priscapennamarina Zhang et Babcock [Qiong] Punctatus He [EM Meis] ⁎Pyrgites Yue (=Punctatus He) Sphenothallus Hall [Qiong–Longw] c Xianguangia Chen et Erdtmann [Qiong] Anabaritids ⁎Anabaritellus Missarzhevsky (=Anabarites Missarzhevsky) Anabarites Missarzhevsky [Meis] Cambrotubulus Missarzhevsky [M Meis–Qiong] ⁎Lobiochrea Val'kov et Sysoev (=Anabarites Missarzhevsky ) Tiksitheca Missarzhevsky [M Meis–Qiong] Conulariids ⁎Aciconularia He (=Carinachites Qian) Arthrochites Chen [E M Meis] ⁎Barbitositheca Qian et Jiang (=Arthrochites Chen) Carinachites Qian [Meis] #Conularia Miller (being Carinachites Qian) ⁎Conulariella He et Yang (=Hexangulaconularia He) Emeiconularia Qian et al. [Meis] #Eoconularia Sinclair (being Carinachites Qian) ⁎Hexaconularia Yang, He et Deng (nomen nudum) (=Hexangulaconularia He) Hexangulaconularia He [E M Meis] ⁎Lagenaconularia He (=Arthrochites Chen) ⁎Mabianoconullus He (=Carinachites Qian) ⁎Paraconularoides He (=Carinachites Qian) ⁎Paranabarites Jiang (=Carinachites Qian) ⁎Primaconulariella He (=Arthrochites Chen) ⁎Quadrosiphogonuchites Chen (=Carinachites Qian) ⁎Sacciella He (=Arthrochites Chen) CTENOPHORANS Batofasciculus Hou et al. [Qiong] c Maotianshanoascus Chen et Zhou [Qiong] c Sinoascus Chen et Zhou [Qiong] c Trigoides Luo et Hu [Qiong] c ANNELIDS Facivermis Hou et Chen [Qiong] c Maotianchaeta Chen [Qiong] c HYOLITHS Hyolithids Aimitus Sysoev [Qiong–Cangl] c Ambrolinevitus Sysoev [L Meis–Qiong] Biligulitheca Qian, Xie et He [Qiong] c Burithes Missarzhevsky [L Meis–Qiong] Dipterygovitus Qian [Qiong] Doliutus Missarzhevsky et Sysoev [Qiong] Eolinevitus Qian, Zhu et Jiang [Qiong] ⁎Glossolites Luo et Hu (=Burithes Missarzhevsky) Haplophrenits Babcock et Robison [Longw] Hyolithes Eichwald [Longw?] Inflatatheca Qian, Xie et He [Qiong] c Linevitus Sysoev [Qiong–Cangl] Microcornus Mambetov [Qiong] Ningqiangethus Qian, Xie et He [Qiong] Paramicrocornus Qian, Xie et He [Qiong] Quadrotheca Sysoev [M Meis] Sulcavitus Sysoev [Qiong] ?Trypanovitus Qian [Qiong] Xixiangethes Qian, Xie et He [Qiong] Yankongovitus Qian [Cangl] Orthothecids Adyshevitheca Mambetov [Qiong] Allatheca Missarzhevsky [M Meis–Cangl] Ancheilotheca Qian [M Meis] Aritstitheca Jiang [Cangl] #Circotheca Sysoev (being Conotheca Missarzhevsky) Coleolella Missarzhevsky [M Meis] Conotheca Missarzhevsky [E Meis–Cangl] Doliutheca Qian [M Meis–Qiong] Eogloborilus Qian [M Meis] Eonovitatus Sysoev [L Meis] Exilitheca Sysoev [Cangl] Globoritubulus Qian et Zhang [M Meis–Qiong] G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 ?Gyrazonatheca Qian [M Meis] Kunyangotheca Qian [E Meis] Laratheca Missarzhevsky [M Meis] ?Leconogloborilus Qian et Jiang (nomen nudum) [M Meis] Leibotheca Qian [M Meis] Lenatheca Missarzhevsky [M Meis] Loculitheca Sysoev [M Meis] Majatheca Missarzhevsky [Qiong] ?Malongtheca Jiang [Cangl] Neogloborilus Qian et Zhang [Qiong] Ovalitheca Sysoev [M Meis–Qiong] ?Paracircotheca Qian [Qiong] Paraeonovitus Qian [L Meis] ⁎Persicitheca Duan (=Neogloborilus Qian et Zhang) Platydorsotus Qian [M Meis] Spinulitheca Sysoev [E M Meis] ⁎Stimulitheca Duan (=Neogloborilus Qian et Zhang) Sulcagloborilus Jiang [M Meis] Turcutheca Missarzhevsky [Meis] ?Xiadongtubulus Qian et Zhang [M Meis] MOLLUSKS and MOLLUSK-LIKE FOSSILS Nonbivalves (originally described as bivalves) ?Cycloconchoides Zhang (being obolellids) [Cangl] ?Hubeinella Zhang (being obolellids) [Cangl] ?Praelamellodonta Zhang (being obolellids) [Cangl] ?Xianfengoconcha Zhang (being obolellids) [Qiong–Cangl] Rostroconchs ⁎Heraultipegma Pojeta et Runnegar (=Watsonella Grabau) Watsonella Grabau [M L Meis] Helcionellids and helcionellid-like fossils Absidaticonus Yue [M Meis] Acutirostriconus Yue [M Meis] Aldanella Vostokova [M L Meis] Anabarella Vostokova [M Meis] Anhuiconus Zhou et Xiao [Cang] Archaeospira Yu [M Meis] ⁎Asperoconus Yu (=Protoconus Yu) Bemella Missarzhevsky [M Meis] Beshtashella Missarzhevsky [Qiong] Cambroconus Yu [M Meis] ⁎Ceratoconus Chen et Zhang (=Pollicina Holzapfel) Coreospira Saito [Longw] ?Dianella Wang [M Meis] Eosoconus Yu [M Meis] ?Gibbaspira He [M Meis] ⁎Ginella Missarzhevsky (=Ilsanella Missarzhevsky) ?Gutticonus Yue [M Meis] c Helcionella Grabau et Shimer [M Meis–Longw] ⁎Huanglingella Chen, Chen et Zhang (=Bemella Missarzhevsky) Huangshandongoconus Yu [M Meis] Hubeispira Yu [M Meis] Hujiagouella Chen et Zhang [M Meis] Igorella Missarzhevsky [M Meis–Qiong] Ilsanella Missarzhevsky [M L Meis] Latirostratus Yu [M Meis] Latouchella Cobbold [Cangl] #Maclurites Lesueur [M Meis] Maidipingoconus Yu [M Meis] ?Masculuconus Feng, Sun et Qian [M Meis] Obtusoconus Yu [M Meis–Qiong] Oelandiella Vostokova [M Meis] Palaeacmaea Hall et Whitfield [M L Meis] ?Papilloconus Feng, Sun et Qian [M Meis] ⁎Paraformichella Qian et Zhang (=Beshtashella Missarzhevsky) Pelagiella Matthew [Qiong] Planuspira Jiang [M Meis] Pollicina Holzapfel [M Meis] Protoconus Yu [M Meis] ⁎Rostroconus Jiang (=Latirostratus Yu) ⁎Sacciconus Yu (=Bemella Missarzhevsky) ⁎Scamboscamna Liu (=Scenella Billings) c Scenella Billings [M Meis–Longw] ?Scutatestomaconus Chen et Zhang [M Meis] Sichuanospira He [M Meis] ⁎Songlinella Chen, Chen et Zhang (not Songlinella Yin) (=Archaeospira Yu) Stenotheca Hicks [M Meis–Qiong] Tannuella Missarzhevsky [M Meis] Tuberoconus Zhou et Xiao [M Meis] Uncinaspira He [M Meis] Xianfengella He et Yang [M Meis] Xilingxiaconus Chen et Zhang [M Meis] Yangtzeconus Yu [M Meis] ⁎Yangtzedonta Yu (=Xianfengella He et Yang) ⁎Yangtzespira Yu (=Archaeospira Yu) Yochelcionella Runnegar et Pojeta [Qiong] ⁎Yunnanospira Jiang (=Archaeospira Yu) Problematic mollusks ?Actinoconus Yu [M Meis] Aegides Jiang [M Meis] ⁎Aegitellus Feng, Sun et Qian (=Aegides Jiang) ⁎Astropolites He et Yang (=Maikhanella Zhegallo) ⁎Canopoconus Jiang (=Maikhanella Zhegallo) ⁎Cassidina Jiang (=Maikhanella Zhegallo) ?Centriconus Yu [M Meis] ⁎Chengjiangoconus He et Yang (=Ocruranus Liu) Codonoconus Chen et Zhang [M Meis] ⁎Cremnodinotus Liu (=Eohalobia Jiang) ?Crestoconus Jiang [M Meis] ?Dengyingoconus Chen et Xiong [M Meis] ⁎Diandongoconus He et Yang (=Emarginoconus Yu) ?Dolichomocelypha Liu [M Meis] ?Dysnoetopla Liu [M Meis] Ebianella He et Lin [L Meis] Emarginoconus Yu [M Meis] ⁎Emeiconus He et Yang (=Purella Missarzhevsky) Eohalobia Jiang [M Meis] #Gotlandochiton Bergenhayn (being Ocruranus Liu) Granoconus Yu [M Meis] ?Hamatoconus Chen et Xiong [M Meis] ⁎Hanshuiella Yue (=Ocruranus Liu) ?Heosomocelypha Liu [M Meis] ⁎Jinkenites Yu (=Maikhanella Zhegallo) ?Kuanchuanella Yue [M Meis] ?Laticonus Yu [M Meis] ⁎Lepidites Zhong (Chen) (=Maikhanella Zhegallo) ⁎Liantuoconus Yu (=Spatuloconus Yu) ?Ligyrokala Liu [M Meis] ⁎Liorichita Liu (=Ocruranus Liu) Maikhanella Zhegallo [Meis] 241 242 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 Maishucunconus Jiang [M Meis] ⁎Mediata Feng, Sun et Qian (=Maikhanella Zhegallo) ⁎Meishucunchiton Yu (=Eohalobia Jiang) Merismoconcha Yu [M Meis] ⁎Microlites Yang, He et Deng (nomen nudum) (=Maikhanella Zhegallo) Minymerisma Yu [M Meis] ⁎Mirusilites Yang, He et Deng (nomen nudum) (=Maikhanella Zhegallo) Ocruranus Liu [M Meis] ?Omalenlina Liu [M Meis] Parascenella Chen et Xiong [M Meis] ?Pileconus Jiang [M Meis] ⁎Poratites Jiang (=Maikhanella Zhegallo) Postacanthella Yue [M Meis] ⁎Postestephaconus Jiang (=Ocruranus Liu) ⁎Protosyringolites He et Yang (=Maikhanella Zhegallo) Purella Missarzhevsky [M Meis] ⁎Ramenta Jiang (=Maikhanella Zhegallo) ⁎Ramentoides Feng, Sun et Qian (=Maikhanella Zhegallo) ⁎Runnegarochiton Yu (=Ocruranus Liu) Securiconus Jiang [M Meis] Sinuconus Yu [M Meis] Spatuloconus Yu [M Meis] ⁎Stephaconus Jiang (nomen nudum) (=Ocruranus Liu) ⁎Stoliconus Jiang (=Ocruranus Liu) Tianzhushanospira Yu [M Meis] Truncatoconus Yu [M Meis] ⁎Tubatoconus Feng, Sun et Qian (=Ocruranus Liu) Xiadongoconus Yu [M Meis] Yangtzemerisma Yu [M Meis] ⁎Yuanjiapingella Yue (=Ocruranus Liu) ?Yunnanoconus Jiang (nomen nudum) [M Meis] Yunnanopleura Yu [M Meis] Mobergellans Mobergella Hedström [M L Meis] Shells ?Archaeotremaria Yu [M Meis] Aviculocephaloconus Chen et Zhang [M Meis] Stictoconus Qian et Bengtson [M Meis] ⁎Striatoconus Feng, Sun et Qian (=Stictoconus Qian et Bengtson) COELOSCLERITOPHORANS Chancelloriids ⁎Adversella Jiang (?=Allonnia Dore et Reid) c Allonnia Dore et Reid [L Meis–Qiong] Archiasterella Sdzuy [L Meis–Qiong] c Chancelloria Walcott [M Meis–Longw] ⁎Dimidia Jiang (=Allonnia Dore et Reid) ⁎Fangxianites Duan (=Chancelloria Walcott) ⁎Onychia Jiang (=Allonnia Dore et Reid) Halkieriids ⁎Acuminachites Qian et Yin (=Halkieria Poulsen) ⁎Dactyosachites He (=Halkieria Poulsen) Halkieria Poulsen [M Meis–Qiong] ⁎Microsachites He (=Halkieria Poulsen) ⁎Sachitellus (=Halkieria Poulsen) Sinosachites He [L Meis] ⁎Tianzhushania Qian, Chen et Chen (=Halkieria Poulsen) Wiwaxia Matthew [Longw] Sachitids Sachites Meshkova [M L Meis] Siphogonuchitids ⁎Aurisella Qian et Xiao (=Ninella Missarzhevsky) ⁎Dabashanites Chen (=Siphogonuchites Qian) ⁎Disulcavichites Qian et Jiang (nomen nudum) (=Siphogonuchites Qian) Drepanochites Qian et Jiang [M Meis] ⁎Lepochites Zhong (Chen) (nomen nudum) (=Lopochites Qian) Lomasulcachites Qian et Jiang [M Meis] Lopochites Qian [M Meis] Ninella Missarzhevsky [Qiong] ⁎Palaeosulcachites Qian (=Siphogonuchites Qian) Siphogonuchites Qian [M Meis] ⁎Trapezochites Qian et Jiang (=Siphogonuchites Qian) BRACHIOPODS Botsfordia Matthew [Qiong] ⁎Brachiovermis Shu, Chen et Zhang (nomen nudum) (=Lingulella Salter) c Diandongia Rong [Qiong–Cangl] Eohadrotreta Li et Holmer [Qiong] Eoobolus Matthew [Qiong] c Heliomedusa Sun et Hou [Qiong] Kutorgina Billings [Longw] Kyrshabaktella Koneva [Qiong] c Lingulella Salter [Qiong–Cangl] c Lingulellotreta Koneva [Qiong–Cangl] #Lingulepis Hall (being Lingulellotreta Koneva) Linnarssonia Walcott [Longw] c Longtancunella Hou et al. [Qiong] Nisusia Walcott [Longw] Obolus Eichwald [Qiong–Cangl] Palaeobolus Matthew [Qiong] ⁎Paleolingula Shu, Chen et Zhang (nomen nudum) (=Lingulella Salter) Paterina Beecher [Qiong] Westonia Walcott [Cangl] c Xianshanella Zhang et Han [Qiong] Stem group brachiopods Lathamella Liu [M L Meis] Tianzhushanella Liu [M Meis] Problematic brachiopods Acidotocarena Liu [M Meis] ?Artimyctella Liu [M Meis] Ernogia Jiang [M Meis] ⁎Parapunctella Jiang (=Ernogia Jiang) ?Plicatolingula Liu [M Meis] ?Protobolus Liu [M Meis] ?Psamathopalass Liu [M Meis] ?Punctella Zhong (Chen) (nomen nudum) [M Meis] ⁎Scambocris Liu (=Acidotocarena Liu) c PHORONIDS Eophoronis Chen (nomen nudum) [Qiong] c Iotuba Chen et Zhou [Qiong] c SIPUNCULANS Archaeogolfingia Huang et al. [Qiong] c Cambrosipunculus Huang et al. [Qiong] c PRIAPULIDS AND OTHER WORMS Acosmia Chen et Zhou [Qiong] c Anningvermis Huang, Vannier et Chen [Qiong] c G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 c Corynetis Luo et Hu [Qiong] Cricocosmia Hou et Sun [Qiong] ? cGangtoucunia Luo et Hu [Cangl] Goettingenia Zhang et Hua [Cangl] Hadimopanella Gedik [Cangl] Houscolex Zhang et Pratt [Qiong] ? cLagenula Luo et Hu [Qiong] c Mafangscolex Hu [Qiong] c Maotianshania Sun et Hou [Qiong–Cangl] ? cOligonodus Luo et Hu [Qiong] ⁎Palaeopriapulites Hou et al. (?=Sicyophorus Luo et Hu) c Palaeoscolex Whittard [Qiong] c Paraselkirkia Hou et al. [Qiong] ⁎Protopriapulites Hou et al. (=Sicyophorus Luo et Hu) c Sandaokania Luo et Hu [Qiong] c Sicyophorus Luo et Hu [Qiong] c Tabelliscolex Han et al. [Qiong] c Tylotites Luo et Hu [Qiong] c Xiaoheiqingella Hu [Qiong] c Xishania Hu [Qiong] c Yunnanopriapulus Huang, Vannier et Chen [Qiong] c CHAETOGNATHS ⁎Eognathacantha Chen et Huang (=Protosagitta Hu) c Protosagitta Hu [Qiong] Protoconodonts Amphigeisina Bengtson [Qiong] ⁎Emeidus Chen (=Protohertzina Missarzhevsky) ⁎Ganloudina He (=Mongolodus Missarzhevsky) Gapparodus Abaimova [Qiong] Hagionella Xie [Qiong] ⁎Hastina Yang et He (=Protohertzina Missarzhevsky) Hertzina Müller [M Meis–Qiong] Mongolodus Missarzhevsky [M Meis] Oneotodus Lindström [M Meis] Protohertzina Missarzhevsky [E M Meis] LOBOPODIANS Cardiodictyon Hou, Ramsköld et Bergström [Qiong] ⁎Eoconcharium Hao et Shu (=Microdictyon Bengtson et al.) Fusuconcharium Hao et Shu Qiong] c Hallucigenia Conway Morris [Qiong] c Luolishania Hou et Chen [Qiong] c Megadictyon Luo et Hu [Qiong] c Microdictyon Bengtson et al. [Qiong] c Miraluolishania Liu et al. [Qiong] c Onychodictyon Hou, Ramsköld et Bergström [Qiong] c Paucipodia Chen, Zhou et Ramsköld [Qiong] Quadratapora Hao et Shu [Qiong] c ANOMALOCARIDIDS Amplectobelua Hou, Bergström et Ahlberg [Qiong] c Anomalocaris Whiteaves [Qiong] c Cucumericrus Hou, Bergström et Ahlberg [Qiong] c Parapeytoia Hou, Bergström et Ahlberg [Qiong] c ARTHROPODS Bradoriids #Aluta (Matthew) (being Liangshanella Huo) Alutella Kobayashi et Kato [Qiong–Cangl] ⁎Anaulaca Shu et Cui (=Tsunyiella Chang) ⁎Antihipponicharion Huo et Shu (=Shangsiella Lee) Auriculatella Tan [Qiong] ⁎Bajiella Jiang (=Nanchengella Huo) ⁎Bicostatella Tan et Li (=Kunmingella Huo) #Bradoria Matthew (being Kunmingella Huo) #Cambria Neckaja et Ivanova (being Shangsiella Lee) Changshabaella Huo et Shu [Qiong] ⁎Chuanbeiella Huo et Peng (=Shangsiella Lee) c Comptaluta Öpik [Qiong] Dahaiella Jiang [Cangl] Dahebaella Zhao et Shu [Qiong] Emeiella Lee [Qiong] ⁎Emeillopsis Huo et Shu (=Emeiella Lee) ?Eotuzoia Shu [Qiong] ⁎Gaoqiaoella Lee (=Tsunyiella Chang) Guangyuanella Chang [Qiong] ?Guizhouella Shu [Qiong] Hanchiangella Huo [Qiong] ⁎Hanchungella Huo (=Liangshanella Huo) Houlongdongella Lee [Qiong–Cangl] ⁎Jinningella Huo et Shu (=Sunella Huo) c Jiucunella Hou et Bergström [Qiong] Jixinlingella Lee [Qiong] c Kunmingella Huo [Qiong] ⁎Kunmingelloides Tong (=Kunmingella Huo) c Kunyangella Huo [Qiong] ⁎Laogongshania Li (=Nanchengella Huo) ⁎Leshanella Li (=Comptaluta Öpik) c Liangshanella Huo [Cangl] Luella Huo [Qiong] ⁎Majiashanella Lin (=Tsunyiella Chang) ⁎Malongella Chang (=Kunmingella Huo) Meishucunella Jiang [Qiong] Mononotella Ulrich et Bassler [Qiong–Cangl] Nanchengella Huo [Qiong] ⁎Neohanchungella Cui (=Liangshanella Huo) Neokunmingella Chang [Qiong–Cangl] Ningqiangella Huo et Shu [Qiong] ⁎Ovaluta Zhang (=Liangshanella Huo) ⁎Paracambria Huo et Shu (=Shangsiella Lee) ⁎Parahipponicharion Shu (=Neokunmingella Chang) ⁎Parakunmingella Chang (=Kunmingella Huo) ⁎Pengshuiella Cui (=Liangshanella Huo] Phasoia Hinze-Schallreuter [Qiong] ⁎Pseudobeyrichona Shu (=Neokunmingella Chang) ⁎Pseudobicostatella Tan et Li (=Kunmingella Huo) ?Pseudodahebaella Shu [Qiong] ⁎Pseudokunmingella Huo et Shu (=Kunmingella Huo) ⁎Qingquanella Cui et Huo (=Houlongdongella Lee) Shangsiella Lee [Qiong] ⁎Shensiella Huo (=Hanchiangella Huo) ⁎Songlinella Yin (=Emeiella Lee) c Spinokunmingella Huo et Shu [Qiong] ⁎Sulcatella Tan et Li (=Tsunyiella Chang) c Sunella Huo [Qiong] c Tsunyiella Chang [Qiong–Cangl] ⁎Varitsunyiella Cui et Huo (=Tsunyiella Chang) Wuchiapingella Huo [Qiong] c Wutingella Chang [Qiong] ?Xingzishanella Tan et Li [Qiong] 243 244 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 ⁎Yaoyingella Chang (=Comptaluta Öpik) Yeshanella Lin [Cangl] Zhenpingella Li [Qiong–Cangl] ?Zhijinella Yin [Qiong] Zhongbaoella Huo et Shu [Cangl] Phosphatocopids Dabashanella Huo, Shu et Fu [Qiong–Cangl] ⁎Eohesslandona Shu (=Dabashanella Huo, Shu et Fu) #Flemingopsis Jones et McKenzie (being Dabashanella Huo, Shu et Fu) ⁎Jingyangella Tan et Li (=Dabashanella Huo, Shu et Fu) ⁎Paradabashanella Shu (=Dabashanella Huo, Shu et Fu) ⁎Paranaviformella Tan et Li (=Dabashanella Huo, Shu et Fu) ⁎Paraphaseolella Tong (=Dabashanella Huo, Shu et Fu) ⁎Phaseolella Zhang (=Dabashanella Huo, Shu et Fu) ⁎Pseudindiana Zhao (=Dabashanella Huo, Shu et Fu) ⁎Pseudodabashanella Shu (=Dabashanella Huo, Shu et Fu) ⁎Shannanella Tan et Li (=Dabashanella Huo, Shu et Fu) ⁎Tricostatella Tan et Li (=Dabashanella Huo, Shu et Fu ) ⁎Xiaoyangbaella Huo, Shu et Fu (=Dabashanella Huo, Shu et Fu) ⁎Zhenbaella Tan et Li (=Dabashanella Huo, Shu et Fu) Trilobites Acanthomicmacca Hupe [Cangl] Anadoxides Matthew [Qiong] Antagmus Resser [Cangl–Longw] Arthricocephalus Bergeron [Cangl] Balangcunaspis Yuan et Zhao [Longw] Balangia Qian [Cangl] Barklyella Shergold [Longw] Bathynotus Hall [Longw] Bergeroniaspis Lermontova [Cangl] Binodaspis Lermontova [Cangl–Longw] Bonnia Walcott [Cangl] Breviredlichia Zhang et Lin [Cangl] Changaspis Lee [Cangl] Changyangia Chang [Qiong–Cangl] Chaoaspis Chang [Qiong] Cheiruroides Kobayashi [Cangl–Longw] c Chengjiangaspis Zhang et Lin [Qiong] Chengkouaspis Zhang et Lin [Cangl] Chengkouia Chien et Yao [Cangl] Chittidilla King [Longw] Chuchiaspis Chang [Longw] Corynexochus Angeln [Longw] Crassifimbra Lochman [Cangl] Danzhaina Yuan [Longw] Dicerodiscus Chang [Cangl] Dinesus Etheridge [Longw] Dipharus Clark [Cangl] Drepanopyge Lu [Cangl] Drepanuroides Chang [Cangl] Duodingia Zhou [Qiong] Duyunaspis Chang et Chien [Cangl] Eodiscus Hartt [Cangl] Eodontopleura Qian et Lin [Cangl] Eodouposiella Yuan et Zhao [Longw] Eokaotaia Yuan et Zhao [Long] Eomalungia Li[Cangl] Eoptychoparia Rasetti [Cangl–Longw] c Eoredlichia Chang [Qiong] Eosoptychoparia Yuan [Longw] Erzishania Zhu et Qian [Cangl] Euarthricocephalus Ju [Longw] Fandianaspis Li [Cangl] Feilongshania Qian et Lin [Cangl] Fuminaspis Zhang et Lin [Qiong] Gaotanaspis Zhang et Li [Longw] Gedongaspis Yuan et Zhao [Longw] Guangyuanaspis Chang et Qian [Qiong] Hoffetella Hupe [Longw] Holocephalina Salter [Longw] Hongjunshaoia Luo [Cangl] Hongshiyanaspis Zhang et Lin [Qiong] Hsüaspis Chang [Cangl] Hunanocephalus Lee [Qiong–Cangl] Hupeidiscus Chang [Qiong–Cangl] Ichangia Chang [Cangl] Jingyangia Chang et Zhang [Qiong] Kermanella Wolfart [Longw] Kootenia Walcott [Cangl–Longw] c Kuanyangia Hupe [Qiong–Cangl] Kueichowia Lu [Cangl] Kunmingaspis Chang [Longw] Longduia Chang et Chien [Cangl] Malongocephalus Zhang et Lin [Qiong] c Malungia Lu [Qiong–Cangl] Maopingaspis Lin et Yin [Cangl] Mayiella Chang [Cangl] Megapalaeolenus Chang [Cangl] Meitanella Lin et Yin [Cangl] Mengzia Lu [Longw] Metaredlichia Lu [Qiong–Cangl] Metaredlichioides Chien et Yao [Cangl] Micangshania Zhou [Qiong] Microryctocara Sundberg et McCollum [Longw] Mufushania Lin [Longw] Nangaoia Zhou [Longw] Nangaops Yuan et Sun [Longw] Neocobboldia Rasetti [Cangl] Ningqiangaspis Zhang et Lin[Qiong] Olenoides Meek [Longw] Oryctocephalites Resser [Longw] Oryctocephalops Lermontova [Longw] Ovatoryctocara Tchernysheva [Longw] Pachyredlichia Chang [Qiong] Pagetia Walcott [Cangl–Longw] Palaeolenus Mansuy [Cangl] Panxinella Lin [Longw] Paokannia Ho et Lee [Cangl] Parabadiella Chang [Qiong] Parachangaspis Liu [Cangl] Paragraulos Lu [Longw] Paraichangia Li [Cangl] Paramalungia Chang [Cangl] Paramecephalus Zhou et Yin [Longw] Parapaokannia Zhang et Lin [Cangl] Paraperiomma Zhou [Longw] Paraprotolenella Zhu et Jiang [Cangl] Parashuiyuella Yuan et Zhao [Longw] Parayiliangella Luo [Cangl] Parayinites Zhang et Lin [Cangl] Pianaspis Saito et Sakakura [Longw] G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 Poulsenia Resser [Longw] Probowmania Kobayashi [Longw] Proichangia Zhang et Zhu [Qiong] Protolenella Chien et Yao [Cangl] Protoryctocephalus Zhou [Longw] Pseudichangia Chu et Zhou [Cangl] Pseudopaokania Yin [Cangl] Pseudoredlichia Zhang et Lin [Cangl] Qiannangraulos Yuan et Zhao [Longw] Qiaodiella Zhang, Li et Zhou [Cangl] Qiaotingaspis Chang [Cangl] Qingkouia Zhang, Lin et Zhou [Qiong–Cangl] Qingzhenaspis Yin [Cangl] Redlichia Cossmann [Cangl–Longw] Resserops Richter et Richter [Cangl] Runnania Lee [Qiong] Saimixiella Li [Cangl] Sanwania Yuan [Longw] Schmalenseeia Moberg [Longw] Shaanxia Zhang et Lin [Qiong] Shabaella Chien et Sun [Cangl] Shatania Chang et Lin [Cangl] Shifangia Chien et Yao [Cangl] Shipaiella Zhang et Qian [Cangl] Shiqihepsis Chien et Yao [Cangl] Sichuanolenus Zhang et Zhu [Cangl] Sinodiscus Chang [Qiong–Cangl] Sinoschistometopus Yuan et Zhao [Longw] Syndianella Lu [Cangl] Syspacephalus Resser [Longw] Szechuanaspis Chien et Yao [Qiong–Cangl] c Tsunyidiscus Chang [Qiong] Ushbaspis Pokrovskaya [Cangl] Wangzishia Sun [Qiong] Wenganaspis Yin [Cangl] Wenganella Yin [Cangl] c Wutingaspis Kobayashi [Qiong] Wuxunaspis Yuan [Longw] Xiangqianaspis Zhou [Cangl–Longw] Xiaomajiella Zhou [Longw] Xilingxia Lu [Cangl–Longw] Xingzishania Zhou[Qiong] Xiuqiella Chien et Yao [Cangl] Yankongia Zhou [Cangl] Yeshanaspis Zhu et Jiang [Cangl] Yiliangaspis Luo [Cangl–Longw] Yiliangella Chang [Cangl] Yiliangellina Chang [Cangl] Yinites Lu [Qiong–Cangl] Yuehsienszella Chang [Cangl–Longw] Yunnanaspidella Chang [Cangl] Yunnanaspis Chang [Cangl] c Yunnanocephalus Kobayashi [Qiong] Zhangshania Li et Zhang [Cangl] Zhenbaspis Chang et Chu [Qiong–Cangl] Zhenpingaspis [Cangl–Longw?] Zhuxiella Zhang et Zhu [Cangl] Other arthropods c Acanthomeridion Hou, Chen et Lu [Qiong] ⁎Almenia Hou et Bergström (=Cindarella Chen et al.) c Apiocephalus Luo et Hu [Qiong] 245 c Branchiocaris Briggs [Qiong–Cangl] ⁎Cambrofengia Hou et al. (=Chengjiangocaris Hou et Bergström) c Canadaspis Novozilov [Qiong–Cangl] Chengjiangocaris Hou et Bergström [Qiong] ⁎Chuandianella Hou et Bergström (=Waptia Walcott) c Cindarella Chen et al. [Qiong] c Clypecaris Hou [Qiong] c Combinivalvula Hou [Qiong] c Cyathocephalus Luo et Hu [Qiong] ⁎Cymbia Jiang (=Isoxys Walcott) c Dianchia Luo et Hu [Qiong] c Diplopyge Luo et Hu [Qiong] c Dongshanocaris Hou et al. [Qiong] ⁎Eonaraoia Zhang et Shu (=Naraoia Walcott) c Ercaia Chen, Vannier et Huang [Qiong] ⁎Ercaicunia Luo et Hu (?=Clypecaris Hou) c Forfexicaris Hou [Qiong] c Fortiforceps Hou et Bergström [Qiong] c Fuxianhuia Hou [Qiong] c Glossocaris Luo et Hu [ Qiong] c Haikoucaris Chen et al. [Qiong] c Isoxys Walcott [Qiong–Cangl] c Jianfengia Hou [Qiong] c Jianshania Luo et Hu [Qiong] c Kuamaia Hou [Qiong] c Kunmingocaris Luo et Hu [Qiong] c Leanchoilia Walcott [Qiong] c Liangwangshania Chen [Qiong] c Mafangia Luo et Hu [Qiong] c Mafangocaris Luo et Hu [Qiong] ⁎Misszhouia Chen, Edgecombe et Ramsköld (=Naraoia Walcott) c Naraoia Walcott [Qiong–Cangl] c Occacaris Hou [Qiong] #Odaraia Walcott (=Pectocaris Hou) c Ovalicephalus Luo et Hu [Qiong] c Parapaleomerus Hou et al. [Qiong] c Pectocaris Hou [Qiong] c Perspicaris Briggs [Qiong] c Pisinnocaris Hou et Bergström [Qiong] Primicaris Zhang et al. [Qiong] c Pseudoiulia Hou et Bergström [Qiong] ⁎Pseudonaraoia Zhang et Shu (=Naraoia Walcott) c Pterotrum Luo et Hu [Qiong] c Pygmaelypeatus Zhang, Han et Shu [Qiong] c Retifacies Hou, Chen et Lu [Qiong] c Rhombicalvaria Hou [Qiong] c Saperion Hou, Ramsköld et Bergström [Qiong] c Shankouia Chen (nomen nudum) [Qiong] c Sidneyia Walcott1 [Qiong] c Sinoburius Hou, Ramsköld et Bergström [Qiong] c Skioldia Hou et Bergström [Qiong] c Squamacula Hou et Bergström [Qiong] ? cSyrrhaptis Luo et Hu [Qiong] c Tanglangia Luo et Hu [Qiong] c Tuzoia Walcott [Qiong–Cangl] c Urokodia Hou, Chen et Lu [Qiong] c Waptia Walcott [Qiong] c Xandarella Hou, Ramsköld et Bergström [Qiong] #Yohoia Walcott (being Leanchoilia Walcott) ⁎Yiliangocaris Luo et Hu (=Canadaspis Novozilov) c 246 G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 c Yunnanocaris Hou [Qiong] ⁎Zhongxinia Luo et Hu (=Leanchoilia Walcott) ECHINODERMS ⁎Dianchicystis Shu et al. (=Ventulocystis Shu et al.) Liyongella Qin et Li [Qiong] Sinoeocrinus Zhao [Longw] c Ventulocystis Shu et al. [Qiong] VETULICOLIANS and CHORDATES Banffia Walcott [Qiong] c Cathaymyrus Shu , Conway Morris et Zhang [Qiong] c Didazoon Shu et Han [Qiong] c Haikouella Chen, Huang et Li [Qiong] c Haikouichthys Luo et Hu [Qiong] ⁎Heteromorphus Luo et Hu (=Banffia Walcott) c Myllokunmingia Shu, Zhang et Han [Qiong] c Pomatrum Luo et Hu [Qiong] c Vetulicola Hou [Qiong–Cangl] c Xidazoon Shu, Conway Morris et Zhang [Qiong] c Yunnanozoon Hou, Ramsköld et Bergström [Qiong] c Yuyuanozoon Chen, Feng et Zhu [Qiong] ? cZhongxiniscus Luo et Hu [Qiong] c Zhongjianichthys Shu [Qiong] c ENIGMATIC METAZOANS FROM THE CHENGJIANG LAGERSTÄTTE #Amiskwia Walcott [Qiong] ? cAnthotrum Luo et Hu [Qiong] ⁎Calathites Luo et Hu (=Phlogites Luo et Hu) ⁎Cambrotentactus Zhang et Shu (=Cotyledion Luo et Hu) ⁎Cheungkongella Shu et al. (?=Phlogites Luo et Hu) ? cConicula Luo et Hu [Qiong] c Cotyledion Luo et Hu [Qiong] c Dinomischus Chen, Hou et Lu [Qiong] ? cDiscoides Luo et Hu [Qiong] #Eldonia Walcott (being Stellostomites Sun et Hou) ? cHippotrum Luo et Hu in Luo et al. 1999 [Qiong] c Jiucunia Hou et al. [Qiong] ? cMaanshania Hou et al. [Qiong] ? cMacrocephalus Luo et Hu [Qiong] c Parvulonoda Rigby et Hou [Qiong] ? cPetalilium Luo et Hu [Qiong] ? cPhacatrum Luo et Hu [Qiong] c Phasganula Luo et Hu [Qiong] c Phlogites Luo et Hu [Qiong] ? cPristioites Luo et Hu [Qiong] ? cRhipitrus Luo et Hu [Qiong] c Rotadiscus Sun et Hou [Qiong] c Shankouclava Chen et al. [Qiong] c Stellostomites Sun et Hou [Qiong] ⁎Yunnanomedusa Sun et Hou (=Stellostomites Sun et Hou) ENIGMATIC TUBULAR FOSSILS Problematic hyoliths ⁎Acutitheca Yue (=Paragloborilus Qian) ⁎Arcitheca Duan (=Cupittheca Duan) Bucanotheca Qian et Jiang [M Meis] Cupittheca Duan [M Meis–Qiong] ⁎Ebianotheca He (=Paragloborilus Qian) ⁎Emeitheca Duan (=Cupittheca Duan) ⁎Ensitheca Duan (non Ensitheca Val'kov) (=Cupittheca Duan) Lophotheca Qian [M Meis–Qiong] Paragloborilus Qian [M Meis–L Meis] ⁎Varitheca Duan (=Cupittheca Duan) Coleolids Coleoloides Walcott [M Meis–Qiong] Coleolus Hall [M Meis] Hyolithelminths Annelitellus Qian [M Meis] Hyolithellus Billings [Meis–Qiong] Pseudorthotheca Cobbold [M Meis] Rushtonia Cobbold et Pocock [M Meis] Torellella Holm [M Meis–Qiong] Other tubular fossils Acanthoclava Li [M Meis] ?Cerabonusoides Qian [Qiong] ?Eoescharopora Jiang [M Meis] ?Flabetheca Qin et Yin [M Meis] ?Micropylepora Jiang [M Meis] ⁎Miratheca Yue (=Sagittitheca Yue) ?Orthangulites Qian et Jiang (nomen nudum) (?=Eohalobia Jiang) [M Meis] ?Petalites Zhong (Chen) (nomen nudum) [M Meis] Quadrochites Qian, Chen et Chen [M Meis] Rugatotheca He [E M Meis] Sagittitheca Yue [M Meis] Scissotheca Yue [M Meis] ENIGMATIC SCLERITES Paracarinachitids ⁎Luyanhaochiton Yu (=Paracarinachites Qian et Jiang) Paracarinachites Qian et Jiang [M Meis] Protopterygotheca Chen [M Meis] Scoponodus Jiang [M Meis] ⁎Yangtzechiton Yu (=Paracarinachites Qian et Jiang) Cambroclaves Cambroclavus Mambetov [Qiong] Deiradoclavus Conway Morris et Chen [L Meis] Deltaclavus Conway Morris et Chen [Qiong] ⁎Heterosculpotheca Jiang (=Zhijinites Qian) ⁎Isoclavus Qian et Zhang (=Cambroclavus Mambetov) Parazhijinites Qian et Yin [M Meis–Qiong] ⁎Phyllochiton Duan (Cambroclavus Mambetov) ⁎Sinoclavus Duan (=Cambroclavus Mambetov) ⁎Tanbaoites Duan (=Cambroclavus Mambetov) Zhijinites Qian [M Meis–Qiong] Tooth-like fossils Cyrtochites Qian [M Meis] Fomitchella Missarzhevsky [M Meis–Qiong] ?Jiangshanodus Yue ( being arthropod spines) [Qiong] #Kijacus Missarzhevsky (being arthropod spines) [Qiong] ⁎Leguminella He (=Paracanthodus Chen) Paracanthodus Chen [E M Meis] Rhombocorniculum Walliser [Qiong] Yunnanodus Wang et Jiang [M Meis] Tommotiids Camenella Missarzhevsky [L Meis] Lapworthella Cobbold [L Meis] Lugoviella Grigor'eva [Qiong] Micrina Laurie [Qiong] Porcauricula Qian et Bengtson [M Meis] G. Li et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 254 (2007) 229–249 Sonella Missarzhevsky et Grigor'eva [Qiong] Tannuolina Fonin et Smirnova [L Meis–Qiong] ⁎Tesella Missarzhevsky et Grigor'eva (=Sonella Missarzhevsky et Grigor'eva) ⁎Yunnanotheca Jiang (nomen nudum) (=Porcauricula Qian et Bengtson) Other sclerites Acanthocassis He et Xie [M Meis] Acanthosphaera Duan [M Meis–Qiong] Acidocharacus Qin et Ding [Qiong] ⁎Amoebinella He et Xie (=Acanthocassis He et Xie) Archicladium Qian et Xiao [L Meis–Qiong] Brushenodus Jiang [M Meis] ?Clypecella Li [M Meis] ?Dentlavula Li [M Meis] Emeithella Qian [M Meis] Fengzuella He et Yu [L Meis–Qiong] ⁎Huizenodus He et Xie (=Acanthocassis He et Xie) Kaiyangites Qian et Yin [Meis] ⁎Lunachites Qian et Yin (=Solenota Qian et Yin) ?Mabiania He [M Meis] ?Ningqiangsclerites Yue [M Meis] ⁎Otoformilites Qin et Li (=Fengzuella He et Yu) Paradoxiconus Qian et al. [M Meis] ⁎Phyllochites Qian et Yin (=Solenota Qian et Yin) ⁎Polycladium Qian et Xiao (=Archicladium Qian et Xiao) ⁎Pteromus Wang (=Brushenodus Jiang) Quadrorites Qian et Ding [M Meis] Rhabdochites He [M L Meis] #Salanacus Grigor'eva (being Acanthocassis He et Xie) Solenota Qian et Yin [M Meis] Tchangsichiton Yu [M Meis] ?Yangtzesclerites Chen et al. [M Meis] ?Zeugites Qian et al. (?=Eohalobia Jiang) [M Meis] Problematic chancelloriids (vase shaped) ⁎Acatomus Duan (=Cambrothyra Qian et Zhang) ⁎Anterosculum Duan, Cao et Zhang (=Cambrothyra Qian et Zhang) Cambrothyra Qian et Zhang [Qiong] ⁎Clinopa Duan (=Cambrothyra Qian et Zhang) ⁎Cyphinites Duan (=Cambrothyra Qian et Zhang ) ⁎Globifructus Geng et Zhang (=Cambrothyra Qian et Zhang) ⁎Horridomus Duan (=Cambrothyra Qian et Zhang ) ⁎Hubeitesta Duan (=Cambrothyra Qian et Zhang) #Lagenochitina Eisenack [E Meis] ⁎Lapistamnia Duan, Cao et Zhang (=Cambrothyra Qian et Zhang) ⁎Mirabichitina Yang et He (=Cambrothyra Qian et Zhang) ⁎Mirabifolliculus Yang et He (=Cambrothyra Qian et Zhang) ⁎Nanjiangochitina Yang et He (=Cambrothyra Qian et Zhang) ⁎Nanjiangofolliculus Yang et He (=Cambrothyra Qian et Zhang) ⁎Parahorridomus Duan, Cao et Zhang (=Cambrothyra Qian et Zhang) ⁎Pollofructus Geng et Zhang (=Cambrothyra Qian et Zhang) SPHEROIDAL FOSSILS (embryos and enigmatics) Aksuglobulus Qian et Xiao [Qiong] Archaeooides Qian [Meis–Qiong] ⁎Mirooides Yang, He et Deng (nomen nudum) (=Olivooides Qian) Olivooides Qian [Meis–Qiong?] ⁎Paramobergella Zhong (Chen) (nomen nudum) (=Olivooides Qian) ⁎Protosphaerites Chen (=Olivooides Qian) Pseudooides Qian [E M Meis] PSEUDOFOSSILS Ambarchaeooides Qian, Chen et Chen [M Meis] 247 Cambrospira Yu [M Meis] Eocucumaria Qian et Ding [E Meis] (being seeds) Huangshandongella Qian, Chen et Chen [ E M Meis] Nephrooides Qian [M Meis] References Babcock, L.E., Zhang, W., Leslie, S.A., 2001. 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