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
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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
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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
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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
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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
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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)
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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]
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?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
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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
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⁎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]
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