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Taxonomic and ecological variations of Permian-Triassic transitional bivalve

communities from the littoral clastic facies in southwestern China

Article in Palaeogeography Palaeoclimatology Palaeoecology · March 2018

DOI: 10.1016/j.palaeo.2018.02.027

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Palaeogeography, Palaeoclimatology, Palaeoecology

journal homepage: www.elsevier.com/locate/palaeo

Taxonomic and ecological variations of Permian-Triassic transitional bivalve

communities from the littoral clastic facies in southwestern China
⁎
Ting Songa, Jinnan Tonga, , Li Tiana, Daoliang Chua, Yunfei Huangb
a
State Key Laboratory of Biogeology and Environmental Geology, School of Earth Sciences, China University of Geosciences, Wuhan 430074, China
b
School of Geosciences, Yangtze University, Wuhan 430100, China

A R T I C LE I N FO A B S T R A C T

Keywords: Immediately after the latest Permian mass extinction, bivalve faunas colonized the Permian-Triassic Transitional
Community turnover Beds (PTTB), especially in littoral clastic facies of South China. The Permian-Triassic (P-Tr) transitional bivalve
Extinction interval fauna is composed of holdovers, long-term survivors and newly evolved taxa, and characterized by moderate
Post-extinction fauna diversity and high dominance. However, the taxonomy and ecology of this bivalve fauna of littoral clastic facies
Temporal variation
have not been well understood, especially in comparison with offshore marine records. This paper presents a
Spatial variation
study on a P-Tr transitional bivalve fauna from five sections of littoral clastic facies in southwestern China, and
as such it contributes to a better knowledge of the ecological processes of benthic communities during the P-Tr
transition. In total, 20 species in 13 genera of bivalves are identified from the PTTB in the lower part of the
Kayitou Formation and the basal part of the Feixianguan Formation of the five sections. Cluster analysis suggests
that the transitional bivalve fauna could be further divided into two successive communities (early and late
communities), and each is composed of five assemblages. Promyalina–Neoschizodus community (early commu-
nity) is characterized by presence of holdover taxa, high taxonomic diversity and evenness, and more abundant
shallow infaunal suspension feeders. In Pteria community (late community), taxonomic diversity and evenness
are much lower, and stationary epifaunal suspension feeders boost at the expense of shallow infaunal suspension
feeders compared with the Promyalina–Neoschizodus community. Furthermore, the Promyalina–Neoschizodus
community shows a clear spatial variation along environmental gradient from brackish lagoon to shallow marine
settings, while the Pteria community is more uniform and homogeneous laterally. The Promyalina–Neoschizodus
community in the studied region is of latest Permian age while the Pteria community should belong to the
earliest Triassic. Consequently, a significant taxonomical and ecological turnover of bivalve communities oc-
curred between the Promyalina–Neoschizodus community and the Pteria community during the extinction in-
terval in southwestern China.

1. Introduction
The Permian-Triassic (P-Tr) mass extinction event has not only re-
sulted in the largest decline in marine biodiversity through the
Phanerozoic history, but also an abrupt ecological switch of marine
benthic ecosystem from the Paleozoic type to the Modern type (Raup
and Sepkoski, 1982; Erwin, 1993; Bambach et al., 2004). Since then
mollusks, especially bivalves and gastropods, became the dominant
taxa in most marine benthic ecosystems (Gould and Calloway, 1980;
Yin, 1985; Hallam and Wignall, 1997; Fraiser and Bottjer, 2005, 2007;
Clapham and Bottjer, 2007a; Chen et al., 2010). Compared to the highly
diverse Late Permian benthic communities prior to the mass extinction,
the Early Triassic marine benthic ecosystems are much more uniform
owning to the predominance of the eurytopic “disaster” taxa, mainly
bivalves (Schubert and Bottjer, 1995; Petsios and Bottjer, 2016; Foster
et al., 2017).
Statistical analysis based on bivalve data from South China suggests
that bivalves only went through a single moderate extinction at genus
level during the ‘latest Permian mass extinction’ (LPME) whilst ex-
tinction rate shows no distinct spikes in the ‘earliest Triassic mass ex-
tinction’ (ETME) (Song et al., 2013; Huang et al., 2014). The interval
between the LPME and the ETME roughly corresponds to the P-Tr
Transitional Beds (PTTB; defined by Yin and Wu, 1985), which contain
abundant bivalves at many P-Tr boundary sections in South China (Yin,
1985; Shen et al., 1995; Tong and Xiong, 2006; Chen et al., 2009,
2010), Guryul Ravine of Kashmir (Nakazawa et al., 1975) and the
southern Alps (Broglio Loriga and Cassinis, 1992; Posenato, 2009),
especially in the clastic facies (Yao et al., 1980; Zhao et al., 1981; Sheng

⁎
Corresponding author.
E-mail address: jntong@cug.edu.cn (J. Tong).

https://doi.org/10.1016/j.palaeo.2018.02.027
Received 30 October 2017; Received in revised form 26 February 2018; Accepted 26 February 2018
0031-0182/ © 2018 Elsevier B.V. All rights reserved.

Please cite this article as: Song, T., Palaeogeography, Palaeoclimatology, Palaeoecology (2018), https://doi.org/10.1016/j.palaeo.2018.02.027
T. Song et al.
et al., 1984; Yin and Wu, 1985; Yang et al., 1987; Chen et al., 2009; Chu
et al., 2018). These P-Tr transitional bivalve faunas are notably dif-
ferent from the Late Permian pre-extinction faunas nor the same as
Early Triassic post-extinction faunas in view of both taxonomy and
ecology (Ciriacks, 1963; Yao et al., 1980; Yin and Wu, 1985; Fang,
1987; Sheng et al., 1987; He and Shen, 1991; Boyer et al., 2004; Tong
et al., 2006; He et al., 2007; Komatsu et al., 2008; Gao et al., 2009;
Hautmann et al., 2013, 2015; Yang, 2015; Foster et al., 2017; Zhang
et al., 2017). The transition from Late Permian fauna to P-Tr transi-
tional fauna, then to Early Triassic fauna documents the evolutionary
and ecological successions of these marine invertebrates associated
with the largest faunal turnover in the Phanerozoic history, and pro-
vides a unique window into the dynamics of benthic evolution during
the extinction interval. However, previous researches were mainly fo-
cused on the taxonomy and biochronology of the P-Tr transitional bi-
valves (Nakazawa et al., 1975; Yin and Wu, 1985; Yang et al., 1987; He
and Shen, 1991; Broglio Loriga and Cassinis, 1992; Shen et al., 1995;
Fang, 2004; Chen, 2004; Posenato, 2009; Chen et al., 2009; Yu et al.,
2010; Zhang et al., 2016; Cui et al., 2017; Chu et al., 2018). To better
understand the evolution of benthic faunas, further studies are needed
for the P-Tr transitional bivalve faunas.
Herein, we present a quantitative analysis of a P-Tr transitional
bivalve fauna from the PTTB of littoral clastic facies in southwestern
China, which documents the taxonomic and ecological changes of bi-
valves during this time in detail. Over five thousands of bivalve fossils
were collected from five P-Tr boundary sections in western Guizhou
and eastern Yunnan (Fig. 1), which deposited in various sedimentary
settings ranging from brackish lagoon to shallow marine facies (Yao
et al., 1980; Wang and Yin, 2001; Bercovici et al., 2015). The study
aims to reconstruct the taxonomical composition and ecological struc-
ture of the P-Tr transitional bivalve communities from the different
facies and understand the taxonomical and ecological evolutionary
process of bivalves throughout the P-Tr transition.
2. Geological setting
P-Tr deposits are well developed and exposed in western Guizhou
Fig. 1. Simplified palaeogeographic map, showing lithofacies types in South China during the P-Tr transition and the location of studied sections (the base map revised from Yin et al.,
2014). CNH, Chinahe; JZ, Jinzhong; JJB, Jinjibang; KL, Kele; WD, Wadu; HZS, Huangzhishan; MS, Meishan.
2
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx
and eastern Yunnan, southwestern China, and have attracted much
research concerning the P-Tr boundary and related events (e.g., Peng
et al., 2005; Shen et al., 2011; Yu et al., 2015; Zhang et al., 2016; Chu
et al., 2016, 2018; Cui et al., 2017; Zhang et al., 2017). Series of P-Tr
boundary sections, from marine to terrestrial facies, have been dis-
covered and studied in western Guizhou and eastern Yunnan, which
exhibit varied lithological successions due to facies variation (Shen
et al., 2011; Chu et al., 2016; Zhang et al., 2017). The Late Permian
succession of the studied area is represented by the Xuanwei Formation,
and the earliest Triassic succession comprises Kayitou Formation or
Feixianguan Formation in different settings.
The Xuanwei Formation of Late Permian age is mainly composed of
greyish-green and brownish-yellow sandstone, siltstone and mudstone,
interbedded by some coal beds (Fig. 2), and contains abundant and
well-preserved Permian plant fossils (Yu et al., 2010, 2015; Chu et al.,
2016). The lithological and fossil compositions of the Xuanwei For-
mation indicate a swamp and shallow lake facies (Bercovici et al.,
2015).
The Kayitou Formation conformably overlies the Xuanwei
Formation in the western part of the studied area, e.g., Chinahe and
Jinzhong sections, and is mainly composed of yellowish-green and
brownish-yellow siltstone and mudstone in the lower part (Fig. 2A–C,
E), and greyish-blue, purple-reddish sandstone and siltstone in the
upper part. The lower part of the Kayitou Formation contains abundant
conchostracans, fossil plants and bivalves, but very few fossils are col-
lected in the upper part. The Kayitou Formation is overlain by the
reddish Dongchuan Formation of terrestrial facies which only yields
few terrestrial fossils.
In the eastern part of the studied area the Xuanwei Formation is
overlain by the Feixianguan Formation, which can be subdivided into
four lithological members (Chen et al., 1979). The basal part of the first
member, i.e. the PTTB, is mainly composed of brownish, yellowish-
green siltstone and mudstone (Fig. 2D, F). The basal part of the Feix-
ianguan Formation contains rich bivalves, and some brachiopods and
gastropods (Yin and Wu, 1985; Yang et al., 1987).
The bivalves are collected from the PTTB in the lower part of the
Kayitou Formation and the basal part of the Feixianguan Formation.
T. Song et al.
Fig. 2. Field photos showing the P-Tr boundary sequences of studied sections. A, B, Chinahe section; C, Jinzhong section (note that strata are overturned); D, Jinjibang section; E, Kele
section; F, Wadu section.
According to the depositional settings and facies analysis (Bercovici
et al., 2015), these bivalves lived in a coastal brackish lagoon (Chinahe,
Jinzhong and Jinjibang sections) or shallow marine (Kele and Wadu
sections) setting, among which the Jinjibang section is located more
oceanward compared to the Chinahe and Jinzhong sections.
3. Methods
Five P-Tr boundary sections were investigated in the littoral clastic
facies of southwestern China. Macrofossils were collected bed by bed in
the field work. Each fossil bed was carefully collected either as much as
possible or until no new species appeared in the outcrop. All the bivalve
fossils were identified and counted in the laboratory. For quantitative
analysis, the number of the bivalve species was inferred by the more
numerous valves (left or right valves). Nearly all the bivalve fossils were
preserved as complete and undamaged valves, and some of them were
articulated, suggesting that they are para-autochthonous.
For the bivalve assemblages, alpha diversity is measured using in-
dices including Shannon index (H), Simpson evenness index (E) and
species richness (S). The Shannon index is calculated as H = −∑ (n/N)
In (n/N), the Simpson evenness as E = 1 − ∑ (n/N)2, where n is the
number of individuals of taxon i, N is the number of total individuals
(Lande, 1996), and S (species richness) is the number of species
3
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx
occurred. However, these diversity indices are not themselves real di-
versities (Jost, 2007). Therefore effective diversity (effective numbers
of species) is applied specially as exp.(H) to intuitively reflect and
compare the real diversity differences between communities (Jost,
2007). Apart from the diversity indices, trophic nucleus was also cal-
culated as the number of taxa that constitute at least 80% of the com-
munity (Neyman, 1967).
Faunal abundance data were analyzed with the software package
PAST (Hammer et al., 2001). Cluster analysis (unweighted paired group
Q-mode) using the Morisita index was performed in order to generate
assemblages of fossil samples on the basis of species' occurrence and
raw abundance, as well as to assess the habitat partitioning of the re-
cognized assemblages along the environment ingredients. Morisita
index is effective in ecological study of palaeocommunities (Hammer
and Harper, 2006). F test was applied here to evaluate the significant
differences between two communities (Hammer et al., 2001), and the
significance level was chosen as 0.05. The difference between two
communities is significant when the F value is < 0.05. The abundance
of the bivalve fossils is given in the raw data (supplemental information
1). In addition, rarefaction analysis with 95% confidence is applied to
account for the effect of sample size on taxonomic richness.
An assemblage is a collection of fossils made from a single bed
(Brenchley and Harper, 1998), or from several successive beds that
 T. Song et al.

4
Fig. 3. Lithological columns and bivalve distributions of the five studied sections and the correlations. Colored shades indicate different bivalve assemblages at each section: pink for the lower assemblage, aqua for the upper assemblage and blue for
the Claraia assemblage of late Griesbachian age. The arrow indicates the last occurrence of coal bed at each section. PTTB, Permian–Triassic Transitional Beds; e-PTB, estimated P-Tr boundary; Xw Fm., Xuanwei Formation. The red dashed line
indicates the estimated P-Tr boundary; the yellow solid line indicates the boundary of the PTTB and the yellow dashed line indicates the tentative boundary of the PTTB. (For interpretation of the references to colour in this figure legend, the reader
is referred to the web version of this article.)
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx
T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Fig. 4. Bivalve fossils from the Chinahe section. All scale bars represent 2.5 mm. A-C. Promyalina schamarae, left valve, CNH-1415, CNH-1428, CNH-1450; D. Promyalina putiatinensis, left
valve, CNH-1467; E. Neoschizodus orbicularis, left valve, CNH-14112; F. Austrotindaria antiqua, left valve, CNH-141269; G-H. Permophorus bregeri, G, right valve, CNH-14269. H, left valve,
CNH-14380; I-L. Pteria cf. ussurica, I, right valve, CNH-14455. J-L, left valve, CNH-14556, CNH-14798, CNH-14946.

Fig. 5. Cluster analysis of the bivalve samples based on the Morisita index in the four studied sections. A, Chinahe; B, Jinjibang; C, Kele; D, Wadu.

share the same species (this paper). Communities (palaeocommunities) palaeocommunities and are defined as the coeval assemblages re-
refer to the fossilized residues of living communities and are assem- cognized by the cluster analysis in the studied sections. An ecological
blages that represent the former living communities (Brenchley and guild is a group of species that exploit the same resources in a similar
Harper, 1998). Communities in this paper are time-averaged way (Simberloff and Dayan, 1991).

5
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4. Results wangi assemblage, composed of samples JJB10-13. The Claraia wangi

assemblage is excluded from further discussion concerning bivalves in
4.1. Chinahe section the PTTB because it is part of the Claraia wangi zone of late Griesba-
chian age and definitely higher than the PTTB. So two bivalve assem-
Chinahe section is located in the Tianba town, Xuanwei City, blages are identified from the PTTB at Jinjibang section: the Neoschi-
eastern Yunnan Province (Fig. 1). Fossil plants are found in great zodus orbicularis assemblage (JJB1-8, lower assemblage) and the Pteria
abundance in the Xuanwei Formation below the uppermost coal bed, cf. ussurica assemblage (JJB9, upper assemblage) (Fig. 5B). Rarefaction
while the Kayitou Formation yields abundant bivalves, and some lin- analysis shows that further sampling would not significantly increase
gulid brachiopods, gastropods, conchostracans and fossil plants. the taxonomical diversity of both assemblages (Fig. 6).
In total, 1290 bivalve specimens were collected from 11 bivalve beds
in the lower part of the Kayitou Formation at this section. Seven species
from five genera are identified, including Pteria cf. ussurica, Promyalina 4.4. Kele section
schamarae, P.putiatinensis, Permophorus bregeri, Austrotindaria antiqua,
Neoschizodus orbicularis and N.laevigatus (Figs. 3, 4). Kele section is located in the Hezhang County, Bijie City, western
Cluster analysis based on the 11 bivalve beds in the Chinahe section Guizhou Province (Fig. 1). The upper Xuanwei Formation yields
identifies three assemblages (similarity > 0.6) (Fig. 5A). Sample CNH8 abundant fossil plants of Late Permian age below the last coal bed,
is shown as an independent assemblage, however, the sample size is too while the lower Kayitou Formation contains abundant bivalves and
small and the rarefaction analysis indicates that the sampling is highly some plants, miniaturized gastropods and brachiopods (lingulid and
inefficient (supplementary information 2A). Therefore, sample CNH8 is Crurithyris).
included into CNH1-7 because they are more similar in taxonomic There were 847 bivalves identified from six beds at the Kele section,
compositions. So two bivalve assemblages are distinguished at the composed of 15 species in nine genera: Pteria cf. ussurica, Promyalina
Chinahe section, and they are Pteria cf. ussurica-Promyalina schamarae schamarae, P.putiatinensis, P.spathi, P.sp., Permophorus bregeri,
assemblage (CNH1-8, lower assemblage) and Pteria cf. ussurica assem- Austrotindaria antiqua, Neoschizodus orbicularis, N.laevigatus, Pernopecten
blage (CNH9-11, upper assemblage) (Fig. 5A). Rarefaction analysis huayingshanensis, Leptochondria sp., Gujocardita sp., G.subquadrata,
suggests that both assemblages are well sampled (Fig. 6). Astartella sp. and A.doulingensis (Figs. 3, 9).
In the Kele section, the bivalves from the six beds show quite high
4.2. Jinzhong section similarity (> 0.6) in the cluster dendrogram (Fig. 5C). However, KL1 is
inefficiently sampled (supplementary information 2B) and comparison
Jinzhong section is located in the Jinzhong Town, Weining County,
western Guizhou Province (Fig. 1). The Xuanwei Formation yields
Permian Gigantopteris Flora below the uppermost coal bed, while the
Kayitou Formation contains abundant bivalves and some lingulid bra-
chiopods, conchostracans and fossil plants.
In total, 216 bivalves were collected from two beds in the lower
Kayitou Formation at the Jinzhong section. Six species from five genera
were identified: Pteria cf. ussurica, Neoschizodus orbicularis, N.laevigatus,
Permophorus bregeri, Promyalina schamarae and Austrotindaria antiqua
(Figs. 3, 7).
All specimens at the Jinzhong section are found only from two
sampling horizons (Fig. 3). But their taxonomic compositions are
clearly different: JZ1 is much more diversified than JZ2, thus they are
taken as two independent assemblages in later analysis, i.e. Pteria cf.
ussurica-Neoschizodus orbicularis assemblage (JZ1, lower assemblage)
and Pteria cf. ussurica assemblage (JZ2, upper assemblage). Rarefaction
analysis shows that the lower assemblage is well sampled. The upper
assemblage has only two taxa occurrence, thus rarefaction analysis is
not conducted. Sampling of the upper assemblage is somehow in-
efficient because the sample size is quite small.

4.3. Jinjibang section

Jinjibang section is located in the Zhenzhou Town, Junlian County,

southern Sichuan Province (Fig. 1). The upper Xuanwei Formation
contains abundant fossil plants of Permian age, whereas the basal part
of Feixianguan Formation yields abundant bivalves, some lingulid
brachiopods, fossil plants and a few conchostracans (Fig. 3).
Altogether 669 bivalves belonging to nine species in seven genera
were identified from 12 beds in the basal part of the Feixianguan
Formation at the Jinjibang section. They are Neoschizodus orbicularis,
N.laevigatus, Pteria cf. ussurica, Austrotindaria antiqua, A.canalensis,
Permophorus bregeri, Promyalina putiatinensis, Claraia wangi and
Eumorphotis sp. (Figs. 3, 8).
In the Jinjibang section, cluster analysis from the 12 bivalve beds
identifies three bivalve assemblages (similarity > 0.6) (Fig. 5B). They Fig. 6. Rarefaction analysis with 95% confidence of the nine transitional bivalve as-
are Neoschizodus orbicularis assemblage, composed of samples JJB1-8, semblages. A, the five lower bivalve assemblages; B, the four upper bivalve assemblages.
CNH, Chinahe; JZ, Jinzhong; JJB, Jinjibang; KL, Kele; WD, Wadu.
Pteria cf. ussurica assemblage, composed of sample JJB9 and Claraia

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T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Fig. 7. Bivalve fossils from the Jinzhong section. Scale bars represent 2.5 mm in all figs. A-C. Neoschizodus orbicularis, left valve, JZ0784, JZ0795, JZ0756; D. Neoschizodus laevigatus, right
valve, JZ07121; E-F. Permophorus bregeri, right valve, JZ07154, JZ07167; G-H. Promyalina schamarae, G, left valve, JZ0813. H, right valve, JZ0819; I-L. Pteria cf. ussurica, I, left valve,
JZ0714. J-L, right valve, JZ0834, JZ0826, JZ0816.

Fig. 8. Bivalve fossils from the Jinjibang section. All scale bars represent 2.5 mm. A. Permophorus bregeri, left valve, JJB0212; B. Pteria cf. ussurica, left valve, JJB0231; C. Austrotindaria
canalensis, left valve, JJB0324; D. Promyalina putiatinensis, right valve, JJB02356; E-H. Neoschizodus laevigatus, right valve, JJB02421, JJB02267, JJB02541, JJB02672; I-L. Neoschizodus
orbicularis, I, right valve, JJB02568. J-L, right valve, JJB02145, JJB02126, JJB02635.

of taxonomic compositions suggests that bivalves from KL1 to KL4
could form an assemblage, and KL5 and KL6 should constitute another
assemblage. Thus two bivalve assemblages are distinguished from this
section, i.e. Pteria cf. ussurica-Promyalina schamarae assemblage (KL1-
4,lower assemblage) and Pteria cf. ussurica- Austrotindaria antiqua as-
semblage (KL5-6,upper assemblage) (Fig. 5C). The rarefaction analysis
indicates that the lower assemblage is a little inefficiently sampled
while the upper assemblage is sufficiently sampled (Fig. 6).
4.5. Wadu section
Wadu section is located in the Wadu village, Panxian County,
western Guizhou Province (Fig. 1). The Xuanwei Formation is rich in
Permian fossil plants below the uppermost coal bed. The basal part of

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T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Fig. 9. Bivalve fossils from the Kele section. All scale bars represent 2.5 mm. A-C, E-F. Promyalina schamarae, A, E, left valve, KL39005, KL39017. B, C, F, right valve, KL39057, KL37012,
KL3706; D. Promyalina spathi, right valve, KL39102; G. Promyalina putiatinensis, right valve, KL39124; H. Pernopecten huayingshanensis, right valve, KL39124; I. Neoschziodus laevigatus, left
valve, KL37124; J. Neoschizodus orbicularis, left valve, KL37294; K-L. Permophorus bregeri, right valve, KL3715, KL37350; M-Q. Pteria cf. ussurica, M, N, left valve, KL37574, KL37265. O-Q,
right valve, KL3767, KL37538, KL37561; R. Leptochondria sp., left valve, KL37631; S-T. Austrotindaria antiqua, left valve, KL37126, KL37348; U-V. Astartella sp., S, right valve, KL3723. T,
left valve, KL3779; W-Y. Gujocardita sp., left valve, KL37105, KL37603, KL37715.

the Feixianguan Formation contains rich bivalves, lingulid brachiopods, In the basal part of the Feixianguan Formation at the Wadu section,
and some rhynchionelliform brachiopods (Crurithyris and Acosarina), 390 bivalves are identified from eight beds, including 15 species in 11
gastropods, and the diversity and abundance of bivalves decline sharply genera: Pteria cf. ussurica, Claraia bioni, C.wangi, Permophorus bregeri,
upward at the Wadu section. Eumorphotis venetiana, Neoschizodus laevigatus, N.orbicularis, Bakevellia

8
T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Fig. 10. Bivalve fossils from the basal part of Feixianguan Formation at the Wadu section. All scale bars represent 2.5 mm. A. Leptochondria virgalensis, left valve, WD19005; B. Towapteria
scythica, left valve, WD801210; C-D. Astartella sp., C, right valve, WD19017. D, left valve, WD19058; E. Permophorus bregeri, left and right valve joint together, WD19211; F-I. Pteria cf.
ussurica, F, H, right valve, WD19024, WD19032. G, I, left valve, WD19098, WD19075; J. Neoschizodus orbicularis, left valve, WD19041; K. Neoschizodus laevigatus, left valve, WD19053; L.
Permophorus bregeri, right valve, WD19101; M. Claraia wangi, left valve, WD19077; N-O. Claraia bioni, N, left valve, WD 801190. O, right valve, WD905002; P-Q. Eumorphotis venetiana, P,
right valve, WD19064. Q, left valve, WD801188; R-S. Promyalina putiatinensis, right valve, WD19057, WD19083; T. Promyalina spathi, right valve, WD19132; U. Promyalina schamarae, left
valve, WD19251.

sp., Promyalina schamarae, P.putiatinensis, P.spathi, Austrotindaria an-
tiqua, Towapteria scythica, Astartella sp. and Leptochondria virgalensis
(Figs. 3, 10).
In the Wadu section, cluster analysis identifies five assemblages
from the eight beds based on similarity > 0.6 (Fig. 5D). Samples WD7
and WD8 form an assemblage characteristic of the Claraia wangi Zone,
which is of the late Griesbachian age and definitely higher than the
PTTB, therefore this assemblage is excluded from further discussion.
Samples WD1 and WD2 clearly constitute a lower assemblage, and
samples WD3 and WD4 form an upper assemblage. Samples WD5 and
WD6 are separately clustered as two independent branches on the
dendrogram, mainly because they are under-sampled. However, both of
them share the same characteristic and dominant bivalve element Pteria
cf. ussurica with the underlying WD3–4 (the upper assemblage), con-
sequently they are included into the upper assemblage. So two bivalve
assemblages are distinguished from the PTTB at the Wadu section in-
cluding Pteria cf. ussurica-Promyalina schamarae assemblage (WD12,
lower assemblage) and Pteria cf. ussurica-Claraia bioni assemblage
(WD3–6, upper assemblage) (Fig. 5D). Rarefaction analysis indicates
that the lower assemblage needs more sampling, but further sampling
would not significantly increase the diversity of the upper assemblage
(Fig. 6).

9
T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Table 1
List of all identified species and their ecology, evolutionary fates and stratigraphical ranges.

Taxa Ecology# Evolutionary status# Stratigraphical range## Indentification after

C PT1 PT2 I O A

Promyalina schamarae Epibys-sf Newly evolved * * * Huang, 2014

Promyalina putiatinensis Epibys-sf Newly evolved * * * * Hautmann et al., 2013
Promyalina spathi Epibys-sf Newly evolved * * * Ciriacks, 1963
Promyalina sp. Epibys-sf Newly evolved * Ciriacks, 1963
Pteria cf. ussurica Epibys-sf Newly evolved * * * * Chen et al., 2009
Bakevellia sp. Sinf-sf Long-term survivor * Huang, 2014
Towapteria scythica Epibys-sf Holdover * * Chen et al., 2009
Claraia bioni Epibys-sf Long-term survivor * Nakazawa and Hapoor, 1981
Eumorphotis venetiana Epibys-sf Newly evolved * * * Chen, 2004
Leptochondria virgalensis Epibys-sf Long-term survivor * * * Huang, 2014
Leptochondria sp. Epibys-sf Long-term survivor * Yin, 1985
Pernopecten huayingshanensis Epibys-sf Holdover * * Yang, 2015
Permophorus bregeri Infmob-sf Long-term survivor * * * * Huang, 2014
Austrotindaria antiqua Infmob-sf Newly evolved * * * * Foster et al., 2017
Neoschizodus orbicularis Infmob-sf Newly evolved * * * * * Hofmann et al., 2015
Neoschizodus laevigatus Infmob-sf Newly evolved * * * * * Hofmann et al., 2015
Astartella sp. Infmob-sf Holdover * Hautmann et al., 2015
Astartella doulingensis Infmob-sf Holdover * * Fang, 1987
Gujocardita sp. End-inf-sf Holdover * Yang, 2015
Gujocardita subquadrata End-inf-sf Holdover * * Fang, 1987

epi, epifaunal; inf, infaunal; sinf, semi-infaunal; bys, byssally attached; mob, mobile; end, endobyssate; C, Changhsingian; PT1, early P-Tr transition; PT2, late P-Tr transition;I, Induan; O,
Olenekian; A, Anisian. #genus level, ##species level. The ecology is partly inferred after Stanley, 1970, the life habit of modern closely related species, and partly cited from Huang,
2014; Hofmann et al., 2015; Hautmann et al., 2015. The evolutionary status and stratigraphical ranges are cited from Huang, 2014.

5. Discussion
5.1. Temporal variation
In total, 20 species from 13 genera of bivalves were recognized from
the PTTB of the five studied sections in southwestern China (Table 1).
At each section, a lower and an upper bivalve assemblage are dis-
tinguished from the PTTB (Figs. 3, 5). The five lower assemblages at the
five studied sections share six dominant components together: Pteria cf.
ussurica, Neoschizodus orbicularis, N.laevigatus, Promyalina schamarae,
Austrotindaria antiqua and Permophorus bregeri, while the five upper
assemblages are predominated overwhelmingly by Pteria cf. ussurica
alone (Fig. 11). Considering all the bivalve fossils are preserved para-
autochthonous, these fossil assemblages are regarded as two time-
averaged communities, and they are named Promyalina–Neoschizodus
community (= early P-Tr transitional bivalve community) and Pteria
community (= late P-Tr transitional bivalve community) according to
their dominant and characteristic components. The Promyalina–-
Neoschizodus community is composed of the five lower assemblages and
the Pteria community contains the five upper assemblages. These two
transitional bivalve communities represent different ecological stages of
marine benthic ecosystem during the P-Tr transition.
Taxonomic richness declines from the Promyalina–Neoschizodus
community to the Pteria community. The Promyalina–Neoschizodus
community contains 15 species in 9 genera while the Pteria community
is composed of 12 species in 10 genera (Fig. 11). Effective diversity
drops about 34%, Shannon index decreases from 1.63 to 1.21 and the
evenness declines from 0.72 to 0.49 (Table 2). Result of the F test shows
that the diversity of the two communities differed significantly
(P = 0.03). In addition, the trophic nucleus is also higher in the Pro-
myalina–Neoschizodus community.
The Promyalina–Neoschizodus community is composed of Permian
holdovers, long-term survivors and newly evolved taxa, whereas the
Pteria community is entirely composed of long-term survivors and
newly evolved taxa (Table 1). There are five Permian holdover species
that occurred in the Promyalina–Neoschizodus community and went
extinct in the Pteria community: Pernopecten huayingshanensis, Gujo-
cardita subquadrata, G.sp., Astartella doulingensis, and A.sp. The other 10
species in the Promyalina–Neoschizodus community, together with the
five newly evolved species in the Pteria community, all appeared for the
first time in the PTTB and persisted into the Early Triassic except for
Claraia bioni. Claraia bioni was previously reported only in the PTTB of
the Huangzhishan section of South China and Guryul Ravine of Kashmir
(Nakazawa et al., 1975; Chen, 2004; Chen et al., 2009), and it did not
extend stratigraphically higher than the PTTB.
Ecological guilds differ greatly between the two communities. Three
ecological guilds are present in the Promyalina–Neoschizodus commu-
nity, including stationary epifaunal suspension feeder (7 species), mo-
bile infaunal suspension feeder (6 species) and endobyssate infaunal
suspension feeder (2 species), while the Pteria community contains
three slightly different guilds, including stationary epifaunal suspension
feeder (7 species), mobile infaunal suspension feeder (4 species) and
stationary semi-infaunal suspension feeder (1 species) (Fig. 11). The
stationary epifaunal suspension feeder and mobile infaunal suspension
feeder are more diverse than other guilds and co-dominate the Pro-
myalina–Neoschizodus community together in absolute abundance,
while stationary epifaunal suspension feeder alone predominates the
Pteria community by over 80%. Both endobyssate infaunal suspension
feeder and semi-infaunal suspension feeder are of low diversity, the
former accounting for 8.2% of individuals in the Promyalina–Neoschi-
zodus community at the Kele section and the latter only for 0.3% in the
Pteria community at the Wadu section (Fig. 11).
The diversity of the two communities differs greatly. The
Promyalina–Neoschizodus community is characterized by the occurrence
of Permian holdover taxa, a high taxonomic diversity and evenness, and
more abundant mobile infaunal suspension feeders. However, upward
within the PTTB, the Permian holdover taxa disappear quickly, sta-
tionary epifaunal suspension feeders begin to predominate and some
new species appear for the first time. Thus, the transition from the
Promyalina–Neoschizodus community to the Pteria community is char-
acterized by decreasing diversity and evenness, and increasing of epi-
faunal suspension feeders at the expense of shallow infaunal suspension
feeders.
5.2. Spatial variations
The PTTB of the studied sections were deposited in a coastal
brackish lagoon (Chinahe, Jinzhong and Jinjibang sections) or shallow

10
T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Fig. 11. Histograms showing the frequencies of bivalves, the trophic nucleus, absolute abundance of ecological guilds and guild species diversity (with number inside the piechart) based
on the number of species representing a certain guild as well as the abundance of the bivalve assemblages. The grey column represents the lower assemblage, and the purple is the upper
assemblage. A, B, Chinahe; C, D, Jinzhong; E, F, Jinjibang; G, H, Kele; I, J, Wadu. (For interpretation of the references to colour in this figure legend, the reader is referred to the web
version of this article.)

Table 2
Diversity indices and effective diversities of the Promyalina–Neoschizodus community and the Pteria community.

Bivalve community Species (genera) Shannon index (H) Effective diversity Simpson evenness index (E) Trophic nucleus (T)

Pteria community 12 (10) 1.21 3.37 0.49 3

Promyalina–Neoschizodus community 15 (9) 1.63 5.09 0.72 4

11
T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

Fig. 12. Cluster dendrograms of the bivalves from the PTTB and diversity of subcommunities at the five sections. A. Cluster dendrogram of the lower assemblages (Promyalina-
Neoschizodus community) showing three subcommunities. B. Diversity (indices) of three subcommunities of the Promyalina-Neoschizodus community. C. Cluster dendrogram of the upper
assemblages (Pteria community), note that they share relatively higher similarities compared to the lower assemblages, and the subdivision of subcommunities might not be significant (in
Fig. 12A). D. Diversity (indices) of three subcommunities of the Pteria subcommunities corresponding to three facies. CNH, Chinahe; JZ, Jinzhong; JJB, Jinjibang; KL, Kele; WD, Wadu.

Table 3
P-Tr transitional sequences and correlation of some classic sections.
Guryul Ravine section
Meishan section
Huangzhishan section (Nakazawa and Hapoor, 1981;
chronostratigraphy (Tong and Xiong, 2006; Yin et al.,
(Chen, 2004; Chen et al., 2009) Sheng et al., 1984; Fang,
2014)
2004)
Claraia wangi zone Claraia wangi zone Claraia concentrica zone
Induan Bed 29
Ophiceras zone Ophiceras zone Ophiceras zone
Triassic
Bed 28 Isarcicella staeschei zone Hindeodus parvus zone Hindeodus parvus zone
Pteria -Eumorphotis
P-Tr Hindeodus parvus zone assemblage Eumorphotis venetiana zone
Bed 27
transitional
bed Peribositra baoqingensis Claraia huzhouensis - Claraia bioni - Etheripecten
Bed 26
Hypophiceras fauna Claraia cf. bioni zone haydeni zone
Permian Bed 25
"Palaeolima" middlemissi
Changhsingian Bed 24 Palaeofusulina zone Clarkina yini zone
zone

The red line represents the latest Permian mass extinction and the earliest Triassic mass extinction, respectively.

marine (Kele and Wadu sections) setting (Fig. 3). Chinahe, Jinzhong
and Jinjibang sections were all situated in a brackish lagoon setting, of
which Jinjibang section should be located more oceanward because the
PTTB at this section are mainly composed of dark mudstone and con-
tains few terrestrial fossils, while the PTTB at Chinahe and Jinzhong
sections are mainly comprised of silty mudstones and contain abundant
terrestrial fossils (see above). Accordingly, the brackish lagoon facies is
further subdivided into two subfacies, i.e. brackish lagoon landward (L;
Chinahe and Jinzhong) and brackish lagoon oceanward (O; Jinjibang).
In contrast, both Kele and Wadu sections represent a shallow marine
facies during the deposition of the PTTB.
Cluster analysis based on the lower assemblages (the
Promyalina–Neoschizodus community) from the five sections suggests
that the five sections can be further grouped into three subcommunities
(similarity > 0.6) (Fig. 12A, B), which correspond to the
palaeoenvironmental positions. The first subcommunity lived at the
Chinahe and Jinzhong sections in a landward brackish lagoon facies,
co-existing with abundant terrestrial fossils, such as conchostracans and
plants. The second subcommunity is at the Kele and Wadu sections in a
shallow marine facies. The third subcommunity from the Jinjibang
section lived in an oceanward lagoon facies and correlates poorly with
other subcommunities. Consequently, these three subcommunities
correspond perfectly with the facies differentiation among the five
sections, and can be assigned to brackish lagoon landward, brackish
lagoon oceanward and shallow marine subcommunities (Fig. 12A, B).
However, the upper assemblages of the Pteria community show rela-
tively high similarities among the five sections (similarity > 0.8)
(Fig. 12C), suggesting a more uniform community without remarkable
differentiation of subcommunities throughout the studied area.
The shallow marine facies records the highest diversity in both

12
T. Song et al. Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx

bivalve communities while the lagoon oceanward facies records the

Yang and Li, 1992

lowest (Fig. 12). Effective diversity, Shannon index and Simpson

Chen et al., 2009

Shen et al., 1995

Posenato, 2009
Yu et al., 2010

Nakazawa and
evenness index in the shallow marine facies are all more than twice of

Kapoor, 1981
Yang, 2015
the lagoon oceanward facies in both communities (Fig. 12). In addition,
This study
Reference

trophic nucleus remains highest value in the shallow marine facies in

both communities (Fig. 12).
The different settings of the studied sections differ in the ecology of

Claraia concentrica
Late griesbachian

the bivalves (Fig. 11). In both the Promyalina–Neoschizodus and Pteria

Claraia wangi

Claraia wangi
Claraia wangi

Claraia wangi

Claraia wangi

Claraia wangi
communities, stationary epifaunal suspension feeders have low di-
versity in the lagoon landward and oceanward facies (1–3 species), and
high diversity in the shallow marine facies (maximum 7 species), while
mobile infaunal suspension feeders maintain a moderate diversity (2–5
–

species) in all the facies. In terms of dominance, mobile infaunal sus-

bregeri, Austrotindaria antiqua, Claraia bioni, Eumorphotis
Pteria cf. ussurica, Neoschizodus laevigatus, Permophorus

Eumorphotis multiformis, Eumorphotis venetiana, Pteria ussurica variabilis, Towapteria scythica, Leptochondria virgalensis,

Eumorphotis venetiana, Eumorphotis bokharica, Claraia

pension feeders dominate in the lagoon landward facies at the Jinzhong
antiqua, Austrotindaria canalensis, Towapteria scythica

ussurica variabilis, Bakevellia costata, Leptochondria

section (56%) and lagoon oceanward facies at the Jinjibang section

Eumorphotis venetiana, Towapteria scythica, Pteria

Towapteria scythica, Promyalina sp., Claraia sp.

(94%) whilst stationary epifaunal suspension feeders dominate in other

facies of the Promyalina–Neoschizodus community. However, in the
Pteria community stationary epifaunal suspension feeders dominate all
the three facies overwhelmingly (Fig. 11). In addition, the ecological
griesbachi, Leptochondria minima

diversity is much higher in the shallow marine facies than in the lagoon
facies. Four ecological guilds are present in the shallow marine facies,
virgalensis, Promyalina sp.
Late bivalve community

which is twice as much as in the lagoon facies.

Eumorphotis venetiana

In short, the Promyalina–Neoschizodus community develops a strong

spatial variation along environmental gradient from the brackish la-
goon to shallow marine facies and could be subdivided into three
venetiana

subcommunities corresponding to the facies differentiation, while the

Pteria community shows a much more uniform distribution without
remarkable differentiation of subcommunities throughout the studied
Pteria ussurica variabilis, Leviconcha orbicularis, Austrotindaria

area. In both communities, the shallow marine setting records the

Claraia bioni, Claraia huzhouica, Towapteria sp., Palaeolima

Claraia bioni, Palaeolima middlemissi, Etheripecten haydeni,

Towapteria, Unionites, Eumorphotis, Bakevellia, Promyalina

highest diversity of bivalves both in terms of taxonomy and ecology.

Pteria cf. ussurica, Neoschizodus orbicularis, Neoschizodus
laevigatus, Promyalina schamarae, Permophorus bregeri,

5.3. PTTB
Major bivalve components of the P-Tr transitional bivalve communities in the studying sections and some P-Tr sections.

The P-Tr boundary, defined by the first appearance datum (FAD) of

Cyrtorostra lunwalensis, Palaeoneilo sp.

conodont Hindeodus parvus, lies in the middle part of the PTTB at many
sections (Tong and Xiong, 2006; Chen et al., 2009, 2010). The PTTB
were officially defined by Yin and Wu (1985) as a transitional strati-
Astartella sp., Gujocardita sp.

graphical sequence yielding a mixed biota composed of “Permian re-

Early bivalve community

licts” and “Triassic type” new taxa in continuous P-Tr boundary sec-
Pteria ussurica variabilis

Claraia guangyuanensis

tions. The bottom of the PTTB is confined by the LPME and the top by
Bakevellia costata

occurrence of abundant bivalve Claraia or ammonoid Ophiceras (Yin

and Wu, 1985; Yang et al., 1987). A later study found that the PTTB
perfectly correspond to the interval between the LPME and ETME at the
Meishan section, which specifically include Beds 25–28 and is also
sp.

consistent with the P-Tr Boundary Stratigraphic Set at the Meishan

section (Peng et al., 2001; Tong and Xiong, 2006; Song et al., 2013)
Shallow carbonate

(Table 3). Therefore, the PTTB is isochronous and can be correlated for
Littoral clastic

Littoral clastic

Littoral clastic
Littoral facies

the P-Tr boundary in South China (Tong and Xiong, 2006). In addition,
Carbonate

Carbonate
platform

platform

the P-Tr boundary lies in the middle of Bed 27 at the Meishan section,
Facies

facies

facies

facies
Basin

which is in the middle part of the PTTB. It is also in the middle part of
the PTTB at the Huangzhishan (Chen, 2004; Chen et al., 2009) and
Zhongzhai (Yang, 2015; Zhang et al., 2014) sections of South China, the
Guryul Ravine section of Kashmir (Nakazawa and Hapoor, 1981;
Jinjibang, Kele, Wadu

Hapoor, 1992) and the Tesero Member of the Werfen Formation from
Cnahe, Jinzhong,

Northern Italy (Broglio Loriga and Cassinis, 1992; Posenato, 2009)

Guryul Ravine
Huangzhishan

Tesero, Bulla
Liangfengya

(Table 3).
Zhongzhai
Section(s)

The P-Tr transitional bivalve fauna has been commonly used to

Shangsi
Mide

distinguish the P-Tr boundary in the littoral clastic facies (Yu et al.,
2010; Zhang et al., 2016; Chu et al., 2016). Although the PTTB have
been widely observed in various facies over the world (Nakazawa et al.,
Western Dolomites,

1975; Yao et al., 1980; Sheng et al., 1987; Peng et al., 2001; Posenato,
2009), bivalves are mostly diverse and abundant in the littoral clastic
South China

South China

South China
South China

South China

South China

facies (Chen et al., 1979; Yao et al., 1980; Yang et al., 1987; Wang and
Italy
Kashmir

Yin, 2001; Gao et al., 2009), in accordance with the fact that bivalves
Region
Table 4

are abundant and predominant in nearshore environments since the

Late Paleozoic (Clapham and Bottjer, 2007b). However, conodonts are

13
T. Song et al.
Fig. 13. Sketch maps showing the paleogeography and ecology of the characteristic bivalves in the PTTB from the studied area (Paleogeographic maps are modified from Bercovici et al.,
2015). A, the Promyalina – Neoschizodus community (early community); B, the Pteria community (late community).
scarcely found in the littoral clastic facies, especially when no limestone
exists in the PTTB, thus the P-Tr boundary cannot be determined by
conodonts in the littoral clastic facies. Consequently, transitional bi-
valve faunas are used to identify the P-Tr boundary in the littoral clastic
facies and further correlate the boundary into the terrestrial facies (Chu
et al., 2016, 2018), although the exact position of the boundary and the
biochronology of the transitional bivalve fauna are still in debate (Shen
et al., 2011; Zhang et al., 2016; Cui et al., 2017; Chu et al., 2018).
5.4. Implications for biotic recovery from mass extinctions
The P-Tr transitional bivalve fauna in the littoral clastic facies of
southwestern China are taxonomically much richer than in most other
regions and/or other facies (Table 4), such as the Mide, Liangfengya,
Zhongzhai, Shangsi and Huangzhishan sections in South China (Yin and
Wu, 1985; Yang et al., 1987; Yang and Li, 1992; Shen et al., 1995; Lai
et al., 1996; Gao et al., 2009; Chen et al., 2009, 2010; Yu et al., 2010;
Yang, 2015), Guryul Ravine in Kashmir (Nakazawa et al., 1975;
Nakazawa and Kapoor, 1981; Hapoor, 1992) and southern Alps (Broglio
Loriga and Cassinis, 1992; Posenato, 2009). Moreover, the taxonomic
richness of the bivalve fauna in the five studied sections is similar to a
partly time-equivalent bivalve fauna from the microbial limestone of
the basal Luolou Formation in South China, which is composed of 20
species in 17 genera (Hautmann et al., 2015). After the LPME, it is not
until the early Spathian that bivalve's diversity re-reached or exceeded
this level, as exemplified by the early Spathian bivalve assemblage in
Utah, which is composed of 27 species in 18 genera (Hautmann et al.,
14
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx
2013). In South China, numerous evidences imply that there might be
an abrupt pause of environmental devastations between the LPME and
ETME, leaving a refuge zone for diverse metazoans to survive and re-
diversify (Song et al., 2014). The diverse transitional bivalve commu-
nities discovered in this study also can be a response to this short en-
vironmental improvement.
Among the 13 bivalve genera occurring in the PTTB of the five
studied sections, only five are newly evolved genera, whereas the other
eight genera all extended from the pre-extinction Permian time
(Table 1). Accordingly, the high diversity of the P-Tr transitional bi-
valve community did not result from the post-extinction evolution of
new taxa, rather from an ephemeral bloom of holdovers that ultimately
failed to survive and re-diversify (dead clade walking; Jablonski, 2002).
5.5. Ecological and evolutionary implications
The Promyalin-Neoschizodus community represented by the five
lower assemblages from the lower part of the PTTB in the studied area
shows a significant spatial variation along the environmental gradient
from the brackish lagoon to the shallow marine facies. Besides, the
taxonomic diversity and evenness are very high, and shallow infaunal
suspension feeders are considerably abundant in the Promyalina-
Neoschizodus community (Fig. 13A). This transitional bivalve commu-
nity is clearly similar to the Late Permian pre-extinction benthic com-
munity with respect to spatial variation, holdover taxa occurrences,
high diversity and evenness, and presence of abundant infaunal sus-
pension feeders (Yin, 1985; Fang, 2004; Biakov, 2006; Clapham and
T. Song et al.
Bottjer, 2007b; Yang, 2015). In addition, the Promyalina-Neoschizodus
community occurred immediately after the LPME, therefore, the Pro-
myalina-Neoschizodus community in the studied region should belong to
latest Permian for sharing same taxonomic and ecological codes with
the Late Permian fauna.
The Pteria community reconstructed from the five upper assem-
blages from the upper part of the PTTB is much more homogenous
through the various facies. Moreover, the diversity and evenness are
lower and epifaunal suspension feeders are much more abundant in the
Pteria community than in the Promyalina-Neoschizodus community
(Fig. 13B). This Pteria community is exceedingly analogous to the Early
Triassic eurytopic “disaster” taxa (e.g., bivalve Claraia) dominated post-
extinction communities, and both are characterized by uniform dis-
tribution, lower diversity and evenness, and more abundant epifaunal
suspension feeders (Fraiser and Bottjer, 2007; Hofmann et al., 2013;
Huang, 2014; Petsios and Bottjer, 2016; Zhang et al., 2017). Further-
more, the Pteria community is highly homogeneous with the bivalve
Eumorphotis venetiana-Towapteria scythica-Pteria ussurica variabilis as-
semblage at the Huangzhishan section (Chen et al., 2009) and Eu-
morphotis venetiana-Eumorphotis aff. Bokharica assemblage at the Guryul
Ravine section (Table 4; Nakazawa and Hapoor, 1981; Hapoor, 1992),
both of which are from the upper part of the PTTB and of earliest
Triassic age. So the late transitional bivalve community in the studied
region should belong to the earliest Triassic age.
Consequently, a significant taxonomical and ecological turnover of
the transitional bivalve fauna happened between the Promyalina-
Neoschizodus community and Pteria community during the extinction
interval in the studied area (Fig. 13). The Promyalina-Neoschizodus
community in the studied region is of latest Permian age while the
Pteria community should belong to the earliest Triassic. Therefore, the
P-Tr boundary in the studied area most likely is between the two P-Tr
transitional bivalve communities (Fig. 3).
6. Conclusions
A diverse bivalve fauna characterizes the PTTB in the littoral clastic
facies of southwestern China. The P-Tr transitional bivalve fauna is
composed of two communities in stratigraphic succession according to
cluster analyses of the bivalve fossils from the five sections in various
facies, which are Promyalina-Neoschizodus community (early commu-
nity) and Pteria community (late community). These two communities
represent two different ecological stages of bivalves' evolution during
the P-Tr transition. The Promyalina-Neoschizodus community is char-
acterized by the existence of holdover taxa, high taxonomic diversity
and evenness, abundant mobile infaunal suspension feeders and a no-
table spatial variation, and these characters are somewhat comparable
to the Late Permian pre-extinction benthic communities. In contrast,
the Pteria community has a lower diversity and evenness, more abun-
dant epifaunal suspension feeders and a more uniform lateral dis-
tribution along the onshore-offshore gradient than the Promyalina-
Neoschizodus community, thus it is more analogous to the Early Triassic
post-extinction benthic communities. Consequently, the Promyalina-
Neoschizodus community in the studied region is of latest Permian age
while the Pteria community should belong to the earliest Triassic. As a
result, a significant taxonomical and ecological turnover of bivalve
communities occurred between the Promyalina-Neoschizodus commu-
nity and the Pteria community during the extinction interval in south-
western China.
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.palaeo.2018.02.027.
Acknowledgments
Ting Song thanks Dr Michael Hautmann for his critical comments of
the early version of this manuscript. Xincheng Qiu, Yuyang Wu, Gan
Liu, Wenwei Guo and Wenchao Shu are thanked for their assistance in
15
Palaeogeography, Palaeoclimatology, Palaeoecology xxx (xxxx) xxx–xxx
field works.
This study was supported by the National Science Foundation of
China (grants nos. 41661134047, 41530104, 41502012, 41602024)
and the State Key Laboratory of Biogeology and Environmental Geology
(GBL21708). This is a China Postdoctoral Science Foundation funded
international exchange program (to Li Tian). It is also a contribution to
IGCP 630.
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