Australian Journal of Earth Sciences (2009) 56, (815–823)

Ostracods (Crustacea) and water oxygenation in the

earliest Triassic of South China: implications for oceanic
events at the end-Permian mass extinction
M-B. FOREL1*, S. CRASQUIN2, S. KERSHAW3, Q. L. FENG4 AND P-Y. COLLIN5
1
Université Pierre et Marie Curie, UMR 7207, Laboratoire de Micropaléontologie, T.46–56, E.5, case 104, 75252
Paris cedex 05, France.
2
CNRS-UMR7207, Université Pierre et Marie Curie, Laboratoire de Micropaléontologie, T.46–56, E.5, case 104,
75252 Paris cedex 05, France.
3
Institute for the Environment, Brunel University, Uxbridge UB8 3PH, UK.
4
State Key Laboratory of Geo-Processes and Mineral Resources, China University of Geosciences, Wuhan
430074, PR China.
5
Université Pierre et Marie Curie, Laboratoire de Stratigraphie, UMR 7072 ‘Laboratoire de Tectonique,’ T. 56–66,
E.5, case 117, 75252 Paris cedex 05, France.
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Ostracods (Crustacea) are benthic inhabitants well known for their consistent qualities as paleoenvir-
onment markers. In particular, they are reliable indicators of water oxygenation level: filter feeders are
more common in poor oxygen conditions, contrasting with deposit feeders, which are abundant in
well-oxygenated settings. In the Permian/Triassic (P/Tr) boundary transition in the Great Bank of
Guizhou, ostracod species are dominated by deposit feeders, showing well-oxygenated conditions
from the latest Permian, through the extinction level into the earliest Triassic. These results are consistent
with ostracod faunas from northwest Guangxi Province. However, these two examples are in contrast
with coeval ostracods from Sichuan, which show lower-than-normal oxygen levels in the earliest Triassic.
The Great Bank of Guizhou forms an isolated platform in the large Nanpanjiang Basin on the south side
of the South China Block; northwest Guangxi is nearby, in a marginal setting: both faced the
Panthalassa Ocean through the P/Tr boundary times according to several published paleogeographic
reconstructions. In contrast, P/Tr boundary transition rocks in Sichuan Province, located *600 km north
of the Great Bank of Guizhou, lie on the Tethyan side of the South China Block. Both the Great Bank of
Guizhou and the Sichuan sites have earliest Triassic microbialites, but these are profoundly different in
structure and composition. The difference between the two areas may reflect contrasts in the nature of
circulating ocean waters, with reduced levels of oxygenation in the Tethys (Sichuan), associated with
modelled slow circulation, in contrast to better circulated Panthalassa ocean waters (Great Bank of
Guizhou and northwest Guangxi). This also may be an argument to show that low oxygenated, or even
anoxic, waters were not the only reason for the P/Tr boundary crisis.
KEY WORDS: China, Dajiang section, ostracods, Permian, Triassic.

INTRODUCTION
Ostracods are calcified benthic crustaceans which form
well-preserved assemblages in carbonate rocks. As
described by Crasquin-Soleau et al. (2007), ostracods
are known to survive mass extinctions and are therefore
good indicators of paleoenvironments in post-extinction
facies. In the case of the end-Permian mass extinction,
ostracods are of great value in identifying the variation
in oxygen levels of the earliest Triassic, and have been
applied in several sites in the Permian/Triassic (P/Tr)
boundary extinction event, where variation in oxygen
levels have been identified in different places.
This paper presents evidence of oxygen levels from
the P/Tr boundary transition in the southern part of the
South China Block (Figures 1, 2), where ostracods show
that the sequence does not demonstrate low-oxygen
conditions across the extinction horizons, in contrast to
what was expected. This discovery provides new data on
the interpretation of earliest Triassic facies of the South
China Block, and in particular adds data to address a
problem with microbialites in the block.
GEOLOGICAL SETTING
During Permian and Triassic times, the South China
Block was a small continent in the eastern Tethys
(Figure 1). The Great Bank of Guizhou lies in the

*Corresponding author: marie-beatrice.forel@upmc.fr

ISSN 0812-0099 print/ISSN 1440-0952 online ! 2009 Geological Society of Australia

DOI: 10.1080/08120090903002631
 816 M-B. Forel et al.
Nanpanjiang Basin which is a giant embayment in the
southern side of South China Block, along the south side
of the Yangtze Platform (Figure 2). During Late
Permian–Middle Triassic time, the Yangtze Platform
was the site of a thick shallow-marine carbonate
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sedimentation with intermittent siliciclastic input
(Enos 1995; Enos et al. 1997, 1998). The study area is
located a few kilometres north of Luodian, south of
Guizhou province, South China (258330 5600 N, 1068390 4100 E:
Figure 1), where the Great Bank of Guizhou displays
Figure 1 Paleogeographic map dur-
ing Middle Permian (modified
from Crasquin-Soleau et al. 2001)
showing the location of the South
China Block.
Figure 2 Schematic paleogeo-
graphic map of South China Block
during the Changhsingian (mod-
ified from Wang & Jin 2000).
 Permian–Triassic ostracods from South China
open shallow-marine shelf carbonates, overlain by a
thick (*13 m) microbialite carbonate layer.
The Great Bank of Guizhou (70 km east–west and
20 km north–south) originated on an inherited Upper
Permian reef margin, and the rocks are folded into a
syncline in the study area. In the Great Bank of
Guizhou, the east side of the syncline exposes a
continuous Permian–Triassic section (Lehrmann et al.
1998). In the Great Bank of Guizhou and other isolated
carbonate platforms of the Nanpanjiang Basin, the end-
Permian mass extinction is recorded across a broadly
conformable Permian–Triassic boundary which is
marked by an abrupt shift in marine biota and in facies
types. During the earliest Triassic, the Great Bank of
Guizhou maintained a similar morphology but a carbo-
nate microbialite occurred everywhere in the bank
during the basal Griesbachian Hindeodus parvus Zone
(the first occurrence of H. parvus is the index of the
base of the Triassic: Yin et al. 1996; Lehrmann 1999;
Lehrmann et al. 2003), marking an abrupt change in the
marine environmental conditions. In the Great Bank of
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Guizhou, H. parvus occurs through the lowermost
metre-thick microbialite carbonate, above the major
facies change that marks the greatest loss in Permian
fossils. So, the Permo-Triassic boundary ‘event horizon’
occurs at the contact between the Wujiaping Formation
and the microbialite (Lehrmann et al. 2003).
METHODS
Field sampling was undertaken in December 2005,
included logging of the P/Tr boundary transition beds
and closely spaced collection of material for ostracod
analysis and sedimentological study. The samples were
processed by hot acetolysis (Lethiers & Crasquin-Soleau
1988; Crasquin-Soleau et al. 2005). This paper presents
results of investigation on ostracods from Dajiang
section in the central part of the Great Bank of Guizhou
(Figure 2).
RESULTS
Twenty-six samples crossing the P/Tr boundary event
horizon were processed for ostracod analysis. In this
study, the biostratigraphic P/Tr boundary was not
identified because Hindeodus parvus was not found in
the samples, but in all the sections analysed in South
China (Crasquin-Soleau & Kershaw 2005, Crasquin-
Soleau et al. 2006, Kershaw et al. 2007), the first
occurrence of microbialites are of lowermost Triassic
age, so it is clear that the P/Tr boundary interval is
included within the sample set. Except for one sample
(05PAJ34 in Triassic sediments), all the samples yielded
ostracods. Figure 3 shows a general field log of the
sequence and the ostracod distribution and taxa.
Eighty-eight species of ostracods belonging to 25
genera were extracted from the latest Permian Changh-
singian limestones sampled at Dajiang. This assemblage
is characterised by the preponderance of the genus
Bairdia with 31 species. Above the event horizon, the
post-extinction assemblage is noteworthy for its diver-
817
sity (30 species belonging to nine genera). The
Early Triassic (Induan) is well-known for its poverty.
However, the lowermost part (Griesbachian) contains
ostracods (Crasquin-Soleau et al. 2004, 2006; Crasquin-
Soleau & Kershaw 2005). One ostracod species has been
found both in Late Permian and Early Triassic, thus
crossing the P/Tr boundary: Paracypris gaetanii
Crasquin-Soleau 2006, discovered in the Griesbachian
of Guangxi (Crasquin-Soleau et al. 2006), is also found in
Dajiang (Figures 3, 4).
Ostracods and paleoenvironment
In general terms, the great majority (98%) of ostracods
are benthic inhabitants, thus dependent on paleoenvir-
onmental changes in sea bottom. In particular, they are
sensitive to variations of bathymetry, salinity, tempera-
ture, nutrients and oxygen level of the water.
The paleoecological requirements of genera, families
and/or superfamilies are now relatively well known for
the Late Paleozoic and Early Triassic (Peterson &
Kaesler 1980; Costanzo & Kaesler 1987; Melnyk &
Maddocks 1988; Crasquin-Soleau et al. 1999) and the
main features are summarised here. The Bairdioidea
are present in shallow to deep, open-marine environ-
ments with normal salinity and oxygenation. The
Kloedenelloidea characterise very shallow, euryhaline
environments. The Kirkbyidae/Amphissitidae group
develops in marine restricted environments while the
Cavellinidae are adapted to euryhaline shallow to very
shallow environments. The Bythocytheridae are indica-
tive of deeper environments (external platform).
All the ostracods reported here are typical of inter-
tropical warm waters (Crasquin-Soleau et al. 1999, 2004).
The distribution of species by families/superfamilies is
shown in Figure 5. Several features should be noted: (i)
there is no significant difference between Late Permian
and Early Triassic associations, in terms of composition
and paleoecological affinities; (ii) both Permian and
Triassic assemblages are dominated (never less than
50% of species) without restriction by open-marine
forms, the Bairdioidea; (iii) the late Permian assem-
blages present shallower forms than Triassic ones, with
Kloedenelloidea (sample 05PAJ21), Amphissitidae and
Kirkbyidae (samples 05PAJ20, 22, 24) and Cavellinidae
(samples 05PAJ21, 24 and 25); (iv) the last levels of the
Permian (05PAJ 26, 27, 28, 29) are represented by only
Bairdioidea (except two species in level 05PAJ28); (v) the
ostracod fauna above the event horizon is exceptionally
well represented and diverse for the considered
time slice, as mentioned above; and (vi) the Early
Triassic assemblages are characteristic of open-marine
environments.
Ostracods and oxygen concentration
A direct relationship between the modes of feeding/
respiration in certain post-Paleozoic ostracods and
oxygen concentration in the water has been established
(Whatley 1990, 1992). Ostracods present two different
modes of feeding and respiration: deposit-feeding
and filter-feeding. Whatley (1990, 1992) showed that,
in contrast to deposit-feeders, filter-feeding ostracods
 818 M-B. Forel et al.
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Figure 3 Lithostratigraphy and stratigraphic distribution of ostracod genera in Dajiang and evolution of the percentage of
filter-feeding ostracods.

create a permanent and enhanced circulation over their places where, oxygen levels are low. Lethiers & Whatley
ventral respiratory surface by virtue of their larger (1994) demonstrated the relevance of these observations
number of branchial plates. This form of feeding confers in the Paleozoic. They showed the increase in filter
advantages on such ostracods during times when, or in feeders matches falling levels of oxygen and vice versa.
 Permian–Triassic ostracods from South China 819
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Figure 4 Ostracods from the Permian–Triassic interval from the Dajiang section, South China. Scale bar is 100 mm. (1)
Bairdiacypris ottomanensis 2004: sample 05PAJ41, #148. (2) Bairdia sp. 3: sample 05PAJ42, #125. (3) Paracypris sp. sensu Chen
1982: sample 05PAJ39, #216. (4) Paraparchitidae indet.: sample 05PAJ44, #82. (5) Paracypris gaetanii Crasquin-Soleau 2004:
sample 05PAJ22, #645. (6) Sulcella sp. 1: sample 05PAJ25, #508. (7) Amphissites? sp. 2: sample 05PAJ24, #571. (8) Petasobairdia
sp. 2: sample 05PAJ26, #609.

Figure 5 Distribution of ostracod species in percentage of families or superfamilies in the Dajiang section.

Neritic filter-feeding ostracods of the Late Paleozoic and
Early Triassic present here comprise the Palaeocopida
(with Paraparchitoidea, Kloedenelloidea, Kirkbyoidea,
Amphissitoidea) and Platycopida. Deposit feeders com-
prise the Podocopida (with Bairdioidea, Cypridoidea,
Cytheroidea, Siggiloidea).
The Lethiers & Whatley (1994) interpretative model
(Figure 6) allows the determination of approximate
oxygen levels in neritic environments using the percen-
tage of filter-feeding ostracods. When the percentage of
filter-feeding ostracods reaches 60%, the oxygen con-
centration should be around 3.5–3.6 mL/L. If the
 820 M-B. Forel et al.
Figure 6 Lethiers–Whatley model of oxygen levels linked to ostracod abundance and type, with the estimations of oxygen
levels of the samples processed in this study (Great Bank of Guizhou, Dajiang) and in Sichuan (Crasquin-Soleau & Kershaw
2005).
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percentage exceeds 85–90%, kenoxia may have been
reached. The term ‘kenoxia’ is used to denote an event
when the oxygen level is below 1 mL/L. The term
‘anoxia’ reflects a complete lack of oxygen. When anoxia
is reached, no ostracods should survive as was shown in
the Dienerian–Smithian of Guangxi (Crasquin-Soleau
et al. 2006) which corresponded to the maximum poverty
of ostracods after the P/Tr boundary events (Crasquin-
Soleau et al. 2007).
We applied this model to the Dajiang section. The
Bairdioidea are deposit feeders, while all the others
groups represented here are filter-feeding ostracods. As
shown by the data presented in this study, there is never
less than 50% of the Bairdioidea in the assemblages, i.e.
the oxygen level was never lower than 4.1–4.2 mL/L. The
percentages of filter-feeding ostracods are shown in
Figure 3. Several peaks are observed. These peaks are
interpreted here as indicating fluctuations in oxygen
concentration. However, oxygen levels did not reach
dysoxia/kenoxia, so the environments did not become
anoxic, at least while the ostracods were alive.
DISCUSSIONS
The discovery in this study of an assemblage of
ostracods that demonstrates well-oxygenated conditions
in Great Bank of Guizhou during the latest Permian and
earliest Triassic has important implications for the
nature of the sedimentary environments throughout the
P/Tr boundary transition in this area. There is now a
range of published studies supporting the view that low-
oxygen conditions played an important part in the
process of the end-Permian mass extinction (Wignall &
Hallam 1992; Isozaki 1994, 1997; Wignall & Twitchett
1996, 2002); furthermore, evidence from biomarkers
suggests that an unusual Early Triassic marine ecosys-
tem existed (Grice et al. 2005a, b). On land, geological
evidence (Sephton et al. 2005) and modelling (Berner
2001, 2002) indicate reduced atmospheric oxygen, prob-
ably associated with the Siberian Traps eruptions. In
contrast to this, there is also growing evidence that not
all areas were subject to lowered oxygen levels in the
Griesbachian. No drop of oxygen content was registered
in earliest Triassic of Tibet (Wignall & Newton 2003), in
Western Taurus, Turkey (Crasquin-Soleau et al. 2004) or
northwest Iran in Abadeh (Kozur 2007).
Kershaw et al. (2007) suggested that the earliest
Triassic microbialites, formed in the immediate after-
math of the mass extinction, were controlled by a com-
bination of low-oxygen marine waters with bicarbonate-
rich supersaturated water upwelled from the deep ocean
during ocean overturn at the P/Tr boundary event
horizon. This proposal was based on: (i) the evidence
presented by Riding (2005) and Riding & Liang (2005)
that supersaturation is the key driving force for
microbial precipitation; and (ii) the current knowledge
of earliest Triassic microbialites which indicates that
their occurrence appears to be largely linked to low-
oxygen conditions (see Kershaw et al. 2007 for full
discussion). A key component in this interpretation is
the abundant evidence of ocean overturn in the earliest
Triassic, drawing anoxic, bicarbonate-rich deep water
to the ocean surface. However, there is also a range of
recent studies which show a drop in atmospheric
oxygen through Late Permian to earliest Triassic
(Weidlich et al. 2003), so that shallow marine
anoxia probably was caused by a combination of both
processes.
The results from the present study demonstrate that
microbialites grew in well-oxygenated water in this
area. Therefore, there are at least two important points
in relation to the upwelling model: (i) if upwelling of
bicarbonate-supersaturated ocean water was a principal
driving force for the formation of the earliest Triassic
microbialites, then the presence of low-oxygen condi-
tions was a subsidiary component of the controls on the
microbialites; and (ii) if lower-than-normal oxygen
conditions played a part in the formation of microbia-
lites (evidence for which is in the Sichuan localities)
then low-oxygen environments were patchy. There is
further evidence of patchy oxygenation [proposed by
Wignall & Newton 2003; Crasquin-Soleau et al. 2004;
Kozur 2007 and also in model results (Kidder & Worsley
 Permian–Triassic ostracods from South China
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Figure 7 Paleogeographic map (adapted from Crasquin-Soleau et al. 2001) with oceanic circulation and sites of upwelling (U)
and sinking (S) (Kidder & Worsley 2004). Inset. Enlargement of South China Block (SCB) showing location of the Great Bank
of Guizhou and Sichuan.
2004; Kiehl & Shields 2005)]. Kershaw et al. (2007) used
the evidence presented by the model results and data to
propose supersaturation was mostly concentrated in
Tethys, as a result of the paleogeographic near-isolation
from Panthalassa, and that overturn of the ocean took
place in the earliest Triassic, providing the key stimulus
for microbialite growth.
Kershaw et al. (2007) particularly drew attention to
the microbialite structures of Sichuan and the Great
Bank of Guizhou: microbialite types are strongly con-
trasting between these two closely located sites, yet
there is clear evidence of lower-oxygen conditions in
Sichuan (Crasquin-Soleau & Kershaw 2005). Such in-
formation emphasises the importance of carbonate
saturation as a control on microbialite formation
(Riding 2005). Kershaw et al. (2007) did not have an
explanation for the differences between sites that are
only about 600 km apart, with no proven land bridge
between them. The present data now provide a potential
solution to this problem, by demonstrating that the
Great Bank of Guizhou in the Nanpanjiang Basin facing
the open ocean of Panthalassa was bathed by well-
oxygenated water, but sites in Sichuan were influenced
by lower levels of oxygen in the seawater in sites on the
Tethyan-facing shelf of the South China Block, which
faced west, since South China Block was rotated around
908 anticlockwise from its present orientation (Figure 7).
The ostracod data accumulating from the South
China Block therefore contribute to the growing dataset
to demonstrate that the development of low-oxygen
conditions in the P/Tr boundary transition was con-
siderably heterogeneous in space. Wignall & Newton
(2003) also showed a diachronism in the history of
anoxia, involving deep shelf sites within Tethys and in
Panthalassa. Modelling by Kidder & Worsley (2004) and
Kiehl & Shields (2005) indicated that Tethys was a slow-
moving, strongly stratified ocean, where low-oxygen
821
conditions would be expected to develop, during ocean
overturn, to a greater extent than happened in Pantha-
lassa. The ostracod data are beginning to provide a test
of these models from interpretation of their oxygen
requirements. While the models are sustainable in
general terms, in detail individual areas depart from
the models. Thus, more data are required to develop a
full test of these models.
CONCLUSIONS
(1) An assemblage of ostracods through the strati-
graphic interval of latest Permian to earliest Triassic
from the central part of the Great Bank of Guizhou,
south China, shows a significant change in taxonomic
composition of ostracods at the extinction event
horizon.
(2) Despite the abrupt change at species level, the
ostracod assemblages below and above the event hor-
izon present the same paleocological characteristics and
are dominated by deposit-feeder forms, which demon-
strate that the seafloor was well-oxygenated throughout
the P/Tr boundary transition.
(3) The South China Block was rotated about 908
anticlockwise relative to its present orientation during
the P/Tr boundary transition, so that the Great Bank of
Guizhou lay in the Nanpanjiang Basin, open to Pantha-
lassa, in contrast to shelf sediments in Sichuan which
faced Tethys. In the Great Bank of Guizhou, earliest
Triassic microbialites formed in well-oxygenated
waters. In Sichuan, time-equivalent deposits show that
microbialites formed in conditions of lowered oxygen,
but not dysoxic. The differences in microbialites be-
tween the two sides of the South China Block may be due
to oxygenation differences of waters bathing the shelf. If
upwelled bicarbonate-rich waters were a key component
 822 M-B. Forel et al.
of control of microbialite forms, then the Sichuan sites
had an additional component of low-oxygen states, while
the Great Bank of Guizhou did not.
(4) This study provides information on the nature of
paleocirculation through the P/Tr boundary times. In
general terms, the differences noted between sites
bathed by the Tethys Ocean and Panthalassa Ocean
agree with models showing a sluggish Tethyan circula-
tion leading to poorly oxygenated surface and bottom
waters, added to the fact that the oxygen concentration
in the atmosphere was also reduced. Circulation in
Panthalassa appears to have been more active, allowing
good ventilation of waters brought to the surface by
upwelling surrounding the South China Block. These
two modes of circulation seem to indicate a relative
isolation of Tethyan water masses from Panthalassa.
(5) Literature survey shows that other areas of the
world varied in the extent to which the earliest Triassic
deposits were oxygenated, and reinforce the view that
oxygenation was patchy. Thus, views that the end-
Permian extinction was driven only by oxygenation of
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Earth’s surface environments are an oversimplification,
and show that the extinction was a complex event.
ACKNOWLEDGEMENTS
We are very grateful to Zhong Qiang Chen (University of
Western Australia, Australia) and Michael Schudack
(Freie Universität of Berlin, Germany) for their con-
structive and helpful reviews. This study is part of the
IGCP 572 ‘Permian–Triassic ecosystems.’
REFERENCES
BERNER R. A. 2001. Modelling atmospheric O2 over Phanerozoic time.
Geochimica et Cosmochimica Acta 65, 685–694.
BERNER R. A. 2002. Examination of hypotheses for the Permo-
Triassic boundary extinction by carbon cycle modelling. Pro-
ceedings of the National Academy of Sciences USA 99, 4172–
4177.
COSTANZO G. V. & KAESLER R. L. 1987. Changes in Permian marine
Ostracode faunas during regression, Florena Shale, Northeast-
ern Kansas. Journal of Paleontology 61, 1204–1215.
CRASQUIN-SOLEAU S., BROUTIN J., BESSE J. & BERTHELIN M. 2001.
Ostracodes and paleobotany from the Middle Permian of
Oman: implications on Pangea reconstruction. Terra Nova 13,
38–43.
CRASQUIN-SOLEAU S., BROUTIN J., ROGER J., PLATEL J-P., AL HASHMI
A., ANGIOLINI L., BAUD A., BUCHER H. & MARCOUX J. 1999. First
Permian ostracode fauna from the Arabian Plate (Khuff Forma-
tion, Sultanate of Oman). Micropaleontology 45, 163–182.
CRASQUIN-SOLEAU S., GALFETTI T., BUCHER H. & BRAYAR A. 2006.
Early Triassic ostracods from Guangxi Province, South
China. Rivista Italiana di Paleontologia e Stratigrafia 112,
55–75.
CRASQUIN-SOLEAU S., GALFETTI T., BUCHER H., KERSHAW S. & FENG Q.
2007. Ostracod recovery in the aftermath of the Permian–
Triassic crisis: Palaeozoic–Mesozoic turnover. Hydrobiologia
585, 13–27.
CRASQUIN-SOLEAU S. & KERSHAW S. 2005. Ostracod fauna from the
Permian–Triassic Boundary Interval of South China (Huaying
Mountains, eastern Sichuan Province): palaeoenvironmental
significance. Palaeogeography, Palaeoclimatology, Palaeoecology
127, 131–141.
CRASQUIN-SOLEAU S., MARCOUX J., ANGIOLINI L., RICHOZ S., NICORA
A., BAUD A. & BERTHO Y. 2004. A new ostracode fauna from the
Permian –Triassic boundary in Turkey (Taurus, Antalya
Nappes). Micropaleontology 50, 281–295.
CRASQUIN-SOLEAU S., VASLET D. & LE NINDRE Y.M. 2005. Ostracods
from Permian–Triassic boundary in Saudi Arabia (Khuff
Formation). Palaeontology 48, 853–868.
ENOS P. 1995. The Permian of China. In: Scholle P. A., Peryt T. M. &
Ulmer-Scholle D. S. eds. The Permian of Northern Pangea,
sedimentary basins and economic resources 2, pp. 225–256. Spring-
er Verlag, Berlin.
ENOS P., WEI J. & YAN Y. 1997. Facies distribution and retreat of
Middle Triassic platform margin, Guizhou Province, south
China. Sedimentology 44, 563–584.
ENOS P., WEI J. & LEHRMANN D. J. 1998. Death in Guizhou—Late
Triassic drowning of the Yangtze carbonate platform. Sedimen-
tary Geology 118, 55–76.
GRICE K., CAO C., LOVE G. D., BOTTCHER M. E., TWITCHETT R. J.,
GROSJEAN E., SUMMONS R. E., TURGEON S. C., DUNNING W. & JIN
Y. 2005a. Photic zone euxinia during the Permian–Triassic
superanoxic event. Science 307, 706–709.
GRICE K., TWITCHETT R. J., ALEXANDER R., FOSTER C. B. & LOOY C.
2005b. A potential biomarker for the Permian–Triassic ecological
crisis. Earth and Planetary Sciences Letters 236, 315–321.
ISOZAKI Y. 1994. Superanoxia across Permo-Triassic boundary:
record in accreted deep-sea pelagic chert in Japan. In: Beau-
champ B., Embry A. & Glass D. eds. Pangea: global environment
and resources, pp. 805–812. Canadian Society of Petroleum
Geologists Memoir 17.
ISOZAKI Y. 1997. Permo-Triassic boundary superanoxia and stratified
superocean: records from lost deep sea. Sciences 276, 235–238.
KERSHAW S., LI Y., CRASQUIN-SOLEAU S., FENG Q., MU X., COLLIN P-Y.,
REYNOLDS A. & GUO L. 2007. Earliest Triassic Microbialites in the
South China Block and other areas: controls on their growth and
distribution. Facies 53, 409–425.
KIDDER D. L. & WORSLEY T. R. 2004. Causes and consequences of
extreme Permo-Triassic warming to globally equable climate
and relation to the Permo-Triassic extinction and recovery.
Palaeogeography, Palaeoclimatology, Palaeoecology 203, 207–237.
KIEHL J.T. & SHIELDS C. A. 2005. Climate simulation of the latest
Permian: implications for mass extinction. Geology 33, 757–760.
KOZUR H. 2007. Biostratigraphy and event stratigraphy in Iran
around the Permian–Triassic Boundary (P/Tr boundary): im-
plications for the causes of the P/Tr boundary biotic crisis.
Global and Planetary Change 55, 155–176.
LEHRMANN D. J. 1999. Early Triassic calcimicrobial mounds and
biostromes of the Nanpanjiang basin, south China. Geology 27,
359–362.
LEHRMANN D. J., PAYNE J. L., MONTGOMERY P., WEI J., YU Y., XIAO J.
& ORCHARD M. J. 2003. Permian–Triassic boundary sections from
shallow-marine carbonate platforms of the Nanpanjiang basin,
south China: implications for oceanic conditions associated with
the end-Permian extinction and its aftermath. Palaios 18, 138–
152.
LEHRMANN D. J., WEI J. & ENOS P. 1998. Controls on facies
architecture of a large Triassic carbonate platform: the Great
Bank of Guizhou, Nanpanjiang Basin, South China. Journal of
Sedimentary Research 68, 311–326.
LETHIERS F. & CRASQUIN-SOLEAU S. 1988. Comment extraire des
microfossiles à tests calcitiques de roches calcaires dures. Revue
de Micropaléontologie 31, 56–61.
LETHIERS F. & WHATLEY R. 1994. The use of Ostracoda to reconstruct
the oxygen levels of the Late Paleozoic oceans. Marine Micro-
paleontology 24, 57–69.
MELNYK D. H. & MADDOCKS R. F. 1988. Ostracode biostratigraphy of
the Permo-Carboniferous of Central and North-Central Texas,
part I: Paleoenvironmental framework. Micropaleontology 34,
1–20.
PETERSON R. M. & KAESLER R. L. 1980. Distribution and diversity
of ostracodes assemblages from the Hamlin Shale and the
Americus limestones (Permian, Wolfcampian) in northern
Kansas. University of Kansas Paleontological Contributions 100,
1–26.
RIDING R. 2005. Phanerozoic reefal microbial carbonate abundance:
comparisons with metazoan diversity, mass extinction events,
and seawater saturation state. Revista Española de Micropaleon-
tologia 37, 23–39.
RIDING R. & LIANG L. 2005. Geobiology of microbial carbonates:
metazoan and seawater saturation state influences on secular
trends during the Phanerozoic. Palaeogeography, Palaeoclima-
tology, Palaeoecology 219, 101–115.
 Permian–Triassic ostracods from South China
SEPHTON M. A., LOOY C. V., BRINKHUIS H., WIGNALL P. B., DE LEEUW
J. W. & VISSCHER H. 2005. Catastrophic soil erosion during the
end-Permian biotic crisis. Geology 33, 941–944.
WANG Y. & JIN Y. 2000. Permian palaeogeographic evolution of the
Jiangnan Basin, South China. Palaeogeography, Palaeoclimatol-
ogy, Palaeoecology 160, 35–44.
WEIDLICH O., KIESSLING W. & FLÜGEL E. 2003. Permian–
Triassic boundary interval as a model for forcing ecosystem
collapse by long-term atmospheric oxygen drop. Geology 31, 961–
964.
WHATLEY R. C. 1990. Ostracoda and global events. In: Whatley R. C. &
Maybury C. eds. Ostracoda and global events, pp. 3–24. Chapman
& Hall, London.
WHATLEY R. C. 1992. The Platycopid signal: a means of detecting
kenoxic events using ostracoda. Journal of Micropalaeontology
10, 181–185.
WIGNALL P. B. & HALLAM A. 1992. Anoxia as a cause of the Permian–
Triassic mass extinction: facies evidence from northern Italy
and Western United States. Palaeogeography, Palaeoclimatology,
Palaeoecology 93, 21–46.
Downloaded By: [BIUS Jussieu/Paris 6] At: 12:50 4 March 2011
823
WIGNALL P. B. & NEWTON R. 2003. Contrasting deep-water records
from the Upper Permian and Lower Triassic of South Tibet and
British Columbia: evidence for a diachronous mass extinction.
Palaios 18, 153–167.
WIGNALL P. B. & TWITCHETT R. J. 1996. Oceanic anoxia and the end
Permian mass extinction. Science 272, 1155–1158.
WIGNALL P. B. & TWITCHETT R. J. 2002. Extent, duration, and nature
of the Permian–Triassic superanoxic event. In: Koeberl C. &
MacLeod K.C. eds. Catastrophic events and mass extinctions:
impacts and beyond, pp. 395–413. Geological Society of America
Special Paper 356.
YIN H. F., WU X. B. & DING M. H. 1996. The Meishan section,
candidate of the Global Stratotype section and Point of Permian–
Triassic boundary. In: Yin H. F. ed. The Palaeozoic–Mesozoic
boundary candidates of the Global Stratotype Section and Point of
the Permian–Triassic boundary, pp. 31–48. China University of
Geosciences Press, Wuhan.
Received 13 September 2007; accepted 3 April 2009