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Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176 – 187

Onset of biotic and environmental recovery from the end-Permian

mass extinction within 1–2 million years: A case study of
the Lower Triassic of the Meishan section, South China
Z.Q. Chen a,⁎, Jinnan Tong b , Kunio Kaiho c , Hodaka Kawahata d
School of Earth and Geographical Sciences, University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia
The Key Laboratory of Bio-geology and Environmental Geology and Faculty of Earth Science,
China University of Geosciences Wuhan 430074, China
Institute of Geology and Paleontology, Tohoku University, Sendai 980-8578, Japan
Ocean Research Institute, University of Tokyo, 1-15-1 Minamidai, Nakano-ku, Tokyo 164-8639, Japan
Accepted 30 November 2006


Paleontological, sedimentary and geochemical data obtained from the Lower Triassic strata of the Meishan section, South
China, suggest that onset of biotic recovery coincided with amelioration of paleoceanographic conditions and an increase in ocean
productivity. The Griesbachian Meishanorhynchia association from Meishan indicates biotic recovery between the recovery stages
1 and 2. Both the initial biotic recovery and palaeoceanic environmental amelioration occurred within 1–2 myr after the end-
Permian mass extinction in the late Griesbachian.
© 2007 Elsevier B.V. All rights reserved.

Keywords: End-Permian; Early Triassic; Mass extinction; Biotic recovery; Meishan; South China

1. Introduction cause, process and pattern of the delayed recovery

following the end-Permian crisis remain speculative
The Permian–Triassic (P–Tr) transition resulted in the despite several recent works (e.g. Twitchett et al., 2004;
most devastating biotic crisis in Phanerozoic history and Bottjer and Gall, 2005, and references therein). Payne et al.
drastically altered the balance of the biosphere and its (2004) proposed that multiple environmental perturbations
subsequent evolution. The severity of this catastrophe as during the Early Triassic may have played a critical role in
well as the prolonged post-extinction recovery of biota in delaying the post-extinction recovery. Their study supports
the Triassic have puzzled paleontologists. It has been the view that deleterious environmental conditions that
proposed that the biotic recovery did not occur until the prevailed in most stages of the Early Triassic may have, at
early Middle Triassic (5–10 myr later) (Erwin, 1993; least in part, controlled the delayed recovery (Hallam,
Hallam and Wignall, 1997; Erwin, 1998). However, the 1991; Hallam and Wignall, 1997; Woods et al., 1999;
Bottjer, 2004, 2005; Fraiser and Bottjer, 2005b; Fraiser et
⁎ Corresponding author. Tel.: +61 864881924; fax: +61 864881037. al., 2005). This also means that biotic recovery could have
E-mail address: (Z.Q. Chen). occurred earlier if Early Triassic oceanic habitats became
0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.
Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187 177

locally and/or intermittently conducive to biotic prolifer- levels, simple fodinichnia (with diameters b5 mm), and
ation. Recently, Twitchett et al. (2004) documented an shelly faunas having very low diversity, high dominance
example in which they proposed that a diverse and and small sizes. The recovery stages 2 and 3 are character-
complex biota recovered in shallow, oxygenated environ- ized by an increase in tiring levels, diversity of trace and
ments of the Neo-Tethys oceans in the middle-late body fossil assemblages, and shell sizes. In addition, the
Griesbachian (Twitchett et al., 2004). Other recent studies presence of small, low diversity vertical domichia and
have also described rapid recovery of fossil groups such as slightly diverse and large (but b20 mm) inchotaxa marks
ammonoids (e.g. McGowan, 2004, 2005), bivalves the recovery stages 2 and 3, respectively. In the benthic
(McRoberts, 2001), gastropods (Wheeley and Twitchett, recovery Stage 4 the epifaunal and infaunal tiering
2005; Nützel, 2005) and brachiopods (Chen et al., 2005b). increases significantly, inchnotaxa become diversified,
Here, we also document an example of rapid recovery complex and large (with diameters N 20 mm), and shelly
from the Meishan section, South China (Fig. 1). faunas embrace high diversity, low dominance and large
At the present, the discussions about the definition of sizes (more details see Twitchett, 2006). This model
biotic recovery are active despite several past reviews attempts to quantify biotic recovery process, and thus
(i.e., Kauffman and Erwin, 1995; Erwin and Pan, 1996; permits comparison of the recovery rates between
Erwin, 1998, 2000; Twitchett et al., 2004; Twitchett, different environmental settings or different regions
2005, 2006). Of these, Twitchett et al. (2004) approach (climate zones) in the aftermath of the P/Tr event.
that subdivides the post-extinction interval into four At Meishan, Chen et al. (2002) reported the oldest
recovery stages is followed herein. Summarizing the Mesozoic brachiopod Meishanorhynchia from the upper
paleoecologic features (including ichnology, dominance, Griesbachian succession. These authors considered that
evenness, body size) of several key post-Permian faunas, Meishanorhynchia is phylogenetically close to the Meso-
Twitchett et al. (2004) defined post-extinction recovery zoic and later brachiopods and its occurrence in the late
patterns composed of four stages from the initial post- Griesbachian signaled initial recovery of brachiopods in the
extinction aftermath to the final recovery of the benthic Mesozoic from taxonomic or systematic points of views.
community. Several key paleoecologic criteria are However, they did not document the community ecologic
employed to discriminate these recovery stages: (1) features of the Meishanorhynchia and its associated faunas.
tiering levels, (2) the presence/absence of key ichotaxa, The ecologic method has been treated as a very important
(3) body (or burrow) size and (4) dominance and evenness measure of biotic recovery following mass extinctions
of shelly assemblages (Twitchett et al., 2004; Twitchett, (Droser et al., 2000; McGhee et al., 2004; Twitchett et al.,
2005, 2006). Of these, the recovery stage 1 occurs in the 2004; Twitchett, 2005). Accordingly, we have restudied the
immediate aftermath with the lowest/shallowest tiering paleontology, sedimentology and geochemistry of the

Fig. 1. Permian–Triassic succession exposed at the Meishan section (Quarry B) showing the P–Tr boundary, end-Permian mass extinction horizon
and initial biotic recovery horizon.
178 Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187

Fig. 2. Composite Lower Triassic lithostratigraphy, biostratigraphy, and sequence stratigraphy of the Meishan section showing lithologic variation,
fossil stratigraphic distributions, stacking pattern of PACs, and possible relative sea-level change curves. Note sequence stratigraphy and inferred
relative sea-level change curves follow Tong and Yin (1998); stratigraphic distributions of conodonts and microflora see Zhang et al. (2007-this
volume); radiometric ages dated from clay beds 25, 28, 37, and 48 are 251.4 ± 0.3 myr, 250.7 ± 0.3 myr, 250.4 ± 0.5 myr, and 250.2 ± 0.2 myr,
respectively (Bowring et al., 1998). Abbreviations: Fm. = Formation, Changhs. = Changhsing, PAC = Punctuated, Aggradational Cycles, P.s. =
Parasequence sets, Seq. = Sequence, SB = sequence boundary, TST = Transgressive System Tract, mfs = maximum flooding surfaces, HST =
Highstand System Tract, b = basin, o = offshore, bw = below wave base, w = fairweather wave base, om = offshore mudstone facies, bs = basinal
black shale facies, ow = offshore wackestone facies, os = offshore siltstone facies; ew = epeiric sea wackestone facies, HCS = hummocky cross
stratification, hb = horizontal bedding.

Lower Triassic strata of the Meishan section (Fig. 1) to radiometric ages have been obtained from numerous
make comparisons with these published recovery patterns claystone beds in this section (Bowring et al., 1998;
(i.e., Twitchett et al., 2004; Twitchett, 2006). In addition, Mundil et al., 2001, 2004). Thus, in this combined study we
Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187 179

are proposed by Zhang et al. (2007-this volume) for Beds

28–51 and Beds 52–62, respectively (Fig. 2). Brachiopod
Meishanorhynchia occurs in Beds 53–55 (equating
to part of Bed 36 of Chen et al., 2002). The asso-
ciated conodonts, ammonoids and bivalves belong to the
tulongensis–carinata conodont zone (Zhang et al., 2007-
this volume), and the Ophiceras–Lytophiceras ammo-
noid zone (or the upper Ophiceras–Claraia griesbachi
zone) (Yin et al., 2001). These biozones suggest a late
Griesbachian age (Tong and Yin, 2002; Zhang et al.,
2007-this volume). The overlying Helongshan Formation
is dominated by thinly bedded limestone; its lower part is
latest Griesbachian in age, and its upper part is Dienerian
(Ding and Wei, 1991; Zhang et al., 2007-this volume).

3. Analytical techniques and paleoecologic and

sedimentologic descriptions

3.1. Paleontology
Fig. 3. Slab surface showing the Meishanorhynchia association from
the upper Yinkeng Formation. Scale bar is 4 mm long. m = Meisha- Macrofossils were collected throughout the Lower
norhynchia, o = ophiceratids, Ly = Lytophiceras. Triassic strata. Most specimens were extracted from a
50 cm thick interval (Beds 53–55) ∼ 9.50–10.00 m above
the end-Permian mass extinction horizon (Figs. 1 and 2).
are able to evaluate biotic recovery rate in conjunction with All specimens were recovered from the surfaces of
environmental changes that potentially affected the Triassic mudstone or muddy limestone slabs split parallel along
recovery. bedding planes to expose the concentrated shell material
(Fig. 3). Specimens derived from 28 slabs, each about 20–
2. Stratigraphic setting 30 cm long and 10–15 cm wide, are included in the
quantitative analysis (Fig. 4). Tiny brachiopod specimens
The Meishan section (Fig. 1) is the well known (b 5 mm in length and 5 mm in width) were extracted from
Global Boundary Stratotype Section and Point for the each sample by physically splitting and abrading the
P–Tr boundary (Yin et al., 2001). The P/Tr boundary is samples. Preservation is variable; brachiopod specimens
placed in the middle part of Bed 27 as defined by the comprise complete shells, whereas ammonoids and
first occurrence of conodont species Hindeodus parvus bivalves are molds and fossil fishes are represented by
(Yin et al., 2001; Fig. 2), whereas the end-Permian mass
extinction horizon coincides with the base of Bed 25
some 10–15 cm below of the P/Tr boundary (Jin et al.,
2000; Kaiho et al., 2001, 2006). In this section the
Lower Triassic strata have received far less attention
than the well-studied P/Tr boundary beds (e.g. Chen
et al., 2002; Tong and Yin, 2002; Zhang et al., 2007-this
volume) even though exposure of the Lower Triassic
succession is also spectacular (Fig. 1).
The succession is divided into the Yinkeng and
Helongshan Formations in ascending order (Fig. 2).
Lithologically, the Yinkeng Formation is characterized by
interbedded black shale, greenish gray mudstone, gray
calcareous mudstones, gray marls, and pale muddy
limestone (Fig. 2). Apart from the Hindeodus parvus Fig. 4. Abundance of the Meishanorhynchia association. Abundance
conodont zone, the Isarcicella staeschei–H. isarcica and of each taxon is represented by the percentage of individuals of each
Neogondolella tulongensis–N. carinata conodont zones taxon among 143 individuals of the association.
180 Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187

fragmented scales. Identification of taxa is based on gross the Punctuated Aggradational Cycles (PACs) (Goodwin
morphology along with details of surface ornament and and Anderson, 1985) recognized by Tong and Yang
interior features. Details of the taxonomy are available (1998) from the Meishan section. Seven cycle types are
from ZQC on request. It should be noted that fishes are recognized (Table 1) and all are characterized by
excluded from quantitative analysis because hundreds of relatively deeper water facies overlain by shallower
scales may come from a single individual fish. Additional facies (Tong and Yang, 1998). Sequence stratigraphic
specimens, primarily of Meishanorhynchia, were also subdivision and interpreted relative sea-level change
collected from five layers within Beds 53–55 (see Chen curves are after Tong and Yin (1998). At the lower
et al., 2002) in order to facilitate identification and quanti- Yinkeng Formation (Beds 31–40), the lower units of
tative paleoecology. All Meishan specimens pertaining to PACs dominated by black shale become thicker and
this study are housed at the School of Earth and Geograp- thicker up section, while the upper units thin up-section.
hical Sciences, The University of Western Australia, Perth, As a result, the entire cycle is dominated by finely
Australia. laminated black shale (i.e., PACs 16–18), indicating that
Diversity indices (Shannon and Simpson indices and the depositional environment deepens upwards. Thus,
Dominance) were calculated using the paleontological these PACs show an overall regradational stacking
statistical analysis package PAST (Hammer et al., pattern characterizing the Transgressive System Tract
2001). The Shannon index evaluates the diversity of (TST) (Tong and Yin, 1998). Beds 41–53 are dominated
Meishanorhynchia and its associations, whereas both by the ML type PACs (PACs 19–43) in the lower and
Simpson index and Dominance (= 1 − Simpson index) ML type PACs (PACs 44–61) in the upper. The former
evaluate the evenness of the association. The analytical cycles are dominated by a thick upper unit of greenish
methods are described in Hammer et al. (2001), and gray mudstone; while the latter are typified by thinly
were also used to evaluate the Early Triassic benthic bedded upper unit of muddy limestone. These PACs do
communities by Twitchett et al. (2004) and Wheeley and not show significant variations throughout the section
Twitchett (2005). and indicate an aggradational stacking pattern, typifying
the early stage of the Highstand System Tract (HST)
3.2. Sedimentology (Tong and Yin, 1998). PACs 62–91 are characterized by
increasing thickness of the upper unit of limestone and
A detailed log of lithology, sedimentary structures suggest a progradational stacking pattern, characteristic
and taphonomy of shell concentrations was measured of the late stage of the HST (Tong and Yin, 1998).
from the uppermost Changhsing Formation to the
Helongshan Formation (Fig. 2). Twenty-five samples 3.3. Geochemistry
were thin sectioned to determine carbonate microfacies.
Samples from the hummocky cross-stratified (HCS) bed Proxies for biomass such as phosphorous and calcium
were also thin sectioned (Fig. 6). Small-scale cycles are contents (e.g. Kaiho et al., 1999) are used to assess
recognized throughout the Lower Triassic succession at variations in oceanic productivity throughout the
Meishan. Generally, black shale, claystone, greenish Griesbachian. The Lower Triassic strata were trenched
gray mudstone, and calcareous mudstone characterizes to obtain fresh samples. These samples were crushed to a
the lower parts of cycles whereas greenish gray fine powder after drying at 40 to 50°C and split into
mudstone, muddy limestone/marlstone, and limestone several small samples for analysis of phosphorous,
for the upper parts. These cycles are comparable with calcium and other major elements by ion coupled plasma

Table 1
Lithology of seven high frequency cycles and their distributions in the Lower Triassic of Meishan
Upper unit Green mudstone Muddy limestone + Calcareous Silt-stone Muddy Limestone Limestone
calcareous mudstone mudstone limestone (micrite)
Lower unit Black shale/ Greenish mudstone Black shale Black shale Black shale/ Calcareous Calcareous
claystone and shale claystone mudstone mudstone
and shale
Occurrence 3, 7–9, 11–35, 36, 42, 49, 68, 70 10 4–6, 66, 69 71 1–2, 63–65,
37–41, 43–48, 51–52–53, 67, 72–91
50, 54, 59, 61 55–58, 60, 62
Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187 181

mass spectrometry (ICP-MS). Following the ICP not been appreciably reworked or transported. These
method, 100 mg of powdered sample were decomposed brachiopods and the associated bivalves, ammonoids,
using 5 mL of ultrapure hydrofluoric acid (HF), 3 mL of gastropods and fishes are named the Meishanorhynchia
ultrapure HNO3 and ultrapure hydrochloric acid (HCl) association, which comprises at least nine species in seven
two times. The solution was diluted with 5 mL of genera of five fossil groups. Its Shannon diversity index is
concentrated ultrapure nitric acid (HNO3) and double- 1.25, Simpson diversity index is 0.55, and mean dominance
distilled MILLI-Q-water to adjust the total volume to is 0.45.
exactly 100 mL. Precision estimated from replicate In this fauna, Meishanorhynchia clearly dominates
analyses is within 3%. Solutions prepared from pure (Fig. 4). Both ammonoids and bivalves are moderately
chemicals and JB-2, JB-3, JA-1, JA-2, JG-1, and JG-2 abundant and diverse, including four bivalve species and
from Geological Survey of Japan were used as standards. three ammonoid species, all of which are characteristic
The reproducibility of these measurements was better of the Lower Triassic globally (McRoberts, 2001;
than 2%. McGowan, 2004). Apart from an apparent increase in
diversity, the taphonomy of ammonoids and bivalves of
4. Results and discussion the Meishanorhynchia association is very different from
those preserved in the lower Yinkeng Formation. Shells
4.1. Earliest Triassic faunas: onset of post-extinction are randomly distributed on slab surfaces, and no shell
recovery concentrations occur at the upper Yinkeng Formation. In
addition, the Meishanorhynchia association faunas have
4.1.1. Community ecologic features throughout the no significant size reduction compared to shells from the
Griesbachian lower Yinkeng Formation.
The earliest Griesbachian (parvus zone) faunas from Another feature of the Meishanorhynchia association
the upper part of Bed 27 are dominated by brachiopod is the emergence of benthic gastropods and fishes. The
Paryphella association (Chen et al., 2006). This fauna is undescribed gastropods are morphologically close to the
moderately diverse and comprises 16 species in 10 genera Early Triassic Bellerophon and the fish scales are compa-
(Chen et al., 2005a, 2006), all of which, however, are rable with the late Early Triassic fishes described from
Permian holdovers and therefore survivors of the end- Chaohu, South China (Tong et al., 2006). In addition,
Permian mass extinction (Chen et al., 2005a,b). Succeed- Wignall and Hallam (1993, fig. 13) stated that both
ing this survival association, both bivalve Claraia and Unionites and Schizodus occur in the upper Yinkeng
ammonoid Ophiceras are characteristic of the lower Formation, equivalent to Beds 53–55 at the same section.
Yinkeng Formation of the lower staeschei–isarcica zone
(Beds 28–30), with 1–2 very rare, small surviving 4.1.2. Comparisons with other Griesbachian recovery
brachiopod taxa (Orbicoelia and Acosarina). Diversity faunas and Middle Triassic recovery faunas
of this fauna is low and dominance is high. No trace fossils As stated above, the earliest Griesbachian Claraia–
and bioturbation structures are detected. Ophiceras fauna of Meishan has low diversity, high
The Claraia–Ophiceras assemblage (Yin et al., 2001) dominance, and lacks trace fossils and bioturbation
also dominates the middle Griesbachian (Beds 31–50, structures. This fauna resembles many coeval post-
upper staeschei–isarcica zone), and the fauna is very low
in diversity although its abundance is locally high. In
particular, Claraia shell concentrations (Fig. 5) are very
conspicuous on outcrop. The earliest major diversification
of the Early Triassic fauna occurs in the upper Yinkeng
Formation (Beds 53–55, lower tulongensis–carinata
zone) and is recognized by the conspicuous Meishanor-
hynchia shell bed (Fig. 3). All Meishanorhynchia shells
are articulated and not abraded; all valves are randomly
orientated with no evidence of sorting. They are very
small and have very thin shells with the radial fibre
textures (Fig. 3). Thus most of the shells are preserved in
situ. Although some Meishanorhynchia shells are pre-
served in the strata near the HCS bed, the taphonomy (all Fig. 5. Claraia concentrations (arrow indicated) from the lower
articulated and randomly oriented), suggests they have Yinkeng Formation (Bed 40). Scale bar is 1 cm long.
182 Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187

extinction assemblages elsewhere in the world, and Section 4.2.3, Fig. 7). An increase in Ca contents in the
overall satisfies the first recovery stage proposed by late Griesbachian oceans is also strengthened by an
Twitchett et al. (2004). increase in carbonate throughout the Griesbachian
The Meishanorhynchia association is composed of stratigraphic records. In South China the lithologic switch
elements of Triassic age. The fauna is moderately from the Permian carbonate successions to the mudstone/
diverse, with a much higher diversity than the underlying shale dominated sequence of Early Triassic age is ob-
Claraia–Ophiceras assemblage. Its mean dominance served in most of the shallow marine P–Tr boundary
(D = 0.45) is lower than the coeval faunas from the sections (e.g. Meishan section) except for the great bank
Dinwoody Formation of the western US (D = 0.53, see of Guizhou (Lehrmann et al., 2003). At Meishan an
also Twitchett et al., 2004), but is higher than Oman important feature of the Griesbachian strata (Beds 28 to
(D = 0.24; Twitchett et al., 2004). In addition, although 79) is that the light gray marlstone and muddy limestone
no trace fossils are observed, small bioturbation increase and thicken up-section, whereas the greenish
structures are visible in Bed 52, indicating a significant gray mudstone and black shale thin (Fig. 2). Bed thickness
increase of tiring in comparison with the earliest Triassic of the marlstone, muddy limestone and limestone changes
community. Thus, the overall paleoecologic features of from typically ∼ 1 cm thick (Beds 29–40) to 1–5 cm thick
the Meishanorhynchia association (increasing diversity (Beds 41–52). Above this level of the number of
and evenness, decreasing dominance, deepening tiring, limestone beds significantly increases and beds thicken
but lacking trace fossils and an increase in body fossil up to 7–15 cm (Beds 53–79). These lithologic variations
sizes) indicate an initial recovery between the recovery record an increase in calcium content throughout the
stage 1 and 2 defined by Twitchett et al. (2004) for the Griesbachian (Fig. 2), indicating an increase in calcium
shallow marine ecosystem of benthos. production in the late Griesbachian oceans.
It should be noted that biotic final recovery (term of Recent studies have revealed that both increasing
Erwin, 1998, 2000) following the end-Permian extinc- ocean CO2 concentration and decreased CaCO3 satura-
tion is generally believed to occur in the early Middle tion of seawater can result in a biocalcification crisis for
Triassic when benthic communities are usually charac- organisms that produce CaCO3 skeletons (Feely et al.,
terized by (1) an increase in biodiversity, (2) reappear- 2004; Sabine et al., 2004). Fraiser and Bottjer (2005a)
ance of Lazarus taxa, (3) reestablishment of reef considered that a biocalcification crisis was ultimately
ecosystems, (4) an increase in body and trace fossil triggered by successive eruption of the Siberian flood
size, (5) radiation of calcareous organisms such as basalts, and together with the associated hypercapnia
dasyclads and scleractinian corals, (6) an increase in (Knoll et al., 1996), global warming (Kidder and
tiering depth/height (Erwin, 1998, 2001; Erwin et al., Worsley, 2004; Kiehl and Shields, 2005) and marine
2002). These ecologic features indicate a return to anoxia (Hallam, 1991; Wignall and Twitchett, 1996;
normal communities, which characterize the final stage Isozaki, 1997), was a major factor in the end-Permian
of benthic recovery (Twitchett, 2006). As a result, the mass extinction and subsequent prolonged recovery.
late Griesbachian Meishanorhynchia association is Indeed, such a biocalcification crisis hypothesis is
interpreted herein to signal the onset of biotic recovery plausible and can explain satisfactorily extinction and
following the end-Permian mass extinction in Meishan. recovery patterns of several skeletonized fossil groups.
Chen et al. (2002) estimated 1 myr for the sedimentation Articulate brachiopods are sessile, shelly elements that
of Beds 29–Bed 55 using the radiometric ages of dominated the Permian benthic communities. Most of
Bowring et al. (1998). Combining these ages with those large, thick-shelled groups became extinct during the
of Mundil et al. (2001, 2004), the Meishanorhynchia end-Permian event except for few lucky survivors with
association is interpreted to have formed within 1–2 myr reduced size and very thin shells (Chen et al., 2005a,b).
after the end-Permian event. This recovery time is Thus, the Early Triassic was bad timing for brachiopod
therefore slightly later than the recovery in Oman faunas (Ager, 1965). This is probably partly because the
(middle-late Griesbachian; Twitchett et al., 2004). Early Triassic oceans could not provide sufficient
calcium satisfying the need of brachiopod shell growth.
4.2. Environmental amelioration and productivity In addition, it should also be noted that increased
recuperation of Early Triassic oceans atmosphere/ocean CO2 in the Early Triassic have also
affected brachiopod respiration. Although rates of
4.2.1. Calcium content variation oxygen consumption in brachiopods are generally very
Measurements of bulk samples show that calcium (Ca) low, slightly reduced ocean oxygen may affect strongly
content increases significantly up-section (see also brachiopod respiration, and thus result in demise of
Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187 183

brachiopods (James et al., 1992). According to Peck Claraia and Ophiceras indicate a low energy marine
et al. (1997), temperature change is one of the primary environment. The muddy limestones and limestones of
factors affecting rates of oxygen consumption of the upper Yinkeng and lower Helongshan Formations
individual brachiopods. An increase of 10° in ocean include rare bioturbation and HCS (Figs. 2 and 6) which
temperature will result in brachiopod oxygen consump- together with multidirectional tool marks suggest an
tion rising nearly 10 times. However, the increased offshore setting between fairweather and storm wave
ocean CO2 caused that ocean oxygen has significantly bases. Thus the Yinkeng and Helongshan Formations
reduced during the Early Triassic (Kidder and Worsley, reflect a broad shallowing-upward trend. This general
2004). Thus, the reduced oxygen and increased seawater shallowing may have provided more hospitable condi-
temperature induced by global warming (Kidder and tions for the post-extinction benthic faunas.
Worsley, 2004) could have strongly hindered brachio- In addition, oceanic anoxia was widespread in the
pod respiration during the P/Tr transition. Paleo-Tethys region during the P–Tr transition (Wignall
Rebounds of the Mesozoic brachiopods are also and Twitchett, 2002). Wignall and Hallam (1993) have
associated with a significant increase in calcium content recognized this anoxic event from the Meishan section.
(Fig. 2). At Meishan, the calcium content of Beds 53–55 These authors considered that the finely laminated black
is still very small compared to the pre-extinction shales and greenish gay mudstone of the lower Yinkeng
Changhsingian limestone (Fig. 7), but has clearly Formation indicates anoxic facies, while the marlstone/
increased compared to the earliest Griesbachian mud- or limestone of the upper Yinkeng Formation indicates
stone-dominated strata. The increasing available calci- dysoxic facies. If this scenario is followed, in Meishan
um in the late Griesbachian oceans is essential for Beds 15–52, dominated by finely laminated black shale and
growth of calcareous skeletons of shelly organisms, and greenish gray mudstone (Fig. 2), indicate a depositional
thus could be hospitable for the post-extinction environment with anoxic or poorly oxygenated seawater;
rebounds of shelly faunas. while Beds 53–71 were deposited at a dysoxic facies
setting. Based on Bowring et al. (1998) radiometric ages
4.2.2. Sedimentological features from Beds 37, 48 (Fig. 2), this anoxia is prolonged and last
The black shales and finely laminated greenish gray at least ∼ 400 kyr within the early-middle Griesbachian.
mudstones with low diversity epifauna dominated by The lithologic switch from the shale/mudstone dominated

Fig. 6. Hummocky cross-stratification in Beds 54–55 of the upper Yinkeng Formation. Scale bars are 3 mm long, and pen is 11 cm long.
184 Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187

sedimentary packages to the carbonate dominated succes- Griesbachian in South China (Cao et al., 2002). The
sions at the upper Yinkeng Formation indicates the ebbing onset of biotic recovery in the late Griesbachian as
of anoxia at the late Griesbachian in Meishan. The return of indicated by the Meishanorhynchia association coin-
relatively oxygenated seawater to the Meishan area in late cides with this interpreted increase in oceanic produc-
Griesbachian is also indicated by the presence of both tivity and is consistent with amelioration of ocean floor
bioturbation structures and the HCS beds at Beds 53–55 and ocean water conditions in the Early Triassic.
(Fig. 6). The late Griesbachian oxygenated sea floor was
probably hospitable for benthos' rebounds in the Early 4.3. Was biotic post-extinction recovery controlled by
Triassic. the paleoceanic environmental conditions?

4.2.3. Geochemical signatures In Meishan some Permian brachiopods survived the

A marked decrease in phosphorus and calcium is end-Permian event (Chen et al., 2005a,b, 2006),
recorded across the end-Permian mass extinction however, they became extinct because of the ongoing
horizon (base of Bed 25) at Meishan (Fig. 7). Above deleterious conditions including deep and anoxic
this level values slowly increase although they do not seawater, undersaturation of seawater with respect to
return to pre-extinction levels. The element reductions calcium carbonate, and low oceanic productivity in the
are consistent with negative shifts in δ13C, δ13C kerogen early Griesbachian oceans. Instead, opportunistic disas-
and pristan/phytane ratios (Fig. 7). These trends are ter taxa such as Claraia and Ophiceras dominated the
consistent with an abrupt mass mortality leading to a early-middle Griesbachian oceans. In late Griesbachian
significant reduction in the flux of organic carbon to the local development of shallow water, oxygenated sea-
seafloor at the end of the Permian (Li, 1999; Cao et al., floor conditions and increasing oceanic productivity
2002). Elsewhere the recognized shifts in δ13Ckerogen promoted growth of sessile benthos. Unfortunately, the
and Pristane/Phytane ratios have been linked to brachiopod-dominated association disappeared in the
widespread photic zone euxinia in the P–Tr transition end-Griesbachian worldwide (Chen et al., 2005b).
(e.g. Grice et al., 2005). Recovery of δ13C values and Curiously, their failure in succeeding repopulation
increases in phosphorus and calcium contents through- coincides with the possible environmental devastation at
out the middle-late Griesbachian suggest that oceanic the end-Griesbachian, which is indicated by the Griesba-
primary productivity increased slowly throughout most chian positive shift followed by an abrupt negative shift of
stages of the Griesbachian. The positive shifting cycle of δ13C values at the end of the Griesbachian (Tong et al.,
δ13C excursions has also been recognized from the 2003; Payne et al., 2004).
middle-late Griesbachian successions from various The recovery pattern described here from the Meishan
depositional settings of South China (i.e., Tong et al., section is consistent with strong environmental control
2003; Payne et al., 2004) and interpreted as slowly (Twitchett et al., 2004; Payne et al., 2004; Woods, 2005).
increasing oceanic productivity during the middle-late Global paleoceanic environmental amelioration during

Fig. 7. Bulk contents of calcite and P and variations of carbon isotope values throughout the P–Tr transition from Meishan.
Z.Q. Chen et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 252 (2007) 176–187 185

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Erwin, D.H., 2001. Lessons from the past: biotic recoveries from mass
involvement in the earlier field trip of this project. We are
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