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Filamentous cyanobacteria fossils and their significance in the Permian-

Triassic boundary section at Laolongdong, Chongqing

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 ARTICLES
Chinese Science Bulletin

© 2008 SCIENCE IN CHINA PRESS

Springer

Filamentous cyanobacteria fossils and their signifi-

cance in the Permian-Triassic boundary section at
Laolongdong, Chongqing
JIANG HongXia, WU YaSheng† & CAI ChunFang
Key Laboratory of Mineral Resources, Institute of Geology and Geophysics, Chinese Academy of Sciences, Beijing 100029, China

The microbial communities blooming immediately after the end-Permian mass extinction represent
abnormally extreme environments, and vary in different areas. In this study, filamentous cyanobacterial
biota was found in the strata after the extinction in the famous Permian-Triassic boundary section at
Laolongdong, Chongqing, southwest China. In thin sections, the filamentous cyanobacterial fossils are
below 1 mm in length, and generally taper to one end, with the widest diameter up to 0.08 mm. Some of
them are curved, indicating that they are soft in life. Their walls are composed of cryptocrystalline to
microcrystalline calcites. The filaments have round cross section, and are internally filled with micrites
and fine sparry calcites, which indicate that the filaments are originally empty. They are randomly dis-
tributed in the rocks, but in some places, they tend to be distributed in radial pattern. The filamentous
organisms are morphologically similar to Rivularia of Rivulariaceae, Cyanobacteria Phylum, but with
calcified sheaths, and are tentatively regarded as an indeterminate new species in Rivularia: Rivularia
sp. Cyanobacteria are photosynthetic autotrophic, and can survive in dysoxic condition. The blooming
of this organism and the absence of other organisms may indicate that the environment was oxygen-
deficient and shallow, since this photosynthetic autotrophic organism needed to live within photic
zone.

microbial fossil, end-Permian, Permian-Triassic boundary, cyanobacteria, Rivularia

The end-Permian mass extinction event killed about
90% ocean lives including 100% reef lives[1,2]. As we
describe the process of the extinction[3 12], the commu-
―
nities coming after that fatal disaster event should be
also taken into our sight. In the famous Permian-Triassic
boundary (PTB) section of Meishan, the community is
inanimate in which conodonts are the main organism,
with some small brachiopods[13]. In the PTB section of
Ziyun, Guizhou Province, the community after the crisis
is dominated by microgastropods or algae. In the Nan-
panjiang basin[14,15], the main lives of the communities
after the extinction in an adlittoral carbonate platform
include microbial calcified cyanobacteria-Renalcis and
microgastopods. In the southwest of Turkey, the post-
extinction community is microbial and is dominated by
stromatolites[16].
GEOLOGY
In this paper a filamentous cyanobacteria microbial
community is reported for the first time from a PTB sec-
tion at Laolongdong, Chongqing, southwest China,
where it occurs in the carbonate stratum just above the
extinction horizon.
1 Geological setting
Laolongdong is a small village, and is 30 km northwest
of Chongqing (Figure 1). A developed Changhsingian
Received November 12, 2007; accepted January 29, 2008
doi: 10.1007/s11434-008-0172-1
†
Corresponding author (email: HTUjianghx@mail.igcas.ac.cnUTH; HTUwys@mail.
igcas.ac.cnUT; cai_cf@mail.igcas.ac.cn)
Supported by the National Natural Science Foundation of China (Grant No.
40472015), the State Key Laboratory of Geological Processes and Mineral Re-
sources China University of Geosciences, Wuhan (Grant No. GPMR2007-01), the
Science Foundation of China Postdoctors (Grant No. 20070420523)

www.scichina.com | csb.scichina.com | www.springerlink.com Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879

Figure 1 The location and geological map of Laolongdong, Beibei,
Chongqing (modified from Kershaw et al.[21]). J3, Upper Jurassic; J2,
Middle Jurassic; T+J1, Triassic+Lower Jurassic; P, Permian. O-C, Ordovi-
cian-Carboniferous.
reef and a Permian-Triassic boundary (PTB) section
have been found there for many years and made it be-
come the focus of geologists[17 26] throughout the world.
― tous fossils are below 1 mm in length, so we regarded
Fan et al.[19]
introduced the Changhsingian reefs of
Laolongdong to the world in the 1990s. Wignall et al.[20]
systematically studied the PTB section at Laolongdong,
and thought that the sea level kept rising during the
Permian-Triassic transition and made the ocean anoxic,
which led to happening of the mass extinction. Later,
Kershaw et al.[21,22] found microbialite-like rock in the
PTB section at Laolongdong. But they did not determine
the microbialite property of the rock. In May 2006, the
authors examined the PTB section at Laolongdong, and
found a filamentous microbial community never de-
scribed before.
The Changhsingian reefs at Laolongdong did not de-
velop into the end of the Changhsingian stage[18], and
they were replaced by carbonate platform facies in the
1872 JIANG HongXia et al. Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879
latest Changhsingian. In the early Triassic, a caesious
tabular marly lime mudstone was deposited on the car-
bonate rocks. Our section begins from the top of the
Changhsingian reef facies, and ends with the tabular
marly lime mudstone of the basal Lower Triassic
Feixianguan Formation.
Our section is divided into 7 beds (Figure 2). The
lithologic succession and main fossils are as follows in
ascending order.
Bed 1, 5 m thick, is a grayish white massive wack-
estone. The weathered outcrops show karstic faces. Cal-
cisponges (Figure 3(a)) have been found in this bed.
They are the main reef-building lives of Permian reefs,
and have calcified skeletons composed of a string of
chambers.
Bed 2, 10.6 m thick, is grayish white thick-bedded
wackestone and packstone, containing abundant fossils.
The skeletal grains are dominated by Colaniella (Figure
3(b)) and echinoderms (mainly crinoids). The secondary
grains include brachiopods, Palaeofusulina, and Tubi-
phytes.
Bed 3, 0.35 m thick, is a medium to thin-bedded
limestone, and is composed of brownish limestone with
grayish or gray-yellow color, and has a speckled struc-
ture (Figure 3(c)). The grayish and gray-yellow part is
original limestone, and is darker in thin sections. The
brownish part on outcrops is lighter in color in thin sec-
tions. On outcrops, there is more brownish limestone
than grayish and gray-yellow limestone. The grayish and
gray-yellow limestone is a wackestone containing os-
trocods and filamentous microbial fossils. The filamen-
them as microbial fossils.
In thin sections, some lighter areas are composed of
micro-spars recrystallized from lime mudstone and large
block calcite cements filling pores, others have abun-
dant ostracods, which were internally filled by large cal-
cite cements. The darker areas contained abundant fila-
mentous microbial fossils, which are the emphasis of
this paper.
This bed contains cubic pyrites, and it is the lowest
horizon to have cubic pyrites. The cubic pyrites are
about 1 mm in size, and can be seen on outcrops and in
thin sections. They cut into fossils, and occur in both the
darker areas and the lighter areas.
Bed 4, 0.1―0.2 m thick, is a thin-bedded speckled
limestone. In thin section, it consists of darker areas
 ARTICLES
GEOLOGY

Figure 2 Lithologic log and main biotic distribution in the Permian-Triassic boundary section at Laolongdong, Chongqing.

JIANG HongXia et al. Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879 1873
Figure 3 Photographs of outcrops and microphotographs of the Permian-Triassic boundary section at Laolongdong, Chongqing. (a) Bed 1, reef limestone
containing thalamid calcisponge fossils; (b) Bed 2, packstone containing Colaniella; (c) Bed 3, outcrop photograph of speckled limestone with mixed
grayish and brown; (d) Bed 4, speckled limestone, with the brownish spots looking as lighter areas in thin section and containing abundant euomphalid
microgastropods; (e) Bed 5, speckled limestone, containing abundant irregular cystiform microbial fossils which cemented by sparry cements; (f) Bed 7,
dendroid limestone consists of dendroid parts altered from original limestone during diagenesis and the residual lime mudstone between the dendroid
branches.

composed of packstone and wackestone with shells of
euomphalid microgastropods (Figure 3(d)), and lighter
areas consisting of coarser grains recrystallized from the
same limestone as the darker areas and sparry cements
filled in the pores. The euomphalid microgastropods
have shells with laminar fabrics. The largest shells are
up to 1.4 mm wide. Cubic pyrite also occurs in this bed,
and has the same size and distributioin as in Bed 3.
Bed 5, 0.55 m thick, is a medium to thin-bedded,
speckled limestone. It contains irregular cystiform mi-
crobial fossils (Figure 3(e)). The cystiform fossils are
mostly distributed in the lighter areas in thin sections
(that is, the darker areas on outcrops), which are com-
posed of coarser calcites recrystallized from lime mud-
stone and pore-filling large calcites. Besides irregular
cystiform fossils, microgastropods are present in this
bed, but not euomphalid. Cubic pyrite grains also occur,
and they have the same size and distribution as in Bed 3.
Bed 6, 0.3 m thick, is thin-bedded, speckled limestone.
The lithology and fossils are the same as in Bed 5.
Bed 7, 0.85 m thick, is a medium-bedded limestone
with dendroid structure (Figure 3(f)). On outcrops, the

1874 JIANG HongXia et al. Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879

dendroid-structured rock consists of grayish limestone
and brownish limestone, with both of them looking as
roughly dendroid. The brownish areas are lighter in
color in thin sections, because they are composed of mi-
cro-spars recrystallized from lime micrites and large
blocky calcites filling in pores. The lighter areas contain
more fossils than the darker areas, and they are mainly
irregular cystiform microbial fossils, which were inter-
nally filled with large calcites.
Kershaw et al.[22] found the conodont Hindeodus
parvus in a sample from 0.3 m below the top of the den-
droid rock. So, they put the P-T boundary at 0.3 m be-
low the top of the dendroid rock. Before more data on
conodonts are available, the P-T boundary in our section
is tentatively placed at 0.3 m below the top of the den-
droid rock of Bed 6.
The beds above Bed 7 are caesious and tabular marly
lime mudstone of the Feixianguan Formation of the
Lowest Triassic, and are not measured in this study.
2 Filamentous fossils
2.1 Characters of the filamentous fossils
The filamentous fossils are distributed in Bed 3 of
Laolongdong section. On outcrops, this bed has no dif-
ference in color to other beds, showing speckled struc-
ture. The fossils in the bed are very small and did not
form special fabrics, so that they are not easy to be
found.
Examination of thin sections reveals that the fossils
are below 1 mm in length. All filaments taper to one end,
with the widest diameter being 0.08 mm (Figure 4(a)).
Some filaments vary in diameter, and a few has a fold
(Figure 4(b)). Some filaments are curved, probably due
to limitation by the contacted objects (Figure 4(c), (d),
(e), (f)), which indicates that they were soft in life. The
walls of the filaments, generally 0.005―0.01 mm, are
composed of cryptocrystalline to microcrystalline cal-
cites, looking as a dark line in longitudinal sections and
as dark circles in cross sections. The thickness of walls
in a single filament is uniform, though it varies among
different filaments, which may reflect difference in the
growth stage or nutritional level they got. All filamen-
tous fossils are filled with micrites or tiny sparry calcites,
which indicates that they are empty originally.
The filamentous fossils are randomly distributed in
rocks, and their longitudinal and cross sections are ran-
domly distributed in thin sections (Figure 4(c)). But, in
JIANG HongXia et al. Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879
ARTICLES
some places, they show radial or bunching arrangement.
2.2 Systematical position of the filamentous fossils
The size of the filamentous fossils belongs to microbes.
Microbes are not a taxonomic group, but a morphologica
group that includes all tiny organisms that must be ob-
served and studied using microscopes or electronic mi-
croscopes. It generally includes bacteria, archaeota,
fungi, tiny algae, and tiny protozoans.
Protozoans are 0.03―0.3 mm in size, but some of
them exceed 100 mm. Only tiny protozoans are regarded
as microbes. The microbial protozoans include four
classes: Mastigophora, Sarcodina, Sporozoa, and Infu-
soria. Of these groups, only the subclass Foraminifera of
the class Sarcodina may have filamentous fossils. We
excluded the possibility that the filamentous fossils be-
long to small foraminifers, the reasons: (1) the fabrics of
their walls do not belong to any types of foraminifers;
(2) they have no chambers; and (3) they are soft in life,
contrast to hard shells of foraminifers.
As green algae are concerned, only the Order Ulotri-
chales have unbranched filamentous type. But they con-
sist of a row of cylindric cells. Our fossils do not show
such feature. In addition, members of Ulotichales are
mostly fresh water type; only a very small number of
them live in sea water. Furthermore, no calcified
Ulotichales members have been reported.
Fungi are usually branched, and the diameter of sin-
gle mycelium is constant. Our fossils taper to one end,
and are unbranched. So they are different from fungi.
Archaeota members are generally spherical or irregu-
larly polygonal. A few of them are rod-like or string-like,
GEOLOGY
and are several to tens of microns in length, and less
than 1 μm in width. They do not taper to one end. Our
fossils are in 1000 μm in length and 80 μm in widths,
and taper to one end. So they do not belong to archaeota.
Bacteria have unbranched filamentous type. But they
have constant diameter, and are very small, generally
0.001 mm in diameter, and less than 0.05 mm in length.
Our fossils are different from bacteria.
The filamentous cyanobacteria have three types: un-
branched, branched, heterocystal types. The unbranched
filament include three types: (1) the mycelium consists
of a string of spherical, elliptic, or drumlike cells, and
has a unsmooth outer surface, generally catenulate, with
constant diameter; (2) the mycelium is composed of a
string of cylindrical cells, and has a smooth outer sur-
face, with a constant diameter; and (3) the mycelium
1875
Figure 4 (a)―(f) microphotographs of filamentous cyanobacterial fossils; (g) Rivularia PCC 7116, the filaments are composed of a string of cells with
sheaths, taper to an end (modified from Boon and Castenholz, 2001); (h) microphotographs of speckled rocks. The lighter area in thin section is the
brownish spot area of outcrops, and consists of large calcite cements containing small rhombic calcite crystals and residual lime mudstone. The darker area
in thin sections is the grayish spot area of outcrops, and is composed of lime mudstone.

1876 JIANG HongXia et al. Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879

consists of a string of cylindrical or drumlike cells, and
has a smooth outer sheath. Its most distinct character is
that the mycelium tapers to one end (but it has constant
diameter in young stage) and its thick end has a spheri-
cal cell. Our filamentous fossils are similar to type (3).
The members of type (3) are originally assigned to
Rivulariaceae. In the newest classification, they are
placed into Subsection IV.II of cyanobacteria[27]. This
1 nous biota, indicating that their environments may be
subsection contains three form-genus: Calothrix Agardh
1824, Rivularia Agardh 1824 (Figure 4(h)), and Toly-
pothrix Kützing 1843.
All our filamentous fossils are characterized by ta-
pering to one end of the mycelium. Other important fea-
tures observed include: (1) a small spherical cell at the
end of the thicker end of the mycelium; (2) the cells are
cylindrical; (3) the young mycelium are constant in
diameter; (4) some mycelium are curved, indicating that
they were soft in life; and (5) the distribution of the my-
celiums in rock are random. All these characters are
similar to Rivularia (Figure 4(g)). The only difference is
that our fossils have calcified sheaths, but Rivularia dose
not. Considering both the similarities and difference, our
filamentous fossils are placed in Rivularia, and are re-
garded as an indeterminate new species, Rivularia sp.
3 Discussion
Figure 3(e), (f) in the paper by Ezaki et al.[23] are possi-
bly the same organisms as our filamentous cyanobacte-
ria. But they labeled them as foraminifer. Groves and
Altiner[8] reported Earlandia sp. in the lower Triassic
stromatolites in Central Taurides, Turkey. They consid-
ered that the fossils they described are the same as Fig-
ure 3(e), (f) of Ezaki et al.[23]. Angiolini et al.[24] studied
the PTB section in Antalya Nappes, Turkey, and de-
scribed some Earlandia distinctly different from those
fossils figured as foraminifer by Ezaki et al.[23]. The fo-
raminifer Earlandia has an initial chamber, but the fos-
sils described by Ezaki et al. and Groves and Altiner
have no chambered feature. Therefore, we do not agree
to their assignment.
Our filamentous fossils can not be placed in fo-
raminifers for three reasons: (1) Foraminifers mostly
live in normal sea, while our filamentous fossils oc-
curred in the extremely abominable environments
after the end-Permian extinction. Although Some re-
searchers found abundant small foraminifers in P-T
transition[28 30], they are multi-chambered, and are dif-
―
JIANG HongXia et al. Chinese Science Bulletin | June 2008 | vol. 53 | no. 12 | 1871-1879
ARTICLES
ferent from our filamentous fossils. The depositional
environment varies from places to places, and small fo-
raminifers may be present in some places. Some re-
searchers reported that some foraminifers can survive
the end-Permian mass extinction[31]. But, all materials
they figured or described are different from our fossils.
(2) Our filamentous organism lived as a very monoto-
very harsh, and in such condition no other organisms but
cyanobacteria could live. Modern microbe ecology
studies show that the microbes in extremely harsh envi-
ronments are mainly cyanobacteria. (3) Groves and
Altiner[8] reported that their fossils were in stromatolites.
Stromatolites are generally formed by cyanobacteria and
bacteria, not foraminifer. No foraminifer has been re-
ported from modern stromatolites[32].
Hips and Haas[33] reported Earlandia, ostrocods, and
2
bioclasts from the 0.5-m-thick thin-bedded lime mud-
stone on the P-T boundary in Bükk, Hungary. Groves
and Altiner[8] thought that Earlandia are distributed in
Italy, Austria, Slovenia, Iran and Turkey, etc. Wang et
al.[34] described two kinds of microbial fossils from the
lowest Triassic microbialite in Chongqing and Hubei
Province, China. One is chambered cyanobacterial
Renalcis, and the other is micro-spherical cyanobacterial
fossil. Their fossils are different from ours in shape and
size.
4 Paleoecologic significance of filamen-
tous cyanobacterial community and their
contribution to rock formation
GEOLOGY
Cyanobacteria are a group having the widest ecological
requirements, and can adapt to different environments
with wide salinity, temperature, pressure ranges. They
are photosynthetic, and can live in extremely poor-nu-
tritional environments[35]. In eutrophic water, cyanobac-
teria are also the main members of microbes[35]. They
may be the most adapted to anoxic environment because
many kinds of cyanobacteria can emit oxygen through
photosynthesis. Due to photosynthesis, cyanobacteria
need to live in shallow water, within photic zone[35].
3
Previous researchers thought that anoxic event hap-
pened in Laolongdong section according to the existence
of pyrite. The authors also found pyrite on the outcrops
and in thin sections. It is generally larger than 1 mm in
size, and can cut fossils, which indicates that they
formed in diagenetic stage, or formed in secondary
1877
enlargement of tiny depositional pyrite grains. Only very
tiny framboidal (<8 µm) pyrite can testify anoxic condi-
tion. Through the Scanning electron microscope (SEM)
we did not find tiny framboidal pyrite. So it is difficult
to determine anoxia in our Laolongdong section. Cras-
quin-Soleaua and Kershaw[25] studied ostrocods in the
PTB section in Chongqing, and concluded that the envi-
ronments were oxygen-deficient after the end-Permian
extinction. The monotonous filamentous cyanobacterial
community found in this study support their conclusion,
since cyanobacteria are photosynthetic, and can adapt to
dysoxic condition. The occurrence of cyanobacterial
fossils also shows the water is shallow since photosyn-
thesis should be carried within photic zone.
Filamentous cyanobacteria are too small to form ef-
fective pores for diagenetic fluid to migrate in rocks.
The rocks with filamentous cyanobacteria fossils show
speckled structure, which, as Figured 4(h), was formed
not by microbes but by multiple diagenetic processes
including dissolution and cementation[36]. In thin sec-
tions, the speckled-structured rock consists of lighter
and darker areas. But both of them contain no microbial
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5 Conclusion
(1) In the PTB section at Laolongdong, Chongqing,
the community immediately after the end-Permian ex-
tinction is a very monotonous one dominated by fila-
mentous cyanobacteria. These fossils are below 1 mm in
length, taper to one end, with widest diameter being 0.08
mm. These features are very similar to those of Rivu-
laria. So, we assigned them to Rivularia, and regarded
them as an indeterminate new species, Rivularia sp.
(2) The stratum containing filamentous microbial fos-
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