全 文 ：Received 14 Jun. 2003 Accepted 20 Oct. 2003
Supported by the National Natural Science Foundation of China (40061001), the State Key Basic Research and Development Plan of China
(2002CCC02600) and the Guizhou Foundation of the Excellent Youth Scientists and Technicians.
* Author for correspondence. E-mail: .
http://www.chineseplantscience.com
Bryophyte-like Fossil (Parafunaria sinensis) from Early-Middle
Cambrian Kaili Formation in Guizhou Province, China
YANG Rui-Dong, MAO Jia-Ren, ZHANG Wei-Hua, JIANG Li-Jun, GAO Hui
(Guizhou University of Technology, Guiyang 550003, China)
Abstract: Abundant well-preserved animal and macroalgal fossils were discovered in the Early-Middle
Cambrian Kaili Formation of Taijiang County, Guizhou Province, China, which constitute the Kaili Biota,
named as one of the three important Burgess Shale-type faunas. The bryophyte-like fossil (Parafunaria
sinensis Yang gen. et sp. nov) was discovered in the horizon of occurrence of Macroalgal fossils. The fossil
of P. sinensis possesses typical characters of bryophyte, such as whorled leaves, capsule and seta, and
complex rhizoid or foot. We infer that the bryophyte-like fossil could be the ancestor of bryophyte. The
discovery provides a new lead for further study on the origin and evolution of bryophyte or land plant.
Key words: bryophyte-like fossil; Kaili Formation; Middle Cambrian; Guizhou Province
The origin and evolution of land plant is one of the
unsolved evolution events of earth’s life. Of the plant fos-
sils being discovered, cryptospore assemblage from the
lower Middle Ordovician (476 Ma) of Saudi Arabia was
considered as the earliest land plant (Strother et al., 1996).
In 1999, Strother, an American paleontologist reported the
oldest cryptospore assemblage from the Middle Cambrian
of North America (Strother and Wood, 1999). Thereafter,
the earliest vascular plant fossil from the Middle Ordovi-
cian of North Europe was determined (Hao et al., 2002).
Pinnatiramosus qianensis (Geng, 1986) from the
Llandoverian (431 Ma) of Early Silurian of Fenggang County,
Guizhou Province, China was claimed to be the oldest macro
land plant fossils (Cai et al., 1996). Isotopic carbon (d13C)
values from the Neoproterozoic Beck Spring Dolomite and
Mesoroterozoic Mescal Limestone paleokarst show that
the land plant was present 1 200 Ma ago (Kenny and Knauth,
2001), whereas molecular clock estimates suggest that the
early land plant appeared by 700 Ma (Heckman et al., 2001).
Obviously, the oldest record of the land plant fossil in the
geological time is much younger than that of the land plant
in the divergence time estimated in molecular clock, d13C
and molecular paleontology. Therefore, paleontologists
constantly consider that the vascular plant fossils should
exist in the pre-Middle Ordovician strata. The discovery of
the bryophyte-like fossil（Parafunaria sinensis）from
the Middle Cambrian Kaili Formation in Guizhou Province,
China not only provides an important evidence for the
above-mentioned viewpoint, but also a new lead for further
study on the origin and evolution of bryophyte or land
plant.
1 Locality and Age of the Fossil
Because sea-level fell during the interval of the Early-
Middle Cambrian，the slop facies, along the range of
Sanhui-Taijiang-Danzhai of the eastern Guizhou Province,
developed the Kaili Formation composed of terrestrial clas-
tic rock, simultaneously, the shallow facies of the range of
Sanhui-Kaili-Danzhai might form some little islands. A lot
of animals and algae lived around the small islands or on
the upper part of the slop. Based on the ecological behav-
iors of animals and algae, and their characteristics of depo-
sition and the geochemical parameter, it can be considered
that most animals and algae in the Kaili Biota lived in the
shallow water, and these organisms were carried into the
slop or shelf by tempest or turbidity flow and buried in the
slop or shelf. Therefore, the Kaili Biota (Lagerstätte) is only
a window to understand the Middle Cambrian life.
The Balang section is 270 km far from Guiyang City,
Guizhou Province of Southwest China (Fig.1A), which con-
sists of mudstones and calcareous mudstones bearing
abundant intact animal and alga fossils which constitute
the Kaili Biota including 12 phyla, and about 120 genera
(Zhao et al., 1994), in which about 20 genera belonging to
macroalgae (Yang et al., 1999; 2000; 2001a; 2001b). The
fossils of the Kaili Biota were collected from the middle part
of the Xingrenaspis-Oryctocephalus Zone in the middle of
the Kaili Formation of the Lower-Middle Cambrian (Fig.1B)
(Zhao et al., 1994). The bryophyte-like fossils described in
this paper were collected from the Kaili Biota, about 1 m
Acta Botanica Sinica
植 物 学 报 2004, 46 (2): 180-185
YANG Rui-Dong et al.: Bryophyte-like Fossil (Parafunaria sinensis) from Early-Middle Cambrian Kaili Formation 181
over the horizon of the first appearance of Oryctocephalus
indicus (key fossil for the Middle Cambrian). So, the age of
the bryophyte-like fossil（P. sinensis）from the Kaili
Formation in Guizhou Province, China certainly is of the
Middle Cambrian.
2 Fossil Description
The bryophyte-like fossil（Parafunaria sinensis Yang
gen. et sp. nov）(Fig.2A-D) from the Kaili Formation of
Taijiang County, Guizhou Province, China was preserved
within the mudstones and calcareous mudstones depos-
ited in the shelf or the upper part of the slop. The bryo-
phyte-like fossil was preserved in carbonaceous form, which
looks like the carbonaceous alga fossils of the Kaili Biota.
Kingdom Bryophyta ?
Division ? Musci?
Class, Order, Family uncertain
Genus Parafunaria gen. nov
Etymology: The fossil is similar to the living Funaria.
Material: 12 specimens with intact thalli collected by
Yang and Zhao from the Kaili Formation, together with many
dissociated fragments.
Holotype: Intact thallus, GTM-9-1-168; Fig.2B, collected
from the Middle Cambrian Kaili Formation of Taijiang,
Guizhou Province, China. Noncalcareous. The color of the
specimens is pale-brown to dark-gray.
Occurrence: The Middle Cambrian Kaili Formation of
Taijiang, Guizhou Province, China.
Type species: Parafunaria sinensis Yang, sp. nov.
(specimen No. GTM-9-1-168; Fig.2B)
Description： Carbonaceous thallus, 4-5 whorled
leaves or crowd leaves, leaf long, obvious costa, no mid-
costa, short stem, complex rhizoid or foot, and with capsule
and seta. The thallus is 2.0 cm in length, 0.8 cm in width.
Stem 1.5 mm in length. Leaf 5-15 mm in length, and 5 mm in
width.
Discussion：The morphology of the fossil has not
been reported. According to its morphology featuring with
4-5 whorled leaves, and obvious costa, complex rhizoid or
foot, capsule and seta, it seems to be comparable with the
living Funaria hygrometrica Hedw (Wu et al., 1990; Taylor,
1981). Therefore, the authors proposed that Parafunaria
sinensis Yang, gen. et sp. nov. could be a bryophyte fossil.
The oldest bryophyte fossil Pallavicinites (=Hepaticites)
from Late Devonian of New York has been previously re-
ported (Huber, 1961; Schuster, 1966). The bryophyte-like
fossil Sporogonites from the Early Devonian of Norway
(Halle, 1916; Andrews, 1960), and Tortilicaulis from the
Late Devonian of South-West Dyfed have certain affinity
for bryophyte (Edwards, 1979; Taylor, 1981). The fossil of
Fig.1. Regional locality and horizon of the bryophyte-like fossil(Parafunaria sinensis) from the Early-Middle Cambrian Kaili
Formation of Taijiang County, Guizhou Province, China.
Acta Botanica Sinica 植物学报 Vol.46 No.2 2004182
Sporogonites yunnanense Hsu from Devonian of eastern
Yunnan, as the cryptospores of genus Andreaea was found
in 1966 (Xu, 1966). Zhang (1988) suggested that
Longfengshania of Proterozoic Changlongshan Formation
is morphologically similar to Sporogonites; thus, he con-
sidered that Longfengshania has affinity to bryophyte. P.
sinensis is similar to the Devonian Pallavicinites and
Sporogonites in structure, such as whorled leaves, and
obvious costa, complex rhizoid or foot, and the feature of
capsule and seta. But it is different from the Devonian
Pallavicinites and Sporogonites in morphology. Fossil P.
sinensis is small, and possesses 4-5 whorled leaves only
which fossils Pallavicinites and Sporogonites from Devo-
nian are larger, and possesses over 10 whorled leaves
(Andrews, 1960; Schuster, 1966; Xu, 1966). Moreover, fos-
sil P. sinensis possesses obvious costa.
The fossil P. sinensis is similar to the living Bryum in the
presence of whorled leaves, and obvious costa, complex
rhizoid or foot, and the feature of capsule and seta, but the
living Bryum (bryophyte) possesses obvious and long stem,
nevertheless the fossils with short and unobvious stem
(Fig.2G, H) from the Kaili Formation is even more similar to
Fig.2. The bryophyte-like fossil Parafunaria sinensis and cryptospore fossils from the Kaili Formation of the Middle Cambrian,
Guizhou Province, China. A. P. sinensis, with complete rhizoid, veins; specimen No. GTM-9-2-113. B. type species: P. sinensis, with
the feature of capsule and seta, specimen No. GTM-9-1-168. C. P. sinensis, with costa, specimen No. GTM-9-1-136. D. P. sinensis,
specimen No. GTM-9-5-123. E. Nodospora sp., specimen No. P-6, X800. F. Dyadospora sp., specimen No. P-6, X800. G and H.
structural sketch of the specimens No. GTM-9-2-113 and No. GTM-9-1-168. Scale in A, B, C, D, G, H for 1 cm.
YANG Rui-Dong et al.: Bryophyte-like Fossil (Parafunaria sinensis) from Early-Middle Cambrian Kaili Formation 183
the living Funaria hygrometrica Hedw (bryophyte).
In 1997, Bian and Ynag① discovered a typical bryo-
phyte fossil Tarimia leei from the Late Ordovician in Tarimia
Basin, Xinjiang, China. This specimen is 2.8 cm in length,
with about 60 leaves, whorled or spiral, the feature of cap-
sule and seta, and no mid-costa. The fossil is possibly the
oldest bryophyte fossil reported previously, and it is simi-
lar to the living Bryum (bryophyte).
3 Significance of the Bryophyte-like Fossil in
the Evolution of the Vascular Plant
The origin and evolution of the vascular plant has been
long-time under debate. Since bryophyte is the oldest land
plant and vascular plant, Cambrian bryophyte fossils be-
come important in the study of the origin and evolution of
land plants. Because bryophyte plant is too small in size for
fossil preservation, and the bryophyte fossils are almost
rare in strata; thus the palaeontologists have difficulty to
define the origin of the bryophyte. Therefore, based on the
morphological and anatomical features of bryophyte, two
views upon the origin of bryophyte have been put forward
(Taylor, 1981; Wu et al., 1990). The first view suggests that
the bryophyte originates from Chlorophyte (Wu et al., 1990),
as both bryophyte and Chlorophyte possess the same chlo-
rophyll and xanthophylls, additionally, in the modern
Chlorophyta, Fritschiella tuberosa and Takakia
lepidozioides have obvious affinities with bryophyte in
morphology and structure (Wu et al., 1990). The second
view considers that the bryophyte originates from
Psilophyton , based on genus Rhynia and genus
Psilophyton of Pteridophyta having obvious affinities with
bryophyte in morphology and structure, and the age
(Silurian) of pteridophyte fossils previously found being
older than that (Devonian) of bryophyte fossils (Taylor,
1981). Nevertheless, the bryophyte fossils discovered are
too rare to satisfy the needs of research.
The bryophyte-like fossil（P. sinensis）discovered
in the Middle Cambrian strata, infer that bryophyte might
have emerged in Cambrian. Since Psilophyton
(Pteridophyta) fossils discovered previously are from the
post-Cambrian, on the basis of geological time of the fossil
record, the oldest Psilophyton (Pteridophyta) fossil dis-
covered previously is in Silurian, and bryophyte fossils
discovered previously are in Ordovician, while the present
bryophyte-like fossil discovered is in the Middle Cambrian,
it infers that bryophyte did not originate from Psilophyton
(Pteridophyta). In contrast, because Psilophyton
(Pteridophyta) possesses obvious and robust stem, as com-
pared with the unobvious and short stem of the bryophyte,
thus, Psilophyton (Pteridophyta) possibly originates from
bryophyte. However this view merits further fossil study
for confirmation.
Gray, Burgess, Strother and Wellman respectively sug-
gested that the plant existed in the Late Ordovician to Early
Silurian based on the findings of cryptospore fossils in the
Late Ordovician and Early Silurian (Gray and Boucot, 1977;
Strother and Traverse, 1979; Burgess and Edwards, 1991;
Wellman, 1995). Volkheimer et al.(1980) reported the bryo-
phyte-like cryptospores from the Mojotoro Formation of
the Early Ordovician and Argenigian of Prague basin
(Vavrdova, 1990). Strother et al. (1996) reported a new evi-
dence for land plants from the lower Middle Ordovician of
Saudi Arabia in 1996. The evidence of fossil P. sinensis
from the Middle Cambrian Kaili Formation of Taijiang and
the cryptospore fossils（Fig.2E, F）from the Middle
Cambrian Kaili Formation of Danzhai County, Guizhou
Province, China indicate that bryophyte was probably
present in the Middle Cambrian.
The cryptospore fossils（Fig.2E, F）from the Middle
Cambrian Kaili Formation of Danzhai County, Guizhou
Province, China consist of imperfectly spores, tetrads and
dyads similar to the plant microfossil Lophodiacrodium
（Strother et al.,1996, pl.1,figs.9,12; pl.2, fig.1）from
Llandoverian and Wenlockian rocks of Pennsylvania
(Strother and Traverse, 1979). Nevertheless, the cryptospore
fossils are still a problem merits further study. The discov-
ery of the Middle Cambrian bryophyte-like fossil and the
cryptospore fossils indicate that bryophyte was probably
present in the Middle Cambrian (520 Ma), and provides a
new lead for further study on the origin and evolution of
bryophyte or land plant.
4 Conclusion
The oldest bryophyte fossils from Devonian (Xu, 1966)
or Late Ordovician (Bian and Yang, 1997) has been previ-
ously reported, but the oldest cryptospore fossil discov-
ered may be traced back to Cambrian (Strother and Wood,
1999). The Middle Cambrian bryophyte-like fossil and the
cryptospore fossils from Guizhou Province, South China
were discovered, indicating that bryophyte was probably
present in the Middle Cambrian (520 Ma), therefore, the
bryophyte-like fossil from Guizhou Province, South China
is the oldest bryophyte fossil ever known. The discovery
infers that the age of origin of bryophyte is 50 Ma earlier
than that being considered before. Based on that
① Bian L-Z(边立曾), Yang R-D(杨瑞东). 1997. The bryophyte
fossils from Late Ordovician in Talimu Basin, Xinjiang. Research
report.
Acta Botanica Sinica 植物学报 Vol.46 No.2 2004184
Pteridophyta and Chlorophyta have affinities with bryo-
phyte in morphology, and on account of the fossil records
of Pteridophyta, Chlorophyta and Bryophyta in the geo-
logical time, we infer that bryophyte originates from
chlorophyte because chlorophyte fossil has been discov-
ered in Precambrian, and pteridophyte originates from bryo-
phyte because the oldest pteridophyte fossil having dis-
covered previously is in Devonian.
Acknowledgements: The authors wish to thank Prof. YIN
Lei-Ming, Prof. ZHAO Yuan-Long and BIAN Li-Zeng for
their help in this study.
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