全 文 ：Received 10 May 2004 Accepted 29 Sept. 2004
Supported by the National Natural Science Foundation of China (30340034).
http://www.chineseplantscience.com
Acta Botanica Sinica
植 物 学 报 2004, 46 (11): 1276-1280
Leaf-like Rhodophycean Fossils in the Early Cambrian from Haikou,
Kunming, Yunnan of China
XU Zhao-Liang
(Laboratory of Systematic and Evolutionary Botany, Institute of Botany, The Chinese
Academy of Sciences, Beijing 100093, China)
Abstract: This paper deals with the new leaf-like rhodophycean fossils in the Early Cambrian from
Chengjiang Biota at Mafang Village of Haikou, Kunming, Yunnan, Southeast China. The new taxa Paradelesseria
sanguinea gen. et sp. nov. is found to have a close relationship with the living Delesseria according to
detailed morphological study between the fossil and modern Delesseria. These new findings show that the
Chengjiang Biota is high in species diversity of fossil algae and might offer new evidence for a better
understanding of the Cambrian explosive biological evolution and its paleoenvironment. The Biota in Haikou
district of Kunming was living in subtidal and lower intertidal marine environment, up to 30 m below surface
of the water based on the comparative study of extant red algae.
Key word: Haikou, Yunnan; Early Cambrian; Chengjiang Biota; leaf-like rhodophycean fossils
Macroscopic carbonaceous film fossils emerged in the
geological history and they are generally preserved in the
shales (Hofman, 1992). The morphology of all fossils
changes from ellipsoids, rod-shaped, ached filamentous,
ribbon-like, tubular to forked and other states. Almost no
one likes leaf-like identifying megascopic algae. However,
in the Proterozoic, these have been distinguished as blade-
like fossils, such as Longfengshania from the Qingbaikou
Group, North China and the Little Dal Group, Northwest
Canada (Du and Tian, 1985; Hofman, 1985). About a de-
cade later, the morphological similar fossils, Palmalga
glumacea and Paraporphyra prota have been described
from Doushantuo Formation (Late Sinian), Miaohe, Zigui,
Hubei of China (Ding et al., 1996). Additionally, two years
ago, the blade-like Plantulaformis sinensis and
Longfengshania cordata were also described from the Early
Cambrian in the Chengjiang Biota at Ercai Village, Haikou,
Kunming, Yunnan, Southwest China (Xu, 2002). These find-
ings are very important for understanding the origin and
evolution of metaphytes and studying their paleoecology.
This paper reports a kind of leaf-like rhodophycean
fossil, Paradelesseria sanguinea in the Yuanshan Member,
Qiongzhusi Formation, Early Cambrian at Mafang Village,
Haikou, Kunming, Yunnan, China.
1 General Geology
The early Cambrian fossil samples studied here were
found in the gray muddy shale in the middle of Yuanshan
Member of Qiongzhusi Formation, Anshan, behind of the
Fengyu Temple, Mafang Village, about 3.5 km southeast
of Haikou Railway Station, Kunming, Yunnan of China
(Fig. 1). The general geology has been briefly described by
Xu(2001a; 2001b).
2 Systematic Paleontology
Rhodophyta
Florideae
Ceramiales
Delesseriaceae
Paradelesseria Xu, gen. nov.
Type species Paradelesseria sanguinea Xu, gen. et sp.
nov.
Diagnosis Thallus brown, composed of “stem” and “leaf”;
“stem” subcylindrical, nodose, the length of internode
greater than the diameter of “stem”; “leaf” single, inserted
in the “stem”, consisting of petiole and blade; petiole
subcylindrical, blade lanceolate or oblanceolate, no veins,
margins smooth.
Discussion There have been no previous reports of algal
fossil morphologically similar to Paradelesseria which pos-
sesses stem and leaf, but fossils with the same structure
have been reported, such as: Punctariopsis from
Qiongzhusi Formation, Ercai Village, Haikou, Kunming of
Yunnan, China (Xu, 2001) and Miaohenella from
Miaohecum, Zigui, Hubei in the Sinian (Late Precambrian)
(Ding et al., 1992). The main features of these fossils are as
follow: single or clumped, consisting of unbranched leaf-
like blade, stipe and holdfast. Paradelesseria differs from
XU Zhao-Liang: Leaf-like Rhodophycean Fossils in the Early Cambrian from Haikou, Kunming, Yunnan of China 1277
them in that mentioned above. Its “leaf ” is larger in size
and very thin in thickness, similar to those of some an-
giosperm in structure, and is inserted directly onto the stem.
Those of Punctariopsis assigned to brown algae are very
long, belt-like, inserted on the holdfast, and those of
Miaohenella are hollow, flatted and also inserted on the
holdfast.
Obviously, there are many analogs of Paradelesseria
among the living higher algae, especially in huge brown
algae and red algae which are complex in structure and
morphology and possess structural differentiation of “root,
stem and leaf” superficially like those in vascular plants.
For example, phaeophycean Sargassum, Laminaria,
rhodophycean Delesseria and Phycodrys are very similar
to the new genus described here in structure, morphology
and size, but the leaves of Delesseria and Phycodrys clearly
possess a mid-vein and that of Paradelesseria does not.
Although Paradelesseria might have close relationships
with Delesseria or Phycodrys, there exist some differences
between them in morphology and structure, so it seems
appropriate to establish the fossil as a new genus.
Paradelesseria sanguinea Xu, gen. et sp. nov.
Figs. 2-5
Diagnosis Stem subcylindrical, 20-30 mm in length, 3.0-
5.0 mm in diameter, nodose; node 8.0 mm in length, 4.0-5.0
mm in diameter; internode 3.0 mm in diameter, 3.5 mm in
length; leaf lanceolate with petiole, tampering to acute or
obtuse shape, 60.0-90.0 mm in length, 9.0-12.0 mm broad;
blade thin, 48.0-65.0 mm in length; petiole subcylindrical,
12.0-25.0 mm in length, 3.0 mm in diameter. Reproductive
structure unknown.
Discussion Samples of P. sanguinea are rarely found in
Chengjiang Biota, only three pieces samples and some frag-
ments are well preserved. Until now, the structure and mor-
phology of P. sanguinea were not reported in the geologi-
cal history, but its characteristics are very similar to those
of extant rhodophycean Delesseria sanguinea and
Phycodrys radicusa.
3 Results and Discussion
The rhodophycean is a large and complex organism in
structure other than some single-celled ones in Bangiales.
Its morphological features can be described as follow: single
and forked filaments, flattened and thallus with blades, pin-
nate or branched, wide or narrow belt forms, hollow
cylindroids and thallus with differentiation of roots, stems
Fig.1. Geographic map of fossil sites. a, village; b, railway station; c, city; d, mountains; e, temple; f, fossiliferous site; g, highway; h,
river; i, railway.
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041278
and leaves which look like those of extant vascular plants.
If the concept of the root, stem and leaf in vascular plants is
extended, the structure, which is flattened and enlarged in
surface area for photosynthesis, could be called leaves.
Thus, all of the higher algae, including chlorophycean,
phaeophycean and rhodophycean have developed to the
level of possessing stems and leaves, unlike the lower algae.
Examples of this form are very abundant in the Phaeophyta,
such as Sargassum, Macrocystis and Postelsia. The typi-
cal example in the Rhodophyta is Delesseria (Smith, 1955;
Zhang and Liang, 1965; Fott, 1971) .
The tissue of giant rhodophyceans is relatively
differentiated. Its chromoplast is mainly mosaiced in the
surface layer of photosynthetic tissue, but no
chromatoplasts occur in storage and conducting tissues.
The huge red algae possess sieve plate-like structures in
the conducting tissue which could be compared with the
phloem of vascular plants in general. However, the xylem-
like structure is not present in the red algae as they perma-
nently live in water. So it is unnecessary to have the trans-
fusion vessels to transfer water for a long distance and the
mechanical tissue to support the erect thallus (Zhang and
Liang, 1965; Fott, 1971) .
Different growth methods exist in red algae. Terminal
growth occurs in the most of rhodophyceans. Intercalary
growth occurs only in Delesseriaceae and Corallinaceae,
and the diffuse growth takes place in a few red algae such
as Porphyra. Concerning their reproduction, several uni-
cellular species assigned to Bangiophycidae have vegeta-
tive propagation. Asexual reproduction mainly produces
haploid tetraspores which sometimes change into abnor-
mal polyspores and monospores. All of these spores are
aflagellar aplanospores. Generally, asexual reproduction
happens relatively rare in the red algae by fragmentation of
thallus and also by attachment of fragments that float away
and develop into new plants. Sexual reproduction is
oogamous, but the process is very complex, having a spe-
cial phenotype in the phytological biology. It is evident
that the red algae have reached a higher plateau in evolu-
tion (Smith, 1955; Zheng and Wang, 1961).
Red algae are a relatively large group, with more than
550 genera and 3 700 species in the Rhodophyta. They are
mostly marine, though some genera and species live in the
fresh water (Smith, 1955; Fott, 1971). They are widely
spreaded in the world, and mainly distributed in the ocean
in the tropics and subtropics. All of them are red in color, as
the benthos emerge near the seashore. The depth of the
water they live in is based on the transparency of the ma-
rine water. In the relatively turbid water, such as the Atlantic,
the rhodophyceans always live up to 30 m below the sur-
face of water. In contrast, in relatively limpid waters, such
as those of the Mediterranean Sea and the Floridian
seashore, red algae can live down to 75-90 m, and some-
times have been found growing as deep as 200 m below the
water surface. There is an obvious distinct vertical zona-
tion where all of the red algae grow only near the seashore.
In general, the members of Delesseria and others grow be-
tween subtidal and lower intertidal zones (Zheng and Wang,
1961; Fott, 1971).
Doubtless water temperature is an important ecological
factor for the geographical distribution of red algae. Most
of the species are sensitive to different water temperatures,
and they can not tolerate temperature changes of more than
5 ℃ range each year. Some others, in contrast, grow well
over a 10 ℃ range of marine water temperature, but only a
few species can live in places where the temperature rang is
greater (Zheng and Wang, 1961).
In conclusion, the new discovery of rhodophycean fos-
sils from the Yuanshan Member of Qiongzhusi Formation
in the Early Cambrian, Anshan of Mafang Village, Haikou
of Kunming, China, shows the high diversity of
Chengjiang Biota, and might offer new evidence for a
Fig.2. Reconstruction sketch of the Paradelesseria sanguinea
XU Zhao-Liang: Leaf-like Rhodophycean Fossils in the Early Cambrian from Haikou, Kunming, Yunnan of China 1279
Figs.3-5. Paradelesseria sanguinea gen. et sp. nov. 3. Holotype, No. 0150, × 2.5. 4. Paratype, No. 0149, × 2.5. 5. Counterpart of
Fig. 4, No. 0148, ×2.5.
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041280
better understanding of the Cambrian explotion of bio-
logical evolution. Comparative studies of the fossils and
extant algae, especially the similarity in their structure and
morphology, suggest that there might be a close relation-
ship between fossil genus Paradelesseria and living
Delesseria. Based on the comparative study of extant red
algae, the present results provide further evidence that the
Chengjiang Biota in Haikou district of Kunming was living
in a subtidal and lower intertidal marine environment within
30 m of the surface of the water.
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