Review of Palaeobotany and Palynology 119 (2002) 125^141

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Fern ecological implications from the Lower Jurassic in

Western Hubei, China
Yongdong Wang 
Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China

Abstract
The Lower Jurassic Hsiangchi Formation in western Hubei, China is well known for its abundant and diverse
fossil ferns, including Marattiaceae, Osmundaceae, Matoniaceae, Dipteridaceae and Dicksoniaceae. Through recent
collections and investigation of the fossil plants in this area, an autochthonous/hypoautochthonous fern community
has been recognised from the upper part of the Hsiangchi Formation in Zigui, Hubei. This community is dominated
by the matoniaceous ferns of the genus Phlebopteris; Dipteridaceous ferns mainly characterised by Dictyophyllum
rank second in abundance. Marattiaceae (Marattia) and Osmundaceae (Todites and Cladophlebis) are often dispersed
spatially among the dominant ferns, whereas the small and slender herbaceous Dicksoniaceae, characterised by
Coniopteris, are sparsely distributed within the community. This fern community colonised the fluviatile peat-forming
environment and indicates a tropical to subtropical climate with warm and humid conditions, which is favourable for
the peat accumulation. It is also a representative fern community recognised from the Southern Floristic Province in
China during late Early Jurassic period. . 2002 Elsevier Science B.V. All rights reserved.
Keywords: fern; ecology; peat accumulation; Hsiangchi Formation; Lower Jurassic; China

1. Introduction
Abundant living ferns characterise the southwestern region of China today including Yunnan,
Sichuan and Hubei Provinces. During the Mesozoic, oras including ferns were well developed in
the Triassic and Jurassic of this region. In western
Hubei, fossil ferns have been described from Early
Mesozoic strata along the Three Gorges Area of
the Yangzte River. Among them, the remarkable
Early Jurassic ora from the Hsiangchi Formation in the Zigui Basin is well known for its abundance and diversity of fossil plants. According to

* Corresponding author. Fax: +86-25-3357026.

E-mail address: y.d.wang@jlonline.com (Y.D. Wang).

the previous studies and the available data, the

Hsiangchi Flora is composed predominantly of
licopsids and bennettitaleans, as well as a sizable
number of ginkgoaleans (Sze, 1949; Wu et al.,
1980; Meng, 1987). Ferns are one of the dominant elements of this ora, including Marattiaceae, Osmundaceae, Matoniaceae, Dipteridaceae
and Dicksoniaceae. Previous studies of ferns in
this ora were based mainly on the gross morphology of impression specimens. In recent years,
a lot of well preserved compression and impression specimens of ferns were collected from Zigui
Basin in western Hubei, where the type locality of
the Hsiangchi Formation is located. Some of
them, such as Marattia and Phlebopteris were investigated on their fertile organs and in situ
spores using light, scanning and transmission elec-

0034-6667 / 02 / $ ^ see front matter . 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 4 - 6 6 6 7 ( 0 1 ) 0 0 1 3 3 - 6

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Y.D. Wang / Review of Palaeobotany and Palynology 119 (2002) 125^141

tron microscope techniques (Wang, 1999; Wang

and Mei, 1999; Wang et al., 2001).
As an important part of the ora and the indicator plants for the environment, the fossil ferns
were less emphasised for their ecology in western
Hubei. The purpose of the present paper is to
study the fern ecological implications from the
upper part of the Lower Jurassic Hsiangchi Formation in western Hubei, based mainly on new
collections and relevant published data. Through
the analysis of the mode of burial, and the indicators of the sedimentary features, a special ecological community dominated by the Matoniaceae
was recognised in this region, from which the climatic conditions can be inferred.

2. Geological setting, sedimentology and

stratigraphy
The present fossil locality is in Zigui County,
western Hubei, the eastern part of the Three
Gorges Area of the Yangtze River. This area is
tectonically located in the southeastern marginal
region of the ancient Yangtze Plate, along the

north coast of the Tethys Ocean (Wang et al.,

1996). The Jurassic strata are well developed in
western Hubei, attaining a maximum thickness
of 6000 m and are mainly distributed in the Zigui
and Jinmen^Dangyang Basins, which crop out,
respectively, along the western and eastern limbs
of the Huangling Anticline (Fig. 1). In Zigui Basin, the Jurassic is principally distributed in Zigui
and Xingshan Counties on both the northern and
southern banks of the Yangtze River. The Jurassic strata consist of an extensive succession of red
clastic sediments of the Middle to Upper Jurassic.
At the base is a coal-bearing sequence of the Lower Jurassic, which was known as the Hsiangchi
Group or the Hsiangchi Coal Series (Lee, 1924).
According to systematic stratigraphic investigations (Chen et al., 1979; Wu et al., 1980; Meng
and Zhang, 1987), the Hsiangchi Group is composed of two biostratigraphic units: the Upper
Triassic Shazhenxi Formation below and the
Lower Jurassic Hsiangchi Formation above (see
also g. 2 of Wang, 1999). The Hsiangchi Formation is composed of grey to greyish sandstones,
siltstones, mudstones and carbonaceous shales, intercalated by coal seams as well as thin layers of

Fig. 1. A sketched geological map showing locality and the distribution of the Jurassic strata in western Hubei, China (simplied
after Meng and Zhang, 1987). 1, Lower Jurassic; 2, Middle Jurassic; 3, Upper Jurassic; 4, granite; 5, Pre-Sinian to Sinian;
6, fault.

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127

Fig. 2. Lithostratigraphy and sedimentology of the Lower Jurassic Hsiangchi Formation in western Hubei.

siderite, with a total thickness of about 200 m.

The base of this formation consists of a layer of
conglomerate and/or conglomeratic sandstone.
Sedimentological analysis indicates that during
the Jurassic period, the Three Gorges Area of the
Yangtze River was characterised by a long-lived
terrestrial depositional environment, resulting in a
series of stream and lake deposits (Gao et al.,
1999). The Lower Jurassic Hsiangchi Formation
represents meandering stream deposits, which
bear the typical normal sequential succession
(Fig. 2). The basal part is the lag gravel of the
stream bed deposit. Upward, the lithology
changes to point bars deposits of thick sandstone,
ood plain deposits of thick mudstone, or uviatile peat marsh with carbonaceous mudstones and

coal seams. This kind of succession repeats in

several rock packages from the basal to the upper
part of the formation.
Abundant fossils are reported from the Hsiangchi Formation, including plants, spore^pollen,
megaspores and bivalves. According to Wu et
al. (1980) and Meng (1987), the plants from the
Hsiangchi Formation are characterised by a Ptilophyllum^Coniopteris assemblage (Table 1). The
spore and pollen grains of the Hsiangchi Formation are represented by a Dictyophyllidites^Classopollis^Cerebropollenites assemblage in the lower
part, and a Dictyophyllidites^Cyathidites^Classopollis assemblage in the upper part of the formation (Zhang and Zhang, 1987) (Table 1). The
megaspores from the lower part of the formation

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Y.D. Wang / Review of Palaeobotany and Palynology 119 (2002) 125^141

Table 1
Fossil assemblage zones recognized in the Lower Jurassic Hsiangchi Formation in western Hubei
Megaplants
(Wu et al., 1980;
Meng, 1987)

Spores and pollen grains

(Zhang and Zhang, 1987)

Megaspores
(Yang and Sun, 1987)

Bivalvas
(Yu, 1987)

Ptilophyllum^Coniopteris
assemblage

Upper part: Dictyophyllidites^

Cyathidites^Classopollis
assemblage
Lower part: Dictyophyllidites^
Classopollis^Cerebropollenites
assemblage

Upper part: Bacutriletes harrisii,

Bacutriletes clavatus, Bacutriletes
onodios, Verrutriletes aurantium
Lower part: Nathorstisporites
hopliticus, etc.

Sibireconcha anodontoides^
Qiyangia xiangxiensis
assemblage

are dominated by Nathorstisporites hopliticus

Jung, Horstisporites planatus (Marcinkiewicz)
Marcinkiewicz and Trileites sp., which indicate
an early Early Jurassic age, whereas the upper
part of the formation is represented by Bacutriletes harrisii Yang and Sun, Bacutriletes clavatus
Marcinkiewicz, Bacutriletes onodios (Harris) Yang
and Sun and Verrutriletes aurantium (Harris) Potonie (Yang and Sun, 1987). The bivalves of this
formation are characterised by a Sibriconcha anodontoides^Qiyangia xiangxiensis assemblage (Yu,
1987) (Table 1). All of this fossil evidence indicates that the geological age of the Hsiangchi Formation is Early Jurassic. The upper part of the
formation is therefore corresponding to the late
Early Jurassic, and is suggested to be Toarcian.

3. Materials and methods

All of the fossil plant material for this study
came from recent collections from the type locality of the Hsiangchi Flora in the town of Xiangxi,
Zigui County, Hubei Province (Fig. 1). Fossil
ferns have been collected from the upper part of
the Hsiangchi Formation. They were mostly preserved as compressions as well as impressions in a
bed of grey shale and black mudstone about 50
cm in thickness (see Wang, 1999). All the fossil
fern material is well preserved including both sterile and fertile pinnae. In addition, some bennettitalean leaves, such as Ptilophyllum, Pterophyllum,
Otozamites, Anomozamites, nilsssonialeans including Nilssonia, ginkgoaleans Sphenobaiera and
Baiera, as well as pteridospermous leaves Cteno-

zamites are frequently found with good preservation associated with ferns.
In addition, the fern specimens described by Sze
(1949), Wu et al. (1980) and Wu (1991) from
the same locality of the Hsianhchi Formation
were also re-examined for comparison and discussion. They were housed in the Palaeobotanical
Collection, Nanjing Institute of Geology and Palaeontology, Academia Sinica, Nanjing, China
(NIGPAS). Besides megaplant material, the palynological data published by Li and Shang (1980),
Zhang and Zhang (1987) were also considered to
reect the oral composition of ferns.
In order to reveal the abundance of the fossil
fern groups, the statistical analysis of the recent
collection is applied giving the percentage of each
fern taxa on generic level.
The selected specimens are gured herein with
registration numbers PB6724, PB18090, PB18139
and PB18894^PB18916, which are housed in Palaeobotanical Collection, NIGPAS, Nanjing, China.

4. Floral composition of the Hsiangchi Formation

and its fern diversity
The Lower Jurassic Hsiangchi Formation is
well known for its southern-type ora in China.
Newberry (1865/1866) was the rst to describe
plant fragments, collected by Pumpelly from Zigui, Hubei (possibly derived from the Hsiangchi
Formation). According to Schenk (1867), the
specimens collected by Richthofen in China,
were derived also from the same formation. Sze

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(1949) made a systematic study on the fossil ora

of the Hsiangchi Group in western Hubei. Afterwards, Wu et al. (1980) and Meng (1987) contributed greatly to the study of the fossil plants from
this locality and horizon. Presently, over 72 species belonging to 31 genera have been reported
from the formation (according to Meng, 1987).
Among them, cycadophyta and ferns are the
dominant groups, ginkgoales and conifers are
common, pteridosperms are quite rare and only
represented by Ctenozamites. The cycadophyta
are characterised by bennettitalean leaves, such
as Pterophyllum, Anomozamites, Otozamites, Ptilophyllum, Tyrmia, Weltrichia, Zamites, Ctenis,
Mironeura, and Nilssonialeans (Nilssonia). The
ginkgoaleans are represented by Ginkgoites,
Baiera, Sphenobaiera, the Czekanowskialeans by
Czekanowskia, Phenicopsis, Ixostrobus and Stenorachis. The conifers are only known as Podozamites, Ferganiella, Swedenborgia and Eladocladus.
It is noteworthy that the fossil plants in the
Lower Jurassic Hsiangchi Formation show a remarkable vertical diversity in western Hubei and
the equatorial horizon in adjacent areas of the
Yangtze River. Available data (Wu et al., 1980;
Huang, 1983, 1988; Meng et al., 1994) and the
present investigation show that ginkgoalean
plants are abundant in the lower part of the
Hsiangchi Formation, whereas licaleans and cycadoaleans are more common and predominant in
the upper part of the formation.
Among the ferns, over 20 species belonging to
nine genera have been reported so far from the
Hsiangchi Formation. They are grouped into ve
families, including Marattiaceae, Osmundaceae,
Matoniaceae, Dipteridaceae and Dicksoniaceae.
Sze (1949) described 18 species belonging to the
genera Marattiopsis ( = Marattia), Sphenopteris
( = Todites), Cladophlebis, Laccopteris ( = Phlebopteris), Dictyophyllum, Clathropteris and Coniopteris. Some of them were revised in the work by Wu
et al. (1980), in which eight species were described, including Marattiopsis asiatica Kawasaki,
Todites princeps (Presl) Gothan, Cladophlebis denticulata (Brongniart) Fontaine, Phlebopteris polypodioides Brongniart, Clathropteris obovata Oishi,
Thaumatopteris sp., Coniopteris cf. hymenophylloides (Seward), Coniopteris cf. murrayana (Brong-

129

niart) Brongniart and Coniopteris. sp. Meng

(1987) also described Phlebopteris ziguiensis from
Zigui Basin as well as Coniopteris chenyuanensis
Meng, Cladophlebis dangyangensis Meng, Cladophlebis complicata Meng from Dangyang Basin
in western Hubei.
Recently, several osmundaceous ferns belonging to Todites and cf. Osmundopsis were described
from the Hsiangchi Formation in Zigui, western
Hubei (Wu, 1991). Wang (1999), Wang and Mei
(1999) and Wang et al. (2001) published detailed
results of the fertile organs and in situ spores of
Marattia asiatica and Phlebopteris polypodioides.
A further study on matoniaceous ferns in this
region is under preparation (Wang and Van Konijnenburg-van Cittert, 2002).
In this investigation, the following taxa (including nine genera of ve families) are recognised
from the upper part of the Hsiangchi Formation
based on recent collections from Zigui County,
western Hubei (Table 2).
4.1. Marattiaceae
Marattia asiatica (Kawasaki) Harris is the only
representative member of the Marattiaceae recognised from the Hsiangchi Formation. Specimens
ascribed to this species were originally described
as Marattiopsis muensteri by Sze (1949) and Marattiopsis asiatica by Wu et al. (1980). They were
all impressions and yield no information of fertile
structures. A recent study by Wang (1999) and
Wang et al. (2001) revealed detailed structures
of fertile organs and in situ spores of this species.
Both sterile and fertile specimens of Marattia
asiatica are found in the upper part of the Hsiangchi Formation. The pinnae are linear and narrow
in shape, and reach a length of over 15 cm (Plate
I, 1). They vary from 24 to 30 mm wide in the
lower region and near the pinna base, and gradually taper from the base to the apical region. The
base is asymmetrical and auriculate (Plate I, 5).
The fertile pinnae are especially well preserved,
including the linear, bivalved synangia, which
are attached on the secondary veins of the pinna
margin (Plate I, 4). A synangium is composed of
25^40 pairs of sporangia, which yield a large
number of monolete spores with a granular exine

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Table 2
List of fern taxa recognized in this study from the upper part of the Lower Jurassic in Western Hubei
Families

Genera and species

(1) Marattiaceae
(2) Osmundaceae

Marattia asiatica (Kawasaki) Harris

Todites princeps (Presl) Gothan
Todites leei Wu
Todites williamsonii (Brongniart) Seward
Cladophlebis denticulata (Brongniart) Fountaine
Phlebopteris ziguiensis Wang and Van Konijnenburg-van Cittert ex. Meng
Phlebopteris muensteri (Schenk) Hirmer and Hoerhammer
Phlebopteris polypodioides Brongniart
Phlebopteris sp.
Dictyophyllum nilssonii (Brongniart) Goeppert
Dictyophyllum nathorstii Zeiller
Dictyophyllum rugosum Lindley and Hotton
Clathropteris obovata Oishi
Clathropteris platyphylla (Goeppert) Nathorst
Hausmannia ussuriensis Kryshtofovich
Hausmannia buchii Andrae
Thaumatopteris spp.
Coniopteris hymenophylloides (Brongniart) Seward
Coniopteris murrayana (Brongniart) Brongniart
Coniopteris chenyuanensis Meng
Coniopteris sp.

(3) Matoniaceae

(4) Dipteridaceae

(5) Dicksoniaceae

(Wang, 1999). Though specimens of M. asiatica

are always found with pinnae in western Hubei, it
seems that this plant has fronds that are at least
bipinnate with a possible third pinna rachis as
reported by Kimura and Tsujii (1980, 1984) of
specimens from the Early Jurassic of Japan. Similar fronds were also found in Marattia anglica
(Thomas) Harris from the Jurassic of Yorkshire
(Van Cittert, 1966).
4.2. Osmundaceae
About four species of the Osmundaceae are
recognised from the Hsiangchi Formation. Todites princeps is the most common one in our
collection. The specimens of this species are
mainly represented by sterile pinnae, though the

fragmentary fertile pinnule were gured by Wu et

al. (1980) and Wu (1991). The frond is bipinnate
with anadromic branching of the pinnules and
sphenopterid venation (Plates I, 2, 6, and II, 1).
The specimens described as Sphenopteris modesta
by Sze (1949), were similar in gross morphology
to T. princeps and were attributed to the latter by
Wu et al. (1980).
Specimens of Todites leei Wu are also bipinnate
with pecopterid frond architecture and simple venation (only fork one time) (Plate II, 3), the fertile
pinnules are strongly reduced (Wu, 1991). Todites
williamsonii (Brongniart) Seward was described by
Wu (1991) from the Hsiangchi Formation. The
sterile pinnae are catadromic in pinna and venation branching. The fertile pinnules are narrow
and much smaller than sterile ones. Another os-

Plate I. Marattia (Marattiaceae) and Todites, Cladophlebis (Osmundaceous) from the Lower Jurassic Hsiangchi Formation. All
gures U0.95 unless otherwise indicated. (1,4,5) Marattia asiatica (Kawasaki) Harris. (1) A larger fertile pinna from the collection of Wu et al. (1980) with a length of over 15.5 cm; PB6724. (4) A fragmentary but well preserved fertile pinna showing the
outline of synangia (from Wang, 1999, plate II, gure 6); PB18090. U2.85. (5) A sterile pinna with asymmetric and auriculate
base (from Wang, 1999, plate I, gure 6); PB18139. (2,6) Todites princeps (Presl) Gothan; PB18894 (2), PB18896 (6). (3) Cladophlebis denticulata (Brongniart) Fountaine; PB18895.

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mundaceous species, cf. Osmundopsis sturi (Raciborski) was described by Wu (1991) based on the
fragmentary specimens. However, re-examination
of the material shows that Wus specimens are
fertile pinnae of Todites (probably T. williamsonii). Unfortunately, the specimens are too small
to show the essential details needed for further
identication. Cladophlebis denticulata (Brongniart) Fountaine is at least bipinnate and has a
larger frond than Todites; and the pinnules are
typically sickle shaped with a denticulate margin
(Plate I, 3).
4.3. Matoniaceae
Phlebopteris is the only representative genus of
the Matoniaceae in the Hsiangchi Formation.
Phlebopteris ziguiensis Wang and Van Konijnenburg-van Cittert ex. Meng is very common in the
present collection. The specimens are characterised by larger fronds (length of pinnae over 14.5
cm, number of pinnae 6^10, even 15, length of
pinnules over 22 mm) with palmate or pedate
shape. From 10 to 17 pinnae arise from the
main rachis of the frond (Plates II, 2, 6, and III,
1, 2). The fertile pinnae are well preserved including sori with ca. eight sporangia. Sze (1949) rst
described one fragmentary specimen as Laccopteris cf. polypodioides, which was transferred as
Phlebopteris polypodioides Brongniart (Wu et al.,
1980) (Plate III, 5). The fertile organs and in situ
spores of P. polypodioides were studied by Wang
and Mei (1999) based on newly collected material.
Another member, Phlebopteris muensteri (Schenk)
Hirmer and Hoerhammer, has a frond of the
same shape and size as P. ziguiensis, but with
longer and narrower pinnules (Plate III, 3). Specimens of P. polypodioides and P. muensteri are less
abundant than those of P. ziguiensis in the present
collection. In addition, the young curved leaves
(genus Spiropteris) are frequently found in close
association with Phlebopteris fronds or pinnae

133

(Plate II, 4, 5). The creeping roots and stems are

found in the same fossiliferous bed (Plate III, 4).
Such evidence probably suggests almost autochthonous fossils.
4.4. Dipteridaceae
In western Hubei, four genera are recognised
for the family Dipteridaceae as Dictyophyllum,
Hausmannia, Clathropteris and Thaumatopteris.
The most common species is Dictyophyllum nilssonii (Brongniart) Goeppert. Specimens of this
species are frequently found with larger pinnae
or fronds (Plate IV, 1). The laminae of the pinnules are thick. The midribs of pinnules are prominent and broad; the main lateral veins arise from
the midrib at an angle of about 60. The secondary lateral veins form a network of irregular polygonal meshes, in which the nest veins are visible (Plate IV, 3). In fertile pinna, the sporangia
are very well preserved, covering the whole part of
pinnules (Plate IV, 4). In addition, Dictyophyllum
nathorstii Zeiller and Dictyophyllum rugosum
Lindley and Hutton are recognised in association
with Dictyophyllum nilssonii, but they are not
abundant. Hausmannia ussuriensis Kryshtofovich
is common in the present collection. The fanshaped lamina shows clear organisation of primary and secondary veins and typical meshes
(Plate V, 1, 2). The petiole and lamina are well
preserved (Plate V, 3).
4.5. Dicksoniaceae
The extant Dicksoniaceae are tropical tree ferns
with erect trunks and arborescent trunk habit.
The Mesozoic dicksoniaceous ferns are, however,
mostly smaller and shorter herbaceous plants.
Only one genus Coniopteris is recorded in the
Hsiangchi Formation, which includes Coniopteris
hymenophylloides Brongniart, Coniopteris murrayana (Brongniart) Brongniart, Coniopteris che-

Plate II. Todites (Osmundaceae) and Phlebopteris (Matoniaceae). All gures U0.95. (1) Todites princeps (Presl) Gothan, showing
the thick rachis and the pinnae; PB18897. (3) Todites leei Wu. A small specimen with thick rachis and clear venation; PB18898.
(2,6) Phlebopteris ziguiensis Wang and Van Konijnenburg-van Cittert ex. Meng. (2) The petiole and 11 pinna rachises; PB18899;
(6) A completely preserved frond with 17 pinnae; PB18902. (4) A young curved leaf Spiropteris (left) and the associated pinnae
of Phlebopteris ziguiensis (right); PB18900. (5) A young curved leaf; PB18901.

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nyuanensis Meng, and Coniopteris sp. Among

them, C. hymenophylloides is the most common
one with both the sterile and fertile pinnae preserved (Plate V, 4^6). All of these species are very
small and slender in growth form, less than 20 cm
tall.

5. Ecology implications
5.1. Fossil preservation and burial mode
As already mentioned, the ferns from the upper
part of the Hsiangchi Formation are preserved in
the peat-forming environments (Fig. 2). Fossils, in
general, are abundant and diversied in these lithologies. All of the fern fossils are found in the
same horizon, including not only the sterile leaves
with clear venation, but also the fertile leaves with
good reproductive organs of sporangia and in situ
spores (Plates I, 1, 4, and IV, 4; Wang, 1999;
Wang and Mei, 1999). The larger fronds of Matoniaceae and Dipteridaceae are nearly completely
preserved including petiole and the radiation of
pinnae (Plates II, 2, 6, III, 1, and IV, 1). Even
smaller fronds of Phlebopteris also show petiole
and pinnae (Plate III, 2). Sometimes, small pinnae
of Todites are found together with Phlebopteris in
the same slab (Plate III, 3). Smaller fronds, such
as those of Coniopteris, are also found with well
preserved sori (Plate V, 4). The young curved
leaves (Plate II, 4, 5) and the roots of ferns (Plate
III, 4) are frequently associated with the fern
fronds. In the fossil-bearing bed, ferns and other
gymnospermous leaves, especially bennettitaleans
are always closely associated and irregularly arranged along the bedding surface. These indicate
that the fossil plants were subject to minimal
transport. Therefore, we consider that the fossil
ferns were deposited near their living area. This

135

suggests an autochthonous or hypoautochthonous

burial in the present fossil-bearing horizon of the
Hsiangchi Formation.
5.2. Structural analysis and the fern community
Statistical analysis based on relative abundance
data were given using over 340 fern specimens
collected from the upper part of the Hsiangchi
Formation (Fig. 3). The Matoniaceae is the dominant group (42.6% of total fern specimens), represented by Phlebopteris. The Dipteridaceae rank
second (26.8%), but with greater of diversity, including Dictyophyllum, Hausmannia, Clathropteris
and Thaumatopteris. The Marattiaceae are relatively abundant (14.7%) and are characterised by
Marattia asiatica. The Osmundaceae (9.1%) and
Dicksoniaceae (5.3%) are less common.
According to palynological data (Li and Shang,
1980; Zhang and Zhang, 1987), dispersed spores
with known licalean anities from the upper
part of the Hsiangchi Formation are represented
by Dictyophyllidites, Concavisporites, Converrucosisporites, Marattisporites, Todisporites, Granulatisporites, Osmundacidites and Cyathidites. Based
on comparative analysis with fossil in situ spores
(e.g. Van Konijnenburg-van Cittert, 1975, 1978,
1989, 1993, 1996; Ash et al., 1982; Litwin, 1985;
Balme, 1995; Schweitzer et al., 1997; Wang, 1999;
Wang and Mei, 1999), the dispersed spores also
indicate the source data as Marattiaceae, Osmundaceae, Matoniaceae, Dipteridaceae and Dicksoniaceae. Furthermore, such data support the
abundance of ferns in the Hsiangchi Formation
similar to that found for megafossils as shown
in Fig. 3.
Comparison with the nearest living relatives
and their living habitats suggest that the fossil
ferns from the upper part of the Hsiangchi Formation are all herbaceous and hydrophytic. They

Plate III. Phlebopteris (Matoniaceae). All gures U0.95. (1) A larger frond of Phlebopteris ziguiensis Wang and Van Konijnenburg-van Cittert ex. Meng, showing the long, thick petiole and nine radiating fertile pinnae; PB18903. (2) A small frond of Phlebopteris showing the petiole and 11 pinnae rachises; PB18904. (3) A pinna of Phlebopteris muensteri (Schenk) Hirmer and Hoerhammer. In the right lower part of the gure (arrow), there is a fragmentary pinna of Todites princeps (Presl) Gothan; PB18905.
(4) The creeping root that is frequently found in association with Phlebopteris pinnae; PB18907. (5) A pinna of Phlebopteris polypodioides Brongniart; PB18906.

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Fig. 3. Statistical result showing the relative abundance (percentage) of the fern specimens collected from the Lower Jurassic Hsiangchi Formation in western Hubei. A, Marattiaceae; B, Osmundaceae; C, Matoniaceae; D, Dipteridaceae;
E, Dicksoniaceae.

are a typical peat-forming fern community dominated by the Matoniaceae. This community is
ecologically characterised by perennial herbaceous
species specialised for growth on shaded oor of a
wet, tropical to subtropical forest. These ferns
mostly prefer high humidity and low light conditions. Phlebopteris and Dictyophyllum with larger
fronds are more abundant and have higher cover
in this community. The Marattiaceae and Osmundaceae are often dispersed among the community.
Coniopteris of the Dicksoniaceae is smaller and
slender in size, and is distributed sparsely in the
community, probably on the marginal land
around the swamp. A reconstruction of the
present fern community is suggested and shown
in Fig. 4.
5.3. Climatic condition
During the Late Triassic to Middle Jurassic, the
western Hubei belonged to the Southern Floristic
Province of China, i.e. the Maritime Floristic
Province proposed by Zhou (1984) or the EuroSinian Region proposed by Vakhrameev (1991).
Fossil ferns are abundant and diverse in the

137

Hsiangchi Formation and are represented by the

present fern community described herein that is
typical of the Southern Floristic Province, which
is indicative of the climatic condition.
The living relatives of the above ferns are
mainly restricted to tropical and subtropical regions. The Matoniaceae is a relict family of the
Malesian region with two genera. The Dipteridaceae is restricted to the Indo-Malayan region and
southern China. The Marattiaceae are tree ferns
ranging in the palaeotropics from Africa to the
Pacic. The Osmundaceae are all perennials,
which includes three extant genera: Osmunda is
distributed throughout most temperate and tropical regions; Todea and Leptopteris are native in
South Africa and Australia. Extant members of
the Dicksoniaceae are mainly tree ferns, distributed in tropical and subtropical regions. Though
the Mesozoic ferns dier from the extant relatives
in gross morphology, their climate tolerances are,
however, very similar. In general, the peat-forming fern community described above dominated
by the Matoniaceae apparently indicates a tropical to subtropical warm and humid climate, which
is conductive for coal formation and accumulation. As western Hubei is located on the north
coast of the Tethys Ocean during the Early Jurassic, the warm and humid marine current is favourable for the development of a fern community in
a coal swamp environment. However, at the same
time, few ferns have been distributed in the
Northern Floristic Province of China due to the
little eect of the marine currents there.
In addition, sedimentary studies (Meng et al.,
1994) indicate that the clay minerals of the upper
part of the Hsiangchi Formation are characterised
by the humid climate indicator minerals, kaolinite
and illite (35% and 20%, respectively) as well as
siderate, whereas the arid or dry climate indicators, such as chlorite and hydromica, are uncommon. These ndings agree with the climatic conditions suggested by the fossil fern community.

Plate IV. Dictyophyllum nilssonii (Brongniart) Goeppert (Dipteridaceae). 1 and 2, U0.95; 3 and 4, U2.85. (1) A larger frond
showing the basal part of four pinnae; PB18908. (2) A pinna near the apical part of a frond; PB18909. (3) Enlargement of 2
showing the venation and the impression of sori. U2.85. (4) Enlargement of a fertile pinna showing two pinnules that are covered by sporangia; PB18910.

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139

Fig. 4. A suggested reconstruction of the fern community dominated by Matoniaceae in the upper part of the Lower Jurassic
Hsiangchi Formation in western Hubei. 1, Matoniaceae; 2, Dipteridaceae; 3, Marattiaceae; 4, Osmundaceae; 5, Dicksoniaceae.

Furthermore, the above-mentioned vertical diversity of fossil plants in the Hsiangchi Formation
shows a variation of climate condition. During
the early stage of Early Jurassic, the sedimentary
basin in Zigui was characterised by high and precipitous terrain as well as developed rivers and
creeks, and the climate was pleasantly cool with
lower humidity. This condition was favourable for
the growth of ginkgoaleans and conifers, whereas
the growth of cycadophytes and ferns was restrained. During the late stage of Early Jurassic,
the basin was marked by smooth terrain, lakes
and swamp were widely distributed. The climate
changed to warm and humid, which was favourable for development of ferns and cycadophytes

as well. The fern community recognised in this

study from the Hsiangchi Formation is, therefore,
a representative fern ecology community in the
Southern Floristic Province of China during the
Early Jurassic.

6. Concluding remarks
(1) Ferns are the conspicuous to dominant
components of the Early Jurassic Hsiangchi Formation in western Hubei, China, including at least
nine genera in ve families, such as Marattiaceae,
Osmundaceae, Matoniaceae, Dipteridaceae and
Dicksoniaceae.

Plate V. Hausmannia (Dipteridaceae) and Coniopteris (Dicksoniaceae). All gures U0.95. (1^3) Hausmannia ussuriensis Kryshtofovich. (1,2) Showing the primary and secondary venation as well as typical meshes; PB18911 (1), PB18912 (2). (3) Part of a frond
showing half of the lamina and petiole; PB18913. (4^6) Coniopteris hymenophylloides (Brongniart) Seward. (4) A bipinnate fertile
frond with sori and a sterile pinna; PB18914. (5,6) Bipinnate sterile fronds with pinnae; PB18915 (5), PB18916 (6).

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(2) Ferns are abundant, diverse and well preserved in the upper part of the Hsiangchi Formation. The nature of the preservation and the associated sedimentary conditions suggest that ferns
from this horizon represent an autochthonous or
hypoautochthonous uviatile peat-forming community dominated by Phlebopteris of the Matoniaceae.
(3) This fern community indicates a warm and
humid, tropical to subtropical climate. It is a remarkable and representative fern community recognised for the late Early Jurassic in the Southern
Floristic Province of China.

Acknowledgements
I would like to offer my sincere thanks to Dr.
Johanna H.A. Van Konijnenburg-van Cittert
(Utrecht) and Dr. Margaret Collinson (London)
for their kind invitation and encouragement of the
present contribution to the special volume, Fern
Ecology through Time. Special thanks are due to
Prof. Zhou Zhiyan (Nanjing) for his kind suggestions and valuable comments on this study. I am
also grateful to Prof. Meng Fansong (Yichang),
Prof. Liu Lujun, Mr. Mei Shengwu, Ms. Leng Qin
(Nanjing) for their help and assistance in collecting
fossil materials. I thank Prof. David L. Dilcher
(Florida Museum of Natural History, University
of Florida), Prof. William A. DiMichele (National
Museum of Natural History, Smithsonian Institution, USA) and Dr. Johanna H.A. Van Konijnenburg-van Cittert (University of Utrecht) for their
critical review and valuable suggestions to the
manuscript. This work was financially supported
by the National Natural Science Foundation of
China (NSFC) (39900007), the Talent Training
Fund from NSFC (990202), the Special Funds of
Palaeontology and Palaeoanthropology (SFPP)
from CAS (990202), as well as the Innovation
Research Fund from NIGPAS, Nanjing, China
(000213). This study was also partly supported by
the Executive Office of State Council Three
Gorges Project Construction Committee, and the
China Yangtze Three Gorges Project Development Corporation of China.

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