全 文 ：Anatomically_preserved Lepidodendralean Plants from Permian Coal
Balls of China:Sigillariopsis Scott
WANG Shi_Jun1 , TIAN Bao_Lin2 , CHEN Gui_Ren2
(1.Inst itute of Botany , The Chinese Academy of Sciences , Beijing 100093 , China;
2.China University of Mining and Technology , Beijing 100083 , China)
Abstract:　Three kinds of anatomically_preserved lepidodendralean leaves from Permian coal balls of China
were studied.They all have double xylem strands , which is consistent with the genus Sigillariopsis Scott.
Compared with the species of Sigillariopsis , they are considered as three new species:S.shanxiensis sp.
nov.and S .taiyuanensis sp.nov.f rom Coal Seam 7 in the upper part of the Taiyuan Formation(lower Lower
Permian), Xishan Coal Field , Taiyuan , Shanxi Province and S .guizhouensis sp.nov.from Coal Seam 1 in
the Wangjiazhai Formation(upper Upper Permian), Shuicheng Coal Mining District , Guizhou Province.Based
on the associated organs of lepidodendraleans other than leaves and the information of the Euramerican lepi-
dodendraleans , the affinities of the three new species of Sigillariopsis are discussed and they are probably leav-
es of Sigillaria Brongniart.In Euramerican Flora , Sigillaria (including its leaves Sigillariopsis)mainly lived
in the Carboniferous period and in the Cathaysian Flora they lived in the Upper Carboniferous to upper Upper
Permian periods in rare localities.The leaves and fertile organs of Sigillaria have not been reported from the
Cathaysian Flora to date.Three new species of the Permian anatomically_preserved leaves of Sigillaria not only
increase the diversity of the Cathaysian sigillarian but also bear important significance on the evolution of sigil-
larian plants and the relationship of the Cathaysian and Euramerican lepidodendraleans.
Key words:　Cathaysian Flora;Permian;coal ball;lepidodendralean leaves;Sigillariopsis;new species
　　Different from Lepidodendron Sternberg which domi-
nated lepidodendralean plants in the Cathaysian Flora and
more than 60 morphological species which have been re-
ported from the Lower Carboniferous to the Upper Permian
in numerous localities
[ 1-5] , Sigillaria Brongniart was far
less developed in the Cathaysian Flora and only several
species have been described from rare localities of the Up-
per Carboniferous to Permian
[ 1 ,2] .Leaves and cones of
Sigillaria have never been reported.In contrast to this
condition in the Cathaysian Flora , Sigillaria has been
well studied in the Euramerican Flora , including the mor-
phology and anatomy of all kinds of organs , such as leav-
es , stems , roots and cones[ 6-12] .
In coal balls of the lower Lower Permian and upper
Upper Permian from North and Southwest China , there are
abundant anatomically_preserved lepidodendralean leaves
including Lepidophylloides Snigirevskaya with single vas-
cular bundles and Sigillariopsis Scott with double vascular
bundles.This paper deals with Sigillariopsis.Lepido-
phylloides will be described in another paper by the same
authors.This is the first detailed study on the anatomical-
ly_preserved leaves of Sigillaria from the Cathaysian Flo-
ra.
1　Materials and Methods
Sigillariopsis studied in this paper is anatomically_
preserved leaves in coal balls from Coal Seam 7 in the up-
per part of the Taiyuan Formation , Taiyuan , Shanxi Prov-
ince and from Coal Seam 1 in the Wangjiazhai Formation ,
Shuicheng , Guizhou Province.The age of the Coal Seam
7 is early Early Permian and the Coal Seam 1 is late Later
Permian
[ 13] .The detailed geological backgrounds of two
localities of coal balls were introduced by Tian and
Wang
[ 13] .The acetate peel technique was employed in
this paper to reveal anatomical information
[ 14 ,15] .
2　Systematics
Genus Sigillariopsis Scott 1904 , non Sigillariopsis
Renault 1879.
Type species:Sigillariopsis cordata Reed 1941.
2.1　Sigillariopsis guizhouensis sp.nov.(Figs.1-
7)
2.1.1 　Diagnosis 　Leaf lamina_like , up to 15 mm
broad and 1.5 mm high , with a narrow apex.Upper sur-
face with a median depress and lower surface with a large
midrib in the apical part of the leaf.Vascular bundle
sheath ω_shaped at the apical part and {_shaped at the
middle and basal part of the leaf with double xylem strand
lying at the inside of the arm of vascular bundle sheath.
Vascular bundle sheath thick and consisting of small
thick_walled fibrous cells.Transfusion tissue present.
Hypodermis thin.
Holotype:Coal ball:F31;Slide:F31_a
Paratype:Coal ball:F31;Slide:F31_b
Received:2001-05-10　Accepted:2001-07-04
Supported by the National Natural Science Foundation of China (39970056)and Special Funds for the Palaeontology and Palaeoanthropology from the Chinese
Academy of Sciences(QTE_98).
植　物　学　报　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　
Acta Botanica Sinica　2002 , 44(1):104-112
Etymology:The specific name is derived from the fossil
locality Guizhou Province
Stratigraphy and location:Wangjiazhai Formation , up-
per Upper Permian;Shuicheng Mining Area , Guizhou
Province
2.1.2　Description　The new species is represented by
only two specimens in cross section in the coal ball F31
from Guizhou Province.One , F31_a , probably represents
a section near the apical part of the leaf , another one ,
F31_b , probably represents a section of the middle or
basal part of the leaf.They are characterized by having aω_or {_shaped vascular bundle sheath of thick_walled
cells-although they are very different in shape and size.
Shape and size:Specimen F31_a has two sections , F31_
a_1 and F31_a_2.In section F31_a_1 , the leaf is entirely
preserved and is 2.2 mm broad and 1.54mm high.There
is a deep V_shaped median depression on the upper sur-
face.The leaf margin turns downward.There is a large
midrib on the lower surface with a short keel (Fig.6).
Specimen F31_b is partly_preserved but its vascular bun-
dle is entirely_preserved(Figs.1 , 2).It is flat and lami-
na_like with a supposed width of 15 mm and a height of
1.5 mm or more.Lateral stomatiferous furrows are absent
in both specimens.Stomatiferous region is located at the
lower surface of the wings of lamina.
Vascular bundle:The vascular bundle sheath is ω_
shaped in specimen F31_a(Figs.1 , 2 , 5 , 6).The vas-
cular bundle sheath is about 620 μm broad and 520 μm
high.The arm of bundle sheath is about 60 μm thick at
the middle part and decrease in thickness toward the end(Figs.5-7).In specimen F31_b , the vascular bundle
sheath is{_shaped and much broader , up to 3.4 mm ,
than that of specimen F31_a.The arm is up to 250 μm
thick at the middle part(Fig.2).Transfusion tissue is in
a continuous zone , 1 -2 cells thick , surrounding the
bundle sheath.Transfusion tracheids possess sparse and
spiral thickening patterns(Fig.3).　　The double xylem strand is located at the inside of
the arm of the bundle sheath.In specimen F31_a , they
are 1-2 cells thick and about 300μm long , consisting of
16-18 tracheids with the smallest ones at the outer ends(Figs.5-7).In specimen F31_b , the xylem strand is
up to 1.1-1.5mm long and 3-4 cells thick , consisting
of more than 40 tracheids (Fig.2).The large tracheids
possess scalariform thickening patterns with fibrils (Figs.
3 , 4).
Phloem is difficult to recognize because of poor pres-
ervation.
Mesophyll:Mesophyll is poorly_preserved in specimen
F31_b and is difficult to describe.In specimen F31_a ,
mesophyll is undifferentiated.Several layers of cells be-
neath the hypodermis on the upper surface are small ,
about 18 -22 μm in diameter , and are compact (Fig.
6).Mesophyll around the vascular bundle consists of
large and thin_walled cells , 22 -46 μm in diameter ,
which are also compact.Mesophyll in other places is
poorly_preserved.
Hypodermis:In specimen F31_a , the hypodermis is thin
and uniform in thickness(Fig.6).In specimen F31_b ,
the hypodermis is about 4-5 cells thick at the lower sur-
face and a little thinner at the upper surface , consisting of
thick_walled fibrous cells(Fig.2).
Stomatiferous region:Stomatiferous regions are located
at the lower surface of the wings of lamina where the hy-
podermis is absent.In specimen F31_a , it is difficult to
determine the width of the stomatiferous region.In speci-
men F31_b , the stomatiferous region is about 1 mm or
more in width.
2.1.3　Comparison 　Present specimens are strongly
characterized by their ω_or {_shaped vascular bundle
sheaths which are very different from those of the Eu-
ramerican species of Sigillariopsis , such as S.cordata
Reed , S .halifaxensis Graham , S .laevis Koopmans ,
S .sulcata Scott and the leaves of Sigillaria brardi(Fig.
21 , a-e).Furthermore , the present specimens are very
large , up to 15 mm broad , compared to other anatomical-
ly_preserved species of Sigillariopsis in the Euramerican
Flora.For example , leaves of Sigillaria brardi , the larg-
est of the Euramerican species of Sigillariopsis , are only 5
-6 mm in breadth[ 16] .
2.2　Sigillariopsis shanxiensis sp.nov.(Figs.8-
14)
2.2.1　Diagnosis　Leaves 1.6-8.0 mm broad and
0.5-2.5 mm high.Upper surface convex to slightly
concave with a V_shaped median depression.Leaf margin
turning downward.Lower surface with a large midrib and
lateral stomatiferous furrows at the wings of lamina.Later-
al stomatiferous furrows with a narrow opening and a broad
and flat bottom , 270 -1 200 (usually 500-700)μm
broad.Xylem strand from a single one at the apical part
to a double one at the middle part and near the base of
the leaf , 1-2 cells thick and consisting of 15 to 40 or
more tracheids.Small tracheids at the outer and inner
ends of the xylem strand.Often the small tracheids at the
inner ends jointing the double xylem strand together.Vas-
cular bundle sheath triangular at the apical part and wide
U_shaped at the other places with a breadth of 0.4-2.7
mm and thickness of 0.2-0.3 mm.Transfusion tissue
present.Mesophyll undifferentiated.Hypodermis moder-
ately thick.
Holotype:Coal ball:T7_85B Bot;Slide:W094
Paratype:Coal ball:T7_74;Slide:WP11 0063
Etymology:The specific name is derived from the fossil
locality Shanxi Province
Stratigraphy and location:Coal Seam 7 , upper part of
the Taiyuan Formation , lower Lower Permian;Xishan
Coal Field , Taiyuan , Shanxi Province
2.2.2　Description　There are about 20 specimens be-
longing to this new species represented by cross sections
and oblique cross sections of the leaf.The leaves vary
greatly in breadth in which the narrow ones are probably
the young leaves or the apical part of the leaf and the wide
ones are probably the adult leaves or the middle and basal
parts of the leaf.Among these specimens three small ones
are somewhat different from others although we consider
WANG Shi_Jun et al:Anatomically_preserved Lepidodendralean Plants from Permian Coal Balls of China:Sigillariopsis Scott 105　
106　 植物学报　Acta Botanica Sinica　Vol.44　No.1　2002
them to belong to this new species.
Shape and size:The leaves have a size of 1.6-8.0
mmwide by 0.5-2.5 mm high.In smaller specimens
with a width less than 2mm , the upper surface is slightly
convex with a shallow V_shapedmedian depression.There
is a triangular midrib on the lower surface , sometimes
with a distinct keel(Figs.8 , 11).In specimens with a
width over 2 mm , the upper surface of the leaf is usually
flat or somewhat concave with a deep U_or V_shaped me-
dian depression.There is a large midrib , sometimes with
a short keel(Fig.9).Both smaller and larger leaves have
jar_shaped lateral stomatiferous furrows on the lower sur-
face (Figs.8 ,9 ,11).
Vascular bundle:In specimens with a breadth of less
than 2 mm , their vascular bundle sheath is inversely tri-
angular in shape with its upper side flat or somewhat con-
cave(Figs.8 ,11).However in specimens with a breadth
of more than 2 mm , their vascular bundle sheath is wide
U_shaped with a width of 0.4-2.7 mm and a height of
22〗0.2-0.3 mm(Fig.9).The sheath consists of 1-2
layer of small thick_walled cells(Fig.10).
The xylem strand is usually 1-2 cells thick and
consists of 15-40 or more tracheids in large specimens
and 10 -12 tracheids in some small specimens.The
small tracheids are located at the outer ends of the xylem
strands and also at the inner ends and connect the double
xylem strands together (Figs.8 , 10).In oblique cross
section , the small tracheids connecting the double xylem
strands bear spiral thickening patterns(Fig.14).A zone
of poorly_preserved tissue encloses the xylem strand which
is probably the phloem(Fig.10).
Transfusion tissue:Transfusion tissue is in a continuous
zone , 1-2 cells thick , and surrounds the bundle sheath.
Transfusion tracheids possess sparse and spiral thickening
patterns(Fig.14).
Mesophyll:Mesophyll is basically undifferentiated and
no palisad tissue is present in most specimens.Mesophyll
cells around the vascular bundle are isodiametric and
densely_arranged , while in other places mesophyll consists
of poorly_preserved chains of parenchymatous cells(Figs.
8 ,11 ,13).
Hypodermis:The hypodermis consists of thick_walled fi-
brous cells and is moderately thick except for the median
depression on the upper surface and the leaf margin where
it is somewhat thicker than at the other places , up to 5-
6 cells in thickness (Fig.9).In some specimens , the
hypodermis of the upper surface is slightly thicker than
that of the lower surface (Fig.11).Occasionally large
secretory cells containing dark_brown content can be seen
in the hypodermis(Fig.9).No hypodermis is present at
the stomatiferous furrows(Figs.8 ,9 ,11).
Stomatiferous furrows:Stomatiferous furrows are jar_
shaped or U_shaped with a narrow opening and a broad
and flat bottom , from 270-450μm broad in small speci-
mens with single xylem strand(Fig.11)to 440-1 200μm broad in large specimens with double xylem strands(Figs.8 ,9).
2.2.3　Comparison　 Three specimens with a single
xylem strand included in this new species are very small ,
only 1.5-1.8 mm broad.Each of them possesses a long
and stalk_like keel on the lower surface , an inversely tri-
angular vascular bundle sheath , a shallow median depres-
sion on the upper surface and narrow stomatiferous furrows(Fig.11).Generally those with double xylem strands are
relatively broad and each of them possesses a short keel ,
a wide U_shaped vascular bundle sheath , a deep median
depress and broad stomatiferous furrows (Fig.9).How-
ever , some specimens with double xylem strands are also
very small and have an inversely triangular vascular bun-
dle sheath(Fig.8), although the two xylem strands are
very close to each other , which makes it possible that the
double xylem strand is formed by the division of a single
one.The specimens with single xylem strand probably
represent the apical part of a leaf and those with double
xylem strand represent the middle and basal part of the
leaf.
The present specimens are larger than most of Eu-
ramerican species of Sigillariopsis.In the latter the leaf is
usually less than 4 mm in breadth , except for the leaves
of Sigillaria brardi Brongn.which has a breadth of up to
6 mm near the base of the leaf.The vascular bundle of
the present specimens is very different from that of the
Euramerican species of Sigillariopsis in shape and size.
In the former it is enclosed and wide U_shaped , while in
the latter it is usually open and narrow(Fig.21 , a ,b ,d ,
e).Although S .cordata Reed also has an enclosed vas-
cular bundle , it is heart_shaped and narrow in cross sec-
tion(Fig.21 , c)and is different from the present speci-
mens.
→
Figs.1-7.　Sigillariopsis guizhouensis sp.nov.1.Cross section , partly_preserved and lamina_like , arrows show vascular bundle , Slide:
F31_b , Scale bar=1 mm.2.High magnification of Fig.1 , showing{_shaped vascular bundle sheath(bs)and xylem strands(xs), Scale bar
=500μm.3.High magnification of the area ② in Fig.2 , showing part of the vascular bundle , t , transfusion tissue , Scale bar=100μm.4.
High magnification of the area ① in Fig.2 , showing scalariform tracheids with fibrils , Scale bar=50μm.5.Cross section of the apical part
of the leaf , partly_preserved , showing well_preserved ω_shaped vascular bundle sheath(bs)and the double xylem strand(xs), Slide:F31_a_
2 , Scale bar=250 μm.6.A slightly obliquely cross section of the apical part of the leaf with ω_shaped vascular bundle sheath , Slide:F31_
a_1 , Scale bar=500 μm.7.High magnification of Fig.6 , showing a part of the vascular bundle , Scale bar=100μm.Figs.8-11.Sigil-
lariopsis shanxiensis sp.nov.8.Cross section near the apical part of the leaf , noting the small tracheids between two xylem strands(arrow ①)
and at the outer ends of the xylem strands(arrow ②), Slide:WP11 0063 , Scale bar=250μm.9.Cross section of the middle part of the leaf ,
noting wide U_shaped vascular bundle sheath(bs)and secretory cells in the hypodermis (arrows), Slide:W094 , Scale bar=500 μm.10.
High magnification of Fig.9 , arrow showing the small tracheids connecting two xylem strands , Scale bar=100μm.11.Probable apical part of
the leaf with single xylem strand , noting the long keel(arrow)on the midrib , Slide:WP11 0062 , Scale bar=250μm.
WANG Shi_Jun et al:Anatomically_preserved Lepidodendralean Plants from Permian Coal Balls of China:Sigillariopsis Scott 107　
108　 植物学报　Acta Botanica Sinica　Vol.44　No.1　2002
2.3　Sigillariopsis taiyuanensis sp.nov.(Figs.15-
20)
2.3.1　Diagnosis　Leaves 5-6 mm broad and 0.6-
1.2 mm high.Upper surface concave with a U_shaped
median depression.Lower surface with a large midrib ,
sometimes prominent into a short keel , and with lateral
stomatiferous furrows near the margin of the leaf.Leaf
margin turning downward.Stomatiferous furrows jar_
shaped with a narrow opening and a wide and protuberant
bottom.Vascular bundle large and wide U_shaped , 1.8-2.6 mm broad.Xylem strand double and thin , 1-3(usually 2)cells thick , with smaller elements at the outer
ends.The double xylem strands far away from each other
with a mass of thick_walled cells between them.Bundle
sheath of thick_walled fibrous cells and thin , only 1-2
cells thick.Transfusion tissue present.Hypodermis thin
and uniform in thickness.
Holotype:Coal Ball:TN_8B;Slide:W113
Paratype:Coal Ball:TN_3;Slide:WP11 0061
Etymology:The specific name is derived from the fossil
locality Taiyuan City
Stratigraphy and location:Coal Seam 7 , upper part of
the Taiyuan Formation , Lower Permian;Xishan Coal
Field , Taiyuan , Shanxi Province
2.3.2　Description　There are four specimens belong-
ing to this new species and they are all cross sections of
the leaf.They probably represent the sections of the mid-
dle and basal part of the leaf.No sections of the apical
part of the leaf have been found.
Shape and size:The shape and size of the present speci-
mens are very close to that of Sigillariopsis shanxiensis
sp.nov.(Figs.15 , 18)and need not be described
here.
Vascular bundle:Vascular bundle sheath is also wide U_
shaped , 1.8-2.6mm broad and consists of thick_walled
fibrous cells.The upper side is slightly thinner , only one
cell thick , than that of the lower side , 1-2 cells thick(Figs.17 , 20).
The leaf has double xylem strands which are only 1-2(occasionally 2-3)cells thick.Each of them con-
sists of more than 30 tracheids (Fig.20).The double
strands are far away from each other and are separated by
a mass of thick_walled fibrous cells(Figs.19 , 20).Sur-
rounding the xylem strands is a continuous zone of thin_
walled cells which may be the phloem (Fig.20).
Mesophyll:Mesophyll is undifferentiated.Around the
vascular bundle are densely_arranged larger and isodia-
metric thin_walled cells.In other places , mesophyll most-
ly consists of poorly_preserved chains of parenchymatous
cells(Figs.19 , 20).
Beneath the hypodermis there is a zone , 1-2 cells
thick , of densely_arranged larger and isodiametric thin_
walled cells(Fig.20).
Hypodermis:The hypodermis is thin and uniform in
thickness.It is a little thicker at the median depression
on the upper side than in other places.Hypodermis is ab-
sent at the lateral stomatiferous furrows.
Lateral stomatiferous furrows:The lateral stomatiferous
furrow is jar_shaped in cross section with a narrow opening
and a broad bottom.The bottom is protuberant and 300-
500μm broad.Cells beneath the epidermis at the bottom
of the lateral stomatiferous furrow are loosely_arranged
with an increasing diameter inwards(Fig.16).
2.3.3　Comparison　The present specimens with dou-
ble xylem strands are very close to Sigillariopsis shanxien-
sis sp.nov.in the shape , size and the vascular bundle
sheath.However , there are distinct differences between
them.Firstly , the bottom of the stomatiferous furrow is
narrower and protuberant in the present specimens , while
it is broader and flat in S.shanxiensis sp.nov.Second-
ly , there are small tracheids between the double xylem
strands which connect them together in S .shanxiensis
sp.nov., while there is a mass of thick_walled fibrous
cells instead of small tracheids between the double xylem
strands such as those in S.shanxiensis sp.nov.Third-
ly , in the present specimens there usually is a zone of
slightly larger , isodiametric and polygonal thin_walled
cells beneath the hypodermis , 1-2 cells thick , which
are different from the mesophyll consisting of chains of
parenchymatous cells , while in S.shanxiensis sp.nov.
this zone of thin_walled cells is absent.
The present specimens are characterized by an en-
closed and wide U_shaped vascular bundle , a mass of
thick_walled cells between the double xylem strand and
the protuberant bottom of the stomatiferous furrows which
are not present in the Euramerican species of Sigillariop-
sis(Fig.21;Table 1).
3　Discussion
3.1　Affinities of three new species of Sigillariopsis
Chaloner
[ 17]
pointed out that the leaves of some spe-
cies of Sigillaria , such as S.elegans , possess double
xylem strands while those of some other species possess a
single xylem strand , such as S .bretoni , and they are not
too much different from the leaves of Lepidophylloides in
structure .It is possible that Lepidophylloides includes the
→
Figs.12-14.　Sigillariopsis shanxiensis sp.nov.12.Oblique section of a leaf , Slide:WP11 0064 , Scale bar=500μm.13.High magnifi-
cation of Fig.12 , showing small tracheids at the inner end of a xylem strand(arrow), Scale bar=250μm.14.High magnification of Fig.13 ,
noting small tracheids (arrows) connecting two xylem strand , Scale bar=100 μm.Figs.15-20.Sigillariopsis taiyuanensis sp.nov.15.
Cross section , arrows showing the lateral stomatiferous furrows , Slide:W113 , Scale bar=500μm.16.High magnification of a stomatiferous
furrow from Fig.15 , noting the protuberant bottom(arrow), Scale bar=100 μm.17.Part of Fig.15 , showing the wide U_shaped vascular
bundle and a mass of thick_walled cells between two xylem strands(arrow), Scale bar=250 μm.18.Cross section , arrow shows a mass of
thick_walled cells between two xylem strands , Slide:WP11 0061 , Scale bar=1 mm.19.High magnification of a part from Fig.17 , showing the
mass of thick_walled cells(arrow)between two xylem strands , Scale bar=140μm.20.Part of Fig.18 , arrow ① shows the mass of thick_
walled cells between two xylem strands and arrow ② shows the large parenchymatous cells beneath the hypodermis , Scale bar=250μm.
WANG Shi_Jun et al:Anatomically_preserved Lepidodendralean Plants from Permian Coal Balls of China:Sigillariopsis Scott 109　
Fig.21.　Line_drawings of species of Sigillariopsis Scott from the Euramerican and Cathaysian floras(all ×15).
a, S .laevis Koopmans;b , S.halifaxensis Graham;c , S.cordata Reed;d , Leaf of Sigillaria brardi Brongn.;e , S .sulcata Scott;f ,
S.guizhouensis sp.nov.;g , S.shanxiensis sp.nov.;h , S.taiyuanensis sp.nov.Dotted area represents thick_walled fibrous cells;Solid
black area represents xylem;Hatch area represents transfusion tissue.(a from Koopmans , 1928;b , d and e from Graham , 1935;c from
Reed , 1941).
leaves of both Lepidodendron and Sigillaria in part.
However , Sigillariopsis only includes the leaves of Sigil-
laria.
In coal balls yielding Sigillariopsis guizhouensis sp.
nov., there is a kind of Sigillaria_like stem without ex-
tra_xylem tissue preserved , which has been preliminarily
studied by Guo and Tian
[ 2]
and identified as S .cf brar-
dii.The stem is obviously different from the Euramerican
S .brardii and is probably a new species.In one coal
ball(F_31), the stem S.cf brardii and the leaves of Si-
gillariopsis guizhouensis sp.nov.are associated with each
other.There is a great possibility that Sigillariopsis
guizhouensis sp.nov.is the leaf of S .cf brardii.　　 In coal balls yielding Sigillariopsis shanxiensis sp.
nov.and S .taiyuanensis sp.nov.from Coal Seam 7 in
the Xishan Coal Field , Taiyuan , Sigillaria_like stem
fragments have been found
[ 18] .It is possible that Sigillar-
iopsis shanxiensis sp.nov.and S.taiyuanensis sp.nov.
are also leaves of Sigillaria.
3.2　About the genus Sigillariopsis Scott
Renault
[ 19]
proposed a new genus Sigillariopsis for a
kind of anatomically_preserved small branch with leaves
and the specific name S.decaisnei was suggested for it.
The leaves have double xylem strands which are very dif-
ferent from previously described lycopod leaves which pos-
sess a single xylem strand.Scott[ 20] used the generic name
Sigillariopsis to describe a kind of lepidodendralean leav-
es with an obvious double xylem strand and considered it
as a new species of Sigillariopsis , naming it as S.sulca-
ta.However , Emberger(1944)was able to make out that
S .decaisnei Renault is probably the leaf of Poroxylon , a
kind of gymnosperm plant , because their tracheids possess
bordered thickening patterns.In this condition , S .cor-
data Reed was chosen as the type species of Sigillariop-
sis
[ 17] .
Up to now , about six species of Sigillariopsis have
been described from the Euramerican Flora:S .cordata
Reed , S .elegans(Calder)comb.nov., S .halifaxen-
sis Graham , S.laevis Koopmans , S.sulcata Scott and
the leaves of Sigillaria brardi Brongn.Together with the
three new species described in this paper from the Cathay-
sian Flora , there are nine species in Sigillariopsis Scott.
All these species are very close in shape in cross section:
upper surface more or less concave with a median depres-
sion , lower surface with a midrib which usually extend in-
to a keel , the wings of lamina are thin.However , in Lep-
idophylloides Snigirevskaya the upper surface of most spe-
cies is convex without median depression , such as L .ac-
iculum , L.alatum , L .angulatum , L.minor , L .tri-
chosulcata , L .latifolium , L .sewardii , L .papillona-
ceum
[ 16 , 21] .Furthermore , the midrib of some species of
Lepidophylloides doesn t extend into a keel , such as L.
alatum , L .minor , L.latifolium , L.sewardii , L.
papillonaceum , L.thomasii , leaves of Lepidodendron
esnotense and L .veltheimianum , Type C of Grah-
am
[ 16 , 21] .L .equilaterale has a triangular shape in cross
section , which is very unusual in lepidodendralean lea-
s ves
[ 16 ] .
Different from the Euramerican species of Sigillari-
opsis which are mostly less than 4 mm in breadth , the
three new species in this paper are larger , up to 5-8mm
broad in S .shanxiensis sp.nov.and S .taiyuanensis
sp.nov.and even more , up to 15 mm broad , in S.
guizhouensis sp.nov.Also , the vascular bundle sheath
of the Cathaysian species is wider than that of the
Euramerican ones , up to 2.6-2.7 mm wide in
110　 植物学报　Acta Botanica Sinica　Vol.44　No.1　2002
Table 1　Comparison of species of Sigillariopsis Scott
Characters
Species
Sigilariopsis
cordata
Sigilariopsis
halifaxensis
Sigilariopsis
laevis
Sigi lariopsis
sulcata
Leaf of
Sigillaria brardi
Sigilariopsis
guizhouensis
Sigi lariopsis
shanxiensis
S igilariopsis
taiyuanensis
Size
3 mm wide , 1
mm high
2 mm wide ,
1.3 mm high
3 - 5 mm
wide , 1.0 -
1.3 mm high
3.5 mm wide ,
1.6 mm high
Up to 5 - 6
mm wide , 1.5
-2.0 mm high
Up to 15 mm
wide , 1.5 mm
high
Up to 8 mm
wide , 2.5 mm
high
Nearly same as
that of S.
shanxiensis
Stomatifer_
ous
furrows
Present Present Absent Present Present Absent Present Present
Vascular
bundle
sheath
Enclosed and
heart_shaped ,
2 - 3 cells
thick
V_shaped
Beneath each
xylem st rand
there is an arc
of thick_walled
cell
Nearly same as
that of S.lae-
vis
Beneath xylem
strand three
bands of thick_
walled cell are
arranged in a
triangular
shape
{_shaped at
the middle and
basal parts , ω_
shaped at the
apical part
Enclosed and
wide U_shaped
Enclosed and
wide U_shaped
Xylem
st rand
Small , consi st_
ing of about 10
tracheids
Small , consist_
ing of about 12
t racheids
Small , consi st-
ing of less than
10 tracheids
Same as that of
S.laevis
Wide , only 1
cell thick , con-
sisting of 16-
18 t racheids
Wide , 1 - 2
cells thick at
the apical part
and 1-4 cells
thick at the
middle and
basal part ,
consisting of
more than 40
tracheids
Wide , 1 - 2
cells thick ,
consisting of
15 - 40 or
more tracheids
Wide , usually
2 - 3 cells
thick
Transfu-
sion tissue
Present ,
around the
vascular bun-
dle
Present ,
around the
vascular bun-
dle
Present ,
around the
vascular bun-
dle
Present ,
around the
vascular bun-
dle
Present , be-
neath the vas-
cular bundle
Present ,
around the
vascular bun-
dle
Present ,
around the
vascular bun-
dle
Present ,
around the
vascular bun-
dle
Hypoder-
mis
Thickest at the
mid_rib and
thinnest at the
middle depres-
sion
5 - 6 cells
thick and uni-
formly dist rib-
uted
3 cells thick
Same as that of
S.laevis
Thick and uni-
formly distrib-
uted
Thin
Moderately
thick
Moderately
thick
Others
There are small
tracheids at the
inner side of
the xylem
strand which
connect the
two xylem
strand together
There is a
mass of thick_
walled fibrous
cells between
the two xylem
st rands
Author(s) Reed , 1941 Graham , 1935 Koopmans ,
1928
Scott , 1904;
Graham , 1935
Graham , 1935;
Renault , 1879 This paper This paper This paper
Locality
and age
Illinois;C2 Britain;C2 Netherlands;C2 Britain;C2 France;P
1
1
Guizhou ,
China;P22
Shanxi ,
China;P11
Shanxi ,
China;P11
S .shanxiensis sp.nov.and S.taiyuanensis sp.nov.
and even more , up to 3.4 mm wide , in S .guizhouensis
sp.nov.In the Euramerian species , the width of the vas-
cular bundle sheath is 1.2-1.3 mm in S .laevis Koop-
mams and S.sulcata Scott , and is even less in S.cor-
data Reed , S .halifaxensis Graham and the leaves of Si-
gillaria brardi , less than 0.5 mm wide[ 16 ,21 , 22] .
References:
[ 1 ] 　Li X_X.Fossil floras of China Through the Geological Age.
Guangzhou:Guangdong Science and Technology Press ,
1995.[ 2 ] 　Guo Y_T (郭英廷), Tian B_L(田宝霖).Anatomical
characteristics of the stem of Sigillaria cf.brardii from coal
balls in Guizhou Province , China.Acta Bot Sin(植物学
报), 1994 , 36:898-900.(in Chinese with English ab-
stract)[ 3] 　Gao Q (高强), Chang J_L(常江林).A new species of
Lepidodendron from the Late Paleozoic Coal_bearing strata in
Huairen , Shanxi Province.Coal Geol China(中国煤田地
质), 1998 , 10:5-7.(in Chinese with English abstract)
[ 4] 　Hu Y_F(胡雨帆), Zhu J_N(朱家楠), Geng B_Y(耿宝
印).New data of Lepidophloios and Lepidodendron in the
Late Carboniferous—Early Permian from the Tumd Banner ,
Inner Mongolia.Geol Rev(地质论评), 1999 , 45:629-
631.(in Chinese with English abstract)[ 5] 　Hu Y_F , Geng B_Y, Lu Y_S.Late Carboniferous fossils
with seeds and other fossil plants found in Heifangou , Inner
Mongolia , North China.Chenia , 1999 , 6:119-126.[ 6 ] 　Delevoryas T.Anatomy of Sigillaria approximata.Amer J
Bot , 1957 , 44:654-660.
[ 7 ] 　Eggert D A.Petrified Stigmaria of Sigillarian origin from
WANG Shi_Jun et al:Anatomically_preserved Lepidodendralean Plants from Permian Coal Balls of China:Sigillariopsis Scott 111　
North America.Rev Palaeobot Palynol , 1972 , 14:85-99.[ 8 ] 　Lemoigne Y.Etudes analytiques et comparees des structures
internes des Sigillaries.Annales des Sciences Naturelles Bot-
aniques , 1961 , 7:473-578.[ 9 ] 　Lemoigne Y.Sur la structure des Sigillaries.Bull Soc Geol
Fr , 1961 , 2:657-660.
[ 10] 　Schopf J M.Contributions to Pennsylvanian Paleobotany:
Mazocarpon oedipternum sp.nov.and sigillarian relation-
ships.Ill Geol Surv , Rep.Invest , 1941 , 75:1-40.
[ 11] 　Feng B C , Rothwell G W.Microsporangiate cones of Mazo-
carpon bensonii (Lycopsida)from the Upper Pennsylvanian
of the Appalachian Basin.Rev Palaeobot Palynol , 1989 ,
57:289-297.
[ 12] 　Chaloner W G.On the megaspores of Sigillaria.Ann Mag
Nat Hist , 1953 , 12:881-897.[ 13] 　Tian B_L , Wang S_J.Paleozoic coal ball floras in China.Li
X_X.Fossil Floras of China Through the Geological Age.
Guangzhou:Guangdong Science and Technology Press ,
1995.163-180.[ 14] 　Phillips T L , Avin M J , Berggren D.Fossil peat from the
Illinois Basin:a guide to the study of coal balls of Pennsyl-
vanian age.Ill StateGeol Surv , 1976.Education ser.II.1-39.
[ 15] 　Galtier J , Phillips T L.The acetate peel technique.Jones T
P , Rowe N F.Fossil Plants and Spores:Modern Tech-
niques.Bath U K:The Geological Society of London ,
1999.67-70.[ 16] 　Graham R.An anatomical study of the leaves of the Carbon-
iferous arborescent lycopods.Ann Bot , 1935 , 49:587 -
608.
[ 17] 　Chaloner W G.Lycophyta.Boureau E.Traite de Paleobota-
nique.II.Paris:Mason , 1967.668-671.[ 18] 　Wang S_J(王士俊), Guo Y_T(郭英廷), Tian B_L(田宝
霖).Palaeocommunity and its succession of peat swamp of
No.7 Seam in Xishan Coalfield.J Chin Coal Soc(煤炭学
报), 1995 , 20:88 -92.(in Chinese with English ab-
stract)
[ 19] 　Renault M B.Structure comparee de quelques tiges de la
Flora Carbonifere.Nouv Arch Mus DHistorie Naturelle ,
1879 , 2:214-348.
[ 20] 　Scott D H.On the occurrence of Sigillariopsis in the Lower
Coal Measures of Britain.Ann Bot , 1904 ,18:519-521.[ 21] 　Reed F D.Coal flora studies:Lepidodendrales.Bot Gaz ,
1941 , 102(4):663-683.
[ 22] 　Koopmans R G.Researches on the flora of the coal_balls
from the “ Finefrau_Nebenbank” horizon of the Province of
Limburg(The Netherlands).Flora en Fauna van het Ned-
erlandsche Karboon , 1928 , 1:1-53.
中国二叠纪煤核中具解剖构造的鳞木类植物:似封印叶属
王士俊1　田宝霖2　陈贵仁2
(1.中国科学院植物研究所, 北京 100093;2.中国矿业大学 , 北京 100083)
摘要:　报道了产于中国二叠纪煤核中的 3 种具解剖构造的鳞木类叶 , 它们都具双木质部束 , 与似封印叶属(Sigil-
lariopsis Scott)特征一致。与该属已有种进行了对比 ,确认它们为 3 个新种:产于山西太原西山煤田太原组上部 7 号
煤层(早二叠世早期)煤核中的山西似封印叶(Sigillariopsis shanxiensis sp.nov.)和太原似封印叶(S .taiyuanensis sp.
nov.)以及产于贵州水城矿区汪家寨组 1 号煤层(晚二叠世晚期)煤核中的贵州似封印叶(Sigillariopsis guizhouensis sp.
nov.)。根据煤核中共生的鳞木类植物其他器官以及欧美植物区鳞木类植物的研究资料推断 ,它们可能属于封印木
属(Sigillaria Brongniart)的叶。在国外(主要是欧美植物区),封印木属植物主要分布于石炭纪 ,见于二叠纪的封印
木很少。华夏植物区的封印木属过去很少发现 ,其叶和生殖器官均未报道过。本文是首次报道华夏植物区具解剖
构造的封印木属的叶 ,它们的发现不仅丰富了华夏植物区封印木属植物的内容 , 而且对于研究封印木属的演化以
及华夏植物区与欧美植物区鳞木类植物之间的关系也具有重要的意义。
关键词:　华夏植物区;二叠纪;煤核;鳞木类叶;似封印叶属;新种
中图分类号:Q914.82　　　文献标识码:A　　　文章编号:0577-7496(2002)01-0104-09
收稿日期:2001-05-10　接收日期:2001-07-04
基金项目:国家自然科学基金项目(39970056);中国科学院古生物与古人类专项经费项目(QTE_98)。
(责任编辑:崔金钟)
112　 植物学报　Acta Botanica Sinica　Vol.44　No.1　2002