全 文 ：古生物学报 , 44(4):505 - 516(2005 年 10 月)
Acta Palaeontolo gica Sinica , 44(4):505 - 516(Oc t. , 2005)
收稿日期:2005-03-19
*国家自然科学基金项目(No. 49872008)、国家科技部 973项目(2006CB701400)资助课题。
甘肃酒泉盆地下白垩统 Pseudof renelopsis
(掌鳞杉科)的发现及其意义*
邓胜徽1) 　杨小菊2) 　卢远征1)
1)中国石油天然气股份有限公司石油勘探开发研究院 ,北京 100083
2) 中国科学院南京地质古生物研究所 ,南京 210008
　　提要　记述甘肃酒泉盆地早白垩世地层中发现的两种掌鳞杉科植物化石 Pseudo f renelopsis dalatzensis
(Chow et Tsao , 1977) Cao ex Zhou和 P. gansuensis sp. nov. 。其中 ,归入 P. dalatz ensis的标本无论是小枝形态
还是表皮构造都与产于吉林延吉盆地大砬子组的模式标本一致。定为新种的标本小枝较粗 , 可能只有一枚叶;节
间角质层厚 25—30 μm ,表面没有乳突和表皮毛 , 气孔器和表皮细胞均规则成行排列;气孔器圆形或椭圆形 , 较大 ,
整体下陷。副卫细胞通常 5—7 个 , 少数可达 8—9 个。保卫细胞仅部分保存。已发现的 Pseudof renelopsis均分布
于北半球 ,时代限于早白垩世 , 且多为早白垩世晚期 ,其中 P. dalatzensis在模式标本产地与丰富的被子植物共生 ,
应为早白垩世 Aptian 到 Albian 期。该种在酒泉盆地中沟组的出现说明该组的时代最高可到 Aptian 或 Albian 期。
已有资料还表明 , Pseudo f rene lopsis主要生活于有季节变化 、盐度较高或较干旱的环境中 , 这与酒泉盆地早白垩世
沉积物和其他门类的化石所指示的环境特征相符合。
关键词　掌鳞杉科　松柏纲　早白垩世　中国甘肃
PSEUDOFRENELOPSIS(CHEIROLEPIDIACEAE) FROM THE LOWER CRETACEOUS
OF JIUQUAN , GANSU , NORTHWESTERN CHINA
DENG Sheng-Hui1) , YANG Xiao-Ju2) and LU Yuan-Zheng1)
1) Research I nsti tute o f Petroleum Exp loration and Development , Beij ing 100083
2) Nan jing In sti tu te o f Geology and Palaeonto logy , the Chinese Academ y o f Sciences , Nan jing 210008
　　Abstract　Two species of cheiro lepidiaceous conifer s are described based on the m ate rials collected f rom the Low-
er C retaceous Xiagou and Zhonggou fo rmations of Jiuquan Basin , Gansu , northwestern China fo r the first time. Nu-
merous specimens assigned to Pseudof renelopsis dalatz ensis (Chow et T sao , 1977)Cao ex Zhou are morphologically
identical w ith the ho lo type from Jilin , Nor theast China. P. gansuensis sp. nov . is characterized by wide shoo ts , thick
internode cuticles (25 —30μm)having a smoo th oute r surface , near ly isodiametrical epiderm al cells a rranged in row s ,
well-defined stomatal files , la rge and sunken stom ata with 6 —7 subsidiary cells. The geographical and stratig raphical
distributions and the eco log ical characters of Pseudof renelopsis are summarized. The present finds also provide new
evidence fo r the Early Cretaceous age o f the fossil-bearing strata. Sedimentological and cuticle analyses indicate that
these conifers were xerom orphic plants adapted to halophy tic habita ts.
Key words　Cheirolepidiaceae , Coniferop sida , Early Cretaceous , Northwestern Ch ina
1　INTRODUCTION
Pseudof renelopsis is an important Mesozoic chei-
rolepidiaceous conifer which has been studied for more
than 100 years. It was first report ed by Nathorst from
the Lower Cretaceous of Mexico based on the type spe-
cies Pseudof renelopsis f eli xii in 1893. About 80 years
later , new materials were described by Watson (1974)
from the Wealden of England as a new genus Manica.
Later , Watson (1977) abandoned this generic name
and chose Frenelopsisvar ians Fontaine from the Lower
Cretaceous of Texas to replace the Mexican species
Pseudof renelopsis f eli xii as the type species of
Pseudof renelopsis , because no cuticle of the Mexico
materials had been obtained. The related male cone
Classostrobus comptonensis , which yielded Classopol lis
pollen was also described(Alvin et al . , 1978). T he
Chinese materials w ere fi rs tly described by Chow and
T sao in 1977 under the genus Manica , and then t rans-
f erred to Pseudof renelopsis (Cao , 1989;Zhou ,
1995). A ccording to a re-examination by Zhou in
1995 , three species Pseudof renelopsis dalatzensis , P.
papil losa and P. heishanensis , were confi rmed. An-
other conifer described as Suturovagina intermedia
(Chow et al. , 1977;Zhou , 1983)was also referred to
genus Pseudof renelopsis by Watson (1988). Howev-
er , due to the obvious dif ference in leaves , we treated
i t as a dif ferent genus.
A total of 7 species have been described in detail
based on cuticles , among which 1 was found in Eu-
rope , North A frica and No rth America , 1 was reported
from Japan , 3 were report ed f rom China and another 2
were reco rded from North America (Fontaine , 1889;
Watson , 1974 , 1977 , 1988;Watson et al. , 1976;Al-
vin , 1977 , 1983;Alvin et al . , 1978 , 1981;Chow et
al . , 1977;Zhou , 1983 , 1995;Srinivasan , 1995;Sai-
ki , 1999;Yang , 2005). Palaeoecological s tudies (Al-
vin , 1982 , 1983;Upchurch et al. , 1981;Watson ,
1988) show that it is a group of extinct conifers , possi-
bly adapted to rather highly saline and arid condi tions.
In this paper , we repo rt a new occurrence of Pseudo-
f renelopsi s in Northwes t China and describe two spe-
cies , including a new one.
Unlike in northeast ern and eas tern China where
the Early Cretaceous coal-fo rming f loras are widely dis-
t ributed and well st udied , in northwestern China the
Lower Cretaceous plant fossils are rare and relatively
poorly s tudied. Some discoveries do indicate however
that conifers dominated the flora (Chow et al . , 1977).
The plant fossils f rom Jiuquan Basin of Gansu P rovince
(Text-fig. 1) have not been sys tematically described
sofar , although the Lower Cretaceous in this basin has
been studied for palaeontology and st ratigraphy for a
long period(Hou , 1958;Ye , 1984;Ma , 1993;Ye et
al. , 1990). Recently , the authors carried out a re-
search project on the s tratig raphical correlation be-
tw een the outcrops and the drill cores of the basin.
Some plant fossils w ere collect ed from both the out-
crops and drill cores. Support ed by the National Natu-
ral Science Foundation of China , t he plant fossils have
been syst ematically studied , and a foliage assemblage
of about 15 species was recog nized. T he assemblage is
clearly dominated by conifers , including Pseud-
of renelopsis dalatzensis (Chow et T sao) Cao ex Zhou
and P. gansuensi s sp. nov. described herein.
2　STRATIGRAPHY , MATERIALS AND METHOD
The Mesozoic st rata in Jiuquan Basin of Gansu
(Text-fig. 1) are ex tensively developed and w ell ex-
posed in the margins of the basin. T he s tratig raphic se-
quence of the Lower Cretaceous , named the Chijinbao
G roup previously , is divided into the Chijinbao Forma-
tion , Xiagou Formation and Zhonggou Formation (Ye
et al . , 1990). T he Chijinbao F ormation is dominated
by fluvial sediment s including conglomerat es , gravel
conglomerates , pebbly sandst ones and sandstones in
the lower member. Lacust rine sandstones and mud-
st ones with intervening units of thin marls occur in the
middle and upper members , rich in hydrobios including
bivalves , gast ropods , ost racodes , charophytes and a-
quatic insect s , as well as some fossil wood. The Xia-
gou Formation is divided int o tw o members , the lower
one charact erized by conglomerates and sandstones ,
and the upper one dominated by dolomitic limestones
that are rich in aquatic organi sms. T he Zhonggou For-
mation consists of red mudstones and argillaceous sand-
st ones at the base and finely laminated
dolomitic limestones , mudstones in the mid-upper
506 古　生　物　学　报 第 44 卷　　　　　　
Text-fi g. 1　Map show ing the localit y of the fossil s
part , with numerous insects , ost racodes , fishes , con-
chost racans , charophy tes , plant s and some bird feath-
ers.
T he Chijinbao Formation is usually correlat ed with
the Jehol G roup of Northeast China , because it con-
tains the representatives of t he Jehol Biota , such as
Ephemeropsis tr isetalis and Eosesthr ia. Its age is still
a matt er of argument , as it is f or the Jehol G roup , al-
though more and more authors now believe it to be
Early Cretaceous (Ye et al . , 1990;Deng et al. ,
2003). Evidence from palaeontology supports t he Early
Cretaceous age of the Xiagou and Zhonggou fo rma-
tions;however , the exact assignment s are s till prob-
lematical.
T he plant fossils described here were collected
from the laminated muds tone of Zhonggou Fo rmation
of the Hanxia section in the southwest of t he basin and
the Xiagou Formation of Well-Chang 101 in eastern ar-
ea of the basin(Tex t-fig. 1). The fossils are compres-
sions wi th cuticles preserved.
T he specimens w ere photographed using optical
and digital cameras fi rs tly and then some small pieces
of t he rock containing the fossils were removed to small
plastic cups. T hese small pieces were t reated with
hydrof luoric acid for 24 hours to separate the f ossil tis-
sues from the mat rix. T he cuticle preparations w ere
obtained by macerating the fossil tissues in concent rat-
ed nit ric acid , follow ed by ammonia or sodium potassi-
um hydroxide. T he cuticles w ere examined under a
light microscope and scanning elect ronic microscope.
For SEM studies , the cuticles w ere fixed to the speci-
men stubs and sput ter coated with gold.
A ll specimens , slides , and SEM stubs studied in
the present paper are housed in the Research Ins ti tute
of Pet roleum Exploration and Development (Beijing).
3　SYSTEMATIC PALAEONTOLOGY
Order　Coniferopsida
Family　Cheirolepidiaceae Takhtajan , 1963
Genus 　Pseudof renelopsis (Nathorst , 1893) emend.
Watson , 1977
Pseudof renelopsis dalatzensis (Chow et Tsao , 1977)
Cao ex Zhou , 1995
(Pl. I , fi gs. 3—11)
　　Basionym　Manica(Manica) dalatzensis Chow et
Tsao , 1977 , p. 171 , pl. III , figs. 5—11;pl. IV , fig.
13;tex t-fig. 3.
　　Synonym　Pseudof renelopsis dalatzensis (Chow
et T sao , 1977) Cao , 1989 , p. 437 , 442 (name only);
Pseudo f renelopsis dalatzensis (Chow et T sao , 1977)
Cao ex Zhou , 1995 , p. 421 , pl. I , figs. 3—5 , 10;pl.
II , figs. 1—8.
　　Description 　There are many specimens of this
507　第 4 期 邓胜徽等:甘肃酒泉盆地下白垩统 Pseudof renelopsis(掌鳞杉科)的发现及其意义　　　　
species in the present collection. T he shoots are about
4 mm wide , and the longes t one is up to 30 mm long
without being branched. T he internodes are quite regu-
lar in length , of 4—5 mm (Pl. I , figs. 3—5), with
fine longit udinal lines on the surface. T he nodes are
flat and s light ly contracted (Pl. I , fig. 4). The f ree
part of leaves are very small , broadly t riangular in
shape , only about 1mm high , with acute apex , one leaf
per node (Pl. I , figs. 3 , 4).
　　The abaxial cuticles of the leaves are usually thin ,
and with a smooth outer surface. No hairs are seen
(Pl. I , fig. 10 , arrow s). The epidermal cells are com-
monly square in shape or irregular in form , and are ar-
ranged in files (Pl. I , fig. 6).
　　The internode cuticles are quit e thick. Stomata are
arranged in distinct files on the outer surface (Pl. I ,
fig. 8), with intervals of 120 μm between two files of
stomatal pit s , separated by about 4 epidermal cells.
T here are about 8—9 stomatal files per mm width.
T he intervals betw een the two adjacent s tomata within
the st omatal row are 50—100 μm (separated by 2—6
cells). The stomatal apparatus are typically sub-round-
ed or elliptical , 40—55 μm in diameter. T he subsidiary
cells , 5 o r 6 in numbers , are s trongly papillate. T he
stomatal pit mouths are usually closed or slight ly
opened (Pl. I , figs. 8 , 9). T he ordinary epidermal
cells are usually papillate , and the papillae are typically
ellip tical or rounded , up to 15 m in diameter.
　　Stomata on the inner surface of t he internode cuti-
cle are arranged in well-defined files (Pl. I , figs. 7 ,
10) which are about 60—80μm wide and with intervals
of 30—50 μm wide , separated by 3 , 4 epidermal cells ,
approximately 8—9 s tomatal files per mm wide. T he
intervals between the two adjacent stomata within the
stomatal file are irregular , ranging between 20 μm and
100 μm , separated by 1—6 ordinary epidermal cells.
T he stomatal apparatus are normally ellip tical , about
50 μm long and 40 μm wide , and are transversally or
slightly obliquely oriented , with 5—6 subsidiary cells.
T he st omatal apertures are elliptical. Guard cells are
deeply sunken. T he ordinary epidermal cells between
stomatal files are usually unclear in fo rm due t o the
presence of t he hypodermis. The periclinal walls are
smooth (Pl. I , fig. 7).
　　The density of stomata decreases downwards and
are absent at the base of the internode (Pl. I , fig. 11).
The ordinary epidermal cells at the base are more regu-
lar (Pl. I , fig. 11) than in the middle of the internode ,
are usually square or rectangular , and arranged in well-
defined files.
　　Comparison and Discussion 　The present speci-
mens are easily distinguished f rom P. var ians , P.
parceramosa , P. nathorstiana , P. glabra , and P.
heishanensis by the presence of thick papillae on the in-
ternode surface and absence of hairs at the leaf mar-
gins. The described specimens are morphologically
similar to P. papi llosa and P. dalatzensis. These two
know n species resemble each other not only in the na-
ture of the shoots but also in some aspects of cuticle ,
such as having densely arranged papillae on the surface
of internode cuticles (Chow et al . , 1977 , Zhou ,
1995). H ow ever , P. papi llosa has sparser and smal-
ler papillae and its stomata on the surface of internode
cuticle have stellate pit mouths(Zhou , 1995). Addi-
tionally , it has numerous hairs on the outer surface and
the margins of leaves , while P. dalatzensis has larger
papillae and no hairs on the outer surfaces and margins
of leaves (Zhou , 1995), and in these respects is identi-
cal with the current materials. T herefore , the present
materials should be referred to P. dalatzensi s.
Pseudof renelopsis gansuensis sp. nov.
(Pl. I , f igs. 1 , 2;Pl. II , fi gs. 1—9;Text-fi g. 2)
　　Holotype　Chang 101-3926-01.
　　Repository　Research Ins ti tute of Pet roleum Ex-
ploration and Development , Beijing.
　　Etymology　The specific epit het is aft er the name
of the fossil locality.
　　Locality　Jiuquan of Gansu Province , China.
　　Stratigraphy　Zhonggou Formation.
　　Age　Early Cretaceous.
　　Diagnosis　Shoot thick , up to 56 mm long without
being branched. Internodes 7—11 mm long , cont rac-
ting at the node , with fine longitudinal ridges and
g rooves on the surface. Int ernode cuticles thick ,
smooth on the out er surface. A t the base of the inter-
node , the epidermal cells mainly rectangular o r square ,
arranged in row s , st omata sparsely dist ributed. In the
middle part of t he internode , epidermal cells typically
square and nearly isodiamet rical , arranged in distinct
508 古　生　物　学　报 第 44 卷　　　　　　
row s. Stomata in well-defined files , separat ed by 2—4
cells , about 7—8 stomatal files per mm wide. Stomata
rounded or sub-rounded , sunken , 90—100μm in diam-
eter;s tomatal apertures elliptical. Subsidiary cells typ-
ically 6—7 in number and occasionally 8—9 , having no
papillae. T he epidermal cells usually square or nearly
isodiamet rical , arranged in rows.
　　Description　There is one specimen in the present
collection. T he shoot is 56 mm long and 9 mm wide ,
without being branched. The internodes are 7—11 mm
long , and contracting at t he node(Pl. I , fig. 1), with
very fine longitudinal ridges and grooves (P l. I , fig.
2). T he leaf i s imperfect ly preserved (Pl. I , fig. 2 ,
arrow).
　　The internode cuticles are thick , about 25—30 μm
(Pl. II , fig. 8). T he outer surface of cuticle is smooth
at t he base , without papillae and hairs (P l. II. fig.
1). T he epidermal cells are mainly rectangular or
square at the base of the internode (Pl. II , fig. 6),
while typically square and nearly isodiametrical in the
middle part of the internode(Pl. II , figs. 1—3). T hey
are about 20—50 μm long and 25—40 μm wide , ar-
ranged in distinct rows (Pl. II , figs. 1 , 2).
　　Stomata are sparsely dis tributed or even absent at
the base (Pl. II , fig. 6), densely and w ell-definedly
arranged in files in the middle part of the internode
(Pl. II , figs. 1 , 4 , 9). T he int ervals between two ad-
jacent s tomata are about 100 μm at the base and be-
come closer upw ards. The intervals between tw o adja-
cent st omatal files are 30—50 μm wide , separated by
2—4 cells. T here are 7—8 files per mm wide. T he
mouths of t he stomatal pits are rounded , elliptical or
square or rectangular (Pl. II , figs. 1—3). The stoma-
ta within a file are closely arranged in the middle part
of the internodes , t he adjacent stomata are separated
by 0—3 epidermal cells , with intervals of 0—50 μm.
No common subsidiary cells are found(Pl. II , fig. 7).
T he stomata are rounded or sub-rounded , 90—100 μm
in diameter and randomly oriented. The st omata are
obviously sunken(Pl. II , figs , 4 , 7—9;Text-fig. 2),
and are randomly oriented. Guard cells are sunken ,
and thickened to f orm a parentheses-like part of about
30—50 μm long and 3—6 μm wide in some cases (Pl.
II , figs. 7 , 9). T he subsidiary cells , typically 6-7 in
number and occasionally up to 8—9 , have no papillae
(Pl. II , figs. 4—9). T he epidermal cells at t he base of
t he internode are approximately square or nearly isodia-
met rical and arranged in files (Pl. II , fig. 6), while
the out lines of t he ordinary epidermal cells between
s tomatal files in the middle of the internode are usually
unclear on the inner surface of the cuticle due to the
presence of hypodermis. T he periclinal walls are
smooth (Pl. II , figs. 4 , 7 , 9).
Text-f ig. 2　Vertical sect ion showing the s t ru cture of a s toma
(A draw ing based on plate II , fig. 8)
　　Comparison and discussion　The present material
is referred to Pseudof renelopsis mainly based on the
characters of its cuticles. The genera Pseudof renelop-
sis and Frenelopsis are morphologically close to each
other , even the epidermal st ructures. Generally speak-
ing , Frenelopsi s has three leaves per node , but some
species , such as Frenelopsis tei xeirae , have two leaves
per node (Alvin et al. , 1978);while Pseudof renelop-
sis is believed to have just a single leaf per node , but
some shoots of P. nathorstiana with two occasionally ,
according to the description (Srinivasan , 1995). T here-
fore , Srinivasan (1995) pointed out that the difference
in phyllotaxy may no longer provide suf ficient grounds
f or generic separation. However , to distinguish these
two genera the epidermal features are helpful. The
subsidiary cells of Pseudof renelopsis are more numer-
ous than in Frenelopsis. A ccording to the dat a f rom 15
species of Frenelopsis by Gomez et al . (2002), the
numbers of subsidiary cells of Frenelopsis are 4—6 ,
and mos tly 4—5 , very few up to 6 , while in the 7
know n species of Pseudo f renelopsis (Table I) the sub-
sidiary cells are 4—9 and typically 5—6 in number.
Some species such as P. var ians and P. glabra have
up to 9 (Watson , 1988;Saiki , 1999). A lvin et al .
(1978) suggested that a small diff erence between the
two genera is that the stomata in Frenelopsis typically
509　第 4 期 邓胜徽等:甘肃酒泉盆地下白垩统 Pseudof renelopsis(掌鳞杉科)的发现及其意义　　　　
Table I　Comparison of the species of Pseudof renelopsis
Characters /Species P . var ians P. parceramosa P. nathorst iana P. papi l losa P. d alatzensis P. heishanensis P. gansuensis P. g labra
Internode length (mm) 1. 5—17 1—11 1—2. 9 5—11 4—10 5—6 7—11 4—8
Internode width (mm) 3—7 1 and upwards 1—2 3—7. 5 3—6. 5 2. 5—4 9 4
Leaf number per node 1 1 1—2 1 1 1 1 1
Maximum length of
f ree leaf(mm) 1. 5 2 1 1. 5 2 2 — 1. 5
Leaf margin
Hai rs up to
60μm
Hairs up to
80μm
Hairs up to
145μm
H airs up to
80μm Without hair — —
Hair up
to 40μm
Su rface of adaxial
cuticle of leaf
Hai ry Hai ry Hairy H airy Without hair Without hair — Without
hair
Internode cuticle
thickness (μm) 50—100 30 10 5—7. 5 5—25 3—5 25—30 3
Su rface of abaxial
cuticle of leaf
With hai rs ,
up to 80μm
Without hai r
and papillae
No mentioned With papillae With papillae
Without pa-
pi llae
Withou t pa-
pillae
S tomatal arrangement
Scattered in
“ closed ”
f orm , row s in
“ open” form
Well-defined
rows
Ill-def ined
rows
Longitudinal
rows usually
w ith scat tered
stomata ,
w ell-defined
rows
Well-defined
rows
Well-defined
rows
Well-defined
row s
Well de-
fined
rows
S tomatal row s per mm 8—10 6—10 4—9 6—10 8—10 7—8 7—9
Diameter of stomatal
apparatu s(μm) 70—100 50—80
50—73 ×
54—62 40—92. 5 55—95 62. 5—100 90—100 8—120
Number of subsidiary
cells
Usually 5—
8 , rarely 4 or
7
Usually 5—
6 , rarely 4 or
9
Usually 5—
6 , rarely 4 or
Usually 5—
6 , rarely 4 , 7
or 8
Usual ly 5—6 Usually 5—
6 , rarely 7
Usually 6—
7 , rarely 8—
9
6—8
Orientation of stomatal
aperture
Random Random Random Random Random Random Random
Subsidiary cell s
With elongate
papillae
Without pa-
pillae to
st rongly pap-
illate
With papillae With papillae
St rongly bul-
ging proxi-
mally
Bulging proxi-
mally surface
f lush w ith
that of epider-
mal cells
Withou t pa-
pillae
Without
papillae
Age Aptian-Albian Berriasian-Al-
bian
Albian
Lower Creta-
ceous
Aptian-Albian
Lower Creta-
ceous ( pre-
Albian)
Aptian-Albi-
an
Albian
Distribution
North Ameri-
ca
North Ameri-
ca , Europe
North Ameri-
ca
China China China China Japan
References Watson , 1977
Alvin , 1977;
Alvin et al . ,
1978; Wat-
son , 1977
Srinivasan ,
1995
Chow and
Tsao , 1977;
Zhou and
Cao , 1979;
Zhou , 1995
Chow and
Tsao , 1977;
Zhou , 1995
Zhou , 1995 The present
paper
Saiki ,
1999
510 古　生　物　学　报 第 44 卷　　　　　　
have papillae ex tending f rom the sides of the pit , while
in Pseudof renelopsis the papillae , if present , are at
the mouth of the pit. A s the leaves of the present spec-
imen w ere incompletely preserved , it is estimated that
there is only one single leaf per node. But the numbers
of subsidiary cell are typically 6—7 , and in some cases
up to 8 or 9 , indicating undoubtedly Pseudof renelopsis
type. Consequent ly , t he present material should be re-
f erred to Pseudof renelopsis.
Text-fig. 3　Dis t rib ution of Pseudo fr enelopsi s in the w orld
1. En gland , 2. Po rtugal , 3. Poland , 4. Ghana , 5. Su dan , 6. Gan su of China , 7. Qing hai of China , 8. Ningxia of
C hina , 9. Jiangsu of Chin a , 10. Zh ejiang of China , 11. Fujian of China , 12. Jiangxi of Chin a , 13. Hubei of C hina ,
14. Jilin of China , 15. Hokk aido of Japan , 16. M exico , 17. T exas of th e United S tates , 18. M aryland and Virginia
of the Uni ted S tates
　　As mentioned above , among the known species of
Pseudof renelopsis (Table I), Pseudof renelopsis vari-
ans , P. papi llosa and P. dalatzensis are very dif ferent
from the described specimen in having many papillae on
the internode cuticles. P. nathorstiana is characterized
by its mostly ill-defined row s of stomata. The most
similar species are P. parceramosa , P. heishanensis
and P. glabra , which are characterized by their
smooth internode cuticles. Among these three species ,
P. parceramosa , with similar t hickness of internode
cuticles , dif fers in having thickened Florin rings in the
stomata , papillae inside the stomatal pit and unclear
outline of the epidermal cells (Watson , 1977). Further
mo re , this species has very long hairs on the outer sur-
f ace and margins of it s leaves (Watson , 1977;Alvin ,
1977). P. heishanensis has the thinnes t internode cuti-
cle (3—5 μm) of all the know n species of Pseud-
of renelopsis , and is characterized by thinner shoot s
(5—6 mm thick) and st ellat e pit mouths (Zhou ,
1995). P. glabra differs f rom the new species in it s
much longer internodes , thickened ring of the s tomata
and st ellate pit mouth(Saiki , 1999). Additionally , the
present mat erial has the largest stomata among the all
know n species of Pseudo f renelopsis. T herefore , we
heret o propose a new specific name fo r the described
material.
4 　DISTRIBUTION AND ECOLOGY OF PSEUDO-
FRENELOPSIS
　　Pseudof renelopsis is a genus of worldwide dist ri-
bution in the Early Cret aceous(Text-fig. 3). P. var i-
ans has been found from Mexico (Nathorst , 1893),
Texas of the United States (Fontaine , 1893;Daghlian
et al . , 1977;Watson , 1977) and England (Watson ,
1974 , 1977). T he American f ossils f rom the Pot omac
Flora of Maryland and Virginia , which were firstly as-
signed to Frenelopsis (Fontaine , 1889) were also re-
moved to Pseudof renelopsi s by Watson (1977). P.
parceramosa is widely dist ribut ed in the Wealden flora
of England (Watson , 1977;Alvin et al . , 1981) and
Portugal (Alvin , 1977). It has also been reported f rom
Ghana , Sudan(Watson , 1977 , 1988)and Poland. Be-
511　第 4 期 邓胜徽等:甘肃酒泉盆地下白垩统 Pseudof renelopsis(掌鳞杉科)的发现及其意义　　　　
sides foliage , some w oods f rom Is le of Wight of Eng-
land have been at tributed to this species (Alvin et al. ,
1981);and some plant debris from the English Weal-
den s tudied by Oldham (1976) was also considered to
relate to this species(Watson , 1977). P. nathorstiana
is another species in the American Potomac f lora pro-
posed by Srinivasan (1995), but only one locality has
been know n so far. Recently , Saiki (1999)described a
new species P. glabra f rom the A lbian of Hokkaido ,
Japan.
　　Including the new one described here , t here are 4
species of Pseudof renelopsis in China. Among these
four species , P. papi llosa has the widest dist ribution ,
which was report ed f rom N ingxia , Qinghai , Jiangsu ,
Zhejiang , Fujian and Jiangxi provinces (Text-fig. 3),
while P. heishanensis has only been found from Daye ,
Hubei , and P. dalatzensis has been recorded in Yanji
Basin , Jilin and the present fossil site.
　　The st ratigraphical dist ribution of Pseudo-
f renelopsi s is res tricted t o the Lower Cretaceous. P.
parceramosa is possibly the older occurrence in the
Berriasian. Mos t other species were found in Aptian to
Albian sediment s (Table I). P. dalatzensis is associ-
ated with many other plant s in the type locality. These
plant s include Otozamites sp. , Zamiophy llum buchia-
num , Cupressinocladus elegans , Elatides curvi f olia ,
Brachy phyl lum obesum , B. parceramosum , Suturo-
vagina intermedia , Frenelopsi s elegans , Elatocladus
sp. and many angiosperm s Saliciphyl lum longi foli-
um , Sterculophyl lum eleganum , Ficophy llum sp. ,
Ranumculophy lum pinnatisectum , Clemati tes lanceo-
latus , Archimagnolia rostratosty lisa , Eucommioi tes
or ientalis , Al loephedra xing xuei , Yanj iphy llum el-
lip ticum , Sapindopsis cf. var iabi lis , S. obtusi f olia
and Rogersia angusti f o lia (Zhou et al . , 1980;Tao et
al . , 2000). V arious angiosperms indicate an Aptian-
Albian age (Tao et al . , 2000). Furthermo re , P.
dalatzensi s has not been f ound f rom the beds w hich are
of Barremian and earlier ages in the adjacent areas.
Consequent ly , P. dalatzensis is a species that possibly
lived during the Aptian-Albian.
　　Pseudof renelopsis remains were always found
from red colored sediments which suggested arid sedi-
ment envi ronments. Pseudof renelopsis parceramosa
from the Potomac F lora indicates coas tal , tidally-inf lu-
enced habi tats and may have adapted t o a range of envi-
ronment s of different salinity or even to non-saline hab-
itats(Upchurch et al . , 1981). Oldham (1976)believed
that t he cheirolepidiaceous conifers of English Wealden
f lora probably lived in river margin swamps. A lvin
(1983) and A lvin et al . (1981) pointed out that P.
parceramosa is likely to have been a f orest t ree than
scrubby or sub-herbaceous plant , and it is t herefore
conceivable that t he Pseudof renelopsis fores ts inhabi t-
ed soils , the water availability of which was variable.
The irregular growth rings of the wood may well be re-
lat ed to periods of drought when g row th was interrupt-
ed. P. varians f rom Texas of the United States seems
t o have been t ruely halophy tic , having inhabi ted salt
marshes(Upchurch et al. , 1981). It was also consid-
ered as a plant adapted to arid environment s by having
cuticle well over 100μm thick (Watson , 1977;Watson
et al . , 1984). Its associated conifer Glenrosa texensis
show s further morphological and cuticle st ructural ad-
aptations to aridity by reduction to a scaleleaf. Papillae
on the epidermal cells would have contained the spread
of t rapped droplets of moisture on the surface , thus re-
ducing the surface area available for evaporation (Wat-
son et al . , 1984). T he geological evidence indicated
that the sediments represent an environment character-
ized by high salinity and high rates of evaporation
(Daghlian et al. , 1977). T herefore , most authors be-
lieve that Pseudof renelopsis was adapted to high salin-
ity and arid environment s.
　　The specimens discussed here w ere preserved in
dolomitic mudstone and mudst one. T he dolomitic mud-
st one was an evaporite which was formed in an environ-
ment of high salinity and high rates of evaporation.
The associated ost racodes are dominated by Cypridea ,
which is adapted to brackish water habitat s. T he pres-
ent plant fossils are characterized by much reduced leaf
with rather thick cuticles. T heir stomata are usually
sunken. The associated plant assemblage with a low
diversity , dominated by scale leaf conif ers , represent s
a semi-arid or arid climate. The distinct grow th rings
in the associated wood fossils indicate a seasonal cli-
mate which w as similar to t hat of English Wealden
(Alvin et al. , 1981). The plant assemblage is marked-
ly dif ferent f rom the typical coal-forming f loras that in-
dicate a warm and humid climate in Northeastern Chi-
512 古　生　物　学　报 第 44 卷　　　　　　
na. Consequent ly , the present Pseudof renelopsis w as
possibly a plant adapted to seasonal and arid or semi-
arid habit at s.
5　DISCUSSIONON STRATIGRAPHY
　　The fossils described herein came from two hori-
zons , the Xiagou Formation and the Zhonggou Fo rma-
tion. T he Xiagou Fo rmation f rom which the new spe-
cies Pseudof renelopsis gansuensis was collected , yields
many microfossils , especially charophyt es , ost racodes ,
and spore and pollen. It is usually considered as early-
middle Cretaceous in age by different authors , about
Hautervian-Barremian. However , t he age of Zhonggou
Formation , which yields Pseudof renelopsi s dalatzen-
sis , is considered to be Barremian based upon charo-
phy tes but it may also be Aptian in age based on paly-
nology. The main evidence is t hat some early angio-
sperm pollenmorphs such as Clavat ipollenites , Pol-
ypor ites and Tricolpi tes are present (Lu et al . , 2002).
As discussed above , Pseudof renelopsis dalatzensis is
associated with many angiosperms in the type locality
and was regarded as Aptian-Albian in age (Tao et al. ,
2000). The appearance of P. dalatzensis in the Zhong-
gou F ormation supports the age interpretation by paly-
nologists (Lu et al . , 2002).
　　Acknowledgement　The authors are especially in-
debted to P rof. Z-Y Zhou of Nanjing Ins ti tute of Geol-
ogy and Palaeontology not only f or his guidance and en-
couragement but also his valuable work on the present
study and his thorough review of the manuscript. We
also express appreciation to D r. Bernard Gomez of t he
University of Rennes and Dr. Joan Watson of the Uni-
versity of Manchester fo r the benefi ted discussion on
the genera Pseudof renelopsis and Frenelopsis and
Prof. Jane Francis of the Universi ty of Leeds for her
linguistic revision of t he manuscript. The work w as
supported by Senior Visi ting Scholarship of Nanjing In-
stitu te of Geology and Palaeont ology , t he Chinese A-
cademy of Sciences. T he paper was finally finished in
the University of Leeds of the United K ingdom when
the autho r (Shenghui Deng) w as a visiting scholar at
the Universi ty f unded by the China Scholarship Coun-
cil.
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Explanation of Plates
Plate I
1— 2. Pseudo f renelopsis gansuensis sp. nov.
1. Vegetative shoot. H oloty pe , Chang-101-3926-01.
2. Enlargement of 1 , show ing app ressed subacute apex of leave
(arrow) and fine paral lel ridges.
3— 11. Pseudo f renelopsis dalazensis (Chow et Tsao) Cao ex Zhou
3—5. Vegetat ive shoots , HX-98-2002-04 , HX-95-2002-08 ,
HX-95-2002-02 , show ing apex of a leave and parallel ridges and
grooves wh ich become convergent tow ards the leaves apex.
6—11. Scanning micrographs of cu ticles. 6. Inner su rface view
of adaxial cut icle , showing the epidermal cells and tw o stomata
(arrow s);7. Inner surface view of internode cu ticle , show ing
the stomata and longitudinal ordinary epiderm al cells;8. Outer
surface view of internode cut icle , show ing s tomatal and non-
stomatal files with papil la of each epiderm al cell;9. Ou ter sur-
face view of a stoma w ith 6 subsidiary cells s trongly bulging
proximally forming a Florin ring;10. Inner su rface view of in-
ternode cu ticle and ou ter surface of abaxial leaf (arrow s),
show ing the regular arranged stomatal and non-stomatal row s;
11. Inner surface view of internode cuticle at the basal part of
internode , show ing few irregularly ar ranged stomata (arrow s)
and regular longitudinal cel ls.
Plate II
1—9. Pseudo f renelopsis gansuensis sp. nov.
Scanning micrographs of cu ticles.
1 , 2. Outer surface view of internode cuticles , showing smooth
su rface , regularly arranged s tomata and more sunken s tomatal
pit . 3. Outer surface view of a stomatal pi t. 4. Inner su rface
view of internode cu ticle , showing well-def ined stomatal fi les
which are normally uniseriate;5. Inner surface view of inter-
node cu ticle , show ing margin epidermal cells and tw o stom ata;
6. Inner su rface view of in ternode cu ticle from the base part of
internode , show ing irregularly arranged s tom ata and epidermal
cells;7. Inner su rface view of in ternode cut icle , showing tw o
s tomata arranged in biseriate;8. Vert ical section and inner view
of a stom a(sp:stomatal pit;sc:subsidiary cel l;ec:epidermal
cell);9. Inner surface view of internode cu ticle , show ing well-
def ined stom atal and non-stomatal row s.
514 古　生　物　学　报 第 44 卷