全 文 ：Paleocene Wuyun Flora in Northeast China:Ulmus furcinervis of Ulmaceae
FENG Guang_Ping1 , Albert G.ABLAEV2 , WANG Yu_Fei1 , LI Cheng_Sen1＊
(1.Inst itute of Botany , The Chinese Academy of Sciences , Beijing 100093 , China;
2.Pacific Oceanological Institute , Russian Academy of Sciences , Vladivostok 600041, Russia)
Abstract:　Ulmus furcinervis(Borsuk)Ablaev is first recognized and described in the Paleocene Wuyun flora
from Heilongjiang Province , China.The combination of key characteristics for identification of these leaf speci-
mens is that they have craspedodromous secondary veins forked near the leaf marginal , tertiary veins percurrent
and marginal teeth convex.Specimens formerly described as leaves of Planera cf.microphylla auct.nonNew-
berry and Sorbaria wuyunensis Tao et Xiong in Wuyun flora are reexamined and assigned to U.furcinervis.
Key words:　Ulmus furcinervis;Ulmaceae;Paleocene
　　The genus Ulmus (Ulmaceae)consists of about 30
species in the North Hemisphere.There are 25 species of
the genus living in China (Chun and Huang , 1998).
Fossil pollen records of the genus can be traced back to
Campanian of northeastern USA (Wolfe , 1976).Fossil
records of foliage and reproductive organ of Ulmus can be
traced back to Paleocene of northwestern USA (Brown ,
1962).The reliable records of Ulmus verified by both
fruits and foliage are not latter than the early Eocene(Manchester , 1987;Manchester , 1989).The early
records of Ulmus in China were documented in the sedi-
ments from Late Eocene of Shaanxi Province(Tao , 1965)
and Eocene of Guizhou Province of China (Zhang ,
1983).In this paper , we reported the earliest record of
Ulmus in China , collected from the sediments of Wuyun
Formation(Paleocene)at Wuyun Coal Mine (49°14′26″
N , 129°28′00″E), Jiayin County , Heilongjiang Province(Feng et al , 2000;Liu , 1983;1990;Luo et al , 1983;
Xiong , 1986).Pollen assemblage of Wuyun Formation is
dominated by angiosperm pollen grains in which tricolpate
pollen grains relating to Betulaceae , Ulmaceae and Jug-
landaceae are superior to others (Liu , 1983).Pollen
grains assigned to Ulmipollenites and Ulmoideipites are
abundant(Liu , 1983).
The leaf specimens assigned to Ulmus furcinervis are
characterized by the dentate margin with broadly triangular
simple or compound teeth , the stout and frequently forked
secondary veins , and the opposite percurrent tertiary
veins.Specimens formerly assigned to Planera cf.micro-
phylla and Sobaria wuyungensis (Tao and Xiong , 1986;
Tao , 2000)from the same locality are also assigned to
U.furcinervis considering their close similarities to the
species.
The terminology of leaf architecture used in this
study follows that of Hickey (1979).
1　Systematics
Family:Ulmaceae
Genus:Ulmus L.1754
Species:U.furcinervis(Borsuk)Ablaev 1974(Figs.1-7)
Synonymia:
Zelkova furcinervis Borsuk:Borsuk , 1952 , P.24 ,
Pl.3 , Fig.7;Pl.4 , Figs.1-11
Zelkova ungeri (Ettingshausen)Goeppert var.du-
plicatodentata Borsuk:Borsuk , 1956 , P.43 , Pl.9 ,
Fig.5
Ulmus furcinervis(Borsuk)Ablaev:Ablaev , 1974 ,
P.102 , Pl.14 , Figs.4 -5;Pl.15 , Figs.2 -10 ,
textFig.23;Sycheva , 1977 , P.28 , Pl.10 , Fig.1;
Ablaev , Iljinskaja , 1982 , P.14 , Pl.6 , Figs.7-11 ,
textFig.6;Kamaeva , 1982 , P.74 , Pl.9 , Fig.3;Ka-
maeva , 1990 , P.42 , Pl.5 , Fig.3v , 4v;Pl.6 , Fig.
7a;Pl.9 , Fig.4;Pl.10 , Figs.1-3;Pl.11 , Figs.6 ,
7 , 8b;Glandekov et al , 1997 , P.147 , Figs.1 -3 ,
Fig.11;Kodrul , 1999 , P.108 , Fig.28 , sine descr.
Ulmus ex gr.furcinervis(Borsuk)Ablaev:Ablaev ,
1985 , P.39 , Pl.3 , Fig.3;Pl.4 , Fig.3;Pl.16 , Fig.4
Planera cf.microphylla auct.non Newberry:Tao
and Xiong , 1986 , P.125 , Pl.5 , Fig.5;Tao , 2000 , P.
145 , Pl.8 , Fig.5
Sorbaria wuyunensis Tao et Xiong:Tao and Xiong ,
1986 , P.127 , Pl.6 , Figs.5 , 6;Tao , 2000 , P.127 ,
Pl.8 , Figs.5 , 6;Tao , 2000 , P.167 , Pl.9 , Figs.5 , 6
Specimens:52141 , 52144 , 52146 , 52168 , 52240 ,
52241 , 52262 , 52278 , 52310 , 52607 , 52611 , 52614 ,
52624 , 52707 , 52714 , 52716 , 52717 , 52718 , 52719 ,
52739 , CBP55000 , CBP55001
Repository:National Museum of Plant History of China ,
Institute of Botany , The Chinese Academy of Sciences ,
Xiangshan , Beijing
Locality:Wuyun Coal Mine , Jiayin County , Hei-
longjiang Province
Stratigraphy:Wuyun Formation
Age:Paleocene
Description:
Leaves symmetrical to slightly asymmetrical (Figs.1-7).Ovate to narrow ovate in outline.Lamina 35-70
Received:2002-10-31　Accepted:2002-12-23
Supported by the National Natural Science Foundation of China(30130030 , 30070056)and Lu_Jia_Xi grant and The Chinese Academy of S ciences(KSCXZ_SW_
108).
＊Author for correspondence.E_mail:.
植　物　学　报　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　

Acta Botanica Sinica　2003 , 45(2):146-151
(110)mm long , 12-35(50)mm wide , and the ratio of
length to width 2.0-3.0.Petiole stout , usually 2-4
mm , occasionally up to 9 mm in the large leaves.Leaf
apex acuminate(Figs.1-5).Leaf base obtuse to shallow
cordate with an obtuse base angle (Figs.1 -3 , 5 , 6 ,
10).Margin dentate or serrate occasionally with simple or
compound teeth , teeth non_glandular with a simple apex ,
broad triangular in shape with convex to acuminate apical
side and convex basal side , sinus right angle , regularly
spaced.Tooth apex broad triangular with convex to
acuminate apical side and convex basal side (Figs.8 ,
10).Small additional teeth on the abmedial side and the
sinus ended by the weak tertiary veins divergent from the
basal side of secondary veins at marginal area (Fig.10).
Primary vein of the teeth central and direct , the accessory
veins of the teeth looped.Texture chartaceous , sometimes
crimpled with veins deeply embedded in the lamina(Figs.1 , 3 , 8 , 9 , 10).Venation pinnate with a stout
midrib(Figs.1-7);secondary veins(7)11-14 pairs ,
stout , craspedodromous , alternately arising from the stout
midrib with the basal 2-3 pairs approximately opposite ,
diverging angle of the secondary veins usually 60°-65°at
the lower part of the lamina with the exception of 45°in
some specimens , and 50°-55°at the upper part of the
lamina , forking one or two times at the marginal area(Figs.1-7 , 10), diverging angle of the basal 2 -3
pairs of secondary veins usually 65°-90°(Figs.2 , 3 , 5 ,
6);tertiary veins opposite percurrent with a straight
course , oblique with an obtuse angle to the midrib(Figs.
1 , 3 , 9 , 10);areoles perfect;veinlets branched (Figs.
9 , 10).
2　Discussion
The fossil leaves described here are superficially
similar to those of living plants in Chaetoptelea , Ulmus
and Planera of Ulmaceae , Rhus of Anacardiaceae and
Sorbaria of Rosaceae.All have serrate_like margin and
pinnate craspedodromous venation with percurrent tertiary
veins.The fossils are readily distinct to the leaflet of liv-
ing Rhus in the dentate teeth with non_glandular apex and
pinnate secondary veins with increasing spacing from base
to apex as well as intersecondary veins absent.While in
the genus Rhus , the marginal teeth of are coarse with
glandular apex , the spacing of the secondary veins is usu-
ally decreasing from base to apex , the intersecondary
veins are usually well developed.The fossils frequently
forked secondary veins at marginal area and orthogonal di-
vergent angle of tertiary veins distinguish from Sorbaria.
The secondary veins of Sorbaria are seldom forked , and
the divergent angles of tertiary veins are usually acute.
Fossils described here are assignable to Ulmaceae because
of the integrated features of their leaves including the
craspedodromous venation with secondary veins forked at
the marginal area , the percurrent tertiary veins , and the
compound teeth and the right_angled sinus which being
ended by the weak tertiary veins that divergent from the
basal side of the secondary veins.Detailed comparison
between the fossils and some living genera of Ulmaceae
are shown in Table 1.The fossils differ from Chaetoptelea
in the forked secondary veins , the opposite percurrent ter-
tiary veins with orthogonal divergent angle , the convex
apical side of the teeth , and the right_angled sinuses.
The fossils are distinguished from Planera in the rather
straight secondary veins forked less than two times , the
missing intersecondary veins , the opposite percurrent ter-
tiary veins , the compound teeth with right_angled sinuses
braced by the fairly strong tertiary veins divergent from the
basal side of the secondary veins.The fossils are closely
similar to Ulmus in the lamina shape , the straight and
forked secondary veins , the percurrent tertiary veins , the
compound teeth with right_angled sinuses.
Z .furcinervis was originally described from Eastern
Sikhote_Alin in Primorye Region , Far East of Russia(Borsuk , 1952).It has been combined to Ulmus by
Ablaev (1974), as the basionym of U.furcinervis ,
based on the specimens from Takhobe Formation (Dani-
an)in the Sobolevka River and from Bogopol Formation(Danian)in the Zerkal River of Eastern Sikhote_Alin(Ablaev , 1974;Ablaev and Iljinskaja , 1982).The
leaves of U.furcinervis are characterized by the dentate
margin with broadly triangular simple or compound teeth ,
the stout and frequently forked secondary veins , and the
opposite percurrent tertiary veins.Z .furcinervis was also
described from the specimens of Boshnjakov Formation
and the Snezhinkin (Paleocene)of the Sakhalin Island(Borsuk , 1956;Sycheva , 1977).These specimens also
belong to U.furcinervis.
Specimens of U .furcinervis are found from the Ear-
ly Paleogene sediments in various localities of East Asia ,
including Kivda Formation (Paleocene)in Zeya_Bureya
Basin of the Armur Region (Kamaeva , 1982;1990),
Shestakov Formation(Eocene)in northern Priokhotye of
western Kamchatka (Ablaev , 1985)and the Zubchatyi
Cape , and Sosopkhan Formation(Paleocene)in the Tigil
region of western Kamchatka (Gladenkov et al , 1997).
Specimens from Wuyun Formation studied in this paper
are assignable to U .furcinervis , because they are closely
similar to the fossils from eastern Sikhote_Alin in leaf
※
Figs.1-10.　1.Leaf , note acutminate apex , the forked secondary veins and percurrent tertiary veins , 52717 , scale bar =10 mm.2.Leaf
formerly assigned to Sorbaria wuyungensis , 52240 , scale bar = 10 mm.3.Ulmus furcinervis from the Takuobe flora , Far East of Russia ,
601/1a, scale bar =10 mm.4.Leaf formerly assigned to Sorbaria wuyungensis , 52262 , scale bar =5 mm.5.Leaf , note the dentate mar-
gin and shallow cordate base , 52739 , scale bar = 10 mm.6.Leaf formerly assigned to Planera cf.microphylla , 52607 , scale bar = 10
mm.7.Fragment of leaf , 52614 , scale bar = 10 mm.8.Magnification of Fig.1 , showing the convex sides of teeth and the right_angled si-
nuses , 52717 , scale bar = 2.5 mm.9.Magnification of Fig.1 , showing perfect areoles and branched veinlets , 52717 , scale bar =2 mm.
10.Magnification of Fig.1 , showing forked secondary veins and percurrent tertiary veins , scale bar =5 mm.
FENG Guang_Ping et al :Paleocene Wuyun Flora in Northeast China:Ulmus furcinervis of Ulmaceae 147　
148　 植物学报　Acta Botanica Sinica　Vol.45　No.2　2003
Table 1　Comparison of the fossils leaves from Wuyun Formation with the leaves of some living genera in Ulmaceae
Fossils Chaetoptelea Planera Ulmus
Apex attenuate(+) + + + +
Base nearly symmetrical(+), or conspicuously asymmetrical(-) + + + -
Pinnate venation with stout midrib(+) + + + +
Secondary veins straight(+)or curved(-) + + - +
Secondary veins forked less than two times (+)near the marginal area or more than two
times(-) + +① - +
Intersecondary veins missing (+)or present(-) + + - +
Tertiary veins mainly opposite percurrent(+)or mainly alternate percurrent(-) + - - +
Divergent angle of tertiary veins right_angled(+)or acute (-) + + - +
Teeth mainly compound(+)or mainly simple(-) + + - +
Sinuses braced(+)by fairly strong or very weak(-)tertiary veins + + - +
Sinuses right_angled(+)or acute(-) + - - +
Apical side of teeth convex (+)or concave(-) + - + +
① usually unforked.
architecture.
Planera cf.microphylla and S.wuyungensis de-
scribed from the same locality as our collection (Tao ,
1986;2000)are also assigned to U.furcinervis consid-
ering their close similarities to the species.The diagnostic
features of Sobaria such as more than three ordered teeth ,
the acute_angled sinuses , the un_forked secondary veins ,
as well as the alternate percurrent tertiary veins are not
visible in the specimens of S.wuyungensis.Features of
Planera including the curved and frequently forked sec-
ondary veins , the conspicuous intersecondary veins as well
as the reticulate tertiary veins are not observed in the
specimens assigned to P.cf.microphylla.
P .microphylla was described from Fort Union For-
mation and Denver Formation(Paleocene)of northwestern
USA based on the frequently forked secondary veins near
the margin(Brown , 1962).This species has been allo-
cated into Chaetoptelea as the synonym of C.microphylla
based on the percurrent tertiary veins and fairly strong ter-
tiary running into the tip of sinuses of leaf margin(Hick-
ey , 1977).C.microphylla has been described from
Golden Valley Formation (Paleocene)of North Dakota ,
USA , and is characterized by more symmetric lamina
shape , irregular and forked secondary veins , moderate to
wide divergent angle of secondary veins and percurrent
tertiary veins(Hickey , 1977).Fossils described here are
comparable to C.microphylla in the frequently forked
secondary veins , the percurrent tertiary and the convex to
acuminate apical side and the convex basal side of
marginal teeth.We suggest that these fossils are probably
conspecific considering their close similarities.Further
works should be done to get more evidence and to verify
this idea.However , the assignment of the fossils from
Northeast China into Chaetoptelea is rejected based on the
forked secondary veins , the opposite percurrent tertiary
veins as well as the convex apical side of marginal teeth.
In living Chaetoptelea , the secondary veins are seldom
forking , the tertiary veins are mostly alternative percur-
rent , the apical side of the marginal teeth is usually con-
cave.
Ulmites ulmifolius has been described from Ny_
lesund Formation , Renardodden Formation and Aspelin-
toppen Formation(Paleocene)of Spitsbergen (Kvacek et
al , 1994).Fossils described here are comparable to the
species.Both have dentate margin , forked secondary
veins and percurrent tertiary veins.The suggestion that
these fossils are conspecific is rejected considering the
acute teeth observed in U.ulmifolius.
Ulmus furcinervis was considered to be one of the
common component in the Kamskyi palaeofloristic complex(late Early Paleocene to early Late Paleocene)based on
its wide distribution in various Paleocene localities(Ko-
drul , 1999).This species represented in younger palae-
ofloristic complex , such as Naibutin and Nizhnesnezhinkin
complex (Early Eocene to Middle Eocene).Fossils
assignable to U.furcinervis were also found from the
Eocene of western Kamchatka (Ablaev , 1985).U.e-
zoana was described from the Eocene of South Sakhalin(Kodrul , 1999), but no figure showed its features.From
the description of U.ezoana , it differs from U .furcin-
ervis in the almost parallel and not frequently forking sec-
ondary veins.This difference is not essential , because the
features are also visible in some specimens of U.furcin-
ervis.
The genus Ulmus seems to have a continuous distri-
bution in East Asia and North America.The close simi-
larities between U.furcinervis from East Asia and C.
microphylla from North America imply that these two pa-
leofloras probably have common component during the ear-
ly Tertiary.The similarities between these two paleofloras
of East Asia andNorth America have also been verified by
the continuous distribution of Metasequoia occidentalis(Liu et al , 1999), Beringiaphyllum pseudoantiquum and
Amersinia obtrullata (Feng et al , 2002), as well as
Nyssidium fruits(Feng et al , 2000).
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(Managing editor:HAN Ya_Qin(intern))
150　 植物学报　Acta Botanica Sinica　Vol.45　No.2　2003
中国东北古新世乌云植物群:榆科叉脉榆
冯广平1　Albert G.ABLAEV2　王宇飞1　李承森1＊
(1.中国科学院植物研究所 , 北京 100093;
2.Pacific Oceanological Institute , Russian Academy of Sciences , Vladivostok 600041, Russia)
摘要:　根据叶结构特征 , 将黑龙江省古新世乌云植物群中的榆科叶化石确定为叉脉榆(Ulmus furcinervis (Borsuk)
Ablaev),叶化石二级脉达缘且在叶缘区分叉 、三级脉及顶 、叶缘锯齿两侧边凸出。重新研究了前人鉴定为小叶普拉
榆(Planera cf.microphylla auct.non Newberry)和乌云珍珠梅(Sorbaria wuyungensis Tao et Xiong)的叶化石并将其归入叉
脉榆。
关键词:　叉脉榆;榆科;古新世
中图分类号:Q914.88　　　文献标识码:A　　　文章编号:0577-7496(2003)02-0146-06
收稿日期:2002-10-31　接收日期:2002-12-23
基金项目:国家自然科学基金(30130030 , 30070056);卢嘉锡基金;中国科学院方向性项目(KSCX2_SW_108)。
＊通讯作者。E_mai l:.
(责任编辑:韩亚琴(实习))
FENG Guang_Ping et al :Paleocene Wuyun Flora in Northeast China:Ulmus furcinervis of Ulmaceae 151