全 文 ：植 物 分 类 学 报 44 (6): 712–720(2006) doi:10.1360/aps040129
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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Received: 26 October 2004 Accepted: 19 August 2005
Supported by the National Natural Science Foundation of China, Grant Nos. 30470117 and 30530050.
* Author for correspondence. E-mail: lics@ibcas.ac.cn; Tel.: +86-10-62836436; Fax: +86-10-62593385.
Paleocene Wuyun flora in Northeast China: Woodwardia
bureiensis, Dryopteris sp. and Osmunda sachalinensis
1, 2, 3WANG Qing 4Albert G. ABLAEV 1WANG Yu-Fei 1, 5LI Cheng-Sen*
1(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany,
the Chinese Academy of Sciences, Beijing 100093, China)
2(College of Life Sciences, Liaoning Normal University, Dalian 116029, China)
3(Graduate School of the Chinese Academy of Sciences, Beijing 100039, China)
4(Pacific Oceanological Institute, Russian Academy of Sciences, Vladivostok 600041, Russia)
5(Beijing Museum of Natural History, Beijing 100050, China)
Abstract Woodwardia bureiensis Fedotov, Dryopteris sp. and Osmunda sachalinensis
Krysht. were first reported in the Paleocene Wuyun flora from Jiayin County, Heilongjiang
Province, China, and they are also new records to China. Woodwardia bureiensis is different
from other species of the genus in that the pinnules of sterile fronds have a closed space with
touching margin. The pinnules of Osmunda sachalinensis Krysht. from Wuyun flora are
broadly cuneate at their base, while those of O. greenlandica (Heer) Brown, described from
the same flora previously, expand at the pinnule base.
Key words Woodwardia, Dryopteris, Osmunda, Paleocene, Wuyun flora, China.
The Wuyun flora found at the Wuyun coal mine (49˚14′26″ N, 129˚28′00″ E), Jiayin
County, Heilongjiang Province, China (Liu, 1983, 1990; Luo et al., 1983; Xiong, 1986; Feng
et al., 2000a, b, 2002, 2003) was first dated as Paleocene-Early Eocene (Jiang, 1963), then
thought to be Late Cretaceous (Hsü, 1983), Maastrichtian-Danian (Zhang, 1983; Tao &
Xiong, 1986) and finally Paleocene (also see Luo et al., 1983; Liu, 1983, 1990; Xiong, 1986;
Feng et al., 2000a, b).
In the Wuyun flora, 75.5% of the genera are angiosperms, and 14.3% are gymnosperms
(Feng, 2000). Only four ferns (macrofossils), i.e. Gleichenites gracilis (Heer) Tanai,
Cladophlebis septentrionalis Hollick, Osmunda greenlandica (Heer) Brown, Thelypterites sp.
were reported in the flora (Tao and Xiong, 1986; Tao, 2000). Palynological investigation
demonstrated the presence of 41 genera of angiosperms, 17 genera of gymnosperms and 16
taxa of ferns in the flora (Liu, 1983).
In this paper, some new specimens of ferns of the Wuyun flora were studied and
assigned to Woodwardia bureiensis Fedotov, Dryopteris sp. and Osmunda sachalinensis
Krysht.
1 Blechnaceae
Woodwardia Smith
Woodwardia bureiensis Fedotov, 1970. Fig. 1: A–D; Fig. 2: A.
Fedotov, 1970, p. 94, pl. 15, figs. 1–9, text-figs. 1a, 1b;
Kamaeva, 1982, p. 73, pl. 8, fig. 1;
Kamaeva, 1990, p. 31, pl. 1, figs. 1–6; pl. 2, figs. 1–4; pl. 3, figs. 1, 2.

No. 6 Wang Q et al.: Paleocene Wuyun Flora in Northeast China 713


Fig. 1. Woodwardia bureiensis Fedotov. A–D. Specimen number 52370–52373. Bar=10 mm.
Acta Phytotaxonomica Sinica Vol. 44 714
Locality: Wuyun Coal Mine, Jiayin County, Heilongjiang Province, China.
Stratigraphy: Wuyun Formation (Paleocene).
Specimen number: 52370, 52371, 52373, 52374.
Repository: National Museum of Plant History of China, Institute of Botany, the Chinese
Academy of Sciences, Beijing.
Description: Sterile fronds 3–7.5 cm long, 3–4.5 cm wide. Rachis 0.8 mm wide. Pinnules
oblong-triangular, sickle-shaped with apices acuminate or acuminate-triangular, 1.1–2.6 cm
long, 0.4–0.6 cm wide. Pinnules sessile, arising from the rachis at approximately 70o.
Pinnules separated from one another by the narrow, rounded sinuses within 1.5–3 mm of the
rachis. Pinnule margin entire, undulate-emarginate. Venation pinnately dictyodromous.
Secondary veins near rachis and near midvein of pinnules forming a chain of elongate areoles,
1.1–4 mm long, 0.25–1.8 mm wide, without free-ending veinlet. Tertiary vein simple or often
1–2 times branched and connected by anastomoses. Tertiary veins diverging to the outside
and their areoles extending to the margin.
Discussion: The genus Woodwardia was established by Smith in 1793 and is represented
by 13 existing species (Kubitzki, 1990). The earliest fossil record of the genus is in Late
Cretaceous (Li & Yang, 1984). During Late Paleocene and Eocene, the ferns of Woodwardia
had a circum-arctic distribution (Collinson, 2001). The fairly common leaf compression of
this genus was found throughout the Tertiary in Europe, North America, and Asia. Over 50
species were listed under Woodwardia or under the variant name Woodwardites in the
Fossilium Catalogus (Pigg & Rothwell, 2001). When the climate changed cool in Quaternary,
the plants of Woodwardia concentrated their distribution area southwards to South Europe,
Asia and America. The diversity center of the genus is in East Asia, especially in the
mountainous regions of central mainland and Taiwan of China (Chiu, 1974; Tryon & Tryon,
1982).
The fossils of Woodwardia (?) were reported in the Eocene Fushun flora at the West
Opencast Coal Mine, Fushun City, Liaoning Province (WGCPC, 1978). The specimens of
Woodwardia sp. were also reported in the Miocene Qiuligou flora, Dunhua County, Jilin
Province, China (Li & Yang, 1984).
The sterile fronds of Woodwardia bureiensis were found from the Eocene Rajczicha
flora in the Zeya-Bureya depression of Priamurye near Rajczicha City, Far East Russia
(Fedotov, 1970). Subsequently, the species was discovered in the Paleocene Kivda flora in the
same depression area (Kamaeva, 1982, 1990). In South Sakhalin, Woodwardia sp. and W.
endoana Oishi & Huzioka from the Paleocene Kamskaya beds and the Eocene
Verkhnesnezhinkinskaya subformation were established (Kodrul, 1999). The fronds from
both localities resemble W. bureiensis, but it would be premature to reassign them to this
species because of their poor preservation.
Two species of Woodwardia, W. endoana Oishi & Huzioka and W. decurrens Oishi &
Huzioka, were originally described from the Eocene Woodwardia flora (Oishi & Huzioka,
1941). According to Endo (1968), both species were referred to W. japonica Sw. var.
eocenica, because of their resemblance to the extant Woodwardia japonica.
It is necessary to emphasize that the sterile fronds of the Woodwardia species from
Priamurye (the Kivda and the Wuyun floras) and Hokkaidao (the Woodwardia flora) are
different in the arrangement of minor pinnules. The pinnules of W. bureiensis are closely
spaced, whereas pinnules of Woodwardia from Hokkaido are widely spaced. And the sori on
the Japanese specimens are restricted to the areoles, while sori on the Kivda material are
found on the margins of the pinnae and vaguely separated from one another (Fedotov, 1970,
p. 95).
No. 6 Wang Q et al.: Paleocene Wuyun Flora in Northeast China 715
A specific feature of W. sasa Oishi & Huzioka from the Eocene Harutori flora (Tanai,
1970) is the distinctly serrulate margin of pinnules. It suggests that the affinity of W. sasa is
close to the extant species of W. japonica. Krassilov (1979) considered that Woodwardia sp.
with the bipinnate fertile frond from the Danian Boshnyakovskaya flora of Western Sakhalin
would be related to W. sasa and also to the North American species W. arctica (Heer) Brown
from the Fort Union series (Brown, 1962). As noted by Krassilov (1979, p.92) Woodwardia
sp. is probably a more ancient representative of the genus, playing a great role in the
composition of the Paleogene floras of the Far East.
2 Dryopteridaceae
Dryopteris Adans.
Dryopteris sp. Fig. 2: B.
Locality: Wuyun Coal Mine, Jiayin County, Heilongjiang Province, China.
Stratigraphy: Wuyun Formation (Paleocene).
Specimen number: 52374.
Repository: National Museum of Plant History of China, Institute of Botany, The
Chinese Academy of Sciences, Beijing.
Description: Sterile pinna of the last order (only one half pinna preserved) linear-
lanceolate, 2.5 cm long, 1.5 cm wide. Costate rachis slightly curved, 0.5–0.8 mm wide,
divided, approximately on 1/3 wide hemi-plate of pinna. The pinnules rounded-triangular in
shape, their margin serrulate. Venation of pinnules pinnate. Thin midvein of pinnules
decurrent to the rachis, and extending to the apex, Midvein producing 5(6) pairs of lateral
veins at acute angle. Lateral veins once, seldom twice forked. In interval between midveins
along the rachis, veins developing and extending from the rachis at right angles. These veins,
close to their base, once forking.
Discussion: The genus of Dryopteris has about 400 extant species. The fossils of
Dryopteris were mainly found from Tertiary in Europe (Wu & Qin, 1991). No fossil of
Dryopteris has been recorded in Far East so far (Krassilov, 1976; Budantsev, 1997; Kodrul,
1999). The specimen of Dryopteris sp. studied in the work is the first fossil record of the
genus in China and it resembles D. serrata Brown (Brown, 1962) from the Paleocene Fort
Union flora in USA. Unfortunately, confirmation of their similarity is impossible owing to the
poor preservation of the Wuyun specimen.
3 Osmundaceae
Osmunda L.
Osmunda sachalinensis Krysht. Fig. 2: C, D.
Osmunda sachalinensis Kryshtofovich, 1936, p. 708, pl. 1, figs. 4–6, text-figs. 1, 2;
Borsuk, 1956, p. 13, pl. 1, figs. 1–3; Tanai, 1970, p. 456, pl. 3, figs. 5–7; Sychova, 1977, p.
19, pl. 14, figs. 6, 7; Ablaev, 1985, p. 35, pl. 1, fig. 2, pl. 15, fig. 7; Kodrul, 1999, p. 80, pl.
43, figs. 1–3.
Osmunda cf. sachalinensis : Ablaev, 1974, p. 53, pl. 1, fig. 1, text-fig. 8; Kamaeva, 1990,
p. 32, pl. 2, fig. 5, pl. 3, fig. 9.
Osmunda regalis L.: Endo, 1968, p. 416, text-fig. 4.
Locality: The Wuyun Coal Mine, Jiayin County, Heilongjiang Province, China.
Stratigraphy: Wuyun Formation (Paleocene).
Specimen number: 52375, 52376.
Repository: National Museum of Plant History of China, Institute of Botany, The
Chinese Academy of Sciences, Beijing.
Acta Phytotaxonomica Sinica Vol. 44 716


Fig. 2. A. Magnification of the part of Fig. 1, C, showing tertiary vein and entire pinnule margin. B. Dryopteris sp.
52374. C, D. Osmunda sachalinensis Krysht. Specimen number 52375,52376. Bar=10 mm.

No. 6 Wang Q et al.: Paleocene Wuyun Flora in Northeast China 717
Description: Sterile pinnule tongue-shaped, asymmetrical, 3.7 cm long, 1.8 cm wide.
Base broadly cuneate and apex broadly triangular. Pinnule margin minutely crenulate.
Venation pinnate, craspedodromous. Midvein thin, 0.8 mm wide at base, slightly sinuous
caused by departing lateral veins, extending to the apex. Lateral veins alternate, emerging
from midvein at an acute angle, up to 12 veins per cm. Lateral veins usually branching
dichotomously twice, with first fork near base and second fork close to margin. Lateral veins
near the apex simple, while lowest ones branching thrice. All veins reach the marginal sinuses
between equal-sized obtuse small teeth.
Discussion: The fronds collected from the Wuyun flora were assigned as Osmunda
greenlandica (Heer) Brown previously (Tao & Xiong, 1986; Tao, 2000). The pinnules of O.
greenlandica are 1.9 cm long, 0.8 cm wide and expand at the base, whereas those of O.
sachalinensis in this work are 3.7 cm long, 1.8 cm wide, broadly cuneate at the base.
The fern of O. greenlandica was found from the Early Tertiary flora of the Fort Union
Formation (lower) and the Denver Formation, in North America (Brown, 1962). Budantsev
(1997) believes that O. greenlandica from North America and Northern Priokhotye (Ablaev,
1985) probably belongs to the genus Osmundastrum (Osmundaceae). The type species
Osmundastrum heislianum (Fotjan.) Budants. was first described by Fotjanova (Sinelnikova
et al., 1976) as Osmunda heisliana from the Pliocene Ermanovskaya flora of Western
Kamchatka.
Osmunda sachalinensis is widely represented in the Paleogene floras of the Far East. It
was first described from the Lower Due flora of Sakhalin (Kryshtofovich, 1936), and was
later discovered in the fossil floras of Primorye (Ablaev, 1974), Priamurye (Kamaeva, 1990),
Japan (Tanai, 1970) and Northern Priokhotye (Ablaev, 1985). According to Fotjanova
(Sinelnikova et al., 1976), O. sachalinensis from the Pliocene Ermanovskaya flora of Western
Kamchatka is represented by a single pinnule, 5 cm long and 1.5 cm wide, with entire margin
and small lateral veins (7 per cm). In the Miocene Medvezhkinskaya flora of Eastern
Kamchatka (western shore of Korf Bay), Chelebaeva (1978) described O. doroshiana Goepp.
from the Early Tertiary Arctic floras (Alaska, Greenland). The specimens of O. sachalinensis
described by Kryshtofovich (1936) were assigned to this species by Chelebaeva (1978). In our
view, it is premature to categorize this species under O. doroshiana①, as the leaf impression
from Kamchatka is not complete (apex and base are broken).
In the Paleogene floras of Western Kamchatka, Budantsev (1983, 1997) applied the
oldest binomial O. macrophylla Penhallow, a species known from a number of
Paleocene-Eocene floras of North America. Its pinnules from Kamchatka are tongue-shaped,
reaching 5 cm long and 1.5–1.7 cm wide, with wavy-crenulate or crenulate margin. Single
pinnulae from the Maastrichtian Augustovskaya flora of Western Sakhalin basin (Augustovka
River) was also referred to as O. macrophylla (Krassilov, 1979). The pinnulae are
tongue-shaped, 4.2 cm long and 1.4 cm wide, with wavy margin, and up to 20 lateral veins
per cm. As underlined by Boulter and Kvacek (1989, p. 26), specific differences between O.
sachalinensis and O. macrophylla are not well defined, being mainly based on the size of the
pinnules and the number of veins.
Chelebaeva (1971) described similar leaf impressions as O. heeri Gaudin, sine descr.
from the Miocene flora in the basin of the Krutogorova River in Western Kamchatka. The
pinnules of O. heeri are different from those of O. sachalinensis in size, to 12 cm long and 3.5
cm wide in the former fern. O. sachalinensis closely resembles the modern O. japonica Thunb
in morphology, the latter grows in Eastern Asia and the Himalayas.

① Subsequently, to Osmunda heeri Gaudin (Iljinskaja, 1968) known from the Neogene floras of Europe the forms from the Arctic,
Kazakhstan, and Sakhalin were also assigned.
Acta Phytotaxonomica Sinica Vol. 44 718
Species diversity of ferns from the Wuyun flora is similar to the floristic composition of
the Kivda flora, Priamurye, Far East Russia, viz. Woodwardia bureiensis, Osmunda
sachalinensis, “Polypodiopteris” kivdensis Fedotov, Cladophlebis arctica (Heer) Seward, C.
aff. arctica (Heer) Seward, C. joergensenii (Heer) Krysht. and C. cf. oerstedtii (Heer) Seward.
It may provide new evidence to show that the two floras have the same age (Paleocene) (Feng
et al., 2000a, 2002, 2003).
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Acta Phytotaxonomica Sinica Vol. 44 720
中国东北古新世乌云植物群: Woodwardia
bureiensis, Dryopteris sp.和
Osmunda sachalinensis

1,2,3王 青 4Albert G. ABLAEV 1王宇飞 1,5李承森*
1(系统与进化植物学国家重点实验室, 中国科学院植物研究所 北京 100093 )
2(辽宁师范大学生命科学学院 大连 116029)
3(中国科学院研究生院 北京 100039)
4(Pacific Oceanological Institute, Russian Academy of Sciences, Vladivostok 600041, Russia)
5(北京自然博物馆 北京 100050)

摘要 在黑龙江省嘉荫县乌云镇古新世乌云植物群中新发现的Woodwardia bureiensis Fedotov、
Dryopteris sp.和Osmunda sachalinensis Krysht.在中国也是新记录。Woodwardia bureiensis与其他种的主要
区别是小羽片排列紧密且边缘彼此靠合。Osmunda sachalinensis小羽片基部宽楔, 而以前乌云植物群所
报道的O. greenlandica (Heer) Brown小羽片基部平展。
关键词 Woodwardia; Dryopteris; Osmunda; 古新世; 乌云植物群; 中国