全 文 ：植 物 分 类 学 报 44 (2): 161–164(2006) doi:10.1360/aps050121
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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Received: 18 August 2005 Accepted: 20 January 2006
Supported by the Science and Technology Foundation of China University of Geosciences (Beijing), Grant No. 200405, and the
National Natural Science Foundation of China, Grant Nos. 49872008, 40372021.
* E-mail: .
** E-mail: .
Parasphenopteris Sun & Deng, a new genus from the Lower
Permian of Wuda, Nei Mongol, China
1SUN Ke-Qin* 2DENG Sheng-Hui**
1(State Key Laboratory of Geological Processes and Mineral Resources, Key Laboratory of Lithosphere Tectonics and
Lithoprobing Technology of Ministry of Education, China University of Geosciences, Beijing 100083, China)
2(Research Institute of Petroleum Exploration and Development, Beijing 100083, China)
Abstract Parasphenopteris orientalis Sun & Deng gen. & sp. nov. is described from the
Shanxi (Shansi) Formation of the Lower Permian in the Wuda area of Nei Mongol, China. The
new genus is characterized by having “stipules” at the base of penultimate pinnae and is quite
different from all other known genera in the Paleozoic.
Key words Parasphenopteris Sun & Deng, Parasphenopteris orientalis Sun & Deng, new
genus, new species, Lower Permian, Wuda, Nei Mongol, China.
The northern part of the Helan Mountain, where Wuda is located, is one of the most
important localities of the Cathaysian flora. In this locality the Carboniferous and Permian are
successionally developed, which are rich in plant fossils. We have collected numerous well
preserved plant fossils from the Wuda area of Nei Mongol during the long-term geological
investigation in the area carried out since the 1990’s. Some of the materials including new
genera and species have been published recently (Sun & Deng, 1999, 2003; Sun et al., 1999;
Deng et al., 2000). Our collection includes eight very interesting and excellently preserved
specimens, which are described in the present paper. It is highly surprising that the fossil plant
has “stipules” borne at the base of each of the penultimate pinnae. The presence of stipules
characterizes some of the living angiosperms, such as Pisum L. of Leguminosae and Smilax
riparia A. DC. of Smilacaceae, but has never been reported in the Palaeozoic plants described
thus far. Such an unexpected character found in a Palaeozoic plant shows not only the
importance of the fossils but also indicates that they represent a hitherto undescribed new
plant. We consider that this plant is worthy of a generic rank. We therefore name the plant as
Parasphenopteris orientalis Sun & Deng, gen. & sp. nov.
It is difficult to make a definite assignment of the new genus at the moment, but based on
the morphology of the pinnate compound leaf, the new genus is most probably close to
sphenopterides of the Filicopsida or the Pteridospermopsida.
1 Description
Parasphenopteris Sun & Deng, gen. nov.
Generic diagnosis: The fronds are very large and at least tripinnate. Main rachis is
smooth and thick. Penultimate pinnae are lanceolate. The rachises of penultimate pinnate are
slender. Ultimate pinnae are narrowly deltoid-lanceolate to ovate-triangular, and closely set.
The pinnules are sphenopteroid. This new genus has “stipules” at the base of each of the
penultimate pinnae. The “stipules” are linear to linear-lanceolate. Pinnae are very small, ovate
or ovoid-triangular.
Acta Phytotaxonomica Sinica Vol. 44 162
Type: Parasphenopteris orientalis Sun & Deng, sp. nov.
Parasphenopteris orientalis Sun & Deng, sp. nov. Fig. 1
The best specimen of this species is indicated in Fig. 1. The figure is in natural size. The
frond is very large and is at least tripinnate. The main rachis is smooth and thick, 3 to 4 mm
wide. Penultimate pinnae are lanceolate, alternate, 3 to 18 cm long, 0.6 to 3 cm wide, forming
an angle of 80o–90o with the main rachis. The rachises of penultimate pinnate are slender, 1
to 1.5 mm wide. Ultimate pinnae are narrowly deltoid-lanceolate to ovate-triangular, closely
set, forming angles of 40o–60o with the rachis. The “stipules” are borne at the base of each
of the penultimate pinnae, linear to linear-lanceolate, 1.5 to 2.5 cm long, and often dissected
into linear lobes. The pinnules are very small, ovate or ovoid-triangular. The pinnules are
crowded and somewhat overlapping. The venation is indistinct.
Locality and horizon: The Shanxi Formation of the Lower Permian in the Wuda area of
Nei Mongol, China.
Etymology: The generic name “Parasphenopteris” indicates that the fossils are similar to
Sphenopteris morphologically, and the specific epithet “orientalis” refers to the locality of the
materials.
Holotype: WD20026 (Fig. 1).
Repository: China University of Geosciences, Beijing, China.
2 Comparison and Discussion
A lot of living angiosperms have stipules. Stipules are commonly borne at the position
where the petioles are attached, usually binate. The functions of stipules are considered as
protecting the leaf buds and enlarging the photosynthesis area. Stipules are in various shapes.
For examples, Pisum of Leguminosae has large stipules similar to the normal leaves, while
Sorbaria (Ser. ex DC.) A. Braun of Rosaceae has small stipules; Polygontum of Polygonaceae
has membranous sheath-like stipules; and the stipules of Smilax riparia of Smilacaceae are
tendril-like, which are very close to those of the new fossil plant here described.
Although the “stipules” of the new fossil plant are very similar to some of the living
plants, we have added quotation marks to the term temporarily because a fossil plant with
distinct stipules has yet never been reported so far from the Palaeozoic. It is noteworthy that
the “stipules” are borne at both the basiscopic and acroscopic bases of each of the penultimate
pinnae instead of at the bases of pinnules.
The present new genus and species are characterized by its multi-pinnate compound
leaves, which are somewhat similar to those in the genus Sphenopteris. However, the
presence of “stipules” in the species shows that it is obviously different from all the known
Palaeozoic plants.
The discovery of Parasphenopteris has not only added a new member for the Cathaysian
flora but also has provided a new clue of the evolution of plant organs in the plant kingdom.
Possessing stipules is an advanced feature of plants. The current discovery indicates that the
stipules have possibly appeared in the Late Palaeozoic, and that the new genus may be a
linking group of the evolution of the plant kingdom.



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Fig. 1. Photograph of the holotype of Parasphenopteris orientalis gen. & sp. nov. (original size), WD20026. Arrows
indicate the “stipules”.
No. 2 SUN K-Q et al.: Parasphenopteris, a new genus from the Lower Permian of Wuda, Nei Mongol, China 163


Acta Phytotaxonomica Sinica Vol. 44 164
References
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内蒙古乌达地区下二叠统植物化石新属——
拟楔羊齿属
1孙克勤* 2邓胜徽**
1(中国地质大学地质过程与矿产资源国家重点实验室和岩石圈构造、
深部过程及探测技术教育部重点实验室 北京 100083)
2(中国石油勘探开发研究院 北京 100083)
摘要 描述了产于内蒙古乌达地区下二叠统山西组的东方拟楔羊齿Parasphenopteris orientalis Sun &
Deng (新属新种)。该植物以末二次羽片基部着生有“托叶”为特征, 区别于晚古生代其他已知植物。
关键词 拟楔羊齿; 东方拟楔羊齿; 新属; 新种; 下二叠统; 乌达; 内蒙古; 中国