全 文 ：植 物 分 类 学 报 45 (4): 576–582(2007) doi:10.1360/aps06030
Acta Phytotaxonomica Sinica http://www.plantsystematics.com
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Received: 28 February 2006 Accepted: 14 February 2007
* Author for correspondence. E-mail: .
Palaeoskapha sichuanensis gen. et sp. nov. (Menispermaceae)
from the Eocene Relu Formation in western
Sichuan, West China
1Frédéric M. B. JACQUES* 2, 3GUO Shuang-Xing
1(Muséum National dHistoire Naturelle, Département Histoire de la Terre, CP 38, UMR 5143 - USM 0203,
8 rue Buffon, 75231 Paris Cedex 05, France)
2(Nanjing Institute of Geology and Palaeontology, the Chinese Academy of Sciences, Nanjing 210008, China)
3(Kunming Institute of Botany, the Chinese Academy of Sciences, Kunming 650204, China)
Abstract Palaeoskapha sichuanensis gen. et sp. nov. of Menispermaceae is described here
for the first time based on a well preserved fossil fruit. The specimen was found in the Relu
Formation of western Sichuan, West China. The specimen, straight, boat-shaped endocarp
with large ventral condyle, clearly belongs to the tribe Tinosporeae. The wide aperture of the
double condyle, combined with a whole shape not deeply invaginated, indicates a genus
different from what was already known to science for this tribe. This fossil widens the
distribution of the tribe during Eocene from North America and Europe to Asia, where it was
formerly unknown.
Key words Eocene, Menispermaceae, Palaeoskapha sichuanensis sp. nov., Relu Formation,
Sichuan.
Modern Menispermaceae (i.e. Moonseed family) consist of approximately 70 genera and
450 species, distributed in tropical lowland regions (Kessler, 1993). They are usually twining
vines, whose fruits are one-seeded drupes with taxonomically useful endocarp (Thanikaimoni,
1986).
Only a few occurrences of Menispermaceae are known in the Cenozoic of Asia (except
India): Menispermoxylon circumvallum Vozenin-Serra & Privé-Gill, 1989, a fossil wood from
Thailand (Miocene), Entoneuron melastomaceum Geyler, 1878, fossil leaf from Borneo
(Eocene), Cocculus flabella (Newb.) Wolfe, 1966 fossil leaves (Guo et al., 1984) from
Xinjiang, Northwest China (Palaeocene), some leaves lacking specific identification, as
Cocculus sp. from Japan (Oligocene, Tanai, 1972; and Miocene, Ozaki, 1991), Menispermum
sp. from Miocene of Japan (Tanai & Suzuki, 1963), Stephania sp. from Pliocene of Japan
(Tanai, 1972). Other occurences are proposed but could not be checked: from the Oligocene
of Japan (Tanai, 1970), from the Palaeocene of Jiayin, Heilongjiang, Huanqu of Shanxi, the
Miocene Chifeng of Nei Mongol (Tao, 2000). The identification of fossil leaves to
Menispermaceae is often doubtful, and the leaves formerly assigned to Cocculus have been
associated with the extinct Trochodendraceous genus Nordenskioldia (Crane et al., 1991;
Manchester et al., 1991). Therefore every discovery of new fossils confirming the Cenozoic
record of the family in Asia is important. Fruits provide some of the most useful characters for
identification of Menispermaceae in the fossil record (Reid & Chandler, 1933; Thanikaimoni,
1986; Jacques & De Franceschi, 2005). The recent recognition of Diploclisia and
Palaeosinomenium endocarps from the Huadian coal mine in Jilin Province, Northeast China
(Manchester et al., 2005) confirms the presence of Menispermaceae, tribe Menispermeae in
the Middle Eocene of northeastern China. In this paper we recognize a new occurrence of the
No. 4 JACQUES & GUO: Palaeoskapha nov. gen. from the Eocene Relu Formation in China 577
Menispermaceae, based on a fossil fruit from the Eocene Relu Formation at Relu village in
Litang County of western Sichuan Plateau, West China. It is assigned to a new genus and is
significant as a member of the tribe Tinosporeae.
1 Material and methods
Two fossil fruits are collected from the Relu Formation
which is situated at Relu village in Maiwa Town (ca. 30° N,
100°32′ E) in Litang County of western Sichuan Plateau,
western China. The strata are 1477 m in total thickness. The
field outcrop section of the Relu Formation is represented in
Fig. 1.
Formerly, the geological age of the Relu Formation,
based only on the stratigraphic sequence, was considered to
be roughly Eocene. Based on more recent geological surveys,
outcrop measurement and collection of fossil plants, the Relu
Formation is primarily considered Eocene. The Eocene
assignment is supported by the composition of the fossil
flora, which is preserved in bed 8th grayish-brown felsic
sandstone. The leaf impressions include Chamaecyparis sp.,
Myrica sp., Palibinia laxifolia, Rhus turcomanica, Arundo
sp. which are shared with the Eocene Er-Oilan-Du3 fossil
plant assemblage in Turkmenistan (Vasilevskaya, 1957). The
extinct Palibinia fossils (second in abundance only to
Myrtophyllum) are restricted to the Eocene strata in Central
China, Middle Asia and South Europe, although they extend
to the early Oligocene in Germany and Austria. Trapa
paulula was found from the Palaeocene Upper Fort Union
formation in North America (Brown, 1962). Therefore, the
flora from the Relu Formation is concluded to represent
Eocene epoch in geological age, and probably Early or
Middle Eocene (GUO Shuang-Xing, unpublished manu-
script).
The specimen studied has two halves: one with the
impression of the ventral face, and one with the locule cast
seen from ventral face and still adherent to the dorsal face.

→
Fig. 1. Section of the Relu Formation. 10, purple-greyish muddy dolomite
intercalated felsic sandstone folium, denuded at top bed; 9, interchanging beds
of greyish muddy dolomite and light purple-greyish dolomitic mudstone; 8,
unequally interchanging beds of purple-greyish and greyish-brown felsic
sandstone intercalated yellow dolomitic limestone, yielding fossil plants; 7,
Unequally interchanging beds of purple-greyish felsic sandstone and siltstone; 6,
light purple-greyish calcspar and acid tuff; 5, unequally interchanged beds of
reddish sandstone, clastic sandstone and calcareous sandstone; 4, purple-reddish
muddy limestone at middle and upper part, purple-reddish conglomerate the
bottom; 3, grey-green thick conglomeratic bed with andesite gravels; 2,
brown-purple felsic sandstone with gravels intercalate purple-grey tuff, yielding
stoneworts; 1, purple-red sandy limestone intercalated with gypsum folium at
middle and upper part, conglomerate at lower part. 0, unconformity, underlying
strata: Upper Triassic. * Fossil position.

Acta Phytotaxonomica Sinica Vol. 45 578
The correct interpretation of the fossil therefore needs a reconstruction of its shape using these
two halves.
2 Results
Palaeoskapha Jacques & Guo, gen. nov.
Type species: Palaeoskapha sichuanensis Jacques & Guo, sp. nov.
Etymology: Palaios (Old Greek = ancient) and Skaphe (Old Greek = boat), referring to
the general shape of the endocarp.
Palaeoskapha sichuanensis Jacques & Guo, sp. nov. Fig. 2: A, B
Syntypes: PB12703 (endocarp) and PB12702 (locule cast).
Repository: Nanjing Institute of Geology and Palaeontology (NIGPAS), the Chinese
Academy of Sciences, China.
Type locality: Relu village (Litang, Sichuan, China).
Stratigraphic horizon: The bed 8th grayish-brown felsic sandstone of the Relu Formation
(Eocene).
Description: Endocarp more or less boat-shaped, elliptic in dorsiventral outline,
bisymmetric, with a sharp ventral median ridge; base rounded and apex slightly pointed, the
median ridge disappearing before the base and apex a lateral wing inferred on each side; large
intrusive condyle on ventral face, separated in two parts by the median ridge, with an elliptic
aperture as broad as the condyle, delimited by a sharp edge; surface smooth, without any
spines; length 8.5 mm, width 4.8 mm; condyle length 4.0 mm, width 3.3 mm. Locule cast
bisymmetric, with a smooth ventral median ridge; base rounded and apex pointed; large
intrusive condyle; surface smooth; length 6.0 mm, width 4.0 mm; condyle length 4.0 mm,
width 3.3 mm.
3 Discussion
This fossil was already studied by one of us in an unpublished work (Guo, 1986,
unpublished MS, p. 67, pl. 18, figs. 7, 8) under the name Carpites bivalves.
Many modern Tinosporeae have spiny projections on the endocarp, as in Parabaena
Miers, but these seem to be lacking in the fossil, as indicated by the absence of any spines on
the margin of the dorsal face. So, we infer and propose that the dorsal surface was smooth.
However, as the locule cast recovers the dorsal surface, this character has not been seen for
the whole surface. The carbonaceous part still visible on the ventral impression is relatively
thin. This indicates that the endocarp wall could have been thin. The margin of the ventral
impression is larger than the locule cast of the seed cavity. It seems to us that this margin
could not represent only the seed cavity, but is the sign of another structure. As the endocarp
wall is thin, it could not have left such an important mark. The presence of a lateral wing,
which is a usual character in the tribe Tinosporeae to which belongs Palaeoskapha, can well
explain this large margin. In modern Tinosporeae, there is no species with a winged endocarp
lacking ventral protruding chamber. Then Palaeoskapha represents a transition form between
two types of endocarp: those having wings and ventral protruding chamber, and those lacking
both.
The boat-shaped endocarp and locule cast, with straight, rather than longitudinally
curved, profile of the species clearly indicates that it belongs to the tribe Tinosporeae Hook. f.
& Thoms. This tribe includes about 19 genera that are distributed today in North and South
America, Africa and Asia. The large and widely open condyle of this fossil is similar to that
of Calycocarpum Nutt. ex Torr. & Gray and Jateorhiza Miers (Fig. 3). However it cannot be
placed into one of these genera: Jateorhiza has a flange that slightly overlaps the condyle;
No. 4 JACQUES & GUO: Palaeoskapha nov. gen. from the Eocene Relu Formation in China 579



Fig. 2. A. Endocarp of Palaeoskapha sichuanensis sp. nov. (PB12703). B. Locule cast of Palaeoskapha sichuanensis sp.
nov. (PB 12702). Scale bar = 5 mm.
Acta Phytotaxonomica Sinica Vol. 45 580
Calycocarpum is more invaginated, being hemispherical; neither of them is winged. The
lateral wings interpreted for Palaeoskapha suggest affinities to winged genera of Tinosporeae,
such as Chasmanthera Hochst. or Leptoterantha J. Louis ex Troupin, but they have a condyle
developed in protruding ventral chamber (Fig. 3). The chamber of Kolobopetalum ovatum
Stapf is the least closed species, but is not totally open. Those reasons justify the placement of
this species in a new genus.



Fig. 3. Comparison of the transversal section of endocarps of the tribe Tinosporeae and Palaeoskapha sichuanensis sp.
nov. s, seed cavity; c, condyle. A. Calycocarpum lyonii (Nutt.) Gray. B. Chasmanthera dependens Hochst. C. Jateorhiza
macrantha (Hook. f.) Exell & Mendonca. D. Leptoterantha mayumbensis (Exell) Troupin. E. Kolobopetalum ovatum Stapf.
F. Palaeoskapha sichuanensis sp. nov.


The specimen underwent a small geological deformation: a slight shift parallel to the
plane of symmetry. Both valves are thus not completely face to face any more.
Although the Palaeoskapha sichuanensis fossil is found in western China, Jateorhiza,
Chasmanthera, Kolobopetalum Engl., Leptoterantha are from Africa whereas Calycocarpum
is a North-American genus. The extant Asian Tinosporeae, including Tinospora, Parabaena,
Aspidocarya and Chlaenandra are readily distinguished in fruit morphology from
Palaeoskapha because they all lack such a widely aperturate condyle. However, usually
European and North-American Eocene Menispermaceae have Asian affinities (Reid &
Chandler, 1933; Manchester, 1994; Jacques & De Franceschi, 2005). The distributions of all
genera with affinities to Palaeoskapha sichuanensis are summarized in Table 1.
No. 4 JACQUES & GUO: Palaeoskapha nov. gen. from the Eocene Relu Formation in China 581
Table 1 Distribution of modern (M) and fossil (F) species of genera with affinites to Palaeoskapha sichuanensis
Africa North America Asia Europe
Aspidocarya M
Calycocarpum M, F F
Chasmanthera M
Chlaenandra M
Jateorhiza M F
Kolobopetalum M
Leptoterantha M
Parabaena M
Tinospora M F M F

Thanikaimoni (1986) proposed that Menispermaceae could originate from Africa, and only
later colonized Asia via India, is difficult to keep, considering the early presence of diverged
species in Eocene of China. A better knowledge of Menispermaceae fossils from Asia will
allow a better understanding of their biogeographical implications.
Other species found along with Palaeoskapha sichuanensis give good clues to the
palaeoecology of the site. The fossil plants from the Relu Formation contain 18 taxa
(including five new species) belonging to 17 genera of 11 families. The fossil taxa include
Chamaecyparis (Cupressaceae); Trapa paulula (Bell) Brown (Trapaceae); Syzygium sp.,
Myrtophyllum (two species) (Myrtaceae); Sorbus sp. (Rosaceae); Leguminocarpon sp.
(Leguminosae); Myrica sp., Palibinia laxifolia Korovin (Myricaceae); Ulmus sp., Zelkova
ungeri Kovats. (Ulmaceae); Evodia sp., Rutaphyllum (Rutaceae); Rhus turcomanica (Krysht.)
Korovin (Anacardiaceae); Cyperacites sp. (Cyperaceae); Arundo sp. (Gramineae). Besides
these this flora includes two formerly unclassifiable taxa, namely Carpites and Antholithes
spp. Among the fossil plants, the form genera Myrtophyllum and Palibinia are most abundant,
occupying 73% of the total. The evergreen and deciduous plants are nearly equal in number of
species; however, the evergreen plants are greater in abundance than the deciduous ones.
Accordingly, the fossil plants are considered to represent an evergreen broad-leaved forest.
4 Conclusion
Palaeoskapha sichuanensis represents a confirmed occurence of Menispermaceae fossil
fruits from the Cenozoic (Eocene) of China. It represents also the only occurence of the tribe
Tinosporeae, however widely present in Asia nowadays. Therefore, this carpological fossil of
Menispermaceae in Asia raises new questions concerning palaeobiogeography of this family.
The fossil shows an interesting combination of characters (a widely opened condyle
associated with a slightly pointed apex) which is not found in modern Menispermaceae. The
palaeobotanical research in Asia could provide new perspectives for our understanding of
Menispermaceae family.
Acknowledgements We are grateful to Steven MANCHESTER (Florida Museum of
Natural History, Gainesville) for making this collaboration possible and reviewing a former
version of the manuscript, Jean DEJAX (Muséum National dHistoire Naturelle, Paris) for
useful help on the manuscript, and Qi WANG (Institute of Botany, the Chinese Academy of
Sciences) for useful remarks on a former version of the manuscript.
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