全 文 ：Received 4 Feb. 2004 Accepted 26 Aug. 2004
Supported by the Shenzhen Municipal Research and Development Foundation (Cycad Collection and Conservation Program) and the Shenzhen
Urban Management Science Foundation (Cycad Fosill Collection and Research Program).
* Author for correspondence. E-mail: .
http://www.chineseplantscience.com
Acta Botanica Sinica
植 物 学 报 2004, 46 (11): 1269-1275
A New Species of Weltrichia Braun in North China with a Special
Bennettitalean Male Reproductive Organ
LI Nan1, LI Yong1, WANG Li-Xia2, ZHENG Shao-Lin3*, ZHANG Wu3
(1. Shenzhen Fairy Lake Botanical Garden, Shenzhen 518004, China;
2. Office of Fossil Protection and Management of Liaoning Province, Shenyang 110032, China;
3. Shenyang Institute of Geology and Mineral Resources, Shenyang 110032, China)
Abstract: We report a new species of the genus Weltrichia Braun with a special Bennettitalean male
reproductive organ—W. daohugouensis Li et Zheng sp. nov., which was discovered from Haifanggou Forma-
tion (Middle Jurassic) in Daohugou Village, Shantou Town, Ningcheng County, Inner Mongolia (Nei Mongol)
Autonomous Region, China. It is a unisexual male strobile which was a laterally compressed preserved
specimen being fan-shaped and appearing as half of a whole. Its base and peduncle are lost while the lower
part of the strobile consists of eleven narrowly wedged segments fused together into a funnel or cup-like.
In the upper part of the strobile these segments separate into free structure (microsporophylls), each
bearing two rows of pollen sacs (synangia). In this paper a new species W. daohugouensis Li et Zheng sp.
nov. is described and the identification of the W. huangbanjigouensis Sun et Zheng is emended.
Key words: Ningcheng County, Inner Mongolia; Haifanggou Formation; Bennettitalean male strobile;
Weltrichia daohugouensis sp. nov.
Along with the recent discoveries of pterosaurs,
saramanders, insects and fossil plants at Daohugou Village
near Ningcheng County, Inner Mongolia (Nei Mongol),
China, controversies over different opinions with regard to
the age and stratigraphic sequence of the fossil-bearing
beds have been raised. Some authors assigned the fossil
beds in the Daohugou area to the Lower Cretaceous Yixian
Formation based on the imagined occurrence of such fossil
as Lycoptera, Peipiaoosteus and Ephemeropsis and as-
serted that the beds are underlain unconformably by the
Jurassic “Tuchenzi” Formation (Wang et al., 2000), while
others considered the fossil beds to be of Middle or Late
Jurassic with uncertainty based on the simultaneous oc-
currence of the long-tailed rhamphorhynchoid pterosaurs
and short-tailed pterodactyloid pterosaurs with primitive
feathers in these beds (Ji and Yuan, 2002)
Recently, based on the field results from biostratigraphic
investigations of the Daohugou area, Ren et al. (2002)
p o in ted o u t t ha t t he insec t a s semb lage and
lithostratigraphic evidence from the Daohugou fossil-bear-
ing beds clearly indicate that they belong to the Middle
Jurassic Jiulongshan Formation.
The biostratigraphic viewpoint adopted in the present
study is condensed and improved by the authors based on
their many years’ work in this area and on the fossils studied.
In order to promote further study on the age, stratigraphic
sequence and nomenclature, we have proposed that the
Daohugou fossil beds would be assigned to the Middle
Jurssic Haifanggou Formation which is of comparable to
the Jiulongshan Formation in northern Hebei Province in
which a lot of plant fossils have been found. Among them
other plants associated with the specimens here studied
are collected from the Daohugou fossil-bearing beds. Since
1995, the Weltrichia daohugouensis Li et Zheng sp. nov.
mainly include Selaginellites chaoyangensis Zheng et Li,
Yanliaoa sinensis Pan, particularly Bennettitalean
Anomozamites haifanggouesis Zheng et Zhang, A. sinensis
Zhang et Zheng, Pterophyllum spp., Tyrmia taizishanensis
Zhang et Zheng, Wliliamsonia sp., Cycadolepis spp., etc.,
which are all important members of the plant assemblage
from the Middle Jurassic Haifanggou Formation or
Tiaojishan Formation (Zhang and Zheng, 1987; Zheng et
al., 2003).
Bennettitalean male strobiles were not found in China
until some time ago, when the first findings on the Weltrichia
were made by Sun et al. (2001). A male reproductive organ
of Weltrichia huangbanjigouensis Sun et Zheng was dis-
covered for the first time, but the specimen was incom-
pletely and without pollen sacs preserved. The inner sur-
face of the cup bears a large number of “resinous sacs”.
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041270
Since the specimens of the new species were well preserved,
their findings pushed forward the study of the male repro-
ductive organs of Bennettitalean plants in China to a higher
level.
1 Description of the New Species
Order Bennettitales
Genus Weltrichia Braun 1849 emend. Harris 1969
(Williamsonia male “flowers” of most authors)
1849. Weltrichia Braun, P. 709.
1969. Weltrichia Braun, Harris P. 158.
Diagnosis Bennettitalean male strobile consisting of a
massive cup with its upper part was divided into numerous
identical lobes or rays; rays of thick substance, tapering to
a point. Outer surface of the cup and rays without
appendages. Cuticles (where existed) showing
syndetocheilic stomata with single subsidiary cell that op-
posites each guard cell. Inner side of cup and rays bearing
pollen sacs, either directly, or on appendages. Pollen sacs
( = synangia) where existed consisting of two equal valves,
each valve with a single row of pollen chambers. Pollen
grains oval, monocolpate.
Type Species Weltrichia mirabilis Braun 1849.
Notes The above-mentioned diagnosis of the genus
Weltrichia was given by Harris (1969, P. 158) to be mainly
based on the specimens from Yorkshire. Saporta (1891) was
the first to point out that there is no difference between
Weltrichia and the Yorkshire Williamsonia flowers we later
recognized as male, but he did not unite the two genera.
Thus, for a long time, the name Weltrichia has been
neglected. Harris (1969) put them partly because Braun’s
original account has been seen by few and partly on
account of Yorkshire male strobiles (Williamson, 1870;
Saporta, 1891; Nathorst, 1909; 1911; Seward,1917) all under
the name Williamsonia overshadowed Weltrichia.
As Weltrichia has priority over Williamsonia (used for
male or female strobiles), Harris (1969) has followed the
strict interpretation of the rules and used Weltrichia for the
male strobiles and kept Williamsonia for the female ones.
It is worthy of special mention that the strange fossil
Cycadocephalus Nathorst (1902; 1909) has much common
with Weltrichia. The type specimen consists of an oval
cluster of 16-18 linear microsporophylls, 9 cm long, spring-
ing from a small circular disc formed of their concrescent
and narrow bases. The whole flower exclusive of the pe-
duncle is 10 cm long and 7 cm broad (Seward, 1917, P. 474,
Fig.574), but as its pollen bearing entirely different, are nu-
merous groups of, often tetrahedral, microspores (Nathorst,
1909), Harris (1969) doubted if it is indeed Bennettitalean.
The following species of Weltrichia are recognized
(Table 1 ): Some flowers that give no clear evidence of hav-
ing borne pollen sacs have been omitted, others may have
been missed. The genus Weltrichia was flourishing from
Rhaetic to Early-Middle Jurassic but until Late Jurassic-
Early Cretaceous, it was replaced gradually by the
Bennettitalean genus Cycadeoidea (Wieland, 1906).
Weltrichia daohugouensis sp. nov.
(Figs. 1-8, 10-13 Li et Zheng)
Diagnosis Male strobile, consisting of small cup bear-
ing at its edge a whorl of about 22 rays (microsporophylls).
Rays close or separate laterally with one another, spreading,
2.5-3.0 cm long, 6-7 mm wide at middle; base narrowed
slightly, apex obtuse-acuminate. Substance of ray rather
thick. Abaxial surface of ray with a number of hairs pointing
Table 1 Known species of Weltrichia
Species Authors Emended author Locality Age
Weltrichia alfredi Krasser, 1917 Hungary Lias
W. blandfordi Feistmantel, 1877 Harris, 1969 India Jurassic
W. campanulata Braun, 1849 Germany Rhaetic
W. daohugouensis sp. nov. Li and Zheng China Middle Jurassic
W. fabrei Saporta, 1891 France Rhaetic
W. huangbanjigouensis Sun and Zheng, 2001 China Late Jurassic
W. mirabilis Braun, 1849 Germany Rhaetic
W. mexicana Wieland, 1909;1916 Harris, 1969 Mexico Jurassic
W. oolithica Saporta, 1891 Italy Lias
W. ovalis Braun, 1849 Germany Rhaetic
W. pecten Leckenby, 1864 Harris, 1969 Britain Middle Jurassic
W. santalensis Sitholey and Bose, 1953 Harris, 1969 India Middle Jurassic
W. setosa Nathorst, 1911 Harris, 1969 Britain Middle Jurassic
W. sol Harris, 1969 Britain Middle Jurassic
W. spectabilis Nathorst, 1909 Harris, 1969 Britain Middle Jurassic
W. whitbiensis Nathorst, 1911 Harris, 1969 Britain Middle Jurassic
LI Nan et al.: A New Species of Weltrichia Braun in North China with a Special Bennettitalean Male Reproductive Organ 1271
forwards and outwards. Adaxial surface of ray in upper part
bearing pollen sacs (synangia) in two longitudinal rows.
Sacs flattened, nearly long-oval or ovate, with thickened
margin, consisting of two identical valves, keel-formed at
their contact position; attached at middle of flat side and
placed obliquely to the ray; each valve of the pollen sac
including 7-8 pollen chambers placed transversely to the
long axis of the pollen sac. Pollen grains oval with a longi-
tudinal sulcus and smooth wall; 40-45 mm long, 15-20 mm
wide. Cuticles unknown.
Holotype DHG-1a-1b (both positive and negative
impressions).
Etymology Species name refers to the fossil locality
name-Daohugou.
Locality and stratigraphy Daohugou village, Shantou
Town, Ningcheng County, Inner Mongolia Autonomous
Region, China; Middle Jurassic Haifanggou Formation.
Description of specimen The specimens of the W.
daohugouensis sp. nov. are relatively complete and well
preserved except their cuticles. Some parts of the speci-
mens were made into balsam transfers, with well preserved
pollen sacs, pollen chambers, pollen grains and vascular
tissues. The shapes of male strobile and its respective tis-
sues are described below.
Male strobile Strobile as half of a whole is compressed
on the same plane of mother rock and is fan-shaped (Figs.
2, 3). Base of strobile is not preserved. The height of
strobile reaches up to 5 cm and is about 8 cm broad in upper
part of the strobile.
Cup Cup of strobile as half of a whole consists of 11
narrowly wedged segments which are fused into a funnel
shaped structure. The cup is 2.5 cm high. On adaxial sur-
face of each segment near the upper part there are two rows
of alternately seated small scars, which may represent as
rudimentary synangia. There are a few vein-like fibers in
the lower part of the segments. (Fig.1A).
Ray (Microsporophyll) Rays arise from the top of the
cup segments and the number of which correspond with
that of the segments. Rays are in contact with or separate
laterally from one another spreading on that same plane
and their upper parts curve outwards. The rays are 2.5 cm
long and 4-5 mm wide at the middle. Apex obtuse or ob-
tuse-pointed. Substance of ray is rather thick. Abaxial sur-
face of ray is densely covered with hairs pointing forwards
and outwards (Figs. 1A, 13). The adaxial surface of ray is
hairless but there are two rows of pollen sac in the upper
part.
Pollen sac Sacs are flattened, narrowly ovoid or ovoid
with a thickened margin and placed obliquely to the ray
axis. Pollen sac is about 2 mm long and 1.0-1.5 mm wide at
Fig.1. Weltrichia daohugouensis sp. nov. A. Restoration of part of a strobile showing microsporophylls with pollen sacs, cup and
bristles at margin of rays. B. A pollen sac showing many pollinic chambers. C. Anastomosing bundles between pollen sacs. D, E. Pollen
grains.
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041272
the middle and is composed of two equal valves. There is a
keel between both valves. Each valve encloses a single
row of transversely elongated pollen chambers (Figs.1B,
11, 12). There are a few of anastomosing bundles in space
between the pollen sacs (Figs.1C, 10).
Pollen grain Many pollen masses are seen in the
pollen chambers but some of them are scattered everywhere
in the pollen sacs. A pollen grain is ovoid , 40 mm×20 mm in
size, with a thin and smooth wall, bearing a longitudinal
sulcus (Figs.1D,E, 6-8).
2 Discussion and Comparison
Discussion In description of the new species we have
followed Harris (1969) who preferred “pollen sac” to
“synangia” for the small 2-valved capsule containing pollen.
Although “synangia” has been considerable, it is insecure
Figs.2-9. Weltrichia daohugouensis sp. nov. 2-3. Bennettitalean male strobile. Its lower part is in a form of an incomplete cup made
of the concrete bases of about 11 narrow and wedged segments; its upper part is divided into separate rays, × 1. 4-5. The upper part
of the rays, showing pollen sacs (synangia) bearing pollen chambers, ×10. 6-8. Pollen masses and pollen grains, ×300. 9. Cycadolepis
sp., together with W. daohugouensis sp. nov. were found in the same fossil-bearing bed.
LI Nan et al.: A New Species of Weltrichia Braun in North China with a Special Bennettitalean Male Reproductive Organ 1273
in morphological implications, particularly in its resemblance
to Marrattiopsis synangia. Similarly he preferred the purely
descriptive term “ray” to “microsporophyll” for the seg-
ments as the segment are separated and spread out from
the cup.
In some species of Weltrichia aside from the rays that
Figs.10-13. Weltrichia daohugouensis sp. nov. 10. Hypodermally anastomosing vascular bundles between the pollen sacs, ×30. 11-
12. Pollen sacs: showing a keel between two valves, and separated pollen masses in each valve, ×60. 13. Hairs on lateral sides of rays
(microporophylls), ×10.
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041274
bear two rows of pollen sacs, some smaller sac-like traces
continue into the cup. Nathorst (1909; 1911) recognized
them in W. whitbiensis and called them as “rudimentary
synangia” because they continue the pollen sac series into
the cup and they resemble pollen sacs. Harris (1969),
however, recognized that the pollen sacs are delicate, eas-
ily detached, and rapidly macerated, while the resinous sacs
are firmly attached and very resistant and even after a long
time they still contain much material which continues to
resist maceration so that the outer cuticles are hard to be
separated.
In the new species presently described we have only
seen the structure that may like to those of “rudimentary
sacs” in the W. whitibiensis (Nathorst, 1911, P. 7; Fig.11)
but no “resinous sacs” were observed as those in W. sol
(Harris, 1969, P. 164; Fig. 69D, E).
In addition, as we know all European, Northern Ameri-
can and Indian species of Wetrichia were associated with
leafy organs, such as Bennettitalean Ptilophyllum, Zamites,
Otozamites and Nilssoniopteris, etc. however, what are
associated with W. daohugouensis sp. nov., are mainly
Pterophyllum, Tymia and Anomozamites and none of the
above mentioned fossils were found in the Daohugou fos-
sil-bearing beds. Thus the new species may not be the
same plant as the known species.
Comparison The new species is very similar to W.
setosa (Nathorst) Harris in that the ray bears some hairs,
but in the latter, the small cup bears a whorl of about 20
rays (microsporophylls) at its edge and inside the cup are a
whorl of numerous ( about 40) delicate sterile scales. W.
setosa is distinguished from other species of Weltrichia by
the inward pointing scales at the top of the cup and by the
toothed bristles or hairy ramenta on the edges of the rays
(microsporophylls).
In the general morphological features the new species
may also be comparable with the type species W. mirabilis
Braun (1849), but they are different in many details. Braun
originally described three species, but he realized the pos-
sibility that in a different type, W. mirabilis which bears
funnel-shaped structures, the lower part of which has the
form of a complete cup made of the concrete bases of about
20 broadly linear segments and the upper part are sepa-
rated into lanceolate lobes each with a midrib and slightly
curved inwards at the apex. The whole, nearly 10 cm long
and 9 cm in diameter at the upper edge, is very similar to the
specimen of the W. spectabilis (Seward, 1917, P. 437,438;
Figs. 551,552). In the latter the cup divides at its top into
about 13 rays, typically 10 mm wide in lower part, tapering
to 3 mm, at 3.3 cm from the cup and then continuing as a
filiform extension curved inwards for about 3 cm. Rays some-
times show midribs. The inner face of rays bears fertile
branches; That are borne in a longitudinal row on either
side of the midrib and at intervals of 3-5 mm. Fertile branches
bear semicircular or broadly elongated pollen sacs that are
morphologically different from the new species.
Weltrichia huangbanjigouensis Sun et Zheng emend
Zheng
2001, Weltrichia huangbanjigouensis Sun et Zheng, P.
82,190; Pl. 12, Fig.3; Pl. 48, Figs.1-3.
Emended diagnosis A Bennettitalean male strobile,
funnel- or cup-shaped impression, borne on top of a small
shoot, about 2 cm long by 4 mm wide. Cup about 2 cm tall
and about 3.5 cm wide in apex; many ovoid resinous sacs
scattered on the inner surface of the cup. Pollen sacs and
pollen grains are unknown.
Discussion This species was established by Sun et
Zheng based on a few specimens discovered near
Huangbanjigou from Upper Jurassic-Lower Cretaceous
lower part of Yixian Formation. It is a representative funnel-
shaped structure, its lower part is in a shape of incomplete
cup made of the truncate bases of rays (microsporophylls)
and pollen sacs were not preserved. But in the inner sur-
face of the cup there are many ovoid, very small impres-
sions very similar to those resinous sacs of Weltrichia sol
Harris (1969, P. 164, Fig.69E) which had been previously
described by Sun et al. (2001, P. 82, 190; Pl. 12, Fig.3; Pl. 48,
Figs.1-3) as “pollen sacs”. In fact they are some resinous
sacs, but we believe that it belongs to the genus Weltrichia.
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