Content uploaded by Toshihiro Yamada
Author content
All content in this area was uploaded by Toshihiro Yamada on Jan 13, 2015
Content may be subject to copyright.
87
GEODIVERSITAS • 2011 • 33 (1)
© Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com
Legrand J., Pons D., Nishida H. & Yamada T. 2011. — Barremian palynofl oras from the
Ashikajima and Kimigahama formations (Choshi Group, Outer Zone of south-west Japan).
Geodiversitas 33 (1): 87-135. DOI: 10.5252/g2011n1a6.
ABSTRACT
e Choshi Group, which crops out in the Outer Zone of south-west Japan, has
been extensively studied for its rich macrofl ora by Makoto and Harufumi Nishida,
among others, and was attributed to the Ryoseki-type Floristic Province by Kimura
(1987). New microfl oras were discovered in muddy, very fi ne-grained sandstones
and mudstones of the marine Ashikajima and Kimigahama formations, represent-
ing the base of the Choshi Group. e authors provide a palynological inventory
for these lithological units, which have been dated as Barremian on the basis of
the ammonites recorded from them, and compare them with the paleofl oristic
associations of the South-Laurasian Province (Brenner 1976) and Euro-Sinian
Region (Vakhrameev 1991). e studied assemblages yielded 53 genera and 89
species of spores and gymnosperm pollen grains, and also marine or freshwater al-
Julien LEGRAND
Denise PONS
Université Pierre et Marie Curie (UPMC),
UMR 7207 CNRS, Centre de Recherche
sur la Paléobiodiversité et les Paléoenvironnements,
Muséum national d'Histoire naturelle, case postale 48,
57 rue Cuvier, F-75231 Paris cedex 05 (France)
jlegrand@snv.jussieu.fr
denise.pons@snv.jussieu.fr
Harufumi NISHIDA
Chuo University,
Faculty of Science and Engineering,
Department of Biological Sciences,
1-13-27 Kasuga, Bunkyo, 112-8551 Tokyo (Japan)
helecho@kc.chuo-u.ac.jp
Toshihiro YAMADA
Kanazawa University,
Graduate School of Natural Science and Technology,
Division of Life Sciences,
Kakuma, 920-1192 Kanazawa (Japan)
tyamada@kenroku.kanazawa-u.ac.jp
Barremian palynofl oras from the Ashikajima
and Kimigahama formations (Choshi Group,
Outer Zone of south-west Japan)
88
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
INTRODUCTION
e backbone of the Japanese archipelago was
largely formed during the Early Cretaceous period.
Before the opening of the Sea of Japan during the
Miocene (28 My), the Inner and Outer Zones
of present-day Japan (Fig. 1) were separated by
a marine basin. e Inner Zone (north-western
MOTS CLÉS
Palynologie,
Barrémien,
Flore Ryoseki,
Groupe Choshi,
SO Japon,
espèces nouvelles.
KEY WORDS
Palynology,
Barremian,
Ryoseki-type Flora,
Choshi Group,
SW Japan,
new species.
gae and some epiphyllous fungi. No angiosperm pollen grain was observed. Four
new species are described: Manumia japonica n. sp., Foveosporites ryosekiensis n. sp.,
Nodosisporites choshiensis n. sp. and N. makotoi n. sp. Other forms, probably new
species, are described here in detail, but the scarcity of the specimens has led us to
place them temporarily in open nomenclature. e spatio-temporal distributions
of the genus Manumia Pocock, reported for the fi rst time in Asia, and Cicatrico-
sisporites sinuosus Hunt, 1985 are plotted on paleogeographical maps. With this
palynological study, we add new data to the present knowledge of Barremian fl oras.
is assemblage probably corresponds to a taphocenose. e authors suggest that
the climate indicated had marked dry and more humid seasons, in accordance with
the hypothesis of a moderate migration of the oceanic islands of the Outer Zone
before their collision with the Eurasian continent, or moderate climatic change
during the Early Cretaceous in Japan.
RÉSUMÉ
Palynofl ores barrémiennes des Formations Ashikajima et Kimigahama (Groupe
Choshi, Zone Externe du sud-ouest du Japon).
Le Groupe Choshi, appartenant à la Zone Externe du sud-ouest du Japon, est
bien connu pour sa riche macrofl ore étudiée entre autres par Makoto et Harufumi
Nishida, et placée dans la Province Floristique « Ryoseki » par Kimura (1987). De
nouvelles microfl ores ont été découvertes dans des grès argileux à grain très fi n et
argiles compactées provenant des Formations marines Ashikajima et Kimigahama
situées à la base du Groupe Choshi. Les auteurs font un inventaire palynologique
de ces unités lithologiques datées du Barrémien par les ammonites, et les compa-
rent avec des associations paléofl oristiques connues de la Province sud-laurasienne
(Brenner 1976) et Région euro-sinienne (Vakhrameev 1991). Les assemblages nous
ont livré 53 genres et 89 espèces de spores et grains de pollen de gymno spermes,
ainsi que des algues marines ou d’eau douce et quelques champignons épiphylles.
Aucune angiosperme n’a été observée. Quatre nouvelles espèces sont défi nies :
Manumia japonica n. sp., Foveosporites ryosekiensis n. sp., Nodosisporites choshiensis
n. sp. et N. makotoi n. sp. D’autres formes, probablement de nouvelles espèces,
sont décrites ici en détails, mais la rareté des individus nous a conduit à les placer
provisoirement en nomenclature ouverte. Les répartitions spatio-temporelles du
genre Manumia Pocock, signalé en Asie pour la première fois, et de Cicatricosisporites
sinuosus Hunt, 1985 sont replacées sur des cartes paléogéographiques. Cette première
étude palynologique permet de compléter la connaissance des fl ores du Barrémien.
Cet assemblage correspond probablement à une taphocénose. Il indique un climat
à saison chaude marquée suivie d’une saison plus humide, ce qui est conforme avec
les hypothèses d’une migration modérée des îles océaniques de la Zone Externe
avant leur collision avec le continent eurasiatique, ou de changements climatiques
modérés au cours du Crétacé inférieur au Japon.
89
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
part, along the Sea of Japan) was connected to
the east of the Korean Peninsula and represented
the margin of the Eurasian continent. e Outer
Zone (south-eastern part, along the Pacifi c Ocean)
of south-west Japan was represented by oceanic is-
lands located further to the south. Due to sinistral
strike-slip movements along the Median Tectonic
Line (MTL), which began in the Late Jurassic,
these islands moved northwards and reached their
present latitude to collide with the Inner Zone
until the close of the Hauterivian (Yaskawa 1975;
Hirooka et al. 1983, 1985; Matsukawa & Obata
1993; Matsuoka et al. 1997; Otoh 1998; Otoh &
Sasaki 1998; Matsukawa & Fukui 2009). It should
be noted, however, that there are many diff erent
interpretations of the speed and period of their
migration and on the position of the Outer Zone
before the strike-slip motion of the plates (Otofuji
et al. 1985; Kojima 1989; Mitsugi et al. 2001;
Ishida et al. 2003; Lee & Kim 2005; Lee 2008).
Some authors even doubt the existence of the ter-
ranes constituting the Outer Zone (Isozaki 1996,
1997; Maruyama et al. 1997; Isozaki et al. 2010).
us, the developmental history of the Japanese
archipelago is still open to question.
e paleofl ora of the Outer Zone is called “Ry-
oseki-(Gondwanian-)type”. It is widely accepted
that it fl ourished under a seasonally dry climate
based on the abundance of microphyllous conifers
and bennettitoid foliage such as Zamites Brongniart
emend. Harris and Ptilophyllum Morris (Kimura
1958, 1987). is fl ora is markedly diff erent from
the warm temperate “Tetori-(Siberian-)type” fl ora
found in the Tetori Group of the Inner Zone,
which is characterized by abundant ferns, ginkgos
and macrophyllous conifers (Kimura & Sekido
1976, 1978; Kimura 1987; Kimura & Ohana
1997). Such pronounced phytogeographical dif-
ferences are often referred by proponents of the
strike-slip hypothesis as evidence supporting the
low-latitudinal origin of the Outer Zone terranes
(Fig. 2). However, similar age of these fl oras has
been increasingly challenged by recent stratigraphic
and paleobotanical studies (Yamada & Uemura
2008; Yamada 2009). Furthermore, the paleocli-
mates noted above were inferred empirically from
the foliar fossils, although these are often diffi cult
to classify even at the generic level because of the
diagenetic loss of their cuticle in Japanese Upper
Jurassic to Lower Cretaceous sediments.
Palynological studies provide stratigraphic and
climatic clues, but few palynological investigations
have been carried out on Lower Cretaceous sedi-
ments in Japan until now (e.g., Umetsu & Matsuoka
2003; Umetsu & Sato 2007). We describe here
palynomorphs from the upper part of the Ashika-
jima Formation and lower part of the Kimigahama
Formation of the marine Choshi Group, located
in the most eastern part of the Chiba Prefecture,
south-west Honshu. e Choshi Group (Shikama &
Suzuki 1972) represents Lower Cretaceous sediments
in south-west Japan and is dated as Barremian to
early Albian based on the ammonite and other mol-
luscan assemblages (Hayami & Oji 1980; Kase &
Maeda 1980; Obata & Matsukawa 2009). is paper
presents the fi rst comprehensive description of a
pre-Aptian palynological assemblage for the Outer
Zone. We compare the assemblage to coeval paly-
nofl oras in adjacent areas to infer the paleoclimate
of the Ryoseki-type fl ora in more detail.
South-
West
North-
East
Inner Zone
Outer Zone
1
40°N
35°N
130°E 140°E
45°N
Tectonic lineCountry boundary
3
200 km
2
F
IG
. 1. — Geological setting of Japan: 1, Itoigawa-Shizuoka tectonic
line; 2, Kashiwazaki-Choshi tectonic line; 3, Median tectonic line.
Map modifi ed from Geological Survey of Japan (2010).
90
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
GEOLOGICAL SETTING
e Lower Cretaceous strata of the Choshi Group
crop out along the eastern coast of the Choshi Pe-
ninsula, Chiba Prefecture (Fig. 3). is group has
been subdivided lithologically into fi ve formations,
in ascending order: the Ashikajima, Kimigahama,
Inubouzaki, Toriakeura and Nagasakihana for-
mations (Obata et al. 1975, 1982; Kase & Maeda
1980). Type biozones were established for the
ammonites of the Lower Cretaceous of south-
west Japan and enabled the age of the Choshi
Group to be determined as Barremian to early
Albian (Obata et al. 1975; Obata & Matsukawa
2009). It was deposited in an off shore to shallow
marine environment (Katsura et al. 1984; Ito &
Matsukawa 1997). Plant macrofossils have been
described from the Ashikajima, Kimigahama, and
Toriakeura formations and contain typical species
of the Ryoseki-type fl ora, such as the cyatheaceous
fern Ptilophyllum cutchense Morris, and the gymno-
sperm Brachyphyllum expansum (Sternberg) Seward
(e.g., Nishida 1960, 1962; Kimura & Ohana 1985;
Kimura et al. 1991).
e Ashikajima Formation overlies unconform-
ably the pre-Cretaceous rocks, and is overlain con-
formably by the Kimigahama Formation (Fig. 4).
Its base is mainly composed of conglomerates with
South Chichibu terrane
Abukuma and South
Kitakami terranes
Tamba, Mino and
Ashio terranes
138°126°
44°
36°
28°
20°
A
500 km
N
Localities of Early Cretaceous floras: Tetori-type; Mixed-type; Ryoseki-type.
coastlines
faults
floras boundaries
pre-Jurassic Asian continental margin
Middle-Late Jurassic terranes
Early Cretaceous terranes
Late Cretaceous terranes
pre-Cambrian (?) -
Early Paleozoic terranes
Choshi
locality
130° 140°
500 km
Mixed-
type Flora
Tetori-type
Flora
Ryoseki-type
Flora
Choshi
locality
I
n
n
e
r
Z
o
n
e
O
u
t
e
r
Z
o
n
e
B
M
i
x
e
d
-
t
y
p
e
Flora
40°
34°
Mixed-type Flora
Tetori-type Flora
Ryoseki-type Flora
FIG. 2. — Paleofl oristic provinces in Japan and eastern Asia during the Late Jurassic-Early Cretaceous period: A, on a paleogeo-
graphical map, which illustrates the relative locations of the Inner and Outer Zones (modifi ed from Golozoubov et al. 1999); B, on a
present map (from Kimura 1987).
91
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
°°
°°
°
vv
Naarai Fm - Miocene
Nagasakihana Fm - early Albian
Toriakeura Fm - latest Aptian
Kimigahama Fm - Barremian
Ashikajima Fm - early Barremian
.
..
Inubouzaki Fm - early Aptian
.
.
.
.
.
.
.
.
.
Atagoyama Group - ~ Jurassic
faults
Choshi Group
°
°
clastic rocks - post-Miocene
no geological data
°
°
°
°
°
°
°
°
°
°
°
°
°
°
°
°
°
°
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
°
°
°
°
v
v
v
°
Ashikajima
Kimigahama
Isejigaura
Toriakeura
Nagasakihana
.
Kimi I1
Ashi II2
.
.
.
.
°
°
.
v
v
v
v
v
v
v
v
v
v
v
v
°
°
°
°
°
°
°
°
°
°
°
°
.
. . . . . . . .
. . . . . . . .
. . . . . . . . .
. . . . . . . .
. . . . . .
. . .
. . .
.
. .
1500 m
N
v
v
v
v
v
v
v
Choshi
Peninsula
140°E
36°N
50 km
141°E
35°5'N
Japon
Jap
Jap
Jap
Jap
Jap
Jap
Jap
p
p
on
JAPAN
RUSSIA
Sakhalin
CHINA
KOREA
Primorye
Hokkaido
Honshu
Shikoku
Kyushu
Chiba
Prefecture
50°N
40°N
30°N
130°E 140°E 150°E
1000 km
e Kimigahama Formation is overlain uncon-
formably by the Inubouzaki Formation. It is mainly
composed of alternating beds of sandstones and
mudstones, containing many calcareous nodules.
e foraminifer and ammonite assemblages have
enabled the assignation of a Barremian age to the
formation (Obata et al. 1975; Hayami & Oji 1980;
Kase & Maeda 1980; Obata & Matsukawa 2009).
We collected our samples from the mudstones along
Kimigahama beach (35°43’N, 140°52’E).
siliceous pebbles, above which are coarse-grained
sandstones with intercalated mudstones, becoming
thinner towards the top (Obata et al. 1975, 1982;
Obata & Matsukawa 2009). e formation is about
200 m thick. It is dated as Barremian based on the
ammonite and belemnite assemblages (Obata et al.
1975; Hayami & Oji 1980). We took our samples
from the muddy, very fi ne-grained sandstones of
the Hatoyama locality, in the south-eastern part of
the Choshi Peninsula (35°41’N, 140°51’E).
FIG. 3. — Location map of the outcrops and geology of the eastern coast of the Choshi Peninsula (modifi ed from Obata et al. 1975,
1982; Ito & Matsukawa 1997; Obata & Matsukawa 2009).
92
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
MATERIAL AND METHODS
e samples, fi rst cleaned under running water and
scrubbed with brush, were left for a few days in the
open air and then broken in a mortar. e ground
samples were sieved and the fraction between 1 and
3 mm in grain diameter was soaked to 10% aqueous
hydrochloric acid until the end of the eff ervescence to
remove calcareous materials. After washing in water,
the samples were submerged in 70% hydrofl uoric
acid for one day and washed again. e residues were
then decanted, covered by a small amount of 10%
hydrochloric acid and heated. When boiling point
was reached they were cooled and then washed. After
sieving, the fraction of each sample with a diameter
between 5 and 100 μm was kept, diluted in water
and submitted to treatment with nitric acid while
heating in a double boiler. e residue was sieved
again, and the fraction with a diameter up to 5 μm
was decanted into a test tube and centrifuged for 10
minutes at 1200 revolutions per minute. e residue
was then fi xed on a slide using a fi lm of Cellosize and
mounted in Canada balsam.
Photographs of the palynomorphs were obtained
using a Nikon D300 digital camera linked to a Nikon
Eclipse 80i microscope, and SEM micrographs were
obtained using a Hitachi TM-1000 tabletop micro-
scope. e formation, the level, the number of the slide
and the graticule (England Finder) are indicated in the
legend for each specimen illustrated in this paper.
Since it is diffi cult to compare the fossil spores
and pollen grains to genera of plants living today, we
followed the classifi cation of sporae dispersae defi ned
by Potonié & Kremp (1954, 1955). For some paly-
nomorphs, we followed slightly modifi ed version of
Potonié & Kremp (1954, 1955) by Dettmann (1963),
Potonié (1956, 1958, 1960, 1966, 1970a, b, 1975),
and Pfl ug (1953). ese classifi cations are based on the
external morphology. However, we tried to determine
the presumed botanical affi nities when possible.
SYSTEMATIC DESCRIPTIONS
We present species that have been identifi ed dur-
ing this investigation (Table 1). e preparations
are housed in the Collection de Paléobotanique
of the Université Pierre et Marie Curie (UPMC),
Paris (France). Only new species and taxa in open
nomenclature are described.
Anteturma PROXIMEGERMINANTES
Genus Converrucosisporites Potonié & Kremp, 1954
TYPE SPECIES. — Converrucosisporites triquetrus (Ibrahim,
1933) Potonié & Kremp, 1954.
Converrucosisporites sp.
(Fig. 5N, O)
Converrucosisporites sp. A, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 136,
pl. III, fi g. 2 (2009).
O
CCURRENCE. — Ashikajima Fm (rare).
DESCRIPTION
Trilete microspore. Amb rounded triangular. e
laesurae are long, rather indistinct, and seem to
extend to the 3/4 of the spore radius. e two faces
are covered by rounded verrucae of various sizes
(0.5 to 3 μm high and 1 to 6 μm in diameter),
distributed side by side or sometimes contiguous.
Near the equator, some elongated verrucae can be
sparsely observed. e exine is about 0.5 μm thick.
Equatorial diameter = 30-35 μm.
BOTANICAL AFFINITIES
Cyatheales?
Genus Manumia Pocock, 1970
TYPE SPECIES. — Manumia verrucata Pocock, 1970.
Manumia japonica
Legrand, Pons, Nishida & Yamada n. sp.
(Fig. 6A-F, I, J)
Manumia sp. A, Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
93
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
paléofl oristiques du sud-est asiatique: 138, 139, pl. III,
fi gs 8-10 (2009).
T
YPE MATERIAL. — Site II, horizon 2, slides b, g; slide
SEM-b; holotype (II2g-O68; Fig. 6C, F), paratypes
(SEM-II2b, SEM-II2b, II2b-N21/4, II2c-K65g; Fig. 6A,
B, D, E, I, J).
Collection de Paléobotanique - UPMC, Paris, France.
E
TYMOLOGY. — e species name is after Japan, where
the Choshi Group is located.
OCCURRENCE. — Ashikajima and Kimigahama Fm.
T
YPE
LOCALITY
. — Hatoyama, SE Choshi Peninsula,
Chiba Prefecture, Japan.
S
TRATIGRAPHIC
H
ORIZON
. — Ashikajima Fm (Bar-
remian).
DIAGNOSIS
Trilete microspore. Amb rounded triangular. e
laesurae are straight and extend to the 3/4 of the
spore radius. ey are bordered by raised lips with
irregular margins (3 to 7 μm wide) that can be
continuous in the apices areas. e ornamenta-
tion of the exine is scabrate to granulate, with
isolated verrucae (3-7 μm in diameter) irregularly
distributed along the laesurae of the proximal
face, wider (3-7 μm) in the interradial areas. At
the equator, some circular or elongated verrucae
are randomly distributed near the angles or be-
tween the laesurae. On the distal face, verrucae
are more or less densely distributed. e exine
is thick (about 2-3 μm). Equatorial diameter =
50-55 μm.
R
EMARKS
A wide range is noted in the ornamentation of this
new species. e laesurae can extend to the 3/4 of
the spore radius or nearly to the equator. On the
proximal face of the largest specimens (Fig. 6A, D),
verrucae can sometimes be coalescent in the inter-
radial areas to form ridges parallel to the equator.
At the equator also, the elongated verrucae can
gather to form an equatorial thickening (Fig. 6D,
I). On the distal face, depending of the specimen,
verrucae or tubercules can be isolated (Fig. 6F) or
more or less densely distributed and coalescent
(Fig. 6B, J). e equatorial diameter ranges from
40 to 55 μm.
e genus Manumia is generally characterized by
strong equatorial thickenings and/or coarse tuber-
cules more or less fused into ridges. e faces are
unequal, with diff erent ornamentations.
Barremian
Ashikajima Fm
Kimigahama Fm
Choshi Group
Kimi I1
Ashi II2
Aptian
185 m
190 m
Lower Cretaceous
pre-Cretaceous
formations
Inu.
Fm
. . . .
. . . . .
sandstones
mudstones
unconformity
conglomerates
siltstones
palynofloras
plant remains
ammonites
bivalves
dinoflagellates
50 m
FIG. 4. — Synthetic stratigraphical column of the Ashikajima and
Kimigahama formations, between the pre-Cretaceous formations
and the Inubouzaki Formation (modifi ed from Obata et al. 1975;
Obata & Matsukawa 2009).
94
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
Four species have been defi ned for the genus
Manumia: M. delcourtii (Pocock, 1970) Dybkjær,
1991, M. irregularis Pocock, 1970, M. variverrucata
(Couper, 1958) Hoelstad, 1985, and M. verrucata
Pocock, 1970. e Japanese species can be distin-
guished from the other species of the genus by the
shape and distribution of its verrucae. Concerning
M. variverrucata, Hoelstad (1985) defi ned it on the
basis of an emendation and new combination of
Concavisporites variverrucatus Couper, 1958. How-
ever, as the diagnosis made by Couper (1958) does
not fi t that made by Pocock (1970) for the genus
Manumia, particularly about the ornamentation,
we can suggest that the defi nition of a new spe-
cies would have been more suitable than the new
combination proposed by Hoelstad (1985).
is spore genus was previously reported from
the Jurassic of Canada (Pocock 1970), Greenland
(Lund & Pedersen 1984; Koppelhus & Hansen
2003), Alaska (Bjærke 1993), and northern Europe
(Couper 1958; Schulz 1967; Guy 1971; Hern-
green & de Boer 1974; Hoelstad 1985; Dybkjær
1991; Koppelhus 1992; Seidenkrantz et al. 1993;
Koppelhus & Nielsen 1994; Koppelhus & Dam
2003; Bøe et al. 2005; Lindström & Erlström 2007;
Stefanowicz 2008), and the Lower Cretaceous of
Norway (Bøe et al. 2005). It was also reported
from the Lower to Middle Jurassic of Afghanistan
(Schweitzer et al. 1987). It seems to characterize
the South Laurasian Province of Brenner (1976).
Legrand (2009: 139, pl. III, fi g. 11) distinguished
a second form, morphologically similar but smaller
(30-35 μm) and with a thinner exine, that could
correspond to a variation of M. japonica.
B
OTANICAL AFFINITIES
Even if the ornamentation suggests an affi nity with
the present Polypodiales (Pteridaceae), particularly
Pteris Linnaeus or Pityrogramma Link fi gured by
Tryon & Lugardon (1991), the absence of any
cingulum should distinguish them.
Genus Uvaesporites Döring, 1965
TYPE SPECIES. — Uvaesporites glomeratus Döring, 1965.
Uvaesporites sp.
Uvaesporites sp. B, Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et prov-
inces paléofl oristiques du sud-est asiatique: 141, pl. IV,
fi g. 4 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb circular. e laesurae are
thin, straight, slightly raised, and extend to the cingu-
lum. Proximal face psilate. e equator is ornamented
by verrucae joined at their base (3-4 μm high and
3-7 μm in basal diameter). e distal face is covered
by verrucae of various sizes, bigger near the equa-
tor than at the pole, often joined, making irregular
and very small closed spaces. Equatorial diameter =
28 μm; width of the «cingulum» = 3-4 μm.
B
OTANICAL AFFINITIES
Cyatheales (Cyatheaceae) or Lycopsida (Selaginel-
laceae) (Takahashi 1988).
Genus Baculatisporites
Pfl ug & omson in omson & Pfl ug, 1953
T
YPE
SPECIES
. — Baculatisporites primarius (Wolff , 1934)
Pfl ug & omson in omson & Pfl ug, 1953.
Baculatisporites sp.
(Fig. 6N)
Baculatisporites sp. B, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
FIG. 5. — Spores from the Barremian of south-west Japan: A, B, Biretisporites potoniaei (Delcourt & Sprumont, 1955) Delcourt, Dett-
mann & Hughes, 1963 (A, proximal face, Ashikajima Fm, II2a-U31/3; B, proximal face, Kimigahama Fm, SEM-I1a); C, Todisporites
miser Takahashi, 1988, proximal face, Ashikajima Fm, II2b-R25hg; D, Todisporites major Couper, 1958, proximal face, Ashikajima Fm,
II2c-H34/1; E, Incertae sedis in Hasenboehler (1981), proximal face, Ashikajima Fm, II2a-Q62/3; F, Cyathidites minor Couper, 1953,
proximal face, Ashikajima Fm, IIa-V64; G, Cyathidites australis Couper, 1953, proximal face, Ashikajima Fm, SEM-II2a; H, Cyathidites
rarus (Bolkhovitina, 1953) Deák, 1964, proximal face, Ashikajima Fm, II2a-G59/1; I, J, Verrucosisporites densus (Bolkhovitina, 1956)
95
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
A
E
H
I
J
M
P
L
CDB
G
K
Q
N
O
F
Pocock, 1970, proximal and distal faces, Ashikajima Fm, II2h-H49g; K, Concavissimisporites variverrucatus (Couper, 1958) Brenner,
1963, proximal face, Ashikajima Fm, II2c-K43-1; L, Osmundacidites wellmanii Couper, 1953, proximal face, Ashikajima Fm, II2b-P32/4;
M, Trilites bossus Couper, 1958, Ashikajima Fm, II2f-H29/1; N, O, Converrucosisporites sp. (N, Ashikajima Fm, II2f-V27/3; O, Ashikajima
Fm, SEM-II2c); P, Cibotiumspora paradoxa (Maljavkina, 1949) Chang, 1965, proximal face, Kimigahama Fm, I1a-F40-g; Q, Concavis-
simisporites punctatus (Delcourt & Sprumont, 1955) Brenner, 1963, proximal face, Ashikajima Fm, SEM-II2c. Scale bar: 10 μm.
96
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
provinces paléofl oristiques du sud-est asiatique: 142-143,
pl. IV, fi g. 8 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb circular. e laesurae are
straight and extend to the 3/4 of the spore radius.
e contact area is psilate. Small verrucae, coni and
bacula (less than 1 μm long) are sparsely distributed
near the equator and on the distal face. e exine is
1 μm thick. Equatorial diameter = 40-45 μm.
B
OTANICAL AFFINITIES
Osmundales, Osmundaceae (Dettmann 1963, 1994).
Genus Neoraistrickia Potonié, 1956
TYPE SPECIES. — Neoraistrickia truncatus (Cookson,
1953) Potonié, 1956.
Neoraistrickia sp. 1 (Fig. 7A, B)
Neoraistrickia sp. A, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 143,
pl. IV, fi g. 12 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb rounded triangular. e
laesurae are straight, slightly raised, and extend to
the 3/4 of the spore radius. e two faces are densely
covered by rounded to slender bacula mingled with
some sharpened coni (3 to 7 μm high). e exine
is 1.5 to 2 μm thick. Equatorial diameter = 50 μm
with the ornamentation.
B
OTANICAL AFFINITIES
Bryophyta or Lycopsida (Selaginellaceae)?
Neoraistrickia sp. 2 (Fig. 7C, D)
Neoraistrickia sp. B, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 144,
pl. IV, fi g. 14 (2009).
O
CCURRENCE. — Ashikajima Fm (rare).
DESCRIPTION
Trilete microspore. Amb circular. e laesurae are
straight and extend to the ⅔ of the spore radius.
e contact area is psilate, followed by more or less
coarse bacula and some elongated, sometimes joined
(particularly at the equator) coni near the equator
and on the distal face. e exine is about 1.5 μm
thick. Equatorial diameter = 45-50 μm.
B
OTANICAL AFFINITIES
Bryophyta or Lycopsida (Selaginellaceae)?
Neoraistrickia sp. 3 (Fig. 7G, J)
Neoraistrickia sp. C, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 144,
pl. IV, fi gs 11, 15 (2009).
O
CCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Trilete microspore. Amb circular. e laesurae cannot be
clearly seen; they seem short and raised in a very small
contact area. e two faces are densely covered by more
or less short and sinuous rugulae topped by bacula,
coni, spines or tubercules (4 to 7 μm high), giving to
the spore a bristling appearance. e exine is about
3 μm thick. Equatorial diameter = 35-45 μm.
R
EMARK
e ornamentation of Neoraistrickia sp. 3 has simi-
larities with that of N. sp. 2, but can be distinguished
from the later by its dense and bristling appearance
and its thicker exine.
B
OTANICAL AFFINITIES
Bryophyta or Lycopsida (Selaginellaceae)?
Genus Tuberositriletes Döring, 1964
TYPE SPECIES. — Tuberositriletes montuosus Döring, 1964.
97
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
AB
G
K
H
C
F
I
J
N
L
M
ED
FIG. 6. — Spores from the Barremian of south-west Japan: A-F, I, J, Manumia japonica n. sp. (A, proximal face, Ashikajima Fm, SEM-
II2b; B, distal face, Ashikajima Fm, SEM-II2b; C, F, proximal and distal faces, Ashikajima Fm, II2g-O68; D, E, proximal and distal faces,
Ashikajima Fm, II2b-N21/4; I, J, proximal and distal faces, Ashikajima Fm, II2c-K65g); G, Pilosisporites verus Delcourt & Sprumont,
1955, distal face, Ashikajima Fm, SEM-II2c; H, L, M, Tuberositriletes sp. (H, proximal face, Ashikajima Fm, SEM-II2a; L, M, proximal
and distal faces, Ashikajima Fm, II2b-U45/4); K, Pilosisporites brevis Delcourt & Sprumont, 1955, proximal face, Ashikajima Fm, SEM-
II2c; N, Baculatisporites sp., proximal face, Ashikajima Fm, II2b-Z40/3. Scale bar: 10 μm.
98
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
Tuberositriletes sp.
(Fig. 6H, L, M)
OCCURRENCE. — Ashikajima Fm (rare).
DESCRIPTION
Trilete microspore. Amb triangular with concave
sides. e laesurae extend to the 3/4 of the spore
radius. On the proximal face, the contact area is
psilate, followed by one row of big welded ver-
rucae (2-4 μm high, 2.5-5 μm in basal diameter)
near the equator. e distal face is ornamented
by welded and wavy rugulae, which are topped
by verrucae. e rugulae are separated by small
irregular spaces that can connect. Equatorial di-
ameter = 28 μm.
B
OTANICAL AFFINITIES
Filicopsida.
Genus Cicatricosisporites
Potonié & Gelletich, 1933 emend. Potonié, 1966
TYPE SPECIES. — Cicatricosisporites dorogensis Potonié &
Gelletich, 1933.
Cicatricosisporites sinuosus Hunt, 1985
(Fig. 8K, L, S, T)
Pollen et Spores 27: 427-428, 430, pl. 2, fi gs 1-3
(1985). — Omran et al., Review of Palaeobotany and
Palynology 66: 301, pl. 1, fi g. 4 (1990). — Schrank &
Ibrahim, Berliner Geowissenschaftliche Abhandlungen,
Reihe A 177: 16, 17, pl. 1, fi g. 14 (1995). — Ibrahim,
Review of Palaeobotany and Palynology 94: 151 (1996). —
Al-Ameri et al., Cretaceous Research 22: 738 (2001). —
Ibrahim et al., Journal of African Earth Sciences 32 (2):
276 (2001). — Mahmoud & Deaf, Rivista Italiana di
Paleontologia e Stratigrafi a 113 (2): 218 (2007). — Mah-
moud et al., Revista Española de Micropaleontología 39
(3): 176, pl. 5, fi g. 14 (2007). — Mejia Velasquez, Floral
composition of a Lower Cretaceous paleotropical ecosystem
inferred from quantitative palynology: 60, pl. VIII, fi g. 81
(2007). — Peyrot et al., Revista Española de Micropaleon-
tología 39 (1-2): 142 (2007). — Legrand, Palynologie des
dépôts Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 156, 157,
pl. VIII, fi gs 2-4 (2009).
Camarozonosporites insignis Norris, 1959, Saad, Pollen et
Spores 20 (2): 274, pl. III, fi gs 5, 6 (1978).
Reticulisporites sp. sensu Batten & Uwins (1985), Bat-
ten & Uwins, Journal of Micropalaeontology 4 (1):
pl. 67, fi gs 1, 2 (1985). — Uwins & Batten, Subsurface
Palynostratigraphy of Northeast Libya: 219, 224, pl. 43,
fi g. 12 (1988). — Masure et al., Géologie Méditer-
ranéenne 25 (3-4): 273, pl. 2, fi g. 20 (1998).
Reticulisporites sp., Pons et al., Géologie de l’Afrique et de
l’Atlantique Sud : Compte rendu des Colloques de géologie
d’Angers: 390 (1996).
Cicatricosisporites daxinganlingensis Pu & Wu, 1985,
Umetsu & Sato, Review of Palaeobotany and Palynol-
ogy 144 (1-2): pl. 1, fi g. 7 (2007).
OCCURRENCE. — Ashikajima and Kimigahama Fm
(common).
D
ISTRIBUTION
. — North Tethys. Upper Jurassic to
Lower Cretaceous of England (Hunt 1985). is species
is known in Japan under the name Cicatricosisporites dax-
inganlingensis Pu & Wu, 1985 (Aptian-Albian, Umetsu &
Sato 2007). It was also reported from the early Aptian
of France (Masure et al. 1998), and late Aptian-early
Albian of Spain (Peyrot et al. 2007).
South Tethys. is species is known in the ?late Hau-
terivian-?middle Barremian and early to middle Al-
bian of Libya under the name Reticulisporites sp. sensu
Batten & Uwins (1985) (Uwins & Batten 1988). It
was also reported in the Barremian of Lebanon (De-
jax, personnal communication), early Aptian of Co-
lombia (Mejia Velasquez 2007), late Aptian of Brazil
(Pons et al. 1996, under the name Reticulisporites sp.),
Aptian-Albian of Egypt (Saad 1978, under the name
Camarozonosporites insignis Norris, 1959; Omran et al.
1990; Schrank & Ibrahim 1995; Ibrahim 1996; Ibra-
him et al. 2001; Mahmoud & Deaf 2007; Mahmoud
et al. 2007), Albian of Tunisia (Pons, personnal com-
munication), and Albian-early Cenomanian of Iraq
(Al-Ameri et al. 2001).
DESCRIPTION
Trilete microspore. Amb rounded triangular. e
laesurae are straight, bordered by lips (1-1.5 μm
wide), and extend to the 3/4 of the spore radius. Ex-
ine striate. Striae (0.5-1.5 μm wide; 0.5-1 μm high)
are separated by spaces 0.5 to 3 μm wide; they are
oblique to perpendicular to the equator, more or less
sinuous and parallel, can bifurcate, and run irregularly
on both faces. On the proximal face, each interradial
area contains 4 to 9 striae. e exine is about 1 μm
thick. Equatorial diameter = 20-35 (40) μm.
99
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
F
IG
. 7. — Spores from the Barremian of south-west Japan: A, B, Neoraistrickia sp. 1, proximal and distal faces, Ashikajima Fm, II2c-
U62/3d; C, D, Neoraistrickia sp. 2, proximal and distal faces, Ashikajima Fm, II2c-H33/4; E, F, I, L, M, Klukisporites variegatus Couper, 1958
(E, proximal face, Kimigahama Fm, SEM-I1a; F, distal face, Kimigahama Fm, SEM-I1a; I, L, proximal and distal faces, Kimigahama Fm,
I1a-H48/1a; M, proximal face, Ashikajima Fm, SEM-II2c); G, J, Neoraistrickia sp. 3, proximal and distal faces, Kimigahama Fm, I1b-S68g;
H, K, N-P, Lycopodiumsporites dentimuratus Brenner, 1963 (H, proximal face, Kimigahama Fm, SEM-I1c; K, N, proximal and distal faces,
Ashikajima Fm, II2f-M56/2; O, lateral view, Kimigahama Fm, SEM-I1a; P, lateral view, Ashikajima Fm, II2e-R56/2). Scale bar: 10 μm.
A
E
H
K
N
O P
L
M
I
J
FG
B CD
100
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
REMARKS
Hunt (1985) attributed this species to Cicatrico-
sisporites. However, it diff ers from all other species
already described in this genus, by its striae per-
pendicular to the equator on the proximal face.
De Haan (1997) proposed in his thesis a new
form-genus, Huntisporis, and a new combination,
Huntisporis sinuosus (Hunt, 1985) de Haan, 1997,
but both are not valid because unpublished.
Cicatricosisporites curvatus Pu & Wu, 1982 (417,
pl. 8, fi gs 19, 20), reported from the ?Valanginian-
Barremian of the Heilongjiang Province of North
China, has a similar size (34-42 μm in diameter)
and distribution of the striae, but the laters are
larger (2-3 μm) than those of C. sinuosus (0.5-
1.5 μm).
Cicatricosisporites sinuosus, reported from the
Neocomian-Barremian of Portugal by Trincão
(1990: 155, pl. 19, fi gs 4, 5, 9), shows a proximal
face with ridges perpendicular to the equator and
a distal face rugate-reticulate, diff erent from C.
sinuosus.
B
OTANICAL AFFINITIES
Cicatricosisporites sinuosus shows similarities with
spores of the genus Saccoloma Kaulfuss (Polypodiales,
Saccolomataceae) (Murillo & Bless 1974; Tryon &
Tryon 1982; Tryon & Lugardon 1991).
Cicatricosisporites sp. 1
(Fig. 9A, B)
Cicatricosisporites sp. B, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 159,
pl. X, fi g. 2 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb rounded triangular. e
laesurae are slightly undulating, bordered by lips,
and extend to the ⅔ of the spore radius. Exine
canaliculate. e proximal face is ornamented by
3 sets of 4 muri parallel to the equator and to the
laesurae. e distal face shows 8 to 10 muri, 3 to
5 μm wide and separated by less than 0.5 μm, which
are parallel among them and to one side (Fig. 11).
e exine is thick at the equator (about 3 μm).
e ornamentation of this form corresponds to the
type VI-B defi ned by Krutzsch (1963). Equatorial
diameter = 40-55 μm.
B
OTANICAL AFFINITIES
Schizaeales, Anemiaceae. Couper (1958) suggested an
affi nity of Cicatricosisporites with spores produced by
Ruff ordia Seward, and Skog (1980, 1982) an affi nity
with Pelletixia Watson & Hill (Schizaeaceae).
Cicatricosisporites sp. 2
(Fig. 9C, D)
Cicatricosisporites sp. D, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 159,
160, pl. VIII, fi g. 5 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb rounded triangular with
convex sides. e spore is bordered by a thin ca-
rina. e laesurae are raised, bordered by a small
thickening, and extend to the equator of the spore.
Exine cicatricose. e proximal face is tetrahedral
and psilate. On the distal face, 3 sets of 4 muri (2
to 4 μm wide, separated by spaces 2 to 5 μm wide)
parallel to the equator form equilateral triangles
centered on the distal pole, where they delimit a
wide triangle (side 15 to 20 μm long) (Fig. 11). e
exine is about 4 μm thick. e ornamentation of
this form corresponds to the type I-A defi ned by
Krutzsch (1963). Equatorial diameter = 45 μm.
B
OTANICAL AFFINITIES
Schizaeales, Anemiaceae.
FIG. 8. — Spores from the Barremian of south-west Japan: A-C, Retitriletes sp. (A, proximal face, Kimigahama Fm, SEM-I1a; B, lateral
view, Kimigahama Fm, SEM-I1a; C, distal face, Kimigahama Fm, SEM-I1c); D, E, H-J, Foveosporites ryosekiensis n. sp. (D, E, proximal
and distal faces, Ashikajima Fm, II2a-U63; H, proximal face, Ashikajima Fm, SEM-II2a; I, proximal face, Ashikajima Fm, SEM-II2b; J, lateral
101
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
A
H
M
M
A-L, N-T
P
QR
I
J
DE
FG
K
L
S
T
N
O
BC
view, Kimigahama Fm, SEM-I1c); F, G, Retitriletes austroclavatidites (Cookson, 1953) Döring, Krutzsch, Mai & Schulz in Krutzsch, 1963,
proximal and distal faces, Ashikajima Fm, II2b-O45/3; K, L, S, T, Cicatricosisporites sinuosus Hunt, 1985 (K, L, proximal and distal faces,
Ashikajima Fm, II2d-D55/4; S, T, proximal and distal faces, Ashikajima Fm, II2b-N33/1); M, P, Microreticulatisporites sp. 1, distal face with
detail, Kimigahama Fm, SEM-I1a; N, O, Microreticulatisporites sp. 2, proximal and distal faces, Ashikajima Fm, II2b-T23d; Q, R, Cicatri-
cosisporites hughesi Dettmann, 1963, proximal and distal faces, Ashikajima Fm, II2h-E52/1. Scale bars: 10 μm.
102
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
Cicatricosisporites sp. 3
(Fig. 9E, F)
Cicatricosisporites sp. E, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 160,
pl. IX, fi g. 8 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb triangular with convex
sides. Tops are slightly raised. e laesurae are thin,
raised, slightly undulating, and bordered by lips in
the contact area. Exine cicatricose. On the proximal
face, we can observe 3 sets of 4 muri which can fork
in each interradial area. ose muri often roll up
slightly before joining with the laesurae. e distal
face shows muri which form triangles centered on
the distal pole; however, those muri are undulating
and intersect the bisecting line, alternating from
one to another (Fig. 11). Equatorial diameter =
40-60 μm.
B
OTANICAL AFFINITIES
Schizaeales, Anemiaceae.
Cicatricosisporites sp. 4
(Fig. 9G, H)
Cicatricosisporites sp. C, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 159,
pl. VIII, fi g. 7 (2009).
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb rounded triangular with
convex sides. e laesurae are raised, slightly undulat-
ing, and extend to the 3/4 of the spore radius. Exine
cicatricose. On the proximal face, a small and psilate
contact area is followed by 3 sets of about 9 narrow
muri (about 0.5 μm wide, separated by furrows of the
same width) parallel to the equator. On the distal face,
many narrow muri are more or less parallel to one set
of the proximal face, and they go from one to another
top of the spore (Fig. 11). e ornamentation of this
form corresponds to the type VI-B defi ned by Krutzsch
(1963). Equatorial diameter = 40-45 μm.
B
OTANICAL AFFINITIES
Schizaeales, Anemiaceae.
Genus Foveosporites Balme, 1957
TYPE SPECIES. — Foveosporites canalis Balme, 1957.
Foveosporites ryosekiensis
Legrand, Pons, Nishida & Yamada n. sp.
(Fig. 8D, E, H-J)
Foveosporites sp., Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique: 149, pl. V, fi g. 16;
pl. VI, fi gs 1, 2 (2009).
TYPE MATERIAL. — Site II, horizon 2, slide a; slides
SEM-a, SEM-b; holotype (II2a-U63; Fig. 8D, E); para-
types (SEM-II2a, SEM-II2b; Fig. 8H, I).
Collection de Paléobotanique-UPMC, Paris, France.
E
TYMOLOGY. — e species name is after the Ryoseki-
type Province of Japan, of which this species seems to
be characteristic.
OCCURRENCE. — Ashikajima and Kimigahama Fm
(common).
T
YPE
LOCALITY
. — Hatoyama, SE Choshi Peninsula,
Chiba Prefecture, Japan.
S
TRATIGRAPHIC
HORIZON
. — Ashikajima Fm (Bar-
remian).
DIAGNOSIS
Trilete microspore. Amb triangular with straight to
slightly convex sides. e laesurae are undulating,
slightly raised (1-2.5 μm), and extend to the 3/4 of
the spore radius. e contact area is psilate or can
sometimes present one tubercule, and is delimited by
a triangular thickening of the exine (3-4 μm wide),
which overhangs and includes the trilete mark, and
can extend to the apices. Beyond this thickening,
the exine is psilate on the proximal face and forms a
broad equatorial fl ange. e equator and the distal
face are densely pitted to foveolate. On the distal
103
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
A
G
K
O
L
N
Q
M
P
H
IJ
B C
D
FE
A-O, Q
P
FIG. 9. — Spores from the Barremian of south-west Japan: A, B, Cicatricosisporites sp. 1, proximal and distal faces, Ashikajima Fm,
II2f-U33d; C, D, Cicatricosisporites sp. 2, proximal and distal faces, Ashikajima Fm, II2a-G40d; E, F, Cicatricosisporites sp. 3, proxi-
mal and distal faces, Ashikajima Fm, II2h-Q57; G, H, Cicatricosisporites sp. 4, proximal and distal faces, Ashikajima Fm, II2f-Y56/1b;
I, J, Cicatricosisporites mohrioides Delcourt & Sprumont, 1955, proximal and distal faces, Kimigahama Fm, I1a-M55; K-M, P, Ruffor-
diaspora australiensis (Cookson, 1953) Dettmann & Clifford, 1992 (K, L, proximal and distal faces, Ashikajima Fm, II2c-D62/4; M, P, distal
face with detail, Kimigahama Fm, SEM-I1a); O, Matonisporites sp. 2, proximal face, Ashikajima Fm, II2a-J46/4; N, Q, Plicatella sp.,
proximal and distal faces, Ashikajima Fm, II2h-J54. Scale bars: 10 μm.
104
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
face, the foveolae are more or less rounded, coarser
near the equator (0.5 to 1.5 μm wide). e exine is
3 to 4 μm thick. Equatorial diameter = 35-45 μm;
equatorial fl ange width = 5-7 μm.
R
EMARKS
Foveosporites labiosus Singh, 1971 has a thickening
on its distal face similar to that observed in F. r yo-
sekiensis, but its location is diff erent. e equatorial
fl ange reminds the Cyatheales (Lophosoriaceae)
fi gured by Tryon & Tryon (1982).
B
OTANICAL AFFINITIES
Lycopodiales, Lycopodiaceae.
Genus Microreticulatisporites
Knox, 1950 emend. Potonié & Kremp, 1954
TYPE SPECIES. — Microreticulatisporites lacunosus (Ibra-
him, 1933) Knox, 1950.
Microreticulatisporites sp. 1
(Fig. 8M, P)
OCCURRENCE. — Kimigahama Fm (one specimen).
DESCRIPTION
Microspore. Amb circular. e laesurae are not
seen. Exine microreticulated and fi nely scabrate.
e mesh is polygonal, of various shapes and sizes
(lumina 0.2 to 1.5 μm in diameter). e exine is
2 μm thick. Equatorial diameter = 40-45 μm.
B
OTANICAL AFFINITIES
e ornamentation of this species shows similarities
with that of some Polypodiales (Dennstaedtiaceae)
fi gured by Tryon & Lugardon (1991). Moreover, the
genus may have sphenophyte affi nities, e.g. with the
Noeggerathiales (Knox 1950; Singh 1964).
Microreticulatisporites sp. 2
(Fig. 8N, O)
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb rounded triangular. e
laesurae are straight, bordered by lips 1 to 1.5 μm
wide, and extend to the ⅔ of the spore radius.
Exine reticulated and slightly rounded. e mesh
is polygonal, of various shapes and sizes (lumina
0.5 to 3 μm in diameter). e exine is 2 μm thick.
Equatorial diameter = 40-50 μm.
B
OTANICAL AFFINITIES
Polypodiales (Dennstaedtiaceae) or Noeggerathiales
(Knox 1950; Singh 1964).
Genus Retitriletes
van der Hammen, 1956 ex Pierce, 1961 emend.
Döring, Krutzsch, Mai & Schulz in Krutzsch, 1963
TYPE SPECIES. — Retitriletes globosus Pierce, 1961.
Retitriletes sp.
(Fig. 8A-C)
OCCURRENCE. — Kimigahama Fm.
DESCRIPTION
Trilete microspore. Amb rounded triangular. e
laesurae are straight, slightly raised, and extend to
the ⅔ of the spore radius. e contact area is psilate,
followed by a reticulum with polygonal lumina of
various shapes and sizes (3-8 μm in dia meter; muri
about 2 μm high) near the equator and on the distal
face. e exine is about 2.5 μm thick. Equatorial
diameter = 40-45 μm.
B
OTANICAL AFFINITIES
Döring et al. in Krutzsch (1963) suggested an affi nity
of Retitriletes with spores produced by the Lycopsida
(Lycopodiaceae), for example Lycopodium Linnaeus.
Genus Matonisporites
Couper, 1958 emend. Dettmann, 1963
TYPE SPECIES. — Matonisporites phlebopteroides Couper,
1958.
105
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
Matonisporites sp. 1
Matonisporites sp. A, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 164,
pl. XI, fi g. 6 (2009).
O
CCURRENCE. — Kimigahama Fm.
DESCRIPTION
Psilate trilete microspore. Amb rounded triangular
with slightly convex sides. e laesurae are straight
and extend to the 3/4 of the spore radius. e exine is
about 4 μm thick. Equatorial diameter = 90 μm.
B
OTANICAL AFFINITIES
Couper (1958) discovered Matonisporites spores in
situ in Phlebopteris Brongniart, Selenocarpus Schenk
and Matonidium Schenk (Gleicheniales, Mato-
niaceae). Ash et al. (1982) and Sukh-Dev (1980)
also discovered in situ Matonisporites spores in Phle-
bopteris sporangia, associated with Dictyophyllidites
Couper, 1958 emend. Dettmann, 1963 spores.
Matonisporites sp. 2 (Fig. 9O)
Matonisporites sp. D, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 165,
pl. XII, fi gs 1, 4 (2009).
O
CCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Psilate trilete microspore. Amb rounded triangular.
e laesurae are straight, bordered by thickenings
(2-4 μm wide), and extend to the ⅔ of the spore
radius. e exine is 2 μm thick. Equatorial dia-
meter = 40-55 μm.
B
OTANICAL AFFINITIES
Gleicheniales, Matoniaceae.
Genus Nodosisporites
Deák, 1964 emend. Dettmann & Cliff ord, 1992
TYPE SPECIES. — Nodosisporites costatus Deák, 1964.
Nodosisporites choshiensis
Legrand, Pons, Nishida & Yamada n. sp.
(Fig. 10G-K)
Nodosisporites sp., Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique: 166, 167, pl. XIII,
fi gs 3, 5, 6 (2009).
T
YPE
MATERIAL
. — Site I, horizon 1, slides SEM-a,
SEM-b; holotype (SEM-a; Fig. 10J); paratype (SEM-b;
Fig. 10K).
Collection de Paléobotanique-UPMC, Paris, France.
E
TYMOLOGY
. — e species name is after the Choshi
Group, from which it was reported.
O
CCURRENCE. — Ashikajima and Kimigahama Fm.
TYPE LOCALITY. — Kimigahama bay, Choshi Peninsula,
Chiba Prefecture, Japan.
S
TRATIGRAPHIC
HORIZON
. — Kimigahama Fm (late
Barremian).
DIAGNOSIS
Tetrahedral trilete microspore. Amb rounded
triangular. Proximal face slightly fl attened, with a
psilate contact area followed by 1 or 2 muri parallel
to the equator. e laesurae are relatively narrow,
slightly raised and undulating, and extend to the
3/4 of the spore radius. e exine is cicatricose.
e distal face is strongly convex (35 to 40 μm in
polar diameter). It shows 3 sets of 3-4 muri paral-
lel to the equator, that join in the apex areas. One
of these sets goes on to form a triangle centered
on the distal pole (Fig. 11). e muri are raised,
2 to 3 μm wide, and are ornamented by tuber-
cules, verrucae or spines (5 to 7 μm long) regu-
larly distributed. e ornamentation of this form
corresponds to the type I-A defi ned by Krutzsch
(1963). Equatorial diameter = 35-50 μm; polar
diameter = 35-40 μm.
R
EMARKS
Appendicisporites spinosus Pocock, 1964 shows
supramural bacula, verrucae or spines as observed
in Nodosisporites, but diff ers from the later in
that these elements are not evenly distributed.
Cicatricose or canaliculate genera Cicatricosis-
porites and Plicatella also show some morpho-
106
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
logical similarities, but both lack supramural
elements.
BOTANICAL AFFINITIES
Schizaeales, Anemiaceae (Dettmann & Cliff ord
1992).
Nodosisporites makotoi
Legrand, Pons, Nishida & Yamada n. sp.
(Fig. 10A-F)
Nodosisporites sp., Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique: 166, 167, pl. XIII,
fi gs 1, 2, 4, 7; pl. XIV, fi g. 7 (2009).
T
YPE MATERIAL. — Site II, horizon 2, slides a, b; holo-
type (II2a-L58/2; Fig. 10A, D); paratypes (II2b-E56/3,
II2b-U33; Fig. 10B, C, E, F).
Collection de Paléobotanique-UPMC, Paris, France.
E
TYMOLOGY
. — e species is named in honour of Prof.
Makoto Nishida, who fi rst studied fossil plants from the
Choshi Group.
O
CCURRENCE. — Ashikajima and Kimigahama forma-
tions.
T
YPE
LOCALITY
. — Hatoyama, SE Choshi Peninsula,
Chiba Prefecture, Japan.
STRATIGRAPHIC HORIZON. — Ashikajima Formation
(Barremian).
DIAGNOSIS
Tetrahedral trilete microspore. Amb rounded
triangular. Proximal face slightly fl attened, with
a psilate contact area followed by 1 or 2 muri
parallel to the equator. e laesurae are rela-
tively narrow, raised (about 3 μm), undulating,
and extend to the 3/4 of the spore radius or to
the equator; they are smooth or ornamented by
small spines (0.5-1 μm high and 1-3 μm in basal
diameter). Near the apices, 5-6 big spines (3-5 μm
high) are present. A translucent equatorial zona
with a dentate margin can be seen. e exine is
cicatricose. e distal face shows 3 sets of 3-4
muri sinuous but parallel to the equator. One of
these sets goes on to form a triangle (sometimes,
more or less a network) centered on the distal
pole (Fig. 11). e ornamentation of this form
corresponds to the type I-A defi ned by Krutzsch
(1963). Equatorial diameter = 40-45 μm.
R
EMARKS
Appendicisporites dentimarginatus Brenner, 1963
shows similarities with Nodosisporites makotoi, but
lacks spines on the apices.
B
OTANICAL AFFINITIES
Schizaeales, Anemiaceae (Dettmann & Cliff ord
1992). e ornamentation shows similarities with
that of Anemia phyllitidis (Linnaeus) Swartz from
Brazil, fi gured by Tryon & Lugardon (1991).
Genus Plicatella
Maljavkina, 1949 emend. Burden & Hills, 1989
T
YPE
SPECIES
. — Plicatella trichacantha Maljavkina,
1949.
Plicatella sp.
(Fig. 9N, Q)
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb triangular with straight
to slightly convex sides and raised tops. e
laesurae are thin, straight, raised, and extend to
the 3/4 of the spore radius. Exine canaliculate.
e proximal face shows a small psilate contact
area, followed by 3 sets of 3-4 muri (1.5 to 4 μm
wide) parallel to the equator; the muri join at the
tops. e distal face is ornamented by 3 sets of
4 wavy muri parallel to the equator, distributed
in a slight interval to form concentric triangles
centered on the distal pole (Fig. 11). e orna-
mentation of this form corresponds to the type
I-B defi ned by Krutzsch (1963). Equatorial di-
ameter = 40-50 μm.
B
OTANICAL AFFINITIES
Schizaeales, Schizaeaceae (Dettmann & Cliff ord
1992).
107
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
F
IG
. 10. — Spores from the Barremian of south-west Japan: A-F, Nodosisporites makotoi n. sp. (A, D, proximal and distal faces, Ashikajima
Fm, II2a-L58/2; B, E, proximal and distal faces, Ashikajima Fm, II2b-E56/3; C, F, proximal and distal faces, Ashikajima Fm, II2b-U33);
G-K, Nodosisporites choshiensis n. sp. (G, H, lateral view, Ashikajima Fm, II2b-K22/2; I, lateral view, Ashikajima Fm, II2b-E40b; J, proximal
face, Kimigahama Fm, SEM-I1a; K, distal face, Kimigahama Fm, SEM-I1b); L-N, Cingulatisporites sp. 1 (L, proximal face, Kimigahama
Fm, I1b-R41; M, N, proximal and distal faces, Ashikajima Fm, II2c-J26/3); O, P, Cingulatisporites psilatus Groot & Penny, 1960, proximal
and distal faces, Kimigahama Fm, I1a-W67g; Q, Cingutriletes sp., proximal face, Ashikajima Fm, II2a-H67g. Scale bar: 10 μm.
A
D
G
OP
Q
J
K
H
L
M
N
I
C
F
B
E
108
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
Genus Cingulatisporites
omson in omson & Pfl ug, 1953
emend. Potonié, 1956
TYPE SPECIES. — Cingulatisporites levispeciosus Pfl ug in
omson & Pfl ug, 1953.
Cingulatisporites sp. 1
(Fig. 10L-N)
Cingulatisporites sp. B, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 168,
pl. XIV, fi gs 4-6 (2009).
O
CCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Cingulate trilete microspore. Amb rounded trian-
gular. e laesurae are undulating and extend to
the cingulum, where they divide into two ends.
Exine scabrate. Equatorial diameter = 30-35 μm;
cingulum thickness = 3-5 μm.
B
OTANICAL AFFINITIES
Filicopsida.
Cingulatisporites sp. 2
(Fig. 12A)
OCCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Cingulate trilete microspore. Amb rounded triangu-
lar. e laesurae are raised, undulating, and extend
to the cingulum. Exine scabrate. Equatorial diam-
eter = 45-50 μm; cingulum thickness = 5-8 μm.
B
OTANICAL AFFINITIES
Filicopsida.
Genus Cingutriletes
Pierce, 1961 emend. Dettmann, 1963
TYPE SPECIES. — Cingutriletes congruens Pierce, 1961.
Cingutriletes sp. (Fig. 10Q)
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Cingulate trilete microspore. Amb rounded triangular
with straight to convex sides. e laesurae are straight
and extend to the ⅔ of the spore radius (a little before
the cingulum). e cingulum is wide (4-5 μm, that
is to say more than 1/5 of the spore radius). Exine
psilate. Equatorial diameter = 30-35 μm.
B
OTANICAL AFFINITIES
Filicopsida (Couper 1958), Lycopsida (Selaginel-
laceae?) (Pierce 1961).
Genus Distaltriangulisporites Singh, 1971
TYPE SPECIES. — Distaltriangulisporites perplexus (Singh,
1964) Singh, 1971.
Distaltriangulisporites sp. (Fig. 12B, C)
Distaltriangulisporites sp. B, Legrand, Palynologie des
dépôts Jurassique supérieur et Crétacé inférieur du Japon,
et provinces paléofl oristiques du sud-est asiatique: 169,
pl. XIV, fi g. 10 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Cingulate trilete microspore. Amb rounded triangular.
e laesurae are slightly raised and extend to the cingu-
lum. Proximal face scabrate. e distal face shows fl at
muri forming a coarse reticulum with wide rounded
lumina. Equatorial diameter = 28-32 μm.
B
OTANICAL AFFINITIES
Schizaeales, Anemiaceae (Dettmann & Cliff ord
1992).
Genus Patellasporites
Groot & Groot, 1962 emend. Kemp, 1970
T
YPE
SPECIES
. — Patellasporites tavaredensis Groot &
Groot, 1962.
109
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
Patellasporites sp.
Patellasporites sp. A, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 172,
pl. XV, fi g. 2 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Trilete microspore. Amb circular. e laesurae
are straight, slightly raised, thicken, and extend
to the inner side of the patella. Proximal face
psilate. Verrucae merge at the equator, forming
a patella, but are separated by thin radial canals.
e distal face is covered by verrucae of various
sizes, bigger near the equator than at the pole,
which merge and let only very small irregular
spaces between them. Equatorial diameter =
30 μm.
BOTANICAL AFFINITIES
Filicopsida.
Cicatricosisporites sp.
cf. C. mohrioides
Cicatricosisporites sp. 1 Cicatricosisporites sp. 2
Cicatricosisporites sp. 3 Cicatricosisporites sp. 4
Nodosisporites choshiensis n. sp.
Plicatella sp.Nodosisporites makotoi n. sp.
FIG. 11. — Ornamentation of the schizaealean forms corresponding to new species or taxa placed in open nomenclature.
110
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
Genus Aequitriradites Delcourt & Sprumont,
1955 emend. Dettmann, 1963
TYPE SPECIES. — Aequitriradites dubius Delcourt &
Sprumont, 1955.
Aequitriradites sp.
(Fig. 12E)
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Zonate trilete microspore. Amb rounded subtrian-
gular. Central body subcircular. Laesurae are not
visible. Central body verrucate; verrucae are very
small and contiguous (about 1-2 μm in diameter).
e exine thickens near the periphery of the central
body (about 3-3.5 μm). e zona is thin, granu-
late, radially folded, with irregular and more or
less transparent margins. It is wider in three areas
(about fi ve times wider), which gives to the spore a
subtriangular shape. No hilum is visible. Equatorial
diameter = 45-70 μm; zona width = 2-12 μm.
B
OTANICAL AFFINITIES
Bryophyta, Marchantiopsida.
Genus Laevigatosporites
Ibrahim, 1933
emend. Schopf, Wilson & Bentall, 1944
TYPE SPECIES. — Laevigatosporites vulgaris (Ibrahim,
1932) Ibrahim, 1933.
Laevigatosporites sp. (Fig. 12J)
Laevigatosporites sp. A, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 177,
pl. XVI, fi g. 11 (2009).
O
CCURRENCE. — Ashikajima Fm (rare).
DESCRIPTION
Psilate monolete microspore. Amb broadly elliptical
in lateral view. e proximal face is slightly concave.
e laesura is straight and runs longitudinally on ⅓ of
the proximal face. Exine about 2 μm thick. Equatorial
diameter = 60 μm; polar diameter = 40 μm.
B
OTANICAL AFFINITIES
Polypodiales (Polypodiaceae, Pteridaceae or Denns-
taedtiaceae).
Incertae sedis
Incertae sedis in Hasenboehler (1981)
(Fig. 5E)
OCCURRENCE. — Ashikajima Fm.
D
ISTRIBUTION
. — Albian-Cenomanian of Portugal
(Hasenboelher 1981).
DESCRIPTION
Psilate trilete microspore. Amb triangular. e laesu-
rae are straight, bordered by lips about 3 μm wide,
and extend to the equator of the spore. Equatorial
diameter = 45-50 μm.
B
OTANICAL AFFINITIES
Filicopsida.
Incertae sedis sp. 1
(Fig. 13N, O)
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Microspore? Amb circular. e proximal face is not
seen. e exine is reticulated with a mesh polygo-
nal, of various shapes and sizes (lumina 1-5 μm in
FIG. 12. — Spores and pollen grains from the Barremian of south-west Japan: A, Cingulatisporites sp. 2, proximal face, Ashikajima Fm,
SEM-II2c; B, C, Distaltriangulisporites sp., proximal and distal faces, Ashikajima Fm, II2c-N53; D, Contignisporites cooksonii (Balme,
1957) Dettmann, 1963, distal face, Kimigahama Fm, I1a-O51/1; E, Aequitriradites sp., Ashikajima Fm, II2c-Y59/2; F, Aequitriradites
verrucosus (Cookson & Dettmann, 1958) Cookson & Dettmann, 1961, Ashikajima Fm, II2h-J66/4; G, M, Contignisporites burgeri Filatoff,
McKellar & Price in Filatoff & Price, 1988, proximal and distal faces, Ashikajima Fm, II2d-P47/4; H, I, Polycingulatisporites reduncus
111
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
A B
E
H
I
P
Q
R
J
K
L
O
G
M
N
F
C D
(Bolkhovitina, 1953) Playford & Dettmann, 1965, proximal and distal faces, Kimigahama Fm, I1a-Z52; J, Laevigatosporites sp., lateral
view, Ashikajima Fm, II2e-Y60; K, Laevigatosporites ovatus Wilson & Webster, 1946, lateral view, Kimigahama Fm, I1c-P35/3; L, Exesi-
pollenites tumulus Balme, 1957, distal view, Kimigahama Fm, I1a-R60/3; N, Callialasporites sp. cf C. ugensis Takahashi & Sugiyama,
1990, Ashikajima Fm, II2f-Y42/3; O, Eucommiidites troedssonii (Erdtman, 1948) Potonié, 1958, Kimigahama Fm, I1b-R39; P, Cerebro-
pollenites mesozoicus (Couper, 1958) Nilsson, 1958, lateral view, Ashikajima Fm, II2a-J61/1d; Q, Alisporites sp., polar view, Ashikajima
Fm, II2d-Q59/2; R, Callialasporites dampieri (Balme, 1957) Sukh-Dev, 1961, Ashikajima Fm, II2b-T46/4m. Scale bar: 10 μm.
112
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
diameter). e exine is about 1.5 μm thick. Equa-
torial diameter = 20-25 μm.
R
EMARKS
Only a group of these elements was observed, and
isolated ones must be found to permit an observation
of the two faces and propose an identifi cation.
B
OTANICAL AFFINITIES
Unknown.
Anteturma
VARIEGERMINANTES = POLLENITES
Genus Callialasporites
Sukh-Dev, 1961 emend. Potonié, 1966
T
YPE
SPECIES
. — Callialasporites trilobatus (Balme, 1957)
Sukh-Dev, 1961.
Callialasporites sp. cf. C. ugensis
Takahashi & Sugiyama, 1990
(Fig. 12N)
OCCURRENCE. — Ashikajima Fm (rare).
DESCRIPTION
Monosaccate pollen grain. Amb oval to subcircular.
e central body is surrounded by a narrow and
undulating equatorial saccus (5-8 μm). e exine is
punctate on the thinned distal pole area. Equatorial
diameter = 35-55 μm.
R
EMARKS
Two morphologically similar but larger species
have been reported by Takahashi & Sugiyama
(1990) from the Santonian of the Taneichi For-
mation, north-east Honshu, Japan (70-84 μm):
Callialasporites ugensis Takahashi & Sugiyama, and
by Kumar (1973) from the Lower Cretaceous of
India (55-68 μm): Callialasporites doringii Kumar
(previously named C. lenticularis (Döring, 1961)
Venkatachala & Kar, 1969 by Singh & Kumar
(1969): pl. 1, fi gs 15, 16). Deák (1962) reported
a grain also showing similarities to ours: Inaper-
turopollenites undulatus Weyland & Greifeld, 1953
(pl. 12, fi gs 10, 13).
B
OTANICAL AFFINITIES
Pinales. Archangelsky & Gamerro (1967) and
Gamerro (1965, 1968) suggested similarities of grains
of the genus Callialasporites with those observed
in cones of Apterocladus lanceolatus Archangelsky
(Podocarpaceae) from the Lower Cretaceous of
Patagonia. Van Konijnenburg-Van Cittert (1971)
reported abnormal grains of the same morphology
in an Araucariaceae.
Genus Alisporites
Daugherty, 1941 emend. Rouse, 1959
TYPE SPECIES. — Alisporites opii Daugherty, 1941.
Alisporites sp. cf. A. australis
de Jersey, 1962
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Bisaccate pollen grain. Central body spherical,
scabrate. e sacci are reticulate and a little longer
than the central body, probably hanging. Total
width = 50 μm. Central body diameter = 25 μm.
Saccates (length × width) = 40 × 10 μm.
REMARKS
is grain is morphologically very similar to
Alisporites australis de Jersey, 1962 previously
reported from the Triassic of Australia (de Jer-
sey 1962).
FIG. 13. — Pollen grains, alga and fungus from the Barremian of south-west Japan: A, Pityosporites sp. cf. P. constrictus Singh, 1964,
lateral view, Ashikajima Fm, II2b-J34/1; B, Alisporites thomasii (Couper, 1958) Pocock, 1962, polar view, Ashikajima Fm, II2b-O42/3;
C, Incertae sedis sp. 2, Ashikajima Fm, II2c-W65/4; D, Balmeiopsis limbatus (Balme, 1957) Archangelsky, 1977, Ashikajima Fm, II2d-
N41/2; E, Inaperturo pollenites sp., Kimigahama Fm, I1b-U64/4; F-H, Classopollis torosus (Reissinger, 1950) Couper, 1958 emend.
113
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
A B
D
I
J
K
L
A-N, P, Q
O
M
E
F
C
G
H
N
O
P
Q
Burger, 1965 (F, distal polar view, Ashikajima Fm, SEM-II2a; G, distal polar view, Ashikajima Fm, II2c-H32/1; H, tetrad, Ashikajima
Fm, SEM-II2a); I, Cycadopites minimus (Cookson, 1947) Pocock, 1970, Ashikajima Fm, II2e-U64; J, Cycadopites sp., Kimigahama
Fm, I1a-X30; K, Ephedripites montanaensis Brenner, 1968, Ashikajima Fm, II2c-N35/4h; L, Gnetaceaepollenites sp., Ashikajima Fm,
II2a-S47/1a; M, Corollina sp., tetrad, Kimigahama Fm, I1a-X42; N, O, Incertae sedis sp. 1, group with detail, Ashikajima Fm, SEM-II2a;
P, Pterospermella sp., Ashikajima Fm, II2b-O65; Q, Microthyriacites sp., Ashikajima Fm, II2e-R63/4. Scale bars: 10 μm.
114
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
BOTANICAL AFFINITIES
Gymnosperms, Pteridospermales?
Alisporites sp.
(Fig. 12Q)
Alisporites sp. A, Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique: 183, pl. XVIII,
fi gs 4, 7, 8 (2009).
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Bisaccate pollen grain. Central body spheri-
cal. e sacci are semi-circular, about as high as
the central body, and wrap the proximal face.
e exine is about 1.5 μm thick. Total width =
43 μm. Central body diameter = 25 μm. Sacci
(length × width) = 15 × 28 μm.
B
OTANICAL AFFINITIES
Gymnosperms, Pteridospermales?
Genus Pityosporites Seward, 1914
TYPE SPECIES. — Pityosporites antarcticus Seward, 1914.
Pityosporites sp. cf. P. constrictus Singh, 1964
(Fig. 13A)
OCCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Bisaccate pollen grain. Central body more or less
spherical, granulate. e sacci are reticulate; they are
small, hanging, narrower at their base, letting a space
of about 12 μm between them. e exine is thin
(about 1 μm). Total width = 50 μm. Central body
diameter = 30 μm. Sacci (length × width) = 25 × 17-
20 μm.
R
EMARKS
is grain is morphologically very similar to
Pityosporites constrictus Singh, 1964 previously
reported from the Aptian-Albian of Korea (Yi
et al. 1993), Albian of Canada (Singh 1964),
and Santonian of the Kuji Group in north-east
Honshu, Japan (Miki 1972; Takahashi & Sugi-
yama 1990).
B
OTANICAL AFFINITIES
Pinales.
Pityosporites sp. cf. P. piniformis
(Zaklinskaya, 1957) Takahashi & Sugiyama,
1990
OCCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Bisaccate pollen grain. Central body oval, psi-
late to fi nely scabrate. Two small sacci are distally
hanging. Proximal crest 1 to 1.5 μm thick. Total
width = 50-55 μm; grain height = 15-20 μm. Sacci
(length × width) = 20 × 15 μm.
R
EMARKS
is grain, fi rstly described in the Eocene-Oligocene
of the Pavlodar Irtysh Basin (western Siberia) by
Zaklinskaya (1957), was found by Takahashi &
Sugiyama (1990) in the Santonian (Taneichi For-
mation) of north-east Honshu, Japan. Zaklinskaya
(1957) had previously placed it in the genus Ce-
drus Tr ew.
B
OTANICAL AFFINITIES
Pinales, probably Pinaceae.
Genus Inaperturopollenites
Pfl ug & omson in omson & Pfl ug, 1953
TYPE SPECIES. — Inaperturopollenites dubius (Pot-
onié & Venitz, 1934) Pfl ug & omson in omson &
Pfl ug,1953.
Inaperturopollenites sp.
(Fig. 13E)
OCCURRENCE. — Ashikajima and Kimigahama Fm.
115
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
DESCRIPTION
Inaperturate pollen grain. Amb subcircular. e
exine is psilate, very folded and thin (about 1 μm
thick). Equatorial diameter = 25-45 μm.
B
OTANICAL AFFINITIES
Pinales.
Genus Corollina Maljavkina, 1949
TYPE SPECIES. — Corollina compacta Maljavkina, 1949.
Corollina sp.
(Fig. 13M)
OCCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Pollen grain. Amb subspherical with poles fl attened. At
the proximal pole, a trilete mark is seen. At the distal
pole, a pseudopore 3-7 μm in diameter is present. e
rimula delimits a cap about 17-24 μm in diameter. No
equatorial ridge is seen. e exine is psilate to slightly
punctate. Equatorial diameter = 22-30 μm.
R
EMARKS
e genus Corollina gathers species from the Keu-
per without ridge, but it still remains used for the
Cretaceous of southeastern Asia.
B
OTANICAL AFFINITIES
Pinales, Cheirolepidiaceae (Couper 1958). Tekl-
eva & Krassilov (2009) suggested an affi nity with
the Gnetales.
Genus Gnetaceaepollenites iergart, 1938
TYPE SPECIES. — Gnetaceaepollenites ellipticus iergart,
1938.
Gnetaceaepollenites sp.
(Fig. 13L)
Gnetaceaepollenites sp. B, Legrand, Palynologie des dépôts
Jurassique supérieur et Crétacé inférieur du Japon, et
provinces paléofl oristiques du sud-est asiatique: 191,
pl. XX, fi g. 15 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Polyplicate pollen grain. Amb ellipsoidal. e exine
is constituted of about 15 ribs running through the
grain, and describing a senestral “helix” of 180°. e
ribs are 1.8-3 μm wide, separated by furrows less
than 0.5 μm wide. e ends of the ribs closely gather
at each end of the long axis of the grain without
joining. Length = 38 μm; width = 27 μm.
B
OTANICAL AFFINITIES
Gymnosperms, Gnetales.
Genus Cycadopites
Wodehouse, 1933 ex Wilson & Webster, 1946
T
YPE
SPECIES
. — Cycadopites follicularis Wilson & Web-
ster, 1946.
Cycadopites sp.
(Fig. 13J)
Cycadopites sp. A, Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique: 192, pl. XXI,
fi g. 2 (2009).
O
CCURRENCE. — Kimigahama Fm.
DESCRIPTION
Monosulcate pollen grain. Amb ellipsoidal fusi-
form. A narrow furrow runs longitudinally through
the grain, slightly widening at the ends; it can be
bordered by folds of the exine (4 to 5 μm wide).
e exine is thin (about 1 μm thick) and psilate.
Length = 45-50 μm; width = 20-25 μm.
B
OTANICAL AFFINITIES
Cycadales or Bennettitales (Balme 1995). e
present Cycadales are distributed in the tropical
to subtropical arid regions. e genus Cycadopites
is often found associated with coal strata, and the
habitat of this group is supposed to have been
116
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
much more diversifi ed than now (Singh 1964).
Nevertheless, grains of the genus Cycadopites were
found in situ in Sahnia Puri (Pentoxylales, Pentoxy-
laceae) (Sukh-Dev 1980) and Lepidopteris Schimper
emend. Townrow (Peltaspermales) (Anderson &
Anderson 1983).
Incertae sedis
Incertae sedis sp. 2
(Fig. 13C)
Incertae sedis sp. G, Legrand, Palynologie des dépôts Juras-
sique supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique: 195, pl. XXI,
fi g. 7 (2009).
O
CCURRENCE. — Ashikajima Fm.
DESCRIPTION
Pollen grain. Amb oval in polar view. An eight
shape wide median sulcus with rounded ends,
runs through the grain. In the middle part of
the grain, the edges of the sulcus strongly widen
(until 10-12 μm), and can fold one over the
other. is thickening can widen progressively
or quite abruptly. e exine is psilate (about
1.5 μm thick). Length × width = 40-50 × 30-
35 μm; longitudinal thickenings (length × width) =
25 × 10-12 μm.
REMARKS
ese grains have similarities with Cycadopites, but
the folded lips that can be observed here on some
grains are much more developped. e genus En-
tylissa Naumova ex Ischenko, shows developped
lips, but slenderer and more ornamented than in
Incertae sedis sp. 2. McGregor (1965) fi gured a pol-
len grain similar to our species: cf. Ginkgocycado-
phytus caperatus (Luber, 1941) Samoilovich, 1953
(pl. 5, fi g. 25), from the Upper Jurassic of Canada.
Monosulcites scaber Kimyai, 1966 reported from the
Cretaceous of New Jersey, USA (Kimyai 1966), also
shows great similarities to our species.
B
OTANICAL AFFINITIES
Gymnosperms.
Algae chlorophyta
CHLOROPHYCEAE
Genus Botryococcus Kützing, 1849
TYPE SPECIES. — Botryococcus braunii Kützing, 1849.
Botryococcus sp. cf. B. braunii Kützing, 1849
OCCURRENCE. — Ashikajima Fm (rare).
D
ISTRIBUTION. — is genus is known from the Pre-
cambrian.
DESCRIPTION
Algae grouped in small bulb-like colonies, very
altered. Colony diameter = 20-25 μm.
B
OTANICAL AFFINITIES
Chlorococcales, Botryococcaceae. Freshwater green
algae.
PRASINOPHYCEAE
Genus Pterospermella Eisenack, 1972
TYPE SPECIES. — Pterospermella aureolata (Cookson &
Eisenack, 1958) Eisenack, 1972.
Pterospermella sp.
(Fig. 13P)
OCCURRENCE. — Ashikajima Fm (rare).
D
ISTRIBUTION. — is genus is known from the Pre-
cambrian to possibly Holocene.
DESCRIPTION
Flattened lenticular body. e central body is
biconvex and darker than the folded equatorial
membrane that surrounds it. Length = 38 μm;
width = 30 μm; central body (length × width)
= 15 × 10 μm; equatorial membrane width =
10 μm.
117
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
BOTANICAL AFFINITIES
Pterospermatales, Pterospermellaceae of marine
environment.
Genus Tasmanites Newton, 1875
TYPE SPECIES. — Tasmanites punctatus Newton, 1875.
Tasmanites sp.
OCCURRENCE. — Ashikajima Fm (rare).
D
ISTRIBUTION. — is genus is known from the Pre-
cambrian to possibly Holocene.
DESCRIPTION
Cyst with a circular equatorial shape. e wall
is thick (about 10 μm), pierced by many dis-
tinct small pores about 1 μm in diameter. A slit
can be seen at the center of the cyst, suggest-
ing an empty phycoma. Equatorial diameter =
83-93 μm.
BOTANICAL AFFINITIES
Pterospermatales, Tasmanitaceae of marine envi-
ronment.
FUNGI ASCOMYCOTA
Genus Microthyriacites Cookson, 1947
TYPE SPECIES. — Microthyriacites grandis Cookson, 1947
Microthyriacites sp.
(Fig. 13Q)
OCCURRENCE. — Ashikajima and Kimigahama Fm.
DESCRIPTION
Disc-like body, brown-colored, with radially ar-
ranged cell rows; the margins of the body are thin
and irregular. e “ostiole” is not distinct, but a
hole in the center of the disc is present, formed by
dissolving of the central cells. Cells are square to
1 %
36 %
4 %
12 %
2 %
11 %
9 %
4 %
21 %
1 %
28 %
23 %
7 %
14 %
12 %
1 %
9 %
5 %
Ashikajima Fm Kimigahama Fm
Psilate spores
Cicatricose spores
Other ornamented spores
Monosulcate pollen grains
Classopollis-Corollina
Bisaccate pollen grains
Araucariaceae
Other gymnosperms
Dinoflagellates
FIG. 14. — Percentages of nine palynomorph groups: A, Ashikajima Formation; B, Kimigahama Formation, south-west Japan.
118
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
elongated, without pore, 3-10 μm long and 2.5 μm
wide. Cells of the central part of the disc are thicker.
Equatorial diameter = 65-70 μm.
B
OTANICAL AFFINITIES
Dothideomycetes, Microthyriales. Epiphyllous
fungi.
Genus Phragmothyrites Edward, 1922
T
YPE
SPECIES
. — Phragmothyrites eocaenicus Edward,
1922.
Phragmothyrites sp.
OCCURRENCE. — Ashikajima Fm.
DESCRIPTION
Disc-like body, brown-colored, with radially arranged
cell rows; the margins of the body are irregular. No
ostiole is present, and the central area is continuous.
Cells are square to elongated, 5-10 μm long and
4-6 μm wide. Cells of the central part of the disc
are thicker. Equatorial diameter = 75-85 μm.
B
OTANICAL AFFINITIES
Dothideomycetes, Microthyriales. Epiphyllous
fungi.
RESULTS
e palynomorphs recovered from the Ashika-
jima and Kimigahama formations of the Choshi
Group show a generally good state of preservation
and considerable diversity. ey are dominated
by spores, mainly represented by species with af-
fi nities to the Anemiaceae, Matoniaceae or Cyat-
heaceae (Table 2). Matonisporites (Matoniaceae)
is represented by three species, some large (90
μm). Cicatricosisporites (Anemiaceae) spores are
abundant and diverse in the assemblages, but may
not refl ect a diversity of the source plants (several
studies have shown the presence, inside the same
sporangium, of variously ornamented spores that
can belong to diff erent “morphographic” species).
Ruff ordiaspora australiensis, Plicatella are common,
and two new species of Nodosisporites have been
described: N. choshiensis and N. makotoi. Several
species of Cyathidites (Cyatheaceae or Dickso-
niaceae), which had a world-wide distribution
during this period, and spores having affi nities
with the Osmundaceae (Biretisporites potoniaei,
Baculatisporites, Todisporites, Osmundacidites)
are present. Deltoidospora (Cyatheales), Undu-
latisporites and Cingulatisporites (Filicopsida)
are common. Contignisporites (Pteridaceae) was
observed, but is rare in the assemblages.
Densely echinate spores, with echinae of vari-
ous shapes and sizes, are common. We observed
Distaltriangulisporites, verrucate spores referable
to Verrucosisporites, Converrucosisporites, Lep-
tolepidites, reticulate spores of Klukisporites and
Ischyosporites, and Pilosisporites. A new species of
Manumia: M. japonica (Pteridaceae), with verru-
cae and ridges more or less randomly distributed,
has been described. is genus is reported here
for the fi rst time in Asia. Monolete spores of the
genus Laevigatosporites are common.
Spores with lycopsid affi nities are common. We
note the presence of Lycopodiumsporites dentimura-
tus in both formations. is species was described
by Brenner (1963) from the Barremian-Albian of
Maryland (USA), but has never been reported
from any other area. Retitriletes austroclavatidites,
Echinatisporis varispinosus, various species of
Neoraistrickia and a new species of Foveosporites:
F. ryo sekiensis, are present. Triporoletes reticula-
tus, Aequitriradites verrucosus and A. spinulosus,
which have affi nities with the Marchantiopsida
(Hepatica), are also recorded.
Gymnosperm pollen grains are mainly rep-
resented by Classopollis (Cheirolepidiaceae),
Araucariacites australis and Balmeiopsis limbatus
(Araucariaceae), and monosulcate grains of Cyca-
dopites. However, proportions of Classopollis are
quite low (9-14%) compared to the 70% reported
by Takahashi (1974) in the Aptian-Albian of the
Miyako Group (north-east Honshu, Japan). Bisac-
cate pollen grains are few but quite diverse, being
represented by Alisporites, Abietineaepollenites,
Cedripites and Pityosporites. We also recorded
119
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
rare Callialasporites (Podocarpaceae) and some
Eucommiidites, Ephedripites and Gnetaceaepol-
lenites (Gnetophyta). Callialasporites is rare in
Japan, whereas this genus is generally abundant
world-wide.
Several types of dinofl agellates and some fo-
raminiferal linings were also observed in the two
formations, and will be reported elsewhere. e
composition of the palynofl oras is summarized
in two diagrams (Fig. 14).
60º N
0º
30º N
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
A
B
Manumia japonica n. sp.
Manumia delcourtii
(Pocock, 1970) Dybkjær, 1991
Manumia verrucata
Pocock, 1970
Manumia irregularis
Pocock, 1970
Manumia variverrucata
(Couper, 1958) Hoelstad, 1985
Fig. 15. — Spatio-temporal distribution of genus Manumia. This genus had a wide distribution across the South-Laurasian Province
(= Euro-Sinian Region): A, during Jurassic time; B, during Early Cretaceous time. Paleogeographical map during the Hauterivian (120
My) in North polar stereographic projection, modifi ed from Smith & Briden (1977) and Masse et al. (1993). Figures of Manumia irregu-
laris Pocock, 1970 and M. verrucata Pocock, 1970 are from Pocock 1970 (source: www.schweizerbart.de); fi gure of M. variverrucata
(Couper, 1958) Hoelstad, 1985 is from Hoelstad 1985 (source: 2dgf.dk); fi gure of M. delcourtii (Pocock, 1970) Dybkjær, 1991 is from
Koppelhus & Batten 1996 (source: www.palynology.org).
120
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
DISCUSSION
Most of the species we identifi ed are distributed
world-wide, and of long stratigraphic range. How-
ever, some species may be specifi c to the Ryoseki-
type Province in Japan (Foveosporites ryosekiensis,
Nodosisporites choshiensis, N. makotoi, Lycopodi-
umsporites dentimuratus) or show diff erent pro-
portions from what is generally observed for the
same period in the adjacent regions, for example
the scarcity of Classopollis or Callialasporites, and
the absence of Appendicisporites and Gleicheniidites
often reported from China and Russia. Spores of
the genus Manumia, reported for the fi rst time in
Asia (Fig. 15), are common in the two formations.
Four species have been described in this genus:
Manumia delcourtii (Pocock, 1970) Dybkjær,
1991 was reported from the Lower to Middle
Jurassic of Denmark (Dybkjær 1991; Koppelhus
1992; Seidenkrantz et al. 1993; Koppelhus &
Nielsen 1994; Koppelhus & Dam 2003), East
Greenland (Koppelhus & Hansen 2003) and
Sweden (Lindström & Erlström 2007), and the
Middle Jurassic of Scotland (Stefanowicz 2008);
Manumia irregularis Pocock, 1970 was reported
from the Lower to Middle Jurassic of East Green-
land (Lund & Pedersen 1984) and Afghanistan
(Schweitzer et al. 1987), and the Middle Jurassic
of Germany (Schweitzer et al. 1987) and Canada
(Pocock 1970); Manumia variverrucata (Couper,
1958) Hoelstad, 1985 was reported from the
Lower to Middle Jurassic of Germany (Schulz
1967) and Holland (Herngreen & de Boer 1974),
the Middle Jurassic of Denmark (Hoelstad 1985),
Sweden (Guy 1971) and Alaska (Bjærke 1993),
and the Middle to Upper Jurassic of England
(Couper 1958); Manumia verrucata Pocock, 1970
was reported from the Middle Jurassic of Canada
(Pocock 1970). Bøe et al. (2005) also reported
Manumia sp. from the Upper Jurassic to Lower
Cretaceous (Barremian) of Norway.
We identifi ed Cicatricosisporites sinuosus in
both formations. is species was reported for
the fi rst time by Hunt (1985) from the Purbeck
Limestone Group of southern England. It has
been reported from the Hauterivian-Barremian
of Japan (Legrand 2009). It seems to have crossed
the Tethys and reached North Africa via Lebanon
(Dejax, personnal communication), Israel (de
Haan 1997) and Libya (Batten & Uwins 1985;
Uwins & Batten 1988) during the Barremian, and
then spread through Egypt (Saad 1978; Omran
et al. 1990; Schrank & Ibrahim 1995; Ibrahim
1996; Ibrahim et al. 2001; Mahmoud & Deaf
2007;
Mahmoud et al. 2007), Tunisia (Pons,
personnal communication) and Iraq (Al-Ameri
et al. 2001) during the Albian-early Cenomanian.
It was recorded from the early Aptian of France
(Masure et al. 1998) and Colombia (Mejia Ve-
lasquez 2007), the late Aptian of Brazil (Pons
et al. 1996), the late Aptian-early Albian of Spain
(Peyrot et al. 2007), and the Aptian-Albian of
north-east Japan (Umetsu & Sato 2007). We
plot these reports of Cicatricosisporites sinuosus
on a paleogeographical map (Fig. 16), where we
note that its distribution is between 0º and 50º
in the Northern Hemisphere. is species seems
to have disappeared during the Cenomanian
transgression. In Africa, it lived in swamps (Al-
Ameri et al. 2001) or near the coast, in warm and
humid areas (Mahmoud et al. 2007). It seems
not to have lived in dry and arid areas further
in the south, and may have reached eastern Asia
by water or by following coastal areas from the
Tethysian regions.
Bisaccate pollen grains are poorly represented
and badly preserved in the assemblages, but we
encountered grains of the genera Abietineaepol-
lenites, Alisporites, Cedripites and Pityosporites.
Cycadopites is also common in the assemblages.
is type of pollen grain was produced by Bennet-
titales, Cycadales, Ginkgoales or Czekanowskiales.
e diff erent species we observed may represent
diff erent origins. Kimura (2000) considered that
Ginkgoales characterize the Tetori-type Prov-
ince. e grains we observed were most prob-
ably produced by Bennettitales or Cycadales,
but only a TEM study can enable more precise
identifi cation than the four orders known from
the macro fl ora.
We reported Eucommiidites troedssonii in our
assemblages. is species has previously been ob-
served in Japan by Umetsu & Sato (2007) from the
late Aptian-early Albian sediments of the Hiraiga
121
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
Formation (Miyako Group in north-east Honshu),
by Takahashi (1988) and Takahashi & Sugiyama
(1990) from the Upper Cretaceous sediments of
the Kuji and Futaba groups (north-east Honshu)
under the name Cupuliferoidaepollenites sp. e
latter, however, is a Tertiary angiospermous tri-
colpate pollen grain, and cannot be compared to
our species, which has a smooth exine.
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
60º N
0º 30º N
A
B
C
D
Cicatricosisporites sinuosus Hunt, 1985
+
F
IG
. 16. — Spatio-temporal distribution of Cicatricosisporites sinuosus Hunt, 1985: A, Purbeckian of England; B, pre-Aptian of Lebanon,
Israel, Libya, China, south-west Honshu (Japan); C, Aptian of France, Colombia and Brazil, Aptian-Albian of Spain, Egypt and north-
east Honshu (Japan), Albian of Libya and Tunisia; D, Albian-early Cenomanian of Iraq. Paleogeographical map during the Hauterivian
(120 My) in North polar stereographic projection, modifi ed from Smith & Briden (1977) and Masse et al. (1993).
122
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
COMPARISON WITH OTHER PALYNOFLORISTIC
DOMAINS
Brenner (1976) did not consider eastern Asia in
his model of palynological provinces, but by ex-
tension of his latitudinally defi ned provinces, we
can suppose that Japan would be included in the
South-Laurasian Province of temperate to humid
subtropical climate. We noted in our assemblages
a dominance of spores, with schizaeaceous spores
particularly diversifi ed. Classopollis, bisaccate pol-
len grains, monosulcate (Cycadopites) or polyplicate
(Ephedripites and Gneatceaepollenites) grains are also
present. ese observations are in accordance with
the characteristics Brenner defi ned for the South-
Laurasian Province.
Vakhrameev (1991) placed Japan in the East
Asia Province of his Euro-Sinian Region, but we
did not observe in our assemblages the abundance
of cheirolepidiaceous pollen grains which generally
characterizes this region. We note that Vakhrameev
(1991) defi ned the East Asia Province based on
studies on macrofl oras of Kimura (1987) and those
on palynofl oras of Takahashi (1974) and Miki
(1972) (Aptian-Albian of the Miyako Group), who
reported, for example, 70% of Classopollis in their
assemblages (Takahashi 1974). is diff erence in
age between the Choshi and Miyako groups can
explain the percentage gap (9-14% in our assem-
blages). However, it should be noted that climatic
change would not be a result of the migration of
the Outer Zone terranes, because cheirolepidiaceous
macrofossils are abundant in the Aptian Sasayama
Group of the Inner Zone (Yamada 2009).
Li (1980, 1983), Chen et al. (1982) and Li &
Liu (1994) noted the abundance and diversity of
bisaccate pollen grains and schizaeaceous spores, and
scarcity of Classopollis in the North China Province
they defi ned. e South China Province is charac-
terized by an abundance of Classopollis (up to 50%)
and cicatricose spores (about 10%) with common
Exesipollenites in the assemblages. e palynofl oras
we have described from the Choshi Group cannot
be linked with those from China.
Cerebropollenites mesozoicus is rare in our assem-
blages, whereas some authors (Herngreen et al.
1996; Nichols 2003) consider this genus to be a
common and distinctive element constituting the
Cerebropollenites Province where Japan has been
included. e scarcity of Cerebropollenites in both
formations can be attributed either to a taphonomic
bias or to local climatic eff ects, but further studies
are needed to test these possibilities.
In short, the palynofl oras of the Ashikajima and
Kimigahama formations can be included in the
South-Laurasian Province of Brenner (1976). For a
period of Upper Jurassic to Lower Cretaceous time,
Japan can also be placed in the East Asia Province
of the Euro-Sinian Region as Vakhrameev (1991)
suggested based on the micro- and macrofossils.
However, even though the macrofl oral data from
Choshi are consistent with the defi nition of this
province, our palynological data suggest that there
could be climatic diff erences on a fi ner scale than
previously considered.
P
ALEOECOLOGY
A shallow marine environment for the succession
has been suggested from sedimentological studies
and the abundance of molluscs in the Choshi area
(Obata et al. 1975; Katsura et al. 1984). Green algae
of the genera Tasmanites and Pterospermella (Prasi-
nophyceae), which lived in marine environments,
confi rm the interpreted sedimentary environments
of the Ashikajima and Kimigahama formations
from the palynological point of view.
In association with these marine elements, we
observed various terrestrial and freshwater paly-
nomorphs: Chomotriletes minor, Ovoidites parvus
(Zygnemataceae) and Botryococcus sp. cf. B. braunii
(Botryococcaceae), which lived in shallow fresh-
water environments (Grenfell 1995). e com-
position of the marine sediments, which contain
abundant organic matter of continental origin, is
consistent with a deposit in a coastal environment
as previously suggested. Terrestrial elements may
have been carried and dispersed by water or wind
more or less far from their habitat, what can ex-
plain the rare occurence of bisaccate pollen grains,
for example.
Pollen grains of the genus Classopollis, related to
the extinct conifer family Cheirolepidiaceae, are
common (9-14%) in the assemblages. Cheirole-
pidiaceous plants have been generally reported to
have lived near or along coastal areas. ey have
123
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
been considered to have formed mangroves, and
also been adapted to arid or semi-arid environments
(Srivastava 1976; Upchurch & Doyle 1981; Pons &
Kœniguer 1985). Classopollis grains are often ob-
served in tetrads in our assemblages, suggesting the
proximity of the producing plants. Cheirolepidia-
ceous twigs of Cupressinocladus obatae (Okubo &
Kimura 1991) and Frenelopsis choshiensis (Kimura
et al. 1985) have been reported from the Choshi
Group. Among the elements associated with the
palynomorphs, we observed fragments of tracheids
with araucarioid cross-fi elds and mixed radial pit-
ting which suggest the presence of this family.
e landscape reconstructed thanks to the paly-
nofl oral results is in accordance with that previously
suggested by the macroremains. However, many
of the species we identifi ed are rare (less than 1%
in the assemblage), and may indicate a diversity
of source areas. For example, the large number of
psilate trilete spores indicates a provenance not
only from coastal areas.
P
ALEOCLIMATE
In some paleoclimatic reconstitutions for the mid-
Cretaceous period, the Outer Zone of Japan has
been located in a humid tropical climatic belt, with
an arid zone in the south (Chumakov et al. 1995).
Kimura (1987) also suggested a tropical to subtropi-
cal and arid climate from the macrofl oral data.
e relative abundance of Classopollis in the Choshi
area and the presence of inaperturate pollen grains
(for example, Araucariaceae) and Gnetales, associ-
ated with low percentages of bisaccate pollen grains
and schizaeaceous spores (even if diversifi ed) suggest
a tropical to subtropical warm climate. Spores of
the genera Cyathidites and Deltoidospora are com-
mon to abundant, epiphyllous fungi of the genera
Microthyriacites and Phragmothyrites are present,
and parasitic epiphyllous fungi (hyphopodia) are
observed on thin cuticle fragments in the Ashika-
jima Formation. ese observations suggest locally
humid climatic conditions in a lacustrine environ-
ment. We suggest that the climate indicated had
marked dry and more humid seasons. Our results
probably correspond to para-autochthonous and
allochthonous elements derived from a variety of
habitats.
Haggart et al. (2006) suggested the existence of
a warm current on the eastern coastal areas of the
Outer Zone. is current may have infl uenced the
climate of these areas, and our results are consistent
with the hypothesis of an unusually warm oceanic
climate instead of a “real” tropical to subtropical
climate. However, while Haggart et al. (2006)
simply focused on the action of marine currents to
explain the fl oral diff erences observed in Japan, we
think these currents were not enough to create the
two distinct fl oristic provinces, even though they
may have infl uenced them (Legrand 2009). Our
results are in accordance with the hypothesis of a
moderate migration of the terranes constituting the
Outer Zone, or moderate climatic change during
the Early Cretaceous in Japan.
CONCLUSION
e Ashikajima and Kimigahama formations yield
rich assemblages, characterized by a predominance
of terrestrial palynomorphs associated with dino-
fl agellates and foraminiferal linings. e associated
particles are mainly woody fragments and altered
cuticles. e assemblages can be attributed to the
East Asia Province of the Euro-Sinian Region or
the South-Laurasian Province. However, they
cannot be linked to any adjacent palynological
area in Asia, and appear to be unique to the re-
gion. Many of the species identifi ed have been
reported previously from Europe, and we report
for the fi rst time the genus Manumia in Asia.
e genera Callialasporites and Eucommiidites,
generally common in the European palynofl oras,
are rare in both Ashikajima and Kimigahama
formations.
Many of the species encountered were originally
described in the palynofl oras of Canada and the
USA. Lycopodiumsporites dentimuratus previously
reported only from the Barremian-Albian of the
Potomac Group by Brenner (1963) is common
in Japan, and the genus Manumia reported from
the Jurassic of Canada and northern Europe is
present. ese North-American elements were
most probably carried by marine currents or wind
up to the Outer Zone of Japan.
124
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
is fi rst report of Barremian palynofl oras from
the Outer Zone of Japan confi rms the composition
of the vegetation described by Kimura (1987), but
not the arid climate that he suggested. We bring
new data to the knowledge of the Ryoseki Floras,
but other palynological studies are needed in Japan
to defi ne characteristics for those fl oras that appear
to be unique to Asia.
Acknowledgements
is study has been fi nanced by a scholarship for
Education and Research off ered by the Japanese
Government (Monbukagakusho: MEXT), and
by the University of Tokyo and Chuo University,
Tokyo, Japan. Many thanks to Prof. Jean Broutin,
Université Pierre et Marie Curie, for his kind advise
and correction of the manuscript. We are thankful
to Prof. David Batten and Prof. Hiromichi Hirano
for accepting to review this manuscript. eir help
greatly contributed to improve it.
REFERENCES
A
L
-A
MERI
T. K., A
L
-N
AJAR
T. K. & B
ATTEN
D. J. 2001. —
Palynostratigraphy and palynofacies indications of
depositional environments and source potential for
hydrocarbons: the mid-Cretaceous Nahr Umr and
lower Mauddud Formations, Iraq. Cretaceous Research
22: 735-742.
A
NDERSON J. M. & ANDERSON H. M. 1983. — Palaeo-
fl ora of southern Africa, Molteno Formation (Triassic).
Volume 1. A.A. Balkema, Rotterdam, 227 p.
A
RCHANGELSKY S. & GAMERRO J. C. 1967. — Pollen
grains found in Coniferous cones from the Lower
Cretaceous in Patagonia (Argentina). Review of Pal-
aeobotany and Palynology 5: 179-182.
A
SH S. R., LITWIN R. J. & TRAVERSE A. 1982. — e
Upper Triassic fern Phlebopteris smithii (Daugherty)
Arnold and its spores. Palynology 6: 203-219.
B
ALME B. E. 1995. — Fossil in situ spores and pollen
grains: an annotated catalogue. Review of Palaeobotany
and Palynology 87: 81-323.
B
ATTEN
D. J. & U
WINS
P. J. R. 1985. — Early-Late
Cretaceous (Aptian-Cenomanian) palynomorphs,
in THUSU B. & OWENS B. (eds), Palynostratigraphy
of north-east Libya. Journal of Micropalaeontology 4
(1): 151-168.
BJÆRKE T. 1993. — Palynological examination of samples
from Alaska. Alaska Geologic Materials Center Data
Report 210: 18 p.
B
ØE R., DAVIDSEN B. & SMELROR M. 2005. — Near-
shore Mesozoic basins along the central Nordland
coast, Norway. Geological Survey of Norway Report
2005.011: 52 p.
B
RENNER G. J. 1963. — e spores and pollen of the
Potomac Group of Maryland. Bulletin of the Depart-
ment of Geology, Mines and Water Ressources, Maryland
27, 215 p.
B
RENNER
G. J. 1976. — Middle Cretaceous fl oral prov-
inces and early migrations of angiosperms, in BECK
C.B. (ed.), Origin and Early Evolution of Angiosperms.
Columbia University Press, New York: 23-47.
C
HEN P., LI W., CHEN J., YE C., WANG Z., SHEN Y. &
S
UN D. 1982. — Sequences of fossil biotic groups
of Jurassic and Cretaceous in China. Scientia Sinica
B 52: 1011-1020.
C
HUMAKOV
N. M., Z
HARKOV
M. A., H
ERMAN
A. B.,
D
OLUDENKO
M. P., K
ALANDADZE
N. N., L
EBEDEV
E.
L., PONOMARENKO A. G. & RAUTIAN A. S. 1995. —
Climatic zones in the middle of the Cretaceous period.
Stratigraphy and Geological Correlation 3: 3-14.
C
OUPER R. A. 1958. — British Mesozoic microspores
and pollen grains, a systematic and stratigraphic study.
Palaeontographica 103 B: 75-179.
D
EÁK M. H. 1962. — Deux nouveaux genres de spore
de la série d’argiles et de marnes aptiennes. Bulletin
de la Société géologique de Hongrie 92: 230-235.
DETTMANN M. E. 1963. — Upper Mesozoic microfl oras
from southeastern Australia. Proceedings of the Royal
Society of Victoria 77 (1): 1-148.
D
ETTMANN M. E. 1994. — Cretaceous vegetation: the
microfossil record, in H
ILL R. S. (ed.), History of the
Australian vegetation: Cretaceous to recent. Cambridge
University Press, Cambridge: 143-170.
D
ETTMANN
M. E. & C
LIFFORD
H. T. 1992. — Phy-
logeny and biogeography of Ruff ordia, Mohria and
Anemia (Schizaeaceae) and Ceratopteris (Pteridaceae):
evidence from in situ and dispersed spores. Alcheringa
16: 269-314.
D
YBKJÆR K. 1991.— Palynological zonation and paly-
nofacies investigation of the Fjerritslev Formation
(Lower Jurassic-basal Middle Jurassic) in the Danish
Subbasin. Danmarks Geologiske Undersøgelse, Series
A 30: 1-150.
G
AMERRO J. C. 1965. — Morfología del polen de Ap-
terocladus lanceolatus Archang. (Coniferae) de la
Formación Baqueró, Provincia de Santa Cruz. Ameg-
hiniana 4: 133-138.
G
AMERRO
J. C. 1968. — Orbiculas (Corpusculos de
Ubisch) y membranas tapetales cutinizadas en cuatro
coníferas del Cretácico Inferior de Santa Cruz, Repú-
blica Argentina. Ameghiniana 5: 271-280.
G
EOLOGICAL
S
URVEY
OF
J
APAN
, A
IST
(ed.) 2010. —
Seamless Digital Geological Map of Japan 1: 200,000.
Feb. 1, 2010 version. Research Information Database
DB084, Geological Survey of Japan, National Insti-
125
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
tute of Advanced Industrial Science and Technology.
Source: http://www.gsj.jp.
G
OLOZOUBOV
V. V. , M
ARKEVICH
V. S. & B
UGDAEVA
E.V. 1999. — Early Cretaceous changes of vegeta-
tion and environment in East Asia. Palaeogeography,
Palaeoclimatology, Palaeoecology 153: 139-146.
G
RENFELL H. R. 1995. — Probable fossil Zygnemata-
cean algal spore genera. Review of Palaeobotany and
Palynology 84: 201-220.
G
UY
D. J. E. 1971. — Palynological investigations in the
Middle Jurassic of the Wilhelmsfålt boring, southern
Sweden. Publications from the Institutes of Mineralogy,
Paleontology, and Quaternary Geology, University of
Lund, Sweden 168: 1-104.
H
AAN P. J. DE 1997. — Early Cretaceous Palynomorphs
from the Southern Coastal Plain of Israel and a Study of
Striate Spores of the Fern Family Schizaeaceae. esis,
University of California, 277 p. (unpublished).
H
AGGART J. W., MATSUKAWA M. & ITO M. 2006. —
Paleogeographic and paleoclimatic setting of Lower
Cretaceous basins of East Asia and western North
America, with reference to the nonmarine strata.
Cretaceous Research 27: 149-167.
H
ASENBOEHLER
B. 1981. — Étude paléobotanique et
palynologique de l’Albien et du Cénomanien du « Bassin
occidental portugais » au sud de l’accident de Nazaré
(province d’Estrémadure, Portugal). èse de 3
e
cycle,
Université Pierre et Marie Curie, Paris, Mémoires des
Sciences de la Terre n° 81-29, 365 p.
H
AYAMI I. & OJI T. 1980. — Early Cretaceous Bivalvia
from the Choshi district, Chiba Prefecture, Japan.
Transactions and proceedings of the Palaeontological
Society of Japan, New Series 120: 419-448.
H
ERNGREEN G. F. W. & BOER K. F. DE 1974. — Paly-
nology of Rhaetian, Liassic and Dogger strata in
eastern Netherlands. Geologie en Mijnbouw 53 (6):
343-368.
H
ERNGREEN
G.F.W., K
EDVES
M., R
OVNINA
L.V. &
SMIRNOVA S.B. 1996. — Chapter 29C. Cretaceous
palynofl oral provinces: a review, in J
ANSONIUS J. &
M
CGREGOR D. C. (eds), Palynology: Principles and
Applications, Volume 3. American Association of
Stratigraphic Palynologists Foundation, Dallas: 1157-
1188.
HIROOKA K., UCHIYAMA S., DATE T., NANAI H., HATTORI
I. & NAKAJIMA T. 1983. — Paleomagnetic evidence
of accretion and tectonism of the Hida and the Cir-
cum-Hida terranes, central Japan, in HOWELL D. G.,
JONES D. L., COX A. & NUR A. (eds), Proceedings of
Circum-Pacifi c Terrane Conference. Stanford University
Publications in Geological Sciences: 115-117.
HIROOKA K., UCHIYAMA S., DATE T., KANAI H. &
N
AKAJIMA
T. 1985. — Paleomagnetic evidence for
accretion and bending tectonics of the Hida and the
Circum-Hida terranes, central Japan, in Howell D. G.
(ed.), Tectonostratigraphic Terranes of the Circum-Pacifi c
Region. Council for Energy and Mineral Resources,
Houston: 391-399.
H
OELSTAD
T. 1985. — Palynology of the uppermost
Lower to Middle Jurassic strata on Bornholm, Den-
mark. Bulletin of the Geological Society of Denmark
34: 111-132.
HUNT C. O. 1985. — Miospores from the Portland
Stone Formation and the lower part of the Purbeck
Formation (Upper Jurassic / Lower Cretaceous) from
Dorset, England. Pollen et Spores 27: 419-451.
IBRAHIM M. I. A. 1996. — Aptian-Turonian palynology
of the Ghazalat-1 well (GTX-1), Qattara Depres-
sion, Egypt. Review of Palaeobotany and Palynology
94: 137-168.
I
BRAHIM M. I. A., ABOUL ELA N. M. & KHOLEIF S. E.
2001. — Palynostratigraphy of Jurassic to Lower Cre-
taceous sequences from the Eastern Desert of Egypt.
Journal of African Earth Sciences 32 (2): 269-297.
I
SHIDA
K., K
OZAI
T., P
ARK
S. O. & M
ITSUGI
T. 2003. —
Gravel bearing radiolaria as tracers for erosional
events: a review of the status of recent research in
SW Japan and Korea. Journal of Asian Earth Sciences
21: 909-920.
I
SOZAKI Y. 1996. — Anatomy and genesis of a subduc-
tion related orogen: a new view of geotectonic sub-
division and evolution of the Japanese Islands. e
Island Arc 5: 289-320.
I
SOZAKI
Y. 1997. — Jurassic accretion tectonics of Japan.
e Island Arc 6: 25-51.
I
SOZAKI Y., AOKI K., NAKAMA T. & YANAI S. 2010. —
New insight into a subduction-related orogen: reap-
praisal on geotectonic framework and evolution of the
Japanese Islands. Gondwana Research 18: 82-105.
I
TO
M. & M
ATSUKAWA
M. 1997. — Diachronous evolu-
tion of third-order depositional sequences in the Early
Cretaceous forearc basins: shallow marine and paralic
successions in the Sanchu and Choshi basins, Japan.
Memoir of the Geological Society of Japan 48: 60-75.
J
ERSEY
N. J.
DE
1962. — Triassic spores and pollen
grains from the Ipswich Coalfi eld. Geological Survey
of Queensland Publication 307, 18 p.
K
ASE T. & MAEDA H. 1980. — Early Cretaceous Gas-
tropoda from the Choshi district, Chiba Prefecture,
central Japan. Transactions and Proceedings of the Pal-
aeontological Society of Japan 118: 291-324.
K
ATSURA
Y., M
ASUDA
F. & O
BATA
I. 1984. — Storm
dominated shelf sea from the Lower Cretaceous
Choshi Group, Japan. Annual Report of the Institute
of Geoscience, University of Tsukuba 10: 92-95.
K
IMURA T. 1958. — Paleofl oristic provinces recognized
in the Onychiopsis Series. Journal of the Geological
Society of Japan 64: 687.
K
IMURA T. 1987. — Recent knowledge of Jurassic and
Early Cretaceous fl oras in Japan and phytogeography
of this time in East Asia. Bulletin of the Tokyo Gakugei
University, Section IV 39: 87-115.
126
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
K
IMURA
T. 2000. — Early Cretaceous climatic provinces
in Japan and adjacent regions on the basis of fossil
land plants, in O
KADA
H. & M
ATEER
N. J. (eds),
Cretaceous Environments of Asia. Elsevier Science BV,
Amsterdam: 155-161.
K
IMURA
T. & O
HANA
T. 1985. — Zamites choshiensis
sp. nov. from the Lower Cretaceous Choshi Group,
in the Outer Zone of Japan. Proceedings of the Japan
Academy, Series B 61: 352-355.
K
IMURA
T. & O
HANA
T. 1997. — Catalogue of the
Late Jurassic and Early Cretaceous plant-taxa in
Japan. Memoir of the Geological Society of Japan 48:
176-188.
K
IMURA T., OKUBO A. & MIYAHASHI H. 1991. — Cu-
ticular study of Ptilophyllum leaves from the Lower
Cretaceous Choshi Group, in the Outer Zone of Ja-
pan, Bulletin of the National Science Museum, Tokyo,
Series C [Geology] 17: 129-152.
KIMURA T., SAIKI K. & ARAI T. 1985. — Frenelopsis
choshiensis sp. nov., a Cheirolepidiaceous conifer from
the Lower Cretaceous Choshi Group in the Outer
Zone of Japan. Proceedings of the Japan Academy,
Series B 61: 426-429.
K
IMURA
T. & S
EKIDO
S. 1976. — Mesozoic plants from
the Akaiwa Formation (Upper Neocomian), the
Itoshiro Group, central Honshu, Japan. Transactions
and Proceedings of the Palaeontological Society of Japan,
New Series 103: 343-378.
K
IMURA
T. & S
EKIDO
S. 1978. — Addition to the
Mesozoic plants from the Akaiwa Formation (Upper
Neocomian), the Itoshiro Group, central Honshu,
Inner Zone of Japan. Transactions and Proceedings
of the Palaeontological Society of Japan, New Series
109: 259-279.
K
IMYAI
A. 1966. — New plant microfossils from the
Raritan Formation (Cretaceous) in New Jersey. Mi-
cropaleontology 12 (4): 461-476.
K
NOX E. M. 1950. — e spores of Lycopodium, Phyl-
loglossum, Selaginella and Isoetes, and their value in the
study of microfossils of Palaeozoic age. Transactions of
the Botanical Society of Edinburgh 35 (3): 207-357.
K
OJIMA
S. 1989. — Mesozoic terrane accretion in north-
east China, Sikhote-Alin and Japan regions. Palaeogeog-
raphy, Palaeoclimatology, Palaeoecology 69: 213-232.
K
OPPELHUS
E. B. 1992. — Palynologisk underssgelse
af nedre jurassiske sedimenter i Anholt-4 boringen.
Dansk Geologisk Forening Årsskrift for 1990-91: 73-78
(in Danish, summary in English).
KOPPELHUS E.B. & NIELSEN L.H. 1994. — Palyno-
stratigraphy and palaeoenvironments of the Lower
to Middle Jurassic Bagå Formation of Bornholm,
Denmark. Palynology 18: 139-194.
K
OPPELHUS
E. B. & B
ATTEN
D. J. 1996. — Chapter 20C.
Application of a palynomorph zonation to a series of
short borehole sections, Lower to Middle Jurassic,
Øresund, Denmark, in JANSONIUS J. & MCGREGOR
D. C. (eds), Palynology: Principles and Applications,
Volume 2. American Association of Stratigraphic
Palynologists Foundation, Dallas: 779-793.
K
OPPELHUS T. & DAM G. 2003. — Palynostratigraphy
and palaeoenvironments of the Rævekløft, Gule Horn
and Ostreaelv Formations (Lower-Middle Jurassic),
Neill Klinter Group, Jameson Land, East Greenland.
Geological Survey of Denmark and Greenland Bulletin
1: 723-775.
K
OPPELHUS
E. B. & H
ANSEN
C.F. 2003. — Palynostratig-
raphy and palaeoenvironment of the Middle Jurassic
Sortehat Formation (Neill Klinter Group), Jameson
Land, east Greenland. Geological Survey of Denmark
and Greenland Bulletin 1: 777-811.
KRUTZSCH W. 1963. — Atlas der mittel- und jungtertiären
dispersen Sporen und Pollen-sowie der Mikroplankton-
formen des nördlichen Mitteleuropas. Lieferung III.
Sphagnaceoide und Selaginellaceoide Sporenformen. Veb
Gustav Fischer Verlag Jena, Berlin, 128 p.
K
UMAR
P. 1973. — e sporae dispersae of Jabalpur
stage, Upper Gondwana, India. e Palaeobotanist
20 (1): 91-126.
L
EE Y. I. 2008. — Paleogeographic reconstructions of
the East Asia continental margin during the Middle
to Late Mesozoic. e Island Arc 17 (4): 458-470.
L
EE
Y. I. & K
IM
J. Y. 2005. — Provenance of the Hayang
Group (Early Cretaceous) in the Yeongyang Subbasin,
SE Korea and its bearing on the Cretaceous palaeo-
geography of SW Japan. Palaeogeography, Palaeocli-
matology, Palaeoecology 228: 278-295.
L
EGRAND
J. 2009. — Palynologie des dépôts Jurassique
supérieur et Crétacé inférieur du Japon, et provinces
paléofl oristiques du sud-est asiatique. èse de 3
e
cy-
cle, Université Pierre et Marie Curie, Paris, 366 p.
(unpublished).
L
I
W.-B. 1980. — Microfl oral areas of Early Creta-
ceous in China, in CHALONER W. G. & SHEERIN A.
(eds), Proceedings of the 5
th
International Palynological
Congress, Cambridge. International Commission for
Palynology: 1-8.
L
I W.-B. 1983. — Microfl oral areas of Early Cretaceous
in China, in L
U Y., MU A. & WANG Y. (eds), Palaeo-
biogeographic Provinces of China. Science Press, Beijing:
142-151 (in Chinese, summary in English).
L
I
W.-B. & L
IU
Z. 1994. — e Cretaceous palynofl oras
and their bearing on stratigraphic correlation in China.
Cretaceous Research 15: 333-365.
L
INDSTRÖM S. & ERLSTRÖM M. 2007. — Dating and
correlating potential aquifers for geothermal energy,
CO
2
-, and energy storage, within the Late Triassic-
Early Cretaceous succession in the Swedish part of
the Danish Basin. Geological Survey of Sweden Report
60-1344/2004: 47 p.
L
UND J. J. & PEDERSEN K. R. 1984. — Palynology of
the marine Jurassic formations in the Vardekløft ra-
vine, Jameson Land, East Greenland. Bulletin of the
127
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
Geological Society of Denmark 33: 371-400.
M
AHMOUD
M. S.& D
EAF
A. S. 2007. — Cretaceous paly-
nology (spores, pollen and dinofl agellate cysts) of the
Siqeifa 1-X Borehole, northern Egypt. Rivista Italiana
di Paleontologia e Stratigrafi a 113 (2): 203-221.
M
AHMOUD
M. S., S
OLIMAN
H. A. & D
EAF
A. S. 2007. —
Early Cretaceous (Aptian-Albian) palynology of the
Kabrit-1 Borehole, onshore northern Gulf of Suez,
Egypt. Revista Española de Micropaleontología 39
(3): 169-187.
M
ARUYAMA S., ISOZAKI Y., KIMURA G. & TERABAYASHI
M. 1997. — Paleogeographic maps of the Japanese
Islands: plate tectonics synthesis from 750 Ma to the
present. e Island Arc 6: 121-142.
M
ASSE J. P., BELLION Y., BENKHELIL J., DERCOURT J.,
G
UIRAUD R. & RICOU L. E. 1993. — Lower Aptian
Palaeoenvironments (114 to 112 Ma), in D
ERCOURT
J., RICOU L. E. & VRIELYNCK B. (eds), Atlas Tethys
Palaeoenvironmental Maps, Maps. BEICIP-FRANLAB,
Rueil-Malmaison.
M
ASURE E., RAYNAUD J.-F., PONS D. & DE RENEVILLE
P. 1998. — Palynologie du stratotype historique
de l’Aptien inférieur dans la région de Cassis-La
Bédoule (SE France). Géologie méditerranéenne 25
(3-4): 263-287.
M
ATSUKAWA M. & OBATA I. 1993. — e ammonite
Crioceratites (Paracrioceras) and Shasticrioceras from
the Barremian of Southwest Japan. Palaeontology
36: 249-266.
M
ATSUKAWA
M. & F
UKUI
M. 2009. — Hauterivian–
Barremian marine molluscan fauna from the Tetori
Group in Japan and Late Mesozoic marine transgres-
sions in east Asia. Cretaceous Research 30: 615-631.
M
ATSUOKA M., TAKAHASHI O., HAYASHI K., ITO M. &
K
ONOVALOY V. P. 1997. — Early Cretaceous paleo-
geography of Japan, based on tectonic and faunal
data, in OKADA H., HIRANO H., MATSUKAWA M. &
K
IMINAMI
K. (eds), Cretaceous Environmental Change
in East and South Asia (IGCP 350) – Contributions
from Japan. Memoir of Geological Society of Japan
48: 29-42.
M
CGREGOR D. C. 1965. — Illustrations of Canadian
fossils Triassic, Jurassic and Lower Cretaceous spores
and pollen of Arctic Canada. Geological Survey of
Canada papers 64-65: 1-32.
M
EJIA VELASQUEZ P. J. 2007. — Floral composition of a
Lower Cretaceous paleotropical ecosystem inferred from
quantitative palynology. esis, University of Florida,
85 p. (unpublished).
M
IKI
A. 1972. — Palynological study of the Kuji Group
in northeastern Honshu, Japan. Journal of the Faculty
of Science, Hokkaido University, Series 4, Geology and
mineralogy 15 (3-4): 513-604.
M
ITSUGI
T., I
SHIDA
K., W
OO
B. G., C
HANG
K. H.,
P
ARK
S. O. & H
IRANO
H. 2001. — Radiolarian-
bearing conglomerate from the Hayang Group, the
Kyongsang Supergroup, southeastern Korea. Journal
of Asian Earth Sciences 19: 751-763.
M
URILLO
M. T. & B
LESS
M. J. M. 1974. — Spores of re-
cent Colombian Pteridophyta. I. Trilete spores. Review
of Palaeobotany and Palynology 18 (3-4): 223-269.
NICHOLS D. J. 2003. — Biodiversity changes in Cre-
taceous palynofl oras of eastern Asia and western
North America. Journal of Asian Earth Sciences 21:
823-833.
N
ISHIDA M. 1960. — Notes on the Cretaceous woods
found in the Choshi Peninsula. Bulletin of the Marine
Laboratory, Chiba University 2: 22-31.
N
ISHIDA
M. 1962. — On some petrifi ed plants from
the Cretaceous of Choshi, Chiba Prefecture. Japanese
Journal of Botany 18: 87-104.
O
BATA
I., H
AGIWARA
S. & K
AMIKO
S. 1975. — Geological
age of the Cretaceous Choshi Group. Bulletin of the
National Science Museum, Tokyo, Series C [Geology] 1
(1): 17-33 (in Japanese, summary in English).
O
BATA
I., M
AIYA
S., I
NOUE
Y. & M
ATSUKAWA
M. 1982. —
Integrated mega- and microfossil biostratigraphy of
the Lower Cretaceous Choshi Group, Japan. Bulletin
of the National Science Museum, Tokyo, Series C [Geol-
ogy] 8 (4): 145-179.
O
BATA I. & MATSUKAWA M. 2009. — Supplementary
description of the ammonoids from the Barremian to
the Albian of the Choshi Peninsula, Japan. Cretaceous
Research 30: 253-269.
O
KUBO
A. & K
IMURA
T. 1991. — Cupressinocladus obatae
sp. nov., from the Lower Cretaceous Choshi Group,
in the Outer Zone of Japan. Bulletin of the National
Science Museum, Series C 17: 91-109.
OMRAN A.M., SOLIMAN H.A. & MAHMOUD M.S.
1990. — Early Cretaceous palynology of three bore-
holes from northern Western Desert (Egypt). Review
of Palaeobotany and Palynology 66: 293-312.
O
TOFUJI
Y., M
ATSUDA
T. & N
OHDA
S. 1985. — Opening
mode of the Japan Sea inferred from the Palaeomag-
netism of the Japan Arc. Nature 317: 603-604.
O
TOH
S. 1998. — Late Jurassic to Early Cretaceous
tectonic inversion in eastern margin of Asia, in KIM
Y. H. (ed.), International Symposium on Earth and En-
vironmental Sciences. Basic Science Research Institute
of Gyeongsang National University, Jinju: 19-36.
O
TOH S. & SASAKI M. 1998. — “Asia” and the “Japa-
nese Islands” in Paleo-Mesozoic time. Memoirs of the
Geological Society of Japan 50: 159-176 (in Japanese,
summary in English).
PEYROT D., RODRÍGUEZ-LÓPEZ J.P., BARRÓN E. &
MELÉNDEZ N. 2007. — Palynology and biostratigra-
phy of the Escucha Formation in the Early Cretaceous
Oliete sub-basin, Teruel, Spain. Revista Española de
Micropaleontología 39 (1-2): 135-154.
PFLUG H. D. 1953. — Zur Entstehung und Entwicklung
des angiospermiden Pollens in der Erdgeschichte.
Palaeontographica 95 B: 60-171.
128
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
PIERCE R. L. 1961. — Lower Upper Cretaceous plant
microfossils from Minnesota. Minnesota Geological
Survey Bulletin 42, 86 p.
P
OCOCK
S. A. J. 1970. — Palynology of the Jurassic
sediments of western Canada. Pt. 1. Terrestrial spe-
cies. Palaeontographica 130 B: 12-136.
P
ONS D., BERTHOU P.-Y., DE MELO-FILGUEIRA J.-B. &
A
LCANTARA SAMPAIO J.-J. 1996. — Palynologie des
unités « Fundão », « Crato » et « Ipubi » (Aptien supérieur
à Albien inférieur-moyen, bassin d’Araripe, Nord-Est
du Brésil) : enseignements paléoécologiques, strati-
graphiques et climatologiques, in J
ARDINÉ
S.,
DE
KLASZ I. & DEBENAY J. P. (eds), Géologie de l’Afrique
et de l’Atlantique Sud : Compte rendu des Colloques
de géologie d’Angers, 1994. Bulletin des Centres de
Recherches Exploration-Production Elf Aquitaine,
Mémoire 16: 383-401.
PONS D. & KŒNIGUER J.-C. 1985. — Les Gymnospermes.
Clartés : l’Encyclopédie du Présent, éditions techniques,
Paris, fascicule 4070, 20 p.
POTONIÉ R. 1956. — Synopsis der Gattungen der Sporae
dispersae. I. Teil: Sporites. Beihefte zum Geologischen
Jahrbuch 23: 1-103.
POTONIÉ R. 1958. — Synopsis der Gattungen der Sporae
dispersae. II. Teil: Sporites (Nachträge), Saccites, Ale-
tes, Praecolpates, Polyplicates, Monocolpates. Beihefte
zum Geologischen Jahrbuch 31: 1-114.
POTONIÉ R. 1960. — Synopsis der Gattungen der Sporae
dispersae. III. Teil: Nachträge Sporites, Fortsetzung
Pollenites mit Generalregister zu Teil I-III. Beihefte
zum Geologischen Jahrbuch 39: 1-189.
P
OTONIÉ
R. 1966. — Synopsis der Gattungen der
Sporae dispersae. IV. Teil: Nachträge zu allen Grup-
pen (Turmae). Beihefte zum Geologischen Jahrbuch
72: 1-244.
P
OTONIÉ
R. 1970a. — Synopsis der Gattungen der
Sporae dispersae. V. Teil: Nachträge zu allen Grup-
pen (Turmae). Beihefte zum Geologischen Jahrbuch
87: 1-222.
POTONIÉ R. 1970b. — Synopsis der Gattungen der
Sporae dispersae. VI. Teil: Nachträge zu allen Grup-
pen (Turmae). Beihefte zum Geologischen Jahrbuch
94: 1-176.
POTONIÉ R. 1975. — Synopsis der Gattungen der Sporae
dispersae. VII. Teil: Nachträge zu allen Gruppen
(Turmae). Fortschritte in der Geologie von Rheinland
und Westfalen 25: 23-150.
P
OTONIÉ R. & KREMP G. 1954. — Die Gattungen der
paläozoischen Sporae dispersae und ihre Stratigraphie.
Geologisches Jahrbuch 69: 111-194.
P
OTONIÉ
R. & K
REMP
G. 1955. — Die Sporae dispersae
des Ruhrkarbons, ihre Morphographie und Strati-
graphie mit Ausblicken auf Arten anderer Gebiete
und Zeitabschnitte Teil I. Palaeontographica 98 B
(1-3): 1-136.
P
U R. & WU H. 1982. — Sporo-pollen from the Late
Mesozoic beds in eastern Heilongjiang Province. Bul-
letin of the Shenyang Institute of Geology and Mineral
Resources, Chinese Academy of Geological Sciences 5:
383-456 (in Chinese, summary in English).
P
U
R. & W
U
H. 1985. — e palynological assemblages
of the Hingganling and Zhalainuouer Groups in
Hinggan Ling Region, northeast China and their
stratigraphical signifi cance. Bulletin of the Shenyang
Institute of Geology and Mineral Ressources, Chinese
Academy of Geological Sciences 11: 47-113 (in Chinese,
summary in English).
S
AAD
S. I. 1978. — Palynological studies in the Egyptian
Western Desert. Pollen et Spores 20 (2): 261-301.
S
CHRANK E. & IBRAHIM M. I. A. 1995. — Cretaceous
(Aptian-Maastrichtian) palynology of foraminifera-
dated wells (KRM-1, AG-18) in northwestern Egypt.
Berliner Geowissenschaftliche Abhandlungen, Reihe A
177: 1-44.
S
CHULZ E. 1967. — Sporenpalaontologische Untersu-
chungen ratoliassischer Schichten im Zentralteil des
Germanischen Beckens. Paläontologische Abhandlungen
2 B (3): 542-633.
S
CHWEITZER H. J., ASHRAF A. R. & WEISS M. 1987. —
Korrelation der Sporenzonen der Oberen Trias und
des Jura im Mittleren Orient und in Süddeutschland.
Geologische Rundschau 76 (3): 923-943.
SEIDENKRANTZ M.-S., KOPPELHUS E.B. & RAVN-SØRENSEN
H. 1993. — Biostratigraphy and palaeoenvironmen-
tal analysis of a Lower to Middle Jurassic succession
on Anholt, Denmark. Journal of Micropalaeontology
12: 201-218.
S
HIKAMA
T. & S
UZUKI
S. 1972. — Stratigraphy and
tectonic development mainly of Cretaceous forma-
tions of Choshi Peninsula, Chiba Prefecture. Science
Reports of the Yokohama National University, Section II
19: 133-157 (in Japanese, summary in English).
S
INGH C. 1964. — Microfl ora of the Lower Cretaceous
Mannville Group, east-central Alberta. Research Council
of Alberta Bulletin 15, 239 p.
S
INGH H. P. & KUMAR P. 1969. — Reappraisal of some
subsaccate fossil pollen genera. e Palaeobotanist
17 (1): 80-92.
S
KOG J. E. 1980. — e genus Ruff ordia (Schizaeaceae)
from the Lower Cretaceous Potomac Group. Botani-
cal Society of America, Miscellaneous Series Publications
158: 106.
SKOG J. E. 1982. — Pelletixia amelguita, a new species of
fossil fern in the Potomac Group (Lower Cretaceous).
American Fern Journal 72 (4): 115-121.
S
MITH
A. G. & B
RIDEN
J. C. 1977. — Mesozoic and
Cenozoic Paleocontinental Maps. Cambridge University
Press, Cambridge, 63 p.
S
RIVASTAVA S. K. 1976. — e fossil pollen genus Clas-
sopollis. Lethaia 9 (4): 437-457.
STEFANOWICZ S. 2008. — Palyno-stratigraphy and
palaeoclimatic analysis of the Lower-Middle Jurassic
129
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
(Pliensbachian-Bathonian) of the Inner Hebrides, NW
Scotland. Master thesis, Lund University, 35 p. (un-
published).
SUKH-DEV 1980. — Evaluation of in situ spores and
pollen grains from the Jurassic-Cretaceous fructifi ca-
tions, in BHARADWAJ D. C., LELE K. M. & KAR R.
K. (eds), Proceedings of the 4
th
International Palyno-
logical Congress, Lucknow. Birbal Sahni Institute of
Paleobotany 2: 753-768.
T
AKAHASHI
K. 1974. — Palynology of the Upper Aptian
Tanohata Formation of the Miyako Group, northeast
Japan. Pollen et Spores 16: 535-564.
TAKAHASHI K. 1988. — Palynology of the Upper Cretaceous
Futaba Group. Bulletin of the Faculty of Liberal Arts,
Nagasaki University, Natural Science 28: 67-183.
T
AKAHASHI
K. & S
UGIYAMA
R. 1990. — Palynomorphs
from the Santonian Uge Member of the Taneichi
Formation, northeastern Japan. Bulletin of the Faculty
of Liberal Arts, Nagasaki University, Natural Science
30: 133-573.
T
EKLEVA M. V. & KRASSILOV V. A. 2009. — Compara-
tive pollen morphology and ultrastructure of modern
and fossil Gnetophytes. Review of Palaeobotany and
Palynology 156: 131-139.
T
RINCÃO P. R . P. 1990. — Esporos e pólenes do Cretácio
inferior (Berriasiano-Aptiano) de Portugal: paleontologia
e biostratigrafi a. Tese, Universidade Nova de Lisboa,
2 volumes, 312 p. (unpublished).
TRYON A. F. & LUGARDON B. 1991. — Spores of the
Pteridophyta: Surface, Wall Structure, and Diversity
Based on Electron Microscope Studies. Springer-Verlag,
New York, 648 p.
T
RYON R. M. & TRYON A. F. 1982. — Ferns and Allied
Plants, with Special Reference to Tropical America.
Springer-Verlag, New York, 857 p.
U
METSU
K. & M
ATSUOKA
A. 2003. — Early Cretaceous
fossil spores and pollen from the Tetori Group in
the upper reaches of the Kuzuryu River, Fukui Pre-
fecture, central Japan. e Journal of the Geological
Society of Japan 109: 420-423 (in Japanese, summary
in English).
U
METSU K. & SATO Y. 2007. — Early Cretaceous ter-
restrial palynomorph assemblages from the Miyako
and Tetori Groups, Japan, and their implication to
paleophytogeographic provinces. Review of Palaeo-
botany and Palynology 144 (1-2): 13-24.
U
PCHURCH G. R. & DOYLE J. 1981. — Paleoecology of
the conifers Frenelopsis and Pseudofrenelopsis (Chei-
rolepidiaceae) from the Cretaceous Potomac Group
of Maryland and Virginia, in R
OMANS C. (ed.), Geo-
botany II. Plenum, New York: 167-202.
U
WINS P. J. R. & BATTEN D. J. 1988. — Early to mid-
Cretaceous palynology of Northeast Libya, in E
L
-
A
RNAUTI
A., O
WENS
B. & T
HUSU
B. (eds), Subsurface
Palynostratigraphy of Northeast Libya. Garyounis Uni-
versity Publications, Benghazi: 215-257.
V
AKHRAMEEV
V. A. 1991. — Jurassic and Cretaceous Flo-
ras and Climates of the Earth. Cambridge University
Press, Cambridge, 318 p.
V
AN KONIJNENBURG-VAN CITTERT J. H. A 1971. — In
situ gymnosperm pollen from the Middle Jurassic of
Yorkshire. Acta Botanica Neerlandica 20: 1-96.
YAMADA T. 2009. — Vegetational history through Mid-
dle Jurassic to Early Cretaceous in Japan. Bunrui 9:
115-121 (in Japanese).
YAMADA T. & UEMURA K. 2008. — e plant fossils
from the Kaizara Formation (Callovian, Jurassic) of the
Tetori Group in the Izumi district, Fukui Prefecture,
central Japan. Paleontological Research 12: 1-17.
Y
ASKAWA K. 1975. — Palaeolatitude and relative posi-
tion of south-west Japan and Korea in the Cretaceous.
Geophysical Journal of the Royal Astronomical Society
43: 835-846.
Y
I
M.-S., C
HOI
S.-J. & Y
UN
H. 1993. — Cretaceous
palynomorphs from the Iljik Formation in the Eui-
seong area, Korea. Journal of the Paleontological Society
of Korea 9 (2): 166-179.
ZAKLINSKAYA E. D. 1957. — Stratigraphic Importance
of Gymnospermic Pollen in Cenozoic Deposits of Pav-
lodarirtish and Northern Voraral Areas. Trudy Geo-
logicheskogo Instituta Akademii Nauk., USSR 6:
1-219 (in Russian).
Submitted on 15 April 2010;
accepted on 13 January 2011.
130
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
APPENDICES
TABLE 1. — List of all the taxa encountered during our investigation, in nomenclatural order, with their occurrence in the Ashikajima
Formation (A) and in the Kimigahama Formation (K) and their fi gurative position. Abbreviations: –, one grain; C, common; P, present;
R, rare.
Occurrence
Figures A K
PROXIMEGERMINANTES
Triletes Azonales
Acavatitriletes
Azonotriletes
Laevigati, Quasilaevigati
Biretisporites potoniaei (Delcourt & Sprumont, 1955) Delcourt, Dettmann & Hughes, 1963 5A, B
PP
Cibotiumspora paradoxa (Maljavkina, 1949) Chang, 1965 5P
RR
Cyathidites australis Couper, 1953 5G
PP
Cyathidites minor Couper, 1953 5F
PP
Cyathidites rarus (Bolkhovitina, 1953) Deák, 1964 5H
–
Deltoidospora hallii Miner, 1935
PP
Todisporites major Couper, 1958 5D
–
Todisporites miser Takahashi, 1988 5C
–
Undulatisporites undulapolus Brenner, 1963
R
Undulatisporites sinuosis Groot & Groot, 1962
–
Apiculati
Granulati, Scabrati
Concavissimisporites punctatus (Delcourt & Sprumont, 1955) Brenner, 1963 5Q
R
Concavissimisporites variverrucatus (Couper, 1958) Brenner, 1963 5K
RR
Osmundacidites wellmanii Couper, 1953 5L
P
R
Verrucati
Converrucosisporites sp. 5N, O
–
Manumia japonica n. sp. 5A-F, I, J
PP
Trilites bossus Couper, 1958 5M
–
Uvaesporites sp.
–
Verrucosisporites densus (Bolkhovitina, 1956) Pocock, 1970 5I, J
–
Baculati
Baculatisporites comaumensis (Cookson, 1953) Potonié, 1956
RR
Baculatisporites sp. 6N
R
Neoraistrickia sp. 1 7A, B
–
Neoraistrickia sp. 2 7C, D
–
Neoraistrickia sp. 3 7G, J
RR
Pilosisporites verus Delcourt & Sprumont, 1955 6G
R
Pilosisporites brevis Delcourt & Sprumont, 1955 6K
R
Tuberositriletes sp. 6H, L, M
–
Nodati
Echinatisporis varispinosus (Pocock, 1962) Srivastava, 1975
PP
Murornati
Cicatricosisporites annulatus Archangelsky & Gamerro, 1966
R
Cicatricosisporites hughesi Dettmann, 1963 8Q, R
RR
Cicatricosisporites mohrioides Delcourt & Sprumont, 1955 9I, J
R
Cicatricosisporites sinuosus Hunt, 1985 8K, L, S, T
PP
Cicatricosisporites sp. 1 9A, B
R
Cicatricosisporites sp. 2 9C, D
R
Cicatricosisporites sp. 3 9E, F
R
Cicatricosisporites sp. 4 9G, H
R
Foveosporites ryosekiensis n. sp. 8D, E, H-J
PP
Klukisporites variegatus Couper, 1958 7E, F, I, L, M
PP
Lycopodiumsporites crassimacerius Hedlund, 1966
–
Lycopodiumsporites dentimuratus Brenner, 1963 7H, K, N-P
CC
Microreticulatisporites sp. 1 8M, P
R
131
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
Occurrence
Figures A K
Microreticulatisporites sp. 2 8N, O
R
Retitriletes austroclavatidites (Cookson, 1953) Döring, Krutzsch, Mai & Schulz in
Krutzsch, 1963
8F, G
RR
Retitriletes sp. 8A-C
R
Ruffordiaspora australiensis (Cookson, 1953) Dettmann & Clifford, 1992 9K-M, P
PP
Triletes Zonales
Auritotriletes
Auriculati
Matonisporites sp. 1
–
Matonisporites sp. 2
9O P P
Appendiciferi
Nodosisporites makotoi n. sp.
10A-F P P
Nodosisporites choshiensis n. sp.
10G-K P P
Plicatella sp.
9N, Q R
Zonotriletes
Cingulati
Cingulatisporites psilatus Groot & Penny, 1960
10O, P R
Cingulatisporites sp. 1
10L-N P P
Cingulatisporites sp. 2
12A R R
Cingutriletes sp.
10Q R
Contignisporites burgeri Filatoff, McKellar & Price in Filatoff & Price, 1988
12G, M R
Contignisporites cooksonii (Balme, 1957) Dettmann, 1963
12D R R
Distaltriangulisporites sp.
12B, C R
Polycingulatisporites reduncus (Bolkhovitina, 1953) Playford & Dettmann, 1965
12H, I R R
Patellasporites sp.
–
Tricrassati
Coronatispora valdensis (Couper, 1958) Dettmann, 1963
–
Zonolaminatitriletes
Zonati
Aequitriradites spinulosus (Cookson & Dettmann, 1958) Cookson & Dettmann, 1961
–
Aequitriradites verrucosus (Cookson & Dettmann, 1958) Cookson & Dettmann, 1961
12F R
Aequitriradites sp.
12E R
Triporoletes reticulatus (Pocock, 1962) Playford, 1971
RR
Monoletes
Acavatomonoletes
Azonomonoletes
Laevigatomonoleti
Laevigatosporites ovatus Wilson & Webster, 1946
12K R R
Laevigatosporites sp.
12J –
Incertae sedis
Incertae sedis in Hasenboehler (1981)
5E –
Incertae sedis sp. 1
13N, O R
VARIEGERMINANTES
Saccites
Monosaccites
Inaperturati (= Saccizonati)
Callialasporites dampieri (Balme, 1957) Sukh-Dev, 1961
12R R
Callialasporites sp. cf C. ugensis Takahashi & Sugiyama, 1990
12N –
Cerebropollenites mesozoicus (Couper, 1958) Nilsson, 1958
12P –
TABLE 1. — Continuation.
132
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
Occurrence
Figures A K
Disaccites
Disacciatrileti
Alisporites thomasii (Couper, 1958) Pocock, 1962
13B R
Alisporites sp. cf. A. australis de Jersey, 1962
–
Alisporites sp.
12Q –
Pityosporites sp. cf. P. constrictus Singh, 1964
13A – –
Pityosporites sp. cf. P. piniformis (Zaklinskaya, 1957) Takahashi & Sugiyama, 1990
––
Aletes and Kryptoaperturates
Azonaletes
Psilonapiti
Inaperturopollenites sp.
13E P P
Granulonapiti
Araucariacites australis Cookson, 1947 ex Couper, 1953
PP
Balmeiopsis limbatus (Balme, 1957) Archangelsky, 1977
13D P P
Spheripollenites psilatus Couper, 1958
RR
Circumpollini
Classopollis torosus (Reissinger, 1950) Couper, 1958 emend. Burger, 1965
13F-H C C
Corollina sp.
13M R R
Plicates
Costates (= Polyplicates)
Costati
Ephedripites montanaensis Brenner, 1968
13K R
Gnetaceaepollenites sp.
13L –
Monocolpates (Monosulcites) and Zonocolpates
Quasilaevigati and Microsculptati
Cycadopites minimus (Cookson, 1947) Pocock, 1970
13I R R
Cycadopites sp.
13J R
Tricolpates, Triptyches
Heterotricolpati, Praecolpati
Eucommiidites minor Groot & Penny, 1960
––
Eucommiidites troedssonii (Erdtman, 1948) Potonié, 1958
12O R R
Poroses
Monoporines
Exesipollenites tumulus Balme, 1957
12L –
Incertae sedis
Incertae sedis sp. 2
13C –
ALGAE CHLOROPHYTA
Charophyceae
Chomotriletes minor (Kedves, 1961) Pocock, 1970
R
Ovoidites parvus (Cookson & Dettmann, 1959) Nakoman, 1966
R
Chlorophyceae
Botryococcus sp. cf.
B. braunii Kützing, 1849
–
Prasinophyceae
Pterospermella sp.
13P –
Tasmanites sp.
–
FUNGI ASCOMYCOTA
Microthyriacites sp.
13Q R P
Phragmothyrites sp.
R
Pluricellaesporites spp.
R
Parasite fungus with hyphopodia R
TABLE 1. — Continuation.
133
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
TABLE 2. — Botanical affi nities of all the taxa encountered during our investigation. Macrofl oral remains of only Lignophyta (Spermato-
phyta) were previously studied. Abbreviations: A, Ashikajima Formation; K, Kimigahama Formation; –, unknown.
BOTANICAL AFFINITY FOSSIL
Phylum Class Order Family Microfl ora
Bryophyta
Marchantiopsida – –
Aequitriradites spinulosus, Aequitriradites
verrucosus, Aequitriradites sp., Triporoletes
reticulatus
–––
Neoraistrickia sp. 1, Neoraistrickia sp. 2,
Neoraistrickia sp. 3
Lycophyta Lycopsida
Lycopodiales Lycopodiaceae
Foveosporites ryosekiensis n. sp.,
Lycopodiumsporites crassimacerius,
Lycopodiumsporites dentimuratus, Retitriletes
austroclavatidites
Selaginellales Selaginellaceae
Cingutriletes sp., Echinatisporis varispinosus,
Neoraistrickia sp. 1, Neoraistrickia sp. 2,
Neoraistrickia sp. 3, Uvaesporites sp.
––
Retitriletes sp.
Monilophyta
Equisetopsida
(= Sphenopsida)
Noeggerathiales –
Microreticulatisporites sp. 1,
Microreticulatisporites sp. 2
Polypodiopsida
(= Filicopsida)
Cyatheales
Cyatheaceae or
Dicksoniaceae
Cibotiumspora paradoxa,
Concavissimisporites punctatus,
Concavissimisporites variverrucatus,
Cyathidites australis, Cyathidites minor,
Cyathidites rarus, Deltoidospora hallii,
Uvaesporites sp.
?
Converrucosisporites sp., Verrucosisporites
densus
Gleicheniales Matoniaceae
Cyathidites rarus, Matonisporites sp. 1,
Matonisporites sp. 2
Osmundales Osmundaceae
Baculatisporites comaumensis,
Baculatisporites sp., Biretisporites potoniaei,
Osmundacidites wellmanii, Todisporites major,
Todisporites miser
Polypodiales
Dennstaedtiaceae
Biretisporites potoniaei, Laevigatosporites sp.,
Microreticulatisporites sp. 1,
Microreticulatisporites sp. 2
Polypodiaceae
Baculatisporites comaumensis,
Laevigatosporites sp.
Pteridaceae
Contignisporites burgeri, Contignisporites
cooksonii, Laevigatosporites sp., Manumia
japonica n. sp.
Saccolomataceae
Cicatricosisporites sinuosus
Schizaeales
Anemiaceae
Cicatricosisporites annulatus,
Cicatricosisporites hughesi, Cicatricosisporites
mohrioides, Cicatricosisporites sp. 1,
Cicatricosisporites sp. 2, Cicatricosisporites
sp. 3, Cicatricosisporites sp. 4,
Contignisporites cooksonii,
Distaltriangulisporites sp., Nodosisporites
choshiensis n. sp., Nodosisporites makotoi
n. sp., Ruffordiaspora australiensis
Lygodiaceae
Concavissimisporites punctatus
Schizaeaceae
Klukisporites variegatus, Plicatella sp.
134
GEODIVERSITAS • 2011 • 33 (1)
Legrand J. et al.
BOTANICAL AFFINITY FOSSIL
Phylum Class Order Family Microfl ora
Monilophyta
Polypodiopsida
(= Filicopsida)
––
Cingulatisporites psilatus, C. sp. 1, C. sp. 2,
Cingutriletes sp., Coronatispora valdensis,
Incertae sedis in Hasenboehler (1981),
Laevigatosporites ovatus, Patellasporites
sp., Pilosisporites brevis, P. verus,
Polycingulatisporites reduncus, Trilites
bossus, Tuberositriletes sp., Undulatisporites
undulapolus, U. sinuosis
–––
Incertae sedis sp. 1
BOTANICAL AFFINITY FOSSIL
Phylum Class Order Family Microfl ora Macrofl ora
Lignophyta (Spermatophyta)
Gymnosperms
Bennettitales, Cycadales or
Pentoxylales
–
Cycadopites minimus,
Cycadopites sp.
Bennettitales:
Ptilophyllum acinacifolium Kimura &
Okubo (K),
P. choshiense Kimura, Okubo &
Miyahashi (K),
P. elongatum Kimura & Ohana (non
Douglas) emend. Kimura, Okubo &
Miyahashi (K),
P. subulatum Kimura & Okubo (K),
P. sp. A in Kimura (1997) (K),
P. sp. B in Kimura (1997) (K),
Zamites choshiensis Kimura & Ohana
(K)
Cycadales: Nilssonia dictyophylla
Kimura & Okubo (K)
Caytoniales –
Sagenopteris inequilateralis Oishi (K)
Erdtmani-
thecales
Erdtmanithecaceae
Eucommiidites minor,
Eucommiidites troedssonii
Gnetales
Ephedraceae
Ephedripites montanaensis
Gnetaceae
Gnetaceaepollenites sp.
–
Classopollis torosus,
Corollina sp.
Coniferales
Araucariaceae
and/or
Podocarpaceae
Araucariacites australis,
Balmeiopsis limbatus,
Callialasporites dampieri,
Callialasporites sp. cf. C. ugensis
Cheirolepidiaceae
or Voltziaceae
Classopollis torosus,
Corollina sp.
Cheirolepidiaceae: Cupressinocladus
obatae Okubo & Kimura (K),
Frenelopsis choshiensis Kimura,
Saiki & Arai (K)
Cupressaceae
or Taxaceae
Exesipollenites tumulus,
Spheripollenites psilatus
Pinaceae
Cerebropollenites mesozoicus,
Pityosporites sp. cf. P. piniformis
Sciadopityaceae
Cerebropollenites mesozoicus
–
Inaperturopollenites sp.,
Pityosporites sp. cf.
P. constrictus
Pterido-
spermales
–
Alisporites thomasii
Stenopteris cyclostoma Saiki,
Kimura & Horiuchi (A)
––
Alisporites sp. cf. A. australis,
A. sp., Incertae sedis sp. 2
T
ABLE 2. — Continuation.
135
Barremian palynofl oras from south-west Japan
GEODIVERSITAS • 2011 • 33 (1)
BOTANICAL AFFINITY FOSSIL
Phylum Class Order Family Microfl ora
Algae
Chlorophyta
Charophyceae Zygnematales Zygnemataceae
Chomotriletes minor, Ovoidites parvus
Chlorophyceae Chlorococcales Botryococcaceae
Botryococcus sp. cf. B. braunii
Prasinophyceae Pterospermatales
Pterospermellaceae
Pterospermella sp.
Tasmanitaceae
Fungi
Ascomycota
Dothideomycetes
Asterinales Asterinaceae Parasite fungus with hyphopodia
Microthyriales –
Microthyriacites sp., Phragmothyrites sp.
–––
Pluricellaesporites spp.
TABLE 2. — Continuation.
