ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No.

172

Palaeontological Society of Japan
December 30, 1993

Co-Editors

Tsunemasa Saito and Kei Mori

Language Editor

Martin Janal (New York)

Editorial Board
Shiro Hasegawa (Hokkaido University), Kunihiro Ishizaki (Tohoku University), Tomoki Kase
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Officers for 1993-1994
President: Hisayoshi Igo
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The fossil on the cover is Isocrinus (Chladocrinus) hanaii Oji, an Early Cretaceous(Aptian) crinoid, which was described from the Hiraiga Formation exposed at Haipe,
Tanohata-mura, Shimo-Hei County, Iwate Prefecture, Northeast Japan. (University
Museum of the University of Tokyo coIl. cat. no. ME6950, paratype specimen, length
about II em)

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 172, pp. 251-263,8 Figs., December 30, 1993

959.

PHYLOGENETIC SIGNIFICANCE OF A NEW
ARAPHID DIATOM SPECIES, DELPHINEISKAMENOOENSIS SP. NOV. *YUKIO Y ANAGISAWA
Geological Survey of Japan, 1-1-3 Higashi,
Tsukuba, Ibaraki, 305

Abstract. A new diatom species Delphineis kamenooensis is described from the Lower
Miocene Kamenoo Formation in the Joban Coalfield of northeast Honshu, Japan. The
species occurs in the upper part of the late Early Miocene Thalassiosira fraga Zone (NPD
2) to the lower part of the latest Early Miocene Crucidenlicula kanayae Zone (NPD 3A) of
the North Pacific diatom zonation and serves as a marker species for this interval. The new
species is characterized by linear valve outline, relatively sparse areolae and partiallydeveloped transverse grooves on valve face. It shows close morphologic resemblance toDelph in eis miocenica (Schrader) Andrews and is probably its direct precursor. D.
kamenooensis sp. nov. has features common to the Delphineis sllrirella, D. Iineala and D.
ovala subgeneric groups proposed by Andrews (1988) and may be an intermediate form
among the three groups.
Key words.

Diatom, fossil, marine, Delphineis, Miocene, Japan.

synthesized its phylogeny and outlined probable evolutionary development and relationships of its subgeneric groups within thegenus. He showed that three distinctive
affinity groups, the Delphineis surirella, D.
lineala and D. ovata groups, had diversified
before the Middle Miocene time and he postulated evolutionary sequences within eachgroup. However, the pre-Middle Miocene
evolutionary history of Delphineis as well as
the evolutionary diversification of the threesubgeneric groups is not well known at present.
In this study, a new Delphineis species, D.
kamenooensis, is described from the upper
Lower Miocene Kamenoo Formation In
Japan on the basis of light and scanningelectron microscopic observations. In addition, Delphineis miocenica (Schrader) Andrews, a species which is quite similar inmorphology to D. kamenooensis and has thesame stratigraphic range as D. kamenooensis,

In troduction

Delphineis is a marine diatom genus erectedby Andrews (1977) for several species formerly assigned to the genus Rhaphoneis Ehr.
The genus Delphineis was then composed ofonly five fossil species (Andrews, 1977), but
subseq uent descri ptions and generic transfersof many additional extant and fossil species
have made Delphineis a moderately large
genus consisting of at least 23 species atpresent (Fryxell and Miller, 1978; Andrews,
1981; Hendey, 1981; Sancetta, 1982;
Akiba, 1986; Prasad, 1986, 1992; Barron
and Baldauf, 1986; Andrews, 1988).
The morphology and phylogeny of Delphineis is fairly well understood mainly
through continuing research effort by Andrews (1977,1981,1988). Andrews (1988)'Received February 5. 1993; revised manuscript
accepted August 16, 1993
251

Ylikio Yanagisawa

252

was also examined in detail by the scanningelectron microscope for comparison. Delphineis kamenooensis has features common toboth the D. sllrirella, D. lineata and D. Ol'ata
subgeneric groups and may be an intermediate species of the three groups.
Materials
Samples studied in this study are as follows(Figures 1, 2) :1. Sample JOB 680; The type material ofDelphineis kamenooensis, laminated siliceous
mudstone of the Lower Miocene Kamenoo
Formation of the Yunagaya Group in the
Joban Coalfield (Sugai et al., 1957), collected

132

135

138

14f

46

44

42

Sea of Japan
40

36

34

Ocean 0

100

200Km

Figure 1. Map showing localities of samples
used in this study.

Figure 2. Map showing the locations of thesamples including Delphineis kamenooensis Yanagisawa sp. nov. and D. miocenica (Schrader) Andrews
in Kurosuno area, lzumi-machi, Iwaki City, Fukushima Prefecture (Topographic map "Iwaki-izumi",
1 : 25,000 scale, Geographical Survey Institute).
JOB 680: the type sample of D. kamenoonsis(Kamenoo Formation). JOB 1526: a sample including Delphineis l11iocenica (Schrader) Andrews
and D. kal11enooensis used for observation (Taira
Formation). Stars indicate the locations of samples
including D. kal11enooensis (Taira Formation).
Closed circles indicate the locations of samples including D. l11iocenica (Taira Formation).

from a small cliff near Kurosuno, lzumimachi, Iwaki City, Fukushima Prefecture (36'
55'54.5"N, 140'50' l4.0"E, Figure 2). The
diatom assemblage is correlative with theupper part of the Thalassiosira fraga Zone---+ Figure 3. LM photomicrographs. Scale bars equal l0.um (A for 1-14, 16 and B for 15). 1-11.
Delphineis kamenooensis Yanagisawa sp. nov., Sample JOB 680, Kamenoo Formation (3. Holotype). 12.
Delphineis kamenooensis Yanagisawa sp. nov., Sample Km 41A, Emi Formation. 13,14. Delphineis
kamenooensis Yanagisawa sp. nov. (slightly broad form), Sample JOB 680, Kamenoo Formation. 15.
Delphineis l11iocenica (Schrader) Andrews, Sample JOB 1526, Taira Fomation. 16. Delphineis miocenica(Schrader) Andrews, Sample Mzn 06, Oidawara Formation.

959.

New diatom species Delphineis kamenooensis

A

I
B

I

253

254

Yukio Yanagisawa(NPD 2) of Akiba (1986).
2. Sample Km 41A; A calcareous concretion taken from the Emi Formation of the
Awa Group distributed in the Kamogawa
area of Boso Peninsula (Nakajima et al.,1981). It was obtained from an outcrop in
Choja River, Wada Town, Awa County,
Chiba Prefecture (3Y04'9.1"N, 140'01'43.5"
E). The occurrence of Actinocyclus ingens
and Crucidenticula kanayae and the lack ofDenticulopsis species indicate that this sample
is correlative with the C. kanayae Zone
(NPD 3A) of Akiba (1986).
3. Sample JOB 1526; Massive mudstone
from the Honya Mudstone Member of the
Taira Formation of the Yunagaya Group in
the Joban Coalfield, sampled from a small
cliff north of Kurosuno, Izumi-machi, Iwaki
City, Fukushima Prefecture (36'56'7.6"N,
140'50' 13.3"E, Figure 2). This sample isincluded in the upper part of Thalassiosira
fraga Zone (NPD 2).
4. Sample Mzn 06; Diatomaceous mudstone from the Oidawara Formation of
Mizunami area, central Japan. It was sampled from a road-cut cliff near Shukunohora,
Hiyoshi-machi, Mizunami City, Gifu Prefecture (35'24'31.22"N, 13rJ6'17.8"E). The
diatom flora of the sample is assigned to thelower part of the Middle Miocene Denticulopsis lauta Zone (NPD 4A) of Akiba (1986).
5. DSDP Leg 57 Holes 438A and 438B
(off northeastern Honshu); These holes were
examined to check the stratigraphic ranges ofDelphineis kamenooensis and D. miocenica.
However, no specimens of D. kamenooensis
were found.

Methods
For each sample, approximately I g of eachdried sediment was placed in a 100 ml beaker.
The sediment was disintegrated in hydrogen
peroxide (15%) and boiled for about 15 minutes, and 20 ml of hydrochloric acid was thenadded and the sediment was boiled for about
10 minutes. The acid was removed by atleast five successive 3-hour decantations with
distilled water. 0.01 N sodium pyrophosphate(Na Z P2 0 7 ) was then added to the liquid andsuspended clays were removed by 3-hourdecantations. This procedure was repeatedat least five times and then sodium pyrophosphate was removed by at least five successive
3-hour decantations with distilled water.
A strewn slide for light microscope (LM)observation was prepared for each sample onan 18 X 18 mm cover glass and the cover glass
was mounted in Pleurax on a glass slide.
For scanning electron microscope (SEM)
observation, diatom valves were picked fromdried samples and mounted on sample stages,
using one hair of a tiny paint brush. SEM
observation was performed on D. kamenooensis and D. miocenica using JOEL T330A of
Geological Survey of Japan.
Taxonomy
In this paper, the classification system ofRound et al. (1990) is adopted for description.

~
Figure 4. SEM photomicrographs of Delphineis kamenooensis Yanagisawa sp. nov. Narrower andbroader scale bars equal 5 Jim and I Jim, respectively. All from sample JOB 680 (Kamenoo Formation). 1.
Outer view of relatively long valve. 2, 3. Outer view of valve apex. Arrows indicate two apical pores andarrow heads indicate external openings of rimoportulae. Note stubby spine-like projections at the shoulderof the valve. 4. Side view of valve. Sa, Sb. Inner view of valve with girdle bands (Stereoscopic pair). 6.
Enlargement of Figure 5a showing valve apex and a lip-shaped internal opening ofrimoportula. 7. Obliqueouter view of valve with a girdle band. 8. Outer view of relatively short valve. 9. Oblique inner view ofvalve with a girdle band. 10. Areolae occluded with rota-type cribra.

959.

New diatom species Delphineis kamenooensis

255

Yukio Yanagisawa

256

Class Fragilariophyceae Round, 1990
Subclass Fragilariophycidae Round, 1990
Family Rhaphoneidales Round, 1990
Subfamily Rhaphoneidaceae Forti, 1912
Genus Delphineis Andrews, 1977,
emend. Andrews, 1981

Delphineis kamenooensis Yanagisawa
sp. nov.
Figures 3-1-14; 4-1-10.

Description.-Valve linear or linear-elliptical to lanceolate, with slightly acute apices,
9-49 pm long, 7-10 pm wide (Figure S).
Valve face flat, punctated by transverse rowsof circular areolae. Cribra of rota-type on
the outer surface of valve and connected to
the valve by two longitudinally oriented
struts (Fig. 4-10). Transverse rows ofareolae fairly sparse, 8-IO.S in 10 pm, parallel
to transapical axis at the center, and becoming radiate near the apices, finer striae curve
completely around the ends of the valve.
Marginal areolae in a row along the edge ofvalve. The other areolae on the valve face,surrounded by slightly raised rims, arranged
secondarily in widely spaced longitudinalrows, usually two rows, but four or six rows

in broader specimens. Some areolae areoccluded with thin siliceous layer on theoutside of the valve and so are invisible on
the external surface of valve. Mantle shallow with a row of areolae of the same size as
valve face areolae. Hyaline axial area indistinct, because of the widely spaced longitudinal rows of areolae. External grooves partially developed, surrounded by slightly
raised rims containing the marginal areolae
and the mantle areolae, but not reaching to
the valve face areolae. Raised areas between
the grooves terminate in stubby spine-like
attachment projections with two heads along
the shoulder of the valve. At each apex, twosmall pores penetrate into the valve interior,and a single rimoportula is located slightly
off-center, occupying the position where oneof the marginal areolae might be positioned.
The two rimoportulae are diagonally placedwith respect to the longitudinal axis. External opening of the rimoportula forms a small
elongated pore with a slightly raised rim.
Internal opening of the rimoportula shows alip-shaped projection. Girdle bands split
and plain.
Holotype.-Figure 3-3 (GSJ FI4S83 deposited in the Geological Survey of Japan).

Type
50

•
•

40

,2,
.c
0;,
c

.3 20
10

•

•
•
• •

•

•

0
6

7

and

locality.-JOB

680

0

•• •
•
•
•• •
• •
I
Ie •
•
• • •
•
••
••

E 30

sample(Kamenoo Formation, Yunagaya Group),
Kurosuno, Iwaki-izumi-machi, Iwaki City,
Fukushima Prefecture (36°SS'S4.S"N, 140 S0'
14.0"E, Figure 2).
Remarks.-Delphineis kamenooensis canundoubtedly be placed in the genus Delphineis on the basis of its general morphologic characteristics such as the presenceof two small pores at the valve apices, rotatype cribra, the number and position ofrimoportulae and the arrangement of areolae,
which agree well with its generic definition by
Andrews (1977, 1981).
This new species resembles Delphineis
miocenica (Schrader) Andrews in having relatively sparse areolae and almost the same sizeand spacing of areolae. Relatively broad
forms of D. kamenooensis with lanceolate

89Width (J.1m)

10II

Figure 5. Length and width plot of Delphineis
kamenooensis Yanagisawa, sp. nov. (Sample JOB
680, Kamenoo Formation).

959.

New diatom species Delphineis kamenooensis

North PacificDiatom zones
Age(Akiba, 1986).---- Stratigraphic
ranges257

Stratigraphic sections

Q

Q".

Ii!":

M.
I

c. nicubu'ILU ISA
Denticulopsis 4B
hyalina

~

c

~

~

.~

E:

v

~

0

.S:
IS

~
15

~

Denticulopsis
4A
lauta

~

'C

~

""

E:

~

~

E:

Denfir1Jlnnoic
n~n,;i/H:fn

~

.5
.=:
,s.

Mhlln'lmi~'Oidawara F.'

~

~

3B

IS

Emi
Fonnation

JJ

.::J
~

Crucidenticula 3A
kanayaeTakakuG.

Shirado

G.

.5
.=:

~
~

~

I,

G.

.::J
.::J
~

I

Taga

I

I

~

~

C

~

v

0

~

"~ Kamenoo F.

0

L:
<'\I

F.

rf Mizunoya F.

....

f;I;l

Taira

:::I

Thalassiosira
jraga

>-

GoyasuNabutoFonnation

F.

2

Iwaki
(Joban)•of

DSDP Holes
438A,4388
Kamogawa

Figure 6. Stratigraphic occurrences of Delphineis kamenooensis Yanagisawa sp. nov. and D.
miocenica (Schrader) Andrews with biochronological distribution of the sections studied in this paper.

outline and four or six longitudinal rows ofareolae on the valve face bear very close
resemblance to D. miocenica (compare Figures3-13, 14 with Figures 3-15, 16). However, D. kamenooensis differs from D.
miocenica in having a more linear valve
outline, and by the presence of externalgrooves and the prominent stub-like projections on the valve shoulder.
Delphineis kamenooensis is also similar to
Delphineis ovata Andrews (1977) and D.
penelliptica Andrews (1977) in valve outline,

but differs from them in having slightly acuteapices, sparsely spaced areolae and indistinct
axial area. The transapical striae of D.
kamenooensis are finer than those of the lattertwo species; D. kamenooensis has 8.0-10.5
striae in 10,um, whereas D. penelliptica andD. ovata have 7 and 5.5-7 striae in 10,um,
respectively. D. kamenooensis shows affinity
to Delphineis lineata Andrews (1977), but isdistinguished by slightly acute apices, theabsence of distinct axial area and more sparsely spaced longitudinal rows of areolae.

258

Yukio Yanagisawa

Delphineis kamenooensis is similar to Delphineis parallelica (Schrader) Andrews 1988(= Rhaphoneis parallelica Schrader in
Schrader and Fenner, 1976) in having a linear
valve outline with slightly acute apices, butdiffers from it by the absence of a clear axial
area.
Stratigraphic occurrence.-Stratigraphic
records in the Joban Coalfield and Kamogawa area indicate that the stratigraphic
range of D. kamenooensis is at least from theupper part of the Thalassiosira fraga Zone
(NPD 2) to the lowermost part of the
Crucidenticula kanayae Zone (NPD 3A)(Figure 6).
The species has been found from several
samples of the Kamenoo and Taira Formations in the Joban Coalfield (See Figure 2)
and also from a calcareous concretion of theEmi Formation distributed in the Kamogawa
area of the Boso Peninsula.
The diatom assemblages of the Kamenoo
and Taira Formations are assigned to theupper part of the T. fraga Zone of Akiba(1986) or the Kisseleviella carina Zone of
Koizumi (1985) on the basis of the absence ofActinocyclus ingens with the abundant occurrence of Kisseleviella carina (Koizumi, 1986,
Yanagisawa et al., 1989; Taketani et aI.,
1990; this study). The first occurrence of D.
kamenooensis therefore ranges down at least
to the T. fraga Zone. The calcareous concretion from the Emi Formation includes a
diatom flora correlative with the lowermost
part of the C. kanayae Zone. Since D.
kamenooensis has not been found in the mainpart of the C. kanayae Zone as well as inmuch younger diatom zones in DSDP Holes

438A and 438B, and also in Joban Coalfield
and Kamogawa area, its last occurrence maylie in the lower part of the C. kanayae Zone.

Delphineis miocenica (Schrader)Andrews 1988
Figures 3-15, 16; 7-1-8
Andrews, 1988, p. 201-202, pI. I, fig. 2.
Basionym: Rhaphoneis miocenica Schrader, 1973, p.
709, pI. 25, figs. I, II
Synonymy: Rhaphoneis amphiceros Ehrenberg sensu
Hanna, 1932, p. 211-212, pI. 15, figs. 3-5; Rhaphoneis miocenica Schrader, Akiba et at., 1982,
pI. 3, fig. 85; Koizumi, 1986, pI. I, figs. 8, 9;
Yanagisawa et at., 1989, pI. 5, fig. 48.

Observation.- Valve lanceolate with slightly acute apices. Valve face flat, punctated byrows of circular areolae. The areolae surrounded by slightly raised rims and occluded
by rotae. Relatively few transverse rows ofareolae parallel to the transapical axis at thecenter, and becoming radiate near the apices.
Marginal areolae in a row along the edge ofvalve. The other areolae arranged secondarily in a few widely spaced longitudinal rows.
Some areolae are occluded with thin siliceous
layer on the outside of the valve and so areinvisible on the external surface of valve.
Mantle shallow with a row of areolae of thesame size as the valve face areolae. Hyaline
axial area indistinct because of the widely
spaced longitudinal rows of areolae. External grooves absent. At the shoulder of valve,
marginal ridge present with projections, but
the projections are unclear. At each apex,
two small pores present, penetrating into thevalve interior, and a single rimoportula locat-~
Figure 7. SEM photomicrographs of Detphineis miocenica (Schrader) Andrews. Narrower andbroader scale bars equal 5 fJ.m and I fJ.m, respectively. 1-4, 6 and 7 from sample Mzn 06 (Oidawara
Formation, Mizumami area) and 5 and 8 from sample JOB 1526 (Honya Member of the Taira Formation,
Kurosuno, Izumi-machi, Iwaki City). 1. Oblique outer view of valve. 2. Outer view of valve apex. Noteunclear spine-like projections at the shoulder of the valve. 3. Inner view of valve apex showing lip-shaped
internal opening ofrimoportula. 4. Outer view offrustule with some girdle bands. 5. Inner view of valve.
Arrow heads indicate internal openings of rimoportulae. 6. Oblique outer view of valve. 7. Outer viewof valve apex. Arrow head indicates external opening ofrimoportula. 8. Inner view of valve apex showing
lip-shaped internal opening of rimoportula (arrow head).

959.

New dia tom species Deiphin eis kamenooensis

259

260

Yukio Yanagisawa

ed slightly off-center, occupying the position
where one of the marginal areolae might bepositioned. The two rimoportulae are diagonally placed with respect to the longitudinalaxis. External opening of the rimoportula
forming a small elongated pore with a slightly
raised rim. Internal opening of the rimoportula has a lip-shaped projection.
Remarks.- The observational results of D.
miocenica of this study agree fairly well withthose by Andrews (1988) except for the presence of the stubby spine-like projections at
the shoulder of the valve. He reported thelack of the projections, but my specimensfrom the Oidawara Formation possess theseprojections although they are very short andunclear. The absence of the projections inAndrews's (1988) specimens is probably due
to very poor preservation of his sample as hesuggested.
Stratigraphic occurrence.-In the northwest Pacific, this species occurs from the Early
Miocene Thalassiosira /raga Zone (NPD 2)
to the upper part of the early Middle Miocene
Denticulopsis hyalina Zone (NPD 4B) of
Akiba (1986) (Figure 6).
In the Joban Coalfied, this species occurs inthe Honya Mudstone Member of the Taira
Formation and the overlying Shirado, Takaku and Taga Groups (Koizumi, 1986; Taketani et al., 1990), but it has not been encountered in the underlying Kamenoo Formation. Therefore, the first occurrence of D.
miocenica might lie near the boundary
between the Kamenoo and Taira Formations(Figure 6). Since the two formations are
assigned to the upper part of the T. fraga
Zone (NPD 2), the first occurrence of D.
miocenica falls within this zone.
At DSDP Holes 438A and 438B, D.
miocenica first occurs from the lower part of
the Crucidenticula kanayae Zone (NPD 3A).
However, the true first occurrence of thisspecies can not be detected because the lowermost part of the C. kanayae Zone and mostof the T. fraga Zone are represented byglauconitic sandstone containng no diatoms(Figure 6). The last occurrence of D.
miocenica is recognized in the upper part of
the D. hyalin a Zone (NPD 4B) at Hole 438A.

Discussion
Among species of the genus Delph in eis, D.
kamenooensis is most similar to D. miocenica.
The two species share many common features
such as the sparsely distributed areolae on thevalve face, the medium-size areolae, the secondarily arranged longitudinal rows of areolae
and the relatively narrow indistinct axialarea, suggesting a very intimate phylogenetic
relationship between the two. Furthermore,
the relatively broad form of D. kamenooensis
bears a very close appearance to D. miocenica.
Stratigraphic data indicate that the first
appearance of D. kamenooensis predates thatof D. miocenica (Figure 6). These stratigraphic ranges along with the morphologic
resemblance strongly suggest that D.
kamenooensis is probably the direct ancestorof D. miocenica. Delphineis kamenooensis
might evolve to D. miocenica with the change
in valve outline from linear to lanceolate
shape, the disappearance of external grooves,and the reduction of the stubby spine-likeprojections at the shoulder of the valve to lessprominent forms.
On the basis of some important characteristics, Andrews (1988) recognized three distinct
subgeneric groups within the genus Delphineis: the Delphineis lineata, D. ovata andD. surirella groups (Table I). The D. lineataand D. ovata groups are both characterized
by the presence of external grooves on thevalve face and the prominent stubby spinelike projections at the shoulder of the valve.
The two groups are distinguished mainly by
areolae size and spacing; the D. lineatagroup has more compactly spaced and finer
areolae than the D. ovata group. On thecontrary, the D. surirella group is clearly
distinguished from the other two groups by
the lack of such grooves and prominent
spine-like projections at the shoulder of the959.

New diatom species Delphineis kal11enooensis

261

Table 1. Comparison of characters among D. miocenica, D. kamenooensis andthe three subgeneric groups in Delphineis.
Subgeneric groups
D. miocenica

D. kamenooensis

AbsentAbsentPresent panially

AbsentPresent(unclear)

Present

Narrow or broad

Narrow

Narrow

CoarseCoarseCoarseCharacters
D.ovata

D. lineata

D. surirella

External grooves

Present

Present

Spine-like projections

Present

Present

Axial area

Narrow

Broad

Areolae

CoarseFinevalve. Although he suggested the morphologic uniqueness of D. miocenica within
the gen us Delphineis, Andrews (1988) tentatively assigned D. l11iocenica to the D. surirelfa group because of the absence of externalgrooves and the lack of strong spine-likeprojections.
As suggested above, the new Delphineis
species herei n descri bed as D. kal11enooensis
is probably the ancestor of D. miocenica(Figure 8). However, D. kamenooensis can-Paleogene

Early Miocene Middle Miocene Late Miocen

bfDb
r
CD.
. . :f.TD.

not be placed in the D. surirella group as D.
l11iocenica can be, because it has the valve facegrooves and the distinct spine-like projections. It can be rather said that D.
kamenooensis is much closer to the D. lineataand D. ovata groups because of the presenceof external grooves and the prominent spinelike projections, but the areolae spacing andindistinct axial area as well as the partial
development of grooves make me hesitate toplace this species in either of these groups.

Pliocene

Pleistocene

Holocene

Groups. lsenata.1

Delphineis
ovata
Group

D. angustata

D. ~enelliptica

D.ovatal

!

';
i

I

? :

Proto·
Delphineis'............

?:

nomeco,,","e"D. karstenii...
,?
....... 1' ............................ •.•••••••••••••.••••••••

D.lineata

Delphineis
lineataGroup!amenoo,"!"~f--ID. miocenica
D. surirella.....

Delphineis
surirellaGroup

D. surirelloides
Figure 8.

Evolutionary sequence in the genus Delphineis modified after Andrews (1988).

Yukio Yanagisawa

262

Thus D. kamenooensis cannot be assignedwith confidence to any of the three subgenericgroups and appears to be phylogenetically anintermediate form between the three groups.
D. kamenooensis may possibly be the common ancestor of the three subgeneric groups.
Much stratigraphic and taxonomic study,
however, needs to be carried out to confirm
such an evolutionary relationship, making
use of more continuous and better preserved
stratigraphic sequences.
Acknowledgements

Special thanks are given to Fumio Akiba(JAPEX Research Center of Japan Petroleum
Exploration Co. Ltd.), George W. Andrews,
John A. Barron (U.S. Geological Survey),
Elisabeth Fourtanier (California Academy of
Sciences) and Itaru Koizumi (Hokkaido
University) for their constructive review ofthe manuscript and helpful suggestions on
diatom taxonomy. I wish to thank Fumio
Yoshida (Geological Survey of Japan) andKonosuke Sawamura for providing samples
used in this study.

References
Akiba, F., 1986: Middle Miocene to Quaternary
diatom biostratigraphy in the Nankai Troughand Japan Trench, and modified Lower Miocene
through Quaternary diatom zones for middle-tohigh latitudes of the North Pacific. In, Kagami,
H., Karig, D.E. and Coulbourn, W.T. et al., In it.
Rep. DSDP, vol. 87, p. 393-481, pis. 1-30. U.S.
Govt. Printing Office, Washington, D.C.
- - , Yanagisawa, Y. and Ishii, T., 1982: Neogene
diatom biostratigraphy of the Matsushima area
and its environs, Miyagi Prefecture, Northeast
Japan. Bull. Geol. Surv. Japan, vol. 33, p. 215239. (in Japanese with English abstract)
Andrews, G.W., 1977: Morphology and stratigraphic
significance of Delphineis, a new marine diatom
genus. Beih. zur Nova Hedwigia, no. 54, p. 243260, pis. 1-5.
--,1981: Reversion of the diatom genus Delphineis
and morphology of Delphineis surirella (Ehrenberg) G.W. Andrews, n. comb. In, Ross R. ed.
Proc. Sixth Symposium on Recent and Fossil

Diatoms, p.81-92, pis. 1-2. Otto Koeltz
Scientific Publishers, Koenigstein.
- - , 1988: Evolutionary trends in the marine
diatom genus Delphineis G.W. Andrews. In,
Round, F.E. ed., Proc. Ninth International
Diatom Symposium, p. 197-206, pI. I. Biopress
Ltd., Bristol and Koeltz Scientific Books, Koenigstein.
Barron, J.A. and Baldauf, J.G., 1986: Diatom stratigraphy of the lower Pliocene part of the Sisquoc
Formation, Harris Grade section, California.
Micropaleontology, vol. 32, p. 357-371.
Fryxell, G.A. and Miller, W. I. III, 1978: Chainforming diatoms: three araphid species. Bacillaria, vol. I, p. 113-136, pis. 1-6.
Hanna, G.D., 1932: The diatoms of Sharktooth Hill,
Kern County, California. Proc. California
Acad. Sci., Fourth Ser., vol. 20, p. 161-263, pis.
2-18.
Hendey, N.I., 1981: Miocene diatoms from thesubantarctic southwest Pacific, Deep Sea Drilling
Project Leg 29, Site 278, Core 10. Bacillaria,
vol. 4, p. 65-124, pis. 1-4.
Koizumi, I., 1985: Diatom biochronology for the lateCenozoic Northwest Pacific. Jour. Geol. Soc.
Japan, vol. 91, p. 195-211.
- - , 1986: Diatom biochronology of the Neogene in
the Joban Coal-field, Northeast Japan- Yunagaya, Shirado, and Takaku Groups-.
In, Nakagawa, H., Kotaka, T. and Takayanagi,
Y. eds., Essays in Geology (Kitamura Commem.
Vol.), p.175-191, pis. 1,2. (in Japanese withEnglish abstract)
Nakajima, T., Makimoto, H., Hirayama, J. and Tokuhashi, S., 1981: Geology of the Kamogawa
district (Quadrangle Series, Scale I : 50,000), 107
p. Geological Survey of Japan. (in Japanese withEnglish abstract)
Prasad, A.K.S.K., 1986: Delphineis livingstonii n. sp.
(Diatomaceae, Bacillariophyceae) from St. George Sound, northeastern Gulf of Mexico.
Botanica Marina, vol. 29, p.517-522.
- - , 1992: Type material of the Miocene diatom
Dimeregramma brandywinensis Patrick and its
transfer to the genus Delphineis Andrews.
Micropaleontology., vol. 38, p.418-420.
Round, F.E., Crawford., R.M. and Mann, D.G., 1990:
The diatoms. 747p. Cambridge Univ. Press,
Cambridge.
Sancetta, c., 1982: Distribution of diatom species insurface sediments of the Bering and Okhotsk
Seas. Micropaleontology., vol. 28, p.221-227,
pIs. 1-6.
Schrader, H.J., 1973: Cenozoic diatoms from thenortheast Pacific, Leg 18. In, Kulm, L. D., von
Huene, R. et al., In it, Rep. DSDP, vol. 18, p. 673797, U.S. Govt. Printing Office, Washington, D.

959.

New diatom species Delphineis kamenooensisc.
- - and Fenner, J., 1976: Norwegian Sea Cenozoic
diatom biostratigraphy and taxonomy. In,
Talwani, M., Udintsev, G., et al., lnit. Rep.
DSDP, vol. 38, p. 921-1099, U.S. Govt. Printing
Office, Washington, D.C.
Sugai, K., Matsui, H., Sato, S., Kitagawa, Y., Sasaki,M., Miyashita, M. and Kawachi, H., 1957:
Explanatory text of the Joban Coal Field, Geological Maps of the Coal Fields of Japan, no.
I, 143p. Geological Survey of Japan. (inJapanese with English abstract)
Taketani, Y., Aita, Y., Ono, T., Okada, H., Hasegawa,

263

S., Maruyama, T., Nemoto, N., Kurihara, S. and
Takayanagi, Y., 1990: Geologic age and depositional environments of the Neogene sequence
distributed in Joban area, Fukushima Prefecture,
Japan. Res. Rep., Fukushima Mus., no. 20, p.
1-99, pis. 1-14. (in Japanese; title translated)
Yanagisawa, Y., Nakamura, K., Suzuki, Y.,
Sawamura, K., Yoshida, F., Tanaka, Y., Honda,
Y., and Tanahashi, M., 1989: Tertiary biostratigraphy and subsurface geology of the Futaba
district, Joban Coalfield, northeast Japan. Bull.
Geol. Surv. Japan, vol. 40, p.405-467, pis. 1-12.
(in Japanese with English abstract)

Kamenoo 1\ /~, Honya *:fi-, Taira.:>jZ, Yunagaya ~*:fi-, Joban 'iit~, Kurosuno m..~If, lzumi-machi +RIBJ, Iwaki ~'h~, Fukushima m~, Emi lIJi!., Awa ~m, Kamogawa"~JII, Boso mf(€., Choja *~, Wada 5flJEIl, Chiba +'l, Oidawara ~~, Mizunami JWijr~,
Shukunohora 1i§11li.I, Hiyoshi-machi B E"IBJ, Gifu U:J1'!., Honshu *,'1'1.

1!!t*lilj'fliJJO) 1 ~~ Delphineis kamenooensis O)**1t~a'iJg~: {~1:ifHI(7) 1 ~:fl Delphineis kamenooensis Yanagisawa sp. nov. z-'iit~hREIl t.:.5j-m-t;o r'€fj)op~*1CI7)1\ / ~Ji ii,
i? ~jfjjI; tA.:. *:flU:, Akiba (1986) t.:. J: ;0 ~t:;t.:>jZi$fE~{~:O*IK5j-(7) Thalassiosira fraga
Zone (NPD 2) (7) J:'€fj) (iltrJtJlOP~i!!:~JtJl) ii'i? Crucidenticula kanayae Zone (iltrJtJlOP~i!!:t.:.ii,t-t-citl±l L, ':'(7)IKFa1I7)f~~:fl C t;.;o. :J't$lJ:~£l't:&.lf~1!tmllJ:Vl¥dfH':'J: ;Ol!l~H':'
J:h~ *:fl~, .Mz-Lk~~ ~.~f~i?~~~ ~J:lf~ • • OO(7).KJ:~-CM
ji--::it-t i?h;o. *:flli, Delphineis miocenica (Schrader) Andrews t.:.~~~t.:. J: <~{J;t L,
JiFf~it;{j(ii'i?, D. miocenica (7)t.Il7t:flc~;ti?h;o. D. kamenooensis Ii, Delphineis ~*)(7)OPKJi!.i?h;o3~17)~~:fl,~-~c~h~hc~D-t;o~.z-M~, **1C~K~':'hi? 3 , Iv - 7.(7) OPFa1 t.:.fftt!:-t ;0.
jljpiR.

*7<::

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 172, pp. 264-276,6 Figs., December 30, 1993

960.

TWO NEOZAMITES SPECIES (BENNETTITALES) FROM THE
LOWER CRETACEOUS OF NORTHEAST CHINA AND
THE INNER ZONE OF JAPAN*
GE SUN
Nanjing Institute of Geology and Palaeontology, Academia
Sinica, Chi-Ming-Ssu, Nanjing, P.R. China

TAKASHI NAKAZAWA
Yoyogi Seminar, 16-27-1 Yoyogi, Shibuya-ku, Tokyo, 151
and

TAMIKO OHANA and TATSUAKI KIMURA
Institute of Natural History, 24-14-3 Takada,Toshima-ku, Tokyo, 171

Abstract. The genus Neozamiles was established on the basis of bennett ita lean pinnate
leaves bearing pinnae with markedly toothed margins. This genus is restricted in geographical distribution to Eastern Siberia, Northeast China, Inner Zone of Japan and Southern
Primorye, and in stratigraphical distribution to the Lower Cretaceous. This paper deals
with the description of Neozamiles elongalus in external and cuticular features on the basisof newly collected leaves from the Hegan Coal-Field, Heilongjiang, Northeast China, and of
N. inlermedius, sp. nov. in external features on the basis of leaves from the Tetori Basin in
the I nner Zone of Japan.
Key words.
type flora.

Neozamiles (Bennettitales), Lower Cretaceous, Hcgan Coal-Field, Tetori-

Neozamites is characterized by its pinnate
leaves [N. lebedevii is known to be bipinnate
according to Zheng and Zhang (1983)J withmarkedly toothed pinna margins and withbennettitalean cuticle.
Until now four species have been recorded.
They are Neozamites verchojanensis Vachrameev, N. lebedevii Vachrameev, N.
denticulatus (Kryshtofovich et Prynada) Vachrameev and N. elongatus Kimura et Sekido.
In this paper, we describe two Neozamites
species: N. elongatus Kimura et Sekido withpreserved cuticle (by Sun, Ohana and Kimura) and N. intermedius, sp. nov. without

Introduction
The bennettitalean genus Neozamiles wasestablished in 1962 by Vakhrameev [spelled
also as Vachrameev in the taxonomic descri ption J. He described the type species Neozamites verchojanensis from the Lower
Cretaceous Batylykh Formation distributed
along the Liampeska River (Figure I-II),
Yak utia, and N. lebedevii from the Lower
Cretaceous Khatylykh Formation distributed
along the Vilui River (Figure 1-15), Yakutia.
"Received June 14, 1993; accepted August 16, 1993
264

960.

Neozamites species from China and Japan

preserved cuticle (by Nakazawa, Ohana andKimura).
Neozamites elongatus was first described by
Kimura and Sekido (1971) on the basis of
pinnate leaves without preserved cuticle fromthe Oguchi Formation in the Inner Zone ofJapan (Figure 1-21 and Figure 2-21). The
present leaves referable to Neozamites elongatus were collected in 1990 from the Lower
Cretaceous Shitouhe Formation (an equivalent to the Chengzihe Formation of nonmarine origin in the Jixi Coal-Field, southeastern Heilongjiang) in the Hegan CoalField (Figure 1-1), northeastern Heilongjiang.
The leaves of Neozamites intermedius, sp.
nov. were collected by one of us (Nakazawa)in 1982 from the fossiliferous beds of the
Akaiwa Formation equivalent of nonmarine
origin (possibly Aptian in age) (Figure 1-22
and Figure 2-22).
The genus Neozamites is restricted in stratigraphical and geographical distributions to
the Lower Cretaceous plant beds in Eastern
Siberia, most of Northeast China and theInner Zone of Japan (Figures 1 and 2).
These regions are characterized phytogeographically by the Siberian and Tetori-typefloras (e.g. Kimura, 1987a, b). In addition,
exceptional occurrence of Neozamites isknown in the Early Cretaceous flora of themixed type from Southern Primorye (Barremian Ussuri and Starosutchan, and Albian
Galenka Formations) (Figure 1-17-20).
It is highly probable that Neozamites did
make its southward migration since the Barremian time together with such genera
characterizing the Siberian and Tetori-typefloras as Ctenis, Ginkgo (including Baiera)
and Podozamites (or Podozamites-like plants)
to Southern Primorye where the Ryoseki-type
plants had extensively flourished (e.g. Kimura, 1987a, b).
The present Neozamites elongatus has wellpreserved cuticle. The specimens were firsttreated with HF (40%), then macerated with
Schulze's solution and followed by diluted

265

KOH. The preparations were observed by
the phase contrast microscope and SEM.
It would be worth mentioning that several
bennettitalean leaves (e.g. Neozamites) andcycadalean leaves (e.g. Ctenis and Nilssonia)
known in the Siberian and Tetori-type floras
do represent markedly toothed or serrate
pinna margins. Such features have not beenrecognized in the bennettitalean andcycadalean leaves known in the Ryoseki-typefloras extensively distributed in the Outer
Zone of Japan, South China and Southeast
Asia.
The Japanese specimens examined are kept
in the National Science Museum, Tokyo and
the Chinese ones in the Nanjing Institute ofGeology and Palaeontology, Academia
Sinica.
Acknowledgements
The first author (Sun) expresses his sincere
thanks to the colleagues of the 9 1020 I-Projectof the Nanjing Institute of Geology andPalaeontology, Academia Sinica for their
help in collecting the fossil plants, to Prof.
S.J. Shui for his kind introduction to thestratigraphy of the Hegan Coal-Field, and to
Dr.K. Saiki for assistance in making theSEM-photomicrographs at the University ofTokyo. His thanks are also extended to the
Kambayashi Scholarship Foundation inTokyo for financial support during his stay inJapan for 9 months, and to the Urban Land
System Institution Co., Ltd., Tokyo for providing him with lodging facilities free ofcharge.
We also thank Dr. Shya Chitaley of theCleveland Museum of Natural History, Ohiofor going over the language of the present
man uscri pt.
This study was supported in part by aGrant-in-Aid for Scientific Research from the
Ministry of Education, Science and Culture(no.02640615 given to Kimura and no.
04640725 given to Kimura and Ohana).

266

Ge Sun el al.

1

>(r

]

150·

l

f."

\

-

110· --40.

960.

Neozamites species from China and Japan

267

Description
Bennettitales
Genus Neozamites Vachrameev, 1962
Neozamites elongatus Kimura et Sekido
Figures 3a, 4a, 5, 6

1971.

Neozamites elongaills Kimura et Sekido:
Kimura and Sekido, p. 192, pI. 24, figs. 1-4, textfigs. 3-4 (Oguchi Formation; Figures 1-21 and2-21).
1980. Neozamites aff. verehojanensis Vachrameev:
Zhang et al., p. 269, pI. 168, figs. 5,8 (Sunjiawan
Formation; Figure 1- 10).

Materia/.-Examined specimens; C-23
(H29-3) and C-27 (H29-2).
Locality and Horizon: Hegan Coal-Field(roughly 47'20'N, 130 17'E; Figure I-I),
Heilongjiang, Northeast China; Lower
Cretaceous Shitouhe Formation (of nonmarine origin).
Occurrence (Frequency): Possibly common.
Description of specimens.- The leaves36']0'-

0I

136' 50'I

136' 55'

Figure 2. Localities of Neozamites in the Inner
Zone of Japan.

+- Figure 1. Localities of Neozamites species in China (Northeast China), Eastern Siberia and Japan(Inner Zone). No Neozamites species has been recorded from outside areas shown in this map. All
specimens of Neozamites species mentioned below were derived from Lower Cretaceous plant sites (F;formation).
1. Hegan, Heilongjiang (Shitouhe F.); the present work; N. eiongaills Kimura et Sekido. 2. Qitaihe,
Heilongjiang (Dongshan F.); Zhang et al., 1980; Zheng and Zhang, 1983; N. denlielilaills (Kryshtofovich
et Prynada) Vachrameev, N. lebedevii Vachrameev and N. verehojanensis Vachrameev. 3. Mudanjiang,
Heilongjiang (Houshigou F.); Zhang el al., 1980; N. verehojanensis Vachrameev. 4. Liangjiang, Antu
and Tongfosi, Jilin (Dalazi F.); Zhang, C. B., 1986 (illustration only), Geological Survey of Jilin, 1989; N.
verehojanensis Vachrameev. 5. Mengjia, Jiutai, Jilin (Yingcheng F.); Yang and Sun, 1982; N. verehojanensis Vachrameev. 6. Fuxin, Liaoning (Shahai F.); Zhang et al., 1980; N. cf. verehojanensis Vachrameev. 7. Har-Zuo, Liaoning (Shahai F.); Zhang et al., 1980; N. cf. lebedevii Vachrameev. 8.
Zalainor, Nei Mongol (Yimin F.); Chen el al., 1988; N. sp. (list only). 9. Heihe, Heilongjiang (Bacheligou F.); Zhang et al., 1980; N. I'erehojanensis Vachrameev. 10. Beipiao, Liaoning (Sunjiawan F.);
Zhang et al., 1980; N. aff. verehojanensis Vachrameev (= N. elongatlls, in the present work). 11. Liampeska River region, Lena Basin (Batylykh F.); Vakhrameev, 1962; N. verehojanensis Vachrameev (originaldescription). 12. Lower course of the Aldan River region (Ekseniakh F.); Vakhrameev, 1962; ditto. 13.
Ganjukha, Zyrianka Basin (Siliap F. ?); Vakhrameev, 1962; ditto. 14. Dzigansk Basin (Khatylykh F. andlower part of Ekseniakh F.); Vassilevskaja, 1966 and Abramova, 1970; N. verehojanensis Vachramcev andN. ? sp. 15. Vilui River region (Khatylykh F.); Vakhrameev, 1962; N. lebedevii Vachrameev (originaldescription). 16. Omsukchan, Magadan (Omsukchan F.); Samylina, 1976; N. verehojanensis Vachrameev. 17. Kanstantinovka, Sujfun Basin (Ussuri and Galenka F.); Krassilov, 1967; N. dentielilatus(Kryshtofovich et Prynada) Vachrameev. 18. Amba-Bir River region, Sujfun Basin (Galenka F.); ditto.
19. Tchigan Cape, west Sutchan Basin (Starosutchan F.); ditto. 20. Big Eljdugo River region, west
Sutchan Basin (Galenka F.); ditto. 21. Mekkodani Valley, Tetori Basin (Oguchi F.); Kimura and Sekido,
1971; N. elongalus Kimura et Sekido (original description). 22. Okurodani Valley, Tetori Basin (Akaiwa
F. equivalent); the present work; N. intermedills, sp. nov.

268

Ge Sun et al.

obtained are fragmental and thus whether theleaves are simply pinnate or bipinnate areunknown, although the leaves of Neozamites
lebedevii described by Zheng and Zhang(1983) (Figure 1-2) are clearly bipinnate.
Therefore, it would be possible that our'leaves' might correspond to the 'penultimate
pinnae' of a bipinnate leaf. In this paper,
however, we use 'leaf or 'leaves' in referring
to our specimens hereafter. The largest leaf
fragment is more than 7.5 cm long and up to
8.2 cm wide with a thin axis, 1.5-2.0 mm
thick, sending off opposite or often
subopposite pinnae at a wide angle. The
angle is reduced distally. The pinnae areattached to the upper part of the axis by asmall central area of pinna base, elongate
rectangular in outline, typically 4.5-5.0 cm
long and 0.7-1.1 cm wide, with bluntly or
subacutely pointed apex and slightly cordate
base, pinna base is not auriculated. Pinna
margins are markedly toothed by 5-6 alternate pairs of teeth or spines directed forward.
Veins are originated from the pinna base,then run in parallel at the proximal half,radiated distally and forked dichotomously at
all levels (Figure 3a); the density is 18-22 in
number per centimeter. Each tooth or spine
receives 1-2 veins. At the proximal medianpart, veins are often grouped together to givea midvein-like appearance (see the holotype ;Kimura and Sekido, 1971).
Cuticle is not preserved in the holotype andothers collected from the Oguchi Formation(Figures 1-21 and 2-21). But fortunately atpresent we have rather well-preserved cuticle
as described below.
Cuticle is essentially hypostomatic. The
upper cuticle is thicker (2-3,um thick measured at a fold) than the lower one, andconsists mostly of ordinary cells which arerectangular, 70-80,um long and 50-70,um
wide, forming longitudinal rows (Figure 5a).
Their anticlinal and end walls are sinuous(wave length is 15-20,um and amplitude is 810 ,urn), but the ordinary cells along the upper
pinna margin are with nearly straight orslightly undulated walls, 80-200,um long and20-25,um wide, and with occasional abortive
stomata [Figures 5b, 5c (left)]. Pericl i nal
walls are unexceptionally smooth.
Lower cuticle consists of alternation of
non-stomatal and stomatal zones. Eachnon-stomatal zone consists of 4-5 longitudinal rows of ordinary cells which are elongate
rectangular with less sinuous anticlinal andend walls, and corresponds to each veincourse (Figure 5d). Each stomatal zone consists of 3-4 intermittent stomatal rows orsometimes disorderly disposed stomata [Figures 5c (right)-5eJ. Stomatal complex issyndetocheilic, round-elliptic in surface view,
about 30-35,um in diameter. Guard cells
are sunken and moderately cutinized, but
their polar extensions are usually exposed;
apertures are usually oblique and occasionally transversely oriented. The ventral(inner) side of subsidiary cell is strongly
cutinized, usually with a single papilla whichis also strongly cutinized, and covers the
middle part of guard cell to some extent(Figures 5f, 6b). The density of stomata is
about 80 in number per square millimeter.
The ordinary cells in the stomatal zones are
not so elongated and irregular (Figure 5e),40-60,um (major) and 30-40,um (minor) insize; their anticlinal and end walls are lesssinuous and are usually delicate and indistinct. Reproductive organs are not known.
Discussion and comparison.-Externally,~ Figure 3. Neozamites species. 3a. Neozamites elongatus Kimura et Sekido. One of the Chinese
specimens drawn partly from Figure 4a, showing the mode of pinna attachment, elongated pinnae withmarkedly developed marginal teeth, and venation. 3b-d, Neozamites intermedius Nakazawa, Ohana etKimura, sp. nov. 3b, Drawn partly from Figure 4c, showing the thick axis (dotted lines), the mode of pinna
attachment, shorter pinnae with markedly developed marginal teeth, and venation. 3e, A pinna fragment
drawn from Figure 4b, showing the toothed margins and venation. 3d, A pinna fragment drawn from NSM
PP-9033 (paratype).

960.

\
---

~\

Neozamites species from China and Japan269

Ge Sun el al.

270

4b

4d

5cm

4c
Figure 4. Neozamites species. 4a, Neozamites elongatus Kimura et Sekid o. One o f the Chinese
specimens [ Reg. no . C-23 (H 29-3)J. 4b- d, Neozamiles intermedius Na kazawa, Oh a na et Kimura, sp. nov.
4b- c, paratypes (Reg. no. NSM PP-9029). 4d, ho lotype ( Reg. no. NSM PP- 9028 ). Its ri ght side o f lea f
is deformed in th e co urse o f preserva ti o n.

960.

Neozamites species from China and Japanthe present leaves agree well with the original
leaves of Neozamites elongatus Kimura et
Sekido. However, they have much more
crowded veins than does the original Neozamites elongatus, this being the only
difference between them. According to additional leaves of the Japanese Neozamites
elongatus newly collected, the vein density is
so variable according to the position of a leaf
or individually that it can not be used as adiagnostic feature to make the specific separation.
The leaves of the type species, Neozamitesverchojanensis resemble externally the present
leaves, but differ from the latter in that in theformer, pinnae are markedly asymmetricalbetween their acroscopic and basiscopic sides
and the pinna margins are much more
toothed, with each tooth further dissected
into 2-3 spines distally.
The leaves regarded by Vassilevskaja(1966) as Neozamites verchojanensis from theLower Cretaceous of Lena Basin (Dzigansk ;Figure 1-14) have large teeth with acutely
pointed tip and much crowded veins (26-30
in number per centimeter in density). Unfortunately in the original leaves of Neozamitesverchojanensis, cuticle has not been recorded.
Neozamites lebedevii was first described by
Vakhrameev (1962) on the basis of externaland cuticular features of the leaves, also fromthe Lower Cretaceous of Lena Basin (Vilui
River region; Figure 1-15). Its leaves are
distinguished from the present ones in thatexternally the former pinnae are short rectangular in form, each with a marked pair ofauricles at the base, and microscopically the
anticlinal and end walls of ordinary cells on
the upper cuticle are strongly sinuous. On
the lower cuticle a large papilla is well developed at the middle of ventral side of a subsidiary cell and the dorsal walls of subsidiarycells are sinuous.
In the leaves of Neozamites denticulatus
known from the Lower Cretaceous of Southern Primorye (Kryshtofovich and Prynada,
1932; first described as Otozamites

271

denticlilatlls; Vakhrameev, 1962; Krassilov,
1967), pinnae are elongated with not auriculated but with cordate base, and the marginal teeth are smaller and sometimes indistinct. Therefore, Neozamites denticulatus iseasily distinguished from N. elongatus.
Under the circumstances, we refer ourleaves to Neozamites elongatlls. Neozamites
aff. verchojanensis illustrated by Zhang et al.
(1980) from the Lower Cretaceous Sunjiawan
Formation (At Beipiao), Liaoning, Northeast
China is referable externally to N. elongatus.
It is highly probable that Neozamites
denticulatus illustrated also by Zhang et al.
(1980) from the Lower Cretaceous Dongshan
Formation (Boli Basin), Heilongjiang is referable to N. elongatus.
Neozamites verchojanensis described by
Yang and Sun (1982) from the Lower
Cretaceous Yingcheng Formation (Jiutai,
Jilin; Figure 1-5) resembles N. elongatlls.
But in the former, pinna teeth are short, triangular and its cuticle is not known. Therefore, it is difficult to make further comparison
between the two.
A dditional diagnosis (cuticular diagnosis
added to the diagnosis of external features) :Cuticle essentially hypostomatic. Upper
cuticle consisting of elongated ordinary cells
forming longitudinal rows. Occasional andabortive stomata present along the pinna
margin. Anticlinal and end walls of ordinary cells sinuous but those along the pinna
margin nearly straight. Periclinal walls
smooth.
Lower cuticle consisting of alternation of
non-stomatal and stomatal zones. Veincourses corresponding to the non-stomatalzone indicated by 3-4 files of much elongatedordinary cells. A stomatal zone consistingof 3-4 intermittent or sometimes disorderly
disposed stomata. Stomata syndetocheilic,round-elliptic in form, obliquely and rarelytransversely oriented. Ventral side of subsidiary cell strongly cutinized, sending off asingle papilla from the middle point, covering
the guard cell to some extent. Ordinary cells

Ge Sun et al.

272

in the stomatal zone irregular in form and lesssinuous in wall but nearly isodiametric.

Neozamites intermedius Nakazawa,
Ohana et Kimura, sp. nov.
Figures 3b-d,4b-dMaterial.-Holotype; NSM PP-9028.
Paratypes; NSM PP-9029, 9033. Examined
specimens: NSM PP-9030, 9032, 9034-9041.
Type locality and horizon: Upper courseof the Okurodani Valley (roughly 36'02'12"
N, l36'53'09"E) (Figures 1-22 and 2-22), a
branch of the Okamigo River, Shokawamura, Ono-gun, Gifu Prefecture: Akaiwa
Formation equivalent (Possibly Aptian inage).
Etymology: According to the intermediate
pinna form between those of Neozamites
elongatus (or N. verchojanensis) and N.
lebedevii.
Occurrence (Frequency): Locally common.
Description.-Leaf is pinnate, but wholeleaf is unknown; preserved leaf is obI anceolate In outline, narrowing gradually
toward the base with a rather thick axis andpetiole. The preserved petiole is 0.5 cm
thick at base (but its very base is unknown),
and bears alternate or subopposite pinnae at awide angle or often perpendicular. The pinnae are attached to the upper sides of axis by
the small region which is 0.5-0.75 mm longand 1.25 mm wide, at or near the middle ofpinna base. The pinnae represented at the
middle portion of a leaf are oblong or rectan-

gular, typically 3 cm long and 1.4 cm wide,with bluntly pointed apex and slightly cordate base. But they are reduced in size,
becoming triangular toward the leaf base, thelowest one being 1 cm long and 0.9 cm wideat the base. The margins of typical pinnae
are markedly toothed except for proximal
one-third where they are usually entire. The
teeth are 3-4 alternate pairs and directed
forward. Each tooth is often further dissected into two lobes by a shallow sinus; thedistal lobe is usually larger and proximal one
smaller in size.
The veins are markedly radiated from thepoint of attachment, forked dichotomously at
all levels; the density is· 26 in number per
centimeter at the middle of pinna. Eachtooth receives 2-6 veins.
Cuticle is not preserved. Reproductive
organs are not known.
Discussion and comparison.-It is certainthat the present leaves belong to the bennettitalean genus Neozamites Vachrameev,
1962, because their Otozamites-like pinnae
are markedly toothed along the margins, andcuticles of two Neozamites species, N. elongatus and N. lebedevii are confirmed by thepresent work and by Vakhrameev's work
(1962) as bennettitaleans respectively.
The present leaves are characterized bytheir intermediate features in pinna size andform between the species with elongated pinnae (Neozamites verchojanensis, N. elongatus
and N. denticulatus) and the species withshorter pinnae (N. lebedevii).
The pinnae of Neozamites lebedevii resem-~
Figure 5. Neozamites elongatus Kimura et Sekido. Prepared from the specimen numbered C-23 (H29-3). Sa, Upper cuticle, showing ordinary cells with sinuous anticlinal and end walls, and smooth
periclinal walls. Sb, Upper cuticle along the pinna margin, consisting of longitudinally elongated ordi narycells with less sinuous cell walls. Sc, Upper cuticle (left side) consisting of marginal ordinary cells and lower
cuticle (right side) consisting of stomata and polygonal ordinary cells with less sinuous cell walls. f; a foldbetween upper and lower cuticles. oc; an occasional abortive stoma. Sd, Lower cuticle, showing scatteredstomata and the vein-course cells (Vc) with thickly cutinized anticlinal and end walls. Se, Lower cuticle,
showing the scattered stomata and the ordinary cells with rather delicate and indistinct anticlinal and end
walls. The stomatal apertures are usually oblique or occasionally transverse in direction. Sf, A stoma insurface view. Gc; sunken guard cell, Sc; thickly cutinized subsidiary cell, P; a thickly cutinized papillaoriginated from the central region of each subsidiary cell to face oppositely with the other, at the generalsurface of lower cuticle.

960.

5c

Neozamites species from China and Japan

n

J

273

Ge Sun et al.

274

0157

- 12kV

20~m

960.

Neozamiles species from China and Japan

ble those of the present specimens, but are
distinguished from the latter by their markedly developed basal acroscopic and basiscopic
auricles as shown by Vakhrameev (1962) and
Zheng and Zhang (1983), instead of having
slightly cordate base as in the latter. Therefore, we (Nakazawa, Ohana and Kimura)here propose Neozamiles intermedius as a
new species to accommodate the present
leaves.
The bennettitalean leaves had been considered to be simply pinnate (except for Nilssoniopteris leaves which have been considered
to be not pinnate) until the discoveries ofbipinnate bennettitalean leaves represented
by Coreanophyllul11 variisegl11entwll Kimuraet Kim (Kimura and Kim, 1982, 1988),
Nipponoptilophyllul11 bipinnatum Kimura et
Tsujii (Kimura and Tsujii, 1984), N.
ryosekiense Ohana, Kimura et Kawazoe(Ohana el al., 1989) and Neozamites lebedevii
Vachrameev (Zheng and Zhang, 1983).
Diagnosis.- Leaf pinnate, medi urn-sized.
(Whole leaf unknown.) Petiole rather thick.
An axis sending off alternate or subopposite
pinnae. Pinnae attached by a small region
near the middle of pinna base on the uppersides of axis, typically oblong or short rectangular in form with blunt apex and slightly
cordate base. Proximal pinnae reduced insize, becoming triangular. Pinna margins oftypical pinnae markedly toothed but marginsof the proximal pinnae nearly entire. The
teeth directed forward and each often further
divided into two lobes by a shallow sinus;
distal one larger in size than proximal one.
Veins numerous and radiated from the small
region of pinna attachment and forked dichotomously at all levels. (Cuticle and reproductive organs not known.)

275

References
Abramova, L.N., 1970: Lower Cretaceous flora of
the Dzigansk and adjacent regions in the Lena
Basin. In 'Study report of palaeontology andstratigraphy, vol. 29'. NIIGA. p.36-57, pis.
1-10. (in Russian with English abstract)
Chen, F., Meng, X.Y., Ren, S.Q. and Wu, c.L., 1988:
The Early Cretaceous flora of Fuxin Basin andTiefa Basin, Liaoning Province. 180 p., 69 pis.
Geol. Publ. House, Beijing. (in Chinese withEnglish summmy)
Geological Survey of Jilin, ed., 1989: Regional Geology of Jilin Province. 698 p., 10 pis. Geol. Pub I.
House, Beijing (in Chinese with English summary).
Kimura, T., 1987a: Geographical distribution of
Palaeozoic and Mesozoic plants East and Southeast Asia. In, Taira, A. and Tashiro, M. eds.,
Historical biogeography and plate tectonic evolution of Japan and Eastern Asia. p. 135-200.
Terrapub., Tokyo.
- - , 1987b: Recent knowledge of Jurassic and
Early Cretaceous floras in Japan and phytogeography of this time in East Asia. Bull. Tokyo
Gakugei Univ., ser. IV, vol. 39, p. 87-115.
Kimura, T. and Kim, B.K., 1982: Coreanophyllum
variisegmenlum gen. et sp. nov. from theDaedong Supergroup, Korea. Proc. Japan
Acad., vol. 58B, no. 6, p. 152-155.
- - and - - , 1988: New taxa in the Late Triassic
Daedong flora, South Korea. Part I. Trans.
Proc. Palaeont. Soc. Japan, N.S., no. 152, p. 603624.
Kimura, T. and Sekido, S., 1971: The discovery of
the cycad-like leaflets with toothed margin fromthe Lower Cretaceous ltoshiro Subgroup, theTetori Group, Central Honshu, Japan. Ibid.,
no. 84, p. 190-195, pI. 24.
Kimura, T. and Tsujii, M., 1984: Discovery ofbipinnate Ptilophyllum leaves (Bennettitales) from theUpper Jurassic Tochikubo Formation, Fukushima Prefecture, Northeast Japan. Proc. Japan
Acad., vol. 60B, no. 10, p.385-388.
Krassilov, V.A., 1967: Early Cretaceous flora ofSouthern Primorye and its significance for stratigraphy. 364 p., 93 pis. Nauka, Moscow. (inRussian)+-- Figure 6. Neozamites elongatlls Kimura et Sekido (SEM micrographs of cuticle). 6a, Upper cuticle,
showing the ordinary cells with faintly preserved sinuous anticlinal and end walls (surface view). 6b, A
broken stoma, showing a pair of oppositely disposed and thickly cutinized papillae. The lower papilla isbroken and the guard cells are missing in the course of maceration (surface view). 6c, Lower cuticle,
showing scattered stomata (inner view). 6d, Lower cuticle, showing a stoma enlarged (inner view). Gc,
guard cell; Sc, thickly cutinized flange of subsidiary cell.

Ge Sun et al.

276

Kryshtofovich, A.N. and Prynada, V.D., 1932: Contribution to the Mesozoic flora of the Ussuriland.
Bull. U. Ceol. Prosp. Sur!'., USSR, vol. 51, no.
22, p. 363-374, pis. 1-2.
Ohana, T., Kimura, T. and Kawazoe, A., 1989: Somefossil plants newly found from the Lower
Cretaceous of Kochi Prefecture, in the Outer
Zone of Japan. Asian Jour. PI. Sci., vol. I, no.
2, p.53-68.
Samylina, V.A., 1976: The Cretaceous flora of
Omsukchan (Magadan District). 208 p., incl. 48
pis. Nauka, Leningrad. (in Russian)Vakhrameev, V.A., 1962: New Early Cretaceous
cycadophytes from Yakutia. Paleont. Zhurnal,
no. 3, p. 123-129, pis. 12-13. (in Russian)Vassilevskaja, N.D., 1966: Some Early Cretaceous
plants from Dzigansk region (Lena Coal-bearing
Basin). Sci. Rep., Inst. Arctic Ceol., no. 15, p.
49-76, pis. 1-7. (in Russian)Yang, X.L. and Sun, L.W., 1982: Fossil plants fromthe Shahezi and Yingcheng Formations in southern part of the Songhuajiang-Liaohe Basin,
NE-China. Acta Palaeont. Sinica, vol. 21, no.
5, p. 588-596, pis. 1-3. (in Chinese with English
abstract)
Zhang, C.B., 1986: The middle-late Early Cretaceous
strata in Yanji Basin, Jilin Province. J. Changchun Ceol. Inst., no.2 (1986), p. 15-27, pis. 1-2.
(in Chinese with English abstract)
Zhang, W., Chang, c.c. and Zheng, S.L., 1980:
Plants. In 'Shenyang Inst. Geol. Min. Res. ed. :Palaeontological Atlas of Northeast China. II.
Mesozoic and Cenozoic volume'. p.221-311,
pis. 103-191. Geol. Publ. House, Beijing. (inChinese)Zheng, S.L. and Zhang, W., 1983: Middle-late Early
Cretaceous flora from Boli Basin, eastern
Heilongjiang Province. Bull. Shenyang Ins!.
Ceol. Min. Res., no. 7, p.68-98, pis. 1-8. (inChinese with English abstract)*

Geographical and stratigraphical names: Japan: Akaiwa iJG!J!'L Mekkodani § Hii-, Oguchi
Okamigo ~J::~~, Okurodani *~ii-, Ryoseki ~E, Tetori -=.f!&. China: Antu
IZI, Bacheligou;,,:$:tJ$j, Beipiao~t~, Boli~flj, Chengzihe:f$;.:r[PJ, Dalazi*lli..:r,Dongshan JlHLJ, Fuxin Jl!.¥JT, Har-Zuo Ilttc, Hegan ~F.lii, Heihe ~[PJ, Heilongjiang ~~IT, Houshigou i~E~, Jilin i5f;t, Jiutai 11 iJ, Jixi ~l1§, Liangjiang fi!UIT, Liaoning ~$, Menjia:iiE
Mudanjiang!j± BIT, Qitaihe -f.:; iJ [PJ, Shahai 7PW, Shitouhe EM [PJ,
Sunjiawan 1**~, Tongfosi ~-Ml~, Yimin fJtliiJ:, Yingcheng ~:f$;, Zalainor tLW~~.
~ P,

*,

J:\=lOO*~t$<!? J: UEl*I*J*O)T$BsE]M'J?f~J?nt: Neozamites ~ (r(::f.TT A 13)
1t1:l2fl: Neozamites~fi~;fTT.A§K.~-t0~:a:-X;J~c Lt-.:*-g~"'e, 1962'¥-, Vak-hrameev f;:' J: I) ~Ij~. t~!1i; ~ nt-.:. *~O)!f!f~Hi, :flJtO)~j\~X'1!f U ,tJ3.i'Ili'rb:t5 0 ,: C,:to J: LJ-tO)£tI:lb\ :/ ~ ~ TlIHm, ,*,ooJluumO)*-ll'il5!:to J: LJ B *Ji'J*O)T-ll'ilS'IE*f;:.~NGn0': U:.t50. ts::tomtaw1'i'iO)T-ll'ils'IE*b'G t, *~f;:.~-t0 c~.:Z. Gn0f.U:£
tI:l-t0.
*~f;:.~-t0f.Ht, ~'ii-ef;:'4flb:ffiGn, 5'tllfl (Nelongatus) fi, 1971,¥-,-=.f!&~~fI'~P~b'G~!1i;~nt-.:b:, R~ts:b:G, -to) cuticle fi*1f~n-C~'ts:b'"?t-.:. 4@], ,*,OOJiUt-ll'ilO)~F.liiERB3b'G, cuticle b:J: <*1f~nt-.: N elongatus b:~Gnt-.:O)-e, ,:,: f;:'-t0)~~F*IH:a:-¥rH5·-t 0. i t-.:, il&:Jl!.Jljt*~ii-f;:.5!;fp-t 0 T-ll'ilS'IE*b' G~ Gnt-.:~*fi,§fJ G b'f;:'*~f;:'~ L, b''0¥JTfl L -ct~1ib Gn 0 0)""(', ,:,: f;:.': G O)~~:;$Xxt L -C N
intermedius, sp. nov. :a:-~Ill§ L t-.:.
1** . {'l'lRfli • *reB;;.:r • :;t:ttJ¥§fJcn

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 172, pp. 277-291, 5 Figs., December 30, 1993

961.

MIDDLE PERMIAN FORAMINIFERS FROMBAN NAM SUAI THA SA-AT, CHANGWATLOEI, NORTHEAST THAILAND*
KA TSUMI UENO
Institute of Geoscience, University of Tsukuba,
Ibaraki, 305
andSUMIO SAKAGAMI
Department of Earth Sciences, Faculty of Science,
Chiba University, Chiba, 263

Abstract. Middle Permian foraminifers including such fusulinaceans as Verbeekina
verbeeki (Geinitz), Pseudodoliolina cf. pselldolepida (Deprat), and Presllmatrina cf. neoschwagerinoides (Deprat) are found from a small limestone outcrop exposed at about 2 km
northwest of Ban Nam Suai Tha Sa-at, east of Loei, northeast Thailand. The present
fusulinacean fauna has a close resemblance with those of the Neoschwagerina simplex (B6)and Presllmatrina schellwieni (B7) Zones of the Ratburi (or Saraburi) Limestone in the Sara
Buri area, central Thailand, and the lower Murgabian (Neoschwagerina simplex Zone) of the
Pamirs, Central Asia. It is referable to early Murgabian (late Middle Permian) of thePermian standard stratigraphic scheme in the Tethyan Province in age. This is the first
discovery of the late Middle Permian foraminiferal fauna characterized by the verbeekinidand neoschwagerinid fusulinaceans in the Loei area.

Key words. Foraminifers, Loei, Middle Permian, Murgabian, Ratburi (or Saraburi)
Limestone, Thailand.

Yakhtashian fusulinaceans (Igo et al., 1993)
were published. However, the Middle Permian fossils as well as their biostratigraphy
and faunal characteristics in these areas are
too poorly known for a detailed discussion,as yet covered only by some preliminary
reports (Pitakpaivan, 1965; Borax and
Stewart, 1966; Toriyama, 1967).
In 1988, we visited the Loei area for thegeologic and paleontologic investigations of
the Upper Paleozoic strata in Thailand. We
collected a limestone rock sample from asmall outcrop near Ban Nam Suai Tha Sa-at,
about 20 km east of Loei, and discriminated a
Middle Permian foraminiferal fauna which is

Introduction

In the Loei and Wang Saphung areas ofnortheast Thailand where the Ratburi (orSaraburi) Limestone is widely distributed, theMiddle Carboniferous to Lower Permian
fusulinaceans were reported and their biostratigraphy and zonation were examined indetail (lgo, 1972). Very recently, two additional contributions concerning the Upper
Carboniferous Gzhe1ian foraminifers (Uenoand Igo, 1993) and the upper Lower Permian• Received December 21, 1992; accepted June 24,
1993
277

Katsumi Vena and Sumia Sakagami

278

400~~

~ 380~}~.
~Ban Nam~~suai~g

M

••

,;°
~D

31 ','

Fossil Locality
Siamo~~3'O~

1I

Figure 1.
Thailand.

Index map showing the fossil locality near Ban Nam Suai Tha Sa-at, east of Loei, northeast

poorly understood in this area.
In the present paper, we discuss the character and geologic age of this foraminiferal
fauna. Moreover, we illustrate most of theforaminifers and describe all thefusulinaceans.
~

Fossil locality and foraminiferal fauna
The fossil locality studied herein is situatedat the roadside of Route 2318, about 2 km
northwest of Ban Nam Suai Tha Sa-at, east ofLoei, northeast Thailand (Figure I), where a

Figure 2. 1: Eotuberitina sp., longitudinal section, IGUT-KU0794. 2: Eotuberitina reitlingerae
Miklukho-Maklay, longitudinal section, IGUT-KU0795. 3: Tuberitina sp., longitudinal section,
IGUT-KU0796. 4: Pseudoammodiscus? sp., axial section, IGUT-KU0797. 5: Eolasiodisclls sp., tangential section, IGUT-KU0798. 6: Spiroplectammina sp., longitudinal section, IGUT-KU0799. 7:
GlobivalvlIlina sp., lateral section, IGUT-KU0800. 8, 9: ParaglobivalvlIlina? sp., lateral sections, IGUTKUOSOl, IGUT-KUOS02. 10, 11. Agathammina sp., 10: longitudinal section, IGUT-KUOS03, 11: transverse section, IGUT-KUOS04. 12, 13. Agathammina pusilla (Geinitz), 12: transverse section, IGUTKUOS05, 13: longitudinal section, IGUT-KUOS06. 14: Pselldoglomospira sp., horizontal section, IGUTKUOS07. 15: Glomospira? sp., horizontal section, IGUT-KUOSOS. 16, 17. Mlillidisclis sp. A., 16: axial
section, IGUT-KUOS09, 17: sagittal section, IGUT-KUOS 10. 18: Pachyphloia sp. A, lateral section,
IGUT-KUOSIl. 19: Nodosaria sp., longitudinal section, IGUT-KUOSI2. 20: Pachyphloia sp. B, longitudinal section, IGUT-KUOSI3. 21: Frondina? sp., longitudinal section, IGUT-KUOSI4. 22-24.
Lunucammina sp., 22, 23: longitudinal sections, IGUT-KUOSI5, IGUT-KUOSI6, 24: lateral section,
IGUT-KUOSI7. 25: Telrataxis sp., axial section, IGUT-KUOSIS. 8, 9: x20, 15,20: x30, 2, 3, 6, 7,10
-14, 16-19,21-25: x40, 1,4, 5: X 100.

961.

Middle Permian joramin!fers from NE Thailand

massive limestone bed crops out. Limestone
microfac ies in this locality is black , impure
a lgae-foraminife ral grainstone. According
to the geologic map of Changwat Loei (scale
I : 250,000) compiled by Vimuktanandana(1988), the Permi an N am M ahoran Formation composed mainly of a limestone, sandstone, shale, and chert is distributed here.
W e identified the following sma ll er for-279

ami n i fers and fusu I i naceans from the samp leco ll ected at the fossi l loca lity st udi ed herein.
Smaller foram inifers are illustrated o n Figures 2 and 3.

Smaller foraminifers: EOlUberilina reillingerae
Miklukho-Maklay, EOlUberilina spp., Tuberilina
sp. , Spiroplectal11l11ina sp. , PseUdOal111110disclIs ?
sp. , Eoiasiodisclis sp. , PselldoglOl11ospira sp. ,

5

1

4

11

6

13

9
15

14
12
10

17

19

20

21

23

24

18

25

Kalsum i Vena and Sumia Sakagami

2S0

1
2

3

4

5

7

6

11

15

9

13
12

10

14

16

Figure 3. 1: EndOlhyra sp., ax ia l secti o n, IGUT-KUOS I9. 2, 3. Neoendolhyra sp., 2 : ax ia l section , IGUT-K U0820, 3: sagitta l section, IG UT-K U082l. 4,5: Climacammina sp. , longitudin a l secti o ns,
IGUT-KU0822, IGUT-KU0823. 6- 11: Multidiscus sp. B, ax ia l secti o ns, IGUT-KU0824, IGUTKU0825 , IGUT-KU0826, IGUT-KU08 27, IGUT-KUOS28, IGUT-KU0829. 12,13. Climacammina gigas
Suleimanov, 12: longi tudinal section , IGUT-K U0830, 13: latera l section , IGUT-KU083 1. 14: Neodiscus cf. permicus (Gro zdilo va), ax ial secti on , IGUT-KU0832. 15, 16: Climacammina valvulinoides
La nge, longi tudinal sectio ns, IG UT-K U0833 , IGUT-K U0834. 13: X 15, 12, 15, 16: X 20, 4- 11, 14 : X
40, 2, 3: X 75 , 1: X 100.

961.

Middle Permian foraminifers from NE Thailand

Glomospira? sp., Globivalvulina spp., Paraglobivalvlllina? sp., Tetrataxis sp., Endothyrasp., Neoendothyra sp., Climacammina gigas
Suleimanov, C. valvlliinoides Lange, Climacammina sp., Lunllcammina spp., Pachyphloia sp. A,
Pachyphloia sp. B, Pachyphloia spp., Nodosariasp., Frondina? sp., Neodisclls cf. permiclls(Grozdilova), Mliitidisclis sp. A, Mliitidisclis sp.
B, Agathammina cf. pusilla (Geinitz), Agathammina sp.
Fusulinaceans: Neofllsulinella? pselldogiralldi(Sheng), Neofllsulinella sp., Yangchenia sp.,
Minojapanella (RlIssiella)? sp., Dlinbarllla?
sp., Parafuslilina sp., Nankinella sp., Staffellasp., Pisolina cf. subspherica Sheng, Presllmatrina
cf. neoschwagerinoides (Deprat), Pselldodoliolina
cf. pseudolepida (Deprat), Verbeekina verbeeki(Geinitz).

Among the foraminifers, fusulinaceans areimportant for the discussion of the geologic
age of this foraminiferal fauna. The mostcharacteristic species of this foraminiferal
fauna is Verbeekina verbeeki (Geinitz).
This species is known to flourish in the late
Middle Permian (Murgabian and Midian) in
the Tethyan Province (Ozawa, 1970 andothers). In the Ratburi Limestone of the
Sara Buri area, central Thailand, it occurs in
the Presumatrina schellwieni Zone (B7)
which is correlated with the lower Murgabian(Toriyama, 1975; Ingavat et al., 1980).
Pseudodoliolina cf. pseudolepida (Deprat) is
also abundant in our foraminiferal fauna.
Pseudodoliolina pseudolepida (Deprat) is awell known species in the Tethyan Province
and widely distributed in its upper Middle
Permian deposits (Ozawa, 1970 and others).
A species closely related to Presumatrina
neoschwagerinoides (Deprat) discriminatedfrom our foraminiferal fauna is worthy ofnote because it is characteristic in the Neoschwagerina simplex Zone (lower Murgabian) of the Pamirs, Central Asia, where
the Permian standard stratigraphic scheme in
the Tethyan Province was established (Leven,
1965, 1967, 1980). Presumatrina neosch wagerinoides (Deprat) was also reported fromthe Neoschwagerina simplex (B6) and
Presumatrina schellwieni (B7) Zones of the281

Ratburi Limestone distributed In the Khao
Phrong Phrab area, Sara Buri, central
Thailand (Toriyama, 1975). These zones arecorrelated with the lower Murgabian (Ingavat, 1984).
Neofusulinella pseudogiraudi (Sheng) was
reported from the Qixia (Chihsia) and Maokou Limestones (Misellina and Cancellina
Subzones) of Guizhou (Kueichow), South
China (Sheng, 1963). In Southeast Asia, it is
known to occur from the limestone distributed in the Sungei Sedili area, Johore, Malaysia(Igo el aI., 1979).
Pisolina subspherica Sheng was originallydescribed from the Nankinella orbicularia
Zone of southern Shanxi (Sensi) which iscorrelated with the lower Qixia Limestone(Sheng, 1956). Later, this species was reported from the upper part of the Cancellina
neosch wagerinoides-Parafusulina multiseplata Zone of Zhejiang, where Parafusulina
multiseptata (Scheliwien) and Cancellina
danneri Skinner and Wilde are associated(Wang and Tang, 1986).
Judging mainly from the joint occurrenceof Verbeekina verbeeki (Geinitz), Pseudodoliolina cf. pseudolepida (Deprat), and
Presuma trina cf. neosch wagerinoides (Deprat), the geologic age of this foraminiferal
fauna is considered to be early Murgabian(late Middle Permian) of the Permian standard stratigraphic scheme in the Tethyan
Province proposed by Leven (1980). As
well, other coexisting fusulinacean and smaller foraminiferal genera and species arealmost concordant with this age determination. Biogeographicaliy, this foraminiferal
fauna is a typical one in the Tethyan ProvInce.

Systematic description
All specimens treated in this paper aredeposited in the paleontological collection of
the Institute of Geoscience, University ofTsukuba. The abbreviation IGUT is theprefix for repository numbers of specimens.

282

Katsumi Ueno and Sumio Sakagami

Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea von Moller, 1878
Family Schubertellidae Skinner, 1931
Subfamily Yangcheninae Leven, 1987
Genus Yangchenia Lee, 1933

Yangchenia sp.
Figures 5-8-9

Remarks.- Two poorly oriented sections
were prepared for this study. Massive
chomata and a Fusulinella-type four-layered
spirotheca in this form possibly suggest it tobe referable to the genus Yangchenia, in spiteof its poor orientation.
Materials.-Diagonal sections; IGUTKU0853,IGUT-KU0854.
Subfamily Schubertellinae Skinner, 1931
Genus Neofusulinella Deprat, 1912

Neofusulinella? pseudogiraudi(Sheng, 1963)

the Qixia and Maokou Limestones of Guizhou, South China.
Materials.-Axial section; IG UTKU0856. Oblique section; IGUT-KU0857.
Sagittal section; IGUT-KU0858.

Neofusulinella sp.
Figure 5-10

Remarks.-Keriothecal structure is barely
discernible in some part of spirotheca in thisspecies. This species can be distinguishedfrom Neofusulinella? pseudogiraudi(Sheng) by its larger shell. It is somewhat
similar to Neofusulinella saraburiensis originally described by Toriyama et al. (1969)
from the Khao Phrong Phrab area of Sara
Buri, central Thailand, in the shell size. The
specific identification is, however, postponed
because only one sagittal section is available
for examination.
Material.-Sagittal section, IGUTKU0855.

Figures 5-11-13

Schubertella pseudogiraudi Sheng (MS), 1962, p. 427428, pI. 1, figs. 8-9.
Schubertella pseudogiraudi Sheng, 1963, p.35, 159,
pI. 4, figs. 14-19; Han, 1976, p.26, pI. 8, fig. 8;
Lin et al., 1977, p.34, pI. 6, fig. 20; Chen andYang, 1978 l p.26, pI. 3, figs. 36-37; Igo et al.,
1979, p. 100, pI. 17, figs. 9-10, pI. 18, figs. 11-14;
Xie, 1982, p. 15, pI. 6, fig. 7; Sun et al., 1983, p.
17, pI. 3, fig. 2.

Remarks.- This species was originallyreferred to the genus Schubertella, and mostfusulinacean students have followed thisgenus assignation. However, it is moreadvisable to include it in the genus
Neofusulinella rather than in Schubertella, in
its spirothecal composition and well developed septal pores, although keriothecal structure, which is a diagnostic shell character inthe former genus, is hardly discernible in the
spiro theca of this species.
The present specimens are quite identicalwith the original ones by Sheng (1963) fromFamily Boultoniidae Skinnerand Wilde, 1954
Genus Minojapanella Fujimotoand Kanuma, 1953
Subgenus Russiella Miklukho-Maklay, 1957

Minojapanella (Russiella)? sp.
Figures 5-1-3

Remarks.- This unidentified species isquestionably assigned to the subgenus Russiella because of its highly elongate shell
shape, regularly and intensely fluted septa,
and weakly developed axial fillings. It canbe distinguished from Minojapanella (Russiella) pulchra Miklukho-Maklay in having asmaller shell and less developed axial fillings.
Materials.-Axial sections; IG UTKU0846, IGUT-KU0847. Sagittal section;
IGUT-KU0848.

961.

Middle Permian foraminifers from NE Thailand

Figure 4. 1- 5. Verbeekina verbeeki (Geinitz), 1,2 : sli ght ly ob li q ue axia l sections, IGUT-KU0835,
IGUT-KU0836, 3, 4a, 5: sagitta l sections, IGUT-KU0837, IGUT-KU0838, IGUT-KU0839, 4b : enl a rged
part of 4a, show in g tightly and skew-coi led inn er vo lution s, a nd minute proloculus. 6. Presumalrina cf.
neoschwagerinoides (Deprat) , 6a: axia l section, [GUT-KU0840, 6b : en larged part of 6a. 7- 11.
Pseudodo/io/ina cf. pseudo/epida (Deprat) , 7 : parallel sect ion , [GUT-KU 084 1, 8 : sag itta l secti o n, [G UTKU0842, 9, II: axia l sections, IGUT-KU0843, [GUT-K U0845 , 10 : tangential section, IGUT-K U0844.
1- 3, 4a, 5, 6a, 7- 11: X 10, 4b, 6b : x 40.

283

Katsumi Veno and Sumio Sakagami

284

Genus Dllnbarula Ciry, 1948

Family Staffellidae Miklukho-Maklay, 1949
Genus Nankinella Lee, 1934

Dunbarula ? sp.
Nankinella sp.

Figure 5-14

Figures 5-6-7

Remarks.- The present form has a fusiform
shell, rather loosely coiled volutions, fluted
septa in polar regions, and slightly recrystallized spirotheca. These shell characters aresuggestive of an assignment to the genus
Dunbarlila. The genus assignment is, however, tentative owing to the poor material.
Material.-Axial section; IGUT-KU0859.
Family Schwagerinidae Dunbarand Henbest, 1930
Subfamily Schwagerininae Dunbarand Henbest, 1930
Genus Parafusulina Dunbar
and Skinner, 1931

Remarks.-Nankinella akiyoshiensis originally described by Toriyama (1958) from the
Neoschwagerina craticulifera Zone (Pmp) of
the Akiyoshi Limestone Group, southwest
Japan, is the closest to this unidentified
species except for having a less angularperiphery of shell.
Materials.- Tangential sections; IGUTKU0851, IGUT-KU0852.
Genus Staffella Ozawa, 1925

Staffella sp.
Figure 5-15

Parafusulina sp.
Figures 5-4-5

Remarks.- This unidentified species somewhat resembles Parafllsulina quasigruperaensis originally described by Sheng (1963) from
the Maokou Limestone of Guizhou, South
China. It is also recorded from the
Presumatrina schellwieni Zone (87) of the
Ratburi Limestone in the Khao Phrong
Phrab area, central Thailand. The specificidentification is postponed until sufficientmaterials are available.
Materials.-Axial section; IGUTKU0849. Parallel section; IGUT-KU0850.

Remarks.-Morphological characters of
this species are not examined precisely
because the material is completely replacedby secondary mineralization.
Material.-Axial section; IGUT-KU0860.
Genus Pisolina Lee, 1933

Pisolina cf. subspherica Sheng, 1956
Figures 5-16-22

Compare.Pisolina subspherica Sheng, 1956, p. 183, 205-206, pi.
2, fig. 6; Sheng, 1966, p. 23, pi. 2, fig. 13 (same aspi. 2, fig. 6 of Sheng, 1956, p. 183,205-206); Lin~
Figure 5. 1-3. Minojapanella (Russiella)? sp., 1,2: axial sections, IGUT-KU0846, IGUT-KU0847,
3: sagittal section, IGUT-KU0848. 4, 5. Parafusulina sp., 4: axial section, IGUT-KU0849, 5: paralielsection,IGUT-KU0850. 6,7: Nankinella sp., tangential sections, IGUT-KU0851, IGUT-KU0852. 8,9:
Yangchenia sp., diagonal sections, IGUT-KU0853, IGUT-KU0854. 10: Neofusulinella sp., sagittal section,
IGUT-KU0855. 11-13. Neofusulinella? pseudogiraudi (Sheng), 11: axial section, IGUT-KU0856, 12:
oblique section, IGUT-KU0857, 13: sagittal section, IGUT-KU0858. 14: Dunbarula? sp., axial section,
IGUT-KU0859. 15: Staffella sp., axial section, IGUT-KU0860. 16-22: Pisolina cf. subspherica Sheng,
16,22: sagittal sections, IGUT-KU0861, IGUT-KU0867, 17,19,21: axial sections, IGUT-KU0862, IGUTKU0864, IGUT-KU0866, 18,20: oblique sections, IGUT-KU0863, IGUT-KU0865. 4,5,20,22: X 10,1619,21: X 15,6,7,15: x20, 8, 9: x30, 1-3, 10-14: x40.

961.

Middle Permian foraminifers from NE Thailand

285

286

Katsllmi Veno and Sllmio Sakagami
et al., 1977, p. 16, pI. 2, fig. 20; Ding, 1978, p.
280, pI. 96, fig. 14; Lin et al., 1979, pI. 1, fig. 7;
Wang et al., 1982, p. 97, pI. 24, fig. 7; Xie, 1982,
p. 8, pI. 3, figs. 1-2; Lin, 1984, p. 174-175, pI. 15,
fig. 21; Chen, 1984, p.59, pI. 14, fig. 13; Zhou
and Zhang, 1984, pI. 3, figs. 6-7.

Description.-Shell medium for genus witha broadly rounded periphery and slightly
shorter axis of coiling. Mature shell of 7 to
9 volutions 2.33 to 3.30 mm in length and 2.87
to 3.90 mm in width. Form ratio ranging
from 0.80 to 0.91, averaging 0.84 for 4 specimens. Shell expanded gradually throughgrowth. Proloculus large for shell size. Itsoutside diameter ranging from 0.220 to 0.370
mm, averaging 0.295 mm for 6 specimens.
Spirotheca replaced by secondary mineralization. Keriothecal structure partly observedin spiro theca of some specimens. Septa
short, numerous and unfluted. Low andbroad chomata developed in all volutions.
Tunnel path straight and narrow.
Remarks.-Our specimens almost agreewith the original one described by Sheng(1956) from the Nankinella orbiclilaria Zone
(upper part of the Qixia Limestone) of southern Shanxi except for having a slightly larger
shell and smaller form ratio. Pisolina cf.
sllbspherica Sheng can be easily distinguishedfrom P. excessa Lee by a different shell shape.
Materials.-Axial sections; IGUTKU0862, IGUT-KU0864, IGUT-KU0866.
Sagittal sections; IGUT-KU0861, IGUTKU0867. Oblique sections; IGUTKU0863, IGUT-KU0865.
Family Verbeekinidae Staff
and Wedekind, 1910
Subfamily Verbeekininae Staff
and Wedekind, 1910
Genus Verbeekina Staff, 1909

Verbeekina verbeeki (Geinitz, 1876)
Figures 4-1-5

Descriptive remarks.-Shell large andalmost spherical. Il1ustrated specimens 4.40

to 6.00 mm in length and 4.95 to 5.80 mm inwidth, gving form ratio of about 1.00.
Mature shell attained a maximum of 10 volutions. Inner few volutions tightly and skew
coiled. Outer volutions uniformly expanded. Proloculus minute and 0.040 mm inoutside diameter. Spirotheca thin and composed of a tectum and fibrous keriotheca.
Parachomata numerous but developed only
adjacent to septa.
In Thailand, Verbeekina verbeeki (Geinitz)
was recorded from the Maoteetang Limestoneof Sara Buri, central Thailand (Pitakpaivan,
1965), from Wat Kirinakratanaram, central
Thailand (Toriyama and Pitakpaivan, 1973),
and from the Presllmatrina sehellwieni Zone
(B7) and the overlying limestone conglomerate formation (Upper C Formation) in the
Khao Phrong Phrab area of Sara Buri, central
Thailand (Toriyama, 1975; Toriyama andKanmera, 1977). The following synonymy
can supplement Ueno (1992).
Verbeekina verbeeki, Nie and Song, 1983, pI. 1, fig. 3;
Kahler, 1988, fig. 77; Vachard, 1990, pI. 5, fig. 3.
? Verbeekina ex gr. verbeeki, Vachard and Miconnet,
1989, pI. 3, fig. 2.

Materials.-Slightly oblique axial sections; IGUT-KU0835, IGUT-KU0836.
Sagittal sections; IGUT-KU0837, IGUTKU0838,IGUT-KU0839.
Subfamily Pseudodoliolininae Leven, 1963
Genus Pseudodoliolina Yabeand Hanzawa, 1932

Pseudodoliolina cf. pseudolepida(Deprat, 1912)
Figures 4-7-11

Descriptive remarks.-Shell cylindricalwith a broadly rounded periphery. Mature
specimens having 12 to 15 volutions, about
5.00 mm in length and 2.55 to 2.83 mm inwidth, giving form ratios of 1.77 to 1.96.
Proloculus small and measuring from 0.095 to
0.125 mm in outside diameter. Spirotheca961.

Middle Permian foraminifers from NE Thailand

thin and composed of a tectum and lower lessdense layer. Parachomata well developed.
The compared synonymies are followed by
Ueno (1992). The present form almost
agrees with Deprat's original. However, thespecific identification is tentative because theformer has a smaller proloculus than thelatter.
Materials.-Axial sections; IGUTKU0843, IGUT-KU0845. Sagittal section;
IGUT-KU0842. Tangential section;
IGUT-KU0844. Parallel section; IGUTKU0841.
Family Neoschwagerinidae Dunbar
and Condra, 1927
Subfamily Sumatrininae Silvestri, 1933
Genus Presumatrina Tumanskaya, 1950

Presumatrina cf. neosch wagerinoides(Deprat, 1913)
Figure 4-6Compare.Doliolina neoschwagerinoides Deprat, 1913, p.52-53,
pI. 10, figs. 1-5, 7 (not pI. 10, fig. 6).
Cancellina primigena, Ozawa, 1925, pI. 2, figs. 3- 7
(same as pI. 10, figs. 1-4, 7 of Deprat, 1913, p. 5253).
Cancellina schellwieni, Chen, 1934, p. 105-106, pI. 16,
figs. 5-12.
Cancellina neoschwagerinoides, Sheng, 1963, p.99100,232, pI. 34, figs. 7,8,10-12; Sheng, 1966, p.
144-145, pI. 25, fig. 2 (same as pI. 16, fig. 5 ofChen, 1934, p. 105-106); Toriyama, 1975,87-90,
pI. 17, figs. 14-23; Lin et al., 1977, p. 89-90, pI.
30, fig. II ; Wang et al., 1982, p. 104, pI. 29, fig.
3; Sun et al., 1983, p. 37-38, pI. 10, fig. 9.
Praesumalrina neoschwagerinoides, Leven, 1965, p.
143, pI. 4, fig. 2; Leven, 1967, p. 196-197, pI. 34,
figs. 2, 3, 5, 7; Chediya et al., 1986, pI. 3, fig. 15 ;Wei et al., 1987, pI. 3, fig. 10.
Presumalrina neoschwagerinoides, Ozawa, 1970, pI. 6,
figs. 3-5 (3,4: same as pI. 10, figs. 1,2 of Deprat,
1913, p. 52-53,5: same as pI. 34, fig. 3 of Leven,
1967, p. 196-197); Chen and Yang, 1978, p. 120121, pI. 31, fig. 3.
Praesumalrina schellwieni, Liu et al., 1978, p. 97, pI.
23, fig. I.
Presumalrina ex gr. neoschwagerinoides, Leven, 1982,
pI. 3, fig. 8.

287? Doliolina cf. neoschwagerinoides(?), Colani, 1924, p.
115, pI. 20, fig. 4.
? Cancellina neoschwagerinoides, Kahler and Kahler,
1979, p. 248-249, pI. 9, fig. 5; Chen, 1984, p.68,
pI. 22, figs. 5-7; Yang, 1985, pI. 3, fig. 14; Wang
and Tang, 1986, pI. 2, fig. 8.
? Cancellina cf. neoschwagerinoides, Sun and Zhang,
1988, pI. 4, fig. 24; Zhu, 1990, p.58, pI. 7, figs.
12-13.
not Cancellina neoschwagerinoides, Zhang and Wang,
1974, p. 291, pI. 151, fig. 4.
not Praesumatrina neoschwagerinoides, Liu et al.,1978, p. 97-98, pI. 23, fig. 5.

Description.-Shell fairly large for genus
and fusiform with bluntly pointed polar ends
and slightly convex lateral slopes. Welloriented axial section having 8 Yz volutions,4.34 mm in length and 2.30 mm in width,
giving a form ratio of 1.88. Shell expanded
gradually through growth. Radius vectors of
the first to eighth vol ution 0.21, 0.29, 0.38,
OA8, 0.61, 0.77, 0.92, and 1.09 mm, and form
ratios 1.11, lAO, 1.59, 1.68, 1.62, 1.63, 1.64 and
1.62, respectively. Prolocul us large for shellsize and measuring 0.270 mm in outside diameter. Spirotheca thin and composed of atectum and keriotheca. Thickness of spirotheca of the first to eighth vol ution 0.020,
0.020, 0.015, 0.025, 0.030, 0.035, 0.030, and0.020 mm. Primary transverse septula thin
and well developed in all volutions. Tips ofprimary transverse septula coated by dark
microgranular calcite materials and slightly
swollen, and in contact with tops of parachomata adjacent to septa. One secondarytransverse septulum occasionally inserted
between 2 adjacent primary transverse septulain outer volutions. Parachomata narrowand high, and well developed in all volutions.
Remarks.-Some fusulinacean students
have referred this species to the genus Cancellina. However, Presumatrina is most suitable for genus assignment of this species as init the transverse septula have slightly
solidified and swollen tips. Our specimen isquite similar to the original ones of
Presumatrina neoschwagerinoides describedby Deprat (1913) from Laos in the shell288

Katsumi Veno and Sumio Sakagami

shape, and nature of chomata and transverse
septula. However, the Loei specimen has aslightly larger shell and prolocul us than
Deprat's original. At any rate, the exact
identification is postponed until additional
specimens are available.
In Thailand, Presllmatrina neoschwagerinoides (Deprat) is reported from the Neoschwagerina simplex (B6) and Presumatrina
schellwieni (B7) Zones of the Ratburi Limestone in the Khao Phrong Phrab area, Sara
Buri, central Thailand, where Neofusulinella
praecursor Deprat, Yangchenia haydeni
Thompson, Parafllsulina granumavenae(Roemer), Chusenella tenuis Toriyama andKanmera, Armenina sphaera (Ozawa), Verbeekina verbeeki (Geinitz), Pseudodoliolina
ozawai Yabe and Hanzawa, Metadoliolina
pinguis Toriyama and Kanmera, Neoschwagerina simplex Ozawa, Presumatrina
schellwieni (Deprat), and Neothailandina
komalmjuni Toriyama and Kanmera areassociated (Toriyama, 1975).
This species can be distinguished fromPresllmatrina schellwieni (Deprat) in having
a larger shell and smaller form ratio, andfrom P. rossica and P. primitiva, both described by Miklukho-Maklay (1957) from the
Crimea, in having a larger shell.
Material.-Axial section; IGUT-KU0840.

Acknowledgmen ts
This study is a part of the project "Biostratigraphy and palaeogeography of theuppermost Permian and the Lower Triassic in
the southern realm of the Asian Tethys" (Project no. 6304199, Leader: Prof. Juichi
Yanagida, Kyushu University), which was
financed by the Overseas Scientific Research
Fund from the Ministry of Education, Science and Culture of Japan. We would liketo thank Mr. Nikorn Nakornsri, Senior Geologist of Geological Survey Division, Department of Mineral Resources, Thailand for hiskind guidance in the field, and to Prof.
Juichi Yanagida and Dr. Takeshi Ishibashi of

Kyushu University, Dr. Koichi Nagai of
Ryukyu University, Dr. Tetsuo Sugiyama of
Fukuoka University, Dr. Kazuyuki Matsusueof Oyo Corporation (Oyo Chishitsu Co.
Ltd.), Ms. Rucha Ingavat-Helmcke and Yaowalak Chaimanee of Geological Survey
Division, Department of Mineral Resources,
Thailand for their kind cooperation in thefield. Thanks are extended to Prof. Hisayoshi Igo of the University of Tsukuba for his
useful suggestions on the fusulinacean taxonomy and critical reading of the manuscript.

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fusulinids from the Kongkashan pass of Xinjiang. Acta Micropalaeont. Sinica, vo!. 5, no. 4,
p.367-378, pis. 1-4. (in Chinese with English

abstract)Sun, X.F., Gao, C.X. and Zhang, L.X., 1983:
Fusulinida. In, Xi'an Inst. Gcol. Mineral

290

Katsumi Ueno and Sumio Sakagami
Resour. ed., Paleontological Atlas of Northwest

China, Shaanxi, Gansu and Ningxia Vol., Part
2 (Upper Paleozoic), p.3-40, pis. 1-10. Geol.
Publ. House, Beijing. (in Chinese)
Toriyama, R., 1958: Geology of Akiyoshi, Part III,
Fusulinids of Akiyoshi. Mem. Fac. Sci., Kyushu Univ., ser. D, vol. 7, p. 1-264, pis. 1-48.
- - , 1967: The Rat Buri Limestone of Thailand.
Jour. Geogr., vol. 76, no. 4, p. 13-20. (inJapanese with English abstract)- - , 1975: Permian fusulines from the Rat Buri
Limestone in the Khao Phrong Phrab area, Sara
Buri, central Thailand (Fusuline fossils fromThailand, Part IX). Mem. Fac. Sci., Kyushu
Univ., ser. D, vol. 23, no. I, p. 1-116, pis. 1-2l.
- - and Kanmera, K., 1977: The Permian fusulines
from the limestone conglomerate formation in the
Khao Phlong Phrab area, Sara Buri, central
Thailand. Geol. Palaeont. Southeast Asia, vol.
18, p. 1-27, pis. 1-3.
- - , - - and Ingavat, R., 1969: Neofusulinella
from Thailand (Fusulinacean fossils fromThailand, Part V). Ibid., vol. 7, p. 15-32, pis. 34.
- - and Pitakpaivan, K., 1973: Middle Permian
fusulines from Wat Kirinakratanaram, central
Thailand. Ibid., vol. 12, p. 43-61, pis. 5-6.
Ueno, K., 1992: Verbeekinid and neoschwagerinid
fusulinaceans from the Akiyoshi LimestoneGroup above the Parafusulina kaerimizensis
Zone, southwest Japan. Trans. Proc. Palaeont.
Soc. Japan, N.S., no. 165, p. 1040-1069.
- - and Igo, H., 1993: Upper Carboniferous foraminifers from Ban Na Din Dam, Changwat
Loei, northeastern Thailand. Ibid., no. 171, p.
213-228.
Vachard, D., 1990: Fusulinoids, smaller foraminifera
and pseudo-algae from southeastern Kelantan(Malaysia) and their biostratigraphic andpaleogeographic value. In, Fontaine, H. ed.,Ten years of CCOP research on the preTertiary of East Asia. CCOP Tech. Bull., vol.
20, p. 143-167, pis. 1-9.
- - and Miconnet, P., 1989: Une association 11
fusulinoides du Murghabien superieur au Monte
Facito (Apennin meridional, Italie). Rev. Micropa!eont., vol. 32, no. 4, p. 297-318, pis. 1-4.
Vimuktanandana, S., 1988: Geological map of Chan-gwat Loei (scale I: 250,000). Geological Survey Division, Department of Mineral Resources,
Thailand.
Wang, J.H. and Tang, Y., 1986: Fusulinids from
Chihsia Formation of Lengwu district in Tonglu
County, Zhejiang. Acta Micropalaeont. Sinica,
vol. 3, no. I, p. 3-11, pis. 1-3. (in Chinese withEnglish abstract)
Wang, Y.H., Wang, L.L., Wang, J.H., Zhu, Z.G., Lin,
G.W., Zhang, L.X. and Quian, Q., 1982:
Fusulinida. In, Nanjing Inst. Geol. Mineral
Resour. chiefly ed., Paleontological Atlas of
East China, no. 2 (Late Paleozoic), p. 5-108, pis.
1-30. Geol. Publ. House, Beijing. (in Chinese)Wei, J.Y., Xiao, W.M., Wang, H.G., Dong, W.L.,
Feng, R.L., Wu, X.G. and Zheng, S.F., 1987:
Carboniferous and Permian stratigraphy in Guizhou. Guidebook of 11th Intern. Congr. Carbo
Strat. and Paleont., Beijing, Excursion 5, p. 169, pis. 1-10. Printing House of Jiangsu Academy of Agricultural Sciences, Nanjing.
Xie, S.G., 1982: Protozoa. In, Geological Bureau of
Hunan ed., The Palaeontological Atlas of
Hunan, Geol. Mem., ser. Z no. 1, p. 2-73, pis. 135. Geol. Publ. House, Beijing. (in Chinese)
Yang, Z.D., 1985: Restudy of fusulinids from the"Maokou Limestone" (Permian) at Datieguan,
Langdai, Guizhou. Acta Micropalaeont. Sinica,
vol. 2, no. 4, p.307-335, pis. 1-3. (in Chinesewith English abstract and description of new
genus)
Zhang, L.X. and Wang, Y.J., 1974: Permian
Fusulinida. In, Nanjing Inst. Geol. Palaeont.

ed., Stratigraphical and Paleontological Atlasof West South China, p.289-296, pis. 150-153.
Sci. Publ. House, Beijing. (in Chinese)Zhou, J.P. and Zhang, L.X., 1984: Fusulinids fromthe Chihsia Formation of Mt. Qixiashan, Nanjingo Acta Palaeont. Sinica, vol. 23, no. 6, p.
716-724, pis. 1-3. (in Chinese with English

abstract)
Zhu, T., 1990: Fusulinacea. In, 121 Coal Geological Prospecting Party of Fujian and Zhu, T. eds.,

The Permian coal-bearing strata andpalaeobiocoenosis of Fujian, p. 40-59, pis. 1-7.
Geol. Publ. House, Beijing. (in Chinese withEnglish abstract)

Akiyoshi f~E, Guizhou (Kueichow) jHI'I, Maokou;t; P, Qixia (Chihsia) fe§ii, Shanxi(Sensi) ~Jm, Zhejiang i'4IfiI.

961.

Middle Permian foraminifers from NE Thailand:91 ~~t:~um. D.I1 ~T AA '71 :9--+j-y- ~ t-tf.J' G~t±:f L- t:9"W3"')"'A*21r1LEE.~:!J 1 oo~t}IHf!l, p =1 ~O)TLA '71!J --tj-y - I- t1J1It3n:."t±l-r-9~f5/f*ljjEw(:EiI'G

Verbeekina verbeeki (Geinitz), Pseudodoliolina cr. pseudolepida (Deprat), Presumalrina cr.
neoschwagerinoides (Deprat) ~"6-tr1'iJL.'RM:*iI;ifit.:: f::'~ ":lil' 0 te.. .:. 0) 1'iJL.'RM:*ItJ:~0);fYj~.'R ~"6- Iv "('I,,' 9 .:. til' G,0) lli'j~atpJtll""' Iv L *c.O)§lJJtIlL -Iv il t." Y :/ t ~:t
G.n9. ,:,.nlt p =1 :t(g~1U~vt9, 7 :r./V I:::'q.- TM:to J: tJ'.t-:t":/ =>. '7 'T ~ TM;fYj~.'R.:c.~~.t-r-9tpJtll"",NLe.JtIlO)1'iR.'RM.O)W~~0).~~~9.
J:!ffIl9J~·JNJ:m~291

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 172, pp. 292-310,5 Figs., December 30, 1993

962.

LATE CRETACEOUS CRABS FROM JAPAN*
JOE S.H. COLLINS
8 Shaw's Cottages, Perry Rise, Forest Hill,
London SE23 2QN, England

Y ASUMITSU KANIE
Yokosuka City Museum, Yokosuka, 238
and

HIROAKI KARASA W A
Department of Earth and Planetary Sciences, School of Science,
Nagoya University, Chikusa, Nagoya, 464-01

Abstract. Seven species in six genera of crabs are recorded from the Upper
CretaceousYezo Supergroup of Hokkaido and Izumi Group of Osaka and Hyogo Prefectures, Japan. The following three species are new: Metahomola brevis. Diaulax yokoii.
Eodorippe binodosa. The existence of Archaeopus shows that the Japanese Late Cretaceous
fauna is closely related to the fauna from the Pacific slope of North America.
Key words.
Izumi Group.

Crustacea, Decapoda, Brachyura, Upper Cretaceous, Yezo Supergroup,

1983, P/agi%phus ezoensis Nagao, 1941, andPithonoton injlatum Collins and Karasawa,
1993.
In this paper, we describe three new species,
Metahomo/a brevis. Diau/ax yokoii andEodorippe binodosa. from the Upper
Cretaceous Yezo Supergroup of Hokkaido
and new descriptions are given of Notopocorystes (Notopocorystes) japonicus (Jimb6,
1894) and A rchaeopus ezoensis (N agao, 1941)
from the Upper Cretaceous deposits of Japan.
In addition, the palaeobiogeography of theJapanese Late Cretaceous decapod fauna is
discussed.
Specimens used in the paper are housed in
the Yokosuka City Museum (YCM), Kanagawa Prefecture; Mizunami Fossil Museum(MFM), Gifu Prefecture; Mikasa City
Museum (MCM), Hokkaido; Toyohashi

Introduction
Since Jimb6 (1894) described EucOlystes
japonicus, the first crab known from theCretaceous of Hokkaido, a further eight
decapod species in eight genera have been
made known through studies of Nagao (1931 ;1932; 1941), Takeda and Fujiyama (1983),
and Collins and Karasawa (1993), these are
Callianassa ezoensis Nagao, 1932, Linuparusjaponicus Nagao, 1931, Paraga/athea miyakoensis Takeda and Fujiyama, 1983,
Hom%psis hachiyai Takeda and Fujiyama,
1983, NotopocOlystes (Eucorystes) intermedius
Nagao,
1931,
Necrocarcinus
undecimtubercu1atus Takeda and Fujiyama,"Received July 28, 1993; accepted September 20,
1993
292

962.

Late Cretaceous crabs from Japan

Museum of Natural History (TMNH), Aichi
Prefecture; Department of Earth and Planetary Sciences, School of Science, Nagoya
University (ESN), Aichi Prefecture; Department of Geology and Mineralogy, Faculty of
Sciences, Hokkaido University (UHR), Hokkaido; The Univeisity Museum, the University of Tokyo (MA).

1400E

293

142'E

144°E

YEZ-I
YEZ-2

44°N ---+----=:R-um-o~i-J..

(
YEZ-I0"--e

Hokkaido

Sap~oro ~ YEZ-U

Acknowledgments------=¢YEZ-I}, ~3
YEZ-14 I/

We thank Messrs. Masatoshi Kera, Yasuji
Kera and Yukinobu Yoshikawa (Hokkaido),
Mr. Mototaka Hayashi (Tokyo Metropolis),
Messrs. Takayuki Yokoi, Shinji Morita,
Yoshiaki Mizuno and Kiichiro Hachiya(Aichi Prefecture), Mr. lunichi Miyamoto(Hyogo Prefecture), and Mr. Susumu Sato
(Kagawa Prefecture) for allowing access to
specimens in their collections. We also
thank Mr. S.F. Morris, formerly of the
Department of Palaeontology, the Natural
History Museum, London, who offered much
valuable advice and Dr. Francisco 1. Vega ofInstituto de Geologia, Universidad Nacional
Autonoma de Mexico for sending us several
papers on fossil crabs.

Localities
1.

Hokkaido (Fig. 1).
Locality YEZ- I : Kitazawa, Pinneshiridake, Nakatonbetsu-cho, Esashi-gun, Hokkaido. Pebble nodule from the U3 Formation of the Upper Yezo Group (Santonian by
Igi, 1959). Notopocorystes (N.) japonicus
collected by Mizuno.
Locality YEZ-2: Kasekizawa, Nakagawacho, Nakagawa-gun, Hokkaido. Calcareous
nodule from the Nigorisawa Formation of theUpper Yezo Group (Santonian; Inoceramus
amakusensis Zone by Osanai et al., 1960).
Diaulax yokoii sp. nov. collected by Yokoi.
Locality Y EZ-3: Sankebetsugawa,
Haboro-cho, Tomamae-gun, Hokkaido.
Pebble nodule from the Upper Yezo Group(Santonian; Inoceramus amakusensis Zone,Urakawa

42'N

50km
~

1300E45'Nf1400.7
30 Km

I Sea of Japan

Osaka

~~o

®,

Awaji Island Izumisano

35'N

~I

. Osaka _""'-.---1!suit~'~--IZM-3

Yo I t:~\ZM-4
IZM-I,2 135°E.1Figure 1. Index map showing theCretaceous decapod localities from Japan.

Lateby Toshimitsu, 1988). Notopocorystes (N.)
japonicus collected by Hachiya.
Locality YEZ-4: Sakasagawa, Kamihaboro, Haboro-cho, Tomamae-gun, Hokkaido. Pebble nodule from the Upper Yezo
Group (Santonian; Inoceramus amakusensis zone by Toshimitsu, 1988). Metahomola
brevis sp. nov. collected by Mizuno.
Locality YEZ-5: Horotatezawa, Haborocho, Tomamae-gun, Hokkaido. Pebble nodule from the Middle Haborogawa Formationof the Upper Yezo Group (Santonian;
Inoceramus amakusensis Zone by Toshimitsu, 1988). Metahomola brevis sp. nov. collected by Hachiya.
Locality YEZ-6: Detofutamatazawa,
Kamihaboro, Haboro-cho, Tomamae-gun,
Hokkaido. Nodule from the Upper Yezo
Group (Santonian; Inoceramus amakusensis Zone by Ueda et al., 1961). Notopocorystes (Eucorystes) intermedius collected by
Hachiya.

294

Joe s.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

Locality YEZ-7: Junction of the Kanajirisawa and the Obirashibe-gawa, Tappu,
Obira-cho, Rumoi-gun, Hokkaido. Pebble
nodule from the Ml? Member of the Middle
Yezo Group (Turonian; Inoceramus hobetsuensis Zone by Tanaka, 1963). Metahomola brevis, Notopocorystes (N.) japonicus,
N. (Eucorystes) intermedius, Archaeopus
ezoensis collected by Hayashi.
Locality YEZ-8: Satoh-zawa, Kamikinenbetsu-gawa, Tappu, Obira-cho, Rumoigun, Hokkaido. Pebble nodule from the Mj
Member of the Middle Yezo Group (Turonian; Inoceramus hobetsuensis Zone by
Tanaka, 1963). Notopocorystes (Eucorystes)intermedius collected by Mizuno.
Locality YEZ-9: Kiritachi, Tomamae-cho,Tomamae-gun, Hokkaido. Pebble nodulefrom the Upper Yezo Group (Santonian;
Inoceramus amakusensis Zone). Archaeopus ezoensis collected by Mizuno.
Locality YEZ-lO: Katsurazawa, Ikushunbetsu, Mikasa City, Hokkaido. Nodule ofthe Mikasa Formation of the Middle Yezo
Group (Cenomanian; Inoceramus
nipponicus-I. pennatulus Zone by Matsumoto,1965). Eodorippe binodosa sp. nov. collectedby Yokoi.
Locality YEZ-l1 : Kumaoizawa, Ikushunbetsu, Mikasa City, Hokkaido. Pebble nodule from the lower part of the Upper Yezo
Group (Santonian; Inoceramus amakusensis Zone by Matsumoto, 1965). Diaulax
yokoii sp. nov., Notopocorystes (N.) japonicus
collected by Kera.
Locality YEZ-12: Hakkinzawa, Oyubari,
Yubari City, Hokkaido. Pebble nodulefrom the upper part of the Middle Yezo
Group (Cenomanian-Middle Turonian by
Hirano et al., 1977). Notopocorystes(Eucorystes) intermedius collected by Kera.
Locality YEZ-13: Kaneobetsu, Oyubari,
Yubari City, Hokkaido. Pebble nodulefrom the upper part of the Middle Yezo
Group (lower Cenomanian-middle Turonianby Iizawa et al., 1980; middle Turonian by
Matsumoto and Suekane, 1987). Notopoco-rystes (N.) japonicus collected by Kera.
Locality YEZ-14: Sanushibe, Hobetsucho, Yufutsu-gun, Hokkaido. Pebble nodule from the upper part of the Upper Yezo
Group (Santonian; Inoceramus amakusensis Zone by Kito et al., 1986).
Notopocorystes (N.) japonicus collected by
Kera.
Locality YEZ-15: Ikandai, Urakawa-cho,
Urakawa-gun, Hokkaido. Pebble nodulefrom the U 1 Member of the Upper Yezo
Group (Santonian; Inoceramus amakusensis Zone by Kanie, 1966). Notopocorystes(N.) japonicus collected by Yoshikawa.
2. Hyogo and Osaka Prefectures (Fig. I).
Locality IZM-I : N adachino, N antan-cho,
Mihara-gun, Hyogo Prefecture. Nodulefrom the Shimonada Formation, Izumi
Group (Lower Maastrichtian; Pachydiscus
aif. subcompressus Zone by Morozumi, 1985).
Archaeopus ezoensis collected by Sato andMiyamoto.
Locality IZM-2: Okawa, Nantan-cho,
Mihara-gun, Hyogo Prefecture. Pebbly conglomerate from the Shimonada Formation,
Izumi Group (lower Maastrichtian; Pachydiscus aff. subcompressus Zone by Morozumi, 1985). A rchaeopus ezoensis collectedby Sato and Miyamoto.
Locality IZM-3: Takinoike, Izumisano
City, Osaka Prefecture. Nodule from the
Matsuo Formation from the Izumi Group(Campanian/Maastrichtian by Matsumotoand Morozumi, 1980). Archaeopus ezoensis
collected by Miyamoto.
Locality IZM-4: Hakosaku, Han-nan-cho,
Sen-nan-gun, Osaka Prefecture. Nodulefrom the Shindachi Formation of the Izumi
Group (Maastrichtian by Kase, 1990). Archaeopus ezoensis collected by Miyamoto.

962.

Late Cretaceous crabs from Japan

Systematic descriptions
Section Podotremata Guinot, 1977
Subsection Dromiacea de Haan, 1833
Superfamily Homolodoromioidea
Alcock, 1899
Family Prosopidae von Meyer, 1860
Subfamily Pithonotinae Glaessner, 1933
Genus Pithonoton von Meyer, 1842

Type species.-By subsequent designationof Beurlen, 1928; Pithonoton marginatum
von Meyer, 1842.
Geologic range.-Middle Jurassic-Late
Cretaceous.
Pithonoton inflatum Collins andKarasawa, 1993
Figures 2-la-c

Pithonoton infiatum Collins and Karasawa, 1993, p.
17, figs. la-c.

Material.- TMNHOl530 (holotype).
Diagnosis.-A Pithonoton with a rounded
I?entagonal carapace, a strongly inflated dorsal surface, an obsolete median course of thecervical furrow and a low ridge on a poorly
delimited cardiac region (Collins and Karasawa, 1993).
Occurrence.-Hokkaido (detail of locality
is uncertain). Middle or Upper Yezo
Group.
Geologic range.-? CenomanianCampanian.
Superfamily Dromioidea de Haan, 1833
Family Diaulacidae Wright and Collins, 1972
Genus Diaulax Bell, 1863

Type species.-By monotypy Diaulax carteriana Bell, 1863; = Platypodia oweni Bell,
1850.
Geologic range.-Albian-Campanian.

295

Diaulax yokoii sp. nov.
Figures 2-4,5a-c

Material.-MFM247,001 (holotype), MCM
A373 (paratype).
Type locality.-Loc. YEZ-2 (Nigorisawa
Formation, Upper Yezo Group; Santonian).
Derivation of name.-From Mr. T. Yokoi
who collected the type specimen.
Diagnosis.-Carapace hexagonal in outline, with regions more or less distinct and awell defined cervical furrow.
Description.- The carapace is hexagonal in
outline, a little broader than long, widest
immediately before the cervical notch about
one-third distant from the front. In longitudinal section, the carapace is strongly downturned from the point of greatest carapace
width and the hind part is flat. In transverse
section, the part in front of the cervical furrow is flatly arched medially and behind that
it is nearly flat. The orbitofrontal margin
occupies about four-fifths of the total width.
The front is down turned ; the rostrum is
obscured by matrix, but appears to be weakly
sulcate, the narrowly raised margins leading
back to the inclined upper orbital margin,terminating in a broadly triangular outerorbital spine. The upper orbital margin is
irregularly beaded and there is a scattering offine granules anteriorly on the dorsal surface.
The orbits are broadly ovate. The lateral
margins are distinct with a thickened edge
continuing as far as the base of the cardiac
region. The anterolateral margins are shortwith a weak spine nearer to the outer orbital
spine than to a stronger one before the cervical notch. There is a similarly strong spine
behind the notch and the posterolateral margins converge to the posterior margin which isnarrower than the orbitofrontal margin andbounded by a thin ridge. A minute node
either side of the midline interrupts the cervical furrow which is broadly V-shaped, deep
and distinct to the outer angles of themesogastric lobe where it broadens and, passing round the proto gastric lobe in a shallow

296

Joe SH. Collins, Yasumitsu Kanie and Hiroaki Karasawa

Figure 2. la- c. PilhonolOn infiatum Collins and Karasawa, 1993, X 2.8. a, dorsal ; b, lateral; c,fron tal view of carapace. 2, 3. Melahomola brevis sp. nov. , dorsal view of carapace, X 3.3. 2, YCMGP854 (holotype), Loc. YEZ-7. 3, MFM247 , 002 (paratype), Loc. YEZ-4. 4, Sa - c. Diaulax yokoii sp.
nov. x3.5 . 4, MCM A373 (paratype) , Loc. YEZ- II , lateral view of right cheliped . 5, MFM247,001(ho lotype) , Loc. YEZ-2, a, dorsa l ; b, fronta l ; c, lateral view. 6. Eodorippe binodosa sp. nov.,
MFM247,003 (holotype), Loc. YEZ-IO, X 3.5 , dorsal view of carapace.

962.

Late Cretaceous crabs from Japan

curve, runs almost transversely to the margin.
A thin furrow branching from the cervicalruns obliquely back and defines a small triangular epibranchial lobe. The otherwise
indistinct branchiocardiac furrow is bounded
laterally by a low ridge which continues
obliquely along the side of the carapace.
The regions are mostly well defined: a weak
depression separates the hepatic region fromthe proto gastric lobes; there is a weak nodeon the anterior mesogastric process a little
anterior to rather more prominent frontal
lobes and a rounded ridge extends almost to
the base of the mesogastric lobe. The
urogastric lobe is divided into lozenge-shaped
portions by a median furrow and laterally
confluent with reniform meso branchial lobes.
A transverse groove separates the urogastricfrom the pentagonal cardiac region which hasa weak median depression anteriorly. Thereis a node on each protogastric lobe opposite
the cervical notch and a granule at the base of
the cardiac region.
The propodus of the cheliped is trapezoidal
in lateral view, with the basal margin about
one-third longer than the dorsal and interdigital margins. There is a wide convexity inthe basal margin at the base of the stout,
moderately deflexed, slightly inturned fixed
finger. The interdigital margin is nearlystraight and bounded by a shallow groove
and a wide, deep groove bounds the carpal
articulation which is rather more oblique inthe left chela, the right and left propodi havethe finely granulated lateral surface smoothly
rounded and have an incipient furrow extending parallel to the dorsal margin, but it iscloser to the midline in the right chela. The
slender dactyl us is about as long as the fixed
finger. The carpus is a little shorter than the
propodus, rounded triangular in section andthe strongly convex lateral surface is smoothwith a median furrow.
Discussion.- The carapace of Diaulax
oweni (Bell, 1850) (lower Albian-lower
Cenomanian) of England is rather more pentagonal in outline and may be further distin-297

guished from D. yokoii sp. nov. in having
poorly differentiated regions and a less conspicuous cervical furrow. The dorsal regions
may be vaguely tumid in some specimens ofD. jeliceps Wright and Collins, 1972 from the
Gault (Albian) of England and its carapace ismore or less similar in outline to D. yokoii sp.
nov., but it lacks the anterolateral spine andan epibranchial boss replaces the spine
behind the cervical notch; the cervical furrow of D. jeliceps is weaker and more sinuous
in its course to the margin. The carapace
length of Dialilax roddai Bishop, 1983a fromthe Glen Rose Limestone (Albian) of Texas
and Diaulax millerae Bishop, 1992 from theMerchantville Formation (Campanian) of
Delaware exceeds the breadth and this character serves immediately to distinguish thosespecies from D. yokoii sp. nov. The medianfurrow on the carpus is common to both D.
oweni and D. jeliceps, but only D. jeliceps hasa groove on the propodus, that of D. oweni
being ornamented by longitudinal rows of
granules continued onto the carpus.
Occllrrence.-Loc. YEZ-2, 11.

Geologic range.-Santonian.
Subsection Archaeobrachyura Guinot, 1977
Superfamily Homoloidea de Haan, 1839
Family Homolidae de Haan, 1839
Genus Metahomola Collins andRasm ussen, 1992

Type species.-By original designation ofCollins and Rasmussen, 1992; Homolopsis
punctata Rathbun, 1917.
Geologic range.-Late Cretaceous.
Metahomola brevis sp. nov.
Figures 2-2,3Material.- YCM-GP854 (holotype),
MFM247,002 (paratype).
Type locality.-Loc. YEZ-7 (Middle Yezo
Group; Turonian).
Derivation oj name.-Referring the rela-

298

Joe S.H. Collins, Yasumitsll Kanie and Hiroaki Karasawa

tively short metabranchial ridges.
Diagnosis.-A Metahomola with shortmetabranchial ridges, three cardiac tubercles
and intestinal nodes on either side of the
midline close to the cardiac region.
Description.- The carapace is subquadrate
in outline, about one-fourth longer than wide
between the linea homolicae at about midcarapace length; in longitudinal section it
rises steeply from the front, becomesdepressed at mid-length, rises and curves
gently to the posterior margin, being moderately arched transversely. Convex anterolateral margins are weakly constricted before
the orbital angle. The orbits are oblique andoccupy half of the carapace width. The rostrum is produced, narrowly triangular andcontinues the downward curve of the carapace; its raised sides lead back to thinupper orbital margins. Almost straight posterolateral margins without spines lead to
sharp posterior angles; the posterior margin
is about as wide as the front and bounded bya fine ridge. Crossing the carapace about
mid-length, the prominent cervical furrow isalmost straight at the base of the mesogastriclobe, turns before the lobe's outer angle and,curving gently round the hepatic region, runs
forward and outward to the margin.
Equally prominent branchiocardiac furrows
run more or less parallel to the cervical andterminate in a deep depression behind ovate,rather tumidmesobranchial lobes. The tipof the anterior mesogastric process extends
between low epigastric lobes and a small
tubercle about mid-length has a sulcus justbehind it. Of the tubercles on the proto gastric lobes, the inner pair flank that on themesogastric process, another lays slightly forward and outward while a third occupies thespace between the hepatic and mesogastriclobes. There is a median tubercle and twobasal ones on the mesogastric lobe and twoon the clearly delimited hepatic region which
is subcircular in outline. The narrowurogastric lobe has a transverse pair of small
nodes. The shield-shaped cardiac region has
three small tubercles in an inverted triangle
and opposite its widest part a short, narrowridge extends onto the metabranchial region.
Each mesobranchial region has one tubercle
behind the cervical furrow and a smaller one
near the urogastric lobe. Flattened, circular
intestinal nodes lay close to the midline oneither side of the cardiac region.
Discllssion.-Metahomola was proposed by
Collins and Rasmussen, 1992 to containhomo lids with an entire rostrum and a ridge
extending from the widest part of the cardiac
region onto the metabranchial lobe. A second genus, Eohomola Collins and Rasmussen, 1992, was described to contain species
also with a metabranchial ridge, but further
distinguished by the presence of a bifurcate
rostrum, thereby leaving those species with anentire rostrum and lacking a metabranchial
ridge in Hom%psis Bell, 1863.
Absence of a metabranchial ridge andjuxtaposition of tubercular ornament immediately distinguishes Homolopsis hachiyai Takeda and Fujiyama, 1983, from the Miyako
Group (lower Aptian) of Iwate Prefecture,from Metahomo/a brevis sp. nov. The
urogastric lobe of M. pllnctata is divided at
the midline, there are no nodes on the cardiac
region and the intestinal nodes are set further
apart. Metahomola gibbosa (SchWter, 1879)
from the Senonian of Germany lacks the
tubercle at the median base of the proto gastric lobe and the metabranchial ridges extend
further towards the lateral margin.
M. brevis sp. nov. closely resembles the~
Figure 3. 1-6. Notopocorystes (Notopocorystes) japonicus (Jimb6, 1894). 1, MCM A374, Loc.
YEZ-14, X 1.5, dorsal view of carapace. 2a-c, ESN83007P, Loc. YEZ-3, X 2.5, a, dorsal; b, lateral; c,frontal view of carapace. 3, YCM-GP856, Loc. YEZ-7, X 2.5, dorsal view of carapace. 4, ESN83008P,
Loc. YEZ-I, X 1.5, dorsal view of carapace. Sa-c, YCM-GP849, Loc. YEZ-8, X 1.5, a, dorsal; b, lateral;c, frontal view of carapace. 6a-c, YCM-GP850, Loc. YEZ-8, X 2.5, a, dorsal; b, lateral; c, frontal viewof carapace.

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Late Cretaceous crabs from Japan

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Joe S.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

Middle Eocene species Prohomola japonica(Yokoyama, 1911) from the Manda Group of
Fukuoka Prefecture (Karasawa, 1992) andcould well be ancestral to it. P. japonica
differs largely in having a flatter carapace
with rather more rounded lateral margins;
the rostrum is well developed, directedstraight forward and there is a pair of large
postrostral spines.
Occurrence.-Loc. YEZ-4, 5, 7,Geologic range.- Turonian-Santonian.
Superfamily Raninoidea de Haan, 1841
Family Raninidae de Haan, 1841
Genus Notopocorystes McCoy, 1849

Type species.-Subsequently designated byWithers (1928), Notopocorystes mantelli
McCoy, 1849 = Corystes stokesii Mantell,1844.
Geologic range.-Early Albian-Santonian.
Subgenus Notopocorystes McCoy, 1849

Notopocorystes (Notopocorystes)japonicus (JimbO, 1894)
Figures 3-1-6

Eucorystes japonicus Jimbo, 1894, p. 101, pI. 9, fig. 7.
Notopocorystes? japonicus (Jimbo). Glaessner,
1929, p. 277.
Notopocorystes (Eucorystes) japonicus (Jimbo).
Nagao, 1931, p.207.
Notopocorystes japonicus (JimM). imaizumi, 1978, p.
32.

Material.-MA 7483 (GT.I-159) (Holotype).
YCM-GP849, 850, 852, 853, 856, 858; MCM
A372, A375; ESN83007P, 83008P (new material).
Diagnosis.-Carapace with a weakly devel-

oped median carina without tubercles on the
midline and protogastric and hepatic tubercles.
Description.- The carapace is truncatedoval in outline, moderately arched longitudinally, reaching its highest point about midlength where it becomes flattened before curving to the posterior margin, and moderately
arched transversely. The orbitofrontal width
is a little more than half the carapace width
measured between the cervical notches. The
rostrum projects slightly in front of theorbital spines, is widely bifurcate with two
blunt terminal spines and is shallowly tomoderately grooved; its sides are raised andlead to a small inner orbital spine. The
orbits are wide and spines separate two evenly
spaced fissures in the upper orbital margin.
Two forwardly directed slender anterolateralspines are slightly upturned and there is asmall one behind the cervical notch. The
posterolateral margins are sharp anteriorly,
becoming rounded and slightly concave tothe posterior margin. The cervical furrow isshallow and fairly broad; it runs forwardfrom the margin and curves back behind themesogastric lobe in a shallow curve. An
equally prominent groove, not quite uniting
with the cervical, delimits the epibranchiallobe. An incipient ridge borders the lateralparts of feeble branchiocardiac furrows whichbecome obsolete before the cardiac region.
A median carina starting as a low, broadridge on the urogastric lobe tapers on the
cardiac region and continues parallel to theposterior margin where it is flanked by two,
short ridges. The tip of the narrow anterior
mesogastric process extends into the base of
the rostrum and the feebly demarcatedmesogastric lobe is smooth. The weakly~
Figure 4. la, b, 2. Notopocorystes (Eucorystes) intermedius Nagao, 1931, x2.5. la, b, ESN83006P,
Loc. YEZ-6, a, dorsal; b, lateral view of carapace. 2, ESN83009P, Loc. YEZ-8, dorsal view of carapace.
3-7. Archaeopus ezoensis (Nagao, 1941). 3, YCM-GP855, Loc. YEZ-7, X 3.0, dorsal view of carapace.
4, ESN830IOP, Loc. IZM-4, X 1.8, male thoracic sterna. 5, ESN83011 P, Loc. YEZ-9, X 3.0, dorsal view ofcarapace. 6a-d, ESN83012P, Loc. IZM-2, X 1.8, a, frontal; b, dorsal; c, ventral; d, lateral view ofcarapace offemale. 7a-c, ESN830 13P, Loc. IZM-2, X 1.8, a, dorsal; b, ventral; c, lateral view of carapaceof male.

962.

Late Cretaceous crabs from Japan

30 1

302

Joe s.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

scabrous urogastric lobe is confluent with the
cardiac region which has a prominent transverse pair of tubercles. There is a tubercle
on the hepatic lobe and a row of minutetubercles from a crescentic ridge by the innerpart of the hepatic region and within the
curve is a larger tubercle on the proto gastriclobe.
Discussion.-Notopocorystes (N.) japonicus
lacks all trace of the 'strap' ornament of furrows and lobes which are diagnostic requirements for Eucorystes. Individuals of thisspecies reach among the largest known of thegenus. The longitudinal curvature becomes
more evenly rounded as growth advances andthe dorsal tubercles are much less in evidence.
Wright and Collins (1972) recognized aphylogenetic progression In the Albian
species Notopocorystes stokesii (Mantell,1844) and erected three subspecies of which
the nominate form, N. stokesii stokesii,
occupies the middle part of the progression.
The earliest known (Douvilleiceras mammillatum Zone) form, N. stokesii praecox Wrightand Collins, 1972 passes into N. stokesii
stokesii (Hoplites spa th i-Dipoloceras cristatum Subzone) which gives rise to N. stokesii serotinus Wright and Collins, 1972,
which was well established in the D. cristatum Subzone, where it exised alongside N.
stokesii stokesii and continued to the Cambridge Greensand (Albian). N. stokesii
praecox has an almost continuous carina,
notched into weak tubercles only just in frontof and just behind the cervical furrow, there isa single, rarely two, tubercle on the hepatic
region and the crescentic ridges and protogastric tubercles are indistinct; the orbitofrontal
width/carapace width ratio averages 58.
N. (N.) japonicus would appear to be aregressive form of N. stokesii praecox for not
only does it have a comparable orbitofrontal/
carapace width ratio of 56.5, but retains someof the characters of that species, while themedian carina is developed only on the branchial region. Characters not possessed by N.
stokesii praecox are the ridges at the base of

the median carina and the transverse pair ofcardiac tubercles, which is unique to thespecies discussed.
Occurrence.-Pankemoyubari River, a lefthand tributary of the Yubari, Ishikari Province; Scaphites beds (Turonian-Coniacian)(type locality). Loc. YEZ-I, 3, 7, 9, II, 13,
14, 15.
Geologic range.-Cenomanian-Santonian.
Subgenus Eucorystes Bell, 1863
Type species.-By monotypy Notopocorystes carteri McCoy, 1854.
Geologic range.-Early AlbianCampanian.
Notopocorystes (Eucorystes)intermedius Nagao, 1931
Figures 4-1,2

Notopocorystes (Eucorystes) intermedius Nagao,
1931, p.207, pI. 14, figs. 4, 4a.
Notopocorystes intermedius Nagao. Imaizumi, 1978,
p. 27, fig. 38.

Material.-UHR3184 (Nagao's type specimen). YCM-GP851, 857; MCM A374;
ESN83006P, 83009P (new material).
Diagnosis.-Carapace without a dorsal
median carina.
Discussion.- This species differs from
Notopocorystes (N.) japonicus (limbO, 1894)
in having the carapace relatively longer thanwide and the dorsal surface without themedian carination. Young individuals of N.
(N.) japonicus have tubercles on the protogastric and hepatic lobes, whereas the dorsal
surface of N. (E.) intermedius is marked only
by weak 'straps' on the gastric region.
Occurrence.-Southeast of Ikushunbetsu(Horonai) colliery and along the tributary of
Ikushunbetsu, Province of Ishikari, Hokkaido (Horonai, Mikasa City); Mikasa Formation, Middle Yezo Group (CenomanianTuronian) (type locality). Loc. YEZ-6, 7, 8,
12.
Geologic range.-Cenomanian-Santonian.

962.

Late Cretaceous crabs from Japan

Superfamily Cyclodorippoidea
Ortmann, 1892(= Superfamily Tymolidae Alcock, 1896)
Family Torynommatidae Glaessner, 1980
(Rectae ICZN, Art, 20 (4), (b) (i))
Genus Eodorippe Glaessner, 1980

Type species.-By monotypy Eodorippe
spedeni Glaessner, 1980.
Geologic range.-Late Cretaceous.
Eodorippe binodosa sp. nov.
Figure 2-6

Material.-MFM247,003 (holotype).
Type locality.-Loc. YEZ-IO (Mikasa Formation, Middle Yezo Group; Cenomanian).
Derivation of name.-With reference tothe node on each protogastric lobe.
Diagnosis. -Carapace su brectangular,slightly longer than wide; lateral margin
without hepatic spine, gently convergent
anteriorly; one node on each protogastriclobe.
Description.- The carapace excluding therostrum, is subrectangular in outline, slightly
longer than wide with the greatest width
across the mesobranchial and cardiac lobes in
the posterior third. The dorsal surface isgently convex in both longitudinal and transverse sections. The wide orbitofrontal margin occupies about 0.8 of the greatest carapace
width; a long, spatulate rostrum with a shallow median sulcus protrudes straight forwards and there is a small, broadly triangular
postrostral spine on either side. The upper
orbital margin is concave, slightly upturnedand bears a sharp outer orbital spine. The
lateral margins, with cervical and branchiocardiac notches, are slightly sinuous andconvergent anteriorly; they form a sharpedge in the anterior half and there is a basal
scar indicating a small spine before the cervical notch. The posterior margin is slightly
convex and bounded by a narrow rim. The
dorsal regions are reasonably well defined by

303

shallow, broad grooves. Large, tumid
protogastric lobes are shallowly separated
from the narrow anterior mesogastric process
and each has a node close to the small,depressed, triangular mesogastric lobe. The
cervical furrow, interrupted at the midline, isbroadly V-shaped medially and curves sharply forward and outward to reach the marginabout one-fourth distant from the front. The
outer course of the branchiocardiac furrow
runs almost parallel to the cervical andencloses a narrow epibranchial lobe, while afaint oblique groove defines a chordate
metabranchial lobe and turns back to form
deeper grooves bordering the urogastric lobe.
The dorsal surface is coarsely granulated,with the granules tending to form spinules atthe lateral margins.
Discussion.- The present new species
differs from Eodorippe spedeni Glaessner,1980 from the Campanian-Maastrichtian of
New Zealand in having a subrectangularcarapace, less obviously narrowing towards
the front. The intermediate oblique grooves
between the cervical and branchiocardiac
furrows are weaker than in E. spedeni and of
the three pairs of protogastric nodes developed in that species only the inner pair ispresent in E. binodosa sp. nov.
There appears to be a certain amount ofdamage about the rostrum of E. spedeni andthe postrostral spines are not in evidence.
Glaessner (1980) said that a significant
difference between Dorippe and Eodorippe
was the apparent absence of the prominent
infraorbital teeth of Dorippe and the shape ofthe rostrum. The postrostral spines noted in
E. binodosa sp. nov. would seem to foreshadow the infraorbital teeth of Dorippe andthe reduction of the rostrum of Eodorippe andfurther development of the postorbital spines
would undoubtedly produce a front in keeping with that of Dorippe.
Geologic range.-Cenomanian.

Joe s.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

304

Table 1. Stratigraphic and geographic distribution of the Late Cretaceous decapods ofJapan (compiled from Bishop, 1986, Collins and Karasawa, 1993, Collins and Rasmussen, 1992,
Feldmann et af., 1993, Glaessner, 1969; 1980; Nagao, 1931; 1932; 1941, Tshudy and Feldmann, 1988 and Wright and Collins, 1972). L, Linuparus; C, Callianassa; P, Pithonoton; D,
Diaufax; M, Metahomofa; E, Eodorippe; N,Notopocorystes s. I.; A, Archaeoplls.
North America
Geologic age

Japan

Madagascar& India

AntarcAustralia
tica

New
Zealand

Pacific
Slope

Western
Interior &Atlantic
Coastal
Plain

Greenland

EuropeLCPDMENA LCPDMENA LCPDMENA LCPDMENA LCPDMENA LCPDMENA LCPDMENA LCPDMENA LCPDMENA

MaastrichtianCampanian

gj~

Santonian

U

Coniacian

~

£

I

~

I

i

I

I

j

Turonian

Cenomanian~J

Early Cretaceous

Section Thoracotremata Guinot, 1977
Superfamily Ocypodoidea Rafinesque, 1815
Family Retroplumidae Gill, 1894
Genus Archaeopus Rathbun, 1908

Type species.-By monotypy Archaeopus
antennatus Rathbun, 1908.
Geologic range.-Early ?-Late Cretaceous.
A rchaeopus ezoensis (Nagao, 1941)
Figures 4-3--7; 5-1--5

P/agi%phlls ezoensis Nagao, 1941, p.97, pI. 26, figs.
I, 2; Imaizumi, 1978, p. 45.

Material.-Nagao's

type

specimen.

YCM-GP855; ESN83010-83018P (new material ).
Diagnosis.-Archaeopus with weak transverse proto gastric ridges.
Description.- The carapace is subquadrate
in outline, almost as long as wide, widest at
the posterior third, moderately arched intransverse section, vaulted and more steeply
downturned in front in longitudinal section.
Straight anterolateral margins converge anteriorly and the posterior margins round into aslightly concave posterior margin which is
thickened, bounded by a depression and,laterally, by a low ridge. The orbitofrontal
margin takes up about four-fifths of thegreatest carapace width, with ovate orbits
taking up the outer thirds. The front, exten-~ Figure 5. 1-5. Archaeoplls ezoensis (Nagao, 1941). 1, ESN83014P, Loc. IZM-4, X 1.8, dorsal
view of carapace. 2a, b ESN83015P, Loc. [ZM·3, x2.5, a, frontal; b, dorsal view of carapace. 3a-c,
ESN830I6P, Loc. IZM-I, X 1.8, a, dorsal; b, posterior; c, ventral view of carapace of female. 4a, b,
ESN83017P, X 1.5, a lateral view of chelipcds; b, dorsal view. 5, ESN83018P, X 1.5, dorsal view.

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Late Cretaceous crabs from Japan

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306

Joe s.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

ding a little beyond the upper orbital margin,
is rounded accuminate, its sides are thickened
and raised, a short median sulcus extends
back between small, elongate epigastric lobes.
The cervical furrow runs almost straight andoblique back from the margin to the mesogastric lobe before turning sharply back andpassing across the midline in a broad curve.
The branchiocardiac furrow is nearly transverse and separates a short urogastric lobe
from the heart-shaped cardiac region. Foursmall, evenly spaced tubercles line the margin
between the furrows. A prominent ridge
extending across the base of the protogastriclobes is interrupted by the tip of the mesogastric lobe; a second ridge normally associatedwith retroplumids is represented by a large
tubercle on each epibranchial lobe with asimilar one stepped inward and backward on
the metabranchial lobes in line with two
smaller nodes at the base of the mesogastriclobe, while a third ridge bound by thebranchiocardiac furrows is carried across the
midline and the median part limited to theanterior pair of three tubercles on the cardiac
region. There is a small tubercle on eachhepatic region and a narrow, lingulate lobe
runs between the cardiac and meso branchiallobes.
Evidently the transverse ridges are a featureof ontogeny, for on a carapace with a widthof 8.0 mm, only the hindmost of the ridges is
to any extent developed, although the metabranchial prominence is circular rather thanlinear; of the middle ridge, only the epi- andmesobranchial tubercles with the latter thelarger as in adults are present; and smooth,tumid protogastric lobes give no indication of
the ridge later developed.
The abdominal sternites are very wide andshallow; the 1st-2nd appear to be broadly
triangular in outline and have a shallow
median depression; The 3rd with a deep
circular median depression is separated fromthe 1st-2nd by a shallow groove; a groove
separates them from the 4th sternites which
are trapezoidal in outline with the latero-posterior angle partly embracing the 5th sternites; the 5th-7th pairs are subrectangular.
The 3rd sternites have a tumidity either sideof the median trough while on the 4th twolarger tumid areas line the trough and a low
ridge bounds the cheliped margin. The 5th
and 6th sternites each have a short ridge
opposite the articulation boss of the corresponding limbs.
The chelipeds are subequal with the rightslightly larger, extended their length would
equal about twice the carapace width; thedactylus, having finely serrated occludent
margins, is slender, de flexed and about half ofthe length; a row of pits lining a shallow
groove runs the length of the fixed finger; thepropodus height is equal to about half its
length. The carpus is about half the lengthof the propodus, its subquadrate lateral surface is weakly rounded and smooth. Theline of inclination of the 1st pereiopods to thechelipeds suggests the presence of a similar
gap between those limbs as discerned by
Rathbun (1908, p.348) on A. antennatus.
The 2nd-4th pereiopods are reduced in length
posteriorly, but the ischia retain much thesame proximal width and taper distally. The
proximal width of the 4th pair is about onethird as long as the ischium. The ischialbase of the 4th pereiopods extends to, orslightly beyond, the posterior angle of thecarapace, thus, the 5th pair of limbs, notpreserved, would probably occupy much thesame (subdorsal) position envisaged by Rathbun (1908) for those limbs of A. antennatus.
Discussion.- There is, in the general outlineof the carapace and juxtaposition of dorsal
characters considerable similarity between A.
ezoensis and A. vancouverensis (Woodward,1896) from the Campanian-Maastrichtian ofBritish Colombia and Vancouver Island.
The anterior transverse ridge of A. vancouverensis is stronger than that of A. ezoensis.
The carapace of A. antennatus Rathbun,
1908 from the Upper Cretaceous Chico Formation of California is comparatively wider,its mesobranchial lobe is larger and ovate in962.

Late Cretaceous crabs from Japanoutline, and the cardiac part of the posteriorridge is continuous rather than limited to apair of tubercles. The sternites of A. antennatus, seen in detail (Rathbun, 1908, pIs. 47,58), are essentially similar to those of A.
ezoensis except that in the former species thecircular pit occupying the Ist-3rd sternites ismore prominent in the male. The lateralangle is more attenuated on the 4th pairsternites and the tumid areas are linear, theone bordering the cheliped is reduced to a
sharp edge.
Via (1982, fig. 2) put forward an acceptable
scheme for the phylogeny and evol ution for
the Retroplumidae. Vega and Feldmann(1992) revised Via's scheme on the basis of thethen known species, Cristipluma mississippiensis Bishop, 1893b from the Maastrichtian
Coon Creek Formation of Union County,
Mississippi, Costacopluma mexicana Vega
and Perrilliat, 1989 from the upper Maastrichtian Potreillos Formation of Mexico,
Costacopluma bishopi Vega and Feldmann,
1992 from the Maastrichtian Mexcala F ormation of Mexico and the new locality recording
for Costacopluma concava Collins and Morris, 1975 from the Maastrichtian of northIndia (Gaetani et al., 1983). Two species,
Costacopluma sp. from the Upper Eocene(Priabonian) of Hungary (MUller and Collins, 1991) and Costacopluma binodosa Collins and Rasmussen, 1992 from the lower
Campanian of Greenland have since been
described. It appears that Costacopluma wasconfined to the Tethyan and that Archaeopus
except for the earliest member, Archaeopus
rathbunae Beurlen, 1965 from the Albian ofBrazil, is characteristic of the North Pacificseas.
Occurrence.-Sanushibezawa, Hobetsucho, Yufutsu-gun, Province of Iburi, Hokkaido; Hakobuchi Group (CampanianMaastrichtian) (type locality). Loc. YEZ-7,
9,IZM-I-4.
Geologic range.- Turonian-Maastrichtian.

307

Discussion
The Upper Cretaceous decapod fauna fromthe Yezo Supergroup of Hokkaido and theIzumi Group of Hyogo and Osaka Prefectures
is represented by nine species in eight genera.
Their stratigraphic range is shown in Table I.
Pithonoton, a predominantly Jurassic genus,
ranges into the Cretaceous; Diaulax, Metahomola, Eodorippe, Notopocorystes s.I., andArchaeopus are confined to the Cretaceous,while Callianassa and Linuparus have living
species.
Among eight genera, Linuparus, Callianassa and Notopocorystes are cosmopolitan;
Pithonoton, Diaulax and Metahomola arecharacteristic of the Tethyan realm, while
Archaeopus was recorded from the Upper
Cretaceous deposits of Japan and the Pacificslope of North America (Vancouver Island,
British Colombia, and California), and namely, it is characteristic of the North Pacificrealm. Considering the occurrences of Pithonoton, Diaulax and Metahomola, theJapanese decapod fauna seems to be related to
the Tethyan fauna from the Cenomanian to
the Santonian, whilst the existence of A rchaeopus shows that it is closely related to thefauna from the Pacific slope of North America. Eodorippe, on the other hand, is known
outside Japan only by its possible derivative,
Eodorippe spedeni Glaessner, 1980 from theCampanian/Maastrichtian of New Zealand.

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Joe S.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

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Animalium, quae in Itinere per Japoniam, Jussu
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Batava Imperium Tenent, Suscepto, Annis 18231830 Colleqit, Notis, Observationibus et A dumbrationibus Il!ustravit. p. ix-xvii, 1-24 (1833);
25-64 (1835); 65-72 (1937); 73-108 (1839);
109-164 (1841); i-xxxi, 165-243 (1849); viixvii (1850), pIs. A, B, 1-8 (1833); C, D, 9-15,17(1835); E. F, 16, 18-24 (1937); G, H, 25-32(1839); 33-37,39-42,47 (1841); I-N, 38,43-46,
48, 51-55 (1844); O-Q, 49-50 (1849).
Lugduni-Batavorum, Leiden.
Hirano, H., Matsumoto, M. and Tanabe, K., 1977:
Mid-Cretaceous stratigraphy of the Oyubari area,
central Hokkaido. Palaeont. Soc. Japan, Spec.
Papers, no. 21, p. 1-10.
Iizawa, K., Oki, M. and Suekane, T., 1980: Upper
Cretaceous fossil from eastern coast of Shuparocho, Yubari City, Hokkaido. Part 2. Tokai
Mem., no. 17, p. 347-368. (in Japanese)
Igi, S., 1959: Pinneshiri. 1/50,000. Explanatory textof the Geological Map of Japan. 41 p. Geological survey of Japan. (in Japanese with English
abstract)
Imaizumi, R., 1978: Fossil decapod crustaceans fromJapan, with special reference to the classificationsof Conchostracans and Malacostracans. Contr.,
Inst. Geol. Paleont., Tohoku Univ., vol. 78, 49 p.,
6 figs. (in Japanese)
Jimbo, K., 1894: Beitrage zur Kenntniss der Fauna
der Kreideformation von Hokkaido. Paliiont.
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Kanie, Y., 1966: The Cretaceous deposits in the
Urakawa district, Hokkaido. Jour. Geol. Soc.
Japan, vol. 72, no. 7, p. 315-328. (in Japanesewith English abstract)
Karasawa, H., 1992: Fossil decapod crustaceans from
the Manda Group (Middle Eocene), Kyushu,
Japan. Trans. Proc. Palaeont. Soc. Japan,
N.S., no. 167, p. 1247-1258.
Kase, T., 1990: Late Cretaceous gastropods from theIzumi Group of southwest Japan. Jour.
Paieonl., vol. 64, no. 4, p.563-578.
Kito, N., Kaiho, K., Takahashi, K. and Wada, N.,
1986: Geologic age of the plesiosaurian fossil
from Hobetsu-cho, Hokkaido, Japan. Bull.

962.
Hobetsu Mus.,
Japanese)no. 3,

p. I ~ 7,

Late Cretaceous crabs from Japanpis. I ~ 13.

(in

Mantell, G.A., 1844: Medals of creation. vol. 2, p. i~viii, 457~IOI6, London.
Matsumoto, T., 1965: A monograph of the Collignoniceratidae from Hokkaido. Mem. Fac.
Sci., Kyushu Univ., ser. D, vol. 16, no. I, p. 1~80,
pis. I ~ 18.
- - and Morozumi, Y., 1980: Late Cretaceousammonites from the Izumi Mountains, southwest
lapan. Bull. Osaka Mus., Nat. Hist., no. 33, p.
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- - and Suekane, T., 1987: Some acanthoceratid
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McCoy, F., 1854: On some new Cretaceous Crustacea. Ann. Mag. Nat. Hist., ser. 2, vol. 14, p.
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Meyer, H. von., 1842: Ueber die in dem dichten
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Morozumi, Y., 1985: Late Cretaceous (Campanianand Maastrichtian) ammonites from Awaji
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I, no. 2, p. 207~214, pI. 14.
- - , 1932: Two Tertiary and one Cretaceous Crustacea from Hokkaido, Japan. Ibid., vol. 2, no. I,
p. 15~22, pI. 4.
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tableau de l'univers et de corps organises.

224

p. Palermo.
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309- - , 1917: New species of South Dakota crabs.
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Seymour Island, Antarctica. Geol. Soc. America, Mem. 169, p. 291~301.
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Cretaceous deposits in the Chikubetsu area,
Hokkaido. Sci. Rep., Fac. Sci. Kyushu Univ.,
Geol., vol. 6, no. 3, p. 15~32. (in Japanese withEnglish abstract)
Vega, F.l. and Feldmann, R.M., 1992: Occurrence of
Costacopluma (Decapoda: Brachyura: Retroplumidae) in the Maastrichtian of southern
Mexico and its paleobiogeographic implications.
Ann. Carnegie Mus., vol. 61, no. 2, p. 133~152.
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de cangrejo del genero Costacopluma (Crustacea: Decapoda: Retroplumidae) del Maastrichtiano del estado de Nuevo Leon. Univ.
Na!. A uton. Mexico, Inst. Geologia, Revista, vol.
8, no. I, p. 84~87.
Via Boada, L., 1982: Nueva contribucion al estudiopaleontologico de la superfamilia Ocypodoidea(CrusUiceos Decapodos). Bo!. Geol. y. Miner.,
vol. 43, no. 2, p. 115~119.
Withers, T.H., 1928: New Cretaceous crabs fromEngland and Syria. Ann. Mag. Nat. Hist., ser.
10, vol. 2, p. 456~462, pI. 13.
Woodward, H., 1896: On some podophthalmatous
Crustacea from the Cretaceous formation of
Vancouver and Queen Charlotte Islands. Jour.
Geol. Soc. London, vol. 52, p. 221~228.
Wright, e.w. and Collins, J.S.H., 1972: British
Cretaceous Crabs. Palaeontological Society
Monographs, Publication no. 533, part of vol.
126 for 1972, 114 p., 22 pis. The Palaeontographical Society, London.
Yokoyama, M., 1911 : Some Tertiary fossils from theMiike Coalfield. Jour. Col!. Sci., Imp. Univ.
Tokyo, vol. 27, art. 20, p. 1~16, pis. 1~3.

Joe s.H. Collins, Yasumitsu Kanie and Hiroaki Karasawa

310

Kitazawa :fUR., Pinneshiridake l!tl:1l"-Lm-ffi, Nakatonbetsu OP~5.JU, Esashi tt¥, Kasekizawa
1tE/'R, Nakagawa opJI[, Sankebetsugawa :=:'=§5.JUJII, Haboro ::f]1\!lt, Tomamae 1illtr, Sakasagawa iJi!JiI, Kamihaboro J::.::f]I\!lt, Horotatezawa I\!lt.lz:/'R, Detofutamatazawa T r =~/'R,
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55ff(, Kaneobetsu 7J ;;f :t- -' '/, Sanushibe -Ij- 3( Y' -', Hobetsu %\Ii:5.JU. Yufutsu ~:tb, Ikandai
Urakawa MM, Nadachino .:ttk!ff, Nantan lWiik. Mihara :=:'1J]l:, Okawa :kJI[,
Takinoike l'I[/ i\!l,. lzumisano 7'RftiJlf. Hakosaku $jl1''f. Han-nan IlRlW. Sen-nan jRlW.

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Joe S.H. Collins • ~lImt:J't • WiiKEfj)§

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 172, pp. 311-327,5 Figs., December 30, 1993

963. MOLLUSCAN FAUNA FROM THE MIOCENEMESHIKUNI FORMATION ON REBUN ISLAND,NORTHERN HOKKAIDO*YOSHIKAZU NODA
Institute of Gelology and Paleontology, Faculty of Science,
Tohoku University, Sendai, 980
and

KOICHI HOY ANAGI
Department of Geology, Faculty of Science,
Shinshu University, Matsumoto, 390

Abstract. The Meshikuni Formation occurs in the northern part of Rebun Island andis composed of sandstone and siltstone in which many molluscan fossils occur. The
Meshikuni molluscan fauna consists of three assemblages; the first one is represented by
Mylillis tichanovitchi Makiyama, the second by "Crassostrea" gravitesta (Yokoyama) and
Glycymeris species, and the third by deep-water mollusks. Mytillis tichanovitchi is one of
the representative species of the Asahi fauna of central Hokkaido, and is reported from theLower to lower Middle Miocene strata of Sakhalin and Kamchatka as well as Japan. The
discovery of MytilliS tichanovitchi suggests that the Meshikuni Formation is correlative with
the Asahi Formation of central Hokkaido.
Key words.

Molluscan fauna, Neogene, My tilliS, Rebun Island, Hokkaido

correlated the Motochi Formation with the
Middle Miocene Chikubetsu Formation ofnorthwestern Hokkaido, and also correlated
the Meshikuni Formation (including the
Kabuka facies) with the Middle to Upper
Miocene Togeshita Formation of northwestern Hokkaido. They also suggested that theHamanaka Formation is correlative with theUpper Miocene Wakkanai stage on the basisof lithofacies characteristics of the so-called"black hard shale." Noda and Hoyanagi(1987) pointed out that the Meshikuni Formation may contain both Chikubetsu andTogeshita molluscan elements and that thepart of the Meshikuni yielding mytilid fossils
may be equivalent to the Motochi Formation,although detailed geological and paleontological studies were not done.

Introduction

Rebun Island, located off the northwest
coast of Hokkaido (Figure 1), consists mainlyof Mesozoic strata of the Rebun Group.
Neogene strata occurring on the island havebeen divided into the Motochi, Meshikuni,
Hamanaka and Kusuko Formations, In
ascending order, by Nagao el al. (1963) (Figure 2). Among these, the Meshikuni Formation includes the Meshikuni facies (mainpart) composed of siltstone and sandstone,
and the Kabuka facies composed of volcanic
conglomerates and hyaloclastites. On thebasis of moll uscan fossils, Nagao et al. (1963)'Received September 21, 1993; accepted October 15,
1993
311

Yoshikazu Noda and Koichi Hoyanagi

312

marine strata in Hokkaido, Honshu, Kamchatka and Sakhalin, this species is very
important for correlation.
The described and figured molluscan specimens are deposited at the Institute of Geology
and Paleontology, Faculty of Science, Tohoku University, Sendai, Japan (lGPS collection catalog numbers 102493-102502).
Acknowledgements

---

o

100km

Figure 1. Map showing Hokkaido and location of Rebun Island.

Nagao et al., 1963
Nonhern
regionQuaternary- Kusuko
Tertiary
Formation
AgeSouthern
regionHamanakaFormation

This study
Nonhern
regionKusukoFormation

Southern
regionHamanaka Hamanaka
Formation Formation

MioceneCretaceous

Figure 2.

Rebun
Group

Rebun
Group

Rebun
Group

Rebun
Group

Stratigraphic sequences on Rebun Island.

The authors express their deep appreciation
to Professor Kenshiro Ogasawara of the
University of Tsukuba for his critical readingof the manuscript. They are also indebted to
Dr. Louie Marincovich, Jr. of the U.S. Geological Survey for his critical review for content and English of the manuscript.
The authors thank very much Mr. Masaaki
Tsuchizawa of Rebun-cho for his willingoffer of molluscan collections. They also
thank Mr. Toshio Kuniyasu of the Control
Office of Rishiri-Rebun-Sarobetsu National
Park and Messrs. Masao Kobayashi andKazunori Morimoto of the Wakkanai Local
Forestry Office for their support to the fieldsurvey.
Thanks are also expressed to Mr. Jun
Nemoto of the Technical Staff of Tohoku
University for his photographic work.
Geological setting

The first author recently investigatedNeogene strata in the northern part of Rebun
Island, including the type locality of theMeshikuni Formation. Subsequently, theauthors concluded that the Meshikuni Formation, which yields mytilid mollusks, iscorrelative with the Asahi Formation (which
includes the Asahi fauna) in central Hokkaido.
The authors verify the presence of Mytilus
tichanovitchi Makiyama in the Meshikuni
Formation, which has already been noted by
Noda and Hoyanagi (1987) and Uozumi and
Akamatsu (1988). Since Mytilus tichanovitchi has been reported from various Miocene

Neogene strata are well exposed on Rebun
Island except for the cliffs along the western
coast. They are subdivided, in ascendingorder, into the Meshikuni, Hamanaka andKusuko Formations in the northern part andthe Motochi, Kabuka and Hamanaka Formations in the southern part, and unconformably overlie the Nairo and Anama Formationsof the Mesozoic Rebun Group (Figures 2 and
3). In this article, the authors deal mainlywith the Motochi and Meshikuni Formations.
The Motochi Formation is restricted in
distribution to the southern part of the island,

963.

Molluscan fauna from Meshikuni Formation

313

CD

CD
LEGENDf~<<I§

Kusuko Fm.
Hamanaka Fm.

[:.::.::.:.::.::.:.::j Meshikuni Fm.

~KabukaFm .

R~~~~~ Motochi Fm.
~ Cretaceous~RebunGr .
...

,./ Fault

X
Xo!

Anticline
Syncline
4km
!

o

1km

•••• I

Figure 3. Geological sketch map of Rebun
Island, mainly based on Nagao et al. (1963) . Volcanic dyke rocks are not shown.

Figur e 4. Map showing routes of examinedsections indicated in Figure 5.

and consists of green-colored tuff and sandstone in association with coal and siltstonelayers. Some moll uscan fossils are obtained
from pebbly sandstone in the lower part of
the formation (Loc. Mo-l).
The Meshikuni Formation crops out in thenorthern part of the island, dips SE, mostly
10-30·, and is composed mainly of conglomerate, sandstone, pebbly mudstone and pebbly
sandstone. Many molluscan fossils occur
from sandstone of this formation.
Along the road connecting the towns of
Uedomari and Funadomari , six geologic
columnar sections were measured andexamined (Figures 4 and 5) . They all show asimilar lithologic succession, beginning withpebbly to cobbly conglomerates at the basal
part of the Meshikuni Formation (which
unconformably overlies the basaltic lava or

consolidated tuff layers of the Mesozoic
Nairo Formation) , which are overlain bypebbly sandstone, pebbly mudstone and/or
medium- to coarse-grained sandstone of themain part of the Meshikuni Formation. The
pebbly sandstone and pebbly mudstone might
have been formed by transgressive shoreface
erosion. The sandstones (Loc. UF-l to 5)
are considered to be deposited in nearshore
environments, such as upper to lower shoreface . Moll uscan fossils occur in sandstonesof the roadcuts (Loc. UF-l , 2, and 3) andfrom nodules in pebbly mudstone that is
typically exposed alon g the Funadomari
Shrine route (Loc. Fd-2).
[n the Teppu district of the west coast, theMeshikuni Formation consists mainly ofalternating beds of siltstone and sandstone,
partly associated with slumping structures inplaces. The sediments were deposited inYoshikazu Noda and Koichi Hoyanagi

314

5

~

c:

......0
G>

c:
Q)

illIm

as
E
L..

0

::E

:::J

Fd-2.3Fd-4.5

@

..r.
~

~

0
~ '(ij E
~

0

Zu..

@t~'::~~'~'::~'iuF-4','.'.' . 3@ITIIITTIiUF...........
t±~s~:1:1:1:~)j~j~\~:~":'::"':'::'.:<...........
..~«:."............

=

I
I III
.''''~

F..·.·.1

Figure 5.

I',',:.
ffiITITl-~~:'::':":'::':":':::":'

I:m::mcoarse-grained orpebbly sandstone
conglomerate or
pebbly sandstonesi~stone:::::: m sandstonewnh pebbly......
',',',~nn:si~stone~ pebbly mudstone....

~: :~ andesne............

:::::::~,',',',............

..........
tmm,.,.,.
..........
,.,.11'..........
'-tit:.,..
cd~2..........
~~~~~~~

~

en
Q)
::E

:::::::
m

~

0

.c:

~

~

" u..

.-

(6)

to medlum[E3 finegrained sandstone

I
I -

3

~

LEGEND

Hnn~,','.',,',',',@

molluscan fossil@!:::::::IUF-2

UF-3

fossil localnies@~UF-14,',','[ill].:':.:.:.:.:
ITillill~_:'.:,::..

Columnar sections of the Meshikuni Formation.

shelf to slope environments.
At the Meshikuni coast, the type locality of
the Meshikuni Formation, a light-greencolored, very coarse-grained tuffaceous sandstone of this formation crops out along thebeach. At its base, it becomes platy, hard,very coarse-grained sandstone and overlies
hard tuff layers of the Mesozoic Anama
Formation. The hard Mesozoic rocks forma cliff-like hill just behind the beach, and it
seems that the Neogene rocks filled valley-likebasins in the older rocks. The Mesozoic
rocks occur topographically higher than the
Neogene beds that crop out almost at sea
level; strata of the latter dip only slightly andare almost horizontal in attitude. This situation suggests that the Mesozoic rocks had
cropped out of the sea and controlled thesedimentation of Neogene strata. The
mytilid fossils occur from coarse-grainedsandstone of the Meshikuni Formation along

hard black
tuffaceous sandstone~n[Localities are shown in Figure 4.

the beach (Loc. Ms-l and 2), and indicate ashallow to intertidal environment.
Figure 6 shows the geologic map of thenortheastern part of the island. The Mesozoic Nairo Formation occurs in the northeastern part of the mapped region, and controls the orientation of the peninsula. The
Tertiary Meshikuni Formation occurs in theneighboring southwest area against the Nairo
Formation. The Hamanaka Formation,
which overlies the Meshikuni Formation, is
not shown in the map, because the boundary
between the two formations is concealed. It
occurs in the western area of the mapped
region (Nagao et al., 1963). There are somefaults striking NW-SE and NE-SW. They
are inferred from lineations in aerial photographs and from the distribution of theboundary between the Nairo Formation and
the Meshikuni Formation.

963.

Molluscan fauna from Meshikuni Formation

315

The second assemblage is represented byCD"Crassostrea" gravitesta (Yokoyama) and
Glycymeris species in association with Turritella species and Laevicardium cf. shiobarense (Yokoyama). These mollusks occur insiltstone with calcareous nodules and occasionally in pebbly sandstones. This assemblage is well exposed along the stream behind
the Funadomari Shrine (Loc. Fd-I-5).
"Crassostrea" gravitesta occasionally forms ashell bed like an oyster bank.
The last assemblage is observed at localities
Tp-l and 2, where fossils also occur in siltstone without pebbly sandstone. Fossil mollusks of this assemblage are represented byrather deep, muddy bottom dwellers such as
Lucinoma acutilineata (Conrad) and
Macoma species. This assemblage is very
different from the other two in having nosandy-bottom dwellers.

LEGENDI I M~~~~~~~nNaira Formation~ Strike and dip-- Inferred faulto
1km

On Mytilus tichanovitchi Makiyama

•

-

-

-

-

I

Figure 6. Geologic map of the northeasternpart of Rebun Island.

Molluscan fauna from theMeshikuni Formation
Molluscan fossils obtained from the Meshikuni Formation are shown in Table 1, andtheir localities in Figure 7. Although themoll uscan material is insufficient for statistic
analysis, three types of molluscan assemblages
corresponding to lithofacies can be recognized.
The first assemblage occurs with mediumto coarse-grained sandstone, and is represented by Mytlius tichanovitchi in association
with pectinids and many fragments of"Balanus" species. These fossils occur sporadically and are not deformed (Loc. UF-I-5,
Ms-I,2).

Specimens of Mytilus tichanovitchi fromthe Rebun were collected from coarse-grainedsandstone at localities UF-3 and MS-2, thelatter of which is near the type locality of theMeshikuni Formation. Mytilus tichanovitchi is one of the characteristic species of Early
to early Middle Miocene faunas of the northwestern Pacific region. It is known from the
Asahi, Fukuroji and Tokomuro Formationsin Hokkaido (Uozumi, 1953; Uozumi, 1966;
Uozumi and Akamatsu, 1988; Matsuno etal., 1964; Ohara, 1966; Kanno et al., 1968 ;Mizuno et al., 1969), the Goyasu Formationin the Joban coal-field (O'Hara and Nemoto,1982), the Kuluven and Ilyin Formations in
Kamchatka and the Chekhov Formation in
Sakhalin (Gladenkov et al., 1984). Recently, M. aff. tichanovitchi was reported from theLower Miocene Hota Group of the 80so
Peninsula, central Japan (Sanada et al.,1983). Radiolarians from the Hota Group
have been assigned to the Calocyc!etta costata Zone of Riedel and Sanfillippo (1978)(Saito, 1992; S. Saito, personal communica-

316

Yoshikazu Noda and Koichi Hoyanagi

Figure 7. Maps showing localities of megafossils.
remaining localities are of the Meshikuni Formation.

Mo-l is from the Motochi Formation.

All

963.

Molluscan fauna from Meshikuni Formation

317

Table 1. List of megafossils collected from the Meshikuni and Motochi Formations UFFd-, Tp- and Ms-series are from the Meshikuni Formation. Mo-l is from the Motochi For~ation:
Localities are shown in Figure 7.
Localities UF·!
Specific nameBiva!via
Vtcharax sp.
'(;lycymeris cisshuensis Makiyama
Glycymeris idensis Kanno
Glycymeris sp.
My/ilus /ichanovi/chi Makiyama
My/itus? sp.
+
hlamys aff. ingeniosa (Yokoyama)+ectinid gen. et sp. inde!.
+"Crassos/rea" gravi/esla (Y okoyama)"Os/rea" sp.
ucinoma acUlilineala (Conrad)
aevicardium cf. shiobarense (Yokoyama)
linocardium shinjiense (Yokoyama)
Serripes aff.loperousii (Deshayes)
pisuJa sp.
MeganguJus /malwn%i (Otuka)+
Macoma izurensis (Yokoyama)
Macoma op/iva (Yokoyama)
Macomasp.
+
Macoma? sp.
+
Myasp.
Bivalvia gen. et sp. indel.
+
Gastropoda
Turri/ella (Halaiella) cf. sagai Kolaka
Turri/ella sp.
"Nalica" sp.
+Nassarius ? sp.
Nep/unea sp.
Barnacle"Balanus" sp.

+

UF·2 UF·3

UF·4 UF·5 Fd·!

Fd·2 Fd·3 Fd·4 Fd·5 Tp-!

Tp-2 Ms·!

Ms·2 Mo-!

+
+
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+

+

+

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+

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+
+

+

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tion), which is Early Miocene in age.
The presence of Mytilus tichanovitchi in
the Meshikuni Formation is important for
paleobiogeographic reconstruction of the
Early Miocene of the northwestern Pacific.
Early Miocene marine strata, such as the
Asahi Formation in the Yubari coal-field andthe Sankebetsu Formation in the Haborocoal-field in Hokkaido, are characterized by
mytilid bivalves. In Kamchatka and Sakhalin, several mytilid mollusks have beenrecorded from Miocene strata, and they seemto be consistent key markers for use in biostratigraphic correlation. The Kamchatkan
mytilid assemblages are consistent in species
composition, and the authors consider them
to be arranged in ascending order as follows:
the Early Miocene Mytilus tichanovitchi-M.
ochotensis- Modiolus wajampolkensis assem-

biage, the early Middle Miocene Mytilus
chejsleveemensis-M. (Plicatomytilus) kakertensis-M. (P.) kavranensis assemblage, andthe Middle Miocene Mytilus expansus-M.
kewi - M. coalingensis assem bl age.
Plicatomytilus species are also preserved in
the so-called Middle Miocene in Hokkaido ;the Kamikineusu and Furanui Formations in
the Hidaka area, central southern Hokkaido.
During the late Middle Miocene to Pliocene,
mytilids such as M. coruscus Gould and/or
M. grayanus Dunker, both of which areextant species, dwelled in Japan. In summary, the Japanese Neogene mytilid assemblages are represented, in ascending order, by
the Early Miocene Mytilus tichanovitchi, byMiddle Miocene Plicatomytilus species, andby Late Miocene-Recent species.

Yoshikazu Noda and Koichi Hoyanagi

318

Discussion
The Meshikuni molluscan fauna is correlative with the Early Miocene Asahi fauna ofcentral Hokkaido on the basis of the presenceof Mytilus tichanovitchi. On the other hand,
Megangulus tmatumotoi, Macoma opt iva andM. izurensis are common in the Early
Miocene Sankebetsu and Middle Miocene
Chikubetsu faunas of northwestern Hokkaido, and the Chikubetsu fauna also
includes Clinocardium shinjiense andLucinoma acutilineata (Noda, 1992). Most
Meshikuni faunal elements indicate a cooltemperate water condition, based on ecologic
data of comparable living species of the samegenera. However," Crassostrea" gravitesta
and Glycymeris cisshuensis suggest that relatively warm water influenced this area, andindicate a possibility of correlation with theTakinoue fauna of central Hokkaido,
although typical warm species of the Takinoue fauna are not present in the Meshikuni fauna.
The Early Miocene Goyasu Formation in
the loban coal-field, Northeast Honshu,
yields molluscan fossils from sandstone, andis correlative with the Meshikuni Formationon the basis of some distinctive species;

Mytilus tichanovitchi, "Crassostrea" gravitesta, Clinocardium shinjiense and Lucinoma
acutilineata (O'Hara and Nemoto, 1982,
1984 ).
No microfossils were obtained from theMeshikuni Formation. A K-Ar age of
18.2± 1.2 Ma from a dolerite outcrop beside
Kusu Lake was reported by Goto and Wad a(1991). The volcanism that produced thedolerite is thought to have been in activity at
almost the same time as the deposition of theMeshikuni Formation, because the dolerite
crops out as a dyke and a lava sheet within
the Meshikuni Formation, although Gotoand Wada (1991) stated that the volcanic
rocks are unconformably overlain by theMeshikuni Formation. This informationsuggests that the Meshikuni Formation is

Early to Middle Miocene in age.
A few molluscan fossils of the Motochi
Formation occur from a very coarse-grainedsandstone (Loc. Mo-l), cropping out at thesea side cliff just behind the hotel near the
Menou Coast on the southern part of theisland (Figure 7). Nagao et al. (l963) reported Chlamys sp., Patinopecten sp., Japelionsp., Neptunea oomurai Otuka and Psephaea
sp. from the same locality. This assemblage
seems to be equivalent to the early Middle
Miocene Chikubetsu fauna. Although there
are no species in common between the Meshikuni and Motochi Formations, these twoformations are considered tentatively to be of
the same age.
The volcaniclastic Kabuka Formation wasconsidered to be a local facies of the Meshikuni Formation by Nagao et al. (l963). It
consists of conglomeratic and hyaloclastic
sediments, and is quite different from theMeshikuni Formation. It unconformably
overlies the Motochi Formation and underlies the Hamanaka Formation. As
mentioned above, the Motochi Formation isconsidered to be the same age as the Meshikuni Formation, which suggests that the
Kabuka Formation should be treated as aseparate stratigraphic unit.

Conclusion
The Meshikuni molluscan fauna may be
divided into three assemblages, two of whichhave shallow marine elements and one ofwhich has rather deep-water elements. The
first assemblage is characterized by Mytilus
tichanovitchi, which indicates that it is suitable to correlate the Meshikuni Formation
with the Early Miocene Asahi fauna of central Hokkaido. The Meshikuni fauna is acool-temperate water fauna and is also correlative with the Sankebetsu and Chikubetsu
faunas of the Haboro coal-field, northwestern
Hokkaido.
The presence of Mytilus tichanovitchi in
the Meshikuni Formation is important for963.

Molluscan fauna from Meshikuni Formation

understanding phylogenic and systematic
problems of mytilid mollusks and understanding the origin and migration of Miocene
mollusks in the North Pacific.
The Motochi Formation seems to be asouthern equivalent of the Meshikuni Formation, although it contains very few fossils.

Description

Turritella (Hataiella) cf. sagai Kotaka, 1951
Figures 8-1 a-b

Compared with.Turritella s-hataii sagai Kotaka, 1951, p.87-88, pI.
12, figs. 13-1 7

Description.-Shell large, turreted, withmore than eight whorls. Whorl profile alittle convex, with rounded shoulder developed at abapical part (J-type of Ida, 1952).
Suture impressed. Whorls sculptured withseven to eight spiral cords, and growth lines
near aperture; spiral cords distinct in notations (u C t6 B s4 A3 r5 rl) in adult; inyoung stage, spiral u obsolete and the area
abapical to spiral C rather excavated; primary spirals flat-topped, and rather quadrate
in section; secondary spiral t narrower thaninterspaces. Base ornamented by two spiralcords of which adapical one is primary spiral
D. Growth line of double-arched type;antispirally directed sinus medium in depth
and at the point of spiral s; spirally directedsinus at the point of spiral D or sutural;
growth-line angle almost zero degree orminus. Apex and protoconch unknown.
Aperture oval in shape, not angulated.
Apical angle about 16 degrees.
Remarks.-Only one specimen was
examined. This Turritella species is the first
discovered from Rebun Island. The shell
has a rather large size compared with hitherto
reported Miocene Turritella in Hokkaido.
This specimen is closely related to Turriteffa (Hataiella )sagai Kotaka in having similar
strength of spiral cords, spiral notation and319

growth lines, but the shortage of the numberof specimens prevents a firm determination.
It is similar to T. shataii Nomura, T.
inflariata Nagao and T. tighilana Iiyina, but
is distinguished by different notation andlarger shell. Turritella kadonosawaensis
Otuka is also similar to the present specimen
in regard to the growth lines, but differs inhaving different spiral notation.
Turritella (Hataiella) sagai was originallydescribed from the Miocene Togari Formation of central Japan (Kotaka, 1951); subsequently, it has been reported mainly fromvarious strata of central Japan and also fromthe Bear Lake Formation of Alaska Peninsula (Marincovich and Kase, 1986).
Dimensions.-Height 72.4 mm. Width ofbody whorl 24.5 mm (lGPS coil. cat. no.
102493).

Gfycymeris cisshuensis Makiyama, 1926
Figures 8-2a-c, -3, -4

Glycimeris cisshuensis [sic] Makiyama, 1926, p. 155156, pI. 13, figs. 2-3; Nagao, 1928, p.29-30, pI.
2, figs. 1-3, pI. 3, figs. 1-3, pI. 4, figs. 22-25.
Glycymeris cisshuensis Makiyama. Nomura andHatai, 1937, p. 125, pI. 17, fig. 7. Itoigawa,1956,
pI. 2, fig. 1 ; Araki, 1960, p. 79-80, pI. 5, figs.
7a-c, 8; Kanno, 1960, p. 207-208, pI. 31, figs. 3435; Hashimoto, 1961, p. 81-82, pI. 9, figs. 3-6 ;Kamada, 1962, p. 62-63, pI. 3, figs. 1-3 ;Okamoto and Nakano, 1963, p.532-536, pI. 57,
figs. 1-3, text-figs. 2-4; Iwasaki, 1970, p.393394. pI. 4, figs. 6-7; Itoigawa in Itoigawa et aI.,
1974, p.57-58, pI. 5, figs. 1-5; Takahashi andNishida, 1974, p. 18, pI. 1, figs. 1-10; Itoigawa etal., 1981, pI. 3, figs.3a-b, 1982, p. 25-26; Tsuru,
1983, p. 53-54, pI. 8, figs. 13-16, pI. 9, figs. 1-2 ;Fuse and Kotaka, 1986, pI. 17, figs. 18-19.
Glycymeris vestitoides Nomura, 1935, p. 33-34, pI.
5(4), figs. 4-7; Nomura and Onisi, 1940, p. 182,
pI. 18, fig. 7; Otuka, 1940, p. 92-93, pI. 11, fig. 4;
Mizuno, 1964, pI. 3, fig. 12; Mizuno, 1965, p.
331, pI. 1, figs. 9-11: Ogasawara, 1973, p. 148, pI.
12, fig. 14; Suzuki et al., 1983, pI. 2, fig. 5.
Glycimeris cisshuensis var. compressa [sic] Nagao,
1928, p. 30, pI. 3, figs. 25, 25a-c.
Glycymeris (Veletuceta) cisshuensis Makiyama.
Matsukuma, 1986, p. 90, pI. 5, figs. 7-8.

Description.-Shell medium to large in size,

320

Yoshikazu Noda and Koichi Hoyanagi

Figure 8. la - b. Turritella (Hataiella) cf. sagai Kotaka , X I, IGPS colI. cat. no. 102493, Loc. Fd-I ,Meshikuni Formation . 2- 4. Glycymeris cisshuensis Makiy a ma , X I, IGPS co il. cat no s. 102494-4, - I
and -2, Loc. Fd- I, Meshikuni Formati o n, 2a : Dorsa l view of intact va lves, 2b: Left va lve, 2c: Right
valve, 3 : Right va lve of intact valves, 4: Right va lve o f intact va lves.

ventricose, transverse ly oval in outline, longer
than high , sli ght ly inequilateral , eq ui va lve,a nd with thick test ; po sterior part a little
longer than a nterior. Beak ort hogyrate.
Anterodorsal margin slightly convex andpassing into regularly rounded anterior m argin ; po sterodorsal margin a little co nvex andmore descending than a nterodorsa l or equal ;ve ntral margin smoothly rounded with bothanterior and posterior margin s. Surface ofshell sc ulptured with more than 30 radial rib s
a nd incremental growth lines , which so metimes making cancellate ornamentations;radial s flat-topped , much wider than interspati a l grooves, becomin g obscure near botha nterior and posterior m argins, a nd with fo ur

963.

Molluscan fauna from Meshikuni Formation

or more numbers of fine threads on the top ofthemselves of well-preserved surfaces. Inner
structure and hinge unknown.
Remarks.-Seven specimens occurred froma calcareous nodulous hard block of siltstone
containing the bank-forming "Crassostrea"
gravitesta (Yokoyama). All valves areintact, and inner characters are not observed.
However, such characters as their thick tests,
ventricose shells, surface sculptures of ribs
and fine threads on ribs are identical withthose of Glycymeris cisshuensis Makiyama.
Two of the specimens show very ineq uilateralshells, but this is considered to be intraspecific variation.
Glycymeris cisshuensis was originally described by Makiyama (1926) from the
Miocene of Korea (Mankodo Formation),and has been reported from various localities
in Japan. But some of these occurrences may
be of species other than G. cisshuensis, as
already stated by Itoigawa et al. (1982).
Matsukuma (1986) thought that G. vestitoides
Nomura, reported from Miocene strata inNortheast Honshu and Hokkaido, is synonymous with the present species, and the authors
agree with him.
This species can be distinguished from the
Recent species G. yessoensis (Sowerby) and
G. vestita (Dunker) and the fossil species G.
idensis Kanno by having a thicker test andmore convex shells. Gladenkov et al. (1984)
stated that Glycymeris slodkewitschi Sinelnikova differs from G. cisshuensis in the presence of chevrons on the cardinal area. Adding to that, this species has a more shouldered shell shape than G. slodkewitschi.
Glycymeris cisshuensis is known from the
Oligocene of Kyushu and from the Mioceneof Japan and Korea.
Dimensions (in mm.).IGPS coIl.
cat. no.

102494-1
102494-1
102494-2
102494-3
102494-4

Length Height

64.3
65.5
56.4
39.8
51.8

52.2
57.0
49.3
33.8
46.0

Thickness

11.6
18.3
28.8
19.6
26.0

Right valve (Loc. Fd-I)
Left valve (Loc. F d·l)
Both valves (Loc. F d-I)
Both valves (Loc. F d-I)
Both valves (Loc. Fd-I)

321

Distribution.-Oligocene: Yamaga, Sakamizu and Wakita Fms. (Formations), Ashiya
G. (Group), Chikuho Coal-field; Yukiaino
Sandstone, Kishima G. and Kyoragi and
Yoshinotani Fms., Ouchi G., Karatsu Coalfield; Funatsu Fm., Takashima Coal-field;
Tochinoura and Kakinoura Fms. (MazeFm.), Nishisonogi G., Sakito-Matsushima
Coal-field. All in Kyushu Island. Miocene:
Mankodo Fm., North Korea (type locality) ;Yamato (Chikubetsu) Fm., northern Hokkaido; Kamikineusu Fm., central southern
Hokkaido; Tanosawa and Nagane Fms.,
Aomori Pref.; Nishikurosawa and Senosawa
Fms., Akita Pref.; Moniwa and Nanakita
Fms., Miyagi Pref.; Taira and Kubota Fms.,
Fukushima Pref.; Itahana Fm., Gunma
Pref.; Ushikubitoge Fm., Saitama Pref.;Mizunami G., Gifu Pref.; Shiodani Sandstone, Tsuzuki G., Kyoto Pref.; Kaisekizan
Fm., Mie Pref.; Togane Fm., Shimane Pref. ;Kiwado and Taoyama Fms., Yamaguchi
Pref.; Lower Fm., Taishu G., Tsushima Is.,
Nagasaki Pref.; Kadogawa Fm., Miyazaki
Pref.
Glycymeris idensis Kanno, 1956
Figures 9-2, -3, -6

Glycymeris idensis Kanno, 1956, p.267-268, pI. 38,
figs. 1-5; Masuda and Takegawa, 1965, pI. I,
figs. 2-3; Mizuno et al., 1969, pI. 28, fig. 10.
Glycymeris (Glycymeris) idensis Kanno. Amano,
1983, p. 41, pI. 2, fig. 7; Amano, 1986, p. 193195, pI. I, figs. I a-b, 2a-b, 4.
Glycymeris i-idensis [sic] Kanno. Gladenkov et al.,1984, p. 177-178, pI. 31, figs. 9-10.

Description.-Shell small to medium insize, equivalve, subequilateral to inequilaterai, moderately convex; length almost equal
to height, ratio of shell height to length is 0.84
to 1.12, average is 0.97; test rather thick.
Beak situated almost center of shell length,
not pointed and rounded. Both antero- andposterodorsal margins almost straight or alittle convex; ventral margin regularlyrounded. Surface ornamented by 27 to more

322

Yoshikazu Noda and Koichi Hoyanagi

Figure 9. 1. Clinocardium shinjiense (Yokoy ama) , X 1.2, right valve inner mold, IGPS call. cat no.
102499, Lac. Mo-l , Motochi Formation . 2, 3, 6. Glycymeris idensis Kanno, X 1.2, IGPS colI. cat. nos.
102495- 1, - 2 and -3, Lac. Fd-2, Meshikuni Formation. 4. Lucinoma acutilineata (Conrad) , X 1, rightva lve of intact va lves, IGPS call. cat. no. !O2500, Loc. Tp-l , Meshikuni Formation. S. Serripes afr.
laperousii (Deshayes), X 1, right val ve, IGPS call. ca t. no. 102501 , Lac. Fd-2, Meshikuni Formation.

than 35 radial ribs and crossing growth lines ;ribs wider than interspatial shallow grooves,sometimes obsolete and invisible ; concentric
growth lines ventrally becoming dense andmaking more or less undulations. Interior
margin crenulated . Ligamental area sculptured with 6 to 7 chevron shaped grooves .
Remarks.-About 20 specimens from theMeshikuni Formation were examined.
Some of the Meshikuni specimens have agreater number of radial ribs than the typespecimen, but they are included in the species
vanatlOn. Amano (1986) studied the specimens from the Upper Miocene Ishiizawa
Formation of eastern Hokkaido , and subdivided them into two morphological types
according to shell shape. Type 1 is a com-

mon shape with a rather inflated and suborbicular shell , and type 2 has a moderately
inflated and longitudinally ovate shell shape.
There exist both types 1 and 2 in the specimen s at hand.
Glycymeris chitanii Yokoyama (Yokoyama , 1929 ; as Glycimeris chitanii) fromthe Bor Formation of Sakhalin is similar tothe present species in its shape, but is distinguished from the prese nt species in having amore rounded and equilateral shell, and more
numerous radials (about 50 in number).
Glycymeris yessoensis (Sowerby) greatly
resembles the present species, but has a thinner shell and more numerous radial ribs(about 40 in number).
Dimensions (in mm .).-

963.

Molluscan fauna from Meshikuni Formation

Figure 10. 1- 3. My lilus lichanovilchi Mak iyama , X I, IG PS co il. ca l. nos. 102497- 3, - 2 a nd - I,
Loc, UF-3 , Meshikuni Formation , 1 : Right va lve ; a : Side view; b : Dorsa l view, 2 : Ri ght va lve ; a :Side view ; b: Do rsa l view ; c : View from bea k, 3a : Side view of intact va lves, 3b : Dorsa l view o f
int act va lves. 4 : Megangulus lmalumofoi (O tuk a) , X I, Left va lve, o uter mold , IG PS coil. cal.n o. 102502,
Loc. UF-I , Meshikuni Formati o n.

323

Yoshikazu Noda and Koichi Hoyanagi

324
IGPS colI.
cat. no.

Length Height102495-1
102495-2
102495-3
102496

30.5
ca.40
27.3
43.4

29.2
39.0
25.3
41.8

Thickness

6.0
6.9
6.2
9.8

Left valve (Loc. Fd-2)
Right valve (Loc. Fd-2)
Right valve (Loc. Fd-2)
Left valve (Loc. Fd-3)

Distribution.-Miocene: Yoshigasawa
Fm., Miyagi Pref. (type locality); Togeshita
and Atsunai (Ishiizawa) Fms., Hokkaido;
Kanagase Fm., Miyagi Pref.; Kakert Fm.,western Kamchatka.
Mytilus tichanovitchi Makiyama, 1934
Figures 10-la-b, -2a-c, -3a-b

Mytilus tichanovitchi Makiyama, 1934, p. 134-135, pI.
4, figs. 11-12; Uozumi, 1953, p. 25, pI. 22, figs.
175a-b; Matsuno et al., 1964, pI. II, figs. 5-6 ;Uozumi, 1966, p.129-130, pI. 9, figs. 2-4, 8;
Kanno et al., 1968, p. 8-9, pI. I, figs. 4-6 ;O'Hara and Nemoto, 1982, pI. I, figs. 5-8 ;Uozumi and Akamatsu, 1988, p.332-333, pI. 3,
figs. 2, 5, 7, pI. 5, figs. 1-5.
Mytilus tihanovichi [sic] Makiyama. Gladenkov elal., 1984, p. 181, pI. 33, fig. 4, pI. 54, fig. II.

Description.-Shell medium to large in size,
equivalve, very inequilateral, very elongate
trigonal in outline, thick tested, and with
distinct dorsal wing. Beak terminal andblunt. Anterodorsal margin narrowly
rounded, making a gentle angle with ventral
margin; ventral margin nearly straight orslightly concave; anterodorsal slightly convex; posterior margin rounded. Dorsal
wing long but not wide, situated from thecorner of anterodorsal and dorsal margins to
the point about 3/5 posterior of shell length,
making a deep groove-like concavity withoverhanging dorsal slope. The peak of theumbonal ridge at about the middle of shelllength or a little anteriorly. Ventral slope
smooth or slightly depressed. Surface ornamented with distinct concentric lines ofgrowth and stair-like undulations. Marginsmooth. Inner surface with radial striations
along margins. Two anterior byssal
retractors, small, situated along the umbonalridge. Muscle scars unknown.
Remarks.- Three well-preserved specimens

and several inner and outer molds were
examined. One of them (Figures 1O-2a-b) is
identical with the specimens of Uozumi(1966) from the Asahi Formation in central
Hokkaido.
Subgenus Tumidimytilus Kafanov, 1984
was proposed on the basis of M. tichanovitchi
as the type species, but there are no descri ptions of inner structures except the dental areaand ligaments. The two small anterior
byssal retractors of this species have not been
observed before; they remind us of the inner
structure of subgenus Plicatomytilus Allison
and Addicott, 1976, which apparently has
anterior byssal retractors. These retractors
suggest that Tumidimytilus is closely related
to Plicatomytilus.
This species is easily distinguished from
Mytilus (Plicatomytilus) species from
Miocene strata of the North Pacific region by
not having plicate folds on the ventral and/or
posterior part of the valve or a wavy plane ofcommissure, and it also differs from the recent
Mytilus species by its remarkably hunchbacked shell shape and overhanging dorsal
slope of shell.
Mytilus ochotensis (Slodkewitsch) from the
Kuluven and Ilyin Formations of western
Kamchatka and the Chekhov Formation of
Sakhalin is very similar to this species in thegeneral outline of the shell and its hunchbacked shell form, but differs in having amore posteriorly situated dorsal wing and lessoverhanging dorsal area than the latter.
Mytilus chejsleveemensis Slodkewitsch fromthe Kakert Formation of Kamchatka (Slodkewitsch, 1938) is another related species, but
has a shorter and much more inflated shellwith a smaller dorsal wing than M. tichanovitchL Mytilus tichanovitchi is also
related to M. furanuiensis Uozumi and
Akamatsu from the Furanui Formation and
M_ shunbetsuensis Suzuki and Uozumi (Suzuki et al., 1983) from the Kamikineusu Formation, both in Hokkaido, but M. furanuiensis
has a dorsal plication between the dorsal
wing and dorsal slope and M. shunbetsuensis

963.

Molluscan fauna from Meshikuni Formation

has a more elongate and less inflated shell.
M. ksakurai Nomura and Hatai from the
Isomatsu Formation of the Tsugaru Peninsula, northern Honshu (Nomura and Hatai,
1936) was once considered to be a synonym of
the present species (Kanno et al., 1968), but
the former has a rather small shell and cooccurs with M. (Plicatomytilus) hidakensis
Suzuki and Uozumi (Noda and Motoyama,1991) originally described from the Kamikineusu Formation (Suzuki et al., 1983), andthe authors are inclined to identify M. ksakurai as a juvenile form of hidakensis.
Dimensions (in mm.).IGPS coil.
cat. no.

102497-1
102497-2
102497-3
102498-1
102498-2

Length HeightThickness

67.4 ca.29
76.1 37.0
97.6 45.9
67.0 ca.33
50.8 28.0

45.4
19.8
26.3+
15.2
ca.12.5

Both valves (Loc. UF-3)
Right valve (Loc. UF-3)
Right valve (Loc. UF-3)
Left valve (Loc. Ms-2)
Left valve (Loc. Ms-2)

Distribution.- ?Oligocene: Machigar Fm.
(?), North Sakhalin (type locality).
Miocene: Kuluven and Ilyin Fms., western
Kamchatka; Chekhov Fm., South Sakhalin;
Asahi (Horomui) and Tokomuro Fms., Hokkaido; "Goyasu" Fm., Fukushima Pref.
References
Allison, R.C. and Addicott, W.O., 1976: The North
Pacific Miocene record of Mytilus(Plicatomytilus), a new subgenus of Bivalvia.
U.S. Geol. Surv., Prof Paper 962, p. 1-22, pis. 13.
Amano, K., 1983: Paleontological study of the
Miocene Togeshita molluscan fauna in the
Rumoi district, Hokkaido. Sci. Rep., Inst.
Geosci., Univ. Tsukuba, Sec. B, vol. 4, p. 1-72,
pis. 1-8.
- - , 1986: Age and characteristics of the so-called"Atsunai-Togeshita fauna" in Hokkaido.
Palaeont. Soc. Japan, Spec. Papers, no. 29, p.
187-198, pI. 18.
Araki, Y., 1960: Geology, paleontology and sedimentary structures (including Problematica) of
the Tertiary formations developed in the environsof the Tsu City, Mie Prefecture, Japan. Bull.
Lib. Arts Dep., Mie Univ., Spec. Vol., no. I, p. 1\18, pis. I-II.
Fuse, K. and Kotaka, T., 1986; The molluscan assem-

325

blages from the Hioki Group, Yamaguchi Prefecture, Southwest Japan. Monogr. Mizunami
Fossil Mus., no. 6, p. 119-141, pis. 17-18. (inJapanese with English abstract)
Gladenkov, Yu,8., Gladikova, V.M., Kafanov, A.I.,
Konova, L. V., Krishtofovich, L. V., Sinelnikova,
V.N. and Popov, S.V., 1984: Marine mollusks.
In, Menner, V.V. ed., Atlas of fauna and floraof the Neogene sediments of the Far East.Tochilinski key section of western Kamchatka-.
Acad. Sci. USSR, Order of the Red Banner ofLabour Geological Institute, Trans., vol. 385, p.
152-250, pis. 27-65. Izdatelstvo "Nauka", Moscow. (in Russian)Goto, Y. and Wada, K., 1991: The middle Miocene
alkali dolerite from Rebun Island, northeasternJapan Sea. Jour. Mineral. Petrol. Econ. Geol.,
vol. 86, p. 375-378.
Hashimoto, W., 1961 : Tertiary molluscan fauna fromthe Kadogawa Formation, Miyazaki Prefecture,
Japan. Rep. Earth Sci. Dept., General Educ.,
Kyushu Univ., vol. 7, p. 69-94, pis. 8-10.
Ida, K, 1952: A study of fossil Turritella in Japan.
Geol. Surv. Japan, Rep., no. 150, p. 1-62, pis. 17.
Itoigawa, J., 1956: Molluscan fauna of the Tsuzuki
Group in the Kyoto Prefecture, Japan. Mem.
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- - , Shibata, H. and Nishimoto, H., 1974: Molluscan fossils of the Mizunami Group. Bull.
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Kafanov, A.I., 1984: System of subfamily Mytilinae(Bivalvia, Mytilidae). In, Nevesskaya, L.A. ed.,Morphology, systematics, phylogeny andecogenesis of bivalve mollusks, p. 43-45. Acad.
Sci. USSR, Paleont. Inst., Moscow. (in Russian)(fide Kafanov, A.I., 1987)- - , 1987: Subfamily Mytilinae Rafinesque, 1815
(Bivalvia, Mytilidae) of the Cenozoic North
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Polar Basin. p. 65-103. Acad. Sci. USSR, Far
East Sci. Cent., Inst. Mar. BioI., Vladivostok. (inYoshikazu Noda and Koichi Hoyanagi

326

Russian)Kamada, Y., 1962: Tertiary marine Mollusca fromthe Joban Coal-field, Japan. Palaeont. Soc.
Japan, Spec. Papers, no. 8, p. 1-187, pis. 1-21.
Kanno, S., 1956: On some new glycymerid molluscs
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- - , 1960: The Teritary System of the Chichibu

Basin, Saitama Prefecture, central Japan. Part
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Makiyama, J., 1926: Tertiary fossils from North
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Matsukuma, A., 1986: Cenozoic glycymeridid
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Mizuno, A., 1964: A study on the Miocene moll uscan faunas of the Kitatsugaru and Nishitsugaru
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stratigraphy of the Kushiro Coal field, eastern
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Chokubetsu formation. Ibid., vol. 20, no. 10, p.
633-649, pis. 27-29. (in Japanese with English

abstract)
Nagao, S., Akiba, C. and Omori, T., 1963: Rebunto(Asahigawa-2 and 6). Explanatory text ofthe geological map of Japan, scale 1: 50, 000,
43p. Hokkaido Development Agency, Sapporo.

(in Japanese with English abstract)
Nagao, T., 1928: Palaeogene fossils of the Island of
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pis. 1-17.
Noda, Y., 1992: Neogene molluscan faunas from theHaboro coal-field, Hokkaido, Japan. Sci. Rep.,
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with Programs, 94th Ann. Meet., Osaka, p.220.

(in Japanese)- - and Motoyama, /., 1991: Geology of the
Isomatsu Formation, northern part of Tsugaru
Peninsula, and its molluscan fauna. Palaeonl.

Soc. Japan, Abstracts with Programs, 140th
Regular Meet., Chiba, p.47. (in Japanese)
Nomura, S., 1935: Miocene Mollusca from Siogama,Northeast Honsyu, Japan. Saito Ho-on K ai
Mus., Res. Bull., no. 6, p. 193-234, pis. 16-17.
- - and Hatai, K., 1936: A note on some fossil
Mollusca from Tugaru Peninsula, Northeast
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- - and - - , 1937: A list of the Miocene Mollusca and Brachiopoda collected from the region
lying north of the Nanakita River in the vicinityof Sendai, Rikuzen Province, Japan. Saito
Ho-on Kai Mus., Res. Bull., no. 13, p. 121-145,
pis. 17-21.
- - and Onisi, H., 1940: Neogene Mollusca fromthe Sennan district, Miyagi Prefecture, Japan.
Japan. Jour. Geol. Geogr., vol. 17, nos. 3-4, p.
181-194, pis. 17-19.
Ogasawara, K., 1973: Moll uscan fossils from the
Nishikurosawa Formation, Oga Peninsula, Akita
Prefecture, Japan. Sci Rep., Tohoku Univ., 2ndSer. (Geol.), Spec. Vo/., no.6 (Prof K. Hatai
Mem. Vol.), p. 137-155, pis. 12-13.
Ohara, S., 1966: Stratigraphy and geologic structureof the Mid-Tertiary System on the western borderof the Yubari Coal-field, Hokkaido, Japan.

Jour. Coli. Arts Sci., Chiba Univ., Nat. Sci. Ser.,vol. 4, no. 4, p. 599-615. (in Japanese with Eng-

lish abstract)
O'Hara, S. and Nemoto, N., 1982: Molluscan fossils
from the "Goyasu Formation" in the Futaba
district of the Joban Coalfield. Jour. Coli. Arts

963.

Molluscan fauna from Meshikuni Formation

Sci., Chiba Univ., B-15, p. 57-64, pis. 1-3.
- - and - - , 1984: Molluscan fossils from thetype Goyasu Formation of the Joban Coalfield.
Ibid., B-I7, p. 45-61, pis. 1-2.
Okamoto, K. and Nakano, M., 1963: Clycymeris andCultel/us from the Teritary Hioki (Ashiya) in the
Yuya-wan area, Yamaguchi Prefecture, Southwest Japan. Ceol. Rep., Hiroshima Univ., no.
12, p. 531-539, pI. 57.
Otuka, Y., 1934: Tertiary structures of the northwest
end of the Kitakami Mountainland, Iwate Prefecture, Japan. Bul/. Eanhq. Res. Inst., vol. 12, pt.
3, p. 566-638, pis. 44-51.
- - , 1940: Miocene Mollusca from Tesio Province,
Hokkaido. Japan. Jour. Ceol. Ceogr., vol. 17,
nos. 1-2, art. 8, p. 91-99, pI. II.
Riedel, W.R. and Sanfilippo, A., 1978: Stratigraphy
and evolution of tropical Cenozoic radiolarians.
Micropaleontology, vol. 24, p.61-96.
Saito, S., 1992: Stratigraphy of Cenozoic strata in thesouthern terminus area of Boso Peninsula, Central Japan. Contr., Inst. Ceo!. Paleont., Tohoku
Univ., no. 93, p. 1-37, pI. I. (in Japanese withEnglish abstract)
Sanada, S., Harada, T. and Nagai, I., 1983: Occurrence of Mytilus from Oyama, Kamogawa City,
Chiba Prefecture, Japan. "Saishuu-to-Shiiku",
vol. 45, no. 12, p.534-535. (in Japanese)
Siodkewitsch, V.S., 1938: Tertiary Pelecypoda fromthe Far East. Paleontology USSR, Academyof Sciences of the USSR, Paleontological Institute, vol. 10, pt. 3, fasc. 18/19, part I, p. 1-508 (inRussian); part 2, p. 1-275, pis. 1-106.

can fossils from the Neogene deposits scatteredalong the western wing of the Hidaka Mountains,
Hokkaido. Jour. Fac. Sci., Hokkaido Unil'.,
Ser. 4, vol. 20, nos. 2-3, p. 225-248, pI. 4.
Takahashi, K. and Nishida, T., 1974: On the molluscan fossils from the lower formation of the
Taishu Group in the Kamiagata district of Tsushima Islands. Bul/. Fac. Lib. Art., Nagasaki
Univ., Nat. Sci., vol. 15, p.15-20, pI. I. (inJapanese with English abstract)
Tsuru, T., 1983: Middle Miocene molluscan faunafrom the Togane Formation in Hamada City,
Shimane Prefecture, Southwest Japan. Bul/.
Mizunami Fossil Mus., no. 10, p.41-83, pIs. 819. (in Japanese with English abstract)
Uozumi, S., 1953: Illustrated Cenozoic fossils ofnorthern Japan, 23. Miocene fossils from Hokkaido (part I). Cenoz. Res., no. 18, p. 356-358,
pI. 22. (in Japanese)- - , 1966: Neogene molluscan fauna in Hokkaido;
Part I. Description of the Asahi fauna associatedwith Mytilus tichanovitchi Makiyama, from Ikushunbetsu district, central Hokkaido. Jour. Fac.
Sci., Hokkaido Unil'., Ser. 4, vol. 13, no. 2, p.
119-137, pIs. 9-10.
- - and Akamatsu, M., 1988: Notes on the four
diagnostic Miocene mytilids of Hokkaido, North
Japan. Saito Ho-on Kai Spec. Publ. (Prof T.
Kotaka Commem. Vol.), p.325-338, pIs. 1-5.
Sendai.
Yokoyama, M., 1929: Moll uscan fossils from Karafto. Jour. Fac. Sci., Imp. Univ. Tokyo, Sec. 2,
vol. 2, pt. 9, p. 369-398, pIs. 71-76.

Suzuki, S., Sakai, A. and Uozumi, S., 1983: Mollus-

Anama 7 T <', Funadomari frJ iB, Furanui 7 =;7 )I. 1', Hamanaka ~ <p, Kabuka ~ i~,Kamikineusu J:.;f'fS, Kusu ~:fl, Kusuko ~:fl1lifl, Meshikuni 15 00, Meshikuni Formation.) :/ !J =-~, Motochi 5[; Jig , Nairo pn~, Rebun ?L:t, Uedomari J:.iB, Teppu ~Iff.
~t;4ijli~t$, tL3t!f,O),*,iIT~Jc~ ~ 7 -=I!O)~{t:o~: ?L:t~~t$vc5t;fjJi" Q <Pti';tIt.) :/!J =-I!;IJ'!?gEt:f:j L-t-::.F!{tEffvCf1, -=-::lO)ff~;IJ~~;f;,<'0!?.:ht-::o --::l11 Mytilus tichanovitchi
vcf;;*~;jl,Qff~-c', =-::l<'011 "Crassostrea" gravitesta t Clycymeris spp. O)ff~-c·t5I),
t (, vc~W-c·t5 I), t (, vc~wO)1i'j;Jfr.ff~-c·a; Qo~-=O)ff~11 Lucinoma ~o)~~~\, 'tie

Jfr.ff~-C·a;Qo Mytilus tichanovitchi 11~tw)lt>a:'lt G<'0-lj-?' ~ '/~7J AT -\' 'Y 7J -C'{'¥IH5-~.:h -C:to I), .):/!J =-I!;IJ'!?O)gEt:f:j11~t*fr=f:Jig:Jgl(O) Mytilus O)ffif~5t;fjJ~~ X.QJ:.-C'£~-C' a; Q .):/!J =- ~ It Mytilus tichanovitchi 0) gEt:f:j vC J: -:> -c~tw~<P:'R:$O)ijiJl B ~ C)<;J it-C'~Q t t {'vc, -t:"O){tEJ1'J:@:;IJ'!?ft, ;r.]iJ!lI/;Jig:Jgl(O)~5.iIj·-==€5.iljtIJ!ffr.lffvcx;Jlt"iiJfili-c·t5Qo
ffE87J5fIJ • i!iI:i1ip)jt0

327

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

New Series No. 172

December 30, 1993

CONTENTS
TRANSACTIONS
959.

Yukio Yanagisawa: Phylogenetic significance of a new araphid diatom
species, Delphineis kamenooensis sp. nov. .............................. 251

960.

961.
962.
963.

Ge Sun, Takashi Nakazawa, Tamiko Ohana, and Tatsuaki Kimura: Two

Neozamites species (Bennettitales) from the Lower Cretaceous of Northeast
China and the Inner Zone of Japan .................................. 264
Katsumi Veno and Sumio Sakagami: Middle Permian foraminifers from Ban
Nam Suai Tha Sa-at, Changwat Loei, northeast Thailand ................ 277
Joe S.H. Collins, Yasumitsu Kanie, and Hiroaki Karasawa: Late Cretaceous
crabs from Japan .................................................... 292
Yoshikazu Noda and Koichi Hoyanagi: Molluscan fauna from the Miocene
Meshikuni Formation on Rebun Island, northern Hokkaido

311