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Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 144

Palaeontological Society of Japan
December 30, 19860031-0204

Co-Editors

Hisayoshi IGO and Takashi HAMADA

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The fossil on the cover is Unuma (Spinunuma) echinatus ICHIKAWA and YAO, a
Middle Jurassic multisegmented radiolaria from Unuma, Gifu Prefecture, central
Japan (photo by A. YAO, x 260).

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 144, pp. 479-489, pI. 95, December 30, 1986

821. EARLY CRETACEOUS PULSIDIS (BIVALVIA) FROM JAPAN*
TAKESHI KOZAI
Higashi Primary School, Kochi 782

Abstract. Five species in the genus Pulsidis Ota including four new species, i.e.,
P. angulata, sp. nov., P. sanchuensis, sp. nov., P. tashiroi, sp. nov. and P. rostrata,
sp. nov., are described from fourteen localities of Lower Cretaceous in Japan on the
ground of my own collections as well as the hitherto treated ones.

Introduction

Localities and geological data

Since the genus PuZsidis was established by
Ota (1964), several species referable to thisbivalve genus have been described from theCretaceous brackish-water deposits in Japan.
However, some of them should be re-examinedtaxonomically. In the present paper, I attempted to the taxonomy and to determine theirstratigraphic ranges based upon the numerous
specimens collected from the various localities
in Japan.
Recently, the correlation for the non-marine
and brackish-water deposits in the Cretaceousof Japan has become much clearer in the lightof the interrelation with the ammonite-bearing
marine formations (Matsumoto et aZ., 1982).
The brackish-water deposits which contain fossilsof PuZsidis occur at various stratigraphic levels.
Therefore, it may be possible to know the geological ages of the classified species.
The specimens described in this paper are
mostly kept in the Geological Collections ofFaculty of Science, Kochi University (KSG),
Kochi and Department of Earth Sciences, Fukuoka University of Education (GT), Fukuoka.

The localities of the fossils described in thispaper are recorded below with a brief note on
the lithology, stratigraphic position and geological age. Readers may refer to Text-fig. 1 forthe geographic position of the localities.
Loc. 1. Black carbonaceous shale in theupper part of the Funagawara Formation at aquarry near Yamabagoe, about 2 km west fromthe center of Of una to City (Onuki and Mori,
1961). The characteristic species are Nanonavis
yokoyamai (Yabe et Nagao) and Isodomella
shiroiensis (Yabe et Nagao) which are marine
and brackish-water molluscs. An ammonite
Crioceratites ishiwarai (Yabe et Shimizu) which
occurs in the same formation indicates Upper
Hauterivian (Obata and Matsumoto, in Matsumoto et al., 1982). Hauterivian.
Loc. 2. Black carbonaceous shale and finegrained sandstone in the upper part of theFunagawara Formation at a roadside exposure,near Yamabagoe. This locality is of the same
stratigraphic position as Loc. 1. Hauterivian.
Loc. 3. Fine-grained sandstone in the upper
part of the Funagawara Formation at a roadside
exposure, about 1.2 km northwest of Funagawara, Of una to City. This is also of the same
stratigraphic position as Loc. 1. Hauterivian.
Loc. 4. Black shale of the lower part of the
Sebayashi Formation on the east side of theRiver Mamonozawa, near Sebayashi, south of* Received July 15, 1985; revised manuscript
received July 19, 1986; read January 22, 1984
at Kyoto University.
479

Takeshi KOZAI480

AYAMABAGOE
N

}

E
N

t
F ------"\

o

;

_ _11

cr - - - 7 ''" --~
1----1 k m-==::::::::"",YOSHIMIG14

KAWANOUCHI

N

ERI

5 km
---

Text-fig. 1.

Map showing fossil localities.

821. Pulsidis from Japan
Kagahara, Nakazato-mura, Tano-gun, Gunma
Prefecture. Brackish-water molluscs such as
Costocyrena radiatostriata (Yabe et Nagao),
Isognomon sanchuensis (Yabe et Nagao), Tetoria
sanchuensis (Yabe et Nagao) etc. occur abundantly in this shale. Upper Barremian or Lower
Aptian (Obata and Matsukawa in Matsumoto
et al., 1982; Matsukawa, 1983).
Loc. 5. Dark gray shale and fine-grained sandstone of the upper part of the Shiroi Formation
at roadside exposure, near Shiroi, Ueno-mura,Tano-gun, Gunma Prefecture. The characteristic
species are Hayamina naumanni (Neumayr),
Costocyrena otsukai (Yabe et Nagao) andIsodomella shiroiensis (Yabe et Nagao) which
are brackish-water molluscs. Hauterivian? (Matsukawa, 1979, 1983).
Loc. 6. Dark gray siltstone in the basal partof the Hibihara Formation at the dam site of
Hibihara, Kahoku-cho, Kami-gun, Kochi Prefecture. The characteristic species are Costocyrena
minor Ohta, Barbatia hibiharensis Tashiro et
Kozai and Brachidontes pyriformis Tashiro et
Kozai. Probably Lower Aptian (Tashiro andKozai, 1984; Tanaka et al., 1984).
Loc. 7. Dark gray siltstone in the lower partof the Yunoki Formation at a river side outcropof Todoronotaki, near Yunoki, Kahoku-cho,Kami-gun, Kochi Prefecture. This horizon iscalled the "Costocyrena Bed". The characteristic
species is Costocyrena radiatostriata (Yabe et
Nagao), and mega-phytofossils are occurred in
the lower part of this shale. Probably Upper
Barremian (dittos).
Loc.8. Fine-grained sandstone in the basal
part of the Hibihara Formation at a roadside
exposure, about 500 m south of Kawanouchi,
Monobe-mura, Kami-gun, Kochi Prefecture. The
characteristic species are Hayamina sp. andCostocyrena minor Ohta. This locality is probably of the same horizon as Loc. 6. Probably
Lower Aptian (Tashiro et al., 1980).
Loc. 9. Siltstone, exposed on roadside, about
800 m south of Ryoseki, Nankoku City, Kochi
Prefecture. This locality was referred to the
Hibihara Formation by Ishizaki (1960). Later
Shasticrioceras aff. S. nipponicus Matsumoto
was collected from the lower horizon of thissiltstone. Judging from this ammonite, thislocality is probably assigned to the Yunoki
Formation. Probably Barremian.

481

Loc. 10. Fine- to medium-grained sandstone
in the upper part of the Ryoseki Formation ata roadside exposure of Yonenomoto, Kochi
City. The characteristic species is Costocyrena
otsukai (Yabe et Nagao). Probably Hauterivian(Tashiro and Kozai, 1984).
Loc. 11. Dark gray, fine-grained sandy shalein the Yoshimo, Shimonoseki City, Yamaguchi
Prefecture. Brackish-water molluscs from theformation had been described by many authors(Hase, 1960). Neocomian.
Loc. 12. Siltstone in the lower part of the
Kawaguchi Formation on a coast of Tanoura,
Tanoura-cho, Ashikita-gun, Kumamoto Prefecture. It contains characteristic brackish-watermolluscs, e.g., Eomiodon matsumotoi Ohta,
Isodomella matsumotoi Ohta and Eoursivivas
matsumotoi (Hase). Neocomian (Ohta, 1977).
Loc.13. Dark-gray sandy shale in the upper
part of the Kawaguchi Formation on the northeast side of the River Kumagawa, near Kawaguchi, Sakamoto-mura, Yatsushiro-gun, Kumamoto Prefecture. This locality is the type-localityof the Kawaguchi Formation, which consistsof dark-gray sandy shale and fine-grained sandstone in alternation. Many brackish-water molluscs are occurred there (Ohta, 1977). Neocomian.
Loc. 14. Dark-gray sandy shale in the upper
part of the Kawaguchi Formation at a roadside
exposure of Sakayoriue, Toyo-mura, Yatsushirogun, Kumamoto Prefecture. The characteristic
species are Hayamina naumanni (Neumayr),
Eomiodon matsumotoi Ohta, Tetoria yoshimoellsis Ohta and Eoursivivas matsumotoi (Hase)(Ohta, 1977). Neocomian.

Systematic description
Family Corbulidae Lamarck, 1818
Subfamily Corbulinae Lamarck, 1818
Genus Pulsidis Ohta, 1964

Type species:-Pulsidis nagatoensis Ota, 1964:
Yoshimo Formation, Yamaguchi Prefecture:
Neocomian.
Diagnosis:-Shell small, inequivalve, inequilateral, elongate trigonal and posteriorly rostrated; right valve larger than left valve; test
thin; surface ornamented with regular concentricTakeshi KOZAI

482

ribs; internal margin of right valve grooved forreception of left valve; cardinal tooth of rightvalve generally small; resilial pit large, deep;chondrophore prominent, triangular, with out
median ridge.
Remarks:-Hase (1960) described Corbula
matsumotoi on the basis of the specimens fromthe Lower Cretaceous Kawaguchi Formationin Kumamoto Prefecture and Yoshimo Formation in Yamaguchi Prefecture. Ota (1964),
however, established the genus Eoursivivas on
the specimens from the Kawaguchi Formation
and Pulsidis on those from the Yoshimo Formation.
In this genus, ratio of H/L is gradually small
with growth. This genus is closely similar to
Caryocorbula Gardner, 1926, but evidently
differs in having no median ridge on the chondrophore. It is similar to Caestocorbula Vincent,1910 in the features of hinge structure, but
discriminated from the latter in its smaller
cardinal tooth of right valve and lack of siphonal
plate.
Ota (1964) described Pulsidis okadai Ota
and Aloides higoensis Matsumoto, but in myopinion, P. okadai and A. higoensis do notbelong to Pulsidis. The former is probably referable to Nipponicorbula because both valves
have weak radial ribs on the disk, and the latterto Caestocorbula because the cardinal tooth ofright valve is large and the test is thick. The
specimens which were described as P. higoensis
by Shikama and Suzuki, 1982 from the Lower
Cretaceous of Choshi, Chiba Prefecture, weretreated as Caestocorbula shikamai Hayami by
Hayami (Hayami and Oji, 1980).
Pulsidis nagatoensis Ota
PI. 95, Figs. 1-4.
1964. Pulsidis nagatoensis Ota, p. 20. text-fig.

1, pI. 20, figs. 1-14, 25.
1975. Pulsidis nagatoensis Ota; Hayami, p. 146.

Material:-The holotype is an internal mouldof right valve (GT. Y6301) illustrated by Ota(1964, pI. 20, fig. 1). It was collected from the

Yoshimo Formation at Yoshimo, Shimonoseki
City, Yamaguchi Prefecture. About 150 specimens from Locs. 9-14 and 7 are also concernedwith the description below.
Diagnosis:-Small-sized species of Pulsidis
characterized by shortly rostrated shell, umbo
situated at mid-length of valve and a shallow
radial groove on area.
Description:-Shell small, moderately inflated,
subtrigonal, a little longer than high, and rostrated to postero-ventral edge; right valve larger
than left; test thin; area demarcated from disk
by a weak posterior carina, divided into twoparts by a shallow median radial groove; anterodorsal margin short, broadly arched or nearlystraight; postero-dorsal margin long, subhorizontal, but weakly concave near umbo; ventral
margin broadly arched; posterior margin subtruncated; umbo located at nearly mid-lengthfrom front of valve, moderately prominent andslightly prosogyrate; surface ornamented withconcentric ribs, which are round-topped, regularly spaced, broader than their interspaces,
denser in posterior half than in anterior; internal
margin of right valve distinctly grooved as a
receptional scar of margin of left valve; innerareal part flat; cardinal tooth of right valve
triangular, relatively large, but narrower androunded at top; resilial pit large, situated on
posterior side of cardinal tooth; anterior adducter scar weakly impressed, long-elliptical;
posterior adductor scar well marked, larger
than anterior one; chondrophore relatively
broad, separated from dorsal margin by a shallow
groove.
Observation:-In the adult specimens, shelllength is about 10 mm in the right valve and9 mm in the left valve. Although the external
concentric ribs are more or less weathered,some specimens show clearly concentric ribs on
the entire surface. The number of ribs about 5
in the distance of 1 mm in height. Apical angle
is generally about 100 0 in the young specimens,
but in the adult specimens 1200 -130 0 in theright valve and 105 0 -115 0 in the left valve,
respectively. H/L of the valve is 0.70 on anaverage in the right valve and 0.73 in the left.

821. Pulsidis from Japan

483

Measurements (in mm):Specimens

L

H

KSG-K001, R.in.m.

8.7
7.3

6.2

KSG-K003, L.in.m.
KSG-K044, L.ex.m.
KSG-K005, R.in.m.
KSG-K006A, R.in.m.

9.8
9.0
9.8
9.0

5.6
6.9
6.3

B

U
4.5
4.0

1.9

6.5

4.6
4.1

H/L
0.61

U/L
0.52

0.77

0.55
0.47

0.70
0.70
0.66

0.46
0.46

6.4

4.5
4.6

0.71

0.51

KSG-K006B,R.in.m.
KSG-K006C, Rin.m.

9.2

6.9

4.5

0.71

0.50

KSG-K006D, L.in.m.

7.4

5.7

3.6

KSG-K007, L.in.m.

8.6

6.2

4.8

0.77
0.72

0.49
0.56

KSG-K008, R.ex.m.

10.7

7.1

4.9

0.66

0.46

1.6

KSG-K009, L.in.m.

7.1

4.5

3.5

0.63

0.49

KSG-K010, R.in.m.
KSG-KOll, L.ex.m.

7.9
9.1

6.0

4.7

6.8

3.0

0.76
0.75

0.59
0.43

KSG-K012, R.ex.m.

9.6

6.5

3.7

0.68

0.47

4.0
4.5

L.=left valve, R.=right valve, ex.m.=external mould, in.m.=internal mould,
H=height, L=length, B=breadth, U=distance from beak to anterior end.
Ratio of U/L is 0.46 on an average in the rightvalve and 0.51 in the left. The umbo of the rightvalve is more anteriorly located than of theleft.
Occurrence:-Abundant at Locs. 9 and 11,
common at Locs. 12 and 14 and rare at Locs. 7
and 10. Total stratigraphic range is Neocomian.

Pulsidis angulata, sp. nov.
PI. 95, Figs. 17- 21.
Material:-Holotype,
KSG-K022, internalmould of left valve; 9 paratypes; KSG-K017
and KSG-K014, internal moulds of right valves;
KSG-K023 and KSG-K019, internal moulds
of left valves; KSG-K021, external mould of left
valve; KSG-K018, KSG-K024, KSG-K025 and
KSG-K020, external moulds of right valves; all
the specimens were collected from Loc. 2.
Diagnosis:-Small-sized species of Pulsia ischaracterized by shortly rostrated right valve,slightly smaller, less rostrated left valve anddistinctly angulate posterior carina.
Description:-Shell small, subelliptical in

outline, moderately inflated; right valve a little
larger than left, shortly rostrated to posterior;
test thin; umbo situated at about mid-length ofvalve, slightly prosogyrate; anterior dorsal margin
short, slightly convex; posterior dorsal margin
nearly straight or gently concave; posterior
margin vertically truncated; anterior margin
semicircular; ventral margin gently arched;
posterior carina distinctly angulated, extended
from umbo to postero-ventral corner, defining
postero-dorsal area from disk; area narrow,nearly flattened, longer in right valve than left;
surface ornamented with fine concentric ribs
which are round-topped, regularly spaced, denser
on posterior part, narrower than their interspaces; interior of right valve grooved to receive
the margin of left valve; pallial line not sinuate,
indistinctly impressed; cardinal tooth of rightvalve triangular, rounded in upper part; resilial
pit triangular, smaller than cardinal tooth,
deeply excavated, located on posterior side ofcardinal tooth; chondrophore prominent, triangular, narrow but elongate, parallel to posterior
dorsal margin.

484

Takeshi KOZAI
Measurements (in mm):Specimens
KSG-K022, L.in.m.
KSG-K017, R.in.m.
KSG-K014, R.in.m.
KSG-K023, L.in.m.
KSG-K021, L.ex.m.
KSG-K019, L.in.m.
KSG-K020, R.ex.m.
KSG-K018, R.ex.m.
KSG-K015, L.ex.m.
KSG-K013, R.ex.m.
KSG-K016, R.ex.m.
KSG-K024, R.ex.m.
KSG-K025, R.ex.m.

L
6.4
7.2
5.6
5.9
5.4
5.2
6.3
6.0
5.2
7.3
6.1
6.3
5.4

Observation:-The outline of this new species
is variable from triangular to pyriform. Surface
ornamentation is well preserved in many specimens. Ratio of U/L is rather constant, being
0040 on an average in right valve and 0.38 in left
valve.
Comparison :-This new species is generally
similar to Pulsidis nagatoensis Ota, 1964, as
described before, but differs from the latter inhaving distinct posterior carina and anteriorly
located umbo. It is similar to Corbula globosa
Tamura, 1959, from the Upper Jurassic Sakamoto Formation in Kumamoto Prefecture, inthe triangular outline, but the angulated posterior carina is distinctly sharper in this species.
Occurrence :-Common at Locs. 1 and 2 andrare at Locs. 3 and 5. Hauterivian.
Pulsidis sanchuensis, sp. nov.
PI. 95, Figs. 22-30.
Material:-Holotype, KSG-K086, internalmould of right valve; three paratypes; KSGK087, internal mould of left valve; KSG-K088,
external mould of right valve; KSG-K089,
external mould of left valve; other 16 specimens
are also at hand; all the specimens were collected

H
3.5
4.3
3.9
3.9
3.6
3.4
4.3
4.0
3.3
4.6
3.5
3.8
3.6

B

1.0
1.4
1.9
1.5
1.9
1.2
1.3
2.0

U
3.0
3.5
2.8
2.9
2.8
2.3
3.3
3.7

H/L
0.55
0.60
0.70
0.66
0.67
0.65
0.68
0.67

2.9
3.7
2.8
2.8
2.6

0.63
0.63
0.57
0.60
0.67

U/L
0.47
0.49
0.50
0.49
0.51
0.44
0.52
0.62
0.56
0.51
0.46
0.44
0.48

from Loc. 4.
Diagnosis:-Small-sized species of Pulsidis
characterized by well rostrated right valve,shortly rostrated left valve, absence of mediangroove on area and umbo situated at one-thirdof shell-length in right valve.
Description:-Shell of moderate size, inequilateral, rostrated posteriorly, inequivaleve; rightvalve larger than left; test thin; anterior dorsal
margin slightly concave; posterior dorsal margin
gently concave; ventral margin gently arched;umbo slightly prosogyrate, situated at about
one-third from anterior extremity in right valve
but about a half of shell-length in left valve;area flattened, longer in the right valve thanin the left; surface covered with fine concentricribs, which are round-topped, regularly spaced,narrower than their interspaces; pallial line
indistinct; posterior adductor scar elliptical,
clearly impressed, faint larger than anterior one;resilial pit deep, large; chondrophore small,slightly prominent.
Observation:-In the adult specimens, averageof shell-length is about 8 mm in the right valve
but 6 mm in the left. Surface ornamentation iswell preserved. The concentric ribs are strong andregularly spaced, numbering about 5 in the distance of 1 mm along height of the adult speci-

821. Pulsidis from Japan

485

Measurements (in mm):Specimens

L

H

KSG-K086, R.in.m.

7.5

4.6

2.4

0.61

0.32

KSG-K087, L.in.m.

3.4

2.6

1.5

0.77

0.44

0.59
0.70
0.59

0.43
0.42
0.34

0.57
0.53
0.72

0.42
0.30
0.46

B

U

KSG-K088, R.ex.m.

6.1

3.6

KSG-K089, L.ex.m.
KSG-K090, R.in.m.
KSG-K091, L.
KSG-K092, R.ex.m.
KSG·K093, R.ex.m.

5.0
5.8
6.6+
7.1+

KSG-K094, R.

6.5
6.6

3.5
3.4
4.5
5.3
3.7
3.5

1.9
1.4
2.2
2.3

2.6
2.1
2.0
3.2
2.7
2.7
2.0

KSG-K095, L.

5.0

3.6

1.2

2.3

mens. In the adult specimens, the apical angle
is 100°-110° in the right valve and 90°-100°in the left.
Comparison:-This new species is similar to
Pulsidis angulata, sp. nov. in its trigonal outlineof the left valve, but is distinguished from thelatter by its more rostrated right valve andindistinct posterior carina. This new species
evidently differs from Pulsidis nagatoensis in its
longer area of the right valve.
Occurrence:-Rare at Loc. 4. Upper Barremian or Lower Aptian.

Pulsidis tashiroi, sp. nov.
PI. 95, Figs. 5-9.

Material:-Holotype, KSG-K042, right valve;nine paratypes; KSG-K038, KSG-K039A and
KSG-K046, internal moulds of left valves; KSGK039B, KSG-K045, KSG-K044, KSG-K047 and
KSG-K048, right valves; KSG-K049, left valve;
all these type-specimens were collected from
Loc.7.
Diagnosis:-Shell medium-sized for the genus,
elongate pyriform in outline, with rostrum
gradually narrowed to posterior end and faintly
grooved area. Right valve slightly larger thanleft.
Descrip tion :-Shell moderately inflated, inequilateral, elongate subpyriform, much longer
than high, gradually rostrated to posterior edge;

1.4
1.6

HjL

UjL

right valve slightly larger than left; test thick forgenus; umbo small, slightly prosogyrate, situatedat about one-third of shell length from front inright valve but somewhat more posteriorly in
left; surface ornamented with concentric ribs;
concentric ribs round-topped, narrower thantheir interspaces; area elongated, faintly grooved
along its median line; a low ridge extended fromumbo to postero-ventral margin, defining
area from disk; anterior dorsal margin slightly
arched; posterior dorsal margin a little concave
near umbo but slightly convex in later half;
posterior end subvertically truncated; ventral
margin broadly arched; interior of area flattened;
inner marginal part of right valve shallowly
grooved to receive the margin of left valve;pallial line indistinct; cardinal tooth of rightvalve triangular, low, small; chondrophore small,thin, triangular, connected with posterior dorsal
margin.
Observation:-The apical angle is measured
120° -130° in the adult specimens. The surface
ornamentation is more or less weathered in most
specimens, but a specimen, KSG-K039B, shows
distinct concentric ribs on the surface.
Comparison:-This new species has smaller
cardinal tooth and chondrophore in the rightvalve than those of Pulsidis nagatoensis. This
new species is somewhat similar to Pulsidis
sanchuensis, sp. nov. in the feature of the ribs,
but differs from the latter having gradually486

Takeshi KOZAI
Measurements (in mm):Specimens
KSG-K042, R.
KSG-K038, L.in.m.
KSG-K039A, L.in.m.
KSG-K039B, R.
KSG-K045, R.
KSG-K044, R.
KSG-K046, L.in.m.
KSG-K04 7, R.
KSG-K048, R.
KSG-K049, L.

L
15.5
14.2
12.6
12.7
15.7

H
5.5
8.0
5.7
7.3
8.0

15.7
13.4
13.2
9.7+
11.6

8.9
7.1

narrowed posterior rostrum. This species is alsosimilar to "Pulsidis" higoensis (Matsumoto,1938), from the Goshyonoura Group of Kumamoto Prefecture, in view of the outline andsurface ornamentation, but has more crowded
concentric ribs. Pulsidis angulata, sp. nov. isalso clearly distinguishable from this new species
by distinct posterior carina.
Occurrence:- Rare at Locs. 7 and 9. Probably
Barremian.

Pulsidis rostrata, sp. nov.
PI. 95, Figs. 10-16.

Material:-Holotype, KSG-K026, externalmould of right valve from Loc. 6; 12 paratypes;
KSG-K028 and -K029, external moulds of rightvalves from Loc. 6; KSG-K030 and -K031
from Loc. 8 and KSG-K032 and -K033 from
Loc. 6, internal moulds of right valves; KSGK034, external mould of left valve from Loc.
6; KSG-K035 and -K036 from Loc. 6 and KSGK037 from Loc. 8, internal moulds of left valves;
KSG-K098, -K099 and other about 100 specimens from Locs. 6 and 8.
Diagnosis:- Shell medium-sized for genus, andcharacterized by a narrowly elongated andpointed posterior rostrum in right valve; densely
spaced concentric ribs on external surface anda distinct radial median groove on area.
Description:-Shell moderate in size, inequi-

6.6
5.0
5.9

B

2.0
2.2
1.9

2.1

U
5.0
5.2
5.5
5.5
6.2
5.1
4.1
4.5
3.8
5.0

H/L
0.35
0.56
0.45
0.57
0.51
0.58
0.53
0.50
0.510.51

U/L
0.32
0.37
0.44
0.43
0.39
0.33
0.38
0.34
0.390.43

valve, highly inequilateral, much longer thanhigh, elongate-trigonal, abruptly narrowed towards posterior part with long and graduallytapering rostrum; right valve larger than left;
test comparatively thin; anterior dorsal margin
short, slightly arcuate; posterior dorsal margin
broadly arched; ventral margin arched in anterior
part but slightly concave or nearly straight inposterior part; posterior end subvertically truncated; umbo situated at more anterior than
one-third of shell length from anterior extremity in right valve, but more posterior than
one-third from anterior extremity in left, slightly
prosogyrate, a little prominent; area with astrong radial median groove and also with regularly spaced concentric ribs; disk covered withregular concentric ribs being rounded at top;interior margin of right valve grooved to receive
the left valve; an internal radial ridge extends
from umbo to margin of inner area; pallial line
indistinctly impressed, curved with an acuteangle under posterior adductor scar; cardinal
tooth of right valve triangular, small; resilial
pit large, deep, located on posterior side ofcardinal tooth; chondrophore narrow, scarcely
separated from dorsal margin by a shallow
groove.
Observation:-The posterior rostrum is constantly long. The surface ornamentation is wellpreserved. The concentric ribs are very fine,numbering about 60 on the whole surface. About

821. Pulsidis from Japan

487

Measurements (in mm):Specimens
KSG-K026, R.ex.m.

L

H

B

U

H/L

U/L

7.9

3.0

1.3

2.8

0.38

0.35

KSG-K028, R.ex.m.

7.5

3.5

1.4

2.9

0.47

0.39

KSG-K029, R.ex.m.

9.9

4.8

1.5

2.9

0.48

0.29

KSG-K030, R.in.m.

9.5

4.3

3.8

0.45

0.40

KSG-K031, R.in.m.

10.8

5.3

3.9

0.49

0.36

KSG-K032, R.in.m.

7.7

3.9

2.5

0.51

0.32

KSG-K033, R.in.m.

8.8

2.9

3.0

0.33

0.34

KSG-K034, L.ex.m.

6.9

3.2

KSG-K035, L.in.m.

8.2

4.5

2.0

2.8

0.46

0.41

4.1

0.55

0.50

KSG-K036, L.in.m.

8.9

3.3

2.9

0.37

0.33

KSG-K037, L.in.m.

10.1

4.7

4.1

0.47

0.41

11 ribs are counted in the distance of 1 mm inheight near the ventral margin of adult specimens. The apical angle at the adult specimens
is 120°-130° in the right valve and 110°-120°
in the left. The angle in younger specimens isgenerally smaller, about 100°.
Comparison:-This new species is different
from Pulsidis angulata, sp. nov. in its finer concentric ribs, radial groove on the area, and lessdefinite posterior carina. The new species issimilar to "Pu Is idis " higoensis in the outline, but
has a median groove on the area. In the adultspecimens, the new species differs from Pulsidis
nagatoensis in its anterior location of umbo,
finer concentric ribs, distinct median radial
groove on much elongated area, but in theyounger specimens this new species is very
similar to Pulsidis nagatoensis in the subtrigonal
outline. For this reason, it could have been
derived from P. nagatoensis.
Occurrence:-Abundant at Locs. 6 and 8.
Probably Lower Aptian.
Acknowledgments
I wish to express my thanks to Professor
Masayuki Tashiro of Kochi University, for his
pertient instruction and reading the first draft.
Thanks are also due to Emeritus Professor
Tatsuro Matsumoto of Kyushu University, for

his critical reading the manuscript with encouragement, Emeritus Professor Yoshihisa Ohta(=Ota) of Fukuoka University of Education, for
his valuable suggestion, and Dr. Keisaku Tanaka
of former staff of Geological Survey of Japan,
Professor Itaru Hayami of the University ofTokyo, Mr. Masao Otsuka of Kogen Juniorhigh School in Kumamoto, and Dr. Tomoki
Kase of the National Science Museum for theirkind helps for this study.
This study was supported in part by Grant-inAid for Scientific Research from the Ministry of
Education, Science and Culture, project nos.
58916031 and 59916034.
References cited
Hase, A. (1960): The Late Mesozoic formations
and their molluscan fossils in west Chugoku
and Kyushu, Japan. Jour. Sci., Hiroshima
Univ., Ser. C, vol. 3, no. 2, p. 283-338, pis.
35-39.
Hayami, I. (1975): A systematic survey of the
Mesozoic Bivalvia from Japan. Bull. Univ.
Mus., Univ. Tokyo, no. 10, p. 1-249, pis.
1-10.
- - and Oji, T. (1980): Early Cretaceous Bivalvia
from the Choshi district, Chiba Prefecture,
Japan. Trans. Proc., Palaeont. Soc. Japan,
N. S., no. 120, p. 419-448, pis. 51-53.
Ishizaki, K. (1960): On the geology and tectonics

488

Takeshi KOZAI

in the area northeast of Kochi City, Shikoku.
Jour. Geol. Soc. Japan, vol. 66, p. 553-565(in Japanese with English abstract).
Matsukawa, M. (1979): Some problems on the
Cretaceous Shiroi Formation of the Sanchu"Graben", Kwanto Mountains, Japan. Jour.
Geol. Soc. Japan, vol. 85, no. 1, p. 1-9(in Japanese with English abstract).
- - , (1983): Stratigraphy and sedimentary environments of the Sanchu Cretaceous, Japan.
Mem. Ehime Univ., Sci. Ser. D (Earth Sci.),vol. 9, no. 4, p. 1-50, pIs. 1-2.
Matsumoto, T. (1938): Geology of Gosyonoura
Islands, Amakusa, with special reference to
the Cretaceous stratigraphy (in Japanese),with preliminary notes on some of the more
important fossils among the Gosyonoura
fauna (in English). Jour. Geol. Soc. Japan,
vol. 45, no. 532, p. 13-26, pIs. 1-2.
- - , Obata, 1., Tashiro, M., Ohta, Y., Tamura,M., Matsukawa, M. and Tanaka, H. (1982):
Correlation of marine and non-marine
formations in the Cretaceous of Japan.
Kaseki (Fossils), no. 31, p. 1-26 (in Japanese with English abstract).
Onuki, Y. and Mori, K. (1961): Geology of the
Ofuna to district, Iwate Prefecture, southernpart of the Kitakami Massif, Japan. Jour.

Geol. Soc. Japan, vol. 67, p. 641-654 (inJapanese with English abstract).
Ota, Y. (1964): On some Cretaceous corbulids
from Japan. Mem. Fac. Sci., Kyushu Univ.,
Ser. D, Geol., vol. 15, no. 1, p. 149-161,
pIs. 20-21.
-(Ohta, Y.) (1977): Stratigraphy of theLower Cretaceous Kawaguchi Formation in
Yatsushiro area, Kyushu, Japan. Bull. Fukuoka Univ. Educ., vol. 27, part 3, p. 45-61(in Japanese).
Tamura, M. (1959): Taxodonta and Isodonta
from the Upper Jurassic Sakamoto Formation in Central Kyushu, Japan. Trans. Proc.,
Palaeont. Soc. Japan, N. S., no. 34, p. 5365, pI. 6.
--(1977): Cenomanian bivalves from the
Mifune Group, Japan. Part. 2. Mem. Fac.
Educ. Kumamoto Univ., no. 26, p. 109144, pis. 1-13.
Tanaka, H., Kozai, T. and Tashiro, M. (1984):
Lower Cretaceous stratigraphy in the Monobe
area, Shikoku. Res. Rep., Kochi Univ., vol.
32, Nat. Sci., p. 215-223 (in Japanese withEnglish abstract).
Tashiro, M. and Kozai, T. (1984): Bivalve fossils
from the type Monobegawa Group (Part 1).
Ibid., vol. 32, Nat. Sci., p. 259-293, pis. 1-

Explanation of Plate 95
Figs. 1-4. Pulsidis nagatoensis Ota . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 482
Gum cast of external moulds (Fig. 1, KSG-K011, x3; Fig. 2, KSG-K012, x3; Fig. 3, KSGK004, x3). Gum cast of internal mould (Fig. 4, KSG-K002, x3).
Figs. 5-9. Pulsidis tashiroi, sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 483
Internal mould (Fig. 5, KSG-K039A, x2.5). Gum cast of internal mould (Fig. 6, KSG-K046,
x3). External moulds (Fig. 7, KSG-K042, x2.5; Fig. 8, KSG-K038, x3.5; Fig. 9, KSG-K039B,
x2.5).
Figs. 10-16. Pulsidis rostrata, sp. nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 486
Gum cast of external moulds (Fig. 10, KSG-K029, x4.5; Fig. 12, KSG-K026, x4; Fig. 13,
KSG-K098, x2.5; Fig. 16, KSG-K027, x2). Internal moulds (Fig. 11, KSG-K031, x3; Fig. 14,
KSG-K036, x2.5; Fig. 15, KSG-K099, x3).
Figs. 17-21. Pulsidis angulata, sp. nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 483
Gum cast of external moulds (Fig. 17, KSG-KOI5, x6; Fig. 18, KSG-K021, x3). Internalmoulds (Fig. 19, KSG-K022, x6; Fig. 20, KSG-K017, x3; Fig. 21, KSG-KOI4, x3).
Figs. 22-30. Pulsidis sanchuensis, sp. nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. 483
Gum cast of internal moulds (Fig. 22, KSG-K096, x7; Fig. 24, KSG-K088, x3.5; Fig. 25,
KSG-K089, x4; Fig. 26, KSG-K092, x3). External moulds (Fig. 23, KSG-K095, x3; Fig. 29,
KSG-K091, x4). Internal moulds (Fig. 27, KSG-K087, x3; Fig. 28, KSG-K090, x4; Fig. 30,
KSG-K086, x5).

KOZAI: Pulsidis from Japan

Plate 95

6

7

10

13
17

14

11

16
18

20

19
15
23
24

25
27

8

3

821. Pulsidis from Japan4.
- - , - - Okamura, M. and Katto, J. (1980):
A biostratigraphical study of the Lower
Cretaceous formations of Monobe area,
Kochi Prefecture, Japan. In, Taira, A. and489

Tashiro, M., eds., Geology and Paleontologyof the Shimanto Belt - Selected Papers in
Honor of Prof. Jiro Katto, p. 71-82, pIs.
10-11. Rinyakosaikai Press, Kochi (inJapanese with English abstract).

Eri :&~n, Funagawara ~;ililJl*, Hibihara El.LtW-, Hinagu B ~?., Kagahara tip 7" JJJt, Kasanokawa ¥i./ JII, Kawaguchi )IIIJ, Kawanouchi )IIO)I*J, Kumagawa );j(Jf:li)11.
Mamonozawa
FI31*,/.Jv\, Of una to :kliYdiilt, Ryoseki ~Ji7cI, Sakamoto W;;f>:, Sakayoriue W. 3 ~ 1:, Sebayashim1*", Shiroi (])t-. Tanoura millL Todoronotaki ,Ji./iO)r1ff, Yamabagoe U-J.H;;..t!!?, Yonenomoto*51;, Yoshimi ~J!, Yoshimo ~ro, Yunoki trtl/;;f.:.

El;;f>:0)111JJtIlE1iln~j£ Pulsidis ~I:'-?l'-C :::;h;l: -r~c.lI\t~;h -C ~t-: Pulsidis~"i:"fI}~
Hlt-:o .7c O)fiL!l?:::;h;l:-c' Pulsidis ~1:.ft"/,l)G;h-Cl't-: P. higoensis lkU P. okadai "i:"*A~/J· GIli7i- l, Wit-:i:' 4 Wi1lfi"i:"1I;tr 5 f.Jl:"i:"~c.lri1i: l t-:o .7c;h G Ii Pulsidis nagatoensis Ota,

P. angulata, sp. nov., P. sanchuensis, sp. nov., P. tashiroi, sp. nov., P. rostrata, sp.

nov.

-c';l;~o

iltl§

ji'\

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 144, pp. 490-515, December 30, 1986

822. PAL YNOLOGICAL STUDY OF 200-METER CORE SAMPLESFROM LAKE BIWA, CENTRAL JAPAN
II: THE PALAEOVEGETATIONAL AND PALAEOCLIMATIC
CHANGES DURING THE CA. 250,000 -100,000 YEARS B. P.*
NORIO FUJI
Department of Earth Science, Faculty of Education,
Kanazawa University, Kanazawa 920, Japan

Abstract. The writer had stated an outline of the palaeovegetational andpalaeoclimatic changes during the last 600,000 years from the point of view ofpalynological investigation of samples taken at intervals of 5 m throughout a 200meter core drilled at the lake bottom 65 m of water depth in Lake Biwa, Japan.
The writer further treated about 200 samples taken at intervals of 25 cm over about
50 m between 55 m- to 110 m-horizons of the 200-meter long core for analyses ofthe detailed vegetational and climatic changes in and around the lake during about
150,000 years between about 250,000 years B.P. and 100,000 years B.P.
On the basis of the pollen diagrams, the lacustrine deposits during the time
250,000 to 100,000 years B.P. are divided palynologically into four pollen zones.
In the global correlation between the palaeoclimate from Lake Biwa, other terrestrial records from Central Europe, the sea level changes from Southern Kanto
in Japan and Western Mediterranean, and the oxygen isotope records from the
Caribbean Sea and Equatorial Pacific etc., the writer found a remarkably noticeablesimilarity between major trends from them.

1972, 1977). The climatic curve and ages ascertainable from Lake Biwa were found to display
a similarity to the palaeotemperature curve(oxygen isotope ratio determination) from the
Caribbean Sea (Emiliani & Shackleton, 1974)
and the Equatorial Pacific (Shackleton &Opdyke, 1976, 1977), and to the environmental
changes (loess, palaeosol and gastropod faunal
records) from Central Europe (Kukla, 1970,

Introduction
The present writer had reported on outlineof the palaeovegetational and palaeoclimaticchanges during the last about 600,000 years
based on the palynological analyses of samples
taken at intervals of about 5 meters throughout
a 200-meter core obtained from Lake Biwa,
Central Japan (Fuji, 1973, 1983, Fuji & Horie,*: Received October 7 1985; revised manuscript received October 11, 1986; Read June 18, 1977 at
Shizuoka University and June 24, 1984 at Akita University.
Contribution from the Department of Earth Science, Faculty of Education, Kanazawa University;
New Series No. 119.
490

822. Palaeovegetation and palaeoclimate, Lake Biwa

1975).
The writer further treated about 200 samples
taken at intervals of 25 cm throughout about 50
meters from 55 meters to 110 meters-horizons in
depth of the 200-meter core for analyses of thedetailed vegetational and climatic changes around
Lake Biwa in the period, about 150,000 years,
between about 250,000 years B.P. and about
100,000 years B.P.
In this article, the vegetational and climaticchanges ascertainable from Lake Biwa in the
period between about 250,000 years ago andabout 100,000 years ago will be discussed fromthe view point of palynological analyses.
Additionally the topography, geology, vegetation, and climate around Lake Biwa, and detailed
description on the method for reconstruction ofpalaeoclimate were already given in the writer's
paper (Fuji, 1983).
Acknowledgments
I am greatly indebted to Emeritus Prof.
Herbert E. Wright Jr. of the LimnologicalResearch Centre, University of Minnesota,
Minneapolis, U.S.A., and to Prof. Shoji Horie ofthe Institute of Palaeolimnology and Palaeoenvironment on Lake Biwa, Kyoto University for
their kind advice and for the helpful critical reading of the manuscript. I am indeed grateful forthe support and encouragement of Emeritus
Prof. Teiichi Kobayashi of University of Tokyo
during this investigation. I also acknowledge
the help of the member of the Lake Biwa Palaeolimnological Research Group, especially thelate Prof. Naoto Kawai of Osaka University,
Prof. Takuo Yokoyama of Doshisha University,
Prof. Sei'ichi Kanari of Hokkaido University,and Prof. Nobuyuki Nakai of Nagoya University. My investigation was possible through a
Scientific Grant from the Ministry of Education,
Japanese Government for fifteen years (1971 to1985) and a help of University of Minnesota,
U.S.A.

491

Palynological investigation(A) Locality of the boring and samples
A 200-meter core used for the present palynological analyses was drilled in 1971 from thebottom of Lake Biwa 65 meters below the present
lake water level. The core samples are composed
mainly of dark greenish blue soft homogeneous
muddy clay with about 30 thin volcanic ash
layers. About 200 samples taken at intervals of
about 25 cm from 55 meters to 110 meters
below the present lake bottom were analyzed.
The samples taken from the same horizons wereanalyzed for geochemical and organic chemicalcomponents, fossil diatom and microanimals, andalso from the view points of palaeomagnetismand granulometric analyses.
(B) Preparation and examination for pollen
analyses
The samples were treated by the HF-KOHacetolysis method, and then treated by a saturated solution of ZnCI 2 •The pollen grains and spores mounted onslides for a microscopic observation were identi-

Siber1Q

ChH\(]

c-r:.

.,

Pacific OceanI,d()- .,
Formosa

12~

I

\\

\

'I
~24'
---+---I---\-,---T,-1360

140'"

\ \ \

-'2!-,
••

IZSO

1320

500

Text-fig.!.
studied area.

Locality!44°

I,OOOkmmap

showingtheNorio FUJI

492

Japan Sea
WAKASA BAY~+

0,.....-_~_~_....::.30km'<---'---'~-35"N

I~
~

plain

bkX)l m r. crea
tectonIC line

Pacific Ocean
136"EQ): Mr. Hira, @: Mt. Hiei, @: Mt. Gozaisho, @: Mt. Fujihara-dake, @: Mt. Ibuki, @: Mt. Mikuniga-dake, (j): Mt. Mikuni-dake, CD: Kaburaki tectonic line, @ : Yanagase tectonic line, @: Suzuka
tectonic line, x : boring locality.
Text-fig. 2.

Topographic map of the Lake Biwa area, Central Japan (after Fuji, 1983).

fied and counted by use of a mechanical stage ofa microscope (Olympus Nomarsky-type tricular
microscope). The counting of pollen grains wascontinued until more than 1,000 arboreal pollen
grains. The identification of pollen grains was
conducted with the aid of a reference collectionof about 500 slides of important trees, shrubs,
and acquatic herbs of the Japanese Islands inpossession of the Department of Earth Science,
Kanazawa University, and also a reference collection of about 9,000 slides kept in the Limnological Research Centre, University of Minnesota,

Minneapolis, the United States of America.
(C) Construction of the diagrams
Shown from the left to the right in the pollen
diagrams are sediment stratigraphy and lithofacies, depth in meter, spectrum number, fissiontrack dates, palaeomagnetic stratigraphy, pollen
zoning, palaeoclimate, summary diagram, pollen
profiles of each taxon and assemblage zones. The
pollen sum was used on the pollen types concerned on the local situation and on the kind of
deposits.
Percentages in the diagrams of this article are822. Palaeovegetation and palaeoclimate, Lake Biwa

shown on two scales; the scale with 10 x exaggeration permits the accurate plotting of minor
curves and minor fluctuations. The summary
diagram is composed of changes in the ratio
between total AP and total NAP, and of the percentage of boreal type plants (Polar - Subpolar
plants), Cool Temperate plants, Cool Temperate- Temperate plants, plants of the middle area of
the Cool Temperate - Warm Temperate zone,
and plants of the southern area of the Temperate- Subtropical zone, calculated on the basis of
the warmth index (month-degress) as describedin a later section.
(D) Method for interpretation of palaeovegetation and climatic history
The writer depends upon the pollen analyses
for the reconstruction of vegetation on the basisof the pollen spectra obtained from the 200meter core samples. He has employed two
methods, namely, (1) pollen spectra of themodern samples collected from Lake Biwa andits vicinity, and also from various localities ofsome climatic zones throughout the JapaneseIslands, and (2) the warmth index (monthdegrees). With regard to this index, the detailed
descriptions were already given (Fuji, 1983).
(E) Dating and sedimentation rates
The dating of the core samples in the representative depth of the core gives important anduseful information. The determination of the age
for the upper 11.5 meters of the core obtained
from Lake Biwa has been carried (Horie et ai.,1971). In the recent study of the 200-meter coresamples, Nishimura and Yokohama (1975)
present the invaluable data concerning the agedetermination of the deeper horizon of the coreby the fission-track method. These results enable
the present writer to determine the date of theboundary between the glacial and interglacialages, or between the stadial and interstadial, orof characteristic events in a climatic fluctuationof the ancient Lake Biwa.
The age was determined by applying a suitable
curve-fitting method to the basic data obtained
from the 14 C and fission-track methods as shownin Table 1. According to Kanari's calculation(Kanari & Takanoya, 1975), the absolute age at

200m

493

UlOOm

o -r-_ _~,:,:bo::Inng:-::'=:-,-bo,rlng....:......-_~

BB~......

t:
-:

I

I

BB425/ Ka
liBiSJ i('--- --!

db. 161
O(J(il(47)

O(J(i
(46

OBi (70)

10.,3,• (Azl (87l. db. 365

100

.

Jl

200

-::;

t.:
11

c3

0 Mr (210)0 Kn3 (220)o Kn2 (235)

.5
0.;

a:i

.

~

~

tL

o Sa

(290)

300
OM. (310)):
Year B.P. in lO·(Fission·trackl(.0 : main volcanic ash
Ka

Ki
Bi
Az
Mr
Kn3
Kn2
Sa
M.

: Kamiogi: Koromogawa: Biotite: Azuki: Murasaki: Konom: Kono [: Sagami: Magata

500

Text-fig. 3. Chronological correlation among
the Quaternary deposits in and around Lake
Biwa (after Fuji, 1983).

the horizon of 200 meters below the bottom surface is 565,000 years B.P., which is not inconsistent with the stratigraphic data and the results
inferred from the palaeomagnetism and palaeoclimate on the basis of the correlation between
the writer's previous curve from Lake Biwa
(samples from 5 meters interval) (Fuji, 1983)and Emiliani's generalized temperature curve(Emiliani & Shackleton, 1974) for the Caribbean
Sea for the last about 700,000 years.
In 1973, a new method of age determination

494

Norio FUJI

was developed for the core under the conditionof a constant sedimentation rate (Yamamoto etaZ., 1973). The age of the respective layer of the
200-meter core is estimated for the constant sedimentation rate of 0.354 mm/yr., which corresponds well to the observed sedimentation rate
at the present (Toyoda et aZ., 1968).
(F) Zoning of pollen assemblage
In order to facilitate description of pollen
assemblages and their discussion, the pollen diagrams are divided into zones and subzones. Thesezones and subzones are based upon conspicuouschanges in pollen percentages. Changes in theratio, Total AP/Total NAP, can be described,
while alternation of the values of one or twopollen types may also lead to the establishmentof pollen zones. Where pollen zones display thesame nature characterized by assemblage ofpollen grains and spores, the present writer has
attempted to indicate them under the same lettercode in order to facilitate comparison. These
pollen zones are restricted to the series of diagrams under discussion. Minor differences of thepollen assemblage in the zones lead to determining subzones.
According to the present writer's analyses,
the deposits are divided into four pollen zones,
Y-1, Y-2, Y-3, and Y-4 in an ascending order, andthe Y-2 pollen zone is classified into six pollen
subzones by minor differences of pollen assemblage and their characters.
(1) Pollen Zone Y-1, depth: 108 - 97 m;
240,000 - 200,000 yr.B.P.
This zone is characterized by a large value of
boreal conifers such as Abies, Picea, Tsuga, Larix
and Pinus hapZoxyZon-type. These conifers reachto 76 - 99% (about 90% on average). The characteristic plants growing only in the Subpolar or
Subalpine zone (warmth index in month degrees:
15 - 55°) show the value of about 5% on average,
though the plants range from 0.2 to 0.7% in the

'"

OJ

'"

"Cl

e

c.

U1

CJ

OJ

55

50

57

rZla
129.1

IJla

58

I3Sa

59

1393

'"

1'30

62

152,

"

63

...
55

66

..
67

II X 10' B.P.

IS,.

x

'"

''''

o

~

I

16Sa

x

167a

16"
rHa

10

17,.

'"
~

I7Sa

12
73

l'

1170
179.1

75

181a

16

18,.

n

18Sa

78
19
80

II

"

187a

18,.
191a

19,.
I9Sa

..

197a

as

1993

86

201,

87

20"
205a
207,

17XI0'B.P.

83

88

89
90
91

92
93

"

95

96
91

"

99

2:11a

21,.
215.
217a
21901

=

22Sb

""
",.

",.

no. 18X la' B.P.

100
101

102
103

10'
IDS

Text-fig. 4. Stratigraphy, spectrum nos.,fission-track data, palaeomagnetism, pollen zoning, and palaeoclimatic change of the 55 to 110
meters samples from a 200-meter core of Lake
Biwa. Arrow part: climatic cord no. t-5

106

101
108
109

110

c::::J ash
c:::J clay

>.

o

IS,.

69

11

~

lAta

25Sa
ZS7a

259.
26101
2.,.

27X la' B.P.

822. Palaeovegetation and palaeoclimate, Lake Biwa
other pollen zones. In contrast to them, Pinus
diploxylon-type, which includes the temperate
Pinus thunbergii and P. densiflora, shows a
low percentage. In this zone, trees such as Ulmus,
Betula and Fagus large type which grow in the
Cool Temperate zone, have the lowest value(average 1.5%) throughout all these pollen zones.
The same phenomena are recognized in the frequency of plants (pollen grains) which growin the Cool Temperate and Temperate zones, inthe middle area of the Cool Temperate andTemperate zones, and in the southern area of theTemperate and Subtropical zones. The main
component of the Y-1 pollen zone is Abies (55 86%), Tsuga (10-27%), Picea (5-15%), andPinus haploxylon-type (4 - 8%).
(2) Pollen Zone Y-2, depth: 97 - 65 m;
200,000 - 126,000 yr.B.P.
The Y-2 pollen zone is characterized by theabundance of plants which grew in the Cool
Temperate zone, and in the Cool Temperate andTemperate zones. However, on the frequency
throughout all this zone, plants adapted to theclimate of the southern part of the Temperate
and Subtropical zones are not always abundant.
Boreal conifers such as Abies, Picea, Tsuga, andLarix are few throughout this zone.
On the basis of fluctuations in the pollen
values of boreal conifers and minor changes ofother types, this zone is divided into six pollen
subzones, Y-2-a, Y-2-b, Y-2-c, Y-2-d, Y-2-e, andY-2-f.
Y-2-a Pollen Subzone, depth: 97-93 m;
200,000 -193, 000 yr.B.P.
This pollen subzone is characterized by a
drastic change of pollen assemblage; accordingly,
boreal conifers including Subpolar plants such as
Pinus haploxylon-type show an abrupt decrease(lowering to 52% from 90%) in comparison with
the Y-1 pollen zone. In contrast to them, the pollen frequency of plants which thrive in the Cool
Temperate and Temperate zones shows a higher
value than that of the Y-1 pollen zone, although
the relative frequency of plants which grow inthe southern part of the Temperate and Subtropical zones do not increase as much as that of
the Cool Temperate plant. This pollen assembl-495

age is shown as follows: Subpolar or Subalpine
plants (only conifers) 0.9%, boreal conifers 52%,
Cool Temperate plants 5.5%, Cool Temperate Temperate plants 0.9%; the main component isAbies (23-51%)-Cryptomeria (6-28%)-Pinus diploxylon-type (12-14%)-Tsuga (9-13%)-Alnus(9%)-Fagus small type (7%)-Lepidobalanus (37%).
Y-2-b Pollen Subzone, depth: 93-90 m;
193,000 - 185,000 yr.B.P.
The main component of this pollen subzone is
Pinus diploxylon-type (13-37%)-Abies (1536%)-Cryp to me ria (9-29%)-Tsuga (26%)-Lepidobalanus (10-15%)-Fagus large type (416%). This pollen assemblage is as follows:
Subpolar or Subalpine plants 0.3-3% (average
1.2%), boreal conifers 31-64% (average 49%),
Cool Temperate plants 2-16% (average 6.2%),
Cool Temperate-Temperate plants 12-32%(average 18%), middle area of the Cool Temperate-Temperate plants 17-47% (average 27%),and south area of the Temperate-Subtropicalplants 0.2-0.8% (average 0.5%). As mentioned
above this subzone is characterized by a little
increase of plants of the Cool Temperate zone
and a decrease of boreal conifers, especially thefrequency of plants of the Cool Temperate zone
has the highest value throughout the samples.
Y-2-c Pollen Subzone, depth: 90-82 m;
185,000 -166,000 yr.B.P.
This pollen subzone is characterized by thesmallest value of Subpolar or Subalpine plants
throughout all the samples treated at the present
and, in contrast to the above-mentioned phenomenon, by the largest value of plants thriving in
the Cool Temperate-Temperate zones over all
the samples. The pollen assemblage of this pollen
subzone is shown as follows: Subpolar plants 0 1% (average 0.2%), boreal conifers 16-70%(average 44%), Cool Temperate plants 0-1.9%(average 0.6%), plants in the Cool Temperate Temperate 14-69% (average 39%), plants growing in the middle area of the Cool Temperate andTemperate zones 7-36% (average 16%), plants
thriving in the southern area of the Temperate
and Subtropical zones 0-3.3% (average 0.8%).
The main element of this pollen subzone isNorio FUJI

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Pucivc:arpus

Summary diagram of the 55 to 110 meters samples from the 200-meter core.

822. Palaeovegetation and palaeoclimate, Lake Biwa

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Text-fig. 6.

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Conifers pollen diagram of the 55 to 110 meters samples from the 200-meter core.

Norio FUJI

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so%

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Conifers and deciduous broad leaved plant pollen diagram from the 200-meter core.

822. Palaeovegetation and palaeoclimate, Lake Biwa

....

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Broad leaved plant pollen diagram from the 200-meter core (1).

Norio FUJI

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Broad leaved plant pollen diagram from the 200-meter core (2).

822. Palaeovegetation and palaeoclimate, Lake Biwa
Cryptomeria (21-69%) - Abies (19-46%)Pinus diploxylon-type (5-22%)-Tsuga (2-16%) - Lepidobalanus (4%) - Sciadopitys (25%).
Y-2-d Pollen Subzone, depth: 82-79 m;
166,000 - 158,000 yr.B.P.
This pollen subzone is characterized by thelargest amount of plants thriving in the southernpart of the Temperate - Subtropical zones. The
percentage of Podocarpus ranges from 0.3 to
1.2% (average 0.3%). The main component of
this pollen subzone is Cryptomeria (17-46%)Abies (22-43%) - Pinus diploxylon-type (1820%) - Tsuga (10-19%); and also, Subpolar or
Subalpine zone 0-3% (average 0.9%), boreal
conifers 30-63% (average 42%), Cool Temperate plants 0.3-3% (average 1.0%), Cool Temperate - Temperate plants 16-48% (average 29%),
plants in the middle part of the Cool Temperate- Temperate zones 15-31% (average 26%), andplants in the southern part of the Temperate Subtropical zones 0.6-9.6% (average 4.5%).
Y-2-e Pollen Subzone, depth: 79-73 m;
185,000 -144,000 yr.B.P.
In this pollen subzone, a decrease of plants inthe southern part of the Temperate - Subtropical zones is matched by a similar increase in thevalue of boreal conifers. The pollen assemblage
is shown by boreal conifers 32-83% (average54%), Subpolar plants 0-1% (average 0.3%),
Cool Temperate plants 0-2.4% (average 0.7%),
plants in the Cool Temperate - Temperate zones
5-48% (average 21%), plants in the middle partof the Cool Temperate - Temperate zones 949% (average 27%), and plants growing in thesouthern part of the Temperate - Subtropicalzones 0-2% (average 0.5%). The main componentof this pollen subzone is Abies (24-70%), Pinus
diploxylon-type (10-35%), Cryptomeria (1427%), and Sciadopitys (about 10%). Besides
these conifers, Sciadopitys is one of evergreen
conifers and grows only in the recent montane
areas of the Kii Peninsula, Shikoku and Kyushu
regions, and also this pollen subzone is characterized by a higher value of this plant.
Y-2-f Pollen Subzone, depth: 73-65 m;
144,000 -126,000 yr.B.P.

501

This pollen subzone is characterized by adecrease of boreal conifers. This decrease ismatched by an increase of plants of the Cool
Temperate - Temperate zones. The pollen assemblage of this pollen subzone is shown by
Cryptomeria (12-46%)-Abies (14-45%)
Sciadopitys (11-29%).
(3) Pollen Zone Y-3, depth: 65 - 59 m;
126,000 - 113,000 yr.B.P.
This pollen zone is characterized by a drastic
increase of boreal conifers (up to about 80%) and
Subpolar plants (up to 2%), and by a decrease ofplants of the Cool Temperate - Temperate zones
and of the middle area of the Cool Temperate Temperate zones. The pollen assemblage isshown by Subpolar or Subalpine plants 0.8 - 5%
(average 2%), boreal conifers 57-88% (average80%), Cool Temperate plants 0-2% (average0.6%), plants of the Cool Temperate - Temperate zones 0.4-7.5% (average 0.2%), plants inthe middle area of the Cool Temperate - Warm
Temperate zones 6-19% (average 11%), andplants in the southern Temperate - Subtropicalzone 0-0.4% (average 0.1%). The main component is Abies (average about 55%)-PiceaTsuga-Cryptomeria.
(4) Pollen Zone Y·4, depth: 59-55 m;
113,000-104,000 yr.B.P.
In this pollen zone, a drastic increase of plants
growing in the Cool Temperate - Temperatezones and in the middle area of the Cool Temperate-Warm Temperate zones is matched by an
abrupt decrease of boreal conifers and Subpolar
plants. The pollen assemblage is' shown as
follows: Subalpine or Subpolar plants 0-4%
(average 1.5%), boreal conifers 34-81% (average 57%), plants of the Cool Temperate zone 0 1.3% (average 0.3%), plants in the Cool Temperate-Temperate zones 2-29% (average 12%),
plants in the middle area of the Cool Temperate-Warm Temperate zones 13-59% (average29%), and plants in the southern TemperateSubtropical zones 0-2% (average 0.8%).

Norio FUJI

502
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Non-arboreal pollen digaram from the 200-meter core (1).

822. Palaeovegetation and palaeoclimate, Lake Biwa

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Non arboreal pollen diagram from the 200-meter core (2).

503

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Text-fig. 12.

20

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30

Non-arboreal pollen diagram from the 200-meter core (3).

822. Palaeovegetation and palaeoclimate, Lake Biwa
Discussion on the interpretation of vegetationand climatic history
Pollen Zone Y-I is characterized by a highpercentage of boreal conifers. The maximum
value for Pinus haploxylon-type, one of therepresentative plants of the Subpolar or Subalpine zone in the Japanese Island, reaches toabout 5% on average in this zone, which is thehighest value over all the samples. In contrast,
trees growing in the other climatic zones are notabundant. In addition, boreal conifers reach tothe highest value of 76-99% (average 90%).
Judging from such pollen assemblages and frequencies of the other pollen zones, the climate atthat time of this zone may have been as cold asthat in the northern part of the Cool Temperatezone. This cold climatic age is correlated with the"c-6" in the present writer's previous paper (Fuji,
1983).
Pollen Subzone Y-2-a is characterized generally by a drastic decrease (low to 50% from 90%)of boreal conifers, and by a high percentage ofplants in the Cool Temperate - Temperate zones
in comparison with the Y-1 Pollen Zone. Abies,
Cryptomeria, Pinus diploxylon-type and Tsuga
are the main component in this pollen subzone. The climate at the time of Pollen Subzone Y-2-a was perhaps similar to that of theCool Temperate-Temperate zones. Therefore,
the climate at that time may have been corresponded to that of the Cool Temperate - Temperate zones.
Pollen Subzone Y-2-b is featured by the abundance of plants growing in the Cool Temperatezone and in the middle area of the Cool Temperate from the view point of palaeovegetation. The
frequency of Podocarpus pollen is more thanthose of Pollen Subzone Y-2-a and Pollen Zone
Y-1. Accordingly, the climate at that time may
have been as warm as that of Pollen Subzone
Y-2-a.
Pollen Subzone Y-2-c is characterized by thesmallest value of Subpolar-type plants and thelargest value of plants thriving in the Cool Temperate - Temperate zones. In addition, plants
in the Cool Temperate zone show a drastic

505

decrease. Judging from these data, the climate atthat time may have perhaps been similar to thatof the southern part of the Cool Temperate Warm Temperate zones.
In the Y-2-d Pollen Subzone, plants in thesouthern area of the Temperate - Warm Temperate zones have the highest value (average 4.5%),
and, in contrast, plants thriving in the Subpolar
and the Cool Temperate zones show a smaller
value. Therefore, the climate at the time ofPollen Subzone Y-2-d was perhaps that of thesouthern area of the Temperate - Warm Temperate zones.
Pollen Subzone Y-2-e is characterized by adecrease of plants in the southern area of theTemperate - Warm Temperate zones and by asimilar increase in the value of boreal conifers.
Therefore, the climate at the time of Subzone
Y-2-e may have been that of the middle area of
the Cool Temperate zone.
Pollen Subzone Y-2-f is characterized by adecrease of boreal conifers and, in contrast to
this phenomenon, plants of the Cool Temperate- Temperate zones and of the middle area of
the Cool Temperate - Warm Temperate zones
decreased. Namely, the climate at the time ofthis pollen subzone may have been similar to thatof the northern area of the Temperate zone. As
mentioned above, from the view point of thepalaeoclimate, Pollen Zone Y-2 may perhaps be
one of the interglacial age, and is divided into six
subzones. The palaeoclimate at the time ofPollen Zone Y-2 was clearly warmer than thoseof Pollen Zones Y-1 and Y-3. Most of this pollenzone correspond to the "t-3" mentioned in thepresent writer's previous paper (Fuji, 1983).
Pollen Zone Y-3 is characterized by a drastic
increase of boreal conifers and by an increase ofplants of the Cool Temperate - Temperate zones
and of the middle area of the Cool Temperate Warm Temperate zones. Therefore, the climate
at the time of this pollen zone may have correspond to a climate of the northern area of theCool Temperate zone and this time was perhaps
one of the glacial age. This pollen zone is correlated with the "c-4" of the present writer's previous paper (Fuji, 1983).

NorioFUJI

506

Pollen Zone Y-4 is characterized by a drastic
increase of plants in the Cool Temperate - Temperate zones and in the middle area of the Cool
Temperate - Warm Temperate zones and by an
abrupt decrease of boreal conifer trees. The
climate at that time may have been similar tothat of the middle area of the Cool Temperatezone or that of the northern area of the Temperate zone.

ing a layer of thickness dS with bulk density
PBO. During these t years, the layer has experi-

enced contractions of A per unit thickness
through consolidation, so that dr = dS (1 - A).
Since the contraction is due to the squeezing ofwater, the equation of mass conservation then isPB (1 - A) = PBO - pA, where P is the densityof water. Thus, the integration of dS over the lastt years, that is, the total thickness if not yet consolidated, isData for "time-stratigraphy"
Ages of boundaries between the pollen zones
can be estimated by some means such as 14C,
fission-track, sedimentary rate, and palaeomagnetic stratigraphy etc. (Horie, 1984).
In regard with radioactive age of an upper partof the 200-meter core, the following data are
obtained by Isotopes A Telddyne Co. (Horie etal., 1971).

0.8±0.05 m in depth . .. 1,430±95 yr.B.P.
4.5±0.15 m in depth
3,650±105 yr.B.P.
11.5±0.2 m in depth .. 14,980±460 yr.B.P.
Within the core samples some volcanic ash layers are intercalated. They are at least about forty
layers, from which Nishimura and Yokoyama(1975) could successfully obtain the following
ages by fission-track method as follows:
37 m in depth ......... 8x104 yr.B.P.
62 m in depth ......... 11x104 yr.B.P.
82 m in depth ......... 17x104 yr.B.P.
99 m in depth ......... 18x104 yr.B.P.
110 m in depth ......... 27x104 yr.B.P.
181 m in depth ......... 46x10 4 yr.B.P.
In a chronostratigraphic investigation of thecore samples, it is necessary to establish therationale of interpolating between the discretevalues of absolute dates. A depositional layer,
which once deposited on the surface of the lakebottom, begins to contrast its thickness throughthe consolidation process under the cumulative
load of deposits having a unit section, a depositional layer of thickness dr with bulk density PB,
which is situated at the depth r beneath the bottom surface at present, is originally that which
had deposited on the surface t years ago, form-S=

J -- J
t

dr

o I-A

P -P

t

=

B
0 PBO -

P

dr ....... (1)While, assuming the sedimentation rate h(t) to beconstant (= h o),S=J:h (t)dt = hot ................. (2)Hence, equating (2) to (1) yields:
t =

(1

)

ho PBO - P

Jt (PB - p)df...... (3)0Equation (3) ensures the age-scaling along thecore sample, when the vertical profile of bulk
density and reference ages are given. According
to Equation (3), a preliminary age-scale for the
200-meter core was composed in reference to the
14C (Horie et al., 1971) and fission-track dates(Nishimura et al., 1975). The age-scale is empirically modified as shown in Text-fig. 13 (Yamamoto, 1976). On the basis of the age-scale figure(Text-fig. 13), ages of every meter of the 200meter core are shown in Table 1 (Yamamoto &Higashihara, 1978).
According to the palaeomagnetic stratigraphy(Horie, 1984), the polality epoch of the 200meter core belongs to the Brunhes normal polality epoch since ca. 0.7 million years ago. Within
the core, there appear four magnetic anomalies.
The age of the youngest event (50 to 55 m indepth of the 200-meter core) in the four magnetic anomaly events is 100,000 yr.B.P., and so close
to the age of the Blake event, which was discovered from the deposits collected from theAtlantic and Caribbean Sea, and of which age

822. Palaeovegetation and palaeoclimate, Lake Biwa

10 5

a:
co
....III
Cl

CII

>.
lJJ
(!)

<

10'

10J~----L-----~----~----~_____

o

100

200

DEPTH(m)Text-fig. 13. An empirically modified agescale (solid line), with nine reference ages including three 1 4 C dates (solid circle) and six fissiontrack dates (open circle) (After Yamamoto,1976).

was found to be between 114,000 and 108,000
yr.B.P.
Correlation between the deep-sea and terrestrial
records, and discussion
The International Subcommission on Stratigraphic Classification recognized generally threeprincipal categories of stratigraphic subdivision
for use through the geologic column. These categories are lithostratigraphy, biostratigraphy, andchronostratigraphy. However, in regard with the
Quaternary stratigraphy, these are imperfect.
The Quaternary possesses rather special features that make it unique in the geologic record.
Therefore, there is a growing consensus that thesubdivisions of the most recent part of the strati-

507

graphic record should follow the geologic procedures, and two further categories have been employed. These are morphostratigraphy and climatostratigraphy.
Climatostratigraphy is undoubtedly usefulconcept and, insofar as the Quaternary at midand high-latitudes tends to subdivided into glacials and interglacials.
In the deep-seas, long sequences of relatively
undisturbed sedimens are preserved records
which frequently extend back beyond the beginning of the Quaternary. Within these sediments,
the microfauna contains a record of changing
oxygen isotope ratios which do not only provide
evidence for the past glacial and interglacial oscillations, but also form the basis for stratigraphic
subdivision and long distance correlation. Sediments from deep-seas hold a number of advantage over terrestrial sequences from the point ofview of stratigraphic subdivision and correlation
throughout the world as follows:(1) The records are more commonly continuous and relatively undisturbed.
(2) A common technique such as oxygenisotope analysis can be used.
(3) As the oxygen isotopic changes are a consequence of climatic changes, they areused for climatostratigraphy.
(4) The sedimentary records can be dated andcorrelated by the independent method ofpalaeomagnetic stratigraphy.
However, if terrestrial sequences provide thefollowing certain characteristic features, correlation betwen the marine and terrestrial successions can be established.
(1) A lengthy stratigraphic record must be
available.
(2) Within terrestrial sequences, evidence ofclimatic change must be clear and unequivocal.
(3) The record of sedimentation must becontinuous.
(4) The terrestrial sequences can be dated bypalaeomagnetic or radiometric method.
The most impressive and important example
of lake sediment sequences as one of standards
for the world-wide correlation of terrestrial

508

Norio FUJI

Table 1

An empirically modified age-scale (After Yamamoto, 1976)
Age

Depth

Depth

Age
(10'yr. B. P.)

:;

(m)

Age
t
(10'yr. B. P.)

Depth

~
(m)

(m)

(lO'yr. B. P.)(m)(lO'yr. B. P.)

t

Depth

(

~

t

Age
.t

1

1. 39

51

95.49

101

214.26151

365.07

2
3

2. ~4

52

97.65

102

216. 98152

368.29

3.24

53

99.82

103

219.73153

371. 54

4

4.31

54

101. 98104

222.44154374.75

5

5.47

55

104.17105

224. 58)155

378.00

6

6.77

56

106. 32106228. GO

381. 25

7

8.09

57

108.50107

230.77

156
157

8

9.50

58

110.69108233.58158

387.69

9

10.98

59

:12.90109

236.39159

390.96

10

12.54

60

115.07110

239.23160

394.19

11

14.14

61

117.28

III

242. 06

161

397.42

12

15.82

62

119.49112244.93162

400.6913

17.48

121. 73113

247.80163

403.91

14

19.22

63
64

123.96114

250.70164407. 1715
16

21. 0422.84

65126. 19115

128. 42

17

24.72

116
117

165
166

410.426667

253.60256.49259.42167

416.89

18

26.59

168

420. 141920
21384.45413.6868130.69132. 95

118

262. 3828.49

69

135.20119265.34169

423.3830.43

70

137.45120

268.29170426.6332.40

71139. 74121271. 28429.90

22

34.36

72

142.06122274.27

23

36.39

73

144.34123

277.28

171
172
173

24
25

38.37

74

146. 66124

280.30174

439.61

40.43
42.48

75

149.01125151. 36
153.70126

283.31286.36175
176

442.8476
77

127

289.40177

449.3278

156.08158.46

128
129
130

292.47295.51

178
179

298.61

131

180
181

452.55455.80459.02

26
27
28
29

44.52
46.59
48.65
50.75

79
80

52.34
54.93

81
32

33
34

57.04

35

61. 25

36
37

63.39
65.52

38

67.64

39
40

69.76

88
89

71. 8790

185.48H

74.02
76. 1591
92

188.01190.57

44

78.29
80.42

93
94

193. 12
195.72

45
46

82.58
84.74

95

198.34195

200.97145
146

345.80349.00

196

47
49
50

91. 1893. 31203. 59
206. 24208.89

147
148
149

352.20355.43358.62

197

48

86.89
89.04

96
97

211. 58ISO

361. 85

30
31
32

42
43

59. 15160. 83
163. 24
165.64132

301. 68304. 7883
84

168.08133

307.91

170.52

134

85
86 -

172.99175.45

135

87

98
99
100

433.14436.37446. 10462.24182465.49183468.70311. 00

184

471. 94314.13185475. 15136

317.29177.92180. 42137
138

320.41323.56186
187

478.39481. 59IS8

182.95139
140

326. 75484.80488.03141

333.07142
143

336.25339. 43
342. 63144

329.89

.,

la9
190
191
192
193
194

198
199
200

491. 23494.45497.65500.87504.06507.28510.47513.68516.86520.08523. '25

822. Palaeovegetation and palaeoclimate, Lake Biwa
record is the record from Lake Biwa, Japan(Lowe & Walter, 1984). Namely, lacustrine sediments obtained from Lake Biwa accept four indispensable conditions as above-mentioned. In
addition, although sediments of both cold andwarm times are complete within sediments fromthe deep-seas, we cannot evaluate changes in climate over short times because of the extremely
small rate of sedimentation in the deep-seas.
The tentative age-climate curve based upon apollen analyses of the 200-meter core samples at
5 meters intervals was carried out by the present
writer (Fuji, 1983, 1984; Fuji & Horle, 1977),
and also more detailed curve checked by pollen
analyses of both the 200-meter and 1,400-metercores samples, and based upon a corrector empirically modified age-scale of the 200-meter core
(Yamamoto & Higashihara, 1978) has beenstated as shown in Text-fig. 13.
In the following part of this section, thewriter will compare tentatively the climaticchange from Lake Biwa with the oxygen isotope
stratigraphy in the Caribbean Sea and Equatorial Pacific, environmental changes in Central
Europe reconstructed by loess, palaeosol andgastropod faunal records, and Mediterranean andJapanese sedimentary cycles and relative sea levelchanges etc. However, both the palaeoclimaticchanges in Lake Biwa and the global climaticchanges have not been elucidated either. Namely,
the global climatic changes should be based
reasonably upon the various records from thedeep-seas and terrestrial sequences, but the majorchanges have not been stated yet. And also, thepresent writer finished only a pollen analysisof the 200-meter core samples at 5 meters intervals, and of the 1,400-meter core samples at 6
meters intervals obtaind from Lake Biwa. He,
therefore, is working the detailed pollen analysis
at 25 cm intervals of both the cores. After his
detailed analysis, the palaeoclimatic changes inand around the lake will be elucidated. And, if
the global climatic changes will be elucidated,
the writer will try the detailed correlation
between the climatic record from the lake andthe major climatic changes throughout the world.
Therefore, the correlation stated in the following

509

elucidation means of couse tentatively.
Concerning the oxygen isotopic records fromthe Carlbbean Sea, Emiliani (1955) and Emiliani& Shackleton (1974) divided the isotopic curves
into sixteen stages. The present writer (Fuji,
1983) compared his palaeoclimatic curve with
Emiliani's isotope temperature record depictedon the the modified time scale, and found arather significant correlation between them.
Especially, during the age about 270,000 to
100,000 years B.P., the c-5 and c-6 cold climatic
periods, and the t-5 temperate climatic period
described in this paper may be correlated with
Emiliani's stage nos. 6, 8, and 7, respectively.
The fact suggests that the palaeoclimatic changearound Lake Biwa was primarily controlled by the
global glacial-interglacial cycles, but in a few
exceptional periods of short duration the climate
was brought under the influence of a strong local
character and/or a problem in regard to absolute
age inferred.
The stratigraphic subdivision of the oxygenisotope record was subsequently extended analysis of sediments in Core V28-238 obtained froma depth of 3,120 m at 01 0 01' N, 160 0 29' E on
the Solomon Plateau in Western Pacific by
Shackleton and Opdyke (1973). Within thiscore, twenty-three isotopic stages younger than
Jaramillo Geomagnetic Event were recognized,
and the record was interpreted as reflecting moreor less continuous sediments since about 870,000
years B.P. And, the isotopic stages were also
recognized in the upper part of Core V28-239.
On comparison between the present writer'spalaeotemperature curve and the record from
V28-238 Core, a noticeable similarity between
major trends of the two curves seems to be
more than similarity between Emiliani's curve(Emiliani & Shackleton, 1974) and the palaeoclimate curve from Lake Biwa (Fuji, 1983; Fuji
& Horle, 1977). That is, at that time described inthis article, the Fuji's c-5 and c-6 cold periods,
and t-5 mild climatic period may be corresponded
to the Shackleton's stage nos. 6, 8, and 7, respectively. And also, in addition, the c-7 cold period
may be correlated with the cold stage between
stages nos. 8 and 9.

C]1

~

o

·
Y rs..
T tme

IPa I aeo-I

B.P.X 10'magne.

L oess cycles
Central EuropeI)

0 xygen Isotope.
2)0 xygen Isotope.
3) - Palaeoclimate.
4)

Caribbean

Pacificw?

wZ

10-1 B L c - - i . s = t 3 = w 3

15-1

•

BI~
20-1

_

w4

b1

I

Iw6

25--l

_

lelO

18~

-5

~

~ ~
el4
Ell

'"~ ~

~

u--l

50--l

I

cl8 cl9

G ,steppe ,glacial

WI

grass

)

1-8

-----F:1-------:. I

. ,"

'

,-' T-d

VI

XI

a

)X1b

·

1-35

~

-----'-_?_

T-e

H<

'

\.. ~?

t-9

~u

.

13
wl3
W14,)

forestinterglaci. ZZ·

Z61::

I ?
-1.0

18

I

C-IZ\
t-l0

I

I

0%0

-Z.O

N.C-T. Temp.

-40

m
+40

~50
m
-40

"':I
~

::::

1-30
)

EI ~--. - .. VZ

0

0

X

T-e

-" 1

1-7

~
::!.

IX

25

----------____ :,E~

IZ

20

'

-'

e-IO

wl2

cl7

I-

II

t

VID

,,"'T-b,-----. EZ .

wlO

)

--)

1-6

10

vn
H5

<'~::"T-a

-----

8

Ho

VI

)

--,)

I

L..,wll

cl6

S

~

--'DI

c-

V

H

--.,

Dz

c-9

=
Ii
i
~
Ol

YZ

?':1'.________
~

I-F§

d)

~

I

_--

e-7

III

"')

e

I >

jll'

./

CI

-6

I

' ) II

..J 0

Y3

---

w8

35--l

Tc-I

)

CZ
I-S

_D\ el2";::=ltS
w9-;"terraces
NewGuinea---.. M'--::_,

? (-

8

30--l

)

'--BI
e-4

Marine 7) T'lme Y rs.
B.P.X 10'6)~p--__________ -)

1-4

7
w7

~ZI_3
B3

:-- __ !l!,

6

c8

CI

c-Z

c

wS

5 ea Ieve I
Kanto, Japan

Maliorca

I-I w- I
I-Z

c-I

II
IZ

5e d'Iment. eye' I es 5)

Lake Biwa

822. Palaeovegetation and palaeoclimate, Lake Biwa

511

On the sedimentary cycles and relative sea
level changes, since changes of eustatic sea levels
are partly related to climatic changes, there
should be some broad correlation between dated
sea level changes in regions not affected byglacio-isostasy and the glacial and interglacial
cycles recorded in the deep-sea cores. There
should be some correspondence between ages oflowered sea level and cold condition, althoughclimatic change and sea level change may beslightly out of phase since time is required forthe melting of the ice sheets following climatic
improvement. Of cource, a following few problems to be solved are in combining the sea levelchange with the climatic change: (a) Eustatic sea
level changes from various regions throughout
the world are affected by both the voluminalchange of ocean water and tectonic movement;and (b) the correlation between the quantity ofice sheets and the global climate has not been
well established.
A work on the Pleistocene sedimentary
records of Mallorca in Western Mediterranean
is a typical example for comparison between
deep-sea and terrestrial records. Parts of the
Mallorca littoral with arid climate are characterized by well-developed calcareous dunes, andwidespread (Butzer, 1962). According to Butzer(1963), these aeolian sediments accumulated
through deflation of freshly-exposed marine
deposits during glacio-marine regressions, andeach regression can be correlated with eachglacial period in areas affected by glacier. Interglacial high sea levels are recorded by marine
terraces and beach deposits. The shoreline stratigraphy in Mallorca is divided into six terrestrial
and six marine hemicycles, which have been
dated partly on the basis of biostratigraphy andradiometric dating of marine shells (Butzer,
1975). These sequence can be correlated with theloess sequence in Central Europe, and with theoxygen isotope records from the deep-sea floors.
As shown in Text-fig. 14, the palaeoclimaticcurve from Lake Biwa (Fuji, 1983; Fuji & Horie,1977) shows a significant correlation with thesedimentary cycles from Mallorca (Butzer, 1975)
from the points of view of chronology and pat-

512

Norio FUJI

tern of change of curve. Namely, the following
facts can be pointed out:(a) The postglacial relative high sea level (Z1,Z2, and Z3) coincides with the warm W-1
and mild climate t-1 periods.
(b) The relative low stands of sea level (B3,
B2, and B1) during the latest Pleistocene
may be corresponded to the cold C-1,
C-2, and C-3 periods, respectively.
(c) And, especially concerning the present
paper, the high sea levels inferred from themarine terraces Y1 and Y3 from Mediterranean are correlated with the cooler ormild climatic age between c-5 and c-4
periods from Lake Biwa, and with t-4
period, respectively.
(d) The lower sea level C1 and C2 (Mediterranean) may be correlated with the slightly
cold or cold c-5 period (Lake Biwa).
(e) The high sea levels Xl and X2 (Mediterranean) coincide with the short temperate
age just after the cold c-6 period, andwith temperate t-5 period (Lake Biwa).
(f) The low sea levels D1 and D2 (Mediterranean) are corresponded to the cold age
between c-7 and c-6 periods (Lake Biwa),
and to the cold c-6 period, respectively.
The fact as mentioned above, suggests that thehigh sea level corresponds to the temperate
period of Lake Biwa, and that the low stand ofsea level to the cold one. The facts as recognized
between the curves from Lake Biwa and Mallorca
can be pointed out by comparing the palaeoclimate curve from Lake Biwa with the sea level
curve of Southern Kanto in Japan (Machida,1975), and with the sea level curve from New
Guinea (Chappell, 1974) as follows:(a) The high sea level P from Japan and I from
New Guinea during the latest Pleistocene
to Holocene is correlated with the warm
w-1 and t-1 periods from Lake Biwa.
(b) The relative low stand of sea level Tc-1
(Japan) and II (New Guinea) coincides
with the cold c-1 period (Lake Biwa).
(c) The high sea level suggested by the marine
Misake M and Obaradai 0 (Japan) and V(New Guinea) may be corresponded to thet-3 period (Lake Biwa).
(d) And, especially concerning the present
paper, the high sea level S (Japan) and VII(New Guinea) in the Shimosueyoshi transgression correlated with the Riss/Wiirminterglacial coincides with the temperate
age between the c-5 and c-3 periods (Lake
Biwa).
(e) The high sea levels inferred from themarine T-a (Japan) and VIII (New Guinea),T-b and IX, T-c and X, T-d and Xlb, andT-e and XII may be corresponded to thetemperate t-5 (Lake Biwa), slightly coolage between the c-7 and c-6, the temperate t-6, temperate t-7 periods, and to thetemperate t-8 period, respectively.
With regard to aeolian deposits, within
lengthy aeolian depositional sequences from
central Europe, one of standards for correlation in the Quaternary was provided. Manysoil layers are intercalated in the loess sequences, and the complete succession appears tocontain a record of glacial and interglaccial conditions (Kukla, 1970, 1975). In the succession,
the loess layers are interpreted as representing
glacial condition, and on the other hand, theinterbedded palaeosoils are considered to be
indicative of interglacial condition. These
records indicate that within the last 1.6
million years, seventeen major glacial andinterglacial conditions have affected central Europe (Text-fig. 14).
As shown in Text-fig. 14, in comparison withthe palaeoclimatic curve from Lake Biwa and thecentral European loess record (Kukla, 1970,1975) for the past 500,000 years, the following
facts can be pointed out:(a) The warm w1 age in the glacial cycle A
from Europe coincides with the w-1 andt-1 periods from Lake Biwa.
(b) The c1, tl, c2, t2, and c3 ages in the
middle and late glacial cycle B (Europe)
are correlated with the c-1, t-2, c-2, t-3
and c-3 periods from the lake, respectively.
(c) The warm w2 and w3 ages (Europe) may
be corresponded to the t-4 period (Lake822. Palaeovegetation and palaeoclimate, Lalle Biwa
Biwa) from the view point of a pattern ofclimatic change.
(d) The cold c5 and c6 ages (Europe) may
be corresponded to the c-4 period (Lake
Biwa) stated in this article.
(e) The cold c7 and c8 ages in the early partof glacial cycle C (Europe) may coinci dewith the c-5 period (Lake Biwa), and thewarm w5 age with the temperate age between the c-5 and c-4 periods, respectively.
(f) The warm w6 and w7 ages in the earlypart of glacial cycle C (Europe) may becorresponded to the temperate t-5
period (Lake Biwa).
As described above, the writer can find a remarkably noticeable similarity between major
trends of the records from Lake Biwa, other terrestrial sediments and the deep-sea~. However, amore reliable conclusion will be arrived at whenmore absolute dates and palynological results will
be obtained for the 200-meter and 1,400-metercores from Lake Biwa.
Conclusions(1) Pollen diagrams were obtained from 200
samples during the 55 meters to about 110
meters-horizons of the 200-meter core
which was drilled at the bottom 65 m
below the present water surface of the lake
in 1971.
According to the calculated age on the basisof the determination of 14C, fission-track
and palaeomagnetic stratigraphy, the age
during the 55 meters- and 110 meters-horizons is from about 100,000 yr.B.P. to about
250,000 yr.B.P.
(2) Judging from the pollen diagrams, thesamples during the 55 meters- to 110 metershorizons are divided into four pollen zones:
Zones Y-1, Y-2, Y-3 and Y-4, and the Y-2
zone is further divided into six pollen subzones.
(3) The palaeoclimate inferred on the basis ofpollen assemblage of every sample is asfollows:

513

periods palaeoclimate inferred
Y-1 Period: Northern part of the Cool Temperate zone; corresponding to
the c-6 period of the present
writer's previous paper (Fuji,
1983), the 8 stage from the
Shackleton and Opdyke's oxygenisotope curve, and to the D2 ofthe relative change curve of sea
level from Mediterranean.
Y-2-a Subperiod: Southern part of the Cool
Temperate - Temperate zones.
Y-2 b Subperiod: Middle part of the Cool
Temperate zone.
Y-2-c Subperiod: Southern part of the Cool
Temperate - Temperate zones.
Y-2-d Subperiod: Southern part of the Temperate - Warm Temperate zones.
Y-2-a to Y-2-d Subperiods correspond to the t-5 period from
Lake Biwa, the 7 stage of theisotope curve, and to the Xl to
X2 from Mediterranean.
Y-2-e Subperiod: Middle part of the Cool
Temperate zone; corresponding
to the c-5 from Lake Biwa, the
6 stage of the isotope curve, andto the C2 from Mediterranean.
Y-3 Period: Northern part of the Cool Temperate zone; corresponding to
the c-4 period from Lake Biwa,
and to the low sea level age between the Y2 and Y3 marine
terraces in Mediterranean.
Y-4 Period: Middle part of the Cool Temperate zone - northern part of theTemperate zone.
(4) Concerning the tentative correlation between the palaeoclimate from Lake Biwa,
palaeotemperature from the deep-seas suchas Caribbean Sea and Equatorial Pacific, environmental changes in Central Europe, andsea level changes in Mediterranean andJapan, the present writer can find a noticeable similarity between the records fromabove-mentioned regions as shown in Textfig. 14.

NorioFUJI

514

References
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Nishimura S. and Yokoyama, T. (1975): Fissiontrack ages of volcanic ashes of core samplesof Lake Biwa and the Kobiwako group (2).
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Shackleton, N. J. and .Opdyke, N. D. (1973):
Oxygen isotope and palaeomagnetic stratigraphy of Equatorial Pacific core V28-238:
oxygen isotope temperature and volumes on
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stratigraphy of Pacific core V28-239, late
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--(1977): Oxygen isotope and paleomagnetic
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Toyoda, Y., Horie, S. and Saijo, Y. (1968):
Studies on the sedimentation in Lake Biwa
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Yamamoto, A. (1976): Paleoprecipitationalchange estimated from the grain size variations in the 200m-long core from Lake

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Biwa: Paleolimnology of Lake Biwa and theJapanese Pleistocene, 4: 179-203.
- - , Kanari, S. and Fukuo, Y. (1973): Grain size
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Limnol., 34: 63-74.
-and Higashihara, M. (1978): Granulometricdata for the 200 meter core samples from
Lake Biwa (IV). Bull. Osaka Electro-Communication Univ., 14: 117-130.

illI*. Garno 7tll1:. Gifu ittJjl.. Hokuriku region~t~.tm:1J. Honshu *1+1. Iga Basin
Ise Bay ffr~~. Kaburaki ql ~11£. Kinki region J1I~.tm:1J. Kyoto ffiff~. Kyushu
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 144, pp. 516-527, pIs. 96-98, December 30,1986

823. LOWER PERMIAN CONODONTS FROM THE KAWANORI FORMATION
OF THE SOUTHWESTERN KANTO MOUNTAINS, TOKYOHISAHARU !GO
Department of Astronomy and Earth Sciences, Tokyo Gakugei University, Koganei, Tokyo 184
and
KEN-ICHIRO HIS ADA
Department of Geology, Osaka Kyoiku University, Tennoji, Osaka 543

Abstract. Lower Permian conodont elements are abundantly recovered fromthe limestone blocks embedded in the Kawanori Formation cropped out in thesouthwestern part of the Kanto Mountains, Tokyo. These conodonts are Hindeodus
minutus (Ellison) Pa element, Diplognathodus sp. A, Idiognathodus sp., Merrillina
sp. A, Neogondolella bisselli (Clark and Behenken), N. gujioensis Igo, N. idahoensis(Youngquist, Hawley and Miller), N. sp. A, Neostreptognathodus exsculptus Igo, N.
? sp., Streptognathodus elongatus Gunnell, Sweetognathus whitei (Rhodes) and
Xaniognathus abstractus (Clark and Ethington).
Most of these conodont elements are indicators of the Neogondolella bisselliSweetognathus whitei Assemblage zone. However, Streptognathodus elongatus andIdiognathodus sp. seem to be reworked elements from the older formation. Some
problems of the Lower Permian conodont zones in Japan and China were briefly
discussed herein.

Introduction and acknowledgement

shale and sandstone containing exotic blocks ofchert and limestone. Hisada concluded that thegroup is an olistostrome and subdivided into the
Nenoyama, Take kawadake , Urayama, Kawanori
and Nippara Formations based on the lithology
of contained exotic blocks.
Gray encrinal limestone block of the Kawanori Formation is exposed in the upstream of
the Ogawadani Valley, a tributary of the Nippara
River. Hisada (op. cit.) marked as F8 in his
geologic map and reported the occurrence ofLower Permian fusulinaceans and smaller foraminifers. The limestone block intercalating tuffaceous shale is about 4 m in thickness andmore than 10 m in length so far as exposed.
The following conodont elements were abun-

Recently, one of the authors, Hisada (1984)
clarified stratigraphy and geologic structureof the Paleozoic and Mesozoic strata in the
Ashigakubo-Kamozawa area, southwestern partof the Kanto Mountains. He subdivided thesestrata into the following three groups, such as
the Permo-Carboniferous Hashidate and Jurassic
Urayama Groups in the Chichibu Belt, and theJurasso-Cretaceous Ogouchi Group in the northern Shimanto Belt.
The Urayama Group is composed chiefly ofReceived November 9, 1985; revised manuscript
received September 10, 1986; read February 3,
1985 at The University of Tokyo.
516

823. Lower Permian Conodonts from Kanto Mountains

517

Text-fig. 1. Map showing locality of condont.
dantly recovered from this limestone, such as
Hindeodus minutus (Ellison) Pa element, Diplognathodus sp. A, MelTillina sp. A, Neogondolellabisselli (Clark and Behenken), N. gujioensis Igo,
N. idahoensis (Youngquist, Hawley and Miller),N. sp. A, Neostreptognathodus exsculptus Igo,
N. ? sp., Streptognathodus elongatus Gunnell,
Sweetognathus whitei (Rhodes), Xaniognathus
abstructus (Clark and Ethington), Idiognathodussp., and the following rami form elements, suchas Enantiognathus sp., Lonchodina spp., Diplododella sp., Hindeodella sp. and Hindeodus
minutus Sc element. These conodonts are particularly abundant in the individual numbers but
mostly fragmental. They seem to be transported
by current and accumulated together with thefragments of crinoid columns. Most of theseelements seem to be derived from the assemblage belonging to the Neogondolella bisselliSweetognathus whitei Assemblage zone (Igo,
1981). However, Streptog~thodus elongatus
and Idiognathodus sp. are older representatives
than the mentioned zone. These conodonts were
thought to be reworked from the older forma-tion.
The authors would like to express their appreciation to Professor Hisayoshi Igo of the University of Tsukuba for his cooperation in the field,
valuable suggestion and critical reading of
early version of this paper. We acknowledge thecomments and suggestions for this study byProfessors Tatsuaki Kimura of Tokyo Gakugei
University and Nobuo Yamagiwa of Osaka
Kyoiku University.
Appraisal of fauna
As already mentiond, conodont elements
recovered from limestone exposed at the
Ogawadani Valley (Text-fig. I) are abundant andcounted about 500 per 1 Kg, but they are mostly
fragmental and more or less abraded. Basedupon the occurrence and lithology of limestone,
these elements were transported by current anddeposited under high energy shallow water.
Associated fusulinaceans are mostly Triticites
and Pseudofusulina which also show some attrition. Most of the present conodont elements

518

Hisaharu IGO and Kenichiro HISADA

were brought from one assemblage, however, theobtained composition may represent a part ofthe orginal assemblage.
Hindeodus minutus (Ellison) is now treated asa long-ranging multielement species. Its Pa element is rather commonly obtained in our collection and similar to the forms described as
Anchignathodus minutus by many authors. This
type of discrete element is very common in theLower Permian limestone in Japan (Igo, op. cit.).
Diplognathodus sp. A is not so common in thepresent collection, but it is identical with Diplognathodus sp. A described from the Permian limestone exposed in "Haper Ranch beds" ofKamloops, southern British Columbia, Canada(Orchard, 1984b) and D. oertlii Kozur describedby Igo (1981) from the upper part of the Lower
Permian in the various localities in Central Japan.
Neogondolella bisselli is abundant in thepresent assemblage. This element is characteristic one in the Lower Permian of North America,
Europe and Asia, and is a leading species in the
middle part of Lower Permian in Japan. Recently, studies of the Permian conodonts in China
have been much progressed (Wang and Wang,
1981a, 1983 and others). This element is characteristic in the lower part of the Chihsia Stage
along with Sweetognathus whitei. The Chihsia
Stage is a classical standard lithogenetic unit of
the Permian in South China and was traditionallycorrelated with the Artinskian of USSR. Recently, however, the lower boundary of the Chihsia
has been placed within the Sakmarian and thebase of the Permian is settled at the base of the
Chihsia in China (Yang et al., 1984 and others).
Wang and Wang (1981a) proposed the assemblage
zones in the Chinese Lower Permian, such as the
Neogondolella bisselli-Sweetognathus
whiteiAssemblage zone and Neogondolella idahoensis
Assemblage zone in ascending order. The sameau thors (Wang and Wang, 1983) also establishedthe following three ass em blage zones in ascending order, such as the Neogondolella bisselliSweetognathus
whitei, Neostreptognathodus
pequopensis-N. sulcoplicatus, and Neogondolella
idahoensis-N. serrata Assemblage zones. The firstmentiond zone is established in the upper half of
the Zone of Pseudoschwagerina in the JapanesePermian. The Streptognathodus elongatus fauna
is found in the lower division of the Pseudo-

schwagerina zone which is characterized by theoccurrence of the primitive species of Pseudoschwagerina. This level is correlated with theAsselian of the Russian standard (Igo, 1981). As
already pointed out by many Permian conodont
specialists, the lowest Permian conodonts aremostly survival elements from the Upper Carboniferous and the occurrence of typical Permianelements are started from above the Asselian.
However, it does not indicate that the evidence issuitable to settle the base of the Permian as theinternational consensus.
Neogondolella gujioensis is common in thisassemblage. This species was originally describedfrom the Akuda Formation exposed in the Gujio
Hachiman district, Central Japan, and it resembles Neogondolella bisselli and N. idahoensis.
This species is characteristic in the uppermost
part of the Zone of Pseudoschwagerina associated with Neogondolella bisselli and Sweetognathus whitei in Japan.
Neogondolella idahoensis is also abundant in
the present collection. This species is classicalspecies of the Permian conodonts, and known
from the Shimadani Formation of the Gujio
Hachiman district, lower part of the Nabeyama
Formation of the Kuzuu district and other localities in Japan. Neostreptognathodus exsculptus is
not so common in our collection but seems to
be one of the important composition in thisfauna. This species was described from the uppermost part of the Zone of Pseudoschwagerina in
the lower subgroup of the Yoro Group andothers in Central Japan.
Streptognathodus elongatus is not so common
in the present fauna. This species was originallydescribed from the Upper Pennsylvanian of
U.S.A. Subsequently, this species has been
known in the Lower Permian of the various
localities in Asia and Europe. In Japan, it isreported from the lowermost Permian Shiraiwa
Limestone exposed at west of Orne City, Tokyo
Prefecture (Igo, op. cit.). Recently, Wang and Li(1984) described this species from the Taiyuan
Formation of North China in association with
Pseudoschwagerina.
The Chinese geologists
placed the Taiyuan in the uppermost division of
Carboniferous.
Sweetognathus whitei is also common in thepresent fauna. This species was originally described from the topmost of the Tensleep Sand-

823. Lower Permian Conodonts from Kanto Mountains
stone, Wyoming and its geologic age was thoughtas the Wolfcampian (Rhodes, 1963). S. whitei isyielded from the Pseudoschwagerina zone associated with Neogondolella bisselli in Central
Japan.
Xaniognathus abstructus is common representatives in the present fauna. This element iscommonly yielded from the upper part of theLower Permian in Japan. The original specimens
were known from the Leonardian Bone Spring
Formation, Texas and also from the Wordian inwestern United States (Clark and Ethington,1962).
As mentioned above, most of the present
conodont elements are known from the Neogondolella bisselli-Swee tognathus whitei Assemblage zone of Japan which was established in theupper part of the Sakamotozawan and correlated
to the lowest part of the Chihsia Stage in China,
and Sakmarian and Wolfcampian of the Russianand American standard divisions, respectively.
The present fauna contains slightly older elements
as the reworked specimens, such as Streptognathodus elongatus and Idiognathodus sp.
Although the present conodont elements wererecovered from the limestone block which isembedded in the Jurassic strata as an olistolith,
but they are additional data to the Lower Permian conodont fauna in Japan.

Systematic description
Genus Hindeodus Rexroad and Furnish, 1964

Hindeodus minutus (Ellison, 1941)
Pa element
PI. 98, Fig_ 14.

Spathodus minutus Ellison, 1941, p. 120, pI. 20,
figs. 50-52.
Spathognathodus minutus (Ellison); Youngquist
and Downs, 1949, p. 169, 170, pI. 30, fig. 4;
Sturgeon and Youngquist, 1949, p. 385, pI.
74, figs. 9-11; Rexroad and Burton, 1961, p.
1156, 1157, pI. 141, figs. 10, 11; Hass, 1962,
p. 209, pI. 34, fig. 36; Murray and Chronic,
1965, p. 606, pI. 72, figs. 29, 30; Igo and
Koike, 1965, p. 88, 89, pI. 9, figs. 16-18;
Dunn, 1965, p. 1149, pI. 140, figs. 15, 21, 24;
Koike, 1967, p. 311, pI. 3, figs. 39-42;

519

Webster, 1969, p. 44, pI. 7, fig. 4;Dunn, 1970,
p. 339, pI. 61, figs. 27,30; Merrill, 1973, p.
305, 306, pI. 1, figs. 1-14, pI. 2, figs. 1-28;
Lane and Strak, 1974, p. 101, fig. 44: 7,
12; Spassov, 1976, p. 135, pI. 2, fig. 1.
Spathognathodus cfr. S. minutus (Ellison);
Huckriede, 1958, p. 162, pI. 10, fig. 8;
Rhodes, 1963, p. 408, pI. 47, fig. 3.
Anchignathodus minutus (Ellison); von Bitter,
1972, p. 65, 66, pI. 6, figs. 2a-2i; Sweet in
Ziegler, 1973, p. 15-17, Anchignathodus-pl.
1, figs. 2a, 2b; Behnken, 1975, p. 297, pI. 1,
figs. 16-18; Tynan, 1980, p. 1300, pI. 2, figs.
8,9; Wang and Wang, 1981a, p. 229, pI. 2, figs.
6, 7; Wang and Wang, 1981b, p. 115, 116, pI.
1, fig. 5; Wang and Dai, 1981, pI. 1, figs. 3, 4;Igo, 1981, p. 26, pI. 10, figs. 5, 8, 11; Zhao,
Han and Wang, 1984, p. 114, pI. 28, figs. 21,
22, 24, 26, 28; Tian, 1983, p. 338, pI. 77, fig.
4; Wang and Li, 1984, pI. 2, figs. 9, 16, 17.
Anchignathodus minutus (Ellison); Kozur, 1975,
p. 5-7, pI. 1, figs. 1-16; Kozur et al., 1975,
p. 3, pI. 1, figs. 2, 3, 5, 7-11, pI. 2, figs. 1, 3,
5, 7-9, pI. 7, figs. 2, 10; Kozur, 1977, p.
1118,1119, pI. 1, figs. 1-16.
Anchignathodus minutus permicus Igo, 1981, p.
26,27, pI. 10, figs. 1-4.
Anchignathodus typicalis Sweet, 1970a, p. 7, 8,
pI. 1, figs. 13, 22; Sweet, 1970b, p. 222, 223,
pI. 1, figs. 13, 20; Sweet et al., 1971, pI. 1,
fig. 34; Sweet in Ziegler, 1973, p. 19, 20,
Anchignathodus-pl. 1, fig. 3; Sweet in Teichert
et al., 1973, p. 427, pI. 11, figs. 8, 9; Behnken,
1975, p. 297, 298, pI. 2, fig. 12; Sweet, 1976,
p. 52,53, pI. 16, figs. 6-8, 9?; Wang, 1978, p.
215, pI. 1, figs. 26-28; Tian, 1983, pI. 77, fig.
9, pI. 79, fig. 4; Zhao et al., 1984, p. 114,115,
pI. 28, figs. 18-20, 23, 27, 29,31.
Ozarlwdina minutus (Ellison), p element(Spathognathodonta), Baesemann, 1973, p.
704, 705, pI. 2, figs. 14 15, 19, 20; Toomey
et al., 1974, pI. 3, fig. 8.
Hindeodus typicalis (Sweet) Pa element, Sweet
in Ziegler, 1977, p. 223, 224, Hindeodus-pl. 2,
figs. 1-6.
Hindeodus ex. gr. Hindeodus minutus (Ellison),Bender, 1980, p. 10, pI. 4, fig. 22.
Hindeodus minutus(Ellison) Pa element, Matsuda,
1981, p. 80-84, pI. 1, figs. 1-13; Orchard,
1984b, p. 212, pI. 23.1, figs. 6, 26.
Spathognathodus echigoensis Igo and Koike,
1964, p. 187, 188, pI. 28, figs. 24, 25?

520

Hisaharu JGO and Kenichiro HISADA

Spathognathodus cristula Youngquist and Miller;
Stibane, 1967, p 335, pI. 35, figs. 21-25.
Spathognathodus rexroadi Webster, 1969, p. 45,
pI. 7, figs. 2,3 (non fig. 1).
Remarks:-Anchignathodus typicalis was distinguished from Spathognathodus minutus by its
ratio of length to width (Sweet, 1970b). Matsuda(1981) stressed that L/W ratio of Spathognathodus minutus from the Pennsylvanian of U.S.A.
(Merrill, 1973) and Lower Triassic of Kashmir isoverlapping with the range of Anchignathodus
typicalis (L/W = 2.0-2.5) by Sweet (1970b).
Spathognathodus minutus is a well-known
long range element and shows broad variation inexternal shape. The shape of posterior end and
L/W ratio do not have any sufficient criteria
to distinguish Anchignathodus typicalis from
Spathognathodus minutus as Pa element of
the multielement species, Hindeodus minutus(Ellison).
Reg. no. TGU. 1744.
Genus Diplognathodus Kozur and Merrill, 1975

Diplognathodus sp. A.
PI. 98, Figs. 10-13.

Diplognathodus sp. A, Orchard, 1984b, p. 210,
pI. 23.1, figs. 7,9.
Diplognathodus oertlii Kozur; Igo, 1981, p. 32,
pI. 8, figs. 9-12, 14-16 (non fig. 13).
Remarks:-Recently, Orchard (1984b) has
described Diplognathodus sp. A from "Harper
Ranch beds" in Kamploops of southern British
Columbia, Canada and discussed the blade andcarina profiles of related species. Igo (1981)
described the similar forms from the Gujio
Hachiman and Samegai districts, Central Japan as
Diplognathodus oertlif Kozur, but they differ
from Kozur's type specimen in the shape of
platform and outline of blade.
Reg. nos. TGU. 1740-1743.
Genus Merrillina Kozur, 1975

Merrillina sp. A
Pi. 98, Fig. 22.

Description:-Unit symmetrical and its posterior part expanded laterally and heart-shape inoral view. Unit with three long, sharply pointed
and discrete posterior denticles. Cusp large,
sharply pointed and steeply inclined posteriorly. Anterior margin of unit anteriorly convex.
Anterior denticles inclined posteriorly. Posterior
margin of unit concave anteriorly. In aboral view,
basal cavity broad, laterally expanded in posterior part.
Remarks:-This unnamed species resembles
Merrillina divergens (Bender and Stoppel), but
the former is distinguished from the latter bylarger cusp. Misikella posthernsteini Kozur andMock superficially resembles this species. However, this Upper Triassic species has almost thesame denticles and cusp in size.
Reg. no. TGU. 1752.
Genus Neogondolella Bender and Stoppel, 1965

Neogondolella bisselli (Clark and Behnken, 1971)
PI. 97, Figs. 10-15.

Gondolella bisselli Clark and Behnken, 1971, p.
429, pI. 1, figs. 12-14.
Neogondolella bisselli (Clark and Behnken);
Behnken, 1975, p. 306, pI. 1, figs. 27, 31;
Clark et al., 1979, pI. 1, fig. 21; Igo, 1981,
p. 37, pI. 1, figs. 1-10 14 17-19' Orchard
1984a, pI. 22.1, fig. 14 (no~ figs. 16: 17).
'
Neogondolella cfr. bisselli (Clark and Behnken);
Malkowski and Szaniawski, 1976, p. 82, pI. 1,
figs. la, b, c.
Description:-Unit characteristically slendertear-drop in shape. Platform tapers gradually to
sharp pointed anterior end and rounded posterior
end. Cusp sharply pointed in early growth stage
and rounded in mature stage. Carina of mature
specimens have 11 to 15 discrete denticles withrounded apex. In gerontic stage, posterior part ofplatform slightly flat, fused denticles developedin posterior half of carina, a few denticles proj-

823. Lower Permian Conodonts from Kanto Mountains
ect in anterior half and cusp inconspicuously
lowered. Oral surface of platform ornamented by
micro-pits.
Remarks:-This species resembles Neogondolella gujioensis Igo, particularly in early growthstage, but the former is distinguished from thelatter in having long and slender platform.
Neogondolella bisselli (Clark and Behnken) has
more numerous nodes on carina than the latter.
Neogondolella idahoensis (Youngquist, Hawley
and Miller) also resembles this species, but theformer has square posterior end, large cusp andsquare loop.
Recently, this species has been described fromthe lower part of Chihsia Formation of Ziyun,
Guizhow in China where it is associated with
Neostreptognathodus pequopensis and Anchignathodus minutus (Wang and Wang, 1981a).
Based upon the occurrence of this species, they
established the Neogondolella bisselli-Sweetognathus whitei Assemblage zone in the Chihsia
Stage.
Reg. nos. TGU. 1721-1726.

Neogondolella gujioensis Igo, 1981
PI. 97, Figs. 1-9.

Neogondolella gujioensis Igo, 1981, p. 37, 38,
pI. 3, figs. 1-19, pI. 4, figs. 1-6.
Description:-Unit tear-drop in shape, slightly arched, and roundly pointed posterior end.
Margin of platform tapers to pointed anterior
end and both sides of unit subparallel in central
part. Platform extends in full length of unit.
In oral view, both sides of platform inflated
upward, and shallow lateral furrows present
along carina. Oral surface of platform with smallpits.
In aboral view, lower surface bears broad keel.
Basal groove connected to small basal cavity surrounded by small loop.
Remarks:-This species resembles Neogondolella bitteri (Kozur), but the former is distinguished from the latter in having short platform
which is 2/3 posterior of full length of unit.

521

This species resembles closely Neogondolellaliangshanensis Wang characterized by deep lateralfurrows and sharply pointed higher anterior
denticles. N. liangshanensis was describedfrom the Upper Permian Wuchiaping Formation,
Liangshan area of southern Shanxi in China(Wang, 1978). N. gujioensis Igo is probably
ancestral form of N. liangshanensis.
Reg. nos. TGU. 1712-1720.

Neogondolella idahoensis (Youngquist, Hawley
and Miller, 1951)
PI. 96, Figs. 1-11.

Gondolella idahoensis Youngquist, Hawley andMiller, 1951, p. 361, pI. 54, figs. 1-3,14,15;
Clark and Ethington, 1962, p. 108, pI. 2, figs.
15, 16; Clark and Mosher, 1966, p. 388, pI.
47, figs. 9-12; Clark and Behnken, 1971, p.
431, pI. 1, fig. 9; Movshovich et al., 1979, pI.
2, figs. 3a, b.
Neogondolella idahoensis (Youngquist, Hawley
and Miller); Behnken, 1975, p. 306, 307, pI.
1, figs. 28-30; Wang, 1978, p. 220, 221, pI.
2, figs. 23-26; Szaniawski and Malkowski,
1979, p. 246, 247, pI. 4, figs. 1-8, pI. 5, figs.
5a, b; Clark et al., 1979, pI. 1, fig. 10; Wang
and Wang, 1981a, p. 230, pI. 2, figs. 15, 23;
Igo, 1981, p. 38, pI. 1, figs. 11-13, 15, 16;
Wardlaw and Collinson, 1984, p. 269, pI. 1,
figs. 10, 11.
Gondolella bisselli Clark and Behnken, Kozur,
1978, pI. 2, figs. 16a-c, pI. 3, figs. 6a-c, 7,
lOa, b; Movshovich et al., 1979, pI. 1, fig. 11,
pI. 2, figs. 1a-c.
Neogondolella bisselli (Clark and Behnken);Orchard, 1984a, pI. 22.1, figs. 16, 17, (nonfig. 14); Orchard, 1984b, pI. 23.1, fig. 10,(non figs. 11, 17).
Description:-Unit slightly arched and asymmetrical. Platform margin tapers to pointed
anterior end. Both sides of unit in posterior half
subparallel. Anterior half of unit lanceolate.
Posterior margin of unit flat. Platform extends
in full length of unit. Carina consists of 15 or 16
rounded denticles in mature specimens. Anterior
half of carina consists of slightly pointed discreteHisaharu !GO and Kenichiro HISADA

522

denticles and posterior half with low and rounded
nodes. Large and stout casp located posterior
end of platform. Flattened posterior margin
rimed by laterally raised carina in some specimens.
Remarks:-This species resembles Neogondolella bisselli (Clark and Behnken), but theformer is distinguished from the latter in having flattened posterior margin. The range of
Neogondolella idahoensis is the Leonardian ofTexas, Utah and Wyoming in U.S.A. and the
Nabeyaman in Japan. This species also reported
from the Upper Chihsia Formation in China.
Behnken (1975) pointed out that Neogondolellabisselli is ancestral form of Neogondolella
idahoensis. Igo (1981) followed his opinion andproposed Neogondolella intermedia which represents an evolutionary link between N. bisselli
and N. idahoensis.
Reg. nos. TGU. 1701-1711.

like denticles. Discrete and pointed denticles
developed in anterior 1/3 of carina. Cusp small
and not located at posterior end of platform.
Carina not reach at posterior end of platform.
Posterior margin of platform rised upward.
Remarks:-This species resembles Neogondolella gujioensis Igo, but the former is distinguished from the latter by the position of cusp.
Cusp of Neogondolella gujioensis is located on
the posterior end of platform, but it is located atslightly anterior side from the posterior endof platform in this species.
Reg. no. TGU. 1700.

Neogondolella sp. A

Description:-In oral view, unit elongate withsharply pointed posterior end. Platform provided
by 11 transverse ridges. Anterior part of platform
pinched laterally. Well developed deep narrow
median trough extends throughout platform andtransverse section of oral surface deeply Vshaped. Anterior blade joints median part ofanterior platform.
In lateral view, unit slightly arched and platform high. In aboral view, expanded basal cavity
large, elongate, deepest at center of platform andextends to anterior free blade as narrow groove.
Remarks:-This species resembles Neostreptognathodus newelli which was described fromPI. 96, Fig. 12.
Neogondolella sp. A; Igo, 1981, p. 39, pI. 4,
fig. 12.
Gondolella cfr. orientalis Barskov and Koloeva;
Suyari, Kuwano and Ishida, 1983, pI. 1, figs.
6a, b.
Description:-In oral view, unit laterally subsymmetrical, lanceolate with sharply pointed
anterior end and roundly pointed posterior terminal. Surface of platform pitted and granular.
Shallow lateral furrows present on both sides ofcarina. Posterior 2/3 of carina composed of node-

Genus Neostreptognathodus Clark, 1972
Neostreptognathodus exsculptus Igo, 1981
PI. 98, Figs. 1, 2.
Neostreptognathodus exsculptus Igo, 1981, p.
40, 41, pI. 5, figs. 2, 3a, b, 4a, b.

Explanation of Plate 96
Fig. 1-11. Neogondolella idahoensis (Youngquist, Hawley and Miller) . . . . . . . . . . . . Page 521
Figs. 1-3: Oral views of broken specimens of anterior part. TGU. 1701, 1702, 1703. Fig.
4: Oral view of twisted specimen. TGU. 1704. Figs. 5, 7, 8, 9, 11: Oral views. TGU. 1705,
1707, 1708, 1709, 1711. Fig. 10: Slightly lateral view. TGU. 1710. Fig. 6: Aboral view ofbroken specimen of anterior part. TGU. 1706.
Fig. 12. Neogondolella sp. A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 522
Oral view of mature specimen. TGU. 1700.
Marker in 4, 7, 8, 9, 11=1000 microns: marker in 1, 2,3, 5,6, 10, 12=100 microns.

IGO and HISADA: Lower Permian Conodonts from Kanto Mountains

Plate 96

823. Lower Permian Conodonts from Kanto Mountains
several localities of Wyoming and the lower partof Meade Peak, Montpelier Canyon section,
Idaho, U.S.A. by Wardlow and Collinson (1984).
Neostreptognathodus newelli Wardlow and Collinson is distinguished from N. exsculptus in having more numerous pairs of transverse ridges. On
anterior part of platform of N. newelli, transverse
ridges fused together to make node-like ridges inparallel with median groove.
Reg. nos. TGU. 1731, 1732.

523

Gnathodus simplex (Gunnell), Movshovich et al.,
1979, pI. 1, figs. 9, 10.

Remarks:-The material examined has characteristically deep V-shaped median through and
13 to 15 transverse ridges.
Reg. nos. TGU. 1737-1739.
Genus Sweetognathus Clark, 1972

Sweetognathus whitei (Rhodes, 1963)
PI. 97, Figs. 16-19.

Neostreptognathodus ? sp.
PI. 98, Figs. 3-6.

Remarks:-The material examined is poorly
preserved and anterior blade is lacking. The unit
has broadly expanded basal cavity and V-shaped
median trough. One rounded node-like ornamentation is developed on the inner side of platform.
Many indistinct transverse ridges exist on bothsides of median trough.
Reg. nos. TGU. 1733-1736.

Genus Streptognathodus Stuffer andPlummer, 1932

Streptognathodus elongatus Gunnell, 1933
PI. 98, Figs. 7-9.
Streptognathodus elongatus Gunnell, 1933, p.
283, pI. 33, fig. 30; Ellison, 1941, p. 130, pI.
22, fig. 9; Rhodes, 1963, p. 405, pI. 47, figs. 5,6, 16-24, 27-34; Sweet in Ziegler, 1975, p.
369, Streptognathodus-pI. 1, fig. 6; Igo, 1981,
p. 43, 44, pI. 7, figs. 10, 12-15; Ishiga, 1982,
pI. 2, figs. 11-13 (non fig. 14); An et al.,1983, p. 179, 180, pI. 32, figs. 3-5; Wan etal.• 1983, pI. 1, figs. 1-4; Wang and Li,
1984, p. 200, 201, pI. 1, figs. 1-7; Zhao etal., 1984, p. 133, pI. 25, figs. 11-20.
Gnathodus elongatus (Gunnell), Kozur, 1978, pI.
1, fig. 4.
Streptognathodus cfr. S. elongatus Gunnell,
Rhodes, 1952, p. 894, pI. 127, figs. 3, 4,8.
Streptognathodus simplex Gunnell, 1933, p. 285,
pI. 33, fig. 40.

Spathognathodus whitei Rhodes, 1963, p. 404,
pI. 47, figs. 4, 9, 25, 26; Clark and Behnken,
1971, pI. 1, figs. 2-6; Merrill, 1973, p. 310,
pI. 3, figs. 8, 9.
Sweetognathus whitei (Rhodes); Clark, 1972, p.
155; Clark, 1974, pI. 2, figs. 12-18; Behnken,
1975, p. 312, pI. 1, fig. 26; Sweet in Ziegler,
1977, p. 547, Neostreptognathodus-pl. 1,
figs. 7a--c; Kozur, 1978, pI. 3, figs. 8, 9a,b; Movshovich et al., 1979, pI. 1, fig. 12;
Clark et al., 1979, pI. 1, fig. 4, Wang andWang, 1981a, p. 231, pI. 1, fig. 20, pI. 2, figs.
8, 10,11,22; Igo, 1981, p. 44, pI. 6, figs. 1722, pI. 7, figs. 1-9; Tian, 1983, p. 334, pI. 77,
figs. 3a, b, 11a, b; Zhao et al., 1984, p. 136,
pI. 26, figs. 1-16; Orchard, 1984b, p. 213,
pI. 23.1, figs. 3-5, 8.

Remarks:-Sweetognathus behnkeni Kozur
with well developed lateral nodes and S.
whitei are yielded together in Japan. Bothspecies transition ally change within the samelevel, therefore, it is very difficult to dis tin:guish as an independent species. Igo (1981)
referred Sweetognathus behnkeni as a gerontic form of S. whitei.
Reg. nos. TGU. 1727-1730.
Genus Xaniognathus Sweet, 1970

Xaniognathus abstractus (Clark andEthington, 1962)
PI. 98, Figs. 15-21.
Subbryantodus abstractus Clark and Ethington,
1962, p. 112, pI. 1, figs. 16, 20 (non fig. 21).

524

Hisaharu lGO and Kenichiro HISADA

Xaniognathus abstructus (Clark and Ethington),
1973, Sweet in Ziegler, p. 473, XaniognathuspI. 1, fig. 1; Igo, 1981, p. 44, 45, pI. 11, figs.
9, 10; Wardlow and Collinson, 1984, p. 271,
pI. 2, figs. 1-12.
Ozarkodina cfr. O. abstructus (Clark and Ethington), Zhao et al., 1984, pI. 27, figs. 7, 8.
Description:-In lateral view, anterior processslightly high and with long numerous den tides.
Anterior den tides 8 to 11 and posterior dentides
3 or 4. Anterior 1 or 2 dentides small. Dentides
long, sharply pointed and discrete one-third of
their length. Cusp large, long and sharply pointed.
Posterior process slightly curved towards inside.
In aboral view, basal cavity small and extended
into both processes as a narrow groove.
Reg. nos. TGU. 1745-1751.
Repository:-All the specimens treated inthis paper are preserved in the Department of
Astronomy and Earth Sciences, Tokyo Gakugei
University.

References cited
An, T., Zhang, F., Xiang, W., Zhang, Y., Xu, W.,
Zhang, H., Jiang, D., Yang, C., Lin, L., Cui,
Z. and Yang, X. (1983): The conodonts ofNorth China and the adjacent regions. Geol.
Publishing House, Beijin China, 223 p., 33
pis. (in Chinese with English abstract).
Baesemann, J. F. (1973): Missourian (Upper
Pennsylvanian) conodonts of northeastern
Kansas. Jour. Paleont., vol. 47, p. 689-710,
pis. 1-3.
Behnken, F. H. (1975): Leonardian and Guadalupian (Permian) conodont biostratigraphy inWestern and Southwestern United States.
Ibid., vol. 49, p. 284-315, pis. 1, 2.
Bender, K. P. (1980): Lower and Middle Pennsylvanian conodonts from the Canadian Arctic
Archipelago. Geol. Suru. Canada, Paper 7915, p. 1-20, pis. 1-4.
Clark, D. L. (1972): Early Permian crisis and its
bearing on Permo-Triassic conodont taxonomy. Geologica et Palaeontologica, vol. 1, p.
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--(1974): Factors of Early Permian conodont
paleoecology in Nevada. Jour. Paleont., vol.
8, p. 710-720, pis. 1, 2.
--and Behnken, F. (1971): Conodonts andbiostratigraphy of the Permian. Geol. Soc.
America, Mem., 127, p. 415-439, pis. 1, 2.
- - , Carr, T. R., Behnken, F. H., Wardlow, B.
R. and Collinson, J. W. (1979): Permian
conodont biostratigraphy in the Great Basin.
In Sandberg, C. A. and Clark, D. L. eds.,
Brigham Young Univ. Geol. Studies, vol. 26,
pt. 3, p. 143-147, pI. 1.
--and Ethington, R. L. (1962): Survey of Permian conodonts in Western North America.
Ibid., vol. 9, pt. 2, p. 102-114, pis. 1, 2.
--and Mosher, L. C. (1966): Stratigraphic, geographic, and evolutionary development of theconodont genus Gondolella. Jour. Paleont.,
vol. 40, p. 376-394, pis. 45-47.
Dunn, D. L. (1965): Late Mississippian conodontsfrom the Bird Spring Formation in Nevada.
Ibid., vol. 39, p. 1145-1150, pI. 140.
- - (1970): Middle Carboniferous conodontsfrom Western United States and phylogenyof the platform group. Ibid., vol. 44, p.
312-342, pis. 61-64.
Ellison, S. P. (1941"): Revision of the Pennsylvanian conodonts. Ibid., vol. 15, p. 107143, pis. 20-23.

Explanation of Plate 97
Figs. 1-9. Neogondolella gujioensis Igo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 520
Fig. 1: Slightly lateral view of juvenile specimen. TGU. 1712.
Figs. 2-6: Oral views of adolescent specimens. TGU. 1713, 1714, 1715, 1716, 1717.
Figs. 7-9: Oral views of mature specimens. TGU. 1718, 1719, 1720.
Figs. 10-15. Neogondolella bisselli (Clark and Behnken) . . . . . . . . . . . . . . . . . . . . . Page 521
Figs. 10, 11: Slightly lateral views of mature specimens. TGU. 1721, 1722.
Fig. 12: Slightly lateral view of adolescent specimen. TGU. 1723.
Figs. 13-15: Oral views of broken specimens of anterior part. TGU. 1724,1725,1726.
Figs.16-19 Sweetognathus whitei (Rhodes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 523
Oral views of boken specimen of blade. TGU. 1727, 1728, 1729, 1730.
Marker in 10,11,12=1000 microns: marker in 1-9,13-19=100 microns.

Jeo and HJSADA: Lower Permian Conodonts from Kanto Mountains

Plate 97

823. Lower Permian Conodonts from Kanto Mountains
Gunnell, F. H. (1933): Conodonts and fish
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Ibid., vol. 7, p. 261-297, pIs. 31-33.
Hass, E. H. (1962): Conodonts. In Moore, R. C.
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Permian radiolarians from the Tamba belt,
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--(1977): Revision der Conodontengattung
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--(1978): Beitrll.ge zur Stratigraphie des Perms
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525

- - , Mostler, H. and Rahimi-Yazd, A. (1975)
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526

Hisaharu !GO and Kenichiro HISADA

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- - , Mosher, L. C., Clark, D. L., Collinson, J. W.
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p. 1282-1309, pIs. 1, 2.

Explanation of Plate 98
Figs. 1, 2. Neostreptognathodus exsculptus Igo . . . . . . . . . . . . . . . . . . . . . . . . . .. Page 522
Oral views of broken specimens of blade. TGU. 1731, 1732.
Figs. 3-6. Neostreptognathodus? sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 523
Oral views of broken specimens of anterior part. TGU. 1733, 1734, 1735, 1736.
Figs. 7-9. Streptognathodus elongatus Gunnell . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 523
Oral views of broken specimens of blade. TGU. 1737, 1738, 1739.
Figs. 10-13. Diplognathodus sp. A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 520
Figs. 10-12: Oral views. TGU. 1740, 1741, 1742.
Fig. 13: Lateral view. TGU. 1743.
Fig. 14. Hindeodus minutus (Ellison) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 519
Pa element, Lateral view. TGU. 1744.
Figs. 15-21. Xaniognathus abstructus (Clark and Ethington) . . . . . . . . . . . . . . . . . . Page 523
Lateral views. TGU. 1745, 1746, 1747, 1748, 1749, 1750, 1751.
Fig. 22. Merrillina sp. A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 520
Lateral view. TGU. 1752.
Markers of all figures = 100 microns.

lao and HISADA: Lower Permian Conodonts from Kanto Mountains

Plate 98

823.

Lower Permian Conodonts from Kanto Mountains

von Bitter, P. H. (1972): Environmental control
of conodont distribution in the Shawnee
Group (Upper Pennsylvanian) of eastern
Kansas. Univ. Kansas Paleont. Contrib., Art.
59, 105 p. 16 pis.
Wan, S., Ding, H. and Zhao, S. (1983): Middle
and Late Carboniferous conodont biostratigraphy of North China. Jour. China Coal
Soc. No.2, p. 62-72, pis. 1, 2 (in Chinese).
Wang, C. and Wang, Z. (1981a): Permian conodont biostratigraphy of China. Geol. Soc.
America Special Paper 187, p. 227-236, pis.
1,2.
--and --(1981b): Permian conodonts fromthe Longtan and Changhsing Formations of
Changxing, Zhejiang and their stratigraphicaland paleoecological significance. Selected
papers on the 1st. convention of Micro·
paleontological Soc. of China. Geol. Publishing House, Beijin China, p. 114-120. pis. 1,
2 (in Chinese).
- - and - - (1983): Review of conodont biostatigraphy in China. Fossils and Strata, No.
15, p. 29-33.
Wang, Z. (1978): Permian-Lower Triassic conodonts of the Liangshan area, southern
Shanxi. Acta Palaeont. Sinica, vol. 17, p.
213-227, pis. 1, 2 (in Chinese with English
abstract).
--and Dai, J. (1981): Triassic conodonts fromthe Jiangyou-Beichuan area, Sichuan Province. Acta Palaeont. Sinica, vol. 20, p.
138-150, pis. 1-3 (in Chinese with English
Kawanori JlI.ilt
Ogouchi 'NiiJJ1'J,

527

abstract).
--and Li, R. (1984): Discovery of Late Carboniferous conodonts from Shanxi Province
and its significance. Acta Palaeont. Sinica,
vol. 23, p. 196-203, pis. 1, 2 (in Chinesewith English abstract).
Wardlow, B. R. and Collinson, J. W. (1984):
Conodont paleoecology of the Permian
Phosphoria Formation and related rocks of
Wyoming and adjacent area. In Clark, D. L.
ed., Conodont biofacies and provincialism.
Geol. Soc. America, Special Paper 196, p.
263-272, pis. 1-5.
Webster, G. D. (1969): Chester through Derry
conodonts and stratigraphy of northern Clark
and southern Lincoln Counties, Nevada.
Univ. California, Pub. Geol. Sciences, vol.
79, 121 p., 8 pis.
Yang, Z., Cheng, Y. and Wang, H. (1986): Thegeology of China, 303 p., 19 pis. Clarendon
Press, Oxford.
Youngquist, W. and Downs, R. H. (1949): Additional conodonts from the Pennsylvanian of
Iowa. Jour. Paleont., vol. 23, p. 161-171,
pis. 30,31.
- - , Hawley, R. W. and Miller, A. K. (1951):
Phosphoria conodonts from southeastern
Idaho, Ibid., vol. 25, p. 356-364, pI. 54.
Zhao, Z., Han, J. and Wang, Z. (1984): The Carboniferous strata and its fauna from southwestern margin of Tarim basin in Xinjiang.
Geol. Publishing House, Beijin China, 187 p.,
35 pis. (in Chinese with English abstract).

Ashigakubo pi T )..f!j!;, Kamozawa ~r~iR, Hashidate t!iirr, Urayama Iililll,
Nenoyama.::y· / Ill, Takekawadake :iitJlIff.-, Nippara S me Ogawadani 'Nil fl.

~JIHW1!H\ij[1lj;lilJII~,U11J'Gi£:l±llJ.:T;'~)':::'\~*:::l /r":/ r : r)~nIHlJ:ltfJ.ml'-4{;f)0)<=t-' ·1,l£l\'?

Lt, )..f1] I:' J: -:> -C, ':::1~1:-::::£i:*0)t!iij'[mi C :; :>- '7 *O)iiIiIW~1lf, :; "- '7 - 1':1 l]i*O)'H'JP'JII'Q
tfl:.lRn\.~nt-.:or p-A-e,rfliLlllffi!llfa, TI:'fi,Ii'J;'L~-,.::y-nlJ~, iR)IIITr~, il1IllI~,7"- -\" - r, lilJ('t:-t.( c·0)7f.*~'i'P,-ts:;l- Y A I, A

JII*)!,

SljRI~I:.;tllln\.~nt-.:o]ltJ?:ffllJllji4!)O)S)jj()IISZ:VltO)'NII:frl:IiiH:.7};(/l-r7.»II*M<=t-'0):&@(:E:11~~0):::l /~~<XO) J: ~ t.d!lIJ~~5JIJ ~f:/ r'i'n t-.:o Hindeodus minutus (Ellison) Pa element, Diplognathodus

sp. A, Merrillina sp. A, Neogondolella bisselli (Clark and Behnken), N. gujioensis
Igo, N. idahoensis (Youngquist, Hawley and Miller), N. sp. A, Neostreptognathodus
exsculptus Igo, N.? sp., Streptognathodus elongatus Gunnell, Sweetognathus whitei(Rhodes), Xaniognathus abstractus (Clark and Ethington), Idiognathodus sp., Enantiognathus sp., Lonchodina spp., Diplododella sp., Hindeodella sp., Hindeodus minutus(Ellison) Sc element. ;:. GO):::l / f:/ r 11 Neogondolella bisselli-Sweetognathus whitei
assemblage zone O)fJ'lfJili:fi.l!-C-;I; 7.> IJ;, Streptognathodus elongatus c Idiognathodus sp.
f1-tnJ: I)E~'llilf\:;'i'7j'; L-. ~i!1j:{I::& c~;tGn7.>omJllB)"~· )..E13fij!-UlInTrans. Proc. Palaeont. Soc. Japan, N.S., No. 144, pp. 528-540, pIs. 99-100, December 30,1986

824. EARLY JURASSIC PLANTS IN JAPAN. PART 7.
FOSSIL PLANTS FROM THE NISHINAKAYAMA FORMATION,TOYORA GROUP, YAMAGUCHI PREFECTURE,
SOUTHWEST JAPANTATSUAKI KIMURATokyo Gakugei University, Koganei, T9kyo 184
GENTARO NAITO
26·9, Higashi·Matsuzaki-cho, Hofu City, Yamaguchi Prefecture 747

and
T AMIKO OHANATokyo Gakugei University, Koganei, Tokyo 184

Abstract. This is the seventh of our serial papers on the Early Jurassic plants
in Japan. In this paper we describe the following fossil plants obtained from the
Nishinakayama Formation of marine origin: Phlebopteris sp., Otozamites micropinnatus, sp. nov., O. sp. A, O. sp. B, Zamites toyoraensis Oishi, Z. sp., Pseudoctenissp., Brachyphyllum ex gr. expansum (Sternberg) Seward, Cupressinoc/adus sp. A,
C. sp. B, Elatides sp. and Geinitzia sp.
As our specimens are mostly of poorly preserved impression, it is difficult togive clear specific names but they are all characteristic constitutional elements ofthis florula.
This florula is quite different in floristic composition from the coeval floras of
the Kuruma Group and Iwamuro Formation in Japan and of the Xiangxi Formationin West Hubei, China.

Introduction

Utano Formation. From the Higashinagano Formation only Nilssonia brevis Brongniart was
described by Oishi (1940). The Nishinakayama
Formation is divided by Hirano (1971, '73) into
Nf-, Np-, Nd- and Na-ammonite Zones in upward
sequence. According to Hirano (1971, '73),
the geological age of the Nishinakayama Formation ranges from Pliensbachian to early Toarcian
based on its rich ammonite-fauna.
The following fossil plants were described by

The Toyora Group of marine origin consistsof the Higashinagano, Nishinakayama and Utano
Formations in upward sequence distributed inwestern part of Yamaguchi Prefecture, Southwest Japan. Invertebrate fossils are very abundant but fossil plants are rather rare except the* Received December 9,1985: Read February 1,
1986 at Tohoku University.
528824. Early Jurassic Plants
Yabe (1922), Oishi (1935, '40) and Huzioka(1938) from the Nishinakayama Formation:
Phlebopteris takahasii Huzioka, 1938; Zamites
toyoraensis Oishi, 1935; Z. yabei Oishi, 1940(= Cf. Zamiophyllum buchianum by Yabe,
1922); Brachyphyllum expansum (Sternberg)
Seward (Oishi, 1940, but not illustrated).
We recognized the following plant-taxa among
the collection made by one of us (Naito), Y.
Kurihara, H. Ishida, T. Fukutomi and others:
Phlebopteris sp., Otozamites micropinnatus
Kimura, Naito et Ohana, sp. nov., O. sp. A,
O. sp. B, Zamites toyoraensis Oishi, Z. sp.,
Pseudoctenis sp., Brachyphyllum ex gr. expansum (Sternberg) Seward, Cupressinocladus
sp. A, C. sp. B, Elatides sp. and Geinitzia sp.
Although the number of plant-taxa is small,
it is worth mentioning that this florula is quite
different in floristic composition from thecoeval floras from the Kuruma Group andIwamuro Formation in Japan (Kimura and529

Tsujii, 1980a, b, '81, '82, '83, '84) and fromthe Xiangxi Formation (s. str.) in West Hubei,
China (Wu, S. Q. et al., 1980).
The floristic features of the Nishinakayama
Formation will be discussed in the last of ourserial papers (Early Jurassic plants in Japan.
Part 8) together with the concluding remarks ofthe Early Jurassic floras in Japan and its adjacentareas. Accordingly this paper deals only with thedescription of fossil plants newly obtained fromthe Nishinakayama Formation.
Figs. lA, B show the distribution of the
Nishinakayama Formation and the main localities of fossil plants described herein.
Systematic description
Filicales: Family Matoniaceae
Genus Phlebopteris Brongniart, 1836: 371

Phlebopteris sp.

131'02'

1A

./

18

-34'10'o

1km- -_ _- - I 'Figs. lA, B

Distribution of the Nishinakayama Formation and localities of fossil plants.

1A; an arrow indicates the location of the Nishinakayama Formation (after Hirano, 1971). 1B; solid

circles show the main localities of fossil plants.

530

Tatsuaki KIMURA, Gentaro NAITO and Tamiko OHANA

Text-figs. 1a-d
Material: N1984. Locality: Ishimachi (Loc.
no. 062). Horizon: Nd-Zone. Occurrence: Rare.
Description: Obtained is a single sterile pinna
fragment, 3 cm long and 1.7 cm wide at its basalpart, narrowing gradually towards its distal part.
The pinna axis is slender, 0.8 mm wide, giving
oppositely off closely set pinnules at nearlyright angle. The pinnules are elongate-triangularin form; the largest one preserved is 8 mm longand 3 mm wide near the base; adjacent pinnules
are connected by a web, 0.5 mm wide. Apicesof pinnules are obtusely pointed; margins are
entire, but most of them are markedly recurved.
Veins are partly visible; midnerve is prominent,
giving off lateral veins forking at once into forward and backward branches which join thoseof adjacent branches to form the 'arches'. The'arches' are usually wider than high, rather flatand never including any branch veins. The'arches' give off 2-3 outer veins running nearly
perpendicularly to the margins. The outer veinsare forking once near the 'arch' or near themargin, and sometimes anastomosing withadjacent branches to form elongate meshes.
In the web region, the 'arches' are along the
pinna axis, small, semi-circular and each giving
off simple or once forked outer veins directed
forwards in the acroscopic base of pinnuleand directed backwards in the basiscopic baseof pinnule.
Remarks: Judging from the characteristic
venation observed in our sterile pinna, it is certain that our pinna is referable to Phlebopteris.
But it is difficult to make its specific identity
because of the insufficiency of our material.
Huzioka (1938) described Phlebopteris takahasii as a new species from Ishimachi (the same
locality as ours) based on a single large-sizedsterile pinna fragment. Huzioka's pinnules arelong and narrow, linear, more than 3 cm long and
5 mm wide, and attached to the thick pinna axis,
4 mm wide at a wide angle. If the venation of
Huzioka's pinnules is properly drawn, Phlebopteris takahasii may be distinct from ours, becausein Huzioka's drawing, there are no 'arches' but

crowded lateral veins directly from the midnerve_
Our Phlebopteris sp. differs from otherPhlebopteris species hitherto described in its
rather flat 'arches' along the both sides of midnerve, instead of angular (as shown by Harris,
1961, figs. 33, 35) in others.
Bennettitales
Genus Otozamites Braun, 1842
Otozamites micropinnatus Kimura,
Naito et Ohana, sp. nov.
PI. 99, Figs. 1-3; PI. 100, Figs. 1-2;

Text-figs. 2a--c
Material: Holotype; N1983 (Narusebashi; Loc.
no. 064). Paratype; N3648 (ditto). Other specimens: 1983A, 3644, 3685, 3687 (ditto). Stratum
typicum: Nishinakayama Formation (Upper Npand Lower Nd-Zones). Locus typicus: Ishimachi,
Toyora-cho, Toyora-gun, Yamaguchi Prefecture(Loc. no. 064)_ Derivatio nominis: According tosmall-sized pinnae. Occurrence: Not rare.
Diagnosis: Typical leaf elongate-oblanceolate
in outline, more than 10.5 cm long, up to 2 cm
wide. (apex and petiole not known.) Rachis
comparatively thick, 2 mm wide measured on
impression, sending off more than 50 alternate
pairs of small-sized pinnae at a wide angle (7075 degrees). Pinnae attached to the upper surfaceof rachis by the middle of base or by the lower
half of semi-amplexicaul base and often imbricate. Pinnae on the apical half of a leaf elongated, 1 cm long and 2.5 mm wide near auricle;
those on the proximal half short-rectangularand those on the proximal part semi-circularor triangular, 2 mm long and 1.2 mm wide atbase, sometimes falcate; apex obtuse; pinna baseasymmetrical and auricula ted. Auricle free anddeveloped near acroscopic basal angle; auricleof the opposite pinna lying below partly covering
basiscopic basal margin of the pinna lying above;
then upper surface of rachis usually concealed.
Veins emerging from the point of attachmentof pinna and radiating into auricle and the restof pinna, 10-12 in number near the base; a few
forking dichotomously near the base_ (Cuticle

824. Early Jurassic Plants
not preserved and reproductive organs notknown.)
Description: PI. 99, Fig. 2 (Text-fig. 2a, c)shows the holotype of Otozamites micropinnatus
represented by its under surface. But fortunately
its rachis was partly stripped off, then we could
recognize both the whole shape and the modeof attachment of pinnae.
The general leaf-form is shown in PI. 99,
Fig. 1 (paratype), although its apex and petiole
are missing. This is under surface view of leaf,
then characteristic pinna bases including developed auricles are all concealed by the crushed
rachis. Such leaf as mentioned above can be
confused with the small-sized leaves of Ptilophy!lum.
PI. 100, Fig. 1 shows also under surface viewof an apical half of the largest leaf in our hand;
its preserved part with 40 alternate pairs of
pinnae is 11 cm long, but its apex is missing.
PI. 99, Fig. 3 and PI. 100, Fig. 2 show two
proximal parts of leaves in under surface view,the former is smaller and the latter is larger insize both with marked petiole.
Discussion and comparison: Otozamites micropinnatus is characterized by its elongate-oblanceolate leaf bearing small-sized and closely
set pinnae with well developed auricles.
Most Otozamites species hitherto known
have far larger leaves than O. micropinnatus. So
far as we know, Otozamites species bearing
small-sized pinnae are; O. bengalensis (Oldham
et Morris), O. bunburyanus Zigno var. indica
Seward et Sahni, O. chuensis Zhou, O. kilpperiana Barnard et Miller, O. margaritaceus
Zhou, O. mattiellianus Zigno, O. mixomorphus
Ye, O. parvus Zhou, O. teneZlus Zhou.
Among them Otozamites be nga Ie ns is, O.
bunburyanus var. indica, O. kilpperiana, O.
parvus and O. tene!lus are clearly distinguishedfrom O. micropinnatus by their leaves all withsemi-circular or nearly circular pinnae and also
O. margaritaceus is distinguished by its leaf
with triangular pinnae.
In Otozamites chuensis originally describedby Zhou (1984) from the early Lower Jurassic
Guanyintan Formation, Southwest Hunan, China,531

its pinnae are similar in form to those on the
distal half of leaf of O. micropinnatus, but it
is difficult to make full comparison of Zhou's
species with our species because Zhou's original
specimen is only represented by 7 pairs ofelongated pinnae.
In Otozamites mattiellianus Zigno (Zigno,
1873-1885; Wesley, 1974) described by Zhou
(1984) from the Guanyintan Formation, its
pinnae are similar in form to those of our elongated ones, but are larger in size and with submucronate (or submucronulate) apex.
Ye (in Wu, S. Q. et aI., 1980) proposed
Otozamites mixomorphus as a new species basedon Sze's O. bengalensis (Oldham et Morris)(Sze, 1949, p. 17, pI. 4, fig. 3b; pI. 11, fig. 4)
from the late Lower Jurassic Xiangxi Formation(s. str.), Western Hubei, China. Otozamites
mixomorphus is most close in size and pinna
form to our O. micropinnatus, but it is difficult
to make full comparison of Ye's species withours because Ye's species is represented only
by broken parts of leaves.
Under the circumstances, we here propose
Otozamites micropinnatus as a new speci!=!s.

Otozamites sp. A
Text-fig. 3

Material: NI-1004, 1008. Locality: Nishinakayama Quarry (Loc. no. 066). Horizon: Nd-Zone.
Occurrence: Rare.
Description: Leaf is pinnate. Pinnae are set
closely, alternate, imbricated, elliptic in form,
1.2 cm long and 0.8 cm wide and attached tothe upper sides of rachis at a wide angle andconcealed the rachis completely; basiscopic baseof pinna is entirely concealed by the acroscopic
base of opposite pinna. Acroscopic basal angle
is not auriculated; upper surface of pinnae is usually convex. Veins are divergent, dichotomously
forking once or twice as shown in Text-fig. 3;
26 per cm in density.
Remarks: In general, form and size of
Otozamites pinnae are different according to
the position of a leaf. Thus it is difficult to makethe specific identity for such leaf-fragments

Tatsuaki KIMURA , G entaro NAITO and T

532

amlO
'k 0 HANA

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F

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824. Early Jurassic Plants
as ours represented only by 7 alternate pairs of
pinnae.
Our pinnae somewhat resemble proximal
pinnae of Otozamites hsiangchiensis Sze (e. g.
Wu, S. Q. et al., 1980, pI. 17, fig. 6) from thelate Lower Jurassic Xiangxi Formation (s. str.),Western Hubei, China.
Leaves with similar pinnae are represented
by Otozamites walkamotaensis Bose et ZabaBano and O. kachchhensis Bose et Banerji
known from the Middle-Upper Jurassic plantbeds in Kachchh (Bose and Banerji, 1984).
But these pinnae are characterized by more orless developed acroscopic angles.
Most of all, Otozamites sp. A resembles
macroscopically O. bunburyanus Zigno (Wesley,
1974, pI. 2, fig. 4) from the Lower Jurassic of
Italy.

Otozamites sp. B
PI. 100 Fig. 3; Text-fig. 4a-b

Material: N3684, IC-20ll (2014; counterpart). Localities: Narusebashi (Np-Zone) andIshimachi (Nd-Zone). Occurrence: Rare.
Description: Obtained are two broken leavesof which pinnae are closely set, imbricated andare attached alternately to the upper surfaceof thick rachis (4 mm wide) at a wide angle,
concealing the rachis completely. Pinnae are
elongate-triangular in form, sometimes falcatewith obtusely pointed apex and with asymmetrical and contracted base possibly with thesmall area of attachment near the basiscopic
margin; acroscopic angle is not so prominent.

533

Basiscopic base and margin are usually concealedby the acroscopic base of an opposite pinna
and the acroscopic margin of pinna of subordinate position respectively. Veins are typical
Otozamites-type. Although their origin is concealed, judging from our observation as shownin Text-fig. 4b, veins are small in number (5-6)
at the base, then repeatedly forking dichotomously near the base; more than 20 in number
at the middle of pinna. Cuticle is not preserved.
Remarks: Although very base of pinnae isusually concealed and acroscopic angle of pinna
is not well developed, it is highly probable that
our leaves fall under the category of Otozamites,
because our pinna base is asymmetrical, contracted, attached to the upper surface of rachis
and the pinnae with typical Otozamites-typevenation.
Our leaf is characterized by its closely imbricated pinnae, but at present we reserve to makeits specific identity and regard it as Otozamites
sp. B. until more satisfactory leaves are available.
The leaves of Otozamites sp. B resemble inform some leaves of O. hsiangchiensis Sze (Wu,S. Q. et al., 1980) known from the late Lower
Jurassic Xiangxi Formation (s. str.), Western
Hubei, China, but is distinguished by its closely
imbricated pinnae and undeveloped acroscopic
pinna angle.
Genus Zamites Brongniart, 1828

Zamites toyoraensis Oishi
PI. 100, Fig. 4; Text-fig. 5a-b

Zamites

toyoraensis

Oishi:

Oishi,

1935,

Text-figs. 1-7(length of solid bar indicates 1 cm, and length of broken bar indicates 0.5 cm).
1 . Phlebopteris sp.: la, b; pinna fragments (N1984). lc; enlarged partly from lao 1 d; ditto, showing
the venation of pinnule. 2. Otozamites micropinllatus Kimura, Naito et Ohana, sp. nov.: 2a; middle
part of a leaf (holotype; N1983) (PI. 99, Fig. 2). 2b; basal part of a leaf (N3685) (PI. 99, Fig. 3). 2c;enlarged from 2a, showing the venation of pinna. 3. Otozamites sp. A (NI-1004), showing the
pinna-form and venation. 4. Otozamites sp. B: 4a; middle part of a leaf (IC-20ll). 4b; enlarged
from 4a, showing the venation of pinna. 5. Zamites toyoraellsis Oishi: 5a; proximal part of a leaf
(N1990) (PI. 100, Fig. 4). 5b; enlarged from 5a, showing the basal part of pinnae and venation of
pinna. 6. Zamites sp.: 6a; a part of leaf (IC-lOOl). 6b; enlarged partly from 6a, showing the venation of pinna. 7. Pseudoctenis sp.: 7a; distal part of a leaf (N1988) (PI. 100, Fig. 5). 7b;enlarged
from 7a, showing the venation.

534

Tatsuaki KIMURA, Gentaro NAITO and Tamiko OHANA

p. 98, text-fig. 1 (Tarai); 1940, p.358 (remarks);
Takahashi, 1973a, pI. 1, fig. 1 (Ishimachi).
Material: N1990, 3684, 3718. Localities:
Narusebashi (Loc. no. 064; Np-Zone) and Sakuraguchidani (Loc. no. 067; Upper Np- and Lower
Nd-Zones). Occurrence: Not rare.
Description: Among three specimens in ourhand one (PI. 100, Fig. 4; Text-fig. 5a-b) isrepresented by the proximal portion of a leaf.
The pinnae are elongate-deltoid or short-rectangular in form below, 1.2 cm long and 0.4 cm
wide near the base, and more elongate-deltoid
above, 1.7 cm long and 0.5 cm wide near thebase, slightly falcate and all with obtusely
pointed apex; attached to the upper surface ofthick rachis (0.5 cm wide measured on impression) by a rounded symmetrical and non-auriculate base with 'callosity' at a wide angle. The
veins are slender and densely crowded; someforking dichotomously near the base, mostly
parallel to each other except at the base where
they are divergent, not converging at apex;
about 16 in number near the base (Text-fig.
5b). Cuticle is not preserved.
Remarks: Zamites toyoraensis was firstdescribed by Oishi (1935) based on a single
broken leaf representing the middle-distal part.
Takahasi's illustration (Takahasi, 1973a) showedalso the same part of a leaf. According to Oishi(1935), in the original leaf, pinna apices areusually acuminate instead of obtusely pointed
in ours and veins are 25-30 in number at thebasal broadest portion of each pinna insteadof 16 in ours. We now think, however, that ourleaf shown in PI. 100, Fig. 4 represents theproximal portion of a leaf belonging to Zamites
toyoraensis, because in other bennettitaleanleaves, such morphological differences as mentioned above are usually recognized according
to the position of pinnae in a leaf.

Zamites sp.
PI. 99, Fig. 4; Text-fig. 6a-b

Material: N3676, IC-100l, IC-1002 (?),
N3647 (?). Locality: Narusebashi (Loc. no. 064;
Np-Zone). Occurrence: Rare.

Description: Obtained are four ill-preserved
leaf-fragments. Thus whole leaf is unknown. In
Text-fig. 6a, leaf is more than 6.5 cm longand up to 4 cm wide with rather thick rachis(3 mm wide). Pinnae are set closely and attachedalternately to the upper sides of rachis at awide angle, but not concealed it completely.
Pinnae are linear, nearly parallel-sided for themost part, typically 2 cm long and 2.5 mm
wide; pinna base slightly contracted and nearly
symmetrical, but its very base and mode of
attachment to the rachis are obscure; pinna apex
is obtusely pointed. Veins are 9 in number atbase, acroscopic and basiscopic one or two veinsare slightly divergent and forking once near thebase; other veins are parallel, not converging
at apx. PI. 99, Fig. 4 may represent the apicalpart of a leaf; its pinnae are small-sized andattached to the rachis at an angle of 45 degrees.
Cuticle is not preserved.
Remarks: Although very base of pinnae isobscure, it is highly probable that our leaves
fall under the category of Zamites because our
pinna base is contracted, symmetrical andattached to the upper sides of rachis and theveins of acroscopic side curving basipetally andthe veins of basiscopic side acropetally.
Our leaf is characterized by its small-size andclosely set pinnae, but we here reserve its specificidentity and regard it as Zamites sp. until moresatisfactory leaves are available.
At first glance, the leaves of Zamites sp.
resemble in external form large-sized leaves of
Pterophyllum nathorsti Schenk (or Tyrmia
latior Ye) and those of Ptilophyllum contiguum
Sze both described by Wu, S. Q. et al. (1980)
from the late Lower Jurassic Xiangxi Formation,
Western Hubei, China. But Zamites sp. is distinguished from Pterophyllum nathorsti of which
pinnae are attached to the lateral sides of rachis
and from Ptilophyllum contiguum of which
pinnae are with asymmetrical base.
Oishi (1940, p. 358, pI. 37, fig. 6) described
Zamites yabei based on Yabe's Cf. Zamiophyllum
buchianum (Yabe, 1922) collected possibly
from Nd-Zone of the Nishinakayama Formation(Takahasi, 1973b). According to Oishi (1940),

824. Early Jurassic Plants
Zamites yabei is characterized by the large leaf
bearing long, narrow and parallel-sided pinnae
attached to the upper surface of the rachis andeach with 15 (?) parallel veins. There is a possibility that Zamites sp. A represents the apicalportion of a leaf of Zamites yabei.

Cycadales
Genus Pseudoctenis Seward, 1911: 19
Pseudoctenis sp.
Pl. 100, Fig. 5; Text-fig. 7a-b

Material: N1988. Locality: Narusebashi (Loc.
no. 064; Np-Zone).
Occurrence:' Rare.
Description: The leaf is imparipinnate withrather stout rachis, 4 mm wide. The whole leaf
is unknown. The pinnae are long and narrow,535

typically 6.5 cm long, nearly parallel-sided forthe most part, 4.5 mm wide, gradually narrowing
to the obtusely pointed apex, constricted near
the base then expanded and decurrent at thebase, and attached remotely to the lateral sidesof rachis at an angle of 40 degrees; angles reducing distally. The veins are typically 5 in number
at base; some of them are forking once or twice
dichotomously, then running parallel to themargins and ending at the distal margins of
pinna; 9 in number (10-11 per cm) at the
middle. Cuticle is not preserved.
Remarks: A single broken leaf was obtained(collected by T. Utamura in 1976). According
to its pinna-form the present leaf has led us toregard it as belonging to the cycadean genus
Pseudoctenis.
Pseudoctenis sp. closely resembles in external
form P. sp. described by Oishi (1940, p. 324,

1
Text-figs. 8-11 (length of solid bar indicates 1 cm).
8 . Brachyphyllum ex gr. expansum (Sternberg) Seward; a part of branching twig
(IC-3003) (PI. 99, Fig. 7). 9. Cupressinocladus sp. A; a pinnately branched shoot(IC-5010) (PI. 99, Fig. 6; PI. 100, Fig. 7). 10. Cupressinocladus sp. B; a branching
twig (IC-5003). 11. Elatides sp.; a branching twig (IC-5012) (PI. 99, Fig. 5). 12.
Geinitzia sp.; an ill-preserved branching twig (IC-2002).

536

Tatsuaki KIMURA, Gentaro NAITO and Tamiko OHANA

pI. 29, fig. 6) from Takaji (Middle Jurassic Utano
Formation). But Oishi's leaf is distinguishable
by its small-size (about one-half of ours) andmore crowded veins (13 in each pinna).
The leaves described by Oishi (1940, p. 322,
pI. 29, figs. 1-3) as Pseudoctenis lanei Thomasfrom the Upper Jurassic Oginohama Formation,
Northeast Japan are also distinguishable fromthat of ours by its pinnae each with acuminate
apex and with 13 veins.
Pseudoctenis nipponica originally describedby Kimura and Tsujii (1983) from the Negoya
Formation, Kuruma Group is distinguished byits pinnae each with acutely pointed apex andwith crowded veins (typically 13 in number).
A Wealden species, Pseudoctenis eathiensis(Richards) Seward (1885, 1917) is one of thecomparable species in size with the present
P. sp., but is distinguished by its pinnae which
are sometimes longer (the longest is 17 cm)
and are more contracted near the base.
Thus, Pseudoctenis sp. is considered to be
distinct, but we at present reserve to give it a
new specific name until the subsequent supplyof material.
Coniferales
Form-genus Brachyphyllum Lindley et Hutton
1836 ex Brongniart, 1828
Brachyphyllum ex gr. expansum(Stenberg) Seward
PI. 99, Fig. 7; PI. 100, Fig. 8; Text-fig. 8Comparable Japanese specimens:

Brachyphyllum expansum (Sternberg) Seward:
Oishi, 1940, p. 391, pI. 39, figs. 10-11; pI. 40,
fig. 9 (Takaji, Utano Formation).
Material: IC3003 (3004), N3636, 3638,
3718, 3719. Locality: Andadani (Loc. no. 031;
Np-Zone). Occurrence: Not rare.
Description: The branching-twig shown in
PI. 99, Fig. 7 (Text-fig. 8) is the largest one in
our collection. In this twig, branches are nearlystraight and main branch is more than 6 cm longand 2 mm wide, giving alternately off closely
set secondary l)ranches in one plane; the secondary branches are more than 2.8 cm long andgiving alternately off short third ones. Leaves
are arranged spirally and mostly appressedto the stem. Leaf is arising from a leaf-basecushion which may be elongated. Leaf with
leaf-base cushion varies from 0.5 cm long and
2 mm wide on the main branch to 2 mm longand 1.5 mm wide on the ultimate branch. Cuticle
is not preserved.
Remarks: In the case of such twigs without
cuticle and reproductive organs as ours, it isdifficult to make precise specific identity, but
our twigs resemble in external form those illustrated by Oishi (1940) as Brachyphyllum expansum from Takaji. Under the circumstances,
we regard our twigs provisionally as Brachyphyllum ex gr. expansum.
So far as we know, such branching twig as
ours has not been recorded from the Upper
Triassic plant-beds in Japan, Korea and fromthe late Lower Jurassic Kuruma Group and its
equivalents in Japan. However, Z. Y. Zhou (1984)
illustrated two small twigs as Brachyphyllum

Explanation of Plate 99(Natural size unless otherwise indicated)
Fig. 1. Otozamites micropinnatus Kimura, Naito et Ohana, sp. nov.: Paratype (N3648).
Fig. 2. Otozamites micropinnatus Kimura, Naito et Ohana, sp. nov.: Holotype (N1983) (Textfigs. 2a, c).
Fig. 3. Otozamites micropinnatus Kimura, Naito et Ohana, sp. nov.: N3685 (Text-fig. 2b).
Fig. 4. Zamites sp.: N3676.
Fig. 5. Elatides sp.: IC-5012 (Text-fig. 11).
Fig. 6. Cupressinocladus sp. A: IC-5012 (Text-fig. 9).
Fig. 7. Brachyphyllum ex gr. expansum (Sternberg) Seward: IC-3003 (Text-fig. 8).

KIMURA, NAITO and OHANA : Early Jurassic Plants

Plate 99

824. Early Jurassic Plants
sp. from the early Lower Jurassic Guanyintan
Formation, Southwest Hunan, China. So thiswould be the oldest record of such-like plants
in East Aia.
Takahasi (1973b) recorded the occurrenceof Brachyphyllum expansum from the uppermost zone (Sinemurian) of Higashinagano
Formation and the Np-Zone of Nishinakayama
Formation and of Brachyphyllum sp. fromthe Nd-Zone of Nishinakayama Formation.
But they were neither described nor illustrated.

537

other pair is free except the basal halves and freepart is at a right angle to the axis, instead of allappressed and not radiated in ours.
Similar branched leafy shoots to ours haveso far been described under various genericnames, but at present it is difficult to makespecific identity and to mention their botanicalaffinity for such sterile impressions as ours.
Cupressinocladus sp. B
Text-fig. 10

Form-genus Cupressinocladus Seward, 1919:307
Cupressinocladus sp. A
PI. 99, Fig. 6; PI. 100, Fig. 7; Text-fig. 9

Material: IC5010, 5011 (5012, counterpart).
Locality: Narusebashi (Loc. no. 064; Np-Zone).
Occurrence: Rare.
Description: PI. 99, Fig. 6 (PI. 100, Fig. 7
enlarged from PI. 99, Fig. 6; Text-fig. 9) showsa pinnately branched shoot with scaly leaves indecussate pairs, more than 12 cm long. The
second branches (in this figure) are alternate,
about 1.5 cm distant; the longest one is 3.5
cm long. The ultimate branches are nearly thesame wide as the secondaries, up to 1.3 cm longand opposite proximally but alternate distally.
The leaves are small-sized, rhomboidal in outline,
all appressed to the axis, weakly keeled dorsallywith acutely pointed apex. Cuticle is not preserved.
Remarks: Among two branched leafy shoots
now in our hand, the illustrated one is wellpreserved. Our branched leafy shoots can not bereferable to Form-genus Brachyphyllum because
Brachyphyllum is defined as its leaves are spirally
arranged (Harris, 1979). Our branched leafy
shoots agree well with the definition of Cupressinocladus given by Seward (1919), Chaloner
and Lorch (1960) and Harris (1969).
Cupressinocladus koyatoriensis originally described by Oishi (1940) from the Upper Jurassic
Oginohama Formation, Northeast Japan differsfrom our C. sp. A in that in Oishi's decussate
leaves, one pair is appressed to the axis but theMaterial: IC5003. Locality: Ishimachi (Loc.
no. 062; Nd-Zone).
Occurrence: Rare.
Description: Obtained is a single ill-preserved
branching twig of which main axis is more than
5.7 cm long sending alternately off shorter
secondaries at an angle of 40 degrees in one
plane; secondary branch is 1.2 cm long sending
off very short third branches. Leaves are rhomboidal in outline with acutely pointed apex anddecussate in arrangement; dorsal and ventral
pairs are usually appressed to the axis but lateralpairs are free, diverging from axis typically at
40 degrees but decurrent proximally. Cuticle
is not preserved.
Remarks: Cupressinocladus sp. B differs fromC. sp. A in its free lateral leaves. Our twig is too
incomplete to compare with other twigs withdecussate leaves hitherto known.
Form-genus Elatides Heer, 1876 em. Harris,
1976: 66
Elatides sp.
PI. 99, Fig. 5; PI. 100, Fig. 6; Text·fig. 11
Material: N1981, 1982, 3638, 3689, IC5012.
Localities: Narusebashi (Loc. no. 064; Np-Zone)
and Sakuraguchidani (Loc. no. 067; Upper Npand Lower Nd-Zones). Occurrence: Not rare.
Description: Obtained are five broken leafy
shoots. The leaves are persistent, spirally andclosely set, diverging radially and arising fromdecurrent cushion. Free part of leaves is falcate

538

Tatsuaki KIMURA, Gentaro NAITO and Tamiko OHANA

and rhomboidal in section each with obtusely
pointed apex and a single keel (vascular bundle?). Cuticle is not preserved.
Remarks: The above description agrees wellwith the emended diagnosis of Elatides given by
Harris (1979).
In Pl. 99, Fig. 5 (Text-fig. 11), only parallelor nearly parallel leaves to the bedding plane
were preserved completely but other leaves
were broken during the collection leaving theircrossing rhomboidal section.
The leafy shoots of our Elatides sp. resemble
in leaf-form those of other Elatides species
such as E. asiatica (Yokoyama) Krassilov, E.
curvifolia (Dunker) Nathorst and those regarded
as Araucarites hitherto described.
The original specimen of Elatides asiatica was
first described by Yokoyama (1906) from theLower Jurassic (?) of Liaoning, China as Palissyamanchurica. Since then similar leafy shoots
have been described by many authors from theLower Jurassic to Lower Cretaceous plant bedsin North and Northeast China and Central Asia
as Elatocladus manchuricus, E. subzamioides
and E. asiatica. Obviously there are some morphological differences among these leafy shoots
but it is difficult to separate them specifically
based only on their external form.
Then it is appropriate to regard Elatides
asiatica as a comprehensive 'species' ranging inage from Early Jurassic to Early Cretaceous.
Elatides sp. is somewhat different in leafform from E. asiatica in its more falcate leaves.
The leafy shoots of Elatides sp. is most closein leaf-form to those regarded as the othercomprehensive species E. curvifolia first de-scribed from the Wealden of Germany, especiallyin its stout and markedly falcate leaves.
The leafy shoots of Araucarites vassilevskiae
Turtanova-Ketova known from the Lower-Upper
Jurassic of Central Asia somewhat resemble inleaf-form those of Elatides sp., but the former
leaves are more delicate and more markedly
curved adaxially than the latter.
The occurrence of leafy shoots of Elatidestype is the first record in the Mesozoic of Japan.
Form-genus Geinitzia Endlicher, 1847: 280
Geinitzia sp.
Text-fig. 12

Material: IC2002. Locality: Ishimachi (Loc.
no. 062; Nd-Zone).
Occurrence: Rare.
Description: Obtained is a single small-sized
branching twig of which secondary branches
are long and narrow, 2.3 cm long and 1 mm
wide, sending off occasional third branches.
Leaves are spirally arranged; free part of leaf isspreading, 1-2 mm long, elongate-triangular inform, possibly equally thick in vertical andhorizontal directions and merging into basal
cushion on stem without contracting. Cuticle
is not preserved.
Remarks: The features of the present leafy
twig, though small fragment, agrees with theemended diagnosis of Geinitzia given by Harris(1979). But it is difficult to make further argument about this twig because of its ill-preservation.
Harris (1979) noticed that the elongated

Explanation of Plate 100(Natural size unless otherwise indicated)
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.

Otozamites micropinnatus Kimura, Naito et Ohana, sp. nov.: N3644.
Otozamites micropinnatus Kimura, Naito et Ohana, sp. nov.: N3687.
Otozamites sp. B: N3684.
Zamites toyoraensis Oishi: N1990 (Text-fig. 5a, b).
Pseudoctenis sp.: N1988 (Text-fig. 7a, b).
Elatides sp.: N3689.
Cupressinocladus sp. A: N5010; enlarged from PI. 99, Fig. 6 (Text-fig. 9).
Brachyphyllum ex gr. expansum (Sternberg) Seward: IC-3004.

KIMURA, NAITO and OHA NA: Early Jurassic Plants

Plate 100

824. Early Jurassic Plants
juvenile leaves of some Brachyphyllum had
the Geinitzia-form.
References
Bose, M. N. and Banerji, J. (1984): The fossil
floras of Kachchh. I-Mesozoic megafossils.
In 'The fossil floras of Kachchh. Part 1.'
Palaeobotanist, vol. 33, pp. 1-189, incl.
pis. 1-55.
Chaloner, W. G. and Lorch, J. (1960): An opposite leaved conifer from the Jurassic ofIsrael. Palaeontology, vol. 2, pt. 2, pp. 236242, pI. 36.
Harris, T. M. (1961): The Yorkshire Jurassic
flora. I. Thallophyta-Pteridophyta. ix +212 pp. Brit. Mus. (Nat. Hist.).
--(1969): Ditto. III. Bennettitales. vi + 186
pp., 7 pis. Ibid.
-(1979): Ditto. V. Coniferales. 166 pp., 17
pis. Ibid.
Hirano, H. (1971): Biostratigraphic study of theJurassic Toyora Group. Part I. Mem. Fac.
Sci., Kyushu Univ., Ser. D, vol. 21, no. 1,
pp. 53-128, pis. 14-20.
--(1973): Ditto. Part III. Trans. Proc. Palaeont.
Soc. Japan, N. S., no. 90, pp. 45-71, pis.
9-10.
Huzioka, K. (1938): On the occurrence of a new
species of Phlebopteris in Japan. Jour. Fac.
Sci., Hokkaido Imp. Univ., Ser. 4, vol. 4,
nos. 1-2, pp. 143-146.
Kimura, T. and Tsujii, M. (1980a): Early Jurassic
plants in Japan. Part 1. Trans. Proc. Palaeont.
Soc. Japan, N. S., no. 119, pp. 339-358,
pis. 38-40.
--and - - (1980b): Ditto. Part 2. Ibid., no.
120, pp. 449-465, pis. 54-56.
--and - - (1981): Ditto. Part 3. Ibid., no.
124, pp. 187-207, pis. 30-32.
--and - - (1982): Ditto. Part 4. Ibid., no.
125, pp. 259-276, pis. 41-43.
--and - - (1983): Ditto. Part 5. Ibid., no.
129, pp. 35-57, pis. 12-14.
and - - (1984): Ditto. Part 6. Ibid., no.
133, pp. 265-287, pis. 54-56.
Oishi, S. (1935): A new species of Zamites fromthe Nishinakayama bed, Yamaguchi Prefecture. Jour. Fac. Sci., Hokkaido Imp. Univ.,
Ser. 4, vol. 3, no. 1, pp. 97-100.
--(1940): The Mesozoic floras of Japan.

539

Ibid., vol. 5, nos. 2-4, pp. 123-480, pis.
1-48.
Seward, A. C. (1895): Catalogue of the Mesozoic
plants in the British Museum (Natural
History). Part II. Gymnospermae. xi + 259
pp., 20 pis. Brit. Mus. (Nat. Hist.).
--(1971): Fossil plants. Vol. 3. xviii + 656 pp.
Cambridge.
-(1919): Ditto. Vol. 4. xvi + 543 pp. Ibid.
Sze, H. C. (1949): Die mesozoischen Flora ausder Hsiangchi kohlen Series in West Hupei,
China. Palaeont. Sinica, N. S., Ser. A, no. 2,
pp. 1-71, pis. 1-15.
Takahasi, E. (1973a): Liassic flora of the HigashiNagano and the Nishi-Nakayama Formationsof the Toyora Group. Sci. Rep., Yamaguchi
Univ., vol. 20, pp. 1-5, pI. 1 (in Japanese).
-(1973b): Stratigraphic distribution of theMine-type and the Toyora-type floras. Ibid.,
pp. 15-28 (in Japanese).
Wesley, A. (1974): On the bennettitalean remains from the Lias of Northern Italy.
In 'Symposium on morphological andstratigraphical palaeobotany'. Birbal Sahni
Inst. Palaeobot., Spec. Publ., no. 2, pp. 6672, pis. 1-6.
Wu, S. Q., Ye, M. N. and Li, B. X. (1980): Upper
Triassic and Lower and Middle Jurassic
plants from the Hisiangchi Group, Western
Hubei. Mem. Nanjing Inst. Geol. Palaeont.,
Acad. Sinica, no. 14, pp. 63-131, pis. 1-39
(in Chinese with English abstract).
Yabe, H. (1922): Notes on some Mesozoic plantsfrom Japan, Korea and China, in the collection of the Institute of Geology and Palaeontology of the Tohoku Imperial University.

Sci. Rep., Tohoku Im.p Univ., 2nd Ser.,vol. 7, no. 1, pp. 1-28, pis. 1-4.
Yokoyama, M. (1906): Mesozoic plants in China.
Jour. Coli. Sci., Imp. Univ. Japan, vol. 21,
art. 9, pp.1-39, pis. 1-12.
Zhou, Z. Y. (1984): Early Liassic plants fromSouthwest Hunan, China. Palaeont. Sinica,
N. S., Ser. A. no. 7, pp. 1-85, pis. 1-34
(in Chinese with English abstract).
Zigno, A. de (1873-1885): Flora fossilis Formationis Oolithicae. Le Piante fossili dell'
Oolite. Vol. 2. Livr. 1, pp. 1-48, pIs. 26-29(1873): Livr. 2, 3, pp. 49-120, pis. 30-37(1881): Livr. 4, 5, pp. 121-203, pis. 38-42(1885). Padova.

Tatsuaki KIMURA, Gentaro NAITO and Tamiko GHANA

540

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542

SYSTEMATIC INDEX(New Series No. 137-No. 144)

PALAEOZOOLOGY

Protozoa
Page
Goto, Hiroya, Maruoka, Kunitaka and Ishii, Ken-ichi: Lepidolina columbiana (Permian
fusulinid) from British Columbia, Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 422
Ishiga, Hiroaki and Miyamoto, Takami: Follicucullus (Radiolaria) from the Upper Permian
Kuma Formation, Kyushu, Southwest Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322
Coelenterata
Omura, Akio, Tsuji, Yoshinori, Ohmura, Kazuo and Sakuramoto Yuji: New data on
hermatypic corals from the Pleistocene limestone on Kikai, Ryukyu Islands . . . . . . . 196
Mollusca
Ehiro, Masayuki and Bando, Yuji: Late Permian ammonoids from the southern Kitakami
Massif, Northeast Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25
Ehiro, Masayuki, Shimoyama, Shoichi and Murata, Masafumi: Some Permian Cyclolobaceae
from the Southern Kitakami Massif, Northeast Japan . . . . . . . . . . . . . . . . . . . . . . 400
Koike, Hiroko: Microstructure of growth increments in the shell of Mercenaria mercenaria(Linne) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
Kozai, Takeshi: Early Cretaceous Pulsidis (Bivalvia) from Japan . . . . . . . . . . . . . . . . . . 464
Majima, Ryuichi: Intraspecific variation in three species of Glossaulax (Gastropoda:
Naticidae) from the Late Cenozoic strata in central and southwest Japan . . . . . . . . . 111
Matsuda, Tomoko: Bivalve fauna from the Miyanohara Formation (Lower Cenomanian)of Sakawa area, Shikoku . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1Matsui, Seiichiro: Recurrent molluscan associations of the Omma-Manganji fauna in the
Gojome-Oga area, Northeast Honshu, Part 1. General discussion of fauna and systematic notes on gastropod and scaphopod species . . . . . . . . . . . . . . . . . . . . . . . . . 149
Matsui, Seiichiro: Recurrent molluscan associations of the Omma-Manganji fauna in the
Gojome-Oga area, Northeast Honshu, Part 2. Systematic notes on bivalve species .... 225
Matsumoto, Tatsuro and Noda, Masayuki: A note on an inoceramid species (Bivalvia)
from the Lower Coniacian (Cretaceous) of Hokkaido . . . . . . . . . . . . . . . . . . . . . . 263
Matsumoto, Tatsuro and Noda, Masayuki: Some inoceramids (Bivalvia) from the Cenomanian (Cretaceous) of Japan - 1. New or little known four species from Hokkaido
and Kyushu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
Matsumoto, Tatsuro, Muramoto, Kikuwo, Takahashi, Takemi, Yamashita, Minoru andKawashita, Yoshitaro: On Neocrioceras spinigerum (Jimbo), a species of Cretaceous
heteromorph ammonoids (Studies of the Cretaceous ammonites from HokkaidoLV) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463
Matsuoka, Keiji: Pliocene freshwater gastropods from the Iga Formation of the Kobiwako
Group, Mie Prefecture, Central Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

543
Noda, Masayuki: A new species of Inoceramus (Cordiceramus) (Bivalvia) from the Upper
Coniacian (Cretaceous) of Hokkaido . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Ohtsuka, Yasuo: Early internal shell microstructure of some Mesozoic Ammonoidea:
Implication for higher taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Sato, Tadashi, Kasahara, Yoshio and Wakita, Koji: Discovery of a Middle Jurassic ammonite Kepplerites from the Mino Belt, Central Japan . . . . . . . . . . . . . . . . . . . . . . 218
Tashiro, Masayuki and Matsuda, Tomoko: Lower Cretaceous bivalves from the Sakawa
area, Shikoku . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366
Arthropoda
Hu, Chung-Hung: Ontogenies of two Middle Cambrian corynexochid trilobites from the
Canadian Rocky Mountains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Ishizaki, Kunihiro: Ostracoda from the Pliocene Ananai Formation, Shikoku, Japan. . . .. 50
Kaneko, Atsushi: A Middle Devonian trilobite fauna from the Kitakami Mountains, Northeast Japan - II. The Calymenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Kobayashi, Teiichi and Hamada, Takashi: Additional Silurian trilobites to the YokokuraYama fauna from Shikoku, Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
Kobayashi, Teiichi and Hamada, Takashi: The second addition to the Silurian trilobite
fauna of Yokokura-yama, Shikoku, Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447

Bryozoa
Nishizawa, Yasuo: Multilaminar colonies of bryozoans from Japan, 1. "Ectoproctalith" ofAntropora tincta (Hastings) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Brachiopoda
Tazawa, Jun-ichi and Kato, Makoto: Striatifera and Gigantoproductus from the Lower
Carboniferous of Fukuji, Central Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 393
Conodonts
Igo, Hisaharu and Hisada, Ken-ichiro: Lower Permian conodonts from the Kawanori Formation of the Southwest Kanto Mountains, Tokyo . . . . . . . . . . . . . . . . . . . . . . . . . . 516

Mammalia
Takahashi, Shizuo, Domning, Daryl, P. and Saito, Tsunemasa: A new ancester of Steller's
sae cow from the Upper Miocene of Yamagata Prefecture, Northeastern Japan . . . . . . 296

PALEOBOTANY
Fuji, Norio: Palynological study of 200-meter core samples from Lake Biwa, Central Japan,
II. The palaeovegetational and palaeoclimatic changes during the CA. 25,000 -100,000
years B.P . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 490
Horiuchi, Junji and Kimura, Tatsuaki: Ginkgo tzagajanica Samylina from the Paleogene
Noda Group, Northeast Japan, with special reference to its external morphology andcuticular features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341

544
Ling, Hsin, Yi: Early Paleogene silicoflagillates and ebridians from the Arctic Oceans
79
Kimura, Tatsuaki, Naito, Gentaro and Ohana, Tamiko: Early Jurassic plants in Japan.
Part 7. Fossil plants from the Nishinakayama Formation, Toyora Group, Yamaguchi
Prefecture, Southwest, Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 528
Matsuoka, Kazumi: Cyst and thecate forms of Pyrophacus steinii (Schiller) Wall et Dale,
1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240
Nishida, Harufumi: On a new Tempskya stem from Japan . . . . . . . . . . . . . . . . . . . . . . 435
Suzuki, Keiji: Larix remains from Pleistocene strata of Northeast Japan, with specialreference to the distribution of Larix in the later half of the Last Glacial age . . . . . .. 64

545

INDEX OF GENERA AND SPECIES
Notes: Words listed are names of genus and species-groups, together with more higher taxa
which are described, discussed or illustrated in the articles in Transaction and Proceedings of
the Palaeontological Society of Japan, New Series, Nos. 137-144. Words in bold-faced type arenames of newly proposed taxa.

A

Acharax tokunagai . . . . . .
228, pI. 31
Acila (Acila) divaricata . . . . . . 225, pI. 31
Acila (Truncacila) nakazimai .. . 225, pI. 31
Acila (Truncacila) sp. . . . . . . . 225, pl. 31
Admete lischkei ....
175, pI. 22
Albaillella triangularis . . . . . . . . . , pI. 64
Alvertella limbata . . . . . . . . . . . . . . .
· . . . .. 141,143, pI. 21, Text-figs. 2A-N
Alvania sp. . . . . . . . . . . . . . . 171, pI. 22
Ammodochium fletcheri ... 85, pIs. 11, 13
Ammodochium rectangulare . . . . 85, pI. 11
Ammonitina . . . . . . . . . . . . . . . . . . 283
Anadara amicula . . . . . . . . . . . 228, pI. 32
Anagaudryceras yokoyamai . . . . . . . .
· . '.' . . . . . . . . . . . . . . 282, pIs. 45, 46
Anaklinoceras . . . . . . . . . . . . . . . . . 471
Ancyloceras kossmati . . . . . . . . . . . . 468
Anthonya apicalis . . . . . . . . . . . . . 9, pI. 1
Antiplanes contraria . . . . . . . . 177, pI. 22
Antiplanes (Rectiplanes) sanctiioannis
177, pI. 22
Antropora tincta ...
20, pI. 4
Apolichas perconvexus ...
215, pI. 30
Apolichas truncatus
459, pI. 92, Text-figs. Bl-2
Araxoceras cf. rotoides . . . . . . . . 40, pI. 7
Araxoceras sp . . . . . . . . . . . . .
41, pI. 7
Arca (Eonavicula) prolata . . . . . . .
371, pI. 74, Text-fig. 3
Arca (Eonavicula) toshiroi . . . . . . . . .
4, pI. 2, Text-fig. 2
Astarte spo cf. A. (Trautscholdia) minor
381, pI. 77
Astarte (Astarte) subsenecta . . . . . . . .
· . . . . . . . . . . . . . . . . . 381, pIs. 75, 77
Aurila hataii .
53
Aurila munechikai . . . . . . . . . . . . . . 53••••••••

0

o'

0

•••

0

o.

•

0

0"

0

Bellamya . ...
. . . . . . . . . . . 186
Bellamya 8uzukii . . . . . .
186, pI. 26
Bicromucythere bisanensis . . . . . . . .. 53
Birostrina . . . . . . . . . . . . . . . . . . . . 411
Birostrina tamurai . . . . . . . 411, pIs. 81-83
Birostorina cf. B. tamurai .
. 11, pI. 85
Bittium binodulosum .
173, pI. 22
Biwamelania . . . . . . . . .
. .... 189
Boreotrophon xestra . . . . . . . . 174, pI. 22
Brachyphyllum ex. gr. expansum . . . . .
· . . . . . . .. 529, pIs. 99-100, Text-fig. 8
Bradleya nuda .... . . . .
. . . .. 52
Buccinum cf. rossicum .
174, pI. 23
pI. 30
Bumastus agmakros .... .
pI. 30
Bumastus kattoi . . . . . . .
pI. 30
Bumastus sakoi .
0

•••

••

0

0

0

0

•

••

0

••••••

0

•

0

•

•••

••••••••o•••••••

0

•

0

•••••

o

•••••••

0

0

•

0

0

•••

••••••0BC• • • • • • • •••••••••

0

Australoscutellum . . . . . . . . . . . . . . . 448
Axinopsida subquadrata . . . . . . 230, pI. 33
Axonoceras . . . .
. . . . . . . . . . . 471

0

0

0

•••••••••

0•

0

Caestocorbula minima . . . . . . . 384, pI. 77
Caestocorbula sp. cf. Co shikamai . . . . .
· . . . . . . . . . . . . . . . . . . . . 385, pI. 77
Callistocythere kattoi . . . . . . . . . . . . 53
Calymene antigonishensis . . . . . . . . pI. 14
Chlamys islandica . . . . . . . . . . 229, pI. 31
Clinocardium chikagawaense ... 231, pI. 33
Clinocardium fastosum ..
231, pI. 33
Clinopegma borealis
175, pI. 23
Collignoniceratidae . . . . .
. . . . . 284
Corbisema apiculata ....
. 81, pI. 10
Corbisema hastata hastata . . . . . 81, pI. 10
Corbisema triacantha . . . . . . 81, pIs. 10, 12
Corbisema sp. cf. C. inermis minor.
· . . . . . . . . . . . . . . . . . . . . . 81, pI. 10
0

••

546

Cosmetodon tomochiensis . . . . . . . . .
· . . . . . . . . . 369, pIs. 74, 76, Text-fig. 2
Crassostrea kawauchidensis . . . . . . . 6, pI. 2
Crenulilimopsis oblonga . . . . . . 228, pI. 32
Cryptonatica clausa . . . . . . . . . 173, pI. 22
Cucullaea (Idonearca) ezoensis ezoensis· . . . . . . . . . . . . . . . . . . . . . . . 3, pI. 2
Cucullaea obliquata . . . . . . . . . : ... .
· . . . . . . . . . . . . 372, pI. 74, Text-fig. 4
Cupressinodadus sp. A . . . . . . . . . . . .
· . . . . . . . . . q29, pIs. 99-100, Text-fig. 9
Cupressinodadus sp. B . . . . . . . . . . . .
· . . . . . . . . . . . . . . . . 529, Text-fig. 10
Cyclocardia aomoriensis . . . . . . 230, pI. 33
Cyclocardia fen''uginea . . . . . . . 231, pI. 33
Cyclocardia sp . . . . . . . . . . . . . 231, pI. 33
Cyclolobus cf. C. walkeri . . . . . . . . .
· . . . . . . . . . . . . . . 37, pI. 5, Text-fig. 7
Cyphoproetus latiaxis . . . . . . . 453, pI. 91
Cytheropteron miurense . . . . . . . . . . 53
Cytheropteron uchioi . . . . . . . . . . . . 52
D

Damesites damesi . . . . . . . 283, pIs. 45, 47
Damesites semicostatus . . . . . . . . . pI. 50
Damesites sugata : . . . . . . . . . . . . pI. 48
Dactylioceratidae . . . . . . . . . . . . . . . 284
Desmocerataceae . . . . . . . . . . . . . . . 283
Desmoceras japonicum . . . . . . 283, pI. 49
Dicranopeltis . . . . . . . . . . . . . . . . . . 457
Dicranopeltis tricornis . . . . . . . . . . . .
· . . . . . . . . . 458, pI. 92, Text-figs. Cl-3
Dictyocha arctios . . . . . . . . . . . 82, pI. 10
Dictyoxha curta . . . . . . . . . . . . 83, pI. 10
Dictyocha deflandrei . . . . . . 83, pIs. 10, 11
Dictyocha fibula . . . . . . . . . . . . 84, pI. 11
Dictyocha obliqua . . . . . . . . . . 84, pI. 11
Dictyocha spinosa . . . . . . . . . . . 84, pI. 11
Dictyocha sp. cf. D. carentis incerta· . . . . . . . . . . . . . . . . . . 83, pIs. 10, 13
Dictyocha sp. cf. D. rotuncata . .. 84, pI. 11
Dictyocha sp . . . . . . . . . . . . 81, pIs. 10, 12
Dindymene (?) megacranidia ... 213, pI. 28
Dindymene . . . . . . . . . . . . . . . . . . . 455
Dindymene (?) sp. . . . . . . . . . 455, pI. 91
Dindymeninae . . . . . . . . . . . . . . . . . 455
Diplognathodus sp. A . . . . . . . 529, pI. 98
Distephanus speculum . . . . . . . . 84, pI. 11
Dosinia (Phacosoma)japonica .. 232, pI. 34
Dusisiren dewana . . . . . . . . . . . . . . .

. 300, pIs. 53-62, Text-figs. 3-15

E
Ebriopsis cornuta . . . . . . . . 85, pIs. 11, 13
Elaeocyma (Splendrillia) braunsi . . . . .
· . . . . . . . . . . . . . . . . . . . . 176, pI. 22
Elatides sp. . . . . . . . . . . . . . . . . . . .
· . . . . . . . . 529, pIs. 99-100, Text-fig. 11
Eleganticeras elegantulum . . . . . . . . .
· . . . . . . . . . . . . . . . . . 284, pIs. 46, 47
Encrinurinae . . . . . . . . . . . . . . . . . . 209
Encrinurus . . . . . . . . . . . . . . . . . . . 209
Encrinurus similis . . . . . . . . . . 212, pI. 29
Encrinurus stenorhachis . . . . . . 212, pI. 29
Encrinurus subtrigonalis . .. 210, pIs. 28, 29
Eumedlicottia sp . . . . 40, pI. 6, Text-fig. 8E
Epitonium (Boreoscala) echigonum ...
. . . . . . . . . . . . . . . . . . . . . 178,pI.22
Exiteloceras . . . . . . . . . . . . . . . . . . . 471
F

Favia . . . . . . . . . .
. . . . . . . . . . . 197
Favites . . . . . . . . . . . . . . . . . . . . . . 197
Felaniella usta . . . . . . . . . . . . 230, pI. 33
Fieldaspis quadriangularis . . . . . . . . • .
· . . . .. 139, 140, pI. 20, Text-figs. 1A-N
Follicucullus bipartitus ... 326, 332, pI. 63
Follicucullus charveti . . . . . . . . . . . .
· . . . . . . . . . . . . . 327,332, pIs. 63, 64
Follicucullus charveti charveti . . . . . . .
· . . . . . . . . . . . . . . . . . 328, pIs. 63, 64
Follicucullus charveti falx .... 329, pI. 64
Follicucullus charveti orthogonus ....
· . . . . . . . . . . . . . . . . . . . . 329, pI. 64
Follicucullus scholasticus . . . . . 330, pI. 64
Follicucullus sp. . . . . . . . . . . . . .. pI. 64
Fragilidinium heterolobum . . . . . . . . . 243
Fulvia mutica . . . . . . . . . . . . . 231, pI. 33
Fusitriton oregonensis . . . . . . . 173, pI. 22
G

Galaxea.197
Gaudryceras striatum . . . . . . . . . . .
· . . . . . . . . 282, 283, pIs. 45,46,48, 50
Gaudryceratidae . . . . . . . . . . . . . . . . 282
Geinitzia sp. . . . . . . . . . 538, Text-fig. 12
Gervillaria haradae . . . . . . . . . 373, pI. 75
Gervillaria (Gervillaria) sp. cf. G. (G.)
forbesiana . . . . . . . . . . . . . . 374, pI. 75
Gigantoproductus sp. . . . . . . . 396, pI. 78

547

Ginkgo . . . . . . . . . . . . . . . . . . 344,347
Ginkgo biloba . . . . . . . . . . . . 349, pI. 67
Ginkgo tzagajanica . . . . . . . . . . . . . .
· . . . .. 344, pIs. 65-68, Text-figs. 2, 3c-j
Ginkgoites . . . . . . . . . . . . . . . . . . . . 344
Glossaulax . . . . . . . . . . . . . . . . 116, 123
Glossaulax hagenoshitensis . . . . . . . . .
· . . . . . . . . . . . . . . . . . 131, pis. 17-19
Glossaulax hyugensis . . . . . 129, pIs. 17-18
Glossaulax nodai . . . . . . . . . . 131, pI. 18
Glossaulax reiniana . . . . . . . . . . .. pI. 17
Glycymeris derelicta . . . . . . . . 228, pI. 32
Glycymeris (Hanaia) goshonouraensis· . . . . . . . . . . . . . . . . . . . . .. 5, pI. 1
Glycymeris yessoensis . . . . .
229, pI. 32
Goldillaeninae . . . . . . . . .
. . . . . 451
Goniastrea . . . . . . . . . .
. .... 197
Goshoraia crenulata. . . . . . .. 9, pI. 3
H

Haplentatinia (?) ichikawai . . . . . .. pI. 64
Hildoceratidae . . . . . . . . . . . . . . . . . 284
Hindeodus minutus . . . . . . . . . 519, pI. 98
Homalopoma amuseltatum .... 171, pI. 22
Hypophylloceras subramosum . . . . . . .
· .......... ".283, pIs. 45, 47-49
I

Ichiyamella . . . . .
. . 456
Ichiyamella megacranidia .
. . 456
Ichiyamella subglobula . . . . . . . . . . .
· . . . . . . . . . 456, pI. 91, Text-figs. Al-4
Igapaludina . . . . . . . . . . . . . . . . . . . 184
Igapaludina stricta . . . . . . 184, pis. 24-26
Illaenidae . . . . . . . . . . . . . . . . . . . . 450
Illaenoid (?) gen. et sp. indet.
. .. pI. 29
IlIaenoides (?) abnormis . . . . . . 453, pI. 90
IlIaenoides (?) magnisulcatus . . . . . . . .
· . . . . . . .
. . . . . . . . 452, pI. 90
Inocermidae . . . . . .
. . . . . 410
Inoceramus . . . . . . . . . . . . . . . . . . . 264
Inoceramus anglicus . . . . . . . . 375, pI. 76
Inoceramus heinzi . . . . . . . 415, pis. 83,85
Inoceramus (Cordiceramus) kawashitai· . . . . . . . . . 355, pis. 69-73, Text-fig. 6
Inoceramus rotundatus ... 264, pIs. 41-44
Inoceramus takahashii . . . . . . . 414, pI. 84
Inoceramus sp. aff. 1. crippsi ... 375, pI. 76
Inoceramus (Cordiceramus) sp. cf. I.
(C.) kawashitai . . . . . . . . . . . . .. pI. 73

Inoceramus sp . . . . . . .
Isognomon sp. . .. .

376, pI. 76
375, pI. 75

JJapanoscutellum genicula tum . . . . . . .
· . . . . . . . . . . . . . . . . . . . . 208, pI. 28
Japonoscuttelum japonicum puteatum· . . . . . . . . . . . . . . . . . . . . 448, pI. 90
Japonoscuttelum japonicum laticephalum ...
449, pI. 90
0

•

•

•

•

•

•

0

•••

K
Kawashitaceras . .
. .......
469
Kepplerites (Seymourites) sp . . . . . . . .
219, Text-fig. 2
Kitchinites japonicus . . . . . . .. pis. 49, 50
0••••••••••

0•••••••••

L

Laevicardium ? corpulentum ... 383, pl._77
Laevicardium ? ishidoense
383, pI. 77
Lamposcutellum . . . . . . . . . . . . . . . . 448
Larix kaempferi . . . . . . . . . . . . . 69, pI. 9
Larix kamtschatica . . . . . .
68, pis. 8-9
Larix cf. kaempferi . . . . . . . . . . . 70, pI. 9? Legumen sp. . ... _ . . . . . . .
10, pI. 3
Leioscuttelum . . . . . . .
448
Lepidolina . . . . . . . . . .
424
Lepidolina columbiana . . . . . . . . . . .
431, pI. 86, Text-fig. 3b
Lepidolina multiseptata . 431, Text-fig. 3a
Leptosolen spo ...
382
Lewyites . . . . . . . . . . .
470
Lichid (?) hypostoma, gen. et sp. indet.
. . . . . . . . . . . . . . . . . . . . . 215,pI.30
Lichidae . . . . . . . . . . . . . . . . . . . . . 457
Limopsis tokaiensis . . . . . . . . . 228, pI. 32
Limopsis sp. A ...
o. pI. 32
Limopsis sp. B . . . . . . . . . . . . . .. pI. 32
Loxoconcha kattoi . . . . .
52
Loxoconcha sinensis .
53
Lucinoma acutilineata . . . . . . . 230, pI. 32
Lucinoma ? sp . . . . . . . . .
380, pI. 77
Lunatia pal/ida . .
. . . . . . 173, pI. 22
0

••

0

•••••••

0••••••

0

•

0

•

0

•••••••••

•••••

0

0

•••••••••••••

0

0

•••

0

•••••

•••••••••

00•••••••••••••••

0

0

•••••

•••

M

Macoma calcarea. . . . . . 232, pI. 34
Macoma tokyoensis . . . . . . . . . 232, pI. 34
Medlicottia . . . . . . . . . . .
38
Medlicottia kitakamiensis . . . . . . . . . .
. . . . . . . . . .. 39, pI. 6, Text-figs. 8A-D
0

•

•

•

•

•

••

548

Meitanillaenus binodosus . . . . . . . . . . 452
Meitanillaenus zhitangensis . . . . . . . . . 452
Menuites sp. . . . . . . . . . . . . . 283, pI. 45
Mercenaria mercenaria . . .. 290, pis. 51-52
Mercenaria stimpsoni . . . . . . . . 232, pI. 34

Neverita incei . . . . . . . . . . . . . . .. pi. 17
Nipponitrigonia kukuchiana . . . . . . . . 378
Nipponocalymene. . . . . . . . . . . . . .. 94
Nipponocalymene hamadai . . . . . . . . .

Meroperixinae . . . . . . . . . . . . . . . . . 450
Merrillina sp. A . . . . . . . . . . . 520, pi. 98
Microstelma sp. . . . . . . . . . . . 172, pI. 22
Miodontiscus nakamurai ..... 231, pI. 33
Mizuhopecten poculum ...... 229, pi. 31
Modiolus falcatus . . . . . . . . . . 373, pi. 75
Mohnia cf. yanamii . . . . . . . . . 174, pI. 23
Montastrea . . . . . . . . . . . . . . . . . . . 197
Montipora . . . . . . . . . . . . . . . . . . . . 197
Mya (Mya) cuneiformis ...... 233, pi. 34
Myadora japonica . . . . . . . . . . 233, pI. 34

Nucula (Pectinucula) kochiensis . ... 2, pi. 1
Nucula (Leionucula) niponica . . . . . . .

Myrtea (Myrtea) amanoi .......•..
· . . . . . . . . . . . . . . . 7, pI. 1, Text-fig. 3

Mytiloides . . . . . . . . . . . . . . . . . . . . 417
Mytiloides mikasaensis . ...... 417, pI. 85
N

Nassarius (Zeuxis) siquijorensis ..... .
. . . . . . . . . . . . . . . . . . . . . 175,pI.22
Nazarovella gracilis . . . . . . . . . . " pI. 64

Neithea (Neithea) syriaca amanoi .
· . . . . . . . . . . . . . . . • . . . . 376, pI. 74

Nemodon sp . . . . . . . . . . . . . ". 370, pI. 74
Neocrioceras . . . . . . . . . . . . . . . . . . 469
Neocrioceras (Schueterella) compressum· . . . . . . . . . . . . . . . . . . . . . . . . . 467

Neocrioceras (Schueterella) mutinodosum· . . . . . . . . . . . . . . . . . . . . . . . . . 468

Neocriocer.as (Neocrioceras) maderi ... 468
Neocrioceras (?) undulosum . . . . . . . . 468
Neocrioceras spinigerum . . . . . . . . . .
· . . . . . . . . . 463, pis. 93, 94, Text-fig. 1

Neogloboquadrina pachyderm a ..... .
· . . . . . . . . . . . . . . . . . . 168, 170, 171

Neogondolella bisselli . . . . . . . 520, pI. 97
Neogondolella gujioensis ..... 521, pI. 97
Neogondolella idahoensis ..•.. 521, pI. 97
Neogondolella sp. A . . . . . . . . 522, pI. 96
Neonesidea villosa . . . . . . . . . . . . . . . . 53
Neostreptognathodus exsculptus .... .
· . . . . . . . . . . . . . . . . . . . . 522, pi. 98

Neostreptognathodus ? sp. . . . . . . . . .
· . . . . . . . . . . . . . . . . . . . . 523, pI. 98

Neptunea intersculpta . . . . . . . 175, pI. 23
Neverita (G/ossaulax) didyma .. 173, pI. 22· . . . . . . . . 99, pis. 14-16, Text-figs. 1a-b

· . . . . . . . . . . . . . . . . . . . . 225, pI. 31

Nuculana (Robaia) robai ..... 226, pI. 31
Nuculana sadoensis . . . . . . . . . 226, pI. 31
Nuculana yokoyamai . . . . . . . . 226, pI. 31
Nuculopsis (Palaeonucula) ishidoensis· . . . . . . . . . . . . . . . . . . . . 368, pi. 74

oOenopota kagana . . . . . . . . . . 178, pI. 22
Ophiodermella pseudopannus .. 11 7, pI. 22
Opoa (?) trinodosa . . . . . . . . . 450, pi. 90
Otozamites micropinnatus . . . . . . . . .
· . . . . . . 530, pis. 99-100, Text-figs. 2a-c

Otozamites sp. A . . . . . . . . 531, Text-fig. 3
Otozamites sp. B . . . . . . . . . . . . . . . .
· . . . . . . . . .. 533, pI. 100, Text-fig. 10

Oxybeloceras . . . . . . . . . . . . . . . . . . 471
P

Pachythaerus nagaoi . . . . . . . . . . . 8, pI. 1
Palliolum (Delectopecten) peckhami ..
· . . . . . . . . . . . . . . . . . . . . 229, pI. 31

Pandora (Pandorella) wardiana ..... .
· . . . . . . . . . . . . . . . . . . . . 233, pI. 34

Parasolenoceras . . . . . . . . . . . . . . . . 470
Paradrillia cf. dainichiensis .... 176, pI. 22
Paratirolites ? sp. . .. 45, pI. 7, Text-fig. 9E
Parvamussium hinagense ..... 376, pi. 76
Parvamussium kattoi . . . . . . . . 378, pi. 76
Parvamussium tosaense . . . . . . 377, pi. 76
Periploma japonica . . . . . . . . . . . 13, pi. 3
Periploma plane . . . . . . . . . . . 233, pi. 34
Periploma cf. besshoensis ..... 233, pI. 34
Peronoceras fibula tum . . .. 284, pis. 46-47
Phlebopteris sp. . . . . . 529, Text-figs. 1a-d
Pholadomya (Bucardiomya) hiratai ...
· . . . . . . . . . . . . . . 11, pi. 3, Text-fig. 4
Pholadomya (Pholadomya) japonica ...
· . . . . . . . . . . . . . . . . . . . . . . 10, pi. 3
Phylloceratidae . . . . . . . . . . . . . . . . 283
Pinna (Pinna) sp. cf. P. robinaldina ... .
· . . . . . . . . . . . . . . . . . . . . 373, pi. 75
Pianiscutelluinae . . . . . . . . . . . . . . . . 447

549

Plicifusus ozawai . . . . . . . . . . 174, pI. 23
Plicatula sp. . . . . . . . . .. .., 378, pI. 76
Polin ices sagamiensis . . . . . . . . . .. pI. 17
Pontocythere japonica . . . . . . . . . . .. 53
Porites . . . . . . . . . . . . . . . . . . . . . . 197
Portlandia hirosakiensis . . . . . . 227, pI. 31
Portlandia (s.l.) nagaseana. . . . . .. 3, pI. 1
Portlandia sanchuensis . . . . . . . 368, pI. 74
Portlandia toyamaensis . . . . . . . 27, pI. 31
Proetus (Gerastos) sugiharensis . . . . . .
· . . . . . . . . . . . . . . . . . . . . 208, pI. 29
Propebela candida . . . . . . . . . . 178, pI. 22
Protocardia (Protocardia) amanoi ....
· . . . . . . . . . . . . 382, pI. 77, Text-fig. 5
Pro to toceras japonicum . . . . . . . . . . .
· . . . . . . . . . . . . . 42, pI. 7, Text-fig. 9A
Prototoceras sp. A .. 42, pI. 7, Text-fig. 9B
Prototoceras sp. B . . . . . . . . . . . . 42, pI. 7
Prototoceras sp. C ... 42, pI. 7, Text-fig. 9C
Protothaca (Ptotocal/ithaca) adamsi ...
· . . . . . . . . . . . . . . . . . . . . 233, pI. 34
Pseudammodochium dictyoides . . . . . .
· . . . . . . . . . . . . . . . . . . . . . 85, pI. 11
Pseudocheirurus gomiensis • . . . . . . . .
· . . . . . . . . . . . . . . . . . . . . 208, pI. 28
Pseudoctenis sp . . . . . . . . . . . . . . . . .
· . . . . . . . . . 535, pI. 100, Text-figs. 7a-b
Pseudogastrioceras sp . . . . . . . . . . 38, pI. 5
Pseudoxybeloceras . . . . . . . . . . . . . . 470
Pterotrigonia (Ptilotrigonia) amakusensis................... 6
Pterotrigonia (Ptilotrigonia) mifunensis· ..........................

6

Pterotrigonia (Ptilotrigonia) miyanoharens is . . . . . . . . . . . . 0 0 . . . . . 0 . . .. 6
Pterotrigonia (? Scabrotrigonia) moriana· . . . . . . . . . . . . . . . . . . 0 . 380, pI. 74
Pterotrigonia (Pterotrigonia) pocilliformis . . . . . . . . . . . . . . . . . . . 379, pI. 74
Pterotrigonia (? Scabrotrigonia) pustulosa . . . . . . . . 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6
Pterotrigonia (? Scabrotrigonia) SPo 0 0 0380
Ptilillaenus lojopingensis 0 . . . . . . . . . 452
Pulsidis . . . . . . . . . . . . . . . . . . . . . . 481
Pulsidis angulata . . . . . . . . . . . 483, pI. 95
Pulsidis nagatoensis . . . . . . . . . 482, pI. 95
Pulsides rostrata . . . . . . . . . . . 486, pI. 95
Pulsidis sanchuensis . . . . . . . . . 484, pI. 95
Pulsidis tashiroi . . . . . . . . . . . 485. pI. 95
Pyrophacus . . . . . . . . . . . . . .. 240, 243

Pyrophacus steinii . . . . . . . . . . . . .. 243,
257,259, pIs. 35-40, Text-figs. 3a-d, 6,8
R

Reesidites minimus
Reticunassa cf. spurca .

284, pI. 45
175, pI. 22

S
Saccel/a (Costanuculana) husamaru· . . . . . . . . . . . . . . . . . . . . . .. pI. 31
Saccel/a (Costanuculana) onoyamai ...
· . . . . . . . . . . . . . . . . . . . . 226, pI. 31
Schizocythere kishinouyei . . . . . . . 52, 53
Schlueterel/a . . . . . . . . . . . . . . . . . . 469
Scutelloid gen. et sp. indet ..... 449, pI. 90
Semisulcospira . . . . . . . . . . . . . . . . 189
Semisulcospira(Biwamelania)
praemultigranosa . . . . . . . . . . . . 190, pI. 27
Siphonodentalium sp. A . . . . . . 179, pI. 23
Siphonodentalium sp. B . . . . . . 179, pI. 23
Solamen spectabilis . . . . . . . . . 229, pI. 31
Sphaerxochinae .. . . . .
. . . . . . . 454
Sphaerxochus . . . . . . . . . . . . . . . . . 454
Sphaerxochus hiratai . . . . . . . . 454, pI. 90
Stacheoceras iwaizakiense . . . . . . . . .
· . . . . . . . . . . . . . 36, pI. 5, Text-fig. 6A
Stacheoceras otomoi . . . . . . . . . . . . .
· . . . . . . . . . . . 404, pI. 79, Text-fig. 3A
Stacheoceras sp . . . . . . . . . . . . . . . . .
· . . . . . . . . . 404, pI. 79, Text-figs. 3B-C
Staurocephalidae . . . . . . . . . . . . . . . 213
Staurocephalus . . . . . . . . . . . . . . . . . 213
Staurocephalus trichochin .... 214, pI. 30
Stephanosel/a vitrea . . . . . . . . . . 21, pI. 4
Streptognathodus elongatus .. .
523, pI. 98
395, pI. 78
Striatifera striata . . . . . .
178, pI. 23
Striodentalium rhabdotum ..
Suavodrillia ? sp. A . . . . . . . . . 177, pI. 22
Suavodrillia ? sp. B . . . . . . . . . 177, pI. 22
Sweetognathus whitei .. .
523, pI. 97

T. . . . . . . 435,440
Tempskya . . . . . .
Tempskya iwatensis . . . . . . . . . . . . .
· . . . . . . . .. 437, pIs. 87-89, Text-fig. 2
Tetragonitaceae . . . . . . . . . . . . . . . . 282
Tetragonites glabrus . . . . . . . . . . . . .
· . . . . . . . . 280, 282, 283, pIs. 46, 48-49
Tetragonitidae . . . . . . . . . . . . . . . . . 283

550

Thetis japonica . . . . . . . . . . . . .. 8, pI. 2
Thracia kakumana . . . . . . . . . 233, pI. 34
Thyasira (Conchocele) bisecta . . . . . . .
· . . . . . . . . . . . . . . . . . . . . 230, pI. 32
Thyasira (Thyasira) tokunagai . . . . . . .
· . . . . . . . . . . . . . . . . . . . . 230, pI. 32
Timorites intermedium . . . . . . . . . . .
· . . . . . . 405, pIs. 79,80, Text-figs. 4A-B
Trachyleberis lungkangensis ... . . . .. 53
Tridonta borealis . . . . . . . . . . 231, pI. 33
Trophonopsis kagaensis . . . . . . 174, pI. 22
Tuberculodinium . . . . . . . . . . . . . . . 241
Tuberculodinium vancampoae . . . . . . .
· . . . . . . . . . . . . . . .. 244, 252, pI. 35
Tulotomoides . . . . . . . . . . . . . . . . . 187
Tulotomoides sanaguensis . . . . . . . . .
· . . . . . . . . . . . . 188, pI. 27, Text-fig. 3
Turbonilla (Ptycheulimella) inscitula ..
· . . . . . . . . . . . . . . . . . . . . 178, pI. 22
Turritella (Neohaustator) nipponica ...
· . . . . . . . . . . . . . . . . . . . . 173, pI. 22
Turritella (Neohaustator) saishuensis ..
· . . . . . . . . . . . . . . . . . . . . 172, pI. 22
U

Unnamed spumellaria . . . . . . . . . . pI. 64. . . . . 448

Un icapeltisv
Verminaria areolae. . . . . . . 21, pI. 4
Volutomitra alaskana . . . . . . . 175, pI. 22x
Xaniognathus abstractus
523, pI. 98
Xenocardita amanoi . . . . . . . . 380, pI. 75
Xenodiscus cf. X. carbonarius . . . . . . .
· . . . . . . . . . . . . . 43, pI. 7, Text-fig. 9DyYabeina . . . . . . . . . . . . . . . . . . . . . 424
Yabeina globosa . . . . . . . 425, Text-fig. 3cYoldia (Cnesterium)johanni ... 227, pI. 31
Yoldia (Megayoldia) thraciaeformis ...
. . . . . . . . . . . . . . . . . . . . . 227,pI.31
Yoldia (Yoldia) kikuchii . . . . . 227, pI. 31

z
Zamites toyoraensis........ .
· . . . . . . . . . 533, pI. 100, Text-figs. 5a-b
Zamites sp . . . . . . . . . . . . . . . . . . . .
· . . . . . . . .. 534, pI. 99, Text-figs. 6a-b

a~"'~1~i-~.:k·~~fTransactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 137 ""No. 144
1985-1986

Palaeontological Society of Japan

The heading in Japanese commemorates the handwriting of Prof. Matajiro YOKOYAMA,
father of Japanese palaeontology, who was a professor of stratigraphy and palaeontology at
the Geological Institute, Imperial University of Tokyo.
The fossil on the cover is Unuma (Spinunuma) echinatus ICHIKAWA andYAO, a Middle Jurassic multisegmented radiolaria from Unuma, Gifu Prefecture, central Japan (photo by A. YAO, x 260).

CONTENTS
Number 137 (published April 30, 1985)
Transactions
Article790. Matsuda, Tomoko: Bivalve fauna from the Miyanohara Formation

Page

Plate

(Lower Cenomanian) of Sakawa Area, Shikoku . . . . . . . . . . . . . . .
791. Nishizawa, Yasuo: Multilaminar colonies of bryozoans from Japan.

1-18

1-3

1. "Ectoproctalith" of Antropora tincta (Hastings) . . . . . . . . . . . ..
792. Ehiro, Masayuki and Bando, Yuji: Late Permian ammonoids from the

19-24

4

southern Kitakami Massif, Northeast Japan . . . . . . . . . . . . . . . . ..

25-49

5-7

793. Ishizaki, Kunihiro: Ostracoda from the Pliocene Ananai Formation,
Shikoku, Japan - Faunal analyses - . . . . . . . . . . . . . . . . . . . . .
794. Suzuki, Keiji: Larix ramains from Pleistocene strata of Northeast
Japan, with special reference to the distribution of Larix in the later
half of the Last Glacial age . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

64-74

Proceedings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..

75-78·

50--638-9

Number 138 (published July 15, 1985)
Transactions

795. Ling, Hsin, Yi: Early Paleogene silicoflagillates and ebridians from the
79-93
Arctic Oceans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
796. Kaneko, Atsushi: A Middle Devonian trilobite fauna from the Kitakami Mountains, Northeast Japan-II. The Calymenidae . . . . . . . . . . 94-110
797. Majima, Ryuichi: Intraspecific variation in three species of Glossaulax(Gastropoda: Naticidae) from the Late Cenozoic strata in central andsouthwest Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111-137
798. Hu, Chung-Hung: Ontogenies of two Middle Cambrian corynexochid
trilobites from the Canadian Rocky Mountains . . . . . . . . . . . . . . . . 138-147

10-13
14-16

17-19
20-,21

Number 139 (published September 30,1985)
Transactions

799. Matsui, Seiichiro: Recurrent molluscan associations of the OmmaManganji fauna in the Gojome-Oga area, Northeast Honshu, Part 1.
General discussion of fauna and systematic notes on gastropod andscaphopod species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149-179 22-23

ii

800. Matsuoka, Keiji: Pliocene freshwater gastropods from the Iga Formation of the Kobiwako Group, Mie Prefecture, Central Japan ....... 180-195 24-27
801. Omura, Akio, Tsuji, Yoshihiro, Ohmura, Kazuo and Sakuramoto,Yuji: New data on hermatypic corals from the Pleistocene limestone
on Kikai, Ryukyu Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196-205
802. Kobayashi, Teiichi and Hamada, Takashi: Additional Silurian trilobitesto the Yokokura-Yama fauna from Shikoku, Japan . . . . . . . . . . . . . 206-217 28-30
803. Sato, Tadashi, Kasahara, Yoshio and Wakita, Koji: Discovery of a
Middle Jurassic ammonite Kepplerites from the Mino-Belt, Central
Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218-221
Proceedings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222-223
Number 140 (published December 30, 1985)
Transactions

804. Matsui, Seiichiro: Recurrent molluscan associations of the OmmaManganji fauna in the Gojome·Oga area, Northeast Honshu. Part 2.
Systematic notes on bivalve species . . . . . . . . . . . . . . . . . . . . . . . 225-239 31-34
805. Matsuoka, Kazumi: Cyst and thecate forms of Pyrophacus steinii(Schiller) Wall et Dale, 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240-262 35-40
806. Matsumoto, Tatuso and Noda, Masayuki: A note on an inoceramid
species (Bivalvia) from the Lower Coniacian (Cretaceous) of Hokkaido . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263-273 41-44
Number 141 (published April 30, 1986)
Transactions

807. Ohtsuka, Yasuo: Early internal shell microstructure of some Mesozoic
Ammonoidea: Implication for higher taxonomy . . . . . . . . . . . . . . . 275-288 45-50
808. Koike, Hiroko: Microstructure of growth increments in the shell of
Mercenaria mercenaria (Linne) . . . . . . . . . . . . . . . . . . . . . . . . . . 289-295 51-52
809. Takahashi, Shizuo, Domning, Daryl, P. and Saito, Tsunemasa: A newancester of Steller's sea cow from the Upper Miocene of Yamagata

Prefecture, Northeastern Japan . . . . . . . . . . . . . . . . . . . . . . . . . . 296-321 53--62
810. Ishiga, Hiroaki and Miyamoto, Takami: Follicucullus (Radiolaria)
from the Upper Permian Kuma Formation, Kyushu, Southwest
Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322-336 63--64
Proceedings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337-340

iii

Number 142 (published June 30, 1986)
Transactions
811. Horiuchi, Junji and Kimura, Tatsuaki: Ginkgo tzagajanica Samylina
from the Palaeogene Noda Group, Northeast Japan, with specialreference to its external morphology and cuticular features ....... 341-353 65- 68
812. Noda, Masayuki: A new species of Inoceramus (Cordiceramus)(Bivalvia) from the Upper Coniacian (Cretaceous) of Hokkaido ..... 354-365 69-73
813. Tashiro, Masayuki and Matsuda, Tomoko: Lower Cretaceous bivalvesfrom the Sakawa area, Shikoku . . . . . . . . . . . . . . . . . . . . . . . . . . 366-392
814. Tazawa, Jun-ichi and Kato, Makoto: Striatifera and Gigantoproductus

74-77

from the Lower Carboniferous of Fukuji, Central Japan . . . . . . . . . . 393- 399

78815. Ehiro, Masayuki, Shimoyama, Shoichi and Murata, Masafumi: Some
Permian Cyclolobaceae from the Southern Kitakami Massif, Northeast
Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400-408 79- 80
Number 143 (published September 30,1986)
Transactions
816. Matsumoto, Tatsuro and Noda, Masayuki: Some inoceramids (Bivalvia)
from the Cenomanian (Cretaceous) of Japan -1. New or little known
four species from Hokkaido and Kyushu . . . . . . . . . . . . . . . . . . . . 409-421
817. Goto, Hiroya, Maruoka, Kunitaka and Ishii, Ken-ichi: Lepidolina

81- 85columbiana (Permian fusulinid) from British Columbia, Canada ..... 422-434

86818. Nishida, Harufumi: On a new Tempskya stem from Japan . . . . . . . . 435-446

87-89

819. Kobayashi, Teiichi and Hamada, Takashi: The second addition to the
Silurian trilobite fauna of Yokokura-yama, Shikoku, Japan ....... 447- 462 90-92
820. Matsumoto, Tatsuro, Muramoto, Kikuwo, Takahashi, Takemi, Yamashita, Minoru and Kawashita, Yoshitaro: On Neocrioceras spinigerum(Jimbo), a species of Cretaceous heteromorph ammonoids (Studiesof the Cretaceous ammonites from Hokkaido-LV) . . . . . . . . . . . . . 463-478 93- 94
Number 144 (published December 30,1986)
Transactions
479-489821. Kozai, Takeshi: Early Cretaceous Pulsidis (Bivalvia) from Japan822. Fuji, Norio: Palynological study of 200-meter core samples from
Lake Biwa, Central Japan II: The palaeovegetational and palaeoclimaticchanges during the CA. 25,000 - 100,000 years B.P. . . . . . . . . . . . . 490-515
823. Igo, Hisaharu and Hisada, Ken-ichiro: Lower Permian conodontsfrom the Kawanori Formation of the Southwest Kanto Mountains,

95

iv

Tokyo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 516-527 96-98
824. Kimura, Tatsuaki, Naito, Gentaro and Ohana, Tamiko: Early Jurassic
plants in Japan. Part 7. Fossil plants from the Nishinakayama Formation, Toyora GrouPl Yamaguchi Prefecture, Southwest, Japan ..... 528-541 99-100
Systematic Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . 542-544
Index of Genera and Species . . . . . . . . . . . . . . . . . . . . . 545-550

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Transactions and Proceedings of the PalaeontologicalSociety of Japan
December 30, 1986

New Senes No. 144

CONTENTS
TRANSACTIONS
821. KOZAI, Takeshi: Early Cretaceous Pulsidis (Bivalvia) from Japan . . . . . . . . . . . . . . 479
822. FUJI, Norio: Palynological study of 200·meter core samples from Lake Biwa, Central
Japan II: The palaeovegetational and palaeoclimatic changes during the CA.
250,000 - 100,000 years B.P. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 490
823. IGo, Hisaharu and HISADA, Ken·ichiro: Lower Permian conodonts from the Kawa·
nori Formation of the Southwest Kanto Mountains, Tokyo . . . . . . . . . . . . . 516
824. KIMURA, Tatsuaki, NAITO, Gentaro and OHANA, Tamiko: Early Jurassic plants inJapan. Part 7. Fossil plants from the Nishinakayama Formation, Toyora Group,
Yamaguchi Prefecture, Southwest Japan . . . . . . . . . . . . . . . . . . . . . . . . . . 528
SYSTEMATIC INDEX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 542
INDEX OF GENERA AND SPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 545