Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No. 92•Palaeontological Society of Japan
December 20, 1973

Editor: Hiroshi U JIlE
Associate editors : Iku w o OBA TA, Yasuji SAITO, Kazuhiko UEMURA

Officers for 1973 - 1974
Honorary Presi den t:
President:

Teiichi KOBAYASH!

Tatsuro MATSUMOTO

Councillors (*Executive) : K a zuo ASAMA,* Ki yoshi ASANo, K iyotaka Cr-II NZEI,* Takashi HAMADA,* Tetsuro HANAI,* Kot ora HATAI, Itaru HAY AMI, Kametoshi
KANMERA,* Tamio KOTAKA, T atsuro MATSUMOTO,* Tok io SI-IIKAMA, Tsugio
SHUTO, Yokichi TAKAYANAGI,* Toshimasa T ANAI, Hiroshi UJIlE*
Executiv e Com mittee:
Genera l Affairs:
Membership:
Finance:

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Kazuo ASAMA, Ienori FUjIY AMA

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Publica tions
Transactions:

Hiroshi UJIIE, Ikuwo OBATA, Yasuji SAITO, Kazuhiko UEMURA

Special Pap ers : Kametoshi KANMERA, Itaru HAY AMI, Tomowo OZAWA" Fossils":

Yokichi T AKAY ANAGI, Toshiaki TAKAYAMA

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P A LAEONTOLOGICAL SOCIETY OF J APAN
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Trans. Proc. Palaeont. Soc. Japan, N. S., No. 92, pp. 163-184, pIs. 27-28, December 20, 1973

624.

EVOLUTION AND MODE OF LIFE OF INOCERAMUS(SPHENOCERAlYIUS) NAUMANNI YOKOYAMA EMEND.,AN UPPER CRETACEOUS BIV ALVE*
KAZUSHIGE TANABE
Department of Geology, Kyushu University, Fukuoka 812

ailli*cl:iNill=tl(j;H Inoceramus (Sphenoceramus) naumanni YOKOYA:-'IA emend. 0)
1ff;1t C. ~iM;~A: Tf.it1itkl*! iJ#l!tl!lJ,J< ':lbb};Q rjll!ilh:ltl'fII;]!;(fiitl!J-Ql( iJ J:. lJ 1:j1 "'lhliif,llimiilJitl!J-gl(r~ 5J;{jiT Q J:l~rlailli*"t /' r .:.:. 7 /'~ 7J /' / ~.:.:. 7 /'il' G;j!H;),,:i'!"JI~l*:!oiU~ ;fct-: Inoceramus (Sphenoceramus) naumanni O)~E'fItl\'l;;lq~ -::J\' 'L, IINI*llfr::.£--5 2: ifli:ik • }jX:ft~~~HUi'I'#~':H::'~%-~
LJ:, {-0)t,'i!!R"t /' r .:.:. 7 /' O)f~,'{*I::' -::J\' 'Lv;t, m\~O)~~ iJ:'il9:;:¥.jr::.M L L'jl'f L v,g;nx:ftf~
Ji:, Q :::. C. ~J, %fi:ti\tJ': ~~ Cn~)~t!M' G, ~lmlm:a: ;mc -C 5E*TmE*~Jiwll¥H~f'J7g Lt-:~WJ!!li'iO):tJ:iMJR:rI::a:i'fL Lv't-: c~;tG;fcQ, U1f~;HL -C 7J /'/~.:.:. 7 /' O)t~*f~-::JV'Lf;t, lfi\lIHa~O) ;ii€~:I-c"{-:tV~ -e 0) I~],L.' P1JJiJ f~lJQ;t L Tjll[>,(J!ijO)j€:£iJ;i,,21ib G;fc, L iJ' t 1R)'PdI c. c. t f~ ik~;~.<mb;fcQJ:.5~~Q,

5J~~o)~~KW~L'il9:~KMTQ'il9:O)~G~'~IO)W~~,':UL'i -c"O)%nx:ftiJ' Gf~nx:ftf::.~b Q -.:. c. -f:', 7·!.1 A'.b. O)"t 1 A iJ;:e.l~:;;k: 2: < tJ': Q :::. c. iJ, G,:::. 0) J:. 5 tJ':ILt:l5i::v;t'il9:O)ID::ru:O) :e.i~tJ': 11:~1Jn \!:' '312: jl1l:::. -9 c. ~;t G;fc Q, L t-:iJ; ~ L *f!lH~iJ It
Mlp!S(JlljO)~1:i;t,

G f~ c. 0)~1ti;J:,~i-¥;~i1;7'GJJ'EE.A..0) ~i~:t*AO)~ftf~I*? i@r.tJf;®,c.%-;t G;fcQ,*f!tO)ffi\lI*j€1~0)~tJ': Gf **xj€:£:~iJv'L t

Lt-:,

~

iPllib Gil, Q :::. c. \!:' a)J GiJ'l::'f!lIJiil1 - fiX.

1969), MATSUMOTO and NODA (1968),
HAY AMI (1969) and STANLEY (1972). In
these studies, HAUFF (1921), and JEFFERIES and MINTON (1965), pointed outthat the mode of life of the Lower
Jurassic inoceramid, Pseudomytiloides
dubius (SOWER BY) was pseudoplanktonic
because some individuals of this species
attached themselves to the surface ofdrift wood. KAUFFMAN (1965, 1967) discussed the mode of life of the Late
Cretaceous inoceramids in the westerninterior of the United States. His discussion was mainly based on the modeof fossil occurrence and the relation
between shell morphology and lithofacies.
HA YAMI (1969) contributed awork on the mode of life of certain
Mesozoic Bivalvia. According to him,

Introduction

Among various Mesozoic Bivalvia. theInoceramidae has been utilized stratigraphically as important index fossils.
because of their diverse shell morphology and wide geographic distributions, but the biological significancessuch as the mode of life and the relation between morphology and function
have not yet been known to our satisfaction.
Studies of the mode of life of the
Inoceramidae have been undertaken by
many paleontologists, such as HAUFF(1921), SEITZ (1962), JEFFERIES andMINTON (1965), KAUFFMAN (1965, 1967,
Received May 9, 1973: Read Oct. 28,
1972 at Matsuyarna.

163

Kazushige TANABE

164

such Mesozoic Bivalvia as Bositra,
Daonella, Monotis, Buchia and Inoceramus
were nektoplanktonic or pseudoplanktonic. STANLEY (1972) mentioned thatsome Cretaceous inoceramids, such as
the subgenus Inoceramus (Mytiloides),exibit definite endobyssate features.
However, most of these studies on themode of life of the Mesozoic Bivalvia
were founded on the mode of fossil
occurrence. Little has been done on
the analysis of relative growth and on
the transportational or buoyancy mechanism of the shells.
In the present paper, I have dealt
with Inoceramus (Sphenoceramus) naumanni YOKOYAMA emend, which almost
corresponds to the group of Inoceramus
naumanni of NAGAO and MATSUMOTO(1940). This species occurs abundantlyin the Coniacian to the Campanian of
the Upper Cretaceous in the JapaneseIslands and Sakhalin, showing a wide
geographic distribution, especially in thenorth Pacific province. The present
species is very interesting for its successive occurrences showing low articulation ratios and a high degree of clustering. The forms from the Santonian
and the Campanian are quite similar toeach other except for the occurrence ofdivergent ribs in the latter.
In this paper, I study the mode offossil occurrence and examine biometrically the shell characters, discussing
the evolution and mode of life of thepresent species. I give special consideration to the numerous specimens whichwere successively collected from theUpper Cretaceous of Hokkaido and south
Sakhalin.
Acknowledgements

express my sincere gratitude to Professor Tatsuro MATSUMOTO of the Kyu-

shu University and Professor R. A.
REYMENT of the University of Uppsalafor their valuable suggestions and critical reading of the manuscript, to
Associate Professors Kametoshi KANMERA and Tsugio SHUTO of the Kyushu
University for their kind advice, to Dr.
!taru HAY AMI for his valuable information and criticism, to Messrs. Tomowo
OZA w A and Hiromichi HIRANO for theiruseful discussions, and to Mr. Masafumi
ARITA for his valuable suggestions on
the transportational mechanism of shells.
Thanks are extended to to Dr. Yasuhide IWASAKI of the University Museumof the University of Tokyo and Mr.
Yasumitsu KANIE of the Yokosuka City
Museum for allowing me to study the
specimens preserved at their respective
museums.
Material
Outline of stratigraphy.-In the central
axial belt of Hokkaido extending to
Sakhalin, marine Cretaceous deposits
are widely distributed. They are divided litho-stratigraphically into fivegroups as follows: the Sorachi group(Jurassic to Neocomian), the Lower Yezo
group (Upper Aptian to Middle Albian),
the Middle Yezo group (Upper Albian
to Turonian), the Upper Yezo group(Turonian to Lower Campanian), theHakobuchi group (Lower Campanian to
Maestrichtian), in ascending order (MATSUMOTO, 1954).
Numerous calcareous nodules are contained in the middle to the upper partof the deposits, and sometimes well-preserved marine fossils such as ammonoids,
bivalves, especially inocerami, gastropods, echinoids, and others, are foundin them. The specimens of Inoceramus(Sphenoceramus) naumanni YOKOYAMA
emend., treated in this paper, were col-

...

624.

Inoceramus (Sphenoceramus) naumanni

lected from the Naibuchi district ofsouth Sakhalin, the Tappu district ofnorthern central Hokkaido, and the
Urakawa district of southern central
Hokkaido (see Text-fig. 1).

165

the Upper Cretaceous in Japan.
Other species of Inoceramus that may
be associated with the present species
in the same nodules, are I. (Platyceramus) amakusensis NAGAO and MATSUMOTO and I. (Sphenoceramus?) pseudosulcatus NAGAO and MATSUMOTO. They

Th~ibuchi

Tappu

The Miho

district

y 20·-35olkm

7' 35'-50·

Text-fig. 2. Map showing the fossil
localities in the Naibuchi district (adaptedfrom MATSUMOTO, 1942).
fhe Mlllo,lh. Second

T"b,,'.'" of tho No.hue .. i, N•. buch, di,hid

Urakawa district·Text-fig. 1. Index map showing the Cre·taceous outcrops and the areas studied.

Sampling method.-Random sampling·was made from a single fossil-bearing·calcareous nodule in most cases, andfrom a fossil-bearing bed in the case of-the samples N19a, N18g and N17b from·the Naibuchi district and the sample
U120 from the Urakawa district.
As there are no data on the absolute:.age of the Upper Cretaceous in the.areas studied, the stratigraphic positionof each sample is expressed by the-thickness (m) in a sequence measured
from the first occurrence of the present
species, as well as by MATSUMOTO'S.(1959) biostratigraphic subdivision of· ..
·· ..; ; '!,OI;ia1\"--=1==-,.-."-,,---.:--..~X
H".ighlin ......

to.- '''''~:~::::.'.:' ~:..
C••••

Text-fig. 3. Stratigraphic position andsize-frequency distribution in samples fromthe Naibuchi district.

166

Kazushige TANABE

are easily distinguishable from I. (S.)
naumanni by the analyses of the individual' relative growth and the apicalangle.
Samples from the Naibuchi district.-7
samples are used in the present study.
They were collected from the Upper
Santonian to the Campanian in the R.
Miho. the second tributary of theR.
Naibuchi. The fossil localities and theirstratigraphic positions are shown in
Text-figs. 2-3. The stratigraphy of theUpper Cretaceous in the district is after
MATSUMOTO (1942).
Samples from the Tappu district.-19
samples are used in the present study.
They were collected from the iIpper
part of the Upper Yezo group (Santoni an) in the R. J ugosen (R 2665a. R
2667b, R 2667c, R 2672a, R 2672f, R 2673a.
R 2673d, R 2680a, R 2680b, R 2680c. R
2680e, R 2681a, R 2686a', R 2686d, R 26~7b,
R 2694a and R 2696a) and the R. Akanosawa (R 2402a-x and R 2403), the t;ibutaries of the Obirashibe Valley. The
fossil localities and their stratigraphic
positions are shown in Text-figs. 4-5.
The stratigraphic division of the Upper
Yezo group in the district is after T ANAKA (1963). According to him, the ageof the group is from Turonian to Lower
Campanian, but in my opinion the upper
limit of the group is probably Upper
Santonian. because of the occurrence of
Inoceramus (Cladoceramus) undLilatoplicatus japonicus NAGAO and MATSUMOTO
in the uppermost horizon.
Samples from the Uralwwa district.12 samples are used in the present study.
They were collected from the upper partIof the Upper Yezo group (Upper Santonian to Upper Campanian) in the uppercourse of the R. Chinomi (U 142, U 143r,
U 147-2, U 120, U 128c and U 115), thelower course of the R. Chinomi (U 4464), the upper course of the R. Urokobe-. )':d':'--- -----

~

[
j

~

: _

Silhtone
Cloy, tone

Bluff

Text-fig. 4. Map showing the fossil
localities in the Tappu district.
----~--------------.

; K. TANABE{1972 )

The

Jugosen, Toppu

district'UI_,",[::~~--,,----.J.. o 0!

HeightText-fig. 5. Stratigraphic position andsize-frequency distribution in samples fromthe Tappu district.

tsu CU 47a, U 47b, U 46, U 42-2 and U42-1) and the coast of Ikandai CU 508).
The fossil localities and their stratigra-

624.

Inoceram1ts (Sphenoceram1ts) na1tmanni

167

N

t
........... Tu f f

Ld Sandstone

The Urokobetsu

13 S;I",one_Text-fiig. 6.

Y. KANIE(19661u

Z

The

The Chinomi~ ConglomerateUSB'--_ _ _....:.500 m

Claystone

Map showing the fossil localities in the Uraka wa district.

CI,;nom; Qnd ,h U,,,kob,,,.u,U,,,lo.awCl c;"';cl

~

:, ]
Z

I'

Text-fig. 7. Stratigraphic position andsize-frequency distribution in samples fromthe Uraka wa district.

phic positions are shown in Text-figs.
6-7. The stratigraphic divisions of thisgroup in the district are based on KANIE(1966).

Mode of fossil occurrence

Mode of fossil occurrence.-The modeof fossil occurrence of the present
species is very similar to that of theLower Jurassic nektoplanktonic or pseudoplanktonic bivalve, Bositra buchi(ROMER) (JEFFERIES and MINTON, 1965;
STANLEY,1972). In the above-mentioned
three districts, the shells occur crowded
in the calcareous nodules throughout thesuccessive sequence. This fact indicatesthat the material is suitable for the
study from the viewpoint of the population concept (NEWELL, 1956). The fossil
occurrence is generally independent of
the lithofacies. The articulation ratio
and the chi-square test of the valves ineach sample from the Naibuchi, theTappu and the Urakawa districts are
summarized in Table 1. The articulation ratio is especially low in the specimenS from the Santonian and it is also
low in the concentric ring stage fromthe Campanian.
In contrast to this,most specimens of certain associatedbivalves, e. g. Nanonavis sachalinensis(SCHMIDT), Nuculana mactraeformis NAGAO, Propeamussium cowperi yubarense

168

Kazushige TANABE(Y ABE and NAGAO) and Inoceramus (Platyceramus) amakusensis NAGAO and MATSUMOTO, are complete and conjoined as
seen in the samples R 2667c, R 2680a-c
and R 2686d. As BOUCOT (1953) pointed
out, the articulation ratio is controlled
by such physical conditions as the velocity of the current on dead shells, thedepositional· process and the strengthof the ligament. The ligament of thepresent species is considered to be weak,
judging from the poorly developed ligamental pits and very thin shell. The
common occurrence of an associate
thin-shelled bivalve, Propeamussium cowperi yubarense with complete conjoined
valves, in spite of weak ligamental intensity, makes it difficult to explain the
low articulation ratio of the present
species only by the weakness of ligamental intensity. A similar clusteredoccurrence, with a low articulation ratio,
has been observed also in the Santonian
sequences along the R. Sakasagawa in
the Kamihaboro district, along the R.
Kotanbetsu III the Kotanbetsu district,nothern central Hokkaido, and along the
R. Shutta-zawa, tributary of the R.
Hobetsu, southern central Hokkaido(see Plate 27). In most cases, the resultof the chi-square test indicates that thedifference between the frequencies of
the two valves is not statistically significant at the 95% level, and, therefore,
the question of the selective transportation and deposition of the valves remains
unanswered.
A definite orientation of the shells is
not usually observed, but in the case of
R 2696a, the fossil-bearing layers are
sharply separated from the barren layers
III a nodule.
This is somewhat similar
to the mode of fossil occurrence of thevery thin shells in some pelagic sediments, as exemplified by the Ohse formation of the Sambosan belt in southern

Table 1. Articulation ratio and chisquare test of the valves in samples fromthe Naibuchi, the Tappu and the Urakawa
districts (Articulation ratio shows thevalue of the concentric ring stage and thatof the divergent rib stage, respectively).
R.V.

sampleL.V.

articulation
ratio

chf-sqare'N

21a

16 (0)

7 ( 0)

9 ( 0)a (----)

0.25 •;.:

19a

11 (3)

7 (2)

4 ( 1)() (0)

0.82 •

!-i

l8p:

l~

(12)

8 (»

10 (7)

0.29 (

n)

O.2~.

laf

69 (n)

26 (9)

4] (18)o (0)

4.19 0lSb

42 (2)

25 OS)

17 ( 8)a (o.n")

1.52.

16h

44 (18)27 (9)

17 ( 9)n (OJ.'.-'i' •

17h

J (3)

1 (1)

'2 (

(l

<J)

'), B •

R 2665.1

73 (0)

49 (0)

21. ( 0)0 (----)~.5h0

R 2667h72 (ll)],", (0)

]-'<

( 0)

!l (----1

().~~

•

R 2667\'87 (0)

46 (0)

41 ( 0)0 (----)

0.H9 •

R 2672.1

43 (0)

2n (0)

23 ( 0)0 (----)

0.21 •

R 2672.f

232 ( 0) 119 (0)ll] ( 0)o (----)

0.02 eR 2673<1

40 (0)

23 (0)

17 ( 0)0 (----)

0.90 eR 2673d39 (0)

2] (0)In ( u)0 (----)

1.26e

R 2680<1

138 (0)

J:l5 (0)5:1 (u)0.01 (----)

7,1120R 2680b7tl (oJ

39 (0)

31 (0)

0.01 (----)

0.9] 0R 2680c

122 (0)70 (0)

52 (0)

0.02 (----)

2.70e

R 2680c172 (0)90 ( 0)82 (0)

0.01 (----)

0.17 eR 2681a]57 (0)82 ( 0)75 ( 0)0 (----)

0.31 eR 2686a55 (0)

32 ( 0)23 (0)(l.u4 (----)

1.47.

R 2686d

29 (0)

16 ( 0)13 ( 0)0 (----)

O.]le

0

N

:-;

n

(

R 2687b

7 (0)

4 (0)

J ( 0)(----)

0.14 eR 2694a]4 (0)

21 ( 0)13 (0)

0.06 (----)

1.88 eR 2696a85 (0)

17 ( 0)48 (0)

0.07*(----)

1.42.

U

142

23 (0)

13 ( 0)10 ( 0)0 (----)

0.44.

U

141r

22 (0)

12 ( 0)10 ( 0)0 (----)

0.18.

u 147-::

122 (7)64 ( 5)51 (2)

0.04 (----)

U120

11 (8)

2 ( 1)

9 ( 7)

U508

21 (8)

11 ( 3)10 ( 5)0 (U 446-4

Js (18)17 ( 9)Itl ( 9)0 (0.50)

0.01 eU47b

126 (56)

6\1 (28)66 (18)n (0.0])

0.29.

U

128c

51 (22)I·. (I:?)

15 (10)n (0.09)11

47a

42 (29)2'-) (16)17 (UI)tl (46

52 (21)15 (9)

I', (I2)

11

42-2

40 (30)23 (17)

II (13)o (I)(l.90 eU42-1

21 (2n)9 (B)12 (12)o (II)

0.43 e(l(0.22)
0)

o.os.

n)

1.52.

( ) ... Divergent rib stage
Valve open position• -.- Not significant at 95% confidence le,-el
o _.. Significant at 95% confidence level

* ...

624.

Inoceramus (Sphenoceramus) naumanni

Kyushu (KANMERA, 1969, Plate 6) andthe Upper Triassic to Jurassic "Posidonia" limestone of the West Carpathians (MISIK, 1966, Plates 33 and 46).
Faunal combination.- The Upper Cretaceous sediment~ in Hokkaido and
Sakhalin are generally poor in bivalves
other than inocerami. The present species holds an important position among
the megafossils contained in nodules,
except for ammonoids. This role has
been ascertained in most nodules collected from the Senonian sequences ofthe districts studied. For example, thepercentages of the number of individualsof this species among the molluscs ineach sample, and also that of the bivalves taken separately from the R.
Jugosen, Tappu district are as follows.
R 2665a: 80% (100%) R 2680b: 89% (100%)
R 2667b: 85% (100%) R 2680c: 85% (94%)
R 2667c: 93% (98%) R 2680e: 97% (100%)
R 2672a: 75% (lOO%) R 2681a: 84% (97%)
R 2672f : 89% (98%) R 2686a': 88% (100%)
R 2673a: 47% (87%) R 2686d: 91% (100%)
R 2673d: 90% (90%) R 2694a : 100% (lOO%)
R 2680a: 94% (97%) R 2696a : 100% (100%)

No other marine megafossils were foundin samples R 2694a and R 2696a. The
percentage is generally high in claystone. Such a dominant occurrence ofthe present species in the Upper Cretaceous fauna can be observed also in theother two districts. This indicates thatthe present species was remarkably
prolific.
Transportational mechanism of shells.The main cause of a reduction in thearticulation ratio is regarded as due totraction and saltation.
VISHER (1969)
indicated that the transportational mechanism consists of a combination ofsuspension, saltation and traction, basedon the grain-size analysis of many recent
and ancient clastic sediments. This169

mechanism of transportation is well
exhibited by the three populations when
the data are plotted on the log-probablity paper. I also tried the grain-size
analysis of some nodules, after having
collected fossils in the following manner.
Crushed nodules were digested in hydrochloric acid in order to dissolve thecarbonate~ and then were neutralizedwith sodium hydroxide. The grains of
the residue were separated by ultrasonic. The results are shown in Textfig. 8. It is considered that the transportational mechanism of the sedimentsof R 2696a (claystone) was suspension
and that of R 2667b (sandy siltstone)
was a combination of traction, saltationand suspension. Although the transportational mechanism may not be thesame between grains and shells, mostshells in the examined samples were
probably transported by currents andeventually fell to the sea bottom.
99.99 r-----r--.------.--,-------.--,------.----,

99

90

c

R2667b~

QJ

u

'QJ
Q.

50

L
O"l
QJ

310

\
R2696a0.1
0.01 "'---'--_.l....---'-_--'---'_~_"'--__'

o

3

4

Q25
0.125
Scale

0.067

2
Phi

05

Scale

mm

Text-fig. 8. Grain-size distributions ofsome nodules from the Tappu district.

Kazushige TANABE

170
Biometrical analysis

Abbreviations.-I use the following
abbreviations in the present paper.
ri b

For univariate analysis
R. V.: Right valve; L. V.: Left valve;
N: Number of individuals in a sample;
O. R.: Observed range; X: Arithmetic
mean; s: Standard deviation; ax: Stan·
dard error of the mean; P: Probability
level; ,,: Degrees of freedom.
For bivariate analysis
R. M. A. : Reduced major axis; a: Growthindex; (3: Initial growth index: r: Correlation coefficient; act: Standard errorof a.
For Repositories
GK. : The prefix means that the specimens
is kept in the type collection of the Kyushu University. The specimens of thesamples U 142, U 143r, U 147-2, U 47b,
U 128c and U 46 were collected by MATSUMOTO in 1940, and U 120, U 508, U 47a,
U ll5, U 42-2 and U 42-1 were collectedby me in 1972.
The specimens of R
number samples (R 2665a-R 2696a, R
2402a-x and R 2403) were collected by
HIRAXO and me in 1971.
MM.: The prefix means that the specimens were collected by MATSU:VIOTO fromthe Naibuchi district in 1937, and arestored in the type collection of the University Museum of the University ofTokyo.
YCM. : The prefix means that the specimen
is stored in the type collection of the
Yokosuka City Museum. The specimensof the sample U 446-4 were collected by
KA:-iIE in 1969.

Basic morphology and measurementsof the present species are diagrammatized in Text-fig. 9. A special comparator designed by SHUTO and a wideview microscope were used for themeasuremen ts.
Size-frequency distributions.- The sizefrequency distributions of the shellheight in the samples from the Naibu-~length

I

TI'

~,.

'

.··T2

DePt~.Tn(
HnTh;.ckne,., ofprismaticloyer)

Text-fig. 9. Basic morphology andmeasurements of the subgenus Inoceramus(Sphenoceramus) .

chi, the Tappu and the U rakawa districts are shown in Text-figs. 3, 5, and7, respectively.
According to BouCOT (1953) and CRAIG(1967), the size-frequency distributions of
the molluscan death assemblages are
originally controlled by interaction of
the mortality and the growth rates ineach living population. The survivorship curve of many marine invertebrates
shows the presence of a large numberof larval and immature individuals, amoderate number of adults and a small
number of gerontic individuals. In thedeath assemblages, the larval individuals
have been damaged mostly by physicalagents and are not preserved. Therefore, the size-frequency distribution indicates strong positive skewness withlarval stages lacking (HALLAM, 1967).
The size-frequency distributions of themolluscan death assemblages are also
controlled by chemical and biologicalconditions, and accordingly, the absolute
growth of each living population andthe representation of the physicochemical conditions are needed for a reasonable interpretation of the size-frequency distributic·ns.
The size-frequency distributions of
the samples from the Santonian in the
Tappu district (Text-fig. 5) indicate that

624.

Inoceramus (Sphenoceram'us) naumanni

the pattern changed with time frommoderate positive skewness (r i g h t
skewed) to a bell-shaped normal distribution curve. Generally speaking, therange of each size-frequency distribution increases as the sequence goes
upward.
In the samples from the
Naibuchi and the Urakawa districts, therange of each size-frequency distribution (Text-figs. 3 and 7) also increases
with time during the Santonian. In thesamples from the Campanian, the largest
limit of the concentric ring stage is lessthan 50 mm in shell height. In comparison with the concentric ring stage,
the range of the divergent rib stage islocated on the right side of each sizefrequency distribution. In the example
U 12Sc, the pattern appears to be bimodal. The largest specimen in the divergent rib stage sometimes exceeds 30
cm in shell height, as observed for the
Urakawa district.
Univariate analysis.-Method of univariate analysis of invertebrate fossils
has been used by SIMPSON and ROE(1939), IMBRIE (1956), SYLVESTER-BRADLEY (195S), SIMPSON, ROE & LEWONTIN(1960), and many others.
It is wellknown that a univariate analysis is
insufficient to charge a growth relationship. In the case of the present species,
the shell form represented by such characters as the simple ratio, D/H and T /H
is changeable with growth. T is the
thickness of the prismatic layer, H theshell height, and D the shell depth (=inflation). Consequently, only the shellheight in the concentric ring stage has
been examined for the univariate analysis.
In the later forms of the present
species in the Campanian, two stagesof growth are distinguished, namely theconcentric ring stage and the divergent
rib stage (see Text-fig. 9). The surface

171

ornament changes from the former tothe latter during ontogeny.
In thespecimens from the Lowest Campanian,
the divergent ribs can be recognized in
the later stage (e. g. Plate 2S, fig. 12),but in the specimens from the Middle
to the Upper Campanian, they can beclearly recognized from the middle oreven early stage (e. g. Plate 2S, figs.
13-1S). The range of variation of the
distan::e from beak to the point of thefirst appearance of the divergent ribs
seems to be nearly constant in eachsample.
Table 2. Measurements of the shellheight in the concentric ring stage insamples from the Naibuchi and the Urakawa districts.
~xobst'fved range

N

IBg

11

17.40 :!: 2.01

}7,::53.00

0.90

1 ~ .iU - 2:'.00

N

1Sf

21

14.'11.

:!: 1.34

19.58

2.93

0.64

I O.f'~

" N

J8b

24

12.25 :!: 1.1)1

19.52

~. 39

0 ... 91.(,(116

11.13 :!: 1. .. 5

2':'. 5~

2. i3

b.57 :!" 0.77':'.90

O. )2

O.ld

b.20

:!: 1.31:,.1'>0

1.;>4

0.51

25.'101.3 ..

14.80 - 25.80

i :!: t o . 05 CT ;sample

N

~

lib

3

U )!.7-:',~;~7 .nS

5.911- ::'1.60- 15.i5- 1... 50- 6.8029,()()

120

"
'08
,
"

,

21.25 :!: 3.28

16.75

3.5&

B

17.i:': 2.71

18.69

3.32

1.17

1 J.35~"6-:'

15l).',u:!: 1.16

15.30

Z.08

0.:; ..

10. JO- ;i .00~7b'I

12.()/' + 0.62

18.36

2.21

0.31

S.W

- 15.9()

U

128c

Ib

10.24

!u

:'7a

"17

9.;h

U

.
]

:!: 1.21

22.27

2.28

0.57

7.45 - 14.35:t 0.53
8. 7~ :t 0.72

n.O)

1. 36

0.26

'.",0 - 11.90

15.93

1.39

0.3:'

h.lO

7.59 : 0.11

1.27

n.IO

0.0!,

7.50 -

U

4,

U

11'

U

42-2

25

7.27 :t 0,62

20.58

1.50

42-1

g

&.19

:- o. ~J

1::.310.76,- 23.65- 11.207.75:'.75 - to.90
O. ~O

4.7S

-

7.75

Therefore, in order to show quantitatively the variation, X (at the 95% confidence level), s, a.t, V and O. R. of the
distance in each sample from the Naibuchi and the Urakawa districts havebeen calculated. The results are summarized in Table 2. The shell size ofthe concentric ring stage was gradually
decreased with time. The chi-square
tests in each sample are as follows.

Kazushige TANABE

172

Samples from the Naibuchi district.-

N 18g: x 2 = 1.32~3.84 = X20.05CV=1l' P~0.05, n.s.
N 18f: x 2 = 0.15~3.84 ="""N 18b: x2 = 0.26«:3.84 ="""N l6b: x2 = 1.41~3.84 ="1/"

Samples from the Urakawa district.U 147-2: x 2 = 1.39~3.84 = X20.05Cv=1l,
U 120 : X2 = 0.24~3.84 ="U 508 : X2 = 0.71~3.84 ="U 446-4: X2 = 0.59~3.84 ="U 47b : X2 = 3.04~3.84 ="U 128c : X2 = 0.29~3.84 ="U 47a : X2 = 0.43~3.84 =.
"U 45: X2 = 4.83~ 6.64

= X20.OICv=1l,

U 115 : X2 = 2.11~3.84 = X20.05CV=1l'
U 42-2 : X2 = 2.23~3.84 ="U 42-1 : X2 = 0.06~3.84 ="n. s.: non-significant

The results of the chi-square test
indicate that the null hypothesis for anormally distributed population is not
rejected for the character considered.
In order to show the difference of X
between the two samples, A (N1' Xl> 51)
and B (N2 , X 2 , 52), a F-test was made.
This is represented by the following
formula: F= 5//5 22 , . : 51 >5 2
The results of the F-test between apair of samples from the Naibuchi andthe Urakawa districts are summarizedin Table 3.
If the ratio 51 2 /5/ between the two
samples is not significant statistically,
the significance of the difference between Xl and X2 is given by the following well known formula (STUDENT'St -test).

If the ratio 5 1 2 /5 2 2 between the two
samples is significant statistically, t iscalculated by the following formulaP~0.05,

n. s.

"
"
"
"
"
"
P~O.Ol,

"
"
"
"
"
"
"
"
"
"

P~O. 05,""(WELCH'S method
variances).

forheterogeneous

The results of the t-test are shown in
Table 4. They indicate that the means
are often significantly different for apair of samples.
Bivariate analY5i5.- As IMBRIE (1956}
pointed out, an assumption of purely
isometric growth is needed for the univariate analysis. Such methods as theregression of Y on X, the regression of
X on Y, BARTLETT'S line, the major
axis and the reduced major axis havebeen applied for the recognition of average relative growth (KERMACK and
HALDANE, 1950; IMBRIE, 1956; SIMPSON,
ROE & LEWONTIN, 1960; SOKAL andROHLF, 1969). Among them, the reduced
major axis is considered a useful approximate method for the representa-

624.
Table 3.

F·test of the shell height in the concentric ring stage between a pair ofsamples from the Naibuchi and the Urakawa districts.
F-testF-testU 147 2

U

147 2

~

8.22

U

120

0

~

U

508

0

U

446-4

U

47b

U

128c

U

47a

U

46

U

115

U

42-2

t;

42-1

173

Inoceramus (Sphenoceramus) naumanni

•
•
•
•
•
•
•
•

120

U

•
•
0

•
0

0
0
0
0

19a

N

~

N

19a

N

18g

N

18f

N

18b

N

16b

0

U

•
•
•

18g

N

18f

N

18b

N

16b

N

1.05

1. 58

1. 21

90.0

~

1.50

1.15

85.5

• ~~
• •
~
1. 30

57.1
74.5

0

0

0

508

U 446-4

7.16

2.81

3.17

3. )5

1. 23

1.15

2.93

2.60

2./,0

~

2.55

2.26

2.12

U

47b

U

128c

U

47a

U

46

U

115

U

42-2
1. 46

0

0

0

0

6.67

6.56

1267

5.63

21. 8

5.80

5.71

1102

4.90

19.0

0

0

0

2.28

2.24

433

1.92

7.47

I .07

2.57

2.53

488

2.17

8.41

2.74

2.69

520

2.31

8.97

190

1.18

3.28

193

1.17

3.33

225

58.0

0

~

0

1.02

0

0

0

0

0

0

0

0

•

ratio of variances not significant at 90% confidt,tlc(' It.'vl,l•significanto

at 95%

oat 99%

tion of average relative growth (IMBRIE,
1956; WALLER, 1969; HAYAMI and MATSUKUMA, 1970). Therefore, I have been
examined the statistical discriminationof allometry by means of the reduced
major axis method. In the present species" polyphasic" (meaning that more than
one allometric relationship exist during
ontogeny) allometry can be expected forthe samples from the Campanian.
Average relative growth: In the studyof the average relative growth, Hand
D were used for the bivariate characters.
If growth is allometric, the relation be·
tween two variables, X and Y, is re-

0.38

1. 20

0

0

42 1

154

• ~~
•
•
• • • ~~
•
•
•
• ~~
~
•
• •
•
1.13

U

1. 25

0

0

0

3.88

presented by the following equation(HUXLEY, 1932).
Y = (3X" or log Y = a log X+log ,B,and according to KERMACK and HALDANE(1950), an is represented by the following formula.

ao=a~ 1f{Changes of the reduced major axis
with time for the samples from the
Naibuchi and the Urakawa districts areillustrated in Text-fig. 10. As one might
expect, a pronounced critical point ofgrowth can be recognized between the174

Kazushige TANABE

Table 4.

I-test of the shell height in the concentric ring stage between apair of samples from the Naibuchi and the Urakawa districts.

STUDENT'S

t-testt-testU 147-2

U 147-2

~
0

U

120

u

508

0

19.

N

N

19a

~

N

18g

N
N

18g

N

18f

N

18b

N

16b

N

2.24

5.46

5.63

(11.73)

0

~

3.39

4.03

(12.58)

18f

0

0

~

1. 37

(10.89)

18b

0

0

0

-..............

( 6.45)

N

16b

0

0

0

0

U

508

U 446-4(4.06)(7.31)

14.7

16.3

~

1. 98

6.40

(6.67)

•

~

3.70

6.31

0

0

U

120

47b

U

U 128c

U

47a

~
46

U

U

115

U

42-2

U

42-1

17.0

29.3

28.5

34.7

29.9

46.6

8.87

(8.82)(9.02)(10.2)(10.1)(11.1)

6.52(7.14)(7.37)(8.64 )

(8.64)(9.70)

U 446-4

0

~

2.42

4.26

8.38

7.84

(11.1)

11.1(12.9)

U

47b

0

0

0

0

~

2.82

(7.12)(7.21)(14.2)(ILl)(15.8)

u

128c

0

0

0

0

0

~

(1. 72)2.30(4.63)(6.69)

47a

0

0

0

0

0

~

0.99

(5.93)

4.78(8.98)

U

46

0

0

0

0

0

•

5.04

U

0

0

~

(3.38)

3.21(6.48)

U

115

0

0

0

0

0

0

0

0

~

(1.06)(6.73)

u

42-2

0

0

0

0

0

0

0

0

0

~

(2.98)

U

42-1

0

0

0

0

0

0

0

0

0

0

~

----- Welch r s methodo----- ratio of variances significallt at 99% confidence levelo

at 95%•

It

non-significant at 80%

concentric ring stage and the divergent
rib stage. Moreover, each critical point
is gradually decreased with time.
According to HAY AMI and MATsuKuMA (1970), the difference of isometry
and allometry in average relative growth
is represented by the following formula,with 955'6 confidence.

K=·O:-:-) .
a ~1=;;2,-1.96~ K ~ 1.96-isometry/( < -1.96-negative allometry/(> 1.96-positive allometry

I have calculated the value of K andtried to determinate the allometric relationship for each sample from the Naibuchi and the Urakawa districts. The
results are summarized in Table 5. The
growth of the concentric ring stage inmost of the samples indicates negative
allometry. On the contrary, that of thedivergent rib stage indicates positive
allometry.
Individual relative growth: As a resultof the average relative growth, it is to
be expected that the abrupt change ofthe growth index is in accord with thechange of ornament from the concentricInoceranius (Sphenoceram~(s) na'umanni

624.
U .4 2-2 R,V.

U .42-2 l.V.

N 16 b

175

R.Y.
D=O. 053H I.J79

0,,0.1 18H1.2H

O::O.106H 1. 26S.. 0.9857

O;;O.252Ho. B80

N1Bb R.V.

U 1 2 a c l.V.

U128c R.V.

D=O,074H UI7

0-0.098 H 1.324 Jl"O,?!\ 74

100.9339

O:O.404Ho 756

0=0.376 H°.l17'.0.95:' 7

r.0.9379
0,0 ~_JL.-'-.l-'..u..LU_--'_LJ.--'-LLJ.>J

U47b

U47bR,V.

0,,0.074 Hl.-493

N 1a fR.V.
0=0,066 H n90

l,V.

0

'sO.985 \I

0.0 ,,""_-L....A-L....Ll..LUCL..-_-'-----'-"'-'...u..uJo
U A.4 6-4 R ,Y.

U446-4l.V.

N 16 bL.Y.

0=0.31 I HO. SSO

0=0.32 9H o.7 76,,,0.961 B,,.0,9789'=0.9852

U147-2 l.V.

U 14 7-2 R.V. 0= D.050H

NIB b
/

l.V.
D;O.OBOH 1•303/.. 0.9656

D=O.315H0 18S
D=O.338Ho. 81J

N 18 fU \.42 l.V.

U142 R.Y.

l.V.
D:Q.06IH 13C9

."
~.,.

/

/"

D=O.351H os : 6

D=O.339 H o:'75/'.0976B

Heigh!

in

Height

in

N23a

l.V.
0= 0.385 HO. ll ]Concentric. ringstage

Divergent

slage

rib

, .. 0.9773

Text.fig. 10. Changes of the reduced major axis between the shell depth (= inflation) andthe shell height for the samples from the Naibuchi and the Urakawa districts.

J{az~Lshige TAN ABE

176

Table 5. Approximate determinationof a llom etry of the average relative growth
between the shell depth and the shell hei g ht
in each sa mples from the Na ibuchi and the
Urakawa districts.
c oncent r ic r ing stage

salllp i e

2),

N

is.

18'

~

18b

~

u

A

z

l Ob

U

- 31 . 92

L.V .

-

R.V.

-------_ .' -- -- --I. , v .

- 20 .94

R.V .

3 . 92L.V ,6 . 21

R. V.

5 . 89l. .V.

8 . 17

R. V.

- 11 . 43

L V.

-

147-2

U 446 - 4

3 . 61

R .\'.

7.t.5

L .v .

3.20

R. \' .

2 . 47l..V .

1.93

L ,V.

8 . 98R.\'.

-

7 . 28) . )6U

12Bc

I. . V .

] . 20U

1,7a R.V.

3 . JLl. V.

5 . 5540 R. V.

4 . 78

L.V .

1.80

R. V.

2 . 79i]

4 . 13L.V .

47b R.V .
u

4 . 893 . 86R.V.

l 42

4 2- 2I• . V.

42 - \

•
•
•
•
•
•
••
•
•
•
•
•
•
•
•
•
•
••
•
•

R. V.

1. 07

@

@
@

dlvcreen t----------------- -- -- - - ---- - 12.58
7.01

6 .43
6 . 397.68

8 . 695 .95--------------t. . 10

6.36
3. 79

9 . 26
9 . 65
6.81
1.68

0
@

6.93

0
0
0
0
0
0

7 . 114 . 57
10.11)--- -- -- -- ---- - -S . l)

1.96 - -- - Pos i tI ve Il il otlct r y@

1. 96 <,, < 1. 96 - ------ I SOl:lct r y .

@

1. 69

L.V.

K< - 1.96 -- -- Negat ive all omet ry

0
0
0
0
0

2 . 685 . 95K >0-----------------_. -----------

•

0
0
0
0
0
0
0

------ ---------

R.V.

O

, 'b stage---------------

244, fig. 1), the outer layer of bivalve
shell substance increases in thickness
with growth . The increase is observedto the point of the disappearance of theinner layer (KADO, 1953, text-fig . 2).
The shell structure of the Inocermidae
is composed of an outer prismatic
layer and an inn er nacreous layer. It
corresponds to the prismatic structureof KOBAYASH I' S (1971) genera li zation.
The outer prismatic layer of thepresent species consists of numerous
columnar calcite prisms. Each prismis bounded by organic matrix. The
longitudinal axis of each prism isarranged nearly perpendicular to theshell surface (see Text-fig. 11). Accordingly, the transverse section along thedirection of the shell growth is suitable
for the measurement of the thickness----1 cm-l cm - -GK. H. l0007

GK . H.l0011

R26721

U46. .'~~=a.'.'

Ol
Ccmf i ticl1ce l e ve l 9St

rings to the divergent ribs.
The shell is ver y thin (a lmost lessthan 100 fl ) and does not become thicker
with growth in the specimens from theSa ntonian (e. g. Plate 28, figs. 1-4).
However , in the specimens from theCampanian, the valu e of the growthindex suddenly incr eases from the concentric ring stage to the divergent ribstage. Ac cording . to WILBUR (1964, p.

Il

Te x t·fig. 11. Shell microstructure ofIno ceramus (Sphenoceram u s) naumanni YoKOYAM A
emend.
OL : Outer prismatic
layer.
IL: Inner nacreous la yer.
OM :Orga nic m at ri x. P: Prism.

624.

Inoceramus (Sphenoceramus) naumanniof the prismatic layer.
The mean value of the growth index
for a population is confined to the following range with 95% confidence byan approximate method (HAY AMI and
MATSUKUMA, 1970), where if and Sa are
the mean and standard deviation of afor a sample.

- 1
a>

to 05 S" < - < 1+ to 05 Sa.
I -~N-= a =-~rN---Isometry

to.05
V' NSa. .
.j7V=- -posItIvea 11 ometry

to 05 S"

Using this approach, I have tried to
distinguish the allometric relationships
between T and H for each sample. The
results are summarized in Table 6. The
growth index of the concentric ringstage in most samples indicates negative
allometry. On the contrary, that of thedivergent rib stage indicates positive
allometry in all the samples. A pronounced critical point of growth between
the two stages is observed from the
ontogenetic allometry of the selected
specimens (see Text-fig. 12). This evidence indicates that an intimate relationship exists between the appearanceof the divergent ribs and the abruptchange in mode of thickening of theprismatic layer.

If the range includes 1, the hypothesisof isometry is not rejected. In otherwords, the ranges of isometry and allometry are represented as follows.

a-<l-

L

T

177

-nega t'Ive a 11 ome t ry

Table 6. Approximate determination of allometry of the individual relative
growth between the thickness of the prismatic layer and the shell height in eachsample from the Naibuchi, the Tappu and the Urakawa districts.
concentric

sempleNex t to.os !it>

R2665a3

R2667cr in 9d ixergent

5 tog e5 tog erib

SaallometryN

0. + tQ05 Sa.

0.4 76' 0.525

0.214

@

-

--

-

-

3

0.533 ± 0.222

0.090

0

-

--

-

R 2672 f

20.797 ± 0.394

0.044

@

-

--

-

R2680b

1

0.803

-

0

-

--

-

-

R2402c

1

0.660

-

0

-

--

-

-

R2402g

1

0.960

-

0

-

--

-

-

-

--

5

1.563 ± 0.401

0.323

Sa

U 47 b

-

--

U128c

2

0.416' 0.302

0.034

0

2

2.429' 0.059

0.007

U46

3

0.623± 0.373

0.150

0

1

2.557

-

N 18 f3

0.4 1 7.0.237

0.095

0

4

1.885 ± 0.346

0.218

N 16 b

-

--

--

1

2.815

-

jWa

-

o

negativeallometry0<1•

po sit i v ea II 0 me try

Ci

> 1 +

@

i so met r y

1 -

~O'

~ a ~ 1 + ~C1'

cllom€'try

-

•
•
•
•
•

Kazushige TAN ABE

178

0.5

GK. H. B 9 B
U.4 6-.4

0.2• concentric

ringo divergent

rib

1 ... 0.0001 H 2 .557
r.O.9779

0.1

0.05

T=O,020Ho. S8O

0,03

r=0.9975

0,02
I

<

.-

.

0.1

0

0,05

3

5

8 10

20

.-

-

100

MM.3966
N1 B '-5 B

~

-

50

1"0.000.4 H 2.046
r=0.9702

0,03

with growth (WILBUR, 1964). Conse·
quently, the thickness of shell of theabove-mentioned specimens from theSantonian would make a remarkablecase of negative allometry. Actually,
the thickness of shell does not increase
with growth (see Plate 28). In recent
bivalves, such a case is found in someswimming and pseudoplanktonic forms(1. HAY AMI'S personal communication).
Theoretical studies of the planktonicmode of life of bivalves were accomplished by JEFFERIES and MINTON (1965)
and GOULD (1971). According to JEFFERIES and MINTON (1965), the physicalforces acting on the planktonic shell are
represented by the following formula.

0

E 0.02T",O.O 15 HO. 473.

r .. O,9740

"

0,01

8 10

I

~

-

0.1

20

50

100

50

100

GK. H. 10024

R2402c-l0~ 0,05~~.- 0.03••

T=O.OIOHo. 66O~~

Vspsg+ VPppg= (Vs+ VP)pwg+D

0,02

r .. 0.9726

0,01

B 10
He i9 h t

20in

Text-fig. 12. Ontogenetic allometry
between the thickness of the prismatic
layer and the shell height of the selected
specimens.

Paleobiological considerations
Mode of life.-Hereconsider themode of life of the present species on
the grounds of the mode of fossil occurrence and the biometrical analysis.
The results of the allometric analysisof the samples from the Santonian indicate the growth of the prismatic layer
in relation to the shell height to be
negative allometric. The thickness of
the inner layer of bivalve shell decreases

where Vs is volume of shell, Vp isvolume of the protoplasm, ps is thedensity of the shell, pP is the densityof the protoplasm, g is the acceleration
due to gravity and D is the drag coefficient. Generally speaking, the growthof the protoplasm is harmonious withthat of the shell. Consequently, the immersed weight of shell (Ws) is represented by the following formula (JEFFERIES and MINTON, 1965).
Ws= Vspsg- Vspwg=gVp(pw-pPHD

According to them, D depends on the
shape of the floating body. As ps ismuch greater than pw (ps is nearly 2.7
and pw is 1), HIS will increase withgrowth. In other words, the gravitational force scales as the cube of theshell length (L 3 ), while the total lifting
forces that could balance it scale atsmaller powers of L (GOULD, 1971).
The physical forces acting on thepseudoplanktonic shell are represented
by the following formula.
Ws= Vspsg- Vspwg-B

624.

Inoceramus (Sphenoceramus) naumanni

where B is the force, affected by byssus.
From these lines of evidence, it isconcluded that the mode of life of thepresent species was planktonic or pseudoplanktonic. at least, for the Santonian.
the individuals of which lacked divergent ribs.
If we scrutinize the early stage of thepresent species, it is found that thecritical point of growth exists at the179of the Santonian stage of 3-4 mm ofthe shell height. The thickness of theprismatic layer increases little (less than
10 fl) from the beak to that stage, but
at that stage, it increases abruptly, andthen keeps a constant thickness (30-40fl).
The growth index of the reduced major
axis in the early stage of the selected
specimens indicates near isometry, as
follows.

GK. H. 10001 (sample R 2672f): T=0.009fl1· 089 , r=0.9805
GK. H. 10002 (sample R 2672f): T=0.017fl1· o0 1, r=0.9797
GK. H. 871-1 (sample U 147-2): T=0.0l2flO· 9GO , r=0.9715

The outline of the shell in the earlystage is similar to that of veliger larvaeof recent bivalves. But in the middle
to the late stages, the outline becomes
mytiliform, having a straight posterodorsal margin, and it is similar to thatof the Lower Jurassic pseudoplanktonic
inoceramid,
Pseudomytiloides dubius(SOWERBY) (HAUFF, 1921). Abyssal
gape is sometimes observed in complete
and conjoined valves (eg. GK. H. 10003,
sample R 2680a; GK. H. 10004, sample
R 2680b).
Consequently, it may be postulated
that the present species changed its
mode of life from planktonic to pseudoplanktonic at a point of around 3-4mm
of shell height. It seems certain thatthe present species was pseudoplanktonic in the middle to the late stagesof growth in the Santonian.
It is presumable that individuals of
the present species attached themselvesby the byssus to flotsam and jetsam.
The growth index of the reduced
major axis of the thickness of the prismatic layer to the shell height also indicates negative allometry in the concentric ring stage of the specimens fromthe Campanian. The morphology of thisstage cannot be distinguished from thatof the Santonian species. On this evidence. it is reasonable to postulate thatthe mode of life of this stage was pseudoplanktonic.
On the contrary, at the beginning of
the divergent rib stage of every specimen, the growth index of the reduced
major axis of the thickness of the prismatic layer to the shell height abruptly
increases and indicates remarkable
positive allometry. In the divergent ribstage, the size of the prisms is muchgreater than in the concentric ring stage(see Text-fig. 11). As HAY AMI (1969)
pointed out, growth of this kind will
result in an increase of the settling velocity of the individual in seawater,
because of the increased weight of theshell. Consequently, the growth of thisstage is not suitable for maintaining a
pseudoplanktonic mode of life. For
example, in Mytilus edulis LINNE, whichattaches itself by byssus threads toblanching object among the recent
bivalves, the reduced major axis of the
thickness of the prismatic layer (T) to
the shell height (H) is
T = 0.013Ho. 9G7 , r = 0.979, N = 20.
The data were measured on a sample
from Nokonoshima, Hakata Bay, Fuku-

180

Kazushige TANABE

oka Prefecture, Kyushu. The groWthindex is smaller than that of the di·vergent fib stage of the present species.
As the change of ornament from theconcentric ring stage to the divergent
rib stage in ontogeny is discontinuous,
it is reasonable· to consider that themode of life changed from pseudoplank·
tonic to benthonic near the boundaryof these two stages. Actually, in theearly stage of growth of many recent
bivalves with radial ribs (e. g. Anadara
subcrenata (LISCHKE), A. infiata (REEVE),
Venerupsis (Amygdara) phillipinarum(ADAMS and REEVE) etc.), radial ribs
occur after the change in the mode oflife from· planktonic to benthonic (Yo.
SHIDA, 1935a, 1935b, 1937).
With respect to the mode of life of
fossil bivalves, there are many studies
such as HAUFF (1921), GUILLAUME (1928),
JEFFERIES and MINTON (1965), KAUFFMAN(1965, 1969), CARTER (1968), STANLEY(1968, 1972), KRIZ (1969), HAY AMI (1969)
and GOULD (1970).
Butovicella migrans (BARRANDE), a
Silurian bivalve, also displays a changeof morphology during ontogeny, similar
to that of the present species (KRIZ,
1969). KRIZ presumed the nepionic stage
to be planktonic and the radial rib stage(his neanic to gerontic stages) as pseu·
doplanktonic, though he gave no con·crete evidence.
JEFFERIES and MINTON (1965) discussed
the mode of life of two Jurassic bival·ves, Bositra buchi (ROMER) and " Posi·
donia" radiata GOLDFUSS. They con·cluded that the two species were nekto·
planktonic, from the analysis of themode of fossil occurrence, the verythin shell, and a model experiment. A
thin shell does not necessarily provide
evidence for a nektoplanktonic mode oflife and their conclusion is perhaps notjustifiable, because they used aluminium

for making a model of the inferredtentacles. Their conclusion was recently
criticized by GOULD (1970) and STANLEY(1972) who regarded the two species as
pseudoplanktonic. The relationship bet·
ween shell thickness and length in
Bositra buchi (JEFFERIES and MINTON,
1965, text·fig. 10) almost corresponds tothat of the present species.
As regards the mode of life of the
Inoceramidae, little direct evidence has
been presented, except HAUFF'S (1921)
observation.
It is very interesting that the shellmorphology of the Cretaceous Inocera·
midae is quite variable.
Moreover,generally speaking, they show a wide
geographic distribution. Such situations
may have given rise to adaptive mor·
phology of this family.
Evoiution.- After having considered
the mode of life of the present species,
the occurrence of the divergent ribs isregarded as an adaptive morphology
with the change of the mode of life
from pseudoplanktonic to benthonic.
The change can be observed not onlyin the ontogeny but also in the phylo·
geny of the present species. Indeed,
there are no morphological difference
between the specimens that were col·lected from the Santonian and thosefrom the Campanian, apart from theoccurrence of the divergent ribs andthe thickening and enlargement of theshell in the latter.
Consequently, the main part of thegroup of Inoceramus naumanni of NA·
GAO and MATSUMOTO (1940) is regarded
as a single chronospecies, Inoceramus(Sphenoceramus) naumanni YOKOYAMA
emend., and it is divided into two chro·nological subspecies, namely I. (5.) nau·manni naumanni YOKOYAMA emend. andI. (5.) naumanni schmidti MICHAEL emend.
The former is represented by the624 . Inoc era?n'us (S1Jh enoce?·a?n~~s) nau?nanni

181

9 00r-----TT.?~--,-----,_----_r----_r----_.----_,r_----.__,I'I

EVOLUTION AND .

MODE OF LI FE

OF

800

Inoceramus

naumanni YOKOYAMA em .

(URAKAWA SECTION)
700

z
«

600

z
«
<l.

500

~

c

«

g 400
"

u
Benthan i c

a

stage

I

300

200

Puudoplanktanic

stag e100-__ _ ?_°0~----t-----~10~---'1~5----~2~0-----f.25~--~30~--~3~5----~40~:J
Heightinmm

Text-fig . 13. Evo lution and mocle of life of t he chronospec ies, Inoceramus(SjJhenoce ramu s) naumanni YOK OYAMA emend. (U r akawa sect ion , show in g themean and the 95% confidence in terva l of the m ean of the s h e ll hei g ht in t hepseudoplankton ic s ta ge).

··specimens from the Coniacian to the-Santonian and is ornamented w ith concentric ri ngs only. The latter is represented by the specimens from th e Ca mpanian and has two stages of g ro wth ,namely, the stage w ith concent ric rings.alone and that with divergent rib s and. concentric rings both.
The evo lution of the present chrono. species is diagrammaticall y illustrated
in Text -fig. 13.
According to the biostratigraphic division of the Upper Cretaceous of Van• COLl ve r Islands and the Gulf Islands,

British Columbia (MU LLER and JELETZKY, 1970) and that in Cali fornia (MA TSUMOTO, 1960), th e same patte rn ofchange in th e morphology with time
can be expected.
As r egards the origin of the present
speci es, I hav e no convincing e vidence
f rom th e T uronian in Japan . At present,
it can onl y be indi cated that Inoceramus(i\lJytiloides) labiatu s (SCHLOTHEIM), a
Lower Turonian cosmopolitan sp eci es,
is morphologicall y somew hat simi lar toth e present s pec ies .

Kazushige TANABE

182
References

BOUCOT, A. J. (1953): Life and death assemblages among fossils. Amer. Jour.
Sci., vol. 251, pp. 25-40.
CARTER, R. M. (1968): Functional studies
on the Cretaceous oyster Arctostrea.
Palaeontology, vol. 11, pt. 3, pp. 458-485,
pIs. 85-90.
CRAIG, G. Y. (1967): Size-frequency distributions of living and dead populationsof pelecypods from Bimini, Bahamas,
B. W. 1. Jour. Ceol., vol. 75, no. 1, pp.
34-45.
GOULD, S. J. (1970): Evolutionary paleontology and the science of form. Earth
Sci. Rev., vol. 6, pp. 77-119.
- - (1971): Muscular mechanisms and theontogeny of swimming scallops. Palaeontology, vol. 14, pt. 1, pp. 61-94.
GUILLAUME, L. (1928): Revision des posidonomyes jurassiques. Bull. Soc. Ceoi.
France, ser. 4, tome 27, fasc. 3, pp. 217234, pI. 10.
HALLAM, A. (1967): The interpretation ofsize-frequency distributions in molluscan
death assemblages. Palaeontology, vol.
10, pt. 1, pp. 2525-2542.
HAUFF, B. (1921): Untersuchung der FossilfundsUitten von Holzmaden im Posidonienschiefer des oberen Lias Wiirtembergs. Palaeontographica, vol. 64, pp.
1-42, pIs. 1-21.
HAYAMI, 1. (1969):
Notes on Mesozoic" planktonic" bivalves (in Japanese).
Jour. Ceol. Soc. Japan, vol. 75, no. 7, p.
375-385.
- - and MATSUKUMA, A. (1970): Variationof bivariate characters from the standpoint of allometry. Palaeontology, vol.
13, pt. 4, pp. 588-605.
HUXLEY, ].S. (1932): Problems of Relative
Crowth. 276 pp., Dial Press, New York.
IMBRIE, ]. (1956): Biometrical methods inthe study of invertebrate fossils. Bull.
Amer. Mus. Nat. Hist., vol. 108, art. 2,
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JEFFERIES, R. P.S. and MINTON, P. (1965):
The mode of life of two Jurassic speciesof " Posidonia" (Bivalvia). Palaeontology,
vol. 8, pt. 1, pp. 156-185, pI. 19.
KADO, Y. (1953): On the scheme of theshell structure of Lamellibranches. Jour.
Sci. Hiroshima Univ., ser. B, div. 1, vol.
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KANIE, Y. (1966): The Cretaceous deposits
in the Urakawa district, Hokkaido (inJapanese). Jour. Ceol. Soc. Japan, vol.
72, no. 7, pp. 315-328.
KANMERA, K. (1969): Litho- and biofaciesof Permo-Triassic geosynclinal limestoneof the Sanbosan belt in southern Kyushu.
Palaeont. Soc. Japan, Special papers, no.
14, pp. 13-39, pIs. 4-8.
KAUFFMAN, E. G. (1965): Taxonomic, ecologic and evolutionary significance of interior shell morphology in the Inoceramidae (Mesozoic Bivalvia). Abst. Prog.
1965 Ann. Meet., Ceol. Soc. Amer., p. 95.
- - (1967): Coloradian macro-invertebrate
assemblages, central western interior
United States. Paleoenvironments of theCretaceous Seaway, a symposium, Colorado
Sch. Mines, Colden, pp. 67~143.
- - (1969): Form, function and evolution.
In MOORE, R. C. (ed.) Treat. Invert.

Explanation of Plate 27
Mode of fossil occurrence in calcareous nodule (x 3).
Figs. 1-5. Inoceramus (Sphenoceramus) naumanni naumanni YOKOYAMA emend.
Fig.!. Sample R 2665a, The R. Jugosen, Tappu district. Fig. 2. Sample H 2003b,
The upper course of the R. I-Iobetsu, Prov. Iburi, southern central Hokkaido. Fig. 3.
Sample R 2680b, The R. Jugosen, am; another species of Inoceramus, probably I. (Piatyceramus) amakusensis NAGAO and MATSUMOTO. Fig. 4. Sample R 2672f, The R.
Jugosen. Fig. 5. Sample R 2686a', The R. Jugosen.
Fig. 6. Inoceramus (Sphenoceramus) naumanni schmidti MICHAEL emend. Sample N 18f, theR. Miho, Naibuchi district.

T ANABE :Evol~~tionand Mode of Life of Inocemmus

Plate 27

624.

Inoceramus (Sphenoceramus) naumanni

Paleont., Part N (Mollusca 6, Bivalvia),
p. N130-N205.
KERMACK, K. A. and HALDANE, J. B. S. (1950) :Organic correlation and allometry. Biometrika, vol. 37, pp. 30-41.
KOBAY ASH!, 1. (1971): Internal shell microstructure of recent bivalvian molluscs.
Sci. Rep. Niigata Univ., ser. E, no. 2, pp.
27-50, pis. 1-8.
KRIZ, J. (1969): Genus Butovicella, 1965 in
the Silurian of Bohemia (Bivalvia).
Sbornik Ceol. Ved., Pal. rada, p. sv. 10,
pp. 105-139, pis. 1-8.
MATSUMOTO, T. (1942): Fundamentals in
the Cretaceous stratigraphy of japan.
Part 1, Mem. Fac. Sci., Kyushu Imp.
Univ., ser. D, vol. 1, no. 3, pp. 133-280.
- - (editor) (1954): The Cretaceous Systemin the japanese Islands. The Cret. Reser.
Commit., japan Soc. Prom. Sci., Ueno.,
Tokyo. 324 pp., pis. 20.
- - (1959): Zonation of the Upper Cretaceous in japan. Mem. Fac. Sci., Kyushu
Univ. ser. D. Geol., vol. 9, no. 2, pp. 5593, pis. 1-2.
- - (1960): Upper Cretaceous Ammonites
of California. Part. 3. Ibid., Special vol.
2, pp. 1-204, pis. 22-27.
- - and NODA, M. (1968): An interesting
species of Inoceramus from the Upper
Cretaceous of Kyushu.
Trans. Proc.
Palaeont. Soc. japan, N. S., no. 71, pp.
317-325, pI. 32.
MISIK, M. (1966): Microfacies of the Mesozoic and Tertiary Limestone of the West
Carpathians. 269 pp., 101 pis., Vydatel'stvo slov. Akad. Vied. Bratislava.
MuLLER, J. E. and jELETZKY, J. A. (1970):
Geology of the Upper Cretaceous Nanaimo Group, Vancouver Islands and Gulf
Islands, British Columbia. Ceol. Surv.
Canada, paper 69-25, 77 pp.
NAGAO, T. and MATSUMOTO, T. (1940): A
monograph of the Cretaceous Inoceramus
of japan. Part 2. jour. Fac. Sci., Hokkaido Imp. Univ., ser. 4 (geol. miner.),vol. 6, pp. 1-64, pis. 1-22.
NEWELL, N. D. (1956):
Fossil populations
In SYLVESTER-BRADLEY, P. C. (ed.) The
SPecies Concept in Palaeontology. The

183

Systematics Assoc. Publ. no. 2, London,
pp. 63-82.
SEITZ, O. (1962): Ueber Inoceramus (Platyceramus) mantelli MERCEY (Barois) aus
dem Coniac und die Frage des Byssus"
Auschnittes bei Oberkreide-Inoceramen.
Beih. Ceo!. jb., no. 79, pp. 353-386, pis. A -:- I
10-13.
SIMPSON, G. G. and ROE, A. (1939): Qauntitative Zoology. 414 pp., New York.
- - and LEWONTIN, R. C. (1960) : Quantitative
Zoology. Revised ed., 440 pp., Harcourt
Brace.
SOKAL, R. R. and ROHLF, F. J. (1969): Biometry. 776 pp., Freeman, San Francisco.
STANLEY, S. M. (1968): Post Paleozoic adaptive radiation of infaunal bivalve molluscs. (A consequence of mantle fusionand siphon formation). jour. Pa/eont.,
vol. 42, no. 1, pp. 214-229.
- - (1972):
Functional morphology andevolution of byssally attached bivalve
mollusks. Ibid., vol. 46, no. 2, pp. 165212.
SYLVESTER-BRADLEY, P. C. (1958): The description of fossil populations.
Ibid.,
vol. 32, no. 1, pp. 214-235.
TANAKA, K. (1963): A study of the Cretaceous sedimentation in Hokkaido, japan.
Ceol. Surv. japan, Rep., no. 197, 119 pp.,3 pis.
VISHER, G. S. (1969): Grain-size distribution
and depositional processes. jour. Sed.
Petrol., vol. 39, no. 3, pp. 1074-1106.
WALLER, T. R. (1969):
The evolution of
Argopecten gibbus stock (Mollusca: Bivalvia), with emphasis on the Tertiary
and Quaternary species of eastern North
America. Paleont. Soc. Mem. 3, 125 pp.,pis. 8.
WILLBUR, K. M. (1964): Shell formation andregeneration.
In WILBuR, K. M. and
YONGE, C. M.(ed.) Physiology of Mollusca.
vol. 1. Academic Press, pp. 243-282.
YOSHIDA, H. (1935a): On the latter pelagic
larvae and early benthonic youngs ofAnadara infiata (REEVE) (in japanese).
Venus, yol. 5, nol. 1, pp. 1-10, pI. 1.
- - (1935b):
On the full-grown veligers
and early young-shell stage of Veneru-

Kazushige TANABE·

184

psis phillipinarum ·(ADAMS and REEVE)(in Japanese). Ibid., vol. 5; pp. 264-273,
pI. l.
(1937): On the veliger larvae and. Abeshinai
Akanosawa
ChinomiHobetsu
IkandaiJugosenKamihaboro
Kotanbetsu
Miho

7--":,/;1-'1

7

"j]

/

iR

¥L
f}!i

:fj:
'}Jlj

j:!:

..L>..
r::l*-I· Ii. *Jl!
J: ~~ $X
J!l ft }Jlj~

f:!!!:

youngs of Anadara subcrenata· (LISCHKE)(in Japanese). Ibid., vol. 7, pp. 5-11,
pI. 1.

NaibuchiObirashibe
Rumoi
Sakasagawa
Shutta-za wa
Tannosawa
Tappu
Urakawa
Urokobetsu

r"J

iiPrl

Ij,

.if ~

fJ1

Wi

)II
i!E
'/=>''Y!llR
(J)
iR
)!
l'ri

m

)oJ
t7 ]:I ::1' lJlj

im·

Explanation of Plate 28
Figs. 1-4, 7-9. Inoceramus (Sphenoceramus) naumanni naumanni YOKOYAMA emend.
Fig. 1. GK. H. 10009, Sample R 2672f, The R. Jugosen (x 10). Fig. 2. GK. H. 10005,
Sample R 2680b, The R. Jugosen (x 10). Fig. 3. GK. H. 10007, Sample R 2672f, The
R. Jugosen (x 10). Fig. 4. GK. H. 10004, Sample R 2680b, The R. Jugosen (x 10).
Fig. 7. Lateral view of right valve, GK. H. 10019, Sample R 2696a, The R. Jugosen(x 1). Fig. 8. Lateral view of left valve, GK. H. 721a, Sample U 143r, The uppercourse of the R. Chinomi (x 1).
Figs. 5-6, 10-18. Inoceramus (Sphenoceramus) naumanni schmidti MICHAEL emend.
Fig. 5. GK. H. 10011, Sample U 46, The upper course of the R. Urokobetsu, Urakawa
district (x 10). Fig. 6. GK. H. 10032, Sample T 1219 P (in a fallen or. rolled nodule),
The R. Tannosawa, Abeshinai district, northern central Hokkaido (T. MATSUMOTO
colI.) , comparative specimen (x 3). Fig. 10. Lateral view of right valve, GK. H. 882,'
Sample U 147-2, the upper course of the R. Chinomi (x 1). Fig. 11. Lateral view ofright valve, GK. H. 871b, Sample and Loc. ditto (x 1). Fig. 12. Left view of theconjoined valves, GK. H. 10034, Sample IA 336, Northwest of Tomiuchi station, Tomiuchi district (A. INO;\'IA colI.) , comparative specimen (x 1). Fig. 13. Lateral view ofleft valve, MM. 3963, Sample N 18f, the R. Miho (x 1). Fig. 14. Lateral view of leftinternal mould, YCM. U 446-4-1, Sample U 446-4, the lower course of the R. Chinomi(x 1). Fig. 15. Lateral view of right valve, GK. H. 906a, Sample U 47b, the uppercourse of the R. Urokobetsu (x 1). Fig. 16. Lateral view of left valve, GK. H. 935,
Sample and loco ditto. (x 1). Fig. 17. Lateral view of right internal mould of theconjoined valves, GK. H. 903, Sample and Loc. ditto. (x 1). Fig. 18a, b. Lateral andventral view of right valve, GK. H. 930, Sample U 128c, The upper course of the R.
Urokobetsu (x 1).
01: Outer prismatic layer, il: Inner nacreous layer, dr: Divergent rib stageTANABE :Evo lution and Mode of Life of Inoceram~~sPlate 285~d~ -'" ~
"
., "}

~.~
.
2
1
, " '--:~.

'~'.

~

16

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 92, pp. 185-199, pI. 29, December 20, 1973

625. LOWER CARBONIFEROUS CONODONTS FROM THE
AKIYOSHI LIMESTONE GROUP, SOUTHWEST JAPAN.*
HlSAHARU lGO
Department of Earth Science, Tokyo Gakugei University

f:ktrfiEk:'5®~'f-O)J1:!:TJ'4IIIlJiO)tlli't~IIHl::fc.W!J L '"( f'1~W(,jji.M~ It, ~1!ff'1 i:. O)t1t1!lli~f1¥~T 0
t:.CJ:;, :;k~f~H1i!ifc.5J;{jjT Q 1f,IEEk:'5:®:Tlt~n1O) ::I /H}fJ"E Lt:'.~.:t·O)*,d*, t1tWi¥J
h::'7J;{jj-99 Gnathodus bilineatus G. commutatus ~f;?tr::I / ~.!:,-' rft::fiil¥~~,~CJ:;t.:o i:.hGO)::I / ".:,-' HtEU'f-f'1, r.iiT1&EEk:'6O)~~n1 c If.m~t,dtt=iil'f-iH? tJ: I), ::It7j,:O) Chesterian
Tf;lI, '*t::f'13 - P ",/~O) Visean J:.f;llfO;J-J:I:~hQo'
nm ~ iti
0r:,-' "

cm

Lower Visean. OKIMURA (1963, 1966)
worked out the endothyroid foraminifers
collected from the lower part of the
Akiyoshi Limestone Group.
His lowermost biostratigraphic unit is
the Endothyra sp. A Zone. He insisted
that this zone can be correlated to theLower Visean.
MINATO and KATO (1963) studied thecorals and brachiopods collected fromthe reddish tuffaceous shale near thebase of the Akiyoshi Limestone Group.
They emphasized that the fauna is apparently younger than the Upper Visean
Onimaru Series and equivalent to theLowest Namurian.
Subsequently, however, Y ANAGlDA(1965) concluded that the mentioned
fauna indicates the Lower to Middle
Visean. OT A (1968) also discussed theage of the lowermost part of the Akiyoshi Limestone and considered that
Zaphrentoides and other corals and brachiopods yielded from the limestone
below the mentioned reddish tuffaceous
shale show the Late Tournaisian to Early
Visean in age.
I have studied the conodont faunacollected from the lowermost Akiyoshi

Introduction and Acknowledgements
After the monumental research by
Yoshiaki OZAWA (1923, 1927), geology
and paleontology of the Akiyoshi Limestone Group distributed in the Akiyoshi
plateau, Yamaguchi Prefecture, Southwest Japan, have been repeatedly studied
by TORIY AMA (1954, 1958) and manyothers.
There are, however, several different
conclusions concerning the geologic
structure, biostratigraphic subdivisionand correlation. The purpose of thisresearch is to elucidate geologic age ofthe lowermost part of the Akiyoshi
Limestone Group based upon the sequence of conodonts. Concerning geologic age and correlation of the lowermost Akiyoshi Limestone Group by
foraminifers, brachiopods and corals
seem to diverge somewhat as summarized below.
MURATA (1961) set the Endothyra
Zone in the basal part of the Akiyoshi
Limestone Group and correlated to the* Received Sept. 2, 1972; revised May 23,
1973; read Sept. 13, 1970 at Tokyo185

Hisaharu IGO

186
1320

131 0

§~ICHERT
vvvv

v vv

v

VVVVv

o

HIRABARUI

Text-fig. 1.

Geological map of the Ohkubo-Hirabaru area, Akiyoshi Plateau.

Limestone Group distributed in the Ohkubo area, southern margin of the Akiyoshi plateau (Text-fig. 1). As a result,
I am inclined to believe that this conodont fauna is apparently correlated to
the Upper Visean of western Europe or
the Chesterian of North American Mississippian.
This research based on a Master's
thesis, is done at the Department ofEarth Science, Tokyo Gakugei University. I wish to acknowledge continuous
supervision of Professor Mosabro KANUMA of Tokyo Gakugei University. I
am greatly indebted to Drs. Hisayoshi
IGO and Toshio KOIKE of Tokyo Uni-

versity of Education for their guidance
and fruitful discussion during the present research. I thank Dr. Masamichi
OTA of the Akiyoshi-dai Science Museum
for his kind facilities in the field.
Stratigraphic Summary

The lowermost part of the Akiyoshi
Limestone Group is distributed in the
Ohkubo area, southern margin of the
Akiyoshi plateau. It crops out typicallyalong the road-side cutting from Hirabaru to Ohkubo and consists mainly of
massive limestone intercalating tuffaceous limestone, basic tuff and lava

625.

Carboniferous conodonts from Akiyoshi

I

76007

I 76008
I 76009
1760106011
177601217601 3a-b
Text·fig. 2. Columnar sections of the
Ohkubo-Hirabaru area showing conodontbearing levels. Index of lithology is thesame with Text-fig. 1.

flows. They are trending N 50·E to N
70·E and dipping 30·S to 50·S. The
following stratigraphic succession canbe observed along the above-mentioned
type section in descending order (Textfig. 2).
Gray crinoidal massive limestone ......... .
.................. more than 20 m thick.
Greenish gray tuffaceous crinoidal well·bedded limestone intercalating reddish
part toward lower .......... 20 m thick.
Reddish tuffaceous shale ................... .
.......... 10 m thick, as far as exposed.

187

Covered .......................... 15 m thick'Gray massive crinoidal limestone ......... .
........... " ... -" ....... _... 30 m thick.
Green and variegated tuff intercalating lava
flows, limestone lens of about 10 m thicknear the top, and thin cherty layer near
the base .......... more than 45 m thick.

The lowermost unit is in contact
with sandstone of the Ota Group by afault. Although conodonts are barren
in the limestone lens intercalated in thisunit, OKIMURA (1965) reported the occurrence of Endothyra sp. A. Next
younger limestone unit is also lacking
in conodonts, but it represents the upper
part of OKIMURA'S Endothyra sp. A
Zone. Gray massive crinoidallimestone
situated above greenish tuff yields Gnathodus bilineatus and other conodonts,
but their occurrence is restricted in thelower part of this lithologic unit. This
unit represents OKIMURA'S Endothyra
similis Subzone.
Reddish tuffaceous shale yields Pleurodictyum dechenianum and other corals
and brachiopods (MINATO and KATO, op.
cit.). According to YANAGIDA et al.
(1971), stratigraphic position of thisfossil bed is included within OKIMURA's
Endostaffella delicata Zone.
The next higher greenish gray tuffaceous limestone occurs numerous
conodonts from various levels. OT A
(1968) reported Zaphrentoides sp., Cyathaxonia sp., Nagatophyllu17! satoi OZAw A, Syringothyris sp.. Gigantoproductus
sp. and other fossils from this unit.
The uppermost lithologic unit of thepresent section also comprises the upper
part of OKIMURA'S Endothyra similis
Subzone and it contains numerous conodonts. This is overlain by massive graylimestone containing A1illerella sp., Pseudostaffella antiqua and other primitive
fusulinaceans.
As already mentioned in earlier lines,

188

Hisaharu Jeo

I

I

I ,II I
'-

I

I~ i

I

';

,

I

I

I

t

III

~

~
\1.1

I

~
IIlU

Zill
u:J

~

I

"'~
I ttI'-ttl
.c:J
UU

III
.:J III C
X :J 111
I1I.cC!),III III

:J:J

~-:d

>~

cttI

oC
u01
III

III :J
:J>
.cttl

-;;;u
c

based on the occurrence of endothyroid
foraminifers. ETO (1967) supported OKIMURA'S view by his sedimentologicalresearch. On the contrary, YANAGIDA
and OTA (1964) concluded that this sequence is in normal stratigraphic orderand the so-called schalstein does not
occupy the base of the Akiyoshi !:-imestone Group.
According to YANAGIDA et al. (op. cit.),
however, the basal part of the Akiyoshi
Limestone Group predominates volcaniclastic. rocks in the Shishidedai area,
northeastern part ot'the Akiyoshi Limestone plateau.

01
III

:J

>

ttl
U

I I
VV VY..Y.
I../YV
vy\
VVV

~V~v
!Vvvy~VVVVI

I V V V:~Vvvv
IVV V IVV Y VIV V V\YV V~
IV Y VVVVV[; Vv ~.~

1/VVv
vvy\

VVVV
';' v VV\lvV/v\

Text·fig. 3. Stratigraphic distribution ofimportant species of conodonts.

the present sequence is dipping southwards, and OKIMURA (1963, 1966) insisted that this sequence is reverse instratigraphic order and overturned

Conodont Fauna and Correlation

Conodonts are recovered from 2 to 3
kg samples of carbonate rocks. Sampling was made from the measured section
at intervals of 3 to 5 m.
The lowest conodont-bearing limestone(76002) y i e Ids Cnathodus bilineatus(ROUNDY), C. commutatus (BRANSON and
MEHL), C. nodosus BISCHOFF, C. akiyoshiensis IGO, n. sp., Ozarkodina delicatula(STUFFER and PLUMMER), O. spp. andSpathognathodus campbelli REXROAD.
Of these species, Cnathodus nodosus
and C. bilineatus are particularly abundant. C. akiyoshiensis, restricted in thislevel, is similar to C. commutatus.
The next higher conodont-bearing
limestone (76006, 76014) is situated immediately above the reddish tuffaceous
shale. It yields the following conodonts,
namely, Cnathodus bilineatus (ROUNDY),
Hindeodella spp., Lonclwdina spp., LigoNeoprioniodus peractus
nodina spp.,(HINDE), Ozarlwdina spp., SPathognathodus campbelli REXROAD and S. cristulus YOUNGQUIST and MILLER. Amongthe listed ones, Cnathodus bilineatus,
Hindeodella spp. and Spathognathodus
campbelli are particularly abundant but

625.

Carboniferous conodonts from Akiyoshi

Table 1.

Distribution of conodonts from the Ohkubo area.
abundant /::;. common x rare

189

o

~
Spec i es

Cavusgnathus convexus
C. charactus
Geniculatus claviger
Gnathodus bilineatusN
M -4".0 e.ge-- (Xl a> 0 _ N M -4".0
o00 0000 00 00

'" '"

X
X
X
I::. 01::. o X X X I::.

G. commutatuso 0

G. homopunctatus

X

0

X

G. aldyoshiensis n.sp.

0

Hibbardella spp.

I::.

Hindeodella spp.

o I::. IJ. I::. X 0

Lonchodina furnishi

X

L. spp.

X I::.

Ligonodina spp.

X I::.

0

X

X

IJ.
X X

X

I::.

0

I::.

X X

N. peracutus
N. scitulus

I::.

0

G. aff. texanus

N. montanaensis

X XX X

00

G. nodosus

Neoprioniodus varians

00 00

00 00 00 00 00 00
e.g
e.g e.g",(0 e.g e.g to'" e-- e-- e-- e-- e--e-- toto '"e--e-- e--e-- e.g
e-- '"e-- e-e-e--r--X

X
X

Ozarkodina delicatula
O.sPP.
S~a thognathodus cristulus

X X

S. campbelli

01::.

X

X

X X

X
X

X X X

Gen.et sp. indet.

other species are subordinate in occurrence.
The localities 76004, 76005, 76012 and76013 represent almost the same stratigraphie level situated about 3 m above
the previous level.
I discriminated the following species,
namely, Gnathodus bilineatus (ROUNDY),
Hindeodella spp., Lonchodina spp., Ligonodina spp., Neoprioniodus peractus(HINDE), N. scitulus (BRANSON and MEHL),
Ozarkodina delicatula (STUFFER and
PLUMMER) and Spathognathodus cristulus
YOUNGQUIST and MILLER.
They are

0

X

X

X

0

X X

X

0

X X X

almost the same in individual number,and there are no predominating species.
The next younger conodont assemblage is yielded from the bedded limestone (76001, 76003, 76008, 76009, 76010)
and it consists of Gnathodus bilineatus(ROUNDY), G. commutatus (BRANSON and
MEHL), Hindeodella spp., Neoprioniodus
montanaensis (SCOTT), Ozarkodina delicatula (STUFFER and PLUMMER), SPathognathodus campbelli REXROAD, S.
cristulus YOUNGQUIST and MILLER, Lonchodina furnishi REXROAD, Cavusgnathus
convexus REXROAD, Gnathodus homopunc-

190

Hisaharu [GOtatus ZIEGLER, Hibardella spp., Neoprioniodus seitulus (BRANSON and MEHL) andOzarkodina spp.. Among them, Gnathodus bilineatus, G. eommutatus, Neoprioniodus
montanaensis,
SPathognathodus eampbelli and unidentified Hindeodella are predominating species inthis assemblage.
Gnathodus homopunetatus is very rare
but this species becomes abundant in
the higher levels.
The following three species are obtained from the limestone (76000) situatedabout 10 m above just aforementioned
one, namely, Cavusgnathus eharaetus
REXROAD, Gnathodus bilineatus (ROUNDY)
and Neoprioniodus varians (BRANSON and
MEHL). Brachiopodal limestone (76007)
4 m above the level (76008) contains afauna of the following species: Genieulatus claviger (ROUNDY), Gnathodus
G. eommutatusbilineatus (ROUNDY),(BRANSON and MEHL), G. homopunetatusZIEGLER, G. aff. texanus ROUNDY, Hindeodella spp., Neoprioniodus montanaensis(SCOTT), Ozarkodina delicatula (STUFFER
and PLUMMER), O. spp. and Spathognathodus eampbelli REXROAD. Abundantand distinctive species in this assemblage are Gnathodus eommutatus, G.
homopunetatus, Hindeodella spp. andSPathognathodus eampbelli.
Although the precise stratigraphic
relationship could not be settled in thefield, Gigmitoproductus-bearing coquinoid
limestone (76015) yields the following
conodonts which are assumed as theuppermost representative of conodont
assemblage in the Ohkubo area.
They are Gnathodus bilineatus (ROUNDY), Hindeodella spp., Lonehodina spp.,
Neoprioniodus seitulus (BRANSON and
MEHL), Ozarkodina spp. and SPathognathodus campbelli REXROAD.
Among the listed ones, the last mentioned species is extraordinarily abun-dant.
As described above, the conodontbearing levels are not entirely continuous except for the middle part ofthe measured section. More extensive
investigation throughout the lowermost
part of the Akiyoshi Limestone Group
distributed outside of the Okubo area
seems to be necessary to settle thedetailed conodont biostratigraphy andzonation. Therefore, detailed zonationof the present fauna is reserved in the
future.
I will discuss the range of the several
important species recovered from thepresent section hitherto pointed out bymany authors in the following lines.
Gnathodus bilineatus occurs from almost
entire part of the measured sections inthis area. Two different types are recognized within numerous specimens
identified as G. bilineatus from thepresent area. The material obtained
from the limestone below the reddish
tuffaceous shale is typical Gnathodus
bilineatus which is characterized bywell-developed bilinear arrangements of
the nodes on the inner platform. However, specimens yielded from higher
levels decrease in number of the nodes
on the inner platform and with shorter
parapet-like transverse ridges on theouter platform compared with those of
the typical one.
This type of G_ bilineatus resembles
MEISCHNER'S (1970) G. bilineatus sehmidti.
He concluded that this subspecies is thedescendant modification of G. bilineatlls
bilineatus.
G. bilineatus is a cosmopolitan species
and has been reported from the Upper
Meramecian Ste. Genevieve Limestone(tothe Middle Chesterian Glen Dean Limestone)or equivalent ones of the USA. This
species is also known from the uppermost Cull 0 (upper part of the Perieyclus

625.

Carboniferous conodonts from Akiyoshi

Zone) to the E2 Zone (upper part of the
Eumorphoceras Zone) in Germany. but
it is characteristic in the Upper Visean.
C. commutatus is abundant and characteristic species in the present fauna.
This well-known species has been reported from the Meramecian Ste. Genevieve Limestone through the Chesterianof the North American Mississippian.
It is also known from the Cull f31r
(Pericyclus Zone) in Germany and theupper part of the D2 to E2 in the British Avonian Succession.
C. nodosus is also an important species
in the present section, but it is restrictedin the lower part. C. nodosus has beenrecorded from the CuIII B (middle Coniatite Zone) of Germany and from the
D3 to E2 of the Avonian Succession.
C. homopunctatus is predominating in
the upper part of the lowermost Akiyoshi Limestone Group. This species
is having almost the same stratigraphic
range with C. commutatus and has been
known from the entire part of the
Visean to the Lower Namurian in Europe.
C. aff. texanus came from the higher
part of the present succession has
slightly different ornamentation of theplatform compared with ROUNDY's typical specimen (HASS, 1953). This species
is ranging from the Upper Osagian to
the Chesterian of the USA. It yields
from the Cull f31r to CuIII a of theGerman Carboniferous and from the C1
to Sl Zones of the British Avonian.
Ceniculatus claviger is yielded fromthe higher part of the present sequence.
This characteristic species was firstdescribed from the Barnett Shale in
Texas (ROUNDY, 1926). Subsequently,
BISCHOFF (1957), VOGES (1959), DRUCE(1969) and others reported this species
from the CUIII a of German Carboniferous and Australia.
Spathognathodus campbelli is common

191

throughout the present section. This
species was described from the upper
D2 to D3 in England and also from theUpper Meramecian to Chesterian of theUSA.
Cavusgnathus convexus occurred fromthe immediately above reddish tuffaceous shale.
This species is rather
long-ranging and recorded from theUpper Osagian (?) to Upper Chesterian.
Many compound types of conodont
are yielded from the present sections
associated with the above mentioned
important species. They are also characteristic species or allied ones withthose of the North American Mississippian or Lower Carboniferous in Europe.
Based upon the foregoing discussion,
I will express my views concerning thecorrelation of the present fauna withthe faunas from other areas in Japanand from North America and Europe.
The present fauna is very similar to
those from the lower part of the Atetsu
Limestone. the Nagoe Formation which
was studied by KOIKE (1967). There is,however, an important difference in thestratigraphic distribution of C. nodosus.
This species is restricted in the lower
part of the present succession, but it
persists within the upper part of the
Nagoe Formation. The other remarkable difference between the Nagoe andthe present Akiyoshi is the occurrenceof Idiognathoides noduliferus. I. noduliferus is dominant in the upper Nagoe
Formation in association with C. bilineatlls, but it is barren in the Ohkubo
fauna. These differences recongnized
between both faunas can not be confirmed whether interregional faunaldifferentiation or chronological problem.
The Uzura fauna of the Akiyoshi
Limestone Group reported by IGO and
KOIKE (1965) consists of quite dissimilarelements which were advocated as the

192

Hisaharu lGO

Lower Pennsylvanian. There is alsostriking difference between the present
fauna and the Lower Pennsylvanianfauna described from the Omi Lime·stone (IGO and KOIKE, 1964).
The correlation of the present faunawith the faunas reported from Europe
and North America is one of the mostimportant purpose of the present investigation. Concerning the intercontinental correlation, as already discussed
in earlier lines, the present fauna containes many species which are characteristic in the Chesterian of the" Upper
Mississippi Valley Region" (COLLINSONet al., 1962 and others). Moreover, thepresent fauna is entirely avoided theimportant Lower Pennsylvanian species.
Also the present conodont assemblage .
is very similar to those from the Gnathodus commutatus and G. nodosus Zones
which represent the Upper Visean ofEurope. In conclusion, the present sequence of conodonts in the lowermost
part of the Akiyoshi Limestone Group
is very similar to those of the Chesterian of the USA and of the Upper
Visean of Europe.

given by RHODES, AUSTIN and DRUCE(1969, p. 79, 80). The specimens obtained
from Akiyoshi are very similar to thepreviously described ones.
Distribrution :-Restricted in locality
76000.
Reg. no. 1057.
Genus Geniculatus HASS, 1953
Geniculatus claviger (ROUNDY)
PI. 29, fig. 28

Polygnathus? claviger ROUNDY, 1926, p. 14,
figs. la-c, 2a, b.
Geniculatus claviger, HASS, 1953, p. 77, pI.
15, figs. 10-19; BISCHOFF, 1957, p.21, pI.
1, figs. 1-6; WIRTH, 1967, p. 25, pI. 19,
fig. 5; DRUCE, 1969, p. 60; MARKS &WENSINK, 1970, p. 257, pI. I, fig. 4.

Remarks :-Detailed synonymy anddescription were given by HASS (1953,
p. 77, 78). The Akiyoshi specimens arequite identical with the previous foreign
ones.
Distribution :-Yielded from locality
76007.
Reg. no. 1082.

Description of Species

Genus Gnathodus PANDER, 1856

Genus Cavusgnathus HARRIS &HOLLINGSWORTH, 1933

Gnathodus akiyoshiensis IGO, n. sp.

Cavusgnathus charactus REXROAD
PI. 29, fig. 26

Cavusgnathus characta REXROAD, 1957, p. 15,
16, pI. 1, figs. I, 2; REXROAD & COLLI:>:SO:<1, 1963, p. 8, pI. 1, fig. 28; THO:VIPSON& GOEBEL, 1969, p. 22, pI. I, figs. 1,4,7.
Cavusgnathus charactus, RHODES, AUSTIN &DRUCE, 1969, p. 79, 80, pI. 13, figs. 6a7d, 13a-c.

Remarks:-A detailed synonymy was

PI. 29, figs. 24, 25

The unit consists of cup with sharply
pointed posterior end and long free
blade. Conspicuous large and highnode-like denticle developed at centerof platform. Surface of platform smooth
and without any ornamentation. Outlineof platform elliptical in shape. The
aboral edge is straight in lateral view,
but oral edge almost straight in theanterior four-fifths of unit and posteriorly declines with steep angle. Ex-

625.

Carboniferous conodonts from Akiyoshi

panded basal cavity is elongated inoutline.
Remarks:- This species resembles
Spathognathodus campbelli REXROAD but
distinguished from the latter in welldeveloped platform and less numerous
denticles. Cnathodus commutatus (BRANSON & MEHL) and C. simplicatus RHODES,AUSTIN & DRUCE are similar to thepresent new species, but C. akiyoshiensis
has characteristic node-like denticle on
the carina in the center of platform.
Distribution :-Restricted in limestoneof 76002.
Reg. no. 1078, (Holotype).

193of the nodes on the inner side of theplatform and has short parapet-like
transverse ridges on the outer platform.
This form resembles MEISCHNER'S C.
bilineatus schmidti.
Distribution:- This species is widelydistributed in this area and was foundfrom limestones 76000, 76001, 76002, 76003
76004, 76005, 76006, 76007, 76009, 76010,
76011, 76012 and 76015.
Reg. nos. 1011, 1012, 1014.

Cnathodus commutatus(BRANSON & MEHL)
PI. 29, figs. 8-13

Cnathodus bilineatus (ROUNDY)
PI. 29, figs. 1-6
Polygnathus biZineatus ROUNDY, 1926, p. 13,
pI. 3, figs. lOa-c.
Gnathodus biZineatus, HASS, 1953, p. 79, pI.
14, figs. 25-29; REXROAD & FURNISII,
1964, p. 670; DUNN, 1965, p. 1148, pI. 140,
figs. 7-9; WIRTH, 1967, p. 205, pI. 19,
figs. 6-9; KOIKE, 1967, p. 296, pI. 1, figs.
9-11; GLOBENSKY, 1967, p. 440, pI. 58,
figs. 9, 13; RHODES, AUSTIK & DRUCE,
1969, p, 94, pI. 18, figs. 14-17; WEBSTER,
1969, p. 30, 31, pI. 5, figs. 11, 12; DUNN,
1970, p. 330, 331, pI. 62, figs. 13, 14;
MARKS & WENSINK, 1970, p. 258, pI. 2,
figs. 3a, b, 4.
Gnathodus biZineatus biZineatus, HIGGI:,\S &BOUCKAERT, 1968, p. 29, pI. 3, fig. 9.

Remarks :-Numerous specimens fromthe present collection show two appreciable morphologic types. The material
from the limestone below the reddish
tuffaceous shale is typical C. bilineatuscharacterized by well-developed nodes
on the inner platform. The transverse
ridges also conspicuous and developed
throughout the outer platform. However, the material yielded from thehigher levels decreases in the numbers

Spathognathodus commutatus BRAKSO:'\ &MEHL, 1941, p. 98, pI. 19, figs. 1-4.
Gnathodus commutatus, REXROAD & BURTON;
1961, p. 1153, pI. 39, figs. 1-3; HIGGINS& BOUCKAERT, 1968, p. 30, pI. 2, fig. 5;
RHODES, AUSTIN & DRUCE, 1969, p. 95,
pI. 19, figs. 9-12; WEBSTER, 1969, p. 31,
pI. 5, fig. 13.
Gnathodus commutatus commutatus, BISCHOFF,
1957, p. 23, pI. 4, figs. 2-6, 15; KOIKE,
1967, p. 296, pI. 1, figs. 12-16; WIRTH,
1967, p. 206, pI. 19, figs. 10, 11; DUNN,
1970, p. 331, pI. 62, figs. 11, 12; MARKS& WENSINK, 1970, p. 285, pI. 3, fig. la, b.
Gnathodus scotiaensis GLOBENSKY, 1967, p.
441, pI. 58, figs. 2-7, 10, 12.

Remarl,s :-A detailed synonymy was
given by RHODES, AUSTIN & DRUCE(1969, p. 95, 96). There are two different
morphologic types in this species. The
specimens collected from the limestone
below reddish tuffaceous shale havevery thin and small unit with sharply
pointed denticles and carina. However,the specimens from the higher levels
are characterized by thick and large
unit and with node-like carina.
Distribution:- This species was foundin limestones 76001, 76002, 76003 and76007.

Hisaharu lGO

194
Reg. nos. 1061, 1064.

Gnathodus homopunctatus ZIEGLER

on both sides of platform.
Distribution:-This species was foundin limestone 76002.
Reg. nos. 1036. 1037.

PI. 29, figs. 18-21
Gnathodus punctatus BISCHOFF, 1957, p. 24,
pI. 4, figs. 7-11, 14.
Gnathodus commutatus homopunctatus, HIG·
GINS, 1961, pI. 10, fig. 9; HIGGINS, 1962,
pI. 2, fig. 21; WIRTH, 1967, p. 206, pI. 19,
fig. 12; MARKS & WENSINK, 1970, p. 259,
pI. 3, fig. 2a, b.
Gnathodus homopunctatus, RHODES, AUSTIN& DRUCE, 1969, p. 103, pI. 19, figs. 5a-8d.

Remarks:-This material is similar to
Gnathodus nodosus, but the former differs from the latter in having platform
with sharply pointed end. G. nodOSllS
has comparatively short, roundly pointed
denticles, but G. homopunctatus shows
sharply pointed higher denticles. This
material shows a broad variation ofnodes and some of them fused together
and constitute parallel ridges on bothsides of platform.
Distribution:-Yielded from limestones
76001 and 76007.
Reg. nos. 1023, 1025, 1030.
Gnathodus nodosus BISCHOFF
PI. 29, figs. 14-17
Gnathodus commutatus nodosus BISCHOFF
1957, p. 23, 24, pI. 4, figs. 12, 13; HIGGINS,
1961, p. 213, pI. 10, figs. 7, 8; HIGGINS,
1962, pI. 2, fig. 19; KOIKE, 1967, p. 297,
pI. 1, fig. 19; WIRTH, 1967, p. 207, 208,
figs. 13-18; MARKS & WENSINK, 1970, p.
260, pI. 3, figs. 3, 4.
Gnathodus nodosus, RHODES, AUSTIN &DRUCE, 1969, p. 104, 105, pI. 19, figs. 16a20e.

Remarks:-This species closely resembles Gnatlwdus commutatus (BRANSON &MEHL). but the former has several nodes

Gnathodus aff. texanlls ROUNDY
PI. 29, fig. 7Aff.

Gnathodus texanus ROUNDY, 1926, p. 12,
pI. 2, figs. 7,8; HASS, 1953, p. 80, pI. 14,
figs. 16-21; BISCHOFF, 1957, p. 25, pI. 2,
figs. 21-25; THOMPSON & GOEI3ELL, 1963,
p; 12, fig. 3; REXROAD & COLLINSON,
1965, p. 8, pI. 1, figs. 33-38; THOMPSON& GOEBEL, 1969, p. 24, pI. 4, figs. 1-3, 5;
THOMPSON & FELLOWS, 1970, p. 89, pI.
2, figs. 15, 16; DUNN, 1970, p. 332, pI. 62,
fig. 21.

Remarks:- A detailed synonymy was
given by THOMPSON & FELLOWS (1970,
p. 89). This material closely resembles
Gnathodus texanus ROUNDY which hasvery narrow platform and its posterior
portion is long and graduallY tapered.
However, G. aff. texauns has a large
platform. Parapet is parallel to carina
on the inner side, and ridge-like ornamentation developed at about rightangle with carina. The present one
also resembles G. texanus pseudo semiglaber THOMPSON described from theupper Osagian Series.
Distribution:-This species is restricted in 76007.
Reg. no. 1021.
Genus Lonchodina ULRICH &BASSLER, 1926
Lonchodina furnishi REXROAD
PI. 29, fig. 33
Lonchodina furnishi REXROAD, 1958, p. 22,
pI. 4, figs. 11-13; HIGGINS, 1961, pI. 11,
fig. 8; RHODES, AUSTIN & DRUCE, 1969,

625.

Carboniferous conodonts from Akiyoshi

p. 141, 142, pI. 24, figs. 20a-23c; MARKS& WENSINK, 1970, p. 226, pI. 1, fig. 5.

Remarlls:- A detailed synonymy was
given by RHODES, AUSTIN & DRUCE(1969, p. 141, 142). No remarkable differences can be recognize<;l in thepresent specimens compared with manyother described ones.
Distribution:- Yielded from limestone
76001.
Reg. no. 1058.

195

Neoprioniodus montanaensis, RHODES, AUSTIN& DRUCE, 1969, p. 160, 161, pI. 22, figs.
5a-8b; MARKS & WENSINK, 1970, p. 226,
227, pI. 1, figs. 9, 10.

Remarks:- A detailed synonymy was
given by RHODES, AUSTIN & DRUCE(1969, p. 160, 161).
Distribution:- Recovered from limestones 76001, 76003 and 76007.
Reg. no. 1069.

Genus Ozarkodina BRANSON &Genus Neoprioniodus RHODES &MEHL, 1933

MULLER, 1956

Ozarkodina delicatula (STUFFER
Neoprioniodus scitulus (BRANSON& MEHL)& PLUMMER)
PI. 29, fig. 29

PI. 29, figs. 30, 31

Prioniodus scitulus BRANSON & MEIIL, 1941,
p. 173, pI. 5, figs. 5, 6.
Neoprioniodus scitulus, REXROAD, 1957, p.35,
pI. 2, figs. 22, 26; WEBSTER, 1969, p. 39,
40, pI. 7, fig. 13; RHODES, AUSTIN &DRUCE, 1969, p. 162, 163, pI. 22, figs. 9alOb, 12a, b.

Remarks:-A detailed synonymy was
given by WEBSTER (1969, p. 39, 40).
Distribution:- This species was foundfrom limestones 76001, 76004, 76011 and76015.
Reg. nos. 1067, 1068.
Neoprioniodus montanaensis (SCOTT)
PI. 29, fig. 32

Lochreia montanaensis SCOTT, 1942, p. 298,
299, pI. 39, fig. 9, pI. 40, fig. 12.
Prioniodus singularis HASS, 1953, p. 88, pI.
16, fig. 4.
Neoprioniodus singularis, HIGGINS, 1961, pI.
11, fig. 5; HIGGINS, 1962, pI. 1, fig. 8;
HIGGINS & BOUCKAERT, 1968, p. 45, pI.
1, fig. 8.

Bryantodus delicatula STUFFER & PLUMMER,
1932, p. 29, pI. 2, fig. 27.
Ozarkodina delicatula, BISCHOFF, 1957, p. 39,
pI. 1, figs. 25-28; HIGGINS, 1961, p. 220,
pI. 12, fig. 13; DUNN, 1965, p. 1149, pI.
140, fig. 18; GLOBENSKY, 1967, p. 446, pI.
56, fig. 19; HIGGINS & BOUCKAERT, 1968,
p. 45, pI. 13, fig. 3; RHODES, AUSTIN &DRUCE, 1969, p. 170, pI. 25, figs. 15, 16;
WEBSTER, 1969, p. 42, 43, pI. 7, fig. 11;
DUNN, 1970, p. 337, 338, pI. 62, fig. 31.

Remarks:-This material has numerous subequal, long and fused denticles.
The apical denticle is subequal in sizewith other denticles. The basal margin
is slightly arched. In aboral view thebasal cavity is a slit-like.
Distribution:- This species is abundant in limestones 76002, 76004, 76007
and 76010.
Reg. no. 1040.
Genus Spathognathodus BRANSON& MEHL, 1941

SPathognathodus campbelli REXROAD

Hisaharu lco

196
PI. 29,fig. 23

Spathognathodus campbelli REXROAD, 1957, p.
37, pI. 3, figs. 13-15; REXROAD, 1958, p.
25, pI. 6, fig. 9; REXROAD & BURTON,
1961, p. 1156, pI. 141, fig. 15; HIGGINS,
1962, pI. 2, fig. 20; REXROAD & FURNISH,
1964, p. 674, pI. 3, figs. 23,24; GLOBENSKY,
1967, p. 447, pI. 55, figs. 12, 20; KOIKE,
1967, p. 310, pI. 3, figs. 26-34; WIRTH,
1967, p. 233, 234, figs. 14-16; WEBSTER,
1969, p. 43, 44, pI. 7, fig. 5; DUNN, 1970,
p. 339, pI. 64, fig. 31; MARKS & WENSINK, 1970, p. 270, pI. 4, figs. 9, 10.
Spathognathodus d. campbelli, RHODES, AuSTI;\; & DRUCE, 1969, p. 223, pI. 8, figs.
1-4.

Remarks:- KOIKE (1967, p. 310) reported that this species has conspicuous
variation in the arrangement of denticles.
This Akiyoshi specimens has a similartendency. This species is related to
Gnathodus akiyoshiensis IGO n. sp., but
the latter is characterized by a welldeveloped elliptical platform.
Distribution:- This species was foundfrom limestones 76001,76002, 76006, 76007,
76009 and 76015.
Reg. no. 1051.
Spathognathodus cristulus
YOUNGQUIST & MILLER
PI. 29, fig. 22

Spathognathodus cristul a YOUNGQUIST &MILLER, 1949, p. 621, pI. 101, figs. 1-3;
GLOBENSKY, 1967, p. 447, pI. 57, figs. 15,
16.
Spathognathodus cristulus, RHODES, AUSTIN& DRUCE, 1969, p. 227, 228, pI. 8, figs.
14a-18d; DU"iN, 1970, p. 339, pI. 64, fig.
30.

Remarks:-A detailed synonymy was
given by RHODES, AUSTIN & DRUCE,(1969, p. 227, 228).
Distribution:- This species is yieldedfrom limestones 76004, 76005, 76006 and76010.
Reg. no. 1083.
References
BRA"iSON, E. B. and MEIIL, M. G. (1941):
Caney conodonts of Upper Mississippianage.
Denison Univ., Bull. Jour. Sci.,
Labs., vol. 35, p. 167-178, pI. 5.
BICSHOFF, G. (1957): Die Conodonten-Stratigraphie des rheno-herzynischen Unterkarbons mit Beriicksichtingung der
Wocklumeria-Stufe
und der
Devon/
Karbon-Grenze. Abh. Hess. Landesamt.
Bodenforschung, vol. 19, 64 p. 6 pis.
COLLINSON et ai., (1962): Six charts showing biostratigraphic zones and correlations based on conodonts from the
Devonian and Mississippian rocks of theUpper Mississippi Valley. Illinois Ceol.
Surv., Circ. 328, 32 p.
DRUCE, E. C. (1969): Devonian and Carboniferous conodonts from the Bonaparte
Gulf Basin, Northern Australia and theiruse in international correlation. Australia Bur. Mineral Res., Ceol. Ceophys.,Bull., vol. 98, 243 p., 5 pIs.
DeN", D. L. (1965):
Late Mississippian
conodonts from the Bird Spring Formation in Nevada. Jour. Paleont., vol. 39,
p. 1145-1150, pI. 140.
- - (1970): Middle Carboniferous conodontsfrom western United States and phylogeny of the platform group. Ibid., vol.
44, p. 312-342, pIs. 61-64.
ETo, J. (1967): A lithofacies analysis of
the lower portion of the Akiyoshi Limestone Group.
Akiyoshi-dai Sci. Mus.,
Bull., no. 4, p. 7-42, pis. 1-5, (in Japanese) .
GLOBENSKY, Y. (1967):
Middle and Upper
Mississippian conodonts from the Windscor Group on the Atlantic provinces ofCanada. Jour. Paleont. vol. 41, p. 432448, pIs. 55-58.
HASS, W. H. (1953): Conodonts of the Barnett Formation of Texas.
U. S. Ceo!.
Surv., Prof. Paper 243 F, p. 69-94, pIs.
14-16.

625.

Carboniferous conodonts from Akiyoshi

HIGGINS, A. C. (1961): Some Namurian conodonts from North Staffordshire. Ceol.
Mag., vol. 98, p. 210-224, pis. 10-12.
- - (1962): Conodonts from the "Griotte"
Limestone of northwest Spain. Espana
Inst. Ceol. y Mineral., Natas y Comun.,no. 65, p. 5-22, 3 pis.
- - and BOUCKAERT, J. (1968): Conodont
Stratigraphy and paleontology of theNamurian of Belgium.
Expl. Cartes.
Minieres, Belgium, Mem. no. 10, 64 p.,
6 pis.
IGo, Hisayoshi and KOIKE, T. (1964): Carboniferous conodonts from the ami Limestone, Niigata Prefecture, central Japan(Studies of Asian conodonts, Part 1).
Palaeont. Soc. Japan., Trans. Proc., N. S.
no. 53, p. 179-193, pis. 27, 28.
. - - and - - (1965): Carboniferous conodonts from Yobara, Akiyoshi Limestone,
Japan (Studies of Asiatic conodonts,
Part 2). Ibid., no. 59, p. 83-91, pis. 8,9.
.KOIKE, T. (1967): A Carboniferous succession of conodont faunas from the Atetsu
Limestone in southwest Japan (Studies
of Asiatic conodonts, Part 6). Tokyo
Kyoiku Daigaku, Sci. Repts., Sec. C, vol.
9, p. 279-318, pis. 1-4.
MARKS, P. and WENSINK, H. (1970): Conodonts and the age of the "Griote"
Limestone Formation in the upper
Aragon Valley (Huesca, Spain). Koninkl,
Nederl, Akad. Westensch. Amsterdam.,
Proc., Ser. B, 73, no. 3, p. 238-273. pis.
1-4.
MEISHNER, K. D. (1970): Conodonten-Chronologie des Deutschen Karbons. Compte
Rendu 6e Congres Intern. Starat. Ceol.
Carbonif., Sheffield 1967, vol. 3, p. 11691180.
.MINATO, M. and KATO, M. (1963): Fossils
with the lowest Namurian aspect newly
found by Dr. Y. HASEGAWA in the Akiyoshi province. Chikyu Kagaku, no. 66,
p. 32-42, pis. 1-2.
MURATA, M. (1961):
On the geologicalstructure of the Akiyoshi Plateau., Inst.
Ceol. Paleont. Tohoku Univ., Contr., no.
53, 46 p. (in Japanese).
,0KIlvIURA, Y. (1963):
Foraminiferal zones

197

underlying the Profusulinella beppensis
Zone of the Akiyoshi Limestone Group.
Hiroshima Univ., Ceol. Rep., no. 12, p.
305-318, pI. 39, (in Japanese).
- - (1965): Endothyroid foraminifera, Endothyranopsis from Japan. Ibid., no. 14,
p. 253-264, pI. 21.
- - (1966): Microbiostratigraphical studies
on the foraminiferal faunas of the Lower
Carboniferous formations of the Chugoku
region, Southwest Japan. Ibid., no. 15,
p. 1-46, pI. 1, (in Japanese).
aT A, M. (1968): The Akiyoshi LimestoneGroup; A geosynclinal organic reef
complex. Akiyoshi-dai Sci. Mus., Bull.,no. 5, p. 1-44, pis. 1-31.
OZAWA, Y. (1923): Stratigraphical study ofthe so-called Upper Chichibu Palaeozoic
system including the Akiyoshi Limestone
Ceol. Soc. Japan., Jour., vol. 30, no. 357,
p. 227-243. (in Japanese).
- - (1927): Palaeontological and Stratigraphical studies on the Permo-Carboniferous limestone of Nagato, part 2,
Palaeontology. Coll. Sci., Imp. Univ.
Tokyo, Jour., vol. 45, art. 6, p. 1-90,
pis. 1-14.
REXROAD, C. B. (1957): Conodonts from the
Chester Series in the type area of southwestern l11inois. Illinois Ceol. Surv.,
Rept. Inv. 199, 43 p., 4 pis.
- - (1958): Conodonts from the Glen Dean
Formation (Chester) of the l11inois Basin.
Ibid., 209, 27 p., 6 pis.
- - and BURTON, R. C. (1961): Conodontsfrom the Kinkaid Formation (Chester)in l11inois. Ibid., vol. 35, p 1143-1158,
pis. 138-141.
- - and COLLINSON, C. (1963): Conodontsfrom the St. Louis Formation (Valmeyeran Series) of l11inois, Indiana andMissouri. Ibid., Circ., 355, 28 p., 2 pis.
- - and - - (1965): Conodonts from the
Keokuk, Warsaw and Salem Formations(Mississippian)' of l11inois, Ibid., Circ.,
388, 26 p. pI. 1.
- - and FuRNISH, W. H. (1964): Conodontsfrom the Pella Formation (Mississippian), south-central Iowa. Jour. Paleont.,
vol. 38, p. 667-676, pI. 3.

198

Hisaharu lGORHODES, F. H. T., AUSTIN, R. L. and DRUCE,
E. C. (1969): British Avonian (Carboniferous) conodont fauna, and their value
in local and intercontinental correlation.
British Mus. (Nat. Hist.) , Bull., Geology,
suppl. 5, 313 p., 31 pis.
ROUNDY, P. V. (1926): The micro-fauna in
Mississippian formations of San Saba
County, Texas. U. S. Ceol. Surv., Prof.
Paper, 146, 63 p., 4 pis.
SCOTT, H. W. (1942) : Conodonts assemblages
from the Heath Formation, Montana.
Jour. Paleont, vol. 16, p. 293-301, pis. 3740.

STUFFER, C. R. and PLUMMER, H. J. (1932):
Texas Pennsylvanian conodonts andtheir stratigraphic relations.
U niv.
Texas Bull., 3201, p. 13-50, 4 pis.
THOMPSON, T. L. and GEOI3EL, E. D. (1963):
Preliminary report of conodont of the
Meramecian Stage (Upper Mississippian)from the subsurface of western Kansas.
Kansas Geo!. Surv., Bull., 165, 16 p. pI. l.
- - and - - (1969): Conodonts and stratigraphy of the Meramecian Stage (Upper
Mississippian) in Kansas. Ibid., 192, p.
1-56, pis. 1-5.
- - and FELLOWS, L. D. (1970) : Stratigraphy

Explanation of Plate 29
Figs. 1-6. Gnathodus bilineatus (ROUNDY).
1. Outer lateral view. 2, 5, 6. Oral views. 3, 4. Aboral views. x 40.
Fig. 7. Gnathodus aft. texanus ROUNDY.
Oral view. x 20
Figs. 8-13. Gnathodus commutatus (BRA!,;SO!,; & MEHL).
8, 11. Aboral views. 9, 12. Oral views. 10, 13. Lateral views. x 40.
Figs. 14-17. Gnathodus nodosus BISCHOFF.
14, 16. Oral views. 15. Lateral view. 17. Aboral view. x40.
Figs. 18-21. Gnathodus homopunctatus ZIEGLER.
18. Lateral view. 19-21. Oral views. x40.
Fig. 22. Spathognathodus cristulus YOUNGQUIST & MILLER.
Lateral view. x 60.
Fig. 23. Spathognathodus campbelli REXROAD.
Lateral view. x 60.
Figs. 24, 25. Gnathodus akiyoshiensis IGo, n. sp.
24. Lateral view. 25. Oral view. Holotype. x60.
Fig. 26. Cavusgnathus charactus REXROAD.
Inner lateral view. x 40.
Fig. 27. Ozarkodina sp.
Lateral view. x 60.
Fig. 28. Geniculatus claviger (ROUNDY).
Oral view. x 40.
Fig. 29. Ozarkodina delicatula (STUFFER & PLUMMER).
Lateral view. x 60.
Figs. 30, 31. Neoprioniodus scitulus (BRANSON & MEI-IL).
Lateral view. x 60.
Fig. 32. Neoprioniodus montanaensis (SCOTT).
Inner lateral view. x 60.
Fig. 33. Lonchodina furnishi REXROAD.
Lateral view. x 40.
Figs. 34, 35. Hindeodella spp.
Lateral views. x 60.

leo.'

Ca'rboniferous condonts fro'Y}t Akiyoshi

Plate 29

625.

Carboniferous conodonts from Akiyoshi

and conodont biostratigraphy of Kinder.
hookian and Osagean (Lower Mississipp·ian) rocks of southwestern Missouri
and the adjacent area. Missouri Ceol.
Surv. Water Resources, Rept. Inv. no. 45,
263 p. 8 pIs.
TORIY A,,!A, R. (1954): Geology of Akiyoshi.
Part 1. Study of the Akiyoshi LimestoneGroup. Fac. Sci., Kyushu Univ., Mem.,
Ser. D, vol. 4, p. 39-97.
(1958): Geology of Akiyoshi. Part 3.
Fusulinids of Akiyoshi. Ibid., vol. 7, p.
1-264, pIs. 1-48.
VOGES, A. (1959): Conodonten aus dem
Unterkarbon I und II (Cattendorfia- und
Pericyclus-Stufe) des Sauerlandes. Palliont. Zeitschr., Bd. 33, no. 4, p. 266-314,
pIs. 33-35.
WEBSTER, G. D. (1969):
Chester throughDerry conodonts and stratigraphy ofnorthern Clark and southern Lincoln
Counties, Nevada. California Univ. Pub.
Ceo!. Sci., vol. 79, 121 p., 8 pIs.
WIRTH, M. (1967): Zur Gliederung des hohern Paliiozikums (Givet-Namur) im
Gebiet des Quinto Real (Westpyrenaen)

Atetsu
Nagoe
Omi
Ota

f,iiJ

'ill

15

;~

..=1"-

I'J

lw

*-

lTI

199

mit Hilfe von Conodonten. Neues Jahrb.
Ceol. Pal. Abh., Bd. 127, p. 179-244, pIs.
19-23.
YANAGIDA, J. (1965):
Carboniferous brachiopods from Akiyoshi, Southwest Japan.
Part 2. Fac. Sci., Kyushu Univ., Mem.,
Ser. D. vol. 16, p. 113-142, pIs. 25-28.
- - et al. (1971): On the geology of thenortheastern part of the Akiyoshi plateau.
Dept. Ceol., Kyushu Univ., Sci.
Rept., vol. 11, no. 1, p. 105-114, pIs. 7-10.
- - and OTA, M. (1964): Geological age ofthe lower part of the Akiyoshi LimestoneGroup. (abst.) Ceol. Soc. Japan., Jour.,vol. 70, no. 826, p. 381.
YOUNGQUIST, W. L. and MILLER, A. K.
(1949): Conodonts from the Late Mississippian Pella beds of south-central
Iowa. Jour. Paleont. vol. 23, p. 617-622,
pI. 101.
All of the specimens treated in thispaper are preserved in the collection of the
Department of Earth Science, Tokyo Gakugei University.

Hirabaru
OhkuboOnimaru
Shishidedai

If!.

)JR

*- 0.. 1*

*
5~

}L

1:11

.L,

I:::)

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 92, pp'. 200-219, pIs. 30, 31, December 20, 1973

626.

FUSULINIDS OF THE NAGAIWA FORMATION*FUMIO KOBAYASHI

Institute of Geology and Mineralogy, Tokyo University of Education~;'fi~O)*,1jj!!p.fl::fil:' "?v'-C:~;1jj!!Rfl::fi:z. ~clj&i-.Qom;;:\;J:t!!O)~:'5~ J:: IJ t~~ U.:: 4 ~~:z.15"ts 9~, 21 ~O)*J\\iH:'i'irt'LJ:: IJ, Eostaffella ultragigantea, Pseudostaffella

antiqua, Eoschubertella sp. A, Pseudostaffella japonica, Parastaffella efr. vlerki 0)
5ill!*iI;~f~7JIJ~h, tJiT=1!fi'i Millerella ;}f\' I:. , f&='1!fl'i ProfusuZinella ::l~:I:.tjj~T.Qo tJ:.to~:'5l\!irf,~;'i/{~-t- ~ 7:/r$I:.,

J::lmli'i.:t::A:I ij17:/rff~;:.M}t~~1,.QoI]-

f* 51: J'(

been published. In the previous paper(KOBAYASHI, 1973), entitled "On the
Middle Carboniferous Nagaiwa Formation ", the writer reported the stratigraphy of the Nagaiwa Formation as
summarized in Text-fig. 2.
In thepresent article, the writer describes
twenty one species, including four new
speCies, from nine genera of fusulinid
fossils ranging from the lower Bashikirian to the lower Moscovian. The
following fusulinid zones and subzones
are established in decending order.

Introduction and acknowledgements
Middle Carboniferous Nagaiwa Formation, distributed in the Southern Kitakami Massif, Iwate Prefecture, Northeast
Japan, has been studied by many workers, such as ONUKI (1937), MINA TO et al.
(1953, 1959), YAMADA (1958, 1959) andothers. Although the Nagaiwa Formation is selected as a standard section of
the Japanese Middle Carboniferous, nocomprehensive paleontological study has

parastaffella efr. vlerki subzone
Zone of Profusulinella \ Pseudostaffella japonica subzone
Eoschubertella sp. A subzone
Pseudostaffella antiqua subzone{
Zone of Millerella
Eostaffella uitragigantea subzone

Detailed discussion of this biostratigraphical subdivision should be referred to the previous paper (KOBA Y ASHI, 1973).
The writer wishes to express his
sincere thanks to Professor Mosaburo
KANUMA, Tokyo Gakugei University
for his kind guidance and reading of

this manuscript. Cordial thanks are
due to Dr. Hisayoshi IGo, Tokyo University of Education for his critical andvaluable suggestions and also readingof this manuscript. The writer is indebted to Assistant Professor Atsushi
ISHII, Tokyo Gakugei University, Dr.
Toshio KOIKE, Tokyo University of
Education, Mr. Kozo WATANABE, Noda
Senior Highschool and Mr. Hisaharu IGO,

* Received Sept. 21, 1972; revised May 30,
1973; read Oct. 24, 1971 at Fukuoka.

200

626.

F1,~su linidsof the Nagaiwa Formation

~~ -

201

N

"

.JL
II
ADO
_-·36 0_._. __ .

Terrace

Deposits

Sa Sakamotozawa FormationiHIT Sakamotozawa F conglomerate
Ng Nagaiwa

Formation

IOn' Onimaru

Formation

Hi

Formation

Hikoroichi

Text -fig. 1. Topographic and g eneralized geo logical map ofNagaiwa area showin g t h e fossil localities_

202

Fumio KOBAY ASH!

Lower G,\,

Midd l e
Nag ai wa
Middle subf .

O n i ma _(~owermost subf.

F

ru(I Lower subforma tio n

. 10

150

~

r

.g~

subformot ion

320~iil~1

:

:

;

,

r r
o 0

,

:,

,

*

(Sakam o t oza wa F.

,

:

:

..

~Lower Perm.

F or ma tion
Uppe r120;,

Car bonifero us

rrrr

b

g g g-

r

~ ~ ~

~

~ g

,g

~~~~

.n

Eosi. ultrogi,,-a,~!ea
I:---.pss. anliqua

rLT
~

II1 I

V~rell~ SP

I
tI
Prof~su"~eJla, cfr

East sp B
Eosi. spp.
-J:. S l affefia sp .
Paraslaffefla (?) sp.

<:;.

I

I

I

mlxta

2'

'"-<=_.
C

Form atio n,Parasl . cfr . vlerki

Paraslaffefla

X Paras laffella

sp.A
sp.B

0
::J

~ chert~ scmdstone B s late
" pyroclastic rock 8
Iiliill

'"

-0

,, ~

a>

lava

Sak amo l ozaw a

(f)

c
-0

;;r

"

"a>

Ea limestone

Ozawolnella cfr brazhnikovae
O~awa'inella sp.A
Siaffefia spp .
Nankinella spp.

X rore::J

Naga iwo

-

;;
.a

~.

iOS fa:ffe/~a posfnagolwoensis

Pseudoslaffelfa (7) spo cammonloca l ity

nagaiwaensis£oslaffella cfr~

sec t ion

PseudoslalfellaI?) sp .

Mlllerefia spp.
E6slaffellaIabundant

1
1"J

Ozawamella spBI
daiyamensis,
I
I marbiensis
MllferellodEoschub erl,efl~ sp:A•~

i MII/erella
Profu~ulin~lIa sp.
Iblgemmicula

N1veflo sp.

Occurrence~ ~ColumnarPseudosloffello Joponica

I];...;1~

units

Thickness 1m)

Representativecollecting~

iLl,,,,,.:,l,'rnl i HI I ' *I I
.n !'I.n

Stra tigraphic

"a>

F.

E23 limestone

'"

C.

g-_.
'"_.
0
::J

~ conglomerate
Onl ma r u

For mation

E23~oslaffeffa

Zone

I

Pss', Easchuber- Pss. japonica
Paraslaffefla
flragiganlea antiqu o lelia sp.A
ofMiller.lla
Profusuline Jl a
Zone
of
I

cfr. vlerki

I

Fusulinine
Zonation

Text-fi g . 2. General ized stratigraphic section of the Nagaiwa Formation
showing the distribution of the fusulinacean species .

Tokyo Gakugei University for their encouragement and val uab le advice during
the course of this study.
All of the figured specimens are stored
in the Institute of Geology and Astro nomy, Tokyo Gakugei Un iversity.

Descript io n of Species

Gen us Eo staffella RAUSERCER NOUSSOV A, 1948

Eostaffe lla ult-ragigantea
KOB A YASH!,

n. sp.

P late 30, figs. 1- 5

Shell very large for the genus, dis-coidal in shape and with straight lateralside. Polar regions flat or w ith shallow
umbilica l depressions. Periphery bluntly
pointed to arched throughout the growth.
Shell coiled inv olute throughout, butrarely evolute in the last coiL Matureshells with 4 to 5 vo lutions.
Axial
length 0.118 to 0.298 mm and medianwidth 0.623 to 1.328 mm. Form ratio of
mature specimens 1: 0.11 to 1 : 0.22.
Shell expands rapidly from the third
volution. Proloculus minute, sphericaland external diameter 0.023 to 0.070 mm.
Spiroti1eca thin, composed of tectum,
upper and lower tectoria, but the las tv olution seems to have a trans lucent
layer between tectum and lower tectoriu m. Septa numerous, slightl y arched626.

Fusulinids of the Nagaiwa Formation

anteriorly, unfluted throughout shell.
Chomata weakly developed or lacking.
Tunnel low and path rather irregularly.
Remarks:-The present new species
somewhat resembles Eostaffella gigantea(KANMERA) and E. japonica (KANMERA),
but the former is easily distinguished203

from the latter two species by its larger
shell, smaller form ratio, and almost
straight lateral side.
Occurrence:-Abundant to common in
Loc. 19, rare in Loc. 21, associated with
Millerela marblensis THOMPSON, Millerella bigemmicula IGo, and M. spp.

Measurements of Eostaffella ultragigantea KOBAYASHI, n. sp. in mm.
Half length

No.
of
sp.

L.

1
2
3
4
5

0.145
0.298
0.118
0.143
0.130

W.

R.

0.648
1. 328
1. 053
O. 770O. 798

0.22
0.22
0.11
0.19
0.16

P.
0.053
0.070
0.028
0.023
0.033

1

2

3

4

.025
.035
.025
.025
.023

.025
.060
.630
.033
.023

.040
.068
.063
.063
.043

.073
.118
.093
.073
.063

5
-

.153
.115
-

Form ratio of vol.

Radius vector
1

2

3

4

.068
.078
.053
.063
.078

.103
.175
.098
.118
.150

.170
.270
.188
.218
.243

.323
.485
.348
.453
.400

5

1

2

3

4

-

.37
.45
.47
.40
.30

.24
.34
.31
.28
.17

.24
.25
.34
.29
.18

.23
.24
.27
.16
.16

.733
.548

-

Reg. Nos. :-2141-1 (Holotype), 2141-2-2141-5 (Paratypes).
Eostaffella nagaiwaensis KOBAYASHI,
n. sp.
Plate 30, figs. 9-13

Shell minute, subspherical to lenticular
in shape with convex lateral side andwith pointed to bluntly pointed periphery.
Polar regions with shallow
umbilical depressions. Shell involute
throughout the growth and almost
planispirally coiled. Mature shells have

5

.21
.21

-

4 to 5 volutions, 0.148 to 0.225 mm inaxial length and 0.270 to 0.368 mm inmedian width. Form ratios of mature
specimens 1: 0.44 to 1: 0.81. but about
1: 1 in inner volutions.
Heights ofvolutions gradually increase. Proloculus
minute, spherical. and external diameter
0.008 to 0.033 mm. Spirotheca thin,composed of tectum, lower and upper
tectoria.
Septa numerous. plane andunfluted. Chomata low. weakly developed or lacking. Tunnel low andpath rather regular.
Remarks:-The present new species
resembles closely Eostaffella rhomboides

Fumio KOBAY ASH!

204

Measurements of Eostaffella nagaiwaensis KOBAYASHI, n. sp. in mm.
No.
of
sp.

L.

Ralf length

1
2
3
4

0.225
0.220
O. 163
O. 148

I

I

W.

R.

0.298
0.273
0.368
0.270

P.

O. 76
0.81
0.44
0.55

0.008
0.010
0.033
0.028

Radius vector
1

2

3

4

.023 .050.023 .063! .018 .040
I
.038
!

.098
.103
.093
.075

.153
.150
. 103
.113

1

2

3

4

.020
.015
.018

.045
.048
.023
.023

.095
.090
.045
.053

.110
.108
.065
.075

5

.078

Tunnel angle (degrees)

Form ratio of vol.
5

1

2

3

4

5

1

25 25
31

-

-

-

-

-

-

.63
.66

.46

-

-

-

-

.-

-

.97
.87
.48
.71

-

-

.90
.76
.58
.61

.72
.72

.170

.87
.65
1. 00

-

-

-

25

-

-

-

RAUSER-CERNOUSSOVA in many respects,
such as form ratio, size of shell, andshape of periphery. However, it differsfrom the latter in poor development ofchomata, thin spirotheca, and large form
ratio of inner volutions of the present
new species. It is also similar to Eostaffella postnagaiwaensis KOBAYASHI, n.
sp .. but the latter has larger and smaller
form ratio than the former. E. nagaiwaensis also resembles Millerella bigemmintla IGo, but it differs in its involute
last volution and more inflated shell.
Occurrence :-Common in Loc. 39, rarein Locs. 25, 29, and 16; upper part of theZone of iV/illerella and lower part of theZone of Pro/usulinella.
Reg. nos. :-2095-2 (Holotype), 2095-1,
2095-3-2095-5 (Paratypes).
Eostaffella postnagaiwaensis

KOBA Y ASHI, n. sp.
Plate 30, figs. 17-20

Shell small, subdiscoidal to lenticular
in shape with convex or rarely concave

2

3

4

5
-

lateral side and broadly rounded periphery. Polar regions with umbilicaldepressions or flat.
Shell involute
throughout the growth and coiled almost
planispirally. Mature shells have 4 to 5 .
volutions and 0.193 to 0.258 mm in axial
length and 0.358 to 0.550 mm in medianwidth. Form ratios of mature specimens 1: 0.45 to 1: 0.71.
Proloculus
minute, spherical and external diameter
0.018 to 0.043 mm. Spirotheca thin, composed of tectum, lower and upper tectoria. Translucent layer between tectumand lower tectorium recognized in thelast volution of some specimens. Septa
numerous, slightly arched anteriorly,
unftuted throught shell. Chomata low
and poorly developed. Tunnel pathrather regular.
Remarks:- Most of the present specimens are deformed. Therefore detailed
comparison· is almost difficult. It resembles Eostaffella ampla (THOMPSON),but the former has more broadly rounded periphery and less dense chomata.
It also resembles Eostaffella infiecta(THOMPSON) and E. circuli (THOMPSON)

626.

Fusulinids of the Nagaiwa Formationin many respects, but the former haslarger form ratio, thinner and slightly
undulated spirotheca.
Occurrence:- Abundant in Loc. 16;

205

Zone of Profusulinella, Pseudostaffella
japonica subzone.
Reg. nos. :-2056-21 (Holotype), 205622-2056-24 (Paratypes).

Measurements of Eostaffella postnagaiwaensis KOBAYASHI, n. sp. in mm.
No.
of I
sp.

Half length

W.

L.

P.

R.

- -I

1
2 I

0.253
0.258
0.193
O. 193

I

3
4

0.358
0.550
O. 3~'8
0.428

O. 71
0.47
0.49
0.45

0.018
0.043
0.020

Radius vector
5.040 .063.025 .053.025 0.48.043 O. 75.120 . 198.088 . 160.090 .150.138 .213

.268
.203

3

3

4

5

.043
.038
.038
.040

.085
.063
.060
.063

.138
.103
.075
.OS8

.138
.098

I Tunnel angle (degrees)

Form ratio of vol.

4

2

.018
.025
.015
.025

2

2

3

.45

.68

1. 00
.60
.58

.72
.71

.71
.72

.53

.67
.46

4

1

5

.72
.64
.50
.46

.52
.48

2

3

4

5

27
25

34
19 20
25
23 24

~--~-~~~~~~~~-

Eostatfella cfr. mixta RAUSER-CERNOUSSOV A
Plate 30, figs. 8, 14-16
Cfr.

Eostat/ella mixta RAUSER-CERAcad. Nauk. Inst. Ceo!.
Sci. Minis!. Prosp., p. 59-60, pI. 1, figs.
34, 35.

1951.

NOUSSOVA,

Shell rather large for the genus, subdiscoidal in shape with straight toslightly convex lateral side, shallowly
umbilicated polar regions, and arched
to pointed periphery. Shell involute
throughout the growth and coiled planispirally. Mature shells have 4 to 5 volutions and 0.138 to 0.118 mm in axial
length, and 0.488 to 0.613 mm in medianwidth. Form ratio of mature specimens
1: 0.23 to 1: 0.31. Proloculus minute,spherical and external diameter 0.018 to

0.025 mm· Spirotheca thin, composed ofthree layers. Septa numerous, slightly
arched anteriorly.
Chomata massive,
low and indistinct in inner volutions.
Tunnel low and path rather regular.
Remarks: - The present specimens
differ from the Russian ones in form
ratio, shape of periphery and othercharacters.
However, the secondary
deformation possibly caused the above
mentioned differences. Thus, the writer
reserved the identification until morewell-preserved specimens accumulate.
Occurrence:- Abundant in Loc. 16 andrare in Locs. 36 and 38; lower part of
the Zone of Profusulinella.
Reg. nos. :-2056-17-2056-20
Eostaffella sp. A
Plate 30, fig. 31

Fumio KOBAY ASH!

206

Measurements of Eostaffella cfr. mixta RAUSER·CERNOUSSOVA in mm.
Half lengthNo.
. ' of
sp.

L.

R.

W.

0.138 0.598 0.23
O. 148 0.488 0.30
0.518 0.188 0.613 0.31
O. 138 0.513 0.27

1
2
3
4
5

P.

2

3

4

2

3

5

4

2

1

4

3

5

.038.065.103.175.295.033.068.100.168.265.038.078.155.258 .038.073.123.203.303.048 .095 .168.278 -Tunnel angle (degs.)
521.010 .013 .028 .050 .075
.013.023.048.050.075.013 .025 .050.075.093. 013 . 030 . 030 . 070 -

0.023
0.025
0.018
0.023
0.025

Form ratio of vol.
1

1

Radius vector

Septal count

3

4

5

2

-

-

-

-

-

-

-

-

-

-

1

3

4

.26 .20 .27 .29 .25.39 .34 .48 .30 .28

-

-

15

-

-

-

19
14

-

-

-

-

-

-

-

6?

10

11

15

.34 .34 .41 .37 .31.27 .32 .18 .25 -

-

-

15

-

-

-

-

-

'-

14

13
16

-

-

-

-

-

-

-

-

-

Remarks:-The present specimens arevery poorly preserved. Thus, the detailed comparison with other species isalmost impossible, but it is slightly
similar to Eostaffella cooperi (ZELLER)
in shape of shell, number of volutions
and arched periphery. E. cooperi is
originally described from the Kinkaid
Limestone, uppermost Chesterian ofU. S. A. However, this American species
differs from E. sp. A in larger proloculus and highly arched periphery.
Occurrence :-Rare in Loc. 17, enclosed
within the nucleus of oolite, lowermost
fossil bed of the Nagaiwa Formation.
Reg. no. :-2179-1
Eostaffella sp. B
Plate 30, figs. 6-7

Shell large for the genus, inflated
discoidal in shape with straight to convex lateral sides; polar regions slightly
depressed or sometimes flat umbili-

cally; periphery arched to highly
arched. Shell involute throughout thegrowth, and coiled planispirally. Matureshells have 5 volutions and 0.253 to 0.280
mm in axial length, 0.625 to 0.750 mm inmedian width. Form ratios of mature
specimens 1: 0.37 to 1: 0.41. Shell expands gradually.
Proloculus minute,spherical, and external diameter 0.025
to 0.045 mm. Spirotheca thin, composedof tectum, lower and upper tectoria.
Septa numerous, slightly arched anteriorly, unfluted and plane throughout
shell. Chomata massive, poorly developed. Tunnel low and path rather
regular.
Remarks:-The present species isslightly similar to Eostaffella gigantea(KANMERA), -but highly arched peripheryand large size of shell of the former
distinguish this species from the latter.
It also resembles Eostaffella ultragigantea KOBA Y ASH!, n. sp., but shape of
periphery, expansion and size of shell
are different.

626.

Fusulinids of the Nagaiwa Formation

Occurrence :-Common in Loc. 30, rarein Locs. 25 and 29: upper part of theZone of Millerella, Pseudostaffella antiqua

207

subzone.
Reg. nos. :-2254-9-2254-10

Measurements of Eostaffella sp. B in mm.

I- -I

Half length

No.
of
sp.
I

2

W.

L.

I

0.280
0.253

I

R.

O. 750
0.625

P.

0.37
0.41

Radius vector

0.045
0.025

.023
.015
I

Form ratio of vol.
5

1

2

3

4

5

.048.088.153.275 .423
.026.085.143.233.363

.48
.60

.40 .35.45 .42

.32
.36

.34
.36

1

2

3

4

Genus Millerella THOMPSON, 1942

2

3

4

5

.035
.038

.053
.060

.088
.083

. 143
. 130

Tunnel angle (degs.)
1
-

2

3

10
13

13
13

4

5

20

-

-

-

straight or convex lateral sides andshallowly depressed polar regions. Periphery arched to pointed. Mature shells
have 4 volutions, 0.078 to 0.103 mm inaxial length, and 0,488 to 0.623 mm inmedian width. Form ratios of themature specimens 1: 0.15 to 1: 0.17.
Shell expands rapidly. The first to
third volutions tend to involute withshallow umbilical depressions, but thelast one is evolute with umbilical depressions. Proloculus minute, sphericaland external diameter 0.028 to 0.040 mm.
Spirotheca thin, smooth and detailed
structures can not observed. Chomata
poorly developed.

Millerella marblensis THOMPSON
Plate 30, figs. 23, 24

1948. Millerella marblensis THOMPSON, Kansas Univ. Paleont. Contr., Protozoa Art
1, p. 76, pI. 23, figs. 1-12, pI. 24, figs.
1-9.
1957. Millerella cfr. marblensis THOMPSON:
IGo, Sci. Rep. Tokyo Kyoiku Daigaku,Sec. C, Vol. 5, Nos. 47, 48, p. 26, 27,
pI. 1, figs. 13, 14, 18, 19.
1964. Millerella marblensis TlImlPsox: MoORE, Jour. Pal., Vol. 38, No.2, p. 294305, pI. 47, figs. 1-24, pI. 48, figs. 1-23.

Shell small, discoidal in shape, with
Measurements of Millerella marblensis THOMPSON III mm.
No.
of
sp.

Radius vector

Half length

L.

W.

R.

P.

0.1030.623 0.17 0.028
1
2 10.0780.4880.160.040
I
3 i 0.0830.550 0.150.028

1

2

3

4

.013.025 .048.048.023.025 .048 .048
.015.025.040.040

1

2

3

Form ratio of vol.
4.053.103.183.338.050.113.188.273.048.088.168.298

1

2

3

4

. 25 . 24 . 26 . 14
.46 .22 . 26 . 18
.31 .28 .24 . 13

Fumio KOBAYASHI

208

10-13.

Remarks:- The present species closely
resembles THOMPSON's typical specimensfrom the Marble Falls Limestone, Texas.
Compared with MOORE'S one from the
Big Saline Formation, Texas, the present
specimens have not so evolute shell.
Occurrence :-Common to rare throughout the Nagaiwa Formation.
Reg. nos. :-2195-1, 2056-25

Shell small, discoidal in shape, withconvex lateral sides, narrowly roundedor acutely pointed periphery and somewhat deeply umbilical depressions.
Mature shells have 4 to 4.5 volutions,
0.098 to 0.178 mm in axial length, and0.263 to 0.450 mm in median width. Formratios of mature specimens 1: 0.24 to
1: 0.57. Shell coiled planispirally throughout the growth. Inner volutions involute with slight umbilical depressions,but the outer ones evolute or partly
evolute. Proloculus minute, sphericaland external diameter 0.018 to 0.048 mm.
Spirotheca thin, composed of tectum,
lower and upper tectoria.
Chomata
massive, asymmetrical and weakly developed.

Millerella bigemmicula IGO
Plate 30, figs. 21, 22

1957. Millerella bigemmicula IGo, Sci. Rep.
Tokyo Kyoiku Daigaku, Sec. C, Vol. 5,
Nos. 47-48, p. 172-174, pI. 1, figs. 1-9,
15-17, 27.
1964. Millerella bigemmicula IGo: SADA,
Jour. Sci. Hiroshima Univ., Sec. C, Vol.
4, No.3, p. 228-229, pI. 21, figs. 5, 6,

Measurements of Millerella bigemmicula IGO in mm.
No.
of
sp.

0.178
0.098
O. 108
O. 150

1
2
3
4

Half length

w.

L.

P.

R.

0.413
0.273
0.450
0.263

0.42
0.34
0.24
0.57

0.025
0.020
0.048
0.018

.018
.010
.025
.030

2

3

4

4.5

.023
.025
.038
.050

.038
.040
.055
.075

.073
.050
.055
.075

.075

I

Radius vector
1

2

3

4

Form ratio of vol.
4.5

.023.048.075.143.218.023 .053 .103.150 .048.083.148.255 .033.058.113 .148 -

I

1

2

.78
.44
.52
.91

.48 .51.47 .39.46 .38.86 .66

Remarks :-It differs slightly from thetypical species from the Ichinotani
Formation in expansion of shell, diameter of proloculus, development ofchomata and form ratio.
Occurrence :-Common to rare throu-

3

4

4.5

.51 .35.33 .22 .51 -I

Tunnel angle (degs.)

I

1

2

3

4

4.5

-

-

22

-

-

-

-

-

-

-

-

17

-

-

-

-

-

-

-

-

ghout the Zone of Millerella and thelower part of the Zone of Profllslllinella.
Reg. nos. :-2111-1, 2096-6.

Genus Ozawainella THOMPSON, 1935

626.

Fusulinids of the Nagaiwa Formation

Ozawainella cfr. brazhnikovae GINKEL

209

Axial length 0.213 to 0.298 mm, medianwidth 0.843 to 0.963 mm, and form ratio
1: 0.25 to 1: 0.31. Mature shells have
4.5 to 5 volutions. Proloculus sphericaland external diameter 0.043 to 0.048 mm.
Spirotheca composed of dense tectum
and thin lower and upper tectoria inouter volutions. This differentiation of
spirotheca in inner volutions obscure.
Chomata weakly developed. or extends
to the poles in some specimens, andpath straight.

Plate 30, figs. 25, 26
Cfr. 1965. Ozawainella brazhnilwvae GINKEL,
Leidse Ceol. Med., Deel. 34, p. 62, 63,
pI. 15, figs. 61, 62.

Shell small, rhomboidal to lenticular
in shape, with convex lateral sides.
Polar regions flat or with broad andshallow umbilical depressions; peripheryarched in the first to second volutions.
angular to bluntly angular in outer ones.

Measurements of Ozawainella cfr. brazhnikovae GINKEL in mm.
No.
of
sp.

L.

1
2

0.213
0.298

Half length

W.

R.

0.843
0.963

0.25
0.31

Radius vector
1

2

3

4

P.
.025
.023

0.043
0.048

.083.115.153.273.430.073.123 .198.308.478

1

I

2

3

.30 .44 .41.32 .19 .29

Remarks :-The present species are
identified with GINKEL's one with somereservation. Detailed specific identification is postponed until more well-preserved specimens accumulate.
Occurrence:- Rather common in Loc.
16; Zone of Profusulinella, Pseudostaffella japonica subzone.
Reg. nos. :-2056-26, 2056-27.
Ozawainella sp. A
Plate 30, fig. 27

Shell small, lenticular to rhomboidal
in shape. Polar regions flat or withslight umbilical depressions; periphery

4

3

4

5

.050
.023

.063
.058

.078
.098

.108
.163

Tunnel angle (degs.)

Form ratio of vol.
5

2

5

.29 .25.32 .34

-

1

2

3

9

9
10

14
10

4

5

-

-

-

-

arched in 1-1.5 volutions, but bluntly
angular in outer ones. Mature shells
have 4 to 4.5 volutions. Axial length
0.431 mm, median width 1.963 mm, andform ratio 1: 0.21. Half length of thefirst to fourth volutions 0.038, 0.058, 0.103,
and 0.138mm. respectively: radius vector
0.075, 0.133, 0.253 and 0.458 mm, respectively. Proloculus small. spherical andexternal diameter 0.075 mm. Spirothecacomposed of tectum. lower and upper
thick layers, but secondary mineralization obliterates most of their original
structure.
Chomata indistinct, andweakly developed.
Remarlls:- This species is distinguished from Ozawainella cfr. brazhni-

Fumio KOBAY ASH!

210

width, giving form ratio 1: 0.34. Detailed
observation is difficult, but the occurrence of this unidentified species isnoteworthy, because this species seems
to be the uppermost representative of
the Zone of Profusulinella of the Nagaiwa Formation.
Occurrence:- Very rare in Loc. 47;
Zone of Profusulinella, Parastaffella cfr.
vlerl,i subzone.
Reg. no. :-2238-1.

kovae from Nagaiwa, in larger shell,
weakly developed chomata, and spirothecal structure.
Occurrence :-Common in Loc. 16;
Zone of Profusulinella, Pselldostaffella
japonica subzone.
Reg. no. :-2056-28.
Ozawainella sp. B
Plate 30, fig. 39

Remarks :-Only one poorly preserved
specimen was obtained. It has 0.263 mm
in axial length and 0.775 mm in median

Genus Pseudostaffella THOMPSON, 1942
Pseudostaffella antiqua (DUTKEVICH)

Measurements of Pseudostaffella antiqua (DUTKEVICH) in mm.
No.
of
sp.
--

R.

--

0.825
0.625
0.900
0.900
0.813
0.675
O. 700
0.498
0.525
0.500

0.59
0.80
0.63
0.64
0.65
0.67
0.64
1. 23
---

Form ratio of vol.
1

P.
2

1
0.488
0.500
0.563
0.575
0.525
0.450
0.450
0.613
--

I

2
3
4
5
6
7
8
9
10

Radius vector

Half length
W.

L.

2

3

0.035
0.055
0.030
0.038
0.035
0.055
0.063
0.050
0.048
0.060

I

--

--

--

--

--

--

--

--

--

--

1

2

3

.62 .63 .57.75 .78 .75.58 .57 .58.54 .54 --- .78 .69 .68-.62 .78 .79 .87 -.23 .79 .76 .61 -.46 .30 . 48 1. 07 1. 32

--

--

13
18
22
22

--

--

20
25
27
20
26

---

2

3

4

5

.035.105.165.260.395.060.085.133.205.338.060.105.195.310.475.060.140.238.350 ?
.050.113.178.303.433.065.103.190.260 ?
.065.095 .143 .248.385.050.093.145 .185.200.050.085.170.208 -.045 .113 .200 -- --Septal countTunnel angle (degs.)
1.37 -.42 .62-- .71.83 .57

5.013 -- .103 .163 .223
.025 .053 .100.160.255-- .075 .113 .175 .275
.050.080.128.190.288--- .138.210 .295
.040.080.150.225 -.015.075.108.150 -.023.028.070.198.263

5

4

4

3

4

5

1

2

3

4

5

25

--

--

6

--

--

--

--

--

--

--

--

--

--

--

--

--

35

--

5

--

--

--

--

--

--

--

--

26

--

?

10

--

--

--

--

18

--

--

--

--

--

--

--

--

--

--

27
55

--

--

--

--

--

--

--

--

21
69

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

--

14
?

--

--

11
12

11

--

8
3?

I

16

--

626.

Fus~dinidsof the Nagaiwa Formation

Plate 31, figs. 1-6
1951. Pseudostaffella antiqua

CDUTKEVICII):

Acad. Nauk.
Inst. Ceol. Sci. Minist. Petrol. Prosp.,
p. 97, 98, pI. 5, figs. 6-8.
1965. Pseudostaffella antiqua CQuTKEvrcr·r):
GI~KEL, Leidse Ceol. Med., Deel. 34, p.
69, 70, pI. 16, figs. 22-25.
RAUSER-CERNOUSSOVA,

Shell small, spherical to subsphericalin shape. Mature shells have 4 to 5
volutions. Axial length 0.450 to 0.613
mm, median width 0.498 to 0.900 mm,
and form ratio 1: 0.59 to 1: 1.23. Inner
first or second volution closely coiled
and commonly at right angle to the axis.
Spirotheca thin, composed of threelayered structures. Proloculus sphericaland external diameter 0.030 to 0.063 mm.
Septa plane and unfluted. Chomata low
and massive. Tunnel low and pathrather regular.
Remarks:- This species is reported
from the Russian Platform, Donetz
Basin, Cantabrian Mountains, Omi andAkiyoshi Limestones of Japan. The
present specimens from Nagaiwa areslightly larger than the specimens fromthe above mentioned localities, but othercharacters are closely similar to these
specimens.
Occurrence:- Abundant in Loc. 30 andrare in Locs. 25 and 29; Zone of Millerella, Pseudostaffella antiqua subzone.
Reg. nos. :-2254-1-2254-6

Pseudostaffella japonica

KOBAYASHI, n. sp.
Plate 31, figs. 7-16

Shell spherical to subspherical inshape. Mature shells have 5.5 to 6, rarely 7, volutions. Axial length 0.703 to

211

1.038mm, median width 0.733 to 1.365mm,
and form ratio 1: 0.51 to 1: 1.19. Inner
first to second, occassionally third, volutions with short axis of coiling, andcoiled commonly at right angle or very
irregularly to the axis. Spirotheca thin,composed of three layered profusulinellid
wall. Proloculus spherical and external
diameter 0.045 to 0.075 mm. Septa numerous, plane and un fluted. Chomata
low and massive. Tunnel path rather
irregular.
Remarks:- Although the present specimens are deformed, this resembles
Pseudostaffella gorskyi (DUTKEVICH) andP. nibelensis RAUSER-CERNOUSSOVA in
many respects, except the weaker development of chomata in the former.
Also this species resembles Pseudostaffella kanumai IGo, but the former haslarger shell and more irregular coilingof the inner volutions than the latter.
Occurrence:- Abundant in Loc. 16 andrare in Loc. 39; Zone of Profusulinella,
Pseudostaffella japonica subzone.
Reg. nos. :-2056-1 (Holotype), 2056-22056-10 (Paratypes).

Pseudostaffella (?) sp.
Plate 30, figs. 28-30

Shell spherical to subspherical inshape, with slightly shallow umbilicaldepressions. Mature shells have 4 volutions. Axial length 0.168 to 0.413 mm,median width 0.208 to 0.425 mm, andform ratio 1: 0.40 to 1: 1.99. Inner firstto second volutions coiled very irregularly or at right angle to the axis.
Spirotheca very thin, composed of dense
layer and thin less dense layer. Proloculus minute, spherical and external
diameter 0.025 to 0.038 mm. Chomata
low and weakly developed or lacking.

I:-'......
t-:)

Measurements of Pseudostaffella japonica KOBA YASm, n. sp. in mm.
No.
of
sp.

Half length

L.

W.

2

4

5

.045 .065 .118 .188.025 .050 .108 .198.035 .063 .160 .263
.093 .150 .275
.050 .108 .188
.038 .068 .108 .193
.030 .050 .098 .198.030
.123 .208
-

.275
.310
.393
.390
.253
.325
.340
.315

1

3

Radius vector

I

P.

R.

6

7

3

4

5

.073 .130 .198
.063 .100 .143
.075 .123 .175.088 .163.088 .150 .248
.053 .098 .125.048 .063 .110.050 .080 .140.095 . 160 .263
.060 .115 .175

.300
.213
.290
.238
.380
.173
.178
.218
.408
.278

.423
.340
.500
.350
.625
.273
.303
.347
.575
.420

1

2

6

,

-O. 703
0.988
0.878
1. 038
O. 703
0.873
0.880
0.825

1
2
3
4
5
6
7
8
9
10

1. 0630.66
1. 243 - 0.80
1. 258 O. 70
1. 025 1. 03
1. 365 0.51
O. 733 1. 19
0.803 1. 10
0.938 0.88
1. 053
1.118
-

-

0.045
0.050
0.070
-

0.073
0.055
0.053
O. 053
0.075

-

-

-

-

-

-

--

-

-

.365
.428 .493
.428
.513
.350
.443
.438
.393
-

-

-

-

-

-

-

-

-

- - - - -

-

-

--

7

.568'
.475 .642
.673
.520
.728
.398
.423
.503
-

-

.593
------

-

~

...
;§l
o

~

----

ttl

;l:..

Form ratio of vol.

1

2

.62
.40
.47

3

4

5

Tunnel angle (degrees)

6"<:

Septal count

;l:..

1

2

3

4

5

-

-

-

-

-

-

-

-

-

-

-

9?

-

-

-

-

-

-

13

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

17
15

27
28
37
55

-

-

-

28
23
30
31
16
24
23
39

38
45

-

16
22
14
28
9
10
21
26

-

-

15
15
12
25
9

-

.77

-

-

-

-

-

-

-

-

7

.50
.50
.51
1. 06
.33
.72 ..69.63 .79.60-.60 .63 .65 .64.76 .93 .91.90.91 .91 .79 .64.92 1. 16 1.11
.99
.44 .50 .48 .48.86 1. 12 1. 19 1.11
.89 1.11 1. 12 1. 04.88 .95 .91
.78

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

6

1

7

8

2

-

3

4

5

6

7

-

--

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

13
15

20
17

24
20

3+
15+

-

-

10
12
12

-

-

8
8

-

~

626.

F1ts1tlinids of the Nagaiwa Formation

Tunnel low and path rather irregular.
Remarks:- The present specimens are
so deformed that the original shape of'shell is obscure. Several important bio-characters are somewhat related to Eos.chubertella. Generic designation is ten-213

tative.
Occurrence :-Common in Locs. 36 and38; Zone of Profusulinella, Eoschubertella
sp. A subzone.
Reg. nos. :-2088-5-2088-6, 2093-1

Measurements of Pseudostaffella (?) sp. in mm.
No.
of
sp.

L.

Half length

1
2
3
4

O. 1900.413
O. 168
0.173

W.

R.

0.423
0.208
0.425
0.343

Radius vector

P.

0.45
1. 99
0.40
0.50

0.030
0.038

1

2

3

4

.030
.018

.063
.088
.038
.048

.075
.148
.058
.070

.098
.198
.088
.103

0.025

Septal countForm ratio of vol.

1

2

3

4

1

2

3

.053
.063
.050
.063

.100
.038
.090
.088

.158
.073
.148
.148

.235
.108
.225
.203

.51
.29

.63
2.32
.42
.55

.48
·2.03
.39
.47

-

Genus Profusulinella RAUSER-CERNOUSSOVA and BELJAEW, 1934
Profusulinella cfr. daiyamensis

HASEGAWA
Plate 31, fig. 17
oCfr. 1967. Profusulinella daiyamensis HASEGAWA, Sci. Rep. Niigata Univ. Ser. E,
No.1, p. 9-15, pI. 1-3.

4

2

1

3

4

.41

-

-

-

-

1. 83

-

-

-

-

.39
.51

-

-

-

-

-

-

-

9

be comparable with HASEGAWA's species
from the Akiyoshi Limestone, but theconclusion is kept open.
Occurrence :-Common to rare in Loc.
39; Zone of Profusulinella, lower partof Pseudostaffella japonica subzone.
Reg. no. :-2095-7.
Profusulinella sp.
Plate 31, figs. 18-19

Shell small, slightly inflated fusiform
in shape. Mature shells have 5 volutions. Axial length 1.033mm and medianwidth 0.413 mm. Inner structures can
not be clarified by ill-preservation andpoorly oriented sections.
Remarks :-Slightly inflated fusiform'shell, concave lateral sides and otherfeatures of the present specimens may

Two well-oriented specimens obtained.
Shell inflated fusiform to elongate fusiform in shape. Mature shells have 5
volutions. Axial length 2.275mm, medianwidth 1.275 mm, and form ratio 1: 1.78.
Half length of the first to fifth volutions
0.103,0.230,0.450, 0.838 and 1.175 mm, respectively. Radius vector 0.053, 0.105,

Fumio KOBAYASHI

214

0.220, 0.465 and 0.740 mm, respectively.
Inner 2 or 3 voluti6ns closely coiled andthe subsequent ones gradually increasein height and width. Proloculus minuteand external diameter 0.055 mm .. Spiro·
theca composed of tectum, lower andupper tectoria. Chomata low and massive. Tunnel angle of the second to
fourth volutions 17, 28, and 47 in degrees, respectively. Tunnel path irregular.
Remarks:- The previous students listed Profusulinella sp. from the Nagaiwa
Formation. Profusulinella sp. describedherein seems to differ from the species
by other authors, because the writer's
specimens are rare in occurrence, andsome of the deformed specimens ofPseudostaffella japonica resembles Profusulinella. Furthermore, the writer'sPseudostaffella japonica is particularly
abundant. Therefore, Profusulinella sp.
listed by the previous authors seems tobe the writer's Pseudostaffella japonica.
Occurrence:- Very rare in Loc. 16;
Zone of Profusulinella, Pseudostaffella
japonica subzone.
Reg. nos. :-2056-29, 2056-30
Genus Verella DALMA TSKA Y A, 1952
Verella sp.
Plate 31, figs. 20-25

Shell elongate fusiform in shape, composed of 3.5 to 4 volutions in mature
specimens. Axial length 1.293 to 2.450
mm, median width 0.353 to 0.558 mm, andform ratio 1: 3.40 to 1: 4.39. Heights ofvolutions increase gradually. The firstto second volutions elongate rhomboidal
and subsequent ones elongate fusiform
or elongate subcylindrical. Proloculus
spherical and external diameter 0.075 to
0.150 mm. Septa weakly fluted in polar
regions. Chomata poorly developed or

Measurements of Verella sp. in mm.
Half length

No.
of
sp.

W.

L.

1
2
3
4
5
6

1. 788

R.

0.488
0.365
0.558
0.405
0.353
0.428

1. 2932.450
1. 558
1. 3001. 775

3.66
3.40
4.39
3.85
3.68
4. 15

Radius vector

i

P.
0.080
0.095
0.075
O. 103
O. 105
O. 150

.130
.148
.100
.100
.125
.205

2

3

4

1

2

3

.050
.063
.050
.050
.060
.058

.098
.125
.080
.088
.115
. 125

.153
.208
.125
.150
.195
.228

.275

2.60
2.35
2.00
2.00
2.08
3.53

3.27
2.50
2.50
3.44
2.50
3.02

3. 76
2.88
3.22
4.17
3.27
3.29

-

.238
.238
-

3

.320
.313
.200
.303
.288
.378

.575
.598
.403
.623
.638
.750

4

.738

Tunnel angle (degs)

Form ratio of vol.

1

2

4

1

2

4

3

-

17

40

-

-

-

-

-

-

3. 10

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

626.

Fusulinids of the Nagai'Wa Formation

lacking. Dense secondary filling obser·
ved in axial regions. Spirotheca thin,probably composed of three layers.
Remarks;- This is the first record of
the occurrence of this genus in Japan.
Comparing with the Russian species andVerella sp. from the Cantabrian Mountains, Spain, the present species is characterized by less number of volutions,
smaller size of shell and weaker septal
fluting. Based on the mentioned dif·ferences, the Nagaiwa species may bebiologically more primitive than thementioned foreign materials.
Occurrence;- Rare in Loc. 16; asso·
ciated with Profusulinella sp., Pseudostaffe.lla japonica, Eostaffella postnagaiwaensis and others; Zone of Profusulinella, Pseudostaffella japonica subzone.
Reg. nos. :-2056-11"'2056-16

with straight axis of coiling. Matureshells have 4 volutions. Axial length
0.358 to 0.523 mm, median width 0.203 to
0.300 mm, and form ratio 1: 1.74 to 1:
1.76. Inner one to one and half volutions irregularly coiled to the axis, subsequent ones regularly coiled. Proloculus spherical, rather large for size ofshell and external diameter 0.038 to 0.050
mm. Spirotheca thin, composed of tectum and a lower dense layer. Chomata
massive, and rather symmetrical. Septa
not fluted and plane.
R emar ks ;-This species resem bles
closely Eoschubertella obscura (LEE andCHEN) in many respects, such as shapeand coiling of shell and rather large
proloculus, but this can be distinguishedfrom the latter in number of volutions
and development of chomata. Also thisspecies resembles Eoschubertella lata
(LEE and CHEN) and E. elliptica (SHENG),
but differing from the latters in having
subspherical shape and irregular coiling.
of the inner volutions.
Occurrence ;-Common in Locs. 36 and38; lower part of the Zone of Profusu-

Genus Eoschubertella THOMPSON, 1937
Eoschubertella sp. A
Plate 31, figs. 28-31

Shell minute, subspherical

shape

III

215

Measurements of Eoschubertella sp. A in mm.
No.
of
sp.

Half length

w.

L.

2

4

1

2

.070 .090 .130 .213
.075 .125 .175 .253
.053 .108 .175

.030
.048
.048
.050

.063
.075
.075
.093

1
0.448 0.258 1. 74 0.048
0.523 0.300 1. 74 0.043
0.358 0.203 1. 76 0.050
0.298
0.038

1
2
3
4

Radius vector

P.

R.

Form ratio of vol.

3

Tunnel angle (degs.)

1

2

3

4

2.33
1. 56
1.10

1. 43
1. 67
1. 44

1. 33
1. 55
1. 67

1. 31
1. 45

-

-

-

1

3

4

.098 .163.113 .175
.105
.158

Septal count
1

2

3

2

3

42
32

44
60

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

7?

14

16

2+

4

4

Fumio KOBAY ASH!

216

finella, Eoschubertella sp. A subzone.
Reg. nos. :-2088-2",2088-4, 2093-2

Genus Parastaffella RAUSERCERNOUSSOVA, 1948
Parastaffella cfr. vlerki GINKEL
Plate 30, figs. 34-38
Cfr. 1965. Parastaffella vlerki GINKEL, Leidse
Ceol. Med., Deel. 34, p. 19, 20, pI. 9,
figs. 7-18, pI. 10, figs. 1-35.

Shell lenticular with convex lateralsides; polar regions having flat or shallow umbilical depressions; peripherybluntly pointed to arched in inner volutions, broadly rounded in outer ones.
Shell involute in inner one to three andhalf volutions, evolute or occasionallyinvolute in subsequnt ones. Matureshells have 4.5 to 5 volutions. Axial
length 0.203 to 0.345 mm, median width
0.833 to 0.925 mm and form ratio 1: 0.22
to 1: 0.41. Chomata massive. low andsometimes extends to poles. Septa unfluted, numerous and inclined anteriorly.

Explanation of Plate 30
Figs. 1-5. Eostaffella ultragigantea KOBAYASHI, n. sp.
1. Axial section of the holotype; 2-4. Axial sections of para types; 5. Sagittal sectionof paratype, x 60; Loc. 20.
Figs. 6-7. Eostaffella sp. B
6-7. Axial sections, x 60; Loc. 13.
Figs. 9-13. Eostaffella nagaiwaensis KOBAYASHI, n. sp.
10. Axial section of the holotype; 9, 11-13. Axial sections of paratypes, x 60; Loc. 39.
Figs. 8, 14-16. Eostaffella cfr. mixta RAUSER-CERNOUSSOVA
8, 14-15. Axial sections; 16. Sagittal section, x 60; Loc. 16.
Figs. 17-20. Eostaffella postnagaiwaensis KOBAY ASH!, n. sp.
17. Axial section of the holotype; 18-20. Axial sections of paratypes, x 60; Loc. 16.
Figs. 21-22. Millerella bigemmicula IGO
21-22. Axial sections, x 60; Locs. 2 and 39.
Figs. 23-24. Millerella marblensis THOMPSON
23-24. Axial sections, x 60; Locs. 16 and 28.
Figs. 25-26. Ozawainella cfr. brazhnikovae GINKEL
25-26. Axial sections, x 40; Loc. 16.
Fig. 27. Ozawainella sp. A
Axial section, x 40; Loc. 16.
Figs. 28-30. Pseudostaffella (?) sp.
28-30. Axial sections, X 40; Locs. 36 and 38.
Fig. 31. Eostaffella sp. A
Tangential section, x 60; Loc. 18.
Figs. 32-33. Parastaffella sp. B
32. Sagittal section; 33. Axial section, x 60; Loc. 45.
Figs. 34-38. Parastaffella cfr. vlerki GINKEL
34-35. Axial sections; 36-37. Tangential sections ; 38. Sagittal section, x 40; Loc. 45.
Fig. 39. Ozawainella sp. B
Axial section, x 60; Loc. 47.
Figs. 40-41. Parastaffella sp. A
40. Axial section, x 40; Loc. 45. 41. Tangential section, x 60; Loc. 40.

KOBAYASHI:

Fusulinids of the Nagaiwa Formation

Plate 30

626.

Fusulinids of the Nagaiwa Formation

Spirotheca composed of very distinct
dense layer and lower and upper lessdense layers. Diaphanotheca indistinct.
Remarks :-GINKEL recognized broad
variations in this species, such as measurements of radius vector, form ratio,
angularity of periphery and height ofchomata. The present material is very217

similar to the original ones from the
Cantabrian Mountains and falls within
the limits of variation of the typespecies, except the spirothecal structure.
Occurrence: - Common in Loc. 45;
Upper part of the Zone of Profusulinella,Parastaffella cfr. vlerki subzone.
Reg. nos. :-2232-1-2232-5

Measurements of Parastaffella cfr. vlerki GINKEL in mm.
No.
of I
L.
sp. i
,
1
2
0.345
0.203
3

Half length

W.

R.

0.918
0.833
0.925

-

0.41
0.22

I

!

i

2

-

-

.56 .54.36 .34

0.043
0.045
0.028

I

Form ratio of vol.
1

Radius vector

P.

3

4

-

-

-

.51 .45.30 .37

.43
.23

1

2

3

4

5

. 1

-

-

-

-

-

.055.123 .220 .363 .460
.045.093 .175.298.450.050 .098 .158.278.450.025.050.090.133 .193
.018.033.048.103.103

Tunnel angle (degs.)

2

3

4

5

Septal count

2

3

-

-

-

-

13

17

20

16
12

20
12

-

-

11

-

-

-

-

-

-

-

-

-

-

-

-

-

1

5

16

4

5

2

1
8

3

4

5

Parastaffella sp. A

Plate 30, figs. 32, 33

Plate 30, figs. 40, 41

Remarks :-Owing to rare occurrence,detailed paleontological work was impossible. Further specimens are necessary to identify.
Occurrence:- Rare in Loc. 45; upper
part of the Zone of Profusulinella, Parastaffella cfr. vlerki subzone.
Reg. nos. :-2096-7, 2096-8

Remarl,s :-Small and involute shells
have 3 to 3.5 volutions; axial length
0.148 to 0.163 mm, median width 0.328 to
0.383 mm and form ratio 1 : 0.43 to 1 : 0.45.·
Because of ill preservation and lack ofwell oriented specimens, specific identification was impossible.
Occurrnce :-Common in Loc. 45, rarein Locs. 41 and 43, associated with Parastaffella cfr. vlerki, Parastaffella sp. Band
others; upper part of the Zone of Projusulinelia, Parastaffella cfr. vlerki subzone.
Reg. nos. :-2232-6, 2096-7.
Parastaffella sp. B

Genus Staf/ella OZAWA, 1925
Staffella sp.
Plate 31, figs. 26, 27

Shell spherical to subspherical in shapewith umbilical depressions. Matureshells have 5 to 5.5 volutions. Axial
length 0.488 to 0.713 mm, median widthFumio KOBAY ASH!

218

irregular.
Remarks:- The present species resembles Staffella breimerai GINKEL in many
respects, but chomata of the former is
not so strong and seldom extends to thepoles.
Occurrence :-Rather common in Loc.
30; Zone of Millerella, upper part of
Pseudostaffella antiqua subzone.
Reg. nos. :-2254-7-2254-80.500 to 0.675 mm and form ratio 1: 0.78
to 1: 1.20. Proloculus rather large,
spherical and external diameter 0.035 to
0.075mm. Volutions involute throughout
the growth. Spirotheca thin, and composed of indistinct tectum, outer thinlayer and inner thick layers which sometimes bear diaphanotheca-like layer.
Chomata low, massive and rather symmetrical. Tunnel low and path rather

Measurements of Staffella sp. in mm.
No.
of
sp.

W.

L.

Radius vector

Half length

P.

R.

2

1

3

4

2

1

5

3

4

5

-0.713 0.673 1. 06 0.045
0.488 0.630 0.78- 0.035
0.600 0.500 1. 20 0.045
0.675
0.075

I

2
3
4

Form ratio of vol.
1

2

3

4

. 038 . 095 . 153 . 230 . 363.048.075.103.153.245.050.088.135.218.300

-

-

5

1

3

4

5

-

-

-

-

-

-

-

17

20

-

-

-

-

-

-

-

-

-

-

-

-

-

-

12

14

18

-

25

-

-

-

33
23
60

-

-

-

-

-

-

Septal count

2

-

-

-

Tunnel angle (degs.)

1. 09 1. 51 1. 02 .96 1. 03.96 .85 .75 .79 .68.74 .78 . 87 1. 07 1. 10---.035.063.150.240.353.050.088.138.195.360.068.113 .155 .203 .273
.050.080.135.203 .305

2

1

9

4

3

5

?

Explanation of Plate 31
Figs. 1-6. Pseudostaffella antiqua (DUTKEVITCH)
1-5. Axial sections; 6. Sagittal section, x 40; Loc. 13.
Figs. 7-16. Pseudostaffella japonica KOBAYASHI, n. sp.
7. Axial section of the holotype; 8-15. Axial sections of para types;tion of paratype, x 30; Loc. 16.
Fig. 17. Profusulinella cfr. daiyamensis HASEGAWA
Oblique section, x 40; Loc. 39.
Figs. 18-19. Profusulinella sp.
18. Axial section; 19. Tangential section, x 30; Loc. 16.
Figs. 20-25. Verella sp.
20-25. Axial sections, x 30; Loc. 16.
Figs. 26-27. Staffella sp.
26-27. Axial sections, x 40; Loc. 13.
Figs. 28-31. Eoschubertella sp. A
28-31. Axial sections, x 40; Locs. 36 and 38.

16. Sagittal sec-KOBAYASHI:

Fusulinids of the Nagaiwa Formation~23
24

Plate 31

626.

Fusulinids of the Nagaiwa Formation

References
CASSITY, P. E. and LANGENHEIM, R. L. (1966) :Pennsylvanian and Permian fusulinidsof the Bird Spring Group from Arrow
Canyon, Clark County, Nevada. Jour.
Paleont., vol. 40, no. 4, p. 931-968, pis.
110-114.
GINKEL, A. C. (1965): Carboniferous fusulinids from the Cantabrian Mountains(Spain). Leise Geol. Med., Deel 34, p.
1-225, pis. 1-53.
HASEGAWA, Y. (1967): Profusulinella daiyamensis n. sp. from the Profusulinella
Zone of the Akiyoshi Limestone Group.
Sci. Rep. Niigata Univ., Ser. E, Geol.
Mineral. no. I, p. 9-15, pis. 1-3.
IGo, H. (1957): Fusulinids of Fukuji, southern part of the Hida Massif, central
Japan. Sci. Rep. Tokyo Univ. Education,
Sec. C, vol. 5, p. 153-246, pis. 1-15.
KANMERA, K. (1952): The Lower Carboniferous Kakisako Formation of southern
Kyushu with a description of Some
corals and fusulinids. Mem. Fac. Sci.
Kyushu Univ., Ser. D, vol. 3, no. 4, p.
157-177, pis. 8-12.
KOBAYASHI, F. (1973): On the Middle Carboniferous Nagaiwa Formation. Jour.
Geol. Soc. Japan, vol. 79, no. 2, p. 69-78.
(in Japanese).
MINATO, M. et al. (1953): Biostratigraphie
des Karbons im Kitakami-Gebirge, nordostliches Honshu, Japan. Jour. Geol.
Soc., Japan, vol. 59, no. 695, p. 385-399.
(in Japanese, with German summary)- - et al. (1959): Zur Biostratigraphie der
Onimaru- und Nagaiwa-Serie. Jour. Fac.
Sci., Hokkaido Univ., Ser. 4, vol. 10, no.

Nagaiwa
Kitakami

E
J:

219

2, p. 337-347.
MOORE, W. L. (1964): Notes on the morphology and taxonomic position of thefusulinid Millerella marblensis THOMPSON. Jour. Paleont., vol. 38, no. 2, p. 294305, pis. 47-48.
ONUKI, Y. (1937): Onimaru and Yukizawa
Series at Sakamotozawa and its vicinity,
Kitakami Mountainland. Jour. Geol. Soc.
Japan, vol. 44, p. 168-186. (in Japanese)
RAUSER-CERNOUSSOVA, D. M. et al. (1951):
Middle Carboniferous fusulinids of theRussian Platform and adjoining regions.
Acad. Nauk. USSR., p. 1-380, pis. 1-58.
SADA, K. (1964): Carboniferous and Lower
Permian fusulines of the Atetsu Limestone. Jour. Sci. Hiroshima Univ., Ser.
C, vol. 4, no. 3, p. 225-269, pI. 21-28.
SHENG, J. C. (1958): Fusulinids from thePenchi Series of the Taitzeho Valley,
Liaoning. Pal. Sinica, N. S., no. 7, p. 1119, pis. 1-16.
THOMPSON, M. L. (1948): Studies of American fusulinids. Univ. Kansas Paleont.
Contr., "Protozoa ", Art. 1, p. 1-184, pis.
1-38.
YAMADA, Y. (1958): On the Nagaiwa Formation.
Prof. H. Fujimoto Jubilee
Publication. p. 74-80. (in Japanese)- - (1959): On the unconformity of the
post-Nagaiwa pre-Sakamotozawa epoch
in the Hikoroichi District. Jour. Geol.
Soc. Japan, vol. 65, no. 771, p. 713-724.
(in Japanese)
ZELLER, D. E. N. (1953): Endothroid Foraminifera and ancestral fusulinids fromthe type Chesterian (Upper Mississippian). Jour. Paleont., vol. 27, no. 2, p.
183-199, pis. 26-28.

Ichinotani
Akiyoshi

/

f:k

220

Palaeontological Society of Japan Special Papers No. 17-Revisionof Matajiro YOKOYA~IA'S Type Mollusca from the Tertiary
and Quaternary of the Kanto Area
By Katura OYAMA.

Issued November 30, 1973, 148 pp., 57 plates.

Price (postage and handling included) ........................................ U. S. S 20.00

The paper presents entirely revised systematics and illustrations of the type specimensof Mollusca from the Tertiary and Quaternary of the Kanto area (Tokyo and vicinities)
which were originallY described by the late Professor Matajiro YOKOYAMA (1920-1927).
Locality records are examined and the ecological conditions of the species are considered.
This would be an indispensable reference for the late Cenozoic biostratigraphy as well as
the molluscan palaeontology.
The special papers are on sale at the Society. Orders must be accompanied by remittance, made payable to Dr. Totsuro MATSUMOTO, Editor of the Special Papers, Palaeontological Society of Japan, c/o Department of Geology, Faculty of Science, Kyushu University,
Fukuoka (Hakata) 812, Japan.

The following backnumbers are on sale at the Society and also purchasable through the
University of Tokyo Press, Hongo, Tokyo 113, Japan.
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Number 7 (Issued Nov. 30, 1961): Japanese Permian Bryozoa. By Sumio SAKAGAMI ....... .
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Number 9 (Issued Dec. 15, 1962): Bibliography of Japanese Palaeontology and Related Sciences, 1951-1960. Compiled by Fuyuji TAKAI ................................ U. S. S 18.00
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Part 1. By Sy6z6 NISIYAMA ................................................ U. S. S 33.00
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SHIKAMA ...... , ............................................................. U. S. $ 17.00
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Part II. By Sy6z6 NISIYAMA .............................................. D. S. S 26.00
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221

PROCEEDINGS OF THE P ALAEONTONLOG ICAL SOCIETY
OF JAPANr] *ti'lo:t7o~{:4;; 112 @]{§U4;;f:t 1973 "t:· 10 fl 20 lei(±) i:}lni~*"{:J:'ll'r~fllIfU.;v,'"(r,mm~7.l,t~ ($lm

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Late Lower Cretaceous flora newly foundfrom the upper beds of the Tetori
Group, Fukui Prefecture, Japan ..... .
· ............................. KI~H;RA, T.
Cryptomeria and Sciadopitys from the Late
Miocene of Akita Prefecture, Japan ..
· ........... HUZIOKA, K. & UEMURA, K.
Fossil pollen group Triprojectacites fromthe Late Cretaceous in northern Japan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . MIIG, A.
Fusulinacean fossils from the Okutama
district (No.2, Fusulinacean fossils in
the intraformational conglomerate. Part
1, Paleofusulina-Reichelina Fauna) ....
.......................... KOBAYASHI, F.
rp ijli~~-&i~1§', h~jt3:YiHJlJjtm g[ Discocyclina~~"'?V\L ............... ·t~"2f(]I. ~~DRftMt=

7 .{ ~ ~. y . ~ Y.9';I-:;;j- ~*ifl\ 0) j.;:'@:fffLpJV)~~f.............................. t~1LOO!!fl"OST" c iiiHili 0) sterraster O)~tiiW: --:::>V''"(· ....................... ;If J:'fffi~ • :'Glijijffi:Jm
J:l'iFllltEElI\lfg[O)ry ;<:,,, qo. "O}."j] -1 (Spirorbisforaminosus MOORE et BUSH) f:--:::>v''"( ....

. • • • . • . . • • • . . • • • . . . . . • • • • . • • • ... . . . f'1rl133 75: 10
Discovery of Late Permian Araxoceras
from the Toyoma Formation in the
Kitakami Massif, Northeast Japan ....
.............. MCRATA, M. & B.<\i\'DO, Y.
Khumerspira, a new genus of bellerophontid, and some Middle Permian gastropods from Cambodia ................. .
. ............... MCRATA, M. & ls I·III , K.
Marine fossils from the Moniwa Formation along the Natori River, Sendai,
Northeast Honshu, Japan. Part 2, Problematica from the Moniwa Formation'" .HATAI, K., MASGDA, K. & NODA, H.
-&i~~'~tl'tllilJjr,Hllil~ 0) :n.ti'illJlrI :J:.; J: U *LlJJlllI g[
fl i t1Cilln: --:::> v' -C .........•........ 'iil'!IlT =-Jl!I;~lz~liijfj'r.(*lt iJ· G J.;.t~ nX: I33J\1lp"~g[ .t=IJt:E 0) rJl7J(
iAiU:--:::>v'-C ...................... tm!11~±ftit
FI*0)://I..-!I..-*c=-~~1u:--:::>J.,·-C............. .

........................ JHj;!~-· ittl33p~±
Neohibolites r:JlOfL T 0 Acrothoracica ....

.................................. 1t;lft'fJl!1)
Spatial pattern of deposit feeding starfish,
Astropecten latespinosus .. MATsuKuMA, A.
Pliocene gobiid fish from Sankago Diatomite Member, Kagoshima Prefecture,
Japan ............ UYENO, T. & lWAo, Y.
~ti4Jj;Q J3 ~ ~J\lfllfip G g[T 0 =-:ill:$c conodont(:--:::>1,0,'"( ................ HHJlf,)...~· JJ'it!1~1(

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POLICY PROVISIONS OF THE TRANSACTIONS AND PROCEEDINGS OF
THE PALAEONTOGICAL SOCIETY OF JAPAN(January 16, 1973)

I

Publishment and EditingIssue-1 The "Transactions and Proceedings of the Palaeontological Society of Japan"(TPPSJ) will be published quarterly.
IB Contents-1 TPPSJ will include original papers and notes that comply with Article 2 of theConstitution of the Society as well as the proceedings of the Society meetings andnews concerning any aspect of palaeontology.
-2 Contributions will be published in the order of the acceptance by the Editorial
Board.
IC Circulation-1 All members of the Society and some organizations specified by the Council will
generally receive TPPSJ free of charge, Non-members and institutions are invited
to become subscribers.
IA

225
ID

Editorial Board
The Editorial Board will be responsible for editing TPPS] according to the policy
provisions of the Society.
-2 The Editorial Board will be composed of the Editor in Chief, who must be a member of the Council, and several members designated by the Executive Committee.
All editorial meetings will be called by the Editor in Chief.
-3 The Editor in Chief will have the authority to submit a copy of the manuscript
under consideration to an appropriate person for reviewing.
-4 The final decision on the acceptance or rejection of submitted manuscript will bemade by the Editorial Board at an editorial meeting. The Editor in Chief willreport the results of the meeting to the Executive Committee.
-5 Rejected manuscript will be returned to the author with an explanation of thereason for its rejection.
-6 An author who disagrees with the decision of the Editorial Board may take hiscomplaint to the Council. Both the Editorial Board and the author must abide by
the final judgement of the Council.
IE Proof Reading-1 Proofs will be read, as a rule, by the Editorial Board. A set of page proofs without
the original manuscript will be sent to the author.
-1

II

Contribution
IIA Eligibility-1 All members of the Society may submit contributions to TPPS]. Contributions
from non-members will be accepted for publication if they are approved by the
Executive Committee.
-2 Manuscript should be written, as far as possible, in English, French or German, andshould have been read at an annual meeting or ordinary meeting of the Society.
IIB Limitation of Manuscript-1 Manuscript should be typewritten, and should be limited to 24 printed pages including tables and text-figures. Two plates may also be added to it. Plates may not be
attached to the articles of up to 4 printed pages only without the approval of the
Editorial Board. Notes may not exceed one printed page, and no plate may be
used for it.
-2 Ten figures or 600 cm 2 of the total printed area will be permitted for a single
article. Two figures or 200 cm 2 of the total for article of less than 4 printed pages.
The original illustrations should be neat and legible to permit reduction to either
the width of one printed page (13.4 cm) or one column (6.4 cm). Figures should
be kept separate from the text of the manuscript.
-3 Excess printing charges for articles exceeding the stated limit must be borne by
the author.
IIC Style of Manuscript-1 Generic and specific names and special words should be indicated in the manuscript.
-2 Suggested positions of tables and text-figures should be indicated in the margin of
the text.
-3 Captions and explanations of text-figures, tables and plates should be submitted on
separate sheets of paper numbered independently from the text.
-4 References cited should be listed in a bibliography at the end of the text in alphabetical order under the author's name, and in chronological order, as follows:
Author's name, year, title of article, name of journal (underlined), vol., no.,pages, plates, etc.

226
-5

It is recommended that an alphabetical list of romanized geographical names be
included with Kanji (Chinese ideograph), if any. The annotated list will be placedat the end of the text.
-6 Name(s) and professional or private present address of the author(s) should appear
below the title of the manuscript.
-7 All manuscripts with the exception of notes should be accompanied by an abstract
in Japanese of SOO-words or less on separate sheet(s) of paper. The abstract should
include the title in Japanese and the author's name. Abstracts of papers by nonJapanese speaking authors will be prepared by the Editorial Board if an English
abstract be submitted with the manuscript.
IlO Obligation of Author-1 Authors are expected to follow the directions of the Editorial Board regarding
editorial matters.
-2 Manuscripts will be returned to an author if he wishes to make large-scale revisions. The revised manuscript must be submitted as a new contribution.
-3 Manuscript should be accompanied by a complete copy including text-figures, tables
and plates. Manuscript should be sent toEditor in Chief
Palaeontological Society of Japan
c/o Business Center for Academic Societies
4-16, Yayoi 2 chome, Bunkyo-ku, 113 Tokyo, JAPAN-4 The author should fill out the register card for contributions and send it to theSociety office under separate cover.
IlE Reprints-1 120 reprints without covers will be furnished free of charge. Excess charges forprinted covers, additional reprint copies, etc. will be borne by the author who must
pay the amount directly to the publisher.
Provisoi) Effecting, amending or rescinding of policy prOVISIOns will be considered by theCouncil, and must be approved by the general meeting of the Society.
ii) The approved policy provisions will nullify and replace existing" Regulations for
Publication in Transactions and Proceedings of the Palaeontological Society of Japan ".

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

New Series No. 92

December 20, 1973

CONTENTS
TRANSACTIONS
624. TANABE, Kazushige : Evolution and mode of life of Ino ceramus (Sphenoceramus) naumanni YOKOYAMA emend., an Upper Cretaceous bivalve .. 163

625. IGo, Hisaharu: Lower Carboniferous conodonts from the Akiyoshi Limestone Group, southwest Japan . . ... ....... . ..... . .... ........ .. ....... 185
626.

KOBAYASHI, Fumio: Fusulinids of the Nagaiwa Formation . . ... ... . ..... 200

PROCEEDINGS ................................................................ 221