[SSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan. New Series No. 184

December 30, 1996

Co - Editors

Kei Mori and Kunihiro Ishizaki

Language Editor

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Editorial Board

Shiro Hasegawa (Hokkaido University), Hiromichi Hirano (Waseda University), Kunihiro Ishizaki (Tohoku
University), Tomoki Kase (National Science Museum), Kei Mori (Tohoku University), Kenshiro Ogasawara(University of Tsukuba), Yoshihiro Tanimura (National Science Museum), Yukimitsu Tomida (National Science
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Cover: A terminally resorbed maxillary tooth of iguanodontids (Ornithischia : Ornithopoda). The specimen isfrom the Kuwajima Formation, the Neocomian section of the Tetori Group, in Shiramine - mura,
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 184, pp. 555-570, 13 Figs., December 30, 1996

© by the Palaeontological Society of Japan 1996

1

1019

1

Five inoceramids (Bivalvia) from the Upper Cretaceousof Hokkaido with some phylogenetic and taxonomic
considerations, Part 1. Introductory remarks, method andsystematic descriptions of one species of
Inoceramus (I.) and one of I. (Volviceramus)

MASAYUKINODA
Fukagochi 5 kumi, Oita, 870 Japan

Received 21 October 1995; Revised manuscript accepted 7 August 1996
Abstract Some of the inoceramid specimens from the upper Turonian to lower middle Coniacian of the
Obira and Ikushunbetsu areas, Hokkaido, are identical or well comparable with certain species, viz.,
Inoceramus (I.) lusatiae Andert, I. (Volviceramus) koeneni Muller, I. (Cremnoceramus) deformis Meek, I.
(Cr.) ernsti Heinz and I. (Cr.) lueckendorfensis Trtiger, all of which are cosmopolitan species that, however,
have not been recorded in Japan. In this paper, I describe these specimens, subject them to somebiometric analyses, and discuss their phylogeny and taxonomic position. Some speCimens show intermediate, or transitional characters between Inoceramus and Cremnoceramus, Cremnoceramus and Cordiceramus, and Cordiceramus and Volviceramus, suggesting their phylogenetic relationships. The
probable succession for the concerned taxa is preliminarily discussed and the taxonomic ranking of
Inoceramus, Cremnoceramus and Volviceramus is also discussed. The above taxa are here regarded assubgenera of Inoceramus. Meanwhile, the widespread occurrence of these species in East Asia hasapplication to international correlation and elucidation of paleobiogeography in the upper Turonian to lower
middle Coniacian. In Part 1, I. (I.) lusatiae and I. (V.) koeneni are described and discussed.
Key words: Inoceramus, Cretaceous, Upper Yezo Group, phylogeny, subgenus

Introduction
I. (I.) uwajimensis Yehara, 1924 and I. (Cr.) mihoensis
Matsumoto, 1957, which are good zonal indices of the upper
lower to middle and upper Coniacian, respectively, were fora long while the only Coniacian inoceramids reported fromJapan. Subsequently, I. (Cr.) rotundatus Fiege, 1930, a
cosmopolitan zonal index of the lower lower Coniacian was
described by Matsumoto and Noda (1985) on the basis of asample from the Ikushunbetsu area, Hokkaido. I. (Co.)
kawashitai Noda, 1986, from the upper Coniacian, I. (PI.)
troegeri Noda, 1992, from the lower to lower middle
Coniacian, and I. (PI.) szaszi Noda and Uchida, 1995, from theupper middle Coniacian were later also recorded. These
species, except for I. (Cr.) rotundatus, are endemic to East
Asia or unknown in other regions, possibly because they
were established only recently.
In 1988, during the construction of the Obira Dam, a large
number of inoceramid fossils were collected from lower tolower middle Coniacian sediments on the riverside andriverbed by cutting and dredging. Most of the specimens
examined were obtained by Toshio Shimanuki. The specimens are mostly well preserved, without secondary deformation. Since then, I inspected the localities together with
Shimanuki and, on another occasion, with Matsumoto andShimanuki, along the main course of the River Obirashibe,and some supplementary specimens were obtained from
fresh outcrops. Another field workers, Kenji Sanada, offered additionally some questionable specimens obtained fromthe lower Coniacian of the Ikushunbetsu Valley.
My preliminary observation suggested that some of themwere conspecific with well-known and widespread speciesof the upper Turonian through middle Coniacian, but were as
yet undescribed in Japan. Further study led to discrimination of five inoceramid species.
The occurrence of these cosmopolitan species from theupper Turonian to lower middle Coniacian of East Asia is
important for international correlation and for the consideration of paleobiogeography in those stages.

Masayuki Noda

556

MethodIn addition to general observation, some biometric analyses applied for selected characters. The basic linear
measurements and angles are shown in Figure 1. For theprocedure of biometric examination, readers may refer toHayami and Matsukuma (1971) and Noda (1992). Evaluations of the normal distribution by means of the chi-square
test and related statistics are omitted here because of theinsufficient size of the samples.
The phylogenetic relationships of the species are

preliminarily discussed on the basis of intermediate or transitional characters shown between Inoceramus (Inoceramus)
and I. (Cremnoceramus), I. (Cr.) and I. (Cordiceramus), and I.
(Co.) and I. (Volviceramus) and their stratigraphic occurrences.

Systematic description
Family Inoceramidae Zittel, 1881
Genus Inoceramus Sowerby, 1814
Subgenus Inoceramus Sowerby, 1814hhhb----1Figure 1. Basic linear measurements and angles. h: shell height, 1: shell length, b: shell breadth, s: lengthof hinge line, H: maximum length between umbo and ventral extremity, L: maximum length perpendicular to H, Q!:
anterior hinge angle, fJ: beak angle, angle of umbonal inflation, y: posterior hinge angle, 0: obliquity, angle
between hinge line and H.
Figure 2. Inoceramus (Inoceramus) lusatiae Andert, all figures natural size. 1a-e. JG.H3095 from lac. Ik 7002. 1a:right valve, 1b: left valve, 1c: anterior view, 1d: posterior view, 1e: dorsal view (call. K. Sanada, 1983). 2a-e. JG.H3096
from lac. Ik 7001, 2a: left valve, 2b: right valve, 2c: dorsal view showing anterior ear, 2d: posterior view, 2e: anterior view
(call. K. Sanada, 1983).

1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1557

Masayuki Noda

558

Type species.-Inoceramus cuvierii Sowerby, 1814, by
subsequent designation by Cox (1969).
Oiagnosis.-See Cox (1969, p. N315), Herm et a/. (1979, p.
52, 53) and Walaszczyk (1992, p. 29).

Inoceramus (Inoceramus) lusatiae Andert, 1911
Figures 2-1a-e, 2a-e; 3-1a-c, 2-4Inoceramus lusatiae Andert, 1911, p. 54, 56, pI. 2, figs.1a, b, pI. 3,
fig. 3, pI. 8, figs. 3-5; Andert, 1934, p.126, pI. 7, figs. 1-3,
text-figs. 14a, b; Dobrov and Pavlova, 1959, pars, p. 151, pI.
4, figs. 5, 6a, b; Trager, 1967, p.73-76, pI. 8, figs. 2, 3 ;Keller, 1982, p.87-89, pI. 6, fig. 3; Szasz, 1985, p.172, pI. 4,
figs. 2, 3; Trager, 1991, p. 29, pI. 3, fig. 6; Tarkowski, 1991, p.
113, pI. 7, fig. 4; Walaszczyk, 1992, p. 32, 33, pI. 27, figs. 1-3,
5,6.
Mytiloides lusatiae (Andert). Kauffman, 1977, p. 240, 243, pI. 9,
figs. 17, 23; Kauffman, Cobban and Eicher, 1978, p. XXXIlI-9,
pI. 9, figs. 17, 23.
MytiJoides ? lusatiae (Andert). Kauffman, 1978b, pi. 2, fig. 5, pi. 4,

fig. 5.
not Inoceramus lusatiae Andert.
23, fig. 1.

Pergament, 1971, p.94, 95, pI.

Lectotype.-The specimen figured by Andert (1911, pI. 2,
figs.1a, b; Trager, 1967, pI. 8, figs. 2a, b) was designated as
the lectotype by Trager (1967, p.73).
Materia/.-JG.H3095, 3096, 3100, 3101, 3102, 3201 and3202 from the Ashiyachi-zawa (so-called Yoshiashi-zawa)in the Ikushunbetsu area, central Hokkaido.
Repository.-JG: Collections of the Jonan Geological
Association, Oita, kept tentatively in Noda's personal laboratory, Oita.
Oescription.-Shell of medium size, nearly equivalve andhighly inequilateral, with very oblique outline. Valve considerably inflated with some variation, from anterior to posterior,and uniformly or geniculately convex along growth axis.
Anterodorsal part steep or perpendicular to commissureplane, posterior one also steep. Posterior auricle well developed, nearly regularly triangular and demarcated clearly fromdisk with a shallow sulcus. A small anterior auricle
devoloped in some specimens. Umbo situated at anterior
end and projected slightly above hinge line. Growth axis
straight, with a considerably large obliquity. Anterior margin
straight or slightly concave, continuing to broadly curved
ventral one; posteroventral part narrowly bent and passing
into a guitar-shaped or hourglass-like concave posterior
margin, which forms an acute angle with hinge line. Hingeline is longer than half of shell length.
Surface ornamented with commarginal undulations andcommarginal rings ("com marginal" may be a more appropriate term than "concentric" for ornamentation of intricate
outline as in this species) in combination, but in umbonal
region major o~nament inconspicuous. Commarginal undulations gradually more predominant with growth, fairly irregular in size, interspacing and intensity, weaken in anterior walland continue to posterior auricle. Commarginal rings superimposed on major ornament and also irregular in size, interspace and intenSity.
Biometry.-Measurements are shown in Table 1.

The individual relative growth of shell length (1) vs. shellheight (h) and shell breadth (b) vs. shell height (h) in eachvalve of the specimen JG.H3095 is shown in Figure 4. In
reference to the basic criteria of Hayami and Matsukuma(1971, p. 150, table 2, p. 151, fig. 8), the relative growth of I vs.
h shows a negative allometry in both valves, and that of b vs.
h is regarded as a positive allometry.
Ontogenetic changes in I/h, b/h and obliquity (0') aredemonstrated in Figures 5A, Band C, respectively. As isclear from Figure 5A, the simple ratio I/h decreases withgrowth and b/h changes abruptly at a certain stage of
growth which is variable among individuals as shown in
Figure 5B. The obliquity is nearly constant throughout thegrowth with about 66 on average.
Profiles of two specimens in two directions are illustrated
in Figure 6. In the cross section A of specimen JG.M3096,
the posterior part is steeper than the anterior slope, and aposterior auricle is clearly demarcated from the disk, and asmall anterior ear is clearly developed.
Remarks.-The specimens JG.H3095 and 3096 are bothclosed valves. JG.H3095 is scarcely deformed secondarily
but the valves are displaced along the commissure plane
and the left valve has been somewhat eroded. JG.H3096 isslightly deformed obliquely, missing the left umbo, with small
anterior ear. JG.H3100 is a right valve, fairly well preservedwith original margin, and somewhat less convex. JG.H3101
in the same nodule as JG.H3100 is a left valve of a different
individual, with a well preserved umbonal portion but lacking
the ventral part. JG.H3102 is an imperfect specimen.
JG.H3201 and 3202 from the upper Turonian near theConiacian localities are fairly large, somewhat imperfectindividuals, lacking the ventral and anterior parts, respectively. The former is characterized by its broad outline, whichsomewhat resembles I. (I.) lamarcki sWmckei Heinz, 1926,
whereas the latter is an elongate form like JG.H3095, 3096,
3100 and 3101.
Comparison and discussion.-The specimens figured byAndert (1911, 1934) show a large extent of variation in marginal
curvature, shape of auricle and obliquity as originally noticed
by Andert himself and then by Trager (1967). The triangularauricle well defined from the disk, guitar-shaped curvature of
the posterior margin, strongly vaulted shell and coarse andirregular commarginal ornamentation are regarded as theessential criteria of I. (I.) lusatiae.
These specimens are agree well with all aspects of I. (/.)
lusatiae described by Andert (1911, 1934) and by Trager (1967),
but the development of a small anterior auricle in JG.H3096
may be exceptional.
According to Trager (personal communication, Oct. 5,
1993) there are two extreme forms in the German specimens,i.e., an elongate form and a subquadrate one, both are linked
by various grades of intermediates. Troger (personal letter ofOct. 5, 1993) attempted a precise comparison between theoriginal specimens of Andert and the plaster replica of theJapanese ones. The data (Figures 7,8) are taken fromTrager's letter with his approval.
Ontogenetic changes in Na/Ha (=L/H), Vo/Ha (length ofanterior margin/H) and Wa (= 0') are shown in Figures 7A, B
and C, respectively. Solid line represents the original spec i-

1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1

Figure 3. Inoceramus (Inoceramus) lusatiae Andert, all figures natural size. 1a- c. JG.H3100 from lac. Ik7003,
1c shows ligament structure (call. K. Sanada, 1982). 2. JG.H3201 from lac. Ik7000 (call. K. Sanada and M. Noda,
1994). 3. JG.H3202 from the same locality (call. K. Sanada and M. Noda, 1994). 4. JG.H3101 from lac. Ik7003(call. K. Sanada, 1982).

559

Masayuki Noda

560

Measurements of Inoceramus (Inoceramus) lusatiae Andert from Hokkaido Linear dimensions in mm.
L
I
b
s
H
a
valve
h
fJ
89
62.0
25.0
79.0
63.0
L.
73.0
86.0
68.0
120
86
81.0
66.0
26.8
R.
120
79
70.0
61.4
24.5
33.0
75.0
57.0
L.
70.0
61.4
57.0
1W
R.
23.0
33.0
75.0
84
76.0
62.7
21.3
43.8
89.0
70.0
100
76
R.

Table 1
specimeny0

JG. H3095
JG. H3096
JG. H3100

0

0

0

-

0

-

0

specimen

valve
L.

60°

0

0

87°

remarks
I/h
s/I
L/H
b/h
closed valves, posterior auricle partially
0.85
0.34
0.80
JG. H3095
R.
0.81
0.79
missing
0.33
L.
0.87
0.35
0.76
0.54
closed valves, anterior auricle present
JG. H3096
R.
0.87
0.33
0.79
0.54
JG. H3100
R.
0.83
0.28
0.79
0.70
somewhat less convex
Abbreviations. L.: left valve, R.: right valve, I/h: simple ratio I versus h, b/h: simple ratio b versus h, L/H: simple ratio L
versus H, s/I: simple ratio s versus I.
cJ

64
66
6T
6T
5T
0

0

A •
B

• JG.H3095L
JG.H3095R

0------0

I.b
mm

60
50

0.901
1= 1.288hr = 0.9993

A

40

1= 1.342ho.889

B

r = 0.9977

30

20

/

10
9
"
8 ,/-

.
/

7
6

/.

/.

5

/.

4
/.

"

/.

/.

/

/

"

b = 0.153h1.129/.

A

r = 0.9300

/.

3

/./

"

B

/
/

2

b = 0.164h1.121
r = 0.9980

....0

/.
/.

7 8 9102030

40 50 60 70 80

h mm

Figure 4. 'Diagram showing individual relative growth
between I and h, and band h (in mm) of Inoceramus (I.)
lusatiae. (Specimen JG. H3095).

mens of Andert, and broken and chain lines the Japanese
ones. As is clear from Figure 7, the original specimens fromthe type locality (Sennenberg, Saxony near the Bohemianborder) show a large extent of variation in the characters
examined, and the Japanese specimens lie within the rangeof variation.
As shown in Figure 8A, at the growth stage 30 mm-50 mm
in H, the size of commarginal undulations of the specimens
examined is indistinguishable between the German andJapanese specimens, whereas in the subsequent stage,
more than 50 mm in H, as is clear from Figure 8B, the size ofcommarginal undulations broadens rapidly in the German
specimens in contrast to their remaining nearly constant in
the Japanese ones. This is the most conspicuous difference between the German and Japanese specimens. On
the basis of the above examination, Trager suggested to me
that the Japanese specimens may be conspecific with
Inoceramus (I.) lusatiae Andert, or a special new subspecies.
The specimens from the Coniacian of the Babadag Basin,
Romania, figured by SZ8.SZ (1985, pI. 4, figs. 2a, b, 3) aresomewhat less convex and predominant in minor ornamentin comparison with the specimens examined.
Keller's specimen (1982, pI. 6, fig. 3) from the upper middle
Turonian of Sack-Mulde of Germany resembles theJapanese specimens in marginal curvature, obliquity, shellconvexity and surface ornamentation, but the stratigraphic
position of his specimen is substantially lower than theothers.
Moskvina's specimen (1959) from the uppermost Turonianof Crimea figured in her pI. 4, fig. 4 also resembles specimen
JG.H3100 (Figure 3-1a c) in the oblique outline, moderately
convex shell and surface ornamentation, but another one (pI.
4, fig. 5) from the uppermost Turonian of Dagestan it is moreor less questionable to assign strictly to I. (I.) lusatiae so far
as I can observe from the photograph.
The specimen from the upper Turonian of the Opole
Trough of Poland figured by Tarkowski (1991, pI. 7, fig. 4) iscomparable with the specimen JG.H3102 in the much higher
outline, small posterior auricle, less obliquity and major1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1

561

I/h

A

1.05
L>:••

1.00

0.;

_----48

JG.H3095L

0--------0

JG.H3095R.... -.-._._....

JG.H3096L

6 .................... -6

JG.H3096R

70

Hmm

0.95
0.90
0.85
0.80

20

10

b/h

50

40

60

80

6 ............ ·4-

B

0,45

30

!~.-.-.;::. . . . . .

l>

....

"

0.40

"

",

·"-<c,,..~~::>...

0.35

..t\,

,,

p'

,1/

0.30
.j.,

0.25

... _.....

#'-0----_-<:1""'.,..

...

.,. . . .a......

'//'....... ""-J''0/.,...,..~-----;~----~--------

10

20

30

40

50

60

70

80

Hmm

J

c

80

75
70
65

10

20

30

40

70

Figure 5. Diagram showing ontogenetic changes in selected characters of Inoceramus (I.) lusatiae.
B: b/h, C: obliquity.

A: I/h,Masayuki Noda

562

B

A

JG.H3095R

JG.H3095R
/

p

a

/
LYL-_________________________________
uJ

,

...

\
\,

JG.H3095L

JG.H3095L\\"' ...

"''''

...

\

\

\

____________________~_~~

L-~P

U

,

V \

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

20mm~
JG.H3096R
p

a

,

/

/

I

I

JG.H3096L

JG.H3096L

I

\
\

p

a

\

U

v

Figure 6. Profiles of two selected specimens of Inoceramus (I.) lusatiae in two directions. A: cross section
at the highest point of shell convexity, B: vertical section along the growth axis. a: anterior, p: posterior, u:umbo, v: venter.

ornament less predominant.
Walaszczyk's specimens (1992, pi. 7, figs. 1-3, 5, 6) fromthe uppermost Turonian through middle Coniacian of theCentral Polish Upland are similar to those of Andert andthose from Japan in the obliquely elongate shell and surface
ornamentation, but unfortunately, as he mentioned (1992), theposterior auricle which is one of the important characters of
the species is lacking. A guitar-shaped, sinuous curvatureof major commarginal undulations is partially perceptible on
the specimen ME: ML1216 (Walaszczyk, 1992, pi. 27, fig. 2),
which suggests clearly the presence of a triangular auricle.
Specimens of Mytiloides lusatiae (Andert) from the uppermost Turonian of the Western Interior figured by Kauffman(1966, pi. 9, figs. 17, 23) and Kauffman et al. (1978, pi. 13, figs.
17, 23) accord well with Andert's specimens in the outline
and triangular and clearly defined posterior auricle, and thespecimen USNM240318 (fig. 17) resembles closely JG.H3100
in the marginal curvature and surface ornamentation withless conspicuous major undulations. Another specimen

USNM240338 (pi. 14, fig. 5) from the same horizon is fairly
distinct from the type specimens, as Kauffman et al. (1978)
noted as a less oblique variety, which closely resembles the
specimen JG.H3102.
The specimen of M. ? lusatiae from the lower lower
Coniacian of Bohemia, figured by Kauffman (1978b, pi. 2, fig.
5), is also somewhat distinct from the type specimens by aless oblique outline, as he remarked.
Pergament's specimen nO.3588-540-123 (Pergament,
1971, pi. 23, fig. 1) from the upper Turonian (lower Coniacian
after Pergament) of northwestern Kamchatka is quite doubtfully assigned to I. (I.) lusatiae as Walaszczyk (1992, p.33)
pointed out, because of its posterior margin without a guitarshaped sinus and the shape of the posterior auricle with no
sharp boundary of the disk. The specimen appears to becloser to I. (I.) lamarcki Parkinson and/or its variety. In
connection with this, Trager (personal letter of Oct. 5, 1993)
suggested that Pergament's specimen is referable to I. (I.)
lusatiae.

1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1

563

Na/Ha

96
100

•
ANDERT, 1911) pl.2, f i g.l

90
80

70

,,

60

'.

o

0

no. NL 8••

no. NL60G

a0

no. NL6JG

I:>

t:;.

nO.89

... -------. JG.H3095...... -._._.-. JG.H3096

Vo/Ha

%

B

80
r

--~ \

70

•

,,

.I

\ y" /~.

_~_-_-e

/

£'

60
50

WA

C

800

--

700
~.~~---

....

---

'

..........

~.-.-

.

....

5
400

10

20

30

40

50

60

70

80

Hmm

Figure 7. Diagram showing ontogenetic changes in selected characters of Inoceramus (I.) lusatiae. Comparison with German specimens (by the courtesy of Prof. Dr. K.-A. Trager). Na/Ha=L/H, Vo/Ha= length of anterior
margin/H, WA=o.

Phylogeny.-On the basis of morphological and stratigraphical evidence, I. (I.) lusatiae from the uppermost Turonian
through lower Coniacian (upper Turonian in the German
scale before Seibertz, 1979) is regarded by Trager (1969)
as an immediate descendant derived from I. (I.) lamarcki
stiimckei Heinz, 1926, of the upper middle Turonian.
Besides, specimens comparable with I. (I.) lamarcki stiimckei,

not described in Japan yet, are obtained from the middle
Turonian of some areas of Hokkaido. JG.H3201 (Figure 32) from the uppermost Turonian of Ikushunbetsu and Pergame nt's specimen from the same stage of Kamchatka resemble the type specimens of I. (I.) lamarcki stiimckei. Thisevidence would help to support the Trager's scheme (1969,
fig. 8). According to Trager (1969), descendants of I. (I.)

Masayuki Noda

564

A

specimen

5~l.:.:.:.:................!. .:.:.:.:.:.:.:.:.:.:.:.~.:.:.:./~:~.:.:.:/(.:. :.:;: : : .:.:. .: : : +Ut~:}~UL:}:,;,;.,;.: : :~:I:.---r-----t

5.01- 6.00

6.01-7.00

7.01 - B.OO B.01 - 9.00 9.01- 10.00

10.01- 11.00 11.01-12.00 mm

specimen:.:.:.:.:.:.:.:.:.:.:.:
I
5j,_~~__i~····~~·····1~zb·······~·········"$;';';l~~:::::~~::~~~;;~;::.:.:.:.:.:.:.:.:.:.:.: .........................:.:.}}:.;:;:::..............................

i5.01 - 6.00

6.01 -7.00 7.01- B.OO 8.01- 9.00

9.01- 10.00 1O.Q1-11.00 1l.01-12.00mm

Figure 8. Diagram showing the wave length of commarginal undulation of Inoceramus (I.) lusatiae. A: growthstage 30 mm-50 mm in H, B: growth stage 50 mm-70 mm in H, black: Japanese specimens, dotted: German
specimens (by the courtesy of Prof. Dr. K.-A. Trager).

lusatiae are unknown, as is the case in Japan, which can be
ascribed either to its terminal extinction or to insufficientcollection up to the present. The species may possibly
represent an offshoot of the main stocks of inoceramids as
schematized by Troger (1969).
On the other hand, Kauffman (1977) regarded the present
species as being closer to the genus Myti/oides than to
Inoceramus (Inoceramus). The specimens I have observed,
however, seem to show that the species is more closely
related to I. (Inoceramus) than to the group of Mytiloides on
account of the triangular and clearly defined posterior auricle, fairly thick hinge plate and coarser and irregular commarginal undulations.
Occurrence.-The locality map is shown in Figure 9.
Loc. Ik7000.- Topographic map "Katsurazawako", Scale
1 : 25,000. A cliff at the confluence of the Ashiyachi-zawa(so-called Yoshiashi-zawa) and the Hatsudenshono-sawa,in usual, under the water level of Lake Katsurazawako.
Sandy siltstone bed of Unit Ilia (Matsumoto, 1965), uppermost
part of the lower Upper Yezo Group. Associated with I. (I.)
teshioensis Nagao and Matsumoto, I. (I.) tenuistriatus Nagao
and Matsumoto and Otoscaphites sp.
Loc. Ik7001.- Topographic map, ditto. Scree of the small
left branch of the Ashiyachi-zawa. Sandy siltstone bed of
Unit IIlb, lower part of the middle Upper Yezo Group (Matsumoto, 1965).
Loc. Ik7002.- Topographic map, ditto. A left bank of thelower reaches of the Ashiyachi-zawa, about 270 m upstream
from the Kumaoi Power Station, submerged at full waterseasons.
Loc. Ik7003.- Topographic map, ditto. A stream floor of
the Ashiyachi-zawa, about 220 upstream from loc. Ik7002,
also submerged at full water seasons. Associated with
Mesopuzosia yubarensis (Jimbo), Gaudryceras denseplicatum
Jimbo and Scaphites sp. No effective zonal index is known,
but at a horizon about 20 m above, I. (I.) uwajimensis Yehara,a good index of the upper lower to middle Coniacian, occurs
abundantly.
Range.- The described specimens were obtained from thementioned horizons. Loc. Ik7000 represents the uppermost

Turonian, and Ik7001, 7002 and 7003 are probably the lower
Coniacian. The Tu/Co boundary is regrettably unclear
because it is below the surface of Lake Katsurazawako.
As discussed above, the present species is recorded fromthe middle Turonian to middle Coniacian of various regionsof Euramerica, with some time lag in each area, but mostly
from the uppermost Turonian to lower Coniacian. The
stratigraphic position of this species in Japan is well inaccordance with these occurrences.
Subgenus Volviceramus Stoliczka, 1871
Type species.-Inoceramus involutus Sowerby, 1828 (InCox, 1969) was originally designated by Stoliczka (1871).
Diagnosis.-See Stoliczka (1871), Meek (1876) and Cox(1969).
Remarks.- Volviceramus was proposed by Stoliczka (1871)
as a full genus, and the original diagnosis was given for thetype species I. involutus. Hattin and Cobban (1977) and
Kauffman (1977) followed Stoliczka and adopted it as a full
genus, whereas MOiler (1887) and Scupin (1913) regarded
Volviceramus as a subgenus of Inoceramus without any
comment. Heinz (1932) separated a group of I. epigonus
Heinz, 1932, which is not so highly inequivalve as I. involutus,
from the genus Volviceramus and established the full genus
Tactoceramus, designating I. epigonus (V. epigonus ofHeinz's sense) as the type species, with a brief definition.
Thus the generic name is available under Art. 13 of the I.C.
Z.N. Vokes (1967,1980) regarded Tactoceramus as an available name, while Cox (1969) regarded it as a junior synonymof Volviceramus. Heinz (1932) also proposed Cymatoceramus as an independent genus and designated I. koeneni
MOiler, 1887, as the type species, but gave no diagnosis. It
is surely invalid. Cox (1969) regarded Cymatoceramus as a
junior synonym of Volviceramus. Tethyoceramus is anotherfull genus proposed by Heinz (1932), who designated I.
emigrans Heinz, 1932, as the type species, but it is also
invalid, because given diagnosis of the type species isinsufficient, with no designation of type specimen or any
illustration.

1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1~

I kushun etsu

144 0
100km

44°+-----J

146

Figure 9.

Locality map of the Ikushunbetsu area.

Ik2714
Ik2799 ..
.. wOetsu-gono'''o565

Masayuki Noda

566

Sornay (1980) gave a revised precise definition, positioning
Tethyoceramus as a subgenus of Inoceramus and designating I. basseae Sornay, 1980 (= Tethyoceramus aft. emigrans
Heinz) from the lower Coniacian of Madagascar as the typespecies. At that date, Tethyoceramus came to be valid
under Art. 13.
As seen in the synonym list of Sornay (1980), some specimens of I. (Cr.) ernsti and I. (V.) cf. involutus were included inTethyoceramus. In reality there are various intermediate
forms between Volviceramus and Tethyoceramus.
I. (Cremnoceramus) is slightly or moderately inequivalve(Herm et al., 1979) whereas I. (Volviceramus) and I. (Tethyoceramus) are highly inequivalve. This is taken as themost important criterion for separating these subgenera, but
the distinction between I. (Volviceramus) and I. (Tethyoceramus) is quite obscure in practice. In my opinion, thedistinction between these two subgenera may be rather
unnecessary, then Tethyoceramus should be included inVolviceramus as a junior synonym.
Woods (1912b, figs. 42, 43) connected a highly inequivalve
variety of Inoceramus lamarcki with I. involutus. This suggests emphatically that I. (Inoceramus) is closely related to I.
(Volviceramus). These forms are hardly assignable either to
Inoceramus or to Volviceramus.
From the above discussion, it would appear natural that
Volviceramus be ranked as a subgenus of Inoceramus.

Lectotype.- The specimen figured by MOiler (1887, pI. 17,
fig. 1) was designated as the lectotype by Trager (1969).
Material.-A single specimen (JG. H3019) from loc. Ob0013
in the Obira area, Hokkaido.
Oescription.-Shell large-sized, highly inequivalve, inequilateral, much higher than long. Left valve strongly
convex from anterior to posterior and along growth axis.
Anterior wall angularly bent from disk and deeply concave tocommissure plane. Posterior half of flank steeply sloped toposterior margin and hinge part, and almost perpendicularnear umbo. Umbo terminal, and left one highly projected
above hinge line, strongly involuted and twisted forwards.
Three radial elevations developed, of which anterior one
demarcates clearly between anterior wall and disk, median
one is prominent and runs posteriorly along growth axis, and

I.b
mm

100
90
80
70

I =1.098ho . 97860

r= 0.9998

00

50

Inoceramus (Volviceramus) koeneni MOiler, 1887

40

Figures 1O-a-d30

Inoceramus (Volviceramus) koeneni MOiler, 1887, p. 412, 413, pI.
17, fig. 1.
Inoceramus lamarcki Parkinson (pars). Woods, 1912a, pI. 52, figs.

20

4,6.
Inoceramus connecting Inoceramus lamarcki Parkinson with
Inoceramus involutus Sowerby. Woods, 1912b, p. 9, figs. 42,
Inoceramus koeneni MOiler. Heinz, 1928b, p.37, pI. 3, fig.2;

Heine, 1929, p.98-100, pI. 15, fig. 63, pI. 17, fig. 66, pI. 18, fig.
67; Andert, 1934, text-fig. 16, pI. 8, figs. 2b, c; Trager, 1969,
p.69-87, pis. 1, 2; Ivanikov, 1979, p.13, list; Ciesliriski andBlaszkiewicz, 1984, p.364, pI. 159, fig.l; Ciesliriski andBlaszkiewicz, 1989, p.255, pI. 160, fig. 1.
Cymatoceramus (Cymatoceramus) cf. koeneni (MOiler). Heinz,
1934, p. 253, 254, pI. 19, fig. 3.
Inoceramus (Cremnoceramus) koeneni MOiler. Hattin and Cobban, 1977, p. 191, 192, fig. 9-3.
Volviceramus? koeneni (MOiler). Kauffman, 1978a, p.IV-8, list.
Volviceramus koeneni (MOiler). Kauffman, 1978b, p. XIII-2, list.

r= 0.9981

6

8 9 102030

40

50 60 708090100

h mm

Figure 11. Diagram showing individual relative growth
between I and h, and band h of Inoceramus (V.) koeneni.
(Specimen JG.H3019).

127.7

s
46.0

specimen
JG. H3019

l/hll-60mm
0.97b/hll 60mm
0.37

Inoceramus (Volviceramus) koeneni MOiler.

= 0.281 h1• 082

7

specimen
JG. H3019

Figure 10.

b

10
9
8

43.

1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1567

Masayuki Noda

568

Biometry.-Measurements are shown in Table 2.
The individual relative growth of I vs. hand b vs. hare
demonstrated in Figure 11. The former shows isometry andthe latter positive allometry.
Ontogenetic changes in I/h, b/h and obliquity (0) are
shown in Figures 12A, Band C, respectively. As is clear
from Figure 12A, I/h is nearly constant throughout growth,
while b/h increases gradually with growth. The obliquity
increases rapidly in the early stage of growth and thengradually, which suggests a spirally twisted growth axis.
Profiles in two directions are illustrated in Figure 13.

posterior one is blunt. Anterodorsal margin long andstraight continuing to broadly rounded anterior one, andnarrowly curved, passing into asymmetrically arcuate ventral
margin; posterior margin fairly short, forming an obtuse
angle with hinge line which is short, about one-third shelllength. Posterior wing narrow and small. Hinge plate fairly
thick, ligament pits rectangular with round ridge.
Surface ornamented with concentric ribs which are fairly
crowded near umbo, prominent, sharp-topped and irregularin strength, and concave interspaces which broaden withgrowth. Minor ornament scarcely perceptible.

I/h

A

1.05
1.00

10

20

30

40

50

60

b/h

70

80

90

100

110

120

130

Hmm

70

80

90

100

110

120

130

Hmm

70

80

90

100

"0

120

130

Hmm

B

0.45

0.40
0.35
0.30

10

20

30

40

50

60

c

105
100
95
90
85
80
75
70
65

10

20

30

40

50

60

Figure 12. Diagram showing ontogenetic changes in selected characters of Inoceramus (V.) koeneni.
A: I/h, B: b/h, C: obliquity.

1019.

Inoceramids from Upper Cretaceous of Hokkaido, Part 1

A

569

B
/

/

/

a
1

b
........

c

20mm

\/

C

/",

."
",,,';",,,,,,,,,,
I,,"
a,,"

I

1\

..... )//

1

/'

",

,,"

I

JG.H3019/' ,,"';

U

I,,"

v

Figure 13. Profiles of Inoceramus (V.) koeneni (specimen JG.H3019) in two directions A-a: cross section at thehighest point of shell convexity, A-b and A-c: cross sections at certain growth stages shown by broken lines in
B. B: vertical section along the growth axis.

Three cross sections a, band c of Figure 13A correspond tochain lines a, band c of Figure 13B, which are perpendicular
to the commissure plane at three different growth stages asshown by the broken lines.
Remarks.- The specimen JG. H3019 is a well preserved
left valve and a fragment of the right valve without secondary
deformation. The prismatic layer is preserved only near thebeak and posterodorsal parts. The precise ornamentation,
therefore, is uncertain. The right valve is visible on the
opposite side of the left valve, but it is too imperfect toassume the original outline. From the left umbo being
strongly involute and twisted forwards, the right umbo ispresumed to have been not so prominent and the shellconvexity may have been weaker than that of the left valve.
Comparison and discussion.-JG. H3019 is characterized
by the strongly involuted left umbo, considerably inflatedshell and twisted growth axis, which are in common withfeatures of the subgenus Volviceramus. It surely resembles
closely the specimens of I. (V.) koeneni described by MOiler(1887) and Trager (1969). In addition, the radial elevations
situated somewhat posteriorly along the growth axis andblunt posterior one in JG.H3019 are closely similar to those
figured by Trager (1969, p. 72, fig. 4), and the sharply concave
anterior wall resembles closely that of the specimen figuredby Trager (1969, pi. 2, fig. 5). Moreover, the cross sections atthree growth stages illustrated in Figure 13A agree well withthose of type I of Troger (1969, p. 72, fig. 3). From the above
comparison, JG. H3019 is referable to I. (V.) koeneni, eventhough having somewhat weaker convexity than the German
speCimens.
The type species of Volviceramus, I. (V.) involutus Sowerby, is clearly distinct from the present species in its extreme
inequivalveness as MOiler (1887) discussed originally.
I. (V.) umbonatus Meek and Hyden, 1858 also resembles
the present species, as MOiler (1887) pointed out. According to the description by Meek (1876) the former shows amuch higher outline than the latter.
The specimen figured by Woods (1912a, pi. 52, figs.4a, c)somewhat resembles I. (V.) koeneni in the highly inequivalve
shell and vaulted outline in anterior view and the welldeveloped posterior wing.
I. (Cr.) ernsti somewhat resembles the present species inthe vaulted shell with a concave anterior wall and coarse
and irregular major ornament, but the former is discriminatedfrom the latter in subequivalveness in contrast to the latterbeing highly inequivalve and having twisted left umbo.
Phylogeny.-Woods (1912a, b) suggested that I. (V.)
involutus derived from I. (I.) lamarcki through a connecting
form figured by himself (1912a, pi. 52, figs. 4, 5; 1912b, p. 9,
figs. 42, 43) is surely comparable with I. (V.) koeneni. Troger(1969, p.74, fig. 6) demonstrated schematically thephylogenetic relationships of I. (V.) koeneni as an immediate
descendant of I. (Cr.) ernsti and the ancestor of I. (V.)
involutus. I support his scheme fully. Collom (1993, personal communication) suggests another lineage in thephylogeny. I. (V.) koeneni in common with I. (Co.) spp. has
a vaulted shell, concaved anterior wall, and twisted umbo,and develops radial elevations. The essential distinction,
though, is that the subgenus Volviceramus became highly
inequivalve with time, whereas the subgenus Cordiceramus isnearly equivalve. I. (V.) involutus, however, has not beenfound in Japan.
As is clear from the above discussion, I. (Cr.) ernsti isregarded as a common ancestor of I. (V.) koeneni and I. (Co.)
kawashitai Noda, 1986, from the upper Coniacian of Hokkaido
and Kyushu (Teraoka et al., 1992; Noda, 1994), and thecharacters of the three species suggest undoubtedly the
relationships among I. (Cremnoceramus), I. (Volviceramus)
and I. (Cordiceramus).
Occurrence.-For the locality map of the Obira area, thereaders may refer to Noda (1992, p. 1312, fig. 1). Loc. Ob0013(=Ioc. NH79 of Tanaka, 1963). Topographic map "Takishita", Scale 1 : 25,000. A left bank about 150 m below
Bridge Takimi-Ohashi, now below the surface of artificial
Lake Obireshibeko. Silty sandstone bed at the base of Unit
Ub (Tanaka, 1963; Tsushima et al., 1958). Associated with
I. (Cr.)deformis Meek, I. (I.) uwajimensis, I. (I.) aff.
pedalionoides, Didymotis akamatsui (yehara), Damesites
damesi (Jimbo), Neophyl/oceras subramosum Spath,
Tetragonites glabrus (Jimbo) and Mesopuzosia yubarensis(Jimbo).
Range.- The specimen handled was obtained from Unit
Ub, associated with I. (Cr.) deformis and I. (I.) uwajimensis.
According to Kennedy and Cobban (1991), I. (Cr.) deformis

570

Masayuki Noda

indicates the lower middle Coniacian of the Western Interior,to Kauffman (1978a, b) and Trager (1981b, 1989) the species
represents the middle Coniacian of Europe, while Seitz (1959)
positioned the species in the middle Coniacian (Unterconiac
before Seibertz, 1979), and Collom (1993, personal communi-

cation) in the upper lower Coniacian. Unit Ub is regarded aslower middle Coniacian deposits (Noda, 1992, p.1313). The
stratigraphic range of the species in Japan should be madeclear on the basis of further field work in various regions.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 184, pp. 571-591,14 Figs., December 30,1996© by the Palaeontological Society of Japan 1996

/1020 / Five inoceramids (Bivalvia) from the Upper Cretaceousof Hokkaido with some phylogenetic and taxonomic
considerations, Part 2. Systematic descriptions of three
species of Inoceramus (Cremnoceramus) andconcluding remarks
MASAYUKINODA
Fukagochi 5 kumi, Oita, 870 Japan
Received 21 October 1995; Revised manuscript accepted 7 August 1996
Abstract Described along with phylogenetic and taxonomic considerations are three species of Inoceramus (Cremnoceramus) from the Upper Turonian to lower middle Coniacian of the Obira and Ikushunbetsu
areas of Hokkaido: Inoceramus (Cremnoceramus) deformis Meek, I. (Cr.) emsti Heinz and I. (Cr.)
lueckendorfensis Trager. Concluding remarks on the significance of occurrence of cosmopolitan species(including two species described in part 1) from Japan are also made.
Key words: Inoceramus, Cretaceous, Upper Yezo Group, phylogeny, subgenus

Introduction
In the present part, three inoceramids from the upper
Turonian to lower middle Coniacian of the Upper Yezo Groupof Hokkaido are described.
Systematic descriptions
Family Inoceramidae Zittel, 1881
Genus Inoceramus Sowerby, 1814
Subgenus Cremnoceramus Cox, 1969
Type species.-Inoceramus inconstans Woods, 1912a,
designated by Cox (1969).
Diagnosis.-See Cox (1969, p. N314) and Herm et al. (1979,p.58).
Remarks.-The name "Cremnoceramus" was originallyproposed as a full genus of the Inoceramidae Heinz fam.
nov., 1932, by Heinz (1932) who gave a brief diagnosis anddesignated "Cremnoceramus absolutus Heinz, 1932" as thetype species. The name was invalid, as Vokes (1967)
pointed out, because neither diagnosis nor illustration wasgiven for the type speCies. Cox (1969) validated Cremnoceramus as a subgenus of Inoceramus Sowerby, 1814 (in Cox,
1969), with a clear revised diagnosis and designated I. incon-stans as the type speCies.

At that date Cremnoceramus
became an available name, with Cox as its author, under Art.
50 of the I.C.l.N. In spite of this situation, Vokes (1980)
wrongly regarded the name as still invalid. Herm et al. (1979)
ranked Cremnoceramus as a full genus of the Inoceramidaewith supplemental description and remarks. However, they
(1979) pointed out the question of whether the group of "I."rotundatus Fiege and "I." weltersdorfensis Andert should be
assigned to Inoceramus or Cremnoceramus. Besides thiswork, in some other papers (Kauffman, 1978a, b; Walaszczyk, 1988; Wiedmann and Kauffman, 1978), Cremnoceramus
is regarded as a full genus, and in still others (Cox, 1969;
Hattin and Cobban, 1977; Ivanikov, 1979; Trager, 1981a;
LOpez, 1989, 1990, 1992 and Kennedy and Cobban, 1991) as asubgenus of Inoceramus. In my opinion, it is adequate that
Cremnoceramus be ranked as a subgenus of Inoceramus
because there is a group of transitional forms which is hardly
assignable either to Inoceramus or Cremnoceramus, as Hermet al. (1979) mentioned. According to Heinz (1932), Cox (1969)and Herm et aJ. (1979), the geniculation of the shell is an
important criterion for Cremnoceramus. In contrast Trager(1981a) pointed out that the geniculation should not be used
for taxonomic purposes because it is a highly flexible featurevarying with mode of life.

Masayuki Noda

572

Inoceramus (Cremnoceramus) deformis Meek, 1871
Figures Ha-d, 2a-<:; 7-4
Inoceramus sp. ? Hall, 1845, p. 310, pI. 4, fig. 2.
Inoceramus deformis Meek, 1871, p. 296; White, 1875, p. 179, 180,

pI. 15, fig. 1; Meek, 1876, p. 146, pI. 14, figs. 4a, b; Stanton,
1893, p.85, pI. 15, fig.2; Heinz, 1928b, p.34, pI. 2, fig. 1,
Seitz, 1959, p. 118; Dobrov and Pavlova, 1959, p. 138, pI. 8,
fig. 3; Scott and Cobban, 1964, pI. 1 ; Trager, 1967, p. 130132, pI. 14, fig. 7; Hattin and Cobban, 1977, p. 191, 192, fig. 9 ;Ivanikov, 1979, p. 48, 49, pI. 6, figs. 2, 3; Cieslifinski andBlaszkiewicz, 1984, p.364, pI. 158, fig. 1; Szasz, 1985, p.161,
162, pI. 6, figs. 1a, b, 2a, b; Cieslinski and Blaszkiewicz,
1989, pI. 159, fig. 1 ; Tarkowski, 1991, p.107, pI. 15, fig. 1.
"Inoceramus" deformis deformis Meek, Kauffman, 1978a, p. IV-8,
list.
"Inoceramus" deformis n. subsp. Kauffman, 1978a, p. IV-8. list.
Inoceramus? deformis Meek. Kauffman, 1978b, p. XIII-2, list.
Inoceramus (?) deformis deformis Meek. Kauffman, 1977, p.
245, pI. 12, fig. 3.
Inoceramus (?) deformis n. subsp. Kauffman, 1977, p. 245, pI. 12,
fig. 2.
Inoceramus (Cremnoceramus) deformis Meek. Seibertz, 1986, p.
1177; Kennedy and Cobban, 1991, p. 9, 10, list.
Cremnoceramus deformis (Meek). Walaszczyk, 1988, pI. 8, figs.
1,2; Kopaevich and Walaszczyk, 1990, pI. 4, figs.4a, b;
Walaszczyk, 1992, p. 52, 53, pI. 29, fig. 4, pI. 30, fig. 4.
Cremnoceramus ? deformis (Meek). Walaszczyk, 1992, pI. 36,
fig. 4.
Holotype.-By monotypy, the specimen drawn by Meek
(1876, pI. 14, fig. 4) from the Niobrara Formation, Pueblo,
Colorado, but its precise locality is uncertain.
Material.-JG.H3016 from loc. Ob0012, JG.H3058 and 3060
from loc. Ob0013 in the Obira area, northwest Hokkaido.
Oescription.-Shell medium-sized, subequivalve and inequilateral. Considerably inflated from anterior to posterior
and also along growth axis sometimes with geniculation.
Anterior side fairly broad and steep or perpendicular tocommissure plane. Posterodorsal part also steep to hinge
without flattened wing-like area. Umbo terminal or subterminal, left one somewhat projected over hinge line andstrongly curved inwards and slightly twisted forwards. Rightumbo scarcely rising above hinge line. Growth axis broadly
concave to anterior. Anterior margin long, nearly straight or
broadly convex, but for a portion near umbo slightly concave; anteroventral margin somewhat narrowly bent. Ventral
margin asymmetrically rounded but subangularly bent at
ventral extremity (at intersecting point of growth axis toventral margin); posterior margin broadly arcuate, forming an
obtuse angle with hinge line which is about half shell length
on average. Hinge plate fairly thick and ligament pits are
deep and scooped with rounded and narrow interspaces,
according to terminology by Crampton (1988, p. 987, text-fig.
3 and p. 985, table 2).
Surface ornamented with major and minor sculpture in
combination; concentric ribs coarse, sharp- or roundtopped and irregular in size and interspaces, showing considerable extent of variation; concentric rings superimposedon ribs and interspaces, weak, round-topped and regular inbreadth and height. One or two radial elevations devel-

oped, one of which corresponds to growth axis or runs
somewhat posteriorly, anterior one demarcates anterior wallfrom main part of disk.
Biometry.-Measurements are shown in Table 1.
Individual relative growth of I vS. hand b vS. h is demonstrated in Figure 2, showing left and right valves of thespecimen JG.H3016. Relative growth of I vS. h shows
negative allometry in both valves, whereas that of b vS. h isregarded as positive allometry.
Ontogenetic changes in I/h, b/h and obliquity (0') are also
demonstrated in Figures 3A, Band C, respectively. The
simple ratio I/h decreases gradually while b/h and obliquity
increase with growth, b/h changes abruptly at a certain
stage of growth in some specimens, and obliquity is considerably variable among individuals.
Profiles of the specimens in two directions are shown in
Figure 4. Weak geniculation is perceptible in the verticalsection of the left valve of the specimens examined. The
cross sections are nearly circular, being usually convex or
bowl-shaped.
Remarks.-The specimen JG.H3016 (Figures Ha----d) consists of well preserved closed valves somewhat displacedalong the commissure plane. Two blunt radial elevations
develop on both valves. The anterior one demarcates indistinctly the anterior wall from the disk and posterior one runs
somewhat posteriorly along the growth axis. Therefore, theshell is subrectangular in general outline. Small hemispherical pits are scattered on the internal mold of the left valve.
These seem to be that the traces of blister pearls produced
to cover holes made by boring barnacles (Seilacher, 1969)
while the animal was alive.
Comparison and discussion.-JG.H3016 resembles closely
the holotype of I. (Cr.) paradeformis SZ8SZ (1985, pI. 17, figs.
1a-d) and other specimens figured in the same paper, fromthe lower Coniacian of the Babadag Basin, Romania, in its
trapezoidal to subquadrate outline, posterodorsal slope tohinge, ontogenetic change of obliquity, coarse and sharptopped concentric ribs and the development of two weakradial elevations. According to SZ8SZ (personal communication, June 15, 1993), "the main difference exists in the shapeof the umbo which is narrow in the Japanese specimen, with the
apex more pOinted and, in general, more highly raised above thehinge line. The shape of ribs in the first growing stage (up to 50
mm-60 mm distance from the apex of the umbo) is also different:
it is almost circular in the Japanese specimen, (cI') being about 85°
whereas in the specimens from Romania, the ribs are strongly
elongated toward the posterior part of the valve, (cI') varying
between 35°-45°. Even in speCimens of I. paradeformis withalmost circular ribs in the umbonal zone (Szasz, 1985, pI. 16, fig.
1) the umbo is more massive than that of the Japanese specimen." Furthermore, SZ8SZ (1985, p. 30) remarked that I.
paradeformis SZ8SZ, 1985, I. dobrogensis SZ8SZ, 1985 and I.
babadagensis SZ8SZ, 1985 from the Babadag Basin, Dobrogea, resemble closely one another and are obviously
related to I. deformis, all of them could constitute a single
biological species. But the three morphotypes are regarded
as three distinct morphological species until necessary information is available.
I. (Cr.) deformis was established on the basis of a Single

1020.

Inoceramids from Upper Cretaceous of Hokkaido, Part 2

Figure 1. Inoceramus (Cremnoceramus) deformis Meek, all figures natural size. 1a- d. JG.H3016 from lac.
Ob0012. 1d: anterior view (call. T. Shimanuki, 1988). 2a- c. JG.H3060 from loc.ob0013, 2a: anterior view (call. T.
Shimanuki and Noda, 1991).

S73

Masayuki Noda

574

Table 1. Measurements of Inoceramus (Cremnoceramus) deformis Meek from Hokkaido.
dimension in mm.
yHL
specimen
valve
h
I
b
s
a
0
L.
99.2
81.8
39.8
38.3
99.6 87.0 132 102
86
JG. H3016
R.
7704
7004
29.7
33.0
81.8 74.8 132 102" 76
29.2
52.0 59.3 123 103
74
JG. H3058
R.
51.8
53.6
14.0
JG. H3060
18.1
53.3 42.7 136 104
L.
53.3
45.1
19.2
8T
0

0

0

0

specimenvalve I/hb/hb/h ll =6omm
l/h H=6omm
L.
0.82
0.94
0.40
0040
JG. H3016
R.
0.91
0.86
0.38
0.31
JG. H3058
R.
JG. H3060
L.
0.85
0.36
For Abbreviations, refer to Noda (1996, table 1 and fig. 1).

A •

I b

mm

• JG.H3016L

B 0------<> JG.H3016R

100
90
80
70

1= 1.206ho. 926

A

r = 0.9988

60
50

1= 1.052ho. 957
r=0.9988

B

40

/
/
/

30

9/
20

f'

f'

/

"
'""

./;5

.

;/

/

7

/

/

'/
/

/

;f

/

'/
'/
/

'/

d

/

;/

10
9
8

P

/

"

;/
;/

/

'/

./

'/

/

/0

/

/

/

/0

6

/

/
/

5

A

/
/

,0

4

b = 0.129h 1.252
r= 0.9976//

b = O. 107h1.268

/

B

/

3

/

/

r= 0.9978

/
/

2

0/

/

/
7 8 910Figure 2.

20

30

40

50 60 708090 100

h mm

Diagram showing individual relative growth
between I and h, and band h of Inoceramus (Cr.) deformis(specimen JG.H3016).

L/H
0.87
0.91

s/I
0047
0047

-

-

0.80

0040

0

0

0

0

0

0

LinearOI'=6omm

84°

70
-

0

-

remarks

secondarily deformed
less than 60 mm in H

imperfect specimen from the lower Coniacian of the NiobraraGroup, Colorado. There has not been much information
subsequently, in regard to this species from the type locality
and adjacent areas, only some figures with no description(Hattin and Cobban, 1977; Kauffman, 1977) and biostratigraphic records by Scott and Cobban (1964), Hattin and Cobban(1977), Kauffman (1977, 1978a, b) and Kennedy and Cobban(1991). Thus the variability of the species is not yet clear atpresent. Collom (personal communication, 1993) is engaged
in research on this species on the basis of a large number of
specimens from several localities in Colorado. Thus theextent of variation of I. (Cr.) deformis will be made clear in thenear future.
The specimens of I. (Cr.) deformis figured by Hall (1845),
Meek (1876), Troger (1967), Kauffman (1977), Hattin andCobban (1977), Walaszczyk (1988,1992), SZ8.SZ (1985)
Kopaevich and Walaszczyk (1990) and Collom (personalcommunication, 1993) from the upper lower to lower middle
Coniacian of various regions around the world are considerably variable in general outline and surface ornamentation
but have the following in common: nearly equivalve, comparatively small beak, not so prominent umbo, vaulted shellwith geniculation, steep posterodorsal slope without distinct
wing-like area and the concave growth axis. Although alsovariable in outline and ornamentation, the present specimens
accord with some forms of I. (Cr.) deformis described or
figured previously in the other characters.
The specimen JG.H3016 is distinct from the hypotype
figured by Kauffman (1977). Collom (Feb. 20, 1993, personal
letter with the comment for the plaster cast of JG.H3016)
regarded it as a fine specimen of I. (Cr.) deformis (Cremnoceramus of his sense). If this assignment is pertinent, theallied species described from various regions may have to bereexamined taxonomically as SZ8.SZ (1985) remarked. I haveno intention of discussing that problem further because of
the insufficiency of the Japanese material.
As Walaszczyk (1992, p.55) regarded I. (Cr.) paradeformis
as a junior synonym of I. (Cr.) ernsti, the holotype of I. (Cr.)
paradeformis truly resembles the lectotype of I. (Cr.) ernsti
illustrated by Woods (1912a, text-fig. 85; =Walaszczyk,

1020.

Inoceramids from Upper Cretaceous of Hokkaido, Part 2
0 - - - -_ _ _

A

I/h
1.00

'.

0 - - - - ----~575
JG.H3016l

6-._._. __ .-...

JG.H3016R
JG.H 3060l

90

)00

110

Hmm

90

100

110

Hmm

90

100

110

Hmm

",\

\
0.95

\.

"\--..~---

0.90
0.85

20

10

30

.040

60

50

b/h

80

70

B

0 . .040---"'-.
~.

"-

/

"-

/

0.35

/

-..-.-

.-<f'

,
,

II'

,0
/

/
/

/

.o-_---...cf

0.30
0.25

/

,<>--

,,/

.P'

/

/

/

,,

"'fr"-------..()"

rf'/

20

10

30

.040

~

50

..

60

70

80

/
'

85
80

C

/

75

.,..- _.6

/
/

/

, /0

/

, ,If'

/
/
/

70

_ct'

,
,,

65
cf

,

/

r, '

" '0-

/--

~ __ --_..cr"'"

/

/

/

/

60

10

20

30

40

50

60

70

80

Figure 3. Diagram showing ontogenetic changes in selected characters of Inoceramus (Cr.) deformis.
A: I/h, B: b/h, C: obliquity.

1992, pI. 32, figs. 2a, b) in general outline and surface ornamentation, but I. (Cr.) ernsti has a concave anterior wall,
which is an essential character and an important criterion of
the species as Woods (1912a) originally remarked; the anterior wall of I. (Cr.) paradeformis is not so conspicuously
concave except for the umbonal region, as SZ8SZ (1985)

described. The two species are thus distinct from eachother.
The specimen of I. (Cr.) schloenbachi pratjei Heinz, 1928
figured by himself (1928b, pI. 6) was regarded as I. (Cr.)
deformis by Seitz (1956, p. 118), but Heinz's subspecies isdistinct from I. (Cr.) deformis in the moderately convex shellMasayuki Noda

576

A

B

20mm

20mm

JG.H3016L

JG.H3016L
p

a

v

u

20 mm

JG.H3060L
pa

Figure 4. Profiles of selected specimens of Inoceramus (Cr.) deformis in two directions. A: cross section at
the highest point of shell convexity, B: vertical section along the growth axis. a: anterior, p: posterior, u: umbo,
v: venter.

especially in the umbonal region, even though the anterior
part and left umbo, important characteristics of the species,
are missing.
Phyfogeny.-On the basis of the chronological change inshell size and surface ornamentation, Walaszczyk (1992, p.
53) regarded f. (Cr.) deformis as the last representative of thelineage of f. (Cr.) waltersdorfensis Andert-f. (Cr.) brongniarti
Mantell-f. (Cr.) deformis Meek. Besides, Kauffman (1978b)considered the lineage of f. (Cr.) deformis as f. (Cr.) rotundatus
Fiege-f. (Cr.) erectus Meek-f. (Cr.) deformis Meek-f. (Cr.)
schfoenbachi Bbhm. The Japanese material is too deficient,at present, to discuss the phylogeny of f. (Cr.) deformis.
Nevertheless the vaulted shell with blunt radial elevations
shown by JG.H3016 may suggest some relationships with I.
(Cordiceramus).
Occurrence.-For the locality map of the Obira area,

readers may refer to Noda (1992, p. 1312, fig. 1).
Loc. Ob0012 (= loc. NH78 of Tanaka, 1963). Topographic
map ''Takishita'', Scale 1 : 25000. A river floor of the Obirashibe, somewhat below Bridge Takimi-Ohashi, now below
the water level of artificial Lake Obirashibeko. Silty sandstone bed at the base of Unit Ub (Tanaka, 1963, Tsushima etaf., 1958). Associated with f. (f.) uwajimensis Yehara, f. (f.) aff.
pedalionoides Nagao and Matsumoto and f. (Cr.) ernsti Heinz.
For Loc. Ob0013. see Noda (1996, Part 1, p.569)
Range.-Locs. Ob0012 and 0013 are located in Unit Ub
which immediately overlies Unit Ua2 (Tanaka, 1963). Unit
Ua2 contains f. (Ct.) rotundatus which is a good cosmopolitan
zonal index of the lower lower Coniacian. Besides, one of
the associated species, f. (I.) uwajimensis, is a representativeof the upper lower to middle Coniacian of Japan and adjacent areas. Thus the stratigraphic position of f. (Cr.) deformis along the River Obirashibe is limited to upper lower and/or lower middle Coniacian. But the true range of thisspecies in Japan should be made clear on the basis offurther stratigraphic work in various regions.
Coniacian inoceramid zonation was previously proposedby Cobban (1951), Scott and Cobban (1964), Kauffman (1978a,b), Kauffman et af. (1978), Cobban (1986) and Szasz and Ion(1988), but there is disagreement among them. According to
the recent work by Kennedy and Cobban (1991) in theWestern Interior, f. (Cr.) deformis indicates practically thelower part of the middle Coniacian. Besides, Kopaevoch andWalaszczyk (1990) regarded the species as a representativeof the upper lower Coniacian of southwestern Crimea.
To sum up, opinions about the range of f. (Cr.) deformis
slightly differ among researchers due to time lag amongareas and/or different views of the specific definition, but therange largely falls within the period from lower to lower
middle Coniacian.

Inoceramus (Cremnoceramus) ernst; Heinz, 1928
Figures 5-a---c; 6-1a, b, 2a-d; 7 -1a---c, 2a---c, 3a-d
Inoceramus lamarcki Parkinson (pars).

Woods, 1912a, p. 325, fig.

85.
Inoceramus ernsti Heinz, 1928a, p. 73, 74; Trager, 1967, p.128-

130, pI. 14, figs. 1-6; Ivanikov, 1979, p. 51, 52, pI. 8, figs. 1, 2 ;Szeisz, 1985, p.172, pI. 29, figs.3a, b; Tarkowski, 1991, p.
108, pI. 14, fig. 5.

~

oI\)

p

:::J

o()

CD

03
3

0:

(j)

a3
c

-0
-0

~

o

~

()

CD

o

c(j)
~

I

o

7'
7'

OJ

0:

_0

IJ
OJ

;::::).

I\)<.h

Figure 5.

Inoceramus (Cremnoceramus) ernsti Heinz.

JG.H3023 from loc.Ob2100, x ca 0.5 (coil. T. Shimanuki, 1985).

-..)
-..)

578

Masayuki Noda

Inoceramus (Inoceramus) ernsti Heinz. Kauffman, 1977, p.242,
244, pI. 11, fig. 5; Kauffman et al., 1978, p.34, pI. 15, fig. 5.
Inoceramus (Cremnoceramus) ernsti Heinz. LOpez, 1990, p. 189-

193, pI. 5, figs.6a, b; LOpez, 1992, p.2-245-2-248, pI. 1,
figs. 4, 5a, b.
Cremnoceramus ernsti (Heinz). Walaszczyk, 1992, p. 55, 56, pI.
32, figs. 1-3.
Tethyoceramus (Protoceramus) ernsti (Heinz). Heinz, 1934, p.
250, pI. 19, fig. 1.
Compare.
Inoceramus cf. ernsti Heinz. Pergament, 1971, p. 119, 120, pI. 33,
fig. 2, pI. 34, fig. 3.
Lectotype.- The specimen hand-drawn by Woods (1912a,
p. 325, fig. 85) was designated as the lectotype by Trager(1967, p.128). The same specimen was photographed byWalaszczyk (1992, pI. 32, figs. 2a, b).
Material.-JG.H3017 from loc.ob0020, JG.H3059 from loc.
Ob0013, JG.H3061 from loc.ob0011, JG.H3203 from loc.
Ob2100 in Obira area, WE.P1312 and 1331 from loc.lk8617a in
Ikushunbetsu area.
Description.-Shell medium to large, slightly inequivalve,inequilateral, much higher than long. Valve considerably
convex from anterior to posterior and also along growth axis
which is straight or somewhat concave to anterior. Anteriorwall angularly bent from disk and deeply concave tocommissure plane, posterior and posterodorsal parts fairly
steep without forming a clearly defined wing. Two or threeblunt radial elevations developed, of which anterior one
demarcates sharply anterior wall from disk, median one
corresponds to growth axis and posterior one is indistinct orhardly perceptible. Umbo small and situated at anterior end,
left one more projected over hinge line than the right andcurved inwards. Anterodorsal margin long and straight,
anterior through ventral ones broadly or moderately rounded
and more or less acutely bent at ventral extremity where
median radial elevation intersect. Posteroventral to posterior ones broadly arcuate and forming an obtuse angle withhinge line which is shorter than half of the shell length. Hinge
plate fairly thick and ligament pits deep, rectangular andinterspaces with round ridge.
Surface ornamented with concentric ribs and rings incombination, the former are round-topped, variable in heightand irregular in breadth and broaden with growth, the latterare superimposed on ribs and interspaces.
Biometry.-Measurements are shown in Table 2.
Figure 8 gives individual relative growth of I vs. h of
JG.H3017 in left and right valves. The I vs. h reduced majoraxes show negative allometry in both valves and b vs. h
positive diphasic allometry with different critical points
between two valves.
Ontogenetic changes in I/h, b/h and obliquity (v) are
shown in Figures 9A, Band C, respectively. The simple
ratio I/h decreases gradually with growth, whereas that of b/h increases gradually or abruptly in some specimens; obliquity increases gradually in early stage of growth andbecomes constant in later growth stage.

Profiles of well preserved specimens in two directions areillustrated in Figure 10. Geniculation is clearly shown in
JG.H3017L and WE.P1321, and anterior wall is angularly bent
from the main part of disk and distinctly concave to commissure plane in all specimens.
Remarks.-JG.H3017L and 3017R (Figures 6-1,2) wereobtained as separate valves in a single nodule. These
probably belong to one and the same individual. These "arefairly well preserved with no secondary deformation but someeroded portions. The hinge structure is clearly observed in
the right valve (Figure 6-2b). The ligamental groove is fairly
broad with scooped pits and angular ridges. Geniculation
at a growth stage of 26 mm in H and a radial elevation
develop distinctly along growth axis. JG.H3203 is a hugeindividual, which was initially assigned to I. (I.) hobetsensis
nonsulcatus, but is distinct from the typical form of thatsubspecies in a deeply concave anterior wall, fairly crowded
ribs, blunt elevation along the growth axis and its higher
stratigraphic occurrence in the upper Turonian.
Comparison and discussion.-Inoceramus ernsti wasestablished by Heinz (1928b) on the basis of the specimen
figured by Woods (1912a). According to Woods (1912a), thespecimen drawn by himself was brought probably from thezone of Holaster planus of the Upper Chalk (upper Turonian),
but its locality is unknown. Trager (1967) designated Woods'
specimen as the lectotype. It is characterized by the
concave anterior wall as Woods (1912a) originally pointed out
and vaulted shell with distinct geniculation, which is an
important criterion of I. (Cremnoceramus). Trager (1967)
described precisely the specific characters and examined
the species biometrically on the basis of more than twenty
specimens from the lower middle Coniacian (= "oberen
Oberturon bis unteren Coniac" before Seibertz, 1979) of theSachsen (Saxony), Bahmen (Bohemia) and Brandenburg
regions.
Specific characters described by Trager (1967) agree withthose of the present specimens, especially in showing two
radial elevations, one of which clearly demarcates theconcave anterior wall from the convex disk as drawn byhimself (Trager, 1967, pI. 14, fig. 3) while the other runs along
the growth axis.
From the above comparison, the specimens available are
identical with I. (Cr.) ernsti Heinz.
The specimens of I. (Cr.) ernsti from the lower Coniacian of
the Babadag Basin of Romania figured by SZ8.SZ (1985, pI. 29,
figs. 3a, b) and also from the lower Coniacian of the NavarroCantabrian Basin of Spain figured by Lopez (1990, pI. 5, figs.
6a, b; 1992, pI. 1, figs.5a, b) closely resemble each other,and both specimens show a somewhat larger I/h in comparison with the Japanese ones.
The specimen from the uppermost Turonian of northwestern Kamchatka, figured by Pergament (1971, pI. 33, fig. 2, pI.
34, fig. 3) somewhat resembles the specimens of Japan in its
biconvex shell and concave anterior wall but has muchcoarser concentric ribs. Although the lateral view of the

Figure 6. Inoceramus (Cremnoceramus) ernsti Heinz. JG.H3017L and 3017R from loc.ObOO20, x 0.86. 1a, b. Leftvalve. 2a-d. Right valve, 2b shows hinge structure (coil. T. Shimanuki, 1989).

1020. Inoceramids from Upper Cretaceous of Hokkaido, Part 2579

Masayuki Noda
580

1020.
Table 2.
sion in mm.
specimen
JG. H3017

Linear dimen-

valve

h

I

b

s

a

y

0

L.

140.0

109.2

46.2

38.4

141.0 102.5

120"

140"

8T

83"

R.

109.0

95.9

33.8

35.9

111.6

121"

138"

81"

78"

H

L
90.0

OH-6omm

L.

52.1

45.3

25.5

19.8

57.6

36.4 136"

90°

86"

-

R.

34.7

30.1

19.1

15.0

35.4

29.5

128"

80"

-

specimen

valve

I/h

s/I

0.33

b/h"-6omm
0.26

L/H

0.78

l/h"=6omm
0.93

b/h

L.

0.73

0.35

R.

0.88

0.89

0.31

0.28

0.81

0.37

JG. H3059

L.

0.87

-

0.49

-

0.63

0.34

secondarily deformed
less than 60 mm in H

WE. P1321

R.

0.86

-

0.55

-

0.83

0.50

less than 60 mm in H

A

00-.- - - - - .

JG.H3017L

B ~-----~ JG.H3017R

100
90
80

1= 1.351ho. 895

A

r = 0.9980

B

r

50
40

Measurements of Inoceramus (Cremnoceramus) ernsti Heinz from Hokkaido.

JG. H3059

I. b
mm

60

58!

WE. P1321

JG. H3017

7

Inoceramids from Upper Cretaceous of Hokkaido, Part 2

I = 1. 024ho. 964=0.998530
20
b=0.028h1.522

B r=0.9838

10
9
8
7
6

A

b= O. 219h1.037
r= 0.9996
b=0.229hl.053

B r = 0.9992

/

/.

,:/'l

f
78910

20

30

40 50 60708090100

h mm

Figure 8. Diagram showing individual relative growth
between I and h, and band h of Inoceramus (Cr.) ernsti(specimen JG.H3017).

11T

remarks

specimen is not shown, his specimen may be rather comparable with I. (I.) iburiensis Nagao and Matsumoto, 1939.
Phylogeny.- The lectotype of I. (Cr.) ernsti was originallyregarded as a variety of I. lamarcki Parkinson which belongs
undoubtedly to I. (Inoceramus). Herm et al. (1979) pointed
out the ambiguity of whether I. rotundatus Fiege and thegroup of "I." waltersdorfensis Andert are assigned to I.
(Inoceramus) or to I. (Cremnoceramus), which suggests a
close phylogenetic relationship between I. (Inoceramus) andI. (Cremnoceramus). The development of blunt radial elevations in some specimens of I. (Cr.) ernsti may foretell thecharacters of I. (Cordiceramus) in the upper Coniacian andsubsequent stages. In other words, it may be a "transient"linking I. (Cremnoceramus) and I. (Cordiceramus). I. (Co.)
kawashitai Noda, 1986, from the upper Coniacian of someareas of Hokkaido and Kyushu is certainly a descendant of
I. (Cr.) ernsti. On the other hand, Trager (1969, p. 74) locatedI. (Cr.) ernsti as a linking species between I. (I.) lamarcki
sWmckei Heinz, 1926, from the middle Turonian and I.
(Volviceramus) koeneni Muller, 1887, from the middle
Coniacian (so-called lower Coniacian before Seibertz, 1979).
I agree with Trager's scheme (1969, p.74, fig.8) for thereason mentioned previously in the phylogeny of I. (Volviceramus) koeneni (Nod a, 1996).
The above discussion materializes a notion that I. (Cr.)
ernsti is important as a common ancestor of I. (Co.) kawashitai and I. (V.) koeneni.
Occurrence.-Loc. Ob0013. See Noda, 1996, p. 569, for the
locality of I. (V.) koeneni.
Loc. Ob0011. The opposite bank of lac. Ob0012 [see p. 576,
for the locality of I. (Cr.) deformis] and the same layer as loc.
Ob0012 which is stratigraphically about 20 m above loc.
Ob0013.
Loc.Ob0020. (=NH54 of Tanaka, 1963). Topographic map

Figure 7. 1a-c. Inoceramus (Cremnoceramus) ernsti Heinz. JG.H3063, X 0.86. Reconstructed plaster cast from JG.
H3017L and 3017R. 2a-c. Inoceramus (Cremnoceramus) ernsti Heinz. WEP1321 from loc.lk8617a. Natural size (coil. A.
Asai). 3a-d. Inoceramus (Cremnoceramus) ernsti Heinz. JG.H3059 from loc.Ob0013, natural size (coil. T. Shimanuki and M.
Noda, 1991). 4. Inoceramus (Cremnoceramus) deformis Meek. JG.H3058, same locality. Natural size (coli. T. Shimanuki
and M. Noda, 1991).

Masayuki Noda

582

A

1/ h

JG.H3017L
0--------0

1.00

JG. H3017R
JG.H3059L

0.95
0.90

--0--

0.85

0.80

,

5'0

40

70

60

t,·

b/h
i

120

90

130

Hmm

B

i

0.45

8'0

i

i

i

I

i

0.40

(

i
I

0.35

/

0.30

95

/ ..... ,

11(/

i

".

.i

--

"~ ._._.1-- ............0 - - __ -0- ___ -<:;-----..0..... .................

cr-- . . / ---0. . . .

0.25

0

/

0_

,

,

10

20

,

,

i

40

50

60

i

30

,-

,

80

70

I

...0---

,

i

90

100

,

,

110

120

130

Honm

110

120

,
130

Hmm

i

C

90
85

r'-'-

.................

--10-._~

80
75

-/

, ,,0', , ...

/... A ..... _.....

____ o____ cr __. . . .0---""""""-O-'----CI'--..L-4
/

Jt

10

20

30

40

50

60

70

80

90

100

Figure 9. Diagram showing ontogenetic changes in selected characters of Inoceramus (Cr.) ernsti.
B: b/h, C: obliquity.

''Takishita'', Scale 1 : 25000. The right bank of the River
Obirashibe, about 50 m below the confluence of theJugosen-zawa, below the surface of the lake at full waterseasons. Silty sandstone bed of Unit Ub. Associated withI. (I.) uwajimensis and I. (PI.) troegeri.

A: I/h,Loc.Ob2100. Topographic map, ditto. A cliff of the River
Kamikinenbetsu at the confluence with the Shigeruno-sawa.
Sandy siltstone bed of Unit Mo (Tanaka, 1963). Frequently
associated with huge individuals of I. (I.) iburiensis.
Loc.lk8617a For the locality map, see Noda (1996, fig. 9).

1020.

Inoceramids from Upper Cretaceous of Hokkaido, Part 2A

B

J G. H3059L
p

583

JG.H3059L

20mm

a

u

20mm

~3017R ~20mm

JG.H3017L

JG.H3017L

p

a

v

u

20mm

WE.P1321Ra

WE.P1321R
pv

JG.H3203L
40",,,,JG.H3203L

Rgure 10. Profiles of selected specimens of Inoceramus (Cr.) ernsti in two directions.
highest point of shell convexity. B: vertical section along the growth axis.

A: cross section at the584

Masayuki Noda

1020.

Inoceramids from Upper Cretaceous of Hokkaido, Part 2c .

Topographic map "Katsurazawako", Scale 1 : 25,000. A cliff
I.b
on a forestry road along the Okuhidarimata-zawa (so-called I b mm
80
Okusamata-zawa), about 450 m west of the fork along the70left branch. Fine sandy siltstone bed of IIIb member (Mat60
sumoto, 1965) of the Upper Yezo Group. Associated with f.
50(f.) uwajimensis.
Range.-The specimens examined occurred in the upper
40
Turonian to the lower middle Coniacian. This is in good
agreement with its range in Europe.
30

Inoceramus (Cremnoceramus) lueckendorfensis

I _1.109h o. 947

C

Dobrov and Pavlova, 1959, p.

10
9
8
7

Inoceramus inconstans lueckendorfensis Trager, 1967, p.102-105,

pI. 11, figs. 1a-c; Tarkowski, 1991, p.109, pI. 13, fig. 8.
Inoceramus "inconstans" lueckendorfensis Trager. Trager, 1981a,
p. 102, 106, 107, pI. 1, figs. 8-11.
Trager.

inconstans

<

/.
/.

/.

/.

I = 0.880h 1• OI8
r= 0.9989

D

/.

/.

20

137, pI. 5, figs.1a, b.

(Cremnoceramus)/.
/.

r = 0.9995

Figures 11-1a---c, 2a---c

Inoceramus. JG.H3097

D <>-------0 JG.H3098

Trager, 1967

Inoceramus inconstans Woods.

585

,

,,

,

.

,

/.
/.

/.

6
5

lueckendorfensis
4

Trager, 1989, p. 917, 918, list.

Hofotype.-Arbeitanr. 92F (frager, 1967, pI. 11, fig.1) byoriginal designation.
Materiaf.-JG.H3097 and 3098 from loc.Ob0003f.
Oescription.-Shell large-sized, subequivalve and inequilateral, highly convex from anterior to posterior and also
along growth axis with geniculation. Anterior part steep or
perpendicular to commissure plane and concave near umbo.
Posterior part sloping steeply and posterodorsal one almost
perpendicular to hinge line. Small posterior wing developed.
Umbo small, situated at anterior end, slightly projectedbeyond hinge line and twisted forwards. A blunt elevation
developed along growth axis. Anterodorsal margin long andstraight; anterior one circularly rounded up to midventer
where narrowly or subangularly bent and continuing tobroadly arcuate posterior margin; posterodorsal margin longand straight, forming an obtuse angle with hinge line which
is more than half of the shell length.
Surface ornamented with concentric ribs and rings incombination, of which the former are considerably variable inshape and intensity, that is, sharp-topped or round-topped,

320

7 8 91030405060708090100

Figure 12. Diagram showing individual relative growth
between I and h, and band h for selected specimens of
Inoceramus (Cr.) lueckendorfensis.

irregular in size, interspacing and intensity, while the latterare superimposed on ribs and interspaces and regular inbreadth and intensity.
Biometry.-Measurements of two specimens are shown in
Table 3. Figure 12 shows individual relative growth of JG.
H3097 and 3098. The reduced major axes of I vs. h show

Table 3. Measurements of Inoceramus (Cremnoceramus) lueckendorensis Trager from Hokkaido.
Linear dimension in mm.

specimen
JG. H3097
JG. H3098
specimen
JG. H3097
JG. H3098

valve
h
L.
105.7
L.
102.8
valve
L.
L.

I/h
0.84
0.87

I
89.2

hmm

s
45.0

89.1

b
45.4
42.0

l/h H =6omm
0.90
0.93

b/h
0.43
0.41

b/h"=6omm
0.24
0.29

37.8

L
H
107.6 87.5
106.1 86.4
L/H
0.81
0.81

a

y

<1

~'=6omm

118°

120°

7T

118

-

73°

72°
66°

0

s/I
0.50
0.42

Figure 11. Inoceramus (Cremnoceramus) lueckendorfensis Trager. All figures natural size. 1a-c. JG.H3097 from loc.
Ob0003f. 2a-c. JG.H3098, same locality (coil. T. Shimanuki, T. Matsumoto and M. Noda, 1993).

Masayuki Noda

586

A

I/h
1. 00

o------e.

JG.H30971

0--------0

JG.H309SL.. "",IJ ........
'",--'"0...

0.95................. _----0

,

'"

/

'-0,.

,,

,,

,
'0,

0.90

"

'0

0.S5

b/h
0.40
0.35
0.30
0.25

75
IP-~

70

/

~.o

/

,/
0 - - - ---0------0....

65

/'

,,/

----cf

I
I

60

I

I
I

/

"

55

10

20

3040

50

60

70

SO

90100110

Hmm

Figure 13. Diagram showing ontogenetic changes in selected characters of Inoceramus (Cr.) lueckendorfensis.
A: I/h, B: b/h, c: obliquity.

negative allometry for JG.H3097 and isometry for JG.H3098,
and those of b vs. h show a negative allometry in young
stage of JG.H3097 and isometry in the same stage of JG.
H3098, while both turn to positive allometry in later stages.
Critical points are different between the two specimens: 70
mm in H for JG.H3097, and 40 mm in H for JG.H3098.
Ontogenetic changes in I/h, b/h and obliquity ( gun) are
shown in Figure 13A, 8 and C, respectively. As is clear from

Figure 13A, the simple ratio I/h decreases gradually withgrowth, whereas, as shown in Figure 138, the simple ratio b/hincreases abruptly at a certain stage of growth which varies
from individual to individual. As a matter of course, convexity abruptly changes at the critical point of individual relative
growth b vs. h in each specimen.
Profiles of the specimens in two directions are illustrated in
Figure 14. In the cross section, somewhat angular eleva-

1020.

Inoceramids from Upper Cretaceous of Hokkaido, Part 2

A

B

JG.H3097L

p

587

20mm

a

JG.H3097L
u

v

20mm

JG.H3098L

JG.H3098L/

P ______________________-aJ

L_~

u

v

Figure 14. Profiles of available specimens of Inoceramus (Cr.) lueckendorfensis in two directions.
section at the highest point of shell convexity, B: vertical section along the growth axis.

tions are perceptible on the flank and in the vertical section,
geniculation in various stages of growth is clearly perceptible.
Remarks.-JG.H3097 and 3098 are fairly well preserved,
with the original outline being a pentagonal margin in general
aspect and the shell convexity, but no precise surface
ornamentation is known because the outer shell layer is lost.
Comparison and discussion.- The pentagonal margin,
vaulted shell with geniculation, development of radial elevation along the growth axis and the presence of a blunt
depression on the posterior flank are common in the present
specimens and the holotype of I. (Cr.) lueckendorfensis
Trager. Moreover, profiles in two directions of the specimens
agree well with those of the holotype (Trager, 1967, p. 104, fig.
27 above; pI. 11, figs.1a and c). The original description
and figures of Trager (1967) show that I. (Cr.) lueckendorfensis
exhibits a considerable extent of variation in outline, shellconvexity and surface ornamentation. The above comparison leads to the conclusion that the present specimens
surely fall within the range of variation of I. (Cr.) lueckendorfensis. Trager (personal letter, May 10, 1993) suggested tome that the Japanese specimens accord well with the typespecimens in outline but are distinct in surface ornamentation.
The present species was originally erected as a subspecies of I. inconstans by Trager (1967). Subsequently he(1981a) listed it as I. "inconstans" lueckendorfensis without
any comment. I think that I. inconstans Woods, 1912a is aheterogeneous species, about which specific definitions
among researchers can be expected to vary. Trager (personal communication, 1988) treated his specimen No. p-6 asan independent species, I. lueckendorfensis, thus I. (Cr.)
lueckendorfensis (personal letter of May 10, 1993). I agreewith him in this treatment.
Phylogeny.- The specimen (JG.H3077 from loc.lk2799) of

A: cross

I. (Cr.) rotundatus resembles the specimens No. p-6 of
Warsaw University from the lower Coniacian of Kolonkh and
RT1062 from the upper lower Coniacian of the Opole Trough,Poland, figured by Tarkowski (1991, pI. 13, fig. 8). On theroute along the Ponbetsu-gono-sawa, loc.Ik2799 is locatedsomewhat above the horizon of loc.lk2798 (= Ik2726, see
Matsumoto and Noda, 1985), which immediately overlies the
Tu/Co boundary containing I. (Cr.) rotundatus. The morphological similarity and stratigraphic relation may imply aphylogenetic relationship between the two species.
Besides, the holotype of I. (Co.) cordiformis purus Seitz,
1961 from the upper Santonian of the LOneburg area, Germany, is similar to the holotype of I. (Cr.) lueckendorfensis in
the pentagonal outline, vaulted shell, sharply demarcatedand broad anterior wall and the development of radial
elevations. The former is, however, discriminated from thelatter in the stronger radial elevations, irregular concentricribs and age of occurrence. The above fact may suggest
some relationship between I. (Cremnoceramus) and I. (Cordiceramus).
Occurrence.-Loc.ob0003f. See Noda (1992, p.1235).
Range.- The specimens handled were obtained from alimited horizon of the lower lower Coniacian of the Obiraarea, Hokkaido, but its stratigraphic range is yet to beclarified. Trager (1981b, 1989) located it in the lower lower
Coniacian. The range of the species in Japan is in good
agreement with that of Germany. Tarkowski (1991) recorded
it from the lower Coniacian of the Opole area, Poland, andfrom the upper Turonian to Coniacian of the Caucasus.

Conclusions
1. The examined specimens from the upper Turonian tolower middle Coniacian of Hokkaido are all identical orcomparable with some species hitherto described, that is, I.

Masayuki Noda

588

(I.) lusatiae Andert, I. (V.) koeneni MOiler, I. (Cr.) deformis
Meek, I. (Cr.) ernsti Heinz and I. (Cr.) lueckendorfensis Trager.

They are well known abroad but this is their first recorded inJapan.
2. Some specimens with intermediate or transitional
characters are important for delineating the phylogenetic
relationships among the following subgenera: I. (Inoceramus), I. (Cremnoceramus), I. (Cordiceramus) and I. (Volviceramus). The discussion of each species leads to an argument in favor of Trager's scheme (1969) and implies, inaddition, that I. (Cordiceramus) may have been derived fromI. (Cr.) ernsti, which is assumed to be the common ancestorof I. (Cordiceramus) and I. (Volviceramus).
3. With respect to taxonomic ranks, Inoceramus, Cremnoceramus and Volviceramus are each regarded herein as asubgenus of Inoceramus. Generally, it may be more reasonable that those species groups which comprise some transitional species with wide individual variation are treated assubgenera than full genera.
4. The species concerned are important for internationalcorrelations and elucidating paleogeography in early
Coniacian time.

Acknowledgments
I express my sincere thanks to Professor Emeritus Itaru
Hayami of the University of Tokyo for his critical reading of
the first draft, and to Professor Emeritus Tatsuro Matsumotoof Kyushu University for his help in the field and laboratory
work and giving instructive suggestions. I wish to thankProfessor Dr. Karl-Armin Trager of Bergakademie Freiberg,
Germany, Dr. Ladislau Szasz of the Central Department of
Geology, Romania and Dr. Christopher J. Collom of Calgary
University of Canada for their courtesy in giving me valuablecomments on I. (I.) lusatiae, I. (Cr.) deformis and I. (Cr.)
lueckendorfensis. I wish to thank Dr. Terufumi Ohno of
Kyoto University who kindly provided me with useful literature
and also Dr. William A. Cobban of the U.S. Geological
Survey for his kindness in providing me with several papers
unavailable in Japan. I am much indebted to Messrs.
Toshio Shimanuki of Kurisawa and Kenji Sanada of Sapporo
for their help in the field work and generous supply of anumber of valuable specimens for this study and also to
Messrs. Takemi Takahashi of Mikasa and Shigehiro
Uchida of Iwamizawa for their cooperation in the field.
Thanks are extended to Professor Dr. Trager and Dr. Akito
Asai of Waseda University who kindly sent me some models
of useful specimens. I thank Dr. Kenichi Saiki of the Mikasa
Museum (now Chiba Museum) for his generosity in offering
facilities for this study and also the staff members of the
Ikushunbetsu Branch of Iwamizawa Forestry Office for providing facilities for the field work.

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Masayuki Noda

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Stanton, T.w., 1893: The Colorado Formation and its invertebrate fauna. United States Geological Survey, Bulletin, no. 106, p.3-288, pis. 1-45.
Stoliczka, E., 1871: Cretaceous fauna of southern India.
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Asahikawa mill, Ashiyachi-zawa 'g~t~rR, Banno-sawa fiO)rR, Bungo :a~, Chiba T~, Fukagochi ~tJiJpg, Hatsudenshono-sawa 9'61tf9T 0) rR, Hidarimata-zawa ft:JBtrR, Ikushunbetsu ~w
5.lU, Inari-zawa filIfo1rR, Iwamizawa *Jl!.rR, Kamiichino-sawa ..t.-O)rR, Kamikinenbetsu ..t.~d~5.lU,
Katsurazawako flrRmJ, Kikumen-zawa §iliiIDrR, Kumaoi fl~~, Kumaoi-zawa fl~~rR, Kurisawa ~rR, Mikasa =~, Nishikatsurazawa ifflflrR, Obira 'PJl, Obirashibe 'P¥~, Obirashibeko 'PJl~
mJ, Oita *9t, Onogawa *lfJII, Okuhidarimata-zawa (Okusamata-zawa) ~ft:JBttR, Ponbetsugono-sawa 'J!F5.lU1i 0) rR, Sapporo ;fL~, Shigeruno-sawa v?' )vO)rR, Takimi-Ohashi i'itJl!.*~,
Takishita i'it~, Tano EHlf, Tappu )j;frj, Uwajima ~fD~, Waseda .!f!.fIllEH, Yezo I!~, Yoshiashizawa )'j''grR, Yubarigoe-zawa :Y'~~rR

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 184, pp. 592-596, 3 Figs., December 3D, 1996
© by the Palaeontological Society of Japan 1996

Latest Permian ammonoid Paratiroiites from the Ofunato
district, Southern Kitakami Massif, Northeast Japan
MASAYUKI EHIRO
Institute of Geology and Paleontology, Graduate School of Science, Tohoku University,
Sendai, 980- 77 Japan

Received 21 April 1996; Revised manuscript accept 5 September 1996
Abstract The latest Permian ammonoids, Paratirolites compressus sp. nov. and P. sp., are describedfrom the Permian strata in the Ofunato district of the Southern Kitakami Massif, Northeast Japan. Thisis the first record of Late Permian fossils from the Ofunato district.
Key words: Paratirolites, uppermost Permian, Kitakami Massif
Introduction

y. Onuki's collection of Kitakami fossils stored in theInstitute of Geology and Paleontology, Tohoku University,
includes one ammonoid and one nautiloid (?) collected by T.
Sugiyama. The attached labels indicate their occurrences
somewhere near Maeda (Figure 1), Ikawa- cho, Ofunato City,but no formation name. The ammonoid is labeled as
Gonioclymenia, which is one of the typical genera of latest
Devonian clymeniids.
On the other hand, Sugiyama (1942) reported the occurrence of Clymenia (=Clymenia spp. of Sugiyama, 1944?) in
the Devonian "Omori Series" (= upper Nakazato Series) in the
Ofunato district of the Southern Kitakami Massif and correlated the series to the Famennian. These clymeniids areimportant for geochronologic consideration of the "Omori
Series", because the series is now generally correlated with
the Middle Devonian. They have hitherto, however, beenneither described nor figured. Ammonoid fossils are rather
rare in Paleozoic strata of the Kitakami Massif, but theMaeda specimen may not be one of those listed by
Sugiyama (1942), because the clymeniids (?) of the "Omori
Series" were collected from a locality at Choanji or Omori,
Hikoroichi- cho, Ofunato City (Sugiyama, 1944), both far from
Maeda (Figure 1). Moreover, reexamination proves that theMaeda specimen is not a clymeniid. It has, in fact, a typicalceratitic suture (Figure 3c) and is considered to be a Late
Permian ammonoid, Paratirolites. This paper describes it as
Paratirolites compressus sp. nov. and discusses its stratigraphic significance. Paratirolites sp. recently collected by theauthor near Maeda is also described.
Geologic settings
On the geologic map of Yabe and Sugiyama (1937), thestrata distributed near Maeda are aSSigned to the Upper

Carboniferous- Middle Permian Maiya Group, which lies infault contact with the Devonian Nakazato Group on thenorthwest. Onuki (1956, fig. 2) treated them as Permian(possibly Lower to Middle Permian) and Minato et al. (1979,
fig. 2k-1) as the Middle Permian Kanokuran Series, although
no formation name had been given. Near Maeda, the Permian strata strike NNW-SSE to NW- SE and dip 40 to 80W.
Observation in the field shows that they are composed
mainly of black to dark gray shale, with subordinate amountsof sandy shale, sandstone and conglomerate. The grayshales, which are similar in lithology to that containing thepresent ammonoid, crop out in the cuttings along the railroadof Iwate Kaihatsu Tetsudo just north of Maeda (Figure 2, Loc.
A). They yield some brachiopods, pelecypods, bryozoans
and crinoid stems. Although no additional ammonoids havebeen collected from there and the age of Paratirolites little
differs from the previous geochronology, as mentioned later,
Sugiyama's specimen of Paratirolites is considered to havebeen collected from Loc. A in Figure 2 or its environs. I
also collected the same fossils as from Loc. A and anammonoid Paratirolites sp. from sandy shales intercalated inblack shales at Loc. B, a road cut on the Sanriku Highway(under construction), about 200 m east of Loc. A.
Stratigraphic significance of Paratiro/ites
Species of Paratirolites are index fossils of the uppermost
Permian. They are known extensively: Dorashamian strata
of Transcaucasia (Ruzhentsev and Sarycheva, 1965; Teichert et aI., 1973, and others) and Central Iran (Bando, 1979),
Changxingian formations of South China (Zhao et al., 1978;
Liang, 1983, and others), undeterminate horizon of Madagascar (Tozer, 1969) and latest Permian Pa/aeofusulina-bearing
bed of Thailand (Ishibashi and Chonglakmani, 1990). One ofthem was also reported, though its generic position is somewhat questionable, from the Senmatsu Formation in the1021.

Latest Permian ammonoids from Kitakami

593

7

6
+

+

+ +
+ .
+ +

5

+ +
+ + +

4

+Omori
+ +

+ + +
+ + +
+ + +
+ + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + + ++ + + + ++ + + + ++ + + + ++ ~ + + + ++\+ + + ++ + + + + ++ + + + + + + +~+ + ++ + + + + + + + + + + +++++++++++r------I+ + + + + + ++ + ++ + + + + + + ~ + + ~+ + + + + + + + + + ++ + + + + + + + + +

3

1 km

Figure 1. Simplified geologic map of the northern Ofunato district (compiled from various sources), withsupposed ammonoid locality. 1. Pre-Silurian Hikami Granites, 2. Silurian, 3. Devonian, 4. Carboniferous, 5.
Permian, 6. Lower Cretaceous, 7. ammonoid locality.

Okago district of the Southern Kitakami Massif (Ehiro andBando, 1985).
Upper Permian formations are widely distributed in thesouthern part of the Southern Kitakami Massif and ones
correlative with the Changxingian (Dorashamian) are disclosed in the northern Kesennuma (Tazawa, 1975; Ishii etal., 1975), Utatsu (Murata and Shimoyama, 1979), and probably Okago (Ehiro and Bando, 1985) districts. To date, noreliable Upper Permian fossil has been found from theOfunato district. The present ammonoid Paratirolites is thefirst record of an Upper (uppermost) Permian fossil from there
and the shales distributed near Maeda are the uppermost
Permian, not the Lower or Middle Permian as previously
thought. The naming of this Paratirolites-bearing strata near
Maeda is left pending, until the stratigraphy of the strata isfurther studied in detail.

Description of species
Family Dzhulfitidae Shevyrev, 1965
Genus Paratirolites Stoyanow, 1910
[Type species: Paratirolites kittli Stoyanow, 1910J

Remarks.-Shevyrev (in Ruzhentsev and Sarycheva, 1965)
established the family Dzhulfitidae to contain three genera:

Dzhulfites Shevyrev, Abichites Shevyrev and Paratirolites
Stoyanow. All these three genera have evolute whorls withlateral ribs and ventrolateral nodes, and a subquadrate to
subtrapezoidal cross section. They also have a uniquesuture with low saddles separating two denticulate laterallobes. Teichert and Kummel (in Teichert et aI., 1973) placed
the genera Dzhulfites and Abichites in Paratirolites based on
their morphological similarities. According to Tozer (1979),
the species of the genus Schizoloboceras described by Zhao
et al. (1978) from the Changxingian beds of South China may
also be included in the genus Paratirolites.
Paratirolites compressus sp. nov.
Figures 3a-c

Material.-A fragmental specimen, IGPS coil. cat. no.
103144 (Holotype), collected by Toshio Sugiyama from theunnamed strata distributed near Maeda, Ikawa-cho, Ofunato
City, Iwate Prefecture.
Diagnosis.-Paratirolites with compressed shell form and alarge umbilicus.
Description.-The specimen consists of parts of two volutions: a phragmocone and a body whorl. It is moderately
large and is considered to attain a diameter of at least 75
mm. The conch is evolute and thinly discoidal in outline,

594

Masayuki Ehiro
pI. 24, figs. 2-3 described as Abichites stoyanowi, p. 180, pI.
23, fig. 4 described as A. mojsisovicsi, p.181, pI. 24, fig. 4
described as A. abichi ; Teichert and Kummel in Teichert etaI, 1973, p.414, pI. 6, figs. 8-9, pI. 7, figs. 1-2, 6, 7; Bando,
1979, p.135, pI. 5, figs. 2, 6, pI. 6, fig. 1, pI. 8, figs. 1, 3) andsomewhat resembles the present species. But the Kitakami
species is more compressed than P. mojsisovicsi. The
present species is also distinguished from the latter and theother species in having a wider umbilicus.
Etymology.-The specific name is derived from its compressed shell form.

Paratirolites sp.
Figures 3d-e

Figure 2. Location map of Permian fossil localities(Locs. A and B) near Maeda, Ofunato City. Base map adopted from Quadrangle "Sakari", 1 : 25,000 scale topographic mapof Japan, Geographical Survey Institute.
with a wide umbilicus. The ratio of the umbilical diameter toshell diameter is estimated to be about 0.48 in the mature
stage. The specimen has a compressed shell form, though
deformed obliquely. The flanks are nearly flat and veryslightly converge toward the umbilicus, with the maximum
width at the ventral shoulder. The ratio of width to height at
the last whorl (living chamber) ranges from 0.3 to 0.4. The
venter is slightly convex. The cross section of the conch issubquadrate, with acutely rounded to edged umbilical andventral shoulders. The sides of the whorl are ornamented
by radial ribs. They are rather short, high and widely
spaced, with maximum height near the center of the side, on
the phragmocone. On the body whorl they are rather
closely spaced, extend from the umbilical to ventral shoulder,becoming wider toward the venter, and are nodose at theventral margin.
The suture is ceratitic, consisting of a large and roundedventrolateral saddle, a considerably low lateral saddle separating two strongly denticulated lateral lobes, and a high androunded second lateral saddle. The ventral part of thesuture is not observed.
Comparison.-Among the species of the genus Paratirolites, P. mojsisovicsi (Stoyanow) has rather compressed shellform, the ratio of width to height ranges from 0.6 to 0.8 (forexample: Shevyrev in Ruzhentsev and Sarycheva,1965, p.
178, pI. 23, figs. 2-3 described as Paratirolites dieneri, p. 179,

Material.-IGPS coil. cat. no. 103276, collected by M. Ehiro
from the unnamed strata distributed near Maeda, Ikawacho, Ofunato City, Iwate Prefecture.
Descriptive remarks.-An outer mould and a small fragmental inner mould of a moderately large specimen were
examined. It attains a diameter of 75 mm. The conch isevolute and has a wide umbilicus. The ratio of the umbilical
diameter to shell diameter is about 0.4 in the inner volution,
but attains 0.47 in the mature stage. The flanks are flat andthe venter is slightly convex. The conch is subquadrate incross section, with acutely rounded umbilical and ventral
shoulders. The ratio of width to height at the last whorl is O.
6 to 0.7, although it is smaller in the inner mould. The sideof the whorl is ornamented by fine radial ribs. They arewidely spaced on the inner whorls, but closely spaced in themature stage.
In having an evolute and compressed shell form with
subquadrate conch section and radial ribs, the present
specimen is considered to belong to the genus Paratirolites
and to be similar to P. mojsisovicsi (Stoyanow) (for example:
Shevyrev in Ruzhentsev and Sarycheva, 1965, p. 178, pI. 23,
figs. 2-3 described as Paratirolites dieneri, p. 179, pI. 24, figs.
2-3 described as Abichites stoyanowi, p. 180, pI. 23, fig. 4
described as A. mojsisovicsi, p.181, pI. 24, fig. 4 described as
A. abichi; Teichert and Kummel in Teichert et al., 1973, p.
414, pI. 6, figs. 8-9, pI. 7, figs. 1-2, 6, 7; Bando, 1979, p.135,
pI. 5, figs. 2, 6, pI. 6, fig. 1, pI. 8, figs. 1, 3) and P. compressum
sp. nov. It is, however, difficult to identify it at the specificlevel, owing to the rather poor state of the preservation.
Moreover, the generic position is somewhat questionable,
because the ventrolateral nodes, characteristic for the genus
Paratirolites, seem to be obscure or missing in the present
species and the suture is not known.
Acknowledgments
The author thanks Dr. Yoshio Onuki for permission to
study his collection. I am also deeply appreciated to Professor Kunihiro Ishizaki of Tohoku University for his review ofthe manuscript. This research was partly supported by aGrant-in-Aid for Scientific Research (C: 07640593) from the
Ministry of Education, Science and Culture of Japan.

1021.

Latest Permian ammonoids from Kitakami

595

cFigure 3. Paratirolites from Maeda, Ofunato district. a- c: Paratirolites compressus sp. nov., IGPS coil. cat. no.
103144, lateral view (a, x 1), cross section (b, X 1) and suture line (c, x 4). d- e: Paratirolites sp., IGPS coil. cat. no.
103276, lateral view of the rubber cast of the outer mould (d, X 1) and cross section (e, X 1).

References cited
Bando, Y., 1979 : Upper Permian and Lower Triassic ammonoids from Abadeh, central Iran. Memoir of theFaculty of Education, Kagawa University, Series II , vol.
29, p. 103-138.
Ehiro, M. and Bando, Y., 1985 : Late Permian ammonoids
from the Southern Kitakami Massif, Northeast Japan.
Transactions and Proceedings of the PalaeontologicalSociety of Japan, New Series, no. 137, p. 25- 49.
Ishibashi, T. and Chonglakmani, C., 1990 : Uppermost Permian ammonoids from northern Thailand. Journal ofSoutheast Asian Earth Sciences, vol. 4, p. 163-170.
Ishii, K., Okimura, Y. and Nakazawa, K., 1975 : On the genus
Colaniella and its biostratigraphic significance. Journalof Geosciences, Osaka City University, vol. 19, p. 107138.
Liang, X. , 1983 : New material of Permian ammonoids withdiscussion on the origin, migration of Araxoceratidae
and the horizon of the Paratirolites. Acta Palaeontologica Sinica, vol. 22, p. 606- 615. (in Chinese withEnglish abstract)
Minato, M., Hunahashi, M., Watanabe, J. and Kato, M. eds.,

1979: The Abean Orogeny, Variscan Geohistory ofNorthern Japan. 427 p. Tokai University Press, Tokyo.
Murata, M. and Shimoyama, S., 1979 : Stratigraphy near thePermian - Triassic boundary and the pre- Triassic unconformity in the Kitakami Massif, Northeast Japan.
Kumamoto Journal of Science, Geology, vol. 11, p. 11 - 31.
(in Japanese with English abstract)
Onuki, Y., 1956 : Explanatory text of the geological map of
Iwate Prefecture, Japan (scale 1 : 100,000). II. Geologyof the Kitakami Massif. 189 p. Iwate Prefecture. (inJapanese)
Ruzhentsev, V.E. and Sarycheva, T.G., 1965: Development
and change of marine organisms at the PaleozoicMesozoic boundary. Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR, vol. 108, p. 166- 182. (inRussian)Sugiyama, T., 1942 : Devonian strata in eastern Asia. Journal of the Geological Society of Japan, vol. 49, p. 260262. (in Japanese with English abstract)
Sugiyama, T., 1944 : On the trilobites from Japan. Studies
from the Geologic and Mineralogic Institute, Tokyo
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Masayuki Ehiro

Tazawa, J., 1975: Uppermost Permian fossils from theSouthern Kitakami Mountains, Northeast Japan. Journal of the Geological Society of Japan, vol. 81, p.629640.
Teichert, C., Kummel, B. and Sweet, w., 1973: PermianTriassic strata, Ku-E-Ali Bashi, northwestern Iran.
Bulletin Museum of Comparative Zoology, vol. 145, p.
359-472.
Tozer, E.T., 1969: Xenodiscacean ammonoids and their
bearing on the discrimination of the Permo-Triassic
boundary. Geological Magazine, vol. 106, p.348-361.
Tozer, E.T., 1979: The significance of the ammonoids Paratirolites and Otoceras in correlating the Permian-Trias-

sic boundary beds of Iran and the People's Republic ofChina. canadian Journal of Earth Sciences, vol. 16, p.
1524-1532.
Yabe, H. and Sugiyama, T., 1937: Preliminary report on thefossiliferous Gotlandian and Devonian deposits newly
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Zhao, J., Liang, X. and Zheng, Z., 1978: Late Permiancephalopods of South China. Palaeontologia Sinica,
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© by the Palaeontological Society of Japan 1996

1

1022

1

Family Dermochelyidae (Superfamily Chelonioidea)from the Upper Cretaceous of North JapanREN HIRAYAMA and TSUTOMU CHITOKU
Faculty of Information, Teikyo Heisei University, 2289, Uruido, Ichihara, Chiba, 290-01 Japan
Hobetsu Museum, 80-6, Hobetsu, Hobetsu-cho, Hokkaido, 054-02 Japan

Received 21 April 1996; Revised manuscript accepted 8 August 1996
Abstract. Mesodermoche/ys undulatus, a new genus and species, is described as the best Mesozoic
representative of the Dermochelyidae (order Testudines, superfamily Chelonioidea). This new taxon is
described from 15 specimens which include cranial fragments, lower jaw, axial and appendicular skeleton,
and a nearly complete shell. These specimens were collected from the Lower Maastrichtian, Upper
Cretaceous of Hobetsu-cho, Central Hokkaido, North Japan. Mesodermochelys shares with Cenozoic
dermochelyids such derived characters as the reduced processus trochlearis oticum formed by prootic,
the anterior process of surangular, and the elongate anterior projection of lateral process of humerus.
The scute sulci of Mesodermochelys are nearly absent, only remaining in the neural region of thecarapace. The bony shell of Mesodermochelys is well developed as a primitive condition. Somecharacters such as the unique configuration of the lower triturating surface with isolated simphyseal andlingual ridges, the undulating medial margin of broad peripherals, the narrow neurals, and the elongate iliac
blade with outward curvature are considered as autapomorphic conditions of Mesodermochelys. Frequent occurrence of this primitive dermochelyid in the Upper Cretaceous of Japan suggests endemic
distribution among Mesozoic chelonioid species.
Key words: Mesodermochelys, Dermochelyidae, Testudines, Reptilia, Upper Cretaceous, North Japan
Introduction
The living leatherback turtle Oermochelys coriacea is themost aberrant and unusual of turtles; the bony plates of thetypical turtle shell are nearly completely lost and jellyfish area major component of its diet (Volker, 1913; Hendrickson,1980). Because of its bizarre morphology and early ideasthat it might be the most primitive of living turtles, the living
leatherback has been the subject of more anatomical
description than any other single species of turtles (Gaffney,1984). Most of the fossil dermochelyids from the Tertiary
have been identified as extinct genera, Eosphargis or Psephohorus in most cases, but no sea turtle from the Mesozoic
has been assigned to the Oermochelyidae with certainty(Gaffney and Meylan, 1988). Allopleuron, a chelonioid fromthe Upper Cretaceous of Western Europe, is adopted as anancestral dermochelyid solely based on shell similarity(Weems, 1988); however, some Cretaceous sea turtles suchas Corsochelys (originally described as an "advanced
cheloniid" by Zangerl, 1960) and an unnamed Japanesechelonioid are currently hypothesized as being very primitive
dermochelyids (Hirayama, 1992, 1995 and in press; Hirayama
and Chitoku, 1994). A portion of this Japanese Cretaceous
sea turtle was originally identified as an indeterminablechelonioid which might be close to both dermochelyid andprotostegid in a preliminary report (Hirayama and Suzuki,1985).
Here we present a detailed description of this new taxonof sea turtle from the Upper Cretaceous of North Japan, anddiscuss its Significance to the phylogeny and evolution of the
Oermochelyidae.
Geological setting
All material described here was contained in sandy calcareous concretions which came from surface outcrop of thelower sandy siltstone formation of the Hakobuchi Group
exposed at Hobetsu-cho, Hokkaido, North Japan (Hirayama
and Suzuki, 1985; Chitoku, 1990; Hirayama and Chitoku,
1994). The Hakobuchi Group as found in Central Hokkaido
is characterized by coarse-grained sediments consisting
mainly of sandstones. It ranges from the Campanian to
Maastrichtian (Upper Cretaceous) (Matsumoto, 1942, 1943,
1954; Toshimitsu et a/., 1995). In Hobetsu-cho, the Hakobuchi Group is exposed from the Inasato district to theTomiuchi district (Figure 1 ; Tanaka, 1960; Takahashi and
Wada, 1985,1987). The middle portion of the lower sandy
siltstone formation consists of dark gray sandy siltstone,
bearing marine invertebrates such as Inoceramus (Cataceramus) shikotanensis, Pachydiscus (Neodesmoceras)

Ren Hirayama and Tsutomu Chitoku

598

142'E

QuaICI'llI l')'D
D

Allu\'ial Depos its
Tcn-acc Deposits•

Upp CI' C I'CI:u:cou.'l: YCl.O S UPl'l" Gmup
Hakobu chi CnlUp: Cu mp:luia u"'" Ma aslli cht i;lll

D

~

~

H±1
~

Upp crYc7.oGI'UUP: T u runi all - C;un plmial1

Mid dl e Yew Gmup
: Ccno m a lli :lIl~ T u ro nian

P re Upper C,,::tacC(Jus (Ace ull certain)•

Hattllonm lllii FO I'm:ttio li

Inl l1Jsi\'c Ro cks•*

Serpe ntin e
Fossilloc:llit),

o

6km

,

Figure 1. Geological map of Hobetsu-cho, showing
localities of Mesodermochelys undulatus gen. et sp. nov. (after
Takahashi and Wada, 1985). See text for the locality of eachspecimen.

japonicus and Neophylloceras hetonaiensis which indicate
an early lower Maastrichtian age. In addition to turtles, amarine reptile, Mosasaurus hobetsuensis, was reported fromthis horizon along the Pankerusanosawa River of Hobetsu(Suzuki, 1985). Obata et al. (1972) mentioned a piece ofbone presumed to be of a plesiosaur from the same locality,
which in fact belongs to an indeterminable turtle (pers.
comm. M. Manabe ; pers. obs. Hirayama).
The fossil turtle localities are scattered along the valley ofthe major rivers of Hobetsu- cho, although 8 of the 15 specimens were concentrated within the basin of the Pankerusanosawa River (Figure 1; localities 6 and 7).

Material and methods
All material described below is housed in the Hobetsu
Museum at Hobetsu- cho, Hokkaido, Japan (Institutional
Abbreviation : HMG). The specimens consist of disarticulated, scattered, uncrushed (except HMG 363 and 365,
the largest individuals) skeletons, although in many instances
some portions of the bone have been eroded (Figure 2).
Specimens were prepared by using both dilute acetic acidand mechanical methods.
Phylogenetic analysis of 135 characters for 18 eucryptodires, including 13 chelonioids, was conducted using PAUP
program (version 3.0) of Swofford (1989). A list of the 135
osteological characters utilized in the analysis, along with
their character states, is provided in Appendices I and II.
The data on chelonioids discussed below are largely basedon Hirayama (1995). In those instances where the characters are drawn from sources other than Hirayama (1995), thereference is shown in Appendix II. All characters werecoded as reversible, and multi state characters coded as
unordered to avoid a priori assumptions of transformation
vectors.
Outgroup polarity among centrocryptodires (eucryptodiresexclusive of plesiochelyids ; Gaffney and Meylan, 1988) was
determined using characters of the plesiochelyids. We
agree with the opinion that the plesiochelyids are the mostprimitive among eucryptodires (Gaffney and Meylan, 1988 ;Gaffney et al., 1991), rather than the former view that theplesiochelyids were the sister group of the remaining
chelonioids (Gaffney, 1976). Information on eucryptodiresexclusive of chelonioids is derived from the following references : Plesiochelyidae, Bram (1965), Gaffney (1975a, 1976),
Rieppel (1980) ; Meiolania , Anderson (1925), Gaffney (1983,
1985a, b, 1990); Chelydridae and Platysternon, Boulenger(1889), Gaffney (1975b, 1979) and observations on the private
collection of Hirayama ; Testudinoidea, Hirayama (1985) ;Trionychoidea, Meylan and Gaffney (1989).
We adopt the higher categorical rank among turtles asproposed by Gaffney (1984), and Gaffney and Meylan (1988).
Character state changes have been optimized on the
resulting trees using PAUP's DELTRAN option. This optimization was selected because we feel it to be slightly moreconservative in terms of assigning synapomorphies to clades
in a data matrix with a substantial amount of missing data (asadopted by Gaudin, 1995).
Systematic paleontology
Order Testudines Linnaeus, 1766
Gigaorder Casichelydia Gaffney, 1975
Megaorder Cryptodira Gray, 1825
Parvorder Eucryptodira Gaffney, 1975
Suborder Polycryptodira Gaffney, 1984
Superfamily Chelonioidea Aggasiz,1857
Family Dermochelyidae Gray, 1825
Type.- Oermochelys coriacea (Linnaeus, 1766)
Known distribution of the family.-Santonian to Maastrichtian of the northwestern Pacific Ocean, Early Campanian of
the Gulf Coast, Late Paleocene to Pliocene of the Atlantic
Sea and Pacific Ocean, and Recent of worldwide tropical tosubtropical sea.
Revised diagnosis .-Large chelonioids with shell length
exceeding 1.3 m ; processus trochlearis oticum formed byprootic reduced ; ossification of rostrum basisphenoidalemuch reduced or lost ; surangular with anterior projection
onto dentary ; lateral process of humerus antero- posteriorly
elongate, forming prominent anterior projection ; pisiform aslarge as ulnare ; fontanel between hyo- hypoplastra as large
as or larger than plastron ; scute sulci of shell rudimentaryeven in primitive forms.
Mesodermochelys gen. nov.
Type Species.-Mesodermochelys undulatus sp. nov.

1022.

Cretaceous Dermochelyidae from Japan

5

599

369

-

eel

Figure 2. Mesodermochelys undulatus gen. et sp. nov., HMG 5 (holotype) and 369. Scale=10 cm. r: right,
I: left. ca: caudal vertebra, ce1-ce8: first to eighth cervical vertebrae, co: coracoid, d1 and d2: first and second
thoracic vertebrae, den: dentary, en: entoplastron, ep: epiplastron, fe: femur, ho: hyoplastron, hp: hypoplastron, hu: humerus, i1: ilium, is: ischium, ne1 : first neural, nu: nuchal, pe1-pe11 : first to 11th peripherals, pI2-pI8:
second to eighth pleurals, pu: pubis, py: pygal, sc: scapula, so: supraoccipital, sp: suprapygal, ul: ulna, xp:
xiphiplastron.

Known distribution.-Upper Cretaceous of northwestern
Pacific Ocean.
Etymology.-Mesozoic relative of Dermochelys.
Diagnosis.-Dermochelyid with moderately developed
triturating surface; lingual ridge of dentary weak, much
lower than labial one, but reduced anteriorly; symphyseal
ridge of dentary high, exposed from lateral view above labial
ridge; lateral process of humerus with primitively more
proximal location, without distal expansion anterior to ectepicondylar foramen; iliac blade elongate, with prominent
outward curvature; shell length up to 1.5 m; shell ossification primitively well developed; scute sulci remain only on
3rd and 5th neurals and proximal portion of adjacent pleurals; neural plates narrowing posteriorly, 7th neural isreduced, 8th neural absent, 7th and 8th pleurals meet
medially; peripherals broad and robust, with undulated
medial border much expanded between costal ribs; Tshaped entoplastron with large lateral wings and rod-likeposterior projection.
Mesodermochelys undulatus sp. nov.
Figures 2-17

Holotype.-HMG 5, supraoccipital, 4th cervical to posterior
extremity of caudal vertebrae, and appendicular skeleton
including partial manus and pes, carapace lacking anterior

portion, and nearly complete plastron.
Type Locality.-Shirafunenosawa River, Inasato, Hobetsu,
Hokkaido, Japan (locality 1 in Figure 1).
Type Horizon.-Lower sandy Siltstone formation (lower
Maastrichtian), Hakobuchi Group.
Distribution.-Same as genus.
Collectors.-M.lshizaki and S. Araki, September, 1980.
Etymology.-From the Latin for the undulating medial
border of the peripheral plates.
Diagnosis.-As for genus.
Referred specimens.-HMG 6, partial carapace andappendicular skeleton. Locality-Ohmagarinosawa River,
Hiraoka, Hobetsu (locality 8 in Figure 1). Collector-M.
Susumago, May, 1977.
HMG 8-thoracic and sacral vertebrae, hind limbs, posterior
portion of shell. Locality-Pankerusanosawa River, Hiraoka,
Hobetsu (locality 7 in Figure 1). Collector-S. Suzuki, October
6, 1981.
HMG 363-prootic, caudal vertebrae, hind limbs andperipherals. Locality-Same as the above. Collector-T.
Chitoku, August 21, 1985.
HMG 365-pectoral girdles and anterior portion of plastron.
Locality-Same as the above. Collector-T. Chitoku, September 12, 1987.
HMG 369-lower jaw, hyoids, almost complete cervicals,
first thoracic and caudal vertebrae, appendicular skeletons,
anterior portion of carapace, and good plastron. Locality-Ren Hirayama and Tsutomu Chitoku

600

Same as the above. Collector-S. Toshimitsu, September 6,
1989.
HMG 9-coracoid and anterior portion of carapace.
Locality-Pankerusanosawa River, Tomiuchi, Hobetsu (locality 7 in Figure 1). Collector-S. Suzuki, October 4, 1981.
HMG 368-sixth peripheral. Locality-Same as the above.
Collectors-T. Chitoku and H. Takahashi, June 11, 1989.
HMG 1053-pelvic girdle, partial carapace. Locality-Sameas the above. Collector-T. Chitoku, September 8, 1992.
HMG 7 -sacral and caudal vertebrae, hind limbs, andposterior portion of shell. Locality-Hobetsu River, Mobetsu,
Hobetsu (locality 4 in Figure 1). Collector-S. Araki, September 4, 1980.
HMG 362-pubis and eighth pleural. Locality-Mobetsu,
Hobetsu (locality 5 of Figure 1). Collector-S. Araki (dateunknown).
HMG 342-cervical and first thoracic vertebrae, scapula,anterior portion of carapace, and partial plastron. LocalityHobetsu River, Kiusu, Hobetsu (locality 3 in Figure 1). Collector-S. Suzuki, September 16, 1984.
HMG 1062-dentary. Locality-Sososhisawa, Inasato,
Hobetsu (locality 1 in Figure 1). Collector-K. Chiyokawa,
November 2, 1994.
HMG 1063-1st and 2nd sacral and caudal vertebrae, pelvic
girdle, right femur. Locality-Same as the above. Collec-

A

tor- T. Furuno, November 2, 1994.
HMG 1064-left pubis, peripherals, and xiphiplastra.
Locality-Same as the above. Collector- T. Furuno, November 2, 1994.

Description and comparisons
Skull (Figure 3)
The only cranial elements of Mesodermochelys are two
fragments.
The posterior portion of the supraoccipital (Figures 3-A, B)
is preserved in the holotype (HMG 5). It includes a very
short crista supraoccipitalis, which projects little beyond theoverlapping area of the parietals. This condition of thecrista supraoccipitalis is quite similar to that of living Dermochelys among chelonioids (Gaffney, 1979). The verticalsheet of bone gradually narrows ventrally, without any swelling, as in Dermochelys.
The left prootic (Figures 3-C, D) was recovered with HMG
363. Its configuration suggests a very weak processus
trochlearis oticum as shown by its smooth dorsal surface
which lacks the rugosity seen in Dermochelys (Gaffney,1979). Both the canalis semicircularis horizontalis and thecanalis semicircularis anterior are confluent with the cavum
labyrinthicum as in Dermochelys (Nick, 1912).

ccanalis semicircularisanterior

BD
canalis semicircularis
horizontalis

Fi~u.re 3. Cranial elements of Mesodermochelys undulatus gen. et sp. nov. A, B. Posterior portion of supra?CClpltal, HMG 5 (holo~pe), A: dorsal view, B: left lateral view. Arrow showing posterior limit of suture withparietal. C, D. Left prootiC, HMG 363, C: anterodorsal view, 0: ventral view. Scale=5 cm.

1022.

Cretaceous Dermochelyidae from Japan

60 1

o

E

H
369.

Figure 4. Stereophotographs of left half of lower jaw of Mesodermochelys undulatus gen. et sp. nov., HMG
A, B. Dorsal view. C, D. Ventral view. E, F. Lateral view. G, H. Medial view.

Ren Hirayama and Tsutomu Chitoku

602
Lower jaw (Figures 4, 5)

The lower jaw is preserved in HMG 369 and 1062. HMG
1062 consists of an isolated dentary, lacking its posterior
portion. HMG 369 (Figure 4) is a nearly complete left half ofthe lower jaw, lacking only a small symphyseal portion of thedentary. HMG 1062 shows a unique configuration of thelower triturating surface among turtles; the symphyseal
ridge is very high. It is visible in lateral view above the labial
ridges (Figure 5- 8). The lingual ridges are very low, becoming reduced anteriorly such that there is no connection
between symphyseal and lingual ridges (Figure 5- A). The
lingual ridge of HMG 369 is identical to that of HMG 1062.
The lateral surface of the dentary is ornamented with numerous longitudinal ridges, suggesting the presence of relatively
thick rhamphotheca. The triturating surface of the dentary
is moderately developed in Mesodermochelys just like living
Eretmochelys (Cheloniidae) which has a secondary palate
and Protostega (Protostegidae) which has only a primarypalate. It is much broader than the dentary in Toxochelys(primitive Cheloniidae) or Oermochelys, both of which have

5CM

B

only a primary palate. Thus, it is not possible to determine
if Mesodermochelys had a secondary palate based solely on
the development of the lower jaw.
The angular is a long and slender sheet of bone as in otherturtles. The surangular has a short anterior process onto
the dentary, which is not seen in any other Mesozoic turtles.
This surangular process of Mesodermochelys, however, ismuch smaller than in living cheloniids (exclusive of Natator
depressus, which lacks the surangular process ; Hirayama,
1995) and Oermochelys (Gaffney, 1979). The coronoid has
the processus coronoideus narrowly exposed from lateralview. It is barely higher than the dentary. The articular has
a definite retroarticular process, on which the foramen
chorda tympani is visible at the base. The prearticular is alarge sheet, covering the posterior half of the fossa meckelii.
Hyoid : In HMG 369 both right and left cornu branchiale arepreserved. Each is 95 mm in length. Their configurationand size relative to the lower jaw are identical to living
chelonioids.

Cervical vertebrae (Figures 6, 7)
The cervical vertebrae are preserved in the holotype (HMG
5), HMG 342 and 369, being the most complete in HMG 369.
Only the atlas intercentrum and the third vertebra are notpreserved in any other specimens. In all of the cervicals thetransverse process is single, and there is no trace of any
cervical rib as is the case in other polycryptodires (Gaffney
and Meylan, 1988). The cervical vertebrae are proportionallyhigh and short as in Oermochelys, Corsochelys and protostegids, and the central articular surface is as high as wide or
higher than wide even in the 5th to 8th cervicals unlike
cheloniids (Zangerl, 1960 ; Zangerl and Sloan, 1960 ; Hirayama, 1995). The neural spine of the 8th cervical is particularly high, with a prominent dorsal articular facet that articulates with the ventral knob of the nuchal plate as in otherchelonioids. The 8th cervical centrum is longer than the 7th
as in protostegids and Oermochelys, not shorter than the 7th
as in the other polycryptodires including cheloniids (Volker,
1913 ; Williams, 1950 ; Zangerl, 1953; Zangerl and Sloan,
1960).
The centra of the cervicals of Mesodermochelys all havefully formed articulations, with cervicals two (and three)opisthocoelous, cervical four biconvex, and cervicals five toeight procoelous. Therefore, the formula of the cervicalcentral articulation is shown as (2 (3 (4) 5) 6) 7) 8). Thispattern is common in the centrocryptodires including
Meiolania (Gaffney, 1985 ; Gaffney and Meylan, 1988).
There is no indication of double or platycoelous articulations
between centra as seen in most living chelonioids (Williams,
1950 ; Hirayama, 1995)

Thoracic vertebrae and ribs

Figure 5.

Composite reconstruction of lower jaw of

Mesodermochelys undulatus gen. et sp. nov., based on HMG

369 and 1062. A. Dorsal view. B. Left lateral view.

The thoracic vertebrae, exclusive of the first one, arepreserved in the holotype. The thoracic centra are rather
slender in both lateral and ventral aspects. The openings
for the spinal nerves are long semilunar embayments extending about 60% of the centrum length as in cheloniids and1022. Cretaceous Dermochelyidae from Japan

603

A

c

B

o

2cm

D

G

H
Figure 6. CeNical (A to G) and first thoracic (H) vertebrae of Mesodermochelys undulatus gen. et sp. nov.,
HMG 369, left lateral view. Third ceNical is missing.

They are not as extensive as in advanced
protostegids such as Archelon (Baird, 1978). The first thoracic is preseNed only in HMG 342 and 369 (Figure 8- H).
The first thoracic vertebra has its anterior articulation with a
Oermochelys.

nearly vertical plane to the axis of centrum, facing anteriorly
as in Oermochelys and protostegids (including Oesmatochelys), rather than ventrally or anteroventrally as incheloniids, including Toxochelys and Allopleuron (Hirayama,

Ren Hirayama and Tsutomu Chitoku

604

2

1

6

5

4

7

8

.

~
N..

"':"

.,'

:~.

10 em
Figure 7. Cervical vertebrae of Mesodermochelys undulatus gen. et sp. nov. First, second and sixth cervicals, HMG 369. Fourth, fifth, seventh and eighth cervicals, HMG 5 (holotype). Third cevical is missing. Upperrow, anterior view. Middle row, posterior view. Lower row, left lateral view.
1995), and most other polycryptodires (Gaffney et aI., 1991).
The isolated first thoracic ribs were recovered from HMG
342 and 369 (65 mm in length). These are relatively shortbones as in cheloniids and Dermochelys (Zangerl, 1953).
Their distal end does not extend beyond the nuchal width
unlike plesiochelyids in which it extends beyond the nuchal(Rieppel, 1980).
Both the ninth and tenth thoracic ribs are well preserved in
the holotype. They both show fusion to the overlying
pleurals as in most turtles.

Sacral vertebrae and ribs (Figures 8-A, 0, E)
Both first and second sacral vertebrae and their attachedribs are preserved in the holotype. The second sacral andright rib are preserved in HMG 7. The first and second
sacral vertebrae are also known from HMG 1063. The
sacral vertebrae are roughly platycoelous, rather thanprocoelous. These vertebrae and ribs are relatively stoutbones in comparison with those of living chelonioids.

Caudal vertebrae and ribs (Figures 8-B, C, F-P, 9)
The caudal vertebrae are preserved in the holotype, HMG

7,363,369 and 1063. All caudals are procoelous, showing
no trace of chevron bones ventrally. In the holotype, ten
disarticulated caudals, which appear to represent the first to
seventh and the last three, are present. The first caudal of
the holotype has small ribs. By comparison with this series
for the holotype, the position of the caudals in the otherspcimens could be determined with some certainty. Threecaudals of HMG 7 are presumed to be the second, third andsixth, the two from HMG 363 are likely the fourth and? tenth,
the one from HMG 369 is the fourth, and the two of HMG(94-206-B) are the first and second. The tail of the male ismuch larger and longer, often twice or more than the female
in most living turtles including chelonioids (Ernst and Barbour,
1989; Hirayama, pers. obs.). Thus, a careful comparison of
the relative size of these caudals with those of living forms
might reveal sexual dimorphism in Mesodermochelys.

Pectoral girdle (Figure 10)
The scapulae have been recovered from five specimens,
the holotype, HMG 6, 342, 365 and 369. The pectoral girdle
shows the characteristic triradiate structure of turtles. The
scapular angle formed by the scapular prong and theacromion is wide, reaching up to 110 degrees. The cor-

1022.

Cretaceous Dermochelyidae from Japan

605

o
B
F

J

c
o

K

L

p

5 em

J

M

N

Figure 8. Sacral and caudal vertebrae of Mesodermochelys undulatus gen. et sp. nov., HMG 5 (holotype). A,
D, E. First and second sacrals with ribs, A: dorsal view, 0 (first), E (second) : anterior view. B, O. First to seventhcaudals in articulation, dorsal and lefi. lateral views. C, P. Last three caudals in articulation, dorsal and left lateralviews. F to L : Anterior views of first to seventh caudals. M, N : Anterior views of two of the last three caudals(last two caudals fused together).

Ren Hirayama and Tsutomu Chitoku

606

A
o

5 em

B

c
D

D

Figure g. Caudal vertebrae of Mesodermochelys undulatus gen. et sp. nov., in left lateral view. A. (HMG 5 :holotype), first to seventh, and last three caudals. B. (HMG 7), second, third and sixth caudals. C. (HMG 369),
fourth? caudal. D. (HMG 363), fourth and tenth? caudals.

acoids are preserved in four specimens, including theholotype, HMG 9, 365 and 369. The coracoid is proportionally narrow and elongate, 248 mm long (right side) in theholotype and 228 mm long (left side) in HMG369, much
longer than the humerus. The coracoid of HMG 365 aspreserved is 380 mm long (left side), and was probably twice
the size of the holotype coracoid. A median keel for the M.
supracoideus is distinct in the holotype, whereas this keel is
not so prominent in the other specimens. The generalmorphology of the pectoral girdle in Mesodermochelys is
identical with those of the pelagic chelonioids such as thedermochelyids including Corsochelys, the protostegids, andsome cheloniids such as Allopleuron.

iliac blade is quite elongate, somewhat like in the trionychids.
It is also laterally curved as in the testudinoids, particularly
the living batagurids (Hirayama, pers. obs.). The pubis of
Mesodermochelys is identical to those of dermochelyids andprotostegids in the possession of the huge lateral process(= pectineal process), which is much extended anteriorly
beyond the medial border of this bone. This is not the casein cheloniids which have a much smaller lateral process(Hirayama, 1995). The ischium has a rudimentary lateralprocess (=metischial process) as in dermochelyids,
advanced protostegids (e.g., Protostega) and living
chelonioids exclusive of Natator (Zangerl et al., 1988)
Forelimb (Figures 12, 13)

Pelvic girdle (Figure 11)
The elements of the pelvic girdle is preserved in nine
individuals, the holotype, HMG 6,7,8, 362, 363, 369, 1063,
and 1064. Of them the pelvic girdle is completely preserved
only in the holotype. The thyroid fenestra, or the obturator
foramen, is small and completely subdivided by the medialpubo-ischium contact as in protostegids and dermochelyids,
and unlike those of cheloniids which have a large, confluent
fenestra (Hirayama, 1995).
The ilia are preserved in HMG 7, 8 and 1053. The ilium of
Mesodermochelys is quite peculiar among chelonioids. The

Only in the holotype and HMG 369 is a part of the forelimbpreserved. The humerus (left) is 195 mm long in theholotype and 172 mm long in HMG 369 (right). It is much
longer than the femur as described below. In HMG 369, theossification of the distal end is incomplete, indicating thepossible immaturity of this specimen. Hirayama (1992) fully
discussed the humeral morphology of sea turtles including
the holotype of Mesodermochelys, and recognized "toxochelyid", "cheloniid", "protostegid" and "dermochelyid"
grades among chelonioids. Thus, we follow this "gradesystem" here.

1022. Cretaceous Dermochelyidae from Japan

B

C

F

E

607

G

H
Figure 10. Pectoral girdle of Mesodermoche/ys undulatus gen. et sp. nov. A. Right scapula, HMG 369,
anterior view. B. Left scapula, HMG 369, posterior view. C. Right scapula of HMG 5 (holotype), posterior view.
D. Right scapula, HMG 342, posterior view. E- G : right coracoid, holotype, dorsal (stereopair) and ventral views.
H, I. Right coracoid, HMG 369, dorsal view (stereopair).

Ren Hirayama and Tsutomu Chitoku

608

5cm

o

5 em

L

o

5cm

o
o

5cm

J
Figure 11. Pelvic girdle of Mesodermochelys undulatus gen. et sp. nov. A- C. Pelvic girdle in articulation(cast), HMG 5 (holotype), dorsal, ventral and right lateral views. D, E. Right ilium, HMG 7, lateral and dorsal view.
F. Pubes, HMG 369, dorsal view. G. Right pubis, HMG 363, dorsal view. H, I. Left ilium, HMG 8, lateral anddorsal views.

1022.

Cretaceous Dermochelyidae from Japan

B

609

c

10cm
Figure 12. Left humerus of Mesodermochelys undulatus gen. et sp. nov., HMG 5 (holotype).
ventral and anterior views (Slightly modified from Hirayama, 1992).

The humerus has a nearly straight shaft extending toward
large caput humeri and high medial process. The greatly
expanded muscle scars for M. coracobrachialis brevis, latissimus dorsi and teres major, and the lateral process arelocated more distally and ventrally than in primitive cheloniidsof the "toxochelyid grade" such as loxochelys or Osteopygis.
The lateral process of the humerus of Mesodermochelys ischaracterized by its long anterior projection where the M.
deltoideus inserts as in dermochelyids or Syl/omus, a
Neogene cheloniid. The lateral process of Mesodermochelys, however, is elongate and nearly straight relative tothe shaft as in other dermochelyids. It lacks the V-shaped
structure of advanced cheloniids such as Syl/omus or the
medial concavity of protostegids. The lateral process in
Mesodermochelys is located at a more proximal position thanin Cenozoic dermochelyids, and its shaft is greatly constricted as in Toxochelys, primitive protostegids (e.g., Desmatochelys) and Corsochelys. It is neither as wide nor as flat asin Cenozoic dermochelyids or advanced protostegids (e.g.,Archelon). In summary, the humeral morphology ofMesodermochelys could be interpreted as a primitive "dermochelyid grade", intermediate between the Cenozoic dermochelyids with an advanced "dermochelyid grade" and theprimitive chelonioids such as Toxochelys (Hirayama, 1992).
The left ulna is present in the holotype. It does not havea distal area with distinct rugosities for articulation with theradius as in all of the cheloniids (Hirayama, 1995; e.g.,
Toxochelys and AI/opleuron).
The manus is preserved in the holotype and HMG 369.
Two carpal elements, the ulnare and pisiform, two metacarpals of the second and fifth digits, and the first distal phalanx,

A-C. Dorsal,

all of the left side, were recovered with the holotype. The
ulnare is roughly rectangular, a rather thick and massive
bone, with a clear articular facet along each border, like inmodern cheloniids rather than Dermochelys. The pisiform issimilar to the ulnare, just slightly smaller than the latter,though it is relatively much larger than in any cheloniid.
One metacarpal, presumed as the third, is preserved in HMG
369. The articular surface of this element is flattened, andhence inmovable as in most chelonioids. Amongchelonioids only some primitive cheloniids such as Toxochelys and one undescribed protostegid primitively retain the
movable artiCUlation at the first and second digits (Hirayama,
1995, in press).

Hind limb (Figure 14)
Parts of the hind limb, including the femur, were recoveredfrom the holotype, HMG 6, 7, 8, 363, 369 and 1063. The
right femur is 145 mm long in the holotype and 136 mm longin HMG 369, much shorter than the humerus as in mostchelonioids. The major and minor trochanters of the femur
are connected by a bony ridge with a straight proximal
border as in Dermochelys, protostegids including Desmatochelys, and living cheloniids (exclusive of Natator; Zangerl etal., 1988). The major trochanter of the femur is not higher
than the minor one as seen in advanced cheloniids including
AI/opleuron and most Cenozoic forms. The tibia is preserved in HMG 7,363, and 369, and the fibula is preserved only
in HMG 363. These are rather stout bones as in cheloniids.
There is no trace of tendon pit on tibia.
Elements of the right pes preserved with the holotype

Ren Hirayama and Tsutomu Chitoku

6 10

A

o

5cm

B

o

5cm

Figure 13. Left ulna and manus of Mesodermochelys
undulatus gen. et sp. nov., HMG 5 (holotype). A. Ulna, dorsal

and ventral views. B. Manus, with ulnare, piciform, ungual I,
metacarpals II and V preserved.

include the astragalus, calcaneum, third metatarsal, and two
phalanges of the first digit. Movable articulations areabsent in these bonse (as in the menues). The pes ispresumed to be about 60% of the legth of the menues, as inliving cheloniids.
Shell (Figures 15, 16, 17)

The shell is partially preserved in all speCimens exclusiveof HMG 1062. The best shell is that of the holotype,although its anterior portion of the carapace is missing. The

anterior portion of the carapace including the nuchal bone is
known from HMG 342 and 369. The plastron of HMG 369
is also essentially complete. Thus, a composite reconstruction of the entire shell is made with certainty.
The shell scute sulci are very poorly developed in
Mesodermochelys. Prominent scute sulci are observed only
on the third and fifth neural plates, then become reduced
and disappear at the proximal portion of the third and fifth
pleurals. Rudimentary furrows, possibly scute sulci, arepreserved at the lateral edge of the sixth to ninth peripherals.
Thus, it is highly probable that Mesodermochelys had shellscutes covering only the neural (and possibly marginal)
reg ion of the carapace in life. This condition is reminiscentof the juvenile of living Carettochelys insculupta (Carettochelyidae; Trionychoidea ; Zanger!, 1959).
The reconstructed carapace of Mesodermochelys is rather
more circular than oval ; its width is only 10% smaller than its
median length. The nuchal plate has a prominent ventral
knob to which the neural spine of the eighth cervical
articulates as in many chelonioids including Oermochelys.
The neural plates are rather rectangular as in the protostegids, Corsochelys and Eosphargis, and unlike thecheloniids which have hexagonal neurals (Hirayama, inpress). The neurals of Mesodermochelys, however, areextremely narrow and reduced particularly in the posterior
part. The pentagonal seventh neural is reduced posteriorly,
and the eighth is absent resulting in the seventh and eighth
pleurals meeting medially as in Cuora (8ataguridae) and thepelomedusids (Pleurodira). The suprapygal is single as inprotostegids, and not subdivided as in most cryptodires
including Corsochelys, Eosphargis, and cheloniids. Contrary
to the reduced neural area, the peripheral plates are robustand broad in Mesodermochelys. The medial portion of theperipherals extends medially between the costal rib ends sothat the medial border of the peripheral series is strongly
undulating and results in a restriction of the pleuro- peripheral
fontanelles. These fontanelles are more restricted than in
Corsochelys in which ossification of the pleurals is comparable to that in Mesodermochelys.
The plastron is best preserved in the holotype and HMG
369. The plastron shows no trace of scute sulci, as is thecondition found in protostegids, Corsochelys and Oermochelys. The maximum thickness of the hyo- hypoplastra isabout 10 mm in the holotype and HMG 369. It is characterized by a high plastral index (the ratio of the minimum lengthof the hyo- hypoplastral bridge to half the width of theplastron), about 110 in the holotype and 120 in HMG 369, andshows a large fontanelle between the hyo- hypoplastra.
Thus, its general appearance is most similar to that ofCorsochelys (ZangElfI, 1960). The epiplastra are preserved in
the holotype, HMG 363 and 369, showing a notable difference among Mesodermochelys. The epiplastron is rather
narrow and slender in the holotype and HMG 369 as inToxochelys, whereas this is a much wider element in HMG
363, the largest specimen. The entoplastron is preserved in
the holotype, HMG 363 and 369. The bone is T - shaped,with large lateral wings and a long posterior process, which
is rod - like without a horizontal sheet. This is unlike otherchelonioids but more similar to Chelydra and Macroclemys

1022.

Cretaceous Dermochelyidae from Japan

6 11

o
"0

5cm

E

o

o

Scm

Scm

H

K

J

p

o

L
5cm

L_ J

o

M
Scm

N

o

Q

o

Scm

R

L==~J

Figure 14. Hind limb of Mesodermochelys undulatus gen. et sp. nov. A- G. Femora of HMG 5 (holotype), A.
C: left side, ventral view (stereopair), B, D : right side, ventral view (stereopair), E : right side, anterior view, F, G :right side, proximal view. H, J. Left femur, HMG 8, ventral view (stereopair). I, K. Right femur, HMG 369, ventral
view (stereopair). L, N. Left tibia, HMG 7, anterior view (stereopair). M, O. Right fibula, HMG 363, anterior view(stereopair). P. Right astragalus and calcaneum, holotype, dorsal view. Q. Right metatarsal II, holotype, dorsal
view. R. Right phalanx and ungual of phalanges I, holotype, dorsal view.

Ren Hirayama and Tsutomu Chitoku

612

A

B

c
o

10 em

D

Figure 15. Carapace of Mesodermochelys undulatus gen. et sp. nov. A. Nuchal, HMG 342, ventral view. B.
Nuchal with left first peripheral attached, HMG 369, ventral view. C. Nuchal, right first and second peripherals, firstpleurals, HMG 342, dorsal view. D. Carapace, HMG 5 (Holotype), dorsal view.

(Chelydridae). Each lateral wing of the entoplastron has afacet for the epiplastron, whereas there is no facet for the
hyoplastron. The medial processes of the hyo- hypoplastra
are not as long as in Corsochelys, possibly not interdigitatingwith corresponding processes across the midline. The lateral processes of the hyo- hypoplastra are also rather short,
not showing any contact with the peripherals as is the casein most chelonioids. The xiphiplastra are nearly straight,

1022.

Cretaceous Dermochelyidae from Japan

6 13

A
o

I De mB

cD

Figure 16. Plastron of Mesodermochelys undulatus gen. et sp. nov. A. HMG 5 (holotype), ventral view. B.
HMG 369, ventral view. C, D. Left epiplastron, HMG 365, dorsal and ventral views.

blade- shaped bones, rather tightly sutured to the hypoplastrain the holotype and HMG 369. There is no trace of a medial
meeting between the xiphiplastra.

Phylogenetc analysis
Results.- A single tree of 252 steps (Figure 18) and aconsistency index of 0.587 was obtained by branch- and -bound search options of PAUP (version 3.0 ; Swofford,1989).
This result supports the monophyly of most higher categorical ranks among eucryptodires, proposed by Gaffney andMeylan (1988), but not the procoelocryptodires (chelonioids,
trionychoids and testudinoids). The inclusion of additional
fossil taxa such as Ordosemys, Sinemys, and Macrobaena
from Asia into the data matrix may be ready to alter the
relationships and character transformations among eu-

614

Ren Hirayama and Tsutomu Chitoku

Ilocm

Figure 17. Composite reconstruction of Mesodermochelys undulatus gen. et sp. nov. largely based on HMG
5 (holotype), 342 and 369. Skeleton in ventral view, carapace in dorsal view.

cryptodires (Brinkman and Gaffney, pers. commun.; Hirayama, in preparation).
The monophyly of chelonioids is supported by 16 characters, of which 14 are unequivocal. Four are related to theformation of paddles, seven are related to reduced shellossification. The monophyly of each family among
chelonioids of Hirayama (1995) is supported; Cheloniidae(including Toxochelys and Ctenochelys) was united by 14 (8unequivocal) characters, Protostegidae by 13 (11 unequivocal)
characters, and Dermochelyidae by 5 (4 unequivocal) characters. Both Mesodermochelys and Corsochelys were placed
as very primitive members of the family Dermochelyidae, as
hypothesized by previous works of Hirayama (1992, 1995, inpress).
Discussion.- The goal of this analysis is to assess thephylogenetic position of Mesodermochelys among
chelonioids. Due largely to missing data and differences ofpreseNation (Appendix I), the phylogeny of the family Dermochelyidae is rather poorly resolved in comparison to those of
the protostegids and cheloniids. In the case of Mesozoic
dermochelyids, there is a significant gap in the data set
because of the poor preseNation of cranial materials. Mostof the skull of Mesodermochelys is unknown, whereas theanterior portion of the skUll and the lower jaw is missing in
Corsochelys. Thus, the large foramen caroticum laterale andthe short subdivided rostrum basisphenoidale are equivocal
in Mesodermochelys, whereas the anterior process of thesurangular is equivocal in Corsochelys. Nonetheless, twocharacters, the poor development of the processus trochlearis oticum formed by prootic and the rudimentary scute sulcion the carapace, could be explained as an unequivocal
synapomorphy uniting them with Cenozoic dermochelyids.
Large size with maximum shell length exceeding over 1.3 m
seems also to be shared by all known dermochelyids (Hi-

Figure 18. Hypothesized phylogeny of the Chelonioidea based on PAUP analysis of 13 extinct and extant taxa and their
5 outgroups in the Eucryptodira. Distribution of apomorphies at each Node is shown by numbers which correspond tocharacters listed in Appendix II and scored in Appendix I as follows. Parallelisms are marked with n, and character reversals
are marked with (*). Node A1: Centrocryptodira (61,64,65-1,81*,120,121*), A2: Polycryptodira (3,4,54,58,59,60,72,762), 81: (7",8-2",34",43-0*,62",63",66",67",68,71"), 82: Chelomacryptodira (32",74",125*), C1: Chelonioidea (14,37,
50",74",85,86,95,96,112,119,122,127,128,129,132), C2: (11",12",16",24",77-2",79",80,87",97",109-2,133"), C3:
Cheloniidae (6,15,19,38",40,62",63",71",77-1,81",93,103",108, 123-2), C4: (8*,16",27,41,44"), C5: (2",18*,24",46",
67"), C6: (6*,66",70",77-2",87",88,97",99",123-1*), C7: (17",54*,79",133"), C8: Protostegidae (22,23,28,29,30,36",
39*,44",72*,89,90,94,112-2"), C9: Dermochelyidae (31, 45, 46", 10H", 125-2"), C10: (35,56",82",91,98",103"), C11 : (5",8*,10",20,26",42,49",92,101-2",112-3,125-3), C12: (50*,86-2,113).

1022.

Cretaceous Dermochelyidae from Japan

615

Piesiochelyidae
Heiolania

Chelydroidea

BI
r----

Testudinoidea---

Trionychoidea

"--

B2

"--

Al

Toxochelys

C3.....-"--

Ctenochelys

A2
Osteopygis'--

FamilyC4"--C5..--"--r--

Allopleuron---

Chelonia IlIYdas

Cheioniidae

C7
'--

C6

CI

Puppigerus

Superfam i Iy-,

Oesll8.tochelys

I
I

C8

Family: Protostegidae_2

Protostega gigas"--

Corsochelys

C2

-i

I

"--

Hesodel'llochelys

C9-Eosphargis

CIO

Del'llochelyidae
"--

CII

-

Psephophorus'---

DeI'llochelys"""""-

Cl2

Cheionioidea

Ren Hirayama and Tsutomu Chitoku

616

rayama, 1995) Mesodermochelys appears. to be ~oreadvanced than Corsochelys in the possession of relatively
large paddles involving a humerus with an elongate ~r~je?­tion of the lateral process and the short crista supraocclpltalis.
The pronounced reduction of shell ossificatio~. and thescute loss are also seen in Allopleuron (Chelonlldae) andadvanced protostegids (Protostega and Archelon). Theseare explained as a result of extreme convergence among
chelonioids as proposed by Hirayama (1995, in press). With
Allopleuron forced into the basal position for both protostegids and dermochelyids as hypothesized by Hirayama(1992), we require 4 more derived characters share~ by thosechelonioids than by others and the tree length Increases
from 252 steps to 259 steps, the remaining cheloniids thencollapse into a paraphyletic group constituting primitive
grades among chelonioids as shown by Gaffney and Meylan(1988) or Hirayama (1992). Furthermore, to force Allopleuron
into the basal position for Cenozoic dermochelyids as
hypothesized by Weems (1988) we require 21 more derived
characters shared by them of which at least 8 are not sharedby Corsochelys and Mesodermochelys and the tree length
increases to 293 steps. The cheloniid affinity of Allopleuron, however, is supported by overwhelming characters,
although Mesodermochelys remains within the dermochelyids as a monophyletic group, in every case.
Conclusions
The phylogeny of the family Dermochelyidae is rather
poorly supported due to missing data and variation in preservation. However, two Mesozoic dermochelyids, Mesodermochelys and Corsochelys, demonstrate that the significantly
reduced shell ossification seen in Cenozoic dermochelyids
including Eosphargis was preceded by shell scute loss andthe development of the characteristic dermochelyid humerus. Mesodermochelys is considered to be more closely
related to Cenozoic dermochelyids than Corsochelys in its
possession of relatively large paddles and a humerus with anelongate projection of the lateral process and a short crista
supraoccipitalis.

The pronounced reduction of shell ossification and thescute loss is also seen in Allopleuron (Cheloniidae) andadvanced protostegids (Protostega and Archelon). Theseare explained as a result of extreme convergence among
chelonioids.
Hirayama (1994, in press), and Hirayama and Chitoku (1994)
demonstrate the peculiarity of the Japanese chelonioid
fauna during the Upper Cretaceous. Primitive dermochelyids such as Mesodermochelys are the most abundantchelonioids from the Santonian to Maastrichtian of Japan,
whereas the cheloniids, which dominate the Cretaceous ofNorth America and Western Europe, are virtually unknown
from Japan. Upper Cretaceous dermochelyids are extremely rare outside Japan. Corsochelys haliniches, known only
from the holotype, is the only Mesozoic dermochelyid knownwith certainty from outside Japan. Thus, Hirayama (in press)
suggests that endemic speciation may have occurred among
the chelonioids during the Cretaceous, resolving into apattern of diversity unlike that for living sea turtles, of whichmost genera including Dermochelys, are cosmopolitan. We
suppose the western part of the Pacific Ocean around Japanmight be the center of diversification of Cretaceous dermochelyids at least since the Santonian. This might be ~hereason why Mesozoic dermochelyids are so rare outSide
Japan. Nonetheless, Mesodermochelys should be a side
branch of dermochelyid phylogeny, not directly ancestral to
any later taxon. It possesses peculiar autapomorphic .characters such as the unique configuration of the lower tnturating surface, unique iliac blade, and peripherals with an
undulating medial border. This is also the case in Corsochelys which has several autapomorphic characters such as
the amphicoelous cervicals (explained as character reversal).
Thus, it seems highly probable that we will discover anotherdermochelyid of Cretaceous age that is more closely related
to the Cenozoic forms.
Acknowledgments
We thank Drs. E.S. Gaffney (American Museum of Natural
History, New York) and PA Meylan (Eckerd College, St.
Petersburg) for encouraging this study and thoughtprovoking
comments on turtle phylogeny. For access to comparative
material we thank Drs. D. Berman (Carnegie Museum, Pittsburgh), J. Bolt (Field Museum of Natural History, Chicago),
D.B. Brinkman (Royal Tyrell Museum of Palaeontology,
Drumheller), M. Darling (Sedgwick Museum, Cambridge), N.
Hotton III (USNM, Smithsonian Institution, Washington D.C.),
E.S. Gaffney (AMNH), W.F. Meijer (Natuurhistorisch Museum,
Maastricht), J.H. Ostrom (Yale Peabody Museum, New
Haven), D.C. Parris (New Jersey State Museum, Trenton), H.
Saegusa (Museum of Human Activity and Nature, Sanda,
Japan), P. Sartner (Institut Royal des Sciences Naturelles
Belgique, Brussels), H.-P. Schultze (University of Kansas,
Natural History Museum, Lawrence), T. Uyeno, Y. Tomida
and M. Manabe (National Science Museum, Tokyo), C.A.
Walker (Natural History Museum, London) and Mrs. K. Kishinami (Bishop Museum, Honolulu). Drs. D.B. Brinkman
and E.L. Nicholls (Royal Tyrrell Museum of Palaeontology)
gave useful comments on the phylogeny among eucryptodires. Dr. N. Kohno (National Science Museum) and Mr. Y.
Yasukawa (Kyoto Univ., Kyoto) gave much useful commentson the methodology of phylogenetic analysis. Messrs. J.
Munemasa and K. Sawaki (former undergraduates of Teikyo
Heisei Univ.) helped draw Figure 7. Mr. G. Kawakami(Hobetsu Museum, Hobetsu) helped draw Figure 1. This
manuscript was substantially improved by Dr. PA Meylan.
This work was partially supported by a Grant-in-Aid from
Teikyo Heisei Univ. (formerly Teikyo Univ. of Technology).
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Williams, ES., 1950: Variation and selection in the cervicalcentral articulations of living turtles. Bulletin of theAmerican Museum of Natural History, vol. 94, p. 505562.
Zangerl, R, 1953: The vertebrate fauna of the Selma Formation of Alabama. Part 3. The turtles of the family
Protostegidae. Part 4. The turtles of the family Toxachelyidae. Fieldiana, Geology, Memoirs, vol. 3, p.61277.
Zangerl, R, 1959: Rudimentare Carapaxbeschuppung bei
jungen Exemplaren von Carettochelys und ihre morphogenetische Bedeutung. Naturforschenden Gesellschaft in Zurich, vol. 104, p. 138-147.
Zangerl, R, 1960: The vertebrate fauna of the Selma Formation of Alabama. Part 5. An advanced cheloniid sea
turtle. Fieldiana, Geology, Memoirs, vol. 3, p.281-312.
Zangerl, R, 1971: Two toxochelyid sea turtles from theLandenian Sands of Erquelinnes (Hainaut) of Belgium.
Memoires de /'Institut Royal des Sciences Naturelles deBelgigue, vol. 169, p.1-32.
Zangerl, R, 1980: Patterns of phylogenetic differentiation in
the toxochelyid and cheloniid sea turtles. American
Zoologist, vol. 20, p.585-596.
Zangerl, R, Hendrickson, L.P. and Hendrickson, J.R., 1988 :A redescription of the Australian flat back sea turtle,
Natator depressus. Bishop Museum Bulletin of Zoology, vol. 1, p.1-69.
Zangerl, R and Sloan, R E, 1960: A new specimen of
Desmatochelys lowii Williston; a primitive sea turtle
from the Cretaceous of South Dakota. Fieldiana, Geology, vol. 14, p. 7-40.

1022.

Cretaceous Dermochelyidae from Japan

APPENDIX I.

619

Data Matrix

The symbol "?" represents missing or equivocal data. The following symbols are used to represent character statesin polymorphic taxa: a=(0,1), b=(1, 2), c=(02), d=(012), e=(1, 3).

Plesiocelyidae

Meiolania
Chelydroidea
Testudinoidea
T rionychoidea

Toxochelys
Ctenochelys
Osteopygis
Allopleuron
Puppigerus
Chelonia mydas
Desmatochelys
Protostega gigas
Corsochelys
Mesodermochelys
Eosphargis
Psephophorus
Dermochelys
Plesiochelyidae

Meiolania
Chelydroidea
Testudinoidea
Trionychoidea

Toxochelys
Ctenochelys
Osteopygis
Allopleuron
Puppigerus
Chelonia mydas
Desmatochelys
Protostega gigas
Corsochelys
Mesodermochelys
Eosphargis
Psephophorus
Dermochelys

5
10
15
20
25
30
35
40
45
50
55
60
65
70
010aO Oa100 aaOOO 00000 OaOOa 00000 00000 1aaaO OOaOO OOOOa 00000 OOO?O 00000 00000
01001 00300 00000 00000 00000 00000 02110 1001 0 001 00 00000 00100 00000 10011 00000
1a111 a1 cOO OOaoa 00000 00000 00000 02010 10110 00000 00010 00010 00111 1111 b 111 aO
0011 a a 1200 aaOOa aaaOO aOOOa OOaOO Ob01 0 1Oa 10 00000 OOaOa aaaaO a0111 111 b2 111 aO
aa11 a 01200 OOaOa OOOOa 00000 00000 01010 1Oa1 0 00000 aOaaa OaOaO 00111 111 de 11100
00010 10100 00011 00010 00000 00000 00000 11111 00100 OOOOa 00010 00111 11111 10000
00110 10000 00011 11 01 0 00000 01000 00000 11111 10110 00001 00010 00111 11111 00000
01110 10000 00011 10110 10010 01000 00000 11111 10110 10100 00011 00111 111?1 ?10?0
00110 01201 11 011 11110 00000 01 000 00000 11111 1011 0 00001 00000 00111 11111 11001
0111 0 00000 00011 1011 0 1001 0 01 000 00000 11111 1011 0 101 01 00010 00111 11111 11 001
0111 0 00000 00011 1111 0 0001 0 01 000 00000 11111 10110 11 001 111 00 10111 11111 11 001
01000 011?0 11010 10000 01110 00111 00000 01000 00110 00001 00010 00111 10021 00000
10111 00010 11110 1??00 01110 10111 00000 01000 00110 00000 00010 00111 100?1 00000
OO??O ?1200 11 ??? ????? ???10 00000 10000 11110 00101 10??? ????? ???11 OOOO? ?OOOO
O???? ????? ????? ????? ????? ????O 1??? 1 ????? ????? ??001 10000 10111 10011 00000
00111 00001 11010 10001 00010 10000 10001 ????O 01 ??? ??011 10000 ???11 ??O?? ?O??O
00111 00001 ??01O 10001 0001 0 1000? ?0001 ????O 01 ??? ??01 0 0001 0 ????? ????? ?????
00111 00001 1101 0 10001 0001 0 10000 10001 01 000 01101 11 01 0 0001 0 11111 10011 00001
75

80

85

90

95

100

105

110

115

120

125

130

135

000?0
00?01
1aa01
11 01 011110
1?? 10
11010
?????
11010
11010
11 01 0
OO???
00010
?????
01010
?????
?????
01 01 0

00000
00000
10000
20000
20000
21 000
21000
??OOO
22010
22000
2201 0
?2011
22111
?201?
22011
?2011
?????
22111

10000
00000
aOOOO
OaOOa
aOOOO
10001
10001
10000
10001
11 001
11 001
00001
01011??O? 1
01101
01001
????1
01 001

00000
00000
00000
00000
aOOOO
10000
10000
00000
11100
111 00111 0011011
11011
11 ?OO
11000
11000
21000
21000

00000
00000
00000
00000
00000
00101
00101
00101
00101
00101
00101
00011
00011
00001
100?1
11001
11??1
11001

00000
00000
00000
OOaOO
00000
10000
10000
10000
11011
11 0111111 0
11000
11100
1???O
11100
11100
11???
111 01

00000
00000
00000
00000
aOOOO
00111
00110
00000
21000
00100
001 00
00000
20100
10000
10100
21100
20100
20100

00000
OO??O
OOOcO
aOaOa
OOOOa
001 0011110
001 0010110
11 01 0
11 01 0
00020
10120
00020
00021
00020
?????
00002

OaOOO
?OOOO
OaOOO
aOooo
aOOOO
01 000
01000
00000
02000
22000
22000
?2000
12000
01000
11001
03000
??1??
0311 0

00001
00000
aOOOO
a0001
OOOOa
OOOaO
10010
0001 000110
11001
11 001
00010
01010
0011 0
00010
00010
?????
0001 02000a
001 02
Oab01
bOOOO
OOdOO
01201
01201
01201
01102
111 00111 00
01002
01011
01002
01002
01003
?????
01 003

10000
O??OO
1aaaO
10000
1aaOO
11110
11110
111 00
11110
00101
00101
11110
11111
11 ? 10
11110
11110
?????
1111 0

00001
0?001
Oa001
00001
Oaa01
01 001
01001
01 00101101
11 011
11 011011?1
01111
011 ? 1
01101
01101
?????
11101

Ren Hirayama and Tsutomu Chitoku

620

APPENDIX II. Characters and character states*: character first identified here.
28.

Skull
1.

2.
3.
4.
5.
6.
7.
8.

9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.

20.
21.
22.
23.
24.
25.
26.
27.

Megacephalic: (0)= no, (1)= yes. Cranial length
reaching up to one third or more of shell length(Boulenger, 1889; Hirayama, in press).
Cranial scute sulci on dermal roofing elements:(0) = present, (1) = absent.
Nasal bone: (O)=present, (1)=lost.
Medial meeting of prefrontals: (0)= absent, (1)=present.
Prefrontal-postorbital contact: (0)= absent, (1)=present.
Orientation of orbital opening: (0)= lateral, (1)=dorso lateral (Fastovsky, 1985).
Parietal-squamosal contact: (0)= present, (1)=absent.
Posterior temporal emargination: (0)= foramen
stapedio temporale concealed from dorsal, (1)=foramen stapedio temporale exposed from dorsal,(2)=entire of processus trochlearis oticum
exposed from dorsal (Gaffney, 1979).
Jugal-quadrate contact: (0)= absent, (1)=present.
Jugal-squamosal contact: (O)=absent, (1)= present.
Medial process of jugal: (0)= present, (1)= lost.
Jugal-pterygoid contact: (0)= present, (1)= lost
Hooked premaxillae: (O)=absent, (1)=present.
Foramen praepalatinum: (O)=present, (1)= lost.
Upper triturating surface involving palatines:(O)=no, (1)= yes.
Upper triturating surface involving vomer: (0)=no, (1)= yes.
Vomerine trough on upper triturating surface:(O)=absent, (1)= present (Kesteven, 1910).
Vomerine pillar concealed by upper triturating
surface: (O)=absent, (1)= present (Zangerl, 1971).
Apertura narium interna entirely formed by vomer
and palatines, excluding premaxillae: (0)= no,(1)= yes.
Pit between premaxillae and vomer: (0)= absent,(1)= present (Gaffney and Meylan, 1988).
Meatus choanae at level of medial process of
jugal: (O)=absent, (1)=present (Zangerl, 1971).
High lingual ridge of maxilla exposed fromlateral*: (O)=absent, (1)=present.
Vomer-pterygoid contact lost, palatines medially
meeting: (O)=no, (1)= yes.
Foramen palatinum posterius enclosed: (0) =present, (1)= lost.
Foramen pa/atinum posterius open posterolateralIy: (0)= absent, (1) = present.
Processus pterygoideus externus: (0)= present,(1)= lost.
Median ventral ridge of pterygoids*: (0)= absent,(1)= present.

29.
30.

31.

32.
33.

34.
35.
36.
37.
38.
39.

40.

41.
42.

43.
44.
45.
46.

47.

Pterygoid extending onto mandibular articulating
surface of quadrate: (O)=no, (1)=yes.
Pterygoid narrow, C-shaped: (O)=no, (1)=yes.
Large processus pterygoideus trochlearis oticum
involving deep notch of quadrate: (0)= no, (1)=yes.
Processus pterygoideus trochlearis oticum involving prominent shelf of prooitc: (0)= yes, (1)=no(Gaffney, 1979).
Incisura collumerae auris closed: (O)=no, (1)=yes(Gaffney, 1979).
Eustachian tube enclosed within elongate incisura collumerae auris auris: (0)= no, (1)=yes(Gaffney, 1979).
Cavum tympani with posterior ventral ridge*:(O)=absent, (1)=present.
Crista supraoccipitalis little projecting beyond
parietal: (O)=no, (1)= yes.
Canalis caroticus internus excluding basisphenoid: (O)=no, (1)=yes (Gaffney, 1979).
Foramen anterius canalis carotici interni lie close
together: (O)=no, (1)= yes.
Foramen basisphenoidale: (0)= present, (1)=absent (Meylan and Gaffney, 1989).
Foramen canalis carotici interni posterius separated from foramen basisphenoidale, enclosed withinpterygoid: (O)=no, (1) = yes.
Ventral surface of basisphenoid with V-shaped
crest emarginated from posterior: (0)= absent,(1) = present.
V-shaped basisphenoid with basipterygoid projection: (0)= absent, (1)= present (Kesteven, 1910).
Large basisphenoid, lying beneath the siginificant
portion of pterygoids: (O)=no, (1)=yes.
High dorsum sellae, not concealing posterior
portion of sella turcia: (O)=no, (1)=yes.
Single rod-like ossification of rostrum basisphenoidale: (0)= absent, (1) = present.
Ossification of rostrum basisphenoidale reduced:(O)=no, (1)=yes (ossification lost in Dermochelys).
Foramen caroticum lateraie much larger thanforamen anterius canalis carotici interni: (0) no;(1) yes.
Foramen caroticum laterale confluent with canalis
cavernosum: (O)=no, (1)=yes.

Lower Jaw
48. Lower triturating surface of dentary enlarged, withsymphysis extending along more than one third ofentire jaw length: (O)=no, (1)= yes (Zangerl, 1971).
49. Hooked dentary: (0)= absent, (1)=present.
50. Symphyseal ridge of dentary: (0)= absent, (1)=present (Gaffney, 1979).
51. High symphyseal ridge of dentary exposed fromlateral beyond labial ridge*: (0)= absent, (1)=1022.

52.
53.
54.
55.

56.
57.
58.

Cretaceous Dermochelyidae from Japan

present.
Denticulated lingual ridge of dentary: (0)=absent, (1)= present.
Lingual ridge of dentary higher than labial ridge' :(O)=no, (1)=yes.
Lingual ridge of dentary: (0)= present, (1)=reduced and lost (Gaffney, 1979).
Foramen dentofaciate majus anterior to lower jaw
symphysis: (0)= no, (1)=yes (Zangerl, 1971).
Surangular process extending anteriorly onto
dentary: (0)= absent, (1)=present.
Coronoid and articular ossification: (0)= present,(1)= lost.
Splenial: (0)= present, (1)= lost (Gaffney, 1979).

Axial Skeleton
59. Cervical rib: (0)= present, (1)= lost (Gaffney et at.,1991).
60. Transverse process of cervicals: (0)= double,(1)= single (Gaffney, 1985a).
61. Central articulation of cervicals formed: (0)= no,amphicoelous or platycoelous, (1)=yes, concavoconvex (Gaffney and Meylan, 1988).
62. Proportion of cervical central articulation: (O)=as
high as wide thoroughout cervicals, (1)= muchwider than high at posterior cervicals (Williams,
1950).
63. Posterior cervicals with thin ventral keel: (0)=absent, (1)= present (Meylan and Gaffney, 1989).
64. Biconvex centrum of anterior cervicals: (0)=absent, (1)=4th cervical, (2)=3rd or 2nd cervical(Williams, 1950).
65. 8th cervical: (O)=amphicoelous, (1)=procoelous,(2)=biconvex, (3)=opistocoelous (Williams, 1950;
Meylan and Gaffney, 1989).
66. 8th cervical centrum much shorter than its neuralarch and 7th centrum: (0) = no, 8th centrum aslong as its neural arch and 7th centrum, (1)=yes(Zangerl, 1953).
67. Double cervical central articulation between 7th
and 8th cervicals: (0)= absent, (1)= present.
68. Double cervical central articulation between 6th
and 7th cervicals: (0)= absent, (1)=present (Williams, 1950).
69. Double cervical central articulation between 5th
and 6th cervicals: (0)= absent, (1)=present (WilIiams, 1950).
70. Platycoelous cervical central articulation between
6th and 7th cervicals: (0)= absent, (1)= present.
71. Anterior central articulation of first thoracic vertebra: (O)=facing anteriorly, (1)=facing ventrally or
anteroventrally.
72. First thoracic rib: (0)= distal ends extending
beyond nUChal, (1)= distal ends limited withinnuchal width (Rieppel, 1980; Gaffney andMeylan, 1988).
73. 10th thoracic rib: (O)=reaching 8th pleural, (1)=free from carapace (Meylan and Gaffney, 1989).
74. Chevrons: (0)= present, (1)= lost (Gaffney and

75.
76.

621

Meylan, 1988).
Opistocoelous caudal: (O)=absent, (1)=present.
Procoelous caudal: (0)= absent, (1)= present only
within anterior portion, (2)=all caudals procoelous(Gaffney and Meylan, 1988).

Appendicular Skeleton
77. Coracoid: (0)= shorter than humerus, (1)=as longas humerus, (2)= much longer than humerus.
78. Elongate coracoid thickened, showing coulm-likeappearance: (0)= no, flattened, (1)=yes.
79. Scapular angle formed by scapular prong andacromion: (O)=about 90°, (1)=exceeding over110°.
80. Lateral process of pubis or pectineal process:(0)= restricted posteriorly to medial portion of
pubis, (1)= enlarged, projecting anteriorly beyond
medial portion of pubis.
81. Thyroid fenestra: (0)= small, subdivided, (1)=huge, confluent.
82. Lateral process of ischium or metischial process :(0)= prominent, as wide as whole iscium, (1)=reduced, often rudimentary.
83. Iliac blade: (0)= nearly straight, (1)= elongate,with medial curvature.
84. Large pelvic girdle, nearly touching coracoids:(0)= no, distant from coracoids, (1)=yes.
85. Humerus: (O)=shorter than femur, (1)= longer
than femur.
86. Location of lateral process of humerus: (O)=atlevel of caput humeri, (1)=distal to caput humeri,(2)= at middle of humeral shaft (Walker, 1973).
87. Shoulder of caput humeri: (0)= present, (1)= lost
(Hirayama, 1992).
88. Lateral process of humerus enlarged into Vshaped structure: (O)=no, (1)=yes.
89. Lateral process of humerus enlarged within theanterior portion of shaft, not well visible fromventral view: (O)=no, (1)=yes.
90. A median concavity of lateral process of humerus: (0)= absent, (1)=present.
91. Elongate lateral process of humerus, extending
into prominent anterior projection: (0)= no, (1)=yes.
92. Ectepicondylar portion of humerus much enlarged
anteriorly: (O)=no, (1)= yes.
93. Ulna-radius contact through prominent distalrugosities: (0)= absent, (1)=present.
94. Radius with middle portion bent toward anterior:(0)= no, nearly straight, (1)= yes.
95. Carpal and tarsal elements flattened, withrestricted joint surfaces: (0)= no, (1)= yes.
96. Elongate 3rd to 5th digits of manus and pes,
without movable articulation: (0)= no, all movable
articulation retained, (1)= yes.
97. Movable articulation of first and second digits:(0)= retained, (1)= lost.
98. Femoral trochanters: (0) = separated by notch,(1) = connected by bony ridge.

Ren Hirayama and Tsutomu Chitoku

622

99.

Major trochanter of femur: (O)=as high as orslightly higher than minor trochanter, (1)= muchhigher than minor trochanter (Hirayama, 1992).
100. Pit on tibia for M. pubi-tibialis and M. flexor tibialis
internus: (0)= absent, (1)= present (Zangerl, 1980).

Carapace101. Scute sulci on carapace: (0)= present, (1)=remained only at neural and marginal area, (2)=entirely lost.
102. Nuchal with prominent anterior emargination:(0)= absent, (1)= present.
103. Nuchal with ventral knob to which the neural archof 8th cervical attaching: (0)= absent, (1)=present.
104. Posterior nuchal fontanelles: (0)= absent, (1)=present.
105. Cervical scute: (0)= narrower than 1st vertebral
scute, (1)=as broad as first vertebral, preventing
contact between first vertebral and first marginal(Hirayama, in press).
106. Neurals thickened and keeled: (O)=no, (1)=yes.
107. Epineural elements between neurals: (0)=absent, (1)= present.
108. Number of neurals*: (O)=eight or less, (1)=nineby splitting of 8th or 7th neurals, (2)=more thanten.
109. Neural shape: (0)= mostly hexagonal, shortsided in anterior, (1)= mostly hexagonal, equallysided, (2)= mostly rectangular (Hirayama, in press).
110. Neural series: (0)= complete, (1)=reduced andlost at posterior portion, (2) = entirely lost.
111. Suprapygal*: (O)=subdivided into two or three,(1)=single.
112. pleural reduction: (0)= lost in adult, (1)= partallypresent, (2)=developed less than halfway to theedge of costal ribs, (3)=entirely lost (Hirayama,
1995).
113. Epithecal ossification of carapace: (0)= absent,(1)= present.
114. peripherals: (0)= present, (1)= lost.
115. prominent undulation along medial border of
peripherals between costal ribs: (0)= absent, (1)=present.
116. peripherals with prominent lateral serration: (0)=absent, (1)= present (Zangerl, 1953).
117. 9th costal separated from 8th costal: (0)=absent, (1)= present (Zangerl, 1953).
118. Suprapygal-pygal contact: (0)= present, (1)= lost.

Plastron
119. Scute sulci on plastron: (0)= present, (1)= lost or
rudimentary (Zangerl, 1953).
120. Axillary buttress: (0)= reaching only peripheral, or
connected with carapace by ligament, (1)= reaching 1st pleural (Meylan and Gaffney, 1989).
121. Inguinal buttress: (0)= reaching only peripheral,or connected with carapace by ligament, (1)=reaching 5th pleural, (2)= reaching 5th and 6th
pleural (Hirayama, 1985).
122. Medial connection of hyo-hypoplastra*: (0)=sutured, (1)= ligamentous.
123. pi astral index (ratio between length and width ofplastral bridge): (0)= more than 100, (1)= between
100 and 60, (2)= less than 60 (Zangerl, 1953).
124. Star-shaped hyo-hypoplastra: (0)= absent, (1)=present.
125. Fontanelle between hyo-hypoplastra: (0)=absent, (1)=smaller than hyo-, or hypoplastron,(1)=as large as hyo, or hypoplastron, (2)= huge,remaining rudimentary hyo-hypoplastra.
126. Dorsal process of epiplastra: (0)= present, (1)=lost (Gaffney and Meylan, 1988).
127. Narrow, blade- like epiplastron: (0)= absent, (1)=present (Zangerl, 1953).
128. Narrow, elongate entoplastron: (0)= absent, (1)=present (Zangerl, 1953).
129. Entoplastron: (0)= connected with hyoplastra by
suture, (1)=connected with hyoplastra by ligament (Zangerl, 1953).
130. Epi-entoplastra fused, T -shaped: (0)= no,epientoplastra separated, (1)=yes.
131. Entoplastron: (0)= present, (1)= lost.
132. Narrow, blade-like xiphiplastra: (0)= no, (1)=yes(Zangerl, 1953).
133. Medial contact between xiphiplastra: (O)=developed along its whole length, (1)= much reduced orlost (Zangerl, 1953).
134. Direction of xiphiplastron: (0)= nearly straight,
extending posteriorly, (1)=curved toward medial(Zangerl, 1953).
135. Mesoplastron: (0)= present, (1)= lost (Gaffney
and Meylan, 1988).

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 184, pp. 623-650, 11 Figs., December 30 1996'

© by the Palaeontological Society of Japan 1996

Permian fusulinaceans from the Akuda andHorikoshitoge Formations, Hachiman town,
Gifu Prefecture, Central JapanHISAHARU IGO
Department of Astronomy and Earth Sciences, Tokyo Gakugei University, Koganei, Tokyo, 184 Japan

Received 18 May 1996; Revised manuscript accepted 15 August 1996

Abstract. Thirty-seven species belonging to 16 genera of fusulinaceans are discriminated from the Akuda
and Horikoshitoge Formations, exposed in Hachiman town, Gifu Prefecture, Central Japan. Among them,
the following six species are newly proposed: Sphaerulina akudensis, Dunbarula uenoi, Schwagerina
shukoae, S. kanumai, Pseudofusulina minoensis, and P. miyamensis. This fusulinacean fauna is similar
in composition to that described from the Nyukawa Group exposed about 60 km northeast of the present
area, and indicates the Yakhtashian to Kubergandian of the standard stratigraphic scheme of the Tethyan
Province.
Key words: Akuda Formation, fusulinaceans, Horikoshitoge Formation, Mino Terrane, Permian
Introduction
The study area is located in Hachiman town, Gujyo
County, Gifu Prefecture, Central Japan. This area belongs
to the northern part of the Mino Terrane of the Inner Zone ofSouthwest Japan, and is underlain by Jurassic clastic sedimentary rocks. The strata consist of turbidites and olistostromes that commonly include allochthonous blocks ofchert, limestone, and greenstone of various age, shape, andsize. The chert blocks yield Triassic conodonts andradiolarians. The limestone blocks are commonly fossiliferous and yield calcareous algae, fusulinaceans, andconodonts that suggest an Early to Middle Permian age.
Kanuma (1958a, b, 1959, 1960) extensively studied this areaand divided the strata into four formations, namely, theAkuda, Kuchibora, Kayukawa, and Shimadani Formations inascending order. The Akuda and Kuchibora Formations
consist mainly of white to pale gray massive limestone andblack bedded limestone, respectively. The Kayukawa Formation consists mainly of clastic sedimentary rocks. The
Shimadani Formation is composed of bedded chert. Kanuma(1958a, b) described late Early Permian fusulinaceans collected from the limestone of the Akuda Formation.
Subsequently, Yoshida (1972) studied the structural geology of this area and analyzed some complicated folding
structures. I restudied this area and discovered many
Triassic conodonts from cherts of the Kayukawa and Kuchibora Formations (Hh. Igo, 1979) and provided new information concerning the geologic age of these formations.
Furthermore, I confirmed that the cherts are allochthonous
blocks or sheets embedded in both formations.

Wakita (1984) made regional rnapping in the Hachimanarea and pointed out that Jurassic clastic sedimentary rocks
are extensively distributed in this area, intercalating allochthonous blocks of various kinds of sedimentary rocks. Moreover, these blocks show various sizes and ages. He subdivided the Jurassic rocks into five such units, the Nabigawa,
Kodaragawa, Kajika, Furumichi, and Samondake Formations.
Concerning the limestone of the Akuda Formation, heshowed that it is a huge allochthonous block embedded in
the Nabigawa Formation and it is designated as the Akuda
Block.
I continued investigation of this area and reported a mixed
conodont fauna preserved within fissure-filling deposits inthe limestone of the Akuda Block (Hh. Igo, 1989). Furthermore, I discussed the origin of this mixing related with thesedimentary environment of the Akuda Limestone. Recently, Horibo (1990) documented the petrography of limestones
which belong to the Akuda and Horikoshitoge Formations,and discussed their depositional environments.
During my consecutive field survey in the Hachiman area,
I have collected many limestone samples from the Akuda
and Horikoshitoge Formations in which abundantfusulinaceans are contained. Paleontological study of thesamples elucidates that the fusulinaceans are similar tohitherto described species from allochthonous limestones
embedded in the accreted Jurassic sedimentary rocks in the
Mino Terrane, Central Japan. The purpose of the present
paper is to describe the fusulinaceans in these two formations and to discuss the age and similarities of the fauna withthat reported from other allochthonous limestones in the
Mino Terrane. I discriminate a total of 37 species ofHisaharu Igo

624

fusulinaceans belonging to 16 genera in this study, among
which six species are new to science and four species left
in open nomenclature.

,"'-';,:"i

Sea of Japan

Geologic settingI

The Akuda Block of Wakita (1984) is an informallydesignated rock unit of limestone previously treated as aformation, the Akuda Formation (Kanuma, 1958, etc. ; Hh.
Igo, 1989). Recently, Horibo (1990) restudied Wakita's Akuda
Block based on the limestone petrography and analysis of
biofacies. He subdivided the Akuda Block into the Akuda(s.s) and Horikoshitoge (newly designated) Formations by
their lithic characters. Furthermore, Horibo showed thatthese formations are almost coeval but that the former was
deposited in lagoons or sand bars while the latter accumulated in the marginal part of the bank. The two formations are
now in fault contact.
In the present study, I follow Horibo's newly proposed
definition of the Akuda and Horikoshitoge Formations. The
Akuda Formation constitutes the main part of the Akuda
Block, which is about 1.7 km in width and 12 km in lateralextent. It strikes E-W and dips 80· to 90·S or N. According
to my observations, the Akuda Formation consists mainly ofwhite to pale gray massive and thick-bedded limestones
including limestone conglomerate and calcarenite, and variegated greenstones and volcaniclastics. The total thick-. -~\ ;. ·. '. t.., . .
\ / ......... -_:' '......,..... \,

,

.'

~I

~_.-,'. :},, --,,);-'_/--"'-~:;.-'~,/<'--;~~m'" ,~/_I)\._, ,"'---"") "-:!. \ ....y ..

\(""':""
}

i

37"

"

,:~::~; ;"'~~"".

.... _' I..... / ...... _--'-~
\

..:

-:.

"'r}-"'·"·

--'
'. ;.

I Pacific Ocean :1,3fi·

1Ja

D

1~O

o

I L........,§Okm
I

140D

14f

Figure 1. Index map of Hachiman town, Gujyo County,
Gifu Prefecture.

ness of this formation was estimated to be 550 m by Hh. Igo(1989) and 500 m by Horibo (1990). The Akuda is well
exposed along the roadcut from Nishi-Akuda to Akuda-

Figure 2. Map showing fusulinacean localities.

1023.

Permian fusulinaceans from Central Japan

Kuchi (Sections 8, C, E) and north of the Horikoshi Pass
(Section A). Limestones in the Sections 8 and E are mostly
pale gray to white and massive. Limestones along theSection C (8-17, 8-11) are bedded and mostly dark gray withbrownish tints.
Intercalations of limestone conglomerate (Horibo's limestone-breccia) are characteristic lithofacies of the Akuda
Formation. This conglomerate consists of various-sizedangular clasts of limestone and dolostone cemented with iIIsorted lime mud, lime sand, and red hematitic clay. The
other type of conglomerate is volcanic conglomerate
exposed along the roadcut at Nishi-Akuda (R-1). Limestone
clasts of the conglomerate are commonly rounded and ofboulder size cemented with variegated tuft.
The Horikoshitoge Formation at Higashi-Akuda consists
mainly of gray to dark gray bioclastic bedded limestone, andattains a thickness of about 200 m. Along the Section D,
limestones (8-37 to 8-48) show typical lithology of the
Horikoshitoge and their microfacies are lime mud/wacke-m625
Secti on E

Secti on D

6-46-

6-47-

R-2

6-466-44-

R-l
6-42-

6-37-

massive limestone

E2525C9 thick-bedded~ limestone

vol cani cl asti cs

SectionC

Figure 4. Geologic columns of the measured section
with indications of fusulinacean-bearing levels. Section D is
located near Nishi-Akuda; Section E is located at HigashiAkuda refer to Figure 2. For lithologic symbols see Figure 3.

~:~if~ limestone conglomerate.:~~'!9.>:,:<:! (tuffaceous matri x)

stone.

These limestones yield abundant fusulinaceans.

Secti on B
Fusulinaceans from the Akuda andHorikoshitoge Formations

Secti on A
6 -2 -~':'2::23

6 -3 -t:~z:::I
6-116-4-h~~

[

A-22

6-17 -1~:::;:X:rI

A-21-

Figure 3. Geologie columns of the measured section
with indications of fusulinacean-bearing levels. Section A is
located north of the Horikoshi Pass; Sections Band Carelocated west of Akuda-Kuchi refer to Figure 2.

Volcanic conglomerate crops out at Nishi-Akuda (R-1) andabundantly includes limestone clasts containing well-preserved fusulinaceans. This conglomerate is assumed to beequivalent with limestone-breccia in the upper part of theColumn 8 (Section 71) of Horibo (1990). This breccia
occupies the upper part of the middle division of the Akuda
Formation. The following fusulinaceans are discriminatedfrom R-1: Pseudoreichelina sp., Pamirina (Levenella) /eveni
Kobayashi, P. (L.) evo/uta Sheng and Sun, Nankinella sp., N.
aft. kotakiensis (Fujimoto and Kawada), Sphaerulina akudensis Hh. Igo sp. nov., Schubertella pseudoobscura Chen,
Neofusulinella giraudi Deprat, Schwagerina shukoae Hh. Igo
sp. nov., S. higashidaniensis Hy. Igo, S. kanumai Hh. Igo sp.
nov., Pseudofusulina minoensis Hh. Igo sp. nov., P. miyamensis Hh. Igo sp. nov., P. regu/aris (Schellwien), P. houziguanica
Sheng, and P. sp. Massive limestone (R-2) situated about 30
m above R-1 yields P. regu/aris (Schellwien) and Cha/aroschwagerina aft. chinensis (Chen). A single species, S.
kwangchiensis Chen, was obtained from R-3 situated about
60 m below R-t
Although the actual stratigraphic position is uncertain, I
collected some samples from limestones exposed south of
the Horikoshi Pass (A-1, A-2). These samples may repre-

626

Hisaharu Igo

sent the lower part of the Akuda Formation and contain

Pseudofusulina norikurensis Hy. Igo and P. paratschernyschewi Kanuma. Collections were also made along the roadcut (Section A) north of the Horikoshi Pass. Limestones
situated in the lower part of this section (A-21 , A-22) yield
Neofusulinefla /antenoisi Oeprat, Minojapanefla (M.) e/ongata
Fujimoto and Kanuma, Dunbaru/a uenoi Hh. Igo sp. nov.,
Pseudofusulina horadaniensis Hy. Igo, Mak/aya pamirica(Leven), M. saraburiensis Kanmera and Toriyama, and Armenia sp.
Limestone samples were collected along the Section 8,
about 300 m north of the above-mentioned localities. They
are at levels 8-2, 8-3, and 8-4 and yield Schwagerina
shukoae Hh. Igo sp. nov., Acervoschwagerina fujimotoi
Kanuma, and others. Massive limestone in the lower part of
the Section C (8-1) includes Parafusulina ? sublineata Hy. Igo
and thickly bedded limestone in the upper part of this section
yields Parafusulina sotensis Hy. Igo and others (8-11).
A single species, Pseudofusulina kraffti (Schellwien), wascollected from a white massive limestone exposed at OHT4, upstream of the Otohime-tani Valley, north of NishiAkuda.
I collected abundant fusulinacean specimens from thinbedded limestones (8-37, 8-42, 8-44, 8-46, 8-47, 8-48)
exposed at Higashi-Akuda (Section 0). These limestones
are equivalent with the lower member of the Horikoshitoge
Formation (Column A, section 59 of Horibo, 1990). The
following fusulinaceans are discriminated from these levels:
Pamirina (Levenella) evo/uta Leven, Neofusulinella giraudi
Oeprat, Toriyamaia /axiseptata Kanmera, Schwagerina muraii
Morikawa, Pseudofusulina norikurensis Hy. Igo, P. paratschernyschewi Kanuma, P. horadaniensis Hy. Igo, P. isomiae Hy.
Igo, P. duplithecata Hy. Igo, P. regu/aris (Schellwien), P. krotowi (Schellwien), and others.
As already mentioned, the Akuda and Horikoshitoge
Formations are a huge allochthonous block embedded inJurassic rocks, and the detailed internal structure of thisblock is unclear. I measured several sections to collectfusulinaceans, but these specimens do not provide enoughbiostratigraphic value, and even the local correlation among
measured sections is difficult. Fusulinaceans discriminatedin this study, however, indicate that the formations are almost
coeval as already pointed out by Horibo (1990).
The present fusulinacean fauna is characterized by
species of Pseudofusulina with a large cylindrical shell, thickspirotheca, and dense axial filling. Characteristic Schwagerina species with a fusiform shell of medium size, thin spirotheca, and delicate septa are also important representatives.
A similar schwagerinid but with an elongate shell identified toParafusu/ina with reservations, is also characteristic.
Although its occurrence is rare, the presence of Acervoschwagerina in this fauna is noteworthy. The fauna also
contains Neofusulinefla, Sphaeru/ina, Schubertefla, Minojapanefla, Nankinefla, and other small fusulinaceans. The
mentioned faunal characters are common features in thefauna reported from the upper Lower Permian (yakhtashian)of the Tethyan Realm. The present fauna excludes typicalParafusu/ina speCies except for Parafusulina sotensis andspeCies of Misellina, which are good indicators of the

80lorian, are also lacking.
In the Section A, limestones of two different levels yield
two species of Mak/aya, and an unidentified species ofArmenia in association with Neofusulinefla and others.
These levels may occupy higher levels than those of theaforementioned species and are correlated with the middle
Middle Permian (Kubergandian).
The Jurassic accreted terrane, the Mino Terrane, inCentral Japan commonly includes various-sized Upper
Paleozoic limestones. Previous to our recognition of accreted terrane schemes, these limestones had long been
considered to be autochthonous strata which were deposited in the shallow carbonate shelf, bank, reef, and othermarine geosynclinal environments. Fusulinaceans of theselimestones were studied by many specialists in the Lake
8iwa area (Morikawa and Isomi, 1961), Akasaka Limestone(Ozawa, 1927, Morikawa, 1958, etc.), Ibuki Limestone(Kobayashi, 1957), Yoganeyama Limestone (Sashida, 1980),
Hachiman area (Kanuma, 1958a, etc.), Nyukawa Group (Hy.
Igo, 1964, etc.), Shirahone Limestone (Choi and Fujita, 1970),
and others. There are other fusulinacean-bearing limestonesin the Mino Terrane of Central Japan such as the Fujiwaradake, Ryozen, and Funafuseyama Limestones.
Fusulinaceans contained in these limestones have not yet
been fully studied, but the present fauna is similar to thefauna described from the Shiroi, Kono, and lower Sote
Formations of the Nyukawa Group (Hy. Igo, 1964, 1965)
exposed about 60 km northeast of the present area.

Systematic paleontology
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea von Moller, 1878
Family Staffellidae Miklukho-Maklay, 1949
Genus Pseudoreichelina Leven, 1970

Type species.-Pseudoreichelina darvasica Leven, 1970

Pseudoreichelina sp.
Figures 10-15-20

Materia/s.-Axial section, TGUFU 1086; tangential sections, TGUFU 1084, 1085, 1087, 1089; sagittal section,TGUFU 1088.
Remarks.- The present specimens are mostly recrystallized and the spirothecal structure and other specific characters are obliterated. The shape of septa in the last uncoiled
volution of the specimens indicates assignment to the genus
Pseudoreichelina
Occurrence.-AII specimens collected from R-1 in Section
E of Nishi-Akuda.
Genus Pamirina Leven, 1970 emend. Kobayashi, 1977

Type species.-Pamirina darvasica Leven, 1970
Subgenus Levenia Ueno, 1991
Subgenus Levenella Ueno, 1994

1023.

Permian fusulinaceans from Central Japan

Type species.-Pamirina /eveni Kobayashi, 1977

Pamirina (Levenella) leveni Kobayashi, 1977
Figures 11-20-31
Pamirina leveni Kobayashi, 1977, p. 11-14, pI. 1, figs. 13-38.
Pamirina (Levenia) leveni Kobayashi. Ueno, 1991, p.746, 747,
figs. 3-8-23.
Pamirina (Levenella) leveni Kobayashi. Ueno, 1994, p. 405.
Materia/s.-Axial sections, TGUFU 1110,1111, 1112,1113,1114,
1115,1116,1117,1120,1121; sagittal sections, TGUFU 1118,1119.
Remarks.-Ueno (1991) proposed the subgenus Levenia,
which is characterized by a finer alveolar keriotheca thanthat of Parnirina Subsequently, Ueno (1994) revised thegenus name Levenia to Levenella. The Akuda specimens
quite agree with the original ones described by Kobayashi(1977) from the Hanagiri Limestone and Otaki, Saitama
Prefecture, Central Japan.
Occurrence.-AII specimens collected from R-1 in Section
E of Nishi-Akuda.

Pamirina (Levenella) evoluta
Sheng and Sun, 1975
Figures 11-32-34
Pamirina ? evoluta Sheng and Sun, 1975, p.46, pI. 1, fig. 15.
Pamirina (Levenella) evoluta Sheng and Sun. Ueno, 1991, p. 747,
748, figs. 4-1-8.
Materia/s.-Axial sections, TGUFU 1122,1123,1124.
Remarks.-Recently, this species has been fully describedby Ueno (1991) on the materials from the Akiyoshi LimestoneGroup. It has a larger proloculus than P. (L.) /eveni. The
specimens referable to this species are very rare in mycollections.
Occurrence.-TGUFU 1122 from 8-47 in Section D near
Higashi-Akuda; TGUFU 1123,1124 from R-1 in Section E of
Nishi-Akuda.
Family Staffellidae Miklukho-Maklay, 1949
Genus Nankinella Lee, 1934
Type species.-Staffella discoides Lee, 1931

Nankinella sp.
Figure 11-16
Materia/.-Tangential section, TGUFU 1106.
Remarks.-The present specimen is completely recrystallized and the inner structure of the shell is obscure, but theangular periphery and umbilicated axial areas indicate
assignment to the genus Nankinella
Occurrence.-One specimen from R-1 in Section E of
Nishi-Akuda.

627

Nankinella aff. kotakiensis(Fujimoto and Kawada, 1953)
Figures 11-17-19
Compare.Hayasakaina kotakiensis Fujimoto and Kawada, 1953, p.119-121,
figs. 1-10; Hy. Igo, 1956a, p. 172, 173, pI. 27, figs. 1, 6, 12-16.
Nankinella kotakiensis (Fujimoto and Kawada). Kanmera, 1958,
p.208-210, pI. 28, figs. 9-13.
Materia/s.-Axial sections, TGUFU 1107,1108,1109.
Description.-Shell very small, discoidal, subspherical withsomewhat broadly rounded periphery. Mature shell consistsof seven to eight volutions. In two specimens, axial length
1.83 to 1.86 mm, median width 1.83 to 1.86 mm, and form ratio
0.716 to 0.749.
Shell mostly recrystallized. Inner four volutions discoidal
and planispiral having slightly angular periphery. Spirothecathin, consisting of a tectum and recrystallized translucent
layer. Proloculus very small and spherical. Chomata present in inner three to four volutions.
Remarks.-The present specimens closely resemble thetype specimens of Nankinella kotakiensis, but the former
differs from the latter in more angular periphery in inner
volutions. Nankinella orientalis Miklukho-Maklay resembles
the present specimens but differs from the latter in fewer
volutions. Nankinella inflata (Colani) also resembles thepresent specimens but the former has a large shell and moreangular periphery in inner volutions.
Occurrence.-AII specimens from R-1 in Section E.
Genus Sphaerulina Lee, 1933
Type species.-Sphaerulina crassispira Lee, 1933

Sphaerulina akudensis Hh. Igo sp. nov.
Figures 11-9-14
Materia/s.-Axial section of holotype, TGUFU 1099; axial
section of paratype, TGUFU 1104; tangential sections,TGUFU 1100, 1101 ; sagittal sections, TGUFU 1102, 1103.
Diagnosis.-Small Sphaerulina with an inflated oval shell,
thick wall, fewer volutions, and unfluted septa.
Description.-Shell small, discoidal, and subspherical withslightly umbilicated axial regions and rounded periphery.
Mature shell having six volutions, 1.03 mm in axial length,1.40 mm in median width, and form ratio 0.736 in holotype.
Inner two volutions tightly coiled and have narrowly
rounded periphery. Outer volutions expand rather rapidly.
Radius vectors of holotype in first to sixth volutions 0.08, 0.15,
0.25, 0.38, 0.53, and 0.73 mm respectively. Proloculus verysmall, spherical, and having external diameter of 0.075 mm inholotype.
Spirotheca rather thick compared with shell size. It
consists of a tectum and recrystallized dark calcite layer. In
spirotheca of outer volutions except last volution appears adiaphanotheca-like transparent layer. Septa rather thick,plane, and having a lighter calcite layer than that of spirotheca. Chomata low, small, and asymmetrical.

Hisaharu Igo

628
Table 112
3
4
5Reg. no.

Fig.

TGUFU1099
TGUFU1100
TGUFU1102
TGUFU1103
TGUFU1104

11- 9
11-10
11-12
11-13
11-14

Measurements of Sphaerulina akudensis Hh Igo sp nov. (in mm)
Radius vector
W.
F.R.
D.P.
L.
4
1
2
3
1.40
1.40
1.50
1.03
0.83

1.03
0.93
...
---

0.68

0.74
0.66
---

--0.82

0.08
?
0.20
0.06
0.06

0.08
?
0.13
0.08
0.08

0.15
0.09
0.18
0.15
0.14

0.25
0.28
0.28
0.24
0.23

0.38
0.38
0.43
0.36
0.30

5

6

0.53
0.53
0.60
0.50
0.45

0.73
0.73
-----

---

Thickness of spirotheca
1
2
3
4
5

1
?
?
?
0.02
?

2
?
?
0.02
0.02
?

3

4

5

6

0.02
?
0.03
0.04
0.02

0.02
0.05
0.05
0.05
0.03

0.05
0.06
0.04
0.03
0.04

0.05
0.08

-------

Remarks.-This new species resembles a species hitherto
described as a Pamirina, but the former has a larger shell andthicker spirotheca than the latter. Recently, Hy. Igo et a/.
(1993) described Sphaerulina cf. croatica Kochansky-Devide
from the Wang Saphung Formation of northeastern Thailand.
The Thailand species resembles S. akudensis but has an
inflated shell and distinct chomata.
Etym%gy.-This species name is derived from a local
name of Hachiman town.
Occurrence.-AII specimens from R-1 in Section E.
Family Schubertellidae Skinner, 1931
Subfamily Schubertellinae Skinner, 1931
Genus Schubertel/a Staff and Wedekind, 1910
Type species.-Schubertella transitoria Staff and Wedekind,1910

Schubertella pseudoobscura Chen, 1934
Figure 11-15
SChubertella pseudoobscura Chen, 1934, p. 16, 17, pI. 1, figs. 1, 2;
Lin, 1984, p. 153, pI. 8, figs. 25, 26.
Material.-Axial section, TGUFU 1105.
Oescription.-Shell small and almost spherical with slightly
umbilicated axial regions. This speCimen has four volutions,0.53 mm in axial length, 0.54 mm in width, and form ratio
0.98. Inner first to second volutions planispiral and COilingat a right angle to axis of outer volutions. Inner two volutions tightly and last one loosely coiled. Proloculus small,
spherical, its outside diameter 0.05 mm. Spirotheca thinand composed of a single structureless layer. Chomata low,asymmetrically developed in third volution.
Remarks.-This specimen resembles the type species but
the former is larger than the latter.
Occurrence.-One specimen from R-1 in Section E.
Genus Neofusulinel/a Deprat, 1912

Type species.-Neofusulinella praecursor Deprat, 1912

Neofusulinel/a giraudi Deprat, 1915
Figures 11-1-4
Neofusulinella giraudi Deprat, 1915, p.11, pI. 1, figs. 6-11; Ozawa,
1927, p. 38, figs. 3-6, 16 c, pI. 39, figs. 4-6.
Schubertella giraudi (Deprat). Sheng, 1963, p. 34, 158, 159, pI. 4,
figs. 1-9; Kobayashi, 1957, p. 263, 264, pI. 1, figs. 1-5; Kanmera, 1963, p. 88, 89, pI. 12, figs. 8-12; Hy. Igo, 1967, pI. 7,
figs. 7, 8; Sash ida, 1980, p. 300-302, pI. 34, figs. 7-9; Wang
ef al., 1981, p. 19, pI. 15, figs. 3, 4; Sun ef al., 1983, p. 17, pI. 3,
fig. 5; Wang and Tang, 1986, pI. 1, figs. 6, 8; Xiao ef al.,1986, p.73, pI. 1, figs. 3, 10, 15, 24, 25; Zhou, 1991, pI. 2, figs.
13,14.
Mesoschubertella giraudi (Deprat). Xia, 1994, pI. 18, fig. 6.
Materia/s.-Axial sections, TGUFU 1090,1092; slightly
oblique axial sections, TGUFU 1091,1093.
Remarks.-This species resembles Neofusulinella praecursor but is easily distinguished from the latter in that its shell
is not elongated. Sheng (1963) transferred this species to
the genus Schubertella based on the general shape, form
ratio, and development of chomata. I assign this species to
Neofusulinella for several reasons. The size of shell in
Schubertella is smaller than that of Neofusulinella The
early stage of Schubertella has sharply changed axis ofcoiling relative to later ones. Chomata of Neofusu/inella areasymmetrical and highly elongate while Schubertella has
asymmetrical massive chomata.
Occurrence.-TGUFU 1090 from R-1 in Section E; TGUFU
1091 from 8-47; TGUFU 1092,1093 from 8-48 in Section 0
near Higashi-Akuda.

Neofusulinella lantenoisi Deprat, 1913
Figures 11-5-8
Neofusulinella praecursor Deprat, 1913, p. 41, 42, pI. 7, figs. 2325; Thompson and Foster, 1937, p. 131, 132, pI. 24, figs. 1-4;
Pitakpaivan, 1965, p. 24-26, pI. 1, fig. 16; Leven, 1967, p. 126,

1023.

Permian fusulinaceans from Central Japan

pI. 1, figs. 9, 10; Toriyama et a/., 1969, p. 28-31, pI. 3, fig. 23,
pI. 4, figs. 1-15; Xiao et a/., 1986, p.75, pI. 1, figs. 11, 16.

Materia/s.-Slightly oblique axial section, TGUFU 1095;
axial sections, TGUFU 1096,1097,1098.
Description.-Shell small to medium in size and thickly
fusiform with straight axis of coiling and bluntly pointed poles.
Mature shell having six volutions, 1.50 to 1.67 mm in axial
length, 0.67 to 0.93 mm in width, and form ratio 1.8 to 2.2.
Proloculus small and spherical with outside diameter of
0.04 to 0.05 mm. Inner two volutions planispirally coiled andmake a right angle with axis of outer volutions. Spirothecathin, consists of a tectum, diaphanotheca, and lower thicker
layer. Septa thin, numerous, and plane. Chomata developed in third to last volutions, low, massive, and asymmetrical. Septal pore distinct in almost all volutions except
for the inner two volutions.
Remarks.-Recently, Li (1985) proposed a new genus
Multiavoella with the holotype M. guangxiensis from theLower Permian at Chongzuo, Guangxi, South China.
Multiavoella may be a synonym of Neofusulinella
Toriyama et a/. (1969) described this species from the
Khao Ph long Phrab section, Changwat Sara Buri, central
Thailand and discussed the taxonomic status, validity, andthe phylogeny of Neofusulinella
Recently, Zhou and Sheng (1993) restudied Permian staffellids and proposed a new concept of classification in thesesubfamilies. They attributed the genera Leella, Haoel/a,Sphaerulina (in part), and Caspiella (in part) to the genus
Neofusulinel/aOccurrence.-TGUFU 1095,1097,1098 from A-22 in Section A; TGUFU 1096 from A-21 in the same section.

629

Type species.-Minojapanel/aKanuma, 1953

e/ongata

Fujimoto and

Minojapanella (M.) elongata Fujimoto andKanuma, 1953
Figures 11-37,38
Minojapanel/a e/ongata Fujimoto and Kanuma, 1953, p. 152, pI. 19,
figs. 1-11; Thompson, 1954, pI. 2, figs. 1-5 (2-3,5; same as
pI. 19, figs. 1, 10, 11 of Fujimoto and Kanuma, 1953, p. 152) ;Morikawa and Isomi, 1961, p.7, pl. 2, figs. 10-15; Choi,1973,
p. 14, 15, pI. 1, figs. 8-11; Xiao et a/., 1986, p. 77, pI. 1, figs. 26,
29; Ozawa and Kobayashi, 1990, pI. 8, fig. 7; Xia, 1994, pI.
18, fig. 20.
Minojapanella (M.) e/ongata Fujimoto and Kanuma. Sheng, 1965,
p.565, pI. 1, figs. 1-6,10-15; Kahler and Kahler, 1980, p.191,
pI. 2, fig. 3; Ueno, 1992, p.1278, figs. 4-2-31.

Subfamily Biwaellinae Davydov, 1984
Genus Toriyamaia Kanmera, 1956

Materia/s.-Oblique section, TGUFU 1127; sagittal section,TGUFU 1128.
Remarks.- The present specimens coincide with the typespecimens which were described from a limestone near
Hachiman town, Gujyo County, Gifu Prefecture, Central
Japan. Fujimoto and Kanuma (1953) did not designate any
holotype in their paper, but Thompson (1954) subsequently
designated as the holotype the specimen illustrated by
Fujimoto and Kanuma in pI. 19, figs. 1a-c; this speCimen is infact the lectotype, not the holotype (Kahler and Kahler 1966,
p. 245). The actual locality of the type specimens was not
mentioned in the original paper. Recently, I examined thetype specimens; and confirmed that the lectotype occurredin a limestone exposed at Houshimaru, Wara village, Gifu
Prefecture.
Occurrence.-TGUFU 1127 from A-22; TGUFU 1128 from
A-21 in Section A.

Type species.- Toriyamaia /axiseptata Kanmera, 1956

Genus Dunbarula Ciry, 1948

Toriyamaia laxiseptata Kanmera, 1956
Figures 10-4, 5Toriyamaia /axiseptata Kanmera, 1956, p.252-255, pI. 36, figs. 114; Sheng, 1963, p. 33, 34, 157, pI. 4, figs. 36, 37; Kanmera,
1963, p.87,88, pI. 11, figs. 1-4, pI. 19, figs. 8, 9; Kanmera andMikami, 1965, p.277-279, pI. 46, figs. 9, 10; Choi, 1973, p.
20, pI. 2, figs. 8-10; Ozawa, 1975, pI. 7, fig. 18; Lin et a/.,1977, p. 19, pI. 3, fig. 17; Kahler and Kahler, 1979, p.226,
227, pI. 4, fig.7; Zhou et a/., 1987, pI. 2, fig. 7; Ueno, 1992,
p.1276-1278, figs. 4-1-7.

Materials.-Tangential sections, TGUFU 1073,1074.
Remarks.-The present specimens closely resemble Kanmera's original ones described from the Kozaki Formation,
Kyushu, Japan.
Occurrence.-AII specimens from B-48 in Section D.
Family Boultoniidae Skinner and Wilde, 1954
Genus Minojapanella Fujimoto and Kanuma, 1953
Subgenus Minojapanella Fujimoto and Kanuma, 1953

Type species.-Dunbarula mathieui Ciry, 1948

Dunbarula uenoi Hh. Igo sp. nov.
Figures 11-35, 36

Materia/s.-Axial section of holotype, TGUFU 1125; axial
section of paratype, TGUFU 1126.
Diagnosis.-Elongate Dunbarula with a thin wall, and septa
only fluted in polar regions.
Description.-Shell small, elongate cylindrical in shape andhaving a gently bent axis of coiling and broadly roundedpolar ends. Juvenile volutions tightly coiled but shell loosely
coiled in last volution. Inner first to second volutions at alarge angle of coiling to the outer ones. Shell of holotype
consists of 5 volutions, 2.13 mm in axial length, 0.43 mm inwidth, with a form ratio of 4.95. Proloculus small andspherical, its outside diameter 0.02 mm in holotype. Radius
vectors of first to fifth volutions in holotype, 0.04,0.05,0.07,
0.12, and 0.20 mm, respectively.
Spirotheca thin and composed of a tectum and less dense
layer. Thickness of spirotheca of first to fifth volutions ofHisaharu Igo

630

holotype 0.010, 0.012, 0.025, 0.027, and 0.020 mm, respectively. Septa numerous and strongly fluted in polar regions.
Septal pores present near polar regions.
Chomata small and developed in second to fourth volutions.
Remarks.-Skinner and Wilde (1967) redescribed topotypesof Ounbarula mathieui Ciry from Tunisia. Ounbarula mathieui
has broad variations in shape, ranging from ellipsoidal
through thickly to slender subcylindrical. The present new
species has a characteristic elongate shell. Other biocharacters show that the species is assignable to the genus
Ounbarula.
Etymology.- This species name is dedicated to Dr. Katsumi Ueno, Assistant Professor of the Institute of Geoscience, University of Tsukuba, for his active research on
fusulinaceans.
Occurrence.-AII specimens from A-22 in Section A.
Family Schwagerinidae Dunbar and Henbest, 1930
Subfamily Schwagerininae Dunbar and Henbest, 1930
Genus Schwagerina Moller, 1877

keriotheca. Proloculus small, spherical, with an outside
diameter of about 0.02 mm. Septa numerous, highly andregularly fluted throughout shell. Septal folds regular inshape with broadly rounded crest.
Chomata lacking except for poor development on thesurface of proloculus. Tunnel single, low, and narrow.
Remarks.- The present new species is characterized byan inflated fusiform shell and regularly fluted septa.
Hy. Igo (1964) proposed Schwagerina higashidaniensis fromthe Shiroi Formation of the Nyukawa Group. He illustrated
one axial and one tangential sections and two sagittal
sections. The specimen questionably identified by him (pI.
104, fig. 2) probably falls within the present new species.
Etymology.- This species name is dedicated to Dr. Shuko
Adachi, Assistant Professor of the Institute of Geoscience,
University of Tsukuba, for her active research on Carboniferous to Triassic foraminifers.
Occurrence.-Holotype TGUFU 1060 and tangential section TGUFU 1059 from R-1 in Section E; paratype TGUFU
1061 from B-2 in Section B near Akuda-Kuchi.
Schwagerina muraii Morikawa, 1955

Type species.-Bore/is princeps Ehrenberg, 1842

Figure 5-4
Schwagerina muraii Morikawa, 1955, p. 104, 105, pI. 14, figs. 1-4,
13; Hy. Igo, 1965, pI. 29, fig. 1.

Schwagerina shukoae Hh. Igo sp. nov.
Figures 9-5-7

Materials.- Tangential section of immature specimen,
TGUFU 1059; axial section of holotype, TGUFU 1060;
slightly tangential axial section of paratype, TGUFU 1061.
Oiagnosis.-Small Schwagerina with an inflated fusiform
shell, small proloculus, regularly fluted septa, and without
axial fillings.
Oescription.-Shell medium in size, tightly coiled, fusiformwith an inflated median part and broadly rounded poles. In
holotype, axial length 6.98 mm, median width 2.90 mm, andform ratio 2.39. Mature shells consist of 6 volutions.
Spirotheca thin, consists of a tectum and fine alveolar

Table 2.

1
2

I

Materia/.-Axial section of mature specimen, TGUFU 1003.
Oescription.-Shell large, elongate fusiform with bluntly
rounded poles. Equatorial part of shell somewhat constricted. This specimen has 6 volutions, and is 9.06 mm in axial
length and 2.73 mm in median width, with a form ratio of
3.32. Radius vectors in first to sixth volutions 0.03, 0.48,
0.67,0.78,1.17, and 1.47 mm, respectively.
Spirotheca thick, consists of a thin tectum and coarse
keriotheca. Thickness of spirotheca in sixth whorl 0.13 mm.
Proloculus large, spherical, its outside diameter is 0.42 mm.
Septa strongly and regularly fluted throughout shell. Heavy
secondary deposition fills in tunnel area and polar regions

Measurements of Schwagerina shukoae Hh Igo sp nov (in mm)

Reg. no.

Fig.

L.

w.

F.R.

D.P.

TGUFU1060
TGUFU1061

9-6
9-7

6.93
9.19

2.90
3.46

2.39
2.66

0.20
0.27?

Radius vector
1

2

3

4

5

6

0.20
0.23

0.33
0.37

0.53
0.83

0.75
1.00

1.00
1.43

1.27
1.83

Thickness of spirotheca
1
2

1

2

3

4

5

6

0.03
0.05

0.04
0.05

0.06
0.05

0.06
0.08

0.07
0.10

0.07
0.10

Figure 5. 1-3. Pseudofusulina norikurensis Hy. Igo, 1 : axial section, TGUFU 1000, 2: slightly oblique axial section
TGUFU 1001, 3: axial section of immature specimen, TGUFU 1002. 4. Schwagerina muraii Morikawa, axial section, TGUFU~003. 5-8. P~eudofusulina paratschernyschewi Kanuma, 5-7: axial sections, TGUFU 1004, 1005, 1006, 8: axial section ofI~mature speCImen, TGUFU 1007. 9-11. Pseudofusulina horadaniensis Hy. Igo, axial sections, TGUFU 1008, 1009, 1010. All
fIgures x 10.

1023.

Permian fusulinaceans from Central Japan

63 1

Hisaharu 19o

632'

except for outermost whorl. Tunnel low and narrow.
Chomata almost absent.
Remarks.-This species resembles Pseudofusulina fusiformis (Schellwien) in many respects, but the former differs fromthe latter in having heavy secondary deposition.
This species was described from the lower Sote Formationof the Nyukawa Group by Hy. 19o (1965). The present
specimen is very similar to 19o's Sote specimen. This
species also resembles Pseudofusulina kraffti (Schellwien),but the former is distinguishable from the latter in its elongate
shell, pointed polar ends, and thin spirotheca.
Occurrence.-From 8-44 in Section D.
Schwagerina kwangchiensis Chen, 1956

Figures 8-7-9
Schwagerina kwangchiensis Chen, 1956, p. 23, 24, pI. 1, figs. 17,
18; Sheng, 1963, p. 61, 62, 187, 188, pI. 14, figs. 19-21.
Materia/s.-Slightly oblique axial sections, TGUFU 1043,
1045; axial section, TGUFU 1044.
Description.-Shell medium in size, elongate fusiform tosubcylindrical, more or less tightly coiled and having broadly
rounded poles. Mature shell with seven volutions, 7.66 mm
in length, 2.06 mm in median width, with a form ratio of 3.72.
Proloculus very small, spherical, 0.12 to 022 mm in outside
diameter. Spirotheca thin, consists of a tectum and keriotheca with fine alveoli. Septa numerous, strongly and regularly fluted throughout shell. Septal folds regularly shaped
and closely spaced.
Axial filling very weak in polar regions of younger volutions.
Chomata almost lacking and tunnel indistinct.
Remarks.-The present species resembles Pseudofusulina
regu/aris (Schellwien), but the former has a smaller
proloculus. It also resembles Pseudofusulina ambigua (Deprat), but is distinguished from the latter in larger size of shelland proloculus.
The present species was originally described from the
Wuhsueh Limestone in Hupeh, China, by Chen (1956). The
limestone equivalent with the Maokou Limestone is exposedin southwestern China.
Occurrence.-AII specimens from R-3 in Section E.
Schwagerina higashidaniensis Hy. 19o, 1964

Figures 8-16, 17
Schwagerina higashidaniensis Hy. Igo, 1964, p. 644, 645, pI. 104,
figs. 1-4.
Materia/s.-Axial sections, TGUFU 1052,1053.
Description.-Shell medium-sized, elongate fusiform tosubcylindrical with broadly rounded poles and straight axis ofcoiling. Mature shell has five to six volutions and 6.32 to
8.00 mm in axial length, 1.53 to 1.83 mm in median width, witha form ratio of 2.27 to 3.45. Inner second to third volutions
tightly coiled with sharply pointed poles. Outer fourth to fifth
volutions increase in height rather rapidly. Proloculus small,
spherical, its outside diameter being 0.16 to 0.17 mm. Spirotheca thin, consists of a tectum and keriotheca with very finealveoli. Septa numerous, thin, strongly and irregularly fluted.
Phrenotheca absent in present specimens. Tunnel singular,
low, and indistinct in outer volutions. Chomata present infirst to second volutions but lacking outer volutions.
Remarks.-The present specimens coincide with 19o's
holotype described from the Siroi Formation, Nyukawa
Group. The present species is easily distinguishable fromsome species of the genus Paraschwagerina The former
has irregular septal fluting, but the latter has regular septal
fluting. The former has extraordinarily thin spirotheca andsepta.
Occurrence.-AII specimens from R-1 in Section E.
Schwagerina kanumai Hh. 19o sp. nov.

Figures 9-1-4
Materia/s.-Axial section of holotype, TGUFU 1055; axial
section of paratypes, TGUFU 1056,1057,1058.
Diagnosis.-Small Schwagerina with an inflated fusiform
shell, small proloculus, and thin spirotheca. Septa regularly
fluted. Axial filling lacking.
Description.-Shell medium in size, inflated fusiform, andbiconical in shape. It has more or less acutely pointed
poles and a straight axis of coiling. Mature shell consists of
6 volutions. Axial length 7.86 mm, median width 3.33 mm,
and form ratio 2.36 in holotype.
Inner three volutions tightly coiled and outer three volutions increasing rather rapidly in height. Proloculus small,
spherical, its outside diameter 0.23 mm in holotype.
Spirotheca thin and composed of a tectum and finealveolar keriotheca. Thickness of spirotheca of each volution in holotype 0.03, 0.03, 0.03, 0.05, 0.06, and 0.08 mm,
respectively. Septa very thin, regularly and intensely fluted.
Septal folds across almost entire chamber in outer volutions.
Phrenotheca weak and rarely observed. Chomata almost
lacking except for rudimentary ones on the surface of
proloculus and first volution.
Remarks.-The present new species is quite similar to
Schwagerina ? annamitica (Deprat), which was described fromthe Shiroi Formation, Nyukawa Group by Hy. 19o (1964). The
former is distinguished from the latter in highly fluted septa.
Etym%gy.- This species name is dedicated to Dr.
Mosaburo Kanuma, Professor Emeritus of the Department of
Astronomy and Earth Sciences, Tokyo Gakugei University.
He contributed to early studies of geology and fusulinaceansin the Hachiman area.
Occurrence.-AII specimens from R-1 in Section E.

Figure 6. 1-7. Pseudofusulina isomie Hy.lgo, 1,3-5,7: axial sections, TGUFU 1011, 1013, 1014, 1015, 1017, 2: sagittal
section, TGUFU 1012, 6: axial section of immature specimen, TGUFU 1016. 8, 9. Pseudofusulina duplithecata Hy. Igo, 8:
axial section, TGUFU 1018, 9: slightly oblique axial section, TGUFU 1019. 10. Pseudofusulina kraffti (Schellwien), axial
section, TGUFU 1020. All figures x10.

1023.

Permian fusulinaceans from Central Japan

633

634

Hisaharu Igo
Table 3

1
2
3
4

1
2
3
4

Measurements of Schwagerina kanumai Hh. Igo sp. nov. (in mm)

Reg. no.

Fig.

L.

W.

F.R.

D.P.

TGUFU1055
TGUFU1056
TGUFU1057
TGUFU1058

9-1
9-2
9-3
9-4

7.83
4.81
4.40
3.92

3.33
1.93
1.83
2.66

2.36
2.50
2.40
1.47

0.23
0.27
0.18
0.27

Thickness of spirotheca
2
3
4
5
0.03
0.03
0.05
0.06
0.03
0.04
0.03
0.04
0.02
0.03
0.05
0.05
...
0.02
0.02
0.04

0.08

1
0.03
0.03
0.01
0.02

1
0.19
0.20
0.17
0.20

2
0.30
0.31
0.24
0.30

Radius vector
4
3
0.46
1.80
0.47
0.73
0.37
0.57
0.47
0.70

5
1.23
1.71
0.88

6
1.68

._-

.. -

...
...

6
...

...
. ..

Genus Parafusulina Dunbar and Skinner, 1931

Type species.-Parafusu/ina wordensis Dunbar and Skinner, 1931

The holotype is also a broken specimen.
Occurrence.-AII specimens from 8-17 in Section D near
Higashi-Akuda.
Parafusulina sotensis Hy. Igo, 1967

Parafusulina? sublineata Hy. Igo, 1965

Figure 10-6

Figures 9-8-15

Parafusulina kawaii Morikawa sotensis Hy. Igo, 1967, p.12, pI. 8,

Parafusulina sublineata Hy. Igo, 1965, p. 221, 222, pI. 30, figs. 1-3,

pI. 32, figs. 1,2.

Materia/s.-Axial sections, TGUFU 1063,1065,1066,1068;
slightly oblique axial sections, TGUFU 1062,1064,1069; tangential section, TGUFU 1067.
Description.-Shell large, highly elongate subcylindrical,
having a slightly straight median part, irregularly undulated
lateral slope, and truncated poles. Axis of coiling slightly
bent. Mature shell consists of eight volutions. Axial length
7.33 to 12.00 mm, median width 1.64 to 3.2 mm, and form
ratio 3.75 to 4.39.
First to third volutions tightly coiled, outer ones rapidly
increase in height. Proloculus small, its outside diameter is
0.17 to 0.23 mm. Spirotheca thin, consists of a thin tectumand keriotheca but spirotheca of inner volutions recrystallized. Septa numerous, highly fluted throughout shell.
Septal folds form rounded crests, commonly extend acrossentire chamber. Phrenotheca almost absent. Vestigial
chomata present in first to second volutions but lacking inouter ones. Axial filling almost lacking.
Remarks.- The present specimens agree closely with Hy.
Igo's holotype speCimen described from the Sote Formation
Nyukawa Group.
'
The Akuda specimens are larger than those of the Sote
specimens. I could not obtain any complete specimens.

figs. 1-6.
Materia/.-Slightly oblique axial section, TGUFU 1075.
Description.-Shell large, elongate fusiform with more orless inflated median part and bluntly rounded poles. Present specimen consists of five volutions. Axial length 9.06
mm, median width 2.93 mm, and form ratio 3.09. Shelltightly coiled with a straight axis of coiling. Proloculus
spherical with an outside diameter of 0.31 mm.
Spirotheca recrystallized, but probably composed of atectum and keriotheca. Septa regularly fluted and septal
loops with bluntly rounded to angular crests. Chomata
lacking. Tunnel single narrow.
Axial filling weakly developed in polar regions of younger
volutions.
Remarks.- The present specimen is weekly recrystallized,
so that I could not determine detailed spirothecal structure.
The Akuda specimen has less dense axial filling than that ofHy. Igo's holotype from the Sote Formation, Nyukawa Group.
Occurrence.-From 8-11 in Section D.
Genus Pseudofusulina Dunbar and Skinner, 1931

Type species.-Pseudofusulina huecoensis Dunber andSkinner, 1931

.Figure 7. 1 9. Pseudofusulina min~ensis H~. Igo sp. nov., 1, 3: axial sections of paratypes, TGUFU 1021, 1023, 2: axial
section of holotype, TGUFU 1022, 4: sagltta~ sectl.on, TGU.FU 1024, 5-9: axial sections of immature specimens, TGUFU 1025,
10~6, 102":, 1028, 1029. 10,11. Pseudofusulma mtyamensls .Hh. Igo sp. nov., 10: axial section of paratype, TGUFU 1030, 11 :axial section of hol.otype, TGUFU 1031. 12-16. Pseudofusulma sp., 13,14: axial sections of mature specimens TGUFU 1033
1034,12,15,16: axial sections of immature specimens, TGUFU 1032, 1035, 1036. All figures X10.
',

1023.

Permian fusulinaceans from Central Japan

635

~
~
.

16

Hisaharu Igo

636

Pseudofusulina norikurensis Hy. Igo, 1959

Figures 5-1-3
Pseudofusulina kraffti (Schellwien) norikurensis Hy. Igo, 1959, p.

244,245, pI. 2, figs. 1-3; Sashida, 1980, p.296-298, pI. 34,
figs. 3, 4.
Pseudofusulina norikurensis Hy. Igo. Morikawa and Isomi, 1961, p.
21, pI. 15, figs. 1-4, pI. 16, figs. 1-7.
Materia/s.-Axial sections, TGUFU 1000,1002; slightly
oblique axial section, TGUFU 1001.
Description.-Shell large, inflated fusiform with broadly
rounded poles. Mature shell consists of 8 to 9 volutions.
Axial length 8.79 to 10.32 mm, median width 4.33 to 5.13 mm,
and form ratio 2.01 to 2.03. Radius vectors in first to eighth
volutions 0040,0.63,0.83,1.17,1.53,1.90,2.30, and 2.76 mm,
respectively.
Proloculus large, spherical with outside diameter of 0.50 to
0.57 mm. Spirotheca thick, consists of a thin distinct
tectum and very coarse keriotheca. Upper keriotheca thinand 1/4 of whole thickness of keriotheca. Septa numerous,
relatively intensely and regularly fluted along axial regions.
Cuniculi well developed in outer volutions. Phrenotheca
poorly developed in lower part of chamber. Axial filling welldeveloped in polar regions of inner five volutions. Tunnel low
and narrow. Chomata lacking throughout shell.
Remarks.- This species resembles Parafusulina japonica(Gumbel), but differs from the latter in having an inflated shellwith broadly rounded poles. This species was originallyproposed by Hy. Igo (1959) as a subspecies of Pseudofusulina
kraffti, but Morikawa and Isomi (1961) raised it to species
level. I consider that this species is closely related toParafusulina japonica in septal fluting and axial filling.
Occurrence.- TGUFU 1000,1001 from A-1 south of the
Horikoshi Pass; TGUFU 1002 from 8-47 in Section D.

Pseudofusulina paratschernyschewi Kanuma, 1959

Figures 5-5-8
Pseudofusulina paratschernyschewi Kanuma, 1959, p. 67, 68, p. 6,

figs. 4-7.
Materia/s.-Axial sections, TGUFU 1004,1005,1006,1007.
Description.-Shell large and fusiform with almost straightaxis of coiling and narrowly rounded poles. It is rather
loosely coiled. Mature shell of 5 to 6 volutions 8.33 mm inaxial length, and 3.40 mm in median width, giving a form ratioof 2.48. Radius vectors in first to sixth volutions, 0047,0.67,
0.93, 1.23, and 1.60 mm, respectively.
Proloculus relatively large, spherical to subspherical, withan outside diameter of 0.58 mm in mature shell. Spirotheca
thick, consists of a thin tectum and coarse keriotheca.

Average thickness of spirotheca in first to last volutions inmature shell 0.05, 0.05, 0.10, 0.11, and 0.15 mm, respectively.
Septa numerous, rather regularly fluted nearly from pole topole. Septal folds narrow, generally high, and commonly
attain top of volutions. Cuniculi developed in outer volutions. Tunnel low and relatively narrow. Chomata almost
lacking. Axial filling poorly developed in polar region of first
to fourth volutions. Phrenotheca not present.
Remarks.- This species was first described by Kanuma(1959). He mentioned that this species accords closely with
Parafusulina japonica (Gumbel) described by Chen (1934).
He considered that Chen's P. japonica is related to
Pseudofusulina tschernyschewi (Schellwien).
This species resembles P. tschernyschewi, but the former
has a larger proloculus. This species also resembles Parafusulina japonica, but differs from the latter in having dense
axial fillings and thick spirotheca.
Occurrence.- TGUFU 1004 from 8-42; TGUFU 1005,1006
from 8- 47; TGUFU 1007 from A-1 south of the Horikoshi
Pass.
Pseudofusulina horadaniensis Hy. Igo, 1964

Figures 5-9-11
Pseudofusulina horadaniensis Hy. Igo, 1964, p. 647, pI. 104, figs.

7-9.
Materia/s.-Axial sections, TGUFU 1008,1009,1010.
Description.-Shell large, cylindrical to elongate fusiform,and having straight median portion and truncated poles.
Outline of shell elongate subhexagonal in axial section.
Axial length 7.13 to 8.59 mm, median width 2.73 to 3.33 mm,
and form ratio 2.10 to 2.58 in mature shells. Mature shellconsists of six to seven volutions.
Radius vectors in first to seventh volution 0.33, 0.48, 0.66,
0.87,1.13,1047, and 1.80 mm, respectively. Shell tightly
coiled in first to third volutions.
Proloculus relatively large, spherical, with outside diameterof 0040 to 0.43 mm. Spirotheca thick, consists of a thintectum and coarse keriotheca with rather coarse alveoli.
Thickness of spirotheca in first to sixth volutions 0.03, 0.05,
0.06,0.08,0.11, and 0.13 mm, respectively.
Septa numerous and intensely fluted. Septal folds about
1/2 as high as chamber. Phrenotheca absent. Tunnel
single, low, and narrow. Chomata lacking. Axial fillingstrong and massive in polar regions. Outer one or twovolutions lack axial filling.
Remarks.-This species closely resembles Pseudofusulina
kraffti but differs from the latter in having pointed truncate
poles and massive axial fillings. This species resembles inmany respects Parafusulina postkraffti Leven (1967) described

Figure 8. 1, 2. Pseudofusulina fusiformis (Schellwien), axial sections, TGUFU 1037, 1038. 3-6. Pseudofusulina regu/aris (Schellwien), 3,5: axial sections, TGUFU 1039,1041, 4,6: slightly oblique axial sections, TGUFU 1040,1042. 7-9.
Schwagerina kwangchiensis Chen, 7, 9: slightly oblique axial sections, TGUFU 1043, 1045, 8: axial section, TGUFU 1044.
10-13. Pseudofusulina houziguanica Sheng, axial sections, TGUFU 1046, 1047, 1048, 1049. 14, 15. Pseudofusulina krotowi(Schellwien), axial sections, TGUFU 1050,1051. 16,17. Schwagerina higashidaniensis Hy. Igo, axial sections, TGUFU 1052,
1053. 18. Chalaroschwagerina aff. chinensis (Chen), axial section, TGUFU 1054. All figures x 10.

1023. Permian fusulinaceans from Central Japan

637

Hisaharu Igo

638

from the Pamirs but the latter has a large shell, intensely
fluted septa, and well developed phrenotheca.
Occurrence.- TGUFU 1008 from 8-48; TGUFU 1010 from
8-44 in Section D; TGUFU 1009 from A-2 south of the
Horikoshi Pass.

Pseudofusulina isomie Hy. Igo, 1965
Figures 6-1-7
Pseudofusulina isomie Hy. Igo, 1965, p. 219, 220, pI. 29, fig. 6, pI.

30, figs. 5, 6, pI. 31, figs. 6, 7.

Materia/s.-Axial section of mature specimens, TGUFU
1011,1013,1014,1015,1017; axial section of immature specimen, TGUFU 1016; sagittal section of mature specimen,
TGUFU 1012.
Description.-Shell large, cylindrical to elongate fusiform,and having slightly concave median part and bluntly roundedpoles. Axial length 8.91 to 11.79 mm, median width 3.40 to
4.33 mm, and form ratio 2.45 to 3.00 in mature specimens.
Mature shell has six to seven volutions.
Radius vectors in first to seventh volutions in largest
specimen (TGUFU 1011) 0.33, 0.47, 0.73, 1.00, 1.37, and 1.73mm, respectively. Shell rather tightly coiled in first to third
volutions and having a straight axis of coiling. Proloculus
large, spherical with an outside diameter of 0.37 to 0.73 mm.
Spirotheca thick, consists of a tectum and relatively
coarse alveolar keriotheca. Upper keriothecal layer thin,
1/4 of whole thickness of keriotheca. Septa numerous,
intensely fluted throughout length of shell and commonly
extend across entire chamber. Fluting regular in medianpart of shell. Phrenotheca almost absent. Chomata lacking, tunnel low and narrow. Axial filling weekly developed inaxial region of inner volutions.
Remarks.-In the Akuda specimens, axial filling is weaker
than in the holotype from the Sote Formation. However,shape of shell and septal fluting are exactly the same inthese specimens.
The Akuda specimens resemble Parafusulina japonica(Gumbel), which was illustrated by Toriyama (1975, pI. 5, figs.
48; pI. 6, figs. 1-8) but the latter specimens have an elongate
fusiform shell and intense septal fluting.
This species closely resembles Schwagerina toyamaensis
Suyari (1962). It differs from the latter in smaller proloculus,
intense septal fluting, and thick spirothecae.
Occurrence.- TGUFU 1011 from 8-37; TGUFU 1013,1014,
1016 from 8-42; TGUFU 1012,1015,1017 from 8-47 in Section D.

Pseudofusulina duplithecata Hy. Igo, 1956
Figures 6-8, 9
Pseudofusulina dupJithecata Hy. Igo, 1956, p. 297-299, pI. 18, figs.

1-10, pI. 19, figs. 1-3, 6; Hy. Igo, 1965, pI. 32, fig. 3.

Materia/s.-Axial section, TGUFU 1018; slightly oblique
axial section, TGUFU 1019.
Description.-Shell large, cylindrical to elongate fusiformwith an almost straight axis of coiling and having slightly
straight median part and bluntly rounded poles. Matureshell has six to seven volutions.
Axial length 9.86 to 11.00 mm and median width 3.36 to
4.06 mm, giving a form ratio of 2.71 to 2.93. Shell tightly
coiled throughout. Radius vectors in first to seventh volutions of largest specimen 0.28, 0.50, 0.70, 0.93, 1.23, 1.63, and2.00 mm, respectively. Proloculus large, spherical, withoutside diameter of 0.40 to 0.33 mm.
Spirotheca thick, consists of a tectum and upper andlower keriothecae. Discontinuous light layer observed inmedian part of the keriotheca. Outer surface of tectum iscorrugated in fifth to sixth volutions of specimen illustrated inFig. 6-9. Septa numerous, weakly fluted in polar regions.
Septal loops low and 1/3 as high as chambers in outer
volutions. Cuniculi developed in outer volutions. Chomata
poorly developed in first to second volutions. Tunnel single,
low, and wide. Axial fillings weekly developed in polar
regions.
Remarks.- This species closely resembles Pseudofusu/ina
isomie in its shell shape, but is distinguishable from the latterin weak and regular septal fluting. This species is characterized by possessing double spirotheca: the inner andouter keriothecae (Hy. Igo, 1956).
This double spirothecae appears in some specimens but isabsent in others.
Occurrence.-AII specimens from 8-47 in Section D.

Pseudofusulina kraffti (Schellwien, 1909)
Figure 6-10
Fusulina kraffti Schellwien, 1909, p.169, pI. 16, figs. 1-9.
Schellwienia kraffti (Schellwien). Ozawa, 1925, p.25-27, pI. 6,

fig. 7, pI. 7, fig. 4; Ozawa, 1927, p.147, fig. 5.
Fujimoto, 1936, p. 80, 81, pI.
14, figs. 3-8; Morikawa, 1955, p. 94, 95, pI. 7, figs. 15-17;
Kanuma, 1959, p. 70, 71, pI. 7, figs. 1-3, 6; Morikawa andIsomi, 1961, p. 20, 21, pI. 14, figs. 1-10; pI. 15, figs. 5, 6;
Sheng, 1963, p. 65, pI. 16, figs. 16-18; Chang, 1963, p. 206, pI.
4, fig.11; Kalmykova, 1967, p. 183, 184, pI. 10, figs. 1-4;
Leven, 1967, p. 146, 147, pI. 10, figs. 5, 6; Choi, 1970, p. 371,
372, pI. 1, figs. 6, 7; Choi, 1973, p.52, pI. 6, figs. 1-5; Sashida, 1980, p. 296-298, pI. 34, figs. 3, 4; Xia, 1994, pI. 15, fig.
5.

Pseudofusulina kraffti (Schellwien).

Figure 9. 1-4. Schwagerina kanumai Hh. Igo sp. nov., 1: axial section of holotype, TGUFU 1055, 2-4: axial sections ofparatypes, TGUFU 1056,1057,1058. 5-7. Schwagerina shukoae Hh.lgo sp. nov., 5: tangential section of immature specimen, TGUFU 1059, 6: axial section of holotype, TGUFU 1060, 7: slightly tangential section of paratype, TGUFU 1061. 8-15.
Parafusulina ? sublineata Hy. Igo, 8,10,15: slightly oblique axial sections, TGUFU 1062, 1064, 1069, 9, 11, 12, 14: axial sections,TGUFU 1063,1065,1066,1068,13: tangential section, TGUFU 1067. All figures X10.

1023. Permian fusulinaceans from Central Japan

~
2

1

14

639

Hisaharu Igo

640
Pseudofusulina kraffti kraffti (Schellwien).

Kanmera and Mikami,
1965, p. 299-301, pI. 44, fig. 7.
Pseudofusulina kraffti (Schellwiei1) forma A. Nogami, 1961, p.
217-219, pI. 10, figs. 5-8.

Material.-Axial section, TGUFU 1020.
Oescription.-Shell large, cylindrical to subcylindrical withbroadly rounded poles. Mature shell has six to sevenvolutions. Axial length 10.66 mm and median width 4.33mm, giving form ratio of 2.46. Shell tightly coiled in the firstto third volutions and having a weekly bent axis of coiling.
Radius vectors in first to sixth volutions 0.27, 0.38, 0.58, 0.90,
1.33? and 1.73 mm, respectively.
Proloculus large, spherical, with outside diameter of 0.37
mm. Spirotheca thick, consists of a tectum and coarse
alveolar keriotheca. Septa numerous, intensely fluted nearaxis of coiling, most septal loops reach ceiling of chambers.
Chomata poorly developed in first to second volutions.
Tunnel single, low, and wide. Axial fillings developed inpolar regions.
Remarks.-In the Akuda specimen, the spirotheca varies inthickness even in the same volution. This well known

Table 4.
Reg. no.

2
3
4
5
6
7
8
9

TGUFU1021
TGUFU1022
TGUFU1023
TGUFU1024TGUFU1025TGUFU1026TGUFU1027TGUFU1028
TGUFU1029

1
2
3
4
5
6
7
8
9

1
0.06
0.03
0.03
0.03
0.03
0.03
0.03
0.06
0.02

~

species has broad variations in shell shape, septal fluting,size of shell, degree of axial filling, and other features.
Many previous authors have already pointed out thesevariations.
Occurrence.-From OHT- 4 at Otohime-Tani.

Pseudofusulina minoensis Hh. Igo sp. nov.
Figures 7-1-9

Materia/s.-Axial section of holotype, TGUFU 1022; axial
sections of paratypes, TGUFU 1021,1023; axial sections ofimmature specimens, TGUFU 1025,1026,1027,1028,1029;
sagittal section of mature specimen, TGUFU 1024.
Oiagnosis.-Large Pseudofusu/ina with an elongate cylindrical shell and large proloculus. Septa regularly fluted.
Oescription.-Shell large, elongate subcylindrical to elongate fusiform, with a slightly flat median part and bluntly
rounded poles. Axial length 11.32 to 10.52 mm, medianwidth 3.33 to 2.73 mm, and form ratio 4.00 to 3.85 in mature
specimens.
Shell rather tightly coiled in first to third volutions andMeasurements of Pseudofusulina minoensis Hh Igo sp nov (in mm)
Radius vector
Fig.
D.P.
L.
W.
F.R.
4
1
2
3
7-1
11.32
3.33
3.40
0.49
0.35
0.50
0.75
1.25
7-2
11.52
2.73
4.22
0.38
0.30
0.47
0.63
0.93
7-3
10.52
2.86
3.68
0.53
0.37
0.53
0.77
1.03
...
...
7-4
3.40
0.43
0.37
0.50
0.73
1.03
7-5
5.73
1.86
3.03
0.43
0.33
0.90
0.50
0.70
7-6
7.79
2.34
3.33
0.37
0.35
0.50
0.73
1.03
7-7
7.33
2.40
3.05
0.49
0.33
1.00
0.50
0.70
7-8
6.79
2.20
3.09
0.42
0.37
0.53
0.73
0.90
7-9
5.40
1.50
3.60
0.26
0.18
0.42
0.60
0.28

Thickness of spirotheca
2
3
4
0.05
0.06
0.06
0.03
0.04
0.08
0.04
0.06
0.09
0.03
0.02
0.07
0.04
0.05
0.06
0.05
0.09
0.03
0.04
0.05
0.07
0.04
0.06
0.08
0.03
0.05
0.04

5
1.45
1.32
1.37
1.40
...

1.33
1.30
1.17
...

5
0.08
0.09
0.08
0.08
...

0.11
0.08
0.08
...

, Figur~ 10. 1 3. Acervoschwagerina fuji:notoi, Kan~ma, 1,2: oblique sections, TGUFU 1070,1071,3: slightly oblique
axial sec~lon, TGU~U 1072, x~O. 4,5. Tonyama/a lax/septata Kanmera, tangential sections TGUFU 1073 1074 x15 6
Par?fusultn~ sotens~s Hy. Igo, slightly oblique axial sect.ion, TGUFU 1075, x 10. 7-10. Maklaya' pamirica (Lev~n), 7, '9: sli' htl .
obliqueaxiald T
sections,
TGUFU
1076,1078
11 12. Maklaya sarab unensls. g .yK.
.
.
' 8, 10: axial sections, TGUFU 1077, 1079 x17. ,anmera an
onyama,. axl~1 sections, TGUFU 1080,1081, X17. 13,14. Armenina sp., axial sections, TGUFU 1082 1083
x17. 15-20. Pseudorelcheltna sp., 15,16,18,20: tangential sections TGUFU 1084 1085 1087 1089 17' axial s~cf 'TGUFU 1086,19: sagittal section, TGUFU 1088.15-19= X 30, 20= X 50.'
,,',.
Ion,

1023.

Permian fusulinaceans from Central Japan641

Hisaharu Igo

642

having a straight or slightly curved axis of coiling.
Proloculus large, spherical, with an outside diameter of 0.53
to 0.26 mm.
Spirotheca thick, consists of a tectum and coarse alveolar
keriotheca. Thickness of spirotheca in first to fifth volutionsof ho\otype 0.03, 0.03, 0.04, O.oS, and 0.09 mm, respectively.
Surface of tectum irregularly undulated.
Septa numerous, more or less regularly fluted, and most
septal loops reach up to about 2/3 height of chamber.
Phrenotheca present in outer volutions. Chomata lacking
except for very poor development in first to second volutions.
Tunnel singular, low, and narrow. Axial filling weekly developed in inner volutions.
Remarks.- This species very closely resembles
Pseudofusulina granumavenae (Roemer) in many points.
The former can be distinguished from the latter, however, inhaving fewer volutions and weak septal fluting.
Etymology.- This species name is derived from theancient province name of Gifu Prefecture.
Occurrence.-Collected from R-1 in Section E of NishiAkuda.

rounded poles and a straight axis of coiling. Axial length
5.66 and 5.S6 mm, median width 1.53 and 1.57 mm, form ratio
3.83 and 3.61. Shell tightly coiled in first to third volutions.
Radius vectors in first to fifth volutions of holotype 0.27, 0.40,
0.53, 0.73, and 0.S7 mm, respectively. Proloculus large
compared with shell size. Spirotheca thin, consists of atectum and fine alveolar keriotheca. Thickness of spirotheca in first to fifth volutions of holotype 0.02, 0.03, 0,04,
0.06, and 0.05 mm, respectively. Septa regularly fluted,
most loops reach to ceiling of chamber. Chomata lacking,but rudimentary ones observed on first volution. Tunnel
singular and low. Axial filling very weekly developed in
inner volutions.
Remarks.- This species resembles Pseudofusulina
minoensis Hh. Igo, but the former has a smaller shell than thelatter.
Etymology.- This species name is derived from the nameof a village in the studied area.
Occurrence.-From R-1 in Section E of Nishi-Akuda.

Pseudofusulina fusiformis (Schellwien, 1909)
Figures 8-1, 2

Pseudofusulina miyamensis Hh. Igo sp. nov.

Materials.-Axial section of holotype, TGUFU 1031; axial

section of paratype, TGUFU 1030.
Diagnosis.-Small Pseudofusulina with an elongate fusiform shell and thin spirotheca, and proloculus rather large
compared with shell size.
Description.-Shell small, elongate fusiform with bluntly

Table 5.
Reg. no.
1
2

1
2

I

TGUFU1030
TGUFU1031

1
0.03
0.02

Pseudofusulina vulgaris var. fusiformis (Schellwien).

Kanuma,1959, p. 75, 76, figs. 7-11.
Pseudofusulina fusiformis (Schellwien). Thompson, 1948, pI. 12,
fig. 3 (same as pI. 15, fig. 2 of Schellwien, 1909, p. 165-168);
Morikawa, 1955, p. 98, 99, pI. 13, figs. 1-7 ; Hy. Igo, 1959, p.
246,247, pI. 3, fig. 5; Morikawa and Isomi, 1961, p. 19, 20, pI.
7, figs. 11, 12, pI. 8, figs. 12, 13, pI. 10, figs. 1-10, pI. 11, figs. 1-10,

Measurements of Pseudofusulina miyamensis Hh Igo sp nov (in mm)
Radius vector
Fig.
L.
F.R.
D.P.
W.
1
2
3
4
0.29
0.20
7-10
5.86
1.53
3.83
0.33
0.47
0.63
0.27
7-11
1.57
3.61
5.66
0.40
0.40
0.53
0.73

Thickness of spirotheca
4
2
3
0.02
0.05
0.02
0.03
0.04
0.06

.

Fusulina vulgaris var. fusiformis Schellwien, 1909, p.165-168, pI.

15, figs. 1-4.

Figures 7-10-11

5
0.83
0.87

5
0.06
0.05

Figure 11. 1-4. Neofusulinella giraudi Deprat, 1,3: axial sections, TGUFU 1090,1092,2,4: slightly oblique axial secx 30. 5-8. Neofusulinella lantenoisi Deprat, 5: slightly oblique axial section, TGUFU 1095, 6-8 :axial sections, TGUFU 1096,.1097, 1~98, x30. 9-14. Sphaerulina akudensis Hh.lgo sp. nov., 9: axial section of holotype,TGUFU 1099,10,11 : tangential sections, TGUFU 1100,1101,12,13: sagittal sections, TGUFU 1102,1103,14: axial section ofparatyp~, TGU~U 1104, x 30. 15. Schubertella pseudoobscura Chen, axial section, TGUFU 1105, x 40. 16. Nankinella sp.
tangential section, TGUFU 110~,. X10. 17-19. Nan~inella aff. kotakiensis (Fujimoto and Kawada), axial sections, TGUFU 1107:
1108,1109, X15. 20-31. Pammna(Levenella) leven! Kobayashi, 20-27,30,31: axial sections, TGUFU 1110 1111 1112 1113 1114
1115, 111~, 1117, 1.120, 1121, 28, 29: sagittal sections, TGUFU 1118,1119, x 30. 32-34. Pamirina (Levenella) 'eva/uta Shen~ andSun, axial sectlo.ns, !GUFU.1122, 1123, 1124, X 60. 35,36. Dunbarula uenoi Hh.lgo sp. nov., 35: axial section of holotype,TGUFU .1125, 36 .. axial section of paratype, TGUFU 1126, x 30. 37, 38. Minojapanella (M.) e/ongata Fujimoto and Kanuma'
37: oblique section, TGUFU 1127, X 30, 38: sagittal section, TGUFU 1128, X50.
tIO~S, TG~FU 1091,1093,

1023.

Permian fusulinaceans from Central Japan

643"'J"'"

, .

, ~ ,

"

30

31

34

"

37

38

Hisaharu Igo

644

pi. 12, figs. 1-10, pi. 13, fig. 5; Sheng, 1963, p. 66, 192, 193, pi.
16, figs. 19-21; Kanmera and Mikami, 1965, p.301,.302, pi.
52, figs. 1-6; Kalmykova, 1965, p. 122, 123, pi. 3, figs. 1-4;
Kalmykova, 1967, p. 181, 182, pi. 9, figs. 1-5; Choi, 1972,'pi. 2,
fig.7; Choi, 1973, p. 46, 47, pi. 9, figs. 1-3, pi. 11,. fig. 4;
Sakaguchi and Yamagiwa, 1975, p. 182, 183, pi. 2, figs. 1, 2,
11 ; Ozawa, 1975, pi. 9, fig. 16; Liu et al., 1978, p. 59, 60, pi.
13, fig. 1 ; Kahler and Kahler, 1980, p. 217-220, pi. 6, fig. 4, pi.
8, fig. 1 ; Ozawa and Kobayashi 1990, pi. 8, fig. 14; Ueno,
1992, p. 1283-1285, figs. 3-7-11.
Schwagerina Fusiformis (Schellwien). Suyari, 1962, p. 26, pi. 8,
figs. 7-9.
Pseudofusulina cf. Fusiformis (Schellwien). Xia, 1994, pi. 16, fig.
15.
non Schellwienia vulgaris var. Fusiformis (Schellwien). Lee, 1927,
p. 67, 68, pi. 9, figs. 3, 5.

Spirotheca thick, composed of a tectum and alveolar keriotheca. Septa fluted rather regularly, most septal loops
attain a height about 2/3 that of the chamber. Cho~ata
poorly developed in first to second volutions. Tunnel singular rather low, and narrow.
Remarks.- The present specimens are similar to SchelIwien's original specimens. This species described by
Kanmera (1958) from the Yayamadake Limestone is very
similar to the Akuda specimens.
Hy. Igo (1972) described this specie~ from North !hailandand discussed biocharacters and synonymous specimens ofprevious papers.
Occurrence.- TGUFU 1039,1041 from 8-47 and R-2 near
Higashi- and Nishi-Akuda; TGUFU 1040,1042 from R-1 in
Section E near Nishi-Akuda.

Materia/s.-Axial sections of mature specimens, TGUFU
1037,1038.
Remarks.- This well known species has been describedfrom many localities in the Tethyan region. The Akuda
specimens are identical to the original specimens describedby Schellwien (1909). Morphological variations in previously
described P. fusiformis are limited, hence we can easily
identify this species. The Hirayu specimen (Hy. Igo, 1959),
Yoganeyama specimen (Sashida, 1980), and the Akuda
specimens (Kanuma, 1959 and this paper) well coincide
morphologically with the original species.
Occurrence.-From 8-37 in Section 0 near HigashiAkuda.

Pseudofusulina houziguanica Sheng, 1963

Pseudofusulina regularis (Schellwien, 1898)
Figures 8-3-6
Fusulina regularis Schellwien, 1898, p.250, pi. 19, figs. 1-6; Deprat, 1912, p. 28, pi. 7, figs. 14, 15.
Schellwienia regularis (Schellwien). Lee, 1927, p. 50, pi. 7, figs.
8-10.
Pseudofusulina regularis (Schellwien). Fujimoto, 1936, p. 94, pi.
10, figs. 9-11, pi. 18, fig.1; Kanmera, 1958, p.194, pi. 33, figs.
1-10; Saurin and Le-Thi-Vien, 1960, p. 396, pi. 1, figs. 16-18;
Morikawa and Isomi, 1961, p. 15, pi. 7, figs. 1-10; Sada, 1964,
p.257, pi. 26, figs. 8-11, pi. 27, figs. 9, 10; Toriyama, 1975, p.
15, pi. 2, figs. 7, 8; Toriyama, 1982, pi. 3, figs. 2-4; Ishibashi,
1984, p. 211, 212, pi. 27, figs. 1-4.
Pseudofusulina (Oaixina) regularis (Schellwien). Hy. Igo, 1972, p.
105, pi. 16, figs. 10-16.
Schwagerina regularis (Schellwien). Toriyama, 1958, p.140, pi.
16, figs. 8-15; Kochansky-Devide, 1959, p.50, pi. 2, figs. 711 ; Suyari, 1962, p. 22, pi. 8, fig. 10; Liu et al., 1978, p. 44, 45,
pi. 8, fig.1; Zhou, 1991, pi. 2, figs. 6, 7.
Materia/s.-Axial sections of mature specimens, TGUFU
1039,1041 ; slightly oblique axial sections, TGUFU 1040, 1042.
Description. -Shell moderate in size, elongate fusiformwith a straight axis of coiling, having slightly inflated medianpart and bluntly pointed poles. Mature shell composed offive to six volutions. Axial length 5.26 to 6.18 mm, medianwidth 2.20 to 2.60 mm, and form ratio 2.38 to 2.39.
Proloculus moderate in size, spherical, 0.33 to 0.28 mm in its
outside diameter. First volution subspherical in shape.

Figures 8-10-13
Pseudofusulina houziguanica Sheng, 1963, p. 67, 193, 194, pi. 16,
figs. 11-14; Wang et al., 1981, p. 36, pi. 3, figs. 6, 7; Zhang
and Dong, 1986, p.100, pi. 14, fig.7; Sun and Zhang, 1988,
pi. 1, figs. 5,14.
Pseudofusulina cfr. houziguanica Sheng. Xiao, Wang, Zhang
and Dong, 1986, p. 100, 101, pi. 10, fig. 17; Ueno, 1992, figs. 53,4; Xia, 1994, pi. 16, fig. 18.
Materia/s.-Axial sections, TGUFU 1046,1047,1048,1049.
Description.-Shell medium in size, rather tightly coiled,inflated fusiform or ellipsoidal, having straight median portionand broadly rounded poles. Mature shell consists of 5 to 6
volutions. Axial length 4.06 to 5.00 mm, median width 2.20
to 2.46 mm, and form ratio 1.65 to 2.20.
Spirotheca rather thick compared with shell size andconsists of a tectum and relatively coarse keriotheca.
Septa numerous and regularly fluted throughout their length.
Septal loops almost reach upper part of chamber.
Proloculus spherical or nearly rectangular, large comparedwith their shell size, thin-walled, and external diameter 0.37
to 0.50 mm. Axial fillings rather heavy except central part ofshell. Chomata lacking except for very poorly developed
ones on surface of proloculus. Tunnel indistinct.
Remarks.-The present species is closely related to Parafusulina funafusensis Matsumaru described from the
Funafuseyama Limestone of the northern Mino Terrane.
Parafusulina funafusensis is probably a junior synonym of thepresent species.
Occurrence.-AII specimens from R-1 in Section E of
Nishi-Akuda.

Pseudofusulina krotowi (Schellwien, 1908)
Figures 8-14, 15
Fusulina krotowi Schellwien, 1908, p.190-192, pi. 20, figs. 1-10.
Schellwienia krotowi (Schellwien). Ozawa, 1925, p. 27, 28, pi. 7,
figs. 5, 6.
Pseudofusulina krotowi (Schellwien). Fujimoto, 1936, p. 82-84,
pI. 15, figs. 1-5, 9-15; Rauzer-Chernousova, 1938, p.143,
144, pI. 9, figs. 1, 2; Morikawa, 1955, p.86, pI. 14, figs. 5, 6;
Kanuma, 1959, p. 78, 79, pI. 8, figs. 1-3; Morikawa and1023.

Permian fusulinaceans from Central Japan

Isomi, 1961, p. 18, 19, pI. 8, figs. 1-11.
Kanmera, 1958, p. 193, 194,
pI. 24, fig. 20; pI. 35, figs. 13, 14; Toriyama, 1958, p.134-138,
pI. 15, figs. 8-19; Kawano, 1961, p. 84, 85, pI. 4, fig. 28; pI. 5,
figs. 1-4; Ishizaki, 1962, p. 113, 114, pI. 29, figs. 6-10.

Schwagerina krotowi (Schellwien).

Materia/s.-Axial sections, TGUFU 1050,1051.
Remarks.-The present specimens closely resemble
Schellwien's original ones. This species was describedfrom a limestone of the Okumyogata Formation about 15 km
north of Hachiman town by Kanuma (1959).
Occurrence.-TGUFU 1049,1050 from 8-46 and 8-48 in
Section D near Higashi-Akuda.

645very thin and consists of a tectum and keriotheca. Septa
numerous and thin, fluting irregular throughout shell.
Chomata poorly developed in first to second volutions.
Remarks.- The present specimens are deformed bycompaction of sediments and the height of the outer volutions has consequently been lowered.
Occurrence.- TGUFU 1070 from 8-2; TGUFU 1071 from
8-3; TGUFU 1072 from 8-4 in Section C near Akuda-kuchi.
Genus Chalaroschwagerina Skinner and Wilde, 1965

Type species.-Cha/aroschwagerina inflata Skinner and
Wilde, 1965

Pseudofusulina sp.
Chalaroschwagerina aff. chinensis (Chen, 1956)

Figures 7-12-16

Materia/s.-Axial sections of mature specimens, TGUFU
1033, 1034; axial sections of immature specimens, TGUFU
1032, 1035, 1036.
Oescription.-Shell medium in size, elongate fusiform,
having somewhat bluntly rounded poles. Mature shellconsists of four volutions.. Axial length 7.60 to 8.59 mm,
median width 2.26 to 2.73 mm, and form ratio 2.78 to 3.80 inmature specimens. Proloculus large, spherical, with outsidediameter of 0.46 to 0.58 mm.
Spirotheca rather thin, consists of a tectum and coarse
alveolar keriotheca, surface of tectum irregularly corrugated.
Septa numerous, strongly fluted throughout shell. Septal
loops rather high, almost reach top of chamber. Axial filling
irregular in axial regions of inner volutions. Tunnel single,
low, and narrow. Chomata lacking except for very poor
development in first volution.
Remarks.-This species closely resembles Pseudofusulina
minoensis Hh. Igo sp. nov., but has slightly stronger axial
filling and septal fluting than in the latter.
Occurrence.-AII specimens from R-1 in Section E of
Nishi-Akuda.
Subfamily Pseudoschwagerininae Chang, 1963
Genus Acervoschwagerina Hanzawa, 1949

Type species.-Paraschwagerina
endoi Hanzawa, 1949

(Acervoschwagerina)

Acervoschwagerina fu}imotoi Kanuma, 1959

Figures 8-18
Compare. -Schwagerina chinensis Chen, 1956, p. 6, 35, pI. 5, fig.

10.

Materia/.-Axial section of mature specimen, TGUFU 1054.
Description.-Shell large, highly inflated fusiform withrounded or elongate node-like poles and a rounded medianpart. Present specimen consists of six volutions. Axial
length 9.32 mm, median width 5.06 mm, and form ratio 1.84.
Inner volutions not so tightly coiled. Proloculus moderate insize, spherical, with an external diameter of 0.27 mm.
Spirotheca consists of a tectum and rather fine alveolar
keriotheca. Thickness of spirotheca moderate with amaximum thickness of 0.08 mm. Septa numerous, moderately fluted throughout shell. Septal folds with bluntly
pointed crests or obtuse crests extend almost halfway up
the chamber. Phrenotheca weakly developed in medianpart of outer volutions. Chomata and axial filling lacking.
Remarks.- The present specimen closely resembles
Schwagerina chinensis, which was described from Hupeh
Province by Chen (1956). However, the former differs fro~the latter in the shape of septal fluting. The present specImen is similar to some species described as Paraschwagerina, but has a larger proloculus, more loosely coiled inner
volutions, and regularly fluted septa.
Occurrence.-From R-2 in Section E of Nishi-Akuda.
Family Neoschwagerinidae Dunbar and Condra, 1927
Subfamily Neoschwagerininae Dunbar and Condra, 1927
Genus Mak/aya Kanmera and Toriyama, 1968

Figures 10-1-3
AceNoschwagerina fujimotoi Kanuma, 1958, p. 62, 63, pI. 4, figs.

6-8; Ishii, 1964, p.125, pI. 1, figs. 1, 2.

Materia/s.-Oblique sections, TGUFU 1070,1071; slightly
oblique axial section, TGUFU 1072.
.
Oescription.-Shell very large, elongate fusiform, andhaving inflated median part and broadly ~r narro~ly roun?ed
poles. Inner three or four volutions very tightly COiled, heightsuddenly increases in outer volutions.
Axial length of present specimens more than 15.98.mm
and median width 6.00 mm or more. Proloculus spherical,
small, and its outside diameter 0.27 to 0.47 mm. Spirotheca

Type species.-Cancellina pamirica Leven, 1967

Mak/aya pamirica (Leven, 1967)
Figures 10-7-10
Gancellina pamirica Leven, 1967, p. 186, 187, pI. 32, figs. 1, 3; Sun

and Zhang, 1988, pI. 4, fig. 21.
Kanmera and Toriyama, 1968, p. 3437, pI. 4, figs. 1-16; Kahler and Kahler, 1979, p'.249, 250, pI.
8, fig. 6; Ishibashi, 1984, p. 222, 223, pI. 31, fig. 16; Yang,
1985, pI. 2, fig. 3; Ozawa and Kobayashi, 1990, pI. 9, figs. 10,
11; Fan et a/., 1990, pI. 7, figs. 19, 27; Ueno, 1991, p.994,

Mak/aya pamirica (Leven).

Hisaharu Igo

646

Armenina sp.

995, figs. 9-5-10.
Materia/s.-Axial sections, TGUFU 1076, 1077, 1O~8, 107~.
Oescription.-Shell medium in size, subsphencal withbroadly rounded poles. Mature specimens consist of 9 to 10
volutions. Axial length 2.10 to 2.56 mm, median widt~ 1.83
to 2.33 mm, and form ratio 1.10 to 1.30. Proloculus minute,spherical, with an external diameter of 0 ..06 to 0.08.mm.
Spirotheca rather thick compared with shell size andcomposed of a tectum and alveolar keriotheca. Thick.nessof spirotheca in first to tenth volutions of mature specimen
0.005, 0.008, 0.020, 0.030, 0.030, 0.050, 0.050, 0.050, 0.070,
and 0.070 mm, respectively. Short, broad, and fan-shaped
primary transverse septula well developed in fourth or succeeding volutions. Septa thick and unfluted thr?ugh?ut
shell. Parachomata nearly triangular in axial section, first
appeaing in third volution. No ~ial septula p~esent.
Remarks.-Recently, this species was descnbed from the
Akiyoshi Limestone by Ueno (1991). The Akuda specimens
closely resemble the Akiyoshi specimens.
Occurrence.-TGUFU 1076,1077,1078 from A-21 ; TGUFU
1079 from A-22 in Section A near the Horikoshi Pass.

Maklaya saraburiensis Kanmera and Toriyama, 1968
Figures 10-11,12
Maklaya saraburiensis Kanmera and Toriyama, 1968, p. 41-43, pI.

4, figs. 17-20 ; Ueno, 1991, p. 993, 994, figs. 9-1-4.
Neoschwagerina saraburiensis (Kanmera and Toriyama).

Han,

1985, p. 685, pI. 2, figs. 14, 15.
Materia/s.-Axial sections of mature specimens, TGUFU
1080,1081.
Oescription.-Shell small, subspherical with bluntly pointed
poles. Mature specimens having seven and nine volutions,1.83 and 1.86 mm in axial length, 1.27 and 1.57 mm in medianwidth, and 1.18 and 1.44 in form ratio respectively. Inner twovolutions planispiral with a short axis of coiling. Proloculus
small, spherical, 0.05 mm in outside diameter. Spirothecacomposed of a tectum and alveolar keriotheca. Thicknessof spirotheca of first to seven volutions of Figure 10-11, 0.007,
0.007, 0.012, 0.025, 0.030, 0.050, and 0.060 mm, respectively.
Short and low primary transverse septula first appear infourth volution. Parachomata observed in third or succeeding volutions. No axial septula present.
Remarks.- This species is distinguished from Mak/ayapamirica (Leven) by a larger shell and greater numbers ofvolutions. The Akuda specimens closely resemble the typespecimen, which was described from the Rat Suri Limestone
in the Khao Ph long Phrab area, central Thailand.
Occurrence.- TGUFU 1080,1081 from A-22 in Section A
near the Horikoshi Pass.

Family Verbeekinidae Staff and Wedekind, 1910
Subfamily Verbeekininae Staff and Wedekind, 1910
Genus Armenina Miklukho-Maklay, 1955
Type species.-Armenina karinae Miklukho-Maklay, 1955

Figures 10-13, 14
Materia/s.-Axial sections of mature specimens, TGUFU

1082,1083.
I
Oescription.-Shell small, subspheri~al with r?unded ~o arregions. Mature specimens having eight to nine. volutlo.ns,
1.67 to 1.76 mm in axial length, 1.33 to 1.40 mm In me?lanwidth, and form ratio 1.26 to 1.76. Inner 1/~ volutl?ns
planispiral, with a short axis of coiling and havl.ng a. nght
angle to outer volutions. Inner three to four volutlons tlg~tlY
coiled and outer ones loosely coiled. Radius vectors of firstto eighth volutions of axial section (Figure 10-13) 0.08,0.11,
0.16, 0.25, 0.34, 0.45, 0.60, and 0.76 mm, re~pectiv~IY. .
Proloculus small, spherical, and 0.05 mm In outSide diameter. Spirotheca very thin for the genus, compo~ed Of. atectum and fine alveolar keriotheca except for the Inner firstto second volutions, which are a dense single layer. Thickness of spirotheca of first to ninth volutions (Figure 10-14)
0.007,0.013,0.013,0.013, 0.010, 0.020, 0.030, 0.030, and
0.030 mm, respectively.
Parachomata small, low, narrow, and reach about 1/4 or
1/3 of chamber height. They appear in third volution. No
axial septula present.
Remarks.- This species closely resembles Verbeekina(Armenina) prisca Toriyama and Kanmera, but differs from thelatter in having a larger shell and rudimentary parachomata.
Occurrence.-From A-21 in Section A near the Horikoshi
Pass.
Repository: All specimens described in this paper are deposited in the Department of Astronomy and Earth Sciences,
Tokyo Gakugei University with the prefix TGUFU.

Acknowledgments
I would like to acknowledge Professor Hisayoshi Igo of theInstitute of Geoscience, University of Tsukuba for his criticalreading of an early version of the manuscript and providing
useful information on fusulinaceans in the Mino Terrane. I
thank Drs. Katsumi Ueno and Shuko Adachi of the same
university for their valuable discussion and help in photographing specimens. I also thank Mr. Junichi Yamamoto, ateacher of Sannou Elementary School in Tokyo. He collaborated with me in the field and prepared many thinsections when he was a student of our university.

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650

Hisaharu Igo

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 184, pp. 651-661, 5 Figs., December 30,1996© by the Palaeontological Society of Japan 1996

1

1024

1

An early Middle Miocene warm-water molluscan
fauna in southwestern Sakhalin - Molluscs fromthe Ausinskaya Formation around Novoselovo-

KAZUTAKA AMANO\ VLADIMIR D. KHUDIK2 and KEN NARITA3'Department of Geoscience, Joetsu University of Education, Joetsu, Niigata Prefecture, 943 Japan

2Far East Geological Institute, Russian Academy of Sciences, Vladivostok 690022, Russia
3Shinshu-Shinmachi Fossil Museum, Shinshu-Shinmachi, Nagano Prefecture 381-24, Japan
Received 29 May 1996; Revised manuscript accepted 20 August 1996

Abstract Thirty-two mollusc species were identified from two localities of the lower Middle Miocene
Ausinskaya Formation around Novoselovo, southwestem Sakhalin. This fauna includes a number ofwarm-water taxa such as Anadara watanabe;' G/ycymeris cisshuensis, Dosinia tugaruana, Tateiwaia sp.,
Euspira meisensis, and Cultellus izumoensis. These taxa were previously unknown in the Ausinskaya
Formation, except for Cultellus izumoensis. A Dosinia-Anadara association made up of subtropical totemperate-water euneritic dwellers is prominent in this fauna, as is a temperate- to cold-water, eu- tomesoneritic Spisula association that has been recognized in lower Middle Miocene strata of northwestern
Hokkaido and northwestern Sakhalin. The cooccurrence of these two ecologically dissimilar faunal
associations implies that adjacent warm (inshore) and cold (offshore) water masses were present within
the temperate marine realm that prevailed in Hokkaido and in southwestern Sakhalin during the early
Middle Miocene.
Key words: Early Middle Miocene, Ausinskaya Formation, warm-water, Mollusca, Sakhalin
Introduction
Neogene strata in southwestern Sakhalin consist of the
Kholmskaya, Nevel'skaya, Chekhovskaya, Verkhneduiskaya,
Ausinskaya, Kurasiiskaya and Maruyamskaya Formations, inascending order (Table 1; Smekhov, 1948; Interdepartmental Committee on Stratigraphy, 1961, 1976). Of these, theupper part of the Chekhovskaya Formation near Novoselovo
has yielded a K-Arage of 16.9Ma (Okamura, 1994), and theAusinskaya Formation in the Makarov Region of eastern
Sakhalin has been correlated with the Denticulopsis lauta
Zone of Koizumi (1985) by Arkhipova et al. (1992). Therefore, we regarded the age of the Ausinskaya Formation insouthwestern Sakhalin as early Middle Miocene.
Molluscs from the Ausinskaya Formation were studied by
Russian workers (llyina, 1954; Krishtofovich, 1964; Zhidkova
et al., 1968; Khudik, 1989). According to Zhidkova et at.
(1968), the stratotype of the Ausinskaya Formation is situatedalong the Bezymyannaya (old name, Ausi; now Tolkaya)
River just south of the Arkansas River. lIyina (1954) reported
abundant mollusc species in the upper part of the Uglegorskaya Formation (= Ausinskaya Formation) between the
Bezymyannaya and Novoselka Rivers, and she describedand illustrated nine new species from there. Khudik (1989)

noted the following species present chiefly in the uppermost
part of the Ausinskaya Formation: Ciliatocardium shinjiense(Khramova), Serripes groenlandicus (Bruguh3re), Peronidia
pulchra (Slodkewitsch), Macoma calcarea (Gmelin), Macoma
nasuta (Conrad), Dosinia ausiensis lIyina, Cultel/us izumoensis Yokoyama, Panomya simotomensis Otuka and Mya
cuneiformis (B6hm). However, the fauna from the main partof the Ausinskaya Formation has never been thoroughly
examined or compared with the early Middle Miocene

Table 1. Neogene stratigraphy of South Sakhalin (Smekhov, 1948: Interdepartmental Committee on Stratigraphy, 1961, 1976).
Age

Formations

Pliocene

Maruyamskaya
Kurasiiskaya
Ausinskaya
Verkhneduiskaya
Chekhovskaya
Nevel'skaya
Kholmskaya

MioceneKazutaka Amano et a/.

652

s

t

LEGEND

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DB
~c

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QJ]o

~ ...
/ TCri:;i Y• '\DE... Mt. Ustaya \

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N-?voselo vo,

y .. hlo.Slk~.Il .. k

LEGEND• Fossil localities~~ Sections examined0lkm

Figure 1. Fossil localities around Novoselovo in southwestern Sakhalin.

=

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'<U

e....
o

~

'"

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'"

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faunas of Hokkaido, which are the geographically nearest
coeval faunas.
In his summary of early Middle Miocene marine climates innorthern Japan, Chinzei (1986) assigned the Chikubetsu
fauna (Uozumi, 1962) of northwestern Hokkaido to the coldwater realm. He put northwestern Hokkaido in the subarctic (?) region on his proposed climato-geographic map. On
the other hand, Ogasawara (1994) judged the Chikubetsu
fauna to be a cool-temperate one, based on the cooccurrences of Spisula equilateralis, Platyodon nipponica andMercenaria sp. along with Anadara, Dosinia and Sinum (Noda,
1992). None of these workers discussed coeval faunas of
Sakhalin, such as the Ausinskaya fauna. The examinationof Ausinskaya molluscs herein is made to address thedifferences in paleoclimatic interpretations noted above. To
this end, we collected many mollusc species from the
Ausinskaya Formation at the localities along the coast near
Novoselovo (Figure 1). The purposes of this study are todescribe these species, to compare them to species fromcoeval faunas in Japan, and to discuss their biogeographic
significance.

Geologic setting and localities
The Chekhovskaya, Ausinskaya and Kurasiiskaya Formations are exposed around Novoselovo (Figure 1). The Chekhovskaya Formation is composed of lapilli tuff, tuff breccia
and basaltic hyaloclastite (Figure 2), whereas the Ausinskaya
Formation consists of fine- to medium-grained sandstoneintercalated with two lignite seams and siltstone. An unconformable relationship between the Chekhovskaya and
Ausinskaya Formations is observed in coastal cliffs about
800 m northeast of the mouth of the Novoselka River, where
the basal conglomerate of the Ausinskaya Formation directly
overlies the lapilli tuff of the Chekhovskaya Formation. The
Ausinskaya sandstone bed in this area strikes N53°E andEJG
Xt.1(2 J

.5'"

'"
<
;::I

.

....
>
'"o
....c
»

"

.c
U

Figure 2. Columnar sections of Neogene formations
around Novoselovo (N and S in Figure 1). Legend: A. mudstone; B. siltstone; C. fine-grained sandstone; D. pebblebearing fine-grained sandstone; E. medium-grained sandstone; F. pebble-bearing medium-grained sandstone; G.
lapilli tuff; H. basaltic hyaloclastite; I. lignite; J. fossil horizons.

dips 16W, while the Chekhovskaya tuff bed strikes N400E
and dips 15°E. The Kurasiiskaya Formation conformably
overlies the Ausinskaya Formation and is subdivided into alower part consisting of dark gray mudstone and an upper
part made up of hard siltstone.
Both the Ausinskaya and Kurasiiskaya Formations havewestward dips in an area between about 800 m east and 1.3
km west of the mouth of the Novoselka River. However, tothe west of the present area, these strata strike N54-79°W
and dip 17-40oE, which suggests the presence of a small
basin-like structure in the area of Novoselovo.
We collected many mollusc fossils from the following two
localities.
Loc. 1. Coastal cliff about 800 m northeast of the mouthof the Novoselka River; tuffaceous pebble-bearing sandstone containing abundant plant debris, just above the basal
conglomerate of the Ausinskaya Formation.
Loc. 2. Coastal cliff about 2.5 km southwest of the mouth

1024.

Miocene Ausinskaya molluscan fauna of Sakhalin653

of the Novoselka River; medium-grained sandstone withcalcareous concretions; upper part of the Ausinskaya Formation.

Most bivalves occur as articulated specimens, which suggest that the fauna is autochthonous. This assemblage
comprises the Dosinia-Anadara association, consisting of
Dosinia (Phacosoma) tugaruana, D. (Kaneharaia) ausiensis,

Molluscan associations in the Ausinskaya Formation

Anadara watanabei, Glycymeris cisshuensis, Cultel/us
izumoensis, Mercenaria yiizukai, and Euspira meisensis.
The species composition of the Dosinia-Anadara association is similar to that of the Mercenaria assemblage (Noda,
1992) from the lower Middle Miocene Chikubetsu Formationin northwestern Hokkaido, and to the Dosinia-Clinocardium
assemblage (Chinzei and Iwasaki, 1967) from the lower

A total of 32 species of molluscs, including 25 bivalves
and 7 gastropods, were collected from the two localities
noted below (Table 2). Thirteen bivalve and four gastropod
species were identified, of which only four species had beenpreviously described and illustrated by lIyina (1954) andZhidkova et al. (1968) from the Ausinskaya Formation:

Middle Miocene Kadonosawa Formation in northeastern
Honshu. According to Noda (1992), the Mercenaria assemblage is characterized by Mercenaria yiizukai, Anadara oga-

Modiolus tetragonalis, Dosinia ausiensis, Cultel/us izumoensis
and Reticunassa ausiensis.
The living counterparts of the species from Loc. 1 dwell invarious kinds of substrates, including rock (Modiolus), sandand gravel (Chlamys), muddy sand (Dosinia), and mud (Tateiwaia). Moreover, most bivalves at this locality occur asdetached valves, suggesting an allochthonous nature.
Many sandy bottom dwellers such as Dosinia and Anadara

and Cultel/us izumoensis. This assemblage is considered
to have lived in the euneritic zone in subtropical to temperatewaters.
The Dosinia-Clinocardium assemblage, on the other hand,is composed of Dosinia (Phacosoma) nomurai, Clinocardium

were obtained from medium-grained sandstone at Loc. 2.

shinjiense, Anadara ninohensis, Clementia iizukai, Glycymeris

wai, Dosinia (Kaneharaia) kaneharai kannoi (=0. ausiensis)

Table 2. Molluscan fossils from the Ausinkaya Formation around Novoselovo.
on the right are specimens from each locality.
Taxa

Localities

Numbers2

Anadara (Anadara) watanabei (Kanehara)

3

A. ? sp.
Glycymeris (Glycymeris) cisshuensis Makiyama
G.(G.) idensis Kanno

1

8

G.(G.) sp.
Modiolus tetragonalis Siodkewitsch
Chlamys (Chlamys) arakawai (Nomura)
C. (Chlamys) otukae Masuda and Sawada

1

3

C. sp.
Nanaochlamys sp.
Clinocardium ct. decoratum (Grewingk)
C. aft. nuttal/ii (Conrad)C. sp.
Macoma sp.
Spisula (Pseudocardium) haboroensis (Yokoyama)
Oxyperas? sp.
Solen sp.
Cultel/us izumoensis Yokoyama
Mercenaria yiizukai (Kanehara)
Dosinia (Phacosoma) tugaruana Nomura
D. (Kaneharaia) ausiensis Ilyina
Saxidomus sp.
Potamocorbula aft. amurensis (Schrenck)
Panopea japonica A. Adams
Thracia pertrapezoidea Nomura
Tateiwaia sp.
Crepidula jimboana Yokoyama
Euspira meisensis (Makiyama)
G/ossaulax didyma coticazae (Makiyama)
Boreotrophon? sp.
Reticunassa ausiensis (llyina)
Plicifusus sp.

2

2
3
12
7

2

5
1
4
1
6

2

2

654

Kazutaka Amano et al.

cisshuensis, Siratoria siratoriensis, and Euspira meisensis,
which are thought to be representatives of a sandy bottom
habitat below the low-tide line (Chinzei and Iwasaki, 1967).
The bivalves, Dosinia (Kaneharaia) ausiensis, Cultellus
izumoensis and Mercenaria yiizukai cooccur in the Dosinia
Anadara association of the Ausinskaya Formation and the
Mercenaria assemblage from the Chikubetsu Formation.
These species are not present in the Dosinia-Clinocardium
assemblage of the Kadonosawa Formation, although somewarm-water taxa, including Clementia iizukai and Siratoria
siratoriensis, are characteristic species only in the Kadonosawa assemblage.

Biogeographic significance of the Ausinskaya fauna
The Ausinskaya fauna contains many subtropical to temperate water species (Uozumi, 1962; Ogasawara, 1994) suchas Anadara watanabei, Glycymeris cisshuensis, Cultellus
izumoensis, Dosinia tugaruana, Tateiwaia sp., and Euspirameisensis. Tateiwaia is an especially characteristic genusof the tropical to subtropical Arcid-Potamid fauna (Tsuda,1965).
It is noteworthy that the fauna around Novoselovo shares
many species in common with the early Middle Miocene
fauna of the Tanosawa Formation in Nishitsugaru district,
Aomori Prefecture (Nomura, 1935; Ogasawara, 1977).
These include Glycymeris vestitoides (=G. cisshuensis),
Chlamys arakawai, Cultellus izumoensis, Dosinia tugaruana,
Thracia pertrapezoidea and Euspira meisensis. The Nishitsugaru district at that time lay in a subtropical zone (Chinzei,1986).
In contrast, only two cold-water species (Uozumi, 1962),
Spisula haboroensis and Mercenaria yiizukai, were obtained
from the upper horizon (Loc. 2) of the Ausinskaya Formation.
Mercenaria yiizukai is one of the characteristic species of the
Chikubetsu fauna of northwestern Hokkaido (Kanno andMatsuno, 1960).
The Chikubetsu fauna is subdivided into the Penitella,
Spisula, and Mercenaria assemblages (Noda, 1992). The
Spisula assemblage is characterized by the dominant occurrence of Spisula equilateralis (Clark) (= S. voyi onnechiuria(Otuka)), in association with Clinocardium spp., Serripes
groenlandicus (Bruguiere), Megangulus tmatumotoi (Otuka)(=M. protovenulosus (Nomura)) and Macoma optiva (Yokoyama). Noda (1992) inferred that this assemblage flourished in the eu- to mesoneritic zone in temperate to cold
waters. We found this assemblage in the stratotype of thelower Middle Miocene Sertunaiskaya Formation along thecoast near Mgachi, northwestern Sakhalin.
As noted above, the species composition of the DosiniaAnadara association in the Ausinskaya fauna resembles thatof the Mercenaria assemblage in the Chikubetsu fauna whichis estimated to have lived in the euneritic zone of subtropical to temperate waters.
The cold-water Spisula and temperate water DosiniaAnadara associations coexisted in Sakhalin and northwestern Hokkaido during the early Middle Miocene (Figure 3). It
is suggested that such climatologically different faunal
associations could hardly be reconciled to dwelling together,• Dosinia-Anadara Assoc.
..SpisuJa Assoc.

Okhotsk Sea

Figure 3. Distribution of Dosinia-Anadara and Spisulaassociations in northern Japan and Sakhalin during early
Middle Miocene time.

but could have lived at distinctive depths, the cold-water
Spisula association inhabiting slightly deeper water than thewarm-water Dosinia-Anadara association. Uchimura andMajima (1992) noted for early Middle Miocene molluscs of theFuranui Formation in central Hokkaido that warm-water
species were abundant in shallow-water deposits above
normal wave base, whereas cold-water species lived indeeper water. These occurrences suggest that adjacent
but thermally different water masses prevailed in both northwestern Hokkaido and southwestern Sakhalin during theearly Middle Miocene. A similar situation occurs today in
mild-temperate waters along the Joban coast, Fukushima
Prefecture, northeastern Honshu (Nemoto and Akimoto,1990). Oxygen isotope paleotemperatures derived from thebivalves Mya cuneiformis and Macoma nasuta in the Ausinskaya Formation are 15-16.5'C (Khudik et aI., 1984), whichapproximates the mean annual surface temperature along
the modern Joban coast.
We conclude that southwestern Sakhalin lay in a temperate zone rather than a subboreal or subarctic zone during theearly Middle Miocene. This climato-geographic scenario
generally agrees with the coeval paleoclimate inferred for1024.

Miocene Ausinskaya molluscan fauna of Sakhalin

northwestern Hokkaido by Ogasawara (1994).

However the

sub~ivision of the early Middle Miocene temperate m~rineprovln~e o! :his region into warm, mild- and cool-temperatezones IS difficult. The absence of clearly distinct temperature-based faunal associations in central Hokkaido andsouthwestern Sakhalin makes such subdivisions uncertain
as mentioned above.
'655

thread. However, Krishtofovich (1964) gave no descriptionof the U~ndi~skaya specimen, so it is difficult to exactlycompare It with Chlamys arakawai.

Measurements (in mm).Specimens
Length Height
JUE no. 15574-1
57.1 +
84.3
JUE no. 15574-2
59.6* number of ribs

NR*

23
23

Valve
left
right

Systematic notes on some species
Class Bivalvia
Family Pectinidae
Genus Chlamys Reding, 1798
Subgenus Chlamys Reding, 1798

Chlamys (Chlamys) arakawai (Nomura, 1935)
Figures 4-14, 15
Pecten islandicus MOiler. Matsumoto, 1930, p. 104, pI. 40, fig. 9(non MOiler, 1776).
Pecten (Pecten) arakawai Nomura, 1935, p.4O, 41, pI. 4, figs. 1, 2.
Pecten (Chlamys) arakawai Nomura. Nomura, 1940, p.17, pI. 2,

figs. 1-3.
Coralichlamys shigemai Hirayama, 1954, p. 51, 52, pI. 3, fig. 2.
Chlamys arakawai (Nomura). Masuda, 1954, p. 150, 152, pI. 19,

figs. 1-6; Sawada, 1962, p.67, 68, pI. 6, fig. 20; Hirayama,
1967, p.390, pI. 1, figs. 13-17; Ogasawara et al., 1989, p.77,
pI. 3, fig. 1 ; Sato, 1991, p. 30-39, pI. 1, figs. 1-9, pI. 2, figs. 1-9,
pI. 3, figs. 1-6; Kobayashi and Ueda, 1991, p.132, pI. 4, figs.
4,5.
Chlamys cf. arakawai (Nomura). Iwai, 1960, pI. 1, fig. 9.
Chlamys (Chlamys) arakawai (Nomura). Kotaka and Noda, 1967,
pI. 2, fig. 3.

Remarks and comparison.- Three specimens wereobtained from Loc. 1. Chlamys arakawai from the Ausinskaya Formation is large in size (maximum height=B4.3 mm),higher than long, and moderately inflated. There are 23
weakly granulated radial ribs with one interstitial thread and23 rounded radial ribs on the left valve. Anterior auricle ofleft valve is large, with eight radial threads.
Krishtofovich (1964, pI. 17, fig. 1) illustrated a species living
around the Japanese Islands, Chlamys (Chlamys) farreri
nipponensis Kuroda, from the Miocene Uandinskaya Formation in northwestern Sakhalin. His Uandinskaya specimen
is closely allied to Chlamys arakawai in its shell size (height=
75.0 mm), proportions (height/length ratio = 1.09), number ofribs (23 ?), and having granulated ribs with one interstitial

Distribution.-Ausinskaya Formation of Sakhalin; Kaigarabashi Member in southwestern Hokkaido; Tanosawa,
Moniwa, Otsutsumi, Yanagawa and Orito Formations in
Honshu; early Middle to late Middle Miocene.

Chlamys (Chlamys) otukae Masuda and Sawada, 1961
Figures 4-16
Chlamys sp. Otuka, 1935, pI. 55, fig. 142.
Chlamys cf. hataii Masuda and Akutsu, 1956, pI. 20, figs. 7-9.
Chlamys otukae Masuda and Sawada, 1961, p. 19, pI. 4, figs. 1-5;

Masuda, 1962, p.182, pI. 19, figs. 13, 14, pI. 21, fig. 12; Ogasawara, 1976, p. 41, 42, pI. 2, figs. 1, 2; Bito et al., 1980, pI. 1,
figs. 8, 10; Masuda et al., 1989, p.16, 17, pI. 2, figs. 7, 9, 10 ;Sato, 1991, p. 52-56, pI. 8, figs. 1-14.

Remarks and comparison.-One right valve was obtained
from Loc. 2. The shell is small in size (length=19.1 mm),slightly inflated and somewhat higher than long. The valve
is ornamented with 22 flat round-topped and smooth radialribs. The radial ribs are broader than their interspaces,
which have one intercalary thread.
The present species shows a wide range of variation in thebifurcation mode of its radial ribs (Sato, 1991). The early
portion of the primary ribs of the Ausinskaya specimen is
subdivided into two or three parts, making it look like a
specimen from the Kaigarabashi Member in Hokkaido illustrated by Masuda and Sawada (1961, pI. 4, fig. 4).
The present species is morphologically very similar toChlamys akitana (yokoyama) but differs from the latter inhaving a less inflated shell and fewer radial ribs.
Measurements (in mm).Specimens
Length Height
JUE no. 15575
18.1
19.1
* number of radial ribs

NR*

22

Valve
right

Distribution.-Ausinskaya Formation in Sakhalin; Kaiga-

Figure 4. All figures natural size, unless otherwise stated. 1. Glycymeris (Glycymeris) idensis Kanno, Loc. 2, JUE no.
15582. 2. Modiolus tetragonalis Siodkewitsch, Loc. 1, JUE no. 15583, x 1.5. 3,8. Dosinia (Kaneharaia) ausiensis lIyina, Loc.
2, JUE nos. 15580-3,15580-1. 4. Anadara (Anadara) watanabei (Kanehara), Loc. 2, JUE no. 15584. 5. Panopea japonica A.
Adams, Loc. 1, JUE no. 15585. 6,7. Dosinia (Phacosoma) tugaruana Nomura, 6: Loc. 1, JUE no. 15577, 7: Loc.2, JUE no.
15578-1, x1.2. 9. Mercenaria yiizukai (Kanehara), Loc. 2, JUE no. 15586. 10. Clinocardium aft. nuttalfii (Conrad), Loc. 1, JUE
no. 15587. 11. Euspira meisensis (Makiyama), Loc.2, JUE no. 15588. 12. G/ossaulax didyma coticazae (Makiyama), Loc.1,
JUE no. 15589. 13. Glycymeris (Glycymeris) cisshuensis Makiyama, Loc.2, JUE no. 15590. 14,15. Chlamys (Chlamys)
arakawai (Nomura), Loc. 1, JUE nos. 15574-1,15574-2. 16. Chlamys (Chlamys) otukae Masuda and Sawada, Loc. 2, JUE no.
15575, X1.5. 17. Nanaochlamys sp., Loc. 1, JUE no. 15576.

656

Kazutaka Amano et at.

1024.

Miocene Ausinskaya molluscan fauna of Sakhalinrabashi Member in Hokkaido; Metoki, Moniwa, Oido, Otsutsumi, Utsuno, Nanao, Ogane Formations in Honshu; early
Middle to late Middle Miocene.
Genus Nanaochlamys Hatai and Masuda, 1953

Nanaochlamys sp.
Figure 4-17
Description.-One left valve was obtained from Loc. 1.
Shell medium in size, flat and subequilateral except forauricles. Apical angle 95°. Surface ornamented with radialribs and a weak concentric constriction. Primary radial ribs
five, provided with six roof-shaped riblets separated bybroader interspaces. Secondary radial ribs four, low, andornamented with three rib lets. Interspaces of primary andsecondary ribs sculptured with three or four roof-shaped
riblets. Anterior auricle much larger than posterior one,
provided with five radial striations. Posterior auricle with six
radial threads.
Measurements (in mm).-

Specimens
JUE no. 15576

Length63.1

Height
69.6

Valve
left

Remarks and comparison.- The present specimen is very
similar in appearance to Nanaochlamys otutumiensis (Nomura and Hatai, 1937), especially to specimens from the Middle
Miocene Suenomatsuyama Formation illustrated by Masuda(1960, pI. 39, fig. 7). However, the present species differsfrom N. otutumiensis in having a flat shell and several roofshaped riblets on the back of the primary and secondary ribs
as well as within the interspaces between the ribs. In its
early juvenile stage, N. otutumiensis sometimes has a flat left
valve like the Ausinskaya specimen. So, it is possible toconsider the Ausinskaya species as an ancestral form of N.
otutumiensis.

It is easy to discriminate the Ausinskaya specimen from
Swiftopecten swiftii (Bernardi), because it has a larger apicalangle, a few secondary ribs, and many roof-shaped riblets.
Family Veneridae
Genus Dosinia Scopoli, 1777
Subgenus Phacosoma Jukes-Brown, 1912

Dosinia (Phacosoma) tugaruana Nomura, 1935
Figures 4-6, 7
Dosinia tugaruana Nomura, 1935, p. 58, 59, pI. 6, fig. 6.
Dosinia odosensis Nomura, 1935, p. 59, 60, pI. 6, fig. 2.
Dosinia (Phacosoma) tugaruana Nomura. Masuda, 1963, p.32,
34, pI. 6, figs. 8-10; Gladenkov et al., 1987, p.35, 36, pI. 7,

figs. 2, 7, 8, 10, pI. 8, figs. 5, 6, 12.
Itoigawa et al., 1974, p.87, pI.
23, fig. 7.
Phacosoma odosensis (Nomura). Itoigawa et al., 1981, pI. 15, fig.
4; Itoigawa et al., 1982, p. 81, 82.

Dosinorbis odosensis (Nomura).

Remarks.-Twelve specimens were obtained from Loc. 2657

and one from Loc.1. The Ausinskaya specimens are small
to medium in size, moderately inflated, and subcircular inform. Pallial sinus narrow, bluntly pointed at end, extending
to central part of valve.
Dosinia tugaruana and D. odosensis were described at thesame time by Nomura (1935) from the Miocene Tanosawa
Formation in Aomori Prefecture, northern Japan. As first
noted by Masuda (1963), the two species are likely to besynonyms.
Iwai (1961) illustrated a specimen as Dosinia tugaruana from
the Miocene Ainaigawa Formation and later (Iwai, 1965)
illustrated it again under the name Dosinia akaisiana Nomura.
Poor preservation of the Ainaigawa speCimen prevents us
from evaluating its species assignment.
Measurements (in mm).-

SpeCimens
JUE no. 15577
JUE no. 15578-1
JUE no. 15578-2
JUE no. 15578-3

Length
33.1
33.1
36.5
17.7

Height
30.3
30.6
35.8
17.1

Depth
12.9
13.7
8.2

Valve
right
both
both
both

Distribution.-Kakertskaya Formation and Ezhovii Horizonof Kamchatka; Ausinskaya Formation of Sakhalin; Tanosawa Formation and Mizunami Group of Honshu; all early
Middle Miocene.

Subgenus Kaneharaia Makiyama, 1936

Dosinia (Kaneharaia) ausiensis lIyina, 1954
Figures 4-3, 8
Dosinia ausiensis lIyina, 1954, p. 228, 229, pI. 18, figs. 7, 8; Zhidkova et al., 1968, p. 107, 108, pI. 5, fig. 2, pI. 18, figs. 7, 8.
Dosinia mirabi/is Uozumi (MS). Fujie et al., 1964, pI. 6, fig. 4.
Dosinia (Kaneharaia) kaneharai kannoi Masuda. Masuda, 1967,
pI. 2, fig. 9; Noda, 1992, p. 83, 84, pI. 4, fig. 1.
Dosinia (Kaneharaia) kaneharai rumoiensis Amano, 1983, p. 50, 51,

pI. 4, fig. 12, pI. 5, figs. 1, 5, 10.
Dosinia (Kaneharaia) kaneharai uandiensis Sinelnikova in Gladenkov et al., 1987, p.35, pI. 7, figs. 4, 6.
Dosinia (Kaneharaia) ausiensis lIyina. Gladenkov et al., 1987, p.

35, fig. 3; Amano and Sato, 1995, p.7, figs. 4-3, 6,12.
Shimizu and Fujii,
1995, fig. 6.

Dosinia (Kaneharaia) kaneharai Yokoyama.

Remarks.-Only one specimen was collected from Loc. 1
and seven from Loc. 2. Dosinia ausiensis is small to
medium in size (maximum length = 50.6 mm) and subcircular
to ovately elongate in shape. Surface is sculptured byrather fine concentric cords which are rounded and elevatednear the ventral margin. Number of cords between 1 cm and2 cm from the beak is 15. Pallial sinus triangular, shallow,
and wide.
The present species was established by lIyina (1954),
based on a specimen from the Uglegorskaya (now Ausinskaya) Formation at Bezymyannii (old Japanese name, Ausi).
When he summarized Japanese Kaneharaia, Masuda (1967)
illustrated a specimen from the Ausi River (type locality of658

Kazutaka Amano et al.

Dosinia ausiensis) under the name D. (Kaneharaia) kaneharai
kannoi. Noda (1992) also described and illustrated D. (K.)
kaneharai kannoi from the early Middle Miocene Chikubetsu

Formation of northwestern Hokkaido. However, judging
from his figure, Noda's (1992) Chikubetsu specimen is better
identified with D. ausiensis, because of its subcircular shellform and many fine concentric cords.
Amano (1983) proposed Dosinia (Kaneharaia) kaneharai
rumoiensis from the Middle Miocene Togeshita Formation of
Hokkaido, and Sinel'nikova in Gladenkov et al. (1987) described D. (K.) kaneharai uandiensis from the Miocene
Uandinskaya Formation in northwestern Sakhalin. However, it is difficult to separate these subspecies from D.
ausiensis using ornamentation, shell size, shell shape or theform of the pallial sinus. Therefore, these subspecies are
considered to be synonyms of the present species.
Comparison .-The present species is closely related to
Dosinia (Kaneharaia) kannoi Masuda in its shell size andnumber of concentric cords (11 -16 between 1 cm and 2 cmfrom the beak). However, this species differs slightly from D.

(K.) kannoi in having a more inflated shell and round - topped

as well as elevated concentric cords near the ventral margin.
It is easy to distinguish D. (K.) kaneharai Yokoyama from D.
(K.) ausiensis, because the former species has an ovate shelland fewer concentric cords (8- 11 between 1 cm and 2 cmfrom the beak). D. (K.) ausiensis ranges from the lower
Middle Miocene to the Lower Pliocene, whereas D. (K.)
kaneharai ranges from Middle to Upper Miocene. Stratigraphically, the range of the latter species is shorter thanthat of the former.
Measurements (in mm). -Specimens
JUE no. 15579
JUE no. 15580-1
JUE no. 15580- 2

Length
50.6
39.2
30.6

Height
54.1
37.2
26.5+

Depth
19.0+

Distribution.- Early Middle Miocene Ausinskaya and Uandinskaya Formations in Sakhalin ; early Middle Miocene

Figure 5. All figures natural size, unless otherwise stated. 1,2. Cultel/us izumoensis Yokoyama, Loc. 2, JUE
nos. 15591 -1, 15591 -2. 3a- b. Tateiwaia sp., Loc. 1, JUE no. 15581, x 2. 4a- b. Oxyperas ? sp., Loc.1 , JUE no.
15592. 5. Thracia pertrapezoidea Nomura, Loc. 2, JUE no. 15594, x 1.45. 6. Saxidomus sp., Loc. 1, JUE no.
15593. 7. Reticunassa ausiensis (llyina), Loc. 1, JUE no. 15595, x 2.05. 8. Spisula (Pseudocardium) haboroensis(Yokoyama), Loc. 2, JUE no. 15596.

1024.

Miocene Ausinskaya molluscan fauna of Sakhalin

Chikubetsu, Middle Miocene Togeshita and Tachikaraushinai Formations in Hokkaido; Late Miocene? "Otogawa
Formation" and Early Pliocene Joshita Formation in Honshu.
Class Gastropoda
Family Potamididae
Genus Tateiwaia Makiyama, 1936

Tateiwaia sp.
Figures 5-3a, b

Description.-Five specimens from Loc. 1 were examined.
Shell small in size and turreted. Post-nucleus whorls sevenin number; spire elevated and narrowly conical. Surface
ornamented with three spiral threads and 14 axial ribs; upperspiral thread rather strong on shoulder, making weak nodes
where it crosses each axial rib; middle spiral thread alsostrong; lowermost spiral thread weak and situated justabove suture. Gently sloping area present above shoulder.
Aperture poorly preserved.
Measurements (in mm).Specimens
Height
JUE no. 15581
15.6

Diameter
7.9

Remarks.- The Ausinskaya species resembles Tateiwaia
yamanarii (Makiyama) in having three spiral threads. However, T. yamanarii has less numerous (8-9) axial ribs andhigher whorls than the present species.
Tateiwaia yamanarii (Makiyama) and T. tateiwai (Makiyama)
are characteristic elements of the subtropical intertidal
association of early Middle Miocene age (Chinzei, 1986).
According to Uozumi (1962), both species were previously
known only from the early Middle Miocene Takinoue Stage
in southwestern and central Hokkaido. Therefore, thepresent specimens record the northernmost occurrence of
this warm-water genus.

Acknowledgments
We are grateful to Dr. Louie Marincovich, Jr., Emeritus Prof.
Koichiro Masuda of the Miyagi University of Education, andProf. Kenshiro Ogasawara of the University of Tsukuba for
their critical reading of the manuscript. We thank Prof.
Albert G. Ablaev of the Pacific Oceanology Institute for giving
us some useful information on the paleoclimatology of
Sakhalin. Emeritus Prof. Koichiro Masuda, Dr. Ryuichi Majima of the Yokohama National University, and Mr. Hirokazu
Takahashi of the Attached Junior High School of the University of Tsukuba at Komaba provided useful advice on thetaxonomy of pectin ids, naticids and potamidids, respectively.

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© by the Palaeontological Society of Japan 1996

SHORT NOTES

Archaeodictyomitra apiarium (Rust), the correct
specific name of A. apiaria, A. apiara andA. apiarum
HISASHI SUZUKI
Department of Geology and Mineralogy, Faculty of Science, Kyoto University, Kyoto, 606-01 Japan

Received 15 May 1996; Revised manuscript accepted 16 August 1996

The radiolarian species Archaeodictyomitra apiarium (Rust)
has occasionally occurred from the Upper Jurassic andLower Cretaceous in the various areas of the world, forexample, Japan, southern Tibet, Alaska, California, southern
Alps and deep sea of the Pacific (e.g., Nakaseko andNishimura, 1981; Wu, 1993; Pessagno, 1977; Kocher, 1981 ;Matsuoka, 1992). Although this species is an important
index fossil of Upper Jurassic and Lower Cretaceous pelagic
strata, its name has been variously rendered as Archaeodictyomitra apiaria, A. apiara, A. apiarium, and A. apiarum (e.g., Baumgartner, 1987; Pessagno, 1977; Matsuoka
and Yao, 1985; Nakaseko et al., 1979 and so on).
Archaeodictyomitra apiarium was first described as Lithocampe apiarium by Rust (1885). His original description of
Lithocampe apiarium is as follows: "Die Schale mit 8 bis 10
Gliedern hat die Form eines langen Bienenkorbes. Die
mittleren Glieder mit 3, die oberen und unteren mit 2 Querreihen von L6chern."
Rust mentioned the derivatio nominis of this species in his
description as "Bienenkorb", namely beehive. The Latin word
"apiarium" is a neuter noun in the nominative singular, not ahadjective, so that inflection is not necessary in case ofchange of generic name. Article 31 b(ii) of the International
Code of Zoological Nomenclature (Ride et aI. eds., 1985)
explains this point as follows: "A species-group name thatis a simple or compound noun (or noun phrase) in apposition
retains the same ending, whatever the gender of the genericname with which it is combined." Consequently, Archaeodictyomitra apiarium is the correct usage following theInternational Code of Zoological Nomenclature.
Acknowledgments.-I would like to express my sincere
thanks to Prof. K. Chinzei of Kyoto University for critical
discussion.

References cited
Baumgartner, P.O., 1987: Age and genesis of Tethyan
Jurassic radiolarites. Eclogae geologicae Helvetiae,
vol. 80, p.831-879.
Kocher! R.N., 1981: Biochronostratigraphische Untersuchun-gen oberjurassischer radiolarienfUhrender Gesteine, insbesondere der SOdalpen. Mitteilungen aus dem
geologischen Institut der Eidg. Technischen Hochschule und der Universitat Zurich, Neue Folge, Nr.234,
p.1-184, pis. 12-17.
Matsuoka, A., 1992: Jurassic and Early Cretaceous radiolarians from Leg 129, Sites 800 and 801, western Pacificocean. In, Larson, RL, Lancelot, Y. et al. eds., Proceedings of the Ocean Drilling Program, Scientific
Results, vol. 129, p.203-220.
Matsuoka, A. and Yao, A., 1985: Latest Jurassic radiolarians
from the Torinosu Group in Southwest Japan. Journalof Geosciences, Osaka City University, vol. 28, p. 125145, pis. 1-3.
Nakaseko, K. and Nishimura, A., 1981: Upper Jurassic andCretaceous Radiolaria from the Shimanto Group inSouthwest Japan. Science Reports, College of General Education, Osaka University, vol. 30, p. 133-203, pis.
1-17.
Nakaseko, K., Nishimura, A. and Sugano, K., 1979: Cretaceous Radiolaria in the Shimanto Belts, Japan. Newsof Osaka Micropaleontologists, Special Vol., no. 2, p.149, pis. 1-8. (in Japanese)
Pessagno, E.A. Jr., 1977: Lower Cretaceous radiolarianbiostratigraphy of the Great Valley sequence and Franciscan complex, California coast ranges. Cushman
Foundation for Foraminiferal Research, Special Publication, no. 15, 63 p., pis. 1-12.
Ride, W.DL, Sabrosky, C.w., Bernardi, G. and Melville, R.V.
eds., 1985: International Code of Zoological Nomenclature Adopted by the XX General Assembly of theInternational Union of Biological Sciences. 338 p.
International Trust for Zoological Nomenclature, London.
ROst, D., 1885: Beitrage zur Kenntniss der fossilen Radiolarien aus Gesteinen des Jura. Palaeontographica, Bd. 31,
p. 273-321, pis. 26-45.
Wu, H., 1993: Upper Jurassic and Lower Cretaceousradiolarians of Xialu chert, Yarlung Zangbo ophiolite
belt, southern Tibet. In, Blueford, J.R. and Murchey, B.
eds., Radiolaria of Giant and Subgiant Fields in Asia, p.
115-136, pis. 1-4.

663

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664
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Ager, D.V., 1963: Principles of Paleoecology, 371 p. Magrwhill Co., New York.
Barron, J.A., 1983: Latest Oligocene through early middle
Miocene diatom biostratigraphy of the eastern tropical
Pacific. Marine Micropaleontology, vol. 7, p. 487-515.
Barron, J.A., 1989: Lower Miocene to Quaternary diatom
biostratigraphy of Leg 57, off northeastern Japan, DeepSea Drilling Project. In, Scientific Party, Initial Reportsof the Deep Sea Drilling Project, vols. 56 and 57, p. 641-

685. U.S. Govt. Printing Office, Washington, D.C.
Burckle, L.H., 1978: Marine diatoms. In, Haq, B.U. andBoersma, A. eds., Introduction to Marine Micropaleontology, p.245-266. Elsevier, New York.
Fenner, J. and Mikkelsen, N., 1990: Eocene-Oligocene
diatoms in the western Indian Ocean: Taxonomy,
stratigraphy, and paleoecology. In, Duncan, A.A.,
Backman, J., Peterson, L.C., et al., Proceedings of the
Ocean Drilling Program, Scientific Results, vol. 115, p.
433-463. College Station, TX (Ocean Drilling Program).
Koizumi, 1.,1972: Marine diatom flora of the Pliocene Tatsunokuchi Formation in Fukushima Prefecture. Transactions and Proceedings of the Palaeontological Societyof Japan, New Series, no. 86, p. 340-359.

Kuramoto, S., 1996: Geophysical investigation for methane
hydrates and the Significance of BSA. Journal of theGeological Society of Japan, vol. 11, p.951-958. (inJapanese with English abstract)

665

SYSTEMATIC INDEX(New Series No. 177-No. 184)

PALAEOZOOLOGY
Protoctista
Page
Funakawa, Satoshi: Intrageneric variation and temporal change in the internal skeletal structure of plagiacanthids(Radiolaria) from Hokkaido, Japan .............................................................. ..
Hasegawa, Shiro and Nomura, Ritsuo: Redescription of Asano and Nakamura's (1937) cassidulinid species based on
their primary types ................................................................... , ......... .
Igo, Hisaharu: Permian fusulinaceans from the Akuda and Horikoshitoge Formations, Hachiman town, Gifu Prefecture,
Central Japan " .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. " .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
Kitazato, Hiroshi and Matsushita, Satoshi: Laboratory observations of sexual and asexual reproduction of Trochammina
hadai Uchio ................................................................................... .
Kobayashi, Fumio: Middle Triassic (Anisian) foraminifers from the Kaizawa Formation, southern Kanto Mountains,
Japan ...................................................................................... ..
Kondo, Yasuo: Density and ralative abundance of microspheric and megalospheric forms of Nummulites assedimentologic and taphonomic indicators ...................................................... ..
Sash ida, Katsuo, Adachi, Shuko, Igo, Hisayoshi, Koike, Toshio and Amnan, Ibrahim B.: Middle and Late Permian
radiolarians from the Semanggol Formation, northwest Peninsular Malaysia ........................... .
Scott, David B., Hasegawa, Shiro, Saito, Tsunemasa, Ito, Koji and Collins, Eric: Marsh foraminiferal and vegetation
distributions in Nemuro Bay wetland areas, eastern Hokkaido ...................................... ..
Suzuki, Hisashi: Reexamination of the age of the Canoptum assemblage (Radiolaria) from acidic tuff in the Hiuchiwan
Unit of the Tanba Group, Southwest Japan ...................................................... ..
Suzuki, Hisashi: Archaeodictyomitra apiarium (Rust), the correct specific name of A. apiaria, A. apiara and A. apiarum

208
89
623
454

528
59
43

282
196

662
Yanagisawa, Yukio: Cenozoic diatom genus Rossiella Desikachary et Maheshwari: An emended description ..... .
Yanagisawa, Yukio: Cenozoic diatom genus Bogorovia Jouse: An emended description ....................... .
Zaw, Win and Sakagami, Sumio: Preliminary note on the rediscovery of "Neoschwagerina (Sumatrina) multiseptata
Deprat" from Akasaka, Japan .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ., .. .. .. .. ..

1
21

547

Mollusca
Amano, Kazutaka, Ukita, Mikiko and Sato, Shinji: Taxonomy and distribution of the subfamily Ancistrolepidinae(Gastropoda: Buccinidae) from the Plio-Pleistocene of Japan ...................................... ..
Amano, Kazutaka, Khudik, Vladimir D. and Narita, Ken: An early Middle Miocene warm-water molluscan fauna insouthwestern Sakhalin - Molluscs from the Ausinskaya Formation around Novoselovo - ............... .
Ehiro, Masayuki: Cephalopod fauna of the Nakadaira Formation (Lower Permian) in the Southern Kitakami Massif,
Northeast Japan .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
Ehiro, Masayuki: Latest Permian ammonoid Paratirolites from the Ofunato district, Southern Kitakami Massif, Northeast
Japan ...................................................................................... ..
Khim, Boo-keun, Nobuhara, Takami and Krantz, David E.: Stable isotope records of the Pliocene fossil Akebiconcha
kawamurai (Bivalvia: Vesicomyidae) from the Tamari Formation, Central Japan: Indication of chemoautotrophic symbiosis ................................................................................ ..
Kondo, Yasuo and Kikuchi, Naoki: "Shelly softground" colonized by an endobyssate bivalve, Modiolus kurilensis: an
example of taphonomic feedback from the Pleistocene of Kanazawa, Japan ......................... .
Matsubara, Takashi: Fossil Mollusca of the Lower Miocene Yotsuyaku Formation in the Ninohe district, Iwate
Prefecture, Northeast Japan, Part 1. General consideration of the fauna ............................ ..
Matsubara, Takashi: Fossil Mollusca of the Lower Miocene Yotsuyaku Formation in the Ninohe district, Iwate
Prefecture, Northeast Japan, Part 2 (1). Bivalvia and Scaphopoda ................................... .
Matsubara, Takashi: Fossil Mollusca of the Lower Miocene Yotsuyaku Formation in the Ninohe district, Iwate
Prefecture, Northeast Japan, Part 2 (2). Gastropoda .............................................. ..
Matsumoto, Tatsuro: A little known species of Inoceramus (Bivalvia) from the Turonian (Cretaceous) of Hokkaido

467
651

184

592

448
296

303
321
361

540

666
Matsumoto, Tatsuro and Asai, Akito: An early Turonian (Cretaceous) new species of Inoceramus (Bivalvia) from
Hokkaido ...... " ................................................... , .. .. .. .. .. .. .. .. .. .. .. .. ..
Niko, Shuji: Pseudorthoceratid cephalopods from the Early Devonian Fukuji Formation of Gifu Prefecture, Central
Japan ........................................................................................
Niko, Shuji, Nishida, Tamio and Kyuma, Yuko: A new Carboniferous cephalopod Bogos/ovskya akiyoshiensis fromSouthwest Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
Niko, Shuji, Pillevuit, Alain and Nishida, Tamio: Early Late Permian (Wordian) non-ammonoid cephalopods from the
Hamrat Duru Group, central Oman Mountains .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....
Niko, Shuji, Pillevuit, Alain and Nishida, Tamio: Triassic aulacocerid (Mollusca: Cephalopoda) from the central Oman
Mountains .....................................................................................
Nishinosono, Shin'ichi: Larval paleoecology of the late Pleistocene naticid gastropod Cryptonatica with two protoconch
types from the Oga Peninsula, northern Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....
Noda, Masayuki: Five inoceramids (Bivalvia) from the Upper Cretaceous of Hokkaido with some phylogenetic andtaxonomic considerations, Part 1. Introductory remarks, method and systematic descriptions of one speciesof Inoceramus (I.) and one of I. (Volviceramus) .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....
Noda, Masayuki: Five inoceramids (Bivalvia) from the Upper Cretaceous of Hokkaido with some phylogenetic andtaxonomic considerations, Part 2. Systematic descriptions of three species of Inoceramus (Cremnoceramus)
and concluding remarks.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....
Noda, Masayuki and Uchida, Shigehiro: Inoceramus (Platyceramus) szaszi sp. nov. (Bivalvia) from the Coniacian(Cretaceous) of Hokkaido .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....
Ozawa, Tomowo, Inoue, Keisuke and Tomida, Susumu: Early to Middle Miocene molluscan fossils from the Megami
and Ebie limestones in Shizuoka Prefecture and their implications for the formation of a Miocene coral-algal
reef within the Japanese faunal realm ............................................................
Sarashina, lsao: Genetic variation within a dimorphic species, Cryptopecten vesiculosus (Bivalvia, Pectinidae) .. .. ..
Takayasu, Katsumi, Gurung, Damayanti D. and Matsuoka, Keiji: Some new species of freshwater bivalves from the
Mio-Pliocene Churia Group, west-central Nepal.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....

375
347
193
522
544
419

555

571
142

175
203
157

Brachiopoda
Saito, Michiko: Early loop ontogeny of some Recent laqueid brachiopods.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

485

Arthropoda
Irizuki, Toshiaki and Matsubara, Takashi: Early Middle Miocene ostracodes from the Suenomatsuyama Formation,
Ninohe City, northeast Japan and their paleoenvironmental significance .............. " .. .. .. .. .. .. .. ..
Karasawa, Hiroaki: Shako, a new Miocene stomatopod Crustacea from Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ....
Kato, Hisayoshi: Miocene decapod crustacea from the Chichibu Basin, Central Japan .... " .. .. .. .. .. .. .. .. .. ..

65
413
500

Conodonta
Igo, Hisaharu: On the occurrence of Upper Paleozoic condonts from central Spitsbergen .. .. .. .. .. .. .. .. .. .. .. ..
Koike, Toshio: The first occurrence of Griesbachian conodonts in Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

79337

Bryozoa
Sakagami, Sumio: Upper Paleozoic bryozoans from the Lake Titicaca region, Bolivia, Part 1. Introductory remarks,
stratigraphy and systematic paleontology .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .... 226
Sakagami, Sumio: Upper Paleozoic bryozoans from the Lake Titicaca region, Bolivia, Part 2. Systematic paleontology.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .... 261

Vertebrata
Hirayama, Ren and Chitoku, Tsutomu: Family Dermochelyidae (Superfamily Chelonioidea) from the Upper Cretaceousof North Japan ............................................................................... .
Jin, Chang-zhu and Kawamura, Yoshinari: The first reliable record of Beremendia (insectivora, Mammalia) in East Asiaand a revision of Peisorex Kowalski and Li, 1963 ..................................................
Jin, Chang-zhu and Kawamura, Yoshinari: A new genus of shrew from the Pliocene of Yinan, Shandong Province,
northern China ............................................................................... .
Kohno, Naoki: Miocene pinniped Allodesmus (Mammalia: Carnivora); with special reference to the "Mito seal" from

597
432
478

667
Ibaraki Prefecture, Central Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
Kiso, Tarou M.: Organic components in enameloid of extant and fossil shark teeth.. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

388
169

Palaeoenvironment
Xu, Xuedong and Oda, Motoyoshi: Surface water changes in Ryukyu Trench slope region, western margin of the North
Pacific during the last 320,000 years .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

105

PALAEOBOTANY
Palaeobotany
Ohana, Tamiko and Kimura, Tatsuaki: Further observations of Cunninghamiostrobus yubariensis Stopes and Fujii fromthe Upper Yezo Group (Upper Cretaceous), Hokkaido, Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..

122

668

INDEX OF GENERA AND SPECIES
Notes: Words listed are names of genus and species-groups which are described, discussed or illustrated in. the articles
in Transactions and Proceedings of Palaeontological Society of Japan, New Series, No. 177 to No. 184. Words In bold-type
are names of newly proposed taxa.

A
Aeervosehwagerina ................................ 645
Aeervosehwagerina fujimotoi .................. 645, fig. 10
Adamnestia sp. indet. ........................ 367, fig. 2
Agathieeras sp............................. 190, figs. 5, 6
Albaillella exeelsa ............................ 52, fig. 10
Albaillella levis .. .............................. 51, fig. 10
Albaillella triangularis .. ........................ 51, fig. 10
Allodesminae gen. et sp. indet. ............ 398, figs. 5, 6
Allodesminae gen. et sp. indet. ............... , 399, fig. 7
Allodesminae gen. et sp. indet. ................ 399, fig. 7
Allodesmus ...................................... 389
Allodesmus kernensis ............................ 398
Allodesmus naorai .......................... 390, fig. 1
Allodesmus sinanoensis .. .......................... 397
Alternifenestella minor .. ...................... 265, fig. 3
Alternifrnestella aft. A. pajerensis .............. 265, fig. 3
Alternifenestella (Cervella) aspera .............. 266, fig. 3
Alternifenestella (Cervella) eervoides ........... , 266, fig. 3
Alternifenestella? sp. Indet. .................. 268, fig. 4
Ammobaeulites exiguus ..................... , 292, fig. 6
Aneistrolepis grammatus ...................... 471, fig. 3
Aneistrolepis masudaensis .................... 470, fig. 2
Aneistrolepis sp. aft. A. hikidai ................ 472, fig. 2
Aneistrolepis sp. . ........................... 472, fig. 3
Anomia ehinesis .. ............................ 322, fig. 1
Aphelaeceras cf. faleiferum .................... 188, fig. 4
Arehaeodietyomitra apiarium .. .. .. .. .. .. .. .. .. .. .. .. 662
Arenovidalina amylovoluta .................... 537, fig. 5
Armenina ........................................ 646
Armenina sp. .. ............................ 646, fig. 10
Artinskia sp. .. .............................. 190, fig. 6
Atractites arabicus .......................... 544, fig. 1
Austrocolomia cf. marschalli .................. 537, fig. 5

B
Baetrites ........................................ 526
Baetrites? sp. . ............................. 526, fig. 3
Beremendia ...................................... 434
Beremendia pohaiensis .. .................. 435, figs. 3-12
Bitaunioceras .................................... 523
Bitaunioceras cf. zonatum .................... 523, fig. 3
Bogorovia ........................................ 25
Bogorovia barronii.. .. .. .. .. .. .. .. .. .. .. .. .. 31, figs. 4, 5
Bogorovia curvata ............................ 37, fig.4
Bogorovia gombosii ........................ 27, figs. 4, 5
Bogorovia lancettula ........................ .. 34, fig. 4
Bogorovia praepaleacea ........................ 36, fig. 4

Bogorovia puncticulata ....... , .......... 32, figs. 4, 12, 13
Bogorovia rostrata ............................ 34, fig. 4
Bogorovia veniamini ........................ 29, figs. 4, 8
"Bogorovia" sparsipunctata .......................... 38
Bogoslovskya .................................... 193
Bogoslovskya akiyoshiensis .................. 193, fig. 1
Boreotrophon sp. indet. ...................... 367, fig. 2
Braehycyc/oceras .................................. 522
Brachycycloceras rustaqense ................ 522, fig. 3
Buchanoeeras sp............................. 358, fig. 8
Burseolina pacifica .. .......................... 102, fig. 4C
Callianopsis ...................................... 507
Callianopsis sp. 1 ............................ 507, fig.4
Callianopsis sp. 2 ............................ 507, fig.4
Cancer ................................. , ........ 510
Cancer sakamotoi .......................... 510, fig. 5
Cancer sp. .. ................................ 512, fig. 5
Cantharidus sp. indet. ........................ 361, fig. 1
Carcinoplax ...................................... 516
Carcinoplax antiqua .......................... 516, fig. 7
Carcinoplax sp. .. ............................ 516, fig. 7
Centropyxis aeuleata ........................ 292, fig. 6
Cerithideopsilla aft. minoensis .................. 364, fig. 1
Chalaroschwagerina .............................. 645
Chalaroschwagerina aft. chinensis .............. 645, fig. 8
Chlamys (Chlamys) arakawai .................. 655, fig. 4
Chlamys (Chlamys) otukae .................... 655, fig. 4
Chlamys sp. A .............................. 180, fig. 3
Chlamys sp. B .............................. 180, fig. 3
Clinocardium sp. indet. ........................ 326, fig. 1
Clinopegma borealis .. ........................ 472, fig. 2
Coptothyris grayi ............................ 490, fig. 4
Corythomelissa .................................... 211
Corythomelissa omoprominentia .............. 212, fig. 5
Corythomelissa pachyostraca ................ 214, fig. 6
Corythomelissa spinosa ...................... 214, fig. 7
Corythomelissa sp............................. 216, fig. 7
Crassostrea gigas .. ........................ 324, figs. 1, 4
Crepidula jimboana .......................... 363, fig. 1
Cryptopeeten .................................... 178
Cryptopecten yanagawaensis .................. 178, fig. 3
Cultellus izumoensis izumoensis .. .... " ........ 328, fig. 1
Cunninghamiostrobus .............................. 140
Cunninghamiostrobus yubariensis .......... 140, figs.1-13A
Cyelina japonica .. ............................ 334, fig. 5
Cymatosira lorenziana .............................. 38
Cymatosira sp. A .................................. 38

669"Cypraea" sp................................. 181, fig. 3

D
Oiplotremina astrofimbriata .................... 538, fig. 5
Oosinia (Kaneharaia) ausiensis .. ................ 657, fig. 4
Oosinia (Phacosoma) kawagensis .............. 330, fig. 4
Oosinia (Phacosoma) tugaruana ................ 657, fig. 4
Ounbarula ........................................ 629
Dunbarula uenoi ............................ 629, fig. 11
Oyscritel/a aft. O. komukensis ................ 258, fig. 18
Oyscritel/a tenuirama ........................ 256, fig. 18
E"Endothyra" cf. kuepperi ...................... 536, fig.4
Entactinosphaera pseudocimelia ................ 53, fig. 11
Entactinosphaera sp. . ......................... 53, fig. 11
Eucal/iax .................................... 507, fig. 4
Euscenarium ...................................... 216
Euscenarium microcapitalium ................ 216, fig. 10
FFabifenestel/a aft. F. huascatayana .............. 268, fig. 4
Fabifenestel/a sp. indet. .................... 270, figs. 3, 4
Felaniel/a ferruginata ...................... 326, figs. 1, 3
Fistulipora carrascoi .................... 242, figs. 10, 11
Fistulipora incrustans ........................ 242, fig. 9
Fistulipora titicacaensis ...................... 244, fig. 11
Fistulipora cf. F. timorensis .................... 242, fig. 9
Fistulipora? multidiaphragma ................ 244, fig. 11
Fistulipora sp. indet. A ........................ 244, fig. 9
Fistulipora sp. indet. B ........................ 244, fig. 9
Fistulipora sp. indet. C ........................ 246, fig. 9
Fistulipora sp. indet. D ...................... 246, fig. 12
Fol/icucul/us monacanthus .................... 53, fig. 10
Foordiceras sp............................. 186, figs. 3, 4
Foordiceras? sp........................... 187, figs. 3, 4
Frenulina sanguinolenta ...................... 492, fig. 5'Frenulina' sp............. " .................. 492; fig. 6
Fukujiceras ...................................... 348
Fukujiceras kamiyai ........................ 348, fig. 2

Hamadaites labyrinthus ...................... 354, fig. 6
Haplophragmoides manilaensis ................ 292, fig. 6
Hiatula minoensis ............................ 327, fig. 1
Hindeodus minutus ........... , .............. 342, fig. 3
Hindeodus parvus ............. , .............. 342, fig. 3
Homa/opoma aft. ena .. ........................ 361, fig. 1

Idiognathodus magnificus ...................... 80, fig. 2
Indonaia .. ........................................ 162
Indonaia churia .............................. 162, fig.4
Indonaia jimuriensis .......................... 164, fig.4
Indonaia narayani .. .......................... 162, fig. 4
Indonaia tenella .. ............................ 164, fig.4
Inoceramus ...................................... 376
Inoceramus ...................................... 558
Inocerams .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 571
Inoceramus kamuy .. ...................... 376, figs. 1-5
Inoceramus aft. I. tuberosus .................... 540, fig. 1
Inoceramus (Cremnoceramus) ...................... 571
Inoceramus (Cremnoceramus) deformis ...... 572, figs. 1, 7
Inoceramus (Cremnoceramus) ernsti ........ 576, figs. 5-7
Inoceramus (Cremnoceramus) leuckendorfensis .. 585, fig, 11Inoceramus (Inoceramus) lusatiae .. .......... 558, figs. 2, 3
Inoceramus (Platyceramus) " ........................ 144
Inoceramus (Platyceramus) szaszi ........ 144, figs. 2, 3, 5
Inoceramus (Volviceramus) ........................ 564
Inoceramus (Volviceramus) koeneni ............ 566, fig. 10
Isarcicel/a isarcica .. .. .. .. .. .. .. .. .. .. .. .. .. .. 344, fig. 3
Ishigaum ? sp................................. 55, fig. 11
Is/andiel/a japonica ............................ 91, fig. 1
Is/andiel/a kazusaensis ........................ 92, fig. 3
Is/andiel/a setanaensis .. ..................... , .. 92, fig. 1
Is/andiel/a sublimbata .......................... 94, fig, 2Is/andiel/a wakasaensis ........................ 94, fig. 1
Is/andiel/a yabei .............................. 96, fig. 2
Itoigawaia ................................. , ...... 512/toigawaia chichibuensis .................. 512, figs. 5, 6

J
Jolonica nipponica ................................ 492

GL

G/obocassidulina brocha.. .. .. .. .. .. .. .. .. .. .. .. 98, fig. 3
G/obocassidulina decorata .. .. .. .. .. .. .. .. .. .. .. 98, fig. 4
G/obocassidulina depressa .................... 100, fig. 4
G/obocassidulina elegans ...................... 101, fig. 4
G/obocassidulina gemma ...................... 101, fig. 4
G/omospira densa ............................ 534, fig. 4
G/omospirel/a irregularis ...................... 536, fig. 4
Goniocladia peruviana ........................ 250, fig. 14

Laevidentalium? sp. indet. .................... 335, fig. 5
Lamel/idens ...................................... 158
Lamellidens arungensis ...................... 158, fig. 3
Lamellidens longiformis ...................... 160, fig. 3
Laperousecythere ishizakii .................... 73, fig. 6
Laperousecythere cf. robusta .................. 73, fig. 6
Laperousecythere yahtsensis .................. 76, fig. 7
Laperousecythere sp........................... 76, fig. 7
Laqueus rubel/us ................................ 486
Laqueus sp. ................................ 488, fig. 1
Liroceras ........................................ 524
Liroceras sp. . ............................... 524, fig. 3
Lithomelissa ...................................... 217

Genus and species indeterminate .............. 350, fig, 3

H

Hamadaites ...................................... 352

670
Lithomelissa trifoliolata ...................... 217, fig. 10
Littorina sp. indet. ............................ 361, fig, 1
Lueinoma ct. aeutilineata ...................... 325, fig. 1
Lunanosorex .................................... 478
Lunanosorex Iii .......................... 479, figs. 2-6

M
Macoma (Maeoma) incongrua .................. 326, fig. 1
Maja ............................................ 518
Maja mor;; .................................. 518, fig. 8
Maklaya .. ........................................ 645
Mak/aya pamirica ............................ 646, fig. 10
Maklaya saraburiensis ........................ 646, fig. 10
Ma/ayspirina cf. fontainei ...................... 536, fig. 4
Meandrospira dinarica ........................ 537, fig. 5"Meandrospira" deformata .................... 537, fig. 5
Medaeus ........................................ 514
Medaeus? sp. . ............................. 516, fig. 7
Meekopora prosseri .. ........................ 246, fig. 13
Meekopora sp. Indet. .................... 250, figs. 12, 13
Meretrix arugai .............................. 333, fig. 5
Mesoc/ermochelys ................................ 598
Mesoc/ermochelys undulatus .............. 599, figs. 2-17
Miliammina fusca .. ................................ 292
Minilya binodata ............................ 268, fig. 3
Minohellenus .................................... 514
Minohellenus? sp............................. 514, fig. 6
Minojapanella .................................... 629
Minojapanella (M.) elongata .................. " 629, fig. 11
Miosesarma ...................................... 518
Miosesarma nagurense ...................... 518, fig. 8
Mitorthoceras? kamitakarense " .............. 355, fig. 7
Mizuhopeetenaff. ehiehibuensis mitsuganoensis .. 321, fig. 1
Munida .. ........................................ 508
Munida sp. cf. M. nishioi ...................... 508, fig. 4
Mursia .......................................... 508
Mursia takahashii ............................ 508, fig. 4
Mya sp. indet. .............................. 334, fig. 5
N

Nanaoehlamys sp............................. 657, fig. 4
Nankinella .. ...................................... 627
Nankinella aft. kotakiensis .................... 627, fig. 11
Nankinella sp. .. ............................ 627, fig. 11
Nassarius (Zeuxis) sp. indet. .................. 367, fig. 2
Nazarovella gracilis .. .......................... 53, fig. 11
Nazarovella inflata ............................ 55, fig. 11
Nebela eollaris .............................. 292, fig. 6
Neoalbai/lella ct. ornithoformis .................. 50, fig. 10
Neofusulinella .................................... 628
Neofusu/inella giraudi ........................ 628, fig. 11
Neofusulinella /antenoisi ...................... 629, fig. 11
Neostreptognathodus sva/bardensis .. .. .. .. .. .. .. 82, fig. 1
Nipponomareia nakamurai ...... " .............. 331, fig. 4o"Ocenebra" katayamai ...................... 367, fig. 2
Oetatormentum ? sp........................... 53, fig. 11
Pamirina (Levenella) evo/uta .................. 627, fig. 11
Pamirina (Levenella) /eveni .................... 627, fig. 11
Paracassidulina nipponensis .............. " .... 96, fig. 3
Paracassidulina sagamiensis .. .. .. .. .. .. .. " .. .. 98, fig. 3
Parafusulina ...................................... 631
Parafusulina ? sublineata .... " ................ 633, fig. 9
Parafusulina sotensis ........................ 633, fig. 10
Parastenopoeeras sp....................... 188, figs. 3, 6
Paratirolites .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 593
Paratirolites compressus .. .. " ................ 593, fig. 3
Paratirolites sp. .. ............................ 594, fig. 3
Parreysia ........................................ 165
Parreysia binaiensis .......................... 165, fig. 5
Parreysia zigzagicostata ...................... 165, fig. 5
Periglypta sp. cf. P. retieulata .................. 180, fig. 3
Phos (Phos) notoensis .. ...................... 366, fig. 2
Physunio ........................................ 166
Physunio chitwanensis .. ...................... 166, fig. 5
Pietothyris pieta .......................... 488, figs. 1, 2
Pietothyris sp. . ........................... 488, figs. 2, 3
Pillueina (Wallueina) cf. striata .. .............. 324, figs. 1, 2
Pleuronautilus yahagiensis ................ 187, figs. 3, 4
Pleuronautilus sp. . ........................... 188, fig. 4
Polypora eyelopora ........................ 272, figs. 5, 6
Polypora elliptica .. .. .. .. .. .. .. .. .. .. .. .. .. 270, figs.4, 5Polypora aft. P. inimica ........................ 272, fig. 6
Polypora cf. P. megastoma .................... 272, fig. 5
Polypora sp. indet. A ........................ 274, fig. 6
Polypora sp. indet. B ........................ 274, fig. 6
Polypora sp. indet. C ........................ 274, fig. 7
Praedeflandrella sp. .. ........................ 55, fig. 11
Pseudobatostomella micropora .............. 258, fig. 19
Pseudobatostomella yanagidai .. .............. 260, fig. 19
Pseudobatostomella sp. indet. ................ 260, fig. 19
Psedofusulina .................................... 633
Pseudofusulina duplithecata .................. 637, fig. 6
Pseudofusulina fusiformis .. .................... 641, fig. 8
Pseudofusulina horadaniensis .. .. .. .. .. .. .. .. .. 635, fig. 5
Pseudofusulina houziguan .................... 643, fig. 8
Pseudofusulina isomie ........................ 637, fig. 6
Pseudofusulina kraffti ........................ 637, fig. 6
Pseudofusulina krotowi ...................... 645, fig. 8
Pseudofusulina minoensis .................... 639, fig. 7
Pseudofusulina miyamensis .................. 641, fig. 7
Pseudofusulina norikurensis .................. 633, fig. 5
Pseudofusulina paratsehernysehewi ............ 635, fig. 5
Pseudofusulina regu/aris ...................... 643, fig. 8
Pseudofusulina sp. .. ........................ 645, fig. 7
Pseudoreiehelina sp. .. ...................... 626, fig. 10
Pseudothurammina limnetis .. .................. 292, fig. 6
Pseudotormentus sp........................... 55, fig. 11
R
Rhombopora eorticata ...................... 262, figs. 1, 2

671

Rhombopora kawabei ........................ 262, fig. 2
Rhombopora /epidodendroides ................ 261, fig. 1

T

Rhombotrypella typica .................... 252, figs. 15, 16
Rhombotrypella aft. R. gigantea ................ 254, fig. 17
Rhombotrypella? sp. indet. .................. 254, fig. 15
Rossiella .......................................... 4
Rossiella e/ongata .. ........................ 13, figs. 4, 15
Rossie//a fennerae .......................... 8, figs. 4, 7
Rossie//a fourtanierae ...................... 9, figs. 4, 9
Rossiella pa/eacea .. ........................ 11, figs. 4, 12
Rossiella symmetrica .. .......................... 5, fig. 4
Ruditapes miyamurensis ...................... 331, fig. 5

Tabulipora cf. T. carbonaria .................. 254, fig. 17
Tabu/ipora sp. indet. ........................ 256, fig. 18
Tainoceras sp............................. 186, figs. 3, 4
Tateiwaia chinzeii .. .......................... 363, fig. 1
Tateiwaia sp................................. 659, fig. 5
Terebratalia coreanica .. .................... 490, figs. 3, 4
Toriyamaia ...................................... 629
Toriyamaia laxiseptata .. ...................... 629, fig. 10
Trachycarcinus .. .................................. 508
Trachycarcinus inf/atus ...................... 510, fig. 5
Trapezium (Neotrapezium) ichinohense ........ 328, fig. 4
Trapezium (Neotrapezium) modiolaeforme .. ...... 330, fig.4
Triplanospongos .................................. 55
Trip/anospongos musashiensis .................. 55, fig. 11
Trochammina inflata .......................... 294, fig. 6
Trochammina macrescens .................... 294, fig. 6
Turbo (Marmarostoma) mekamiensis ............ 181, fig. 3
Typh/ocarcinodes .. ................................ 518
Typh/ocarcinodes? sp ......................... 518, fig. 7sSchubertella.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 628
Schubertella pseudoobscura .................. 628, fig. 11
Schwagerina .. .................................... 630
Schwagerina higashidaniensis .................. 631, fig. 8
Schwagerina kanumai .. ...................... 631, fig. 9
Schwagerina kwangchiensis .. .................. 631, fig. 8
Schwagerina muraii .......................... 630, fig. 5
Schwagerina shukoae .. ...................... 630, fig. 9
Sear/esia kurodai .. .. .. .. .. .. .. .. .. .. .. .. .. .. 366, fig. 2
Septopora andeana .. ........................ 276, fig. 8
Septopora incaica ............................ 276, fig. 8
Septopora lineata ........................ 276, figs. 7, 8
Shako .. ........................................ 413
Shako tomidai .............................. 415, fig. 1
So/en sp. indet. .............................. 327, fig. 1
Sphaerulina ...................................... 627
Sphaeru/ina akudensis ...................... 627, fig. 11
Spondy/us sp................................. 180, fig. 3
Spyroceras fukujiense .................... 350, figs. 3, 4
Spyroceras me/olineatum .. ................ 352, figs. 4, 5
Stearoceras sp............................. 190, figs. 3, 6
Stenodiscus a/tip/ana .. ...................... 250, fig. 15
Streb/atrypa sp. indet. ........................ 262, fig. 1uUpogebia ........................................ 506
Upogebia sp................................. 506, fig.4

v
Vicaryella "bacu/a" .. .......................... 364, fig. 1
Vicaryella otukai .. ............................ 366, fig. 1
Vicaryella sp. indet. .......................... 366, fig. 1

yYabeina (Lepidolina) mu/tiseptata .............. 547, fig. 2
Yo/dia (Yo/dia) aft. omorii ...................... 321, fig. 1

Transactions and Proceedingsof thePalaeontological Society of Japan

New SeriesNos.177 -184
1995-1996

Cover: A terminally resorbed maxillary tooth of iguanodontids (Ornithischia: Ornithopoda). The specimen isfrom the Kuwajima Formation, the Neocomian section of the Tetori Group, in Shiramine-mura; Ishikawa
County, Ishikawa Prefecture, the west coast of Central Honshu, Japan (Shiramine-mura Board of
Education, Coil. Cat. No. SBEI 001, 23 mm in dorsoventral length).

CONTENTS

Number 177 (published April 30, 1995)

TRANSACTIONS
982.
983.
984.
985.
986.

Page Figures

Yukio Yanagisawa: Cenozoic diatom genus Rossiella Desikachary et Maheshwari: An emended
description .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
Yukio Yanagisawa: Cenozoic diatom genus Bogorovia Jouse: An emended description .. .. .. .. ..
Katsuo Sash ida, Shuko Adachi, Hisayoshi Igo, Toshio Koike, and Ibrahim B. Amnan: Middle and Late
Permian radiolarians from the Semanggol Formation, northwest Peninsular Malaysia .. .. .. .. .. .. ..
Yasuo Kondo: Density and relative abundance of microspheric and megalospheric forms of Nummulites as sedimentologic and taphonomic indicators ........................................
Toshiaki Irizuki and Takashi Matsubara: Early Middle Miocene ostracodes from the Suenomatsuyama
Formation, Ninohe City, northeast Japan and their paleoenvironmental significance ............. .

1-20
21-42

1-16
1-18

43-58

1-11

59-64

1- 4

65-78

1- 7

79-83

1- 2

SHORT NOTES
31.

Hisaharu Igo: On the occurrence of Upper Paleozoic conodonts from central Spitsbergen

PROCEEDINGS.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 84-87

Number 178 (published June 30, 1995)
TRANSACTIONS
987. Shiro Hasegawa and Ritsuo Nomura: Redescription of Asano and Nakamura's (1937) cassidulinid
species based on their primary types .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 89-1 04
988. Xuedong Xu and Motoyoshi Oda: Surface water changes in Ryukyu Trench slope region, western
margin of the North Pacific during the last 320,000 years .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 105-121
989. Tamiko Ohana and Tatsuaki Kimura: Further observations of Cunninghamiostrobus yubariensis Stopes
and Fujii from the Upper Yezo Group (Upper Cretaceous), Hokkaido, Japan .................... 122-141
990. Masayuki Noda and Shigehiro Uchida: Inoceramus (Platyceramus) szaszi sp. nov. (Bivalvia) from theConiacian (Cretaceous) of Hokkaido ..................................................... . 142-153

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1- 8
1-13
1- 8

Number 179 (published September 30, 1995)
TRANSACTIONS
991. Katsumi Takayasu, Damayanti D. Gurung and Keiji Matsuoka: Some new species of freshwaterbivalves from tne Mio-Pliocene Churia Group, west-central Nepal ............................ 157-168 1- 5
992. Tarou M. Kiso: Organic components in enameloid of extant and fossil shark teeth .............. 169-174 1- 2
993. Tomowo Ozawa, Keisuke Inoue and Susumu Tomida: Early to Middle Miocene molluscan fossils from
the Megami and Ebie limestones in Shizuoka Prefecture and their implications for the formation of a
Miocene coral-algal reef within the Japanese faunal realm .................................. 175-183 1- 3994. Masayuki Ehiro: Cephalopod fauna of the Nakadaira Formation (Lower Permian) in the Southern
Kitakami Massif, Northeast Japan ....................................................... . 184-192 1- 6

SHORT NOTES
Shuji Niko, Tamio Nishida and Yuko Kyuma: A new Carboniferous cephalopod Bogoslovskya akiyoshiensis from Southwest Japan ......................................................... . 193-195
33. Hisashi Suzuki: Reexamination of the age of the Canoptum assemblage (Radiolaria) from acidic tuff inthe Hiuchiwan Unit of the Tanba Group, Southwest Japan .................................. 196-200 1- 3
32.

PROCEEDINGS .. .............................................................................. 201

Number 180 (published December 30, 1995)
TRANSACTIONS
995. lsao Sarashina: Genetic variation within a dimorphic species, Cryptopecten vesiculosus (Bivalvia,
Pectinidae) ............................................................................ 203-207
996. Satoshi Funakawa: Intrageneric variation and temporal change in the internal skeletal structure ofplagiacanthids (Radiolaria) from Hokkaido, Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 208-225
997. Sumio Sakagami: Upper Paleozoic bryozoans from the Lake Titicaca region, Bolivia, Part 1. Introductory remarks, stratigraphy and systematic paleontology .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 226-260
998. Sumio Sakagami: Upper Paleozoic bryozoans from the Lake Titicaca region, Bolivia, Part 2. Systematic paleontology.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 261-281
999. David B. Scott, Shiro Hasegawa, Tsunemasa Saito, Koji Ito and Eric Collins: Marsh foraminiferal andvegetation distributions in Nemuro Bay wetland areas, eastern Hokkaido ...................... 282-295
1000. Yasuo Kondo and Naoki Kikuchi: "Shelly softground" colonized by an endobyssate bivalve, Modiolus
kurilensis: an example of taphonomic feedback from the Pleistocene of Kanazawa, Japan .. .. .. 296-302
1001. Takashi Matsubara: Fossil Mollusca of the Lower Miocene Yotsuyaku Formation in the Ninohe district,
Iwate Prefecture, Northeast Japan, Part 1. General consideration of the fauna.. .. .. .. .. .. .. .. .. 303-320
1002. Takashi Matsubara: Fossil Mollusca of the Lower Miocene Yotsuyaku Formation in the Ninohe district,
Iwate Prefecture, Northeast Japan, Part 2 (1). Bivalvia and Scaphopoda.. .. .. .. .. .. .. .. .. .. .. .. 321-335

1-12
1-19
1- 8
1- 6
1- 4
1- 7
1- 5

Number 181 (published April 30, 1996)
TRANSACTIONS
1003. Toshio Koike: The first occurrence of Griesbachian conodonts in Japan.. .. .. .. .. .. .. .. .. .. .. 337-346
1004. Shuji Niko: Pseudorthoceratid cephalopods from the Early Devonian Fukuji Formation of Gifu Prefecture, Central Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 347-360
1005. Takashi Matsubara: Fossil Mollusca of the Lower Miocene Yotsuyaku Formation in the Ninohe district,
Iwate Prefecture, Northeast Japan, Part 2 (2). Gastropoda ................................. . 361-374
1006. Tatsuro Matsumoto and Akito Asai: An early Turonian (Cretaceous) new species of Inoceramus(Bivalvia) from Hokkaido ............................................................... . 375-387
1007. Naoki Kohno: Miocene pinniped Allodesmus (Mammalia: Carnivora); with special reference to the"Mito seal" from Ibaraki Prefecture, Central Japan .......................................... 388-404
In Memoriam: Teiichi Kobayashi (1901-1996) .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 405-406

1- 3
1- 8
1- 2
1- 6
1- 9

PROCEEDINGS.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 407-412

Number 182 (published June 30, 1996)
TRANSACTIONS
1008. Hiroaki Karasawa: Shako, a new Miocene stomatopod Crustacea from Japan .................. 413-418
1009. Shin'ichi Nishinosono: Larval paleoecology of the late Pleistocene naticid gastropod Cryptonatica withtwo protoconch types from the Oga Peninsula, northern Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 419-431
1010. Chang-zhu Jin and Yoshinari Kawamura: The first reliable record of Beremendia (Insectivora, Mammalia) in East Asia and a revision of Peisorex Kowalski and Li, 1963.. .. .. .. .. .. .. .. .. .. .. .. .. 432-447
1011. Boo-keun Khim, Takami Nobuhara and David E. Krantz: Stable isotope records of the Pliocene fossil
Akebiconcha kawamurai (Bivalvia: Vesicomyidae) from the Tamari Formation, Central Japan: Indicationof chemoautotrophic symbiosis ......................................................... . 448-453
1012. Hiroshi Kitazato and Satoshi Matsushita: Laboratory observations of sexual and asexual reproductionof Trochammina hadai Uchio ........................................................... . 454-466
1013. Kazutaka Amano, Mikiko Ukita and Shinji Sato: Taxonomy and distribution of the subfamily Ancistrolepidinae (Gastropoda: Buccinidae) from the Plio-Pleistocene of Japan ........................ 467-477
1014. Chang-zhu Jin and Yoshinari Kawamura: A new genus of shrew from the Pliocene of Yinan, Shan dong
Province, northern China ................................................................ 478-483

1- 4
1- 7
1-14

1- 3
1-10
1- 5
1- 6

Number 183 (published September 30, 1996)
TRANSACTIONS
1015. Michiko Saito: Early loop ontogeny of some Recent laqueid brachiopods.. .. .. .. .. .. .. .. .. .. .. 485-499
1016. Hisayoshi Kato: Miocene decapod crustacea from the Chichibu Basin, Central Japan .......... 500-521
1017. Shuji Niko, Alain Pillevuit and Tamio Nishida: Early Late Permian (Wordian) non-ammonoid cephalopods from the Hamrat Duru Group, central Oman Mountains ............................ 522-527
1018. Fumio Kobayashi: Middle Triassic (Anisian) foraminifers from the Kaizawa Formation, southern Kanto
Mountains, Japan ..................................................................... . 528-539

1- 7
1- 8
1- 3
1- 5

SHORT NOTES
34. Tatsuro Matsumoto: A little known species of Inoceramus (Bivalvia) from the Turonian (Cretaceous) of
Hokkaido .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 540-543
35. Shuji Niko, Alain Pillevuit and Tamio Nishida: Triassic aulacocerid (Mollusca: Cephalopoda) from thecentral Oman Mountains ................................................................ 544-546
36. Zaw Win and Sumio Sakagami: Preliminary note on the rediscovery of "Neoschwagerina (Sumatrina)
multiseptata Deprat" from Akasaka, Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 547-550 1- 2
PROCEEDINGS ................................................................................ 551-554

Number 184 (published December 30, 1996)
TRANSACTIONS
1019. Masayuki Noda: Five inoceramids (Bivalvia) from the Upper Cretaceous of Hokkaido with somephylogenetic and taxonomic considerations, Part 1. Introductory remarks, method and systematic
descriptions of one species of Inoceramus (I.) and one of I. (Volviceramus).. .. .. .. .. .. .. .. .. .. .. 555-570
1020. Masayuki Noda: Five inoceramids (Bivalvia) from the Upper Cretaceous of Hokkaido with somephylogenetic and taxonomic considerations, Part 2. Systematic descriptions of three species of
Inoceramus (Cremnoceramus) and concluding remarks ..................................... . 571-591
1021. Masayuki Ehiro: Latest Permian ammonoid Paratirolites from the Ofunato district, Southern Kitakami
Massif, Northeast Japan ............................................................... . 592-596
1022. Ren Hirayama and Tsutomu Chitoku: Family Dermochelyidae (Superfamily Chelonioidea) from theUpper Cretaceous of North Japan ........................................................ 597-622
1023. Hisaharu Igo: Permian fusulinaceans from the Akuda and Horikoshitoge Formations, Hachiman town,
Gifu Prefecture, Central Japan.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 623-650
1024. Kazutaka Amano, Vladimir D. Khudik and Ken Narita: An early Middle Miocene warm-water molluscan
fauna in southwestern Sakhalin-Molluscs from the Ausinskaya Formation around Novoselovo- .. 651-661
SHORT NOTES
37. Hisashi Suzuki: Archaeodictyomitra apiarium (Rust), the correct specific name of A. apiaria, A. apiara andA. apiarum ........................................................................... . 662

PROCEEDINGS.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 663-664
SYSTEMATIC INDEX .......................................................................... 665
INDEX OF GENERA AND SPECIES .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 668

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'Transactions and Proceedings of the PalaeontologicalSociety of Japan '..

,December 30, 1996

New Series No. 184
CONTENTS
TRANSACTIONS

1019. Masayuki Noda: Five inoceramids (B ivalvia) from the Upper Cretaceous of Hokkaido
with some phylogenetic and taxonomic considerations, Part 1.

Introductory remarks,

method and systematic descriptions of one species of Inoceramus (I.) and one of I.
(Volviceramus)...... .... .. .. .... .. ... . .. .. .. .. ........ .. .. .. ........ .... ........ .. 555

1020. Masayuki NOda : Five inoceramids (Bivalvia) from the Upper Cretaceous of Hokkaido
with some phylogenetic and taxonomic considerations, Part 2.

Systematic descriptionsof three species of Inoceramus (Cremnoceramus) and concluding remarks.. .. . . .. .. . . .. 571
1021 . Masayuki Ehiro : Latest Permian ammonoid Paratiro/ites from the Ofunato district, Southern Kitakami Massif, Northeast Japan...... .. ...... .. .. .... .. .................. .... 592

1022. Ren Hirayama and Tsutomu Chitoku : Family Dermochelyidae (Superfamily Chelonioidea),from the Upper Cretaceous of North Japan .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 597
1023. Hi?aharu Igo: Permian fusulinaceans from the Akuda and Horikoshitoge Formations,
Hachiman town , Gifu Prefecture, Central Japan .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 623
1024. Kazutaka Amano , Vladimir D. Khudik and Ken Narita: An early Middle Miocene warmwater molluscan fauna in southwestern Sakhalin- Molluscs from the Ausinskaya Formation . around Novoselovo-

651

SHORT NOTES
37.

Hisashi Suzuki: Archaeodictyomitra apiarium (Rust), the correct specific name of A.
apiaria, A. apiara and A. apiarum

662

PROCEEDINGS ............ .. .......... .... .. .. .. .. ...... , . .. ..
SYSTEMATIC INDEX••••• •

••

• •

••

' 0

INDEX OF GENERA AND SPECIES

,0

••••

' 0

••••••••••••••• •••••• •

663••••••• •• •••••

665
668