ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No.

163

Palaeontological Society of Japan
September

30 , 1991

Co-Editors

Tsunemasa Saito and Kei Mori
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The fossil on the cover is Trilophodon sendaicus Matsumoto, an extinct elephant, which
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 163, pp. 807-827, 6 Figs., September 30, 1991

922. UPPER CARBONIFEROUS FUSULINACEANS FROM THE
AKIYOSHI LIMESTONE GROUP, SOUTHWEST JAPAN*
KA TSUMI UENO
Institute of Geoscience, University of Tsukuba,
Ibaraki 305

Abstract. Twelve species belonging to seven genera of fusulinaceans from theProtriticites sp. and Quasifusulinoides toriyamai Zones of the Akiyoshi Limestone Group
distributed in the Akago-Managatake area in the northeastern part of Akiyoshi Plateau,
Yamaguchi Prefecture are described herein. Among them, Protriticites robustus, Obsoletes
horridus and Quasifusulinoides toriyamai are new taxa. This fusulinacean fauna is characterized by the genera Protriticites, Obsoletes and Quasifusulinoides, and is closely related
to those reported from the Moscow Syneclise, Donetz Basin, southern Fergana of Central
Asia and Tadushi River Basin of southern Sikhote-Alin in the generic composition. Itsgeologic age is referable to the early Kasimovian.
Key words. Akiyoshi Limestone Group,
Quasifusulinoides, Upper Carboniferous.

Kasimovian,Obsoletes,

Protriticites,

paleontological and stratigraphical knowledge of the Upper Carboniferous in the
Akiyoshi Limestone Group still remains to
be poor.
Recently, I clarified a foraminiferal biostratigraphy of the lower part of the Akiyoshi
Limestone Group distributed in the AkagoManagatake area of the northeastern part of
Akiyoshi Plateau, and established theProtriticites sp. and Quasifusulinoides toriyamai Zones in a stratigraphic interval
between the Beedeina akiyoshiensis andTriticites "simplex" Zones (Ueno, 1989).
In the present paper, I describe 12 species offusulinaceans including three new ones,
Protriticites robustus, Obsoletes horridus andQuasifusulinoides toriyamai, from theProtriticites sp. and Quasifusulinoides toriyamai Zones.
All of the described specimens are kept in
the Akiyoshi-dai Museum of Natural History.
The abbreviation· ASM is prefixed to thespecimens' repository number.

Introduction

Until the discovery of fusulinacean faunacharacterized by Triticites (s.l.) matsumotoi in
the Akiyoshi Limestone Group by Ota(1968), the Upper Carboniferous strata had
been considered to be entirely lacking not
only in the Akiyoshi Limestone Group but
also in the most of limestone plateaus ofSouthwest Japan. At that time, he established the Triticites (s.l.) matsumotoi Zone,
which overlies the Middle Carboniferous
Beedeina akiyoshiensis Zone. Later, Ota etal. (1973) described a fusulinacean fauna of
this zone and correlated it to the Kasimovian
of USSR or the lower part of the Kawvian ofNorth America.
The distribution of the Triticites (s.l.)
matsumotoi Zone, however, is much
restricted and has been known only in a small
area about 200 m north of the check point on
the Akiyoshi-dai toll road. Therefore,"Received September 4, 1990; revised manuscript
accepted May IS, 1991
807

Katsumi Uena

808

Geologic setting

In the Akago-Managatake area situated in
the northeastern part of Akiyoshi Plateau,
Yamaguchi Prefecture, the Akiyoshi Limestone Group comprises a thick b asa ltic basement rock and the overlying white to gray,pure, massive limestone. The limestone inthis area has nearly a complete succession
from the late Visean to Gzhelian and yields
rich shallow-marine organic remain s.
Based on mainly fusulinacean foraminifers,
Ueno (1989) subdivided the Carboniferous
sequence of the Akiyoshi Limestone Groupin this area into the following 13 zones indescending order.
Triticiles "simplex" Zone
Quasifusulinoides toriyamai Zone

Protriticites sp. Zone
Beedeina akiyoshiensis Zone
Pseudofusulinella taishakuensis Zone
Fusulinella biconica Zone
Akiyoshiella ozawai Zone
Pseudostaffella anliqua Zone
Millerella yowarensis Zone
Eostaffella bigemmicula Zone
Eostaffella mosquensis Zone
Mediocris breviscula Zone
Endothy ra Zone

These zones are distributed in a trendparallel to an almost NW to SE direction andbecome younger southwestwards. Strikes
an d dips measured in the field and also fromoriented rock samples trend N 10 -50 W and
20 -40 NE, respect ively. Judging from theavailable data , the Akiyoshi LimestoneGroup in the Akago-Managatake area is completely overturned and dips monoclinally
0

0

1
KAMA

2

II

3

0

0

1_ 1

4

115
•

6
_ 200mIo_ _ _
100

UEYAMA
Figure 1. Map showing the di stribution of foraminiferal zones and fossil localities stud ied herein. I:
Pseudofusulinella taishakuensis Zone, 2: Beedeina akiyoshiensis Zone, 3: Protriticites sp. Zone, 4:Quasifusulinoides IOriyamai Zone, 5: Triticires "simp/ex" Zone, 6: fossil loca lity.

922.

Carboniferous fusulinaceans from Akiyoshi

northeastwards.
The distribution of the Protriticites sp. andQuasifusulinoides toriyamai, and their overlying and underlying zones are shown in
Figure 1. The Protdticites sp. Zone, about
15 m thick, conformably overlies the Moscovian Beedeina akiyoshiensis Zone and isrecognized in the US, BS and KU sections.
This zone is defined as the stratigraphic interval between the first occurrence of the genus
Protriticites and that of Quasifusulinoides
toriyamai. An algal sparitic limestone ispredominant in this zone with a subordinate
amount of algal micritic limestone.
The Quasifusulinoides toriyamai Zone,
about 50 m in maximum thickness, conformably overlies the Protriticites sp. Zone and isobserved only in the US and BS sections.
Lithology of the limestone is similar to thatof the Protdticites sp. Zone.
The Triticites "simplex" Zone has a clinounconformable contact with the underlying
Quasifusulinoides todyamai, Protriticites sp.
and Beedeina akiyoshiensis Zones. Consequently, it directly overlies the Beedeina
akiyoshiensis Zone in the MT section.

Faunal affinity and correlation
The following fusulinaceans are discriminated in the Protdticites sp. andQuasifusulinoides todyamai Zones.

809

Among these fusulinaceans, the following
three genera, Protriticites, Obsoletes andQuasifusulinoides, are characteristic and areimportant to discuss faunal affinity and age
determination.
At present, the fusulinacean fauna characterized by the genera Protdticites, Obsoletes
and Quasifusulinoides is known in a stratigraphic interval between the Middle Carboniferous Fusulina (or Beedeina) Zone and theUpper Carboniferous Triticites Zone in several regions of USSR, such as the Moscow
Syneclise, Donetz Basin, southern Fergana ofCentral Asia and Tadushi River Basin ofsouthern Sikhote-Alin (Figure 2). This
stratigraphic interval corresponds to thelower Kasimovian. The stratigraphy of the
Kasimovian in the above-mentioned areas issummarized as follows:
Moscow Syneclise: The stratotype of the
Kasimovian Stage is located in the Moscow
Syneclise of the Russian Platform. The
Kasimovian is represented mainly by marine
carbonate rocks and conformably overlies the
Moscovian. It is subdivided into the following four horizons, namely, Kreviakinsky,
Khamovnichesky, Dologomilovsky and Yauzsky in ascending order. Each horizoncorresponds to the Protriticites pseudomontiparus - Obsoletes obsoletus Zone (C 3 A 1 ),Protriticites sp. Zone
Protriticites robustus, sp. nov., P. sp., Obsoletes horridus, sp. nov., O. cf. obsoletus(Schellwien), Pseudofusulinella (Kanmeraia)
praeantiqua Nassichuk and Wilde, P. (K.) cf.
delicata Skinner and Wilde, and Staffella
pseudosphaeroidea Dutkevich.
Quasifusulinoides toriyamai Zone
Quasifusulinoides toriyamai, sp. nov.,
Pseudofusulinella (Kanmeraia) praeantiqua
Nassichuk and Wilde, Schubertella aff. kingi
Dunbar and Skinner, S. sp., Eostaffella sp. A
and E. sp. B.

Figure 2. Geographic distribution of theProtriticites-Obsoletes-Quasifusulinoides fauna. 1:
Moscow Syneclise, 2: Donetz Basin, 3: Southern
Fergana, 4: Tadushi River Basin, 5: Akiyoshi.

810

Katsumi Ueno

Montiparus montiparus Zone (C aA2)'
Triticites arcticus-T. ohioensis Zone (CaBl)
and T. irregularis-T. acutus Zone (C aB2)'
respectively. The stratigraphic distributionof fusulinaceans in the stratotypic section wasshown by Rauser-Chernoussova et al. (1979).
According to them, the genus Protriticites (P.
globulus, P. pseudomontiparus and P. sphaericus) is abundant in the Kreviakinsky Horizon, but its actual range is slightly wider andthe genus first appears in the upper Myachkovsky Horizon of Moscovian. The genus
Obsoletes (0. obsoletus and O. gapeevi) whichflourished in the Kreviakinsky Horizon
ranges up to the Khamovnichesky Horizon.
The genus Quasifusulinoides (Q. fusiformis)
appears in the Myachkovsky Horizon, but
becomes common in the Kreviakinsky Horizon. These three genera coexist only in the
Kreviakinsky Horizon.
Donetz Basin: The Donetz Basin contains
an almost complete and practically uninterrupted Carboniferous succession of strata up
to 12,000 m in thickness. The Carboniferous
stratigraphy in the Donetz Basin described by
Aisen berg et al. (1979) indicates that the
Kasimovian in this basin corresponds to theinterval from the N2-Na limestones to limestone Pl. It is further subdivided into threezones, C~, C~ and C~. The C~ Zone contains
Quasifusulinoides and Protriticites with someother genera that ranged upward from the
Moscovian. In the lower part of q Zone
(limestones 0 1 -02), the genera Protriticites,
Obsoletes and Quasifusulinoides are abundant, whereas the upper part of C~ Zone
(limestones Oa-04) contains the genus
Montiparus (M. ex gr. montiparus). The C~
Zone is characterized by the abundant occurrence of Triticites.
Southern Fergana of Central Asia:
Bensh (1972) reported the Upper Carboniferous and Lower Permian stratigraphy andfusulinaceans of southern Fergana and established the following three zones in the
Kasimovian: the Protriticites pseudomontiparus-Obsoletes obsoletus, Montiparus

montiparus and Triticites arcticus-T. acutus
Zones. Several species of Protriticites, Obsoletes and Quasifusulinoides are contained
only in the lowest zone.
Tadushi River Basin of southern SikhoteAlin: In the Tadushi River Basin of southern
Sikhote-Alin, a fusulinacean fauna consistingof the genera Obsoletes, Protriticites andQuasifusulinoides was reported by Nikitina(1969) from a limestone lens in the basal partof the Upper Carboniferous. From a somewhat lower horizon in the same section, typical Moscovian fusulinaceans, such as Fusiellacf. typica. Fusulinella bocki, F. pseudobocki
and others, are recorded together with
Protriticites? sp. and Obsoletes? sp. She
concluded that the Obsoletes-Protriticites
Zone characterized by Obsoletes, Protriticites
and Quasifusulinoides can be established in
the basal part of the Upper Carboniferous.
In the Akiyoshi Limestone Group, thefusulinacean zones characterized by the genera Protriticites, Obsoletes and Quasifusulinoides are situated between the Beedeina (B.
akiyoshiensis) and Triticites (T. "simplex")
Zones, although a stratigraphic hiatus exists
between the Triticites "simplex" andQuasifusulinoides toriyamai Zones. The
fusulinacean faunal assemblage of theProtriticites sp. and Quasifusulinoides toriyamai Zones has a close similarity to thosereported from the Moscow Syneclise, Donetz
Basin, southern Fergana of Central Asia andTadushi River Basin of southern SikhoteAlin and is surely referable to the lower
Kasimovian of USSR.

Systematic paleontology
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea
von Moller, 1878
Family Eostaffellidae Mamet, in Mamet,
Mikhailov and Mortelmans, 1970
Genus Eostaffella Rauser-Chernoussova, 1948

922.

Carboniferous fusulinaceans from Akiyoshi

Eostaffella sp. A
Figure 5-1

Material studied.-Axial section; ASM25078A from Loc. BS2l8.6.
Description.-Shell small, involute andhighly lenticular. Periphery angular, andumbilical regions slightly depressed in thelast volution. Specimen having 2 ~ volutions, measuring 0.09 mm in length and 0.35
mm in width, with a form ratio of 0.26.
Proloculus spherical and 0.05 mm in its outside diameter. Spirotheca very thin, consisting of a tectum and upper and lower tectoria.
Chomata low, broad and asymmetrical.
Tunnel angle of the second volution 8
degrees.
Remarks.- The present species is characterized by a highly lenticular shell andresembles somewhat Eostaffella mutabilis
described by Rauser-Chernoussova (inRauser-Chernoussova et al., 1951) from the
Moscovian of the Russian Platform. However, the former has a fewer number of volution and smaller form ratio than the latter.
Occurrence.-Rare in the upper part of the
Quasifusulinoides toriyamai Zone.
Eostaffella sp. B
Figure 5-2

Material studied.-Axial section; ASM25062 from Loc. BS2l5.
Description.-Shell small and discoidal
with rounded periphery. Illustrated specimen having 3 volutions, measuring 0.13 mm
in length and 0.31 mm in width, giving a form
ratio of 0.42. Proloculus spherical and relatively large for shell size, being 0.055 mm inits outside diameter. Composition of spirotheca not clear owing to its poor preservation.
Chomata present in the last volution. Tunnel angle of the third volution 22 degrees.
Remarks.- The present form is similar to
Eostaffella pseudostruvei forma minima described by Kireeva (1949) from central Donbass. However, specific identification of this811

specimen is postponed because of its illpreservation. Eostaffella sp. B can be easily
distinguished from E. sp. A of this study by
the broad shell periphery and large form ratioof the former.
Occurrence.-Rare in the upper part of the
Quasifusulinoides toriyamai Zone.
Family Schubertellidae Skinner, 1931
Genus Schubertella Staff and Wedekind, 1910
Schubertella aff. kingi Dunbar
and Skinner, 1937
Figures 5-4-11

Compare.Schubertella kingi Dunbar and Skinner, 1937, p.
610-611, pl. 45, figs. 10-15; Thompson and
Wheeler in Thompson et al., 1946, p. 24-25, pl.
8, figs. 6-10; Thompson and Hazzard in Thompson et al., 1946, p. 40-41, pl. 10, figs. 1-9 ;Thompson, 1948, pl. 4, fig. 21 (same as pl. 45, fig.
12 of Dunbar and Skinner, 1937, p. 610-611);
Thompson, 1954, p. 33-34, pl. 5, figs. 11-42, pl. 7,
figs. 11-13; Igo, 1957, p. 192-194, pl. 4, figs. 915; Sakagami and Ornata, 1957, p.249-250, pl.
19, fig. 1; Toriyama, 1958, p. 73-75, pl. 7, figs. 18; Kochansky-Devidi!, 1959, p. 17-18,47, pl. 1,
figs. 7-12; Suyari, 1962, p. 8, pl. 1, figs. 10-11 ;Douglass in Mudge and Y oche1son, 1962, p. 6061, pl. 8, figs. 5-7; Sabins and Ross, 1963, p.
360-361, pl. 38, figs. 9-12; Skinner and Wilde,
1965, p.25, pl. 27, figs. 4-11 ; Kauffman andRoth, 1966, p. 6, pl. 1, figs. 1-4; Stewart, 1966, p.
84, pl. 1, figs. 13-14 (same as pl. 45, figs. 14, 12 of
Dunbar and Skinner, 1937, p. 610-611); Stewart,
1968, p.326, pl. 41, figs. 13-14 (same as pl. 45,
figs. 14, 12 of Dunbar and Skinner, 1937, p. 610611); Han, 1976, p. 25, pl. 3, figs. 2-3; Igo et ai,
1977, p. 100, pl. 17, figs. 9-10, pl. 18, figs. 11-14;
Leven and Scherbovich, 1978, p. 85, pl. 1, figs. 78; Chen and Wang, 1983, p.35-36, pl. 1, figs.
31-33, pl. 2, figs. 1-3; Han and Zhao, 1984, p.
62-63, pl. 2, figs. 37-40; Sheng and Wang, 1984,
p. 33-34, pl. 1, figs. 25-26; Niko and Watanabe,
1987, p.46, figs. 4-A-H; Han, 1988, pl. 1, fig. 5.

Material studied.-Axial sections; ASM25070, ASM25079, ASM25067, ASM25073,
ASM25078B from Loc. BS218.6 andASM25059B from Loc. BS201.5. Sagittal
sections; ASM25077 and ASM25065 from
Loc. BS218.6.

Katsumi Ueno

812

Description.-Shell small and fusiform toelongate fusiform. Polar regions bluntly to
sharply pointed. Mature specimens having 4
to 5 volutions, measuring from 0.60 to 0.80
mm in length and 0.28 to 0.40 mm in width.
Form ratio ranging from 2.03 to 2.57, averaging 2.12 for 6 specimens.
Inner 2 or 2 Yz volutions coiled almost
perpendicularly to remaining planispiralones. Average radius vectors of the first to
fifth volutions for 8 specimens being 0.04,
0.06,0.09,0.14 and 0.19 mm. Average form
ratios of the first to fifth volutions for 6
specimens being 0.91, 0.99, 1.20, 1.81 and 2.09,
respectively.
Pro loculus small and almost spherical. Itsoutside diameter ranging from 0.03 to 0.045
mm, averaging 0.04 mm for 8 specimens.
Spirotheca thin and composed of a single
homogeneous layer in inner 2 or 3 volutions,but a tectum and lower discontinuous lessdense layer (diaphanotheca) in outer ones.
A verage thickness of spiro theca of the first to
fifth volutions for 8 specimes being 0.005,
0.008,0.010,0.010 and 0.Ql5 mm, respectively.
Septa short, straight and unfluted in central
part of shell, but weakly fluted in extreme
polar regions. Massive chomata well developed in outer 3 vol utions. A verage tunnel
angles of the third to fifth volutions for 6
specimens measuring 17, 31 and 35 degrees,
respecti vel y.
Remarks.- This species resembles Schubertella kingi Dunbar and Skinner which was
originally described from the Wolfcampian ofTexas. However, development of the lower
less dense layer (diaphanotheca) in the present specimens seems to be more feeble anddiscontinuous than that in the original ones.
In addition, the former shell is smaller thanthe latter.
Occurrence.-Abundant in the Quasifusulinoides toriyamai Zone.
Schubertella sp.
Figures 5-12-13

Material studied.-Axial sections; ASM25063 from Loc. BS218.3 and ASM25080
from Loc. BS221.
Description.-Shell small and bulged tothickly fusiform with rounded polar regions.
Mature specimens having 4 Yz volutions,
measuring from 0.73 to 0.78 mm in length and0.43 mm in width. Form ratio varying from1.70 to 1.81. Inner 2 volutions tightly coiled
at a large angle to outer ones. Form ratiosof the first to fifth volutions of typical specimen (Figure 5-13) being 0.75,1.17,1.20,1.48
and 1.91, respectively. Proloculus small andspherical. Its outside diameter ranging from0.04 to 0.06 mm. Spirotheca thin, composedof a single homogeneous layer in inner few
volutions, and an extremely thin tectum and a
discontinuous lower less dense layer (diaphanotheca) in outer ones. Septa almost
plane. Chomata weakly developed in outer
3 volutions.
Remarks.- The present species is similar to
Schubertella aff. kingi of this study, butdiffers from the latter in having a smaller
form ratio in the mature stage. This unidentified species has a shell smaller than thatof Schubertella kingi exilis described from theLower Permian of South Urals by Suleimanov (1949).
Occurrence.-Rare in the upper part of the
Quasifusulinoides toriyamai Zone.
Family Fusulinidae von Moller, 1878
Subfamily Fusulinellinae Staff
and Wedekind, 1910
Genus Protriticites Putrja, 1948
Protriticites Putrja, 1948, p.90-91.

Diagnosis.-Shell small to moderate insize, inflated fusiform to subcylindrical inshape, with rounded to bluntly pointed polarends. Axis of coiling straight throughout.
Proloculus commonly small. Spirothecathin to rather thick, composed of 4 layers in
inner volutions, a tectum, diaphanotheca andupper and lower tectoria, but of 3 layers in922.

Carboniferous fusulinaceans from Akiyoshi

outer ones, a tectum, alveolar keriotheca andthin , inconstant upper tectorium . Upper
tectorium not seen in the last volution. In
well-preserved specimens, minute pores penetrating all 4 layers in inner volutions. Septa
weakly to moderately fluted in polar regions,but unfluted in central part of shell.
Chomata well developed and massive. Tunnel path straight. Axial fillings absent.
Remarks. - The genus Protriticites wasintroduced by Putrja (1948) with Protriticites
globulus as the type species. This genu s ischaracterized by its peculiar spirothecal structure that has a Fusulinella-like spirotheca in

8 13

the inner volutions and Triticites-like spirotheca in the outer ones. In the original genericdiagnosis, Putrja described minute pores (finealveolar structure) penetrating all 4 layers in
the inner volutions, but this delicate structure
can be seen only in well-preserved specimens.

Protriticites robustus Ueno , sp. nov.
Figures 3- 1- 4

Material studied. - Axial section ofholotype ; ASM25040B. Slightly oblique
axial section of paratype; ASM25040A.
Sagittal section of paratype ; ASM25042.

Figure 3. 1- 4. Protriticites robustus , sp. nov., 2a: axial section of holotype, ASM25040B, 1 : slightly
oblique axial section of paratype, ASM25040A , 3: tangential section o f paratype, ASM25035 , 4: sagittal
section of paratype, ASM25042, X 15, 2b : enlarged part of la, x 75 . 5,6: Protriticites sp., oblique sections,
ASM25081 , ASM25029, X 20.

Katsumi Ueno

814

Table 1.

Measurements of Protriticites robustus, sp. nov. (in mm).
Radius vector

Specimen number

FigureLength

Width

F.R.

D.P.
2

I

3

4

5

6

I

ASM25040A

3-1

3.88

2.13

1.82

0.12

0.12

0.20

0.33

0.52

0.81

1.15

2

ASM25040B

3-2

4.45

2.03

2.19

0.16

0.13

0.22

0.38

0.68

1.03

lAO'

Form ratio

I

2

I

2

3

4

5

6

25

17

23

26

39

-

0.030 0.060 0.090 0.070 0.090" 23

20

27

28

-

-

I

2

3

4

5

6

1.29

1.49

1048

1.52

1.60

1.70

-

0.040 0.050 0.065

1.85"

0.QI5

1.28

1.82

1.89

2.06

Tunnel angle (degrees)

Thickness of spirotheca

2.17

I

2

4

3

5

6

0.090 0.090

F.R.: Form ratio, D.P.: Diameter of proloculus, " : 5 ~ volution

Tangential section of paratype; ASM25035.
All specimens from Loc. BS199.
Descrzption.-Shell medium in size andtypically fusiform with bluntly pointed polarends. Lateral slopes almost straight.
Holotype having 5 Yz volutions, measuring
4.45 mm in length and 2.03 mm in width,
giving a form ratio of 2.19. Axis of coilingstraight. The first volution tightly coiled,
but later ones becoming expanded gradually.
Radius vectors of the first to fifth volutions of
the holotype being 0.13, 0.22, 0.38, 0.68 and1.03 mm, giving form ratios of 1.28, 1.82, 1.89,
2.06 and 2.07, respectively.
Prolocul us spherical, ranging from 0.12 to
0.16 mm in its outside diameter.
Spirotheca thin, composed of a tectum,
thin, indistinct diaphanotheca and upper andlower tectoria in the second volution, but
relatively thick and composed of a tectum,
thick lower alveolar keriotheca and inconstant upper tectorium in subsequent 2 volutions, and composed of a tectum and alveolar
keriotheca in outermost one. Thickness of
spirotheca of the first to fifth vol utions of theholotype measuring 0.015,0.030,0.060,0.090
and 0.070 mm, respectively.
Septa nearly plane in central part of shell,
but weakly to moderately fluted in polar
regions. Sep~al counts of the first to sixth
volutions of illustrated specimen (Figure 310) being 8, 12, 16, 18,24 and 22, respectively.

Chomata broad, massive and observed
from the first to penultimate volutions. Tunnel angles of the first to fourth volutions of
the holotype being 23, 20, 27 and 28 degrees,
respectively. Axial fillings absent.
Remarks.-Protriticites robustus, sp. nov.
can be distinguished easily from Protriticites
globulus Putrja by its less-developed chomata,larger proloculus and smaller form ratio.
Ota (1977) illustrated Triticites (s.l.) matsumotoi from the Triticites (s.l.) matsumotoi
Zone of the Akiyoshi Limestone Group.
Triticites (s.l.) matsumotoi can be distinguished from the present new species in having a smaller shell and well-defined,
Fusulinella-like, 4-layered spirotheca in the
inner volutions.
Protriticites robustus, sp. nov. resembles
some species of Montzparus in the shellmorphology, but differs from the latter inhaving a 4-layered spirotheca In the inner
volutions.
Measurements.-See Table 1.
Occurrence.-Rare in the uppermost partof the Protriticites sp. Zone.

Protriticites Sp.
Figures 3-5-6

Material studied.-Oblique sections;
ASM25081 from Loc. KU184 and ASM25029
from Loc. US21O. Some additional, but922.

Carboniferous fusulinaceans from Akiyoshi

poorly oriented specimens from theProtriticites sp. Zone were also studied.
Description.- Two oblique sections (Figures 3-5,6) having 6 volutions, measuring2.60 and 2.70 mm in length, and 1.45 and1.50 mm in width. Pro loculus spherical,
measuring 0.10 and 0.08 mm in outside diameter. Spirotheca composed of a single homogeneous layer in inner 2 volutions, a tectum,
diaphanotheca and upper and lower tectoria
in middle ones, and a tectum and finely
alveolar keriotheca in outer 2 volutions.
Septa almost plane. Chomata present.
Remarks.-Because all the examined specimens are poorly oriented, specific identification is impossible. The present form canbe distinguished from Protriticites robustus,
sp. nov. in having a smaller shell.
Occurrence.-Common in the Protriticites
sp. Zone.
Genus Obsoletes Kireeva, 1950
Obsoletes Kireeva, 1950, p. 201-202.

Diagnosis.-Shell small to moderate insize, elongate fusiform to cylindrical in shape.
Axis of coiling almost straight. Proloculus
spherical, small to medium. Spirotheca usually thin and quite complicated, composed ofa tectum, diaphanotheca and very thin, discontinuous lower tectorium in inner volutions. Outer ones consisting of a tectum,
rather thick alveolar keriotheca and thin,inconstant upper tectorium. Very minutepores penetrating a diaphanotheca observablein well-preserved specimens. Septa notfluted in central part of shell, but weakly
fluted in extreme polar regions. Chomata
present, weak to massive. Axial fillings
absent.
Discussion and remarks.- The genus Obsoletes was established by Kireeva (1950) with
Fusulina obsoleta Schellwien as the typespecies. She described the ontogenetic
change of spirothecal structure in this genus.
Later, Rauser-Chernoussova and Fursenko

815(1959) briefly stated that the spirotheca of thegenus Obsoletes is thin, porous and composedof a tectum, pro theca and inconstant, verythin upper tectorium. They did not refer to
its peculiar spirothecal structure which Kireeva had pointed out. Some fusulinacean
students, however, recognized that a diaphanotheca and discontinuous thin lower tectorium can be observed in the inner volutionsof the genus Obsoletes (e.g., Chen, 1963;
Bensh, 1972; Rozovskaya, 1975). Such a
spirothecal structure somewhat resembles thatof the genus Protriticites, and this causes
ambiguity in the generic status of Obsoletes.
Rozovskaya (1975) stated that the genus
Obsoletes can be distinguished from the genus
Protriticites in having a spiro theca with thealveolar structure (keriotheca) in the outermost volution. However, the alveolar keriotheca also exists in the genus Protriticites.
As mentioned above, the taxonomic status
between these two genera is confused, especially in their spirothecal composition.
However, the genus Obsoletes can be distinguished from the genus Protriticites in having
a more elongate shell, slightly larger form
ratio, less developed chomata and thinnerspirotheca. Moreover, concerning theirspirothecal composition in the inner volutions, the former genus has a 3-layered spirotheca, composed of a tectum, diaphanotheca, and thin discontinuous lower tectorium.
On the other hand, the latter genus has a
4-layered spirotheca, composed of a tectum,
diaphanotheca, and upper and lower tectoria.
The spirothecal composition in the outer
volutions is closely allied to each other.

Obsoletes horridus Veno, sp. nov.
Figures 4-1-8

Material studied.-Axial section of holotype; ASM25037 A. Axial section of immatureparatype
specimen; ASM25038.
Slightly oblique axial sections of paratypes ;ASM25043, ASM25039, ASM25033A andASM25041. Sagittal sections of paratypes;

8 16

Katsumi Veno

12b

922.

Carboniferous fusulinaceans from Akiyoshi

ASM25045 and ASM25041. Sagittal sections of paratypes; ASM25045 and ASM
25033B. In addition, some other specimens
were studied. All specimens from Loc.
BS199.
Description.-Shell large for the genus andelongate fusiform in shape. Polar regions
bluntly pointed. Mature shell having 5 Yz to
6 volutions, measuring from 4.30 to 5.08 mm
in length and 1.53 to 2.03 mm in width.
Form ratio of complete axial section(holotype) being 3.12.
Axis of coiling straight. Shell expanding
gradually through growth. Average radius
vectors of the first to sixth volutions for 8
specimens being 0.14, 0.21, 0.33, 0.50, 0.75 and0.82 mm. Form ratios of the first to sixth
volutions of the holotype being 1.84, 2.17,
2.51,2.38,2.73 and 2.98, respectively.
Proloculus large and spherical. Its outside
diameter varying from 0.16 to 0.24 mm, aver-817

aging 0.20 mm for 13 specimens.
Spirotheca thin and composed of a tectum,
diaphanotheca and thin lower tectorium in
inner volutions. Spirotheca in outer 2 volutions consisting of a tectum, finely alveolar
keriotheca and thin, inconstant upper tectori urn. A verage thickness of spirotheca of thefirst to sixth volutions for 8 specimens being
0.D15, 0.020, 0.030, 0.037, 0.054 and 0.080 mm.
Septa straight and not fluted in central partof shell, but weakly to moderately fluted inpolar regions. Septal counts of the first to
sixth volutions of illustrated specimen (Figure4-6) numbering 7,15,17,17,19 and 18,
respectively.
Chomata massive and well developed in all
volutions except for the last one. Tunnel
angles of the first to sixth volutions for 6
specimens averaging 24,27,33,51,57 and 71
degrees. Axial fillings absent.
Remarks.- The present new species is

Measurements of Obsoletes horridus, sp. nov. (in mm).

Table 2.

Radius vector
Specimen number

Fig.

Length

Width

F.R.

D.P.
I

2

3

4

5

6

I

ASM25037A

4-1

5.08

1.63

3.12

0.19

0.13

0.18

0.28

0.42

0.62

0.86

2

ASM25043

4-3

4.43

1.53

2.90

0.14

0.11

0.18

0.25

0.38

0.57

0.82

3

ASM25033A

4-5

5.00

1.90

2.63

0.24

0.19

0.23

0.44

0.67

0.93

4

ASM25038

4-8

3.00

1.14

2.63

0.18

0.11

0.18

0.26

0.39

0.59*

Form ratio

Thickness of spiro theca

Tunnel angle (degrees)

I

2

3

4

5

I

1.84

2.17

2.51

2.38

2

1.55

1.60

1.66

1.97

3

1.68

2.58

2.18

2.69

2.63

0.Ql5

0.020

0.040

0.065

0.070

26

27

39

50 61

4

1.64

2.28

2.55

2.92

2.50*

0.010

0.015

0.020

0.030

0.060'

30

27

27

41 54*

6

I

2

3

4

I

2

3

4

2.73

2.98

0.020

0.020

0.030

1.80

2.38

0.Ql5

0.020

0.035

0.035

0.050 0.070 24

39

39

59 65

71

0.030

0.045 0.090

17

18

37 53

70

5

6

18

5

6

* : 4 ~ volution+- Figure 4. 1-8. Obsoletes horridus, sp. nov., 1: axial section of holotype, ASM25037 A, 2, 6: sagittal
sections of paratypes, ASM25045, ASM25033B, 3-5a, 7: slightly oblique axial sections, paratypes,
ASM25043, ASM25039, ASM25033A, ASM25041, 8: axial section of immature paratype specimen,
ASM25038, X 15, Sb: enlarged part of Sa. X 75. 9-13. Obsoletes cf. obsoletus (Schellwien), 9: sagittal
section, ASM25037B, 10-12a, 13: axial sections, ASM25036, ASM25044, ASM25032, ASM25034, X 15,
12b: enlarged part of 12a, X 75.

Katsumi Ueno

818

allied to Obsoletes fusiformis described by
Bensh (1972) from the Kasimovian Protriticites pseudomontiparus - Obsoletes obsoletus Zone of southern Fergana. However, theformer possesses a larger shell than the latter.
The present form also resembles Obsoletes
magnus Kireeva from the C ~ Suite of theDonetz Basin, but differs from the latter inhaving a larger shell.
Measurements.-See Table 2.
Occurrence.-Common in the uppermost
part of the Protdticites sp. Zone.

Obsoletes cf. obsoletus (Schell wien, 1908)
Figures 4-9-13

Compare.Fusulina obsoleta Schellwien, 1908, p. 167-168, pI. 19,
figs. 5-7.
Neofusulina obsoleta (Schellwien); Lee, 1927, p. 1819, pI. 2, fig. 19.
Protriticites obsoletus (Schellwien); Putrja, 1948, p.
94, pI. 1, fig. 7; Rozovskaya, 1950, p. 10-11, pI.
I, figs. 1-4.
Obsoletes obsoletus (Schellwien); Rauser-Chernoussoya and Fursenko, 1959, pI. 7, fig. 8 (same as pI.
1, fig. 2 of Rozovskaya, 1950, p. 10-11); Chen,
1963, pI. 1, figs. 2-3, pI. 2, fig. 3; Pasini, 1965, pI.
7, fig. 1 (same as pI. 19, fig. 7 of Schell wi en, 1908,
p. 167-168); Rozovskaya, 1975, pI. 12, figs. 5-6(fig. 5: same as pI. 19, fig. 7 of Schellwien, 1908,
p. 167-168, fig. 6: same as pI. 1, fig. 2 of Rozovskaya, 1950, p. 10-11); N iikawa, 1978, p. 562563, pI. 12, figs. 1-2; Ozawa and Kobayashi,
1990, pI. 3, figs. 16-18.
Obsoletes obsoleta (Schellwien); Loeblich andTappan, 1988, pI. 267, figs. 10-12 (same as pI. 19,
figs. 6-7,5 of Schellwien, 1908, p. 167-168).
Protriticites aft'. obsoletus (Schellwien); Sheng, 1958,
p. 36-37, 95-96, pI. 10, fig. 12.

Material studied.-Axial sections;
ASM25036, ASM25044, ASM25032 andASM25034. Sagittal section; ASM25037B.
Some additional specimens were also studied.
All specimens from Loc. BS199.
Description.-Shell large for the genus andelongate fusiform to cylindrical in shape withbluntly pointed or rounded polar ends.
Lateral slopes slightly convex. Mature specimens of 6 volutions, measuring from 5.08 to

5.30 mm in length and 1.53 to 2.05 mm inwidth. Form ratio varying from 2.58 to 3.48,
averaging 3.01 for 4 specimens.
Axis of coiling straight throughout andshell expanding gradually through growth.
Average radius vectors of the first to sixth
vol utions for 5 specimens being 0.12, 0.21,
0.31,0.47,0.71 and 0.93 mm. Form ratios of
the first to sixth volutions for 4 specimens
being 1.45,1.81,2.05,2.32,2.58 and 2.57,
respectively.
Proloculus small and spherical. Its outside diameter ranging from 0.12 to 0.16 mm,
averaging 0.14 mm for 4 specimens.
Spirotheca thin, composed of a tectum,
diaphanotheca and thin lower tectorium in
inner volutions, but a tectum and finely alveolar keriotheca in outer ones. Inconstant
upper tectorium sometimes observed in outer
volutions. Average thickness of spirothecaof the first to sixth volutions for 5 specimens
measuring 0.022, 0.027, 0.031, 0.040, 0.062 and0.105 mm, respectively.
Septa weakly fluted only in axial regions.
Septal counts of the first to fifth volutions ofillustrated sagittal section (Figure 4-9) numbering 7, 15,21,20 and 15.
Chomata low and small. A verage tunnel
angles of the second to fifth volutions for 4
specimens measuring 23, 32, 41 and 57
degrees, respectively. Axial fillings absent.
Remarks.- The Akiyoshi specimens arealmost identical with Obsoletes obsoletus
which Schellwien (1908) originally describedfrom the Upper Carboniferous in the Donetz
Basin of USSR, in their essential morphological characters. However, the former
has a larger shell and less-developed chomata
than the latter. Exact identification is postponed pending the availability of better
materials.
Obsoletes cf. obsoletus differs from 0. horridus described herein in having less developed chomata and smaller proloculus.
Occurrence.-Common in the uppermost
part of the Protriticites sp. Zone.

922.

Carboniferous fusulinaceans from Akiyoshi

Genus Pseudofusulinella Thompson, 195 I
Subgenus Kanmeraia Ozawa, 1967
Pseudofusulinella (Kanmeraia) praeantiqua
Nassichuk and Wilde, 1977
Figures 5-15-26
Pseudofusulinella praeantiqua Nassichuk and Wilde,
1977, p.24, pI. 1, figs. 7-13.

Material studied.- Axial sections; ASM
25046, ASM25048, ASM25058, ASM25051,
ASM25052, ASM25056, ASM25050,
ASM25057, ASM25047 and ASM25049.
Sagittal sections; ASM25054 andASM25053. All specimens from Loc. BS200.
In addition to the illustrated specimens, several others were studied.
Description.-Shell small for the genus,
elongate fusiform in shape with bluntly
pointed polar ends and slightly concave lateral slopes. Mature specimens having 6 Yz to
7 volutions, measuring from 3.70 to 4.55 mm
in length and 1.38 to 1.93 mm in width.
Form ratio varying from 2.13 to 3.15, averaging 2.54 for II specimens.
Axis of coiling straight throughout. Innerfew volutions tightly coiled. Average radius
vectors of the first to seventh volutions for 12
specimens being 0.10,0.17,0.25,0.36,0.50,
0.70 and 0.89 mm. Average form ratios of
the first to seventh volutions for 10 specimens
being 1.61, 1.89,2.01,2.08,2.27,2.31 and 2.29,
respectively. Chamber heights almost constant throughout shell length.
Proloculus spherical and small. Its outside diameter ranging from 0.11 to 0.18 mm,
averaging O. 14 mm for 27 specimens.
Spirotheca thin and composed of a tectum,
diaphanotheca and lower thin and upper
thick tectoria related to dense deposits ofchomata. Thickness of tectoria usually inconstant and decreasing toward polar ends.
In some specimens, very fine perforations
observable in a diaphanotheca of outer volutions. A veiage thickness of spirotheca of thefirst to seventh volutions for 12 specimens
measuring 0.020, 0.025, 0.027, 0.031, 0.036,

819

0.040 and 0.035 mm, respectively.
Septa almost plane, but weakly fluted only
in extreme polar regions. Septal counts ofthe first to sixth volutions of typical sagittal
section (Figure 5-26) numbering 10, 14, 15,
17,20 and 21.
Tunnel path narrow and straight. A verage tunnel angles of the first to seventh volutions for 10 specimens measuring 9, 12, 13, 14,
17,21 and 21 degrees, respectively. Chomata
prominent and highly asymmetrical. Tunnelsides of chomata very steep, overhanging insome specimens. Outside slopes very gentle
and extending into polar regions. Chomata
reaching nearly roof of chamber in central
part of shell. Axial fillings absent or very
weakly developed in some specimens.
Remarks.- Pseudofusulinella (K anmeraia)
praeantiqua was originally described by
Nassichuk and Wilde (1977) from thePseudofusulina plana Assemblage Zone of
the Belcher Channel and Hare Fiord Formations in Ellesmere Island of Canada, which
they correlated with the Asselian. The
Akiyoshi specimens are quite referable to theoriginally described ones.
This species resembles somewhat
Pseudofusulinella (Kanmeraia) antiqua andP. (K.) meeki, which were described by Skinner and Wilde (1965) from the McCloud
Limestone of California. However, the former has a smaller shell than the latter two.
Pseudofusulinella (Kanmeraia) praeantiqua can be distinguished from P. (K.) japonica described by Ozawa (1967) from theRaidenyama Formation of the Kwanto
Mountains in having more concave lateralslopes, slightly larger shell and narrower tunnel path.
Occurrence.-Common in the uppermost
part of the Protriticites sp. Zone and rare in
the lower part of the Quasifusulinoides toriyamai Zone.

Pseudofusulinella (K anmeraia) cf. delicata
Skinner and Wilde, 1965

820

Katsumi U enG

922.

Carboniferous fusulinaceans from Akiyoshi

821

Figures 5-14

Quasifusulinoides Rauser-Chernoussova and Rozovskaya in Rauser-Chernoussova and Fursenko,
1959, p.21O.

Pseudofusulinella delicata Skinner and Wilde, 1965,
p. 28-29, pI. 8, figs. 23-28.

Remarks.- The genus Quasifusulinoides
closely resembles the genus Fusulina, but the.former is distinguished from the latter inhaving fine perforations in a diaphanotheca.
Moreover, the former occupies a slightly
younger stratigraphic position than the latter.

Compare.-Material studied.- Axial section; ASM
25055 from Loc. BS200.
Description.-Shell small for the genus andelongate fusiform in shape with bluntly
pointed poles. Lateral slopes straight.
Specimen having 7 volutions, measuring 5.07
mm in length and 1.70 mm in width, giving aform ratio of 2.98. Inner 3 volutions tightly
coiled. Radius vectors of the first to sixth
volutions being 0.08,0.13,0.21,0.30,0.47 and0.68 mm, and form ratios 2.00, 2.23, 2.73, 3.08,
2.79 and 2.72, respectively.
Spirotheca thin and composed of a tectum,
diaphanotheca and upper and lower tectoria.
Chomata prominent and highly asymmetrical. Tunnel angles of the second tosixth volutions measuring 17,18,19,27 and22 degrees, respectively.
Remarks.- The present species is characterized by a slender shell and is similar to
Pseudofusulinella (K anmeraia) delicata Skinner and Wilde. However, the former has aslightly larger shell and smaller form ratio
than the latter.
Occurrence.-Rare in the uppermost partof the Protdticites sp. Zone.
Subfamily Fusulininae von Moller, 1878
Genus Quasifusulinoides
Rauser-Chernoussova andRozovskaya in Rauser-Chernoussova
and Fursenko, 1959

Quasifusulinoides toriyamai
Ueno, sp. nov.
Figures 6-1-13

Material studied.-Axial section ofholotype; ASM25068 from Loc. BS218.6.
Axial sections of paratypes; ASM25059A
from Loc. BS201.5, ASM25060 from Loc.
BS212.5, ASM25071, ASM25074, ASM25072,
ASM25064, ASM25076 from Loc. BS218.6
and ASM25031 from Loc. US236. Slightly
oblique axial sections of paratypes;
ASM25061 from Loc. BS214 and ASM25069
from Loc. BS218.6. Sagittal sections of paratypes; ASM25075 and ASM25066 from Loc.
BS218.6. In addition, many other specimensfrom the Quasifusulinoides toriyamai Zone
were studied.
Description.-Shell medium for the genus
and fusiform in shape with bluntly pointed
polar regions. Mature specimens having 6 to
7 Yz volutions, measuring from 4.90 to 6.35mm in length and 2.03 to 2.80 mm in width.
Form ratio ranging from 2.25 to 2.57, averaging 2.40 for 9 specimens.
Inner I or I Yz volutions tightly coiled, but
outer ones becoming gradually expanded.
Axis of coiling straight throughout. A ver-+-- Figure 5. 1: Eostaffella sp. A, axial section, ASM25078A, X 100. 2: Eostaffella sp. B, axial
section, ASM25062, X 100. 3: Staffella pseudosphaeroidea Dutkevich, axial section, ASM25030, X 30. 411. Schubertella aff. kingi Dunbar and Skinner, 4-7, 10, 11: axial sections, ASM25067, ASM25073,
ASM25078B, ASM25070, ASM25079, ASM25059B, 8, 9: sagittal sections, ASM25077, ASM25065, X40.
12, 13: Schubertella sp., axial sections, ASM25063, ASM25080, X 40. 14: Pseudofusulinella (Kanmeraia)
cf. delicata Skinner and Wilde, axial section, ASM25055, X 15. 15-26. Pseudofusulinella (Kanmeraia)
praeantiqua Nassichuk and Wilde, 15-23a, 24: axial sections., ASM25058, ASM25057, ASM25046,
ASM25051, ASM25056, ASM25052, ASM25047, ASM25049, ASM25048, ASM25050, 25,26: sagittal sections, ASM25054, ASM25053, X 15, 23b: enlarged part of 23a, X 75.

822

Katsumi Veno

922.

Carboniferous fusulinaceans from Akiyoshi

823

Measurements of Ouasifusulinoides toriyamai, sp. nov. (in mm).

Table 3.

Radius vector

SpecimennumberFig.

Length

Width

F.R.

D.P.
1

2

3

4

5

6

7

1

ASM25074

6-3

6.50

2.80

2.32

0.24

0.24

0.35

0.49

0.68

0.89

1.15

1.40

2

ASM25064

6-4

6.13

2.58

2.38

0.24

0.20

0.24

0.35

0.51

0.70

1.96

1.28

-

3

ASM25031

6-7

4.90

2.03

2.42

0.19

0.13

0.20

0.29

0.42

0.58

0.80

4

ASM25060

6-8

6.30

2.45

2.57

0.31

0.23

0.32

0.44

0.59

0.78

0.98

1.23

5

ASM25072

6-10

6.15

2.40

2.56

0.24

0.18

0.28

0.41

0.56

0.80

1.02

-

6

ASM25068

6-11

6.32

2.70

2.34

0.35

0.27

0.38

0.50

0.72

0.97

1.25

-

7

ASM25059A

6-12

5.90

2.43

2.43

0.28

0.22

0.31

0.46

0.67

0.89

1.10

1.28*

8

ASM25076

6-13

6.35

2.68

2.37

0.30

0.21

0.33

0.47

0.68

0.99

1.25

1.38*

Form ratio

1

Thickness of spiro theca

1

2

3

4

5

6

7

1

2

3

4

5

6

7

1.29

1.42

1.68

1.65

1.65

1.72

1.88

0.020

0.025

0.030

0.030

0.040

0.045

0.040
0.060

2

1.33

2.13

2.31

2.38

2.38

2.37

2.41

0.020

0.025

0.025

0.030

0.040

0.Q45

3

1.77

1.65

2.00

2.32

2.46

2.41

-

0.020

0.020

0.D25

0.035

0.030

0.040

-

4

1.26

1.65

1.99

2.39

2.53

2.62

2.65

0.035

0.055

0.040

0.035

0.055

0.045

-

5

1.67

1.93

2.12

2.32

2.06

2.36

-

0.015

0.025

0.030

0.040

0.045

0.045

-

6

1.33

1.64

1.87

2.01

2.23

2.40

-

0.025

0.030

0.030

0.035

0.045

0.060

-

7

1.27

1.85

2.12

2.16

2.22

2.48

2.31*

0.020

0.025

0.D25

0.030

0.040

0.D25

0.020*

8

1.33

1.94

2.26

1.95

1.87

2.08

2.29*

0.020

0.025

0.030

0.045

0.055

0.020

-

6

7

Tunnel angle (degrees)
1

2

3

4

5

1

14

18

14

24

-

-

-

2

14

12

12

13

22

14

-

3

19

16

20

19

23

20

-

19

20

24

23

-

23

-

-

-

-

-

4
5

18
14

15
22

6

8

16

15

14

-

-

7

13

21

27

23

-

-

-

8

16

18

22

24

-

-

-

* : 6 ~ vo1ution<- Figure 6. 1-13. Quasifusulinoides toriyamai, sp. nov., 11: axial section ofholotype, ASM25068, 1,9:
sagittal sections of paratypes, ASM25075, ASM25066, 2-4,7,8, 10, 12, 13: axial sections of paratypes,
ASM25071, ASM25074, ASM25064, ASM25031, ASM25060, ASM25072, ASM25059A, ASM25076, Sa, 6:
slightly oblique axial sections of para types, ASM25069, ASM25061, X 10, 5b: enlarged part of Sa, showing
the peculiar spirothecal structure, X 100.

824

Katsumi Ueno

age radius vectors of the first to seventh volutions for 11 specimens being 0.21,0.30,0.42,
0.58, 0.80, 1.03 and 1.23 mm, and averageform ratios being 1.40,1.75,2.03,2.17,2.19,
2.25 and 2.22, respectively. Chamber heights
almost constant throughout shell length.
Proloculus spherical and medium to large.
Its outside diameter ranging from 0.19 to 0.46mm, averaging 0.28 mm for 56 specimens.
Spirotheca thin, composed of a tectum andupper and lower tectoria in inner 2 or 3
volutions, but becoming relatively thick andcomposed of a tectum, diaphanotheca andthin upper and rather thick lower tectoria inouter ones. Clear fine perforations sometimes observable in a diaphanotheca. Average thickness of spirotheca of the first to
seventh volutions for II specimens measuring0.022, 0.028, 0.030, 0.035, 0.043, 0.041 and
0.052 mm, respectively.
Septa intensely and regularly fluted
throughout length of shell, except for its
central part. Rudimentary chomata developed in inner 2 or 3 volutions. Tunnel path
almost straight. Average tunnel angles ofthe first to sixth volutions for II specimens
measuring 14,17,20,20,23 and 18 degrees,
respectively. Axial fillings prominent.
Remarks.- The present new species caneasily be distinguished from Quasifusulinoides
fusiformis (Rozovskaya), Q. parafusiformis
Bensh and Q. juvenatus Kireeva in having asmaller form ratio and the prominent axial
fillings.
Quasifusulinoides toriyamai, sp. nov. issomewhat similar to Q. turgida described by
Nikitina (1969) from the Obsoletes-Protritidtes Zone of the Tadushi River Basin ofsouthern Sikhote-Alin. However, the former
has a larger shell than the latter.
The present form also resembles Q. oblonga
Nikitina which occurred in association with
Q. turgida. However, the former can be
distinguished from the lat~er by a smaller
form ratio.
Fusulina kurikiensis and F. otanii described by Kanmera (1954) from the Fusulina

Zone of the Yayamadake Limestone in Kyushu somewhat resemble Quasifusulinoides
toriyamai, sp. nov. in the shell morphology,
but the former two species have a different
spirothecal composition from the latter.
Measurements.-See Table 3.
Occurrence.-Abundant and restricted in
the Quasifusulinoides toriyamai Zone.
Family Staffellidae A.D. MiklukhoMaklay, 1949
Genus Staffella Ozawa, 1925
Staffella pseudosphaeroidea Dutkevich, 1934
Figure 5-3

Staffella pseudosphaeroidea Dutkevich, 1934, p.1722, 66-68, pI. 3, figs. 2-10; Kanmera, 1954, p.
123-126, pI. 12, figs. 1-13; Chen and Wang,
1983, p. 34, pI. I, figs. 9-10.
Staffella ex gr. pseudosphaeroidea Dutkevich; Ginkel, 1965, p. 15-16, pI. 6, figs. 1-10.
Staffella cr. pseudosphaeroidea Dutkevich; Ginkel,
1965, p. 14-15, pI. 5, figs. 8-11.
Parastaffella pseudosphaeroidea (Dutkevich);
Rauser-Chernoussova et al., 1951, p. 152-153, pI.
13, figs. 1-2:
Pseudoendothyra pseudosphaeroidea (Dutkevich);
Grozdilova and Lebedeva, 1960, p. 105, pI. 12,
fig. 3; Kahler and Kahler, 1979, p.206-207, pI.
I, fig. 1.
Pseudostaffella? pseudosphaeroidea (Dutkevitch);
Ross and Dunbar, 1962, p. 15-17, pI. 2, figs. 1-8.

Material studied.- Axial section; ASM
25030 from Loc. US211.
Description.-Shell spherical, relatively
large for the genus and having 5 Yz volutions.
Axial regions slightly depressed. Length
1.29 mm and width 1.39 mm, giving a form
ratio of 0.93. Shell loosely coiled. Radius
vectors of the first to fifth volutions being
0.10,0.17,0.27,0.41 and 0.58mm, and form
ratios 0.64, 0.76,0.89, 1.02 and 1.01, respectively.
Proloculus spherical and 0.11 mm in outside diameter. Spirotheca highly recrystallized and probably composed of a thintectum, thick diaphanotheca and upper andlower tectoria. Thickness of spirotheca of
the second to fifth volutions measuring 0.02,

922.

Carboniferous fusulinaceans from Akiyoshi

0.03,0.06 and 0.05 mm, respectively. Septa
not fluted. Chomata present but inconspicuous. Axial fillings absent.
Remarks.- The present specimen is quite
identical to Dutkevich's original ones in theessential shell characters.
Staffella pseudosphaeroidea described by
Kanmera (1954) from the Yayamadake Limestone of Kyushu and S. ex gr. pseudosphaeroidea described by Ginkel (1965) from the
Cantabrian Mountains of Spain have a more
quadrate shell than that of the Akiyoshi specimen.
Occurrence.-Rare in the lower part of theProtriticites sp. Zone.

Acknowledgments
I would like to express my appreciation toProf. Sumio Sakagami of Chiba University
for reading the manuscript of this paper. I
am much indebted to Messrs. Akihiro
Sugimura and Takehiko Haikawa of Akiyoshi-dai Museum of Natural History for
their kind permission to examine thin sections deposited in the museum and giving
warm encouragement, and to Mr. Yasuyuki
Miyama of Sumitomo Cement Co., Ltd. forfruitful discussions on the fusulinacean taxonomy.

References cited
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Vdovenko, M.V., Gorak, S.V., Dunaeva, N.N.,
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The

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Protriticites, Quasifusulinoides i Obsoletes iz pogranichnykh otlozheniy srednego i verkhnego825karbona. Voprosy Mikropaleont., vol. 7, p.7184, pIs. 1-6. (in Russian)Chen, X. and Wang, J.H., 1983: The fusulinids of theMaping Limestone of the Upper Carboniferous
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Dunbar, C.O. and Skinner, J.W., 1937: The Geologyof Texas, Volume III, Part 2. Permian
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Dutkevich, G.A., 1934: Novye vidykh fuzulinid iz
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Grozdilova, L.P. and Lebedeva, N.S., 1960: Foraminifery kamennougol'nykh otlozheniy zapadnogo sklona Urala i Timana. Trudy Vses. Neft.

Nauchno-Issled. Geol.-Razved. Inst. (VNIGRl),vol. 150, p. 1-264, pis. 1-33. (in Russian)Han, J.X., 1976: Fusulinida. In, Geol. Bureau,
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Carboniferous strata and its fauna from southwestern margin of Tarim basin in Xinjiang, p.
55-95, pIs. 1-14. Geol. Publ. House, Beijing. (inChinese with English abstract)
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Tokyo Kyoiku Daigaku, Sec. C, vol. 5, nos. 4748, p. 153-246, pIs. 1-15.
- - , Rajah, S.S. and Kobayashi, F., 1979: Permian
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Kahler, F. and Kahler, G., 1979: Fusuliniden (Foraminifera) aus dem Karbon und Perm vonWestanatolien und dem Iran. Mitt. Osterr. Geol.
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Kanmera, K., 1954: Fusulinids from the Yayamadake Limestone of the Hikawa Valley, Kumamoto
Prefecture, Kyushu, Japan, Part I. Fusulinids
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Jour. Geol. Geogr., vol. 25, nos. 1-2, p. 117-144,
pIs. 12-14.

826

Katsumi Ueno

Kauffman, A.E. and Roth, R.I., 1966: Upper Pennsylvanian and Lower Permian fusulinids from
north-central Texas. Cushman Found. Foram.
Res., Spec. Publ., no. 8, p. 1-49, pis. 1-11.
Kireeva, G.D., 1949: Nekotorye novye vidy fuzulinid
iz kamennougol'nykh izvestnyakov tsentral'nogo
rayona Donbassa. Trudy Geologo-issledovatel'skogo Byuro, Glavnoe Upravlenie po Razvedkam
Uglya, Ministerstvo Ugol'noy Promyshlennosti
SSSR, vol. 6, p. 25-55, pis. 1-2. (in Russian)- - , 1950: Novye vidy fuzulinid iz izvestnyakov svit
C! i C~ Donetskogo basseyna. In, Materialy po
stratigrafii i paleontologii Donetskogo Basseyna,
p. 193-214, pis. 1-4. Geologo-issledovatel'skie
Raboty, Glavnoe Upravlenie po Razved kam
Ugly a, Ministerstvo Ugol'noy Promyshlennosti
SSSR. (in Russian)
Kochansky-Devide, V., 1959: Die Fusuliniden Foraminiferen aus dem Karbon und Perm im Velebit
und in der Lika (Kroatien). Unteres Perm.
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Lee, J.S., 1927: Fusulinidae of North China.
Palaeont. Sinica, vol. 4, no. I, p. 1-173, pis. 1-24.
Leven, E.Ya. and Scherbovich, S.F., 1978:
Fuzulinidy i stratigrafiya asselyskogo yarusa
Darvaza. 162 p., 21 pis., Izdatel. <Nauka>, Moskva. (in Russian)Loeblich, A.R., Jr. and Tappan, H., 1988: Foraminiferal genera and their classification. 970
p., 847 pis., Van Nostrand Reinhold Co., NewYork.
Mudge, M.R. and Yochelson, E.L., 1962: Stratigraphy and paleontology of the uppermost Pennsylvanian and lowermost Permian rocks in Kansas. U.S. Geol. Surv., Prof Paper 323, p. 1-213,
pis. 6-17.
Nassichuk, W.W. and Wilde, G.L., 1977: Permian
fusulinaceans and stratigraphy at Blind Fiord,southwestern Ellesmere Island. Geol. Surv.
Canada, Bull. 268, p. I-59, pis. 1-5.
Niikawa, I., 1978: Carboniferous and Permian
fusulinids from Fukuji, central Japan. Jour.
Fac. Sci., Hokkaido Univ., ser, IV, vol. 18, no. 4,
p. 533-610, pis. 1-14.
Nikitina, A.P., 1969: Pervye predstaviteli Obsoletes,
Protriticites i Fusulina iz verkhnego karbona
Primorskogo kraya. Paleont. Zhurnal, 1969,
no. 4, p. 3-11, pis. 1-2. (in Russian)
Niko, S. and Watanabe, A., 1987: Late Carboniferous and Early Permian fusulinids from Kuzuryu
Lake district, Fukui Prefecture (Part I, Ozawainellidae; Schubertellidae; Fusulinidae).
Sci. Pap., Coli. Art. Sci., Univ. Tokyo, vol. 37,
no. I, p.37-58.
Ota, M., 1968: The Akiyoshi Limestone Group: A
geosynclinal organic reef complex. Bull. Akiyoshi-dai Sci. Mus., no. 5, p. 1-44, pis. 1-31. (inJapanese with English abstract)- - , 1977: Geological studies of Akiyoshi, Part I.
General Geology of the Akiyoshi LimestoneGroup. Bull. Akiyoshi-dai Mus. Nat. Hist., no.
12, p. 1-33, pis. 1-3.
- - , Toriyama, R., Sugimura, A. and Haikawa, T.,
1973: Restudy on the geologic structure of the
Akiyoshi Limestone Group, southwest Japan.
Japan. Jour. Geol. Geogr., vol. 82, no. 3, p. 1\5135. (in Japanese with English abstract)
Ozawa, T., 1967: Pseudofusulinella, a genus of
Fusulinacea. Trans. Proc. Palaeont. Soc.
Japan, N.S., no. 68, p. 149-173, pis. 14-15.
- - and Kobayashi, F., 1990: Carboniferous toPermian Akiyoshi Limestone Group. In, Organizing Committee Benthos '90 ed., Fossil andRecent benthic foraminifera in some selected
regions of Japan. Guidebook for field trips,
4th International Symposium on Benthic Foraminifera, Sendai, 1990, p. EI-31, pis. 1-13.
Pasini, M., 1965: Fusulinidi, una chiave analitica perla determinazione del generi. 107 p., 18 pis.
Palaeontogr. Italica, Pisa.
Putrja, F.S., 1948: Protriticites-Novyy rod
fuzulinid. Trudy L'vovsk. Geol. Obsche. pri
Gosud. Univ. im Ivana Franko, ser. Paleont., vol.
I, p. 89-96, pI. 1. (in Russian)Rauser-Chernoussova, D.M. and Fursenko, A.V.,
1959: Osnovy paleontologii, Obschaia chast',
Prosteyshie. 482 p., 13 pis. Izdatel. Akad. Nauk
SSSR, Moskva. (in Russian)- - , Ivanova, E.A., Grozdilova, L.P. and Makhlina,
M. Kh., 1979: The Upper Carboniferous series.
In, Wagner, R.H., Higgins, A.c. and Meyen, S.V.
eds., The Carboniferous of the U.S.S.R., Yorkshire Geol. Soc., Occ. Publ., no. 4, p. 147-174.
- - , Kireeva, G.D., Leontovich, G.E., Gryzlova, N.
D., Safonova, T.P. and Chern ova, E.I., 1951:
Srednekamennougol'nye fusulinidy Russkoy
Platformy i sopredel'nykh oblastey. 380 p., 58
pis. Akad. Nauk SSSR, Inst. Geol. Nauk, Minist.
Neft. Promyshlennosti SSSR, Izdatel. Akad.
Nauk SSSR, Moskva. (in Russian)Ross, C.A. and Dunbar, C.O., 1962: Faunas andcorrelation of the Late Paleozoic rocks of N ortheast Greenland, Part II. Fusulinidae. Medd.
Grq,nland, vol. 167, no. 5, p.I-55, pis. 1-7.
Rozovskaya, S.E., 1950: Rod Triticites, ego razvitie istratigraficheskoe znachenie. A kad. Nauk
SSSR, Trudy Paleont. Inst., vol. 26, p. 1-78, pis.
1-10. (in Russian)- - , 1975: Sostav, sistema i filogeniya otryada
Fuzulinida. Ibid., vol. 149, p. 1-227, pis. 1-35.
(in Russian)
Sabins, F.F., Jr. and Ross, C.A., 1963: Late
Pennsylvanian-Early Permian fusulinids fromsoutheast Arizona. Jour. Paleont., vol. 37, no.

922.

Carboniferous fusulinaceans from Akiyoshi

2, p. 323-365, pis. 35-40.
Sakagami, S. and Ornata, T., 1957: Lower Permian
fusulinids from Shiraiwa, northwestern part ofOrne, Nishitama-gun, Tokyo-to, Japan. Japan.
Jour. Geol. Geogr., vol. 28, no. 4, p. 247-264, pis.
19-20.
Schellwien, E., 1908: Monographie der Fusulinen,
Teil I. Die Fusulinen des russisch-arktischen
Meeresgebietes. Palaeontogr., vol. 55, p. 145194, pis. 13-20.
Sheng, J.Z. (Sheng, J.e.), 1958: Fusulinids from thePenchi Series of the Taitzeho valley, Liaoning.
Palaeont. Sinica, N.S.B, no. 7, p. 1-119, pis. 116.
- - and Wang, R.N., 1984: Fusulinid-bearing formations and fusulinid fauna from the Dadun
coalfield of Pei Xian, northern Jiangsu. Bull.
Nanjing Inst., Geol. Palaeont., A cad. Sinica, no.
7, p. 1-68, pis. 1-10. (in Chinese with English
abstract)
Skinner, J.W. and Wilde, G.L., 1965: Permian biostratigraphy and fusulinid faunas of the Shasta
Lake area, northern California. Univ. Kansas,
Paleont. Contr., no. 39 (Protozoa, art. 6), p. 1-98,
pis. 1-65.
Stewart, W.J., 1966: Schubertellinae volfkempa (nizhnyaya permi) iz gor Franklina Tekhasa. Voprosy Mikropaleont., vol. 10, p.80-88, pis. 1-2.
(in Russian)- - , 1968: Schubertellinae from the Wolfcamp,
Lower Permian, Franklin Mountains, Texas.

827

Jour. Paleont., vol. 42, no. 2, p. 322-328, pis. 4142.
Suleimanov, I.S., 1949: Novye vidy fuzulinid podsemeystva Schubertellinae Skinner iz kamennougol'nykh i nizhnepermskikh otlozheniy k
Bashkirskogo Priural'ya. Akad. Nauk SSSR,
Trudy Inst. Geol. Nauk, vol. 105, GeoL ser. (no.
35), p. 22-43, pI. 1. (in Russian)Suyari, K., 1962: Geological and paleontologicalstudies in central and eastern Shikoku, Japan,
Part 2. Paleontology. Jour. Gakugei, Tokushima Un iv, vol. 12, p. 1-64, pis. 1-12.
Thompson, M.L., 1948: Studies of American
Fusulinids. Univ. Kansas, Paleont. Contr., no.
4 (Protozoa, art. 1), p. 1-184, pis. 1-38.
- - , 1954: American Wolfcampian fusulinids.
Ibid., no. 14 (Protozoa, art. 5), p. 1-226, pis. 152.
- - , Wheeler, H.E. and Hazzard, J.e., 1946: Permian fusulinids of California. Geol. Soc. America, Mem. 17, p. 1-77, pis. 1-18.
Toriyama, R., 1958: Geology of Akiyoshi, Part III.
Fusulinids of Akiyoshi. Mem. Fac. Sci., Kyushu Univ., ser. D, vol. 7, p. 1-264, pis. 1-48.
Ueno, K., 1989: Carboniferous and Lower Permian
foraminiferal biostratigraphy in the Akiyoshi
Limestone Group. (Studies of the Upper Palaeozoic foraminifers in the Akiyoshi LimestoneGroup, southwest Japan, Part I). Bull. Akiyoshi-dai Mus. Nat. Hist., no. 24, p. 1-39, pI. 1-8.
(in Japanese with English abstract)

Akago *'~. Akiyoshi f:k-a, Akiyoshi-dai f:k-a-El'. Kyushu flJli, Kwanto ~*. Managatake
Ji;i5 ?- ffi. Raidenyama ~~0.!, Yamaguchi 0.! I=l. Yayamadake ~0.!ffi.
f:k-a;o19(;fiIi~tJ, G(j) J:1iB;ohR**-O~Ea~: f:k-a-El'~t*1iB(j)*,~-Ji;i5 ?-ffi:l:t!!~t;:.5t;:fp
1" Q f:k-a;o 19(;fi Ii fF (j) Protriticites sp. c Quasifusulinoides toriyamai
IJ. 3 *'T~**.t(Protriticites robustus. Obsoletes horridus. Quasifusulinoides toriyamai) ~~tr 7 ~ 12 ~(j)i1i!i~Ea~~~1t1" Q(j)iIi!i~EafF~lt Protriticites. Obsoletes. Quasifusulinoides (j) 3 ~ ~
0

*'

;:

:1 -~rPJ~.
r j1 &:l:t!!. ~1iB 7 Iv 7fT. i'&ria1'rl
JII&:l:t!!.t IJ £1" QiIi!i~Ea~~ c ~.RX;t;:.:to~'-C~~,~1tl1.l~~1" -t(j)Ilif~lt;0 hR$c.f&M (j) iltrM Kasimovian c ~ ;t G.n Q
J:!I!J/l9J~~tr;:c -c-~il"::)Vt G.n -C:to IJ. -'E- A(j)!J r ? : /

:L

0

0

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 163, pp. 828-851, 5 Figs., September 30, 1991

923. CAMPANIAN INTERMEDIATE WATER BENTHICFORAMINIFERA FROM CENTRAL HOKKAIDO, JAPAN*
KUNIO KAIHO
Institute of Geology and Paleontology,
Tohoku University, Sendai 980

Abstract. Although Late Cretaceous sublittoral and bathyal mudstones are widelydistributed in central Hokkaido, Japan and contain abundant calcareous microfossils,
studies on benthic formaminifera have been few. I obtained many well-preserved foraminifera numbering about 50,000 specimens from two 10 kg-samples from the CampanianKashima Formation of the Yezo Group. This study presents a description of benthic
foraminiferal faunas that include three new species and clarifies a faunal composition in theCampanian intermediate water. The benthic foraminiferal faunas are dominated by
Silicosigmoilina Jutabaensis Asano (85-90%), Notoplanulina rumoiensis Takayanagi, Nuttallinella florealis (White), Haplophragmoides asanoi Takayanagi, and Dorothia hokkaidoana
Takayanagi. They are also characterized by a high species diversity of the suborder
Lagenina (66% Lagenina species) including such genera as Len ticulin a, Frondicularia,
Saracen aria and Astacolus, and by a low species diversity of the suborder Rotaliina (9%Rotaliina species). These faunas indicate an upper bathyal depositional water depth.
Key words.
kaido, Japan.

Benthic foraminifera, Campanian, Cretaceous, intermediate water, Hok-

the Campanian intermediate water depth.
This is the first description of Campanian
benthic foraminifera from Hokkaido, Japan,
with the exception of a benthic foraminiferal
study of northern Hokkaido done by Yasuda(1986). All the specimens described herein
are deposited in the collections of Institute ofGeology and Paleontology, Tohoku University, Sendai (IGPS) with their repository
numbers being from IGPS 101470 to 101539.
The planktonic foraminifera and Ostracoda
will be reported elsewhere by Kaiho (in press,a) and Ishizaki (in press), respectively.

Introduction
Late Cretaceous mudstones deposited in
the intermediate water depth are widely distributed in central Hokkaido, Japan. These
strata contain abundant calcareous microfossils. Takayanagi (1960) described foraminifera from the Late Cretaceous formations of central Hokkaido. As most of his
specimens are not well preserved, I tried toobtain well-preserved foraminifera by washing a large quantity of samples from theCampanian mudstone of the Kashima Formation (Motoyama et aI., 1991).
'The purpose of this paper is to describe
these well-preserved benthic foraminiferal
species and to clarify faunal compositions inGeological setting
I selected the Sanushube River section for astudy of calcareous microfossils out of various sections measured in Hokkaido. The
section is located in the northwestern part of"Received May 17, 1991 ; revised manuscript accepted
August 10, 1991
828

923.

Campanian benthic foraminifera from central Hokkaido

Hobetsucho, south-central Hokkaido (Figure1). The upper part of the thick marine
Upper Cretaceous sediments developed along
this section is stratigraphically separated as
the Kashima Formation, which represents theuppermost part of the Yezo Group. The
Kashima Formation consists of a dark graymassive soft mudstone which contains Inoceramus shell fragments, foraminifera, Radiolaria, calcareous nannofossils, ostracodes, ammonites and plesiosaurian fossils (N akaya,1989). It is conformably overlain by shallowo829

marine sediments of the Hakobuchi Group,characterized by sandstone, alternations ofsiltstone and sandstone, and coal measures,that contain rare fossils. The Kashima Formation would have been about 1000 m inthickness if it were not disturbed by faulting,but, in reality, may be separated and overlapped by faults (Kito et al., 1986). From
the upper half of the Kashima Formation, I
collected studied samples. Kito et al. (1986)
assigned an early Campanian age to the
middle part of the Kashima Formation (sam-

lkm
,

I

142'1'30"

4iss... :.

~:;"A
\
\

\ KS

HK4\

SNS08

H
OJ

:>

'rl

p:;

...:j

"

N

OJ
.0

p::

;:l

...c:
Ul
;:l

I=:

ru

U)

~

HOBETSU

Figure 1. Location map of the Sanushube River section and sample localities. KS: Kashima Formation; HK: Hakobuchi Group; PR: Poronai Formation (upper Middle-Upper Eocene); ms: mudstone;
ss: sandstone; alt: alternation of siltstone and sandstone; F: fault.

Kunio Kaiho

830

pIe SNSOI : plesiosaurian localilty) based onradiolarians, planktonic foraminifera, ammonites and Inoceramus. Kaiho (in press, a)assigned a Campanian age to the upper halfof the Kashima Formation (samples SNS02
and SNS07) exposed along the studied section. Calcareous nannofossils from theuppermost part of the Kashima Formation(sample SNS08) from the same section also
show a Campanian age (Motoyama et al.,1991 ).
Materials and methods
Although I obtained in 1979 well-preserved
calcareous foraminifers from five localities
along this section, the number of specimens
was not sufficient for detailed studies.
Accordingly, I collected a large quantity ofsample (about 10 kg) from each of the two
localities (SNS02 and SNS07) (Figure 1) in
the following year.
In the laboratory, 10 kg of dry sedimentwere disaggregated by using a sodium sulphate and naphtha solution (Maiya andInoue, 1973), wet sieved through a 63-J.lm
screen and dried again. Foraminifera werethen picked up from the processed samples
and identified. The type specimens of Takayanagi (1960) were examined for comparative
studies. All the identified species are shownin the accompanying plates. These species
were also illustrated in Kaiho (1990) without
detailed descriptions.
Benthic foraminiferal fauna anddepositional water depth
The benthic foraminiferal fauna in the two
samples SNS02 and SNS07 show a very similar specific composition (Table 1). I recognized 47 benthic foraminiferal species inabout 50,000 specimens from these two samples. Of these species, 55% of them are discovered for the first time from Japan. Three
species are new species. Silicosigmoilina

Jutabaensis Asano occupies 85-90% of thetotal specimens. Other abundant taxa
include Notoplanulina rumoiensis (Takayanagi), Nuttallinella florealis (White), Haplophragmoides asanoi Takayanagi and Dorothia hokkaidoana Takayanagi. The benthic foraminiferal faunas are also characterized
by a high species diversity of the suborder
Lagenina, comprISing such genera as
Lenticulina, Frondicularia, Saracen aria, andAstacolus, and a low species diversity of thesuborder Rotaliina, represented by 66%
Lagenina, 9% Rotaliina, and 25% Textulariina species. Based on the bathymetric distribution of Cretaceous benthic foraminifera
reconstructed by Sliter and Baker (1972) withdata from southern California, most of thegenera occurring in the Sanushube section areinterpreted to have inhabited the upperbathyal zone.
Systematic description
In this section, I describe 47 species including three new species which were encounteredduring the course of present study. Amongthese, six tentatively identified species are
given their names in alphabetic letters.
Although some of these species may represent
a new species, no scientific name is given tothem pending examination of additional specimens in order to establish their range of
vanatlOn. At the suprageneric levels, thepresent classification follows the scheme
proposed by Loeblich and Tappan (1988).
Order Foraminiferida Eichwald, 1830
Suborder Textulariina Delage
and Herouard, 1896
Superfamily Astrorhizacea Brady, 1881
Family Bathysiphonidae Avnimelech, 1952
Genus Bathysiphon M. Sars, 1872
Bathysiphon vitta Nauss
Figure 2-1

Bathysiphon vitta Nauss, 1947, p.334, pI. 48, fig. 4;
Takayanagi, 1960, p. 64, 65, pI. 1, figs. Sa, b;

923.

Campanian benthic foraminifera from central Hokkaido

Table 1.

Abundance of benthic foraminifera from two Campanian samples.
Each sample weighs about 10 kg in dry weight. Number of specimens shown by the following symbols. VA: > 10,000; A: 1001000; C: 10-99; F: 3-9; R: 1-2; - : barren.
SampleSpecies
Bathysiphon vitta
Rhizammina indivisa
Ammodiscus sp. A
Glomospira charoides
Glomospira gordialis
Silicosigmoilina Jutabaensis
Haplophragmoides asanoi
Ammobaculites sp.
Cyclammina asanoi
Gaudryina pyramidata
Dorothia hokkaidoana
Marssonella sp. A
Dentalina basiplanata
Dentalina gracilis
Nodosaria affinis
Nodosaria aspera
Frondicularia bicorsis
Frondicularia durrelli
Frondicularia mucronata
Frondicularia whaingaroica
Frondicularia sp. A
Lenticulina extruatus
Lenticulina haboroensis
Lenticulina obirashibensis
Lenticulina rotulata
Lenticulina williamsoni
Lenticulina yabei
Lenticulina sp. A
Lenticulina sp. B
Lenticulina sp. C
Saracen aria triangularis
Saracen aria tunesiana
Pravoslavlevia sp.
Astacolus elongatus, n. sp.
Astacolus nagaoi
Astacolus polandensis
Marginulina bullata
Citharina multicostata
Psilocitharella ezoensis, n. sp.
Psilocitharella triangulata, n. sp.
Globulina lacrima
Ramulina aculeata
Ramulina tetrahedralis
Stilostomella alexanderi
Stilostomella stephensoni
Nuttallinella jiorealis
Notoplanulina rumoiensis

SNS02

SNS07

F
F
C
F
R
VA
A

C
F
C
-

-

C
F
A
-

F
C
R
R
C
F
-

C
-

C
C
R
C
-

F
C
C
C
C
R
R
-

F
-

R
C
F
F
C
C
A

-

VA
A
R
C
-

A
R
C
R
-

-

C
F
C
R
R
C
F
C
C
C
R
R
F
C
C
R
C
C
C
R
C
F
R
C
F
C
C
A
A

831

832

Kunia Kaiha

923.

Campanian benthic foraminifera from central Hokkaido

Trujillo, 1960, p.302-303. pI. 43, figs. 2a-b ;Tappan, 1962, p. 128-129, pI. 29, figs. 6-8; Graham and Church, 1963, p. 17-18, pI. 1, figs. la2b; Sliter, 1968, p.40,41, pI. 1, figs. 13a-15b.

Type.-Hypotype, IGPS 101470, sample
SNS07.
Description.- Test large, elongate, tubular,
compressed, with transverse growth constrictions, but no internal partitions; wall finely
agglutinated, with considerable amount of
cement, surface rather smoothly finished, grayto white in color; aperture at open end of
tube.
Remarks.- This finely walled large species
is common in the Upper Cretaceous of Japanand has a cosmopolitan distribution in
Cretaceous oceans.

Family Rhabdamminidae Brady, 1884
Subfamily Rhabdammininae Brady, 1884
Genus Rhizammina Brady, 1876
Rhizammina indivisa Brady
Figures 2-2,3Rhizammina indivisa Brady, 1884, p.277. pI. 29, figs.
5-7; Sliter, 1980, p.368, pI. 1, fig. 1 ; Yasuda,
1986, p. 40, 41, pI. 1, fig. 4.

Types.-Hypotype, figure 2-2, IGPS
101471; hypotype, figure 2-3, IGPS 101472.
Both from sample SNS02.
Description.- Test elongate, tubular,
irregulary curved, with transverse growth constriction; wall coarse, thin, made of firmly
cemented coarse sand grains, surface roughly
finished; aperture terminal and rounded.
Remarks.- This species has a wall and test
diameter similar to those of Bathysiphon
akanosawaensis Takayanagi.

833

Superfamily Ammodiscacea Reuss, 1862
Family Ammodiscidae Reuss, 1862
Subfamily Ammodiscinae Reuss, 1862
Genus Ammodiscus Reuss, 1862
Ammodiscus sp. A
Figure 2-4Type.-Hypotype, IGPS 101473, sample
SNS07.
Description.- Test large, planispiral, closely coiled, evolute, very compressed, peripheryrounded; chamber increasing gradually anduniformly in size as added with many coils;
spiral suture distinct, deep; wall coarsely
arenaceous, surface rather roughly finished;
aperture formed at open end of tube.
Remarks.- The diameter of the figured
hypotype is 0.96 mm and its thickness 0.11
mm. This species has a test smaller than thatof A. pennyi Cushman and Jarvis. It differsfrom A. cretaceous in having a coarser wall.
Subfamily Ammovertellinae Saidova, 1981
Genus Glomospira Rzehak, 1885
Glomospira charoides (Jones and Parker)
Trochammina squamata Jones and Parker var. charoides Jones and Parker, 1860, p. 304.
Trochammina charoides Jones and Parker. Carpenter, 1862, p. 141, pI. 11, fig. 3.
Ammodiscus charoides (Jones and Parker). Brady,
1884, p. 334, pI. 38, figs. 10-16.
Glomospira charoides (Jones and Parker). Cushman, 1925, p. 25, pI. 2, fig. 12; Galloway andMorrey, 1931, p.331, pI. 37, figs. 1-2; Marie,
1937, p. 260; Gandolfi, 1942, p. 31; Noth, 1951,
p. 28, pI. 2, fig. 7; Takayanagi, 1960, p. 69, pI. 1,
figs. 17a-18b.
Gordiammina charoides (Jones and Parker). Franke, 1928, p. 15, pI. 1, fig. 16.

Type.-Hypotype, IGPS 101474, sample
SNS02.

~ Figure 2. Scale bar= 100 }Lm. 1, Bathysiphon vitta Nauss. SNS07. 2,.3, Rhiz~m.mina indivisa
Brady. 2; SNS02, 3; SNS07. 4, Ammodiscus sp. A. SNS07. 5, Glomospzra gordzalzs (Jones andParker). SNS02. 6-9, Silicosigmoilina Jutabaensis Asano. 6-8; SNS07. 9; S.NS02. 10,11, Haplophragmoides asanoi Takayanagi. 10; SNS07. 11; SNS02. 12, Ammobaculztes sp. SNS07. 13,
Cyclammina asanoi Takayanagi. SNS07. 14, Gaudryin~ pyramidata Cushman. SNS02. 15, Marssonella
sp. A, SNS07. 16,17, Dorothia hokkaidoana Takayanagl. 16; SNS07. 17; SNS02.

Kunio Kaiho

834

Remarks.- Three juvenile specimens arefound in sample SNS02. It is a cosmopolitan, long-ranging species.
Glomospira gordialis(Jones and Parker)
Figure 2-5

Trochammina squamata gordialis Jones and Parker,
1860, p. 304; Brady, 1876, p. 77, pI. 3, figs. 1-3.
Ammodiscus gordialis Jones and Parker [sic].
Brady, 1884, p. 333, pI. 38, figs. 7-9.

Type.-Hypotype, IGPS 101475, sample
SNS02.
Description.- Test convolute, compressed,
oval, asymmetrical, composed of a tube ofnearly even diameter, and exhibiting low
trochospiral and twisted coiling arrangement; wall very finely arenaceous, with muchcement, surface smooth, colored in white;
aperture formed at open end of tube.
Remarks.-A single juvenile specimen isrecognized in sample SNS02.
Superfamily Rzehakinacea Cushman, 1933
Family Rzehakinidae Cushman, 1933
Genus Silicosigmoilina Cushman
and Church, 1929
Silicosigmoilina futabaensis Asano
Figures 2-6-9

Silicosigmoilina futabaensis Asano, 1950, p. 159, pI. I,
figs. 6, 7; Trujillo, 1960, p. 303, pI. 44, figs. 8a, b.
Bramletteia ezoensis Takayanagi, 1960, p.84, pI. 3,
figs. 14a-17.

Types.-Hypotype, figure 2-6, IGPS
101476, sample SNS07; hypotype, figure 2-7,
IGPS 101477, sample SNS07; hypotype,figure 2-8, IGPS 101478, sample SNS07;
hypotype, figure 2-9, IGPS 101479, sample
SNS02.
Description.- Test compressed, oval toelliptical in outline, sigmoid in end view;
periphery broadly rounded, edge subrounded
to subacute; chambers planispiral in earlystage, sigmoidal in adult stage; wall finely
arenaceous, with much siliceous cement,smoothly finished, white in color; suturesgenerally indistinct; aperture simple, oval,
terminal.
Remarks.- This species resembles Silicosigmoilina californica Cushman and Church
described from California and from Maastrichtian and Danian intervals of Hokkaido,
but differs in having a more flattened andsmaller test. Silicosigmoilina futabaensis is
the most dominant species in the Campanianintermediate water in Hokkaido.
Superfamily Lituolacea
de Blainville, 1827
Family Haplophragmoididae
Mayne, 1952
Genus Haplophragmoides
Cushman, 1910
H aplophragmoides asanoi
Takayanagi
Figures 2-10, 11

Haplophragmoides asanoi Takayanagi, 1960, p.6970, pI. I, figs. 19a-b.

Types.-Hypotype, figure 2-10, IGPS
101480, sample SNS07; hypotype, figure 211, IGPS 101481, sample SNS02.
Description.- Test compressed, planispiral,
involute, biumbilicate, nearly circular in outline, nearly parallel in side view, peripheral
margin broadly rounded, peripheral outline
commonly lobulate; six to seven chambers infinal whorl, slightly inflated, increasing gradually in size as added; sutures straight,
radial, slightly depressed; wall coarsely agglutinated, surface not smoothly finished, pale
brown in color; aperture equatorial, a lowinteriomarginal arch.
Remarks.- This species resembles both
Haplophragmoides incognatus Martin and
Haplophragmoides fraseri Wickenden ofTrujillo (1960), but differs in having fewer numbers of chamber. It resembles Haplophragmoides rugosus Cushman and Waters, butdiffers in having finer wall surface and chambers that increase more slowly in size.

923.

Campanian benthic foraminifera from central Hokkaido

835

Family Lituolidae de Blainville, 1827
Subfamily Ammomarginulininae
Podobina, 1978
Genus Ammobaculites Cushman, 1910
Ammobaculites sp.

Superfamily Verneuilinacea Cushman, 1911
Family Verneuilinidae Cushman, 1911
Subfamily Verneuilininae Cushman, 1911
Genus Gaudryina d'Orbigny, 1839
Gaudryina pyramidata Cushman

Figure 2-12

Figure 2-14

Type.-Hypotype, IGPS 101482, sample
SNS07.
Description.- Test elongate, compressed,
early portion close-coiled, with five chambers
per coil, later portion with a few uniseriallyarranged chambers, nearly parallel sided,
peripheral margin subangular; chambers
inflated, approximately twice as broad asheight in uniserial portion; sutures depressed, radial in coiling portion, right angle
to test axis in uniserial portion; wall coarsely
agglutinated, surface roughly finished; aperture terminal, ovate, centrally located.

Gaudryina laevigata Franke var. pyramidata Cushman, 1926, p. 587, pI. 16, fig. 8.
Gaudryina (Pseudogaudryina) pyramidata Cushman.
Cushman and Goudkoff, 1944, p. 56, pI. 9, figs. 7,

Superfamily Loftusiacea Brady, 1884
Family Cyclamminidae Marie, 1941
Subfamily Cyclammininae
Marie, 1941
Genus Cyclammina Brady, 1879
Cyclammina asanoi Takayanagi
Figure 2-13

Cyclammina asanoi Takayanagi, 1960, p. 75, 76, pI.
II, figs. Ila, b, 16.

Type.-Hypotype, IGPS 101483, sample
SNS07.
Description.- Test medium in size, muchcompressed, planispiral, involute, biumbilicate, periphery subacute, lobulate; chambers seven to eight in the final whorl; sutures
distinct, slightly depressed, radial, slightly
curved or slightly sigmoidal; wall finely
arenaceous, smoothly finished, interior labyrinthic; aperture a curved slit at base of apertural face.
Remarks.- This species is characterized
by its lobulate peripheral outline and inhaving seven to eight chambers in the final
whorl.

8.

Gaudryina pyramidata Cushman. Trujillo, 1960, p.
308, pI. 44, figs. 9a-c; Graham and Clark, 1961,
p. 109, fig. 2, I ; Sliter, 1968, p. 48,49, pI. 3, fig. 9.

Type.-Hypotype, IGPS 101484, sample
SNS02.
Description.- Test elongate, triangular intransverse section, early stage triserial, later
becoming biserial; chambers distinct, low,increasing gradually in size, slightly inflated
in biserial portion; sutures oblique, slightly
curved, depressed; wall finely agglutinated,surface smooth, white to pale brown in color
with black minerals; aperture, a low interiomarginal arch.
Remarks.- This species is similar to Gaudryina sp. a described by Takayanagi (1960)
from the Cretaceous of Hokkaido, but thelatter is too poorly preserved to establish thepositive identification.
Superfamily Textulariacea Ehrenberg, 1838
Family Eggerellidae Cushman, 1937
Subfamily Dorothiinae Balakhmatova, 1972
Genus Dorothia Plummer, 1931
Dorothia hokkaidoana Takayanagi
Figures 2-16,17

Dorothia hokkaidoana Takayanagi, 1960, p. 83, pI. 3,
figs.9a-12b.

Types.-Hypotype, fig. 2-16, IGPS 101485,
sample SNS07; hypotype, fig. 2-17, IGPS
101486, sample SNS02.
Description.- Test small, elongate, narrow,sides nearly parallel, compressed in adult;
early portion with four or more chambers per

Kunio Kaiho

836

whorl, later becoming triserial, then being
regularly biserial; chambers in earlier portion almost indistinct, later rather distinct,
inflated, relatively low and broad, very gradually increasing in size as added; sutures
obscure in early portion, later becoming
rather distinct, depressed; wall coarsely
arenaceous, surface rather roughly finished;
aperture indistinct, a low arch at base of lastchamber.
Remarks.- The present specimens havemore inflated chambers and more roughly
finished surface than the holotoype and paratypes, partly because the present specimens
are much better preserved. Common species
in the Upper Cretaceous of Hokkaido.
Genus Marssonella Cushman, 1933
Marssonella sp. A
Figure 2-15
Dorothia crassa (Marsson). Yasuda, 1986, p. 55, pI.
4, figs. lOa, b (not of Marsson).

Type.-Hypotype, IGPS 101487, sample
SNS07.
Description.- Test small, elongate, the earliest portion rounded, later portion nearlyparallel sided, circular in transverse section,
early trochospiral portion with four to fivechambers per whorl, followed by biserial
stage, terminal face concave; chambers distinct, low very slightly inflated; suturesslightly depressed; wall finely arenaceous,surface smooth, colored in white; aperture
low, at inner margin of last-formed chamber.
Remarks.- This species is similar to Marsson ella ellisorae Cushman and Dorothia
beloides Hillebrandt, but differs in having alower trochospiral test in the earliest portion.

It resembles Gaudryna crassa Marsson, butdiffers in having a concave terminal face.
Suborder Lagenina Delage andHerouard, 1896
Superfamily Nodosariacea
Ehrenberg, 1838
Family Nodosariidae Ehrenberg, 1838
Subfamily Nodosariinae
Ehrenberg, 1838
Genus Dentalina Risso, 1826
Dentalina basiplanata Cushman
Figure 3-1
Dentalina basiplanata Cushman, 1938. p. 38-39, pI. 6,
figs. 6-8; Cushman, 1946, p. 68, pI. 24, figs. 1-6;
Takayanagi, 1960, p.94, pI. 5, fig. 6; Graham
and Church, 1963, p. 27, pI. 2, fig. 11.

Type.-Hypotype, IGPS 101488, sample
SNS07.
Description.- Test large, elongate, gently
arcuate, gently tapering toward rounded initial end, which is circular in cross section;chambers 11 in total, initial eight chambers
being low, overlapping, not inflated, later
becoming slightly inflated and slightly elongated; sutures mostly normal to long axis of
test, initially flush, limbate, later slightly
depressed; wall calcareous, smooth, finely
perforate; aperture radiate, placed toward
concave side.
Remarks.-Length of hypo type, 2.l8 mm;
diameter, 0.39 mm. Compared with the
holotype, studied specimens have a larger test,
thicker chambers, and relatively longer andnon-lobulate initial portion. They have amuch larger test and lower and thicker chambers than the specimens from the KjT boundary section of Hokkaido (Kaiho, in press, b)
and that of Takayanagi.

~ Figure 3. Scale bar= 100 j./m. 1, Dentalina basiplanata Cushman. SNS07. 2, Dentalina gracilisd' Orbigny. SNS02. 3, Nodosaria affinis Reuss, SNS02. 4, Nodosaria aspera Reuss. SNS02. 5, 6, Frondicularia mucronata Reuss. SNS07. 7, Frondicularia whaingaroica Stache. SNS07. 8,9, Frondicularia
bicornis Reuss. 8; SNS07. 9; SNS02. 10, Frondicularia sp. A. SNS07.
11, Frondicularia durrelli
Trujillo. SNS07. 12, 13, Lenticulina extruatus (Cushman). SNS07. 14, Lenticulina sp. A. SNS07. 15,
Lenticulina yabei Takayanagi. SNS02. 16, Lenticulina rotulata (Lamarck). SNS02. 17, Lenticulina
obirashibensis Takayanagi, SNS02.

923.

Campanian benthic foraminifera from central Hokkaido

837

Kunio Kaiho

838

Dentalina gracilis d'Orbigny
Figure 3-2
Dentalina gracilis d'Orbigny, 1840, p. 14, pI. 1, fig. 5(Inaccessible, fide Catalogue of Foraminifera) ;Cushman, 1946, p. 65, pI. 23, figs. 3-6.
Dentalina inornata d'Orbigny, 1846, p.44, pI. 1, figs.
50, 51 ; Papp and Schmid, 1985, p. 28, pI. 9, figs.
5-8.

Type.-Hypotype, IGPS 101489, sample
SNS02.
Description.- Test long, slender, uniserial,
somewhat arcuate; chamber inflated except
for early stage of microspheric forms; sutures
distinct, oblique; wall calcareous, smooth,finely perforate; aperture radiate, placed
toward concave side.
Remarks.-It resembles Dentalina luma
Belford from the Campanian of western
Australia, but differs in having more inflatedchambers in the early stage and more oblique
sutures.
Genus Nodosaria Lamarck, 1812
Nodosaria affinis Reuss
Figure 3-3
Nodosaria affinis Reuss, 1845, p. 26, pI. 13, fig. 16.
Nodosaria paupercula Ruess. Cushman, 1946, p. 75,
pI. 27, figs. 10-12 (not of Reuss).

Type.-Hypotype, IGPS 101490, sample
SNS02.
Remarks.- This fragmented specimen possesses a uniserial rectilinear chamber arrangement and wall ornamented by 16 longitudinal
and nearly continuous costae.

Nodosaria aspera Reuss
Figure 3-4
Nodosaria aspera Reuss. Graham and Church, 1963,
p. 41, pI. 4, figs. 17, 18; Cushman, 1946, p. 72, pI.
26, fig. 6.

Type.-Hypotype, IGPS 101491, sample
SNS02.
Description.- Test uniserial, rectilinear,rounded in cross-section, chambers slightly

overlapping, increasing rather uniformly insize as added; sutures normal to long axis of
test, depressed; wall ornamented with small,closely set spines covering entire surface;
aperture rounded with a short cylindricalneck projecting well beyond outline of final
chamber.
Remarks.- The single specimen has moreelongate and less overlapping chambers thanthe holotype of Nodosaria aspera Reuss.
Subfamily Frondiculariinae Reuss, 1860
Genus Frondicularia Defrance, 1826
Frondicularia bicornis Reuss
Figures 3-8, 9Frondicularia bicornis Reuss, 1845, p. 32, pI. 13,
fig. 45.
Frondicularia bicornis Reuss var. etiola Marie, 1941,
p. 129, pI. 15, fig. 185.
Frondicularia bicornis Reuss var. rhomboidalis Marie,
1941, p. 129, pI. 15, fig. 184.
Frondicularia cf. elegans d'Orbigny. Takayanagi,
1960, p. 112, pI. 6, figs. 21a, b. (not of d'Orbigny)
Frondicularia verneuiliana d'Orbigny. Sliter, 1968,
p. 62, 63, pI. 6, figs. 8-lOc. (not of d'Orbigny)

Types.-Hypotype, IGPS 101492, figure 38 (megalospheric), sample SNS07; hypotype,
IGPS 101493, figure 3-9 (microspheric), sample SNS02.
Description.- Test elongate-leaf-shaped,
compressed, sides flat or slightly concave,
periphery truncate, greatest breadth near theapertural end; chambers elongate, low,increasing gradually in size; sutures distinct,
oblique, limbate, slightly elevated; wall calcareous, finely perforate, surface smoothly
finished, with a single costa on globular
proloculus and sometimes very fine shortcostae on later chambers; aperture terminal,
radiate.
Remarks.- The megalospheric test has abroader width than the microspheric test.

Frondicularia durrelli Trujillo
Figure 3-1
Frondicularia durrelli Trujillo, 1960, p.323, pI. 46,

923.

Campanian benthic foraminifera from central Hokkaido

839

fig. 8; Sliter, 1968, p.60, pI. 6, figs. la, b.

(Inaccessible, fide Catalogue of Foraminifera).

Type.-Hypotype, IGPS 101494, sample
SNS07.
Description.- Test elongate-leaf-shaped,
compressed, sides nearly parallel, peripheral
margins truncate, carinate; chambers low,increasing gradually in size; sutures flush,
oblique; wall calcareous, finely perforate,with three to six, sinuate, high, longitudinalcostae extending length of test; aperture terminal, radiate.
Remarks.-It has a much slender test than
Frondicularia striatula Reuss described by
Takayanagi (1960).

Type.-Hypotype, IGPS 101497, sample
SNS07.
Description.- Test large, e10ngate-leafshaped, flat, broadest near middle; chambers
numerous, angularly curved at longitudinalaxis, very narrow, with a longitudial depression along longitudinal axis; septal sutures
depressed; wall calcareous, finely perforate,surface smooth except for a longitudinal andsutural depression; aperture terminal.
Remarks.- This species is distinguishedfrom other species in having a large, thin testwith numerous narrow chambers which arevery angularly curved at the longitudinal axis.

Frondicularia mucronata Reuss
Figures 3-5, 6
Frondicularia mucronata Reuss, 1845, p.31, pI. 13,
figs. 43, 44.
Frondicularia mucronata Reuss subsp. costata Balakhmatova, 1960, p. 93, pI. 16, figs. I, 2, 5 (Inaccessible, fide Catalogue of Foraminifera).

Type.-Hypotype, figure 3-5, IGPS
101495; hypotype, figure 3-6, IGPS 101496;
Both from sample SNS07.
Description.- Test leaf-shaped, flat, with abasal spine, greatest breadth of test not at
middle but rather near aboral end, peripherytruncate, narrow, flat; initial chamber nearly
globular with a few distinct short longitudinal costae, remaining two to four, angularly
curved chambers narrow; sutures limbate,slightly curved; wall calcareous, finely perforate; aperture distinct, on a long quadrangular neck.
Remarks.- This species is characterized byhaving a basal spine and the greatest breadth'of the test near the aboral end.

Frondicularia sp. A
Figure 3-10

Type.-Hypotype, IGPS 101498, Sample
SNS07.
Description.- Test large, leaf-shaped, flat,
periphery truncate, greatest breadth about
middle, length approximately twice as long as
width; chambers elongate, curved, low,increasing gradually in size; sutures curved,
depressed; wall calcareous, finely perforate
with numerous short fine longitudinal costae
on each chamber; aperture terminal, radiate.
Remarks.- This specimen is similar to F.
bicornis Reuss, but differs in having a broader
test and depressed sutures.
Family Vaginulinidae Reuss, 1860
Subfamily Lenticulininae Chapman,
Parr and Collins, 1934
Genus Lenticulina Lamarck, 1804
Lenticulina extruatus (Cushman)
Figures 3-12,13

Frondicularia whaingaroica Stache
Figure 3-7
Frondicularia whaingaroica Stache, 1865, p. 210, pI.
22, fig. 43 (Inaccessible, fide Catalogue of Foraminifera).
Frondicularia fragilis Karrer, 1870, p. 175, pI. 2, fig. 3

Robulus navarroensis (Plummer) var. extruatus Cushman, 1938, p. 31, pI. 5, fig. 1 ; Cushman, 1941,
p. 56, pI. 15, fig. 2; Cushman, 1946, p. 52, pI. 16,
figs. 9-lOb, pI. 17, fig. 2a, b.
Lenticulina spissocostata (Cushman). Perlmutter
and Todd, 1965, p. 11, pI. 1, fig. 13; Sliter, 1968,
p. 67, 68, pI. 7, figs. 7a-8b.

Kunio Kaiho

840

Types.-Hypotype, figure 3-12, IGPS
101499; hypotype, figure 3-13, IGPS 101500.
Both from sample SNS07.
Description.- Test large, lenticular, periphery acute, with a keel; chambers distinct,
nine in last whorl, gradually increasing in size
as added: sutures curved, distinctly limbate,strongly raised; wall may have several
obliquely curved costae, which are nearlyparallel to outer edge of test; aperture radiate, at peripheral angle.
Remarks.- This species is distinguishedfrom other species in having strongly raised
sutures. It has a more elliptical test and highapertural face than Robulus venustus Takayanagi which has a rounded outline.
Lenticulina haboroensis (Takayanagi)
Figures 4-4, 5
Hemicristellaria haboroensis Takayanagi, 1960, p.
109, pI. 6, figs. 14a, b.

Types.-Hypotype, figure 4-4, IGPS
101501; hypotype, figure 4-5, IGPS 101502.
Both from sample SNS07.
Description.- Test ovate in outline,
flattened, side roughly parallel, dorsal periphery keeled, apertural face narrow, elongate,flat, depressed between two marginal keels;chambers broad and low, enrolled in very
early stage, later uncoiling, added on slightly
curved axis, increasing gradually in size;
sutures curved, raised; wall calcareous, finely
perforate, often with short longitudinal costae
in early portion; aperture radial, at dorsal
edge.
Remarks.- The studied specimens have asmaller coiling part and a broader uncoilingpart than the holotype.
Lenticulina obirasibensis Takayanagi
Figure 3-17
Lenticulina obirasibensis Takayanagi, 1960, p. 103, pI.
5, figs. 28a, b; Yasuda, 1986, p. 66, 67, pI. 6, figs.
7a-c.
Lenticulina sp. cf. L. lobata (Reuss) of Egger. Graham

and Church, 1963, p. 34, pI. 3, figs. 17a, b.
Robulus sp. A. Martin, 1964, p.69, pI. 7, figs. la, b.

Type.-Hypotype, IGPS 101503, sample
SNS02.
Description.- Test planispiral, involute,completely lenticular, elliptical in outline,
compressed, periphery acute with slight todistinct keel; seven to nine chambers in lastwhorl, gradually increasing in size; sutures
gently curved, limbate, flush with surface orvery slightly raised; wall calcareous, finely
perforate, surface smooth; aperture radiate atperipheral angle, usually with a ventral slit on
apertural face.
Remarks.- This species is characterized by
its elliptical outline, seven to nine chambersin the final whorl, gently curved suture and byhaving a thickness/diameter ratio of the test
being about two-fifths. This species has amuch compressed test than Lenticulina williamsoni (Reuss) occurring in the K/Tboundary section of Hokkaido (Kaiho, inpress, b) and Robulus sorachiensis Takayanagi.
Lenticulina rotulata (Lamarck)
Figure 3-16
Lenticulites rotulata Lamarck, 1804, p. 188; 1806, pI.
62, fig. 11 (Inaccessible, fide Catalogue of Foraminifera).
Lenticulina rotulata (Lamarck). Cushman, 1927, p.
142, pI. 28, figs. 7a, b (holotype redrawn); Frizzell, 1943, p.341, pI. 56, figs. 2a, b; Cushman,
1946, p.56-57, pI. 18, fig. 19, pI. 19, figs. 1-7;
Takayanagi, 1960, p. 103, 104, pI. 5, figs. 29a, b ;Yasuda, 1986, p. 67, 68, pI. 6, figs. lOa, b.
Lenticulina muensteri (Roemer). Sliter, 1968, p.66,
pI. 7, figs. 9, 13.

Type.-Hypotype, IGPS 101504, sample
SNS02.
Description.- Test, lenticular, closely
coiled, roughly circular in outline, biumbonate, periphery acute with slight to distinct
keel; chambers eight to eleven in final whorl,
increasing gradually in size; sutures gently
curved, limbate, flush with surface or slightly
raised; wall calcareous, finely perforate, sur-

923.

Campanian benthic foraminifera from central Hokkaido

face smooth; aperture radiate at peripheral
angle, slightly protruded, with a ventral slit
on an apertural face.
Remarks.- This species is characterized byits circular outline, biumbonate and distinct
keel. It has a thicker keel and larger test
than L. rotulata of Takayanagi (loc. cit.). It
resembles Lenticulina tanakai Takayanagi,
but differs in having a more compressed test,
better developed keel and circular umbonal
boss.

841

whorl, chambers increasing very slowly inthickness, but rapidly in breadth; sutures
limbate, curved, flush with surface; wall calcareous, finely perforate, thick, surface
smooth; aperture radiate at peripheral angle.
Remarks.- This species is characterized byits semi-heart-shaped outline, thick wall, distinct keel and large proloculus followed byfour to five chambers in the megalospheric
forms.

Lenticulina sp. A
Lenticulina williamsoni (Reuss)
Figure 4-2

Cristellaria williamsoni Reuss, 1862, p. 327, pI. 6, figs.
4a-b.
Robulus williamsoni (Reuss). Cushman, 1931, p. 37,
pI. 5, figs. 2a, b; Cushman, 1946, p. 54, pI. 18,
figs. 2a-3b.

Type.-Hypotype, IGPS 101505, sample
SNS02.
Description.- Test lenticular, thick, periphery slightly lobulate, slightly keeled, large
apertural face; chambers six in final whorl,
inflated, rapidly increasing in size; sutures
curved, depressed; wall calcareous, finely
perforate, surface smooth; aperture radiate,
at peripheral angle.
Remarks.- The present specimens differsfrom L. williamsoni occurring from the KIT
boundary section of Hokkaido (Kaiho, inpress, b) in having depressed sutures andinflated chambers.
Lenticulina yabei Takayanagi
Figure 3-15

Lenticulina yabei Tkayanagi, 1960, p. 106, pI. 6, figs.
2a-3b: Yasuda, 1986, p. 68, pI. 6, figs. 13a-c.

Type.-Hypotype, IGPS 101506, sample
SNS02 (megalospheric form).
Description.- Test semi-heart-shaped,
lenticular, compressed, periphery distnctly
keeled; large proloculus followed by four tofive chambers in megalospheric forms andeight chambers in microspheric forms in last

Figure 3-14

Type.-Hypotype, IGPS 101507, sample
SNS07.
Remarks.- This species differes from L.
extruatus Cushman in having more weakly
raised sutures and longer apertural face.
Lenticulina sp. B
Figure 4-1

Type.-Hypotype, IGPS 101508, sample
SNS02.
Description.- Test long elliptical, compressed, periphery acute, without distinct
keel; chambers increasing rapidly in size,
somewhat inflated in last chamber; sutures
gently curved, flush with surface; wall calcareous, finely perforate, surface smooth; aperture radiate, at peripheral angle, with a ventral slit on apertural face.
Remarks.- This species is characterized bya long elliptical outline and lack of distinct
keel. It has an outline similar to that ofLenticulina yabei Takayanagi, but differs inhaving a slit on the apertural face and more
thicker chambers in later portion. It differsfrom L. obirashibensis in having a more elongate test, inflated last chamber and relatively
higher apertural face.
Lenticulina sp. C
Figure 4-3

Type.-Hypotype, IGPS 101509, sample

842

Kunio Kaiho

923.

Campanian benthic foraminifera from central Hokkaido

SNS07.
Description.- Test lenticular, thick, roughly
ovate in outline, planispiral, involute; chambers seven in final whorl, increasing graduallyin size; sutures curved, raised; wall calcareous, finely perforate, surface smooth except
for raised sutures; aperture radiate at peripheral angle.
Remarks.- This specimen is distinguishedfrom L. williamsoni Reuss in having raised
sutures.

843

verse section, dorsal angle acute, apertural
face broad and inflated; chambers low andbroad, increasing gradually in size; sutures
much curved in enrolled portion, but becoming gently curved later, flush with surface;wall calcareous, finely perforate, surface
smooth; aperture radial, at edge of dosal
side.
Remarks.- This species is characterized byits smooth surface and acute non-keeled dorsal periphery. Common in the Upper
Cretaceous of Hokkaido.

Genus Pravoslavlevia Putrya, 1970
Pravoslavlevia sp.

Saracenaria tunesiana ten Dam and Sigal

Figure 4-12

Figures 4-7-9,13

Type.-Hypotype, IGPS 101515, sample
SNS07.
Remarks.- The specimen differs from
Saracen aria tunesiana ten Dam and Sigal in
having a larger, more closed, and biumbonate
initial coiling stage, narrower chambers in
lectilinear stage, and smooth surface.
Genus Saracen aria Defrance, 1824
Saracen aria triangularis (d'Orbigny)
Figure 4-6
Cristellaria triangularis d'Orbigny, 1840, p.27, pI. 2,
figs. 21, 22. (Inaccessible, fide Catalogue of Foraminifera)
Saracenaria triangularis (d'Orbigny). Cushman and
Church, 1929, p.505, pI. 37, figs. 13, 14; Cushman, 1946, p. 58, pI. 28, figs. la-3.
Saracenaria cf. triangularis (d'Orbigny). Takayanagi, 1960, p. 106, pI. 6, figs. 8a, b.
Saracen aria sp. A. Yasuda, 1986, p. 70, 71, pI. 7, figs.
6a-c.

Type.-Hypotype, IGPS 101510, sample
SNS07.
Description.- Test planispirally enrolled inearly stage, later flaring and tending tobecome rectilinear, chestnut-shaped in trans-

Saracen aria tunesiana ten Dam and Sigal, 1950, p.
36, pI. 2, fig.21; Trujillo, 1960, p.316,317, pI.
45, figs. 11 a, b.

Types.-Hypotype, IGPS 101511, figure 47; hypotype, IGPS 101512, figure 4-8; hypotype, IGPS 101513, figure 4-9; hypotype,
IGPS 101514, figure 4-13. All from sample
SNS07.
Description.- Test elongate, triangular intransverse section, angled acute, keeled inearly dorsal portion, early stage closely
coiled, later three to four chambers uncoilingand becoming rectilinear, with broad inflated
ventral face of last chamber and strongly
depressed middle part of ventral side; chambers broad and low, highest at dorsal margin;
sutures gently curved, slightly depressed;
wall calcareous, finely perforate, with longitudinal costae, curved and almost parallel withdorsal periphery; aperture radiate at dorsal
angle.
Remarks.- This species is distinguished inhaving a long uncoiling part, deeply depressed middle part of the ventral side, andseveral distinct longitudinal costae.

~ Figure 4. Scale bar= 100.um. 1, Lenticulina sp. B. SNS02. 2, Lenticulina williamsoni (Reuss).
SNS02. 3, Lenticulina sp. C. SNS07. 4, 5, Lenticulina haboroensis (Takayanagi). SNS07. 6, Saracenaria triangularis (d'Orbigny). SNS07. 7-9,13, Saracenaria tunesiana ten Dam and Sigal. SNS07.
10, 11, Astacolus elongatus, Kaiho, n. sp. 10; SNS07. 11; SNS02. 12, Pravoslavlevia sp. SNS07. 14,
15, Astacolus nagaoi (Takayanagi). SNS07. 16, 17, Astacolus polandensis Trujillo. SNS07.

Kunio Kaiho

844

Subfamily Marginulininae Wedekind, 1937
Genus Astacolus de Montfort, 1808
Astacolus elongatus Kaiho, n. sp.
Figures 4-10, II

Types.-Holotype, figure 4-10, IGPS
101516, sample SNS07; paratype, figure 4-11,
IGPS 101517, sample SNS02.
Description.- Test elongate, flattened,
periphery rounded; chambers numbering up
to eight, broad and low, but last chamber
relatively high, added on a slightly curved
axis, sutures strongly oblique, curved; wallcalcareous, finely perforate, surface smooth;aperture radiate at dorsal angle.
Remarks: This new species is distinguished from other related species in having avery elongate test. It resembles Astacolus
jarvisi (Cushman), but differs in having amore slender test.

Astacolus nagaoi (Takayanagi)
Figures 4-14,15

Hemirobulina nagaoi Takayanagi, 1960, p. 117, pI. 7,
figs.13a-14b.

Types.-Hypotype, figure 4-14 (microspheric form), IGPS 101518; hypotype, figure
4-15 (megalospheric form), IGPS 101519.
Both from sample SNS07.
Description.- Test ovate in outline, nearly
three times as long as broad, flattened, periphery subacute to rounded, not keeled, apertural
face very narrow, high, flat, truncate; chambers five to seven, broad and low, enrolled invery early stage, later becoming uncoiled,added on a slightly curved axis, increasing
very slowly in size, strongly oblique, dorsalside higher than ventral; sutures distinct,
strongly oblique, very slightly curved, flushwith surface; wall calcareous, finely perforate, very thin, translucent, surface smooth;aperture radiate, terminal, at dorsal angle.
Remarks.- The microspheric form is composed of a small pro10culus followed by six
low chambers. The megalospheric form hasa large proloculus followed by four higher
chambers. The megalospheric form has athicker test than the microspheric form.

Astacolus polandensis Trujillo
Figures 4-16, 17

Astacolus polandensis Trujillo, 1960, p. 317, 318, pI.
45, figs. 14a, b.

Types.-Hypotype, figure 4-16, IGPS
101520; hypotype, figure 4-17, IGPS 101521.
Both from sample SNS07.
Description.- Test ovate in outline, nearly
three times as long as broad, flattened, dorsal
periphery acute with narrow keel in early
portion, apertural face very narrow, high, flat,
truncate; large proloculus followed by four
triangular chambers, increasing rapidly insize, oblique, dorsal side higher than ventral,slightly inflated; sutures distinct, oblique,slightly curved, depressed; wall calcareous,
finely perforate, surface smooth; apertureradiate, terminal, at dorsal angle.
Remarks.- This species differs from A.
nagaoi in having a keeled dorsal periphery,
depressed sutures and inflated higher chambers.
Genus Marginulina d'Orbigny, 1826
Marginulina bullata Reuss
Figures 5-1, 2

Marginulina (Marginulina) bullata Reuss, 1845, p. 29,
pI. 13, figs. 34-38.
Marginulina bullata Reuss. Cushman, 1946, p.62,
pI. 21, figs. 32a-37; Bandy, 1951, p.498, pI. 72,
figs. 13a, b; Graham and Church, 1963, p. 37, pI.
4, figs. la, b; Martin, 1964, p.63-64, pI. 5, figs.
lOa-b; Sliter, 1968, p. 70, pi 8, figs. 6a-7.
Marginulina cf. M. glabra d'Orbigny. Cushman,
1951, p. 18, pI. 5, figs. 25-27. (not of d'Orbigny)
Marginulina texasensis Cushman. Takayanagi,
1960, p.90, pI. 4, figs. 13a, b (not of Cushman).
Marginulina troedssoni Brotzen. Trujillo, 1960, p.
325-326, pI. 46, figs. 14a-c (not of Brotzen).

Types.-Hypotype, figure 5-1 (microspheric form), IGPS 101522; hypotype, figure 5-2(megalospheric form), IGPS 101523. Both

923.

Campanian benthic foraminifera from central Hokkaido

from sample SNS02.
Description.- Test elongate, coiled in earlystage, later stage becoming uncoiled, circularin cross section, initial edge compressed in
microspheric forms, rounded in megalospheric form; five or six chambers, slightly inflated
in early portion, more inflated in later portion; sutures distinct, flush in early stage,
later depressed, nearly transverse; wall calcareous, finely perforate, smooth; apertureradiate, terminal, at dorsal angle.
Remarks.- This is a cosmopolitan, Late
Cretaceous species.
Subfamily Vaginulininae Reuss, 1860
Genus Citharina d'Orbigny
Citharina multicostata (Cushman)
Figure 5-3
Vaginulina multicostata Cushman, 1930, p.28, pI. 4,
fig. 4; Cushman, 1946, p. 79, pI. 29, figs. 9-16.
Citharina sp. A. Graham and Church. 1963, p. 26, pI.
2, fig. 9.
Citharina multicostata (Cushman). Sliter, 1968, p.
55, 56, pI. 5, fig. 6.
Citharina strigillata (Reuss). Loeblich and Tappan,
1988, p. 116 pI. 452, figs. 1,2.

Type.-Hypotype, IGPS 101524, sample
SNS07.
Description.- Test elongate, much compressed throughout, dorsal edge straight, ventral edge convex, tapering on both sides,
periphery truncate; chambers elongate,
curved, strongly oblique, low and broad;sutures not depressed, very oblique, curved;
wall ornamented with numerous costae, parallel to straight dorsal edge of test, and continuous in early portion, but independent of
individual chambers in later portion; aperture terminal.
Remarks.- The present specimens havestronger costae in early portion than theholotype. This species differs from Citharina suturalis Cushman described from theCretaceus of Arkansas, but differs in having a
truncate periphery, independent costae of theindividual chambers in later portion and845

strong continuous costae in early portion.
Genus Psilocitharella Loeblichand Tappan, 1986
Psilocitharella ezoensis Kaiho, n. sp.
Figures 5-4, 5

Types.-Holotype, fig. 5-4, IGPS 101525;
paratype, fig. 5-5, IGPS 101526. Both fromsample SNS07.
Description.- Test elongate, two sides nearly parallel, rectilinear, compressed, peripherytruncate, margin carinate, straight on dorsalside, lobulate in ventral side; chambers
inflated, half as low as breadth in early portion, but as high as breadth in later portion;
sutures oblique, highest at dorsal side,straight, depressed; wall calcareous, finely
perforate, surface ornamented by longitudinal
or slightly oblique costae not continued
beyond sutures; aperture radiate at the dorsal angle, protruded.
Remarks.- This new species has a muchlarger test and lower chambers than those of
Citharina geisendorferi (Franke) costata
Takayanagi.

Psilocitharella triangulata Kaiho, n. sp.
Figure 5-6Type.-Holotype, IGPS 101527; paratype,
IGPS 101528. Both from sample SNS07.
Description.- Test subtriangular in outline,
elongate, compressed, periphery truncate,margin carinate; chambers broad, low, notinflated; sutures elevated, curved oblique inearly portion, straight, strongly oblique inlater portion; wall calcareus, finely perforate,surface smooth, without elevated costae;
aperture radiate, at dorsal angle, slightly
protruded.
Remarks.- This new species resembles
Citharina cf. recta Reuss of Takayanagi(1960), but differs in having broader chambers.

846

Kunia Kaiha

923.

Campanian benthic foraminifera from central Hokkaido

847

Family Polymorphinidae d'Orbigny, 1839
Subfamily Polymorphininae d'Orbigny, 1839 .
Genus Globulina d'Orbigny, 1839
Globulina lacrima (Reuss)

and with remnants of connecting tubes atboth ends.

Figure 5-7

Figure 5-9

Polymorph ina (Globulina) lacrima Reuss, 1845, p. 40,
pI. 12, fig. 6, pI. 13, fig. 83.
Globulina lacrima Reuss. Cushman and Ozawa,
1930, p.77, pI. 19, figs. 1-2; Cushman, 1946, p.
96,pI.40,figs.lla-12; HoiKer, 1957,p.170, 171,
text-figs. 212, 213; Sliter, 1968, p.77, pI. 9, figs.
17a-c, pI. 10, figs. la-c.
Globulina lacrima lac rima Reuss. Graham andChurch, 1963, p.48, pI. 5, figs. 15a-c.

Type.-Hypotype, IGPS 101529, sample
SNS07.
Description.- Test subglobular, ovate incross section; chambers strongly appressed,
extending nearly to base, becoming sigmoidal; suture distinct, flush; wall calcareous, finely perforate, surface smooth; aperture terminal, radiate.

Ramulina tetrahedralis Ludbrook

Ramulina tetrahedralis Ludbrook.
1974, p. 178, pI. 34, figs. 4, 5.

Scheibnerovli,

Type.-Hypotype, IGPS 101531, sample
SNS02.
Remarks.- This species has a spherical testwith four cylindrical arms and very finely
hispid surface.
Suborder Rotaliina Delage
and Herouard, 1896
Superfamily Stilostomellacea Finlay, 1947
Family Stilostomellidae Finlay, 1947
Genus Stilostomella Guppy, 1894
Stilostomella alexanderi (Cushman)
Figure 5-10

Subfamily Edithaellinae Fuchs, 1967
Genus Ramulina T.R. Jones, 1875
Ramulina aculeata (d'Orbigny)
Figure 5-8
Dentalina aculeata d'Orbigny, 1840, p. 13, pI. I, figs.
2,3 (Inaccessible, fide Catalogue of Foraminifera); Cushman, 1946, p.67, pI. 26, figs. 17,
18?; Martin,1964, p.60, pI. 4, figs. 12a, b.
Ramulina sp. a Takayanagi, 1960, p. 118, pI. 7, figs.
17,18.
Ramulina pseudoaculeata (Olsson). Sliter, 1968, p.
79, pI. 10, fig. 8.
Ramulina sp. Scheibnerovli, 1974, p. 180, pI. 35, fig. 1.

Type.-Hypotype, IGPS 101530, sample
SNS02.
Remarks.- These segmented specimens havea pyriform test with numerous short spines~

Ellipsonodosaria alexanderi Cushman, 1936, p. 52, pI.
9, figs. 6-9; Cushman, 1946, p. 135, pI. 56, figs.
12-15.
Ellipsonodosaria alexanderi var. impensia Cushman,
1938, p.48, pI. 8, figs. 4, 5; Cushman, 1946, p.
136, pI. 56, figs. 16-18; Cushman, 1949, p.9, pI.
4, fig. II.
Stilostomella alexanderi (Cushman). Skinner, 1962,
p.47, pI. 5, fig. 31 ; Yasuda, 1986, p.81, pI. II,
figs. 2a, b.

Type.-Hypotype, IGPS 101532, sample
SNS07.
Remarks.- Those segments characterized
by a wall ornamented with short backwardly
pointing spines, which are irregularly scattered over the wall surface, are referred to thepresent species. It is a cosmopolitan Creta-

Figure 5. Scale bar= 100 j.lm. 1,2, Marginulina bullata Reuss. SNS02. 3, Citharina multicostata(Cushman). SNS07. 4, 5, Psilocitharella ezoensis Kaiho, n. sp. SNS07. 4; holotype. 5; paratype. 6,
Psilocitharella triangulata kaiho, n. sp. SNS07, holotype. 7, Globulina lacrima (Reuss). SNS07. 8,
Ramulina aculeata (d'Orbigny). SNS02. 9, Ramulina tetrahedralis Ludbrook. SNS02. 10, Stilostomella alexanderi (Cushman). SNS07. 11, Stilostomella stephensoni (Cushman). SNS07. 12, 13,
Nuttallinella florealis (White). SNS07. 14-17, Notoplanulina rumoiensis (Takayanagi). 14, 16; SNS07.
15, 17; SNS02.

Kunio Kaiho

848

ceous species.

Stilostomella stephensoni (Cushman)
Figure 5-11
Ellipsonodosaria stephensoni Cushman, 1936, p. 52,
pI. 9, figs. 10-15; Cushman, 1946, p. 134, pI. 56,
figs. 2-7; Cushman, 1949, p. 9, pI. 4, fig. 10.
Stilostomella stephensoni (Cushman). Frizzell,1954,
p. 121, pI. 18, figs. 22, 23 (in text as Ellipsonodosaria stephensoni Cushman); Said and
Kenawy, 1956, p. 146, pI. 4, fig. 37; Takayanagi,
1960, p. 121, pI. 8, fig. 3; Skinner, 1962, p. 47, pI.
5, fig. 30; Yasuda, 1986, p. 82, pI. 11, figs. 3a-4.
Siphonodosaria pseudoscripta (Cushman) of Graham
and Church, 1963, p. 56, pI. 6, fig. 13.

Type.-Hypotype, IGPS 101533, sample
SNS07.
Description.- Test elongate, rectilinear,
tapering, with short, longitudinal, costae-likespines surrounding mostly lower half of its
pyriform chambers.
Remarks.- This species resembles Stilostomella stephensoni Cushman of Takayanagi(1960) and Siphonodosaria pseudoscripta(Cushman) of Graham and Church (1963),
but has a much larger test (about twice inlength).
Superfamily Asterigerinacea
d'Orbigny, 1839
Family Epistomariidae Hofker, 1954
Subfamily Eponidellinae Seiglie
and Bermudez, 1965
Genus Nuttallinella Belford, 1959
Nuttallinella florealis (White)
Figures 5-12,13
Gyroidina florealis White, 1928, p. 293, pI. 40, fig. 3.
Nuttallina florealis (White). Takayanagi, 1960, p.
128, pI. 9, figs. 3a-5.

Type.-Hypotype, figure 5-12, IGPS
101534; hypotype, figure 5-13, IGPS 101535.
Both from sample SNS07.
Description.- Test planoconvex, spiral side
flattened, umbilical side strongly convex, withumbilical boss, nearly flat, peripheral outline
lobulate, peripheral margin acute, keeled;chambers five in final whorl; sutures limbate,strongly curved, flush in spiral side, radial,slightly curved, slightly depressed in umbilical side; wall calcareous, finely perforate,surface smooth; aperture interiomarginal, an
elongate slit opening near periphery.
Remarks.-Abundant in the CampanianKashima Formation, in the Oyubari area,
Hokkaido.
Superfamily Chilostomellacea Brady, 1881
Family Gavelinellidae Hofker, 1956
Subfamily Gyroidinoidinae Saidova, 1981
Genus Notoplanulina Malumianand Masiuk, 1976
Notoplanulina rumoiensis (Takayanagi)
Figures 5-14-17
Planulina rumoiensis Takayanagi, 1960, p. 139, pI. II,
figs. la-c.

Types. - Hypotype, figure 5 -14, I G P S
101536, sample SNS07; hypotype, figure SIS, IGPS 101537, sample SNS02; hypotype,figure 5-16, IGPS 101538, sample SNS07;
hypotype, figure 5-17, IGPS 101539, sample
SNS02.
Description.- Test flattened, low trochospiral, planoconvex, two whorls visible on
spiral side, umbilicus open, bordered with a
lip and partly covered by umbilical chamber
extensions, periphery angular and carinate;seven to eight chambers in final whorl, low
and broad, inflated on umbilical side;sutures strongly curved and depressed onumbilical side; wall calcareous, finely perforate except for imperforate thickened sutures;
aperture, an elongate interiomarginal slit,
extending from near periphery to open umbilicus.
Remarks.-I recognized that the holotypeof this species is indeed a juvenile form
because of the discovery of abundant adultspecimens from the two samples treated herein. This species resembles Notoplanulina
rakauroana Finlay, but differs in having asmaller, less evolute test.

923.

Campanian benthic foraminifera from central Hokkaido

Acknowledgments

I would like to thank Y. Takayanagi forreviewing the manuscript and T. Saito forreading it. I am thankful to K. Takahashi
and N. Wada of the Geological Survey of
Hokkaido for their help during my field workand to S. Otomo for his assistance in thephotographic work.
References cited
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850

Kunio Kaiho

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Jour. Paleont., vol. 34, p. 290-346, pis. 43-50.
White, M.P., 1928: Some index foraminifera of the923. ,Campanian benthic foraminifera from central Hokkaido
Tampico Embayment area of Mexico. Part II.
Ibid., vol. 2, p. 280-317, pis. 38-42.
Yasuda, H., 1986: Cretaceuos and Paleocene for-851

aminifera from northern Hokkaido, Japan. Sci.
Rep., Tohoku Univ., 2nd Ser. (Geol.), vol. 57, p.
1-101, pis. 1-14.

Hobetsu-cho f,I.5JljlllJ.
~trta:@:t:p:R:WO) j]:/ /~.=. 7 :/:M0)t:pJli7.l\.JtI;;~1i::fLlR: ~trta:@:t:p:R:Wf,I.5JIjIllJ~;:,7t;(jji' G~~JIi~Jj;'[!1bJliO) 10 kg 0)~fl2 "'Y/J' I?, j]:/ /~.=. 7 :/:MO)7$JhUE1i::fLlR~;:,lliRffi' G t:pJli7.l\.
Jtl;;1:.1i::fLlRift-J 50,000 M{;$:a-~tI:I L t.:o .: O)JtI;;~~$a-fII\:nl(;i' Go)ft, M{;$:~O) 85-90% a-d:i

Ih G Silicosigmoilina Jutabaensis c, Notoplanulina rumoiensis, Nuttallinella florealis,
Haplophragmoides asanoi, Dorothia hokkaidoana tJ:, c', 47 fi-e t!; Go .:.h I? O)fia-g~t1t L,~$fII\:nl(;a-1JJ3 I? iH;:, L t.:o ~~i.J' I? J! G C, Lagenina BE El i.J~ 66% a- d:i Ih, Rotaliina BE El ft,
9% co/tJ:,l"o ':O)~$ft, J:.W,*i*rta1l'f (200-500m) 0)7.I\.i*a-lf-i'0¥taf*!FJ):7\:

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 163, pp. 852-860, 4 Figs., September 30, 1991

924.

THALASSINA A NOMA LA (HERBST, 1804) (THALASSINIDEA:
DECAPODA) FROM THE MIOCENE BIHOKUGROUP, SOUTHWEST JAPAN*
HIROAKI KARASA W A
Department of Earth Sciences, School of Science,
Nagoya University, Chikusa, Nagoya 464-01
andISAO NISHIKAWA
Yuki, Yuki-cho, Hiroshima Prefecture 720-18

Abstract. Thalassina anomala (Herbst, 1804), a thalassinid decapod, is describedfrom the early Middle Miocene Bihoku Group exposed at Munekane, Yuki-cho, Jinseki-gun,
Hiroshima Prefecture, Southwest Japan. Thalassina anomala is a living species which
inhabits mangrove swamps and this occurrence shows the existence of mangrove swampy
paleoenvironment in the Middle Miocene. It is closely related to the tropical spike recognized in the early Middle Miocene of Southwest Japan.
Key words.
anomala.

Bihoku Group, early Middle Miocene, Hiroshima Prefecture, Thalassina

the Pleistocene to Holocene formations inAustralia (Etheridge and MacCulloch, 1916;
Forster and Barthel, 1978, etc.), New Guinea(Van Straelen, 1928) and Japan (Imaizumi,1953). Imaizumi (1969) also reported
Thalassina sp. from the early Middle
Miocene Kawaminami Formation in Kaga
City, Ishikawa Prefecture. The occurrenceof fossil Thalassina in the Bihoku Group is
the second record from Japanese Miocene
formations.
In this paper, we describe Thalassina
anomala from the Bihoku Group and discuss
its paleoecological and geological significance.

Introduction

The burrowing mangrove mud lobster,Thalassina anomala (Herbst), is one of thecharacteristic thalassinid decapods which
inhabits mangrove swamps in the tropicalIndo-West Pacific regions. It is well known
to be a large, active burrower living in mangrove swamps on the high spring tide zone,
forming large mounds.
The junior author, Nishikawa, has collected and reported many thalassinid shrimps
from the Miocene Bihoku Group exposed at
Munekane, Hiroshima Prefecture (Nishikawa, 1972; 1974). As a result of our subsequent study, these decapod fossils wereidentified with Thalassina anomala (Herbst).
Fossil Thalassina has been described from

Occurrence and geological noteThalassinid decapods were collected from
the Miocene Bihoku Group exposed at Mune-"Received July 4, 1991; accepted July 16, 1991
852

924.

A decapoda (Thalassina anomala) from Miocene Bihoku Group, Japan853

Pa1eon to1ogica1 description
Toyomalsu mura
130'E 133'E 136'E

1000 m31'

Figure 1.

•1

Locality map.

Order Decapoda Latreille, 1803
Suborder Pleocyemata Burkenroad, 1963
Infraorder Thalassinidea Latreille, 1831
Superfamily Thalassinoidea Latreille, 1831
Family Thalassinidae Latreille, 1831
Gen us Thalassina Latreille, 1806

Type species.- Cancer (Astacus)
malus Herbst by original description .

ano-

Thalassina anomala (Herbst, 1804)
Figures 3-1-3,4-1-4kane, Yuki-cho (Figure 1). In Yuki area,
the Bihoku Group overlies the basement
rocks belonging to the Cretaceous Inai Formation and the Paleozoic Y oshii Group.
Stratigraphic studies of the Bihoku Group in
this area have been made by Imamura (1953),
Itoigawa and Nishikawa (1976) and Matsuoka (1978). In this paper, we use the stratigraphic classification of Matsuoka (1978), who
divided the Group into the Yuki Conglomerate, the linseki Formation and the Kibi
Formation in ascending order.
Thalassina anomala, associated with many
plants and the brachyuran decapod, "Sesarma" sp., occurs in an ill-sorted siltstone bedof the lower member of the linseki Formation.
Itoigawa and Nishikawa (1976) have reported a molluscan fauna represented by the
Cyclina-Hiatula-Siratoria assemblage fromthe Kibi Formation exposed at Kozuike near
the decapod-bearing locality. Matsuoka(1978) has also reported Operculina complanata japonica Hanzawa and such molluscan fossils as the Batillaria and "Vasticardium"-Phacosoma assemblages from the Kibi
Formation. Brackish-water molluscs, Batissa bihokuensis Matsuoka, Melanoides sp.,
Novaculina sp., and plant Comptonia
naumanii (Nathorst) are also known from thelinseki Formation developed at Kozuike(Matsuoka, op. cit.).

Cancer (Astacus) anomalus Herbst, 1804, p.45, pI.
LXII. (non bidi.)
Thalassina scorpionoides Latreille, 1806, p. 51. (nonbidi.)
Thalassina antiqua Bell, 1844, p.455. (nomen
nudum)
Thalassina emeryi Bell, 1845, p.93, figs. a, b, c; A.
Milne Edwards, 1860, p. 349, pI. 15, figs. la, b.
Thalassina gracilis Dana, 1852, p. 514, pI. 32, figs. 5ag.
Thalassina maxima Hess, 1865, p. 37, pI. 7, fig. 18.
Thalassina anomala (Herbst), de Man, 1886, p. 260;
Ortmann, 1891, p. 52; Borradai1e, 1903, p. 541 ;Balss, 1914, p. 88; Etheridge and MacCulloch,
1916, p.7, p.l, figs. 1-3; de Man, 1928, p. 5 ;Van Straelen, 1928, p.64, pI. 12, figs. 1-5; Imaizumi, 1953, p.68, pI. 9; Sakai, 1965MS, p.53,
fig. 11 ; Sankolli, 1970, p. 242, figs. 3, 4; Hirata,
1973, p. 24; Miyake, 1982, p.91, pI. 31, fig. 2;
Kamesaki et al., 1988, p. 25.
Thalassina sp., Imaizumi, 1969, N-7, figs. 8,9 (non.
figs. 7a, b).
Thalassina sp., Forster and Barthel, 1978, p. 302, fig.
8.
Crustacea, Nishikawa, 1972, p. 217, figs. 1-3.
Thalassinoidea fam. gen. et sp. indet., Nishikawa,
1974, p. 3, figs. 17-19.

Material. - MFM391 0 1-MFM391 06.
Repository. - Mizunami Fossil Museum.
Description. - Carapace: rostrum, gastricregion, anterior cardiac region, hepatic region
and anterior branchial region well-preserved.
Rostrum with a weak median furrow, large,
acute triangular, about one-fifth the length of
the gastric region; the lateral margins weakly
dentate or smooth, extending onto the gastricHiroaki Karasawa and Isao Nishikawa

854

1 + - - - - abdomen---4I1IE-E-cara pace-1
posterior
cardiac region

anterior
cardiac region
regionposterior spine
abdominal somite

telsonexopoditebranchial region second
cerbical groove
cerbical groove
oblique ridge
line "b"chelipedline "d" . /__ ----j"---.r1'rf~~[2~~~~~~~-_ / ' dactylus

fixed finger

1st pereiopod
ischium
3rd pereiopod
2nd pereiopod
Figure 2. Terminology of Thalassina anomala (Herbst, 1804) (modified by SankoIIi, 1970)

region as a distinct carina reaching one-third
the length of the gastric region. Anterior
margin of the carapace bears a small spine
behind the orbit, and a carina runs from thebase of the spine parallel with the rostralcarina. Gastric region slightly convex,
smooth except posteriorly where it is covered
with transverse ridges. Cervical groove distinct, deep. Anterior cardiac region divided
into two (AI and A2 regions of Sankolli,
1970; see Figure 2) by the first transverse
furrow; A 1 region armed with 5 small

granulose tubercles on each side; A2 region
covered with 4 small tubercles. Anterior
half of branchial region coarsely decorated
with granulose tubercles which diminish insize posteriorly, the posterior half smooth.
Oblique ridge on the branchial region visible,
finely dentate and the upper part not preserved. Hepatic region densely covered with
granulose tubercles. The region between theline "b" and "d" (see Figure 2) smooth, andthe anterior margin fringed with fine teeth.
Dactylus, fixed finger, propodus, carpus,~ Figure 3.
Thalassina anomala (Herbst, 1804). Bihoku Group. X 1.0. 1, Carapace, abdomen,chelipeds and 1st to 3rd pereiopods, MFM39101 ; 2, Left cheliped, outer view, MFM39101; 3, Rightcheliped, inner view, MFM39103.

924.

A decapoda (Thalassina anomala) from Miocene Bihoku Group, Japan

855

85 6

Hiroaki Karasawa and [sao Nishikawa

924.

A decapoda (Thalassina anomala) from Miocene Bihoku Group, Japan

merus and ischium of cheliped are preserved
and the impressions of some parts remain.
Chelipeds subchelate and equal or subequal ;dactylus long, elongate and the apex sharp;
upper margin bears 15-20, low, small teeth
diminishing in size towards the front, thelower margin is finely dentate. Fixed finger
very short, one-third to one-fourth the lengthof the dactylus; upper margin denticulate
and lower margin smooth. Palm 1.2 to 1.4
times as long as the dactylus and rectangular·in outline; the length of the larger cheliped
about 1.3 times as long as the width, and thatof the smaller one 1.8 to 2.0 times as long as
the width; upper outer margin being provided with 16 to 20 sharp spines directed
forward; upper inner border armed with 20
spinules; lower margin also spinelike; distal
margin near the articulation of the dactylus
bearing fine serrations; outer surface densely
granulate, the denticules varying in size, andbecoming denser on the lower half; a medianlongitudinal carina with conical tubercles
fj.mning from the carpus to the articulation of
the dactylus; inner surface also covered withmany granules; longitudinal row of similar
tubercles on the inner surface along the lower
margin; surface below a longitudinal row oftubercles densely decorated with granules.
Carpus about two-fifths to one-half as long as
the propodus, rectangular, tapering proximally; lower and upper margins marked by somespines; outer distal surface sparsely granulate. Merus as long as or slightly longer thanthe carpus; lower inner and outer margins
bearing some spines. Ischium short, about
three-fourths as long as the merus; lower
margin armed with two sharp spines distally,
and the upper margin irregularly denticulate.
Basis very short with the outer inner margincomposed of a row of small conical granules.
1st pereiopods: dactylus, merus, ischiumand basis are ill-preserved. Dactylus slender,
elongate; upper margin smooth and lower

857

margin finely denticulate; on the outer surface a longitudinal ridge extends along theupper margin. Lower margin of the ischiumis provided with some spines of which the
distal one is the largest. Lower margin of
the basis is also provided with some small
spines.
2nd pereiopods: merus, ischium, basis andcoxa remain. Merus flattened. Ischiumalso flattened about five-ninths as long as themerus. Basis short about seven-tenths aslong as the basis. Both margins of the merus,
ischium and basis smooth.
3rd pereiopods: proximal part of themerus, ischium and basis remain; they areflattened with smooth margins.
Abdomen: telson and six abdominal segment remain. Telson about 1.4 times as longas broad, convex; lateral margin rounded
and the dorsal surface smooth. Uropods
without suture on the endopodite and the
exopodite are not preserved. Sixth abdominal segment about 1.5 times as long as broad,and the dorsal surface longitudinally vaulted,
smooth; both pleura missing.
Remarks. - The living species, Thalassina
scorpionoides Latreille, T. gracilis Dana andT. maxima Hess are synonyms of T. anomala(Herbst) (Borradaile, 1903; de Man, 1928).
One fossil species, Thalassina emeryi Bell ( =T. antiqua Bell, nomen nudum) is also synonymous with T. anomala (Glaessner, 1929;
Van Straeien, 1928).
Imaizumi (1969, figs. 7a, b, 8, 9) reported
the cheliped of Thalassina sp. from the early
Middle Miocene Kawaminami Formation
developed at Kawaminami, Kaga City, Ishikawa Prefecture; his figures 8, 9 are clearly
identified with T. anomala, whereas figures
7a, b are clearly distinguishable from T.
anomala by the presence of a longitudinalcarina on the outer surface of the merus, thelong carpus and its flattened outer surface,
and this specimen can safely be placed in+- Figure 4. Thalassina anomala (Herbst, 1804). Bihoku Group. X 1.0. 1, Abdomen, cheJipeds and1st to 3rd pereiopods, MFM39105; 2, Gastric region, MFM39106; 3, CheJipeds and 1st pereiopods,
MFM39102; 4, Carapace and chelipeds, MFM39104.

858

Hiroaki Karasawa and Isao Nishikawa
tralia (Etheridge and McCulloch, 1916; Forster and Barthel, 1978), New Guinea (Bell,1844, 1845; Van Straelen, 1928). Recent;Japan (Okinawa-jima, Kume-jima, Ishigakijima, Iriomote-jima), India (Bombay, Maha,
Nicobar Islands), Sri Lanka (Ceylon), Burma(Mergui Archipelago), Indonesia (Pulau
Simeulue, Pulau Nias, Sumatra, Tandjoingpandan, Borneo, Celebes, Java, Seram, Lesser
Sunda Islands, Teluk Sarera), Singapore,
New Guinea, Australia (Thursday Island,
Port Moresby, Nicol Bay, Port Curtis,
Sy'dney), Fiji Islands, Philippine Islands (deMan, 1928; Imaizumi, 1953; Sakai, 1965MS).
Measurements. - See Table 1.

Callianassa.
De Man (1928) discriminated two varieties,
T. anomala var. gracilis Dana, 1852 and T.
anomala. var. squamifera de Man, 1915.
Poore and Griffin (1979) elevated one variety,
T. anomala var. squamifera to species level as
T squamifera. Though they quoted dissimilarities in the scaphocerite, sternal ridge
between the pleopods on pleonites 2 to 5 andthe fixed finger, we consider it to be doubtful
as to whether or not these differences deserve
specific rank.
Distributions. - Miocene; Japan (Ishikawa,
Fukui, Hiroshima, Tanegashima) (Imaizumi,
1969; Karasawa, 1990). Pleistocene to Holocene; Japan, Ehime (lmaizumi, 1953), AusTable 1.

Measurements of specimens, in millimeters
MFM39101 MFM39102 MFM39103 MFM39104

Rostrum length
Gastric region length
Gastric region width
Telson length
Sixth abdominal segment:
lengthwidth
Dactylus length of cheliped
Fixed finger length ofcheliped
Propodus length ofcheliped
Pro pod us length ofcheliped
Carpus length of cheliped
Carpus width of cheliped
Merus length of cheliped
Ischium length of cheliped
Dactyl us length of 1st
perciopod
Merus length of 1st
pereiopod
Ischium length of 1st
pereiopod
Basis length of 1st
pereiopod
Ischi urn length of 2nd
pereiopod
Basis length of 2nd
pereiopod
Coxa length of 2nd
pereiopod
Ischi urn length of 3rd
pereiopod
Basis length of 3rd
pereiopod

3.1
31.8
19.7
20.3

22.8
14.0

25.4
17.2

12.6

10.8

9.4

42.6

39.4

34.9

33.7

47.8

26.2

19.2

17.8

21.7

21.3

13.1
14.2
14.5

16.3
14.8
18.8

19.7
14.8

R

29.8
21.8

24.9

26.3

11.0

11.4

5.7

5.8

11.2

9.4

5.2

5.4

8.5

9.1

9.5
5.7

L

L

L
30.6
12.0

R

L
35.2

R

21.7
10.4
26.3

924.

A decapoda (Thalassina anomala) from Miocene Bihoku Group, Japan
Discussion

Disassociated chelipeds of thalassinoid fossils are usually the only parts preserved;
most of our specimens, however, also retainparts of the carapace, walking limbs, or both.
Fossil Thalassina commonly found in the
Pleistocene to Holocene deposits are preserved in a phosphatic hard nodule and they are
considered to be moult or in moulting position (Forster and Barthel, 1978). Preservations are near to that of the Pleistocene to
Holocene Thalassina though our specimens
occur in a siltstone. Judging from theirproximal parts, the limbs clearly remain in
their natural position. The abdomen is bent
forwards. The gastric region is reversed, andthe apex of the rostrum is directed backwards.
Nevertheless, the hepatic and branchialregions remain in the natural position andthey are situated on both sides of the thoracic
sternum, separated from the cardiac region at
the linea thalassinica. Most of the fossil
specimens show their ventral side, and posterior cardiac regions and the first to fifth
abdominal segments are lacking. Considering the nature of preservation, our specimens
also are possibly moults and are thought to besemi-autochthonous.
The decapod crustacean assemblage fromthe stated locality is characterized by the
predominance of Thalassina anomala. Living forms inhabit mangrove swamps in thetropical Indo-West Pacific regions and are
distributed in the northwestern Pacific Ocean
south of Okinawa-jima. They produce several mounds (1 m or more in maximum
height) around the high spring tide zone and.seem to live at depths of 1-2 m below thesurface (Hirata et al., 1973; Macintosh,1988). Referring to these ecological data, theThalassina assemblage at the stated locality isthought to indicate a mangrove swampy
paleoenvironment at the high tide level. The
occurrence of Thalassina, together with molluscs, reef building corals, mangrove pollen
and others (Itoigawa, 1978; Itoigawa and859

Tsuda, 1986; Tsuda et al., 1986; Itoigawa,1989) supports the existence of the tropicalspike in early Middle Miocene time (16 Ma)
in Southwest Japan.
Acknowledgments

Special thanks are due to Professor Junji
Itoigawa of Department of Earth Sciences,
Nagoya University for his kind guidance andhis reading of the manuscript. We wish to
express our thanks to Dr. J.S.H. Collins of
London who gave useful suggestion and who
critically read our manuscript. We arethankful to Dr. Hisakatsu Minei of Department of Agriculture, Kyushu University who
provided us with the living specimens for ourstudy. We thank also Dr. Keiji Matsuoka of
Toyohashi Museum of Natural History for
his useful discussion.
References cited
Balss, H., 1914: Ostasiatische decapoden II. Die
Natantia und Reptantia. Abhand. Math.Physik. Kl. Konig. Bayer. Akad. Wissensch.,
suppl.-vol. 2, no. 10, p. 1-IOL
Bell, T., 1844: On a fossil crustacean from New
Holland. Ann. Mag. Nat. Hist., vol. 14, p.455.
- - , 1845: On the Thalassina emeryi, a fossil crustacean, forwarded by Mr. W.S. MacLeay, from
New Holland. Quart. Jour. Geol. Soc., vol. 1, p.
93-94.
Borradaile, L. A., 1903: On the classification ofThalassinidea. Ann. Mag. Nat. Hist., vol. 7, no.
12, p. 534-55 I.
Dana, J.D., 1852: Crustacea. United States Exploring Expedition During the Years 1838-1842
under the Command of Charles Wilkes, U.S.
N., vol. 13, 1620 p. Philadelphia.
Etheridge, R. and McCulloch, A.R., 1916: Sub-fossil
crustaceans from the coasts of Australia. Rec.
Australian Mus., vol. 11, p. 1-14, pis. 1-7.
Forster, R. and Barthel, K.W., 1978: On recent andfossil Thalassina (Decapoda, Crustacea) andtheir burrows. Neues Jahrb. Geol. Paliiont.
Monath., vol. 1978, no. 5, p. 291-306.
Glaessner, M.F., 1929: Crustacea Decapoda. Fossilium Catalogus, part 41,464 p.
Hirata, Y, Nakasone, Y. and Shokita, S., 1973:
Okinawan decapods and molluscs in color. 144
p. Fudokisha, Okinawa. (in Japanese)

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Hiroaki Karasawa and Isao Nishikawa

Hess, W., 1865: Beitrage zur Kenntnis der
Decapoden-Krebse Ost-Australiens. Arch.
Naturgesch., vol. 31, p. 127-173.
Imaizumi, R., 1953: Thalassina anomala (Herbst)
from Japan. Short papers, IGPS (lnst. Geol.
Pal., Tohoku Univ., Sendai), no. 5, p. 67-76.
- - , 1969: Neogene crustacea from the vicinity ofKanazawa, Ishikawa Prefecture. Atlas ofJapanese fossils, no. 4, N-7, Tsukujishokan,
Tokyo. (in Japanese)
Imamura, S., 1953: In, Imamura et al.. Guide bookof geological excurusion, Kamie, Funasa, Miyoshi, Mirasaka, Shobara and Shokozan. 50 p.
Inst. Geol. Min., Fac. Sci., Hiroshima Univ. (inJapanese)
Itoigawa, J., 1978: Evidence of subtropical environments in the Miocene of Japan. Bull. Mizunami
Fossil Mus., no. 5, p.7-21.
- - , 1989: Tropical spike in early Middle Miocene(ca. 16 Ma) of Southwest Japan. Proc. International Symposium on Pacific Neogene Continental and Marine Events, p.19-26. Nanjing
University Press, Nanjing.
- - and Tsuda, K., 1986: Paleontological significance of tropical molluscan assemblages inJapanese Miocene, with special reference tomangrove swamp assemblages. Monogr. Mizunami Fossil Mus., no. 6, p. 171-182. (in Japanesewith English abstract)- - and Nishikawa, I., 1976: A few problems on
the Miocene Setouchi Series in the northern partof Okayama-Hiroshima Prefectures, southwest
Japan. Bull. Mizunami Fossil Mus., no. 3, p.
127-149. (in Japanese with English abstract)
Kamesaki, S., Nomura, K., Hamano, T. and Mimae,
H., 1988: Crustacea (shrimps and hermit
crabs). Okinawa Kaichuu Seibutsu Zukan, vol.
8, 232 p. Shinseitoshosshuppan, Okinawa. (inJapanese)
Karasawa, H., 1990: Miocene decapod crustacean
assemblages from Southwest Japan with specialreference to the intertidal assemblages of muddy
bottom. Monogr. Mizunami Fossil Mus., no. 7.
(in Japanese with English abstract)
Macintosh, D.J., 1988: The ecology and physiology
of decapods of mangrove swamps. Symp. Zool.

Soc. Lond., no. 59, p.315-341.
Man, J.G. de, 1886: Report on the Podopthalmous
crustacea of the Mergui Archipelago, collected
for the trustees of the Indian Museum, Calcutta,
by Dr. John Anderson, F.R.S. superintendent of
the Museum. Jour. Linn. Soc. London, vol. 22,
p. 1-312.
- - , 1928: The decapoda of the Siboga Expedition.
VII. The Thalassinidae and Callianassidae collected by the Siboga Expedition with some
remarks on the Laomediidae. Siboga Exped.
Monogr., vol. 39a6 , p. 1-187.
Matsuoka, K., 1979: Brackish-water molluscan faunafrom the Miocene Bihoku Group, Hiroshima
Prefecture, Southwest Japan. Bull. Mizunami
Fossil Mus., no. 6, p.27-40.
Miyake, S., 1982: Japanese crustacean decapods
and stomatopods in color. Vol. I, 261 p. Hoikush a, Tokyo. (in Japanese)
Nishikawa, I., 1972: Crustacean fossils from the
Miocene formation of the eastern part of Hiroshima Prefecture. Chigakukenkyu, vol. 23, no.
7,8, p. 217-220. (in Japanese)- - , 1974: Note of fossils from the Bihoku Group(4). Hibakagaku, no. 100, p. 1-4. (in Japanese)
Ortmann, A., 1891: Die Decapoden-Krebse des·Strassburger Museums. Zool. Jahr. Syst., vol. 6,
p.I-57.
Poore, G.C.B. and Griffin, D.J.G., 1979: The Thalassinidea (Crustacea: Decapoda) of Australia.
Rec. Australian Mus., vol. 32, no. 6, p. 217-321.
Sakai, K., 1965MS: Taxonomic studies on Thalassinoidea (Decapoda, Crustacea) from Japan andits adjacent waters. 155 p. Faculty of Agriculture, Kyushu University.
Sankolli, K.N., 1970: The Thalassinoidea (Crustacea, Anomura) of Maharashtra. Jour. Bombay Nat. Hist. Soc., vol. 67, no. 2, p.235-249.
Van Straelen, V., 1928: Crustaces decapodes subfossiles de Merauke (Nouvelle Guinee). Res. Exp.
Sci. Nederland, Nouvelle Guinee, vol. 6, p. 63-68,
pI. 12.
Tsuda, K., Itoigawa, J. and Yamanoi, T., 1986:
Mangrove swamp fauna and flora in the Middle
Miocene of Japan. Palaeont. Soc. Japan, Spec.
Papers, no. 25, p. 129-134.

JA~JIil<O)9=t~fi1tV;~~lIfFJ: 'HiH.. t.:;>J-4'-7 '7 77:/"\",:l : JA~JIil<f1I!;omrm*l!lJ7irMn::.5t;(pi"" GV;~~lIfFf1I!;o ~lIr'8'B (9=tWl9=t~tltW:BJO J: I) g[ 1... t.:;>J- 4'- 7 '7 77:/ "\",:l Thalassinia anomala (Herbst) ~~tI: 1... t.:o T. anomala fi:m1:.fI-r:, 1';/ r- -l1!i:;k~i$0)~*:!{g.00~;/yp-~~1:.~.-r:~I), ~O)g[lli~9=t~tlti-r:~~~t.:o ~O)flO).lli~ 9=t
M9=tmtltWMO)l1!ilfI B*O)~*l¥Jii}jll~~~i"" GomiR*~ . l1!iJlI ;IJ

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 163, pp. 861-878, 9 Figs., September 30, 1991

925. TWO NEW GENERA FROM THE OMMA-MANGANJIOSTRACODE FAUNA (PLIO-PLEISTOCENE) OF
JAPAN-WITH A DISCUSSION OF
THEORETICAL VERSUS PURELY DESCRIPTIVEOSTRACODE NOMENCLATURE
TETSURO HANAI
Osaka Gakuin University, Suita, Osaka 564
and

NORIYUKI IKEY A
Institute of Geosciences, Faculty of Science,
Shizuoka University, Shizuoka 422

Abstract. In this paper, descriptive terms derived from the one continuous sheet theoryof the ostracode carapace are proposed to replace traditional, purely descriptive terms suchas the trichoid sensilla. Traditional sieve versus open, and normal versus radial pore canalclassifications are replaced by a new classification based on constant (c-type) versus variable(v-type) pore canals. The normal versus radial pore canal system of classification is only asubdivision of the v-type pore canal system. The description of surface ornamentation
becomes more theoretical when the disposition of the underlying epidermal cells is taken into
consideration.
Two new genera, Yezocythere (type species, Y. hayashii) and Johnnealella (type species,
J. nopporensis), are described in terms of this proposed idiographic approach.
Key words. Cold water species, idiographic approach in taxonomy, Johnnealella
nopporensis, Omma-Manganji Ostracoda, trichoid sensilla, Yezocythere hayashii.

tive observation, certainly exist and SWIm
actively under our microscope. Therefore
when we try to understand these organisms
via taxonomy, our approach should be idiographic. An idiographic approach is required not simply due to the influence of inductive reasoning in the narrow sense, but rather
as a consequence of the nature of the objects
being observed. In this paper, we develop
this theme of an idiographic approach toostracode classification within the generalcontext of the "continuous sheet theory" ofthe ostracode cuticle and specifically apply tothe description of two new genera.

Introduction

In the process of taxonomic classification of
ostracodes, what we try to classify is not thereal nature of ostracodes, but rather the
ostracodes that we are able to observe andunderstand. This is due in part to the factthat it is impossible to comprehend the natureof ostracodes only by means of a single or a
few biological laws. Yet, ostracodes, which
are understood partly in terms of biologicallaws but mostly by means of purely descrip"Received July 29, 1991 ; accepted August 10, 1991
861

862

Tetsuro Hanai and Noriyuki Ikeya

The history of the study on ostracodes fromthe time of Linnaeus to the present suggests
two fundamental notions. One way of viewing the ostracode organism is that, after
Darwin's theory of evolution through natural
selection found wide acceptance, classification became a method with poor resolving
power for presenting the tested hypothesis ofevolutionary relations among taxa in the formof a classificatory hierarchy. This hierarchy
consists, more or less, of arbitrary "packages"of convenience (Gould and Johnston, 1972),
but it does offer an efficient means of referringto information stored in the detailed biological descriptions (Hanai, 1988, p. 17, 18; Mayr,
1988, p. 270).
The second notion is that the knowledge
resulting from understanding ostracode morphology, which is stored in taxonomic
descriptions, has been advanced through new
findings based on field observations andaided by laboratory techniques. This knowledge has been advanced more throughhypothetical-deductive generalization at thedescriptive level than through explanation by
generalities known as biological laws.
These two notions have vital importance intaxonomy because they inevitably suggest thatthe construction of the classification is merely
a short term goal of ostracode taxonomy,
whereas the ultimate purpose is to understandthe very nature of ostracodes. Further, theimproved comprehension of ostracodes is aprerequisite for a better classification.
To understand the nature of any taxon
through generalization at the descriptive levelis important, because the hypothesis
produced through such generalization may
always have the possibility of not being
confined to one taxon. It may turn out to bea law of great generality covering a wide
range of higher taxa.
As biological understanding of ostracodes
has been advanced mainly through generalization at the descriptive level, the shift frompredominately descriptive to increasingly theoretical methodology has occurred. In theprocess, the purely descriptive terms such as"normal pore" have been replaced with termsof theoretical derivation, such as "opening oftrichoid sensillum" (Hempel, 1965, p. 140).
This has been discussed by the senior author
with documentation and solid evidence fromostracode studies (Hanai, 1988, p. 19).
Viewed from this modern idiographic
approach, our descriptions of ostracodes
appear to be still too conservative and modest
to adequately express the present status of ourunderstanding of ostracodes. In fact, ourknowledge is vast and covers wide fields ofthe biological sciences. It seems, therefore,
appropriate to reconsider the systematic
descriptions of ostracodes in terms of an idiographic approach. The following discussionof the "continuous sheet theory" of the
ostracode carapace will serve as an example.
We hope that our proposed methodology
improves the way ostracode taxonomists present descriptions of new taxa.

Continuous sheet theory of
ostracode carapace
First, we wish to briefly outline the one
continuous sheet theory of ostracode carapaces. This theory was first proposed by
Harding (1964, p. 1,2) and later developed byKornicker (1969, p. 109-119). It holds thatthe valves of an ostracode shell and the soft
cuticle joining them together are one continuous piece of cuticle. This theory replaces
descriptive terms with terms of theoreticalderivation. For example, the descriptive
term calcified portion of the inner lamella isreplaced by the more theoretical term marginal infold of the calcified procuticle. The
same is true for the terms upper and lower
elements of the hinge structure, which are
replaced by the two elements of hinge structure that are the outside and the inside of the
body cavity. Thus, the theory may provide
an explanation for a wide variety of facts (foradditional examples, refer to Hanai, 1988, p.
20-23) that may even include the occurrenceTwo new genera of Omma-Manganji ostracodesof exclusively intact valves in Cambrianostracodes, and the continuous shield-like
carapace of nauplius and metanauplius in
Manawa staceyi (Swanson, 1989, p. 16).
The one continuous sheet theory of the
ostracode carapace, and, the viewing of themarginal area with this theory in mind, will
also suggest as its logical extension the possibility that the basic structure of radial pore
canals is essentially the same as that of normal pore canals. This is simply because
both normal and radial pore canals are openings through one and the same continuous
sheet of the ostracode carapace. The marginal areas are merely an infold of the marginally thickened area of the carapace. The
differences are found only in the elongate
structure and marginal location of the radialpore canals. This was shown by the comparative study of the marginal and normal pore
canals of Bicornucythere bisanensis by Okada(1982, p. 254).
A natural extension of this theory leads us
to reconsider the traditional classification of
normal versus radial pore canals in terms ofanother fact - that two other types of pores
occur that are always quite distinct on asingle carapace, regardless of their location.
This distinction is always sharp and detectable through a wide range of higher taxonomic groups. Thus, we propose here a
new classification of two types of openings forsensilla that is based on probable function.
The first type of pore is termed theconstant-type (c-type) pore, which keeps constant its relative position and its morphologyof the sensilla on one carapace. If it
increases in number through moltings, there is
only an addition of one or so pores. There
are usually about thirty total c-type pores on
the carapaces of genera that have been investigated.
The second is termed the variable-type(v-type) pore, which varies its morphology inaccordance to its location on one carapace
ranging from sieve to sunken sieve type trichoid sensilla. It increases its number dis-863

tinctly during moltings. V-type pores number from about 400 to 500 on a single carapace. The contrast between c- and v-typesof sensilla is seen in the lip type and sieve
type openings for trichoid sensilla in Cythereomotenipponica (Hanai, 1988, p.20-23).
The type 3 pore system of Tsukagoshi (1990,
p. 228) in Cythere schornikovi is equivalent

0

00

0

•

00

•

0
0

0

0

0

0
0

0

3.2
0

0

0

•

0

0
0

0

00

0

0

0

... 3.1

•

0

0

00

0

.r

0

.

0
02.2

0
... 2.3
o
0

'
0

o

0

0

o

0
0

0

. . . . °. . .

0

0-·

0
o / 000

6

0 ...............

/0-- 0
o

0

•
0
0
0
0
/
/\
0
\ / 0 - '0....... 0 - 0
0-000
0

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0

Figure 1. Distribution of c-type (black circles)
and v-type (open circles) sensilla on carapace. 1.
Bicornucythere bisanensis (Okubo, 1975) from
Recent, Shibukawa, Tamano-shi, Okayama Prefecture. Left lateral view of complete carapace of male
(IGSU-O- 102). 2a. Xestoleberis hanaii Ishizaki,
1968 from Recent, Aburatsubo Cove. Complete
carapace of male (IGSU-O-682). Left lateral(slightly dorsal) view. 2b. Left ventrolateral (slightly posterior) view. The lines connecting the sameseries of pores in Figures 1-2a and 1-2b are forreference only to show the relative position of sensillabetween the two figures. Numbers refer to SEM
photos in Figures 2 and 3. Scale bars indicate 100
p.m.

864

Tetsuro Hanai and Noriyuki Ikeya

Two new genera of Omma-Manganji ostracodes
to the c-type, and Tsukagoshi's types 1,2,4
and 5 are equivalent to the v-type. Tsukagoshi's classification of v-type pores is probablya reflection of secondary modification. The
peculiar Ben-type openings without trichoid
structure, and the sunken sieve type openings
for trichoid sensilla in Bicornucythere
bisanensis (Hanai, 1982, p. 10; Hanai et al.,
1985, p.428) correspond to the c-type andv-type, respectively (Figure 1-1). Kamiya's"twisted" type bristle of Loxoconcha(Kamiya, 1989, p.40-42) corresponds to our
c-type sensillum. The distinction between
these two types of sensilla is even more clearly
noticeable in the both pores and setae of
Xestoleberis hanai~ which is illustrated inthis paper as an additional example in Figures 1-2,2,3. Compare the distal half ofseta of c-type pore (Figure 2-2) with that ofv-type pore (Figure 2-3).
As mentioned above, the c-type pores areconstant in morphology, location on carapace
and are virtually constant in number within a
species. However, the c-type pore may varymorphologically from lip type in one genus
to Ben-type in another. In contrast to c-type
pores, the v-type varies in the shape of the
openings from the sieve type to the sunken
sieve type that have variously differentiatedsensory setae. However, their structure isessentially quite constant, always keeping theform of the trichoid sensillum even among
different higher taxa. The origination of the
distinction between c- and v-type pores istraceable ontogenetically back into the young
instars, and may be in a stage younger thanthe differentiation found among v-type pores.
Thus, it is quite likely that the classificationof v type pores based on the various shapes ofthe pore openings and the setae seems to be aclassification based on secondary modifi-

865

cation.
Intermingled distribution of differentiatedv-type pores is seen in various forms of setae.
This can be seen, for example, by comparing
a pore with a long non-branched sensory seta
and a pore with a branched sensory seta in
Bicornucythere bisanensis. The long setae
without apparent branches are mechanoreceptors which might evaluate stimuli from thesurrounding objects, while the short ramifying setae are those that sense weak currents
and vibrations of water. In other cases, amore or less regular transition from the sieve
type to the sunken sieve type pores is traceable on the lateral side of a carapace from thecenter toward the venter, suggesting that the
sunken sieve of bottom-dwelling species ismerely an adaptation of sieve type pores toparticular types of substratum. An example
is seen in Xestoleberis hanaii (see Figure 3).
The traditional sieve versus openclassification of pore canals in several cytheraceans might actually mark the adaptation of
taxa to sand or rocky bottom habitats withturbulent waters versus mud bottom habitats
with calm waters. Therefore, the open or
simple type pores in the traditional sense are
not homologous, including both v-typesunken sieve pores and c-type open pores.
The adaptive distinction appears to reflect thedifference in habitat preference (Hanai et al.,
1985, p.428). This distinction may be formed through traditional phenotypic selection.
Ornamentation and epidermal cells
A second part of this study describes ornamental structures on carapaces in light of thefact that carapaces are the calcified cuticle ofthe integument that directly reflects thearrangement of the underlying epidermal cells+- Figure 2. Xestoleberis hanaii Ishizaki, 1968. Aburatsubo Cove. Recent. Location of four sensillaon carapace is shown in Figure 1-2a, b. I. Figure shows coexistence ofc- and v-type sensilla on anterolateral area of carapace. 2. A complete external view of a c-type sensillum. 3. A complete external view of av-type sensillum. 4. An external view of pore opening of c-type sensillum. 5,6. Tips of two v-typetrichoid sensilla on posteroventral area of carapace. Scale bars of I and 2 indicate 5 j1.m, and bars of 3-6
indicate 0.5 j1.m.

866

Tetsuro Hanai and Noriyuki Ikeya

Two new genera of Omma-Manganji ostracodes(Okada, 1982, p. 232). This situation is quite
unlike the valves of bivalves and it supports
Triebel's (1958, p. 206) classification of broad
and fine ornamentation of the carapace surface. :rhus, the surface ornamentation of
ostracodes can be classified into two elements.
One is the primary relief, which is controlled
mainly by the shape of the main "soft" body(internal organs), and the other is the secondary relief or ornamentation whose
configuration is regulated mainly by the calcium carbonate secretion of underlying epidermal cells.
The overall shape of the main "soft" bodyis determined by the shape of the constricted
appendages and reproductive organs, and its
relief is controlled by the location of various
muscles. The sulcus and central node of thecytheracean genera may directly reflect theconstricted and domed reliefs of the bodysurface, representing the different constructional reaction of the carapace against
the tensile force of the muscles. The generalshape of the "soft" body is a conservative
character being consistent at least within thesame genus. An example is seen in the genus
Paijenborchella where the shape of "soft"body remains constant, whereas the shape ofthe carapace varies from spinose, bean-shaped
to winged (Han ai, 1970, p.697-700). This
type of variation seems also to be foundamong forms of the family Trachyleberididae.
The s~rface of the epidermis which keeps theconservative bean-shape just after ecdysis,
seems to develop into variously shaped adultforms by secretion of the carapace material.
The hard carapace is formed by secretion of
calcium carbonate into the space called the
procuticle between the epicuticle and themembranous layer. The basic pattern of
ridges of reticulation corresponds to the pattern of the boundaries between epidermal867

cells, and is therefore controlled by thearrangement of the underlying epidermal
cells. A plausible explanation for this is thefact that the trichoid sen sill a always open on
the ridge or side of wall that is the boundary
between two adjacent epidermal cells. Whenthey open on the floor of the reticulum, theyoften develop a bridge between the opening
and the ridge. This is simply because a nerve
cell will run between epidermal cells or will
be surrounded by an epidermal cell with asingle nucleus, a.nd will never pierce theepidermal cell.
Architectural construction and taxonomic
comparison of certain features of carapace
ornamentation, including reticulations,spines, and ridges, can be described in relation to the underlying epidermal cells.
Thus, the analysis of reticulation pattern andof the distribution of pore canals acquires abiological meaning. When the numbering ofeach reticulum is carried out in terms of theone sheet theory of the ostracode carapace, we
recommend that the reticulation is traced inV-shaped concentric patterns running parallel to the free margin of the carapace. Thiswas done on certain paleocopid ostracodes byJones (1988, p. 268).
Other complex structural patterns may be
developed secondarily to re-enforce carapace
strength. A sophisticated architectural explanation of these ornamental structures has
been given by Benson (1975, p. 25-44). Typical examples are seen in box-frame ridges of
Schizocythere kishinouyei and in the concentrically disposed corrugate ornamentationof Loxoconcha optima.
In summary, we have applied an idiographic approach to the way in which we study
and classify higher taxonomic groups of
ostracodes, with special reference to the continuous sheet theory of the carapace. We~ Figure 3. All v-type pores of Xestoleberis hanaii Ishizaki, 1968. Aburatsubo Cove. Recent.
Figureshows transition from sieve type trichoid sensillum to open type trichoid sensillum. Location of four
sensilla is shown in Figure 1-2a, b. 1,2. Sieve type. 3,4. Partially sunken sieve type. 5. Completely
sunken sieve type. 6. Marginal trichoid sensilla. Scale bars indicate 0.5 JIm.

868

Tetsuro Hanai and Noriyuki Ikeya

find this approach unifies many aspects ofcarapace morphology-pore types and distributions, ornament, and epidermal cell distributions, for example-allowing a more cohesive and biologically meaningful explanationof the ostracode organism.

The sequence of presentation oftaxonomic descriptions
In describing the carapace of the two new
genera below, the familiar and widely usedcharacter nomenclature will be retained as far
as possible, and the characters will be treatedin a standardized sequence. Thus, thedescription will be given in the order, first on
the general outline, the lateral view fromanterior to posterior, the dorsal and then theventral view, and finally on the internal view.
This is because this more or less standardized
sequence facilitates the comparative study
and is also an efficient way to prevent overlooking anything important (Mayr, 1969, p.
268). At the same time, it is desirable thatthe sequence of characters in the description
will also be biological, coordinating with thedescription of living forms. This positionreflects even on the paleontological description of carapaces. The carapace can be described in order of epidermal organ (integument), sense organs (eyes, sensilla) andlocomotory organs (scars of various muscles).
Thus, although the description of the carapace in paleontology is necessarily a partial
and imperfect presentation of the entire animal, we must keep in mind that we are trying
to present as accurate a restoration of theanimal as is possible based on its carapace
morphology.
~

Systematic descriptions
All illustrated and measured specimens
have been deposited in the collections of theInstitute of Geosciences, Faculty of Science,
Shizuoka University (lGSU). Abbreviations used in the descriptions include: 0=
ostracodes, L = length, H =height, W =width,
S =specimens, R V =right valve, LV = left
valve, X = arithmetic mean, and OR = observed range. All measurements are in mm.
Family Hemicytheridae Puri, 1953
Subfamily Hemicytherinae Puri, 1953
Genus Yezocythere Hanai
and Ikeya, n. gen.
Type species. - Yezocythere hayashii, n.
gen., n. sp.
Etymology. - After Yezo, the ancient nameof Hokkaido.
Diagnosis. - Large tumid carapace. Subcentral tubercle low but distinct. Reticulation of carapace surface divided into three
general areas, anteroventral, posterodorsal
and posteroventral. A nearly straight andsomewhat prominent ridge divides the last
two areas. Bean-shaped eye tubercle marked
posteriorly with a narrow depression but notprojected in lateral view. Middle of threefrontal scars distinctly small. Younger instars with more prominent subcentral tubercle
and posteroventral and posterodorsal swellings.
Remarks. - Urocythereis Ruggieri, 1950 isperhaps the closest relative to this new genus.
In fact, the lateral outline, general pattern ofreticulation, distributional pattern of muscle
scars, and hinge structure are all closely simi-

Figure 4. Yezocythere hayashii. sp. nov. Loc. Hayashi-8l8a, Setana Formation. Lower Pleistocene.
la-b. Stereo pair of lateral view, LV of male (Holotype, IGSU-0-150). x53. 2a-b. Stereo pair of lateralview, RV of male (IGSU-0-151). X 53. Ja-b. Stereo pair of interior lateral view, RV of male (lGSU-O151). x53. 4a-b. Stereo pair of interior lateral view, RV of female (IGSU-0-153). x53. 5. Right lateralview, young instar (Adult-I) (lGSU-0-154). X 53. 6. Right lateral view, young instar (Adult-2) (IGSU0-155). X 69. 7. Right lateral view, young instar (Adult-3) (IGSU-0-156). X 88. Sa-b. Stereo pair of
dorsal view, LV of male (Holotype, IGSU-0-150). X48. 9a-b. Stereo pair of dorsal view, RV of male(lGSU-0-151). x48.

Two new genera of Omma-Manganji ostracodes

869

870

Tetsuro Hanai and Noriyuki Ikeya

Two new genera of Omma-Manganji ostracodes
lar to each other. However, the presence of a
low but distinct subcentral tubercle and thedistinctly small size of the middle muscle scaris diagnostic of this new gen~s. The generalmuscle scar pattern is similar to that ofElofsonella Pokorny, 1955. As far as the
distributional pattern of the frontal scars isconcerned, Normanicythere Neale, 1959 andBaffinicythere Hazel, 1967 are close to thisnew genus in having a distinctly small middle
scar, but are quite different in the distributional pattern of adductor muscle scars.

Yezocythere hayashii Hanai
and Ikeya, n. sp.
Figures 4-1-4-9,5-1-5-5,6.
Urocythereis gorokuensis: Ishizaki, 1971, p.83, 84,
pI. 3, figs. 4, 5.
Urocythereis ? sp. Tabuki, 1986, p. 74, pI. 6, fig.!.
Urocythereis sp. A. Cronin and Ikeya, 1987, p. 82, pI.
I, fig. 17.

Type.-Holotype, LV of male, IGSU-O-150 (Figure 3-1 a, b, 3-8a, b, L =0.99 mm ;H=0.51 mm), lower Pleistocene Setana Formation, loco Hayashi-818a.
Illustrated specimens. - RV of male, IGSU
-0-151 (Figure 4-2a, b, 4-3a, b, 4-9a, b);
RV of female, IGSU-0-153 (Figures 4-4a, b,
5-3a, b, 5-4a, b); RV of young instars, IGSU
-0-154-156 (Figure 5-5-5-7); lower Pleistocene Setana Formation, loc. Hayashi818a. LV of female, IGSU-0-152 (Figure
5-la, b, 5-2a, b, 5-5a, b), lower Pleistocene
Setana Formation, loco Hayashi-45.· RV of
male, IGSU-0-681 (Figure 6), Recent, Mutsu
Bay, station M-lO.
Etymology. - After Mr. Kei-ichi Hayashi
who provided specimens of this species andcontributed to the stratigraphy of the PlioPleistocene formations, southwestern Hok-871.

.- .-..-.-.
Figure 6. Yezocythere hayashii, sp. nov. from
Recent Mutsu Bay (M-IO). RV of male (IGSU-O681). Interior lateral view. Scale bar indicates 100)lm.

kaido.
Diagnosis. - Species is characterized by regular reticulum over most of the carapace
surface, running in parallel longitudinal rows
posterodorsally. Genus is monotypic as far
as is known.
.
Description. - Adult form: Carapace large,
tumid, elongate and sub-quadrangular.
Anterior margin obliquely rounded; dorsal
margin nearly straight, slightly inclined posteriorly; ventral margin slightly sinuate at
middle; posterior margin dorsally andventrally convex and thus slightly sinuate at
middle; posteroventral margin slightly projecting posteriorly. Maximum height atanterior cardinal angle. Viewed dorsally,
carapace elliptical, sides parallel with a low
projection of subcentral tubercle. Subcentral tubercle low but distinct, locatedslightly anterior to middle. Surface coarsely
reticulate. Reticulation tending to align inparallel rows along anterior to anteroventral
margins, in nearly horizontal rows in posterodorsal area, and in posteroventrally directed
oblique rows in posteroventral area.
Marginal area moderately wide, inner margin nearly parallel to free margin. Vestibule+- Figure 5. Yezocythere hayash ii, sp. nov. Specimen IGSU-0-152 from loc. Hayashi-45 and specimen
IGSU-0-153 from loc. Hayashi-818a, Setana Formation. Lower Pleistocene. la-b. Stereo pair of lateralview, LV of female (IGSU-0-152). x53. 2a-b. Stereo pair of interior lateral view, LV of female (IGSU0-152). x53. 3a-b. Stereo pair of lateral view, RV of female (IGSU-0-153). x53. 4a-b. Stereo pair ofnormal pore canal opening with sunken sieve type on lateral surface of central muscle scar area, RV of female(IGSU-0-153). X 1600. Sa-b. Stereo pair of adductor muscle scars, LV of female (IGSU-0-152). X 230.

872

Tetsuro Hanai and Noriyuki Ikeya

moderately deep, crescent-shape along anterior inner margin, shallow along posteroventral margin. Hinge amphidont, withsmooth posterior element. Muscle scars consisting of three frontal scars of which the
middle one is distinctly smaller, four adductor muscle scars of which the two middle
scars are distinctly divided. Two dorsal
muscle scars and a mandibular scar distinct.
Eye tubercle bean-shaped, marked with anarrow depression just behind it. V-typepore canals large, distributed over lateralsurface concentrically, surrounding central
muscle scar area, and opening close to side of
reticulum wall; pores sunken sieve, having acentral protruded micro-pore for the sensory
bristle connected with wall of pore by apier-like ridge. Pore canals opening along
marginal contact zone are straight and numerous along anterior and posteroventral margins. Relatively few c-type simple small
pores also scattered on the ridges.
Sexual dimorphism distinct, male moreelongate and slender than female.
Young instars: Young instars from adult-4 to adult-l were available for study. Lateral outline of carapace changes fromsubtriangular in adult-3 to subquadrangularin adult-I. Subcentral tubercle moving from
central in adult-3 to slightly anterior to
middle in adult-I. Posteroventral and posterodorsal nodes varying from visible in adult-4 to obscure in adult-I. Reticula of anterior half of carapace have a nearly constantnumber; the pattern of arrangement is parallel to anterior margin in about 5 concentricrows. Reticulation of posterior half changing from irregular pattern in adult-3 to regularly radiating pattern in adult-I. Small

second order reticulations dividing large firstorder reticulation into a few to several partitions In dorsocentral area of carapace.
Transformation through ecdysis more prominent in younger instar.
Dimensions. - Measurements of specimensfrom localities Hayashi -818a and 45 are as
follows:s
Adult -¥Adult rl'
Adult-I
Adult-2
Adult-3
Adult-4

RV
LV
RV
LV
RV
LV
RV
LV
RV
LV
RV
LV

N

LX

HX

2
3

0.95
0.95

0.57
0.48

2
4

0.76
0.75
0.58
0.57
0.46
0.46
0.36
0.37

0.43
0.43
0.34
0.34
0.29
0.28
0.24
0.23

10
II
8
8
2
4

LOR

H OR

0.89
0.56-0.57
0.94-0.97
0.99
0.74-0.78
0.74-0.76
0.55-0.60
0.54-0.61
0.43-0.47
0.42-0.47
0.36
0.35-0.39

0.53
0.94-0.96
0.46-0.50
0.51
0.43
0.41-0.45
0.32-0.36
0.33-0.36
0.27-0.30
0.25-0.30
0.23-0.24
0.22-0.25

Stratigraphic range. - Lower Pleistocene to
Holocene.
Remarks. - This species is characteristic ofJapanese lower Pleistocene cold water
ostracode assemblages.
Genus Johnnealella Hanai
and Ikeya, n. gen.

Type species. - Johnnealella nopporensis, n.
gen., n. sp.
Etymology. - In honor of Professor John W.
Neale who gave us much information andhelpful comments on related genera.
Diagnosis. - Large, elongate and with thickly calcified carapace. Hingement holamphidont. Surface foveolate with large coneshaped pits. Most pits have a large pore
canal opening the center. Hemicytherid ad-~ Figure 7. Johnnealella nopporensis, sp. nov. Loc. Hosoda-N2, Shimonopporo Formation. Lower
Pleistocene. la-b. Stereo pair of lateral view, LV of male (IGSU-0-158). x53. 2a-b. Stereo pair oflateral view, RV of male (IGSU-0-159). X 53. 3a-b. Stereo pair of lateral view, LV of female (Holotype,
IGSU-0-160). x53. 4a-b. Stereo pair of lateral view, RV of female (IGSU-0-161). x53. 5. Rightlateral view, young instar (Adult-I) (IGSU-0-164). x53. 6. Right lateral view, young instar (Adult-2)(IGSU-0-165). x69. 7. Left lateral view, young instar (Adult-3) (IGSU-0-J66). x88. Sa-b. Stereo
pair of dorsal view, LV of male (IGSU-0-163). x48. 9a-b. Stereo pair of dorsal view, RV of male (IGSU0-162). x48.

Two new genera of Omma-Manganji ostracodes

873

874

Tetsuro Hanai and Noriyuki Ikeya

ductor muscle scars with widely spaced double scars, and elongate and almost separated
lower adductor scars. Frontal scars consisting of two round scars, of which the upper
scar is small but distinct. Two large mandibular scars are elongate and easily observed; mandibular scar widely separated from
central muscle scar field.
Remarks. - Although the holamphidont
hingement of this genus is quite different fromthat of the genus Fin march in ella Swain, 1963,
the distributional pattern of the muscle scars,
especially that of the widely separated double
adductor scars, and the wide spacing of themandibular scars from central muscle field,
and marked sexual dimorphism suggest a very
close relationship of this genus to the merodont Fin m arch in ella.
Johnnealella nopporensis Hanai
and Ikeya, n. sp.
Figures 7-1-7-9,8-1-8-5,9.
Urocythereis ? gorokuensis: Tabuki, 1986, p. 71, pI.
6, figs. 10-17.
Urocythereis sp. C. Cronin and Ikeya, 1987, p. 82, 84,
pI. 3, fig. 16.

Type.-Holotype, LV of female, 10SU-0-160 (Figures 7-3a, b, 8-3a, b, L= 1.02 mm;
H=0.57 mm), lower Pleistocene Shimonopporo Formation, loco Hosoda-N2.
Illustrated specimens. - R V of female,
IOSU-0-161 (Figures 7-4a, b, 8-4a, b); LV
of male, IOSU-0-158 (Figure 7-1a, b); RV
of male, IOSU-0-159 (Figures 7-2a, b, 9);
LV of male, IOSU-0-163 (Figures 7-8a, b, 82a, b, 8-5a, b); RV of male, IOSU-0-162(Figures 7-9a, b, 8-1a, b); RVofadult-l andadult-2, IOSU-0-164 and 165 (Figure 7-5,7-6); LV of adult-3, IOSU-0-166 (Figure7-7), lower Pleistocene, Shimonopporo For-mation, loco Hosoda-N2.
Etymology. - From the type locality,
Nopporo, near Sapporo City, Hokkaido.
Diagnosis. - Species is characterized byheavily calcified carapace, a raised muscle
platform, large posterodorsal and posteroventral nodes, and ovate fossae. Oenus ismonotypic as far as is known.
Description. - Adult form: Carapace verylarge and heavily calcified. Lateral outline
subquadrangular. Anterior margin broadly
and obliquely rounded; dorsal and ventral
margins nearly straight, slightly sinuate andnearly parallel to each other but slightly
tapering posteriorly; posterior margin nearlystraight or slightly sinuate in its upper half
and narrowly rounded in its lower half.
Highest at anterior cardinal angle. Viewed
dorsally, carapace fusiform with a distinct
subcentral tubercle slightly anterior to middle, strong posteroventral node posterior to
middle and a small posterodorsal node nearposterior cardinal angle. A thick ridge runs
close to anterior margin; another ridge starts
at the eye spot and runs also along the anterior margin, leaving a rather depressed area inbetween the two ridges. Posterior area ofcarapace compressed. Surface foveolate withlarge pits. Pits cone-shaped; most pits witha central pore canal opening. Walls of pits
minutely, horizontally corrugated. Strongly
calcified surface of carapace in between pits
ornamented with extremely fine microreticulations. Ridges of microreticulations
low and minute pits align along inner marginof the ridges.
Marginal infold moderately wide, nearlyparallel to free margin. Vestibule crescentshaped along anterior margin, reaching todepth of two-thirds of width of marginal
infold. Deep vestibule also developed along~ Figure 8. Johnnealella nopporensis, sp. nov. Loc. Hosoda-N2, Shimonopporo Formation. Lower
Pleistocene. la-b. Stereo pair of interior lateral view, RV of male (IGSU-O-162). X 54. 2a-b. Stereo pairof interior lateral view, LV of male (IGSU-O-163). X 54. 3a-b. Stereo pair of lateral surface ornamentation on posterodorsal area, LV of female (Holotype, IGSU-O-160). X 178. 4a-b. Stereo pair of normal
pore canal opening on lateral surface of centroventral area, RV of female (IGSU-O-161). X 880. 5a-b.
Stereo pair of adductor muscle scars, LV of male (IGSU-O-163). X 178.

Two ne w genera of Omma-Manganji ostracodes875

Tetsuro Hanai and Noriyuki Ikeya

876- ••-.

•e•Figure 9. Johnnealella nopporensis, sp. nov.
from lower Pleistocene Shimonopporo Formation,
Loc. Hosoda-N2. RV of male (IGSU-O-IS9).
Interior lateral view. Scale bar indicates 100 jim.

adult-I. Subcentral tubercle being locatedat center of carapace in adult-3. Posterodorsal node is less well developed than the posteroventral node in adult-3, becoming stronger in adult-I. Micro-reticulation first
appearing in adult-3 .
Dimensions. - Measurements of specimensfrom locality Hosoda-N2 are as follows:
Adult .If
Adult d"A:dult-I
Adult-2

posteroventral margin. Hinge holamphidont. Muscle scar field located on wallof subcentral cavity.
Eye spot large and round; located justbelow anterior cardinal angle. V-type pore
canals large and round, scattered over thevalve surface, and always located on floor of
the cone-shaped pits and occupy nearly theentire floor surface. Pores sunken sieve
type; sieve plate forms a dome with a central
micropore for the sensory bristle. Pier-likeridge connecting inner wall of pore with areasurrounding central micropore. Pore canals
with opening along marginal contact zone
straight, numerous along anterior andposteroventral margins. Relatively few simple small c-type pores also scattered over the
micro reticulated surface.
Muscle scar consisting of a row of four
adductor scars of which two middle scars very
widely separated; lower scar elongate andalmost but not entirely separated into twoscars. Two distinct round frontal scars.
Two dorsal musCle scars and two mandibular
scars also easily observed.
Sexual dimorphism distinct; male moreelongate in lateral outline and has a more
pronounced surface ornamentation.
Young instars: Young instars from adult-3 to adult-l were available for investigation.
Lateral outline of carapace changes fromsubtriangular in adult-3 to subquadrate in

Adult-3

S

N

LX

HX

LOR

H OR

RV
LV
RV
LV
RV
LV
RV
LV
RV
LV

3
1
6
5
2

1.00

0.55

0.99
1.00
0.81

0.51
0.54
0.46

0.98-1.01
1.02
0.92-1.02
0.99-1.01
0.78-0.83

0.54-0.56
0.57
0.46-0.53
0.54-0.55
0.43-0.48

2
2

0.64
0.64

0.38
0.38

0.62-0.65
0.62-0.65

0.36-0.39
0.36-0.39

0.50

0.31

Stratigraphic range. - Lower Pleistocene to
Holocene.
Remarks. - This species is characteristic ofJapanese middle Pleistocene cold water
ostracode assemblage.
Acknowledgments

We thank Kei-ichi Hayashi, Ichiro Hosoda
and Ryoichi Tabuki for specimens and stratigraphic documentation; John W. Neale,
Joseph E. Hazel, Thomas M. Cronin and
Katsumi Abe for reviews of various versionsof the manuscript. Hazel and Cronin kindly
read the final manuscript and gave us important advice.
Localities

Materials for this study came from thefollowing outcrops and Recent bottom sampIes:
Hayashi-4S: Lower Pleistocene Setana Formation(2nd fossil bed from the bottom of 2nd cycle ofsedimentation). A junction of Kaigarazawagawa and an unnamed stream, west of Kuromatsun ai, Suttsu-gun, Hokkaido (Lat. 42°39'26"N,Long. 140°16'28" E). Second fossil bed above
the base of the Upper Cycle (Hayashi, 1988).
Greenish gray, very fine-grained sandstone.

Two new genera of Omma-Manganji ostracodes
IGSU-O-152.
Hayashi-818a: The same horizon. Northern
entrance of the Kuromatsunai Tunnel of JR
Hakodate Main Line, south of Kuromatsunai,
Suttsu-gun, Hokkaido (Lat. 42'38'39"N, Long.
140'18' 29" E). Greenish gray, very fine-grai ned
sandstone. IGSU-O-150, 151, 153-156.
Hosoda-N2: Lower Pleistocene Shimonopporo Formation. 1,800 m east of Prefectural Library,
Ebetsu City, Hokkaido (Lat. 43'03'58"N, Long.
14I'32'57"E). Basal fine-grained sandstone.
IGSU-O-158-166.
Recent bottom sample: St. M-IO. 5 km SE of Mutsuyokohama, 31 m in depth, Mutsu Bay, Aomori
Prefecture (Lat. 4I'03.0'N, Long. 14I'12.I'E).
Substratum, blue black compact mud. Sedimentwater interface oxidized and yellowish brown,
and with abundant bivalve shells (Pecten,
Macoma, predominate). IGSU-O-681.
Recent tide pool sample: Rocky tide pool, backyardof the Marine Biological Station, Tokyo University. Aburatsubo Cove, Miura Peninsula, Kanagawa Prefecture (Lat. 139'37'N, Long. 35'09'40"
E). Rocky bottom with calcareous (Corallina
pi/uli/era and Amphiroa dilatata) and otheralgae. IGSU-O-682.
Recent bottom sample: 20 m off the shore of Shibukawa, Tamano-shi, Okayama Prefecture (Lat.
133'48'N, Long. 34'18'E). Mud, water depth
about 10 m. IGSU-O-102.

877

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Kamiya, T., 1989: Differences between the sensory
organs of phytal and bottom-dwelling Loxoconcha (Ostracoda, Crustacea). Jour. Micropalaeont., vol. 8, no. I, p. 37-47.
Kornicker, L.S., 1969: Relationship between the free
and attached margins of the myodocopid
ostracod shell. In, Neale, J.W. ed., The taxo-

878

Tetsuro Hanai and Noriyuki Ikeya

nomy, morphology and ecology of Recent
Ostracoda, p. 109-135, pis. 1-7. Oliver andBoyd, Edinburgh.
Koshimizu, S., Akamatsu, M. and Kitagawa, Y.,
1988: Fission-track dating of the PliocenePleistocene Uranosawa Formation in the
Nopporo Hills, Hokkaido, Japan. Jour. Geol.
Soc. Japan, vol. 94, no. 6, p.461-463. (in Japanese and English abstract)
Mayr, E., 1969: Principles of systematic zoology. 428
p. McGraw-Hill Co. Inc., New York.
- - , 1988: Toward a new philosophy of biology.
Observation of an evolutionist. 564 p. Belknap
Press, Harvard Univ., Cambridge.
Neale, J.W., 1959: Normanicythere gen. nov. (Pleistocene and Recent) and the division of the
ostracod family Trachyleberididae. Palaeontology, vol. 2, pt. I, p. 72-93, pis. 13, 14.
Okada, Y., 1982: Ultrastructure and pattern of thecarapace of Bicornucythere bisanensis (Ostracoda, Crustacea). In, Hanai, T. ed., Studies on
Japanese Ostracoda. Univ. Mus., Univ. Tokyo,
Bull., no. 20, p. 229-255, pis. 16-30.
Ruggieri, G., 1950: Gli Ostracodi delle Sabbie Grige

Quaternarie (Milazziano) di Imola, Partie I.
Giorn. Geol., ser. 2, vol. 21, p. I-57, pI. 1.
Sars, G.O., 1923-1928: An account of the Crustacea
of Norway, with short description and figures of
all the species. Bergen Mus., Bergen, Norway,
vol. 9, Ostracoda. 277 p., 118 pis.
Swain, F.M., 1963: Pleistocene Ostracoda from the
Gubik Formation, Arctic Coastal Plain, Alaska.
Jour. Paleont., vol. 37, no. 4, p. 798-834, pis. 9599, 13 text-figs.
Swanson, K.M., 1989: Ostracod phylogeny and evolution-a manawan perspective. Cour. Forsch.Inst. Senckenberg, no. 113, p. 11-20, pis. 1-3.
Tabuki, R., 1986: Plio-Pleistocene Ostracoda of the
Tsugaru basin, north Honshu, Japan. Bull. Coli.
. Educ., Univ. Ryukyus, no. 29, part 2, p.27-160.
Triebel, E., 1958: Speziel\e Arbeutsgebiete der Mikropalaeontologie. 2, Ostracoden. In, Freund,H. ed., Handbuch der Mikroskopie in der Technik, vol. 2, no. 3, p. 191-236, 8 pis.
Tsukagoshi, A., 1990: Ontogenetic change of distributional patterns of pore systems in Cythere
species and its phylogenetic significance. Lethaia, vol. 23, p.225-241.

*

Aburatsubo nIl~, Ebetsu iI.'6lj, Kaigarazawa Jl.ID!:iR., Kuromatsunai ~mi*J, Mutsuyokoham a ~!!Ii!: ~~, Omma-Manganji~ -7] IM~, Setana lfJi tIlIJ, Shibukawa iY< )11,
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Palaeontological Society of Japan (PSJ) Council Actions
During its meeting on June 21, 1991, the PSJ Council enacted the following changes
to its membership.
New members elected:
Osamu Fujiwara,
Y oshihito Kamata,
Sumito Mizushima,
Tsutomu Nakazawa,
Hiroshi O'shima,

Yasushi Furuta,
Masaichi Kimura,
Akihiro Murata,
Masakazu N ara,
Haruki Sato,

Motoko Higuchi,
Tetsuo Matsumura,Hisao Nakagawa,
Ken Narita,
Shin'ichi Sato,881

Takumi Sato,
Hideyuki Uematsu,

Hiroyuki Shinzawa,
Nobuyuki Yagi.

Hiroyuki Sugie,

Minoru Harada,

Kyoichi Nagata.

Norio Iyoda,
Akio Miki,
Bai-ming, Zhen.

Y oshiyuki Kansha,
Yasuo Sumi,

Resigned members:(Fellow)
Masuoki Horikoshi.
(Ordinary members)
Hiroshi Funatsu,
Membership revoked:(Fellow)
Kouji Minoura.
(Ordinary members)
Toshiaki Futagawa,
Tatsuya Matsumoto,
Hitoshi Ueda,------------------------------- 0 -------------------------------

ErrataKIT AMURA, AKIHISA
Paleoenvironmental transition at 1.2 Ma in the Omma Formation, central
Honshu, Japan. Trans. Proc. Palaeont. Soc. Japan, N.S., no. 162, p. 767780, June, 1991.
On page 780, left-hand column, lines 28-34. Reference of Takayama etal., 1988, printed in an out-of-order sequence, should replace the one
incomplete reference of Takayama et al. printed on the same page,
right-hand column, lines 29-32.

-------------------------------0-------------------------------

1991 if. 8 }jk. t.:.

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

September 30, 1991

New Series No. 163

CONTENTS
TRANSACTIONS
922. Katsumi Ueno: Upper Carboniferous fusulinaceans from the Akiyoshi Limestone Group, Southwest Japan ........................................ 807
923. Kunio Kaiho: Campanian intermediate water benthic foraminifera from
central Hokkaido, Japan .............................................. 828
924. Hiroaki Karasawa and Isao Nishikawa: Thalassina anomala (Herbst, 1804)(Thalassinidea: Decapoda) from the Miocene Bihoku Group, Southwest
Japan .............................................................. 852
925. Tetsuro Hanai and Noriyuki Ikeya: Two new genera from the OmmaManganji ostracode fauna (Plio-Pleistocene) of Japan - with a discussion oftheoretical versus purely descriptive ostracode nomenclature .............. 861
PROCEEDINGS .......................................................... 879