ISSN 0031-0204

a~'~i.~~~.~.!~,

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 118

Palaeontological Society of Japan
June 30, 1980

Co-Editors
Associate editor!taru HAY AMI and Tsunemasa SAITO
Tomowo OZAWA

Officers for 1979-1980

President: Tetsuro HANAI
Councillors: Kazuo ASAMA, Kiyotaka CHINZEI, Takashi HAMADA, Tetsuro HAN AI,
Itaru HAYAMI, Hisayoshi IGO, Kametoshi KANMERA, Tamio KOTAKA, Tatsuro MATSUMOTO, Ikuwo OBATA, Tsunemasa SAITO, Tsugio SHUTO, Yokichi
TAKA YANAGI, Toshimasa T ANAl, Ryuzo TORIY AMA
Members of Standing Committee:

Kazuo ASAMA (Finance), Kiyotaka CHINZEI(General Affairs), Takashi HAMADA (Planning), Itaru HAY AMI (Transactions),
Hisayoshi IGO (Foreign Affairs), Ikuwo OBATA (Membership), Tsunemasa
SAITO (Transactions), Tsugio SHUTO (Special Papers), Y okichi T AKAY ANAGI("F ossils' ')Secretaries: Yoshihiro T ANIMURA (Membership), Kunihiro ISHIZAKI ("Fossils"),
Tomowo OZAWA (Transactions), Toshiyuki YAMAGUCHI (General Affairs),
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Auditor: Sumio SAKAGAMI

The fossil on the cover is Nipponitella explicata HANZAWA, an aberrant
uncoiled fusulinacean from the Lower Permian Sakamotozawa Formation,southern Kitakami, Northeast Japan.

All communications relating to this Journal should be addressed to the
PALAEONTOLOGICAL SOCIETY OF JAPAN,
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Trans. Proc. Palaeont. Soc. Japan, N. S., No. 118, pp. 269-289, pIs. 31-33, June 30, 1980

715.

PERMIAN ECTOPROCTA (BRYOZOA) FROM THE
ABADEH REG ION, CENTRAL IRAN*SUMIO SAKAGAMI
Department of Earth Sciences, Chiba University,
Chiba City, 260

Abstract. The biostratigraphic distributions of some Permian bryozoans in the
Abadeh region, Central Iran are studied. These bryozoans are closely related tothose from the Gnishik·Khachik·Lower Dzhulfian Stages of Armenian Dzhulfa, andsome of them are known also from the Permian of Pamir, Russian platform, Salt
Range, Kashmir, South China and Timor. Fifteen species, including four new
species, of the bryozoans in all are described and illustrated.

1972.

Introduction and Acknowledgements
The present article deals with theresult of biostratigraphic and paleontological studies of bryozoans collected from the
Abadeh region during our field survey madeas one of the research projects on the
Permian/Triassic boundary problems.
Before going further, I would like toexpress my sincere thanks to the staff
members of the Geological Survey of
Iran in Tehran for their hearty arrangement and help in making the research
successful, especially Dr. H. T ARAZ and
Mr. F. GOLSHANI for their kind guidancein the field. I would also like to thankin particular the members of the Japanese party: Drs. Y. BANDO, K. ISHII, M.
MURATA, K. NAKAZAWA, K. NAKAMURA,
Y. OKIMURA and D. SHIMIZU for theirkind cooperations in the field. The fieldsurvey was financed by the Overseas
Scientific Research Fund of the Ministryof Education, Japan for this project inBryozoan biostratigraphy

Most of the bryozoans studied here
were collected by us in the Abadeh region from D and C Sections, which contain
Unit-3 and Unit-4 by T ARAZ (1971, 1974).
These two units were considered to be
the upper part of the "Guadalupian" andthe lower part of the Abadehian by him.
Only three bryozoan species were foundsporadically in the lower parts of A, H
and J Sections corresponding to the lower
part of Unit-6 (the Araxilevis bed of thelower Dzhulfian) by T ARAZ (1971, 1974).
The biostratigraphic distributions of
the Permian bryozoans in the Abadeh
region are shown in Table 1. Fifteen
species in seven genera, including four
new and one indeterminable species werediscriminated. Of eleven known species,
five have been reported originally fromArmenian Dzhulfa, three species from
Pamir, two species from the Russian
platform and Salt Range, and one from
Kashmir, South China and Timor.

* Received December 17, 1979; read Jan. 22,
1979 at the Annual Meeting of 1979 at Fukuoka.
269

270

S'u,mio SAKAGAMI
Table 1.

Biostratigraphic distribution of t he br yozoans in the A badeh sections.
(Columnar section was comp il ed by the Research Group)
IIGuada lupian"Gnishik

AbadehianeKhachikL. Dzhulfian

CApaxi lev i s II. )Speciesx

Fistuiipom elegantula NI Kl FOROVAFi stulipora pseudomonticulosQ SA10\Gf\\lI, n . sp .

x

EridopoY"'Q c f . pal'Qsi tica (h'AAGEN ~U1d \\'FJVr2 EL)

Hexa gone lla toptuosa hlAAGEN ruld WENTZELx

Stenopora ? n. sp .
Al'Qxopo1"Q a 2~axensis ~ KJRO ZOVAx

Septop02'Q tar>Q z i SAKAGA\II, n . sp .

XxSeptopora lineata NIKIFOROI'A
Polypora tubulosa NIKI FOROVA
POlyp01'Q soyanensis~ IOROZO\'A

PolypoI'Q abadehensis SAKAGA'II , n. sp .

Polypom magni cava mROZOVA

x- -- -x-- x--- - X--- X
x--- -x
X

PolYPoY'Q str iata SAKAC~t\i\ II, n . sp .

x

Polypom aff . damshamensis NIKI FOROVAXPolypom ? s p. indet .

Some of the present specimens are
identical with the original specimen of
Hexagonella tortuosa WAAGEN and WE NTZEL which was described from the Middle
Productus Limestone of the Trans-Indus,
Pakistan.
Araxopora araxensis NIKIFOROVA was originally described fromthe Gnishik to Khachik of Armenian
Dzhulfa. Recently it was reported also
from the Permian of Sainbeyli, T urk ey
by SAKAGAMI (1976). Further, according
to MOROZOVA (1970), Stenodiscus granularis YANG from the Permian of China
may be identical with A. araxensis. Polypora tubulosa NIKIFOROVA and Septopora
lineata NIKIFOROVA were both described
originally from the Gnishik of Armenian
Dzhulfa and the former species also fromthe Permian of Sainbeyli, T urkey byx

SAKAGAMI (1976). Eridopora parasitica(W AAGEN and WE NTZEL) was described
originally from the Middle Productus
Limestone. I am of the opinion that
E1'idopora major BASSLER from Timor
may be synonymous with E. parasitica.
Stenopora? n. sp. coincides with S.? lwshmirensis SAKAGAMI (1980, MS) from the lower
part of the Zewan Formation of Kashmir.
Thus it is noted that all of the bryozoan
species from C-O horizon of the lower partof Unit-4 have been hitherto known fromthe Pre- Abadehian. However, this unit
contains Codonofusiella and Reichelina,
which characterize the Abadehian age asshown in Table 1 that was compiled by our
Research Group (M S). Polypora soyanensis MOROZOVA and Polypora magnicava
MOROZOVA were both described originally715.

Permian Ectoprocta from the Abadeh region

from the lower Kazanian of the Russian
platform, USSR. According to MOROZOVA(1970), the Kazanian of the Russian platform can be correlated to the Khachikof Armenian Dzhulfa. Among two bryozoan species from the C-15 horizon of
Unit-4, one is thought to be an affinis of
Polypora darashamensis, which was described originally from the lower Dzhulfian of Armenian Dzhulfa, and the otheris a new species: P. striata. Accordingly
the geological age of this horizon can not
be decided certainly. From Unit-5, any
bryozoan remains have not been found.
The occurrence of Fistulipora elegantula
NIKIFOROVA, which has been known fromthe Lower Dzhulfian of Armenian Dzhulfa,
ensures that Unit-6 in the Section H canbe correlated to the lower Dzhulfian.
According to MOROZOV A (1970), this species may be identical with Dybowslliellasinensis YANG and Loo which was described from the upper part of the Nuoyinhe Formation (U pper Permian) of Qilianshan, North China. The other two species:
Fistulipora pseudomonticulosa andPolypora
sp. were collected from the lower Dzhulfian of the Sections Hand ], respectively.
Description of Species

Genus Fistulipora McCoy, 1850

Fistulipora elegantula NIKIFOROVA
PI. 31, Figs. 1-3.

1933. Fistulipora elegantula NIKIFOROVA, p.
11, 12, pI. 2, figs. 1-9, text-figs. 5, 6.
1962. Dybowskiella sinensis YAXG and Loo,p. 23, pI. 3, figs. 10, 11.
1970. Fistulipora elegantula: MOROZOYA, p.
62, pl. 1, figs. 3, 4.
1975. Fistulipora elegantula: GORJUNOVA, p.
44, pI. 1, fig. 4.

In surface specimen, zoarium consistingof straight, cylindrical hollow stem, about271

45 mm in length and 13 mm in diameter.
Well developed monticules regularly arranged, their intervals about 6 to 7 mm.
Zooecial apertures densely distributed
without any special regularity except for
the flat areas of monticules. Diameterof monticule about 2 mm. Thickness ofzoarium ranging from 2.5 to 3.3 mm.
In tangential section, zooecial tube
nearly circular with well developed lunarium, the inside longitudinal diameter excluding lunarium 0.303 mm average, ranging from 0.280 to 0.320 mm (12 measurements); outside diameter including lunarium 0.391 mm average, ranging from0.350 to 0.420 mm (30 measurements) ; andinside transverse diameter 0.272 mm average, ranging from 0.230 to 0.300 mm (20measurements); outside diameter 0.452
mm average, ranging from 0.400 to 0.550
mm (30 measurements). Usually 4 to 4.5
zooecia per 2 mm diagonally. Well developed lunarium, typically crescentic shapedin tangential section, usually disposed at
proximal part of zooecial margin, occupying about one-fourth of zooecial circumference. Thickness of lunarium including
fibrous tissue ranging from 0.050 to 0.070
mm in mature region, thickness of darkline of lunarium about 0.0l5 mm, length
between the tips of lunarium ranging
from 0.200 to 0.230 mm. Vesicular tissue
regularly arranged, usually one vesicle
between adjacent zooecia near surface,but one to three in inner section, 7 to 9
vesicles per mm diagonally.
In longitudinal section, zooecial tubes
run for a short distance along coenelasma,
but curve gradually upward, making aright angle in mature region. Diaphragms
thin, nearly straight or slightly concave,usually 5 to 6 diaphragms countable in a
tube. Interspaces between diaphragms
usually 0.200 to 0.250 mm, but 0.500 to
0.750 mm in middle part of tube. Interzooecial tissue consisting of regularly

Sumio SAKAGAMI

272

arranged vesicular tissue which is quadrate but seems to be depressed in outerzone. 9 to 11 vesicles in inner zone, but
13 to 16 in outer zone per mm longitudinally.
Remarks.-Conspicuously developed lunarium with very sharp and juted tips,
and regularly arranged vesicular tissue
are characteristic of Fistulipora elegantula.
The present specimen agrees with Fistulipora elegantula which was originallydescribed by NIKIFOROVA (1933) from thePermian of the Djhulfa region. Later
MOROZOVA (1965 in RUZHENCHEV andSARYCHEVA; 1970) revised it and madethe horizon of its occurrence clear, thatis the lower Dzh ulfian (the A raxilevis
horizon). The present species was also
described from the Pamirian (Upper Permian) of Southwestern Darvaz of Pamir
by GORJUNOV A (1975).
Horizon and SPecimen 110.- The Araxilevis horizon of Section H. Reg. nos.
DESC-80001, DESC-80002 and DESC80003.
Fistulipora pseudomonticulosa
SAKAGAMI, n. sp.
PI. 31, Figs. 4-6.

In surface specimen, zoarium consistingof straight, subcylindrical hollow stem,
more than 70 mm in length and 10 to 12
mm in diameter. Monticules more or lessregularly arranged in space of 6 to 9 mm.
Zooecial apertures densely distributed
without any special regularities except
for the fiat areas of monticules. Thickness of zoarium ranging from 1.0 to 1.5mm.
In tangential section, zooecial tube subcircular or ovate. Its inside longitudinal
diameter excluding lunarium 0.339 mm
average, ranging from 0.290 to 0.390 mm;
outside diameter including lunarium 0.364

mm average, ranging from 0.310 to 0.410
mm (25 measurements) and inside transverse diameter 0.191 mm average, ranging
from 0.l70 to 0.210 mm; outside diameter
0.269 mm average, ranging from 0.240 to
0.310 mm (25 measurements). Usually 4
to 5 zooecia per 2 mm diagonally. Thin
lunarium disposed at proximal part ofzooecial margin, occupying about onefourth of zooecial circumference, its thickness about 0.010 to 0.020 mm. Vesicular
tissue rather coarse and regular in size,
usually one occasionally two or threevesicles between adjacent zooecia, and 6
to 8 vesicles per mm diagonally.
In longitudinal section, zooecial tubes
run for nearly parallel direction to coenelasma, curve gradually upward, making
a large angle to surface. Usually three
diaphragms disposed in lower to middle
part of a zooecial tube. Interzooecial
tissue consisting of regularly arranged
vesicular tissue which is quadrate but insome cases depressed. About 10 vesicles
per mm longitudinally.
Remarks.-The present new species is
not unlike Fistulipora monticulosa NIKIFOROVA which was known from the lower
Dzhulfian of Trans-Caucasia in the generalappearance, and especially it is similar tothe specimen illustrated by MORozov A(1970, fig. 2 of plate I). However, thepresent species differs from F. monticulosa
by the fewer number of diaphragms,
namely, there are usually only 3 diaphragms in a zooecial tube of the present
new species but about 10 or more diaphragms in F. monticulosa.
Horizon and SPecimen no.-The Araxilevis horizon of Section A. Reg. nos.
DESC-80004 (holotype), DESC-80005 and
DESC-80006.
Genus Eridopora BASSLER, 1929

715.

Permian Ectoprocta front the A.badeh region

Eridopora cf. parasitica (W AAGEN
and WENTZEL)
PI. 31, Figs. 79.

Compare :1886. Fistulipora

parasitica WAAGEX andp. 923, 924, pI. 45, fig. 6; pI.
105, figs. 1-4.
1929. Eridopora major BASSLER, p. 52, pI.
225, figs. 1-4.
1975. Eridopora major: G(jRJlT~O\'A, p. 45, 46,
pI. 3, fig. 1.
WE:-iTZEL,

Zoarium incrusted on the obverse surfaceof Septopora lineata and Polypora tubulosa,and composed of relatively thick layer,
varying from 2 to 2.5 mm in thickness.
Surface of the zoarium could not be observed.
In tangential section, zooecial tube
roundly triangular with moderately developed lunarium. Its inside longitudinal
diameter excluding lunarium 0.416 mm
average, ranging from 0.360 to 0.460 mm(23 measurements) in outer part of zooecia I tube, 0.316 mm average, ranging from0.280 to 0.360 mm (20 measurements) in
the inner part; and inside transverse
diameter 0.351 mm average, ranging from0.300 to 0.400 mm (23 measurements) inouter part of zooecial tube, 0.313 mm
average, ranging from 0.290 to 0.350 mm(20 measurements) in the inner part.
Usually 4 to 4.5 zooecia, but 4.5 to 5 ininner part, per 2 mm diagonally. V-shaped
lunarium disposed at proximal end of zooecial tube. The thickest part of lunarium
measured 0.052 mm average, ranging from
0.040 to 0.070 mm (23 measurements) inouter part of zooecial tube, but very thin
in inner part. In the opposite side oflunarium in a tube, a pair of small projections occasionally visible. Vesicular
tissue not so regularly in size and arrangement, usually one to two vesicles between
adjacent zooecia in outer part, but two to
three in inner part, and 8 to 10 vesicles per273

mm diagonally.
In longitudinal section, zooecial tubes
run for a short distance along coenelasma.
but curve rather rapidly upward, growstraightly and making a right angle inmature region. Many diaphragms disposed
throughout tube. Interval between diaphragms usually 0.100 to 0.250 mm. There
are 6 to 8 diaphragms in inner part oftube, but not any diaphragms in about
0.5 mm of outer part of tube. Interzooecial tissue consisting of not so regularly arranged vesicular tissue and depressed quadrate in form. About 9 to 11
vesicles per mm longitudinally.
Remarlcs.- The present species is characterized by the subtriangular zooecial
aperture with lunarium developed at theposterior end of aperture in tangential
section. These characters show that thepresent species belongs to Eridopora.
Further, the most conspicuous characterof the present species is the existence ofa pair of small projections at the opposite
side of lunarium in the tangential sectionof tube. BASSLER (1929) recognized it asa lunarium in the illustration of Fistulipora parasitica by W AAGEN and WENTZEL(1886), and he distinguished Eridopora
major from F. parasitica by the differenceof the mode of lunaria. However, in theillustration of E. major by BASSLER (1929),
the small but distinct projections disposedat the opposite side of lunarium can beobserved. Therefore, Eridopora major
may be a junior synonym of E. parasitica.
E. major was recently described also fromthe upper part of the Lower Permian ofCentral Pamir by GORJUNOVA (1975).
Horizon and Specimen no.-C-O bed of
Unit-4 in Section C. Reg. nos. DESC80007, DESC-80008a and DESC-80009a.
Genus Ifexagonella W AAGEN
and WENTZEL, 1886

Sumio SAKAGAMI

274

proximally parallel to mesotheca for avery short distance, then curve upward,
making a right angle with outer surfaceof zoarium. Diaphragms usually one to
two times of zooecial diameter, irregularly spaced, 5 to 7 per 2 mm length ofzooecial tube. Vesicular tissue relatively
coarse and irregular in form, size andarrangement in immature region, but moredepressed, rectangular or rounded quadrate and regularly arranged in longitudinal series in middle region, but becoming relatively coarse again in mature
region, 10 to 12 vesicles in the middle.
Vesicular tissue replaced by dense lamellar tissue near surface.
Remarli~"·.-Although some thin sections
made from only one zoarium were examined, the present form can be easily
identified with Hexagonella tortuosa
W AAGEN and WENTZEL which was originally described from the middle divisionof Productus Limestone at Bilot andGulami of the Trans-Indus of Pakistan.
The present species can be distinguishedfrom Hexagonella ramosa by the differentmode of growth and the more irregular
network of hexagonellid ridge.
Horizon and SPecimen no.-Base of
Unit-3. Reg. nos. DESC-80010, DESC80011 and DESC-80012.

Hexagonella tortuosa W AAGEN

and WENTZEL
PI. 31, Figs. 10-12.
lortuosa
WAAGE:'-iandp. 914, 915, pI. 108, figs. 1,

1886. Hexagonella
WE:->TZEL,

3-5.

Zoarial surface not observed. Zoarium
bifoliate, consisting probably of subcylindrical branch. In tangential section
near surface, the lines of hexagonellidridges consisting of polygonal areas but
not regular pattern can be observed.
Thickness of zoarium 6 to 7 mm.
In tangential section, zooecial tube
nearly circular to oval, their inside diameter 0.195 mm average. ranging from
0.160 to 0.270 mm (200 measurements on
two sections), usually 4.5 to 5 zooecia per
2 mm diagonally. Vesicular tissue irregular in shape and size in inner zone,
usually one to three vesicles between
adjacent zooecia, but undeveloped in outerzone because it is filled with thin fibrous
tissue. Usually 7 to 9 vesicles per mmhorizontally. Lunarium usually absent
but occasionally developed weakly, occupying about one third of zooecial circumference, the thickness very thin.
In longitudinal section, zooecial tubes

Table 2.
Specimen no.

80013

Orientation of section
L
Diameter of zoarium
5.0
Diameter of immature zone
2.2
Thickness of mature zone
1. 3-1. 5
Diameter of zooecium (shorter)
I O. 160-0.240
Diameter of zooecium (longer)
I 0.160-0.310
Diameter of mesoecium (shorter)! 0.030-0.070
Diameter of acanthoecium (outer)I less than 0.030
No. of zooecia per 2 mm longitudinally i ca. 6-7

Measurements of Stellopom?
80014

Tr
5.5
2.0
1.6-1. 9-----L: longitudinal section,

Tr: transverse section,

T: tangential section,

80015aL
5.0
2.0
1. 5-1. 6
0.200-0.260. 0.200-0.300

0.040-0.080
ca.6
Ob: oblique section.

715.

Genus Stenopora LONSDALE, 1844

Stenopora? n. sp.
PI. 32, Figs. 1, 2.

19S0. Stenoporc.?

275

Permian Ectoprocta from the Abadeh region

kashmirensis

SAKAGA~II(MS)

Among more than ten thin sections ofzoaria, seven sections were examined.
Zoaria consisting of cylindrical stems,
rarely branched, ranging from 4.0 to 5.5
mm in diameter. Diameter of immaturezone 2.0 to 2.5 mm and length of maturetube ranging from 0.8 to 1.9 mm. Ratioof half diameter of zoarium to thicknessof mature zone varying from 1.3: 1 to
2.5: 1 measured on seven specimens.
In tangential section, zooecial tube
nearly circular or rounded polygonal inmature region. Diameter of zooecial tube
measured along transverse direction ranging from 0.160 to 0.280 mm and anotherdiameter measured along longitudinal
direction ranging from 0.160 to 0.310 mm.
Zooecial apertures may be arranged
without special regularities. Mesoecia
rarely present, rounded, oval or roundedpolygonal, the shorter diameter varying
from 0.030 to 0.100 mm. In mature region, thick walls of adjacent zooecial

tubes separated by a dark divisional wall.
Small acanthoecia usually disposed at eachpoint of intersection of zooecial tube, theouter diameter less than 0.030 mm andthe central pore is extremely small.
In longitudinal section, zooecial tubes
trend parallel to longitudinal direction of
zoarium in inner region, curve more
rapidly outward at boundary between
inner and outer regions of zooecial tube
and making nearly right angle with surface. Zooecial wall thin in inner region,but making rapidly thick in outer region.
No diaphragm but usually one superior
hemiseptum, occasionally two, well developed at the boundary between inner andouter regions of zooecial tube.
In transverse section, inner part ofzooecial tube polygonal with very thinzooecial wall. Other characters in thissection can also be observed in longitudinal section.
Measurements.-See Table 2.
Remarks.- The present form is identicalwith S.? kashmirensis SAKAGAMI (MS)from the lower part of Zewan Formationof Kashmir, India in the essential characters, but the diameters of zoaria andzooecia of the present form are bothslightly larger than those of the Kashmir
specimen.

kashmirensis SAKAGA~II (in mm).

SOO15b

SOO16a

S0017

S0016b-------

Ob(L)
5.0
2.2
1. 4-1. 5
0.IS0-0.270
0.180-0.300
0.050-0.100
less than 0.030
5-6

L,T
5.5
2.0-2.2
1. 5-1. S
0.170-0.2S0
0.lS0-0.300
O. 040-0. 100
less than 0.030
5-6
~~-

--

----~-

L,T
4.0
2.3

0.8-0.90.160-0.280
0.180-0.300
0.040-0.100
less than 0.030
5-6

Tr

5.2
2.5
1. 2-1. 4
0.170-0.260

S1tmio SAKAGAMI

276

Horizon and SPecimen llo.-C-O bed of
Unit-4 in Section C. Reg. nos. DESC80013, DESC-800l4, DESC-80015a, b,
DESC-80016a, band DESC-80017.

Genus Araxopora MOROZOVA, 1965
A raxopora araxensis (NIKIFOROV A)
PI. 32, Figs. 3-5.

1933. Batostomella spinigera var. araxensis
NIKIFOROVA, p. 13, 14, 36, pI. 4, figs.
1-4.
1958. Stenodiscus granularis YA"G, p. 133,
134, pI. 11, figs. 2-8.
1965. Araxopora araxensis: MORozovA, p. 186,
pI. 25, figs. 4, 5, pI. 26, fig. 5, text-fig.
20.
1975. Araxopora araxensis: GORJU);OVA, p. 59,
pI. 9, fig. 3.
1976. Araxopora araxensis: SAIC\.G,\~II, p. ·102,403, pI. 42, figs. 8-11.

Two typically longitudinal, one tangential and one oblique sections were examined. Zoarium consisting of straight,
cylindrical stem, 5.3 mm in diameter. Allof inner region obliterated by the secondary alteration.
In longitudinal section, zooecial tubes
may run parallel to longitudinal directionof zoarium in inner part, but rapidly bendat the inner edges of mature region, andTable 3.

go straight throughout mature region
reaching the surface at an angle of about900. Length of zooecial tube of mature
region ranging from 0.9 to 1.1 mm. Zooecial wall very thin in immature region
but becoming thick rapidly and consisting of dark, coarse, laminated fibrous
tissue in mature region. Diaphragm disposed at outer edge of immature tube
and one to two, occasionally three inmature tube.
In tangential section, zooecial tube oval
in mature region but irregularly polygonal in the nearest part to surface.
Zooecial diameters at the middle level ofmature tube, one measured along horizontal (shorter) direction 0.156 mm average, ranging from 0.110 to 0.200 mm (50measurements) and another measured
along longitudinal (longer) direction 0.245
mm average, ranging from 0.200 to 0.300
mm (50 measurements). Zooecial apertures arranged longitudinally but not so
regularly. Mesoecia irregularly arranged,
oval to somewhat irregular in form,
variable in size, 0.070 mm average. ranging from 0.030 to 0.110 mm (15 measurements). Usually 8 to 12 zooecial apertures and 5 to 8 mesoecia in a field of one
sq. mm. Acanthoecia present, 5 to 8 in
one sq. mm, its inside diameter ranging
from 0.010 to 0.030 mm.

Measurements of Araxopora araxensis (NIKIFORUVA) (in mm).

Specimen no.

80018

80019

80020
--1-

Orientation of zoarium
Diameter of zoarium
Diameter of inner region
Thickness of outer region
Diameter of zooecia (shorter)
Diameter of zooecia (longer)
Inner diameter of mesoecium
Outside diameter of acanthoecium
Inner diameter of acanthoecium

T

L(T)

L(T)

5.3
3.5
0.9-1. 1
0.110-0. 180

5.3
3.5
0.9-1. 1
0.140-0.200

0.030-0.090

0.040-0. 110
0.050-0.100
0.010-0.030---------

I

0.110·0.200
0.200-0.300
0.040-0. 110
0.050·0.120
0.010·0.030----------.--

715.

Permian Ectoprocta jTom the Abadeh region

Measurements.-See Table 3.
Remarks.-Owing to the very thin immature wall, the inner zone of zoariumof the present species is obliterated inmany occasions. The present form isidentical not only with the originallydescribed specimen from Armenian
Dzhulfa of Trans-Caucasia but also with
the Turkish specimens, which was recently described by SAKAGAMI (1976), in allof the essential characters. The present
species was also described from the
Murgavskian (the middle part of Upper
Permian) of southeastern Pamir by GORJUNOVA (1975).
Detailed comparisons
between the specimens hitherto described
were discussed by SAKAGAMI (1976).
Horizon and SPecimen no.-D-16 bed of
Unit-3 in Section D. Reg. nos. DESC80018, DESC-80019, DESC-80020 and 80039.
Genus Polypora McCoy, 1844
Polypora tublliosa NIKIFOROVA
PI. 32, Fig. 7.
1933. ?ulypora tubulosa NIKIFOROVA, p. 15,
20-22; 37,38, pI. 3, figs. 1-6, pI. 4, figs.
1-3.
1970. ?olypora tubulosa: MOROZO\,A, p. 213,
216, pI. 49, figs. 3, 4.
1976. ?olypora tubulos1-: SAKAGA\H, p. 403405, pI. 42, fIg. 5.

Three tangential sections of zoaria were
examined. Zoarium expanded laterally,
may be fan shaped. Straight branches
connected by dissepiments at regular intervals ranging from l.000 to 1.200 mm
measured from center to center of dissepiments. Bifurcation of branch not frequent. Width of branch wider than that offenestrule, ranging from 0.550 to 0.750
mm, after bifurcation about 0.350 mm,
before bifurcation about 0.900 mm, usually
8 to 11 branches per 10 mm horizontally.

Intervals between branches ranging from
0.875 to 1.250 mm measured from centerto center of branches. Fenestrule elongated elliptical in outline, width usually
ranging from 0.375 to 0.500 mm, occasionally 0.625 mm, length ranging from 0.625
to 0.750 mm, 9 to 10 fen est rules per 10 mm
length of branch. Dissepiment narrower
than that of branch, width ranging from0.300 to 0.650 mm. Zooecial tubes arranged usually in 4 to 5 intersecting
longitudinal rows, but 3 rows in shortdistance after bifurcation and 6 rows at
just before bifurcation. In tangential
section, zooecial tube hexagonal, rhomboidal or somewhat irregularly formed atlower and middle levels of branch, andcircular at upper level. Zooecial diameternear surface ranging from 0.100 to 0.120
mm. Zooecial apertures 14 to 18 per 5mm length of one row, usually 4 apertures per fenestrule. Interval between
zooecial apertures in longitudinal series
from center to center ranging from 0.250
to 0.380 mm. Nodes may be disposed ateach intersection of zooecial apertures
but indistinct. Interzooecial materials
consisting of dark, very fine fibrous tissue
with closely arranged fine granules.
Stereom covering the reverse side consisting of inner tissue with 3 to 5 prominent capillary canals on which large
capillaries regularly arranged in longitudinal series, 10 to 12 per mm length of
one row, and outer thick, dark, fine fibrous
tissue with very fine granules.
Meshwork !ormllla.-8-11/9-10/ /14-18/
4(3-6).
J\;/easllrements.-See Table 4.
Remarlls.-The most conspicuous characteristic of Polypora tuiJlI/osa is in
having the very prominent capillary
canals. The present form agrees wellwith the original specimen in all the essential characters. Polypora tllimiosa wasoriginally described by NIKIFOROVA (1933)

278

Sumio SAKAGAMI

Table 4.

Measurements of Polypora tubulosa NIKIFOROVA (in mm).

Specimen no.

80021

80022

80008bNo. of branches per 10 mm horizontally
No. of fenestrules per 10 mm length
No. of zooecia per 5 mm length
No. of rows of z(oecia
No. of zooecia per fenestrule

8-10
9-10
15-16
4 (3-5)
3-4

10-11
9-10
16-18
4-5 (3-6)40.550-0.600(0.350-0.900)
1. 000-1. 075
1. 000-1. 200
0.300-0.250
O. 100-0. 110
0.250-0.280
ca.0.025

9-11
9-10
14-16
3-4
3-4

Width of branch

0.625-0.875

Interval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)Distance between nodes (c. to c.)

1. 125-1. 250
1. 025-1. 125
0.375-0.500
0.100-0.120
0.300-0.380

from the "Upper Carboniferous" of Armenian Dzhulfa, but later MOROZOVA (1965,1970) revised it as an Upper Permian
Dzhulfian species.
Horizon and Specimen no.-D-16 andD-17 beds of Unit-3, and C-O bed of
Unit-4 in Sections D and C. Reg. nos.
DESC-80021, DESC-80022, DESC-80008b
and DESC-80009b.
Polypora soyanensis MOROZ OVA

0.600-0.750

O. 875-1. 250
1. 000-1. 200
0.375-0.650
0.100-0.120
0.320-0.380

PI. 32, Fig. 6.
1970. Polypora soycznensis MORozovA, p. 211,

212, pI. 48, figs. 3, pI. 49, fig. 2.

Five tangential sections of fragmentary
zoaria were examined. Straight branches
nearly parallel to each other, connectedby dissepiments at regular intervals ranging from 0.875 to 1.250 mm measured from
center to center of dissepiments. Bifurcation of branch very rare. Width ofbranch wider than that of fenestrule,
Table 5.

Measurements of Polypora
80023

80024b

14-16
9-10
18-19
3-4
4

14
10
19?
4
4

Width of branch

0.350-0.500

0.375-0.450

Interval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)

0.625-0.875
1. 000-1. 100
0.375-0.500
0.090-0.100
0.250-0.325

0.625-0.750
O. 950-1. 000
0.300-0.350
0.090-0.100
0.250-0.300

Specimen no.
------~----------

No. of branches per 10 mm horizontally
No. of fenestrules per 10 mm length
No. of zooecia per 5 mm length
No. of rows of zooecia
No. of zooecia per fenestrule715.

Perrnian Ectoprocta frorn the Abadeh region

ranging from 0.300 to 0.500 mm, usually
13 to 16 branches per 10 mm horizontally.
Interval between branches ranging from0.500 to 0.750 mm measured from centerto center of branches. Fenestrule elongated elliptical in outline, width ranging
from 0.160 to 0.475 mm, length ranging
from 0.625 to 0.950 mm and S to 10 fenestrules per 10 mm length of branch measured on 4 specimens. Dissepiment relatively wide but narrower than that ofbranch, ranging from 0.275 to 0.500 mm
in width. Zooecial tubes arranged usually
in 3, occasionally 4 intersecting longitudinal rows. In tangential section, zooecial tubes hexagonal or rhomboidal except
for the both sides whose tubes are pentagonal or triangular at lower to middle
levels of branch. Zooecial tubes nearsurface circular, ranging from O.OSO to
0.100 mm in diameter. IS to 20 zooecial
apertures arranged per 5 mm length of
one row, usually 4 apertures per fenestrule. Interval between zooecial apertures in longitudinal series from centerto center ranging from 0.250 to 0.350 mm.
Stereom covering the reverse side conslstmg of inner tissue with 6 to S prominent capillary canals on which capil-

laries are not developed, and outer finegrained fibrous tissue.
Meshwork formula.-13-16/S-10/ /lS-20/3(4).
Measureillents.-See Table 5.
Remarks. -The present species is nearest to Polypora magnicava MOROZOVA in
the essential characters. The differences
between these two species will be discussed in the description of P. magnicava.
Horizon and SPecimen no.-C-ll, C-1213 and C-13 beds of Unit-4 in Section C.
Reg. nos. DESC-S0023, DESC-S0024b,
DESC-S0025, DESC-S0026 and DESCS0027b.
Polypora magnicava MOROZOV 1\
Pi. 32, Fig. S.
1970. Polypora magn"icava lv!OIWZOYA, p. 210,
pI. 49, fig. 1.

A single tangential section of fragmentary zoarium was examined. Branches
usually straight, connected by dissepiment
at regular intervals ranging from 0.S75
to 1.000 mm measured from center tocenter of dissepiments, but occasionallyslightly zigzag and anastomosing.] LWidth

soyanensis MUIWZOVA (in mm).

S0025

S0026

S0027bca. 14
9.5-10
lS-20
3-4

IS

10
19-21
3-5

16-17
S-9
lS-20
3-4

4

4

4

0.375-0.450

0.375-0.425(0.300-0.500)
0.500-0.625,.000-'1. 100
0.325-0.450
0.OSO-0.100
0.250-0.2S0

O. 575-0. S75
1. 000-1. 125
0.250-0.300
O. OSO-O. 090
0.250-0.300

279

0.300-0.375
0.500-0.750
1. 125-1. 250
0.250-0.325
0.OSO-0.100
0.250-0.350----------------------- -------

280

Sumio SAKAGAMI

of branch as wide as that of fenestrule,
ranging from 0.300 to 0.400 mm, usually
12 to 16 branches per 10 mm, horizontally.
Interval between branches ranging from0.550 to O.SOO mm measured from centerto center of branches. Fenestrule irregularly elongated elliptical or rectangularwith rounded corners, width ranging from0.250 to 0.475 mm, length ranging from
O.SSO to 0.950 mm and 10 fenestrules per
10 mm length of branch. Dissepimentrather weak, ranging from 0.150 mm to0.250 mm in width. Zooecial tubes arranged usually in 3, occasionally 3 weaklyintersected longitudinal rows in a branch.
In tangential section, zooecial tubes rounded triangular in both sides and roundedrhomboidal in central row at lower levelof branch, becoming kidney shaped tocircular at middle to upper levels, 0.100
to 0.125 mm in diameter near surface.
Zooecial apertures IS to 20 per 5 mm
length of one row, usually 4 apertures
per fenestrule. Interval between zooecial
apertures in longitudinal series from
center to center 0.250 to 0.300 mm.
Stereom covering the reverse side consisting of inner thin tissue with relatively
fine striations and outer darker tissue
with fine granules of about 0.003 mm inTable 6.

diameter.
Meshwork formula.-12-16/10/ /lS-20/
3(3).
Measllrements.-See Table 6.
Remarks.-The present form is identicalwith the original specimen which was
described from the lower Kazanian of
the Russian platform except that the
numbers of branches per 10 mm horizontally is 12 to 16 in the present form
instead of 10 in the Russian specimen.
As MOIWZOVA (1970) pointed out, thepresent species resembles Polypora soyanensis MOROZOVA which occurs in association with the present species in the Russian platform and also in the present
locality. P. magnicava can be distinguished from P. soyanensis by the larger
fenestrule, narrower dissepiment, larger
zooecial aperture and in the shape ofzooecial tube in tangential section.
Horizon and SPecimen no.-C-13 bedof Unit-3 in Section C. Reg. no. DESCS0027a.
Polypora abadelzensis SAKAGAMI, n. sp.
PI. 33, Fig. 1.

Four tangential sections of zoaria were
examined. Straight branches connected

Measurements of Polypora magnicava MORozovA (in mm).

Specimen no.

80027aNo. of branches per 10 mm horizontally
No. of fenestrules per 10 mm length~o. of zooecia per 5 mm length
No. of rows of zooecia
No. of zooecia per fenestrule
Width of branchInterval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)ca. 161020----------------~---

3--4
4(5)

0.300-0.350
0.550-0.800
0.875-1.000
0.200-0.300
0.090-0.100
0.250-0.300

715.

Pe1'mian Ectoprocta from the Abadeh 1'egion

by dissepiments at regular intervals, ranging from 0.875 to 1.150 mm measured
from center to center of dissepiments.
Interval between branches from centerto center 0.750 to 1.000 mm. Bifurcationof branch infrequent. Width of branch
wider than that of fen est rule, ranging
from 0.375 to 0.625 mm, 10 to 13 branches
per 10 mm horizontally. Fenestrule elongated elliptical in outline, width ranging
from 0.250 to 0.450 mm, length ranging
from 0.500 to 0.825 mm, 9 to 12 per 10 mm
length of branch. Dissepiment strong,width ranging from 0.325 to 0.500 mm.
Zooecial tube arranged usually in 5 intersecting longitudinal rows, but 4 rows
after bifurcation and 6 rows before bifurcation in a short distance. In tangential
section, zooecial tubes rhomboidal or
elongated hexagonal at lower to middle
levels of branch, and circular near surface, separated by a nearly straight line,
ranging from 0.090 to 0.110 mm in diameter. Zooecial apertures 17 to 20 per
5 mm length of one row, almost perfectly
stabilized in position of aperture in relation to dissepiments, 4 apertures per
Table 7.
Specimen no.

281

fen est rule. Interval between zooecial
apertures in longitudinal series from
center to center ranging from 0.220 to0.320 mm. Stereom covering the reverse
side consisting of inner tissue 4 to 6 prominent capillary canals on which any
capillaries can not be observed, and outerfibrous tissue with coarse granules.
Meshwork fonnula.-10-13/9-12/ /17-20/5(4-6).
Measurements.-See Table 7.
Remarks.-The present new species is
not unlike Polypora darashamensis NIKIFOROVA which was described originallyas a variety of Polypora timorensis BASSLER from the Permian of Armenian
Dzhulfa in the general appearance, but
the present species differs from P. darashamensis in the meshwork formula.
According to MOROZOVA (1970), the meshwork formula of P. darashamensis is 811/5-6/ /14-15/4(3-6).
Horizon and Specimen no.-C-12-13 andC-13 beds of Unit-4 in Section C. Reg.
nos. DESC-80024a, DESC-80028 (holotype),
DESC-80029 and DESC-80030.

Measurements of Polypora abadehensis SAKAGAMI, n. sp. (in mm).

i
180024a

80028

80029
- ---

--

80030
-

-- -

----

1

No. of branches per 10 mm
10horizontallyI
9-11
No. of fenestrules per 10 mm length!
I
20
No. of zooecia per 5 mm length
No. of nodes per 5 mm lengthI
No. of rows of zooecia
I5(!)
No. of zooecia per fenestrule
I o. 550 (0.675)
Width of branchInterval between branches (center I O. 875-1. 000
to center)
Interval between dissepiments
0.875-1. 125(c. to c.)
0.325-0.350
Width of dissepiment
1
Diameter of zooecia near surface
0.100-0.110
Distance between zooecia (c. to c.) 0.250-0.300

12-13

12

ca. 11

10
17-19

11-12
. 18-193-5
40.450-0.550

9
18-20
5
4
0.500-0.625

18
4 (3-6)40.375-0.475

0.750-0.950

0.750-0.900

0.700-1. 000

Io. 950-1. 125

1. 050-1. 150

0.875-1. 000

I0.325-0.350
0.100-0.110
0.270-0.320

0.450-0.500
0.090-0.110
0.250-0.300

0.325-0.375
0.100-0.110
0.220-0.280

-

282

Sumio SAKAGAM I

Polypora aff. darashamensis NIKIFOROVA
PI. 32, Fig. 9.

Compare.1933. Polypora til1lorensis var. darashamensis
NIKIFoRovA, p. 22-24, 38, pl. 4, figs.
1-4.
1970. Polypora darashamensis: MOROZOVA, p.
219, 220, pI. 51, fig. 1.

Two tangential sections of fragmentary
zoaria were examined. Straight branches
connected by dissepiments at regular intervals, 1.300 to 1.400 mm measured from
center to center of dissepiments. Interval
between branches from center to center
ranging from 0.625 to 1.375 mm. Bifurcation of branch not observed. Width of.
branch almost as wide as that of fenestrule, usually ranging from 0.400 to 0.550mm, 8 to 11 branches per 10 mm horizontally. Fenestrule elongated elliptical in
outline, width ranging from 0.250 to 0.750
mm, length ranging from 0.800 to 1.125mm and about 7.5 per 10 mm length ofbranch. Dissepiments not so strong, ranging from 0.375 to 0.475 mm in width.
Zooecial tubes arranged normally in 4 to
5 intersecting longitudinal rows. In tangential section, zooecial tubes rhomboidal
Table 8.

at lower and middle levels of branch, andcircular near surface, separated by a finewavy line, ranging from 0.100 to 0.120 mm
in diameter. Zooecial apertures 15 to 16
per 5 mm length of one row, usually 5
apertures per fenestrule. Interval between zooecial apertures in longitudinal
section from center to center 0.300 to
0.350 mm. Stereom covering the reverse
side consisting of inner tissue with 6 to
7 prominent capillary canals and outerfibrous tissue with very small granules.
jvleshwork formula.-8-11/7.5/ /15-16/4-5.
Measurements.-See Table 8.
Remarks.-Although Polypora darashamensis was originally described from thePermian of Armenian Dzhulfa by NIKIFOROV A (1933), all of the illustrations
were only those of surface specimens.
Later, MOROZOV A (1970) revised thisspecies and made clear that its geologicalrange is from the Dzhulfian up to theLower Triassic.
The present form is nearest to P.
darashamensis in the general appearance,but slight differences in the meshwork
between them are recognized, especiallyin the numbers of zooecial rows: usually
5 in the present species instead of usually

Measurements of Polypora aff. darashamensis NIKIFoRovA (in mm).

Specimen no.
No. of branches per 10 mm horizontally
No. of fenestrules per 10 mm length
No. of zooecia per 5 mm length
No. of rows of zooecia
No. of zooecia per fenes.trule
Width of branchInterval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)

80031b80032

11
ca.7.515-16
3-5
50.400-0.550
0.625-1. 050
1. 300-1. 400
0.375-0.475
0.110-0.120
0.300-0.350

8ca.7.5
15-16
4(3-6)
50.500-0.550
1. 125-1. 375
1. 3750.375
0.100-0.110
0.300-0.350

715.

Permian EctOJJ1"Octa fl'om the Abadeh region--1 inP. darashamensis.
Horizon and SPecimen no.-C-15 bed of
Unit-4 in Section C. Reg. nos. DESC80031 band DESC-80032.
Polypora striata SAKAGA:YII, n. sp.
PI. 33, Fig. 2.

Two tangential sections of fragmentary
zoaria were examined. Straight branches
connected by very strong dissepiments atregular interval, ranging from 1.250 to
1.500 mm measured from center to centerof dissepimen ts. Bifurcation of branch
infrequent. Width of branch wider thanthat of fen est rule, usually ranging from0.500 to 0.750 mm, 8 to 10 branches per
10 mm horizontally. Fenestrule slightly
elongated elliptical in outline, width ranging from 0.375 to 0.625 mm, length ranging from 0.550 to 1.075 mm, about 8 per
10 mm length of branch. Dissepiment
ranging from 0.500 to 0.750 mm in width.
Zooecial tubes arranged usually in 5 intersecting longitudinal rows. In tangential section, zooecial tubes of the central
three rows rhomboidal and those of bothsides pentagonal or hexagonal at middle
level of branch, and circular near surface,
Table 9.

283

ranging from 0.110 to 0.130 mm in diameter. Zooecial apertures usually 18 per
5 mm length of one row, usually 5, occasionally 6 apertures per fen est rule. Intervals between zooecial apertures from
center to center ranging from 0.250 to0.300 mm. Stereom covering the reverse
side consisting of inner tissue with 7 to
8 strongly developed capillary canals andouter tissue with very coarse fibrous
striations and fine granules.
Meshwork formula.-8-10/8/ /18/5.
Measurements.-See Table 9.
Remarks.-The most conspicuous character of the present new species IS thepresence of the outer tissue with very
remarkable, fibrous striations and finegranules. In the meshwork formula it isnear to the precedently described species:
Polypora darashamensis NIKIFOROVA.
Horizon and SPecimen no.-C-15 bed of
Unit-4 in Section C. Reg. no. DESC80031a (holotype) and DESC-80033.
Polypora? sp. indet.
PI. 33, Figs. 3, 4.

Two tangential sections of fragmentary
zoaria were examined. Zoarium frequen-

Measurements of Polypora striata SAKAGAMI, n. sp. (in mm).

Specimen no.
No. of branches per 10 mm horizontally
No. of fenestrules per 10 mm length
No. of zooecia per 5 mm length
No. of rows of zooecia
No. of zooecia per fenestrule
Width of branchInterval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)

80031a

80033

ca.9
ca.8
18
4 (occas. 5)
5-6 (?)
0.625-0.750
0.875-1. 125
1. 250-1. 500
0.625-0.750
0.110-0. 130
0.250-0.300

8-10
818 (?)
450.500-0.625
1. 000-1. 250
1. 250-1. 500
0.450-0.500
0.110-0. 120
0.250-0.300

284

S1tmio SAKAGAMI
Table 10.

Specimen no.

Measurements of Polyporc sp. indet. (in mm).

I

S0034
-~I-- 7-S-

No of branches per 10 mm horizontally
No. of fenestrules per 10 mm length
No. of zooecia per 5 mm length
No. of rows of zooecia
No. of zooecia per fenestrule
Width of branchInterval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)tly bifurcated and expanded laterally may
be fan shaped. Nearly straight or weakly
zigzag branches anastomosing or connected by very short dissepiments atregular intervals ranging from 1.875 to2.125 mm measured from center to centerof dissepiments. Width of branch almost
as wide as that of fenestrule, ranging
from 0.700 to 0.850 mm, usually 7 to 8
branches per 10 mm horizontally. Intervals between branches ranging from 1.125
to 1.625 mm measured from center tocenter of branches. Fenestrule usuallyelongated elliptical but occasionally irre-

5-6
14-15
5
5
O. 700-0. SOO
1. 125-1. 500
1. S75-2. 125
0.550-0.700
0.120-0.130
0.300-0.400

S0035

7
5.5
15-16
5
5-6
0.750-0.S50
1. 500-1. 625
1. S75-2. 125
O. 600-0. S50
O. 1l0-0. 130
O. 250-0. 3S0

gularly shaped, width ranging from 0.675
to 0.875 mm, length ranging from 1.125
to 1.550 mm, and 5 to 6 fenestrules per
10 mm length of branch. Dissepimentusually nothing but occasionally very
short and wide, 0.550 to 0.850 mm inwidth. Zooecial tubes arranged usuallyin 5 intersecting longitudinal rows. In
tangential section, zooecial tube rhomboidal at lower to middle levels of branch
and circular at upper level. Zooecial
diameter near surface ranging from 0.110
to 0.130 mm. Zooecial apertures 14 to 16
per 5 mm length of one row, usually 5 toExplanation of Plate 31(All figures are in x 20)
Figs. 1-3. Fistulipora elegantula NIKIFoRovA
1, longitudinal section of zoarium, DESC-SOOOL 2, transverse section of zoarium, DESCS0002. 3, tangential section of zoarium, DESC-S0003.
Figs. 4-6. Fistulipora pseudomonticulosa SAKAGAMI, n. sp.
4, longitudinal section of zoarium, DESC-80004 (holotype). 5, transverse section of zoarium,
DESC-S0005. 6, tangential section of zoarium, DESC-S0006.
Figs. 7-9. Eridopora d. parasitica (WAAGEN and WENTZEL)
7, longitudinal section of zoarium, DESC-S0007. S, tangential section (inner part) of zoarium,
DESC-S0009a. 9, tangential section (outer part) of zoarium, DESC-SOOOSa.
Figs. 10-12. Hexagonella tortuosa WAAGEN and WENTZEL
10, longitudial section (half side) of zoarium, DESC-S0010. 11, 12, tangential sections ofzoaria, DESC-SOOll and DESC-S0012, respectively.

SAKAGAMI:

Permian Ectoprocta from Abadeh

Plate 31

715.

Permian Ectoprocta from the Abadeh reuion

6 apertures per fenestrule. Intervals between zooecial apertures in longitudinal
series from center to center ranging from0.250 to 0.400 mm. Interzooecial materials
consisting of dark, dense tissue with
abundant capillaries. Stereom covering
the reverse side composed of inner coarse
fibrous tissue and outer dark, dense tissue
consisting of 4 to 5 layers with abundantcapillaries. Diameter of capillaries ranging from 0.003 to 0.005 mm.
Meshwork formula.-7-S/5-6/ /14-16/5.
Measurements.-See Table 10.
Remarl,s.-The present form is notunlike Polypora? macrops YANG and Loo
which was described from the upper partof the Bayinhe Formation (the upper partof Lower Permian) in the zoarial structure, i. e. the dissepiments are very shortor almost absent and branches anastomosing each other in many cases. However, the present species can be distinguished from P. macrops in the meshwork
formula. The specific denomination must
be postponed until more materials areaccumulated.
Horizon and SPecimen no.-The Araxilevis horizon of Section J. Reg. nos.
DESC-S0034 and DESC-S0035.
Genus Septopora PROUT, lS59
Septopora lineata NIKIFOROVA
PI. 33, Fig. 5.
var. lineatap. 28, 29, 39, pI. 6, fig. 7,
text-figs. 9, 10.
1970. Septopora lineata: MOROZ OVA, p. 199,
200, pI. 44, figs. 1, 2.
1933. Septopora

subquadrans

NIKIFOROVA,

Two tangential sections made out of thesame zoarium were examined. Straightbranches connected by dissepiments atregular intervals ranging from 0.700 to
0.S50 mm measured from center to center285

of dissepiments, but occasionally amalgamated to adjacent branch and in somecases, branch detached from old one. No
branch arising directly from the centerof dissepiment observed. Width of branch
ranging from 0.375 to 0.750 mm, 10 to 14
branches per 10 mm horizontally. Intervals between branches varying from 0.500
to 2.250 mm measured from center tocenter of branches. Fenestrule sphericalto somewhat squeezed elliptical but incase of that surrounded by long dissepiments crescentic with rounded corners inoutline, width varying from 0.250 to l.250
mm, length ranging from 0.250 to 0.425
mm, and 13 to 14 fenestrules per 10 mm
length of branch. Dissepiment narrower
than that of branch, ranging from 0.375
to 0.450 mm, in which zooecial tubes
arranged in intersecting longitudinal rows,
bearing usually 5 to S, sometimes evento 10 zooecia. Zooecial apertures arranged in 2 longitudinal rows, divided bywell developed carina, IS to 20 per 5 mm
length of one row, perfectly stabilized inposition of aperture in relation to dissepiments, 3 apertures per fenestrule.
In tangential section, zooecial tube
rhomboidal with rounded corners tokidney shaped at lower to middle levels
and circular near surface. Zooecial diameter near surface 0.120 to 0.140 mm. Intervals between zooecial apertures inlongitudinal series from center to center
ranging from 0.240 to 0.300 mm. Stereom
covering the reverse side consisting of
inner thin translucent tissue and outer
thick tissue with coarse fibrous striations.
Meshwork formula. -10-14/13-14/ /lS20.
Measurements.-See Table 1l.
Remarks.-Although there are slightdifferences between the present form andthe originally described specimens of
Septopora lineata which was describedfrom Armenian Dzhulfa as a variety of286

Sumio SAKAGAMI
Table 11.

Measurements of Septopora lineata NIK IFOROV A (in mm).

Specimen no.

I,

S0036

I

10-14
ca.13
lS-20
3
0.375-0.500
0.500-1. 100
0.700-0.S50
0.375-0.450
0.120-0.140
0.260-0.300
5-6

I

S0037

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

No. of branches per 10 mm horizontally
No. of fenestrules per 10 mm length
No. of zooecia per 5 mm length
No. of zooecia per fenestrule
VVidth of branchInterval between branches (center to center)
Interval between dissepiments (c. to c.)
VVidth of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)No. of zooecia in dissepiment

Septopora subquadrans ULRICH by NIKIFOROV A (1933) in the measurements ofthe meshworks, the present form can be
identified with Septopora lineata. Especially, there is a wide range of variationsin the width of fenestrule in the samezoarium. In other microscopic characters
the present specimens agree well withthose from Armenian Dzhulfa described
by NIKIFOROV A (1933) and MOROZOVA(1970).
Horizon and SPecimen no.-C-O bed of
Unit-4 in Section C. Reg. nos. DESC-

I

I

9
ca. 1419-20

30.625-0.750
1. 000-2. 250
0.700-0.750
0.375-0.425
0.120-0.140
0.240-0.300
5-1080036 and DESC-80037.
Septopora tarazi SAKAGAMI, n. sp.
PI. 33, Figs. 7, S.

A single tangential section was examined. Zoarium consisting of nearlystraight, parallel branches connected bydissepiments at regular intervals ranging
from 0.625 to 0.675 mm measured from
center to center of dissepiments. Usuallya new branch detached infrequently from

Explanation of Plate 32(All figures are in x 20)
Figs. 1, 2. Stenopora? n. sp.
1, longitudinal section of zoarium, DESC-S0017. 2, tangential section of zoarium, DESCS0016a.
Figs. 3-5. Araxopora araxensis (NIKIFOROVA)
3, longitudinal section (half side) of zoarium, DESC-S001S. 4, tangential section (inner part)of zoarium, DESC-S0020. 5, tangential section (outer part) of zoarium, DESC-S0039.
Fig. 6. Polypora soyanensis MOROZ OVA
Tangential section of zoarium, DESC-S0023.
Fig. 7. Polypora tubulosa N IKIFOROV A
Tangential section of zoarium, DESC-S0022.
Fig. S. Polypora manicava MOROZ OVA
Tangential section of zoarium, DESC-S0027a.
Fig. 9. Polypora aft. darashamensis NIKIFOROV A
Tangential section of zoarium, DESC-S0032.

SAKAGAMI:

Permian Ectoprocta from Abadeh

Plate 32

715.

Permian Ectoprocta from the Abadeh region

old one. Width of branch ranging from
0.425 to 0.500 mm, 8 to 10 branches per
10 mm horizontally. Intervals between
branches usually ranging from 0.750 to
1.250 mm occasionally even to 1.500 mm
measured from center to center of branches. Fenestrule usually spherical orsomewhat squeezed, but in case of fenestrule surrounded by long dissepiments,
depressed arrow-head shaped in outline,
width varying from 0.325 to 1.000 mm,
length ranging from 0.250 to 0.325 mm,
and 16 to 17 fenestrules per 10 mm lengthof branch. Dissepiment narrower thanthat of branch, ranging from 0.250 to0.325 mm, in which zooecial tubes arranged in 2 intersecting longitudinal rows,
bearing usually 3 to 5 but varying 2 to 9
zooecial tubes. Zooecial apertures arranged in 2 longitudinal rows which are
divided by well developed carina, 21 to
22 per 5 mm length of one row, almost
perfectly stabilized in position of aperturein relation to dissepiment, 3 apertures per
fenestrule.
In tangential section. zooecial tube
Table 12.

~1easurements

28'/

rhomboidal or somewhat deformed quadrilateral at lower level, kidney shaped
to elliptical with irregular margin at
middle level and circular near surface.
Zooecial diameter near surface 0.100 to
0.110 mm. Intervals between zooecial
apertures in longitudinal series from
center to center ranging from 0.250 to0.300 mm. One hemiseptum visible insome zooecial tubes at middle level ofbranch. Prominent nodes arranged on
the carina, elongated elliptical in tangential section, their shorter and longer
outside diameters ranging from 0.125 to0.150 mm and 0.250 to 0.300 mm, respectively and intervals between them 0.450
to 0.600 mm measured from center tocenter. Shape of hollow in carina elongated elliptical with very strong notches.
Also in a dissepiment, 1 to 4 nodes arranged in one row, circular in tangential
section, about 0.100 to 0.120 mm in outside
diameter, with star-like hollow. Stereom
covering the reverse side rather thin,consisting of inner light-colored dense
tissue and outer dark and coarse fibrousof Septopora tarazi S,\KAGA?vII, n. sp. (in mm).

Specimen no.
No. of branches per 10 mm horizontally="10. of fenestrule per 10 mm length
No. of zooecia per 5 mm length
No. of nodes per 5 mm length
No. of zooecia per fenestrule
Width of branchInterval between branches (center to center)
Interval between dissepiments (c. to c.)
Width of dissepiment
Diameter of zooecia near surface
Distance between zooecia (c. to c.)
Outside diameter of node (longer)
Outside diameter of node (shorter)
Distance between nodes (c. to c.)No. of zooecia in dissepiment

80038
8-10
16-17
21--229-10
30.425-0.500
0.750-1. 250
0.625-0.675
0.250-0.325
O. 100-0. 110
0.250-0.300
0.250-0.300
0.100-0.130
0.450-0.600
3-9

288

Swrnio SAKACAMI

tissue with 2 to 3 rows of capillaries.
about 0.030 mm in diameter.
Meshworl, formllla.-8-10/16-17/ /21-22.
Measurements.-See Table 12.
Remarks.- The present species can be
distinguished from Septopora lineata
NIKIFOROVA which was described fromthe Armenian Dzhulfa and also from C-O
horizon of the Abadeh Permian in themeshwork formula and microscopic characters. The thin stereom covering reverse
side and the prominent nodes arranged in
one row on straight carina are conspicuous characters of the present new
species.
Horizon and SPecimen no.-D-16 bed of
Unit-3 in Section D. Reg. no. DESC-80038(holotype).

Repository.-All of the specimens treated in this paper are preserved in thecollection of the Department of Earth
Sciences, Faculty of Science, Chiba University, Chiba City, Japan.

References
B,\SSLER, R. S. (1929): The Permian Bryozoa
of Timor. Paliiont. Timor, vol. 16, no.
28, p. 37-89, pI. 225 (1) -247 (23) .

GOR]C);OYA, R. V. (1975): The Permian
Bryozoa of Pamir. Akad. Nauk SSSR.,
Trudy Paleont. Inst., vol. 148, p. 1-127,
pIs. 1-29 (in Russian).
!\1oRozOVA, I. P. in R UZIIE?\CII EV, V. E. andSARYCHEVA, T.G. (1965): Bryozoa in"The development and change of marineorganisms at the Palaeozoic-Mesozoic
boundary". Akad. Nauk SSSR., Trudy
Paleont. Inst., vol. 108, p. 57-62, 183-197,
pIs. 25-28 (in Russian).
MOROZOYA, I. P. (1970):
Late Permian
Bryozoa. Akad. Naull SSSR., Trudy
Paleont. Inst., vol. 122, p. 1-347, pIs. 164 (in Russian).
NIKIFOROVA, A. I. (1933): Upper Paleozoic
Bryozoa from the Djulfa region. Trans.
United Ceol. Prospect. Sen'. USSR., Fasc.
364, p. 1-44, pIs. 1-6 (in Russian withEnglish abstract).
SAKAGAMI, S. (1976): On the Permian Bryozoa from the northern part of Sainbeyli,
central Turkey. Trans. Proc. Palaeont.
Soc. Japan, N. S., no. 103, p. 398-405, pl.
42.
- - (1980): Upper Permian Bryozoa from
Guryul ravine spur three kilometre northof Barus. Palaeont. Indica (in press).
TARAz, H. (1971): Uppermost Permian and
Permo· Triassic transition beds in central
Iran. Amer. Assoc. Petrol. Ceol. Bull.,vol. 55, no. 8, p. 1280-1294.
- - (1974): Geology of the Surmaq-Deh Bid

Explanation of Plate 33(All figures are in x 20)

Figs. 1, 6. Polypora abadehensis SAKAGA~II, n. sp.
1, tangential section of zoarium, DESC··80028 (holotype). 6, showing the lower level of thesame zoarium.
Fig. 2. Polypora striata SAKAGAl\Il, n. sp.
Tangential section of zoarium, DESC-80031a (hoiotype).
Figs. 3, 4. Polypom? sp. indet.
Tangential sections of zoaria, DESC-80034 and DESC-80035, respectively.
Fig. 5. Septopora lineata NIKIFOROVA
Tangential section of zoarium, DESC-80037.
Figs. 7, 8. Septopora tarazi SAKAGAMI, n. sp.
7, tangential section of zoarium, DESC-80038 (hoiotype). 8, showing the arrangement of
nodes of the same zoarium.

SAKAGAMI:

Permian Ectoprocta from Abadeh

Plate 33

715.

Permian Ectoprocta from the Abadeh regionarea, Abadeh region, central Iran. Geol.
Surv. Iran Rep., no. 37, p. 1-148.
WAAGEN, W. and WENTZEL, J. (1886): Salt
Range fossils. Palaeont. Indica, Ser. 13,
vol. 1, pt. 6, p. 854-892, pIs. 105-U5.
YANG, K. C. (1958) : Stenoporidae from Upper

Acta Palaeollt.
Palaeozoic of China.
Sillica, vol. 6, no. 2, p. 122-139, pIs. 1-5.
~- and Loo, L. H. (1962): Paleozoic Bryozoa of Qilianshan. Geology of Qilianshall,
vol. 4, no. 5, p. 1-114, pIs. 1-24 (inChinese) .

-1 '7/ 'l'ir15 7 /~ T - jlliJglZG')=1£$L;7~nHn (;: vt .r1trn): :>'iH, C=:~\fl:*o)JftWr:UJtHv:.[#J;iili L.
-C, -1 '7 / 'I,m) 7 /~ T -JlliJl!l;G')=~\!Hc;: ~-.r1t1t:P G')~II'1'iJFf~(I~Ij}f'1E~D'''? t.::.o ,liJX'\JU ~nt.::.;: vt!l!.1t"£i (1 7 Iv -" :=. 7;( Iv 7 7 .Jlli1~G') Gnishik-Khachik-Lower Dzhulfian I~r/J' G ¥IH~f~nt.::.

b G') c 'd'ijiU.drHJ)~~ b i?, -s: t.::./< " - IV,=~m:$L;;: rt!:H fU'I c b J~;IJl·f,fI~~ Iv -C'I.' Q 0

Hi-:d.::.o289

r 1/ /7', :/J '/ " - Iv ,IH '/:)-, 7- -'E- - IVG')
4 Yfr~~{l;LI5f,ni-""-n:. 01. '-C'!I~l:rY.l~I'I'gi;Gjljl(
ftU: iU:~

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290

Palaeontological Society of Japan Special Papers No. 17

Revision of Matajiro Yokoyama's Type Mollusca from the
Tertiary and Quaternary of the Kanto Area
By Katsura Oyama. Reprinted May 30, 1980, 148 pp., 57 pis.
Price (postage and handling inc!.) ¥6,OOO or equivalent US dollars
This publication is a reprint of Dr. K. OYAMA'S (1973) revision of the late Professor
Matajiro YOKOYAMA'S type and illustrated specimens of Mollusca from the Tertiary
and Quaternary of the Kanto area (Tokyo and its vicinity). It is concerned not onlywith the systematics but gives also the locality records, geographic and bathymetric
distribution and ecology of each species. The paper proved to be so valuable andindispensable for Late Cenozoic biostratigraphy and molluscan paleontology that it ran
out of stock in five years. It has just been reprinted to meet readers' demand.
The Special Papers are on sale at the Society. Orders must be accompanied byremittance made payable to Dr. Tsugio SHUTO, Editor of the Special Paper, Palaeontological Society of Japan, c/o Department of Geology, Faculty of Science, Kyushu
University 33, Fukuoka (Hakata) 812, Japan.

Trans. Proc. Palaeont. Soc. japan, N.S., No. 118, pp. 291-324, pis. 34-36, june 30, 1980

716.

EARLY CRETACEOUS GASTROPODA FROM THE CHOSHI
DISTRICT, CHIBA PREFECTURE, CENTRAL J AP AN*TOMOKI KASE
Department of Geology, National Science Museum,
Hyakunin·cho, Shinjuku-ku, Tokyo 160
AND

HARUYOSHI MAEDA
Geological Institute, Faculty of Science,
the University of Tokyo, !-longo, Bunkyo-ku, Tokyo 113

Abstract: This paper deals with a gastropod fauna from the Lower Cretaceousof Choshi district, Chiba Prefecture, central japan. The following twelve species
belonging to eleven genera are described: Calliostoma? ojii KASE, sp. nov., Ataphrus (s. str.) nipponieus KASE, sp. nov., Hayamia rex KASE, gen. et sp. nov.,
Haya/llia ehoshiensis KASE, gen. et sp. nov., Oolitica sp., Amberleya (Eueyclus)
japollica IC\sE, sp. nov., Metriolllphalus nagasakiensis KASE, sp. nov., Perissoptera
eiegalls I\.,\SE, sp. nov., Pietteia eretaeea KASE, sp. nov., Ceratosiphon densestriatus
KASI·:, sp. nov., Vanikoropsis deeussata (DESIIA YES) and Eriptyeha japollica KASE,
sp. nov.
The genus Hayamia is proposed as new. The operculum of Haycm.ia rex,
preserved nearly ill situ in the aperture, is characterized by the elliptical outline
without a quadrangular process at the adaxial edge. The opercular feature of
Hayamia rex is apparently identical with that of Natieopsis. This fact suggeststhat Hayamia has a close relationship to Naticopsis and evolved along a phylogenetic
line independent from Neritopsis. Pietteia eretaeea is the first record of occurrenceof this genus from Cretaceous beds. Discrimination between the genus Ceratosiphon
and Tessarolax is also presented.
It is noteworthy that the gastropod fauna of the Choshi Group includes several
species closely related to or, in one case, identical with the species from the Barremian, Aptian or Albian of France and England.

of Choshi district, Chiba Prefecture, central Japan. Molluscan fossils have been
considered to be rare and the fossils obtained are, in general, in the form ofmoulds and casts, so that paleontologicalstudies on gastropods could not previously
be undertaken on a firm basis. In 1977,
however, five students of the Geology-

Introduction
Aside from a faunal list and illustrations provided by SHIKAMA and SUZUKI(1972), little has been known on the gastropod fauna from the Lower Cretaceous* Received February 5, 1980, read October
20, 1979, in Nagoya.

291

Tomoki KASE and Haruyoshi MAEDA

292

rately by I. HAYAMI and T. 0]1.

Astronomy Club, the University of Tokyo
including the junior author found a large
number of exceptionally well preserved
invertebrate fossils at some localities.
They entrusted identification of gastropods to the senior author. Supplementarycollections were made by I. HAY AMI of
the University Museum, the Universityof Tokyo and the senior author in 1977
and 1978. As a part of continuing studies
on the Cretaceous gastropod faunas inJapan, the senior author herein describes
twel ve species of gastropods including
two species which belong to a new genusof the Neritopsinae.
The collections
under discussion also include a numberof bivalves which will be described sepa-

Geological Setting

The Lower Cretaceous strata of the
Choshi disrict, named the Choshi Groupby SHIKAMA and SUZUKI (1972), crop out
narrowly along the eastern coast of the
Choshi Peninsula. Previous interpretations of the stratigraphic sequence in thisarea vary greatly among different workers. This is probably because the exposures of Cretaceous rocks are limited
in size and number, and because the contained fossils have not been sufficiently
studied to serve as horizon markers andno key beds are found. Recently, OBATA
,,

,

...,

36'

--',

" - .......

. Inub6 -zakia2km

Text-fig. 1.

Map showing the fossil localities in the Choshi district, Chiba
Prefecture, central Japan.

716.

Early Cretaceous Gast1'opoda from Choshi

Loc.!

l \;a

c!J

298

0

lsejigaura

,;>
L>

0
0

Text-fig. 2.

100m

0

Loc.3

100m

~

:\1ap showing the fossil localities in the coast of Isejigaura, northern coastof Kimigahama and coast of Nagasaki harbour, Choshi City.

et al. (1975) described the stratigraphy of
the Choshi Group, subdividing, from baseto top, into the five formations, as follows:
1) Isejigaura Formation, the lowest unit ofthis group, composed of gray to blueish graymudstone and alternating mudstone and grayfine- to medium-grained sandstone or siltstone.
Barremian based on Barremites (B.) aff.
difficilis (D'ORI3IG);Y) and Crioceratites d.
emerici (LEVEILLE).
2) Ashikajima Formation, overlying Isejigaura Formation with unconformity. Baseconsisting of pebble- to cobble-sized conglomerate, following in upward sequence bybrownish gray cross-laminated medium- tocoarse-grained sandstone, alternating mediumto coarse-grained sandstone. Lower Aptian suggested by TropaeulI! aff. bowerbanki (SOWERBY), Australiceras aff. gigas (SOIn:R13Y) andDufrenoyia aff. dufrenoyi (D'ORIlIG);Y).
3) Kimigahama Formation consisting ofalternation of blueish gray mudstone and lineto medium-grained sandstone. Trigoniid andother bivalves, plant fragments and ambersometimes occurring in calcareous concretions,and trace fossils very common. Lower Aptian
based on Cheloniceras (s. str.) d. proteus
CASEY.

4) Nagasakihana Formation consisting ofcross-laminated, medium-grained sandstone.
Very poor in fossils except for some trace

fossils and plant fragments. Lower Aptian
based on Cheloniceras (s. str.) lI1eyendor,iii(D'ORIllGNY) from lower horizon.
5) Toriakeura Formation consisting ofalternation of light gray mudstone and fineto medium-grained sandstone, fault contact
with underlying Nagasakihana and Kimigahama Formrtions, overlain by Middle Miocene:\letogahana and Pliocene Naarai Formations.
Mudstone with echinoderms, bivalves, ammonites, plant fragments and trace fossils.
Lower Upper Aptian suggested by Cheloniceras (Epicheloniceras) sp.

Locality GUIde
Loc. 1 ~ = Loc. 7306 in OBAT,\ et al., 1975"
Location: coast of Isejigaura, Ashikajima-cho,
Choshi City [35°43'13"N, 140043'19"EJ.
Stratigraphic position: near the base of the
Isejigaura Formation by OIlAT/\ et al.
(1975) .
Age: Barremian.
Lithology and associate fauna: gray sandy
siltstone with plant fragments and lenticular, highly calcareous concretions. Gastropods occur in the concretions together
with bivalves and ammonites. The concretions are parallel to bedding. They
are composed of micrite and contain asmall amount of quartz grains.
The
delicate surface sculpture, such as crowded growth lines of Amberleya (Eucyclus)

294

Tomoh ]{ASE and HCLnLyoshi MAEDA

japo n ica, is we ll preserved, but the s he ll s
are r ecrysta lli zed to coa rse -g ra in ed ca lc it e_
S pecies: AmberLeya (Eueyclus) japonica K.\ 51·: ,sp. nov. and Doliliea sp.
Co ll ectors : I-l. lvIAI': DA and T. 0./1.

Loc. 2 t: = Loc. 09 in SII IKAMA a nd SLZ L' KI,
1972; Loc. 73 16 in OI3 ATA e l aL., 1975J
Location: northern coast of Kimi gahama,As hik aj ima-cho, Chos h i Cit y [ 35°42' 541/N,
140 052 ' 241/E ~ .
Strat igraphi c posit ion : about 50 m above thebase of t he I<im igahama Formation by
OS ATA e l al . ( 1975) .
Age: Barrem ian. One of the ammon ites col lected at this loca li t y is assigned to thegenus B ;; rremil es , suggestin g a Barremianage (personal comm unicat ion b y Dr. I.
OSATA ) . Pre vio us to this report no foss ilavailable for age determ in ation was reported from the I<imigaham a Formation
exposed on the northern coast of Kimigahama. Further study must be done tosolve the strat igrap hi c relation between
the format ions at Nagasa ki a nd 1< imiga ham a.
Litho logy and associate fauna: a lternation oflaminated quartzose fine·gra in ed sandstone
and da rk gray si ltstone (Text·figs. 3, 4) .
The sandstone contains carbonitized plant
fra gments, coa l, an d is poor in fossils;
the si ltstone yie ld s deform ed mo lluscs

s Li ch as NucuLa sp ., echinoderms, andca rbon iti zed plant fragments. The gastropods described in this paper occurred ina s il ty laye r of the basal pa rt of a sandstone about 40 cm in maximum thickness,
w hi c h abruptly th ins out lateral ly . Microscopicall y, fossil bearing sed imen t is composed of 50 to 60 percent micrite, 40 to
50 percent angu lar quartz gra ins and asmall amo unt of ang ul ar pla g ioclase grains
and rounded c hert grains. T he foss ils areaggrega ted in th is la yer and bi va lves are
the most dominant const it uent, i. e. Nano nevis sp., NucuLa sp., vener id , etc . Gas tropods a re common, and ammon ites andasteroid ec hinoderms occur rarel y. Delicate surface scu lp t ure of these fossi ls andthin, lo ng sp ines of aporrha id gastropods
are well preserved, although the shells
are recr ysta lli zed to coarse -g ra ined calcite .
Man y indi v iduals of NUClda sp. and Nano -bedded. gray, sandy
sills lone.

fine-grained sandstone:molluscs abundanl in silty
layer a l base .
gray sandy sillslone: plan I
fragm ents and bivalves rare .
laminated fine - grainedsandston e .

finel y lamina led graysandy sills lone; burrowing rare .

fine-grained sandstone ;large-scale cr055 -Iaminalion
al m iddle: coal and planl
fragments common .

finely laminaled gra ysandy sills lone .

-. . .e -*Text ·fig.3 . Photograph s hows the foss ilbearing sandstone at Loc. 2, northern coastof 1< imi gahama .

Text-fi g. 4. Co lum nar section at Loc. 2,
northern coast of Kimigahama.

716.

Early Cretace011S Gastrol)oda from Choshinavis sp. in this layer are articulated,suggesting that these fossils are nearly
autochthonous in origin.
Species: Call iostoma? ajii KASE, sp. nov.,!ltaphrus (s. str.) nippanieus KASE, sp.
nov., Hayamia rzx KASE, sp. nov.,
Hayamia choshimsis KASE, sp. nov.,
Perissoplera elegans KASE, sp. nov., Pie/·leia eretaeea KASE, sp. nov., Ceratosiphon
densestriatus KASE, sp. nov., Vanikoropsis
deeussata (DESIlA YES) and Eriptyehajaponica K:\5E, sp. nov.
Collectors: 1. HAYAMI, T. KASE, Y. KASE, H.
\IAEDA, S. MORIYA, T. 0.11, T. TSL((IIand S. U\II:\O.
Loc. a [=Loc. 13 in SIIII,'\\IA and SCZl:"I,
1972; Loc. 7028 in OIlATA el a/., 1975J
Location: coast of Nagasaki harbour, Nagasaki·cho, Choshi City [35°41'30"N, 140 51'54"E_.
Stratigraphic position: about 50 m above thebase of the Kimigahama Formation by
OBATA et al. (1975).
Age: Lower Aptian.
Lithology and associate fauna: Gray calcareous
sandy siltstone.
Many molluscs werereported in this bed (see Table 5 in Sill·"/\\IA and SUZL"I, 1972).
Species: Me/riomphalus nagasal,iensis K,YSE,
sp. nov.
Collector: S. SAKl:\L\.
0

General Remarks of the Gastropod
Fauna of the Choshi District

The Barremian to Lower Aptian gastropod fauna of the Choshi district consists of ten new, one already known and
one indeterminable species, as follows:
Callios/ol1w? ajii KASI':, sp. nov.
A/aPhrus (Atcphrus) nipponieus KASE, sp.
nov.
Hayamia rex KASE, gen. et sp. nov.
Ifayamia ehashiensis K,\SE, gen. et sp. nov.
Amberleya (Eueyclus) japonica KASIC, sp.
nov.
Go! itica sp.
Me/riamphalus nagasakimsis KASI':, sp. no\'.
Perissaptera elegans KASE, sp. nov.

~95

Pietleia ere/aeea KASE, sp. nov.
Ceratasiphon denses/ria/us KASE, sp. nov.
Vanikorapsis deeussata (DESIlA YES)
Eriptyeha japanica KASE, sp. nov.

Of these species AmiJerleya (Eucyclus)
japonica and Oolitica sp. are from theBarremian Isejigaura Formation at Lac. 1,
and all the other species are from theKimigahama Formation. The fossils fromthe Kimigahama Formation were mostly
obtained at Lac. 2, but the material of
Metriomplwlus nagasal?iensis was only
collected at Lac. 3. Ten gastropod species
were identified by SIIIKAMA and SUZUKI(1972). However. the material from Loc.
3 is so fragmental that specific and generic
identification is difficult, except for
Metriomphalus nagasa!?iensis.
In Japan the Lower Cretaceous gastropods have not been sufficiently studied.
Marine deposits of the Aridan (=Aritan,
approximately Upper Neocomian) age
bear some gastropods from the Outer
Zone of the Southweat Japan, but mostof them remain undescribed. The upper
part of the Ishido Formation (lower Upper
Barremian according to OBATA et al.,1976) of the Sanchu area in central Japanconsists of shallow marine sediments andcontains several gastropod species thatwere described by Y ABE, NAGAO andSHIMIZU (1926) and revised taxonomicallyby HAY AMI and KASE (1977). Only one
species, Aporrhaidae gen. et sp. indet.
illustrated by MATSUKAWA (1977) fromthe Ishido Formation is related to Ceratosip/wn densestriatus sp. nov., but they
are distinguishable at the specific level.
The Miyako Group (Upper Aptian to
Lower Albian) of Northeast Japan ismainly composed of neritic reef sediments
and contains numerous gastropods in calcareous sandstones. These were described and illustrated by NAGAO (1934) and
HANAI, OBATA and HAYAMI (1968) and296

Tomoh K ASE c£ncl Han~yoshi NIAEDA

wer e taxonomicall y re vised by HAY AlVlI
and K ASE (1977), but still there are alarge numbe r of undescribed species.
The t wo groups ha ve no species in common, but onl y on e genus AtaphTUs (A tajJhrus) occurs in both g roups. T his genus,
however, is so lon g ranging and widelydistributed that the occurren ce of Atapln'us (AtajJhrus) is not indicative of an y
direct r elationship w ith t he Miyako fauna .
Furth er stud y is necessar y to solve th equestion of w hether t he difference is du e
to a difference in age . between the twogroups or because different depositional
environm ents are represented.
In contrast, the fauna of th e Choshi
Group contains t he followin g characteristic gastropods which are identical withor closely related to the species from theLower Cretaceous in western Europe:
The specimens of Vani lwropsis decussa ta (DESHAYES) are indist in g uishable
from those from the Gault of Cosne inFrance.
Hayamia rex sp. no v. is closel y related
to Naitopsis spiralicrenata COSSiVlANN
from the Barremian of France.
Mel riol'nphalus nagasakiensis sp. no v.
and Ceratosiphon dense s fi' iatus sp. no v.
seem to be related to T urbo coquandi
PICTET and CAMPICHE and A ponhais
ebrayi de LORIOL from the Gault of St.
Croix and Cosne in F rance, respectively.
Perissoptera elegans sp. no v. and Cerato siphon densestriatus sp. nov. may be also
intimately related to Anchura (PerissojJtera) 71wTginata (SOWERBY) and R ostellaria
retusa SOWERBY from the Albian of England, respectively.

A

B

Repository'Specim ens with the prefi x NS M are. Te x t-fi g. 5. Calliostolna? ojii K ASE,
sp. no v. (x 7.4) .

C

716.

Early Cretaceous Gastl'opoda j1'om CIIOS/I1~

preserved in the Department of Geology.
National Science Museum, Tokyo. Specimens with the prefix GIYU are preservedin the Geological Institute, Faculty of
Education. Yokohama National University.
Systematic Description(By Tomoki KASE)

Morphologic terminology in the present
paper conforms to that adopted in the"Treatise on Invertebrate Paleontology"
by KNIGHT et at. (1960, p. I129-I135).
Synonymy and type-species of the genus
and other information appearing in theTreatise are omitted unless otherwise
mentioned.
Class Gastropoda
Order Archaeogastropoda
Superfamily Trochacea
Family Trochidae
Subfamily Calliostomatinae
Genus Calliastoma Sw AINSON, 1840
Calliostoma? ajii KAsE, sp. nov.
PI. 34, Figs. 5-7; Text-fig. 5.

Material.-Holotype: NSM PM 15000,
the largest specimen (Pi. 34, Figs. 7a, b).
Para types: NSM PM 15001-15004.
Diagnosis.-Shell very small, trochiformwith carinated basal periphery. Heightalmost equal to width. Incremental angle
on penultimate and body whorls about
60°. Base weakly convex with a narrowand shallow medial depression. Depression smooth and covered by thin callus.
Surface sculptured by spiral cords andcolla bra I flat-topped ribs. Ribs strongest
near sutures. Columella short, thick and297

lobed anteriorly.
Description.- Very small trochiform andanomphalous shell with height almost
equal to width. Spire obtusely pointed,
occupying a little less than two-thirds oftotal shell height. Protoconch consistingof about three depressed conical whorls,
separated from each other by linear
suture, and with almost straight or very
weakly inflated whorl sides in early
growth stage, but with very weakly
concave sides in later stage. Penultimate
whorl about three times wider than high.
Body whorl wider than high with more distinctly concave side than those of spire
whorls and sharply separated from baseby a carinated periphery. Base broad
and weakly convex in overall, but umbilical part around columella shallowly
grooved and covered by a thin callus.
Aperture subquadrangular in outline,
prosoclinely inclined to shell axis andabout 1.4 times as wide as high. Outer
lip short, thin, sharp at its edge, nearlystraight, and abruptly turned to weakly
concave and wide basal lip with a sharp
angulation. Columella short, thick, concave and more or less inclined to shellaxis, coated 'by thick callus and lobed
anteriorly. A narrow fissure sometimes
present behind columellar lip. Parietal
region coated by thin callus. Surface
sculptured with fine spiral cords and flattopped collabral ribs that cover whorlsides. Spiral cords distinct and somewhat
crowded near suture. Collabral ribs
somewhat imbricated anteriorly, distinct
near suture and incised at basal periphery. Base sculptured by about 15 spiralcords that gradually become coarser
towards umbilicus.
Remarks.-Five specimens are available
for study. The holotype (Pi. 34, Figs. 7a,b) is the largest specimen, but the delicate surface sculpture of the whorl sides
and base was damaged during prepara-

Tomoki KASE and Haruyoshi MAEDA

298
Table l.

i'-Ieasurements in mm. [Cclliasto1l1c?
ajii KASE, sp. nov.]

Specimen
NSM PM 15000
Holotype
NSM PM 15001
Paratype
NSM PM 15002
Paratype
NSM PM 15003
Paratype
NSM PM 15004
Paratype

HeightWidth

Height ofaperture

7.2

7.4

2.8

7.6

6.9

2.7

6.8

6.5

2.5

5.4

6.1

2.0

5.5

5.9

2.0

tion. The surface sculpture is better
observed in one of the para types (PI. 34.
Figs. 6a-c).
The generic assignment of the present
species is doubtful. The thick, inclined
and anteriorly lobed columellar lip agrees
well with that of the genus Calliostoma.
This genus is very common in the Cenozoic and Recent. The genus ranges fromthe Lower Cretaceous to the present according to WENZ (1938) and KNIGHT et al.
(1960). COSSMANN (1918) listed several
Eocene species as the oldest forms, but
he suggested that the certain record of
the appearance this genus may range backto the base of the Jurassic or Triassic.
The umbilical character of the present
species, however, differs from that ofCalliostoma; that is, the umbilical part ofthe present species is weakly depressed
medially and smooth owing to the deposition of thin callus. The new species isprobably separable from Calliostoma at
the genus level, but a new genus name is
not proposed here, because thoroughrevision of the Cretaceous trochiform
gastropods is needed to define the delimitation of the genus Calliostoma.
The present species may also be related
to the geuns Proconulus COSSMANN, 1918.
However, Proconulus is typically characterized by having larger shell size, coel-conoid shell profile, convex base, andsimple and weakly grooved columellar lip,and, therefore, is considerably different
from the new species.
Africoconulus Cox, 1965 (type-species:
Proconulus spinatus DuBAH, 1948 fromthe Domerian of Morocco) has similarbasal characters to the present species.
The new species, however, cannot bereferred to that genus owing to the almost
straight whorl sides and simple ornamentation.
Calliostoma? ojii is probably congenericwith Trochus rozeti d'ARCHIAC, 1847 fromthe Cenomanian Tourtia beds of Belgium,
because the shell outline and basal characters are quite similar. However, it iseasily distinguishable from that species
by the finer and more delicate ornamentation on the whorl sides.
Occurrence.-Known only from the type
locality [Loc. 2].
Family Ataphridae
Genus Ataphrus GABB, 1869
Subgenus Ataphrus s. str.
Ataphrus (Ataphrus) nipponicus

KAsE, sp. nov.
PI. 34, Figs. 3, 8-11.

Material.-Holotype: NSM PM 15007, alarge specimen (PI. 34, Figs. 11a, b). Paratypes: NSM PM 15005, 15006, 1500815029).
Diagnosis.-A moderate-sized species ofAtaphrus (Ataphrus) with a shell about
1.2 times wider than high in adult, broadand elliptical aperture and very finenumerous spiral threads on shell surface.
Description.-Moderate-sized, thin andanomphalous trochiform shell with widthabout 1.2 times height. Spire obtusely
pointed and occupying about three-fifths

716.

Early Cretaceous Gastropoda from Choshi

of total shell height, having weakly inflated and ad pressed whorls. Teleoconchof about four, rapidly expanded whorls
separated by very weakly impressed
sutures. Protoconch smooth, depressed,
composed of about one volution. andseparated from teleoconch by a prosocline
groove(?). Body whorl about 1.4 times
as wide as high, weakly inflated aboveand broadly rounded at periphery withslightly convex base. Aperture broad,about 1.5 times as wide as high, strongly
prosoclinely inclined to shell axis andelliptical in outline with almost continuous
peristome. Outer lip long, arcuate andbeveled with sharpened edge, and continues to a slightly concave basal lip withbasal angulation. Parietal region smoothand concave, and coated by thick callus.
Columellar lip long, slightly concave andstrongly inclined to shell axis. Outerface of columellar lip broad and weakly
depressed medially with an obtuse denticle facing towards aperture. Sculpture
consisting of numerous and microscopic
spiral threads over shell surface. Spiralthreads coarser on basal periphery thanon other part of shell surface. Growthlines prosocline and somewhat rugose.
Remarks.-Twenty-five specimens areavailable for study. These specimensshow complete shell outline, very finesurface ornamentation and delicate characters of apertural area.
Although the number of specimens in
the present collection is not large enoughto evaluate the variability of shell form,
the ratios of W max./H and HA/H are
rather constant among these specimens.
Furthermore, the convexity of whorls,
the apical angle and the shape of theaperture is rather uniform compared withother species of this group. Although
the shell form is rather constant among
the specimens of nearly the same size,
smaller specimens seem to have larger

299

ratio of W max./H than larger ones.
Individual relative growth for the total
shell height (H) against total shell width(W) in five specimens shows slightly
positive allometry (Text-fig. 6).
Thecharacteristics of the protoconch are wellshown in one of the para types (PI. 34,
Figs. lOa, b). It consists of about one
whorl, but the boundary between the
protoconch and teleoconch is not clearlyobservable; probably they are separated
by a shallow groove.
This new species is similar to Ataphrlls(Ataphrus) yokoyamai NAGAO, 1934 fromthe uppermost Aptian Hiraiga Formationof the Miyako Group of Northeast Japan.
As noted by HAY AMI and KASE (1977, p.
36), Ataphrus kitakamiensis NAGAO, 1934
from the Miyako Group is regarded as asynonym of A. (A.) yokoyamai. According
to NAGAO'S description Ataphnls (AtapInus) yokoyamai is apparently characterized by smooth shell surface, but somespecimens collected near the type locality
possess distinct spiral threads over theshell surface. Some specimens withsmooth shell surface were also obtained
from the same locality, but they weremore or less abraded. The new species
differs from Ataphrus (Ataphrus) yokoyamai by the smaller shell size, moreconical shell outline, thinner test, broader
aperture and much finer spiral threads.
Most of the Jurassic and Lower Cretaceous species belonging to Ataphrus (Ataphrus) possess an apparently smooth shellsurface, and may be distinguishable from
Ataphrus (Ataphrus) nipponicus. However,
it is not necessarily known whether thesmooth shell surface of these species isoriginal or it is only due to unfavorable
preservation as exemplified by AtaphrZls(Ataphrus) yokoyamai.
The new species is quite similar to
Ataphrus adriaticus PARONA, 1909 andalso to Ataphrus pillai PARONA, 1909 from

Tomoki KASE and Ha1'uyoshi MAEDA

300

c

A
...o

H

H=0.6114W1.1559
r=0.999B

E
EM

H

NSM PM 15006~~.4~~~----T2~~4~5~~ltO--~2~0-- W
in mm.

B

H

...o

...

o

H=0.6B19W1.0931
r=0.9962Ho

H=0.7010W1.0921
r= 0.9972

•..r

E

EM

NSM PM 15023

NSN PM 15009
a~~~+---~-4-4~++~~--~

0.4

.2

In

3 4 5

mm.

10

20

w

M

~i+.4~~~----+-~3-4~5~~1~0--~2~.0-W

in

mm.

Text-fig. 6. A: Linear measurements of Ataphrus (s. str.) nipponicus KASE, sp. nov.
adopted in the present study. (H: total shell height parallel to the axis of shell coiling;
W: total shell width perpendicular to the axis of shell coiling). 8-F: Double logarithmic
diagrams for five samples of Ataphrus (s. str.) nipponicus KASE, sp. nov.

the "Albian" of Italy, especially to theirsmall specimens. The new species, however, distinguishable from the two Italian
species by its much smaller shell size,
more strongly convex base, broader aperture and the presence of fine spiralthreads on the shell surface.
Occurrence.--Known only from the type
locality [Loc. 2J.

Superfamily Neritacea

Early C1"etaceo1ts:Castropoda j1'om Choshi

716.

E

H
H=O.6585W1.l248
r=O.9950

NSM PM 15024
0.4

.2

In

3

4

10

mm.

W

20

F

H

H ::;O.6337W

1.1622/r=O.9983

NSM PM 15025
d+-~~H----+--~~~H4+------W
0.4

.2

In

3

4 5

10

mm.

20

Family Neritopsidae
Subfamily Neritopsinae
Genus Hayamia KASE, gen. nov.
[= Turbinopsis PCELINCEV, 1963;

non CONRAD, 1860J
Type-species.-Hayamia rex KASE, sp.
nov. from the Barremian of Choshi district, central Japan.

301

Diagnosis.- Large-sized and relatively
thin-shelled neritopsid with height almost
equal to width and more or less protruded
spire. Surface sculptured by numerous
and more or less irregularly spaced spiralthreads, in some species, also by costellae.
Inner lip long, narrow and concave
without a notch behind.
No parietal
tubercle. Operculum of naticopsid type,elliptical, lacking a quadrangular process
at adaxial edge.
Remarks.-Neritopsis is one of the mostcommon gastropods in the Jurassic andCretaceous. WENZ (1938, p. 412) wasaware of the difference of inner lip structure between living and Mesozoic species,
stating "-in der Mitte einer Einbuchtung,die den Vorsprung des Deckels aufnimmt
und bei den altenen Formen wenig, bei
den jlingeren starker ausgepragt ist".
Fortunately, the opercula of Hayamia rex
are preserved in the apertures of three
specimens at hand (PI. 35, Fig. 10). Theirorientation in the aperture are nearly thesame among the three specimens; thatis, the concave surfaces and the thickmargins face the ventral and abaxial
directions, respectively. This fact suggests that the opercula were entombed
nearly in situ in the apertures.
Neritopsid opercula have been describedby many authors from Mesozoic andCenozoic rocks. Some earlier authors
considered them variably as brachiopods,
cephalopods, echinoderms, and so on(BROSAMLEN, 1909). Several specimens
were described under Peltarion DESLONGCHAMPS and DESLONGCHAMPS, 1858, Cyclidia ROLLE, 1862, Scaphanidia ROLLE,
1862 and Rhynchidia LAUBE, 1870.
Three types of opercula were recognized on the basis of their morphologicalcharacters (Text-fig. 7). The first type,
exemplified by Recent Neritopsis radula(LINNE), is characterized by a very solid,
thick and trapeziform outline with aTomoki KASE and Har1tyoshi MAEDA

302
Table 2.

l'vleasurements in mm. [Ataphrus (Ataphrus) nipponicus KASE, sp. nov.J- - - - - - _ ... _.-NSM PM 15005 Paratype
NSM PM 15006 Paratype
NSM PM 15007 Holotype
NSM PM 15008 Paratype
NSM PM 15010 Paratype
NSM PM 15011 Paratype
NSM PM 15012 Paratype
NSM PM 15013 Paratype
NSM PM 15014 Paratype
NSM PM 15015 Paratype
NSM PM 15016 Paratype
NSM PM 15017 Paratype

HeightMaximum
width

Minimum
width

Height ofaperture

Apical
angle

9.6
9.6
9.5
9.2
7.4
7.3
6.8
6.4
5.6
4.4
4.3
3.9

11. 7
11. 3
11. 3
11. 4
8.8
8. 7
8.3
8.2
7.3
5.2
5.4
5.1

10.0
10.2
9.1
9.2
7.7
7.6
7.0
6.7
6.0
4.2
4.4
4.2

5.7
5.9
5.6
ca.5.7
4.5
4.4
4.0
3.7
3. 7
3.2
3.2
2.9

122
110
102
118
106
114
108
111
124
109
109
120

prominent process at adaxial side andwith a well arched abaxial margin. The
outer surface is even, smooth and coveredby callus, and has a prominent quadrangular process. The inner surface is, on
the contrary, convex, also smooth andcovered by callus, and the quadrangularprocess is radially striated. Cyclidia described by ROLLE (1862) may also be included in this type. Scaphanidia described by ROLLE (1862) and Rhynchidia
described by LAUBE (1870) may questionably be referred to this type. The second
type is exemplified by Neritopsis jurensis(MUNSTER) in BROSAMLEN (1909) from theKimmeridgian of Germany. Peltarion
DESLONGCHAMPS and DESLONGCHAMPS,
1858 seems to have been originally namedfor opercula of this type. The operculaof this type are characterized by the
angularly elliptical outline and has a quadrangular process at the adaxial edge.
The outer surface possesses fine concentric lamellae and some symmetrical andradial ridges. The wide abaxial part ofthe inner lip surface is covered by thickcallus, its adaxial margin has chevronshaped boundary, and the narrow adaxial

0

0

0

0

0

0

0

0

0

0

0

0

part of the inner surface possesses finelamellae which are nearly parallel to the
adaxial margin. It is considered that
opercula of this type are common in theUpper Triassic to Lower Cretaceous species which are of relatively small shellsize and bear cancellated ornamentation.
The first and second types are similar toeach other in having a prominent process
and in the structure of the outer andinner surfaces. The third type is represented by Hayamia rex and probably manylarge-sized and spirally striated species.
The opercula of this type are quite different from those of the above-mentioned
two types. The most distinct differenceis the lack of the process at the adaxialedge, and, therefore, shows an ellipticaloutline. It is thin in the adaxial part but
thick in the abaxial part. The outersurface of the operculum is coated bythick callus, concave, undulated at the
middle and ornamented by both radialand concentric elements. The two parts
are bordered by a chevron-shaped boundary. The convex inner surface is, on
the contrary, smooth and covered by callus
in the abaxial part, and is ornamented by716.

Early Cretaceous Gastropoda frorn Choshi

concentric lamellae in the adaxial part.
An operculum of this type was also reported by HUDLESTON (1894) from the
Bajocian of England, and rests in situ in
the aperture of Neritopsis bajocensis
D'ORBIGNY. The operculum of Neritopsis
deslongchampsii BEAUDOUIN, 1868 fromthe Oxfordian of France may be referable
to this type. Peltarion moreausi DESLONGCHAMPS, 1863 from the Oxfordian ofFrance is regarded as an operculum of
this type. The inner lip of Hayamia rex
does not possess a prominent notch, into
which, as seen in Neritopsis radula, opercular process fits. This fact is in harmony
with the lack of such a process in the
operculum of Hayamia rex. Hayamia 1'ex
is also similar to Neritopsis bajocensis in
its larger shell size, comparatively thinshell, narrow labial area and numerous
and irregualrly spaced spiral cords andcostellae. It is interesting that the opercular feature of Hayamia rex is apparently similar to that of Naticopsis. This
fact suggests that Hayamia has a close
relationship to Naticopsis and evolved ina phylogeneric line independent of Neritopsis.
Hungariella KUTASSY, 1933 is close inhaving a similar shell outline but differsby possessing a more acute spire, tubercles on the parietal and columellar lips
and a different type of surface sculpture.
Neritoptyx OPPENHEIM, 1892 differs bypossessing a prominent parietal tubercle.
PCELINCEV (1963) proposed the genus
Turbinopsis (type-species: Neritopsis multicostulata PCELINCEV, 1931). Turbinopsis
is probably considered to be asy nonym ofHayamia, but the genus name Turbinopsis is preoccupied by CONRAD (1863) for
an unrelated cerithiacean gastropod.
The following species formerly described from the Jurassic and Cretaceous beds
can be included in Hayamia, judging fromthe shell outline and surface sculpture:303

Neritopsis mutabilis HAY AMI, 1960 fromthe Lower Lias of west Japan.
Neritopsis philea D'ORBIGNY, 1847
(d'ORBIGNY, 1852) from the Lias ofFrance.
Neritopsis bajocensis D'ORBIGNY, 1852
from the Bajocian of France.
Neritopsis tricostata D'ORBIGNY, 1852
from the Bajocian of France.
Neritopsis baugierana D'ORBIGNY, 1852
from the Bathonian of France.
Neritopsis sulcosa D'ARCHIAC, 1843
(MORRIS and LYCETT, 1851) from the
Bathonian of England.
?Neritopsis striata MORRIS and LYCETT,
1851 from the Bathonian of England.
Neritopsis inaequalicosta D'ORBIGNY,
1847 (D'ORBIGNY, 1852) from the Callovian
of France.
?Neritopsis deslongchampsii BEAUDOUI:\,
1868 from the Oxfordian of France.
Peltarion moreausi DESLONGCHAMPS.
1863 from the Oxfordian of France.
Neritopsis spiralicrenata COSSMAN:\',1907 from the Barremian of France.
Neritopsis multicostulata PCELINCEV,
1931 (DRUSCHlTS and KUDRYAVSTEV,
1960; PCELINCEV, 1963) from the Barremian of Crimea.
The generic position of these species,
however, will be ensured when the opercular and inner lip features are clarified.
Hayamia rex KASE, sp. nov.
PI. 35, Figs. 5-10; Text·fig. 7, Figs. 2a, b.

Mate1'ial.-Holotype: NSM PM 15031, alarge specimen (Pl. 35, Figs. 9a-c). Paratypes: NSM PM 15030, 15032-15039.
Diagnosis.-A large-sized and prominently shouldered species of Hayamia
characterized by ovately naticiform shell
outline, broad ramp and considerably
protruded spire with width almost equal
to height. Surface sculpture consistingT O'mok'i J(ASE cLnd HaTuyoshi MAEDA

304

Text -fig_ 7. Co mparison and form of opercu la in three ner itops id
genera and subge ne ra . la, b : Neritopsis (s. str.) radula ( LI -'~E) , aft er
WE"Z (1938) . (x 1. 0) ; 2a, b: H ayamia 1'ex KASE , sp. no v . ( x 2.4) ; 3a,b: "Pellarion" bilobaLu1I1 D ESLOi'lGC II A ~ I PS a nd D " S L(J ~GC ll l\~'IP S, after
KNIGIIT eL al. (1 960) . ( x 2.3) .

of abo ut a dozen strong spiral cords w ith
much finer subordinate ones in their interspaces.
Description. - Large-size, thick-shelled
a nd ovately naticifo rm shells with heightalmos t eq ual to w idth and with spire
about a fourth of total s hell height in
adult. Teleoconch distinctl y s houldered
an d inflated, consisting of about two
w horls, and se parated from each otherby impressed suture. Pro toconch consisti ng of 1.5 smooth a nd naticiform
Table 3.

w horls, and delimited from teleoconch by
an indistinct line. Bod y who rl large,-'slightly w ider t han high and plumply
inflated; sut ural ramp of body w horl
weakly inclined abaxially and somewhat
concave. Apert ure rather narrow, prosocline to s hell axis, oval in outline withalmost continuous peristome, and weakly
angulated above and regularly rounded
below. Outer lip thin and beveled, broadly rounded , but digitated in accordancewith spiral cords, and merged into rounded

Meas ure ments in mm. [H ayamia rex K ASE , sp. nov. ]HeightWidth

Height ofa perture

NS M PM 15030 Parat ype
SM PM 15031 Ho]ot y pe
NSM PM 15032 Paraty pe
NSM PM 15033 Parat y pe
NS M PM 15034 Parat ype

33 . 929 . 4
29 +
21. 9

39.1
32. 4
30. 1
26 +
24. 4

25. 1
22.4
22. 5+
19.3

31. 9
28 . 9NSM PM 15035 Parat ype
NSM PM 15036 Parat ype

23 . 1
12.7

22. 5

20 . 0

13.9

11. 2

22 . 3
12.6

Spec imen

Hei g ht ofbody wh orl

29.5

27 +
21. 3

716.

Early Cretaceons Gastropoda j1'mn Choshi

basal lip without any angulation. Innerlip smooth and concave, and extended
abaxially to opposite side of aperture
forming a narrow labial area. Parietal
lip smooth and covered by thin callus.
Surface ornamented by a dozen roundtopped and variably strengthened spiralcords, and by much finer subordinate
cords in their interspaces. Growth lines
distinct, more or less prosocline and somewhat rugose near aperture. Umbilicus
absent except for a weak depression atcorresponding place. Operculum calcareous, thick, symmetrical and elliptical in
outline, lacking a quadrangular process.
Remarks.- Ten specimens are available
for the present study. Four of them,illustrated in Plate 35, Figures 6-9, arenearly complete. The number of spiralcords and the width of their interspaces
are variable among the specimens. The
holotype possesses twelve spiral cords on
the body whorl, but one of the paratypes(Plate 35, Figures 7a-c) has 16 spiral cords
and those of the base are much finer andcrowded. One or two fine subordinate
spiral cords are sometimes intercalated
between the spiral cords on the whorlsides. The height of the spire in proportion to the total shell height gradually
increases with growth; that is. the bodywhorl abruptly abuts on the precedent
whorl anteriorly, and, therefore, the sutureline greatly increases its angle with theaxis and runs obliquely downward acrossthe face of the previous whorl. Similarchange of shell form is also known in
the Recent species Neritopsis radula(LI;-\KE).

The new species is closely related to
Neritopsis spiralicrenata COSSMANN, 1907
from the Barremian of France. DELPEY(1940) assigned Neritopsis spiralicrenata
to the genus Nododelphinula COSSMANN,
1916 and referred it to Nododelphinula
neritopsoides (BLAKCKENHORN, 1890). Cos-305

SMANN's species, however, belongs to thegenus Hayamia, because the labrum isnearly holostomous and it lacks an umbilicus in addition to the similar surface
sculpture. The French species is ofsmaller shell size, has coarser crenulated
spiral cords and much coarser and lessnumerous spiral threads between the
spiral cords than Hayamia rex.
Occurrence.-Known only from the type
locality [Loc. 2],

IJayamia choshiensis KASE, sp. nov.
PI. 35, Figs. 3, 4.

Alaterial.-Holotype: NSM PM 15040, anearly complete specimen (PI. 35, Figs.4ac). Para type : NSM PM 15041.
Diagnosis.-A small species of Haya171ia
with distinctly shouldered body whorl,
moderately protruded spire and obliquelyelongated aperture. Surface sculptured
by about 40 spiral cords with several
subordinate ones, and by prosocline weakcostellae on posterior part of body whorl.
Description.-Small and obliquely elongate turbiniform shells with moderately
thick test. Height almost equal to width.
Spire occupying about 0.3 of total shellheight. Whorls well rounded, rapidly
expanded and at least three in number
including protoconch, and separated fromeach other by an impressed suture. Protoconch poorly known, consisting ofsmooth rounded whorls. Body whorllarge, a little wider than high, with awide and weakly concave sutural ramp
near aperture. Aperture moderately wide,
higher than wide, angularly circular inoutline and slightly prosocline to shellaxis with almost continuous peristome.
Outer lip moderately thick, beveled andangularly convex. Surface ornamented
by regularly spaced spiral cords withseveral subordinate ones in interspaces;306

Tomo1ci ]{ASE and Ha1'uyoshi MAED A
Table 4.

Measurements in mm. [Hayamia choshiensis K ASE , sp. nov. J

Specime n
NS M P fv! 15040 Holot y pe

Hei g ht

Width

Height ofaperture

Height ofbody whorl

Ap ical
angle

19.1

20.3

16. 3

18. 5

ca. 180·

spiral cords narrower than interspaces,
about 40 in number on bod y whorl, andbecoming finer to wards base; costellae
distinct on posterior part of bod y whorl.
Growth lines fine , distinct and prosocline.
Operculum unknown.
Remarlcs.-Two specimens are available
for stud y. The protoconch of th e holot y pe is deformed and its delicate surface
scu lpture is more or less obliterated. The
paratype (Plate 35, Figures 3a, b) lacks its
outer lip, but clearly reveals the surface
ornamentation which consists of fine,reg ularl y spaced spiral cords and distinct
prosocline axial coste llae on the posterior

part of the bod y whorl. A s discussed in
the precedent pages, this group changes
its shell profile through growth. Judging
from the condition of overlappin g of thebod y whorl on the penultimate whorl,
however, the holot ype seems to be an
adult.
The present species is related to Neritopsis multicostulata PCELINCEV, 1931
(DRUSCH ITS and K UDln AV TSEV, 1960;
PCELI 'CEV, 1963) from the Barremian of
Crimea . Neri topsi s mult icostula ta , however, has larger shell size, indistinctly
shou ld ered whorls, slightl y thicker spiralcords, circular aperture, and distinct andmore numerous axial costellae on eachwhorl.
Hayamia rex differs from Ha yamia
choshiensis by much larger shell size,
circul ar aperture and different type of
surface sculpture.
Occu1"rence.- Known onl y from the t ype
localit y [Loc. 2].
Suborder doubtful
Superfamily Am berleyacea
Famil y Am berleyidae
Ge nus flmberleya MORR IS and
LYCETT, 1851
Subgenus Eucyclus Eum:sDESLONGC HAMPS, 1860fl mberleya (Eucyclus) japo1lica
KASE, sp. nov
Amberleya (Eucyc!lI s) ja/Jon.ica
K AS E, sp. nov. (x 1. 2) .

Te x t-fig. 8.

PI. 34, Figs. 15, 16 ; Te x t-fig. 8.

716.

Early Cretaceous Gastrol)oda from Choshi

Material.-Holotype: NSM PM 15042, alarge specimen (PI. 34, Figs. 15a, b). Paratype: NSM PM 15043.
Diagnosis.-A large and elongate speciesof Amberleya (Eucyclus) characterized bybroad ramp and sculpture consisting ofsmooth and round-topped spiral cords.
Description .-Shell large-sized, moderately thick-tested, anomphalous and elongate turbiniform with height less than
twice width. Spire conical with an acuteapex being less than a half total shellheight. Apical angle about 55°. Protoconch not preserved. Suture impressed.
Whorls at least five in number. Spirewhorls angulated at middle, concave overposterior half forming a broad ramp,
peripherally flattened and bearing sevenor more strong and round-topped spiralcords. Spiral cords gradually weakened
anteriorly. Aperture subovate, subparallelto shell axis, acute above and roundly
projected below. Outer lip thin at edge,
obliquely straight above, angulated at
middle and rounded below. Parietal lip
vertical and thick. Growth lines distinct
and rugose.
Remarks.-Two specimens are available
for this study. The holotype (NSM PM
15042; Height 57 mm+ ; Width, 37.5 mm;
Height of aperture, 26 mm+; Apicalangle, ca. 55°) is poorly preserved andits apical part is slightly deformed, but
the shell outline and surface sculpture are
well shown except for the basal part.
Spiral cords on the body whorl of theholotype cannot be counted precisely
owing to poor preservation, but the
number at least seven. Basal characteristics of this species are well revealed inanother fragmental specimen which showsan anteriorly projected, rounded, basal lip
lacking an anterior siphonal canal.
A number of species assigned to AI/lberleya (Eucyclus) reported from theCretaceous strata of the world do not307

agree with the new species in shell sizeand surface sculpture. Among these
species, A771berleya dilleri STANTON, 1895
from the Tithonian Knoxville beds ofCalifornia is somewhat similar to thepresent species in having a smaller shellsize (maximum height is less than 35 mm)
and thinner granulated and crest-topped
spiral cords. The present new species
also somewhat resembles Turbo faucignyanus PICTET and Roux, 1849 from theAptian of France. But the French species
has smaller shell size and wider shell
outline than our species, and has a biangulated body whorl with much finer spiralcords on the base and ramp.
Occllrrence.-Known only from the type
locality [Loc. 1].
Genus Oolitica COSSlvlANN, 1893
Oolitica sp.
PI. 34, Figs. 4a, b.

Relllarks.-This species is represented
only by a poorly preserved specimen(NSM PM 15044; Height, ca. 13 mm ;Width, 10 mm; Height of aperture, 8.5mm; Apical angle, ca. 80'). This species
is characterized by littoriniform shell
outline, broad and steeply inclined ramp,
and six or more spinous and crest-topped
spiral cords on the body whorl. The
umbilical area is poorly preserved, having
a wide and subelliptical aperture anddiscontinuous peristome. The shell surface is covered by prosocline and wrinkled
growth lines. These features are common to the genus Ooli/ica COSSlvlANN,
1893. The fragmentary nature of theavailable material precludes placement inhitherto known species assignable to thisgenus.
Occllrrence.-Loc. 1.

Tomoki KASE and Haruyoshi MAEDA

308

separated from each other by weakly
impressed suture. Protoconch missing.
Body whorl large, as wide as high orslightly wider than high and weakly convex above, with rounded periphery. Basebroad, weakly convex peripherally but
concave at umbilical part. Umbilicus
narrow and grading to base without
angulation. Aperture subcircular, slightly
higher than wide and inclined to shellaxis with discontinuous peristome. Outer
lip rounded and weakly crenulated inaccordance with peripheral cords. Inner
lip regularly con cave and subverticalwith slightly reflected margin. Surface
sculpture consisting of finely tuberculated
spiral cords; number of spiral cords fouron the spire whorls and about 14 on bodywhorl. Spiral cords of body whorl becoming thicker and more finely tuberculated towards umbilicus and slightly
narrower than their interspaces. Sculpturein umbilicus not observed.
Remarks.-A number of specimens areavailable for study, but most are incomplete. The holotype (Plate 34, Figures
14a-c) is a large and comparatively wellpreserved specimen, which shows well
the surface ornamentation of the adapertural part, but the characters of theumbilical and apertural areas are poorly
known. Umbilical characters are well
exhibited in one of the paratypes (Plate
34, Figures 2a, b). The variability of shellform cannot be appreciated because of
the scarcity of well preserved specimens.
However, the smaller specimens (e. g.

Family Nododelphinulidae
Genus Metriomphalus COSSMANN, 1916
iv!etriomphalus nagasalziensis
KASE, sp. nov.
PI. 34, Figs. 1, 2, 14.

1972. Margardes (?) sp., SIIIKA:-IA and St;ZUKI, Sci. Rep. Yokohama Nat. Unit'.,sec. II, no. 19, pI. 6, fig. 15.
?l972. Margarites aff. sachalinensis NAGAO:SHIKAMA and SUZUKI, Sci. Rep. Yokohama Nat. Univ., sec. II, no. 19, pI.
6, fig. 16.
1977. Margarites? sp., HAYA:-'" and KASE,
Univ. Mus. Univ. Tokyo, Bull., no.
13, p. 35.
?1977. Margarites? sp., HAYA!dl and KASE,
Univ. Mus. Univ. Tokyo, Bull., no.
13, p. 35.

jvfaterial.-Holotype: GIYU-MM-104, alarge specimen (PI. 34, Figs. 14a-c). Paratypes: GIYU-MM-105-107.
Diagnosis. - A medium-sized species ofk!etriomphalus characterized by low spire,
weakly impressed suture and ornamentation consisting of coarse tuberculated
spiral cords.
Description. - Medium-sized, narrowly
phaneromphalous and moderately thickturbiniform shells with height greaterthan width in small specimens but nearly
equal to width in large ones. Spire small
and low, comprising about half of total
shell height. Whorls weakly inflated, atleast 4.5 in number, rapidly expanded andTable 5.
-

Measurements in mm. [J'vletriomphalus nagasakiensis KASE, sp. nov.~-----

Specimen------

GIYU-MM-I04 Holotype
GIYU-MM-I05 Paratype
GIYU-MM-I06 Paratype
GIYU ·MM-I07 Paratype

~

~---

-

~

Height

Width

-----

12.0
ca.9
ca.8
----

-- ------Height of aperture------~--.-

7.0
4.3
ca.4
5.0

11. 6
8.9
7.5
9.0
-----

716.

Early Cretace ow; Gastropoda from Choshi

Plate 34, Figures 2a, b) differ in shapefrom the larger ones in having a higher
shell outline and biangulated body whorl.
The surface sculpture, apertural and umbilical characters do not differ among
these specimens. The observation ofmany specimens of various growth stages
leads me to conclude that the angulationof the body whorl becomes weaker androunded through the stages of growth.
The present species was once illustrated
by SHIKAMA and SUZUKI (1972) under thename of Margarites (?) sp., one of theparatypes (Plate 34, Figures 2a, b) is theillustrated specimen (SHIKAMA and SUZUKI, 1972, pI. 6, fig. 15). They also illustrated one specimen from the same locality
under the name of jIargariles aff. sachalinensis NAGAO, 1939. Specimens of thatspecies are, however, so poorly preservedthat the specific as well as generic determination is difficult.
This species may be closely related toTurbo coquandi PICTET and CAl'vII'ICHE,1864 from the Gault of St. Croix, Switzerland. COSSl'vIANN (1915) assigned thisspecies to the genus Metri07nphalus.
Turbo coquandi possesses larger shell size,
deeply impressed suture, narrower aperture and more numerous spiral ccds thanthe present new species.
Turbo goupilianus D'ORBIGNY, 1843 fromthe Cenomanian of France, is related to
the new species, but is distinguished fromit by the narrower aperture, finely tuberculated spiral cords, higher spire andmore inflated whorls which are separated
by deeply impressed suture.
Occurrence.-Known only from the type
locality [Loc. 3].
Order Mesogastropoda
Superfamily Strombacea
Family Aporrhaidae

309

Genus Perissoptera TATE, 1865
Type-species. - Rosleliaria
par!?insolli
1822 from the Lower Albianof England. [original designation]~ANTELL,

Perissoptera elega7Js KASE, sp. nov.
PI. 36, Figs. 1115.

Malerial.-Holotype: NSIVI PM 15045, anearly complete specimen (PI. 36, Fig. 14).
Paratypes: NSM PM 15046-15049.
Diagnosis.-A medium- or small-sized
species of Perissoptera characterized byonly spirally ornamented body whorl andopisthocyrt sigmoidal axial ribs on spire
whorls. Body whorl with a prominentkeel. Simple and slender anterior digitation of extended outer lip projected
obliquely downwards.
Description. - Medium- or small-sized
and thick-shelled aporrhaid with highly
turriculate spire attaining less than athird of total shell height. Incremental
angle in neighbourhood of later whorls
about 35', while apical angle about 55°.
Spire whorls ten in number including
protoconch, regularly expanded in size,
roundly convex, and separated from eachother by impressed suture. Protoconch
small, rounded, smooth, naticiform, consisting of about four whorls, graduallychanging into ornate teleoconch without
a din tinct boundary. Sculpture of spire
whorls consistently composed of 13 in
number on penultimate whorl, and their
inclination against shell axis gradually
becomes stronger through growth. Spiralcords fine and numerous (about 30 on
penultimate whorl of holotype). Body
whorl angularly inflated above, but abruptly becomes narrower anteriorly, forminga moderately long, pointed and nearlystraight anterior siphonal canal. Body
whorl sculpture consisting of spiral cords,
but very weak sigmoidal axial ribs some-310

TO?no1ci KASE and Han~yoshi MAEDA
wards initially but graduall y turning upward, regularly curved , and becomesalmost parallel to shell axis at its posterior end. Pcsterior spine also curved
ventrall y. Anterior spine narrow , moderately long, extends obliquely downwards
and encounters posterior spine almost rightangle.
RemaTli'.'·.- This species is represented
by a large numter of specimens collected
at Loc. 2, but most of them are fragmental or immature, judging from thedevelopment of the outer ilp spines.
Three well preserved specimens, illustrated on Plate 36, show the nearly complete shell outline, surface sculpture andthe characteristic outer lip spine. The
extended outer lip spine of these specimens indicates that these individua ls are
mature. The holotype (Plate 36, Figure14) lacks th e protoconch and posterior
end of the posterior digitation of thebifurcatedouter lip , but
exhibits
the apertural surface. This specimen
possesses a complete anterior canal andthe mode of the bifurcation of the outer
lip spine is well observable. These features of the outer lip are considered ascharacteristic of Perisso pte ra. One of theparat ypes (Plate 36, Figure 15), reveals theadapertural surface and pos!:esses analmost complete outer lip spine and anterior siphonal canal. The mode of bifurcation of the outer lip spine is notdifferent from that of the holotype. In
another para types (Plate 36, Figure 13) the

Text-fi$7. 9. Piet teia crela cea KASE,
sp. nov. ( x 1.7) .

times occur, and axial ribs abruptl y disappeared near boundary of penultimate andbody whorls, whe re a prominent spiralround-topped keel appears. Aperture narrow and lenticu lar. Outer lip extended
into a bifurcated spine. Posterior spine
lon g and narrow, extending slightly down Table 6.

Measurements in mm. [P n issop lera eiega. ns K ASE , sp. nov. ]Specimen
NS M PM 15045 Holot ype
NSM P'vl 150'16 Paratype
NSM PM 15047 Para type
NSM PM 15049 Para type

- ----

Hei ght

Width

I-Ieight ofaperture

Incremental angle
in neighbourhoodof later w horls

45+
44 +
37 +
6. 4

ca.25
27 +
25. 5+

ca. 3029.7
25 +ca. 25 'ca . 35'ca _30'2.8

Ap ical
angle

55 '

716.

Early Cr'etaceous Gastropoda j1'om Choshi

anterior digitation of the outer lip spineis broken off, but the early whorls including protoconch are perfectly preserved.
The present species is very similar to
Anchura (Perissoplera) marginata (SowERBY, 1836), (GAR[):\ER, 1875; COSSMAN1\',
1904) from the Albian of England, butdiffers from the European species by thesmaller shell size, more flexed axial ribs,
more prominent spiral keel on the upper
part of the body whorl, and simple andnarrower anterior digitation of the outer
lip spine.
Anchura (Perisso{1lera) hannai ALLISON,
1955 from the middle Albian of Baja California is distinguishable from the present
new species by the much smaller shellsize, weakly prosocline axial ribs and thepresence of short and wide anterior digitation which is differentiated into twoparts.
Occurrence.-Known only from the type
locality [Loc. 2].
Genus Pietteia COSSMAN1\', 1904
Type-species.- Roslellaria humus DESLONGCHAMPS, 1842 from the Bajocian ofFrance. [original designation].
Diagnosis.-Medium- to large-sized aporrhaids with high and turriculate spire.
Protoconch consisting of about two smooth whorls. Spire whorls angularly inflated at middle with prominent transverse
ribs. Body whorl only spirally sculptured
with a posterior prominent and anterior
weak spiral carination. Anterior siphonal
canal long. Inner lip covered by thick andwell-delimited callus.
Remarks.-Pietteia corresponds to thesecond group of Alaria of PIETTE (1891),
and was established by COSSMANN (1904)
as a subgenus of Di(:rolonw GABB, 1868.
WENZ (1938), on the other hand, placed
Pietteia as a subgenus of Anchura CONRAD,
1860 together with Dicroloma. Pietteia:311

differs decidedly from Anchura by thepresence of a simple, narrow and longouter lip spine, posterior prominent andanterior weak spiral carinations on the
body whorl, angulated spire whorls andpresence of a rather long spine at theramp angle of the body whorl. The presence of the long spine at the ramp angleof the body whorl is also found in certain
species of Dicroloma. Dicroloma, however,is distinguished from Pietteia by some
important points; that is, Dicroloma possesses a bifurcated outer lip spine, only
spirally sculptured whorls, a prominent
anterior spiral carination on the bodywhorl and the lack of thick and welldelimited callus on the inner lip. For
these reasons, Pietteia is here accepted
as a distinct genus.
Geological range of Pietteia, according
to COSSMAN1\' (1904), is from the Sinemurian to Sequanian.
No Cretaceous
species has been assigned to Pietteia except for one species, Anchu ra (Pietteia)
schweifurthi (QUAAS, 1902) from theMaestrichtian of Egypt (ABBAS, 1963).
However, the Egyptian species, according
to the descriptions and illustrations by
QUAAS (1902) and ABBAS (1963), cannot
be assigned to Pietteia because of itsdifferent surface sculpture and lack of aspine on the ramp angle of the body whorl.
The new species described in this paper,
therefore, is the first record of true
Pie tteia from the Cretaceous.
Pietteia cretacea KASE, sp. no\'.
PI. 36, Figs. 4-10; Text-lig. 9.

Material.-Holotype: NSM PM 15050,(PI. 36, Fig. 10). Paratype: NSM PM
15051-15055.

Diagnosis.-A large-sized species of
Pietteia characterized by long outer lip
spine and a long spine at ramp angle ofTomoki J{ASE and Haruyoshi MAEDA

312

body whorl three-fourths volution backfrom outer lip. Aperture broad. Spirewhorls with broad subsutural ramp andornamented by eight to ten slightly prosocline raised axial ribs.
Description.-Shell larse-sized having
high, evenly tapering and slightly coelconoid spire. Incremental angle in neighbourhood of earliest teleoconch about 20·.
Spire attaining about one half of total
shell height with eleven angularly inflated
whorls. Suture impressed. Protoconch
poorly known, consisting of about two
small and smooth whorls. Each spire
whorl with concave and steeply inclined
broad ramp. Spire whorls sculptured byaxial ribs and spiral cords. Axial ribs
strong, weakly prosocline, round-topped,much narrower than their interspaces andten in number on penultimate whorl buteight or less on early whorls. Axial ribs
obscure on ramp, but abruptly appearing
near ramp angle. Spiral cords very fineand regularly spaced, two of which prominent at middle position of whorl andbecoming more or less tuberculated atintersections with axial ribs. Posteriorcarination continuing with outer lip spine
and appearing near boundary between
penultimate whorl and body whorl andgradually strengthened towards aperture.
Several prominent spiral cords present
below anterior carination. Outer lip spine
thick, stout, and extending nearly horizontal at first but rather abruptly turned
slightly backwards with pointed end.
Aperture broad, elongate subovate, rounded
Table 7.

posteriorly, and angulated anteriorly
in harmony with siphonal canal. Outer
lip rounded; posterior part of outer lip
thick and opisthocyrt, and anterior partthin and sharp at its edge. Inner lip
regularly concave with a well-delimited
heavy callus. On body whorl a long andthick spine situated at the ramp angle
three-fourths volution back from outer lip.
Remarlzs.-Five specimens are available
for study. The two characteristic carinations on the body whorl are well seen on
one of the para types (Plate 36, Figures
8a, b), although its apical part and anterior
siphonal canal are broken. The long spineon the ramp angle in this specimen lacks
a free end which presumably curves towards ada pert ural direction and seems toextend only a little judging from its
termination. The presence of the spineon the ramp angle is also shown in theholotype and two specimens (Plate 36,
Figures 5, 9), which bear the mark ofspine at three-fourths volution back fromthe outer lip. The sculpture of the bodywhorl is well represented on the holotype
and two of the para types (Plate 36, Figures4, 5), comprising two prominent spiralcarinations and several subordinate ones,and very fine spiral cords. The spine at
the ramp angle of the body whorl areconsistently present at three-fourths volution back from the outer lip.
Compared with Anchura (Pietteia) hamlls(DESLONGCHAMPS, 1842) (PIETTE, 1891;
WENZ, 1940) from the Bajocian of France,
the new species is larger and has a longer

Measurements in mm. [Pietteia cretacea KJ\SE, sp. nov.]

716.

Ea1'ly Cretaceous Gastropoda from Choshi

outer lip spine, wider aperture, smaller
spical angle and finer spiral cords.
Occurrence.-Known only from the type
locality [Loc. 2].
Genus Ceratosiphon GILL, 1870[=Ornithopus GARDNER, 1875;
non HIcHcacK, 1848J
Type-species. - Pterocera moreausianai)'ORBIGNY, 1848 from the Neocomian ofFrance. [original designationJ
Diagnosis.-Moderately large-sized andshort spindle-shaped aporrhaids with low
spire. Whorls generally about seven in
number and prominently angulated at
middle with only spiral sculpture. Protoconch consisting of two or three, initial,
smooth, naticiform whorls, and a succeeding rounded whorl with reticulated ornamentation. Body whorl bicarinated. Outer
lip extended to three long, narrow andtapering spines, grooved internally withcorresponding external ridges; posterior
spine adnate to spire. Inner lip coveredby a thin but widely spread callus. Anterior siphonal canal long and thin with(or without) a thin spine branching fromneck to opposite side of aperture.
Remarks.-Several aporrhaid gastropods
from the Cretaceous beds from Europe,
which were listed by CassMANN (1904, p.
57, 58) as members of Tessarolax GABB,1864 are not typical for the genus asstated by STEW ART (1926, p. 364). Ceratosiphon was established by GILL (1870)(STEWART, 1926) on the basis of Pterocera
moreausiana D'ORBIGNY from the Neocomian of France as the type-species. Subsequently, GARDNER (1875) proposed thegenus Ornithopus on the basis of Rostellaria 1'etusa SaWERBY, 1836 as the typespecies, but the generic name was preoccupied by HrCHcaCK (1848) (CassMANN,
1904) for a reptile. CassMANN (1904), as fol-313

lowed by WENZ (1940), regarded Ceratosiphon as the junior synonym of Tessarolax,
stating "***En ce qui concerne Ceratosiphon GILL, dont Ie type est Pterocera
N/oreausiana D'Orbigny, je ne puis apercevoir absolutment aucune difference
sectnelle entre cette coquille et Tessarolax:l'absence de gibbosite opposee au labren'est probablement que fortuire chez P.
Moreausiana, Ie labre est peut-etre unpeu plus parme que chez T. distortum,mais les digitations sont absolument semblables. Par son sequent, Ceratosiphon est
synonym posterier de Tessarolax."
Since then, several aporrhaid gastropod
species characterized by biangulated bodywhorl and five long and narrow spines
mainly from the Lower Cretaceous inwestern Europe were regarded as Tessarolax. This is probably because of theincomplete preservation of the materialof Tessarolax distortum and incomplete
generic definition of this genus. In my
opinion a prominent axial projection on
the opposite side of the body whorl of theaperture in the genus Tessarolax is conservativeof this genus. Tessa1'olax,in general,
has larger shell size than Ceratosiphon, more
numerous spire whorls, higher spire, thickdeposition of callus over the shell surface
and the anterior siphonal canal without abranching spine. The two, therefore,
must be generically distinguished. Tessarolax is probably well represented in
the Upper Cretaceous of North America,
Japan and southern India. Ceratosiphon,on the other hard, previously was known
only from the Lower Cretaceous and Cenomanian of western Europe.
The following species appear to belong
to Ceratosiphon:
Pterocera moreausiana D'ORBIGNY, 1843
(GARDNER, 1875) from the Neocomian ofFrance.
Rostellaria retusa SaWERBY, 1836 (GARDNER, 1875; CassMANN, 1907) from the314

TO?11,Oki KflSE and Han~yoshi MAEDA

Lower Albian of England .
Pteroeera fittoni FORBES, 1845 from theLower Aptian of England.
Rostellaria biearinata DESHAYES in
LEYlVlERIE, 1842 from the Albian of France.
D'ORBIGNY (1843) described one aporrhaid
gastropod species under the name of
Pteroeera bicarinata from the Albian ofFrance. His figure has a hig her spire
than a t ype specimen of R oste!Laria biearinata DESH AYES and has a distinct
projection on the opposite side of the
body w horl of the aperture, and is ve ry
different from DESHA YES' S figure.
Aporrhais histoehila GARDNER, 1875
from the A lbian Upper Greensand of
England .
Aponhais oligochila GARD NER, 1875
from the Cenomanian Gray Chalk ofEngland.
Aponhais ebrayi DE LORIOL, 1882 fromthe Gault of France.
? Aponhais (Te ssarolax) gardneri KEEPING, 1883 from the Upper Ap tian ofEngland.
Aporrhaid gen. et sp. indet., MATSUKA W A, 1977 from the upper part of the
Ishido Formation (Barremian) of Sa nchu
area, central Japan.

Ceratosiphon densestriatus
KASE, sp. nov.
PI. 36, Figs. 1- 3; Text ·fig. 10.

NJaterial.- Holot ype : NSM PM 15056,
a nearl y complete specimen (PI. 36, Fig. 1).
Paratypes: NSM PM 15057-15062.
Diagnosis.-A large-sized species of
Ceratosiphon characterized by a proportinally higher spire, prominentl y biangulated body whorl and impressed suture.
Posterior spine extended slightly beyond
spire. Anterior siphonal canal thin witha thin spine branching from posterior partat nearl y a right angle. Surface sculptured

Text-fig. 10.
K ASE ,

CeratosipllOn densestriatlls

sp. nov. (x 2.5).

by numerous very fine spiral threads.
DesC1'iption.-Shelllarge but thin-shelledwith more or less abruptly tapered spire.
Incremental angle in neighbourhood ofbody and penultimate whorls about 40°
but apical angle about 90°. Volutions
seven or eight in number. Protoconch of
three smooth and well rounded naticiform
w horls, followed by a fourth well rounded
w horl bearing cancellated ornamentationconsisting of six distinct and regularly
spaced fine spiral cords and similarly
strengthened opisthocyrt axial threads.
Spire whorls reg ularly expanded, separated by impressed suture. Whorls abruptly
changing from protoconch wi thout distinct
boundary. Second and third teleoconch
w horls prominently angulated at middle
vvith broad ramps and nearl y verticalw horl sides. Surface sculpture of second
teleoconch whorl consisting of three spiralcords on ramp and nine finer on es on thew horl side. Third teleoconch w horl with

716.

Early Cretaceous Gastropoda from Choshi

much finer and numerous spiral cords orthreads. Axial sculpture of these whorls
much finer than that of first teleoconch
whorl. Body whorl prominently biangulated at periphery, convex and anteriorly
constricted, and ornamented by muchfiner and numerous spiral threads. Aperture wide and semilunar in outline. Outer
lip with three extended outer lip spines.
Posterior spine thick, ad nate to spire andmore or less extended posteriorly beyond
spire; two other spines extended fromtwo ridges of body whorl and moderately
long for this group. Anterior siphonal
canal thin and slightly curved with a thinspine branching from posterior part atnearly a right angle. Inner lip long andnearly straight with a thin callus that
extends onto body whorl.
Remarks.-Six specimens are available
for this study. The holotype is a nearlycomplete specimen and reveals well theshell characters of the apertural side, but
the ends of the anterior spine of the outer
lip and the thin spine branching from theanterior siphonal canal are not preserved.
All other specimens are incomplete andfragmental, two of which illustrated in
Plate 36 show the apertural and adapertural
surface characters and the characteristic
protoconch. The presence of a long andslender spine branching from the neck of
the anterior siphonal canal is well preservedin the holotype and one of the paratypes(Plate 36, Figure 2a). Several other specimens, which lack anterior siphonal canal,
evidently show the trace of the branching spine.
Aporrhais ebrayi DE LORIOL, 1882 fromthe Gault of Cosne, France is very similar
to Ceratosiphon densestriatus in the shell
outline and the arrangement of the outer
lip spines and anterior siphonal canal, butpossesses much coarser spiral sculpture,less prominent spiral angulations on the
body and spire whorls and shorter poste-

315

rior outer lip spine than Ceratosipnon
densestriatus.
The new species is easily distinguishable
from Rotellaria retusa SOWER BY, 1836
(GARDNER, 1875; COSSMANN, 1907) from theAlbian of England by its higher spire, muchfiner spiral sculpture, impressed sutureand shorter and slender outer lip spines.
Occurrence.-Known only from the type
locality [Loc. 2].
Superfamily Hipponicacea
Family Vanikoroidae
Genus Vanikoropsis MEEK, 1876
Type-species.-Natica toumeyana MEEK
and HAYDEN, 1856 from the Senonian ofNorth Dakota. [original designation]
Vanikoropsis decussata (DESHA YES)
PI. 35, Figs. 1, 2.

1842. Littorina decussata DESHA YES in LEYMERlE, Mhn. Soc. geol. France, vol. 5,
fase. 1, p. 13, pI. 17, fig. 16.
1842. Turbo deeussatus (DESEAYES): D'ORIlIGNY, Paleont. fran,., terr. eret., vol.
2, p. 219, pI. 184, figs. 11-13 (nonMONTAGU, 1815).
1850. Turbo alsus D'OR13IGNY, Prodrome dePateont., vol. 2, p. 130.
1863. Turbo alsus D'ORIllGNY: PICTET and
CAMPICHE, Mater. Pateont. Suisse, ser.
2, p. 496.
1882. Turbo alsus D'ORBIGNY: DE LORIOL,
Mhn. Soc. paleont. Suisse, vol. 9, p. 3133, pI. 3, figs. 16-20.

Material.-NSM PM 15063, 15064.
Diagnosis.-A very small species of
Vanikoropsis characterized by having thintest and surface sculpture of narrow spiralribbons and transverse ribs.
Description.-Shell very small, thin,phaneromphalus and elongate turbiniform

816

Tomoki KASE and Ha1"uyoshi MAEDA
Table 8.

Specimen--~

Measurements in mm. [Vanikoropsis decussata (DESIIAYES)]
HeightMaximumwidth--_._----NSM PM 15063
NSM PM 15064

Minimum
width

ca.6.8ca.7.95.3

with conical and accuminate spire. Spireoccupied about two·fifths of total shellheight. Whorls at least five in number
and regularly expanded. Spire whorls
moderately convex and separated by impressed suture. Body whorl large, slightly wider than high with regularly rounded
whorl sides. Umbilicus widely open, deep
and delimited from roundly convex basewithout any angulation. Aperture moderately wide, obtusely acute above androunded below, and almost parallel toshell axis, with height about 1.4 times
width. Parietal area with thin callus.
Inner lip weakly arched over parietal andcolumellar area and well rounded below.
Surface of spire whorls sculptured by five
narrow ribbons and slightly prosocyrt
transverse ribs, and tuberculated at theirintersections. Transverse ribs 20 to 24 in
number on penultimate whorl. Body whorl
sculptured by narrow spiral ribbons andsomewhat sigmoidal transverse ribs. Spiralribbons wider on anterior half than thoseon posterior half and 18 to 20 in number.
Transverse ribs gradually weakened towards base and also near aperture.
Remarks.- Two specimens are available
for study. One illustrated specimen (Plate
35, Figures la-c) is well preserved and well
exhibits the apertural features and delicate
surface sculpture, but its protoconch is
missing. The other illustrated specimen(Plate 35, Figures 2a-c) lacks the aperturalpart, and the shell surface is somewhat
abraded. The crowded growth lines near
the aperture, which are well observed infirst specimen, are assumed to be a feature

4.2
ca.5.4Height ofapertUi e~~--

4.0
ca.4.6

Incremental angle inneighbourhood of penul.
timate and body whorls----- ------

70·
80·

of maturity. The first specimen differs
in shape from the second by smaller apicalangle and more slender shell outline, butthey must be conspecific, judging fromthe surface sculpture and umbilical character.
In every essential character the present
specimens agree well with Turbo alsus(DESHA YES) sensu DE LORIOL (1882) fromthe Gault of Cosne in France. The shellsize, ornamentation and elongate turbiniform shell outline are especially similar
to those of the specimens from Cosne.
That species was established by DESHA YES
in LEYMERIE (1842) from the Gault of
Aube in France. The present specimens
have wider shell outline than those of
The specimen illustrated by
Aube.
d'ORBIGNY (1842), on the contrary, hasmuch higher shell outline than that of
Aube, but is similar to those of Cosne.
DE LORIOL (1882) stated that the discrimination of the specimens between Aubeand Cosne is very difficult so far as therange of variation of the specimens from
Cosne was examined.
Taxonomic position of the genus Vanikoropsis and assignment of several Jurassic and Cretaceous species to this genus
were discussed by SOHL (1967). According to him, only two species, Vanikoropsis
foumeyana (MEEK and HAYDEN) from theMaestrichtian of. North Dakota and Vanikoropsis nebrascensis (MEEK and HAYDEN)
from the Maestrichtian of Wyoming,
Montana, South Dakota and Colorado,
have been known, and several species
once assigned to Vanikoro or Vanikoropsis

71 6.

EaTly CTe ta ce o~~s GastT01Joda from ChoshiA
Te x t·fi g . 11.

317

B
E riplycha jap onica [( AS E, sp. no v. (x S.7) .

belong to oth er genera. Recentl y, HAY Aand KASE (1 977) assig ned N atica(Lunatia) ainuana NAGAO, 1939 from theConiacian or Santonian Upper Ammonite
Bed (Upper Yezo Group) of south Saghalin
to Vanilzoropsis. The present species agrees well with the above-mentioned Upper
Cretaceous species in shell outline, surface
sculpture and umbilical feature , and isprobably congeneric with them. However, it differs from the three Upper Cretaceous species by the much smaller shellsize, thinner shell and sculpture consistingof spiral ribbons and transverse ribs.
Occunence.-Loc. 2
MIOrder Cepha laspidea
Subfamily Acteonacea

Diagnosis.- Large- to small-sized andlow-spired ringiculid shell. Much thickened outer lip denticulate within. Columellar lip with one fold which sometimes
bifid. Parietal lip much thickened bycallus with a various-shaped prominent
axial ridge. Surface sculptured by punctate spiral grooves. Anterior canal moreor less defined.
Remar12s.-Discrimination of the genus
Avellana from the genus Eriptycha andsome other Cretaceous ringiculid genera
is somewhat controversial. This is partly
because the characteristics of the inner
lip structure of the genus Avellana are
not clear. Eriptycha is very similar to
A vellana in shell outline, s urface sculpture
and outer lip structure, but differs fromthe latter genus by the parietal lip whichbears a strong axial ridge.

Family Ringiculidae
Genus Eriptycha MEEK, 1876[= Euptycha MEEK, 1863; non
HUBNER, 1826J
Type-species.-Au ricula decU1'taia SowERBY, 1831 from the Turonian of Gosauin Austria. [original designationJ

E riptycha japonica KASE, sp. no v.
Pi. 34, Figs . 12, 13 ; Te x t·fi g . 11.

Ma terial.- Holot y pe: NSM PM 15065(PI. 34, Figs. 12a, b), Paratype: ISM PM
15066
Diagnosis.-A very small species ofEriptycha characterized by short spire318

Tomoki KASE and Haruyoshi MAEDA
Table 9.

Measurements in mm. [Eriptycha japonica KASE, sp. nov.]

Specimen
NSM PM 15065 Holotype
NSM PM 15066 Paratype

HeightMaximum width

Minimum width

Height of aperture

5. 0
5. 2

4. 2
4.7

3. 1
3.7

4. 6
4. 7

and cylindrical shell outline. Denticles
along inner margin of outer lip weak andabout 20 in number. Parietal axial ridge
reinforced with a prominent V-shaped
projection on lower part. Anterior siphonal
canal deep and wide. Spiral grooves ofbody whorl about 30 in number.
Description.-Shell very small, lowspired and subelliptical in front view.
Height slightly greater than width. Teleoconch regularly rounded and rapidly
expanded with about 2.5 volutions. Suture
impressed.
Heterostrophic protoconchvery small, pointed above teleoconch, andconsisting of at least one smooth whorl.
Body whorl very large, occupied about
0.9 of total shell height, with regularlyrounded sides. Aperture narrow, constricted by prominent fold and ridge,
narrowed and tapered posteriorly andbroadened anteriorly with reversed comma-shaped outline. Outer lip long, almost
parallel to shell axis, reinforced, moderately thickened by callus that is reflected
out of aperture back onto body whorl.
Inner surface of outer lip with about
20 weak denticles. Inner lip covered bywell-margined callus, long, convex inposterior but strongly concave below andcontinues to outer lip forming a posterior
canal. Parietal wall bearing a J-shaped
strong ridge with a vertical bar as aridge subparallel to posterior canal andits abaxial edge is prominently reinforcedby callus. A prominent V-shaped projection is present on lower part of parietal
wall. Columella with a strong and nearlyhorizontal fold which extended to aperture. Surface of body whorl sculptured

by about 30 punctate spiral grooves that
separeted by flat-topped spiral cords.
Remarl?s.-In this collection the present
species is represented only by two specimens. The holotype is complete andshows the characteristic features of innerand outer lips. The para type also shows
the apertural features, but the shellsurface is more or less abraded. The
heterostrophic protoconch is well preserved in the holotype aFld consists of asmooth whorl, but its earlier whorl(s)seems to be submerged below the surfaceof the teleoconch.
Avellana minima NAGAO, 1934 from theAptian Miyako Group of Northeast Japan
is very similar to the new species inmany essential points. Many well preserved specimens of Avellana minima collected near the type locality reveal thatthe much thickened parietal lip has areinforced axial ridge in the upper partand a plication in the lower part, thecolumellar lip has an almost horizontal
fold at its middle and the outer lip isdenticulate internally. These featuresindicate that the two species are congeneric. Avellana minima, however, differsfrom the new species by the wider shell
outline, smaller number of denticles oOn
the outer lip (eight in number) and smaller
number of spiral grooves (15 or 16 on
the body whorl).
The new species resembles Euptycha
globata STOLICZKA, 1868 from the Turonian or Coniacian of southern India, butdiffers from that species in the ellipticaland much smaller shell, lower spire, one
prominent V-shaped projection on the716.

Early Cretaceous Gastropoda !1'om Choshi

lower part of the parietal wall and finersurface ornamentation.
Occurrence.-Known only from the type
locality [Loc. 2].
Acknowledgements

We express our deep gratitudes to Professor Tetsuro HANAI (Geological Institute,
University of Tokyo) and Dr. Itaru HAy AMI (University Museum, University ofTokyo) for their invaluable suggestions
and encouradgements which stimulated
the present work. Deep gratitude is also
expressed to Dr. Norman F. SOHL (U. S.
Geological Survey, Washington, D. C.,
U. S. A.) for critical reading the manuscript and gave us invaluable advice. We
are much obliged to Dr. Tadashige HABE
and Dr. Ikuwo OBATA (National Science
Museum, Tokyo) for their useful suggestions, and also to Mr. K. OZAKI (Yokohama National University) and Mr. S.
SAKUMA (Chikyu Gijutsu Kaihatsu Co.)for their permission to observe the specimens preserved in the collection room of
the Yokohama National University. Acknowledgments are also due to Messrs. T.
On, S. MORIYA, S. UMINO and M. TSUKUI(University of Tokyo), and also to Mr. Y.
KASE (Choshi Senior High School) who
kindly offered their collections. Dr. K.
CHINZEI (University of Tokyo) and Mr.
A. MATSUKUMA (National Science Museum, Tokyo) kindly sent us copies ofimportant papers. Miss. Y. WATANABE
drew the sketches.
This study was supported in part by
the Grant in Aid for Co-operative Research (No. 334043) of Ministry of Education.
References
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320

Tomoki EASE and Har1,~yoshi .lV[AEDA

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1. Front view of a paratype (x 2.0) from the Kimigahama Formation at Loc. 3. GIYUMM-107.
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5a-c. Front, back and basal views of a paratype (x 3.0) from the Kimigahama Formation
at Loc. 2. NSM PM 15003.
6a-c. Front, back and basal views of a paratype (x 3.0) from the Kimigahama Formation
at Loc. 2. NSM PM 15002.
7a, b. Back and front views of the holotype (x 3.0) from the Kimigahama Formation at
Loc. 2. NSM PM 15000.
Figs. 3a, b, 8a, b, 9a, b, lOa, b, lla-c. Ataphrus (s. str.) nipponicus KASE, sp. nov.
3a, b. Apical and front views of a paratype (x 2.5) from the Kimigahama Formation at
Loc. 2. NSM PM 15017.
9a, b. Front and back views of a para type (x 2.5) from the Kimigahama Formation at Loc.
2. NSM PM 15011.
lOa, b. Front and back views of a paratype (x 2.5) from the Kimigahama Formation at
Loc. 2. NSM PM 15012.
lla-c. Back, front and apical views of the holotype (x 2.5) from the Kimigahama Formation
at Loc. 2. NSM PM 15007.
Figs. 15a, b, 16. Amberleya (Eueylus) japonica KASE, sp. nov.
15a, b. Front and back views of the holotype (x 1.0) from the Isejigaura Formation at
Loc. 1. NSM PM 15042.
16. Front view of a para type (x 1.0) from the Isejigaura Formation at Loc. L NSM PM
15043.
Figs. 12a, b, 13a, b. Eriptyeha japonica KASE, sp. nov.
12a, b. Front and back views of the holotype (x4.0) from the Kimigahama Formation at
Loc. 2. NSM PM 15065.
13a, b. Back and front views of a para type (x 4.0) from the Kimigahama Formation at
Loc. 2. NSM PM 15066.

KASE and MAEDA:

Early Cretaceous Gastro1Joda !1'orn Choshi

Plate 34

716.

Early Cretaceous Gastropoda fro11t Choshi

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Tomoki KASE and Har1~yoshi MAEDA

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SOllL, N. F. (1967): Upper Cretaceous gastropods from the Pierre Shale at Red
Bird, Wyoming. U. S. Ceol. Surv., Prof.
Paper, 393-B, p. 1-46, pIs. I-II.
ST.'\:--:TO:,\, T. W. (1895): Contributions to theCretaceous paleontology of the Pacificcoast. The fauna of the Knoxville beds.
U. S. Ceol. Su rv., Bull., 133, p. 1-132, pIs.
1-20.
STE\\-ART, R. B. (1926): GABB'S California
fossil type gastropods. Proc. Acad. Nat.
Sci. Philadelphia, vol. 78, p. 287-447, pIs.
20-32.
STOLlC2KA, F. (1868): Cretaceous fauna ofsoutern India, vol. 2. The Gastropoda.
Palaeont. Indica, 497 pp., 28 pIs. Culcutta.
WE"2, W. (1938-1944): Gastropoda, Teil 1,
Allgemeiner Teil und Prosobranchie, in
SCHIl\DEWOLF, O. H., I-landbuch der Paliiozoologie, 7 vols., 1639 pp. Berlin.
YABI':, H., NAGAO, T. and SI"'1I2U, S. (1926) :Cretaceous Mollusca from the Sanchu
graben in the K wanto mountainland, Japan.
Sci. Rep. Tohoku Imp. Univ., 2nd ser.,vol. 2, no. 2, p. 33-79, pIs. 12-15.

Ashikajima-cho rftil.m!1bIllJ
Choshi ~n'"Kimigahama ;g- 7" r~
Kurohae-cho f!(l.1=.IIIj"
Toriakeura jl\i IIfJ !IIi

Inubo-zaki *11);J{,j
Nagasaki-cho *~I(,j-I1Ij"

Isejigaura ffr~n7"!lliNagasakihana *~~:..

Explanation of Plate 35
Figs. 1a-c, 2a--c. Vanikoropsis decussata (DESHA YES)
la-c. Front, apical and back views of a specimen (x 4.0) from the Kimigahama Formation
at Loc. 2. NSM Pl'vI 15063.
2a-c. Apical, front and back views of a specimen (x 4.0) from the Kimigahama Formation
at Loc. 2. NSM PM 15064.
Figs. 3a, b, 4a-c. I-layamia choshiensis KASE, gen. et sp. nov.
3a, b. Back and apical views of a paratype (x 1.5) from the Kimigahama Formation at Loc.
2. NSM PM 15041.
4a-c. Front, apical and back views of the holotype (x 1.5) from the Kimigahama Formation
at Loc. 2. NSM PM 15040.
Figs. 5a, b, 6a, b, 7a-c, 8a-c, 9a-c, 10. I-layamia rex KASE, gen. et sp. nov.
5a, b. Ventral and dorsal surfaces of an operculum (x 1.5) from the Kimigahama Formation
at Loc. 2. NSM PM 15039.
6a, b. Front and back views of a paratype (x 1.2) from the Kimigahama Formation at Loc.
2. NSM PM 15034.
7a-c. Front, back and apical views of a paratype (x 1.2) from the Kimigahama Formation
at Loc. 2. NSM PM 15035.
8a-c. Back, front and apical views of a paratype (x 1.2) from the Kimigahama Formation
at Loc. 2. NSM PM 15032.
9a-c. Front, back and apical views of the holotype (x 1.2) from the Kimigahama Formation
at Loc. 2. NSM PM 15031.
10. Operculum preserved nearly in situ in the aperture (x 1.5) from the Kimig1,lhama Formation at Loc. 2. NSM PlIl 15038.

KASE and MAEDA:

Early Cretaceous Gast1·opoda from Choshi

Plate 35

716.
----

323

Early Cretaceous Gastropoda !1'om Choshi--_ .. _ - - - - - - - - - - - - - - - - - - - - - - - - - - - - - --T-~!HMJt {-J1l!.)jrJ)jjlrWli'.ljlli*cJJ1Utm:-=r-3U~~E-n1l!.)j rJ) jjirWl [jilli*cl~ J: I) iii} I?;/e t-:(l~ Y-HikltdJ:f1>!i"iI)!/71ft:j-j ~ j;)Zii1 Lt.:*h.~~,10ffj[

fill~il;tJ'l1h1l1211nrJ)IJl.Ue.~Ht:r1(Cal/iostoma? ojii KASE, sp. nov., Ataphrus (s. str.)
nipponicus KASE, sp. nov., Hayalllia rex KASE, sp. nov., Hayamia choshiensis KASE,
sp. nov., Amberleya (Eucyclus) japonica KASE, sp. nov., Golitiea sp., Metriomphalus
nagasaliiensis IC\SE, sp. nov., Perissoptera elegans KASE, sp. nov., Pietteia cretacea
KASE, sp. nov., Ceratosiphon densestriatus IC'\sE, sp. nov.,
Vanilioropsis decussata(DESIL\ YES) and Eriptycha japoniea KASE, sp. no\'. ~~'llUJJj • jj~Y'E L t-:rJ)-c'i!c'rJi:"t" ~;/el?rJ)II'lcli, IJil!t!.JJiirJ)jf,j'if(5tJJil"tJ:, H:§"t"«@"l' <":)1J,rJ)tml);il;itL'n'~o Hayamialil!,0::li)[,[rJ) 7Hr~, ;kj(ri'.'Milli, ~PJJ~rJ) 7 !J ~#":) C C, to J: Vq)~J~O)II";k:ffl;vCr.II.I;z,.. ~j;ft-:t.el ').',i,-C'~ I?fC Hayalllia J~rJ) 7!J Ii Naticopsis kiirJ)k

Neritopsinaelll~f:frJ)ftUrJ)~~IJ,I?I~}jIJ~n,nl.:.~i(J;.("t"~o J;J.J:rJ)!J;:~Ii, Hayamia ~c Naticopsis h~rJ),j:jt,Mt.em~llkJ*IJ;II,.'i;iJ,~;Ie,
H ayalilia kIll); N eritopsis J,1l lib'U.e . :d-:ffi)ii]v.:.iict "') -C jlHt L t-: :: ~1I1'\'7~"t" ~ 0 Piet teia
cretacea li~;IJ~,c flllu~t.:7Pff'':'lIl3C"t" ~ JMt~f#":)!f!H'U.e -'(: ~ :; 'J 7' rJ)-ffI!JJii-c', ~'OIE*~Ii:; "" :7 *~ i':' rJ).J,.. J:fll?;Ie -C \" t-:IJ;, £1' 1"1 rJ)Wi5 ,.:. J: ,), ninDJ 1'1 illi,;j:~ ,.:. l(, ffit"t" ~ :: IJ; 11f] I? IJ' ,.:.
t.e "') t-:o Ceralosiphon }~Ii COSSIvIAKi': (1907)
Tessarolax N:ll,rJ) '/ / "'- j,
tiLt.e~;/L-C @" t-:IJ;, ,,1'l!1!,1J;5!I~j!.rJ) ~""C j; ~ :: ~ 5]; Lt.:o 1iH' fI'l'i.ftf!ii';'IJ1UtlWtrJ) 'I" ~.:., i, 7 '7 '/ ;<, ~-(=¥ ~ 7, rJ)/~ 1/ ~ 7'/, 7 -j' -J- 7'/ j; ~ H i 7 Iv ~ 7'/ rJ) l(, rJ) c J/;:jul. j; ~ Hii'l:~t.c:flfllJ;~
fl;'i;}t -Cl'~oJJIIi'lifi 1J:.Y.!.:· 'lii!T! 1Ij'f1~

c

c

c

J;)*

c

c

<

324

Tomoki J{ASE and Haruyoshi MAEDA

Explanation of Plate 36
Figs. 1, 2a, b, 3. Ceratosipholl densestriatus KASE, sp. nov.
1. Front view of the holotype (x 1.5) from the Kimigahama Formation at Loc. 2. NSM P:\I15057.
2a, b. Front view (x 1.5) and enlargement (x 4.5) of a paratype from the Kimigahama
Formation at Loc. 2. NSM PM 15058.
3. Back view of a para type (x 1.5) from the Kimigahama Formation at Loc. 2. ;'IISM P:\·I
15059.
Figs. 4-7, 8a, b, 9, 10. Pietteia cretacea KASE, sp. nov.
4. Back view of a paratype (x 1.5) from the Kimigahama Formation at Loc. 2. NSM PM
15053.
5. Front view of a paratype (x 1.5) from the Kimigahama Formation at Loc. 2. NSM PM
15052.
6. View of the outer lip spine of a paratype (x 1.5) from the Kimigahama Formation at
Loc. 2. NSM p~.! 15056.
7. Front view of a paratype (x 1.5) from the Kimigahama Formation at Loc. 2. ;'IISM P~'!
15054.
8a, b. Lateral and front views of a paratype (x 1.5) from the Kimigahama Formation at
Loc. 2. NSM PM 15051.
9. Front view of a paratype (x 1.5) from the Kimigahama Formation at Loc. 2. NSM Pl\!
15055.
10. Front view of the holotype (x 1.5) from the Kimigahama Formation at Loc. 2. NS:\!
PM 15050.
Figs. 11, 12a, b-15. Perissoj!tera elegans KASE, sp. nov.
11. Lateral view of a paratype (x4.0) from the Kimigahama Formation at Loc. 2. NSl\!
PM 15049.
12a, b. Front and back views of a paratype (x 1.2) from the Kimigahama Formation at
Loc. 2. NSM PM 15048.
13. Back view of a paratype (x 1.2) from the Kimigahama Formation at Loc. 2. NSM Pl\'1
15047.
14. Front view of the holotype (x 1.2) from the Kimigahama Formation at Loc. 2. NSM
PM 15045.
15. Back view of a paratype (x 1.2) from the Kimigahama Formation at Loc. 2. NSM Pl\!
15046.

KASE and MAEDA:

Early Cretaceous Gastropoda from Choshi

Plate 36

Trans. Proc. Palaeont. Soc. japan, N. S., No. 118, p. 325-338, pI. 37, june 30, 1980

717. CEPHALOPOD FAUNULE FROM THE CRETACEOUS YATSUSHIRO FORMATION (KYUSHU) AND ITS IMPLICATIONS*TATSURO MATSUMOTO
Seinan·gakuin University, Fukuoka 814

KAMETOSHI KANMERA
Dept. of Geology, Kyushu University 33, Fukuoka 812
and

YOSHIHISA OHTA
Dept. of Earth·Sci., Fukuoka Univ. of Education, Fukuoka 811-41

Abstract. This is a report on the cephalopod faunule from the Cretaceous
Yatsushiro Formation in southwestern Kyushu. It consists of 5 species from three
localities, which are assigned to the middle part of the Yatsushiro Formation. One
is a new species of Heminautilus, a peculiar nautiloid which deviates much inshell·form from H. saxbii (MORRIS), being apparently similar to certain species of
Paracenoceras. Others are ammonites, Brewericeras n. sp., allied to B. hlilenense(ANDERSOI") from the Pacilic Coast of North America, Prolyell ice ras (?) sp., E piley.
meriella aft. hitzeli (JACOB) and Plaliknemiceras n. sp., allied to P. gracile (Dot;·
ViLLE). Palaeontological descriptions are given by the senior author. The faunule
is interesting in that both the North Pacific and the Tethys Sea elements intermingled. The ammonites indicate the Lower Albian. As the Yatsushiro Formation contains land flora and some non-marine to brackish water molluscs in its lower andupper parts, the cephalopod faunule gives a key to correlate the fossils of different
facies.

mountainous area to the south of Yatsushiro, Kumamoto Prefecture. Kyushu.
One of them is the Albian Yatsushiro
Formation, from which several cephalopod
fossils were obtained. This paper is toreport a result of our study on them.
For the palaeontological descriptions oneof us (T. M.) alone is responsible.

Introduction
As has already been described (MATSUMOTO, ed., 1954 ; MATSUMOTO and KANMERA, 1964), folded and thrust-faulted
Cretaceous formations are exposed in the* Received February 18, 198J; read june 10,
1979 at Tatsunokuchi.

325

32G

Tatsnro MATSUJI.IOTO. Kametoshi KANMERA and Yoshihisa OHTA
~1

MiyajiCJ:J 2
~3

•

4

1<

5
6

)<

Fig. 1. Geological route map and the stratigraphic profile between Miyaji and
Joguzan (prepared by Y. OIIT1\). 1: shale, 2: sandstone, 3: conglomerate, 4:
locality of ammonites (Brewericerc.s n. sp. and Prolyelliceras sp.), 5: marine in·vertebrate fossils, 6: brackish water fossils. Inset is an outline map of Kyushu,
indicating the location of the Yatsushiro Formation.
Bisho

2km
I

2
~~

~~

3

4

5

6

m [ill]J]] illIIIJJ WRmtFig. 2. Geological map of the Yatsushiro Formation and the surrounding rock units.
(simplified from MATsnioTo and KA:'OIERA, 1964). 1: Miyaji Formation, 2: Yatsushiro
Formation, 3: Hinagu Formation, 4: Hachiryuzan Formation, 5: Kawaguchi Formation(altogether Lower Cretaceous), 6: igneous, metamorphic and Palaeozoic rocks (undifferentiated), A: area of Fig. 1, B: area of Fig. 3, . : locality of fossil nautiloid.

717. Cretaceous Cephalopods 11'0111. Yatsu,shiro
Geological Setting
As is clearly shown in the publishedgeological map (MATSUMOTO and KAJ';'MER A, 1964) the Yatsushiro Formation is
exposed in two tectonic belts, the Hinagu
belt in the south and the Miyaji belt in
the north, which are separated by a thrustzone of serpentinite (Fig. 2). In the
Hinagu belt, the Yatsushiro Formation
rests on the Aptian Hinagu Formation
with a distinct unconformity, although
the relation may be modified by later
faulting in some places. The strata of
the latter form a partly overturned syncline, whereas those of the former are ina normal syncline. In the Miyaji belt,
the Yatsushiro Formation is disconformably overlain by the Miyaji Formation;
there the strata incline northward withrather low angled undulations.
The Yatsushiro Formation itself isabout 580 m in thickness and consists ofa cyclic sequence of clastic rocks. It is
subdivisible into three members, theLower, Middle and Upper. In the Lower
and Upper Members, conglomerate andcoarse-grained sandstone predominate,with red beds, subordinate shale, carbonaceous seams and plant beds. In the
Middle Member there are many intercalations of sandy shale and certain parts ofthe shale and nearby sandstone containshallow marine (and partly brackish water)
invertebrate fossils (molluscs, echinoids,
corals etc.), in which ammonites and a
nautiloid were rarely found.
In the Hinagu belt the lower threefifths of the Yatsushiro Formation are
exposed, whereas in the Miyaji belt theupper half of the formation is exposed.
Therefore the fossiliferous beds of the
Middle Member are accessible in both
belts, although beds of sandy shale arethinner in the northern belt. In Fig. 1,
ammonite and some relevant fossil locali-

327

ties of the northern (i. e. Miyaji) belt isshown. Two other ammonites were obtained from a large block certainly fallenfrom the nearby exposure of the lower
part of the Middle Member in the southern (i. e. Hinagu) belt at a locality indicated in Fig. 3. A nautiloid locality is
about 6 km east of this ammonite locality
within the same Hinagu belt (Fig. 2) and

o
I

300m
J

Fig. 3. Geological route map, north of
Shimo-fukami (area B in Fig. 2) (preparedby K. KA;-';~IERA), showing the location ofammonite bearing large block (fall) by+(loc. Km 1843). Formation names.-Ya:
Yatsushiro, Hi: Hinagu, Ha: Hachiryuzan,
Ka: Kawaguchi. Lithological symbols sameas in Fig. 1.

328'l'atsU1"O MATSUMOTO, Kametoshi KANMERA and Yoshihisa OHTA

PI. 37, Fig. 1; Text-fig. 4

Dec. 1956, when he was a student of
Kyushu University. This is the holotype.
Description :-The holotype is small,about 13.5 mm in diameter, but a part ofthe body-chamber (about 90°) is preserved. It is involute, showing a verysmall umbilicus (less than 1 mm).
Its shell-form is characteristic. The
whorl in the late stage is subtrapezoidal
in cross-section, having a rather flat venter, angular ventrolateral shoulders, generally convergent flanks, abruptly roundedumbilical shoulders and steeply inclined
umbilical walls. The body-chamber isnearly as high as broad and broadest immediately above the umbilical shoulder.
On the body-chamber alone there is ashallow median groove and the upper partof the flank is slightly concave below the
keel-like ventrolateral shoulder, whereas
the lower part is inflated.
In earlier growth-stage this characteristic shell form is not well displayed,
namely the venter is more rounded, theventrolateral shoulder is subrounded, theflanks are more gently rounded, converging towards the venter and the umbilicalshoulders are gently rounded.
The surface of the internal mould issmooth. The outer surface of the shell,
as impressed on the external mould, has
numerous, dense, fine radial lirae and veryfine longitudinal lirae, showing a finely
reticulate ornamentation. The radiallirae
are flexuous and rursiradiate on the ventral part. They are sometimes bifurcate
or virgate on the upper part of the flankand also near the umbilical shoulder.
The septal suture in the late growthstage has a well developed, broad but
moderately deep lateral lobe, which passes
to an asymmetric saddle on the ventro-

Material :-GK. H6909A, B, which consists of the internal (A) and external (B)moulds, obtained by Takeshi AKATSU inlateral shoulder and then to a very shallow
lobe on the venter. At about the umbilicalshoulder there is a small but distinct
lateral saddle which passes to a concave

assignable to the lower part of the Middle
Member. The Yatsushiro Formation nearthese two localities are rather narrowly
and incompletely exposed, owing to thefaulting on both wings of the syncline.
The cephalopod specimens described below
are now kept in the Type Collections,
Kyushu University (GK).
Palaeontological Descriptions(by Tatsuro MATSl:~JOTO)Class Cephalopoda
Subclass Nautiloidea
Order Nautilida
Familly Nautilidae V'ORBIGNY
Genus Heminautilus SPATH, 1927
Type species :-Nautilus saxbii MOl{!{IS,1848.
Renwrks :-1 depend primarily on KUMMEL (1956, p. 434) for the redefinition ofthis genus. There is, however, a questional point as to the relationship of
Heminautilus with Paracenoceras SPATH,
1927 (type-species Nautilus hexagonus J.
DE C. SOWERBY, 1826), as has alreadybeen discussed by SPATH (1924, p. 25) andconversely by KUMMEL (1956, p.435). A
single small specimen from the Yatsushiro
Formation shows interesting features
which may be involved in the same question.
JIemill(Jutilus akatsui MATSUMOTO,
sp. nov.

717.

329

Cretaceous Cephalopods from Yatsushiroline on the umbilical wall. The sutureof the immature stage has no ventral lobe,
a shallower lateral lobe and a lower saddle at the umbilical shoulder. It is generally descending.
Position of the siphuncle is unknown.
as I do not dare to cut this single specimen.
Occurrence :-In a silty very fine-sandstone exposed on the west side of a road,about 2 km south of Bisho, Toyo-mura.
Yatsushiro-gun, Kumamoto Prefecture,
together with some trigonians and othermolluscs.
Comparison and discussion:- With respect to the shell-form this species is rather
similar to certain species of Paracenoceras
SPATH, 1927, such as P. hexagonum(SOWER BY) (see Fig. 5) and especially P.
prohexagonum SPATH, 1935. Those twoand many other species of Paracenoceras
are mostly Middle or Upper Jurassic in age.
The exception may be Nautilus rhodani
Roux (in PICTET and Roux, 1847, p.275,
pI. 1, fig. 4) from the Albian of Switzerland, which was referred to Paraceno-

ceras by SPATH (1927, p. 25) and KUMMEL(1956, p. 403). As its holotype is an internal mould, the ornamentation of its
shell surface is not known. If there were

(
..
,,

.

"~.
:•
I .

Paracenoceras hexagonum
Diagrammatic cross section and suture (after KUl\1yIEL, 1956,
figs. 18 and 20B). Bar indicates 10 mm.
Fig. 5.

(SO\\,ER!3Y) •E'I·t.sh(U,S'.

Fig. 4. Heminautilus akatsui MATSt:'sp. nov. Diagrammatic sketch ofholotype, showing frontal and lateral views
and the last suture. Bar indicates 1 mm.
MOTO,

Fig. 6. Heminautilus saxbii (MORRIS).
Diagrammatic two views of holotype represented by KU~IMEL, 1956, pI. 10, figs. 1, 2.
Bar indicates 10 mm (cited by permission).

330

Tatsll1'o MATSUMOTO, Kametoshi KANMERA and Yoshihisa OHTA

radial ribs on the outer shell, it could bereferable to Paracymatoceras SPATH, 1927
(type-species N. asper (OPPEL) ZITTEL,
1868) and would be allied to P. tllnghaicllm
MATSUMOTO and AMANO, 1964, from theLower Cenomanian (possibly uppermost
Albian) of Kyushu. Anyhow, it is distinguished from the present species by its
broader whorl, broader venter, and shallower lateral lobe without perceptible
saddle at the umbilical shoulder.
The suture of Paracenoceras is, in most
species, less strongly sinuous than that ofHeminautilus, but some species, e. g. P.
rollieri (LOESCH, 1914) (see JEANNET, 1951,
fig. 29), has a moderately sinuous suture,
which is somewhat similar to that of thepresent species. However, it cannot be
overlooked that the suture of the present
species in the late growth stage resembles
that of Heminautilus saxbii (MORRIS) (seeKUMMEL, 1956, pI. 10, figs.!, 2) (also Fig.
6 of this paper), despite the difference inthe degree of compression of the shell.
H. saxbii and other species of Heminautilus normally have a compressed shell,
with high whorls and a flat or slightly
concave, narrow venter. Therefore thepresent species much deviates from themin the shell-form, although the flanks areconvergent as in those species of thatgenus.
Heminautilus was provisionally placedin the subfamily Paracenoceratinae SPATH,
1927 by SPATH (1927) with a query, but
has been replaced to the subfamily Cymatoceratinae SPATH, 1927 by KUMMEL(1956), because the latter author recognizes the flexiradiate ribs in Heminalltilus
like those of Cymatoceras. In the present
species, as clearly seen on the externalmould of the holotype, there are fine radiallirae, which are rursiradiate on the outerpart of the flank and show a sinus on theventer. The so-called ribs in Cymatoceras
are not impressed on the internal mould

and, therefore, should not be comparedwith the ribs of the Ammonoidea. They
are better called thick lirae, if the terminology in the Ammonoidea is applied. In
fact, I see on some undescribed examplesof Cymatoceras, from the Upper Cretaceous of Hokkaido and South Saghalien,
very fine lirae on the surface of the immature shell, which are developed into
the rib-like thick lirae on that of themature shell. That fine Iirae on the immature shell show a finely reticulate
ornament by the combination with veryfine longitudinal lirae or striae. This
ornament is essentially the same as thatseen on the external mould of the holotypeof the present species. On the other hand,
Paracenoceras is smooth or has only
growth-lines on the shell surface.
To sum up, I regard the present species
as an atypical example of Heminautilus,
despite its similarity to Paracenoceras inshell-form. It is still questionable whether
the small holotype is immature or mature.
For the reasons of its small size and finely
reticulate ornament, I would presume it
as being immature, but further material
is needed to confirm the conclusion.
Subclass Ammonoidea
Order Ammonitida
Family Desmoceratidae ZITTELGenus Brewericeras CASEY, 1954
Type species :-Ammonites breweri GABB,
1869.
Remarks:- This genus was defined by
CASEY (1954, p. 112), who assigned it to
the Desmoceratidae. On the basis ofnumerous specimens, JONES et al. (1965)
reexamined this genus and Leconteites
CASEY, 1954 (type-species Desmoceras
lecontei ANDERSON, 1902) and suggested
its possible derivation from the latter.

717.

Cretaceous Cephalopods j1'om Yatsushi1'oThey referred the two genera to the
Beudanticeratinae of the Desmoceratidae,
although CASEY assigned Leconteites to
the Cleoniceratinae of the Hoplitidae.
Brewericeras has been known characteristically from the upper part of Lower
Albian in the Pacific Coast of North
America. The species described below is
the first record of this genus from japan.
It has some features which seem to support the opinion of JONES et al.
Brewericeras enorme MATSUMOTO,

sp. nov.
PI. 37, Fig. 2

klaterial :-A single specimen, GK.
H6907, the holotype, found by Y. OHTA,
who donated it to Kyushu University
through T. MATSUMOTO.
Description :-The holotype is secondarily compressed and the original breadthof its whorl is difficult to estimate. The
measurements (in mm) on the unrestored
specimen are as follows: Diameter=102.0,
whorl-height=50.0, Umbilicus=18.8 (0.18).
Presumably, the whorl was originallyhigher than broad, with a narrowly arched
venter; the sharpened venter of the fossil
is probably due to the secondary compression.
The umbilicus is fairly narrow, surrounded by a low but vertical wall andangular umbilical shoulder. The flank is
flattened, especially on its inner half.
There are numerous, fairly crowded,
falcate or sinuous ribs on the flank. In
the last half whorl preserved, the ribs
are mostly simple and distinct only on
the ventral half of the flank; there areoccasionally shorter ribs which are intercalated or branched on the ventrolateralpart; on the inner half of the flank, weaklirae and faint extensions of a few ribs
are discernible, which are prorsiradiate331

and nearly straight or sometimes gently
curved. On the venter ribs are much
weakened.
On the flank of the preceding half
whorl several longer ribs arise from theumbilical shoulder and are flexiradiate on
the main part of the flank, with bifurcalation or intercalation of shorter ribs. On
the interval between the longer ribs there
are several flexuous ribs of unequal length,
which appear at various distances fromthe umbilical margin. The ribs are, therefore, fairly crowded on the outer half ofthe flank, numbering about 25 within ahalf whorl. They are much weakened on
the venter.
The ornamentation of the next inner
whorl is not well shown, but no strong
ribs are perceptible on the inner half ofthe flank.
The suture is only partly seen at about
the middle of the last whorl preserved;
it shows typically desmoceratid pattern.
Occurrence :-The holotype was foundat a locality indicated as C in Fig. 1, at
the altitude of about 200 m on an ascending path from Miyaji to Mt. joguzan. It
was in a dark grey shale which is referred to the upper part of the Middle
Member of the Yatsushiro Formation.
Comparison:- This species resembles
the ribbed variety of Brewel'iceras Illtlenense (ANDERSON), as redefined by McLEARN (1972), from the upper part of theLower Albian of the Pacific Coast of
North America, between California andAlaska (ANDERSON, 1938; JONES et aI.,
1965; McLEARN, 1972; IMLA Y, 1960;
JONES, 1967), in its mode of ribbing of thelast part as well as in its shell-form, but
is distinct from that species in its characteristic ribbing of the middle growthstage.
The last mentioned character reminds
us of the ribbing of Leconteites lecontei
whiteavesi JONES, MURPHY and PACKARD

332

1'atIi1l1'o MATSUMOTO, Kametoshi KANMERA and Yoshihisa OHTA

(1965, p. F13, pI. 6, figs. 1-3, 4-9, 12-14).
In that subspecies, as well as in typicalL. lecontei. the intercalation and branchingof the ribs persist to the late stage andthe long ribs normally have bullae or
tubercles at the umbilical shoulder; ribs
are therefore, more numerous than in thepresent species, although there is variation in the rib density. The forms whichwere regarded as intermediate between
B. hulenense and L. lecontei (JONES et aI.,
1965, pI. 11, figs. 7-12,15-17) are not identical with the present holotype, in thatsome of the ribs in those forms haveweak umbilical tubercles or bullae andalso in that the irregularity of the ribbing by branching or intercalation and byintensity difference is manifested in thelimited period of the growth-stage in our
specimen.
To sum up, I would regard the described specimen from the Yatsushiro
Formation as representing an atypicalnew species of Brewericeras.
Family Lyelliceratidae SPATH
Genus Prolyelliceras SPATH, 1930
Type species :-P. peruvianum SPATH,
1930.
Prolyelliceras(?) sp.
PI. 37, Fig. 3

Material :-GK. H6908, an imperfect
external mould of the right side of anammonite.
Description :-This specimen has, on its
outer whorl, fairly coarse radial ribs
which start from the umbilical marginand have tubercles at the umbilical andthe ventrolateral shoulders. There areregularly intercalated, weaker and shorter,
secondary ribs. On the inner whorl, theribs are finer, more numerous and more
crowded, with longer secondaries, andhave less prominent umbilical bullae andsmall ventrolateral tubercles. The ribs
are nearly rectiradiate in the late growthstage but gently sinuous in the earlier
stage. No sign of lateral tubercles isperceptible.
Occurrence:- This specimen was obtained by Y. OHTA at a locality indicatedas B in Fig. 1, in the fossiliferous sandstone stratigraphically about 40 m below
the Brewericeras-bearing shale. Nanonavis
yokoyamai (Y ABE and NAGAO), Pterinella
shinoharai HAY AMI, Neithea matsumotoi
HAY AMI, Astarte subsenecta YABE andNAGAO, Nemocardium yatsushiroense HAy AMI, trigonians etc. occur there.
Comparison :-Because of the imperfectpreservation, this ammonite is hardly
identified with precision. As the venter
is not preserved, and accordingly thetrituberculate character is not confirmed,
the specimen is provisionally referred to
the genus Prolyelliceras SPATH, 1930, witha query. The hitherto described speciesof Prolyelliceras, e. g. P. peruvianum
SPATH, 1930 (see BENAVIDES-CAcERES,
1956; RIEDEL, 1937), from the Albian ofPeru and Colombia, and P. radenaci(PERVINQUIERE, 1907), from the Lower
Albian of Tunisia and also Peru (seeSOMMERMEIER, 1910), have no secondary
ribs. Lyelliceras cotteri SPATH, 1930,
from the Albian of Hazara, has secondary
ribs but also lateral tubercles.
Genus Epileymeriella BREISTROFFER, 1947
Type-species: - Parahoplites
hitzeli
JACOB, 1908.
Remarks :-1 follow CASEY (1957, p.31)
in assigning this genus to the subfamily
Lymeriellinae BREISTROFFER, 1951 of the
Lyellicera tidae.

717. Cretaceous Cephalopods from Yatsushiro
Epileymeriella sp. aff. E. hitzeli(JACOB)
PI. 37, Fig. 4Compare :1908. Parahoplites hitzeli JACOB, Mem. Soc.
Ceoi. France, Paleont., vol. 15, no. 38,
p. 48, pI. 8, figs. 1-3.
1954. Leymeriella (Epileymeriella) sp. d.
hitzeli (JACOB), CASEY, Paiaeontoi.,vol. 1, p. 57, pI. 9, figs. 5, 5a.

Materia} :-GK. H6906 A, B, a small
fragmentary whorl, represented by aninternal mould (A) and also an externalmould (B).
Description:- The whorl is higher thanbroad, with height=12.0 mm and breadth=7.5 mm. The suture is indistinct.
On the flank there are numerous ribs
which are curved forward on the ventrolateral part. Normally two ribs are
branched from an umbilical bulla andthere are, at least partly, still more branching or doubling on the ventrolateral part,as seen on the external mould. Occasionally, there is a simple rib which is accompanied by a constriction-like deeper
interspace. Along the siphonal line there
is a groove which interrupts the ribs, with
their peripheral ends being disposed alternately.
Occurrence :-This specimen was obtained by K. KANMERA at loco Km 1843,
north of Shimo-fukami, Sakamoto-mura,
Yatsushiro-gun, Kumamoto Prefecture.
It came from the silty fine-sandstone of
the Middle Member of the Yatsushiro
Formation in the Hinagu belt (see Figs.
2, 3).
Comparison :-This ammonite was listed
as Hoplites d. dentatus (SOWERBY)by
MATSUMOTO and KANMERA (1964), but
this was a misidentification, because thatspecies should be somewhat more inflated
and have stronger umbilical tubercles.

333

Dr. R. I. KIRBY (personal communication, August 1979) told me that this specimen apparently resembles some specimens (e. g. Univ. Oxford Geol. K5615)
which he provisionally calls Analwplites(Daghestanites) d. daghestanensis GLAZUNOVA, 1953 and Dr. J. A. JELETZKY (personal comm., Sept. 1976) suggested methat it looks similar to "Anahoplites"
haidaquensis (WHITEA VES). The latter is,
however, the type-species of Pseudo leymeriella CASEY, 1957, which is a trochleiceratid (see WIEDMANN, 1966). However, in view of the other ammonites
associated with this specimen, it is more
likely that the specimen is a species of
Lymeriellinae, as Dr. J. W. KENNEDY (personal comm., Sept. 1976) suggested me.
lt indeed resembles Epileymeriella hitzeli(JACOB), from the Lower Albian of France,
but has a groove instead of a smooth zone
on the middle of the venter.
Family Engonoceratidae HYATTGenus Platiknemiceras BATALLER, 1954
Type-species :-Knemiceras (Platiknemiceras) bassei BATALLER, 1954.
Remarks :-Platiknemiceras was clearly
redefined by CASEY (1961), who ranked it
up to an independent genus and listed anumber of species in addition to P. bassei.
They occur characteristically in the Albianand distribute in the Tethys region andits extensions in the Americas (fromTexas to Peru). A species from Kyushu(Japan), described below, gives an addition to the distributional area of the genus.
Platiknemiceras caseyi MATSUMOTO,
sp. nov.
PI. 37, Fig. 5

Material :-A small specimen, GK.
H6905, holotype, represented by an in-334

Tats~[1'O MATSUMOTO, Kametoshi KANMERA and Yoshihisa OHTA

ternal mould and also a part of the external mould. This species is dedicatedto Dr. Raymond CASEY.
Description:- The holotype is about 27
mm in diameter, flatly discoidal, involuteand very narrowly umbilicate. Its narrow
venter is bicarinate.
There are sinuous, very low ribs on theflank, but they are much weakened andalmost obsolete on the inner half. On thepreserved last half whorl the ribs arerather rursiradiate on the middle part of
the flank and then gently curved forward
on the outer part. No tubercles aredeveloped along the umbilical margin andin the middle of the flank, but the peripheral ends of the outer ribs from a trainof clavi on the ventral keel.
The suture is incompletely exposed atwhorl-height of 5.5 mm, a little more than
one volution behind the preserved end of
the shell. It consists of less numerous
elements than in the suture of Engonoceras and the saddles are not so entire
as in that genus.
Occurrence :-The holotype was obtainedby K. KANMERA (in 1948) at loco Km
1843b from a huge block of fine-sandy
siltstone derived from the nearby exposureof the Middle Member of the Yatsushiro
Formation. This locality is close to Km
1843 where Epileymeriella aff. hitzeli wasfound (see Figs. 2, 3).
Comparison:- This fossil was listed
previously (MATSUMOTO ed., 1954; MATSUMOTO and KANMERA, 1964) as Engonoceras aff. stolleyi (BOHM, 1898) on account of the resemblance in the muchcompressed shell-form and mode of ribbing on young whorls (see HYATT, 1903,
pI. 23, figs. 7-8). However, it should not
be referred to that genus, because its
suture is not of the Engonoceras pattern.
The present species rather resembles
Platiknemiceras gracile (H. DOUVILLE,
1916), from the Lower Albian of Egypt

and adjacent areas. Its ventral keels arecrenulated as in that species. Although
the shell of P. gracile is described as
being smooth, the figure in DOUVILLE(1916, pI. 16, fig. 9) seems to show weakribs on the outer part of the flank in the
middle growth-stage. There are, however,difference in the curvature of the ribs andthe details of the suture. The suture of
the present specimen resembles that of
Platiknemiceras sp. as figured by CASEY(1961, p. 354, text-fig. Ie), although its
elements are less numerous than in thatspecies probably because of its immature
stage.
To sum up, the present specimen represents a new species of Platiknemiceras
which is characterized by sinuous low
ribs on the outer part of the flank.
Implications of the Faunule

Despite the incomplete preservation andpaucity of the fossils, the cephalopods
described provide valuable information,
as follows: (1) Geological age :-Brewericeras hulenense is an index of the Zone
of B. hulenense in the Albian sequencesof the Pacific Coast of North America.
In the Haida Formation of British Columbia this species is associated with Douvilleiceras spinigerum (WHITEAVES) and weagree with McLEARN (1972) and also with
JONES et al. (1965) in referring the Zone
of B. hulenense to the upper part of theLower Albian in terms of the international
scale. The close affinity of the describednew species of Brewericeras with B.
hulenense and also the fact that it has
some characters which remind us of theancestral Leconteites strongly suggest the
middle or upper part of the Lower Albian
for the Middle Member of the Yatsushiro
Formation.
Prolyelliceras is considered to range
from the Lower Albian to Middle Albian,717. Cretaceous Cephalopods from Yatsushiro
although no representative is known from
northwest Europe. Epileymeriella hitzeli,
to which our specimen is allied, occurs in
the Zone of Leymeriella tarde/urcata,which indicates the lower Lower Albianin France and England.
Most of the hitherto known species of
Platiknemiceras occur in the Lower Albian,as explained by CASEY (1961). Our species, although new, would have no contradiction against the view to regard it
as a Lower Albian ammonite.
Heminautilus is rather common in theLower Cretaceous, from Barremian toAlbian. Although the species describedin this paper has some peculiar features,
there is no discrepancy in regarding theage of its host rock as Lower Albian.
Summarizing the above, the age of the
cephalopod bearing Middle Member of the
Yatsushiro Formation is concluded as
Lower Albian, probably the middle or
upper part of the Lower Albian. Consequently, the allocation of the Yatsushiro
Formation in the correlation chart of
OBATA and MATSUMOTO (1976, fig. 2)
should be removed slightly downward toaccomodate its middle part (indicated bya solid circle) within the scale of K3bl.
(2) Palaeobiogeography:-Brewericeras
has been known to typify the Albian
northeastern Pacific province, from Alaska
to Carifornia. Its occurrence in Kyushu(Southwest Japan), though represented bya rare species, should be noted for its
expanded distribution around the North
Pacific Ocean.
Knemiceras and Platiknemiceras arecharacteristic of the Tethys Realm (including the extensions to the Americas),being distributed widely in these tropicalto subtropical equatorial seas (e. g. CASEY,
1961, fig. 2). Although very poorly represented, the occurrence of an exampleof this Tethyan element in Kyushu isnoteworthy. The coexistence of Platik-

335

nemiceras n. sp. (aff. P. gracile) with
Brewericeras n. sp. (aff. B. hulenense),
together with Prolyelliceras(?) sp., Epileymeriella aff. hitzeli and Heminautilus n.
sp., in the same Middle Member of the
Yatsushiro Formation, exemplifies an interesting case of a lucky encounter ofdifferent faunal elements in the Japaneseprovince. This would provide a key forthe inter-regional correlation between
different provinces.
(3) Significance 0/ cephalopod bearing
interbeds :-For some geological reasons
Kyushu is endowed with favourable conditions in which marine beds are interfingered with non-marine ones. The
Yatsushiro Formation is one of such
examples. Plant beds occur in the Lower
Member and also in the Upper Member,brackish-water or non-marine mollusca in
the lower part of the Middle Member (asindicated by loco A in Fig. 1) as well as
in other members. These non-marine
fossils are referable to the Albian (probably lower half of the Albian) on accountof the cephalopod species from the mainpart of the Middle Member.
The fossil flora of the Yatsushiro Formation (mainly Lower Member) was
studied by KIMURA (1978), who has shownthat the Yatsushiro flora contains noangiosperms and that it generally resembles the Lower Cretaceous Ryoseki florain the Outer Zone of Southwest Japan,
but has certain species which are particular to it. Dr. KIMURA (personal comm.,
May 1979), furthermore, tells to one ofus (T. M.) that the Yatsushiro and theso-called Ryoseki floras belong to the
Wealden flora of the northern hemispheredistributed on the north side of the Early
Cretaceous equatorial belt and that they
are characterized by xenophytes of "aridedaphic condition". The plant fossils weretransported from a land of such a condition to the Lower Cretaceous basins (one

336

Tatsuro MATSUMOTO, Kametoshi KANMERA and Yoshihisa OHTA.

of which was the Yatsushiro basin) andit should be noted that the Yatsushiro
Formation contains red beds in its lower
part. This may be generally in harmony
with the presence of some Tethyan elements in the marine fauna. Marine bivalves which occur in the Middle Member
were described by HAY AMI (1965-66),
together with those from other Lower
Cretaceous formations of Japan, and echinoids by TANAKA and OKUBO (1954).
Colonial corals were identified by M.
EGUCHI (cited in MATSUMOTO ed., 1954,
p. 114). The environmental conditions
during the time of the Yatsushiro Formation should be interpreted from the comprehensive study of these fossils as wellas the sediments.
Acknowledgements: - We thank Mr.
Takeshi AKATSU who provided us a nautiloid fossil which he obtained during his
field work, Drs. W. J. KENNEDY, R. I.
KIRBY, J. A. JELETZKY and Tatsuaki
KIMURA, who have given to one of us(T. M.) valuable information from their
specialities, Drs. K. TANABE, I. OBATA
and I. HAY AMI, Miss Mutsuko HAY ASH!DA and Miss Kazuko HARA, who helped
us in preparing the paper. Dr. KIRBY
kindly read the first draft, when one ofus (T. M.) visited England. This is acontribution to the "International Correlation of the Cretaceous System",supported by the Science Research Fund
(No. 334043) of lvIonbusho.

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Totsuro lVIATSUMOTO, Kametoshi KANMERA and Yoshihisa OHTA

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Hachiryuzan Ai1!!11
Hinagu B :;R7\,
Joguzan
Nekodani ~Iii:fr
Sakamoto:lN*
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Explanation of Plate 37
Fig. 1. fiell1inautiius akatsui MATSUNIOTO, sp. nov ................................... Page 328
Ilolotype, GK. H 6909A, B. Two lateral (a, b) and ventral (c) views of internal mould (A)and surface ornamentation on replica (d) from external mould (B).
Fig. 2. Brewericeras enorme MATSl:ilIOTO, sp. nov ................................... Page 331
Holotype, GK. H6907. Lateral view.
Fig. 3. Proiyelliceras (?) sp ......................................................... Page 332
Replica from GK. H6908. Lateral view.
Fig. 4. Epileymeriella sp. aff. E. hitzeli (JACOI3) .................................... Page 333
GK. H6906A (internal mould). Lateral (a) and ventral (b) views.
Fig. 5. Plaliklle1lliceras caseyi MATSU~.JOTO, sp. nov ................................. Page 333
Holotype, GK. H6905A (internal mould). Lateral view of the preserved whole part (a) andthat of a younger part (b); external suture of a young stage (at whorl-height=6 mm) : twoparallel wavy lines at the top are keels of a compressed venter, where suture is unexposed.
Bar indicates 10 mm in almost every figure, except 1 mm in Fig. 5c. Kyushu University photos(K. 'L\~i\BE), without whitening.

MATSUMOTO et al. :Plate 37

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

June 30, 1980

New Series No. 118

CONTENTS
TRANSACTIONS
715. SAKAGAMI, Sumio: Permian Ectoprocta (Bryozoa) from the Abadeh Region,
Central Iran .......................................................... 269
716. KASE, Tomoki and MAEDA, Haruyoshi: Early Cretaceous Gastropoda fromthe Choshi District, Chiba Prefecture, Central Japan .................. 291
717.

MATSUMOTO, Tatsuro, KANMERA, Kametoshi and OTA, Yoshihisa: Cephalopod Faunule from the Cretaceous Yatsushiro Formation (Kyushu) and
its Implications ...................................................... " 325