ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 149

Pa laeonto logica l Society of Japan
April 30, 1988

Co-Editors

Hisayoshi IGO and Tsunemasa SAITO

Editorial Board: Yoshikazu HASEGAWA, Itaru HAYAMI, Hisayoshi IGO (chief), Makoto KATO,
Itaru KOIZUMI, Yasumochi MATOBA, Kei MORI, Hiroshi NODA, Kazunari TANABE,
Tsunemasa SAITO, Kazuhiko UEMURA, Akira YAO
Officers for 1987-1988
President: Tatsuaki KIMURA
Honorary President: Teiichi KOBA YASH I
Councillors:

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Hisayoshi IGO, Junji ITOIGAWA, Tadao KAMEl, Tatsuaki KIMURA, Tamio KOTAKA,Kei MORI, Ikuwo OBATA, Tsunemasa SAITO, Yokichi TAKAYANAGI, Kazushige
TANABE, Toshimasa TANAIMembers of Standing Committee: Kiyotaka CHINZEI (Membership), Takashi HAMADA (Foreign Affairs), ltarn HAYAMI (Finance), Hisayoshi IGO (Co-Editor of Transactions),
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Secretaries:

Katsumi ABE (planning), Takeshi ISHIBASHI (Special Paper), Kunihiro ISHIZAKI("Fossils"), Tomoki KASE (General Affairs and Membership), Hiroshi NODA (Editorof Transactions), Ternfumi OHNO (Membership), Tatsuo On (Planning), Kazuhiko
UEMURA (General Affairs), Toshiyuki YAMAGUCHI (Finance)
Audi tor: Noriyuki IKEY A

The fossil on the cover is Vicarya yokoyamai Takeyama, an Early Middle
Miocene gastropod from the Kurosedani Formation at Kakehata, Yatsuo-cho,
Nei-gun, Toyama Prefecture, central Japan (Collected by T. Kotaka and K.
Ogasawara, IGPS No. 99075, photo by S. Ohtomo and Y. Kikuchi, xO.9).

All communication relating to this journal should be addressed to the

PALAEONTOLOGICAL SOCIETY OF JAPAN% Business Center for Academic Societies,
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 149, pp. 361-370, 5 Figs., April 30, 1988

851. EARLY CARBONIFEROUS (VISEAN) GASTROPODSFROM THE HIKOROICHI FORMATION OF THE KITAKAMIMOUNTAINS, NORTHEAST JAPAN*TOMOKI KASE
Department of Geology, National Science Museum, Tokyo, 160

Abstract. Six Early Carboniferous (Visean) gastropods are described from a sandstone bedof the Hikoroichi Formation exposed in the Onimaru Quarry of Ofuna to City, the Kitakami
Mountains, northeast Japan. This is the first report of Carboniferous gastropods from the
Kitakami Mountains. Kawanamia is proposed as a new genus of the family Lophospiridae. The
species described herein are Straparollus (Euomphalus) asanoi, n. sp., S. (E.) sp., Kawanamia
onimarensis, n. sp., Baylea yvanii (Leveille), Littorinides sp., and Pseudozygopleura(Stephanozyga) nishimurai, n. sp. The occurrence of these gastropods suggests Visean age andsupports the previous age assignment of the sandstone bed exposed in the Onimaru Quarry.

has been obtained by the efforts of the staff
members of the Onimaru Quarry Co. Ltd., andhas been made available to specialists for identifi·
cation and description. Additional material was
also collected by myself from 1985 to 1987. The
gastropods described in this paper were collectedfrom a fine- to coarse-grained, ill-sorted, tuffaceous sandstone bed, and occurred in association
with ubiquitous valves of brachiopods andcrinoid stems. These fossils may have been transported for a considerably long distance fromtheir original habitats by bottom currents, since
they are commonly fragmented, eroded andarranged parallel to the laminations. Kawamura(1984) has shown that the sandstone of the H2Member of the Hikoroichi Formation was
deposited on a shallow marine bottom aroundvolcanic islands away from continents.
Mollusks are less dominant fossils in the
Onimaru Quarry, although a number ofgastropod specimens have been collected there.
Although the described species are too few, they
can nevertheless be used to assess previous age
assignments of the bed.
All the specimens used in this paper arehoused in the collection of the Section of
Invertebrate Paleontology, National Science

Introduction

The biota of the Early Carboniferous
Hikoroichi Formation in the Hikoroichi area ofthe Kitakami Mountains, northeast Japan ishighly diverse and has been the subject of anumber of works describing or illustrating taxa
from it (Minato et al., 1979; Tazawa, 1981,
1984a,b). The formation is about 500 m thick inthe type area and consists of such shallow marine
sediments as sandstone, slate and impure
limestone, which are intercalated with thick tuff
beds (Tazawa, 1981; Kawamura, 1983). A richly
fossiliferous sandstone bed crops out at the
Onimaru Quarry in the Hikoroichi area (Fig. 1).
Fossils from this sandstone bed have not yet
been fully described, but Tazawa (1984a)reported the occurrence of two Gigantoproductus species. The bed belongs to Tazawa's(1981, 1984a,b) uppermost part of the Lower
Member of the Hikoroichi Formation and to theupper part of Kawamura's (1983) H2 Member ofthe Hikoroichi Formation.
A large collection from the Onimaru Quarry*Received October 28, 1987; revised manuscript
accepted March 15, 1988; Studies on Japanese Trilobites and Associated Fossils, no. LXIV.

361

Tomoki KASE

362.....•

N

1
"

..

~#.-

....

.~

4

ONIMARU

+

4

-".

OMORI

".
......

39'06'

Figure 1. Fossil locality (NSM-PCL6-53-3) in the
Hikoroichi area.

Museum, Tokyo.

western European regions. These records suggest
a Visean age for the sandstone bed in the Onimaru Quarry.
Recent paleontological studies have indicatedthat the Hikoroichi Formation ranges from thelate Tournaisian to Visean (Mori and Tazawa,
1980; Tazawa, 1981 , 1984a, b; Kawamura, 1983).
Based on the occurrence of two Gigantoproductus species from the same sandstone bed in the
Onimaru Quarry, Tazawa (1984a) dated it as the
Visean. The occurrence of the gastropods described herein supports these age assignments.
Systematic description
Superfamily Euomphalacea
Family Euomphalidae
Genus Straparollus de Montfort, 1810
Subgenus Euomphalus J. Sowerby, 1814
Straparollus (Euomphalus) asanoi, n. sp.
Figures 2-1-4

The gastropod fauna
Six gastropod species described herein are as
follows;
Straparollus (Euomphalus) asanoi, n. sp.
Straparollus (Euomphalus) sp.
Kawanamia onimarensis, n. gen. and n. sp.
Baylea yvanii (Leveille)
Littorinides sp.
Pseudozygopleura (Stephanozyga) nishimurai,
n. sp.
also found four additional species, Bellerophon? sp., Retispira sp., pleurotomariacean gen.
et sp. indet., and Platyceras sp., but they are too
poorly preserved to describe.
One is referred to Baylea yvanii (Leveille),
which has been known to occur from theTournaisian of Belgium, the lower Visean ofWales and the Lower Carboniferous of England(Batten, 1966). Among five other species,
Straparollus (Euomphalus) asanoi, n. sp. has aclose relationship to Straparollus (Euomphalus)
acutus (Sowerby) known from the Visean of thewestern Europe (Batten, 1966). Species of Littorinides and Pseudozygopleura (Stephanozyga)
are also known to occur from the Visean of

Diagnosis: A large and trochiform Straparollus(Euomphalus) having a weakly stepped andconcave to flat sub sutural ramp with maturity, arather narrow umbilicus and a trapeziform whorl
section of body whorl. Spiral angulation on
upper whorl surface thick and round-topped,with deep sinuation on it.
Description: Large trochiform euomphalids
with shell height about 3/5 of shell diameter andwith height of spire slightly smaller than 1/2 ofshell height. Pleural angle in later spire whorls
about 90°. Protoconch not preserved. Early
whorls with regularly rounded upper whorl
surface. Sutures impressed. A weak spiralangulation positioned at midwhorl on last threewhorls in mature specimens, then it becomingstronger toward body whorl. Body whorl with an
obtuse trapeziform whorl section. Upper whorl
surface with flat to weakly concave sub sutural
ramp being delimited by a round-topped, thickspiral angulation. Basal periphery of body whorl
obtusely angulated and whorl surface between
the spiral angulation and basal periphery steeply
inclined and slightly flattened. Base rather
narrowly phaneromphalous and weakly convex,

851. Carboniferous gastropods from Kitakami

363

3

Figure 2. Straparollus (Euomphalus) asanoi, n. sp. 1, apertural view of the ho lotype , NSM-PM15385 ,shell diameter , 88.4 mm , height , 53 .5 mm , x 1.0; 2, top surface of the holot ype showing the dorsal sinu s andgrowth lines, x 1.0; 3, lateral view of a paratype, NSM-PM15387 showing ea rly whorl profile, shell di ame ter ,27.9 mm , height , 27 .5 mm , x 1.0, thi s specimen is strongly dis torted laterall y so that the apical angle is smaller
than the original; 4, oblique view of a paratype, NSM-PM15386 , shell diameter , 93 mm , height , ca. 55 mm , x
0.95 , the spiral angulation on the right half of the body whorl is accentu ated by diagenetic defor mation.
All specimens are silico n rubber casts.

with roundly convex umbilical wall. Umbilicalsutures impressed . Width of umbilicus about 1/ 3of base diameter. Shell surface ornamented
entirely by rugose growth lines; these lines beingorthocline on upper whorl surface above spiralangulation, rather deeply sinuous on spiralangulation, then bending prominently to be comeprosocyrt on upper whorl surface below spiralangulation. Growth lines also gently curving tobecome prosocline across basal periphery andinto umbilicus , then turning to curve orthocline
toward umbilical sutures.
Discussion : In the general outline, Straparollus(£uomphalus)asanoiclosely
resembles

TomokiKASE

364

Straparollus (Euomphalus) acutus (Sowerby,1818) from the Visean of Belgium, England,
Ireland and Germany (de Koninck, 1881; Batten,1966). S. (E.) asanoi also has an ontogenetic
change of whorl profile similar to that of S. (E.)
acutus; both species have a roundly convex whorl
surface in the early growth stage and have a spiralangulation at midwhorl in the later growth stage.
However, the size of S. (E.) asanoi is more than
three times that of S. (E.) acutus.
Straparollus (Euomphalus) asanoi resembles
Euomphalus crotalostomus (M'Coy, 1844), (deKoninck, 1881) from the Visean of Ireland andBelgium in shell size and characteristics of whorl
shape. The Hikoroichi species is distinct as theheight-to-width proportion is much larger thanthat of E. crotalostomus.
Straparollus (Euomphalus) asanoi also
resembles Omphalotrochus kalmiussi Zernetskaja(1967) from the lower Visean of Donetz Basin,
U.S.S.R., but differs in having a more adaxiallylocated spiral angulation on the upper whorl
surface than the latter species.
Etymology: This species is named after Mr. T.
Asano who donated his collection to the
National Science Museum, Tokyo.
Figured specimens: Holotype, NSM-PMI5385;
paratypes, NSM-PMI5386, 15387.
Straparollus (Euomphalus) sp.
Figures 3-4,5Discussion: This species is found commonly in
the Onimaru Quarry. The specimens show
depressed discoidal shell outline. They also haveangulated upper whorl surface and rounded
lower whorl surface. The spiral angulation on theupper whorl surface seems to appear in the later
growth stage. The maximum shell diameter is 40
mm in the largest specimen. Although most of
the specimens of tli1s species are severely
distorted to make comparison with other species
difficult, they differ definitely from S. (E.)
asanoi in its small shell size and in having
planispiral shell form.
Figured specimens: NSM-PMI5394, 15395.

Superfamily Pleurotomariacea
Family Sinuopeidae
Subfamily Omospirinae
Genus Baylea de Koninck, 1883
Baylea yvanii (Leveille, 1835)
Figures 3-1,6-9
Trochus yvanii Leveille, 1835, p. 39, pI. 2, fig. 24.
Pleurotomaria yvanii: de Koninck, 1843, p. 390, pI. 37,
figs. 1,7.
Baylea yvanii: de Koninck, 1883, p. 69, pI. 27, figs.
1-5, pI. 32, figs. 8,9; Knight, 1941, p. 51, pI. 21,
figs. 1a-c; Batten, 1966, p. 25, pI. 3, fig. 4; Peel,
1985, p. 104, pI. 6-6-18.
Yvania yvanii: Fischer, 1885, p. 851.
Baylea communis de Koninck, 1883, p. 70, pI. 27, fig.
11.
Baylea spirata de Koninck, 1883, p. 71, pI. 32, fig. 10.
Baylea duplicosta de Koninck, 1883, p. 71, pI. 25, figs.
11, 12.
Baylea leveillei de Koninck, 1883, p. 73, pI. 27, figs.
6-10.
Baylea simplex de Koninck, 1883, p. 74, pI. 27, figs.
12-15.

Description: Shell thin, gradate and turbiniform, with height greater than width. Height ofspire about 1/2 of total shell height. Pleural angle
about 80°. Protoconch and earlier teleoconchs
not preserved. Sutures distinct, not impressed.
Spire whorls prominently keeled at midwhorl,with a gently inclined and weakly concave upper
whorl surface and vertical outer whorl surface.
Body whorl also possessing a broad, gently
inclined and concave upper whorl surface; its
outer whorl surface having a vertical and veryslightly concave area just below shouldered edge,
then it becoming convex at basal periphery. Baseregularly convex, with a tiny umbilical chink.
Aperture subquadrate in shape, being acutely
angled above and broadly rounded below. Outer
lip thin and angularly convex. Inner lip regularly
concave and slightly reflexed. Basal lip roundly
concave and very slightly reflexed. Surface withfine, sharp and rather regularly spaced spiralcords and with some subordinate ones in theirinterspaces; spiral cords numbering from four tosix on upper whorl surface and about 15 at base,respectively; outermost spiral cord on upper
whorl surface being stronger than others.

851. Carboniferous gastropods from Kitakami

365

2

9

Figure 3. I, 6 - 9 Baylea yvanii (Leveille). 1. apertural view of a specimen, NSM-PM 15 3 88 , shell diamet er ,13.4 mm , height, 13.5 mm, x 1.35 ; 6, apertural views of two specimen s, NSM-PMI5389 , x 1.5; 7, adapertural
view of a specimen, NSM-PMI5390, shell diameter , 12 .5 mm , height , 13.5 mm , x 1.5 ; 8 , oblique view ofa specimen, NSM-PM15391, shell diameter, 14.5 mm, x 1.5; 9, adapertural view of the largest specimen, NSM-PMI5392,
shell diameter, 20.0 mill , x 1.5. 2, 3, Littorinides sp., apertural and apical views of a specimen, NSM-PMI5393 ,shell diameter , 14.5 111m, height , 11.0 111m , x 1.5. 4, 5, Straparollus (EuomphaluS') sp. 4, apica l view of a
distorted specimen, NSM-PMI5394, maximum shell diameter , 23.3 mm , x 1.5 ; 5, apical view of a di storted
specimen, NSM-PMI5395, maximum shell diameter , 22.4 mm , x 1.5. 10 - 14 , Kawanamia onimarensis, gen. et sp.
nov. 10, 11, apertural and basal views of the holotype , NSM-PMI5396 , showing cha racteristics of basal
ornamentation and columellar lip , max imum shell diameter , 24.5 mm, height , 18 .6 mm , x 1.4 ; i2 , latera l view ofa paratype, NSM-PMI5397 , x 1.4; 13 , 14 , apical and lateral views of a paratype , NSM-PM15398 , showing theshallow V-shaped sinus at the upper whorl angul ation , shell diameter, 62 .3 mm , x 1.0.
All specimens are silicon rubbe r casts.

TomokiKASE

366

Selenizone seemingly located just above
shouldered angulation, delimited by outermost
spiral cord and shouldered edge , and represented
by a wide, very gently inclined concave
depression ; one or two very fine spiral cords
present within the selenizone.
Discussion: This species is found commonly in
the Onimaru Quarry . Specimens of this species
all occur as a slightly distorted external mold.
The upper whorl surface is ornamented by spiralcords numbering from four to six; the outermost
one is most prominent. In their interspaces, there
are one or two very fine spiral cords. Although
the coarse-grained nature of the matrix is notsuitable to show delicate characteristics of theselenizone, the selenizone appears to be positioned between the outermost spiral cord andshouldered angulation. The presence of a veryfine spiral cord in the selenizone is also known in
the European specimens of Baylea yvanii (e.g.,
Knight , 1941, pI. 21, fig. Ib).
The Hikoroichi specimens can be referred toBaylea yvanii (Leveille, 1835) which is known tooccur from the Tournaisian of Belgium, theLower Carboniferous of England and the lower
Visean of Wales (Batten, 1966, p. 26). Batten(1966) stated that B. yvanii shows a large
variation in form as to the height of spire and therapidity of whorl expansion among Belgian
specimens and synonymized other five Baylea
species described by de Koninck (1883) with B.
yvanii. As to the shell outline , the Hikoroichi
specimens seem to fall within the range ofvariation of B. yvanii.
Figured specimens: NSM-PMI5388-15392.

around columella with an obtuse angulation
around it. Parietal area covered by thin callus.
Columellar lip wide, semilunar in form andreflexed. Ornamentation consisting of fine lirae
on subsutural area and over base.
Discussion: Kawanamia is erected for only thetype species. The shape of the whorl and theV-shaped shallow notch at the upper whorl
angulation simulate those of Lophospira, but its
low turbiniform shell shape readily distinguishes
from Lophospira. Superficially, Kawanamia
resembles Trochonema and Trochonemopsis inshell form , but differs in having the V-shaped
labral notch at the upper whorl angulation and inlacking a deep umbilicus.
Etymology: The genus is named after Mr. T .
Kawanami , Director of the Onimaru Quarry Co.
Ltd ., who donated his collection to the National
Science Museum, Tokyo and enabled me tocollect the material from that quarry.

Kawanamia onimarensis, n. Sp.
Figures 3-10-14; 4Diagnosis: Same as for the genus .
Description: Large , anomphalous and low
turbiniform shells with height slightly greaterthan width. Spire low conical with height about
2/5 of total shell height . Protoconch and early
whorls not preserved. Spire whorls embracing
previous one just below the upper angulation ,bu t the area of inpingement moving downward in
final whorl. Spire whorl surface broad, gently

Family Lophospiridae
Subfamily Lophospirinae
Genus Kawanamia, n. gen.

Type species: Kawanamia onimarensis, sp. nov.
Diagnosis : Large, anomphalous and low
turbiniform shell with a biangulated outer whorl
surface. Ramp broad, gently inclined and moreor less concave. Labral sinus culminating at upper
keeled angulation in a shallow V-shaped notch.
Base evenly convex, but shallowly depressed

Figure 4_ Reconstruction of Kawanamia onimarensis, n. gen. and n. sp. (x ca. 1).

851. Carboniferous gastropods from Kitakami

inclined, weakly concave and flushed. Body
whorl with a broad and concave ramp and abiangulated outer whorl surface; the upper
angulation being more acute than the lower one
whose position being situated slightly above
rounded basal periphery; outer whorl surface
vertical and concave. Base with evenly convex
surface in main part, but the area aroundcolumella shallowly depressed with an obtuse
angulation at abaxial margin. Aperture weakly
tangential to body whorl and subquadrangulate
in outline. Parietal callus thin and extending
slightly from inside the aperture. Columellar lip
wide, semilunar in shape and reflexed.
Upper whorl surface ornamented by fine,prosocline lirae that follow growth lines, arise at
the upper suture and die out at the middle ofupper whorl surface; they become obsolete in the
final whorl. Base covered by fine, sinuous lirae.
Growth lines fine, steeply prosocline on theupper whorl surface, forming an acute sinus at
upper angulation, then curving to form a verticaland roundly prosocyrt trend on the outer whorl
surface; the growth lines on base widelyopisthocyrt on the main part and turning tosinuate around the columella.
Discussion: Comparison of this species withrelated forms is made in the discussion part of
the genus.
Figured specimens: Holotype, NSM-PM15396;
paratypes, NSM-PM 15397, 15398.
Superfamily Neritacea?Family P1agiothyridae
Genus Littorinides Knight, 1937
Littorinides sp.
Figures 3-2,3Discussion: One specimen is referrable to
Littorinides, but it is not enough to ascertain its
specific identity. Although no aperture ispreserved in this specimen, its shell size, shell
outline and surface ornamentation are quite
similar to those of Littorinides solidum (deKoninck, 1843), the type species of the genus,
described from the Lower Carboniferous of
Belgium, which was redescribed and illustrated367

by Knight (l941, pI. 82, figs. 1a,b). The
Hikoroichi taxon differs from L. solidum inhaving coarser spiral cords.
The shell is small in size and turbiniform, andhas well inflated whorls that are separated byimpressed sutures. The whorl inflation is weak in
the upper whorl surface and strong at the base.
The surface ornamentation consists of fine, sharp
spiral cords and prosocline growth lines. The
growth lines are very finely granulated at place
where these spiral cords intersect.
Figured specimen: NSM-PMI5393.
Superfamily Loxonematacea
Family Pseudozygopleuridae
Genus Pseudozygopleura Knight, 1930
Subgenus Stephanozyga Knight, 1930
Pseudozygopleura (Stephanozyga) nishimurai,
n. sp.
Figures 5-1- 3

Diagnosis: A very large species of Pseudozygopleura (Stephanozyga) having higher proportion of whorl and slightly opisthocline collabral
ribs. Whorls adpressed. Number of ribs about 14
on each whorl.
Description: Shell attains more than 15 cm inheight, high spired, with apical angle of about
20°. Protoconch and early whorls unknown.
Whorl profile asymmetrically inflated, slightly
concave from suture to upper one-third of whorl,then it gradually becoming convex downward;
the highest point located at lower one fifth ofwhorl. Early whorls adpressed, but suture
gradually becoming deeper between later whorls.
Preserved early four whorls in the holotype
seemingly smooth, with whorl profile similar tothat of later whorls, with axial ribs appearing in
the fifth whorl. Curvature of the upper whorl
surface of body whorl similar to that of previous
whorls, with weakly convex and steeply inclined
base, no detectable angulation occurring between
outer whorl surface and base. No umbilicus.
Outer lip and anterior extremity poorly preserved, but the former widely convex and thelatter weakly projected anteriorly to form an
ill-defined anterior sulcus. Columellar lip almost

368

TomokiKASE

2Figure 5. Pseudozy gop/eura (Stephanozy ga) nishimurai, n. sp. 1, lateral view of the holotype,
NSM·PM 15399 , showing early four smooth and later six strongly ribbed whorls, height , ca. 89 mm x 1.0; 2,apertural view of a paratype, NSM·PM 15400, showing the reflexed columellar lip , she ll diameter , 22.7 mm ,height , 57.5 mm , x 1.0; 3, lateral view of a large paratype, NSM-PMI5401 , showing the ribbed spire who rl s andthe smooth body whorl , shell diame ter , 48.3 mm , height , 83 .0 mm , x 1.0.

straight, vertical and reflexed . Surface sculptured
by collabral ribs and very fine growth lines. Ribs
strong, 14 in number in each whorl of theholotype , more or less aligned in vertical rows
from whorl to whorl , very slightly opisthocline
and sinuous, round-topped, slightly narrower
than their interspaces, suppressed above andbelow, and seemingly missing or very weak inbody whorl of adult specimens. Growth lines
orthocline for a short distance below upper
suture , then turning to opisthocyrt below.
Discussion: This species is represented by fiveincomplete external molds. They are considerably distorted by diagenetic compression except
for the holotype , so that they show slightly
wider shell profile than their original state. The
holotype preserves four early smooth whorls andsix strongly ribbed spire whorls. Characteristicsof the whorl profile , collabral ribs and sutures areclearly represented in this specimen. Two large
and probably ma ture specimens, al though they
are poorly preserved, possess a seemingly smooth
body whorl. The loss of ribs, costae or nodes atthis point is also known in such species as
Pseudozygopleura(Stephanozyga)
rugifera(Phillips, 1836), (Longstaff, 1933 , pI. 7, fig. 7) ,P. (S.) subnodosa Knight (1930), and Loxonema
murchisonianum de Koninck (1843) , (deKoninck, 1881).
The size of this species reconstructed from thelarge and small specimens may attain more than
15 cm in height. This species is the largest among
the members of Stephanozyga and can be
distinguished from all other species primarily by851. Carboniferous gastropods from Kitakami
its large shell size. Pseudozygopleura (Stephanozyga) nishimurai resembles P. (S.) rugifera fromthe Lower Carboniferous of the United Kingdom
and Belgium, but the loss of collabral ribs in the
body whorl of the specimen illustrated byLongstaff (1933) suggests that P. (S.) rugifera
is much smaller than P. (S.) nishimurai even in
its adult stage. Furthermore, P. (S.) nishimurai
has much stronger collabral ribs than P. (S.)
rugigera.
Pseudozygopleura (Stephanozyga) nishimurai
resembles Pseudozygopleura magna Wang (1987)
from the Lower Carboniferous of Guizhou,
China, but differs from the latter in its muchlarger shell size and in having coarser axial ribs on
the spire whorls. P. (S.) nishimurai is also
distinguished from P. (S.) nodosa (Girty, 1915)
and P. (S.) subnodosa Knight (1930), both fromthe Upper Carboniferous of Missouri, primarily
by its much larger shell size.
Etymology: This species is named after Mr. S.
Nishimura who donated his collection to the
National Science Museum, Tokyo.
Figured specimens: Holotype, NSM-PM 15399;
paratypes, NSM-PMI5400, 15401.
Acknowledgments
Messrs. T. Kawanami, T. Asano, S. Nishimura
and S. Kinno generously donated their collections to the National Science Museum, Tokyo. I
thank Messrs. T. Ono, A. Kaneko and S. Niko for
their help in the field. I also thank Dr. K.
Uemura who reviewed the manuscript.
References
Batten, R. L. (1966): The Lower Carboniferous
gastropod fauna from the Hotwells Limestone ofCompton Martin, Somerset. Part 1. Monogr.
Palaeont. Soc. London, no. 509, p. 1-52, pis. 1-5(published as a part of volume 119 for 1965).
Fischer. P. (1887): Manuel de conchyliologie et depaleontologie conchyliologique ou histoire
naturelle de mollusques vivants et fossiles. 1369 p.,
23 pis., Libraille F. Savy, Paris.
Girty, G. H. (1915): Invertebrate paleontology. In
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of the Pennsylvanian Series in Missouri. MissouriBur. Geol. Mines, 2nd Ser., vol. 13, p. 263-375,
pis. 27-32.
Kawamura, T. (1983): The Lower Carboniferous
formations in the Hikoroichi region, southern
Kitakami Mountains, northeast Japan (Part 1) Stratigraphy of the Hikoroichi Formation -. Jour.
Geol. Soc. Japan, vol. 89, no. 12, p. 707 -722 (inJapanese with English abstract).
--(1984): The Lower Carboninferous formations intheHikoroichi region, southern Kitakami
Mountains, northeast Japan (Part 2) - Sedimentological study of sandstone and limestone -. Ibid.,
vol. 90, no. 11, p. 831-847, pis. 1-3 (in Japanesewith English abstract).
Knight, B. J. (1930): The gastropods of the St. Louis,
Missouri, Pennsylvanian outlier: the Pseudozygopleurinae. Jour. Paleont., vol. 4, supplement 1, p.
1-88, pis. 1-4.
- - (1941): Paleozoic gastropod genotypes. Geol.
Soc. Amer., Spec. Papers, no. 32, p. 1-510, pis.
1-96.
Koninck, L. G. de (1843-1844): Description de animauxfossiles, que se trouvent dans Ie terrain carbonifere
de Belgique. 651 p., 54 pis., Liege.
--(1881, 1883): Faune de calcaire carbonifere de laBelgique. Gasteropodes. Ann. Mus. Roy. Hist.
Natur. Belgique, Ser. Paleont., vol. 6, p. 1-157,
pis. 1-21; vol. 8, p. 1-240, pis. 22-54.
Leveille, C. (1835): Aperr,;e geologique de quelques
localites tres riches en coquilles sur les frontiers deFrance et de Belgique. Mem. Soc. geol. France, vol.
2, p. 29-30, pI. 2.
Longstaff, J. D. (1933): A revision of the British
Carboniferous Loxonematidae, with descriptionsof new species. Quart. Jour. Geol. Soc. London,
vol. 89, p. 87-124, pis. 7-12.
M'Coy, R. (1844): A synopsis of the characters of the
Carboniferous limestone fossils of Ireland 207 p.,
29 pis., Dublin.
Minato, M., et al. (1979): The Abean Orogeny. 427 p.,
144 pis., Tokai Univ. Press, Tokyo
Mori, K. and Tazawa, J. (1980): Discovery andsignificance of Visean rugose corals andbrachiopods from the type locality of the Lower
Carboniferous Hikoroichi Formation. Jour. Geol.
Soc. Japan, vol. 86, no. 2, p. 143-146 (inJapanese).
Peel, J. S. (1985): Class Gastropoda. In Murray, L. W.
(ed): Atlas of invertebrate macrofossils, p.
102-113. John Wiley and Sons, New York.
Phillips, J. (1815-1818): Illustration of the geology ofYorkshire, part 2. The Mountain Limestone
district, 253 p., 24 pis., London.
Sowerby, J. (1815-1818): The mineral conchology ofGreat Britain. London, vol. 2, p. 1-251, pis.

TomokiKASE

370

103-203.
Tazawa, J. (1981): Notes on some Carboniferous
brachiopods from the Kitakami Mountain, northeast Japan. Saito Ho-on Kai Mus. Res. Bull., no.
49, p. 53-60, pI. 4.
- - (1984a): Occurrence of Gigantoproductus (Carboniferous Brachiopoda) from the lower Hikoroichi
Formation, Kitakami Mountains, northeast Japan.
Earth Sci. (Chikyu Kagaku) , vol. 38, no. 2, p.132134, pI. 1 (in Japanese).
--(1984b): Early Carboniferous (Visean) brachiopods from the Hikoroichi Formation of theKitakami Mountains, northeast Japan. Trans. Proc.
Palaeont. Soc. Japan, N.S., no. 133, p. 300-312,
pI. 6l.
Wang, H. (1987): Molluscs from Lower Carboniferous
Bojiwan Formation of Weining, Guizhou. Acta
Palaeont. Sinica, vol. 26, no. 3, p. 319-322, pI. l.
Zernetskaja N. V. (1967): Gastropod fauna of theLower Visean of Donetz Basin. In Dunaev, N. N.
(ed.): Fauna of the Lower Visean deposits ofGreater Donbas, p. 30-42, pIs. 16-24. Naukova
Dumka, Kief (in Russian).

Of una to ::i;;J1Hil1t, Onimaru *1L, Hikoroichi B~riJ .

:lR:lt B*:ltJ::Wttl!.O) B ~riJJ!l!i.i£::fiER*C.lltrJtIl (\::"-j!'-tit) Jm)E~ : :lR:lt B*:ltJ::Wttl!., jdYdil1triJ':.hi?(i:ltJ::Wttl!.-C:'(iW~-c~cillli: ~.ht.:::fiER*c.O)Jm)E~-C:';l; ~o Kawanamia ~fi Lophospiridae NO)¥r ~ c L -C §cillli: Lt.:~c tit L t.: fi fi Straparollus (Euomphalus) asanoi, n. sp., S. (E.) sp" Kawanamia
0)*jLl*::fi~1:.n:±l-t~ B~riJJ!l!iO)r&:fiJ: I) 6fiO)Jm)E~z.§cillli:U:o0conimarensis, n. gen. and n, sp., Baylea yva.nii (Leveille), Littorinides sp. Pseudozygopleura(Stephanozyga) nishimurai, n. sp. -c:';l;~o ':.hi?0) 5 ~O)-fifi-</I..4"'-o) ~ 11..;f. -titc 14'"\I A 0) 1::' -j!' - titl)' i?%I i?.h -C l' ~ Baylea yvanii 1:' JqJ;<E ~.h, S. (E.) asanoi fi -< I\.. 4'" -, I-" 1';J, 14'" \I A c 71 1\..7 /' 1-"0) t:'-j!'-jjjx;;Pi?.h~ S. (E.) acutus (SowerbyH:'.iti~ c "If; x.i?.h~o ':.hi?O)Jm)E~O).i£:±l~*1Ll*::fi~o)B~riJJ!l!iO)~:fi~~~-tit-C;l;~,:cz.~~L.~*
O).:O)~:fiO)~ft.z.~~-t~o~.~.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 149, pp. 371-377, 3 Figs., Apri130, 1988

852. FISHERITES, A RECEPTACULITID FROM THE
MIDDLE ORDOVICIAN OF PYONGYANG COAL-FIELD*
SHun NIKO
Department of Earth Science and Astronomy, College of Arts and Sciences,
The University of Tokyo, Komaba, Tokyo 153
andNOBUOIKEBE
6-4, Hama-machi, Ashiya City 659

Abstract. A single specimen of Middle Ordovician receptaculitid from the Josen
Supergroup of North Korea was previously called as Receptaculites coreanicus Ikebe (nom.
nud.) by Ikebe in 1939. We reinvestigated the specimen, and found that it was a new species of
the genus Fisherites, F. coreanicus Niko and Ikebe on account of having relatively large thallus
and asymmetrical intercalary meroms.

mostly of Archeozoic gneisses, the Upper
Proterozoic Sangweon Group and the CambroOrdovician Josen Supergroup. The receptaculitid
described here was collected by Ikebe in 1936
from the uppermost part of the Josen Supergroup, and was documented only in a report ofgeological investigation of eastern district of the
Pyongyang Coal-field under the name of
Receptaculites coreanicus (Ike be, 1939; Ikebe in
Nakamura et al., 1957). The original specimen,
which has been deposited in the paleontologicalcollection of Kyoto University, is restudied here
and assigned to the genus Fisherites Finney andNitecki, 1979.
Receptaculitids are rare to occur in Far East
Asia, and so far as we know there are only two
published records of receptaculitids besides our
specimen; they are Calathium frechi Endo, 1932
from the Santao Formation (Canadian, Early
Ordovician) in the Liaoning Province, ofNortheast China and Receptaculites sp. (Hamada
and ltoigawa, 1983) of the Lower (to Middle?)
Devonian Fukuji Formation, central Japan.
The specimen described in this paper isIntroduction
Receptaculitids are problematic organisms of
Early Ordovician to Devonian, possibly of latePaleozoic, possessing discoidal to columnar
skeletons consisting of the assemblages ofmeroms. Since the beginning of the nineteenth
century, this organism has been interpreted invarious ways by various authors as algae, pinecones, foraminifers, sponges, corals, cystoids,tunicates, transitional form between sponges andcorals, or an independent extinct phylum(Kesling and Graham, 1962), however, an algal
hypothesis becomes prevailing after Kesling andGraham (1962), Byrnes (1968), Nitecki (1969,
1970, 1972) and Rietschel (1969, 1977). The
purpose of this paper is to describe themorphology of a Middle Ordovician receptaculitid specimen from the Pyongyang Coal-field.
The Pyongyang Coal-field occupies a large
area of North Korea with Precambrian andLower Paleozoic basement, which is composed*Received June 25, 1987; revised manuscript accepted
March 15, 1988
371

Shuji NIKO and Nobuo IKEBE

372

registered and deposited in the Department of
Geology and Mineralogy, Faculty of Science,
Kyoto University (KUlP).
Geological setting and fossil occurrence
The Cambro-Ordovician sequence, which iscomposed mainly of carbonate rocks withsubordinate amounts of shale and quartzite, iscalled collectively the Josen Supergroup in
Korea, and is commonly divided into two major
types on the basis of their rock types and faunas,i.e., the Duwibong-type sequence and the
Yeongweol-type sequence (Kobayashi, 1928,
1966; Yosimura, 1940). The Duwibong-type
Josen Supergroup is characterized by continental
shelf deposits. On the other hand the Yeongweoltype was formed in the axial region of the
Ogcheong basin and contains Chiangnan type
fauna of South China (Kobayashi, 1967). The
Josen Supergroup in the Pyongyang Coal-field,

from which the receptaculitid fossil wascollected, belongs to the Duwibong-typesequence including faunas with the Hwangho
affinity of Northeast China.
Fisherites coreanicus occurs with Actinoceras
sp. in the weakly recrystallized and dolomitized
limestone (peloidal grainstone) of the uppermost
Josen Supergroup at the locality 1.4 km south of
Samtung (Figure 1). Geology and stratigraphy ofthis area were described by Ikebe (1939) andNakamura et al. (1957) in detail. According tothem, the Ordovician sequence of the J osen
Supergroup is made up of, in ascending order,
the Kotyongsan Formation, the Sanbatsmol
Formation and the Kangjinsan Formation, in
which the fossil site lies in the middle part. The
equivalent strata (Unhak Formation) of the Mt.
Mantal area with the Kangjinsan Formation
contains many cephalopods including Actinoceras, Armenoceras, Stereoplasmoceras, Vagino-

Josen Supergroup
Kangjinsan Fm.
Fm.
Kotyongsan Fm.
Imchyon Fm.
Syonglla Fm.
Sangweon Groupkilometers

Fa uI tFigure 1. Map showing collecting locality (X), and geology of the Samtung area, North Korea (modified
from Ikebe, 1939).

852. Fisherites from Pyongyang Coal-field
ceras and others (Kobayashi, 1928, 1929, 1930),
gastropods (Kobayashi, 1929, 1930) and stromatoporoids (Yabe and Sugiyama, 1930). As notedby Kobayashi (1929, 1930), this fauna shows anaffinity to the Toufangkou fauna and Machiakou
fauna of Northeast China. Biostratigraphicallythe Kangjinsan Formation spans from the late
Chazyan to the Ashbyan age, Middle Ordovician.

Systematic paleontology
Class Receptaculitaphyceae Weiss, 1954
Order Receptaculitales James, 1885
Family Receptaculitaceae Ej~hwald, 1860
Genus Fisherites Finney and Nitecki, 1979
Fisherites coreanicus Niko and Ikebe, sp. nov.
Figures 2-1-3; 3-A-D
1939 Receptaculites coreanicus Ikebe (nomen nudum),
p.436.
1957 Receptaculites coreanicus Ikebe. Nakamura,
Matsushita, Kobatake and Ikebe, p. 13 (listed).

Etymology: - The specific name is derived
from Korea.
Diagnosis: -Medium- to small-sized species ofFisherites characterized by possessing subglobular thallus with small corniculum, inflated
proximal end of merom shafts and three distinct
types of intercalation of meroms.
Material:-Description of this new species isbased upon only a single deformed specimen(KUJP 10201) in which no lacunar hemisphere ispreserved.
Terminology:- The terminology in this paper
follows Fisher and Nitecki (1982).
Description: - Thallus: Thallus is medium- tosmall-sized for the genus. The preserved portionof nuclear hemisphere is shallow dish shape,
approximately 6.0 cm x 4.7 cm across. The
nucleus and its immediate vicinity area of approximately 4 mm in diameter are missing. However, we can assume that there was a small
corniculum at the nuclear part, because of a
weak projection of inner wall is recognized at thesuspected point.

373

Meroms: Meroms are arranged in circlets
around a reconstructed nucleus. Each merom isdivided into head and shaft, and individual heads
consist of thick and rhomboidal plates, which
form the outer wall with the juxtaposition of
adjacent meroms and four ribs of stellate structure. Ribs are short and stout. Among themselves
the abapical medional ribs are situated nearest
the outer wall. The proximal ends of columnar
shaft are inflated and form a thick inner wall.
The size of shaft and head decreases toward
the nucleus; the length of the merom attains
15.0 mm at least. Small elliptic perforations arerecognized on a diagonal line of the plate. Eachmerom is enveloped in a dark thin micritic layer(Hiillschicht; Rietschel, 1969), and its inner partconsists of sparry calcite. Except for the meroms,
intervallum is filled with the sediment (Figure
2-3).
Intercalation of meroms: The merom heads
are not preserved in the vicinity of nucleus
because of weathering, nevertheless it is possiblethat the plate boundaries are reconstructedemploying a pattern of merom shafts. In thisspecimen, three distinct types of intercalation are
distinguishable; (1) a symmetrical intercalation at
normal interpositum and triangulum (Figure3-A); (2) a symmetrical intercalation atinterpositum-triangulum triplet (Figure 3-B); and(3) an asymmetrical intercalation (Figures 3-C,

D).
The latitudinal distribution of those types isalmost mutually exclusive. There are 6 (+ 3?)
visible pairs of the normal interpositum andtriangulum causing a symmetrical intercalation,and they appear to be restricted to the vicinity of
nucleus. The interpositum-triangulum triplets are
distributed more lacunarward of the thallus thannormal one; and their visible numbers are 9(+3?). The latitudinal distribution of asymmetrical intercalations is confined to over a distance ofapproximately 24 mm from the suspected nucleus; there are 2 asymmetrical intercalary parastichies, of which direction is opposite
Discussion: - The present specimen is referred
to Fisherites Finney and Nitecki, 1979 on
account of a relatively large thallus among the374

Shuji N IKO and Nobuo IKEBE

852. Fisherites from Pyongyang Coalfield

c

0

375

V
A,B

'------_I

C, D

Figure 3. Intercalation in Fisherites coreanicus Niko and Ikebe, sp. nov., camera lucida drawing of plateboundaries or outline of merom shafts in the neighbourhood of intercalation. Interposita, triangula and theinitial plates of asymmetrical intercalary parastichies are indicated by stipling.
A, interpositum-triangulum pair. B, interpositum-triangulum triplet. C, asymmetrical intercalation of sinistral
direction. D, asymmetrical intercalation of dextral direction. All bar scales equal 4mm.

known receptaculitids and having the asymmetrical intercalary meroms. As noted by Fisher andNitecki (1982) in Fisherites occidentalis (Salter),F. reticulatus (Owen) and Calathium egerodae
Nitecki, the cooccurrence of both symmetricaland asymmetrical intercalation is recognized in
the examined specimen.

Fishentes coreanicus, sp. nov. is somewhat
similar to F. occidentalis (Salter) (Hinde, 1884;
Fisher and Nitecki, 1982) from the Middle
Ordovician of Quebec, Canada, in the generalappearance of meroms and the shape of thallus,
but differs from it in having a smaller corniculum
and inflated proximal end of the merom shafts.

~ Figure 2.
Fisherites coreanicus Niko and Ikebe, sp. nov., from the Kangjinsan Formation (Middle
Ordovician) of the Josen Supergroup, North Korea; holotype, KUJP 10201. 1, nuclear hemisphere, submerged inwater. 2, nuclear vicinity of Figure 2-1, coated with ammonium chloride. 3, thin section parallel to merom
shafts, central axis on right. All bar scales equal 1 cm.

Shuji NIKO and Nobuo IKEBE

376

Moreover, the interpositum-triangulum triplet
causes a symmetrical intercalation, which ischaracteristic feature of the examined specimen,
and rare in F occidentalis on the contrary.
Fisherites reticulatus (Owen) of the Middle
Ordovician Galena Group, North America(Finney and Nitecki, 1979; Fisher and Nitecki,1982) is clearly distinguished from the present
form in that F reticulatus possesses a larger
thallus than the F coreanicus and lacks the interpositum-triaugulum triplet.
Fisherites burmensis Rietschel and Nitecki,
1984 of the Middle Ordovician in the Shan
Plateau region, Burma differs from F coreanicus
in having the clustered intercalation.
Acknowledgments
We thank Drs. T. Kobayashi of the Japan
Academy and T. Hamada of The University ofTokyo for fruitful discussions and critical reviewsof the manuscript. Also we wish to thank Drs. K.
Chinzei and D. Shimizu of Kyoto University for
their permission to study the material in thecollection in the Department of Geology andMineralogy. Part of this research was supported
by a grant from the Japan Academy.
References
Byrnes, J. G. (1968): Notes on the nature andenvironmental significance of the Receptaculitaceae. Lethaia, vol. I, p. 368-381.
Endo, R. (1932): The Canadian and Ordovician
formations and fossils of South Manchuria. Bull. U.
S. Nat. Mus., 164, p. 1-152, pis. 1-40.
Finney, S. C. and Nitecki, M. H. (1979): Fisherites n.
gen. reticulatus (Owen, 1884), a new name for
Receptaculites oweni Hall, 1861. Jour. Paleont.,
vol. 53, p. 750-753.
Fisher, D. C. and Nitecki, M. H. (1982): Standardizationof the anatomical orientation of receptaculitids.
Mem. Paleont. Soc., 13, p. 1-40.
Hamada, T. and ltoigawa, J. (1983): Japanese fossils.
Shougakukan, Tokyo, p. 1-168 (in Japanese).
Hinde, G. J. (1884): On the structure and affinities ofthe family of the Receptaculitidae, including
thereinthe genera Ischadites,
Murchison(=Tetragonis, Eichwald); Sphaerospongia, Pengelly;
Acanthochonia, gen. nov.; and Receptaculites,

Defrance. Quart. Jour. Ceol. Soc. London, vol. 40,
p. 795-849, pis. 36-37.
lkebe, N. (1939): Geological structure of the Santo and
Shogen districts to the east of Phyongyang. Jour.
Ceo!. Soc. Japan, vol. 46, p. 433-444 (in Japanese).
Kesling, R. V. and Graham, A. (1962): Ischadites is adasycladacean alga. Jour. Paleont., vol. 36, p.
943-952, pis. 943-952.
Kobayashi, T. (1928): Ordovician fossils from Coreaand South Manchuria. Japan. Jour. Ceo!. Geogr.,
vol. 5, p. 173-212, pis. 18-22.
--(1929): On the Ordovician formations in South
Manchuria and North Korea (Part 1). Jour. Ceo!.
Soc. Japan, vol. 36, p. 520-535 (in Japanese).
- - (1930): Ordovician fossils from Korea and South
Manchuria. Part II. On the Bantatsu Bed of theOrdovician age. Japan. Jour. Ceol. Ceogr., vol. 7,
p. 75-81, pis. 8-H.
-(1966): The Cambro-Ordovician formations andfaunas of South Korea, Part X. Stratigraphy of the
Chosen Group in Korea and South Manchuria.
Sect. A. The Chosen Group of South Korea. Jour.
Fac. Sci. Univ. Tokyo, Sec. II, vol. 16, p. 1-84.
--(1967):The Cambro-Ordovician formations andfaunas of South Korea, Part X. Stratigraphy of the
Chosen Group in Korea and South Manchuria.
Sect. C. The Cambrian of eastern Asia and otherparts of the continent. Ibid., vol. 16, p. 381-535.
Nakamura, S., Matsushita, S., Kobatake, N. and lkebe,N. (1957): Studies on the geologic structure of the
Phyongyang Coal Field, Korea. Mem. Coli. Sci.
Univ. Kyoto, Ser. B., vol. 23, Special Number, p.
1-63, pis. 1-24.
Nitecki, M. H. (1969): Redescription of Ischadites
koenigii Murchison, 1839. Fieldiana Ceol., vol. 16,
no. 13, p. 341-359.
--(1970): North American cyclocrinitid algae. Ibid.,
vol. 21, p. 1-182.
--(1972): North American Silurian receptaculitid
algae. Ibid., vol. 28, p. 1-108.
Rietschel, S. (1969): Die Receptaculiten. Senckenbergiana Lethaea, vol. 50, p. 465-517.
--(1977): Receptaculitids are calcareous algae but nodasyclads. In Fossil algae, edited by Flugel, E., p.
212-214. Springer- Verlag, Berlin.
--and Nitecki, M. H. (1984): Ordovician receptaculitid algae from Burma. Palaeont., vol. 27, part 2,
p.415-420.
Yabe, H. and Sugiyama, T. (1930): On some Ordovician
stromatoporoids from South Manchuria, North
China and Chosen (Corea), with notes on two new
European forms. Sci. Rep. Tohoku Imp. Univ., Ser.
2, vol. 14, part 1, p. 47-62, pis. 17-23.
Yosimura, I. (1940): Geology of the Neietsu district
Kogendo, Chosen. Jour. Ceol. Soc. Japan, vol. 47:
p. 112-122, pI. 8 (in Japanese).

852. Fisherites from Pyongyang Coal-field

377

Pyongyang'fl!1M. Josen lji!j!~L Sangweon tH1C San tao .:::: .ilL Liaoning ~~:[, Fukuji Nil.tt!l.
Duwibong 4 UM ~. Yeongweol '¥ ~. Ogcheong i7C}lI. Chiangnan ft iI. Hwangho !It iOJ •Samtung .:::: ~. Syonglla i'~ litg. Imchyon f* H. Kotyongsan ~ ~ W. Sanbatsmol !f!; EB im •Kangjinsan i:n~il-'. Unhak ~rt.rL \1antal I!lt~. Toufangkou liJ5HIIL Machiakou .~*@\'.

:>f!1MiREB.ltll)l.Jl:. cpffl5::t /L.- F' e.'AifdJ' Gil)Fisherites (v-l:! 7' j1=\,- "" :7 -{ TAm) : ilI!.J!2iM936ip
v -l:! 7' j1 =\'- "" :7 -{ T A mil) 1-'f:il)1#:. Receptaculites coreanicus Ikebe (nom. nud,) !:: T~[t'~i'::1flr:t '!!:ht.: (ilI!.:i?l. 1939)0 ~'@]~:q~:\IJt~til)~3*. J:tti:l¥Jj;:~il)~:!*1* (thallus). to J: cnr-M¥J;jtffA? p J." (asymmetricalintercalary merom) ~M'/)::!:: 7J' G. :: il) {IM-t ~. Fisherites JI.1H,:: -@;6b t.:. '!!: G t,:: Fisherites
coreanicus Niko and Ikebe, sp, nov,!:: L L -'f:-il)if~'UJ~~c~ L t.:oI)c f-M&1'f) • it!l.:i?lJ~1:
i'::::ItijifJ~. ~vJ!f,t]Jil!11:U I) l*~ Lt.:cpM::t /L.- F' e:' A*e.

ool*,t.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 149, pp. 378-395, 9 Figs., April 30, 1988

853. SOME INOCERAMIDS (BIVALVIA) FROM THE
CENOMANIAN (CRETACEOUS) OF JAPAN - III
THREE SPECIES OCCURRING COMMONLY
IN THE NORTH-WEST PACIFIC REGION*
T ATSURO MATSUMOTO
c/o Department of Geology, Kyushu University 33, Fukuoka 812

AKITO ASAI
Division of Geology, Department of Mineral Industry,
School of Science & Engineering, Waseda University, Tokyo 160

HIROMICHI HIRANO
Institute of Earth Science, School of Education, Waseda University, Tokyo 160
and

MASA YUKI NODA
5-Kumi, Fukagochi, Oita 870

Abstract. This is Part III of the serial descriptions of inoceramid species from the
Cenomanian of Japan. In this part we describe the following three species which occur
commonly not only in the Pacific areas of the USSR (Pergament, 1966) but also in Hokkaido.
(1) Inoceramus pennatu[us Pcrgament. As the specific name indicates, this species ischaracterized by a well developed posterior wing. For some reasons Pergament presented
sufficient examples of the left valve but few of the right. This is supplemented by the specimensfrom Hokkaido. I. pennatu[us occurs in both the Upper and Middle Cenomanian. (2) l.
reduncus Pergament. Several examples of this species have been acquired from the Upper
Cenomanian of Hokkaido. In addition, there are deficient specimens which are comparable withthis species. If they are taken into consideration, the species may range down to the middle partof the Cenomanian. A question remains that l. reduncus Pergament, 1966 may be a juniorsynonym of I. korjakensis Terechova, 1965. (3) I. ginterensis Pergament. In addition to theoriginal specimens from the Pacific areas of USSR, typical examples have been reported fromthe Upper. Cenomanian of the Interior Province of the U.S.A. This species occurs also fairly
commonly in the Middle to Upper Cenomanian of Hokkaido. Although there is some extent ofvariation, no marked distinctions have been noticed between examples of different provinces or
stratigraphic positions.

of our collections from the Cenomanian rocks ofJapan. In this paper (Part III), we describe thethree species which have been already establishedby Pergament (1966) on plentiful material fromthe Pacific region of the USSR and occur also

Introduction

Succeeding to Part II, we continue to report
the result of our study on inoceramid specimens*Received December 7,1987; accepted March 3, 1988.
378

853. Cenomanian inoceramids from Japan - II!
fairly commonly in Hokkaido.
The material of this study consists partly of
the recent acquisitions by students of Waseda
University during the field work conducted by
Hirano in the Oyubari area, central Hokkaido. In
addition to previous papers (Hirano et al., 1981),
the description in detail of the stratigraphy ofthis area is being under preparation for
publication in which route maps and illustrated
stratigraphic sections will be included (Hirano etal., 1988).
Another part of the material is selected
specimens obtained through the field works bymembers of the regional working group of theCretaceous Stratigraphy and Mid-Cretaceous
Events conducted by Matsumoto, with whom M.
Noda, H. Hirano, K. Tanabe, H. Okada, Y.
Miyata and S. Toshimitsu cooperated. The works
were carried on the Oyubari, Ikushumbets, Obira(=Obirashibe) and northerly adjacent Kotanbetsu
areas. As to the stratigraphic subdivisions andfossil locations, readers may refer to Matsumoto
et al. (1978), Hirano et al. (1977), Matsumoto(1965), Tanabe et al. (1977) and Matsumoto andOkada (1973).
As another part of the material of this study
some specimens of the older collections by
Matsumoto are reexamined. They are from thecelebrated Saku area in the transverse mid-valleyof the River Teshio and also from the Shiyubariarea, i.e., northeasterly contiguous with the
Oyubari area. The stratigraphy of these areas was
described by Matsumoto (1942), in which fossil
localities were tabulated and shown in routemaps.
The locality numbers in this paper have theprefix R for the Obira and adjacent areas, theprefix T for the mid-valley of the River Teshio,
the prefix Y for the Oyubari-Shiyubari area andthe prefix Ik for the Ikushumbets area.
The repositories of the described specimens
are as follows with abbreviation in parentheses:
Geological Collections, Kyushu University,
Fukuoka (GK)
Jonan Geological Association, c/o M. Noda,
Fukagochi, Oita (JG)
PaleontologicalCollections,
Schoolof379

Education, Waseda University, Tokyo (WE)
The technical terms for the morphologic
characters and their abbreviations are the same as
those used in Part I (Matsumoto and Noda, 1986,
p. 410) and Part II (Matsumoto et al., 1987).
Systematic Descriptions
Genus Inoceramus Sowerby, 1814

Inoceramus pennatulus Pergament, 1966
Figures 1-1,2; 2-1, 2; 31966 Inoceramus pennatulus Pergament, p. 35, p1.5,
figs. 1,2; pI. 6, figs. 1,2; pI. 7, figs. 1-3.

Holotype:-No. 10-3580-33 (left valve) designated and illustrated by Pergament (I 966,
pI. 5, fig. 1), from the Mamechinsoko Group(Cenomanian) of northwestern Kamchatka.
Material:-GK. H89 (Fig. 1-1), internal mouldof right valve, from a drift in a small gully
between locs. T831 and T832 where the lower
part of Member lIb is exposed, a small branch
No. 2 of the River Teshio near Chirashinai(=Tirasinai in Matsumoto, 1942, pI. 12), ColI. T.
Matsumoto. WE. POOl Y (Fig. 1-2), internalmould of right valve, with test attached partially,
from loco Y250079, i.e. 170m southeast of loco
Y168 of Matsumoto (1942), on the right bank of
the Hikage-zawa, Member My7 of the ShiyubariOyubari area, obtained by T. Sugiyama. JG.
H2492 (Fig. 2-1), internal mould of left valve,with deficient postero-dorsal part, from loco
Ik1040 of Matsumoto (1965), Member IIc of theMikasa Formation (ColI. M. Noda and T.
Takahashi); JG. H2955 (Fig. 2-2), somewhat
distorted left valve, from loco R5400j5, Kanajirizawa, Member Mh of the Obira area (ColI. M.
Noda, S. Toshimitsu and T. Matsumoto).
WE. P020Y (Fig. 3-A) and WE. P021Y (Fig.
3-B), left and right valves, with deficient apex ofbeak and incomplete ventral part, in one and thesame nodule from loco Y070161a84, Hakkinzawa, middle part of Member My7, Oyubari area
(ColI. A. Asai, Y. Tanaka and H. Hirano). They
are to be called Inoceramus cf. pennatulus.
Description:-Shell normally large, inequivalve
and inequilateral. Left valve more convex in the

380

TatsuroMATSUMOTO etal.

Figure 1. Inoceramus pennatu!us Pergam ent.
I: GK. H89 (RV) , drift in a gully between lac. T831 and T832, derived from Member lIb, Chirashinai ,mid-valley of the Teshio (Call . T. Matsumoto); lateral (a) and anterior (b) views. Scale bar = 20 mm . 2: WE.
POOl Y (RV) , from lac. Y250079 , Member My7 , Hikage-zawa , Shiyubari-Oyubari area (Call. T. Sugiyam a) ;M. Noda photos.
lateral view . Scale bar = 20 mm .

umbonal part than the right , with prominent
umbo projecting over the hinge line and curved
inward and forward . Anterior part steep andnearly vertical , abruptly falling down from themain part of the flank , which in turn inclines
gradually to the broad and flat posterior wing.
Hinge line long, being more than a half of theshell length. Axis of growth oblique and nearlystraight or broadly arcuate (i. e. convex forward) ,forming an angle (0) of 65° to 75 ° with thehinge line in late stages.
Surface of the shell ornamented with concentric subcostae and ribs of irregular strengthand density. In the young stage ribs are weak ,853. Cenomanian inoceramids from Japan - III

381

--- -

---

,,
1

Figure 2. Inoceramus pennatulus Pergamen t.
1: JG . H2492 (LV) from loe. Ik1040, Member lIe of the Mikasa Formation, River Ikushumbets (Coli. M. Nod aand T. Takahashi) ; lateral view, x 4 /5.2: JG . H2955 (LV) from lo e. R5400j5 , Kanajiri-zawa, Member Mh , Obiraarea (ColI. M. Noda , S. Toshimitsu and T. Matsumoto); lateral (a) and posterior (b) view s, x 1. M. Noda photos.

but dense and fme lirae or rings may be welldiscernible. The subcostae and ribs are generally
distinct in the middle growth-stage and lowered
and weakened later. Irregularity may also occur
in the course of the ribs , showing sometimes
bifurcation , intercalation or disappearance . Flat

posterior wing nearly smooth or with much
weakened ribs. The ribs generally form an asymmetric curvature , being long and broadly arcuate
posteriorly , more rounded ventrally and rather
abruptly truncated by the nearly straight or
broadly arcuate anterior margin.

Tatsuro MA TSUMOTO et al.

382

T ab le l. Measurement s on selected specimens of I. pennatu!usSpecimens

ll h

h

H

L

L/H

Holot ype (L) '

110

67

.61

104

70

.67

G1\.. H89 (R)

137

101
107

.74
. 77

142

108

.76

141

109

. 77

.76

184

127

. 69

WE. POOl Y (R) 139
JG . H2942 (L) ~165 ~125

b

b/ h

48

.35

~43

. 26

s

sl l

a

~

r

0

~43

. 64 ~100 ' ~ 53' ~125 °

~55

.54

90 '

~52

.49

103 '

~80

.64

~91 '

135

~130 '

70'

~70 '

0

70

0

~ 75 '~74 °• Measured on the photograph (Pergament, 1966, pI. 5, fig . 1) at the midd le growth-stage. - Measurement on a
res tored o utline.

Measurements: -See Tabl e 1.
Comparison and discussion :-We have given
above a free translation of diagnosis written in
Russian by Pergament (1 966, p. 35) and further
supplemented it from our observation.
As the shell is large, the fossil specimens areoften incomplete . This is true even for the types
illustrated by Pergament. Of his nine specimens,
eight are more or less incomplete left valves and asingle right valve which is deficient for lack of
the ventral half. This is supplemented by ourbetter preserved specimens , although they are by
no means complete.
Noda and Matsumoto (1976) once interpreted
this species too much broadly. We now define
this species strictly, regarding "I. pennatulusinterjectus" Pergament (1966 , p. 37, pI. 8, fig. 1;
pI. 9 , figs. 1-3) as distinct from I. pennatulus,

Figure 3. Inoceramus cf. 1. pennatu!us Pergamen t.
A: WE. P020Y (LY); B: WE. P021 Y (RY) . Both in one and the same nodule fro m lac. Y070 161 a84 ,Hakkin-zawa, Member My? , Oyubari area (Ca ll. Y. Tanaka , H. Hirano and A. Asai) ; lateral view, x 2/ 3. Scale bar= 30mm.
A. Asai photos.

853. Cenomanian inoceramids from Japan - III
because the posterior wing is narrow in I.
interjectus and the axis of growth is more
oblique, with narrower angle of inclination thanthat of I. pennatulus. The two specimens whichwere illustrated as examples of I. pennatulus by
Noda and Matsumoto (1976, Cr-31, figs. 4, 5)
should be revised to I. interjectus. Sornay (1978,
p. 508) has already remarked that "I. pennatulusinterjectus" is closely allied to I. conicus
Gueranger (see Sornay, 1978, pI. 1, figs. 1-3).
This implies that I. interjectus is independent of
I. pennatulus.
Occurrence:-Loc. Y250079, Hikage-zawa,
Oyubari area, Member My7 by Hirano et al.
(1988); this locality is 170m southeast of loco
Y168, lower part of Member Mf by Matsumoto(1942). From near locs. T831 and T832 at
Chirashinai, 4km southeast of Saku, in thetransverse mid-valley of the River Teshio, lower
part of Member lIb by Matsumoto (1942). These
two localities are in rather lower part of the
Cenomanian but may be referred to the lower
Middle Cenomanian on the international scale.
Loc. Y070161a84 of the Hakkin-zawa (Oyubariarea), where examples of I. cf. pennatulus wereobtained, is assigned to the middle part ofMember My7, Middle Cenomanian, because of
the occurrence of Acanthoceras takahashii
Matsumotofromthenearbylocality(Y070 163b83). In all the above cases the hostrock is dark grey mudstone or sandy siltstone.
Loc. Ik1040 on the River Ikushumbets, 470m
downstream from the Katsura-zawa Dam,
Member IIc, i.e. middle part of the Mikasa
Formation; loco R5400j5, Kanajiri-zawa, Member
Mh of the Obira area. These two localities areevidently ascribed to the Upper Cenomanian.
The host rock is again dark grey mudstone.

Inoceramus reduncus Pergament, 1966
Figures 4; 5-2-4; 61966 Inoceramus reduncus Pergament, p. 40, pI. 16,
figs. 1, 2; pI. 17, figs. 1, 2; pI. 18, figs. 1-3.
1966 Inoceramus reduncus Pergament subsp. nov. ?, p.
42, pI. 19, figs. 1,3.

Holotype:-No. 9-3580-54,

designated and383

illustrated by Pergament (1966, p. 40, pI. 16, fig.
1), from the Ginterobskaya Group (Cenomanian)of the Ugolnaya Bay area in the northeasternpart of the Koryak Mountains.
Material:-GK. H25 (Figs. 4A-C), somewhat
displaced both valves, i.e. internal mould of left
valve, with shell attached partially, and internaland external moulds of right valve with inner andouter shell layers attached respectively, from loco
T7b, Rubeshibe-zawa, upper part of Member lIbof the Abeshinai-Saku area (Matsumoto, 1942),
ColI. T. Matsumoto; GK. HlO146 (Fig. 5-2),
right valve and partly cropping out left valve,with shell preserved for the major part, from loco
R5243, right side of the River Obirashibe,
Member Mh of the Obira area (Tanabe et al.,1977), ColI. T. Matsumoto; GK. HlO147 (Fig.
5-3), left valve with shell layer and hinge plate
preserved partly but lacking in posterior part,from loco R5407c, Kanajiri-zawa, Member Mh of
the Obira area (Ditto, 1977), Coli. T. Matsumoto;JG. H2960 (Figs. 6A, B), internal mould ofleft valve, with shell preserved partly, from loco
R5400T, Kanajiri-zawa, Member Mh of the
Obira area (Ditto, 1977), ColI. M. Noda, S.
Toshimitsu and T. Matsumoto.
GK. H8267 (Fig. 5-4), umbonal part of left
valve, partly composite, internal mould, fromloco RlOO on Highway 239, between Soeushinai
and Kotanbetsu, 800m west of the Kiritachi Pass,
Member My5 of Hashimoto et al. (1965), ColI. T.
Matsumoto and H. Okada. This is I. cf. reduncus(see discussion below).
Description: -Shell medium-sized normally,
but occasionally large, inequilateral and highly
inequivalve. Left valve inflated, with muchconvex and prominent umbonal part andmarkedly incurved beak, as the specific nametells. The convex curvature of the left valve
profile along the axis of growth changes at the
beginning of the middle stage where concentric
constriction occurs normally; another constriction may occur later. Right valve less inflated
than the left and the change of convexity on the
profile along the axis of growth may be
expressed weakly by the concentric constriction
or deeper interspace of ribs at the position

384

Tatsuro MATSUMOTO et al.

Figure 4. Inoceramus redullcus Pergament.
A-C: GK. H25 (BV) from loco T7b, Rubeshibe-zawa , upper part of Member lIb , Abeshinai-Saku area (Co lI. T .
Matsumoto) ; lateral (A) and posteri or (B) views of LV ; lateral view of RV (C) , xL
M. Noda photos.

corresponding to that of the left valve. Anteriorside abruptly bent from the disk and nearly
perpendicular to the plane of the valves, withslightly concave anterodorsal part Posterior
wing-like area narrow and not clearly defined, forit passes gradually from the convex main part of

853. Cenomanian inoceramids from Japan - III
disk.
Outline of shell roughly asymmetric-triangulardorsally and broadly sub rounded ventrally. H
somewhat but not much larger than L, with L/Habou t 0.7 in LV and 0.8 in RV on the average.
Anterior margin roughly straight, with slightanterodorsal concavity and anteroventral gentle
convexity; ventral margin more or less broadly
rounded, passing to gently convex to nearlystraight posterior margin. Hinge line of moderate
length. Anterior hinge angle (a) nearly 90° in the
typical form but may be somewhat larger inothers. Posterior hinge angle ("y) obtuse and mayvary with growth or between individuals.
Shell of early growth-stage up to the first
constriction nearly smooth or with faintconcentric undulations; but dense and fineconcentric lines on the surface of outer shelllayer or corresponding rings on that of inner shelllayer may be well discernible. In the middle andlate growth-stages ribs and subcostae develop
distinctly, being somewhat irregular in strengthand density. Minor concentric rings may be
combined with them when preservation isfavourable. Concentric constrictions or deeper
interspaces occur at periodic intervals, asmentioned above. In the gerontic stage theornament may weaken.
Measurements:-See Table 2.
Comparison and discussion: - The specimen
briefly
described
andillustrated under
Inoceramus koriakensis Terechova (in Verechagin
et aI., 1965, p. 36, pI. 23, fig. I), which was
acquired from the Ginterobaskaya Group in thearea of the Ugoinaya Bay, looks very similar tothe holotype and other typical examples of thisspecies. Therefore, we would suggest that 1.
reduncus Pergament, 1966 could be a junior

385

synonym of 1. korjakensis Terechova. 1965. As
we have not studied the actual specimens
themselves in USSR and as Pergament (1966)
described 1. reduncus on many specimens withfine illustrations, we use in this paper the specificname 1. reduncus provisionally. We expect that apalaeontologist in the USSR would give adefinite answer to our question.
The left valve of GK. H25 (Figs. 4A, B) isessentially similar to that of a para typeillustrated by Pergament (1966, pI. 18, fig. 1)
and the right valve of GK. H25 (Fig. 4C), if its
deficient beak is adequately restored, resembles
that of another paratype (Pergament, 1966, pI.
16, fig. 2). A small specimen, GK. H8267 (Figs.
5-4a, b) is probably the inflated umbonal part,showing a constriction at the beginning of the
middle aged part; its later part must have been
destroyed away. It is comparable with the one
illustrated by Pergament (1966, pI. 18, fig. 2a).
GK. HlO147 (Fig. 5-3), secondarily compressed
left valve with apparently broad disk in younger
half, resembles again the one figured by
Pergament (1966, pI. 17, fig. 2). The ligamental
structure of the hinge plate is observable in thetwo specimens, but the conjugated right valve is
not preserved in ours.
The right valve of GK. HlO146 (Fig. 5-2) isquite similar to the form represented by two
specimens of Pergament (1966, pI. 19, figs. 1,3).
The latter was regarded with a query as an
unnamed subspecies of 1. reduncus by Pergamenton account of some difference in the ontogenetic
change of L/H, but we do not think the
difference so significant as deserving subspecificseparation. Our example occurs in the same
Member Mh of the Obira area as the typicalform. Therefore, it is regarded as showing a

Table 2. Measurements on selected specimens of I. reduncus
SpecimenshHolotype (L)
(R)

75
58

GK. H25 (L)GK. HI0146 (R)]G. H2960 (L)

-90
73
60

H

L

L/H

b

b/h

.73 -91.86 -71

-63
-53

.69
.75

22

.29

98 -68
75
63
61
68

.70
.84
.89

33
25
18

llh
55
50

78 .8760 .8262 1. 03

s

sl I

a

36

.68

93

.37 -46
.33
36
34
.26

.59
.60
.56

r

f3
_70

0

_50

0

135

0

98
94°
0

89°

135
148
159

ii

0

0

0

0

45

0

45
50-70°
63°0Tatsuro MA TS UMO TO et al.
386

853.

Cenomanian inoceramids from Japan - III

387

Figure 6. Inoceramus reduncus Pergament.
JG . H2960 (LV) from loc. R5400T , Kanajiri-zawa, Member Mh , Obira area (Coil. M. Noda , S. Toshimitsu and T .
M. Noda photos.
Mat sumoto) ; lateral (A) and anterior (B) views , x I.

variation within this species. Incidentally the left
valve of GK. HI0146 is somewhat displaced
toward the venter , with beak concealed in therock matrix (see Fig. 5-2b) .
The affinity of I. reduncus with I. gradilis
Pergament (see Matsumoto et at. , 1987 , p. 155)
should be considered. The latter may be a
specialized offshoot from the former or the two
species may have been derived from a common
ancestor. We presume that their origin could bethought in such species as I. tenuis Mantell andeventually in Birostrina concentrica (Parkinson)on the ground of change in morphologic characters and also stratigraphic occurrence.
Occurrence: - The representative specimens
described above are from loco T7b, Rubeshibezawa , upper part of Member lIb of theAbeshinai-Saku area (Matsumoto, 1942) ; locs.
R5243 , R5400T and R5407c , all in Member Mhof the Obira area (Tanaka, 1963 ; Tanabe et at.,~ Figure 5.
Inoceramus ginterensis Pergament (1) and Inoceramus reduncus Pergament (2-4).
1: GK. H8260 (RV) from loc. R5408, Kanajiri-zawa, Member Mh , Obir a area (Coil. T. Matsumoto). Internal (a)and external (b) mould s, x l. 2: GK. H 10146 (RV) from loc. RS243 , the River Obirashibe, Member Mh , Obiraarea (Coli. T. Matsumoto) ; lateral (a) and anterior (b) views, x l. A portion of LV shown on the basal surface of
2b . 3: GK. HIOI47 (LV) from loc. RS407c, Kanajiri-zawa, Member Mh, Obira area (Coil. T. Matsumoto); lateral(a), anterior (b) and dorsal (c) view s, x 1. 4: GK. H8267 (young part of LV) from loc. RIOO , 800 m west of the
Kiritachi Pass on Highway 239 between Soeushinai and Kotanbetsu (Co il. T. Matsumoto and H. Okada); lateral(a) and anterior (b) views, x 1. It is b etter to call this I. cf. reduncus.
M. Noda photo s.

Tatsuro MATSUMOTO et al.

3881977). These are in the Teshio Mountains ofnorthwestern Hokkaido and referred to theupper part of the Cenomanian. A comparable
specimen is from loco RlOO on Highway 239
across the Teshio Mountains between Soeushinai
and Kotanbetsu, Member My5 of Hashimoto etal. (1965), which is referred to the upper part of
the Middle Cenomanian.

Inoceramus ginterensis Pergament, 1966
Fil,\ure 5-1; 7-1-4; 8-1-5; 91966 Inoceramus ginterensis Pergament, p. 50, pI. 27,
figs. 1, 2; pI. 28, figs. 1, 2; pI. 29, fig. l.
1977 Inoceramus ginterensis Pergament; Cobban, p.
15, pI. 19, fig. 3.
1977 Inoceramus ginterensis Pergament; Kauffman andPowell, p. 64, pI. 3, figs. 1-4, 6; pI. 4, figs.
1,3-6; pI. 5, figs. 2, 3,5; pI. 7, fig. 4.
1983 Inoceramus ginterensis Pergament; Cobban, p. 5,
pI. 2, figs. 17, 18.

Holotype: -No. 16p-3580-87, designated andfigured by Pergament (1966, p. 50, pI. 27, fig. 1)(copied by Kauffman and Powell, 1977, pI. 4,
fig. 4), from the Cenomanian in the Ugoinaya
Bay area in the northeastern part of the Koryak
Mountains.
Material:-GK. H28A (BV) (Figs. 7-4a-c), B(LV of BV)(Fig. 7 -3) and C (LV)(Figs. 7 -2a-b),internal moulds with inner shell layer adhering toconsiderable parts; also fragmentary pieces of external moulds, with outer shell layer adhering,from loco T843, the Chirashinai, a small tributary
to the middle course of the River Teshio, unit
lIb-c (~) of Matsumoto (1942, p. 189), ColI. T.
Matsumoto; GK. H417A (RV) (Fig. 8-5) and B(LV), internal moulds and part of external mouldwith shell adhering, from loco Y508, theTengu-zawa, Member lIm (Matsumoto, 1942, p.
230),
Shiyubari-Oyubari
area,
CoIl.
T.
Matsumoto; GK. H8260A (RV) (Fig. 5-1a),internal mould with inner shell layer adhering,and GK. H8260B (Fig. 5-1b), external mould ofA, with outer shell layer adhering, from loco
R5408, Kanajiri-zawa, Member Mh of the Obiraarea (Tanabe et at., 1977, fig. 7), ColI. T.
Matsumoto.

WE. P002Y (LV) (Fig. 7-1), composite
internal mould from loco Y070135m82, lower
part of Member My7, ColI. H. Hirano, N. Katoand Y. Nakanishi; WE. P037Y (RV) (Fig. 8-1),
internal mould, with uncleaned shell adhering on
the umbonal part, from loco Y070135f82, lower
part of Member My7, ColI. H. Hirano and Y.
Nakanishi; WE. P023Y (Fig. 8-4), large but
deficient LV, with shell adhering for major part,from loco Y070l35k84, lower part of Member
My7, CoIl. A. Asai; WE. P024Y (PV) (Fig. 8-2),
internal mould from loco Y070137e83, lower
part of Member My7, CoIl. A. Takagi and H.
Sekine; WE. P022Y (RV) (Fig. 8-3), composite
internal mould from loco Y070119, upper part ofMember My7, ColI. H. Ando; WE. P040Y (Fig.
9), large right valve from loco Y070137j83,
Member My7, Coli. A. Takagi and H. Sekine; allon the routes (stream and forestry roads) along
the Hakkin-zawa, Oyubari area. It may be betterto call the last one I aff. I. ginterensis (seediscussion below).
Description: -Shell normally medium-sized
and weakly inequivalve; the left valve somewhat,
but not much, more convex and slightly higher
than the right, with its beak slightly projecting
over hinge line.
Outline of shell subovate to subrectangular,
fairly higher than long, axis of growth forming
angle (8) of 60° to 65° with the hinge line;
anterior margin nearly straight, ventral margin
asymmetrically and moderately rounded, passing
to broadly arcuate (convexly) posterior margin,
which forms an obtuse angle ('y) of about 135°with the fairly long hinge line.
Valves gently inflated, with the maximum
convexity somewhat ventrally from the umbonalpart, passing gradually to the flat, postero-dorsal
wing-like part. Anterior side steep, abruptly bent
from the disk and may be somewhat concave
near the umbonal part.
Shell in the early growth-stage ornamented
with faint concentric undulations and fine anddense concentric rings or lirae which may bediscernible under favourable condition of preservation. In the middle to late growth-stages
concentric ribs and subcostae develop with

853. Cenomanian inoceramids from Japan - III

Figure 7. Inoceramus ginterensis Pergament.
1: WE. P002Y (LV), from loc o Y070135m82 , Hakkin-zawa, lower part of Member My? , Oyubari area (ColI. H.
Hirano, N. Kato and Y. Nakanishi). A. Asai photo. 2: GK. H28C (LV), lateral (a) and anterior (b) view s, x 1; 3:
GK. H28B (LV), lateral view , xl; 4 : GK. H28A (BV) , right (a) and left (b) lateral and anterior (c) views, x 1.
The above three (2 - 4) in one and the same nodule from mudstone at lo co T843 , Chirashinai, Member Ilb-c
Ci3 facies) , mid-valley of the Teshio (ColI. T. Matsumoto).
M. Noda photos.

389

390

Tatsuro MA TSUMOTO et al.

Figure 8. Inoceramus ginterensis Perga ment.
1: WE. P037Y (LV) from loc o Y070135f82 (Co lI. Y. Nakani shi and H. Hirano); 2: WE. P024Y (RY) from loc.
Y070137e83 (Coli. A. Takagi and H. Sekine); 3: WE. P02 2Y (RY) from loc. Y070119 (Co lI. H. Ando) ; 4: WE .
P023Y (LY) from loco Y070135k84 (ColI. A. Asai). Th ese four (1 - 4) on the Hakkin-zawa route , Member My7(1,2,4 lower part; 3 upper part) , Oyubari area, x 2/ 3. A. Asai photo s. 5: GK. H417A (RY) from loc o Y508 ,Tengu-zawa , Member 11m of the Shiyubari-Oyubari area (Co lI. T. Mat sumoto), x I.
M. Noda photo .

853. Cenomanian inoceramids from Japan - III

391

Table 3. Measurements on selected specimens of I. ginterensis

Specimens
Ho!otype (L)GK. H28A (L)(R)
GK. H28B (L)GK. H417A(R)WE. P002Y (L)WE. P022Y (R)WE. P023Y (L)WE. P024Y (R)

h

81
48
50
56
90
50
135
120
86

60
36
36
44
73
38
93
88
66

l/h

H

.74
.75

86 -5854 -3857
40
60
45
98 -675236
140 -100
132
96
97
74

.72
.79
.81
.76
.69
.73
.77

L

L/H

b

23
.67
.70
17
12
.70
.75
15
.68 -21.69 -14.71.73 -30
19.76

irregular intensity and density, showing sometimes bifurcation or intercalation; interspaces
may be occasionally deep; fine concentric lines
or rings may be discernible where shell isfavourably preserved.
Measurements:-See Table 3.
Comparison and discussion:-In addition to
Pergament (1966), Kauffman and Powell (1977)
described at full length the diagnostic charactersof this species and gave remarks on its
relationships with other species. The specimensfrom Oklahoma, figured by Kauffman andPowell (1977), are mostly typical examples,
except for a small specimen (Ditto, pI. 1, fig. 2)
which shows "abnormally close, regular juvenile
rugae." Cobban (1977, 1983) also reported theoccurrence of this species in other areas of
the Interior Province.
The specimens from Hokkaido figured in thispaper are also typical, although there may beslight or unimportant differences which can beinterpreted as being within the extent ofvariation or modification caused by the mode ofpreservation.
On the ground of above observations we canstate that no significant distinctions have been
noticed between populations of such separate
provinces as the Koryak-Kamchatka belt and S.
Sakhalin-Hokkaido or the Northwest Pacific andthe Interior Province of North America. Should
this be warranted, 1. ginterensis would be foundmore extensively. A probable example from
Madagascar (=1. [lavas Sornay, 1965, pI. A, fig. 1b/hs.28 -36
.35
26
26
.24
30
.27
.23 -52.28 -28-56.25 -68.22 -50

s/t

93°
86°
86°
.68 _85°96°.76.78 _83°
.60

.72
.72

.60
.77
.76

(3

r

0

67°
53°

138°
130°
130°
138°
138°
135°
_128°

61°
57°
5r
60°
60°
56°
65°
60°
60°

a

93°
85°
85°

65°

115°
135°

non fig. 2) as pOinted out by Kauffman andPowell (1977, p. 66) could be evaluated on thisaccount.
WE. P022Y (Fig. 8-3) from the upper part ofMember My7 of the Oyubari area is quite similar
to WE. P024Y (Fig. 8-2) from the lower part ofthe same member. This may suggest that there isno marked distinction between the forms fromdifferent stratigraphic. positions. Member My7 of
Hirano et ai., (1988) is mainly referred to Middle
Cenomanian, because Turrilites acutus Passy and
Acanthoceras takahashii Matsumoto occur in the
middle part of the member. Although the schemeof stratigraphic division of Matsumoto (1942) is
not the same as that of Hirano et al. (1988), GK.
H417 is from Member lIm of the Shiyubari area
by Matsumoto (1942), which is the uppermost
unit of the Cenomanian sequence, and GK.
H28A-C came also from the upper part of the
Cenomanian in the Teshio Mountains. There is,however, no significant difference between the
GK specimens from the upper part of the
Cenomanian and the WE specimens from the
middle part of the same stage.
The above may be a tentative result frombetter preserved specimens at our disposal. It
would be desirable to examine the subject on
population samples of enough number. In thispaper we exclude atypical or doubtful specimens
described under 1. ginterensis by Pergament(1966, pI. 25, fig. 5; pI. 26, fig. 1; pI. 30, fig. 1)
and also 1. aff. ginterensis of Pergament (1966, p.
52, pI. 32, fig. 2; pI. 33, figs. 2, 3; pI. 34, figs. 1,

392

Tatsuro MATSUMOTO et al.

Figure 9. Inoc eramus aff. 1. ginterensis Pergamen t.
WE . P040Y (R V) from lac. Y070137j83 , Hakkin-zawa, Member My7, Oyubari area (Call. A . Takagi and H.
Sekine) , x 1. Scale bar = lOmm .
A. Asai photos.

2) from I. ginterensis in our proper sense.
Likewise there are several doubtful specimens in
our collection which deviate from the typicalform of I. ginterensis. WE. P040Y (Fig. 9) may
be an example of them , since it resembles theone illustrated by Pergament (1966, pI. 34, fig.

853. Cenomanian inoceramids from Japan - III2) under I. aff. ginterensis. How they should betreated and what they mean remain unsettled.
They need further study on sufficient material.
As to the phylogenie origin of I. ginterensis
we recall such a species as I. takahashii
Matsumoto et Noda (1986, p. 414, pI. 84, fig. 1),
from the Lower Cenomanian of Hokkaido, in thesubrectangular outline, weakly inequivalve state,less inflated valves and irregular ribbing. I.
ginterensis is more oblique, with more roundedventral margin and has stronger major ribs in the
middle to late growth-stages in addition to theweaker subcostae.
I. heinzi Sornay (1965, p. 7, pI. B, fig. 4) (also
Matsumoto and Noda, 1986, p. 415, pI. 83, fig.
4; pI. 85, figs. 2, 3), from near the CenomanianTuronian boundary, is somewhat allied to I.
ginterensis but has more projected and morestrongly incurved left umbo, shorter hinge line
and more oblique axis of growth.
Occurrence:-The described specimens of thetypieal form are from (1) loco T843 in the upper
reaches of the Chirashinai, a tributary to the
middle course of the River Teshio, mudstone in
the lower part of Unit IIb-c (~) of Matsumoto(1942), upper part of the Cenomanian,
Abeshinai-Saku area, Teshio Mountains; (2) loco
Y508, above the first water-fall of the Tenguzawa, mudstone of Member lIm of Matsumoto(1942), upper part of the Cenomanian,Shiyubari-Oyubari area; (3) loco R5408, Kanajirizawa, mudstone of Member Mh of Tanaka(1963), upper part of the Cenomanian of the
Obira area; (4) locs. Y070135 and Y070137,
mudstone in the lower part of Member My7 of
Hirano et al. (1988), Hakkin-zawa, Middle
Cenomanian; (5) loco Y070119, mudstone in theupper part of Member My7, Hakkin-zawa,
Oyubari area.
Summary
As a summary the species described in thispaper are listed below with indication of theirknown age and area in Hokkaido.
I. pennatulus Pergament: Upper Cenomanianof the lkushumbets and Obira areas; com-

393

paratively lower part of the Cenomanian in the
Oyubari and Saku areas, which may be lower
Middle Cenomanian on the international scale.
I. reduncus Pergament: Upper Cenomanian of
the Saku and Obira areas; comparable form (I. cf.
reduncus) in the upper Middle Cenomanian of
the Kotanbetsu area.
I. ginterensis Pergament: Upper Cenomanianof the Saku, Obira and Shiyubari areas; also
Middle Cenomanian of the Oyubari area.
To sum up, the three species show fairly longranges in stratigraphic occurrence. This is similar
to, if not quite identical with, the situation in the
Pacific region of the USSR (Pergament, 1966,
figs. 4, 6, 8, 10; 1977, p. 91; 1981, table 5). The
Zone of I. pennatulus in the Upper Cenomanianof Japan (Matsumoto, 1977) is merely a local
zone in the lkushumbets area. The same named
zone is set in the middle part of the Cenomanianby Pergament, but I. pennatulus occurs also in
the upper part of the stage in the Pacific USSR.
Acknowledgements
We are much indebted to many persons who
worked with us or with one of us in thestratigraphie investigations of the Saku,
Kotanbetsu, Obira, lkushumbets and ShiyubariOyubari areas, where inoceramids were collectedalong with other fossils. Their names arerecorded in the item Material of the descriptionsof species as well as in the Introduction.
Thanks are also due to the authorities of theForestry Bureau Branch Offices of the respective
districts for permission and help to us to dogeological field works in the areas of their care.
Hirano received financial support of theGrant-in-Aid for Individual Research of the
Ministry of Education, Science and Culture (C,No. 57540464 for 1982 and No. 61540577 for1986 and 1987), Research Fund of theGeographical Society of Tokyo for 1983 and1987 and Grant-in-Aid for Fundamental Science
Research of Waseda University (No. 58A-29
for 1983, No. 59A-56 for 1984, No. 60A-46
for 1985, No. 61A-69 for 1986 and No.
62A-79 for 1987) for this work (Parts II & III)Tatsuro MATSUMOTO et al.

394

and related stratigraphic investigations by Hirano
et al.

References cited
Cebban, W. A. (1977): Characteristic marine melluscan
fessils frem the Daketa Sandstene and intertengued Mances Shale, west-central New Mexice.
U. S. Geol. Surv. Prof Pap., 1009, p. 1-30, pIs.
1-21.
_ _ (1983): Melluscan fessil recerd from the nertheastern part .of the Upper Cretaceeus seaway,
Western Interier. Ibid, 1253-A, p. 1-25, pIs.
1-15.
Hashimete, W., Nagae, S. and Kanne, S. (1958):
Explanatery text .of the Geel. Map .of Japan,
Seeushinai sheet, Scale 1:50000. Hekkaide
Develep. Agency. (1).
Hirane, H., Ande, H., Hirakawa, M., Merita, R. andIshikawa, T. (1981): Biestratigraphic study .of theCretaceous System in the Oyubari area, Hekkaide.
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Waseda Univ., Ser. Bioi. & Geol., vel. 30, p. 33-45(J).

Matsumete, T. and Tanabe, K. (1977):
Mid-Cretaceeus stratigraphy .of the Oyubari area,
central Hekkaide. Palaeont. Soc. Japan, Spec. Pap.,
ne. 21, p. 1-10.
- - , Takizawa, K. and Tsuchida, S. (1988): Biestratigraphy .of the Upper Cretaceeus in the Oyubariarea, central Hekkaide. Bull. Sci. Engin. Res. Lab.
Waseda Univ. (in preparatien).
Kauffman, E. G. and Pewell, J. D. (1977): Paleentelegy. In Kauffman, E. G., Hattin, D. E. and Pewell,J. D. (1977): Stratigraphic, paleentelegic andpaleeenvirenmental analysis .of the Upper
Cretaceeus reeks .of Cimmarren Ceunty, nerthwestern Oklahema. Mem Geol. Soc. A mer. , vel.
149, p. 47-150, pIs. 1-12.
Matsumete, T. (1942): Fundamentals in the Cretaceeus
stratigraphy .of Japan. Part I. Mem. Fac. Sci.,
Kyushu Imp. Univ., Ser. D, Geol., vel. 1, ne. 3, p.
130-280, maps in pIs. 5 -19.
- - (1965): A menegraph .of the Cellign.oniceratidae
fr.om H.okkaid.o. Part I, Mem Fac. Sci., Kyushu
Univ., ser. D, Ge.ol., vel. 16, n.o. 1, p. 1-80, pIs.
1-18.
--(1977): Z.onal c.orrelati.on .of the Upper Cretace.ous
in Japan. Palaeollt. Soc. Japan, Spec. Pap., n.o. 21,
p.63-74.
Asai, A. and Hiran.o, H. (1987): S.ome
in.oceramids (Bivalvia) fr.om the Cenemanian(Cretace.ous) .of Japan. Part II. Three species from

Hekkaide, well kn.own abr.oad but hitherte
underdescribed in Japan. Trans. Proc. Palaeollt.
Soc. Japan, N. S., n.o. 147, p. 146-164,8 fIgs.
_ _ and N.oda, M. (1986): Ditto. - I. Newer little
kn.own f.our species fr.om H.okkaid.o and Kyushu.
Ibid., n.o. 143, p. 409-421, pIs. 81-85.
_ _ and Okada, H. (1973): Saku F.ormati.on .of the
Yezo ge.osyncline. Sci. Repts. Dept. Geol., Kyushu
Univ., vel. 11, n.o. 2, p. 275-309 (1).
_ _ , _ _ , Hiran.o, H. and Tanabe, K. (1978):Mid-Cretace.ous bi.ostratigraphic successi.on in
H.okkaid.o. In Matsum.ot.o, T. (c.ompil.): MidCretace.ous zenati.on in Japan. Ann. Mus. d'Hist.
Nat. Nice, vel. 4 (fer 1986), pt. xxxiii, p. 1-23.
N.oda, M. and Matsum.ot.o, T. (1976): Mes.oz.oic
m.olluscan f.ossils .of Japan 4 (Cretace.ous
Inoceramus). Atlas of Japanese Fossils, n.o. 45,
Cr-31-Cr-36, Tsukiji-Sh.okan, Teky.o (1).
Pergament, M. A. (1966): Z.onal stratigraphy andInoceramus .of the l.owerm.ost Upper Cretace.ous .of
the Pacific c.oast .of the USSR. Trans. A cad. Sci.
USSR, Geol. Illst., vel. 146,83, p. 36 pIs. (R).
- - (1977): Stratigraphy and c.orrelati.on .of MidCretace.ous .of the USSR Pacific regi.ons. Palaeont.
Soc. Japan, Spec. Pap., n.o. 21, p. 85-95.
--(1981): Pacific regi.ons .of the USSR. In Reyment,
R. A. and Bengts.on, P.: Aspects of Mid-Cretaceous
Regional Geology, Academic Press, L.ond.on, p.
69-102.
S.omay, J. (1965): La faune d'in.oceramides duCen.omanien et du Turenien inferieur du sud-.ouest
de Madagascar. Ann. PaIeont., Invertebres, vel. 51,
p. 1-18, pIs. A-C.
--(1978): Precissiens pale.ont.ol.ogiques et stratigraphiques sur divers inecerames Cen.omaniens et,en particulier, sur ceux de la Sarthe fIgures par E.
Gueranger en 1867. Geobios, vel. 11, n.o. 4, p.
505-515, 2 pIs.
Tanabe, K., Hiran.o, H., Matsum.ot.o, T. and Miyata, Y.
(1977): Stratigraphy .of the Upper Cretace.ous
dep.osits in the Obira area, n.orthwestern H.okkaide.
Sci. Repts. Dept. Geol., Kyushu Univ., vel. 12, n.o.
3, p. 181-202 (J).
Tanaka, K. (1963): A study en the Cretaceeus
sedimentatien in H.okkaid.o, Japan. Rept. Geol.
Surv. Japan, n.o. 197,122, p_ 2 maps, 3 pIs.
Verechagin, V. N., Kinas.ov, V. P., Parakech.ov, K. V.
and Terecheva, G. P. (1965): Field Atlas of theCretaceous Faunae from the Northeast USSR,
Magadin, 215, p. 74 pis. (R).
(1) : inJapanese
explanatien.
(R): in Russian.

with

Englishabstract

.or

brief

853. Cenomanian inoceramids from Japan - III

395

Abeshinai 7 -" -/ -j- -1 (~:ifZ;t P'.J). Chirashinai 7- '7 -/ -j- -1 (~.& it; P'.J). Fukagochi ~ ii'J P'.J •Fukuoka/lITJ. Hakkin-zawa B ~ IR. Hikage-zawa ElIl.f11IR. Hokkaido ~I: $: ~. Ikushumbets(=Ikushunbetsu) ~~5JIj. Kanajiri-zawa ~m:IR. Katsura-zawa i'11R. Kiritachi fJJlL. Kotanbetsut; f]- 5J1j. Mikasa .=: 11. Obira 'j,:ifZ. Obirashibe 'j':ifZ ~. Oita :*: 7t. Oyubari :*:!7 iji .
Rubeshibe-zawa Iv -" -/-"IR. Saku fti:~. Shiyubari (Shuyubari) -/ =z. - / { ! I (::t!7iji). Soeushinai
rmlfP'.J. Tengu-zawa J(friJlR. Teshio J(J;i'K. Tirashinai = Chirashinai. Tokyo :l!tJjC Waseda !f!ft1i

m

133.

<

*fBB!I~H:!/ "?=-7 :/1l~iI1-1 / -I! '7 A7--ill o ~tim;;tc:ifZtf.ttl!.~~':'J: iI1-t-:, 3 ~ : ::'O)fflfU:X:""('
ft:; :ilO);;tc:ifZtf.ttl!.lKt.:lt""('t.: ~1:$:~iI' 6 t J: iI1-t -:,?XO) 3 ~~~cill\( Lt.:o<<( 1) Inoceramus pennatulus Pergament : *~-Cti:ti~,*mt. ti:mtO)mtlJHt~lli-t -:, ; mto)lW
6'<;'cp~~. t&.$b~$~' ; }jj]lt~mlliJ""(';l;-:,o Pergament (1966)ftti:mtO)~{9IjHUl(~ff- Lt.:b~. timtO)(jl]tJ\z. Lb, ') t.:o ~1:$:~O)ft*4ft::. h ~flIlJ.e. -t -:, 0 ~~5J1j c ::t \:::' '7 -/ -" ""('ft -I! / "? =7:/ 1:$~':'!fH&Ef:Jta~. :*:!7~N:~fti:~ ""('ltcp$0)~~rO):1Jb'6 til1 L -n,-:,o(2) I, reduncus Pergament : cp~~~*~""(,ti:..t:i~~mt ; ti:mtftIW6'<;'b~~ L <. mtlJH~IHt~
lli L P'.Jlltr:1J~.:.iJ!ll <}ffi illl-t -:, Rltffi:O).ifEcp-C1W 6 .<;.O)l!t(til~;l; I). <l,"h ~(f ? Rltffi::mJtllO)mt~
iID~':'lt*Hl*! . *Hlil\M~ § lL ~. J1jJltcpif:t).il1t~.:.~~iJ!ll ~ ,0) b~;l; -:, b~ • .fll!t L -c~mltlj""(';l; -:, !JI!~{9IJ
It -I! / "?=-7:/ 1:$~.:.iI1 L. ~7t*f!Ilc ~,;t-:, to)ltcp$~':' t;l;-:,o000(3) I, ginterensis Pergament: !JI!~Ef:Jt.:to)ftcp~-C. ~~~~mt. 1W6'<;'b~~tl""('ti:mtrnt#<
I)~I:I:\ L -n,t.:~'o iiIfU~lti\§N~M~i\§ffi::1JM""(';l;-:,o Iff.I'L,}JjJltifi{9u:mJtll~.:.~~<. ffi:JtIlt).il1tltcpW

1Jt00iJ!ll~iI'~~~mltlj; *Hl*!' *Hl~(HiJif:JtIl~':'J: <ff-~h. CPJtllt).il1t""(t(Jf,f?b~J:'thlf~1!b6h-:,o !JI!~Ef:Jt.: t o)b~:; :il;;tc:ifZtflK c ~t$:~t.:lt""(,t.:~t*P'.J~.ttl!.IK~.:. t tfifi~.:.7t;{fj L -C ~'-:'o~t$:~""('ft -I! / "? =- 7 :/ 1:$~':'~
cp$~':' til1-t -:'0 -t:- L -Ccp$ c 1:$0) c O)r,,'-CM~1:O)~<.

<•

It~~':'~1!b 6ht.: ~ '0 *~O)9Ct.lHt I. takahashii Mataumoto et ~oda Xa-t:- h~.:.j[[~, t 0) ""('Itt.:iI'1;; ? ii' ; j; t.: I. heinzi Somay It*~~.:.j[[~ 6 l..l '0t).1: 3 ~O)~FfEf:JiI1lliil!lirHlItil't.: I) ffi:~ '0 1ff.I~0)::' c (t :; :ilO);;tc:ifZi:FIJtIj.ttl!.~-C t ii?2.1!b 6h -C ~,-:'0 fJ(:.;IU. pennatulus*~~t$:~-c,t -I! / "? =- 7 :/ 1:$~':'. :; :il;;tc:ifZtflK""('ltcptm~.:.l¥iJt-c ~'-:'

b~. JZ''fLt~l;1]""('t.:~'ot~*~ftl"~jHl)iA' :ifZlIlf5!,ii[ ·lIlf133*2.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 149, pp. 396-416, 11 Figs., April 30,1988

854. BARREMIAN AMMONITES FROM THE ISHIDO FORMATION, JAPAN- SUPPLEMENTS AND FAUNAL ANALYSIS*MASAKI MATSUKAWA
Department of Earth Sciences, Faculty of Science,
Ehime University, Matsuyama 790

Abstract. Six species belonging to five genera of Barremian ammonites. are describe~
the Ishido Formation of the Sanchu Terrane in the Kwanto Mountal~s. They are.
~;I~phYllOCeras sp., Karsteniceras asiaticum (Yabe et Shimizu), K. obatal Matsukawa,Heteroceras (Argvethites) sp., Barremites (Barremites) strettostom~, and Va.ldedorsella(Puezalpella) sp. The ammonite fauna of the Ishido Formation (the Ishldo fauna) I~ made up
essentially of two groups of ammonites, the smooth or weakly. ornate no~malY cOIled forms
and various heteromorphs, a characteristic feature of the BarremIan ammon! te faunas. generally
in the Tethyan realm. The Ishido fauna shows a high sim~larity. to .tha: of the. MedJter~anean
region of the Tethyan realrr.., and contains also one specIes, 51mblrskltes (Mllan~wskla) sp.,which migrated from the northeast European province in the European-Boreal regIOn throughthe Arctic epicon tinental sea.
f--------------------Ishido Formation. This completes the descriptionof the 15 species belonging to 10 genera ofammonites obtained from this formation so far.
The characteristics of the Ishido fauna are
discussed and a comparison made with somefaunas of other regions.
I thank Drs. I. Obata (National Science
Museum, Tokyo), P. F. Rawson (University
College, London), J. P. Thieuloy (Universite deGrenoble), J. Wiedmann (Universitat Tiibingen),and H. G. Owen [British Museum (Natural
History) 1 for their critical reading of themanuscript. I am also indebted to Drs. K_ Tanabe(University of Tokyo), and M. Futakami(Kawamura-gakuen Women's University, Chiba)for providing fruitful discussion on certain problems. Thanks are extended to Drs. K. Tanaka(Shinshu University), T. Kotaka (Tohoku University) and I. Fujiyama (National Science
Museum, Tokyo), and Messrs. T. Kinoshita(Tachikawa Dainana Junior High School, Tokyo),
H. Tsuda (Shinagawa-ku, Tokyo) and Y. Ogawa(Manaus, Brazil) for their kind help in collecting
the fossils described herein.

Introduction
The Ishido Formation is one of the best
known Lower Cretaceous sequences seen in
Japan. Since Yabe, Nagao and Shimizu (1926)
described the stratigraphy and molluscan fossilsof the Sanchu Cretaceous, many stratigraphical,
sedimentological and paleontological studies havebeen published (see bibliography in Matsukawa
1983, Obata and Matsukawa 1984). Matsukawa(1983) has established the lithostratigraphy andmolluscan biostratigraphy, and presumed sedimentary environments on the basis of ageological survey over the area of the Cretaceous
formations. He used the cephalopods collectedfrom the Ishido Formation in the inter-regionalcorrelation. These cephalopods were identified
and described by Obata et al. (I976) and Obata
and Matsukawa (1984). However, some ammonites remain to be described.
This paper gives a description of five genera
comprising six species of ammonites from the*Received October 29, 1987; revised
accepted March 15, 1988.

manuscript396

854. Barremian ammonites from Ishido Formation
This work is dedicated to the memory of mylate mother who died suddenly during thewriting of this paper. The study was financially
supported in part by the Science Research Fundof the Japanese Ministry of Education, Science
and Culture (Monbusho) (Obata, no. 56340041
for Cooperative Research in 1981 - 1982;
Matsukawa, no. 56740335 in 1981 and no.
61740463 in 1986).
Geological setting

According to Matsukawa (1983), the Cretaceous strata of the Sanchu Terrane comprise clastic
rocks that are almost 2700 m in thickness (Figs.
1, 2). These are characterized by three cyclicsequences, each one of which is thick withcoarse-grained sediments at the base fining
upward; each attain more than several hundred
meters in thickness and show a changing environ-397

mental facies from nonmarine to marine beds.
Although three sedimentary cycles are recognizable, the whole sequence is divided into four units
named in ascending order the Shiroi, Ishido,
Sebayashi and Sanyama Formations (Fig. 2).
Ammonites occur in the Ishido and Sanyama
Formations. Those in the Ishido Formation
indicate as early to late Barremian age. Thosefrom the Sanyama Formation indicate a late
Aptian to early Cenomanian age. Hence the
Sebayashi Formation, which lies between these
two formations, is probably referable to theUppermost Barremian to Aptian.
The nonmarine Shirai Formation and overlying marine Ishido Formation represent the firstcycle. These two formations successively onlap
the structurally complex basement consisting ofJurassic and older racks, and are interpreted as atransgressive sequence passing from fluvial to

Tok yo10l• 200/ ·.~~3o

1kmo\Sebayash i

o

~:~

1km

,,!e

D· ....

~

R299"', Ishido'- r\~
Figure 1.

':~Ccl~~!~. . . . . O~~4km'~:
p~........
. '.::... B,c/.: •..

Koya
dam

\632

!

'.~- .
621

60~/

•

1.....

... /.

7.:.,~D Cretaceous .~'.
deposits''~o
Map showing the ammonite localities. The prefix SA of the locality number is omitted.

Masaki MATSUKA WA

398
E
0

C

GJ

0

u:

C
<1l

<1l

..0

<1l

~

r--c

E
>.

e

c

<1l
(/)

<1l

..r= ::;;.

0.

<1l:::l~<1l::;;(/)Cl.

>..

..0"
GJ 0
(/)...J

c

<1l

.§

u.:
0

~

'0

<1l

.r=

~

co

(/)

:> u.:

m
:; ~
<1l

I
.~

.r=
(/)

C': Qi,n

(/)

c

:::l

<1l

~E

(/) 0 O~~--e GJ_
'0

-,0

ct

~1
F

9

~2~3D4B5~6D7g§§8•

10

Figure 2. Individual columnar sections with the indication of the horizons which yielded ammonites. 1:
Shale, 2: Alternating beds of sandstone and shale, 3: Muddy sandstone, 4: Sandstone, 5: Conglomerate, 6:
Chert, 7: Pyroclastic rocks, 8: Limestone, 9: Fault, 10: Horizon yielding ammonites. a: Miyakozawa, b: Ishido,
c: Shiroi, d: Myokezawa, e: Mamonozawa, f: Niheizawa, g: Mamyodaira.

epeiric embayment deposits.
Barremian ammonites were found in the
Ishido Formation at 9 localities (Fig. 1) representing three different horizons. These horizons
were confirmed in the Mamonozawa and Myokezawa valleys (Fig. 2). The characteristic species
are as follows: Barremites (B.) difficilis (d'Orbigny), Pulchellia ishidoensis Yabe et Shimizu,
Simbirskites (Milanowskia) sp. and others in thelower horizon; Shasticrioceras sp. aff. S. patricki
Murphy and Phylloceras sp. from the middle
horizon, and Barremites (B.) strettostoma (Uhlig),
Pseudohaploceras japonicum Obata et Matsukawa, Heteroceras (H.) sp. aff. H. (H.) astierid'Orbigny and others from the upper horizon.
These ammonites are all of Barremian age.
Depository

The specimens with the prefix are preserved in

the following institutions:
National Science Museum (NSM-PM), Tokyo
Institute of Geology and Paleontology,
Tohoku University (IGPS), Sendai
Department of Geology, Faculty of General
Education, Shinshu University (SGE),
Matsumoto
Systematic description

Order Ammonoidea Zittel, 1884
Suborder Phylloceratina Arkell, 1950
Su perfamily Phyllocerataceae Zittel, 1884
Family Phylloceratidae Zittel, 1884
Subfamily Calliphylloceratinae Spath, 1927
Genus Calliphylloceras Spath, 1927
Type species: Phylloceras dispu tabile Zittel,
1869, by original designation.

854. Barremian ammonites from Ishido Formation
Calliphylloceras sp.

Calliphylloeras (Wright, 1957, Ll89) on the basis

Figure 3-1

Material: NSM-PM 7531, an imperfect moldof shell.

Description: The specimen is about 45 mm inshell diameter. The shell is discoidal, involute andsmooth. It has a very narrow and shallow
umbilicus (U/D = 0.12) but with a low steep wall.
The whorl is of compressed form being higher
than broad (B/H = 0.27). The flanks are slightly
inflated at the middle of whorl height, and slope
rapidly to the periphery. The venter is narrowand sharp. The test is thin. Numerous subcostae
are very densely arranged, rectiradiate with lesssinuous curvature on the flank, and are gently
arched on the venter. They are weak in theumbilical side, but become gradually conspicuousin the ventral side of flank. Each rib is as broadas the intercostal spaces. The constrictions are
regular and frequent: four in a half volution.
They show weak flexuosity on the internalmold, and are nearly parallel to the ribs. The
suture is unknown.
Measurements (in mm):
Specimen
NSM·PM 7531

Diameter(D)
46.2(1)

Umbilicus Height(H)(U)
5.4{0.12) 24.9

399

Breadth(B)
ca 6.7

B/H
ca 0.27

Remarks: The illustrated specimens ofPhylloceras aldersoni Anderson (Anderson, 1938,
p. 143, pI. 11, figs. 3-6) from the Aptian ofCalifornia are similar to the present specimen in
the umbilical ratio (U/D), the density andsigmoid form of costae and the weakly flexuous
constrictions. The whorl section of the former is,
however, much broader than the latter: theformer having a ratio of breadth to height of
0.81 and the latter, 0.27. Furthermore, thenumerous costae are well marked on the whole
whorl flank, whereas they are present only on
the outer half flank in the latter.
Phylloceras sp. illustrated by Obata andMatsukawa (1984, p. 16, pI. 4, figs. la, b) fromthe Upper Barremian of the Ishido Formationdiffers from the present specimen in having
flexuous constrictions.
The present specimen belongs to the genus

of smooth, involute and compressed shell,
periodic sigmOid constrictions on internal moldand numerous costae.
However, this form from the Ishido Formation can not, by reason of insufficient preservation, be assigned to a definite species at present.
Occurrence: Loc. SA-203.
Suborder Ancyloceratina Wiedmann, 1966
Superfamily Ancylocerataceae Meek, 1876
Family Ancyloceratidae Meek, 1876
Subfamily Ancyloceratinae Meek, 1876

Remarks: The classification of the Ancyloceratinae adopted here has been discussed already in Matsukawa (1987).
Genus Karsteniceras Royo y Gomez, 1945

Remarks: The generic characters are discussed
in Matsukawa (1987).
Type species: Ancyloceras beyrichii Karsten,
1858, by original designation.
Karsteniceras asiaticum (Yabe et Shimizu)
Figures 3-4, 5, 6a, b; 4

1926 Leptoceras asiaticum Yabe et Shimizu, p. 73, pI.
15, fig. 21.

Material: IGPS 22849, the holotype, described by Yabe and Shimizu (1926) in Yabe etal. SGE-008 and 009 specimens collected by K.
Tanaka.
Description: The shell is small, about 18 mm
in maximum diameter, coiled in a loose spire.
The whorl is crioceratid with regular open spire
and with a part of the last volution closely
approaching the next inner one until at least one
and half volutions. The whorl section issubrectangular, nearly flat on the flanks androunded on the venter which bears a siphonalgroove. The whorl is ornamented with numerous,
almost radial, narrow ribs which are separated bybroader interspaces. The intercostal areas aretwice or three times as broad as the rursiradiate

400

Masaki MATSUKA WA

854. Barremian ammonites from Ishido Formation

~I

10

I0::
;

:

:......... /Figure 4. Karsteniceras asiaticum (Yabe et
Shimizu). (Right) Lateral view; (Left upper) Ventral
view; (Left lower) Whorl section. The diagram also
represents the measuring-distances.

ribs on the outer volution (adult stage), but are
crowded in the inner volution of the young stage,
where they are interrupted on the venter. There
are about 20 ribs on the outer volution. The
suture exhibits a characteristic trifid lateral lobe.
Measurements: (in mm);
In Fig. 4, Dx = maximum diameter of loose
spire; Dn = minimum diameter of loose spire; H =whorl height in maximum diameter of loose
spire; B = whorl breadth in maximum diameterof loose spire; Ux = maximum umbilical gap; Un= minimum umbilical gap.
Specimens
IGPS 22894
SGE·008
SGE·009

Ox On H B(B/H) Ux
18.0 12.6 4.4
8.7
16.0 14.0 3.6 1.5(.42) 8.5
16.2 8.5 4.9
7.9

Un
5.8
7.9
4.5

Ux/Ox Un/On
.48
.46
.53
.56
.48
.52

Variation: The percentage ratio of maximum
umbilical gap to maximum diameter of the loose
spire shows a range from 0.48 to 0.53, thepercentage ratio of minimum umbilical gap tominimum diameter of loose spire is from 0.46 to
0.56.
Remarks: The present species is similar to
Karsteniceras beyrichii (Karsten) (Royo yGomez, 1945, p. 460-461, pI. 1, fig. 1), the type401

species of the genus from the Barremian ofColombia, in its coiling characteristics. However,
the whorl section of the former is sub rectangularwith nearly flat flanks throughout all stages.
Furthermore, the interspaces between ribs of theSanchu specimen are broader than in the
Colombian form, the latter being twice or threetimes as broad as the ribs in the former, butequal to the breadth of the ribs in the latter. On
the basis of the subsequent description of K.
beyrichii (Karsten) by Etayo Serna (1968, p.
54-56, pI. 1, figs. 1-3, 5, 7), this species can bealso distinguished by the pattern of its rursiradiate ribs.
Occurrence: Loc. SA-621.

Karsteniceras obatai Matsukawa
Figures 3-7a, b, 8a, b, 9; 51926 Ancyloceras? sp., Yabe and Shimizu in Yabe etal., p. 71, pI. 15, figs. 12, 13.
1987 Karsteniceras obatai Matsukawa, p. 349-354,
fig. 3.

Material: SGE-005, collected by H. Kumai;
SGE-006, -007, -010 collected by K. Tanaka.
Measurements: (in mm); (See Fig. 4 forexplanation of the symbols used).
Specimens Ox On H
SGE·005 14.1 11.2 4.0
SGE·007 19.8 11.4 5.4
SGE·OIO 13.3 9.8 3.6

B

-

-

Ux Un
7.2 5.1
10.2 5.4
6.7 4.6

Ux/Ox Un/On
.51
.45
.52
.47
.50
.47

Comparison: The present species is allied, in
its coiling, to Karsteniceras balernaense Rieber(Rieber, 1977, p. 779-781, pI. 1, fig. 2) fromthe Barremian of Switzerland, but is distinguished by different ornamentation. In K.
obatai, surface of the later shell (e.g. about 10
mm in diameter of NSM-PM9589) is covered
with numerous lirae. In K. balernaense, surface«- Figure 3. 1, Calliphylloceras sp., late Barremian, uppermost part of the formation, from loc. SA-203,
NSM-PM 7531, lateral view, x 1.2. 2, Valdedorsella (Puezalpella) sp., late Barremian, upper part of theformation, from loc. SA-307, NSM-PM 7532, ventral (a) and lateral (b) views, x 1.0.3, Heteroceras (Argvethites)sp., late Barremian, uppermost part of the formation, from loc. SA-203, NSM-PM 7294, lateral (a, d), ventral (b)and dorsal (c) views, x 1.0. 4, 5, 6, Karsteniceras asiaticum (Yabe et Shimizu), early Barremian, lower part of theformation, from loc. SA-621. Holotype, IGPS 22847 (5) and SGE-009 (4), SGE-008 (6): rubber cast (b) frommold (a), lateral views, x 1.7. 7, 8, 9, Karsteniceras obatai Matsukawa, early Barremian, lower part of theformation, from loc. SA-621. SEG-005 (7), SEG-007 (8) and SEG-006 (9): rubber cast (8a) from mold (8b);lateral (7a, 8a, b, 9) and ventral (7b) views, x 2.5 (7), x 1.8 (8) and x 2.0 (9).

MasakiMATSUKAWA

402

Type species: Heteroceras (Argvethites)
lashense Rouchadze , by original designation .
Heteroceras (Argvethites) sp.
Figures 3-3a, b, c, d; 6

1976 A nahamulino aff. subcy lindrica
Obata et al., figs. 6-2 , pI. 2, fig. 2.

:,
0
,

'

:
,

'
'

:

>.

}

.!

Figure 5. Karsteniceras obatai Matsukawa. (Left)
Lateral view; (Right upper) Ventral view; (Right lower)
Whorl sec tion.

of the later shell (e.g. about 20 mm in diameterof the type specimen) is covered with ribbing.
Occurrence: Loc. SA·62 J.
Genus Heteroceras d' Orbigny , 1850

Type species : Heteroceras emericianum, subsequent designation by Meek , 1876.
Subgenus Argvethites Rouchadge , 1933

(2)

(3)

o

2cm

Figure 6. Heteroceras (Argvethites) sp. A: Helix,
B: First shaft, C: Second shaft. (1) Internal mold of
sec ond shaft, (2) Helix , fir st shaft and second shaft of
NSM-PM 7294 , (3) Drawing of the same specimen toshow the posi tion of the helix and fir st sha ft.

(d 'Orbigny) ,Material : NSM·PM7294, an imperfect internalmold of shell , collected by H. Tsuda.
Description : This specimen consists of two
main parts, an initial helix and shaft. The last
helical whorl passes directly into the first shaft
which is flattened laterally. The first shaft islong, being about four-fifth as long as the second~haft and almost parallel to the second one.
The helix is small in size. The general outlineof the helix is roughly triangular, with an apicalangle of about 60 degrees. The helical whorls arecontiguous and ornamented by twisted single
ribs.
The second shaft is located on a level with the
top of breadth along the length. The second shaft
is nearly circular in cross section, having a ratio
of l.l in B/H at the larger end and a ratio of 1.2
at the smaller end. The annular ribs are elevated ,slender, slightly sharpened at the top , but are lowon the dorsum. In the second shaft at the laterstage , the ribs are slightly flexuous or rectiradiateon the flanks , and separated by interspaces which
are three times as broad as the ribs. At the bent
part of the second shaft , the whorl has a shallow
groove , and the ribs weaken on the dorsum andare divided into primaries and secondaries
branching at the ventral-shoulder. The suture line
is of Ancyloceras type but is only partially visible
and not in detail.
Measurements (in mm) of the specimen are as
follows : height of total helix = 3.2, length ofsecond shaft = 50.2, whorl-height at larger end of
second shaft = 11.3 , whorl-breadth at larger endof second shaft = 12 . 1, whorl·height at smaller
end of second shaft = 7.8, whorl-breadth atsmaller end of second shaft = 9.3.
Remarks: The present specimen is similar, in
its whorl section, to the illustrated specimen of
Crioceras furcatum (Simionescu, 1898, p. 146 ,854. Barremian ammonites from Ishido Fonnation

pI. 4, figs. 7 a, b) from the Barremian of Romania.
The latter was listed subsequently by Patrulius(1969) as Toxoceratoides subfurcatum, sp. nov.
This is a view also held by J. P. Thieuloy(personal communication, Descember 1986).
However, the venter of bent part of the former isslightly grooved without interruption of ribs.
Anyhow, the present specimen belongs to thesubgenus Argvethites (Wright, 1957, L212) on
the basis of aperture level with top of helix.
However, this form from the Ishido Formation
cannot, by reason of insufficient preservation, be
assigned to a definite species at present.
Occurrence: Loc. SA-203.
Suborder Ammonitina Hyatt, 1900
Superfamily Desmocerataceae Zittel, 1895
Family Eodesmoceratidae Wright, 1955
Subfamily Eodesmoceratinae Wright, 1955
Genus Barremites Kilian, 1913
Type species: Ammonites difficilis d 'Orbigny
1841, by original designation.

Subgenus Barremites Wright, 1953
Type species: Ammonites difficilis d'Orbigny
1841, by original designation.

403

Wernsdorfer Schichten in Silesia (Poland), andthe several hypotypes of Desmoceras strettostoma figured by Karakasch (1907, p. 72-73, pI.
5, figs. 3-5; pI. 6, fig. 5), from the sequence in
the Sably Mine in Crimea. The species from the
Sanchu Cretaceous has ratios of 0.13 to O. 18
in U/D and 0.36 to 0.60 in B/H; the Silesian
species has ratios of 0.10 to 0.11 in U/D and
0.41 in B/H by estimate; the Crimean species
has ratios of 0.10 to 0.16 in U/D and 0.46 to
0.60 in B/H. Although the Sanchu specimens are
smaller than the Silesian and Crimean ones, they
may be included within the range of variation ofa single species. Thirty four specimens from loco
SA-203 are less than 25 mm in diameter. Besides,
the subangular shoulder of ventro-lateral
periphery, which is the other difference between
the Japanese specimens and the Crimean ones, isrecognizable in the all of Japanese specimens but
only in the middle portion of one of the Crimean
ones (Karakasch, 1907, pI. 5, fig. 3).
To sum up, the Sanchu specimens represent
Barremites (Barremites) strettostoma.
Subfamily Puzosiinae Spath, 1922
Genus Valdedorsella Breistroffer, 1947
Type
Anthula,1947).

species:Desmoceras
1899 (designated byakushaense
Breistroffer,

Barremites (Barremites) strettostoma (Uhlig)
Figures 8-4, Sa, b, c

1976 Barremites (Barremites) sp. aff. B. strettostoma
Obata et ai., p. 128-129, pI. 1, fig. 1, pI. 2, fig.
4.
1984 Barremites (Barremites) sp. aff. B. strettostoma
Obata and Matsukawa, p. 21, pI. 1, figs. la, b, 3.

Subgenus Puezalpella Dimitrova, 1967
Type species: Desmoceras uhligi Haug, 1889,by original designation.
Valdedorsella (Puezalpella) sp.
Figures 3-2a, bMaterial: See Obata et al. (1976).
Remarks: Barremites (Barremites) sp. aff. B.
strettostoma from the Sanchu Cretaceous has
been described by Obata et al. (1976) and Obata
and Matsukawa (1984). This form was distinguished from B. (B.) strettostoma by
differences in certain minor characters shown by
the two syntypes of Haploceras strettostoma by
Uhlig (1883, p. 225, pI. 17, figs. 3,4), from theMaterial: NSM-PM 7532, an imperfect moldof a shell, in which the inner whorl is notpreserved.
Description: The specimen is about 46 rom indiameter of shell. The shell is compressed,
evolute, smooth, having a somewhat angularwhorl section. The whorl is high and muchcompressed (B/H = 0.52). The breadth is at a

Masaki MATSUKA WA

404

the Sanchu specimen is too incomplete to make aspecific determination.
Occurrence: Le. SA-307.

maximum about two-thirds of the flank height
above the umbilical margin. The flanks areslightly inflated. The umbilicus is of narrow
width (19%) in proportion to the diameter and issurrounded by a sloping wall. The shell surface isornamented with four distinct straight constrictions which are concave on the whorl flank,
becoming strongly prorsiradiate near the venter
over which they cross. The suture line is
unknown.
Measurements (in mm);
Diameter(D)

Specimen

NSM-PM 7532

Umbilicus Height(U)(H)

8.8(0.19)

46+

Faunal analysis

Breadth B/H

17.7
15.9

(B)

8.3

0.52

Remarks: The present specimen is dis·tinguished from the illustrated specimen of
Puzosia melchioris (Karakash, 1907, p. 75-76,
pI. 7, fig. 4a, b) from the Barremian of Crimea inthe umbilical ratio (U/D): the former is narrow(0.19), but the latter is wide (0.30).
Valdedorsella (Puezalpella) uhligi (Haug,
1889) illustrated by Dimitrova (1967, pI. 69,
figs. 4, 4a; pI. 72, fig. 1, la, 2, 2a) from theAptian of Bulgaria differs from the present
specimen in having a wide umbilicus. However,Table 1.
~

Specific name[1] Characteristics of assemblages.
The ammonite assemblages of the Ishido
Formation are made up of three morphotypes:
smooth or weakly ornate ammonites (66individuals being classified with at least 7 species
belonging to 5 genera of the Phylloceratidae,
Desmoceratidae and Eodesmoceratidae), heteromorph ammonites (40 individuals, with at least
6 species of belonging to 5 genera of the Ancyloceratidae and Heteroceratidae) and ornate
ammonites (6 individuals representing 2 speciesof 2 genera belonging respectively to theOlcostephanidae and Pulchellidae) (Table 1). The
assemblages show a predomination of smooth or
weakly ornate ammonites with the heteromorphammonites providing a higher range of morphological diversity (Fig. 7).
The composition of species, number ofindividuals and preservation of ammonite
assemblages in the Ishido Formation differ fromList of ammonite species from the Ishido Formation.

Local i ty number

632

621

604

416

100

101

Phyllocerus sp.

~OO

203

307

802

Total
I

1

Ca.ZLiphylloCBl'aS sp.

1

Phy lloceratidae gen. et sp. indet.

1

1

Boehi ani t idae gen. et sp. indet.

1

1

Shasticrioaer>as sp. aif. S. patrioki MURPHY

1

Kar'sceniael'a8 QBiaticu.m (YABE et SIIIMlZU)
K.

3

5 (I)
1

5

1

1

3

4

.ff. H. a8tieri d ORB IGNY

H.

(R. ) sp.

H.

(Al'gvethi tes) sp.

1

3(1 )

obatai MATSUKAWA

Ancyloceratidae gen. et sp. indeL
HeteY'Oaera.s (8. ) sp.

4

I

1

Ptychoceratidae gen. et sp. indet.
I(B. ) sp.

I

5

5
1

I

2
2

2

4(1)

4
1

PseudohapZocel'ao japoniawn OBATA et MATSUKAWA
PulcheZlia ishicioel1sis YABE et SHIMIZU
Total

9

I

Va!dedorse!!a (l'I<eza!pe!!a) sp_
Ba1'l'emites (B.) difficilis (d' ORBIGNY)(B. ) strettostoma (UHLIG)

5

1

1

Simbil'skites (Milanowskia) sp.

B.

10

8

Heteroceratidae gen et sp. indet.

B.

1

34

1

16

1

2

2

2

4 (I)
1

18

38
17

4

I

Numbers in parentheses are taken from Yabe, Nagao and Shimizu (1926).

1

I

4

2

77

5

2

112

854. Barremian ammonites from Ishido Formation

Genus

FamilyPhylloceratidae

405

SuborderPhylloceratina

Figure 7. Circular diagrams displaying the occurrence of ammonite species in the Ishido Formation at
genus, family, suborder levels and proportions of morphotypes of smooth or weakly ornate ammonites 0),
ornate ammonites (2) and heteromorph ammonites (3). They are shown in the frequency of individual
occurrences of ammonite species belonging to genus, family and suborder.

place to place and horizon to horizon. In
particular, at Loc. SA-621 (lower part of theformation in the western area) and Loc. SA-203(eastern uppermost part in the eastern area),many ammonite species occur along with a large
number of bivalve species including trigonians(Table 1; see table 2 in Matsukawa, 1983). The
body chambers, phragmocones and fine ornament of these ammonite specimens are generally
well preserved, although the body chambers andphragmocones of large-sized specimens from Loc.
SA-203 are partly broken. Judging from thefeatures described above, these assemblages may
be showing evidence of biocoenose or thanatocoenose conditions. The degree of crushing of
the inner whorl of specimens from Loc. SA-621
is different according to which genera the specimens belong (Fig. 8). The specimens of Barremites (smooth or weakly ornate ammonites) andKars ten iceras (heteromorph ammonites) are
crushed flat, but the specimens of Pulchellia(ornate ammonites) nearly retain their originalform. This is interpreted as an indication of differences in the strength of the shells between
various genera or coiling types against the crushing forces of the compacting sediments in which
they have been buried. On the other hand, as

the broken specimens of ammonites of large sizeand loosely spiral shells from Loc. 203, possessfine ornament well preserved, it is suggested thatthey have been transported a short distance andto have been broken up by wave action upon ashore. These occurrences contrast strikingly withthose of bivalves collected from the same
localities (described below).
So far as the associated bivalves which show
epifauna or infauna in mode of living areconcerned, they are probably still close to their
original living places. This is suggested by their
good state of preservation with conjoined shells
in poorly sorted muddy sandstone, the numberof large shells, that is, Gervillaria haradae andPterotrigonia (P.) hokkaidoana, at Loc. SA-203,
and those with unabraded finely ornamented
valves, that is, Grammatodon (Nanonavis)
yokoyamai and Astarte (A.) subsenecta, at Loc.
SA-621. The bivalve fauna is considered toconsist of shallow marine inhabitants.
Consequently, ammonites and bivalves can beinterpreted as inhabiting a shallow marine
embayment, and to have migrated eastward in
response to the transgression of the Ishido Sea.

Masaki MA TSUKAWA

406

I

I

I
Sa

8b

I
9b

3a

3b

854. Barremian ammonites from Ishido Formation[2] Comparison with some facies and environments containing ammonite assemblages inother regions.
According to Kakabadze (1979), in thesouthern USSR (Crimea, Caucasus and Middle
Asia) the ammonite assemblages, lithofacies andenvironments of the Hauterivian to Barremianare summarized as follows: (1) the abundantassemblages are made up of heteromorphammonites, smooth or weakly ornate ammonites
and ornate ammonites in ascending order, (2) the
assemblages are included in the lime-richlithofacies which are divided into five types
interpreted to be deposited in relatively shallow
sublittoral and infralittoral environments, (3) the
heteromorph ammonites show the highest
morphological diversity, and (4) the smooth or
weakly ornate ammonites tend to increase in
numbers proportionally to the occ'.lrrence offiner deposits which indicate a deeper water
environment.
Thus, features of ammonite occurrences in thesouthern USSR are similar to those of the Ishido
Formation in Japan. There are, however, somedifferences in lithofacies and presumed environments: the sequence in the USSR is considered
to be sublittoral and infralittoral, but theJapanese sequence indicates an epeiric shallow
marine environment. The similar relation
between ammonite assemblages and lime-richlithofacies in southern USSR is also recognized in
the Barremian stratotype at Angles in France(Busnardo, 1965).
The features as described above suggest thatthe Barremian ammonite fauna generally in thesegeographic regions can be interpreted as being
made up of three morphological groups ofammonites, the heteromorphs and smooth or
weakly ornate ammonites predominating, theornate ammonites being of secondary importanceand the fauna as a whole indicates a shallow
marine habitat.
It is worthy of note that the habitats of407

ammonite faunas in the Upper Cretaceous in
Japan have been divided also into different
environments based on morphotypes (Matsumoto, 1965; Matsumoto and Okada, 1973;
Tanaka, 1963; Tanabe, 1979; Tanabe, Obata andFutakami, 1978). Smooth or weakly ornate ammonites (e.g. desmoceratacean and lytoceratacean) are considered to indicate an offshore
inhabitat, whereas ornate morphotype ammonites (e.g. the hoplitaceans) are considered
to indicate coastal sea habitat.
From the facts as described above, there is apossibility that the habitat of ammonite faunas
splits off during the period ranging from the
Early to Late Cretaceous.
[3] Comparison with some faunas of otherregions.
The ammonite bearing strata of the Ishido
Formation is considered here to be a key to theinter-regional correlation among the Mediterranean, Circum Pacific and European-Boreal
regions based on the occurrence of some characteristic genera: the Mediterranean, Tethyan region is represented by Barremites and Pulchellia,
the Circum Pacific region by Shasticrioceras andthe European-Boreal region by Simbirskites(Matsukawa, 1983; Obata and Matsukawa, 1988).
Here, the following facts are noted: (1) The
genus Simbirskites, one of the characteristic
genera in the Upper Hauterivian of the EuropeanBoreal region extending into the early Barremian,
and the genus Barremites and Pulchellia which
are indices of the Barremian in the Mediterranean
region, are contained in one and the same bed ofthe lower part of the Ishido Formation, (2) The
Ishido Formation in Japan and the Ono Formation in California contain some Tethyan ammonites though these two areas belong to the
Circum Pacific region.
As to the first problem, Simbirskites (Milanowskia) sp. was collected from the Loc. SA-62I(in the western area) and Loc. SA-lOi (in the~ Figure 8.
Comparison of the mode of occurrence of ammonites from two localities, Loc. SA-621 (1-3b,
6-8b, 10) and Loc. SA-203 (4-Sc, 9a, b, 11). 1, 2: Barremites (Barremites) difficilis; 3a, b, 6: Pulchellia
ishidoensis; 4, Sa, b, c: Barremites (B.) strettostoma; 7: Simbirskites (Milanowskia) sp.; 8a, b, 10: Karsteniceras
obatai; 9a, b: Pseudohaploceras japonicum; 11: Heteroceras (H.) sp. aff. H. astieri. Scale bars show 1 cm.

Masaki MATSUKA WA

408

1
3a
3b

4c
Figure 9. Comparison with Simbirskites (Milanowskia) sp. (INSM-PM 9515 ; 2a, b: rcps 22847) from the
Ishido Formation and Simbirskites (Milanowskia) speetonensis (Young et Bird) fro m the Speeton Clay (3a, b)
and Ulyanovsk (formerly Simbirsk) , Russia (4a, b, c). Taken from Obata and Matsukawa (1984) for 1, 2a, band
Raw so n (1971a) for 3a - 4c. The figure s are x 2.00, 2a, b) and natural size (3a, b, 4a, b, c).

eastern area). From each locality only one specimen was collected by Yabe and Shimizu , and by
the author, re spectively (Table 1). This species issomewhat similar to Simbirskites (Milanowskia)
speetonensis (Young et Bird) from the part of
the Speeton Clay (assigned to the lowest quarterof the Upper Hauterivian) in England (Obata
and Matsukawa , 1984) (Fig. 9).
Loc. SA-621 is a cliff section , 30 m high,exposing muddy sandstone beds. In the lower
part , at about 10 m above the base of the cliff,
there is a horizon which yields mollusks andammonites. If the Simbirskites (Milanowskia) sp.
which was originally described by Yabe et al.
(1926) as Simbirskites kochibei, was collectedfrom this fossiliferous bed, the boundary
between the Hauterivian and Barremian can beplaced below the bed.
However, a thick pile of marine clastic

sediments accumulated rapidly in the Sanchu
area (for instance, the Ishido Formation
corresponding to one part of the Barremianattains 500 m in thickness) and it might bedifficult to make an exact comparison between
the horizon at 10 m in SA-621 and about 27 m
of C6 to LB5 bed (Upper Hauterivian to Lower
Barremian) in the Speeton Clay.
Until now only two specimens of the genus
Simbirskites have been found at two localities in
the lower part of the Ishido Formation. The
genus is one of those characteristic of the Upper
Hauterivian of the European-Boreal region
extending into the earliest Barremian (Rawson
1971a, 1983 ; Rawson and Mutterlose, 1983 ;Kemper 1973): with a distribution , geographically, mainly in northern Europe, the Urals,
Arctic Canada, northern Siberia and the northern
Pacific Coast regions of the Arctic-Boreal

854. Barremian ammonites from Ishido Formation......-

~

Japan

I

Yo r k. Was h .

Bar.

Barrem.

Sim.

1------+ -

I

Sim.

Spit.
Sim.

- -- - +-._--+-.:..-r..----i

Spit .

•

Hauter.

o*• Simbirskites
Simbirskites
Barremites•.&(Haut.)(Barrem.)

Sim.
Spit.
Bar.

I

Simbirskites
Spitidiscl,ls
Barremites

Shasticrioceras?(Hau te r i v.)
Shasticrioceras (Barrem.)York.: Yorkshire,

Wash.: Washington

Inferred distribution of Simbirskites\::::::llnferred currents around the Arctic

D

land

Figure 10. Occurrence and comparison of stratigraphical range of the genus Simbirskites. Continental
position after Smith et al. (1981). Coastlines compiled from Jeletzky (1970), Kauffman (1973) and Ziegler(1982). Biostratigraphy is based on Rawson (1971b, 1983), Imley (1960), Kemper et al. (1981) and Matsukawa(1983). Oceanic circulation adapted from Obata and Matsukawa (1987).

409

410

Masaki MA TSUKA WA

Table 2.

GeneraDistribution of Barremian fauna. 1: Yorkshire, 2: Hannover, 3: Wernsdorfer, 4: Angles, 5: Bulgaria, 6:
north Caucasus and Crimea, 7: Madagascar, 8: the Arctic Canada, 9: Pacific slope of Canada, 10: Ono,
11: Boyaca, 12: Ishido. For references see text.
' " Faunas

Phy lloaems
Partsahiaeras
Phy i lopaahyaems
Calliphy lloaems
EuphyUoaeras
SalfedieUa
Sowerbyaems
Diasa loael'as
Lytoaems
Eulytoaems
Al"gonauticeras?

Eotetmgonites
Protetragonites
Pseudotetragonites
Costidisaus
Maaroscaphi tes
Aaa>1tholytoaems
Cpi.oceratites
Toxoceras

Hop loarioaems
Shastiarioaems
Pedioaeras
Par<zay loaeras
Karstenic81'aS
Simancy loce ras
Protacrioceras

Acrioceras

Dissimi li tes
Uhligia
Paraspinoceras
A'lcyloceras
Hami ticeras

Toxoaeratoides
Heteroaeras
Hemibaaulites
Colahidites
Imel'ites
Balearites
Hemihoplites
Mathel'onites
Anahamulina
Hamulina
Hamu lini tes
Euptyahocel'as
Ptyahoaeras
Leolissoaeras
Aconeceras

1

2

3

4

5

6

7

8

9

10

11

12

•• •
• • •
•
••• • ••••• • •• ••• •• •• • • • • •••••• • •••• •• • ••• -.-• •• • • •• •••••• •• •• ••• • • • •• ••• •• •• • • • ••• • • • • • -.-• • ••• •• •• • • •
• •
•
••
• • •
• •
• • •
• ••• •• ••• ••• • •• ••• •(to be continued)

854. Barremian ammonites from Ishido Formation
Olcostephanus
Simbirskites
Ba:rl"emites

Subsynetla
ValdedorseUa
Pe eudohap Zoceras

CaZZizoniaeras
Melchiorites

Spitidiscus
Ftesiospi tidiscus
Holcodiscus
Parasaynoceras
Astieridiscus

Sitesites

Nicklesia
PuZcheUia
Cal'S tenia

Subpulchetlia
PsiZotissotia

Paraspiticeras

ParahopZites

•

• • •
• • •• ••• ••• • • •• • •• •• • • •• ••• • •••• • •••• • •province (Fig. 10). It extends into the early
Barremian, however, and the coincidence of themany specimens of the genera Barremites,
Pulchellia, Karsteniceras and other species in thelower part of the Ishido Formation, having
something in common with the Barremianammonite faunas in the Mediterranean region,
indicates not only an early Barremian age for theJapanese examples of Simbirskites (Milanowskia),
but also an intermingling of Boreal and Tethyan
forms in this region.
The Simbirskites species in the Ishido
Formation probably migrated from the northwest European province via the Arctic regions to
Sanchu. The occurrence of Simbirskites with
Mediterranean ammonites in the lowest
Barremian has been recognized also in northwest
Europe (Kemper et al., 1981) and in the
Caucasus and Crimea (Kakabadze, 1983).
As to the second problem, I list Barremianammonite genera from the world (Table 2):
Yorkshire and Hannover in the northwest
European province and the Arctic Canadianprovince as the European-Boreal region,
Wernsdorfer in Silesia (Poland), Angles in France,
Bulgaria and north Caucasus and Crimea as the411• • ••• ••••• •• ••

Mediterranean region, Madagascar as the Indian
region, and Pacific slope of Canada and Ono,
California in the northeast Pacific province,
Boyaca, Colombia in the southeast Pacificprovince, and Ishido, Japan in the northwest
Pacific province as the Circum Pacific region.
The data on faunas outside Japan are based on
the following references: for the Yoskshire
fauna Spath (1924), Rawson (1971a, b) andRawson and Mu tterlose (1983), the Hannover
fauna Koenen (1902) and Kemper (1976), theWernsdorfer fauna Uhlig (1883) and Vasicek(1972), the Angles fauna Busnardo (1965), theBulgarian fauna Dimitrova (1967), the north
Caucasus and Crimean fauna Drushtchits andKudryavtseva (1960), the Madagascar fauna
Collignon (1962), the Arctic Canadian fauna
Jeletzky (1971), Pacific slope of Canadian fauna
Jeletzky (1971), the Ono fauna Murphy (1975)
and the Boyaca fauna BurgI (1954). By thesereferences, I confirmed individual species shownby plates and descriptions on the Hannover,Wernsdorfer, Bulgarian, northern Caucasus andCrimean, Madagascar and Ono faunas, whereas I
followed the specific names by lists on theYorkshire, Angles, Arctic and Pacific slope ofMasakiMATSUKAWA

412.4 Shasticrioceras@

Karstenicerasl::. Shasticrioceras?(• Barremites

Hauterivian)* Pu/chelJia• Heteroceras

Figure 11. Map showing the distribution of the Barremian ammonite genera: Shasticrioceras, Heter~ceras,
Karsteniceras, Barremites and Pulchellia, found also in the Ishido Formation. Presumed land dotted. Contmental
position after Smith et al. (1981). Coastlines from Harrington (1962), Gordon (1973), Jeletzky (1970),
Kauffman (1973) and Ziegler (1982).

Canada and Boyaca faunas.
From the geographical distribution of theammonite faunas in the early Barremian, it isapparent from Table 2 that the majority of theammonites in the Circum-Pacific localities are of
Tethyan affinity. The Arctic Canadian fauna is
more characteristic of the Boreal-realm, which
includes also northwestern Europe. However, thePacific slope of Canada and Japan have a fauna inwhich there are elements of Boreal affinity in a
predominantly Tethyan assemblage (Fig. 11).
Conclusions
1. Five genera including six species ofammonites from the Ishido Formation, whichwere not included in the descriptions by Obata etal. (1976) and Obata and Matsukawa (1984), aredescribed in this paper. The total number of theammonites now known from this formation
amounts to 15 species belonging to 10 genera.
2. The ammonite fauna of the Ishido
Formation is characterized by two Tethyan
genera, Ba"emites and Heteroceras, and also by
the association of smooth or weakly ornate
ammonites and heteromorph ammonites whichcharacterises faunas of this age in the Tethyan
province as a whole including those in thesouthern USSR and the type sequence of Angles,
France.
3. The Simbirskites species from the lower
Ishido Formation is a migrant from thenorthwest European province via the Arctic
region.
4. The ammonite fauna of the Ishido
Formation shows a high similarity to the854. Barremian ammonites from Ishido Formation
Californian and Colombian faunas in the Circum
Pacific region and to the Silesian and Bulgarianfaunas in the Mediterranean region. These faunas
from the Circum Pacific region are essentially
Tethyan in character. However, the Pacific slopeof Canada and Japan have a fauna in which there
are elements of Boreal affinity in a predominantly Tethyan assemblage.
AppendixCollection localities
Loc. SA-IOI
Location: About 1000 m north of Myoke, Nakazato
village, Tano county, Gunma Prefecture.
Horizon and age: Basal part of the Ishido Formation.
Early Barremian.
Occurrence: Dark bluish gray sandstone in which a
fossiliferous lens of gray calcareous sandstone occurs.
Ammonites occur in the lens together with bivalves
which were reported by Matsukawa (1983; table 2). A
specimen of Simbirskites (Milanowskia) sp. wascollected from this horizon. Although fragmentary, its
fine ornament is well preserved. Fossil bivalves are
represented by disarticulated broken valves.
Collector: M. Matsukawa.
Loc. SA-200
Location: North of Ichinosebashi, to the north of
Sebayashi hamlet, Nakazato village, Tano county,
Gunma Prefecture.
Horizon and age: Upper part of the Ishido
Formation. Early Barremian.
Occurrence: Bluish gray, poorly stratified muddy
sandstone. A fragmentary specimen of Shasticrioceras
sp. aff. S. patricki Murphy was collected from this bed
and is not accompanied by other mollusks.
Collector: M. Matsukawa.
Loc. SA-203 [= loco 9, Oze, in Yabe et al. (1926); =loco N9 in Takei (1963); = Hy4008 in Hayami (1966)]
Location: Ichinosebashi, north of Sebayashi,
Nakazato village, Tano county, Gunma Prefecture.
Horizon and age: Uppermost part of the Ishido
Formation. Late Barremian.
Occurrence: Dark bluish muddy sandstone consistingof thick-bedded to massive strata, and alternating bedsof dark bluish muddy sandstone and shale. Manymollusks were reported from this bed (see table 2 in
Matsukawa, 1983). Fossils are represented by casts andmolds. In the small specimens (e.g. Barremites (B.)
strettostoma, a smooth to weakly ornate ammonite),the body chamber and phragmocone are generally wellpreserved.
In some large-size specimens (e.g.

413

Pseudohaploceras japonicum, a weakly ornate ammonite
and Heteroceras (H.) sp. aff. H. (H.) astieri, aheteromorph ammonite), phragmocones and bodychambers are separated in most cases. Many bivalves are
generally in good preservation with conjoined valves.
Ammonite species: Calliphylloceras sp., Heteroceras(H.) sp. aff. H. (H.) astieri, Heteroceras (H.) sp.,Heteroceras(Argvethites)sp.,
Barremites(B.)
strettostoma, Barremites (B.) sp., Pseudohaploceras
japonicum, Phylloceratidae gen. et sp. indet.,
Ancyloceratidae gen. et sp. indet., Heteroceratidae gen.
et sp. indet. and Ptychoceratidae gen. et sp. indet.
Collectors: H. Tsuda, Y. Ogawa, M. Futakami, and
M. Matsukawa.
Loc. SA-307 [~ loco 10, Kawarazawa, in Yabe et al.
(1926)]
Location: Niheizawa, being located at about 1 km
north of Sakamoto hamlet, Oganomachi town, Chichibu
county, Saitama Prefecture.
Horizon and age: Upper part of the Ishido
Formation. Late Barremian.
Occurrence: Dark blue, calcareous, thick-bedded andfine-grained sandstone with intercalated thin-bedded
muddy sandstone. Many mollusks were reported by
Matsukawa (1983, table 2). The mode of occurrence
and preservation of bivalves differ generally dependingon the type of lithology. Casts and molds, and disjoint
shells that are not completely broken, are contained in
the thick-bedded, fine-grained sandstone; fine wellpreserved ornament, and articulated valves are found in
the thinly-bedded muddy sandstone. Fine spines ofmany Ceratosiphon sp. (gastropods) are also wellpreserved in the thinly-bedded muddy sandstone. As far
as the ammonites are concerned, the ribs on the main
helical whorl of Heteroceras (H.) sp. aff. H. (H.) astierifrom the thick-bedded fine-grained sandstone are wellpreserved. However, the fine ornament of Barremites(B.) sp. collected from the thinly-bedded muddy
sandstone is fairly crushed toward the lateral margin.
Ammonite species: Heteroceras (H.) sp. aff. H. (H.)
astieri, Barremites (B.)
sp. and Valdedorsella(Puezalpella) sp.
Collector: M. Matsukawa.
Loc. SA-416
Location: Tonedaira, Ueno village, Tano county,
Gunma Prefecture.
Horizon and age: Upper part of the Ishido
Formation. Late Barremian.
Occurrence: Thick-bedded black muddy sandstone.
A single specimen of Barremites (B.) strettostoma wascollected. It is partly erased and is not accompanied byother mollusks.
Collector: M. Matsukawa.

414

MasakiMA TSUKA WA

Loc. SA·604
Location: Sinzaburomaezawa located about 2 km
west of Jukkoku pass, Saku town, Minamisaku county,
Nagano Prefecture.
Horizon and age: Upper part of the Ishido
Formation. Latt: Barremian.
Occurrence: Black muddy sandstone. A specimen of
Heteroceras (H.) sp. was collected and is a fragment of
curved shaft with well preserved fine ornament.
Pholadomya sp. is also present in the same bed
preserved as disarticulated valves.
Collector: M. Matsukawa.
Loc. SA-621 [= loc. 4, Ishido, in Yabe et al. (1926);= loc. Hy 4015 in Hayami (1966); = loc. 11 in Takei etal. (1977) I
Location: Ishido, about 4 km east of Koya, Saku
town, Minamisaku county, Nagano Prefecture.
Horizon and age: Lower part of the Ishido
Formation. Early Barremian.
Occurrence: Thick-bedded black muddy sandstone.
Mollusks are occasionally found in the muddy sandstone
to the exclusion of other fossils. Many mollusks werereported from the bed: Yabe et al. (1926), Hayami(1965, 66), Takei et al. (1977) and Matsukawa (1983).
Ammonite specimens collected are represented bycasts and molds. Their body chambers and phragmocones are generally preserved, although the inner whorlof the younger stages of many specimens are flattened
laterally. Bivalves are characterized by small specimens.
These specimens are mostly preserved as disjointed
valves, although the fine ornament is well preserved.
Fine spines of gastropods are preserved intact as casts.
Ammonite species: Karsteniceras asiaticum K
obatai, Simbirskites (Milanowskia) sp., Barremite; (B.)
difficilis, Pulchellia ishidoensis and Ancyloceratidae gen.
et sp. indet.
Collectors: K. Tanaka, H. Kumai, S. Usuda, 1.
Fujiyama and M. Matsukawa.
Loc. SA-632 ['" loc. 18 in Takei et al. (1977) I
Location: The upper stream of Miyakozawa, about
2.5 km south of Koya, Saku town Minamisaku county
Nagano Prefecture.
',
Horizon and age: Uppermost part of the Ishido
Formation. Late Barremian.
Occurrence: Black muddy, bedded-sandstone. A
single poorly preserved ammonite specimen, here
identified as Ancyloceratidae gen. et sp. indet wascollected from this bed. The specimen is flattened
laterally, without the inner whorls of younger stages.
Some bivalves, Nanonavis (N.) yokoyamai and Astarte(A.) subsenecta, occur in the same layer as disarticulated
valves.
Collector: M. Matsukawa.

Loc. SA-S02
Location: Mamyodaira, Oganomachi town, Chichibu
county, Saitama Prefecture.
Horizon and age: Upper part of the Ishido
Formation. Late Barremian.
Occurrence: Greenish, medium-grained sandstone, inwhich a lens of fossiliferous black calcareous mudstone
occurs with well preserved ammonite specimens.
Bivalves wi th fine ornament are also well preserved,
although they occur mostly as disjointed valves.
Ammonite species: Barremites (B.) strettostoma.
Collector: T. Kinoshita.

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Smith, A. G., Hurley, A. M. and Briden, J. C. (1981):
Phaner~zoic
paleocontinental
world
maps.
Cambridge, Cambridge Univ. Press. p. 1-102.
Spath, L. F. (1924): On the ammonites of the Speeton
Clay and the subdivisions of the Neocomian. Ceol.
Mag., 61, p. 73-89.
Takei, K. (1963): Stratigraphy and geological structureof the Cretaceous system in the eastern part of theSanchu Graben, Kwanto Mountains. Jour. Ceol.
Soc. Japan, 69, p. 130-146 (in Japanese withEnglish abstract).

416

Masaki MATSUKA WA- - , Takizawa, F., Takeuchi, T. and Fujiwara, H.
(1977): Cretaceous system in the western part of
the Sanchu graben, Kanto mountains. Ibid., 83, p.
95-113 (in Japanese with English abstract).
Tanabe, K. (1979): Palaeoecological analysis ofammonoid assemblages in the Turonian Scaphites
facies Hokkaido, Japan. Palaeontology, 22, 3, p.
609-630.
- - , Obata, 1. and Futakami, M. (1978): Analysis ofammonoids assemblages in the Upper Turonian of
the Manji area, central Hokkaido. BUll. Nat. Sci.
Mus., C, 4, p. 37 -62,1 pI.
Tanaka, K. (1963): A study on the Cretaceous
sedimentation in Hokkaido, Japan. Rep. Ceol.
Surv. Japan, 197, p. 1-119,3 pis.
Uhlig, V. (1883): Die Cephalopodenfauna der Wernsdorfer schichten. Denkschr. K. Akad. Wiss. Wien.,

Math. Nat. Kl., 46, p. 127-290, pis. 1-32.
Vasicek, Z. (1972): Ammonoidea of the Tesin-Hradiste
Formation(LowerCretaceous)intheMoravskoslezske Beskydy Mts. Rozipravy, 38, p.
5-103, pis. 1-16.
Wright, C. W. (1957): In Moore, R. C. (ed.), Treatise on
Invertebrate Paleontology, part L, Mollusca,
Cephalopoda, Ammonoidea, Ll-L490, Ceol. Soc.
A mer. & Univ. Kansas Press.
Yabe, H., Nagao, T. and Shimizu, S. (1926): Cretaceous
mollusca from the Sanchu Gaben in the Kanto
Mountainland, Japan. Sci. Rep. Tohoku Imp.
Univ., 9, 2 p. 33-76, pis. 12-15.
Ziegler, P. A. (1982): Geological Atlas of Western andCentral Europe. l30p., 40 figs., Shell Intern. Petrol.
Maatschap. B. V.

is:jil:j']:li!:I!!Ii£O)/~ V ~ 7 Yj\)J7 Y'£-7"1 r-Jfi1Jnctb¥!.JllfO)fWtJT : LlJ<=Pi1!!..!ilXO)/~ v ~ 7 y:j']:li!:1!!I1J, Gi£1±l Lt-: 7 Y'£- 7" 1 r 5 1m 6 'fifl~il~~ Lt-:o :: td-=- J: I), is:1!!I1J' Gi£1±l Lt-: 7 Y '£- 7" 1
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llft-=- fi Simbirskites (Milanowskia) 0) -'fiflIJ~1!; i.h ~ iJ~, :: .hfi 3 - P .y /{ 0) <J' v 7 ;L--}1l!..!ilXIJ, G~t~ifllHI-C'¥$illb L -C ~ t-: ~ o)-C;lt) ~ c flUE~.h ~t'~JIIIEtJi,J
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 149, pp. 417 -429, April 30, 1988

855. PLIOCENE FRESHWATER BIVALVES (LAMPROTULA AND CUNEOPSIS:UNIONIDAE) FROM THE IGA FORMATION, MIE PREFECTURE,
CENTRAL JAP AN*
KEIJI MATSUOKA
Toyohashi Museum of Natural History, 1-238 Oana, Oiwa, Toyohashi 441-31

Abstract. The following six species of freshwater bivalves are described from the
Pliocene Iga Formation in the Ueno Basin, central Japan: Lamprotula (Sinolamprotula) sp. A,
L. (Parunio) sp. B, L. sp., Cuneopsis (Cuneopsis) gracilenta, n. sp., C. (Procuneopsis) okuyamai,n. sp., and C. (Tchangsinaia) praemaxima, n. sp. These unionid species of the genera
Lamprotula and Cuneopsis become extinct in the Japanese Islands in Recent times, but they arestill living in China. These bivalves are the only record of this age known from the JapaneseIslands.

sp. A, L. (Parunio) sp. B, L. sp., Cuneopsis(Cuneopsis) gracilenta, n. sp., C. (Procuneopsis)
okuyamai, n. sp., and C. (Tchangsinaia)praemaxima, n. sp. are described and figured.
This is one of a series of papers dealing withinvertebrate fossils from the Kobiwako Group.

Introduction
The freshwater fossil flora and fauna fromthe Iga Formation of the Kobiwako Group arequite unique among the Pliocene fossil biota ofJapan (Okuyama, 1981b, c, 1983-1986; Nakajima et al., 1983; Matsuoka, 1985a, 1987; Tanaka
and Matsuoka, 1985; Nakajima, 1986). Matsuoka(1985a) proposed a name of the Iga non-marine
fauna for freshwater fossils from the Iga Formation in the eastern part of the Ueno Basin, Mie
Prefecture.
Both of Iga non-marine molluscan fauna I andII (Matsuoka, 1987) belonging to the Iga
non-marine fauna differ from the Miocene
Sasebo non-marine molluscan fauna (Suzuki,1941) and other Pleistocene non-marine
molluscan faunas (Matsuoka and Nakamura,
1981 ; Matsuoka, 1983, 1986, 1987) in the faunalcompositions. The Iga non-marine molluscan
faunas consist of some 20 species including many
new species. Among them, one new genus andfour new species of the viviparid and pleurocerid
have been already described (Matsuoka, 1985b).
In this paper, Lamprotula (Sinolamprotula)

Geology and localities
The Iga Formation is distributed mainly in the
Ueno (Iga) Basin. Fossiliferous sequences of thenon-marine Iga Formation are well exposed in
the Ohyamada area in the eastern part of thebasin. Stratigraphic studies of the Iga Formation
were made by Takaya (1963), Kondo (1968),
Okuyama (1981a), and Yokoyama et al. (1980).
The Iga Formation consists mainly of silt, sand,and gravel, and intercalates many volcanic ash
layers. It is divided into three lithological units inascending order: Ohyamada Clays, Kashikimura
Sands, and Aitagawa Alternations. The
Ohyamada Clays yield abundant diatoms,
sponges, molluscs, fishes, and other fossils. The
fossil molluscs described in this paper wereobtained from six localities of the Ohyamada
Clays (Figure 1). The Hattorigawa volcania ash
layer in the lower part of the Ohyamada Clays*Received October 21, 1987; Revised manuscript
accepted March 18, 1988.
417

Keiji M4 TSUOKA

418.:"~ molluscs~v volcanic asho. graveli:;:c:: sand-:- silt

34'

Loc.6(I25)-~~

Ohyamada-mura..~~- - -......

-- ~I"I,=34' 45'.JL -_~-=~~

Ueno City"-"

I

ii.... -: _

_

..1'_

-~---:.

_-.1\_.£\-- :.

"\

\

-~-

'"'-'.:I~~~

R.Kume0136·12'

Figure 1.

I

!wwI

!wwI

!wwl

'"

\
L--. "'

1 km!wwl136·14'

Locations of six geologic sections where fossil molluscs were collected"

856. Freshwater bivalves from Iga Formation

was dated 4.6 Ma by the fission track method(Yokoyama et al., 1980).
Locality 1(12): A riverbed on the north sideof the Hattori River at Hata, Ohyamada-mura,
Ayama-gun (lat. 34°46'N, long. 136° 12'7"E).
A gray silt exposed on the riverbed of the
Hattori River includes two volcanic ash layers:
Hattorigawa I and II, and yields wood, leaves,diatoms, protozoans, sponges, molluscs, fishes,and reptiles. The fossil molluscs were collectedfrom a gray silt about 1 to 2 m below the
Hattorigawa II volcanic ash layer. The molluscan
assemblage consists of Igapaludina stricta,
Anodonta (Pleioanodonta) sp., Semisulcospira(Biwamelania)
praemultigranosa,
Bellamya
suzukii, Psilunio sp., Acuticosta sp., Lamprotulasp., Cuneopsis gracilenta, n. sp., C. okuyamai, n.
sp., Unio (Nodularia) sp., Inversidens sp., andLepidodesma sp.
From this locality, Okuyama (1981b, c,
1983-1986) collected and illustrated many
fossils such as macroscopic plants, molluscs,
cyprinid fishes, turtles, and crocodiles.
Kobayakawa and Okuyama (1984) reported asilulid fish, Silurus sp.
Locality 2 (l4a): A riverbed on the south sideof the Hattori River, Hirata, Ohyamada-mura,
Ayama-gun (lat. 34°45'35"N, long. 136°12'59"
E).
A silty sand below 5 to 16 cm thick volcanic
ash layer yields the following species, Igapaludina
stricta, Semisulcospira (Biwamelania) praemultigranosa, S. (B.) sp., Lepidodesma sp., Unio
(Nodularia) sp., Inversidens sp., Acuticosta sp.,
Cuneopsis gracilenta, n. sp., and Lamprotula(Sinolamprotula) sp. A. They are preserved asmolds in silty sand including coarse grains ofquartz and plant fragments.
Locality 3 (l4b): A riverbed on the north sideof the Hattori River, Hirata, Ohyamada-mura,
Ayama-gun.
This locality is on the opposite side of the
Locality 2. The following well-preserved molluscs
from silt blocks, which had been derived fromupstream outcrops of the Hattori River, werecollected by Okuyama: Igapaludina stricta,
Lamprotula (Parunio) sp. B, Acuticosta sp.,

419

Anodonta (Pleioanodonta) sp., Psilunio sp.,
Cuneopsis okuyamai, n. sp., C. praemaxima, n.
sp., C. gracilenta, n. sp., Unio (Nodularia) sp.,
Lepidodesma sp., and Inversidens sp. The
horizon of the fossiliferous bed is probably a few
meters above the Nakamura volcanic ash layer.
Locality 4 (15): An exposure on the northside of the Hattori River, about 1 km southeastof Hirata, Ohyamadamura, Ayama-gun (lat. 34°
45'29"N, long. 136°13'37"E).
A gray silt exposed on the riverbed yields
abundant sponges, molluscs, and fishes. The
molluscan assemblage consists of Igapaludina
stricta, Semisulcospira (Biwamelania) praemultigranosa, Cuneopsis okuyamai, n. sp., C. graci/enta,
n. sp., and Inversidens sp. An interbedded volcanic ash layer with the fossiliferous bed issituated stratigraphically several meters above the
Nakamura volcanic ash layer.
Locality 5 (16): A riverbed on the west side of
the Hattori River, Nakamura, Ohyamada-mura,
Ayama-gun (lat. 34°45'19"N, long. 136°13'44"E).
A fossiliferous silt bed is exposed on theriverbed. Igapaludina stricta, Bellamya suzukii,
Semisulcospira (Biwamelania) praemultigranosa,
Cuneopsis okuyamai, n. sp., C. gracilenta, n. sp.,Inversidens sp., and Acuticosta sp. were obtained
from a gray silt about 1 m above the Nakamura
volcanic ash layer. The volcanic ash is intercalated in the lower part of the Ohyamada Clays.
From this locality, the following fossils havebeen recorded: diatoms (Tomoda and Negoro,
1979; Negoro, 1981; Tanaka and Matsuoka,
1985), and fishes (Nakajima et al., 1983;
Nakajima, 1984).
Locality 6 (125): A cliff about 500 m north of
Hasuike, Ueno City (lat. 34°44'23"N, long. 136°
12'51"E).
A gray silt exposed in this cliff yields wood
fragments, sponges, molluscs, and fishes, andincludes one volcanic ash layer. There occur suchwell-preserved molluscan fossils as Igapaludina
stricta, Semisulcospira (Biwamelania) praemultigranosa, Cuneopsis praemaxima, n. sp., Inversidens sp., and Anodonta (Pleioanodonta) sp. The
volcanic ash layer is probably correlated with

Keiji M4 TSUOKA420

the Hattorigawa II volcanic ash.
Systematic paleontology
All the illustrated specimens are deposited in
the collection of Department of Earth Sciences,
Nagoya University, Nagoya (ESN) and in the
Mizunami Fossil Museum, Gifu (MFM).

Lamprotula (S.) leai (Gray), Early Pleistocene
Sanmen Formation in Sanmenxia, Hunan
Province and Early Pleistocene Nihowan Formation in Sanggan He, Hubei Province (Wan, 1961),
Latest Pleistocene Zhenpiyan Cave Deposits in
Guilin, Guangxi Province (Huang, 1981), andLamprotula (S.) obovata Huang from the Latest
Zhenpiyan Cave Deposits in Guilin, Guangxi
Province (Huang, 1981).

Class Bivalvia
Family Unionidae Fleming, 1828
Genus Lamprotula Simpson, 1900
Type species: - Chama plumbea Chemnitz, byoriginal designation. Recent. Southeast Asia.
Geologic range:- Jurassic to Recent.
Discussion:-The genus Lamprotula wasoriginally proposed as a subgenus of Quadrula
Rafinesque, 1820 by Simpson (1900). The
monograph "Fossil Lamellibranchia of China,"
published in 1976 by Nanjing Institute ofGeology and Palaeontology, divided the genus
Lamprotula into nine subgenera including two
new subgenera: Lamprotula (s. str.) Simpson,
1900; Eolamprotula Ku, 1962; Sinolamprotula
Gu et Huang, 1976; Odhnerella Modell, 1964;
Sulcatula Leroy, 1940; Cuneolamprotula Ku et
Huang, 1976; Paruino Ping 1930; Scrip to lamprotula Modell, 1964.
Recent species of the present genus distributed in Korea, China, and Vietnam.
Subgenus Sinolamprotula Gu et Huang, 1976
Type species:- Unio leai Gray, 1836, byoriginal designation. Recent. Vietnam, China,
and Korea.
Geologic range:-Pliocene to Recent.
Discussion:-The subgenus Sinolamprotula
was proposed as a replacement name ofRichthofenia Modell, 1964 by Gu and Huang(1976). This name is equivalent to Simpson's(1900) Quadrula leai group. The juvenile shell issimilar to that of the genus Inversidens (Modell,1964).
Fossils of the subgenus Sinolamprotula havebeen reported only from China as follows:

Lamprotula (Sinolamprotula) sp. A
Figures 2, 4-3

1985 Lamprotula sp., Matsuoka, Monogr. Assoc. Ceol.
Call. Japan. No. 29, p. 79.

Material studied:-ESN-40 10 1
Description and discussion:-This description
is based on inner and outer molds of one left
valve. The ventral and posterior margins of theinner mold are distorted, but it preserves its
characteristic shape, hinge and muscle scars of
the subgenus Sinolamprotula.
Shell rhomboidal, attaining 54.5 mm inlength, and moderately inflated. Beak low,
located at anterior part of shell. Dentition
imperfectly known; left valve possessing two
pseudocardinal teeth, two lateral teeth, andnarrow interdentum. Dorsal pseudocardinal
tooth stout, compressed trianguloid, and ventral
one feeble with a wide socket. Lateral teeth
weakly curved, subparallel to shell axis. Angle
formed between pseudocardinal tooth and lateral

1 em
Figure 2. Inner surface of Lamprotula (Sino·
lamprotula) sp. A.

856. Freshwater bivalves from Jga Formation

one measuring 17 degrees. Anterior adductor
muscular scar deep, subtrianguloid, and attaining
8.4 mm in maximum diameter. Pedal protractor
muscle scar adjoined posteriorly adductor scar,and reaching 3.3 rnm in maximum diameter.
Pallial line indistinct.
Hinge and muscular scars of this species
resemble those of the Recent Lamprotula (S.)
leai (Gray).
Stratigraphic tange:-Restricted to the Ohyamada Clays, Iga Formation.
Occurrence:-Locality 2 (I4a).
Subgenus Parunio Ping, 1931
Type species: -Parunio crassus Ping (= Lamprotula antiqua Odhner), by original designation.
Pliocene. Tungur, Mongolia.
Geologic range:-Oligo-Miocene to Recent.
Discussion: - Those Oligocene to Miocene
species identified under the subgenus Parunio
have been reported from the Tanhoi Formation,southern district of Lake Baikal (popova, 1981),
northwestern district of Lake Barun-Torei,
Siberia (Popova et al., 1974), Nojima Group,
Nagasaki, southwest Japan (Ueji, 1934), andKoura Formation, Shimane, southwest Japan(Matsuoka, 1985a). For the Pliocene, tworecords come from Mongolia (Ping, 1931) andJapan (in this paper). Those species from
Pleistocene sediments, as well as the Recent
species of this subgenus, are limited to theeastern district of China (Odhner, 1925; Leroy,
1940; Otuka, 1940; Chow, 1958; Wan, 1961).
Lamprotula (Parunio) sp. B
Figures 3; 4-1a, b
1985 Lamprotula (Parunio) sp., Matsuoka, Monogr.
Assoc. Ceol. Coil. Japan, No. 29, p. 79.

Material studied:- ESN-40084
Description:-Shell ovate, medium in size,
thick, somewhat inflated. Postero-dorsal margin
slightly curved, forming a somewhat sharp angle
with inclined ventral margin. Anterior margin
narrow, slightly truncated. Hinge ligament
conspicuous, located posteriorly to beaks,

421

1em
Figure 3.
sp. B.

Inner surface of Lamprotula (Parunio)

reaching 29 mm in length. Posterior ridge
inconspicuous. Beaks prominent, oblique, andsituated in anterior part of shell. Dentition
imperfectly known; left valve bearing at least two
pseudocardinal teeth; dorsal one large and long,ventral one short; lateral one unknown. Hinge
plate wide. Beak cavities moderately deep,
possessing several muscle scars. Anterior
adductor muscle scar subtriangle, deep, irregularwith serrate-surface, attaining 8.8 rnm in
maximum length, and pedal protractor muscle
scar located within muscle scar, elliptical, deep,
and reaching 2.6 mm in maximum length.
Anterior pedal retractor muscle scar situatedbelow lower pseudocardinal tooth, elongated,and measuring 2.7 mm in maximum length.
Surface of shell shiny bearing irregularly
undulated concentric with pustules.
Discussion: -Obtained shell is a pair ofimperfect attached valves measuring 44.7 rnm inlength and 43.5 mm in width. This is the firstrecord of this subgenus from the Pliocene strata
in Japan.
The shape of Lamprotula (Parunio) sp. B has asimilarity to L. nojimensis, which has been
described by Ueji (1934) from the Miocene
Nojima Group, Nagasaki, southwest Japan. The
present Pliocene species is more ovate in outline
with a curved dorsal margin and pustulated
SCUlpture. On the shell surface Lamprotula(Parunio) sp. B is distinguished the shell surface
from L. antiqua Odhner by its ventral margin andbeak SCUlpture. The latter species from the EarlyKeijiMATSUOKA
422

856. Freshwater bivalves from Jga Formation

Pleistocene Sanmen Formation in Sanxi
Province, China has a more rounded ventral
margin and well-developed beak sculpture. This
species resembles L. wangi from the Pleistoceneof Sanxi Province, China with respect to its shell
outline, but the former lacks the undulated
sculpture on the shell surface and possesses
angular posterior end.
Stratigraphic range:-Ohyamada Clays of Iga
Formation.
Occurrence:-Locality 1 (12).
Lamprotula sp.
Figure 4-2

Discussion :-Two
incomplete
specimens(ESN-40 136, MFM-I11207) of Lamprotula
which do not appear to belong to any of theabove-listed species were collected from the
Ohyamada Clays.
Stratigraphic range:-Ohyamada Clays of Iga
Formation.
Occurrence:-Locality 1 (12).

Genus Cuneopsis Simpson, 1900
Type species:-Unio celtiformis Heude, byoriginal designation. Recent. Jiangxi and Hunan
Province, China.
Geologic range:-Jurassic to Recent.
Discussion:-Modell (1964) divided the genus
Cuneopsis into three types on the basis of shell
outline: long-knife type (including Cuneopsis
celtiformis), long-tongue type (including C.
pisciculus), and wedge type (including C.
heudei). Starobogatov (1970) created threesubgenera, Cuneopsis s. str., Tchangsinaia, andProcuneopsis, which correspond to the three
types described by Modell (1964), respectively.
Five Recent species of this genus are known
from central China and north Vietnam (Haas,
1969). In Japan, the geological distribution of423

the genus Cuneopsis ranges from Miocene to
Pliocene (Mizuno, 1966 and Matsuoka, 1985a).
Subgenus Cuneopsis s. str.
This subgenus is similar to the genus
Lanceolaria in its shell outline. Lanceolaria ismore inequilateral, and its posterior and isangular.
The fossil species which can be included in thesubgenus Cuneopsis are as follows: Cuneopsis
zhenpiyanensis and C. subceltiformis from thelatest Pleistocene Zhenpiyan Cave Deposits inGuangxi Province, China (Huang, 1981), C.
nagahamai from Fukazuki Formation of theNojima Group, Kurosaki, Kitamatsuura-gun,Nagasaki Prefecture, Japan (Mizuno, 1966), andC. wershutskyi from the Oligo-Miocene Tanhoi
Formation to the south of Lake Baikal, U.S.S.R.
(popova, 1981).
Cuneopsis (Cuneopsis) gracilenta, n. sp.
Figures 5-1-4
1985 Cuneopsis gracilenta, Matsuoka, Monogr. Assoc.
Ceol. Coil. Japan, No. 29, p. 79 (nomen nudum).

Type locality: - Hata (locality(12)),
Ohyamada-mura, Ayama-gun, Mie Prefecture.
Derivation of name:-gracilentus, Latin for
slender.
Material studied:-ESN-40097-40099, 40117,
40139, MFM-I10026, 110027, 111066-111068,
111090,111091,111204,111205.
Diagnosis:-Beaks located anteriorly at one
fifth the length of shell. Length about threetimes the height. Anterior margin narrowly
rounded, truncated above the medial line.
Description:-Shell medium, reaching 86 mm
in length. Outline elongated trapezoid, about
lance-head shaped, width being three times aslong as height. Valves compressed, thickened+- Figure 4. All figures are x 1. la, b. Lamprotula (Parunio) sp. B. ESN-40084, left valve; a, outer surface; b,
inner surface. 2. Lamprotula sp. ESN-40136, attached valves. 3. Lamprotula (Sinolamprotula) sp. A. ESN-40101,
inner mold of left valveAa-c. Cuneopsis(Tchangsinaia) praemaxima, n. sp. MFM-ll0018 (paratype 1), attachedvalves; a, left side; b, dorsal side; c, right side. Sa, b. Cuneopsis (Tchangsinaia) praemaxima, n. sp. ESN-40081(holotype), attached valves; a, left side; b, right side. ESN-40084, 40136, and MFM-ll0018 were collected by
Okuyama.

424

Keiji M4 TS UOKA

5b

6a

6b

856. Freshwater bivalves from Jga Formation

anteriority, inequilateral. Dorsal margin long,straight, sloping downward. Posterior margin
narrowly rounded, forming a broad angle withthe obliquely descending posterior margin.
Ventral margin almost straight, sometimes
moderately convex medially. Anterior margin
slightly truncated. External ligament distinct,
long, located posteriorly of beaks. Posterior ridge
conspicuous, bluntly pointed. Posterior slope,narrow, compressed. Beaks low, elevated above
the hinge line, situated anteriorly at one fifth theshell length. Surface of shell marked only withconcentric lines of growth. Right valve with one
pseudocardinal tooth, compressed triangular, nointerdentum, and one lateral tooth; left valve
with two compressed pseudocardinal teeth, its
anterior one large, and two lateral teeth, ventral
one distinct. Beak cavities shallow. Anterior
adductor muscle scar deep, trianguloid; posterior
muscle scar sub circular , anterior side distinct;
pedal protractor located in the front of theanterior adductor muscle scar, orbiculoid, distinct. Pallial line distinct anteriorly in particular.
Discussion: -In its general shape, Cuneopsis
gracilenta is similar to Cuneopsis celtiformis(Heude) distributed in Jiangxi and Hunan
Provinces of China (Heude, 1874), C. subceltiformis Huang and C. zhenpiyanensis Huang,both from the latest Pleistocene of theZhenpiyan Cave Deposits in Guangxi Province ofChina (Huang, 1981), C. nagahamai from the
Miocene Nojima Group, Nagasaki, southwest
Japan (Mizuno, 1966). This new species isdistinguished from four other species by theshapes of posterior margin, beak, and hinge plate.
Measurements (in mm):ESN-40117 (holotype)
ESN-40097 (para type 1)
ESN-40099 (para type 2)

Length
84.2
70.3
75.1

Height
32.0
26.0
30.3

MFM-110026 (paratype 3)
MFM-110027 (paratype 4)
MFM-110036 (paratype 5)
MFM-I11067
MFM-III091
MFM-I11090
ESN-40098
MFM-111068

425
64.8
56.2
61.5
86.4
68.0
77+
68.7
62.2

26.4
24.2
27.3
29.8
24.4
31.9
27 +
23.2

Stratigraphic range:-Iga Formation to Koka
Formation (Matsuoka, 1987).
Occurrence:-Localities 1 (12), 4 (15), and 5(16).

Subgenus Procuneopsis Starobogatov, 1970
Type species:-Unio heudei Heude, 1874 byoriginal designation. Recent. Hebei, Shan dong,
Zhejiang, Jiangsu, Anhui, Jiangxi, Hubei, andHunan Provinces in central China.
Discussion:-Ten species from U.S.S.R. are
included in Cuneopsis (Procuneopsis) by Popova(1981). This subgenus also contains probably
Cenozoic species, C. barbouri King, 1926, C.
heudeinihowanensis
Otuka,
1942,
C.
shanghaiensis Huang et Lan, 1976, C. crassata
Huang et Wei, 1976, and C. subovata Huang etWei, 1976. The latter four have been reported
from China (Nanjing Inst. Geol. and Palaeont.,
1976). The geographical distribution of Procuneopsis in the Eocene to Pliocene interval isrestricted to Siberia (north China, Mongolia,
south Baikal, and south Primorskij).
Cuneopsis (Procuneopsis) okuyamai, n. sp.
Figures 5-5a, b, 6a, b, 7-9
1985 Cuneopsis okuyamai, Matsuoka, Monogr. Assoc.
Geol. Coli. Japan, No. 29, p. 79 (nomen nudum).

Type locality: - Hirata,
Ayama-gun, Mie Prefecture.

Ohyamada-mura,+- Figure S. All figures are x 1. 1-4. Cuneopsis (Cuneopsis) gracilenta, n. sp. 1, ESN-40117 (holotype). 2,
ESN-40097 (paratype 1), Right valve, outer mold. 3, MFM-l10036 (paratype 5), Left valve. 4, ESN-40139, innermold. 5-9. Cuneopsis (Procuneopsis) okuyamai, n. sp. Sa-b, ESN-40086 (holotype), attached valves. 6a-b,
MFM-110021 (paratype 2). 7, MFM-110022 (paratype 3), inner surface of right valve. 8, ESN-40137, attachedvalves. 9, ESN-40138, attached valves. ESN-40086, 40097, 40117, 40137-40139, MFM-110021, 110022, and110036 were collected by Okuyama.

426

KeijiMATSUOKA

Derivation of name:- Named in honor of Mr.
Shigemi Okuyama who expended great efforts incollecting molluscs from the Iga Formation.
Material studied:-ESN-40086-40090, 40137,
40138, MFM-I10021, 110022, 111059-111061,
111092-111094, 111206.
Diagnosis:-Shell medium, sub angular. Beaks
situated at extreme end of antero-dorsal margin.
Posterior margin medially pointed. Ventral
margin straight.
Description:-Shell subtriangular, medium,
reaching 50 mm in length. Valves subinflated,
especially inflated in area below beaks, solid,
inequilateral. Dorsal margin slightly convex.
Posterior margin medially pointed. Ventral
margin straight, with moderate slope in anterior
part. Anterior margin broadly rounded or slightly
truncated. Hinge ligament short and inconspicuous, located posteriorly of beaks. Posteriorridges fairly well defined near beaks, becoming
faint toward posterior margin. Posterior slope
slightly concave. Beaks full, oblique, elevatedabove hinge line, located near anterior end ofshell, their sculpture consisting of a few weakradial ridges. Surface of shell generally smoothwith fine growth lines. Right valve with onecompressed, triangular, and serrated pseudocardinal tooth; no interdentum; almost straight,
long, main lateral tooth, and another short toothin postero-ventral hinge plate. Left valve withtwo compressed pseudocardinal' teeth and probably two lateral teeth. Beak cavities moderately
deep. Anterior adductor muscle scar deep,
semicircular. Anterior pedal retractor muscle scarrough, distinct, small, located in front of anterior
adductor one. Pedal protractor muscle scar under
the hinge plate. Posterior adductor muscle scar
sub quadrangular, smooth, defined, especiallyanterior side. Posterior pedal retractor muscle
scar deep, small, trapezoid, located underposterior end of lateral tooth. Pallial line distinct
anteriorly.
Discussion: -Cuneopsis okuyamai is very
similar to the Recent species Cuneopsis capitata
which is distributed in Anhui, Zhejiang, Jiangsu,
Jiangxi, Hubei, and Hunan Provinces of central
China. The present new species differs from the

Recent in having a narrow hinge plate andstraight ventral margin having a wider hinge plateand concaved ventral margin. Cuneopsis
shanghaiensis from the Quaternary of Shanghai,
Jiangsu Province, China differs from Cuneopsis
okuyamai, in possessing major ridges on the shellsurface and a concaved ventral margin. Those
specimens figured by Huang (1964) as Cuneopsis
martinsoni from the Tanhoi Formation to thesouth of Lake Baikal is closely related to thisnew species. However, the posterior margin of
the latter species is moderately pointed, and theventral margin is straight.
Measurements (in mm):ESN-40086 (holotype)
ESN-40087 (paratype 1)
MFM-l10021 (paratype 2)
MFM-l10022 (paratype 3)
MFM-l11059
ESN-40088
MFM-l11094
ESN-40085

Length
40.0
45.0
34 +
39.5
48.2
44 +
34.7
14.8

Height Width
26,6
19,8
27,0
20.3
21.8
31.7
30.3
12.2
24.7
14.6
22.0
24.4
15 +
13.1

Stratigraphic range:-Ohyamada Clays to
Aitagawa Alternations of Iga Formation.
Occurrence:-Localities 1(12), 3(14b), and5(16).
Subgenus Tchangsinaia Starobogatov, 1970
Type species:-Unio p iscicu Ius Heude, byoriginal designation. Recent. Jiangsu, Anhui,
Zhejiang, Jiangxi, Hubei, and Hunan Provinces incentral China.
Discussion:-Among Neogene bivalves of the
Kyzylgir Formation in the Chujskaya Basin, Altai
Highlands, Popova (1981) assigned three species
to the present subgenus, namely Cuneopsis
tuerykensis Popova et Star, C. elongata Kursal,
and C. pliocaenica Kursal. Cuneopsis teilhardi
Leroy, 1940 and C. maxima Odhner, 1925 fromthe Tertiary of China are herein included in thissubgenus. AnlOng the Recent species of
Cuneopsis, only C. piscicula belongs to thepresent subgenus.

856. Freshwater bivalves from /ga Formation
Cuneopsis (Tchangsinaia) praemaxima, n. Sp.
Figures 4-4a-c, 5a-b

1985 Cuneopsis praemaxima, Matsuoka, Monogr.
Assoc. Geol. Coli. Japan, No. 29, p. 79 (nomen
nudum).

Type loca/ity:-Hojiro (locality 6(125», Ueno
City, Mie Prefecture.
Derivation of name:-Prae-, Latin for before,+ maxima, Latin for large.
Material studied:-ESN-40081 and MFM110018.
Diagnosis:-Shell medium, comparatively
solid, elliptical. Length about two times larger
than height. Beaks situated anteriorly at one
seventh to one eighth the shell length. Dorsal
margin moderately curved. Hinge ligament
occupied about half of dorsal margin.
Description:-Shell medium, reaching 79 mm
in length, elliptical, asymmetric about two times
as long as height. Commissure of plane dextrally
twisted. Left valve overhanging right valve.
Valvesinequilateralcomparatively
solid,
especially anterior part compressed, inflated inarea below beaks. Dorsal margin regularly andslightly convex. Posterior margin broadly
rounded or somewhat truncated. Ventral margin
slightly curved, with concave middle part, andresuming the same curvative up to anterior
margin. Anterior margin broadly rounded. Hinge
ligament conspicuous, long, nearly half thelength of dorsal margin, located posteriorly of
beaks. Posterior ridge bluntly rounded. Posterior
slope, narrow, straight or concave. Beaks full,prosogyrate, eroded, somewhat elevated above
hinge line, and located anteriorly at one seventh
to one eighth the shell length. Surface of shell
bearing only concentric lines of growth. Dentition partially known; left valve with one stout
pseudocardinal tooth; right valve possessing atleast one pesudocardinal tooth, and a long andslender lateral tooth. Beak cavities shallow.
Anterior adductor muscle scar deep, ovaltrianguloid, serrated. Pedal protractor muscle
scar elliptical, compressed, and located behindanterior adductor muscle scar. Anterior pedal
retractor muscle scar defined. Posterior adductor

427

muscle scar unknown. Pallial line distinct.
Discussion:-In the shape of shell outline, the
new species is very similar to Cuneopsis maxima,
a species reported from the Early Pleistocene
Sanmen Formation in Shanxi Province (Odhner,
1925; Leroy, 1940; Otuka, 1940), and from the
Early Pleistocene Nihowan Formation in
Qingshuihe, Hei Monggol Province (Zhu, 1976,
1940). Cuneopsis praemaxima differs from C.
maxima in having a smaller shell, more anteriorly
situated beaks, no beak sculpture, and narrower ahinge plate. The shell shape suggests its
relationship with Cuneopsis maxima, which has
been considered to be an ancestor of three
Recent species, Cuneopsis heudei, C. piscicula,and C. capitata (Odhner, 1925). I believe that
Cuneopsis maxima is also a descendant of C.
praemaxima.
The Recent species Cuneopsis piscicula(Heude) distributed in Anhui, Zhejiang, Jiangsu,
Jiangxi, Hubei and Hunan Provinces in China, hasa twisted shell similar to that of C. praemaxima,
but it differs from the present species having ashorter elliptical shell.
Measurements (in mm):ESN-40081 (holotype)
MFM-II0018 (paratype 1)

Length79.1
65.8

Height Width
34.3
19.4
40.1
26.3

Stratigraphic range: Ohyamada Clays of Iga
Formation.
Occurrence: Localities 3 (l4b) and 6 (125).
Acknowledgments
I am grateful to Professor J. Itoigawa of
Nagoya University for reading the manuscript
and offering valuable suggestions. I wish to thankMr. S. Okuyama for placing those fossil bivalvesfrom the Iga Formation at my disposal, andProfessor B. Huang of Nanjing Institute ofGeology and Palaeontology for providing theliterature used in this study.
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Keiji MA TSUOKA

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429

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implications for vicariance event ······Ishizaki, K.

Acanthohoplites

from

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fJ1:aJlJO)1:~~

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15El(~:T /' /7' '7 '/ lJ I- /'1:' t C -j <iJal7Ll*lt15m'
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If(;~mtJT··,······························,·······,**0@'~Fa~+~~

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1:.*'&P.l'11lfO) A it15O)M:!¥;tl1:lTh n·········¥~J~fID!2-nl;!J.7ot/~-~J5"x.G............. "1II: I~; A'R)jj Jif !:IJ • J!i ~ Eli'!=.

433

New members approved by the Council Meeting held on January 27, 1988
Ordinary Members:Baba, Kenji; Inoue, Keisuke; Kamikawa, Yoichi; Morino, Yoshihiro; Okuyama, Shigemi;
Shin, Jai Hoon; Yamaguchi, Keiko.
New fellows approved:
Akamatsu, Morio; Ando, Hisao; Kawamura, Yoshinari; Kikuchi, Yoshibumi; Hada,
Shigeki; Hase, Yoshitaka; Makino, Tooru; Nishiwaki, Niichi; Sato, Yoshio; Tanaka,
Kunio; Tanimura Yoshihiro; Tsugo, Yoshihiro; Yasuda, Hisato.
Withdrawing members:
Sawamura, Konosuke; Yoshida, Michio; Maeda, Shiro; Siida Isao; Talent, John A.;
Jordan, Reiner; Loeblich, Alfred R., Jr.
Deceased Member:
Takahashi, Eitaro (August 20, 1987)

ErrataNo. 148,844. KOJIMA, Satoru and MIZUTANI, Shinjiro:
Triassic and Jurassic Radiolaria from the Nadanhada Range, Northeast China
page

column

line

read

for

256

right

5

257

left

11

wide
age.

a wide
age

258

left

34

(delete)

269

right

28

Eucyritidiellum

fusulinaceans
Eucyritidie!um

No. 148,847. KONDO Yasuo:
Burrowing depth of in faunal bivalves-Observation of living species and its relation to shellmorphology
page
323

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

New Series No . 149

April 30, 1988

CONTENTS
TRANSACTIONS
851. KASE, Tomoki: Early Carboniferous (Visean) gastropods from the Hikoroichi Formation of the Kitakami Mountains, Northeast Japan . ...... .. . ... ... ... .. .. 361
852. NIKO , Shuji and IKEBE, Nobuo: Fisherites, a receptaculitid from the Middle
Ordovician of Pyongyang Coal-field .. ... . . .. . .. . . ... .... . .. .. .. ..... 371
853 . MATSUMOTO, Tatsuro , ASAI, Akito, HIRANO, Hiromichi and NODA, Masayuki :Some inoceramids (Bivalvia) from the Cenomanian (Cretaceous) of Japan-III , Three
species occurring commonly in the North-West Pacific region... ..... .. . .... 378

854 . MATSUKAWA, Masaki: Barremian ammonites from the Ishido Formation , Japan- Supplements and faunal analysis - . . . . ...... .. .. . . .. .. .. . . . . . . . . . . 396
855 . MATSUOKA , Keiji: Pliocene freshwater bivalves (Lamprotula and Cuneopsis: Unionidae) from the Iga Formation, Mie Prefecture , central Japan . . .. .. . . . . . . . . . . . 417
PROCEEDINGS .. .. .... .. . . .. . .... . ... .. .. . . . . . . . ... . .. ..... .. ... 430