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The fo ss il on the cover is Trilophodon sendaicus Mats umoto , an extinct eleph a nt, which
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 161, pp. 681-696, 7 Figs., April 30, 1991

912. EARLY TRIASSIC RADIOLARIANS FROM THE
OGAMATA FORMATION, KANTO MOUNTAINS,CENTRAL JAPAN. PART 2*
KA TSUO SASHIDA
Institute of'Geoscience, University of Tsukuba,
Ibaraki, 305

Abstract. Chert beds of the Ogamata Formation exposed in the upper reaches of the
Nakatsugawa River, Kanto Mountains, yield abundant Early Triassic radiolarians. A partof this radiolarian fauna including the spicular type Palaeoscenidiidae was already describedby the present author. As an addition to the study of this fauna, several newly discriminatedSpumellaria and Nasellaria are described in this paper. The families Palaeoscenidiidae
with a latticed shell, Pantanellidae, and Sponguriidae are included in the Spumellarians.
The families Eptingiidae having a spumellarian-like affinity and Acanthodesmiidae areclassified as Nassellaria in this study. Five new species, Archaeothamnulus okuchichibuensis, Parentactinia virgata, Pactarentinia koikei, Pantanellium? virgeum, and Spongostephanidium longispinosum, are proposed herein.
Key words.

Kanto Mountains, Ogamata Formation, radiolarian, Triassic.

(1985), and Yoshida (1986). These works,
however, were mainly focused on Middle toLate Triassic radiolarians. Biostratigraphic
and taxonomic works on Early Triassic
radiolarians are quite scarce except for myprevious study on the spicule type radiolarians from the Ogamata Formation (Sashida,1983).
Recently, I have succeeded to recover many
conodonts and radiolarians from the samerock sample and also another chert bed
exposed in the middle course of the Oyamazawa Valley near the type section of the
Ogamata Formation. These two samples
contain the same radiolarian fauna, which ischaracterized by abundant spumellarians,
such as those belonging to the families Palaeoscenidiidae of both spicule and latticed
shell types, Pantanellidae, Sponguriidae andseveral unidentified genera and species ofEntactiniidae. The families Eptingiidae of
spumellarian-like N assellaria and Acanth-Introduction

Our knowledge of Mesozoic radiolarianbiostratigraphy has been rapidly increased invarious land sections and through the DeepSea Drilling Project for the past 15 years.
However, Triassic radiolarian biostratigraphy
seems to be rather behindhand compared withJurassic and Cretaceous ones because of thelack of continuous rock sequence including
well-preserved radiolarians except for theUpper Triassic in the Pacific Coast Side ofNorth America (e.g., Blome, 1984) and Middle Triassic calcareous rocks in the European
Tethys (e.g., Kozur and Mostler, 198 I ;Dumitrica, 1982). In Japan, several biostratigraphic works through the taxonomic
studies of Triassic radiolarians were also
undertaken by Nakaseko and Nishimura(1979), Yao (1982), Kishida and Hisada'Received March 26, 1990; revised manuscript accepted December 20, 1990
681

Katsuo Sash ida

682

odesmiidae are also discriminated in thisfauna.
The geologic age of this radiolarian faunais estimated as a late Spathian based on theco-occurring conodonts.

River in Saitama Prefecture, was investigatedby Fuj imoto et al. (1950, 1957), Ishii (1962)
and, Ishii and Matsukawa (1980). Theseauthors regarded that the Ogamata Forma-

Acknowledgments
I thank Professor Hisayoshi Igo (Universityof Tsukuba) for his valuable suggestion andcritical reading of this manuscript. I also
thank Professor Toshio Koike (Yokohama
National University) for his identification of
conodonts. The present report was fundedin part by the Grant-in-Aid for Cooperative
Research from the Ministry of Education,
Science and Culture (Grant no. 01304008)
and the Project Research of the University ofTsukuba in 1988.

LegendOl

l@jA

t------~B--

coGeologic setting
The Ogamata Formation exposed in thearea of the upper reaches of the N akatsugawac::J. ...... .

Figure 1.

Map showing the sample localities.

Figure 2. Generalized columnar section of
the Ogamata Formation along the Oyamazawa
Valley. A: chert block, B: dark green to graysiliceous shale, C: chert, D: limestone, E:
basaltic rock, F: alternation of sandstone andshale, G: massive sandstone, H: black shale.

912.

Early Triassic Radiolarians from Kanlotion occupies the highest stratigraphic position of the so-called Chichibu System and is
the Upper Permian. Sato el al. (1981) and
Sash ida el al. (1982) , however, reported theoccurrence of Jurassic radiolarians from argillaceous rocks of this formation and pointed
out the necessity to revise the stratigraphy,
geologic structure, and geochronology of the
Ogamata Formation .
I summarized the stratigraphy of this formation in the type section, as follows (Sashida , 1983). The Ogamata Formation , about
1,200 m thick , consists mainly of argillaceous
rocks including abundant exotic blocks of
Carboniferous to Middle Permian limestone,
Early Triassic to Early Jurassic chert, basaltic
rock and sandstone. Based on the sedimentological features , the Ogamata Formation ispresumed to be a subduction product ofoceanic plate during the Middle Jurassic
Period .
Recently, I have established the detailed
superficial stratigraphy of this formation in
the Oyamazawa Valley (Figures I, 2) as follows in ascending order.

683

1. Medium- to fine-grained massive sandstone; more th a n 200 m thick .
2. Alternation of sandstone and shale;
about 40 m thick.
3. White massive limestone with basaltic
rock in the lower part ; about 50 m thick .
4. Black chert; about 40 m thick .
5. Dark green to gray siliceous shale;
about 30 m thick.
6. Fine-grained sandstone including chert
block s of about 5 m in the maximum diameterand intercal ati ng black shale in the lower
part ; about 60 m thick.
7. Dark gray to green chert; about 120 m
thick.
8. Coarse- to medium-grained sandstonefrequently including black shale beds andchert blocks in the lower to middle parts;
about 200 m thick.
9. Gray to green chert; more than ISO m
thick .
A thin-bedded black chert (OG - 2) of theuppermost part of Unit 4 yields Early Triassic
radiolarians, which are described in thispaper. Late Middle Jurassic radiolarians

Figure 3. 1- 3, Neospalhodus err. homeri (Bender), 1: OG - I ; 2: OG - 2 ; 3: OG - l. 4, Ozarkodina
sp. , OG - 2. 5, Ellisonia sp. , OG-I. Scale bars, A to C equal to 100,urn; A app lies to 2, B to 1,3,4, andC to 5.

Katsuo Sash ida

684

occur abundantly in Unit 5 and a black shaleinterbedded with the sandstone of Unit 8.
Chert beds of Unit 7 yield Middle to Late
Triassic radiolarians. Early Jurassic radiolarians were recovered from a green chert ofUnit 9. Each unit mentioned above is incontact with a northward dipping fault with agouge of less than 20 cm thick except for thecontact between Units 1 and 2. The alternation of sandstone and shale of Unit 2 conformably overlies the sandstone of Unit I.
Geologic age of samples

Two chert samples OG-I and OG-2 arestudied in this study. The former one is thesame sample, which was studied in my previous paper (Sashida, 1983). As mentioned
above, OG-2 was collected from Unit 4,
which crops out along the Oyamazawa Valley. These two samples yield abundant but
poorly preserved conodonts, such as
Neospathodus cfr. homeri (Bender),Xaniognathus sp., Ellisonia sp. and many
unidentified ramiform elements. N. homeri
is a well known age diagnostic species andhas been reported from an interval from the
Spathian to lower Anisian (Aegean) of
Greek, Nevada, Italy, West Pakistan, British
Columbia, Japan, and other areas.
Neospathodus timorensis (Nogami) is frequently associated with N. homeri in the
Aegean of the above mentioned areas, but theformer species does not found in samples
OG-I and OG-2. The geologic age of twochert samples including these conodonts is
considered to be Spath ian rather than early
Anisian.
Systematic description

All specimens described herein are deposited in the collection of Institute of Geoscience,
the University of Tsukuba (IGUT).

Suborder Spumellaria Ehrenberg, 1875
Superfamily Entactiniacea Riedel, 1967
Family Palaeoscenidiidae Riedel, 1967
emend. Furutani, 1983

Remarks. - Although the family Palaeoscenidiidae is one of the representative groupsof the Paleozoic Radiolaria, there are some
confusions regarding the diagnosis of thisfamily. The study of this group has beenreviewed by Furutani (1983, 1990), Goodbody (1986), and Wakamatsu et al. (1990).
The main differences of diagnosis of the family Palaeoscenidiidae between Furutani(1983) and Goodbody (1986) are obvious in
number and disposition of spicule and withor without latticed shell. Goodbody (1986)
emphasized the presence or absence of latticed shell as a distinguishing criterion
between the families Palaeoscenidiidae andEntactiniidae. Namely, the latter family ischaracterized by a latticed or spongy andessentially spherical shell. Recently, Furutani (1990) has discriminated seven types ofthe skeletal construction in Silurian to
Devonian spicule radiolarians. According
to his study, the basic skeleton of many genera belonging to the family Palaeoscenidiidae
differs clearly from that of the genera Haplentactinia Foreman, Entactinia Foreman,and others of the family Entactiniidae.
Among the Furutani's seven types of spicule,the Type 4 is a typical skeleton of the genus
Palaeoscenidium Defrandre, because it has
four apical and basal spines and a tetrahedral
projection, which is a perpendicular plane tothe median bar. This type is also almost thesame as that of the genera Pactarentinia
Furutani and Tlecerina Furutani bearing acompletely spherical outer shell. Furutani(1990) emphasized the skeletal similarity
among these genera and insisted that thediscriminative criteria between the families
Palaeoscenidiidae and Entactiniidae should
not be the presence or absence of latticed
outer shell but the basic nature of spiCUle
such as the number and disposition of912.

Early Triassic Radiolarians from Kanto

685

PA

T
AS

LB

AH

LA

_______A~ __ --.L
BH

BH

LSB
BS
BS

WB~

4

Figure 4. Schematically illustrated and measured parts of the subfamily Palaeoscenidiinae, and their letter
indices. 1: Archaeosemantis sp. 2: Pactarentinia koikei Sashida, n. sp. 3: Parentactinia virgata Sashida, n. sp. 4: Archaeothamnulus okuchichibuensis Sash ida, n. sp. Abbreviations; AH: apical hemisphere, BH: basal hemisphere, AS: apical spine, BS: basal spine, P A: primarily apical spine, LA: lengthof apical spine, LB: length of basal spine, WB: width of basal hemisphere, DS: diameter of shell, LSB :length of spine on the basal spine.

spicule. Wakamatsu et al. (1990) followed
the mentioned Furutani's opinion.
Dumitrica (1982) suggested that the postPaleozoic palaeoscenidiid-like taxa such as
the genera Archaeosemantis Dumitrica, Aarchaeothamnulus Dumitrica, and Palhindeolithus Deflandre should belong to two oreven three families. Wakamatsu et al. (1990)
also pointed out the possibility that these
post-Paleozoic forms may be transferred toanother or other families. However, thepresent Early Triassic forms with the basic
skeleton of four basal and apical spines havealmost the same skeletal features (Types 3 and4) shown by Furutani (1990) and the post-

Paleozoic palaeoscenidiid-like taxa have notyet been sufficiently documented. Under
these circumstances, I tentatively included
Early Triassic spicule radiolarians into thefamily Palaeoscenidiidae.
The measured parts and terminology of thefamily Palaeoscenidiidae treated in this paper
are shown in Figure 4.
Genus Archaeosemantis Dumitrica, 1978

Remarks. - Dumitrica (1978b) included
various morphological types of this genus
into one species, Archaeosemantis pterostephanus Dumitrica. Subsequently, he split

Katsuo Sash ida

686

this species into three species and pointed out
the homologies of skeletal disposition ofsome species of this genus with that of N assellaria (Dumitrica, 1982). One of the redefined species, A. pterostephanus is characterized by having one to three rings caused by
the fusion of spines, while A. cristianensis
Dumitrica does not have any ring. The
species criteria of these two species are thepresence or absence of ring formed by fusionof two or more spines. Another species, A.
longivirga Dumitrica has a long median bar
and eight dissimilar spines. The species of
the genus Archaeosemantis recovered fromthe Ogamata Formation do not have any
fused spines.

Archaeosemantis venusta Sashida
Figures 5-4-8

Archaeosemantis venusta Sashida, 1983, p. 171, pI. 36,
figs. I, 2, 4-9, non fig. 3.

Remarks. - In my previous paper, I incl uded the specimens with three apical and basal
spines into this species but excluded themfrom this species in the present study. This
species is easily distinguished from otherspecies of the genus Archaeosemantis byhaving four apical and basal spines and ashort median bar.
Occurrence. - Common in 00-1 and 002.
Types. - III ustrated specimens, Figure 5-4,00-1, lOUT -KS4231 ; Figure 5-5, 00-1,
IOUT-KS4216; Figure 5-6, 00-2, 10UTKS4239; Figure 5-7, 00-2, lOUT -KS4241 ;Figure 5-8, 00-1, lOUT -KS4240.
Archaeosemantis aff. venusta Sashida
Figure 5-9
Aff. Archaeosemantis venusta Sashida, 1983, p.171,
pI. 36, figs. I, 2, 4-9.

Description. - Spicule skeleton with eightspines ansmg from a short median bar.
Four basal spines are usually longer and the
distal half curved inward. Two from eachend of bar. Two basal spines are divergent
rather laterally and curved downward on ahalf. These two basal spines form an angleof 110-120 degrees with their opposite spines.
The divergent apical spines are rod-like anddistally tapering. The length of apical spines
is about a fourth to sixth of the basal spines.
Remarks. - The present specimens slightly
resemble A. cristianensis Dumitrica in having
laterally directed and slightly curved twobasal spines, however they commonly have
two apical spines. These specimens differ
from A. venusta in having two basal spines,
which are laterally directed and slightly
curved downward.
Dimensions.-Based on four specimens (in,urn): length of apical spine, 20-35 (average30); basal spine, 95-130 (average 105) ;width of basal hemisphere, 110-150 (average
130).
Occurrence. - Rare in 00-1 and 00-2.
Types. - Illustrated specimen, Figure 5-9,00-1, lOUT -KS3615.

Archaeosemantis sp.
Figures 5-1- 3

Aechaeosemantis venusta Sashida, 1983, pI. 36, fig. 3.
non figs. I, 2, 4-9.

Description. - Spicule commonly with fiveto six spines arising from a very short medianbar. Three basal spines having graduallytaperd end are long and distal half of two ofthem curved inward. One of the basal spines
is straight or its distal part curved outward.
Apical spines are two to three, long, conicaland diverge on a plane including a medianbar. These apical spines form an angle of
120 degrees with their neighborhood. In thecase of two apical spines, they form a line inalmost the same direction as a median bar.
All of spines are commonly armed with

912.

Early Triassic Radiolarians from Kanto

numerous very short spinules or nodes on
their lateral and external sides.
Remarks. - In the present study, the specimens having five to six spines are excluded
from Archaeosemantis venusta Sash ida, andthese forms are included into the present
unnamed species. This unnamed species isquite similar to Archaeosemantis cristianensis
Dumitrica in number of spines and generalfeature. The latter species, however, hasstrongly curved apical spines.
Dimensions. - Based on six specimens (in,urn): length of apical spines 18-22 (average20); length of basal spines 140-180 (average160); width of basal hemisphere 70-80 (average 75).
Occurrence. - Rare in OG-I and OG- 2.
Types. - III ustrated specimens, Figure 5-1,
OG-I, IGUT -KS4218; Figure 5-2, OG-2,
IGUT -KS4219; Figure 5-3, OG-I, IGUTKS4229.
Genus Archaeothamnulus Dumitrica, 1982
Archaeothamnulus okuchichibuensis
Sash ida, n. sp.
Figures 5-10- 14

Diagnosis. - Skeleton formed with four
sturdy basal spines and one short apicalspine. Basal spines commonly armed withnumerous spinules.
Description. - Skeleton consists commonlyof five spines diverging from the center ofshell. Four basal spines are rod-like, sturdy
and long. They diverge downward at anangle of about 30 degrees to the horizontal
plane including the center of shell, andcurved inward at about one third of basal
spines toward the end of basal region. Basalspines from the turning point to their end
stretch almost parallel to each other andgently taper. Numerous long conical spinesarise from the basal spines at a right angle tothem. The longest spine is present at theturning point of basal spines. The length ofthe longest spine is less than one third of687

basal spine from the turning point to end.
The length of verticils being shorter toward
the end of basal spine. Ten spines are counted on one basal spine in well-preserved specimens. Apical spine is short and conical, andits length is less than one tenth of basal spine.
Remarks. - This new species is characterized by its basal spines, which are parallel
each other, many long conical spines, and ashort apical spine. The present new species
is more or less similar to the specimen described by Dumitrica (1982, pI. 7, fig. 7) as
Archaeosemantis cfr. longivirga Dumitrica ingeneral feature and having verticils on thebasal spines. However, the former species
has only one short apical spine and rod-likespines. This new species also resembles
Archaeothamnulus ramosus Sashida, however, the latter species is easily distinguishedfrom the former by having outwardly diverging basal spines.
Dimensions. - Based on ten specimens (in,urn): length of basal spines 160-250 (average200); length of apical spines 15-25 (average20); length of spines on basal spines 20-25(average 23); width of basal hemisphere 120170 (average 150).
Etymology. - The species name is from thegeographical name, okuchichibu where the
Ogamata Formation crops out.
Occurrence. - Common in OG-I and OG2.

Types.-Holotype, Figure 5-13, OG-I,
IGUT -KS3773: Paratypes, Figure 5-10,
OG-2, IGUT -KS3623; Figure 5-11, OG-2,
IGUT-KS8993; Figure 5-12, OG-I, IGUTKS8994; Figure 5-14, OG-l, IGUT -KS3622.
Genus Parentactinia Dumitrica, 1978
Parentactinia nakatsugawaensis
Sashida, 1983
Figures 5-15, 16; 6-1,3-6

Parentactinia nakatsugawaensis Sashida, 1983, p.
172-173, pI. 37, figs. 1-9.

688

Ka/suo Sash ida

912.

Early Triassic Radiolarians from Kanto

Remarks. - This species shows rather wide
variation in size of incomplete shell. Some
specimens (e.g. Figure 5-15) have a very small
shell. The intersection of basal spine andshell is quite near to the junction of spines.
This form rarely occurs, less than 15%, butother specimens (e.g. Figures 6- I, 4) have alarge shell, whose intersection with the basal
spines is almost at the middle of basal spines.
Dimensions.-Based on 25 specimens (in,urn): length of basal spines 160-220 (average200); length of apical spines 30-120 (average100); diameter of shell I 10- I 70 (average130); width of basal hemisphere 220-300(average 245).
Occurrence. - Common in 00-1 and 002.
Types. - Illustrated specimens, Figure 5- I 5,
00-1, lOUT -KS3701 ; Figure 5-16, 00-1,
IOUT-KS3640; Figure 6-1, 00-2, 10UTKS3608; Figure 6-3, 00-2, lOUT -KS4242;
Figure 6-4, 00-1, IOUT-KS3601; Figure 65,00-1, IOUT-KS3690; Figure 6-6, 00-1,
lOUT -KS3677.
Parentactinia virgata Sashida, n. sp.
Figure 6-7

Diagnosis. - Skeleton consists of shortmedian bar and eight stout spines, of whichfour rather long. Basal spines bear stout andinwardly directed branches. These branches
trifurcate or much more and in some cases
fused each other to make incomplete cubic
shell.
Description. - Four apical spines conical,
obliquely directed, one of them (PA spine)rather long, rod-like and tapers. Other threeapical spines short. The length of the lon-689

gest apical spine is one third to half that ofbasal spine. Four basal spines cylindrical,
stout and straight. At the end or near theend of basal spine, one or two, rarely three
stout and rod-like branches present. The
thickest branch directed inward. Thisbranch bears verticils of two to five, commonly four spinules near the end of branches.
Spinules are straight, anastomosed with theother spinules to form a very loose reticulate
shell.
Remarks. - This new species is characterized by its very loosely reticulated cubic incomplete shell bordered by four stout
branches. The diagnosis of the genus Parentactinia Dumitrica is having a single globular incomplete shell. The present new
species does not bear any globular shell but
the basic disposition of both apical and basal
spines is quite identical to that of Parentactinia. Therefore I tentatively included
this species under the genus Parentactinia.
At first sight, this species has a close similarity
to Tandarnia arachnoconcha Dumitrica,
however, the latter species has a different
disposition of spines.
Dimensions.-Based on four specimens (in,urn): length of PA spines 40 to 70 (average50); length of short apical spine 15 to 30(average 20); diameter of incomplete shell
89-130 (average 105); width of basal hemisphere 120-150 (average 135).
Etymology. - Latin, virgatus, made of twigs.
Occurrence.-Rare in 00-1 and 00-2.
Type.-Holotype, Figure 6-7, 00-1,
lOUT -KS3699.

Genus Pactarentinia Furutani, 1983
Remarks. - This genus differs from Par-~
FigureS. 1-3, Archaeosemantis sp., 1: OG-I, IGUT KS-4218; 2: OG-2, IGUT-KS4219; 3:
OG-I, IGUT-KS4229. 4-8, Archaeosemantis venusta Sashida, 4: OG-2, IGUT-KS4231; 5: OG-I,
IGUT-KS4216; 6: OG-2, IGUT-KS4239; 7: OG-2, IGUT-KS4241 ; 8: OG-I, IGUT-KS4240; 9, Archaeosemantis aff. venusta Sashida, OG-I, IGUT - KS3615. 10-14, A rchaeothamnulus okuchichibuensis
Sashida, n. sp., 10: Paratype, OG- 2, IGUT - KS3623; 11: Paratype, OG-2, IGUT - KS8993; 12: Paratype,
OG-I, IGUT-KS8994; 13: Holotype, OG-I, IGUT-KS3773; 14: Paratype, OG-I, IGUT-KS3622. 15,
16, Parentactinia nakatsugawaensis Sashida, 15: OG-2, IGUT -KS4242; 16: OG-I, IGUT -KS3640.
Scale bars, A and B equal to 100 Jim; A applies to 10-16 and B to 1 to 9.

690

Katsuo Sash ida

912.

Early Triassic Radiolarians from Kanto

entactinia in having a complete sphericalshell, which cover a part of basal spines.
The present Triassic Pactarentinia slightly
differs from that of the Devonian. Namely,
the former has a spongy spherical shell but
lacks tent-like lamellae at the junction of
apical and basal spines.
Pactarentinia koikei Sashida, n. sp.
Figures 6-2,8-12

Diagnosis. - Shell consists of rather large
spongy spherical shell and rod-like long basal
and apical spines.
Description. - Shell comprises eight spines
and a bar-centered spicule with a single spherical spongy shell. The spicule consists of ashort median bar and four downward rodlike basal spines and four upward rod-likeapical spines of which PA spine is predominant. The longest apical spine (P A spine)
attains more than ISO ,urn. The basal spines
are also long and each of them forms an angleof approximately 90 degrees with its neighbors. The basal spines gently taper, rarely
they fuse directly to the spongy shell to forma rather large spongy fabric. The spongyshell is almost spherical and sometimes slightly compressed in the apical and basal directions, and its internal surface is usuallysmooth.
Remarks. - The present new species is quite
similar to Pactarentinia holdsworthi Furutani
in general shell shape. However, P. holdsworthi has a small latticed and tent-like shellformed by lamellae connecting with the proximal portions of basal spines and median bar.
The basal spine of P. holdsworthi has small
needle-like spines in latticed shell.

691

Dimensions.-Based on 20 specimens (in,u m): diameter of spongy shell 120-ISO (average 130); length of basal spines 130-170(average 13S); length of short apical spines
100-120 (average 110); length of PA spine
12S-ISO (average 13S); width of basal hemisphere 370-430 (average 410).
Etymology. - The species name koikei isnamed for Professor Toshio Koike of Yokohama National University in honor of his
contribution to the study of Triassic conodonts.
Occurrence.-Common in OG-I and OG2.
Types. - Holotype, Figure 6-8, OG-2,
IGUT -KS3893; Paratypes, Figure 6-9, OG2, IGUT-KS3894; Figure 6-10, OG-I,
IGUT -KS3707; Figure 6-11,12, OG-2,
IGUT -KS3706; Figure 6-2. OG-l.
IGUT -KS3709.
Superfamily Shaerollacea Haeckel, 1881
emend. Pessagno, 1977
Family Pantanellidae Pessagno, 1977
Subfamily Pantanellinae Pessagno, 1977
Genus Pantanellium Pessagno, 1977

Pantanellium? virgeum Sashida, n. sp.
Figures 7-9 -14

Diagnosis. - Shell comprises first medullaryand spherical cortical ones with bipolar rodlike primary spines.
Description. - Cortical shell spherical withlarge hexagonal and rarely pentagonal poreframes. All pore frames have rather highnodes at bars between pore frame vertices.
Bars of pore frames rather thin along Y and
two to three times thicker along Z. Six pore~
Figure 6. 1, 3-6, Parentactinia nakatsugawaensis Sashida, 1: OG-2, IGUT -KS3608; 3: OG-2,
IGUT-KS4242; 4: OG-I, IGUT-KS3601 ; 5: OG-I, IGUT-KS3690; 6: OG-I, IGUT-KS3677; 2, 812, Pactarentinia koikei Sash ida, n. sp., 2: OG-I, IGUT-KS3709; 8: Holotype, OG-2, IGUT-KS3893;
9: Paratype, OG-2, IGUT -KS3894; 10: Paratype, OG-I, IGUT -KS3707; 11-12: a stereo pair showinginternal feature, OG- 2, IGUT - KS3706. 7, Parentactinia virgata Sash ida, n. sp., Holotype, OG-I, IGUTKS3699; 13,16, Pou/pus sp., 13: OG-I, IGUT-KS3654; 16: OG-2, IGUT -KS3655. 14,15, Saitoum sp.,
14: OG-2, IGUT - KS3665; 15: OG-I, IGUT - KS3604. Scale bars, A and B equal to 100,urn; A applies
to 1, 2, 5, 8-10, 15, and B to 3, 4, 6, 7, 11-14, 16.

692

Katsuo Sash ida

912.

Early Triassic Radiolarians from Kanto

frames visible along both AN and BB'.
Shorter polar spines about three-fourths ofthe length of longer one. Almost all portions of spines circular in the axial section,
but usually the proximal one-fourth to twofifths portion of shorter spine tends to become
triradiate. Grooves become wider proximally. The first medullary shell seems to havehexagonal and pentagonal fragile pore frames
and is connected with two rod-like polar
spines and several thin secondary radial
beams.
Remarks. -One of the diagnostic characters of the genus Pantanellium Pessagno is
having triradiate polar spines. Although thepresent new species has a triradiate portion
only at the proximal part of shorter polar
spines, the longer spine and the middle anddistal parts of shorter spine lack longitudinalridges and grooves. These characters may
indicate that of the ancestor of typical
Pantanellium. However, the shell morphology of the present new species is quite identical to the criteria of the genus Pantanellium.
I tentatively incl uded this new species in thegenus Pantanellium.
The present new species is slightly similar
to Pantanellium browni Pessagno and Blome
described from the Rhaetian (?) to HettangianBlack argillite Member of the Kunga Formation, Queen Charlotte Islands, Canada.
However, they are distinguished from eachother by the above described character ofpolar spines.
Dimensions. - System of dimension described below is based on that of Pessagno
and Blome's description (1980, p. 241, text-fig.
5). Based on 14 specimens (in .urn): AN
82- 108 (average 94); N S' 92-115 (average693100); AS 108-135 (average 118); BB' 93-110(average 105); CC' 41-68 (average 48); dd'41-58 (average 49).
Etymology. - Latin, virgeus means rods.
Occurrence. - Common in OG-I and OG2.
Types.-Holotype, Figure 7-13, OG-I,
IGUT -KS4209; Paratypes, Figure 7-9, OGI, IGUT-KS3616; Figure 7-10, OG-2,
IGUT-KS4232; Figure 7-11, OG-2, IGUTKS4238; Figure 7-12, OG-2, IGUTKS4236; Figure 7-14, OG-I, IGUT-KS3603.
Superfamily Spongodiscacea Haeckel, 188 I
emend. Pessagno, 1971, 1973
Subsuperfamily Pseudoaulophacilae
Riedel, 197 I emend. Pessagno, 1971
Family Sponguriidae Haeckel, 1862
emend. Pessagno, 1978
Subfamily Archaeospongopruninae
Pessagno, 1973
Genus Protopsium Pessagno
and Poisson, 198 I

Remarks.- The diagnostic character of thisgenus is the presence of ellipsoidal spongyshell with two polar spines and patagium-likemass. The genus Protopsium has clear fourtypes in shell character. Namely, the first
one has a flattened ellipsoidal fine spongyshell with long polar spines with or without
alternating grooves and ridges (e.g. Pro topsium ehrenbergi Pessagno and Poisson).
The second is characterized by having fine toloose spongy meshwork and a conical polar
spine with deep grooves and ridges (e.g. P.
ispartaense Pessagno and Poisson, P. gesponsa De Wever). The third one has very finespongy meshwork with three to five fine coni-~
Figure 7. 1- 8, Spongostephanidium /ongispinosum Sashida, n. sp., 1: Holotype, OG-l, IGUTKS3643; 2: Paratype, OG-2, IGUT - KS3663; 3: Paratype, OG-l, IGUT - KS3664; 4: Paratype, OG-2,
IGUT-KS3653; 5: Paratype, OG-I, IGUT-KS3675; 6: OG-2, IGUT-KS3698; 7: Paratype, IGUTKS3661 ; 8: OG-2, IGUT -KS3669; 9-14, Pantanellium? virgeum Sash ida, n. sp., 9: Paratype, OG-l,
IGUT-KS3661; 10: Paratype, OG-2, IGUT-KS4232; 11: Paratype, OG-2, IGUT-KS4238; 12: Paratype, OG-2, IGUT - KS4236; 13: Holotype, OG-l, IGUT - KS4209. 15-18, Protopsium sp., 15: OG-2,
IGUT-KS3672; 16: OG-l, IGUT-KS3685; 17: OG-2, IGUT-KS3645; 18: OG-l, IGUT-KS3676.
Scale bars, A to C equal to 100,urn; A applies to 15, B to 2, 6, 9, 12, 14, 16-18 and C to 1, 3- 5,7,8, 10,
11, 13.

Katsuo Sashida694

cal polar spines (e.g. P. libidonosum De
Wever). The last one is characterized byhaving shell form like the genus Gorgansium
Pessagno and Blome, which has an ellipsoidal
to spherical shell with three polar spines (e.g.
P. posinos De Wever). However, the fourth
one has a slightly different morphology inhaving a lattice shell. The taxonomic position of this type may be removed into anothergenus.

skeletal structure of this group except for theJurassic genus Perispheridium (e.g., Pessagno
and Blome, 1982; MacLeod, 1988). I discriminated several forms belonging to thisfamily in the present study, but only one
genus Spongostephanidium was describedherein because the detailed internal skeleton
of other forms was not determined.
Genus Spongostephanidium Dumitrica, 1978

Spongostephanidium longispinosum
Protopsium sp.

Sashida, n. sp.

Figures 7-15-18

Figures 7-1-3

Description. - Primary shell small, sphericalto ellipsoidal with tetragonal pore frameswith small nodes at vertices. Secondaryspines extending from primary test into circular to ellipsoidal spongy shell. Polar spines
rod-like, long but usually not equal in length.
Remarks. - This unidentified species is easily distinguished from other species of thisgenus by having rod-like polar spines. Thisis slightly similar to Protopsium ehrenbergi
Pessagno and Poisson reported from the Liasof Giimiislti, Turkey in general shell shape.
However, the former is distinguishable fromthe latter by having long and thin rod-likepolar spines.
Dimensions.-Based on 15 specimens (in,urn): diameter of spongy shell 130-163 (average 146); length of polar spine 74-200 (average 175).
Occurrence. - Rare in OG-I and OG-2.
Types. - Illustrated specimens, Figure 7-15,
OG-2, IGUT -KS3672; Figure 7-16, OG-I,
IGUT-KS3685; Figure 7-17, OG-2, IGUTKS3645; Figure 7-18, OG-I, IGUT-KS3676.

Diagnosis. - Spongostephanidium withspherical rather coarse spongy-like cephalis
and three rod-like horns.
Description. - Cephalis spherical withcoarse spongy mesh. Arches of basal skeleton included within the inner parts of spongy
cephal is. Three horns rod-like with pointed
ends. They are usually unequal and shorter
than the diameter of cephalis and situated in
the same plane. The longest horn rarelyattains two times in length that of shorter one
in a specimen. One of the horns L is commonly shorter and making with the apicalhorn an angle smaller than with the otherhorn L. Sometimes wide and short threegrooves present near the base of horn. Internal surface of spongy shell is rather smooth.
Remarks. - The internal skeleton of thisspecies is not fully observed and it could not
be determined whether this species bears adorsal spine or not. I tentatively included
this species in the genus Spongostephanidium.
S. longispinosum slightly resembles S. spongiosum Dumitrica.
Dimensions.-Based on 20 specimens (in,u m): diameter of cephalis 110-140 (average125), length of longest horn 110-160 (average130), length of shorter spines 45-80 (average
65).
Etymology. - Latin, longi means long,spinose spine.
Occurrence. - Common in OG-I and OG-

Suborder Nassellaria Ehrenberg, 1875
Family Eptingiidae Dumitrica, 1978

Remarks.-Since Dumitrica (1978) set up
the family Eptingiidae, many radiolarians areincluded in this family. However, most of
the authors did not illustrate any internal

912.

Early Triassic Radiolarians from Kanto

2.

Types.-Holotype, Figure 7-1, OG-l,
IGUT -KS3643; Paratypes Figure 7-2, OG2, IGUT-KS3663; Figure 7-3, OG-l,
IGUT -KS3664; Figure 7-4, OG-2, IGUTKS3653; Figure 7-5, OG-l, IGUT-KS3675;
Figure 7-6, OG-2, IGUT -KS3698; Figure 77, OG-l, IGUT-KS3661; Figure 7-8, OG-2,
IGUT -KS3669.
Family Acanthodesmiidae Haeckel, 1862
Subfamily Poulpinae De Wever, 1981

Remarks.-De Wever (1981) set up the
subfamily Poulpinae under the family
Pylentonemidae Deflandre. Recently, Takemura (1985) investigated the internal
skeletal structure of Jurassic N assellaria, andhe defined Poulpinae De Wever as the subfamily of the family Acanthodesmiidae
Haeckel, because of the existence of a sagittalring.
Genus Poulpus De Wever, 1979
Poulpus sp.
Figures 6-13, 16

Remarks. - I obtained three specimens referable to this unidentified species. Poulpus
sp. slightly resembles P. oculatus De Weverdescribed from the Lias of Turkey and Greek
in general shell shape. The latter, however,has triradiate tripods and an apical spine.
Giving the specific name is rendered until a
number of well-preserved specimens are accumulated.
Occurrence. - Rare in OG-l and OG-2
Types. - Illustrated specimens, Figure 6-13,
OG-l, IGUT-KS3654; Figure 6-16, OG-2,
IGUT -KS3655.
Genus Saitoum Pessagno, 1977
Saitoum sp.
Figures 6-14, 15

Remarks. - Ten ill-preserved specimens are

695

obtained. Owing to silica fillings in the
cephalis, the disposition of morphologicalelements was not determined. However, ageneral view of these specimens is assigned to
the genus Saitoum Pessagno. This unidentified species is slightly similar to S. curuipedatum De Wever and S. keki De Weverin general shell shape.
Occurrence.-Rare in OG-l and OG-2.
Types.-Illustrated specimens, Figure 6-14,
OG-2, IGUT-KS3665; Figure 6-15, OG-l,
IGUT -KS3604.

References cited
Blome, CD., 1984: Upper Triassic Radiolaria andradiolarian zonation from Western North America. Bull. A mer. Paleontol., vol. 85, no. 318, p.
1-88, pis. 1-17.
De Wever, P., 1981: Une nouvelle sous-famille, lesPoulpinae, et quatre nouvelles especes deSaitoum, Radiolaires mesozoiques tethysiens.
Geobios, no. 24, p. 5-15, pI. I.
Dumitrica, P., 1978a: Family Eptingiidae n. fam.,extinct Nassellaria (Radiolaria) with sagittal
ring. D.S. Inst. Geol. Geoftz., vol. 64 (19761977), Part 3, Palaeo nt., p. 27-38, pis. 1-4.
- - , 1978b: Triassic Palaeoscenidiidae andEntactiniidae from the Vicentinian Alps (Italy)
and the Eastern Carpathians (Romania). Ibid.,
p. 39-54, pis. 1-5.
- - , 1982: Middle Triassic spicular Radiolaria.
Revista Espanola Micropal., vol. 14, p.401-428,
pis. 1-7.
Fujimoto, H., Kawata, K., Miyazawa, S., Morikawa,
R., Arai, F., Takano, T., Yoshida, S., Hara, K.,
Tazuke, H. and Mado, H., 1950: Geologicalstudy of Okuchichibu. Bull. Chichibu Mus.,
Nat. Hist., no. 1, p. 1-28. (in Japanese withEnglish abstract)- - , Watanabe, K., Akagi, S., Iijima, H., Kaneko,
S., Takano, J. and Matsuzaki, S., 1957: Geologyof the north-western part of the Kanto Mountainland. Ibid., no. 7, p. 17-28. (in Japanese withEnglish abstract)
Furutani, H., 1983: Middle Palaeozoic Palaeoscenidiidae (Radiolaria) from Mt. Yokokura,
Shikoku, Japan. Part 1. Trans. Proc. Palaeont.
Soc. Japan, N.S., no. 130, p. 96-116, pis. 21-27.
- - , 1990: Middle Paleozoic radiolarians from Fukuji area. Gifu Prefecture, central Japan. Jour.
Earth Sci., Nagoya Univ., vol. 37, p. I-56, pis. 114.

Katsuo Sash ida

696

Goodbody, Q.H., 1986: Wenlock Palaeoscenidiidae
and Entactiniidae (Radiolaria) from the Cape
Phillip Formation of the Canadian Arctic Archipelago. Micropaleontology, vol. 32, no. 2, p.
129-157, pis. 1-7.
Ishii, A., 1962: Upper Paleozoic of the N akatsugawa
area, Okuchichibu, Kanto-massif. Bull. Chichibu Mus.. Nat. Hist., no. 20, p. 178-195. (inJapanese with English abstract)- - and Matsukawa, M., 1980: Ogamata Formation, Kanto Mountains, central Japan. Bull.
Tokyo Gakugei Univ.• Ser. 4, vol. 32, p. 195-207.
(in Japanese with English abstract)
Kishida, Y. and Hisada, K., 1985: Late Triassic to
Early Jurassic radiolarian assemblages from the
Ueno-mura area, Kanto Mountains, central
Japan. Mem. Osaka Kyoiku Univ .• Ser. 3, vol.
34, no. 2, p. 103-129, pis. 1-4.
Kozur, H. and Mostler, H., 1981 : Beitrage zur Erforschung der mesozoischen Radiolarien. Teil IV:
Thalassosphaeracea Haeckel, 1862, Hexastylacea
Haeckel, 1882 emend, Petrusevskaja, 1979, Sponguracea Haeckel, 1862 emend. und weiter triassische Lithocyc1iacea, Trematodiscacea, Actinommacea und Nassellaria. Geol. Paliiont. Mitt.
Insbruck, vol. II, p. 1-208, pis. 1-69.
MacLeod, N., 1988: Lower and Middle Jurassic
Perispyridium (Radiolaria) from the Snowshoe
Formation, east-central Oregon .. Micropaleontology, vol. 34, no. 4, p. 289-315, pis. 1-5.
Nakaseko, K. and Nishimura, A., 1979: Upper Triassic Radiolaria from southwest Japan. Sci.. Rep .•Coli. Educ. Osaka Univ., vol. 29, no. 2, p.61109, pis. I-II.
Pessagno, E.A. and Blome, C.D., 1980: Upper Triassic and Jurassic Pantanellinae from California,
Oregon and British Columbia. Micropaleontology, vol. 26, no. 3, p. 225-273, pis. I-II.
- - and - - , 1982: Bizarre Nassellariina (Radiolaria) from the Middle and Upper Jurassic ofNorth America. Ibid., vol. 28, no. 3, p. 289-318,
pis. 1-8.
- - and Poisson, A., 1981: Lower Jurassic Radiolaria from the Giimlislli allochton of southwest
Turkey (Taurides Occidentales). Min. Res.
Explor. Inst. Turkey. Bull., 92, p.47-69, pis. 115.
Sash ida, K., 1983: Lower Triassic Radiolaria fromthe Kanto Mountains, central Japan Part I :Palaeoscenidiidae. Trans. Proc. Palaeont. Soc.
Japan. N.S., no. 131, p. 168-176, pis. 36-37.
- - , Igo, Hh., Igo, Hy., Takizawa, S., Hisada, K.,
Shibata, T., Tsukada, K. and Nishimura, H.,
1982: On the Jurassic Radiolaria from the Kanto
Mountains. News Osaka Micropal.. Special
Vol., 5, p. 51-66, pis. 1-2.
Sato, T., Takizawa, S., Kuwahara, T., Sashida, K.,
Hisada, K., Aono, H., Fuseya, M. and Arakawa,Y., 1981: Exotic nappes of probably sliding
origin in the South Belt of the Kanto Mountains.
Ann. Rep.• Inst. Geosci.• Univ. Tsukuba, no. 7, p.
56-59.
Takemura, A., 1986: Classification of Jurassic Nassellarians (Radiolaria). Palaeontgr. A, vol. 195,
p. 29-74, pis. 1-12.
Wakamatsu, H., Sugiyama, K. and Furutani, H.,
1990: Silurian and Devonian radiolarians fromthe Kurosegawa Tectonic Zone, southwest Japan.
Jour. Earth Sci.. Nagoya Univ., vol. 37, p. 157192, pis. 1-13.
Yao, A., 1982: Middle Triassic to Early Jurassic
radiolarians from the Inuyama area, central
Japan. Jour. Geosci. Osaka City Univ., vol. 25,
p. 53- 70, pis. 1-4.
Yoshida, H., 1986: Upper Triassic to Lower Jurassic
radiolarian biostratigraphy in Kagamigahara
City, Gifu Prefecture, central Japan. Jour.
Earth Sci.• Nagoya Univ., vol. 34, p. 1-21, pis. 117.

Kanto ~*. Nakatsugawa cp$JII. Okuchichibu !illI:~)(. Ogamata *-}i'"7!!, Oyamazawa*LlJ1.R. Saitama JiIt.:E.
~*LlJ~*ff'"7!!~£~MM~.~~ft~~E. ~~2:~*LlJ~!illI:~)(K~~~~*
it-<!! ~tJ, Ga*:ffl~jTtJ:MM~.*c~ft~1~EtJ;~~~.:.£~ ~o .:. ~~ft;E.1~Etl¥~ 5~. cp$JIIVlEj$H.:.~~~ ~ =J- l' - ~ tJ, Gt!} Gnt..:tj-:lfdt,j§t": vttJ, G tJ: ~ Palaeoscenidiidae
M~-'iffilt~"e~.:.~~~.:.J: IJ~J!OC·*15~n-n'~o *liIf~lt~t..:I.:.*LlJ1.R~=J-1'- ~~~*,H) 110
Spumellaria t. Nassellaria ~.:. ":>l '-c~W L. 5 ~f]H:'~~IJ LgcJ!OC L t..:o Spumel-*-,laria -e Itl;;:!n~~ ~ ~ t ":> Palaeoscenidiidae fit, Pantanellidae M. Sponguriidae M tJ;~ i
Nassellaria -Cit Spumellaria fj!H~1rn~ t ":> Eptingiidae M t. Acanthodesniidae MtJ;~~IJ ~ nt..:o .:. ~~ft;E.1~Etl¥~7J';~i:j::~lt, ':liiff~ ~ ::z / f:/ ~ tJ, G MM~.*c~ late
Spathian t. ~ *- Gn~otl'iffiWJ~n.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 161, pp. 697-713, 12 Figs., April 30, 1991

913.

DISCRIMINATION OF TWO NEW ANADARA SPECIESUSING SHELL GROWTH PARAMETERS AND
FOURIER DESCRIPTORSOSAMU SASAKI
Institute of Geology and Paleontology,
Tohoku University, Sendai, 980

Abstract. Two new Anadara species, Anadara (Scapharca) omaruensis, n. sp. and A.
(S.) takanabensis, n. sp., are discriminated statistically from allied species using new
methods. The ontogenetic changes in shell characters are described to interpret theirdifferences in form within the framework of the spiral tube model. It becomes clear thatthese species differ in the spiral mode around the coiling axis. The commissure shapes ofthese species are compared in the Fourier Descriptors space. The variants of these species
are distributed in the one-dimensional region of the space and they can be discriminated byusing the clustering method. It is therefore possible to discriminate these allied species
based on the degree to which the commissure shape is inequilateral.
Key words.
tube model.

Anadara. clustering method, Fourier Descriptors, Fourier Distance, spiralcurves, but there is theoretical and experimental evidence that the Fourier Descriptors are
one of the most useful techniques (Ehrlichand Weinberg, 1970; Zahn and Roskies,
1972; Granlund, 1972). Persoon and Fu(1977) used the Euclidean metric in the spaceof the Fourier Descriptors to compute thedistance between different closed curve
shapes. One advantage of the application ofthis method to the problem facing morphological taxonomists is that it puts eachform in the Euclidean space that makes possible statistical analysis of the distribution ofmorphological variation involved in a taxonomic unit.
The purpose of this work is the applicationof new methods to discriminate two new
Anadara species collected from the Koyu
Formation in Miyazaki Prefecture. The new
species are allied to Anadara (Hataiarca)
castellata (Yokoyama) and Anadara (Scapharca) ommaensis (Otuka), but differ from

Introduction
Various spiral tube models have been
proposed for the description of the geometryof the molluscan shell, which is a product ofaccretionary growth (Lison, 1949; Owen,
1953; Rudwick, 1959; Stasek, 1963; Raupand Michelson, 1965). Raup (1966) estimated the morphological effects of change inthree parameters of the spiral tube model.
Sasaki (1990) applied the knowledge of the
spiral model to the classification of the gen us
Anadara.
There have been a few successful methods
for the description of the generating curve
shape, though several sets of parameters havebeen proposed. The description of the shapeof objects in a scene is one of the important
problems in picture processing. There aremany techniques available to describe closed'Received April 12, 1990; revised manuscript accepted December 20, 1990
697

Osamu Sasaki

698

these species. The differences and similarities of the shell morphologies of these allied
species will be interpreted in the frameworkof the spiral tube model.

•

Anadaro tatunokutiensis•

Anadara amicula elongata

Geological setting and occurrence
Four populations of Anadara species studied here were collected from 4 localities in 3
areas; Iwasebashi and Toriyama in the Takanabe area in Miyazaki Prefecture, Dainichiin the Kakegawa area in Shizuoka Prefecture,
and Omma at the Kanazawa area in Ishikawa
Prefecture (Figure 1) .
(1) Takanabe area in Miyazaki Prefecture
The Miyazaki Group is divided into the
Saito and Koyu Formations, which are distributed around Takanabe Town in Miyazaki
Prefecture (Suzuki, 1987: Figure 2). The
geological age of the upper part of the Koyu
Formation is 3.4-2.5 Ma, based on microfossil data (Suzuki, 1987). The two new Anadara species were collected from the Koyu
Formation at 5 localities around Takanabe
Town, where they were associated with suchcharacteristic species of the Kakegawa Fauna
as Megacardita panda and Suchium suchiense suchiense. Several specimens of Anadara(Scapharca) omaruensis, n. sp. were collected
at each of the localities at Iwasebashi, Namikake and Kurotai, and A. (S.) takanabensis,
n. sp. at Toriyama and Nihonmatsu (Figure3).

o Anadara castellata and its allied sp,cAnadara ommaensis and its allied sp.

Figure 1. Index map of localities of Anadara specimens.

TAKANABE

KAKEGAWA(Suzuki, 1987). fTsuchi andl1 Ibaraki,1981)

Ma

KANAZAWA(Tsuchi and
Ibaraki,1988)

ftI

~

0

u:

V!

~c:::1il

l1
ftI

2

~

u:

ftI

~

::l

>-

..lI:

l1lo::

lo::

·0

4

..lI:
ftI
•
NUo

~o
.-::E.;
....
V!

6

ftI

11
..c:
III ftI
ftI

E

.eftl
G/>111·ftI ..

~ 0

-I-

l1

.
ftI

ftI

Cl
ftI

V!

J

tl

:§

.- ]l.,
..c:

.~

tl

oS

~~
oE i;.!!l.,

CJ

.f: E
ftI tl
O"t;j

tl

s::
-.::

tl

. Gqa. asanoi
Toptl.. Grt. tosaensis
Base

s::

-'"E
;:j tl

.~

E E

tl 0

E
E
tl tl

"t;j"t;jtltl

s::s::-.::-.::
Figure 2. Stratigraphic occurrences of Anadara specimens.
indicate shell-bearing horizons.

Black arrows

913.

Discrimination of two new Anadara species

At Iwasebashi, Namikake and Kurotai,
mudstone interbedded with thin, fine-grainedsandstone is well exposed, and it is correlatedwith the middle part of the Koyu Formation.
Molluscan fossils are scattered in the finegrained sandstone. A nadara (Scapharca)
omaruensis is a dominant species in themolluscan assemblages at these localities, andspecimens occur in a range of sizes from small
to large. Preservation of the shells is fairly
good.
At Toriyama and Nihonmatsu, alternationof siltstone and fine-grained sandstone is
exposed and correlated with the upper part of
the Koyu Formation. Molluscan fossils arefound in closely packed shell beds in thefine-grained sandstone. Anadara (Scapharca) takanabensis is dominant in these molluscan assemblages. Preservation of these699

fossils is fairly good.
(2) Kakegawa area in Shizuoka Prefecture
The Dainichi Sandstone of the Kakegawa

a

TL

J

,

b

\l

,

beak position

L

TL

.J

o Hagenoshita
6 Sakamoto@) Kurotaio Namikake@ IwasebashiG) Toriyama

6 Nihonmatsu

Figure 3. Index map of localities of Anadara specimens at Takanabe area in Miyazaki
Prefecture.

Figure 4. Linear dimensions measured forthe analysis of ontogenetic change in shell form.
TL: length of the toothed portion of the hingeline. AM, PM: heights of the anterior and posterior margins of the commissure. CH: height ofthe cardinal area. EV: length of the divergenttoothed portion of the posterior teeth segment.
PT, CT, AT: lengths of the posterior, central andanterior segmental teeth series.

700

Osamu Sasaki

Group at Dainichi was correlated with theHorinouchi Formation (3.2-2.0 Ma) (Tsuchi
and Ibaraki, 1988). Molluscan fossils arefound as closely packed shell beds in sandstone at Dainichi. Anadara castellata is adominant species of Dainichi assemblages.
Shells are well preserved, but very fragile.
(3) Kanazawa area in Ishikawa Prefecture
The Omma Formation is composed mainlyof fine- to medium-grained sandstone around
Kanazawa City. The geological age of thelower part of this formation is 1.17-1.02 Ma,
based on microfossils and on fission track
dating (Tsuchi and Ibaraki ,1988). Molluscan fossils form shell beds in fine-grainedsandstone, and there are more than 25 shell
beds in the Omma section along the Sai River.
Three Anadara species zones were proposedby Ogasawara (1977) in the Omma Formation. These are the Anadara (Anadara)
amicula elongata Zone, the Anadara (Scapharca) ommaensis Zone, and the Anadara(Hataiarca) pseudosubcrenata Zone, in
ascending order. Specimens of A. (S.) ommaensis were collected from the middle partof the Omma Formation. Preservation of
the Anadara shells is fairly good.

Statistical description of shell growth
The shape of the Anadara shell has traditionally been described with respect to thesize of shell, shape of cardinal area, degree ofshell inflation, positIOn of beak, ratio
between height and length of shell, degree of
curvature of posterior, anterior and ventral
margins of commissure, and arrangement ofhinge teeth (Reinhart, 1943).
Nine intervals shown in Figure 4 will bedefined here in order to represent these shellcharacters. TL is the length of toothed portion of hinge line. CH is the height of cardinal area. PM and AM are the heights of theposterior and anterior margins of commissure. PT, CT and AT are the lengths ofposterior, central and anterior segments of
toothed portion of hinge line. EV is thelength of divergent toothed portion of theposterior segment of hinge line.
Most of the shell characters mentioned
above can be represented by using the ratios
between these intervals. The ratio CH/TL
represents the shape of cardinal area anddegree of shell inflation. The ratio AT /TL
corresponds to the position of beak. The
ratio AM/PM represents the ratio between
the degrees of curvature of anterior and posterior margins of commissure. The ratio EV /PT represents the arrangement of hinge teeth.
I) Size-dependent change in the cardinal
area (CH-TL relationship)
Figure 5a shows the relationship between
TL and the general logarithm of CH of
Anadara (Scapharca) ommaensis and A. (S.)
omaruensis. Figure 5b shows the same relationship of A. (Hataiarca) castellata and A.
(S.) takanabensis. A symbol in the diagram
represents the values of one specimen, and thebroken lines represent the approximate rangeof linear distribution of the symbols. The
plotting of the relationship produces a linear
distribution in the diagram if the height (CH)increases against the length (TL) of the cardinal area in a logarithmic mode.
The approximate lines of Anadara (Scapharca) ommaensis and A. (S.) omaruensis
are at a higher angle to the TL axis than thatof A. (H.) castellata and A. (S.) takanabensis. The cardinal areas of the high-angle
species are taller than those of the low-angle
species throughout ontogeny. For example,
A. (S.) omaruensis, a high-angle species, is
CH=2.2mm at TL=20mm, while A. (s.)
takanabensis, a low-angle species, is CH =0.5- 1.5 mm at the same TL.
In Figure 5, the range of distribution of
A nadara (Scapharca) omaruensis overlapsthat of A. (S.) ommaensis, and the range of A.
(H.) castellata partly overlaps that of A. (S.)
takanabensis. A degree of overlap between
the ranges of these species in the diagrams isthought to be due to the similarity of these
species in the shape of their cardinal areas.

913.

::c

~;~~~

+
0.8

1

. . -~a:~ +0:;;: 0.5"- 0.4
:>

~~@

~

-c

701

a

a

E
5

Discrimination of two new Anadara species

.....

...

-1

0.2

8

0

-2

20

0

40

0

00

20

b

E
5

::c

AlfJltifo\@~o; i... ~ -::-L'."(p6o"ZK!¥l

0

c

00

- -1

~6

40

60

0.8

0

:;;: 0.6

0"- 0.4:>oo°'ti:Do""o

~

00

b

O~-*': .P-

1

40

T l I mm I

T II mml

.....

0

0

0.2

0
0

0

-2
0

20

40

00

T II mml

020

T llmm I+ Anadam (Scapharca) ommaensis«)

A nadara (Scaphm'ca) omaruensist:.

Anadara (llataiarca) castel/alaoAnadera (Scapharca) takanabensis

Figure 5. Distribution of ontogenetic variation in the relationship between the height (CH)and length (TL) of the cardinal area.

Therefore, A. (5.) omaruensis is similar to A.
(5.) ommaensis in the shape of its cardinalarea, and also A. (5.) takanabensis is similar
to A. (H.) castellala.
2) Size-dependent change in the teeth
arrangement (EV jPT - TL relationship)
The relationship between EV jPT and TL
represents the ontogenetic change in thearrangement of hi nge teeth. N oda (1966)
classified the arrangement of Anadara hinge
teeth into 3 types: the convergent type, divergent type, and divergent type with v-shaped
teeth. At EV jPT=O, Anadara shells havethe convergent type of teeth arrangement, andat EV jPT>O, it has the divergent type withv-shaped teeth. A nadara species change
their teeth arrangement during shell growth.
It is therefore impossible to represent
Anadara species by using only one type ofteeth arrangement. The critical size can be
determined for the growth of a species, atFigure 6. Distribution of ontogenetic variation in teeth arrangement represented by change inthe value of EV /PT. A shell has the convergent
type of teeth arrangement at EV /PT = 0, and thedivergent type with v-shaped teeth at EV /PT>O.
Symbols as in Figure 5.

which its teeth arrangement is transformed
from the convergent type to divergent typewith v-shaped teeth. Therefore, the growthof teeth arrangement that is peculiar to each
Anadara species can be represented by using
the critical size (Sasaki, 1990).
Figure 6a shows the relationships between
EV jPT and TL of Anadara (Scapharca)ommaensis and A. (S.) omaruensis. A. (5.)
omaruensis changes its teeth arrangement at
TL= 13 mm, and A. (5.) ommaensis transforms it at TL = 21 mm, as shown by theshaded zones in the scatter diagram. Figure
6b shows the same relationships of A. (H.)
castellata and A. (5.) takanabensis. A. (H.)
castellata transforms its teeth arrangement at
TL=20 mm, and A. (5.) takanabensischanges at TL = 30 mm.
The difference in their critical size represents a difference in the ontogenetic change inteeth arrangement between these species, sothat both A. (5.) omaruensis and A. (S.)

Osamu Sasaki

702

takanabensis differ from both A. (s.) ommaensis and A. (H.) castellata.
3) Size-dependent change in the positionof the beak (AT /TL-TL relationship)
The ratio of AT /TL corresponds to theposition of beak on the hinge line. The
position of beak represents proximity of theshell shape to equilateral. The beak position of Anadara species migrates during
growth, and the change can be represented by
the relationship between AT /TL and TL. It
is therefore concluded that the extent towhich the shell is equilateral can be describedby using the relationship of AT /TL to TL.
Figure 7a shows the relationship between
AT /TL and TL of both A. (s.) ommaensis
and A. (s.) omaruensis. Figure 7b shows therelationships between AT /TL and TL of
both A. (H.) castellata and A. (S.) takanabensis. The relationships of these species
are similar to each other throughout
ontogeny.
a4) Size-dependent change in the shape ofthe commissure (AM/PM - TL relationship)
AM/PM is the ratio between the heights of
the anterior and posterior margins of thecommissure, which partly represents the shapeof the commissure. Figure 8 shows the relationship between AM/PM and TL of the four
populations studied here. The ranges of
AM/PM of these species are smaller than
AM/PM = 1, so that the commissure shapes ofthese species are somewhat expanded towards
the posterior margin. The difference in theratios of these species therefore will express
the difference between the degree of expansion of the posterior and anterior parts. At
the same value of TL, the AM/PM values ofA. (S.) ommaensis and A. (s.) omaruensis are
greater than those of A. (H.) castellata and A.
(s.) takanabensis. The large ratio species
are A. (s.) ommaensis and A. (S.) omaruensis, which are somewhat equilateral in theheight of the commissure. The small ratio
species are A. (H.) castellata and A. (S.)
takanabensis, which are more expanded posteriorly than anteriorly.

0.8

Shape discrimination using
Fourien Distance

~ 0.8

"-

I - 0.4

<

0.2

o

o

40

20

eo

T L I mm,

In the light of the mathematical models, a
difference in the shape of the commissure willproduce an important difference in the shell
shape. However, it has been difficult to

b

0.'

0.8

~ 0.8

~ 0.8

"

<

"

~ 0.4

I - 0.4

<

0.2

0.2

o

0

o

40

20

eo

T L I mm,Figure 7. Ontogenetic change in the ratio of
the length of the anterior teeth segment (AT) to thelength of the toothed portion of the hinge line(TL); the ratio represents the migration of thebeaks. Symbols as in Figure S.

0

40

20

eo

T L I mm,Figure 8. Ontogenetic change in the ratio
between the heights of the anterior (AM) andposterior (PM) margins of the commissure. Symbols as in Figure S.

913.

Discrimination of two new Anadara species

measure the degree of difference between
shapes of commissures, and also to analyse
statistically the distribution of different
shapes in a population. In this paper, theshape of the commissure will be described byusing the Fourier Descriptors (Granlund,1972), and the difference between the shapes
will be measured by using the Fourier Distance in the Fourier Descriptors space (Persoon and Fu, 1977).
The Fourier Descriptors (FDs) given in
Granlund (1972) are defined as follows: weassume r is a clockwise-oriented simple
closed curve of the commissure with parametric representation [xU), y(l) ] = z(l),where I is the arc length along r. A point
moving along the boundary generates thecomplex function u(l) = x(l) + iy(/) which isperiodic with period L. The FDs becomenow

If we denote by {an} the FDs of a curve of
the commissure a and by {b n } the FDs of acurve of the commissure 13 and M harmonics
are used, then the distanced (a, 13) given in
Persoon and Fu (1977) are defined as follows:

dCa, f3)=[n~-t.n*o la n- bnI2}/2
Suppose a and 13 are the commissure of
specimens so we will scale (s), rotate (cP), andadjust the starting point (a) such that the
distance is minimized. We have then to
determine $', cP, a such thatis minimized.
The defined distance is the Euclidean metric in the space of the Fourier Descriptors{an}. The commissure shape of each specimen will be transformed by the Fourier
Function to plot in the Fourier Descriptors
space. Hence, the Fourier Distance between
the place in the space represents the degree ofdifference between the commissural shapes.

1 (La n = T Jo u(/)e- i (21C!L)nl

and

Table 1.

TOI
T02
T03
T04
T05

703

Exmample of the Fourier Distance between Anadara specimens analyzed here. TOI is a
specimen of Anadara (Scapharca) takanabensis, n. sp. from Toriyama, and 019 is a
specimen of Anadara (Scapharca) ommaensis (Otuka) from Omma.
T05

T06

T07

......... 015

T02

T03

T04

14.69

11.10
13.85

9.06 9.22 11.26 8.53 .........
15.00 10.16 8.80 8.36 .........
16.05 7.82 7.24 7.58 .........
16.58 17.38 13.43 .........
5.18 6.15 .........

19.92
11.59
16.92
26.49

016

017

30.95

16.00

22.64
24.31

13.72
11.60
23.80

018
21.29
15.78

019

020

14.83

20.23

11.53
13.96

13.89
17.30
27.10

10.53

14.68
29.09
15.14

22.01
9.79

13.91

39.03
25.44

T06

5.28 ......... 12.25

23.42

9.72

13.46

10.00

13.65

014
015

10.67

7.58
15.60

12.65
10.05
16.98

7.25
10.29
11.35

14.34
8.02

10.26
5.83
14.30

016
017
018
019

8.46

18.70
9.07
10.09

15.06

28.92
8.00
6.82

Osamu Sasaki

704

40
W

U

Z

-<
..... 30
(/')

Cl

a:::

20

w

GroupOa::::::::>

0

10

u..

o

(S~p""~o) 1""" 1

A=dwo
takanabensis.....

... ..

Anadara (Scapharca)

omaruensis

Anadara (Hataiarca)

A nadara (Scapharca)

castellata

ommaensis

Figure 9. Dendrogram calculated by using the complete linkage clustering method
based on the matrix of the Fourier Distance, of which an example is shown in Table 1.

The value of the Fourier Distance is 0
between specimens with the same shape ofcommissure.
Table I shows part of the matrix of theFourier Distances between the specimens
selected from the four populations, which arecalculated by using 16 harmonics. The number of specimens used here is 15 of A nadara(Scapharca) ommaensis, 7 of A. (s.) omaruensis, 6 of A . (H.) castellata, and II of A. (s.)
takanabensis, all of which are left valves ofa lmost the same size.

Figure 9 shows the dendrogram constructedby using the complete linkage clustering
method applied to the values in Table 1.
Figure 10 shows typical commissural shapes
ana lysed here. The numbers of each form in
Figure 10 correspond to the numbers in thedendrogram in Figure 9.
As shown in Figure 9, these specimens can
be classified into 4 groups based on the
Fourier Distance. The least similarity is
between the groups A and D, and the greatest
similarity is between the specimens in the913.

Discrimination of two new Anadara species

Figure 10. Examples of commissural shapes
analysed by the Fourier Descriptors. The numbers correspond to the numbers in Figure 9, showing the positions of these examples in the dendrogram.

group B.
The shape of the commissure placed in thegroup A is characterized by a narrowly
rounded anterior margin and a broad, truncated posterior margin, as is shown by the
specimen 1 in Figure 10. The shape of thecommissure placed in the group D is withbroadly rounded anterior and ventral margins, as is shown by the specimen 4 in Figure
10. The specimens placed in the groups B
and C have intermediate shapes between
those in the groups A and D, as is shown by
the specimens 2 and 3 in Figure 10.
Specimens of a species are distributed in a
restricted range of the space, so that eachgroup in the dendrogram includes only one or
two species. Specific elements of thesegroups in the dendrogram are as follows; thegroups A and B-1 are composed of Anadara(Scapharca) takanabensis and A. (H.) castellata, the groups B-2 and C are mainly composed of A. (S.) omaruensis and A. (S.)
ommaensis, and the group D is composed ofA. (S.) ommaensis.
The possibility of discrimination of eachspecies is dependent on the degree of overlapping of the ranges of distribution in the705

Fourier Descriptors space. As shown by thedendrogram in Figure 9, the range of Anadara (Hataiarca) castel/ata is completely
overlapped by that of A. (S.) takanabensis.
It is therefore impossible to discriminate these
species based on the shape of the commissure.
A. (s.) takanabensis and A. (S.) omaruensis
in the group B have very similar commissural
shape, but the range of A. (S.) takanabensis
does not overlap that of A. (S.) omaruensis,
so these species can be discriminated. It isdifficult to discriminate A. (S.) omaruensis
from A. (S.) ommaensis based on their commissural shapes because the ranges of these
species overlap in the groups Band C.

Discussion
In this paper, shell form is described withrespect to shell growth and the shape of thecommissure. Shell growth is described by
the size dependent change in the characters of
the shell: 1) size-dependent change in CH,2) size dependent change in EV /PT, 3) size
dependent change in AT /TL, and the shapeof the commissure is described by the parameters of 4) size-dependent change in AM/PMand 5) Fourier Descriptors. Differencesamong the Anadara species described byusing these parameters will be interpreted in
the framework of the spiral tube model, andthe applicability of these parameters to thediscrimination of species will be appraised.
Several spiral tube models have been
proposed in studies of molluscan shell geometry (Owen, 1953; Rudwick, 1959; Raup andMichelson, 1965; Raup,1966). Raup andMichelson (1965) proposed 4 parameters forthe description of shell form: 1) D, distancefrom the coiling axis to the generating curve,2) W, rate of expansion of the generating
curve, 3) T, rate of translation of the generating curve along the coiling axis, and 4) S,
shape of the generating curve. The first threeof these are parameters for the description ofshell coiling.
The ratios between the defined intervals

706

Osamu Sasaki

used for the description of Anadara shells
correspond to the parameters of the model ofRaup and Michelson (1965), which represents
the basic geometry of any moll uscan shells.
CHjTL and EV jPT corresponds to DjW,
representing the mode of shell coiling aroundthe coiling axis. AT jTL corresponds to T,translation of the generating curve along theaxis, and AMjPM can be compared with S,
the shape of the generating curve of thismodel (Sasaki, in 1990).
1) Shell coiling around the coiling axis
In the framework of the spiral tube model,
the CH-TL relationship directly represents
ontogenetic change in the mode of shell coiling around the coiling axis, and the EV jPT TL relationship indirectly represents the samecharacter of coiling.
As shown in Figures 5a, b, the linear distribution in the diagram of the CH - TL relationship suggests that the shell forms a logarithmic spiral around the coiling axis (Sasaki,1990). The angle which the approximateline forms to the TL axis can be interpreted as
the degree of openness of a logarithmic spiral.
Therefore, Anadara (Scapharca) ommaensis
and A. (s.) omaruensis have more open logarithmic spiral shells than A. (H.) castellata
and A. (S.) takanabensis have.
The A nadara shells change their teeth
arrangement during ontogeny. The change
in teeth arrangement is caused by the change
in the shell coiling mode during growth. At
EV jPT>O, the Anadara shell is growing in alogarithmic spiral around the coiling axis
(Sasaki, 1990.). Therefore, the critical size is
the size at which the Anadara shells change
their mode of shell coiling to a logarithmic
spiral. The logarithmic spiral model explains well the rapid increase in the inflationof a shell in old age compared with its slow

expansion of the commissure. The criticalsize appears to be a useful index of thematurity of a shell. The ontogenetic change
in teeth arrangement measured for an A nadara species always produces a restricteddistribution in the scatter diagram of therelationship of EV jPT to TL. In manycases, however, the measurement in a population of two or more species produces a wide
range of distribution in a scatter diagram(Sasaki, 1990). The EV jPT - TL relationshipof the Anadara species can be interpreted as
the characters of growth peculiar to eachspecies, hence the critical size is a useful
criterion for discrimination of these Anadara
species. Therefore, each Anadara population studied here could contain a single
species, because each population is distributed in a restricted range in the diagram. Alsothese species differ in their critical size of thechange in teeth arrangement, hence they are
different species.
2) Translation of the generating curvealong the coiling axis
The AT jTL-TL relationship represents thetranslation along the coiling axis. As shownin Figures 7a, b the relationships of these
species are very similar during growth, so thatthese species are similar in the degree of thetranslation of the generating curve along thecoiling axis.
3) Shape of the generating curve
In the Fourier Descriptors space, variationof a population will be characterized by theextent and the number of dimension of theregion where the specimens of the population
are distributed. In this analysis, specimensof these Anadara species are distributed in an
one-dimensional region in the Fourier Descriptors space, and the typical forms in~
Figure 11. la, b: Anadara (Scapharca) ommaensis (Otuk a) from the Omma Formation (x 1). 2a,b: Anadara (Scapharca) omaruensis, n. sp. from the Koyu Formation, IGPS colI. cat. No. 10325(Holotype) (X 1). 3a, b: Anadara (Hataiarca) castellata (Yokoyama) from the Kakegawa Group (Dainichi Sandstone) (X 1). 4a, b: Anadara (Scapharca) takanabensis, n. sp. from the Koyu Formation, IGPS
colI. cat. No. 101327 (Holotype) (X 1).

913.

Discrimination of two new A nadara species

707

708

Osamu Sasaki

different posItIOn in the dendrogram are
different in the degree to which their commissure shape is inequilateral. It is therefore
possible to represent most of the variation in
the commissure shape of these species byusing the ratio AM/PM.
In the AM/PM-TL diagram (Figure 8), therange of Anadara (Scapharca) ommaensis
overlaps that of A. (S.) omaruensis, and therange of A. (H.) castellata overlaps that of A.
(S.) takanabensis. This shows that A. (S.)
ommaensis is similar to A. (S.) omaruensis incommissural shape, and A. (H.) castellata issimilar to A. (S.) takanabensis. Therefore,
the ratio AM/PM is a useful parameter forthe discrimination of a new species from its
allied species.
Anadara (Scapharca) takanabensis from
Toriyama was first classified as Anadara
inflata by Yokoyama (1928). Shuto (1961)
reported it under the name Anadara castellata and Noda (1966) identified it as A. (S.)
iwashibaraensis. Yoshida (1987) placed thespecies in A. (H.) castellata. The comparison in this paper suggests that A. (S.) takanabensis is allied to A. (H.) castellata but
is a different species.
Anadara (Scapharca) iwashibaraensis was
originally described by Noda (1965) based ona specimen from the Dainichi area. According to the original description, this species ischaracterized by a narrowly rounded anterior
margin and a broadly truncated posterior
margin. Therefore the commissure shape ofA. (S.) iwashibaraensis seems to be similar
not only to that of A. (S.) takanabensis but
also to that of A. (H.) castellata, whichplotted in the group A in the dendrogram(Figures 9, 10). It is therefore impossible todiscriminate these species on the basis ofcommissural shape.
The range of rib numbers is an importantmeasurement in specific determination of
Anadara species (Reinhart, 1943). The shellsurface of Anadara (Scapharca) iwashibaraensis is sculptured with 31 radial ribs.
However, A. (S.) takanabensis ranges in rib
numbers from 36 to 39. These species aredifferent in rib numbers provided it is remembered that on some specimens it is difficult todecide which threads should be counted asribs. It is therefore possible to discriminate
A. (S.) takanabensis from A. (S.) iwashibaraensis by the number of ribs.
Conclusion

1) Anadara (Scapharca) omaruensis, n.
sp. from Iwasebashi is allied to A. (Hataiarca) castellata and A. (S.) ommaensis. It
differs, however, from allied species in themode of shell coiling around the coiling axis,
which is represented by the size-dependentchange in EV /PT. A. (S.) omaruensis is
more allied to A. (S.) ommaensis than to A.
(H.) castellata in the shape of the commissure, which is represented by the size-dependentchange in AM/PM.
2)

Anadara (Scapharca) takanabensis,

n. sp. from Toriyama is closely allied to A.
(H.) castellata in commissural shape, butdiffers from in the mode of shell coilingaround the coiling axis.
3) Anadara (Scapharca) takanabensis isallied to A. (S.) iwashibaraensis in commissural shape, but it differs from the latter in thenumber of radial ribs.
Acknowledgments
The author wishes to express deep gratitude
to Prof. Kenshiro Ogasawara of Institute of
Geoscieme, University of Tsukuba for his
continuous guidance and kind encouragement during this study. Deep appreciation isdue to Dr. Alan G. Beu of New ZealandGeological Survey for his kind suggestions
and critical reading of the manuscript.

~
Figure 12. 1, 2a-4b: Anadara (Scapharca) takanabensis, n. sp. from the Koyu Formation, IGPS
coil. cat. No. 10326, (x I). 5a-Sb: Anadara (Scapharca) omaruensis, n. sp. from the Koyu Formation,
IGPS coil. cat. No. 101328 (X I).

913.

Discrimination of two new Anadara species

-,

6a

6b

709

Osamu Sasaki

710

Systematic discription
Family Arcidae Lamarck, 1809
Subfamily Anadarinae Reinhart, 1935
Genus Anadara Gray, 1842
Subgenus Scapharca Gray, 1847
Anadara (Scapharca) omaruensis, n. sp.

minami-machi, IGPS colI. cat. No. 101325(Holotype), IGPS colI. cat. No. 101326;
Kurotai, Kawaminami-machi, N amikake,
Kawaminami-machi, Koyu Formation in the
Miyazaki Group, Pliocene.
IGPS is the abbreviation for Institute ofGeology and Paleontology, Tohoku University, Sendai.

Figures 11-2a, b; 12-5a-8b

Description. - Shell medium in size, moderately thick, slightly inequivalve and inequilateral; beaks prosogyrate, situated atanterior third of shell; shell elongate in outline, hinge margin straight, interrupted byprominent narrow umbones; ventral margin
broadly 'curved, anterior margin smoothly
rounded, posterior margin sharply curved
below, straight above without depressed area
along posterior side.
External surface sculptured with 36-39
radial ribs (commonly 37-39); surface of ribs
strongly granulated on left valve, rathersmooth on right valve; most ribs narrower
than their interspaces.
Cardinal area narrowly trigonal, withchevron-shaped grooves. Hinge somewhat
arched, rather narrow, slightly wider at
extremities than at center of shell; taxodont
teeth small and numerous; teeth diverge
ventrally near center of hinge line, and converge with v-shaped teeth at extremities inadult shells; all teeth convergent in juvenile
shells. Inner ventral margin sculptured accordance with external ribs.
Comparison. - The present species resembles Anadara (Scapharca) ommaensis(Otuka) from the Omma Formation andAnadara (Hataiarca) castellata (Yokoyama)
from the Dainichi Sandstone, but as shown
above the teeth arrangement of A. (S.) omaruensis is different from those of these allied
species at the same shell size. A. (s.)
brough ton ii (Schrenck) is allied to A. (s.)
omaruensis, but has a larger number of radialribs than this species.
Localities. - Iwasebashi, Matsubara, Kawa-

Anadara (Scapharca) takanabensis, n. sp.
Figures 11-4a, b; 12-1,4b

A nadara (Scapharca) iwashibaraensis, N oda, 1966,
pI. 2, fig. 26, pI. 12, figs. 16-17.

Description. - Shell large, inflated, Inequilateral, beaks prosogyrate, situated atanterior third to fourth of length; shell elongate, hinge margin long and straight, anterior
margin narrowly rounded, ventral margin
broadly rounded, posterior margin truncated,with depressed area along posterior side.
External sculpture similar on right and left
valves, consisting of 35-39 radial, rather flattopped ribs (commonly 36-37); interspaces
between ribs of same width as one rib, crossedby commarginal growth lines. Ligamental
area rather narrow. Hinge straight and narrow; taxodont teeth small and numerous;
inner margin crenulated in accordance withexternal ribs.
Comparison. - The present species resembles Anadara (Hataiarca) castellata (Yokoyama), but as shown above these species differ
in their ontogenetic change of the teeth
arrangement. This species is allied to A. (S.)
iwashibaraensis Noda, but the latter has only
31 radial ribs. A. (S.) taiwanica Noda resembles A. (S.) takanabensis, but its shell ismore inflated and more equilateral than thisspecies.
Localities. - Toriyama, Kawaminami-machi, Koyu-gun, Miyazaki Prefecture, IGPS
colI. cat. No. 101327 (Holotype), IGPS colI.
cat. No. 101328: Nihonmatsu, Takanabemachi, Koyu-gun, Miyazaki Prefecture, Koyu

913.
Table 2.

Discrimination of two new Anadara species

711

Measurements of Anadara (Scapharca) omaruensis, n. sp.

Specimen

L

H

C

TL

CH

EV

PT

AT

PM

AM

Ribs

L/R

101328- 1

9
10
11.5
16
20
20
25
28.5
46
15
16
15
26

8
9
10
13
16
15
20
24
34

2.5
3

5.9
7
7.5
9.3
12.5
12.1
15.8
17.8
28
8.8
10.1
9.1
15
19
28
32
32
24.8
15.3
32
7.4
9.1
8.6
8.7
11
11
10.5
12.4
12.5
14.1
15.5
16.9
36.7
35.5
32.2
35
29.5
31
20.2
23.7
22.1
18.2
25
17.4
13.2
31.5

0.2
0.3
0.3
0.5
0.7
0.5
0.4
1.5
3.3
0.5
0.6
0.4
0.8
1.6
2.7
3.7
3.9
1.8
1.1
3.5
0.4
0.5
0.4
0.5
0.5
0.5
0.5
0.8
0.6
0.6

o
o

3.8

1.6
1.7
1.9
1.9
3.7
3.4

5.5
6.5

3.3

8.5
10
10
13
16
20
8.5
8.5
8
12.5
17

4

38
37
37
37

14
4
4.5
4
8.5
11

24
27

16
18

25
6.5
7.5
6.5

18
3.5
4
4

39
36

38
38

R
L
R
L
R
R
L
L
L
L
L
L
L
L
L
L
R
R
R
R
R
L
R
L
R
R
R
L
R
L
R
L
L
R
L
R
L
R
L
L
L
L
R
R
L
R

-2
- 3
- 4

- 5
- 6
- 7
- 8
- 9
-10
-11
-12
-13
-14
-15
-16
-17
-19
-20
-21
-30
-31
-32

-33
-34
-35
-36
-37

-38
-39
-40
-41
-51
-52
-53
-54
-55
-56
-57
-58
-59
-60
-61
-62
-63
101327

13
14
15
19

3
4
6
5.5
7
8
12
4
5
3.5
7

52
54

42
40

14
15

54
12
14

41
10
11.5
11
11.5
14

15
4
4.5

13.5
14
16.5
17

13.5

20
20
23
25

16
16
18
19.5

61
58
55
55
49
50
37
40
37
34
40
29
23
51

45
56
40
42
35
36
28
30
28
27
30

4
4
5
6
6
6
6.5
7
8
16
17
16
15
14
17
10
10
10
9

22

11
8

19
40

6.5
14

1
1.2
3
3
3
3
3
3
1.9
2.1
2.1
1.8
1.8
1.2
0.9
3.5

o
o
o

o

o
5.2
11.2

o
o
o
1.9
5.5
8
9.3
12.3
5.5
1.5
15.5

o

o

o
o
o
o
o
o
o
o
o

3.1
12
12
10
10
11.3

11
5.4
6.2

5.5
4
6.3
2.7

4.4
4.8
6.5
7.8
7.8
10.8
11.8
18.7
5.9
6.6
5.8
9.8
12.3
17.3
20.4
21.2
14
10.3
21.9
5
5.7

5.5
6
7.2
7.5
6.9

8.3
8
9.5
10.5
10.9
24
23.1
20.3
23.6
19
20.5

13.5
15.5
14.5
12.3
16.2
11.3

o
9.5

All specimens from the Koyu Formation, Miyazaki Prefecture

20.2

4
5
7.5
1.9
2.5
2.4
3.9
5.4
8.9
9.1
8.5
7.3
3.9
8
1.5
2.6
2.4
1.9
2.8
2.5
2.7
3.2
3.4
3.7
4.9
12

11
10
10
8.5
8.5
6
7
6.5
4.8
7.4
4.5
3.6
8.8

3.5

6.5
6
8

7

38
38
38
38
36
37
36
39

38
37
39
39

38
38
38

38

8

4

10
8

6
5.5

10
10
12
12

6
7
7.5
7.5

27
28
25
25
21

22
22

37

19
19
17
16
13
13

37
37

13

39
37
39
39

36
37

38
37
39
39

38

22
17
19
18
16
17
13
11.5
26

11

12
9.5
8.5
17

38

38
38
39

38

38
39

Osamu Sasaki

712

Table 3.
Specimen
101326- 1
- 2
- 3
- 4
- 5
- 6
- 7
- 8
- 9
-10
-11
-12
-13
-14
-15
-16
-17
-18
-19
-20
-21
-22
-23
-24
-25
-26
-27
-28
-29
-30
-31
-32
-33
-34
- 35
-36
-37
-38
-39
-40
-41
-42
-43
-44
-45
-46
-47
-48
-49
-50
-51
-52
-53
-54
-55
-56
-57
-58
-59

-60
-61
-62
-63
101325L
8

H
6.2

Measurements of Anadara (Scapharca) takanabensis, n. sp.
C
2

9.2

12.5

16
11
18
22
19
29
16
23
27
26
30
31
33

12
13.5
14
17
15
24
12.5
18.5
22
20
23
25
26.5

4
4.5
4.5
5.5
5
8
4.5
6.5
7
8
9
8.5
9

26.5
32
30
37.5
35
38
37
45
55
49
48
50
49
50
55
53
55
54
57
62

21.5
24
23
28
26
30
28
35
40
35
35

8
8.5
8
11
10
10
10
13
14
14
12.5
13.5
13.5
13
15
15
15.5
14
16
17

40
40
40
40
42
44
41
49

55
61
60
63
63
63
63
59
68
65
63
61
70
68
70
71

74
70
74
66
66

45
45
46
46
51
45
44
45
50
50
49
52
55
52
53
54

58
57
57
53
51

16
17.5
18
18
18

22
19
17
19
21
18
19
19
22
22
19
22.5
19
20
23
21
23
21
17.5

TL
5.3
5
7.6
6.1
7.6
10.7
9.9
12.2
14.2
12.1
19.2
10.5
15.8
18.9
17.4
19.5
20.9
20.5
16.3
17.8
21.5
20
24.5
23
25.5
25.2
30
35.7
33.5
30.1
31
34.9
33.6
34.2
34.4
37
36.5
37.4
41.8
36
36.5
38
43
40.3
40.8
42
39
39
42
42.5
19.9
42.5
42.5
47
47
43.5
47.5
46.8
46.6
48.4
47
46
44
41.5

CH
0.2
0.1
0.3
0.5
0.5
0.5
0.7
0.4
0.8
0.7
1.1
0.5
0.9
0.8
0.8
1.2
1.1
1.5
0.8
1.2
1
0.9
1.4
0.8
1.5
1.1
1.8
2.4
2
1.8
2
2.1
2
2.1
2.2
2.5
2.8
3.3
3.1
3.3
2.3
3
3.3
3.5
3.1
4.8
2.5
2.7
2.2
4.7
4.1
3.8
2.7
5.3
5.3
4.3
5.4
4.8
3.4
5.1
5.2
5.3
2.8
3

EV
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6.5
3.4
0
0
3.8
3.1
7.5
1.5
5
7.4
9.5
10
8
4.5
8.2
13
15.7
15.1
17
8
10
9.9
15.1
13
15.3
12.5
21
21.3
17.3
21.3
19
20
22.5
21.5
18.5
14.5
14

PT
3.9
3.7
5.5
4.3
5.8
7.2
7.4
8.7
10.1
9
12.9
7.2
11.3
12.8
12.8
13.9
14.2
14
11
12.8
15
14
17
15.2
17.8
17.4
20.4
23.7
21.2
20.9
21
21.5
22
21.8
20.7
23
22.1
23.4
25.4
22.3
22.6
23.3
24.2
25.3
24.5
25
24.6
24.8
25.5
25.3
25.5
26.1
25.5
29.3
27
27.7
28.3
27.5
28.9
29.8
28
26.4
26.5
25.5

AT
1.4
1.3
2.1
1.8
1.8
3
2.5
3.5
4.1
3.1
5
3.3
4.5
6.1
4.6
5.6
6.7
6.5
5.3
5
6.5
6
7.5
7.8
7.7
7.8
9.6
11.3
11.6
9.2
10
11.2
11
11.8
12
13.2
13.1
12.9
14.4
12.8
13
13.3
15.2
13.4
13.1
15.7
13.3
13.3
14.8
15.4
14.2
14.5
16
16
17.3
14.7
16
15.7
15.4
16.2
17.3
18.5
15.5
14.5

All specimens from the Koyu Formation, Miyazaki Prefecture

PM
5.3

6.5
7.9
10
9.7
11.3
13
10.5
17
9
14
16
15
17
18
20
14
16
17
17
20
19
20.5
20
25
30
27
26
27.5
27
28
28
30
29
30
34

35
30
33
32
33
32
30
33
34
32
33
34
35
36
32
36

40
37
40
37
32

AM
2.5

2.5
3.4
4.3
5
5.5
5.5
4.8
7
4
6
6
6
8
10
10
8
7.5
9
7.5
11
10
11
11
15
15
15
12
13
16
17
15
18
19
16
20

20
17
19
18
21
21
18
19
20
21
22
21
22
22.5
22
23

23
23
27
23
21

Ribs
35
36
37
36
35
36
35
37
38
37
36
38
37
38
38
37
37
37
37
37
36
35
37
36
39
37
37
37
36
37
36
35
37
37
36
39
37
36
36
37
35
38
39
37
35
36
37
35
36
35
36
37
36
37
36
37
37
37
36
35
36
38
39
38

L/R
R
L
L
L

R
R
L

R
L

R
L

R
L

R
L

R
L
L
L
L
L

R
R
L
L

R
L
L

R
L

R
L
L

R
L
L
L

R
L

R
L

R
R
R
R
R
R
R
R
L
L

R
R
L

R
L

R
L
L

R
L

R
L

R

913.

Discrimination of two new Anadara species

Formation in the Miyazaki Group, Pliocene.

References
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for characterization of grain shape. Jour. Sed.
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1294-1295.
Reinhart, P.W., 1943: Mesozoic and Cenozoic Ar-

713

cidae from the Pacific Slope of North America.
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73-206, pis. 11- I3.
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37-41.
- - and - - 1988: Notes on the Omma-Manganji
Molluscan Fauna: Its geologic age andpaleoceanographic implications. Saito Ho-on

Kai Spec. Pub. (Prof T. KotakaVo!.), p.557-565.

Yokoyama, M., 1923: Tertiary Mollusca from Dainichi in Totomi. Jour., Coll. Sci., Tokyo Imp.
Univ., vol. 44, p. 1-18, pis. 1-2.
Zahn, c.T. and Roskies, R.Z., 1972: Fourier descriptors for plane closed curves. IEEE Trans.
Computers, vol. C-21, p. 269-281.

ID!:RX;:lHt~ C 7 - ~ I. gcJitt-J-?:' flh ,t-: A nadara ~ 2 ~:fiO)'Ftl5Jlj: =":)0) 7J$?:' flh' -C,
Anadara ~O) 2 ~:fi, Anadara (Scapharca) omaruensis c Anadara (Scapharca) takanabensis ?:'.:t.n-t'.n0) ~f!;J:fi iJ, ~ 'F1j5J1j L-I-:o D- c ":)0) 7J$ U.ID!: 0) RX;ffi:~1r. ?:,gcJitt L, RX;ffi:~1r.0)il§:~iJ' ~ID!:O)t!f:~-1:- T /v O)wm~O)<=P-c'ID!:*Mi~O)il§:~ ?:'§'lZfIffi"t" Q 7J$-C';,5 Q 0.:.

.n ~ O)*5!Jlt 2 ~:fi?:,"'3-tr 4 :fiftID!:O)~$ffi@] iJ -C·O)t!f:EiEO)~A f~jfj); l' iJ~;,5 Q .:. c iJ~hiJ'-0t-:o t l)D-c":)O)7J$ft, ID!:JjfJP'ifllO)M!~?:'7- ~ I.gcJitt?:'flh,-c~cJittL, 2 {l9Iwia9 0) ID!:1jfJ
p'ifllO)MfrnO)il§:~?:, 7 - ~ I.gcJitt-J-ia90)jfj);l'?:'~"t"7 - ~ I.gE~!lXJ: -0-Cif;"t"7J$-C';,5
Q04:fi?:,"'3-tr.*~O)OOW~0)7-~I.~.?:,9~~~~~V$KJ:-0-CT~~PV~*

AKic~k*5*~,4:fiO)MfrnO).f!;J~ID!:IjfJP'ifll0)~.ruO)W~KJ:-0-C~~.n,*~<2 ":) 0) 9- /v - 7 f~ 5t vt ~.n Q .:. c ?:' if; L -C l ' Q00

Commem.

{Zi>'<l'!I[Trans. Proc. Palaeont. Soc. Japan, N.S., No. 161, pp. 714-719,3 Figs., April 30, 1991

914. MIDDLE CARBONIFEROUS BACTRITOIDEA (MOLLUSCA:CEPHALOPODA) FROM THE AKIYOSHI LIMESTONE GROUP,YAMAGUCHI PREFECTURE (MOLLUSCAN PALEONTOLOGY
OF THE AKIYOSHI LIMESTONE GROUP-X)*
SHUJI NIKO
Department of Environmental Studies, Faculty of Integrated Arts
and Sciences, Hiroshima University, Hiroshima 730

T AMIO NISHIDA
Department of Earth Sciences, Faculty of Education,
Saga University, Saga 840
and

YUKO KYUMA
Nishiurakami Elementary School,
Nagasaki 850

Abstract. A Middle Carboniferous bactritoid fauna is described from the Akiyoshi
Limestone Group at Isa, Mine City, Yamaguchi Prefecture. The components of this fauna
and those age based on the associated foraminifers are: Bactrites nagatoensis, sp. nov. (Late
Bashkirian-Early Moscovian), B. sp. (Late Bashkirian), and Ctenobactrites sp. (Early
Moscovian). This is the first detailed systematic report of bactritoids in Japan.
Key words. Middle Carboniferous bactritoids, Bactrites, Ctenobactrites, Akiyoshi
Limestone Group.

assignment (Bactrites spp.) of the bactritoids.
Specimens utilized in this report are reposited in the paleontological collections of
the Akiyoshi-dai Museum of Natural History(ASM), Yamaguchi Prefecture.

Introduction
Many well-preserved bactritoids, whichhave not previously been described in Japan,
were recovered by two of us (T. Nishida andY. Kyuma) from the Akiyoshi LimestoneGroup at Isa, Mine City, Yamaguchi Prefecture (Figure 1). On the basis of these specimens, the present report describes three
species including a new taxon. Details of themode of occurrence and the associated molluscan fauna were previously reported by
Nishida and Kyuma (1986) with preliminary

Systematic paleontology
Su bclass Bactritoidea Shimansky, 1951
Order Bactritida Shim an sky, 1951
Family Bactritidae Hyatt, 1884
Genus Bactrites Sandberger, 1843

"Received April 18, 1990; revised manuscript accepted March I, 1991
714

914.

Carboniferous Bactritoidea from Akiyoshi

132'

,
,,,

N

.
,

,,

t
,,

,

Figure 1. Index map of Isa, Yamaguchi Prefecture, showing the bactritoid localities.

Bactrites nagatoensis Niko,
Nishida and Kyuma, sp. nov.
Figures 2, 3-1 - 5

Material. - Twenty-nine incomplete phragmocones were examined. Holotype, ASM
51310; paratypes, ASM 51311-51313,5135151355, 51384.
Diagnosis. - Large-sized Bactrites withcompressed cross section in mature shell andwell-developed dorsal carina; camera relatively long with deeply concaved septa;
sutures simple.
Description. - Phragmocone longiconic
orthocone, large; largest paratype (ASM
51355) having maximum diameter 62 mm;cross section of shell circular in juvenilephragmocone and laterally compressed inlater stages; holotype, 108 mm in length,
expands from 17 to 25 mm over 82 mm. A
single dorsal carina well-developed, pointed
in early juvenile phragmocone, later it

715

becomes indistinct ridge; weak transverse
lirae developed in later juvenile and matureshell. Septal concavity deep; sutures simple, slightly oblique, forwards aperture onventer, possess a ventral lobe at siphuncle;
cameral length relatively long, width/length
ratio of dorsoventral section through siphuncle ranges from 0.6 to 0.7; siphuncle ventral
and marginal with orthochoanitic necks.
Remarks. - Bactrites nagatoensis, sp. nov.
most resembles B. peytonensis Mapes, 1979,
from the Chesterian (Lower Carboniferous)of Arkansas, in that both taxa possess a compressed cross section in the later stages, adorsal carina and simple sutures. However,the longitudinal surface lirae and the transverse ribs on the mature shell of the latter
species are not observed in B. nagatoensis.
This new species also resembles Bactrites
finisensis Mapes, 1979, from the Virgilian(Upper Carboniferous) of Texas and B.
mexicanus Miller, 1944, from the Guadalupian (Upper Permian) of Las Delicias, Mexico in the possession of a dorsal carina. B.
finisensis differs from B. nagatoensis by thelower ratio of cameral length/shell diameter.
B. mexicanus can be differentiated from B.
nagatoensis in having a dorsal saddle.
Occurrence and age. - Bactrites nagatoensis, sp. nov. is locally a common componentof the Middle Carboniferous strata of the
Akiyoshi Limestone Group. With the
exception of a paratype (ASM 51384), which
was collected from a bioclastic grainstone in
the Profusulinella beppensis Zone (Late Bashkirian) at loc. 1M 64, specimens were found ina coquinoid limestone of the Fusulinellabiconica Zone (Early Moscovian) at loc. 1M
60.
Etymology. - The species is named for
Nagato, which is a medieval country name of
the type locality.

Bactrites sp.
Figures 3-6-9

Shuji Niko e/ at
7 16

7

914.

Carboniferous Bactritoidea from Akiyoshi

7 17

3

7

5

9

Figure 3. 1- 5, Baclriles nagaloensis, sp. nov. 1- 4, paratype, ASM 51312. 1, ventral view, X 3 ; 2,lateral view, venter on left, X 3 ; 3, dorsal view, X 3 ; 4, sep tal view, ve nter down , X 4. 5, paratype, ASM
Sl3II , dorsal view, X O.98. 6- 9, Bac/riles sp. , ASM 5 1385. 6, ve ntral view , X 2 ; 7, lateral view, ve nteron right, X 2; 8, dorsal view, X 2 ; 9, septa l view , ve nter down , X 4. 10, 11, Ctenobaclriles sp . 10, ASM
51223, side view, X 2; 11, ASM 51220, side view, X I.
~
Figure 2. Baclrires nagaloensis, sp. no v. 1- 6, ho lo type, ASM 51310. 1, ve ntral view, X I ; 2, lateralview, venter on right, X I ; 3, dorsal view, X I ; 4, septal view, venter down , X 2 ; 5, details of siphuncle
and ornamentation from adoral part of the holotype as viewed in Figure 2- 1, X 4, 6, dorsoventral polishedsection showing septal neck (arrow) , X 6. 7, paralype, ASM 5 1351 , do rsa l view, X 2. 8, paraty pe, ASM
51313 , dorsal view, X 3. 9, para type, ASM 51352, dorsa l view, X 3. 10, paratype, ASM 51353, ventral
view, X I .

Shuji Niko et al718

Material.-Single specimen (ASM 51385)
of incomplete phragmocone.
Remarks. - The specimen of an incomplete
phragmocone, 25 mm in length, is a longiconic orthocone with the circular crosssection and smooth shell surface. It can be
distinguished from the associated Bactrites
nagatoensis by the absence of dorsal carina.
The diameter of the conch is 8 mm at near the
apical end and expands up to approximately
10 mm over 19 mm. The siphuncle is relatively small and ventral position.
The morphology of the specimen is closely
related to some Carboniferous forms including Bactrites carbonarius Smith, 1903, B.
Jayettevillensis Mapes, 1979, B. longocameratus Shimansky, 1968, B. milleri Mapes, 1979,
and B. quadrilineatus Girty, 1909. However,the specimen is not complete enough for a
confident specific identification.
Occurrence and age. - Bactrites sp. wasrecovered from a bioclastic grainstone in the
ProJusulinella beppensis Zone (Late Bashkirian) at loc. 1M 64.

Ctenobactrites mirus Shimansky, 1954, fromthe Asselian to Sakmarian (Lower Permian)of the Urals, Soviet Union. The Russian
species is represented by a juvenile phragmocone. Thus, direct comparison with the present specimens is not possible.
The Virgilian (Upper Carboniferous)
species from Texas, Ctenobactrites sp. recorded by Mapes (1979) can be distinguished fromthe present form in that the latter possesses
much coarser rib spacing.
The combination of the transverse ribs anda longitudinal ridge in ornamentation of thepresent specimens also is reminiscent of
mature modification of some Bactrites, suchas B. peytonensis Mapes, 1979. However, theribbing on septate part of the shell and theflat-topped rib shape of the present specimens
are diagnostic of Ctenobactrites.
Occurrence and age. - Ctenobactrites sp.
was recovered from a coquinoid limestone in
the Fusulinella biconica Zone (Early Moscovian) at loco 1M 60.
Acknowledgments

Genus Ctenobactrites Shimansky, 1951
Ctenobactrites sp.
Figures 3-10, II

Material.-Four specimens (ASM 51220,
51223-51225) of fragmentary phragmocones.
Description. - Phragmocone 10ngiconic
orthocone, moderately large; largest specimen (ASM 51220) having maximum diameter
34 mm. Surface ornamentation consists offlat-topped transverse ribs and a single longitudinal ridge; width of grooves separatingribs amounts to 1.2 mm in largest specimen;
each rib ranges from 1.2 to 2.9 mm in width,and forming shallow sinus at ridge. Septa
moderately curved with simple, slightly
oblique sutures.
Remarks. - The present specimens havecharacteristic ornamentation of the flattopped transverse ribs with a longitudinalridge that is the typical morphology ofWe thank Drs. Takashi Hamada and Kazushige Tanabe (University of Tokyo) for helpful comments. Dr. Royal H. Mapes (Ohio
University) reviewed the manuscript and provided the North American bactritoid specimens. Laboratory research was supported
by a grant from the Japanese Ministry of
Education, Science and Culture (No.
01790346) to S. Niko.

References
Girty, G.H., 1909: The fauna of the Caney Shale ofOklahoma. U.S. Geol. Surv., Bull., 377, 106 p.
Hyatt, A., 1883-1884: Genera of fossil cephalopods.
Proc. Boston Soc. Nat. Hist., vol. 22, pt. 3, p.
253-338.
Mapes, R.H., 1979: Carboniferous and Permian
Bactritoidea (Cephalopoda) in North America.
Univ. Kansas Paleont. Contrib., Art. 64, 75 p., 41
pIs.
Miller, A.K., 1944: Geology and paleontology of the914.

Carboniferous Bactritoidea from Akiyoshi

Permian area northwest of Las Delicias, southwestern Coahuila, Mexico, Part IV, Permiancephalopods. Geol. Soc. America, Special
Paper, 52, p. 71-127, pis. 20-45.
Nishida, T. and Kyuma, Y., 1986: On the molluscan
fauna. Abstracts of Palaeontological Societyof Japan, 135th Annual Meeting at Kitakyushu, p. 41-42. (in Japanese)
Sandberger, G., 1843: Schilderung der paHiontologischen Verhaltnisse der alteren Formationen Nassa us. Amtl. NaturJ Aerzte Mainz, p. 154-160.
Shimansky, V.N., 1951: K voprosu ob evoliutsii
verkynepaleozoishikh pryamkh golovongikh.

Akad. Nauk SSSR Dokl., vo!' 79, no. 5, p. 867870. (in Russian)- - , 1954: Pryamye nautiloidei i baktritoidei sakmarskogo i artinskogo yarusov Yuzhnogo Urala.
Akad. Nauk SSSR, Paleont. Inst. Trudy, vo!. 44,
156 p., 12 pIs. (in Russian)- - , 1968: Kamennougoinye Orthoceratida, Oncoceratida, Actinoceratida i Bactritida. Ibid.,
vo!.117, 151 p., 20 pIs. (in Russian)Smith, J.P., 1903: The Carboniferous ammonoids ofAmerica. U.S. Geol. Surv., Monogr., vol. 42,
211 p.

Akiyoshi f;kE". Isa W-fti:. Mine ~iil. Nagato *F~.
f;kE":Ow(:E~~I.J' G 0) '*':Jtil:OER*C/~ 9 ~ ~ T .A m:

ilJ I:::J !If-:W-fti:jfuJW(O)f;kE":Ow(:E~~I.J'

Gft. ~~0)/~9 ~ ~ T.Am~~t±:l-r-~o .::.O)tb!lo/.J~O)mij\(;:fj. :t;J: tfIliR1f-r-~:.ffR~1t:0 J: I) ~ljrtJT ~;h ~ ffif~ ft. Bactrites nagatoensis, sp. nov. (/~ -/ =f- -/vi!t1&:JtilI.J, G.'f:-.A:2 - i!tlitr:Jtil), Bactrites sp. (/~ -/=f- - /vi!t1&:Jtil). Ctenobactrites sp. ( .'f:- .A :2 - i!tlitr:Jtil) -e it>~o¥a 15' ft. OOF9I.J'GO)/~ 9 ~ ~ T .AmO):m~cllljU: tJ:~o
;L

*

719

;L

r,'C..:r~l'f] • ®rnR,tllt • 0..r,,9rG.:rTrans. Proc. Palaeont. Soc. Japan, N.S., No. 161, pp. 720-738, II Figs., April 30,1991

915.

EARLY JURASSIC RADIOLARIANS FROM THE NANJOMASSIF IN THE MINO TERRANE, CENTRAL JAPANPART 1. TRICOLOCAPSA, STICHOCAPSA,AND MINOCAPSA, N. GEN.*A TSUSHI MATSUOKA
Department of Earth Sciences, College of General Education,
Niigata University, Niigata 950-21

Abstract. Well-preserved Early Jurassic radiolarians are obtained from a manganeseband in the Nanjo Massif, Mino Terrane, central Japan. This paper is Part I of a serial
descriptive work on an Early Jurassic radiolarian fauna from the Nanjo Massif and deals
with nassellarians which possess a constricted or closed distal end. One new genus,
Minocapsa, is erected, and 9 new species and 2 new subspecies are described: Tricolocapsa
minoensis, T. (?) megaglobosa, T. (?) jera, Stichocapsa plicata plicata, S. plicata semiplicata, S. elegans, S. biconica, S. nanjoensis, Minocapsa cylindrica and M. globosa.
The fauna from the Nanjo Massif is very similar to that of the upper Archicapsa
pachyderm a Zone of Jurassic radiolarian zones in Japan. The fauna is correlative with thelower Toarcian fauna in east-central Oregon, North America.
Key words.

Early Jurassic, Mino Terrane, Nanjo Massif, Nassellaria, Radiolaria.

composition of assemblages needs to beclarified.
Manganese bands within Early to Middle
Jurassic chertsi1iceous mudstone sequences in
the Mino Terrane, central Japan yield excellently preserved radiolarian tests. Since Yao(1972) introduced some "spongosaturnalids"
from the manganese bands in the Inuyama
area, several radiolarian workers have described new species from the manganesebands in the Mino Terrane (Baumgartner,
1984; Ichikawa and Yao, 1976; Isozaki andMatsuda, 1985; Takemura, 1986; Takemura
and N akaseko, 1982, 1983, 1986; Yao, 1979).
In a series of works by this author, a wellpreserved radiolarian fauna from a manganese band of Early Jurassic age (early Toarcian) in the Nanjo Massif, Mino Terrane,
central Japan is systematically described. In
this paper (Part 1 of this series), one new
genus is erected, and 9 new species and 2 new

Introduction
In this decade, Early Jurassic radiolarians
have been described from Turkey (De Wever,
1981 a, 1981 b, 1982; Pessagno and Poisson,1981), from North America (Carter et at.,1988; MacLeod, 1988; Pessagno and Blome,
1980; Pessagno and Whalen, 1982; Pessagno
et at., 1986; Yeh, 1987a, 1987b) and fromJapan (Hori, 1988; Hori and Otsuka, 1989;
Hori and Yao, 1988; Isozaki and Matsuda,
1985; Kishida and Hisada, 1985: Sash ida,
1988; Takemura, 1986; Takemura and Nakaseko, 1982, 1983, 1986; Yao, 1982). In
spite of these descriptive works, we often
encounter undescribed forms in Lower Jurassic rocks. In order to establish more detailed
radiolarian zonation and to recognize faunal
provinces of radiolarians, the entire specific"Received August 14, 1990; revised manuscript accepted January 10, 1991
720

915.

Jurassic radiolarians from the Nanjo Massif

subspecies of nassellarians with a constricted
or closed distal end (i.e. Tricolocapsa, Stichocapsa, and Minocapsa, n. gen.) are described.
Acknowledgments: The author is gratefulto Drs. Akira, Yao and E.S. Carter for theirsuggestion to this study and for critical reading of the manuscript. This work is supported by the Grant-in-Aid for Scientific Researchof the Ministry of Education, Science andCulture of Japan (Grant No. 02740392).

Geologic setting
The Mino Terrane, one of the Jurassic-

721

Early Cretaceous accretionary complexes inJapan, is distributed in the central part ofJapan (Figure I). This terrane is composed
mainly of Jurassic-Early Cretaceous clastic
rocks associated with chert (Permian-Early
Jurassic), limestone (Carbonifero us- Permian)
and greenstone. This terrane is considered
to be constructed through accretionary processes at a convergent plate boundary during
Jurassic and Early Cretaceous time. The
Nanjo Massif (Figure I) occupies the northwestern part of the Mino Terrane. The Jurassic complex in this area is divided into threeunits characterized by distinct litho-facies;
namely Kasugano, lmajo, and Kohkura facies

Figure 1. Map showing the fossil locality, MNA-IO (using I: 50,000 topographic map of "Imajo" published by Geographical Survey Institute of Japan).
The dotted area in the inset map shows the distribution of the Jurassic-Early
Cretaceous accretionary complex of the Mino Terrane, central Japan.

Atsushi Matsuoka

722(Hattori and Yoshimura, 1982). According
to Hattori and Yoshimura (1982), theKasugano unit consists of Early Jurassic
mudstone, greenstone, and olistoliths of
Paleozoic limestone, greenstone and chert.
The Imajo unit is composed of sandstone andmudstone with a lesser amount of siliceous
mudstone. The chert encountered in thisunit is Triassic in age and occurs as majorolistoliths in Jurassic sequences. The Kohkura unit is characterized by coarse-grained
massive sandstone. This unit also belongs to
the Middle Jurassic and contains laterallytraceable thick chert of Triassic age.
Since Ito and Shiratake (1980), and Ito andMatsuda (1980) reported Jurassic radiolarians
from the Nanjo Massif, the fossil record ofJurassic radiolarians has rapidly accumulated(Hattori, 1984, 1987, 1988, 1989; Hattori and
Sakamoto, 1989; Hattori and Yoshimura,
1982, 1983; Takamura and Hayami, 1985;
Yoshimura et al., 1982). Among these,
Hattori (1987, 1988, 1989), and Hattori and
Sakamoto (1989) figured a great number ofradiolarians from mudstone, siliceous mudstone and manganese band samples, and discussed the geologic age of the terrane basedon the radiolarian assemblages.

Material and methods
Radiolarian fossils treated throughout thisseries of studies are from a manganese band
contained in a siliceous mudstone layer
belonging to the Imajo unit. The band has alenticular shape with a maximum thickness of
20 cm. It was obtained from an outcrop
along a road cutting. Figure I shows the
locality of rock sample MNA-IO. This band
contains manganese carbonate spherules ranging 0.5-2.0 mm in diameter from which wellpreserved radiolarian tests were extracted.
The rock sample was broken into small
fragments of a few centimeters size and put inHel solution for one day. The residue wasgathered using # 35 and # 200 sieves. For
scanning microscope observation, radiolarian

tests were picked under a binocular microscope and mounted on aluminium stubs.
For optical microscope observation, theun picked residue containing radiolarian tests
was mounted on a slide glass with the synthetic medium "Entellan New".

Radiolarian fauna and its age assignment
Sample MN A -10 contains more than 200
species of radiolarians. The following taxa
are identified at species level: Acanthocircus
bispinus (Yao), Archicapsa pachyderm a (TanSin Hok), Bistarkum rigidium Yeh, Canoptum artum Yeh, Canoptum poissoni Pessagno, Canoptum spinosum Yeh, Cuniculiformis aristotelis De Wever, Hsuum acutum
Yeh, Hsuum gratum Yeh, Hsuum lucidum
Yeh, Hsuum (?) unicum Yeh, Katroma kurusuensis Hori, Paracanoptum anulatum (Pessagno and Poisson). In addition to them,
forms belonging to genera Higmastra Baumgartner, Canutus Pessagno and Whalen,
Combusta Yeh, Lantus Yeh, Pleesus Yeh
and Pseudopantanellium Yeh are obtained.
Because sample MNA-IO contains no agediagnostic fossils other than radiolarians, age
assignment is inferred from comparisons withradiolarian faunas in regions where age iscontrolled by fossil groups such as ammonites. The fauna of sample MNA-IO contains many taxa which Yeh (1987a, 1987b)
described from the Hyde Formation and thebasal Warm Springs Member of the Snowshoe Formation in the Suplee-Izee area, eastcentral Oregon, North America, as listed
above. The Hyde Formation and the basal
Warm Springs Member are regarded as lower
Toarcian based on stratigraphic relationships
with the under- and overlying formations
which yield age-diagnostic ammonites (Pessagno et al., 1986; Yeh, 1987a). This fauna
includes forms assignable to the genus
Canutus Pessagno and Whalen which Pessagno et al. (1987) used as a corporeal taxon
disappearing within the Subzone 01B (Lower
Toarcian) of their Jurassic radiolarian zona-

915.

Jurassic radiolarians from the Nanjo Massif

tion in North America. The fauna also contains forms assignable to the genus Higmastra
Baumgartner, a supplementary zonal marker,
which appears near the base of the Subzone
01B. In addition, no mesosaturnalids are
obtained from the sample, although various
other forms of saturnalid spumerallians arepresent. Species of Mesosaturnalis (M. hexagonus Yao and M. sp. cf. M. septispinusYao) are found in the middle to upper Toarcian of the Maude Group, Queen Charlotte
Islands, British Columbia (Carter et al., 988).
Above-mentioned data suggest that the faunaof sample MNA-1O is correlative with thelower Toarcian fauna in eastcentral Oregon,
North America.
Matsuoka and Yao (1986) proposed eightradiolarian zones for the Jurassic of Japan on
the basis of biostratigraphic investigations on
continuous stratigraphic sequences in Southwest Japan. These zones are the Parahsuum
ovale (= Parahsuum sp. C in Matsuoka andYao, 1986), Archicapsa pachyderma, Laxtorum (?) jurassicum, Tricolocapsa plicarum,
Tricolocapsa conexa, Stylocapsa (?) spira lis,
Cinguloturris carpatica and Pseudodictyomitra primitiva Zones in ascendingorder. The faunal composition of sample
MN A -10 is very similar to that of the upper
Archicapsa pachyderma Zone (middle Lower
Jurassic).

Systematic paleontology
The genera treated in this paper are notassigned to families because classification at
the family level is not adequately settled atpresent.
Type and figured specimens are registeredand deposited in the Department of Earth
Sciences, College of General Education,
Niigata University. In the systematic description and explanation of figures, the NU
MR number is the register number of thespecimens in that Department. In explanation of figures, the number in parentheses
following the NU MR number indicates the723

registered film number.
Subclass Radiolaria Muller, 1858
Superorder Polycystina Ehrenberg, 1838,
emend. Riedel, 1967
Order N assellaria Ehrenberg, 1875
Genus Tricolocapsa Haeckel, 1881
Type species: Tricolocapsa theophrasti
Haeckel, 1881
Tricolocapsa minoensis
Matsuoka, n. sp.
Figures 2-1a-5b
Tricolocapsa sp. cf. T. plicarum Yao, Y 0shimura et al., pI. 6, fig. 4.
1982 Tricolocapsa (?) sp. A, Yoshimura et al., pI. 6,
fig. 5.
1985 Tricolocapsa sp. A, Takamura and Hayami, pI.
3, fig. 4.
1987 Tricolocapsa sp. A, Hattori, pI. 13, fig. 1.
1987 Tricolocapsa sp. B, Hattori, pI. 13, fig. 2.
1989 Tricolocapsa sp. A, Hattori, pI. 10, fig. A.
1989 Tricolocapsa spp., Hattori, pI. 10, fig. D.
1982

Description. - Shell of three segments,
drop-like shaped. Cephal is hemispherical,
poreless. Thorax truncate conical. Abdomen large, subspherical with a constricted
aperture. Collar and lumber stricturesslightly recognizable or indistinct externally.
Outer surface of shell ornamented with continuous longitudinal plicae. Eleven to 15
moderately spaced plicae visible on outershell. One row of pores present between
longitudinal plicae. Pores small to moderate
in size, circular and uniform in shape. Aperture moderate in size, circular.
Measurements (in ,um).-Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell, DA; diameter of aperture.
Holotype max. mm. mean
TH
115
115
93
105
(15)
MW
94
94
72
83
(15)
DA
13
14
12
13
(6)
Remarks. - Pores vary in size among specimens, from small (Figures 2-1 a, b) to moderate (Figure 2-3).

724

Atsushi Matsuoka

Figure 2. Tricolocapsa minoensis Matsuoka, n. sp. 1- 4; Scanning electron micrographs, S ; transmitted light micrographs. All figures X 380. la, b. Holotype, NU MR 0001 (a ; 11859, b ; 11860), b ;basal view. 2a, b. Paratype, NU MR 0002 (a ; 11 801 , b; 11802), b; bas al view. 3. Paratype, NU MR
0003 (11415). 4. Paratype, NU MR 0004 (11057). Sa, b. Paratype, NU MR 0005 (a ; 93 - 21 , b ; 93- 22) .

Tricolocapsa minoensis, n. sp. differs fromT. plicarum Yao by lacking a dish-like basal
appendage, by having more spaced plicae
and by its smaller size.
Ety mology.- This species is named for the
Mino Terrane which includes the type area,
Nanjo Massif.
Type specimens .-Holotype NU MR 0001
(Figures2- 1a, b); Paratypes NU MR 0002
(Figures 2-2a, b), NU MR 0003 (Figure 2- 3),
NU MR 0004 (Figure 2-4), NU MR 0005
(Figures 2-5a, b).
Type locality.-MNA - IO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence . -Nanjo Massif, Mino Terrane,
central Japan . Kuma area, Southern Chichibu Terrane, western Kyushu (Matsuoka,
unpublished data) .
Range. - Archicapsa pachyderma Zone
(middle Lower Jurassic) as far as known .

Tricolocapsa (?) megaglobosa
Matsuoka, n. sp.
Figures 3- 1a - 5b

Description. - Shell of three segments with adish-like basal appendage. Cephal is hemispherical, poreless. Thorax truncate-conicalwith circular, densely spaced pores. Abdomen large, inflated, barrel-shaped with circular, densely spaced pores larger than those inthorax. Collar stricture distinct externally.
Lumber stricture slightly recognizable externally . Joint between abdomen and basal
appendage marked by a row of pores slightly
larger than those in abdomen . Basal
appendage half to a third of abdomen in
width, with circular to subcircular densely
spaced pores smaller than those in remaining
part of shell surface.

915.

Jurassic radiolarians from the N anjo Massif

725

Figure 3. Tric%capsa (?) megag/obosa Matsuoka, n. sp. 1- 4; ~cannlng electron mIcrographs, S ;tran smitted light micrographs. All figures X 380. la, b. Holotype, NU MR 0006 (a ; 11827, b ; 11828), b ;basal view. 2a, b. Paratype, NU MR 0007 (a ; 11825, b ; 11826), b ; basa l view. 3a, b. Paratype, NU MR
0008 (a ; 11771 , b ; 11772), b; basal view. 4. Paratype, NU MR 0009 (11810). Sa, b. Paratype, NU MR
0010 (a ; 96- 05 , b ; 96- 06).

726

A tsushi Matsuoka

Measurements (in ,um).-Numbers of specimens measured are in parentheses. TH;
total height of shell , MW ; maximum widthof shell, A W ; width of basal appendage.
Holotype max. min . mean
TH
153
160
120
145
(12)
MW
105
117
88
101
(12)
AW
55
61
47
55
(II)
Remarks. - This species is questionably
assigned to Tricolocapsa beca use it possesses
a dish-like basal appendage. This species isdistinguished from Tricolocapsa (?) fusiformis Yao by having a large, inflated abdomen ,densely spaced pores and a distinct collar
stricture.
Etymology. - This specific name is derived
from the Latin mega ( = large) and globosus-

a-um (= spherical).
Type specimens. - Holotype NU MR 0006
(Figures 3-la, b) ; Paratypes NU MR 0007
(Figures 3-2a, b), NU MR 0008 (Figures 33a, b), NU MR 0009 (Figure 3-4), NU MR
0010 (Figures 3-5a, b).
Type locality.- MNA - lO , Nanjo Massif,
Mino Terrane, central Japan .
Occurrence. -Nanjo Massif, Mino Terrane,
central Japan .
Range. - Archicapsa pachyderm a Zone
(middle Lower Jurassic) as far as known.
Tricolocapsa (?) fera Matsuoka, n. sp.
Figures 4-la - 3b

Figure 4. Tricolocapsa (?) fera Matsuoka, n. sp. 1- 2; Scanning electron micrographs, 3 ; transmitted light micrograph s. All figures X 380. la, b. Holotype, NU MR 0011 (a; 12299, b ; 12300), b; obliquebasal view. 2a, b. Paratype, NU MR 0012 (a; 11735, b ; 11736), b ; oblique basal view. 3a, b. Paratype,
NU MR 0013 (a ; 90- 17, b ; 90- 18).

915.

Jurassic radiolarians from the Nanjo Massif

Description. - Shell of three segments,
pyriform. Cephal is hemispherical, porelesswith a small apical horn. Thorax truncate
conical or cylindrical with small pores.
Abdomen large, subspherical with a constricted aperture. Collar stricture slightly recognizable externally. Lumber stricture pronounced. Pores moderate in size, circular to
subcircular, densely spaced and set inpolygonal (largely hexagonal) pore frames.
Abdomen possessing small pointed spines
situated at the pore frame vertices. Aperture
moderate in size, circular with a shortprotruding rim.
Measurements (in Jim). - Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell, DA; diameter of aperture.
Holotype max. mm. mean
TH
139
146
125
133
(6)
(6)
MW
100
125
96
107
(5)
II
14
8
II
DA
Remarks. - This species is questionably
assigned to Tricolocapsa because it possessesa small apical horn. However, it is uncertain
whether the presence of an apical horn is an
important criterion for classification at
generic level of three-segmented nassellaria.
The horn seems to be an extension of apicalspine. Tricolocapsa (?) fera, n. sp. is similar
to T. tetragona Matsuoka and T. sp. cf. T.
ruesti Tan Sin Hok (in Yao, 1979) in generaloutline but differs from both by having anapical horn and by densely spaced pores on
the abdomen.
Etymology. - This specific name is derived
from the Latin ferus-a-um (=wild).
Type specimens. - Ho10type NU MR 0011
(Figures 4-la, b); Paratypes NU MR 0012
(Figures 4-2a, b), NU MR 0013 (Figures 43a, b).
Type locality.-MNA-IO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence. - Nanjo Massif, Mino Terrane,
central Japan.
Range. Archicapsa pachyderm a Zone
(middle Lower Jurassic) as far as known.

727

Genus Stichocapsa Haeckel, 1881
Type species: Stichocapsa
jaspida RUst, 1885

Stichocapsa plicata Matsuoka, n. sp.
Figures 5-la-5b: 6-la-6b
1989
1989
1989
1989

Tricolocapsa sp. 8, Hattori, pI. 10, fig. B.
Tricolocapsa spp., Hattori, pI. 10, fig. F.
Tricolocapsa sp. D, Hattori, pI. 29, fig. D.
Tricolocapsa sp., Hattori and Sakamoto, pI. 19,
fig. B.

Description. - Shell of four to six segments,
drop-like or spindle shaped. Cephalis
hemispherical, poreless. The last segment
large, inverted hemispherical or inverted conical with a constricted aperture. Remaining
segments including thorax and abdomen truncate conical. Strictures between segments
indistinct externally. Fifteen to 22 densely
spaced longitudinal plicae visible on outershell except for cephalis. The longitudinal
plicae distinct or partly obscure. One row ofpores present between the plicae. Pores uniform, small and circular. Aperture small,circular, occasionally with a short protrudingnm.
Measurements (in Jim).-Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell, DA; diameter of aperture.
Holotype max. mm. mean
TH
154
170
110
142
(29)
MW
108
124
80
96
(29)
DA
8
8
6
7
(12)
Remarks. - Two subspecies are included
under this species; these are S. plicata
plicata, n. subsp. and S. plicata semiplicata,n. subsp. S. plicata, n. sp. differs from S.
convexa Yao by having longitudinal plicae
between which one row of pores is present.
Etymology. - This specific name comesfrom the Latin plicatus-a-um (= plicate).
Type specimens. -Holotype NU MR 0014
(Figures5-la, b); Paratypes NU MR 0015
(Figures 5-2a, b), NU MR 0016 (Figures 5-

728

Atsushi Matsuoka

Figure 5. Slichocapsa plicala plicala Matsuoka, n. subsp. 1-4; Scanning electron micrographs,S ;transmitted light micrographs. All figures X 380. la, b. Holotype, NU MR 0014 (a; 11823, b; 11824), b ;basal view. 2a, b. Paratype, NU MR 0015 (a ; 11757, b ; 11758), b ; basal view. 3a, b. Paratype, NU MR
0016(a; 11776,b; 11777), b ; oblique basal view. 4. Paratype, NU MROOI7(11731). 5a,b. Paratype,
NU MR 0018 (a; 94- 23, b; 94-24).

915.

Jurassic radiolarians from the Nanjo Massif

3a, b), NU MR 0017 (Figure 5-4), NU MR
0018 (Figures 5-5a, b).
Type locality.-MNA-lO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence. - N anjo Massif, Mino Terrane,
central Japan.
Range. - Archicapsa pachyderm a Zone
(middle Lower Jurassic) as far as known.

Stichocapsa plicata plicata
Matsuoka, n. subsp.
Figures 5-1 a - 5b

1989
1989
1989
1989

Tricolocapsa sp. B, Hattori, pI. 10, fig. B.
Tricolocapsa spp., Hattori, pI. 10, fig. F.
Tricolocapsa sp. D, Hattori, pI. 29, fig. D.
Tricolocapsa sp., Hattori and Sakamoto, pI. 19,
fig. B.

Description. - Shell of four to five segments,
drop-like or spindle shaped. Cephalis
hemispherical, poreless. The last segment
large, inverted hemispherical or inverted conical with a constricted aperture. Remaining
segments including thorax and abdomen truncate conical. Strictures between segments indistinct externally. Fifteen to 22 densely
spaced longitudinal plicae visible on outershell. The plicae distinct on whole shell
except for cephalis. One row of pores present between the plicae. Pores uniform, small
and circular. Aperture small, circular, occasionally with a short protruding rim.
Measurements (in pm).-Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell, DA; diameter of aperture.
Holotype max. mIn. mean
TH
154
170
110
142
(20)
MW
108
124
82
102
(20)
DA
8
8
6
7
(9)
Remarks. - Shape varies among specimens;
some are drop-like shaped (Figure 5-la, 3a, 4,
5a, b) and others spindle shaped (Figure 52a). Stichocapsa plicata plicata, n. subsp.
differs from S. plicata semiplicata, n. subsp.
by having distinct longitudinal plicae that729

extend from the thorax to the distal end.
Etymology. - This subspecies is thenominotypical subspecies of Stichocapsa
plicata, n. sp.
Type specimens.-Holotype NU MR 0014
(Figures 5-1a, b); Paratypes NU MR 0015
(Figures 5-2a, b), NU MR 0016 (Figures 53a, b), NU MR 0017 (Figure 5-4), NU MR
0018 (Figures 5-5a, b).
Type locality.-MNA-lO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence. - N anjo Massif, Mino Terrane,
central Japan.
Range. Archicapsa pachyderma Zone
(middle Lower Jurassic) as far as known.

Stichocapsa plicata semiplicata
Matsuoka, n. subsp.
Figures 6-la-6b

Description. - Shell of five to six segments,
drop-like shaped. Cephalis hemispherical,
poreless. The last segment large, truncate
subspherical with a constricted aperture.
Remaining segments including thorax andabdomen truncate conical. Strictures
between segments indistinct externally. Fifteen to 22 densely spaced longitudinal plicae
visible on outer shell except for cephalis andthe middle part of shell. Pores small, circular and arranged longitudinally. One row ofpores present between the plicae. Aperturesmall, circular, occasionally with a shortprotruding rim.
Measurements (in pm).-Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell, DA; diameter of aperture.
Holotype max. mIn. mean
TH
139
149
123
142
(9)
MW
95
99
80
94
(9)
DA
7
8
7
7
(3)
Remarks. - Stichocapsa plicata semiplicata, n. subsp. is compared to S. plicata
plicata, n. subsp. under the latter subspecies.
Etymology. - The subspecific name comes

730

Atsushi Matsuoka

Figure 6. Slichocapsa plicala semiplicala Matsuoka, n. subsp. 1-5 ; Scanning electron micrographs,
6' transmitted light micrographs. All figures X 3S0. la, b. Holotype, NU MR 0019 (a ; IISI9, b ; IIS20),
b ; basal view. 2a, h. Paratype, NU MR 0020 (a; 12296, h ; 12297), h ; oblique basal view. 3. Paratype,
NU MR 0021 (1IS05). 4. Paratype, NU MR 0022 (11072). 5. Paratype, NU MR 0023 (11349). 6a, h.
Paratype, NU MR 0024 (a; 80- 02, h ; 80- 03).

915.

Jurassic radiolarians from the Nanjo Massiffrom the Latin semi ( = half) and plicatus-aum ( = plicate).
Type specimens.-Holotype NU MR 0019
(Figures 6- 1a, b); Paratypes NU MR 0020
(Figures 6- 2a, b), NU MR 0021 (Figure 6-3),
NU MR 0022 (Figure 6- 4), NU MR 0023
(Figure 6- 5), NU MR 0024 (Figures 6-6a, b).
Type locality. - MNA - IO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence. - Nanjo Massif, Mino Terrane,
central Japan .
Range. Archicapsa pachyderm a Zone731

(middle Lower Jurassic) as far as known.

Stichocapsa elegans Matsuoka, n. sp.
Figures 7- 1a - 5b

Description. - Shell of four to five segments,
elongate ovoidal. Cephal is hemispherical,
poreless. Thorax, abdomen and the fourth
segment in the case of five-segmented specimens truncate conical. All segments but thelast one form conical proximal part; the last

Figure 7. Stichocapsa elegans Matsuoka, n. sp. 1- 4; Scanning electron micrographs, S; transmitted
light micrographs. All figures X 380. la, b. Ho1otype, NU MR 0025 (a ; 12233, b ; 12234), b ; basal view.
2a, b. Paratype, NU MR 0026 (a ; 12237, b ; 12239), b; oblique apical view. 3. Paratype, NU MR 0027
(11834). 4. Paratype, NU MR 0028 (11835). Sa, b. Paratype, NU MR 0029 (a ; 95 - 14, b ; 95- 15).

732

Atsushi Matsuoka

segment large, hemispherical with a strongly
constricted aperture. Strictures between segments indistinct externally. Ten to 12 longitudinal plicae observed on the conical proximal portion of shell in lateral view. One
row of pores runs on the plicae. The distal
portion of shell with smooth, perforated surface . Pores small and circular.
Measurements (in ,um).-Numbers of specimens measured are in parentheses. TH ;total height of shell, MW ; maximum widthof shell, DA ; diameter of aperture.
Holotype
max . min . mean
TH
170
179
152
165
(IS)
MW
(15)
89
105
81
89
DA
(2)
8
8
6
7
R emarks. - Stichocapsa elegans, n. sp. dif-fers from S. plicata , n. sp. by its slender formand by possessing perforated longitudinal
plicae on the conica l proximal portion of theshell.
Etymology. - The specific name comes fromthe Latin elegans ( = elegant).
Type specimens. -Holotype NU MR 0025
(Figures7 - la, b) ; Paratypes NU MR 0026
(Figures 7- 2a, b), NU MR 0027 (Figure 7-3 ),NU MR 0028 (Figure 7- 4), NU MR 0029
(Figures 7- 5a, b) .
Type locality. - MNA - IO, Na njo Massif,
Mino Terrane, centra l Japan .
Occurrence. - Nanjo Massif, Mino Terrane,
centra l Japan .
Range. Archicapsa pachyderma Zone
(middle Lower Jurassic) as far as known.

Figure 8. Slichocapsa biconica Matsuoka, n. sp. 1- 4 ; Scanning electron micrographs, S ; transmitted light micrographs. All figures X 380 except for Ib and 2b ( x 500) . la, b. Holotype, NU MR 0030 (a ;11727, b ; 11728), b ; basal view. 2a, b. Para type, NU MR 0031 (a ; 11725, b; 11726), b ; basal view. 3.
Paratype, NU MR 0032 (11722). 4. Paratype, NU MR 0033 (11778). Sa, b. Paratype, NU MR 0034. (a ;90- 13, b ; 90- 14).

915.

Jurassic radiolarians from the Nanjo Massif

733

Figure 9. Stichocapsa nanjoensis Matsuoka, n. sp. 1- 2 ; Scanning electron micrographs, 3- 4; transmitted light micrographs. All figures X 380. 1. Holotype, NU MR 0035 (12298). 2. Paratype, NU MR
0036 (12295). 3. Paratype, NU MR 0037 (80- 11). 4a, b. Paratype, NU MR 0038 (a; 90- 04, b; 90- 05).

Stichocapsa biconica Matsuoka, n. sp.
Figures 8- 1a - 5b

Description. - Shell of five segments,
spindle-shaped . Cephalis hemispherical,
poreless, occasionally with rough surface.
Thorax and abdomen truncate conical.
Forth segment barrel-shaped. The last segment inverted conical with a constricted aperture. Collar stricture rather disti nct. Strictures between segments, except for the collar
one, indistinct. Pores small to moderate andcircular to subcircular. Aperture moderate
in size, circular.
Measurements (in ,um). - Numbers of specimens measured are in parentheses. TH;
total height of shell , MW; maximum widthof shell , DA ; diameter of aperture.
Holotype
max. min. mean
( IS)
153
153
110
130
TH
( IS)
77
77
57
68
MW
(2)
9
999
DA
Remarks. - This species differs fromCy rtocapsa (?) kisoensis Yao by consisting offive segments rather than four and by lacking
an apical horn.
Etymology. - The specific name is derived
from the Latin bi ( = two) and conicus-a-um( = conical) .
Type specimens. - Holotype NU MR 0030
(Figures 8- la , b) ; Paratypes NU MR 0031
(Figures 8- 2a , b), NU MR 0032 (Figure 8- 3),

NU MR 0033 (Figure 8-4), NU MR 0034
(Figures 8- 5a, b) .
Type locality. - MNA - IO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence. - Nanjo Massif, Mino Terrane,
central Japan .
Range. - A rchicapsa pachyderma Zone
(middle Lower Jurassic) as far as known.

Stichocapsa nanjoensis Matsuoka , n. sp.
Figures 9- 1 - 4b

Description. - Shell of four segments,
spindle-shaped. Cephalis hemispherical,
poreless. Thorax truncate conical , abdomen
barrel-shaped and fourth segment inverted
conical with a constricted , circular aperture .
Collar stricture pronounced externally.
Other segmental joints indistinct externally.
Eight to 10 weakly developed longitudinal
plicae run from thorax to the distal end inlateral view. One row of pores arranged
between the longitudinal plicae. Pores circular to subcircular, small and uniform in
size.
Measurements (in ,um). - Numbers of specimens measured are in parentheses. TH;
total height of shell, MW ; maximum widthof shell , DA ; diameter of aperture.

A tsushi Matsuoka

734

Figure 10. Minocapsa cylindrica Matsuoka, n. gen . et n. sp. 1- 4 ; Scanning electron micrographs, S ;transmitted light micrographs. All figures X 250. la, b. Holotype, NU MR 0039 (a ; 11853, b; 11854), b;
oblique basal view . 2a, b. Paratype, NU MR 0040 (a ; 12247, b; 11248), b ; oblique basal view. 3.
Paratype, NU MR 0041 (12275). 4. Paratype, NU MR 0042 (12246). Sa, b. Paratype, NU MR 0043 (a ;94- 11 , b ; 94- 12).

Holotype
104
62

max.
104
63

min.
90
54

mean
96
(10)
59
(10)
8
(5)
8
7
Remarks. - Stichocapsa nanjoensis , n. sp.
is distinguished from S. biconica , n. sp. by its
small size, by consisting of four segments
rather than five and by having longitudinal
plicae. It also differs from Cyrtocapsa (?)
kisoensis Yao by having longitudinal plicae
and by lacking an apical horn.
Etymology. - The species is named for the
Nanjo Massif, its type locality.
Type specimens.-Holotype NU MR 0035
(Figure 9-1) ; Paratypes NU MR 0036 (Figure 9- 2) , NU MR 0037 (Figure 9-3) , NU MR
0038 (Figures 9- 4a , b).
TH
MW
DA

Type locality.-MNA - IO, Nanjo Massif,
Mino Terrane , central Japan .
Occurrence. - Nanjo Massif In the Mino
Terrane, central Japan.
Range.- Archicapsa pachyderma Zone
(middle Lower Jurassic) as far as known .
Genus Minocapsa Matsuoka , n . gen .

Type species : Minocapsa cylindrica Matsuoka , n. sp.

Diagnosis. - Closed cyrtid. Shell consi sting of four or more segments, pyriform to
ovoidal. Cephalis hemispherical without
apical horn. Thorax and abdomen truncate
conical. Final segment large, hemi sphericalwithout aperture . Pores, circular to subcir-

915.

Jurassic radiolarians from the Na njo Massif

735

Figure 11. Minocapsa globosa Matsuo ka, n. ge n. et n. sp. 1- 3; Scanning electron micrographs, 4 ;transmitted light micrographs. All figures X 250. la, b. Holotype, NU MR 0044 (a ; 11843, b ; 11844), b ;oblique basa l view. 2. Paratype, NU MR 0045 (12244). 3. Paraty pe, NU MR 0046 (12276) . 4a, b.
Paratype, NU MR 0047 (a; 95 - 22, b ; 95 - 23).

cular and closely spaced.

Remarks. - Minocapsa, n. gen . is di stinguished from Stichocapsa Haeckel by lacking
aperture. It also differs from Zhamoidellum
Dumitrica and Cryptamphorella Dumitrica
by not being cryptothoracic and by consistingof four or more segments rather than three.
Etymology. - The generic name is named
for the Mino Terrane which includes the typearea, N anjo Massif.
Minocapsa cylindrica Matsuoka, n. sp.
Figures 10- 1a - 5b
1987
1989

Bagotum sp. E, Hattori, pI. 15, fig. 4.
BagolUm sp. atf. B. modeslUm Pessagno andWhalen , Hattori and Sakamoto , pI. 13, fig . K.

Description. - Shell of five se gments,
ovoidal. Cephalis spherical; thorax andabdomen truncate conical. The proximal
three segments form a conical proximal part.
The distal two segments form a cylindrical to
ovoidal distal part. The last segment hemispherical without aperture. Segmental joints
generally indistinct externally; joint between
abdomen and fourth segment faintly marked
by change in contour from the conical proximal part to the cylindrical to ovoidal distal
part. Pores circular to subcircular, rather
densely spaced and set in tetragonal to hexagonal pore frames. Pores on the distal parttend to be arranged longitudinally. Poresaround distal end slightly larger than thoseon the rest of shell; these pores surroundedby circular rims. Small nodes or spines situ-

Atsushi Matsuoka

736

ated at the pore frame vertices.
Measurements (in ,um).-Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell.
Holotype max. mm. mean
181
(6)
165
193
193
TH
125
(6)
115
135
120
MW
Remarks. - This species differs from
Minocapsa globosa, n. sp. by consisting of fivesegments rather than four, by possessing amore slender distal part and by the pores ofthe distal part tending to be longitudinallyarranged.
Etymology. - The specific name comes fromthe Latin cylindricus-a-um (=cylindrical).
Type specimens. - Holotype NU MR 0039
(Figures 1O-1a, b); Paratypes NU MR 0040
(Figures 1O-2a, b), NU MR 0041 (Figure 103), NU MR 0042 (Figure 10-4), NU MR 0043
(Figures 1O-5a, b).
Type locality.-MNA-lO, Nanjo Massif,
Mino Terrane, central Japan.
Occurrence.-Nanjo Massif, Mino Terrane,
central Japan.
. Range. - Archicapsa pachyderm a Zone
(middle Lower Jurassic) to
Laxtorum (?)
jurassicum Zone (upper Lower Jurassic) as
far as known.

Minocapsa globosa Matsuoka, n. sp.
Figures ll-la-4b

Description. - Shell of four segments,
pyriform. Cephal is spherical; thorax andabdomen truncate conical. The proximal
three segments form a conical proximal part.
The fourth segment large, subspherical without aperture. Collar and lumber stricturesindistinct externally. Joint between abdomen and fourth segment marked by rapidchange in contour from the conical proximal
part to spherical distal part. Pores circular
to subcircular, densely spaced and set inpolygonal (largely hexagonal) pore frames.
Pores around distal end slightly larger thanthose on the rest of shell and surrounded byrims. Small nodes situated at the pore frame
vertices in some specimens.
Measurements (in ,um).-Numbers of specimens measured are in parentheses. TH;
total height of shell, MW; maximum widthof shell.
Holotype max. min. mean
177
(6)
170
185
TH
173
142
(6)
149
135
MW
140
Remarks. - This species is compared to
Minocapsa cylindrica, n. sp. under the latter
species.
Etymology. - The specific name is derived
from the Latin globosus-a-um (= spherical).
Type specimens. - Holotype NU MR 0044
(Figures ll-1a, b); Paratypes NU MR 0045
(Figure 11-2), NU MR 0046 (Figure 11-3),
NU MR 0047 (Figures 11-4a, b).
Type locality.-MNA-lO, Nanjo Massif,
Mino terrane, central Japan.
Occurrence. - N anjo Massif, Mino Terrane,
central Japan.
Range. - Archicapsa pachyderm a Zone
(middle Lower Jurassic) as far as known .

References
Baumgartner, P.O., 1984: A Middle Jurassic-Early
Cretaceous low-latitude radiolarian zonation
based on Unitary Associations and age of Tethyan radiolarites. Ec/ogae geol. Helv., vol. 77,
no. 3, p. 729-837, pis. 1-10.
Carter, E.S., Cameron, B.E B. and Smith P.L. 1988:
Lower and Middle Jurassic radiolarian biostratigraphy and systematic paleontology, Queen
Charlotte Islands, British Columbia. Geol.
Surv. Canada, Bull., 386, p. 1-109, pis. 1-18.
De Wever, P., 1981a: Hagiastridae, Patulibracchiidae
et Spongodicidae (Radiolaires Polycystines) duLias de Turquie. Rev. Micropa!eont., vol. 24,
no. I, p.27-50, pis. 1-7.
- - , 1981 b: Parasaturnalidae, Pantanellidae etSponguridae (Radiolaires Polycystines) du Liasde Turquie. Ibid., vol. 24, no. 3, p. 138-156, pis.
1-5.
- - , 1982: Nassellaria (Radiolaires Polycystines)
du Lias de Turquie. Ibid., vol. 24, no. 4, p. 189232, pis. 1-13.
Hattori, I., 1984: Alternating clastic limestone andred chert as olistolith in the Mino Terrane, cen-

915.

Jurassic radiolarians from the Nanjo Massif

tral Japan. Jour. Geol. Soc. Japan, vol. 90, no.
I, p.43-54, pis. 1-2.
- - , 1987: Jurassic radiolarian fossils from the
Nanjo Massif, Fukui Prefecture, central Japan.
Bull. Fukui Municipal Museum, Natural History, no. 34, p.29-10I, pis. 1-23. (in Japanesewith English abstract)- - , 1988: Radiolarian fossils from manganese nodules at the upper reach of the Tarumigawa in the
Nanjo Massif, Fukui Prefecture, central Japan,
and the tectonic significance of the northwestern
Mino Terrane. Ibid., no. 35, p. 55-10 I, pis. 114. (in Japanese with English abstract)- - , 1989: Jurassic radiolarians from manganese
nodules at three sites in the western Nanjo
Massif, Fukui Prefecture, central Japan. Mem.
Fac. Educ. Fukui Univ., no. 39, p. 47-134, pis. 145. (in Japanese with English abstract)- - and Sakamoto, N., 1989: Geology and Jurassic
radiolarians from manganese nodules of the
Kanmuriyama-Kanakusadake area in the Nanjo
Massif, Fukui Prefecture, central Japan. Bull.
Fukui Municipal Museum, Natural History, no.
36, p. 25-79, pis. 1-19. (in Japanese with English
abstract)- - and Yoshimura, M., 1982: Lithofacies distribution and radiolarian fossils in the Nanjo area in
Fukui Prefecture, central Japan. News ofOsaka Micropaleontologists, Spec. Vol., no. 5, p.
103-116, pis. 1-4. (in Japanese with English
abstract)- - and - - , 1983: Late Triassic to Middle Jurassic ages for greenstones within the Mesozoic
Nanjo Massif of the Mino Terrane, central Japan.
Mem. Fac. Educ. Fukui Univ., no. 32, p.67-80,
pis. 1-9. (in Japanese with English abstract)
Hori, R., 1988: Some characteristic radiolarians fromLower Jurassic bedded cherts of the Inuyama
area, Southwest Japan. Trans. Proc. Palaeont.
Soc. Japan, N.s., no. 151, p.543-563.
- - and Otsuka T., 1989: Early Jurassic radiolarians from the Mt. Norikuradake area, Mino terrane, central Japan. Jour. Geosci., Osaka City
Univ., vol. 32, p. 175-199, pis. 1-4.
- - and Yao, A., 1988: Parahsuum (Radiolaria)
from the Lower Jurassic of the Inuyama area,
central Japan. Ibid., vol. 31, p. 47-61, pis. 1-2.
Ichikawa, K. and Yao, A., 1976: Two new genera of
Mesozoic cyrtoid radiolarians from Japan. In,
Takayanagi, Y. and Saito, T. eds. Progress in
micropaleontology, p. 110-117, pis. 1-2, Amer.
Mus. Natural History, New York.
Isozaki, Y. and Matsuda, T., 1985: Early Jurassic
radiolarians from bedded chert in Kamiaso,
Mino Belt, central Japan. Earth Sci. (Chikyu
Kagaku), vol. 39, no. 6, p.429-442, pis. 1-3.
Ito, M. and Matsuda, T., 1980: Discovery of Triassic

737

and Jurassic radiolarians and Triassic conodontsfrom the Nanjo Mountains, western Mino Belt,
central Japan. Bull. Fukui Municipal Museum,
Natural History, no. 27, p. 7-12, pI. 1. (in Japanese)- - and Shiratake, T., 1980: A review of the Paleozoic radiolarian fossils found around Mt. Kanmuri on the Fukui-Gifu Prefectural border incentral Japan - New findings Triassic-Jurassic
period radiolarians -. Ibid., no. 27, p. 1-6, pI.
1. (in Japanese)
Kishida, Y. and Hisada, K., 1985: Late Triassic to
Early Jurassic radiolarian assemblages from the
Ueno-mura area, Kanto Mountains, central
Japan. Mem. Osaka Kyoiku Univ., Ser. 3, vol.
34, no. 2, p. 103-129, pis. 1-4.
Matsuoka, A. and Yao, A., 1986: A newly proposedradiolarian zonation for the Jurassic of Japan.
Marine Micropaleontology, vol. II, p. 91-106, pis.
1-3.
MacLeod, N., 1988: Lower and Middle Jurassic
Perispyridium (Radiolaria) from the Snowshoe
Formation, east-central Oregon. Micropaleontology, vol. 34, no. 4, p. 289-315, pis. 1-5.
Pessagno, E.A. Jr. and Blome, C.D., 1980: Upper
Triassic and Jurassic Pantanelliinae from California, Oregon, and British Columbia. Ibid., vol.
26, no. 3, p. 225-273, pis. I-II.
- - , - - , Carter, E.S., MacLeod, N., Whalen, P.A.
and Yeh, K.-Y., 1987: Preliminary radiolarian
zonation for the Jurassic of North America.
Part II of studies of North American Jurassic
Radiolaria. Cushman Found. Foram. Res.,
Spec. Pub., no. 23, p. 1-18.
- - and Poisson, A., 1981 : Lower Jurassic radiolaria from the Gumuslu Allochton of southwestern
Turkey (Taurides Occidentales). Bull. Mineral
Research and Exploration Institute of Turkey,
no. 92, 1979, p. 47-69, pis. 1-15.
- - and Whalen, P.A., 1982: Lower and Middle
Jurassic Radiolaria (multicyrtid Nassellariina)
from California, east-central Oregon and theQueen Charlotte Islands, B.c. Micropaleontology, vol. 28, no. 2, p. 111-169, pis. 1-13.
- - , - - and Yeh, K.-Y., 1986: Jurassic Nassellariina (Radiolaria) from North American Geologic Terranes. Bull. Am. Paleontology, vol. 91,
no. 326, p. 1- 75, pis. 1-11.
Sashida, K., 1988: Lower Jurassic multisegmented
N assellaria from the Itsukaichi area, western partof Tokyo Prefecture, central Japan. Sci. Rep.,
Inst. Geosci., Univ. Tsukuba, Sec. B, vol. 9, p. 127, pis. 1-4.
Takamura, Y. and Hayami, T., 1985: On the Paleozoic and Mesozoic strata in the eastern area of
Imajo-cho, Fukui Prefecture, central Japan.
Bull. Fukui Municipal Museum, Natural His-

Atsushi Matsuoka

738

tory, no. 31, p. 1-16, pis. 1-6. (in Japanese)
Takemura, A., 1986: Classification of Jurassic Nassellarians (Radiolaria). Palaeontographica,
Abt. A, Bd. 195, p.29-74, pis. 1-12.
- - and Nakaseko, K., 1982: Two new Jurassic
genera of Family Palaeoscenidiidae (Radiolaria).
Trans. Proc. Palaeont. Soc. Japan, N.S., no. 128,
p. 452-464, pis. 70-73.
- - and - - , 1983: Perseus, a new genus of Jurassic Palaeoscenidiidae and the phylogeny of Subfamily Hilarisirecinae. Mem. Fac. Sci., Kyoto
Univ., (Ser. Geol. Mineral.), vol. 49, nos. 1-2, p.
111-118, pis. 9-10.
- - and - - , 1986: The cephalic skeletal structureof Jurassic "Eucyrtidium" (Radiolaria). Jour.
Paleontology, vol. 60, no. 5, p. 10 16-1024.
Yao, A., 1972: Radiolarian fauna from the Mino Beltin the northern part of the Inuyama area, central
Japan. Part I: Spongosaturnalids. Jour. Geosci., Osaka City Univ., vol. 15, p. 21-64, pis. I-- - , 1979: Radiolarian fauna from the Mino Belt in
the northern part of the Inuyama area, central
Japan. Part II: Nassellaria I. Ibid., vol. 22, p.
21-72, pis. 1-12.
- - , 1982: Middle Triassic to Early Jurassic
radiolarians from the Inuyama area, central
Japan. Ibid., vol. 25, p. 53-70, pis. 1-4.
Yeh, K.-Y., 1987a: Taxonomic studies of Lower
Jurassic radiolaria from east-central Oregon.
Nat. Museum Nat. Science, Spec. Pub., no. 2, p.
1-169, pis. 1-30.
- - , 1987b: A revised classification for Family
Canoptidae (Radiolaria). Mem. Geol. Soc.
China, no. 8, p.63-72, pis. 1-2.
Yoshimura, M., Kido, S. and Hattori, I., 1982:
Stylolitic cherts and radiolarian fossils in the
Imajo area of the Nanjo Massif, Fukui Prefecture, central Japan. Bull. Fukui Municipal
Museum, Natural History, no. 31, p.65-77, pis.
1-6. (in Japanese with English abstract)

II.

Imajo 4'Et., Kasugano ~ B!I!f, Kohkura ii1ii~, Nanjo m~.
~IHlrm~Wi1!!tJ'I?O)iitrM'; '" '7*cntltlc;e. (.:t-O) I) Tricolocapsa, Stichocapsa, Minocapsa, n. gen.: ~111Hlri¥i~Wi1!!0)"";/ jf;/ /~;/ r-tJ'I?~I?.ht.:iitrJtlj,; '" '7*cnti!c;e.~~O)~
I $fi c L. "'C, ~JmO) Nassellaria3 ~ (Minocapsa ft~~) ~;:: "?\, ''''C, J;), "f 0) 9 t1T~ (2 t1Tl!Ii:~~~tr) ~-$'~, ~~ L. t.:o T. minoensis, T. (?) megaglobosa, T. (?) fera, S. plicata plicata,
S. plicata semiplicata, S. elegans, S. biconsica, S. nanjoensis, M. cylindrica, M. globosa. .:0) "" ;/ 71';/ /~;/ r tJ, 1?il:1±l L. t.:ntltlc;e.~~Hl, btJ~OO-e~li ~.ht.:,; '" '7 re1~E*0) ? 't;,Archicapsa pachyderma*l::$o)~~f;::mfbl"t7.Jo it.:, ':o)~~ft, ~t7;1 !i71O):;tv::{;/ ~'Ii 9='*$0) ~ 7 Iv "/ 7 ;/1li"f$tJ, 1?il:1±l"t 7.J ntltlc;e.~~~;::.Jttt~.h 7.J o~fIlil ~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 161, pp. 739-750,4 Figs., April 30, 1991

916. PAMIRINA (PERMIAN FUSULINACEA) FROM THE
AKIYOSHI LIMESTONE GROUP, SOUTHWEST JAPAN*
KA TSUMI UENO
Division of Environmental Science, Graduate Schoolof Science and Technology,
Chiba University, Chiba 260

Abstract. The genus Pamirina is considered as the direct ancestor of verbeekinid andneoschwagerinid fusulinaceans, and two distinct groups of Pamirina which have different
spirothecal structure each other are so far known in this genus. The different stratigraphic
distribution of two groups of Pamirina in the Akiyoshi Limestone Group distributed in the
Kaerimizu area, the northeastern part of Akiyoshi Plateau, supports to subdivide the genus
into two subgenera. In this paper, Levenia, n. subgen. is proposed and three species ofPamirina; P. (P.) darvasica, P. (Levenia) leveni, P. (L.) evaluta, and Misellina (Brevaxina) ? sp. coexisting with P. (P.) darvasica from the Pamirina (Levenia) leveni Zone aredescribed and illustrated. This Pamirina fauna is referable to the late Artinskian or the late
Yakhtashian age.
Key words.
Yakhtashian.

Akiyoshi Limestone Group, Artinskian, Kaerimizu, Levenia, Pamirina,

the middle volutions, and tectum and finealveolar keriotheca in the outer volutions.
The other one is represented by Pamirina
tethydis and P. leveni, and has a thickly
lenticular shell. Their spirotheca is composed of a structureless single layer in the
inner volutions, and tectum and pro theca in
the outer ones. Kobayashi (1977) considered that the latter group is more primitive
than the former one and recognized the following four evolutionary stages from thegenus Pamirina to the most primitive memberof the family Verbeekinidae, the genus Misellina, through the detailed observation of
ontogenic change of spirothecal structure andgrowth curve of Pamirina and also certain
species of Misellina. Namely, primitive
Pamirina (P. tethydis and P. leveni),
advanced Pamirina (P. darvasica), primitive
Misellina (M. dyhrenfurthi - M. otakiensis
group, which are now referred to the subgenus Brevaxina of the genus Misellina), andIntroduction

The family Staffellidae had been long
regarded as an ancestral stock of the family
Verbeekinidae until Leven (1970) establishedthe genus Pamirina with Pamirina darvasica
as the type species. He explained that thegenus is the direct ancestor of the verbeekinid
fusulinaceans, and the family Verbeekinidae
is derived from the family Ozawainellidae.
Later, Kobayashi (1977) described three
species of Pamirina from the Itsukaichi,
Shomaru and Otaki areas in the Kwanto
Mountains and confirmed the presence of two
distinct groups of Pamirina having different
spirothecal structure. One group includes
Pamirina darvasica, the type species of thisgenus, and has a subspherical shell and spirotheca composed of a single structure less layer
in the inner volutions, tectum and pro theca in'Received September 4, 1990; revised manuscript
accepted February 25, 1991
739

Katsumi Ueno

740

advanced Misellina (M. claudiae etc.). He
concluded that these four evolutionary stages
from Pamirina to Misellina construct thesingle evolutionary series. All of the materials treated in Kobayashi (1977), however,were obtained from limestone blocks or limestone pebbles in the middle and southernbelts of the Chichibu Terrain, which are now
thought to have comprised accretionary complexes during Mesozoic time (Ozawa andKo bayashi, 1985 and others). Therefore,
their phylogeny based on stratigraphic evidence is still incomplete.
In the course of the reexamination of thefusulinacean biostratigraphy of the Akiyoshi
Limestone Group in the Kaerimizu area, thenortheastern part of Akiyoshi Plateau, I
found many individuals of Pamirina from thePseudofusulina ambigua Zone by Ota (1977).
In this paper, Levenia, n. subgen. is introduced for the group of Pamirina without afine alveolar keriotheca in the spirotheca, andthree species of Pamirina besides Misellina(Brevaxina)? sp. coexisting with Pamirina(P.) darvasica, which is considered to be thetransitional form between Pamirina andBrevaxina, are described and illustrated.
The fusulinacean biostratigraphy and stratigraphic distribution of Pamirina in the
Akiyoshi Limestone Group are also briefly
noted.
All of the illustrated specimens are kept rn
the Department of Earth Sciences, Faculty of
Science, Chiba University.
Geologic setting and fusulinacean
biostra tigraphy
The Kaerimizu area is situated in the northeastern part of Akiyoshi Plateau (Figure I).
The Akiyoshi Limestone Group ranging fromthe late Middle Carboniferous to the late
Middle Permian which contains rich shallow~

AKIYOSHI-OAI

KAMA

N

o

AKIYOSHI-DA

2KIII
I

Figure 1. Map showing the Kaerimizu areain the Akiyoshi Plateau.

marine organic remains such as foraminifers,
algae, corals and mollusks is well exposed inthis area. It has been investigated by many
students in the focus of geologic structure andfusulinacean biostratigraphy (Ozawa, 1923;
T oriyama, 1954, 58; Hasegawa, 1958, 63; Sakaguchi et al., 1966; Ota et at., 1973; Ota,
1977). Ota (1977) divided the Akiyoshi
Limestone Group in this area into the following 10 fusulinacean zones in ascending order:
the Fusulinella biconica, Triticites simplex,
Pseudofusulina vulgaris, Pseudofusulina ambigua, Misellina claudiae, Parafusulina kaerimizensis, Afghanella schencki, Neoschwagerina craticulifera, Verbeekina verbeeki andColania douvillei Zones.
My recent investigation in the Kaerimizu
area makes it possible to divide Ota's
Pseudofusulina ambigua Zone into the lower
Pseudofusulina ex gr. kraffti and upper Pamirina (Levenia) leveni Zones. The distribu-

Figure 2. Distribution of biozones and Pamirina localities studied herein. 1: Chalaroschwagerinavulgaris Zone, 2: Pseudofusulina ex gr. kraffli Zone, 3: Pamirina (Levenia) leveni Zone, 4: Misellina(Brevaxina) dyhrenfurlhi olai Zone,S: Misellina (Misellina) claudiae Zone (M: Misellina (M.) claudiae
swarmed layer), 6: Parafusulina kaerimizensis Zone, 7: fossil locality.

916.

Pamirina from Akiyoshi

741

Figure 2Ao,(

MANAGATAKE~

Legend

&1
2

.4
L2J3
~5

o

Figure 2B

Em6
100m

'• _c::=::=J1

•

7

742

Katsumi Uenotion of biozones in the Kaerimizu area IS
shown in Figure 2.
The Pamirina (Levenia) leveni Zone isdefined as the stratigraphic interval from thefirst appearance of Pamirina (Levenia) leveni
and P. (L.) evoluta to that of Misellina(Brevaxina) dyhrenfurthi otai. Algal oncoidal limestone is predominant in this zone
with a subordinate amount of bioclastic sparitic limestone and muddy limestone. The
following fusulinaceans are discriminated:
Pamirina (P.) darvasica Leven, P. (Levenia)
leveni Kobayashi, P. (L.) evoluta Sheng andSun, Toriyamaia laxiseptata Kanmera, Minojapanella (M.) cf. elongata Fujimoto andKanuma, Schubertella spp., Pseudoreichelina
darvasica Leven, P. spp., Pseudoendothyra
obiniouensis Leven, P. spp., Pseudofusulina
ex gr. kraffti (Schellwien), P. fusiformis(Schellwien), P. spp., Chalaroschwagerinavulgaris (Schellwien), Misellina (Brevaxina)
aff. otakiensis (H uzimoto) and M. (B.)? sp.
Accordingly, its geologic age is referable tothe late Artinskian or the late Yakhtashian.
Concerning the stratigraphic distributionof Pamirina in the Akiyoshi LimestoneGroup, Pamirina (Levenia) leveni and P. (L.)
evoluta appear first in the basal part of the
Pamirina (Levenia) leveni Zone and range up
to the middle part of the zone. While Pamirina (P.) darvasica is obtained from the upper
part of the Pamirina (Levenia) leveni Zone
associated with Misellina (Brevaxina)? sp.
From immediately above this horizon, Misellina (Brevaxina) aff. otakiensis is detected
and this horizon is succeeded by the Misellina(Brevaxina) dyhrenfurthi otai Zone througha short stratigraphic interval. In the Akiyoshi Limestone Group, Levenia occupies
slightly lower stratigraphic position than thatof Pamirina (s.s.). This fact is concordant
with Kobayashi (1977)'s conclusion on thephylogenetic relationship between Pamirina(Levenia) leveni and P. (P.) darvasica.
In the Permian Period of the Tethyan
region, eight zones by the fusulinacean genera
can be recognized. They are Pseudoschwagerina, Pseudofusulina-Chalarosch wagerina, Misellina, Cancellina-Maklaya, Neosch wagerina, Lepidolina- Yabeina, Codonofusiella-Reichelina and Palaeofusulina Zones
in ascending order. Judging from the known
occurrence of Pamirina together with its
stratigraphic distribution in the Akiyoshi
Limestone Group, the genus Pamirina is mostcharacteristic in the upper part of
Pseudofusulina-Chalarosch wagerina Zone,
which indicates the late Artinskian or the late
Yakhtashian age, although some forms arefound together with Misellina. Moreover, as
far as the present knowledge of its geographic
distribution is concerned, it seems that Pamirina is paleobiogeographically restricted to the
Tethyan Realm.
Systematic paleontology
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea von Moller, 1878
Family Ozawainellidae Thompson
and Foster, 1937
Subfamily Ozawainellinae Thompson
and Foster, 1937
Genus Pamirina Leven, 1970 emend.
Kobayashi, 1977

Type species. - Pamirina darvasica Leven,1970.
Pamirina Leven, 1970, p. 23; Rozovskaya, 1975, p.

~
Figure 3. 1-7. Pamirina (Pamirina) darvasica Leven, 1,2: axial sections, GK6-18, SRI2-2, 3a,4,
5: tangential sections: SRI2-l2, GK6-24, GK6-10, 6,7: sagittal sections: GK6-7, GK6-1, 3b: enlarged
part of 3a. 8-23. Pamirina (Levenia) leveni Kobayashi, 8, 9a. 10-16: axial sections, T A 122-14, T A 122-5,
TAI22-la, KS94-1, TAI22-29, TAI22-32, TAI22-24a, TAI21-2, TAI22-25b, 17-20: tangential sections,
TAI22-25a, TAI22-1b, TAI22-7, TAI22-24b, 21-23: sagittal sections, TAI22-40, TAI22-47, TAI22-A,
9b: enlarged part of 9a. 24. Misellina (Brevaxina) ? sp., 24a: axial section, GK6-11, 24b: enlarged photograph of 24a. All X 40, exept for 3b, 9b and 24b; X 100.

916

Pamirina from Akiyoshi

743

3b

12

~
--.9(a ~"" ' ,

R~'C'i2016

Katsumi Veno

744

114- 115; Lin el al. , 1977, p. 19- 20; Kobayashi ,1977, p. 10- 11 ; Liu et al., 1978, p. 80; Sun elaI., 1983, p. 9; Loeblich and Tappan, 1988, p.
286; Sheng el al., 1988, p. 175.
Chinlingella Wang and Sun, 1973, p.152, 171-172;
Sheng et al., 1988, p. 176.

Diagnosis. - Shell small , thickly lenticular
to subspherical , rarely spherical with broadly
rounded periphery. Axial regions slightly tomoderately umbilicated. Mature shell usually having 3 to 6 volutions. Axis of coilingusually straight throughout except forjuvenarium of some forms. Shell expands
gradually. Proloculus spherical, small tolarge for shell size. Spiro theca thin and 2
types in structure. Spirotheca of primitive
forms composed of structureless single layer
in inner volutions, and tectum and lower lessdense layer (protheca) in outer ones. Besides
outer volutions of advanced forms have spirotheca, consisting of tectum and fine alveolar
keriotheca . Septa straight and plane. Tips
of septa swollen in sagittal section. Chomata weak and low, sometimes rudimental.
Tunnel path low and broad.
Remarks. - The original diagnosis ofPamirina by Leven (1970) is summarized asfollows. "Shell small, subspherical, more orless depressed in axial regions. Coiling axis
in inner few volutions unstable. Rate ofshell expansion uniform. Proloculus spherical and small. Spiro theca with fine alveolar
keriotheca. Aperture single and chomata
distinct. "As stated by Lin et al. (1977), the gen us
Chinlingella Wang and Sun is quite identicalwith Pamirina and is a junior synonym of thelatter.
The genus Pamirina can be divided into thefollowing two subgenera, Pamirina andLevenia, n. subgen., by the difference in general shell shape, the spirothecal structure andstratigraphic occurrence.
Subgenus Pamirina Leven , 1970

Type species. - Pamirina darvasica Leven,1970.

Diagnosis . - Shell small , subspherical torarely spherical with broadly rounded periphery and slightly umbilicated axial regions .
Mature shell consists of 5 to 6 volutions.
Axis of coiling usually straight throughout
except for juvenarium of some forms.
Prolocul us small and spherical. Spirothecathin and composed of single structurelesslayer in inner volutions, tectum and lower lessdense layer (protheca) in middle ones, andtectum and fine alveolar keriotheca in outer 1
or 2 volutions. Septa straight and plane.
Tips of septa slightly swollen in sagittal section . Chomata low and broad. Tunnel
path regular and wide.
Remarks. - The following species are included in this subgenus, Pamirina darvasica
Leven , 1970 (= Chinlingella chinlingensis
Wang and Sun, 1973; C. nobilis Wang and
Sun , 1973); P. pulchra Wang and Sun, 1973 ;P. orbiculoidea Sheng and Sun, 1975; P.
staffellaeformis Zhou, Sheng and Wang, 1987.
Geologic age. - Late Artinskian or late
Yakhtashian (Early Permian).
Geographic distribution. - Carnic Alps (Kahler and Kahler, 1980). Darvas Range(Leven, 1970), North Thailand (Sakagami
and Iwai , 1974), Chinling Range (Wang andSun, 1973), Guizhou (Liu et al., 1978),
Guangxi (Lin et al., 1977), Qinghai (Shengand Sun, 1975), Yunnan (Zhou et al ., 1987),
Xinjiang (Sun and Zhang, 1988), Kwanto
Mountains (Kobayashi , 1977) and Akiyoshi(present study).
Pamirina (Pamirina) darvasica
Leven, 1970
Figures 3- 1 - 7

Pamirina darvasica Leven, 1970, p. 23-24, pI. 1, figs.
1- 9, 23-24 (not pI. 1, figs. 10-11); Rozovskaya,
1975, pI. 35 , figs. 4-5 (same as pI. 1, figs. 1- 2 ofLeven , 1970, p. 23-24) ; Kobayashi , 1977, p. 1415, pI. 2. figs. la, Ib, 2- 3; Liu el al., 1978, p.
80, pI. 18, figs. 8, 12; Kahler and Kahler, 1980, p.
187- 188, pI. 3, fig. 8 (not pI. 3, fig. 7) ; Loeblich916.

Pamirina from Akiyoshi

a nd Tappan , 1988, pI. 295, fig. II (not pI. 295, fig.
12, same as pI. I, fig. I of Leven, 1970, p. 23 - 24).
Chinlingella chinlingensis Wang and Sun , 1973, p.
152- 153, 172, pI. I, figs. 12, 17-32, pI. 3, figs. I, 5,
10.
Chinlingella nobilis Wang and Sun , 1973, p. 153- 154,
173, pI. I, figs. 1- 5, 9-11.
Staffella aff. mo1teri Ozawa, Sakagami and Iwai ,1974, p. 54-55, pI. 4, figs. 1- 13.
Pamirina nobitis (Wang and Sun) , Lin et at., 1977, p.
20, pI. 3, figs. 21 - 22; Sun e/ at., 1983, p. 9, pI. 2,
fig. 15 (same as pI. I, fig. I of W ang a nd Sun ,1973, p. 153- 154, 173) ; Zhou et at., 1987, pI. 2,
figs. 3, 4.
Pamirina chintingensis (Wang and Sun), Sun el at. ,1983, p.9, pI. 2, fig. 17 (same as pI. I, fig. 19 ofWang a nd Sun, 1973, p. 152- 153, 172) ; Zhou elat., 1987, pI. 2, figs. 1, 2; Loeblich and Tappan,
1988, pI. 295, figs. 8- 10 (same as pI. I, figs. 19- 20,
31 of Wang and Sun, 1973, p.152 - 153, 172) ;Sun and Zhang, 1988, pI. 3, figs. 7, 15.

Material studied. - Axial sections; GK618 from Loc. GK6 and SR 12-2 from Loc.
SRI2. Sagittal sections; GK6-1 and GK6 7 from Loc. GK6. Tangential sect ions ;GK6- 10 a nd GK6-24 from Loc. GK6 andSR12- 12 from Loc. SRI2. Additional specimens were obtained from Locs. GK6 and
SRI2 .
Description. - Shell small, involute throughout and subspherical with broadly rounded
periphery. Axial regions slightly depressed .
Mature specimens having 5 to 5 Yz volutions
0.53 to 0.63 mm in axial length and 0.77 to
0.92 mm in median width, giving form ratio
of about 0.70.
Axis of coiling slightly rotated in juve narlum. Shell expands gradually throughgrowth. Average radiu s vectors of the first to
fifth vol utions for 4 specimens 0.05 , 0.10, 0.17 ,0.28 and 0.43 mm . Average form ratio s ofthe first to fifth volutions for 2 specimens 0.70,
0.72,0.81,0.76 and 0.70, respectively.
Proloculus small and sp herical. Its outside diameter about 0.05 mm .
Spirotheca thin a nd composed of single
structureless layer in inner I or 2 volutions,tectum a nd lower less den se layer (protheca)in the next 2 volutions, and tectum and fine745

alveolar keriotheca in outer or 2 volutions.
Average thickness of spirotheca of the first to
fifth volutions for 4 specimens 0.009, 0.011 ,0.0 IS, 0.020 and 0.022 mm.
Septa straight and plane. Tips of septa in
sag ittal sectio n swollen. Septal counts of thefirst to fifth volutions of specimen illustrated
In Figure 3- 6; 6, II , 13 , 14 and 14.
Chomata low , broad and well developed inouter vol utions. Tunnel path wide and regular.
Remarks. - The Akiyoshi specimen s a requite similar to the D arvas ones in the essential morphological characters.
Pamirina (P .) darvasica somewhat resembles P. (P.) eostaffellaeformis originally descri bed by Zhou et al. (1987) from the Pamirina chin/ingensis and Misellina ova/is Zonesof eastern Yunn a n. However, the former ha sa smaller shell than the latter.
Staffella afr. molleri Ozawa described by
Sakagami a nd Iwai (1974) from the Ph a Duk
Chik Limesto ne and Huai Hin Lat limestone
conglomerate of North Thailand has a subsp herical shell and its spirotheca composed ofa tectum a nd fine a lveolar keriotheca , a nd
coexists with typical Artinskian fusulinaceanssuch as Cha/arosch wagerina vulgaris, Pseudo-

fusulina krotowi, Mesoschubertella thompsoni
and others. Judging from the above
mentioned facts , the Thailand specimens areapparently assigned to the genus Pamirina
and identical with P. darvasica.
Occurrence. - Common in the upper part of
the Pamirina (Levenia) /eveni Zone.
Subgenus Levenia Ueno, n. subgen .

Type species. - Pamirina le veni Kobayashi ,1977.

Diagnosis. - Shell small and thickly
lenticular with broadly rounded periphery.
Axial regions slightly to deeply depressed.
Mature shell consists of 3 to 5 Yz volutions.
Coiling planispiral throughout, but slightly
rotated in juvenarium of some forms. Shell
expands gradually. Proloculu s spherical

746

Katsumi Venoand sm a ll to large for shell size. Spirothecathin , composed of single structureless layer ininner volutions, but tectum and lower lessdense layer (protheca) in outer ones. Septa
straight and plane. Tips of septa slightly
swollen in sagittal section. Chomata hardly
observed or rudimentarily developed . Tunnel path low and regular.
Remarks. - The following five species canbe referred to the subgenus Levenia, Pamirina
tamanouchiensis (Sakagami , 1956) ; P.
evoluta Sheng and Sun , 1975 ; P. leveni
Kobayashi , 1977; P. tethydis Kobayashi ,1977 ; P. ? hataii (Suyari , 1962) .
Thi s new subgenus differs from Pamirina(s.s.) in having a thickly lenticular shell,smaller form ratio and different spirothecal
structure. Moreover the former occupies
slightly lower stratigraphic position and isconsidered the direct ancestor of the latter as
Kobayashi (1977) pointed out.
Geologic age. - Late Artin skian or late
Yakhtashian (Early Permi a n).
Geographic distribution. -Qinghai (Shengand Sun, 1975), Kwanto Mounta ins (Sa-kagami , 1956 ; Takaoka, 1966 ; Kobayashi ,1977), Mino Mountains (Kanuma, 1960),
Kochi (Suyari , 1962) and Akiyoshi (present
study) .

Pamirina (Levenia) leveni
Kobayashi , 1977
Figures 3- 8 - 23

Pamirina pulchra (Wang and Sun), Sheng and Sun ,1975, p. 46, pI. I, fig s. 16- 18.
Pamirina leveni Ko bayashi , 1977, p. 11 - 14, pI. I, fi gs.
13- 38.

Mat erial studied. - Axial section s;
TA121 - 2 from Loc. TA121 , TAI22 - la ,TAI22- 5, TAI22- 14, T A 122-24a, T A 12225 a, TAI22 - 25b, TAI22-29, TAI22 - 32 from
Loc. T A 122 and KS94- 1 from Loc. KS94.
Sagittal sections; T A 122- 40, T A 122- 47 andTAI22- A from Loc. TA122. Tangential sections ; TAI22- lb, TAI22- 7 and TA122 - 24bfrom Loc. T A 122. Additional specimen swere a lso obtained from some localiti es in the
Pamirina (Levenia) leveni Zone.

Figure 4. 1- 8. Pamirina (L evenia) evaluta Shen g and Sun , 1- 5, 8 : ax ial sections, KS46 - 1, NS77- 2a,
TA I 16- 2, NS77- 2b, NS75 - 1, NS98- 2, 6,7 : sagittal sectio ns, KS79- 2, NSIOO- l. 3a,6a, X 40, 1,2, 3b, 4, 5,
6b- 8, X 100.

916.

Pamirina from Akiyoshi

Description. - Shell small, involute throughout and thickly lenticular with broadly
rounded periphery. Axial regions almost
straight to slightly depressed. Mature specimens of 5 volutions 0.31 to 0.43 mm in axial
length and 0.69 to 0.895 mm in median width,
giving form ratios of 0.40 to 0.57.
Axis of coiling straight throughout but
slightly rotated in juvenarium of some specimens. Shell expands gradually throughgrowth. Average radius vectors of the first to
fifth volutions for II specimens 0.05, 0.09,
0.15,0.26 and 0.41 mm. Average form ratiosof the first to fifth volutions for 9 specimens
0.69, 0.58, 0.51, 0.48 and 0.42, respectively.
Proloculus small and spherical. Its outside diameter ranges from 0.04 to 0.06 mm,
averaging 0.05 mm for II specimens.
Spirotheca thin and composed of single
structureless layer in inner few volutions, andtectum and lower less dense layer (protheca)in outer ones. Average thickness of spirotheca of the first to fifth volutions for 12
specimens 0.006,0.009,0.013,0.016 and 0.017
mm.
Septa straight and plane. Tips of septa
swollen in sagittal section. Septal counts ofthe first to fifth volutions of illustrated specimen (Figure 3-23) 7?, 10, 12, 12 and 13.
Chomata inconspicuous or rudimentarily
developed. Tunnel path low and broad.
Remarks. - The Akiyoshi specimens arequite referable to the original ones describedby Kobayashi (1977) in the· essential morphological characters, except for having aslightly larger shell in the former.
Sheng and Sun (1975) described Pamirina
pulchra (Wang and Sun) from the Qinghai
district of China. According to theirdescription and illustration, it has a thickly
lenticular shell and two layered spirothecacomposed of a tectum and lower homogeneous tectorium. This spirothecal structure is
identical with that in the subgenus Levenia
rather than Pamirina (s.s.). Judging fromthe shell shape and spirothecal structure, thisspecies is referable to Pamirina (Levenia)

747

leveni Kobayashi.
Occurrence.-Abundant in the lower andmiddle parts of the Pamirina (Levenia) leveni
Zone.
Pamirina (Levenia) evoluta
Sheng and Sun, 1975
Figures 4-1 - 8

Pamirina? evo/uta Sheng and Sun, 1975, p. 46, pl. 1,
fig. 15.

Material studied.-Axial sections; KS46I from Loc. KS46, NS75-1 from Loc. NS75,
NS77-2a and NS77-2b from Loc. NS77,
NS98-2 from Loc. NS98 and T A 116-2 from
TA116. Sagittal sections; KS79-2 from
Loc. KS79 and NS 100-1 from Loc. NS 100.
Description. - Shell small and thickly
lenticular with broadly rounded periphery.
Mature specimen having 3 ~ volutions (Figure 4-3) 0.21 mm in axial length and 0.46 mm
in median width, giving a form ratio of 0.46.
Specimens of 2 ~ volutions 0.13 to 0.19 mm
in axial length and 0.225 to 0.38 mm inmedian width, giving form ratios of 0.41 to
0.58.
Coiling planispiral and involute in inner
2~ volutions but evolute in last one. Shell
expands gradually through growth. Radius
vectors of the first to third volutions ofmature individual 0.08, 0.13 and 0.21 mm,
and form ratios 0.69, 0.62 and 0.57, respectively.
Proloculus spherical and large for genus.
Its outside diameter ranges from 0.07 to 0.125mm, averaging 0.09 mm for 9 specimens.
Spirotheca thin and composed of single
homogeneous layer in inner volutions, but
tectum and thin lower less dense layer (protheca) in outer ones. Thickness of spirotheca of the first to third volutions of typical
specimen (Figure 4-3) 0.01, 0.015 and 0.Dl5mm.
Septa straight and plane. Septal counts ofthe first and second volutions of specimen

Katsumi Vena748

illustrated in Figure 4-6; 7 and 12.
Chomata present but inconspicuous.
Remarks. - This species is characterized byan evolute shell and umbilicated axial regionsin the mature stage and larger proloculus forthe genus. It was originally decribed by
Sheng and Sun (1975) from the Chihsian of
the Qinghai district in China. In the Akiyoshi specimens, deep umbilicuses are observedin only one axial section (Figure 4-3). The
rest is considered as immature individuals of
this species.
Occurrence. - Common in the lower andmiddle parts of the Pamirina (Levenia) leveni
Zone.
Family Verbeekinidae Staff
and Wedekind, 1910
Subfamily Misellininae
Miklukho-Maklay, 1958
Genus Misellina Schenck
and Thompson, 1940
Subgenus Brevaxina Schenck
and Thompson, 1940
Misellina (Brevaxina)? sp.
Figure 3-24

layer in inner 2 volutions, tectum and lower
less dense layer (protheca) in the third one,and tectum and fine alveolar keriotheca inouter 2 volutions. Thickness of spirothecaof the first to fifth volutions 0.005, 0.01, 0.02,
0.02 and 0.02 mm. Septa not fluted. Rudimentary parachomata developed in the thirdand fourth volutions.
Remarks. - General shell shape and the
spirothecal structure of the present specimen
are similar to those of Pamirina (P.) darvasica, but the presence of rudimentary parachomata in the present specimen indicates theintimate relationship to the subgenus Brevaxina of the genus Misellina. The present specimen is seemingly intermediate between
Pamirina and Brevaxina. In this paper, it istentatively assigned to Brevaxina.
Leven (1970) illustrated two specimens ofPamirina darvasica with rudimentarily developed parachomata in the outer volutions(Leven, 1970, pI. 1, figs. 11-12). The Akiyoshi specimen is quite referable to those
Leven's ones.
Occurrence. - Rare in the upper part of the
Pamirina (Levenia) leveni Zone and coexisting with P. (P.) darvasica.

ComparePamirina darvasica Leven, 1970, p. 23-24, pI. I, figs.
11-12 (not pI. I, figs. 1-10,23-24); Loeblich andTappan, 1988, pI. 295, fig. 12 (not pI. 295, fig. II,
same as pI. I, fig. II of Leven, 1970, p.23-24).

Material studied. - Axial section; GK6-11
from Loc. GK6.
Description. - Shell small, subsphericalwith slightly depressed axial regions. Axial
length 0.52 mm and median width 0.66 mm,
giving a form ratio of 0.79. Shell consists of
5 volutions and inner 1 Yz volutions skew
coiled but remains planispirally coiled.
Radius vectors of the first to fifth volutions
0.065, 0.11, 0.17, 0.25 and 0.34 mm, and form
ratios 0.77, 0.82, 0.71, 0.74 and 0.74, respectively. Proloculus spherical and measures
0.05 mm in outside diameter. Spirothecathin and composed of single structure lessAcknowledgments
I would like to express my appreciation toProf. Sumio Sakagami of Chiba University
for reading of the manuscript. Thanks arealso extended to Messrs. Akihiro Sugimura
and Takehiko Haikawa of Akiyoshi-dai
Museum of Natural History for their continuous advice and encouragement, to Mr.
Yasuyuki Miyama of Chiba University forthe useful discussion on the fusulinaceantaxonomy, and to Prof. Hisayoshi Igo of the
University of Tsukuba for his kind loan of
the necessary literatures.
References cited
Hasegawa, Y., 1958: Note on the geological structureof the Akiyoshi Plateau, S.W. Honshu. Earth916.

Pamirina from Akiyoshi

Science (Chikyu Kagaku), no. 39, p. 15-18. (inJapanese with English abstract)- - , 1963: New find of fossils in the reddish
tuffaceous shale in the Akiyoshi Province. Ibid.,no. 64, p. 32-36.
Kahler, F. and Kahler, G., 1980: Fusuliniden aus
den Kalken der Trogkofel-Schichten der Karnischen Alpen. In, Fliigel, E. ed., Die TrogkofelStufe im Unterperm der Karnischen Alpen.
Sonderhefte der Carinthia II, 36, p. 183-254, pis.
1-9.
Kanuma, M., 1960: Stratigraphical and paleontological studies of the southern part of the Hida
Plateau and north-eastern part of the Mino
Mountainland, central Japan, Part 2. Paleontology, No.4, Ozawainellinae, Schubertellinae, Verbeekininae and Neoschwagerininae.
Bull. Tokyo Gakugei Univ., vo!' II, p. 55-73, pis.
10-13.
Kobayashi, F., 1977: Some considerations on theancestor of the family Verbeekinidae (Fusulinacea). Trans. Proc. Palaeont. Soc. Japan,
N.S., no. 105, p. 1-16, pis. 1-2.
Leven, E. Ya., 1970: 0 proiskhozhdenii vysshikh
fuzulinid [On the origin of the advanced
fusulinids]. Paleont. Zhurnal., 1970, no. 3, p.
18-25, p!. I. (in Russian)Lin, J.X., Li, J.X., Chen, G.X., Zhou, Z.R. andZhang, B.F., 1977: Fusulinida. In, Hubei Inst.,
Geo!. Sci. et al. eds., Pa laeontological Atlasof Central South China, no. 2 (Late Paleozoic),
p.4-96, pis. 1-30. Geo!. Pub!. House, Beijing.
(in Chinese)Liu, Z.A., Xiao, X.M. and Dong, W.L., 1978: Protozoa. In, Working Group of Stratigraphy andPaleontology of Guizhou ed., Palaeontological
A tlas of Southwest China, Guizhou vo!., no.2
(Carboniferous to Quarternary), p. 12-98, pis. 124. Ibid. (in Chinese)
Loeblich, A.R., Jr. and Tappan, H., 1988: Foraminiferal genera and their classification. 970
p., 847 pis. Van Nostrand Reinhold Comp., NewYork.
Ota, M., 1977: Geological studies of Akiyoshi, Part I.
General geology of the Akiyoshi LimestoneGroup. Bull. Akiyoshi-dai Mus. Nat. Hist. , no.
12, p. 1-33, pis. 1-3.
- - , Toriyama, R., Sugimura, A. and Haikawa, T.,
1973: Restudy on the geologic structure of the
Akiyoshi Limestone Group, Southwest Japan.
Jour. Geogr., vo!. 82, no. 3 (795), p. 115-135. (inJapanese with English abstract)
Ozawa, T. and Kobayashi, F., 1985: Stratigraphy
and geological structure of the Paleozoic andMesozoic strata in the southern part of the
Kwanto Mountains, central Japan. Hyogo
Univ., Teacher Educ. Jour., Ser. 3 (Natural

749

Science and Practical Life Studies), vo!. 6, p.
103-141, p!. I. (in Japanese with English
abstract)
Ozawa, Y., 1923: Stratigraphical study on the socalled Upper Chichibu Paleozoic System including the Akiyoshi-dai Limestone. Jour. Geol.
Soc. Japan, vo!' 30, no. 357, p.222-243. (inJapanese)
Rozovskaya, S.E., 1975: Sostav, sistema i filogeniya
otryada fuzulinida [Composition, phylogeny andsystem of the order FusulinidaJ. A kad. Nauk
SSSR, Trudy Paleont Ins!., vo!. 149, p.I-267,
pis. 1-35. (in Russian)Sakagami, S., 1956: Fusulinids from the limestone
conglomerate in the Yagooki valley, Tamanouchi, Hinode-mura, Nishitama-gun, Tokyo-to,
Japan. Trans. Proc. Palaeont. Soc. Japan, N.S.,
no. 24, p. 259-265, p!. 37.
- - and Iwai, J., 1974: Fusulinacean fossils fromThailand, Part VIII. Permian fusulinaceansfrom the Pha Duk Chik Limestone and in thelimestone conglomerate in its environs, North
Thailand. Geol. Palaeont. Southeast Asia, vo!.
14, p. 49-81, pis. 4-9.
Sakaguchi, S., Sugano, K., Nakamura, T. andKunugi, K., 1966: A study on the biostratigraphy of the Kaerimizu doline, Akiyoshi Limestone Plateau, Yamaguchi Prefecture, Japan.
Mem. Osaka Gakugei Univ., B, Nat. Sci., no. 14,
p. 220-229. (in Japanese with English abstract)
Sheng, J.Z. and Sun, D,D., 1975: Fusulinids of
Qinghai. 97 p., 14 pis. Geo!. Pub!. House,
Beijing. (in Chinese)- - , Zhang, L.X. and Wang, J.H., 1988:
Fusulinida. 240 p. Sci. Pub!. House, Beijing.
(in Chinese)Sun, Q.L. and Zhang, L.X., 1988: Early Permian
fusulinids from the Kongkashan pass of Xinjiang. Acta Micropalaeont. Sinica, vo!. 5, no. 4,
p.367-378, pis. 1-4. (in Chinese with English
abstract)Sun, X.F., Gao, C.X. and Zhang, X.L., 1983:
Fusulinida. In, Xi'an Inst., Geo!. Mineral
Resour. ed., Paleontological Atlas of Northwest
China, Shaanxi, Gansu and Ningxia VO!., Part 2(Upper Palaeozoic), p.3-40, pis. 1-10. Geo!.
Pub!. House, Beijing. (in Chinese)
Suyari, K., 1962: Geological and paleontologicalstudies in central and eastern Shikoku, Japan,
Part 2. Paleontology. Jour. Gakugei, Tokushima Univ., vol. 12, p. 1-63, pis. 1-12.
Takaoka, Y., 1966: Fusulinid from the Mt. Tatoro,
Mt. Kano, Mt. Futago and Mt. Shiraishi areas of
the K wanto-massif, central Japan. Bull. Chichibu Mus., Nat. His!., no. 13, p. 39-70, pis. 5-16.
(in Japanese with English description of species)
Toriyama, R., 1954: Geology of Akiyoshi, Part I.

750

Katsumi Ueno

Study of the Akiyoshi Limestone Group. Mem.
Fac. Sci., Kyushu Univ., Ser. D, Geal., vol. 4, no.
I, p. 39-97.
- - , 1958: Geology of Akiyoshi, Part 3. Fusulinids of Akiyoshi. Ibid., vol. 7, p. 1-264, pis. 148.
Wang, K.L. and Sun, X.F., 1973: Carboniferous andPermian foraminifera of the Chin ling Range andits geologic significance. Acta Geol. Sinica,
1973, no. 2, p. 137-178, pis. 1-7. (in Chinese withEnglish abstract and description of new species)
Zhou. T.M., Sheng, 1.Z. and Wang, Y.l., 1987:
Carboniferous-Permian boundary beds andfusulinid zones at Xiaodushan, Guangnan, eastern Yunnan. Acta Micropalaeont. Sinica, vol.
4, no. 2, p. 123-160, pis. 1-6.

Akiyoshi fJ\E, Chinling ~~, Guangxi $[!!l, Guizhou .tlt Itsukaichi 1i Bm, Kaerimizu
1m I) 7.K. Kochi ~9;0. Kwanto 1*1*, Mino ~1J., Otaki ::k~. Qinghai lI'ffflf:. Shomaru IEYi.
Yunnan ~m, Xinjiang ~iJI.
fJ\EEWZ:'fi~ffiJ' i? O)=-~*ciltJ5j;iJ!!R~ Pamirina: =-~*c9=tJtfj V;:.~* L- k. Verbeekinidae
W:to J: u: Neoschwagerinidae WiltJ5j;iJ!!R~O)J1[*O)m)t c ~.h -C \" J.J Pamirina hi v;::. It7!-$
1itE@lm;v;::' fine alveolar keriotheca :a:-1#--=>.J' Iv - 7' C1#t-.: t.n'.J' Iv - 7'iJ~9;o i?.h -C:to I). ~1fiO):JJiJ~ J: I) )~J&rfre iI; J.J colt x.i?.h -C \" J.J *1fiIHJ\E-Ei', 1m I) 7.Kjt£~O)tkEEWZ:'fi~
ffJ: I) 3 flO) Pamirina :a:-t9t:±l L--tO)£:±l~~:a:-t9tW L- t-.: c .:. 6. fine alveolar keriotheca:a:-1#--=>.J'IV - 7' O):JJiJ~1#t-.: tJ:\".J' Iv - 7' J: I) §jji? iJ> v;::.J::{lz: 0) ~~iJ> i?£:±l-t J.J .:. c Ht~~
L- t-.:o .:. O)*5:5lIHt. Kobayashi (1977) iJ~o1t'fi€ L- t-.: Pamirina 0) **ff. c -3$( L- -C:to I), 7!-$
1itE@lm;v;::. fine alveolar keriotheca :a:-1#t-.: tJ: \" f Iv - 7' v;::.)(;J L- -C ~~hi Levenia :a:-i:Jl!:P/§ -t J.J
00~ i?v;::.*~-eIHJ\EEWZ:'fi~ffO) Pamirina (Levenia) leveni *iJ>i?~i?.ht-.: Pamirina(P.) darvasica, P. (Levenia) leveni, P. (L.) evaluta, Misellina (Brevaxina) ? sp. :a:-llc1i:-t J.J 0.:..h i? O)iltJ5j;iJ!!R~IHtiltrJtfj=-~*cO)~Jtfj Artinskian iI; J.J \, 'It~Jtfj Yakhtashian :a:-~-t 0
J:.!llfW¥~

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 161, pp. 751-755,3 Figs., April 30, 1991

SHORT NOTES

25.

A NEW SPECIES OF PETALAXIS (RUGOSA) FROM THE
HUANGLONG FORMATION, ZHEJIANG,
SOUTHEAST CHINA *NOBUO Y AMAGIW A
Department of Earth Science, Faculty of Education,
Osaka Kyoiku University, Osaka 543

ANDE WANG
Department of Geology, Zhejiang University,
Hangzhou 310027, ChinaandSHIRO MAEDA
c/o Department of Earth Sciences, Faculty of Science,
Chiba University, Chiba 260

A new rugose coral species, Petalaxis
hangzhouensis is described in this paper.
We collected the present specimen from alimestone bed, exposed along the Manjuelong
Highway near Longjing-Wengjiashan, Hangzhou City, Zhej iang Province, Southeast
China (Figure 1). According to Wang, Lin,Luo and Chen (1987), this bed, 0.88 m thick,consists of sparry echinoderm and micritic
limestone with bioclasts, and is the third
higher bed from the base of the Upper Huanglong Formation, which is assigned to theWeiningian Stage (Moscovian), Middle Carboniferous. Besides the present new speciesof coral, this limestone yields Fusulinella
mosguensis Rauser and Safonova, F. sp.,
Eofusulina sp., Textularia sp., Nodosaria sp.,
Uraloporella sp., and Anthracoporella sp.
Acknowledgments: We thank to Dr. Guowei Lin (Zhejiang Institute of Petroleum
Geology), Messrs. Hending Wang and Yanshao Chen (Zhejiang University) for theirkind advice and suggestion in the field survey.

We acknowledge Prof. Hideo Ishikawa(Chiba University), Messrs. Ushio Fujimoto,Tadashi Endo, Hideo Shimizu and Michiji
Goto for their assistance in the field. We
also thank Dr. Yoichi Ezaki (Osaka City
University) for his discussion, and Prof.
Hisayoshi Igo (University of Tsukuba) and• Received October 3, 1989; revised manuscript accepted January 10, 1991

Figure 1. Map showing the locationof Hangzhou City (arrrow).

12 0

1-=-l!=-=-*~:5=i'~~c-='-::lC~=320
EAST'

h

'CHINA'
SEA·..·I~:(

'j?

o

50km

~

751

Nobuo Yamagiwa et al.

752

Prof. Toshio Koike (Yokohama National
University) for their offer of references.

Description of species
Family Petalaxidae Fomichev, 1953
Genus Petalaxis Milne-Edwards
and Haime, 1852

Petalaxis hangzhouensis Yamagiwa,
Wang and Maeda, n. sp.
Figures 2-1-2; 3-1-2

Diagnosis. - Small Petalaxis with thincolumella, lonsdaleoid and concentric dissepiments and relatively long minor septa.
Tabulae slope gently upwards to center.
Description. - A single corallum obtained.
Corallum compound, massive, cerioid andattains 8 cm in diameter.
Transverse section: Corallites polygonal,
five to six sided and 3.0 to 5.0 mm in diameterin mature stage. Wall thick and rather regularly zig-zagged; median dark line present.
Septa of two orders, major and minor inalternation, thin and straight or slightly sinuous. Septal spines on the wall always correspond to major and minor septa in disposition. Fine structure of septa is diffusotrabecular to fibro-normal. Major septa are
II to 13 in number, long and often attain tocenter of corallite. One of them is continuous with columella. Minor septa are relatively long, ranging from 1/2 to 4/5 length ofmajor ones in mature stage. Dissepimentarium relatively wide and consists of one to tworows of lonsdaleoid dissepiments of varying
size and one to two or three rows of concentric ones. Besides, several rows of lonsdaleoid dissepiments rarely observable.
Tabularium wide and clearly differentiatedfrom dissepimentari urn by inner wall.
Columella consists of a thin axial plate andstraight or slightly sinuous.
Longitudinal section: Dissepimentarium
occupies less than 1/2 to 1/3 radius of coral-

lite and consists of large, medium and occasionally small vesicles of dissepiments. Dissepiments generally arranged in one row, but
occasionally in a few rows. Their convex
sides facing upwards as well as inwards.
Wide tabularium consists of complete or incomplete tabulae, which gently ascending
towards the center. Clinotabellae occasionally present. Two to four tabulae are counted in a vertical distance of 1 mm. Columellathin, straight or slightly sinuous.
Comparison. - The present new species
resembles Petalaxis orboensis (de Groot),which was described from the Upper Moscovian Orbo Limestone, northern Palencia,
Spain in having a thin columella, similar
character of dissepiments, relatively longminor septa, and gently ascending tabulae.
The former, however, differs from the latter inhaving smaller corallites and less numerous
septa.
Fan (1978) also described Petalaxis orboensis (as Lithostrotionella orboensis) from theWeiningian of Sichuan Province, southwestern China. The present new species issimilar to Fan's specimen, but the former
differs from the latter in having smaller corallites, less numerous septa, some welldeveloped lonsdaleoid dissepiments and amore simple columella. Wu and Zhao
(1989) also reported Petalaxis orboensis fromthe Weiningian of eastern Yunnan. The
present our species is easily distinguishedfrom the latter by the above mentioned character.
This new species is similar to P. mohikanus(Fomichev) described from the Moscovian of
the Donetz Basin, but the present species has
smaller corallites, less numerous septa and amore simple columella. It is also distinguished from P. stylaxis (Trautschold) reported from the Middle Carboniferous of the
Moscow Basin and other areas (Stuckenberg,
1888; Dobrolyubova, 1935a,b; Fomichev,
1939; Sando, 1983, Wu and Zhao, 1974;
Wang, 1978) by lack of complex columella
and having more weakly developed lons-Short Notes 25.

New species of Petalaxis

Figure 2. Pelalaxis hangzhouensis Yamagiwa, Wang and Maeda , n. sp. 1, 2, Transverse
sections, x 14, x 5, respectively (Reg. no. CF- 8701a).

753

Nobuo Yamagiwa el al.

754

2
Figure 3. Petaloxis hangzhouensis Yamagiwa, Wang and Maeda, n. sp. 1, 2, Longitudinal
sections, x 5 (Reg. nos. CF - 8701c, 8701d, respectively).
daleoid dissepiments and gently ascending
tabulae towards the center.
Elymology.- This species is named for
Hangzhou City.
Occurrence .- Near the base of the Upper
H uanglong Formation exposed along the
Manjuelong Highway, Hangzhou City, Zhejiang Province, China.
Geological age. - Weiningian (Moscovian) ,Middle Carboniferous.
Reposilory .- Reg . no. CF - 8701 (Holotype) , housed at the Department of Geology,
Zhejiang University , China.
References cited
Dobro ly ubova, T.H., 1935a: Determinant of Middle
Carboniferous colonial Rugose co rals of the
Mosco w Basin. Vses. Naucho-Issled. Inst. Mineral. Syrya, p. 1- 14. (in Russian)- - , 1935b: Rugosa colonial co ra ls in the Middle
Carboniferous of the Moscow Basi n. Ibid. ,Trudy, no. 81 , p. I- 50, pIs. 1- 14. (in Russian withEnglish summary)F an, Y.N ., 1978 : Rugosa . p. 149- 207, pis. 50- 76. In,
Paleontological A lIas of sOlllhweSlern China,
Sichuan Provin ce, vol. 2, Geological Press, Beijing. (in Chinese)
Fomichev, V.D. , 1939 : Ph ylum Coelenterata. p.5064, pIs. 6- 11. In , Gorskiy, l.I. , ed., A tlas of lheleading forms of the fossil faunas of che
USSR. , vol. 5, Middle and Upper Carboniferous. TSNIGRI , Leningrad. (in Russian)- -, 1953: Rugose corals and stra tigraph y of Middle and Upper Ca rbonifero us and Permi an
deposits of the Donetz Basi n. VSEGEI, Tr ., p.
1- 622, atlas, pis. 1- 64. (in Russian)Groot, G .E., de, 1963 : Rugo se corals from the Carboniferous of northern Palenci a (Spain). Leidse
Geol. Meded. , vol. 29, p. 1- 123, pis. 1- 26.
Sa ndo , W.J. , 1983: Revision of Lilhoscrolionella(Coelenterata, Ru gosa) from the Carboniferous
a nd Permian. U.S. Geol. Sun>., Pro! Paper
1247, p. I-52, pis. 1- 20.
Stuckenberg, A.A. , 1888: Anthozoen und Bryozoen
des oberen mitterlrussisc hen Kohlenkalk s. Russian Mem. Com. Geo/. , vo l. 4, p . I - 54, pIs. 1- 4.

Short Notes 25.

New species of Petalaxis(in Russian with German summary)Wang, H.D., 1978: Tetracoralla. p. 106-189, pis. 2961. In, Paleontological atlas of southwestern
China, Guizho Province, vol. 2, Carboniferous Quaternary. Geological Press (Beijing). (inChinese)Wang, Y., Lin, G., Luo, Z., and Chen, X., 1987:
Hangzhou district. p.24-30. In, Carboniferous
and Permian stratigraphy in Jiangsu and Zhejiang. Guide Book for Excursion 8 (11th International of Carboniferous Stratigraphy andGeology).

755

Wu, W.S. and Zhao, 1.M., 1974: Carboniferous
corals. p.265-273, pis. 134-139. In, Handbookof the stratigraphy and paleontology of southwest China. Nanjing Inst. Geol. Pal., Acad.
Sinica. (in Chinese)- - and - - , 1989: Carboniferous and early Early
Permian Rugosa from western Guizhou and eastern Yunnan, SW. China. Pal. Sinica, no. 177,
p.I-230, pis. 1-63. (in Chinese with English
summary)

756

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New members approved at the Council Meeting on January 30, 1991
Mariko Miuchi,
Saneatsu Saito,
Akihisa Kitamura,Hisashi Suzuki,
Kiyohiro Kakusaka,
Keiichi Takahashi,
Tomio Adachi,
Kazuo Nagasawa,
Mitsuru Marukawa,
Hiroshi Takahashi,
Shigeo Shinozaki
Norie Tanaka,
Kazuhiro Nakamura,
Masamichi Yamazaki,
Tetsuya Shimazaki
Pothuri Divakar N aidu,
Takashi Nakamura,
Takuya Tanikawa,
Isao Sarashiya,
Naohiro Sato,
Hideshi Wakabayashi,
Naoki Suzuki,
Satoko Tateno,
Ayahide Kamemaru,
Satoshi Funakawa,
Yasuto Ariga,
Takayoshi Fukudomi,
Shigenori Ogihara,
Kenichi Takahashi.

759

New fellows approved
Masayuki Ehiro,
Norihisa Inuzuka,
Atsushi Kaneko,
Atsushi Takemura.
Seceding members(Fellow)
Takehiko Iwai.
(Ordinary members)
Ryuichi Arikawa,Rie Tajiri,
Deceased member(Fellow)

Hiroshi Furutani,
H iroaki Ishiga,
Masanori Shimamoto,Ken-ichiro Hisada,
Takahiro Kamiya,
Hirokazu Takahashi,

Fumitoshi Kojima,
Yataro Yamada.

Keiji Matsubara,

Akira Morishita.

PALAEONTOLOGICAL SOCIETY OF JAPAN, SPECIAL PAPERS, NUMBER 34 0)j]'(fiWi~0~ L-:;I; 9 0 *~~jfiC"~~td:j]'(fiWi~ trt~i? oy1n:J:,

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HTransactions and Proceedings of the PalaeontologicalSociety of Japan

April 30, 1991

New Series No. 161

CONTENTS
TRANSACTIONS
912. Katsuo Sashida: Early Triassic radiolarians from the Ogamata Formation,
Kanto Mountains, central Japan. Part 2. . ............................. 681
913. Osamu Sasaki: Discrimination of two new Anadara species using shell
growth parameters and Fourier Descriptors. . ........................... 697
914. Shuji Niko: Middle Carboniferous Bactritoidea (Mollusca: Cephalopoda)
from the Akiyoshi Limestone Group, Yamaguchi Prefecture (Molluscan
Paleontology of the Akiyoshi Limestone Group-X) ...................... 714
915. Atsushi Matsuoka: Early Jurassic radiolarians from the Nanjyo Massif in the
Mino Terrane, central Japan. Part 1. Tricolocapsa, Stich ocapsa , andMinocapsa, n. gen. . ................................................. 720
916. Katsumi Veno: Pamirina (Permian Fusulinacea) from the Akiyoshi Limestone Group, southwest Japan. . ....................................... 739
SHORT NOTES
25.

Nobuo Yamagiwa, Ande Wang and Siro Maeda: A new species of Petalaxis(Rugosa) from the Hunanglong Formation, Zhejiang, southeast China .... 751\

PROCEEDINGS .......................................................... 756~