ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No .

165

Palaeontologica l Society of Ja pa n
April 30, 1992

Co-Editors

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Science Museum)

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The fossil on the cover is Trilophodon sendaicus Matsumoto, an extinct elephant, which
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1009-1023, 3 Figs., April 30, 1992

932.

MOLLUSCAN FAUNA FROM THE LATE MIOCENE HATS USE
FORMATION IN THE MIURA PENINSULA,
KANAGAWA PREFECTURE, JAPAN*
KIYOSHI OKUMURA
Department of Natural Science, Naruto University of Education,
Naruto City, Tokushima Prefecture, 772
and

YUT AKA Y AMAGISHI
Kurihama Junior High School, Yokosuka City, Kanagawa Prefecture, 238

Abstract. The Hatsuse Formation distributed in the southern part of the Miura
Peninsula mainly consists of well-bedded marine tuff breccia, lappili tuff, and scoriaccous
sandstone and siltstone of marine origin with intercalated thin layers of pumice and conglomeratic sandstone. The molluscan fossils occur from a rather coarse-grained sandstonc
associatcd with fragments of pyroclastic rocks. The Hatsuse Fauna named herein is composed of three species of Gastropoda, one species of Scaphopoda, and 15 species of Bivalvia.
The Hatsuse Fauna is characteristically an admixture of euneritic and subneritic forms,
though all of them are warm-water dwellers. In consideration of specics composition andtheir mode of occurrence, the shallow-water dwellers arc interpreted to bc the ones whichwere displaced into the outcr neritic environment.
The geologic age of the Hatsuse Fauna indicates Late Miocene, and is correlated with
the Zushi Fauna distributed in the northern part of the Miura Peninsula, Kanagawa
Prefecture and the Senhata Fauna of the Boso Peninsula, Chiba Prefecturc.
Key words.

Hatsuse, Miura, Zushi, Senhata, Mollusca.

ed in the southern part of the Miura Peninsula have been lithologically divided into the
Misaki, Aburatsubo and Hatsuse Formationsin ascending order, and those formations havetentatively been correlated with Tertiary
strata in the northern part of the Miura
Peninsula (Akamine el al., 1956; Mitsunashi and Yabe, 1968). The precise geologicalage of these strata has remained undetermined. One important reason for this is that,
hitherto, no fossils have been found from the
Hatsuse Formation.
Recently, numerous fossil molluscs werecollected by the present authors from the
Hatsuse Formation. This is probably the

Introduction

The Hatsuse Formation was originallydescribed by Suzuki (1932) for the marine
strata distributed around Hatsuse-Machi inthe northern part of Miura City, southernmost part of the Miura Peninsula, Kanagawa
Prefecture. He also reported a large numberof molluscan fossils from this formation,
though these fossils were undoubtedly collected from part of the Pleistocene Miyata
Formation.
Subsequentely, the Tertiary strata distribut'Received Sepember I, 1991; accepted Decembcr 20,
1991
1009

1010

Kiyoshi Okumura and Yutaka Yamagishi

first fossil collection from this formation.
This fauna i ncl udes some characteristic
species as shown in Table 1. It can be considered that those fossil moll.uscs carry very
important information for age determinationand correlation of the Tertiary strata in thisarea.
The purpose of the present work is todiscriminate the molluscan fossils from the
Hatsuse Formation, to attempt to determine
the geological age, and to correlate thesestrata with molluscan faunas in the northernpart of the Miura and Boso Peninsulas.

Acknowledgments
The authors wish to express their appreciation to Dr. Saburo Kanno, former Professorof the University of Tsukuba for his kindencouragement during the present study andcritical reading of the manuscript.

Brief note on the geology aroundthe fossil localities

Formation (Pleistocene) consists mainly ofloose sandstone.
The Hatsuse Formation shows a strike trending generally from N 60"W to EW with the
exception of the N agahama and Arasaki areas
where it trends from N 30"E to N60"E. Also,
marked folds and faults which trend in anorthwest-to-southeast direction are developed in this formation.
There are three major synclines in Mito,
Nagahama and Kaneda and two anticlines inKurosaki-no-Hana and Kami-Miyada.
Their axes cross the line of strike of the
Hatsuse Formation (Figure I). The
Minami-Shitaura and Hikihashi faults arereverse faults thrusting up from southwest tonortheast and can be observed in several
places. The Hikihashi Fault, one of thethrust faults, is the boundary between the
Hatsuse and Aburatsubo Formations.
The maximum thickness of the Hatsuse
Formation is estimated to be about 460 m by
Akamine et al. (1956).

Fossil localities and fossil occurrences
The Miocene strata, in which the present
fossil localities are incl uded, are overlain
unconformably by the Pleistocene Miyata
Formation. According to Akamine et al.
(1956), the Miocene strata distributed in thesouthern part of the Miura Peninsula can bedivided into three stratigraphic units: Misaki, Aburatsubo and Hatsuse Formations inascending order. These formations are mainly characterized by pyroclastic rocks.
The Misaki Formation is overlain conformably by the Aburatsubo Formation, whichis overlain partially by the Hatsuse Formation. The Hatsuse Formation consists mainly of brownish or yellowish andesitic tuff
breccia and lapilli tuff which are partiallyinterbedded with scoriaceous sandstone, pyroclastic siltstone and thin layers of pumic andconglomeratic sandstone. Cross-laminations
are well developed in this formation and serve
to distinguish it from the underlying Aburatsubo Formation. The overlying Miyata

Molluscan fossils were collected from two
localities near the Wakamiya Shrine, Kanda,
Hatsuse-machi, Miura City. One (Loc. A) isa small exposure on the east side of the PublicHall of Kanda region, Miura City and otherone (Loc. B) is a small cutting exposed on themountain foot about 250 m south of theshrine, and about 450 m northeast of Loc. A(Figure I). Both fossil localities can be
regarded as of equivalent stratigraphic position.
Loc. A: Molluscan fossils occurred from a
massive, conglomeratic tuffaceous sandstone.
Most of the molluscs are sporadically scattered, and are mostly inner or outer molds,
except for the following species; Glycymeris
rotunda (Dunker), Chlamys miurensis (Yokoyama), Miyagipecten matsumoriensis
Masuda, and Amussiopecten akiyamae
Masuda.
The dominant species at Loc. A include

932.

Molluscan Fauna from the Hatsuse Formation

Glycymeris rotunda, Chlamys miurensis,
Amllssiopecten akiyamae, Miyagipecten matSlimoriensis, Laevicardium sp., and Clinocardillm sp.
Loc. B: Molluscan fossils occured from apyroclastic, conglomeratic sandstone. F ossils are sporadically scattered as in Loc. A,
but are more dense in this locality than in
Loc. A. The dominant species from thislocality are as follows; Glycymeris rotunda,
In docrassatella cf. tenuilirata (Shuto),
Laevicardium a ngustum (Y 0 koyama),
Oxyperas bernardi (Pilsbry), and Anisocorbula cf. venusta (Gould).
The molluscan fossils from both localities
are poorly preserved; even such thick-shelledmolluscs as Cypraea, Clinocardium andAnisocorbula are represented by water-worn
fragments or incomplete specimens, though a
few species are rather well preserved.
Characteristics of the Hatsuse Fauna
The molluscan fossils discriminated fromthe Hatsuse Formation incl ude three speciesof Gastropoda, one species of Scaphopoda
and 15 species of Bivalvia. Among them,
two species of Gastropoda, one species of
Scaphopoda, and II species of Bivalvia are
identified specifically. One species ofBivalvia is discriminated as a new species(Table I).
From a bathymetrical viewpoint, most of
the fossil molluscan species from Loc. A are
considered to be shallow-water dwellers infine-grained sandy or silty bottoms because of
the presence of such genera as Cypraea andA zorin us whose present-day distribution
ranges from the intertidal zone to 50 m depth.
However, other of the molluscs could possibly live in deeper waters down to about 300
m. The molluscan fossils from Loc. Barealso represented by rather shallow-water
inhabitants in fine-grained sandy or silty
bottoms except for Mikadotrochus and Cre-pidula.
The former genus dwells on deep-water

1011

rocky or gravelly bottoms at depths ranging
from about 50 to 200 m (Habe and Kosuge,1967), whereas the latter inhabits rocky orshelly bottom from littoral depths to 180 m(Keen, 1963). Another rather deep-water
dweller is Fissidentalium found at Loc. Band
this species presently lives from about 50 to
200 m depth (Habe and Kosuge, 1967).
In terms of faunal similarities between the
A and B localities, only a few species arecommon to both, namely Glycymeris rotunda,

Chlamys miurensis, Amussiopecten akiyamae
and Azorinus abbreviatlls. Other than thesefour, the two localities are characterized bydifferent molluscs.
J udgi ng from the foregoi ng, it is clear thatthe present molluscan association represents a
mixture of shallow and rather deep-water
dwellers, and of rocky- or shelly-bottom
dwellers and sandy- or muddy-bottom
dwellers. These species are considered tohave been transported for some distance fromthe initial living site in shallow to deep
waters.
Most of the species constituting the present
molluscan association are shallow, warmwater dwellers with the exception of a few
deep-water dwellers. The three extant
species among the fauna, namely Glycymeris
rotunda, Oxyperas bernardi, and Azorinus
abbreviatus, are known to be distributed in
the warm Kuroshio Current region and intemperate waters off the coast of southern tocentral Japan (south of latitude 3YN, namely
south of the Boso Peninsula).

Geological age and correlation
Some species of the Hatsuse Fauna are
restricted to strata of Miocene age.
Miyagipecten matsumoriensis has been reported from middle to late Miocene strata distributed in areas north of the Kanto region.
Amussiopecten akiyamae has been reported
from late Miocene formations in Inagozawa
and Zushi (Masuda, 1962) and also in Senhata (O'Hara and Ito, 1980; described as A.

Kiyoshi Okumura and Yutaka Yamagishi

1012

Table 1.

Geologic distributions of the molluscan fossils from the Hatsuse Formation.

~

Local.

Miocenelocali ties

Species

1 Mikadotrochus cf. yoshiwarai
2 Crepidula cf. nidatoriensis
3 Cypraea sp.
4 Fissidentalium cf. yokoyamai
5 Glycymeris rotunda
6 Chlamys miurensis
7 Amussiopecten akiyamaea
8 Miyagipecten matsumoriensis
9 Indocrassatella cf. tenuili rata
10 Laevicardium angustum
11 L. sp.
12 Clinocardium cf. hataii
13 C. hatsusense, n. sp.
14 C. sp.
15 Pitar kaniei
16 Oxyperas bernardi
17 Azorinus abbreviatus
18 Ani socorbura cf. venusta
19 A. sp.

Plio.
E

M

L

Pleist. Recent
A

h-

+
+

hr-hr--

+

hr--

r--

+
+
+

hf--

hr-hf-hr--

+
+
+

hIhI-

+
+
+
+
+
+
+

+
+
+

hr-f--

B++++

Abbreviation Plio.: Pliocene, Pleist.: Pleistocene, E: Early, M: Middle, L: Late, Local.: Fossil localities, h: Known only from the Hatsuse Formation.

praesignis).

To the best of the present
authors' knowledge, this species seems to havea restricted stratigraphic range from the latest
middle Miocene to late Miocene. Chlamys
miurensis was originally described by Y 0koyama (1920) from the basal part of theZushi Formation in Zushi City. Subsequently, Shikama (1973) reported it from thebasal part of the Zushi, O'Hara and Ito (1980)
from the Late Miocene Senhata Formation of
the 80so Peninsula, Ozaki (1954) from thebasal conglomerate of the "Tyosi" Pliocene in
Choshi City, and Kamada and Hayasaka(1959) from the Pliocene of Futaba-Tomiokaarea in Fukushima Prefecture. Mikadotrochus yoshiwarai was originally describedfrom the "Tyosi" Pliocene by Ozaki (1954).
It was also recorded from the late Miocene

Zushi Formation (Shikama, 1973) and the
Senhata Formation (O'Hara and Ito, 1980).
From these data, the molluscan fauna fromthe Hatsuse is correlated with the Zushi
Fauna (Shikama, 1973) in the Miura Peninsula and with the Senhata Fauna (O'Hara
and Ito, 1980) in the 80so Peninsula.
This correlation is supported by the geologic ranges of the respective constituent
species of the Hatsuse Fauna (Table I).
Moreover, this correlation agrees well withthat proposed by Mitsunashi and Yazaki(1968) and Mitsunashi et al. (1979), whobased their correlation on the tephra distribution in the southern Kanto region including
the 80so and Miura Peninsulas. According
to their result, the typical HK and OK key
tephra beds are recognized in the Hatsuse and932.

Molluscan Fauna from the Hatsuse Formation

Senhata Formations in the Miura and Boso
Peninsulas (Omori et al., 1986).
Systematic description
Family Pleurotomariidae
Genus Mikadotrochus Lindoholm, 1927
Mikadotrochus cf. yoshiwarai (Ozaki, 1954)
Figure 2-1 a, b

1013

basal part of the Zushi Formation (lateMiocene) of the Miura Peninsula (Shikama,1973) and the Senhata Formation (lateMiocene) of the Boso Peninsula (O'Hara andIto, 1980), besides the type locality in Choshi
City (probably from the basal part of the
Naarai Formation, late Pliocene).
Registration number.-NUH [Naruto University palaeontological collections from the
Hatsuse Formation] 87001.

Compared with.Pleurotomaria yoshiwarai Ozaki, 1954, p. 9-10, pis.
1-2.
Mikadotrochus yoshiwarai (Ozaki). Shikama, 1973,
p.92, pI. 16, figs. 3-4; O'Hara and Ito, 1980, p.
138-139, pI. 7, figs. la-c.

Remarks.- The specimen at hand is incomplete. The characteristic features of the present species are as follows; shell rather thin,depressed trochiform with more than six
whorls; each whorl slightly convex; surfaceof whorls ornamented with many spiralthreads bearing granules which are formed bytheir intersection of growth lines.
The original description of the species by
Ozaki (1954) states "surface with fine, smooth,revolving striae crossed by growth lines."
However, the Hatsuse specimen differs slightly from the type specimen in its whorl sculpture. The difference between these two specimens is possibly due to their state of preservation; actually, the type specimen appears to
be an abraded shell. A narrow and obscureslit band revolves at a position somewhat
anterior to the middle of the whorl. In otherwords, five spiral striae above (poste.rior of)
the slit and three spirals beneath the slit arecounted on the fifth whorl. Base and umbilical portions are missing.
The present species is closely related to
Mikadotrochus yoshiwarai (Ozaki, 1954)
described from the Neogene of the Choshi(Tyosi) district, Chiba Prefecture, but theincomplete, small, immature specimen underdiscussion does not permit more precise comparison with the species.
M. yoshiwarai has been reported from theFamily Crepidulidae
Genus Crepidula Lamarck, 1799
Crepidula cf. nidatoriensis Otuka, 1934
Figure 2-5a, b, c

Compared with.Crepidula nidatoriensis Otuka, 1934, p. 626, pI. 48,
figs. 63a-b.

Shell medium for the genus, semi-ovate inoutline, with two whorls; apex strongly turned posteriorly and nearly touching bodywhorl; surface almost smooth except for fineradial lines running from central part of
dorsal area to aperture. Length 21 mm,
height 16 mm, width 20 mm, minimum diameter of aperture 16 mm.
Remarks.- The single specimen in hand issomewhat abraded. The present specimen isclose to C. nidatoriensis Otuka, 1934, reported from the Kadonosawa Miocene, but thepresent species is distinct by having a more
conspicuous apex, which more strongly
curves posteriorly than that of the latter, andthe fine radial lines running from the central
part of the dorsal area to the aperture andcrossing it at a right angle. C. nidatoriensis
sogai Kamada, 1962, reported from the
N umanouchi and Kokozura Miocene in theloban district is similar to the present specimen, but the strongly curved apex and fineradial lines on the dorsal part in Crepidula cf.
nidatoriensis serve to distinguish it fromKamada's subspecies. This specimen possibly represents a new form, but is insufficientto permit the establishment of a new species.

Kiyoshi Okumura and Yutaka Yamagishi

1014

Registration number.-NUH 87002.
Family Cypraeidae
Genus Cypraea Linnaeus, 1758
Cypraea sp.

Family Glycymeriidae
Genus Glycymeris da Costa, 1778
Subgenus Glycymeris s.s.

Glycymeris (Glycymeris) rotunda(Dunker, 1882)

Figure 2-4a, b, c

Figure 2-2a, bRemarks.-More or less depressed incomplete specimens are at hand. These specimens are characterized by having no visible
spire, body whorl covered completely by
callus, and narrow aperture with denticulated
outer lip. Cypraea is usually distributed inwarm temperate to tropical regions.

Glycymeris rotunda (Dunker). Makiyama, 1927, p.
29, pI. 1, figs. 1-4; Makiyama, 1952, p. 131-138;
Hayasaka, 1962, pI. 45, figs. 8a-9b; Habe andKosuge, 1967, p.126, pI. 46, fig. 15; Kuroda,
Habe and Oyama, 1971, p.532, pI. 71, fig. 6;
Matsukura 1986, p.89, pI. 6, figs. 14, 15;
Okumura, 1988, p. 39, pI. 12, fig. I.
Pectunculus yamakawai Yokoyama, 1922, p. 190, pI.
16, figs. 4, 5.

Registration number.-NUH 87003.
Family Dentaliidae
Genus Fissidentalium Fischer, 1885
Subgenus Fissidentalium s.s.
Fissidentalium (Fissidentalium) cf. yokoyamai (Makiyama, 1931)
Figure 2-9Compared with.Dentalium (Fissidentalium) yokoyamai Makiyama,
1931, p.44, pI. 1, fig. I.

Shell slender with elongate tube, graduallyattenuating toward apex; slightly curved
dorsally; shell very thin with large numbersof flat-topped, slightly elevated longitudinal
riblets crossed by fine growth lines; longitudinal ribs bifurcated, separated by interspaces which are wider than the ribs. Longest specimen at hand; 40 mm in length,
diameter of aperture 7 mm, and 3 mm at apex.
Remarks.- The present species is closely
allied to Makiyama's species described fromthe Kakegawa Pliocene, Shizuoka Prefecture,and D. (F.) watanabei Kanno from the Chichibu Basin, Saitama Prefecture, but the present specimens differ from the latter two by
having numerous and fine longitudinal ribletson the external surface. However, the few
rather abraded specimens at hand do not
permit the establishment of a new species.

Registration number.-NUH 87004.

Remarks.- The present species is represented by the following characteristics: I)
thick and solid, obliquely suborbicular shell,
medium in size, more or less inflated shell; 2)
beak pointing slightly forwards; 3) ligamental area narrow, triangular with subequilateral chevron grooves; 4) about ten
strong teeth arranged obliquely in anterior
and posterior sides; 5) surface sculptured bynumerous low radial ribs, and inner margin
with about 40 crenations.
Several well preserved articulated as well as
nonarticulated valves were collected. This
species occurs rather frequently at both localities A and B.
Geographic distribution.-N. lat. 31" to 42"(Kuroda and Habe, 1952). This species lives
in fine-grained sandy bottoms in a depth
range of 30-300 m.
Geologic range.-Late Miocene to Recent.
Registration number.-NUH 87005.
Family Pectinidae
Subfamily Chlamydinae
Genus Chlamys R6ding, 1798
Chlamys miurensis (Yokoyama, 1920)
Figure 2-8, 10, 11

Pecten miurensis Yokoyama, 1920, p. 157-158, pI. 12,
figs. 2-6.
Chlamys miurensis (Yokoyama). Ozaki, 1954, p. 15,
pI. 9, figs. 1-2; Kamada and Hayasaka, 1959, p.

932.

Molluscan Fauna from the Hatsuse Formation

1015

N

t
Tokyo Bay~[J]J 2

D
I

Mito

3
I

4
5

* 6

~

o
I

0.5
I

Figure 1. Fossil localities in the Miura Peninsula
1 : Hatsuse Formation (Miocene) 2: Miyata Formation (Pleistocene)
5: Syncline 6: Fossil localitylkm

I

3: Alluvium deposits

4: Fault

10 16

Kiy oshi Okumura and Yulaka Yamagishi

932.

Molluscan Fauna from the Hatsuse Formation

21, pi. 2, fig. 2; Masuda, 1962, p. 177, pi. 23, fig.
I ; Shikama, 1973, p. 166, pi. 17, fig. 4; O'Hara
and Ito, 1980, pi. 16, fig. 2.

Remarks.- This species was originally described by Yokoyama (1920) from the
Tagoegawa Conglomerate Member (the basal
conglomerate of the Zushi Formation) distributed in the northern part of the Miura
Peninsula. This species is characterized byhaving 22-25 rather compressed radial ribs on
the surface which bear several riblets on therespective radials; the radial ribs appear to
be flush with the surface owing to abrasion,
but there are some scaly ri blets as seen in theventral margin illustrated in Figures 2-9, 10,II.

Two or three intercalary riblets bearing
distinct scales appear in the interspaces of theradial ribs. This species closely resembles
Chlamys kaneharai which has a wide distribution essentially in middle Miocene strata in
the Kanto region, but C. miurensis is distinguished by having several riblets on eachradial ribs, whereas only three riblets aredeveloped on each radial of C. kaneharai.
Moreover, among the three radial riblets of C.
kaneharai, the central one is most conspicuous. I n addition, this species possesses twoor three intercalary riblets in the interspacesof the radials, whereas C. kaneharai is provided with only one scaly riblet between
adjacent radial ribs.
Geographic distribution.-Pacific coast,south of Fukushima Prefecture to Shizuoka
Prefecture.
Geologic range.-Late Miocene to late
Pliocene.
Registration number.-NUH 87006,87007,

1017

87008.
Subfamily Pectininae
Genus Amussiopecten Sacco, 1897
Amussiopeclen akiyamae Masuda, 1962
Figure 3-4a, b
Amllssiopecten planicostlilatlis (Nomura and Niino).
Akiyama, 1957 (non Nomura and Niino, 1932),
p. 32, pi. 7, figs. 2-4.
Amllssiopecten akiyamae Masuda, 1962, p.224, pi.
27, figs. 1-3; Masuda, 1986, pi. 2, fig. 4.
Amllssiopecten iitomiensis (Otuka). Shikama, 1973,
pi. 17, fig. 6.
A I11l1ssiopecten praesignis (Yokoyama). 0' Hara andIto, 1980, p. 131-132, pi. 14, figs. 5-7; pi. 15, figs.
1-5.

Remarks.- This species was originally described by Masuda (1962) based on specimens
collected from the Inagozawa Formation in
Chiba Prefecture. Amussiopecten planicostulatus described by Akiyama (1957) basedon specimens collected from the Zushi Formation is also included in the present species.
A. iitomiensis (Otuka) can be distinguishedfrom the present species by having fewer but
more prominent radial ribs than those of thepresent species and no intercalary threads
between the radials. A. praesignis (Yokoyama) also differs from the present species
by having squarish, rather high, flat-topped
radial ribs on the right valve and rather broad
and roof-shaped radials on the left valve.
Geographic distribution.- This species havebeen reported only from the southern Kanto
region, the 80so and Miura Peninsulas.
Geologic range.-Late Miocene.
Registration number.-NUH 87009.
Genus Miyagipecten Masuda, 1952~
Figure 2. Molluscan fossils from the Hatsuse Formation (l). la-b. Mikadotrochlls cf. yoshiwarai
Ozaki, X 1.2, Loc. B, Reg. No. NUH 87001. 2a-b. Glycymeris (Glycymeris) rotllnda (Dunker), X I.l, Loc.
A, Reg. No. NUH 87005. 3a-b. Clinocardillm cf. hataii Hayasaka, X 1.1, Loc. A, Reg. No. NUH 87015.
4a-c. Cypraea sp., X 1.3, Loc. A, Reg. No. NUH 87003. Sa-c. Crepidlila cf. nidatoriensis Otuta, X 1.2,
Loc. B, Reg. No. NUH 78002. 6a-b. Laevicardilll11 sp., xO.8, Loc. A, Reg. No. NUH 87014. 7. Clinocardilll11 sp., X I.l, Loc. A, Reg. No. NUH 87017. 8,10,11. Chlal11Ys l11illrensis (Yokoyama), xO.6, 8: Loc.
A, Reg. No. NUH 87006, 10: Loc. B, Reg. No. NUH 87007, II: Loc. A, Reg. No. NUH 87006. 9.
Fissidentalilll11 (s. s.) cf. yokoyamai Makiyama, X l.1, Loc. B, Reg. No. NUH 87004.

Kiyoshi Okumura and Yutaka Yamagishi

1018

Miyagipecten malsumoriensis Masuda, 1952
Figure 3-6
Miyagipeclen malsumoriensis Masuda, 1952, p.252,
pI. 24, figs. 4a-7b; Masuda, 1957, p.31, pI. 4,
figs. I a-4; Masuda, 1962, p. 226, pI. 27, fig. 13 ;Masuda, 1977, p.5-6, pI. I, figs. 1-4; Masuda,
1986A, pI. 2, fig. I ; Masuda, 19868, pI. 8, fig. 9 ;Masuda and Noda, 1977, pI. 2, fig. 6; Masuda
and Takegawa, 1965, pI. I, fig. II ; O'Hara andIto, 1980, pI. 16, figs. I a -c.

Remarks.-Only one left valve is at hand.
This species is characterized by having alarge, circular outline and low convex thinsheiL The shell surface is nearly smooth
except for fine, concentric growth lines andradial striations on the umbonal area. The
inner surface is smooth without inner rib lets.
The anterior auricle of left valve is rather
smaiL Miyagipecten matsumoriensis is
known from Middle and Upper Miocene
strata in Northeast Honshu (Masuda, 1952,
1957,1977; Masuda and Noda, 1977;
Masuda and Takegawa, 1965; Iwasaki, 1970)
and the Boso Peninsula (O'Hara and Ito,
1980).
The genus Miyagipeclen has been reported
from the lower part of the Yakataga F ormation of southern Alaska by Kanno (1971).
Geographic distribution.- Northeast Honshu and Kanto region.
Geologic range.-Middle to Late Miocene.
Registration number.-NUH 87010.
Family Crassatellidae
Genus lndocrassatella Chavan, 1952lndocrassatella cf. tenllilirata (Shuto, 1957)

The anterior shell margin is rounded and theposterior one truncated. The species somewhat resembles Crassalella pauxilla (Yokoyama, 1925; Kanno, 1960; Kanno andChung, 1973), but it differs from the latter byhaving a more inflated large sheiL However,a more precise comparison between these
allied species is prevented due to the poor
condition of the materiaL
Registration number.-NUH 87011, 87012.
Family Cardiidae
Genus Laevicardium Swainson, 1840
Laevicardillm anguslllm (Yokoyama, 1925)
Figure 3-3a, b

Cardium anguslUlll Yokoyama, 1925, p. 12, pI. 4, figs.
2a-b.
Cardium (Laevicardium) anglislUm Yokoyama.
Kuroda, 1931, p. 53, pI. 5, fig. 10.
Laevicardium anguslUm Yokoyama. Kanno, 1960,
p.259, pI. 39, figs. 14a-b.

Remarks.-An internal mold is in thecollection. This species is characterized byhaving an extremely high shell compared with
its length (1.5 times the length), a strongly
convex and longitudinally ovate outline,
su bequilaterally concave anterodorsal margin, strongly convex posterodorsal margin,
and prominent umbo.
Geographic distribution.-Shigarami Formation in Nagano Prefecture, Hatsuse Formation in Kanagawa Prefecture, and Chichibu
Basin in Saitama Prefecture.
Geologic range.-Early Miocene to Early
Pliocene.
Registration number.-NUH 87013.

Figure 3-5, 12Compared with.Crassalelliles lenuiliralUm Shuto, 1957, p. 72-73, pI.
22, figs. 6-8, 12; O'Hara and Ito, 1980, pI. 16, fig.
5.

Remarks.- Two abraded specimens are athand. This species is characterized by having a subtrapezoidal, moderately inflated thinshell with regularly rounded concentric ribs.

Laevicardillm sp.
Figure 2-6a, b

Remarks.-An incomplete left valve is athand. This unnamed species has a strongly
abraded shell from the central disk to theventral border, and lacks shell material fromthe umbonal area as shown in Figure 2-6a.
The characteristic features of this species are932.

Molluscan Fauna from the Hatsuse Formation

10 19

Figure 3. Molluscan fossils fro m the Hatsuse Formation (2). I. Pilar kaniei Shikama, X I.I , Loc. A ,Reg. No. NUH 870 19. 2. C/inocardium halsusense n. sp. , X 1.5, Loc. A , Reg. No. NUH 87016. 3a - b.
Laevicardium anguslUm (Yokoyama), x l.l , Loc. B, Reg. No. NUH 870 13. 4a - b. Amussiopeclen akiyamae
Masuda , X 0.5, Loc. A , Reg . No. N UH 87009. 5, 12. indocrassale//a cf. lenui/iala (Shuto), X 1.1 , 5: Loc. A ,Reg. No. NUH 870 12, 12: Loc. A , Reg. No. NUH 870 11. 6. Miyagipeclen malsumoriensis Masuda, x O.7 ,Loc. B, Reg. No. NUH 870 10. 7,10. Azorinus abbrevialus (Gould), X I, 7: Loc. B, Reg. No. NUH 8702 1,
10 : Loc. A , Reg. No. NUH 87022. 8. Anisocorbu/a cf. venUSla (Gould), X 1.7, Loc. B, Reg. No. NUH 87023.
9. Oxyperas bernardi (Pi lsbry), X I, Loc. A, Reg. No. NU H 87020. II. Anisocorbu/a sp. , X 1.3 , Loc. B, Reg.
No . NUH 87024.

Kiyoshi Okumura and Yutaka Yamagishi

1020

its medium size and inflated-ovate and rather
thick shell. Hinge apparatus is missing.

Registration number.-NUH 87014.
Genus Clinocardium Keen, 1936
Clinocardium cf. hataii Hayasaka, 1956
Figure 2-3a, bCompared with.ClinocardiunJ hataii Hayasaka, 1956, p. 18, pI. 2, figs.
3a-b.

Remarks.- The present species was originally described by Hayasaka (1956) from the
Pliocene of the Futaba district, Fukushima
Prefecture. The characteristic features of thisspecies include: shell medium in size, equilateral, suborbicular and moderately inflatedwith conspicuous beak; anterodorsal marginrounded, merging with rounded ventral border; posterior and posterodorsal margins
broadly rounded; surface sculptured withmore than 20 strong radial ribs, but an accurate determination of the number of radials isdifficult to obtain because of their poor stateof preservation; hinge unknown.
This species somewhat resembles L.
shiobarense described from the Kanomatazawa Miocene of Tochigi Prefecture, but thelesser number of strong radials in the present
species serve to distinguish it from L. shiobar-

ense.
Registration number.-NUH 87015.

squarish radial ribs, separated by interspaces
which are narrower than breadth of radialribs; hinge inaccessible; inner margin
crenulated corresponding to external radialribs.
Measurements (in mm.).Specimens
NUH 87016
NUH 870171
NUH 870172
NUH 870173

Length
14.0
13.0
11.0
10.0

Height
10.5
10.0
8.0
7.5

Thickness
3.7
3.0
2.5
2.5

Valve
right
right
right
left

Type
Holotypc
Para type

Remark.- This new species is characterized by its quadrilateral shape, which serves
to distinguish it from other allied species.
Registration number.-NUH 87016(Holotype); 870171, 870172, 870173 (Paratype).
Clinocardium sp.
Figure 2-7

Remarks.-An inner mold is at hand.
This specimen is medium-sized, its width aslong as high, moderately convex, and equilateral with prominent umbo. The surface isornamented with many radial ribs, how manyis uncertain owing to the inadequate state ofpreservation.
Registration number.-NUH 87018.
Family Veneridae
Gen us PitaI' Romer, 1857
PitaI' kaniei Shikama, 1973

Clinocardium hatsusense, n. sp.

Figure 3-1

Figure 3-2a, b

Pilar kaniei Shikama, 1973, p. 201, pI. 17, figs. 12-13.

Description.-Shell small for the genus,
rather flattened, thin, quadrilateral shape;
posterodorsal margin nearly straight meetingwith narrowly rounded posteroventral border; ventral margin broadly and regularly
arched, ascending to anterior margin; anterodorsal margin short, making a rather distinct
corner with anterior margin; umbo prominent; apical angle about lOY; surface ornamented with about 50 flat-topped andRemarks.- The present species was originally described by Shikama (1973) from thelate Miocene Zushi Formation of the Miura
Peninsula. The following characters of thepresent species correspond well with Shikama's species; shell medium in size andmoderately inflated; beak prominent andturned forward; ventral margin broadly
arched; posteroventral corner slightly angulated, anterior margin narrowly rounded;

932.

Molluscan Fauna from the Hatsuse Formation

length about 28 mm., and thickness about
6.5- 7.0 mm.
Geographic distribution.-Miura Peninsula,
Kanagawa Prefecture.
Geologic range.-Late Miocene.
Registration number.-NUH 87019.
Family Mactridae
Genus Oxyperas March, 1853
Oxyperas bernardi (Pilsbry, 1904)
Figure 3-9

Oxyperas bernardi (Pilsbry). Kuroda, Habe andOyama, 1971, p.670, pI. 95, figs. 1-2; Matsuura
1977 pI. 9, fig. 19; O'Hara and Ito, 1980, pI. 16,
fig. 6.
Oxyperas aff. bernardi (Pilsbry). Hayasaka, 1973,
pI. 6, fig. 4.

Remarks.-An incomplete, strongly abraded right valve missing the anterior quarter ofthe shell is at hand.
Geographic distribution of living species:
South of the 80so Peninsula (about N. lat.
35') to N. lat. 25' (Kuroda and Habe, 1952),
and living from littoral depths to 100 m (Kuroda, Habe and Oyama).
Geologic range.- Late Miocene to Recent.
Registration number.-NUH 87020.

1021

Geologic range.-Late Miocene to Recent.
Registration number.-NUH 87021, 87022.
Family Corbulidae
Gen us A nisocorblila Iredale, 1930
Anisocorblila cf. venllsta (Gould, 1861)
Figure 3-8Compared with.Anisocorblila venslisia (Gould). Hayasaka, 1961, p.
63, pI. 5, figs. 7a-b; pI. 6, figs. 8a-b; Matsushima, 1969, pI. II, fig. 14; Kuroda, Habc andOyama, 1971, p.707, pI. 102, fig. 15; Itoigawa(in ltoigawa, Shibata and Nishimoto, 1974), p.
103, pI. 32, figs. 7a-8b.

Remarks.- Two specimens are at hand.
The present species is characterized by having
a solid, slightly inflated, subtrigonal shellwith a small, weakly pointed beak situatedmore or less anteriorly, and a blunt ridge
running from the beak to posteroventral corner. The present species closely resembles
Anisocorbula venusta, but these much abraded specimens do not permit definite
identification with typical A. venllsta.
Registration nllmber.-NUH 87023.
A nisocorbula sp.
Figure 3-1 I

Family Solecurtidae
Genus Azorinus Recluz, 1869
Azorinus abbreviatus (Gould, 1861)
Figure 3-7, 10

Azorinlls abbreviailis (Gould). Habe and Kosugc,
1967, p. 160, pI. 60, fig. 18; Hayasaka and Hashimoto, 1970, p. 20, pI. I, figs. 10-11 ; Kuroda,
Habe and Oyama, 1971, p. 681, pI. 99, fig. 5.

Remarks.- Two abraded specimens are athand. The shell is thin, weakly swollen, andcharacterized by a slight depression runningfrom the umbo to ventral margin. Living
specimens of this species inhabit muddy bottoms in waters from 10 to 80 m deep, and are
distributed south of the 80so Peninsula to theEast China Sea, Southeast Asia and Indian
Ocean.

Remarks.- Two poorly preserved specimens are at hand. Shells are small in size,
ovate-trigonal in shape with a blunt ridge
running from beak to posteroventral corner.
Concentric surface sculpture is slightly visible
near the ventral margin. The hinge apparatus is inaccessible.
Various species of this genus are distributedin the west Pacific regions south of southern
Hokkaido in littoral to inner neritic waters.
Registration number.-NUH 87022.
References cited
Akamine, H., Iwai, S., Koike, K., Naruse, S., Ogose,
T., Omori, M., Seki, Y., Suzuki, K. and
Watanabe, K., 1956: Geology of the Miura
Peninsula. Earlh Sci., no. 30, p.I-8. (inKiyoshi Okumura and Yutaka Yamagishi

1022Japanese with English abstract)
Akiyama, M., 1957: Amussiopecten iitomiensis(Otuka) and its allies from Japan. Trans. Proc.
Palaeont. Soc. Japan, N.S., no. 25, p. 31-39, pis.
6-7.
Habe, T. and Kosuge, S., 1967: Shells. 223 p., 64 pis.
Hoikusha, Osaka. (in Japanese)
Hayasaka, S., 1956: Pliocene Mollusaca from the
Futaba district, Fukushima Prefecture, Japan.
Saito Ho-on Kai Mus., Res., Bull., no. 25, p. 1320, pI. 2.
- - , 1961: The Geology and paleontology of theAtsumi Peninsula, Aichi Prefecture, Japan. Sci.
Rep., Tohoku Unil'., 2nd Ser. (Ceol.), vol. 33, no.
I, p. I-IOJ, pis. 1-12.
- - , 1962: Chevrons of glycymcrid shell. Trans.
Proc. Palaeont. Soc. Japan, N.S., no. 47, p. 291297, pI. 45.
- - , 1973: Pliocene marine fauna from the Tancga-Shima, South Kyushu. Sci. Rep., Tohoku
Univ., 2nd Ser. (Ceol.), Spec. Vol., no. 6, p.97108, pis. 6-7.
- - and Hashimoto, S., 1970: Discovery of marine
fossil from immediately bclow thc Ata pyroclastic flow deposits, Onejime-Cho, Osumi Peninsula,
Kagoshima Prefecture. Rep., Fac. Sci., K agoshima Univ., no. 3, p. 17-22, pI. l.
Itoigawa, J., Shibata, H. and Nishimoto, H., 1974:
Molluscan fossils of the Miocene Mizunami
Group, Central Japan. Bull., Mizunami FossilMus., no. 2, p.43-203, pis. 1-63. (in Japanesewith English abstracl)
Iwasaki. Y., 1970: The Shiobara-type Molluscan
Fauna. An ecological analysis of fossil Molluscas. Jour. Fac. Sci., Univ. Tokyo, Sec. 2, vol.
17, pt. 3, p. 351-444, pis. 1-7, 25 figs.
Kamada, Y., 1962: Tertiary marine mollusca fromthe Joban coalficld, Japan. Palaeont. Soc.
Japan, Spec. Papers, no. 8, p. 1-187, pis. 1-2l.
- - and Hayasaka, S., 1959: Remarks on a fossil
marine fauna from Tateishi, Futaba district, in
the Joban coalfield, Fukushima Prefecture.
Saito Ho-on Kai Mus., Res. Bull., no. 28, p. 1728, pI. 2.
Kanno, S., 1960: The TertiGlJI System of the Chi-chibu basin, Saitama Prefecture, central Japan.
Part II. Paleontology. p. 123-396, pls.31-5l.
Japan Soc. Promot. Sci., Tokyo.
--,1971: Tertiary molluscan fauna from the Yakataga district and adjacent area of southern Alaska. Palaeont. Soc. Japan, Spec. Papers, no. 16,
p. 1-154, pis. 1-18.
- - and Chang, L. S., 1971: Amussiopecten and its
associate fauna from northern Taiwan. Ceol.
Palaeol1l. Southeast Asia, vol. 12, p. 189-206,
pis. 30-3l.
- - and Chung, C. T., 1973: Molluscan fauna fromthe so-called Paleogene Formation in northern
Taiwan. Ibid., vol. 13, p. 91-127, pis. ID-l2.
Kcen, M., 1963: Marine molluscan genera of western North America. 126 p. Stanford Univ.
Press, Stanford, Calif.
Kuroda, T., 1931: Fossil Mollusca. In Honma,
Shinano Chubu Chishitsu-Shi (Geology of Central Shinano), Part 4, p. 1-90, pis. 1-13. (inJapanese)- - and Hahe, H., 1952: Check list and bihliography of the Recent marine mollusca of Japan.
Hosokawa Print. Co., Tokyo, 21D p.
- - , Habe T. and Oyama, K., 1971 : The sea shells of
Sagami Bay. BioI. Lab. Imp. Household. 741 p.
(in Japanese), 489 p. (in English), 121 pis. Maruzcn, Tokyo.
Makiyama, J., 1927: Molluscan fauna of the lower
part of the Kakegawa Series in the province ofTotomi, Japan. Mem. Coli. Sci., Kyoto Imp.
Univ., Ser. B., vol. 3, no. I, art. I, p. 1-1479, pis.
1-6.
- - , 1931 : Stratigraphy of the Kakegawa Pliocene
in Totomi. Ibid., vol. 7, no. I, art. I, p. I-53, pis.
1-3.
- - , 1952: On the mutation of the fossil C(l'cymerisrOll/nda. Trans. Proc. Palaeonl. Soc. Japan, N.
S., no. 5, p. 131-138.
Masuda, K., 1952: On some Miocene Pectinidae fromenvirons of Sendai. Ibid., no.8, p.249-254,
pI. l.
- - , 1957: A note on Miyagipecten matsumoriensis
Masuda. Saito Ho-on Kai Mus., Res. Bull., no.
26, p. 31-39, pI. 4.
--,1962: Tertiary Pectinidae of Japan. Sci. Rep.,
Tohoku Univ., 2nd Ser. (Ceol.), vol. 33, no. 2, p.
117-238, pis. 18-27.
- - , 1977: Miocene molluscs from the Shimokurosawa Formation, Ichinoseki City, Iwate Prefecture, Northeast Honshu, Japan. Saito Ho-on
K ai Mus., Res. Bull., no. 45, p. 3-10, pI. l.
- - , 1986A: Historical change of the Neogene
mollusca fauna of Japan with special refercnce topectinids. Monogr. Mizunami Foss. Mus., no. 6,
p. 1-21, pis. 1-3. (in Japanese with English

abstract)- - , 1986B: Notes on origin and migration of
Cenozoic pectinids in the North Pacific.
Palaeonl. Soc. Japan, Spec. Papers, no. 29, p.
95-110, pis. 7-10.
- - and Noda, H., 1977: Remarks on the Miocene
marine fauna from Watarimachi. Miyagi Prefccture, Japan. Saito Ho-on Kai Mus., Res. Bull.,no. 45, pI. 13-16, pI. I.
- - and Takegawa, 1965: Remarks on thc Miocene
mollusca from thc Scnnan district, Miyagi Prefecture, Northeast Honsh u, Japan. Ibid., no. 35, p.
1-14, pis. 1-2, I text-fig.

932.

Molluscan Fauna from the Halsuse Formation

Matsukawa, A., 1986: Cenozoic Glycymerid bivalvesof Japan. Palaeont. Soc. Japan, Spec. Papers,
no. 29, p. 77-94, pis. 5-6.
Matsushima, Y., 1969: On the molluscan fossils fromthe Alluvial deposits in Yokohama City. Bull.
Kanagawa Pref Mus. no. 2, p. 79-96, pis. 10-13.

(in Japanese with English abstract)
Matsuura, N., 1977: Molluscan fossils from the late
Pleistocene marine terrace deposits of Hokuriku
region, Japan Sea side of central Japan. Sci.
Rep.. Kanazawa Vniv., vol. 22, no. I, p. 117-162,
pis. 1-20.
Mitsunashi, T. and Yazaki,K., 1968: Geologic mapof the Miura Peninsula, Scale I: 25,000.

Geological Survey of Japan.
- - et al., 1976: Geological map of Tokyo Bay andadjacent area, Scale I: 100,000. Geological
Survey of Japan.
O'Hara, S. and Ito, M., 1980: Molluscan fossils fromthe Senhata Formation in Boso Peninsula.
Prof S. Kanno Memorial Vol., p. 121-136, pis.
14-17.
Okumura, K., 1988: Molluscan f()ssils from the Pleistocene Ninomiya Group in the Oiso Hills,
Kanagawa Prefecture, central Japan. Cullllral

Properties Rep.. Ninomiya Town Educ. Commitee, no. 16, p. 1-99. (in Japanese with English
abstract)
Omori, M., Hayama, K. and Horiguchi, M., 1986:
Kanto region (Geology of Japan. no. 3). 335 p.
Kyoritsu Publ. Co., Tokyo. (in Japanese)
Otuka, Y., 1934: Tertiary structures of the northwestern end of the Kitakami Mountainland, Iwate
Prefecture, Japan. Bull. Earthquake Res. Inst ..

Vnir. Tokyo, vol. 12, p. 566-638.
- - , 1934: On some fossils from the northern foot of
Mt. Minobu and the Hayakawa tutlite of Hakone.
Jour. Geol. Soc. Japan, vol. 41, no. 492, p.562568. (in Japanese)
Ozaki, H., 1954: On the palaeontology of the basal
conglomerate of Pliocene, in Tyosi City, Kanto
Region. Bull. Natn. Sci. Mus .. Tokyo. N.S.,
vol. I, no. I (no. 34), p. 9-21, pI. 1-10.
Shikama, T., 1973: Moll uscan assemblages of thebasal part of the Zushi Formation in Miura
Peninsula. Sci. Rep.• Tohoku Vnil'.• 2nd Ser.
(Geol.), Spec. Vol., no. 6, p. 179-204, pis. 16-17.
Shuto, T., 1957: Crasatellites and Venericardia from
the Miyazaki Group (Paleontological study of
the Miyazaki Group). Mem. Fac. Sci.• Kyushu
Vnil'.. Ser. D, vol. 6, no. 2, p. 69-90, pI. 22.
Suzuki, K., 1932: Pliocene formation north of
Atsugi-Machi, Kanagawa Prefecture (II). Jour.
Geol. Soc. Japan, vol. 39, no. 462, p. 97-13 I. (inJapanese)
Yokoyama, M., 1920: Fossils from the Miura Peninsula and its immediate north. Jour. Coli. Sci..
Imp. Vnir. Tokyo., vol. 39, art. 6, p. 1-193, pis. 120.
--,1922: Fossils from the Upper Musashino of the
Kazusa and Shimosa. Ibid., vol. 59, art. I, p. 1200, pis. 1-17.
- - , 1925: Mollusca from the Tertiary Basin of
Chichibu. Jour. Fac. Sci.• Imp. Vnil'. Tokyo.
Sec. 2, vol. I, pt. 3, p. 111-126, pis. 14-15.
- - , 1926: Tertiary mollusca from Shiobara in
Shimotsuke. Ibid, vol. I, pt. 4, p. 127-136, pis.
16-20.

Boso m1$., Kami-miyada J·.'8EE, Kaneda ~EE. Kurihama ;7.JI.U~, Kurosaki-no-hana ~~ /14, Misakiguchi -=~ I I, Mito -=P. Miura .=:00. Miurakaigan -:':iflji'liH¥, Nagahama
ffi:i%, Sagami t§t!i'!. Wakamiya %'8.
-=rffi~!,:!1'b~tmt}]F~il€l/i,kff.:iM?o{r.Em:: Ij~J~il€l\iJ\1*iJJ¥J.l{r.EO) 19 t~~~5J1j L. :?CO) 5~. g~.3g, ~~.IM~~a~.15~(5~I~g~~~)0)~.~ff0kom~

fiJj:JJJ(:glt.""t"'"' -c ll£i'lJ.El=.g""(-;I; Q -=- C, neritic 1:,~,M C outer neritic 1:,~,g 0)i1t.gIe::. ~ 0 -c -=- 0) {r.ElWt;;fiJj:IJJ(: ~.h -c l' Q -=- C, t t.:, 4'!fu&gt;, 0 -=- O)lW~ft.!=r~tJl:f~MO) {,0)""(-;1; Q -=- !.: ~§Jj 07')'1'::' Lt.:olJJi!:H il'f· IUi¥ f1ZO)~!~(;,

1023

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1024-1039, 8 Figs., April 30,1992

933.

INTRASPECIFIC VARIATION AND HETEROCHRONY
OF PHANEROLEPIDA PSEUDOTRANSENNA OZAKI(GASTROPODA: TURBINIDAE) FROM THE
PLIOCENE NOBORI FORMATION, PACIFICSIDE OF SOUTHWESTERN JAPAN*
RYUICHI MAJIMA
Geological Institute, Yokohama National University, Yokohama, 240
andAKIHIRO MURATA
Institute of Geosciences, Shizuoka University, Shizuoka, 422

Abstract. The recent discovery of 79 specimens of Phanerolepida pseudotransenna
Ozaki from the Pliocene Nobori Formation indicates that the species shows a very wide
range of individual variation in its surface ornamentation. This variation is interpreted tobe caused by differences in timing of the onset of ornamental development among individuals
in the population. An evolutionary lineage, Nehalemia hieroglyphica --> Phanerolepida
oregonensis --> P. pseudotransenna --> P. transenna, is established on the basis of theirstratigraphic occurrences, and by comparison of ontogenetic developments of shell ornamentation. At least the first two speciation events are considered to be the results of heterochrony.
Phanerolepida rehderi MacNeil is designated as a junior synonym of P. pseudotransenna. Four opercula are reported and illustrated in association with their own shells, near totheir life position.

Key words. Phanerolepida, intraspecific vanatlOn,
Pliocene, Nobori Formation, southwestern Japan.

heterochrony,

Gastropoda,

North America (Hickman, 1972) to its lateroccurrences which have been restricted to theJapanese Neogene and Recent faunas, are of
special interest.
In Japan, the following five species of
Phaneroleida have previously been known:
Phaneroleida transenna (Watson, 1879),
Pliocene (Otuka, 1949; Okutani, 1968)Recent (Watson, 1879 and 1886; Dall, 1907;
Ok utani, 1964, 1966, 1968, 1969; N oda andOgasawara, 1976);
P. expansilabrum Kuroda, 1931, Miocene(Kuroda, 1931; Watanabe et al., 1950;
Shibata, 1974 [as P. cf. expansilabrumJ;

Introduction

Phanerolepida is a very interesting genus
from the malacological, conchological andpaleontological viewpoints. Ecologically it
is a homalopomatine, exceptionally living in
the very deep sea, and morphologically itbears a peculiar rhombohedral network pattern of ornamentation. The evolution andmigration of Phanerolepida, from its first
appearance in the early Oligocene of western'Received September 17, 1991; revised manuscript
accepted December 20, 1991
1024

933.

Intraspecific variation and heterochrony of Phanerolepida

Noda and Kikuchi, 1980; Itoigawa el al.,
198 I, 1982 [as P. cf. expansilabrllmJ) ;P. pselldotransenna Ozaki, 1956, ? Miocene(Hatai and Masuda, 1962) - Pliocene (Ozaki,
1956; N oda and Ogasawara, 1976; Mimoto,1986).
P. rehderi MacNeil, 1961, Pliocene (MacNeil, 196 I ; N oda and Ogasawara, 1976;
Noda, 1980, 1988); andP. okinawana Noda, 1988, Pliocene (Noda,
1988).
Their morphological comparison and taxonomic validity have not been well
documented because some of them were
proposed on the basis of poorly preserved
material in which the type material of P.
pselldotransenna is included. From summerof 1987 to April of 1990, we collected 79
specimens of P. pselldotransenna, including
many that are well preserved. The aim of
this study is to describe P. pseudotransenna
in detail, to show the mode of individual
ontogenetic variation, to explain the origin of
this variation, and to evaluate the role ofheterochrony in the evolution of the genus.

1025

Abbrevi a tions
The following abbreviations are used inthis paper:
IGSU, Institute of Geosciences, Shizuoka
University, Shizuoka, 422 Japan; andNSM, National Science Museum, Shinzyukuku, Tokyo, 169 Japan.

Collecting localities
Specimens of Phanerolepida pseudotransenna examined for this study were collectedfrom the massive mudstone of the Pliocene
Nobori Formation of the Tonohama Group,
exposed in a quarry at Nobori, Hanemachi,
Muroto City, Kochi Prefecture, on the Pacificside of southwestern Japan (Figure I). Our
three localities in the quarry from which thespecies was collected are as follows:
IGSU loc. no. 62-35- I : a cliff located insouthern end of the quarry, consisting ofmassive, pale gray mudstone intercalated withlenticular molluscan shell beds about IO cm
in maximum thickness and over 6 m in maxi-

SEA OF JAPANPACIFIC OCEAN

Figure 1. Index map showing the fossil localities of Phanerolepida pseudotransenna Ozaki from theupper Pliocene Nobori Formation. The fossil localities are plotted on a I : 25,OOO-scale topographic map ofJapan, Quadrangle "Hane," Geographical Survey of Japan.

1026

Ry uichi Majima and Akihiro Murala

Figure 2. A view of the qua rry cliff ( I) a nd the mode of occurrence of Phanerolepida pseudotransenna
Oza ki (2 - 4) at IGS U loc. no. 62- 35 - 3.

mum width. The matri x of the lentic ul a r
shell beds is composed of mudstone.
IGSU loc. no. 62 - 35 - 2: a cliff, si tu a ted
near the center of the quarry, consist in g ofmassive, pale brownish gray, weak ly weat hered mud stone.
IGSU loc. no. 62 - 35- 3: a large cliff (Figure 2- 1) exposed in nort hern end of thequarry consist in g of massive, pale gray mudsto ne.
Except fo r the len ticul ar she ll beds at IGSU
loc. no. 62 - 35- 1, th e moll usca n fo ssi Is fromthese loca lities a re considered to have beenpreserved in SiIU, because: ( I) they occ ur
sporadica ll y in massive mud stone (Figure 22 - 4) ; (2) many bi va lves are preserved inarticul ated condition; and (3) four specime ns of Phanerolepida pseudolransenna bearo percu la within their own apertures near totheir life position (Figure 8).
The massive mud sto ne of the three loca liti es sporad ica ll y yie ld s the following deepwate r species: Tenuileda ikebei (Suzuki andKanehara, 1936), Portlandia sp., Limopsis
tajimae Sowerby, 1914, L. chilaniana Yokoyama, 1926, Glycymeris nipponica (Yokoyama , 1920), Acesla golialh (Sowerby, 1883),
Deleclopeclen peckhami (Gabb , 1869),
Nemocardium sp., Cuspidaria hirasei K uroda , 1948, Fissidentalium y okoyamai (Makiyama , 193 1), Balhy bembix aeola (Watson ,1878), Ginebis argenteonitens (Lischke, 1872),
Phanero/epida pseudolransenna Oza ki , 1956,
Aulacofusus cerulescens Kuroda a nd Habe,
1966, Neplunea noboriensis Ozak i, 1956,
Turrancilla sua vis (Yokoyam a, 1926), Fulgoraria (Musashia) cancellala Kuroda a nd
Ha be, 1950, F. (Saolomea) delicata (Fulton ,933.

Intraspecific variation and heterochrony of Phanerolepida1940), an d Makiyamaia coreanica (Adams
and Reeve, 1849) .
The lenticul ar shell beds at IGSU loc. no.
62 - 35 - 1 bear the spec ies A nadara (Tosarca)
tosaensis (N oda , 1965), Glycymeris cf. yessoensis (Sowerby, 1886), Chlamys satoi (Yokoyama, 1928), Chlamys sp ., Cryptopecten
vesiculosus (Dunker, 1877) , Amussiopecten
praesignis (Yokoyama , 1922) , Venericardia
panda (Yokoyama, 1926), Dimya sp., Crassatellites sp. , Paphia sp. , Placamen tiara (Dillwyn , 18 17), Dentalium sp., Umbonium (Suchium) suchiense Yokoyama , 192 3, Phan erolepida pseudotransenna Ozaki, 1956, Turri/ella perterebra Yokoyama, 1923, Glossaulax

..y

1a

(SRFsRFsFRFsF)

(SRF)

1027

hyugensis (Shuto , 1964), G. hagenoshitensis(Shuto , 1964) , Mammilla sp. , Oyptonatica
andoi (Nomura , 1935), C. adamsiana (Dunker, 1859), Babylonia ela/a (Yokoyama ,1923) , Siphonalia tonohamaensis N omura ,1937, S. y abei Nomura , 1937, Baryspira albocallosa (Lischke, 1873), Oliva mustellina(Yokoyama , 1923), Granulifusus dualis (Yokoyama , 1928) , Conus sp., Orthosurcula pervirgo (Yokoyama , 192 8), Spirotropis subdec!ivis (Yokoyama , 1926), and A rchitectonica
sp. Because these mollu scs are composed ofboth sha ll ow a nd deep water species, they are
considered to have been more or less reworked .

(SRF)

Figure 3. Phanero/epic/a pseuc/olransenna Ozaki. l a - 3b, IGS U loco no. 62- 35 - 1, X 1.7 : l a - lb, IGSU
M37 ; 2a-2 b, IGSU M3 8; 3a - 3b , IGSU M39. Letters around shel l indica te successive ornament patterns on
body whorl. 4, 5, IGS U loc. no. 62- 35 - 3, X 2.6: 4, IGSU M60; 5, IGSU M64. Arrows R, F , and s indicate
th e extent of the respective patterns on bod y whorl.

Ryuichi Majima and Akihiro Murata

1028

Table 1.

Comparison for letter formula between Shinzato and Nobori specimens.

NOBORI FORMATION(IGSU LOC, NO. 62-35-3)
25 SPECIMENSSHINZA TO FORMATIONOKINAWA (NODA AND OGASAWARA, 1976)
43 SPECIMENSLETTER FORMULA

NUMBER OF
SPECIMENS

See text for details.

LETTER FORMULA

NUMBER OF
SPECIMENS

SRF

10

SRF

11

SRFsF

5

SRFsF

2

SsF
SRFs

21

SRFs

SRs

2

SsFs

1

Ss

3

?Ss
SR

2

SRFRF
SRFsRsSRFRFRFSRFRFRFs
SRFRFRFRF
SRFsRFsRF
SRFsFRFRF
SRFRFRFsRFs(A)

LETTER FORMULA

LETTER FORMULA

SRF

SRF

SRF(s)F -+ SRFRF

SRF(s)F -+ SRFRF

S(s)F-+SRF
SRF(s)-+SRFR

SRF(s)-+SRFR

SR(s)-+SR
S(s)F(s)-+SRFR
S(s)-+SR?S(s)-+ ?SRSR
SRFRF
SRF(s)R(s)-+SRFRSRFRFRFSRFRFRF(s)-+ SRFRFRFRSRFRFRFRF
SRF(s)RF(s)RF -+SRFRFRF
SRF(s)FRFRF -+ SRFRFRFRF(B)

SRFRFRF(s)RF(s)-+SRFRFRFRFRLETTER FORMULA(C)NUMBER OF
SPECIMENS5LETTER FORMULA

NUMBERS OF
SPECIMENS

SR
SRF

11

SRFR

22

SRFR

2

SRFRF

5

SRFRF

4

SRFRFRFSRFRFRFRSRFRFRFRF

2

SR
SRF

SRFRFRFRFR(1 +?1)
11

2

933.

Intraspecific variation and heterochrony of Phanerolepida

Intraspecific variation of Phanerolepida
pseudotransenna Ozaki
Ornamentation of the teleoconch of
Phanerolepida pseudotransenna is divided
into four patterns: three or four spiral cords
on the smooth surface (S in abbreviation;
Figure 3-4-5 [arrow SJ), a roughly developed rhombohedral network (R; Figure 34- 5 [arrow RJ), a finely developed rhombohedral network (F; Figure 3-4 - 5 [arrow
FJ), and a smooth surface occasionallybearing very fine spiral striae (s; Figure 3-5[arrow sJ). The pattern boundaries are commonly sharply separated from each other, but
the pattern R, when R immediately follows
the pattern S, is sometimes superposed byweak spiral cords that continue from thespiral cords of the pattern S (Figure 3-la)and the R is sometimes gradually changed
into the s (see boundary between the Rand s
in Figure 3-5). In the following discussion,
we use a letter formula for the description of
the shell ornamentation for respective individuals. The above four abbreviations of thepattern are arranged ontogenetically. For
example, when the teleoconch is sculptured
from earlier to later with S -> R -> F -> s,then we use the letter formula "SRFs."
Noda and Ogasawara (1976) showed theindividual variety of the shell ornamentationof Phanerolepida rehderi MacNeil, 1961, here
considered to be a junior synonym of P.
pseudotransenna Ozaki, 1956, by using material from the upper Pliocene Shinzato Formation, Okinawa Prefecture, southern Japan.
Noda and Ogasawara's (1976) work is summarized as follows.
Surface ornamentation of teleoconchchanges ontogenetically. The first, second,
and rarely third teleoconchs have the pattern
S, and its later teleoconchs are sculptured
with R, F, and/or s. The degree of development of these four patterns (S, R, F, and s) isgreatly variable in individuals. The 43 Shinzato specimens were classified into the following 7 different development types (Table 1-

1029

A): SRF (10 specimens), SRFsF (5 specimens), SsF (I specimen), SRFs (21 specimens), SRs (2 specimens), SsFs (I specimen),and Ss (3 specimens).
We observed 79 specimens (see Appendix)
from the Nobori Formation, of which weanalyzed 25 well preserved specimens fromthe IGSU loc. no. 62-35-3, where P.
pseudotransenna occurs autochthonously andmost abundantly. Among the 25 specimens(Table I-A, Figures 4,5), 11 specimens areclassifiable into Type SRF (Figure 5-9alOb), 2 specimens into SRFsF (Figure 5-8a8b), I specimen into SRFs, and I specimen
into a questionable Ss (Figure 5-12a-12b).
The remaining 10 specimens bear ornamentation that is not classifiable into any of thetypes of Noda and Ogasawara (1976) (Table
I-A). They consist of the Type SR (I specimen; Figure 5-11 a - II b), SRFRF (2 specimens; Figure 5-6a - 6b), SRFsRs (I specimen; Figure 5-7a - 7b), SRFRFRF (I specimen; Figure 5-5a-5b), SRFRFRFs (I specimen; Figures 3-4,5-4a-4b), SRFRFRFRF(I specimen; Figure 5-1 a - I b), SRFsRFsRF(I specimen), SRFsFRFRF (I specimen;
Figure5-3a-3b), and SRFRFRFsRFs (1specimen; Figures 3-5, 5-2a-2b). The
ranges of individual variation of the shellsurface ornamentation of the Shinzato andNobori specimens are quite different (Table
I-A).
We propose herein a sorting scheme for thesurface ornamentation. As noted above, thepattern R sometimes becomes coarser andchanges gradually into the pattern s (Figure3-5). Thus, the pattern s is considered to be
the coarsest part of the pattern R. In thefollowing discussion of pattern arrangement
the s is replaced by R (Table 1-8).
In this sorting scheme (Table 1-8), thethree patterns S, R, and F are regularly arranged in both the Shinzato and Nobori specimens: that is, the teleoconch ornamentation
always begins with the pattern S; the S isalways followed by the pattern R; the R isalways succeeded by the pattern F; and the R

Ryuichi Majima and Akihiro Murata

1030

SPECIMEN(IGSU·M) EARLY2In

LETTER FORMULA

VOLUTION
2

LATE

I

64
69
72

SRFRFRFRF
SRFRFRFsRFs
SRFsRFsRF
SRFsFRFRF

6~

SRFRFRFsm

SRFRFRF

65
74
75
97
77

H

SRFRF
SRFRF
SRFsRs
SRF
SRF
SRFsF

91
71
63
59
76
94
66
70
6l
67
83

SRF
SRFs

68

SR

SRF
SRF
SRF
SRF
SRFsF
SRF
SRF
SRF
SRF

LEGEND

D1

02

rn3
~4

U5
~6

I7

-,~8

933.

Intraspecific variation and heterochrony of Phanerolepida

and F appear repeatedly in some specimens(Table I-C).
These differences among individual ornamental patterns are not considered to becontrolled by age differences because there is
no relationship between number of whorls
and ornament type (Figure 4; Noda andOgasawara, 1976, fig. 3).
We propose a model to explain the
different ranges of individual variation
between the Shinzato and Nobori specimens.
This model maintains that the differencesamong these individual ornamental types arecontrolled by the differences in timing of theonset of ornamental development in individuals and that the onset timing of respective
individuals differs successively (Figure 6-1).
In Figure 6-1, if the onset timing is extremely
accelerated, then an SRFRFRFRF-type ornamentation results, while if the onset timing isgreatly delayed, then an SR-type ornamentation appears. In this model, the range ofindividual variation on shell ornamentation
is primarily controlled by the range of variation in timing of the onset of the ornamentdevelopment (Figure 6-1). The Nobori
specimens are, therefore, considered to have awider range in timing of the onset of ornament development than the Shinzato specimens. This mode of individual variation
controlled by such a mechanism has beenreported by Majima (1985, 1988), who studied
the heterochronic evolution of three naticid
gastropods.
The patterns Rand F in the later adultstage of some specimens (lGSU M69, M72,

1031

M60, MIOO and M65 in Figure 4) are very
closely arranged in comparison with theothers. We consider these close arrangements to be caused by slow growth rate due
either to the later adult stage, worse trophic
conditions, or reproductive activity.
Evolution of Phanerolepida
We hypothesize that the Recent Phanerolepida transenna evolved from Eocene Nehalemia hieroglyphica through Oligocene P.
oregonensis and Pliocene P. pseudotransenna.
In the speciation events represented, at least
two events are considered to have been the
results of heterochrony, as discussed below.
Phanerolepida is considered to have
evolved from the western North American
species Nehalemia hieroglyphica Hickman,
1974. Nehalemia hieroglyphica Hickman
shares characteristics of both Phanerolepidaand Homalopoma and occurs near the top of
the Eocene Cowlitz Formation, approximately 450 m stratigraphically beneath the oldest
occurrence of Phanerolepida, P. oregonensis
Hickman, in the lower Oligocene Keasey
Formation (Hickman, 1974). On the
teleoconch surface, N. hieroglyphica is ornamented with strong spiral ribs separated byapproximately equal interspaces (Hickman,
1974). This ornamentation is considered tobe comparable to the pattern S of P.
pseudotransenna (Figure 6-3).
Phanerolepida oregonensis is considered tohave evolved from N. hieroglyphica, and has a
net-like, coarse rhombohedral pattern of sur-+- Figure 4. Individual variation of ornamentation on shells of Phanerolepida pseudotransenna Ozaki
at IGSU loc. no. 62-35-3. Respective lateral columns illustrate successive ornamental patterns. The "0"position of the volution coincides with the boundary between the patterns Rand S but not with protoconch/teleoconch boundary. In many specimens, the protoconch/teleoconch boundary is indistinct due toweathering. Legend: I, protoconch or pattern S (three or four spiral cords on the smooth surface) indeterminable due to weathering; 2, protoconch; 3, pattern of three or four spiral cords; 4, roughly developedrhombohedral network pattern; 5, finely developed rhombohedral network pattern; 6, smooth surface
occasionally bearing very fine spiral striae; 7, sculptural boundary preserved; 8, sculptural boundary or
early conchs indeterminable due to weathering.
IGSU M86 specimen (Figure 5-12a- b; Table I-A, ? Ss) is not illustrated in this figure because of its
imperfect preservation of ornamentation. So the total number of specimens in this figure is 24, but not 25
mentioned in the text.

Ry uichi Majima and Akihiro Murata

103 2

F

~"

2a
/

(SRFRFRFRF)(SRFsFRFRF)(SRFRFRFsRFs)(SRFRFRFs) 4a2b"(SRFRF)S(SRFsF)(SRFsRs)

/'

7b
8b

(SRF)

(SRF)

(SR)

11a

(5?s?s)933.

Intraspecific variation and heterochrony of Phanerolepida

face ornamentation, and its early teleoconchs
have a faint spiral sculpture superposed on
the rhombohedral network (Hickman, 1972)(Figure 6-2). The coarse rhombohedral network pattern is considered to be comparable
to the pattern R of P. pseudotransenna, andthe superposed faint spirals are probably
comparable with the spiral ribs of N. hieroglyphica, and also with the pattern S of P.
pseudotransenna.
Phanerolepida pseudotransenna is inferredto have evolved from P. oregonensis in a
migration process from the eastern Pacific to
the western Pacific during the late Oligocene
or early Miocene. The former species showsa very wide range of individual variation inornamentation and has a variant bearing
SR-type ornamentation which is interpreted
as a relic of P. oregonensis. In some specimens of P. pseudotransenna, the pattern R
that immediately follows the pattern S issuperposed by weak spiral cords that contin ue from the spiral cords of the pattern S(Figure 3-la), just as with the early teleoconchs of P. oregonensis. This similarity supports the above interpretation for the lineage.
Phanerolepida transenna is considered tohave evolved from P. pseudotransenna, andcommonly shows an ontogenetic ornamental
sequence on the teleoconch, i.e., faint spiralsculpture --> coarse rhombohedral network
pattern --> fine rhombohedral network pattern, which are considered to be comparablewith the patterns S, R, and F of P.
pseudotransenna, respectively. The derivation of P. transenna from P. pseudotransennais supported by evidence that P. pseudotransenna has a variant bearing SRF-type ornamentation that is considered to be a forerunner of P. transenna. The precise individual variation of the ornamentation of P.
transenna is not well known because of thelack of available specimens, but the following1033

can be stated. The faint spiral sculpture of
P. transenna may be indistinct in some specimens, and the fine rhombohedral network
pattern is commonly widely developed, finerin grain size than the pattern F of P.
pseudotransenna, and may be interrupted by
the coarse rhombohedral network pattern.
The evolutionary lineage Nehalemia hieroglyphica (ornament pattern: S) --> P.
oregonensis (S + R --> R) --> P. pseudotransenna (S --> R <- variation --> S --> R --> F --> R--> F --> R --> F --> R --> F --> R) is considered
to be an example of peramorphosis (Figure6). Peramorphosis is the occurrence of theancestral adult morphology in a descendantjuvenile stage of development (McNamara,1986). A highly variable onset timing of
ornament development in individuals occurs
in the evolutionary process from P. oregonensis to P. pseudotransenna. Although thewide range of individual ontogenetic variation in P. pseudotransenna makes interpretation of the heterochrony complicated, thisevolution is evidently a peramorphosis
because many specimens of P. pseudotransenna have the ancestral patterns Sand R in
their younger stages.

Systematic Paleontology
Class Gastropoda Cuvier, 1797
Subclass Prosobranchia Milne-Edwards, 1848
Order Archaeogastropoda Thiele, 1925
Family Turbinidae Rafinesque, 1815
Subfamily Homalopomatinae Keen, 1960
Gen us Phanerolepida Dall, 1907
Type species. - Turbo transenna Watson,
1879, by monotypy. Holocene, Pacific sideof central and southwestern Japan (Noda andOgasawara, 1976).

~
Figure S. Phanero/epida pseudotransenna Ozaki. IGSU loco no. 62-35-3. X 1.7. la-lb, IGSU
M73; 2a-2b, IGSU M64; 3a-3b, IGSU M72; 4a-4b, IGSU M60; 5a-5b, IGSU MIOO; 6a-6b, IGSU
M74; 7a-7b, IGSU M75; 8a-8b, IGSU M61; 9a-9b, IGSU M94; lOa-lOb, IGSU M67; lla-llb,
IGSU M68; 12a - 12b, IGSU M86. Letters around the shell indicate the successive patterns on body whorl.

Ryuichi Majima and Akihiro Murata

1034---------Ontogeny--------Early whorl ~..- - - - - - - - - - - - late whorl

t
::0

'"

:::>

"'"'"

~

i

:::>

=:c.

g:

'"
-=

'"
'"

~

i' gO!!!

§:

I

'~"

o

..

Phanerolepida
pseudotransenna

PERAMORPHOSIS•i? XXXXXXXXXXXXXJ(S+R) . . .

(R)

PhanerolepidC!
oregonensis(S)

Nehalemia
hieroglyphica

PERAMORPHOSIS

?

•

2

3

Figure 6. Schematic illustration of a model for the explanation of individual variation of Phanerolepida
pseudotransenna Ozaki (I), and a heterochrony of Nehalemia hieroglyphica Hickman (3) ---> Phanerolepida
oregonensis Hickman (2) ---> P. pseudotransenna Ozaki (I) lineage. Note that the difference of individual variation between Shinzato and Nobori specimens is interpreted to be caused by variation in the onset timing of
ornament development for each individual.

Phanerolepida pseudotransenna
Ozaki, 1956
Figures 3,5, 7, 8

Phanerolepida pseudotransenna Ozaki, 1956, p. 1-2,
pI. I, figs. 1-2; Noda and Ogasawara, 1976, pI. I,
fig. 13;
Masuda and Noda, 1976, p. 246 ;Mimoto, 1986, p. 36, pI. 15, figs.5a-6b.
? Phanerolepida pseudotransenna Ozaki. Hatai and
Masuda, 1962, pI. 40, figs. 26, 27.
Phanerolepida rehderi MacNeil, 1960, p. 30, pI. 7, figs.
6-8, 11-13; Hickman, 1972, figs. 3,7, II, 13;
Noda and Ogasawara, 1976, pI. I, figs. la-lOb;
Masuda and Noda, 1976, p. 246; Noda, 1980, p.
II, pI. 6, fig. I ; Noda and Kikuchi, 1980, pI. I,

933.

Intraspecific variation and heterochrony of Phan erolepida

Figure 7. Four views of the ho loty pe of
Phan ero/epida pseudolransenna Oza ki. NSM PI
4368. x 1. 8. Nisinotani , Nobori , Ha ne- mach i, Muroto C ity, Koc hi Prefecture, Pac ifi c side of so uth western J a pa n. Uppe r Pli ocene N o bo ri Formation.

fi gs. 2a, 2b, 13, ?4 [ figs. 3, 4, show in g operc ul a
iso lated from th e shell ] ; N oda , 1988, p. 34, pI. 5,
figs. I a - 3c.
Phan ero/epida Iransenna (W a tso n) . Aoki a nd Baba ,198 4, p. 73, figs. I a, I b [ not Ph anero/epida lransenn a (Watson , 1879)].

Type malerial. - NSM PI 4368 (h o lotype:Fi g ure 7) from Ni sin o ta ni , N o bo ri , H a nemac hi (Ozaki , 1956), Muro to City, Koc hi
Prefecture, Pacific side of so uth western Japa n.
Pli oce ne Nobori F orm a tion of th e Ton oha m a Group .
Stratigraphic occurrence. - ? Mi oce ne: T o kigawa Formati o n, Sa ita ma Prefecture (H ata i
a nd Mas uda , 1962 ). Pliocene: N o bo ri Form ation , K oc hi Prefecture ; Shinzato F o rm ation , Okin a wa Prefecture (M ac N e il , 1960;
N o d a and Ogasawa ra, 1976 ; N oda , 1980,
1988 ).
R emarks. - W e collected 5 opercula of P.
pseudolransenna from IGSU loco no. 62 - 35 3, of which 4 are preserved in the ir ow na pertures near to their life posi tion (Figure 81, 8- 3 - 5). These o percula a re indi stingui shable from tho se of the Recent spec ies, P.
transenna.
Phanerolepida rehderi M ac Neil , 1960, a1035

Figure 8. Opercula of Phan e ro/ep ida
pseudOiransenna Ozaki. IGSU loc. no . 62 - 35 - 3.
X 1.7. 1, 3 - 5, operc ul a preserved in th e ape rture nea r
to th ei r life positi o n. 2, operc ulum occ urrin g isola ted from its ow n shell . I, IGSU M 73; 2, IGSU
M117 ; 3, IGS U M112 ; 4, IG SU M69 ; 5, IGSUMill.

Plioce ne speci es of Okina wa, so uthern Jap a n,is here designated as a junior sy no nym of th epresent speci es for the first time . The tw ospec ies a re ide ntica l to each other in shellform . Although th e tw o spec ies h ave
differe nt ra nges of indi vidu a l o rn a menta lvariations (Ta ble I - A) , th ese differences a reinterpreted as intras pecific var iati o n res ultin g
fr o m the variational ra nge of onset timing o fo rname nt deve lo pme nt (Fi g ure 6- 1).
Phanerolepida expa nsilabrum Kuroda ,1931 from th e Mioce ne o f J a pa n is intermediate in age between P. oregonensis a nd P.
pseudolra nsenna. Because P. expansilabrum
has prev iou sly been kno w n o nl y by poorly
preserved spec imen s, spec ific comparisonsa mon g the three species mu st awa it availability of numero us well preserved spec imen s o f
the Mi oce ne spec ies.

Acknowledgments
W e wo uld like to th a nk Prof. H . N oda ,University o f T sukub a , for hi s kind di sc us-

1036

Ryuichi Majima and Akihiro Murata

sion. We are grateful to Dr. J. Mark Erickson, St. Lawrence University, Dr. T. Koike,
Yokohama National University, and an
anonymous reviewer for their review of theearly or final version of the manuscript andproviding helpful comments. Dr. T. Kase,
National Science Museum, kindly sent us
photographs of the holotype of Phanerolepida pseudotransenna. Dr. Warren C. Blow,
Smithsonian Institution, kindly allowed
access to the type specimens of Nehalemia
hieroglyphica and Phanerolepida oregonensis
in his care. Messrs. R. Uchimura and H.
Ishimoto, Shizuoka University, kindly assisted in collecting fossil specimens. This study
was funded by Grants-in-Aids for Science
Research from the Ministry of Education,
Science and Culture, the Government ofJapan (Nos. 02740394 and 03640654).
References cited
Aoki, N. and Baba, K., 1984: Additions to themolluscan fossils from the Nobori Formation,
Shikoku. Ann. Rep., Ins!. Geosci., Univ. Tsukuba, no. 10, p. 73~79.
Dall, W.H., 1907: Descriptions of new species ofshells, chiefly Buccinidae, from the dredgings of
the U.S. "Albatross" during 1906, in the northwestern Pacific, Bering, and Japanese Sea.
Smithsonian Misc. Coli., vol. 50, pt. 2, p.
139~ 173.
Hatai, K. and Masuda, K., 1962: Megafossils fromnear Higashi-Matsuyama City. Saitama Prefecture, Japan. Trans. Proc. Palaeont. Soc. Japan,
N.S., no. 46, p. 254~262, pI. 40.
Hickman, C. S., 1972: Review of the bathyal gastropod Phanerolepida (Homalopomatinae) anddescription of a new species from the Oregon
Oligocene. Veliger, vol. 15, no. 2, p. 107~112.
- - , 1974: Nehalemia hieroglyphica, a new genus
and species of archaeogastropod (Turbinidae:
Homalopomatinae) from the Eocene of Oregon.
Veliger, vol. 17, no. 2, p. 89~91.
Itoigawa, J., Shibata, H., Nishimoto, H., and Okumura, Y., 1981 : Miocene fossils of the Mizunami
Group, central Japan. 2. Molluscs. Monogr.
Mizunami Fossil Mus., No. 3~A, p. 1~53, pis. I~
52. (in Japanese).
- - , - - , - - , and - - , 1982: Miocene fossils of
the Mizunami Group, central Japan. 2. Molluscs (continued). Monogr. Mizunami FossilMus., No. 3~B, p. 1~330. (in Japanese).
Kuroda, T., 1931: Fossil Mollusca. In Homma, F.,
Shinano Chabu Chisitsu-Shi (Geology of Central Shinano), part 4, p. I ~90, pis. I ~ 13. (inJapanese).
MacNeil, F.S., 1960: Tertiary and Quaternary Gastropoda of Okinawa. u.s. Geol. Surv., Prof
Paper 339, p. I ~ 148, pis. I ~ 19. (published in1961).
McNamara, K.J., 1986: A guide to the nomenclatureof heterochrony. Jour. Paleont., vol. 60, no. I,
p. 4~13.
Majima, R., 1985: Intraspecific variation in three
species of Glossaulax (Gastropoda: Naticidae)
from the late Cenozoic strata in central andsouthwest Japan. Trans. Proc. Palaeont. Soc.
Japan, N.S., no. 138, p. III ~ 137, pis. 17~ 19.
1988: Evolution of Japanese Glossaulax
species (Gastropoda: Naticidae). II: G.
hyugensis, G. hagenoshitensis and G.nodai. Fossils (Palaeont. Soc. Japan), no. 45, p. 1~7. (inJapanese with English abstract).
Masuda, K. and Noda, H., 1976: Checklist andBibliography of the Tertiary and Quaternary
Mollusca of Japan, 1950~ 1974. 494 p. Saito
Ho-on Kai, Sendai.
M imoto, K., 1986: Fossils in Kochi Prefecture.
Illustrations of the specimens kept by membersof the Kochi Fossil Society. 54 p. Kochi Fossil Society, Kochi. (in Japanese).
Noda, H., 1980: Molluscan fossils from the Ryukyu
Islands, southwestern Japan. Part I. Gastropoda and Pelecypoda from the Shinzato Formation in southeastern part of Okinawa-jima. Sci.
Rep., lnst. Geosci., Univ. Tsukuba, Sec. B, vol. I,
p. 1~95, pis. 1~12.
- - , 1988: Molluscan fossils from the Ryukyu
Islands, southwest Japan. Part 2. Gastropoda
and Pelecypoda from the Shinzato Formation in
the middle part of Okinawa-jima. Sci. Rep.,
Ins!. Geosci., Univ. Tsukuba, Sec. B, vol. 9, p.
29~85, pis. 5~ 19.
- - and Kikuchi, Y., 1980: Phanerolepida expansilabrum (Kuroda) (Gastropoda) from the
Miocene N antaisan Volcanic Breccia, Ibaraki
Prefecture, northeastern Japan. Venus, vol. 39,
no. 1, p. 69~ 73, pI. 1. (in Japanese with English
abstract).
- - and Ogasawara, K., 1976: On the variation ofPhanerolepida rehderi MacNeil. Contr., Dept.
Geol. Miner., Niigata Univ., no. 4, p. 431 ~436, pI.
1. (in Japanese with English abstract).
Okutani, T., 1964: Report of the archibenthal andabyssal Mollusca mainly collected from Sagami
Bay and adjacent waters by the R.V. Soyo-Maru
during the year 1955~ 1963. Jour. Fac. Sci.,
Univ. Tokyo, Sec. 2, vol. 15, p. 371~447.

933.

Intraspecific variation and heterochrony of Phanerolepida- - , 1966: Archibenthal and abyssal Mollusca collected by the R.V. Soyo-Maru from Japanesewaters during 1964. Bull. Tokai Reg. Fish.
Res. Lab., vol. 46, p. 1-32.
- - , 1968: Systematics, ecological distribution andpalaeoecological implication of archibenthal andabyssal Mollusca from Sagami Bay and adjacentareas. Jour. Fae. Sci.. Unil'. Tokyo. Sec. 2, vol.
17, pt. I, p. 1-98.
- - , 1969: Synopsis of bathyal and abyssal megaloinvertebrates from Sagami Bay and the south off
Boso Peninsula trawled by the RjV Soyo-Maru.
Bull. Tokai Reg. Fish. Res. Lab., vol. 57, p. 1-61.
Otuka, Y., 1949: Fossil Mollusca and rocks of theKiyosumi Group exposed at Minato-machi,
Chiba Prefecture, and its environs (1st paper).
Japan. Jour. Geol. Geogr., vol. 21, nos. 1-4, p.
295-309, pI. 13.
Ozaki, H., 1956: Some new and interesting molluscs
from Miocene Nobori Formation in the eastern

1037

Koti Prefecture, Japan. Bull. Natn. Sci. Mus..
Tokyo, vol. 3, no. I (no. 38), p. 1-6, pI. 1.
Shibata, H., 1974: Phanerolepida cf. expansilabrum
Kuroda. In, J. Itoigawa, H. Shibata, and H.
Nishimoto, Geology and Paleontology of
Mizunami City. Bull. Mizunami Fossil Mus.,no. I, p.43-203, pis. 1-63. (in Japanese).
Watanabe, K., Arai, J. and Hayashi, T., 1950:
Tertiary geology of the Chichibu basin. Bull.
Chiehibu Mus. Nat. Hist., no. I, p. 29-92, pis. 16. (in Japanese).
Watson, R.B., 1879: Mollusca of H.M.S. 'Challenger'
expedition. IV. Trochidae continued, viz. thegenera Basilissa and Troehus, and the Turbinidae, viz. the genus Turbo. Jour. Linn. Soc.
London, vol. 14, p.692-716. [not seen].
- - , 1886: Report of the Scaphopoda and Gastropoda collected by H.M.S. Challenger during
the year 1873-1876. Res. Voy. Challenger. Rep.
Sci.• Zoology, vol. 15, p. 1-722.

Hane 3fJfN. Kochi ~:ttl!, Muroto ~p, Nishinotani g§ /1:)-, Nobori E;, Okinawa rq,~g,
Saitama ~~. Shinzato ~1[, Tokigawa m~JII. Tonohama Jl!trA.
~~iJj!,O)!'f~*lCE;~.i£: Phanerolepida pseudotransenna Ozaki (.im:.@.ilJ:

1) "'" ? T""/-ij--tf'=fil-) O)fiP'l~~c~lF.fli: Phanerolepida pseudotransenna Ozaki O)JI:ilf:96~ ~ht.::. 75
j~lt*f:I:.~~rnH\uflit-:'~F~t-:.* ~ t.e~~>a:-~i"o .: o)~~li{fEI-(;$:r,,'O)~~96~O)Im~IF.fWlO)~~t-:.~IZSIi"."6 c m~ ~ h."6 o ~ff.:1¥J.i£:lli c ~O)~~O)OO-(;$:961:.J:O)~~O).Itti:iJ' I? Ne-

halemia hieroglyphiea .... Phanerolepida oregonensis .... P. pseudotransenna .... P. tansenna
c \" ? im1~*7IJiJ~Jt5E ~ ht.::.o -l" 0) ? t;" 1't.e < c HJJtlbO) 2 """:)O)fi53'1~li~1F.f1tO)t.t*-Ctl;."6 c1f;;tl?h."6 oPhanerolepida rehderi MacNeil >a:-OO-(;$:~~O).l:~~t-:.i1i\;":)I,,-C P. pseudotransenna O)~*.::. A C L- t.::.o ~ c f*"'? t.::. 4 """:)o)~iJ~1:..f,.!,IF.fO){ftlit-:.:ilf:\' ,tkfrn"C'.i£:lli L- t.::.o:/ /r,,'~~Ili-. HEHIIB~

Ryuichi Majima and Akihiro Murata

1038
Appendix.

Measurements, letter formula and remarks for all examined specimens of Phanerolepida
pseudotransenna Ozaki from the Nobori Formation.
Numberofwhorl

Letter formula ofshell-surface
ornamentation

4 1/2+
4+
4 1/2+

5+
3 1/2+

SRFsRFsFRFsF
SRF
SRF
SRF
SRF
SsRF
SRF?F
SRF
?F
?F
?F
?F
?RF
?R?R
?F
SRF?F
?F
?F?F?
?FR
?F?
?F
?F?
SRF
SRFRFRF
SRFsF
SRF
SRF
SRFRFRFsRFs
SRFRF
SRF
SRF
SR
SRFsRFsRF

12.4
12.4
11.1+
12.0+

4+
4+
3 1/2+5

SRF
SRF
SRFsFRFRF
SRFRFRFRF

12.1+
12.4
9.3+

6 1/2
3 1/2+4+

SRFRF
SRFsRsSRFsFSpecimen
IGSU

Locality
IGSU

Shell
height

Shell
width

Apertureheight

M37
M38
M39
M40
M41
M42
M43
M44
M45
M46
M47
M48
M49
M50
M51
M52
M53
M54
M55
M56
M57
M58
M59
M60
M61
M62
M63
M64
M65
M66
M67
M68
M69

62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-1
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-2
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3

17.0
17.8
17.7
16.9
16.4
15.6
17.8+
10.9+
12.3+
11.3+
11.0+
14.0+
9.9
14.7
15.1
15.8
12.7+
11.3+
8.4+
11.0+
8.8+
8.8+
16.3
13.5
15.8
14.2
15.9
17.2
14.8
17.2
17.0
14.1
16.3

20.0
20.3
19.8
20.2
19.0+
17.2+
17.7+
16.5+
16.9
15.3+
18.5
20.0+
15.0
17.1+
19.5+
18.4
17.0
15.8+
15.2+
19.1+
13.8+
11.2+
18.8
16.7
18.1
17.0
17.4
19.0
18.2
20.3
19.5+
18.0
17.8

12.4
13.1
12.4
12.3
10.8
11.4
12.5+

10.7
9.8
11.6
10.6
11.7
11.9
10.5
13.0
12.5
11.2
10.9

M70
M71
M72
M73

62-35-3
62-35-3
62-35-3
62-35-3

16.8
16.3
15.6+
15.0+

19.1+
19.6+
16.7+
17.1+

M74
M75
M76

62-35-3
62-35-3
62-35-3

16.3+
15.0
14.0

16.4+
17.6+
19.5

4+
3 1/2+
4 1/2+
3 1/2+

3+
11.8
9.8+
13.5
9.5+
8.0
12.2
10.1
11.9
11.5
11.0+
8.0+

1/2+
1/2+
1/2+
3+
3 1/2+

3+
4 1/2+
4 1/4
2+
2+
2 1/2+
1+
1/2+

3+
5+
4 1/2+
4 1/2+
3 1/2+
4+
4 1/2+

3+4+
3 1/2+

Remarks
Fig. 3-1
Fig. 3-2
Fig. 3-3

Deformed

Deformed
Deformed
Deformed
Deformed
Deformed
Deformed
Deformed
Deformed
Deformed
Deformed
Deformed
Fig.4
Figs. 3-4, 4, 5-4
Figs. 4, 5-8
Fig. 4
Fig. 4
Fig. 3-5,4, 5-2
Fig. 4
Fig. 4
Figs. 4, 5-10
Figs. 4, 5-11
Figs. 4, 8-4,
with operculum
Fig. 4
Fig. 4
Figs. 4, 5-3
Figs. 4, 5-1, 8-1,
with operculum
Figs. 4, 5-6
Figs. 4, 5-7
Fig. 4

933.

Intraspecific variation and heterochrony of Phanerolepida

SpecimenrGSU

Locality
rGSU

Shell
heightShell Apertureheight
width

M77
M78
M79
M80
M81
M82
M83
M84
M85
M86
M87
M88
M89
M90
M91
M92
M93
M94
M95
M96
M97
M98
M99
MIOO
MIOI
MI02
MI03
MI04
MI05
MI06
MI07
MI08
MI09
MIlO
Mill

62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3
62-35-3

15.4
14.5
16.2
16.6
14.9+
14.3+
14.0
14.2
14.7+
14.5
16.4
13.5
16.7
13.3+
14.2+
12.2+
13.1+
11.6+
11.4+
14.1+
12.7+
12.7+

17.3
18.4
19.3
19.5
17.3+
19.7+
17.8
18.8+
18.4+
16.5+
18.6
18.3+
18.1+
17.3+
15.1+
15.3+
17.5+
14.1+
13.4+
14.8+
18.4+
18.0+
21.4
15.4+
19.0+
12.7+
15.3+
13.4+
14.8+
11.2+
11.4+
15.0+
14.5+
13.3+
14.9+

11.7
11.9
12.8
13.1
11.6+
10.8+
11.3
10.2+
13.0
11.3
13.0
12.8+
13.8
10.6+
10.8+
10.0+
10.9+
8.9+
9.3+
10.2+

MI12

62-35-3

13.3+

9.5+

MI13
MI14
MI15
MI16

62-35-3
62-35-3
62-35-3
62-35-3

11.9+
19.2+
15.3+
10.5+

8.7+

13.3+
13.6+
11.9+
9.3+
15.0+
14.1+
12.5+
10.6+
13.9+
13.2+

11.6+
12.6+
7.1+

Numberofwhorl

Letter formula ofshell-surface
ornamentation

3

SRF
?RF
?FR?s
?RF
?RFsR?F
?F
SRFs
SRF?F
?FRF?F
S?s?s
SRF?F?F
?FR?F
SRF?F
?F
SRF
SR?
S?F?
SRF
SR?
SRF?F
SRF
?R?
?RF
SRFRFRF
?RF?F
?F
?F?F
?F
?RF
SR?F
SR?FRF?F
SRF?Fs
SRF?FsRF?
?F?
?F?F?

1/2+

3+
3
4

1/2+
1/2+

3+
3+
3 1/2+
3+2+
4+
3 1/2+
3 1/2+
3 1/2+
3+
5 1/2
4+
4+
6
4 1/2+
5+
3+

11.0
12.6
5
11.0+
11.2+
9.8+
8.3+
10.3+
8.9+

10.3+

9.5+
7.0+

1/2+

3+
2

1/2+

3+
3 1/2+
4+
4+
2 1/2+
2 1/2+
4 1/2+?F?
3 1/2+
3+
4+
3+

SRF?
S?F?F
SRF?F
?RF?

1039

Remarks
Fig. 4

Deformed
Deformed
Deformed
Fig. 4
Deformed
Fig. 5-12
Deformed
Deformed
Deformed
Fig. 4
Deformed
Figs. 4,5-9

Fig. 4
Deformed
Figs. 4, 5-5Deformed
Deformed
Deformed

Fig. 8-5,
with operculum
Fig. 8-3,
with operculum
Deformed

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1040-1069, 12 Figs., April 30, 1992

934. VERBEEKINID AND NEOSCHW AGERINID FUSULINACEANS FROM
THE AKIYOSHI LIMESTONE GROUP ABOVE THE PARAFUSULINAKAERIMIZENSIS ZONE, SOUTHWEST JAPAN*
KATSUMI UENO
Institute of Geoscience, University of Tsukuba, Ibaraki, 305

Abstract. The Akiyoshi Limestone Group above the Paraflislilina kaerimizensis Zone
in the Kaerimizu area can be divided into the Paraflislilina kaerimizensis, Afghanellaozawai, Neoschwagerina craticlilifera robusta, Verbeekina verbeeki-Afghanella schencki,
Neoschwagerina fusiformis, Verbeek ina verbeeki and Colania dOllvillei Zones in ascendingorder, mainly on the basis of the stratigraphic distribution of verbeekinid and neoschwagerinid fusulinaceans. In this paper, 10 species of verbeekinid and neoschwagerinid
fusulinaceans including one new subspecies, Neoschwagerina craticlilifera robusta, aredescribed. Also, the fusulinacean biostratigraphy of the Akiyoshi Limestone Group above
the Parafllslllina kaerimizensis Zone in the Kaerimizu area is briefly noted.
Key words. Akiyoshi Limestone Group, fusulinacean biostratigraphy, Middle Permian, Parafllslllina kaerimizensis Zone, Neoschwagerinidae, Verbeekinidae.

Concerning the biostratigraphic zonation byfusulinaceans in the Permian sequence of the
Akiyoshi Limestone Group, Ota (1977) subdivided it into II zones. Recently, Ozawa
and Kobayashi (1990) established 17
fusulinacean zones in the same part.
The fusulinacean family Verbeekinidae
arose from the family Ozawainellidae at the
beginning of the Middle Permian and thefamily Neoschwagerinidae was derived fromthe former in the middle of that period.
These two families flourished through Middle
Permian time. It is known that they havesignificant value for biostratigraphic zonationin the Middle Permian of the Tethyan realm.
About 30 species of the families Verbeekinidae and Neoschwagerinidae have beenrecorded from the Akiyoshi LimestoneGroup.
Very recently, I have discussed the origin
and early evolution of verbeekinid and neoschwagerinid fusulinaceans, based on biostratigraphic evidence obtained from theIntroduction
The Akiyoshi Limestone Group, covering
almost the whole area of the Akiyoshi Plateau situated in the central part of Yamaguchi
Prefecture, southwest Japan, is considered tohave formed as a large organic reef complex
during Late Paleozoic time (Ota, 1968).
This limestone contains rich shallow marine
fossil remains such as foraminifers, algae,
sponges, corals, brachiopods, mollusks andothers. Since Ozawa's fundamental work in
1923, fusulinacean foraminifers have often
been studied in connection with the biostratigraphic subdivision of the Akiyoshi Limestone Group, except for its lowermost part(e.g. Ozawa, 1925b; Toriyama, 1954,1958;
Hasegawa, 1958, 1963; Murata, 1961; Sakaguchi et al., 1966; Yanagida et al., 197 I ;Ota et al., 1973; Ota, 1977; Matsusue, 1986 ;Ueno, 1989; Ozawa and Kobayashi, 1990).
'Received September 17, 1991; accepted December
20, 1991
1040

934.

Kdma

Verbeekinids and neoschwagerinids from Akiyoshi(;)

aerimizu t ?<oAkiyoshiI2km...........

Figure 1. Map showing the Kaerimizu area in
the Akiyoshi Plateau.

Akiyoshi Limestone Group below the Misellina (M.) claudiae Zone in the Kaerimizu
area (Ueno, 1991 a, 1991 b).
This paper deals with the biostratigraphyof verbeekinid and neoschwagerinid
fusulinaceans from the Akiyoshi LimestoneGroup above the Parafusulina kaerimizensis
Zone in the Kaerimizu area, and 10 species ofthem including one new subspecies, Neoschwagerina craticulifera robusta, are described. In addition, the fusulinacean biostratigraphy of the Akiyoshi LimestoneGroup above the Parafusulina kaerimizensis
Zone is briefly noted.
Fusulinacean biostratigraphy
The Kaerimizu area is a large doline located in the northeastern part of the Akiyoshi
Plateau (Figure I). Ota (l977) divided the
Akiyoshi Limestone Group in this area into
the following 10 fusulinacean zones inascending order: the Fusulinella biconica,
Triticites simplex, Pseudofusulina vulgaris,
Pseudofusulina ambigua, Misellina claudiae,

1041

Parafusulina kaerimizensis, Afghanella
schencki, Neoschwagerina craticulifera, Verbeekina verbeeki and Colania douvillei
Zones.
I recognized seven fusulinacean biohorizons characterized by the first and last occurrence of some marker species in the stratigraphic interval between Ota's Parafusulina
kaerimizensis and Colania douvillei Zones(Figure 2). Each biohorizon is well recognized in the examined sections of the Kaerimizu area and defines the base of eachfusulinacean zone established herein.
The fusulinacean zones are distributed
almost concentrically in the Kaerimizu
Doline, being younger toward the bottom of
the doline, except for the southern part of thesurveyed area where the Afghanella ozawai
and Neosch wagerina craticulifera robusta
Zones are shown as the inlier in the Parafusulina kaerimizensis Zone (Figure 3).
Consequently, the Akiyoshi LimestoneGroup in this area lies almost horizontallyand is in reverse order.
A total of II species of verbeekinid andneoschwagerinid fusulinaceans were identified from the Akiyoshi Limestone Groupabove the Parafusulina kaerimizensis Zone in
the Kaerimizu area. Their stratigraphic distribution is shown in Figure 4. The
fusulinacean biostratigraphy of the Parafusulina kaerimizensis Zone through the
Colania douvillei Zone is summarized as follows.
Parafusulina kaerimizensis Zone: This
zone is defined as the stratigraphic interval
between the first occurrence of Parafusulina
kaerimizensis and that of Afghanella ozawai,
and is about 15 m in thickness. Stratigraphic relationship with the underlying Misellina (M.) claudiae Zone is conformable.
Well sorted foraminiferal sparitic limestone
and algal foraminiferal sparitic limestone are
predominant. The following fusulinaceans
are discriminated: Rauserella aff. breviscula
Sosnina, Rauserella sp., Minojapanella (Russiella) pulchra Miklukho-Maklay, Neo-

Katsumi Veno

1042

FU SULINACEAN

Z 0 N EBIOHORIZON

Coiania douvillei.... F. Colania douvillei

Verbeekina verbeeki
Neoschwag. fusiformis
Verbeekina verbeekiAfghanella schenck.i
Neoschwagerina
craticulifera robusta.... L. Neoschwagerina fusiform is &Pseudodoliolina pseudolepida.... F. Neoschwagerina fusiform is.... F. Verbeek ina

verbeeki &Afghanella schencki.... F.

Neoschwagerina craticulifera robusta.... F.

Afghanella ozawai

Afghanella ozawai
Paraf. kaerimizensis
Mise/lina (1.4.) c/audiae(PART).... F. Parafusulina kaerimizensis.... F. Neoschwagerina simplex.... F. "'aklaya pamirica
F.: First occurrence

L.: Last
Figure 2.

fusulinella giraudi Deprat, N. pseudogiraudi(Sheng), Neofusulinella sp., Pseudoreichelinasp., Pseudoendothyra spp., Parafusulina
kaerimizesis (Ozawa), P. lutugini (SchelIwien) and Chusenella alpina (KochanskyDevide and RamovS).
Afghanella ozawai Zone: Afghanella ozawai Zone, about IO m thick, conformably
overlies the Parafusulina kaerimizensis Zone.
Limestone facies of the present zone is similar
to that of the Parafusulina kaerimizensis
Zone. The fusulinacean fauna incl udes
Rauserella aff. breviscula Sosnina, Rauserellasp., Schubertella rara Sheng, Neofusulinella~

occurrence

Fusulinaeean zones and biohorizons above the Parafusulina kaerimizensis Zone.

giraudi Deprat, N. pseudogiraudi (Sheng),
Pseudoendothyra spp., Parafusulina kaerimizensis (Ozawa), Pseudodoliolina cf. ozawai
Yabe and Hanzawa, Afghanella ozawai Hanzawa and Afghanella sp.
Neoschwagerina craticulifera robusta
Zone: This zone is in conformable contact
with the subjacent Afghanella ozawai Zone
and is about 20 m in thickness. The lower
limit of the present zone is defined by the firstoccurrence of Neoschwagerina craticulifera
robusta, subsp. nov. Algal foraminiferal
sparitic limestone and foraminiferal micritic
limestone are predominant. The identified

Distribution offusulinacean zones and fossil localities in the Kaerimizu area. I: Misellina
2: Parafusulina kaerimizensis Zone (Pk), 3: Afghanella ozawai Zone (Ao),
4: Neoschwagerina craticutijera robusta Zone (Nc), 5: Verbeekina verbeeki-Afghanella schencki Zone
(V-A), 6: Neoschwagerina fusijormis Zone (Nf), 7: Verbeekina verbeeki Zone (Vv), 8: Cotania douvillei
Zone (Cd), 9: fossil locality studied herein.
Figure 3.

(M.) claudiae Zone (Me),934.

Verbeekinids and neoschwagerinids from Akiyoshi

1043

100m
I

(nnnnH~ 2
•

3

7

8

:::::::::::::::...............

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

6!TITIillJs•

9

Katsumi Ueno

1044

with a subordinate amount of algal micritic
limestone and muddy limestone. The
fusulinacean fauna includes Rauserella cf.
sphaeroidea Sosnina, Rauserella sp., Dunbarula schubertellaeformis Sheng, Dunbarulaspp., Schubertella sp., Neofusulinella sp.,
Staffella sp., Pseudoendothyra spp., Parafusulina edoensis (Ozawa), P. gigantea (Deprat), Parafusulina spp., Chusenella? cf.
pindingensis (Sheng), Chusenella sp., Verbeekina verbeeki (Geinitz), Pselldodoliolina
pseudolepida (Deprat), Afghanella schencki
Thompson and A. cf. sumatrinaeformis (Gubler).
Neoschwagerina fusiform is Zone: Neoschwagerina fusiformis Zone, about 10 m in
maximum thickness, is in conformable contact with the underlying Verbeekina
verbeeki-Afghanella schencki Zone. In thiszone, foraminiferal sparitic limestone is pre-fusulinaceans are as follows: Rauserella alf.
breviscula Sosnina, Rauserella spp., Dunbarula schubertellaeformis Sheng, Neofusulinella
pseudogiraudi (Sheng), Pselldoreichelina sp.,
Pselldoendothyra spp., Parafuslilina kaerimizensis (Ozawa), Chusenella referta Skinnerand Wilde, C. alpina (Kochansky-Devide andRamovs), C. schwagerinaeformis Sheng,
Pselldodoliolina cf. ozawai Yabe and Hanzawa, and Neosch wagerina craticulifera robllsta, subsp. nov.
Verbeekina verbeeki-Afghanella schencki
Zone: This zone, about 10 m in maximum
thickness, is defined as the stratigraphic interval between the first occurrence of both
Verbeekina verbeeki and Afghanella
schencki and that of Neoschwagerina fusiformis, and conformably overlies the Neoschwagerina cra tic lllifera robusta Zone.
Foraminiferal sparitic limestone is dominant

1>

<I

2

o

(')

<
<

1>

(")

c..

N

o

:J

CD

444
•

Maklaya pamirica•

Neoschwagerina

simplex
Parafusulina kaer imizensis

."

c:::

Afghanella ozawaienAfghanella sp.

c:::

Pseudodoliolina ct. ozawai
Verbeekina

verbeeki

Neoschwag. craticulifera
robusta--- ....... - - -Afghanella schencki
Afghanella ct. sumatrinaeformis
Pseudodoliolina pseudolepida
Neoschwagerinafusiform is

Neoschwagerina haydeni
Colania

douvillei

4 : Biohorizon
Figure 4. Stratigraphic distribution of verbeekinid, neoschwagerinid and some related fusulinaceans.
(abbreviations of zone names: see legend of Figure 3)

934.

Verbeekinids and neoschwagerinids from Akiyoshi

dominant with a subordinate amount of foraminiferal micritic limestone and muddy
limestone. The following fusulinaceans are
identified: Rauserella sp., Dunbarula
schubertellaeformis Sheng, Dunbarula spp.,
Schubertella sp., Neofusulinella sp.,
Pseudoendothyra spp., Parafusulina sp.,
Sch wagerina sp., Pseudodoliolina pseudolepida (Deprat), Neoschwagerina fusiformis
Skinner and Wilde, and N. haydeni Dutkevich and Khabakov.
Verbeekina verbeeki Zone: The lower
limit of the Verbeekina verbeeki Zone isdefined by the last occurrence of Neosch wagerina fusiformis and Pseudodoliolina
pseudolepida. In the same horizon, Verbeekina verbeeki appears again, which is diagnostic in the Verbeekina verbeeki-Afghanella
schencki Zone but barren in the Neoschwagerina fusiformis Zone. The thicknessof the present zone is about 10 m. A stratigraphic hiatus may exist between the Verbeekina verbeeki Zone and subjacent ones, judging from the distribution of fusulinaceanzones. The following fusulinaceans are discriminated: Rauserella sp., Dunbarula
schubertellaeformis Sheng, Dunbarula spp.,
Neofusulinella sp., Pseudoreichelina sp.,
Pseudoendothyra spp., Chusenella referta
Skinner and Wilde, and Verbeekina verbeeki(Geinitz).
Colania dOllvillei Zone: The Colania
douvillei Zone occupies the highest stratigraphic position in the Kaerimizu area and only
crops out at the bottom of the doline. It is atleast 5 m in thickness and in conformablecontact with the Verbeekina verbeeki Zone.
The lower limit of the present zone is definedby the first occurrence of Colania douvillei.
The upper limit, however, can not be
confirmed in the Kaerimizu area. Algal
sparitic limestone and muddy limestone aredominant. The present zone is rather poorin foraminifers and only two species offusulinaceans, Pseudoendolhyra sp. andColania dOllvillei (Ozawa), are identified.
The geologic age of the Parafusulina kaer-

1045

imizensis Zone through the Colania douvillei
Zone is considered to be the Murgabian of
the standard stratigraphic scheme in the Tethyan province proposed by Leven (1980).

Systematic paleontology
All specimens identified in this paper aredeposited in the paleontological collectionsof the Akiyoshi-dai Museum of Natural
History. The abbreviation ASM is the prefixof the specimen repository number.
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea vonMoller, 1878
Family Verbeekinidae Staff
and Wedekind, 1910
Subfamily Verbeekininae Staff
and Wedekind, 1910
Genus Verbeekina Staff, 1909
Type species. - Fusulina verbeeki Geinitz,
1876.

Verbeekina verbeeki (Geinitz, 1876)
Figures 5-1-6,6-1-5

Fusulina I'erbeeki Geinitz, 1876, p.399-400.
Schwagerina l'erbeeki (Geinitz). Staff, 1909, p.506,
pI. 7, figs. 5, 7; Deprat, 1912, p. 40-41, pI. I, figs.
7-11; Colani, 1924, p. 108-113, pI. 15, fig. 14, pI.
18, figs. 2,5,16-21,25-27, pI. 19, figs. 1-7, II, 13.
Verbeekina verbeeki (Geinitz). Ozawa, 1925b, p. 4851, pI. 10, figs. 6-7; Lange, 1925, p. 260-261, pI.
3, fig. 66a, pI. 5, fig. 66b; Ozawa and Tobler,
1929, p. 48, pI. 5, fig. 5; Chen, 1934, p. 101-102,
pI. 16, fig. I ; Gubler, 1935, pI. I, figs. 10-16;
Thompson, 1936, p. 197-200, pI. 24, figs. 1-8 ;H uzimoto, 1936, p. 10 I-I 04, pI. 9, fig. 8, pI. 21,
figs. 1-3; Thompson, 1948, pI. 16, figs. 7-12
(same as pI. 24, figs. 1-2,4,6-8 of Thompson,
1936, p. 197-200); Kochansky-Dcvide andRamovs, 1955, p. 389,416, pI. 4, figs. 1-3; Chen,
1956, p.9,47-48, pI. 9, figs. 5-6, pI. 13, figs. 1-2;
Miklukho-Maklay, 1957, p. 113-114, pI. 3, fig. 2;
Kobayashi, 1957, p.301-302, pI. 9, figs. 5-7 ;Toriyama, 1958, p.205-208, pI. 37, figs. 1-6, pI.
38, figs. 1-6; Rauser-Chcrnoussova and Fursen-

1046

Katsul17i Vena

Figure S. 1- 6. VerbeekinG verbeeki (Geinitz), la, 2- 4 : ax ia l sections, ASM25156, ASM25162,
ASM25161 , ASM25155 , 5,6 : sagitta l secti o ns, ASM25164, ASM25165 , x 10, lb : en larged part of la, x 40.

934.

Verbeekinids and neoschwagerinids from Akiyoshi

ko, 1959, pI. 10, fig. 5; Nogami, 1961, p.167169, pI. 2, figs. 1-4; Sheng, 1963, p. 85-86, 215216, pI. 26, figs. 1-5; Hanzawa and Murata,
1963, pI. 4, fig. 8; Igo, 1964, p. 62-63, pI. 2, fig.
I ; Ishii and Nogami, 1964, p. 24, pI. 8, figs. 4-6;
Pasini, 1965, pI. 16, fig. I ; Leven, 1965, p. 140,
pI. 4, fig. 1 ; Ishii, 1966, pI. 6, fig. 9; Sheng, 1966,
p. 138-139, pI. 23, fig. I (same as pI. 13, fig. 1 ofChen, 1956, p. 9,47-48); Skinner and Wilde,
1966, pI. 17, fig. 6; Igo, 1967, pI. 6, figs. 1-2 ;Leven, 1967, p. 205, pI. 38, fig. I ; Choi, 1970, p.
348-349, pI. 15, figs. 1-3; Ozawa, 1970a, pI. 2,
fig. I; Tien, 1970, p.35-37, pI. 6, figs. 1-10;
Toriyama and Pitakpaivan, 1973, p. 50-53, pI. 6,
figs. 1-5; Zhang and Wang, 1974, p. 293, pI. 152,
fig. 3; Toriyama, 1975, p.66-68, pI. 14, fig. 12;
Rozovskaya, 1975, pI. 27, figs. 6-7; Sheng andSun, 1975, p.47, pI. 2, fig. 13; Ozawa, 1975, pI.
10, figs. 10-11 ; Toriyama and Kanmera, 1977, p.
8-9, pI. I, figs. 8-10; Lin et at., 1977, p.83-84,
pI. 25, fig. 9; Ota, 1977, pI. I, figs. 5-6; Liu etat., 1978, p. 81, pI. 18, fig. 17; Ding, 1978, p. 288,
pI. 100, fig. 2; Chen and Yang, 1978, p. 105, pI.
27, fig. 3; Kobayashi, 1979, pI. 4, figs. 9-10;
Hasegawa et at., 1979, pI. 85, figs. 1-5,7; Tien,
1979, p. 130-132, pI. 21, figs. 3-9, pI. 22, fig. I B,
pI. 23, fig. 3B; Kahler and Kahler, 1979, p. 215217, pI. 2, fig. 4, pI. 10, figs. 1-2, 4-5; Wang etat., 1981, p.58-59, pI. 19, fig. 6; Wang et at.,1982, p. 99, pI. 27, figs. 1-2; Xie, 1982, p. 66, pI.
32, fig. 3(same as pI. 25, fig. 9 of Lin et at., 1977,
p. 83-84); Sun et aI., 1983, p. 33, pI. 9, fig. 5 ;Chen, 1984, p.64, pI. 17, figs. 1-2; Yang, 1985,
pI. 2, fig. 34; Tien, 1986a, pI. 7, fig. 13; Tien,
1986b, pI. 12, fig. II ; Ishibashi, 1986, p. 105-107,
pI. 15, figs. 2-8, pI. 16, figs. 1-3; Wang and Tang,
1986, pI. 2, fig. 16; Kobayashi, 1986, p. 145, pI. 4,
fig. 8, pI. 9, fig. 6; Wang and Zhou, 1986, pI. 2,
figs. 1-2; Han el at., 1987, pI. 2, figs. 1-2,6;
Loeblich and Tappan, 1988, pI. 298, figs. 1-5(same as pI. 24, figs. 1-2,6-8 of Thompson, 1936,
p. 197-200); Kobayashi, 1988a, p.8, pI. 11, figs.
10-14; Kobayashi, 1988b, p.443, figs. 5-3 -4;
Tien, 1989, pI. 33, figs. 7-8; Ozawa and Kobayashi, 1990, pI. 10, fig. 11.
Verbeekina verbeeki forma A, Pitakpaivan, 1965, p.
53-54, pI. 6, fig. 5.

Material.-Axial sections; ASM25157
from Loc. SF77, ASM25158 from Loc. SF79,
ASM25155, ASM25156 from Loc. SF80,
ASM25160 from Loc. SA 120, ASM25161,
ASM25162, ASM25163 from Loc. SA 152.
Sagittal sections; ASM25164 from Loc.
SA96, ASM25165 from Loc. SA152. Axial

1047

section of polyvalent individual; ASM25159
from Loc. T A256.
Description. - Shell large and almost spherical with very shallow umbilicus. Mature
specimens having 13 to 15 volutions, 5.40 to
7.20 mm in length and 5.15 to 7.00 mm inwidth, with a form ratio of about 1.00.
Inner 3 to 5 vol utions eostaffelloid andtightly coiled with rotated axis of coiling insome case. Beyond the fifth volution, shell
becomes spherical and loose rather rapidly.
Chamber height almost constant throughout
length of shell. Radius vectors of the first to
twentieth volutions of well oriented axial
section (Figure 5-1) 0.05, 0.08, 0.12, 0.20, 0.34,
0.60,0.88, 1.19, 1.56, 1.91, 2.23 and 2.51 mm,
and form ratios 0.50,0.53,0.67,0.92,0.97,
1.05, 1.06, 1.09, 1.05, 1.03, 1.03 and 1.04,
respectively.
Proloculus spherical and exceedingly minute for shell size. Its outside diameter varies
from 0.030 to 0.060 mm, averaging 0.045 mm
for 8 specimens.
Spirotheca thin and composed of a single
structureless layer in inner 3 volutions, but ofa tectum and lower fibrous alveolar keriotheca beyond the fourth volution. Alveolar
keriotheca finer than that observed in neoschwagerinids. Thickness of spirotheca of
the first to thirteenth volutions of abovementioned specimen 0.010, 0.010, 0.010, 0.020,
0.015,0.025,0.030,0.040,0.055,0.065,0.090,
0.065 and 0.050 mm.
Septa thin, unfluted throughout shelllength and bending anteriorly. They arecomposed of the same elements as the spirotheca. Average septal counts of the first to
thirteenth vol utions for 3 specimens 6, 9, 11,
11, 8, 8, 10, 12, 18, 19, 25, 30 and 36, respectively.
Parachomata numerous, small and low,and semicircular in transverse section. They
are well developed in inner juvenile volutions, but sporadically and inconstantly observed in some outer ones, because they
develop only on both sides adjacent to septa
and decrease in height rather rapidly toward1048

Katsumi Vena

the cente r of the cham bers.

Remarks. - Verbeekina verbeeki (Geinitz)
is one of the most fami liar verbeekinids in the
Middle Permian of the Tethyan province.

The Akiyoshi specimens are quite simil ar to
the topotypes restudied and redescri bed byThompson (1936).
In the Kaerimizu area, the distribution of

Sb
Figure 6. 1- 5. Verbeekina verbeeki (Geinitz), 1- 4: axia l sections, ASM25l57, ASM25l58 , ASM25l60,
ASM25163 , ASM25159 , Sa: axi a l section o f pol yva lent indi vidu a l, X 10, 5b: enlarged pa rt of Sa, X 40.

934.

Verbeekinids and neoschwagerinids from Akiyoshi

Verbeekina verbeeki (Geinitz) is stratigraphically separated into two intervals, namely
the Verbeekina verbeeki-Afghanella sclzencki
Zone and the Verbeekina verbeeki Zone.
Most of the individuals found in the former
zone are generally smaller in size and havefewer volutions than those in the latter one,
although other important morphologicalcharacters are in close agreement.
Occurrence.-Abundant in the Verbeekina
verbeeki-Afghanella schencki and Verbeekina verbeeki Zones.
Subfamily Pseudodoliolininae Leven, 1963
Genus Pseudodoliolina Yabeand Hanzawa, 1932
Type species. - Pseudodoliolina ozawai
Yabe and Hanzawa, 1932.

Pseudodoliolina pseudolepida(Deprat, 1912)
Figures 7-1- 10
Doliolina pseudolepida Deprat, 1912, p. 46, pI. 5, figs.
6-9, pI. 6, fig. 4.
Doliolina lepida mut. pseudolepida Deprat, Deprat,
1913, p. 50.
Doliolina lepida (Schwager). Ozawa, 1927, p. 152153, pI. 45, figs. 1-2 (not pI. 37, figs. Ie, 7b, 8b,9b); Gubler, 1935, p. 95-99, pI. 4, figs. 3-10, II?,
12?, pI. 5, figs. 14-15; Ciry, 1941, pI. I, fig. 2.
Pseudodoliolina pseudolepida (Deprat). Thompson
and Foster, 1937, p. 141-142, pI. 25, figs. 2-4;
Thompson, 1948, pI. 17, figs. 3-4 (same as pI. 25,
figs. 2,4 of Thompson and Foster, 1937, p. 141142); Kochansky-Devide and Ramov~, 1955, p.
416-417, pI. 4, figs. 7-10, pI. 8, fig. 6; Toriyama,
1958, p.211-212, pI. 39, figs. 20-25; Nogami,
1961, p.163-165, pI. I, figs. 7-11; Ishii and
Nogami, 1961, pI. 25, fig. 7 (same as pI. I, fig. 7 ofNogami, 1961, p. 163-165); Suyari, 1962, p.32,
pI. 12, fig. 10; Hanzawa and Murata, 1963, p. 23,
pI. 20, figs. 1-5; Sheng, 1963, p. 97-98,229-230,
pI. 29, figs. 1-9; Sheng, 1966, p. 143, pI. 26, fig.
4; Ozawa, 1970a, pI. 3, figs. 7-8; Tien, 1970, p.
39-40, pI. 7, figs. 6-9, pI. 8, figs. 1-4; Toriyama
and Pitakpaivan, 1973, p. 53-57, pI. 6, figs. 6-13 ;Zhang and Wang, 1974, p. 294, pI. 152, fig. 8 ;Sheng and Sun, 1975, p.50, pI. 14, figs. 1-2;
Ozawa, 1975, pI. 10, fig. 13; Toriyama and Kan-

1049

mcra, 1977, p. 12-14, pI. I, figs. 13-18, pI. 2, figs.
1-6; Lin el al., 1977, p.89, pI. 27, fig. II; Liuel al., 1978, p.83, pI. 20, fig. 5; Ding, 1978, p.
288, pI. 98, fig. 6; Toriyama and Kanmcra, 1979,
p. 54-56, pI. 8, figs. 7-13; Hasegawa el al., 1979,
pI. 83, figs. I, 2,4; Ticn, 1979, p. 132-134, pI. 20,
figs. 4-6; Kahler and Kahler, 1979, p. 224-225,
pI. 3, fig. 3; Sun et al., 1983, p.37, pI. 9, fig. 3;
Chen, 1984, p. 68, pI. 19, fig. 4; Yang, 1985, pI. 2,
fig. 28; Tien, 1986a, pI. 7, fig. II (same as pI. 20,
fig. 5 ofTien, 1979, p. 132-134); Tien, 1986b, pI.
11, fig. 10, pI. 12, figs. 1-3, 5D, liB; Han et al.,1987, pI. 2, fig. 4; Kobayashi, 1988a, p. 10, pI. 5,
figs. 18-21, pI. 6, fig. 18; Kobayashi, 1988b, p.
443-445, figs. 6-1 - 8; Tien, 1989, pI. 32, figs. 67; Ozawa and Kobayashi, 1990, pI. 10, fig. 8.
Pseudodoliolina cf. pseudolepida (Deprat). Kawano,
1961, p. 105-107, pI. 10, figs. 8-10.
Pselldodoliolina aIr. pselldolepida pseudolepida (Dcprall. Kanmera, 1963, p. 108-109, pI. 14, figs.
15-18.
Pselldodoliolina ozawai Yabe and Hanzawa. Chen
and Yang, 1978, p. 112, pI. 30, figs. 4-5.
Pseudodoliolina sp., Hasegawa et al., 1979, pI. 83, figs.
3,5 -7; Tien, 1989, pI. 32, figs. 8-10 (9: same as
pI. 11, fig. 1 of Tien, 1986b).

Material. - Axial sections; ASM25167 from
Loc. AL169, ASM25166 from Loc. ALl80.5,
ASM25168 from Loc. KN4, ASM25169 from
Loc. T A26l, ASM25170 from Loc. SA 119,
ASM25171 from Loc. EB98, ASM25172 from
Loc. MT253. Sagittal sections; ASM25l73,
ASM25174 from Loc. T A263. Tangential section; ASM25175 from Loc. KS201.
Description. - Shell medium to large forgenus, cylindrical with broadly roundedpolar regions and convex lateral slopes.
Mature specimens of II to 14 volutions, 4.55
to 6.23 mm in length and 1.90 to 3.10 mm inwidth. Form ratio ranges from 1.97 to 2.51,
averaging 2.20 for 16 specimens.
Shell expands gradually through growthwith an almost straight axis of coiling. The
first volution spherical, but later ones become
cylindrical rather rapidly. Radius vectors of
the first to thirteenth volutions of illustrated
axial section (Figure 7-1) 0.11,0.17,0.21,
0.27,0.32,0.41,0.49,0.59,0.71,0.85,1.06,1.25
and 1.41 mm, and form ratios 1.00,1.97,2.10,
2.30,2.63,2.71,2.80,2.83,2.83,2.74,2.50,2.31
and 2.17, respectively.

1050

Katsumi Vena

13

"'4 •

15

Figure 7. I- tO. Pseudodoliolina pseudolepida (Oeprat) , 1-3,5 - 8: axial sections, ASM25170 ,ASM25169 , ASM25167 , ASM25171 , ASM25166, ASM25168, ASM25172 , 4,9 : sagitta l sections, ASM25173,
ASM25174, 10 : tangential section, ASM25175 . 11, 12. Colania douvillei (Ozawa), 11: diagonal section ,ASM25213 , 12 : axial section , ASM25212. 13- 15. Pseudodoliolina cf. ozawai Yabe and Han zawa, 13:
tangential sect ion, ASM25177 , 14 : oblique section , ASM25178, 15 : sag itta l section, ASM25176. All
X 10.

934.

Verbeekinids and neoschwagerinids from Akiyoshi

Proloculus commonly spherical but rarely
irregular. Its outside diameter varies from
0.150 to 0.300 mm, averaging 0.211 mm for 22
specimens.
Spirotheca thin and composed of a single
structureless layer in inner few vol utions, but
consists of a tectum and a lower, less dense
layer in outer ones. Alveolar structure notobservable even in outermost volution.
Thickness of spirotheca of the first to thirteenth volutions of above-mentioned specimen 0.010,0.005,0.010,0.010,0.010,0.015,
0.015,0.015,0.010,0.020,0.020,0.020 and
0.025 mm.
Septa numerous and slightly bending anteriorly. They consist of the same elements as
the spirotheca.
Parachomata narrow and high, well developed in all volutions, and attain about Yz to% of chamber height.
Remarks. - Pseudodoliolina pseudolepida(Deprat) is one of the most common speciesof the genus and widespread in the Middle
Permian of the Tethyan province. The Akiyoshi specimens are quite identical with
Deprat's original ones.
This species is easily distinguished from
Pseudodoliolina ozawai Yabe and Hanzawa
by a larger shell and greater number of volutions.
Hasegawa et al. (1979) reported Pseudodoliolina sp. from the Verbeekina heimi Subzone (equivalent to the Verbeekina verbeekiAfghanella schencki Zone in this study) in
the Kaerimizu area. Judging from theirillustrations, Pseudodoliolina sp. has a slightly smaller proloculus than P. pseudolepida(Deprat). However, this difference is hardly
sufficient to warrant the specific separation ofthese forms.
Occurrence. - Abundant in the Verbeekina
verbeeki-Afghanella schencki and Neosch wagerina fusiformis Zones.

Family Neoschwagerinidae Dunbar
and Condra, 1927

1051

Subfamily Neoschwagerininae Dunbar
and Condra, 1927
Gen us Neosch wagerina Yabe, 1903
Type species. - Schwagerina craticulifera
Schwager, 1883.

Neosch wagerina craticulifera robusta
Ueno, subsp. nov.
Figures 8-1-10, 10-6-11

Material. - Axial section of the holotype;
ASM25179 from Loc. T A246. Axial sectionsof paratypes; ASM25180 from Loc. T A246,
ASM25181 from Loc. AL98, ASM25182 from
Loc. ALIOO, ASM25183 from Loc. ALIOI,
ASM25184 from Loc. ALl08, ASM25185
from Loc. KS-AD7, ASM25186 from Loc.
KS-AD9, ASM25187 from Loc. KS158,
ASM25188 from Loc. NS254. Sagittal sections of paratypes; ASM25189 from Loc.
TA246, ASM25190 from Loc. TA247,
ASM25191 from Loc. T A250, ASM25192
from Loc. EB68, ASM25193 from Loc.
KSI55.5. Slightly oblique axial section ofparatype; ASM25194 from Loc. ALl03.
Diagnosis. - Large Neoschwagerina having
an inflated fusiform shell with bluntly
pointed polar ends, almost straight or slightly
convex lateral slopes, very small proloculus,
and low, broad and fan-shaped primary transverse septula. Inner few volutions skewcoiled, outer ones planispiral. Parachomata
narrow and high.
Description. - Shell large for genus andinflated fusiform with bluntly pointed polarends. Lateral slopes almost straight orslightly convex. Mature specimens having
16 to 19 volutions, 4.50 to 5.95 mm in lengthand 3.55 to 4.85 mm in width. Form ratio
ranges from 1.06 to 1.44, averaging 1.27 for 15
specimens.
Inner 2 or 3 volutions tightly and skewcoiled. Beyond the third or fourth volution,
shell expands gradually with a straight axis ofcoiling. Radi us vectors of the first to seventeenth volutions of the holotype 0.05,0.08,

1052

K alsumi Vena

934.

Verbeekinids and neoschwagerinids from Akiyoshi

0.12,0.17,0.22,0.29,0.39,0.51,0.67,0.77,0.91,
1.05,1.25,1.43,1.64, 1.83 and 1.97 mm, andform ratios 0.90, 1.03, 1.25, 1.24, 1.45, 1.62,
1.62, 1.57, 1.5 I, 1.58, 1.52, 1.49, 1.43, 1.43, 1.38,
1.36 and 1.38, respectively.
Prolocul us very small and spherical. Itsoutside diameter ranges from 0.040 to 0.090
mm, averaging 0.054 mm for 17 specimens.
Spirotheca thin to medium, composed of atectum and alveolar keriotheca. Thicknessof spirotheca of the first to seventeenth vol utions of the holotype 0.010,0.010,0.015,
0.015,0.015,0.030,0.040,0.065,0.055,0.030,
0.020, 0.030, 0.050, 0.020, 0.030, 0.020 and
0.025 mm. Primary transverse septula low,broad and fan-shaped, and well developed
beyond the fourth volution. They are observed immediately above each parachomata
and abut tops of parachomata adjacent to
septa. No secondary transverse septula
developed even in outermost volution.
Septa composed of downward deflectionsof tectum and alveolar keriotheca. Septal
counts of the second to seventeenth vol utionsof typical sagittal section of para type (Figure8-8) 3?,9, 14, 17,20, 19,22,21,26,26,25,26,
29, 30, 34 and 33, respectively. Axial septula
present beyond the sixth volution. Twoaxial septula occasionally inserted between 2
adjacent septa in outer volutions.
Parachomata high, rather narrow, andbegin to occur in the third volution. They
attain a height about % that of chambers.
Remarks. - Neosch wagerina cra ticulifera
robusta, su bsp. nov. is closely similar to N.
craticulifera (s.s.), especially in the nature of
the primary transverse septula and parachomata, but differs from the latter in having
a larger shell, more vol utions and a smaller
form ratio. The present new subspecies also
resembles Neoschwagerina margaritae Deprat, but differs from the latter in having asmaller shell and a different shell shape. The

1053

morphological characters of Neoschwagerina
craticulifera robust a, subsp. nov. suggest thatthe present new subspecies is phylogeneticallyintermediate between N. craticulifera (s.s.)and N. margaritae Deprat.
Measurements. - See Table I.
Occurrence.-Abundant In the Neoschwagerina craticulifera robusta Zone.

Neoschwagerina fusiformis
Skinner and Wilde, 1967
Figures 9-1- 14

Neoschwagerina fusiformis Skinner and Wilde, 1967,
p. 16-17, pI. 19, figs. 1-8; Ding, 1978, p. 288289, pI. 98, fig. 3; Wang et al., 1981, p. 64-65, pI.
18, figs. 1-2, II ; Chen, 1984, p.69, pI. 19, fig. 3(same as pI. 98, fig. 3 of Ding, 1978, p. 288-289).

Material. - Axial sections; ASM25195,
ASM25196, ASM25197 from Loc. KN2,
ASM25198, ASM25199A,B from Loc. KN4,
ASM25200 from Loc. KN21, ASM25201,
ASM25202 from Loc. T A265, ASM25203A,B
from Loc. AL168. Sagittal sections;
ASM25204 from Loc. KN2, ASM25205 from
Loc. KN5, ASM25206 from Loc. ALI80.
Description. - Shell medium to large forgenus and fusiform to elongate fusiform withbluntly pointed poles and almost straight toslightly convex lateral slopes. Mature specimens of 14 to 18 volutions 4.65 to 7.90 mm inlength and 2.70 to 4.70 mm in width. Formratio ranges from 1.43 to 1.90, averaging 1.65
for 17 specimens.
Axis of coiling straight through growth
except for inner few volutions of some specimens. Radius vectors of the first to fifteenth
vol utions of typical axial section (Figure 912) 0.08, 0.11,0.13,0.18,0.24,0.29,0.35,0.42,
0.51, 0.69, 0.79,0.97, 1.13, 1.29 and 1.46 mm,
and form ratios 0.53,0.76, 1.15, 1.66, 1.49,\.69,1.89, 2.02, 1.98, 1.71, 1.99, 1.84, 1.77, 1.78
and 1.75, respectively.

Figure 8. 1-10. Neoschwagerina craticulifera robusta. subsp. nov., 6a: axial section of the holotype,
ASM25179, 1,2,4, 5, 9, 10: axial sections of para types, ASM25184, ASM25180, ASM25181, ASM25188,
ASM25186, ASM25182, 3,7,8: sagittal sections of paratypes, ASM25189, ASM25191, ASM25190, X 10, 6b :enlarged part of 6a, X 40.
+----

o
V>
+>-

Measurements of Neoschwagel'ina craticulifera I'obusla, subsp. nov. F.R.: From ratio, D.P.: Diameter of proloculus (in mm).

Table 1.

Radius vector
Reg. no.
I

ASM25179

2

ASM25180
ASM25181
ASM25183
ASM25184

3
4
5
6
7

ASM25187
ASM25186

Figure Length Width
8-6
8-2
8-4
10-9
8-1
10-10
8-9

5.69

4.20

5.53
5.50
5.60

3.85
4.05
4.35

1.35
1.44
1.36
1.29

5.10
5.18
5.15

4.05
4.00
4.85

1.26
1.30
1.06

-

-

D.P.

F.R.

0.060
0.065
-

0.050
0.050
0.045
0.045

I

2

3

4

5

6

7

8

9

10

II

12

13

14

15

16

17

18

0.05
0.06

0.08

0.12

0.29
0.28

0.39
0.37

0.51
0.49

0.67
0.61

0.91
0.92

-

CUI

0.52
0.44
0.52

0.65
0.57
0.64

1.86
1.88
1.78

1.99

2.19

0.10
0.11

0.13
0.15

0.40
0.34
0.40
0.34
0.42

1.65
1.68

0.10
0.11

0.22
0.19
0.21
0.17

1.47
1.49
1.40

-

0.17
0.13
0.15

1.25
1.29
1.27
1.22

1.97

0.07
0.07
0.08
0.06

1.05
1.10
1.11

1.83

0.05
0.04
0.05
0.05
0.05

0.77
0.77
0.81

1.64

0.12
0.12

0.22
0.21

1.43

0.09

0.17
0.16

0.46
0.53

0.57
0.67

1.27
1.15
1.29

1.46
1.31
1.46

1.82
1.71
1.86

2.03
1.92
2.18

2.39

0.08

0.20

0.25
0.30
0.25
0.29

0.72
0.75
0.70
0.81

0.95
0.89
0.88
0.83

1.04
1.08
1.00

0.95

1.12

1.55
1.64
1.46
1.67

-

-

-

-

2

3

4

5

6

7

8

9

10

II

12

13

14

15

16

17

18

I
2
3

0.90
0.73

1.24
0.94
1.47
1.15
1.00
1.00
1.13

1.45
1.09
1.52
1.33
1.27
1.28

1.62
1.28
1.56
1.58
1.16

1.62
1.45
1.63
1.66
1.37
1.31
1.35

1.57
1.43
1.52
1.78
1.36
1.28
1.40

1.51
1.44
1.57
1.66
1.32
1.34

1.58
1.42
1.60
1.58
1.44

1.52
1.48
1.56
1.46
1.42

1.43
1.41
1.52
1.34
1.39

1.30
1.31

1.43
1.38
1.47
1.36
1.30
1.24

1.38
1.39
1.48
1.40
1.29

1.33
1.32

1.49
1.44
1.58
1.40
1.37
1.28
1.23

1.36
1.40
1.44
1.36
1.26
1.24
1.17

-

0.60
0.70
0.60
0.70

1.25
0.71
1.25
0.85
0.87
0.75
0.87

1.38

4
5
6
7

1.13
0.56
1.28
0.64
0.76
0.77
0.75

1.28

1.31
1.50

1.37

1.25
1.21

1.20

1.27
1.18

19

-.
~

-

~
::::

-

1.23
1.10

Cl

-

1.09

-

---

Thickness of spirothcca

I

I

2

3

4

5

6

7

8

9

10

II

12

13

14

15

16

17

18

0.010

0.015
0.010

0.015

0.015

0.030

0.040
0.025

0.065

0.055

0.025

-

0.020
0.025
0.030

0.040
0.025

0.030
0.050

0.030
0.040

0.080
0.040

0.040
0.040

0.030
0.040

0.070
0.030

0.030
0.040
0.045

0.020

0.030

0.050
0.045

0.020

0.055

0.020
0.060

0.030

0.020
0.030

0.030
0.045

0.055

0.030

0.025

0.060
0.030

0.025

0.020
0.030

0.060
0.050
0.040

0.030
0.035

0.030
0.040

0.035

0.025
0.030

0.025
0.025
0.040

0.030

2

0.010
0.005

3
4

0.005
0.005

5
6
7

0.005
0.005
0.010

0.005
0.005
0.005
0.005

0.010
0.010

0.010
0.015

0.010

0.010
0.010

0.010
0.010
0.015

0.010

0.015

0.020

0.015

0.020

0.015
0.015

0.020
0.015

0.020
0.015

0.025
0.020

0.020

0.045

0.030
0.035

0.025

0.040

0.030

-

-

0.040

-

0.075
0.020

0.035
CUl25

>::

s:::,

c;:;

s:::

-

1.21

2.52

"-----.- - -From ratio
I

-

19

19

0.035

0.040

0.030

-

0.050

0.030

0.040

-

934.

Verbeekinids and neoschwagerinids from Akiyoshi

Proloculus small and spherical. Its outside
diameter varies from 0.030 to 0.130 mm, averaging 0.081 mm for 21 specimens.
Spirotheca rather thin for genus and composed of a tectum and alveolar keriotheca.
Thickness of spirotheca of the first to fifteenth
volutions of specimen mentioned above 0.005,
0.005,0.005,0.010,0.025,0.030,0.030,0.025,
0.030,0.030,0.025,0.025,0.025,0.035 and
0.040 mm. Primary transverse septula thin
and well developed beyond the third volution. They occur immediately above eachparachomata and abut tops of parachomata
adjacent to septa. One secondary transverse
septulum rarely and rather inconstantly inserted between 2 adjacent primary transverse
septula.
Septa thin, bending anteriorly and composed of downward deflections of spirothecal
element. Septal counts of the third to sixteenth volutions of typical sagittal section(Figure 9-2) 12, 14, 15, 15, 16, 17, 24, 23, 27,
27, 30, 28, 30? and 28? Axial septula begin
to occur in the fifth vol ution. One axial
septulum inserted between 2 adjacent septa ininner volutions, but usually 2, rarely 3 inouter ones.
Parachomata narrow and high, reaching
about % the height of chambers.
Remarks. - Neoschwagerina fusiformis was
originally described by Skinner and Wilde(1967) from the Permian of Tunisia. The
holotype specimen (Skinner and Wilde, 1967,
pI. 19, fig. I) has a minute proloculus andskew-coiled inner volutions, and is considered to be a microspheric form of the species.
Most of the Akiyoshi specimens are quite
similar to one of the paratypes (Skinner and
Wilde, 1967, pI. 19, fig. 2).
This species somewhat resembles Neoschwagerina haydeni Dutkevich and Khabakov in its essential morphological characters,
especially in the nature of septula and parachomata. However, the former can be distinguished from the latter in having a moreelongate shell and larger form ratio.
Occurrence. - Abundant In the Neo-1055

schwagerina fusiformis Zone.
Neoschwagerina haydeni Dutkevich
and Khabakov, 1934
Figures 10-1 - 5

Neoschwagerina craticulifera (Schwager).

Hayden,1909, p. 248-249, pI. 21, figs. 1-7.

Neoschwagerina craticulifera haydeni Dutkeyich andKhabakoy, 1934, p.94-99, pI. 2, figs. 6-8, pI. 3,
figs. 1-2; Thompson, 1946, p.155-156, pI. 23,
figs. 12-13; Thompson, 1948, pI. 20, figs. 3-4(same as pI. 23, figs. 12-13 of Thompson, 1946);
Kanmera, 1957, pI. 19, fig. 24 (same as pI. 23, fig.
12 of Thompson, 1946); Toriyama, 1958, p. 220223, pI. 41, figs. 7-8 (not pI. 41, fig. 6); Suyari,
1962, p. 35-36, pI. 11, fig. 5; Ishizaki, 1962, p.
173-175, pI. 12, figs. 4-6.
Neoschwagerina haydeni Dutkeyich and Khabakoy.
Honjo, 1959, p. 147-149, pI. 3, fig. 10; Minato
and Honjo, 1959, pI. 3, fig. 2; Sheng, 1963, p.
103-104,236-237, pI. 32, figs. 1-8; Zhang andWang, 1974, p. 292, pI. 151, fig. 7; Chen andYang, 1978, p. 116, pI. 32, figs. 3-5; Liu et al.,1978, p. 89, pI. 21, fig. 6; Toriyama andKanmera, 1979, p.77-80, pI. 12, figs. 19-22, pI.
13, figs. 1-8; Hasegawa et al., 1979, pI. 84, figs.
1-8; Wang et al., 1981, p. 63, pI. 15, figs. 1-2,5,
7; Wang et al., 1982, p. 106, pI. 30, fig. 7; Sunet al., 1983, p. 39, pI. 10, fig. 13; Lin, 1984, p.
171-172, pI. 14, fig. 12; Chen, 1984, p. 69, pI. 21,
fig. 2; Yang, 1985, pI. 2, fig. 12; Kobayashi,
1988b, p.445-447, figs.7 - 2-11; Ozawa andKobayashi, 1990, pI. 10, figs. 12-14.
Neoschwagerina cf. haydeni Dutkeyich and Khabakoy. Ding, 1978, p. 289, pI. 101, fig. 2.

Material. - Axial sections; ASM25207
from Loc. ALl7S, ASM2520S from Loc.
ALISO, ASM25209 from Loc. KN21,
ASM25210 from Loc. SAI4S. Sagittal section; ASM25211 from Loc. KN 19.
Description. - Shell medium to large forgenus and thickly fusiform with roundedpolar ends and almost straight lateral slopes.
Mature specimens having 15 to 17 volutions,5.10 to 6.00 mm in length and 3.S0 to 4.15 mm
in width, giving form ratios of 1.31 to 1.54.
Axis of coiling straight except for inner few
volutions of some specimens. Shell expands
gradually through growth. Radius vectors of
the first to sixteenth vol utions of illustrated

1056

Katsumi Vena

934.

Verbeekinids and neoschwagerinids from Akiyoshi

axial section (Figure 10-1) 0.06,0.10,0.14,
0.20,0.26,0.35,0.43,0.52,0.63,0.76,0.88, 1.03,
1.16, 1.31, 1.46 and 1.66 mm, and form ratios
0.58,0.74,0.81, 1.10, 1.29, 1.31, 1.53, 1.63, 1.66,
1.61, 1.59, 1.53, 1.54, 1.58, 1.53 and 1.50,
respectively.
Proloculus small and spherical, being 0.060
to 0.070 mm in outside diameter averaging
0.063 mm for 3 specimens.
Spirotheca thin for genus and composed ofa tectum and alveolar keriotheca. Thicknessof spirotheca of the first to sixteenth volutionsof above-mentioned specimen 0.005, 0.005,
0.010,0.015,0.015,0.020,0.015,0.020,0.020,
0.035,0.035,0.030,0.025,0.025,0.025 and
0.030 mm. Thin primary transverse septula
first appear in the third or fourth vol uti on.
They develop immediately above each parachomata and abut tops of parachomata adjacent to septa. One secondary transverse septulum sometimes inserted between 2 adjacent primary ones.
Septa thin, bending anteriorly and composed of downward deflections of tectum andalveolar keriotheca. Axial septula begin tooccur in the fifth volution. Beyond the seventh volution, 2 axial septula commonly
observed. Three axial septula rarely inserted
between 2 adjacent septa in outer few volutions.
Parachomata narrow and high adjacent to
septa, but low and semicircular in central partof chamber.
Remarks. - Neoschwagerina haydeni Outkevich and Khabakov somewhat resembles N.
craticulifera robusta, subsp. nov., describedabove, in shell shape. However, the former
can be easily distinguished from the latter inhaving a thinner spirotheca, more slendertransverse septula and parachomata.
Occurrence. - Common in the Neoschwagerina fusiformis Zone.

1057

Subfamily Lepidolininae
Miklukho-Maklay, 1958
Genus Colania Lee, 1934 emend.
Ozawa, 1970
Type species. - Colania kwangsiana Lee,
1934.
Colania douvillei (Ozawa, 1925)
Figures 7-11-12

Neoschwagerina globosa (Yabe). Douville, 1906, p.
182, pI. 17, pI. 18, figs. 1-2; Deprat, 1912, p.51,
pI. 4, figs. 1-2 (not pI. 4, figs. 3-4); Deprat, 1913,
p. 55; Deprat, 1914, p. 29-30; Ozawa, 1922, p.
368-372; Colani, 1924, p. 152-153, pI. 23, figs.
1-2,4-14,22-34,36-38, pI. 25, figs. 9, 13, pI. 26,
figs. 3-5.
Neosch wagerina douvillei Ozawa, 1925a, pI. 3, fig.
6, pI. 4, fig. 5; Ozawa, 1925b, p.55-57, pI. II,
figs. 5-7; Huzimoto, 1936, p.114-115, pI. 23,
figs. 1-5; Toriyama, 1947, p. 78-79, pI. 17, fig. 8 ;Chen, 1956, p. 12,58-59, pI. 13, figs. 3-4, 6-7 (notpI. 13, fig. 5, pI. 14, fig. 7); Toriyama, 1958, p.
223-227, pI. 41, figs. 9-13, pI. 42, figs. 1-6; Sakagami, 1958, p.92-93, pI. 14, figs. 7-10;
Kanuma, 1960, p. 70- 71, pI. 13, figs. 5- 7 ;Nogami, 1961, p. 180-183, pI. 5, figs. 1-5; Sada,
1961, p. 123-125, pl. 12,figL 1-8; Sheng, 1963, p.
102-103,235-236, pI. 33, figs. 1-7; Hanzawa and
Murata, 1963, pI. 4, figs. 3-4, pI. 17, figs. 1-10;
Sheng, 1966, p. 146, pI. 25, fig. I ; Sheng and Sun,
1975, p.51-52, pI. 13, figs. 11-14; Lin el al.,
1977, p. 92, pI. 28, fig. 12; Liu el al., 1978, p. 89,
pI. 24, fig. I ; Han, 1980, p. 90, pI. 34, figs. 13-15 ;Xie, 1982, p. 69, pI. 34, fig. I ; Wang et al., 1982,
p. 106, pI. 30, fig. 5.
Neosch wagerina cf. douvillei Ozawa. Kawano, 1960,
pI. 25, figs. 12-13; Kawano, 1961, p. 107-108, pI.
II, figs. 2-5; Yoshimura, 1961, pI. 3, fig. 4 ;Suyari, 1962, p. 36, pI. 10, fig. 14.
Colania douvillei (Ozawa). Ozawa, 1970a, pI. 7, figs.
8-10; Choi, 1972, p.371-372, pI. 46, figs. 1-6;
Toriyama, 1975, p. 103, pI. 20, figs. 22-26; Toriyama and Kanmera, 1977, p. 17-24, pI. 2, figs. 717, pI. 3, figs. 1-16; Ota, 1977, pI. I, figs. 3-4;
Sakagami and Miyama, 1988, fig. 1-15; Kobayashi, 1988a, p. 13, pI. 7, figs. 7-9, pI. 8, figs. 1-8 ;Ozawa and Kobayashi, 1990, pI. II, figs. 1-2.
Gifuella douvillei (Ozawa). Hasegawa el al., 1979,+-- Figure 9. 1-14. Neoschwagerina fusiformis Skinner and Wilde, 1,3-7,9-11, 12a, 13: axial sections,
ASM25203A, ASM25198, ASM25196, ASM25195, ASM25197, ASM25200, ASM25203B, ASM25201,
ASM25199A, ASM25199B, ASM25202, 2,8,14: sagittal sections, ASM25204, ASM25205, ASM25206, X 10,
12b: enlarged part of 12a, X 40.

1058

Katsumi VenD

Figure 10. 1- 5. Neoschwagerina haydeni Dutkevich and Khabakov, 1,2,4,5: ax ia l sect io ns,
ASM25209 , ASM25208 , ASM2521O, ASM25207 , 3 : sag itta l section , ASM252 ll. 6- 11. Neoschwagerina
craticulifera robusta, subsp. nov. , 7,9, 10 : ax ia l sect io ns of para types, ASM25l85 , ASM25l83 , ASM25 l 87 , 6,
11: sag itta l sections of paratypes, ASM25 l 93, ASM25 l92, 8: sli ghtly ob liqu e axial secti o n of paratype,
ASM25l94. A ll X 10.

934.

Verbeekinids and neoschwagerinids from Akiyoshi

pI. 86, figs. 1-6.
not Neoschwagerina dOllvillei Ozawa. Gubler, 1935,
p. I I I-I 13, pI. 6, fig. 2, pI. 7, figs. 7-8,10-1 I, pI.
8, fig. 6; Ishii, 1966, pI. 5, fig. 4; Igo, 1967, pI. 7,
figs. 6-8, pI. 8, fig. 7; Tien, 1970, p. 48-49, pI. I I,
figs. I - I 5, pI. 12, figs. 1-13 ; Wang et aI., 198 I, p.
62-63, pI. 18, fig. 5; Yang, 1985, pI. 2, fig. 14.
not Colania dOllvillei (Ozawa). Ozawa, 1970b, p.
35-38, pI. 3, figs. 1-10, pI. 4, figs. 1-10, pI. 5, figs.
1-5, pI. 6, figs. 1-9; Tien, 1979, p. 137- 139, pI.
22, figs. 1- 10, pI. 23, figs. 1-2; Tien, 1986a, pI. 8,
fig. 2 (same as pI. 22, fig. 2 of Tien, 1979); Tien,
1986b, pI. I I, figs. 1-6, 9A; Tien, 1989, pI. 33,
figs. 1-6(2-3: same as pI. I I, figs. 5, I of Tien,
1986b).
not? Neoschwagerina dOllvillei Ozawa. Chen andYang, 1978, p. 117, pI. 32, figs. 7-9.

1059

Taishaku, Atetsu and Omi. The present
specimens agree closely with ones reported
from these limestone plateaus.
Occurrence. - Rare in the Colania douvillei
Zone.
Subfamily Sumatrininae Silvestri, 1933
Genus Afghanella Thompson, 1946
Type species. - Afghanella schencki
Thompson, 1946.

Afghanella ozawai Hanzawa, 1954
Figures I I-I - 17

Material. - Axial section; ASM25212.
Diagonal section; ASM25213. Both from
Loc. EBIII.
Description. - The following description isbased on one axial section (Figure 7-12).
Shell large and inflated fusiform withbroadly rounded polar ends and straight axisof coiling. Length and width of shell having
13 volutions, 6.80 mm and 4.55 mm, respectively. Form ratio about 1.50. Shell
expands gradually through growth. Radius
vectors of the first to twelfth volutions 0.22,
0.33,0.44,0.54,0.67,0.79,0.96,1.16,1.36, 1.56,
1. 74 and 1.95 mm, and form ratios 0.91, 1.15,
1.22,1.28,1.37,1.48,1.52,1.48,1.57, 1.49, 1.53
and 1.51, respectively. Prolocul us somewhat
irregular, being 0.210 mm in outside diameter.
Spirotheca thin and composed of a tectum
and alveolar keriotheca. Thickness of spirotheca of the first to twelfth volutions 0.015,
0.030, 0.020, 0.025, 0.020, 0.025, 0.040, 0.030,
0.025,0.035,0.035 and 0.040 mm. Primarytransverse septula thin and begin to appear in
the second volution. No secondary transverse septula present. Parachomata narrowand high, meeting the tops of primary transverse septula adjacent to septa.
Remarks. - Colania douvillei (Ozawa) iswidespread in the Middle Permian rocks inthe eastern part of the Tethyan province, andalso in the huge limestone masses in the Inner
Zone of southwest Japan, such as Akiyoshi,

Neoschwagerina (Yabeina) schellwieni (Deprat).
Ozawa, 1925a, pI. 2, figs. 8- 10.
Yabeina schellwieni (Deprat). Ozawa, 1925b, p.6061, pI. 10, figs. 3a, 4 (4: same as pI. 2, fig. 9 of
Ozawa, 1925a).
Cancellina schellwieni (Deprat). Ozawa, 1927, p.
161, pI. 34, fig. 18 (same as pI. 2, fig. 9 of Ozawa,
1925a), pI. 44, fig. I b, pI. 45, fig. 3.
Afghanella ozawai Hanzawa, 1954, p. 3-7, pI. I, figs.
1-6, pI. 2, figs. 1-4; Kanmera, 1957, pI. 19, figs.
12-13; Toriyama, 1958, p.252-254, pI. 48, figs.
I 1,13-14 (not pI. 48, figs. 12, 15-17), Kawano,
196 I, p. I 17- I 19, pI. 15, figs. 3-11 ; Hanzawa and
Murata, 1963, pI. 2, figs. 1-2; Ozawa, I 970a, pI.
6, figs.6-7 (same as pI. 19, figs.12-13 ofKanmera, 1957); Lin et al., 1977, p. 95, pI. 30,
fig. 3; Hasegawa et al., 1979, pI. 82, figs. I I - I 6 ;Xie, 1982, p. 72, pI. 35, fig. 6 (same as pI. 30, fig.
3 of Lin et al., 1977, p. 95); Kobayashi, 1988a,
p. 14, pI. 5, figs. 22-25; Ozawa and Kobayashi,
1990, pI. 10, figs. 1-2.
Afghanella schencki Thompson. Toriyama, 1958, p.
250-25 I, pI. I?, 2,4-5, 10 (not pI. 48, figs. 3,6-9) ;Ota, 1977, pI. 2, figs. 1-2.

Material. - Axial sections of megalospheric
form; ASM25214 from Loc. EB49,
ASM25215, ASM25216 from Loc. EB50,
ASM25217, ASM25218 from Loc. EB56,
ASM25219, ASM25220 from Loc. T A231,
ASM25221 from Loc. MT210, ASM25222
from Loc. NS220, ASM25223 from Loc.
SF49, ASM25224A from Loc. KS-AD24.
Sagittal sections of megalospheric form;
ASM25224B from Loc. KS-AD24, ASM25225
from Loc. SF33, ASM25226 from Loc. AL92,

1060

Katsumi Vena

934.

Verbeekinids and neoschwagerinids from Akiyoshi

ASM25227 from Loc. EB49. Axial sections of
microspheric form; ASM25228 from Loc.
SA56, ASM25229 from Loc. AL90.
Description. - Shell small to medi urn for
genus and inflated fusiform to fusiform withbl untly pointed poles. Dimorphism wellobserved. Megalospheric specimens having
7 to 9 volutions, 2.55 to 4.20 mm in lengthand 1.53 to 2.53 mm in width. Form ratio
ranges from 1.38 to 1.99, averagi ng 1.67 for 20
specimens. Microspheric form of 9 to 10
volutions 2.95 to 4.55 mm in length and 1.40
to 2.30 mm in width, giving form ratios of1.98 to 2.11.
Axis of coiling straight throughout inmegalospheric form. Inner 2 or 3 volutions
tightly and skew coiled in microspheric form.
Radius vectors of the first to seventh volutionsof well oriented axial section of megalospheric form (Figure 11-2) 0.15,0.22,0.32,0.44,
0.58, 0.73 and 0.92 mm, and form ratios 1.40,
1.64, 1.62, 1.66, 1.72, 1.67 and 1.65, respectively.
Proloculus of megalospheric form large forshell size, spherical to somewhat irregular,
being 0.165 to 0.320 mm in outside diameter,
averaging 0.239 mm for 49 specimens.
Proloculus of microspheric form minute andspherical, being 0.030 to 0.040 mm in outside
diameter, averaging 0.036 mm for 5 specimens.
Spirotheca thin to medium, composed of atectum and alveolar keriotheca. Thickness of
spirotheca of the first to seventh vol utions ofabove-mentioned specimen 0.010,0.025,
0.040,0.025, 0.070?, 0.050 and 0.030 mm.
Primary transverse septula developed in all
volutions. Their tips slightly swollen.
Lower edges of primary transverse septula incontact with tops of parachomata adjacent to1061

septa. One secondary transverse septulum
inserted between 2 adj acent primary transverse septula in outer volutions.
Septa consist of downward deflections oftectum and anterior and posterior extensionsof keriotheca. Lower half of septa usuallysolidified and pendant-shaped in cross section. Septal counts of the first to eighth
volutions of one illustrated sagittal section(Figure 11-4) 8, 18,23,23,24,26, 28? and
31? Axial septula begin to appear in thesecond or third volution. Two axial septula
usually inserted between 2 adjacent septa.
Parachomata massive and triangular in transverse section, rarely narrow and high, andwell developed in all volutions.
Remarks. - Small shell size and the natureof axial and transverse septula in Afghanellaozawai Hanzawa indicate that it is one of themost primitive species in this genus. The
stratigraphic position of the present species
also supports this conclusion.
Afghanella ozawai Hanzawa somewhat
resembles A. robbinsae and A. african a, both
described by Skinner and Wilde (1967) fromthe Permian of Tunisia. However, the former differs from the latter two in having athicker spirotheca and fewer axial septula
between 2 adjacent septa.
Occurrence. - Abundant in the Afghanellaozawai Zone.

Afghanella schencki Thompson, 1946
Figures 12-1 - 11

Afghanella schencki Thompson, 1946, p. 153-155, pI.
25, figs. 1-12; Thompson, 1948, pI. 18, figs. 1-5(same as pI. 25, figs. 1-3,7,10 of Thompson,
1946, p. 153-155); Chen, 1956, p. 14,67-68, pI.
12, figs. 4-9; Toriyama, 1958, p. 250-251, pI. 48,
figs. 3, 6-9 (not I?, 2, 4-5,10); Sheng, 1958, p.

+-- Figure 11. 1-17. Afghanella ozawai Hanzawa, 1-3,5,7,9,10,12-15: axial sections of megalospheric form, ASM25217, ASM25215, ASM25224A, ASM25218, ASM25220, ASM25214, ASM25219,
ASM25216, ASM25221, ASM25222, ASM25223, 4,6,8,11: sagittal sections of megalospheric form,
ASM25226, ASM25225, ASM25227, ASM25224B, X 10, 16a, 17: axial sections of microspheric form,
ASM25229, ASM25228, X 20, 16b: enlarged part of 16a showing minute proloculus and skew coiled
juvenile volutions, X 100. 18-20: Afghanella sp., tangential sections, ASM25230, ASM25231,
ASM25232, X 10.

1062

Katsumi Vena

Figure 12. 1- 11. Afghanella schencki Thompson , 1,2,4- 8,10,11 : axial sect ion s, ASM25238 ,ASM25235A, ASM25235B, ASM25234, ASM25240, ASM25236, ASM25237, ASM25239, ASM25233, 3,9:
sagittal sections, ASM25242, ASM25241. 12- 16. Afghanella cf. sumatrinaeformis (Gubler), 12- 15: axial
sections, ASM25244, ASM25245, ASM25246, ASM25243, 16: sagitta l section, ASM25247. All X 10.

277, 289- 290, pI. 4, figs. 9- 10 ; Sheng, 1963, p.
109-110, 244-245, pI. 36, fig. 20 ; Pasini, 1965, pI.
17, fig. l a, Ib (same as pI. 25, fig. 12, 7, ofThompson , 1946, p. 153-155); Sheng, 1966, p. 151 , pI.
26, fig. 6; Leven, 1967, p. 198, pI. 36, figs. 4, 7- 8,
pI. 37, fig . 3; Zhang and Wang, 1974, p. 291 , pI.
151 , fig. I ; Rozovskaya , 1975, pI. 33, figs. 5- 7
(same as pI. 25, figs. 2- 3, 7 of Thompson, 1946, p.
153- 155); Lin et al. , 1977, p.95 , pI. 30, fig. 4 ;Chen and Yang, 1978, p. 121 , pI. 34, figs. 3- 4 ;Liu el al. , 1978, p. 96, pI. 23, fig. 4; Kahler andKahler, 1979, p. 251 - 252, pI. 8, fig. 5, pI. 9, fig. 2 ;Hasegawa et al., 1979, pI. 82, figs. 5- 10; Da andSun, 1983, p. 113, pI. 30, fig. 3 ; Sun et al., 1983,
p. 39- 40, pI. 10, fig. 5 ; Chen, 1984, p. 71 - 72, pI.
22, fig. I ; Sun and Zhang, 1985, p. 506, pI. I, figs.

7, 9- 10 ; Yang, 1985, pI. 3, fig. 21 ; Loeblich andTappan , 1988, pI. 306, figs. 3- 6 (same as pI. 25,
figs. 2- 3, 6- 7 of Thompson, 1946, p. 153- 155) ;Ozawa and Kobayashi , 1990, pI. 10, figs. 5- 7.
Afghanella ozawai Han zawa. Toriyama, 1958, p.
252-254, pI. 48 , figs. 12, 15 - 17 (not pI. 48, figs. II ,13- 14).
Afghanella schencki schencki Thompson. Toriyama and Kanmera , 1979, p. 58 - 61 , pI. 9, figs. 824.

Material. - Axial sections; ASM25233
from Loc. MT240, ASM25234 from Loc.
MT243, ASM25235A ,B from Loc. SA 119,
ASM25236 from Loc. SAI20, ASM25237

934.

Verbeekinids and neoschwagerinids from Akiyoshi

from Lac. SA 130, ASM25238 from Lac.
SF69, ASM25239 from Lac. T A254,
ASM25240 from Lac. EB97. Sagittal sections;
ASM25241 from Lac. SA 130, ASM25242
from Loc. MT239.
Description. - Shell small to medi urn for
genus and elongate fusiform with bluntly
pointed polar regions and almost straight toslightly convex lateral slopes. Mature shellof 9 to 11 volutions 3.33 to 4.55 mm in lengthand 1.50 to 2.35 mm in width. Form ratio
ranges from 1.74 to 2.37, averaging 1.96 for 18
specimens.
Shell expands gradually through growth.
Axis of coiling straight throughout. Radius
vectors of the first to ninth vol utions of typical axial section (Figure 12-1) 0.12, 0.16, 0.25,
0.33, 0.43, 0.55, 0.68, 0.83 and 0.99 mm, andform ratios 1.48,1.81,1.80, 1.85,2.00, 1.96,
1.96, 1.94 and 1.86, respectively.
Prolocul us small and spherical. Its outside diameter varies from 0.070 to 0.195 mm,
averaging 0.121 mm for 19 specimens.
Spirotheca thin and consists of a tectumand probably very thin alveolar keriotheca.
Thickness of spirotheca of the first to ninth
vol utions of specimen mentioned above 0.0 15,
0.025, 0.020, 0.015, 0.025, 0.015, 0.020, 0.025
and 0.025 mm.
Primary transverse septula very thin anddeveloped in all volutions. Tips of primarytransverse septula slightly swollen and incontact with tops of parachomata adjacent to
septa. One secondary transverse septul urn
inserted between 2 adjacent primary transverse septula in outer 3 or 4 volutions.
Septa thin and bending anteriorly. Tips ofsepta solidified and pendant-shaped. Septal
counts of the first to tenth volutions of illustrated sagittal section (Figure 12-3) 9,14,16,
18,16,18,21,19,21 and 26, respectively.
Axial septula first appear in the fifth volution.
Two, rarely 3 axial septula observed in outer3 or 4 volutions. Parachomata narrow andhigh.
Remarks. - Afghanella schencki was originally described by Thompson (1946) from1063

the Bamian Limestone of Afghanistan. The
proloculus in the Akiyoshi specimens isslightly smaller than that in the original ones.
Afghanella schencki Thompson can beeasily distinguished from A. ozawai Hanzawa
in having a more elongate shell, more volutions, smaller prolocul us, thinner spiro thecaand more developed primary transverse andaxial septula. The stratigraphic distributionof these two species is also different.
Occurrence. - Abundant in the Verbeekina
verbeeki-Afghanella schencki Zone.

Afghanella cf. sumatrinaeformis(Gubler, 1935)
Figures 12-12-16

CompareNeaschwagerina annae (Volz). Hayden, 1909, p.
250-251, pI. 22, figs. 8-14.
Neascll\vagerina suma/rinae/armis Gubler, 1935, p.
123-127, pI. 5, figs. 3-4,10,17, pI. 7, fig. 2.
Suma/rina suma/rinae/armis (Gubler). Ciry, 1941,
pI. I, figs. 8-9.
A/ghanella sumalrinae/armis (Gubler). Kochansky-Devide and Ramov5, 1955, p.391-392,417,
pI. 5, figs. 1-10; Chen, 1956, p.14,68-69, pI. 7,
figs. 9-11 ; Kanmera, 1957, pI. 19, figs. 16-17(same as pI. 5, figs. 3-4 of Gubler, 1935, p. 123127); Sheng, 1966, p. 152, pI. 22, fig. II (same as
pI. 7, fig. 9 of Chen, 1956, p. 14, 68-69); Leven,
1967, p. 198-199, pI. 37, figs. 1-2; Ozawa, 1970a,
pI. 6, figs. 10-11 (same as pI. 5, figs. 3-4 of Gubler, 1935, p. 123-127); Tien, 1970, p. 55, pI. 13,
figs. 9-10, pI. 14, figs. 5-9; Rozovskaya, 1975, pI.
33, figs. 8-9 (same as pI. 5, figs. 3-4 of Gubler,
1935, p. 123-127); Toriyama and Kanmera,
1979, p.63-68, pI. 10, figs. 14-24; Yang, 1985,
pI. 3, fig. 22; Loeblich and Tappan, 1988, pI.
306, figs. 7-9 (same as pI. 5, figs. 3-4, 10 of Gubler, 1935, p. 123-127).
A/ghanella cf. schencki Thompson. Kanmera, 1957,
pI. 19, figs. 14-15.

Material. - Axial sections; ASM25243
from Lac. T A256, ASM25244 from Lac.
SA 122, ASM25245 from Loc. SA 124,
ASM25246 from Loc. MT238. Sagittal section; ASM25247 from Loc. MT239.
Description. - Shell medi urn to large forgenus and thickly fusiform with bluntlyKatsumi Ueno

1064

pointed to rounded axial regions and slightly
convex lateral slopes. Mature specimens having 11 to 13 volutions, 4.05 to 4.98 mm inlength and 2.58 to 3.50 mm in width, giving
form ratios of 1.26 to 1.53. Shell expands
gradually through growth. Proloculus small
to medium, being 0.090 to 0.175 mm in outside diameter, averaging 0.124 mm for 4 specimens. Spirotheca thin and composed of atectum and thin alveolar keriotheca. Primary transverse septula observed in all volutions. Tips of primary transverse septula
solidified and pendant-shaped, joining withtops of parachomata. One, rarely 2 secondary transverse septula inserted between 2
adjacent primary transverse septula. Septa
thin but usually coated with dense calcareousmaterial. Tips of septa slightly swollen. One
axial septulum usually occurs between 2 adjacent septa beyond the fifth volution, but 2
axial septula rarely inserted in outer ones.
Parachomata prominent and well developedin all volutions.
Remarks. - The present species quite agrees
with Afghanella sumatrinaeformis (Gubler)in the essential shell characters except forhaving a smaller proloculus.
Afghanella cf. sumatrinaeformis (Gubler)differs from A. schencki Thompson in having
a more thickly fusiform shell, smaller form
ratio and more volutions.
Occurrence. - Common in the Verbeekina
verbeeki - Afghanella schencki Zone.

Afghanella sp.
Figures 11-18-20

Material. -Tangential sections
ASM25230 from Loc. KS 175, ASM25231
from Loc. AL89, ASM25232 from Loc. EB56.
Remarks. - Three tangential sections arereferable to Afghanella sp. This unidentified species resembles Afghanella ozawai
Hanzawa, which coexists with it, in the natureof the transverse septula. However, the former has a larger and more elongate shell thanthose of the latter. At any rate, the exact

specific identification is postponed until better material is accumulated.
Occurrence. - Rare in the lower part of the
Afghanella ozawai Zone.

Acknowledgments
I would like to express my deep appreciation to Prof. Hisayoshi Igo of the Universityof Tsukuba for critical reading of the manuscript. Thanks are also extended to Prof.
Sumio Sakagami ofChiba University, Messrs.
Akihiro Sugimura and Takehiko Haikawa of
the Akiyoshi-dai Museum of Natural History
for their contin uous advice and encouragement, and Mr. Yasuyuki Miyama of
Sumitomo Cement Co., Ltd. for fruitful discussion on fusulinacean taxonomy.

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1065

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1066

Katsumi Vena

Kahler, F. and Kahler, G., 1979: Fusuliniden (Foraminiferen) aus dem Karbon und Perm vonWestanatolien und dem Iran. Mitt. OSterr. geol.
Ges., vol. 70 (1977), p. 187-269, pis. 1-10.
Kanmera, K., 1957: Revised classification of Cancellina, Neoschwagerina, and evolution of
Sumatrininae and Neoschwagerininae. Mem.
Fac. Sci., Kyushu Univ., ser. D, vol. 6, no. I, p.
47-64, pis. 19-20.
- - , 1963: Fusulines of the Middle Permian Kozaki
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11-19.
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 II.
Paleontology, no. 4. Ozawainellinae, Schubertellinae, Verbeekininae and Neoschwagerininae.
Bull. Tokyo Gakugei Univ., vol. II, p. 55-83, pis.
10-13.
Kawano, M., 1960: Some fusulinids from the Aralani
Conglomerate in the northeastern part of
Yamaguchi City. Sci. Rep., Tohoku Un iv. , 2ndser. (Geol.), Spec. Vol., no. 4, p. 223-230, pis. 2425.
- - , 1961: Stratigraphical and paleontologicalstudies of the Paleozoic formations in the westernpart of the Chugoku Massif. Bull. Fac. Educ.,
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II, spec. no., p. 1-133, pis. 1-15.
Kobayashi, F., 1979: Petrography and sedimentary
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Japan. Jour. Geol. Soc. Japan, vol. 85, no. 10,
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C, vol. 12, no. 4, p. 131-163, pis. 1-10.
- - , 1988a: Middle Permian foraminifers of the Omi
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- - , 1988b: Late Paleozoic foraminifers of the
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Japan, N.S., no. 150, p.435-452.
Kobayashi, M., 1957: Paleontological study of the
Ibukiyama Limestone, Shiga Prefecture, central
Japan. Sci. Rep., Tokyo Kyoiku Daigaku, sect.
e, vol. 5, no. 48, p.247-331, pis. 1-10.
Kochansky-Devide, V. and Ramovs, A., 1955: Die
Neoschwagerinenschichten und ihre Fusulinidenfauna bei Bohinjska Bela und Bled (Julische
Alpen, Siowenien, NW-Jugoslawien). Siov.
A kad., 4, p. 359-424, pis. 1-8.
Lange, E., 1925: Eine mittelpermische Fauna von

Guguk Bulat (Padanger Oberland, Sumatra).
Verh., Geol.-mijnb. Genootsch. Nederland en
Kolonien., geol. ser., Deel 7, p. 157-239, pis. 1-5.
Leven, E Ya., 1965: 0 stratigraficheskom znachenii
roda Polydiexodina Dunbar et Skinner, 1931.
Voprosy Mikropaleont., vol. 9, p. 129-146, pis. 16. (in Russian)- - , 1967: Stratigrafiya i fuzulinidy permskikh otlozheniy Pamira. Akad. Nauk SSSR, Trudy
Geol. Inst., vol. 167, p. 1-224, pis. 1-39. (in Russian)- - , 1980: Ob'yasniitel'naya zapiska k stratigraficheskoy shkale permskikh otlozheniy oblasti
Tetis. 51 p., Minist. Geol. SSSR, Vses. Orden a
Lenina Nauchno-Issled. Geol. Inst. (VSEGEI),
Leningrad. (in Russian)Lin, J.X., 1984: Protozoa. In, Yichan Inst., Geol.
Mineral Resour. chiefly ed., Biostratigraphy of
the Yangtze gorge area, (3), Late Paleozoic Era,p. 110-177, pis. 1-16. Geol. Publ. House, Beijing.
(in Chinese with English abstract and description of new species)- - , Li, J.X., Chen, G.X., Zhou, Z.R. and Zhang, B.
F., 1977: Fusulinida. In, Hubei. Inst., Geol.
Sci. et al. eds., Paleontological Atlas of Central
South China, no. 2 (Late Paleozoic), p. 4-96, pis.
1-30. Geol. Publ. 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., Paleontological
A tlas of South west China, Guizhou vol., no. 2(Carboniferous to Quaternary), p. 12-98, pis. 124. Geol. Publ. House, Beijing. (in Chinese)
Loeblich, A.R., Jr. and Tappan, H., 1988: Foraminiferal genera and their classification. 970
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Matsusue, K., 1986: Foraminiferal biostratigraphy ofthe lower part of the Akiyoshi Limestone Group.
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Miklukho-Maklay, A.D., 1957: Nekotorye
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Minato, M. and Honjo, S., 1959: The axial septula ofsome Japanese Neoschwagerininae with special
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Murata, M., 1961 : On the geological structure of the
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Verbeekinids and neoschwagerinids from Akiyoshi

Nogami, Y., 1961: Permische Fusuliniden aus dem
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Ota, M., 1968: The Akiyoshi Limestone Group: A.
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- - , Toriyama, R., Sugimura, A. and Haikawa, T.,
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- - , 1970b: Variation and relative growth of
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- - , 1975: Stratigraphy of the Paleozoic and Mesozoic strata in the Tamagawa area, southeasternpart of the K wanto mountains. Sci. Rep., Dept.
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7-10. (in Japanese with English abstract)- - and Kobayashi, F., 1990: Carboniferous toPermian Akiyoshi Limestone Group. In, Organizing Committee Benthos '90 ed., Fossil andrecent benthic foraminifera in some selected
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Upper Chichibu Paleozoic system including the
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Jour. Coli. Sci., Imp. Univ. Tokyo, vol. 45, art. 4,
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- - , 1925b: Paleontological and stratigraphicalstudies on the Permo-Carboniferous limestone of
Nagato, Part II, Paleontology. Ibid., vol. 45,
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- - , 1927: Stratigraphical studies of the fusulina

1067

limestone of Akasaka, province of Mino. Jour.
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- - and Tobler, A., 1929: Permian Fusulinidae
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I, p. 45-49, pI. 5.
Pasini, M., 1965: Fusulinidi, una chiave analitica perla determinazione del generi. 107 p., 18 pIs.
Palaeontgr. Italica, Pisa.
Pitakpaivan, K., 1965: Fusulines of the Rat Burilimestone of Thailand (Fusuline fossils fromThailand, Part I). Mem. Fac. Sci., Kyushu
Unil'., ser. D, vol. 17, no. I, p. 3-69, pIs. 1-6.
Rauser-Chernoussova, O.M. and Fursenko, A. V.,

1959: Osnovy paleontologii, Obschaya chast;
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SSSR, Moskva. (in Russian)Rozovskaya, S.E., 1975: Sostav, sistema i filogeniya
otryada fuzulinida. Akad. Nauk SSSR, Trudy
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Group in the Atetsu Limestone plateau. Jour.
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117-129, pis. 11-14.
Sakagami, S., 1958: Fusulinids from the Upper Permian conglomerates of the northern part of Itsukaichi, Tokyo-to, Japan. Jour. Hokkaido Gakllgei Univ., ser. 2, vol. 9, no. 2, p. 72-97, pis. 14.
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Formations in the Atetsu Limestone Plateau,
southwest Japan. Proc. Japan Acad., vol. 64,
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Sakaguchi, S., Sugano, K., Nakamura, T. andKunugi, K., 1966: A study on the biostratigraphy of the Kaerimizu doline, Akiyoshi limestone Plateau, Yamaguchi Prefecture, Japan (Preliminary). Mem. Osaka Univ., Liberal Arts
and EdIlC., B, no. 14, p.220-229. (in Japanesewith English abstract)
Sheng, J.Z. (Sheng, J.c.), 1958: Some fusulinids from
the Maokou Limestone of Chinghai Province,
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- - , 1963: Permian fusulinids of Kwangsi, Kueichow and Szechuan. Palaeont. Sinica, N.S., ser.
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- - ed., 1966: Fusulinids of China. 177 p., 27 pis.
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97 p., 14 pis. Geol. Publ. House, Beijing. (inChinese)Skinner, J.W. and Wilde, G.L., 1966: Permian
fusulinids from Sicily. Univ. Kansas, Paleont.
Contr., Papers., no. 8, p. 1-16, pis. 1-20.

1068

Katsumi Uena

- - and - - , 1967: Permian foraminifera from
Tunisia. Ibid., no. 30, p. 1-22, pis. 1-32.
Staff, H. von, 1909: Beitrage wr Kenntnis derFusuliniden. Neues Jahrb. Min. Geol.
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Sun, Q. L. and Zhang, L.X., 1985: Early Permian
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Acta. Palaeont. Sinica, vol. 24, no. 5, p. 503-510,
pis. 1-2. (in Chinese with English abstract)
Sun, X.F., Gao, C.X. and Zhang, X.L., 1983:
Fusulinida. In, Xi'an Inst., Geol. Mineral
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Suyari, K., 1962: Geological and paleontologicalstudies in central and eastern Shikoku, Japan,
Part II. Paleontology. Jour. Gakugei, Tokushima Univ., vol. 12, p. 1-64, pis. 1-12.
Thompson, M.L., 1936: The fusulinid genus Verbeekina. Jour. Paleont., vol. 10, no. 3, p. 193-201, pI.
24.
- - , 1946: Permian fusulinids from Afghanistan.
Ibid., vol. 20, no. 2, p. 140-157, pis. 23-26.
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Kansas, Paleont. Contr., Protozoa, art. I, p. 1184, pis. 1-38.
- - and Foster, c.L., 1937: Middle Permian
fusulinids from Szechuan, China. Jour.
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Tien, N.D., 1970: Quelques fusulinides de NGi Com,sud Viet-Nam (Zone a Neoschwagerina). Archives Geol. Viet-Nam, no. 13, fasc. I, p. I-70,
pis. 1-15.
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Permian of Southeast Asia, CCOP Tech. Bull.,vol. 18, p.116-137, pis. I-10.
- - , 1986b: Foraminifera and algae from the Permian of Guguk Bulat and Silungkang, Sumatra.
Ibid., p. 138-147, pis. 11-15.
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Toriyama, R., 1947: On some fusulinids from
Tosayama, Koti-ken, shikoku, with a note on thestratigraphical range of Neosch wagerina.
Japan. Jour. Geol. Geogr., vol. 20, nos. 2-4, p.
64-82, pis. 26-27.
- - , 1954: Geology of Akiyoshi, Part I. Study of
the Akiyoshi Limestone Group. Mem. Fac.
Sci., Kyushu Univ., ser. D, vol. 4, no. I, p. 39-97.

1958: Geology of Akiyoshi, Part III.
Fusulinids of Akiyoshi. Ibid., vol. 7, p. 1-264,
pis. 1-48.
- - , 1975: Permian fusulines from the Rat Buri
Limestone in the Khao Phlong Phrab area, Sara
Buri, central Thailand (Fusuline fossils fromThailand, Part IX). Ibid., vol. 23, no. I, p. 1116, pis. 1-21.
- - and Kanmera, K., 1977: The Permian fusulines
from the limestone conglomerate formation in the
Khao Phlong Phrab area, Sara BUfi, central
Thailand (Fusuline fossils from Thailand, PartX). Geol. Palaeont. Southeast Asia, vol. 18, p.
1-27, pis. 1-3.
- - and - - , 1979: Permian fusulines from the
Ratburi Limstone in the Khao Khao area, Sara
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4-14.
- - and Pitakpaivan, K., 1973: Middle Permian
fusulines from Wait Kirinakratanaram, central
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Ueno, K., 1989: Carboniferous and Lower Permian
foraminiferal biostratigraphy in the Akiyoshi
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Akiyoshi-dai Mus. Nat. Hisl., no. 24, p. 1-39, pis.
1-8. (in Japanese with English abstract)- - , 1991a: Pamirina (Permian Fusulinacea) fromthe Akiyoshi Limestone Group, southwest Japan.
Trans. Proc. Palaeont. Soc. Japan, N.S., no. 161,
p.739-750.
- - , 1991 b: Early evolu'tion of the families Verbeekinidae and Neoschwagerinidae (Permian
Fusulinacea) in the Akiyoshi Limestone Group,southwest Japan. Ibid., no. 164, p.973-1002.
Wang, J.H. and Tang, Y., 1986: Fusulinids from
Chihsia Formation of Lengwu district in Tonglu
County, Zhejiang. Acta Micropalaeont. Sinica,
vol. 3, no. I, p. 3-11, pis. 1-3. (in Chinese withEnglish abstract)
Wang, Y.H., Wang, L.L., Wang, J.H., Zhu, Z.G., Lin,
G.W., Zhang, L.X. and Quian, Q., 1982:
Fusulinida. In, Nanjing Inst. Geol. Mineral
Resour. chiefly ed., Paleontological A tlas of
East China, no.2 (Late Palaeozoic), p.5-108,
pis. 1-30. Geo!. Pub!. House, Beijing. (inChinese)Wang, Y.J., Sheng, J.Z. and Zhang, L.X., 1981:
Fusulinids from Xizang of China. In, NanjingInst. Geol. Palaeont. ed., Palaeontology of
Xizang, Book 3, p. I-80, pis. 1-21. Sci. Pub!.
House, Beijing. (in Chinese with English
abstract)- - and Zhou, J.P., 1986: New material of934.

Verbeekinids and neoschwagerinids from Akiyoshi

Bull.. NanjingInsl. Ceol. Palaeont.• Acad. Sinica. no. 10, p.
141-156, pis. 1-4. (in Chinese with English
abstract and description of new species)
Xie, S.G., 1982: Protozoa. In, Geol. Bureau, Hunan
ed.. The Palaeontological A tlas of Hunan. Ceol.
Mem .. ser. 2, no. I, p.2-73, pis. 1-35. Geol.
Publ. House, Beijing. (in Chinese)
fusulinids from Xainza, Xizang.

Yanagida, J., Ota, M., Sugimura, A. and Haikawa, T.,1971 : On the geology of the northeastern part of
the Akiyoshi limestone plateau. Sci. Repts..
Dept. Ceol.. Kyushu Univ., vol. II, no. I, p. 105114, pis. 7-10. (in Japanese with English

abstract)
Yang, Z.O., 1985: Restudy of fusulinids from the1069"Maokou Limestone" (Permian) at Oatieguan,
Langdai, Guizhou. Acta Micropalaeonl. Sinica.
vol. 2, no. 4, p. 307-335, pis. 1-3. (in Chinesewith English abstract and description of new
genus)
Yoshimura, N., 1961 : Geological studies of the Paleozoic groups in the Oga Plateau, central Chugoku, Japan. Ceol. Rep.. Hiroshima Unil'., no.
10, p. 1-36, pis. 1-4. (in Japanese with English

abstract)
Zhang, L.X. and Wang, Y.l., 1974: Permian
Fusulinida. In, Nanjing Inst., Geol. Palaeonil.

ed.. Stratigraphical and Paleontological Atlasof West South China. p.289-296, pis. 150-153.
Sci. Pub I. House, Beijing. (in Chinese)

Akiyoshi f.:k6', Akiyoshi-dai f.:k6'iJ. Atetsu ~iiJ1Y. Kaerimizu jffi I) 7.K. Omi WfliJ. Taishaku

7iffR. Yamaguchi ill p.
f.:k 6' E I!k.;fi ~ ff. Parafusulina kaerimizensis '* tJ Ili'f: 0) Verbeekinidae:f4 to J: If Neoschwagerinidae :f4f,jj~£E.~: f.:k6'iJ;lt*$O)jffi I) 7Jc~~I=-5t;fjj-t Q Parafusulina kaerimizensis '* J: I) 1::. {lL 0) fKi5 E I!k.;fi ~ ff t.J' G II flO) Verbeekinidae :f4 to J: U' Neoschwagerinidae fJj.iiiO~£E.~~~5J1j L t-.:o ~ G v=- c..h Gf,jj~£E.~O) ~ J1':5t;fjj v=-~--j\;, -C. Parafusulina
kaerimizensis '* tJlli'f: 0) f.:k6'E I!k.E ~ff ~ T {lL t.J' G Parafusulina kaerimizensis m. Afghanella ozawai '*. Neoschwagerina craticulifera robusta
Verbeekina verbeeki-Afghanella
schencki '*. Neoschwagerina fusiformis '*. Verbeekina verbeeki m. Colania douvillei m0) 71tE'*v=-5tm L t-.:o *~tfli-efi I ~BEfl (Neoschwagerina craticulifera robsta, subsp,nov.) ~%tsc.O) ') 'f:,0) 10 fl~~G$t-t Q c. c. <b v=-. Parafusulina kaerimizensis ,*tJIli'f:O)f,jj~£E.~~J1':v=-":)\;'-C <b ~!jtv=-71!:""Qo
1::.!!1TII9J~m.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1070-1089, 7 Figs., April 30, 1992

935.

A NEW CRYPTIC SPECIES OF PYCNODONTE FROMRYUKYU ISLANDS: A LIVING FOSSIL OYSTER *ITARU HAYAMI
Geological Institute, University of Tokyo,
Hongo 7-3-1, Bunkyo-ku, Tokyo, 113
and

TOMOKI KASE
National Science Museum, Hyakunin-cho
3-23-1, Shinjuku-ku, Tokyo, 169

Abstract. A new enigmatic oyster was found alive from several poorly lighted submarine caves of Miyako and Okinawa Islands, southern Japan. It belongs to the genus
Pycnodonte, which occurs commonly in the Cretaceous-Miocene of the Tethyan realm but
had been believed extinct. Its shell shape is variable, but the right valve is paper thin andhas an extremely wide depositional surface of flexible outer prismatic layer. Nevertheless,every diagnostic character of Pycnodonte (s.s.) is well recognized (e.g. long dorsal margin,
dorsally located and subcircular adductor muscle insertion, wide commissural shelf, vermicular chomata, vesicular shell structure and remarkable radial gashes on the right valve).
It is suggested that this oyster is a significant example of a living fossil, and that Pycnodonte
has survived for many millions of years by transforming itself into a cryptic organism.
Key words.

cryptic, Pycnodonte, submarine cave, living fossil.

High levels of endemism in cryptic organisms have attracted the attention of many
biologists. Riedl (1966) made a comprehensive review on the biota of coastal caves andillustrated the spectral distribution of variousorganisms (including some bivalves) in somewave-excavated intertidal grottoes of theMediterranean region. Since Hartman and
Goreau (1970) discovered sclerosponges
resembling stromatoporoids (problematic
Paleozoic and Mesozoic fossil organisms) insome sublittoral crevices of Caribbean coral
reefs, several species of articulate brachiopods, bryozoans, polychaetes anddecapod crustaceans have been recognizedalso as unique constituents of cryptic communities, and their origin and evolutionary
significance have been discussed by many

Introduction

The technical development and popularization of scuba diving in recent years enabled us
to examine the diversity and ecology of various organisms indigenous to sublittoral
sheltered environments. Submarine cavefaunas, in particular, have emerged as aninteresting subject of study because they seem
to offer substantial evidence to test various
theories of systematics, evolutionary ecology
and marine biogeography. Cryptic faunas
also may contain paleontologically interesting species because such sheltered places serve
as suitable refuges to archaic and anachronistic organisms.
'Received January 10, 1992; accepted February 3,
1992
1070

935.

A living fossil oyster

investigators (Jackson, Goreau and Hartman,
1971; Jackson and Winston, 1982; Iliffe,
Hart and Manning, 1983; Kobluk, 1988;
etc.). However, the unique features of molluscs in such sheltered environments havebeen little studied.
On the fore-reef slopes of Shimoji-shima
and Ie-jima of the Ryukyu Islands, southern
Japan, there are numerous limestone caves
which seem to have been formed by underground water during some low sea-levelstages of the Pleistocene and drowned during
the post-glacial sea-level rise. Since a few
years ago, we have been studying the cryptic
faunas (especially molluscs) in these caveswith the vigorous assistance of several skilled
divers. As reported preliminarily (Hayami
and Kase, 1991 ; Kase and Hayami, in press.),
several characteristic moll uscs incl uding Neritopsis radula (a living fossil snail) have beenfound alive in some of these caves, and thecalcareous muddy sediments therefrom bear
numerous dead shells of bivalves and gastropods which are different from the shallowwater faunas of ordinary exposed environments. The diversity, ecology, origin andevolutionary significance of these cryptic
faunas may be an interesting subject to study.
In spite of the apparent great species diversity, most of these molluscs are very small insize (commonly less than 5 mm in maximum
length). Some sclerosponges, corals, soft
sponges, articulate brachiopods, bryozoans,
ostracods and decapod crustaceans were also
found alive in some of these caves, but thebiomass per unit space is generally muchsmaller than that of the exposed environment
around the caves.
The new oyster described here is an exceptionally large-sized species among the cavebivalves. It was first discovered by M.
Taniguchi, a skilled diver of Miyako Island,at the poorly lighted walls and ceilings ofseveral caves (about 20 meters below sea
level) along the western coast of Shimojishima of Miyako Islands (Figure 1). The
same species has been found also from simi-

1071

larly sheltered environments of Okinawa (Iejima and Seragaki beach). In spite of its
enigmatic shell morphology, many essential
characters indicate that this oyster belongs to
the genus Pycnodonte, which flourished in
Cretaceous and Paleogene periods in the
Tethyan realm and had been believed to be an
extinct taxon. Its cryptic habitat is alsomeaningful, because such a sheltered place
may offer a refuge for archaic organisms. In
this article we describe its morphology andlife habit and discuss its evolutionary
significance as a "living fossil".
On the genus Pycnodonte
Before going into the description, previous
systematic studies of pycnodonteine oysters
are briefly reviewed because the newly discovered cryptic species seems to belong to
Pycnodonte.
In spite of great intraspecific morphologic
variability and frequent convergence, naturalclassification of fossil and extant oysters has
been attempted by many investigators.
Among others, Stenzel's (1959, 1971) scheme
may be the most elaborate and influential.
He classified oysters into two families,
Gryphaeidae and Ostreidae, and further subdivided the former into three subfamilies,
Gryphaeinae, Pycnodonteinae and Exogyrinae. Stenzel's scheme has been substantiallyfollowed by many subsequent authors,
though Torigoe (1981) raised the Pycnodonteinae as a distinct family, Harry (1985)
subdivided the Pycnodonteinae into threetribes (Pycnodontini, Neopycnodontini andHyotissini), and Carter (\ 991) regarded the
Gryphaeidae and the Ostreidae as constituting distinct superfamilies.
Important criteria for these family- or
subfamily-level classifications are 1) shellcharacters (particularly, presence or absenceof chalky deposit and vesicular shell structure, commissural shelf, chomata, and shapeof muscle scar) and 2) anatomical charactersof soft parts (particularly, the status of theItaru Hayami and Tomoki Kase

1072

N

t

"CDI
\
e
*

-..

' ...... -.Daidokutsu

B~ \. ... _..,... .....'oAma~i .••

f~Miii"na••
Ie "- ,.,c, .~ Okinawa )(

....

•

:z
:z:

ShimDji• :\ •• MiyakD

:z

....
....

c
Q

:

: ~ '",I'I/,i.,'6_k~m'. .L:. -._- -,c: . '=" :~_I. .J:. .:1.

__i_X__LO________ _ _ _ _---'c.~{:) IkemaoMiyako-o

6 km

Figure 1. Map showing the localities (indicated by asterisks) of Pycnodonte (Pycnodonte) taniguchii n.
sp. in Ryukyu Islands of Japan. A: Index map, B: Central part of Okinawa Island, C: Miyako Islands.

promyal passage, heart-kidney-rectum complex, digestive tract, labial palp fusion andanal appendage) and 3) reproductive strategy (incubatory or non-incubatory). In

accordance with Stenzel's system (with somemodifications), Torigoe (1981) described theshell and soft parts of all the extant oysters inJapanese waters, and Harry (1985) gave a935.

A living fossil oyster

synopsis of all the supraspecific taxa of extant
oysters in the world.
Stenzel (1971) distinguished the Pycnodonteinae from the Gryphaeinae by the presence of a well defined commissural shelf,vermiculate chomata and vesicular shell structure, but treated the former as derived fromthe latter, regarding Gryphaea (Bilobissa)
from the Middle-Upper Jurassic as transitional between the two subfamilies. Subsequent studies of shell microstructure, however, do not necessarily support Stenzel's
scheme. According to Siewert (1972) andHudson and Palmer (1976), Pycnodonte andHyotissa considerably differ from the
Gryphaeinae but resemble the Ostreidae inthe structural chambering and in lacking welldefined crossed foliated structure. Siewert(1972) supposed that Pycnodonte together
with many ostreid genera was derived from
Lopha which constituted a stock independent
from Gryphaea throughout the Jurassic.
Carter (1991) critically reviewed all theavailable data about the shell microstructureof fossil and extant oysters, and admitted thestructural resemblance between the Pycnodonteinae and the Ostreidae, although he did
not change Stenzel's (1971) system in which
the Pycnodonteinae is regarded as a subfamily of the Gryphaeidae. Contrary to Stenzel's description, "No prismatic shell layer
except in Neopycnodonte': an outer layer ofcalcitic simple prisms is commonly observed
in RV of many fossil and extant
pycnodonteine species. Vesicular shell structure, which was regarded by Stenzel (1971) as
diagnostic of the Pycnodonteinae, exists also
in a lophine oyster, Dendostrea folium.
Therefore, the origin and the familial position of the Pycnodonteinae is still controversial. If much weight is given to the shellstructure and structural chambering, thePycnodonteinae may be considered to be
more closely related to the Ostreidae (especially the Lophinae) than to the Gryphaeinae.
Nevertheless, as to the content and limit of
the Pycnodonteinae, most recent investigators

1073

seem to support Stenzel's (1971) scheme.
Discrimination of the Pycnodonteinae
from the Ostreidae is justified also by comparative anatomy of extant species. As illustrated by Torigoe (1981), the Pycnodontinae
shares a Z-shaped digestive tract with theLophinae, but the rectum penetrates (orcomes into contact with) the pericardium of
the heart in Neopycnodonte and Hyotissa,
while it passes the dorsum of the heart in all
the genera of the Ostreidae (including theLophinae). Adductor muscle insertion is
subcircular in the Pycnodonteinae (like theGryphaeinae) but reniform in the Ostreidae.
The status of the promyal passage and labial
palp fusion may be also important to discriminate the Pycnodonteinae from the Ostreidae.
Habe (1977) regarded Pretostrea as a pycnodonteine genus and referred several Japanese
species to it, but Torigoe (1981) clarified on
the basis of anatomy that some of thembelong to Hyotissa and other species to
Dendostrea (of the Lophinae). These anatomical characters are useful to see throughshell convergence.
Pycnodonte Fischer de Waldheim, 1835, as
defined by Stenzel (1971) and many recent
workers, had been regarded as an extinct
genus, though its descendant(?) genus
Neopycnodonte Stenzel, 1971, is represented
by a single extant circumglobal species, N.
cochlear (Poli). Hyotissa Stenzel, 1971,
which comprises several tropical-subtropical
species and corresponds to the tribe Hyotissini Harry, 1985, is another extant pycnodonteine genus, but the Lopha-like plicate
valves, subequivalve shell, internal surface
with a moire luster and huge adductor muscle
insertion are dissimilar from those of
Pycnodonte and Neopycnodonte.
A large number of fossil species of
Pycnodonte (including Gigantostrea,
Phygraea and Costeina as subgenera) havebeen known mainly in the Tethyan realm
from late Early Cretaceous to early Miocene.
Probably owing to biased fossil preservation,
only left valves were described in many fossil

1074

Itarll Hayami and Tomoki Kase

species. Though the intraspecific variationand detailed shell characters of the typespecies are poorly known, the gen us
Pycnodonte, as defined by Stenzel (1971 : p.
N 1106), seems to be characterized by strongly
convex LV, irregular but generally subcircular to semicircular outline, wide commissural shelf, vermicular chomata, gently undulated LV rarely with well defined radial
costae and flat to concave RV. Radial
gashes of R V is a very conspicuous feature of
Pycnodonte, though they sometimes occur
also in other genera of the Pycnodonteinae (e.
g. Harry, 1985: p. 136, fig. 16) and somespecies of the Gryphaeinae (e.g. Duff, 1978:
pI. 8, fig 7b).
Subgeneric division of Pycnodonte is controversial and somewhat confusing.
Pycnodonte (s.s.) [type species: Pycnodonte
radiala from the Upper Cretaceous of the
Crimea] is generally defined by the improminent umbo, long dorsal margin, developed auricular parts and absence of radialribs on LV. Gigantostrea Sacco, 1897 [typespecies: Ostrea gigantica Solander in Brander, 1766, from the Eocene of England], wasregarded as synonymous with Pycnodonte(s.s.) by Stenzel (1971 : p. NI107), but Palmer
and Brann (1965: p. 149) and Wilson (1987 :p. 13) applied this subgeneric name to several
species from the Eocene-Miocene of the Atlantic Coastal Plain of USA, and Fleming(1966: p. 23) also to two species from the
Paleogene of New Zealand. Gigantostrea
seems to differ from Pycnodonte (s.s.) in theplanoconvex to biconvex outline, less inequivalve shell and undeveloped auricles.
Phygraea Vyalov, 1936 [type species:
Phygraea frauscheri Vyalov, 1936 (=Gryphaea pseudovesicularis GUmbel, 1861),
from the Paleocene of eastern Europe] shows
planoconvex to concavoconvex outline andconspicuous radial gashes of RV like
Pycnodonte (s.s.), but its subgeneric distinction can be based on the Gryphaea-like outline (though the resemblance is due to convergence), much thicker LV, prominent umbo of
L V, short dorsal margin, and undeveloped
auricles. This subgenus possibly includes
Pycnodonte vesicularis (Lamarck, 1806) fromthe Upper Cretaceous of western Europe, west
and north Africa, south India and New
Caledonia. Woods (1913) and Freneix(1960,1972) described the wide morphologic
variation of P. vesicularis, and many European authors regarded P. radiata, the typespecies of Pycnodonte, as synonymous with P.
vesicularis. This view was rejected and P.
vesicularis was referred to Phygraea by Stenzel (1971) and some subsequent authors,chiefly because its subcircular shell form
looks different from the figured original specimen of P. radiata. The shape difference
between Pycnodonte (s.s.) and Phygraea, as
discussed later, may be related to di fferent
modes of life.
In addition, Costeina Vyalov, 1965 [typespecies: Pycnodonte (Costeina) costei Coquand, 1869, from the Upper Cretaceous ofnorth Africa] may be characterized by thepresence of dichotomous radial ribs on thesurface of LV, and Crenostrea Marwick, 1931
[type species: Ostrea (Crenostrea) wuellerstorfi Zittel, 1864, from the Oligocene of New
Zealand] seems to be subgenerically distinguished by the pointed umbo of LV and verystrong chomata, though R V of their typespecies are insufficiently known. Squires
and Demetrion (1990) proposed Pegma [typespecies: Pycnodonte (Pegma) bajaensis
Sq uires and Demetrion, 1990, from the
Eocene of Baja California] as a subgenus of
Pycnodonte, but we think that it is, if notsynonymous with, closely related to Hyotissa.

~
Figure 2. Pycnodonte (Pycnodonte) tanigllchii n. sp., Holotype (UMUT RM 18908), brownish individual. la: Exterior of LV with many dead shells of juvenile individuals, lb: Interior of LV, thedepositional surface of middle layer showing pepper-and-saJt appearance, Ie: Exterior of RV, showing
remarkable radial gashes. Id: Interior of RV, showing wide depositional surface of prismatic outer layer.
All figures xO.7.

935.

A living fossil oyster

1075

Itaru Hayami and Tomoki Kase

1076

No.

Reg. no.

I
2
3
4
5
6
7
8

RM18908
RM18909
RMI8910
RM18911
RM18912
RMI8913

Table 1. Observed specimens of Pycnodonte (Pycnodonte) taniguchi/, n. sp.
LengthHeight Thickness Phenotype
Locality
Type
I03mm
87 mm
I09mm
96mm
97mm
97mm
123 mm
81 mm

138mm
124mm
I05mm
84mm
76mm
101 mm
125mm
84mm

71 mm
71 mm
68 mm
45mm
39mm
53mm
66mm
28mm

brownish
whity
whity
brownishbrownishbrownishbrownishbrownish

W-ArchBlack Hole
Devil's Palace
W-ArchSeragaki Beach
W-ArchW-ArchW-ArchHolotype
Paratype
Paratype
Paratype
Paratype
Paratype

Remarks
dried
dried
in alcoholin alcoholin alcohol
sectioned
anatomized
anatomized

Registered specimens are in the University Museum, University of Tokyo (UMUT).
Speciment Nos. 7 and 8 were anatomized by Dr. Torigoe of the Hiroshima University.

The geographic distribution of Pycnodonte
was said to be worldwide (Stenzel, 1971 : p.
N 1107), but this genus has been known
almost exclusively from low-middle latitudinal regions. Cretaceous species are common
in western Europe, Crimea, north and west
Africa, south India and Gulf Coast regions.
Paleogene species show almost the same geographic distribution as Cretaceous ones. In
Japan large species of Pycnodonte seems to berare, but Tashiro (1978) described a small
species from the Santonian of west Kyushu,
and Hanai and Oji (1981) noticed numerous
individuals (mostly LV) of an undescribed
species adhering to an Aptian fossil beachrock in north Honshu. As revised by
Oyama, Mizuno and Sakamoto (1960: p.
129), Ostrea crassis Nagao, 1928, from the
Oligocene of west Kyushu seems to belong to
Pycnodonte, although Nagao (1928) wrongly
described the right and left valves in reverse.
All the extant representatives of the
Pycnodonteinae (Neopycnodonte and Hyotissa) are also tropical to warm-temperate
inhabitants (Harry, 1985,1986; Nicol, 1991).

Systematic description? Family Gryphaeidae Vyalov, 1936
Subfamily Pycnodonteinae Stenzel, 1959
Genus Pycnodonte Fischer
de Waldheim, 1835
Subgenus Pycnodonte s.s.

Pycnodonte (Pycnodonte) taniguchii
Hayami and Kase, n. sp.
Figures 2-7

Material. - Several living specimens fromthe submarine caves on the western coast of
Shimoji-shima, Miyako Islands, Ryukyu(Table I). The holotype (UMUT RM 18908)
from one of the caves, a diving point called"W-Arch" (Lat. 24°51'43"N, Long. 12Y09'41"
E, ca. 20 m below sea level).
Diagnosis. - A large and concavo-convex
extant species of Pycnodonte (s.s.), the adultshell of which is characterized by an unusually wide commissural shelf, an attachment
area extending over the long geniculated
dorsal margin of LV, distinct vermicularchomata, dorsally situated subcircular adductor insertion, variably developed vesicular
structure, moderately thick LV, and more orless hollowed RV with a paper-thin andflexible distal area and remarkable radial
gashes on the external surface.
General shell features. - Shell large for the
genus, often exceeding 100 mm in length andheight, very inequivalve, more or less prosocline, nonplicated, presumably dimorphic incoloration (commonly purplish brown but
sometimes ivory white). Outline variable,
usually flat and subcircular in young stage
but becoming linguiform or fan-shaped in
adult. LV strongly convex, solid, relatively
thin except for middle-ventral area in adultstage, characterized by improminent umbo,
broadly undulated and puckery surface and935.

A living fossil oyster

10

10 77

1b

Pycnodonte (Py cnodonte) wniguchii n. sp. , Holotype (UMUT RM 18908). la: Umbonal
area of LV , showing the liga ment area a nd muscle inse rtion s, Ib: Umbonal area of RV , showing the ligament
Figure 3.

area a nd muscle insertions.

All figures X 1.5.

generally well developed auricular parts,
sometimes having a posterior radial sulcus
but lacking any well-defined radial sculptureand hyote spines. Attachment area , which
occupies the whole external surface of LV inearly stage (up to 20- 30 mm in height) ,extends along the entire dorsal margin in
adult, geniculated at the umbonal area in
dorsal view. R V broadly and deeply concave except flattened or feebly convex umbonal area, sticking to the inner surface of LVwith flexible distal area, extremely thin andlight (in the holotype, L V is 221 gram and RV
is 26 gram in weight) , characteri zed byremarkable radial gashes on the external surface. Ligament area relatively narrow,subtriangular with an apical angle of about
60' in LV, though its apical part was commonly corroded in RV ; resilifer small , subcentral, slightly opisthocline. Subumbonal
cavity of LV moderately deep. Inner surface
of both va lves shiny but does not show amoire lu ster. Adductor muscle insertion
comparatively small (with a diameter about

1/ 10 of shell height) , subcircular, without
elevated rim , slightly longer than high in LV
but nearly as long as high in RV , with its
center placed at one-fifth to one-fourth ofshell height from the dorsal margin in adult(unusually dorsally situated) . Gill protractor muscle scar (Quenstedt muscle insertion
by Sten ze l, 1971) distinct in both valves,situated a nteriorly in LV but subcentrally
nearly below the resilifer in RV. Chomata
relatively weak but distinct, moderate inlength , vermicular, situated on the outerslopes of a pair of small ridges. Commissural shelf unusually broad , occupying almost
the distal half of shell; its proximal margin
markedly angulated on the internal surface of
RV.
Shell microstructure. - The outer layer of
L V often ragged but consists of accumulated
regularly foliated calcite which dips slightly
relative to depositional surface. The middle
laye r of LV composed of thick accumulationof composite crossed foliated calcite. In
purplish-brown individuals thi s layer catches1078

flaru Hayami and Tomoki Kase

935.

A living fossil oyster1079____ umbo (corroded)~ resilifer"-j --~~
If
It ~ ,ermicular chomala~~tQuenstedt muscle insertion

Q"'---;-~ "..

0.,.,.--\ \ -'I'- - /
U)'-)
./ \
. J
/

/ _\

b

adductor muscle insertion
proximal margin ofcommissural shelf'~ distal margin of/ / j/

radial gash

/

~

~

-,\,~----',-- de~~~:~~i~ :~~:~cl~y~r

\ ...

\\
- - - - ventral margin (incomplete)Figure 5.

Sketch of the internal surface of RV (based on the holotype).

many yellowish lenses which bear numerous
ellipsoidal vacuoles of about 40 pm in maximum diameter (Figure 6-6). The brownish
and yellowish parts irregularly intermingled,
so that the depositional surface shows apepper-and-sait appearance (Figure 2-1 b).
Development of vesicular structure quite variable in LV; in some individuals there arelarge cavities with very coarse vesicular structure between the outer and middle layers, but
in many other individuals this structure is notobserved at all. The outer layer of RV comparatively thick in the middle-ventral area
(attaining almost I mm in total), composed ofseveral sublayers of simple prismatic calcite
which are interrupted by thin conchiolin seatsor cavities (Figure 6-2). Calcite prisms variable in size, but commonly 20-50 pm indiameter and reclined about 50· to the sur-face. Its depositional surface extremely
wide, occupying almost the entire distal halfof shell. Owing to this structure, the distal
part of RV without backing of middle layer
very flexible when alive. Radial gashes
occur only in this outer layer. A number offlat cavities exist in and below the prismatic
outer layer, in which vesicular structure iscommonly developed (Figure 6-3, 4). Middle layer of R V cross foliated, solid and rarely
vacuolated, easily separable from outer layer.
The inner layer of both valves composed ofcomplex crossed foliated calcite but generally
very thin.
Soft part. - The soft part relatively small,as recognized from the unusually widecommissural shelf. Promyal passage extends
between RV and visceral mass, reaching theumbonal part. Visceral mass relatively+- Figure 4. Pycnodonte (Pycnodonte) taniguchii n. sp. 1: Holotype (UMUT RM 18908), dorsal viewof articulated valves, 2a-c: Paratype (UMUT RMI891O), left, right and dorsal views of a whity articulated
individual. 3a-c: Paratype (UMUT RMI8911), left, right and dorsal views of a brownish articulated
individual. All figures X 0.7.

1080

fl aru Hay ami and Tomoki Kase

935.

A living fossil oyster

small, orange in color. Labial palp broadly
fused, resembling that of Neopycnodonte cochlear. Digestive tract relatively thick, Zshaped, never looped; rectum does not penetrate the pericardium of the heart but comesin contact with its posterodorsal part; anal
papilla very long. In summary, every anatomical character resembles that of Neopycnodonte and Hyotissa and differs significantly
from that of the Ostreidae. (The descriptionof anatomical characters is mainly based on
Torigoe's personal communication.)
Comparisons. - The present new oystershows every essential character of the
Pycnodonteinae called for by Stenzel (1971),
Torigoe (1980) , Harry (1985) and Carter(1991). In the shell morphology as well as
the cryptic habitat, however, the present
species is so unique that its distinction fromother extant oysters can be based on various
characters. The extreme inequivalveness,broad commissural shelf, dorsally situatedadductor muscle, and thin and deeply concave RV with broad and flexible distal area
(owing to the shell consisting of simple prismatic outer layer) in the adult stage are themost striking diagnostic characters of thisspecies.
Among extant pycnodonteine oysters,
Neopycnodonte cochlear (Poli, 1795) may bemost closely related to the present species
because the two species share very dorsally
situated adductor muscle, vermicularchomata, radial gashes and limitedly distributed vesicular structure in RV. According to Torigoe (pers. comm.), who anatomized an individual at our request, many anatomical characters are not much different

1081

from those of N. cochlear, although the rectum does not pierce the pericardium in thepresent species. N. cochlear is regarded as a
circumglobal species, having been recorded
from various regions of the Indo- West
Pacific, eastern and western Atlantic, andMediterranean realms. It extends to greater
depths (27 to 2,100 m) than any other extant
oyster (Stenzel, 1971; Harry, 1985). In
Japan, though it often was treated under thename of Ostrea musashiana Yokoyama,
1920, the species occurs commonly on thelower sublittoral to upper bathyal gravelly
bottom (Kuroda, 1931; Habe, 1977; etc.).
Although the soft parts are relatively small,the present species grows much larger than N.
cochlear in shell size. The shape of the adultshell is also quite different because the attachment area of the present species extends
entirely over the long dorsal margin of LV.
Moreover, unlike N. cochlear, the simple
prismatic outer layer of RV is well developed,
and its depositional surface is much wider.
The shell of N. cochlear is fragile, while the
L V and the middle-inner layer of R V are
generally solid in the present species.
The present species resembles the typespecies of Pycnodonte (s.s.) more closely thanthat of Neopycnodonte in various shell characters, e.g. concavo-convex valves, long dorsal margin and linguiform to fan-shaped
outline in the adult stage and well developed
radial gashes on the external surface of RV.
Some specimens of the present species are
surprisingly similar in shell shape to thefigured original specimen of its type species,
P. radiata Fischer de Waldheim, 1835 (reproduced by Stenzel, 1971 : fig. J80), from the~
Figure 6. SEM photomicrographs showing shell microstructure of Pycnodonte (Pycnodonte)
laniguchii, n. sp. Unit of scale in microns. 1: Depositional surface of simple prismatic outer layer near theventral margin of RV (UMUT RM 18913); the venter is toward the lower, X 250. 2: Oblique internal viewof fractured edge of multilayered simple prismatic outer layer near the ventral margin of RV (the same
individual); the venter is toward the right upper, X 500. 3: Fractured external surface of middle part of
RV (UMUT RMI8909); vesiculate cavity below simple prismatic outer layer, X 100. 4: Vesiculate structure in a cavity between outer and middle layers of RV (the same individual), X 60. 5: Depositional surfaceof crossed foliated middle layer of LV (UMUT RM 18913), X 1,000. 6: Slightly etched surface of subvertical section of LV (the same individual), vacuolated lenticular part embedded in crossed foliated middle
layer; the venter is toward the right and the external surface toward the lower, X 100.

1082

ftaru Hayami and Tomoki Kase

Upper Cretaceous of the Crimea. In a number of fossil species of Pycnodonte RV looks
much smaller than LV, as described andfigured by many authors. Judging from theconcordant ventral margins of two valves inall the living specimens of the present species,the "size discordance" in those fossil species
may be only superficial, because the marginal
part of R V was physically weak and may havebeen selectively lost before fossilization.
Even in living specimens of the present
species, this portion, if dried and not strengthened, is apt to be broken into pieces, just likethe marginal apron of RV in living
propeamussiids (see Hayami, 1988). Flexible distal area of RV, therefore, may be aubiquitous feature in the genus Pycnodonte.
The apparently wide range of variation ofshell form of the present species is partly
attributable to ontogenetic change. Some
immature specimens of the present species(e.g. UMUT RM18911) show shorter andarcuate dorsal margin and subcircular outline, resembling some species of the subgenus
Phygraea, especially Pycnodonte (Phygraea)
mutabilis (Morton, 1828) from theCampanian-Maastrichtian of the Gulf andAtlantic Coastal Plain, which was describedby Gardner (1916) and Wade (1926) as
Gryphaea vesicularis and by Stephenson(1941) as Gryphaea mutabilis. In thisrespect, some specimens of Pycnodonte(Ph ygraea) vesicularis (Lamarck, 1806), a
widely distributed Late Cretaceous species in
Europe, north and west Africa, India and
New Caledonia (Stoliczka, 1870-1871;
Woods, 1913; Dartevelle and Freneix, 1957 ;Freneix, 1960, 1972), also exhibit a similar
outline. Those Cretaceous species of
Phygraea, however, have much thicker shells,
and, as interpreted by Jablonski and Bottjer(1983), probably were cup-shaped recliners
on soft substrata.
On the other hand, fossil species of
Pycnodonte (s.s.) were mostly lifelong sessile
animals, because the shell is not very thickand because the attachment area seems toextend over the long dorsal margin as seen in
the present species. All fossil Pycnodonte(s.s. and Phygraea) share with the present
species conspicuous radial gashes on R V,
which indicate the presence of a well developed simple prismatic outer layer.
Vesicular shell structure is widespread in
the fossi I and extant species of the
Pycnodonteinae and has been regarded by
Stenzel (1971) as diagnostic of this subfamily.
This structure is actually observed in thepresent species (Figure 6-3,4). In RV it iscommonly seen in small cavities beneath theouter layer, although in LV its development isquite variable among individuals. As Carter(1991: p.355, fig.2E) ascertained the presence of locally developed vesicular structurein the middle layer of a lophine oyster,Dendostrea folium, taxonomic evaluation ofthis structure should be reconsidered. Chel'tsova (1969) mentioned the presence of
vacuolated and "pinnate" structure 1TI
Pycnodonte radiata. The type species ispossibly similar to the present species in thisfeature, though a more detailed comparisonof shell microstructure has yet to be workedout.
Distribution.-In addition to the type locality, the present species was found alive insubmarine caves on fore-reef slope ("Black
Hole" and "Devil's Palace" as called bydivers) along the western coast of Shimojishima of Miyako Islands, and similarly
sheltered places at "Daidokutsu" of Ie-jimaand at Seragaki near Manza Beach of Okinawa Island (all 20-30 m in depth) (Figure 1).

Observations and discussions
Cryptic habitat. - The habitat of the present oyster is hardly accessible to us, but its life
habit can be recognized through divers' talkand underwater videotapes. Various caveorganisms seem to show spectral distribution
from the entrance to the innermost part inaccordance with the change of physical andbiological factors. Individuals of this oyster

935.
umbo

A living fossil oyster1083

cavity with vesicular structuresimple prismatic outer layer
epibiont (scleractinian)I--

1/..-::

c:

~

E

.r.
o

ftI
f tI

abandoned venter

" ,

:'<:

Cave waif, ' ,geniculation

adductor muscle

epibiont (sclerosponge)

vacuolated lens
10mm

Figure 7. Subvertical section of Pycnodonle (Pycnodonle) lanigllchii, n. sp. in living position (based ona sectioned articulated individual, UMUT RM 18913).

always grow on gloomy walls or ceilings ofsubmarine caves and other sheltered places
open to fore-reef slopes under normal salinity
and temperature. Such a large-sized suspension feeder as this oyster probably cannot live
unless a considerable amount of phytoplankton is supplied from the exposed environment. It does not live in the totally dark
innermost part of such caves, where Neritopsis
radula and several characteristic minute molluscs are still found alive.
The present oyster, on the other hand, does
not dwell in the ordinary exposed environment. This is probably the main reason why
such a large-sized conspicuous bivalve had
not been discovered by scientists and collectors until quite recently. Moreover, owing to
the wide attachment area the LV seems toadhere to the cave wall for a long time after
death, and the paper-thin RV, if disarticulated, is easily broken. Dead shells of N.
radula are found on exposed bottoms andbeaches, but they were certainly transported
from sheltered environments by hermit crabs.

Various epibionts are observed on the surface of both valves of this oyster. Bryozoans
and serpulids are most common, and certain
species of scleractinian coral, barnacle andsclerosponge are occasionally found (Figure7). A characteristic community consisting ofsclerosponges, articulated brachiopods,
bryozoans, annelids and scleractinian corals(commonly noncolonial) are found at or near
the habitat of the present oyster. Suchsclerosponge-bearing communities have been
discovered at similarly sheltered places (mainly crevices and caves) in a number of Caribbean and Indo-West Pacific coral reefs, andtheir ecological and paleontological significance as well as their origin have been
discussed by many authors (Hartman and
Goreau, 1970; Jackson, Goreau and Hartman, 1971; Jackson, 1977,1979; Kobluk,
1988; etc.). The present oyster is possibly
also a constituent of such a cryptic community at least in this region.
Color dimorphism. -Of eight large specimens examined, two are ivory white and six

1084

Itaru Hayami and Tomoki Kaseare purplish brown in shell coloration (Table1). There is no intermediate individual, andthe color type of infant individuals is difficult
to determine. The two phena are notdifferent in various characters other than shellcoloration. This fact suggests the presence ofdimorphism, but the sample size at each locality is too small to ascertain their strictly
sympatric relation and to clarify the relative
frequency. Because the habitat of this oyster
is very dark, selection can scarcely act on thevariation of shell coloration.
Shell growth. - The shell shape of the present oyster changes significantly with growth.
In the initial stage (up to 20-30 mm in shellheight) every individual attaches itself to thecave wall or some other hard object (including the shell surface of larger individuals)
with the whole surface of LV. In this stage
both valves are nearly flat or only feebly
convex, the commissural shelf is still narrow,and the test thickness is almost the same
between the two valves. Although theattachment area extends along the entire dorsal margin of LV until the latest stage, themain part of the generated shell margin of LVsits up suddenly apart from the wall surface,
and then the shell becomes very inequivalve
and concavoconvex, as shown in Figure 7.
In some cases numerous infant shells of thesame oyster species adhere to the shell surface(Figure 2-la, c), but such individuals probably cannot grow larger because we havenever seen any cluster of large individuals.
Functional morphology. - As seen in thedorsal view of many specimens, the attachment area is commonly geniculated at theumbonal area (Figure 4-1). The chevronlike attachment area seems to strengthen theadherence. Divers could not collect any
specimen without using a hammer. In many
grown individuals of this oyster the ventral
surface of LV is strongly puckered, so that
this portion often looks branched in crosssection (Figure 7). This feature indicatesthat a preexisting ventral margin was abandoned, and that a new margin grew from theinside of the shell. The formation and function of the strongly puckered venter are probably similar to those of hyote spines in
Hyotissa (Stenzel, 1971: p. N 1026).
Repeated reconstruction of ventral margins
result in the stronger convexity of LV andshell thickening of the middle-ventral portion. Judging from the occasionally imbricated simple prismatic outer layer, reconstruction of venter may occur also in RV,
though the abandoned venter must be readily
broken.
The RV of this oyster is extremely thin andhas almost no physical strength, but its flexible distal part sticks so closely to the innersurface of LV that we can hardly examine thesoft parts without breaking the shell. The
strongly puckered venter of LV and unusuallybroad commissural shelf may be advantageous, because, as interpreted by Stenzel(1971 : p. N 1025), they make it more difficult
for shell-breaking predators to reach the soft
part. In fact, individuals with inj ured-andrepaired shell are often met with in thisspecies.
Radial gashes of RV, which were called"tuck grooves" by Harry (1985), are a very
characteristic feature in many pycnodonteineoysters. As observed in the present oyster,
they occur exclusively in the simple prismatic
outer layer (Figure 2-lc, d, 5). Because thislayer is commonly multilayered, the gashes
rarely penetrate the shell except near theventral margin. They may also contribute to
the increase of flexibility of the distal part.
In many fossil species of Pycnodonte, RV
looks much smaller than LV. The presenceof radial gashes, however, strongly suggeststhat those species had originally the flexible
distal part of R V and concordant valve margins like the present species.
Evolutionary ecology. - In view of the very
conservative morphology, the present oyster
can be regarded as directly descended fromsome Cretaceous and Paleogene species of
Pycnodonte (s.s.). This, however, does notnecessarily mean that fossil species of935.

A living fossil oyster

Pycnodonte were also cryptic animals. In
fact, many small individuals of a pycnodonteine oyster, as recognized by Hanai andOji (1981), attach themselves to the uppersurface of a fossil beachrock in the Aptian ofnorth Japan. Bottjer, Roberts and Hattin(1978) ill ustrated many individuals of
Pycnodonte kansasense [sic] encrusting thesurface of an inoceramid valve in the Turonian Greenhorn Limestone of Kansas.
Kauffman (1967) and Tashiro (1978) also
recorded similar occurrences respectively for
Pycnodonte? conjesta from the Colorado
Group in the Western Interior andPycnodonte amakusaensis from the Santonian muddy sediments of west Japan.
Kauffman and Sohl (1973) mentioned that
small individuals of Pycnodonte often attached themselves to the surface of lower
valves of rudists in the latest Cretaceous
rudist reefs of the West Indies. Jablonski
and Bottjer (1983), on the other hand, regarded Pycnodonte (Phygraea) mutabilis from theLate Cretaceous offshore chalky sediments of
the Gulf and Atlantic Coastal region as an
iceberg strategist (cup-shaped recliner by
Seilacher, 1984). This is probably true
about many species of the subgenus
Phygraea, because of their gryphaeate morphology. These Cretaceous species of
Pycnodonte, both sedentarists and recliners,
are thus not considered to have been crypticorganisms.
Typical Cretaceous and Paleogene speciesof Pycnodonte (s.s.) are generally characterized by a longer dorsal margin, better developed auricular part and commonly thinnershell than those of Phygraea. If the attachment area extends over the entire dorsal
margin with growth, those fossil species
should be regarded as lifelong sessile organisms like the present species. Cryptic life ofthese fossil species is still unlikely, because
they occur abundantly together with manyother epifaunal and infaunal bivalves.
Palmer and Ftirsich (1974) examined themicrogeographic distribution of various en-1085

crusting organisms on hardground and insmall crevices at the base of the Bradford Clay
(Bathonian) in England. As mentioned by
Boucot (1981), this seems to represent an
exceptional preservation of ancient shelteredbiota. Palmer and Ftirsich recorded that theupper (exposed) surface and the lower(sheltered) surface of the crevices were inhabited by more or less different organisms. An
unnamed species of Plicatula, a brachiopod,
some serpulids and bryozoans could be cryptic organisms because they predominantly
occur on the lower surface of crevices. Some
oysters belonging to Liostrea, Exogyra,? Nanogyra and Lopha are occasionallyfound adhering to the lower surface, but theyoccur more abundantly on the exposed surface of the hardground. These crevices are
only about 25 cm deep and measure up to 5
cm from roof to floor; their scale must be by
far smaller than that of the submarine caves inwhich the present oyster dwells.
Why is the present oyster found only in
sheltered places? If Cretaceous-Paleogene
species of Pycnodonte were noncryptic, whatdrove the descendant to such a cryptic environment? There are two possible answers.
One is an explanation regarding the change of
habitat as a result of overgrowth competition
among reef-building organisms; that is, inmodern coral reefs, as Jackson (1977) interpreted for the restricted distribution ofsclerosponge-fauna, rapidly encrusting colonial organisms predominantly occupy thesurface of exposed hard bottom, excluding
therefrom slowly growing sessile organisms.
The other explanation is based on theassumption that the predation pressure onbivalves is relatively low in such a shelteredenvironment.
Many sedentary bivalves of similar size are
known in and around Indo-West Pacific coral
reefs. Spondylus varius, Hyotissa hyotis andChama lazarus are often found alive on the
exposed fore-reef bottom around the caves.
Although little is known about the growth
rates of these bivalves, it is unlikely that these1086

ftaru Hayami and Tomoki Kase

noncryptic bivalves grow much more rapidly
than the present oyster. On the other hand,
they are characterized by much thicker shells
and more strongly armoured upper valves.
The strongly puckered venter and broad
commissural shelf may be effective to someextent against predation, but the shell of this
oyster appears to be more delicate anddefenseless in comparison with such massive
sedentary bivalves in the exposed environment. Though nothing is known about theactual predators of this oyster, these lines ofevidence seem to suggest that the second
explanation is more plausible in this case.
Submarine cave faunas often containdeeper-water elements. Kobluk (1988)
attributed the cause of resemblance between
cryptic and deeper-water faunas mainly toanalogous physical factors, especially to similarly low illumination level. Some submarine caves of Ryukyu Islands yield many
incredibly extraneous bivalve genera and families, the bathymetric distributions of which
extend to or are restricted to lower neritic,
bathyal and even abyssal bottoms (Hayami
and Kase, 1991 ; Kase and Hayami, in press).
Deep-sea origin of the present oyster, however, is rather unlikely, because there is noevidence for Pycnodonte to have migrated todeep seas after the Paleogene. It may havesurvived for a long geologic period by wandering sublittoral cryptic environments.

Conclusion: significance as a living fossil
Although there are several extant representatives of the Pycnodonteinae (Neopycnodonte and Hyotissa), this subfamily seems
to have attained its acme in Late Cretaceousand Paleogene times. The genus Pycnodonte(including several subgenera) was a leadinggroup of this subfamily during these periods,
but there is no fossil record after the Middle
Miocene. As described above, the newly
discovered archaic oyster is considered to be adirect descendant of Pycnodonte (s.s.). Its
shell morphology may look enigmatic at a

glance, but many important characters of thissubgenus are well retained. In addition to
the archaic morphology and evolutionary
stasis, its cryptic habitat strongly suggests thatthe present species is a significant example of"living fossils". Its morphology, physiology
and ecology must be informative for understanding the paleobiology of fossil
Pycnodonte and related oysters.
Many Cretaceous and Paleogene species of
Pycnodonte, however, were probably noncryptic organisms, because they occur very
abundantly together with other bivalves invarious kinds of sediments. Two different
life habits are assumed for fossil species of
Pycnodonte. One is iceberg strategists (secondary cup-shaped recliners), and the other islifetime sedentarists like the present species.
Iceberg strategy is assumed in various familiesof Mesozoic bivalves (Jablonski and Bottjer,
1983; Seilacher, 1984), but is believed tohave become much rarer in the Cenozoic,
probably as the result of the increase of
powerful predators (Hayami and Hosoda,
1988).
As advocated by Vermeij (1977,1987) andothers, the increase of predation pressure after
the Mesozoic seems to have had a serious
impact on the evolution of adaptive strategy
of molluscs. Bivalve evolution seems to havefollowed several trends not only in morphology but also in behavior and habitat. In
many free-living bivalves escaping ability
from predators (e.g., by burrowing and byswimming) has increased. Some sedentarygroups seem to have developed armoredsculptures against the attack of durophogous
predators. Another effective way of surviving must be to change habitat to a place oflower predation pressure. Neopycnodonte,
which is probably also a descendant of
Pycnodonte, now survives in deeper water
than other oysters. It is generally supposedthat submarine caves and other cavities aswell as exposed deep-sea bottoms often provide suitable refuges for archaic and relatively
defenseless organisms.

935.

A living fossil oyster

The inferred habitat change of Pycnodonte
is noticeably similar in many respects to thatof Neritopsis, which is a famous "living fossil"
gastropod (Batten, 1984). Neritopsis radula,
which is often found alive together with thepresent oyster in the same caves, shows very
conservative morphology. Fossil records of
Neritopsis are common in the Mesozoic andPaleogene (especially in the Tethyan realm),
but N. radula from the Indo-West Pacific anda closely related Caribbean extant species are
the only known representatives after the
Miocene. To conclude, it is strongly suggested that Pycnodonte and Neritopsis have beensurviving in shallow and warm seas for many
millions of years by transforming themselves
into cryptic animals. We may be able tohave a glimpse of Mesozoic-type biota in sucha cryptic environment. Further studies oforganisms in submarine caves would provide
concrete and significant evidence to test various theories in evolutionary biology.

Acknowledgements
We thank sincerely Dr. Kenji Torigoe ofHiroshima University for his kind information about the anatomical characters of thepresent oyster. Our cordial thanks are also
due to skillful divers, Messrs. Mitsutoshi
Taniguchi (Miyako-jima), Shuichi Ohashi,
Shigemitsu Kinjo and Hiroyuki Kinjo (Okinawa), and Miss Miyoko Uchima (Okinawa)for their self-devoting cooperation in collecting the material.
POSTSCRIPT. At the preparation of this manuscript weoverlooked an important work (Dhondt, 1984: The
unusual Cenomanian oyster Pycnodonte biauriculatum. Geobios, Mem. special, no. 8, p. 53-61), in
which the morphology, distribution and paleoecology
of a Cenomanian oyster Pycnodonte (Pycnodonte)
biauriculatum (Lamarck, 1819) were described anddiscussed in detail. It was said that the species
distributes widely (though episodically) along thenorthern margin of the Tethys from the Iberian Peninsula to Central Asia, sometimes forming crowded
fossil banks as a monospecific community. The shell
shape and long dorsal margin of P. (P.) biauriculatum

1087

resembles those of the present species, but its LV isvery thick like Phygraea. It is unlikely that the
Cenomanian oyster was a cryptic organism, as Dhondt regarded the species as having been an iceberg
strategist on a soft substrate. She treated the genderof the generic name Pycnodonte as neuter, though weare ignorant about the reason.

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935.

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1089

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pIs. 55-62.

Ie-jima WiI~, Miyako Islands -gtl7iJ~, Okinawa Island i'rpi':llli~, Ryukyu Islands l'frE:E:j(
Seragaki Beach miN)'i[~' -7-, Shimoji-shima Ti1!!.~.

~~,

l'frE:E:j(~~tJ'G9£JiI.~nt-:'t:·;7 / r"/T~O)Iffi.1:.f! (1:.§"·"Cl'91~:O;l;""'): -gtl7IJ~0)
Ti1!!.~:t-o J: lfi'rpi':llli~ ® 1*:0) iWl.iti1PJilIl (i*1tfJ 20 m) 0)7W1fili l '~iii v.::. tJ, t.>: I) :kml 0) JiI,,\§}1I nt.>: l,;I; "'" tJ~9£JiI, ~ nt-:.o IDtMjffi· ~l'UIfl:l'tI\:~ • ~ff{ffl>a:~W L tda:llt .: O)f! ft ~ 'Y ::I '7 ;iJ'""'" v.::.~ {, iii l ,tJ~, :m1:.O)Jlt~f! f.::.~~i- 9 {, 0) ftt.>:
B llUc -tl5t=*cv'::' T 7- A iW~-c'iJ\?i
L, ,*,ffit!fx~ll.jI( L L!: 'j!j xJ:d't -C §" t-:. Pycnodonte ~O)JEtff!-c' t5 9': C tJ~1ljj G tJ'v'::'t.>:--:;t-:. o Pycnodonte (Pycnodonte) laniguchii [TP~: :;;J-:;;J-~ 'Y ::I '7 ;1;'''''' (ffif;i];)] c~fHt<,*-C~cax: L,{-0)*1~tl1:.!\'kJ"f:l¥J~~>a:~~-t" 9 0{-O)Mtk· IDt13ftllt~v.::.@;tstJ~,

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<

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~O)7i-~t-': vttJ, G t.>: 9 :tiIDto) mH~mlfHl\;o) J: ') v.::.fi
1:.Bifv'::'Ii*~-c·, "jiif L IAl 'MQltiii>a:~-t"o LtJ'L Pycnodonte(s.s.) 0)-t""'-C0)!f!fW: (17UxJi, *l''W~V.::.i''d--:;-ClAtJ~9~~iii, 'W{J!U v.::. {:Ii: !i:-t" 9 F9MO) F,fJIDi:~njJt $.l' commissural shelf, ~tko) chomata, i[!!O)*tk

O).m~ >a: {, --:>IDi:F9 O)~i1PJ, :tiIDi:~iii 0) "jiif L l' 1.&MtkO)j!liJn EI)

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1090

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1092

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Jf; ..................... i19tj"IJ.Ij'f • ,*,tlttl¥iX&~ •• 133~5\
Benthic foraminiferal assemblages of two piston cores!tflll'ffittl~.:ttfJ.Jl~G').~............................................................ 1tE!jff',!'fA? '" -T;/, ::[1- '7;/ F &l;G')://vlv*lij¥,11:E~!5G')tI~~,*,Mlij¥,~!ill*i!ll:.l::'to vt;., &ii:~vt"""J't!jfft;*~WG')l:ltW.G')ft.m.ilf.G')~~~.

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7 ~.;; l' :1 ~1\:lI:.E G')ili'<# '7 ;/ ~ t tlffiJ:l[fi'i~............................................................ il'Ii~ imtaken from south of the Ishigaki Island·································Xu Xuedong· Ujiie Hiroshi

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Cenomanian/Turonian J:l[~I::'tovt ;.,l!7t* ~ &1*lt A /~~Hil?rSoft bottom l::.tovt;.,1jt1*~MIIlIIl\\JlG') growth form....................................................... ··~l33bl*c'f............................................. i')Jj(I!j!f~ • Lll Q jlfz

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Palaeontological Society of Japan (PSJ) Council Actions
During its meeting on January 24, 1992, the PSJ Council enacted the followingchanges to its membership.
New mem bers elected :Yuuji Abe,

Toshiaki Futagawa,Kouzi Hasegawa,Tomoteru Hiyoriyama,

Yoshiyuki Hayashi,

Yoshinori Hikida,

Haruki Horikawa,

Masumi Ichikawa,

Shigeru lnaba,

Seiji Kakuta,

Takafumi Katoh,

Sanao Kashikura,

Hakuichi Koike,

Minoru Koitabashi,

Ken-ichi Kouko,

Katsue Koya,
Takehiro Morishige,

Yukihide Matsumoto,
Masafumi Murayama,

Hiroshi Matsuo,
Takehiro Nakamura,

[093Takashi Nakashima,
Itsuro Oshiro,
Takeshi Saito,
Takehiro Sato,
Harumi Sugawara,
Isamu Terui,
Hideo Yabe,

Ricardo Franco-Nieto,
Johann G. Rigor,
Takeyoshi Saito,
Chieko Shimada,Ken Takagi,
Yutaka Tsubaki,
Koichi Yoshiba

Hiroyuki Oishi,
Kazuyoshi Ryuzaki,
Tak uj i Sasaki,
Hiroko Suzuki,
Michio Takahashi,
Shigeki Togashi,

New Fellows elected;
Kazumi Akimoto,
Tomoko Matsuda,
Tomio Nakagawa,
Hiroshi Nokariya,
Osamu Sakamoto,Susumu Tomida,

Ren Hirayama,
Atsushi Matsuoka,
Hideo Nakaya,
Masayuki Oishi,
Tokiyuki Sato,
Akifumi Tomizawa,

Ryuichi Majima,
Keij i Matsuoka,
Harufumi Nishida,
Kiyoshi Okumura,Toshio Takagi,
Y oshitaka Yabumoto

Resigned members;(Fellow)
Kazumi Suyari(Ordinary members)
Tsutomu Kinoshita,

Hidekazu Yoshida

Deceased members;(Fellow)
Toru Makino(Foreign members)
Li-Sho Chang

1094

Bibliography 1986-1990 (7) flJ ff ,: ""'J \, '-r
B*<5i£4&.J't~-C'f:l:.,1986-1990~0) 5~rs'IC0~~ht.:~Jt(mt'!AAliJtuJ: (J'T ~:;/7 ~ 71 O)~o'l'$~~IfffJt'l¥) ~tlHJIHcJ: I) flJrr-f1.>':' C Ict! I) i Lt.:o ':'0) Bibliography ICf:l:.ltt*illi I) B*<5i£4&.J't~~Jl.O)
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1[if.5 0298-51-4320·51-4210[ fJU]WA TANABE Kozo (1973): Proflislilinella assemblage in the Omi Limestone, Niigata Prefecture, central
Japan (Studies of Carboniferous fusulinacean of Omi, part I),

Transactions and Proceedings of the

Palaeontological Society of Japan, New Series, no. 95, pp. 371- 394, Plates 51-53, Figure I, Tables I-I L
Carboniferous

Fusuline

BANDO Yuji and KATTO Jiro (1980): On the Upper Triassic ammonoids from the Sampozan Group at
Hitsuzan, Kochi City in Shikoku.
Jiro Katto.

In A. Taira and M. Tashiro (eds.): Selected papers in honor of Prof.

Geology and Paleontology of the Shimanto Belt.

(~:mm~LIJ 0) :=:::i:llJ!illf J: I) Il€tfj Lt.::=::~*e1&JtIlo) 7 /'.f: 7 1

Triassic

Ammonoidea

Rinyakosaikai Press, Kochi, pp. 95-100.

r IC"':) 0'--C) (J,E,)

1095

HANZA W A Shoshiro (1961): Cretaceous and Tertiary three-layered larger Foraminifera and their allied
forms: their classification and geographical and stratigraphical distributions.
Society of Japan), no. 2, pp. 1-24, Figures 1-29, Tables 1-2.

Fossils (Palaeontological(f&JtIlB.ilUc· ~=:*c=:Jl'Il*~1l1L!R&V'-i"O)ilIfbl:fif!0)5H]lt.;: IS> V'~.:.Jm:F1f1,¥i¥J • Jl'IlilL'¥i¥J5j';fp) (J.)
Cretaceous to Tertiary

Larger Foraminifera[~~t.,*, O)*)(~JHj# ~ ;fO)(~Iiii)( O){JU]

KOBA Y ASH 1 lwao and KAKIZAKI Takeo (1978): [Preservation of vertebrate bones in nodules from the
Tsurushi Formation]'

Saito Ryojiro Sensei Taishoku Kinenshi, pp.43-49, Plates 1-2.

(~TJW'@: /:: ~ -/H.:.§J:!!!~nt.::1'tftEO)i*m (J.)
MioceneMammalia

Taxa index for the bibliography
ANIMALIA

GeneralProblematica, Trace fossils
and others
CHORDATA

Mammalia
Proboscidea
Artiodactyla

ANNELIDA
MOLLUSCA

Cephalopoda
Coleoidea
Ammonoidea
Nautiloidea
Mollusca excl. Cephalopoda
Bivalvia

Aves

Scaphopoda

Reptilia

Gastropoda

Amphibia
PiscesPolyplacophora and Monoplacophora
COELENTERATA

Osteichtyes

Anthozoa

Chondrichtyes

Zoantharia

PROTOCHORDATA

Tabulata

Graptolithia

Scleractina

Conodontochordata

Tetracorallia

ECHINODERMATA

Holothuroidea

Octocorallia
Stromatoporata

Echinoidea

Hydrozoa

Crinoidea

Scyphozoa

BRACHIOPODA

PORIFERABRYOZOAPROTOZOA excl. MASTIGOPHORA

ARTHROZOARadiolaria

Insecta

Rhizopoda

Crustacea

Foraminifera

Malacostraca
Cirripedia
Ostracoda
Branchiopoda
Trilobita

Fusuline
Larger Foraminifera
Benthic Smaller Foraminifera
Planktonic Foraminifera

1096

VEGETABILIAPTERIDOPHYT AGeneralProblematica and others
Palynology

Pteropsida
Articulatae
NON TRACHEOPHYTAANGIOSPERME

Charophyta
Calcareous Algae

Dicotyledoneae
GYMNOSPERME
Coniferopsida

Coccolithophoridae
Dinoflagellata

Ginkgopsida
Cycadopsida

Silicoflagellata

Pteridospermopsida

OthersDiatomae

Geologic age index for the bibliography
GENERAL OR INDEPENDENT TO AGE
PRE-CAMBRIAN
PALEOZOICOLD PALEOZOICCAMBRIAN

LATE CRETACEOUSMESOZOIC TO CENOZOICLATE CRETACEOUS TO TERTIARYCENOZOICTERTIARYCAMBRIAN TO ORDOVICIANPALEOGENEORDOVICIANSILURIAN
SILURIAN TO DEVONIANNEW PALEOZOICPALEOCENE
PALEOCENE TO EOCENE
EOCENE
EOCENE TO OLIGOCENE

DEVONIANOLIGOCENE

CARBONIFEROUS
CARBONIFEROUS TO PERMIANPERMIANOLIGOCENE TO MIOCENE
NEOGENE
MIOCENE
MIOCENE TO PLIOCENE

PALEOZOIC TO MESOZOICPERMIAN TO TRIASSICMESOZOICTRIASSICJURASSICJURASSIC TO CRETACEOUSCRETACEOUS
EARLY CRETACEOUS

PLIOCENE
PLIOCENE TO PLEISTOCENE
NEOGENE TO QUATERNARYQUATERNARYPLEISTOCENE
PLEISTOCENE TO HOLOCENEHOLOCENE----------------------------------0 ----------------------------------

Erratum

MA TSUMOTO, T A TSURO
On some acanthoceratid ammonites from the Turonian of Hokkaido.
Trans. Proc. Palaeont. Soc. Japan, N.s., no. 164, p. 910-927, December, 1991.
Figure 6 caption, printed on a gummed-label and inserted in this number,should be substituted for the erroneous figure caption appearing on
page 920 of No. 164.
----------------------------------0 ----------------------------------

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HTransactions and Proceedings of the PalaeontologicalSociety of Japan

New Series No. 165

April 30, 1992

CONTE:NTS
TRANSACTIONS
932.

933.

934.

935.

Kiyoshi Okumura and Yutaka Yamagishi: Molluscan fauna from the Late
Miocene Hatsuse Formation in the Miura Peninsula, Kanagawa Prefecture,
Japan .............................................................. 1009
Ryuichi Majima and Akihiro Murata: Intraspecific variation and heterochrony in Phanerolepida pseudotransenna Ozaki (Gastropoda: Turbinidae)
from the Pliocene Nobori Formation, Pacific side of southwestern Japan ... 1024
Katsumi Ueno: Verbeekinid and Neoschwagerinid fusulinaceans from the
Akiyoshi Limestone Group above the Parafusulina kaerimizensis Zone,
southwest Japan. . ................................................... 1040
Itaru Hayami and Tomoki Kase: A new cryptic species of Pycnodonte from
Ryukyu Islands: A living fossil oyster. . ............................... 1070

PROCEEDINGS .......................................................... 1090