ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 136

Pa laeontologica l Society of Japan
December 30, 1981

ltaru HAY AMI and Ikuwo OBATA

Co-Editors

Officers for 1983-1984
President: Y okichi TAKA YANAGI
Councillors:

Kiyotaka CHINZEI, Takashi HAMADA, Tetsuro HANAI, Yoshikazu HASEGAWA,

Itaru HAYAMI, Hisayoshi IGO, Junji ITOIGAWA, Tadao KAMEl, Tatsuaki KIMURA,
Tamio KOTAKA, Ikuwo OBATA, Tsunemasa SAITO, Tsugio SHUTO, Yokichi TAKAYANAGI, Toshimasa TANAIMembers of Standing Committee: Kiyotaka CHINZEI (General Affairs, Co-Editor of "Fossils"),
Takashi HAMADA (Foreign Affairs), Tetsuro HANAI (Member-at-large), Yoshikazu
HASEGAWA (Membership), Itaru HAYAMI (Co-Editor of Transactions), Hisayoshi IGO(Planning), Tatsuaki KIMURA (Finance), Ikuwo OBATA (Co-Editor of Transactions),
Tsugio SHUTO (Editor of Special Papers), Yokichi TAKAYANAGI (Co-Editor of"Fossils ")Secretaries: Katsumi ABE (General Affairs), Ienori FUJIYAMA and Juichi YANAGIDA (Special
Papers), Kunihiro ISHIZAKI ("Fossils"), Hiroshi NODA (Planning), Tatsuo OJI
actions)
Auditor: Toshio KOIKE

The ostracod carapace on the cover is an adult specimen of Manawa konishii
NOHARA (Suborder Palaeocopina, Family Punciidae) from the East China
Sea. (photo by K. ABE, x190)

All communication relating to this journal should be addressed to the

PALAEONTOLOGICAL SOCIETY OF JAPAN
c/o Business Center for Academic Societies,
Yayoi 2-4-16, Bunkyo-ku, Tokyo 113, Japan(Trans-Trans. Proc. Palaeont. Soc. Japan, N.S., No. 136, pp. 445-454, pIs. 82, 83, December 30, 1984

786. ECHINOIDS FROM THE CRETACEOUS HAIDATEYAMA GROUP,
EASTERN KYUSHU*KEISAKU TANAKA
Geological Survey of Japan, Ibaraki 305
MASA YUKI NODA
Minamioita Junior High School, Oita 870
andHITOSHI TANAKA
Institute of Geology and Mineralogy, Hiroshima University, Hiroshima 730

Abstract.
In this paper four species of spatangoid echinoids are describedwhich came from the Upper Formation of the Haidateyama Group in Oita Prefecture. They are three species of He teras ter, of which one is new, and one species
assigned to Pseudowashitaster. For the description of a new species, material fromthe Lower Cretaceous Arida Formation in Wakayama Prefecture also is complementally dealt with here. Three of the echinoid species occur in common in theLower Barremian of several areas in the Chichibu Terrane, Southwest Japan. Therefore, this combined with already reported ammonites strongly suggests an Early
Barremian age for the Upper Formation of the Haidateyama Group.

Introduction

echinoids have been obtained from the Haidateyama Group, some of which have been preliminarily reported by Noda (1977) withoutpalaeontological descriptions. This is a usefulpiece of information about the Lower Cretaceousechinoid faunas of Japan which are very inferior
to the molluscan faunas. The echinoid specimens
available for the present study were collectedby Seiji Kohi, a former pupil of one of us (M. N.)and by another of us (H. T.), but their specificidentification has been done by the senior author(K. T.) in cooperation with M. N. Thus, four
species belonging to two genera have been distinguished; two of them are identified with

The Lower Cretaceous Haidateyama Group in
the Chichibu Terrane of eastern Kyushu hashitherto been investigated from the standpointof stratigraphy by Fujii (1954), Teraoka (1970),
Tanaka and Ohta (1980) and Tashiro et al.
(1983). Nevertheless, palaeontological works
have not yet been completed, except for thedescription of an ammonite species by Noda
(1977) and that of trigonian species by Tashiro
and Matsuda (1983).
Recently, numerous specimens of fossil* Received August 15, 1983
445

Keisaku TANAKA et al.

446

Geological Setting

hitherto known species and another is new. In
this paper the four echinoid species are described
and their biostratigraphic implications are discussed. Furthermore, .the specimens from theBarremian Arida Formation of the Yuasa area
Wakayama Prefecture are also dealt with here ~complemental material for a new species.
The specimens described here are kept at the
Geological Survey of Japan, Tsukuba (GSJ)
and the Institute of Geology and Mineralogy,
Hiroshima University, Hiroshima (IGH) , with
their respective abbrevations in parentheses.
Before going further, we wish to express our
sincere gratitude to Professor Emeritus Tatsuro
Matsumoto of Kyushu University for his valuablesuggestions and kindness in critical reading of thefirst draft. We also thank Mr. Seiji Kohi forsupplying the material for this study, and Mr.
Yoshio Masai of the Geological Survey of Japanfor photographing some of the specimens.

The Haidat~yama area represents one of theCretaceous outcrop belts in the Chichibu Terraneof eastern Kyushu. The most recent stratigraphic
context of the Haidateyama Lower Cretaceous
has been described by Tashiro et al. (1983).
The Lower Cretaceous clastic sequence rests
on pre-Cretaceous strata with an angular unconformity, forming an east-plunging syncline. It
is collectively called the Haidateyama Group.
According to Tashiro et al. (1983), this group is
stratigraphically divided into four units, provisionally named I (nonmarine), II (mostly brackish water), III (marine) and IV (marine) inascending order, totalling more than 900 m
thick. These four units are here termed theLower, Middle, Upper and Uppermost Formations. The lower three formations (Hauterivian? - Barremian) are in conformable sequence andthe Uppermost Formation (Upper Albian) is
faulted against them.
The Upper Formation from which the echi/

I-'"

SUKU BO ('<'Su-l02H-l06
Mt. Ho~doteyomo x~ ;~.,.

\

•

/"

"

~-~1-Su-l04\
t, KAMIKq.S.H IGOEr" ,--',/>.
1 ~ ....... _... __ . . . . ~. . . . . . .,....J-l,f"""'"
tJ"
~

'J\

H-l05
HK-204

t

','I
~,
~

'

o1...._ _ _-L-_ _ 2km~I

Text-fig. 1. Localities of echinoid specimens from the Haidateyama Group. Inset is a map of Kyushu, showing location of thestudied area (solid circle).

786. Cretaceous Echinoids from Kyushu

noid specimens examined came, consists chieflyof thickly alternating beds of sandstone andmudstone and is about 200 m thick. It yields
abundant shallow marine molluscan fossils.
The echinoid specimens for the present study
were obtained from locs. HK-204, H-105, H-106,
Su-102 and Su-104 of the Upper Formation(Lower Barremian) of the Haidateyama Group(Text-fig. 1). Of these localities, Noda's locality
is indicated by prefix HK and H. Tanaka's onesby Hand Suo
Locs. HK-204, H-105 (much the same as thepreceding one) and H-106 are situated at Honjomura, Minamiamabe-gun, Oita Prefecture. Loc.
HK-204 is occupied by siltstone with subsidiary
mudstone, and locs. H-105 and H-106 by siltstone to fine sandy siltstone. Locs. Su-102 andSu-104 are situated at Notsu-machi, Ono-gun,
Oita Prefecture. The former is occupied by finesandy siltstone and the latter by siltstone.
Especially locs. HK-204, H-105 and H-106 yielda number of echinoid fossils.
At all the echinoid localities various molluscan
species also occur abundantly. The occurrenceof Ancyloceras (Ancyloceras) sp. aff. A. (A.)
vandenheckii Astier at loco HK-204 (Noda, 1977)
and Crioceratites (Crioceratites) sp. aff. C. (C.)
koechlini (Astier) at loco H-105 (Matsumoto etal., 1982), among others, is worthy of mention.
Systematic Descriptions(by Keisaku Tanaka and Masayuki Noda)
Order Spatangoida Claus, 1876
Suborder Toxasterina Fischer, 1966
Family Toxasteridae Lambert, 1920
Genus Heteraster d'Orbigny, 1853
Heteraster macroholcus (Nisiyama)
PI. 82, Figs. 1, 2

1950. Washitaster (?) macroholcus Nisiyama,[nst. Geol. Palaeont. Tohoku Univ.,
Short Papers, no. 1, p. 44, text-figs. 4-6.
1965. Heteraster macroholcus, Tanaka, Trans.
Proc. Palaeont. Soc. Japan, N. S., no. 59,447

p. 133, pI. 16, figs. 3,4, text-fig. 4.
1968. Paraheteraster macroholcus, Nisiyama,
Palaeont. Soc. Japan, Special Papers, no.
13, p. 190, pI. 21, fig. 3; ibid., no. 11(1966), pI. 18, figs. 3, 12, 13.
1982. He teras ter macroholcus, Tanaka andKozai, Trans. Proc. Palaeont. Soc. Japan,
N. S., no. 126, p. 344, pI. 55, fig. 2,
text-fig. 2.
Materia/:-GSJ. F6024 and 6025, fromloco HK-204 (colI. Kohi); IGH:Hi-a-001 and
002, from loco H-105 (colI. H. Tanaka); IGH:Hib-001 and 002 from loco H-106 (colI. H. Tanaka).
Ten comparable specimens from loco HK-204,
two from loco H-105, one from loco H-106 andone from loco Su-102 were also examined.
Descriptive Remarks:-The specimens available are secondarily deformed. They are characterized by the deep and very long frontal sinus,
distinct anterior notch, very posteriorly placed
apical system, superficial paired ambulacra, anddeeply sunken peristome. Moreover, the frontal
ambulacrum shows an irregular alternation of
two to eight longer slit-shaped pores and one
shorter slit-shaped pore in the outer row andsomewhat inward disposition of pores (slightly
smaller or shorter than the normal round to,oval pores), opposite to shorter outer pores, inthe inner row (see Tanaka, 1965, text-fig. 4),
thus the pore pairs lining up in three (or betterto say rather four) files. The number of longer
pores alternating with a shorter pore varies fromplace to place in one specimen and also differs
between individuals, ranging from two to eightwith the commonest occurrence of two to five.
Furthermore, in a specimen (IGH:Hi-a-002)
represented by an external mould of the aboral
surface, a broad, diffuse peripetalous granularband or pseudo-fasciole is seen which is covered
with streaks of granules very slightly smaller thanthose covering the rest of the aboral surface anddecorated with tubercles.
Occurrence:-Locs. HK-204 (siltstone to mudstone), H-105 (siltstone to fine sandy siltstone)
and H-106 (siltstone to fine sandy siltstone);
Upper Formation of the Haidateyama Group,
Lower Barremian. One specimen probably re-448

Keisaku TANAKA et al.

ferable to this species is found in the fine sandy
siltstone of the same formation at loco Su-102.
Heteraster bungoensis Tanaka et Noda, sp. nov.
PI. 82, Figs. 3-6; PI. 83, Fig. 4; Text-figs. 2, 3

Materia/:-Holotype, GSJ. F6013, represented
by an internal mould, from loco HK-204 (see p.
447), (colI. Kohi); paratypes, GSJ. F6014 and
6036, represented by an internal mould, respectively, from the same locality (colI. Kohi). Designated as paratypes are also GSJ. F6033A, B(A, internal mould; B, imperfect external mouldof aboral surface) and GSJ. F6039A, B (A,internal mould; B, imperfect external mould of
aboral surface), from loco K-613 (= loco 1 in
Tanaka and Okubo, 1954), west of Kumai,
Yuasa-cho, Arida-gun, Wakayama Prefecture,
Upper Member of the Arida Formation, Lower
Barremian (colI. K. Tanaka). Three comparable
specimens from locs. H-106 and Su-104 (colI.
H. Tanaka) are also available.
Diagnosis:-Test medium-sized, heart-shapedin outline, somewhat rounded polygonal, weakly
emarginate in front, more or less longer thanwide, evenly rounded at the antero-Iateral
margins, rather straight at the postero-Iateral
margins, widest slightly in front of the midpoint,
contracted behind, relatively low; frontal sinus
rather short, somewhat broad, shallow, notsharply limited, expanding towards the ambituswith constant depth, extending to the peristome;
anterior notch shallow, moderately broad.

Aboral surface gently arched with steeper anterior slope, highest slightly behind the apicalsystem, abruptly truncated behind; posterior
truncated surface slightly concave; posterior
margin very shallowly emarginate. Oral surface
more or less inflated in the plastronal region andantero-Iateral regions, depressed around theperistome. Apical system small, slightly posterior,
ethmophract, with four gonopores arranged ina quadrate form; posterior genital plates mutually contiguous.
Frontal ambulacrum rather short, shallowly
sunken throughout its length, subpetaloid;
petaloid part extending about four-fifths theway to the anterior margin; poriferous zones
about half as wid~ as the interporiferous zone
at the middle; near the middle part double pairsof longer pores alternating with a pair of shorter
pores; near the proximal part and also near the
distal part a pair of longer pores alternating witha pair of shorter pores; longer pore pairs slitshaped, somewhat longer in the outer row thanin the inner row, somewhat circumflexed, widelyset; shorter pore pairs small, elongate oval tooval more or less longer in the outer row thanin the inner row, close together; pore pairs lining
up in four files.

.....§i§

-

~ a

c-'"
,-]

c

~m
L.......J

Text-fig. 3. Heteraster bungoensis Tanaka et Noda, sp. nov.,

Text-fig. 2. Apical system of Heteraster bungoensis
Tanaka et Noda, sp. nov.,
GSJ. F6039A, paratype.

GSJ. F6039B, paratype. a, Frontal
ambulacrum. b, Right anterior
ambulacrum. c, Right posterior
ambulacrum.

lmm

786. Cretaceous Echinoids from Kyushu

Paired ambulacra flexuous, superficial, subpetaloid. Anterior paired ambulacra nearly as
long and wide as the frontal ambulacrum;petaloid
parts rather long, extending about two-thirds
the way to the ambitus, open distally; poriferous
zones unequal, widest near the middle, tapering
towards the ambitus and the apical system;
anterior poriferous zones much narrower thanthe posterior ·zones which are narrower than theinterporiferous near the middle; pores in theanterior poriferous zones oval in the inner row,
short slit-shaped in the outer row, close together;pores in the posterior poriferous zones shortslit-shaped in the inner row, very long slit-shaped
and acuminated inwards in the outer row, rather
close together; pores of each pair somewhat
circumflexed, nonconjugate.
Posterior paired ambulacra somewhat shorter,
narrower, less flexuous and diverging at a smaller
angle than the anterior; petaloid. parts short,extending about halfway to the margin, nearly
two-thirds .as long as those of the anterior paired
ambulacra, open distally; poriferous zones unequal, widest near the middle, tapering towards
the ambitus and the apical system; anterior
poriferous zones much narrower than the posterior zones which are somewhat narrower than theinterporiferous near the middle; pores in theanterior poriferous zones oval in the inner row,
short slit-shaped in the outer row, closely set;pores in the posterior poriferous zones of slit
shape, much longer in the outer row than in the
inner row, rather close together; pores of eachpair more or less circumflexed, nonconjugate.
Peristome near the anterior fourth, small,moderately deep, subpentagonal, wider thanlong. Plastron mesamphistemous. Periproct rather large, transversely oval, high up on the posterior truncated surface, at half the height of thetest, partly visible from above. Tubercles small,
perforate, crenulate, scrobiculate; closely spacedon the plastron, widely scattered elsewhere;interporiferous zones of the paired ambulacra
dotted with smaller tubercles; numerous granules
between tubercles. No fascioles.
Measurements:-The specimens available are
deformed by the subsequent distortion. The

449

holotype represented by the least deformed
specimen is 23.4 mm long, 22.2 mm wide, and
9.2 mm high.
Remarks:-The holotype and other specimens
available from the Haidateyama Group are small(probably young) individuals. Accordingly, two
specimens (GSJ. F6033 and 6039; coli. K.
Tanaka), one of which is a much larger individual, and a comparable specimen (GSJ. F6008;
coli. K. Tanaka) represented by a relatively large
individual, from the Barremian Arida Formationin Wakayama Prefecture are also dealt with here.
The largest of the Arida specimens, though incomplete and secondarily depressed, probably
attains about 47 mm in length.
Comparison:-This new species resembles
Heteraster nexilis Nisiyama (1950, p. 42, textfigs. 1-3) from the Barremian of several areas in
the Chichibu Terrane, Southwest Japan, butdiffers from that species in having a test of
cordate outline, an apical system which is slightly
behind the centre, a rather short and shallow
frontal sinus, a shallow anterior notch and asubpentagonal peristome.
The present species is easily distinguishable
from the other two Japanese contemporary species, Heteraster macroholcus (Nisiyama) (1950,
p. 44, text-figs. 4-6; Tanaka, 1965, p. 133,
pI. 16, figs. 3, 4, text-fig. 4) and H. yuasensis(Tanaka et Okubo) (1954, p. 223, pI. 7, fig. 6,
text-fig. 7; Tanaka, 1965, p. 136, pI. 16, figs. 6,
7, text-fig. 6), in many respects. The apicalsystem is much less posterior and the frontal
sinus is much shorter and much shallower in the
new species with a test of cordate outline thanin Heteraster macroholcus with a test of ovate
outline. He teras ter bungoensis sp. nov. has
superficial paired ambulacra as against sunken
in H. yuasensis. The frontal sinus extends towards the anterior margin with constant depth,
forming a shallow but dis.tinct anterior notchin the present species, whereas in Heterasteryuasensis it becomes much shallower and indistinct near the ambitus, thus the anterior notch,
if any, being very indistinct. The new species has
a subpentagonal peristome, and both Heteraster
macroholcus and H. yuasensis have a transversely

450

Keisaku TANAKA et al.

oval peristome. As to the mode of arrangementof pore pairs in the frontal ambulacrum, Heteraster bungoensis as well as H. yuasensis shows
the Enallaster type (four files) of Devries (1955),
whereas H. macroholcus does show the Heteraster type (three files).
This species is similar to Heteraster lepidus(de Loriol) (1887-88, p. 91, pI. 15, fig. 4) fromthe Lower Barremian (Rey, 1972) of Portugal,
in the cordate outline of test, the shallow frontal
sinus and the shallow anterior notch. However,it differs from that species in that the frontal
ambulacrum shows Enallaster-type arrangementof pore pairs, the anterior paired ambulacra have
longer petaloid parts, and in that both the anterior and the posterior paired ambulacra havewider posterior poriferous zones whose outerpores are much longer than those of the anteriorzones.
The present species also resembles Heterastercouloni (Agassiz) (d'Orbigny, 1853, p. 179, pI.
848) from the Barremian of France and Switzerland, but is separated from that species by its
lower test, shallower frontal sinus, somewhat
shorter petaloid parts of the paired am bulacra ,less divergent posterior paired ambulacra, and by
its features of pore pairs in all the ambulacra.
Particularly, the new species shows Enallastertype arrangement of pore pairs in the frontal
ambulacrum as against Heteraster-type in Heteraster couloni.
Heteraster bungoensis is also akin to H.
pomeli (Ficheur) (1900, p. 592, pI. 10, figs.
5-8) from the Lower Aptian of Algeria, in theoutline of the test, position of the apical systemand features of the pore pairs in all the ambulacra. However, it differs from that species
in that the test is relatively low, the frontal
sinus is shallow and not constricted immediately
behind the anterior margin, and in that theanterior notch is shallow.
Occurrence:-Loc. HK-204; Upper Formationof the Haidateyama Group, Lower Barremian;siltstone. This species occurs also in the siltstoneof the Upper Member (Lower Barremian) of theArida Formation in the Yuasa area, Wakayama
Prefecture. Moreover, several specimens probably

identical with this species are found in thesiltstone of the Upper Formation of the Haidateyama Group at locs. iI-l06 and Su-104.

Heteraster sp.

PI. 83, Fig. 5
Material:-GSJ. F6027, represented by aninternal mould, from loco Su-l04 (colI. H.
Tanaka).
Description:-The specimen is poorly preserved and longitudinally compressed secondarily. Nevertheless, it measures about 12.5 mm
in length.
The test seems to have a rather heart-shaped
outline. The frontal sinus is rather shallow,
becoming much shallower towards the ambitus,
thus the anterior notch, if any, is probably
indistinct. The apical system is somewhat posterior and has four gonopores.
The frontal ambulacrum is rather shallowly
sunken and subpetaloid. Each poriferous zone
consists of about twenty pore pairs. A pair ofslit-shaped pores widely spaced regularly alternates with a pair of elongate oval pores close
together, the pore pairs lining up in four files.
However, at least in the left poriferous zone a
double pair of longer pores is placed at the
middle.
The paired ambulacra are somewhat flexuous,slightly sunken and subpetaloid. The anterior
paired ambulacra consist of anterior poriferous
zones with oval to elongate oval pores close
together and much wider posterior poriferous
zones with slit-shaped pores widely set. The
posterior paired ambulacra have much shorter
petaloid parts than do the anterior paired ambulacra. They consist of similar poriferous zones
with rather slit-shaped pore pairs.
Remarks:-The present specimen is so poorly
preserved and very small (probably young) thatthe specific identification is impossible. However,
it is very closely similar in many respects to andprobably the same species as certain echinoid
specimens that have been referred to as Heteraster sp. (Tanaka and Kozai, 1982, p. 345, pI.

786. Cretaceous Echinoids from Kyushu

55, fig. 4), from the Yunoki Formation (Upper
Barremian) of the Monobe area, Kochi Prefecture. It may also be noted that the present formshows some resemblance to a Heteraster species,
from the Upper Member (Lower Barremian)of the Arida Formation of the Yuasa area,
Wakayama Prefecture (Tanaka and Okubo, 1954,
p. 224, pI. 7, fig. 7, text-fig. 7), which has been
compared with Heteraster bohmi (de LorioI) (presumably synonymous with H. bravoensis (Bose)
according to Cooke, 1955), as is the case withthe above Monobe specimens (Tanaka andKozai, 1982). Anyhow, we refrain from further
taxonomic discussion of the present form,
because of the very small size and poor preservation of the material available.
Occurrence:-Loc. Su-104; Upper Formationof the Haidateyama Group, Lower Barremian;siltstone.
Suborder Hemiasterina Fischer, 1966
Family Hemiasteridae Clark, 1917
Genus Pseudowashitaster Tanaka, 1982
Pseudowashitaster mysticus Tanaka

PI. 83, Figs. 1-3
1982. Pseudowashitaster mysticus Tanaka in
Tanaka and Kozai, Trans. Proc. Palaeont.
Soc. Japan, N. S., no. 126, p. 348, pI. 55,
figs 5-7; pI. 56, figs. 1, 2, text-figs. 3, 4.
Material:-GSJ. F6021, 6022 and 6023, fromloco HK-204 (colI. Kohi); GSJ. F6026A, B (A,internal mould; B, imperfect external mouldof aboral surface), from loco H-105 (colI. H.
Tanaka); IGH: Hi-b-003A, B (A, internal mould;
B, external mould of aboral surface), from loco
H-106 (colI. H. Tanaka).
Descriptive Remarks:-The test, though incomplete and secondarily deformed, appears tohave an oval outline. The frontal sinus is very
long and deep, forming a conspicuous notch in
the anterior edge of the test. The apical system
is very eccentric behind and ethmophract, withfour gonopores. The frontal ambulacrum is very
long, deeply sunken and composed of oval inner451

pores and slit-shaped outer pores which arescarcely or occasionally set more'or less inwards.
The paired ambulacra are very unequal in length
and width, flexuous, subpetaloid and open distally. The anterior paired ambulacra are very
long, considerably narrow almost throughout theway but rapidly widened near the ambitus andare very slightly sunken. The posterior paired
ambulacra are short, broad and flush. All thepaired ambulacra consist of anterior poriferous
zones with minute, round pore pairs and muchwider posterior poriferous zones with longer andelongated pore pairs. The peristome is far anterior. The periproct is oval, vertically elongate
and high up on the posterior truncated surface.
The plastron is mesamphisternous. The peripetalous fasciole is distinct, narrow and continuous.
The largest one (GSJ. F6022) of the specimens, though transversely compressed secondarily, probably attains a little more than 70 mm
in length.
It has been pointed out by Tanaka and Kozai(1982) that Pseudowashitaster mysticus shows
intraspecific variation in arrangement of outerpores in the frontal ambulacrum. In one (GSJ.
F6021) of the specimens, for example, a shorter
pore is placed somewhat inwards three times atintervals of 12 to 14 longer pores within a seriesof 52 pores in the middle part of the rightpbriferous zone, and also three times at intervalsof 9 to 15 longer pores within a series of 33
pores in the middle to distal parts of the left
poriferous zone. In GSJ. F6026 one shorter poreis set more or less inwards every 7 to 11 longer
pores in the middle part of the right poriferous
zone. On the other hand, IGH: Hi-b-003 has
only two shorter pores within a series of about
50 pores in the middle part of the right poriferous zone.
Occurrence:-Locs. HK-204 (siltstone to mudstone), H-I05 (siltstone) and H-I06 (siltstone);
Upper Formation of the Haidateyama Group,
Lower Barremian.
Concluding Remarks
Among the echinoid specimens examinedKeisaku TANAKA et aZ.

452

from the Upper Formation of the Haidateyama
Group, the following four species are describedin this paper:
Heteraster macrohoZcus (Nisiyama)
Heteraster bungoensis Tanaka et Noda, sp.
nov.
Heteraster sp.
Pseudowashitaster mysticus Tanaka
Of the above four, Heteraster macrohoZcus isnumerically dominant. This species is known tooccur in the Middle to Upper Members (= Am to
Au in Obata and Ogawa, 1976) of the Arida
Formation of the Yuasa area, Wakayama Prefecture (Nisiyama, 1950; Tanaka and Okubo, 1954),
in the lower part of the Ishido Formation of theSanchu Graben, 8aitama - Gumma Prefecture(Tanaka, 1965), and also in the Monobe Formation of the Monobe area, Kochi Prefecture(Tanaka and Kozai, 1982). These three echinoid
horizons are assigned to the Barremian, strictly
speaking, to the Lower Barremian respectively(Obata and Ogawa, 1976; Obata et aZ., 1976;
Tashiro et aZ., 1980).
Pseudowashitaster mysticus was reported
from the Yunoki Formation of the Monobe area
(Tanaka and Kozai, 1982), the lower part of the
Ishido Formation (Tanaka, 1965), and also fromthe Upper Member of the Arida Formation(Tanaka and Kozai, 1982). The first of theseformations is correlated with the Upper Bar-

remian (Tashiro et aZ., 1980).
Heteraster bungoensis sp. nov. is found not
only in the Haidateyama Group but also in theUpper Member (Lower Barremian) of the Arida
Formation.
To sum up, so far as the material available isconcerned, it ,is evident that all the three species
mentioned above are restricted to the Barremianand moreover, occur in common in the Lower
Barremian.
Various molluscan species occur abundantly
at the echinoid localities in the Upper Formationof the Haidateyama Group. Among them,
AncyZoceras (AncyZoceras) sp. aff. A. (A.)
vandenheckii Astier, though somewhat imperfect, was considered as suggestive of the uppermost Barremian or the lowermost Aptian (Noda,
1977), but is not convincing. Very recently
another diagnostic ammonite, Crioceratites(Crioceratites) sp. aff. C. (C.) koechlini (Astier),
was obtained from about the same locality as
the above ammonite, and it suggests an Early
Barremian age (Matsumoto et aZ., 1982). Thus,
the Haidateyama echinoid faunule is compatible
with the ammonoid-based correlation. It is also
not inconsistent with the bivalve-based age assignment by Tashiro et aZ. (1983). In short, it
is reasonable to correlate the Upper Formationof the Haidateyama Group with the Lower
Barremian.

Explanation of Plate 82
Figs. 1,2. Reteraster macroholcus (Nisiyama) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 447
1. GSJ. F6024 from loco HK-204, Haidateyama area, aboral (a) and left lateral (b) views,
x1.3.

2. GSJ. F6025 from loco HK-204, aboral (a) and oral (b) views, x 1.1.
Figs. 3-6. Heteraster bungoensis Tanaka et Noda, sp. nov . . . . . . . . . . . . . . . . . . . . Page 448
3. GSJ. F6013, holotype, from loco HK-204, Haidateyama area, aboral (a), oral (b), leftlateral (c), anterior (d) and posterior (e) views, x 1.4.
4. GSJ. F6036, paratype, from loco HK-204, aboral (a) and oral (b) views, x 1.6.
5. GSJ. F6033, paratype, from loco K-613, Yuasa area, aboral (a), oral (b) and left lateral(c) views, xL 7.
6. GSJ. F6039A, paratype, from loco K-613, aboral view, xL
All specimens illustrated here are internal moulds.

Plate 82

TANAKA et al. : Cretaceous Echinoids from Kyushu

30

3c

3d

3e
5c

786. Cretaceous Echinoids from Kyushu
References
Cooke, C. W. (1955): Some Cretaceous echinoids
from Americas. U. S. Geol. Suru., Prof.
Paper, 264-E, p. 87-112, pIs. 18-29.
Devries, A. (1955): Note sur Ie genre Heteraster.
Bull. Soc. Geol. France, ser. 6, vol. 5, p.
315-323.
Ficheur, E. (1900): Notes sur quelques Echinides
nouveaux de l'Aptien d'Algerie. Bull. Soc.
Geol. France, ser. ~, vol. 28, p. 590-596,
pIs. 10, II.
Fujii, K. (1954): .Stratigraphy and geologicalstructure of the Usuki area, Oita Prefecture,
Kyushu. Part I. Jour. Geol. Soc. Japan, vol.
60, p. 413-427 (in Japanese with English
abstract).
Loriol, P. de (1887-88): Recueil d'etudes
paleontologiques sur la faune cretacique duPortugal. Vol. 2: Description des Echinodermes. Comm. Trau. Geol. Portugal, 122 p.,
22 pIs.
Matsumoto, T., Obata, I., Ohta, "I., Tamura, M.,
Matsukawa, M. anq. Tanaka, H. (1982):
Correlation of marine and . non-marine
formations in the Cretaceous of Japan.
Fossils, no. 31, p. 1-26 (in Japanese withEnglish abstract).
Nisiyama, S. (1950): Two new species of the
Toxasteridae from Japan. Inst. Geol. Palaeonto Tohoku Uniu., Short Papers, no. 1, p.
42-47.
--(1966): The echinoid fauna, from Japan andadjacent regions. Part I. Palaeont. Soc. Japan,
Special Papers, no. 11, p. 1-277, pIs. 1-18.
- - (1968): The echinoid fauna from Japan andadjacent regions. Part II. Palaeont. Soc.
Japan, Special Papers, no. 13, p. 1-491, pIs.
19-30.
Noda, M. (1977): A brief note on Ancyloceras
from the Haidateyama Formation, Kyushu.
Trans. Proc. Palaeont. Soc. Japan, N. S., no.
104, p. 418-423, pI. 44.
Obata, I., Matsukawa, M., Tsuda, H., Futakami,
M. and Ogawa, Y. (1976): Geological ageof the Cretaceous Ishido Formation, Japan.
Bull. Natn. Sci. Mus., ser. C (Geol.), vol. 2,
p. 121-138, pIs. I, 2 (in Japanese withEnglish abstract).
- - and Ogawa, Y. (1976): Ammonites biostratigraphy of the Cretaceous Arida Forma-453

tion, Wakayama Prefecture. Bull. Natn. Sci.
Mus., ser. C (Geol.), vol. 2, p. 93-110, pIs.
1-4 (in Japanese with English abstract).
Orbigny, A. d' (1853-60): Paleontologie franraise: Description des animaux inuertebres.
Terrain cretace, Echinides. Vol. 6, p. 1-598,
pIs. 801-1006 [Completed by G. H. Cotteau] .
Rey, J. (1972): Recherches geologiques sur Ie
Cretace inferieur de I 'Estramadura (Portugal).
529 p., 22 pIs. Laboratoire de Geologie del'Universite Paul Sabatier.
Tanaka, H. and Ohta, Y. (1980): Geology of the
Haidateyama area, Ohita Prefecture. Part II,
Stratigraphy and structure of the Early
Cretaceous Haidateyama Group. Bull. Fukuoka Uniu. Educ., vol. 30, pt. 3, p. 91-102(in Japanese with English abstract).
Tanaka, K. (1965): Cretaceous echinoids fromthe Sanchu graben, central Japan. Trans.
Proc. Palaeont. Soc. Japan, N. S., no. 59,
p. 126-142, pIs. 15,16.
- - and Kozai, T. (1982): Some Cretaceousechinoids from the Monobe area, central
Shikoku. Trans. Proc. Palaeont. Soc. Japan,
N. S., no. 126, p. 341-355, pIs. 55, 56.
--and Okubo, M. (1954): On some echinoids
from the Palaeo-Cretaceous of the Yuasa
district in the Kii Peninsula and of the
Yatsushiro district in Kyushu. Jour. Geol.
Soc. Japan, vol. 60, p. 215-227, pI. 7 (inJapanese with English abstract).
Tashiro, M. and Matsuda, T. (1983): A study of
the Pterotrigoniae from Japan. I, Taxonomy.
Mem. Fac. Sci., Kochi Uniu., ser. E, Geol.,
vol. 4, p. 13-52, pIs. 1-13.
- - , Okamura, M. and Katto, J. (1980): A
biostratigraphical study of the Lower
Cretaceous formations of Monobe area,
Kochi Prefecture. In Taira, A. and Tashiro,M., eds., Geology and Paleontology of the
Shimanto Belt - Selected Papers in Honor
of Prof. Jiro Katto, p. 71-82, pIs. 10,11 (inJapanese with English abstract).
- - , Tanaka, H. and Matsuda, T. (1983): The
stratigraphy of the Cretaceous System ofHaidate-yama area, Oita Prefecture. Res.
Rept. Kochi Uniu., vol. 32, Nat. ScL, p. 4754 (in Japanese with English abstract).
Teraoka, Y. (1970): Cretaceous formations in454

Keisaku TANAKA et al.
the Onogawa basin and its vicinity, Kyushu,
Southwest Japan. Rept. Geol. Surv. Japan.

no. 237, 87 p. (in Japanese with English
abstract).

Arida 1im, Banjo 1I1lE, Haidateyama mtlUJ, Honjo *1lE. Ishido :fi'§t, Itaya :tlii~, Kamikoshigoe
ffi$. Ono :k!f. Sanchu UJ9=>,
Shinkai ¥I'M. Sukubo ~RI*, Yuasa
Yunoki til!/
J:1Jl~, Koshigoe 1lI!~. Monobe ~$. Minamiamabe mita$. Notsu

rim.

*

:tL1'li~$t7)""f$S!IE*m;flUJII~iI' b.i£: L.t..: ry =. {t:fi: :k;t!Ji!<Il)f;J;:5(:1fit;::;t;{fl-t 0 r$S!lI!*mtlUJTitlfll) J:$/iliI' b.i£: L.t..: ry =. {t:fil;:: "?~ ''L. Heteraster Il) 3:f!i (I~HIl:f!i·~:f!i·
F<iJ~T-~~:f!i) ]10: Pseudowashitaster Il) 1 flI (ret~fI) ~ilIll~1j 1..., il1=.~~~~c.~rr.." t..:.
Heteraster 1l)~:f!i'tfllllf.l:UJ!Ji!<rlm:!t!!~ Il) ;fj'm~Il)J:1IIl1I <r$ ... ~ v ~ 7 /') ii' b ~.i£:-t o.
Heteraster 1l)fl!E?;illllt~X:1fill)ft!!:I1!!.lit1l) rllll"'~ v ~ "7 /'iI' b. Pseudowashitaster Il) 1 :f!i ~F<iJ C< rh'll·1:$ ... ~ v ~ 7 /'iI'b~b;h.'L~'o. m;flUJ.rtJ~Il)J:$II't7 /'~T1 H;::J:..,,'L
rilll"'~ v ~ 7 /,1;:::MLt ~;h. 'L:to I). ry =. {tllll)l\!I{;ZIt ':'Il)%j' Jt~~ C ~MI L.t.c: ~ '.
m9=>~1ft·ffm$z·m9=>f0

Explanation of Plate 83
Figs. 1-3. Pseudowashitaster mysticus Tanaka . . . . . . . . . . . . . . , . . . . . . . . . . . . Page 451
1. GSJ. F6022 from loco HK-204, Haidateyama area, external cast. aboral (a) and oral (b)views. x 1.2.
2. GSJ. F6026 from loc. H-105, Haidateyama area. 2a. Aboral view of an internal mould(A), xL 2b, Aboral view of an external mould (B). x 3. A peripetalous fasciole is seen
between two arrows.
3. GSJ. F6021 from loco HK-204, aboral (a) and oral (b) views. x 1.1.
Fig. 4. Heteraster bungoensis Tanaka et Noda. sp. nov. . . . . . . . . . . . . . . . . . . . . . Page 448
GSJ. F6014, paratype, from loco HK-204, Haidateyama area, aboral (a), oral (b) and leftlateral (c) views, x1.4.
Fig. 5. Heteraster sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 450
GSJ. F6027 from loco Su-104, Haidateyama area, aboral view, x2.2.
All specimens illustrated here are internal moulds, unless otherwise stated.

TANAKA et al.: Cretaceous Echinoids from Kyushu

Plate 834cTrans. Proc. Palaeont. Soc. Japan, N.S., No. 136, pp. 455-473, pis. 84-86, December 30, 1984

787. NOTES ON MYTILOIDES INCERTUS (CRETACEOUS BIVALVIA) FROMTHE UPPER TURONIAN OF THE POMBETS AREA, CENTRAL HOKKAIDO*MASA YUKI NODA
Minamioita Junior High School, Oita 870

Abstract. In this paper, Inoceramus incertus Jimbo, 1894 is restudied biometrically as well as taxonomically and biostratigraphically. At first normal distribution of various characters is evaluated for each sample from Pombets of
Hokkaido by the method of chi-square test, and the specific diagnosis is revised.
Then the extent of variation for selected characters are numerically shown. Finallythe biostratigraphic significance of the species is discussed.
The results obtained are as follows: (1) Inoceramus incertus Jimbo should be
assigned to Mytiloides. Almost all the specimens of so-called 1. incertus from Pombets are assigned to M. incertus which shows a considerable extent of variation inshell form and surface ornamentation. The extent of variation tends to expand inascending the stratigraphic sequence. (2) A few specimens from loc. Ik2014 are
distinctly outside the variation of M. incertus and should be called Mytiloides sp.
aff. M. mytiloidiformis (Trager). (3) M. fiegei (Trager) from the Upper Turonianof various regions of the world is probably regarded as a junior synonym of M.
incertus. Thus, M. incertus can be evaluated as one of the widespread indices of theUpper Turonian.

In troduction

date.
About 45 years later, Nagao and Matsumoto(1940) redescribed Inoceramus incertus Jimbo
on a number of specimens, including the syntypes and some subsequent collections from the
mudstone of particular exposures on the riverside of the Pombets, and ascribed the stratigraphic position of this species to the lowest partof the Upper Ammonite Beds.
Subsequently, Matsumoto (1959) referred
the species to the lineage of 1. labiatus - 1. latus
and ascribe.<i its range probably to the Upper
Gyliakian in the Japanese scale which is approximately correlated with the Upper Turonianin the international scale on the ground of theassociated ammonoids.
Since 1975, I have investigated the exposure

Inoceramus incertus was established by Jimbo(1894) on the basis of three incomplete specimens from a single calcareous nodule. According to the original description, the nodule wasobtained in the pebbles of the River Pombets.
As pointed out by Matsumoto (1963), Jimbo's
original figure (pI. 8, fig. 7) was syntheticallyreconstructed based on these specimens. Jimbo
mentioned that the complete outline was exactly unknown, but that it was distinguishedfrom Inoceramus sp. (Yokoyama, 1890) in thecrowded minor concentric ornamentations. The
geological age was not precisely known at that
*Received Oct. 7, 1983: Read June 29, 1980 at
KochL
455

Masayuki NODA

456

with Matsumoto with assistance of a numberof co-operaters. On the large samples in theseand previous collections, I examined statisticallythe variation and the ontogenetic change of
selected characters. In this paper, I described
the result of this statistical study, giving remarkson the relations with previously known species.
The stratigraphic implication of this and related
species is discussed.
Incidentally, a part of the result of this study
has been preliminarily reported in a short paper
by Matsumoto and Noda (1983), with emphasison biostratigraphic aspects. This paper gives afull account for the palaeontological aspects.
Before going further, I thank Emeritus Professor Tatsuro Matsumoto of Kyushu University
for his cordial help throughout the present study
including comments on manuscript and supply
of a large number of specimens registered at
Kyushu University (GK) for this study. I am
indebted to Professor Itaru Hayami of Universityof Tokyo for his kindness in giving facilities forthe access to the syntypes of Jimbo and usefuladvice, and also to Professor Masayuki Tashiro of
Kochi University for his generosity to let mequote his unpublished data. My gratitude isextended to Messrs. Kikuo Muramoto of Mikasa
Museum, Takemi Takahashi of Mikasa andShigehiro Uchida of Horomui for their help in
the field work. I express my sincere gratitude toProfessor Karl-Armin Trager of Bergakademie
Freiberg for his kindness in offering the replicaof certain specimens from the Upper Turonianof East Germany. Furthermore, I am muchobliged to Mr. Yoshihiro Matsubara, the Chief ofIwamizawa Forestry Office and other staff
members for giving facilities in the field work.
A part of this study was supported by scientific research subsidy of the Shimonaka Commemorative Fundation (1978, 1980).
Locality
The outcrops of localities Ik2012-2014
are on the right bank of the River Pombets,
about 3 km upstream from Ikushumbets,
central Hokkaido, as shown in Text-fig. 1. They

are in Ikushumbets Quad. (1:25000); Long.
141°58'46"N-50"N, Lat. 43° 16'20"E-23"E.
The main part of Ik2011 is on the opposite bankof Ik2012, the cliff of a forest road along the left
side of the River Pombets. The sequence of strata
exposed on the cliff is shown in Text-fig. 2 (seealso Matsumoto, 1965; Matsumoto et al., 1981).
The stratigraphy and the geological age of thisarea have al~eady been reported by several researchers (Matsumoto, 1959; Matsumoto, 1965;
Matsumoto, 1971; Matsumoto et al., 1981;
Matsumoto and Noda, 1983). The whole sequence of the fine-sandy siltstone exposed fromloco Ik2012b to Ik2014, below the 5 m bed ofgreen sandstone at loco Ik2012a and above themassive or thickbedded pebbly coarse-grainedsandstone at loco Ik2015, is referred to Unit
IlIa', the lower part of the Upper Yezo Group.
Considering the lithology and the mode ofpreservation, Jimbo's type specimens must havebeen derived from somewhere in the cliff ofIk2012-2014.
The associated species of ammonoid and othermegafossils are listed with their frequency by
Matsumoto et al. (1981).
Material and Method
The sample size and the stratigraphic positions
are shown in Table 1. The repositories of theexamined specimens are as follows: GK: Kyushu
University; HK: Hokkaido University; JG: Jonan
Geological Association, Oita; IGPS: Tohoku University; TTC: Takemi Takahashi Collection,
Mikasa; UMUT: University Museum, Universityof Tokyo.
In this paper, at first, the specific characters
are examined by means of traditional method,in addition to the numerical analyses of variation
and relative growth, and then compared withallied species.
At first, several well preserved specimens ofMytiloides. incertus are selected, and their linear
dimensions and angles are measured as shown in
Text-fig. 3.
Based on the measurements, (1) the meanvdue, standard deviation and Pearson's coeffi-

787. Mytiloides incertus from Hokkaido

/

457

i

./

!

/
Tertiary

IlIa'

IId'

o' ' - - - - -_ _---l,100mi.
Text-fig.1.

Route map along the River Pombets (after Matsumoto, 1965).

Qr.... / ,,"@Q:r:./nr~~.". '. 'V. :;-/:.::~..

.......-

i-c-Lb..La-J

L'klOIl----.l. . . o~Lc-.J
Llk20111'k2012J

Text-fig. 2.
1981).

1-------____

11<2013_ _ _ _ _ _ _ _ _..;....._ _ _ _"'20'. ___--'-:'"'0"

Sketch of the outcrops where Mytiloides incertus occurs (after Matsumoto et al.

cient of variation of the selected biometricalcharacters are obtained for each sample, andthe normal distribution for each character isevaluated by means of chi-square test, (2) the
difference between the mean values of eachcharacter is evaluated by Student's t-test, (3)for the samples of which the t-value is not
significant, the two samples are combined andthe mean value, standard deviation and Pearson's

coefficien t of variation are obtained, (4) thesignificant difference from some samples of allied
species is examined by t-test for each biometricalcharacter, (5) the two variables, shell height (h)and shell length (1), and H and L for eachspecimen are concerned with examination ofrelative growth, plotting h, (H) in abscissa and 1(L) in ordinate on logarithmic graph paper,and the striking features of the reduced major

Masayuki N ODA

458

Table 1. Sample size of each outcropping unit.
Ik2014

exposure

Ik2011

subdivision

c

d

e

g

h

b

d

f

9

2

1

6

15

2

4

2

numbertotal

Ik2012

Ik2013

(2)

(1)

(10)

11

1

34

total

41
(13)

8

54

( ): not allocated to a subdivided unit.

Text-fig. 3.
ments.

Basic morphology for measure-axes and the coefficients of correlation arecalculated.
For the method of measurements and theprocedure of statistics and their analyses, refer toHayami (1969), Hayami and Matsukuma (1971)
and Noda (1975).
And also, for the comparison with the allied
species described by Trager (1967), the relative
growth between 8 and h , and that between l /hand h are shown in the same way as Trager's.
Palaeontological Description
Family Inoceramidae Giebel, 1852
Genus Mytiloides Brongniart, 1822

Mytiloides incertus (Jim bo)
PI. 84 ; PI. 85; PI. 86 , Figs. 1-8.

1894 . Inoceramus

incertus

Jimbo.

Palaeont.

Abhandl., Ne ue Folge , vol. 2, no. 3, p.
189 [43], pI. 24 [8], fig. 7.
1930. Inoceramus inconstans inconstans Woods ;Fiege, Palaeontographica, vol. 73, p . 38, pI.
5, figs. 16, 17, pI. 6, fig. 18.
1940. Inoceramus incertus Jimbo; Nagao andMatsumoto, Jour. Fac . Sci., Rokkaido
Imp. Univ., ser. 4, vol. 6, no. 1, p. 10-13,
pI. 3, figs . 1-3, 5, pI. 10, fig . 2.
non 1940. Inoc eramus incertus Jimbo ; Nagao
and Matsumoto, op . cit. , ser. 4, vol. 6,
no. 1, pI. 3, fig. 4.
non 1940. Inoceramus incertus var. yubarensis
Nagao and Matsumoto , op. cit. , ser. 4,
vol. 6 , no. 1, p . 11 , 12, pI. 6, fig. 1.
1959. Inoceramus incertus Jimbo ; Matsumoto,Mem. Fac . Sci., Kyushu Univ., ser. D,
vol. 9, no. 2, p. 85.
1963 . Inoceramus incertus Jimbo; Matsumoto,
A survey of th e fossils from Japan illus trated in classical monographs. Part 10,
p . 45 , pI. 67 , fig. 7.
non 1966 . Inoceramus sp. cf. incertus; Tamura
and Tashiro, Mem . Fac . Educ ., Kuma moto Univ ., no . 14, sec . 1, pI. 1, figs. 1113.
1967 . Inoceramus fiegei fiegei Trager. A bh.
Staatl. Mus . Mineral-Geol., vol. 12, p .
105-108, pI. 11, fig . 3, pI. 13 , figs. 14,
15, 17,20.
1976 . Inoceramus incertus Jimbo ; Noda andMatsumoto , Atlas of Japan ese fossils. no .
4 5, sheet 270, fig. 7.
1977 . My ti/oides(= "Inoceramus")incertus(Jimbo); Kauffman, Palaeont. Soc . Japan,
Special Papers. no. 21, p . 179, 180.
1978. Mytiloides fiegei fiegei (Trager); Kauffman, Ann. Mus . N at. d 'Rist. Nic e, vol.
4, no . 23 , p. 9, pI. 15, fig . 1, pI. 16 ,

7B7. Mytiloides incertus from Hokkaido
fig. 4.

1982. Inoceramus fiegei fiegei Troger; Keller,
Geol. Jahrb., A, no. 64, p. 110, pI. 7,
fig. 5.

1983. Mytiloides incertus (Jimho); Matsumoto
and Noda, Proc. Japan. Acad., vol. 59,
ser. B, no. 5, p. 111, figs. 2-4.

Lectotype.-UMUT.MM7535. One of the syntypes (designated by Matsumoto and Noda,
1983, p. 111). Pebble of the River Pombets,
Mikasa City.
Material.-UMUT.MM7535-3 (= MM7481 =TK.I-151, including four specimens), GK.H10041,
10048, 10056, 10061-10064, 10066-10068,
10108, 10109, 10111-10129, HK7242, 7246,
JG.H2128, 2850-2855, TTCOOOI and 0002.
Diagnosis.-Shell of medium size, equivalve,inequilateral. Valve moderately inflated, anterior
part steep and posterior half gradually flattened,passing into a wing-like area without sharpboundary. Umbo terminal and not prominent.
Hinge-line rather long. Marginal outline withlarge extent of variation, being from oblique oval
to subquadrate. Surface ornamented with a combination of major concentric undulations andminor concentric rings. The former being low,broad and regularly spaced and the latter numerous, sharp and much crowded.
Description.-Shell of medium size, large

A

individual about 95 mm in H (JG.H2855, GK.
H10087 and 10109). Growth axis straight orslightly concave to anterior and considerably
oblique to the hinge-line. Valve gently inflated,
anterior part comparatively steep but someindividuals perpendicular to the valve plane(TTC0002), posterior half gradually flattened
and continuing to the wing-like area without
sharp boundary. The section along the growthaxis uniformly convex but abrupt change of convexity are observed in a specimen of TTC0002.
Outline considerably variable from oval (TTC0001) to subquadrate (TTC0002). The specimens
with subquadrate outline generally flat on flankand steep to the marginal part in adult, hingeline fairly long, about a half of shell length on
an average, forming an angle of about 1300 withthe posterior margin.
Concentric undulations mostly low and roundtopped with some extent of variation in size,interspace, intensity and density, for example,
as in the specimens of GK.H10109, GK.H10087,
GK.H10094, GK.H10123, GK.H10118 and TTC0001. Concentric rings distinct, sharp andcrowded in all specimens, and continuing fromthe main part of flank to the wing-like area.
Measurements and statistics.-The result of
the chi-square test for selected characters of eachsample is shown in Table 4. The chi-square valuesL

1=1. 489h

459

0.903

B
L= 1.570Ho. 850
r_ 0.9(J23r = 0.9300

.

,
.'
3

.'

"

2

1~1~0------~20----~--~-r~~~~~~

30

Text-fig. 4.

40

50 60708090100 h

1~1~0------~20----_~30---4~0~5rO-6~0~~~0-8rO~90~1~00~H

Averaged relative growth of Mytiloides incertus. A: I and h, B: Land R

Masayuki NODA460

TTCOOO2

GK.H10130&'\~

TTCOOO1

~

10mm
~

~

~

~

~

~ ~ ~
~0o ~ ~
Text-fig. 5. Ontogenetic change of shell convexity, showing the cross sections
at the growth stages of 7 mm, 10 mm, 20 mm and 30 mm in H.
Table 2. Biometrical characters of the Samples Ik2013 and Ik2011, and the result of Student's t-tests.

~

a

llh

'Y

HL/h

L/H

HL/I

Ik2013 IK2011 Ik2013 Ik2011 Ik2013 Ik2011 Ik2013 Ik2011 Ik2013 Ik2011 Ik2013 Ik2011

N

21

9

11

6

21

9

21

9

16

7

16

9

m

113°

114°

147.3

146.7

1.04

1.05

0.87

0.86

0.49

0.52

0.49

0.49

s

6.9

5.8

4.84

4.97

0.111

0.134 0.078

0.088

0.075

0.088

0.076

0.037

u

6.106

5.088

3.286

3.388 10.673 12.762 8.966 10.232 15.306 16.923 15.510

7.557

F

4.219

1.415

1.054

1.457

1.273

1.377

0

0

0

0

0

•

0.380

0.202

0.213

0.310

0.836

0.000

0

0

0

0

0

0

t

N: sample size, m: mean value, s: standard deviation, u: Peason's coefficient of variation, F: value ofF-test, t: value of Student's t-test, solid circle: significant, open circle: not significant. significancy showing at 95% confidence level.

of a, 'Y, 1/h, 1/hH=40mm, L/H and HL/h are lessthan 7.81 which is 5 percent significant limit at
the degree of freedom 3, and those of T/h and
HL/h are less than 3.84, the 5 percent significant
limit at the degree of freedom 1, thus the hypothesis of the random sampling from a normally
distributed population can be adopted. Therefore various numerical values, i. e. mean value,
standard deviation and coefficient of variation,
may be meaningful.
The measurement values for every specimen

are omitted for brevity, but those of the selected
specimens, that is, the lectotype, JG .H2853-2855
which are most common form and TTC0001,
0002, the extreme forms, are shown in Table 3.
Then F-test and t-test are adopted for characters
between the samples Ik2011 and 2013, the results are not significant for all characters examined. Thus the two samples can be combined
into a single one. The statistical result is showniii Table 4.
The sample Ik2014 is small, and its specimens

787. Mytiloides incertus from Hokkaido------

461

Table 3. Measurements of selected specimens of M. incertus. linear dimension in mm .

.

h

I

H

L

T

HL

Q(

'Y

{j

UMUT.MM7535-3
lectotype

38.4

35.5+

41.0

32.6

-

20.0

106°

148°

56°

JG.H2853
JG.H2854

54.0
54.0

58.9
55.5

66.0
61.5

53.0
49.3

-

27.0
27.4

116°
116°

150°

47°

24.6

150°

46°

TTC.OOOI

63.6

61.9

74.1

58.6

31.8

25.0

TTC.0002

65.1

65.5

69.0

67.0

30.0

30.9

106°
119°

150°
145°

54°
52°

--------

l/h

l/h H =40mm

L/H

T/h

HL/h

HL/I

UMUT.MM7535-3
lectotype

0.92+

-

0.80

-

0.52

0.56-

JG.H2853

1.09

0.46
0.50

0.50
0.51

TTC.OOOI

1.03
0.97

0.80
0.80
0.79

-

JG.H2854

1.08
1.12

TTC.0002

1.01

0.97

0.46

0.98
1.13

0.39
0.47

0.46
0.49
0.40
0.47

Table 4. Biometric characters of M. incertus.
Q(

'Y

l/h

l/h H =40mm

L/H

T/h

HL/h

HL/I

N

30

17

30

15

30

10

23

23

m

113.3

147.0

1.043

1.036

0.860

0.494

0.501

0.498

s

6.51

4.60

0.12410.10050.08080.0646 . 0.0784

0.0669v

5.745

3.129 11.898

9.701

9.395

13.077 15.649

13.681

X2

Table 5.

4.66

5.82

1.34

7.36

6.81

3.30

7.69

3.26

n=3

n=3

n=3

n=3

n=3

n=l

n=3

n=l

Student's t·test of the Sample Ik2014 for the Samples Ik2013 and Ik2011.

~

Ik2014
Q(

l/h

L/H

HL/h

HL/I

0.425

2.729

2.766

3.622

2.681

Ik2013
0

•

•

•

•

0.654

2.266

3.881

3.295

5.309

0

...

•

•

•

Ik2011are often of unfavourable preservation. In general
aspects, however, a form elongated along the
growth axis is more common than the lesselongated usual form. The results of t·test
between the sample Ik2014 and the sample

Ik2011 plus 2013 are significant with respects
to certain characters (Table 5).
Then the average relative growth of thesamples Ik2011 and 2013 are examined, plotting
h in abscissa and 1 in ordinate on loga,rithmic

Masayuki NODA

462

TTCOO 0 1

A

TTC0002
GK.Hl0119
JG . H285J.....

. . .-. - - -._-~:.=... . . . ~___""""--.--o-~--. -. -.-.

GK.Hl0130....................•
20

60

.40

80mm

H

B
100
8

....

TTC 000
TTC0002'.

------

GK.Hl0119~~-=.-.

JG.H2853
GK .Hl0130

60

20

.40

60

80mm

H

Text-fig. 6A. Ontogenetic ch an ge of obliquity. 6B. Ontogenetic change of L /H .

T ext-fig. 7.

Variation of shell form in My til oides incertus.

787. Mytiloides incertus from Hokkaido

graph paper, and also Hand L, and the reduced
major axes and the coefficient of correlation
are calculated. These are demonstrated in Textfig. 4A, B.
Text-fig. 5 shows the cross section at thegrowth stages of 7 mm, 10 mm, 20 mm and 30
mm in H and Text-fig. 6A shows the change of [)with growth and Text-fig. 6B that of 1/h.
Variation.-Although Jimbo's syntypes arevery incomplete, a considerable extent of variation in the outline is suggested by the curvatureof the concentric ornaments. In fact, examining
a large number of specimens there is considerably
wide variation in shell form as is demonstrated in
Text-fig. 7. The two extreme individuals may be
hardly identified with each other. There are,however, various grades of intermediate forms,
and normal distribution is confirmed in various
characters by the method of chi-square test.
Therefore, these various forms can be regarded
as representing one and the same species. On theother hand, some of the specific characters which
are not dealt with numerically are rather important as taxonomic criteria. As Matsumoto(1981, personal information) suggested, thesurface ornamentation of the specimen TTCOOOI
has its own characteristics, that is, concentricribs are low, crowded, round-topped and regularin size, interspace and intensity, whereas theminor rings rather inconspiquous. It differs
considerably from the typical form of thisspecies. The specimens GK.H10119 and JG.
H2850 show also similar ornaments. But in thespecimen GK.H10115 which is closed valves,the ornaments of right valve show an intermediate character whereas the left valve has
typical ornament. Furthermore, the right valveof GK.H10109 shows TTC0001-like ornamentin the umbonal part but has typical ornamentin adult stage. This fact proves a considerable
extent of variation even in the surface ornamentation.
Remarks. -Despite of the high variability,
no specimen of M. incertus resembles Jimbo's
illustration (reproduced here as Text-fig. 8)
which was inadequately restored.
The specimens described under I. incertus by

Text-fig. 8.

463

Restoration of My-

tiloides incertus based on three syntypes by Jimbo (1894).

Nagao and Matsumoto (1940, p. 10) are mostly
identified with M. incertus defined in this paper.
For instance, the specimens HK7246 and HK7242(Nagao and Matsumoto, 1940, pI. 3, figs. 1, 2)
are certainly so. The specimen IGPS22740(Nagao and Matsumoto, pI. 3, fig. 3) has comparatively sharp concentric ribs but can be
regarded as being within the extent of variationof M. incertus. HK7162 (pI. 3, fig. 5 in Nagao
and Matsumoto), which is an internal mould,
has also sharp concentric ribs but can be likewise
included in M. incertus, despite a different
opinion by Kauffman (1977, p. 180), because it
closely resembles the young part of JG.H2851(Pl. 84, Fig. 7). On the internal mould the surface
ornament would not be precisely impressed.
An exceptional specimen, IGPS22738 (Nagao
and Matsumoto, 1940, pI. 3, fig. 4), which wasnoted by Nagao and Matsumoto as an abnormal
form, is too much elongated along the growthaxis and probably identified with M. sp. aff. M.
mytiloidiformis (Trager) described below.
The specimen HK5960 (/. incertus var.
yubarensis Nagao and Matsumoto, pI. 6, fig. 1)
from Kamihobets is clearly distinct from M.
incertus as has already mentioned by Matsumoto·and Noda (1968).
Comparison with allied species. -Nagao andMatsumoto (1940) compared this species to M.
labiatus (Schlotheim) from the Lower Turonianand M. hercynicus (Petrascheck) from the comparatively upper part of the Lower Turonian toMasayuki NODA

464

pI. 13, figs. 14, 15, 17, 20) and that M. fiegei
is probably synonymous with M. incertus. The
measurements of the specimens including theholotype of M. fiegei fiegei and the specimens
JG.H2853 and JG.H2855 are shown in Table 6,
and the result of t-test between the two nominal
species in Table 7.
The numerical values in Tables 6 and 7 aresomewhat inaccurate, since the German specimens are not sufficiently numerous and theirmeasurements were made on the photographs
except for the paratype (replica).
As was illustrated by Troger (1967, pI. 13,
figs. 17, 20) a considerable extent of variation

the lower part of the Middle Turonian, andclarified the similarity and difference between
them. Matsumoto and Noda (1968) compared
the species to M. teraokai from the Middle
Turonian and they (1975, p. 205) examined theontogenetic change of obliquity in M. labiatus,
M. teraokai and M. incertus, and clarified thecharacteristics of respective species.
Matsumoto (1981, personal information)suggested me that the typical form of the present
species represented by the specimens JG .H2853
closely resembles the holotype of M. fiegei
fiegei (Troger) (Fiege, 1930, pI. 5, figs. 16, 17,
pI. 6, figs. 18, 19; Troger, 1967, pI. 11, fig. 3,

Table 6. Measurements of the typical form of M. incertus and the specimens of I. fiegei fiegei.
linear dimension in mm.
h

1

H

L

HL

a

'Y

li

c.i
.!:;

JG.H2853

54.0

58.9

66.0

53.0

27.0

116°

150°

47°

~

JG.H2855

88.1

71.7

95.0

67.5

34.5

112°

149°

51°

holotype

30.0

31.2

32.6

27.8

14.8

102°

151°

48°

No. 455D

50.6

47.0

57.8

45.8

21.9

116°

145°

47°

USNM 240344

58.4

59.5

67.2

55.3

24.8

107°

152°

47°

~

;;::.
...;

c.i
.!:;

~

~

;;::.

l/h

l/h H =40mm

L/H

HL/h

HL/I

JG.H2853

1.09

1.08

0.80

0.50

0.46

JG.H2855

0.81

0.97

0.76

0.39

0.47

holotype

1.04

-

0.85

0.49

0.47

No. 455D

0.93

0.94

0.79

0.43

0.47

USNM 240344

1.02

1.13

0.82

0.42

0.42

...;

Table 7. Student's t-test between M. incertus and I. fiegei fiegei.

acharacterspecies
N

m
s

l.f

M.i.

l/h

'Y

l.f

M.i.

I.f

M.i.

HL/h

L/H

I.f.

M.i.

I.f

M.i.

HL/I

I.f

M.f,

3
30
3
23
3
23
50
17
3
3
30
3
113.3 108.3 147.0 149.3 1.043 0.997 0.860 0:820 0.501 0.447 0.489 0.470
6.51
7.11 4.60 3.79 0.1241 0.0583 0.0808 0.0300 0.0784 0.0374 0.0669 0.0000

t-value

1.261

0.813

0.629

significancy

0

0

0

I

0.842

1.161

0.484

0

0

0

787. Mytiloides incertus {rom Hokkaido
in marginal outline is presumed in that species.
The ontogenetic changes of selected characters
are demonstrated in Text-figs. 5 and 6 in thesame way of Trager's expression for comparison
between the two species. The ontogenetic
changes of 8 and L/H (WA, Na/Ha by Trager)
confirm well between them. The abrupt changeof shell convexity on the ventral part as shownin p. 108, fig. 28 (Tr5ger, 1967) is sometimes
found also in M. incertits.
An extreme form represented by the specimens TTCOOOI and GK.H10119 apparently
resembles M. dresdensis dresdensis (Trager) in
the outline and surface ornamentations, but as isclear from Text-fig. 6A and B, the ontogenetic
changes of 6 and L/H differ from those of M.
dresdensis dresdensis (Trager, 1967, p. 205,
fig. 28).
I. striatoconcentricus striatoconcentricus GUmbel (1868) from the Upper Turonian of Europe
and Africa is similar to M. incertus in crowded
and sharp concentric iings, but differs in its
smaller ratio of L/H (0.70-0.85), smaller beakangle, prominent umbo and clearly demarcatedwing.
Another extreme form with subquadrate
outline represented by a specimen TTC0002
resembles M. aviculoides (Meek and Hayden)
from the Upper most Turonian of the WesternInterior Province (Kauffman, 1977b, pI. 12, fig.
1, pI. 13, fig. 2), but M. aviculoides is much more
oblique than M. incertus.
Matsumoto (1981, personal information)suggested me to compare I. sp. figured by BUrgI(1957, pI. 15, fig. 3) from the Lower Coniacianof Colombia to the present species but theformer is discriminated from the present species
in the smaller obliquity even in the early stageof growth and the posteriorly placed umbo.
Kauffman (1977a, p. 180) pointed out
the similarity between M. incertus and 1. (Pl.)
chouberti Sornay from the Lower Coniacian of
Tarfaya, Morocco and regarded the latter as asubspecies of the former. But there are somequestions for the subspecies which is clearly
age-inequivalent. In other alternative, 1. (PI.)
chouberti may be regarded as a chronological465

transient of M. incertus.
Although Kauffman (1977a) regarded thespecimen HK7162 (Nagao and Matsumoto, 1940,
pI. 3, fig. 5) as closely similar to or conspecificwith M. labiatiodi{ormis (Troger) (= 1. dresdensis
labiatoidi{ormis Trager), they are clearly discriminated from each other. Namely, the antero-dorsal margin of M. incertus is nearly straight
regardless of a large extent of variation in theoutline, whereas that of M. labiatoidi{ormis isconsiderably convex, and in addition, in the
ontogenetic change of L/H, M. labiatoidi{ormis
becomes gradually elongated along the growthaxis with growth.
Kauffman (1977a) pointed out that the specimen IGPS 57821 (Nagao and Matsumoto, 1940,
pI. 10, fig. 2) differs from the typical form of
M. incertus in the outline and surface ornamentation and that the specimen is very closely
allied to the specimens of M. {iegei {iegei. But
Kauffman's concept depended on the originalillustration of Jimbo (1894) is clearly inadequate, as has been mentioned above. Kauffman
regarded M. {iegei {iegei as a junior synonym of
M. meekianus (Anderson) from the "Coniacian"(age uncertain) of California. As is examined inTable 7, M. {iegei {iegei is a synonym with M.
incertus. Thus the specimen IGPS 57821 is also
ascribed to a form of M. incerus.
Some specimens (i. e. GK.H10119, TTC0001)
of M. incertus are similar to the holotype of
M. modaliaensis (Sornay) (Sornay, 1981, pI. 1,
fig. 1) from the Lower Turonian of Colombia in
the orientation of growth axis and shell convexity but M. modaliaensis differs from M.
incertus in its short hinge-line, convex anterodorsal margin and less conspicuous concentricrings.
Phylogenetic relationship. -Matsumoto (1959)
pointed out that the present species is closely
related to M. labiatus from the Lower Turonianand especially akin to M: latus ranging from theLower to Upper Turonian.
Kauffman (1975, 1978a, b) attempted precise
zonation of the Turonian of Europe and theWestern Interior Province by means of Mytiloides
species.

Masayuki NODA

466

A
90
80
70
60

B

L
I ~2.477h

90
80
70
60

0.699

r = 0.9995

L= 2.438H O•646

r= 0.9620

50

50

30

20

10~~~------~--~--~~~~~~~

7 8 9 10

Text-fig. 9.

20

30

40

SO 60708090

h

10

20

30

40

50 60 708090

H

Individual relative growth of Mytiloides sp. aff. M. mytiloidiformis. A: I and h,

B: Land H.

Matsumoto and Noda (1975) did not establish
zonal subdivision but recognized that "broadform" (with larger L/H) becomes gradually
predominant as we ascend the sequence. Although Kauffman (1977a, fig. 1) tentatively
suggested the tripartite zonation for the Lower
Turonian of Japan, it is practically difficult in
the case of Hokkaido.
In Shikoku, Aoki and Tashiro (1982) reported abundant occurrence of Mytiloides froma shaly bed of a,bout 20 m thickness at a locality
called Doganaro. They tentatively used thespecific names 1. (M.) opalensis (Bose), 1. (M.)
mytiloides (Mantell), 1. (M.) labiatus (Schlotheim), 1. (M.) subhercynicus (Seitz), 1. (M.?)
hercynicus (Petrascheck) and 1. (M.?) "latus"(Mantell). These species are overlapping considerably in stratigraphic range. *Furthermore, in the typical exposure of theLower Turonian in Saumur of France, Badillet
and Sornay (1980) recorded various species of
Mytiloides associating with one another andpointed out the impossibility of finer zonationof the Lower Turonian by means of Mytiloides
species.
Matsumoto and Noda (1968) pointed out thesimilarity of M. teraokai (Matsumoto et Noda)*Palaeon tological discussion on these species
should be made on another opportunity.

from the Middle Turonian of Kyushu, to M. latus(Sowerby) or to M. hercynicus (Petrascheck)(= M. latus by Seitz, 1921, p. 101) from thecomparatively upper part of the Lower Turonian.
The well-preserved specimens GK.H6836 (Matsumoto and Noda, 1968, pI. 32, fig. 4), JG.H2125(Matsumoto and Noda, 1975, pI. 18, fig. 7) of
M. teraokai closely resemble the typical form of
M. incertus, but M. teraokai has an anterior ear.
The morphological resemblance and the successive occurrence suggested the intimate relationship between the two species. There are, however, some questions to regard M. teraokai as
an immediate ancester of M. incertus. It would
be unnatural that the once acquired anterior ear
disappeared against the law of irreversibility. To
sum up, M. teraokai is probably an off-shoot,specialized from the main stock of evolution in
Mytiloides.
Seitz (1934) discriminated six varieties in a
species of M. labiatus (s. 1.), of which var.
subhercynica occurs in the upper part of thesequence (Troger, 1967; Kauffman, 1975) andshows a greater extent of variation (Seitz, 1934,
fig. 18).
In the case of M. incertus, the broad form
with a flat valve is also rare in the lower part
(Ik2014) and various forms occur in the upperpart.

787. Mytiloides incertus from Hokkaido

Considering the lineage of Mytiloides, various
forms of the genus from the Lower Turonian are
successively linked as species or subspecies,
although there is some disagreement amongauthors with respect to the taxonomic ranking.
According to Kauffman (1978a, b), M. subhercynicus, M. hercynicus and M. "latus" (sensu
Woods, 1911, fig. 41), which are all broad form,
are found in the Middle Turonian of the WesternInterior of U. S. A. and England.
From the above discussion, it is reasonable toregard M. incertus as an immediate descendantof a Middle Turonian species with a broad outline such as M. hercynicus or others.
Mytiloides sp. aff. M. mytiloidiformis (Trager)
PI. 86, Fig. 91940. Inoceramus incertus Jimbo; Nagao andMatsumoto, Jour. Fac. Sci., Hokkaido
Imp. Univ .• ser. 4, vol. 6, no. 1, p. 10-13,
pI. 3, fig. 4.
Material. -GK.H10130 from loco Ik2014. IGPS
22738 from Pombets.
Description. -Shell medium in size, probably
equivalve, much elongated along the growth axis,
with tongue-like outline. Umbo terminal, some·what projecting above the hinge-line. Umbonal
part considerably inflated. Antero-dorsal andanterior parts steep to the commissure plane,
posterior half decreasing the convexity, but
inclined fairly steep near the umbo and clearlybounded with wing-like area. Ventral part
gradually flattened. Growth axis fairly oblique
about 30°-50° in early stage of growth, at thegrowth stage of about 38 mm in H, the axis
nearly straight forming an angle about 60° withthe hinge·line. Antero-dorsal and anterior margins
broadly arcuate and bent abruptly continuing toacutely curved ventral margin, which in turn
passes into broadly rounded posterior margin.
Postero-dorsal margin nearly straight, forming
an obtuse angle about 150° with the hinge-line.
Hinge-line fairly short being about one thirdof shell length.
Surface ornamented with concentric ribs and467

rings in combination. The concentric ribs low,round-topped and regular in strength. The
concentric rings sharp-headed and crowded.
Wing-like area smooth without any- ornamentation. Weak radial ornaments discernible on
inner surface.
Measurements. -The measurements of thespecimen GK.H10130 are shown in Table 8 incomparison with those of the holotype of
M. my tilo idiform is. Then the individual relative
growth for hand 1, and Hand L are demonstrated in Text-fig. 9A and B respectively. The
ontogenetic change of shell convexity, I) andL/H are shown in Text-figs. 5 and 6 respectively
after Trager's (Troger, 1967, p. 201, fig. 24).
(Remarks: The value of L/H shown in Table 8
may be somewhat larger in actual specimen,
because the anterior part is lacking for a few
mm.)Discussion.-The specimen GK.H10130 was
firstly regarded as an extremely elongate formof M. incertus and was used for statistic analysis
altogether with other specimens. However, thevalue of L/H of GK.H10130 is outside thepossible range (m ± 3s) of M. incertus population, and the specimen, therefore, may have to
be excluded from the sample.
The result of measurements as shown in Table
8 and the ontogenetic change of selected charac·ters demonstrated in Text-fig. 9 are well agreeable with those of the holotype of M. mytiloidiformis (Trager) and also age-equivalent. But thefollowing distinctions are pointed out between
the two specimens. (1) The shell of the holotypeof M. mytiloidiformis is more strongly inflated
than that of GK.H10130. (2) The antero-dorsal
margin nearly straight in the holotype but
broadly convex in the Japanese specimen. (3)The surface ornamentation continues to thewing-like area in the holotype but the area issmooth in the Japanese specimen. (4) The majorribs regular in strength, size and interspace inthe holotype but irregular in the Japanese one.
To sum up, the present specimen GK.H10130
must be closely allied to but is not identified
with M. my tilo idiform is (Troger). However,
it is reserved, at present, to give a specific name

468

Masayuki NODA

Table 8.

Measurements of M. mytiloidiformis (Troger) and the specimen GK.H10130. linear
dimension in mm.

-------

Q

'Y

5

h

I

H

L

holotype

97°

150°

50°-70°

54.0

47.0

(64)

(36)

GK.H10130

110°

160°

60°

61.4

44.9

68.0

34.0

HL

l/h

l/h H =40mm.

L/H

HL/h

HL/I

holotype

17.0

0.87

(0.87)(0.54)

0.31

0.36

GK.H10130

12.7

0.73

0.87

0.50+

0.21

0.28

for the specimen because of the insufficientmaterial.
Troger (1967) established M. fiegei mytiloidiformis on four specimens from the Upper
Turonian of Sachsen, East Germany, and herecognized also M. fiegei fiegei from the same
bed. The sympatric occurrence of the two
subspecies, however, is very doubtful on biological ground. M. fiegei my tilo idiform is isclearly discriminated from M. fiegei fiegei by its
fairly prominent umbo, more oblique and longer
growth axis, and these distinctions are regarded
as specific criteria. From the above discussion,
therefore, it is more reasonable to separate thetwo forms as independent species.
Comparison with allied species. -The present
specimen is clearly discriminated from the comparatively "elongated" form of M. incertus inits still more "elongated" outline, prominent

umbo and fairly inflated umbonal region.
In comparison with such "elongated" formsof Mytiloides as M. labiatus, M. mytiloides(Mantell) and M. opalensis (Bose), the anterodorsal to anterior margin of the present specimen
is much more broadly arcuate. The results oft-test for selected characters between the specimen GK.H10130 and M. my tilo idiform is areshown in Table 9. As is clarified from Table 9,
there are certain characters with significant difference such as larger angle of 5 in GK.H10130
and small value in HL/h.
A specimen (Heine, 1929, pI. 5, fig. 27) of
M. sublabiatus (Muller) is similar to the holotypeof M. mytiloidiformis in the outline, shell convexity and obliquity of the growth axis, butother specimens (pI. 5, figs. 28, 29) have sharp
concentric ribs which continue to the wing-likearea. Heine ascribed these characters within an

Explanation of Plate 84(All figures natural size)
Figs. 1-10.Mytiloides incertus (Jimbo)
1. UMUT.MM7535-3, lectotype, one of the syntypes of the Jimbo's specimens, from
pebble in the River Pombets, Mikasa City, central Hokkaido (coll. Kotora Jimbo)
2. UMUT.MM7535-2, paralectotype, same nodule of the lectotype
3. UMUT.MM7535-4, non paralectotype, same nodule of the lectotype
4. UMUT.MM7535-1, paralectotype, same nodule of the lectotype
5. GK.H10124, right valve. Loc. Ik2013 (coll. T. Matsumoto, 1954)
6. JG.H2855, right valve. Loc. Ik2013h (colI. M. Noda, 1982)
7. JG.H2851, left valve. Loc. Ik2011c (coll. M. Noda, 1980)
8. JG.H2850, left valve. Loc. Ik2013 (coll. K. Muramoto, 1975)
9. GK.H10087, right valve. Loc. Ik2013g (colI. T. Matsumoto)10. GK.H10109, articulate valves. Loc. Ik2011c, dorsal view (colI. T. Matsumoto)NODA : Mytiloides incertus lrom Hokkaido

Plate 84

787. Mytiloides incertus from Hokkaido

extent of variation. These characters, however,are clearly distinct from those of the specimen
GK.H10130.
A specimen of M. striatoconcentricus striatoconcentricus (GUmbel) from Peru, figured byHeinz (1928, pI. 4, fig. 3) resembles the present
specimen in somewhat elongated outline andcrowded concentric ornaments, but the former
is clearly discriminated from the latter by welldeveloped posterior wing.
M. striatoconcentrlcus aff. carpathicus (Simionescu) (Heine,. 1928; Trager, 1967; Keller,
1982), from the Upper Turonian of Europe, issimilar to the present specimen in· ontogenetic
changes of 0 and l/h, but its value of L/H isabout 0.65-0.80 (Trager, 1967, p. 197, fig. 27)
as compared with 0.50 in the present specimen.
As is illustrated in the original paper, M. striatoconcentricus carpathicus (Simionescu) (1899, pI.
2, fig. la, b) has a smaller beak angle, long andstraight antero-dorsal margin, fairly strongly
convex ventral part, smaller posterior-hinge
angle and smaller obliquity with 0 about 90° .
M. labiatoidiformis (= 1. dresdensis ? labiatoidiformis Trager) is similar to the present
specimen in the mytiloid outline elongated along
the growth axis, the ontogenetic change ofobliquity and the presence of weak redial ornament on the inner surface of flank. But theformer is distinct from the latter in its convex
anterior margin, somewhat 'posteriorly placed
umbo, crowded concentric ribs which are sharptopped and regular in size and strength, andsimple ratio of L/H ranging 0.6-0.9.
Table 9.

469

Phylogenetic relationship. -Various species of
Mytiloides with elongated outline morphologically resemble the present form. According to
Kauffman (1977a, 1978a), the characteristic
species of Mytiloides are, in ascending order,
M. submytiloides (lowest part of the Lower
Turonian), M. opalensis (lower part), M. mytiloides (middle part), M. labiatus (upper part),
and broad form of Mytiloides becomes predominant in the lower part of the Middle Turonian.
On the other hand, the species of elongated typefrom the Upper Turonian are M. striatoconcentricus aff. carpathicus, M. labiatoidiformis
and M. mytiloidiformis, and further upward,
from the Lower Coniacian, M. sublabiatus.
The species with elongated form from theUpper Turonian may be related to one of thespecies of the Lower Turonian. As anotheralternative, they are not derived from a common
ancestor but of polyphyletic origin. The similarity may be caused from a similar mode oflife as Matsumoto (1959) suggested.

Geological range of Mytiloides incertus
Nagao and Matsumoto (1940) recorded thestratigraphic position of M. incertus as the lowest
part of the Upper Ammonite beds in the Pombets
area, and Matsumoto (1959) regarded it as representing the upper part of the Upper Gyliakian(approximately Upper Turonian in international
scale). Nevertheless, Kauffman (1977a) interpreted that the so-called "M. incertus" consistsof several heterogeneous species, and they may

Student's t-test for selected characters of M. labiatus and the specimen GK.H10130
from Hokkaido.
01

'Y

8

l/h

L/H

HL/h

HL/I

....'":s

N

5

5

5

5

5

5

5

.::

.c

m

114.0°

155.0°

36.4°

1.262

0.588

0.572

0.458

~

s

4.18

5.65

5.59

0.07300.10100.06560.0707.S!

GK.H10130

110°

160°

60°

0.73

0.50

0.21

0.28

t

0.874

0.807

3.854

6.652

0.795

5.038

2.298

significancy

0

0

•

•

0

•

A

470

Masayuki NODA

range from the Lower Turonian to possibly the
Middle Coniacian. Matsumoto et al. (1981)
examined the associated species of ammonites
from locs. 2014-2012 in ascending order, withthe result that the whole sequence comprisingthese localities belongs to the Upper Turonian.
The ammonites include such excellent indicesas Subprionocyclus neptuni (Geinitz), S. norma lis(Anderson) from loco Ik2014, Reesidites minimus(Hayasaka et Fukada) from loco Ik2012, Lymaniceras planulatum Matsumoto, Prionocyclus aberrans Matsumoto from locs. Ik2014-2012 andP. novimexicanus (Marc ow) from loco Ik2012.
Loc. Ik2011, shown in text-fig. 2 of Matsumoto
et al. (1981) should be read as 2011a' as corrected by Matsumoto and Noda (1983, fig. 1).
The three specimens recorded in Matsumoto etal. (1981, table 2) were obtained in talus probably originate from c bed of Ik2011, as is shownin Text-fig. 2b. Eleven specimens examined inthis study are all collected by Matsumoto fromthe c bed of the same locality. Matsumoto etal. (1981) pointed out the uppermost shaly
layer of about 5 m in thickness (loc. Ik2011a),which is below the unconformably overlying
Tertiary, is referred to the Lower Coniacian on
the basis of the occurrence of I. uwajimensis
Yehara (younger shell), Didymotis akamatsui(Yehara) and Baculites cf. yokoyamai Tokunaga
et Shimizu.
Meanwhile, the definition on the TuronianConiacian boundary and also that of the subdivisions of the Turonian are not always agreeable among authors. For example, the German
scale had long been used in a different way fromthe international one, until it was corrected bySeibertz (1979). We should take these points
into consideration 'in reading the German papers
published before 1980. As a matter-of-course,
Troger (1981) corrected his previous paper(1967), but his 1981 scheme is not exactly thesame as Seibertz's (1979).
To sum up, the lower part of the Upper
Turonian in the sense before 1979 corresponds
to the Upper Turonian and the upper part of theUpper Turonian in the same sense to the Lower
Coniacian.
According to Troger (1981), M. fiegei ranges
throughout the Upper Turonian and barely
persists to the Lower Coniacian, and M. mytiloidiformis is common in the lower part of theUpper Turonian and a few individuals survived
to the Lower Coniacian. According to the precise
data of Keller (1982), M. fiegei occurs in theUpper Turonian only.
In this paper, I regard M. fiegei fiegei (Troger)
as a junior synonym of M. incertus (Jimbo) andM. fiegei mytiloidiformis as an independent
species. The stratigraphic occurrence of these
species based on corrected scheme of Troger(1981) in Germany are quite in harmony withthose in Hokkaido.
Moreover, M. fiegei [= M. incertus] or M. aff.
fiegei occurs from various region of the worldsuch as central Europe (Keller, 1982; Troger,1967), the Western Interior of North America(Kauffman et al., 1978), Afghanistan (Sornay,1974) and Brazil (?). It characterizes the sequence of the Upper Turonian. It can be evaluated as one of the widespread indices of theUpper Turonian.

Explanation of Plate 85(All figures natural size)
Figs. 1, 2.. Mytiloides incertus (Jimbo)
1. TTCOOOL closed valves. An extreme form with oval outline, concentric ribs crowded
and concentric rings inconspicuous. Loc. Ik2013 (colI. Takemi Takahashi, 1978) A:
lateral view of the right valve, B: lateral view of the left valve, C: anterior view, D:
posterior view, E: dorsal view
2. TTC0002. closed valves. An extreme form with sub quadrate outline. Loc. Ik2013 (colI.
T. Takahashi, 1978) A: lateral view of the right valve, B: lateral view of the left valve,
C: posterior view showing the abrupt change of shell convexity, D: dorsal view

NODA: Mytiloides incertus from Hohkaido

Plate 85

787. Mytiloides incertus from Hokkaido
References cited
Aoki, T. and Tashiro, M. (1982): A stratigraphic
study of the Cretaceous Shimanto Belt (The"Doganaro" and Uwagumi Formation) at
Uwagumi, Kagami-machi, Kami-gun, Kochi
Prefecture, Shikoku. Res. Rept. Kochi Univ.,
vol. 31, (Nat. Sci.), p. 1-24, pIs. 1-7 (inJapanese with English abstract).
Badillet, G. and Sornay, J. (1980): Sur Quelques
formes du groupe d'Inoceramus labiatus
decrites par O. Seitz. Impossibilite d'utiliser
ce groupe pour une datation stratigraphique
du Turonien inferieur du Samuruis (France).
C. R. Acad. Sci. Paris, vol. 290, ser. D, p.
323-325.
Biirgl, H. (1957): Biostratigrafia de la Sabana de
Bogota y sus alrededores. Bol. geol. Bogota,
vol. 5, no. 2, p. 113-185.
Fiege, K. (1930): tJber die Inoceramen desOberturon mit besondere Briicksichtigung
der in Rheinland und Westfalen vorkommenden Formen. Palaeontographica, vol. 73,
p. 31-47, pIs. 5-8.
Giimbel, C. W. (1868): Verzeichnis der in derSammlung des Zool. Mineral. vereins in
Rengensburg vorfindlichen Versteinrungen.
Corresp. Bl. d. zool.-mineral. Ver. Rengensburg, 22, Jahrg.
Hayami, I. (1969): Mensuration of fossils andstatistics - An example of palaeontological
exercise -. Sci. Repts. Dept. Geol. Kyushu
Univ., vol. 10, no. 2, p. 67-90 (in Japanesewith English abstract).
- - and Matsukuma, A. (1971): Mensuration offossils and statistics - Analysis of allometry
and variation -. Ibid., vol. 10, no. 3, p.
135-160 (in Japanese w'ith English abstract).
Heine, F. (1929): Die Inoceramen des mittel
westfiilischen Emschers und unteren Unterosenons. Abh. Preuss. Geol. Landesanstalt, [N. F.], no. 120, p. 1-124, pIs. 1-20.
Heinz, R. (1928): Uber die Oberkreide Inoceramen Sudamerikas und ihre Beziehungen zu
dener Europas und anderer Gebiets, pt. 5,
Kenntnis ober Kreide Inoceramen. Mitt.
Min.-Geol. Staatsinst. Hamburg, 1928, no.
10, p. 1-39, pIs. 1-3.
Jimbo, K. (1894): Beitriige zur Kenntnis derFauna die Kreideformation von Hokkaido.

471

Palaeont. Abhandl., N. F., vol. 2, no. 3, p.
140-198, pIs. 17-25.
Kauffman, E. G. (1975): Dispersal and biostratigraphic potential of Cretaceous benthonicbivalvia in the Western Interior. Geol. Assn.
Canada, Special Paper, no. 13, p. 163-194.
--(1977a): Systematic, biostratigraphic andbiogeographic relationships between Middle
Cretaceous Euramerican and North PacificInoceramidae. Palaeont. Soc. Japan, Special
Papers, no. 21, p. 169-212.
- - (1977b): Illustrated guide to biostratigraphically important Cretaceous macrofossils,Western Interior Basin, U. S. A. The Mountain Geologist, vol. 14, nos. 3 and 4, p. 225274, pIs. 1-32.
--(1978a): (for 1976) British Middle Cretaceous inoceramid biostratigraphy. Ann. Mus.
d'Hist. Nat. Nice, vol. 4, no. 4, p. 1-12.
--(1978b): (for 1976) An outline of Middle
Cretaceous marine history and inoceramid
biostratigraphy in Bohemian Basin, Czechoslovakia. Ibid., vol. 4, no. 8, p. 1-12, pIs.
1-5.
- , Cobban, W. A. and Eicher, D. L. (1978):(for 1976) Albian through Lower Coniacianstrata and biostratigraphy, Western Interior
United States. Ibid., vol. 4, no. 18, p. 1-52.
Keller, S. (1982): Die Oberkreide der Sack-Mulde
bei Alfeld (Cenoman-Unterconiac.), Lithologie, Biostratigraphie und Inoceramen. Geol.
Jahrb., A., vol. 64, p. 3-171, pIs. 1-8.
Matsumoto, T. (1959): Zonation of the Upper
Cretaceous in Japan. Mem. Fac. Sci., Kyushu
Univ., ser. D, vol. 9, no. 2, p. 55-93, pIs. 610.
(1963): Matsumoto, T. ed. A survey of thefossils from Japan illustrated in classicalmonograph, part 10, p. 60-68, Palaeont.
Soc. Japan.
- - (1965): A monograph of the Collignoniceratidae from Hokkaido, part I. Mem. Fac.
Sci., Kyushu Univ., ser. D, vol. 16, no. 1,
p. 1-80, pIs. 1-18.
- - , Futakami, M., Tanabe, K. and Obata, I.
(1981): Upper Turonian ammonite assemblages in the Pombets area, Central Hokkaido.
Bull. Kitakyushu Mus. Nat. Hist., vol. 3,
p. 1-10 (in Japanese with English abstract).
- - and Noda, M. (1968): An interesting speciesof Inoceramus from the Upper Cretaceous

472

Masayuki NODA

of Kyushu. Trans. Proc. Palaeont. Soc.
Japan, N. S., no. 71, p. 317-325, pI. 32.
__ and - - (1975): Notes on Inoceramus
labiatus (Cretaceous Bivalvia) from Hokkaido. Ibid., no. 100, p. 188-208, pI. 18.
- - and - - (1983): Restudy of Inoceramus
incertus Jimbo with special reference to its
biostratigraphic implication. Proc. Japan
Acad .• vol. 59, ser. B, no. 5, p. 109-112.
Nagao, T. and Matsumoto, T. (1940): A monograph of the Cretaceous Inoceramus ofJapan, part 2, Jour. Fac. Sci., Hokkaido
Imp. Univ., ser. 4, vol. 6, no. 1, p. 1---64,
pIs. 1-22.
Noda, M. (1975): Succession of Inoceramus
from the Upper Cretaceous of Southwest
Japan. Mem. Fac. Sci., Kyushu Univ., ser.
D, vol. 23, no. 2, p.211-261, pIs. 32-37.
- - and Matsumoto, T. (1976): Atlas of Japanese fossils. no. 45, sheet 265-270 with
6 pIs. Tsukiji-Shokan, Tokyo (in Japanese).
Seibertz, E. (1979): Probleme der Turon-Gliederung Nordeuropas (Oberkreide) im iiberregional en vergleich. Ne wsl. Stratigr., vol. 7,
no. 3, p. 166-170.
Seitz, O. (1921): Die stratigraphisch wichtigen
Inoceramen des norddeutschen Turons.
Zeitsch. Deutsch. Geol. Gesell., vol. 73,
p.99-107.
--(1934): Die variabilitiit des Inoceramus
labiatus v. Schloth. Jahrb. Preuss. Geol. L.
A., 1934 (1935), vol. 55, no. 1, p. 426474, pIs. 1-9.
Simionescu, I. (1899): Fauna Cretacica Superiora

de la Urmos (Transylvania). Acad. Romana,
Pub. Fondului Vasille Adamachi, no. 4, p.
240-274.
Sornay, J. (1974): Inocerames Turoniens d'Afghanistan. Ann. Paleont. (Invertebres), 1974,
vol. 60, p. 27-34, pIs. 1, 2.
- - (1981): Inocerames (Bivalvia) du Turonien
Inferieur de Colombia (Ameriqui du Sud).
Ibid., vol. 67, no. 2, p. 135-148, pIs. 1, 2.
Tamura, M. and Tashiro, M. (1966): Upper
Cretaceous system of Kumamoto. Mem.
Fac. Educ., Kumamoto Univ., no. 14, sec.
1, p. 24-34, pI. 1 (in Japanese with English
abstract).
Troger, K.-A. (1967): Zur Paliiontologie, Biostratigraphie und faziellen Ausbildung derUnteren Oberkreide (Cenoman bis Turon)
Teil 1. Paliiontologie und Biostratigraphie
der Inoceramen des Cenoman bis Turon
Mitteleuropas. Abh. Staatl. Mus. Mineral.Geol., vol. 12, p. 12-127, pIs. 1-14.
- - (1981): Zur Problemen der Biostratigraphie
der Inoceramen und Untergliederung desCenoman und Turon in Mittel-und Osteuropa. Newsl. Stratigr. vol. 9, no. 3, p. 139156.
Woods, H. (1910-1911): A monograph of theCretaceous Lamellibranchia of England.
Inoceramus. Palaeontgr. Soc. Mon., London,
vol. 2, parts, 7, 8, p. 250-338, pIs. 45-54.
Yokoyama, M. (1890): Versteinerungen derJapanischen Kreide. Palaeontgraphica, vol.
36, p. 159-202, pIs. 18-25.

Explanation of Plate 86(All figures natural size)
Figs. 1-8, Mytiloides incertus (Jimbo)
1. JG.H2852, closed valves. Loc. Ik2013 (colI. M. Noda, 1980)
2. GK.H10094, closed valves. Loc. Ik2013d (colI. T. Matsumoto)
3. JG.H2853, right valve. Loc. Ik2013 (colI. M. Noda, 1980)
4. GK.H10112, articulate valves. Loc. Ik2013 (colI. T. Matsumoto)
5. GK.H10123, closed'valves. Loc. Ik2013 (colI. T. Matsumoto)
6. GK.H10118, closed valves. Loc. Ik2013g (colI. T. Matsumoto)
7. GK.H10111, articulate valves. Loc. Ik2011c (colI. T. Matsumoto)
8. GK.H10121, internal mould of closed valves. Lo':. Ik2013 (colI. T. Matsumoto)Fig 9.
Mytiloides sp. aff. M. mytiloidiformis (Troger) GK.H10130, left valve. Loc. Ik2014(colI. T. Matsumoto)NODA: Mytiloides incertus from Hohhaido

Plate 86

473

787. Mytiloides incertus from Hokkaido
Doganaro 'lit: 7" ~N~, Horomui .!J/iJ, Ikushumbets ~ff}Jlj, Iwamizawa;fM- J!1R, lonan iktm,
Kagami-machi 1t.fJ/; ~II/J. Kami-gun fi'Jyll1\, Kamihobets J:1!!!}]IJ, Mikasa ,::::1(t, Minamioita'f}, Pombets ;r-}jIJ, Uwagumi J:,fflm*~tw.@1H!u!1!!~'1- ='- - p :=. 7 /' J:i'lflEEO) Mytiloides incertus I:''?l '''C: *~-Clt Inoceramus inc(Jrlus Jimbo, 1984 1-=''?l,-e?t*O)liIf3C1.ii*I:.1JU:z. "C1:.4'hil!tl:lE~B':JJ!ttl!/H), it.:.., 1tE Jli'f"f:Ll'~J!:It!!1.J' i? jIj:~Jt Lt.:... i 1', 9¥}]IJ.!1!!~O)M'tllJi1.J' i? 0) -iT /' 7' IL.-I:'~f L "C, 7J
1 2 *~:lEI::' J: ") "C .f}JfHQ):. '?l '''C tEm0';fp0)r.i(~~11·l', J; ty t.:.\i)"CfilO):lE~-?:t~ L t.:...
;XI:.l' <'?1.J'O)MJ!{I:' '?l '''C, -to)%r~~~1t1:i¥J1:.1!Jl i?1.J'I:' Lt.:... ~ l <1:.~M0)1ttiOOff.~J¥J:®':.i'iH:' '?l '''C ilftli& Lt..:. .:c 0)#lP~11;;(0) J: 0 -C J;;o.

1. ~t*, Inoceramus incertus Jimbo c: ~J'l"Cl't.:..{,o)ltMytiloides incertus (Jimbo)
c: 1JSf';;;~ ~ -cJ; Q. .:;It i -C I. illcertus C: l 'hit "C l ,t.:..~}]IJEEO)~*O):7dm'f.tlt M. illcertus"'C' J:<, Jf~, :}H(fj'~~P.fIi C: t 1.J,tJ: I):*: ~ l '%rnta::'1j;-t. i t.:..J:(,zO)~l(IiO);t /' 7' IH1 cJ£r~1.Jt:*:~<tt.Q.
2. MO)}E~-?itl::''?l'''Clt*Jt (p. 459)

ta::'J!J:.

3. EEJi!! Ik20l4 1.J' i?.llffl:!l Lt.:.. Mytiloides 0) !il'l*0) 9=t I:' It I!Jl i?1.J'I:' M. illcertus O)lfr11<0) ? i? C: It~.Y,) i?ntJ:l' t O)1.J'l''?1.J'J; Q • .: nit Mytiloides sp. aff. M. mytiloidi/ormis(Trager) C: bl'?~~{,o)-CJ;7.J.
4. •• M. fiegei (nager)" C: 1.J' .. M. aff. fiegei" C: 1.J'P¥Iin "C l' Q b 0) /Jt1!!:J\!.O) l' -0l'-0tJ::It!!~O)'1- '" - p :=. 7 /' J:)')f)/J'i?¥Ri'i' ~n "Cl 'Zi. *1j}f3CO)~til!tll:' {, C: -? Or~~O)~Hi:~tdli
tJT~, -to)ft!!o;>t?ij·t/J'i? M.. fiegei (Trager), 1967 11:'l!:i? < M. illcertus 0)%4T~:/ I:=. A<

C:J!tJ:~;ltQ.

Lf.:..;Ij',,)"C, M. incertus It '1- '" - p:=. 7 /'l:iiIlta::'*H&-?'tQJJ.;;l,:It!!~I:.bt.:..

Q :fj'~.lJtJ:'lffJ'r./j~1tl1 C: l,x.. Q.

~ff:H~ZTrans. Proc. Palaeont. Soc. Japan, N.S., No. 136, pp. 474-491, pIs. 87-89, December 30, 1984

788. A MIDDLE DEVONIAN TRILOBITE FAUNA FROM THE KITAKAMIMOUNTAINS. NORTHEAST JAPAN -- I. THE LICHIDAE*
ATSUSHI KANEKO
15-7, Fukae-honmachi l~home, Higashinada-ku, Kobe, 658

Abstract. A trilobite fauna has been discovered from the Middle Devonian
Nakazato Formation in the southern part of the Kitakami Mountains in northeastern
Honshu, Japan. The fauna is composed of lichid, scutellid, dechenellid, calymenid,
phacopid, dalmanitid and odontopleurid trilobites from the Higuchi-zawa andOhmori-zawa, Hikoroichi~ho, Ofuna to City, Iwate Prefecture. The purpose of
the present paper is to give a description of the trilobites from the Higuchi-zawa
locality of the fauna. A new lichid genus Nipponarges gen. nov. and its type-species
Nipponarges mediosulcatus, gen. et sp. nov. and a new lichid species Acanthopyge(Acanthopyge) duplicispinata, sp. nov. are proposed in this part.
Thysanopeltella paucispinosa (Okubo, 1951) and Dechenella (Dechenella)minima (Okubo, 1951), which had been previously considered to be reliable fossil
evidences of the Eifelian in the Ohmori-zawa and of the Givetian in the Higuchizawa :respectively, co-exist at the Higuchi-zawa locality. In the present state ofknowledge it is only Thysanopeltella Kobayashi 1957 that can be used as an age
indicator of this trilobite fauna. Therefore, the geological age of the fauna may be
Eifelian.

Introduction

the basis of a fragmental cephalon and a left
eye from the southwestern slope of Takainariyama, Hikoroichi-cho, Ofunato City, and then
Sugiyama (1944) gave it a new specific name
Phacops (s. so) okanoi. Their locality has beencalled the "Phacops Zone", but its stratigraphic
horizon is uncertain. The author considers thatthe locality may be situated near the Kuronbora
at the middle part of the Ohmori-zawa, Hikoroichi-cho, Ofunato City, where Bronteus (Thysanopeltis) paucispinosa was newly documented
on the basis of a pygidium by Okubo (1951).
According to Okubo (1950), the paucispinosabearing horizon belongs to his 49 bed in theupper part of the Middle Devonian III Group of
his Takainari System, which corresponds to

Until recently, only a few Devonian species oftrilobites had been reported from the Kitakami
Mountains in northeastern part of Honshu,
Japan.
In the Devonian System in the Kitakami
Mountains, most of trilobites have been reported
from the Middle Devonian formation named
the Nakazato Formation by Yabe and Sugiyama
in their preliminary note (1937).
In 1941, Sugiyama and Okano described thefirst discovery of Phacops (s. s.) sp_ indet. on* Received October 8,1983.
*J.T.A.F. (Studies on Japanese Trilobites andAssociated Fossils) contrib. no. 28.
474

788. A Middle Devonian Trilobite Fauna

the Nakazato Fonnation of Yabe and Sugiyama(1937). At the same time, Okubo (1951) also
proposed a new species Dechenella minima on
the basis of a cranidium from the middle part of
the Higuchi-zawa, Hikoroichi-cho, Ofuna to Cityand pointed out an occurrence of Phacops (s. s.)differing from Phacops okanoi. In 1956, Okubo
described a new species Phacops nonakai on thebasis of a cephalo-thorax and two cranidia
from the Higuchi-zawa locality.
The above-mentioned four species of trilobites
have been reported from the Devonian Systemin the Kitakami Mountains. The stratigraphic
horizon at which these Devonian trilobites occurred can be assigned to the Middle Devonian
Nakazato Formation.
In 1957, Kobayashi redescribed Okubo's(1956) specimens. Namely Kobayashi assigned
Bronteus (Thysanopeitis) paucispinosa to his newsubgenus Thysanopeitella within the genus
Scutellum (Le. a senior synonym of Bronteus)
and Dechenella minima to the subgenus Dechenella (Dechenella) Kayser, 1880. Kobayashi regarded the former as Couvinian in age or thereabout and the latter as Givetian in age becausein Europe Thysanopeitella occurs only in the
Eifelian Stage and the subgenus Dechenella(Dechenella) in the Givetian Stage.
In 1974, Kobayashi and Hamada assigned
Scutellum(Thysanopeitella)pauclspmosa(Okubo) to the subgenus Thysanopeitella (Septimopeltis) Pfibyl & Vanl:!k 1971.
In 1977, Kobayashi and Hamada have regarded the geological age of Thysanopeitella(Septimopeltis) paucispinosa (Okubo) as Eifelianbecause the subgenus occurs only in the Eifelianin Bohemia, and at the same time they transferred Phacops nonakai Okubo to the genus
Reedops R. & E. Richter, 1925.
Recently, judging from the faunal assemblageof corals and brachiopods, Minato et al. (1979)
also pointed out that the Calceola-bearing
horizon at the Ohmori-zawa locality, from which
they reported an occurrence of fragmental
trilobites such as lichid, scutellid, dalmanitid
and phacopid, can be regarded as Late Couvinianand the fossiliferous horizon at the Higuchi-zawa475

locality as Givetian. According to their opinion,"The forms from Loc. A (Le. the Ohmori-zawa
locality) may be Couvinian, since they include
Calceola, "Favosites" of polymorph us groupand Heliolites ex. gr. porosus, all with "Eifelian"affinities. However, it also carries Macgeea, which
is nonnally Givetian-Frasnian in age. Also commonly Givetian, Chascothyris is found amongst
the brachiopods from Loc. A.... (several passages
omitted) ... Fossils from Loc. B in Higuchi-zawa
contain the Givetian Dechenella (M. Okubo,1951) and Undispirifer (Nakazatothyris) whichclosely resembles an "Eifelian" to Givetian form
from the USSR (M. Minato and M. Kato, 1978).
The fossil horizon at Loc. B may thus be Givetian." Minato et al. (1979) introduced Haga's
stratigraphic subdivision of the Nakazato Formation into four members, the third named N3
which contains the fossiliferous horizons at the
Ohmori-zawa and Higuchi-zawa.
In the early summer of 1980, the author
discovered a trilobite-bearing fossiliferous horizon at a small tributary of the Higuchi-zawa,
Hikoroichi-cho, Ofunato City, Iwate Prefecture.
The horizon is composed of fine-grained sandstone with tuffaceous intercalations and canbe assigned to the Middle Devonian Nakazato
Fonnation and Haga's N3 member. The locality
may be the same as that from which Okubo
reported the occurrence of Dechenella minima
in 1951 and Phacops nonakai in 1956. The
author's painstaking fossil hunting at that locality resulted in a discovery of various kinds oftrilobites. They belong to the Lichidae, Scutelluidae, Dechenellidae, Calymenidae, Phacopidae,
Dalmanitidae, Odontopl~uridae and so on. In
addition to this trilobite fauna, various kinds ofbrachiopods including atrypoids, gastropods,
tabulate and rugose corals including Calceolasp., orthocone cephalopods and plants are alsoabundantly represented. Dechenella (Dechenella)minima (Okubo) and Thysanopeitella paucispinosa (Okubo), which consists not only of thepygidium but also of the cranidium and librigena,
co-exist at the above-mentioned locality. Therefore, both horizons at the Higuchi-zawa locality
and Ohmori-zawa locality can be considered to476

Atsushi KANEKO

be synchronous.
The subgenus Dechenella (Dechenella) can be
no longer used as an indicator of Givetian age
because Onniston (1967) has described several
species of the subgenus from the Eifelian Blue
Fiord Fonnation in the Canadian Arctic. Furthermore, according to Ochs and Wolfart (1961),
the subgenus Dechenella (DecheneUa) has been
discovered even from the Upper Eifelian of the
Eifel district of Gennany. The other membersof the present Devonian trilobite fauna except
Thysanopeltella are either endemic or longranging, so it is difficult to use them as indicatorsof geological age. On the other hand, since thegenus Thysanopeltella has been never known
from strata younger than the Upper Eifelian in
the European succession, it is safe to say that thegeological age of the present Devonian trilobite
fauna may be no younger than late Eifelian.
Very recently, Copper, Tazawa, Mori andKato (1982) have come to a similar conclusionon the basis of their study of the atrypoid
brachiopods from Haga's N3 member of the
Nakazato Fonnation. They have pointed outthat the Atrypa-bearing bed of the Higuchi-zawa(Le. the minima horizon sensu Kobayashi and
Hamada (1977, p. 11» is correlatable to thatof the Ohmori-zawa (i.e. the paucispinosa horizon) since Atrypa pauciplicata Okubo, 1956
from the Higuchi-zawa could be regarded asa junior synonym of Atrypa desquamata mut.
japonica Sugiyama, 1942 from the Ohmori-zawa.
Those authors assigned them to the subgenus
Atrypa (Planatrypa) and suggested that it isquite possible for the Japanese species to bea junior synonym of Atrypa (Planatrypa)squamifera from the Upper Eifelian of the Eifel
district of Gennany and that at least the N3
member of the Nakazato Formation is middle
to late Eifelian in age because of the Atrypa(Planatrypa) never ranging into Givetian.
The Kitakami Devonian trilobite fauna showsa considerable degree of endemism at the genericlevel. As for the Japanese Devonian trilobite
faunas, there are no correlative elements at thegeneric level with the exception of the genus
Acanthopyge (s. 1.), which is almost cosmo-

politan, between the Kitakami Devonian trilobite
fauna and the Hida Devonian trilobite faunadescribed by Kobayashi and Hamada (1977)
from the Fukuji Fonnation and its equivalentsin the Hida Massif in central Honshu. The age of
the Fukuji Fonnation was considered to be
Gedinnian to Eifelian or probably lower Eifelian
by Kobayashi and Hamada (1977) or to be
Siegenian to Emsian by Copper et al. (1982).
At the familial level, the Kitakami Devonian
trilobite fauna is most remarkably different fromthe Hida Devonian trilobite fauna in being
dominated by Phacopidae and Dechenellidae,
whose representatives are much more numerous
than all other faunal members of the trilobites.
The Hida Devonian trilobite fauna has no elements of the Phacopidae and Dechenellidae.
Furthennore, the fonner trilobite fauna commonly also contains Dalmanitidae, but lacks
Cheiruridae. By contrast, the latter trilobite faunalacks Dalmanitidae, but contains Cheiruridae
frequently. Calymenidae are common to thetwo faunas, but the two calymenids belong todifferent subfamilies proposed by Siveter (1977).
The calymenid from the Kitakami Mountains
shows evidently the papillate-buttress structureon the cranidium typical of the subfamily
Calymeninae (s. s.), while that from the Hida
Massif shows the absence of that structure
typical of the subfamily Flexicalymeninae.
Kobayashi and Hamada (1977) described
flexicalymeninids from the Hida Devonian
trilobite fauna as Gravicalymene yamakoshii
and pointed out that their new species is very
closely allied to Gravicalymene maloungkaensis(Mansuy), which is widely distributed in South
China, north Viet-Nam and possibly East Bunna(p. 131). The above-mentioned two species areclosely allied to Gravicalymene zhenzishanesis
Nan, 1980 from the Givetian (Di) Heitai Fonnation at Heitai near Mishan near the northwestern
shore of Lake Hsingkai (= Khanka), Northeast
China. These Devonian species of Gravicalymene
evidently show that these faunas can be correlated to one another. Chatterton and Campbell(1980) proposed to apply their new generic name
Apocalymene to the post-Ordovician species of788. A Middle Devonian Trilobite Fauna
Gravicalymene from Australia, Morocco andTurkey. Chatterton et al. (1979) suggested theEast to Southeast Asian species of the Devonian
Gravicizlymene to be assignable to Apocalymene.
The Kitakami Middle Devonian trilobite fauna
can not be considered to be descended from the
Hida Lower Devonian trilobite fauna , which, on
the other hand , may possibly be one of theancestors of the Heitai Givetian fauna.

Systemlltic Palaeontology
Family Lichidae Hawle & Corda, 1847
Subfamily Ceratarginae Tripp, 1957
Genus Nipponarges Kaneko, gen . nov.
Derivation of name: Nippon = Japan + aninvalid trilobite name Arges Goldfuss, 1839,
which was replaced by Ceratarges GUrich, 1901
as a junior homonym of Arges DeHaan, 1835 in
the Crustacea from Japan.
Type-sp ecies:
N ipponarges m ediosulcatus
Kaneko, gen. et sp. nov. from the Middle Devonian (probably Eifelian) Nakazato Formation in
the southern part of the Kitakami Mountains,
northeastern Japan.
Species assigned: Presently only the typespecies is known.
Generic diagnosis : A large ceratarginid with
2p and 3p transglabelJar furrows clearly definedon median glabelJar lobe. Cranidium vaulted,
with a median glabelJar lobe which consists offrontal and 2p and 3p glabellar lobes, a pair ofbullar lobes and a pair of posterolateral cranidial
lobes , which are all circumscribed. Frontal
glabelJar lobe weakly inflated , much wider (trans.)
than long, expanding in front of bulJar lobe.
BulJar lobe subreniform in outline , weakly
inflated , much wider (trans.) than central area of
median glabellar lobe. Posterolateral cranidial
lobe short (exsag.) , highly inflated with a very
steep slope , slightly overhanging surrounding
furrows anteriorly and posteriorly , bearing
closely spaced , thick, obtusely spinose tubercles.
Preoccipital depressed area clearly defined , very
profound , transversely elongate, with a pair of

4 77

tubercles. Palpebral lobe relatively small, mesiallyfurrowed. Anterior border almost flat , short (sag.)medially , but well-developed anterolaterally.
Anterior pit in front of anterolateral corner ofbullar lobe. Occipital ring relatively narrow (sag.
and exsag.) moderately arched transversely,
widest (sag.) medially , narrowing (exsag.) laterally , with very steep anterior slope , bearing
two rows of tubercles ; one on anterior slope ,the other on the crest. Librigena unknown.
Hypostome unknown .
Thorax unknown .
Pygidium strongly spinose on margin , withno marginal border. Axis strongly inflated,bluntly rounded off in posterior, gently tapering
anteriorly along elongate (exsag.) adaxial marginof second pleural segment to in front of secondring furrow, with extremely thickened, short,obtuse spines posterolaterally. First axial ringstrongly convex, with vertical posterior slope .
Second axial ring moderately inflated laterally.

T ext· fig. 1. Diagram of morphological terminology o f cerat arginid cranidium. (1) anteriorborder, ( 2 ) anterio r border furrow , (3) anterior
pi t, (4) ax ial furrow , ( 5 ) lo ngi t udin al furrow, (6)bull ar lobe, (7) pregla be ll ar furrow, (8 ) m e dianglabellar lobe «9)+(11)+(13)) , (9) frontal glabellar lobe, (10) 3p t ransgla bella r furrow , ( 11) 3p
glabellar lobe, (12) 2p · transglab ellar furrow ,( 13) 2p gla bell ar lobe , (1 4 ) Ip gla bellar furrow,(15) p reoccipital depressed area, (16) posterolat eral cranidial lobe, (1 7 ) palpebral furrow , (18)palp ebral lobe , (19) posterior border furrow,(20) posterior border , (21) occipital furrow, (22)occipital rin g478

Atsushi KANEKO

Pleural region elongate (exsag.), at least withthree pairs of pleural segments and cylindricalmarginal spines. First two pleural segments
subdivided by pleural furrows into two bands,
the posterior ones of which are massive andcontinue into spines laterally. Pygidial surface
coarsely tuberculose.
Discussion: On the basis of the observationof the ontogenic development of "bicomposite
lobes" sensu Reed (1902, p. 66) of three taxonomically diverse lichid species, Lichas laciniatus(Wahlen berg, 1818), Acanthopyge (Mephiarges)
bifida Edgell, 1955 and Amphilichas minganensis(Billings, 1865) shown by Temple (1969), Chatterton (1971) and Shaw (1968) respectively,
Temple (1972, pp. 373-374) pointed out thatthey are not bicomposite in orgin, i.e. they are
not formed by the fusion of 2p and 3p lateralglabellar lobes and by backward prolongationof the inner ends of the 3p glabellar furrows,
and so proposed the term bulla for the rudimentary swelling in axial furrow on its adaxial
side opposite the 2p glabellar lobe in the earlylichid grawth stage and the term bullar lobe,
instead of "bicomposite lobe", for the forwardly
grown-up lobe along the side of the glabella.
According to his interpretation of lichid morphology, the 2p and 3p glabellar furrows arepresent only on the median glabellar lobe.
Temple (1969, pp. 213-214, Text-fig, 2a, pI. 2,
fig. 11) showed that in the earliest known(presumably meraspid) growth stage of Lichas
laciniatus the median glabellar lobe bears the 2ptransglabellar and 3p lateral glabellar furrows.
And then Chatterton (1971, pp. 34-35, fig. 9B)
showed that in the early metaprotaspid stage ofAcanthopyge (Mephiarges) bifida the glabella
without the bulla, which corresponds with themedian glabellar lobe in the later growth stage,
bears the 2p and 3p transglabellar furrows.
Recently Tripp and Evitt (1981, pp. 667, fig. 1a)also showed that in the early metaprotaspid
stage of Hemiarges tumeri rasetti Tripp & Evitt,
1981 two (2p and 3p) lateral glabellar furrows
are occasionally indicated. These furrows areusually lost during the ontogeny in the family
Lichidae. Therefore, Nipponarges gen. nov. canbe easily distinguished from all other membersof the family Lichidae by the presence of theclearly defined 2p ap.d 3p transglabellar furrows
on the median glabellar lobe. In the basic structural pattern of lobation of the glabella and thepresence of the clearly defined preoccipitaldepressed area with a pair of tubercles at thebase of the median glabellar lobe, Nipponarges
is comparable with some genera in the subfamily
Ceratarginae Tripp, 1957.
The basic morphology of the genus Nipponarges gen. nov. is most similar to that of the
Middle Silurian to Middle Devonian genus
Acanthopyge Hawle & Corda, 1847, which isgeographically wide-spread. However, there areseveral distinctive features of generic importancein addition to 2p and 3p transglabellar furrows
on the median glabellar lobe, which are as
follows: 1) a much less inflated frontal glabellar
lobe; 2) extremely wider (trans.) bullar lobe
than the central area of the median glabellar
lobe; 3) much more highly inflated, exsagittallyshortened posterolateral cranidial lobe withclosely spaced, prominent, obtusely spinose
tubercles; 4) relatively well-developed anteriorborder; 5) more anteriorly situated anterior
pit; 6) relatively small palpebral lobe which isstrongly furrowed mesially; 7) more strongly
inflated, massive posterior bands of the firsttwo pleural segments whose distal extensions
form conspicuous, cylindrical spines; 8) much
more elongate (exsag.) adaxial margin of second
pleural segment; 9) conspicuous, short, obtuse
spines on the pygidial axis.
In so far as the massiveness of the pleuralsegments including the cylindrical marginal
spines of the pygidium and the large size for a
ceratarginid are concerned, Nipponarges gen.
nov. is similar to the Lower to Middle Devonian
genus Terranovia Maximova, 1977 from Novaya
Zemlya and Salair of U.S.S.R. and the Canadian
Arctic rather than Acanthopyge. However, inNipponarges the first two pleural segments are
inclined more posteriorly and the second pleuralsegment, of which adaxial margin is extremely
elongate (exsag.), widens more rapidly laterally
and is subdivided into two bands by the pleural788. A Middle Devonian Trilobite Fauna

furrow except its anteroadaxial portion. Furthermore, in Terranovia, according to Ormiston(1982), the cranidium bears two conspicuous
pairs of upwardly and backwardly directed
spines, instead of the transglabellar furrows,
on the median glabellar lobe and the pygidium
bears two pairs of upwardly directed pleuralspines, and there is a transverse union of posterolateral cranidial lobes, instead of the preoccipitaldepressed area, a character typical of Acanthopyge, Nipponarges and some of other ceratarginids, at the base of the median glabellar lobe
as in the case of Ceratarges Gtlrich, 1901 fromthe Lower to Middle Devonian of Germany andFrance; the median and bullar lobes are a little
more inflated, but the posterolateral cranidial
lobe is much less inflated than those of Nipponarges.
Radiolichas Reed, 1923 from the Middle
Devonian of Germany is more similar to Nipponarges gen. nov. than Terranovia in the weak
inflation of the frontal glabellar lobe, in theextreme development of the bullar lobe, in theclearly defined, transversely elongate preoccipitaldepressed area with a pair of tubercles and in
the inclination of the first two pleural segmentsof the pygidium. However, Radiolichas has notransglabellar furrows on the median glabellarlobe, but has much less inflated and more laterally situated posterolateral cranidial lobe andthe oar-shaped pygidial marginal spine whichis mesially furrowed.
At first glance, the pygidium of the Nipponarges gen. nov. looks similar to that of themonotypic genus Terataspis Hall, 1863 in thesubfamily Lichinae on the basis of Lichas grandis
Hall, 1861 from the Emsian Schoharie Formation and its equivalent in New York State, U.S.A.
in the strong spinosity on margin and in the large
size (Hall and Clarke (1888), pp. 73-77, pis.
17-19). However, Nipponarges has exsagittallymore elongate pleural region as in the case of
Acanthopyge and Radiolichas in the subfamily
Ceratarginae than Tera tasp is, and has the conspicuous, short, obtuse spines on the axis,
whose analogues can be seen in Acanthopyge, forexample, such as Acanthopyge consanguinea(Hall, 1894) illustrated by Whittington (1956,479

p. 1203, pI. 131, figs. 14-17, 19), Acanthopyge(Mephiarges) bifida Edgell, 1955 illustrated by
Chatterton (1971, p. 34, pI. 6, figs. 19-20, 22)
and Acanthopyge (Acanthopyge) duplicispinata
sp. nov. (p. 481, PI. 89, Figs. 3a-c) and in Terranovia illustrated by Ormiston (1982, p. 1264,
pI. 2, figs. 1-4, 10). Ormiston (1982) assigned
the pygidium which Corgan (1963) reported
as Terataspis sp. indet. from the Lower Devonian(Onesquethawan) in the Canadian Arctic to thegenus Terranovia and regarded it as conspecificwith its type-species Terranovia nalivkini Maximova, 1977, and points out that Terataspis
can be regarded as a genus restricted to theEmsian rocks of eastern North America (personalcommunication from Dr. A. R. Ormiston,
6 October, 1981).
The author comes to conclusion that the
Nipponarges morphology was derived from an
Acanthopyge morphology, probably from the
Asian species morphology characterized by aslender median glabellar lobe and a well-developed bullar lobe as discussed below.
Distribution: Confined to the Nakazato
Formation in northeastern Honshu, Japan.

Nipponarges mediosulcatus Kaneko,
gen. et sp. nov.
PI. 87, Figs. 1-3, PI. 88, Figs. 1-4, Text-fig. 2.

Derivation of name: From the Latin words
medius, median + sulcus, furrow.
Holotype: Incomplete cranidium, KACoT0101 [PAl7144] (PI. 87, Figs. 1a-c, Pl. 88,
Figs. la-e), coli. A. Kaneko, 1980.
Specific diagnosis: As for the genus.
Description: Two pieces of the holotype
cranidium are orientated in due considerationof both a plane of cleavage of matrix and ceratarginid morphology: Cranidium moderately
vaulted; anterior outline straight in front ofanterior pits, and then directed posterolaterally
from each of anterior pits at an angle of about
60 degrees from sagittal line. Median glabellar
lobe with 2p and 3p transglabellar furrows;
frontal glabellar lobe slightly inflated, roughly
pentagonal in outline, much wider (trans.) than

480

Atsushi KANEKO

long, expanding in front of bullar lobe, isolated
by 3p transglabellar furrow from semi-cylindricalcentral area of median glabellar lobe; 3p glabellar
lobe rectangular in outline, about three times aslong (sag. and exsag.) as wide; 2p glabellar lobe
slightly divergent posteriorly and sloping downnearly vertically at its base. Bullar lobe subreniform in outline, slightly inflated, extremely
large, much wider (trans.) than central area of
median glabellar lobe, sloping down nearlyvertically at its posterior end. Posterolateralcranidial lobe roughly elongate (trans.) oval in
outline, shortening (exsag.) adaxially, much
shorter (exsag.) than bullar lobe, expanding
strongly dorsolaterally so that its raised portion
slightly overhangs surrounding furrows anteriorly
and posteriorly. Palpebral lobe short (exsag.),elongate (trans.) oval in outline, directed outward and slightly forward and downward, deeply
furrowed mesially, highly elevated with analmost vertical slope except adaxial side whichslopes down gently for a short distance tomost raised portion of narrow (trans.) palpebral
furrow which runs on nearly vertical inflationof fixigenal carapace to form a butress between palpebral lobe and anterolateral comerof posterolateral cranidial lobe. Ip glabellar
furrow relatively wide (sag. and exsag.); outside
longitudinal furrow Ip glabellar furrow diverging
anteriorly at an angle of about 60 degrees fromsagittal line, deepest and clearly U-shaped at its
most raised portion in front of anteroadaxial
comer of posterolateral cranidial lobe, fromwhich Ip glabellar furrow slopes down steeply
anterolaterally to merge axial furrow and slopes
down slightly less steeply posteromedially tointersect longitudinal furrow at funnel-shaped,
profound concavity behind posteroadaxial comerof bullar lobe; inside longitudinal furrow Ip
glabellar furrow shallowing rapidly medially,running in a broad arc along distal end of medianglabellar lobe. Longitudinal furrow narrow(trans.), sharply incised in front of Ip glabellar
furrow, but represented by change of slope without incised furrow behind Ip glabellar furrow;from concavity to level of 2p transglabellar
furrow on median glabellar lobe longitudinalfurrow moderately deep, converging slightly,
rising nearly vertically, and then becoming
deeper, sloping down gently; beside 3p glabellar
lobe longitUdinal furrow deep, deflected slightly
outward; from level of 3p transglabellar furrow
to anterior pit longitudinal furrow becoming
progressively shallower, curving more or lesssharply outward and downward along anterior
margin of bullar lobe; behind Ip glabellar furrow
longitudinal furrow rising steeply on posterior
slope of concavity for a short distance to adaxial
extremity of posterolateral cranidiallobe, thereafter merging with occipital furrow. Transglabellar furrows on median glabellar lobe moderately
incised, more or less wide (sag. and exsag.),
curving downward and forward distally; 2ptransglabellar furrow running into abaxial extremity of slight deflection of longitudinalfurrow. Preoccipital depressed area clearly defined by Ip glabellar furrow behind medianglabellar lobe, elongate transversely, profound,surrounded by steep walls, shortest (sag.) medially, with a pair of distinct, large tubercles onposteroadaxial slopes of concavities; slightswelling continuing adaxially from large tubercle,
but narrowing (exsag.) rapidly and dying out
medially. Anterior border separated only by
preglabellar furrow from frontal glabellar lobe,
almost flat, but very slightly inflated transversely
inside anterior pit, shortest (sag.) medially,
becoming rapidly longer (exsag.) and attaining
maximum length in front of, or slightly inside,
anterior pit and then shortening as far as a (+ (3).
Preglabellar furrow weakly incised, curving
gently along anterior margin of frontal glabellar
lobe. Anterior pit situated in front of anterolateral comer of bullar lobe, profound, relatively
large, joining anterior border furrow and preglabellar furrow and .axial furrow and longitudinal furrow; on ventral surface anterior pit
represented by prominent boss. Anterior borderfurrow on fixigena wide (exsag.), shallow, risingslightly posterolaterally at an angle of about 80
degrees from sagittal line. Anterior portion of
fixigena very broad (trans.), convex behindanterior border; known part of middle portion
risin'I with a very steep slope and progressively

788. A Middle Devonian Trilobite Fauna
inflated posteriorly between anterior extensionof ·palpebral furrow and axial furrow, running
into antero-abaxial corner of posterolateralcranidiallobe.
Occipital ring wide (trans.), slender, moderately arched transversely, longest (sag.) medially,
narrowing (exsag.) laterally, with very steep
anterior slope and almost straight posterior
margin. Occipital furrow merging with 1p glabellar furrow medially; behind a pair of tubercles
and its adaxial extensions occipital furrow wide
(exsag.), shallow; behind posterolateral cranidial
lobe occipital furrow deep,. U-shaped, progressively narrowing laterally. Librigena unknown.
Hypostome unknown.
Thorax unknown.
Pygidium strongly spinose on margin, withno marginal border. Axis strongly inflated,bearing extremely prominent, dorsolaterally
directed, short, obtuse spines posterolaterally,
whose subhorizontai section reveals ellipticaloutline; articulating half-ring short (sag. andexsag.), separated from first axial ring by changeof slope without incised articulating furrow;first axial ring strongly convex, sloping down
vertically posteriorly; second axial ring moderately inflated laterally. Axial furrow most deeply
excavated in front of first ring furrow, moderately convergent from antedor margin to in frontof second ring furrow, thereafter curving out
gently. Pleural region except border spines
narrow (trans.), considerably elongate (exsag.);first two pleural segments directed posterolaterally, widening (exsag.) rapidly laterally; firstpleural segment subdivided by narrow (exsag.),shallow pleural furrow into short (exsag.),slightly convex anterior band and much more
highly inflated, longer (exsag.), massive posterior
band whose anterior and posterior slope is almost
vertical adaxially and whose distal extension
forms massive, cylindrical, long (trans.), gently
upturned marginal spine; second pleural segment
except marginal spine with posterolaterally
elongate sub-rhomboid outline resulting fromextremely elongate (exsag.) adaxial margin along
axial furrow, subdivided by narrow (exsag.),shallow pleural furrow, which becomes shallower

481

and curves outward distally, into anterior andposterior bands except its anteroadaxial portion
which is shorter (exsag.) than posterior band of
first pleural segment; anterior band of second
pleural segment about as long (exsag.) as posteiorband except margin, a little less inflated adaxiallybut sloping gently away laterally; third pleuralsegment longitudinally elongate sub-rhomboid in
outline, less inflated than the preceedings, withmassive spine on posteromost margin. First
interpleural furrow deep, narrow (exsag.), Vshaped, almost parallel with second interpleuralfurrow, which is less deep and narrow; thirdinterpleural furrow less deep and narrow (exsag.)than the second one, nearly longitudinal.
Surface ornamentation: posterolateral cranidial lobe and at least posteromost portions ofmedian and bullar lobes covered with dense,
thick, obtusely spinose tubet:cles; judging fromthe plugged tubercle bases on the externalmould of the holotype, rest of median and bullar
lobes bear the same type as above-mentioned orless prominent, thick tubercles; medium.£ized
tubercles scattered on anterior border andfixigena and posterior border; occipital ring
bearing two rows of medium.£ized tubercles, oneof which is somewhat spinose and present on the
crest, the other of which is present on anterior
slope. Small to large, more or less pointed
tubercles scattered on pleural region including
marginal spine of pygidium; much larger ones on
third pleural segment; smaller ones on marginal
spine.
Genus Acanthopyge Hawle & Corda, 1847
Subgenus Acanthopyge Hawle & Corda, 1847
The subgenera of Acanthopyge have been
discussed recently by Chatterton et al. (1979).

Acanthopyge (Acanthopyge) duplicispinata
Kaneko, sp. nov.
PI. 88, Figs. 5-7, PI. 89, Figs. 1-3, Text-fig. 3.

Derivation of name: From the Latin words
duplex, paired + spina, spine.

482

Atsushi KANEKO

Text-fig. 2. Restoration of Nipponarges mediosulcatus Kaneko, gen. et sp.
nov . Two pieces of the cranidium are slightly differently orientated from Plate 87 ,Figs. la-b .
Holotype : Incomplete cranidium , KACnT10106 [PAl7149] (PI. 89, Figs. la-e), colI. A.
Kaneko , 1980.
Specific diagnosis: An Acanthopyge species
with a slender median glabellar lobe flanked
by well-developed bullar lobe and with paired

and unpaired, spinose tubercles and spines on
cranidium, some of which represent spines 1, 2,
3?, 4, 5, A1 sensu Chatterton (1971). Palpebral
lobe highly elevated, with rudimentary PI sensu
Chatterton. Ventral surface of hypostome apparently smooth. Pygidium strongly spinose on

788. A Middle Devonian Trilobite Fauna

margin, with no marginal border, and withelongate (exsag.) pleural region, and with welldeveloped axis.
Description: Cranidium vaulted, subtrapezoidal in outline, with a median glabellar lobe, apair of bullar lobes and a pair of posterolateralcranidial lobes, which are all circumscribed.
Glabella strongly convex; median glabellar lobe
about one-third as wide as (trans.) occipital ringat its base, elongate (sag.), highly arched transversely, rising above and expanding in front ofbullar lobe, geniculate anteriorly; bullar lobe
roundly subtriangular in outline, moderately
convex. Posterolateral cranidiallobe roundly subrhomboid in outline, about as long (exsag.) asbullar lobe, strongly inflated, with very steep
posterior slope; its anterolateral portion slightly
ridging on internal mould, descending very
gently from prominent spine. pair to palpebral
furrow, from bottom of which short (exsag.)palpebral lobe rises vertically above the crest of
posterolateral cranidiallobe excluding prominent
spine pairs (AI). Eye ridge unknown. Longitudinal furrow in front of Ip glabellar furrow slightly
converging anteriorly toward the highest pointof glabella at about one-third the length (sag.)of median glabellar lobe from its base, and thenslightly deflected outward, and then becomingslightly shallower and curving sharply outward
and downward to merge preglabellar furrow andaxial furrow; longitudinal furrow behind Ip
glabellar furrow gently diverging posteriorly tomerge with occipital furrow. Ip glabellar furrow
directed posteriorly diagonally from axial furrow
to in front of anterolateral corner of posterolateral cranidial lobe, intersecting longitudinalfurrow at profound concavity behind posteroadaxial corner of bullar lobe, merging with occipital furrow on sagittal line; its adaxial portion
outside longitudinal furrow deeply incised, sothat posterolateral cranidial lobe is completely
separated from bullar lobe. Preglabellar and axial
furrows lightly incised. Preoccipital depressedarea behind median glabellar lobe profound,clearly defined by lightly incised adaxial portionof Ip glabellar furrow, by posterior portion of
longitudinal furrow and by occipital furrow,

483

divided into a symmetrical pair of slightly swelling, subtriangular areas by junction of Ip
glabellar and occipital furrows on sagittal line.
Occipital ring wide (trans.), moderately archedtransversely, longest (sag.) medially, shortening(exsag.) abaxially; its posterior margin nearlystraight. Librigena unknown. Hypostome subtrapezoidal in outline, about three-quarter aslong as wide (trans.); median body gently convex, slightly less than a half of hypostomallength(sag.), a half as long as wide (trans.); medianfurrow short (exsag.), deep, subtransverse, only
incised abaxially in posterior one-third of median
body; abaxial portion of posterior lobe of
median body slightly inflated to form macula;lateral border furrow deep, wide (trans.), directed posteriorly diagonally to merge deep,short (sag. and exsag.), transverse posteiorfurrow; anterior wing short (exsag.), directed
upward and outward opposite median furrowof median body; posterior border long (sag.
and exsag.), wide (trans.), most convex anteromedially; doublure extending beneath lateral and posterior borders; axial one-third of
doublure excavated to form an anteriorly
deepening pit, whose posterior slope has a pair ofelongate (exsag.), narrow (trans.), low ridges.
Thorax unknown.
Pygidium excluding marginal spines semielliptical in outline. First two pleural segments
clearly defined except very slightly inflated
medial part of second axial ring, transverse
medially, flexed posterolaterally near abaxial
end of axial ring; flexed part of second axial
ring inclined more posteriorly than that of firstaxial ring. Axis wide (trans.), semi-cylindrical,
moderately arched transversely, most convex
posteromedially, rounded off abruptly but
continued by low, convergent post-axial ridge
posteriorly; articulating half-ring short (sag. andexsag.), moderately convex, separated from restof axis by short (sag. and exsag.), transverse,
moderately incised articulating furrow; first axial
ring strongly convex, sloping down verticallyposteriorly, with narrow (trans.), slightly backwardly flexed adaxial part, only behind which
second axial ring moderately inflated; ring

484

Atsushi KANEKO

furrows very faint behind medial part of firsttwo axial rings , but becoming distinct behindremainder part of them. Axial furrow distinct,
slightly converging posteriorly from anterior
margin to in front of second ring furrow, there ·after slightly diverging to posterolateral cornerof axis, and then converging in a gentle curve,
then flexed backward and converging slightly
beside post-axial ridge. Pleural region elongate(exsag.), horizontal; first two pleural segments
represented by long (exsag.), strontly convex,
posteior pleural band which continues into
prominent marginal spine distally; anterior
pleural band · of first pleural segment short(exsag.), almost flat; anterior pleural band of
second pleural segment slightly longer (exsag.)than posterior pleural band, represented byslight swelling between very faint pleural furrow
and more distinct interpleural furrow; third
pleural segment elongate (exsag.), with promi nent, upturned marginal spine at posterior end
which very slightly diverges posteriorly and with
five, small, irregularly sized spines on postero lateral margin.
Surface ornamentation: Most prominent spine
pair (A1) upwardly directed on posterolateralcranidial lobe behind abaxial extremity of bullar
lobe; median glabellar lobe bearing two prominent spine pairs (4 , 5); one (4) on the highest
point of glabella more prominent than the otherat level of anterior extremity of bullar lobe, andsmall to medium-sized median spinose tubercles
in a row on sagittal line; small- to medium-sized
spinose tubercles, some of which may havea tendency to be subsymmetrically arranged,
scattered on bullar and posterolateral cranidial
and rest of median glabellar lobes; small spinose
tubercle pair present on slight swelling of pre occipital depressed area; palpebral lobe bearingslight tuberculate swelling (reduced PI); a
medium-sized median spine flanked by a pair ofmore prominent spines on occipital ring; ventral
surface of hypostome apparently smooth, but
one specimen (PI. 88, Figs. 5a-d) showing a few
pits on median body; widely to moderately
spaced, medium-sized, spinose tUbercles, which
may be subsymmetrically arranged, scatteredText-fig. 3. Restoration of Acanthopyge(Acanthopyge) duplicispinata Kaneko, sp. nov.
on pygidial axis, post-axial ridge and pleuralregion; most prominent ones on most convex
posteromedial part of axis.
Discussion: Acanthopyge species with spinose
carapaces are occasionally included in the subgenus Acanthopyge·(Mephiarges) R. & E. Richter,
1930. R. & E. Richter (1917) described a spiny
lichid cephalon as Lichas (Euarges) mephisto
from the Middle Devonian Crinoiden-Zone at
Auburg near Gerolstein in the Eifel region inGermany. In 1930, they proposed for this forma new subgenus Mephiarges within the genus
Acanthopyge Hawle & Corda, 1847, which isa senior synonym of the genus Euarges GUrich,
1901, and gave the following diagnosis; "eineAcanthopyge mit stacheitragenden Augen und

788. A Middle Devonian Trilobite Fauna

gelappten Freiwangen (p. 39)". Later Edgell(1955) emended the subgeneric diagnosis when
he described Acanthopyge (Mephiarges) bifidafrom the limestone of the Murrumbidgee Groupat Wee Jasper, New South Wales, Australia, which
was considered to be lower Middle Devonian(Couvinian). He diagnosed A. (Mephiarges) as
follows; "An Acanthopyge with eyes on spiny
stalks, barbed genal spines and spines on theanterior glabellar lobe". Whittington (1956)
did not accept Edgell's aSsignment of the species
bifiaa to the subgenus Mephiarges and pointed
out many similarities to Arges consanguineus
Clarke, 1894, from the Lower Devon·ian New
Scotland limestone (the Helderberg group) in
Albany County in New York State, U.S.A.,
which he redescribed as an Acanthopyge species.
On the contrary, in 1971, Chatterton redescribed
Acanthopyge (Mephiarges) bifida Edgell from theupper Emsian to probably lower Eifelian Taemas
Formation of the Murrumbidgee· Group at the
Taemas-Cavan area near Yass, New South Wales.
He accepted Edgell 's em~ndation of the subgeneric diagnosis and emphasized more similarities between A. (Mephiarges) bifida Edgell andA. consanguinea (Clarke) than Whittington(1956) did and so suggested that the North
American species might belong to the subgenus
Mephiarges (pp. 40-41). Perry and Chatterton(1976) described a ceratarginid with a spinyc~anidium as Acanthopyge (Mephiarges) sp. fromthe Emsian Delorme Formation, Mackenzie
Mountains, northwestern Canada and considered
it to be very closely related to A. consanguinea.
Kobayashi and Hamada (1977), however, pointed
out that there is no link between A. (M.) bifida
and A. consanguinea (p. 52). In 1979, Chatterton,
Johnson and Campbell discussed as follows;
Acanthopyge (Mephiarges) bifida "has many
features in common with A. consanguinea(Clarke) and the Acanthopyge sp. of Perry andChatterton (1976), which do not show the distinguishing characteristics of A. (Mephiarges) ,though they do have an ornament of pits rather
than tubercles on the hypostome indicating thatthe other similarities may not really indicate a
relationship as close as has been previously

485

suggested (Chatterton 1971; Perry and Chatterton1976). It is nevertheless becoming increasingly
difficult to diagnose A. (Mephiarges) , whosetype species A. (M.) mephisto seems to be an
extreme form at the end of a range of variation
in a number of characters (pp. 819-820)".
Compared with A. (M.) bifida Edgell and A.
consanguinea (Clarke), the present new species
has several similarities. They are prominent spine
pairs (4, 5) on the median glabellar lobe, prominent spine pair (AI) on the posterolateralcranidial lobe, more or less well-developed occipital spines (though median one much lessprominent than that of A. (M.) bifida), a highly
elevated palpebral lobe to butress the stalk-likeeye socle, a relatively large axis of the pygidium,narrow anterior bands on the first two pleuralsegments and large median tubercles on theposterior portion of the axis of the pygidium.
However, in the present new species the palpebral
spine (PI), which is a diagnostic character of A.
(Mephiarges) and can be observed to some extent in A. consanguinea, A. (M.) bifida and A.
(Mephiarges) sp. by Perry and Chatterton (1976),
is quite rudimentary, the posterior bands of thefirst two pleural segments of the pygidium bearno prominent, dorsally directed spine pairs,
the median body of the hypostome is neither
tuberculate nor pitted, but almost smooth,though one specimen (PI. 88, Figs. 5a-d) showsa few "pits", which may be attributable topreservation. Furthermore the Japanese species
differs from the above-mentioned overseas
species in the degrees of development of themedian glabellar and bullar lobes. Namely, A.
(A.) duplicispinata sp. nov. shows a slender
median glabellar lobe flanked by well-developed
bullar lobes, whereas A. consanguinea and A.
(M.) bifida show a well-developed median glabellar lobe attended by relatively reduced bullar
lobes. It is a remarkable f!lct that the glabellar
morphology of a similar type to the present new
species can be observed in most Asian (Turkestan
including Kazakhstan, the Kuznetsk Basin)
species of Acanthopyge (s. I.). As opposed tothis, in most extra-Asian species with the exceptions such as Lichas (Arges) contusus Hall, 1863

486

Atsushi KANEKO

and Lichas devonianus Whidborne, 1889 whichwere compared to the genus Radiolichas Reed,
1923 by Reed (1923) and Pfibyl and Erben(1952), the glabellar morphology shows a similar
type to the above-mentioned, two overseas
species.
It should be noted that similar prominentspinose tubercle or spine pairs on the cranidium
can be observed also in some of lichid genera andsubgenera, for example, such as the Middle
Ordovician species Hemiarges turneri Chatterton& Ludvigsen, 1976 (pI. 19, figs. 3-16, 21, 25,26), the Wenlockian species Hemiarges mikulici
Perry & Chatterton, 1977 (pi. 5, figs. 23-25,
pI. 6, figs. 1-2) and the Lower Devonian species
Acanthopyge (Lobopyge) sinuata (Ratte, 1886)
ilhistrated by Chatterton et aZ. (1979, pi. 109,
figs. 1-7, 12, 14).
Therefore the author considers that thepresent new species is not assignable to thesubgenus Acanthopyge (Mephiarges) and is not
related to A. consanguinea and A. (M.) bifida,but is most closely allied to the Asian species
discussed below rather than to most extra-Asian
species.
The specimens illustrated as Lichas haueri
Barrande, 1846 by Weber (1932) from the Lower
Devonian in Turkestan is probably not conspecific with Lichas haueri Barrande, a seniorsynonym of type-species Acanthopyge Zeuchtenbergii Hawle & Corda, 1847 (pi. 1, figs. 5-7)
from the Eifelian, so-called Acanthopyge Limestone which is the shallow-water equivalent of
the Chotec Limestone of the Barrandian in
Bohemia (personal communication from Dr. Ivo

Chlupac, 7, March, 1983) because of the difference in the degree of development of the medianglabellar and bullar lobes as mentioned above.
The Turkestan specimens are closely comparable
to the present new species rather than with theBohemian species. Figure 37 of Weber's plate 1
from the locality no. 285 is too poor to observe
the surface ornamentation, but is similar to thepresent new species in glabellar morphology indorsal view. However, the Turkestan specimens
has a parallel-sided median glabellar lobe andsomewhat wider (trans.) bullar lobes. Figure 26
of Weber's plate 1 from the locality no. 268
show somewhat less elongate (sag. and exsag.)median glabellar lobe and wider (trans.) longitudinal furrow than those of the present new
species, but shows the signs of tubercle pairs on
the median glabellar lobe, which is slightly convergent posteriorly as in the present new species.
As an exception to the extra-Asian species,
Acanthopyge contusa (Hall, 1863) illustrated byHall and Clarke (1888, pi. 19B, figs. 3--6) ismore comparable with the present new species
and the Turkestan specimens than to Lichas(Euarges) sibiricus Tschernysheva, 1951 from theMiddle Devonian (D~, DD in the Kuznetsk Basin
which had been compared to it by Tschernysheva(1951). In the North American species thecentral area of the median glabellar lobe is parallel-sided (fig. 3) as in the case of figure 37 ofWeber's plate 1 or slightly divergent at its base(figs. 4, 5); the bullar lobe is more rounded thanthat of the Turkestan specimens; the cranidium
is ornamented with only scattered tubercles.
In Acanthopyge sibirica (Tschernysheva,

Explanation of Plate 87
Figs. 1-3. Nipponarges mediosulcatus Kaneko, gen. et sp. nov.
Figs. la-e. Holotype, KAC-DT-OlOl [PAl7144]. Rubber cast from external mould, which is acounterpart of internal mould shown in Figs. lb, c of Pi. 88 of incomplete cranidium; a, b,
dorsal stereo pair, x 1.5. c, posterior view, x 1.5. d, e, oblique lateral stereo pair, x 1.5.
Figs.2a-d. Paratype, KAC-DT-Ol02 [PAl7145]. Rubber cast from external mould, which is acounterpart of internal mould shown in Figs. 2a, b of PI. 88 of incomplete pygidium; a, b,
dorsal stereo pair, x 1.5. c, d, right lateral stereo pair, x 1.5.
Fig. 3. Rubber cast from external mould of another fragmental pygidium, KAC-DT-Ol03 [PA17146] , dorsal view, xL 5.

KANEKO: A Middle Devonian Trilobite Fauna

Plate 87

788. A Middle Devonian Trilobite Fauna
1951), the cranidium is gently inflated and thebullar lobe is extremely well-developed and the
median glabellar lobe seems extremely slender
as in the case of Radiolichas araneus (Holzappfel, 1895); the associated pygidium shows awell-developed axial lobe with a rounded outline
as in the case of the present new species, but thepleural region is poorly known.
Lichas (Euarges) planus Weber, 1932 from theMiddle Devonian at the locality no. 37 in the
district of Khodjent in Turkestan, which has aconsiderably large and weakly inflated carapace
for the genus Acanthopyge (s. I.), shows a glabellar morphology of similar type to the present
new species.
In Lobopyge limbata Maximova, 1968 fromthe Lower Devonian (D:, DD of Central Kazakhstan, the glabella has a more or less slender
median glabellar lobe flanked by well-developed
bullar lobes as in the case of the present new
species, but the anterior border is relatively
well-developed as in Craspedarges Gtlrich, 1901
and the associated pygidium is typical of Acanthopyge (Lobopyge) Pfibyl & Erben, 1952.
Acanthopyge (Lobopyge ?) sp. indet. illustrated by Kobayashi and Hamada (1977, pI. 4,
fig. 2) from the Lower Devonian of the Hida
Massif in central Honshu, Japan, thougha fragment of a cranidium, seems to show arelatively slender median glabellar lobe to beflanked by well-developed bullar lobes.
In Acanthopyge latelobe Nan, 1976 from theupper Middle Devonian (DD at Dong Ujimqin
Qi in Inner Mongolia and an' extremely convex
cranidium illustrated as Lichas (Euarges) parvulus
Novak var. convexa by Tschernysheva (1951)
from the upper Lower Devonian (DD in the
Kuznetsk Basin, the median glabellar lobe ismore or less wider (trans.) than those of theabove-mentioned Asian species, but the bullar
lobe is well-developed.
As an exception to the Asian species, Acanthopyge aff. contusa (Hall) illustrated by Maximova (1960) from the lower Middle Devonian(DD in Rudny Altai seems to be dissimilar to
A. contusa (Hall) from North America in having
considerably reduced bullar lobe as do most

487

extra-Asian species.
A glabellar morphology of the extra-Asiantype can be seen also in the South American
species Acanthopyge ballivianii (Kozlowski)
redescribed by Baldis (1967) from the Middle
Devonian Chavela Formation of the Province
San Juan, Argentina, but the pygidium of it has
four pairs of pleural segments terminating in
sharp spines.
Acknowledgmen ts
The author is greatly indebted to Dr. Allen
R. Ormiston of Amoco Production Company,
Tulsa, Oklahoma, for his useful paper (1982) inmanuscript, invaluable suggestions and kindness
in critical reading of the present manuscript; toProfessor Dr. Takashi Hamada of Universityof Tokyo, for his reading over the manuscript
and kind instruction and encourgement; toformer Curator Dr. Eiji Matsumoto of National
Science Museum, Tokyo, for his helpful criticism
on the manuscript and constant encourgement;
to Professor Dr. Harry B. Whittington of University of Cambridge, for his invaluable suggestions;and to Dr. Ivo Chlupac of Central Geological
Survey, Prague and to Dr. Jun-ichi Tazawa of
Tohoku University, for useful informations andreferences. He also expresses his sincere gratitude
to the following persons for so kindly giving him.many useful references in their charge: Dr. I. A.
Dubrova of Palaeontological Institute, Moscow;
Dr. M. J. Copeland of Geological Survey ofCanada; Dr. Radvan Horny of National Museum,
Prague; Dr. Derek J. Siveter of University of
Hull; Mr. Ko-ichi Nakamura of Geological Surveyof Japan; Mis Eleanor A. Gossen of New YorkState Museum and Science Service, Albany.
Special thanks are due to Mr. Hitoshi Koizumi
of Kesen Chishitsu Kenkyo.sho (K.K.), for his
financial help and many useful informations
and references.
Deep appreciation is also expressed to Mr.
Kazuo Kitagawa, for his kind cooperation andlending of the specimens; to Teruo Ono, for hiskind support in the course of this work; to Mrs.
Tomoko Tsuda, for her reading of the earlyAtsushi KANEKO

488

version of the rough draught; to the librarians ofDepartment of Geology and Mineralogy, Facultyof Science, Kyoto University, for offering convenience to the author; and Messrs. Shigeru
Dezawa, Shigeki Yamamoto and Takeshi Tsuda,for their assistance in various ways.
Repository of material: University Museum,
University of Tokyo ([PA]). Kaneko Collection(KAC-DT); 15-7 Fukaehonmachi 1-chome,
Higashinada-ku, Kobe, 658, Japan. Kitagawa
Collection (KKfc); 422 MyohOin-Maegawa-cho,Higashiyama-ku, Kyoto, 605, Japan.
References
Barrande, J. (1846) Notice preliminaire sur Ie
Systeme Silurien et les trilobites de Boheme.
Leipzig, i-vi, 97 p.
Baldis, B. A. J. (1967) Some Devonian Trilobitesof the Argentine Precordillera. International
Symposium on the Devonian System,
Calgary, vol. 2, p. 789-796, pIs. 1-2.
Billings, E. et al. (1861-1865) Paleozoic fossilsof Canada. Geol. Surv. Canada (not seen).
Chatterton, B. D. E. (1971) Taxonomy andontogeny of Siluro-Devonian trilobites fromnear Yass, New South Wales. Palaeonto-

graphica Abt. A, Bd. 137, p. 1-108, pIs. 124.
- - and Ludvigsen, R. (1976) Silicified Middle
Ordovician Trilobites from the South
Nahanni River area, District of Mackenzie,
Canada. Ibid., Abt. A, Bd. 154, p. 1-106,
pIs. 1-22.
- - , Johnson, B. D. and Campbell, K. S. W.
(1979) Silicified Lower Devonian Trilobitesfrom New South Wales. Palaeontology,
vol. 22, pt. 4, p. 799-837, pIs. 104-11l.
Clarke, J. M. (1894) The Lower Silurian trilobitesof Minnesota. preprint from Geology of
Minnesota, vol. 3, pt. 2, Paleontology, 1897.
p. 694-759, fig. 82 (not seen).
Copper, P., Tazawa, J., Mori, K. and Kato, M.
(1982) Atrypa (Devonian Brachiopoda)
from Japan. Trans. Proc. Palaeont. Soc.
Japan, N. S., no. 127 (749), p. 368-374,
pI. 59.
Corgan, J. X. (1963) Lower Devonian (Onesquethawan) marine faunules of Yukon andNorthwest Territories, Canada. Bull. Am.
Ass. Petrol. Geol., vol. 47, no. I, p. 150157, figs. 1-7.
DeHaan, W. (1933-1850) Crustacea in Siebold's
fauna Japonica (not seen).
Edgell, H. S. (1955) A Middle Devonian Iichid

Explanation of Plate 88
Figs. 1-4. Nipponarges mediosulcatus Kaneko, gen. et sp. nov.
Figs. la-e. Holotype, KAC-DT-0101 [PAl7144]. Incomplete cranidium. a rubber cast fromexternal mould, enlargement of part of glabella shown in Figs. la, b' of PI. 87 to show
junctions between longitudinal and transglabellar furrows, about x5.5. b, c, internal mould,
dorsal stereo pair, xL d, e, rubber cast from internal mould shown in b, c, ventral stereo
pair, xL
Figs. 2a, b. Internal mould of paratype pygidium, KAC-DT0102 [PAl7145]. a, b, dorsal stereo
pair, xL
Fig. 3. Internal mould of paratype, incomplete pygidium, KAC-DT-0104 [PA1747], dorsal
view, xl.
Figs. 4a, b. Fragmental pygidium, KAC-DT-0105 [PAl7148]. a, dorsal view of internal mould,
xL b, dorsal view of cast from external mould showing third marginal spine, xL
Figs. 5-7. Acanthopyge (Acanthopyge) duplicispinata Kaneko, sp. nov.
Paratype, KAC-DT-0108 [PAl7151]. Incomplete hypostome. a, b, ventral stereo pairFigs. 5a-d.
of rubber cast from external mould showing a few "pits" on the median body, which may be
attributable to preservation, x 5.5. c, d, ventral stereo pair of internal mould, x 5.5.
Figs. 6a, b. Incomplete hypostome, KAC-DT-0109 [PAl7152]. a, b, ventral stereo pair of internal mould, x5.5.
Figs. 7a, b. Incomplete hypostome, KKfc-0604 [PAl7154]. a, b, ventral stereo pair of internalmould, x1.5.

KANEKO : A Middle Devonian Trilobite Faune

Plate 88

788. A Middle Devonian Trilobite Fauna
trilobite from South-Eastern Australia. Paliiont.·Zeitshr. Bd. 29, p. 136-145, pI. 14.
Goldfuss, G. A. (1839) "Beitrage zur Petrefactenkunde", B. Beitrage zur familie der fossil en
Crustaceen. Nova Acta Phys. Acad. Caes.
Leop. Car., vol. 19, pt. 1, p. 353-364, pI.
33.
Glirich, G. (1901) Uber eine neue Lichas-Art aus
dem Devon von Neu-Sud-Wales und liber
die Gattung Lichas Uberhaupt. Neues Jahrb
fur Min., Beil., Bd. 14, p. 519-539, pis. 18,
19.
Hall, J. (1863) Preliminary notice of the fauna of
the Potsdam Sandstone. New York State
Cabinet Nat. Hist. (Albany), 16th Ann.
Rept., Appendix D, p. 119-226.
--and Clarke, J. M. (1888) Descriptions of
the Trilobites and other Crustacea of the
Oriskany, Upper Helderberg, Hamilton,
Portage, Chemung and Catskill Groups.
New York Geological Survey,' Palaeontology
of New York, vol. 7, p, 1-236, pl~. 1-59.
Hawle, I. and Corda, A. J. C. (1847) Prodrom
einer Monographie der bohmischen Trilobiten. 176 p., 7 pis., Prague.
Holzapfell, E. (1895) Das Obere Mittldevon(Schichten mit Stringocephalus burtini und
Maeneceras terebratum) im Rheinischen
Gebirge. Abh. preu(J. geol. Landesanst.,
N.F. Bd. 16, p. 1-459, pis. 1-19.
Kayser, E. (1880) Dechenella, Ejne Devonische
Gruppe der Gattung Phillipsia. Zeitschr.
Deutsch. Geol. Gesell., Bd. 32, p. 703-707,
pI. 27.
Kobayashi, T. (1957) Notes on two Devonian
trilobites from the Kitakami mountains inJapan. Jour. Fac. Sci. Univ. Tokyo, sec.
2, vol. 11, pt. 1, p. 1-10, pI. 1.
--and Hamada, T. (1974) Silurian Trilobites ofJapan in comparison with Asian, Pacific andother Faunas. Palaeont. Soc. Japan, Sp.
Pap. no. 18, p. 1-155, pis. 1-12.
- - and - - (1977) Devonian Trilobites ofJapan in comparison with Asian, Pacific andother Faunas. Ibid., no. 20, p. 1-202, pis.
1-13.
Maximova, Z. A. (1960) Palaeontological basis of
the Palaeozoic stratigraphy of the Rudny
Altai 7: Devonian and Carboniferous trilobites of the Rudny Altai. Trudy, VSEGEI,
vol. 7, p. 1-123, pis. 1-9 (in Russian).

489--(1968) Middle Palaeozoic trilobites ofCentral Kazakhstan. Trudy, VSEGEI, N.S.,
vol. 165, p. 1-208, pis. 1-35 (in Russian).
- - (1977) Devonian trilobites from Novaya
Zemlya and other regions of the Soviet
Arctic. Ezheg. Vses. Paleordol. Obsh., vol.
20, p. 140-185 (in Russian, not seen).
Minato, M. et al. (1979) The Abean Orogeny.
427 p. 144 pis. Tokai Univ. Press, Tokyo.
Nan, J. (1976) Trilobita: in Bureau of Geology,
Autonomic Province of Inner Mongolia andNorth China Institute of Geological Science(ed.) Palaeontological Atlas of North China.
Inner Mongolia, pt 1. Paleozoic. 431 p.,
232 pis., Geological Publishing House,
Peking (in Chinese).
(1980) Trilobita: in Shenyang Institute ofGeology and Mineral Resources (ed.) Palaeontological Atlas of Northeast China, pt. 1.
Paleozoic. 686 p., 261 pis., Geological
Publishing House, Peking (in Chinese).
Ochs, G. and Wolfart, W. (1961) Geologie derBlankenheimer Mulde (Devon, Eifel). Abh.
Senckenberg. Naturf Gesell., Bd. 50, p. 1100, pIs. 1-5.
Okubo, M. (1950) On the Gotlandian and
Devonian Deposits of Hikoroichi, KesenDistrict, Iwate-Prefecture. Jour. Geol. Soc.
Japan, vol. 56, no. 657, p. 345-350 (inJapanese with English abstract).
--(1951) Trilobites from Japan. ChikyuKagaku, no. 4, p. 133-139, pI. 1 (in Japanese with English description of species).
- - (1956) Some Devonian Brachiopods andTrilobites of the Southern Kitakami Mountinland. Japan. Jour. Geol. Geogr., vol. 27,
p. 37-45, pIs. 1-3.
Ormiston, A. R. (1967) Lower and Middle
Devonian trilobites of the Canadian Arctic
Islands. Geol. Surv. Canada, Bull. 153, p.
1-148, pis. 1-17.
--(1982) Terrano via, Arctic Vicariant of the
Devonian Lichid Ceratarges (Trilobita). Jour.
Paleont., vol. 56, no. 5, p. 1257-1265, pis.
1,2.
Perry, D. G. and Chatterton, B. D. E. (1976)
Phacops and other trilobites from Emsian
age beds of the Delorme Formation, Mackenzie Mountains, Northwest Territories.
Can. Jour. Earth Sci., vol. 13, p. 14661478, pIs. 1, 2.

490

Atsushi KANEKO-and - - (1977) Silurian (Wenlockian)trilobites from Baillie-Hamilton Island, Canadian Arctic Archipelago. Ibid., vol. 14, p.
285-317, pis. 1-7.
Plbyl, A. and Erben, H. K. (1952) Uber einige
neue oder wenig bekannte Acanthopyginae(tril.) des bohmischen und des deutschen
Devons. Paliiont. Zeitschr., Bd. 26, p. 141174, pis. 9-12.
.
and Vanek, J. (1971) Studie liber die Familie
Scutelluidae Richter et Richter (Trilobita)
und ihre phylogenetische Entwicklung. Acta
Univ_ Carolinae, Geol., no. 4, p. 361-394,
pis. 1-10.
Reed, F. R. C. (1902) Notes on the genus Lichas.
Quart. Jour. Geol. Soc. London, vol. 58,
p.59-82.
-(1923) The subgenera of Lichas. Geol. Mag.,
vol. 60, p. 454-462.
Richter, R. and Richter, E. (1917a) Uber einzelne
Arten von A cidasp is, Lichas, Cheirurus,
Aristozoe, Prosocoelus, Terebratula andSpirophyton aus der Eifel. Jarb. Nassau. Ver.
Naturk. Wiesbaden, Bd. 70, Jg., p. 143-161.
and - - (1917b) Die Lichadiden des Eifeler
Devons. N. Jarb. Miner. Geol. Paliiont.,
Jg. Bd. 1, p. 50-70, pis. 5, 6.
- - and - - (1925) Unterlangen zum Fossilium
Catalogus, Trilobitae III. Senckenberg., Bd.
7, p. 239-244.
- - and - - (1930) Neue Horner-Trilobiten
in unserer Schausammlung. Natur undMuseum, Bd. 60, p. 31-45, figs. 1-10.
Shaw, F. C. (196S) Early Middle Ordovician
Chazy trilobites of New York. Mem, N. Y.
St. Mus. Nat. Hist., vol. 17,p. 1-163.
Siveter, D. J. (1977) The Middle Ordovician of
the Oslo Region, Norway, 27. Trilobites ofthe family Calymenidae. Norsk Geol. Tids-skr., vol. 56 (for 1976), p. 335-396, pis.
1-13.
Sugiyama, T. (1942) New find of Atrypa fromJapan. Jour. Geol. Soc. Japan, vol. 49, no.
581, p. 73-75 (in Japanese).
- - (1944) On the Trilobites from Japan.
Studies, Geol. Min. Inst., Tokyo Bunrika
Daigaku, vol. 1, p. 21-30, pI. 1 (in Japanese).
--and Okano, H. (1941) On the Discovery of
Phacops (s. s.) from the Nakazato Series(Middle Devonian) of the Kitakami Mountainland, Japan. Jour. Geol. Soc. Japan,
vol. 48, no. 574, p. 357-361.
Temple, J. T. (1972) Essay Review: Ontogenies
of Devonian trilobites from New South
Wales. Geol. Mag. vol. 109, no. 4, p. 373376.
Tripp, R. P. (1957) The classification and evolution of the superfamily Lichacea (Trilobita).
Ibid. vol. 94, no. 2, p. 104-122.
and Evitt, W. R. (1981) Silicified Lichidae(Trilobita) from the Middle Ordovician of
Virginia. Ibid. vol. 118, no. 6, 665-677,
pis. 1-2.
Tschernysheva, N. E. (1951) Upper Silurian and
Devonian trilobites from the Kuznetsk
Basin. 72 p., 3 pis., VSEGEI (Gosgeolizdat),
Moscow (in Russian).
Wahlen berg, G. (1818) Petrifact Telluris Svecanae.
Nova Acta R. Soc. Sci. uppsal., vol. 8, p. 1116 (not seen).
Weber, V. N. (1932) Trilobites of the Turkestan.
Trudy Vses. Geol. Razv. Objed. NKTP,
SSSR, p. 1-148, pis. 1-4 (in Russian withEnglish summary).
Whidborne, G. F. (1889) A monograph of the
Devonian fauna of the south of England. I.
The fauna of the limestones of Lummaton,

Explanation of Plate 89
Figs. 1-3. Acanthopyge (Acanthopyge) duplicispinata Kaneko, sp. nov.
Figs. la-f. Holotype, KAC-DT-0106 [PAl7149]. Incomplete cranidium. a-c, rubber cast; a, b,
dorsal stereo pair, x9.5. c, posterior view, x9.5. d-f, internal mould; d, e, dorsal stereo pair,x9.5. f, left lateral view, x9.5.
Fig. 2. Rubber cast from external mould of fragmentary cranidium showing anterior border,
KAC-DT-0107 [PAl7150], dorsal view, x9.5.
Figs. 3a-c. Paratype, KAC-DT-0110 [PA17153]. Rubber cast from external mould of incomplete
pygidium. a, b, dorsal stereo pair, x9.5. c, left lateral view, x9.5.

KANEKO: A Middle Devonian Trilobite Fauna

Plate 89

788. A Middle Devonian Trilobite Fauna
Wolborough, Chircombe Bridge and Chudleigh. Palaeontogr. Soc., vol. 42, no. 202,

p.1-46.
Whittington, H. B. (1956) Beecher's lichid
protaspis and Acanthopyge consanguinea(Trilobita). Jour. Pal., vol. 30, no. 5, p.

491

1200-1204, pI. 13!.
Yabe, H. and Sugiyama, T. (1937) Preliminary
report on the fossiliferous Gotlandian and
Devonian deposites newly discovered in the
Kitakami mountainland. Proc. Imp. Acad.
Tokyo, vol. 13, no. 10, p. 417-420.

Dong UjimqinQi JR.!i1;I'l<f~i;G&:Il;. Heitai ~f1. Hida Massif 1!l~W.It!!. Higuchi-zawa MiQ IR, Hikoroichi B ~llm, Hsingkai J!!~Jt, Iwate 'NT-, Kitakami Mountains ~t.1:lll.lt!!, Kuronbora !l p /';f. '7,
Mishan \filll, Nakazato .pflt Of una to *~HiJ!t, Ohmori-zawa *~iR, Takainari-yama ~fmr.1jJli~tJ:W.It!!.pffll7';f. /'*.p!!!IIJi:O)=:'iltlil,ib¥Qll¥'1::.-?~,-c--to) 1. Lichidae f4: <f:!$7';f.
/' *<f:! Ef!.1\lf N. nBI\lf II , G, Lichidae, Scutelluidae, Dechenellidae, Calymenidae, Phacopidae.
Dalmanitidae "&r1 Odontopleuridae O)-X-.f+t::.~-t 0 l!lMt.e=:.w.ib~II-H:·reYl L t.: • .:. 0)1/)~llfl:l:.!ti'f~*RaiJ!tm B I;f{mIllTO)*~IR&'r1tiir..lIRP-ffO)=:~W.II' >..>Jll(;o. f,r;jJi:.It!!I:!:..:..h"i -C-7i~,t::.-tO)Ji:tUlIl(Iia-~I::.-to>(,O)c ~;tG.h-C~t.:. flP~. Thysanopeltella paucispinosa(Okubo, 1951) 7Jl..r1 Dechellella (Dechl'llella) minima (Okubo. 1951) Lt. -t.h'CJl*~IR
1::.:(£-:>-C71 7 ",lvjtl:a-,tffir:Jif{I.:{E-:>-C ;;"-!I1:H~jf;-jo >(,0) c ~;:tG.h -C ~t::.. LII'L,~1ij1H:tt;iIi n IRO) iI¥..It!!I::.Jj'~-C Jlif{--j 0':' c 1I~1lJJ G 11'1': t.e -:> t.:O) -r: -&; o. !fifE -r:1 ;l'.JI#t\;;O)mjf,~ c
L-Cffl~'o':' cO)-r:~o >(,0)1:1:. Thysanopeltella O).;z,.-r:-&;o. -tJtI&*=:'ft.!t.tb~llf.O) 1=..~,
1f.f\;;I:l:.717 ",Iv llta-rGt.e~'>(,O)c :!5·;:tG.ho • .:..hl:l:. Copper et al. (1982) O)/tlljJEJ'Ji
Atrypa O)l'*iH*,~,~c Urit-to.
*=:.!p'fjI)~lI'f.O)t~fll1;l::'r,.g L -C 1:1:. • .\!is v "It.- -r:1:t.JR~t 13 *1.:rn:ltfO)~~ C ~;:t G.ho >(, O)II~
Ij,'t.e < t.e <, :lltlmlW:I\f!O)rffll7';f. :/*=:1t!1tmb¥!.Jlffcl:l:.t.:t-':iJLtll:W-JI~O) Acanthopyge (s.l.)
1 JlltO).;z,.a-~jijjl::" -t 0 I::'~ ~t.e~ '. H7Jl..r111P.f;~O)HltPl1;II' G.;z,. -C >(, *=:W.!t.ib¥!.JlifltJl~!WlIl.It!!=:~.!t.i!iJ¥!.JlI-fO)~~-r:-&; 0 c It~·;:t Gilt! ~ '.
*xrKlc.!n-Ca Lichidae HI::.~-t 0 2m1!ia-~ctfit 1..-, j1;Jr} c -?I::.:x-t LmJllta-Wp~ LNippoll.
arges c {.piS Lt::.. ~~I:!:.ii.i\fj;<f:!Jl:J:a-~IJ~U.eliJ{.n;_ (2p 7Jl..rJ 3p) 1I~jJ'/t1;1Jo.~, "c*Hc L-C
IHH~-r:~o. mAt\tO)¥rf!i Nipponarges mediosulcalus t)'71-1::..: 0) J: Ij t.dt'i!fa-n.;-t >(, 0)l.t)/U-':ifl G.h -C~ ,t.e~ '. ft!!O) 1 ¥r~I:I:. Acanthopyge (Acallihopyge) duplicispinata -r:-&; o.
~p.:~I:I:., tffinIRPl\:O)=:~!Ua-Jlffi;*;\c~xL-Cfj< T'~-C·.:l?o.
1i.J.

m

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 136, pp. 492-501, pIs. 90-92, December 30, 1984

789. CASSIDULINIDAE (FORAMINIFERIDA) FROM THE EASTERNPART OF LOTZOW-HOLM BAY, ANTARCTICA*
RITSUO NOMURA
Department of Earth Sciences, Faculty of Education,
Shimane University, 1060 Nishi·kawatsu, Matsue 690

Abstract.

Four species of Cassidulinidae, Globocassidulina biora (Crespin),porrectus (Heron-Allen and Earland), C. parvus (Earland) and
Ehrenbergma glabra Heron-Allen and Earland, from the raised beaches and modern
bottom sediments in the eastern part of Liitzow-Holm Bay, Antarctica are described and revised on the basis of ontogenetic and new anatomic information.
The ontogenetic development of the aperture in G. biora is divided into thefollowing three steps: I-shaped, L-shaped, and =-shaped, so that G. biora is morphologically similar to Globocassidulina subglobosa (Brady) and G. crassa rossensis
Kennett in immature stages respectively. Uncoiled forms, such as C. porrectus andC. parvus, are also similar to G. subglobosa in globular immature stages.
Cassidulin~ides

Introductionof their having a similar external morphology.
Ever since Loeblich and Tappan (1964a) erected
the family Islandiellidae for the forms having
radial wall texture and a toothplate, and distinguished it from the family Cassidulinidae
there has been much confusion in foraminifer~systematics. However, their suprageneric classification has been questioned by several subsequent
workers as described by Nomura (1983b).
Further pertinent information is needed toresolve this controversial problem, such as theinternal character of the test and its morphologic
variability, in addition to the wall structure. Thus
the purpose of the present work is to provide
systematic infQrmation on the Antarctic cassidulinid foraminifera by integrating several
taxonomic characters, including the ontogenetic
and anatomic observations, in order to document
the sil.lilarities and differences in internal andexternal morphology within the Cassidulinidae.

During systematic studies of the foraminiferal
family Cassidulinidae, many specimens wereobtained from the bottom sediments and Pleistocene raised beaches in the eastern part of LiitzowHolm Bay, Antarctica. As a result of detailed
examination, four species assigned to three
genera were determined: Globocassidulina biora(Crespin), Cassidulinoides porrectus (Heron-Allenand Earland), Cassidulinoides parous (Earland)
and Ehrenberginaglabra Heron-Allen and Earland.
They are the most characteristic species in the
Antarctic foraminiferal province.
In recent years, a number of paleontologists
have focused attention on the wall structure andinternal character of foraminifera. Among
foraminiferal taxa, the family Cassidulinidae
has attracted the most attention because members possessing different wall structures in spite*Received October 14, 1983
492

789.

Cassidulinidae from Liitzow-Holm Bay, Antarctica

Materials and Methods
Materials used in this study were taken fromthe raised beaches (9 and 11.5 m above sea
level) in Skarvsnes and Langhovde and Recent
bottom sediments at the depths 31.5 and 98 m
around East Ongul Island, which were collectedby Mr. M. Hayashi of Geographical Institute,
Shimane University, during the 16th Japanese
Antarctic Research Expedition, 1975-1976.
Their localities are as follows:
Sample 75012901: Raised beach sand, 11.5
m above sea level, Skarvsnes (69°29.2'S,39°38,4'E).
Sample 75110201: Raised beach sand, 9 m
above sea level, East Ongul Island (69°0.7'S,39° 34.5'E).
Sample 75112101: Recent bottom sediments, 31.7 m in depth, northeast of East
Ongul Island (68°59.7'S, 39°37.3'E).
Sample 75112102: Recent . bottom sediments, 98 m in depth, north of East Ongul
Island (69°00'S, 39°33.1'E).
The materials from the raised beaches consistof massive, poorly sorted sand containing molluscan fossils, Laternula elliptica (King andBoderip), Adamussium colbecki Smith, andother fossils such as serpuloid tubes. The material
studied here is from the foraminiferal assemblages
from the Pleistocene raised beach sediments
described by Nomura (1983d).
For anatomical observations using a SEM,
the specimens were prepared using the Canadabalsam-xylene method described by Nomura(1983a).
Systematic Description
Before going further, several morphologic
terms for apertural structures based on comparative anatomy, which were defined by
Nomura (1983b) in the study of Japanese Cassidulinidae need to be explained.
Primary tongue (PT): internal free portion of
toothplate.
Cristate tooth (CT): external free portion of
toothplate.

493

Copula (CP): conjugate portion of primary
tongue and cristate tooth.
Cavity (CA):depressed portion between
septal chamber wall and primary tonque.
Sulcus (S): depressed portion between septal
chamber wall and cristate tooth or lip.
Lip (L): overturned structure of septal chamber wall, which is linked with the cristate
tooth.
Secondary tongue (ST): free portion formedon the anterior comer of the aperture.
Apertural ridge (AR): ridge-like structure at
the base of the aperture, which is formedon the preceding chamber.
Apertural flap (AF): imperforate plate formedon the lower side of the aperture.
Capitals in brackets indicate the abbreviation
used in illustrations.
Family Cassidulinidae d'Orbigny, 1839
Genus Globocassidulina Voloshinova, 1960
Globocassidulina biora (Crespin)
Pl. 90, Figs. la, b-3, 12jPl. 91, Figs.1-7j
Text-fig. 1

Cassidulina biora Crespin, 1960, p. 28, 29, pi. 3,
figs. 1-10.

Cassidulina crassa d'Orbigny. Crespin, 1960,
p. 29, pl. 3, figs. 12,13.

Cassidulina sp. Crespin, 1960, p. 29, 30, pl. 3,
fig. 11.

Globocassidulina biora (Crespin). FilIon, 1974, p.
139, pi. 1, figs. 9-12, 14, 15; Osterman and
Kellogg, 1979, p. 264, pi. 2, fig. 1; Finger andLipps, 1981, p. 129, pl. 4, figs. 6a~, 7a, b.
Globocassidulina biora ? (Crespin). Filion, 1974,
p. 139, pi. 1, figs. 8, 13.
Globocassidulina ? sp. Filion, 1974, p. 140, pl.
4, fig. 4.
Anticieina antarctica Saidova, 1975, p. 331,
pi. 89, fig. 14, pl. 114, fig. 5.
Globocassidulina cf. biora (Crespin). Milam andAnderson, 1981, pi. 10, figs. 2, 3a, b.
Globocassidulina crassa rossensis Kennett. Milamand Anderson, 1981, pi. 10, figs. la, b.
Globocassidulina sp. Finger and Lipps, 1981,
p. 129, pl. 4, figs. 1a-c, 2a~, 3a-c, 4a~.

494

Ritsuo NOMURA

External morphology:-The external morphology of Globocassidulina biora has been
described in detail by Crespin (1960) in her
original description. The test is generally subglobular, but somewhat prolonged and compressed-oval in side and edge views respectively
in mature stage. Chambers in final whorl areusually five pairs, and slightly inflated or not; sothat the periphery of the test is lobulate orsometimes smooth. The umbilical rep,ion isslightly depressed. Sutures are almost straightand narrowly limbate. The aperture, as described
below, consists of two openings parallel witheach other at the middle portion of the final
apertural face. However, its shaped varies froma vertical slit to tripartite in the immature stage.
The test wall is polished and translucent andpierced with fine pores.
Through ontogenetic development, the external morphology of the test is divided into
three growth stages: nepionic to neanic, ephebic
and gerontic stages. In nepionic to neanic stages,
two forms, the megalospheric and microspheric
are easily distinguished as follows: the former ischaracterized by a globular test with proloculus
diameter of 72 .urn in average, whereas the lattershows a compressed test having a proloculus
diameter of less than 10 .urn. In the ephebic
stage, the test shows the shape described above .
The gerontic stage is characterized by having amore prolonged test with a tendency toward an
uncoiled appearance.
Apertural structure:-The peculiar structureof the aperture in both external and internalviews is the most diagnostic feature of thisspecies. As noted in Crespin's original description,
the aperture is divided into two narrow openings
both parallel to the periphery of the preceding
chamber margin; the longer one is usually situatedon the proximal side.
SEM observation reveals that the aperturalcompartment is formed by the junction of astrongly developed cristate tooth and partly
protruded lip (Text-fig. 1, nos. 4, 5). Text-fig.
1 shows the developmental mode of the cristate
tooth and lip. As seen in these figures, the development of the cristate tooth through ontogeny is larger than that of the lip. Similar
developmental pattern is noted by Finger andLipps (1981) as described later. Owing to theconspicuous elongation of the cristate tooth
toward the lip, the sulcus is not significant.
Each tip of the cristate tooth and lip is nodulate,
and an apertural ridge is also formed in early
growth stage.
When viewed from the interior in dissected
specimens, it is noted that the anterior andposterior corners of the aperture are devoid of
free structures corresponding to primary andsecondary tongues (PI. 91, Fig. 4).
Another notable character of the aperture isthe variable feature in the outline of apertural
openings through the nepionic to gerontic.
Nepionic to neanic apertures are almost verticallyarranged loop-like slits to the basal suture of theapertural face . Subsequently the early ephebic
stage aperture shows a tripartite feature consisting of basal and areal apertural branches, in

Text-fig. 1. Apertural development of Globocassidulina biora (Crespin). CT=cristate tooth,
L=lip , AR=apertural ridge . 1; neanic stage, 10 /lm scale. 2 ; early ephebic stage, lO.um scale. 3; early
ephebic stage, 10 .urn scale. 4; ephebic stage, 10 .urn scale. 5; late ephebic stage, 100 .urn scale.

789. Cassidulinidae from Liitzow-Holm Bay, Antarctica
which the cristate tooth is formed on the posterior side of the areal one. Such mature stages
as ephebic and gerontic demonstrate the parallel
openings (Text-fig. 1).
Wall texture:-Fragments of test wall crushed
in Lakeside cement reveal a jagged-granular
texture under crossed nicols. However, the septal
wall of the proloculus, being very thin, is also
sometimes similar to the mosaic structure (PI. 90,
Fig. 12).
Perforation:-Pores are slit-shaped, parallelwith each other on the earlier portion (PI. 91,
Fig. 1), irregular in later (PI. 3, Fig. 2), androunded on the final chamber; length of slitshaped pores is 3.3 J.l.m in average; pore density
is 16 pores per 20 J.l.m x 20 J.l.m.
Remarks:-Since Crespin (1960) described
this species from the Vestfold Hills area of
Antarctica, it has been reported as being very
common in the Antarctic region. However, theapertural morphology of this species is not
uniform and the ontogenetic variation of theaperture used to distinguish this species fromother related species has not been well known
until now. Owing to the morphological changeof the aperture through ontogeny, much confusion, particularly in the form of immature
stages, is seen in recent literature.
In the original description of this species,
Crespin was not aware of its ontogenetic variation, so that seemingly immature forms whichshould have been assigned to this species were
not included in the continuous range of thevariation (e. g., Cassidulina sp. in her pI. 3, fig.
11 clearly shows a rudimentary cristate tooth).
All dissected specimens here examined clearly
demonstrate that continuous variation of theaperture is divided into three basic steps throughontogeny: slit (I-shaped) -+ tripartite (L-shaped)-+ parallel openings (=-shaped). Finger andLipps (1981) similarly noted these ontogenetic
changes of the aperture in the Globocassidulina
crassa plexus (particularly G. crassa rossensis
and G. biora), in stating that the G. subglobosa-like aperture grades into the aperture characterized by one or two apertural branches.
In fact, this ontogenetic variation has per-495

plexed the species identification and the relationship of G. biora, G. crassa rossensis Kennett,
and G. subglobosa (Brady) as discussed byFillon (1974), Kellogg et al. (1979), and Finger
and Lipps (1981). Fillon (1974) considered thatG. biora and G. crassa rossensis are in close
phylogenetic relationship because of their close
similarity in adult forms. On the other hand,pointing out the resemblance of immature formsof G. biora and G. crassa rossensis, Finger andLipps (1981) stated "dissection reveals thatearlier chambers of the larger (adult) G. crassa
rossensis and G. biora display their characteristic(fully developed) apertures without any indication of apertural growth stages". So far as G.
biora, however, their observations are not inagreement with the present ones. As alreadystated, G. biora shows the variation of apertural
shape during ontogeny.
Close examination of the final apertures of
the holotype and paratypes illustrated by Kennett (1967, p. 134, pI. 11, figs. 4-6) and subsequent workers including Finger and Lipps(1981) shows that G. crassa rossensis possesses
a tripartite aperture with a very long basal andless developed cristate tooth. These aperturalfeatures indicate that the two are clearly differentiated in the mature stage. However, G. biora
and G. crassa rossensis have a similar ontogenetic
development in the shape of the test (Text-fig. 2)
and G. biora possesses a tripartite (L-shaped)
aperture like that of G. crassa rossensis in immature stage. Taking these features into consideration, the two taxa are not conspecific, butthey are expected to be grouped in the closest
phylogenetic relationship as stated by Fillon(1974).
Genus Cassidulinoides Cushman, 1927

Cassidulinoides porrectus (Heron-Allen andEarland)
PI. 90, Figs. 4a, b-7, 13;PI. 92, Figs. 6-9

Cassidulina crassa d'Orbigny var. porrecta HeronAllen and Earland, 1932, p. 358, 359, pI. 9,
figs. 34-37.

Ritsuo NOMURA

496

I'm
1000

900

(A)

•

• =- shaped aperture

(B)

)'m
1000

(micro- 8. megalospheric forms)- L - shaped (microspheric)o L - shaped (megalospheric).. 1 - shaped(microspheric)A I - shaped(megalospheric)

•

900

•
•
•

700

•
700

J:

•••

• •••

J:

~500

1-500

o

Z

••

W

-'

.~

.W· •
••••

~
-(:

300

-

300

* Globocassidulina
crassa rossensis

••

0

o

0

__ a_A""•:.;=after KENNETT(1967)

100

••

"h

100

100

500

300

WIDTH

700

}1m

100

300

500

THICKNESS

Text-fig. 2. Interrelations of test length and width (A) and test width and thickness (B) for
Globocassidulina biora (Crespin) showing the apertural type. Sample from 75012901.

Explanation of Plate 90
Figs. 1-3. Globocassidulina biora (Crespin)la, b; ephebic stage (DESS 82038), x60. 2; early ephebic stage (DESS 82039), x105. 3;
neanic stage (DESS 82040), x105.
Figs. 4-7. Cassidulinoides porrectus (Heron-Allen and Earland)
4a, b; ephebic stage (DESS 82041), x45. 5; neanic stage (DESS 82042), x 60.6; neanic stage(DESS 82043), x 60. 7; neanic stage (DESS 82044), x 60.
Figs. 8, 9. Ehrenbergina glabra (Heron-Allen and Earland)
8; megalospheric form (DESS 82045), x65. 9a, b; microspheric form (DESS 82046), x80.
Figs. lOa-c. Cassidulinoidesparvus (Earland), (DESS 82047), x80.
Figs. 11-14. Wall textures, 40 /J.m scale.
11; jagged-granular texture (Ehrenbergina glabra). 12; mosaic-granular texture (neanic stageof Globocassidulina biora). 13; indistinctly radial texture (Cassidulinoides porrectus). 14;
distinctly radial texture (Cassidulinoides parvus).
DESS: Department of Earth Sciences, Shimane University

NOM URA: Cassidulinidae from Antarctica

Plate 90

4c

4b

9b

10a

10c

5

789. Cassidulinidae from Liitzow-Holm Bay, Antarctica
Cassidulinoides porrectus (Heron-Allen and Earland). Parr, 1950, p. 344, pi. 12, fig. 26;
Filion, 1974, p. 139, pi. 4, figs. 1-3.
Cassidulina crassa porrecta Heron-Allen andEarland. McKnight, 1962, p. 126, 127, pi.
21, fig. 139.
Cassidulinoides porrecta (Parr). Osterman and
Kellogg, 1979, p. 264, pi. 2, figs. 8, 9.
Cassidulinoides porrecta (Heron-Allen and Earland). Milam and Anderson, 1981, pi. 6, fig. 6.
Ex ternal morphology:-Cassidulinoides porrectus has been considered to have a pyriform
test, though in the immature (nepionic to neanic)
stage it has a globular test like that of Globocassidulina subglobosa (B"rady). Therefore, the relative growth shows isometry in the immature
stage and positive allometry in the mature stage(Text-fig. 3). These features are described in thefollowing three growth steps.
Nepionic to neanic tests showing isometric
growth are globular to oval, and have three pairsof chamber just before attaining the ephebic
stage (PI. 90, Figs. 5, 6). Ephebic tests, showing
positive allometry, form an uncoiled chamber(-100) pm
10
9
8
7
6
5

J:4

lt!)

Z 3
W
oJ

2

1~----~--~--~~~~r--

1

2

3

WIDTH

4

5 6 7 8 pm(-100)

Text-fig. 3. Interrelation of test length andwidth for Cassidulinoides porrectus (Heron-Allenand EarIand), indicating the isometry in immature stage and positive allometry in mature stage.
Sample from 75012901.

497

arrangement. In the gerontic growth stage,
tests are somewhat cylindrically prolonged(Nomura, 1983d, pI. 2, figs. 2a, b).
In contrast to the original description, chambers are distinct and somewhat inflated throughthese growth stages. Sutures are distinct, slightly
depressed and limbate. The last-formed chamber
is narrowed. The aperture is a loop-like slit
arranged almost at right angles to the basal
suture of the apertural face, and situated at the
middle portion of the face. The wall in wellpreserved state is transparent, polished, pierced
by numerous fine pores, and earlier chambers in
the preceding whorl are visible externally. However, poorly preserved walls are opaque as in theoriginal description.
Apertural structure:-Detailed examination of
the aperture reveals that the loop-like slit opening is surrounded by a well-developed cristate
tooth on the posterior side and a lip on theanterior side (PI. 92, Fig. 6). In the immature
stage, the cristate tooth is attached to the lip on
its anterior end, but not in mature stages (PI. 92,
Figs. 6, 8, 9).
Dissected specimens show that the primary
tongue is very narrow even in the mature stage,
but it appears to be present from the nepionicaperture onwards; a rudimentary primary tongue
with cavity is clearly observable in the first whorl(PI. 92, Fig. 8). On the contrary, the secondary
tongue is not formed throughout ontogeny.
Morphological change in the aperture throughontogeny is very slight and not serious for
specific distinction of this species from othersimilar forms; the loop-like slit gradually increases in length as chambers are added.
Wall texture:-In polarized light, fragments oftest wall reveal indistinctly radial texture (PI. 90,
Fig. 13).
Perforation:-Pores are rounded in all growthstages (PI. 92, Fig. 7); pore diameter is 1.3 /.lm
in average and pore density is 7 pores in 20/.lm x
20/.lm.

Remarks:-In at least immature forms such as
the nepionic and neanic stages, there are someapparent similarities between C. porrectus andGlobocassigulina subglobosa. The two species

498

Ritsuo NOMURA

have subglobular tests and the apertures areperpendicular to the basal suture. However,
the wall texture differs; the wall of G. subglobosa
is of jagged-granular texture (Nomura, 1983c).
The ephebic to gerontic specimens of thisspecies are apparently close to Cassidulinoides
parkerianus (Brady), illustrated in the Challenger
Report, in the size and shape of the test, having
a protruded final chamber. However, C. porrectus differs from C. parkerianus in having amore compressed test and narrower apertualface.
Cassidulinoides parvus (Earland)

PI. 90, Figs. 10a--c, 14; PI. 92, Figs. 1-5
Ehrenbergina parva Earland, 1934, p. 139, 140,

pI. 6, figs. 28-32.
Cassidulinoides parkerianus (Brady). Parr, 1950,

p. 344, pI. 12, fig. 25; McKnight, 1962, p.
127, pI. 22, fig. 141; FilIon, 1974, p. 139,
pI. 4, fig. 5; Finger and Lipps, 1981, p. 129,
pI. 2, fig. 8.
External morphology:-Test is smali for the
genus; test width is up to 120 p.m in microspheric
form and 170 p.m in megalospheric, and length is
up to 600 p.m in microspheric form and 400 p.m
in megalospheric. As described by Earland(1934), this species possesses a biseral arrangement throughout, and it is usually found at thecrosier-shaped state consisting of a very small
coiled or loosely coiled early portion and linear
or arcuate later one. Many microspheric forms

here examined indicate the following two growthstages: Nepionic to neanic tests are very small(140 p.m in length, 120 p.m in width, and 100 p.m
in thickness) and globular in shape; ephebic togerontic stages show uncoiled-linear development(PI. 92, Fig. 3).
Chambers are inflated, particularly distinct
in later growth stages; sutures are therefore
depressed and the test periphery is lobulate.
The aperture is a loop-like slit set obliquely inthe middle portion of the final apertural face,but in mature stages it is usually situated subterminally on the apertural face. Well-preserved
walls are transparent, showing the earlier chambers.
Apertural structure:-Viewed from the exterior, the apertural structure is simple (PI. 92,
Fig. 1). The cristate tooth is prominent, but its
anterior and posterior ends are linked to'. thelip (PI. 92, Fig. 2). Internally no free structures
are observed (PI. 92, Figs. 3-5); a copula without the primary tongue originates from the
middle portion of the anterior apertural face,close to the preceding aperture. The proximal
end of the aperture is not fused with the preceding or previous chamber, so that an oval apertural
opening is formed in the subterminal face. Thus,
there is apparently no marked change in apertural structure and shape of the apertural opening through ontogenetic growth.
Wall texture:-All fragments of test wallshow distinctly radial texture in polarized light(PI. 90, Fig. 14).
Perforation:-Pores on coiled and uncoiled

Explanation of Plate 91
CT=cristate tooth, L=lip
Figs. 1-7. Globocassidulina biora (Crespin)
1; outer test surface showing slit-shaped pores, 10 p.m scale. 2; outer test surface showing
irregular pores, 10 p.m scale. 3; dissected specimen, 100 p.m scale. 4; enlargement of Fig. 3,
100 p.m scale. 5; internal structure showing the change of apertural shape, 20 p.m scale. 6;
internal structure showing proloculus aperture, 10 p.m scale. 7; internal structure of neanic
stage, 20 p.m scale.
Figs. 8, 9. Ehrenbergina glabra Heron-Allen and Earland
8; dissected specimen, 100 p.m scale. 9; internal structure showing apertural development,
10 p.m scale.

NOMURA: Cassidu linidae from Antarctica

Plate 91

789. Cassidulinidae from Liitzow-Holm Bay, Antarctica

chamber walls are rounded without decoration.
Pore diameter is 0.5 p.m in average and poredensity is 8 pores in 20 p.m x 20 p.m.
Remarks:-This species was originally described under the genus Ehrenbergina by Earland(1934). Compared with other typical Ehrenbergina species, however, such morphologicalaspects as chamber shape and apertures indicatethat this species is more reasonably put in
Cassidulinoides; the genus Cassidulinoides ischaracterized by having a non-{!ompressed test
on both dorsal and ventral sides, globular chambers and subterminal loop-shaped aperture,
whereas the genus Ehrenbergina is characterizedby compressed, laterally prolonged chambers
and elongate slit aperture with apertural flap.
The presence of a tooth plate has been used as
one of the basic criteria for the genus Cassidulinoides (Loeblich and Tappan, 1964a). However,C. parous does not possess the tooth plate as described above. From this point of view, Nomura(1983b) stressed that the structure of the toothplate is variable between respective species aswell as within one species. Therefore the presence or absence of a toothplate is not sufficient
to justify generic distinction. The same is true
for Cassidulinoides parous, which is devoid of a
primary tongue in spite of the presence of acristate tooth.
It is noted that C. parous has not been reported so often from the Antarctic region. In
the original description, Earland pointed that
this species is confined to the Bransfield Strait,
South Shetlands and Palmar Archipelago, andthat it is easily overlooked owing to the small
size. However, this species is commonly foundin Liitzow-Holm Bay area.
Genus Ehrenbergina Reuss, 1850
Ehrenbergina glabra Heron-Allen and Earland

PI. 90, Figs. 8, 9a, b, 11, PI. 91, Figs. 8,9Ehrenbergina hystrix Brady var. glabra HeronAllen and Earland, 1922, p. 140, pI. 5, figs.
1-6,11.

499

Ehrenbergina glabra Heron-Allen and Earland.
Uchio, 1960, pI. 1, figs. 1-3; McKnight,
1962, p. 127, pI. 22, figs. 142a, b; Fillon,
1974, p. 139, pI. 5, figs. 9, 10; Anderson,
1975, p. 86, pI. 11, figs. 2a, b; Saidova, 1975,
pI. 89, fig. 13; Osterman and Kellogg, 1979,
p. 264, pI. 2, fig. 4; Milam and Anderson,
1981, pI. 9, fig. 7.
External morphology:-The test is triangular,
with laterally prominent spines. Spines are welldeveloped in later ontogenetic stages, as they
gradually increase in length in harmony withchamber development. In both megalo- andmicrospheric forms, the earliest portion is coiled,
but its roundness is different due to the size of
proloculus; microspheric forms appear to be
sharpened, whereas megalospheric ones are morerounded. Chambers are low and broad, notinflated; and overlap about half of the breadthon the dorsal side. Sutures are limbate and wellshown on the dorsal side, but on the ventral side
the limbation is not distinct. The aperture isa long-narrow arcuate slit almost parallel to thedorsal line in apertural view. The wall is translucent, pierced by fine pores, but not perforateon the apertural flap.
Test length is up to 765 p.m; width up to
740 p.m; thickness up to 380 p.m.
Apertural structure:-Owing to the development of an apertural flap, the aperture of thisspecies is a simple slit. However, many dissected
specimens reveal that a primary tongue andcristate tooth are formed on the dorsal side of
the aperture. These structures, which are formedby the turnover of the septal chamber wall on
the dorsal side, are internally connected witheach other.
Ontogenetically viewed, a rudimentary structure is already formed in the earliest stages,presenting a very shallow cavity beneath theprimary tongue. But compared with the development of the cristate tooth, the primary tongue
is not so conspicuous (PI. 91, Figs. 8, 9).
Wall texture:-Examination of the test throughpolarized light shows that walls are of jaggedgranular texture (PI. 90, Fig. 11).

Ritsuo NOMURA

500

Perforation:-Pores are slit-shaped on earlier
chambers, rounded to irregular on later portion,
but absent on the apertural flap as well as theapertural margin. Length of slit-shaped pores is
1.2 /J.m in average; pore density is 8 pores in 20/J.m x 20/-tm.
Remarks:-Heron-Allen and Earland (1922)
first described this taxon from many stations
around the seas of New Zealand's sUbantarctic
islands and the Antarctic, as one of the mostthriving forms in Antarctic cold water. At thattime, they considered this species to be a varietyof E. hystrix, pointing out such differences asabsence of apertural grooves and a more inflated
apertural face. As discussed by Loeblich andTappan (1964a, b), however, this species isdistinctly differentiated from E. hystrix (= Reissia hystrix, Loeblich and Tappan, 1964b) inhaving the test wall of jagged-granular texture.
Recognition of E. glabra as a distinct species issafely accepted.
So far as examined in the literature, thisspecies seems to bear a greater resemblance toE. pupa (d'Orbigny). Some specimens show very
short peripheral spines, but typical pupa-likeforms are not found from Lutzow-Holm Bay
area.
Acknowledgements
I express my sincere gratidude to Mr. M.
Hayashi of Shimane University, for providing
the materials. I also thank Associate Professor
Yasumochi Matoba of Akita University, for his

valuable suggestions and critical reading themanuscript; Dr. B. W. Hayward of New ZealandGeological Survey, Lower Hutt, for kindly reading over the manuscript.
References
Anderson, J. B. (1975): Ecology and distributionof foraminifera in the Weddell Sea of Antarctica. Micropafeontology, vol. 21, no. 1,
p. 69-96, pis. 1-11.
Crespin, 1. (1960): Some Recent foraminiferafrom Vestfold Hills, Antarctica. Sci. Rept.
Tohoku Univ. 2nd ser. (Geol.), Spec. Vol.,
no. 4, p.19-31, pis. 1-3.
Earland, A. (1934): Foraminifera, Part III. The
Falklands sector of the Antarctic (excludingSouth Georgia). Discovery Repts., vol. 10,
p. 1-208, pis. 1-10.
Fillon, R. H. (1974): Late Cenozoicforaminiferal
paleoecology of the Ross ~ea, Antarctica.
Micropaleontology, vol. 20, no. 2, p. 129151, pis. 1-6.
Finger, K. L. and Lipps, J. H. (1981): Foraminiferal decimation and repopulation in an
active volcanic caldera, Deception Island,
Antarctica. Micropaleontology, vol. 27, no.
2, p. 111-139, pis. 1-4.
Heron-Allen, E. and Earland, A. (1922): Protozoa, Part II. Foraminifera. Brit. Antarctic("Terra Nova") Exped., 1910, Zool., vol. 6,
no. 2, p. 25-268, pis. 1-8.
- - and - - (1932): Foraminifera, Part 1. The
ice-free area of the Falkland Islands andadjacent seas. Discovery Repts., vol. 4, p.
291-460, pis. 6-17.

Explanation of Plate 92
C=cavity, CP=copula, CT=cristate tooth, L=lip, P=proloculus, PT=primary tongue, S=sulcus
Figs. 1-5. Cassidulinoides parvus (Earland)
1; external structure of aperture in mature stage, 10 /J.m scale. 2; external structure ofaperture in immature stage, 10 /J.m scale. 3; dissected specimen, 50 /J.m scale. 4; internal
structure of uncoiled portion, 20 /J.m scale. 5; enlargement of Fig. 3, showing aperturaldevelopment, 20 /J.m scale.
Figs. 6-9. Cassidulinoides porrectus (Heron-Allen and Earland)
6; external structure of aperture, 100 /J.m scale. 7; outer test surface showing rounded pores,
10 /J.m scale. 8; internal structure of coiled portion, 20 /J.m scale. 9; internal structure of
uncoiled portion, 30 /J.m scale.

NOMURA: Cassidulinidae from Antarctica

Plate 92

789. Cassidulinidae from Lutzow-Holm Bay, Antarctica
Kellogg, T. B., Osterman, L. E. and Stuiver, M.
(1979): Late Quaternary sedimentology andbenthic foraminiferal paleoecology of theRoss Sea, Antarctica. Jour. Foram. Res.,
vol. 9, no. 4, p. 322-335.
Kennett, J. P. (1967): New foraminifera fromthe Ross Sea, Antarctica. Contr. Cushman
Found. Foram. Res., vol. 18, pt. 3, p. 133135, pI. 11.
Loeblich, A. R. Jr. and Tappan, H. (1964a):
Sarcodina. In Moore, R. C. (ed.): Treatiseon Invertebrate Paleontology, Protista. 2, pt.
c. Kansas University Press, 900 p.
- - and - - (1964b): Foraminiferal classification and evolution. Jour. Geol . Soc. India,
vol. 5, p. 5-40.
McKnight, W. M. Jr. (1962): The distribution offoraminifera off parts of the Antarctic coast.
Bull. Amer. Paleont., vol. 44, no. 201, p.
65-158, pIs. 9-23.
Milam, R. W. and Anderson, J. B. (1981): Dis·
tribution and ecology of Recent benthonicforaminifera of the Adelie·George V con·tinental shelf and slope, Antarctica. Marine
Micropaleont. vol. 6, p. 297-3~5, pIs. 1-11.
Nomura, R. (1983a): An embedding techniquefor observation of internal microfossil structure by scanning electron microscopy.
Micropaleontology, vol. 29, no. 1, p. 1-5,

501

pI. 1.
- - (1983b): Cassidulinidae (Foraminiferida)from the Uppermost Cenozoic of Japan,
Part 1. Sci. Rept. Tohoku Univ. 2nd ser.
(Geol.), vol. 53, no. 1, p.l-l0l, pIs. 1-25.
(1983c): Ibid., Part 2, Ibid., vol. 54, no. 1,
p. 1-93, pIs. 1-6.
- - (1983d): Foraminifera from the raisedbeach deposits on the east coast of LUtzowHolm Bay, Antarctica. Mem. Nat. [nst.
Polar Research, Spec. Issue, 28, p. 219-228,
pIs. 1, 2.
Osterman, L. E. and Kellogg, T. B. (1979):
Recent benthonic foraminiferal distributions
from the Ross Sea, Antarctica: Relation toecologic and oceanographic conditions.
Jour. Foram. Res., vol. 9, no. 3, p. 250269, pIs. 1-3.
Parr, W. J. (1950): Foraminifera. B. A. N. Z.
Antarctic Res. Exped., 1929-31, Repts.,
ser. B. (Zool., Bot.), vol. 5, pt. 6, p. 233392, pIs. 3-15.
Saidova, K. M. (1975): Benthonic foraminiferaof the Pacific Ocean. Acad. Sci. USSR, P. P.
Shirshov Inst. Oceanology, 875 p. pIs. 1116. (in Russian).
Uchio, T. (1960): Benthonic foraminifera of the
Antarctic Ocean. Bio. Res. Japan Antarct.
Res. Exped., 12, p. 3-20, pI. 1.

i¥ffi., V "- '/ ot • ;t;,J..-A~JI'i~iI'l?tD Cassidulinidae *,~tDff:fL!H.: i¥ft~tD V "- ';1* • ;t;<ifiHJx 1l.5, 9, -31.5, -98 m) iI'l? Cassidulinidae
HtD 4 fil!tD-1Hl.!t., Glbboeassidulina biora (Crespin), Cassidulinoides porreetus (Heron·'J..-.l,~*J¥:tD~~~lnJRc:mtll:iftjJttJj/J{i¥1.J

Allen and Earland), Cassidulilloides parvus (Eariand), Ehrellbergina glabl'a Heron·
Allen and Earland 1::"?\"LI~pj,:JE~:t" J: L)(Mi';IJ"f:I't-J~j!tD ~ C I::~c~ U-':oG. biora, C. porrectus, C. parvus tD 3 fiili, II'oH*1e~~ jlfI C-CJf3t!!W1:ftl.J' -t.n-t.nit~~~,hZ:.o t-.:c;Ur.. G. biora tDn:fLlillMf*1e~~jlHCLIM--L~--=~~tD3f~lIM'tD~~~ft~ff;-tt-.:clO, I ~1C L ~ltD YJJ~:l!:JtIJ tD 7Hffljf~t~a G. subglobosa (Brady) c G. crassa
rossensis Kennett I::-t.n-t.n MW.-t z:'o i t.:, Jll(;~iI;I1flUfii@l~ jf;-t C. porreetus c C.
parvus IiYJJ~JtlJI:: G. subglobosa 1::W.Z:.o LiI- L, .:.nl?tDtI!cMW.tI!cli~.f'lUli:c n:fLtD

1':1 rffl~'ljj\jtD ~H:: J: -:> L I~JJIJ-t Z:. .: c iI,liff~-C't; 1.10

502

8 *is'!E~~fr~~IJ.J}(7)mu~ • •PALAEONTOLOGICAL SOCIETY OF JAPAN. SPECIAL PAPERS. NUMBER 29 ~ 1986::P!l:t:.
flffL~<. ~~~~~~»Lt~~~~~~~H~~~~~. ~~.~~~h~~.~~~~~L. ~~~.L~.~~. ~~2 .~m.gR~ ~ffl*~~• • • • • • • ~~ B*~~_.~~~~m.
~Jl~ (R~:ilHtBi~!I5) ~l:.$ Li6Iv-Cr~"'.

(1)~~_I:.~-t~~ilB:)(-c. 1't-:)(~*.5jljlliJl&l:'b ~h L"'I*J~~ t~. ~-~*llli 13 ~rl:.!it:lia~~~:)(~.~t.: t ~ (l9iRI::::/:/:f.:; '7 J. ~Bi::)(~::i:.) -ct .n,. 7ti!:I:t~*~rr~*.5jIj%I:'IlHlJ:'~'!1?ff£-t~:. c • • ~t.J.7'\-iJ-1? tl';H~iJ':;1[lli~.tL~J!i6~if.>~iI.t~I:'lt. ~~~tfi:I:.rt: r.~J:~c.t I) t7t:l:iJ'~ <~ .t" '.

f'lJAilUfiJj-I:.-It<ii L ~lElitt.d!fl{ I) ~~lli-C~ ~:tk1rnt:.
$i6i!tc c tl:.£,-rl*~~~L~m:lli L~r~"'. (ffli1r~J:I't;gj;pftLi-t).
(3) $iMfll'litEiEl3-C-tiJ'. (1.:~$~~gJj~L~. ( ) 1*J~l±~~"1'-:>~r~",.
(a) $i6~.Ef:;~; fiJj-jim~it':ltji~fl:fiJj-· 'llnli:jjj:~·. (*~~~ -Cif.>~:: eJ.
(b) =i=~~; ~lfijxggl3. (5flJ~R..~#~c-t~:: c).
(e) 1jIf~1*J~~~\'r. (800-1.200::f:f!il1l:. 5flJX~"iiJ).
(d) l*J~t.c: I? V:1:.BkXiJ,+7t~Wi*-Cif.> ~ :: c ~IDEgJj. (t'lIJlj~~'fl'l~. L -ct .t H.
(e) *x~Jm (AilUJ: I) J!i6JtIU t':ltl*ir.i-CI't-X~ -1 725rr~~~~~ttl!l:-t.:t': L. /~-1 -j]-iJ·:r..
Y - r *f;$:iJ-~.~c-t ~ .:: c); i t.:*x*/N~* (8 :f.ti!...1,..) 1:'1~~-t" ~.7tiJ'if.> ~ c ~ It.
(2)

I*J~.:)(. c t I:' +7t~Wi*~ftiilt Lt.:l*ir.i-C.

if.>~:: c.

~~H.t~~I*J~ (~Jtl:.~-t~/~-~:/r); ~1ZI·~~~"'~l!l:cAilIJJ:I)fiJj-~JtI!l:; ~J{IZIJl&(0~ttl!l:.
fmiJ-l?~f'lJAilU!'i!'It:;1[lli~J!i6~tj1W<.

(g)~.L~.~ ~r~H.
~~fm~~$~.

(4)

(5)

~~T.tfi:.

1lllilf..

(~~~~~IDEgJjct.c:~~Mi;'':lt~$i6N.V:~ir.im:lli~j;IJ 19R5::P 5 fJ 31 B (,lZ·~). !~ri!fltf.liil.~ ~~iJ'£'~I:.rt: r. v 7 '" Y - c ~§~~J:I*J~L. 19861j=.1 fJ ~~l'~~~-c'ilF~i'f-:g~J:. $i6:i,iI:'@]~~T~-C-t. t.:t!. L~~iitrit.:ltf~I:'. $i6~C~$.~3(~~. 1I.~~iJ·I?*~~:: ciJ'if.>~iJ-t L.tLi..ltlv.
J:~c (0 ~fmiJ-l?~f'lJAilU!'i!~1llli~J!i6..1,..iJ't.c:"'~~lt. 19861j=.I.t~X$~fIJITJj;I]iilt~ (rlj}f~iilt:JHurrJitmJj;l]~J) ~$~f"'t.: L i-t~-C. J:~c (2) ~"dlj:iJ'..1,..t.:~.tL n' ~lf.;~I:::'~.7,..~!f.~.tL

i-t.
(6)

(7)

~~XiJ'ft~t.c:iJ.t~t:'lt. ~,¥~~-c~iR:gf!k-C~ ~ tDtlF- < f'lJAilUt:. c I) iJ-iJ- ~ T~-C-t. :)(.~~furr
Jj;I]iilt~~$~lt • • ~iJ-l?rrt.c:'" (f7tl1j=.ltll fJ*I:'$~t'$j;IJ). ~~~~. ~M:t.c:c·i'f-:gf~f'lJAilUI:'cI) iJ-iJ·1J t-t. -t-~~~It:)($~ c ~*'g3l0:'.t IJ. ~~1j=.~t'\ (iitrf7u-clt 10 fJ 20 B) i -C1:,fJJt'liJ'~.llit.c:I1.tLli. tmWJ~~3(#iJ~*.il:.~.tL~:: cl:.t.c:-:>~"'i-t.
*.5jIj%~I*~lt~tt:f:'*r.. iitrl9iJ~$~c L~I'FiiltL~r~"'. 7f'gJj~)i1:'t*liii.~j4.~I:.rR''''~h-lt~r~",.

11ntiJ'iJ-iJ-~~-C. ~.5jljt.c:~~~~~ • .5j!JAilljltl'Fiilt-ltf. *AilIJ 25 .~=i=~I:.1W<Mi!§! L
i-t. ~.tLt.J.J:ltIlJlJA (@L~~I:'lt~HI) C",?:,cl:.t.c:'Ji-t. "'<"?iJ-~~ifB::t~.~~lfllH:.
-t~ C ~1':'lt. 1!!:~A.~~iJ-I?:t~7f- L~. f,$:~J:~7f'Ki1E-~t.c:"'.t? I:. L~r~"'. f'lJAilIJJ:~m7f-$~iJ'~A -C~ ~ .t ? 1*ir.i~ti::tijilff{l!U· J:"'fl:. +7t~ B~ c -:> ~. ~ -1 7'-Cr-fttli= L ~r ~"'.

O::i:.~lliJl&llhiilt~iJ'~ l?.tLtd- -:> ;'.::t;!;.g. I':::' It. lliJl&Hi}j£"~:' c iJ'if.> ~ iJ- t L.tL i ~ Iv.
O*7t,Ql(;~I*~:tJj;.~~~c Itt.c: IJ i -ltA.. 7t~I*jj/lj~I'Fiilt~J:$i6Iv-Cr~ "'.

503

SYSTEMATIC INDEX(new Series No. 129-No. 136)

PALAEOZOOLOGY

Protozoa
Page
Furutani, Hiroshi: Middle Palaeozoic Palaeoscenidiidae (Radiolaria) from Mt. Yokokura,
Shikoku, Japan. Part I. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Ibaraki, Masako: Middle-Late Eocene planktonic foraminiferal faunas from limestones of
the Setogawa Group, central Japan . . . . . . . . . . • . . • . . . . . . . . . . . . . . . . • . . . 401
Ishiga, Hiroaki: Morphological change in the Permian Radiolaria, Pseudoalbaillella scalp rata
in Japan . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , 1
Mizutani, Shinjiro and Kido, Satoshi: Radiolarians in Middle Jurassic siliceous shale from
Kamiaso, Gifu Prefecture, central Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
Nomura, Ritsuo: Cassidulinidae (Foraminiderida) from the eastern part of Liitzow-Holm
Bay, Antarctica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 492
Sada, Kimiyoshi, Nomura, Kazuyoshi and Oho, Yukimasa: Primitive Fusulinacea from
Dangyokei of Taishaku . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 388
Sashida, Katsuo: Lower Triassic Radiolaria from the Kanto Mountains, central Japan
Part 1: Palaeoscenidiidae " . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • 168
Coelenterata
Omura, Akio: Uranium-series ages of some solitary corals from the Riukiu Limestone on
the Kikai-jima, Ryukyu Islands . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . • . . . . 117
Omura, Akio: Oxygen and carbon isotopic composition in the skeleton of an ahermatypicscleractinian coral, Dendrophyllia japonica Rehberg . . • . . . . . . . . . . . . . . . . . . . . 159
Omura, Akio: Uranium-series age of the Riukiu Limestone on Hateruma island, southwestern Ryukuys . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . 415
Brachiopoda
Tazawa, Jun-ichi: Early Carboniferous (Visean) brachiopods from the Hikoroichi Formation of the Kitakami Mountains, northeast Japan . . . . . . . . . . . . . . . . . . . . . . . . . 300
Tazawa, Jun-ichi, Gunji, Yukio and Mori, Kei: A Visean brachiopod fauna from the Mano
Formation, Soma district, Abukuma mountains, northeast Japan . . . . . . . . . . . . . . . 347
Mollusca
Goto, Michiharu: Some bivalves from the Lower Jurassic Kuruma Group of central Japan . 79
Hanai, Tetsuro: A story of the belemnite - an approach based on the method of inventing auxiliary assumptions in paleontology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Majima, Ryuichi: Observations on occurrence of Japanese Neogene naticids (Gastropoda)
bearing calcareous opercula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361

504
Matsumoto, Tatsuro: Cretaceous nautiloids from Hokkaido - I . . . . . . . . . . . . . . . . . "9Matsumoto, Tatsuro and Miyauchi, Toshiya: Cretaceous nautiloids from Hokkaido - III
Part 3. Some nautiloids from the Campanian of Soya, northern Hokkaido . . . . . . . . . 220
Matsumoto, Tatsuro, Miyauchi, Toshiya, Kanie, Yasumitsu, Miyata, Yuichiro and Ueda,
Yoshiro: Cretaceous nautiloids from Hokkaido - V . . . . . . . . . . . . . . . . . . . . . . . 335
Matsumoto, Tatsuro and Muramoto, Kikuwo:Cretaceous nautiloids from Hokkaido - II
Part 2. Three nautiloid species from the Santonian and Campanian of Hokkaido . . . .. 85
Matsumoto, Tatsuro, Takahashi, Takemi, Obata, Ikuwo and Futakami, Masao: Cretaceous
nautiloids from Hokkaido - IV Part 4. An interesting nautiloid species from the
Cenomanian of Hokkaido . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 288
Noda, Masayuki: Notes on the so-called Inoceramus japonicus (Bivalvia) from the Upper
Cretaceous of Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . 191
Noda, Masayuki: Notes on Mytiloides incertus (Cretaceous Bivalvia) from the Upper
Turonian of the Pombets area, central Hokkaido . . . . . . . . . . . . . . . . . . . . . . . . . 455
Tabuki, Tyoichi: Paleoenvironment of the Plio-Pleistocene Daishaka Formation, Tsugaru
basin, northeast Japan . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . • . . .. 61
Yang, Seong Young: On the subgenus Wakinoa (Cretaceous non-marine Bivalvia) from
Gyeongsang Group, Korea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177

Arthropoda
Hu, Chung-Hung: Ontogenic and phylogenic development of two Upper cambrian trilobitesfrom the Nolichuky Formation, Tennessee . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Hu, Chung-Hung: Ontogenesis of Ehmaniella burgessensis Rasetti (Trilobita) from Bur"ess
shale, Middle Cambrian, Yoho Park, British Columbia . . . . . . . . . . . . . . . . . . . . . . 394
Ishizaki, Kunihiro: Ostracoda from the Pliocene Ananai Formation, Shikoku, Japan Description - . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. .. 135
Kaneko, Atsushi: A Middle Devonian trilobite fauna from the Kitakami Mountains,Northeast Japan I. The Lichidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474

Echinodermata
Tanaka, Keisaku: Hemiasterid echinoids from the Upper Cretaceous of Japan. . . . . . . . 427
Tanaka, Keisaku, Noda, Masayuki and Tanaka, Hitoshi: Echinoids from the Cretaceous
Haidateyama Group, eastern Kyushu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445
PALAEOBOTANY.
Fuji, Norio: Palynological study of 200-meter core samples from Lake Biwa, central
Japan I: The palaeovegetational and palaeoclimatic changes during the last 600,000
years . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
Fuji, Norio: Palynological investigation of the postglacial deposits in Lagoon Kahokugata, Kanazawa, central Japan . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
Kimura, Tatsuaki and Tsujii, Masanori: Early Jurassic plants in Japan. Part 5 . . . . • . . . .. 35
Kimura, Tatsuaki and Tsujii, Masanori: Early Jurassic plants in Japan, Part 6 . . . . . . . . . . 265
Matsuoka, Kazumi: Some dinoflagellate cysts from the Nanggulan Formation in central
Java, Indonesia . . . . . . . . . . . • . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . 374

505

INDEX OF GENERA AND SPECIES
Notes: Words listed are names of genus- and species-groups, which are described, discussed
or illustrated in the articles in Transactions and Proceedings of the Palaeontological Society ofJapan, New Series, Nos. 129-136. Words in bold-faced type are names of newly proposed taxa.

A

Abies . . . . . . . . . . . . . . . . . . . 238,319
Abies Mariesii . . . . . . . . . . . . . . . . . 238
Abies Veichii . . . . . . . . . . . . . . . . . . 238
Abies firma . . . . . . . . . . . . . . . . . . . 238
Abies homolepis . . . . . . . . . . . . . . . . 238
Abies sachalinensis . . . . . . . . . . . . . . 238
Abies sachalinensis uar. mayriana ..... 238
Acabthastrea echinata . . . . . . . . . . . . 120
Acanthocircus hexagonus . . . . . . . . . . 255
Acanthocircus suboblongus . . . . . . . . 255
Acanthopyge . . . . . . . . . . . . . . . . . . 481
Acanthopyge balliuianii . . . . . . . . . . . 487
Acanthopyge consanguinea . . . . . . . . . 479
Acanthopyge contusa . . . . . . . . . . . . 487
Acanthopyge aff. contusa . . . . . . . . . . 487
Acanthopyge latelobe . . . . . . . . . . . . 487
Acanthopyge leuchtenbergii . . . . . . . . 486
Acanthopyge (A can thopyge ) duplicispinota . . . . . . . . . . . . . 481, pis. 88, 89
Acanthopyge (Lobopyge) . .......... 487
Acanthopyge (Lobopyge?) . . . . . . . . 487
Acanthopyge (Lobopyge) sinuata .... 486
Acanthopyge (Mephiarges) . . . . . . . . . 484
Acanthopyge (Mephiarges) bifida ..... 478
Acanthopyge (s.l.) . . . . . . . . . . . . . . 476
Acarinina . . . . . . . . . . . . . . . . . . . . 405
Acarinina punctocarinata ..... 405, pI. 77
Acarinia rotundimarginata . . . . . . . . . 407
Acesta goliath . . . . . . . . . . . . . . . . . 71
Acila nakazimai . . . . . . . . . . . . . . .. 71
Adamussium colbecki . . . . . . . . . . . . 493
Albaillella sp. . . . . . . . . . . . . . . . . . .. 5
Alnus . . . . . . . . . . . . . . . . . . . 239,325
Alocopocythere goujoni . . . . . . . . . . . 139
Amaltheus sp. . . . . . . . . . . . . . . . . . 79
Amphilichas minganensis . . . . . . . . . . 478
Anapachydiscus deccanensis yezoensis. . . . . . . . . . . . . . . . . . . . . . . . . . . 92

(A.) uandenheckii . . . . . . . . . . . . . . 447
A nglonautilus . . . . . . . . . . . 89,226,289
Anglonautilus japonicus . . . . . . . . . . .
. . . . . . . . . . . . . . . 289, 341, pis. 57-60
Anglonautilus mamiyai . . . . . . . . . . .
. . . . . . . . . . . . . . 226,292, 341, pI. 50
Anglonautilus undulatus . . . . . . 226, 291
Angulithes . . . . . . . . . . . . .. 12, 87, 343
Angulithes sowerbianus . . . . . . . . . .. 15
Angulithes triangularis . . . . . . . . . . 14,88
Anticleina antarctica . . . . . . . . . . . . . 493
Aphelaspis . . . . . . . . . . . . . . . . . . .. 26
Aphelaspis subditus . . . . . . . . . . . . .. 32
Aphelaspis tarda . . . . . . . . . . . . 27, pI. 10
Aphelaspis tumifrons . . . . . . . . . 29, pI. 11
Aphelaspis walcotti . . . . . . . . . . . . .. 32
Aphelaspis sp. . . . . . . . . . . . . . . . . . 31
Apocalymene . . . . . . . . . . . . . . . . . . 476
Arca boucardi . . . . . . . . . . . . . . . . . 75
Archaeosemantis . . . . . . . . . . . 102, 171
Archaeosemantis cristianensis . . . . . . . 171
Archaeosemantis longiuirga . . . . . . . . . 171
Archaeosemantis uenusta ..... 171, pI. 36
Archaeosemantis sp . . . . . . . . . 171, pI. 36
Archaeospongoprunum? . . . . . . . . . . 103
Archaeothamnulus . . . . . . . . . . . . . . 173
Archaeothamnulus ramosus ... 173, pI. 37
Archaeothamnulus uerticillatus . . . . . . 173
Archaeothamnulus? sp . . . . . . . 173, pI. 36
Arges . . . . . . . . . . . . . . . . . . . . . . . 477
Arges consanguineus . . . . . . . . . . . . . 485
Astarte borealis . . . . . . . . . . . . . . . . 71
Astroentactinia ..... , . . . . . . 103,110
Atrypa . . . . . . . . . . . . . . . . . . . . . . 476
Atrypa (Planuatrypa) . . . . . . . . . . . . 476
Atrypa (Planuatrypa)squamifera ..... 476
Atrypa desquamata mut.japonica .... 476
Atrypa pauciplicata . . . . . . . . . . . . . . 476

506

B
Baculites cf. yokoyamcii . . . . . . . . . . . 470
Baiera? sp. . . . . . . . . . . . . . . . . . . . . 266
Bairdppilata . . . . . . . . . . . . . . . . . . . 141
Bairdoppilata alcyonicola . . . . . . . . . . 142
Bairdoppilata sp. . . . . . . . 141, pIs. 33, 34
Bakevellia . . . . . . . . . . . . . . . . . . .. 83
Bakevellia (Neobakevellia) sp . . . . 83, pI. 15
Basslerites taiwanensis . . . . . . . . . . . . 137
Bathypsammia tintinnabulum . . . . . . . 160
Baticasphaera . . . . . . . . . . . . .. 376, 387
Baticasphaera explanata ...... 376, pI. 74
Baticasphaera hirsuta . . . . . . . . . . . . . 377
Baticasphaera micropapillata ... 377, pI. 74
Betula ... , . . . . . . . . . . . . . . 239, 325
Betula ermani . . . . . . . . . . . . . . . . . 239
Betula platyphylla . . . . . . . . . . . . . . 239
Bisiphytes . . . . . . . . . . . . . . . . . . .. 15
Bissylentactinia . . . . . . . . . . . . . . . . 103
Bissylentactinia quadriramosum ..... 104
Bissylentactinia rudicula ...... 103, 172
Blountia nixonensis . . . . . . . . . . . . .. 32
Bolbaster . . . . . . . . . . . . . . . . . . . . 441
Brachythyris . . . . . . . . . . . . . . . . . . 353
Brachythyris kitakamiensis . . . . . . . . . 354
Brachythyris pseudovalis . . . . . . . . . . 354
Brachythyris sp . . . . . . . . . . . . 353, pI. 67
Bradleya albatrossia . . . . . . . . . . . . . 139
Bron~~ . . . . . . . . . . . . . . . . . . . . . 475
Bronteus (Thysanopeltis) paucispinosa. . . . . . . . . . . . . . . . . . . . . . . . . . 474

Buntonia . . . . . . . . . . . . . . . . . 135,148
Buntonia hanaii . . . . . . . . . . . . . . . . 149
Buntonia parascorta ..... 148, pis. 28, 35
Buntonia scorta ..... 149, pis. 28, 29, 35
Buntonia u-carinata ...... 151, pis. 29,35
Bythoceratina dipleura . . . . . . . . . . . 137
C

Calceola . . . . . . . . . . . . . . . . . . . . . 475
Caligodinium . . . . . . . . . . . . . . . . . . 377
Caligodinium amiculum . . . . . . . . . . . 377
Caligodinium granulatum . . . . . . . . . . 378
Caligodinium sp. cf. C. amiculum .... .
. . . . . . . . . . . . . . . . . . . . . 377,pI.74

Callistocythere . . . . . . . . . . . . . . . . . 143
Callistocythere alata . . . . . . . . . . . . . 146
Callistocythere ananaiensis . . . . . . . . .
. . . . . . . . . . . . . . . 143, pis. 30, 31, 35

Callistocythere dorsotuberculata ..... 145

Callistocythere kattoi .... 144, pis. 31,35
Callistocythere ovata . . . . . . . . . . . . . 137
Callistocythere puri . . . . . . . . . . . . . . 144
Callistocythere undata . . . . . . . . . . . . 146
Callistocythere sp. . . . . . . . . . 145, pI. 32
Cameraspis convexa . . . . . . . . . . . . . 33
Canavaria sp. . . . . . . . . . . . . . . . . .. 79
Canoptum (?) sp. A . . . . . . . . . . . . . . 258
Carinautilus . . . . . . . . . . . . . . . . . . . 342
Carinonautilus . . . . . . . . . . . . . . . .. 17
Carinonautaus ariyalurensis . . . . . . . . 17
Carpinus . . . . . . . . . . . . . . . . . . . . . 239
Cassidulina biora . . . . . . . . . . . . . . . 493
Cassidulina crassa . . . . . . . . . . . . . . . 493
Cassidulina crassa var. porrecta ...... 495
Cassidulina sp. . . . . . . . . . . . . . . . . . 493
Cassidulinoides . . . . . . . . . . . . . . . . . 495
Cassidulinoides parvus .... 498, pis. 90, 92
Cassidulinoides porrecta . . . . . . . . . . . 497
Cassidulinoides porrectus .. 495, pis. 90, 92
Catapsydrax dissimilis . . . . . . . . . . . . 408
Cenoceras . . . . . . . . . . . 15,88,223,341
Cenoceras trechmanni . . . . . . . . . . .. 15
Ceratarges . . . . . . . . . . . . . . . . . . . . 477
Ceratoikiscum sp. . . . . . . . . . . . . . .. 97
Ceratoikiscum spp. . . . . . . . . . . . . .. 98
Chamaecyparis . . . . . . . . . . . . . . . . . 244
Chascothyris . . . . . . . . . . . . . . . . . . 475
Chiloguembelina . . . . . . . . . . . . . . . . 411
Chiloguembelina cubensis . .... 411, pI. 77
Chlamys cosibensis . . . . . . . . . . . . .. 71
Chlamys jousseaumei . . . . . . . . . . . . . 161
Chlamys swiftii . . . . . . . . . . . . . . . . 74
Choristites . . . . . . . . . . . . . . . . . . . . 307
Choristites bisulcatus. . . . . . . . 308, pI. 61
Choristites mosquensis . . . . . . . . . . . . 308
Chytroeisphaeridea . . . . . . . . . . . . . . 377
Cimomia . . . . . . . . . . . . . .. 12,89,341
Cimomia burtini . . . . . . . . . . . . . .
13
Cimomia schloderi .... . . . . . . . . .. 13
Cimomia vaughani . . . . . . . . . . . . . . 13
Cimomia wylliei . . . . . . . . . . . . . . .. 13
Cladoceramus . . . . . . . . . . . . . . . . . 202
Cladophlebis . . . . . . . . . . . . . . . . . , 79
Cleiothyridina . . . . . . . . . . . . . . . . . 307
Cleiothyridina fimbriata . . . . . . . . . . . 307
Cleiothyridina sp . . . . . . . . . . . 307, pI. 61
Clisiphyllum awa . . . . . . . . . . . . . . . 389
Clisiphyllum ofukuensis . . . . . . . . . . . 389
Conchyliolith~ (Anomites) resupinatus

507. . . . . . . . . . . . . . . . . . . . . . . . . . 304

Coniopteris . . . . . . . . . . . . . . . . . . . 284
Coniopteris hymenophylloides ...... 284
Coniopteris sp. A . . . . . . . . . . . . . . . 284
Coosella convexa . . . . . . . . . . . . . . . 32
Coosia albertensis . . . . . . . . . . . . . .. 32
Cordosphaeridium uncinispinosum . ... 382
Cory Ius . . . . . . . . . . . . . . . . . . . . . . 321
Craspedarges . . . . . . . . . . . . . . . . . . 487
Crassifimbra walcotti . . . . . . . . . . . . . 395
Crenella columbiana . . . . . . . . . . . .. 75
Crenotrapezium . . . . . . . . . . . . . . .. 79
Crepicephalus deadwoodensis . . . . . .. 32
Crioceratites (Crioceratites) sp. aff. C.
(C. ) koechlini . . . . . . . . . . . ...... 447
Cryptomeria . . . . . . . . . . . . .. 238,319
Cryptomeria japonica . . . . . . . . . . . . 238
Cryptonatica adamsiana ...... 365, pI. 69
Cryptonatica clausa . ..... 363, pIs. 69, 70
Cryptonatica ichishiana ... 365, pIs. 69, 70
Cryptonatica janthostoma . . . . . . . . .
· . . . . . . . . . . . . . . . . . 362, pIs. 68, 70

Cryptonatica janthostomoides . ... 71,362
Cryptonatica sp. 1 . . . . . . . . . . . . . . 362
Ctenis. . . . . . . . . . . . . . . . . . . . . .. 35
Ctenis japonica. . . .
. . . . . . . . .. 36
Ctenis kaneharai . . . . . . . . . 35, pIs. 12, 13
Ctenis nipponica. . . . . . . . . . . . . . .. 36
Ctenis sp. . . . . . . . . . . . . . . . . . . . . 36
Cyathaxonia sp. . . . . . . . . . . . . . . . . 349
Cycadites . . . . . . . . . . . . . . . . . . .. 47
Cycadites manchurensis . . . . . . . . . .. 47
Cycadites saladini . . . . . . . . . . . . . .. 47
Cycadites cfr. saladini . . . . . . . . 47, pI. 12
Cycadocarpidium nagatoense . . . . . . . 280
Cyclobalanopsis . . . . . . . . . . .. 239, 325
Cymatoceras . . . . . . . . . . . . 90, 225, 336
Cymatoceras atlas. . . . . . . . . . . . 88, 228
Cymatoceras carlottense . . . . . . . . . . 337
Cymatoceras cenomanense . . . . . . . . . 225
Cymatoceras elegantoides . . . . . . . . . . 339
Cymatoceras honmai . . . . . . . . . . . . .
· . . . . . . . . . . . .. 225,341, pIs. 48, 49

Cymatoceras loeblichi . . . . . . . . . . .. 89
Cymatoceras madagascarense ....... 94
Cymatoceras mikado . . . . . . . . 339, pI. 65
Cymatoceras sp. cf. C. mikado ...... 339
Cymatoceras pacificum . . . . . . . . . . .
· . . . . . . . . . . . . . . . 92,341, pIs. 18-20

Cymatoceras patens . . . . . . . . . . . . . 88
Cymatoceras perstriatus . . . . . . . . . .. 90

Cymatoceras pseudoatlas .. 90,341, pI. 18
Cymatoceras pseudoneganum . . . . . .. 88
Cymatoceras pseudoneokomiense .... 338
Cymatoceras sakalavum . . . . . . . . . . . 337
Cymatoceras sp. cf. C. sakalavum .... .
. . . . . . . . . . . . . . . . . . . . . 337, pI. 64

Cymatoceras saussureanum. . . . . . . .. 88
Cymatoceras semilobatum . . . . . . . .. 88
Cymatoceras senomanense . . . . . . . .. 88
Cymatoceras sharpei . . . .. 93,340, pI. 66
Cymatoceras tourtiae . . . . . . . . . . . . 88
Cymatoceras tskaltsithelensis ..... .. 90
Cymatoceras virgatum . . . . . . . . . . . . 337
Cymatoceras sp. aff. C. virga tum .... .
. . . . . . . . . . . . . . . . .. 336, pIs. 62-64

Cymatonautilus . . . . . . . . . . . . . . .. 89
Cyphastrea sp . . . . . . . . . . . . . . . . . . 120
Cyrtia laminosa . . . . . . . . . . . . . . . . 355
Cyrtocapsa mastoidea . . . . . . . . . . . . 255
Cytherella . . . . . . . . . . . . . . . . . . . . 140
Cytherella japonica . . . . . . . . . 140, pI. 32
Cytherella sp . . . . . . . . . . . . . . . . . . . 141
Cytherelloidea sabahensis . . . . . . . . . . 139
Cytheropteron . . . . . . . . . . . . . . . . . 135
Cytheropteron rhombea . . . . . . . . . . . 137
Czekanowskia . . . . . . . . . . . . . . . . . 272
Czekanowskia blacki . . . . . . . . . . . . . 273
Czekanowskia furcata . . . . . . . . . . . . 273
Czekanowskia hartzi . . . . . . . . . . . . . 273
Czekanowskia rigida . . . . . . . . . . . . . 272
Czekanowskia ex gr. rigida .... 272, pI. 56
Czekanowskia setacea . . . . . . . . . ... 273
D

Daena . . . . . . . . . . . . . . . . . . . . . . . 374
Daena? sp . . . . . . . . . . . . . . . . . . . . . 374
Dechenella . . . . . . . . . . . . . ...... 475
Dechenella (Dechenella) . . . . . . . . . . . 475
Dechenella (Dechenella) minima ..... 475
Dechenella minima . . . . . . . . . . . . . . 475
Delthyris aff. D. clarksvillensis ...... 355
Deltocymatoceras . . . . . . . . . . . . . .. 89
Deltoidonautilus. . . . . . . . . . . . . . .. 14
Dendrophyllia japonica . . . . . . . . . . . 160
Desmoceras kossmati . . . . . . . . . . . . . 337
Dibunophyllum cf. D. biparticum .... 349
Diceratigalea . . . . . . . . . . . . . . . . . . 105
Diceratigalea hemisphaera . . . . . . . . . 105
Dictiomitrella (?) kamoensis ... 258, pI. 53
Dictyomitrella . . . . . . . . . . . . . . . . . 258
Dictyomitrella sp. A . . . . . . . . . . . . . 253

508

Dictyomitrella (?) kamoensis ....... 258
Dictyophyllum . . . . . . . . . . . . . . . .. 79
Didymotis akamatsui . . . . . . . . . . . . . 470
Diphyes . . . . . . . . . . . . . . . . . . . . . 379
Diphyes colligerum . . . . . . . . . . . . . . 380
Diphyes spinulum . . . . . . . . . . 379 pI. 73
Discocyclina . . . . . . . . . . . . . . . . . . 376
Discocyclina sp . . . . . . . . . . . . . . . . . 404
Dora to cy there taiwanensis . . . . . . . . . 137
Doratophyllum decoratum . . . . . . . .. 50
Dunderbergia . . . . . . . . . . . . . . . . .. 26
Dunderbergia anyta. . . . . . . . . . . . .. 33
Duvalia . . . . . . . . . . . . . . . . . . . . . . 129
Dytremacephalus granulosa ........ 33
E

Ehmaniella . . . . . . . . . . . . . . . . . . . 396
Ehmaniella burgessensis ...... 396, pI. 76
Ehmaniella sp . . . . . . . . . . . . . . . . . . 395
Ehrenbergina . . . . . . . . . . . . . . . . . . 499
Ehrenbergina glabra ..... 499, pis. 90, 92
Ehrenbergina hystrix . . . . . . . . . . . . . 500
Ehrenbergina hystrix var. glabra ..... 499
Ehrenbergina parva . . . . . . . . . . . . . . 498
Ehrenbergina pusa . . . . . . . . . . . . . . 500
Elatocladus . . . . . . . . . . . . . . . . . . . 275
Elatocladus sp. A . . . . . . . . . . 275, pI. 56
Elatocladus sp. B . . . . . . . . . . . . . . . 275
Elephas shigensis . . . . . . . . . . . . . . . 235
Elrathia permulta . . . . . . . . . . . . . . . 396
Elvinia . . . . . . . . . . . . . . . . . . . . .. 26
Elvinia roemeri. . . . . . . . . . . . . . . .. 33
Enallaster . . . . . . . . . . . . . . . . . . . . 450
Endothyra . . . . . . . . . . . . . . . . . . . . 390
Entactinia cometes . . . . . . . . . . . . . . 102
Entactinia diversita . . . . . . . . . . . . . . 102
Entactinia monalloea . . . . . . . . . . . . . 102
Entactinia? additiva . . . . . . . . . . . . . 102"aff. Entactinia sp." . . . . . . . . . . . . . 103
Entactinosphaera euthlasta . . . . . . . . . 102
Entactinosphaera fredericki ........ 101
Entactinosphaera palimbola ........ 102
Entactinosphaera riedeli . . . . . . . . . . . 102
Entactinosphaera unimana . . . . . . . . . 101
Entactinosphaera? diplostraca ....... 102
Eomiodon . . . . . . . . . . . . . . . . . . .. 79
Eostaffella kanmerai ........ 390, pI. 75
Epicymatoceras . . . . . . . . . . . . . . .. 89
Epigondolella postera . . . . . . . . . . . . 169
Etremophyllum . . . . . . . . . . . . . . . . 270
Etremophyllum glandulosum ....... 270

Etremophyllum tetoriense . . . . . . . . . 270
Eucymatoceras. . . . . . . . . . . . . . . .. 88
Eurocymatoceras . . . . . . . . . . . . . . , 89
Eutrephoceras . . . . . . . . . . . 12, 223, 342
Eutrephoceras ahltensense . . . . . . . .. 16
Eutrephoceras balchistanense .... 16,224
Eutrephoceras darupense . . . . . . . . .. 87
Eutrephoceras dekayi . . . . . . . . . . .. 87
Eutrephoceras indicum . . . . . . . . . . . 22
"Eutrephoceras" japonicum . . . . . . .. 90
Eutrephoceras kobayashii . ... 87, 223, 341
Eutrephoceras montanense . . . . . . . .. 15
Eutrephoceras nodai . . . . . . . . . . 22, pI. 9
Eutrephoceras pseudobouchardianus .. 225
Eutrephoceras sloani . . . . . . . . . . . .. 87
Eutrephoceras soyaense . . . . . . . . . . .
. . . . . . . . . . . . . . 223,341, pis. 47, 48

Eutrephoceras ef. soyaense . . . . . . . . . 225
Eutrephoceras sublaevigatum ...... .
. . . . . . . . . . . . . . . . . . . . 16,87,224

Eutrephoceras tawaense . . . . . . . . . . . 223
Eutrephoceras sp. . . . . . . . . . . . . . .. 87
Exochosphaeridium bifidum . . . . . . . . 381
Exochosphaeridium brevispinosum .. .
. . . . . . . . . . . . . . . . . . . . . 381,pI.72

Exochosphaeridium reticulatum .....
. . . . . . . . . . . . . . . . . . . . . 380, pI. 72

Ezocallista brevisiphonata . . . . . . . . . 71

F
Fagus . . . . . . . . . . . . . . . . . . . . . . . 239
Fagus . . . . . . . . . . . . . . . . . . . . . . . 325
Fc;gus crenata . . . . . . . . . . . . . . . . . . 239
Fagus crenata . . . . . . . . . . . . . . . . . . 319
Fagus japoniea . . . . . . . . . . . . . . . . . 239"Favosites" . . . . . . . . . . . . . . . . . . . 4 7 5Flabellum rubrum . . . . . . . . . . . . . . . 118
Follicucullus . . . . . . . . . . . . . . . . . .. 5
Follicucullus monacanthus . . . . . . . . .. 5G

Genevievella plesiochielus. . . . . . . . .. 32
Gikgo lepida . . . . . . . . . . . . ...... 266
Ginkgo . . . . . . . . . . . . . . . . . . . . . . 265
Ginkgo concinna . . . . . . . . . . . . . . . 267
Ginkgo digitata var. huttoni . . . . . . . . 267
Ginkgo flabellata . . . . . . . . . . . . . . . 267
Ginkgo efr. lepida . . . . . . . . . . . . . . . 267
Ginkgo pusilla . . . . . . . . . . . . . . . . . 267
Ginkgo schmidtiana . . . . . . . . . . . . . 267
Ginkgo sibirica . . . . . . . . . . . . . . . . . 266

509,Ginkgo ex gr. sibirica . .... 265, pIs. 54, 56
Ginkgoidium . . . . . . . . . . . . . . . . . . 271
Ginkgoidium (?) longilobatum . . . . . . 270
Ginkgoidium nathorsti . . . . . . . . . . . . 270
Ginkgoidium? sp. A . . . . . . . . . . . . . 271
Ginkgoites digitata . . . . . . . . . . . . . . 265
Ginkgoites sibirica . . . . . . . . , ..... 267
Ginkgoites? sp. " . . . . . . . . . . . . . . . . 265
Glaphyrocysta . . . . . . . . . . . . . . . ' .. 378
Glaphylocys.ta circularis . . . . . . 378, pI. 71
Glaphylocysta dentata . . . . . . . 378, pI. 71
Glaphylocysta intricata .... : . . . . . . 378
Glaphylocysta retiintexta . . . . . . . . . . 378?Glaphylocysta reticulosa . . . . . . . . . . 378
Globanomariina micra . . . . . . . . . . . . 409
Globigerapsis index . . . . . . . . . . . . . . 408
Globigerina . . . . . . . . . . . . . . . . . . . 407
Globigerina boweri . . . . . . . . . . . . . . 409
Globigerina collactea . . . . . . . . . . . . . 407
Globigerina frontosa . , . . . . . . . . . . . 409
Globigerina inaequispira ' . . . . . . . . . . 410
Globigerina linap~rta ..... '. . . . . . . . 410
Globigerina (Subbotina) linaperta .... 410
Globigerina officialis . . . . . . . . 407, pI. 77
Globigerina pera . . . . . . . . . . . 407, pI. 78
Globigerina praebulloides . . . . . . . . . . 407
Globigerina praebulloides praebulloides. . . . . . . . . . . . . . . . . . . . . 407, pI. 77
Globigerina pseudoeocaena . , . . . . . . . 411
Globigerina pseudoeocaj?na var. pseudoeocaena . . . . . . . . . . . . . . . . . . . . 411
Globigerina senni . . . . . . . . . . . . ... 408
Globigerinatheka . . . . . . . . . . . . . . . 408
Globigerinatheka index index . , 408, pI. 78
Globigerinatheka (Globigerapsis) index
index . . . . . . . . . . . . . . . . . . . . . . 408
Globigerinella micra . . . . . . . . . , ... 409
Globigerinita . . . . . . . . . . . . . . . . . . 408
Globigerinita globiformis ..... 408, pI. 78
Globigerinoides index . . . , , . . . . . . . 408
Globocassidulina . . . . . . . . . . . . . . ' 493
Globocassidulina biora ... , 493, pIs. 90, 91
Globocassidulina cf. biora .. ', ..... 493
Globocassidulina crassa rossensis ..... 493
Globocassidulina sp. . . . . . . . . . . . . . 493
Globorotalia collactea . . . . . . . . . . . . 407
Globorotalia tosaensis .... , .... , , . 135
Glossaulax didyma . . . . . . , ..... , . 369
Glycymeris yessoensis , . . . . . . . . . .. 71
Glyphaspis parkensis . , . . . . . . . . , . , 395
Glyphaspis cf. parkensis . . . . . . . . . , , 395

Glyphaspis cf. walcotti ... " .. " .. 397
Glyptostrobus , , . , . , , .. , .... , , . 238
Glyptostrobus pensilis ..... , ... , , . 238
Goniophoria angulata ..... , ... , .. 352
Grammoceras sp. . ... , . . . . . . " . , 79
Gravicalymene , .. , ... , , , ... , . , .476
Gravicalymene maloungkaensis ..... , 476
Gravicalymene yamakoshii . , , , ..... 476
Gravicalymene zhenzishanesis .. , .. , , 476
Guembelina cubensis . , . , , ..... , .. 411
H

Hantkenina , . , ..... , ... , .. , ... 374
Haplentactinia . . . . . . . . . . , .... " 96
Haplentactinia (s. 1.) sp, ... ,., ... , 97
Haplentactinia arrhinia . . . . . . . . ,., 101
Haplentactinia rhinophyusa ,.,. 101, 172
Hastigerina micra , ..... , , . . . . . . . 409
Helioentactinia, .... , . . . . . . . , ... 103
Heliolites ex, gr. porosus .. , , .. " ... 475
Hemiarges mikulici .. , , , .. , ... , .. 486
Hemiarges turneri ... , .. , .. , ..... 486
Hemiarges turneri rasetti . , , .. , .. , .478
Hemiaster , .. , . , ... , .. , .. , .. , . 428
Hemiaster indicus . . . . . . , ..... , .. 438
Hemiaster integer . . . . . . . . . . , .. , . 433
Hemiaster punctatus . , , , . , ... , ... 432
Hemiaster regulusanus . , .. , , ..... , 437
Hemiaster regulusi ... , . . . . . . . . ,. 437
Hemiaster stella ... , . , , . , . . . . . . . 430
Hemiaster tamulicus .... , , . . . . . . . 432
Hemiaster uwajimensis, , .. , . , . , , , . 428
Hemiaster (Hemiaster) frontacutus . ... 430
Hemiaster (Hemiaster) similaris . , , , , , 430
Hemiaster (Hemiaster) stella, ... , ... 432
Hemiaster (Hemiaster) uwajimensis . , .
· .. , '. , . , .. , .... , , . , , . 428, pI. 80
Hemiaster (Hemiaster) sp. B aff. H, (H,)
punctatus . , ... , ... , .... 432, pI. 80
Hemiaster (Hemiaster) sp. C .. , 433, pI. 80
Hemiaster (Hemiaster) sp. D ... 434, pI. 80
Hemiaster (Hemiaster) sp. aff. H (H.)
stella .. , . , . '. , . , , .. , ... 430, pI. 80
Hemiaster (Leymeriaster) polygonalis· , . , , , , . . . . . , , , , , ... , 435, pI. 81
Hemiaster (Leymeriaster) regulusanus . . 438
Hemiaster (Leymeriaster) sexangulatus· .. , , .... , . . . . . . . . . . . , , , , .437
Hemiaster (Leymeriaster) sp. aff. H (L,)
regulusanus ... , . , ... , .. , 437, pI. 81
Hemiaster (Mecaster) indicus .... , .. 439

510

Hemiaster (Mecaster) sp. aff. H. (M.) indicus . . . . . . . . . . . . . . 430, pIs. 80, 81
Hemicytherura cava .............. 139
Hemicytherura radiata . . . . . . . . . . . . 139
Heminautilus . . . . . . . . . . . . . . . . .. 89
Hercoglossa . . . . . . . . . . . . . . . . . .. 13
Hermanites posterocostatus ........ 147
Heteraster . . . . . . . . . . . . . . . . . . . . 447
Heteraster bohmi . . . . . . . . . . . . . . . 451
Heteraster bravoensis . . . . . . . . . . . . . 451
Heteraster bungoensis .... 448, pIs. 82, 83
Heteraster couloni . . . . . . . . . . . . . . 450
Heteraster lepidus . . . . . . . . . . . . . . . 450
Heteraster macroholcus . . . . . . 447, pI. 82
Heteraster nexilis . . . . . . . . . . . . . . . 449
Heteraster pomeli . . . . . . . . . . . . . . . 450
Heteraster yuasensis . . . . . . . . . . . . . 449
Heteraster sp . . . . . . . . . . . . . . 450, pI. 83
Hilarisirex . . . . . . . . . . . . . . . . . . . . 105
Hilarisirex quadrangularis . . . . . . . . . . 105
Hindeosphaera . . . . . . . . . . . . . . . . . 103
Hirsutocythere? . . . . . . . . . . . . . . . . 135
Homotryblium . . . . . . . . . . . . . . . . . 383
Homotryblium oceanicum ......... 383
Homotryblium pallidum ..... 383, pI. 74
Homotryblium plectilum ..... 383, pI. 74
Homotryblium valium . . . . . . . . . . . . 384
Homotryblium sp. . . . . . . . . . . . . . . 380
Homotryblium sp. cf. H. oceanicum .. .
· . . . . . . . ; . . . . . . . . . . . . 383, pI. 74?Homotryblium {loripes . . . . . . . . . . . 383
Hyphantoceras orientale . . . . . . . . . . 341
I

Inoceramus . . . . . . . . . . . . . . . . . . . 203
Inoceramus cycloides vanuxemiformis· . . . . . . . . . . . . . . . . . . . . . . . . . 202
Inoceramus cycloides var. undulata ... 215
Inoceramus digitatus . . . . . . . . . . . . . 202
Inoceramus diversus . . . . . . . . . . . . . 212
Inoceramus dresdensis labiatoidiformis· . . . . . . . . . . . . . . . . . . . . . . . . . 465
Inoceramus dresdensis? labiatoidiformis· .... ' . . . . . . . . . . . . . . . . . . . . . 469
Inoceramus ezoensis . . . . . . . . . . . . . 191
Inoceramus ezoensis vanuxemiformis .. 202
Inoceramus fiegei {iegei . . . . . . . . . . . 458
Inoceramus incertus . . . . . . . . . . . . . 458
Inoceramus incertus var. yubarensis .. .
· . . . . . . . . . . . . . . . . . . . . . 202,463
Inoceramus inconstans . . . . . . . . . . . . 214

Inoceramus inconstans inconstans .... 458
Inoceramus jciponicus . . . . . . . . . . . . 191
Inoceramus japonicus mammillatus ... 212
Inoceramus japonicus var. antsohaensis· . . . . . . . . . . . . . . . . . . . . . . . . . 212
Inoceramus labiatus . . . . . . . . . . . . . 455
Inoceramus latus . . . . . . . . . . . . . . . 455
Inoceramus mantelli .............. 202
Inoceramus michaeli . . . . . . . . . . . . . 202
Inoceramus naumanni . . . . . . . . . . . . 202
Inoceramus striatoconcentricus striatoconcentricus . . . . . . . . . . . . . . . . . 465
Inoceramus undulatoplicatus ....... 191
Inoceramus uwajimensis . ....... 21,470
Inoceramus sp . . . . . . . . . . . . . . . . . . 455
Inoceramus sp. cf. incertus . . . . . . . . . 458
Inoceramus (Cataceramus) japonicus .. 204
Inoceramus (Cladoceramus?) japonicus· . . . . . . . . . . . . . . . . . . . . . . . . . 205
Inoceramus (Mytiloides) mytiloides ... 466
Inoceramus (Mytiloides) opalensis .... 466
Inoceramus (Mytiloides) subhercynicus· . . . . . . . . . . . . . . . . . . . . . . . . . 466
Inoceramus (Mytiloides?) hercynicus .. 466
Inoceramus (Mytiloides?) "latus" .... 466
Inoceramus (Platyceramus) amakusensis· . . . . . . . . . . . . . . . . . . . . . . . . . 92
Inoceramus (Platyceramus) chouberti .. 465
Inoceramus (Platyceramus) cycloides
cycloides . . . . . . . . . . . . . . . . . . . . 215
Inoceramus (Platyceramus) cycloides
wegneri . . . . . . . . . . . . . . . . . . . . . 215
Inoceramus (Platyceramus) higoensis ..
· . . . . . . . . . . . . . . . .. 206, pIs. 44-46
Inoceramus (Platyceramus) japonicus .. 203
Inoceramus (Platyceramus) japonicus
hokkaidoensis . . . . . . .. 205, pIs. 41-43
Inoceramus (Platyceramus) japonicus
japonicus . . . . . . . . . . . . . . 203, pI. 41
Inoceramus (Platyceramus) rhomboides· . . . . . . . . . . . . . . . . . . . . . . . . . 215
Inoceramus (Platyceramus) rhomboides
rhomboides . . . . . . . . . . . . . . . . . . 215
Inoceramus (Platyceramus) rhomboides
heinei . . . . . . . . . . . . . . . . . . . . . . 215
Inoceramus (Platyceramus) yubarensis· . . . . . . . . . . . . . . . . . . . . . . . . . 213
Integraster . . . . . . . . . . . . . . . . . . . . 441
Irvingella . . . . . . . . . . . . . . . . . . . .. 26
Irvingella major . . . . . . . . . . . . . . .. 33

511

JJanishewskiella (?) angulata . . . . . . . . 352
Juglandsriesii . . . . . . . . . . . . . . . . . . 239
Jugosocythereis . . . . . . . . . . . . . . . . 147
Jugosocythereis? sp.
. . 147, pis. 30,35
K

Kallosphaeridium . . . . . . . . . . . . . . . 377
Kallosphaeridium brevibarbatum ..... 377
Kallosphaeridium sp. . . . . . . . . 377, pI. 74
Kangarina cava . . . . . . . . . . . . . . . . . 139
Karavankina . . . . . . . . . . . . . . . . . . 307
Karavankina jizodoensis . . . . . . . . . . . 307
Karavankina praepermica . . . . . . . . . . 307
Karavankina sp . . . . . . . . . . . . 306, pI. 61
Kingstonia ara . . . . . . . . . . . . . . . .. 32
Komaspidella laevis . . . . . . . . . . . . .. 32
Koreanaia . . . . . . . . . . . . . . . . . . .. 177
Koreanaia cheongi . . . . . . ~ . . . . . . . 178
Koreanaia yunnanensis . . . . . . . . . . . 178
Krithe . . . . . . . . . . . . . . . . . . . . . . 135
Krithe hanaii . . . . . . . . . . 142, pis. 33, 35
Krithe producta . . . . . . . . . . . . . . . . 139
Kuandiania crassicaulis . . . . . . . . . . . 36
Kumamotoa . . . . . . . . . . . . . . . . . . 177
Kumamotoa matsumotoi . . . . . . . . . . 178
Mumamotoa mifunensis . . . . . . . . . . . 178
Kumamotoa suzukii . . . . . . . . . . . . . 178
Kummeloceras . . . . . . . . 12, 85, 223, 341
Kummeloceras clementinum . . . . . . .. 16
Kumeloceras f/euriausianus. . . . . . . .. 15
Kummeloceras kamuy . . .. 86, 341, pI. 16
Kummeloceras splendens . . . . . . . . 14, 87
Kummeloceras turcicum . . . . . . . . . . 88
Kummeloceras yamashitai . . ...... .
. . . . . . . . . . . . . . . 18,87,341, pis. 4·7
Kummeloceras yezoense . . . . . . . . . .
. . . . . . . . . . . . . . 19,87,341, pis. 7,8
Kummelonautilus . . . . . . . . . . . . . . . 345
L

Larix . . . . . . . . .
. . . . . . . . . . . 321
Laternula elliptica . . . . . . . . . . . . . . . 493
Lepidobalanus . . . . . . . . . . . .. 239, 325
Lepidodendropsis sp. . . . . . . . . . . . .. 98
Leptaena analoga . . . . . . . . . . . . . . . 350
Leptaena cf. L. analoga . . . . . . . . . . . 350
Leptaena depressa var. analoga ...... 350
Leptaena rhomboidalis . . . . . . . . . . . 350
Leptaenella analoga . . . . . . . . . 350, pI. 67
Leptaenella cf. L. analoga . . . . . . . . . . 350

Leptagonia . . . . . . . . . . . . . . . . . . . 350
Leptagonia distorta . . . . . . . . . . . . . . 351
Leptagonia cf. L. analoga . . . . . . . . . . 351
Leptophloeum rhombicum . . . . . . . . , 98
Leymeriaster . . . . . . . . . . . . . . . . . . 435
Lichas devonianus . . . . . . . . . . . . . . . 486
Lichas grandis .... . . . . . . . . . . . . . 479
Lichas haueri . . ... .
. .. 486
Lichas laciniatus . . . . . . . . . . . . . . .. 478
Lichas (Arges) contusus . . . . . . . . . . . 485
Lichas (Euarges) mephisto . . . . . . . . . 484
Lichas (Euarges) parvulus var. convexa. . . . . . . . . . . . . . . . . . . . . . . . . . 487

Lichas (Euarges) planus . . . . . . . . . . . 487
Lichas (Euarges) sibiricus . . . . . . . . . . 486
Limopsis tokaiensis . . . . . . . . . . . . .. 71
Lindleycladus lanceolatus . . . . . . . . . . 278
Lingulosphaera spinula . . . . . . . . . . . 379
Lithocampe(?) nudata . . . . . . . . . . . . 256"Lithocannosphaeropsis fallax" . ..... 1 04
Lithostrotionella taishakuensis . . . . . . 389
Lobactinocapsa . . . . . . . . . . . . . . . . 101
Lobopyge limbata . . . . . . . . . . . . . . . 487
Lonsdaleia sp. . . . . . . . . . . . . . . . . . 349
Lymaniceras planulatum . . . . . . . . . . 470
MMacgeea . . . . . . .
. . . . . . . . . . . 475
Macoma calcarea . . . . . . . . . . . . . . . 71
Macoma sp. . . . . . . . . . . . . . . . . . .. 75
Macrocypris decora . . . . . . . . . . . . . . 139
Macrotaeniopteris jourdyi . . . . . . . . . 50
Makiyama chitanii . . . . . . . . . . . . . . 62
Mammites sp. . . . . . . . . . . . . . . . . .. 23
Marattia asiatica. . . . . . . . . . . .. 51, 283
Mecaster . . . . . . .
. . . . . . . . . . . 438
Megateuthis . . . . . . . . . . . . . . . . . . . 128
Melosira . . . . . . . . . . . . . . . . . . . . . 248
Melosira solida . . . . . . . . . . . . . . . . . 248
Metasequoia . . . . . . . . . . . . . . . . . . 238
Micrabacia japonica . . . . . . . . . . . . . . 118
Millerella kanmerai . . . . . . . . . . . . . . 390 .
Millerella sp. . . . . . . . . . . . . . . . . . . 389
Mirifusus baileyi . . . . . . . . . . . . . . . . 253
Mizuhopecten . . . . . . . . . . . . . . . . . 71
Mizuhopecten yessoensis yokoyamae .. 71
Monia macroschisma . . . . . . . . . . . .. 71
Monoceratina dipleura . . . . . . . . . . . . 137
Montastrea annularis . . . . . . . . . . . . . 159
Montipora verrucosa . . . . . . . . . . . . . 159
Muricoglobigerina . . . . . . . . . . . . . . . 408

512

Muricoglobigerina senni ...... 408, pI. 78
Mutilus nodulosum . . . . . . . . . . . . . . 139
Myrica . . . . . . . . . . . . . . . . . . . . . . 325
Mytiloides . . . . . . . . . . . . . . . . . . . . 458
Myti/oides aviculoides . . . . . . . . . . . . 465
Mytiloides dresdensis dresdensis ..... 465
Mytiloides fiegei fiegei . . . . . . . . . . . . 458
Mytiloides fiegei mytiloidiformis ..... 468
Mytiloides hercynicus . . . . . . . . . . . . 463
Mytiloides incertus . . . . .. 458, pis. 84-86
Mytiloides labiatiodiformis . . . . . . . . . 465
Mytiloides labiatus . . . . . . . . . . . . . . 463
Mytiloides latus . . . . . . . . . . . . . . . . 466
Mytiloides meekianus . . . . . . . . . . . . 465
Mytiloides modaliaensis . . . . . . . . . . . 465
Mytiloides mytiloides .. . . . . . . . . .. 23
Mytiloides mytiloidiformis . . . . . . . . . 467
Mytiloides opalensis . . . . . . . . . . . . . 468
Mytiloides striatoconcentricus aff. car·pathicus . . . . . . . . . . . . . . . . . . . . 469
Mytiloides submytiloides . . . . . . . . . . 469
Mytiloides teraoka . . . . . . . . . . . . . . 464
Mytiloides teraokai . . . . . . . . . . . . . . 466
Mytiloides sp. aff. M. mytiloidiformis. . . . . . . . . . . . . . . . . . . . . 467, pI. 86
N

Nagatophyllum sp. . . . . . . . . . . . . . . 389
Nagdongia soni . . . . . . . . . . . . . . . . 181
Natica burtoni . . . . . . . . . . . . . . . . . 361
Natica duncani . . . . . . . . . . . . . . . . . 361
Natica epiglottina . . . . . . . . . . . . . . . 361
Natica marochiensis ......
361
Natica pachyope . . . . . . . . . . . . . . . . 361
Natica rostalina .......
361
Natica vitellus . . . . . . . . . . . . 366, pI. 69
Natica (Cryptonatica) oregonensis .... 362
Natica (Cryptonatica) russa ....
362
Naticarius niasensis . . . . . . . . . . . . . . 366
Naticarius cf. N. niasensis ......
366"Naticarius" sp. . .
366, pI. 69
Nautilus .
223, 341
Nautilus angustus . . . . . . . . . . . . . .
12Nautilus asper ...... . . . . . . . . . .. 89
Nautilus bryani . . . . . . . . . . . . . . .
12Nautilus burtini .
13
Nautilus butonensis. . . . . . . . . . . . .. 90
Nautilus cantabrigensis . . . . . . . . . . . 16
Nautilus clementinus . . . . . . . . . . . .
16Nautilus cookana . . . . . . . . . . . . . .. 12
Nautilus dekayi ..
16
0

0

0

0

0

•

0

••••

0

•

••••••

0

•••

0

•••

•••••••••••••••••••

0

•

0

0

0

0

•

•

•

•

•

•

•

•

•

•

•

•

••

0

0

••••••••

0

•

•

••

Nautilus deslongchampianus . . . . . . .. 88
Nautilus elegans ...
88
Nautilus fleuriausianus . . . . . . . . . . " 14
Nautilus intermedius. . . . . . . . . . . .. 15
Nautilus ju~ii . . . . . . . . . . . . . . . . .. 89
Nautilus justus . . . . . . . . . . . . . .
15
Nautilus laevigatus . . . . . . . . . . . . .. 87
Nautilus largilliertianus .. . . . . . . . .. 17
Nautilus leiotropis ....
89
Nautilus mikado . . . . . . . . . . . . . . . . 339
Nautilus neocomiensis . . . . . . . . . . .. 88
Nautilus orbignyi . . . . . . . . . . . . . .. 15
Nautilus plicatus . . . . . . . . . . . . . . .. 89
Nautilus pompilius . . . . . . . . . . . 12, 298
Nautilus pseudobouchardianus ...... 225
Nautilus pseuQoelegans . . . . . . . . . . . 88
Nautilus saxbii .... '. . . . . . . . . . . " 89
Nautilus sharpei . . . . . . . . . . . . . . . . 340
Nautilus sowerbianus . . . . . . . . . . . 13, 87
Nautilus sowerbyi . ~ . . . . . . . . . . . " 14
Nautilus splendens . . . . . . . . . . . . . . 12
Nautilus sublaevigatus . . . . . . . . . . . . 224
Nautilus subtruncatus . . . . . . . . . . .. 89
Nautilus suciensis . . . . . . . . . . . . . . . 228
Nautilus texanus . . . . . . . . . . . . . . " 89
Nautilus turcicus . . . . . . . . . . . . . . . 15
Nautilus undulatus . . . . . . . . 89, 228, 289
Nautilus vaelsensis
89
Nautilus (Cymatoceras) carlottensis ... 228
Nautilus (Cymatoceras)pseudo·atlas .. 90
Nautilus (Cymatoceras) virgatum ..... 336"Nautilus neubergicus" ...
87
Neobakevellia
83
Neocalamites ......
283
Neocalamites carrerei . . . . . . . . . . . . 283
Neocalamites hoerensis . . . . . . . . . . . 283
Neocalamites spo A . . . . . . . . . . . .
283
Neocalamites sp. efr. N. hoerensis .... 283
Neocalamites? sp.· ....
283
Neocymatoceras tsukushiense . ',' . . .. 90
Neohibolites miyakoensis . . . . . . . . . . 125
Neonesidea villosa . . . . . . . . . . . . . . . 139
Neospath.odus sp.
169
Nilssonia . . . . . . . . . . .
38
Nilssonia Schaumburgensis . . . . . . . .. 39
Nilssonia acuminata . . . . . . . . . . . . . 45
Nilssonia bargi·bidi . . . . . . . . . . . . .. 42
Nilssonia borealis
44
Nilssonia brevis . . . . . . . . . . . . . . . . 38
Nilssonia brogniarti . . . . . . . . . . . . " 44
Nilssonia complicatus . . . . . . . . . . . . 42
0

•

•

•

•

0

0

0

0

•

•

•

•

•

•

•

•

•

•

•

••

0

••

•

••

•••••••••

0

0

•

•

•

••

•••••••••••••

0

••

0

•

•

•

"

•••••••••••

0

0

0

••

••••

0

••

••••••••••••••

0

0

0

•

0

••••

•••••••

o'

0

••

•

•

•

•

•

•

513

Nilssonia compta . . . . . . . . . . . . . " 47
Nilssonia efr. compta. . . . . . . . . . . .. 45
Nilssonia elegans. . .
. ...... .
39
Nilssonia gigantea . . . . . . . . . . . . . " 42
Nllssonia grandifolia . . . . . . . . . . . .. 42
Nilssonia grandifolia f. rarinervis .
42
Nilssonia grossinervis . . . . . . . . . . . " 42
Nilssonia incisoserrata..... "
47
Nilssonia inouyei ...
38
Nilssonia efr. inouyei .
. .. 38, pI. 13
Nilssonia japonica .
. 39, pI. 14
Nilssonia kendalli '.. .
47
Nilssonia kotoi .... .
44
Nilssonia macrophylla
42
Nilssonia magnifolia
42
Nilssonia mediana ... .
46
45
Nilssonia minor .... .
Nilssonia muensteri . . . . . . . . . .
43
Nilssonia nigracollensis . . . . . . . . . . . 38
Nilssonia orientalis var. minor. . . .
39
Nilssonia efr. orientalis . . . . . . . . . . . 42
Nilssonia ex gr. orientalis
38
39
Nilssonia parvula .....
Nilssonia polymorpha . .
45
Nilssonia pteropylloides ..... '.
45
Nilssonia revoluta . . . . . . . . . . . . . .. 39
Nilssonia saighanensis . . . . . . . . . . .. 42
Nilssonia schaumburgensis . . . . . . . . , 45
Nilssonia schmidtii. . . . . . .. 45
Nilssonia simplex . . . . . . . . .
39
Nilssonia sinensis .. . . . . .
44
Nilssonia stenophylla. . . . . . .
42
Nilssonia taeniopteroides . . . . . . . . .. 43
Nilssonia taeniopteroides !Jar. bifurcata. . . . . . . . . . . . . . . . . . . . . . . . . . 43

Nilssonia tenuicaulis
45
Nilssonia thomasi . . . . . . . . . . . . . .. 43
Nilssonia undulata ....
39
Nilssonia vittaeformis . . . . . . . . . . .. 39
Nilssonia sp. . . . . . . . . . . . . . . . . . . 39
Nilssonia sp. 1. . . . . . .. 45
Nilssonia sp. A . . . . . . . . . . . . . 43, pI. 14
Nilssonia sp. B . . . . . . . . . . . . . . . .. 45
Nilssonia sp. C . . . . . . . . . . . . . . . .. 45
Nilssonia sp. D . . . . . . . . . . . . . . . .. 46
Nilssonia sp. E . . . . . . . . . . . . . 47, pI. 13
Nilssoniopteris . . . . . . . . .
50
Nilssoniopteris ajorpokensis . . . . . . .. 50
Nipponarges . . . . . . . . . . . . . . . . .. 477
Nipponarges mediosulcatus .
479, pis. 87, 88

Nippononaia . . . . . . . . . . . . . . . . . . 177
Nipponanaia ryosekiana . . . . . . . . . . . 183"Nippononaia" sengokuensis . . . . . . . 183"Nippononaia" wakinoensis . . . . . . . . 177
Nonion mircus . . . . . . . . . . . . .. .. 409
Nuculana . . . . . . . . . . . . . . . . . . . . 80
Nuculana (Dacryomya) konishii .... .
· . . . . . . . . . . . . . . . . . . . . . 80, pI. 15

Nuculana (Dacryomya) ef. konishii ...
· . . . . . . . . . . . . . . . . . . . . . 80, pI. 15
71
71
75

Nuculana ikebei . . . . . . . . . . 'Nuculana pernula sadoensis
Nuculana yokoyamai ..

o. .. 17,342
Obinautilus .....
17
Obinautilus pulchra ... .
. . . . . . 26
Olenus . . . . . . . . . . . .
Olenus gibbosus . . . . . . . . .
. .. 32
Oligosphaeridium complex. .. . .... 384
Operculodinium . . . . . . . . . . . . . . . . 381
Operculodinium cf. hirsutum .
. 383
Operculodinium tiara . . . . . . . . . . . . 383
Operculodinium uncinispinosum .... .
· . . . . . . . . . . . . . . . . . . . . 382, pI. 73

Orbiculoidea sp. . . . . . . . . . . . . . . .. 98
Orthotetes . . . . . . . . . . . . . . . . . . . . 305
Orthotetes australis. . .
. .... 305
Orthotetes ef. australis . . . . . . . . . . . . 305
Orthotetes sp. . . . . . . . . . . . . 305, pI. 61
Oryza. . . . . . . . .
. . . . . . . . . . . 319
pPachyoncus . . . . . .
. . . . . . . . . 257
Pachyoncus crassus .
. . . . . . . . . 257
Pachyoncus kamiasoensis .. 257, pIs. 52, 53
Pactarentinia . . . . . . . . . . . . . . . . . . 109
Pactarentinia holdsworthi .. 109, pIs. 22, 23
Palacantholithus . . . . . . .
. ... 104
Palacantholithus stellatus . .... , 104, 174
Palacantholithus? . . . . . . . . . . . . . . . 101
Palaeoscenidium . . . . . . . . . . . . . 96, 169
Palaeoscenidium (s. I.) quadriramosum· . . . . . . . . . . . . . . . . . . . . . . . . . 107

Palaeoscenidium cladophorum ... 99, 171
Palaeoscenidium? quadriramosum .... 104
Palaeoscenidium sp. A . . . . . . . 107, pI. 21
Palaeoscenidium spp. . . . . . . . . . . . .. 98
Palhindeolithus .... :..
. ... 104
Palhindeolithus ambiguus .
. .. 104. ... 171
Palhindeolithus paululus ..

514

Palhindeolithus pulcher. . . . . . . . 172
Palmenella . . . . . . . . .
. . . . . . . . 135
Palmoconcha . . . . . . . . . . . . . . . . . . 135
Panomya arctica . . . . . . . . . . . . . . .. 70
Panomya arctica . . . . . . . . . . . . . . .. 75
Pantane/lium . . . . . . . . . . . . . . . . . . 256
Pantanellium baileyi . . . . . . . . . . . . . 257
Pantanellium foueatum ... 253, pis. 51, 52
Pantane/lium malheurense . . . . . . . . . 257
Pantanellium sp. A . . . . . . . . . . . . . . 253
Parabolinoides . . . . . . . . .
. . . . .. 26
Parabolinoids contractus ., . . . . . . .. 33
Paracycas . . . . . . . . . . . . . . . . . . .. 47
Paracymatoceras. . . . . . . . . . . . . 89, 343
Paracymatoceras asper . . . . . . . . . . . . 292
Paracymatoceras texanum . . . . . . . . . 292
Paracymatoceras tunghaicum .... 89, 292
Paraheteraster macroholcus . . . . . . . . 447
Parajaubertella imlayi . . . . . . . . . . . . 337
Parasaturnalis sp. . . . . . . . . . . . . . . . 255
Parasepsagon . . . . . . . . . . . . . . . . . . 103
Parentactinia . . . . . . . . . . . . .. 103, 170
Parentactinia nakatsugawaensis ..... .
. . . . . . . . . . . . . . . . . . . . . 172, pI. 37
Parentactinia pugnax . . . . . . . .. 108, 172
Parentactinia uetustum ... 108, pis. 21, 22
Parentactinia sp. . . . . . . . . . . . . . . .. 98
Passaloteuthis . . . . . . . . . . . . . . .. 127
Pcillopora damicornis . . . . . . . . . . . . 159
Pemphigaspis bullata .
. ..... , 32
Pentactinocapsa ...
. . . . . . 101
Pentactinocarpus ..
. . . . . . . 101
Pentactinorbis . . . .
. . . . . . . 101
Phacops nonakai . . . . . . . . . . . . . . . . 475
Phacops (s.s.) okanoi . . . . . . . . . . . . . 474
Phacops (s.s.) sp . . . . . . . . . . . . . . . . 474
Phoenicopsis . . . . . . . . . . . . . . . . . . 273
Phoenicopsis angustifolia . . . . . . . . . . 273
Phoenicopsis ex gr. angustifolia ..... .
. . . . . . . . . . . . . . . . .. 273, pis. 54-56
Phoenicopsis speciosa . . . . . . . . . . . . 275
Phoenicopsis sp . . . . . . . . . . . . . . . . . 273
Phoenicopsis? sp. . . . . . . . . . . . . . . . 273
Picea . . . . . . . . . . . . . . . . . . . 238,319
Picea bicolor . . . . . . . . . . . . . . . . . . 238
Picea bicolor var. reflex . . . . . . . . . . . 238
Picea glehnii . . . . . . . . . . . . . . . . . . 238
Picea jezoensis . . . . . . . . . . . . . . . . . 238
Picea jezoensis var. hondoensis . . . . . . 238
Picea polita . . . .
. ..... 242
Pinus . . . . . . . .
. ..... 238
Pinus densiflora . . . . . . ..
. ..... 238

Pinus diploxylon . . . . . . .
. . . . . . 319
Pinus himekomatsu . . . . . .
. . . . . 238
Pinus karail!nsis . . . .
. 238
Pinus pentaphylla . . . . . . . .
. .... 238
Pinus pumila . . . . . . . . . . . . . . . . . . 238
Pinus thunbergii . . . . . . . . . . . . . . . . 238
Pityophyllum (Pityocladus) sp . . . . . . . 268
Pityophyllum longifolium . . . . . . . . . 277?Pityophyllum longifolium . .
. . 277
Plagiostoma . . . . . . . . . . . . . . . . . .. 79
Platyceramus . . . . . . . . . . . . . . . . . . 203
Plectambonites rhomboidalis uar. analoga . . . . . . . . . . . . . . . . . . . . . . . 350
Plicatounio . . . . . . . . . . . . . . . . . . . 177
Plicatounio (s.s.) . . . . . . . . . . . . . . . . 181'Podocarpites'mentoukouensis . . . . . . 277
Podocarpushii . . . . . . . . . . . . . . . . . 242
Podozamites . . . . . . . . . . . . . . . . . . 277
Podozamites agardhianus var. acuminatus . . . . . . . . . . . . . . . . . . . . . . 278
Podozamites distans . . . . . . . . . . . . . 278
Podozamites distans var. nagatoensis .. 278
Podozamites distans var. ozawae ..... 278
Podozamites ex gr. distans . . . . . . . . .
. . . . . . . . . . . . . . . . . . 277, pis. 55, 56
Cfr. Podozamites gracilis . . . . . . . . . . 278
Podozamites lanceolatus . . . . . . . . . 277
Podozamites schenki . . . . . . . . . . . . . 278
Podozamites sp. A . . . . . . . . . . . . . . 279
Podozamites sp. B . . . . . . . . . . . . . . . 279
Podozamites sp. cf. P. griesbachi . . . . . 277
Podozamites sp. cfr. astartensis . . . . . . 279
Podozamites (Cycadocarpidium?) sp. C. . . . . . . . . . . . . . . . . . . . . 280, pI. 56
Polyentactinia . . . . . . . . . . . . . . . . . 103
Polyentactinia craticulata . . . . . . . . . . 103
Polynemamussium alaskense . . . . . . .. 71
Porites sp . . . . . . . . . . . . . . . . . . . . . 120
Portlandia japonica . . . . . . . . . . . . .. 71
Prionocyclus aberrans . . . . . . . . . . . . 470
Prionocyclus nouimexicanus .
. . 470
Procymatoceras . . . . . . . . . . . . . . .. 89
Procymatoceras baberi . . . . . . . . . . .. 89
Producta analoga . . . . . . . . . . . . . . . 350
Pseudoalbaillella . . . . . . . . . . . . . . . .. 2
Pseudoalbaillella globosa . . . . . . .
5Pseudoalbaillella lomentaria .... .
5
Pseudoalbaillella ornata . . . . . . . .
5Pseudoalbaillella rhombothoracata .
2
Pseudoalbaillella scalprata . . . . . .
2Pseudoalbaillella scalprata m. postscalprata . . . . . . . . . . . . . . . . . 3, pI. 2

515

Pseudoalbaillella scalprata m. rhqmbothoracata . . . . . . . . . . . . . . . . 3, pI. 3
Pseudoalbaillella scalprata m. scalprata. . . . ... . . . . . . . . . . . . . . . .. 2, pI. 1
Pseudoalbaillela simplex. . . . . . . . . . .. 5
Pseudocenoceras. . . . . . . . . . . .. 16, 342
Pseudocenoceras applanatum ....... 17
Pseudocenoceras archiacianus . . . . . .. 17
Pseudocenoceras fittoni . . . . . . . . . .. 17
Pseudoctenis . . . . . . . . . . . . . . . . .. 36
efr. Pseudoctenis brevipennis . . . . . .. 38
Pseudoctenis capensis .. . . . . . . . . .. 38
Pseudoctenis carteriana . . . . . . . . . .. 38
Pseudoctenis cteniforme . . . . . . . . . . 38
Pseudoctenis herriesi . . . . . . . . . . . .. 38
Pseudoctenis lanei. . . . . . . . . . . . . .. 38
Pseudoctenis nipponica .... 36, pIs. 12-14
Pseudoctenis weberi . . . . . . . . . . . .. 38
Pseudoctenis sp. . . . . . . . . . . . . . . .. 38
Pseudocycas . . . . . . . . . . . . . . . . . . 47
Pseudohastigerina . . . . . . . . . . . . . . . 409
Pseudohastigerina micra . ...... 409, pI. 78
Pseudohyria sp. . . . . . . . . . . . . . . . . 183
Pseudowashitaster . . . . . . . . . . . . . . . 451
Pseudowashitaster mystic us ... 451, pI. 83
Pteroperna . . . . . . . . . . . . . . . . . . . 83
Pteroperna kitadaniensis ...... 83, pI. 15
Pterophyllum . . . . . . . . . . . . . . . . .. 36
Pterophyllum jaegeri . . . . . . . . . . . .. 43
Pterophyllum ? sp. aff. Nilssonia tenuicaulis . . . . . . . . . . . . . . . . . . . . .. 46
Pterygocythereis scalaris . . . . . . . . . . 139
Ptychaspis . . . . . . . . . . . . . . . . . . .. 26
Ptychaspis bullasa . . . . . . . . . . . . . " 33
Ptychoparia permulta . . . . . . . . . . . . 396
Pustula . . . . . . . . . . . . . . . . . . . . . . 305
Pustula pustulosa . . . . . . . . . . . . . . . 306
Pustula cf. P. pustulosa ...... 306, pI. 61
Pustula cf. P. tesuipustulata . . . . . . . . 306
Pycnodonte musashiana . . . . . . . . . . . 161
Q

Quercus . . . . . . . . . . . . . . . . . . . . . 239
Quercus acuta . . . . . . . . . . . . . . . . . 239
Quercus acutissima . . . . . . . . . . . . . . 239
Quercus dentata . . . . . . . . . . . . . . . . 239
Quercus glauca . . . . . . . . . . . . . . . . . 239
Quercus phillyraeoides . . . . . . . . . . . . 239
Quercus serrata . . . . . . . . . . . . . . . . 239
Quercus variablilis . . . . . . . . . . . . . . . 239

R

Radimella nodulosa . . . . . . . . . . . . . . 139
Radiolichas . . . . . . . . . . . . . . . . . . . 479
Radiolichas araneus . . . . . . . . . . . . . . 487
Reedops . . . . . . . . . . . . . . . . . . . . . 475
Reesidites minimus . . . . . . . . . . . . . . 470
Reissia hystrix . . . . . . . . . . . . . . . . . 500
Rhipidomella . . . . . . . . . . . . . . . . . . 305
Rhipidomella sp. . . . . . . . . . . 305, pI. 61
Ristola . . . . . . . . . . . . . . . . . . . . . . 259
Ristola(?) turpicula . . . . . . . . . 259, pI. 53
Robaia robai . . . . . . . . . . . . . . . . .. 71
Rugosochonetes . . . . . . . . . . . . . . . . 352
Rugosochonetes celticus . . . . . . . . . . 352
Rugosochonetes cf. R. celticus ..... .
. . . . . . . . . . . . . . . . . . . . . 352, pI. 67
Rugosochonetes aff. R. hardrensis .... 352
Rugosochonetes speciosus . . . . . . . . . 352
Rugosochonetes sp . . . . . . . . . . . . . . . 352
S

Salix . . . . . . . . .
. . . . . . . . . . . 242
Sao hirsuta . . . . . . . . . . . . . . . . . . . 395
Schizophoria . . . . . . . . . . . . . . . . . . 304
Schizophoria resupinata ...... 304, pI. 61
Scutellum . . . . . . . . . . . . . . . . . . . . 475
Scutellum (Thysanopeltella) paucispinosa . . . . . . . . . . . . . . . . . . . . . . . 475
Sepsagon . . . . . . . . . . . . . . . . . . . . . 102
Sequoia sempervirens . . . . . . . . . . . . 23
Somphoentactinia . . . . . . . . . . . . . . . 103
Sphaeroidinella senni . . . . . . . . . . . . . 408
Sphaeroidinellopsis senni . . . . . . . . . . 408
Sphenobaiera . . . . . . . . . . . . . . . . . . 268. Sphenobaiera eurybasis . . . . . . . . . . . 268
Sphenobaiera cfr. eurybasis ... 268, pI. 54
Sphenobaiera gyron . . . . . . . . . . . . . 270
Sphenobaiera huangi . . . . . . . . . . . . . 270
Sphenobaiera ikorfatensis . . . . . . . . . . 271
Sphenobaiera longifolia . . . . . . . . . . . 271
Sphenobaiera nipponica . .. 268, pIs. 54-56
Sphenobaiera ophioglossum . . . . . . . . 270
Sphenobaiera cfr. ophioglossum .... .
. . . . . . . . . . . . . . . . . . 270, pIs. 54-56.
Sphenobaiera pecten . . . . . . . . . . . . . 268
Sphenobaiera pulchella . . . . . . . . . . . 271
Sphenobaiera spectabilis . . . . . . . . . . 271
Sphenoceramus . . . . . . . . . . . . . . . . 202
Sphenoceramus orientalis . . . . . . . . .. 94
Sphenoceramus schmidti ...... '.' .. 94

516

Spirifer. . . . . . . . . . . . . . . . .. 307, 353
Spirifer bisulcatus . . . . . . . . . . . . . . . 308
Spirifer ex gr. bisulcatus . . . . . . . . . . . 308
Spirifer crassus . . . . . . . . . . . . . . . . . 353
Spirifer liangchowensis . . . . . . . . . . . 307
Spirifer sp. . . . . . . . 307, 353, pIs. 61, 67
Spirifer (Tylothyris) laminosa ....... 355
Spirifera bisulcata . . . . . . . . . . . . . . . 308
Spirifera laminosa . . . . . . . . . . . . . . . 355
Spongentactinnella . . . . . . . . . . . . . . 103
Stegodon orientalis . . . . . . . . . . . . . . 235
Stephanodiscus . . . . . . . . . . . . . . . . 248
Stephanodiscus carconensis . . . . . . . . 248
Stichocapsa japonica . . . . . . . . . . . . . 255
Stigmaulax guppiana . . . . . . . . . . . . . 361
Storgaardia . . . . . . . . . . . . . . . . . . . 277
Storgaardia hissarensis . . . . . . . . . . . . 277
Storgaardia spectabilis .... 277, pIs. 54, 56
Strophomena analoga . . . . . . . . . . . . 350
Strophomena rhomboidalis var. analoga. . . . . . . . . . . . . . . . . . . . . . . . . . 350
Stylosphaera? . . . . . . . . . . . . . . . . . 103
Subbotina . . . . . . . . . . . . . . . . . . . . 409
Subbotina bakeri . . . . . . . . . . . . . . . 403
Subbotina frontosa boweri .... 409, pI. 79
Subbotina inaeguispira . ...... 410, pI. 79
Subbotina linaperta . . . . . . . . . 410, pI. 79
Subbotina pseudoeocaena .... 410, pI. 79
Subprionocyclus neptuni . . . . . . . . . . 470
Subprionocyclus normalis . . . . . . . . . 470
Sweden borgia . . . . . . . . . . . . . . . . . 280
Sweden borgia attenuata . . . . . . . . . . . 282
Swedenborgia coreanica . . . . . . . . . . . 282
Swedenborgia cryptomerioides ...... 280
Swedenborgia lata . . . . . . . . . . . . . . . 282
Sweden borgia longifolia . . . . . . . . . . . 282
Swedenborgia major . . . . . . . . . . . . . 282
Swedenborgia megasperma . . . . . . . . . 282
Swedenborgia minor . . . . . . . . . . . . . 282
Sweden borgia onoyamai . . . . . . . . . . 282
Sweden borgia rigida . . . . . . . . . . . . . 282
Swedenborgia rigid a var. acuminata ... 282
Swedenborgia tuttosperma . . . . . . . . . 282
Sweden borgia sp. A . . . . . . . . . . . . . . 280
Sweetognathus whitei . . . . . . . . . . . .. 5
Syringothyris . . . . . . . . . . . . . . . . . . 354
Syringothyris fontanalis . . . . . . . . . . . 354
Syringothyris japonica . . . . . . . . . . . . 354
Syringothyris kitakamiensis . . . . . . . . 354
Syringothyris laminosa . . . . . . . . . . . 355
Syringothyris aff. S. laminosa ....... 355

Syringothyris sakamotoi . . . . . . . . . . 354
Syringothyris sp. . . . . . . . . . . 354, pI. 67

T
Taenicephalus . . . . . . . . . . . . . . . . . 26
Taenicephalus shumardi . . . . . . . . . .. 33
Taeniopteris . . . . . . . . . . . . . . . . . . 48
Taeniopte,ris daintreei . . . . . . . . . . .. 48
Taeniopteris gracilis . . . . . . . . . . . . . 39
Taeniopteris jourdyi . . . . . . . . . . . .. 50
Taeniopteris efr. jourdyi ... 50, pI. 13, 14
Taeniopteris parvula . . . . . . . . . . . .. 48
Taeniopteris efr. parvuls. . . . . . . . . .. 48
Taeniopteris richthofeni . . . . . . . . . .. 50
Taeniopteris stenophylla . . . . . . . . . . 50
Taeniopteris sp. 1 . . . . . . . . . . . . . .. 48
Taeniopteris sp. A . . . . . . . . . . . 50, pI. 12
Taeniopteris sp. B . . . . . . . . . . . . . .. 51
Taeniopteris sp. C . . . . . . . . . . . . . .. 51
Taeniopteris sp. D . '. . . . . . . . . . . . .. 51
Taeniopteris sp. cfr. T. nabaensis . . . .. 50
Tandarnia arachnoconcha . . . . . . . . . . 171
Tandarnia recoarense . . . . . . . . . . . . . 171
Tandarnia recoarensis . . . . . . . . . . . . 172
Tandarnia simplicima . . . . . . . . .... 174
Taniopteris inouyei . . . . . . . . . . . . .. 50
Taxodium . . . . . . . . . . . . . . . . . . . . 238
Taxodium distichum . . . . . . . . . . . . . 238
Terataspis . . . . . . . . . . . . . . . . . . . . 479
Terebratulina peculiaris . . . . . . . . . . . 161
Terranovia. . . . . . . . . . . . . . . . . . . . 478
Terranovia nalivkini . . . . . . . . . . . . . 479
Tetragonites popetensis . . . . . . . . . .. 94
Tetragonites cf. T. timotheanus . ..... 337
Thysanopeltella . . . . . . . . . . . . . . . . 475
Thysanopeltella (Septimopeltis) ..... 475
Thysanopeltella (Septimopeltis) paucispinosa . . . . . . . . . . . . . . . . . . . . . 475
Tlecerina . . . . . . . . . . . . . . . . . . . . . 110
Tlecerina exilis . . . . . . . . . 111, pIs. 25, 26
Tlecerina fenestrata. . . . .. 110, pIs. 26,27
Tlecerina horrida' . . . . . . . . 110, pIs. 23-25
Tlecerina sp. A . . . . . . . . . . . . . 113, pI. 27
Tomiopsis . . . . . . . . . . . . . . . ..... 308
Tomiopsis sp . . . . . . . . . . . . . . . 308, pI. 61
Trachyle.beris lungkangensis . . . . . . . . 139
Triassocampe . . . . . . . . . . . . . . . . . . 259
Triassothamnus . . . . . . . . . . . . . . . . 101
Tricolocapsa ef. rusti . . . . . . . . . . . . . 255
Tricolocapsa plicarum . . . . . . . . . . . . 255
Trigonioides . . . . . . . . . . . . . . . . . . 180

517

Trigo1'!ioides (Trigonioides) jaehoi ....
. . . . . . . . . . . . . . . . . . 183, pIs. 39, 40
Trigonioides (Wakinoa) wakinoensis ...
. . . . . . . . . . . . . . . . . . . . . 183, pI. 39
Trigonioides (Wakinoa) sp. cf. T. (W.)
tamurai . . . . . . . . . . . . . . . . 180, pI. 38
Trigonioides kodairai. . . . . . . ...... 178
Trigonioides kodairaiformis .. . ..... 178
Trigonioides paucisulcatus . . . . . . . . . 183
Trigonioides tetoriensis . . . . . . . . . . . 181
Trochocyathus hanzawai . . . . . . . . . . 118
Truncorotaloides . . . . . . . . . . . . . . . 407
Truncorotaloides collactea .... 407, pI. 78
Trymataspis convexus . . . . . . . . . . . . 395
Tsuga . . . . . . . . . . . . . . . . . . . . . . . 319
Tsuga diversifolia . . . . . . . . . . . . . . . 238
Tsuga sieboldii . . . . . . . . . . . . . . . . . 238
Turritella fortilirata. . . . . . . . . . . . .. 71
Tylothyris . . . . . . . . . . . . . . . . . . . . 355
Tylothyris cf. T. laminosa ... "...... 356
Tylothyris laminosa . . . . . . . . 355, pI. 67
Tylothyris planimedia .". . . . . . . .' ... 356
Tylothyris transversa . . . . . . . . . . . . . 356
Tylothyris sp. . . . . . . . . . . . . . . . . . 356
Ulflmus . . . . . . . . . . . . . . . . . . 244, 325
Umbonium . . . . . . . . . . . . . . . . . . . 131
Undispirifer (Nakazatothyris) . . . . . . . 475
Unuma echinatus . . . . . . . . . . . . . . . 253

v
Venericardia ferruginea . . . . . . . . . .. 71
Viviparus sp . . . . . . . . . . . . . . . . . . . 181
W

Wakinoa . . . . . . . . . . . . . . . . . . . . . 180
Wakinoa tetoriensis . . . . . . . . . . . . . . 181
Washitaster (1) macroholcus . . . . . . . . 447
Welleraspis lata. . . . . . . . . . . . . . . .. 32
Welleraspis lochmanae . . . . . . . . . . . , 32
Welleraspis sWartizi . . . . . . . . . . . . .. 32
Wilson ella pennsylvanica . . . . . . . . . . 32

x
Xestoleberis . . . . . . . . . . . . . . . . . . . 152
Xestoleberis suetsumuhana . . . . . . . . .
. . . . . . . . . . . . . . . . . . 153, pIs. 34,35
Xestoleberis sp. . . . . . . . . . . . . . . . . 152
Xiphachistrella . . . . . . . . . . . . . . . . . 104
Xiphocladiella cuja . . . . . . . . . . . . . . 173yYanishewskiella . . . . . . . . . . . . . . . . 352
Yanishewskiella cf. Y. angulata ..... .
. . . . . . . . . . . . . . . . . . . . . 352, pI. 67
Yanishewskiella? sp . . . . . . . . . . . . . . 353
Yoldia notabilis . . . . . . . . . . . . . . . , 71
Yuknessaspis santaquinensis . . . . . . . . 395
Z

Zamites . . . . . . . . . . . . . . . . . . . . . 36
Zelko va . . . . . . . . . . . . . . . . . 239, 321