ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 148

Palaeontological Society of Japan
September 30, 1986

Co-Editors

Hisayoshi 100 and Takashi HAMADA

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The fossil on the cover is Unuma (Spinunuma) echinatus ICHIKAWA and YAO, a
Middle Jurassic multisegmented radiolaria from Unuma, Gifu Prefecture, central
Japan (photo by A. YAO, x 260).

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 409-421, pis. 81-85, September 30,1986

816. SOME INOCERAMIDS (BIV AL VIA) FROMTHE CENOMANIAN (CRETACEOUS) OF JAPAN-I
NEW OR LITTLE KNOWN FOUR SPECIESFROM HOKKAIDO AND KYUSHU*TATSURO MATSUMOTO
c/o Department of Geology, Kyushu University 33, Fukuoka 812
and
MASAYUKINODA
Minami·Oita Junior High School, Oita 870

Abstract.
In a series of papers in 3 parts some new and little known ino·
ceramid species from the Cenomanian of Japan are to be described. In this part we
describe the following four species including three new ones. (1) Birostrina sp.
nov., which belongs to the group of B. concentrica but has a radial groove on theposterior part and ribs of moderate intensity in the late growth-stage. It is represented by a number of specimens from the Middle Cenomanian of the Mifune(central Kyushu), Mikasa (central Hokkaido) and Kotanbetsu (northwestern Hokkaido) areas. (2) Inocermus sp. nov., which is allied to 1. comancheanus but has asubrectangular outline with less convex valves and weaker subcostae. Its holotype
came from the Lower Cenomanian of the Mikasa area. (3) 1. heinzi Sornay, which isrepresented by several specimens from the Upper Cenomanian and/or lowest part of
the Lower Turonian of the Saku area (northwestern Hokkaido). They are very
similar to the holotype from Madagascar. (4) Mytiloides sp. nov., which has a larger
beak angle and more convex anterior margin than M. submytiloides and is characterized by regular concentric ribs combined with fine concentric lirae. Its holotype came from the Zone of Euomphaloceras septemseriatum (Upper Cenomanian)of the Mikasa area.

Introduction

commISSiOn on Cretaceous Stratigraphy of thelUGS (1976-1984), we have obtained a greatamount of mega-fossils which may be usefulfor biostratigraphic and other aims. The majorparts of the collections are those of ammonites
and inoceramid bivalves and some of them havebeen and would be evaluated in improving thetaxonomy and biostratigraphic zonation andcorrelation.

Through the field works in various areas ofJapan to cooperate in the IGCP Project No. 58
Mid-Cretaceous Events (MCE) (1975-1982) andalso in the Regional Working Group for the Sub-

*Received March 20, 1985; Read June 16, 1985
at Osaka City University.
409

410

Tatsuro MATSUMOTO and Masayuki NODA

In a series of papers entitled at the heading,
we are going to describe some species of the
Inoceramidae from the Cenomanian of Japan,
which have been recorded only preliminarilyor left untouched for full description. As the
persons concerned with the cooperative study
vary case by case, the results are to be publishedsuccessively in three parts by different authorships.
For some reasons, some species from the
Cenomanian of Japan are represented by a small
number of specimens. This is probably due toour collection failure and indeed unfavourable
for getting an adequate conception of a species.
It would be, however, better to describe themat this moment than to leave them unrecorded
for a long time, so that someone may obtain asatisfactory sample of population in future torevise the present definition. In Part I we describe four species of which three belong to such
category. The material of this study is the
specimens from the Mikasa, Saku and Kotanbetsu areas of Hokkaido, but one of the species
depends also on those from the Mifune areaof Kyushu which were once treated by Tamura
and Matsumura (1974).
The repositories of the described or mentioned specimens are as follows with abbreviation in parentheses:
Geological Collections, Kyushu University,
Fukuoka (GK)
Geological Collections, Faculty of Education,
Kumamoto University, Kumamoto (KE)Institute of Geology and Palaeontology,
Tohoku University, Sendai (IGPS)
University Museum, University of Tokyo,
Tokyo (UMUT)
Systematic descriptions
Family Inoceramidae Zittel, 1881
As to the authorship of this family we follow
Vokes' (1980) indication.
Kauffman (in Kauffman et al., 1977 andelsewhere) commented that the current taxonomy of this family at generic level based

primarily on external characters should be
revised on the ground of more important interiorfeatures, but has not yet given fully his schemeof classification nor his description of genericdiagnosis, except for a few genera, e.g. Myti[oides. At present, we have no competent ideaof our own scheme of overall taxonomy andhave to depend on the existing concepts of
Dobrov and Pavlova (1959), Cox (1969), Seitz(1961, 1965, 1967, 1970), Sornay (1966),
Dhondt (1983) etc., as well as Kauffman (op.
cit; 1977), with some modification.
For the terms to describe morphologicalcharacters we follow primarily those definedby Nagao and Matsumoto (1939), which havebeen used by Noda (1983,1984 etc.) and Matsumoto and Noda (1985) with some modification.
Readers may refer to Efremova (1978) to compare them with other authors'.
The symbols (or letters) to abbreviate thedimensions and angles in the tables of measurements in this paper are as follows:
h=height, l=length, H=maximum linear dimension from beak to posteroventral extremity,
L=ditto perpendicular to H, s=length of
hinge line (previously indicated as HL), b=
breadth (previously indicated as th or T),
a=anterior hinge angle (angle between hingeline and anterodorsal margin), ~=angle ofumbonal inflation, 'Y=posterior hinge angle(angle between hinge line and posterior
margin), r=obliquity (angle between axis of
growth and hinge line); L=left valve, R=rightvalve
The terms to describe the concentric ornamentation vary between authors and also inaccordance with the characters. In this paper we
use the following terms:
concentric lines or lirae: fine markings on thesurface of shell, which are not impressed on
the internal mould, except in the case of thecomposite internal mould
concentric rings: fine concentric elevationsof the third order, which may correspond tothe concentric lines or lirae but are somewhat
more elevated than them and may be discernible even on the internal mould

816. Cenomanian Inoceramids from Japan-I
concentric subcostae or riblets (minor ribs):
concentric elevations of the second order,
well recognized even on the internal mould,
being larger (i.e. thicker and higher) thanthe concentric rings but somewhat smaller(narrower and lower) than and often branchedfrom or intercalated between the concentricelevation of the first orderconcentric ribs or ridges (major ribs): concentric elevations of the first order. Whenthey are broad but low or weak the term
concentric undulations may be employed.
concentric depressions: concentric furrowalong the constricted or geniculated part ofthe shell in the case when the shell shows anabrupt change in convexity.
Radial or divergent ornaments do not occur
in the described species, except for a radialsulcus or groove in a species.
Radial markings which reflect the interiorshell morphology may appear on some wellpreserved specimens, but they should not betreated as ornamentation.
Genus Birostrina J. Sowerby, 1821

Type species:-Inoceramus sulcatus Parkinson,
1819.
Diagnosis:-Shell small, inequivalve, withnarrow and prominent left umbo and prosogyrous beak; hinge line short, without or with very
narrow posterodorsal auricle; surface ornamented
with fine concentric lirae and with or without
concentric ribs, riblets and/or radial folds.
Remarks:-Presence of the radial folds is
not a generic criterion and the genus is best
represented by Inoceramus concentricus Parkinson, 1919. Although the group of 1. concentricus
was taken too much comprehensively by Nagao
and Matsumoto (1939), Birostrina evidently
ranges up to the Cenomanian in the North
Pacific region.
Birostrina tamurai sp. nov.
PI. 81, Figs. 1-6; PI. 82, Figs. 1-3;
PI. 83, Figs. 1-3; PI. 85, Figs. 4-5

1974. Inoceramus concentricus costatus Nagao

411

et Matsumoto; Tamura and Matsumura,p. 49, pI. 1, figs. 1-5.
1976. Inoceramus concentricus nipponicus
Nagao et Matsumoto; Noda and Matsumoto, pI. Cr-31, fig. 2.
1976. Inoceramus concentricus costatus Nagao
et Matsumoto; Noda and Matsumoto, pI.
Cr-31, fig. 3.
1977. Birostrina n. sp., Kauffman, p. 175.

Material:-Holotype, KE 1976 (PI. 81, Fig. 1),
composite internal mould of left valve from loco
4 (Yaseto, Mifune area, Kyushu) of Tamura andMatsumura (1974). Paratypes, KE 1977 fromloco 1 (Omine), KE 1978 from loco 5 (Okadake
Playground), KE 1979 from loco 2 (Asanoyabu)
and KE 1706 from loco 3 (Subayashi) of the
Mifune area; also GK. H 10164 and H 10165
both valves, GK. H 10166, small left valve, andothers (GK. H 10167-H 10169) from loco Ik
2021a, Pombets gorge, Mikasa district (colI. M.
Noda and T. Matsumoto); GK. H 10082, H
10157-H 10159 from loco R 100 and GK. H
10153-H 10156, H 10160-H 10162 from loco
R 101 of the Kotanbetsu area (colI. T. Matsumoto and H. Okada).
Diagnosis:-Rather small, inequivalve, asymmetrically ovoid shell of the group of Birostrina
concentrica, characterized by the presence of aradial sulcus on the posterior part and thedevelopment of rather irregular concentric ribs
and riblets in late growth stage in addition tofiner concentric lirae or rings.
Description:-The holotype (PI. 81, Fig. 1)
is small and probably not a full-grown individual, but it does show diagnostic characters.
Tamura and Matsumura (1974, pI. 1, figs. 1a1b) have carefully given two figures in somewhat
different views, which clearly show the posterior
radial sulcus and the details of the posterodorsal
part as well as other features, although theauthors did not mention these two characters in
the text. KE 1979 (Tamura and Matsumura,
1974, pI. 1, fig. 5; this paper PI. 81, Fig. 3) issomewhat larger than the holotype but deficientin the anteroventral and posteroventral portions.
KE 1977 (Tamura and Matsumura, 1974, pI. 1,
fig. 2; this paper PI. 85, Figs. 4-5) is also in-412

Tatsuro MATSUMOTO and Masayuki NODA

complete but the largest of the Mifune specimens, attaining to about 75 mm in H in a restored outline. GK. H 10164 (PI. 81, Fig. 6;
PI. 82, Fig. 1) from Mikasa and GK. H 10153(PI. 82, Fig. 2) from Kotanbetsu may represent
the normal size of this species. The right valveof the former is badly eroded, but this deficiency
is supplemented by another specimen, GK. H
10165 (PI. 81, Fig. 5; PI. 82, Fig. 3) from Mikasa,
which however, is somewhat distorted. GK.
10154 from Kotanbetsu has both valves andmore right valves are preserved in the Kotanbetsu sample (e.g. GK. H 10155-10157, H
10159-10161). Shell layer is partly preservedin some Kotanbetsu specimens.
On the basis of a considerable number ofthese specimens the specific characters are wellrecognized. Evidently the left valve is moreconvex than the right, having a fairly inflated,prominent but narrow umbonal part whose beakis considerably curved inward and forward,
exceeding the hinge line.
The hinge line is nearly as long as or slightly
longer than a half of shell length, forming anangle (a) of about 80° with the anterior margin
and that (r) of about 140° with the posterior
margin. The axis of growth extends towards
the posteroventral extremity nearly straightly
or obliquely with a forward gently concaveof about 40°
curvature, forming an angle
to 50° with the hinge-line. The anterior marginmis gently concave below the umbo and broadly
convex in the lower part towards the anteroventral edge, where it is bent to the asymmetrically rounded ventral margin, which passes
to the gently arcuate posterior margin. The
narrow posterodorsal edge is somewhat flattened.
A radial sulcus runs on the posterior partof the disk near the margin of the umbonal
inflation in the left valve, extending towards
the posteroventral part, where the ribs may
show slight sinuosity on crossing the shallow
sulcus. A faint, corresponding sulcus is discernible on some right valve specimens, e.g.
GK. H 10154 and H 10155.
The shell is ornamented with numerous
concentric lirae or rings and rather irregularly
developed concentric ribs and riblets. The
lirae or rings are distinct on the young part(e.g. up to 25 mm in H in the holotype) as fineand dense markings on the shell surface andsometimes even on the internal mould, someof which may be somewhat stronger than others.
Major ribs and branched or intercalated minor
ribs are developed in late growth stages withrather irregular intensity and distance, beingcombined with the finer rings on the interspaces. Major ribs extend to the posterior part,
whereas minor ribs or riblets and rings on
the main part of the disk fade away as they
approach the zone of radial sulcus. Some of the

Explanation of Plate 81
Figs. 1---U. Birostrina tamurai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 411
1. Holotype, KE 1976 (LV), from Mifune; lateral (a), anterior (b), posterior (c)an d dorsal (d) views, X 1. 2.
2. Replica of holotype in the lighting somewhat different from 1a to show theposterior sulcus, x 1.
3. KE 1979 (LV) from Mifune, lateral view, x 1.
4. GK. H 10166 (LV), from Mikasa; lateral (a), anterior (b) and posterior (c) views,
xl.
5. GK. H 10165 (BV) from Mikasa; left (a) and right (b) lateral, anterior (c) andposterior (d) views, x 1.
6. GK. H 10164, left lateral view in the disposition a little different form PI. 82,
Fig. la, x 1.1.
BV: both valves, LV: left valve, RV: right valve.
Photos. (Pis. 81-85) by M. Noda.

MATSUMOTO and NODA: Cenomanian Inoceramids from Japan-I

ld

Plate 81

816. Cenomanian Inoceramids from Japan-I

413

Table Measurements of selected specimens.
Specimens
KE 1976

h
(L)

34.5

l/h

H

L

L/H

b

b/h

26.0

.75

37.0

26.7

.72

14.0

.41

1B.0

27.0

.45

GK. H 10164 (L)

59.6

36.7

.62

62.5

3B.0

.61

GK. H 10164 (R)

52.B

36.7

.70

56.B

3B.0

.67

4B.4

34.2

.71

1B.5

GK. H 10153 (L)

ex

i3

r

Ii

.61

BOo

60°

140°

50°

19.0

.52

BO°

55°

19.0

.52

130 0

56°

130°

56 0

1370

50°-65°

145°

50 0

55°

GK. H 10157 (R)

40.0

30.B

.77

42.0

31.6

.75

12.0

.30

GK. H 100B3 (.L)

50.0

42.0

.B4

5B.6

37.B

.65

21.4

.42

ribs extended from the main part of the disk
reach the narrow, flat, posterodorsal edge with
gradual weakening, as seen in the well preserved
specimens (e.g. holotype and GK. H 10082).
Etymology:-This species is dedicated toProfessor Minoru Tamura of Kumamoto University who has contributed much to the palaeontology of the Mesozoic Bivalvia.
Comparison and discussion:-We owe muchto ErIe G. Kauffman (1977, p. 175) who suggested that the presence of the posterior radialsulcus should be evaluated as a specific character in Birostrina. In fact, the sulcus is observable in a number of examined specimens fromparticular beds of the three areas. In somespecimens (e.g. GK. H 10082 figured by Nodaand Matsumoto, 1976; this paper PI. 83, Fig.
3) the sulcus is faint.
The lectotype of Inoceramus concentricus
var. costatus Nagao et Matsumoto, 1939 was
designated by Tamura and Matsumura (1974, p.
49) to the one illustrated by Nagao and Matsumoto, 1939, pI. 24, figs. 1a---"1!, i.e. UMUT. MM
6480 [=1-690]. It has no posterior sulcus and isdistinguished from B. tamurai. It resembles
closely Inoceramus concentricus var. porrectus
Woods (1917, pI. 4, figs. la-b), as Nagao andMatsumoto (1939, p. 271) have already pointed
out. Kauffman considered it as identical with
B. porrecta (Woods) from New Zealand. We
would defer the conclusion about this point
until we could examine more carefully the
specimens from New Zealand and Japan.
Kauffman (op. cit.) compared the species
represented by Tamura and Matsumura's specimens (that is now called B. tamurai) with anundescribed species, "Birostrina n. sp. (sulcate)",

s/l

~15.4

21.4

~.50

100°

.51

90°

55°

from the Albian of England. This is interesting,but B. tamurai has concentric ribs of moderateintensity in the late growth-stage. The youngshell of B. tamurai ornamented with fine concentric lirae and rings may be similar to that
Albian species, which has not yet been clearly
described nor illustrated. The sulcus may be
the same kind of radial plication as those of
B. sulcata (Parkinson, 1819) and B. subsulcata(Wiltshire, 1879) (see Woods, 1911).
Occurrence:-The type locality is Yaseto,
Toyono-mura, Shimomashiki-gun, Kumamoto
Prefecture (central Kyushu). This and othernearby localities are recorded by Tamura andMatsumura (1974). They are all in the lithic
sandstone of the Lower Formation of the Mifune
Group. Eucalycoceras sp. cf. E. spathi (Collignon) is recorded to occur in the same formation (Matsumoto in Tamura and Matsumura,1974). On the evidence of this ammonite andother associated fossils (Tamura, 1979) the
Lower Mifune is assigned to the Middle Cenomanian.
Other described specimens came from twoplaces. One is loco Ik 2021a at Pombets gorge
in a tributary of the River Ikushumbets, Mikasa
district (central Hokkaido), in the Mikasa Sandstone between the layers rich in trigonians.
Calycoceras cf. C. asiaticum (Jimbo) occurs
fragmentarily at loco Ik 2021b in a trigonian
bed. For the route map and stratigraphic section
see Matsumoto (1965, figs. 3 and 5). The otheris locs_ R 100 and R 101, on road side of Highway 239, about 800 m linearly west of Kiritachi Pass, Kotanbetsu area (northwestern
Hokkaido), from calcareous nodules in the
mudstone of the Middle Yezo Group. Caly-Tatsuro MATSUMOTO and Masayuki NODA

414

than long later. Hinge plate fairly thick; hingeline moderately long, somewhat more than halfof shell-length. Anterior hinge angle (O!) about

coceras sp. of newboldi group was found fromloco R 100. For the map and stratigraphic sec·tion of this route see Matsumoto and Okada(1973, figs. 7-8).
To sum up, the known age of Birostrina
tamurai is Middle Cenomanian. The fossils
occur separately in the Mifune area but in groups
often with both valves conjugated in the Mikasa
and Kotanbetsu areas.

1050 and angle of umbonal inflation «(3) 85 0 •Anterior margin nearly straight, with very gently
arcuate curvature towards rounded anteroventral
edge; ventral margin rather broadly and asymmetrically rounded, passing gradually to gently
arcuate posterior margin. Posterior hinge angle(1) from 120 0 (in young stage) to 1400 (later).
Axis of growth runs along the central line offlank, forming angle (0) of about 65 0 with thehinge line.
Shell in the main growth-stage ornamented
with numerous, fairly dense, rather weak concentric subcostae, which become very weak in
the last stage. Major ribs of the first order scarcely developed, except for the indistinct ones inthe early middle grwoth-stage, but the occurrence of comparatively more distinct subcostae
at periodic intervals associated with somewhat
deeper interspaces may imply incipient or
hidden major ribs. Fine concentric lirae of thethird order discernible on the surface of theouter shell layer which is preserved on theumbonal part of the holotype.
Etymology:-This species is dedicated to Mr.
Takemi Takahashi of Mikasa, who is one of themost eminent naturalists in collecting scientifically valuable fossils from Hokkaido.
Remarks:-The holotype itself is fairly good,
but no more specimen has been obtained fromthe same bed and the extent of variation is un-

Inoceramus takahashii sp. nov.

PI. 84, Figs. 1a-c
Holotype:-GK. H. 10145, both valves, collected by Takemi Takahashi from the upper
reaches of the 8th branch of the Kami-ichi-nosawa, a tributary to the River Ikushumbetsu,
Mikasa district, central Hokkaido. Outer andinner shell layers preserved on the umbonal
part; inner shell layer attached on some contiguous part; the rest main part of left valve andsome portions of right valve internal mould.
Specific characters:-Shell subequivalve; theumbo of left valve slightly more projecting andincurved than the right. Valve gently convex
in the main part, but for somewhat more inflated
umbonal part. Posterior part flattened, passing
gradually from the main part of the disk, except
the umbonal part, where there is a faint auriculardepression. Anterior side steep, with concave
anterodorsal part.
Shell subrounded rectangular in rough outline,
nearly as high as broad in early stage and higher

Measurements of holotype (left valve).
Specimen

GK. H 10145

h

92.0

71.0

l/h

H

L

L/H

b

b/h

.77

102.0

72.0

.71

24.0

.26

r

sll
41.0

.58

105

0

85

0

1400

Explanation of Plate 82
Figs. 1-3. Birostrina tamurai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 411
1. GK. H 10164 (BV) from Mikasa; left lateral (a), right lateral (b), anterior (c),posterior (d) and dorsal (e) views, X 1.
2. GK. H 10153 (LV) from Kotanbetsu; lateral (a), anterior (b), posterior (c) anddorsal (d) views, x 1.
3. GK. H 10165 (BV) from Mikasa; anterior view in the lighting somewhat different
from PI. 81, Fig. 5d, x 1.

65 0

MATSUMOTO and NODA : Cenomanian Inoceramids from Japan-I

Plate 82

816. Cenomanian Inoceramids from Japan-I

known at present. There are, however, somespecimens from other areas within Hokkaido,
which are comparable but not quite identicalwith the holotype. They will be described inPart 3.
Comparison:-This species is somewhat similar
to Inoceramus comancheanus Cragin, 1895 fromthe Albian of Texas. By courtesy of Dr. W. A.
Cobban, there are plaster casts (GK. H 9194 andH 9195) of the two syntypes (USNM. 32686 A,B) (for their and other illustrations see Hill,
1901 and Reeside, 1923) from the Duck Creek
Limestone. Furthermore, Pergament (1964)
described an example from the Albian of Kamchatka under Inoceramus cf. I. comancheanus
and then Eigenheer and Sornay (1974) anotherexample from the basal Cenomanian of Montlaux(Alpes-de-Provence, France) under Inoceramus
aff. I. comancheanus. Our species differs fromthem in having more subrectangular outline,with larger umbonal angle (~) and more broadly
rounded ventral margin and weaker and denser
concentric su bcostae. Especially, the concentricribs are distinctly stronger in the late growthstage of I. comancheanus. In the two syntypes(left and right valves) of that species the mainpart of the flank is more convex and slopes
down less gradually to the posterior flat part,with a very faint, incipient, radial sulcus between
the two parts. The specimen from the basal
Cenomanian of Montlaux, mentioned above,is more compressed than the American types
and is similar to ours in that respect, but is extended more obliquely and has more irregularand stronger ribs than the holotype of our
species. Anyhow, that French species is noteither identical with ours.
I. takahashii is somewhat similar to I. prefragilis Stephenson, 1953 (see also Kauffman
et al., 1977), from the Middle Cenomanian ofTexas and Oklahoma, in the general outline
and ornamentation, but the latter has a moreconvex main part of the flank, more distinct
auricular sulcus or break, and finer but more
distinct concentric subcostae.
The specimens described long ago by Etheridge(1872) from the "Marathon Formation" of415

Queensland (Australia) under I. multiplicatus
Stoliczka var. elongatus Etheridge, I. pernoides
Etheridge and I. sp. aff. I. pro blema ticus
d'Orbigny seem to be allied to I. comanchean us.
The third one is especially similar to I. takahashii in the shell-form but different in having
more distinct concentric subcostae on its younger half and stronger concentric ribs on its later
half. The age of the Marathon Formation wasconsidered as Cenomanian by Etheridge but
must be Albian in the light of recent knowledge
about the Cretaceous stratigraphy of Australia(Ludbrook, 1978).
To sum up, the described specimen found by
T. Takahashi represents a new species belonging
to the group of I. comanchean us.
Occurrence :-In a calcareous nodule derived
from the sandstone referable to the lower partof the Mikasa Formation in the upper reachesof the 8th branch (Hachi-no-sawa in Japanese)of the Kami-ichi-no-sawa, a tributary of theRiver Ikushumbets. As Sharpeiceras sp. was
obtained by T. Takahashi form a nearby locality,
the sandstone is most probably referred to theLower Cenomanian.
Inoceramus heinzi Sornay, 1965

PI. 83, Figs. 4a-t\; PI. 85, Figs. 2-3b
1933. Inaequiceramus inaequiualuis (SchlUter);Heinz p. 246, pI. 18, fig. 1.
1965. Inoceramus heinzi Sornay, p. 7, pI. B,
fig. 4.

Material:-GK. H 10148, internal (PI. 83, Fig.
4) and external moulds of left valve from loco T
5046 B; GK. H 10149 (PI. 85, Fig. 2), left valve,with shell attached on the main part, from loco
T 5046 A; GK. H 10150, internal (PI. 85, Fig. 3)
and external moulds of right valve from loco T
5045 B; poorly preserved GK. H 10151, fromloco T 5046 B (I. cf. 1. heinzi); all collected by
T. Matsumoto.
Description :-Shell medium-sized, inequivalve
and inequilaterai. Beak of left valve projectedconsiderably beyond hinge and curved remarkably inward and forward, as seen in GK. H

Tatsuro MATSUMOTO and Masayuki NODA

416

somewhat larger anterior hinge angle, longer
hinge line and larger ratio of L/H than our
described specimens and also than the holotype.
The specimens IGPS. 22709 and UMUT. MM
6492 [=GT. 1-710] from pebbles of the River
Obirashibe, described under Inoceramus sp.
indet. (sp. nov.?) by Nagao and Matsumoto

10148. Valve moderately convex in the umbonalpart, inclined abrupty forward to the almost
vertical and somewhat concave anterior side andgently towards the flattened posterodorsal wing
like part. Convexity along the growth·axis
gradually decreased towards ventral margin.
Outline suboval, extending obliquely towards
the posteroventral extremity, with axis of growthforming angle (8) of about 50 0 with hinge line;
beak anterior, hinge line slightly shorter than ahalf of shell length, forming angle (a) of 7580 0 with anterior margin and that (')') of 1450
to 1500 with the posterior margin; anterior
margin broadly arcuate with gentle an terodorsal concavity; ventral margin asymmetricallyrounded, passing gradually to gently convex,
arcuate posterior margin.
Shell ornamented with rather low concentricmajor ribs or undulations and also concentricrings, the latter of which may not be impressedon internal mould but well discernible on shellsurface or external mould. Posterior muscle
impression is observable slightly behind the
midline in GK. H 10150.
Comparison and discussion :-The described
specimens are very similar to the holotype
illustrated by Heinz (1933, pI. 18, fig. 1) andalso by Sornay (1965, pI. B, fig. 4) of 1. heinzi
Sornay from Madagascar.
The specimens (no. 534 A, B) from Ankomaka, drawn by Sornay (1965, fig. 3), have

(1939, p. 280, pI. 27, fig. 1 and pI. 28, fig. 1),
are somewhat similar to this Sornay's form, as
Kauffman (1977, p. 176) has pointed out, butthey are larger, "broader" and flatter. Probably
they represent a distinct species, but we have
not yet been successful to decide the stratigraphic position of this peculiar form.
Occurrence :--Loc. T 5046 A, B, Saku-gakkono-sawa, lower part of Member II c (clayey mudstone); also loco T 5045 B, close to T 5046,
uppermost part of Member II b (fine-grained
sandy siltstone), Saku area, Teshio Mountains(see route map of Matsumoto and Okada, 1973).
The main part of II b, lower than loco T 5045, isreferred to the Middle Cenomanian on the evidence of ammonites, Desmoceras (Pseudouhligella) japonicum Yabe, Calycoceras spinosum (Kossmat), C. cf. C. bathyomphalum (Kossmat) and Euomphaloceras asura Matsumoto et
Muramoto [= ?E. multicostatum (Basse)], andthe main part of II c, higher than loco T 5046,
is referred to the Lower Turonian on the evidence of Mytiloides mytiloides (Mantell), Fagesia
cf. F. rudra (Stoliczka) and F. thevestensis

Measurements of described specimens.
Specimens

h

l/h

H

L

LiH

b

b/h

sll

a

!3
65°

GK. H 10148

70

65

.93

78

52

.67

19

.27

32.0

.49

85°

GK. H 10150

47

42

.89

55

42

.76

12

.26

19.5

.46

90°

r

0

145°

50°

150°

50°

Explanation of Plate 83
Figs. 1-3. Birostrina tamurai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 411
1. GK. H 10155 (RV) from Kotanbetsu; lateral view, x 1.
2. GK. H 10158 (LV) from Kotanbetsu; lateral (a) and posterior (b) views, x 1.
3. GK. H 10082 (LV) from Kotanbetsu; lateral (a), anterior (b), posterior (c) anddorsal (d) views, x 1.
Fig. 4. Inoceramus heinzi Sornay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 415
GK. H 10148 (LV) from Saku; lateral (a), dorsal (b), posterior (c) and anterior (d)views, x 1.

MATSUMOTO and NODA: Cenomanian Inoceramids from Japan-I

Plate 83

816. Cenomanian Inoceramids from Japan-I

Pervinquiere. Therefore, the beds exposed atlocs. T 5045 and T 5046 are probably referable
to the Upper Cenomanian and/or lowest partof Lower Turonian.
Genus Mytiloides Brongniart, 1822
Type species:-Ostracites labiatus Schlotheim,
1813.
Remarks:-For generic diagnosis and distinction from Inoceramus Sowerby, 1814 readers
may refer to Kauffman et al., 1977 and Keller,
1982, whom we follow.
Mytiloides mikasaensis sp. nov.

PI. 85, Figs. 1a-c
Holotype:-GK. H 10045, composite internalmould of right valve, with test attached partly,collected by Tatsuo Muramoto from loco Ik
1038 on the right bank of the River Ikushumbets, Mikasa district, central Hokkaido.
Specific characters:-Valve gently convex
from beak to ventral margin and also from front
to rear, inclined fairly steeply on anterior sideand gently to posteior, passing to rather narrow
flat wing. Hinge plate thin. Hinge line rather
short, a little more than a third of shell length.
Beak nearly but not exactly at the anterior endof hinge line, suberect, projecting slightly.
Shell extended obliquely towards the posteroventral extremity, with H fairly longer than L
and with axis of growth conyexly curved forward, showing angle (0) of 45° to 35° with hingeline. Anterior hinge angle (a) and angle of umbonal inflation (~) large. Anterior margin convex,
with forward arcuate anteroventral margin, passing gradually to broadly and asymmetricallycurved ventral margin, and then curved rather
abruptly at the posteroventral extremity togently arcuate, long, posterior margin. Posterior
hinge angle (r) very large in late growth-stage.
Shell ornamented with regular concentric417

ribs, which are of moderate intensity in the
middle growth-stage and gradually weakened
later; surface with fine and regular concentric
Iirae, which may be somewhat prominent on thesummit of concentric ribs.
Test very thin. Faint radial striae, which may
be tracks of mantle retractor muscles, discernible
on the main part of internal mould, but for anarrow zone along the growth axis, which may
be posterior adductor muscle insertion area.
Remarks:-Fragmentary shells with regularconcentric ribs are contained in the calcareous
nodule together with the holotype, but they are
too incomplete to show the entire outline.
Comparison :-This species is distinguishedfrom Mytiloides submytiloides (Seitz, 1935)(Holotype; Seitz, 1935, p. 444, pI. 37, fig. 1,
text-fig. 8a), from the Upper Cenomanian andlowest Turonian of Europe and North America(Wiedmann and Kauffman, 1985), in its larger
anterior hinge angle (a) and larger umbonal
angle (~) more convex and forward projecting
anterior margin, and more regular concentricribbing combined with finer and more regularconcentric lirae.
M. mikasaensis is similar in shell-form to M.
goppelnensis (BadiIIet et Sornay, 1980) (Keller,
1982), from the Lower Turonian of Europe andNorth America, especially to the form describedunder M. goppelnensis tourtenayensis (Badillet
et Somay, 1980) from France, but the former
has less arcuate growth-axis and more regularand less crowded concentric ribs combined moreregularly with finer lirae.
For so~e reasons, Seitz (1935), Kauffman(in Kauffman et al., 1977 and other papers) and
Troger (1981) seem to have misunderstood
Inoceramus opalensis Bose, 1923 in the original
sense, which is represented by Bose (1923, pI.
13, figs. 1-3) and never so oblique as the socalled M. opalensis by them. This has beenrevised by BadiIIet and Somay (1980), to whose
opinion we are quite agreeable. W. A. Cobban

Measurements of holotype.
Specimen
GK. H 10045

h
41.0

51.0

I/h

H

L

L/H

b

b/h

1.24

51.8

37.6

.73

11.0

.27

19.0

5/1

a

.37

140

~
0

130

5

'Y
0

155

0

45

0

35 0

418

Tatsuro MATSUMOTO and Masayuki NODA

(1983, personal communication) told us thatexamples of correctly identified M. opalensis
occur near the boundary of Lower and Middle
Turonian, i.e. near the boundary of Mammites
nodosoides Zone and Collignoniceras woollgari
Zone.
Mytiloides modeliaensis (Sornay, 1981), fromthe Lower Turonian of Colombia and North
America, is similar to M. mikasaensis with respect
to the shell-form, especially to the convex anterior margin, but the former has on the averageless oblique growth-axis and more crowded
and sharper concentric ribs.
Occurrence:-Loc. Ik 1038, right bank of themain stream of the River Ikushumbets, Zone of
Euomphaloceras septemseriatum, Upper Cenomanian of the Mikasa district, central Hokkaido(see Matsumoto, 1965, figs. 2 and 4 for the location ofIk 1038).
Summary
As a summary the species described in thispaper are listed below with indication of theirknown age and area.
Birostrina tamurai sp. nov., Middle Cenomanian of the Mifune (central Kyushu),
Mikasa (central Hokkaido) and Kotanbetsu(northwestern Hokkaido) areas
Inoceramus takahashii sp. nov., Lower Cenomanian of the Mikasa area
Inoceramus heinzi Sornay, Upper Cenomanianand (?) Lower Turonian of the Saku area
(northwestern Hokkaido)
Mytiloides mikasaensis sp. nov., Upper Cenomanian of the Mikasa area.
We hope that more examples of these species
would be found from other localities in the
future, so that their biostratigraphic and biogeographic implications may become clearer.

Acknowledgements
We wish to thank Professors Minoru Tamuraand Hakuyu Okada and Messrs. Takemi Takahashi, Tatsuo Muramoto and Kikuwo Muramoto
for their kind help to this study, including the
supply of valuable specimens. Thanks are extended to Professors Itaru Hayami of the University of Tokyo and Tamio Kotaka of Tohoku
University, who gave us facilities to study there
some type specimens in their charge, and also toDrs. W. A. Cobban, E. G. Kauffman and Jacques
Sornay who gave us good suggestions.
References cited
Badillet, G. and Sornay, J. (1980): Sur quelques
formes du groupe d'Inoceramus labiatus
decrites par O. Seitz. ImpossibiIite d'utiliser
ce groupe pour une datation stratigraphiquedu inferieur du Saumurois (France). C.R.
Acad. Sci. Paris, vol. 290 D, p. 323-325.
Bose, E. (1923): Algunas faunas cretacicas deZacutecas, Durango y Guerrero. Inst. Gesl.
Mexico, Bol., vol. 42, p. 1-219, pIs. 1-19.
Cox, L. R. (1969): Inoceramidae. In Moore, R.
C. (ed.), Treatise on invertebrate paleontology, Part N, Mollusca 6, p. N 314-N 321,
Geol. Soc. Amer. & Univ. Kansas Press.
Cragin, F. W. (1892): A contribution to theinvertebrate paleontology of the Texas
Cretaceous. Geol. Surv. Texas, 4th Ann.
Rep., no. 2 (inaccessible).
Dhondt, A. V. (1983): Campanian and Maastrichtian inoceramids: a review. Zittliana,
vol. 10, p. 689-701.
Dobrov, S. A. and Pavlova, M. M. (1959): Inoceramidae. In Moskvin, M. M. (ed.), Atlasof the Upper Cretaceous fauna of Northern
Caucasus and Crimea. Trudy vses. nauchnoissled. Inst. prir. Gazov, p. 130-165, pIs. 123 (in Russian).

Explanation of Plate 84
Fig. 1. Inoceramus takahashii sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 414
Holotype, GK. H 10145 (BV) from Mikasa; right (a) and left (b) lateral and anterior(c) views, x 1.

MATSUMOTO and NODA: Cenomanian Inoceramids tram Japan-I

Plate 84

816. Cenomanian Inoceramids from Japan-I
Efremova, v. I. (1978): On methods and unification of measurements of morphologicalelements of inoceram tests. In Pergament,
M. A. (ed.), Jurassic and Cretaceoous Inocerams and their Stratigraphic Importance,
p. 99-104, tables, Acad. Sci. USSR (inRussian).
Eigenheer, R. and Sornay, J. (1974): Sur une
forme apparenteee a Inoceramus comancheanus Cragin dans Ie Cenomanien basal deMontlaux (Alpes-de-Provence). Bull. Mus.
Natn. d'Rist. Nat., 3 ser., no. 229, p. 141144.
Etheridge, R. (1872). Description of the Paleozoic and Mesozoic fossils of Queensland.
Quart. Jour. Geol. Soc. London, vol. 28, p.
317-350, pIs. 14-25.
Heinz, R. (1933): Inoceramen von Madagascar
und ihre Bedeutung fiir die Kreide-Stratigraphie. Zeits. Deut. Geol. Ges., vol. 85, p.
241-259, pIs. 16-22.
Hill, R. T. (1901): Geography and geology of theBlack and Grand Prairies, Texas, with detailed description of the Cretaceous formations and special reference to artesian
waters. Rep. U.S. Geol. Surv., vol. 21, no. 7,
666 p., 71 pIs. (inaccessible).
Kauffman, E. G. (1977): Systematic, biostratigraphic, and biogeographic relationships between Middle Cretaceous Euramerican andnorth Pacific Inoceramidae. Palaeont. Soc.
Japan Spec. Pap., no. 21, p. 169-212.
- - , Hattin, D. E. and Powell, J. D. (1977):
Stratigraphic, paleontologic and paeoenvironmental analysis of the Upper Cretaceous
rocks of Cimarron County, northwestern
Oklahoma. Mem. Geol. Soc. Amer., vol. 149,
150 p., 12 pIs.
Keller, S. (1982): Die Oberkreide der Sack-Mulde
bei Alfeld (Cenoman-Unter-Coniac). Lithologie, Biostratigraphie und Inoceramen.
Geol. Jahrb., Reihe A, Heft 64, 154 p., 8
pIs.
Ludbrook, N. H. (1978): Australia. In Moullade,M. and Nairn, A. E. M. (eds.), The Phanerozoic Geology of the World II The Mesozoic,
A, Chapt. 7, p. 209-249, Elsevier, Amsterdam.
Matsumoto, T. (1965): A monograph of the
Collignoniceratidae from Hokkaido. Part I.
Mem. Fac. Sci., Kyushu Univ., Ser. D, vol.

419

16, no. 1, p. 1-80, pIs. 1-18.
--and Noda, M. (1985): A new inoceramid(Bivalvia) species from the Upper Campanian(Cretaceous) of Hokkaido. Proc. Japan
Acad., vol. 61, ser. B, p. 1-3.
--and Okada, H. (1973): Saku Formation of
the Yezo geosyncline. Sci. Repts. Dept.
Geol., Kyushu Univ., vol. 11, no. 2, p. 275309 (in Japanese with English abstract).
Nagao, T. and Matsumoto, T. (1939): A monograph of the Cretaceous Inoceramus ofJapan. Part I. Jour. Fac. Sci., Rokkaido
Imp. Univ., Ser. 4, vol. 4, p. 241-299,
pIs. 23-34.
Noda, M. (1983): Notes on the so-called Inoceramus japonicus (Bivalvia) from theUpper Cretaceous of Japan. Trans. Proc.
Palaeont. Soc. Japan, N.S., no. 132, p. 191219, pIs. 41-46.
- - (1984): Notes on My tiloides incertus (Cretaceous Bivalvia) from the Upper Turonian of
the Pombets area, central Hokkaido. Ibid.,
no. 136, p. 455-473, pIs. 84-86.
--and Matsumoto, T. (1976): Mesozoic molluscan fossils of Japan 4 (Certaceous Inoceramus). In Matsumoto, T. (ed.), Atlasof Japanese Fossils, no. 45, Cr 31-Gr 36,
Tsukiji Shokan, Tokyo.
Pergament, M. A. (1966): Zonal stratigraphy
and inocerams of the lowermost Upper
Cretaceous of the Pacific coast of the
USSR. Trans. Acad. Sci. USSR, Geol. Inst.,
vol. 146, 83 p., 36 pIs. (in Russian).
Reeside, J. B. (1923): The fauna of the so-called
Dakota Formation of northern central
Colorado and its equivalent in southeastern
Wyoming. U.S. Geol. Surv. Prof. Pap., no.
131 H, p. 199-207, pIs. 45-50.
Seitz, O. (1935): Die Variabilitiit des Inoceramus
labiatus v. Schloth. Jahrb. Preuss. Geol.
Landesan., vol. 55 (for 1934), p. 429-474,
pIs. 36-40.
--(1961): Die Inoceramen des Santon von
Nordwestdeutschland. I Teil. Beih. Geol.
Jahrb., Heft 47,186 p., 15 pIs.
--(1965): Die Inoceramen des Santon und
Unter-Campan von Nordwestdutschland. II.
Ibid., Heft 69, 194 p., 26 pIs.
--(1967): Die Inoceramen des Santon und
Unter-Campan von Nordwestdeutschland.
III. Ibid., Heft 75, 171 p., 27 pIs.

420

Tatsuro MATSUMOTO and Masayuki NODA--(1970): tiber einige Inoceramen aus derOberen Kreide. Ibid., Heft 86, 171 p., 28
pis.
Sornay, J. (1965): La faune d'inocerames duCenomanien et du Turonien inferieur dusud-oest de Madagascar.. Ann. PaIeont.,
Invertebres, vol. 51, p. 1-18, pis. A-G.
--(1966): Idees actuelles sur les inocerames
d 'apres divers travaux recents. Ibid., Invertebres, vol. 52, p. 59-92.
--(1981): Inocerames (Bivalvia) du Turonien
inferieur de Colombie (Amerique du Sud).
Ibid., Invertebres, vol. 67, p. 135-148, pis.
1-2.
Stephenson, L. W. (1953): Larger invertebrate
fossils of the Woodbine Formation (Cenomanian) of Texas. U.S. Geol. Surv. Prof.
Pap .• 242 (for 1952), p. 1-226, pIs. 1-59.
Tamura, M. (1979): Cenomanian bivalves from
the Mifune Group, Japan. Part 3. Mem. Fac.
Educ. Kumamoto Univ .• no. 28, Nat. Sci.,
p. 59-74, pIs. 1-3.
--and Matsumura, M. (1974): On the age ofthe Mifune Group, central Kyushu, Japan,
with a description of ammonite from thegroup by T. Matsumoto. Ibid .• vol. 23, no.
1, p. 47-56, pI. 1.
Trager, K.-A. (1967): Zur Paliiontologie, Biostratigraphie und faziellen Ausbildung derunteren Oberkreide (Cenoman bis Turon).
Teil I Paliiontologie und Biostratigraphie
der Inoceramen des Cenoman bis Turons
Mitteleuropas. Abh. Staatl. Mus. Min. Geol.•
vol. 12, p. 13-207 (incl. 14 pis.).
Vokes, H. E. (1980): Genera of the Bivalvia:
A systematic and biostratigraphic catalogue(Revised and updated), 307 p., Palaeonto.
Res. Inst., Itahaca, N.Y.
Wiedmann, J. and Kauffman, E. G. (1978): MidCretaceous biostratigraphy of northern
Spain. Ann. Mus. d'Hist. Nat. Nice, vol. 4(for 1976), III, p. 1-84 (incl. 12 pIs.).
Woods, H. (1911): A monograph of the Cretaceous Lamellibranchia of England. Vol. 2,
pt. 7. Palaeontogr. Soc .• 1910, p. 261-284,
pis. 45-50.
--(1917): Certaceous faunas from the northeast of South Island, New Zealand. N.Z.
Geol. Suv. Paleont. Bull., no. 4, p. 1-41,
pis. 1-20.

Asanoyabu T:\1;O)ii. Hachi-no-sawa i\O)iR. Ikushumbets [IkushunbetsuJ ~lF5.m. Kami-ichiKumamoto
Mifune f,l.jJ~H.
no-sawa J::-O)iR. Kiritachi ~:ll:. Kotanbetsu i1:f'HJI].
Mikasa :=:~. Obirashibe 'P¥Ii. Okadake I8iJffi. Omine
Pombets [PonbetsuJ 9"f-}]JJ.
Saku f.£~. Saku-gakko-no-sawa fti:~'¥=t.lCO)iR. Shimomashiki Tt.d:l1iX;, Subayashi !Itt"" Teshio;'klji, Toyono-mura ~!ilfN, Yaseto i\ifjip.

**.

Jm*.

Explanation of Plate 85
Fig.1. Mytiloides mikasaensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 412
Hoiotype, GK. H 10045 (RV), from Mikasa; lateral (a), anterior (b) and posterodorsal views, x 1. Fragmentary left valve is attached on the opposite side.
Figs. 2-3. Inoceramus heinzi Sornay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 415
2. GK. H 10149 (LV) from Saku; lateral view, x 1.
3. GK. H 10150 (RV) from Saku; lateral (a) and anterior (b) views, xL
Figs. 4-5. Birostrina cf. B. tamurai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 411
4. KE 1977 (LV), incomplete, secondarily compressed, large example (internalmould), x 1.
5. Rubber-cast from the external mould (posteroventral portion) of KE 1977, x 1.

MA TSUlvIOTO and NODA: Cenomanian Inoceramids tram Japan-I

Plate 85

816. Cenomanian Inoceramids from Japan-I

421*fBl~r~*-l! / ""7 =- 7 :/1\M'E[1' / -l! '7 A , A - I. ~tifij.ii! CfL1'/iE[0) J: <:m I?.h -c ~ ,t.eli' -:,f-:Hi: *:f!S0) -l! / ""7 =- 7 :/li' I?ll, t. tf. J: <~~~.h -c ~,t.eli' "? t-:1'jli t li' t.e IJ il!> GO) ,"(", k-.h
I? ~ ;i¥/xIl~JliJn... -0' <• ;: 0) iffiljx'"("Il/X 0) 4 fi~IlG$lt L t-:. (1) Birostrina tamurai sp. nov. ;tJlitrl':' Inoceramus concentricus costatus Nagao et Matsumoto c Iif-'i.h -C ~ ,t-: ~ 0)0)-$'"(", B. concentrica ~I.:.~-t Glit, Wl:0)~$1.:.1tdlt~O)Mlitil!> IJ, hX:~f~MI.:.l!1Jlit;e~-t Go fL1+1 O)~ti1\1lrff$~ Utifij)1HIH~~5]IJN.O'itft5]JjO) -l! / ""7 =- 7 :/ cp$E[o (2) Inocera-

mus takahashii sp. nov.; I. comanchean us.7,..li~~~<'~I.:.~-r G li~, Wl:O)~!1l\1.J~l(t~1.i}f;'"(", IJ~I?

!lli1!1J~~'!I~'o ~Ui~5]IJo)-l!/""7=-7:/T$E[,

J"1Ji(E[O)~O)I':' i'1J:<E'"("~Go(3) I. heinzi Sornay;

7(li!f!i:~0)-l! / ""7=-7:/·7-,,- -""7!J"-j/,A'j]=-7 :/!f[Hi1IE[. (4) Mytilo'ides mikasaensis sp. nov., M. submytiloides (Seitz) J: IJ ~Wl:m1't'lli;*~ <. iiHZo
li~litr1.il.:.80)~H/ti <~~'"(". !IU\lJIE LH'i].C"I!1JI.:.:!JOx. -C *Dili'~ ,1lij.c,'~~litWl:~ITii~fii? ;: c'"("~5]IJ ~.hGo ~~5]IJO) -l! / ""7 =- 7:/ J:$O) Euomphaloceras septemseriatum ;HHCE[-t Go
t~*it!'l~ • !ffm!fftzPTrans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 422-434, pI. 86, September 30,1986

817. LEPIDOLINA COLUMBIANA (PERMIAN FUSULINID)FROM BRITISH COLUMBIA, CANADA*HIROYA GOTO
Institute of Geosciences, College of Liberal Arts, Kobe University; Kobe 657
KUNITAKA MARUOKA
Higashinada High School, Kobe 658
and
KEN·ICHI ISHII
Institute of Geosciences, College of Liberal Arts, Kobe University, Kobe 657

Abstract. Nine species of Yabeina have been reported from British Columbia.
On the basis of statistical study of specimens collected from Marble Canyon andUpper Hat Creek, it is concluded that our specimens as well as specimens of thenine species of "Yabeina" (Y. columbiana, Y. minuta, Y. obesa, Y. dawsoni, Y.
gracilis, Y. dunbari, Y. cylindrica. Y. ampla and Y. paruula) belong to one species
which should be referred to the genus Lepidolina. They are described under the
species L. columbiana (Dawson).

Introduction

Recently Ishii and others (1985) reexamined
distribution of neoschwagerinids in the Tethys.
According to them, Neoschwagerina and Yabeina
are distributed allover the Tethys, while distri·bution of Colania and Lepidolina is restricted
to East Asia including Southeast Asia, China
and Japan, which forms the Paleopacific bio·
geographic province in Middle Permian time.
Our paleontological restudy of "Yabeina"
columbiana from British Columbia has revealed
that this species does not belong to the genus
Ya be ina, but is assignable to the genus Lepi·
dolina. In this paper we intend to present a resultof our study on species of the genus Yabeina
described from Marble Canyon, British Columbiaon the basis of specimens collected by Goto.

It has been suggested that Middle Permian
fusulinids of the Cordilleran region including
British Columbia, Oregon and California resem·
ble Tethyan fusulinids unlike those of the Mid·
Continent, North America (Dunbar, 1932,
Thompson and Wheeler, 1942, Ross, 1967 andothers). With this suggestion in mind, one ofus (Goto) visited the Marble Canyon region
during his stay in British Columbia in 1980
and collected limestone samples containing
many specimens of "Yabeina" columbiana.

*Received April 15, 1985; Revised manuscript
received June 19, 1986: Read January 22, 1984
at Kyoto University.
422817. Lepidolina columbiana from B.C. Canada
Historical reviews:-According to Duffell and
McTaggart (1951), the study of fusulinids ofthe Marble Canyon limestone, British Columbia
was started in 1872 by Selwyn. In 1877 Dawson
revisited Selwyn's original locality in Marble
Canyon and on the basis of the fusulinids
assigned a Carboniferous age to the Marbl~
Canyon limestone. Dawson (1879) described
Loftusia columbiana from Marble Canyon.
Although Loftusia Brady is originally based on
a Tertiary species, Dawson concluded that L.
columbiana was a Carboniferous form rather
than Cretaceous or Eocene because of theaccompanying fusulinids.
Dunbar (1932) restudied Dawson's thin sections of specimens of Loftusia columbiana anddescribed the species under the genus Neoschwagerina. He pointed out that the limestone
at Marble Canyon is Permian in age. Based on a
study of well oriented section of Loftusia columbiana Dawson, Thompson and Wheeler (1942)
pointed out that this species should be referredto the genus Yabeina Deprat. They also describedYabeina minuta, Schwagerina pavilioensis var.
acris, Nankinella? and Staffella in addition to
Yabeina columbiana from samples of talus in
Marble Canyon. Thompson, Wheeler and Danner(1950) redescribed Y. columbiana and Y. minuta
associated with Yabeina ? n. sp. Schwagerina
acris, S. andersoni ? and Codonofusiella duffeli
from the Marble Canyon region. Skinner and
Wilde (1966) described seven new species of thegenus Yabeina listed below in addition to Y.
columbiana, Y. minuta, Schwagerina canadensis
and Chusenella spicata from the same !ocalityof Marble Canyon material. The seven speciesof the genus Yabeina are Yabeina obesa Y
dawsoni, Y. gracilis, Y. dunbari, Y. cylind'rica:
Y. ampla and Y. parvula.
Acknowledgement:-We wish to express ourthanks to Dr. Raymond C. Douglass of U.S.
Geological Survey, Prof. Wilbert R. Danner ofUniversity of British Columbia, Canada andProf. Koichiro Ichikawa of Osaka City University
for their kind advice.

423

Geological setting
Geological mapping of the area including the
Marble Canyon region, British Columbia wasmade by Duffell and McTaggart (1951). Trettin(1980) described Permian Cache Creek Groupof the Marble Range. According to them the
Marble Canyon Formation is a constituent of
Cache Creek Group. The Marble Canyon Formation is composed of the massive and mostly
recrystallized limestone with intercalations of
volcanic rocks, argillites and cherty quartzites,
typically exposed in the Marble Canyon andPavilion Mountains (Text-fig. 1). In the western
limit of the distribution of the limestone inPavilion Mountain, strata are dipping mainly
to the west and southwest at about 60 degrees,
but near the eastern boundary, dips are to thewest at about only 20 degrees. Though therepetition of strata resulting from minor folding and faulting is conspicuous, it is probablethat the limestone has a thickness of about
1800 m. The Marble Canyon Formation isflanked on both sides by chert, argillite, greenstone, minor limestone, quartzite and their
metamorphic equivalents, which are constituents of the Cache Creek Group. The Marble
Canyon Formation yields Permian foraminifers,
crinoids, blastoids, corals and brachiopods,
and this formation has been assigned to the
Guadalupian Formations of the other regions
by many investigators (Duffel and McTaggart,
1951 and Trettin, 1980).
Our samples were collected from the following two localities (Text-fig. 1).
Loc. 1, Marble Canyon: Talus on the northshore of the Pavilion Lake. Massive recrystallized
limestone. Type locality of "Yabeina" columbiana (Dawson). Sample obtained from Loc. 1
are grayish limestone, containing fossils of
fusulinids, smaller foraminifers and algae. The
preservation of these fossils is generally poor,
because of recrystallization of limestone owing
to regional metamorphism.
Loc. 2, Upper Hat Creek: About 30 km southof Loc. 1. Bedded limestone, partly recrystallized. Samples obtained from Loc. 2 are dark

Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII

424

[[[]]
[[I]]] 2

500 45'Nq~1-_ _-_19 km51...!

i!

Text-fig.!. Geological sketch map of the Marble Canyon region. Loc. 1: Marble Canyon, Loc.
2: Upper Hat Creek, 1: Pavilion beds, 2: western belt of Cache Creek Group, 3: central belt of Cache
Creek Group (Marble Canyon Formation), 4: eastern belt of Cache Creek Group. (after Trettin, 1980)
gray limestone showing micritic texture. Only
fusulinids are found as fossils. This locality is
newly found by Goto.
Morphologic variation
Before going over the systematic paleontology, some results of morphological analysisof yabeinid specimens from two localities arepresented in order to examine the scope of
specific variation.
Morphometry of the specimens from Marble
Canyon and Upper Hat Creek is carried out
with respect to the following shell characterssuch as form ratio, diameter of proloculus,
thickness of spirotheca and first appearanceof secondary transverse septula.

First, the comparison of individuals treatedthis time with those of species of Yabeina andLepidolina described previously is made withreference to the form ratio and the diameterof proloculus (Text-fig. 2). The individuals
treated this time and those of nine species of
Yabeina described by Skinner and Wilde fromthe same region are recognized to be situatedon the intermediate position between Lepidolina multiseptata from East Asia (cf. Ozawa,1975), and the Yabeina group including Y.
globosa, Y. packardi, Y. kaizensis, Y. igoi, Y.
katoi, etc. In Text-fig. 2, measured values of
proloculus of the present specimens and thoseof nine species of Yabeina mentioned alreadyare overlapping with that of Lepidolina asiatica
but are not overlapping with that of most of

817. Lepidolina columbiana from B.C. Canada
19. Lepldalino mulliseptata multiseptata

/
6

400

20. L. m. shiraiwensis

E 300

o
o

o

2.0

10

form

rotio

30

4.0

425

Text-fig. 2. Diam eter of proloculus
against form ratio for present specimensfrom Upper Hat Creek (open circle) andMarble Canyon (solid circle). Variation
ranges of species of L epido lina and Yab eina
are shown as vertical bars and squares. 1:"Yabeina" columbiana, 2: "Y." minuta, 3 :"Y." obesa, 4: "Y." dawsoni, 5: "Y."gracilis, 6: "Y." dunbari, 7: " Y . " cylindrica ,8: "Y." ampla, 9: "Y. " parvula, 10: Y.
cascadensis, 11: Y. decora, 12: Y. fu siformis, 13 : Y. paclwrdi, 14: Y. kaize nsis,
being indicated by symbol of black triangle,
15: Y. igoi, 16 : Y. inouy e i, 17: Y. haloi,
18: Y. globosa, 19 : L epidolina multiseptala
muitiseptata, 20: L. m. shiraiwensis, 21: L .
asiatica, 22: L . japonica. Variation rangeof the species of Yab eina (s. str.) falls withinth e part below the broken line .

Text-fig. 3. Comparison of spiroth eca,primary and secondary transverse septula
among L epidolina multiseptala multiseplala ,L. columbiana and Yab einaglobosa. a : L. 111 .
multiseptata, topotype, loc o Phnom Bak,
Sisophon, Cambodia (coll. Ishii), b: L.
columbiana, KU 3002, enlarged part of pI.
fig. 2, loco Marble Canyon , Briti sh Columbi a(colI. Goto) , c: Y. globosa, topotype, loco
Mameishi limeston e, Akasaka, Gifu Prefec ture, Japan (colI. Ishii) .

Hiroya GOTO, Kunitaka MAR UOKA and Ken-Ichi ISHII

426

the Yabeina group. This text-figure shows thatsome specimens, which were referred to "Yabeina" columbiana and "Y." ampla by Skinnerand Wilde, are below 100 microns in diameterof proloculus. However, there is a possibility
that thin sections of these specimens were not
cut through the center of proloculus. Since
diameter of proloculus of representative speciesof Yabeina such as Yabeina globosa, Y. cascadensis, Y. decora, Y. fusiformis, Y. packardi, Y.
kaizensis, Y. igoi, Y. inouyei and Y. katoi ismostly below 100 microns, we regard the genus
Yabeina as fusulinids having the prolocular
diameter of less than 100 microns.
Secondarily, the present specimens andthose of nine species of "Yabeina" from Marble
Canyon have the generic character of Lepidolina
that secondary transverse septula are thin bar
and occasionally pendant club-shape in crosssection (Text·fig. 3). In view of these differences,
Marble Canyon 8 Upper Hat Creek

20

n = 43m = 1.81
en
0

15

:::J

-0

>

n = 29-0C

m= 1.78
10

't-'

0

"-

n = 14Q).0E

m= 1.86

5

:::J
C

o

l

for mral' i l'r";;;:;::~;:::::;"Y."columbiana, "Y. "ampla

I/y' il obesa, "yN dunbari, I'y/lparvu/a

Text-fig. 4. Frequency
distribution
curves of form ratio for the present specimens from Marble Canyon and Upper Hat
Creek. n: measured individual number, m:mean value.

we described that present specimens and thoseof nine species by Skinner and Wilde belong to
the genus Lepidolina.
Next, the frequency distribution curves for
magnitude of form ratio of the specimens fromMarble Canyon and Upper Hat Creek show abell shape (Text-fig. 4). The bell shape in thefrequency distribution curves of specimensfrom Marble Canyon and Upper Hat Creek may
be attributed to that the specimens come froma population. Most of the specimens have a form
ratio of 1.5 to 2.0. The specimens of "Yabeina"
columbiana described by Skinner and Wilde(1966) are within this range of form ratio (Textfig. 4).
Incidentally several morphologic characters
including form ratio, diameter of proloculus,
thickness of spirotheca, first appearance of
secondary transverse septula are said to change
phylogenetically in the subfamily Neoschwagerininae (Ozawa, 1970). For example, the moreadvanced species of the genus Lepidolina whichis included in the Neoschwagerininae havelarger form ratio, larger proloculus, thinnerspirotheca and the earlier appearance in ontogeny of secondary transverse septula. In addition,
the magnitude of form ratio also changes according to change of environment such as the change
in speed of water current and in wave action.
It seems that species of Eopolydiexodina andMonodiexodina having larger form ratio arefound in limestone of higher energy environment such as sparite, while the species of Yabeina and Verbeekina having smaller form ratio
are observed in limestone of low energy environment such as micrite. The larger value of form
ratio in the Marble Canyon specimens is larger
than that of Upper Hat Creek. In view of resembling lithology of limestone in the two
localities, the above mentioned feature may
reflect the phyletic increase in form ratio.
The specimens from both localities are compared with one another by plotting thicknessof spirotheca and volution number on which
secondary transverse septula first appears (Textfigs. 5, 6). On these two text-figures, present
specimens and those of nine species of "Yabe-817. Lepidolina columbiana from B.c. Canada

thickness of spirotheca. in microns

Ip30

first appearance of secondary transverse septu Ia

50

40

I

-1

I

427

volution number
5,

I

-2

-3
-4
5-6

02
04

,

~7

07

-8

-:9
,
~11

""',--15
,
,,

11 0
I

I

!------16

190--0 :20d--o
21 ;

r- 21

20 ,
I

en
Ci

:::J
~

>

:.a

.~

~I

...

<lJ

E
:::J
C

II

I

014

I
I

718
I
I

,I
?22

Upper Hat Creek

'0 10

:1

I I 111111

0

,

~10
012
?13
015

!II II
Upper Hat CreekII

11.1111

10

I

I

20

II

Q)

~ 5

I

I

'-

..0

c
I

II

I

40
30
50
thickness of spirotheca. in microns

0

I

I

Marble Canyon

0

.D

170

I

18

~5

I

:

--~--17

--::i

19

06

08
90

---10

--:12
----13:

,

10

01

~I_

o '------.1...-1

5

Text-fig. 5. Variaton in thickness of
spirotheca of the present specimens (lower
figure) in comparison with that of various
species of Lepidolina and Yabeina (upper
figure). For numbers 1 to 21 see Text-fig. 2.
The measurement values for the present
specimens fall within the range between thedotted lines of the upper figure.

ina" are still situated between the Yabeina
species group and the Lepidolina species group.
Finally, we made an investigation about
allometry on which the half length and theradius vector were adopted as two parameters.
If the two characters of each individual havethe relation of allometry, the following formulais expected between values of these characters

10

volution number
Text-fig. 6. Stage of the first appearance of secondary transverse septula of thepresent specimens (lower figure) in comparison with that of various species of Lepidolina and Yabeina (upper figure). For
numbers 1 to 22 see Text-fig. 2. The measurement values for the present specimens
fall within the range between the dottedlines of the upper figure.
(x, y). Y = bXa , where, a is a relative growthcoefficient and b is a initial growth index. The
formula Y = bXa is transformed to log Y = a log
X + log b. The latter formula is expected to belinear line having the inclination of a.
Text-figs. 7 and 8 show the respective mean

Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII

428

MarbleTable 1.

Canyonform ratio

A 0..1-1.0.
B 1.0.-1.5
C 1.5-2.0.

5.0.
3.0.
E
E

A

D 2.0.- 2.5
E 2.5- 3.0

2.0.

D

c

-

1.0.

II'>

0..5

(;

Relative growth coefficients (a)for the present specimens, Lepidolina multiseptata multiseptata,
L. m. shiraiwensis, L. asiatica(after Ozawa, 1975) and Yabeina
globosa (after Hanzawa and Murata,1963).

E

Form ratioaMarble Canyon

1.0-1.5
1.5-2.0
2.0-2.5
2.5-3.0
3.0-3.5

1.02
0.82
0.76
0.79
0.69

Upper Hat Creek

0.5-1.0
1.0-1.5
1.5-2.0
2.0-2.5
2.5-3.0

0.99
0.76
0.85
0.85
0.79

u

Q)

>

~

'0

~

0..3
0..2

0.1

0.2

0.3

0..5

2.0. 3.0

1.0.

5.0

half length, in mm

Text-fig. 7. Growth curves of thepresent specimens from Marble Canyon.
Line A is a mean curve of specimens whose
form ratio are within 1.0 to 1.5, and soforth.

Lepidolina m. multiseptata
L. m. shiraiwensis
L. asiatica
Yabeina globosa

0.63
0.77
0.82
0.98

Upper Hat Creek
farm ratia
5.0.

A 0.1- 1.0.
B 1.0.- 1.5

3.0.
E
E
c

...0

u

2.0.

C 1.5- 2.0.
D 2.0.- 2.5
E 2.5- 3.0.

A
C

D
E

1.0.

Q)

>

II'>
~

0..5

'0

... 0.3
0

0..2

0.1

0.2

0.3

0..5

1.0.

2.0. 3.0.

5.0.

half length, in mm

Text-fig. 8. Growth curves of thepresent specimens from Upper Hat Creek.
For line A to E see Tex t-fig. 7.

values which were calculated for all individuals
classified by range of form ratio in each locality.

Table 1 demonstrates the relative growth coefficients - that is inclination of a - of therespectively classified groups, which is led byapplying method of least squares. As seen from
Table 1, all. the individuals from Marble Canyon
and Upper Hat Creek are within the range ofabout 0.7 to 1.0. It suggests that these twogroups would belong to the same species. Moreover, they differ from those of Lepidolina m.
multiseptata, while they are difficult to be distinguished from L. asiatica, L. m. shiraiwensis
and Yabeina globosa by the relative growthcoefficient only. However, they can be distinguished from specimens of the latter two species
on basis of prolocular diameter.
In conclusion, all specimens from Marble
Canyon and Upper Hat Creek treated this time
as well as those of nine species of "Yabeina"
described by Skinner and Wilde are regarded
to belong to the same species. They belong
undoubtedly to the genus Lepidolina and not
to Yabeina. They are described under the species
Lepidolina columbiana (Dawson).

817. Lepidolina columbiana from B.C. Canada

429

Table 2. Measurement of Lepidolina columbiana (Dawson)

KU 3001, from Marble Canyon, PI. 86, Fig. 1.
vn

1

2

3

4

5

6

7

8

9

10

11

12

13

hI

0.18 0.30 0.40 0.50 0.65 0.85 1.00 1.20 1.45 1.68 1.91 2.18 2.53

rv

0.25 0.35 0.41 0.50 0.64 0.79 1.00 1.15 1.45 1.65 1.88 2.13

fr

0.72 0.85 0.97 1.00 1.00 1.08 1.00 1.04 1.00 1.02 1.02 1.02

14

15

Length: 5.08 mm, width: 4.45 mm
Thickness of spirotheca: 30 microns
Diameter of proloculus: 250 microns
First appearance of sts: 5th volution
vn: volution number, hI: half length (mm), rv: radius vector (mm), fr: form ratio,
sts: secondary transverse septula. Note: Thickness of spirotheca is measured at 10th
volu tion on all specimens.

KU 3002, from Marble Canyon, PI. 86, Fig. 2.
vn

1

hI

0.28 0.34 0.45 0.66 0.91 1.20 1.65 2.00 2.40 2.79

rv

0.20 0.30 0.40 0.50 0.65 0.85 1.05 1.20 1.43 1.65

fr

1.38 1.13 1.11 1.31 1.39 1.41 1.57 1.67 1.68 1.69

2

3

4

5

6

7

8

9

10

11

12

13

14

15

11

12

13

14

15

Length: 5.70 mm, width: 3.30 mm
Thickness of spirotheca: 12 microns
Diameter of proloculus: 302 microns
First appearance of sts: 4th volu tion

KU 3003, from Marble Canyon, PI. 86, Fig. 3.
4

5

6

7

8

9

10

vn

1

hI

0.29 0.48 0.65 0.83 1.20 1.50 1.85 2.40 2.83 3.15 3.50 3.80

rv

0.24 0.33 0.40 0.45 0.53 0.60 0.69 0.80 0.98 1.23 1.43

fr

1.19 1.46 1.63 1.83 2.29 2.50 2.68 3.00 2.90 2.57 2.45

23Length:' 7.40 mm, width: 3.10 mm
Thickness of spirotheca: 28 microns
Diameter of proloculus: 320 microns
First appearance of sts: 6th volution

Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII430

KU 3004, from Marble Canyon, PI. 86, Fig. 4.
vn

1

hI

0.26 0.42 0.65 0.95 1.20 1.58 1.95 2.53 2.95 3.60 4.37 4.93

rv

0.15 0.29 0.33 0.43 0.50 0.65 0.81 0.96 1.13 1.30 1.50 1.66

fr

1.79 1.45

2

3

~.OO

4

5

6

7

8

9

10

11

12

13

14

15

13

14

15

14

15

14

15

2.24 2.40 2.42 2.41 2.63 2.62 2.77 2.91 2.97

Length: 10.5mm, width: 3.60 mm
Thickness of spirotheca: 25 microns
Diameter of proloculus: 250 microns
First appearance of sts: 6th volution

KU 3005, from Marble Canyon, PI. 86, Fig. 5.
2

3

4

5

6

7

8

9

10

11

12

vn

1

hI

0.12 0.29 0.49 0.83 1.10 1.35 1.63 1.80 2.36 2.65 3.30 3.60 3.93

rv

0.10 0.19 0.23 0.28 0.35 0.44 0.49 0.58 0.65 0.77 0.91 1.02 1.20

fr

1.20 1.53 2.16 3.00 3.14 3.10 3.36 3.13 3.62 3.44 3.65 3.53 3.28
Length: 7.30 mm, width: 2.57 mm
Thickness of spiro theca : 15 microns
Diameter of proloculus: 123 microns
First appearance of sts: 6th volution

KU 3006, from Upper Hat Creek, PI. 86, Fig. 6.
vn

1

hI

0.10 0.19 0.28 0.35 0.52 0.62 0.80 0.97 1.09 1.20 1.30 1.43 1.53

rv

0.13 0.20 0.35 0.45 0.59 0.82 0.95 1.15 1.25 1.45 1.58 1.75 1.90

fr

0.77 0.93 0.79 0.78 0.88 0.75 0.84 0.84 0.87 0.83 0.83 0.81 0.80

2

3

4

5

6

7

8

9

10

11

12

13

Length: 3.10 mm, width: 4.00 mm
Thickness of spirotheca: 28 microns
Diameter of proloculus: 121 microns
First appearance of sts: 6th volution

KU 3007, from Upper Hat Creek, PI. 86, Fig. 7.
vn

1

hI

0.07 0.19 0.25 0.34 0.43 0.58 0.74 0.94 1.20 1.45 1.70 1.98 2.25 2.53 2.80

rv

0.09 0.16 0.25 0.29 0.45 0.57 0.70 0.90 1.03 1.35 1.66 1.90 2.15 2.41 2.80

fr

0.78 1.23 1.00 1.16 0.96 1.02 1.05 1.04 1.17 1.07 1.02 1.04 1.05 1.05 1.04

2

3

4

5

6

Length: 5.60? mm, width: 5.35 mm
Thickness of spirotheca: 26 microns
Diameter of proloculus: 144 microns
First appearance of sts: 8th volution

7

8

9

10

11

12

13

817. Lepidolina columbiana from B.C. Canada

431

KU 3008, from Upper Hat Creek, PI. 86, Fig. 8.
vn

1

2

3

4

5

6

7

hi

0.25 0.35 0.54 0.85 1.15 1.35 1.60 1.85 2.15 2.45 2.75 3.28

rv

0.16 0.27 0.38 0.48 0.60 0.70 0.85 101 1.16 1.32 1.47 1.66

fr

1.56 1.30 1.43 1.79 1.92 1.93 1.88 1.84 1.85 1.86 1.86 1.98

8

9

10

11

12

13

14

15

13

14

15

13

14

15

Length: 6.67 mm, width: 3.10 mm
Thickness of spirotheca: 25 microns
Diameter of proloculus: 260 microns
First appearance of sts: 8th volution

KU 3009, from Upper Hat Creek, PI. 86, Fig. 9.
vn

1

hi

0.17 0.30 0.43 0.60 0.85 1.10 1.39 1.98 2.32 2.55 3.00

2

3

4

5

6

7

8

9

10

11

12

rv

0.15 0.19 0.23 0.30 0.40 0.51 0.66 0.80 0.95 1.10 1.23 1.45

fr

1.13 1.58 1.87 2.00 2.13 2.17 2.10 2.47 2.44 2.32 2.44
Length: 6.20 mm, width: 2.60 mm
Thickness of spirotheca: 20 microns
Diameter of proloculus: 200 microns
First appearance of sts: 8th volution

KU 3010, from Upper Hat Creek, PI. 86, Fig. 10.
2

3

4

5

6

7

8

9

10

11

12

vn

1

hi

0.28 0.40 0.61 0.95 1.17 1.61 1.94 2.50 3.00 3.55 3.95 4.40 5.00

rv

0.15 0.23 0.31 0.43 0.55 0.60 0.83 0.98 1.10 1.25 1.45 1.60 1.76

fr

1.83 1.78 1.97 2.21 2.12 2.68 2.35 2.65 2.73 2.84 2.72 2.75 2.85
Length: 10.00? mm, width: 3.65 mm
Thickness of spirotheca: 21 microns
Diameter of proloculus: 214 microns
First appearance of sts: 7th volution

Systematic paleontology
Family Verbeekinidae Staff and Wedekind, 1910
Subfamily Neoschwagerininae
Dunbar and Condra, 1928
Genus Lepidolina Lee, 1933

Lepidolina columbiana (Dawson)
PI. 86, Figs. 1-10.

Loftusia colum biana Dawson, 1879, p. 69-75,
pI. 6, figs. 1-7; Silvestri, 1932, p. 87, pI. 2,
figs. 4-7.
Neoschwagerina columbiana, Dunbar, 1932, p.
46-48, pI. 1, figs. 1-4.
Yabeina minuta Thompson and Wheeler, 1942,
p. 707-708, pI. 106, figs. 6-10; Thompson,
Wheeler and Danner, 1950, p. 62, pI. 8, figs.
4-7; Ishii and Nogami, 1964, p. 22, pI. 4,
figs. 4-7; Skinner and Wilde, 1966, p. 309-

432

Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII

311, pI. 34, figs. 3-7; Yang, 1978, p. 120,
pI. 34, figs. 1-2.
Yabeina columbiana, Thompson and Wheeler,
1942, p. 708-710, pI. 106, fig. 5, pI. 107,
fig. 5, pI. 108, fig. 1, pI. 109, figs. 1-4;
Thompson, Wheeler and Danner, 1950, p.
61, pI. 8, figs. 1-3; Kanmera, 1954, p. 1618, pI. 3, figs. 1-5, 7, questionable specimens,
pI. 3, fig. 6; Nogami, 1958, p. 101-102, pI. 1,
figs. 9-10; Chisaka, 1960, p. 249-250, pI.
7, fig. 3, p. 54, fig. 18c; Skinner and Wilde,
1966, p. 47-48, pI. 38, figs. 1-6; Yamagiwa
and Saka, 1972, p. 268-269, p. 31, fig. 4;
Zang, 1982, questionable specimens, p. 209,
pI. 34, figs. 3, 7.
Yabeina obesa Skinner and Wilde, 1966, p. 50,
pI. 40, figs. 1---7.
Yabeina dawsoni Skinner and Wilde, 1966, p.
50, pI. 41, figs. 1-7.
Yabeina gracilis Skinner and Wilde, 1966, p. 5051, pI. 42, figs. 1-9.
Yabeina dunbari Skinner and Wilde, 1966, p.
51-52, pI. 43, figs. 1,.
Yabeina cylindrica Skinner and Wilde, 1966, p.
52, pI. 44, figs. 1-7.
Yabeina ampla Skinner and Wilde, 1966, p. 5253, pI. 45, figs. 1-4, pI. 46, fig. l.
Yabeina parvula Skinner and Wilde, 1966, p.
53-54, pI. 46, figs. 2-11, pI. 47, figs. 1-2.

Description:-Shell large, inflated fusiform toelongate fusiform in. shape, with straight toslightly curving axis of coiling, gently convex
to concave lateral slopes. Mature specimens of
11 to 13 volutions or more. 5.0 to 10.0 mm
long, and 2.6 to 5.4 mm wide, form ratios of
0.8? to 3.3
Outside diameter of proloculus 100 to 330
microns, mostly 150 to 250 microns. Spirotheca composed of a tectum and a keriothecawith very fine alveoli. Thickness of spirotheca
10 to 30 microns in outer volutions. Septa
and primary transverse septula present throughout shell and essentially slender. Axial septulathin, first appear in 4th to 5th volution. Secondary transverse septula first appear in 4th to 8th
volution, mostly in 5th to 7th volution, usually
one septulum between adjacent primary transverse septula throughout shell, but two septulain most outer volutions.

Measurement:-See Table 2.
Remarks:-Morphologic variation within thespecies is given in detail in the preceding chapter.
The present species differs from Lepidolina
multiseptata multiseptata and L. m. shiraiwensis
in having rather smaller proloculus. Moreoverthe secondary transverse septula appear inslightly later stage and the thickness of spirotheca is larger in the former than in the latters.
The secondary transverse septula of thepresent species appear in earlier stage than thoseof L. asiatica. Although measured values of sizeof proloculus and form ratio of the present
species overlap those of L. asiatica, these valuesof L. asiatica are concentrated on the end of themorphometrical variation of L. columbiana(Text-fig. 2).
The present species resembles L. japonica(Huzimoto, 1936) in size of proloculus, but theformer has generally a smaller form ratio thanthe latter (Text-fig. 2), so far as Huzimoto's
original description of L. japonica is concerned.
Occurrence:-The specimens at hand havebeen collected from the type locality, Marble
Canyon, and from Upper Hat Creek, British
Columbia, Canada (Marble Canyon Formationof the Cache Creek Group). They occur together
with many specimens of Schwagerina in the two
localities, and also with specimens of Colania
in Upper Hat Creek.
Materials:-Reg. no. KU 3001, KU 3002, KU
3003, KU 3004 and KU 3005 from Marble
Canyon. KU 3006, KU 3007, KU 3008, KU
3009 and KU 3010 form Upper Hat Creek. Thesematerials are collected by H. Goto, and aredeposited in Institute of Geosciences, College of
Liberal Arts, Kobe University.

References
Chisaka, T. (1960): On some Permian Fusulinids from the Takagami Conglomerate,
Choshi Peninsula, Chiba Prefecture, Japan.
Jour. Call. Arts and Sci. Chiba Univ., vol. 3,
no. 2, p. 235-254, pIs. 1-9.
Dawson, G. M. (1879): On the new species of
Loftusia from British Columbia. Geol. Soc.

817. Lepidolina columbiana from B.C. Canada
London Quart. Jour., vol. 35, p. 69-75, pI.
6.
Duffel, S. and McTaggart, K. C. (1951): Ashcroft
Map - Area, British Columbia. Ceol. Suru.
Canada, Mem. 262.
Dunbar, C. O. (1932): Neoschwagerina in the
Permian faunas of British Columbia. Royal
Soc. Canada, Proc. and Trans., 3rd ser., sec.
4,vol. 26, p. 45-49, pI. 1.
Geological Institute of Southwest China (ed.).
Yang (1978): Paleontological monograph ofSouthwest China. Part 2 (Sichuan). Ceol.
[nst. Southwest China, Pub., p. 1-684, pis.
1-191 (in Chinese).
Hanzawa, S. and Murata, M. (1963): The paleontologic and stratigraphic considerations on
the Neoschwagerininae and Verbeekininae,
with the descriptions of some fusulinid
foraminifera from the Kitakami Massif,
Japan. Sci. Rep. Tohoku Uniu. (Ceology),
vol. 35, no. I, p. 1-31, pis. 1-20.
Huzimoto, H. (1936): Stratigraphical and palaeontological studies of the Titibu System of
the Kwanto-mountainland, Part 2. Palaeontology. Sci. Rep. Tokyo Bunrika Daigaku,
sec. C, vol. I, no. 2, p. 29-125, pis. 1-26.
Ishii, K. and Nogami, Y. (1964): Contributions
to the geology and paleontology of Cambodia. Part 1. Permian fusulinids. Jour.
Ceosci., Osaka City Uniu., vol. 8, Art. 2,
p. 9-70, pis. 1-8.
- , Okimura, Y. and Ichikawa, K. (1985): Notes
on Tethys biogeography with reference to
Middle Permian fusulinaceans. In Nakazawa,
K. and Dickins, J. M., (eds.), The Tethys,
her paleogeography and paleobiogeography
from Paleozoic to Mesozoic, p. 139-155,
Tokai Uniu. Press.
Kanmera, K. (1954): Fusulinids from the Upper
Permian Kuma Formation, Southern Kyushu, Japan, with special reference to thefusulinid zone in the Upper Permian ofJapan. Mem. Fac. Sci. Kyushu Uniu., Ser. D,
Ceol., vol. 4, no. 1, p. 1-38, pis. 1-6.
Morikawa, R. and Suzuki, Y. (1961): Fusulinids
form the Akasaka Limestone (Part 2). Sci.
Rep. Saitama Uniu., ser. B, vol. B, no. 1, p.
43-74, pis. 4-22.
Nogami, Y. (1958): Fusulinids from the Maizuru
Zone, Southwest Japan. Part 1. Ozawainellinae, Schu bertellinae and N eosch wagerininae. Mem. Coil. Sci. Kyoto Uniu., ser. B,

433

vol. 25, no. 2, p. 97-114, pis. 1-2.
Ozawa, T. (1970): Notes on the phylogeny andclassification of the Superfamily Verbeekinidea. (Studies of the Permian Verbeekinoidean foraminifera-I). Mem. Fac. Sci.
Kyushu Uniu., Ser. D, Ceol., vol. 20, no. 1,
p. 17-58, pis. 1-9.
- - (1975): Evolution of Lepidolina multiseptata (Permain foraminifer) in East Asia.
Ditto, vol. 23, no. 2, p. 117-164, pis. 2226.
Ross, C. A. (1976): Development of fusulinid(Foraminiferida) faunal realms. Jour. Paleont., vol. 41, no. 6, p. 1341-1354.
Sada, K. and Yokoyama, T. (1966): Upper
Permian fusulinids from the Taishaku
Limestone in West Japan. Trans. Proc. Palaeonto Soc. Japan, N.S., no. 63, p. 303-315,
pis. 33-34.
Selwyn, A. R. C. (1872): Journal and report ofpreliminary explorations in British Columbia. Ceol. Suru. Con., Pept. Prog., 187172, p. 16-72.
Skinner, J. W. and Wilde, G. L. (1966): Permian
fusulinids from Pacific Northwest andAlaska. Uniu. Kansas Paleont. Contr., Paper4, p. 1-64, pis. 1-49.
Thompson, M. L. and Wheeler, H. E. (1942):
Permian fusulinids from British Columbia,
Washington and Oregon. Jour. Paleont.,
vol. 16, p. 700-711, pis. 105,-109.
- - , - - and Danner, W. R. (1950): Middle
and Upper Permian fusulinids of Washington and British Columbia. Contr. Cushman
Foundation Foram. Res., vol. 1, pts. 3-4,
p. 40-63, pis. 7-8.
Trettin, H. P. (1980): Permian rocks of theCache Creek Group in the Marble Range
Clinton Area, British Columbia. Canada
Ceol. Suru. Paper, 79-17, p. 1-17.
Yamagiwa, N. and Saka, Y. (1972): On the
Lepidolina zone discovered from the Shima
Peninsula, Southwest Japan. Trans. Proc.
Palaeont. Soc. Japan, N.S., no. 85, p. 260274, pIs. 31-32.
Zang, L. (1982): Fusulinids of eastern QinghaiXizang plateau. In Nanjing Institute ofGeology and palaeontology (eds.), Stratigraphy and Palaeontology in W. Sichuan
and E. Xizang. Acad. Sinica, Sichuan
People's Pub., p. 119-244, pIs. 1-36 (inChinese with English abstract).

Hiroya GOTO, Kunitaka MAR UOKA and Ken-Ichi ISHII

434

i::." 7 piC Lepidolina columbiana (~/t.- L, ~~*JJw.!:I:~): 7' ~ T ~ 'Y -/9,fiJ!1l) Yabeina mtl.J~¥Ilfi~n-C~'Qo <';1" 1. l:iicJ-lillJijtll)<"-7'
It.- 4'1'.::.*:.;c7'Y/~- " ' Y r !J~-!JI.J'~ Yabeina ~c~nt-:WjW.!1;{I::{:i~M!tH...
7' ~ T.{ 'Y -/ """":I P :.;:I P:'; i::."7j1!!lJijtl.J'~~HE.

~ni~¥Il~~nk~ll)cM~-cm~MIl)~.~AIl).N~fi~ko ~Il)M~. ~*~~.I.J'~i1GlI\t~nt-: Yabeina ~11 Lepidolina Hii~<6IJ, 1.J'-?i'ij-f1!I::::.~-tQc*IJil'JiLt-:o ~ll)fIi~ Lepidolina columbiana (DAWSON)
l.. -C ~G1liX Lt-:o
1~~!tf:5!r. • JLii7d~;f. • 1j;JI:{~-c

Explanation of Plate 86
Lepidolina columbiana (Dawson)
All figures x10
Figs. 1, 2, 6, 7. Specimens of inflated fusiform.
1. KU 3001, 2. KU 3002 from Marble Canyon.
6. KU 3006,7. KU 3007 from Upper Hat Creek.
Figs. 3, 8. Specimens of fusiform.
3. KU 3003 from Marble Canyon.
8. KU 3008 from Upper Hat Creek.
Figs. 4, 5, 9, 10. Specimens of elongate fusiform.
4. KU 3004, 5. KU 3005 from Marble Canyon.
9. KU 3009, 10. KU 3010 from Upper Hat Creek.

GOTO et al: L epido lina columbiana lrom B.C. Canada

Plate 866

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 435-446, pis. 87-89, September 30,1986

818. ON A NEW TEMPSKYA STEM FROM JAP AN*
HAR UFUMI NISHIDAThe International Budo University,
841 Monomizuka, Shinkan, Katsuura, Chiba 299-52

Abstract. Tempskya iwatensis sp. nov. was described from the Taneichi district, Iwate Prefecture, Japan. The age of the specimen was dated as Santonianbased on marine mollusca. Stratigraphically, this is the youngest Tempskya species
known with certainty. The past distribution of Tempskya is plotted on paleogeographic map in relation to the Cretaceous paleoclimate.

North America, chiefly in the Rocky Mountains. One species from Atlantic Coastal Plain(T. whitei Berry) has also been excluded byearlier workers for anatomical comparison because this species is also based on poorly preserved material. Endo (1926) reported a Tempskya stem from the Cretaceous of Kii (Wakayama Pref.). His specimen was based on a root
mass with no embedded rhizomes. Ogura (1927)
reidentified that specimen as part of a fern stemwith Cyatheaceous affinity, highly possibly
Cyathocaulis Ogura.
About 12 years ago Biichi Fukkiri, an amateurrock collector, found a petrified trunk at theseashore of Oh-hama, Yagi, Taneichi Town,
Sannohe-Gun, Iwate Prefecture (locality, 40°21'12"N, 141°45'30"E; Text-fig. 1). The areaaround the fossil locality, north of Kuji City iscalled here the Taneichi district. Two thirdsof the trunk was embedded in sandstone which
is widely distributed along the coast. Fukkiri
used the specimen to decorate a corner of hisgarden. Ryozo Sugiyama of the Ichinohe SeniorHigh School (Taneichi Senior High School atthat time) noticed the peculiar shape of thespecimen and advised Fukkiri to donate it to
the Iwate Prefectural Museum. The specimen
is now deposited in that museum. I was asked
to identify the specimen by the curator of theIntroduction

Tempskya is a genus of leptosporangiate
ferns flourished during the Cretaceous andprobably becoming extinct by the end of theperiod. It had a tree habit sometimes attaining a height of up to six meters. Some species
are thought to have had a prostrate or vinelike habit with dorsiventral stems. The stem of
Tempskya is generally described as a "false
stem" consisting mainly of a mass of adventitious roots in which are embedded slenderrhizomes and petioles of the plant.
A comprehensive reference to the history
and taxonomy of Tempskya was prepared by
Ash and Read (1976). According to them ten
species of Temp$kya have been described up to
1976. This excluded most of the European
species described prior to T. rossica Kidston et
Gwynne-Vaughan (1911), which were too
poorly preserved to allow anatomical comparison with later described tempskyas. Nine species,
other than T. rossica, have been described in*Received July 10, 1985; Revised manuscript
received July 15, 1986; Read June 24, 1984
at Akita University; Contributions from theLaboratory of Phylogenetic Botany, Facultyof Science, Chiba University, No. 106.
435

Harufumi NISHIDA

436

Text-fig. 1.

Map showing fossil locality.

museum. Through anatomical studies the specimen has been identified as a new species ofTempskya and this paper deals with its description and taxonomical treatment.
Geological setting and the age of the fossil:In the Taneichi district the Taneichi Formation oflate Cretaceous period is exposed along the
Pacific coast between Kuji City and Taneichi
Town. The Taneichi Formation was formerly
thought to be Miocene. It consists mainly ofshallow-marine sediments and can be subdivided
into three members.
The lowest member unconformably overlies
the granite basement. It consists of basal conglomerate and overlying sandstone in whichcoal seams and tuff beds are intercalated. The
middle member, which is a major part of theTaneichi Formation, consists of greenish graysandstone. Oyster beds partly intercalate in
the lower part of this member. From this member Terui et al. (1975) reported the occurrenceof a Cretaceous marine fauna including Ino-

ceramus (Sphenoceramus) naumanii Yokoyama
and Polyptycoceras cf. subundulatum (Yokoyama) that clearly indicate Santonian age (Matsumoto, 1977). This major sandstone member is
exposed along the coast at Yagi where Fukkiri
found the stem embedded in the sandstone of
the intertidal zone. It is highly probable thatthe age of the stem is Santonian. The upper
member consists of alternating layers of coarsegrained sandstone and conglomerate.
In the Kuji district, about 20 km south of
the Taneichi district, the Upper Cretaceousshallow marine to terrestrial sediments called
the Kuji Group are exposed. The Kuji Groupis divided into three formations, the Tamagawa,the Kunitan and the Sawayama in ascendingorder. Three members of the Taneichi Formation
can be stratigraphically correlated with threeformations of the Kuji Group, respectively(Matsumoto et al., 1982).
Tanai (1979) described a fossil flora fromthe Tamagawa and Sawayama Foramationsof the Kuji Group. The former is dated as Late
Coniacian and the latter as Early Campanianbased on both marine faunas and comparisonsof floral composition with that of other localities, especially that of North Saghalien. The
middle member of the Taneichi Formation is,therefore, correlated with the Kunitan Formation, which is Santonian in age.
Off the Pacific coast of the Taneichi district
the Paleogene sediments of the Noda Formation
are exposed. Petrified woods are also known tooccur from the Noda Formation, although they
have not yet been studied taxonomically. It
seems unlikely that the Tempskya specimen
could have been derived from the Paleogene
sediments.
Ikeya (1981) examined ESR (Electron Spin
Resonance) of the petrified woods that had
been collected as rolled stones at the coast of
Taneichi district. He reported that other biological materials of the same age show characteristic ESR patterns. Ikeya compared the
ESR of the rolled stones of Taneichi district
with that of in situ petrified woods collectedfrom both the Taneichi and the Noda Forma-818. New Tempskya from Japan

tions. The result indicated that the ESR of therolled stones is very similar to that of the Cretaceous in situ material. To ascertain the age of
the Tempskya specimen the examination of its
ESR pattern will provide useful evidence.
Material and method
The specimen before cutting was a petrified
columnar trunk, yellowish white on the surface,about 50 cm long, with diameters ranging from
7 to 11.5 cm. One end is almost smooth but isslightly knobby. The trunk weakly tapers to
one end. It was cut into five pieces as is shownin Text-fig. 2.
Peels prepared by etching the material aboutfour minutes in 20 to 25 percent hydrofluolic
acid solution were made into slides for study.
Systematic description
Order Filicales
Family Tempskyaceae Read and Brown, 1937
Genus Tempskya Corda, 1845
Tempskya iwatensis H. Nishida, sp. nov.

PIs. 87-89; Text-fig. 2.
Description :-Other than some longitudinalgrooves the specimen has a rather smooth external surface. It constitutes a so-called false
stem that is characteristic of Tempskya, consisting of numerous adventitious roots and a small
number of solenostelic rhizomes and petioles.
In the false stem the rhizomes give rise to petioles in all directions. Therefore the false stem
does not exhibit dorsiventrality (Text-fig. 2).
In section 1 (Text-fig. 2) of the false stem norhizome was present. Rhizomes appear andincrease in number toward the other end ofsection 1 indicating that it is nearer the baseof the false stem. Throughout the length of
the false stem the exact number of rhizomes
could not be determined because most of therhizomes had been decayed before fossilization.
The rhizomes and petioles are poorly preserved because of decay and some disruption

437

by roots. The anatomical description which
will be given below is derived from observations of these incompletely preserved components. Description of certain diagnostic featuresthat are thought to be taxonomically important,for example, the relative length of internodes
and amount of xylem parenchyma, are in conformity with the characteristics used in theartificial key to the species prepared by Ash
and Read (1976).
The rhizomes are very slender, 2.7 - 3.6 mm
in diameter, with long to medium internodes,
with slight overlapping leaf bases (usually twoor three). Leaf traces depart from only one
side of a rhizome alternately in two rows (PI.
87, Fig. 1). The epidermis is not clearly defined.
No dermal hairs nor their remnants are present.
A hypodermal layer that is comparable to theouter cortex as described by Ash and Read
(1976) was not clearly defined. The only evidence of a hypodermal layer is one or two external layers of the outer cortex that consistof less sclerotized cells (PI. 87, Fig. 3). The
cortex consists of two layers. The outer cortex,
that is comparable to the middle cortex cells
in other species, is sclerotic, about six or sevencells wide, with cells that are simple-pitted andcontain reddish brown contents. The innercortex is 15 - 17 cells wide, and composed ofthick-walled parenchyma cells that become
smaller near periphery of the layer.
The stele is solenostelic with a well-developedpith (PI. 87, Fig. 4). The endodermis is a single
layer of tangentially flattened cells. The pericycleis two to three cells thick. Internal to the pericycle is a layer of very small cells. This layer is
not always detected but when present probably
represents protophloem. Metaphloem is developed internally to the protophloem and is alayer three to five cells thick. The xylem plateof the concentric bundle is slightly thicker on
the ventral side and the tracheids of which it
is composed are scalariform. The position of the
proto xylem is not clear because of nearly equal
size of tracheids. However, smaller elements
tend to be located at the outer edge of thexylem cylinder. Xylem parenchyma is relatively

Harufumi N ISHID A

438

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818. New Tempskya from Japanabundant (PI. 88, Fig. 1). Parenchyma cells areoften alined in groups of five to more thaneight cells. The cells of endodermis, pericycleand xylem parenchyma contain reddish brown
contents similar to those occur in outer cortex.
The periphery of pith, which is about threecells thick, consists of thick-walled parenchyma
cells (PI. 87, Fig. 4). Toward the center of thepith cells become smaller and thick·walled. The
preservation of the pith is quite poor and thepith area is often invaded by adventitious roots.
However, at places the pith consists of sclerotic
cells as in other species of Tempskya (PI. 87,
Fig. 2).
Roots depart endogenously from the solenostele of the rhizome. The size, shape, stateof development and state of preservation ofroots vary considerably (PI. 89, Fig. 2). They
are equal to the roots of other species of Tempskya in general structure. Root hairs are present(PI. 89, Fig. 4). The cortex consists of outerand inner layers. The outer cortex is parenchymatous, with cells that are more thick-wallednear the periphery. The inner cortex is generally
sclerenchymatous, although the degree of sclerotization varies. In some roots the inner cortex
is completely thick-walled parenchymatous, orin rare cases only a layer adjacent to the endodermis is parenchymtous.
In mature roots the xylem is diarch, whereas
in juvenile roots such as shown in PI. 89, Fig. 3,
the central xylem mass is too small to be clearly
defined. In such young roots the inner cortex
is not typically sclerotized, although the degreeof sclerotization is not always correlated withthe age of roots.
Petioles are very slender, 1.2 x 1.6 mm indiameter, and do not persist in the tissue of thefalse stem after departure from the rhizome.
However, the petiole is better preserved than
the rhizome and persists for a greater distancein the false stem than the parent rhizome.
~439

The epidermis is decayed and thus lost. The
outer cortex is sclerenchymatous and consistsof cells similar in structure and contents to thecells of the corresponding layer in the rhizome.
It is 15 - 17 cells thick at petiole base. The
inner cortex consists of thick-walled parenchyma
cells. The stele is C-shaped and consists of anamphiphloic concentric bundle with a continuo
ous C·shaped xylem plate. Both adaxial margins
of xylem plate curve inward. The abaxial arcof the xylem strand is rather flat. Near the
point of insertion of the petiole its xylem strand
is thicker and lacks apparent proto xylem groups(PI. 88, Fig. 3). Distally in the petiole the xylem
strand diminishes in thickness until it becomes
one or two cells thick except for its adaxial
margins where xylem is three or more cells
thick. As the xylem strand thins three protoxylem groups become apparent (PI. 88, Fig. 2).
At places endodermis, pericycle, protophloem
and metaphloem are preserved (PI. 89, Fig. 1).
The center of pith consists only of thick-walledparenchyma cells (PI. 89, Fig. 1).
Diagnosis of the species:-False stem radiallysymmetrical, 7 to 12 cm in diameter at base.
Rhizome slender, 2.7 to 3.6 mm in diameter.
Internodes long to medium usually two or threeoverlapping petiole bases in one section. Cortex
two-layered; outer layer sclerotic, inner layer
thick-walled parenchymatous. Much xylem parenchyma in xylem strand. Proto xylem indefinite in xylem of rhizome. Pith of the petiole
consists of thick-walled parenchyma.
Locality:-Oh-hama, Yagi, Taneichi Town,
Sannohe-Gun, Iwate Prefecture; 40021'12''N,
141 °45'30"E.
Horizon and age:-Taneichi Formation; Upper
Cretaceous (Santonian).
Specimen:-Holotype (IPMM 31003) is deposited in the Iwate Prefectural Museum. Bothends of the holotype specimen and their micropreparations are deposited in the Laboratory of

Text-fig. 2. External features of specimen IPMM 31003 (E) and serial cross sections of upper part of false trunk (A-D) showing its radially symmetrical constitution. A
to D correspond to the surfaces indicated as a to c. Arrowhead indicates the level where
the section 1 mentioned in the text was made.

Harufumi NISHIDA

440

Phylogenetic Botany, Faculty of Science, Chiba
University.
Affinity and discussions

Affinity:-Nine species of Tempskya deserve
anatomical comparisons with the specimen represented here (Table 1). According to the separation of species by Ash and Read (1976) the
specimen belongs to the intragenetic group thathas radially symmetrical false stems. Among
species with radially symmetrical false stem T.
rossica is compared favorably with the Iwate
specimen in the histology of the cortex, amountof xylem parenchyma and the length of internodes, which are characters regarded by earlier
workers to be taxonomically valuable. The newform is distinguished from T. rossica by thesmaller size of the rhizome and the absence ofa central depression in the abaxial arc of a leaf
trace. Although T. zelleri also has a histology
similar to the Iwate specimen, it is highly characteristic in having angular stelar strands thatnever occur in the species described here, as wellas other early described species.
Apart from the radially symmetrical composition of the false stem the specimen anatomically resembles T. knowltoni and T. minor,
both belonging to the dorsiventral group. Smaller
size of rhizome, simple disposition of corticaltissues and long to medium internodes are thecommon features. The specimen is distinguishedfrom T. knowltoni in having many free petioles
in the false stem and in having a larger quantityof parenchyma in xylem strand. T. minor has
many free petioles in the false stem but lacks
xylem parenchyma. The petiolar pith consistingof thick-walled parenchyma is characteristicof the specimen and is a feature not found inother species. Although the Iwate specimen ispartially characterized by poorly delineatedcharacters such as the variable amount of xylem
parenchyma, the combination of features described here lead me to designate the specimen
as a new species.
Size of the plant:-Absence of well-preserved
rhizomes and petioles in whole length of thefalse stem indicates that the specimen represents
a basal part of the false stem. Based on careful
examination of numerous false stems, Andrews
and Kern (1947) estimated that T. wesselii
attained a height of 3.6 m if the basal diameteris 25 cm. The thickest false stem of T. wesselii
attains a diameter of 40 cm or more. With thisdiameter as a basis for calculation the plant
would have been nearly 5.8 m high. If it is postulated that the height of a plant is in proportion to the basal diameter of its stem, the heightof T. iwatensis with a false stem 10 - 12 cm indiameter at base, is estimated as 1.45 - 1.74 m.
Andrews and Kern also mentioned that thelower one sixth of the false stem consists of aroot mass alone. The lower 25 to 30 cm of the
Iwate specimen appears to consist of a root
mass without rhizomes.
Age of Tempskya:-Most Tempskya species
known to date are derived from the Lower
Cretaceous strata. Ash and Read (1976) revised
the age of Tempskya-bearing beds in North
America including the Patapusco Formation
bearing T. whitei. Most beds are now believed
to be Late Albian in age (Table 1). Some species
extend from Late Albian into Early Cenomanian.
According to Read and Brown (1937), European
tempskyas occur mainly in Wealden (Berriasian
to Barremian), but in part in Aptian (Lower
Greensand). Perucer beds in Czechoslovakia,
which yield T. varians Velenovsky, are considered to be Cenomanian in age (Read andBrown, 1936). Tempskya rossica from the Karaganda River Basin is the only specimen that isderived from unconfirmed Upper Cretaceous
strata. With the possible exception of T. rossica,
T. iwatensis is presentely the only Tempskya
known to occur in the Upper Cretaceous.
Ferns associated with Tempskya:-Fern leaves
are lacking in the Tempskya beds of the Taneichi
Formation. However, from the Kuji Group richfern flora has been recorded by Tanai (1979).
Although the Kunitan Formation, which is correlated with the Tempskya-bearing bed of theTaneichi Formation does not contain plantfossils, both the Tamagawa and Sawayama
Formations are rich in ferns. The Gyliakian

Table 1.

Anatomical comparisons of Tempskya iwatensis, sp. nov., with nine species of anatomically preserved Tempskya.

False stem*

Size ofrhizome(mm)

Internodelength**Outereorlex***

InnercortexSize offalse slem(em in diam.)

Xylem
paren·
ehyma

Senonian?

R

6.0-7.0

0

se.

pa.

5.5-9.5

2****

T. Imowltolli Seward

L. AlbianE. CenomanianD

2.5-3.5

0

se.

pa.

1.5-6.52

T. grallciis Read et Brown

L. AlbianR

4.6-6.0-se.

pa. + irr. se.

10.4-]2.2

3

T. millor Read et Brown

L. AptianAlbianD

2.0-3.5

0

se.

pa.

9.0-14.4

1

90

T. wesselii Arnold

L. AlbianR

4.0-5.0

+

se.

pa. +
doub. se.

7.0-40.0
or more3

E;

T. wyomillgellsis Arnold

L. AlbianR

6.0-8.0

0

se.

pa. +
doub. se.

?

3~

T. superba Arnold

L. Albian?

R

10.0-15.0-

sc.

pa. + irr. se.

more than
12.0

3SpeciesT. rossiea

Age

Kidsloll et Gwynne·Vaughan

T. reesiciei Ash et Read

L. AlbianR

3.0-5.00pa.

pa. island
in se.

10.0-30.0

00
.....

~
~

i'"

~

c

~

2

T. zelleri Ash el Read

L. AlbianR

3.0-8.0

+

se.

pa.

17.0

2

T. iwaiellsis, sp. nov.

SantonianR

2.7-3.6

0

se.

pa.

7.0-12.0

3

~

'g
;:s

*R: radially symmetrical, D: dorsiventral.
**+: very long, 0: long to medium, -: very short.
***In some species this layer is originally described as middle cortex.
****3: much, 2: little, 1: absent.
>I>->I>.....

442

Harufumi NISHIDA

flora of North Saghalien of the same age as T.
iwatensis contains 21 species of ferns, of whicheight are common to the Tamagawa and Sawayama floras. Because the affinities of Tempskya
are still uncertain it is impossible to relating
Tempskya stems to the isolated remains offern leaves.
In looking for possible affinities of Tempskya
to fern families several earlier workers haveemphasized the occurrence of Anemia fremonti
Knowlton foliages (Schizaeaceae) in a horizon
the same as or close to that bearing Tempskya(Seward, 1924; Read and Brown, 1937). The
Sawayama flora contains A. elongata (Newb.)
Knowlton and the Gyliakian flora contains A.
cf. elongata (Kryshtofovich and Bykovskaya,1960).
Many anatomically preserved fragments offerns embedded in Late Cretaceous calcareous
nodules have been described from Hokkaido(Stopes and Fujii, 1910; Ogura, 1927, 1930;
Hashimoto, 1971; Nishida and Nishida, 1979;
Nishida, 1981a, b). Most of these belong to
Cyatheaceae. The age of the strata bearing
such nodules extends from Cenomanian toCampanian, and is partly comparable with theage of the Taneichi Formation. Only one sporebearing fragment, Schizaeopteris mesozoica
Stopes et Fujii, however, suggests the presenceof Schizaeaceae. There is no record of Tempskya
from either Hokkaido or Saghalien. To avoid
further speculation I emphasize here the needfor much wider and detailed investigations onthese Cretaceous ferns in an effort to clarify
the taxonomic position of Tempskya.
Other plants associated with Tempskya:Seward (1924) reported the occurrence of anaraucarian root associated with T. knowltoni.
When Arnold (1945) described T. wyomingensis
he recorded Cyacadeoidea sp. from the same
locality. According to Andrews and Kern (1947)
Cupressinoxylon sp., Cycadeoidea sp. and dicotyledonous woods occur with T. wesselii in
Idaho.
From the Taneichi Formation four speciesof gymnospermous woods have been recognizedby M. Nishida (pers. com.). Although the detailed taxonomic accounts will be publishedelsewhere, he tentatively recognizes Araucarioxylon kiiense Ogura, Cupressinoxylon vectense
Barber, Taxodioxylon albertense Shimakura anda new species of Piceoxylon. Cycadeoidea wasdescribed from the Late Cretaceous Upper Yezo
Group of Hokkaido (Kryshtofovich, 1920;
Endo, 1925). The floral components associatedwith Tempskya in Japan and those in North
America are very similar and indicate similar
paleoecological conditions at these localities.
A possible habitat for Tempskya, suggested byAndrews and Kern (1947) is a tropical montane
cloud-forest, where most species of the tree
ferns occur today.
Distribution of Tempskya and its relation topaleoclimate:-Parrish et al. (1982) predict rain-

Explanation of Plate 87
Fig. 1. C.s. of dorsal half of a half-decayed rhizome, showing leaf traces and free petioles (arrowheads) departing in two rows. X 15.6. Bar 1 mm.
Fig. 2. C.s. of a best preserved rhizome. Sclerotic pith is partly preserved (arrowhead). Outercortex is decayed but inner cortex consisting of thick-walled parenchyma is well preserved.
X 29.0. Bar 0.5 mm.
Fig. 3. C.s. of a rhizome showing construction of tissues. Outer cortex is preserved only at its
periphery. Cells of outermost layer of outer cortex are larger and thinner-walled than inner
cells, indicating the presence of a hypodermal layer. ic; inner cortex, oc; outer cortex, vb;
vascular bundle, p; pith. X 30.7. Bar 0.5 mm.
.
Fig. 4. Vascular tissue in fig. 3 enlarged. Protoxylem indefinite. Note abundant xylem parenchyma. Phloem is badly preserved. Periphery of pith consists of thick-walled parenchyma.
e; endodermis, pe; pericycIe, ph; phloem, xy; xylem. X 61.4. Bar 0.1 mm.

NISHIDA : New Tempskya from Japan

Plate 87

818. New Tempskya from Japan
fall patterns, based on atmospheric circulationsof the past, to explain the distribution of coals
and evaporites in the Mesozoic and Cenozoic.
Paleobotanical data are useful in testing such ahypothesis. Text-fig. 3 shows a distribution of
Tempskya plotted on a paleogeographic map.
The base map (Owen, 1983) shows the distribution of t he continents in the Turonian stage .
Contours of rainfall patterns (relative values
only) are based on Cenomanian data by Parrish
et al. (1982) who also present Maestrichtian data.
Although the age of Tempskya extends from
Berriasian to San to nian, I preferred here to use
Cenomanian data as being represen tative of thepaleoclimate of Tempskya age. Because judging
from the paleoclimate maps by Parrish et al.
(1982), the paleogeography and paleoclimate ofTempskya-bearing localities had been subject tominor change through the Cretaceous period .
The worldwide occurrence of Tempskya is

o· -

443

reviewed by Read and Brown (1937). North
American localities are mapped by Ash andRead (1976). In North America, Temps llya
occurs in the Rocky Mountain Geosyncline ,Mexican Geosyncline and Atlantic CoastaloPlain, extending from 3I N to 4SoN and 103° W
to 118°W. In Europe Tempsllya-bearing beds0
are located from 50 N to 53° N and 2° W to
65° E. The locality of T. iwatensis is approxi omately 40 N, and 141° E.
The Rocky Mountains and Mexican Geosynclines, which are major sources of No rth American Tempsllya, are developed in the easterninterior of the Rocky Mountains that provided
a rain shadow against the westerlies. AnotherNorth American locality faces the Atlantic coast.
In Europe, England and Ireland localities andFrench to Bohemian localities are all on islands.
The Karaganda River Basin near t he Ural Mountains is situated at the ma rgin of the Asian conti-.,. -Text-fig. 3. Distribution of Tempshya on pal eogeographic map showing rainfall
patterns in Cenomanian. Paleogeography an d rainfall patterns are redrawn after Parrish
et al. (1982) on base map by Owen (1983). Numbers show relative values of rainfall. 200;
high, 200-100; moderately high, 100-50; modera tely low, 50; low, shaded; approximate
contour of land masses.

-1

444

Harufumi NISHIDA

nental lowland on the north coast of Tethys.
The Taneichi district is located at the east
Pacific margin of Asia continent.
Tree ferns need high humidity for theircontinuous growth and thus the postulated
habitat for Tempskya is a tropical or subtropicalhumid forest developed at high elevations (1,200- 2,100 m) on undulating hills or mountains(Andrews and Kern, 1947). In tropical to subtropical South America, cloud-forests rich inferns are well developed on the Andean slopes
at higher elevations (1,800 - 2,600 m).
Species of Tempskya are distributed in lowlands at rather high latitudes except in North
America where several localities occur at nearly
31°N. In these localities rainfalls are predicted
to be moderately high as well as moderately
low without affecting the distribution of Tempskya. These paleogeographical and paleoclimatological conditions differ somewhat from thepostulated habitat of Tempskya.
From my observations of living species of
Cyatheaceae in South America, Australia andsouth Japan, continuous atmospheric moistureseems to be a most important factor limiting
their growth rather than total amount of rainfall, if the temperature is moderate. Under
moderate temperatures coastal cliffs or mountains of no more than 400 - 500 m seems to behigh enough to produce sufficient humidity fortree-fern growth. The Juan Fernandez Islands
and Pacific coast of South Chile are examples
of such localities. According to Parrish et al.
(1982) the climate in a given area is regarded
to be wet when precipitation exceeds evaporation, even though the total amount of precipitation is low. All localities are placed either
near the coast or on islands that are influenced

by wet oceanic winds. The possible habitat ofTempskya could be a humid low to high montane forest in subtropical or even a warm temperate climate. The fact that evaporites arenever associated with these localities supports
the idea of a lower evaporation rate resulting
in high humidity (Parrish et al., 1982).
Tempskya has not yet been described from
Asian inlands at latitudes comparable to thoseof other localities. This may be the product of
a dry climate. Much more information must
be obtained, however. Wide latitudinal distribution of the American Tempskya is in marked
contrast to the restricted distributional patternof the European members. In North America
a moist habitat occurs along the northward
extension of the north Atlantic, whereas inEurope land masses are confined to latitudes
around 50o N. This may account for restricteddistribution of Tempskya in Europe.
On the naturalness of intragenetic groups:Whether the false stem of Tempskya is radiallysymmetrical or dorsiventral has been used as
one of important criteria in classifying species
in~o two subgeneric groups (Read and Brown,
1937; Andrews and Kern, 1947; Arnold, 1958;
Ash and Read, 1976). However, Andrews andKern (1947) questioned this grouping andsuggested that dorsiventrality reflects either ajuvenile stage or only a part of an entire false
stem of a plant. No fossil evidence supportingthese suggestions has been discovered as yet.
The dorsiventral group (T. knowltoni and T.
minor) has slender rhizomes less than 3.5 mm indiameter. The radially symmetrical group has
thicker rhizomes usually more than 4. 0 mm indiameter, except for T. reesidei and T. zelleri
where diameters decrease to 3.0 mm in the lower

Explanation of Plate 88
Fig. 1. C.s. of rhizome. Part of a xylem strand. Abundance of xylem parenchyma is better displayed. X 170. Bar 0.1 mm.
Fig. 2. Enlargement of a free petiole shown in Plate 87, Fig. 1. Thin xylem strand with three
protoxylem groups (arrowheads) is shown. Outer cortex is lost. X 140. Bar 0.1 mm.
Fig. 3. C.s. of a petiole base showing a vascular strand without central depression on its abaxialarc. Protoxylem is not clear in xylem plate. Upper side is abaxial. X 43. Bar 0.5 mm.

N ISHIDA: New Tempskya from Japan

Plate 88

818. New Tempskya from Japan
extremities of the rhizome. The simple structureof cortical tissues is another feature common
to the dorsiventral group. The consistent rela·tionship between internal structure and thedorsiventrality appears to support the natural·
ness of the grouping.
T. iwatensis, however, displays characteristics
that are intermediate between the two groups.
This species belongs to the radially symmetricalgroup, but at the same time displays a slenderrhizomes and a simple disposition of corticaltissues as in the dorsiventral group. This combi·
nation of characteristics in T. iwatensis suggeststhat there is no strict boundary between thetwo groups. The construction of the false stem
is still, however, a good feature in identifying
Tempskya species.
The rhizomes of Tempskya are dorsiventral.
Therefore, the fern that is hypothetically ancestral to Tempskya could have had prostrate,
dorsiventral rhizomes like those of the living
schizaeaceous ferns. The false stems first formedby such an ancestor probably was a dorsiventral
type, so that the primitive form of Tempskya
would have structure similar to T. minor withits dorsiventral stem and that exhibits the simplest structure of all. The slightly younger age ofT. minor supports this idea. However, it remains
uncertain whether or not T. minor and anotherdorsiventral species, T. knowltoni, are direct
descendants of the ancestral form. Tempskya
is a highly variable group that diversified froma simple ancestral form in the Cretaceous. T.
iwatensis is a species, that in the course of its
evolution, has retained the simple internal
structures of some other tempskyas.
Acknowledgement
I am grateful to Prof. Makoto Nishida of
Chiba University for his continuous encouragement and suggestions. Dr. Wilson N. Stewart,
Prof. Emeritus of the University of Alberta,
Canada, kindly reviewed the manuscript and gave
me valuable suggestions. Thanks are due to Mr.
Ryozo Sugiyama of Ichinohe High School forgiving me an opportunity to examine the speci-

445

men that is now deposited in the Iwate Prefectural Museum to which I also acknowledge forthe loan of the specimen. This study was partly
supported by a Grant-in-Aid for Encouragementof Young Scientists from the Ministry of Education, Science and Culture, No. 60740385.
References
Andrews, H. N. and Kern, E. M. (1947): The
Idaho tempskyas and associated fossil
plants. Ann. Missouri Bot. Gard., vol. 34,
no. 2, p. 119-183, app. p. 185-186, pis.
15-27.
Arnold, C. A. (1945): Silicified plant remains
from the Mesozoic and Tertiary of western
North America. 1. Ferns. Papers Michigan
Acad. Sci. Arts and Letters, vol. 30, p. 334, pis. 1-11.
--(1958): A new Tempskya. Contr. Michigan
Univ. Mus. Paleontology, vol. 14, no. 8, p.
133-142, pis. 1-3.
Ash, S. R. and Read, C. B. (1976): North American species of Tempskya and their stratigraphic significance. U.S.G.S. Prof. Paper,
874, p. 1-42, pis. 1-13.
Endo, S. (1925): Nilssonia-bed of Hokkaido
and its flora. Sci. Rep. Tohoku Imp. Univ.,
2nd Ser., vol. 7, no. 3, p. 57-72, pis. 11-17.
--(1926): On the genus Tempskya from Kii.
Chikyu, vol. 6, p. 5-6 (in Japanese).
Hashimoto, W. (1971): On a new Cretaceous tree
fern from Nakagawa-machi, Teshio Province,
Hokkaido. Sci. Rep. Tokyo Kyoiku Daigaku,Sec. C, vol. 11, no. 103, p. 1-10, 3 pis.
Ikeya, M. and Miki, T. (1981): ESR (Electron
Spin Resonance) of fossil bones, woods
and natural minerals. Earth, vol. 3, no. 8,
p. 477-483 (in Japanese).
Kidston, R. and GWynne-Vaughan, D. T. (1911):
On a new species of Tempskya from Russia.
Verh. Russ. K. min. Gesell., vol. 48, p. 120, pis. 1-3.
Kryshtofovich, A. N. (1920): A cycadean trunk
from Hokkaido. Jour. Geol. Soc. Tokyo,
vol. 27, no. 325, p. 1-8, pI. 19.
--and Baykovskaya, T. N. (1960): Mesozoic
flora of Saghalien, 122 p., 21 pis., Academia
Nauka, Moscow (in Russian).
Matsumoto, T. (1977): Zonal correlation of theUpper Cretaceous of Japan. Pal. Soc. Japan,

Harufumi NISHIDA

446

Spec. Paps .• no. 21, p. 63-74.
- - , Obata, I., Tashiro, M., Ohta, Y., Tamura,M., Matsukawa, M. and Tanaka, H. (1982):
Correlation of marine and non-marine formations in the Cretaceous of Japan. Fossils,
no. 31, p. 1-26 (in Japanese):
Nishida, H. (1981a): Anatomical studies of a new
specimens of Yezopteris polycycloides
Ogura from the Upper Cretaceous of Hokkaido, Japan. Jour. Jap. Bot., vol. 56, p.
169-180, pI. 8.
- - (1981b): A revision of the genus Cyathorachis in Japan. Bot. Mag. Tokyo, vol.
94, p. 249-259.
--and Nishida, M. (1979): Thyrsopterorachis.
gen. nov., a tree fern stipe from the Upper
Cretaceous of Hokkaido, Japan. Ibid .• vol.
92, p. 187-195.
Ogura, Y. (1927): On the structure and affinities
of some fossil tree-ferns from Japan. Jour.
Fac. Sci. Imp. Uniu. Tokyo. Bot.. vol. 1, p.
351-380, pIs. 2-8.
-(1930): On the structure and affinities ofsome Cretaceous plants from Hokkaido.
Ibid .• vol. 2, p. 381-412, pI. 18-21.
Taneichi :fl1!m. Kuji 7-.~.
Kunitan 1'!!Ift. Noda !!fm.

Owen, H. G. (1983): Atlas of continental displacement, 200 million years to the Present.
159 p., Cambridge Univ. Press, London.
Parrish, J. T., Ziegler, A. M. and Scotese, C. R.
(1982): Rainfall patterns and the distribution of coals and evaporites in the Mesozoic and Cenozoic. Palaeogeogr.. Palaeoclimatol.• Palaeoecol .• vol. 40, p. 67-101.
Read, C. B. and Brown, R. W. (1937): American
Cretaceous ferns of the genus Tempskya.
U.S.G.S. Prof Paper. 186-F. p. 105-131,
pis. 16-17.
Stopes, M. C. and Fujii, K. (1910): Studies on
the structure and affinities of Cretaceous
plants. Phil. Trans. Roy. Soc. London, B.
vol. 201, p. 1-90, pIs. 1-9.
Tanai, T. (1979): Late Cretaceous floras fromthe Kuji district, northeastern Honshu,
Japan. Jour. Fac. Sci. Hokkaido Uniu .• Ser.
4. vol. 19, no. 1-2, p. 75-136, pIs. 1-14.
Terui, K., Terui, K., Yanagisawa, H. and Kobayashi, T. (1975): Cretaceous fossils from theTaneichi Formation in the north coast of
Rikuchu, Iwate Prefecture. Jour. Geol. Soc.
Japan, vol. 81, no. 12, p. 783-785.

Oh-harna *~. Vagi /\*.
Hokkaido ~tila~.

Tarnagawa :li)II.

Sawaya rna /Rlli.

B *£7;/ 7 7-=\'- 70) ~:fl1!t.:."?~'-C: !ft=F~~PlillllmlBJ /\**~O) ~JIr..'-KHNIO) fijl!ft.t~ • • ~ • • • ~U.Lk.~*~. *~*~.O)*~~~ 7;/77-=\,-7 (Thm~~a).

ccO)ifi.:il:tIl~ nn' -? t.:o t~*li:fl1!mJ\1<p$~n' I? 0)£1±l ~ I?h. II'.'H~lilfil~fJt£O)jfb~{t15C. 7-.~.~t.:.7};ffl-to7-.~Jf/tt'f.CO)~mft~I't-.J!t~n'l? J:$B!lli*-lj-;/".:=. 7;/ cHfjE~c.

c.

~. - It.-1!t.:. J: 0 mtilfiWfilJO);f6:!ill:. im:.:il:¥a.:. ~JUil1n;f!\Ii~';:*ml*mtiln;~~';:~mO)M:t.:.~~mtiln;9(jit-lt-r. n'n ~ t.:.H9i;O)I'J~,*m til n;;6 0 ;:ts:. en' I? ffr:&l.
Tempskya iwatensis L -C~G. Lt.:o Iiil.Q)(lP.:fl1!O)7};(fla-~6l). E1 !lli*cO)i~-A~~IJ:-eO)7}

ht.:o

c

c

Tempskya O)~1f:mma-Jt:iELt.:o ¥Bff:¥.£O)(g;llln'I?~~iWi!1l!t':'~E'ffL. tt*~;tI?ht.: J: ? tdMW~J: ~ litl' L7::>. 11jilJW~t.:.7};ffl Lt.::;: c n;? n'n;nhoo .1*J007}1tillFt.:. "?

;{fin'l?

~'-C\-dNFa-ffifUr:t.:ol§1Ri'int

Explanation of Plate 89
Fig. 1. Enlargement of a petiolar vascular bundle at its adaxial margin. Pith consists of thickwalled parenchyma (arrowhead). e; endoderm is. X 227. Bar 0.1 mm.
Fig. 2. Adventitious roots displaying various degrees of preservation and construction of tissues.
X 31. Bar 0.5 mm.
Fig. 3. Adventitious root with inner cortex composed of cells having less thickened walls. Xylem
mass is poorly developed. X 81. Bar 0.1 mm.
Fig. 4. Long intermingled root hairs. X 163. Bar 0.1 mm.

NISHIDA : New Tempskya from Japan

Plate 89

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 447-462, pIs. 90-92, September 30, 1986

819. THE SECOND ADDITION TO THE SILURIAN TRILOBITEFAUNA OF YOKOKURA-Y AMA, SHIKOKU, JAP AN*
TEIlCHI KOBAYASHIJapan Academy, Ueno Park, Taito-ku, Tokyo
and
TAKASHI HAMADA
Department of Earth Science and Astronomy, College of Arts and Sciences,
University of Tokyo, Komaba, Meguro-ku, Tokyo

Abstract.
Nine species of trilobites including six new species and two newsubspecies from Mt. Yokokura are described in this article. A new genus, Ichiyamella, is instituted on Ichiyamella subglobula, sp. nov. Besides this genus Japonoscutellum and Apolichas are indigenous to Japan or Eastern Asia. Some notes are
given on the Scutelluidae, Meroperixinae, Illaenidae, Goldillaeninae, Sphaerexochinae, Dindymeninae and the Lichidae in addition to Sphaerexochus, Dindymene
and Dicranopeltis and Dindymene (?) megacranidia.

In the monograph on Silurian Trilobitesof Japan etc. (1974), 26 species of trilobites
were described from Mt. Yokokura. Recently
11 species were added to them (1984, 1985).
A further addition is made here with 9 species
and subspecies as listed below. The faunal analysis is postponed to another occasion.
Palaeontological notes are given on the families
and subfamilies marked with plus in the list
below beside a few genera. Ichiyamella is a new
genus proposed on this occasion.

and Hamada.
Japonoscutellum japonicum puteatum, subsp.
Japonoscutellum japonicum laticephalum, subsp.
nov.
Subfamily Scutelluinae R. and E. Richter (?)
Scutelluid, gen. et sp. indet.
Subfamily.Meroperixinae Kobayashi and
Hamada+Opoa (?) trinodosa, sp. nov.
Family Illaenidae Hawle and Corda+
Subfamily Goldillaeninae Balashova+
Illaenoides (?) magnisulcatus, sp. nov.
Illaenoides (?) abnormis, sp. nov.
Family Proetidae Salter
Subfamily Cyphoproetinae Pillet
Cyphoproetus latiaxis, sp. nov.
Family Cheiruridae Salter
Subfamily Sphaerexochinae Opik+Sphaerexochus hiratai Kobayashi and Hamada
Family Encrinuridae Angelin
Subfamily Dindymeninae Kielan+

List of additional trilobites
Family Scutelluidae R. and E. Richter+
Subfamily Planiscutelluinae Ko bayashi*Received September 17, 1985; Studies on
Japanese Trilobites and Associated Fossils,
No. XXXIX.
447

448

Teiichi KOBA YASHI and Takashi HAMADA

Dindymene (?) sp. indet.
Ichiyamella subglobula, sp. nov.
Ichiyamella megacranidia (Kobayashi and Hamada)
Family Lichidae Hawle and Corda+
Subfamily Lichinae Hawle and Corda
Dicranopeltis tricornis, sp. nov.
Subfamily Homolichinae Phleger
Apolichas truncatus Kobayashi and Hamada

Ack:lOwledgments
Here the authors record their best thanks to
Messrs. S. Asami, I. Fujiyama, A. Kaneko, F.
Kojima, Y. Mizuno, S. Nakajima, K. Noda andT. Uchida for collecting the trilobites dealt withhere, and to the Japan Academy for the grant tothis study.
Description of trilobites
Family Scutelluidae Richter and Richter, 1956
Leioscutellum and Lamposcutellum were
described from China respectively by Wu (1977)
and Yin (1980). The latter is one of the oldest
scutelluids that bear some characteristics of theEobronteinae as well as the Planiscutelluinae.
The former may be the oldest genus of the
Paralejurinae.
Lamposcutellum Yin, 1980, whose type is
L. guizhouense Yin, 1980, from Middle Ordo·
vician in Guizhou, is allied to. Eobronteus,
Protobronteus and Planiscutellum in one or
another characteristic. Its glabella is constricted
at about one-third of the cephalon from theposterior margin, more expanded in anterior,reaching the frontal margin and provided withthree pairs of lateral furrows; occipital furrow
transversal; neck ring uniform in sagittal length;
axial furrow well pronounced in middle and posterior; small paraxial nodes occasionally present
at the constriction; eyes of medium size opposedat posterior furrows and lobes; eye-ridge extending from middle lateral furrow to eye; fixedcheek anterior to this ridge divided into two
subequal parts by another ridge; lateral marginalborders extending from two sides of glabella

and produced into genal spines. Pygidium isbell-shaped; axial lobe nearly as long as broad, alittle shorter than one-third of pygidium; medianrib bifurcate in posterior; 7 pairs of pleduralribs present on its two sides. Lamproscutellum
shaanxiense Zhou, 1982 and L. latilimbatum
Zhou, 1982, occur in Middle Ordovician, Shaan xi.
Leioscutellum Wu, 1977 founded on L.
tenuicaudatum Wu, 1977 from Lower Silurian
in Guizhou, resembles Paralejurus, but its glabella
is less expanded in anterior, occipital furrow describing two arcs, eye large and genal angle
rounded on cephalon and 8 pairs of pleural ribs
on two sides of the simple somewhat broad
median rib on the bell-shaped pygidium.
Australoscutellum Chatterton and Campbell,
1980 was instituted on Bronteus longispinifer
Mitchell, 1887 from the Ludlovian in New
South Wales, Australia. According to the authorsof the genus it is allied to Microscutellum inthe glabellar pattern, but it has only 5 or 6 pairsof pleural ribs, besides a median rib which seems
to be trisected. This genus indicates probably
an early isolate branch of the Thysanopeltinae.
Un icapeltis Maximova, 1977, whose typespecies is Unicapeltis unica Maximova, 1977
from Middle Devonian of Salair, is quite distinctive in its glabellar configuration, particularlyin the connection of the posterior pair of profound lateral furrows in describing a backwardly
convex arc in the axial part. A pair of shallow
paraxial furrows extend forward from bothsides of this arc delimiting middle lateral lobes.
Nevertheless, this genus resembles Thysanopeltella, Arctopeltis and Altaepeltis. It would
be another aberrant genus of the Thysanopeltinae.
Subfamily Planiscutelluinae Kobayashi
and Hamada, 1974
Genus Japonoscutellum Pftbyl and Vanek, 1971
Japonoscutellum japonicum (Kobayashi
and Hamada) puteatum, subsp. nov.
PI. 90, Figs. la-c.

819. Silurian Trilobites from Shikoku, 2nd Addition
This cephal on agrees with J. japonicum inmajor aspects, but it disagrees with the typicalform of the species in the middle and anterior
lateral furrows which are disconnected fromthe axial furrow and strongly pitted, somewhat
smaller eyes and the distinct eye-ridge and ala.
The cheek is narrow and depressed, but
provided with a narrow convex lateral border.
A narrow ridge runs along the lateral margin of
the eye. The anterior branch of the facial suture
extends from the eye and then diagonally crosses
the cheek to the marginal border, interrupting
the above ridge. Fine linear ornaments are seenon the glabella, particularly well in its anterior
part.
Occurrence:-Gomi quarry, Yokokura-yama;collected by Kojima.

Japonoscutellum japonicum (Kobayashi
and Hamada) laticephalum, subsp. nov.
PI. 90, Figs. 2a-b.

Description :-Cranidium broader than long,most expanded laterally in anterior, gently
inflated and becoming subvertical along frontal
margin with forward convexity; sagittal profile
nearly straight except for occipital furrow andanterior bent of frontal lobe. Glabella fungiform,
most expanded at frontal lobe, strongly contracted at posterior lateral furrow and as wide
as occipital ring at middle lateral furrow; posterior furrow expanded adaxially, separating
posterior and middle lateral lobes of subequal
length; middle furrow longer and anterior furrow
longest; both linear; anterior lateral lobe shorter
than middle or posterior lobe; these three pairsof lateral furrows all interrupted by axial zone
which is a little narrower than posterior furrow;un furrowed axial zone becoming a little wider
between anterior pair of furrows; occipital
furrow deeper than these lateral furrows; occipital ring somewhat shorter than posterior laterallobe on lateral side; frontal lobe arching downin anterior and becoming subvertical; axial
furrow deep. Fixed cheek narrow and depressed;
eyes of medium size opposed at posterior laterallobes; eye-ridge extending from middle lateral449

furrow to palpebral lobe; ala small, suboval,located on lateral side of posterior lateral lobe
of glabella. Facial suture diagonal posterior toeye; its anterior branch extending from eye tolateral end of frontal glabellar lobe. Fine striae
like terrace-lines on doublure seen most of dorsal
test which are subparallel anterior margin on
glabella, particularly dense and distinct onfrontal lobe, more or less longitudinal on neckring and subparalltl to axial furrow on fixedcheek.
Comparison :-This cranidium resembles thoseof Planiscu tellum and Kosouopeltis, but theessential distinction from them is the absenceof the preglabellar depression. Planiscutellum
has the large concave preglabellar depression
in front of the glabella. The depression is very
narrow in Kosouope/tis, but it separates theglabella from the subvertical doublure clearly.
This cranidium reveals the characteristics ofJaponoscutellum in the glabellar pattern, but it
disagrees with J. japonicum s. str. in the lateralexpansion of the cranidium in anterior, eyes
located more posteriorly and possession of
distinct eye-ridges and smaller alae. The linear
ornamentation is well developed in this subspecies.
Occurrence :-The Yokokura-yama limestone;
collected by Noda.
(?) Subfamily Scutelluinae R. andE. Richter, 1956
Scutelloid, gen. et sp. indet.
PI. 90, Fig. 3.
A small dorsal shield, about 14 mm long.
Thorax is composed of ten segments, most
expanded laterally at about posterior one-third,
axial lobe broader than pleural lobe in anterior,
but regularly narrows back as far as the axial
and pleural lobes become subequal in breadth.
Pleural furrows are absent.
Pygidium is suboval, wider than long andmoderately convex, convexity being emphasizedin one-third near anterior margin; axial lobe
sm~l subtriangular and well inflated; its medianpart smooth, parallel-sided and slightly elevated450

Teiichi KOBA YASHI and Takashi HAMADA

above lateral parts. Pleural ribs seven on eachside; median rib a little broader than others;
all of them flat·topped and separated from one
another by narrow furrows; doublure about
half as wide as pleural lobe.
The pygidium disagrees with other pygidia
from the Yokokura-yama limestone in theoutline and convexity of the pygidium and theflat ribs and narrow furrows. The median rib
may be simple. Because the cephalon is illpreserved, its generic reference cannot be made.
Occurrence:-Gomi quarry, Yokokura·yama;collected by Kojima.
Subfamily Meroperixinae Kobayashi
and Hamada, 1974
Ligiscus Lane and Owen, 1982, whose typespecies is L. arcanus Lane and Owen, 1982 fromthe Silurian near the Llandovery-Wenlock boundary, in western North Greenland, is an effaced
scutelluid probably referable to the Meroperixinae. In this genus the occipital ring is
ill-defined, but a median tubercle is present in
the palce.

Genus Opoa Lane, 1972
Opoa (?) trinodosa Kobayashi and Hamada,sp. nov.

PI. 90, Figs. 4a--c.
1974. Microscutellum primigenium Kobayashi
and Hamada (pars), p. 74, pI. 5, figs. 9a--c
only
Descrip tion :-Glabella three-fourths as long as
broad, semi elliptical transversally expanded inanterior one-third of sagittal length and conicaland tapering back as far as its half breadth in
the remainder, gently ascending to one-third ofsagittal profile from posterior margin where alarge median tubercle exists; lateral slope steep
in conical part, whereas it is gentle in expanded
anterior; frontal rim narrow with terrace lines
separated from glabella by narrow marginal
furrow; irregular linear ridges distributed inanterior and middle parts of glabella which aresubparallel to frontal margin, but they become

wavy backward; many tubercles spread in wholeglabella, mostly in interspace between ridges,
but ridges are absent in posterior part; mediantubercle at two-thirds from front very large; apair of tubercles on its postero-Iateral sides,
second largest; axial furrow deep; basal laterallobes subtriangular; eyes of medium size opposedin posterior.
ObselVation and comparison:-This species isrepresented by an imperfect cranidium whose
fixed cheeks and neck ring are largely gone,
but a semicircular scar of the right eye-bandsuggests that the fixed cheek is very narrow.
It is certain that this cranidium belongs to
the Scutelluidae, but it does not fit in any
generic diagnosis so far known. The nearest to
it is Opoa whose glabella is, however, simply
parallel-sided in posterior, but it is truncateconical and provided with a pair of the basal
lateral lobes in this species. The test is granulate
as in Opoa, but the interspace reveals a honeycomb structure in Opoa, while it is ridged in theanterior side and three large tubercles are present
in the posterior part of the glabella in thisspecies. Therefore, it is probable that this species
will be found to be a distinct genus in future.
Among the pygidia referred to Microscutellum primigenium there are two kinds. With thisfind it becomes probable that the strongly
granulate pygidium having geniculate pleuralarea like Opoa damesi Lane to which the authors
have compared belongs probably to this species.
Occurrence :-Yokokura-yama limestone, atGomi quarry, collected by Asami.
Family Illaenidae Hawle and Corda, 1874
Raymond's bipartition of the Illaenidae into
the Illaeninae and Bumastinae (1916) has been
widely accepted by Hupe (1955), Balashova(1975) and others. In Moore's Treatise (1959)
Jaanusson discriminated the Ectillaeninae in thefamily and provisionally added the Theamataspidae Hupe, 1953 to it as a subfamily. Later
Bruton (1970) erected the Panderininae for
Panderia. In additing the Goldillaeninae Balashova, the authors (1974) classified the family819. Silurian Trilobites from Shikoku, 2nd Addition
into the Illaeninae, Panderininae, Ectillaeninae,
Buamastinae, Goldillaeninae and (?) Theamaspidinae beside three genera whose subfamilyreference is uncertain.
Balashova (1960) instituted the Goldillaeninae
in the Scutelluidae to include Goldillaenus andGoldillaenoides the latter of which had theoccipital ring. While Jaanusson (1979) accepted
Goldlllaenus as a subgenus of Illaenoides Weller,
1908, Lane (1972) brought back the Goldillaeninae to the Scutelluidae.
Chatterton and Ludvigsen (1976) instituted
a new genus, Failleana with Failleana clava as
its type species and suggested that the Bumastinae be placed in synonymy with the Illaeninae
as they found the rostral flange in Bumastus(Bumastoides) aplatus. Lane and Thomas (1978)
on the other hand referred Bumastus to the
Scutelluidae and added three new Bumastuslike Silurian genera, i.e. Cybantyx, Litotix andRhax the last of which is based on Rhax pollincitrix Lane and Thomas from the LlandoveryWenlock of Queensland, Australia. Thus thetaxonomy of the family Illaenidae and its relation to the Scutelluidae is now a moot question.
The Typhlokorynetidae Shaw, 1966 and thePortaginidae Lesperance, 1968 are represented
respectively by Typhlokorynetes Shaw, 1966
and Portagius Lesperance, 1968. If they areincluded in the Illaenacea, the superfamily iscertainly comprehensive and much remains tobe solved particularly of the effaced illaenoids.
The Illaenidae appeared already in the early
Ordovician and died out before the Devonian
period. The Scutelluidae on the contrary first
appear in the Middle Ordovician rocks in China
and the wide divergence has taken place fromthe Planiscutelluinae during the late Silurian
and Devonian periods. The trend of evolution
was quite different between the two families.
It is improbable that the Scutelluidae were
derived from the effaced Illaenidae or viceversa. In the Scutelluidae the effacement of
the cephalic segmentation for example is mostadvanced in the Meroperixinae (Upper andMiddle Silurian), but still the occipital lobe is
generally well defined. In the Illaenidae, on451

the contrary, the occipital furrow is completely
effaced, at least in the grown stage, although it
may remain in the immature stage as seen in
Faillaena clava Chatters ton and Ludvigsen,
1976, in fig. 34, pI. 6. Among the Illaenidae
Bumastus (Middle Ordovician-Silurian) reveals
particularly high specialization in the verybroad glabella and almost complete obliteration of the trilobation in the thorax and pygidium before the appearance of the Scutelluidae.
Distribution :-In the Ordovician fauna of
China this family is well represented by eightor nine genera in four or five subfamilies asfollows:
Illaenus, Stenopareia, Nanillaenus, Cekovia,
Trigocekovia and Spinillaenus in Illaeninae
Zdicella and Zbirovia in Ectillaeninae
Bumastus and Dysplanus in Bumastinae
Meitanillaenus in Goldillaeninae(?) Theamatopsis in Theamatopsinae
Trigocekovia Xia, 1978 whose monotypic
species is T. fonicatus Xia, 1978, resembles
Cekovia, but its cephalon is triangular in dorsal
view and hem~sphaeric in frontal view. Spinillaenus Xia, 1978, of which S. sinensis Xia, 1978
is monotypic is similar to Illaenus and OcWlaenus, particularly to the latter, but it has theconcave marginal border which is produced
postero-Iaterally into a triangular genal spine as
far as the third thoracic segment. The thorax
consists of 10 segments, instead of 8 in OcWlaenus. Meitanillaenus which appeared first in
Hunan in the late Ordovician age survived in
the Silurian period when Ptilillaenus andWuchuanella also occur.
In Japan the family is represented by many
Silurian species in two subgenera, Bumastus
and Bumastella of Bumastus beside a species
of Goldillaenus and two species of "Illaenoides"
which belong probably to a new genus.
Subfamily Goldillaeninae Balashova, 1959
In the Silurian fauna of China the family
Illaenidae is represented by the following threeendemic genera whose type-species are cited inbrackets.

452

Teiichi KOBA YASHI and Takashi HAMADA

Ptilillaenus Lu, 1962 (Ptilillaenus lojopingensis Lu, 1962)
Meitanillaenus Chang, 1974 (Meitanillaenus
binodosus Chang, 1974)
Wuchuanella Wu, 1978 (Wuchuanella quadrata Wu, 1978)
Ptilillaenus lojopingensis was primarily founded on cranidium and pygidium. A free cheekfrom Lojoping- which Chang (1974, pI. 18, fig. 8)
referred to this species has the lateral borderwhich is rounded at the genal angle and becomes
broader near the angle. A shallow marginal
furrow runs along the border. The axis of theassociate pygidium is long, conical and composedof 7-8 rings and a round terminal lobe. The
marginal border is nearly flat and moderate inbreadth.
Meitanillaenus binodosus from Meitan, Guizhou has the cranidium nearly as long as wideand strongly contracted at about one-fourthfrom posterior, much more expanded in anterior
than posterior where it is about half as broad as
the cranidium; a pair of semicircular nodes
present on fixed cheeks at glabellar constriction;eyes there opposed, fairly large; free cheek withnarrow lateral border and short genal spine.
Like Bumastus hypostoma is provided with apair of large triangular wings; central body subcircular and divided into two parts by middle
furrow and maculae. Thorax distinctly trilobed
by axial furrow; axial ring slightly broader thanpleuron. Pygidium semicircular; axis short andtrigonal; articulating facet distinctly limited bypleural furrow.
Meitanillaenus zhitangensis Wu, 1983 fromthe Upper Ordovician of Xijiang which has adistinct occipital furrow and lobe may be excluded from this genus. Wuchuanella quadrata
Wu is represented by two cranidia resembling
Meitanillaenus, but lacking alae.
Finally, the Brontocephalidae Kolobova,
1973, including Brontocephalus Tschugaeva,
1975, and Brontocephalina Tschugaeva, 1975,
but excluding Bulanaspis Tschugaeva, 1956 isallied to the Goldillaeninae. Its glabella is strongly expanded in anterior. Brontocephalina marginatula Tschugaeva, 1975, from the Upper

Ordovician of Northeast USSR has the pygidiumwith axial lobe composed of tre trapezoidal
short anterior part and the longer posterior
part but ill-defined near the terminal part.
Genus Illaenoides Weller, 1907
Illaenoides (?) magnisulcatus Kobayashi and
Hamada, sp. nov.

PI. 90, Figs. 5a-b.
Description:-Illaenoid pygidium gently convex except for marginal inclination, semiellipticalin outline, widest a little behind anterior margin
where axis is a little projected beyond pleurallobes; anterior margin forming a broad obtuse
angle near median point of pleural part; convexity of pygidium reaching maximum at about
one-third from posterior end. Axial furrow completely effaced; axial projection of anterior margin one-third as broad as pygidium. First pleuralfurrow extending postero-laterally from lateralend of axial projection and confluent with marginal furrow. Lateral border relatively thick for
illaenid. Facetted area defined by obtuse ridge
extending postero-laterally from median angulation of anterior pleural margin; lateral marginal
furrow tending to shallow backward. Sagittal
profile of pygidium in exfoliated part simply
arching down to posterior margin from highest
point. Test smooth.
Observation:-The pygidium is 8.5 mm longand 11 mm wide. The axial projection of theanterior margin measures 5.8 mm in breadthand the lateral border exclusive of marginal
furrow 1.8 mm.
Comparison:-The convex marginal border
and well developed marginal furrow are mostdistinctive of this pygidium, suggesting that
this pygidium belongs to an undescribed genus.
Among the Japanese trilobites it is similar toBumastus (Bu mas tella) bipuncatus, but thelateral margins converge more rapidly and themarginal border is depressed. It resembles also
the pygidium of Ptilillaenus lojopingensis Lu in
the presence of marginal border which is, however, not convex and the marginal furrow isabsent in that species.

819. Silurian Trilobites from Shikoku, 2nd Addition
Occurrence:- Yokokura-yama limestone; collected by Fujiyama.
Illaenoides (?) abnormis Kobayashi and
Hamada, sp. nov.
PI. 90, Figs. 6a-c.

Description :-Pygidium parabolic in outline,
broader than long in ratio of 1:0.7, regularly
and gently convex with narrow marginal borderand narrow marginal furrow; posterior one-thirdof pygidium distinctly arching down below thegeneral base level.
Observation and comparison:-The margin of
this pygidium is not perfect, but if perfect, it isprobably 19.3 mm long and 27 mm broad. The
axial part is about 13 mm wide on the articulating margin. The lateral border is 3 mm across.
Like the preceding pygidium the axial furrow
is completely effaced in this pygidium, but themarginal border is evidently narrower and not
so convex as in that pygidium. This pygidium
is more convex and distinctly bent down inposterior one-third. Because it looks the original
curvature, it is quite an unique aspect among
illaenids.
Occurrence:- Yokokura-yama limestone; collected by Mizuno.
Family Poetidae Salter, 1864
Subfamily Cyphoproetinae Pillet, 1969
Genus Cyphoproetus Kegel, 1927

Cyphoproetus latiaxis Kobayashi and
Hamada, sp. nov.
PI. 91, Figs. 1a-g, 2a-c.
Description:-Cephalon semicircular in outline, well convex except for distinctly concaveand smooth marginal border; glabella half as wide
as cephalon, subconical, but much more expanded in posterior and broadly arcuate in front;
anterior lateral furrow (2p) weak and short; posterior lateral lobe isolated by strong lateralfurrow (lp), which is long and straight, butpostero-laterally bent near its posterior end anddisconnected from occipital furrow; neck ring

453

composed of large median lobe and a pair of verysmall lateral lobes; occipital and axial furrows
profound; eye medium sized, semicircular closeset glabella behind anterior lateral furrow andprovided with distinct eye-socle; palpebral lobe
flat and simply slanting to axial furrow; preglabellar field short, arching down to concave
frontal furrow which is in turn bent up toward
frontal edge; facial sutures widely divergent
diagonally from eyes with postero·lateral convexity, incurved through margina,l furrow andrim a little beyond eyes' limit; posterior branchrunning backward along axial furrow, then becoming diagonal to cut the cheek border onparallels through eyes; narrow ridge runningalong posterior facial suture till it becomes
confluent with posterior cheek border; test
densely granulose.
Pygidium two·thirds as long as broad; anterior pleural margin obtusely angulated atmedian point; postero-lateral margin broadly
arcuate; posterior margin nearly transversal.
Axial lobe one-third as wide as pygidium, wellconvex, rapidly tapering back to rounded posterior end whence a short postaxial ridge issues;
six axial rings separated from one another by
deep ring furrrows. Pleural lobe divided into
five flat-topped ribs, all geniculate, less slanting
in inner part than outer part; first rib distinctly
facetted; succeeding ribs forked by narrowinterpleural furrows which are deepened in
distal part. Marginal border flat and smooth;
its inner side depressed where pleural and interpleural furrows die out. Test granulose.
Observation:-Because the occipital lobe ismesially exfoliated, it is indeterminable whether
or not a median tubercle is present. The test is
broken also near the genal angles. Therefore theweak incurvature of the cephalic margin, commonly seen near the angle in Cyphoproetus
cannot be ascertained in this specimen. Note·worthy is that a round-topped narrow ridge
resembling the so-called eye·socle by Owens,
1963 is running in this cephal on along the posterior branch of the facial suture as far as theposterior cheek border. On the left cheek anarrow ridge is clearly seen behind the eye.

454

Teiichi KOBA YASHI and Takashi HAMADA

The associate pygidium has a flat marginalborder of moderate breadth, but no distinct
marginal furrow on its inner side where pleuralribs and furrows terminate. The pleural ribs are
distinctly flat-topped in the inner part.
Comparison:-This cranidium belongs to thegroup of Cyphoproetus with a narrow preglabellar area and granulose test like Wenlockian
C. strabismus Owen, 1963, but the granulation is coarser and the cranidium and glabella
are broader and the eyes smaller in this species.
Occurrence:-Yokokura-yama limestone atGomi quarry; collected by Uchida.
Family Cheiruridae Angelin, 1854
Subfamily Sphaerexochinae tJpik, 1937
Beside Sphaerexochus and Pompeckia Whittington (1963, 1965), this subfamily included
Heliomera Raymond, 1905, Kawina Barton,
1915, Cydonocephalus Whittington, 1963, andXystocrania Whittington, 1965. Kolymella andParasphaerexochus are two additional genera
which were instituted by Tschugaeva, 1973,
Kawina plana Tschugaeva, 1964, and Parasphaerexochus galeatus Tschugaeva, 1973, being
the respective type-species. The former occurs
in Middle Ordovician and the latter in late
Lower Ordovician, both in Eastern Siberia,
Northeast USSR.
Genus Sphaerexochus Beyrich, 1845
This is an Ordovician-Silurian genus of Trilobita whose type-species is Sphaerexochus mirus
Beyrich, 1845. According to Lane (1971) itcomprises 23 other species including one doubtful reference. Most of them are European species,
but a quarter is American and one, namely,
Sphaerexochus taimyricus Balashova, 1960, is
an Asian species. S. mirus is reported fromTurkestan by Weber (pygidium, 1932, cranidium
and pygidium, 1951) and also from Australia
and North America. The authors added one
Silurian (up. Llandoverian-low. Wenlockian)species from Malaysia (1971) and two Ludolovian species from Japan (1974, 1976). Re-

cently Sphaerexochus fimbrisulcatus Lu, 1975,
and S. granosus Ju, 1983, were described from
the Middle Ordovician in East China, S. dongzhuangensis Zhou, 1982, from Middle Ordovician
in Shaan-Xi, Northwest China, S. luoheensis
Nan, 1980, from Upper Ordovician, Heilongjiang,Northeast China, and S. guizhouensis Wu, 1977,
from Lower Silurian, Guizhou, Southwest China.
Sphaerexochus tisulcatus Tschugaeva, 1973
occurs in Lower Ordovician of Northeast USSR.
Further additions were made from Canada
and Australia as follows:
Sphaerexochus arenosus Chatterton & Ludvigsen, 1976, from Middle Ordovician,CanadaSphaerexochus dimorphus Perry & Chatterton, 1977, from Middle Silurian, Canada
Sphaerexochus molongoensis Chatterton &Campbell, 1980, from Middle Silurian,Australia
Sphaerexochus lorum Chatterton & Campbell,
1980, from Upper Silurian, Australia
Sphaerexochus glaber Holloway, 1980, from
Middle Silurian, U.S.A.
Sphaerexochus brundlyi Chatterton, 1984,
from Middle Silurian, Canada
Sphaerexochus johstoni Chatterton & Perry,
1984, from Middle Silurian, Canada
These species are represented by many excellent specimens, some of which are silicified
and show their ontogeny. Thus our concept of
the genus was greatly clarified.
Finally, supplemented with Sphaerexochus
seto Lane and Owens, 1982, from the Silurian(up. Llandovery-low. Wenlock) from Greenland
it is nearly cosmopolitan, although none is reported from South America, so far as the authors
are aware.
Sphaerexochus hiratai Kobayashi and
Hamada, 1974
PI. 90, Figs. 7a-c.
1974. Sphaerexochus

hiratai Kobayashi and
Hamada, p. 88, pI. 6, figs. 6-10, pI. 7,
figs. 1-8, pI. 8, figs. 1-7, text-fig. 6F.

819. Silurian Trilobites from Shikoku, 2nd Addition

This species is well represented by many
cranidia and pygidia, but its free cheek, hy·
postoma and thorax were unknown. In a small
complete rolled shield from the same locality
the pygidium is seen to fit nicely the broadsinuate frontal margin of the cranidium.
Its thorax is tapering back slowly and composed of ten segments. The axial ring is nearly
as wide as the pleuron which is unfurrowed
and abruptly rounded near lateral extremity.
The anterior segments whose pleurae are moreor less geniculate are more convex than theposterior ones. Two small round bosses areseen on the left side of the sixth and seventh
axial rings at their antero-lateral ends, insteadof the postern-lateral ends.
The free cheek has a narrow rim and a broadconcave border outside the relatively small
cheek-roll. The anterior branch of the facial
suture runs forward and a little inward fromthe eye. Its posterior branch in form of a sutureridge extends from the eye to the genal angle
taking a broad arcuate course and meets thelateral margin just in front of the angle wherea very short rudimentary spine issues.
Occurrence:-Yokokura-yama limestone; collected by Kojima.
Subfamily Dindymeninae Kielan, 1959
non Henningsmoen, 1959
Family Dindymeninae was proposed in 1959
by Kielan to include Dindymene Hawle andCorda, 1847 and a new genus, Eodindymene
whose type-species is Dindymene pulchra Olin,
1906, but she excluded Plasiaspis Prantl andpHbyl, 1948. In the same year Henningsmoen
proposed also the Dindymeninae as a subfamilyof the Encrinuridae, but he included Plasiaspis
and possibly Prosopiscus Salter, 1865 beside
Dindymene in his subfamily. Because only theyear, 1959, is cited for their dates of publication, it is a question which of the family or
subfamily possesses the priority. The present
authors, however, consider that Plasiaspis andProsophiscus are to be eliminated out of thesubfamily, because these two genera have wellsegmented glabellae and Plasiaspis has more455

segments in its thorax and pygidium than Dindymene. Furthermore, the difference in theventral facial suture between Dindymene andPlasiaspis is emphasized by Kielan.
Incidentally, Dindymenella Lu, 1976, instituted on Dindymenella sulcata Lu, 1976, fromthe early Upper Ordovician in Yunnan differsfrom Dindymene as well as Eodindymene in thepossession of 12 segments in thorax, insteadof 10 segments in those two genera and theaxial lobe of the pygidium is comparatively
small and its spines are fairly stout and long.
The distinct lateral furrows in three pairs on its
glabella are quite distinctive from those twogenera.
Genus Dindymene Hawle and Corda, 1847
Recently Dindymene orientalis Zhou andD. deflecta Ju were described from the Upper
Ordovician, respectively in Jiangxi and Xijiang,
both Central China (Chou et al., 1983). Dindymene brevicaudatus Kolobova, 1972, is known
from the Upper Ordovician in Kazakhstan.
Dindymene (?) sp. indet.

PI. 91, Fig. 4.
A blind cephal on composed of large vaulted
glabella and less convex cheeks of moderate
size. Glabella suboval, broader in anterior, gradually narrowing in middle and posterior parts;
posterior margin transversal and nearly straight;
lateral furrows absent; a large median tubercle
present at a point slightly anterior to center ofglabella. Cheeks separated from glabella bynarrow straight axial furrows and opposed at
posterior half of glabella. Marginal furrow andrim very narrow. Small tubercles densely distributed on whole surface except for axial andmarginal furrows where they are absent or
sparce; medium sized tubercles scattered irregularly among small ones.
The genal and posterior parts of the cephalon
are obscure, but the antero-lateral outline can be
figured out by tracing the tuberculate marginal
rim and smooth marginal furrow. Little is knownof facial suture.

456

Teiichi KOBA YASHI and Takashi HAMADAThis cephalon belongs most probably to
Dindymene or the Dindymeninae s. str., but
it is quite distinct from most species of thegenus in its proportionally large glabella tothe cheeks and accordingly the much greater
forward projection of the glabella. The antero·lateral outline of the cheek is also different fromtypical Dindymene. These differences suggest
its subgeneric or generic distinction from Upper
Ordovician Dindymene in Europe.
Occurrence:-Gomi quarry of the Yokokura·yama limestone; collected by Fujiyama.
Genus IchiyameZla Kobayashi and Hamada,gen. nov.

Diagnosis:-Cephalon strongly vaulted, moreor less hemisphaeric and divided into a large
glabella and two small cheeks by gently arcuate
or nearly straight and forwardly divergent axial
furrows; no lateral furrows on glabella, but apair of short linear furrows cutting into glabella
from frontal margin; occipital ring relatively
large; cheek lacking eyes and facial stures.
Type-species:-Ichiyamella subglobula Koba·
yashi and Hamada, sp. nov.
Remarks:-The authors are of opinion that
Dyndymene (?) megacranidia is referable to thisgenus. No median tubercle is present on theglabella in these two species.
Distribution :-Silurian in Japan.
Ichiyamella subglobula Kobayashi and
Hamada, sp. nov.
PI. 91, Figs. 3a-f; Text-figs. AI-4.

Description :-Cephalon strongly vaulted, alittle broader than long. Glabella very large,strongly expanded forward from narrow baseand well rounded in anterior; lateral furrows
absent, but a pair of weak short linear furrows
inserted from frontal margin in two lateralparts of glabella; occipital ring much wider
than glabellar base and abruptly narrowing
near lateral ends; axial furrow diagonal andgently arcuate. Cheek ovoid, very small in comparison with glabella; marginal border thickened

A

2
Text-figs. A-C. Restoration of Three
Silurian Trilobites from Mt. Yokokura. (videp. 457)
Al-4: Ichiyamella subg/obu/a.

toward genal angle where it is remarkably hanging down below general level of cephalon; eyes
and facial suture absent; test densely granulate.
Observation :-The holotype cephalon is 14.7mm long, 16 mm broad and 7.5 mm high; its
glabella 11.5 mm long, 13 mm at the widest
and 4.7 mm at the base; occipital ring 3 mmlong and 7.5 mm wide; and its cheek 4.6 mm
wide, 5 mm high and 7.3 mm in diagonal length.
Because the diagonal furrows are confluent
with the occipital furrow far inside of the lateralextremities of the neck ring, they look similar
to the preoccipital furrows of Sphaerexochus.
In this cephalon, however, there are neither
axial furrows nor cheeks, if they be really theposterior lateral glabellar furrows. Therefore
these diagonal furrows are evidently the axial
furrows.
Occurrence :-Ichiyama, near Ochi-town; collected by Nakajima.

Ichiyamella megacranidia Kobayashi and
Hamada, 1985
1985. Dindymene (?) megacranidia Kobayashi

and Hamada, p. 213, pI. 28, figs. 3a--c.
This species agrees with the preceding in thetrilobation of the strongly convex cephalon, apair of short linear frontal furrows on the glabella, small cheeks and granulate test, but it is different from that species in the roundly quadrate
glabella limited laterally by nearly straight axial819. Silurian Trilobites from Shikoku, 2nd Addition
furrows. A very narrow marginal rim and alittle broader and deep intramarginai furrow
are also present in this species. The posterior
mrgin of this cephalon is unknown.
Occurrence :-Gomi quarry of the Yokokurayama limestone.

Tsungilichas Chang, 1974 from the Lower
Silurian of Guizhou, China was synonymized
with Dicranopeltis by Thomas (1978). Recently
Dicranopeltis cf. polytoma (Angelin) was reported by Zhou and Zhou (1982) to occur in
the Upper Ordovician of Inner Mongolia.

Family Lichidae Hawle and Corda, 1874
In the Ordovician fauna of Asia this family isrepresented by Metopolichas, Amphilichas, Acrolichas (?) and Lyralichas in China, Indochina,
Burma and Turkestan. Acanthopyge, Lobopyge,
Graspedarges and Lichas s.l. are reported fromthe Devonian of Western, Central and Eastern
Asia where in the last Acanthopyge is known
from the Amur region and Radiolichas (?) from
Mongolia, Lichas, s.l. from Viet-Nam. Graspedarges and Acanthopyge (Lobopyge ?) were
found in Japan. Compared to them, the occurrences of Silurian lichids are scarce in Asia.
Lichas (Gorydocephalus) in two species aredescribed from the Middle Silurian of Western
Mongolia by Tschernysheva (1937) and Acanthopyge (3 spp.), Acanth?ma, Tetralichas andDiscrnopeltis (?) from the Kuznetsk region andKazakhstan, Turkestan by Weber (1932, 1951),
Tchernysheva (1951), Maximova (1960) andBalashova (1968). Gorydocephalus is now
synonymized with Trochurus.

2
BI-3:

Apolichas trucatus.

CI-3:

Dicranopeltis tricornis.

Subfamily Lichinae Hawle and Corda, 1874
Genus Dicran op Ie tis Hawle and Corda, 1974
Phleger (1936) erected two new genera,
Dicranopletoides and Makromuktis on Lichas
decipiens Winchell and Marcey, 1865 andDicranopeltis nasutus Weller, 1907, respectively,
but they were synonymized with Dicranopeltis
by Warburg (1939) and this opinion was upheldby Tripp (1957 and in Moore's Treatise, 1959).
Makromuktis is Dicranopeltis having a mediananterior projection on the glabella. It is noteworthy that such forms occur not only in the
Niagaran in North America but in the Wenlockian in Britain and also in the Yokokurayama limestone in Japan.

457

Teiichi KOBA Y ASH! and Takashi HAMADA

458

Dicranopeltis tricornis Kobayashi and
Hamada, sp. nov.
PI. 92, Figs. la-g; Text-figs. Cl-3.

Description :-Cephalon large, broad, pentagonal, well vaulted, most elevated at a little anterior to glabellar center whence a stout medianspine issues, subvertical anterior to spine, but
gently sloping down backward; its length aboutfive-eights of cephalic breadth exclusive ofgenal spines; anterior margin nearly straight,
diagonal and forming a broad obtuse blunt
angle of about 110 degrees, slightly more protruded forward in median part than anterolateral parts; posterior margin almost as broadas anterior one, transversal, but more extended
back at neck ring and forming obtuse angles atlateral ends; antero-lateral and postero-lateral
margins meeting at genal angle whence a stout
long lateral spine issues.
Glabella large, occupying more than a half ofcephalon, broadest through a point a little posterior to median spine and broadly sinuate

toward mid-length of lateral glabellar lobes;
front-median lobe composed of laterally expanded frontal part, long conical middle partnarrowing back, and small posterior part expanding toward neck ring; lateral lobes quite
different in size; anterior lateral lobe long andoccupying major part; posterior lateral lobe
small but a little larger than basal lobe; dorsal
and occipital furrows stronger than lateral ones.
Fixed and free cheeks well fused; eyes small,located a little anterior to cheek-center; no eyeridge discernible; suture ridge extending laterally
from eye, describing an arc on cheek, thenfacial suture cutting diagonally through lateralborder as far as near intergenal angle; anterior
branch of facial suture apparently runningforward and inward from eye and crossing
doublure between frontal and antero-lateralparts of marginal border. Genal spine stout,
protruding in same line with antero-lateralborder and then slightly bent backward; posterior cheek border divided into two parts byintergenal angle at its median point.

Explanation of Plate 90
Japonoscutellum japonicum (Kobayashi and Hamada) puteatum, subsp. nov .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 448
a: dorsal, b: frontal and c: left lateral views of the holotype cranidium. PA 18071,
x2.0, colI. Kojima.
Figs. 2a--c. Japonoscutellum japonicum (Kobayashi and Hamada) laticephalum, subsp. nov .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 449
a: dorsal, b: frontal and c: right lateral views of the holotype cranidium. PA 18072,
x2.8, colI. Noda.
Fig. 3. Scutelloid, gen. et sp. indet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 449
A dorsal view of an almost complete shield with 10 thoracic segments, though they
were exfoliated. PA 18073, x3.4, colI. Kojima.
Figs. 4a-c. Opoa (?) trinodosa Kobayashi and Hamada, sp. nov. . . . . . . . . . . . . . . . Page 450
a: dorsal, b: frontal and c: right lateral views of the holotype cranisium. PA 18074,
x1.8, colI. Fujiyama.
Figs. 5a, bIllaenoides (?) magnisulcatus Kobayashi and Hamada, sp. nov. . . . . . . . . Page 452
a: dorsal and b: left lateral views of the holotype pygidium, PA 18075, x3.2, colI.
Fujiyama.
Figs. 6a--c. Illaenoids (?) abnormis Kobayashi and Hamada, sp. nov. . . . . . . . . . . . . Page 453
a: dorsal, b: posterior and c: left lateral views of the holotype pygidium showing anabnormal slant of the margin, PA 18076, x1.4, coli. Mizuno.
Figs. 7a-c. Sphaerexochus hiratai Kobayashi and Hamada . . . . . . . . . . . . . . . . . . . Page 454
a: left lateral, b: pygidial and c: cephalic dorsal views of an enrolled specimen ofsmall size, PA 18077, x4.1, colI. Kojima.
Figs. la-c.

KOBA YASHI and HAMADA: Silurian Trilobites from Shilwku

Plate 90

819. Silurian Trilobites from Shikoku, 2nd Addition

Fine granules distributed on test from glabella
to convex marginal border and three spines,
excluding various smooth furrows; smoothinterspace, however, wider than granules intotal space.
Observation :-This species is represented by ahalf of a cephalon which is, however, scarcely
deformed. If complete, the cephalon exclusiveof three spines is about 26 mm long, 40 mm
broad and 16 mm high.
These spines are all broken, but they arepresumably fairly long. The left eye is indicatedby an eye-scar whose outline is obscure on theaxial side. Its lateral and posterior margins are,however, traceable along the peripheral narrow
furrow of the eye-socle and also with reference
to the suture ridge behind the eye. The posterior
branch of the facial suture is seen in the further
extension, while its anterior branch is recognizable only on the doublure and its vicinity.
Comparison:-This species agrees with Lichas
scabra scabra (Beyrich, 1845), the type-speciesof Dicranopletis Hawle and Corda, 1947 in the
broad pentagonal outline of the cephalon with apair of lateral spines and the essential configuration of the cranidium, although the glabella ofthis species disagrees with the lectotype of
Dicranopeltis scabra in fig. 1, pI. 4 and Dicranopettis scabra propiqua (Barrande, 1846) restored
in text-fig. 13, both in Vanek (1959) in theoutline of the median lobe, the proportional
length of bicomposite lobe with the posterior
lobe and other minor details. The most important distinction of this species from Dicranopeltis
scabra is the possession of the median glabellar
spine.
Dicranopeltis polytomus (Angelin, 1854) withwhich Lichas elegans Tornquist, 1884 was synonymized by Warburg (1939) has also a similar
cranidium which, however, lacks the medianspine. Tsungilichas pustulosus Chang, 1974,
from the Lower Silurian of Guizhou is represented by a strongly vaulted cranidium resembling this species, but it has no spine on the somewhat longer glabella.
In regard of the median projection this species
is comparable to Makromuktis nasuta (Weller,

4591907), although it is not a proboscis-like projection in this species. Dicranopeltis woodwardi(Reed) and Dicranopeltis sp. I by Thomas (1981)
has an upwardly directed spine on the medianglabellar lobe which is bifurcated at a distancefrom its root. Lichas (Metopolichas) celorhin
Angelin var. coniceps Herz von Leuchtenberg byReed 1917 form the Ordovician of Shihtien,
Yunnan also has a median spine on the frontal
lobe of the glabella.
Occurrence:-Yokokura-yama limestone; collected by Asami.
Subfamily Homolichinae Phleger, 1936
Genus Apolichas Kobayashi and Hamada, 1974
Apolichas truncatus Kobayashi and Hamada,1974

PI. 92, Figs. 2a-c; Text-figs. Bl-3.
truncatus Kobayashi and
Hamada, p. 80, pI. 8, figs. 9-12; text-fig.

1974. Apolichas
6A.

Little has been known on the cheek of thisspecies. A new cephalon shows that a left cheekis separated from the tricomposite lateral lobeof the glabella by a deep axial furrow and overhanging the narrow marginal border. It is reniform and strongly vaulted, but its top is lower
than the lateral lobe. Its eye is very large forthe Lichidae.
Compared to the type specimen this cephalon
is somewhat different in the anterior outline.
The main lobe of its glabella is protruded forward, forming weak concavity on each side of
the frontal margin of the cephalon. Further·more the convexity of the glabella is a little
stronger, attaining the maximum shortly behind
its center. The posterior margin of the cephalon
is not well preserved and the genal spine which
existed on the left cheek of the type specimen
is lost in this cephalon. On the basis of thesetwo cephala combined the dorsal aspect of the
cephalon can now be restored completely.
Occurrence :--Yokokura-yama limestone. KACSt-0032(a) ; collected by Kaneko.

460

Teiichi KOBA YASHI and Takashi HAMADA
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Explanation of Plate 91
Figs. la-g. Cyphoproetus latiaxis Kobayashi and Hamada, sp. nov . . . . . . . . . . . . . . Page 453
a, e: dorsal, b, f: frontal, c: right lateral and d, g: left lateral views of the holotype
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a: dorsal, b: right lateral and c: posterior views of a pygidium, PA 18079, x7.0,
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a: dorsal, b: posterior, c: left oblique, d: frontal, e: right lateral and f: left lateralviews of the holotype cephalon, PA 18080, x2.0, coIl. Nakajima.
Fig. 4. Dindymene (?) sp. indet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 455
A fragmentary cranidium showing the granulation of various sizes, PA 18081, x3.3,
coIl. Fujiyama.

KOBA YASHI and HAMADA: Silurian Trilobites from Shilwku

Plate 91

819. Silurian Trilobites from Shikoku, 2nd Addition
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462

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1 pI.
See also Kobayashi and Hamada (1974 and1984) for other references here omitted.

Ertaogon ='~i'fl', Gomi 1\.1*, Guizhou yUH, Heilongjiang ,~1lI!,iL
iIlI.§, Kweichou (=Guizhou) iVH, Shaan·Xi MlI.§, Shihtien 1i(!i1J],~l¥l,
Xijiang WriI.

lchiyama mill,Tienshan xLII,

jiangxi
Yunnan* x:

pLlmt'lt~UlO) c/)v )vK:c='lt!Rltr.A.,O);~1 2 !fl1JQ :~(,ij I':' Ii 5 HI':' !'~-t 0 9 ~TlO)= ~ 1(1); ,ie
1I'~~n-n'0o .:to)5'b1':'1:l:6:lfrfl!t2:lfrilli:fillil;fl;;l:n-n'L,:lfr~ Ichiyamella {';l;0 o.:t 0) {un.:. f*ff:'f:¥c~O)t':6br::'lEliYllt.d!:7EO) [113*t.( 1" 2 {[iii;;/; 0 0 .:-. I? (1) ==::i;t;b. t I*l l'f-O);/; 0
flL I/.l.it.:.JiltI, 'L {, ET~frn& L L 1" 00 Ichiyamella, Japonoscutellum, Apolichas I:l:m{f.l:pj,:n*ltO)~-C';/;00'N:.j;fo-· ~Ill~.&±Explanation of Plate 92
Figs. la-g. Dicranopeltis tricornis Kobayashi and Hamada, sp. nov . . . . . . . . . . . . . . Page 458
a, d: dorsal, b, e: left lateral and c, f: frontal views of the holotype cranidium showing an inflated shield with long and stout spines, PA 18082, a, b, c x2.0, d, e, f x1.0,
coll. Asami.
Figs. 2a-c. Apolichas truncatus Kobayashi and Hamada . . . . . . . . . . . . . . . . . . . . Page 459
a: dorsal, b: frontal and c: left lateral views of a cranidium to show the trilobationand the large eye, PA 18083, x3.5, coIl. Kaneko.

KOBA YASHI and HAMADA: Silurian Trilobites tram Shikoku

Plate 92

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 463-474, pIs. 93-94, September 30,1986

820. ON NEOCRIOCERAS SPINIGERUM (JIMBO),
A SPECIES OF CRETACEOUS HETEROMORPH AMMONOIDS(STUDIES OF THE CRETACEOUS AMMONITESFROM HOKKAIDO-LV)*TATSURO MATSUMOTO
c/o Department of Geology, Kyushu University, Fukuoka 812
KIKUWO MURAMOTO, TAKEMI TAKAHASHI,
MINORU YAMASHITA and YOSHITARO KAWASHITA
c/o Museum of Mikasa City, Hokkaido 068-21

Abstract. Neocrioceras spinigerum (Jimbo), the type species of Neocrioceras,
is redefined on the basis of the specimens in our collections as well as the hitherto
described ones. Its adult body-chamber is spirally coiled in the same way as thephragmocone. It is quadrituberculate at intervals and the disposition of the ventrolateral tubercles is variable (opposite or alternate or irregular) among individuals
and also with growth-stages. The quadrilobate suture is florid; I trifid and as deepas U; other elements bifid and of dove-tail form. Reliable records show that thisspecies occurs in the Santonian of Hokkaido and Sakhalin. Comparison with otherspecies is described at length, with a result that certain species previously referred
to Neocrioceras should be transferred to other genera.
In an appendix the genus Neocrioceras is redefined and its relationships withother genera are discussed.

to describe this species in detail with more
illustrations.
In this study the senior author is primarily
responsible for the paleontological description
and discussion; all the authors were engaged
in field work of collecting fossils in respective
areas concerned and in developing specimensfrom calcareous nodules.

In troduction
For some reasons, the genus Neocrioceras
Spath, 1921 has been understood inadequately
by some authors. To remove this misunderstanding, the first step is to redescribe its typespecies, Crioceras spinigerum Jimbo. This has
been done concisely in a recently publishedpaper (Matsumoto, 1985), in which only two
syntypes of Jimbo have been photographedbecause of the page limitation. This paper is

Description of a species

Neocrioceras spinigerum (JimbQ)
PI. 93, Figs. 1-8; PI. 94, Figs. 1-16;
Text-fig. 1*Received July 17, 1985; Read February 2, 1986
at Sendai.
463

464

Tatsuro MATSUMOTO et ai.

1894. Crioceras spinigerum Jimbo, p. 184, pI.
24, figs. 1, la, 1 b.
1921. Neocrioceras spinigerum (Jimbo); Spath,
p.51.
1933. Neocrioceras spinigerum (Jimbo); Shimizu, p. 13, pI. 2, figs. 4-9.
1963. Neocrioceras spinigerum (Jimbo); Matsumoto, p. 45, pI. 67, figs. 1, la, lb.
1981. Neocrioceras spinigerum (Jimbo); Tanabe
et al., p. 220, pI. 37, figs. 2a, b; text-fig.
5.
1985. Neocriocnas spinigerum (Jimbo); Matsumoto, p. 56, text-figs. 1, 2.
Lectotype:-UMUT. MM7521 (Jimbo, 1894,
pI. 24, fig. 1, 1a; Matsumoto, 1985, figs. lA, 1B;
this paper, PI. 93, Figs. 1-2) from Ikandai of the
Urakawa area (southern central Hokkaido) ,designated by Matsumoto, 1985 (p. 56).
UMUT.
Other
materiai:-Paralectotype,MM7522 (Jimbo, 1894, pI. 24, fig. 1b; Matsumoto, 1985, figs. 2A, 2B; this paper PI. 93, Figs.
3-5) from Ikandai. TTC_ 400819 (this paper,
PI. 94, Figs. 1-2) from Ikdandai; UMUT.
MM7640 from the Sosushi, a branch of theSanushibe, Hobetsu area (H. Yabe CoIl.); TTC.
360806 (PI. 94, Figs. 6-9) from a branch stream
called the Y-sawa of the Kikume-zawa, a tributary of the Ikushumbets, Mikasa district; UMUT.
1-3531 (TMC_) from loc_ T 55 p and UMUT. 13582 (TMC) from loco T 277 e of the Saku-Abeshinai area (see Matsumoto, 1942, for the numbered localities of TMC); TTC. 360815 (PI. 94,
Figs. 14-16) from the Saku-gakko-no-sawa, twoother unnumbered specimens of MYC. and
YKC., all the above six from the Saku-Abeshinai
area in the Teshio Nakagawa district; MC. Hbo3695 (PI. 93, Figs. 6-8) from the Haboro area.
YKC. (without number) from the Obira area;
H. Maeda's collection from the Onkono-sawa,
Kotanbetsu area_ Also UMUT. MM7708 (M.
Kawada ColI.) and TMC from the Naibuchi area;
IGPS_ 36836 from Namikawa (Shimizu, 1933,
pI. 2, figs_ 4-9; this paper Text-fig. 1); GK.
H5204, GK. H5205 (PI. 94, Figs. 3-5) and
GK. H5218 from Kawakami (S. Nagaoka CoIl.),
all the above six from South Sakhalin; NSM.
PM9491 (Tanabe et ai., 1981, pI. 37, figs. 2a, b;

text-fig. 5) from somewhere in Sakhalin (?).
Repositories are as follows with abbreviationin parentheses:
Geological Collections, Kyushu University,
Fukuoka (GK).
Institute of Geology and Palaeontology,
Tohoku University, Sendai (IGPS)
National Science Museum, Tokyo (NSM)
University Museum, University of Tokyo(UMUT)The Muramotos' Collection (on display at theMuseum of Mikasa City), Mikasa (MC)Takemi Takahashi's Collection, Mikasa (TTC)
Minoru Yamashita's Collection, Mikasa (MYC)
Yoshitaro Kawashita's Collection, Mikasa(YKC)
The specimens collected by T. Matsumoto(TMC) are kept at GK and UMUT.
Diagnosis:-Shell medium-sized, consisting ofslightly open crioceratoid planispiral whorls,
which are depressed in cross-section_ Bodychamber about three fifths of the last whorl.
Ribs fine and numerous, gently projectedon the dorsum, sinuous on the dorsolateral part,somewhat prorsiradiate on the flank, crossing
the venter nearly vertically. Spiny tubercles infour rows at intervals, with looping of minor
ribs. The ventrolateral tubercles normally opposite, resting on low major ribs, but sometimes
deviate slightly or even considerably from thisdisposition to give rise to irregular or alternate
configuration, thus variable among individuals
and/or with growth-stages_ The ventrolateral
tubercles often more numerous than the lateral
tubercles by addition of intercalated ones. Depending on the frequency of tuberculation, theintervening nodeless ribs vary from one to four,occasionally showing outward branching_
Sutural formula E, L., U, I; lateral saddles
and lobes (L, U) bifid and of dove-tail shape;
I nearly as deep as U and trifid; all the elements
finely and deeply incised.
Description:-A completely preserved adultshell is very rare_ MC Hbo-3695, about 75 mm
in diameter, is probably an adult shell, in which
body-chamber, although partly destroyed, occupies about 2100 and its last septum is at 48 mm

820. Neocrioceras spinigerum (Jimbo)

465

Measurements (in mm):
Specimens

DiameterUmbilicus

HeightBreadth

B/H

Lectotype
Paralectotype*
Paralectotype -180°
TTC.400819
TTC. 360806*
TTC.360815
TTC.360815*
MC. Hbo-3695
MC. Hbo-3695*
TTC.370817*
UMUT. MM7640
GK. H5205
GK. H5204
GK. H5118
IGPS. 36836*

34.4
39.5
31.0
24.4
26.6
42.5
36.0
75.0
48.0

18.3 (.53)
19.7 (.50)
13.8 (.45)
11. 7 (.48)
12.8 (.48)
18.0 (.42)
16.8 (.47)
37.0 (.49)
22.5 (.47)~22.5 (.52)
11.4 (.54)
15.6 (.47)
13.2 (.48)
16.0 (.47)
22.8 (.45)

9.5(.28)
11.6 (.29)
8.5 (.27)
7.8 (.32)
9.0 (.34)
13.8 (.32)
12.4 (.34)
~21
(.28)
16.3 (.34)
~15
(.35)
5.6 (.27)
10.8 (.33)
8.4 (.30)
10.0 (.29)
16.5 (.33)

14.5 (.42)
15.3 (.39)
11.0 (.35)
11.6 (.48)
14.4 (.54)(distorted)
18.2 (.51)
34.5 (.46)
24.2 (.50)~23(.53 )
8.0 (.39)
15.2 (.46)
11.7 (.42)
14.4 (.42)
24.5 (.49)

1.53
1.32
1.29
1.49
1.60

~43

20.4
33.2
27.7
34.0
50.0

1.47
~1.6

1.48
~1.5

1.43
1.41
1.40
1.44
1.48

* measured at the end of the phragmocone
approximate~

in shell diameter. It should be noted that the
body-chamber is coiled in the same form as thephragmocone and never extended to a straightshaft as in Anisoceras. This denies the presumption of Spath (1921, p. 51).
IGPS. 36836, described by Shimizu (1933,
p. 13, pI. 2, figs. 4-9), is also an adult shell,
but its body-chamber is incompletely preserved
and secondarily squashed.
The lectotype is wholly septate. Some otherspecimens, e.g. paralectotype, TTC. 360815,
TTC. 370817, TTC. 360806, GK. H5118 and
GK. H5204, show the last septum at smaller
diameters, from 40 mm to 25 mm, and only aportion of the body-chamber is preserved. They
may be more or less immature but some ofthem may be small adult shells. On the probably
adult body-chamber the ribs and tubercles are
somewhat coarser than those on immature shells,
but the difference is by no means great and it
is difficult to know the variation in size.
The ammonitella and early immature shell
are not preserved in mot cases. NSM. PM 9491,
reported by Tanabe et al. (1981), may be a comparatively better preserved example, but is yet
incomplete in lacking the very initial conch.

The observable part of the ammonitella is subhelical and soon followed by a nearly planispiralcrioceratoid whorl.
Shimizu's (1933) specimen was actually reconstructed with artificial connection between
the outer and inner whorls. Therefore, theslightly helicoidal form in his diagram is doubtful.
The whorl is characteristically depressed, but
the ratio of B/H is considerably variable from anindividual to another and also with growth-stage.
The measurements show that B/H ranges from
1.3 to 1.5 in many cases but occasionally exceeds
beyond that range. Anyhow, the ventral part isbroad and its central zone between the tubercles
is nearly flat, whereas the flanks are convex,
with the maximum breadth between the rows oflateral tubercles.
In the diagram of whorl-section by Jimbo(1894, pI. 8, fig. 1a) the four tubercles areillustrated as equidistant, but in the actual
specimen the distance between the left andright ventrolateral tubercles is wider than thatbetween the lateral and ventrolateral ones. The
same feature is observable in almost all the
specimens examined. In this respect the diagrams

466

Tatsuro MATSUMOTO et al.

by Shimizu (1933, pI. 2, figs. 7, 8; disorderedin the explanation of plate) as well as Text-fig.
1C in this paper are correct.
The disposition of tubercles is considerably
variable in this species. Some authors regarded
the alternate disposition of the ventral (i.e.
ventrolateral) tubercles as a diagnosis of Neocrioceras. This came probably from Shimizu's(1933) specimen whose illustration was citedin the first edition of the Treatise (Wright in
Moore (ed.), 1957). In this specimen, IGPS.
36836, the ventrolateral tubercles are alternate
on the figured part (Shimizu, 1933, pI. 2, fig.
5), i.e. on the last half of the phragmocone
and the first 30° of the body-chamber, butthey are opposite or nearly so on other observable parts. At the changing point from thealternate to opposite disposition on the bodychamber a few ribs cross the venter obliquely
for adjustment (Text-fig. 1B). Incidentally thehighly squashed last part of the body-chamberof this specimen was excluded from the illustration by Shimizu (1933, pI. 2, fig. 4).
In MC. Hbo-3695, the ventrolateral tubercles
are opposite on the body-chamber but irregular,
if not quite alternate, on the late part of thephragmocone (see PI. 93, Figs. 8, 7). In thelectotype and many other specimens (e.g. TTC.
360815, TTC. 370817, TTC. 360806, UMUT.
MM7640, UMUT. 1-3581, GK. H5118, etc.)
the ventrolateral tubercles are opposite ornearly so. The paralectotype has also the opposite ventrolateral tubercles in its young stages
but on the preserved last half whorl there is noleft ventrolateral tubercle and the ribs run
irregularly. This is unusual and interpreted asprobably pathologic. In GK. H5205, which iswholly septate, the ventrolateral tubercles aredisposed irregularly or rather alternate in thelater part, but in the younger part they areopposite and rest on the low major rib. In GK.
H5204 they are opposite on the phragmocone
and tend to deviate from that disposition on the
succeeding part of the body-chamber.
The lateral tubercles number 7 or 8 in a half
whorl, occurring as a rule at regular intervals
but occasionally at a longer or shorter interval

than the normal or average ones. In addition to
the ventrolateral tubercles which correspond to
the lateral ones, more ventrolateral tubercles
occur in an intercalated disposition variably
in some specimens or at some growth-stages.
The tubercles in the late growth-stage have along spine with a septate node at the base. The
lateral nodes are somewhat larger than theventrolateral ones. The ventrolateral spines
stretch upward and forward; the lateral ones
sideward.
The ribs are mostly simple and some of them
are looped at the tubercles. Occasionally theribs are bifurcated outward.
The curvature of the rib, as described in the
diagnosis, is indeed characteristic of this species,
but the degree of curvature varies to someextent. The feature of the curvature, such as
dorsal projection, dorsolateral sinuosity andforward inclination on the flank, are generally
weak on younger whorls, becoming more distinct
later. The ribs cross the siphonal zone of theventer nearly vertically between the rows ofventrolateral tubercles and some of them, loopedby the tubercles, show a faint sinus (i.e. backward arc). There are also indistinct constrictions behind some of the major ribs which have
opposite tubercles.
The sutural pattern described in the diagnosis
is kept constantly. Jimbo (1894, pI. 24, fig. 1b)
illustrated correctly the general pattern but
ignored minor incisions, probably because thespecimen (paralectotype) was somewhat eroded.
Shimizu's (1933, pI. 2, fig. 9) illustration showsminor details as well as the general pattern.
Some more figures (Text-figs. lA, D) are addedin this paper. Even in small, immature specimens
the suture is fairly deeply and finely incised.
In mature shells the suture is florid, with deep
and fine incisions and narrowed stems andbranches of the elements. Spath (1921) misunderstood that suture of this species was simple.
Occurrence :-Some of the above described
specimens are not precisely located in the stratigraphic sequence; others from the Upper Urakawan (K6b), approximately Santonian, andoften associated with Sphenoceramus naumanni

820. Neocrioceras spinigerum (Jimbo)

467

0':': : . _ .

c

B
L

,,
,

I

,,
I I

, I

I

",

P
I

I

,

I,,'

I

,

I

I

I

I

o
Text-fig. 1. Some features on IGPS. 26836, an example of Neocrioceras spinigerum(Jimbo).
A. External lobe of the last third suture. B. Disposition of ribs and tubercles on posterior
half of the body-chamber. C. Diagrammatic whorl-section at D=50 mm. (Spines restored inwhich left ones (dotted lines) projected on the same plane). D. Suture at a young stage(H=5.4mm, B=7.5 mm). Scale bar=10 mm (A-C); 1 mm (D). These figures are to supplement Shimizu's (1933, pI. 2, figs. 4-9).
(T.M. de lin. )(Yokoyama) and S. nagaoi (Matsumoto et Ueda).
Besides the described specimens, fossils of
this species may be found occasionally from theUpper Urakawan (Santonian) of various areasin Hokkaido and also Sakhalin, although they
are often hardly taken out in a complete shapefrom calcareous nodules.
The true stratigraphic range of this species
beyond the Santonian and the geographic distribution outside Hokkaido and Sakhalin are
yet uncertain.
Comparison:-The specimens from Pondoland
described by Spath (1921, p. 52, pI. 7, figs. 6ac) under Neocrioceras cf. spinigerum are not
identified with the named species, because they

have compressed or nearly circular whorls.
They seem to be closer to N. (?) undulosum
Matsumoto, 1977, but are too incompletely
preserved for a precise identification.
Neocrioceras(Schlueterella) compressum
Klinger (1976, p. 74, pI. 33, fig. 5; text-figs. 8j,109), from the Lower Santonian of Zululand,
was established on a fragmentary whorl, curved
gently on a plane. The holotype clearly showsa change in the curvature which suggests a subelliptical coiling; its whorl is laterally compressed
and its suture has a very small I. These arefundamentally different from the charactersof N. spinigerum. As in N. spinigerum the holotype has ventrolateral and inner lateral tubercles

468

Tatsuro MATSUMOTO et al.

connected by looped ribs and intervening two
to three nontuberculate ribs. As only a single
fragmentary specimen was available, no remark
was given on the variation. This Zululand species
is provisionally referable to Schlueterella Wiedmann, 1962 (emend. Matsumoto and Miyauchi,1984).
The specimens from the Lower Santonian of
the Gosau Group (Austria) described by Immel
et al. (1982) under Neocrioceras (Schlueterella)compressum seem to show an open subellipticalcoiling (op. cit., pI. 10, fig. 1a), but this might
have been affected by secondary deformation
as suggested by some of the normally coiled
ammonites from the Gosau. A fragmentaryphragmocone M 2 illustrated in the same paper(op. cit., pI. 9, fig. 3) indeed resembles the holotype from Zululand, but the body-chamber
and also the late part of the phragmocone (op.
cit., pI. 10, figs. 2-4; pI. 11, fig. 3) show a zigzag
arrangement of tubercles in the lateral andprobably also ventrolateral rows. This may be
another characteristic of S. compressa. The samecharacter occurs more distinctly in S. kawadai
Matsumoto et Miyauchi, 1984, from the Campanian of Hokkaido and Sakhalin, which differsfrom contemporary European species S. pseudoarmata (SchlUter, 1872) at this point. Regrettably the suture is not well exposed on the Gosau
specimens, but it is essential for a precise identification. In this paper, however, the Gosau specimens are provisionally identified with Schlueterella compressa. Anyhow, it is evidently distinct
from Neocrioceras spinigerum.
N. (Schlueterella) mutinodosum (SchlUter,
1872) (see Wright, 1979), from the Turonian ofEurope, should be referred to Pseudoxybelocerasor P. (Christophoceras), because four rows oftubercles occur not only on major ribs, withlooped riblets, but also on intervening minor ribs.
Neocrioceras (Neocrioceras) maderi Immel,
Klinger et Wiedmann, 1982 (p. 24, pI. 9, fig. 2;
pI. 11, figs. 1-2), from the Lower Santonianof the Gosau Group, is distinguished from N.
spinigerum in its more gently arcuate and compressed instead of depressed whorls and outward shifted rows of tubercles (i.e. ventrolateraland ventral rows). Its ribs do not show such a
dorsal projection and dorsolateral sinuosity as
those of N. spinigerum. Again its suture is notknown. This Gosau species is somewhat similar
to Kawashitaceras obiraense Matsumoto (1984b,
p. 342, figs. 2, 3A, 3B, 4A, 4B), but whether
the summit of the ventral tubercle is minutely
subdivided or not should be examined on better
preserved specimens. Anyhow, it is doubtful torefer this species (maderi) to Neocrioceras.
Ancyloceras kossmati Simionescu, 1899 (p.
257 [21], pI. 1, figs. 6-8), from the Upper Cretaceous of Transilvania (Roumania) was referred
to Neocrioceras (Schlueterella) by Wiedmann(1962, p. 205). It has widely open whorls coiled
subelliptically on a plane and the whorl seems
to be higher than broad, although the specimens
may be secondarily deformed. Quadrituberculate
major ribs and intervening finer ribs without
tubercles encircle the whorl radially without
sinuosity and the ribs are looped only occasionally at a few tubercles. The ventral tubercles
are opposite on the figured specimen (Simionescu, 1899, pI. 1, figs. 8A, B). Again the suture
is not illustrated. This species is similar to S.
compressa in shell-form but differs in its infrequent occurrence of looped ribs at a few tubercles. It is provisionally referred to Schlueterella,
unless enough evidence be obtained for theestablishement of a new genus or subgenus.
There are a few specimens (e.g. Hokkaido
University Collection 9467 with uncertain
record of locality) which may represent an
unnamed new species of Neocrioceras. This isdistinguished from N. spinigerum by a larger
size (about 100 mm in diameter of the phragmocone), more open spiral form, subcircular whorlsection and outward shifting tendency of lateral
tubercles in a late growth-stage. More specimens
with reliable locality records are wanted todefine clearly this species.
N. (?) undulosum Matsumoto, 1977, fromthe Turonian of Hokkaido, is atypical in thatits immature shell shows a low helicoid shapeand is followed by an open spiral shell in an
undulating plane, but its body-chamber is gently
curved on that plane without a retroversal hook.

820. Neocrioceras spinigerum (Jimbo)

Its ornamentation is fundamentally similar tothat of N. spinigerum.
Appendix
Remarks on the genus Neocrioceras
TATSURO MATSUMOTO
The genus Neocrioceras was proposed by
Spath (1921, p. 51), but the legal designationof Crioceras spinigerum Jimbo, 1894 as its typespecies was settled by Diener (1925, p. 192).
Since then this genus has been comprehended
widely and was even misunderstood by someauthors.
In the preceding pages the type species has
been defined clearly and compared with otherspecies. In this appendix I attempt to give ageneric diagnosis of Neocrioceras and remarkson the relationships of this genus with othergenera. The latter problem is still difficult, because heteromorph ammonite species of thiskind are often based on incomplete material.
Therefore, the following remarks, which arein part cited from the recently issued papers(e.g. Matsumoto, 1977; 1984a, b; Matsumotoand Miyauchi, 1984) with some modification,
may still be tentative.
Generic diagnosis:-Whorls in the main
growth-stages more or less open crioceratoid,
coiled in a plane or nearly so, ornamented withnumerous ribs and four rows of spiny tubercles,
i.e. the lateral and ventrolateral ones, at intervals, with intervening nodeless ribs; typicallyribs looped at the tubercles. Suture deeply andfinely incised, consisting of E, L, U, I; I trifid
and as deep as U; other elements bifid and of
dove-tail pattern.
Distribution:-From Turonian to Santonian(possibly to Lower Campanian) in Japan and
Sakhalin, so far as reliable records are concerned.
Relationships of Neocrioceras with othergenera:-The phylogenetic origin of Neocrioceras is uncertain. It was tentatively sought in
the plastic and long-ranging genus Hyphantoceras Hyatt, 1900 (Matsumoto, 1977), but this469

is a working hypothesis suggested by N. (?)
undulosum Matsumoto. Neocrioceras has noaffinity with Anisoceras Hyatt, because its
body-chamber is spirally coiled in the same
way as its phragmocone without straightly
extended arm and because the suture is different in its well developed I and also because
there is no species which could suggest the connection.
Schlueterella Wiedmann, 1962 is based on
the incompletely known type species, Anisoceras pseudoarmatum SchWter, 1872 from theCampanian of Germany, and supplemented bya closely allied species S. kawadai Matsumotoet Miyauchi, 1984, from the Campanian of thenorthern Pacific region. Its revised genericdiagnosis has recently been given by Matsumotoand Miyauchi (1984, p. 59). It is similar to Neocrioceras in the general mode of ornamentation(four rows of tubercles at intervals with intervening untuberclate ribs and with looping ofribs at the tubercles) but different in its open,
subelliptically (in stead of spirally) coiled whorls,
which consist of nearly straigt or gently arcuate
long shafts and U-curved parts, and also inhaving very small I in the suture. The mode ofdisposition of ventral tubercles does not work
as a criterion. The shell-form of the type species
and S. kawadai in early immature stage is notknown precisely and the phylogenetic originof this genus is uncertain. Anyhow, on the
ground of the distinction mentioned above,it is better to treat Schlueterella as an independent genus rather than to affiliate it to
Neocrioceras as a subgenus.
Kawashitaceras Matsumoto, 1984 was proposed for the type sepcies, Neocrioceras dentatum Matsumoto et Obata, 1981, represented
by a peculiar body-chamber from the Upper
Turonian of Hokkaido, and another species
K. obiraense Matsumoto, 1984, from the Upper
Turonian (or possibly Coniacian) of Hokkaido.
It has also quadrituberculate looped ribs andintervening nodeless ribs, but the tubercles
are distributed restrictedly on the ventral partand of dental form with minute crenulations
on their backward sloping summit. The tuber-

470

Ta tsuro MA TS UM OTO e tal.

cles have long spines in Neocrioceras and
Schlueterella, which would work as protection
against enemies. The body-chamber of Kawashitaceras is long and broadly arcuate. The
shape of the preceding phragmocone is notcompletely known but may be loosely open
and subelliptically coiled as judged from somepreserved pieces of curved parts. The peculiar
tubercles of a dental form distributed on theventral part may be favourable for this animalliving in a shell of very unstable shape to repose
himself (or herself) on the bottom sediments
and also for him to start for occasional locomotion back upward in the sea water (Matsumoto, 1984b). The phylogenetic origin of
Kawashitaceras is yet uncertain. Anyhow,
Kawashitaceras is distinct enough to be treatedas a genus independent of Schlueterella or
Neocrioceras.
Pseudoxybeloceras Wright et Matsumoto,
1954, established on the type species, Hamites
quadrinodosus Jimbo, 1895 (emended by Matsumoto, 1977), had nearly straight parallel shaftsconnected by U-curved parts. Numerous simple
ribs encircle the shell, of which some are periodically stronger than others. Every rib on theshell of late grwoth-stages have tubercles infour rows, ventrolateral and lateral ones. The
major or stronger ribs have stronger tubercles
where riblets are looped in some case but notalways so. Anyhow, the presence of tubercles
on every rib is one of the criteria to distinguish
Pseudoxybeloceras from Neocrioceras and
Schlueterella. The suture of P. quadrinodosum
figured by Jimbo shows shallow I, but that of
P. lineatum (Gabb) (Matsumoto, 1959, fig. 79)
clearly shows that I is as deep as U, being fairly
similar to that of Neocrioceras spinigerum. I issomewhat smaller than other lobes as Matsumoto(1959, p. 162) described previously but not sosmall as that of Schlueterella kawadai. With
respect to shell-form, Pseudoxybeloceras may becloser to Schlueterella than to Neocrioceras,
but the shell-form in the early immature stage
is not precisely known in both Pseudoxybeloceras and Schlueterella.
Parasolenoceras Collignon, 1969, based on thetype species P. splendens Collignon, 1969, fromthe Campanian of Madagascar, has ventrolateral
tubercles alone on every rib but no lateral tubercle at any growth-stage. In its suture I is as deepas U. With respect to the shell-form and suture
Parasolenoceras is similar to Pseudoxybeloceras.
Periodic major ribs with stronger tubercles
are not recognized in the holotype of P. splendens, but they appear more or less clearly in otherspecies, e.g. P. periodicum Matsumoto et Miyauchi, 1984, P. compressum (Henderson, 1970)
and P. nanaimoense (Ward et Mallory, 1977)
and fine ribs mayor may not be looped at the
tubercle. As in the case of Pseudoxybeloceras,
this character alone would not be sufficient toseparate a genus from Parasolenoceras.
Lewyites Matsumoto et Miyauchi, 1984 (p.
64), established for Idiohamites (?) oronensis
Lewy, 1969 (type species), I. (?) circularis
Lewy, 1969, and Hamites (?) taylorensis Adkins,
1929, is distinct in that periodic major ribs
have ventrolateral tubercles where minor ribs
are looped and that intervening minor ribs haveno tubercles. As in the suture of Schlueterella,
I is much smaller and shallower than U.
Hitherto reported species of Lewyites andParasolenoceras are all Campanian in age, where·
as Schlueterella and Pseudoxybeloceras appeared
earlier and ranged up to the Campanian. The
lateral tubercles do not appear in young shellsof Ps. quadrinodosum and occur infrequently
in Ps. linea tum. These facts suggest that Parasolenoceras may have derived from Pseudoxybeloceras. Likewise, Lewyites may have branchedfrom Schlueterella.
The degree of differentiation into major
and minor ribs varies from a species to another.
Therefore, there is little reason to maintain
Christophoceras Collignon 1969 (type species
C. ramboulai Collignon, 1969) as a genus independent of Pseudoxybeloceras. Some authors(e.g. Klinger, 1976) may keep Christophoceras
as a subgenus of Pseudoxybeloceras, but in thetype species of Pseudoxybeloceras itself someribs are somewhat (but not much) stronger thanothers and can be regarded as a kind of majorribs.

820. Neoerioeeras spinigerum (Jimbo)

As suggested by Ward and Mallory (1977),
Solenoeeras Conrad, 1860 (type species Hamites
annulifer Morton, 1842) may be related to Parasolenoeeras. It is characterized by a smaller shellwith greater appression of shafts. Lewy (1967)
has shown a gyro conic spiral form of a youngshell in a species of Solenoeeras from Israel.
Oxybeloeeras Hyatt, 1900 (type species
Ptyehoeeras erassum Whitfield, 1880) was
suppressed by some authors as a synonym of
Solenoeeras but is kept by Matsumoto andMorozumi (1980, p. 20) as a subgenus of Solenoeeras for its larger size and absence of constrictions. Solenoeeras is world-wide in the Campanian and Maastrichtian.
Exiteloeeras Hyatt, 1894 (type species Aneyloeeras jenneyi Whitfield, 1880) has also
ventrolateral tubercles on every rib and seems
to be loosely coiled earlier in a subelliptical form
and later in a crioceratoid spiral (according to
the restroration by Scott and Cobban, 1966;
Gill and Cobban, 1973, p. 10, fig. 6). The sutureof E. jenneyi (cited by Wright, 1967, fig. 251-7c
from Whitfield, 1880) is similar to that of Neoerioeeras spinigerum in being deeply and finely
incised and having bifid, dove-tail major elements and pointed I as deep as U. Exiteloeeras
may have an affinity with Parasolenoeeras, but
its true phylogenetic origin is uncertain.
The very initial stage of E. jenneyi has notbeen known precisely, but Gill and Cobban(1973, p. 10) observed nearly straight (or broadly arcuate) limbs connected by a semicircularbend in a juvenile (but not initial) stage. Thissuggests an affinity with such species as Parasolenoeeras tomitai Matsumoto, 1984a, which,however, is based on incomplete specimens.
The two species are nearly contemporary (Upper
Campanian), but known in much separate provinces, one from Wyoming and the other fromJapan. E. uneiforme Lewy, 1969, from the Campanian of Israel, shows a crioceratoid phragmocone and an unciforme body-chamber. E. etequenee Lewy, 1969 is peculiar in showing looped
ribs at the tubercles and inervening nodeless ribs
as in Lewyites, to which genus it could be transferred.

471

Axonoeeras Stephenson, 1941, established for
A. eompressum Stephenson, 1941 (type species)
and other species from the Neylandville Marl(possibly uppermost Campanian) of Texas, looks
similar to Exiteloeeras in the mode of coiling,ribbing and tuberculation, but its whorls are
irregularly in contact or partly open. In bothgenera the initial part is hooped. The suture has
not been illustrated for any species of Axonoeeras. In the type species almost all the ribs haveventral tubercles on either side of the siphonalzone, but occasionally a few nodeless ribs may
be intercalated. On the last part (probably adultbody-chamber) of A. pingue Stephenson, 1941,
the tuberculate ribs are coarsen and alternatedwith one or two nodeless finer ribs. This character reminds us of the ornament of Nostoeeras
Hyatt, 1894. The available evidence, however,is by no means sufficient to connect Axonoeeras
phylogenetically with Nostoeeras.
Anaklinoeeras Stephenson, 1941 (type species
A. reflexum Stephenson, 1941, from the Neylandville Marl of Texas) and Planostoceras
Lewy, 1967 (type species P. rehavami Lewy,
1967, from the Upper Campanian of Israel)are evidently offshoots of N ostoeeras, taking
the posture of body-chamber in a plane, as
Pravitoeeras Yabe, 1902 (type species P. sigmoidale Yabe, 1902 from the Upper Campanianof Japan) is a probable offshoot of Didymoeeras
taking a plane shell-form in the main growthstages (see Matsumoto et al., 1980; Morozumi,1985). Some authors affiliate Anaklinoeeras
and Planostoeeras to Nostoeeras as subgenera,
but the changes in shell-form seem to haveoccurred abruptly in these three examples.
To sum up, the tuberculate heteromorphs
whose main or adult shells posture nearly orstrictly in one plane are probably polyphyletic.
Neocrioeeras and certain other possibly related
genera with four rows of tubercles and theirpresumed descendants with two rows of tubercles could be grouped into a subfamily, forwhich Spath (1953) suggested the Neocrioceratinae. I would evaluate this suggestion, if theintimate relationships among these genera weremade clear with sufficient evidence. With regard

472

Tatsuro MATSUMOTO et ai.

to many of them, however, the phylogenetic
origin is uncertain in our present knowledge.
Acknowledgements
We wish to thank Professors Itaru Hayami,
Tamio Kotaka and Kametoshi Kanmera and
Dr. Ikuwo Obta for their offer of facilities to
one of us (T.M.) for the study of the specimens
kept at UMUT, IGPS, GK and NSM respectively,
and also Dr. Kenshiro Ogasawara and Mr. Haruyoshi Maeda for their kind help at IGPS and
UMUT. We are also indebted to Dr. Masayuki
Noda for his kindness in taking photographs,
to the authorities of the Japan Academy for
permission to reproduce the photographs of
Jimbo's type specimens printed in p. 57 (figs.
1-2) of the Proceedings of the Japan Academy,
vol. 61, ser. B, no. 2 (1985), and to Miss Yoshimi
Tanigawa in preparing the typescript.
References Cited
Adkins, W. S. (1929): Some Upper Cretaceous
Taylor ammonites from Texas. Univ. Texas
Bull., 2901, p. 203-242, pis. 5, 6.
Collignon, M. (1969): Atlas des Fossiles Carac·
teristiques de Madagascar (Ammonites), fasc.
15 (Campanien infedeur), p. 1-216, pis.
514-606, Servo Geol., Tananarive.
Diener, C. (1925): Ammonoidea neocretacea.
Fossilium Catalogus, 1 Animalia, part 29,
p. 1-244, Junk, Berlin.
Gill, J. R. and Cobban, W. A. (1973): Stratigraphy and geologic history of the Montana
Group and equivalent rocks, Montana,
Wyoming, and North and South Dakota.

U.S. Geol. Surv. Prof. Paper 776, p. 1-37.
Henderson, R. A. (1970): Ammonoidea from theMata Series (Santonian-Maastrichtian) of
New Zealand. Special Papers in Palaeont.,
no. 6, p. 1-82, pis. 1-15.
Hyatt, A. (1894): Phylogeny of an acquired
characteristic. Proc. Amer. Phil. Soc.,
Philadelphia, vol. 32, p. 349-647, pis. 114.
--(1900): Cephalopoda. In Zittel, K. A. von.
1896-1900: Textbook of Palaeontology,translated by Eastman, C. R., p. 502-604,
London.
Immel, H., Klinger, H. C. and Wiedmann, J.
(1982): Die Cephalopoden des Untereren
Santon der Gosau von Brandenberg/Tirol,
Osterreich. Zitteliana, vol. 8, p. 3-32,
pls.1-11.
Jimbo, K. (1894): Beitrage zur Kenntnis derFauna der Kreideformation von Hokkaido.
Paliiont. Abh., N. F., vol. 2, p. 149-194[3-48], pis. 17-25 [1-9].
Klinger, H. C. (1976): Cretaceous heteromorphammonites from Zululand. Dept. Min. Geol.
Surv. Mem., no. 69, 142 p., 43 pis., Pretoria.
Lewy, Z. (1967): Some late Campanian nostoceratid ammonites from southern Israel.
Israel Jour. Earth-Sci., vol. 16, p. 165-173.
--(1969): Late Campanian heteromorph ammonites from southern Israel. Ibid., vol. 18,
p.109-135.
Matsumoto, T. (1942): Fundamentals in the
Cretacenous stratigraphy of Japan. Part 1.
Mem. Fac. Sci. Kyushu Imp. Univ., ser. D,
vol. 1, p. 129-210, pis. 5-20.
--(ed.) (1963): A Survey of Fossils from Japan
Illustrated in Classical Monographs, 57 p.,
68 pis., Palaeont. Soc. Japan, Tokyo.
--(1959): Upper Cretaceous ammonites ofExplanation of Plate 93
Neocrioceras spinigerum (Jimbo)
Figs. 1, 2. Lectotype, UMUT. MM7521, from Ikandai, Urakawa. Lateral (1) and ventral (2)views, xl. 5.
Figs. 3-5. Paralectotype, UMUT. MM7522, from Ikandai, Urakawa. Lateral (3) and ventral viewsof pathologic body-chamber (4) and normal phragmocone (5), x 1.6.
Figs. 6-8. MC. Hbo-3695, from Haboro. Lateral view (6) with Sphenoceramus nagaoi (Matsumoto et Ueda) and an ammonite, ventral views of a portion of phragmocone (7) and bodychamber (8), xl.
Photos by courtesy of Mr. H. Maeda (1-5) and Dr. M. Noda (6-8).

MA TSUMOTO et al: Neoc rioceras spinigerum (Jimbo)

7

Plate 93

820. Neocrioceras spinigerum (Jimbo)
California. Part 2. Mem. Fac. Sci. Kyushu
Univ., ser. D, Spec. vol. 1, p. 1-172, pIs.
1-41.
--(1977): Some heteromorph ammonites fromthe Cretaceous of Hokkaido. Ibid., ser. D,
vol. 23, no. 3, p. 303-366, pIs. 43-61.
--(1984a): Some ammonites from the Cam·panian (Upper Cretaceous) of northern
Hokkaidq. Part 1. Ammonites from theUpper Campanian of the Teshio Mountains.
Palaeont. Soc. Japan, Spec. Pap., no. 27, p.
5-32, pIs. 1-9.
--(1984b): An aberrant ammonite genus fromthe Cretaceous of Hokkaido. Proc. Japan
Acad., vol. 60, ser. B, p. 341-344.
- - (1985): Restudy of Crioceras spinigerum
Jimbo, a Cretaceous ammonite species. Ibid.,
vol. 61, ser. B, p. 56-59.
- - and Miyauchi, T. (1984): Some Campanianammonites from the Soya area. In Matsu·
moto, T. (1984a): Some ammonites fromthe Campanian (Upper Cretaceous) ofnorthern Hokkaido. Part 2. Palaeont. Soc.
Japan, Spec. Pap., no. 27, p. 33-76, pIs.
10-31.
--and Morozumi, Y. (1980): Late Cretaceousammonites from the Izumi Mountains,
Southwest Japan. Bull. Osaka Mus. Nat.
Rist. no. 33, p. 1-31, pIs. 1-16.
- - , - - , Bando, Y., Hashimoto, H. andMatsuoka, A. (1980): Note on Pravitoceras
sigmoidale Yabe (Cretaceous heteromorphammonite). Trans. Proc. Palaeont. Soc.
Japan, N.S., no. 123, p. 168-178, pIs. 2226.
--and Obata, 1. (1981): A new heteromorphammonite from Hokkaido in the collectionof Yoshitaro Kawashita. Bull. Natn. Sci.
Mus., Tokyo, ser. C, vol. 7, no. 3, p. 115118, pIs. 1, 2.
Morozumi, Y. (1985): Late Cretaceous (Cam·panian and Maastrichtian) ammonites from
Awaji Island, Southwest Japan. Bull. Osaka
Mus. Nat. Rist., no. 39, p. 1-58, pIs. 1-18.
Schliiter, C. (1871-76): Cephalopoden deroberen deutschen Kreide. Palaeontographica,
vol. 21, p. 21-24, pIs. 1-18 (1871); p. 25120, pIs. 19--35 (1872); vol. 24, p. 121263, pIs. 36-55 (1876).
Scott, G. R. and Cobban, W. A. (1965): Geologic
and biostratigraphic map of the Pierre Shale473

between Jarre Creek and Loveland, Colorado.
U.S. Geol. Surv. Miscell. Geol. Invest. Map,
1-439,1 map with text p. 1-4.
Shimizu, S. (1933): Note on two interesting
Senonian ammonites from Hokkaido andSouth Saghalin. Jour. Shanghai Sci. Inst.,
sec. 2, vol. 1, p. 11-15, pI. 2.
Simionescu, 1. (1899): Fauna cretacica superiora
de la Urmos (Transilvania). Acad. Romana,
vol. 4, p. 237-274 [1-38], pIs. 1-3.
Spath, L. F. (1921): On the Upper Cretaceous
Ammonoidea from Pondoland. Ann. DurbanMus., vol. 3, p. 39--57, pIs. 6-7.
--(1953): The Upper Cretaceous cephalopod
fauna from Graham Land. Falkland lsi.
Depend. Surv. Sci. Rep., no. 3, p. 1-60,
pIs. 1-13.
Stephenson, L, W. (1941): The larger invertebrate fossils of the Navarro Group of Texas.
Univ. Texas Pub., no. 4101, p. 1-641(incl. pIs. 1-95), folded tables 1-6.
Tanabe, K., Obata, 1. and Futakami, M. (1981):
Early shell morphology in some Upper
Cretaceous heteromorph ammonites. Trans.
Proc. Palaeont. Soc. Japan, N.S., no. 124,
p. 215-234, pIs. 35-38.
Ward, P. D. and Mallory, V. S. (1977): Taxonomy
and evolution of the lytoceratid genus
Pseudoxybeloceras and relationship to thegenus Solenoceras. Jour. Paleont., vol. 51,
p. 606-618 (incl. 3 pIs.)
Whitfield, R. P. (1880): Paleontology of theBlack Hills of Dakota. In Newton, H. andJenney, W. P.: Report on the geology andresources of the Black Hills of Dakota.
U.S. Geog. Geol. Surv. Rocky Mtn. Region,p. 325-468, 16 pIs. (inaccessible in Japan).
Wiedmann, J. (1962): Ammoniten aus der
vascogotischen Kreide (Nordspanien). 1.
Phylloceratina,
Lytoceratina. Palaeonto·
graphica, vol. 118, A, p. 119--237, pIs.
8-14.
Wright, C. W. (1957): In Moore, R. C. (ed.):Treatise on Invertebrate Paleontology. Part
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l-L 490, Geol. Soc. Amer. & Univ. Kansas
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Tatsuro MATSUMOTO et al.

474

Wright, C. W. and Matsumoto, T. (1954): Some
doubtful Cretaceous ammonite genera fromJapan and Saghalien. Mem. Fac. Sci. Kyushu

Univ., ser. D, vol. 4, no. 2, p. 107-134,
pIs. 7-8.

Abesbinai 'it.:fZ:J*J=7"'c/71,
Haboro +Jrm,
Hobetsu 1l1l71I],
lkandai ;tf~hlkushum·bets ~~:tf71I].
Kawakami JIII..
Kikume-zawa 1M lHi iR.
Kotanbetsu [KotambetsJ 1~"fl71lJ.
Mikasa =~, Naibuchi r"JiJ:iI. Nakagawa 9")11. Namikawa ~JII. Obira 'J'f. Onkono-sawa::z / iR, Saku &7-.., Saku-gakko-no-sawa itI:7-..$t:cO)iR.
Sanushibe -if":>( c/ "'. Sosushi '/;<. c/oTeshio ::Rlll. Urakawa iJlif<iJ.

* /'13ill!*c~~~i;~7 /,-1':-71 r 0) 1 fill Neocrioceras spinigerum (JIMBO) I:''':)I,'L: Neocrioceras ml,O)ti4:a::fifl"c';!; Q N. spinigerum (JI\\BO) O)i"J,;JRil;.iI G < IIJ!!f(, '2!'-:h L 1" t.n ' lW.{til;;!;~kO)~ • • :a:~*K~~. ~O).m~.il-GMm'2!'-:hk~*~t.MG, ~&O)~m~~i~ U': o $5'\-0) ~~rT*il;~O) iR71IH:.i~~ Q C GL 1" Q ,~, (5'HfliJ 0) ~ O)~~il;MlHiil-x~!iI'iCyIJil-)Ii, ill\lftq:.J: I), it.:liiJ-il!\l{-4:J*JO)!EftJl:IH:'J: 1)Il!:~il;;!;Q;: c~J!:{*(191:'~U,:o it.:lj\(;i f
WjO)tl:J.1H *flii:~JV10)r& C rnlt!(H:. ~ ~ -t '7 ;<.~O)-t ~ i O);!; QflHi G-tt Iv~ ~~WML 1" Q*;: C ~ ~ Gil-I:. U': o la-iIii'i U~Qi&.ttEITc£jHe:;Mi0 <C, *:f!THiiJlil6J~/LI:.$ (:to t Ie: -if" /' I· =. 7/') I:.i& G. Sphenoceramus naumanni (Yokoyama) ~ S. nagaoi (Matsumoto et Ueda)~I-'¥?;: ct);~I,'ofIt* Neocrioceras ~1:'A:hG:hL\,'t.:[!tW.%~O)iJj\f&cO)tt"t~~tA.;r,...

*& c il-tJ: I) "'tJ: Q {, 0) Ii ftl!o)~f:'f$ U.:1Jil; J:\, -;: c ~~i1Ui!tt U': ot~::4':~J'l~· N*tt7-../!$ '1(1;j~~~· UIT
fJ~

Neocrioceras ~f9TJiI,: ;:O)~O)%m~;~cL~. )IITLIJ:;tJ'l~

ftll.~cO)mlJ;J.,s~UIH~<.~~I}fj~G.

fit*0)~J5L U':Jjf!;~~tlt~1j • ~.iI U': o;!;Q ~I:' ,,:)1,'LIHfl1LO)m~~t%O)i.iJfjgtt~~~ U': oU- G*tfCl¥Jmf$~ ~ Gil-I:' -t Q 1:'li~m-t Q iJj\f;j\JH:. ":)\, 'L O)~JiI,I:'/Gviilil;;!; Q ;: !: ~1I1ji/ii G,~~~.gI¥JM.~~<O)~~G~~~&*~M

Explanation of Plate 94

Neocrioceras spinigerum (Jimbo)
Figs. 1-2. TTC. 400819, from Ikandai, Urakawa. Lateral (1) and ventral (2) views of an immature shell, x 1.5.
Figs. 3-5. GK. H5205, S. Nagaoka's ColI. from South Sakhalin. Ventral (3), lateral (4) andfron tal (5) views, x 1.
Figs. 6-9. TTC. 360806, from the Kikume-zawa, Mikasa. Two lateral (6, 8) and ventral (7, 9). views, x 1. 2.
Figs. 10-13. TTC. 360815, from Nio-no-sawa, Toyosato. Two lateral (10, 12) and ventral (11,13) views, xL
Figs. 14-16. TTC. 370817, from the Saku-gakko-no-sawa, Saku. Lateral (14) and ventral views ofphragmocone (15) and a portion of body-chamber (16).
Photos by courtesy of Dr. M. Noda.

MATSUMOTO et al: Neocrioceras spinigerum (Jimbo)

1

4

3

8

15

Plate 94

5

9

475

PROCEEDINGS OF THE PALAEONTOLOGICALSOCIETY OF JAPAN

a*i5~4~¥~~ 135 @]19~~:;lj103~)o

@

A

~

;1l

(iifiif1TtrJO 0) Squalodon 'IJI1:'-::J
I,'L .............................. 1l7;l~l!.f~&
Recognition of the genus Enterodon from
Iwaki Formation of Joban Coalfield andobliteration of "Anthracothema tsuchiyai"(nom. nud.) ................ , ... Y. Tomida
tl*:1 l/:!l '/ 3 /' '1'0) "):n\.!lfJ~\J3nlllrpH" 11i'U{',~1tEI:'-::J\"L .................... lI1li;j,;~*On the genus Palaeoloxodon from the East
China Sea ........................ H. Otsuka
1'l1j)j iH..f (,/$l;/iR i:%) 0) Yl Jf; !E. fR .......... '5ll:.i ~ ~~1
iH..f~fl']f3!E. 0) f'Utl.l , Gj'iiJI.I;:hI.l, Z, 1.1' ? .......... .
....................... , Ironmn#E' fillfill£fr:b;~tJL1Hp~J\l'iI:r77~.0)*~~~:1-¥f~ ff~0)3-00~W~~0)....... ' ...................... f!~:fr ~ffr~;

1lt~1jl7HJc'IT;;fMff)0)""lv·qc~9iiJEffl7 ot -T,'/ of- ? ;< :J11 f]~ (Trochidae) O)lJli:n<.n::.
-::J\"L ........................... . :;!CJ1iRlifitr.l£ft~~~7/'~~1bO)~~~~O)W~ .. iIIrl* ~A record of Menabites (Ammonoidea) fromthe Cretaceous of Hokkaido ............... .
· .. .. .. ... T. Matsumoto and T. Takahashi
Little known ammonite Grandidiericeras
from the Cretaceous of Hokkaido ......... .
· ............... T. Matsumoto and R. Saito
Note on an ammonite species of Pachydiscus
from Awajj Island, Southwest Japan ..... .
T. Matsumoto, Y. Morozumi and T. Ozawan~JEm.=.

E *i11:!ffrr¥:~m 1351"1WIJ~I.I;1986ff 6 Ji 14' 15
E ~tJLHlnl:il: L3 ~;ttlf.¥/.JMi7:l-*-e~~llt ~htc. ($JJnt <0) r~-~J:.¥.!rJIl!..f!lli'!'~:,®,:;m ......... , .. III iRMj1:IiHtEr::"ic~J ~ htc.IL\~t1:4~0) JJJi:*~cf,R t bJi::R:I:'~

I:.{f? 1JifjiJt;(t"t;;0)~1t ......... , ... ,1H/r{~'~;
E *0) Nipponitrigonia kikuchiana I::' -::J\"f!!lm................................ tlHl~'T·.~ • • D·~M7~~7/'0)iH..f1:=ttYl~E........................ rnf\;.iIz· i'lIi~ 1m/Il:i..!I1frrmljj[~llfO)J:jUf(lJj'!.tt ...... EB{I::.iIZ •L~~.~·~H X'~EB~ft'MEBma
Restudy on "Inoceramus yubarensis NAGAO

et MATSUMOTO" from the Upper Cretaceousof Hokkaido ...................... M. Noda
p~1tEilJ~llfl:'-::J\"L ......... , .... §~lX5:jy /v ~ l/ /,y(/{of-~ ~/')1:.:t,;vtZ,~/vA$C-lj-/' :1"1tE ..... , ....... , .. iII~t.j-{'f- • 1JI1IW IDP.
~,1iMrjHi'i~EIK:E l

Fusulinella. FusulinaI)

1tE[il)~llf~O)feJ~I:'-::J\"L............... .
........................ riIIljJJ!t· f!i:1Il~~:r~.W.It!l~~mlK~~~z,&/.~Mm~MO)~

1!i!E.:¥fl:: ...... ji'fIUl!#lJ1 • :E111~ff~91!. ~lHjlll.7..~"fmlELTITI*7-,\,- ~O)ji(lIx£t\~11~ti:l'lr,~xHt ....
................. , ..... , .......... Il7;JH J'~
i: mlfti1%f19\:-"fml'I' %Jrl'flO:.:t-ovt z, I¥jQfj~HrfL!'u 1t
E;lIH:.-::J\"L ...................... illi 'iL,Mir.!
fi~A.~~Ji!(~m~M~~O)~1:~R!E.Wm(~m) .. , ........ , ... " ............. , ·f:i\~til~ILl p l)jl.~iY'J;~liy:ilftf;H·iffif'0 (3)..... ·I)'lIWil*:kAA
III ml it.lt!l J:Jf'il 0) ;:fk!Jly !(E'i ;fl3.~l J!i1l!!! fu nt ........... .

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

*HjfflJl·*{ERT·~1Jl(~

YHtf"fmlS!lli*l I) i1f!±l Ltc. 3 ft]0)l!!!f'01tE ....

· .. .. .. .. .. .. .. .. . . .. . .. *tt8~1j)j· :;!c{EJ3.';'fcp. ~tmlJL1'IH&WJ%f~{1::0)1tEifHfullf ~O) 3 )(W.lfutdZ , .. , .......... " .. *~~/l£k' !5~Ij)jT~:,...;f- ~ rJ.4 ltl.:s1:iEi{:Effli~Hi 1!R;f,C{!::1:i:fJ
tlt~,5A : tliPlBjI;y-' :k1B..iIJ11~. /' HtE'ltH:. -::J\, 'L~tiH..f.ilfi:l1t~-¥,,*,~rlrj.;J.It!lJi!(O)i'fJil/lllJi'l 0) Yl~Htfi W~(~0)1) ........................ ~*1j)j&

fl'fL!E.:t-o Hh /;T ~ 7 " n~=txYl ~ 7J" ,/;j; ~ TO)j@1.rG}f;n.~...... f~UiJ~~ 'l'icJllif1::rt.... illilBJ3.';m· .7..IImtFJ'IllilJEfUtEllff::'-::Jl'L ............... ·jljPH3/Y::. 11' .'E.1tE llfl:' -::Jl' L ...... m:J::i~1( • f~HllBiJL"fml!!O)Jttfi'lIDI:'-::J\"L '" .:kIB.iI.ilfi:· ***,-.................................. iill7K~1v'::1tE!i'H:'-::JI,'L

m~~~Yl~ill • • ~O)~~.~~.MK~~<~

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r~*tilZ 'l'icJllft\:;jft476
New members approved by the Council Meeting held on June 13,1986
Yen-Nien, Cheng; Satoshi, Chiba; Chikara, Hiramatsu; Toshiaki, Irizuki; Megumi, Kobayashi;
Fumio, Kubota; Hiroshi, Kurita; Takashi, Murota; Jun, Nakano; Yukihiro, Numabe; Yoshitaka, Ohta; Ikuo, Sunouchi; Kiyoshi, Takatsuka; Yuichiro, Tanaka; Yasuhiro, Taniuchi;
Yutaka, Toda; Akira, Tsukagoshi; Michio, Yoshida.
Members resignatedJuzo, Arai (Fellow); Yoshio, Hojo; Tadashi, Horiuchi.

Bibliography U)flj:(j(;:?\. \-C

B *~1:.!J&.J~~"('f:!:, 1981-1985%0) 5 ~r,~I':::'~~ ~.hf-:NiftrxO) Bibliography >£ffifJIH.:::. J: IJ flJrri" 7.l .:c re_ts: IJ i Lt..:o .: 0) Bibliography t':::'11V't*JlH IJ J-1J1:!J&.J~~~~O)lVf7C~iftrX>£'I:J,L.'I':::', II ;{:O)'tiH>£;iS"J
iJ' ",:,f..:;jl'~ .wO)~x~ i3"&I)7.l .: eeL i i"
xiiikO)~l(ji;11~@IAO) $ttr~£~ c L -C frl ,t..: l' c ftl,l' i i"0)"(', J:,1~ 5 -:f7J'0)~iftr:)( ~ 7, ~ YfO) (lIH.:::.ftt-:> -c f'Fl.ix L, 1986~12~ 3113 i -r::t-oiZS IJ T ~ l '00

W AT ANABE Kozo (1973) Pro!usulinella assemblage in the Omi Limestone, Niigata Prefecture,
central Japan (Studies of Carboniferous fusulinacean of Omi, part 1). Transactions andProceedings of the Palaeontological Society of Japan, New Series, no. 95, pp. 371-394, Plates
51-53, Figure 1, Tables 1-11.
Fusuline
Carboniferous[,*~cp O)*Jt~XO){7~]BANDO Yuji and KATTO Jiro (1980) On the Upper Triassic ammonoids from the Sampozan
Group at Hitsuzan, Kochi City in Shikoku. In A. Taira and M. Tashiro (eds.): Selected
papers in honor of Prof. Jiro Katto. Geology and Paleontology of the Shimanto Belt. Rinyakosaikai Press, Kochi, pp. 95-100. (j','Ij~-(Jm*WO):=::<iJllJlf,fJ: I)~!±l Lt..::=::llHi:l&J!/lO) 7 :/-'C-:T1
H.:::. "Jl ,-C) (J. E.)
Triassic
Ammonoidea[!jtfJ*'pO)*x~*9ff ~ 0) 5fIlX iiiftrx 0) fJlj]

HANZA W A Shoshiro (1961): Cretaceous and Tertiary three-layered larger Foraminifera andtheir allied forms: their classification and geographical and stratigraphical distributions.
Fossils (Palaeontological Society of Japan), no. 2, pp. 1-24, Figures 1-29, Tables 1-2. U:f(H EI~e·~:=::e~*~~R~&~~O)~~.O)~.~b~K~~~~·~~~~~~)

Cretaceous to Tertiary~)

Larger Foraminifera[,*~cpO)*XJH~ff ~5fIlX~iftrXO){7Ij]

KOBA YASHI [wao and KAKIZAKI Takeo (1978): [Preservation of veretebrate bones in nodules
from the Tsurushi Formation] Saito Ryojiro Sensei Taishoku Kinenshi, pp. 43-49, Plates 1-2.
(i!lT~~/;;:L -JH.:::.'e2jjj1~.ht..:tj·1t:OO)f*ff-) (J.)

MioceneMammalia[t.6Ux~<p0)~x1HgO) t.r.l '5fIlX~filjXO) fJlj]

i'± 1. aiftrxI1~~0)~f(;II~I':::'.?I17·L-CT~l'0 ~0)7,.?I1 J!.-11ftt*O)i=I~,JH.:::.V'tl'ii"0
2. '*ltt.:g,111tl'm~ Lt.r.l '"('ill '-C <t!. ~ l '0
3. ~xcp"(';iS"JiJ' -:> t..:I:I'1~O)~mln • HH~11 Index ~*.I@ L, ~,f,1~ A L -C <t!. ~ l
4. ~.h:f.hO)w:r'~!IlH·, ;jf~ ~ U/'H.:::.mi1lA) O)lVf'9l:~t.:::. J: 7.l13 *O)iff:jSf~ ;is''JiJ' -:> t..:~ffiJX>£ t-3j\\0~ 0)~frI1:t-o~ b-l± <t!. ~ l '0
'0

477

Hrilix ~ :A I- O)jzsN:'t'T980 {Iiltnnl'f #;111

JldtA:':r:Jlj!"f:ffllJt1l1!{~f'r~/J:.47/r'1:r~-:r;( I}~

1:1 :4':J'i~I'-:'1&.}"¥=0)(mk 0 nmtls~.l't 0::(jilIi.fl'mr:~ • R1~~

TAXA INDEXANIMALIA
GeneralProblematica, Trace fossils
and Others
CHORDATA
Mammalia
Proboscidea
Artiodactyla
Aves
Reptilia
Amphibia
PiscesOsteichtyes
Chondrichtyes
PROTO CHORDATA
Graptolithia
Conodontochordata
ECHINODERMATA
Holothuroidea
Echinoidea
Crinoidea
BRACHIOPODA
BRYOZOAARTHROZOA
Insecta
Crustacea
Malacostraca
Cirripedia
Ostracoda
Branchiopoda
Trilobita
ANNELIDA
MOLLUSCA
Cephalopoda
Coleoidea
Ammonoidea
Nautiloidea
Mollusca excl. Cephalopoda
Bivalvia
Scaphopoda
Gastropoda
Polyplacophora and Monoplacophora

COELENTERATA
Anthozoa
Zoantharia
Tabulata
Scleractina
Tetracorallia
Octocorallia
Stromatoporata
Hydrozoa
Scyphozoa
POLYFERAPROTOZOA Excl. MASTIGOPHORA
Radiolaria
Rhizopoda
Foraminifera
Fusuline
Larger Foraminifera
Benthic Smaller Foraminifera
Planktonic Foraminifera
VEGET ABILIA
GeneralProblematica and Others
Palynology
ANGIOSPERME
Dicotyledoneae
GYMNOSPERME
Coniferopsida
Ginkgopsida
Cycadopsida
Pteridospermopsida
PTERIDOPHYT A
Pteropsida
Articulatae
NON TRACHEOPHYTA
Charophyta
Calcareous Algae
Coccolithophoridae
Dinoflagellata
Diatomae
Silicoflagellata
Others478
GEOLOGIC AGE INDEXGENERAL OR INDEPENDENT TO AGE
PRE-CAMBRIAN
PALEOZOICOLD PALEOZOICCAMBRIANCAMBRIAN TO ORDOVICIANORDOVICIANSILURIAN
SILURIAN TO DEVONIANNEW PALEOZOICDEVONIAN
CARBONIFEROUS
CARBONIFEROUS TO PERMIANPERMIANPALEOZOIC TO MESOZOICPERMIAN TO TRIASSICMESOZOICTRIASSICJURASSICJURASSIC TO CRETACEOUSCRETACEOUS
EARLY CRETACEOUSLATE CRETACEOUSMESOZOIC TO CENOZOICLATE CRETACEOUS TO TERTIARYCENOZOICTERTIARY
PALEOGENE
PALEOCENE
PALEOCENE TO EOCENE
EOCENE
EOCENE TO OLIGOCENEOLIGOCENEOLIGOCENE TO MIOCENE
NEOGENE
MIOCENE
MIOCENE TO PLIOCENE
PLIOCENE
PLIOCENE TO PLEISTOCENE
NEOGENE TO QUATERNARYQUATERNARYPLEISTOCENE
PLEISTOCENE TO HOLOCENEHOLOCENE

~~

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Transactions and Proceedings of the PalaeontologicalSociety of Japan
September 30, 1986

New Series No. 143

CONTENTS
TRANSACTIONS
816. MATSUMOTO, Tatsuro and NODA Masayuki: Some inoceramids (Bivalvia) from the
Cenomanian (Cretaceous) of Japan-I. New or little known four species from Hokkaido and Kyushu . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . 409
817. GOTO, Hiroya, MARUOKA, Kunitaka and ISHD, Ken-Ichi: Lepidolina columbiana(Permian fusulinid) from British Columbia, Canada ...........•.....•....• 422
818. NISHIDA, Harufumi: On a new Tempskya from Japan .....••.....•........ 435
819. KOBAYASHI, Teiichi and HAMADA, Takashi: The second addition to the Silurian
trilobite fauna of Yokokura-yama, Shikoku, Japan . . . • . . . . . . . . . . . . . . . • . . . 447
820. MATSUMOTO, Tatsuro, MURAMOTO, Kikuo, TAKAHASHI, Takemi, YAMASHITA,
Minoru and KAWASHITA, Yoshitaro: On Neocrioceras spinigerum (Jimbo), a speciesof Cretaceous heteromorph ammonids (Studies of the Cretaceous ammonites from
Hokkaido-LV) . . . . • . . • . . . . . . . . . . . . . . . . • . . • . . . . . . . . . . . . . . . . . . . 463
PROCEEDINGS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . 475