ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No.

164

Palaeontological Society of Japan
December 30, 1991

Co-Editors

Tsunemasa Saito and Kei Mori

Language Editor

Martin Janal (New York)

Editorial Board
Kunihiro Ishizaki (Tohoku University), Yukio Isozaki (Yamaguchi University), Tomoki Kase
Itaru Koizumi (Hokkaido University), Yasumochi Matoba (Akita
University), Kei Mori (Tohoku University), Hiroshi Noda (University ofTsukuba), Tsunemasa Saito(Tohoku University), Y okichi Takayanagi (lshinomaki Senshu University), Kazushige Tanabe(University of Tokyo), Yukimitsu Tomida (National Science Museum), Kazuhiko Uemura (National(National Science Museum),

Science Museum)

Officers for 1991-1992
President: Kiyotaka Chinzei
Honorary President: Teiichi Kobayashi
Councillors: Kiyotaka Chinzei, Takashi Hamada, Yoshikazu Hasegawa, Itaru Hayami,
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Ikuo Obata, Kenshiro Ogasawara, Tomoo Ozawa, Tsunemasa Saito, Yokichi
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Members of Standing Committee: Hiromichi Hirano (General Affairs), Tomoki Kase
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Auditor: Kuniteru Matsumaru
The fossil on the cover is Trilophodon sendaicus Matsumoto, an extinct elephant, which
was described from the Pliocene Tatsunokuchi Formation developed in the vicinity of
Sendai, Northeast Honshu, Japan. (IGPS coil. cat no. 87759 (A), length about 18.5 cm)
All communication relating to this journal should be addressed to the

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, pp. 883-909,20 Figs., December 30, 1991

926.

ON THE EVOLUTIONARY CLASSIFICATION OF THE FAMILYLEPIDOCYCLINIDAE (FORAMINIFERIDA)*
KUNlTERU MATSUMARU
Department of Geology, Faculty of Education,
Saitama University, Urawa, 338

Abstract. The nepionic types of the microspheric juvenarium of almost all genera of
the family Lepidocyclinidae are described, giving the framework of the evolution andclassification of the family Lepidocyclinidae. From the viewpoint of the nepionic types, thedevelopment of periembryonic chambers of megalospheric forms, and the chronologicaloccurrences of various taxa included in the family Lepidocyclinidae, all the lepidocyclinids
are grouped into three lineages. As such, the family Lepidocyclinidae is thought to beheterogeneous in origin. The family Lepidocyclinidae comprises three subfamilies: Lepidocyclininae, Helicolepidininae, and Eulepidininae, the last one a new subfamily; theseare briefly described.
Key words. Uniserial nepiont, reduced uniserial nepiont, asymmetric biserial nepiont,
symmetric biserial nepiont, Lepidocyclinidae, Eulepidininae, n. subfamily.

lensis intergraded with E. semiradiata Barker
and Grimsdale, which had numerous equatorial chambers in cyclical arrangement. The
latter gave rise to Polylepidina chiapasensis(Vaughan), with one or two principal auxiliary chambers. Moreover, Helicostegina
gyralis Barker and Grimsdale was anqther
ancestor to either Middle to Upper Eocene
Helicostegina polygyralis (Barker), which had
a longer and tightly coiled uniserial portion
and a thickened spiral wall, or Upper Eocene
to lowest Oligocene Helicostegina paucispira(Barker and Grimsdale), which had arcuatechambers in cyclical arrangement. According to Cole (1960), Helicostegina polygyralis(Barker) is a senior synonym of Helicostegina
soldadensis Grimsdale, 1941 (= HelicosteginoMes Caudri, 1975), but the present author
does not accept Cole's views regarding thesynonymy of Helicostegina polygyralis, andrecognizes both Helicostegina polygyralis(Figure 8-1) and H. soldadensis (Figure 101-3). Helicostegina polygyralis (Barker),comprising various forms with different-

Introduction

The ongm and evolution of the family
Lepidocyclinidae Scheffen, 1932 has beenreported on by Barker and Grimsdale (1936),
Tan (l936b), Rutten (1941), Hanzawa(1964,1965), Matsumaru (l97Ia, b), Frostand Langenheim (1974), Sirotti (1982),
Butterlin (1977, 1987), and others. The fundamental route of the evolution of lineages
within the family Lepidocyclinidae has been
described by Barker and Grimsdale (op. cit.).
Their thesis has been developed by many
authors as follows: Lower Eocene Helicostegina gyralis Barker and Grimsdale, which
evolved from Upper Paleocene Eoconuloides
lopeztrigoi (Palmer), equipped with counter
septa, then evolved into Middle Eocene
Eulinderina guayabalensis (Nuttall), which
had only a single whorl of nepionic chambers
followed by arcuate equatorial chambers incyclical arrangement. Eulinderina guayaba"Received December 4, 1990; revised manuscript
accepted September 19, 1991
883

884

Kuniteru Matsumaru

length spiral chambers, evolved into Helicolepidina spiralis (Tobler), with one or two
principal auxiliary chambers and radialstolons.

After the publication of Barker and Grimsdale (1936), Rutten (1941) thought that
Pseudolepidina Barker and Grimsdale, 1937
could have arisen from Helicostegina Barker

1c

Y=7
X=14

Z=15

2b

Figure 1. A drawing showing the ontogenetic change from spiral to concentric growth of microspheric
Lepidocyclina (Eulepidina) papulifera Douville from Rembang, Java, Indonesia (Tan Sin Hok, 1934, fig. 4; laof Fig. I). The numbering of spiral to concentric nepionic and postnepionic chambers, including the firstchamber, proloculus (I b of Figure I), and the schematic diagram of chamber budding formation of microspheric L. (Eulepidina) papulijera follows Tan's idea (I b-c of Figure I). Chambers of the first spiral and several
concentric cycles are easily shown by single lines. The lower three are an alternative interpretation by theauthor (2a L c of Figure I). In 2b, the concentric growth is shown at the 14th budding stage. The fundamental
chamber budding formation is relevantly shown at the lower right (2c of Figure I). In order to explain thegrowth pattern, it is useful to distinguish some parameters. The number of spiral chambers with only one
chamber forming the aperture is designated by Y (Y = 7 in 2a-c of Figure I). The total number of initial
spirally coiled nepionic chambers, excluding the symmetric chamber, is designated by X (X = 14 in 2a-c ofFigure I). There is the closing chamber or symmetrical chamber in the place where the initial spire meets thesecond spire. The position of the closing chamber is shown as Z (Z= 15 in 2a-c of Figure I).

926.

Classification of the Family Lepidocyclinidae

and Grimsdale, 1936, while Actinosiphon
Vaughan, 1929, Triplalepidina Vaughan andCole, 1938, LepidocyC/ina (LepidocyC/ina)
GUmbel, 1870, and L. (Pliolepidina) Douville,
1911 could have evolved from L. (Polylepidina) Vaughan, 1924. Frost and Langenheim (1974) considered that Lepidocyclina(Polylepidina) was an ancestor to Pseudolepidina, LepidocyC/ina (Lepidocyclina), L.
(Nephrolepidina) Douville, 1911, and L.
(Eulepidina) Douville, 1911. According toSirotti (1982), LepidocyC/ina mantelli (Morton) and Eulepidina ocalana (Cushman),both of the subfamily Lepidocyclininae
Scheffen, 1932, were considered to have derived from Upper Eocene Lepidocyc/ina guber-885

nacula Cole, remarkably developing radial
equatorial stolons lying on different planes.
The latter evolved from LepidocyC/ina proteiformis Vaughan. Also, it is noted that Lepidocyclina protei/ormis derived from Polylepidina chiapasensis Vaughan, which
evolved from Eulinderina semiradiata Barker
and Grimsdale. Sirotti (op. cit.) also
thought that Nephrolepidina ecuadorensis(Hofker) and Helicolepidina spiralis (Tobler),both of the subfamily Helicolepidininae Tan,
1936, evolved from Helicostegina paucispira
Barker and Grimsdale, acquiring radial equatorial stolons lying on the same plane. Thisis based on Hofker's hypothesis (1968) of
Nephrolepidina springing from Helicostegina.

Lineage 1

Y

=7

=9

y
Polylepidina
zeilmansi

180)r /" \
l.trr.C'

1"'.J~

QJ
C
QJ
U

110

,J,

v

Y=8

1 X.16.1S

J hl1

17~

o

'G

';«;",,1 '1'\I('(=10

Y=11)

y

=12

Polylepidina
chiapasensis

I\1\

Type C

11,,'1 Y.9

~

':!(
.

16"1

/"

Eulinderina guayabarensisw

l1"'k~10OG \y,8( ( f"') I) X:15.1:l
~l.16

QJ

0.
0.

T

ype C

:::J

Helicolepidina spiralisoPseudolepidina trimera
QJ

"0
"0

L

'1

......./"lJ-a.'4t

13f'1 .... 11~\I~~O>.. ~13 Y,7
'\ ~,

18~

Type B

QJ

C
QJ

u

o
w

HelicosteQina Qyralis

Type A

~

QJ

~

o

...J

Figure 2. Schematic diagram of chamber budding formation of microspheric forms of Helicostegina
gyralis Barker and Grimsdale (topotype, Sabaneta, Veracruz, Mexico, BM (NH) P39150), Pseudolepidina
trimera Barker and Grimsdale (Adams, 1987, pI. 3, fig. 14), Helicolepidina spiralis (Tobler) (EI Alto, NW
Peru, BM (NH) P30248), Eulinderina guayabalensis (Nuttall) (Barker and Grimsdale, 1936, pI. 34, fig. II),
Polylepidina zeilmansi (Tan Sin Hok) (Tan Sin Hok, 1936a, pI. I, fig. 8), and Polylepidina chiapasensis(Vaughan) (1. Butterlin's 1182 specimen, Corinto no. I well, 930-935(8), Mexico). These forms belong tolineage I of the Lepidocyclinidae.

886

Kuniteru Matsumaru

Recently, Butterlin (1987) stated that Lepidocyclina (Polylepidina) was an ancestor to
all the subgenera Lepidocyclina, Eulepidina,and Nephrolepidina. In the recent classification of the family Lepidocyclinidae, Adams(1987) removed the Paleocene genera
Actinosiphon Vaughan, 1929 and Orbitosiphon Rao, 1940 from the family Lepidocyclinidae, placing the first genus in a new
family, the Actinosiphonidae, while the otherwas included in the family Orbitoididae
Schwager, 1876. The Middle Eocene Pseudolepidina was referred to the subfamily
Helicolepidininae. However, in Loeblichand Tappan (1987), Actinosiphon and Orbitosiphon, as a synonym of the former, werereferred to the subfamily Lepidorbitoidinae
Vaughan, 1933, while Pseudolepidina wasreferred to the subfamily Lepidocyclininae.
Thus these differing opinions have caused
considerable chaos around the question of theorigin, evolution, and classification of thefamily Lepidocyclinidae. The author has
had an alternative opinion on these problems
and gives his considerations in this paper.
Before proceeding to describe the family
Lepidocyclinidae, some problems should be
mentioned.
We should expect more abundant fossils of
all the taxa of the family Lepidocyclinidae
even though many authors have reported
them. Unfortunately, we have never seen
microspheric forms of Actinosiphon and Orbi-

tosiphon, and no one has observed the upper
Maastrichtian and Danian orbitoids thatmust straddle the Cretaceous/Tertiary boundary. Such gaps have to be bridged largely by
speculation, and paleontological or biological interpretation is necessary. In particular,
it is important to recognize the embryonicand nepionic (or periembryonic) types representing the megalospheric and microspheric forms of all the taxa of the family Lepidocyclinidae, and Cretaceous and Paleogene
orbitoidal Foraminifera.
As far as we know to date, we have fortunately discovered the following Upper
Cretaceous and Tertiary orbitoidal Foraminifera in the Indo-Pacific region (= IndoPacific of Figure 5): Lepidorbitoides bisambergensis (Jaeger), L. minor (Schlumberger),
L. socialis (Leymerie), Siderolites? sp., Orbitocyclina sp., Orbitocyclinoides schencki
Bronnimann, Pseudorbitoides sp., Sulcorbitoides sp., Sulcoperculina inaequalis Ho, S.
cubensis Palmer, S. vermunti Thiadens,
Actinosiphon tibetica (Douville), Orbitoclypeus suruaensis (van der Vlerk), O. molluccan a (Henrici), O. kimurai Matsumaru,
Polylepidina zeilmansi (Tan), P. birmanica
Rao, P. antillea (Cushman), and Helicostegina cf. paucispira (Barker and Grimsdale)(Douville, 1916; Tan, 1936a; van der Vlerk,
1923; Davies and Pinfold, 1937; Henrici,
1934; Rao, 1942; Smout and Haque, 1956;
Ho Yen et al., 1976; Hashimoto et al., 1978,_
Figure 3. Schematic diagram of chamber budding formation of microspheric form of Orbitocyclina
minima (Douville) (Gorsel, 1978, fig. 18), Lepidocyclina ocalana Cushman (Cole, 1941, pI. 13, fig. 6),
Lepidocyclina proteiformis (Vaughan) (Tantoyuca, Veracruz, Mexico, BM (NH) P32-630), Lepidocyclina
tobleri (Douville) (A. Sirotti's 63/3/8 specimen, Vistabella-Point, Trinidad), Lepidocyclina pustulosa
Douville (Butterlin, 1971, pI. 2, fig. 8), Lepidocyclina mantelli (Morton) (Marianna Limestone, Little Stave
Creek, Alabama, BM (NH) P47331), Lepidocyclina canellei Lemoine and Douville (left: A. Silvestri, ANFR
74-111 specimen, Cuba-Martinico; right: Cole, 1961, pI. 37, fig. 5), Nephrolepidina japonica (Yabe) (left toright: Topotype, Abuta Limestone (Matsumaru, 1969), Japan, slide 34-16; Ibid., slide 27; Kamiyokoze
Formation, Une, Japan, slide 13; Saginosu Formation, Kuroya, Japan, slide I ; Saginosu Formation, Kimo,
Japan, slide 32), Nephrolepidina orientalis (van der Vlerk) (Sarawak specimen, SK-2065), Nephrolepidina
luxulians Tobler (Tabanio no. I well, 2,600-2,610 feet, Java sea) and Nephrolepidina rutteni (van der Vlerk)(Shimoshiroiwa Formation, Japan, SH-3, slide 5). The terms pliolepidine, isolepidine, nephrolepidine,
trybliolepidine and multi lepidine in parentheses have been properly understood as the important charactersof the megalospheric nucleoconch of each species, and they have been of primary importance in generic orsubgeneric classification. These forms belong to lineage 2 of the Lepidocyclinidae.

926.

Classification oj the Family Lepidocyclinidae

887

endemic taxa, although no Eocene Lepidocyclina, Pseudolepidina, or Helicolepidina haveas yet been discovered in this region.
Almost all of the forms referred to thefamily Lepidocyclinidae have an equatorialchamber layer and a lateral chamber layer
superposing both above and below the for-

Premoli Silva and Brusa, 1981; Hashimotoand Matsumaru, 1984; Matsumaru andKimura, 1989). Therefore, these foraminiferal genera or species from the CaribbeanGulf coast of Mexico and Europe-Mediterranean regions (= America-Europe of Figure5) existed in the Indo-Pacific region as
Lineage 2

Y=6

Y=5

Y=4

Y

=7

Y =8

Nephroleidina
luxulians(multilepidine)

Q)

c:

Q)

C~~O

.Q
~

...
0.
0.

\...J

Type D

2:13

1

12

japonica ,~(nePhro~to trybryolepidine)

Nephrolepidina

::J

....o

~\(trYbrYOlePidine)

X_II
2:12

V·'/.·

1 2 @ Y:6
10 V
X:l0~, Z:11Q)

Nephrolepidina rutteni'~OAr~""0~~~2
C9~P0Y~7
U

Nephrolepidina orientalis(t rybryolepidi ne)otffG"o,
)Y-6
10
X;,1

\1.. . . ."2='2\....'2')(i;O/,11,\
9"~9 Y:7 10-9 ... 10I
G~
X=11 ~C) Y,8
10..
)02:12
X,13

9

,~

8~)~8

Iv) Y=6
\'
X:l1
"-,2

,J

2=12.......,2'"

4

2=14

Type D

14

Lepidocyclina canallei---(jsolepidine)
Q)

c:

6

I

"-,0

Q)

o
o
Cl

---

~

o

~...

~

'::J

) ~~~

.ft,
::::J7

I

Y-S

lLO

X;9.8
2=10

Z=10

Type D

Lepidocyclina
mantelli
7 (isolepidine)

o
Q)

?G7~6

~6Ce)6 ~~~9

2=9--..,.--;-:---(0:-8\,.9;.s--S.,.J

l.eRidocyclina Lepidocyclina.
,
ocafan.a"l'~I"o:\" protelformls~8[1'1{..f)~(L~'0l Gj
9~y:s.\~ ~~~

Z=9~3

Type D

Lepidocyclina
pustul<;>sa tpl;>leri Lepidocyclina pustulosa(pl/olepldJne)
10(isolepidine)r;.C~9G)~~161 C ) ~~l2

I'" "'
9.... 9

(
\..'2

9

Z.'2

13

Type D

2=13

~~~~

Ul

::J

oQ)
o

Orbitocyclina minima

1''1'\

co
Q)

9~r'9

()

~'3 Z·,3

...

...

Q)

0.
0.

::J

Y:7

\ G 1 X='2.11

Type CKunileru Malsumaru

888

mer. The morphological characters of theembryonic apparatus and periembryonicspires accompanied by the development of aprincipal auxiliary chamber and adauxiliarychamber of the megalospheric form of thisfamily have been regarded as being of maintaxonomic importance. As such, the studyof the microspheric form of the family LepidocyC\inidae has made little progress since
Tan's research (1934-36, 1939).
The author studied the microspheric formof the family Lepidocyc\inidae to obtain
phylogenetic information about budding
chamber formations; this information isdrawn from the ontogenetic stages developedby the microspheric generation, because eachform appears to recapitulate some of theimportant events in its phylogenetic history.
Also, the author feels that the nepionic types
give the framework of a natural classification.
As a result of the present study, the origin of
the family LepidocyC\inidae is thought to beheterogeneous, and all of the lepidocyC\inids
are grouped into three lineages according to
the nepionic types of their microspheric form,
although the morphological features of theembryonic and periembryonic chambers ofthe megalospheric form have to be added.
The present paper was read October 2, 1990
at the Fourth International Symposium on
Benthic Foraminifera, BENTHOS '90, Sendai, Japan.

Lineage 3

...

ClJ

0.
0.
~ClJ
C

OclJ

..... u

"'0
clJOl

3·0-

-10

Eulepidina dilatata(8,\l \.) )X-9/

Eulepidina favosa(9,",7/t"'l~7 Y:05
\

Y =7

Y=6

Y=5

Z:10

10

8Y'C. . .' ~~~O

~

)

11

8

Z=ll

Eulepidina gigantea
,?9\
8-;'8

~u) ~~~,

___

Z:.12

12

Eu~e
idina omphalus
15

8~(r ~'0 Y:?

'0)_ _ _ '~'3

TypeD

X=le
Z·13
_

__

clJ

c
clJ
u

Orbitoclypeus nummuliticus;010,\o/ 8~--G~)9 ~:,7,

W

9

clJ

Type D

Z:12

12
(/J

:J

o
clJ
u

co
.....

...

clJ

o

clJ

Lepidorbitoides

min 0 r8.ff~~
Type C( r\2) Y-5
X~1O.9
11

Z.11

Y=7

Type D

X:.12
Z=13

0.
0.

:::::lFigure 4. Schematic diagram of chamber budding formation of microspheric forms of Lepidorbitoides
minor (Schlumberger) (left: MacGillavry, 1963, pI. 6, fig. 2; right: Gorsel, 1978, fig. 14 (c», Orbitoclypeus
nummuliticus (GUmbel) (Sirotti, 1987, pI. 2, fig. 3), Eulepidina dilatala (Michelotti) (A. Sirotti's specimen,
Sciacca, Sicily), Elilepidina Javosa (Cushman), 1919 (Huasteca Formation, Tamismolon, Veracruz, Mexico),
Eulepidina gigan/ea (Martin) (Tan Sin Hok, 1935a, pI. 3, fig. 4) and Elilepidina omphalus (Tan Sin Hok)
(Ibid, pI. 4, fig. 5). These forms belong to lineage 3 of the Lepidocyclinidae.

926.

Classification of the Family Lepidocyclinidae

The nepionic types of each form of thefamily Lepidocyclinidae andthree lineages
Tan (1934) interpreted the budding chamber formation of microspheric Eulepidina
paplilijera (Douville) as follows: as seen in
the upper row (la-c) of Figure I, two nepionic spires, starting from a retrovert aperture recognized in the first chamber (la ofFigure I ; 8th chamber, including proloculus,
recounted by the author, I b-c), which developed from a single coil of periembryonicchamber, meet each other from opposite
directions to make a symmetric chamber (6thchamber of Tan, la; 13th chamber of theauthor's recounting, I b-c). But Tan unfortunately presumed a wrong course for the successive budding spires that lead up to annularly disposed chambers (Figure I, la-c).
This made it difficult for later scientists toread and eval uate his study.
In contrast with Tan's interpretation, as
seen in the lower row (2a-c) of Figure I, theauthor found that the 14th chamber, counting
from the proloculus, gave rise to annular
growth. As a matter of fact, the 14th chamber is defined as the first ann ularly disposedchamber, and it is the frontal chamber of thefirst symmetrical chamber.
Generally speaking, the juvenarium or
nepionic chambers of the microspheric form
consist of a minute spherical proloculus and anumber of succeeding chambers spirally disposed. The number of spirally disposedchamber is designated as Y. They are surrounded by a set of primary and secondary
spires of nepionic chambers, and both spires
join each other to make a symmetrical chamber. X is the number of chambers developed
from a pair of primary and secondary spires,
respectively. Z is designated for the firstsymmetrical chamber counting from theproloculus. Then these nepionic chambers
are enclosed by the first ann ularly disposedchambers (14th chamber, 2a-c of Figure I).
In all of the forms in the family Lepidocy-889

clinidae and its relations, there are four
different nepionic types:
A: uniserial nepiont (or nepionic chambers); as seen in Figure 2, the postnepiont
buddings after the first whorl of nepioniccham bers are 0 bserved at least in the 13th
budding stage of Helicostegina gymlis Barker
and Grimsdale. The uniserial nepiont groupis found in Helicostegina. Spiral thickeningof the primary coil can be seen.
B: reduced uniserial nepiont; the first
whorl of nepionic spires became increasingly
shortened. Helicolepidina and Pseudolepidina belong to this group (Figure 2). Thickening of the spiral wall can be seen.
e: asymmetric biserial nepiont; a pair ofprimary and secondary spires are introduced
as a newly acquired character as shown byasymmetric spires of different length (Figures2-4). Orbitocyclina, Lepidorbitoides (part),
Elilinderina, Polylepidina, and Lepidocyclina(part) belong to this group.
0: symmetric biserial nepiont; both primary and secondary spires as shown by symmetric spires of equal length (Figures 3-4).
Lepidorbitoides, Lepidocyclina, Orbitoclypeus,and Eulepidina belong to this group.
Lineage I has an ancestor, Helicostegina
gym lis Barker and Grimsdale, possessing
uniserial nepiont (type A). Helicolepidina(type B nepiont), Pseudolepidina (type B nepiont), Elilinderina (type e nepiont), andPolylepidina (type e nepiont) are known inlineage I (Figures 2, 5). The nepionic chambers of microspheric Polylepidina chiapasensis (Vaughan), the type species of Polylepidina, is unfortunately incompletely known,so that no exact determination is possible.
Fortunately the author could regard the nepionic chambers of microspheric Polylepidina
zeilmansi (Tan) as the type C nepiont. The
four genera Helicolepidina, Pseudolepidina,
Eulinderina and Polylepidina are supported
by both the reduced number of uniserial
nepionic chambers and the development of
postnepionic spires (Figure 2). The microspheric form with an asymmetric biserial

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::::l
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Acti nosi phonsem mesl_-\Ajelicorbitoides-Lepidor bi oides
,?

o
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Orbitoclypeus

I

I

J \

'=.seu~rbi!OideS\~,E'phrOlepidina vaughan'osa

L.vurn [aun~nsis,\,N.tournoueri
Lineage 2\
L
d.au
It. 'n antE'r"1.\. L. :"mallei
L. mi raflorensis=EulE'Pidi na dil tata,E.ur dos~e~1 .
_' - _ __ _ _ __Lineage 3~L.ocalana-~iw=, Ohal'

;::.;
~,
~

....
:::::

~

I:l

t::;

Suleoperculina~ Sulcorbitoi des\ Pseudorbitoi des~ Orbitocycli Ina§_-S Eulepidina dilat~t~,~.:..fo~o~ te. Lineage--_f=~--?~ ~siphon

tibetica

T1r

F=- ~F=-mT,na,a l"" =__--=

3

Ne~phrolepid'na anoulosal~_____

Nm"''"'

liN. t 0 urn~uE'ri(:\j. ia no nical- Li nE'age 2

N.s matren is- -- - -I:l---- -------

Orbito lypeus

0...
I

. u

\'Sulcorbitoides- Lepidorbitoides
1- -?
U

Uo~~~J~lei

Lineage 1\. N.rut eni~N.isolep\dinOides - =-

?

- --

PolylE'pidina anfl lea

l~mansi, p.~a-~ica~O_regi n~audsPlra------

Lineigel-

Figure 5. Pedigree chart showing the tentative evolution of the family Lepidocyclinidae (in bold), and Cretaceous andPaleogene orbitoidal foraminiferal genera and species (ordinary lettering) related to the Lepidocyclinidae.

~

926.

Classification of the Family Lepidocyclinidae

891

nepiont (type C) originates from each form
either with an uniserial nepiont (type A) orwith a reduced uniserial nepiont (type B).
According to Barker and Grimsdale (1936),
Eulinderina guayabalensis (Nuttall) evolved
from Helicostegina gyralis Barker and Grimsdale. As such, the form with type C nepiont
originated from one with a type A nepiont.
Lineage 2 is most exhibited by the symmetric biserial nepiont (type D) of each microspheric form of both Lepidocyclina andNephrolepidina (Figures 3, 5). The micro-

spheric form of Upper Paleocene to upper
Lower Eocene Actinosiphon is obscure, butthat the microspheric Actinosiphon was asymmetric biserial nepiont can be inferredfrom the fact that the symmetric biserial
nepiont, which was recognized in each formof Middle Eocene Lepidocyclina pustulosa
Douville, L. tobleri (Douville), L. ocalana
Cushman, and L. pro teiform is Vaughan,
could have evolved from the asymmetricbiserial nepiont of Maastrichtian Orbitocyclina minima (Douville), on which Vaughan(1929a) established Orbitocyclina as a valid
genus, and that Actinosiphon existed in thegeologic time span between Lepidocyclina

Figure 6. Helicostegina gyralis Barker andGrimsdale. 1. Equatorial section of a microspheric
form (BM (NH) P39150) equipped with counter
septa, showing the uniserial nepiont, type A. 2-3.
Vertical sections of both microspheric and megalospheric forms (both of BM (NH) P33356). Scalebars= 100 microns.

Figure 7. Helicolepidina spiralis (Tobler). 13. Equatorial sections. 1. A microspheric form
(BM (NH) P302-48) with radial stolons, showing the
reduced uniserial nepiont, type B. 2-3. Megalospheric forms (2: BM (NH) P30247; 3: BM (NH)P30249). Arrow showing a radial stolon. Scalebars= 100 microns.

892

Kllniterll Matsllmarll

926.

Classification of the Family Lepidocyclinidae

893

and Orbitocyclina. Also, as stated later, thepresence of radial stolons, recognized in equatorial chambers of Orbitocyclina mlmma(Douville), Orbitocyclinoides schencki Bronnimann, Actinosiphon semmesi Vaughan, A.
tibetica (Douville), Lepidocyclina ariana
Cole and Ponton, and L. proteijormis(Vaughan), can represent a bridge in thephylogenetic development from Orbitocyclina

Figure 9. I. Lepidocyclina mantelli (Morton).
la-c. Equatorial sections of megalospheric and microspheric forms (la: BM (NH) P47328; Ib: BM(NH) P47326; Ie: BM (NH) P47330). la-b. Bothmegalospheric forms showing the development ofadauxiliary chambers. Ie. A microspheric formshowing the symmetric biserial nepiont, type D. Id.
Vertical section of a megalospheric form (BM (NH)P47324). 2. Lepidocyclina proteiformis Vaughan.
2a-b. Equatorial sections of both megalosphericand microspheric forms (BM (NH) Ex P32630). 2a.
A megalospheric form showing 4 nepionic spires ofperiembryonic chambers. 2b. A microspheric formshowing the symmetric biserial nepiont, type D. 2e.
Vertical section. BM (NH) Ex P32630. Scalebars= 100 microns.

3

Figure 10. 1-3. Helicostegina soldadensis
Grimsdale. 1,3. Equatorial sections of megalospheric forms (I: BM (NH) P38379; 3: BM (NH)P4915). 2. Vertical section of a megalospheric form
(BM (NH) P4916). 4. Helicostegina paucispira(Barker and Grimsdale). Equatorial section of amegalospheric form (BM (NH) P47311). Scalebars= 100 microns.

~ Figure 8. 1. Helicostegina polygyralis (Barker).
la-b. Eqatorial section of megalospheric forms(both of BM (NH) Ex P32627) equipped with counter septa. Arrows showing radial stolons. Ie. Verticalsection of amegalospheric form (BM (NH) Ex P32627). 2. Eulinderina sp. 2a. Equatorial section of amegalospheric form (BM (NH) P33325) equipped with counter septa. 2b. Vertical section of a megalospheric form (BM (NH) P33325), showing an equatorial chamber layer divided into some zones near peripheryof test. 3. Polylepidina zeilmansi (Tan). 3a-e. Equatorial sections of microspheric and megalospheric
forms (Tan, 1936, pI. 1, figs. 3b-e, 8). 3a. Asymmetric biserial nepiont, type C. 3b-e. Megalospheric
forms, showing 4 nepionic spires of periembryonic chambers. Scale bars= 100 microns.

894

Kuniteru Matsumaru

through Actinosiphon to Lepidocyclina.
From Middle Eocene to Upper Miocene
megalospheric lepidocyclinids in lineage 2,
the embryonic apparatus evolved from isolepidine through nephrolepidine to trybliolepidine. The pliolepidine and multi lepidine
embryonic apparatus can be regarded as apeculiar structure in which some series of isoto nephrolepidine and nephro- to trybliolepi dine embryonic apparatus fuse into a cohesive whorl.
Cole (1963) observed Lepidocyclina oca/ana Cushman, which posesses a wedge-likeequatorial plate or wedge-shaped compact
layers in the equatorial chamber layer. As
such, this species is closely related to Trip/a/epidina veracruziana Vaughan and Cole, from

Figure 11. 1-4. Pseudolepidina trimera Barker
and Grimsdale. 1-2. Equatorial sections ofmegalospheric forms (1: BM (NH) P33031; 2: BM (NH)P33033) showing radial stolons or stoloniferous foramina (arrow). 3-4. Vertical sections of megalospheric forms (3: BM (NH) P33037; 4: BM (NH)P33032). Scale bars= 100 microns.

the Upper Eocene beds of Mexico, the typespecies of Trip/a/epidina Vaughan and Cole,
1938. Recently, Trip/a/epidina was abandoned by Cole (1963) as a synonym of Lepidocyclina GUmbel, 1870, while the former
was distinguished by Hanzawa (1964) fromthe latter based on the addition of a new
structural constituent. Based on the present
view of the nepionic chambers of the microspheric form of Lepidocyclina oca/ana, thisFigure 12. 1. Actinosiphon semmesi Vaughan.
Equatorial section of a megalospheric form (BM(NH) P32633). Arrows showing apertural foramina
or radial stolons. 2-3. Actinosiphon tibetica(Douvilli!). 2a. Vertical section of a megalospheric
form (Lepidorbitoides tibetica Douvilli! of BM (NH)F949). 2b-c. Equatorial sections of megalospheric
forms (Lepidorbtoides tibetica of BM (NH) F944 andF949). Arrows showing radial stolons or stoloniferous foramina. 3a-b. Equatorial sections of megalospheric forms (Lepidocyclina (Polylepidina) punjabensis Davies of BM (NH) P34987). Arrows
showing radial stolons or stoloniferous foramina.
Scale bars = 100 microns.

926.

Classification oj the Fami/y Lepidocy clinidae

species posesses the same symmetric biserial
nepiont (type D) as Lepidocyclina spec ies andis regarded as the genus Lepidocyclina.
The nepionic type of lineage 3 developed
from both the asymmetric bi serial nepiont
with spiral thickening of the prim ary coi l,and the symmetric biserial nepiont without
spiral thickening recogni zed in the microspheric Lepidorbitoides minor (Schlumberger), to the symmetric biseri a l nepiont with out spiral thickening of micro spheric Orb itoclypeus Silvestri, 1907 . The la tter gave riseto the same nepiont as micro sp heric Eu/epidina Douville, 1911 (Figures 3, 5). The
common morphological features among Le-

895

pidorbitoides Si lvestri , 1907, Orbitoclypeus ,and Eu /epidina are their genera l shape andthe number and disposition of the lateralchambers. The main evo luti on of lineage 3
is not completely c lear, but the evolution
involves the development of adauxiliarychambers. We can see the development ofadauxi li ary chambers from Lepidorbiloides
minor (Schlumberger) to L. socia/is (Leymerie) , whereas the adauxiliary chambers ofOrbitoclypeus and Eu /epidina are numerou s
in the equatoria l plane.
The origin of lineage I starts from Upper
Paleocene Eoconu /oides, but that of lineage 2
and lin eage 3 seems probably to be Upper
Cretaceous " ROla/ia", jUdging from the fossi lrecords of Sidero/ites?, Su/copercu/ina, Su/corbitoides, a nd Pseudorbitoides in the I ndoPacific region. Gorsel (1978) has a lreadyinferred thi s from the fossi l occurrences in theAmerica- Europe region. Detailed consideration of thi s point is beyond the sco pe of thispaper (Figure 5).

Discussion of some megalospheric formsof Actinosiphon, Lepidocyclina,
Nephrolepidina, and Eulepidina

Figure 13. 1- 2. AClinosiphon semmesi Va ughan. I. Equatorial section of ho loty pe (USNM
321783). Arrows showing radial stolon s. 2. Vertical secti on of cotype (USNM321782). Sca le bars =100 microns.

AClinosiphon Vaughan , 1929 was describedwith the type species Actinosiphon semmesi
Vaughan from the Lower Eocene Chicontepec Form ation , Mexico. Its arcuate- to ogiva l- shaped equatorial chamber wall s carryradial stolons, which are characteristic of thi s
genus. A s stated earli er, Rutten (1941) supposed th at Actinosiphon evo lved from Po/y /epidina Vaughan , 1924 as it reduced the size ofthe nep ionic chambers and developed radia l
sto lons. De Cizancourt (1951) consideredthat Lepidorbitoides cf. p/anasi Rutten , described from the Paleocene San Juan Limestone, Venez uela , by Caudri ( 1944) , a nd
Po/y/epidina barbadensis (Vaughan), described from Pa leocene blocks derived fromloes River mudflows, Barbados, by Vaughan(1945) , are synonymous; she placed thespecies in Actinosiphon. Grimsdale, in van

896

Kuniteru Matsumaru

Raadshooven (1951, p. 488), considered "Lepidocyclina" barbadensis Vaughan, describedfrom the Paleocene Guasare Formation, Venezuela, by van Raadshooven, and Actinosiphon semmesi Vaughan as the same species.
Also, Rutten, in van Raadshooven (op. cit.),suggested that if the genera Lepidorbitoides
and Orbitocyclina were included as subgenerain the genus Lepidocyclina, there could not bemuch objection to also including the genus
Actinosiphon as a subgenus. Rao (1940,1944) treated L. (Polylepidina) punjabensis
Davies, 1937 and Lepidorbitoides tibetica
Douville, 1916, from Kampa Dzong, Tibet, as
the same species, and he established the newgeneric name Orbitosiphon; Smout andHaque (1956) also abandoned L. (Polylepidina) punjabensis as a synonym of Lepidorbitoides tibetica, but thought tibetica belonged in Actinosiphon.
The author saw the holotype (USNM
321783) and topotype (BM (NH) P51967) of
Actinosiphon semmesi Vaughan, and the
topotype (BM (NH) P51969, P34987) andother specimens (BM (NH) P34987, F467) of
LepidocyC/ina (Polylepidina) punjabensis
Davies, from the Paleocene Ranikot Beds of
the Punjab Salt Range, Pakistan. Although
Adams (1987) differentiated Orbitosiphon
praepunjabensis from L. (P.) punjabensis(BM (NH) P51969), radial stolons were visible in the equatorial chambers in all of theforms listed above (Figures 12, 13). The
author regards L. (P.) punjabensis Davies andOrbitosiphon praepunjabensis Adams, 1987 as
Actinosiphon tibetica (Douville), because of
the existence of radial stolons in the equatorial chambers.
Moreover, the author studied Lepidocyclina
ariana Cole and Ponton, from the Middle
Eocene Punta Mosquito Formation, Venezuela, in which radial stolons were found(Figures IS, 16). As such, the author consid-

ered that Upper Paleocene to Lower Eocene
Actinosiphon semmesi Vaughan could have
evolved into Middle Eocene Lepidocyclina
ariana Cole and Ponton. Also, Vaughan(1936) has described upper Middle to Upper
Eocene Lepidocyclina proteiformis Vaughan
as having radial stolons.
Bronnimann (1944, p. 39) described Upper
Cretaceous Orbitocyclinoides schencki Bronnimann, from Kerman, Iran, which possessedradial stolons and more nepionic spires thanOrbitocyclina. As such, Bronnimann considered that there are close phylogenetic relationships between "Lepidocyclina" punjabensis(Davies) and Orbitocyclinoides schencki
Bronnimann. As stated before, the former
species is a junior synonym of Actinosiphon
tibetica (Douville) (Figure 12-3a - b). In
light of present knowledge, the author supposes that the phylogenetic relationships
among Orbitocyclina, including Orbitocyclinoides, Actinosiphon, and Lepidocyclina are
a recognition of an underlying community
during different geologic stages.
The next problem concerns the phylogenyof Eulepidina Douville, 1911. Hanzawa(1964, p. 308) considered that Eulepidina
might have derived from Nephrolepidina withthe remarkable development of the deuteroconch, whereas Adams (1987, p. 312)
thought that Lepidocyclina (Eulepidina)originated from Lepidocyclina s. str., which
possessed equatorial chambers arranged incircles, and that it could not have arisen fromany known species of Lepidocyclina (Neph rolepidin a ). Sirotti (1982) supposed that
Upper Eocene Eulepidina ocalana (Cushman) derived from Upper Eocene Lepidocyclina gubernacula Cole, whereas Butterlin(1987) considered that Lower Oligocene Lepidocyclina (Eulepidina) undosa Cushman
evolved from Lower Oligocene Lepidocyclina
chaperi Lemoine and Douville.

~ Figure 14. Lepidocyclina ariana Cole and Ponton. I. Equatorial section. 2-3. Equatorial views.
3. Arrow showing the position of radial stolons (Figure IS-la-c). 4. Vertical section. Middle to Upper
Eocene Punta Mosquito Formation, Venezuela. Scale bars= 100 microns.

926.

Classification of the Family Lepidocyclinidae

897

898

Kuniteru Matsumaru

The author wants to indicate that Eulepidina, possessing an increased number ofadauxiliary chambers, is essentially different
from both Lepidocyclina and Nephrolepidina,which possess from only a few to ten adauxiliary chambers, though some species have
twelve or rarely fourteen. In order toobserve the megalospheric nucleoconch, S.E.
M. pictures were taken from specimens of
Lepidocyclina ariana Cole and Ponton, from
the Middle to Upper Eocene Punta Mosquito
Formation, Venezuela, L. ocalana Cushman,from the Upper Eocene Ocala Limestone,
Florida, U.S.A., Nephrolepidina morgani(Lemoine and Douville), from the upper
Aquitanian to lower Burdigalian at SaintPaul-les-Dax, SW France, and Nephrolepidina
japonica (Yabe), from the Lower Miocene
Kamiyokoze Formation, Japan (Figures 14,
17 -19). The protoconch and deuteroconchof the megalospheric embryonic chambers ofthese Lepidocyclina and Nephrolepidina
species are pierced by numerous very small
perforations (1-3 microns) or several large
stoloniferous foramina (10-12 microns) thatconnect the adauxiliary and lateral chambers(Figures 17-19).
In Eulepidina dilatata (Michelotti), fromthe Aquitanian at Escornebeou, SW France,
the deuteroconch embracing the circumference of the protoconch is pierced by manylarge stoloniferous foramina (15-20 microns),
which are arranged in a few rows in theequatorial plane and which connect theadauxiliary chambers (Figure 20). This morphogenetic pattern, present in megalospheric
forms of Eulepidina dilatata (Michelotti), is
the same as the one well investigated in
Orbitoclypeus nummuliticus (Gi.imbel) by
Sirotti (1987). Accordingly, the author considers that an evolutionary path from
Paleocene-Eocene Orbitoclypeus to Oligocene-Lower Miocene Eulepidina exists (Fig-ure 5).

The classification of thefamily LepidocycIinidae
Both the nepionic types of the microsphericjuvenarium and the peculiar embryonic andperiembryonic chambers of megalospheric
forms characterize the following classificationof the family Lepidocyclinidae.

Generic distinction
Family Lepidocyclinidae Scheffen, 1932
Test discoidal to lenticular, with vacuolar
and rarely developed or well-developed lateral chambers and distinct equatorial layer ofchambers; chamber walls perforated bystolons and perforations. Paleocene toUpper Miocene.
Subfamily Lepidocyclininae
Scheffen, 1932
Test lenticular, with a distinct equatorialchamber layer equipped with lateral layers oneach side. Paleocene to Upper Miocene.

Lepidocyclina Gi.imbel, 1870
Type species. - Nummulites mantelli Morton, 1833
Bilocular megalospheric apparatus subequal or equal in size; a multilocular nucleoconch is present. Periembryonic chambers have more than quadriserial nepionicspires, and arcuate, ogival, rhombic,
spatulate, or hexagonal equatorial chambers
are arranged cyclically. Microspheric generation is exhibited by symmetric to rarely
asymmetric biserial nepionts. An equatorial~
Figure 15. 1-2. Lepidocyclina ariana Cole and Ponton. la-c. Equatorial views of a megalospheric
form (Figure 14-3) showing radial stolons (arrows) and diagonal stolons. 2. Equatorial views of amegalospheric form (Figure 14-2) showing a radial stolon (arrow). Scale bars= 100 microns.

926.

Classification of the Family Lepidocyclinidae

899

900

Kuniteru Matsumaru

chamber layer is rarely divided into somezones near the periphery of the test. Lateralchambers present.
Figure 3 (pars): Schematic diagram ofchamber budding formation of microspheric
forms of Lepidocyclina ocalana Cushman, L.
protei/ormis (Vaughan), L. tobleri (Douville),
L. pustulosa Douville, L. mantelli (Morton),
and L. canallei Lemoine and Douville.
Figure 9: Equatorial and vertical sections ofmegalospheric and microspheric forms of
Lepidocyclina mantelli (Morton) and L.
proteiformis Vaughan. Figures 14-17 :Equatorial and vertical sections, and equatorial views of megalospheric forms of Lepidocyclina ariana Cole and Ponton. Figure 18-1-2: Oblique views of megalospheric
forms of Lepidocyclina ocalana Cushman.

Actinosiphon Vaughan, 1929
Type species. - A ctinosiphon semmesi Vaughan, 1929b
Bilocular embryo, unequal or rarely equalin size; periembryonic chambers have 2 to 4
nepionic spires. Equatorial chambers arcuate, ogival, spatulate, or hexagonal, somewhat circularly arranged. Radial stolons inequatorial chambers are present. Lateralchambers are well developed.
Figure 12: Equatorial and vertical sectionsof megalospheric forms of Actinosiphon semmesi Vaughan and A. tibetica (Douville).
Figure 13: Equatorial and vertical sectionsof megalospheric forms of Actinosiphon semmesi Vaughan.
Subfamily Helicolepidininae Tan, 1936b

Nephrolepidina H. Douville, 1911
Type species. - Nummulites marginata Michel otti, 1841
Bilocular embryo, deuteroconch distinctly
larger than protoconch; a multilocular nucleoconch is present. Periembryonic chambers have 6 to 28 nepionic spires. Equatorialchambers are arcuate, ogival, rhombic,
spatulate, or hexagonal, arranged cyclically topolygonally, with the exception of a stellate
form. Microspheric juvenarium developed
into a symmetric biserial nepiont is present.
Lateral chambers present.
Figure 3 (pars): Schematic diagram ofchamber budding formation of microspheric
forms of Nephrolepidina japonica (Yabe), N.
orientalis (van der Vlerk), N. luxulians Tobler and N. rutteni (van der Vlerk). Figure
18-3: Oblique view of a megalospheric formof Nephrolepidina morgani (Lemoine andDouville). Figure 19: Equatorial andoblique views of megalospheric forms ofNephrolepidina japonica (Yabe).

Test discoidal to lenticular, with bilocular
embryo surrounded by thickened wall withvacuolar and rarely developed or well-developed lateral chambers, no or one adauxiliarychamber, one to four or six? periembryonicspires are present. Thickening of the spiralwall is present or absent. Radial stolons are
rarely present. Middle Eocene to Lower
Oligocene.

Helicostegina Barker andGrimsdale, 1936
Type species. - Helicostegina dimorpha
Barker and Grimsdale, 1936
Early chambers in an involute trochospiralcoil, later ones are subdivided ventrally or on
both sides to form arcuate chamberlets.
Adult stage consists of annularly disposedarcuate chamberlets. Microspheric uniserial
nepiont is present.
Figure 2 (pars): Schematic diagram ofchamber budding formation of a microspher-~ Figure 16. Lepidocyclina ariana Cole and Ponton. 1,3. Equatorial sections of a megalospheric
form (Figure 14-1) showing radial and diagonal stolons. 2. Equatorial view of a megalospheric form
(Figure 14-2) showing radial stolons (arrows). Scale bars= 100 microns.

926Classification of the Family Lepidocyclinidae

901

Kuniteru Matsumaru

902

ic form of Helicostegina gyralis Barker andGrimsdale. Figure6: Equatorial and vertical sections of megalospheric and microspheric forms of Helicostegina gyralis Barker andGrimsdale. Figure 8-la-c: Equatorial
and vertical sections of megalospheric forms
of Helicostegina polygyralis (Barker). Figure
10: Equatorial and vertical sections ofmegalospheric forms of Helicostegina soldadensis Grimsdale and H. paucispira(Barker and Grimsdale).

Helicolepidina Tobler, 1922
Type species. - Lepidocyclina (Helicolepidina) spiralis Tobler, 1922
Bilocular embryo followed by planispiralcoil of one or two whorls, termed the helicolepidine string. Microspheric reduced uniserial nepiont is present. Lateral chambers arewell developed.
Figure 2 (pars): Schematic diagram ofchamber budding formation of a microspheric form of Helicolepidina spiralis (Tobler).
Figure 7: Equatorial sections of megalospheric and microspheric forms of Helicolepidina spira lis (Tobler).
Eulinderina Barker andGrimsdale, 1936
Type species. - Planorbulina (Planorbulinella) guayabalensis Nuttall, 1930
Bilocular embryo followed by trochoid coilof one whorl, spire with thick outer wall
followed by arcuate chambers arranged
cyclically. An equatorial chamber layer israrely divided into some zones near the
periphery of the test. Microspheric asymmetric biserial nepiont is present. Lateralchambers vacuolar and rarely developed orwell developed.
Figure 2 (pars): Schematic diagram ofchamber budding formation of a microspheric form of Eulinderina guayabalensis(Nuttall). Figure 8-2a-b: Equatorial andvertical sections of megalospheric forms of
Eulinderina sp.

Polylepidina Vaughan, 1924
Type species. - Lepidocyclina (Polylepidina) chiapasensis Vaughan, 1924
Bilocular embryo, subequal in size: periembryonic chambers have 2 to 4 unequal
nepionic spires. Arcuate equatorial chambers are arranged cyclically. Microsphericasymmetric biserial nepiont is present. Lateral chambers present, but sometimes only a
few layers.
Figure 2 (pars): Schematic diagram ofchamber budding formation of microspheric
forms of Polylepidina zeilmansi (Tan Sin
Hok) and P. chiapasensis (Vaughan). Figure 8-3a- b: Equatorial sections of microspheric and megalospheric forms of Polylepidina zeilmansi (Tan Sin Hok).
Pseudolepidina Barker andGrimsdale, 1937
Type species. - Pseudolepidina trimera
Barker and Grimsdale, 1937
Bilocular embryo, subequal or equal insize; large third principal auxiliary chamberon the convex side of the equatorial plane ispresent. All periembryonic chambers have 4
unequal nepionic spires. Arcuate equatorialchambers are arranged cyclically. An equatorial chamber layer is divided into somezones near the periphery of test. Microspheric reduced uniserial nepiont is present.
Figure 2 (pars): Schematic diagram ofchamber budding formation of a microspheric form of Pseudolepidina trimera Barker andGrimsdale. Figure 11 : Equatorial and ver-~Figure 17. Lepidocyclina ariana Cole and Ponton. 1-2. Equatorial views of megalospheric forms(Figure 14-3) showing the structure of embryonic chambers, development of periembryonic chambers (4nepionic spires) and very small perforations (2-3 microns) of embryonic chambers. Scale bars = 100
microns.

926.

Classification of the Family Lepidocyclinidae

903

Kuniteru Matsumaru

904

tical sections of megalospheric forms of
Pseudolepidina trimera Barker and Grimsdale.
Eulepidininae Matsumaru, new subfamily
This new subfamily of the family Lepidocyclinidae is established to accomodate only thegroup of Eulepidina, which possesses alenticular test with a distinct equatorial chamber layer equipped with lateral chamber
layers at each side. Lower Oligocene to
Lower Miocene.

Eulepidina H. Douville, 1911
Type species. - Orbitoides dilatata Michelotti, 1861
Generic description: Large bilocularmegalospheric nucleoconch from 300 to 1,850
microns in diameter with cuboidal protoconch surrounded nearly or completely by
deuteroconch, both surrounded by a thickwall from 22 to 110 microns thick pierced bynumerous stolons that lead to many adauxiliary chambers that have more than 30 nepionic spires. Microspheric juvenarium
developed into a symmetric biserial nepiont ispresent. Equatorial chambers are arcuate,
ogival, spatulate, or hexagonal, arranged
cyclically to polygonally. Numerous lateralchambers are well differentiated from theequatorial layer. Lower Oligocene to Lower
Miocene.
Remarks. - Eulepidina Douville, 1911 isdistinguished from LepidocyC!ina GUmbel,
1870 and Nephl'olepidina Douville, 1911 by alarge embryonic apparatus, a thick-walled
megalospheric nucleoconch, and numerous
nepionic spires. Also the former can be
distinguished from Actinosiphon Vaughan,

1929 and all forms of the subfamily Helicolepidininae by numerous nepionic spires. A
serious look should be taken at thephylogenetic relationship between Orbitoc!ypeus and Eulepidina, because they bothpossess the same periembryonic nepionicspires.
Figure 1 : Equatorial section of a microspheric form of Eulepidina papulijel'a(Douville) and a drawing of the same showing the ontogenetic change of growth of
microspheric Eulepidina papulijera. Figure
4 (pars): Schematic diagram of chamber
budding formation of microspheric forms ofEulepidina dilatata (Michelotti), E. favosa(Cushman), E. gigantea (Martin) and E.
omphalus (Tan Sin Hok). Figure 20 :Oblique view of a megalospheric form ofEulepidina dilatata (Michelotti).
Acknowledgments
During the author's Foraminifera study inEurope, through financial assistance from the
Ministry of Education of Japan in 1985, andthe courtesies of JAPEX (Japan Petroleum
Exploration Co., Ltd.) and Modena University, Italy in 1989, the author gained much
helpful information, examined papers andobserved type and valuable specimens of theUpper Cretaceous and Tertiary orbitoidal
Foraminifera, and conducted discussions ofmany problems with European scientists.
The author was enabled to collect many fossil
specimens through the courtesy of a numberof scientists. As such, the author expresses
his cordial thanks to these organizations andacknowledges the help of the following individuals: Professors M. Neumann, J. Butterlin, A. Blondeau, Dr. A. Poignant (Universite
Pierre et Marie Curie (Paris VI)); Dr. M.- T.

~
Figure 18. 1-2. Lepidocyclina ocalana Cushman. Oblique views of embryonic and periembryonicchambers, very small perforations (2-3 microns) and stoloniferous foramina (10-12 microns) can be seen into
the embryonic chambers. 3. Nephrolepidina morgani (Lemoine and Douville). Oblique view of embryonic and periembryonic chambers of a megalospheric form (upper Aquitanian to lower Burdigalian beds at
Saint-Paul-Ies-Dax, France) showing stoloniferous foramina scattered into the embryonic chambers. Scalebars= 100 microns.

926.

Classification of the Family Lepidocyclinidae

905

906

Kunilel'U Malsumal'u

926.

Classification of the Family L epidocyclinidae

907

SAZU , Ljubli a na ).

References Cited
Ada ms, C. G. , 1987: On the classi fication of th e
Lep id ocyc li nidae (Foraminiferida) w ith redescriptio ns of th e unre lated Pa leocene genera
Actinosiphon a nd Orbitosiphon . Micropaleontology , vo l. 33, no . 4, p. 289- 317.
Barker, R.W. and Grimsdale, T.F ., 1936: A co ntribution to the ph y logeny of the orb ito id a l fora minifera , wi th descriptions o f new forms fromth e Eocene of Mexico. Jour. Paleonl. , vo l. 10,
no.4 , p.231 - 247.
- - , 1937: Studies of Mex ica n fossil foraminifera.

Anna/s and Magazine of Natural History , Ser.

Figure 20. Eulepidina dilatata (Michelotti).
Oblique view of a mega losp heric form (Aquitanianat Escornebeou , France) show in g th e embryon ic andper iembryon ic chambers a nd a few rows of many big
sto lo nifero us foramina ( 15- 20 mi c ro ns). Scaleb ar = 100 microns.

Venec-Peyre (Mu se um Nati o nal d ' Hi stoire
N at urell e, Pa ri s); Dr. D . Decro uez (Museumd ' Hi stoire Naturelle d e G e neve); Prof. H .
H ag n, Dr. K . K arl-Hein z (MUnch en University); Dr. F. Rag l (Natural Hi story Mu se um ,Vienna); Dr. e.G. Ad ams, Mr. R.L. H o d gkin so n (British Mu se um (N at ura l Hi sto ry)) ;Prof. A. Sirotti (Modena University); Prof.
F . Schiavinotto (Roma Unive rsity) ; Prof. R.
Ra ttge r (Kiel University) ; Dr. K . Dro bn e
(Paleontoloski In st itut Iva na Rakovca ZRC,

10, vo l. 19, p. 16 1- 178.
Bronnimann, P. , 1944: E ine neues Subgenus vonOrbilOcyc!ina aus Iran nebst Bemerkungen Uber
He/icolepidin a Tobler und verwa ndte F or men.
Schweiz. Paldonl. Abhandl. , vo l. 64, p.2 - 42.
Butterlin , J. , 197 1 : Con tribu ti o ns a la co nnai ssa nce
du Paleogene mar in du Nord-Ouest de la Co lo mbie, basee sur les Macroforaminiieres. Ec!og.
Geol. Helv., vo l. 64, no. I, p. 13- 27 .
- - , 1977: Problemes co ncernant la repartition
stratigraph ique et les relations phyletiques desmacrofo ra mini ieres du Pa leogene de la reg ion desCarai"bes. Memoria Segundo Congreso Latino

Americano de Geologia, Caracas,

Venezuela,

11 - 16, Noviemb. , 1973, Tomo 3, p. 1367 - 1379.
- - , 1987: Origine et evo luti on des Lepidocycl in es
de la region des Carai"bes. Co mp a ra iso ns etrelations avec les Lep id ocyc lin es des a utres
regions du monde. Rev. Micropa/., vo l. 29, no.
4, p. 203 - 2 19.
Caud ri , C. M.B. , 1944 : The la rger foraminifera of San
Ju an de los Morros, State ofG ua ri co , Venezuela.
Bul!. A mer. Po leon !. , vo l. 28, no. 11 4, p. 355 - 404.
- - , 1975: Geology and paleontology of Soldado
Rock , T rinidad (Wes, Indi es). Part 2: The larger foraminifera. Ec!og. Geo!. He/v., vo l. 68, no.
3, p. 533 - 589.
C iza nco urt, M. de, 1951: Grands foraminiieres d u
Pa leocene in ferie llr et de l' Eoce ne moyen duVenezuela. Mem. Soc. Geo!. France , vo l. 30,
no. 64, p. 1- 68.
Co le, W.S. , 1941: Stratigraphic and paleontologic~
Figure 19. Nephrolepidina japonica (Yabe). 1,3. Equatoria l views of megalospheric forms show in g
the embryo nic a nd periembryonic chambers, sto lo niferous foramina (arrow), a nd diagonal a nd an nul ar
sto lons of equa tor ia l chambers. 2a - b. Oblique views of megalospheric forms show in g sto lon iferous forami na (arrow) co nn ectin g between dellteroconch and adaux ili ary c ha mbers, a nd diagonal sto lo ns between
equator ia l a nd la tera l cham bers. The Lowe r Miocene Kam iyokoze Formation , Japan. Scale bars = 100
microns.

908

Kuniteru Matsumaru

studies of wells in Florida-no. 1. Bull. Florida
Geol. Surl'., no. 19, p. 1-91.
- - , 1960: Revision of Helicostegina, Helicolepidina
and Lepidocyclina (Polylepidina). Contr. Cushman Found. Foram. Res., vol. II, no. 2, p. 57-63.
- - , 1961 : An analysis of certain taxonomic problems in larger foraminifera. Bull. A mer.
Paleon!., vol. 43, no. 197, p. 373-407.
- - , 1963: Illustrations of conflicting interpretations of the biology and classification of certain
larger foraminifera. Ibid., vol. 46, no. 205, p. 164.
Davies, L.M. and Pinfold, E.S., 1937: The Eocene
beds of the Punjab Salt Range. Mem. Geol.
Surl'. India, Palaeon!. Indica, N.S., vol. 24, p. 179.
Douville, H., 1911: Les foraminileres dans Ie Tertiaire des Philippines. Philippine Jour. Sci., vol.
6, no. 2, p. 53-80.
- - , 1916: Le Cretace et l'Eocene du Tibet central.

Mem. Geol. Surl'. India, Palaeon!. Indica, N.S.
vol. 5, no. 3, p. 1-84.
Frost, S.H. and Langenheim, R.L., 1974: Tertiarylarger foraminifera and scleractinian corals from
Mexico. In, Cenozoic reef biofacies, p. 1-387.
Northern Illinois Univ. Press, Dekalb.
Gorsel., T.T. van, 1978: Late Cretaceous orbitoidal
foraminifera. In, Hedley, R.H. and Adams, C.
G. eds, Foraminifera 3, p.I-120. Academic
Press, London.
Grimsdale, T.F., 1941: Appendix: new species of
Helicoslegina from Soldado Rock. In, Vaughan, T.W. and Cole, W.S., Preliminary report on
the Cretaceous and Tertiary larger foraminifera
of Trinidad, British West Indies. Geol. Soc.
A mer. Spec. Paper 30, p. 1-137.
GUmbel, C.W., 1870: Beitrage zur Foraminiferenfauna der Nordalpinen Eocangebilde. Abhand.

Math. Physik. KI. Konig. Bayer. Akad. Wissensell., vol. 10, pt. 2, p. 581- 730.
Hanzawa, S., 1964: The phylomorphogeneses of the
Tertiary foraminiferal families Lepidocyclinidae
and Miogypsinidae. Sci. Rep., Tohoku Unil'.,
2nd Ser. (Geol.), vol. 35, no. 3, p. 295-313.
- - , 1965: The ontogeny and the evolution of larger foraminifera. Ibid., vol. 36, no. 2, p.239256.
Hashimoto, W., Matsumaru, K. and Kurihara, K.,
1978: Larger foraminifera from the Philippines
6. Larger foraminifera found from the Pinugay
Hill Limestone, Tanay, Rizal, Central Luzon.
Geol. Palaeont. Southeast Asia, vol. 19, p. 65-72.
- - , and Matsumaru, K., 1984: Mesozoic and Cenozoic larger foraminifera of the Philippines and areference to those found from Borneo by the
A PRSA's pa leontological recon na issance.
Ibid., vol. 25, p. 147-166.

Henrici, H., 1934: Foraminiferen aus dem Eozan und
Altmiozan von Timor. PalaeonlOgraphica, Supplement-Band 4, p. I-56.
Ho, Yen, Zhang Ping-kao, Hu Lan-ying and Shengling-chang, 1976: Mesozoic and Cenozoic foraminifera from the Mount lolmo Lungma region.

A report of scientific expedition in the Mount Jolmo Lungma region (1966-1968), Fasc. 2, p. 1-88.
Hofker, 1., 1968: Tertiary foraminifera of coastal
Ecuador. Palaeontographica, Band 130, Liefg.
1-6, p. I-59.
Loeblich, A.L. and Tappan, H., 1987: Foraminiferal
genera and their classification. p. 1-970,847 pis.
Van Nostrand Reinhold Co., New York.
MacGillavry, H.l., 1963: Phylomorphogenesis andevolutionary trends of Cretaceous orbitoidal foraminifera. In, von Koenigswald, G.H.R.,
Emeis, 1.D., Buning, W.L., and Wagner, C.W.
eds., Evolutionary trends in foraminifera, p. 139197. Elsevier Pub. Co., Amsterdam.
Matsumaru, K., 1967: Geology of the Tomioka area,
Gunma Prefecture, with a note on "Lepidocyc1ina", from the Abuta Limestone Member. Sci.
Rep., Tohoku Unil'., 2nd Ser. (Geo!.), vol. 39,
no. 2, p. 113-147.
- - , 1971a: Studies on the genus Nephrolepidina inlapan. Ibid., vol. 42, no. 2, p. 97-185.
- - , 1971 b: The genera Nephrolepidina and Eulepidina from New Zealand. Trans. Proc.
Palaeont. Soc. Japan, N.S., no. 84, p. 179-189.
- - , and Kimura, K., 1989: Larger foraminiferafrom the Eocene Shimizu and Miocene Misaki
Formations in Tosa Shimizu City, Kochi Prefecture, Shikoku, lapan. Ibid., no. 156, p. 255-269.
Michelotti, G., 1841: Saggio storico dei Rizopodi
caratteristici dei terreni sopracretacei. Mem.
Soc. Italian a, Sci. Res., vol. 22, p.253-302.
- - , 1861: Etudes sur Ie Miocene inferieur del'ltalie septentrionale. Natuurk. Verhandl. HoIland, Maatsch. Wetensch., vol. 2, pt. 15, p. 1-183.
Morton, S.J., 1833: Supplement to the "Synopsis of
the organic remains of the ferruginous sand formation of the United States", contained in Vols
17 and 18 of this journal. A mer. Jour. Sci. andArts., vol. 23, p. 288-294.
Premoli Silva, I. and Brusa, c., 1981 : Shallow-waterskeletal debris and larger foraminifers from DeepSea Drilling Project site 462, Nauru Basin, western equatorial Pacific. Initial Rep. Deep Sea
Drilling Project, vol. 61, p. 439-473. U.S. Govt.
Printing Office, Washington, D.C.
Raadshooven, B. van, 1951 : On some Paleocene andEocene larger foraminifera of Western Venezuela.
Proc., World Petrol. Congr., Leiden, Sect. I, p.
476-489.
Rao, S.R.N., 1940: On Orbitosiphon, a new genus of
Orbitoidal foraminifera from the Ranikot beds of926.

Classification of the Family Lepidocyclinidae

the Punjab Salt Range (N.W. India). Current
Sci., Bangalore, vol. 9, p.414-415.
- - , 1942: On Lepidocyclina (Polylepidina) birmanica, sp. nov., and Pselldophragmina (Asterophragmina) pagoda, subgen. nov. et sp. nov.,
from the Yaw Stage (Priabonian) of Birma.
Geol. Surv. India. Records, vol. 77, Prof. Paper
12, p. 1-16.
- - , 1944: A revision of some foraminifera described by Douville from the Kampa System ofTibet. Proc. Nat. Acad. Sci.• India. Allahabad,B, vol. 14, no. 3, p. 93-101.
Rutten, M.G., 1941 : A synopsis of the Orbitoididae.
Overdruk Geol. Mijnbollw, no. 2, p. 34-62.
Scheffen, W., 1932: Ostindische Lepidocyciinen I.
Wetensch. Meded. Dienst Mijnb. Ned. OostIndie, no. 21, p. 5-76.
Sirotti, A., 1982: Phylogenetic ciassification of Lepidocyciinidae: a proposal. Boll. Soc. Paleont.
Italiana, vol. 21, no. I, p.99-112.
- - , 1987: Ontogeny in Tethyan Discocyclinidae
and Orbitociypeidae (Foraminifera). Ibid., vol.
26, no. 3, p. 269-278.
Smout, A.H. and Haque, A.F.M., 1956: A note on
the larger foraminifera and Ostracoda of the
Ranikot from the Nammal Gorge, Salt Range,
Pakistan. Records Geol. Surv. Pakistan, vol. 8,
no. 2, p. 49-60.
Tan Sin Hok, 1934: Ober mikrosphare Lepidocyciinen von Ngampel (Rembang, Mitteljava).
Ing. Ned.-Indie; 4. Mijnbouw Geol., Band I, p.
203-211.
- - , 1935a: Ober Lepidocyclina gigantea K. Martin
von Stid-Priangan (Westjava), Tegal (Mitteljava)
und Benkoelen (Stidsumatra). Ibid., Band 2, p.
1-8.
- - , 1935b: Zwei neue mikrosphare Lepidocy-909

clinen von Java. Ibid., Band 2, p.9-18.
- - , 1936a: Lepidocyclina zeijlmansi nov. sp., eine
polylepidine Orbitoidide von Zentral-Borneo,
nebst Bemerkungen tiber die verschiedenen
Einteilungsweisen der Lepidocyciinen. Ibid.,
Band I, p. 7-14.
- - , 1936b: Zur Kenntnis der Lepidocyciiniden.
Natllurk. Tijdsch. Ned.-Indie·, vol. 96, no. 4, p.
235-280.
- - , 1939: On Polylepidina. Orbitocyclina and Lepidorbitoides. Ing. Ned.-Indie; 4. Mijnbollw.
Geol., Band 5, p.53-84.
Tobler, A., 1922: Helicolepidina, ein neues Subgenus
von Lepidocyclina. Eclog. Geol. Helv., vol. 17,
p.380-384.
Vaughan, T.W., 1924: American and European Tertiary larger foraminifera. Bull. Geol. Soc.
Amer., vol. 35, p.785-822.
- - , 1929a: Studies of Orb ito ida I foraminifera: thesubgenus Polylepidina of Lepidocyclina andOrbitocyclina, a new genus. Proc.• Nat. A cad.
Sci., vol. 15, no. 3, p. 288-295.
- - , 1929b: Actinosiphon semmesi, a new genus
and species of orbitoidal foraminifera, andPseudorbitoides trechmanni H. Douville. Jour.
Paleont., vol. 3, no. 2, p. 163-169.
- - , 1945: American Paleocene and Eocene larger
foraminifera. Geol. Soc. A mer.• Mem. 9, pt. I,
p.I-75.
- - , and Cole, W.S., 1938: Triplalepidina veracruziana, a new genus and species of orbitoidal
foraminifera from the Eocene of Mexico. Jour.
Paleont., vol. 12, no. 2, p. 167-169.
Vlerk, I.M. van der, 1923: Een Overgangsvorm tusschen Orthophragmina en Lepidocyclina uit het
Tertiair van Java. Verh. Geol. Mijnb. Gen. 7, p.
91-98.

Lepidocyclinidae f'j- (fl'fL.!R El )1l)J1tir.~i¥J5:HIH.:.~ ~ -C ; ~!E::ij\[,1l) Lepidocyciinidae W
c..."tn~.:.Jlf:~tJ::EnUc.:to J: aE~-=~c.1l) "Orbitoids" fl'fL.!RIl)~:BjCm!~*~,:,·':n'-c. ~UII
tdl1.~;fa-rrtJ:: "? td6*. 11!l~1l) f,!iJJtIl~~Il)!J l' 7"f1.. 4 m! (uniserial nepiont, reduced uniserial nepiont, asymmetric biserial nepiont :to J: a symmetric biserial nepiont) tJ, I? tJ:: .0 .:. c. tJ~
IjJ3 I? tJ,~.:. tJ:: "? t-:o ~~f1.. .:.
I? 1l)~5J1j c.lYi:BjCm!~* Il) 1ff~~ -'tr' f,!iJ1ff~~ 1l)geJi:to J: a
Lepidocyciinidae WIl) a; I? 11>.0 7t~~ c..:t 1l)~f\;i¥JtJ::gtt±l ~iHdi~;~.:.~~t-t.o c., \, 'i"~. Lepidocyciinidae Wll)li}j;f1., 3 7t~*kift (Lepidocyciininae !!IfW, Helicolepidininae!!lf
f'j-, Eu\epidininae!!lfW. tJTf~\) ~.:.FJf}j;~-C\".o,:, c.tJ~ljJ3l?tJ,~.:.tJ::"?t-:o it-:, ~~f1., 3 !!If
W~.:.FJf}j; -t.o ~}j;1l) 7t~a-~1i: ~ t-:o .:.
i -C- ~.:., Lepidocyciinidae W f1., E~-=ij\[, Il)
Eoconuloides }j;tJ, 1?ge1=.~, J1tir. ~ t-: c. ~ -C ~ t-:tJ~, *1i1f~~.:. J: "? -C ~filib-c 3 !!IfWf1.,
Eoconuloides }j;tJ, I?, a;.o \" f1. B !!If~c.1l) "Rotalia" }j;tJ, I? Maastrichtian t!!:1l) Orbitocyclina}j;a-~-C 83*~, i t-:I1'iIt!!:1l) Lepidorbitodies}j;, ~ 1?~.:.aE~-=~c.1l) Orbitoclypeus }j;",1l)J1tir. a-~-C 83* ~ t-: ~ Il) -C- a; "? -c, .:t I? f1.~11§.i¥J ~.:.v1.iinX;1l) ~ Il) -c' a; .0 .:. c. ~ IjJ3 I? tJ,~.:.

n

n

n
n

~ nt-:on101LOO~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, pp. 910-927,8 Figs., December 30, 1991

927.

ON SOME ACANTHOCERATID AMMONITES FROM
THE TURONIAN OF HOKKAIDO(STUDIES OF THE CRETACEOUS AMMONITESFROM HOKKAIDO-LXIX)T A TSURO MATSUMOTO
c/o Department of Earth & Planetary Sciences,
Kyushu University 33, Fukuoka 812

Abstract. New or revised information is given on the taxonomy and stratigraphic
occurrence of the following five species: (1) Pselldaspidoceras jiexlIoslim Powell, 1963,
which has recently been found at four localities in the lowest part of Member lIn, i.e. basal
Turonian, in the Oyubari area. One of the examined specimens was the holotype of
Pseudaspidoceras (Ampakabites) kawashitai Matsumoto and Obata, 1982, which now falls
into synonymy with Ps. jiexlIoslim. (2) Neomphaloceras pselidomphallim (Matsumoto,1975), whose revised diagnosis is given. It occurs in the lower part of the Middle Turonian.
(3) Neomphaloceras costatllm (Matsumoto and Kawashita, 1978), which was erroneously
assigned to Mammites or Morrowites. It occurs in the upper part of the Middle Turonian.
(4) Mammites afT. nodosoides (SchlUter), which is more evolute than typical M. nodosoides.
(5) Mammites n. sp. (?), which has peculiar bar-like ribs across the venter and spatulate
ventrolateral horns on the body chamber.
Key words.

Pselldaspidoceras, Neomphaloceras, Clinningtoniceras, Mammites, Morr-owites, Turonian.

the biostratigraphic papers (Tanabe et al.,
1975; Matsumoto et al., 1991). They are
stored in the following institutions, withabbreviations in parentheses:
Department of Earth & Planetary Sciences[formerly Department of Geology],
Kyushu University, Fukuoka (GK)Mikasa City Museum, Mikasa (MCM)
University Museum, University of Tokyo(UMUT)
Faculty of Education, Waseda University(WE)
Yokosuka City Museum, Yokosuka(YCM)
The terminology used in the description ofammonites is the same as that in my recent
papers (e.g. Matsumoto, 1988, p.4), which
followed essentially that in the Treatise (Arkell et al., 1957) with some modifications.

Introduction

While engaged in a biostratigraphic study
aimed at establishing an integrated scheme ofzonation by means of ammonites, inoceramids and foraminifera, I became aware ofsome erroneous points in the taxonomic andstratigraphic assignment of some ammonites
from the Turonian of Hokkaido. There
were also new finds of important species
which are useful for zonation and correlation.
In this paper I describe five selected Turonian
species, giving the revised or new informationand discussing their implications.
The investigated specimens came mostly
from the localities indicated in the maps of'Received January 26, 1991; accepted September 24,
1991

910

927.

Turonian acanthoceratid ammonites

Palaeon tological descriptions
Family Acanthoceratidae
de Grossouvre, 1894
Genus Pseudaspidoceras Hyatt, 1903

Type species.-Ammonites footeanus
Stoliczka, 1864.
Remarks.-See Kennedy et al. (1987, p.
34-41) for the generic characters and scope ofvariability. I now agree with them in regarding Ampakabites Collignon, 1965 as a synonym of Pseudaspidoceras. Cobban et al.
(1989, p.40) have also given the diagnosis.
Pseudaspidoceras flexuosum Powell
Figures 1-3

Pseudaspidooeras flexuosum Powell, 1963, p. 318, pI.
32, figs. 1,9, 10; text-figs. 2a-c, f, g.
Pseudaspidoceras (Ampakabites) kawashitai Matsumoto and Obata, 1982, p. 75, pI. 3, fig. 1 ;text-fig. 2.
Pseudaspidoceras flexuosum Powell; Kennedy etai., 1987, p. 34, pI. 2, figs. 1-4, 8-13, 16-17; textfigs. 3A-C, 5, 6C, D, 7A, C (with full synonymy).
Pseudaspidoceras flexuosum Powell; Cobban et ai.,
1989, p. 41, fig. 91L.

Material.-(1) MCM. A83-1 (plaster cast
GK. H9629) (Figure I) collected by Y. Kawashita (May 30, 1978) at loc. Y5599 on theright branch in the upper reaches of the
Kamimaki-zawa; (2) WE. A I lOY coil. by
H. Hirano at his loc. Y070077d (89), i.e. T.
M.'s loc. Y5232 (east end) on the right bankof the Hakkin-zawa; (3) MCM. A193-1(Figure 2) colI. by Y.K., T.M. & M. Noda
(Aug. 24,1983) at loco Y5165b; (4) GK.
H8367 (Figure 3) colI. by H.H., K. Tanabe,
M.N. & T.M. (Aug. I, 1975) at loc. Y6015c,
both on the right side of the Taki-no-sawafrom the lowest part of Member lIn (Matsumoto, 1942), Middle Yezo Group of the
Oyubari area.
Description. - The specimens (1) and (2)have the body chamber for about half a whorl
and are fairly large, viz. about 250 mm indiameter at the preserved end. The speci-

911

mens (3) and (4) are incomplete but, if restored, they would be very large, roughly
twice as large as the former two.
In these four specimens the whorl is moreor less higher than broad and subrectangularor subtrapezoid in section, with nearly flat,
subparallel flanks, abruptly bent umbilicaledge and vertical umbilical wall. The venter
looks more or less arched, but this may be due
partly to secondary compression and partly tothe original variability.
The ornament on the body chamber andthe preceding last part of the phragmocone iswell shown by the I st and 2nd specimens.
They have primary ribs at wide intervals,
numbering 5 or 6 per half whorl. Eachprimary rib arises from the umbilical bulla,which is prominent on the late part of the
body chamber. Often the rib is indistinctly
doubled on the flank, consisting of a stronger,
rectiradiate or gently flexuous major rib and aweaker, somewhat flexuous branched minor
one, both of which are looped at the outerventrolateral, more or less clavate tubercle,
whereas the inner ventrolateral node or bullate swelling is on the major one.
On the interspace between the primary ribs
there are a few weaker ribs, some of whicharise from weak umbilical bullae singly or
doubled and looped at the small and weak
inner or outer ventrolateral nodes, dependingon the presence or absence of either of thelatter two. Some other weak ribs have noumbilical node. These weak ribs or subcostae are rather irregular in curvature, but areoften gently convex on the flank and somewhat rursiradiate on the ventrolateral part.
Also faint, inner or outer ventrolateral nodes
or swellings mayor may not exist on these
subcostae. The arched, weak ribs, sometimes
doubled, seem to occur on the venter but are
obscured in these specimens by erosion.
The major ribs on the next inner whorl are
weaker and less distant than those on theouter whorl.
The third specimen shows a part of the
body chamber and the fourth one consists of9 12

Tatsuro M a lsum oLO

Figure 1. Pselldaspidoceras .flexuos lll1l POlVe 11. T he lirst spec imen MC M. A1l3 - I fr o m Y5599,
Kamim ak i-zalVa ofO yub a ri a rea. A, B : lateral and ven tral vielVs ; C : th e las t suture o f' G K. H9629, whic his a pl aster cast o f' th e ho lo type of Ps. ( Ampak abiles) kalVash ilai Matsum oto a nd Obata, 1982. A rr ow
indi cat es th e pos iti o n of th e last suture, or IV hi ch E a nd El L sadd le arc no t we ll exposed. (P ho tos by M.
Noda; T. M. de/in. )fr agmenta ry pieces o r a la te se pt ate who rl.
Th ey show simil ar mode o f' ribbin g to th at inth e o uter who rl o f the ro rego in g two spec ime ns. An a pp arent differen ce in th e o rn ament is th at both the o uter a nd innerve ntro latera l tubercl es are di stin ct in th e 1st
a nd 2nd spec imens but o nl y o ne o f' th em is

we ll deve lo ped in th e 3rd and 4th. Actu a ll y,tn th e 3rd spec im en th e clava te o uterve ntro latera l tub ercles a re well deve lo ped but
the inn er o nes are o bscure, whereas in th e 4th
spec im en th e inn er ventro latera l nodes arelarger. Fa int rib s run across th e ve nter witha gentl e fo rwa rd co nvexit y, show in g so me-927.

Turonian acanlhoceralid ammoniles

9 13

o

30mm
Figure 2. Pselldaspidoceras jlexliOSlIl11 Powe ll. The third specimen MC M. A 193 - 1 fr o m Y SI6Sb,
T a ki-n o-sawa o f Oyubari a rea. A, B: latera l and ve ntral vie ws; C : whorl secti o n at Q ; D : the firstlateral lo be L o n the inner who rl. (Ph otos by M. Noda ; T.M . de/in.)times a d o ubled feature . An obscure zo ne ofelevation is di sce rnible on the mid- ve nter in apart of the 4th specimen.
Sutures of the late growth- stage are observab le on the specimens (I) , (3) a nd (4). They
are simil a r to those illu strated finel y by Co llignon (1965 , pI. 389, fig . 1664) in sho w ing

tall a nd rather n a rrowed stems of El L and Lj
U2 sa ddles, fairly deep lobul es and so mewh a tphylloid termin a ls of foliole s. The breadthof L, howe ve r, see m s to be variab le in thi s
species, ra ngi ng from very broad as inPowell 's (19 63 ) tex t-fig. 2f to fa irly narrow as
in Co llignon 's (/oc. cit.). In thi s respect ourTatsuro Matsumoto

914
Table 1.
Specimen, remarks

Measurements (in mm) of Pseudaspidoceras fiexuosum.
D

U

H

B

B/H

E
E-90' (LS+25')
LS+ ISO'
restored
Q in Fig. 2E-90'-200(1)205(1 )
-250(1)-525(1)76(.32)60(.29)81(.32)185(.35)

103(.43)92(.35)95(.38)205(.39)-78(.33)
72(.35)-70(.28)

0.76
0.77
0.74

188

-\08

066*

\08

85

0.79

Holotype (Powell, 1963)
1/
Paratype ( II)Collignon, 1965, f. 1662
Cobban & Scott, 1972

-230( I)
145(1 )
132(1)
165(1 )

83(.36)49(.34)46(.35)48(.29)

0.84

(I)
1/

(2)
(3)
1/

(4)

94(.41)49(.34)34(.26)48(.29)99(.43)57(.39)62(.47)73(.44)

0.87
0.74
0.66*

D=diameter, U = width of umbilicus, H=whorl-height, B=whorl-breadth; E=at the preserved end,
E-90' = at the point 90' adapically from E, LS = at the last septum, LS + ISO' = at the point ISO' ado rally
from LS,* = secondarily compressed; - = approximate or inferred. Hokkaido specimens (1) - (4) as
indicated in Material. For comparison, measurements on selected foreign specimens are shown below
the line (cited from literature).

examples (Figures IC, 20 and 3E) andanother in Kennedy et al. (1987, fig. 3C) are
intermediate. Also the subdivision of L
varies from bipartite to asymmetrically tripartite.
Dimensions.-See Table 1.
Remarks.- When Matsumoto and Obata(1982, p. 75) established P. (Ampakabites)
kawashitai on specimen (I), they depended
on Collignon (1965) and Cobban and Scott(1972) in evaluating Ampakabites as a distinct taxon of generic rank.
Now, I however follow Kennedy et al.
(1987, p. 34-41), who have shown a great
extent of variation in P. flexuosum Powell,1963, with which Kamerunoceras (Ampakabites) auriculatum Collignon, 1965 andAmpakabites collignoni Cobban and Scott,1972 have been synonymized. Similarly, P.
(A.) kawashitai Matsumoto and Obata, 1982
should be also regarded as a synonym (seeabove description and comparison below).
Comparison. - The fairly large specimens(1) and (2) from Hokkaido have well separated inner and outer ventrolateral tubercles on
the primary ribs on the outer whorl. Theyclosely resembles the holotype of P. fle-xuosum designated by Powell (1963, pI. 32,
figs. 1,9 and text-fig. 2c) in shell size (0=230mm estimated by Powell; "medium-size" byKennedy et al., 1987, p.38 is inadequate,
because the photograph in Powell, 1963 is
reduced to X 0.5), whorl shape and ornament,
although the latter is only a fragmentary bodychamber. With respect to the distinctness of
inner and outer ventrolateral tubercles, our
specimens (1) and (2) are comparable with
Ampakabites collignoi Cobban & Scott (1972,
p. 81, pI. 29, figs. 1-3; text-figs. 39,40), fromthe Bridge Creek Limestone Member of theGreenhorn Limestone of Colorado, although
the latter is smaller and seems to be secondarily compressed.
A fragmentary septate whorl, UMUT.
MM6890, from the Obirashibe area, which
was called Mammites sp. by Yabe (1909, p.
411) and described by Matsumoto et al.
(1978, p.14, pI. 6, fig. 2; text-fig. 6) under
Ampakabites (?) sp., is similar to the form
mentioned above in the outline of whorl
section, major ornament and sutural pattern,
but I hesitate to make the specific identification until more specimens are obtained
from the corre1atable stratigraphic level in the927.

Turonian acanthoceralid ammonites

Figure 3. Pseudaspidoceras flexuosul11 Powell. Fragmentary pieces of the fourth specimen GK.
H8367 from Y6015c, Taki-no-sawa of the Oyubari area. A - D: late part of the sep tate whor l in lateral view
(A, C), whor l-section at Q (8) and ventral view (D). E: rubber model taken from the composite externa lmould of a middle-aged part of the same indi v idu a l, showing impressed sutures. (T.M. de/in; photos by M.
Noda)

915

916

Tatsuro Matsumoto

Obirashibe area.
In the 3rd specimen from the Oyubari area,
the inner ventrolateral (i.e., outer flank) tubercles are weakened or obsolete and the outerventrolateral clavi are distinct. In this respect
this form resembles the Madagascar specimens described by Collignon (1965, p. 29, pI.
388, fig. 1662; p. 31, pI. 389, fig. 1664) andsome examples from western Texas (Kennedy
et al., 1987, text-figs. 7A-C). The large but
still septate 4th specimen shows distinctly the
inner ventrolateral (or rather outer flank)
tubercles but the outer ventrolateral clavi are
obscure.
Despite a large number of specimens fromwestern Texas, no example of complete preservation has been ill ustrated by Kennedy et al.
(1987). An example from Hokkaido, shownin Figure I, may therefore represent one of thebest preserved specimens of P. flexuosul11. Inthat specimen the last two sutures are approximated and situated at about 175 to 185 mm
in diameter. Its body chamber occupies atleast half a whorl, although its final ventral
and outer lateral portions were destroyed.
Discussion. - On the basis of the two morphotypes from Hokkaido, I would suggest as
a working hypothesis that they might represent the dimorphs, viz. a microconch whichshows distinctly both the inner and outerventrolateral tubercles even in the adult stage
and a macroconch in which either of theinner and outer ventrolateral tubercles are
reduced in more or less late growth-stages.
The diameter of the latter in the adult stage isvery large, being roughly twice as large as theformer. Whether this interpretation could
be maintained in a large number of specimensfrom the type locality in western Texas andcontiguous areas or the feature is merely avariation should be decided by further investigation.
Occurrence. - Summarizi ng the local ity
records in the item Material, the describedfour specimens occur in the lower part ofMember lIn, which is assigned certainly to thebasal Turonian. Integrated data of biostrati-

graphy in this area are described in anotherpaper by Matsumoto et al. (1991).
P. flexuosul11 is one of the most diagnostic
species in the basal Turonian of western
Texas (Kennedy et al. 1987) and has beenreported to occur in New Mexico, Arizona,
Colorado, Germany, Nigeria and Madagascar. This is the first record of P. flexuosum
from Japan. The fact that four individuals
were found within a limited area seems tosuggest that this species is not rare in Japan.
We should search for more examples in thisand other areas in Hokkaido.
Genus Neol11phaloceras Matsumoto
and Obata, 1982
Type species. - Yubariceras pseudol11phaIum Matsumoto, 1975.
Diagnosis. - Very similar to Cunningtoniceras Collignon, 1937 in the shell-form, ornament and suture, but distinguished in having
an additional row of lateral tubercles.
Discussion. - Cunningtoniceras Collignon,
1937 has been revived by Cobban (1987, p. 9),
who noticed a peculiar sutural pattern inEuol11phaloceras of the correct sense, whichshows narrow ElL saddle and very broad,roughly bipartite L.
Cunningtoniceras has the normal acanthoceratid sutures and is regarded as a direct
derivative of Acanthoceras (see Wright and
Kennedy, 1987). It is characterized by multiple ribbing and tuberculation on the outerpart of the whorl at least in part and also
development of ventrolateral horns on theadult whorl. Cobban et al. (1989) havedescribed three species of Cunningtoniceras
recently found from the late Cenomanian
Metoicoceras mosbyense Zone of New Mexico, whereas a rare example of Cunningtoniceras has been recorded from the Middle
Turonian of Japan (Matsumoto and Suekane,1989).
In my view, Neol11phaloceras can possibly
be regarded as a descendant of Cunningtoniceras acquiring an additional row of lat-

927

Turonian acanthoceratid ammonites

eral tubercles. It retain s ancestral charactersin showing extra ribs and tubercles on theventral part of immature whorls and horned
ventrolateral tubercles In the adult. Its
suture is also quite similar to th at of
Cunningtoniceras. A weak point in thisinterpretation is the apparent missing link in
the latest Cenomanian and earliest Turoni a n(see discussion below). Possibly so me
marine biota may have been affected by frequent unfavourable conditions, such as anoxic events, during this time interval. One may
recall here the almost absence of fossil
nautiloids in Pliocene and Pleistocene strata

9 17

until the rejuven ated appearance of the
Recent species of Nautilus (Teichert andMatsumoto , 1988).

Neomphaloceras pseudomphalum(Matsumoto)Figures 4 and 5

Ylibariceras pseudomphalum Matsumoto , 1975, p.
146, pI. 22, fig. I.

Neompha loceras

pseudomphalum (Matsumoto):
Mats umoto and Obata, 1982, p. 73, pI. 2, fig. I ;pI. 5, fig. I.
Neomphaloceras pseudomphallim (Matsumoto):
Matsumoto el al., 1985, p. 162, pI. 5, fig. I.

Figure 4. Neomphaloceras pselldomphalum (Matsumoto). An immature spec imen , YCM . GP716.
A, B: lateral and ve ntra l views; C: external suture at S; ribs a nd tubercles are drawn by dotted lines;
constriction-like groove hatc hed. (reproduced from Matsum oto a nd Suekane, 1987, pI. 4, figs. 1, 2 a nd
text-fig. 3 of Sci. Rept. Yok os uk a City Museum, no. 35 by permission).

918

Tatsuro Matsumoto-.. ---------~.., ............
-

I

'. I

I,

I

I

'\
"

\

:,

1/

'I

I

"
-~."

:\
,/

'"

B

Figure 5. Neomphaloceras pselidomphallll1l (Matsumoto). An example of an adult shell, MCM.
A 175-1, from the Kaneobetsu-zawa of the Oyubari area. A: lateral view; B: whorl-section at Q; C: thesecond last suture. (T.M. delin.)

Neomphaloceras pselidomphallim (Matsumoto);
Matsumoto and Suekane, 1987, p. 10, pI. 4, figs.
1-2,4-7, 10-11 ; text-fig. 8.

Material.-In addition to the eight specimens described in the above-listed papers,
here is a fine specimen, MCM. A175-1 (Figure 5) collected by Koji Hasegawa from an
outcrop of the Kaneobetsu-zawa, Oyubari
area.
Remarks. - The medium-sized holotype
and YCM. 717, one of Suekane's specimens,
represent the shell at middle growth-stage.
The smaller para type and the three specimens
described by Matsumoto and Suekane (1987)
show the characters of younger stages. The
two larger specimens described by Matsumoto
and Obata (1982) and Matsumoto et al.

(1984) are adult but their body chambers areincomplete. K. Hasegawa's specimen represents a nearly complete adult shell. I haveseen more specimens in Y. Kera's private
collection from the Oyubari area. On thebasis of these specimens the specific diagnosis
may be given as follows:
Diagnosis. - The shell is fairly large in theadult and rather evolute, showing a moderate
rate of whorl expansion, a little (i.e. about II4) involution and a moderately wide umbilicus. The whorl is somewhat broader
than high (e.g., B/H= 1.24 in measured examples) and widely subrectangular or subquadrate in cross section, with a broadly archedor nearly flat venter, subparallel flanks,
abruptly bent umbilical shoulders and vertical umbilical wall.

927.

Turonian acanthoceratid ammonites

At first the shell looks nearly smooth, onlywith constrictions and associated flares; soon
ribs appear on which tubercles are developedin nine rows. The young and middle-aged
whorls have moderately strong primary ribs
and also occasionally intercalated or branched, weaker or somewhat shorter ribs.
These ribs are nearly rectiradiate on the mainpart of the flank and curved forward to forma ventral projection. There are furthermore
branched or intercalated extra riblets normally with tubercles in three rows on theventer, some of which may be projected considerably and accompanied by a constrictionlike groove. The tubercles are in nine rows,of which the three rows (i.e. umbilical, lateraland inner ventrolateral) on each flank arenearly equidistant (see Matsumoto, 1975, pI.
22, fig. 1 and Figure 4 of this paper).
The outer whorl of an adult shell, of which
the body chamber occupies slightly more thanhalf (i.e. about 200'), has distant major ribs
with occasionally intercalated weaker andsomewhat shorter ribs. The tubercles on the
major ribs are prominent on the flank but
much weakened or almost obsolete on theventer. The inner ventrolateral tubercles on
the body chamber are especially strengthened
into horn-like protuberances, amalgamating
the weakened outer ventrolateral tubercles
into their thickened basal part. A riblet
associated with a narrow and shallow constriction may remain persistently on the ventral part of the last septate whorl. On thelast portion of the body chamber a few nodeTable 2.

919

less ribs may occur probably as flanges on theapertural margin.
The suture is of typical Acanthoceras pattern, showing massive, bipartite EjL saddle
and fairly narrow L.
Dimensions.-See Table 2.
Comparison and discussion. - The young
to middle-aged form of this species resembles"Acanthoceras" carvertense Powell (1963, p.
315, pI. 33, figs. 8-9; pI. 34, figs. 6, 9; textfig.2e), from the Lower Turonian of west
Texas-north Mexico, in shell form and ornamentation, but for the presence of an additional row of lateral tubercles in the former.
As to the latter species Powell (1963, p. 136)
suggested its affinity with Kanabiceras, i.e.
Euomphaloceras redefined by Cobban (1987),
whereas Kennedy et al. (1987, p. 33) referred
it to Kamerunoceras.
Matsumoto et al. (1989, p. 30) have notedthat there are two types of sutural pattern inthe species of Kamerunoceras defined by
Kennedy and Wright (1979); one is the normal type like that of Acanthoceras, as seen in
K. eschii (Solger), the type species, and K.
turoniense (d'Orbigny) and the other is thepeculiar type like that of Euomphaloceras (inthe sense of Cobban, 1987) as seen in therevived Sch in de wolfites. As I have not examined the specimens of 'Ac.' carvertense, I
can hardly decide which type of sutural pattern is shown by this species. The too brief
description of the suture without illustration
by Kennedy et al. (1987, p.34) does notenlighten me. If its suture has narrow EjL

Measurements (in mm) of Neomphaloceras pseudomphalum.

Specimen, remarks

DU

Holotype, E(c)II
E-45'(ic)93.0( I)

30.0(.32)

Paratype (c)Fig. 5 LS+ 70'(c)
II
LS(c)

19.3(1 )6.6(.34)

206.0(1 )167.0(1)62.0(.30)
49.0(.29)

H

B

BjH

MR:VR
7: 15

35.0
7.5(.39)88.0(.43)

42.0
10.2(.55)

1.20
1.24

71.0(.43)-88 (.53)

1.24

7:8
6:9

MR: VR=number of major ribs versus that of all ribs on venter per half whorl, (c)=costal, (ic)=intercostal. See Table I for others.

920

Tatsuro Matsumoto

30mm...............

A

Figure 6. Neomphaloceras costalum (Matsumoto and Kawashita) .
Lateral (A) and ventra l (8) views of the holotype, MeM. A45- 4. (Photos by M. Noda)

saddle and very broad L li ke that of Euomphaloceras , it sh ould be assig ned toSchindewoljites or rather to Paraburroceras
Cobban , Hook and Kennedy , 1989. If it isnormal Acan/hoceras type, that species could
be either Kamerunoceras or possibly immature Cunningtoniceras. In the latter casethat spec ies could be ancestral for N. pseudomphalum , being a candidate which cou ld filla part of the missing link mentioned in theforegoing page.
Occurrence. - So far as reli a ble locality
record s are concerned , this species occurs in
the unn a med zone immediately below thezone where the typical form of In oceramus
hobe/sensis Nagao and Matsumoto is characteristic in the Obirashibe, A shibetsu andOyubari areas. The known age is early
Middle Turonian , although the true vertica l

range should be determined by further investigation.

Neompha loceras cos/atwn(Matsumoto and Kawashita)
Figures 6 and 7

Mammil es cos/allis Matsumoto and Ka was hita in
Ma tsum o to e/ al. , 1978 , p. 5, pI. I, figs . 1, 2; pI.
2, figs . 1,2; te xt-figs. 1, 2.

Mammi/es lVingi lVingi Mo rrow ; Wright a nd Kenned y, 198 1, p. 80- 8 1 (pars.).

Neomphaloceras cf. pseudomphalum (Matsu moto) ;Ma tsu mo to el al. , 1985 , p. 163, pI. 6, fig. 1

Material. - The following specimens arereexamined , in which the temporary numbering in previous papers is indicated 111
brackets; holotype, MCM. A45-4 [Yo Kawashita's ColI. 51 - 8- 20J (Figures 6, 7) in a nod-

927.

Turonian acanthoceratid ammonites

ule fallen loose in a small stream called the
Shogakko-no-sawa, Kashima; paratype,
MCM. A45-5 [Kawashita's Coll.51-11-14J(plaster cast GK. H9630) (Matsumoto et al.,1978, pI. 1, fig. 2; pI. 2, fig. 2) from the outcrop of mudstone with interbeds of sandstone
at loco Y5201 on the right side of the Hakkinzawa; another illustrated specimen under
Neomphaloceras cf. pseudomphalum by Matsumoto et al. (1985, pI. 6, fig. 1), from theoutcrop of mudstone at loco Y5240 on theright bank of the Kaneobetsu-zawa, close to
the confluence with the Shuparo River.
Diagnosis. - The whorl is rather evolute,
expanding at a moderate rate, and encircles
an umbilicus of moderate width. The adultshell is large, with diameter well over 300 mm
in the holotype. The whorl is slightly
broader than high and subquadrate in section, with a roughly flat venter, subparallel
flanks; abruptly bent umbilical edge andnearly vertical umbilical wall.
The ribs are distant on the outer whorl, as-------

~~

10 ••
~

Figure 7. Neomphaloceras costa tum (Matsumoto and Kawashita). A: whorl-section at Q of
Fig. 6; B: the second last suture of the holotype.
(T.M. delin.)921

a rule alternately long and short on the septate part and almost all long on the adultbody chamber, with occasional exceptions.
The tubercles are in nine rows, viz. those at
the umbilical edge on the long ribs, more orless bullate; those somewhat inside themidflank, conical earlier but bullate later andwhich may weaken or become obsolete especially on some shorter ribs; those at the inner
ventrolateral shoulder are strong and thick,enlarging with growth, and horn-like on theadult body chamber; the outer ventrolateralones weakening and finally amalgamated by
the basal part of the horns; those on thesiphonal line which may remain as traces(indistinct elevations) on the broadened but
lowered ribs on the venter.
The ribs are nearly recti radiate on the mainpart of the flank, gently curved forward on theouter flank toward the large ventrolateral
nodes, and lowered and broadened on theventer, showing a broad convexity. A few
extra riblets may remain faintly even on theventer of the adult stage. Near the apertural
margin of the body chamber there are a few
ribs which are not so widely separated as theribs on the main part and the tubercles onthem are blunted. They may have worked asflanges along the peristome at the final
growth-stage.
The suture is of normal acanthoceratid
pattern, showing deep E, massive, broad andbipartite ElL saddle, relatively narrow andfairly deep L, rather obscurely bipartite L/U2
saddle of moderate breadth and smaller U2
and auxiliaries.
Dimensions.-See Table 3.
Discussion. - Mammites costatus Matsumoto and Kawashita, 1978 was regarded by
Wright and Kennedy (1981, p. 80) as a syno~
nym of Mammites wingi Morrow, 1935
because of the good similarity in size, whorl
section, ribbing and tuberculation. There is,however, an unmistakable difference in the
suture. M. wingi has narrow (taller thanwide) ElL saddle and peculiarly broad, bifid
L. For this and other reasons Cobban andTatsuro Matsumoto

922

Table 3.

Measurements of Neomphaloceras cos/a tum

Specimen, remarks

D

U

H

Holotype, E-300(e)324.0( 1)95.0(.29)

132.0(.41 )

B

B/H

MR:VR
5:7

1/

Q in Fig. 7(e)

108.0

119.0

1.10

5:7

1/

2nd LS (ie))

105.0

112.0

1.07

5:7

Hook (1983, p. 9) have proposed the genus
Morrowites, with M. wingi Morrow as its typespecies. I agree with them. Another pointis that this species has a row of flank tubercles, which are added to and not displaced
from the umbilical tubercles. Moreover, it
has a siphonal row of tubercles, even if they
weaken in the late growth-stage.
In short the said similarity is a homeomorphy and this species cannot be referred toMorrowites wingi nor to any species ofMammites. The development of Mammites-like horned ventrolateral tubercles on its
subquadrate outer whorl is anotherhomeomorphy. I was sorry to have carelessly assigned it to Mammites, but now I shouldrefer it to Neomphaloceras.
Comparison. - N. costa tum resembles N.
pseudomphallim but they can be discriminated; viz. N. pseudomphalum has broader
whorls with larger ratios of B/H, narrower,
less predominent ribs, more prominent tubercles, of which the laterals are more separated
from the umbilical ones and the ventrolateralhorns are more pronounced, as comparedwith N. costatum. They seem to be stratigraphically separated.
This species looks somewhat similar to
Mammites daviesi Spath (1935, p. 415, fig. I;
pI. 32), from Ramri island (Southeast Asia),
and to M. mocamedensis Howarth (1966, p. 6,
pl. 3), from Angola, in the coarse ribbing, but
the latter two species should be referred toMorrowites because of their pecliar sutures.
They show no lateral tubercles or united
ventrolateral horns.
Occurrences. - The age of this species wasincorrectly inferred as early Turonian.
Through the stratigraphic reinvestigation(Matsumoto et al., 1991) it should be revised
to Middle Turonian. The localities of theexamined specimens are all in the Inoceramus
hobetsensis zone of the Oyubari area andsome of them (e.g., Y5240) are rather in ahigher part of this zone. I should, however,investigate further to know the true stratigraphic range of this species and its phylogenic
relationship with N. pseudomphallim.
Genus Mammites Laube
and Bruder, 1887
Type species. - Ammonites nodosoides
Schlliter, 1871.
Diagnosis.- The adult shell is fairly to verylarge in the type and related species but
medium-sized in some others. The whorl issubquadrate to subrectangular or subtrapezoidal in section. The ribs are coarse andmore or less distant but may be more crowdedwith branched or intercalated ones. The
umbilical tubercle is conical and/or bullate;the inner and outer ventrolateral tubercles
normally more or less clavate, of which theinner one is prominent, enlarging on theouter whorl to a strongly protruded horn andabsorbs the outer one. The suture is similar
to that of Acanthoceras, showing fairly deepE, massive or broad, bipartite E/L saddle andrather narrow L.
Discussion. - Wright and Kennedy (1981, p.
75) stated that the relative width of E/L and
L varies considerably in Mammites. I do
not think the said variation is gradational,
and recognize two different types. I agreewith Cobban and Hook (1983, p.9), whohave established Morrowites for the Mammites-like ammonites in the shell-form and927.

Turonian acanthoceratid ammonites

ornament but have an "unusually broad, bifid
lateral lobe (L)" and rather narrow and tall
ElL saddle. The difference is clearly shownby their illustrations (op. cit., text-figs. 4, 5, 7
in comparison with text-fig. 2). How theunusual pattern was formed seems to be suggested by their text-fig. 6A, B. This is quite
similar to what I stated (in Matsumoto et al.,
1989, p.30) about the sutures of Cunningtoniceras, Lotzeites and Euomphaloceras.
Cobban and Hook (1983) described,
besides the type species, Morrowites wingi(Morrow), Morrowites depressus (Powell),Morrowites subdepressus Cobban and Hookand Morrowites cf. dixeyi (Reyment). I havepointed out above that Mammites daviesi
Spath and M. mocamedensis Howarth shouldbe also referred to M orrowites.
Remarks. - Examples of Mammites do
occur in the Cretaceous of Japan, but they arestill few or poorly preserved. I should, however, provide here some information so that
more material can be added in the future.

Mammites aff. nodosoides (SchlUter)
Mammites aff. nodosoides (Schlotheim); Matsumoto,
1981, p. 58, fig. 2.

Descriptive remarks.-I examined in the
Mikasa City Museum a specimen collected by
H. Oyobikawa from the Nakakinembetsu, atributary of the Obirashibe River. Although
its right side is dissolved in the rock matrix
for a taphonomical reason, its left half is
fairly well preserved. The dimensions (inmm) at the preserved end (intercostal) are as
follows:
D=257 (I) U=98 (.38) H=9l (.35)
B (restored)=61 (.24) BjH=0.67
The body chamber is preserved for about half
a whorl and is nearly complete. This ammonite is thus similar in size to some adult shellsof Mammites nodosoides (SchlUter) (seeWright and Kennedy, 1981), but its whorl ismore evolute and has a wider umbilicus than
any illustrated or measured examples of M.

923

nodosoides.
The ribs on the outer whorl of this specimen are distant and especially widely separated on the last part of the septate whorl andthe early half of the body chamber. I ntercalated shorter ribs occur only on the inner
whorl. The umbilical tubercles are bullate
and strengthen in the late stage. On the bodychamber the ventrolateral horns are welldeveloped, stretching upward and somewhat
laterally.
The suture is of normal Acanthoceras type.
To sum up, this specimen generally resembles some examples of M. nodosoides in theornament and suture, but there is somedifference in the frequency of shorter ribs anddegree of involution. Although there are a
few more but poorly preserved specimens of
Mammites from loco R2307 (Nakakinembetsu) of the Obirashibe area and loco Y60l7(Taki-no-sawa) of the Oyubari area, they are
insufficient to know the extent of variation in
our form. I should call the described specimen provisionally Mammites aff. nodosoides(SchlUter).
Occurrence. - Although the Lower Turonian Member Mj of the Middle Yezo Groupis exposed fairly extensively along the River
Naka-kinembetsu of the Obirashibe area, theexact point of the locality is not recorded forthe described specimen. Loc. R2307 where acomparable specimen was obtained is certainly assigned to Member Mj and My t ilo ides
goppelnensis Badilet and Sornay occurs in thesame member at a nearby locality. Anotherspecimen at loc. Y6017 came from the Lower
Turonian Member lIn in the Taki-no-sawasection of the Oyubari area.
Mammiles sp. nov. (?)
Figure 8

Material. - A single incompletely preserved specimen, GK. H8371 (Fig. 8) collected by
M. Koshizaka (August 10, 1972) from a transported nodule in the lower course of theTaki-no-sawa (640 m upstream from its

Tatsuro Matsumoto

924

mouth) , Oyub a ri area.
Description . - Although the specimen IS Incomplete, it shows a part of the bod y chamberand some pa rt of the septate whorl as sho w n
in the photograph. For a taphonom ical reason the left ha lf is obscured but can be resto red from the right half. Thu s, its ap proxim ateintercostal dimen sions (in mm) are as fol lows :D = 133 (I) U = 48 (.36) H = 49 (.37)
B= 56 (.42) Bj H = 1.14
The whorl was originally so mewhat
bro ader than high . broadest at a bout the
mid-height a nd subrounded in the interco stal
section , with a broadl y rounded to nea rl y fl at
venter a nd gent ly convex flank , w hich passes
to a steep ly inclined umbilical wall , without
forming sub ang ular edge. The whorl is----------

10 mm'"'-----'

B,............

..... ,l

\ I

AFigu re 8.
K os hi za ka .
M. de/in.)

Mammiles sp. nov. (?). GK. H 837 1 from the Taki-no-sawa, collected and donated by M.
A : latera l view; B: restored w ho rl- secti on ; C: su ture a t S of A . (Photo by M. N oda; T.

927.

Turonian acanthoceratid ammonites

rather evolute and the umbilicus is of moderate width.
The shell is ornamented with simple, rectiradiate, long and rather narrow ribs, which
are widely separated on the outer whorl andtheir interspaces are somewhat wider than theribs even on the next inner whorl. The ribs
have bullate tubercles on the inner flank of
the septate whorl and considerably high,bullate ones at the mid-flank of the bodychamber. The inner ventrolateral tubercles
are spinose on the septate whorl, stretchingalong the umbilical wall of the outer wall.
The outer ventrolateral tubercles, whichshould exist at this stage, are not observableowing to the unfavourable state of preservation. On the body chamber the inner andouter ventrolateral tubercles are united into ahigh but rather narrow spatulate protuberance, which is not clavate but extends to abar-like rib, running straight across theventer, though with decreasing height toward
the mid-venter.
The suture is like that of normal Mammites; viz. EjL saddle is broad, L deeper
than wide; LjU2 saddle situated on the row
of inner lateral tubercles.
Comparison.- This specimen is distinct
from the lectotype and other specimens of
Mammites nodosoides in its subroundedintercostal whorl section, narrower major ribs
without branched or intercalated minor ribs,
narrowly bullate tubercles on the inner to
middle part of the flank instead of rather
conical umbilical nodes and the spatulate
ventrolateral protuberances extending to thebarlike ribs on the venter, instead of rather
clavate ventrolateral horns, on the bodychamber. Therefore, this specimen probably
represents a new species, but I choose topostpone proposition of a new name until
better preserved specimens can be obtained insitu from a well-defined stratigraphic position.

Discussion. - Pseudaspidoceras cornucostale Morrow (1935, p. 469, pI. 51, fig. I; textfig. 5), from the Greenhorn Formation of925

Kansas, seems to have somewhat spatulate
ventrolateral horns on the outer whorl, but its
ribs are thicker and have nodes at the umbilical edge. Its suture is quite dissimilar to thatof normal Mammites, showing narrow andasymmetric EjL saddle and very broad L.
Occurrence.-As for Material. The place
where Koshizaka collected this ammonite as aboulder is occupied by Middle Turonian
strata. As the specimen is much eroded, it
may have transported from a fairly upstream
part where Lower Turonian strata are well
exposed.

Acknowledgements
Director Mamoru Kera and Mr. Kikuo
Muramoto of the Mikasa City Museum kindly afforded me facilities to study there, especially for the study of the valuable specimens
which Messrs. Yoshitaro Kawashita, Koji
Hasegawa and Hiroshi Oyobikawa collected.
Mr. Masatada Koshizaka generously presented an interesting ammonite from his collection to Kyushu University for this study andProfessor Hiromichi Hirano and Mr. Yasuharu Kera kindly showed me other interesting
specimens with necessary information. I am
grateful to Dr. W.A. Cobban and Dr. M.K.
Howarth for their instructive suggestions.
Dr. Masayuki Noda kindly helped me in thefield work and also in taking photographs.
Miss Akiko Murakami and Miss Kazuko
Hara assisted me in preparing the typescript.

References cited
ArkelI, W.J., Kummel, B. and Wright, C.W., 1957:
Mesozoic Ammonoidea. In. Moore, R.C. ed..
Treatise on Invertebrate Paleontology. Part L,
p. LSO-L.465. Geo!. Soc. America and Univ.
Kansas Press.
Cobban, W.A., 1987: Some middle Cenomanian(Upper Cretaceous) acanthoceratid ammonites
from the Western Interior of the United States.
u.s. Geol. Surv.. Prof Paper 1445, p. 1-28, pis.
1-13.
- - , 1988: Tarrantoceras Stephenson and related
ammonoid genera from Cenomanian (Upper

926

Tatsuro Matsumoto

Cretaceous) rocks in Texas and the WesternInterior of the United States. U.S. Geol. Surv.,
Prof Paper 1473, p. 1-30, pis. 1-10.
- - and Hook, S.c., 1983: Mid-Cretaceous (Turonian) ammonite fauna from Fence Lake area ofwest-central New Mexico. New Mexico Bureau
Mines & Mineral Res., Mem. 41, p. I-50 (incl.
14 pis.).
- - , - - and Kennedy, W.J., 1989; Upper
Cretaceous rocks and ammonite faunas of southwestern New Mexico. Ibid., 45, p. 1-137 (incl.
96 pis.)- - and Scott, G.H., 1972: Stratigraphy and ammonite fauna of the Graneros Shale and Greenhorn
Limestone near Pueblo, Colorado. U.S. Geol.
Surv., Prof Paper 645, p. 1-108, pis. 1-41.
Collignon, M., 1937: Ammonites cenomaniennes dusud-ouest de Madagascar. Ann. Geol. Servo
Mines, vol. 8, p. 31-69, pis. I-II.
- - , 1965: A tlas des fossiles caracteristiques deMadagascar (Ammonites), vol. 12 (Turonien), p.
1-82, pis. 376-413. Service Geol., Tananarive.
Howarth, M.K., 1966: A mid-Turonian ammonite
fauna from the Mo~amedes desert, Angola.
Garcia da Drla (Lisboa), vol. 14, p. 217-228, pis.
1-3.
Kennedy, W.l. and Wright, C.W., 1979: On Kamerunoceras Reyment, 1954 (Cretaceous Ammonoidea). JOllr. Paleont., vol. 53, p. 11651178 (incl. 4 pis.).
- - , - - and Hancock, J.M., 1987: Basal Turonian ammonites from West Texas. Palaeontology, vol. 30, p. 27-74 (incl. 10 pis.).
Matsumoto, T., 1942: Fundamentals in the Cretaceous stratigraphy of Japan. Part I. Mem. Fac.
Sci. KYllShu Imp. Univ., Ser. 0, vol. I, no. 3, p.
129-280, pis. 1-20.
- - , 1975; Additional acanthoceratids from Hokkaido. Mem. Fac. Sci. Kyushu Univ., Ser. 0,
vol. 22, p. 99-163, pis. 11-23.
- - , 1981 : Plastotypes of Cretaceous ammonites
from abroad donated subsequently to Kyushu
University. Sci. Rept., Dept. Geol., Kyushll
Univ, vol. 14, p. 57-62 (in Japanese with English
abstract ).
- - , 1988: A monograph of the Puzosiidae (Ammonoidea) from the Cretaceous of Hokkaido.
Part I. Major classification. Palaeont. Soc.
Japan., Spec. Papers, no. 30, p. 3-28.
- - , Y. Kawashita, Fujishima, Y. and Miyauch, T.,
1978: Mammites and allied ammonites from theCretaceous of Hokkaido and Saghalien. Mem.
Fac. Sci. Kyushu Univ., Ser. 0, vol. 24, p. 1-24,

pis. 1-6.
- - , Maiya, S. and Noda, M., (1991): Towards an
integrated ammonoid-, inoceramid- and foraminiferal biostratigraphy of the Cenomanianand Turonian (Cretaceous) in Hokkaido. Jour.
Geography., vol. 100, no. 3, p. 378-398 (inJapanese with English abstract)- - and Obata, /., 1982: Some interesting acanthocerataceans from Hokkaido. Bull. Natn. Sci.
Mus., Tokyo, Ser. C, vol. 8, p. 67-92.
- - , - - and Kawashita, Y., 1985: Notes on someacanthoceratid ammonites from the Oyubari area,
Hokkaido. Ibid., vol. 11, p. 155-182.
- - and Suekane, T., 1987: Some acanthoceratid
ammonites from the Yubari Mountains, Hokkaido-Part I. Sci. Rept., Yokosuka City Mus.,no. 35, p. 1-14, pis. 1-4.
- - , - - and Kawashita, Y., 1989: Ditto-Part 2.
Ibid., no. 37, p.29-44.
Morrow, A.L., 1935: Cephalopods from the Upper
Cretaceous of Kansas. Jour Paleont., vol. 9, p.
463-473, pis. 49-53.
Powell, J.D., 1963 : Cenomanian-Turonian (Cretaceous) ammonites from Trans-Pecos Texas andnortheastern Chihuahua, Mexico. Ibid., vol. 37,
p. 309-322, pis. 31-34.
Spath, L.E., 1935: On a Turonian ammonite (Mammites daviesi) from Ramri Island, Burma.
Records, Geol. Surv. India, vol. 68, p.414-416,
pI. 32.
Tanabe, K., Hirano, H., Matsumoto, T. and Miyata,Y., 1977: Stratigraphy of the Upper Cretaceous
deposits in the Obira area, northwestern Hokkaido. Sci. Rept., Dept. Geol., Kyushu Univ.,
vol. 12, no. 3, p. 181-202 (in Japanese with English abstract).
Teichert, C. and Matsumoto, T., 1987 : The ancestryof the genus Nautilus. In, Saunders, W.B. andLandmass, H.H. eds., Nautilus, Chapter 2, p. 2532. Plenum Press, New York.
Wright, C.W. and Kennedy, W.J., 1981: The Ammonoidea of the Plenus Marls and the Middle
Chalk. Monogr. Palaeontogr. Soc., London, p.
1-148, pis. 1-32 (Publ. No. 560, part of Vol. 134
for 1980).
- - and --,1987: The Ammonoidea of the Lower
Chalk. Part 2. Monogr. Palaeontogr. Soc., London, p. 127-218, pis. 41-55 (Publ. No. 573, partof Vol. 139 for 1985).
Yabe, H., 1909: Zur Stratigraphie und Palaontologie
der oberen Kreide von Hokkaido und Sakhalin.
Zeitsch. Deutsch. Geo!. Gesell., vol. 61, p. 402444.

Ashibetsu p'jJlj, Hakkin-zawa B~i'R, Hokkaido ~twJ:l[, Kamimaki-zawa l::.~i'R, Kaneo-

927.

Turonian acanthoceratid ammonites

927

betsu ~~)JIj, Kashima J1[i1I;, Mikasa -=~, Naka-kinembetsu 9='~c~)JIj, Obirashibe IPF-~,
Oyubari :k:95lf, Shogakko-no-sawa IJ,~jjtO)iR, Shuparo -./ "- - /{ ", Taki-no-sawa lito)
iR, Yokosuka .~Jt
~twJiO)t:L!II!*7"- "- "=-7 '/iff.7 7J '/ r?-7 7-.f47 '/-<:-:T1 r ~c"?\',-C: ~twJiO)J::$
tl!ll!*7"- "- " =- 7 '/ (tJ r Tu c W~~) O)Jt!!Ji 7'1> I? iff. L. t.:lkO) 51'10) 7 '/ -<:-:T 1 r ~c "? \"-C, .:t 0) 5} M tJ: I? rHc Ji Ff I¥J mUjj 0) ~ Ji!. Z' j! ~ L. -C ~c).ffi L. t.:o (I) Pseudaspidoceras
flexuosllm Powello Tu ~r$O)~*1'1-c';i; 9 *1'17'1~fl~Jt!!,~O) I"J [Ji* (:k:95lfO)$Ji
lIn ~r$) 7'1> I?iff.:±l L. t.:o ~11 P. (Ampakabites) kawashitai Matsumoto & Obata 11*1'10)-./ / =- A1:;i;9 0 (2) Neomphaloceras pselidomphallim (Matsumoto)o Y4J. 9='. nIt~:MO)~*~C£0 ~~j~IC'a:'~~ L. t.:o 9='$ Tu O)r$~ciff.i' 9 0 *~0)*ki1E11 Cunningtoniceras ~~-c';i; 1':> ?
~~ L. t.:o (3) Neomphaloceras coslallim (Matsumoto & Kawashita)o *1'111~fJJ Mammites O)~1'1, 1&~cI1 Morrowites wingi (Morrow) 0) -./ / =- A C ~;h, -C \, ,t.:7'1~, ¥}~WO)*6* Neomphaloceras ~cA;h, 9 '"' ~ 1:, Ji* t 9='$ Tu O)./:t~I¥JJ:: $1:;i; 7.>0 (4)Mammites aff. nodosoides (SchlUter)o *~O) Mammiles It B *~C t \, ,t.:o M. nodosoides~C1);.l97'1~~~7'1~~<~7'1~J.t\,'0 fi\'jf":7'1;r$Tu (:t!t'(Et~tr) ~Ciff.i'90 (5) Mammitessp.
nov. (?) ~ffl[.iE~7'1~:k:95lf¥itO)iR1:J*~ • ~~0)~*-C·ft*~;h,90 7i-1JIUO)~0)~:!*~~

c7'1~"" 1?:!*1:~ ~ O)jJ[I:i] 'c~3C, .:t O)ft!!.~~ tdJ~Z'~i'7'1;, ~:O-C'W:ff

1'1O).ftZ'~L.~~,~~·~.L.-C~1&0)~~.MZ':Mi'90

t :>G5G~tJ:O)-c', ~~*.M

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, pp. 928-943,4 Figs., December 30, 1991

928.

LATE PERMIAN FUSULINACEAN FAUNA OF
DOl PHA PHLUNG, NORTH THAILAND*
KA TSUMI UENO
Institute of Geoscience,
University of Tsukuba, Ibaraki 305
andSUMIO SAKAGAMI
Department of Earth Sciences, Faculty of Science,
Chiba University, Chiba 260

Abstract. Eight species and seven genera of fusulinaceans are identified from two
localities in the lower member of the Huai Thak Formation in the Doi Pha Phlung area,
northeast of Lampang, north Thailand. They include Reichelina cf. changhsingensis Sheng
and Chang, Gallowayinella gllidingensis Liu, Xiao and Dong, Palaeofllslliina sinensis Sheng,P. cf. laxa Sheng and Codonofllsiella aff. kwangsiana fllsiformis Sheng. This fusulinacean
fauna is characterized by abundant Gallowayinella gllidingensis Liu, Xiao and Dong, issimilar to that in the Changxing Limestone and its equivalents in South China, and indicatesan early Changxingian or early Dorashamian age.
Key words. Changxingian, Doi Pha Phlung, Dorashamian, Gallowayinella. Huai
Thak Formation, Upper Permian.

north Thailand (Figure I), and Sakagami and
Hatta (1982) reported the Late Permian
Palaeofusulina-Colaniella Fauna from limestone blocks collected at the eastern foot of
Khao Doi Pha Phlung.
In this paper, a unique fusulinacean fauna
found at two localities in the lower memberof the Huai Thak Formation, in which Gallowayinella guidingensis Liu, Xiao and Dongis abundant, is described. The faunal affinity
and correlation are also discussed with special reference to the stratigraphic distributionof the genus Gallowayinella.
All specimens identified in this paper aredeposited in the Collection of the Departmentof Earth and Planetary Sciences, Faculty of
Science, Kyushu University. The abbreviation GK.D is the prefix for the specimens'
repository numbers.

Introduction

The Upper Permian formations inThailand are very restricted and paleontological information about them is limited incomparison to those of the Lower and Middle
Permian. To date, Upper Permian foraminifers have been reported only from sevenlocalities (Baum et at., 1970; Bri:innimann etal., 1978; Sakagami and Hatta, 1982; Hahn
and Siebenhtiner, 1982; Ingavat, 1984).
In the summer of 1988, we visited the Doi
Pha Phlung area and collected many rocksamples containing abundant Upper Permian
fossil remains from the Huai Thak Formation. This area is situated about 10 km
southwest of Ngao in the Lampang district of"Received June 25, 1991; revised
accepted October 25, 1991

manuscript

928

928.

Permian fusulinacean fauna of North Thailand

Acknowledgments

The present study was part of the project"Biostratigraphy and palaeogeography of theuppermost Permian and the Lower Triassic in
the southern realm of the Asian Tethys" (Project no. 6304199, Leader: Professor Juichi
Yanagida, Kyushu University), which was
financed by the Overseas Scientific Research
Fund of Ministry of Education, Science andCulture of Japan. We express our cordialappreciation to the members of the project
team: Professor J uichi Yanagida, Drs. Takeshi Ishibashi, Kazuyuki Matsusue of Kyushu
University, Drs. Nikorn Nakornsri, Chonpang Chonglakmani, Ms. Yaowalak
Chaimanee of Geological Survey Division,
Department of Mineral Resources, Thailand,
Dr. Tetsuo Sugiyama of Fukuoka University
and Dr. Koichi Nagai of Ryukyu University
for their kind cooperation in the field. Our
special thanks to Professor Yanagida whokindly read the manuscript. We would also
like to thank Professor Yuji Okimura ofHiroshima University for his helpful suggestions on the smaller foraminifers. Professor
June R.P. Ross of Western Washington University kindly read this manuscript.
Previous works and geologic setting

In the Lampang district, the Permian formations, generally called the Ratburi Groupin Thailand, are widely distributed. Piyasin(1972) divided the Ratburi Group in this area
into the following three formations in ascending order, namely, the Kiu Lorn, Pha Huat
and Huai Thak. The first two yield Pseudoschwagerina sp. and Neoschwagerina sp., andare correlated with the Lower and Middle
Permian, respectively.
The Huai Thak Formation in the Doi Pha
Phlung area may be divided into lower andupper members. In the lower member, blackshale is predominant with a subordinate
amount of calcareous shale, sandstone andlenticular limestone. The upper member

929

consists mainly of massive to weakly stratified
limestone (the "Doi Pha Phlung Limestone")with black shale. The general strike of the
Huai Thak Formation is almost north tosouth and its dips gently to the east.
Sakagami and Hatta (1982) identified the
Palaeofusulina-Colaniella Fauna from four
limestone blocks collected at the eastern footof Khao Doi Pha Phlung, which may havecome from the "Doi Pha Phlung Limestone".
They compared this fauna to the late Late
Permian Palaeofusulina- Colaniella Fauna
and most probably to the Palaeofusulina
sinensis- Cola n iella parva Fauna of Ishii,
Okimura and Nakazawa (1975).
Later, Waterhouse (1983) reported a richbrachiopod fauna from three localities included in two horizons in the lower part of
the Huai Thak Formation. This brachiopod
fauna is characterized by abundant Oldhamina squamosa Huang, and is closely similar to that in the early Changxingian of South
China, which is assigned to the early Dorashamian.
Recently, Ishibashi and ChongIakmani(1990) reported a small ammonoid faunacomprising Pseudogastrioceras aff. szechuanense Chao and Liang, Paratirolites nakornsrU Ishibashi and Chonglakmoni and
Xenodiscus ? sp. from the shale bed in theuppermost part of the Huai Thak Formation.
They concluded that the geologic age of thisfauna is late Dorashamian.
The fusulinacean fauna characterized byabundant Gallowayinella guidingensis Liu,
Xiao and Dong was found at two localities in
the lower part of the Huai Thak Formation(Figure I). One locality (Loc. HT 1) is at
the left bank of the Huai Mae Phlung river,about 1.5 km north of the summit of Khao
Doi Pha Phlung, and is one of Waterhouse's
brachiopod localities. At this locality, ablack shale member of about 3 m in thickness
crops out, and abundant brachiopods andforaminifers, and sparse trilobites, bryozoans,bivalves and corals were found.
Fusulinaceans were found in the lower part ofKatsumi Ueno and Sumio Sakagami

930

this outcrop.
Another locality (Loc. HT 2) is situated atabout 750 m southwest of Loc. HT 1. It is athin lenticular conglomeratic limestone
exposed on the northern ridge of Khao Doi
Pha Phlung. This limestone is mainly composed of coarse crinoidal and algal remains
with micritic matrix.
The stratigraphic relationship between
these two localities is not confirmed directly
because of the poor exposure. Judging fromthe general dip and strike, however, the former horizon may be slightly lower than thelatter. The latter one is succeeded by the"Doi Pha Phlung Limestone" with a considerable stratigraphic interval.

Faunal affinity and correlation
As shown in Table 1, eight species among

seven genera of fusulinaceans were identified
from Locs. HT 1 and HT 2.
In the fusulinacean fauna, Gallowayinella

Table 1

List of fusulinaceans from thelower member of the Huai Thak
Formation.
SpeciesReichelina cf. changhsingensis
Primoriina sp.
Gallowayinella guidingensis
Palaeofusulina sinensis
P. cf. laxa
Codonofusiella aft'. kwangsiana
fusiformis

HT2+++ ++++++ +++++ ++++Kahlerina? sp.
++++ : abundant, ++ : common, + : rare

Sphaerulina sp.

Gulf af
SiaM

o

HTI

2Km

I

Figure 1. Map showing the fossil localkties.

928.

Permian fusulinacean fauna of North Thailand

guidingensis Liu, Xiao and Dong is abundantat both localities. Reichelina cf. changhsingensis Sheng and Chang is abundant at Loc.
HT I, and common at Loc. HT 2.
Palaeofusulina sinensis Sheng is common at
Loc. HT I, but very rare at Loc. HT 2.
Codonofusiella aff. kwangsiana fusiformis
Sheng is identified from only one specimen
from Loc. HT I. Smaller foraminifers arevery rich at both localities and these assemblages are similar.
Fusulinacean foraminifers are one of themost important and reliable fossils for
examining the geologic age of Permian strata.
Palaeofusulina sinensis was originally described by Sheng (1955) from the Changxing(Changhsing) Limestone of Guizhou (Kueichow), Sichuan (Szechuan) and Jiangxi(Kiangsi) districts in South China. This
species has hitherto been reported from many
localities of the Changxing Limestone or its
eq uivalents in the eastern part of the Tethyan
region, and its biostratigraphic distribution isrestricted to the Changxingian or Dorashamian of the Upper Permian. Palaeofusulina laxa was originally described by
Sheng (1963) also from the Changxing Limestone.
The genus Reichelina flourished in the Late
Permian in the Tethyan region, although its
first appearance is in the late Artinskian(Toriyama, 1971). Reichelina changhsingensis Sheng and Chang, one of the most common species of the genus in the eastern
Tethys, has been reported from the Wujiaping(Wuchiaping) Limestone and Changxing
Limestone in South China (Sheng andChang, 1958 and others).
The biostratigraphic distribution and geologic age of the genus Gallowayinella havebeen discussed by many fusulinacean students
since the middle of this century. Sheng(1955) reported fusulinacean faunas from theUpper Permian Changxing Limestone in
Jiangxi, southern Sichuan and central Guizhou districts, and he stated that the Gallowayinella Horizon is possibly correlated931with that of the Palaeofusulina Horizon,
although the stratigraphic relationshipbetween Gallowayinella and Palaeofusulina is
not known exactly.
Sheng and Lee (1974) showed that thegenus Gallowayinella is one of the main
constituents of the Codonofusiella Zone inSouth China, which is equivalent to the
Wujiapingian of the Upper Permian.
Sheng and Rui (1980) biostratigraphicallyreexamined 22 Gallowayinella localities inseven areas in South China. They concluded
that the Gallowayinella meitienensis Subzone
can be established in the lower part of the
Palaeofusulina Zone, and the genus Gallowayinella is one of the marker fossils whichcharacterize the beginning of the Changxingian.
Lin (1980) discussed the age and stratigraphic significance of the genus Gallowayinellain South China. He established the following three fusulinacean zones in the LopingianStage: Codonofusiella lui, Gallowayinella
meitienensis and Palaeofusulina sinensis
Zones in ascending order. According tohim, the first zone corresponds to the Wujiapingian, while the other two correspond to
the Changxingian. He concluded that thestratigraphic distribution of the genus Gallowayinella is restricted to the lower part of
the Changxing Limestone, and the boundary
between the Changxing Limestone and the
Wujiaping Limestone can be drawn at thebase of the Gallowayinella meitienensis Zone.
Luo and Zhang (1987), on the other hand,emphasized the biostratigraphic importanceof the genus Gallowayinella not only in thelower part of the Palaeofusulina Zone but
also in the upper part of the Codonofusiella
Zone in South China, based on a stratigraphic evidence observed in Shanggao, the Jiangxi district.
As mentioned above, the genus Gallowayinella has been reported from the upper
Wujiapingian and lower Changxingian ofSouth China, although it is common in thelower Changxingian.

932

Katsumi Vena and Sumia Sakagami

928.

Permian fusulinacean fauna of North Thailand

In the Doi Pha Phlung area, Gallowayinella
guidingensis Liu, Xiao and Dong occurs inassociation with Reichelina cf. changhsingensis Sheng and Chang, Palaeofusulina sinensis
Sheng and P. cf. laxa Sheng. This fusulinacean fauna is quite similar to that in thelower Changxingian reported from many
areas in South China and is considered to beof early Changxingian or early Dorashamianage. Consequently, the lower part of the
Huai Thak Formation is correlated with thelower Dorashamian. This conclusion is consistent with that obtained from the brachiopod fauna by Waterhouse (1983).
Systematic paleontology
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea
von Moller, 1878
Family Ozawainellidae Thompson
and Foster, 1937
Subfamily Ozawainellinae Thompson
and Foster, 1937
Genus Reichelina Erk, 1942

Reichelina cf. changhsingensis
Sheng and Chang, 1958
Figures 2-1- 11, 4-3

Compare.Reichelina changhsingensis Sheng and Chang, 1958,
p. 207, 211-212, pI. 1, figs. 1-11 ; Sheng, 1963, p.
27-28,150-151, pI. 1, figs. 23-29; Sheng, 1966, p.
18, pI. 1, fig. 11 (same as pI. 1, fig. 1 of Sheng andChang, 1958, p. 207, 211-212); Lin et al., 1977,
p. 13, pI. 2, fig. 7; Igo and Igo, 1977, p. 93, pI. 13,
figs. 21-24; Liu et al., 1978, p. 14, pI. 1, figs. 6-8(6-7: same as pI. 1, figs. 30, 39 of Sheng, 1963, p.
27-28,150-151); Chen and Yang, 1978, p.22,
pI. 3, figs. 26-27; Sun, 1979, pI. 1, figs. 8-9, 14;933

Rui, 1979, pI. 1, fig. 5; Sakagami, 1980, figs. 2-12, 5-6, 10; Zhao et al., 1981, pI. 4, figs. 1-6 ;Wang et al., 1981, p.15-16, pI. 21, figs. 3-8;
Wang et al., 1982, p. 11, pI. 1, figs. 31-32; Sun etaL, 1983, p.7, pI. 1, figs. 12-13; Igo, 1983, p.
104, pI. 1, figs. 7-8; Sheng and Rui, 1984, p. 33,
pI. 1, figs. 1-2; Chen, 1984, p. 17, pI. 1, fig. 18;
Yang, 1985, pI. 1, figs. 19-20; Yamagiwa and
Hisada, 1986, p. 181-182, pI. 1, figs. 1-15 ;Kobayashi, 1986, p.141, pI. 3, figs. 20-25, 28;
Gu, 1987, pI. 5, figs. 1-3; Zhu, 1989, pI. 5, figs. 78; Zhu, 1990, p.40, pI. 1, fig. 1.
Reichelina aff. changhsingensis Sheng and Chang,
Sakagami and Hatta, 1982, p.5, pI. 1, figs. 1-3.

Material studied. - Axial sections; G K.D
14727b, GK.D 14727c from Loc. HT I and
GK.D 14743 from Loc. HT 2. Tangential
sections; GK.D 14731c, GK.D 14718, GK.D
14725 from Loc. HT I and GK.D 14735 from
Loc. HT 2. Parallel sections; GK. D 14728b
and GK.D 14721 from Loc. HT 1. Diagonal
sections; GK.D 14715, GK.D 14728a from
Loc. HT I and GK.D 14734 from Loc. HT 2.
Additional specimens were also available
from both localities.
Description. - Shell minute, discoidal withangular periphery and straight axis of coiling.
Mature specimens having 4 to 4 Yz volutions,0.19 to 0.22 mm in length and 0.61 to 1.55 mm
in width, giving form ratios of 0.13 to 0.36.
Inner 3 Yz to 4 volutions uniformly coiled,
but the last half one becomes expanded rapidly and forms a cockscomb-like uncoiling part.
It attains a size 2 or 3 times larger than innercoiling part. Radius vectors of the first to
fourth volution of illustrated typical axial
section (Figure 2-4) 0.06,0.14,0.21 and 1.30mm, and form ratios 0.33,0.36,0.38 and 0.08,
respectively.
Proloculus spherical and minute, being
0.03 to 0.04 mm in outside diameter.
Spirotheca thin, composed of a tectum and+- Figure 2. 1-11. Reichelina cf. changhsingensis Sheng and Chang, 3a, 4, 5: axial sections, GK.D
14727b, GK.D 14727c, GK.D 14743, 6,8, 9: tangential sections, GK.D 14735, GK.D 14731c, GK.D 14718,
7,10: parallel sections, GK.D 14728b, GK.D 14721, 1,2, 11: diagonal sections, GK.D 14715, GK.D 14728a,
GK.D 14734, X 40, 3b: enlargement of 3a, X 75. 12-16. Palaeofusulina sinensis Sheng, 12, 14, 15: axial
sections, GK.D 14729a, GK.D 14722, GK.D 14729c, 16: sagittal section, GK.D 14714, 13: tangential
section, GK. D 14729b, X 30. 17: Palaeofusulina cf. laxa Sheng, axial section, GK.D 14720, X 30.

Katsumi Ueno and Sumio Sakagami

934
Table 2.

M.easurements of Reichelina cf. changhsingensis Sheng and Chang.
Diameter of proloculus (in mm).

F.R.: Form ratio, D.P. :Radius vector

Reg. no.

Fig.

2

GK. D 14727b
GK. D 14727c

2-3
2-4

3

GK. D 14743

2-5

1Length

Width

F.R.

D.P.
1

2

3

.4

4Y2

0.15

0.22

0.53

0.21
0.19

1.30
0.39

0.20

0.75

0.27

0.04

0.06

0.19
0.22

1.55
0.61

0.13

0.04

0.06

0.09
0.14

0.36

0.03

0.07

0.11

Form ratio
2

3

4

4Y2
0.09

0.36

0.39

0.40

0.41

2

0.33

0.36

0.38

0.08

3

0.46

0.51

0.46

0.27

probably diaphanotheca. Septa also thinand not fluted. Chomata low and broad,extending to axial regions. Tunnel pathstraight.
Remarks. - The present form closely resembles Reichelina changhsingensis which was
originally described by Sheng and Chang(1958) from the Changxing Limestone on thesouthern slopes of Dameishan (Tameishan)
and Daoduishan (Taotuishan) of South
China, except for having a more expanded
last volution.
Sakagami and Hatta (1982) described
Reichelina aff. changhsingensis Sheng andChang from the limestone blocks at the eastern foot of Khao Doi Pha Phlung. They
stated that Reichelina aff. changhsingensis
Sheng and Chang is seemingly very close to
R. changhsingensis Sheng and Chang in its
general features, except for a more expanded
last volution. The present form is probably
con specific with Reichelina aff. changhsingensis Sheng and Chang described by Sakagami
and Hatta (1982).
Measurements. - See Table 2.
Occurrence.-Abundant in Loc. HT 1 andcommon in Loc. HT 2.
Genus Primoriina Sosnina, 1981
Primoriina sp.

Figures 4-6-7

Material studied. - Axial sections; GK.D
14740 and GK.D 14742 from Loc. HT 2.
Descriptive remarks. - Shell minute andthick lenticular with angular periphery.
Specimen of 4 volutions (Figure 4-6) 0.26
mm in length and 0.37 mm in width, giving aform ratio of 0.70.
The present form is similar to Primoriina
rotunda described by Sosnina (1981) from theMetadoliolina lepida Zone of Sikhote-Alin in
USSR. However, the specific identificationis postponed owing to the insufficiency of Doi
Pha Plung materials.
Occurrence. - Rare in Loc. HT 2.
Family Boultoniidae Skinnerand Wilde, 1954
Genus Gallowayinella Chen, 1934
Gallowayinella Chen, Thompson, 1964, p. C406; Linet al., 1977, p. 43; Wang et al., 1982, p. 30;
Sheng et al., 1988, p. 74-75.
Gallowaiina Chen, 1934, p. 237; Rozovskaya, 1975,
p. 62; Loeb1ich and Tappan, 1988, p.261.
Gallowaiinella Chen, Dunbar and Skinner, 1937, p.
571-572; Dunbar and Henbest, 1942, p. 85 ;Thompson, 1948, p. 45; Sheng, 1955, p. 292 ;Chen, 1956, p. 2, 21 ; Rauser-Chernoussova andFursenko, 1959, p. 211-212; Sheng, 1966, p. 69;
Leven, 1967, p. 131; Liu et aI., 1978, p. 25;

928.

Permian fusulinacean fauna of North Thailand

Chen and Yang, 1978, p.41.
Gallowayina Chen, Thompson, 1964, p. C406 (notGallowayina Ellis, 1932).

Type species. - Gallowaiina meitienensis
Chen, 1934.
Remarks. ~ The generic name Gallowaiina
dedicated to 1.1. Galloway was originallyproposed by Chen (1934). Dunbar andSkinner (1937) proposed the new name Gallowaiinella for Gallowaiina, since the name
Gallowaiina may be construed as an inadvertent error in spelling this surname as a basefor the generic name and is automaticallycorrectable to Gallowayina which has been
preoccupied by Ellis (1932). After that,
Thompson (1964) noted that the generic name
Gallowaiinella is also defective in the same
way as Gallowaiina (ICZN Arts. 32(d) and52(f) and should be emended to Gallowayinella. He also stated that this seemed
to be the most acceptable nomenclatural solution, although he commented that it would benecessary to recognize the validity of Gallowaiina, since it differs by a single letter from
Gallo wayin a Ellis (ICZN Art. 56(b».
Recently, Loeblich and Tappan (1988) restored the generic name to Gallo waiin a, which
Chen was originally proposed in 1934.
In the case of Gallowaiina Chen, 1934,
ICZN Art. 52(f) would be more effective
because it is named in honor of 1.1. Galloway,although both ICZN Arts. 52(f) and 56(b)come into existence. Consequently, thegeneric name Gallowayinella seems to be suitable instead of Gallowaiina.
Up to the present, the following 16 species
and one variety from the Lopingian of South
China have been proposed by the Chinese
students. Among them, nine species arevalid as follows.
Gallowayinella meitienensis Chen, 1934
G. meitienensis var. evoluta Chen, 1934 (= Palaeofusulina evoluta)
G. wutuensis Kuo, 1948 (= Wutuella wUlUensis;
type species of Wutuella Sheng, 1963)
G. minima Chen. 1956 (= Lantschichites minima)
G. decora Lin, 1977

935

G. cylindrica Deng, 1977
G. fusiformis Zhou, 1977
G. megalocula Yang, 1978 (= G. meitienensis Chen)G. cervicalis Yang, 1978 (= G. meitienensis Chen)
G. angulata Yang, 1978
G. guidingensis Liu, Xiao and Dong, 1978
G. quasicylindrica Ding, 1978 (= Ziguiella quasicylindrica; type species of Ziguiella Lin, 1980)
G. laxa Lin, 1980 (= G. guidingensis Liu, Xiao andDong)
G. jiaogensis Zhang, 1981
G. ellipsoidalis Wang, 1982 (= G. meitienensis Chen)
G. histoformis Xie, 1982
G. lepingensis Sheng and Rui, 1984

Geologic age.-Late Wujiapingian to early
Changxingian or late Dzhulfian to earlyDorashamian (Late Permian).
Geographic distribution.-Pamir, USSR
(Leven, 1967), North Thailand (this study),South China (Chen, 1934 and others).
Gallowayinella guidingensis Liu,
Xiao and Dong, 1978
Figures 3-1 - 15, 4-8 - 10
Gallowayinella guidingensis Liu, Xiao and Dong,
1978, p.25-26, pI. 3, figs. 8-9; Lin, 1980, pI. 1,
fig. 2.
Gallowayinella cylindrica Deng, Lin, 1980, pI. 1, fig.
3.
Gallowayinella laxa Lin, 1980, p. 41, pI. 1, figs. 4-5.

Material studied.-Axial sections; GK.D
14731a, GK.D I 4724a, GK.D 14730, GK.D
14723, GK.D 14716, GK.D 14731b, GK.D
14726, GK.D 14711, GK.D 14713 from Loc.
HT I and GK.D 14738, GK.D 14737, GK.D
14741, GK.D 14733 from Loc. HT 2. Sagittal
sections; GK.D 14719, GK.D 14717 from
Loc. HT I and GK.D 14739 from Loc. HT 2.
Slightly oblique axial section; GK.D 14736
from Loc. HT 2. Axial section of immature
specimen; GK.D 14727a from Loc. HT l.
In addition, some other specimens from Locs.
HT I and HT 2 were studied.
Description.-Shelliarge for genus, subcylindrical to elongately fusiform with almost
straight to gently arched median portion andbluntly pointed polar ends. Mature specimens having 6 to 7 volutions, 4.33 to 5.60 mm

936

Katsumi Vena and Sumia Sakagami

928.

Permian fusulinacean fauna of North Thailand

937

3

Figure 4. 1: PalaeoJusulina sinensis Sheng, tangenti a l secti o n, GK.D 14732, X 30. 2: CodonoJusiella afT. kwangsiana Jusiformis Sheng, ax ia l secti on, GK.D 14724b, x 40. 3 : Reichelina cf. changhsingensis
Sheng and C hang, tangenti a l section , GK .D 14725, x 40. 4 : Kahlerina ? sp., sagittal section, GK.D
14712, x 40. 5 : Sphaerulina sp., tangential sectio n, GK.D I4728c, x 20. 6, 7: Primoriina sp., ax ial
sections, GK.D 14740, GK.D 14742, X 75. 8- 10. Gallowayinella guidingensis Liu, Xiao and Dong, 8 :sagittal section, GK.D 14717, 9,10: axial section s, GK.D 14711 , GK.D 147 13, x IS.

in length and 1.10 to 1.50 mm in width. Formratio ranges from 3.41 to 4.36, averaging 3.80
for 26 specimens.
Inner I to 2 volutions tightly coiled, but
outer ones expanded uniformly . Axis ofcoiling straight throughout. Average radius
vectors of the first to seventh volutions, for 10
specimens 0.12,0.17, 0.25, 0.34, 0.46,0.57 and0.69 mm, and form ratios 2.28, 3.10, 3.53, 3.96,
4.08 , 4.02 and 3.62, respectively .
Proloculus commonly spherical , but slightly compressed in some specime ns. Outside
diameter of prolocul us ranges from 0.11 to
0.24 mm , averaging 0.16 mm for 54 specimens.
~

Spirotheca thin , composed of an upper thintectum and lower hyaline layer (diaphanotheca). Average thickness of spirotheca of the first to seventh vol ution for 10
specimens 0.014, 0.014,0.016,0 .020,0.021,
0.021 and 0.0 18 mm , respectively.
Septa intensely and regularly fluted
throughout length of shell exept for central
part of it, attaining to more than 2/ 3 ofchamber height. Septal counts of the first to
fifth volution of illustrated sagittal section(Figure 3-5) 13, 27, 33,39 and 41.
Rudimentary chomata observed only in thefirst volution of so me specimens. Tunnel

Figure 3. 1- 15. Galloway inella guidingensis Liu, Xiao and Dong, 1,2,4,6- 13: axial sectio ns, GK.D
14738, GK.D 14731 a, GK.D 14737, GK.D 14741 , GK.D 14733, GK.D I 4724a , GK.D 14730, GK .D 14723,
GK.D 14716, GK .D 14731b, GK .D 14726, 3,5 : sag ittal sections, GK.D 14739, GK.D 147 19, 14: slightly
oblique axia l section , GK.D 14736, 15 : axial section of immature specimen , GK .D 14727a. All x IS .

Katsumi Ueno and Sumio Sakagami

938

Table 3

Measurements of Gallowayinella guidingensis Liu, Xiao and Dong.
Diameter of proloculus (in mm).

F.R.: Form ratio, D.P. :Radius vector

Fig.

Reg. no.

Length

Width

D.P.

F.R.

1

2

3

4

5

6

7

I

GK.014731a

3-2

4.50

1.13

3.98

0.18

0.13

0.17

0.25

0.34

0.44

0.56

-

2

GK. D 14737

3-4

5.20

1.40

3.71

0.13

0.08

0.12

0.20

0.29

0.41

0.54

0.71

3
4

GK.014733

3-7

4.90

1.23

3.98

0.15

0.11

0.14

0.21

0.28

0.37

0.49

0.65

GK.OI4724a

3-8

4.60

1.20

3.83

0.12

0.12

0.20

0.27

0.38

0.51

0.63

5

GK.014730

4.45

1.18

3.77

0.35

0.45

0.57

4.95

1.28

3.87

0.13
0.12

0.27

GK.014716

0.18
0.13

0.20

6
7

3-9
3-11

0.18

0.27

0.37

0.50

0.65

GK.OI4726

3-\3

5.60

1.40

4.00

0.16

0.32

0.43

0.55

0.70

8
9

4-9

1.35

0.17
0.21

0.57

0.71

0.12

0.17

0.24
0.24

0.45

4-10

3.56
3.85

0.33

GK.014713

4.80
5.20

0.16
0.18

0.23

GK.OI4711

0.11
0.12

0.31

0.44

0.56

-

1.35

Form ratio

Thickness of spiro theca

1

2

3

4

5

1
2

2.24
2.25

2.88
3.17

3.98
3.10

4.03

4.00

3.93

-

3.62

3.85

3.89

3.66 0.010 0.010 0.Dl5 0.015 0.020 0.020 0.020

3

1.45

3.07

3.67

4.79

4.84

4.80

-

4

2.75

3.03

3.48

3.49

3.63

3.68

0.015 0.015 0.020 0.020 0.025

-

5

2.00

2.75

3.37

3.86

4.40

4.23

0.015 0.015 0.010 0.Dl5 0.Dl5

-

6

2.26

2.74

3.22

3.70

4.00

3.85

0.015 0.010 0.020 0.020 0.025 0.020

7

2.91

3.88

3.60

3.98

3.90

4.00

3.86 0.020 0.020 0.020 0.020 0.025 0.025

8

2.50

2.89

3.23

3.76

3.97

3.73

3.33

9

1.78

2.76

3.11

4.03

3.91

4.11

-

6

7

path narrow and almost straight. Axial
fillings present in the first to fourth or fifth
volution.
Remarks. - The present form is quite identical with Gallowayinella guidingensis described by Liu et af. (1978) from the Changxing Limestone in the Guizhou district ofSouth China. In its original description, Liuet af. (1978) have illustrated only two specimens. In the Doi Pha Phlung specimens,
there are some variations in the shell shape,size, form ratio, diameter of proloculus anddensity of axial fillings. They all can be
included in the intraspecific variations.
Gallowayinella guidingensis Liu, Xiao andDong somewhat resembles G. meitienensis
Chen, the type species of this genus from the

1

2

3

4

5

6

0.010 0.010 0.015 0.020 0.020 0.025

7-0.010 0.010 0.010 0.020 0.020 0.015 0.020-

0.Dl5 0.Dl5 0.020 0.025 0.020 0.025 0.015
0.010 0.Dl5 0.015 0.020 0.015 0.Dl5 -Meitian (Meitien) Limestone of Hunan,
South China. However, the former can be
distinguished from the latter in having a moreelongately fusiform shell and larger form
ratio.
The present species resembles Gallowayinella cylindrica originally described by
Deng (in Lin et af., 1977) from the Longtan
Formation of Guangdong. However, theformer has a smaller form ratio than thelatter.
Gallowayinella faxa described by Lin(1980) is considered to be the immature stageof G. guidingensis Liu, Xiao and Dong.
Measurements. - See Table 3.
Occurrence.-Abundant in Locs. HT I and
HT 2.

928.

Permian fusulinacean fauna of North Thailand

Genus Palaeofusulina Deprat, 1912
Palaeofusulina sinensis Sheng, 1955

939

from Loc. HT 1. Additional specimens werealso available from both localities.
Description. - Shell small, bulged fusiformwith almost straight axis of coiling and bluntly pointed polar ends. Mature specimens
having 4 to 4 Yz volutions, 1.85 to 2.10 mm inlength and 1.20 to 1.48 mm in width. Formratio ranges from 1.54 to 1.74, averaging 1.61
for 4 specimens.
The first volution tightly coiled. Beyond
the second volution, shell expands rather
rapidly and becomes loose with growth.
Average radius vectors of the first to fourthand fourth and a half volutions for 3 specimens 0.09, 0.18, 0.36, 0.57 and 0.69 mm, andform ratios 1.70, 1.91, 1.73, 1.10 and 1.39,
respectively.
Proloculus almost spherical, being 0.08 to
0.14 mm in outside diameter, averaging 0.11
mm for 4 specimens.
Spirotheca thin and two-layered, composedof a tectum and diaphanotheca. Very minute fibrous structure observable in a diaphanotheca of some specimens. Average
thickness of spirotheca of the first to fourth,and fourth and a half volutions for 3 specimens 0.018,0.020,0.022,0.015 and 0.020 mm,
respectively.
Septa intensely and narrowly fluted. Septal counts of the first to third volutions of

Figures 2-12-16,4-1

Paiaeofusulina sinensis Sheng, 1955, p. 295-296, 305306, pI. 1, figs. 13, 16, pI. 4, figs. 1-15; Sheng,
1963, p. 50-51,175, pI. 9, figs. 1-9; Sheng, 1966,
p. 67, pI. 9, fig. 3 (same as pI. 4, fig. 2 of Sheng,
1955, p.295-296, 305-306); Zhang and Wang,
1974, p.295, pI. 153, fig. 8; Lin et ai., 1977, p.
43, pI. 9, fig. 4; Liu et ai., 1978, p. 22, pI. 2, fig.
15; Chen and Yang, 1978, p.40, pI. 8, figs. 1-2;
Sun, 1979, pI. 1, figs. 22-23; Rui, 1979, pI. 3, figs.
20-21; Wang et ai., 1981, p. 20-21, pI. 21, figs.
15, 17-18,20; Sakagami and Hatta, 1982, p. 5-7,
pI. 1, figs. 8-22; Wang et aI., 1982, p. 28, pI. 4,
fig. 22; Sun et ai., 1983, p. 20- 21, pI. 4, fig. 17 ;Lin, 1984, p. 158, pI. 9, fig. 30, pI. 10, fig. 7 ;Sheng and Rui, 1984, p.37, pI. 2, figs. 17-19;
Chen, 1984, p. 26, pI. 3, fig. 4; Yang, 1985, pI. 1,
figs. 23-25; Gu, 1987, pI. 6, figs. 1-2; Zhu, 1989,
pI. 8, figs. 12-13; Fan et ai., 1990, pI. 6, figs. 1416.
Paiaeofusulina cf. sinensis Sheng, Sheng and Chang,
1958, p. 207-208,212, pI. 1, figs. 21-25; Zhao etai., 1981, pI. 4, figs. 30-31; Gu, 1987, pI. 6, fig. 3.
? Paiaeofusulina sinensis Sheng, Xie, 1982, p. 17, pI. 7,
fig. 1.

Material studied.-Axial sections; GK.D
14729a, GK.D 14722 and GK.D 14729c.
Sagittal section; GK.D 14714. Tangential
sections; GK.D 14729b and GK.D 14732.
All of the illustrated specimens were obtained

Table 4. Measurements of Paiaeofusulina sinensis Sheng.
pro1oculus (in mm).

F.R.: Form ratio, D.P.: Diameter ofRadius vector

Fig.

Reg. no.

Length

Width

F.R.

D.P.
1

2

3

4

4Yz

1

GK. D 14729a

2-12

2.10

1.2:

1.74

0.14

0.12

0.22

0.41

0.67

2

GK. D 14722

2-14

1.96

1.23

0.10

0.34

0.52

0.69

GK. D 14729c

2-15

1.85

1.20

0.08
0.08

0.17

3

1.59
1.54

0.16

0.34

0.52

0.69

Thickness of spirotheca

Form ratio

1
2
3

1

2

3

4

4Yz

1.83

1.91
1.94
1.88

1.71
1.84
1.65

1.68
1.83
1.60

1.43
1.35

1.38
1.88

0.08

1

2

3

4

4Yz

0.020

0.020
0.020
0.020

0.020
0.025
0.020

0.Ql5
0.Ql5

0.020
0,020

0.015
0.020

-

Katsumi Ueno and Sumio Sakagami

940

illustrated sagittal section (Figure2-16) 10,
20 and 31. Chomata present but inconspicuous. Tunnel path narrow and almost
straight. Axial fillings absent.
Remarks. - The Doi Pha PhI ung specimens
can be fairly assigned to Palaeofusulina
sinensis Sheng, which has been reported frommany localities in the Changxing Limestone
and its equivalents of South China, based on
the important morphological characters.
Measurements. - See Table 4.
Occurrence. - Common in Loc. HT I and
rare in Loc. HT 2.

Palaeofusulina cf. laxa Sheng, 1963
Figure 2-17

Compare.Palaeofusulina laxa Sheng, 1963, p. 54, 179, pI. 11,
figs. 1-9; Li n et at., 1977, p. 42, pI. 8, fig. 23 ;Liu et al., 1978, p. 24, pI. 2, fig. 21 (same as pI. 11,
fig. 2 of Sheng, 1963, p. 54, 179); Chen andYang, 1978, p.38, pI. 7, figs. 8-9; Wang et at.,1982, p. 28, pI. 4, fig. 23; Sun et al., 1983, p. 21,
pI. 4, fig. 19; Zhu, 1989, pI. 8, figs. 9-10.

Material studied. - Axial section; G K.D
14720 from Loc. HT I.
Description. - Shell small, subsphericalwith bluntly pointed polar ends. Length andwidth of shell having 4 volutions, 1.73 mm
and 1.31 mm, with a form ratio of 1.32.
Shell expands rather rapidly. Outer 2 volutions loosely coiled as compared with inner
ones. Radi us vectors of the first to fourth
volution 0.12,0.25,0.44 and 0.71 mm, andform ratios 1.67, 1.56, 1.43 and 1.30, respectively. Prolocul us spherical, being 0.13 mm
in outside diameter. Spirotheca thin, composed of a tectum and rather thick diaphanotheca. Thickness of spirotheca of thefirst to fourth volutions 0.020,0.025,0.030
and 0.025 mm, respectively. Chomata notobserved. Axial fillings absent.
Remarks. - The present species closely
resembles Palaeofusulina laxa originally descri bed by Sheng (1963) from the Changxi ng
Limestone of Guizhou, in the shell shape andrate of expansion. However, the exact identification is postponed until the sufficient materials are accumulated.
Rui and Sheng (19SI) included Palaeofusulina laxa Sheng into the genus Parananlingella which they newly established at thattime. Howevere, the expansion of the outermost volution in Palaeofusulina laxa Shengis not so distinct as compared with Parananlingella acervula Lui and Sheng, the typespecies of the genus Parananlingella. The
former species, laxa, should be left in thegenus Palaeofusulina, to which it was originally referred.
Occurrence. - Rare in Loc. HT I.
Genus Codonofusiella Dunbar
and Skinner, 1937
Codonofusiella aff. kwangsiana
fusiformis Sheng, 1963
Figure 4-2

Compare.Codonofusiella kwangsiana fusiformis Sheng, 1963, p.
45,170, pI. 6, figs. 10-15; Lin et aI., 1977, p. 40,
pI. 8, figs. 9-10 (same as pI. 6, figs. 11, 15 ofSheng, 1963, p.45, 170); Liu et at., 1978, p.27,
pI. 3, fig. 7.

Material studied. - Axial section; GK.D
14724b from Loc. HT 1.
Description. - Shell small, fusiform withbluntly pointed poles with 4 volutions.
Length and width of shell 1.03 mm and 0.49mm, respectively, and giving a form ratio of
2.10. The first volution coiled almost perpendicularly to outer ones. Prolocul us
small, being 0.45 mm in outside diameter.
Radius vectors of the first to fourth volutions
0.05, O.OS, 0.14 and 0.27 mm, and form ratios
1.11,1.40,2.14 and I.S9, respectively. Spirotheca thin, composed of a tectum and diaphanotheca. Septa strongly and regularly
fluted. Chomata distinct.
Remarks. - The present form is similar to
Codonofusiella kwangsiana fusiformis Shengin the shell shape, but differs from the latterin having more fluted septa.

928.

Permian fusulinacean fauna of North Thailand

Occurrence. - Rare in Loc. HT 1.
Family Staffellidae
Miklukho-Maklay, 1949
Genus Sphaerulina Lee, 1934
Sphaerulina sp.
Figure 4-5

Material studied. - Tangential section;
GK.D 14728c from Loc. HT 1.
Remarks. - Greneral shell shape, the spirothecal composition and presence of very finealveolar structure in the present specimen
indicates that it is included in the genus
Sphaerulina, although it is not properly oriented.
Occurrence. - Rare in Loc. HT 1.
? Family Verbeekinidae Staff
and Wedekind, 1910
Subfamily Kahlerininae Leven, 1963
Genus Kahlerina Kochansky-Devide
and Ramovs, 1955
Kahlerina? sp.
Figure 4-4

Material studied.-Sagittal section; GK.D
14712 from Loc. HT 1.
Remarks. - The generic assignment of thepresent specimen is tentative, because only
one sagittal section is insufficient for study.
Occurrence. - Rare in Loc. HT 1.
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Katsumi Ueno and Sumio Sakagami

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p. 181-189, pis. 5-14. Geo!. Publ. House, Beijing. (in Chinese with English abstract anddescription of new species)

Changxing (Changhsing) :lltJi!, Dameishan (Tameishan) ::ktltlll, Daoduishan (Taotuishan)fffi:Jt III , Guangdong $Jlt Guizhou (Kueichow) iV'I'!, Jiangxi (Kiangxi) iIg§, Hunan i'Iifl
1¥i, Longtan iEi'J., Meitian (Meitien) tialIl, Shanggao J::ii1ii. Sichuan (Szechuan) ~)II.
Wujiaping (Wuchiaping) ~*W.
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Z'~t±lLt..:o ;cO)?i?fiO)"'-'5EII~-e~t..:~o)fiReichelina cf. changhsingensis Sheng andChang, Gallowayinella guidingensis Liu, Xiao and Dong, Palaeofusulina sinensis Sheng, P.
cf. laxa Sheng, Codonofusiella aff. kwangsiana fusiformis Sheng 0) 5 fi-c: ih 9 0 ~ 0)il!15~£R
ff~f;t Gallowayinella guidingensis Liu, Xiao and Dong Z'~l1:fi C L, <J:tW1¥i$O):lltJi!::fi
WZ!fi:to J: Lf;cO) "'-'lIi'ft~}jf;::J! 6;/'t9il!15~£Rff~f;::~{j;J,"" 9 0 ;cO)IIi'f~fHtrWl Changxingianih9~'f;tlltrWl Dorashamian C~*-6;/'t90
J:lIlJJm~' :!tl[J::m~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, pp.944-963, 13 Figs., December 30, 1991

929.

PERMINERALIZED OTOZAMITES LEAVES (BENNETTITALES)FROM THE UPPER CRETACEOUS OF HOKKAIDO, JAPAN*TAMIKO OHANA and TATSUAKI KIMURA
Institute of Natural History, 24-14-3 Takada,Toshima-ku, Tokyo 171

Abstract. The present paper deals with the description of external and internal featuresof two types of permineralized Otozamites leaves. We establish Otozamites kerae, sp. nov.
and 0. takahashii, sp. nov. to accommodate the present leaves. The former is characterized
by small-sized leaf and pinnae with denser veins, and by the presence of roofing cells overthe stomata and the absence of papillae and trichomes on both sides of cuticles. The latteris characterized also by small-sized leaf and pinnae with less dense veins, and by the presenceof roofing cells over the stomata and of ramen tum-like trichomes on the lower cuticle.
Well-preserved leaf cuticles of both species and histological features of the former species aredescribed in detail. The occurrence of both species is one of the youngest records of
Otozamites in the Mesozoic.
Key words.

Permineralized Otozamites, Upper Cretaceous, Hokkaido (Japan)

the Triassic, Jurassic and Early Cretaceous
plant sites, but are rare in the Late Cretaceous
plant sites. The occurrence of both species is
one of the youngest records of the genus
Otozamites in the Mesozoic.
In order to make a histological study of the
Kumaoizawa and Kamimakizawa specimens,
they were sliced along various surfaces with adiamond saw with a 0.4 mm thick blade.
Then cellulose acetate peels were made.
Cuticular preparations were made from all
specimens at hand.
The examined specimens and slides (peels
and cuticular preparations) for Otozamites
kerae and cuticular preparations for O. takahashii are kept in the Institute of Natural
History, Tokyo.

Introduction
The Upper Yezo Group of marine origin(mostly Coniacian-Santonian in age) is distributed widely in the median part of Hokkaido and yields various fossil plants in thecalcareous (often ammonite-bearing) nodules
it contains. Although they are represented
mostly by tiny fragments, most of them arepermineralized by calci urn carbonate andtherefore their internal details are rather wellpreserved.
This paper gives the description of two new
species of permineralized Otozamites leaves
collected by Messrs. Yasuj i Kera, Masatoshi
Kera, Ken'ichi Saiki and Hiroshi Takahashi
from the Upper Yezo Group exposed at theKumaoizawa and Kamimakizawa Valleys
and Nakakinenbetsu (Obira) Valley (FigureI). They are all fragmental, but show
unique external and internal features. Otozamites species are abundant and varied in

Acknowledgements
We thank Messrs. Yasuji Kera, Masatoshi
Kera, Ken'ichi Saiki and Hiroshi Takahashi
who collected and provided the examined
specimens for the present study. Our thanks*Received July 5, 1991; revised manuscript accepted
September 17, 1991
944

929.

Permineralized Otozamites leaves

are extended to Dr. Shya Chitaley of theCleveland Museum of Natural History, Ohiofor going over the language of the present
manuscript.
This study was supported in part by the
Grant-in-Aid for Scientific Research from the
Ministry of Education, Science and Cultureof the government of Japan (nos. 6340623 and02640615).

Description
Class Cycadopsida
Order Bennettitales
Genus Otozamites Braunin M iinster, 1842
Otozamites kerae Ohana et Kimura,sp. nov.
Figures 2-7

Material.-Holotype; INH-IA (collectedby Y. Kera from the Kamimakizawa at talus).
Other specimens examined; INH-lB (counterpart of INH-IA), INH-2 (collected by M.
Kera from the Kamimakizawa at talus) andINH-3 (by K. Saiki from the Kumaoizawa
from rolling rock). Stratum typicum:
Upper Yezo Group (mostly ConiacianSantonian in age). Locus typicus: Middle
course of the Kamimakizawa (roughly 142'08'
05"E, 43'07' 43"N; an eastern branch of the
Shuparo River), Kashima, Yubari City.
Derivatio nominis: After Y asuj i Kera, whocollected the holotype. Occurrence: Rather
rare.
Description. -1. External features (Figures 2A-E): The piece of leaf is pinnate withthick rachis. Its petiole is not preserved.
Judging from the leaf fragments obtained, theleaf is small, long and narrow, with nearlyparallel margins for the most part. It is up to
2.2 cm across in its widest part, narrowing
gradually towards both ends. The pinnae
are alternate, set closely, attached to uppersurface of rachis at an angle of 40 degrees.
They are oblong or nearly rectangular inform, typically 1.2 cm long and 4.5 mm wide945

at middle; acroscopic basal part is markedly
expanded to form a rounded angle (auricle),
but the basiscopic basal margin is usuallyconcealed by the rounded part of the pinna of
the opposite side. Margins are entire withbroadly rounded apex. The veins are densely crowded, radiated from the basiscopic
basal part and are dichotomously forked
repeatedly at all levels; the density is 40-42
per cm at the middle of pinna. Reproductive
organs are not known.
2. Internal features: 1) Rachis (Figures
3A-J, 5A-H, 6A-D): In cross section, therachis is obpentagonal in form (Figures 3AJ), 2.6 mm X 2.4 mm thick. It shows epidermal and hypodermal layers, ground tissue
and vascular bundles. Epidermal layer (Figures 5C, H) is one cell thick. The cells arerectangular in cross section, 8-10 pm thickadaxially and 15 pm thick abaxially. Cuticle is obscure in cross section.
Hypodermal layer (Figures 5C, E, F, H) is
6-8 cells thick (120-140 pm thick) in crosssection. Hypodermal cells are thick-walled
and sclerenchymatous, but those of adaxial
side are rather thin-walled (Figure 5H).
Ground tissue consists mostly of large,
thin-walled parenchymatous cells. They areround or oval in form (Figures 58-H, 68) incross section, (20)-50-(90) pm in diameter,
and with small intercellular spaces in
between. A few rather thick-walled cellswith cellular contents, isolated or grouped[Cc in Figure 5C; secretary cells or idioblast; cf. Rao and Achuthan, 1968, text-fig.
9], 40-50 pm in diameter, are scattered in the
ground tissue (Figures 58-H, 68).
Vascular bundles are arranged in a double
series in the form of a 'U' (Figures 3A-J, 5A,6A); the concavity of the 'U' faces upward
and the bundles vary from 12 to 13 in number
separating from each other laterally. Someof the bundles are in a state of incomplete
division. The bundles are collateral, endarch, with poorly preserved phloem (Figures
58-H) and well-preserved xylem with spiraland scalariform tracheids (Figures 6C-D).

Taml"ko Ohana an d Tatsua kl" Kimura

946

1A

100km,The Japan
Sea-:2't The Pacific
)

/

1C

929.

Permineralized Otozamites leaves

The xylem in the outer and inner sides of the'u' faces in opposite direction.
Thick-walled sclerenchymatous cells(mechanical tissue or fibres?) are present
outside and inside the phloem (Figures 5BH). They do not extend to the space between
the adjacent bundles. These sclerenchymatous cells are less developed in the
inner bundles of the 'u' (Figure 50).
2) Pinnae (Figures 4, 7 A-L): Structurally the pinna consists of a single cell layer ofupper epidermis with well-preserved cuticle, asingle cell layer of upper hypodermis consist-20

ing of thick-walled cells followed by one or
two layers of vertically elongated thin-walled
palisade cells, spongy tissue with large intercellular spaces in between , a single cell layerof lower hypodermis and a single cell layer oflower epidermis with well-preserved cuticle.
Vascular bundles (veins) are located in a row
in the spongy tissue.
The pinna is 0.3- 0.36 mm in vertical thickness (Figures 4, 7A-C, G) and is somewhat
thicker along the margins. The pinna margin is usually broadly rounded in sectional
view.

2A

1em

2C
2em

1
28

94 7

1 em

1cm

r\

~­
~.

//2E

Figure 2. OlOzal11iles kerae, sp. nov. 2A. A leaffragment (holotype; enlarged twice). Its upper partis buried in the rock matrix. INH-I A (collected by Y. Kera from the Kamimakizawa). 2B. Counterpartof the holotype. The arrow indicates an associated ammonite. Two median ridges in this figure cover thelateral ridges of the concealed rachis. 2e. Imbricated pinnae with marked acroscopic basal angles (auricles)
drawn partly from the holotype. Both dotted lines indicate real position of the lateral edges of a concealedrachis. 2D. Venation of a pinna drawn from the holotype. 2E. Venation of small pinnae from INH-3(collected by K. Saiki from the Kumaoi zawa).
~
Figure I. Localities of Olozal11ileS leaves described in this paper. IA. A median region of Hokkaido
Island , showing the exposed area s of Upper Cretaceous sediments (dotted areas) and three localities of
OlOzal11iles leaves. Nk ; Nakakinenbetsu Valley (Tappu , Obira-cho), Ku ; Kumaoizawa Valley, Km;
Kamimakizawa Valley. lB. Detailed plant sites of Kumaoizawa (Ku , roughly at long. 142' 03'OO" E, lat. 43'
14'27"N) and Kamimakizawa (Km). Ie. Detailed plant site of Nakakinenbetsu (Tappu, Obira-cho) (Nk).

948

Smm

Tamiko Ghana and Tatsuaki Kimura

929.

Permineralized Olozamiles leaves

949~ 3C-b y
\ i

x

3D-b

5mm

4
200l-lm

I
Figure 3 (left and top) and Figure 4. Otozamiles kerae, sp. nov. 3A - J. Serial cross sections of the buriedrachis with pinnae (holotype) cut and photographed successively, 0.1 mm apart, in ascending order from Figure 3A
to Figure 3J. They show the arrangement of vascu lar bundles (double series in the form of a ' U' ) and the modeof vascular supply to a pinna (details, see text). Figures 3B- b, 3C- b, 3D- b, 3E- b, 3F- b and 3G-b are drawn from
Figures 38-a, 3C-a, 3D- a, 3E- a, 3F - a and 3G- a, respectively. Figure 4. A cross section of a pinna (holotype)enlarged.

---

Tamiko Ghana and Tatsuaki Kimura

950

50

.---t----,...-5C

-----

5E

L------'-5H

Figure 5. OlOzamilas kerae, sp. nov. SA. A cross section of the buried rachis made between those of
Figure 3D and Figure 3E, indicating the areas of the corresponding detailed micrographs (Figures 5B- F, H).
In each vascular bundle, black part shows xylem and the other part shows phloem with thick-walledsclerenchymatous cells (mechanical tissue or fibres ?). 5B. Adaxial vascular bundles in the left side of the
double ' U'-shape series. Two terminal bundles (x and y) are about to unite with each other. Sc. Adaxial
side of the rachis. It shows poorly preserved epidermal cell layer (Ep) , thick hypodermal layer (Hy) , groundtissue (Gt) and cells with cellular contents (secretary cells or idioblast ; ec). SO. Vascular bundles on theinside of the ' U' . In each phloem part, thick-walled sclerenchymatous cells (mechanical tissue or fibres?)are less developed than in each phloem part of the outer bundles of the ' U'. SE. Vascular bundles on theoutside of the ' U'. SF. Vascular bundles on the abaxial side of the outside of the ' U'. SG. Drawn partly
from Figure SF. Xy; xylem, Ph; phloem, Twc; thick-walled sclerenchymatous cell (mechanical tissue or
fibres ?). SH. Abaxial side of the rachis. The cells of hypodermal layer are rather thin-walled.

929.

Permineralized Otozamites leaves

95 1

6A
a

b

2.5mm i

\\
c'
250~m

a,Hy,

Gt

,Ph, Xy,

Gt

Ph

Xy

Gt

Xy

P~b

,d

60

Figure 6. OlOzamiles kerae, sp. nov. 6A. C ross sect io n of the buried rachis w ith pinn ae, e nl a rged
fro m Figu re 3G. Showing the a rea of the detailed microg raph (Figu re 6B) a nd the ve rti ca ll y c ut lin es (p q and r- s) . 6B. Pinna-trace (Tr) origin a ted from a united bund le at the termin a l position o f the left sideof the ' U' (see Figu res 3F- b and 3G - b) (in both Figu res 6A a nd 6B, th e ri ght side is really th e upper side).
6C. A verti cal secti o n of the rachis c ut a long the p- q lin e in F igure 6A between a a nd b. H y ; hypodermis ,Gt; ground ti ss ue, Ph; phl oem, Xy; xy lem. 6D. A ve rtica l secti on of the rac hi s cut along th e r- s lin e in
F igur e 6A between c a nd d . In thi s micrograph , the left-side xy lem is made up of sp ira l trac he ids (primary
xy lem part ?) a nd the right-sid e xy lem of sca lar iform tracheids (secondary xy lem part ?) .

952

Tamiko Ghana and Tatsuaki Kimura

Cells of the upper epidermis are rectangularin cross section, 25,um long and 7.5,umhigh, but they are larger and squarish aroundthe marginal region. In surface view, they
are rectangular, 50-85,um long and 30,um
wide, with marked sinuous anticlinal walls;
the sinuosity is 15,um long (wave-length) and20,um wide (amplitude) in anticlinal walls,
and is lO,um long and lO,um wide in end
walls (Figure 7F).
In our observation of upper cuticle, it consists only of ordinary cells, and no stoma,trichome or papilla have been found on theouter periclinal walls.
Cells in the upper and lower layers of
hypodermis are polygonal and thick-walled;
their lumina are often markedly reduced involume. The lower layer of hypodermis isinterrupted at the stomatal region (Figures
7A-C, E).
The palisade cells are well developed, 95125,um long and 20-25,um wide in crosssection, but the area above a vascular bundle
is devoid of palisade cells (Figures 7A-D).
The spongy cells are irregular in form andlarger, 30-50,um in diameter, but the area
below the vascular bundle is usually devoidof spongy cells (Figures 7A-E).
In our observation of cross section of pinnae and their lower cuticle, stomata are distributed between vein courses by a single orsometimes 2-3 rows. Cells on the veincourses are elongate-rectangular, with lesssinuous walls, and those between the stomatalrows are isodiametric with less sinuous walls.
The marginal zone of pinna, ca. 250 ,urn wide,
is devoid of stoma (Figure 71).
Stomata are syndetocheiric. Guard cells
are sunken and each cell is 50,um long and 12,urn wide, and strongly cutinized; aperture istypically 17,u m long and transversely oriented to the veins (Figure 7H). Subsidiarycell is 20,um wide and its walls are not sinuous but straight (Figures 7J-K).
Stomatal apparatus is also sunken in thebottom of a large oval pit formed by surrounding 8-10 epidermal cells. In addition,these epidermal cells form a dome-shapedroof, 80-100,um long, 70-90,um wide and up
to 35 ,urn high from the surface of guard cell,and with a rounded or stellate mouth. Roof
cells are somewhat thickened toward themouth which does not lie just above thestoma, but at the end toward the rachis, while
the stoma is at distal end to the rachis (Figures 7H-K). No papilla has been found on
the ordinary cells except for the roof cells,
and no trichome or trichome base has beenfound on the epidermal cells (Figures 7H-I).
In cross section, there is a large chamber
inside each stoma (Figures7A-C, E). In
longitudinal cross section of a pinna, thevascular bundles are 3\-35 in number (thesame as the number of veins), each of which islocated at the lower part of spongy tissue andconsists of small masses of adaxial xylem andpoorly preserved abaxial phloem. They are
irregularly surrounded by a parenchymatous
sheeth consisting of large cells (Figures 7AE).

Between the adaxial hypodermis and vascular bundle (vein), there develops a wedgeshaped mechanical tissue consisting of largesclerenchymatous cells in 2-4 horizontal rows
(Figures 7A-D); the uppermost row consistsof 4-5 cells and the lowermost row of 1-2
cells. These mechanical tissue cells are notseen in cuticular preparations. Figure 7L
shows inside view of a stomatal apparatus(SEM micrograph).
3) Probable course of vascular supply tothe pinna: Figures 3A-H and 5B show serial
cross section of the rachis and obliquely cut
three pinnae buried in the rock matrix (thelower extension of Figure 2A) for I mm inlength of the rachis. In Figures 3B (a-b) and3C (a-b), at first two bundles (x and y) fromthe left adaxial corner come to be elongatedvertically and are constricted in the middle.
Then upper portions of both bundles come incontact and unite with each other [Figures
3D (a-b), 5B] to form a single bundle which
is separate from lower parts of the constriction [Figures 3E (a-b), 3F (a-b)]. A pinna

929.

Permineralized Otozamites leaves

Figure 7. Olozamiles kerae, sp. nov. 7A. A longitudinal section of a buried pinna (holotype) cut
nearly a long its median portion (enlarged). 78. Drawn from Figure 7A. It shows upper and lower
epidermal layers (UEp and LEp), upper and lower hypodermal layers (UHy and LH y), palisade and spongy
tissues (Pa and Sp) , vascu lar bundle (vein ; Vb), wedge-shaped mechanical tissue (Mt), stomata l chamber(Stc), guard cells (Gc) , stomatal pit (Stp) and dome formed by roof cell s (Rc) with mouth. In Figures 78
and 7C, shaded areas show the cell walls. 7C. A longitudin al section of a buried pinna (holotype). The
marginal region is rather thickened compared to Figure 7A. 7D. Wedge-shaped mechanical ti ssue (Mt)made up of thick-walled and large cell s extending upward from the vascu lar bundle (vein; Vb). 7E.
Abaxial part of a pinna (longitudinal section) en larged partly from Figure 4. It shows two large-spaced
stomatal chambers (Stc), guard cells (Gc), stomata l pit (Stp) and a dome formed by roof cells (Rc) with amouth (Mo). 7F. Enlarged upper cuticle made up on ly of ordinary cell s. Their long axis is in parallel tothe veins. The anticlinal and end wa ll s are markedly sinuous and thickened and with rounded loops (often
mushroom-shaped) reaching nearly to the middle of the cell. Cells o n the vein courses are not specia li zed.

953

Tamiko Ghana and Tatsuaki Kimura

954

7G
I

100\-lm
2mm

L -______________

~I

7K

7H
200\-lm

200\-lm[
100!-,m929.

Permineralized Otozamites leaves

receives this single bundle [Figures 3F (a-b),
3G (a-b), 6BJ at the pinna base, but unfortunately, details of the mode of branching of
this single bundle at the very base of pinna
are not visible.
Discussion and comparison. - Externally,
the present Otozamites leaves are small, longand narrow having comparatively thick rachis and falcate, closely set, imbricate andoblong or nearly rectangular pinnae withbroadly rounded apex and densely crowded
and radiated veins.
A number of Otozamites species have been
described from Mesozoic plant beds.
Among them, we could not find the leaves
externally similar in habit to the present ones.
Therefore, even on the basis of external formof the present leaves, it would be reasonable
to create Otozamites kerae, sp. nov. to accommodate the present leaves.
Apart from the present specimens, internalfeatures of the bennettitalean leaf-rachises or
petioles have been described for the following
taxa:
Pti/ophyllllm amarjo/ense Bose: Bose, 1953; Boseand Kasat, 1972a.
P. clIlchense Morris: Bancroft, 1913; Rao andAchuthan, 1967.
P. gllplai Sharma: Sharma, 1967.
P. sahnii Gupta et Sharma: Bose and Kasat, 1972a.
Diclyozamites fa/ca/us (Morris) Medlicott et Blanford: Bancroft, 1913; Bose and Zeba-Bano, 1978.
D. indictls Feistmantel: Bose and Zeba-Bano, 1978.
Dictyozamites sp.: Bose and Kasat, I 972b.
Williamson ian-type stems: Bancroft, 1913.

The arrangenent of vascular bundles In
Ptilophyllum rachises is a double series of

955'U'-shape (Rao and Achuthan, 1967; Boseand Kasat, 1972a), but the number of bundles
is larger (20-32 in Ptilophyllum amaljolense,
23-32 in P. sahnii, 25-32 in P. cutchense)than that of the present Otozamites kerae.
In Ptilophyllum guptai, pinna trace-bundles
are two in number at their origin (Sharma,1967), instead of a united one as in the present
Otozamites kerae. While, in Ptilophyllum
cutchense the number of trace-bundle is single at its origin (Rao and Achuthan, 1967).
According to Bose (1953) and Bose andKasat (l972a), in the petiole of Ptilophyllum
amarjolense, the vascular bundles are I I in
number (5 abaxial, 2 lateral and 4 adaxial)
and arranged in a reniform manner. Unfortunately the petiole of Otozamites kerae is
not known.
In the rachis of Dictyozamites falcatus, thevascular bundles are 25-30 in number and arealso arranged in a double 'U'-shaped series,
surrounded by an incomplete layer of thickwalled mechanical tissue (Bose and ZebaBano, 1978).
As mentioned above, the arrangement ofvascular bundles in the rachises of the bennettitalean genera, Ptilophyllum, Dictyozamites and the present Otozamites is similar, but the latter differs from the anatomically known rachises of Ptilophyllum and Dictyozamites in having small number of vascularbundles and less developed mechanical tissue
surrounding them.
The mechanical tissue of pinna in association with vascular bundle (vein) is unique in
Otozamites kerae, and is considered to begood for reinforcement of the weakness of the+--- Figure 7. Otozamites kerae, sp. nov. 7G. An oblique longitudinal section of a pinna (holotype).
The rachis is located under the left edge. 7H. Stomatal rows between adjacent veins (lower cuticle;
holotype). Each stoma is covered with a dome formed by roof cells of which the mouth is situated at thelower side of this micrograph (toward the rachis-side). Apertures are directed transversely or sometimes
somewhat obliquely. Guard cells are invisible, because they are sunken. A pair of dark parts showsheavily cutinized subsidiary cells. Anticlinal and end walls of the ordinary cells are less sinuous than thoseof upper ordinary cells. 71. Lower cuticle (holotype). The arrow shows non stomatal marginal zone
where the sinuosity of ordinary cells is nearly the same as that of upper ordinary cells. 7J. Three stomata
with their own domes formed by roof cells of which mouths are stellate. S; subsidiary cell, Rc; roof-cell,
Mo; mouth of a dome. 7K. Two stomata with the same dome as shown in Figure 7J. 7L. Inner view ofa stoma (SEM-micrograph). A; aperture, Gc; guard cell, Sc; stomatal chamber.

956

Tamiko Ghana and Tatsuaki Kimura

1cm

1

/
Be
1cm

1cm
I
Figure S. Otozamiles takahashii, sp. nov. SA. A broken leaf (holotype, INH - 4A indicated by 'a'(surface view)] collected by Hiroshi T akah as hi at the Nak ak inenbetsu (Obira) locality (Figure IC) inassociation with an ammonite indicated by the arrow. S8. A sketch of the holotype, showing welldeveloped auricles concea lin g the rachis entirel y. Dotted lines indicate position of rachis. Sc. A part ofa leaf enlarged from the holotype, showing the ve nation . SD. The a uricle of a pinna (indicated by 'X' )stripped away to show the o rigin of rad iated veins [ INH - 48 indicated by ' b' in Figure SA (wrong-sideview) ].

929.

10

Permineralized Otozamites leaves957

20 0 fJ m

100fJm~\.,----.,v~--~'---v--

Mar

Mr

Pm

Uc

Figure 9 a nd Figure 10. OLOzal17ileS takahashii, sp. nov. 9A. Upper c uticle co nsis ting o nl y of theord in ary ce ll s with sinu ous antic lin a l a nd end wa ll s (holotype). 9B. Sinuou s wa ll s of upper ordinary ce ll s
enlarged part ly from Figure 9A. Periclinal wa ll s are a ll smooth in surface view . Figure 10. Marginal partof a pinna. Upper a nd lower c utic les were unfolded on the same plane (Ma r ; main regio n of the lower
c ut icle w ith stom a ta , Mr ; margina l reg ion of the lowe r c uticle consisting on ly of ord in ary ce ll s, Pm ; partof pinna marg in , Uc ; upper cu ti c le) (holo type).

958

Tamiko Ghana and Tatsuaki Kimura

929.

Permineralized Otozamites leaves

pinna architecture due to the existence of alarge space (stomatal chamber) inside eachstoma (Figures 4, 7 A-C, E).
Stomatal roofing cells (or roof cells) similarin structure to those of Otozamites kerae
were described by Harris (1969) for O. beani(Lindley et Hutton), O. graphieus (Leckenby) and O. leekenbyi Harris, and by Barale(1987) for O. pterophylloides Schimper.
Such roofing cells are thought to be unique tosome of the Otozamites species hitherto described, and have not been recorded in theleaves of other bennettitalean genera.
A stellate or oval-shaped mouth of a
stomatal pit formed by 8 or more outermonocyclic or dicyclic encircling cells has
been recorded in other gymnospermous genera, like Tieoa, Ruflorinia and Mesosingeria(Archangelsky, 1963) and Otozamites parviaurieulata Menendez (Menendez, 1966) andother cycadalean genera (e.g. Harris, 1964).
But each mouth in those cases unexceptionally opens just above a stomatal aperture.
Diagnosis. - Leaf small, long and narrow,nearly parallel-sided for the most part, up to
2.2 cm wide. Pinnae alternate, closely set,
imbricate, oval or rectangular in form, typically 1.2 cm long and 4.5 mm wide, withmarkedly auriculated acroscopic base andbroadly rounded apex, and attached to uppersurface of rather thick rachis. Veins densely
crowded, 40-42 per cm in density, and radiated.
Rachis obpentagonal in cross section, 2.6
mm wide and 2.4 mm high, being made up ofan epidermal layer of one cell thick, ahypodermal layer 6-8 thick-walled cells thick(120-140 J.1.m thick), parenchymatous groundtissue, vascular bundles, 12-13 in number,arranged in a double 'U'-shaped series.

959

Vascular bundle made up of spiral and scalariform tracheids and phloem with thick-walledsclerenchymatous cells (mechanical tissue or
fibre ?). Xylem in the outer and inner sidesof the 'U' facing in opposite directions.
Thin-walled cells with dark contents are scattered in the ground tissue (secretary cells or
idioblast). (Cuticle of the rachis poorly
preserved.) Pinnae 0.3-0.36 mm thick, upperand lower epidermal layers one cell thick,upper and lower hypodermal layers one cell
thick, palisade and spongy tissues present,
vascular bundles (veins) and wedge-shaped
mechanical tissues, each extending above eachbundle upward. Epidermis (cuticle) hypostomatic. Upper epidermis rectangular, consisting of uniform ordinary cells with markedly sinuous walls. Lower epidermis consisting of main part and nonstomatal marginalzone. In the main part, stomata distributed
between adjacent veins In 1-3 rows.
Stomata syndetocheiric. Guard cells sunken,
stomatal pit covered by roof cells originatedfrom encircling cells. The mouth of a dome
formed by roof ce II s oval or stellate, locatedon the rachis side. Stomatal aperture transverse or sometimes oblique to the vein course.
Trichomes and papillae absent on both epidermis. (Petiole, stem and reproductive
organs not known.) Registered slides (Allfrom the holotype): INH-1-C-13 (Figure
3A), C-14 (Figure 3B), C-16 (Figure 3C), C18 (Figure 3D), C-20 (Figure 3E), C-25 (Figures 5B-F, H), C-28 (Figures 3F, 7 A, C), C30 (Figure 3G), C-32 (Figure 3H), C-33 (Figure 31), C-34 (Figures 3J, 4, 6B, 7E), C-35
(Figures 7B, D), R-6 (Figure 6C), R-II (Figure 60), L-4 (Figure 7G), Cu-I (Figure 71),
Cu-2 (Figures 7H, J, K), Cu-3 (Figure 7F),
Cu-4 (Figure 7L).

~
Figure 11. Otozamites takahashii. sp. nov. ttA. Main region 01' the lower cuticle (holotype),
showing irregularly disposed stomata with covering dome (or sac)-shaped roof cells and ramentum-liketrichomes. The rachis is located at the left hand of this figure. 11B. A sketch of the lower cuticle(holotype), showing most of main region with stomata and trichomes (Tr) and a part of marginal rim (orzone) (Mr). Open circle indicates the roof with a mouth (dotted open circle) located unexceptionally at its
left hand corner. Solid dash line shows position and direction of stomatal aperture. In this figure, general
direction of veins is in parallel to the x-y line and the rachis is located on the left.

960

Tamiko Ohana and Tatsuaki Kimura

128
12A

~----------------I1100fJm

13A

138

929.

Permineralized Otozamites leaves

Otozamites takahashii Ohana
et Kimura, sp. nov.
Figures 8-13

Materials. - Holotype; INH-4, with counterparts (collected by H. Takahashi at talus)
Stratum typicum: Upper Yezo Group (mostly Coniacian-Santonian in age). Locus
typicus: Lower course of the Nakakinenbetsu Valley (a tributary of the ObirashibeRiver) (roughly 141055'51''E, 44°03'30"N),
Tappu, Obira-cho, Rumoe-gun, Hokkaido.
Derivatio nominis. - After Hiroshi Takahashi, who collected the holotype.
Occurrence. - Rare (at present).
Description. - Two tiny leaf fragments wereobtained. They are preserved on the samerock slab with an ammonite as shown in
Figure 8A.
1. External features (Figure 8A-D) :Leaves are nearly parallel-sided, 2 cm wide
but whole length is unknown. Pinnae areclosely set, alternate, imbricate and attached
to upper surface of rachis at an angle of 55
degrees. Pinnae are generally rectangular inform, typically 1.25 cm long and nearly parallel-sided for the most part, 3.5 mm wide andslightly falcate; acroscopic margin is nearlystraight throughout and apical basiscopic
margin is rounded; apex is obtusely pointed
or rounded. Acroscopic angle is remarkably
expanded to form an auricle which conceals
the rachis entirely. Veins are delicate, densely crowded, originate from middle of base(Figure 8C), then radiated and forked dichotomously at all levels, and end with themargins; the density is 40 per cm at the
middle of pinna.

961

2. Cuticular features: Cuticle is hypostomatic, fairly sturdy and hard to fold in
preparation; therefore, it is difficult to know
its thickness.
Upper cuticle consists only of ordinary
cells (Figures 9A, B) which are typically rectangular, 25-65,um long and 20-30,um widewith sinuous anticlinal (lateral) and end
walls. The sinuosity of anticlinal walls istypically 15,um long (wave-length) and 10,urn wide (amplitude), and that of end wall isless marked. Peripheral walls are smoothand no papillae and trichomes are recognized.
Lower cuticle is divided into a marginalrim (Figure 10) and main region (Figures
IIA, B). Both vein course and interveinal
region are less marked. The marginal rim
consists of ordinary cells which are different
in size and sinuosity from those of upper
cuticle. They are irregular in size, 40-150,urn long and 15-30,um wide, and their sinuosity is also irregular and less developed.
The main region consists of ordinary cells
and stomata (Figures 10, II A, B). The ordinary cells are irregular, 20-80,u m long and
20-50,um wide, rectangular, square to nearly
triangular in form; their sinuosity is lessdeveloped than that of upper ordinary cells.
The stomatal complex is syndetocheiric.
The stomata are densely crowded, but do notappear to form stomatal files and bands;
their density is 70-90 per square mm. In a
stoma, guard cells and subsidiary cells are
sunken in a stomatal pit with surrounding
ordinary cells. Guard cells are 40-50,um
long and thickly cutinized on the inside andsubsidiary cells are rather small, each 50-60,urn long and 15,um wide, and less cutinized.

~
Figure 12 and Figure 13. Otozamites takahashii, sp. nov. 12A. A roof (or sac) consisting of roof
cells originated from encircling cells surrounding a stoma. The mouth (Mo) is stellate in form. The roof
cells near the mouth are generally much cutinized (holotype). 128. A sketch of a roof and a sunken stoma.
As a matter of fact, the roof and sunken stomatal apparatus cannot be seen in one focus. In this sketch, roof
cells (Rc) and a stomatal apparatus are drawn with the same clearness. Guard cell ends (Ge) on theleft-hand side adhere closely to one another, but those at the right-hand side are much separated possiblyin the course of preservation. Subsidiary cell outlines are shown by the dotted lines. Guard cells (Gc) areheavily cutinized along their lateral margins. 13A. A typical roof with stellate mouth (holotype). The
stomatal apparatus is out of focus. 138. A sketch of a roof shown in Figure 13A. In both figures (Figures
13A and 138), the rachis is located upward.

962

Tamiko Ohana and Tatsuaki Kimura

The stomatal aperture is transverse to theveins (Figures 12A, B, 13A, B). The stomatal pit is covered by a dome-shaped or sacshaped roof consisting of 10-12 ordinary cellextensions surrounding a stoma. The mouthof each roof is stellate and more or less cutinized and is not located above the stomatal pit(or aperture) but unexceptionally at or near
the rachis-side edge of a roof (Figures 10,
llA, B, 12A).
The ramentum-like trichomes are present
randomly on the lower cuticle. Their preserved part consists of 5 cells in a row. They
are club-shaped and nearly parallel-sided,
100-150 pm long ahd 20 pm wide; most ofthem are at right angles to the vein course andare 13 per square mm in density (Figures IIA,
B). No papilla is recognized on the ordinaryand subsidiary cells. Petiole, stem and reproductive organs are not known.
Discussion and comparison. - Two leaf
specimens from Obira are small, elongatedwith slender pinnae having marked auricles
and crowded delicate veins. Therefore, theObira leaves are different in external form of
pinnae from those of Otozamites kerae described in this paper. Unfortunately wefailed to get sectional peels of pinnae andrachis of the Obira leaves.
In the Obira leaves, upper ordinary cells are
irregular in form and size, and their anticlinal
and end walls are rather thin and their sinuosity less marked than Otozamites kerae.
The Obira leaves have stomatal files andbands undeveloped, but in Otozamites kerae,
stomatal bands are well marked in the interveinal region. Ramentum-like trichomes arepresent on the lower cuticle in Obira leaves,
but trichome is not observed in Otozamites
kerae. Thus, both leaves are specificallydistinct. The presence of the so-called roof
(or sac cells, or roofing cells) covering thestomatal pit is a characteristic feature of many
Otozamites species (e.g. Harris, 1969; Barnard and Miller, 1976). Figures 12 and 13
show well-preserved roofs, but the stomatal
apparatus is sunken, and so it is impossible to

make microscopic observation of both organs
together.
In some species, such as Otozamites pulcher
Barnard et Miller, the mouth is open at itsedge on the rachis-side. Such a roof is notknown in the leaves of other bennettitalean
genera.
The internal structure of many Otozamites
species from Mesozoic plant sites has been
described (e.g. Harris, 1969; Archangelsky
and Baldoni, 1972; Barnard and Miller,
species, Ot1976). However, both theozamites kerae and o. takahashii, describedherein are different in their cuticles being
without papillae and trichomes (except for
the ramentum-like trichomes in O. takahashii).
The present ramentum-like trichomes are
similar in form to the imprinted ramenta on
the peduncle of Williamsonia hildae (Harris,
1969, fig. 580).
Diagnosis. - Leaf small in size. Pinnae set
closely, alternate, imbricate, attached toupper surface of rachis, rectangular withobtusely pointed apex and well-marked auricle concealing the rachis entirely. Veins
delicate, crowded and radiated.
Cuticle hypostomatic. Upper cuticle consisting only of ordinary cells with less marked
sinuous walls. Lower cuticle with marginal
rim devoid of stomata, main region consistingof ordinary cells with syndetocheiric stomata
not in files or bands. Roof formed by theextension of 10-12 encircling cells with its
stellate mouth open at rachis side.
Ramentum-like trichomes present on thelower cuticle. Other trichomes and papillae
absent on both cuticles.
Registered slides (All from the holotype).-INH-4-01 (Figures 9B, 10, liB, 13A),
4-02 (Figures IIA, 12A), 4-03 (Figure 9A).
References
Archangelsky, S., 1963: A new Mesozoic flora from
TicD, Santa Cruz Province, Argentina. Bull.
Brit. Mus. (Nat. Hist.). Geol.• vol, 8, no. 2, p. 1-

929.

Permineralized Otozamites leaves

92, pis. 1-12.
Archangelsky, S. and A. Baldoni, 1972: Revisi6n delas Bennettitales de la Formaci6n Baquero (Cretacico Inferior), Provincia de Santa Cruz. I.
Hojas. Rev. del Museo de La Plata. N.S., vol. 7,
no. 44, p. 195-165, pis. 1-16.
Barale, G., 1987: New observations on Otozamites
pterophylloides Schimper 1872, emend. Saporta
1874, from the Jurassic of France and Denmark.
Rev. Palaeobot. Palynol., vol. 51, p. 117-126.
Barnard, P.D.W. and J.e. Miller, 1976: Flora of theShemshak Formation (Elburz, Iran), Part 3 :Middle Jurassic (Dogger) plants from Katumbargah Vasek Gah and Imam Manak. Palaeontographica. Bd. 155B, Lfg. 1-4, p. 31-117, pis. lIS.
Bancroft, N., 1913: On some Indian gymnosperms.
Trans. Linn. Soc. London. Sec. ser. Bot., vol. 8,
p. 69-86, pIs. 7-9.
Bose, M.N., 1953: Ptilophyllum amarjolense, sp. nov.
from the Rajmahal Hills, Bihar. Proc. Natn.
Inst. Sci. India, vol. 19, no. 5, p. 605-6 I 2, pis. 13.
Bose, M.N. and M.L. Kasat, I972a: The genus Ptilophyllum in India. Palaeobotanist, vol. 19, no. 2,963

p. 115-145, pis. 1-14.
- - and - - , 1972b: On a petrified specimen of
Dictyozamites from the Rajmahal Hills, India.
Ibid., vol. 19, no. 3, p. 248-250, pis. 1-2.
Bose, M.N. and Zeba-Bano, 1978: The genus Dictyozamites Oldham from India. Ibid., vol. 25, p.
79-99, incl. pIs. 1-8.
Harris, T.M., 1964: The Yorkshire Jurassic flora.
l/. Caytoniales. Cycadales and Pteridosperms.
191 p., 7 pis. Brit. Mus. (Nat. Hist.).
- - , 1969: Ditto. III. Bennettitales. 186 pp., 7 pis.
Menendez, e.A., 1966: Fossil Bennettitales from theTic6 flora, Santa Cruz Province, Argentina.
Bull. Brit. Mus. (Nat. Hist.), Geol., vol. 12, no. I,
p. 1-42, pis. 1-19.
Rao, A.R. and V. Achuthan, 1967: Further contribution to our knowledge of the anatomy of Ptilophyllum. Palaeobotanist, vol. 16, no. 3, p.249257, incl. pis. 1-2.
Sharma, S.D., 1967: Investigation on the Jurassic
flora of Rajmahal Hills, India. 3. A review ofthe genus Ptilophyllum of Morris, with description of two new species from Amarjola in the
Rajmahal Hills. Palaeontographica. Bd. 120B,
Lfg. 5-6, p. 139- I 50.

Hokkaido ::ft$~, Kashima IN.tiiJ, Kamimakizawa J:~iR., Kumaoizawa ~~~iR., Nakakinenbetsu r:p*,c.~,'i]Ij, Obira IPP., Obirasibe 'J'fli, Rumoe (or Rumoi) JjJtij, Tappu Jt;fp, Yezo
Group 1J!;~l!tW.
::ft$~J:Jmt:L!Il!*'£~1~ Otozamites ~1~::Q: ::ft$~O)r:pfdl*iZ91JlUK.7t;fp'" QJ:JmIJ!;~
I!tWO)::QW<:j!{ / -:; "- J\..-r:p ~ut. !¥JT Jt-C;-Ittl; QiJ\ *IH.Ifr!lm~iJ~ J: <f*ff ~.n·n' QM¥l.J1~::Q~.£"'Qo ::;fs:glfU)'(li. g5imJ:~iR.. =~m~~~iR.:toJ: L1/J'flBp:p*,c.~,'i]Ij)lliJ'I?,,*~~.nt.: Otozamites ~~::;fs:O)?'i-:m:to J: L1P1JmO)*IH.Ifr!lm~~tHIH~::'WI.~ L. O. kerae:to J: L1 O.
takahashii 0) 2 ~lI~~,'i]IJl..-t.:o Otozamites kerae -c'li~0)=tX7C1¥J:mj!{iJ~ J: <f*ff ~.n.
~iWIl~::'li double-'U' ml~::.re7U'" Q*It'1t:5R C -to)~~ ~ -Jm~J/iJli'" Q .:. c iJ~-e ~ t.:o i t.:~0)J{mHUi~11~1If? r - AmlO)*IH.Ifr!l~~,'i]1j L t.:o Otozamites takahashii -eli. ~O)=tX
7C1¥J7fJj!{~~ l?iJ'~::'", Q':' c li-c' ~ t.t.iJ'"? t.:iJ~. ~:rL~1If? r - AmlO)*IH.Ifr!l~~,'i]Ij'" Q C
C ~ ~::.. ~b60-C ~11tl; Q~=§O)fftEiJ~~,'i]Ij-c' ~ t.:o Otozamites kerae ~::'~~60I?.nt.:~O)=tX7C1¥J:mj!{li. Otozamites ~-c'ItJ!fJJO)WI.~-c'tl; IJ. '" -* 7-7 A § O)~~::.~'" Q~~~*~~~Q,:,cKK.Lt.:o*n~~·*MJt~

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, pp. 964-972,9 Figs., December 30, 1991

930.

STENOPTERIS CYCLOSTOMA SAIKI, KIMURA ET HORIUCHI
SP. NOV. (POSSIBLE PTERIDOSPERM), FROM
THE LOWER CRETACEOUS CHOSHI GROUP,
IN THE OUTER ZONE OF JAPAN*
KEN'ICHI SAIKI
Department of Earth Science and Astronomy, University ofTokyo, 3-8-1 Komaba, Meguro-ku, Tokyo 153

T ATSUAKI KIMURA
Institute of Natural History, 24- 14-3 Takada,Toshima-ku, Tokyo 171
andJUNJI HORIUCHI
Attached School, Tokyo Gakugei University (Oizumi Campus),
1-22-5, Higashi-Oizumi, Nerima-ku, Tokyo 178

Abstract. Stenopteris cyclostoma Saiki, Kimura et Horiuchi, sp. nov. is based on leaf
fragments obtained from the Choshi Group of marine origin. This species is characterized
by its fern-like pinnate leaves with wedge-shaped pinnae divided into narrow segments
directed forward, each with a single vein, and cycad-like leaf cuticle bearing thick-walled
cells. Such a leaf form has not been recorded in any extant or fossil cycads. We regard thepresent plant as a possible pteridosperm. Stenopteris cyclostoma is the first reliable recordof a pteridospermous plant in the Ryoseki-type flora in the Outer Zone of Japan except for
the record of Sagenopteris. This paper deals with the detailed macro- and microscopicaldescription of Stenopteris cyclostoma.
Key words.

Pteridosperm, Stenopteris, Lower Cretaceous, Choshi Group.

Hatoyama, Choshi Peninsula (possibly belonging to the Ashikajima Formation; localities are shown in Figure I) and were described by Nishida (1960) as Czekanowskia
rigida Heer.
Fortunately, in the three specimens
mentioned above the cuticle is well preserved.
According to our observation, their cuticle israther cycad-like, but in external forms they
are far apart from known extant and fossil
cycads and are rather fern-like. Hence weregard the present leaf fragments as a possible

Introduction

A single broken pinnate leaf was obtained
from the Barremian Ashikajima Formation of
the Choshi Group, which is mostly of marine
OrIgm. In addition, two specimens kept at
the National Science Museum, Tokyo were
examined. They were collected by Nihei
Takagi from the sandy shale bed at*Received August 15, 1991; accepted September 24,
1991
964

930.

Stenopteris cyclostoma sp. nov.

CHOSHI-+-I,"---35'N
'40'E

I

//

PENINSULAPACIFIC OCEAN'-------' 1k m

B

1Figure 1. Localities of Stenopteris cyclostoma
Saiki, Kimura et Horiuchi, sp. nov., in the Ashikajima Formation, Choshi Group. A; Ashikajima
locality. B; Hatoyama locality.

pteridosperm and refer them to Stenopteris
Saporta em. Harris (1964), and propose thename S. cyclostoma, sp. nov. This paper
deals with its detailed macro- and microscopical description.
In fact, it is quite difficult to determine thetaxonomic position for such leaf fragments,
depending only on their external morphology. In many cases, such leaf fragments
would unavoidably be placed in form-genera
such as Sphenopteris or Acrostichopteris forconvenience's sake. The Yorkshire
Stenopteris had also been assigned toSphenopteris, Hymenophyllites and even to
Ruffordia (Harris, 1964).
As repeatedly mentioned by one of us(Kimura), there were two distinct floras during Late Jurassic and Early Cretaceous timein Japan and its adjacent regions: theTetori-type flora distributed in the Inner
Zone of Japan and its extension, which flourished under a warm-temperate environment,
and the Ryoseki-type flora distributed typically in the Outer Zone of Japan and its
extension, which flourished under a subtropical-tropical and rather arid environment(e.g., Kimura, 1987a, b).
The Ryoseki-type flora is characterized by
the abundance of matoniaceous (including
Weichselia, restricted in occurrence to the965

Early Cretaceous plant sites) and schizaeaceous ferns, of varied bennettitaleans such as
Zamites and Ptilophyllum (incl udi ng
Nipponoptilophyllum with leaves of bipinnate
habit), of varied cycadaleans such as Nilssonia
with coriaceous leaves (incl udi ng N.
schaumburgensis-type leaves) and of variedcheirolepidiaceous and other conifers, such as
Frenelopsis, Brachyphyllum, Cupressinocladus and Parasequoia. The above genera
and groups have not been recorded in the
Tetori-type flora except for schizaeaceous
ferns. It is worth mentioning that so far to
our knowledge, ginkgoaleans, czekanowskialeans and Podozamites leafy shoots havenot been recognized in the Ryoseki-type flora.
The Lower Cretaceous Choshi Group ismostly of marine origin and yields abundantfossil plants of the Ryoseki type in association with various fossil animals. Althoughmost fossil plants are fragmental, most
gymnospermous plants fortunately have wellpreserved cuticles.
The Ryoseki-type flora is known from
Upper Jurassic-Lower Cretaceous plant sites
mainly along the Pacific Coast of Japan forabout 1,500 km (roughly from 131'E to 142'E
and from 33'N to 40'N), but unfortunately
fossil ferns or fern-like plants have lost detailsof their reproductive organs and leaf cuticles,of the gymnosperms, except for the Choshi
plant sites. Therefore, fairly a large numberof plants in the Ryoseki-type flora have beenleft undetermined taxonomically or they havebeen unavoidably placed in form-genera for
convenience. In fact, no fossil plants assignable to pteridosperms had been recorded in
the Ryoseki-type flora except for Sagenopterisleaves or leaflets and Pachypteris ? sp. (Kimand Kimura, 1987; Oishi, 1940, p. 422).
Under the circumstances, the presence ofthe present pteridospermous Stenopteris
cyclostoma with cycadalean cuticle would besignificant for the taxonomy of the Mesozoic
fern-like plants in the Ryoseki-type flora inJapan. It is highly probable that we canobtain additional specimens in future from966

Ken'ichi Saiki, Tatsuaki Kimura and Junji Horiuchi

5
Figures 2- S. Slenopleris cyclosLOma Saiki , Kimura et Horiuchi , sp. nov.: 2. External rorm or theholotype (NSM PP-9009). X 1.5. 3. Drawn rrom the holotype, showing veins dichotomously rorked. 4.
External rorm or one or paratypes (NSM PP-75 16). Rein se rted rrom Ni shida (1960, pI. 2, fig. 12). X I. S.
Drawn rrom Figure 4, showing alternately lobed lamina. X I.

the Choshi Group which would be taxonomically useful for the plants at present
remaining unclassified or undetermined in the
Ryoseki-type flora.

Acknowledgements
We express our sincere gratitude to Dr.
Ikuwo Obata of the National Science
Museum, Tokyo who provided the specimens
kept at the museum on loan for the present
study. The junior author (Saiki) is muchindebted to Prof. Takashi Hamada of the
University of Tokyo for his continuous
encouragement. Our thanks are extended to

Dr. Shya Chitaley of the Cleveland Museumof Natural History, Ohio, for checking thelanguage of our manuscript. We thank Mr.
Satoshi Kimura of the Atagawa Tropical andAlligator Garden for his kindness in providing extant cycad leaves.
This study was supported in part by the
Grant-in-Aid for Scientific Research from the
Ministry of Education, Science and Culture(no. 02640615 given to Kimura) .

Material and methods
Three leaf- fragments investigated wereobtained from the Ashikajima Formation~
Figure 6. Cuticles or Slenopteris cyclosLOma Saiki , Kimura et Horiuchi, sp. nov. 1, 2, 8 an d 9 are
SEM-photographs prepared rrom the holotype (NSM PP- 9009). 1. Internal view or the upper (adaxial)
cuticle. Rows or the thick-walled cells are indicated by arrows. X 65. 2. Internal view or the lower(abax ial) cuticle. Note that the thick-wa lled cells are absent. X 65. 3. Distribution and orientation or
stomata, drawn rrom Figure 1. Orientation or stomatal ape rture is ignored in the stoma figured by 'open
circle'. 4. Distribution a nd orientation or stomata, drawn rrom Figure 2. S. A stoma prepared rrom NSM
PP-7516. x 500. 6. A stoma photographed by phase contrast microsco pe (NSM PP-7516). x 400. 7.
Stomata photographed by light microsco pe, showing thickly cutinized subsidiary cells (NSM PP- 7516). X
200. 8. Internal surrace or thin-walled cell or the upper cuticle, showing granules. X 20,000. 9. Inside
view or upper cuticle, showing thick- and thin-walled cells. X 550. K ; thick-w alled cell , N ; thin-walled
cell.

930.

Stenopteris cyclostoma sp. nov.

967

Ken'ichi Saiki, Tatsuaki Kimura and Junji Horiuchi

968(Barremian In age) of the Choshi Group(Obata et al., 1982). The holotype (NSM
PP-9009; Figures 2-3) was collected from
Ashikajima, Choshi City, Chiba Prefecture(Figure lA). Other specimens (paratypes)(NSM PP-7513 and 7516) were collected from
Hatoyama (Figure IB) and were kept in the
National Science Museum, Tokyo (Figures 45). The holotype, its SEM-stages andcuticular preparations are also kept in the
National Science Museum, Tokyo.
Fossil leaves were treated by Schulze's solution followed by dil uted N aOH and mounted
in Canada balsam. Eight genera of extant
cycads (Bowenia, Ceratozamia, Cycas, Dioon,
Encephalartos, Macrozamia, Stangeria andZamia) used for cuticular comparison weretreated in 10% chromic oxide for 24 hours atroom temperature.
For the SEM observation, cuticles were
coated by gold with a JEOL FC-lIOO apparatus and photographed by JEOL J-15.
Description of specimens
Three leaf-fragments were examined (Figures 2-5). The largest specimen (NSM PP7516) is 5.5 cm long. Pinnae are wedgeshaped and are dissected alternately up to
three times or more to form slender segments.
The segment is linear, up to 30 mm long andabout 2 mm wide, narrowing gradually
toward the acute apex. Each segment bears asingle median vein, approximately 0.5 mm
wide. Veins are dichotomously forked at anangle of 15-30 degrees and the angle tends to
get reduced upward. Basal portion of seg~ments is narrowed slightly. Lateral vein isabsent.
Scanning electron micrographs (Figures 61,2) show upper and lower cuticles of thesame part of the holotype. Cuticle is amphistomatic. Stomata are distributed approximately 20 per square mm on both sides.
Stomatal bands are rather well defined, separated by nonstomatal bands 5-13 cells wide
(11O-200.um in width) (Figures 6-1, 2, 3,4).
Papilla or trichome base is absent.
Upper (adaxial) cuticle consists of stomata
and two types of ordinary cells; one withcutinized periclinal walls, about 8 Jim thick(thin-walled cells) and the other with periclinal walls consisting of two cuticular layers(thick-walled cells) (Figures 6-1, 9). Innersurface of periclinal walls is disorganized.
Thin-walled cells are elongate-rectangularor short-rectangular or sometimes triangularin shape, 60-160.um long and 1O-35.um
wide. Anticlinal walls of the thin-walled
cells are straight, rather broad, ca. 2.um highand ca. 5.um wide. Inner surface of periclinal walls is granulated; the granules are
ca. 0.2.um in diameter (Figure 6-8).
Thick-walled cells are 15-50.um long and6-25.um wide, rectangular or triangular inshape, consisting of both external and internal cuticular layers. The external layer isabout 8.um thick, corresponding to that ofthin-walled cells, and the internal layer is lessthan l.um thick. The thick-walled cells
form longitudinal rows, 2-14 cells long (60400.um) and typically a single cell (sometimes up to 4 cells) wide. The rows arescattered between thin-walled cells. The

Figure 7. Stenopteris cyclostoma Saiki, Kimura et Horiuchi, sp. nov., the holotype (NSM PP-9009).
1. Internal view of a stoma, showing sunken aperture. X 650. 2. External view of a stoma. Groove
surrounding the stomatal aperture is indicated by an arrow. X 650. 5. Cross section of a thick-walled cell,showing external and internal layers. X 5,500. 6. A figure drawn from Figure 5. e.l.; external layer ofthick-walled cell, Ll.; internal layer of thick-walled cell. X 5,500. 8. Light microscope photograph of cells.
Note well stained thick-walled cells. X 500. Figures 7-3, 4. Stangeria eriopus Nash, for comparison. 3.
Internal view of a stoma for comparison. X 650. 4. External view of a stoma. X 650. Figures 7-7, 9.
Dioon spinulosum Dyer, for comparison. 7. Cross section of cuticle, showing external and internal layersof thick-walled cell. X 1,700. 9. Thick-walled cells stained by alcoholic safran in, for comparison. X 500.
Figures 7-1, 2, 3, 4, 5, 7 are SEM-photographs.

930.

Stenopteris cyc!ostoma sp. nov.

969

e.!.

6

970

Ken'ichi Saiki, Tatsuaki Kimura and Junji Horiuchi

morphology of the present thick-walled cells
is similar to those of some extant cycads in
two-layered construction (Figure 7-7). The
thick-walled cells are well stained by alcoholic safranin as seen in some extant cycads(Figures 7-8, 9).
Lower (abaxial) cuticle is thinner than theupper. It consists of thin-walled cells andstomatal apparatuses. Thin-walled cells of
upper and lower cuticles are similar to eachother.
Stomatal apparatus is haplocheiric, 50-80,urn in diameter (Figures 6-5, 6, 7; Figure 71). Subsidiary cells are thickly cutinized,
consisting of polar and 4-6 lateral cells.
Polar cells are rectangular or polygonal inshape, 27-46,um long. Lateral subsidiarycells are bean-shaped, about 20-45,um long.
Guard cells are slightly sunken, each cell 3550,um long and 5-7,um wide. Stomatal
aperture is fusiform, about l2,um long.
Outer stomatal ridge (rim) is thickened, forming a fusiform opening, about lO,um long.
Papilla is absent around the stomatal aperture(Figure 7-2). Morphology of the present
stomatal apparatus is similar to those of someextant cycads, especially of Stangeria eriopus
Nash (Figures 7-3,4).

Discussion and comparison
Although the present leaves are quite fragmental, they are doubtless of pinnate habit
and what we regard as a pinna has an external
resemblance to that of some ferns. However,the leaf cuticle is gymnospermous and is likethose of some extant cycads. Hence weregard the present leaves as those of pteridospermous Stenopteris.
On the basis of two specimens now kept at
the National Science Museum, Tokyo, Nishida (1960) assigned them to Czekanowskia
rigida Heer. However, they are clearly distinguishable from the leaves of Czekanowskia
by the presence of the thick-walled cells. In
addition, Nishida (1960) mentioned the presence ·of dwarf (or short) shoots covered byscale-leaves at the base of a wedge-shaped
leaf-bundle. But we found neither suchdwarf shoots nor scale-leaves in his specimens.
Among the Mesozoic pteridosperms known
from the present Northern Hemisphere, several leaves assigned to genera such as Pachypteris Brongniart, Stenopteris Saporta andThinnfeldia Ettingshausen have a rather close
external resemblance to the present leaves.
Of these, the leaves of Pachypteris and Thinnfeldia are clearly distinguished from the present leaves by having pinnate venation.
Externally the present leaves are most close to
those of Stenopteris and their external andcuticular features agree generally with Harris'
emended diagnosis of Stenopteris (Harris,1964). Harris (1932, 1937) described two
species of Stenopteris from Scoresby Sound,
East Greenland and three species of this genus
from the Yorkshire Jurassic (Harris, 1964).
However, the present leaves are different fromthose of Harris' five species because of thepresence of thick-walled cells and lack oftrichomes.
From the Choshi Group, we collectedmany Ryoseki-type gymnosperms with preserved cuticles. Generally, they are characterized by the absence of trichomes or less developed trichomes, if present. It seems thatthese plants might have flourished under an
oceanic environment during Early Cretaceoustime.
Thomas and Bancroft (1913) recognized
thick-walled cells in the cycads. According
to Pant and Nautiyal (1963), the thick-walled
cells are present in extant cycads excluding
Cycas and some fossil cycads, and they havethickened anticlinal walls and are easily
stained by alcoholic safranin (Figure 7-9).
According to our SEM-observation, thethick-walled cells in Dioon spinulosum Dyer
are not caused by a thickened cuticle, but by
two cuticular layers (Figure 7-7). Such a
two-layered structure is also recognized in
Bowenia, Ceratozamia, Encephalartos, Macrozamia and Zamia, but it has not been recog-

930.

Stenopteris cyclostoma sp. nov.

nized in Stangeria and Cycas. The thickwalled cells of the present fossil leaves have
similar structure to the above and are alsoeasily stained by alcoholic safranin (Figures
7-5,6,8). Therefore, we could not differentiate between the thick-walled cells of thepresent fossil leaves and those of extant
cycads.
The present cuticle is very close to that ofAlmargemia incressata originally describedby Archangelsky (1966) from the Baquero
Formation, Argentina in the presence ofthick-walled cells and cycad-like stomata,respectively. Almargemia was supposed byArchangelsky (1966) as a cycadalean, because
both thick- and thin-walled cells were present
on the same epidermal plane. But the ultimate
segments of Almargemia are distinguishedfrom those of the present leaves by having 34 veins.
So far as we know, the present leaves differ
in external form from those of extant andfossil cycads hitherto known, but they resemble in their external form those of some ferns
or pteridosperms. On the other hand, thecuticle in the present specimen bears characteristic thick-walled cells which have been
said to be a characteristic feature of most
extant cycads. The present leaves are notthose of fern, because in ferns a cuticle isgenerally absent or less developed.
Under the circumstances, it seems to beappropriate to give the present plant a newgeneric name. But at present we refrain fromdoing it and regard the present plant as a new
species of pteridospermous Stenopteris,
because the present specimens are all fragmental and the plant is most close to
Stenopteris in external morphology. In fact,
as stated by Harris (1964, p. 31), the taxonomic division of the non-bennettitalean
cycadophyte leaves into Cycadales and Pteridospermales is difficult when depending only
on their leaf-form and leaf-cuticle, since wehave no single character to mark the twoclasses from one another.

971

Taxonomy
Pteridospermales (?)
Genus Stenopteris Saporta,
1873 em. Harris, 1964

Stenopteris cyclostoma
Saiki, Kimura et Horiuchi, sp. nov.
Figures 2-6,7-1,2,5,6,8

Czekanowskia rigida Heer: Nishida, 1960, p. 191, pI.
2, figs. 12, 13.

Holotype: NSM PP-9009. Para types :NSM PP-7513, 7516. Stratum typicum:
Ashikajima Formation (Barremian in age),
Choshi Group. Loclls typicus: Ashikajima,
Choshi City, Chiba Prefecture. Derivatio
nominis: After circular stomatal complex.
Occurrence. ~ Rather rare at present.
Diagnosis. ~ (Whole leaf unknown). Pinnae wedge-shaped, dissected alternately up to
three times or more to form slender segments.
A segment linear in shape, narrowing gradually toward the acutely pointed apex. Eachsegment bearing a single median vein. Veins
dichotomously forked at an angle of 15-30
degrees, the angle tending to reduce upward.
Basal portion of segments slightly narrowed.
Lateral vein absent.
Cuticle amphistomatic. Stomata approximately 20 per square mm on both sides, forming well-defined rows. Papilla or trichomebase absent. Upper (adaxial) cuticle consisting of stomata, and two types of ordinary
cells: One with cutinized periclinal walls(thin-walled cells) and the other with periclinal walls consisting of two cuticular layers(thick-walled cells). Lower (abaxial) cuticle
thinner than the upper, consisting of thinwalled cells and stomata. Thin-walled cells
similar to those of the upper cuticle.
Stomatal apparatus haplocheiric. Subsidiary cells thickly cutinized, consisting of 2
polar cells and 4-6 lateral cells. Guard cells
slightly sunken. Stomatal aperture lon-Ken'ichi Saiki, Tatsuaki Kimura and Junji Horiuchi

972

gitudinally orientated and sunken. Externalsurface of stoma surrounded by well-definedgroove. Outer stomatal ridge (rim) thickened, forming fusiform opening. (Reproductive organs not known.)
References
Archangelsky, S., 1966: New gymnosperms from theTic6 flora, Santa Cruz Province, Argentina.
Bull. Brit. Mus. (Nat. Hist.), Geol., vol. 13, no. 5,
p. 261-295, pis. 1-8.
Harris, T.M., 1932: The fossil flora of Scoresby
Sound East Greenland. Ill. Medd. om Gr9!
nland, Bd. 85, no. 3, p. 1-112, pis. 1-19.
- - , 1937: Ditto. V. Ibid., Bd. 112, no. 2, p. 1114, pI. 1.
- - , 1964. The Yorkshire Jurassic flora. I/.
Caytoniales, Cycadales and pteridosperms.
Brit. Mus. (Nat. Hist.), ix + 185 pp., 7 pIs.
Kim, J. H. and T. Kimura, 1987: Cuticle ofSagenopteris (Caytoniales), an extinct gymnospermous plant, first observed in Japan. Proc.
Japan Acad., vol. 63B, p. 179-182.
Kimura, T., 1987a: Recent knowledge of Jurassic and
Early Cretaceous floras in Japan and phytogeo-

graphy of this time in East Asia. Bull. Tokyo
Gakugei Univ., IV, vol. 39, p. 87-115.
- - , 1987b: Geographical distribution of Palaeozoic and Mesozoic plants in East and Southeast
Asia. In, Taira, A. and M. Tashiro eds.: Historical biogeography and plate tectonic evolutionof Japan and Eastern Asia, p. 135-200. Terrapub., Tokyo.
Nishida, M., 1960: Lower Cretaceous plants in Choshi Peninsula, Chiba. Journ. Coll. Arts andSci., Chiba Univ., vol. 3" no. 2, p. 187-194, pis.
1-3 (in Japanese with English description of a
new species).
Obata, I., Maiya, S., Inoue, Y. and M. Matsukawa,
1982: Integrated mega- and micro-fossil biostratigraphy of the Lower Cretaceous Choshi
Group, Japan. Bull. Natn. Sci. Mus., Tokyo,
C, vol. 8, no. 4, p. 145-179, pis. 1-7.
Oishi, S., 1940: The Mesozoic floras of Japan.
Journ. Fac. Sci., Hokkaido Imp. Univ., IV, vol.
5, nos. 2-4, p. 123-480, pis. 1-48.
Pant, D.D. and D.D. Nautiyal, 1963: Cuticle andepidermis of recent Cycadales. Leaves, sporangia and seeds. Senck. BioI., Bd.44, no. 4, p.
257-347, pis. 19-29.
Thomas, H.H. and N. Bancroft, 1913: On the cuticles
of some recent cycadean fronds. Trans. Linn.
Soc., Ser. 2 (Bot.), vol. 8, p. 155-204, pis. 17-20.

Ashikajima #iHllth, Choshi &J~T, Hatoyama iBl1.l:.Jll, Ryoseki flJt:o, Tetori =Fl&..
TfffiBlII!*&J~T~lW.il: Stenopteris ~~fi: ~1;:.:htdr.:Oft."t""L, :/ ?~~O)~Jt-c'a;-to)J:.lIiiO)jjlit*IBOOHt., *$7tO)m~ 'J T"/~:ta J: Lf-$O){r.:o 'J T"/ ~~;::~~~ I;:.
:h Q 2 ~ ii' I;:. tJ:. Q cuticle ~iI'fHE"t Q0 *{r.:O~ft., 7i-~ft.:/ ?(~-c: a; Q ,~~;::%,ji(iI~a; IJ ,-t O)7i-~fHOH[] I;:.:h L~' Q {r.:O~O)r:p-c::/ ?·fiT~ ~.:t I;:.:h L ~'Q Stenopteris ~;::lil

Q iI~,~Jli~'O)-c:,c;: ;:~;:: Stenopteris cyclostoma a-~fic L-LtJi!:P~"tQo~*~-. *ttj!~ . WlP'lIIIfii'El

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, pp. 973-1002, 10 Figs., December 30, 1991

931. EARLY EVOLUTION OF THE FAMILIES VERBEEKINIDAE
AND NEOSCHW AGERINIDAE (PERMIAN FUSULINACEA) IN
THE AKIYOSHI LIMESTONE GROUP, SOUTHWEST JAPAN*
KA TSUMI UENO
Institute of Geoscience, University of Tsukuba,
Ibaraki 305

Abstract. Fifteen species of verbeekinid and neoschwagerinid fusulinaceans are discriminated from the Akiyoshi Limestone Group below the Misellina (M.) clalldiae Zone in
the Kaerimizu area. Among them, Misellina (Brevaxina) nipponica, sp. nov., M. (M.)
postclalldiae, sp. nov. and M. (M.)? ventricosa, sp. nov. are new to science. Stratigraphic
distribution of the early representatives of verbeekinid and neoschwagerinid fusulinaceans,besides their ancestral forms, makes it possible to establish 10 fusulinacean biohorizons inthe stratigraphic interval between the first occurrence of Pamirina (Levenia) leveni and thatof Parafllslliina kaerimizensis. Of them, three biohorizons, which are well recognized in thesurveyed area, define the base of the Pamirina (Levenia) leveni, Misellina (Brevaxina)
dyhrenfllrthi olai and Misellina (M.) clalldiae Zones established herein. Biostratigraphic
evidence observed in the Akiyoshi Limestone Group provides many important data concerning the origin and early evolution of the families Verbeekinidae and Neoschwagerinidae.
Key words. Akiyoshi Limestone Group, Kaerimizu, Middle Permian, Neoschwagerinidae, Verbeekinidae.

Leven, 1982; Yang, 1985). Ozawa (1970)
rethought the phylogenetic relationship of theneoschwagerinid species and confirmed thatthree distinct lineages are present in the family Neoschwagerinidae. He also gave diagnoses of all genera referred to the Verbeekinidae and Neoschwagerinidae. Up to thepresent, about 300 species have been proposed
belonging to these families.
In the Akiyoshi Limestone Group below
the Misellina (M.) claudiae Zone, the following six species of verbeekinid and neoschwagerinid fusulinaceans, Misellina (M.)
claudiae (Deprat), M. (M.) ovalis (Deprat),
M. (M.) parvicostata (Deprat), M. (Brevaxina) dyhrenfurthi otai Sakaguchi and
Sugano, Maklaya pamirica (Leven) and M.
akiyoshiensis Ota (MS), occur, although
totally about 30 species of them are recordedin the whole extent of the Akiyoshi Lime-

Introduction

The families Verbeekinidae and Neoschwagerinidae flourished and diversified in
the Middle Permian of the Tethyan realm,
and are considered to be the most importantfusulinacean groups in determining biostratigraphic zonation and geologic age in that
period. Since the beginning of this century,
dozens of works have been devoted to thephylogeny and classification of the Verbeekinidae and Neoschwagerinidae (e.g. Deprat,
1913; Ozawa, 1925a, 1927; Lee, 1933; Gubler, 1935; Thompson, 1948; Kanmera, 1957 ;Miklukho-Maklay et al., 1958; Minato andHonjo, 1959; Morikawa and Suzuki, 1961;
Hanzawa and Murata, 1963; Ozawa, 1970;'Received September 5, 1991 ; accepted October 28,
1991
973

Katsumi Ueno

974

Geologic setting
Shishide-dai

1:J

Akago

Edo

N

o
I

+
2km
r

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

stone Group (Toriyama, 1958; Ota, 1977;
Ozawa and Kobayashi, 1990 and others).
Recently, I reported three species referable
to the genus Pamirina and one species tentatively assigned to the subgenus Brevaxina of
the genus Misellina from the Pamirina(Levenia) leveni Zone in the Akiyoshi Limestone Group of the Kaerimizu area, anddiscussed the evolution of the genus Pamirina, which is considered to be an ancestral
phylogenetic stock of the family Verbeekinidae (Ueno, 1991). In that paper, I
proposed the subgenus Levenia for primitive
Pamirina without alveolar structure in the
spirotheca.
In the present paper, I discuss the early
evolution and phylogeny of verbeekinid andneoschwagerinid fusulinaceans in the Akiyoshi Limestone Group below the Misellina(M.) claudiae Zone in the Kaerimizu area.
Moreover, 14 species of verbeekinid and neoschwagerinid fusulinaceans including three
new ones; Misellina (Brevaxina) nipponica,
M. (M.) postclaudiae and M. (M.)?
ventricosa, are also described.

The Kaerimizu area is situated in the northeastern part of the Akiyoshi Plateau (FigureI). Since the discovery of large-scale overturned structure by Ozawa (1923), more thana dozen geological and paleontologicalstudies have been carried out (Ozawa, 1925b;
Hanzawa, 1954; Toriyama, 1954, 1958; Hasegawa, 1958, 1963; Sakaguchi et al., 1966;
Sakaguchi and Sugano, 1966; Kawano,
1967; Ota, 1968, 1977; Ota et al., 1973; Hasegawa et al., 1979; Ozawa and Kobayashi, 1990; Ueno, 1991). Ota (1977) divided
the Akiyoshi Limestone Group in this area
into the following 10 zones: the Fusulinellabiconica, Triticites simplex, Pseudofusulina
vulgaris, Pseudofusulina ambigua, Misellina
claudiae, Parafusulina kaerimizensis, Afghanella schencki, Neoschwagerina craticulifera,
Verbeekina verbeeki and Colania douvillei
Zones in ascending order.
According to my recent investigation, Ota's
Pseudofusulina ambigua Zone can be subdivided into the lower Pseudofusulina ex gr.
kraffti and the upper Pamirina (Levenia)
leveni Zones, and Misellina claudiae Zone
into the lower Misellina (Brevaxina) dyhrenfurthi otai and the upper Misellina (M.)
claudiae Zones, respectively,
Figure 2 shows the distribution of fusulinacean zones in the Kaerimizu area. The
zones are distributed concentrically and getyounger downward. Consequently, the
Akiyoshi Limestone Group in this area lies
almost horizontally and is in reverse order.
In the stratigraphic interval between thefirst occurrence of Pamirina (Levenia) leveni
and that of Parafusulina kaerimizensis, 10
fusulinacean biohorizons, which are definedby the first occurrence of the characteristic
species of verbeekinid, neoschwagerinid andtheir ancestral fusulinaceans, are recognized.
They are shown in Figure 3. Of these, threebiohorizons characterized by the first occurrence of Pamirina (Levenia) leveni, Misellina(Brevaxina) dyhrenfurthi otai and Misellina

931.

Evolution of Verbeekinidae and Neoschwagerinidae(M.) claudiae, are well recognized in theexamined sections and define the base of threefusulinacean zones, respectively. Others areobserved only in some sections.
The Pamirina (Levenia) leveni Zone, about
25 m thick, conformably overlies the Pseudofusulina ex gr. kraffti Zone. This zone isdefined as the stratigraphic interval between
the first occurrence of Pamirina (Levenia)
leveni and that of Misellina (Brevaxina)
dyhrenfurthi otai. Algal oncoidal limestone
is predominant with a subordinate amount ofalgal sparitic limestone and muddy limestone.
I have previously listed the identified fusulinaceans from the Pamirina (Levenia) leveni
Zone (Ueno, 1991).
The Misellina (Brevaxina) dyhrenfurthi
otai Zone is about 15 m in thickness, andconformably overlies the Pamirina (Levenia)
leveni Zone. The limestone facies of thiszone is similar to that of the Pamirina(Levenia) leveni Zone. The following fusulinaceans are discriminated: Toriyamaia laxiseptata Kanmera, Schubertella sp., Mesoschubertella sp., Neofusulinella sp.,
Pseudofusulina postkraffti (Leven), Pseudoreichelina spp., Pseudoendothyra yobarensis(Ozawa), Pseudoendothyra spp., Misellina(Brevaxina) dyhrenfurthi otai Sakaguchi and
Sugano, and Misellina (M.) sp. A.
The Misellina (M.) claudiae Zone, about
30 m thick, is defined as the stratigraphic
interval from the first occurrence of Misellina(M.) claudiae to that of Parafusulina kaerimizensis. The stratigraphic relationship between the present zone and the subjacent
Misellina(Brevaxina)

dyhrenfurthi

otai

Zone is conformable. Foraminiferal algal
sparitic limestone is dominant in the lower
part, algal micritic limestone and muddy
limestone are predominant with a subordinate amount of algal sparitic limestone in the
middle and upper parts. The identified
fusulinaceans are: Mesoschubertella sp.,
Neofusulinella giraudi Deprat, N. pseudogiraudi (Sheng), Neofusulinella sp., Minojapanella (Russiella) pulchra Miklukho-975

Maklay, Pseudofusulina cf. quasifusuliniformis Leven, Nagatoella orientis (Ozawa),
Nagatoella sp., Chusenella tingi Chen,
Pseudoendothyra spp., Misellina (M.)
claudiae (Deprat), M. (M.) postclaudiae, sp.
nov., M. (M.) spinosa Han, M. (M.)?
ventricosa, sp. nov., Misellina (M.) sp. B,
Armenina salgirica Miklukho-Maklay, A. cf.
asiatica Leven, Maklaya saraburiensis Kanmera and Toriyama, M. pamirica (Leven),
Maklaya sp. and Neoschwagerina simplex
Ozawa.
The geologic age of these three zones is
considered as follows: the Pamirina(Levenia) leveni Zone is almost late Yakhtashian, the Misellina (Brevaxina) dyhrenfurthi
otai Zone is early Bolorian, and the Miselliana (M.) claudiae Zone is late Bolorian to
Kubergandian of the standard stratigraphic
scheme for the Tethyan province proposed byLeven (1980).

Some considerations on the early
evolution and phylogeny of thefamilies Verbeekinidae
and N eoschwagerinidae
A total of 15 species of verbeekinid andneoschwagerinid fusulinaceans are identified
in the Pamirina (Levenia) leveni, Misellina(Brevaxina) dyhrenfurthi otai and Misellina(M.) claudiae Zones. The stratigraphic distribution of the verbeekinid and neoschwagerinid fusulinaceans are shown in
Figure 4.
Concerning the early evolution andphylogeny of the families Verbeekinidae and
Neoschwagerinidae, the occurrences of Misellina (Brevaxina)? sp., Misellina (M.) sp. A,
M. (M.)? ventricosa, sp. nov. and Misellina(M.) sp. B in the Akiyoshi Limestone Groupare significant and interesting. They suggest
an intimate phylogenetic relationshipbetween Pamirina (s.s.) and Brevaxina,
Brevaxina and Misellina (s.s.), Misellina (s.s.)and Armenina, and Misellina (s.s.) and Maklaya, respectively (Figure 5).

Katsumi Vena976.... .... : :::: :: .. : ..
~:<EB~~~~l~n·.

o

200m

I

LEGEND
Kaerimizu
.6.

1
2

3
4

5
6

•

7

931.

Evolution of Verbeekinidae and Neoschwagerinidae

ZO N E
Paraf. kaerimizensis
Mise/lina (M.)
c/audiae

977

FUSULINACEANBIOHORIZON.... F. Parafusulina kaerimizensis.... F. Neoschwagerina simp/ex.... F. Mak/aya pamirica.... F. M. saraburiensis & Armenina
sa/girica.... F. Mise/lina (M.) postc/audiae.... F. Misellina(M.) c/audiaeM.(B.) dyhrenfurthi otai
Pamirina (Levenia).... F. Mise/lina (M.) sp.A.... F. M. (Brevax ina) dyhrenfur thi otai.... F. Mise/lina (Brevaxina) nipponica.... F. Pamirina (P.) darvasica/eveni.... F. Pamirina (Levenia) /eveni

Pseudof. ex gr. kraffti
F.: First occurrence

Figure 3. Fusulinacean biohorizons in the Pamirina (Levenia) leveni, Misellina (Brevaxina)
dyhrenfurthi atai and Misellina (M.) claudiae Zones.

Based on biostratigraphical and morphological observations, it has been considered that certain species of Levenia, Pamirina(s.s.), Brevaxina and Misellina (s.s.) comprisea single evolutionary series, and that Misel!ina (s.s.) is the ancestral stock of later verbeekinid and neoschwagerinid genera (Ozawa,
1970; Kobayashi, 1977 and others). The
occurrences of some transitional forms orclosely related species between two distinct
genera together with the biostratigrapic evidence in the Akiyoshi Limestone Group
prove the above-mentioned considerations.
Evidently, the Akiyoshi Limestone Group
has an almost continuous record in relation
to the origin and early evolution of the fam~ilies Verbeekinidae and Neoschwagerinidae.

Systematic paleontology
Among the illustrated fusulinaceans, four
species; Pamirina (P.) darvasica Leven,
Pamirina (Levenia) leveni Kobayashi, P. (L.)
evoluta Sheng and Sun, and Misellina(Brevaxina) ? sp. have already been described(Ueno, 1991). All specimens identified inthis paper are deposited in the paleontological collections of the Akiyoshi-dai
Museum of Natural History. The abbreviation ASM is the prefix ofthe specimen repository number.

Figure 2. Distribution of foraminiferal zones and fossil localities in the Kaerimizu area. I:
Chalaraschwagerina vulgaris Zone, 2: Pseudafusulina ex gr. kraffti Zone, 3: Pamirina (Levenia) leveni
Zone, 4: Misellina (Brevaxina) dyhrenfurthi atai Zone, 5: Misellina (M.) claudiae Zone (M: Misellina(M.) claudiae swarmed layer), 6: Parafusulina kaerimizensis Zone, 7: fossil locality studied herein.

2'

'"c

:=:""l~ ciQ''"o ..=
to

A

I»

~ f>o

A AA

A

A

I»

>~.

...
-.1»
,<'0
~

_.

o ::r
'"

2:
r8:
_.'"
'"3 ::t
0

•

_.

~

C

...
-<

ID

;;:
'0
;;:

o
::r
o

::l

N

~

g

:;:-

o
::J

'"'"

o...,
<
'"c:r...
~

tn

c::

- - - - - - - -. M.(M.) claudiae• M. (M.)? ventricosa• M. (M.) spinosa

A

•
I»
'0
'"

"c::

M.(S.) dyhrenfurthi otai
M. (M.) sp. A'" 0::lM.(M.) postclaudiae
M. (Mol sp. B •Maklaya saraburiensis •M. sp. •Armenina salgirica •A. cf. asiatica •M. pamirica •Neoschwagerina simplex.
Parafusulina kaerimizensi s

::::s
~

Q)

I.::)

n

t::;
I::

-.
~

CD

~

Q)

::s
o

::::s
tn

~

5·
5:
I»

o

0-

Artinskian

Kungur.

UfimianKazan.1 Urals

o

'"
o

~

::r
:E

Yakhtashian

Bolorian

Ku ber.l Mu rgab.1 Tethys

I»

0iJ...

5·
5:

Chang moan

Ch

'"

-.I
00

A A AA

M. (Sol nipponica•'" cc:r_.
o::l ~o~o...,A

Paraf.
Ikaerimiz ' A k i yo s h i

Pseudofusulina ex gr. kraffti
Pamirina (Levenia) leveni
P. (L.) evoluta• P. (P.) darvasica• Misellina (Srevaxina)? sp.

5·
;.~
'"

Mise/lina (M.)
c/audiae

Psf. ex gr'IPamirina(Leve""'M. (Brev.)
kraffti/eveni Idyh. otai

hSlan

Maokou.

Chi na

c

en
en
::JJ

931.

Yakhtash.

Bolori a n,
G),Kubergand. Murgabian

Pamirina (5.5.),0,Brevaxina
Mise/lina (5.5.)

: i

@

G):

979

Levenia,,,

Evolution of Verbeekinidae and Neoschwagerinidae

Misel/ina (Brevaxina)? sp.

0: Misellina (Mol sp. A@: Mise/tina (Mol? ventricosa@: Misellina(Mol sp. B:, ', Mak/aya@, Neoschwag.

,
,
,

,

,
,

, Armenina, VerbeekinaiL

Figure 5. Reconstruction of early phylogeny of some verbeekinid, neoschwagerinid andtheir ancestral fusulinacean genera.

Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea
von Moller, 1878
Family Verbeekinidae Staff
and Wedekind, 1910
Subfamily Misellininae
Miklukho-Maklay, 1958
Genus Misellina Schenck
and Thompson, 1940
Subgenus Brevaxina Schenck
and Thompson, 1940
Type species. - Doliolina compressa Deprat,
1915.

Misellina (Brevaxina) nipponica
Ueno, sp. nov.
Figures 7-1-12

Brevaxina ibukiensis Kobayashi, Zhou et al., 1987,
pI. 6, fig. 13.
Misellina (Brevaxina) aff. otakiensis (Huzimoto),
Vena, 1991, p. 742 (listed).

Material. - Axial section of the holotype;
ASM25082. Axial sections of paratypes;
ASM25083A, B, ASM25084, ASM25085,
ASM25086, ASM25088B. Slightly oblique
axial sections of paratypes; ASM25087,
ASM25088A, ASM25089A. Sagittal sections of paratypes; ASM25089B, ASM25090.
All specimens from Loc. GK9. Some otherspecimens from Loc. GK9 were studied.
Diagnosis. - Small Brevaxina having a
spherical shell with shallow umbilicus, andtightly and skew-coiled inner volutions.
Spirotheca relatively thin. Septa short andstraight. Parachomata small and incipient.
Description. - Shell small for gen us and980

Katsumi Vena

Figure 6. 1-4. Pamirina (Pamirina) darvasica Leven, 1,2: axial sections, ASM25007, ASM25005, 3:
tangential section, ASM25008, 4: sagittal section , ASM25002, x 40. 5: Misellina (Brevaxina) ? sp. , axial
section , ASM25004, X 75. 6- 10. Pamirina (Levenia) /eveni Kobayashi , 6- 9: axial sections, ASM 25019,
ASM25018A , ASM25021 , ASM25025, 10: sagittal section , ASM25028, X 40. 11- 14. Pamirina (Levenia)
eva/uta Sheng and Sun , 11- 13: axial sections, ASM25014, ASM250 13A , ASM25016, 14: sagitta l section ,ASM25010, X 100. (after Ueno , 1991).

almost spherical with slightly umbilicatedaxial regions . Mature specimens having 5 to
6 volutions, 0.82 to 0.97 mm in length and0.78 to 1.07 mm in width. Form ratio ranges
from 0.91 to 1.10, averaging 1.02 for 9 specimens. Holotype specimen of 6 volutions
0.92 mm in length and 0.96 mm in width,
giving a form ratio of 0.96.
Inner 2 volutions tightly and skew-coiled,outer ones planispirally coiled. Shell
expands gradually through growth. Radius
vectors of the first to sixth vol utions of theholotype 0.06,0.10, 0.15, 0.24, 0.36 and 0.50mm, and form ratios 0.83, 0.75, 1.00, 1.00, 1.03

and 0.94, respectively.
Proloculus small and spherical. Its outside diameter varies from 0.045 to 0.065 mm,
averaging 0.054 mm for 9 specimens.
Spirotheca thin and composed of a single
structureless layer in inner few volutions, butof a tectum and fine alveolar keriotheca inouter ones. Thickness of spiro theca of thefirst to sixth vol utions of the holotype 0.005,
0.010,0.015,0.020,0.030 and 0.035 mm.
Septa short, straight and composed of
downw ard deflections of spirothecal elements. Tips of septa slightly solidified.
Septal counts of the second to fifth volutions

931.

Evolution of Verbeekinidae and Neoschwagerinidae

Table 1.

981

Measurements of Misellina (Brevaxina) nippanica, sp. nov.
Radius vector

Specimen

FigureLength

Width

F.R.

D.P.
1

2

3

4

5

6

0.36

0.50

ASM25082
ASM25083A

7-1

0.92

0.96

0.15

0.89

0.95

0.050
0.060

0.10

7-2

0.96
0.94

0.06

2

0.05

0.07

0.14

0.24
0.26

0.41

0.49'

3

ASM25084

7-3

0.93

0.86

1.08

0.045

0.06

0.10

0.16

0.24

0.32

0.46

4

ASM25088B

7-4

0.84

0.80

1.05

0.055

0.06

0.10

0.17

0.28

0.44

5

ASM25085

7-5

0.97

1.07

0.91

0.045

0.07

0.11

0.17

0.27

0.41

6

ASM25083B

7-6

0.89

0.89

1.00

-

0.06

0.11

0.20

0.32

0.45

1

0.58

Thickness of spirotheca

Form ratio
I

2

3

4

5

6

1.03 0.94

0.005

0.010

0.015

0.020

0.030

0.035

0.010

0.015

0.025

0.040

0.040'

0.015

0.025

0.030

0.040

0.040

0.010
0.010

0.020

0.030

0.005

0.015

0.030

0.040
0.040

0.050

0.010

0.020

0.020

0.025

0.030

1

2

3

4

5

1 0.83

0.75

1.00

1.00

6

2

0.80

0.86

0.93

0.89

0.95

3
4

0.83

0.90

0.88

1.00

1.06 0.98

1.10'

0.005
0.010

0.91

0.85

1.06

1.00

0.93

0.010

5

0.69

0.86

0.91

0.82

0.90

6

0.67

1.00

0.91

0.89

0.92

0.87

F.R.: Form ratio, D.P.: Diameter of pro1ocu1us (in mm).

of one sagittal section of paratype (Figure 711) 6,9, 12 and 15.
Parachomata small, semicircular and incipient. They developed beyond the third volution, being about J;l to }:3 as high as chambers.
Remarks. - Small shell size, rudimentary
parachomata and the stratigraphic distribution of the present new species indicate it to
be one of the most primitive species in thesubgenus Brevaxina.
Misellina (Brevaxina) nipponica, sp. nov.
somewhat resembles M. (B.) otakiensis originally described by Huzimoto (1936) from the
Otaki district of the Kwanto Mountains.
However, the former has a smaller form ratio
and less developed parachomata than thelatter.
Misellina (Brevaxina) ibukiensis illustrated
by Zhou et al. (1987) has a more sphericalshell, smaller number of volutions and weaker parachomata than the original specimensof Kobayashi (1957) from the Ibukiyama': 5 Yz vo1ution

Limestone, and is quite identical with thepresent new species.
Misellina (Brevaxina) nipponica, sp. nov.
somewhat resembles M. (B.) minima described by Lin (in Lin et al., 1977) from theLower Permian Chihsia Formation of Guangxi, but differs from the latter in having alarger form ratio.
The present new species can be easily distinguished from Misellina (Brevaxina)
dyhrenfurthi (Dutkevich) in having a smaller
shell and more rudimentary parachomata.
Measurements. - See Table 1.
Occurrence. - Rare in the uppermost part of
the Pamirina (Levenia) leveni Zone.

Misellina (Brevaxina)
dyhrenfurthi otai
Sakaguchi and Sugano, 1966
Figures 7-13-25

Misellina alai Sakaguchi and Sugano, 1966, p. 145147, pI. 1, figs. 1-12.

982

K atsumi Veno

931.

Evolution of Verbeekinidae and Neoschwagerinidae

Misellina (Brevaxina) sp. nov., Ozawa, 1970, pI. 2,
figs. 12-14.
Brevaxina parva Wang and Sun, 1973, p. 155, 174, pI.
2, figs. 1-3; Sun et al., 1983, p. 35, pI. 8, fig. 9(same as pI. 2, fig. I of Wang and Sun, 1973, p.
155,174).
Misellina cf. dyhrenfurthi (Dutkevich), Ozawa, 1975,
pI. 10, figs.4-5 (same as pI. 2, figs. 12-13 of
Ozawa, 1970).
Brevaxina dyhrenfurthi (Dutkevich), Lin et aI., 1977,
p. 86, pI. 26, figs. 12-13.
Misellina dyhrenfurthi (Dutkevich), Ozawa andKobayashi, 1990, pI. 9, figs. 1-2.

Material. - Axial sections; ASM25091
from Loc. NS122, ASM25092, ASM25096,
ASM25101 from Loc. SRI3, ASM25093,
ASM25094 from Loc. NS126, ASM25095
from Loc. SA-AD9, ASM25097 from Loc.
EB8 (float block), ASM25098 from Loc.
NS130, ASM25099 from Loc. NS124, ASM
25100A from Loc. NS 129. Sagittal sections;
ASM25100B from Loc. NS129, ASM 25102
from Loc. NS 131. Additional specimens
were abundantly obtained from the Misellina(Brevaxina) dyhrenfurthi otai Zone.
Description. - Shell almost spherical orslightly quadrate with shallowly depressed
axial regions and convex lateral slopes.
Mature specimens of 6 Yz to 8 volutions 1.03
to 1.58 mm in length and 1.04 to 1.53 mm inwidth. Form ratio ranges from 0.97 to 1.15,
averaging 1.04 for 25 specimens.
Axis of coiling straight through growth
except for inner I or 2 volutions of somespecimens. Chamber height almost constantthroughout length of shell. Radius vectorsof the first to seventh volutions of typical
axial section (Figure 7-18) 0.07,0.10,0.15,
0.23, 0.34, 0.50 and 0.67 mm, and form ratios
0.71,0.80,0.90,1.39,1.32,1.18 and 1.12,983

respectively.
Pro loculus small and spherical. Its outside diameter ranges from 0.045 to 0.110 mm,
averaging 0.070 mm for 36 specimens.
Spirotheca thin and composed of a single
structure less layer in inner 2 or 3 volutions,but consists of a tectum and lower alveolar
keriotheca in outer ones. Thickness of spirotheca of the first to seventh vol utions ofabove-mentioned specimen 0.008,0.010,
0.005,0.010,0.020,0.025 and 0.020 mm.
Septa unfluted and slightly inclined anteriorly. They are composed of downward
deflections of the tectum and extensions of thealveolar keriotheca on either sides of thetectum. Septal counts of the first to eighth
volutions of one illustrated sagittal section(Figure 7-24) 5,10,12,14,14,18,18 and 15,
respectively.
Parachomata small, semicircular to triangular in shape with obtuse apex, and well developed beyond the third volution. They attainabout X to Yz of chamber height.
Remarks. - This subspecies was originallydescribed by Sakaguchi and Sugano(1966) asa distinct species of the genus Misellina fromthe upper part of the Pseudofusulina vulgaris
Zone in the Kaerimizu area. They statedthat the present species resembles Misellina(Brevaxina) compressa (Deprat), the typespecies of the subgenus Brevaxina, in having
an almost spherical shell and well-developed
umbilical depressions even in the outer vol utions of some specimens, but differs from thelatter in having a smaller and more sphericalshell.
Sakaguchi and Sugano (1966) did not referin the original description to the fact that thepresent species closely resembles Misellina~
Figure 7. 1-12. Misellina (Brevaxina) nipponica, sp. nov., 1: axial section of the holotype,
ASM25082, 2-7: axial sections of paratypes, ASM25083A, ASM25084, ASM25088B, ASM25085,
ASM25083B, ASM25086, 8-10: slightly oblique axial sections of paratypes, ASM25088A, ASM25089A,
ASM25087, 11,12: sagittal sections of paratypes, ASM25089B, ASM25090, X 30. 13-25. Misellina(Brevaxina) dyhrenfurthi atai Sakaguchi and Sugano, 13-23: axial sections, ASM25097, ASM25098,
ASM25092, ASM25096, ASM25099, ASM25093, ASM25100A, ASM25101, ASM25095, ASM25094,
ASM25091, 24,25: sagittal sections, ASM25100B, ASM25102, x20. 26-29. Armenina salgirica
Miklukho-Maklay, 26: slightly oblique axial section, ASM25127, 27: tangential section, ASM25128, 28:
sagittal section, ASM25129, 29: axial section, ASM25130; 26,28,29: X 15, 27: X 10.

Katsumi Ueno

984

(Brevaxina) dyhrenfurlhi (s.s.), which was
originally described by Dutkevich (in Likharev el al., 1939) from the Darvas region of the
USSR. In most of the important characters,such as the shell shape, form ratio and development of parachomata they are quite similar
to each other. However, the Akiyoshi specimens differ from the Darvas ones in having alarger shell and more volutions. Hence the
Akiyoshi form, otai, is treated as a subspeciesof Misellina (Brevaxina) dyhrenfurthi (s.1.).
It is considered that Misellina (Brevaxina)
dyhrenfurlhi olai Sakaguchi and Sugano is
more advanced than M. (B.) dyhrenfurthi
dyhrenfurthi (Dutkevich), and the former is a
descendant of the latter.
Misellina (Brevaxina) sp. nov. illustratedby Ozawa (1970) and M. cf. dyhrenfurthi(Dutkevich) of Ozawa (1975) have 8 volutions in the mature stage and a slightly larger
shell than that of M. (B.) dyhrenfurlhi (s.s.).
Ozawa's specimens are almost identical with
Misellina (Brevaxina) dyhrenfurthi otai Sakaguchi and Sugano.
Brevaxina parva originally described by
Sun and Wang (1973) from the Lower Permian Yazi Formation of the Chinling Range
in South China quite resembles M. (B.)
dyhrenfurlhi olai Sakaguchi and Sugano in
essential morphological characters. The former is considered to be a junior synonym of
the latter.
Occurrence. - Abundant in the Misellina(Brevaxina) dyhrenfurlhi olai Zone.
Subgenus Misellina Schenck
and Thompson, 1940
Type species. - Doliolina o valis Deprat,
1915.

Misellina (Misellina) claudiae(Deprat, 1912)
Figures 8-1 - 13
Doliolina claudiae Deprat, 1912, p.44-45, pI. 4, figs.
5-9; Lee, 1933, pI. 2, fig. 5, 5a; Chen, 1934, p.
99-100, pI. 16, figs. 13-20; Huzimoto, 1936, p.

104-105, pI. 21, figs. 4-9.
Verbeekina claudiae (Deprat), Ozawa, 1925a, pI. 2,
figs. 1-2; Ozawa, 1925b, p. 52-53, pI. II, figs. 9II.
Misellina aff. claudiae (Deprat), Kanmera, 1956, pI.
36, fig. 15; Nogami, 1961, p. 169-171, pI. 7, figs.
7-9.
Misellina cf. claudiae (Deprat), Kobayashi, 1957, p.
296-297, pI. I, fig. 19.
Misellina claudiae (Deprat), Toriyama, 1958, p.208211, pI. 39, figs. 1-19; Sakagami, 1958, p. 89-90,
pI. 4, figs. 1-2; Kanuma, 1960, p.64-65, pI. II,
figs. 2-9; Ishizaki, 1962, p. 166-167, pI. 10, figs.
II-I7; Kanmera, 1963, p. 110-112, pI. 14, figs.
7-14; Sheng, 1963, p. 91, 222-223, pI. 28, fig. 15,
pI. 30, figs. 12-19; Sheng, 1966, p. 140-141, pI.
22, fig. 8 (same as pI. 4, fig. 5 of Deprat, 1912, p.
44-45); Takaoka, 1966, p.62-63, pI. 9, fig. 16;
Leven, 1967, p.181-182, pI. 30, figs. 7-8; Igo,
1967, p. 13-14, pI. 7, fig. 9, pI. 8, fig. 7; Douglass,
1967, pI. 6, figs. 17-20; Koike et al. 1968, p.
206-207, pI. 32, figs. 3-4; Ozawa, 1970, pI. 3, figs.
16-18 (18: same as pI. 2, fig. 5 of Lee, 1933);
Choi, .1972, p.637-639, pI. I, figs. I, 2?; Wangand Sun, 1973, pI. 2, figs.4-5, 13, 15; Ozawa,
1975, pI. 10, fig. 9 (same as pI. 3, fig. 16 of Ozawa,1970) ; Ota, 1977, pI. 2, figs. 3-4; Lin et al.,
1977, p. 85, pI. 26, fig. 5; Kobayashi, 1977, pI. 2,
figs. 15-16; Liu et al., 1978, p. 84, pI. 19, fig. 6;
Minato et al., 1979, pI. 41, fig. 2(same as pI. I, fig.
2 of Choi, 1972, p. 637-639); Hasegawa et al.,
1979, pI. 77, figs. 4-5,8-10, 12-14, 16-22; Lin etal., 1979, pI. 4, figs. 8-12; Xie, 1982, p. 66, pI. 32,
figs. 4-5; Leven, 1982, pI. 2, figs. 4, 7 (7: same as
pI. 30, fig. 7 of Leven, 1967, p. 181-182); Wanget al., 1982, p. 10 I, pI. 28, figs. 6-7; Sun et al.,1983, p. 34, pI. 9, fig. 6; Zhou, 1984, p. 118-119,
pI. I, figs. 14-15; Zhou and Zhang, 1984, pI. 2,
figs. 8-11 ; Chen, 1984, p. 65, pI. 18, figs. 5-7 ;Xia and Zhang, 1985, p. 150, pI. 29, fig. 7; Sun
and Zhang, 1985, p.505, pI. I, figs. 2-5; Han,
1985, p. 683, pI. I, fig. 18; Yang, 1985, pI. 2, figs.
1-2; Wang and Tang, 1986, pI. 2, fig. 18; Zhang
and Li, 1987, p.405-406, p1.3, figs. 10, 20, 24;
Sun and Zhang, 1988, pI. 4, figs. 4-6, 8-10;
Ozawa and Kobayashi, 1990, pI. 9, figs. 5-6 ;Tong et al., 1990, pI. 4, fig. 5.
Misellina (Misellina) claudiae (Deprat), Toriyama,
1975, p.55-57, pI. 13, figs. 1-2; Ishibashi, 1984,
p. 221-222, pI. 31, figs. 6-15.
Misellina minor (Deprat), Chen and Yang, 1978, p.
107, pI. 26, fig. 10.
? Misellina claudiae (Deprat), Kalmykova, 1967, p.
215, pI. 30, figs. 13-14; Chen and Yang, 1978, p.
107, pI. 26, figs. 8-9.
non Misellina claudiae (Deprat), Hanzawa and Murata, 1963, pI. 6, figs. 7-8; Han, 1976, p.62, pI.

931.

Evolution of Verbeekinidae and Neoschwagerinidae

15, figs. 10-13.

Material. - Axial sections of megalospheric
form; ASM25103 from Loc. MI51, ASM
25104 from Loc. MI9, ASM25105 from Loc.
MI39, ASM25106 from Loc. AL35.5, ASM
25107 A,B from Loc. MI23, ASM25108 from
Loc. SA31, ASM25109 from Loc. MI43.
Sagittal sections of megalospheric form;
ASM 25110 from Loc. MI8, ASM25111 from
Loc. MI58, ASM25112A from Loc. MI60.
Axial sections of microspheric form; ASM
25113 from Loc. MIl3, ASM25114 from Loc.
MI28. In addition, many specimens from theMisellina (M.) claudiae Zone were studied.
Description. - Megalospheric shell smalland short ellipsoidal with broadly rounded tosomewhat umbilicated axial regions and convex lateral slopes. Microspheric shell slightly larger than megalospheric one, and subglobular to fusiform with broadly rounded tobluntly pointed poles. Mature shell consistsof 8 to 9 volutions in megalospheric form, but
10 to 11 volutions in microspheric one.
Length and width of megalospheric form 1.68
to 2.73 mm and 1.38 to 2.23 mm, respectively.
Form ratio ranges from 1.18 to 1.38, averaging 1.27 for 23 specimens. Microsphericspecimens 2.05 to 2.29 mm in length and 1.73
to 2.48 mm in width. Form ratio varies from
1.16 to 1.43, averaging 1.28 for 7 specimens.
Axis of coiling straight throughout inmegalospheric form. Inner few volutions
tightly and skew coiled in microspheric form.
Height of chamber almost constant in bothforms. Radius vectors of the first to eighth
volutions of typical megalospheric form (Figure 8-6) 0.11,0.16,0.24,0.34,0.46,0.60,0.77
and 0.93 mm, and form ratios 0.82, 1.00, 1.08,
1.12,1.20,1.16,1.10 and 1.24, respectively.
Proloculus of megalospheric form large forgenus and spherical to somewhat irregular.
Its outside diameter ranges from 0.105 to
0.220 mm, averaging 0.135 mm for 30 specimens. Proloculus of microspheric form minute and measuring 0.030 to 0.070 mm in outside diameter, averaging 0.048 mm for 8 speci-

985

mens.
Spirotheca thin and composed of a single
structureless layer in inner 2 volutions, butmoderate in thickness and composed of atectum and alveolar keriotheca beyond thethird volution. Lower surface of spirotheca
uneven in some forms. Thickness of spirotheca of the first to eighth volutions ofabove-mentioned megalospheric specimen
0.010,0.020, 0.030, 0.040, 0.045, 0.075, 0.075
and 0.040 mm.
Septa numerous, bending anteriorly andcomposed of downward deflections of tectum
and anterior and posterior extensions of keriotheca. Septal counts of the first to eighth
volutions of one illustrated sagittal section ofa megalospheric form (Figure 8-10) 7, 10, II,
16, 15, 18, 21 and 26, respectively.
Low and broad parachomata well developed in all volutions except for the first one
in megalospheric form and inner 2 or 3 volutions in microspheric one. Height of parachomata about Ys to Yz that of chambers.
Remarks. - Dimorphism is well observedin the Akiyoshi specimens prepared for thisstudy. The microspheric form is characterized by a larger shell, more volutions andsmaller proloculus than the megalosphericform. Moreover, the inner volutions of the
microspheric form are compactly and skew
coiled.
The Akiyoshi specimens quite agree withthe original ones of Deprat (1912) fromYunnan, and no further comparison is necessary.
Occurrence.-Abundant in the lower partof the Misellina (M.) claudiae Zone and rarein the middle part of the same zone.

Misellina (Misellina) postclaudiae
Veno, sp. nov.
Figures 8-15-18

Material. - Axial section of the holotype;
ASM25120 from Loc. MI29. Axial sectionsof paratypes; ASM25119 from Loc. AL61,
ASM25121 from Loc. AL67. Sagittal sec-

Katsumi Vena
986

931.

Evolution of Verbeekinidae and Neoschwagerinidae

tion of paratype; ASM25122 from Loc. AL
67. Some additional specimens from Loc.
AL67 were studied.
Diagnosis. - Large Misellina having a subglobular to short ellipsoidal shell withrounded polar ends, almost straight axis ofcoiling and numerous volutions. Inner few
volutions rather tightly coiled. Spirothecathin, with slightly undulated lower surface insome case. Septa slightly bending anteriorly.
Well-developed parachomata beyond thethird volution.
Description. - Shell large for genus and subglobular to short ellipsoidal with rounded
axial regions and broadly convex lateralslopes. Mature specimens having 10 to 11
volutions, 2.63 to 2.96 mm in length and 2.13
to 2.65 mm in width, giving form ratios of
1.09 to 1.24. Holotype specimen of 10 Y2
volutions 2.96 mm in length and 2.38 mm inwidth, giving a form ratio of 1.24.
Axis of coiling almost straight throughout
growth. Inner few volutions tightly coiled,
but later ones expanding gradually. Radius
vectors of the first to tenth volutions of theholotype 0.07,0.12,0.16,0.23,0.31,0.42,0.56,
0.71, 0.93 and 1.14 mm, and form ratios 0.71,
0.87,0.94, 1.22, 1.39, 1.50, 1.38, 1.44, 1.29 and
1.25, respectively.
Proloculus small and spherical. Its outside diameter ranges from 0.050 to 0.080 mm,
averaging 0.065 mm for 5 specimens.
Spirotheca thin to medium, and composedof a single structureless layer in inner 3 volutions, but of a thin tectum and rather thickalveolar keriotheca in outer ones. Lower
surface of spirotheca slightly undulated in

987

middle volutions of some specimens. Thickness of spirotheca of the first to tenth volutions of the holotype 0.010, 0.010, 0.015,
0.020, 0.020, 0.050, 0.045, 0.040, 0.065 and
0.040 mm.
Septa slightly bending anteriorly and composed of downward deflections of spirothecal
elements. Septal counts of the third to seventh vol utions of ill ustrated sagittal section ofparatype II, 13, 16, 17 and 19, respectively.
Parachomata well developed in all volutions except for inner 2, and semicircular to
triangular in transverse section. Height ofparachomata about % to Y2 as that of chambers.
Remarks. - Misellina (M.) postclaudiae, sp.
nov. is somewhat similar to the microspheric
form of M. (M.) c1audiae (Deprat). However, the former can be distinguished from thelatter in having a slightly larger shell and analmost straight axis of coiling even in thecompactly coiled inner volutions.
The present new species also resembles
Maklaya cutalensis (Leven) from the Kubergandian of the Pamir in general shell morphology. However, the former can be easily
distinguished from the latter in having no
primary transverse septula.
In the present new species, low and broad
protrusions of the lower surface of the spirotheca are observed in the middle volutions ofsome specimens. However, these are too
poorly shown to recognize as the primarytransverse septula.
Measurements. - See Table 2.
Occurrence. - Rare in the middle part of
the Misellina (M.) claudiae Zone.

+- Figure S. 1-13. Misellina (Misellina) claudiae (Deprat), I-S: axial sections of megalospheric form,
ASM25103, ASM251 04, ASM25105, ASM25106, ASM25107 A, ASM25 108, ASM25107B, ASM251 09, 9-11 :sagittal sections of megalospheric form, ASM2511O, ASM25111, ASM25ll2A, 12a, 13: axial sections of
microspheric form, ASM25ll3, ASM25ll4, 12b: enlargement of 12a; 1-11, 12a, 13: X 10, 12b: X 20. 14:
Misellina (Misellina) sp. A, axial section, ASM25l23, X 20. 15-lS. Misellina (Misellina) postclaudiae, sp.
nov., 18: axial section of the holotype, ASM25120, 15: sagittal section of paratype, ASM25l22, 16,17:
axial sections of para types, ASM25l19, ASM25l21; 15: X 10, 16-18: X 15. 19-24. Misellina (Misellina)?
ventricosa, sp. nov., 23: axial section of the holotype, ASM25Il5A, 19,22: axial sections of paratypes,
ASM25ll5B, ASM25ll6A, 20, 21, 24: slightly tangential sections of paratypes, ASM25ll7, ASM25ll6B,
ASM25ll8, X 15.

Katsumi Ueno

988
Table 2.

Measurements of Misellina (M.) postclaudiae, sp. nov.

Specimen

FigureLength

Width

D.P.

F.R.

1

ASM25119

8-16

2.63

2.42

1.09

0.050 ?

2

ASM25121

8-17

2.63

2.13

1.23

0.075

3

ASM25120

8-18

2.96

2.34

1.24

0.075

Radius vector
1

2

3

4

5

6

7

8

9

10

1

0.08

0.12

0.17

0.26

0.39

0.51

0.67

0.82

1.03

1.25

2

0.08

0.13

0.18

0.26

0.36

0.47

0.63

0.81

1.00

-

3

0.07

0.12

0.16

0.23

0.31

0.42

0.56

0.71

0.93

1.14

8

9

10

Form ratio
1

2

3

4

5

6

7

1

0.56

0.71

0.89

1.13

1.00

1.04

1.07

1.18

1.06

-

2

0.60

0.69

0.79

1.04

1.32

1.50

1.35

1.27

1.27

-

3

0.71

0.87

0.94

1.22

1.39

1.50

1.38

1.44

1.29

1.25

8

9

10
0.065

Thickness of spiro theca
1

2

3

4

5

6

7

1

0.010

0.020

0.025

0.040

0.050

0.050

0.075 ?

0.070

0.080

2

0.010

0.Q15

0.020

0.030

0.045

0.060

0.070

0.075

0.060

-

3

0.010

0.010

0.Q15

0.020

0.020

0.050

0.045

0.040

0.065

0.040

F.R.: Form ratio, D.P.: Diameter of pro loculus (in mm).

Misellina (Misellina)spinosa Han, 1976
Figure 9-15

Misellina spinosa Han, 1976, p. 61-62, pI. 11, figs. 911 ; Yang, 1985, pI. 3, fig. 1.

Material. -Axial section; ASM25112B
from Loc. MI60.
Description. - Shell small, short ellipsoidal
with rounded axial regions, and consisting of
6 volutions. Length 0.86 mm and width 0.70mm, giving a form ratio of 1.23. Inner 2
volutions skew coiled, but outer ones
planispirally coiled. Radius vectors of thefirst to sixth volutions 0.04,0.09,0.12,0.20,

0.28 and 0.39 mm, and form ratios 0.75,0.94,
1.13,1.08,1.18 and 1.19, respectively. Spirotheca thin and composed of a single structureless layer in inner 3 volutions, but of a tectumand fine alveolar keriotheca in outer ones.
Thickness of spiro theca of the first to sixth
volutions 0.005,0.005,0.010,0.015,0.020 and
0.025 mm. Parachomata small and rudimental, developed merely in outer 3 volutions.
Remarks. - Misellina (M.) spinosa wasoriginally described by Han (1976) from theLower Permian Sanmianjing Formation of
Neimenggu (Inner Mongolia). It has a thinner spirotheca and less developed parachomata than any other known species of the931.
Table 3.

Evolution of Verbeekinidae and Neoschwagerinidae

989

Measurements of Misellina (M.)? ventricosa, sp. nov.

Specimen

FigureLength

Width

F.R.

D.P.

I

ASM25115B

8-19

2.08

1.89

1.10

0.050

2

ASM25116A

8-22

2.55

2.15

1.19

0.075

3

ASM25115A

8-23

2.87

2048

1.16

0.065

Radius vector
I

2

3

4

5

6

7

8

9

10

II

I

0.05

0.10

0.16

0.24

0.37

0.53

0.71

0.89

2

0.08

0.13

0.18

0.25

0.38

0.51

0.67

0.85

1.06

3

0.06

0.11

0.14

0.20

0.29

0041

0.55

0.73

0.92

1.10

1.29

9

10

II

Form ratio
I

2

3

4

5

6

7

8

I

0.80

0.60

0.81

0.92

1.08

1.07

1.10

1.12

2

1.30

1.00

1.05

1.27

1.28

1.28

1.29

1.33

-

3

0.50

0.52

0.57

1.15

1.17

1.26

1.25

1.23

1.21

1.20

1.14

9

10

II

0.055

0.055

Thickness of spirotheca

I

I

2

3

4

5

6

7

8

0.005

0.010

0.020

0.040

0.050

0.060

0.060

0.060

0.030

0.060

0.050

0.040

0.050

0.075

0.065

0.080

2

0.010

0.010

0.010

0.025

0.040

3

0.005

0.010

0.010

0.020

0.030

F.R.: Form ratio, D.P.: Diameter of proloculus (in mm).

subgenus Misellina.
Occurrence. - Rare in the lower part of theMisellina (M.) claudiae Zone.

Misellina (Misellina)?
ventricosa Ueno, sp. nov.
Figures 8-19 - 24

Material. - Axial section of the holotype;
ASM25115A from Loc. MIl7. Axial sections of paratypes; ASM25115B, ASM
25116A from Loc. MIl7. Slightly tangential
sections of paratypes; ASM25116B, ASM
25117 from Loc. MIl7, ASM25118 from Loc.
KS 127. Some additional specimens from
Loc. MII7 were studied.

Diagnosis. - Small verbeekinid having
intermediate shell characters between the genera Misellina and Armenina. Shell shortellipsoidal with broadly rounded polar ends.
Inner volutions tightly and skew coiled, outerones rather loosely coiled. Spirotheca moderately thick, with fine alveolar keriotheca.
Parachomata prominent and rather high.
Description. - Shell small, short ellipsoidal
with broadly rounded axial regions and convex lateral slopes. Mature specimens having
8 Yz to II volutions, 2.08 to 2.87 mm inlength and 1.89 to 2.48 mm in width. Formratio ranges from l.l 0 to 1.27, averaging 1.19
for 6 specimens. Holotype specimen of II
volutions 2.87 mm in length and 2.48 mm in

990

Katsumi Ueno

width with a form ratio of 1.16.
Inner 2 or 3 volutions tightly and skew
coiled with a short axis of coiling. Outer
volutions expanding gradually and becoming
loose. Radi us vectors of the first to eleventh
volutions of the holotype 0.06, 0.11,0.14, 0.20,
0.29,0.41,0.55,0.73,0.92,1.10 and 1.29 mm,
and form ratios 0.50,0.52,0.57, 1.15, 1.17,
1.26, 1.25, 1.23, 1.21, 1.20 and 1.14, respectively.
Proloculus small and spherical with outside diameters of 0.045 to 0.075 mm, averaging 0.059 mm for 4 specimens.
Spirotheca moderately thick and composedof a single structureless layer in inner 3 volutions, but of a tectum and fine alveolar keriotheca in outer ones. Thickness of spirotheca of the first to eleventh vol utions of theholotype 0.005, 0.010, 0.010, 0.020, 0.030,
0.050, 0.075, 0.065, 0.080, 0.055 and 0.055 mm.
Parachomata prominent and rather high,and first appear in the third vol ution. Parachomata attain nearly Yz to % of chamberheight.
Remarks. - The present new species is characterized by tightly and skew-coiled inner
volutions, rather loosely coiled outer ones,
and high and prominent parachomata.
These biocharacters are somewhat heretical in
the genus Misellina, and rather common in
the genus Armenina, although its shell shapeis slightly different from that of typical Armenina. From the morphological point ofview, the present new species is considered to
be transitional between the genera Misellina
and Armenina, and here assigned tentatively
to the genus Misellina.
Misellina (M.) ? ventricosa, sp. nov. can be
distinguished easily from M. (M.) postclaudiae, sp. nov. in having more tightly
coiled inner vol utions, thicker spirotheca forcorresponding volutions and finer alveolar
keriotheca.
This new species somewhat resembles Armenina sa/girica Miklukho-Maklay. However, the latter has a thinner spirotheca, more
spherical shell, more tightly coiled inner volu-

tions and more prominent parachomata thanin the former.
Measurements. - See Table 3.
Occurrence. - Rare in the lower part of theMisellina (M.) claudiae Zone.
Mise/lina (Misellina) sp. A
Figure 8-14

Materia/.-Axial section; ASM25123 from
Loc. NS120.
Descnption. - Shell small, short ellipsoidal
with rounded axial regions and inflated ventral sides, and having 9 volutions. Length
1.80 mm and width 1.55 mm with a form ratioof 1.16. Inner 2 vol utions tightly coiled witha slightly rotated axis of coiling. Radius
vectors of the first to ninth vol utions 0.08,
0.11,0.14,0.20,0.26, 0.32, 0.48, 0.63 and 0.81 ?
mm, and form ratios 0.63, 0.71, 0.93, 1.00,
1.15, 1.25, 1.08, 1.03 and 1.05, respectively.
Proloculus small and spherical, being 0.075
mm in outside diameter. Spirotheca composed of a single structureless layer in inner 2
volutions, but of a tectum and alveolar keriotheca beyond the third one. Thickness of
spirotheca of the first to ninth volutions 0.005,
0.005,0.010,0.020,0.020,0.020,0.025,0.040
and 0.030 mm. Parachomata prominent andrather high, attaining about % the height ofchambers. They are observed beyond thesecond volution.
Remarks. - Misellina (M.) sp. A is considered to be one of the most primitive forms inthe subgenus Misellina. This species is somewhat similar to Misellina (M.) parvicostataand M. (M.) minor, both originally describedby Deprat (l915). The specific identification, however, is postponed until additional materials become available.
Occurrence. - Rare in the middle part of
the Misellina (Brevaxina) dyhrenfurthi otai
Zone.
Misellina (Misellina) sp. B
Figures 9-11-13

931.

Evolution of Verbeekinidae and Neoschwagerinidae

Material. -Axial sections; ASM25124,
ASM25125 from Loc. TAI96, ASM25126
from Loc. NS159. Some other specimensfrom Loc. T A 196 were studied.
Description. - Shell small and short ellipsoidal with rounded polar regions. Mature
specimens of 9 to 10 Yz volutions 2.08 to 2.88mm in length and 1.66 to 2.45 mm in width.
Form ratio varies from 1.17 to 1.25, averaging
1.22 for 4 specimens. Shell expands gradually throughout growth. Axis of coilingstraight. Radi us vectors of the first to ninth
volutions of typical axial section (Figure 912) 0.05,0.11,0.15,0.22,0.28,0.40,0.51,0.64
and 0.78, and form ratios 1.11, 1.00, 1.07, 1.12,
1.25, 1.27, 1.23, 1.25 and 1.37, respectively.
Proloculus small and spherical, being 0.060 to
0.095 mm in outside diameter, averaging 0.078
mm for 5 specimens. Spiro theca thin andcomposed of a single structureless layer ininner 3 volutions, but of a tectum and alveolar keriotheca in outer ones. In some specimens, lower surface of spirotheca slightly
undulated and forms low and broad downward protrusions roughly corresponding toparachomata. Thickness of spiro theca of thefirst to ninth volution of above-mentioned
specimen 0.010, 0.010, 0.0 I 0, 0.020, 0.015,
0.020, 0.040, 0.040 and 0.040 mm. Parachomata massive, broad and semicircular intransverse section, and developed beyond thesecond volution.
Remarks. - The present species somewhat
resembles Maklaya saraburiensis Kanmera
and Toriyama, which accompanies the former. However, Misellina (M.) sp. B has no
primary transverse septula like that observedin the latter, although incipient downward
protrusions are developed in some specimensof M. (M.) sp. B.
Occurrence. - Rare in the middle part of
the Misellina (M.) claudiae Zone.
Subfamily Verbeekininae Staff
and Wedekind, 1910
Genus Armenina Miklukho-Maklay, 1955

991

Type species. - Armenina karinae Miklukho-Maklay, 1955.
Armenina salgirica
Miklukho-Maklay, 1955
Figures 7-26-29

Armenina sa/girica Miklukho-Maklay, 1955, fig. 1 ;Miklukho-Maklay, 1957, p. 120, pI. 4, fig. 3 (sameas fig. 1 of Miklukho-Maklay, 1955); Leven,
1965, p. 141, pI. 4, fig. 3, pI. 6, figs. 1-3; Leven,
1967, p.203, pI. 35, figs. 2-3, pI. 38, figs. 2, 6, pI.
39, fig. 4; Ozawa, 1970, pI. 2, fig. 6 (same as pI.
38, fig. 2 of Leven, 1967, p. 203); Yang, 1985, pI.
2, fig. 29.

Materia/.-Axial section; ASM25130 from
Loc. T AI96. Slightly oblique axial section;
ASM25127 from Loc. NS159. Sagittal section; ASM25129 from Loc. TAI96. Tangential section; ASM25128 from Loc. NS
159. Some additional specimens from Loc.
NS159 were studied.
Description. - Shell small for genus andsubspherical with broadly rounded polar
regions. Mature specimens having 9 to 11
volutions, 2.58 to 3.00 mm in length and 2.09
to 2.93 mm in width. Form ratio of welloriented axial section (Figure 7-29) 1.16.
Inner 2 or 3 volutions tightly and skew
coiled with a short axis of coiling. Beyond
the third or fourth volution, coiling becomesloose as compared with inner ones. Radius
vectors of the first to tenth vol utions of abovementioned axial section 0.07,0.09,0.12,0.17,
0.26,0.35,0.47,0.64,0.85 and 1.07 mm, andform ratios 0.53,0.72,0.92, 1.18, 1.29, 1.29,
1.28, 1.24, 1.15 and 1.14, respectively.
Proloculus small and 0.060 to 0.080 mm inoutside diameter, averaging 0.070 mm for 5
specimens.
Spirotheca thin and composed of a single
structureless layer in inner 3 volutions, but ofa tectum and fine alveolar keriotheca in outerones. Lower surface of spirotheca slightly
undulated in middle volutions of some specimens. Thickness of spiro theca of the first totenth volutions of typical axial section

992

Katsumi Uena

• . e,

931.

Evolution of Verbeekinidae and Neoschwagerinidae

mentioned above 0.005,0.005,0.010, 0.Ql5,
0.020, 0.020, 0.025, 0.035, 0.035 and 0.040 mm.
Septa slightly bending anteriorly and consist of downward deflections of tectum andanterior and posterior extensions of keriotheca. Septal counts of the third to ninth
volutions of illustrated sagittal section (Figure 7-28) 9, 11, 14, 14, 17, 18 and 20 ?
Parachomata rather narrow and high, reaching about % the height of chambers, andbegin to occur in the third volution.
Remarks. - Armenina sa/girica was introduced by Miklukho-Maklay (l955), based ona specimen from the Crimea, USSR. The
Akiyoshi specimens are quite identical withthe original one.
Leven (l967) also reported Armenina salgirica Miklukho-Maklay from the Kubergandian and lower Murgabian of the Pamir.
His specimens have a larger shell and more
volutions than those of the Crimean andAkiyoshi ones.
Occurrence. - Rare in the middle part of
the Misellina (M.) claudiae Zone.

993

and skew coiled, remaining volutions planispirally and rather loosely coiled. Radius
vectors of the first to eighth volutions 0.09,
0.13,0.19,0.25,0.37,0.52,0.67 and 0.81 mm,
and form ratios 0.72,0.88, 1:03, 1.04, 1.07,
1.06, 1.04 and 1.15, respectively. Proloculus
small, being 0.045 mm in outside diameter.
Spirotheca composed of a single structurelesslayer in the first volution, but of a tectum andfibrous alveolar keriotheca beyond the second
one. Lower surface of spirotheca uneven.
Parachomata low, semicircular and welldeveloped beyond the third volution.
Remarks. - Armenina asiatica was originally described by Leven (1967) from the
Kubergandian of the Pamir. The Akiyoshi
specimen is very similar to the Pamir ones
except for having a smaller shell and fewer
volutions, and is considered to be an immature individual of Armenina asiatica
Leven. The specific name, however, hasbeen left tentative until sufficient materials areavailable.
Occurrence. - Rare in the middle part of
the Misellina (M.) claudiae Zone.

Armenina cf. asiatica Leven, 1967
Figure 9-16

CompareArmenina asiatica Leven, 1967, p.204, pI. 38, fig. 3,
pI. 39, figs. 2-3; Ozawa, 1970, pI. 2, fig. 9 (sameas pI. 39, fig. 3 of Leven, 1967, p. 204); Tien,
1989, pl.20, figs. 1-2.

Material.-Axial section; ASM25131 from
Loc. TA196.
Description. - Shell small for gen us andsubspherical with a slightly longer axis ofcoiling. Specimen having 8 volutions, 1.93
mm in length and 1.55 mm in width with aform ratio of 1.25. Inner 2 volutions tightly

Family Neoschwagerinidae Dunbar
and Condra, 1927
Subfamily Neoschwagerininae
Dunbar and Condra, 1927
Genus Maklaya Kanmera
and Toriyama, 1968
Type species. - Cancellina pamirica Leven,1967.

Maklaya saraburiensis Kanmera
and Toriyama, 1968
Figures 9-1 - 4~ Figure 9. 1-4. Maklaya saraburiensis Kanmera and Toriyama, la, 2, 3: axial sections, ASM25132,
ASM25134, ASM25133, 4: sagittal section, ASM25135, Ib: enlarged part of la; la, 2-4: X 10, Ib: x40.
5-10. Maklaya pamirica (Leven), 5a, 6: axial sections, ASM25136, ASM25137, 7,8: tangential sections,
ASM25139, ASM25138, 9, lOa: sagittal sections, ASM25140, ASM25141, Sb, lOb: enlarged parts of5a andlOa; 5a,6-9,IOa: X 10, Sb,IOb: X 40. 11-13: Misellina (Misellina) sp. B, axial sections, ASM25125,
ASM25124, ASM25126, X 10. 14: Maklaya sp., axial section, ASM25142, 14a: X 10, 14b: enlargement of
14a, X 20. 15: Misellina (Misellina) spinosa Han, axial section, ASM25112B, X 30. 16: Armenina cf.
asiatica Leven, axial section, ASM25131, x20.

994

Katsumi Vena

Maklaya saraburiensis Kanmera and Toriyama, 1968,
p.41-43, pI. 4, figs. 17-20; Ozawa, 1970, pI. 4,
figs. 14-15 (15: same as pI. 4, fig. 17 of Kanmera
and Toriyama, 1968, p. 41-43); Toriyama, 1975,
pI. 18, figs. 21-24(same as pI. 4, figs. 17-20 of
Kanmera and Toriyama, 1968, p.41-43).
Neosc!Jwagerina saraburiensis (Kanmera and Toriyama), Han, 1985, p. 685, pI. 2, figs. 14-15.

Material.-Axial sections; ASM25132
from Loc. NS159, ASM25133, ASM25134
from Loc. T A 196. Sagittal section; ASM
25135 from Loc. NS 159. Some additional
specimens from Locs. NS 159 and T A 196
were studied.
Description. - Shell small for genus, subspherical with rounded polar ends and convex lateral slopes. Mature specimens having
8 to 9 volutions, 1.90 to 2.20 mm in lengthand 1.68 to 2.05 mm in width. Form ratio
ranges from 1.13 to 1.29, averaging 1.18 for 4
specimens.
Shell expands gradually throughout
growth. Inner few vol utions tightly coiled
with a short and slightly rotated axis of coiling. Radi us vectors of the first to eighth
volutions of typical axial section (Figure 9-2)
0.09,0.14,0.22,0.33,0.43,0.57,0.77 and 0.88mm, and form ratios 0.78,0.93,0.95, 1.02,
1.09,1.11,1.06 and 1.17, respectively.
Proloculus small and spherical. Its outside diameter ranges from 0.060 to 0.110 mm,
averaging 0.088 mm for 6 specimens.
Spirotheca composed of a tectum and alveolar keriotheca. Lower surface of spirotheca
undulatory with short but broad downward
protrusions of keriothecal element, forming
rudimentary primary transverse septula.
They are observed beyond the third volution,and occur immediately above each parachomata joining with tops of parachomata
adj acent to septa. Thickness of spirotheca of
the first to eighth volutions of abovementioned specimen 0.010,0.020,0.020,0.035,
0.030,0.035,0.080 and 0.030 mm.
Septa consist of downward deflections oftectum and anterior and posterior extensionsof keriotheca. Septal counts of the second toeighth volutions of illustrated sagittal section(Figure 9-4) 6,9,12,13,15,18 and 21. No
axial septula developed.
Parachomata observed beyond the third
volution, being nearly Yz as high as chambers
in inner volutions, but a little more than Yz
to about % as high in outer ones.
Remarks. - Maklaya saraburiensis was
originally described by Kanmera and Toriyam a (1968) from the Rat Buri Limestone in
the Khao Phlong Phrab area, central
Thailand, and is considered to be the mostprimitive species of the genus. The Akiyoshi
specimens quite agree with the T!Jailand ones.
Occurrence. - Rare in the middle part of
the Misellina (M.) claudiae Zone.

Maklaya pamirica (Leven, 1967)
Figures 9-5-10

Neosc!Jwagerina sp. (n. sp.), Sakaguchi, 1963, p. 115116, pI. 8, figs. 7-10.
Cancellina pamirica Leven, 1967, p. 186-187, pI. 32,
figs. 1, 3; Rozovskaya, 1975, pI. 29, fig. 6 (sameas pI. 32, fig. 1 of Leven, 1967, p. 186-187) ;Leven, 1982, pI. 3, fig. 3 (same as pI. 32, fig. 1 ofLeven, 1967, p. 186-187); Sun and Zhang, 1988,
pI. 4, fig. 21.
Maklaya pamirica (Leven), Kanmera and Toriyama,
1968, p. 34-37, pI. 4, figs. 1-16; Ozawa, 1970, pI.
4, figs. 12-13 (same as pI. 4, figs. 1, 7 of Kanmera
and Toriyama, 1968, p. 34-37) ; Toriyama, 1975,
pI. 18, figs. 16-20(same as pI. 4, figs. 1-2, i 1 ofKanmera and Toriyama, 1968, p. 34-37); Kahler
and Kahler, 1979, p. 249-250, pI. S, fig. 6; Ishibashi, 1984, p. 222-223, pI. 31, fig. 16; Yang, 1985,
pI. 2, fig. 3; Loeb1ich and Tappan, 1988, pI. 300,
figs. 8-11 (8-9: same as pI. 4, figs. 1, II of Kanmera and Toriyama, 1968, p. 34-37; 10-11:
same as pI. 32, figs. 1,3 of Leven, 1967, p. 186187); Ozawa and Kobayashi, 1990, pI. 9, figs.
10-11; Fan el al., 1990, pI. 7, figs. 19,27.

Material. - Axial sections; ASM25136 andASM25137. Sagittal sections; ASM25140
and ASM25141. Tangential sections; ASM
25138 and ASM25139. All specimens from
Loc. NS173.
Description. - Shell medi urn to large forgenus and almost spherical with rounded
periphery. Mature specimens having 10 to931.

Evolution of Verbeekinidae and Neoschwagerinidae

13 volutions, 2.75 to 3.60 mm in length and2.60 to 3.25 mm in width, giving form ratiosof 1.06 to 1.11.
Shell expands gradually throughout
growth. Inner few volutions tightly coiled
with a short and slightly rotated axis of coiling. Radius vectors of the first to thirteenth
volutions of illustrated axial section (Figure9-5) 0.04,0.05,0.09,0.13,0.18,0.27,0.39,0.54,
0.73,0.92, 1.19, 1.42 and 1.69 mm, and form
ratios 0.71,0.60,0.53,0.65,0.94,1.08, 1.08,
1.07, 1.04, 1.05, 1.02, 1.01 and 1.01, respectively.
Proloculus small and spherical, being 0.045
to 0.060 mm in outside diameter.
Spirotheca rather thick and composed of atectum and alveolar keriotheca. Short but
broad, fan-shaped rudimentary primary transverse septula well developed beyond thefourth or fifth volution. They occur immediately above each parachomata joining withtops of parachomata adjacent to septa.
Thickness of spirotheca of the first to thirteenth volutions of above-mentioned specimen 0.005,0.005,0.005,0.010,0.015,0.040,
0.040, 0.070, 0.065, 0.070, 0.115, 0.060 and
0.060 mm.
Septa composed of downward deflectionsof spirothecal elements. Septal counts of thethird to tenth volutions of one illustrated
sagittal section (Figure 9-9) 7?, 9?, 12, 16,
19,22,25 and 25. No axial septula present.
Parachomata semicircular or triangular intransverse section, and first appear in the third
volution. They attain about Yz as high aschambers.
Remarks. - Maklaya pamirica (Leven)differs from M. saraburiensis Kanmera andToriyama in having a larger shell, more volutions and more developed primary transverse
septula.
Neoschwagerina sp. (n. sp.) of Sakaguchi(1963) from the Tamba district has a sphericalshell, relatively thick spirotheca, short andbroad primary transverse septula and no axial
septula. Judging from these shell characters,
the Tamba specimens should be assigned to995

Maklaya pamirica (Leven).
Occurrence. - Rare in the upper part of theMisellina (M.) claudiae Zone.
Maklaya sp.
Figure 9-14

Material. - Axial section; ASM25142 from
Loc. NS159.
Description. - Shell small for genus andspherical with convex lateral slopes. Specimen consisting of 10 volutions, 2.10 mm inlength and 2.00 mm in width, giving a form
ratio of 1.05. Inner few vol utions tightly
coiled with a short and slightly rotated axis ofcoiling. Radius vectors of the first to ninth
volutions 0.06,0.09,0.14,0.20,0.27,0.37,0.49,
0.64 and 0.81 mm, and form ratios 0.42, 0.56,
0.74,0.82,0.93, 1.07, 1.00, 1.00 and 1.06,
respectively. Proloculus small and spherical,
being 0.050 mm in outside diameter. Spirotheca rather thin and composed of a tectumand keriotheca. Thickness of spirotheca of
the first to ninth volutions 0.005,0.010,0.015,
0.035, 0.020, 0.020, 0.035, 0.040 and 0.040 mm.
Short but broad, fan-shaped incipient primary transverse septula occur facing eachparachomata beyond the fourth volution.
They become inconspicuous in outer few
volutions. Parachomata massive and attainabout % the height of chambers.
Remarks. - The present species somewhat
resembles Maklaya saraburiensis Kanmera
and Toriyama, but differs from the latter inhaving a smaller form ratio and more massive
parachomata.
Occurrence. - Rare in the middle part of
the Misellina (M.) claudiae Zone.
Genus Neoschwagerina Yabe, 1903
Type species. - Schwagerina craticulifera
Schwager, 1883.

Neoschwagerina simplex Ozawa, 1927
Figures 10-1-12

Neoschwagerina simplex Ozawa, 1927, p. 153-154, pI.

996

Katsumi Veno

Figure 10. 1- 12. Neoschwagerina simplex Ozawa, la, 2- 7 : ax ial sections, ASM25143, ASM25144,
ASM25147, ASM25146, ASM25149, ASM25148, ASM25145 , 8a, 9: sagitta l sectio ns, ASM25150, AS M
25 151 , 10- 12: tangential sectio ns, ASM25153 , ASM25154, ASM25152, 1b, 8b : en la rged parts of la and 8a ;la, 2- 7, 8a, 9- 12: x 10, lb, 8b : x 40.

931.

Evolution of Verbeekinidae and Neoschwagerinidae

34, figs. 7-11, 22-23, pI. 37, figs. 3a, 6a; Chen,
1956, p. 11, 55-56, pI. 12, figs. 13-16; Kanmera,
1957, pI. 19, figs. 22-23 (same as pI. 34, figs. 7-8of Ozawa, 1927, p. 153-154); Honjo, 1959, p.
l39-142, pI. 3, figs. 1,4-5, pI. 4; Minato andHonjo, 1959, pI. 1, fig. 2; Suyari, 1962, p. 36, pI.
11, fig. 1; Ishizaki, 1962, p. 175-177, pI. 12, figs.
7-8; Kanmera, 1963, p. 112-113, pI. l3, figs. 1-6,
pI. 14, figs. 1-6, pI. 19, fig. 15; Leven, 1965, p.
143, pI. 4, fig. 4; Sheng, 1966, p. 145, pI. 25, fig. 8(same as pI. 12, fig. 13 of Chen, 1956, p. 11, 5556); Leven, 1967, p. 189-190, pI. 32, figs. 8-10;
Igo, 1967, p. 14-15, pI. 5, figs. 10-11 ; Otuka andKaneda, 1967, p.26, pI. 3, figs. 3-5; Ozawa,
1970, pI. 4, figs. 9-11 (11 : same as pI. 34, fig. 7 of
Ozawa, 1927, p. 153-154); Choi and Fujita,
1970, p. 376-377, pI. 1, figs. 10-12; Ozawa, 1975,
pI. 10, fig. 16 (same as pI. 4, fig.9 of Ozawa,1970); Lin et al., 1977, p. 91, pI. 28, fig. 3; Liuet al., 1978, p. 90, pI. 21, fig. 8; Sash ida, 1980, p.
305-306, pI. 35, figs. 8-10; Leven, 1982, pI. 2, fig.
9 (same as pI. 32, fig. 10 of Leven, 1967, p. 189190); Yang, 1985, pI. 2, fig. 5; Kobayashi,
1988a, p. 11, pI. 6, figs. 1-17; Kobayashi, 1988b,
p.447-449, fig. 6-17.
Cancellina cf. simplex (Ozawa), Dutkevich andKhabakov, 1934, p. 91-94, pI. 2, figs. 4-5.
Cancellina primigena (Hayden), Mik1ukho-Mak1ay,
1957, p. 121-122, pI. 5, figs. 2-3.
Cancellina sphaerica Mik1ukho-Mak1ay, 1957, p.
122-125, pI. 5, fig. 1.
Neoschwagerina cf. simplex Ozawa, Kanmera, 1957,
pI. 20, fig. 1; Kanuma, 1960, p. 67-68, pI. 11, figs.
1, 10-11; Sakaguchi and Yamagiwa, 1973, p.
75-76, pI. 1, figs. 1a-b; Sakagami and Miyama,
1988, fig. 1-14; Tien, 1989, pI. 20, figs. 4-5.
Neoschwagerina sphaerica (Mik1ukho-Mak1ay),
Honjo, 1959, p. 159, pI. 3, fig. 3; Chen andYang, 1978, p. 115, pI. 31, figs. 7-8.
Neoschwagerina simplex simplex Ozawa, Toriyama,
1975, p. 99-101, pI. 19, figs. 25-28, pI. 20, figs. 121.
Neoschwagerina simplex tenuis Toriyama and Kanmera, Toriyama, 1975, p.97-99, pI. 19, figs. 1424; Leven, 1982, pI. 2, fig. 6.
Neoschwagerina cf. simplex tenuis Toriyama andKanmera, Toriyama and Kanmera, 1979, p.74,
pI. 12, fig. 10.
? Maklaya sp., Li, 1986, p. 89, pI. 2, fig. 13.
? Neoschwagerina simplex Ozawa, Sun and Zhang,
1988, pI. 4, fig. 20; Ozawa and Kobayashi, 1990,
pI. 9, fig. 12.
non Neoschwagerina simplex Ozawa, Sheng, 1963, p.
101-102,234-235, pI. 34, figs. 14-15; Han, 1980,
p. 90, pI. 33, figs. 7-8; Chen, 1984, p. 70, pI. 21,
fig. 9.
non Neoschwagerina cf. simplex Ozawa, Koike et al.,997

1968, p.207-208, pI. 32, figs. 6-10.
non Neoschwagerina simplex simplex Ozawa, Han,
1985, p. 685, pI. 2, fig. 11.
non Neoschwagerina simplex tenuis Toriyama andKanmera, Han, 1985, p. 685, pI. 2, figs. 12, 16.

Material. -Axial sections; ASM2S143,
ASM2S146 from Loc. TA204, ASM2S144,
ASM2S14S, ASM2S147, ASM2S148, ASM
2S149 from Loc. NSI73.S. Sagittal sections;
ASM2SISO from Loc. NSI73.5, ASM 2SIS1
from Loc. T A204. Tangential sections;
ASM 2SIS2, ASM2SIS4 from Loc. TA204,
ASM 2SIS3 from Loc. NSI73.S. Additional
specimens from Locs. T A204 and N S 173.S
were also studied.
Description. - Shell small to medi urn for
genus and almost spherical. Mature speci"
mens having 13 to IS volutions, 3.48 to 4.18
mm in length and 3.28 to 4.1S mm in width.
Form ratio ranges from 0.99 to 1.13, averaging LOS for 12 specimens.
Inner 3 or 4 volutions tightly coiled with ashort and slightly rotated axis of coiling.
Outer volutions expanding gradually.
Radi us vectors of the first to fifteenth vol utionof well oriented axial section illustrated by
Figure IO-S; O.OS, 0.09, 0.11, O.IS, 0.20, 0.29,
0.38,0.49,0.64,0.82, LOS, 1.26, I.S0, 1.97 and2.20 mm, and form ratios 0.67, 0.S3, 0.S7, 0.76,
0.90,1.02,1.11,1.18,1.11,1.08,0.98,0.97,0.94,
0.87 and 0.86, respectively.
Proloculus small and spherical. Its outside diameter ranges from O.OSO to 0.080 mm,
averaging 0.OS8 mm for II specimens.
Spirotheca rather thick and consists of atectum and alveolar keriotheca. Thicknessof spirotheca of the first to fifteenth volutionof above-mentioned specimen O.OOS, O.OOS,
O.OOS, 0.010, 0.02S, 0.030, 0.060, 0.030, 0.040,
0.060, 0.120, 0.110, 0.070, 0.100 and 0.04S mm.
Septa composed of downward deflectionsof tectum and anterior and posterior extensions of keriotheca. Septal counts of thesecond to fourteenth volution of typicalsagittal section (Figure 10-8) 6, 8, 9, 10, 13,
IS, 18, 17,21,2S,29,33 and 36, respectively.
Low but broad, fan-shaped primary trans-

Katsumi Ueno

998

verse septula well developed beyond the fifth
volution. They are observed immediately
above each parachomata joining with tops ofparachomata adj acent to septa. Rudimentary axial septula sporadically and inconstantly occur in middle and outer volutions.
Parachomata semicircular or triangular intransverse section and well developed beyond
the fifth vol ution.
Remarks. - This species is considered to be
the most primitive representative of the genus
Neoschwagerina, and the direct descendant of
Maklaya pamirica (Leven). The former
differs from the latter in having a larger shell,
more vol utions and incipient axial septula.
Toriyama (1975) illustrated rich materials
of Neoschwagerina simplex Ozawa (N. simplex simplex Ozawa and N. simplex tenuis
Toriyama and Kanmera) from the Rat Buri
Limestone in the Khao Phlong Phrab area,
central Thailand. The Thailand specimens
have much larger proloculus than the Akiyoshi ones, although the other biocharacters areclosely allied.
Occurrence. - Rare in the uppermost part of
the Misellina (M.) claudiae Zone.
Acknowledgmen ts
I would like to express my sincere appreciation to Prof. Hisayoshi Igo of the Universityof Tsukuba for critical reading of the manuscript. Thanks are also extended to Prof.
Sumio Sakagami of Chiba University, andMessrs. Akihiro Sugimura and TakehikoHaikawa of the Akiyoshi-dai Museum of
Natural History for their continuous advice
and encouragement. Special thanks are due
to Mr. Yasuyuki Miyama of Sumitomo
Cement Co., Ltd. for fruitful discussion on
the fusulinacean taxonomy.
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Buri, central Thailand (Fusuline fossils fromThailand, Part IX). Ibid., vol. 23, no. I, p.l116, pis. 1-21.
- - and Kanmera, K., 1979: Permian fusulines fromthe Ratburi Limestone in the Khao Khao area,
Sara Buri, central Thailand (Fusuline fossils
from Thailand, Part XII). Geol. Palaeont.
Southeast Asia, vol. 20, p. 23-93, pis. 4-14.
Ueno, K., 1991: Pamirina (Permian Fusulinacea)

Katsumi Ueno

1002

from the Akiyoshi Limestone Group, southwest
Japan. Trans. Proc. Palaeont. Soc. Japan, N.
S., no. 161, p. 739-750.
Wang, J.H. and Tang, Y., 1986: Fusulinids from
Chihsia Formation of Lengwu district in Tonglu
county, Zhejiang. Acta Micropalaeont. Sinica,
vol. 3, no. I, p. 3-II, pis. 1-3. (in Chinese withEnglish abstract)
Wang, K.L. and Sun, X.F., 1973: Carboniferous andPermian foraminifera of the Chinling range andits geologic significance. Acta GeoL Sinica,
1973, no. 2, p. 137-178, pis. 1-7. (in Chinese withEnglish abstract and description of new generaand species)
Wang, Y.H., Wang, L.L., Wang, 1.H., Zhu, Z.G., Lin,
G.W., Zhang, L.X. and Quian, Q., 1982:
Fusulinida. In, Nanjing Inst., Geol. Mineral
Resour. chiefly ed., Paleontological A tlas of
East China, no. 2 (Late Paleozoic), p. 5- 108,
pis. 1- 30. Geol. Publ. House, Beijing. (inChinese)Xia, G.Y. and Zhang, Z.e., 1985: Protozoa. In,
Tianjin Inst., Geol. Mineral Resour.
ed.,
Paleontological Atlas of North China, no. I(Paleozoic vol.), p.32-150, pis. 1-35. Geol.
Publ. House, Beijing. (in Chinese)
Xie, S.G., 1982: Protozoa. In, Geol. Bureau, Hunaned., The Palaeontological Atlas of Hunan, Geol.

Mem., ser. 2, no. I, p.2-73, pis. 1-35. Geol.
Publ. House, Beijing. (in Chinese)
Yang, Z.O., 1985: Restudy of fusulinids from the"Maokou Limestone" (Permian) at Oatieguan,
Langdai, Guizhou. Acta Micropalaeont. Sinica,
vol. 2, no. 4, p.307-335, pis. 1-3. (in Chinesewith English abstract and description of new
genus)
Zhang, L.X. and Li, W.Y., 1987: Carboniferous andlower Lower Permian (Chihsia Stage) fusulinidsof Oafeng district, northern liangsu. Acta
Palaeont. Sinica, vol. 26, no. 4, p.392-41O, pis.
1-3. (in Chinese with English abstract anddescription of new species)
Zhou, J.P. and Zhang, L.X., 1984: Fusulinids fromthe Chihsia Formation of Mt. Qixiashan, Nanjing. Ibid., vol. 23, no. 6, p. 716-724, pis. 1-3. (inChinese with English abstract)Zhou, T.M., Sheng, 1.Z. and Wang, Y.l., 1987:
Carboniferous-Permian boundary beds andfusulinid zones at Xiaodushan, Guangnan, eastern Yunnan. Acta Micropalaeont. Sinica, vol.
4, no. 2, p. 123- I 60, pis. 1-6.
Zhou, Z.R., 1984: Middle and late Chihsia (Permian) fusulinids of southeastern Hunan. Acta
Palaeont. Sinica, vol. 23, no. I, p. 107-123, pis.
1-4. (in Chinese with English abstract anddescription of new species)

Akiyoshi fJ\E, Akiyoshi-dai fJ\E-E;-, Chihsia :f2!j~, Chinling .it, Guangxi J1.\il9, Ibukiyama ffTllj;:LlJ, Kaerimizu im IJ 7.1<., Kwanto I*I~, Neimenggu P'I~~, Otaki *i\'1t, Sanmianjing =mj;I:J:, Tamba ftiBl, Yazi :t~T, Yunnan ~i¥i.
tkEEW(!Ei~fft;:,~ i?.tL G Verbeekinidae f4:N J: rJ Neoschwagerinidae f4*Oiill.!RMO):fJJ
Wl~~: fJ\E-E;-~t~1fllO)im IJ 7.l<.itB~t;:,5t;(P..., G Misellina (M.) claudiae 'Iff):.J T O)fJ\EEw(!Ei~ffi»i? 3 ~fi;Misellina (Brevaxina) nipponica, sp. nov., M. (M.) postclaudiae, sp.
nov., M. (M.)? ventricosa, sp. nov. ~~tr 15 fiO) Verbeekinidae f4:N J: rJ Neoschwagerinidaef4Wiiill.!RM~~,8JjL, -to)q:,0) 14fi~~~Lt.:o ~i?t;:" ::"0) 15fic, jilfff4O)m9C~-Ct.;G Pamirina ~O)*oiill.!RMt;:'J: '?-C Pamirina (Levenia) leveni Kobayashi O)l±lmiJ>
i? Parafusulina kaerimizensis (Ozawa) 0) I±lm i -Ct;:, 10 O)lIifra'lID (Biohorizon) ~~;Zn:: L
t.:o ::.. 0) ? '1;" 3 ,,?O)lIifra'lIDft-t.tL.f ic Pamirina (Levenia) leveni 'Iff, Misellina (Brevaxina)
dyhrenfurthi otai'lff, Misellina (M.) claudiae 'lffO)£Jf£~n::~L-C\"Go fK~Ew(!Ei~fft;:,j! i?.tL G ::...tL i? Verbeekinidae f4, Neoschwagerinidae f4, -t L -C Pamirina ~*oiill.!RMO)~~5t;(P, :N J: rJ~n::O) 2 ~ra'O) *~I*If*~IFs~"" G q:,ra'I¥:J~~~~"?fiO).i£1±l ft, Ver~~ L-C
beekinidae f4:N J: rJ Neoschwagerinidae f4*oiill.!RMO)~~ c WWlO) *~~ ? i\" G
l::!flm~<0Trans. Proc. Palaeont. Soc. Japan, N.S., No. 164, p. 1003, December 30, 1991

SHORT NOTES

27.

MELONIS UeHIOI, A NEW NAME FOR A HOMONYM IN THE
FORAMINIFERAL GENUS MELONIS FROM THE PACIFIC*SHIRO HASEGAWA
Institute of Geology and Paleontology, Facultyof Science, Tohoku University, Sendai 980

In the course of a study of benthic foraminifera in the Pacific Ocean, I have recognized a homonym among species of the genus
Melonis. Nonion parkerae was proposed by
Uchio in 1960 for a Recent species inhabitingwaters off the coast of California (Uchio,
1960, p. 60, pI. 4, figs. 9, 10). Matoba (1967)
transferred Uchio's species to the genus
Melonis and this combination was used bymany authors. However, Uchio's speciesname was preoccupied by Nonion parkerae
Le Calvez, established in 1959 for a species
recovered from the Bay of Biscay (Berthois
and Le Calvez, 1959, p. 362, pI. I, figs. 13, 14).
Because Uchio's species name was a juniorprimary homonym of Le Calvez's, it wasinvalid and should be replaced by anothername (International Code of Zoological
Nomenclature, 1985, 3rd ed., Articles 52b, 57b
and 60). I therefore propose Melonis uchioi,
new name, for this species.
References cited
Berthois, L. and Le Calvez, Y., 1959: Deuxieme
contribution a l'etude de la sedimentation dans Ie
Golfe de Gascogne. Rev. Trav. Insl. Peches
Maril., vol. 23, no. 3, p. 323-377, pI. I.
Matoba, Y., 1967: Younger Cenozoic foraminiferal
assemblages from the Choshi district, Chiba
Prefecture. Sci. Rep., Tohoku Univ., 2nd ser.
(Geol.), vol. 38, no. 2, p. 221-263, pis. 25-30.
Uchio, T., 1960: Ecology of living benthonic foraminifera from the San Diego, California, area.
Cushman Found. Foram. Res., Spec. Publ., no.
5, p. 1-72, pis. 1-10.

·Received August 20, 1991; accepted September 5,
1991

1003

Trans. Proc. Palaeont. Soc. Japan, N.S. , No. 164, pp. 1004- 1008, 3 Figs. , December 30, 1991

SHORT NOTES

28.

A PECULIAR FUSULINACEAN FAUNA FROM THE YASUBACONGLOMERATE, KOCHI PREFECTURE, SHIKOKU*
KATSUMI UENO
In stitute of Geoscience, University of Tsukuba, Ib araki 305

A peculiar fusulinacean fauna is found in asmall limestone pebble contained in the
Yasuba Conglomerate, which has long been
regarded as an Upper Permian limestone

conglomerate (Toriyama, 1942 and others).
It crops out typically north of Shingai, Tosayamada Town, Kochi Prefecture, Shikoku(Figure I) . Recently, Isozaki (1985) intensively restudied this conglomerate embedded
in his Shingai Formation and insisted thatthe conglomerate can be subd ivided into
three types, Type I to Type III, which are all
exotic in origin.

Shingai
133° 42 ' E
33°37'N+o

2km

Tosa-yamadaoFigure L Geological map around the fossil
local ity. I : Cretaceous, 2: Kamiyakawa Formation ,3: Shingai Formation, 4: "Shirak idani Group" , 5 :Gonyu Formation , 6 : Agekura Formation , 7: serpentinite, 8: fossil loca lity. (Geological map after
Isozaki , 1985)

Figure 2, Thin sect ion of Waagenophy//um
biolithite conta inin g the fusulinaceans studied herein , x 3.

'Received September 13, 1991 ; accepted September
24, 1991

1004

28.

Fusulinaceans from the Yasuba Conglomerale

I disclosed the present fusulinacean faunafrom a pebble contained in Isozaki's Type II
conglomerate. This fauna is characterized
by the abundant occurrence of Lantschichites
elegans Sosnina, which was reported from theMiddle Permian of southern Primor'e(Sikhote-Alin). Isozaki (op. cit.) consideredthat the Type II conglomerate was deposited
in some area adjacent to the ancient K urosegawa land (arc or microcontinent).
The limestone pebble containing this fusulinacean fauna is a Waagenophyllul11 biolithite with an algal bioclastic micritic matrix(Figure 2). Fusulinaceans and smaller foraminifers abundantly occur in the intercorallite sediments of this fasciculate coral biolithite, such as Rauserella sp., Lanlschichiles
elegans Sosnina, Nankinella sp., Chusenellasp., Tetrataxis sp., Climacaml1lina sp.,
Neoendothyra sp., Pachyphloia sp., Nodosaria sp. and Frondina sp. Among the fusulinaceans, Lantschichites elegans Sosnina ismost characteristic and abundant. This
species was originally described from theMetadoliolina lepida Zone of southern
Sikhote-Alin, which is correlated with thelate Midian of the Middle Permian (Kotlyar
et al., 1989). Lanlschichites was introduced
by Tumanskaya (1953) as a subgenus of thegenus Codon ojusiella , with Codonojusiella(La n tsch ich iles) l11aslenn ikol'i Tumanskayaas the type species. It has been reported
mainly from the latest Middle Permian of
Sikhote-Alin (Tumanskaya, 1953; Sosnina,1968), South China (Chen, 1956; Sheng,
1963; Wang el al., 1981) and Texas (Skinnerand Wilde, 1954). Two species of Lan/schichites have been reported in Japan; one isLanlschichiles inllboensis (Chisaka) from the
Takagami Conglomerate of the Choshi
Peninsula (Chisaka, 1960) and the lwaizaki
Limestone of the southern Kitakami Mountains (Choi, 1970), and the other is Lanlsc/zichites sp. from the Mt. Nabekoshi area in thesouthern Kitakami Mountains (Tazawa,1975). The present fusulinacean fauna
found in a limestone pebble of the Yasuba

1005

Conglomerate is considered to be latest Middle Permian (late Midian) in age and has
close similarity with that of Sikhote-Alin.
Acknowledgments: I would like to express
my appreciation to Prof. Hisayoshi Igo of the
University of Tsukuba for reviewing thismanuscript. Thanks are also extended to
Mr. Yutaka Tanaka for his kind cooperationin the field.

Systematic description
All specimens identified in this paper aredeposited in the collection of Institute of
Geoscience, the University of Tsukuba(IGUT).
Family Boultoniidae Skinnerand Wilde, 1954
Gen us Lanlsc/zichites Tumanskaya, 1953

Lantschichiles elegans Sosnina, 1968
Figures 3-1 ~ 16

Lantschichiles elegans Sosnina, 1968, p. 126-127, pI.
31, figs. 1- 3.

Description. - Shell small, elongate fusiform with bluntly pointed polar ends.
Mature specimens having 4 Yz to 5 Yz volutions, 2.18 to 2.78 mm in length and 1.28 to1.95 mm in width. Shell of regularly coiled
part 0.65 to 0.89 mm in width, giving form
ratios of 3.12 to 3.87.
The first I or I Yz volutions tightly coiled
at large angles to outer ones. Beyond thethird volution, shell becomes elongate rather
rapidly. Last halfvolution abruptly expanded, forming an uncoiled flaring lip about
twice as large as earlier regularly coiled part.
Radi us vectors of the first to fifth, and fifth
and a half volutions of well-oriented axial
section (Figure 3-la) 0.05,0.08,0.12,0.20,
0.35 and 0.52 mm. Form ratios of the second
to fifth, and fifth and a half volution of thesame specimen 1.25,2.67,3.10,3.33 and 2.87.
Proloculus small and spherical. Its out-

1006

K a lsumi Ve na

28.

Fusulinaceans from the Yasuba Conglomerateside diameter ranges from 0.045 to 0.070 mm,
averaging 0.057 mm for 7 specimens.
Spirotheca thin, slightly recrystallized andcomposed of a tectum and lower transparent
layer (diaphanotheca). Thickness of spirotheca of the first to fifth vol utions of abovementioned axial section 0.005, 0.005, 0.015,
0.020 and 0.030 mm, respectively.
Septa intensely and regularly fluted
throughout length of shell, forming high andnarrow, omega-shaped septal loops.
Cuniculi well developed in outer volutions.
Chomata not observed. Weak axial fillings
developed in inner 2 or 3 volutions.
Remarks. - This species can be distinguished from Lantschichites maslennikol'i(Tumanskaya) in having a larger shell andform ratio, and from L. inuboensis (Chisaka)in having a larger shell.
The inner regularly coiled part of Lantschichites elegans is somewhat similar to that ofL. splendens (Skinner and Wilde) from theuppermost Guadalupian of Texas. However, the former has a larger and more developed uncoiled flaring part than the latter.
Lantschichites minima (Chen) descri bed
and ill ustrated by Sheng (1963) from theupper part of the Maokou Limestone ofGuangxi (Kwangsi) also resembles thisspecies. However, the former differs fromthe latter in having a more slenderly cylindrical shell and larger form ratio.
Material. - Axial sections; IG UT - K U
0001, IGUT-KU0002, IGUT-KU0003,
IGUT -KU0004. Axial section of immature
specimen; IGUT -KU0005. Sagittal sections; IGUT-KU0006, IGUT-KU0007.
Parallel sections; IGUT -KU0008, IGUTKU0009, IGUT -KUOOIO, IGUT -KUOOII,
IGUT -KUOOI2, IGUT -KU0013, IGUT+----

KU0014, IGUT-KUOOI5.
tion; IGUT -KUOOI6.

1007

Tangential

sec-

References
Chen, S., 1956: Fusulinidae of South China, Part 2.
Palaeont. Sinica. N.S. B, no. 6, p. 1-71, pIs. 1-14.
Chisaka, T., 1960: On some Permian fusulinids fromthe Takagami Conglomerate, Choshi Peninsula,
Chiba Prefecture, Japan. Jour. ColI. Arts andSci., Chiba Univ., vol. 3, no. 2, p. 235-254, pIs. 19.
Choi, D.R., 1970: On somc Permian fusulinids from
Iwaizaki, N.E. Japan. Jour. Fac. Sci., Hokkaido Univ., Ser. 4, vol. 14, no. 3, p. 313-325, pIs.
5-8.
Isozaki, Y., 1985: Yasuba conglomerate and its modeof occurrence. Jour. Geol. Soc. Japan, vol. 91,
no. 8, p. 535-551, pI. 1. (in Japanese Wilh English
abstract)
Kotlyar, G.V., Zakharov, Yu. D., Kropatcheva, G.S.,
Pronina, G.P., Chedija, 1.0. and Burago, V.I.,
1989: Pozdnepermskiy etap el'olyutsii organicheskogo mira, Midiyskiy yarus SSSR (ProjectNos. 203 and 272 of thc International Geological
Correlation Programmc). Akad. Nauk SSSR,
Dal'nevostochnyy Gcol. Inst., 182 p., 28 pIs.
"Nauka", Leningradskoe otdel., Leningrad. (inRussian)Sheng, J.Z. (Sheng, J.c.), 1963: Permian fusulinids ofKwangsi, Kueichow and Szechuan. Palaeont.
Sinica. N.S. B, no. 10, p. 1-247, pIs. 1-36.
Skinner, J.W. and Wilde, G.L., 1954: The fusulinid
subfamily Boultoniidae. Jour. Paleont., vol. 28,
no. 4, p. 434-444, pIs. 42-45.
Sosnina, M.I., 1968: Novyc pozdnepermskie fuzulinidy Sikhote-Alinya. In, Novye vid)' drevnikh
rasteniy i bespozvonochnykh SSSR, Vyp.2,
Chast' Pervaya, p.99-l28, pIs. 25-31, Vscs.
Nauchno-Issled. Geol. Inst. (VSEGEI), Moscow.
(in Russian)
Tazawa, J., 1975: Uppermost Permian fossils fromthe southern Kitakami Mountains, northeast
Japan. Jour. Geol. Soc. Japan, vol. 81, no. 10,
p. 629-640, pIs. 1-3.
Toriyama, R., 1942: The fusulinids of the Yasuba
conglomeratc in the Province of Tosa. Japan.

Figure 3. 1-16. Lanlschichiles elegans Sosnina, la, 2-4: axial scctions, IGUT-KUOOOl, IGUTKU0002, IGUT-KU0003, GUT-KU0004, 5: axial section of immature specimen, IGUT-KU0005, 6-7:
sagittal sections, IGUT -KU0006, IGUT -KU0007, 8-15: parallel sections, IGUT -KU0008, IGUT -KU0009,
IGUT-KUOOlO, IGUT-KUOOll, IGUT-KU0012, IGUT-KU0013, IGUT-KU0014, IGUT-KU0015, 16:
tangential section, IGUT -KU0016, X 30, Ib: enlarged part of la, X 75. 17: Rallserella sp., axial section,
IGUT-KU0017, x30. 18: Nankinella sp., tangential section, IGUT-KU0018, x30. 19: Chusenella sp.,tangential section, IGUT-KU0019, X 10.

1008

Katsumi Ueno

Jour. Ceol. Ceogr., vol. 18, no. 4, p. 237-247, pis.
24-25.
Tumanskaya, O.G., 1953: 0 verkhnepermskikh fuzulinidakh Yuzhno-Ussuriyskogo kraya. Trudy
Vses. Nauchno-Issled. Ceol. Inst. (VSECEl), p.
3-25, pls.I-15. (in Russian)Wang, YJ., Sheng, J.Z. and Zhang, L.X., 1981:
Fusulinids from Xizang of China. In, NanjingInst. Geol. Palaeont. ed., Palaeontology of
Xizang, Book III, p. 1-80, pis. 1-21. Sci. Publ.
House, Beijing. (in Chinese with English abstractand description of new subgenus)

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

New Series No. 164

December 30, 1991

CONTENTS-TRANSACTIONS
926. Kuniteru Matsumaru: On the evolutionary classification of the family
Lepidocyclinidae (Foraminiferida) .................................... 883
927. Tatsuro Matsumoto: On some Acanthoceratid ammonites' from the Turonianof Hokkaido (Studies of the Cretaceous ammonites from Hokkaido-LXIX) 910
928. Katsumi Veno and Sumio Sakagami: Late Permian Fusulinacean fauna of
Doi Pha Phlung, North Thailand .................................... 928
929. Tamiko Ohana and Tatsuaki Kimura: Permineralized Otozamites leaves(Bennettitales) from the upper Cretaceous of Hokkaido, Japan .. . . . . . . .. 944
930. Ken'ichi Saiki, Tatsuaki Kimura and Junji Horiuchi: Stenopteris cyclostoma
Saiki, Kimura et Horiuchi, sp. nov. (possible Pteridosperm), from the lower
Cretaceous Choshi Group, in the outer zone of Japan .................. 964
931. Katsumi Veno: Early evolution of the families Verbeekinidae and Neoschwagerinidae (Permian Fusulinacea) in the Akiyoshi Limestone Group,
Southwest Japan
973
SHORT NOTES
27.

Shiro Hasegawa: Melonis uchioi, a new name for a homonym in the for-28.

aminiferal genus Melonis from the Pacific .............................. 1003
Katsumi Veno: A peculiar fusulinacean fauna from the Yasuba Conglomerate, Kochi Prefecture, Shikoku ........................................ 1004