ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No.

169

Palaeontological Society of Japan
April 30, 1993

Co-Editors

Tsunemasa Saito and Kei Mori

Language Editor

Martin Janal (New York)

Editorial Board
Shiro Hasegawa (Hokkaido University), Kunihiro Ishizaki (Tohoku University), Tomoki Kase
(National Science Museum), Toshiaki Maruyama (Yamagata Univer,ity), Kei Mori (Tohoku University), Kenshiro Ogasawara (University of Tsukuba), Tsunemasa Saito (Tohoku University), Y okichi Takayanagi (Ishinomaki Senshu University), Kazushige Tanabe (University of Tokyo), Yukimitsu Tomida (National Science Museum), Kazuhiko Uemura (National Science Museum), Akira Yao(Osaka City University)

Officers for 1993-1994
President: Hisayoshi Igo
Honorary President: Teiichi Kobayashi
Councillors: Kiyotaka Chinzei, Takashi Hamada, Y oshikazu Hasegawa, Itaru Hayami,
Hiromichi Hirano, Hisayoshi Igo, Noriyuki Ikeya, Junji Itoigawa, Tomoki
Kase, Makoto Kato, Tatsuaki Kimura, Itaru Koizumi, Kei Mori, Hiroshi Noda,
Ikuwo Obata, Kenshiro Ogasawara, Tomowo Ozawa, Tsunemasa Saito, Yokichi
Takayanagi, Kazushige Tanabe, Akira Yao
Members of Standing Committee: Kenshiro Ogasawara (General Affairs), Tomoki Kase
(Finance), Tsunemasa Saito (Editor in Chief, TPPSJ), Kei Mori (Co-Editor, TPPSJ),
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Auditor: Hisaharu Igo
The fossil on the cover is Isocrinus (Chladocrinus) hanaii Oji, an Early Cretaceous(Aptian) crinoid, which was described from the Hiraiga Formation exposed at Haipe,
Tanohata-mura, Shimo-Hei County, Iwate Prefecture, Northeast Japan. (University
Museum of the University of Tokyo coil. cat. no. ME6950, paratype specimen, length
about 11 cm)

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 169, pp. 1-14, 4 Figs., April 30, 1993

948.

EARLY MIOCENE (AQUITANIAN) LARGER FORAMINIFERAFROM THE SHIMIZU FORMATION, ASHIZURI CAPE,
KOCHIPREFECTURE,SHIKOKU,JAPAN*
KUNITERU MATSUMARU
Department of Geology, Faculty of Education, Saitama
University, Urawa, 338

MYINT THEIN
Department of Geology, University of Mandalay,
Mandalay, Myanmar
and

YUJIRO OGAWA
Institute of Geoscience, University of Tsukuba,
Tsukuba, 305

Abstract. Early Miocene (Aquitanian) larger Foraminifera are first described from thecalcareous matrix of volcanic conglomerate and sandstone of the Shimizu Formation in the
Shimanto Belt, Tosa Shimizu City, Ashizuri Cape, Kochi Prefecture, Shikoku. The joint
occurrence of Miogypsinoides dehaartii (van der Vlerk), Spiroclypeus margaritata (Schlumberger) and Victoriella conoidea (Rutten) enables the age of the calcareous matrix of volcanic
conglomerate and sandstone of the Shimizu Formation to be assigned to the Early Miocene(Aquitanian). Detailed stratigraphic studies, which were carried out in Ashizuri Cape,
Tosa Shimizu City, Shikoku, show that the mudstone yielding the Late Eocene (Priabonian)larger Foraminifera described by Matsumaru and Kimura (1989) is not in the Shimizu
Formation, but in the Kurusuno Formation (Myint Thein et al., 1991; Myint Thein, 1992).
As such, the newly described larger Foraminifera from the Shimizu Formation indicate an
age for the formation of Early Miocene (Aquitanian).
Key words. Miogypsinoides, Miogypsina, Nephrolepidina, Spiroclypeus, Aquitanian,
Shimizu Formation.

Introduction
The Shimanto Belt in Kochi Prefecture,
Shikoku, which is known as a tectonic belt
along the outermost zone of Southwest Japan,
consists of highly deformed flysch deposits
and melange complex with some exotic
blocks (e.g. basalt, red pelagic shale, etc.).
'Received April 24, 1992; revised
accepted January 25, 1993

manuscript

In Ashizuri Cape, Kochi Prefecture, the Shimizu Formation, which occupies the southernmost part of the Shimanto Belt, exhibits
syndepositionally deformed flysch and debris
flow deposits, and rarely yields fossils.
Recently, Matsumaru and Kimura (1989)
found Late Eocene (Priabonian) larger Foraminifera from a gray tuffaceous mudstone of
the Shimizu Formation at Takahata Village,
Tosa Shimizu City, Kochi Prefecture, but

Kuniteru Matsumaru et al.

2

Myint Thein et al. (1991) and Myint Thein(1992) consider it as belonging to the Kurusuno Formation from the viewpoint of lithologic characteristics. Therefore, the Kurusuno Formation contains Asterocyclina stella(Gumbel), Orbitoclypeus kimurai Matsumaru, O. sp., Discocyclina sella (d'Archiac),
and the age of the formation is Late Eocene(Priabonian) (Matsumaru and Kimura, 1989).
On the other hand, the age of the Shimizu
Formation had to be reexamined. Myint
Thein et al. (1991) fortunately found Upper
Cretaceous radiolarians such as Protoxiphotractus perplexus Pessagno, etc. fromdeformed mudstone of the Shimizu Formation exposed along the coastal bench at Tsuro
Village, Tosa Shimizu City in Ashizuri Cape,
Kochi Prefecture. However, these fossils are
considered to be allochthonous. Yujiro
Ogawa, one of the Myint Thein et al.
research group, sent nine thin sections fromthe calcareous matrix of volcanic conglomerate and sandstone of the Shimizu Formation,
Tosa Shimizu City, Ashizuri Cape, toKuniteru Matsumaru for study of the larger
Foraminifera. The fossil locality for thepresent study is along the sea coast on thesouthern part of Ohki Beach, Tosa Shimizu
City in Ashizuri Cape, Kochi Prefecture (Lat.
37'47' 15.4"N., Long. 132°57'34.6"E.) (Figure
1).

The purpose of this paper is primarily tomake the first description and discussion of 5
genera and 5 species of larger Foraminifera,
and 1 genus and 1 species of smaller Foraminifera from the Shimizu Formation, andsecondarily to propose a geological age basedon the presence of age-diagnostic species.
All specimens described herein have been
deposited in the collections of the Department of Geology, Faculty of Education,
Saitama University.

Geological age
The larger and smaller Foraminifera foundfrom the calcareous matrix of volcanic con-glomerate and sandstone of the Shimizu
Formation, Tosa Shimizu City, Ashizuri
Cape, Kochi Prefecture were determined asfollows: Miogypsinoides dehaartii (van der
Vlerk), Miogypsina sp., Nephrolepidina praejaponica Matsumaru, Spiroclypeus margaritata (Schlumberger) and Victoriella conoidea(Rutten), which belong to the larger Foraminifera, and Pararotalia sp. belonging to
the smaller Foraminifera. Miogypsinoides
dehaartii and Spiroclypeus margaritata are
diagnostic larger Foraminifera indicative of
the Tertiary e5 of the East Indian Letter
Classification (Matsumaru, 1974). The present fauna in the calcareous matrix of the
Shimizu Formation at this location is especially characterized by the joint occurrence ofMiogypsinoides dehaartii (van der Vlerk),
Spiroclypeus margaritata (Schlumberger) andVictoriella conoidea (Rutten).
Miogypsinoides dehaartii was first described by van der Vlerk (1924) from Dr. F.
Weber's sample from the OrbitoidenMergelkalk and Foraminiferenkalk, Larat
Island, Tanimbar Islands, Indonesia. In 42
years later, van der Vlerk was led to theconclusion that the stratum of type locality of
Miogypsinoides dehaartii is of Aquitanianage, based on biometrical investigations on
Miogypsinoides dehaartii, Lepidocyclina sp.
and Cycloclypeus sp. This judgement is
especially based on statistical comparison of
the value Factor A showing the degree ofdevelopment of embryonic chambers between
Larat Lepidocyclina from Indonesia andLepidocyclina of Aquitanian age in sample
FR475 from Grand Nid-Sausset, Marseille,
France (van der Vlerk, 1964).
Spiroclypeus margaritata (Schlumberger,
1902) was described from the "Oligocene"
limestone of Teweh, Barito Basin, Borneo,
Indonesia. Schlumberger stated that thisspecies resembles Heterostegina sp. of Prof. K.
Martin, University of Leiden, from the"Oligocene" beds near Dax, Aquitaine Basin,
France. Recently, Butt (1966) described
Spiroclypeus blankenhorni Henson var. or-

Early Miocene larger Foraminifera from Shikoku

948.

Pacific Ocean

..
iii

~
••• Alluvium and terrace deposits
•

,0

o

Misaki Group
Coherent Facies
Shimizu Formation

F1
L:=:J Chaotic Facies--

Coherent Facies
Kurusuno Formation

Chaotic Facies

r+.:+l Ashizuri Granite

lLJ_Fault_. _.

Uncon formity---+- SynclineoI

Figure 1. Map showing the fossil locality (arrow) at the southern part of Ohki Beach, Tosa Shimizu
City, Ashizuri Cape, Kochi Prefecture, and geological map of the study area.

3

4

Kuniteru Matsumaru et al.

948.

Early Miocene larger Foraminifera from Shikoku

nata Henson from Escornebeou near Dax,
where the Aquitanian stratotype is located,and Martin's Heterostegina sp. from near Dax
can be assigned to this species.
Spirocypeus blankenhorni and S. blankenhorni var. ornata were originally describedfrom the Lower Miocene of Aintab, Turkish
Syria (Henson, 1937). Adams et al. (1983)
state that Spiroclypeus blankenhorni var.
ornata is known from the Chattian Stage in
the Mediterranean region, while Spiroclypeus
blankenhorni and S. margaritata are common in the Aquitanian Stage in both theMediterranean and Indo-Pacific regions.
They do not, however, give any evidence or
criticism of the species concepts among Spiroclypeus blankenhorni, S. blankenhorni var.
ornata and S. margaritata. The present
authors consider that the former two may be
a junior synonym of the latter, judging from
Henson's description and illustration. In
Kita-Daito-Zima (North Borodino Island),Japan, Hanzawa (1940, p.764, p.790) once
noted that Spiroclypeus margaritata occurs in
Zone 4 (302.31 m-394.98 m) of drill cores of
the Daito Limestone as an Aquitanian form,
its geological horizon being assigned to thatstage based on Spiroclypeus attaining its climax of development in the Aquitanian.
Debourle and Delmas (1955) described
Victoriella aquitanica from the Oligocene(probably Aquitanian) of Mimizan, Landes,
France. They differentiated Victoriellaaquitanica from Victoriella "plecte" (Chapman), which is a junior synonym of Victorielfa conoidea (Rutten), by its regularly trochospiral form through ontogeny. Glaessnerand Wada (1959, p. 199) were of the opinion
that Victoriella aquitanica would prove to bea synonym of Victoriella conoidea, and also
state that Victoriella conoidea is rare in thelate Eocene and widespread in the Oligocene
to Aquitanian. Recently, Chaproniere

5(1984) described Victoriella conoidea fromsample 70614 of the Puketi Formation,
Waitakere Group from Kaipara Harbour of
Huketere Peninsula, New Zealand. He expressed the age of the Puketi Formation as
Zone N. 5 in terms of the planktonic foraminiferal zonal scheme of Blow (1969).
The Puketi Formation is then correlated with
the Aquitanian based on the same zonal
scheme (Blow, op. cit.).
Therefere, the conspicuous presence of suchage-diagnostic species as Miogypsinoides dehaartii, Spiroclypeus margaritata and Vic toriella conoidea enables the occurrence of thepresent larger foraminiferal assemblage in the
Shimizu Formation to be set as Aquitanian.
As such, an Early Miocene (Aquitanian) age,
Tertiary e5, is assigned to the calcareous
matrix of volcanic conglomerate and sandstone of the Shimizu Formation forming the
Shimanto Belt, in the southern part of Ohki
Beach, Tosa Shimizu City, Ashizuri Cape,
Kochi Prefecture.
Description of species
Family Miogypsinidae Vaughan, 1928
Genus Miogypsinoides Yabe andHanzawa, 1928
Miogypsinoides dehaartii(van der Vlerk, 1924)
Figures 2-1,3-2 left, 3-3-5

Miogypsina dehaartii van der V1erk, 1924, p. 429-431,
text-figs. 1-3.
Miogypsina abunensis Tobler, 1927, p. 323-330, textfigs, 3, 5.
Miogypsinoides dehaartii (van der Vlerk) var. pustulosa Hanzawa, 1940, p. 780-782, pI. 39, figs. 2021, pI. 40, figs. 2-29, pI. 42, fig. 13, text-fig. 5.
Miogypsinoides lateralis Hanzawa, 1940, p. 783, pI.
39, figs. 10-14.
Miogypsinoides sp. aff. dehaartii (van der Vlerk).
Bronnimann, 1940, p. 80-81. pI. 8, fig. 19.
Miogypsina (Miogypsinoides) dehaartii van der Vlerk.

+- Figure 2. 1. Miogypsinoides dehaartii (van der Vlerk). Oblique section.

2. .Spiroc!rpeus marg~r­
itata (Schlumberger). Vertical section. 3. Victoriella conoidea (Rutten). Tangenllal sectIOn near ~xlal
plane. 4. Nephrolepidina praejaponica Matsumaru. Vertical section. 5,6. Pararotalia sp; 5, Obliquesection, 6. Vertical section. 1, 5, 6, X 100; 2, 3, 4, X 40.

Kuniteru Matsumaru et al.

6

Drooger, 1953, p.llO-114, pI. I, figs. 15-26;
Raju, 1974, p. 80-81, pI. I, figs. 19-25, pI. 3, fig. 8,
pI. 4, figs. 2-4; Adams and Belford, 1974, p. 497,
pI. 73, figs. 12-14; Matsumaru and Barcelona,
1982, p.84-89, pI. 14, fig. 6 (left); Hashimoto,Matsumaru and Alcantara, 1982, p. 33-37, pI. 10,
fig. 8, pI. II, fig. 22; Chaproniere, 1984, p. 46-47,
pI. 7, figs. 7a-b, pI. 8, figs. 1-3, pI. 17, figs. 15-17,
text-fig. 17.
Miogypsinoides dehaartii (van der Vlerk). Cole,
1954, p. 602, pI. 220, figs. 1-8; Cole, 1957a, p.
339-340, pI. 111, figs. 5-7, 9-16 (non fig. 8);
Cole, 1957b, p. 769, pI. 243, figs. 1-2 (non fig. 3),
pI. 244, fig. 3; Hanzawa, 1957, p.92, pI. 15, figs.
3,7,9,12, 19,22, pI. 19, fig. 3, pI. 21, fig. 4;
Coleman, 1963, p. 13, pI. 2, figs. 13-20; van der
Vlerk, 1966, p. 422-423, pI. I, figs. 1-4 (non fig.
5), 7-10, pI. 2, fig. I (non fig. 3); Bielle andCaralp, 1967, p.36-42, pI. 4, figs. 1-2; Cole,
1969, p. CIO-\2, pI. I, figs. 2-12,15,20 (non figs.
1,13-14,16-19); Matsumaru, 1974, p.107-109,
pI. 17, figs. 2-5, 10-\2,14-15,18-19,21,24-25;
Matsumaru, 1976, p. 196-198, 204, pI. 4, figs. 7,
14, 17 (non figs. 3, 23).
Miogypsinoides sp., Coleman, 1963, p. 13-14, pI. 3,
figs. 1-6.
Miogypsinoides sp. cf. dehaartii (van der Vlerk).
Bielle and Caralp, 1967, p. 36-42, pI. 4, figs. 3-6.
Miogypsinoides abunensis (Tobler). Matsumaru,
1974, p.107-109, pI. 17, fig. 22; Matsumaru,
1976, p. 196-198, pI. 4, figs. 8, II.
Miogypsinoides formosensis Yabe and Hanzawa.
Matsumaru, 1976, p. 196-198, pI. 4, figs. 6, 12, 19.
Miogypsinoides bantamensis Tan. Matsumaru, 1976,
p. 196-198, pI. 4, figs. 15, 18.

Description.- Test is of moderate size, 1.0
to 1.4 mm long, more than 0.6 mm wide and
0.4 to 0.5 mm thick, fan-shaped in outline andbiconvex. The surface of the test is smoothor covered with pustules of 40 to 80,um indiameter.
The embryonic chambers consist of spherical protoconch, 117,um in diameter, andkidney-shaped deuteroconch, more than 88,urn in diameter, which are at the begining ofa single whorl of subquadrate-shaped nepionic chambers. The protoconch is separated from the apical border of test by interven~tion of the deuteroconch, and y parameter(Drooger, 1952) shows a negative value ofabout 30 degrees.
The number of nepionic chambers is morethan 8. Eight rows of ogival to rhombic
equatorial chambers are developed, 94 to 104,urn in radial length and 80 to 88,um in tangential length on the frontal side of nepionicstage, gradually enlarging toward frontal
margin of the test. Internal height of theequatorial chambers is 120 to 208,um invertical sections.
The lateral layers are compact and laminated, with vertical canals of 6 to 8 ,urn in diameter penetrating lateral layers from septa ofequatorial chambers. The rudimentary lateral chambers of 80 to 112,um width and 24
to 32,um height occur sometimes over the
nucleoconch near the surface of the test.
Remarks.-Although there are no equatorial sections of specimens for study, the present form is easily identified as Miogypsinoides
dehaartii (van der Vlerk) in every respect:apical position of embryonic chambers, negative y parameter, single whorl of nepionicchambers, equatorial chamber rows and compact laminated structure of lateral layers. vander Vlerk (1924, 1966) confirmed the presenceof rudimentary lateral chambers in his
Miogypsina dehaartii from the type locality.
This species is, however, classified as a
Miogypsinoides by many authors and also wasplaced in the subgenus Miogypsinoides of
Miogypsina by van der Vlerk (1966).
Tan Sin Hok (1936, p. 51-52) established a
new subgenus Conomiogypsinoides based on
Miogypsina abunensis Tobler, but Hanzawa(1940, p. 768-769) regarded Conomiogypsinoides to be nothing but a synonym of
Miogypsinoides, whose type species is Miogypsina dehaartii van der Vlerk. van der Vlerk(1966) investigated again an assemblage of
Miogypsinoides dehaartii from the type local-

Figure 3. 1. Spiroclypeus margaritata (Schlumberger). Vertical section. 2 (left), 3-5. Miogypsinoides dehaartii (van der Vlerk). All figures of vertical sections, except for oblique section of Figure 3-4.
2 (right). Miogypsina sp. Oblique section. 6,7,8. Nephrolepidina praejaponica Matsumaru; 6, 7. Obliquesections, 8, Vertical section. All figures x40.

948.

Early Miocene larger Foraminifera from Shikoku

7

Kuniteru Matsumaru et al.

8

ity and regarded Conomiogypsinoides to be asynonym of Miogypsinoides. The present
authors synonymize Miogypsinoides abunensis (Tobler) with Miogypsinoides deha artii,
because both species have in common thedevelopment of compact laminated structureof lateral layers.
Genus Miogypsina Sacco, 1893
Miogypsina sp.
Figure 3-2 right

Description.-Several random thin sections
assigned to the genus Miogypsina are in thepresent collection, but none of them adequately exhibits the equatorial and verticalplanes. The oblique sections indicate asmall test with a diameter of 1.0 to 1.5 mm
and there are 4 to 5 regular layers of lateralchambers on each side of the equatorial layer.
Some pillars with diameters of as much as 40
pm are scattered throughout the inflated
portion of the test.
Remarks.-Due to the lack of equatorial
sections the present form is not identified, but
may be used as Miogypsina borneensis Tan,because this species is in association with
Miogypsinoides dehaartii in the type locality.
Also, in the fauna of Zones 3 and 4 ofKita-Daito-Zima, Miogypsina borneensis isassociated with Miogypsinoides dehaartii var.
pustulosa Hanzawa, M. bantamensis Tan, M.
lateralis Hanzawa, Miogypsina polymorpha(Rutten), LepidocyC/ina (Nephrolepidina)
tournoueri Lemoine and R. DouvilltS, L. (N.)
plicomargo Hanzawa, Spiroclypeus margaritata (Schlumberger), Cycloclypeus communis Martin, Operculina gaimardi d'Orbigny, Heterostegina depressa d'Orbigny,
Rotalia gaimardi d'Orbigny, Planorbulinella
larvata (Parker and Jones), Austrotrillina
howchini (Schlumberger), Borodinia septentrionalis Hanzawa, and others (Hanzawa,1940). The present larger foraminiferal
fauna with Miogypsina sp. from the Shimizu
Formation in Ashizuri Cape can therefore becorrelated in this manner with a partial faunafrom Zones 3 and 4 of Kita-Daito-Zima.
Family Lepidocyclinidae Scheffem, 1932
Genus Nephrolepidina H. DouvilltS, 1911
Nephrolepidina praejaponica Matsumaru,
1989
Figures 2-4; 3-6-8.

Nephrolepidina praejaponica Matsumaru. Matsumaru and Kimura, 1989, p. 265, 267, figs. 6-113.

Description.- Test small, lenticular, withdiameter of 1.6 to 2.3 mm and thickness of 0.7
mm. Conical pillars are from 42 to 83 pm indiameter, and distributed on the central partof the test surface.
The embryonic chambers exposed in
oblique sections are nephrolepidine, and theprotoconch is subcircular with a diameter ofmore than 136 pm. The second chamber or
deuteroconch, which embraces the protoconch, has an internal diameters of more than 176
pm. The ratio (= DII/DI) of the innerdiameter of deuteroconch (II) to that ofprotoconch (1) is 1.3. The distance acrossboth the protoconch and deuteroconch is 232
pm. The outer wall of the embryonic chambers is about 20 pm thick.
The equatorial chambers of arcuate form
near the periembryonic chambers change
from ogival to short hexagonal near the
periphery. Those near the periphery have aradial diameter from 56 to 70 pm and a transverse diameter from 52 to 80 pm. Heightincreases gradually towards periphery, about
40 pm near the center and about 65 pm near
the periphery.
The lateral chambers are arranged in a tierof 6 to 10 layers over the center. Chambers
over the center and at the periphery have alength of 77 to 120 pm, a height of 20 to 30
pm, and floors and roofs 15 to 20 pm thick.
Remarks.- Though a sufficient number ofthin sections was not available for study, thepresent specimen has the same features ofsmall embryonic chambers and short hexagonal equatorial chambers as Nephrolepidina

948.

Early Miocene larger Foraminifera from Shikoku

Figure 4. Spiroclypeus margaritata
oblique sections. All figures X 40.

(Schlumberger).

1, 4, 5, 7, 8, 9, verticalsections.

9

2, 3, 6,

Kuniteru Matsumaru et al.

10

praejaponica Matsumaru from the Lower
Member of· the Misaki Formation, Tosa
Shimizu City, Kochi Prefecture, Shikoku(Matsumaru and Kimura, 1989). The present form is thus assigned to Nephrolepidina
praejaponica.
Nephrolepidina praejaponica is considered
to exhibit more primitive features than Nephrolepidina japonica (Yabe) as already indicated by Matsumaru and Kimura (op. cit.).
According to Matsumaru (1971, p. 164),
Nephrolepidina japonica is considered to havederived from Nephrolepidina tournoueri, but
the former may be a junior synonym of thelatter. In Europe, Early Miocene (Burdigalian) Nephrolepidina tournoueri evolved
from Late Oligocene (Chattian) to Early
Miocene (Aquitanian) Nephrolepidina morgani (Lemoine and R. Douvilll!) (de Mulder,1975). Nephrolepidina praejaponica may
then be assigned to Nephrolepidina morgani.
As the nepionic spiral of Nephrolepidina
praejaponica is obscure, the name praejaponica is used in the present instance.
Nephrolepidina praejaponica was found in
the Late Early Miocene (Burdigalian) Misaki
Formation (Matsumaru and Kimura, 1989)
and the Early Miocene (Aquitanian) Shimizu
Formation in this paper, both in Tosa Shimizu City. As both formations are boundedby faults, the stratigraphical relationshipbetween them cannot be demonstrated (Figure 1). It is, however, inferred that both the
Misaki and Shimizu Formations, which arecharacterized by Nephrolepidina praejaponica, are in close stratigraphical relationship.
Family Nummulitidae de Blainville, 1827
Genus Spiroc/ypeus H. Douvilll!, 1905
Spiroc/ypeus margaritata(Schlumberger, 1902)
Figures 2-1-9; 3-1.

Heterostegina margaritata Sch 1umberger, 1902, p.
252-253. pI. 7, fig. 4.
Spiroclypeus orbitoideus H. Douvill15, 1905, p.460-

462, pI. 14, figs. 1-6; Tan, 1937, p. 183-184, pI. 1,
figs. 2-4, pI. 2, figs. 1-13, pI. 3, figs. 1-7 (non figs.
8,24), pI. 4, fig. 1 ; Cole, 1957a, p. 332-333, pI.
95, figs. 6-12; Matsumaru, 1976, p. 200, pI. 1,
figs. 1,8, 10; Hashimoto, Matsumaru andSugaya, 1981, p. 59, pI. 13, fig. 8.
Spiroclypeus leupoldi van der Vlerk, 1925, p. 14-15,
pI. 2, fig. 16; pI. 5, figs. 41, 48; Yabe and Hanzawa, 1929, p. 188, pI. 24, fig. 9; Cole, 1954, p.
577-578, pI. 208, figs. 1-19; Hanzawa, 1957, p.
45-46, pI. 5, figs. 7-13; Matsumaru, 1974, p. 108,
pI. 15, figs. 2-4,10, 13-15,21-23,28; Matsumaru, 1976, p. 199-200, pI. 1, figs. 4-7,14-15,21,2324; Hashimoto, Matsumaru and Alcantara,
1982, p. 34-36, pI. 10, figs. 18-20, pI. II, figs. 1-7,
9.
Spiroclypeus wolfgangi van der Vlerk, 1925, p. 15-16,
pI. 2, fig. 15, pI. 5, figs. 39,49; Tan, 1937, p. 183,
pI. 1, fig. I.
Spiroclypeus yabei van der Vlerk, 1925, p. 16, pI. 2,
fig. 19, pI. 5, figs. 40, 50; Tan, 1937, p. 183, pI. 1,
figs. 5-6, pI. 3, figs. 10-11, pI. 4, figs. 8-10, text-fig.
1; Cole, 1954, p. 580-581, pI. 207, figs. 1-14, pI.
208, figs. 20-26; Cole, 1957b, p. 764, pI. 239, figs.
9-10.
Spiroclypeus tidoenganensis van der Vlerk, 1925, p.
16-17, pI. 1, fig. 12, pI. 5, figs. 42,47; Tan, 1937,
p. 183, pI. 1, fig. 10, pI. 2, figs. 4-5, pI. 3, fig. 12,
pI. 4, figs. 2-5, 19-21 ; Hanzawa, 1957, p.46-47,
pI. 3, figs. 1-6, pI. 4, figs. 1,8-10; Cole, 1957a, p.
332, pI. 95, figs. 13-15; Matsumaru, 1976, p. 200,
pI. 1, figs. 3, 9, 12, 18-20,22, pI. 6, fig. 15; Hashimoto and Matsumaru, 1978, p.85-86, pI. 11,
fig. 2; Hashimoto, Matsumaru and Sugaya, 1981,
p. 60-61, pI. 13, figs. 9, 12.
Spiroclypeus margaritata (Schlumberger). Yabe andHanzawa, 1925, p.627-630, pI. 2, figs. 10, pI. 3,
figs. 8-9, pI. 4, figs. 3-8, text-figs. 1-4; Yabe andHanzawa, 1929, p. 187, pI. 23, figs. 1,3-4, pI. 24,
figs. 1-5; Krijnen, 1931, p.89, pI. I, figs. 1-3;
Tan, 1937, p. 182-183, pI. 2, fig. 12, pI. 3, fig. 9, pI.
4, figs. 6-7; Hanzawa, 1940, p. 789-790, pI. 42,
figs. 3-9; Cole, 1954, p.578-580, pI. 206, figs.
10-25, pI. 207, figs. 15-16; Matsumaru, 1974, p.
108, pI. 15, figs. 16,24,26; Hashimoto and Matsumaru, 1975, p.122, pI. 13, figs. 11-12; Hashimoto, Matsumaru and Sugaya, 1981, p. 59-60,
pI. 13, fig. 3; Hashimoto, Matsumaru and Alcantara, 1982, p. 34-36, pI. 11, fig. 8.
Spiroclypeus margaritata (Sch1umberger) var. umbonata Yabe and Hanzawa, 1929, p. 187-188, pI.
124, figs. 5-8.
Spiroclypeus higginsi Cole, 1939, p. 185, pI. 23, figs.
10-15, pI. 23, fig. 13; Hanzawa, 1957, p. 45, pI. 5,
figs. 1-6, 14; Cole, 1957a, p. 332, pI. 95, figs. 1-5,
pI. 109, fig. 16; Cole, 1957b, p. 763-764, pI. 239,
figs. 11-12, 14; Matsumaru, 1974, p.108, pI. 15,

948.

Early Miocene larger Foraminifera from Shikoku

figs. 1,5,8,12,18-19; Matsumaru, 1976, p.199,
pI. I, figs. 2, II, 16-17.

Description.- Test small, biconvex, bordered by a rather thin flange, central area 1.8 to
3.3 mm in diameter and 0.70 to 0.95 mm inthickness. The central portion has rather
evenly distributed small pillars having a
diameter of about 40 j1m. In some specimens the umbonal pustule becomes larger andhas a diameter of 270 to 350 j1m.
The embryonic chambers consist of a spherical protoconch followed by a reniform
deuteroconch; protoconch has diameter of
114 to 250 j1m, deuteroconch of 220 to 225
j1m.
The height of the embryonic chambers
in vertical section is 125 to 170 j1m. The
embryonic chambers are followed by an undivided operculine chamber and the following chambers are divided into heterostegine
chamberlets. The chamberlets near the
periphery of the test are rectangular withradial length of 96 to 166 j1m and with tangential length of 64 to 104 j1m.
The lateral chambers are low to rather
high, appressed between thick and thin floors
and roofs. There are 6 lateral chambers oneach side of the equatorial plane at the central
part of the test. Lateral chambers have alength of 73 to 208 j1m and a height of 6 to 27
j1m.
The floors and roofs have thickness of
20 to 94 j1m.
Remarks.- There is a confusion in thedefinition of the species of Spiroclypeus.
Although there is no equatorial section of thestudied specimen, the present form should be
identified as Spiroclypeus margaritata(Schlumberger, 1902) for the following reasons.
Tan (1937, p. 182) divided the species of
Spiroclypeus into the pustulate group, which
is characterized by features of prominentpillars of the central portion of the test, andthe reticulate group, which is characterized by
the development of an externally reticulate
appearance of the septa of the lateral chambers on the central part of the test. He regar-II

ded Spiroclypeus margaritata and S. tidoenganensis van der Vlerk, 1925, as belonging to
the pustulate group, while S. leupoldi van der
Vlerk, 1925, and S.orbitoideus H. Douville,
1905, belong to the reticulate group. Spiroclypeus margaritata from Kita-Daito-Zimahas, however, been revised as a member of thereticulate group by Hanzawa (1940, p.789790).
On the other hand, van der Vlerk (1925)
identified a fragile form of Spiroclypeus fromthe N aintoepo Beds, Tidoengsche Landen,
East Borneo as Spiroclypeus yabei, but Cole
(1969, p. 8) assigned S. yabei as a synonym of
S. margaritata. Though Hanzawa (1957, p.
46) distinguished S. tidoenganensis from S.
margaritata from the Tagpochou Limestone,
Saipan by the development of pustules andpillars traversing lateral chambers, Cole
(1969, p. 8-10) regarded the former as a synonym of the latter. Also, Krijnen (1931, p. 94)
described S. leupoldi, S. margaritata and S.
tidoenganensis from Loc. 633L of the upper
Naintoepo Beds, East Borneo, but Tan (1937,
pI. 2, fig. 12; pI. 4, figs. 6-7) identified themas S. margaritata. Spiroclypeus higginsi
Cole found from Guam by Cole (1939), waslater a synonym of S. leupoldi in Cole (1969,p. 9), but the former was regarded as a primitive species of the latter by Matsumaru (1974),
based on the specimens from the Mangagoy
and Bislig Formations, Mindanao Island,
Philippines.
From the above, it is inferred that the
Spiroclypeus in the west Pacific region should
be identified as Spiroclypeus margaritata.
Family Victoriellidae Chapman
and Crespin, 1930
Genus Victoriella Chapman
and Crespin, 1930
Victoriella conoidea (Rutten, 1914)
Figure 2-3

Carpenteria conoidea Rutten, 1914, p.47, pI. 7, figs.
6-9.
Carpenteria proteiformis Goes var. plecte Chapman,

Kuniteru Matsumaru et al.

121921, p. 320, pI. 51, fig. 3.
Carpenteria protei/ormis Goes. Cushman, 1921, p.
361-362, pI. 73, figs. 2-3; Yabe and Hanzawa,
1929, p. 178, pI. 15, fig. 15.
Victoriella conoidea (Rutten). Glaessner and Wade,
1959, p. 199, pI. I, figs. 1-5, pI. 2, figs. 1-5,7-10,
pI. 3, fig. 3, text-figs. 1-4; Hornibrook, 1968, p.
64, fig. II ; Chaproniere, 1984, p. 50, pI. 13, figs.
5a-b.

Description.- Test conical, with a small
attachment area near the apex; juvenile stage
free, a low trochospiral coil, the coiling being
high-spired, with three or four subsphericalchambers per whorl in the adult stage, thenerect conical in growth, 1.50 mm in lengthand 0.84 mm in width. Aperture umbilical,
surrounded by a thick lip. Wall coarsely
perforate, except for the imperforate region
around the aperture, with pillars which displace the pores; wall structure calcareous,bilamellid, consisting of an inner lining andan outer lamella.
Remarks.- The present form is referred toVictoriella conoidea (Rutten) by peculiar features of its trochospirally coil, which differs
in the direction of the axis of coiling from thenepionic stage, bilamellid walls and remarkable regular chamber shape and arrangement.
Glaessner and Wade (1959) described Carpenteria proteiformis Goes· and var. plecte
Chapman, 1921 as junior synonyms of Victoriella conoidea (Rutten). The authors follow
their opinion.
Family Rotaliidae Ehrenberg, 1839
Subfamily Pararotaliinae Reiss, 1963
Genus Pararotalia Y. Le Calvez, 1949
Pararotalia sp.
Figures 2-5-6

Description.- Test in a trochospiral coil,biconvex, periphery subacute, serrated, thedorsal side less vaulted than the ventral side,
0.7 to 0.8 mm in diameter and 0.5 to 0.6 mm
in thickness; chambers flat to elevated on the
spiral side, with an umbilical shoulder surrounding a large solid umbilical plug, smoothknob surrounded by a groove. The plug is

200 to 320,um in diameter. The wall isthick, calcareous and perforate.
Remarks.- The illustrations of the present
specimen show an oblique section of theventral side of the test and a vertical section.
They do not reveal the detailed structure, but
the present form is similar to Rotalia mecatepecensis Nuttall, which was described fromthe Tagpochau Limestone, Saipan by Hanzawa (1957, p. 59-60), or Rotalia floscula
Todd and Post from Bikini Drill Holes (Toddand Post, 1954). Judging from the description and illustration of Rotalia mecatepecensis and R. floscula, both species are within thegenus Pararotalia by Le Calvez (1949). As
such, the present form is identified as Pararotalia sp.

References cited
Adams, e.G. and Belford, D.l., 1974 : Foraminiferal
biostratigraphy of the Oligocene-Miocene limestones of Christmas Island (Indian Ocean).
Palaeontology, vol. 17, part 3, p.475-506.
- - , Gentry, A.W. and Whybrow, P.l., 1983:
Dating the terminal Tethyan event. In, Meulenkamp, 1.E. ed., Reconstruction of marine
paleoenvironments, Utrecht Micropal. Bull. 30, p.
273-298.
Bielle, M. and Caralp, M., 1967: Les Miogypsinidae
de deux coupes profondes des Landes. Bull.
Soc. Geol. France, ser. 7, vol.9, p.36-42.
Blow, W.H., 1969: Late Middle Eocene to Recent
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Planktonic Microfossils, Geneva, 1967, vol. I, p.
199-421. E.l. Brill, Leiden.
Bronnimann, P., 1940: Ober die tertHiren Orbitoididen und die Miogypsiniden von Nordwest
Marokko. Schweiz. Pal. Ges., Abh., vol. 63, art.
I, p. 1-113.
Butt, A.A., 1966: Late Oligocene Foraminifera from
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'
Chapman, F., 1921: Report of an examination ofmaterial obtained from a bore at Torquay.
Geol. Surv. Victoria, Record, no. 4, p. 315-324.
- - , and Crespin, 1., 1930: Rare Foraminifera fromdeep borings in the Victorian TertiariesVictoriella gen. nov., Cycloclypeus communis
Martin, and Lepidocyclina borneensis Provale.

948.

Early Miocene larger Foraminifera from Shikoku

Proc. Royal Soc. Victoria, new ser. 42, p. 110115.
Chaproniere, G.C.H., 1984: Oligocene and Miocene
larger Foraminiferida from Australia and New
Zealand. BMR Jour. Australian Geology andGeophysics, 188, p. 1-98.
Cole, W.S., 1939: Larger Foraminifera from Guam.
Jour. Paleont., vol. 13, no. 2, p. 183-189.
- - , 1954: Larger Foraminifera and smaller diagnostic foraminifera from Bikini drill holes. U.S.
Geol. Surv., Prof Paper 260-0, p. 569-608.
- - , 1957a: Larger Foraminifera of Saipan. Ibid.,
280-1, p. 321-360.
- - , 1957b: Larger Foraminifera from Eniwetok
Atoll drill holes. Ibid., 260-V, p. 743-784.
- - , 1969: Larger Foraminifera from deep drill
holes on Midway Atoll. Ibid., 680-C, p. 1-15.
Coleman, PJ., 1963: Tertiary larger Foraminifera of
the British Solomon Islands, southwest Pacific.
Micropaleontology, vol. 9, no. I, p. 1-38.
Cushman, J.A., 1921: Foraminifera of the Philippine
and adjacent seas. U.S. Nat. Mus., Bull. 100,
vol. 4, p. 1-608.
de Blainville, H.M.D., 1827: Manuel de malacologie
et de conchyliologie (1825). Paris: F.G.
Levrault.
Debourle, A. and Delman, M., 1955: Victoriellaaquitanica, nouveau foraminirere de l'oligocene
d'Aquitaine. Bull. Soc. Geol. France, ser. 6, vol.
5, p.47-49.
de Mulder, E.F.J., 1975: Microfauna andsedimentary tectonic history of the Ionian islands
and western Epirus (Greece). Utrecht Micropal.
Bull. 13, p. 1-139.
Douville, H., 1905: Les foraminireres dans Ie Tertiaire de Borneo. Bull. Soc. Geol. France, ser. 4,
vol. 5, p. 435-464.
- - , 1911: Les foraminireres dans Ie Tertiaire desPhilippines. Philippine Jour. Sci., vol. 6, no. 2,
p.53-80.
Drooger, e.W., 1952: Study of American Miogypsinidae. Thesis Univ. Utrecht, Vonk and Co.,
Zeist., p. 1-80.
- - , 1953: Some Indonesian Miogypsinidae.
Proc. Koninkl. Ned. Akad. Wetensch., ser. B, 56,
p. 104-123
Ehrenberg, e.G., 1839: tiber die Bildung derKreidefelsen und des Kreidemergels durch unsichtbare Organismen. Phys. Abhandl. Konig.
Akad. Wissensch. Berlin, 1838, p. 59-147.
Glaessner, M.F. and Wade, M., 1959: Revision of theforaminiferal family Victoriellidae. Micropaleontology, vol. 5, no. 2, p. 193-212.
Hanzawa, S., 1940: Micropaleontological studies ofdrill cores from a deep well in the Kita-DaitoZima (North Borodino Island). Jubilee Pub.
Comm. Prof H. Yabe's 60th Birthday, vol. 2.

13

Tohoku Imp. Univ., Sendai, p. 755-802.
- - , 1957: Cenozoic Foraminifera of Micronesia.
Geol. Soc. America, Mem. 66, p. 1-163.
Hashimto, W. and Matsumaru, K., 1975: Larger
Foraminifera from the Philippines. Part 3.
Limestones from Eastern Coastal Ranges ofNorth and Central Luzon. Geol. Palaeont.
Southeast Asia, vol. 16, p. 117-125.
- - and - - , 1978: Larger foraminifera from thePhilippines. Part 8. Larger foraminifera fromCentral Samar. Ibid., vol. 19, p. 81-88.
- - , - - , and Alcantara, P.M., 1982: Larger Foraminifera from the Philippines. Part 13. Larger
Foraminifera from the Trankalan Limestone andthe Escalante (Toboso) Formation, West of
Lanao River Valley, Northeastern Occidental
Negrons. Ibid., vol. 24, p. 31-38.
- - , - - , and Sugaya, M., 1981: Larger Foraminifera from the Philippines. Part II. On
the Coal Harbor Limestone, Cagraray Island,
Batan Island Group, Albay province. Ibid., vol.
22, p. 55-62.
Henson, F.R.S., 1937: Larger Foraminifera from
Aintab, Turkish Syria. Eclog. Geol. Helv., vol.
30, no. I, p.45-57.
Hornibrook, N. de 8., 1968: A handbook of New
Zealand Microfossils (Foraminifera andOstracoda). New Zealand Geol. Surv., Handbook, ser. 62, p. 1-136.
Krijnen, W.F., 1931: Het genus Spiroclypeus in het
Indo-Pacifische gebied. Geol.-Mijnb. Genoot.
Ned. Kolonien, Verc., Geol. ser., vol. 9, p. 77-111.
Le Calvez, Y., 1949: Revision des foraminireres
Lutetiens du Bassin de Paris. II. Rotaliidae etfamilles affines. Mem. Servo Carte Geol. Detailtee, France, p. I-54.
Matsumaru, K., 1971 : Studies on the genus Nephrolepidina in Japan. Sci. Rep., Tohoku Univ., 2nd.
ser. (Geol.), vol. 42, no. 2, p. 97-185.
- - , 1974: Larger Foraminifera from East Mindanao, the Philippines. Geol. Palaeont. Southeast Asia, vol.. 14, p. 101-115.
- - , 1976: Larger Foraminifera from the islands ofSaipan and Guam, Micronesia. In, Takayanagi
and Saito, T. eds., Progress in Micropaleontology.
Micropaleontology Press Spec. Pub!., New York.,
p.190-213.
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Bondoc Peninsula, Luzon, the Philippines andlarger Foraminifera. Geol. Palaeont. Southeast
Asia, vol. 23, p. 77-90.
- - and Kimura, K., 1989: Larger Foraminiferafrom the Eocene Shimizu and Misaki Formations
in Tosa Shimizu City, Kochi Prefecture, Shikoku, Japan. Trans. Proc. Palaeont. Soc. Japan,
N.S., no. 156, p. 255-269.

14

Kuniteru Matsumaru et al.

Myint Thein, Ogawa, Y. and Akiyama, T., 1991:
Finding of. Cretaceous radiolarians from theblock of sheared olistostrome in the southern partof the Shimanto Belt near the Ashizuri Cape,
Kochi Prefecture. Jour. Geol. Soc. Japan, vol.
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Myint Thein, 1992: Analysis of sedimentary facies
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Supergroup, SW Shikoku, lapan. Mem. Fac.
Sci., Kyushu Univ., ser. D, Earth and Planetary
Sciences, vol. 27, no. 2, p.75-99.
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p. I-Ill.
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von Niederlandisch Neu-guinea. Uitkomsten
Nederlandsch Nieuw-Guinea Expeditie in 1903,
Geol., vol. 6, p.21-51.
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Leiden, ser. I, vol. 6, p. 250-253.
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3, no. 3, p.45-61.
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vermicularis nov. sp. from Koetai in east Borneo.
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bei Miogypsina. Eclog. Geol. Helv., vol. 20, p.
323-330.
Todd, R. and Post, R., 1954: Smaller Foraminiferafrom Bikini drill holes. U.S. Geol. Surv., Prof
Paper 260-N, p. 547-568.
van der Vlerk, I.M., 1924: Miogypsina dehaartii nov.
spec., de Larat (Moluques). Eclog. Geol. Helv.,
vol. 18, p. 429-431.
- - , 1925: A study of Tertiary Foraminifera fromthe "Tidoengsche Landen" (E. Borneo). Dutch
East Indies, Dienst Mijnb., Wetensch. Meded.,no. 3, p. 13-32.
- - , 1964: Biometric research on European Lepidocyclines. Proc. Kon. Ned. Akad. Wetensch., Amsterdam, ser. B, vol. 67, no. 5, p. 1-10.
- - , 1966: Miogypsinoides, Miogypsina, Lepidocyclina et Cycloclypeus de Larat (Moluques).
Eclog. Geol. Helv., vol. 59, no. I, p. 421-429.
Vaughan, T.W., 1928: Subfamily Miogypsininae
Vaughan. In, Cushman, l.A., Foraminifera:
their classification and economic use. Spec. Pub!.
Cushman Lab. Foram. Res., vol. I, p. 354.
Yabe, H. and Hanzawa, S., 1925: A Lepidocyclinalimestone from Klias Peninsula, B.N. Borneo.
Geo!. Mijnb. Genoot. Nederl. Kolonjen, Verh.,
Geo!. ser., vol. 8, p. 617-632.
- - and - - , 1928: Tertiary foraminiferous rocksof Taiwan (Formosa). Proc. Imp. Acad.
Tokyo., vol. 4, no. 9, p. 533-536.
- - and - - , 1929: Tertiary foraminiferous rocksof the Philippines. Sci. Rep., Tohoku Imp.
Univ., 2nd ser. (Geol.), vol. II, no. 3, p. 137-190.

Ashizuri Cape JEmiIJljI, Misaki Formation =w.rr~I!, Ohki Beach :k1lezi'taJ¥, Shimanto Belt
1ZB7]-J-*, Shimizu Formation m7.K~I!, Takahata ~ffil, Tosa Shimizu City ±-fti:m7.KrIJ.

=~~D1~UEmillljl(7)5~*mP.l'it.J' i?ii"9 .QfJJJtIll=j:liITtit (7 =\=- ~ 7 /WD *~~fL!B: m7j(~
I! 0) ;k LU~m\O)h ~;fi, ii');fi$O):fiW(:1t3'E~1WiJ' Git L t.::kml~:j1.!R 5 ~ 5 :fl ,H'ml~:}L.!R 1 ~ 1 :fl~gC~ L t.:. :kml~:j1.!R Miogypsinoides dehaartii, Spiroclypeus margaritata
Victoriella conoideaiiJ4'fzlfFvi, fJJ:JtIl$~i!t (7=\'-;$1 .::.7:/:JtIl) ~~L, S;$:7"UJ1i,iJ'Gvif}]
66 '"(¥j~:* ~ ht.:. I'UfF$O) Nephrolepidina praejaponica vifJJ:JtIl$~i!t (7' /v T -{ tf ~ 7:/:JtIl) O)=w.rr~l!iJ' t:%l Gh '"( l' ~ t.:66, m7.K~111U:: =w.rr~I!O)I!w.f*li*vi, :flli66'"(J1I~ L
t.:f*li*vcJ; ~ ;:. c iJ':Jt:>E ~ hk.
t0JLffil~ • Myint Thein· 'Nil ~=N~

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 169, pp. 15-43, 9 Figs., April 30, 1993

949.

LOWER PERMIAN FUSULINACEANS FROM BAN PHIA,
CHANGWAT LOEI, NORTHEASTERN THAILAND*HISA YOSHI IGO, KA TSUMI UENO, and KA TSUO SASHIDA
Institute of Geoscience, University of Tsukuba, Ibaraki, 305

Abstract. Dark gray, thin-bedded limestones intercalated with calcareous sandstoneand shale are exposed at about 3 km southwest of Ban Phia, Changwat Loei, northeastern
Thailand. These limestones yield well-preserved fusulinaceans which indicate a late Early
Permian (late Yakhtashian) age and represent a new additional Permian fusulinacean faunain Thailand. The following species are new to science; Darvasites ingavati,
Pseudofusulina siamensis, Staffella ? o valis, Nankinella? /oeiensis, and Pseudoendothyra?
constricta.
Key words.
Yakhtashian.

Fusulinacean, Loei, Lower Permian, Saraburi Limestone, Thailand,

about 3 km southeast of Ban Phia, northeastof Loei (Figure 1).
We found an interesting fusulinacean faunain these limestone beds, which indicate a
Yakhtashian age of the Tethyan standard(late Early Permian). This fauna represents
a newly recognized Permian fusulinaceanzone in Thailand. We describe these materials in this paper.
All specimens identified in this paper aredeposited in the paleontological collectionsof the Institute of Geoscience, University ofTsukuba (lGUT).

Introduction
Fusulinaceans are one of the most intensively studied fossil groups in the Upper
Paleozoic of Thailand (e.g., Pitakpaivan,
1965; Igo, 1972; Toriyama, 1975; Toriyama and Kanmera, 1979; Ingavat andDouglass, 1981). Toriyama (1984) summarized these previous works and established 20
almost successive fusulinacean zones of late
Early Carboniferous to Late Permian age.
Although three or four faunal breaks wererecognized, these zones were regarded as thestandard biostratigraphy of the Upper Paleozoic in Southeastern Asia.
In November and December of 1990,
Hisayoshi Igo, Katsuo Sashida, and Kenichiro Hisada of the University of Tsukuba,
and Nikorn Nakornsri and Apsorn Ampornmaha of the Geological Survey Division,
Department of Mineral Resources, Thailand
made a field survey in the Changwat Loeiarea, northeastern Thailand; they collected
limestones exposed along a roadside gutter,'Received June 17, 1992; revised
accepted January 25, 1993

Acknowledgments
Financial support for our field survey wasprovided by a grant under the Monbusho
Research Program No. 02140210 to H.Igo.
We are grateful for the field guidance of Mr.
Nikorn Nakornsri, Senior Geologist of theGeological Survey Division, Department ofMineral Resources, Thailand. We also
thank Dr. Ken-ichiro Hisada and Miss
Apsorn Ampornmaha of the Institute of
Geoscience, University of Tsukuba for theirmanuscript15

Hisayoshi [go et al.

16

.-'

,,

.... -', ....

,

Locality',_',------*I---\...._ ...

\

....

,\

\

\ ,

"

,

o

1

2km ,

===-_-===__

f-I

L--

Figure 1.

To
Loei

,,

,,

I

",

......

I\\.... \_- \-,
\

\

,

\

\

,

"

" ,

'"
\

,

\

,I

Map showing the fossil locality near Ban Phia, northeast of Loei, northeastern Thailand.

assistance with field work. The present
manuscript was linguistically benefitted by
Ms. Jean Thyfault.

Geologic setting
The Loei area is geologically called the"Loei Folded Belt" and is underlain by
Middle to Upper Paleozoic rocks and widely
overlain by thick Mesozoic non-marine strata
called the "Khorat Series." Recently, theUpper Paleozoic rocks exposed in this area
were divided into the Carboniferous Wang
Saphung Formation and Permian SaraburiGroup. The latter was further subdivided
into the Dan Sai Shale, Saraburi Limestone,
and Khao Luak Formation by Bunapas(1983) in descending order. According to

the geologic map "Changwat Loei," scale 1 :250,000 (Charoenpravat et al., 1976), thePermian rocks in this area are grouped as the
Ratburi Group and subdivided into the Pha
Dua, E-Lert, and Nam Mahoran Formations
in descending order. The Ratburi Group
has long been used as the Permian. lithologic
unit in Thailand, but it is now restricted to
the limestone exposed in PeninsularThailand, and the Saraburi Group (or Limestone) is commonly accepted to be in the Loei
Folded Belt (Bunapas, 1983).
The Nam Mahoran Formation (or Saraburi
Limestone) consists mainly of pale gray towhite, massive to thickly bedded, and fossiliferous limestone, and it overlies the Wang
Saphung Formation. Although actual stratigraphic contact between both formations has

949.

Lower Permian fusulinaceans from NE Thailand

never been reported, their relation is regarded
as conformable. The Nam Mahoran Formation constitutes high hills surrounded bysteep cliffs, and the Wang Saphung Formation topographically occupies the low land.
Furthermore, the Nam Mahoran yields Early
Permian fusulinaceans (Igo, 1972), brachiopods (Yanagida, 1967), and other fossils,
but the Wang Saphung yields Upper Carboniferous fossils (e.g., Igo, 1972; Yanagida,
1974, 1975; Toriyama et al., 1975). Subsequently, Fontaine et al. (1981) pointed out
the occurrence of Devonian and Carboniferous fossils in the north of the Loei area andsubdivided the Paleozoic rocks into the Ban
Sa Ngao and Ban Khok Phai Formations inascending order. These two formations areapparently equivalent with the Nong Dok
Bua and Wang Saphung Formations in thepresent area.
Fusulinacean limestones exposed along aroadside gutter near Ban Phia are alternated:-:.:.:-..'...: .... ::,-.. \ .... Loe 34:.-:.,<.:: .:.. ::.:':." .... Loe 35::' ':",:: :',:.:.:.: .: ::.... Loe 36
L0 e 3 7~.,...,...,-!--J.,-..;-,J....

','," <:: ....:. :-:. :::.:

]

m

17

with calcareous sandstone and shale; theexposed area is about 30 m thick (Figure 2).
On the basis of lithologic character, thissequence belongs to a part of the Wang
Saphung Formation, and newly obtained
paleontological data show that it apparently
represents a much higher sequence than that
exposed in Ban Sup, Ban Na Nam Man, along
the Huai Nam Kai River, north of Ban Phi a(lgo, 1972).
The present paleontological collections
were made at five different levels (Loe-34, 35,
36,37, and 38). Limestone beds are thinbedded (30-50 cm), black to dark brown,
mostly muddy fusulinacean and algal wackestone with thin intercalations of shale, but alimestone bed in the middle part (Loe-36) iscoarse-grained, muddy, and fusulinaceancrinoidal grainstone. As is discussed below,
the geologic age of the fusulinacean fauna
discriminated in the present study is Yakhtashian (late Early Permian), and this age ismuch younger than previous dating of the
Nam Mahoran Formation by means offusulinaceans and brachiopods (Asselian,
early Early Permian). Recently we havefound a Gzhelian (latest Carboniferous)
fusulinacean fauna in a limestone of the N am
Mahoran Formation exposed near Ban Na
Din Dam, east of Loei. These newpaleontological data show that the stratigraphic relationship between the Wang Saphungand Nam Mahoran Formations is not simple
superposition, but rather their upper andlower parts interfinger each other.

Appraisal of fusulinacean fauna,',',,::;. : .', .. "" .. :~ 1.... Loe 38:-::: .-:. 2CIJ

D 3Figure 2. Columnar section with indication ofsampling levels. I: muddy limestone, 2: calcareous sandstone, 3: covered.

Fusulinaceans are rather rare in the lower
levels (Loe-37, -38) but are abundant in theupper levels (Loe-34, -35, -36) of the section
studied in this paper (Table 1). The present
fusulinacean fauna is characterized by theabundant occurrence of the species belonging
to· the genera Staffella ?, Nan kin ella ?, Pamirina, Darvasites, and Pseudofusulina. The
genera Pamirina and Darvasites have beenHisayoshi Jgo et al.

18

known extensively in various areas of the
Tethyan n~alm. The present fauna is similar
to the Pamirina (Pamirina) darvasicaDarvasites contractus fauna reported fromthe upper Yakhtashian of Darvas (e.g., Kalmykova, 1967; Leven, 1970a, 1980) in the
joint occurrence of the two zonal species.
Pseudoreichelina darvasica Leven and speciesof Schubertella and Toriyamaia are also common in the Darvas fauna. A similar faunacharacterized by Pamirina and Darvasites
was reported from the Carnic Alps (Kahler
and Kahler, 1980), Qinghai (Sheng and Sun,1975), and Xiaodushan and Guangnan, eastern Yunnan (Zhou et al., 1987), both ofSouth China. In these districts, however, thefauna is characterized mainly by the occurrence of well-known species of schwagerinids
such as Chalaroschwagerina vulgaris (Schel1wien), Pseudofusulina kraffti (Schellwien),
and P. fusiformis (Schellwien). The
schwagerinids in the present fauna are represented mostly by Pseudofusulina
siamensis, sp. nov., which is characterized bya small-sized shell. Staffella? ovalis, sp.
nov. and other staffellids are rather common
in the present fauna, but they are rare in thefaunas reported from the above-mentioned
districts.
Toriyama (1984) summarized thefusulinacean biostratigraphy in Thailand andestablished 20 almost continuous

Table 1. List of fusulinaceans from Ban Phia, Changwat Loei.
Loe 38SpeciesPamirina (Pamirina) darvasica Leven
Pamirina (Levenia) sp.
Schubertella me/onica Dunbar and Skinner
S.
sp. A
S.
sp. B
Toriyamaia sp.

+

Minojapanella (Minojapanella) sp.
Darvasites contractus (Schellwien)D.
ingavati, sp. nov.
Pseudofusulina siamensis, sp. nov.
P.
sp. A
P.
sp. B
P.
sp. C
P.
sp. D

+

+

Nankinella? /oiensis, sp. nov.
Sphaerulina cf. croatica Kochansky-Devidi!
Pseudoendothyra? constricta, sp. nov.
P.?
sp.

+

++: common

Loe 36

Loe 35

Loe 34

++

+++

+++
+
++

++
+
+
+
+
++
++
+++

++
+
++

+

Pseudoreichelina darvasica Leven
Staffella? ova/is, sp. nov.
S.
sp. A
S.?
sp. B

+ + + . abundantLoe 37

+

+
+
++
+
+
++

+
+
+++
+

+
++
++
+

+ : rate

949.

Lower Permian fusulinaceans from NE Thailand

fusulinacean zones by his own and collaborators' studies. Among them, 14 zones are
regarded to be Permian, and he pointed out
three or four major faunal breaks in the
Sakmarian- Yakhtashian, Dzhulfian, andprobably in the uppermost Dorashamian.
The Sakmarian-Y akhtashian interval is represented in between the Asselian Triticites
ozawai-Paraschwagerina yanagidai Zone
(Igo, 1972) and the upper Yakhtashian Misellina otai-M. cfr. termieri Zone (Toriyama,1975). According to a recent study by
Watanabe (1991), however, the Triticites ozawai-Paraschwagerina yanagidai Zone
should be correlated with the latest Carboniferous (Gzhelian) rather than the earliest Permian (Asselian). Ingavat and Douglass(1981) reported the occurrence of Monodiexodina sutschanica (Dutkevitch) and M.
shiptoni (Dunbar) from northwest Thailand,
and they correlated this fauna with those of
the post-Artinskian beds of the Salt Range,
Pamirs, and others. The occurrence of thegenus Monodiexodina is restricted mostly tocalcareous sandstone or coarse-grained,
siliceous limestone interbedded witharenaceous rocks. It is entirely absent fromor very rarely occurring in ordinary limestonefacies; therefore, the biostratigraphic relationship between the Monodiexodina Zone
and other fusulinacean zones established inlimestone is not clear. Toriyama (1984)
placed the Monodiexodina shiptoni Zone ofnorthwest Thailand in the upper Yakhtashian, but this zone may represent an age
younger than Yakhtashian. The Misellina
otai-M. cfr. termieri Zone also represents ayounger age than Toriyama's proposed one
and can be correlated with the lower Bolorianbased on the first appearance of Misellina(s.s.) in other Tethyan regions (e.g., Ueno,
1991b).
As mentioned above, the fusulinaceanzones apparently indicating the Asselian to
Yakhtashian are absent in Thailand according to previous reports. The present upper
Yakhtashian fauna, however, is a new addi-

19

fion to the fusulinacean biostratigraphic unitsof Thailand and fills in a part of the Early
Permian interval.
Systematic paleontology

Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea von Moller, 1878
Family Ozawainellidae Thompson
and Foster, 1937
Subfamily Ozawainellinae Thompson
and Foster, 1937
Genus Pamirina Leven, 1970
emend. Kobayashi, 1977

Discussion. -There are two diverse opinions on the taxonomic position of the genus
Pamirina. One is that Pamirina belongs to
the family Ozawainellidae (e.g., Leven,
1970a) and the other that it belongs to thefamily Staffellidae (e.g., Rozovskaya, 1975).
This problem is worth discussing as it also
concerns the classification and phylogeny of
the higher multiapertural fusulinaceans, thefamilies Verbeekinidae and Neoschwagerinidae.
The genus Pamirina was established byLeven (1970a) with the occurrence ofthe typespecies of Pamirina darvasica Leven in theupper Yakhtashian (Artinskian) Safetdaron
Suite of southwestern Darvas. Based on theshell shape and spirothecal composition,
Leven classified this genus in the family
Ozawainellidae and excluded staffellied genera as the ancestor of Pamirina because theirspirothecal structure is quite different fromthat of the latter. He considered an ozawainellid genus such as Eostaffella or
Pseudostaffella a possible ancestor of thegenus Pamirina, although their occurrence israre in the Upper Carboniferous and Lower
Permian.
In the interpretation of Rozovskaya (1975),
Pamirina is morphologically intermediate
between the genera Pseudoendothyra andMisellina, although the spirothecal structure

20

Hisayoshi Jgo et al.

of Pseudoendothyra is different from those of
the other two genera. She regarded Pamirina as belonging to the family Staffellidae.
Kobayashi (1977) first pointed out the existence of two distinct groups in Pamirina.
One has a thickly lenticular shell, and theother has a subspherical shell. The spirotheca of the former consists of a single structureless layer in the inner volutions, and a tectumand lower, less dense layer (protheca) in theouter ones. The other subspherical type hasa spiro theca consisting of a single structureless layer in the inner volutions; a tectumand lower, less dense layer in the middle
ones; and a tectum and fine alveolar keriotheca in the outer volutions. He interpreted
the former type to be more primitive than thelatter, but their stratigraphic occurrence was
not definitely confirmed. Kobayashi (op.
cit.) assigned the genus Pamirina to the family Ozawainellidae and insisted that the family Verbeekinidae is the direct descendant of
the family Ozawainellidae.
Recently, Rauser-Chernoussova (1988) described some "pamirinids" from the Artinskian of the Caspian Basin in Central Asiaand emphasized that the genus Pamirina issurely referable to the family Staffellidae
rather than to the Ozawainellidae, because its
spirothecal composition is identical with thatof staffellids. The Caspian "pamirinids",
however, have a spirotheca completely replaced by secondary mineralization, which isquite different in composition compared withthat of original Pamirina described by Leven(1970a). Accordingly, all of the Caspian"pamirinids" of Rauser-Chernoussova (1988)
should be referred to such staffellid genera as
Staffella, Sphaerulina, or Pseudoendothyra.
Very recently, Ueno (1991a) studied materials collected from the Akiyoshi LimestoneGroup of western Japan and split a newsubgenus Levenia from Pamirina. This newsubgenus does not have any fine alveolar
layer in the spiro theca. He explained that
Levenia is an ancestor of Pamirina (s.s.)based on the previously mentioned spiroth-ecal structure and stratigraphic occurrence.
According to Ueno (1991a), both Pamirina(s.s.) and Levenia have a microgranularspirotheca similar to that of the ozawainellids. No evidence of secondary mineralization has been observed in their spirotheca.
All specimens referable to the genus Pamirina (s.l.) studied herein have a microgranularspirotheca (see Figures 3-3b, 3-9b, and 3lOb), whereas staffellids (e.g., Staffella?
ovalis, sp. nov. and others), even in the samethin section, have a light-brownish spirotheca
which has been completely replaced by secondary mineralization. This suggests that the
spirothecal compositions of Pamirina (s.l.)and staffellids fundamentally differ. Consequently, it is invalid to include the genus
Pamirina in the family Staffellidae. Judging
from the spirothecal composition and generalshell shape, it seems most suitable, as Leven(1970a) originally considered, that the genus
Pamirina should be referred to the family
Ozawainellidae.
Huang and Zeng (1984) proposed the subgenus Nanpanella for the genus Pamirina,
with the type of Pamirina (Nanpanella) laxa
Huang and Zeng from the Lower PermianLonglinian of Longlin, Guangxi of South
China. Nanpanella, however, has a spirotheca replaced by secondary mineralization andis apparently assignable to the family
Staffellidae (Sheng et al., 1988).
Subgenus Pamirina Leven, 1970
Pamirina (Pamirina) darvasica
Leven, 1970
Figures 3-1-9,4-1-33
Pamirina darvasica Leven, 1970a, p. 23-24, pI. I, figs.
1-9, 23-24 (not pI. 1, figs. 10-11); Rozovskaya,
1975, pI. 35, figs. 4-5 (same as pI. I, figs 1-2 ofLeven, 1970a); Kobayashi, 1977, p. 14-15, pI. 2,
figs. la-b, 2-3; Liu et al., 1978, p. 80, pI. 18, figs.
8,12; Kahler and Kahler, 1980, p. 187-188, pI. 3,
fig. 8 (not pI. 3, fig. 7); Loeblich and Tappan,
1988, pI. 295, fig. 11 (not pI. 295, fig. 12, same as
pI. I, fig. I of Leven, 1970a); Xi a, in Ding et al.,1991, pI. 10, fig. 22.

949. Lower Permian fusulinaceans from NE Thailand

~
",
~~
.

.

~:~

Figure 3. 1-9. Pamirina (Pamirina) darvasica Leven, 1- 3a, 4- 7 : axial sections, IGUT-KU0231,
KU0232, KU0233, KU0234, KU0235, KU0236, KU0237, 8, 9a: sagittal sections, IGUT-KU0263,
KU0264, x 40, 3b, 9b : enlarged parts of 3a and 9a, x 100. 10. Pamirina (Levenia) sp., lOa : axial
section,IGUT-KU0273, x 40, lOb: enlargement of lOa, X 100. 11-13; Toriyamaia sp., 11 : tangential
section, IGUT-KU0274, 12: sagittal section, IGUT-KU0275, 13: axial section, IGUT-KU0276, X 30.
14 : Minojapanella (Minojapanella) sp., diagonal section, IGUT-KU0277, x 40. 15-19 : Schubertella sp.
B, axial sections, IGUT-KU0278, KU0279, KU0280, KU0281, KU0282, x 40. 20- 25 : Schubertellamelonica Dunbar and Skinner, axial sections IGUT-KU0283, KU0284, KU0285, KU0286, KU0287,
KU0288, X 40. 26,27. Schubertella sp. A, 26 : tangential section, IGUT-KU0289, 27: axial section,
IGUT-KU0290, x 40.

21

22

Hisayoshi [go et al.

949.

Lower Permian fusulinaceans from NE Thailand

Chinlingella chinlingensis Wang and Sun, 1973, p.
152-153,172, pI. I, figs. 12,17-32, pI. 3, figs. 1,5,
10.
Chinlingella nobilis Wang and Sun, 1973, p. 153-154,
173, pI. I, figs. 1-5,9-11.
Staffella aff. mb11eri Ozawa. Sakagami and Iwai,
1974, p. 54-55, pI. 4, figs. 1-13.
Pamirina orbiculoidea Sheng and Sun, 1975, p. 45-46,
pI. 1, figs. 11-14; Zhou et al., 1987, pI. 2, figs. 56; Ding et al., 1989, pI. 2, figs. 27-28.
Pamirina nobilis (Wang and Sun). Lin et al., 1977,
p. 20, pI. 3, figs. 21-22; Sun et al., 1983, p. 9, pI.
2, fig. 15 (same as pI. I, fig. 1 of Wang and Sun,1973); Xia et al., 1986, pI. 9, figs. 6, 8; Zhou etal., 1987, pI. 2, figs. 3-4; Ding et al., 1989, pI. 2,
fig. 21.
Pamirina chinlingensis (Wang and Sun). Sun et al.,1983, p.9, pI. 2, fig. 17 (same as pI. 1, fig. 19 ofWang and Sun, 1973); Nie and Song, 1983, pI. I,
figs. 23-24; Xia et al., 1986, pI. 10, fig. 9; Zhou
et al., 1987, pI. 2, figs. 1-2; Ding et al., 1987, pI.
I, figs. 4-5; Loeblich and Tappan, 1988, pI. 295,
figs. 8-1 0 (same as pI. 1, figs. 19-20, 31 of Wangand Sun, 1973); Sun and Zhang, 1988, pI. 3, figs.
7,15; Ding et al., 1989, pI. 2, figs. 23-26.
Pamirina (Pamirina) chinlingensis (Wang and Sun).
Huang and Zeng, 1984, pI. 4, fig. 7.
Pamirina (Pamirina) darvasica Leven. Huang andZeng, 1984, pI. 4, fig. 14; Ueno, 1991a, p.744745, figs. 3-1-7.
Pamirina angulata Xia, in Ding et al., 1991, p. 128,
pI. 9, figs. 8-9.
Pamirina adulta Xia, in Ding et al., 1991, p. 127-128,
pI. 9, figs. 10-11, 14.

Description.-Shell small and almost
spherical with slightly umbilicated axial
regions and broadly rounded periphery.
Mature specimens having 5 to 6 volutions,0.47 to 0.81 mm in length and 0.48 to 0.87 mm
in width. Form ratio ranges from 0.81 to
1.06, averaging 0.92 for 39 specimens.
Inner few volutions lenticular in shape andalmost planispirally to skew coiled. Shell
becomes expanded and spherical beyond thethird or the fourth volution. Radius vectorsof the first to the fifth, and the fifth and a half
volution of well oriented axial section (Fig-23

ure4-2) 0.03,0.05,0.12,0.19,0.30, and 0.36mm, and form ratios 0.56,0.63,0.92,0.91,
0.89, and 0.95, respectively.
Pro loculus small and spherical. Its outside diameter ranges from 0.030 to 0.055 mm,
averaging 0.043 mm for 55 specimens.
Spirotheca thin and composed of a single
structureless layer in inner volutions, of atectum and lower, less dense layer (protheca)in middle ones, and of a tectum and clearly
defined fine alveolar keriotheca in outer volutions. Thickness of spiro theca of the first to
the fifth and the fifth and a half volution of
the above-mentioned specimen 0.005, 0.010,
0.015,0.025,0.030, and 0.025 mm, respectively.
Septa straight and plane. Tips of septa insagittal section slightly swollen. Septal
sutures commonly depressed. Septal countsof the first to the fifth volution of typicalsagittal section (Figure 4-31) 8,9,12,13, and15, respectively. Chomata low, broad, commonly asymmetrical, and developed in outer
volutions. Tunnel angles of the third to thefifth volution of the above-mentioned axial
section 24,37, and 49 degrees, respectively.
Remarks.-Broad morphological variations, especially in the coiling nature of thejuvenarium, are observed in the Loei specimens. Form having a straight axis of coiling(e.g., Figure 4-6) is identical with Pamirina(P.) darvasica originally described by Leven(1970a) from the Yakhtashian of Darvas,
whereas that having a rotated axis of coilingin the juvenile volutions (e.g., Figure 4-1) isquite similar to Pamirina (P.) orbiculoidea
proposed by Sheng and Sun (1975) from the
Qinghai district of South China. The Loei
specimens seem to comprise a single population, hence the former species, darvasica, isconsidered to be a senior synonym with thelatter one, orbiculoidea.

~
Figure 4. 1-33. Pamirina (Pamirina) darvasica Leven, 1-25: axial sections, IGUT-KU0238,
KU0239, KU0240, KU0241, KU0242, KU0243, KU0244, KU0245, KU0246, KU0247, KU0248, KU0249,
KU0250, KU0251, KU0252, KU0253, KU0254, KU0255, KU0256, KU0257, KU0258, KU0259, KU0260,
KU0261, KU0262, 26-33: sagittal sections, IGUT-KU0265, KU0266, KU0267, KU0268, KU0269, KU0270,
KU0271, KU0272, x40.

Hisayoshi [go et at.

24

Pamirina (P.) adulta and P. (P.) augusta
described by Xia (in Ding et at., 1991) fromthe Lower Permian Nagatoella-Darvasites(Nd) and Pamirina globosa (Pv) Subzones of
the Pamirina Zone (upper part of the Beiyangshan Limestone) in Zhen'an, Shaanxi are
identical with P. (P.) darvasica Leven.
Materials.-Axial sections: IOUTKU0231-KU0234, KU0236-KU0245,
KU0247-KU0250, KU0252, KU0253,
KU0255-KU0262 from Loe-34; KU0254
from Loe-35; KU0235, KU0246, KU0251
from Loe-36. Sagittal sections: IOUTKU0263, KU0265, KU0267-KU0271 fromLoe-35, KU0266 from Loe-36.
Subgenus Levenia Ueno, 1991
Pamirina (Levenia) sp.
Figure 3-10

Description.-Shell minute and thickly
lenticular with slightly depressed axial
regions and broadly rounded periphery.
Specimen having 5 volutions, 0.34 mm inlength and 0.49 mm in width, giving a form
ratio of 0.70. Axis of coiling slightly rotatedin inner volutions. Radius vectors of thefirst to the fifth volution 0.02,0.04,0.07,0.12,
and 0.19 mm, and form ratios 0.57,0.73,0.70,
0.72, and 0.73, respectively. Proloculus minute and spherical, being 0.035 mm in outside
diameter. Spiro theca very thin and composed of a single structureless layer in inner
volutions, but of a tectum and lower, lessdense layer (protheca) in outer ones. Thickness of spirotheca of the first to the fifth
volution 0.005,0.005,0.010, 0.Ql5, and 0.020
mm, respectively. Very small chomata discernible in outer volutions.
Remarks.- The present species somewhat
resembles Pamirina (Levenia) tethydis originally described by Kobayashi (1977) from the
Hanagiri Limestone of the Kwanto Mountains, central Japan. The former can be
distinguished from the latter in having aslightly rotated axis of coiling in the inner
volutions. The specific identification, how-ever, is postponed because only one specimen
was obtained in this study.
Material.-Axial section: IOUT-KU0273
from Loe-34.
Family Schubertellidae Skinner, 1931
Subfamily Schubertellinae Skinner, 1931
Oen us Schubertella Staff and Wedekind, 1910
Schubertella melonica Dunbar and Skinner,
1937
Figures 3-20-25
Schubertella me/anica Dunbar and Skinner, 1937, p.
611-613, pI. 57, figs. 10-14; Chang, 1963, p.44,
pI. 1, figs. 9-10,13; Chen and Wang, 1983, p.
38-39, pI. 2, figs. 16-17; Nie and Song, 1983, pI.
2, fig. 11 ; Da and Sun, 1983, p. 16, pI. 25, fig. 13.
Schubertella ex gr. me/anica Dunbar and Skinner.
Leven and Scherbovich, 1978, p. 85, pI. 1, fig. 6.

Description.-Shell small and fusiform withbluntly pointed polar ends and straight orslightly convex lateral slopes. Mature shellof 4 Yz to 5 volutions 0.70 to 1.10 mm inlength and 0.45 to 0.61 mm in width. Formratio varies from 1.57 to 1.80, averaging 1.68
for 8 specimens.
Inner 1 Yz to 2 volutions skew coiled witha short axis of coiling. Shell expands gradually beyond the third volution. Radius
vectors of the first to the fifth volution ofillustrated axial section (Figure 3-24) 0.05,
0.08,0.14,0.22, and 0.34 mm, and form ratios
0.89,0.88, 1.56, 1.74, and 1.49, respectively.
Proloculus small and spherical, measuring0.035 to 0.050 mm in outside diameter.
Spirotheca thin and composed of a tectumand lower, less dense layer. Thickness of
spirotheca of the first to the fifth volution of
the above-mentioned specimen 0.010,0.015,
0.025, 0.030, and 0.035 mm, respectively.
Septa unfluted and almost plane. Massive
chomata developed in outer vol utions. Tunnel path straight. Tunnel angles of the third
to the fifth vol uti on of axial section
mentioned above 29, 38, and 23 degrees, respectively.
Remarks.- The Loei specimens are almost

949.

Lower Permian fusulinaceans from NE Thailand

identical with the original ones described by
Dunbar and Skinner (1937) from the Lower
Permian of Texas, except for a slightly smaller
shell in the former.
Materials.-Axial sections: IGUTKU0286 from Loe-34; KU0283-KU0285,
KU0287, KU0288 from Loe-36.

Schubertella sp. A
Figures 3-26-27

Descriptive remarks.-Shell small and elongate fusiform with bluntly pointed poles.
Axial section of 4 Yz volutions (Figure 3-27)
1.21 mm in length and 0.49 mm in width,
giving a form ratio of2.50. Inner 2 volutions
skew coiled. Radius vectors of the first to
the fourth and the fourth and a half volution
0.05,0.07,0.13,0.21, and 0.27 mm, and form
ratios 0.67, 1.00, 1.46,2.05, and 2.32, respectively. Proloculus small and spherical, being
0.045 mm in outside diameter. Spirothecathin and composed of a tectum and lower, lessdense layer. Thickness of spiro theca of thefirst to the fourth volution 0.010,0.010,0.020,
and 0.025 mm, respectively. Chomata asymmetrical and developed in outer volutions.
Tunnel angles of the second to the fourth
volution 19,29 and 42 degrees, respectively.
This species resembles Schubertella kingi
Dunbar and Skinner, but differs from thelatter in having a slightly larger shell.
Materials.-Axial section: IGUTKU0290. Tangential section: IGUTKU0289. Both from Loe-36.
Schubertella sp. B
Figures 3-15-19

Remarks.-Schubertella sp. B can be easily
distinguished from S. melonica Dunbar andSkinner in having a smaller shell and form
ratio.
Materials.-Axial sections: IGUTKU0278, KU0280-KU0282 from Loe-34;
KU0279 from Loe-36.

25

Genus Toriyamaia Kanmera, 1956
Toriyamaia sp.
Figures 3-11-13

Remarks .-Several poorly oriented specimens were obtained and identified with Toriyamaia sp. The specific identification is
postponed until better materials are available.
Materials.-Sagittal section: IGUTKU0275 from Loe-34. Tangential section:
IGUT-KU0274 from Loe-34. Oblique section: IGUT-KU0276 from Loe-36.
Family Boultoniidae Skinnerand Wilde, 1954
Genus Minojapanella Fujimotoand Kanuma, 1953
Subgenus Minojapanella Fujimotoand Kanuma, 1953
Minojapanella (Minojapanella) sp.
Figure 3-14

Remarks.-Although the present material
is poorly preserved, the spirothecal composition and intensely fluted septa of the specimen
indicate it to be referable to the genus Minojapanella.
Material.-Diagonal section: IGUTKU0277 from Loe-36.
Family Schwagerinidae Dunbarand Henbest, 1930
Subfamily Schwagerininae Dunbarand Henbest, 1930
Genus Darvasites Miklukho-Maklay, 1959
Darvasites contractus (Schellwien, 1909)
Figures 5-1-13

Fusulina contracta Schellwien (MS). Staff, 1909, pI.
7, fig. 8.
Fusulina contracta Schellwien, 1909, p. 159-163, pI.
13, figs. 9-12.
Triticites contractus (Schellwien). Chen, 1934a, p.
34-35, pI. 5, fig. 13, pI. 7, figs. 1-4; Chen, 1934b,
p. 41-42, pI. 7, figs. 2-3.
Triticites sinensis Chen, 1934a, p. 36-38, pI. 7, figs. 8,
12.
Hemifusulina contracta (Schellwien). Chang, 1963,
p.49-50, pI. 3, figs. 1-3, 10; Sheng, 1966, p.73,

26

Hisayoshi /go et al.

41\)
9Figure 5. 1- 13. Darvasites contractus (Schellwien), 1,5,7,8, 10 : axial sections of megalosphericform, IGUT-KU0291, KU029s, KU0297, KU0298, KU0300, 2, 6,12 : slightly oblique axial sections ofmegalospheric form, IGUT-KU0292, KU0296, KU0302, 3,4: sagittal sections of megalospheric form,
IGUT-KU0293, KU0294, 9,11: tangential sections of megalospheric form, IGUT-KU0299, KU0301 ,x iS, 13 : axial section of microspheric form , IGUT-KU0303, X 30. 14- 20. Darvasiles ingavati Igo,
Ueno and Sashida, sp. nov. , 14: axial section of the holotype, IGUT-KU0304, 15: sagittal section ofparatype, IGUT-KU030s, 16- 20: axial sections of pa ra types, IGUT-KU0306, KU0307, KU0308, KU0309,
KU0310, X IS. 21,22 : Pseudofusulina sp. A, axial sections IGUT-KU0311 , KU0312, 21 : X IS, 22 :
X 10.

949.

Lower Permian fusulinaceans from NE Thailand

pI. 10, fig. 4 (same as pI. 7, fig. 1 of Chen, 1934a).
Triticites (Darvasites) cf. contractus (Schellwien),
Kochansky- Devide, 1964, pI. 18, fig. 5.
Darvasites ex gr. contractus (Schellwien). Ramovs
and Kochansky-Devide, 1965, p. 331-332, pI. 4,
fig. 1 (same as pI. 18, fig. 5 of Kochansky-Devide,
1964).
Hemifusutina sinensis (Chen). Sheng, 1966, p. 75, pI.
7, fig. 7 (same as pI. 7, fig. 8 of Chen, 1934a).
Darvasites contractus (Schellwien). Leven, 1967, p.
136, pI. 4, figs. 5-6; Kochansky-Devide, 1970, p.
196,231-232, pI. 5, figs. 9-11; Ramovs andKochansky-Devide, 1979, pI. 3, fig. 12; Kahler
and Kahler, 1980, p. 195, pI. 5, fig. 2; Kahler,
1983, p.59, pI. 7, fig. 7; Kahler, 1985, p.49-51,
pI. 3, figs. 2-4
Triticites (Darvasites) contractus (Shellwien). Kalmykova, 1967, p. 166-168, pI. 2, figs. 9-13.
Eoparafusulina contracta (Schellwien). Sheng andSun, 1975, p. 38-39, pI. 5, fig. 11 ; Lin et at.,1977, p.45, pI. 9, fig. 15 (same as pI. 7, fig. I ofChen, 1934a); Da and Sun, 1983, p. 98, pI. 8, fig.
16; Xia and Zhang, 1985, p. 130-131, pI. 28, fig.
11.

Eoparafusulina belluta Skinner and Wilde. Shengand Sun, 1975, p. 38, pI. 5, figs. 1-3.
Eoparafusulina sinensis (Chen). Wang et at., 1982,
p. 81-82, pI. 19, figs. 3,5 (3: same as pI. 7, fig. 8of Chen, 1934a).
? Eoparafusulina contracta (Schellwien). Liu et at.,1978, p. 67, pI. 15, fig. 12; Wang et at., 1982, p.
81, pI. 19, fig. 1 ; Chen and Wang, 1983, p. 83, pI.
14, figs. 13, 17; Wang, 1987, pI. 7, fig. 14; Luoet at., 1989, pI. 2, fig. 7.
not Triticites (Rauserites) contractus (Schellwien).
Rozovskaya, 1958, p. 101, pI. 9, figs. 10-12.

Description.-Shell of megalospheric form
small for the genus and ellipsoidal withrounded polar ends and almost straight ventral part. Microspheric form smaller thanmegalospheric one. Mature specimens ofmegalospheric form having 7 to 8 volutions,2.25 to 3.95 mm in length and 1.30 to 1.80 mm
in width. Form ratio varies from 1.73 to
2.51, averaging 1.94 for 7 specimens.
Microspheric form of 7 Yz volutions (Figure5-13) 1.38mm in length and 0.83mm inwidth, giving a form ratio of 1.67.
In megalospheric form, axis of coilingstraight throughout growth and inner 3 or 4
volutions rather tightly coiled. In
microspheric form, inner few vo1utions tight-27

ly and skew coiled. Radius vectors of thefirst to the eighth volution of typical axial
section of megalospheric form (Figure 5-1)
0.09, 0.13, 0.19, 0.26, 0.37, 0.49, 0.68, and 0.82mm, and form ratios 1.44, 1.85, 1.90, 2.11,
2.08,2.27,2.04, and 2.02, respectively.
Radius vectors of the first to the seventh
volution of ill ustrated axial section of microspheric form 0.04,0.07,0.10,0.14,0.20,0.27,
and 0.38 mm, and form ratios 1.00, 1.00, 1.20,
1.50, 1.80, 1.89, and 1.71, respectively.
Proloculus of megalospheric form smalland spherical. Its outside diameter ranges
from 0.100 to 0.140 mm, averaging 0.115 mm
for 19 specimens. Proloculus of microspheric form spherical and minute, being 0.030 mm
in outside diameter.
Spirotheca composed of a tectum and lower
structure less layer in inner few volutions, butof a tectum and coarse keriotheca in outerones in both forms. Thickness of spirothecaof the first to the eighth volution of theabove-mentioned megalospheric specimen
0.010,0.010,0.025,0.030,0.030,0.065,0.075,
and 0.055 mm, respectively.
Septa moderately and regularly fluted
throughout length of shell except for central
part. Septal counts of the first to the eighth
volution of one illustrated sagittal section(Figure 5-4) 5,9,13?, 15, 19, 18, 16, and 18,
respectively.
Chomata massive and well developed in all
volutions except for the last one in somespecimens. Tunnel path straight. Tunnel
angles of the first to the seventh volution of
the above-mentioned megalospheric specimen
29,44, 34, 32, 38, 40, and 32 degrees, respectively. Axial fillings absent.
Remarks.-Darvasites contractus was originally described by Schellwien (1909) fromthe Lower Permian of Darvas. Subsequently, Kalmykova (1967) also described andill ustrated this species from Darvas. The
Loei specimens are identical with
Schellwien's original and Ka1mykova's
topotype specimens, and a further discussionis unnecessary.

Hisayoshi [go et al.

28

Darvasites sinensis proposed by Chen(l934a) from the lower part of the Chuanshan
Limestone of southern Jiangsu (Kiangsu) has
an ellipsoidal shell, massive chomata
throughout growth, and moderately and regularly fluted septa; it is considered to be ajunior synonym of Darvasites contractus(Schellwien).
Dimorphism is recognized in the Loei specimens. The microspheric individual is characterized by a skew-coiled juvenarium andhas a smaller shell and much smaller
proloculus than do the megalospheric individuals.
Materials.-Axial sections of megalospheric form: IGUT-KU0291, KU0297 from Loe34; KU0295, KU0298, KU0300 from Loe-36.
Slightly oblique axial sections of megalospheric form: IGUT-KU0292, KU0296 fromLoe-34; KU0302 from Loe-36. Sagittal sections of megalospheric form: IGUT-KU0294
from Loe-34; KU0293 from Loe-36. Tan-

Table 2. Measurements of Darvasites ingavati Igo, Ueno and Sash ida, sp. nov (in mm)
Radius vector
FigureReg. no.

Length

Width

F.R.

D.P.
1

2

3

4

5

6

1

IGUT-KU0304

5-14

2.88

1.55

1.85

0.110

0.09

0.13

0.21

0.34

0.52

0.69

2

IGUT-KU0306

5-16

2.90

1.45

2.00

0.115

0.09

0.14

0.22

0.35

0.55

0.77

3

IGUT-KU0307

5-17

2.45

1.38

1.78

0.115

0.08

0.14

0.19

0.31

0.50

0.71

4

IGUT-KU0308

5-18

2.58

1.35

1.91

0.110

0.09

0.16

0.27

0.47

0.70

5

IGUT-KU0309

5-19

2.80

1.35

2.07

0.125

0.12

0.19

0.33

0.53

0.73

6

IGUT-KU03lD

5-20

2.00

1.00

2.00

0.120

0.09

0.15

0.23

0.37

0.53

Form ratio

Thickness of spirotheca

1

2

3

4

5

6

1

2

3

4

5

6

1

1.11

1.61

2.09

2.03

1.98

1.99

O.OlD

0.015

0.030

0.065

0.070

0.080

2

1.55

1.79

2.00

2.00

1.98

1.96

0.015

0.D25

0.045

0.060

0.070

-

3

1.13

1.14

1.50

1.66

1.56

1.68

0.D15

0.020

0.030

0.055

0.070

-

4

1.56

1.81

1.68

1.48

1.81

0.015

0.025

0.040

0.070

0.080

5

1.33

1.50

1.65

1.64

1.87

-

0.030

0.045

0.065

0.060

6

1.56

1.67

1.96

2.00

1.96

0.010

0.030

0.030

0.055

0.070

Tunnel angle (degrees)
1

2

3

4

5

6
23

1

32

25

32

44

39

2

44

32

40

36

36

-

3

-

-

42

48

45

-

4

-

44

51

42

44

5

-

-

38

25

-

6

-

30

33

48

-

F.R.: Form ratio
D.P.: Diameter of pro10cu1us

949.

Lower Permian fusulinaceans from NE Thailand

gential sections of megalospheric form:
IGUT-KU0299 from Loe-34; KU0301 fromLoe-36. Axial section of microspheric form:
IGUT-KU0303 from Loe-34.

Darvasites ingavati Igo, Ueno
and Sashida, sp. nov.
Figures 5-14-20

Diagnosis.-Small Darvasites with oval
shell, broadly rounded median portion, small
rudimentary chomata, and regularly fluted
septa.
Description.-Shell small for the genus andoval in shape with bluntly pointed axial
regions and broadly rounded median portion.
Mature specimens of 5 to 6, rarely 6 Yz volutions 2.00 to 2.90 mm in length and 1.00 to
1.55 mm in width. Form ratio ranges from1.78 to 2.07, averaging 1.94 for 6 specimens.
The holotype of 6 Yz volutions 2.88 mm inlength and 1.55 mm in width with a form
ratio of 1.85.
The first volution rather tightly coiled.
Axis of coiling straight throughout growth.
Radius vectors of the first to the sixth volution ofthe holotype 0.09, 0.13, 0.21, 0.34, 0.52,
and 0.69 mm, and form ratios 1.11, 1.61,2.09,
2.03, 1.98, and 1.99, respectively.
Proloculus small and spherical. Its outside diameter ranges from 0.100 to 0.125 mm,
averaging 0.114 mm for 7 specimens.
Spirotheca composed of a tectum and lower
structureless layer in inner few volutions, butof a tectum and coarse keriotheca in outerones. Thickness of spiro theca of the first to
the sixth vo1ution of the ho10type 0.010,
0.Ql5, 0.030, 0.065, 0.070, and 0.080 mm, respectively.
Septa moderately and regularly fluted
throughout length of shell except for central
part. Intensity of septal fluting increases
toward polar ends. Septal counts of the firstto the sixth volution of sagittal section ofparatype (Figure 5-15) 6,11,10,14,18, and24?, respectively.
Chomata small but developed in all volu-

29

tions except for the last one in some specimens. Tunnel path straight. Tunnel anglesof the first to the sixth volution of theholotype 32, 25, 32, 44, 39, and 33 degrees,
respectively. Axial fillings absent.
Remarks.- The present new species is characterized by an oval shell and rather small
chomata. It can be distinguished easily from
Darvasites contractus (Schellwien) by its
shell shape and development of chomata.
Darvasites ingavati, sp. nov. somewhat
resembles D. citrus originally described by
Ramovs and Kochansky-Devide (1965) fromthe Permian of Croatia. The former, however, differs from the latter in having a larger
shell.
Etymology.- This specific name is dedicated to Mrs. Rucha Ingavat, former paleontologist of the Geological Survey Division,
Department of Mineral Resources, Thailand,for her contribution to the study offusulinaceans in Thailand.
Materials.-Axial section of the holotype :IGUT-KU0304. Axial sections of paratypes: IGUT-KU0306-KU0310. Sagittal
section of paratype: IGUT-KU0305. All
specimens from Loe-36.
Measurements.-See Table 2.
Genus Pseudofusulina Dunbar
and Skinner, 1931
Pseudofusulina siamensis Igo, Ueno andSashida, sp. nov.
Figures 6-1-15

Diagnosis.-Pseudofusulina having a small
elongate fusiform shell with tightly coiled
juvenarium and small chomata in all volutions except for the last one. Septa
moderately and regularly fluted.
Description.-Shell small for the genus andelongate fusiform with bluntly pointed polarends and almost straight lateral slopes.
Mature shell having 7 to 8 volutions, 4.88 to
6.70 mm in length and 2.10 to 2.73 mm inwidth. Form ratio ranges from 2.18 to 2.73,
averaging 2.35 for 7 specimens. The

30

Hisayoshi /go et al.

6

..
9

Figure 6. 1-15. Pseudofusulina siamensis Igo, Ueno and Sashida, sp. nov., 1: axial section of theholotype, IGUT-KU03l3, 2,5- 7,13: axial sections of paratypes, IGUT-KU0314, KU0317 , KU03l8,
KU0319, KU0325 , 3,4,9,10,15: sagittal sections of paratypes, IGUT-KU03IS, KU03l6, KU0321 ,KU0322, KU0327 , 8, 11: slightly oblique axial sections of paratypes, IGUT-KU0320, KU0323 , 12, 14:
tangential sections of paratypes, IGUT-KU0324, KU0326. 16,17. Pseudofusulina sp. B, 16: tangential
section, IGUT-KU0328, 17 : axial section, IGUT-KU0329. 18 : Pseudofusulina sp. D, slightly oblique
axial section, IGUT-KU0330. 19-23. Pseudofusulina sp. C, 19: tangential section IGUT-KU0331 , 2022: axial sections, IGUT-KU0332, KU0333 , KU0334, 23 : sagittal section, IGUT-KU033S . All X 10.

949.
Table 3.

Lower Permian fusulinaceans from NE Thailand

31

Measurements of Pseudofusulina siamensis Igo, Ueno and Sashida, sp. nov. (in mm)
Radius vector

Reg. no.

FigureLength

Width

D.P.

F.R.

I

2

3

4

5

6

7

8

I IGUT-KU0313

6-1

5.18

2.28

2.27

0.120

0.11 0.15 0.21 0.34 0.51 0.77 1.06

2 IGUT-KU0314

6-2

5.70

2.48

2.30

0.115

0.10 0.16 0.20 0.30 0.53 0.81

l.l5 1.39

3 IGUT-KU0317

6-5

4.88

2.15

2.27

0.125

0.10 0.15 0.23 0.36 0.59 0.86-

4 IGUT-KU0318

6-6

4.90

2.25

2.18

0.120

5 IGUT-KU0319

6-7

5.05

2.10

2.40

0.110

0.11 0.16 0.25 0.40 0.59 0.87 1.24
0.08 0.14 0.21 0.34 0.55 0.80 l.l3

6 IGUT-KU0325

6-13

5.25

2.28

2.30

0.150

0.10 0.16 0.24 0.34 0.54 0.79 1.03

Form ratio

Thickness of spirotheca

I

2

3

4

5

6

7

8

2

I

3

4

5

6

7

0.010 0.015 0.025 0.040 0.080 0.090 0.110

1.00

1.60

2.41

2.46

2.45

2.47

2.43

2 1.l0

1.25

1.76

2.29

2.13

1.99

1.91

3 1.30

1.67

1.76

2.11

1.88

2.08

-

4 1.00

1.24

1.50

1.66

1.80

2.06

1.96

5 1.75

2.07

2.41

2.43

2.58

2.54

2.41

0.015 0.025 0.030 0.045 0.090 0.110 0.010 0.020 0.035 0.035 0.060 0.130 0.120

6 l.l0

1.44

1.92

2.37

2.48

2.28

-

0.015 0.020 0.025 0.050 0.080 0.110 0.100

I

0.010 0.025 0.035 0.050 0.050 0.100 0.120 0.120
0.020 0.020 0.035 0.050 0.095 0.095

1.99

Tunnel angle (degrees)
I

2

3

4

5

6

7

I

-

46

44

50

53

47

30?

2

-

-

35

38

41

46

-

3

30

40

55

43

43

37

-

4

-

-

25

27

35

42

-

49

45

-

52

50

-

5

29

41

45

51

6

-

34

37

46

8

-

Septal counts
Reg. no.

FigureI

I IGUT-KU0315

6-3

2

8

3

4

5

6

7
-

8

12

15

15

18

21

II

14

15

19
21

2 IGUT-KU0316

6-4

7

II

3 IGUT-KU0321

6-9

8

11

13

14

18

4 IGUT-KU0322

6-10

8

12

18

17

21

-

5 IGUT-KU0327

6-15

7

II

12

17

18

21

32

Hisayoshi /go et al.

holotype of 7 vol ution s 5.18 mm in lengthand 2.28 mm in width, giving a form ratio of
2.27.
The first I or 2 volutions rather tightly
coiled. Beyond the seco nd or the third vo lution , shell expands uniformly and becomesfu siform to elongate fusiform. Axis of coiling straight throughout growth. Radius
vectors of the first to the seventh vol ution of
the holotype 0.11,0.15, 0.21,0.34, 0.51,0.77,
and 1.06 mm , and form ratios 1.00, 1.60, 2.41,
2.46, 2.45, 2.47, and 2.43, respectively.
Proloculus small an d spherical. Its outside diameter varies from 0.100 to 0.150 mm ,averaging 0.119 mm for 22 specimens.
Spirotheca thin a nd consists of a tectumand keriotheca. Thickness of spirotheca of
the first to the seve nth volution of theholotyp e 0.0 10,0.0 15 , 0.025 , 0.040 , 0.080,
0.090, and 0.110 mm , respectively.
Septa thin and regularly fluted throughout
length of shell except for central part. Septal
counts of the first to the sixth volution ofsagittal section of one of the paratypes (Figure6-4) 7, II, 11 , 14, IS , and 19, respectively.
Chomata small and observed in the first to
penultimate volution. Tunnel path almost
straight. Tunnel angles of the first to thesixth volution of axial section of one paratype(Figure 6-7) 29, 41 , 45, 51,59, and 45 degrees,
respectively. No axial fillings present.
Remarks. - Pseudofusulina siamensis, sp .
nov . is a species of this genus having a characteristic small shell , whose size is more or lesssimil ar to that of the genus Triticites . The
former , ho wever, has more fluted septa thanFigure 7. . 1- 4. Pseudoreichelina darvasica Leve n, 1,2: axia l section s, IGUT-KU0336, KU0337, 3:
tan.gentlal sectIon, IGUT-KU033 8, 4: parallel section , IGUT-KU0339. 5- 13. Pseudoendothyra ? constrz~ta Igo, Ueno an d Sash Ida, sp. nov. , 5 : ax ial section of the hol otype, IGUT-KU0340, 6- 9, 11- 13: axia l
sectIOn s of paratypes, IGUT-KU0341 , KU0342, KU0343, KU0344, KU0346, KU0347, KU0348, 10 :parallel sectIon of paratype, IGUT-KU0345. 14 : Pseudoendothyra? sp., axia l secti on, IGUT-KU0349.
All x 40.

949.

Lower Permian fusulinaceans from NE Thailand

does the latter. It is also allied to the genus
Daixina in some respects but differs from thelatter in having a smaller and more compactly
coiled shell. The stratigraphic position of
the present species and Daixina is different.
Etymology.- The specific name comes from
Siam, the ancient name of Thailand.
Materials.-Axial section of the holotype :IGUT-KU0313. Axial sections of paratypes: IGUT-KU0314, KU0317-KU0319,
KU0325. Sagittal sections of paratypes:
IGUT-KU0315, KU0316, KU0321, KU0322,
KU0327. Slightly oblique axial sections ofparatypes: IGUT-KU0320, KU0323. Tangential sections of paratypes: IG UTKU0324, KU0326. All specimens from Loe36.
Measurements.-See Table 3.

Pseudofusulina sp. A

33

KU0332-KU0334. Sagittal section:
IGUT-KU0335. Tangential section:
IGUT-KU0331. All specimens from Loe-34.

Pseudofusulina sp. D
Figure 6-18

Remarks.- The present species can be distinguished from Pseudofusulina siamensis, sp.
nov. and Pseudofusulina sp. C in having alarger proloculus.
Material.-Slightly oblique axial section:
IGUT-KU0330 from Loe-34.
Family Staffellidae Miklukho-Maklay, 1949
Genus Pseudoreichelina Leven, 1970
Pseudoreichelina darvasica Leven, 1970
Figures 7-1-4

Pseudoreichelina darvasica Leven, 1970b, p. 19-20, pI.
Figures 5-21-22

Remarks.- The present species somewhat
resembles Pseudofusulina krotowi (Schellwien) but differs from the latter by its smaller
shell and less fluted septa.
Materials.-Axial sections: IGUTKU031l, KU0312 from Loe-34.
Pseudofusulina sp. B
Figures 6-16-17

Remarks.-Pseudofusulina sp. B is similar
to P. kraffti (Schellwien) to some extent but
differs from the latter in having a smaller
shell.
Materials.-Axial section: IGUTKU0329. Tangential section: IGUTKU0328. Both from Loe-34.
Pseudofusulina sp. C
Figures 6-19-23

Remarks.-Pseudofusulina sp. C differsfrom P. siamensis, sp. nov. in having a slightly larger proloculus and less fluted septa.
Materials.-Axial sections: IGUT-

I, figs. 6-13; Lipina and Rozanov, 1973, pI. 6,
figs. 8-10 (same as pI. I, figs. 7-8, II of Leven,1970b); Rozovskaya, 1975, pI. 35, fig. 6 (same as
pI. I, fig.8 of Leven, 1970b); Loeblich andTappan, 1988, pI. 254, figs. 17-18 (same as pI. I,
figs. 8,13 of Leven, 1970b); Ueno, 1992, p. 8-10,
figs. 5-1-16.

Descriptive remarks.-Shell consists of 2
parts. Early part of 4 volutions, lenticularand coiled planispirally with shallowly umbilicated axial regions and angular periphery.
Later one uniserial and rectilinear, consistingof 6 to 9 chambers. Total length of shell 1.28
to 1.40 mm. Spirotheca light-brownish colored and replaced by secondary mineralization. Aperture basal in coiled part, but
median in rectilinear one.
Pseudoreichelina darvasica Leven is known
from the upper Yakhtashian (Artinskian)
Safetdaron Suite of southwestern Darvas(Leven, 1970b) and the upper Yakhtashian of
the Akiyoshi Limestone Group, southwest
Japan (Ueno, 1992). The Loei specimens are
identical with both of the Darvas and Akiyoshi ones.
Materials.-Axial sections: IGUTKU0336, KU0337 from Loe-34. Tangential

34

Hisayoshi [go et al.

Figure 8. 1- 4. Staffella? sp. B, 1,2, 4 : ax ial section s, IGUT-KU0350, KU0351 , K U0353, 3 :sagittal section, IGUT-KU0352. 5- 14. Nankinella? loeiensis Igo, Ueno and Sashida, sp. nov., 5 : axia l
section of the holotype, IGUT-KU0354, 6: sagittal secti o n of paratype, IGUT-KU0355 , 7- 14 : ax ial
sections of paratypes, IGUT-KU0356, KU0357 , KU0358 , KU0359, KU03 60, KU03 6 1, KU03 62, KU03 63.
15 : Staffella sp. A, tangential section, fGUT-KU0364. All X 30.

949.

Lower Permian fusulinaceans from NE Thailand

section: IGVT-KV033S from Loe-34. Parallel section: IGVT-KV0339 from Loe-3S.

35

the holotype O.OS, 0.13, 0.19, 0.26, 0.35,0.48,
0.63, and 0.76? mm, and form ratios 0.S8,
0.77,0.79,0.77,0.83, 0.S3, 0.79, and 0.78,
respectively.
Proloculus spherical or somewhat irregularin shape, being 0.050 to O.OSO mm in outside
diameter.
Spirotheca light-brownish colored andrather thin, replaced by secondary mineralization. Fine alveolar structure visible in spirotheca of outer volutions. Thickness of spirotheca of the first to the eighth volution of theholotype 0.010, 0.015,0.020 ?, 0.020, 0.020,
0.030,0.040, and 0.040 mm, respectively.
Septa thin and planar. Septal counts ofthe fourth to the seventh volution of sagittal
section of one paratype (Figure 9-10) 9, 11,
12, and 14, respectively.
Chomata small, asymmetrical, and developed in all volutions. Tunnel path straightand narrow. Tunnel angles of the second to
the seventh volution of axial section of one
paratype (Figure 9-13) 24,21,21,21,23, and28 degrees, respectively. Axial fillings present in some specimens.
Remarks.-Staffella ? transiens originallydescribed by Kochansky-Devide (1965) fromthe PV 1 (Lower Permian) of Croatia, is the

Genus Staffella Ozawa, 1925
Staffella ? ovalis Igo, Veno andSash ida, sp. nov.
Figures 9-9-15

Diagnosis .-Small Permian Staffella having a thickly lenticular or oval shell withbroadly rounded periphery and slightly umbilicated polar regions. Coiling planispiralthroughout growth. Spirotheca with finealveolar structure. Chomata distinct.
Description.-Shell small for Permian representatives of this gen us and thickly
lenticular or oval with broadly rounded
periphery and shallowly depressed polar
regions. Mature specimens having 7 Yz to S
Yz volutions, 1.20 to 1.45 mm in length and I.
33 to 1.75 mm in width. Form ratio varies
from 0.79 to 0.91, averaging 0.S3 for 7 specimens. Holotype specimen of S volutions
1.20 mm in length and 1.53 mm in width,
giving a form ratio of 0.79.
Shell expands uniformly throughout
growth. Axis of coiling straight. Radius
vectors of the first to the eighth volution of

Measurements of Staffella? ovalis Igo, Ueno and Sashida, sp. nov. (in mm)

Table 4.

Radius vector
Reg. no.

FigureLength Width

F.R.

D.P.
1

2

3

4

5

6

7

8

1 IGUT-KU0376

9-12

1.23

1.43

0.86

2 IGUT-KU0377

9-13

1.23

1.50

0.82

3 IGUT-KU0378
4 IGUT-KU0379

9-14

1.25

1.50

0.83

0.09 0.14 0.19 0.27 0.34 0.44 0.58 0.76
0.080? 0.08 0.13 0.19 0.26 0.36 0.49 0.63
0.090 0.10 0.16 0.20 0.28 0.36 0.49 0.60 0.75

9-15

1.20

1.53

0.79

0.050

Form ratio
1

2

Thickness of spirotheca
3

4

5

1 0.78 0.64 0.65 0.64 0.74 0.84 0.90 0.85 0.010 0.Q15

0.020

0.030

0.025

2 0.50 0.54 0.65 0.85 0.81 0.82 0.82
0.015 0.020
3 0.60 0.56 0.71 0.79 0.81 0.80 0.84 0.82 0.015 0.015
4 0.88 0.77 0.79 0.77 0.83 0.83 0.79 0.78 0.010 0.Q15

0.020

0.020
0.020

0.025 0.030 0.D35
0.030? 0.030 0.050 0.060
0.020 0.030 0.040 0.040

34

0.08 0.13 0.18 0.25 0.34 0.47 0.61 0.74

5

6

7

8

1

2

-

0.020? 0.020

6

7

8

0.025 0.040 0.030

36

Hisayoshi [go el al.

Figure 9. 1- 8. Sphaerulina cf. croalica Koch ansky-Devide, 1,2,4- 8: axial sectio ns, IGUTKU0365 , KU0366, KU0368 , KU03 69, KU03 70, KU03 71, KU0372, 3 : ta nge nt ial section , IGUT-KU0367.
9-15. Slaffella? ovalis Igo, Ueno and Sash ida, sp. nov. , 15 : axia l section of the holotype, IGUT-KU0379,
9,12- 14 : ax ial sections of para types, IGUT-KU0373, KU03 76, KU03 77, KU0378, 10, 11: sagittal sectionsof paratypes, IGUT-KU0374, KU03 75. All X 30.

949.

Lower Permian fusulinaceans from NE Thailand

closest to the present new species. The former, however, has a larger shell, more volutions, and slightly smaller form ratio than thelatter.
Staffella? ovalis, sp. nov. somewhat resembles S. yaziensis originally described by
Wang and Sun (1973) from the Yazi Formation of the Qinling (Chin ling) Range, South
China. However, the former has a larger
shell and more volutions than does the latter.
Presence of fine alveolar structure in the
spirotheca of the present form is inconsistent
with the original diagnosis of the genus
Staffella. The present new species is, therefore, tentatively assigned to the genus

Staffella.
Etymology.- This specific name is derived
from the Latin ovalis, meaning oval.
Materials.-Axial section of the holotype:
IGUT-KU0379. Axial sections of paratypes: IGUT-KU0373, KU0376-KU0378.
Sagittal sections of paratypes: IGUTKU0374, KU0375. All specimens from Loe34.
Measurements.-See Table 4.

37

Staffella sp. A
Figure 8-15

Remarks.-Staffella sp. A differs from
Staffella? ovalis, sp. nov. in having a larger
form ratio and absence of fine alveolar structure in the spirotheca.
Material.- Tangential section: IGUTKU0364 from Loe-37.

Staffella ? sp. B
Figures 8- 1-4

Remarks.- This species is characterized byhaving a Staffella-like rounded periphery in
the inner volutions and Nankinella-like angular periphery in the last one, and a replaced spirotheca composed of a tectum andfine alveolar keriotheca. Therefore, it istentatively assigned to the genus Staffella.
Materials.-Axial sections: IGUTKU0350, KU0351, KU0353. Sagittal section: IGUT-KU0352. All specimens fromLoe-36.

Table 5. Measurements of Nankinella? loeiensis Igo, Ueno and Sash ida, sp. nov. (in mm)
Radius vector
Reg. no.

FigureLength Width

F.R.

D.P.
I

2

3

4

5

6

I IGUT-KU0354

8-5

0.90

1.65

0.55

0.060

0.08

0.15

0.26

0.43

0.66

0.87

2 IGUT-KU0356

8-7

0.83

1.58

0.52

0.070

0.08

0.15

0.24

0.40

0.61

0.79

3 IGUT-KU0357

8-8

0.88

1.53

0.57

0.065

0.09

0.16

8-9

0.95
0.80

1.70
1.40

0.56
0.57

0.085
0.070

0.09

0.15
0.12

0.24
0.21

0.35
0.32

0.49

4 IGUT-KU0358
5 IGUT-KU0361

0.19

0.29

0.67
0.61
0.64

8-12

Form ratio

Thickness of spirotheca

I

2

3

4

5

6

0.50

0.50

0.50

0.50

0.49

0.46

2 0.75

0.67

0.56

0.56

0.54

0.53

3 0.44
4 0.56
5 0.50

0.50

0.45
0.55

0.53
0.58
0.57

0.58
0.51

0.57

0.53

0.62
0.47

0.55

I

0.53
0.50

0.50

0.06

0.48
0.45

0.48

I

7

2

3

4

5

0.015 0.020 0.025 0.035
0.010 0.020 0.020 0.035 0.035
0.040
0.010 0.QI5 0.025
0.010 0.010 0.020 0.QI5
0.030
0.035
0.040
0.005
-

6

7

0.020?
0.040
0.050
0.030
0.040

0.025
0.040

7

0.86
0.81

Hisayoshi Jgo et al.

38

Genus Nankinella Lee, 1934
Nankirtella ? loeiensis Igo, Ueno
and Sashida, sp. nov.
Figures 8-5-14

Diagnosis.-Permian Nankinella having asmall lenticular shell with angular peripheryand almost straight lateral slopes. Axis ofcoiling straight throughout growth. Spirotheca with fine alveolar structure. Chomata
distinct.
Description.-Shell small for Permian representatives of the genus and lenticular withangular periphery and almost straight lateralslopes. Axial regions slightly depressed.
Mature specimens of 6 to 7, rarely 7 Yz volutions 0.73 to 1.05 mm in length and 1048 to
1.75 mm in width. Form ratio ranges from0049 to 0.63, averaging 0.55 for II specimens.
Holotype specimen of 6 volutions 0.90 mm inlength and 1.65 mm in width with a form
ratio of 0.55.
Shell involute throughout except for innerfew volutions and expands gradually. Axisof coiling straight throughout growth.
Radius vectors of the first to the sixth volution of the holotype 0.08, 0.15, 0.26, 0043, 0.66,
and 0.87 mm, and form ratios 0.50, 0.50, 0.50,
0.50,0049, and 0046, respectively.
Proloculus small and spherical. Its outside diameter varies from 0.060 to 0.085 mm,
averaging 0.067 mm for 9 specimens.
Spiro theca light brown and replaced bysecondary mineralization. Fine alveolar
structure discernible in spirotheca. Thickness of spiro theca of the second to the sixth
volution of the holotype 0.015,0.020,0.025,
0.035, and 0.020 ? mm, respectively.
Septa not fluted and slightly bending anteriorly. Septal counts of the first to the sixth
volution of illustrated sagittal section ofparatype (Figure 8-6) 9?, 11, 13, 12, 16, and19, respectively.
Small but rather prominent chomata present in all volutions. Tunnel path narrow
and almost straight. Tunnel angles of thesecond to the seventh volution of axial sec-tion of one paratype (Figure 8-8) 12, 12, 15,
15, 12, and 14 degrees, respectively.
Remarks.-Nankinella? loeiensis, sp. nov.
somewhat resembles N. quasihunanensis described by Sheng (1963) from the Wujiaping(Wuchaping) Limestone of the Guizhou(Kueichow) district. The former, however,can be distinguished from the latter in having
a smaller shell. Moreover, the present new
species occupies a much lower stratigraphic
position than that of Sheng's species.
The present new species also differs from
Nan kin ella inflata (Colani) in having a smaller shell.
In this study, loeiensis is questionably assigned to the genus Nan kin ella, because it hasa fine alveolar structure in the spirotheca,
which is inconsistent with the original diagnosis of the genus.
Etymology.- This species is named for the
local geographical name Loei, from where thespecimens were collected.
Materials.-Axial section of the holotype:
IGUT-KU0354. Axial sections of paratypes: IGUT-KU0356-KU0363. Sagittal
section of paratype: IGUT-KU0355. All
specimens from Loe-36.
Measurements.-See Table 5.
Genus Sphaerulina Lee, 1933
Sphaerulina cf. croatica
Kochansky- Devide, 1965
Figures 9-1-8Compare. Sphaerulina croatica Kochansky-Devide, 1965, p.
123-125, 143-144, pI. 5, fig. 9, pI. 6, figs. 1-14, pI.
7, figs. 1-8, pI. 8, fig. 1 ; Rui, 1979, pI. 1, figs. 2021.

Description.-Shell small and almost spherical with slightly umbilicated axial regions
and broadly expanded periphery. Mature
specimens having 5 to 6 volutions, 1.03 to 1.50
mm in length and 1.13 to lAO mm in width.
Form ratio ranges from 0.91 to 1.11, averaging 1.02 for 7 specimens.
Axis of coiling straight throughout growth.

949.

Lower Permian fusulinaceans from NE Thailand

Outer volutions expanded rather rapidly andbecome loose compared with inner ones.
Radius vectors of the first to the sixth volution of typical axial section (Figure 9-8) 0.08,
0.14,0.21,0.36,0.54, and 0.72 mm, and form
ratios 0.88,0.86, 1.05, 0.97, 1.06, and 1.09,
respectively.
Prolocul us small and almost spherical,
being 0.060 to 0.080 mm in outside diameter.
Spirotheca light brown, rather thick andrecrystallized by secondary mineralization.
Fine alveolar structure discernible in spirotheca. Thickness of spirotheca of the first to
the sixth volution of the above-mentioned
specimen 0.010,0.020,0.025,0.070,0.090, and0.070 mm, respectively.
Septa rather thick, plane, and slightly bending anteriorly. Low and small chomata observed in outer volutions.
Remarks.-Sphaerulina croatica was originally described by Kochansky-Devide (1965)

Table 6.

39

from the PV 1 and Pds (Lower and Middle
Permian) of Croatia. The Loei specimens
have a slightly smaller shell and fewer volutions than the latter.
This species can be easily distinguishedfrom Sphaerulina crassispira Lee, the typespecies of the genus, in having a more loosely
coiled shell.
Materials.-Axial sections: IGUTKU0365, KU0366, KU0368-KU0370,
KU0372 from Loe-34; KU0371 from Loe-36.
Tangential section: IGUT-KU0367 fromLoe-34.
Genus Pseudoendothyra Mikhailov, 1939
Pseudoendothyra? constricta Igo,
Ueno and Sashida, sp. nov.
Figures 7-5-13

Diagnosis-Small Permian Pseudoendothyra having a lenticular shell with bluntly

Measurements of Pseudoendothyra ? constricta Igo, Ueno and Sash ida, sp. nov. (in mm)

Reg. no.

FigureRadius vector
Length

Width

F.R.

D.P.
I

2

3

4

5

I

IGUT-KU0340

7-5

0.36

1.03

0.35

0.060

0.07

0.13

0.19

0.35

0.55

2

IGUT-KU0341

7-6

0.34

0.90

0.38

0.060

0.06

0.12

0.18

0.32

0.49

0.30
0.27

0.97

0.31

0.14

0.24

0.36

0.050
0.050

0.08
0.05

0.045

0.16
0.23

0.46

0.41

0.10
0.14

0.40
0.28

0.34

0.76
0.74

0.47
0.32

0.055
0.060

0.11
0.12

0.16
0.19

3

IGUT-KU0342

7-7

4

IGUT-KU0343

5

IGUT-KU0344

7-8
7-9

6
7

IGUT-KU0347

7-12

0.41

0.87

IGUT-KU0348

7-13

0.37

1.16

Form ratio

0.07
0.06
0.07

Thickness of spirotheca

I

2

3

4

5

I

2

3

4

5

I

0.86

0.69

0.60

0.56

0.46

0.010

0.Ql5

0.025

0.025

2
3
4

0.67
0.75

0.67
0.64

0.48
0.49
0.55
0.46

0.44

0.010
0.015
0.005
0.010

0.015
0.020
0.020

0.025

0.47

0.010
0.010
0.010

0.67

0.50
0.55

0.58
0.56
0.59
0.54

0.025
0.010
0.030
0.020

0.53

0.50

0.020
0.025

0.57

0.50

0.50

0.44

0.42
0.39

0.020
0.Ql5
0.015

0.020

5
6
7

0.60
0.57

0.60

0.010
0.010

0.020

0.015
0.030
0.020

0.010
0.020

0.38
0.27
0.35

0.49
0.52

40

Hisayoshi [go et al.

pointed periphery and shallow umbilicus.
Outermost volution commonly evolute.
Spirotheca with fine alveolar structure.
Chomata present.
Description.-Shell small for Permian representatives of the genus and lenticular withbluntly pointed periphery and slightly umbilicated axial regions. Mature specimens having 4 Yz to 5 Yz volutions, 0.27 to 0.41 mm inlength and 0.76 to 1.16 mm in width. Formratio ranges from 0.32 to 0.47, averaging 0.37
for 7 specimens. The holotype of 5 volutions
0.36 mm in length and 1.03 mm in width witha form ratio of 0.35.
Axis of coiling almost straight. Shell
involute except for the last volution andexpands uniformly. Radius vectors of thefirst to the fifth volution of the holotype(Figure 7-5) 0.07, 0.l3, 0.19, 0.35, and 0.55mm, and form ratios 0.S6, 0.69, 0.60, 0.56, and0.46, respectively.
Proloculus small and spherical, being 0.045
to 0.060 mm in outside diameter.
Spirotheca light brown and slightly recrystallized. Fine alveolar structure observed inspirotheca. Thickness of spiro theca of thefirst to the fifth volution of the holotype
0.010, 0.015, 0.025, 0.025, and 0.025 mm,
respectively.
Septa unfluted. Chomata small and asymmetrical, developed in all volutions except for
the first one in some specimens. Tunnel pathstraight. Tunnel angles of the first to the fifth
volution of axial section of one paratype(Figure7-S) 17, 17, 15, 17, and 25 degrees,
respectively.
Remarks.-Fine alveolar structure is observed in the spiro theca of the present new
species. This seems inconsistent with theoriginal diagnosis of the genus Pseudoendothyra. Accordingly, the present new species istentatively assigned to Pseudoendothyra.
Pseudoendothyra? constricta, sp. nov.
somewhat resembles P. obiniouensis which
was originally described by Leven (l970b)from the Yakhtashian of Darvas. However,the former differs from the latter in having aslightly larger shell with more volutions,
smaller form ratio and more rounded periphery, and fine alveolar structure in the spirotheca.
Etymology. - The specific name is derived
from the Latin constrictus, meaning constricted.
Materials.-Axial section of the holotype :IGUT-KU0340. Axial sections of paratypes: IGUT-KU034l-KU0344, KU0346-KU034S. Parallel section of paratype:
IGUT-KU0345. All specimens from Loe-35.
Measurements.-See Table 6.

Pseudoendothyra ? sp.
Figure 7-14

Remarks.- Fine alveolar structure is discernible in the spirotheca of the present
species. This species is closely allied to
Pseudoendothyra? constricta, sp. nov., butdiffers from the latter in having a slightly
longer axis of coiling and larger form ratio.
Material.-Axial section: IGUT-KU0349
from Loe-35.
References cited
Bunapas, S., 1983: Paleozoic succession in Thailand.
Proceedings of the Workshop on Stratigraphic
Correlation of Thailand and Malaysia, vol. 1,
Technical Paper, p. 29-76.
Charoenpravat, A., Wongwanich, T., Tantiwanit, W.
and Theetiparivatra, U., 1976: Geological Map
of Changwat Loei, Sheet NE47-12, ser. 1 : 250,000. Geol. Surv. Div., Dept. Min. Res. Thailand.
Chang, L.H. (Zhang, L.X.), 1963: Upper Carboniferous fusulinids from Keping and its adjacentregions, southwestern Sinkiang (I). Acta
Palaeont. Sinica, vol. 11, no. 1, p. 36-70, pIs. 1-3.
(in Chinese and Russian)Chen, S (Chen, X), 1934a: Fusulinidae of South
China, Part I. Palaeont. Sinica, ser. B, vol. 4,
fasc.2, p. 1-185, pIs. 1-16.
- - , 1934b: Fusulinidae of the Huanglung and
Maping limestones, Kwangsi. Mem. Nat. Res.
Inst. Geol., no. 14, p. 33-54, pIs. 6-8.
- - and Wang, 1.H., 1983: The fusulinids of theMaping Limestone of the Upper Carboniferous
from Yishan, Guangxi. Palaeont. Sinica, N.S.
H, no. 19, p. 1-139, pis. 1-25. (in Chinese with

949.

Lower Permian fusulinaceans from NE Thailand

English abstract)
Da, Y.T. and Sun, Q.L., 1983: Fusulinida. In,
Regional Geology Surveying Team of Xinjiang
et ai, chiefly eds., Paleontological Atlas ofNorthwest China, Xinjiang Weiwuer Aut.
Region Vol., no. 2 (Late Paleozoic), p.7-113,
pis. 1-30. Geol. Publ. House, Beijing. (inChinese)Ding, P.Z., lin, T.G. and Sun, Y.F., 1987: An excursion guide to Permian geology of Xikou area,
Zhen'an County, Shaanxi. Bull. Xi'an Inst.
Geol. Mineral Resour., Chinese A cad. Geol. Sci.,
no. 15, p. 113-138, pis. 1-5. (in Chinese withEnglish abstract)- - , - - and - - , 1989: Carboniferous-Permianboundary of Xikou area of Zhen'an, south
Shaanxi, east Qinling Range. lIe Congr. Internat. Strat. et Giol. Carbon., Beijing, 1987,
Compte Rendu, Tome 2, p. 199-206, pis. 1-4.
Ding, Y.J., Xia, G.Y., Li, L., Yu, X.G., Zhao, S.Y.
and Zhao, Z., 1991: The Carboniferous-Permianboundary and faunas from Xikou area, Zhen'an,
Shaanxi in eastern Qinling Range. Bull. Tianjin
Inst. Geol. Mineral Resour., Chinese A cad. Geol.
Sci., no. 24, p. 1-202, pis. 1-31. (in Chinese withEnglish abstract and description of new genera
and family)
Dunbar, C.O. and Skinner, l.W., 1937: The Geologyof Texas, Volume III, Part 2. Permian
Fusulinidae of Texas. Univ. Texas Bull., no.
3701, p. 517-825, pis. 42-81.
Fontaine, H., Poumot, C. and Songsirikul, B., 1981 :New Upper Paleozoic formations in Northeast
Thailand in Devonian and Lower Carboniferous.
CCOP Newsletter, vol. 8, no. 4, p. 1-7.
Huang, Z.X. and Zeng, X.L., 1984: The early Early
Permian (Longlinian Stage) fusulinid fauna from
Longlin, Guangxi. Earth Sci.-Jour., Wuhan
Coll. Geol., vol. 26, no. 3, p. 11-24, pis. 3-4. (inChinese with English abstract and description of
new genera and subgenus)
Igo, H., 1972: Fusulinacean fossils from Thailand.
Part 6, Fusulinacean fossils from North
Thailand. Geol. Palaeont. Southeast Asia, vol.
10, p. 63-116, pis. 9-19. Univ. Tokyo Press.
Ingavat, R. and Douglass, R.C., 1981 : Fusuline fossils from Thailand, Part 14, The fusulinid genus
Monodiexodina from Northwest Thailand.
Ibid., vol. 22, p. 23-34, pis. 6-10.
Kahler, F., 1983: Fusuliniden aus Karbon und Perm
der Karnischen Alpen und der Karawanken.
Sonderhefte der Carinthia II, vol. 41, p. 1-107,
pis. I-II.
- - , 1985: Oberkarbon und Unterperm der Karnischen Alpen, Ihre Biostratigraphie mit Hilfe derFusuliniden. Ibid., vol. 42, p. 1-93, pis. I-II.
- - and Kahler, G., 1980: Fusuliniden aus den

41

Kalken der Trogkofel-Schichten der Karnischen
Alpen. In, Fliigel, E. ed., Die Trogkofel-Stufeim Unterperm der Karnischen Alpen, Ibid., vol.
36, p. 183-254, pis. 1-9.
Kalmykova, M.A., 1967: Permskie fuzulinidy Darvaza. Trudy Vses. Nauchno-Issled. Geol. Inst.
(VSEGEI). nov. ser., vol. 116, Biostratigraficheskiy sbornik, no. 2, p. 116-287, pis. 1-32.
(in Russian)
Kobayashi, F., 1977: Some considerations on theancestor of the family Verbeekinidae(Fusulinacea). Trans. Proc. Palaeont. Soc.
Japan, N.S., no. 105, p. 1-16, pis. 1-2.
Kochansky-Devide, V., 1964: Die Mikrofossilien desjugoslawischen Perms. Paliiont. Zeitschr., Bd.
38,3/4, p. 180-188, pI. 18-19.
- - , 1965: Karbonske i permske fuzulinidne foraminifere Velebita i Like.-Srednji i gornji perm.
Acta geol. Jugosl. akad., vol. 5, p. 101-150, pis.
1-13.
- - , 1970: Permski mikrofosili zahodnih Karavank.
Geologija-Razprave in Porocila, vol. 13, p. 175256, pis. 1-26.
Leven, E.Ya., 1967: Stratigrafiya i fuzulinidy permskikh otlozheniy Pamira. Akad. Nauk SSSR,
Trudy Geol. Inst., vol. 167, p. 1-224, pis. 1-39.
(in Russian)- - , 1970a: 0 proiskhozhdenii vysshikh fuzulinid.
Paleont. Zh urn ai, 1970, no. 3, p. 18-25, pI. I. (inRussian)- - , I 970b : Novyi rod permskikh aberrantnykh
fuzulinidy. Ibid.• 1970, no. 4, p. 16-20, pI. I. (inRussian), 1980: Ob'yasniiel'naya zapiska k stratigraficheskoy shkale permskikh otlozheniy oblasti
Tetis. 5Ip., Minist. Geol. SSR. Vses. Ordena
Lenina Nauchno-Issled. Geol. Inst. (VSEGEI),
Leningrad. (in Russian)- - and Scherbovich, S.F., 1978: Fuzulinidy iStratigrafiya asselyskogo yarusa Darvaza. 162
p., 21 pis. Izdatel. (Nauka), Moskva. (in Russian)Lin, l.X., Li. l.X., Chen, G.X., Zhou, Z.R. andZhang, B.F., 1977: Fusulinida. In, Hubei Inst.
Geol. Sci. et al. eds., Paleontological Atlas ofCentral South China, no. 2 (Late Paleozoic), p.
4-96, pis. I-30. Geol. Publ. House, Beijing. (inChinese)
Lipina, O.A. and Rozanov, A.Y., 1973: 0 gomologicheskoi izmenchivosti foraminifer i arkheotsiat.
Akad. Nauk SSSR, Sibirskoe Otdel., Trudy Inst.
Geol. i Geofiz., vol. 49, p. 13-31, pis. 3-6. (inRussian)Liu, Z.A., Xiao, X.M. and Dong, W.L., 1978: Protozoa. In, Working Group of Stratigraphy andPaleontology of Guizhou ed., PaleontologicalAtlas of Southwest China, Guizhou Vol., no. 2

42

Hisayoshi [go et al.

(Carboniferous to Quaternary), p. 12-98, pis. 124. Geol. Publ. House, Beijing. (in Chinese)
Loeblich, A.R., Jr. and Tappan, H., 1988: Foraminiferal genera and their classification. 970
p., 847 pis. Van Nostrand Reinhold Co., NewYork.
Luo, J.D., Hong, Z.Y., Xu, H.Q., Lin, X.S., Yi, W.J.,
Li, Y.L. and Wu, F.B., 1989: On the
Carboniferous-Permian boundary in Fujian.
lIe Congr. Internat. Strat. et Geol. Carbon.,Beijing, 1987, Compte Rendu, Tome 2, p.216228, pis. 1-4.
Nie, Z.T. and Song, Z.M., 1983: Fusulinids of Lower
Permian Qudi Formation from Rutog of Xizang(Tibet), China. Earth Sci.-Jour., Wuhan Coll.
Geol., vol. 19, no. 1, p.29-42, pis. 1-2. (inChinese with English abstract)
Pitakpaivan, K., 1965: Fusulinacean fossils fromThailand, Part I. Fusulines of the Rat Buri
Limestone of Thailand. Mem. Fac. Sci., Kyushu Univ., ser. D, Geol., vol. 17, no. I, p. 1-69,
pis. 1-6.
Ramovs, A. and Kochansky-Devide, V., 1965:
Razvoj mlajsega Paleozoika v okolici Ortneka na
Dolenjskem. Siovenska Akad. Znanosti Umetnosti (Ljubljana), CI. 4, Historia Nat. Medicina,
Razprave, Dissert. 8, p. 323-416, pis. 1-18.
- - and - - , 1979: Karbonske in permske plasti v
severnih Julijskih Alpah. Geologija-Razpravein Porocila, vol. 22, no. I, p. 21-54, pis. 1-6.
Rauser-Chernoussova, D.M., 1988: 0 polozhenii
roda Pamirina v sisteme fuzulinid. Paleont.
Zhurnal, 1988, no. 4, p. 13-19, pis. 1-2. (in Russian)Rozovskaya, S.E., 1958: Fuzulinidy i biostratigraficheskoe raschlenenie verkhnekamennougol'
nykh otolozheniy Samarskoy Luki. Akad. Nauk
SSSR, Trudy Geol. Inst" vol. 13, p. 57-120, pis.
1-15, (in Russian)- - , 1975: Sostav, sistema i filogeniya otryada
fuzulinida. Akad. Nauk SSSR, Trudy Paleont.
Inst., vol. 149, p. 1-227, pis. 1-35. (in Russian)Rui, L., 1979: Upper Permian fusulinids from western Guizhou. Acta Palaeont. Sinica, vol. 18,
no. 3, p.271-297, pis. 1-4. (in Chinese withEnglish abstract)
Sakagami, S. and Iwai, J., 1974: Fusulinacean fossils
from Thailand, Part 8. Permian fusulinaceansfrom the Pha Duk Chik Limestone and in thelimestone conglomerate in its environs, North
Thailand. Geol. Palaeont. Southeast Asia, vol.
14, p. 49-81, pis. 4-9, Univ. Tokyo Press.
Schellwien, E., 1909: Monographie der Fusulinen,
Teil II, Die asiatischen Fusulinen. A. Die
Fusulinen von Darwas. Palaeontogr., Bd. 56, p.
137-176, pis. 13-16.
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.
- - , 1966: Fusulinids of China. 177 p., 27 pis., Sci.
Publ. House, Beijing. (in Chinese)- - and Sun, D.D., 1975: Fusulinids of Qinghai.
97 p., 14 pis., Ibid. (in Chinese)- - , Zhang, L.X. and Wang, J.H., 1988: Fusulinida.
240 p., Ibid. (in Chinese)
Staff, H. von, 1909: Beitrage zur Kenntnis derFusuliniden. Neues Jahrb. Min., Geol.,
Paliiont., Beil., Bd. 27, p. 461-508, pis. 7-8.
Sun, Q.L. and Zhang, L.X., 1988: Early Permian
fusulinids from the Kongkashan pass of Xinjiang. Acta Micropalaeont. Sinica, vol. 5, no. 4,
p. 367-378, pis. 1-4. (in Chinese with English
abstract)Sun, X.F., Gao, C.X. and Zhang, X.L., 1983:
Fusulinida. In, Xi'an Inst. Geol. Mineral
Resour. ed., Paleontological Atlas of Northwest
China, Shaanxi, Gansu and Ningxia Vol., no. 2( Upper Paleozoic), p. 3-40, pis. 1- 10. Geol.
Publ. House, Beijing. (in Chinese)
Toriyama, R., 1975: Fusuline fossils from Thailand,Part 9. Permian fusulines from the Rat Buri
Limestone in the Khao Phlong Phrab area, Sara
Buri, central Thailand. Mem, Fac. Sci., Kyushu
Univ., ser. D, Geol., vol. 23, no. I, p. 1-116, pis.
1-21.
- - , 1984: Summary of the fusuline faunas inThailand and Malaysia. Geol. Palaeont. Southeast Asia, vol. 25, p.137-146. Univ. Tokyo
Press.
- - , Hamada, T., Igo, H., Ingavat, R., Kanmera, K.,
Kobayashi, T., Koike, T., Ozawa, T., Pitakpaivan, K., Piyasin, S., Sakagami, S., Yanagida,J., and Yin, E.H., 1975: The Carboniferous andPermian Systems in Thailand and Malaysia.
Ibid., vol. 15, p. 39-76.
- - and Kanmera, K., 1979: Fusuline fossils fromThailand, Part 12. Permian fusulines from the
Ratburi Limestone in the Khao Khao area,
Saraburi, central Thailand. Ibid., vol. 20, p. 2393, pis. 4-14.
Ueno, K., 1991 a: Pamirina (Permian Fusulinacea)
from the Akiyoshi Limestone Group, southwest
Japan. Trans. Proc. Palaeont. Soc. Japan, N.S.,
no. 161, p. 739-750.
- - , 1991b: Early evolution of the families Verbeekinidae and Neoschwagerinidae (Permian
Fusulinacea) in the Akiyoshi Limestone Group,southwest Japan. Ibid., no. 164, p.973-1002.
- - , 1992: Permian aberrant fusulinaceans from the
Akiyoshi Limestone Group, southwest Japan.
Sci, Repts., Inst. Geosci., Univ. Tsukuba, Sec. B,
vol. 13, p. 1-13.
Wang, K.L. and Sun, X.F., 1973: Carboniferous andPermian foraminifera of the Chinling Range and949.

Lower Permian fusulinaceans from NE Thailand

its geologic significance. Acta Geol. Sinica,
1973, no. 2, p. 137-178, pIs. 1-7. (in Chinese withEnglish abstract and description of new species)
Wang, Y.H., 1987: Late Carboniferous fusulinids in
middle and lower Yangtze valley. Bull. Nanjing Inst. Geol. Mineral Resour., Chinese A cad.
Geol. Sci., vol. 8, no. 2, p. 85-91, pis. 1-2. (inChinese with English abstract)- - , Wang, L.L., Wang. J.H., Zhu, Z.G., Lin, G.W.,
Zhang, L.X. and Quian, Q., 1982: Fusulinida.
In, Nanjing Inst. Geol. Mineral Resour. chiefly
ed., Paleontological Atlas of East China, no. 2(Late Paleozoic) , p.5-108, pis. I-3~. Geol.
Publ. House, Beijing. (in Chinese)
Watanabe, K., 1991 : Fusuline biostratigraphy of theUpper Carboniferous and Lower Permian ofJapan, with special reference to the
Carboniferous-Permian boundary. Palaeont.
Soc. Japan, Spec. Papers, no. 32, p. 1-150.
Xia, G.Y., Li, J.X., Wang, Y.H. and Dong, W.L.,
1986: The fusulinid zones and the boundary of
Carboniferous-Permian in Longlin region, Guangxi. Bull. Yichang Inst. Geol. Mineral Resour.,

Chinese Acad. Geol. Sci., no. II, p. 67-104, pIs.
I-II. (in Chinese with English abstract)- - and Zhang, Z.e., 1985: Protozoa. In, Tianjin
Inst. Geol. Mineral Resour. ed., PaleontologicalAtlas of North China, no. 1 (Paleozoic vol.), p.
32-150, pis. 1-35. Geol. Publ. House, Beijing.
(in Chinese)
Yanagida, J., 1967: Early Permian brachiopods from
central Thailand. Geol. Palaeont. Southeast
Asia, vol. 3, p.46-97, pis. 11-23. Univ. Tokyo
Press.
- - , 1974: Middle Carboniferous brachiopods from
Loei, North Thailand. Ibid., vol. 14, p.7-23,
pis. 2-3.
- - , 1975: Upper Carboniferous brachiopods from
Wang Saphaung, North Thailand. Ibid., vol.
16, p. 1-31, pis. 1-3.
Zhou, T.M., Sheng. J.Z. and Wang, Y.J., 1987:
Carboniferous-Permian boundary beds andfusulinid zones at Xiaodushan, eastern Yunnan.
Acta Micropalaeont. Sinica, vol. 4, no. 2, p. 123160, pis. 1-6.

m,Guangxi]A j1§,
Akiyoshi fj(jl:f, Beiyangshan ~t:F lll, Chuanshan Wd lll, Guangnan]A
Guizhou (Kueichow) JtH/, Hanagiri itffOJ, Jiangsu (Kiangsu) lIi*, Kwanto ~J:j~:, Longlin!lift, Qinghai WW:, Qinling (Chinling) ~fii!t, Shaanxi ~j1§. Wujiaping (Wuchaping) lJ'l:
Xiaodushan rN.91111. Yazi tltr. Yunnan ~m. Zhen'an ~*.

*ff.

~1~tJlnmDI1 ~1:: 7*'l'ii:T$A.)lr.b.**1i~.'il.: !J 11E~t*tffi p.:r..1 iJf<0) l:: 7tHuJ'
m.K.w~~~~~.~~~.~~tK.L. ~m~~~*O)?~~~-~.O)-tffit~ Gn ~o 4-@J,: O)~~~tJ, G .: n i "C!J- II//~.A ~<=plEmtffio) Ttffi~ Iv A *0) Yakhta ..
shian (Artinskian) ~tffitJ, G¥IH!i' ~ n"n, ~~~.'R{r.~ll'H.::.(ijji!;J:t ~ ~ O)Z'~J.iJj L 1-:0': 0)iF,ft\;bj~~~~.'R {r.~MI1.!J 11E "'(·I1.1J].l0-C -t- O):ff:tEtJ:lit~;g ~ n t..:o it..: 4-@J 0) Je~~.::.
J: -:> -C~~*~.::.~Gn~ t~Gn -Cl't..:? ~~7·-~. t ~/vA*:T A,,;t< '7 ~.IH~3(~ {H'::';5 ~ 'iiJ~gtl: tJ:591 < tt. -:> t..:o ~ t..: ~.::.m:PI'l L t..:~l1 11. Darvasites ingavati, Pseudofusulina siamensis, Staffella ? ovalis, Nankinella ? loeiensis, Pseudoendothyra ? constricta 0)511~;5~043

R.~.·~~.~·mffi.~

Trans. Proc. Palaeont. Soc. Japan, N .S., No. 169, pp. 44- 72, 23 Figs., April 30, 1993

950.

SKELETAL STRUCTURES OF LOWER AND MIDDLEMIOCENE LOPHOPHAENIDS (RADIOLARIA)
FROM CENTRAL JAPAN*
KAZUHIRO SUGIYAMA
Department of Earth and Planetary Sciences, School of Science,
Nagoya University, Nagoya, 464-01

Abstract. Nine species of five lophophaenid genera are described from the Lower
Miocene Toyohama Formation, Morozaki Group, Aichi Prefecture and the lowest Middle
Miocene Oidawara Formation, Mizunami Group, Gifu Prefecture, central Japan. A special
attention is given to describing and illustrating the details of their skeletal structures which
possess one or two horizontal rings, since it is believed that these details are important inestablishing the taxonomy and clarifying the phylogenetic relationships. The examination
discriminates seven types of skeletal structure. Four species are newly described, and two
new genera, Steganocubus and Cryptogyrus, are also erected herein.
Key words. Radiolaria, lophophaenids, skeletal structures, Lower and Middle
Miocene, central Japan

should be necessary.
However, it seems that the taxonomic position of H. vema is essentially problematic.
The problems can be summarized in the following two points: One concerns whether
the genus Helotholus Jyjrgensen, 1905, is ajunior synonym of the genus Ceratocyrtis
Biitschli, 1882, or not, and the other iswhether H. vema really belongs to this genus
or not. Concerning the former problem,
Petrushevskaya (1971 b) was the first who
formally synonymized these two genera, andher action seems to be reasonable. Thus the
problem is out of interest for this paper.
However, the latter problem remains unsettled although over the past two decades many
papers have discussed the taxonomy of H.
vema, although Weaver (1983), Chen (1975),
Lazarus (1990), and others kept the original
generic assignment for this problematic
species. On the contrary, Petrushevskaya(197lb), Dumitrica (1973), Sanfilippo (1988),
and others referred H. vema to the genus

Introduction

I recently reported the occurrence of wellpreserved radiolarians in the Lower Miocene
Toyohama Formation, Morozaki Group,
Aichi Prefecture (Sugiyama, 1992), and illustrated one characteristic species identified as
Helotholus vema Hays, 1965. This species,
having a unique basal ring (Hays, 1965) in theskeleton, was originally described from Antarctic sediments, and is one of the most characteristic species of the Pliocene radiolarianfauna of the Antarctic region (e.g. Hays,
1965; Chen, 1975). Moreover, Kellogg(1975) and Weaver (1983) mentioned that H.
vema was derived from an ancestral elongatedform (H. praevema Weaver, 1983) byincreases of thoracic width during the Late
Miocene to Pliocene interval. My previous
identification is, therefore, stratigraphicallyproblematic and a taxonomic reexamination'Received July 21,1992; revised manuscript accepted
October 19, 1992
44

950.

Lower and Middle Miocene lophophaenids

Pseudocubus Haeckel, 1887, based on thesimilarity of their skeletal structures.
Moreover, one can point out that the taxonomic problem of H. vema concerns not
only Pseudocubus but also other genera.
For example, some species of the genus
Antarctissa Petrushevskaya, 1967, seem tohave a skeletal structure similar to H. vema
and Pseudocubus, although their precise
skeletal structure has not been fully describedyet (e.g., Antarctissa ewingi Chen, 1974, pI. 3,
fig. 6; A. whitei Bj¢rklund, 1976, pI. 13, fig.
13). In addition, a certain external similarity
is present between the genera Antarctissa,
Botryopera and Ceratocyrtis (= Helotholus)
as discussed by Petrushevskaya (1986). Consequently, we cannot determine the genericassignment of H. vema without establishing aconsistent taxonomy for all the genera
mentioned above. There is no need to stressthat perfect understanding of their skeletal
structures is necessary before everything forthat purpose. Our present state of knowledge about their skeletal structures is, however, too poor to consider their precise taxonomy, much less their phylogeny.
During the subsequent investigations onradiolarian assemblages from the Toyohama
Formation and the lowest Middle Miocene
Oidawara Formation, Mizunami Group, Gifu
Prefecture, I have fortunately obtained wellpreserved materials whose speletal structure is
the same or similar to the above mentioned
problematic taxa, and, as a result of the examination, I realized that to stress ring structurein the skeleton is very useful for their taxonomy. In the present study, therefore, I focus
on describing the structures in detail using ascanning electron microscope (SEM) in orderto present fundamental taxonomic information. The results should contribute to ourbetter grasp of taxonomy of all nassellarians
in future.
Geologic setting and age
The Morozaki Group is widely distributed

45in the southern part of the Chita Peninsula,
Aichi Prefecture, and is divided into theHimaga, Toyohama, Yamami and Utsumi
Formations in ascending order (Shibata,1977). It abundantly yields a deep sea fossil
fauna composed chiefly of Echinodermata,
Crustacea, Mollusca and Pisces which characterizes a bathyal and mesopelagic oceanic
environment (Shibata, 1977; Hachiya et al.,1988). The Toyohama Formation, the second lowest one of this group, is composed
mainly of thick, tuffaceous mudstone accompanied by a subordinate amount of sandstone and tuff layers. The tuffaceous mudstone sometimes intercalates with calcareous
nodules which are approximately 10 cm inmajor diameter and usually contain chelipedsof the thalassinidean decapod Callianassa
titaensis Nagao. Radiolarians are very wellpreserved in the nodules, and a preliminary
report by Sugiyama (1992) revealed that they
are relatively rich in cool-temperate indicators. Four rock samples treated in this study
named TH-l, 2, 3 and 4, the first three samples
were already used in Sugiyama (1992), arealso decapod-bearing calcareous nodules embedded in a tuffaceous mudstone of the Toyohama Formation (Figure 1). They wereobtained approximately from the middle partof the Toyohama Formation. More than
150 radiolarian species have been recoveredfrom these samples.
The precise geologic age of the Toyohama
Formation is not known yet. However, I
obtained the following radiolarian species in
my subsequent investigations: one individual of Lychnocanoma elongata (Vinassa deRegny) from TH-2, one individual of Stichocorys sp. aff S. delmontensis (Campbell
and Clark) from TH-4 and four individualsof S. armata (Haeckel) from TH-2,3 and 4.
They indicate that part of the Toyohama
Formation corresponds to an interval fromthe Lower Miocene Stichocorys delmontensis
Zone to the Middle Miocene Dorcadospyrisa/ata Zone of Riedel and Sanfilippo (1978),according to Sakai (1980). My preliminary

K azuhiro Sugiyama

46

TH-4----

Mikawa
Bay

OCEAN

N300+
E130°

A

135°

B
D~

Fig. 2Ise BayN

TH-3
E 1360 57' 18"

N 340 41'48"

o

2km

..1'

I . '_ _ _ _ _ _Figure 1. Map showing localities of the samples TH-l, 2,3 and 4 from the Toyohama Formation,
Morozaki Group.

investigation on the co-existing diatoms hasrevealed that the flora is dominated by Kisseleviella carina Sheshukova-Poretzkaya andlacks Denticulopsis species. The Toyohama
Formation is, therefore, correlative with the
Early Miocene Kisseleviella carina Zone to
Actinocyclus ingens Zone (Koizumi, 1985).
Although further detailed investigations
should be necessary for precise age determination, I tentatively consider that the age of thestudied samples is of late Early Miocene inthis study.
The Lower to Middle Miocene Mizunami
Group distributed in the Mizunami basin,
Gifu Prefecture, is considered as a contemporaneous heterotopic facies of the MorozakiGroup. According to Itoigawa (1974), theMizunami Group is divided into four formations of which the Oidawara Formation,
composed mainly of tuffaceous anddiatomaceous mudstone, disconformably
overlies the other three formations. Itoigawa
et al. (1974) recognized the Nelionella-Phanerolepida molluscan assemblage fromthis formation, and inferred a lower sublittoral depositional environment. Mori (1974)
examined diatoms from this formation, andestimated an inshore to coastal depositional
environment affected by oceanic water.
The studied route covers approximately thewhole section of the Oidawara Formationfrom the lowest disconformity to the uppermost unconformity (Figure 2). The Oidawara Formation in this route is composed of
two lithologic units divided by a sharpboundary: lower unit consisting of
pumiceous sandstone to sandy siltstone, withor without underlying basal conglomerate,and intercalating several tuff layers; andupper unit composed of monotonous
diatomaceous siltstone. The lower unit may
represent a shallower depositional environment than that of the upper unit jUdging fromits coarser lithology. The occurrence ofradiolarians in the lower unit is very rare,
whereas the upper unit yields a large quantityof siliceous microfossils such as diatoms,
silicoflagellates and radiolarians. This study

950.

Lower and Middle Miocene lophophaenids.0.

~
lliWilliN

Pl iocen e

conglomerate
Upper Oidawara F.
(diatomaceous
siltstone)~

Lower Oidawara F.
(siltstone. cgl.. tuff)§

Hazarna F.
(pumiceous tuff)

vv v

47

Figure 2. Route map and geologic column of the Oidawara Formation, Mizunami Group in thestudied area, showing the locality a nd hori zo n of samples 00-17, IS, 21 , 26,27 and 2S.

deals with six rock samples 00-17,18,21,26,
27 and 28 from the upper unit. More than
120 species of radiolarians have been recovered from these samples.
Sugano (1976) already reported the occurrence of 34 species of radiolari ans in the
Oidawara Formation, and correlated it to the
Meliltosphaera magnaporulosa Zone (Nakaseko and Sugano, 1973) which ranges inage from Early to early Middle Miocene. In
this study, I obtained the following index
species throughout the upper unit: Calocyclelta costata (Riedel), C. virginis (Haeckel),
Stichocorys delmontensis (Campbell and
Clark) , Dorcadospyris Jorcipata (Haecke l)
and Ceratocyrtis stoermeri Goll and Bj¢-

rklund . It is, therefore, concluded that thisformation is correlative with the Calocyclelta
costata Zone of Riedel and Sanfilippo (1978)
and the Ceratocyrtis broeggeri Zone of Golland Bj ¢ rklund (1989). The six studied samples are thu s of an Early to Middle Miocene
age, according to Riedel and Sanfilippo(1978) , Sakai (1984) , Goll and Bj¢rklund(1989) and so on.

Descriptive terminology
As mentioned in the introduction, the main
purpose of the present study is to understandthe skeleta l structures of some lophophaenids
from the Lower to Middle Miocene of centralK azuhim Sugiyama

48

Figure 3. C haracteristic index species from the Toyohama and Oidawara Formations. 1 :Lychnocanoma elongala (Vinassa de Regny) , ESN 146431 , TH-2. 2: Stichocorys sp. aff. S. delmontensis(Campbell and Clark), ESN 146432, TH-4. 3: Dorcadospyris jorcipata (Haeckel), ESN 146451 , 00-26.
4: Stichocorys delmontensis (Campbell and Clark), ESN 146452, 00-28. 5 : Calocy cletta costala(Riedel), ESN 146453,00-26. All sca le bars = 50 ,urn.

Japan . They are, however, so complicatedgeometrically that some specia l terms arenecessary to formu late them. In the firstplace, descriptive terms used in this study,
adding traditional ones, are listed and briefly

I"A

explained below. Schematic illustrations arealso shown in Figure 4.
Arch: Skeleton connecting two basic rays.
The expression follows Petrushevskaya( 1968).

I"

D
MB PR
Dorsal view

Figure 4.

Oblique lateral view

Basal view

Schematic illustrations of idealized skeletal structure.

950.

Lower and Middle Miocene lophophaenids

Sagittal plane: A meridional plane passing
through MB.
MB: Median bar.
A: Apical ray (spine).
A': Lateral branch of A from the junction of
PRo It always lies in the sagittal plane.
D: Dorsal ray (spine).
D': Lateral branch of D from the j unction of
MB and A. The angle between D and D' iscommonly 30·-45". Two D', one at each sideof D, may be present in a specimen. Welldeveloped D' is undiscerned from proper
secondary lateral ray (1).
L: Primary lateral ray (spine). The orientation is shown as Lr, meaning right primarylateral ray, in due place. This follows not
Petrushevskaya (1968, fig. 1) but De Wever(1981, fig. 1) and Dumitrica (1991, fig. 1).
L': Long lateral branch of L. The angle
between Land L' is commonly less than 45·.
One L usually has one L' on the dorsal side.
l': Stout, laterally directed bar arising from
PRo It lies between D and L in basal view.
I": Longitudinal bar extending from thejunction of l' and PRo In basal view, two I"
usually make a larger angle than two l' do,and commonly lie in a plane perpendicular tothe sagittal plane.
V: Vertical ray (spine).
V': Stout, laterally directed bar arising from
PRo It is nearly in the sagittal plane.
V": Longitudinal bar extending from PRo
It commonly arises from the same position as
V and V', but rarely has a different position.
Ax: Axial spine.
L-R: Longitudinal bar typically connecting
the middle part of Land PR situating at thejunction ofL and L'. The length is variable,
so it is in some cases hardly recognized. The
position is also more or less unstable, especially when L' is strongly developed. Thenit connects PR not with L but with L'. This
skeleton possibly corresponds to bb' and cc'of Swanberg and Bj¢rklund (1987).
PR: Proximal ring which can be connectedwith A, A', two 1', two I", two L-R, V, V' andV". It is situated on a horizontal plane near49

the stricture between the first and second
segments. According to Petrushevskaya(1968), PR+two L-R are homologous witharches two ap + two pj when V is present,
whereas they correspond to two ap + pp
when V is absent. They are also
homologous with arcs b' x' c' + b'dec' of
Swanberg and Bj¢rklund (1987). Further
comments are given in the next section alongwith comments on DR.
DR: Distal ring which connects the distal
parts of A, two I" and V".
X: Distance between PR and DR.
Y: Distance between DR and the top of the
eucephalis.
Notes on skeletal ring and taxonomy
It is well known that nassellarians have aninternal skeleton, consisting of MB and basic
rays (A, D, L, I, V and Ax) with or withoutarches, and that many radiolarian taxonomists consider the skeletal structure as themost important criterion for classification.
For example, the main purpose of studies by
Riedel (1967, 1971), Petrushevskaya (1971 a,
1971b,1981), Takemura (1986), Nishimura(1990), and others is to establish the natural
system of nassellarians based on their skeletal
structure. Their common conception is thatthe morphology occurring in earlier
ontogenetic stages may be a more conservative for phylogeny than features formed inlatter stages. That is to say, their conception
conforms to that of heterochrony(McNamara, 1986).
Besides the arrangement of the basic rays of
the internal skeleton, Petrushevskaya (1968)
was the first who recognized the significanceof arches for all nassellarian classification.
Her comprehensive studies (Petrushevskaya,
1971a, 1971b, 1981, etc.) appear to haveinfluenced many subsequent investigators.
Among them a special mention must be paid
for the result of Swanberg and Bj¢rklund(1987), who examined the incipient ontogenyof the living cannobotryid Amphimelissa

50

Kazuhiro Sugiyama

setosa (Cleve) in detail, and revealed thatmost arches (corresponding to the term "arcs"
in their paper) occur prior to the developmentof D, I, V and Ax. From this, one can immediately suppose that the arch structure ofnassellarian skeleton is a more conservative
feature than the final pattern of arrangementof the basic rays.
Among the arches, those developed nearlyhorizontally at or near the collar region areeasy to observe even if the shell wall is thickly
developed on specimens, since it normally has
a rather thin or incomplete wall and thearches are easily recognizable as ridges orswells on the wall in most cases. Thosearches look as if to form a "ring" and to havea distinctive character as an independent
skeletal element in basal view. However,except for Nishimura (1990) who used theterm "basal ring" for this structure of allnassellarians, most investigators have paid
attention to it as a characteristic of the skeleton mainly for two taxa: acanthodesmiids
and those nassellarians with the
Pseudocubus-type skeletal structure whose"rings" are not directly attached to the shellwall but connected with it through intervening connector bars. They are tentatively
called Pseudocubus group, which does notcorrespond to any present nomenclatures.
For acanthodesmiids, the relevant authors are
Haeckel (1887) and Goll (1958) who used theterm "basal ring", whereas Haeckel (1887),
Popofsky (1908, 1913), and Dumitrica (1973)
used the term "ring" for the Pseudocubus
group. Moreover, Hays (1965) and Lazarus(1990) used similar terms including the word
"ring", which are "basal horizontal ring" and"internal ring", respectively. They areapparently for the Pseudocubus group.
It is obvious that PR used in this paper isapplicable only to the Pseudocubus group, as
defined in the way explained in the third
section. Of the nine species described in thispaper, it has revealed that seven, excluding
two Ceratocyrtis species, have such PR in
their skeleton (Table 1). Then one canTable 1. Correlation of seven typicalskeletal structures discriminated in the present
study.
~racteristi

A'

I'

V

V' Y{X shellP••udocubus obel'acus

0 0 -

0

0

-

S

-

Sleganocubus subllll.

0 0

0

0

-

0

S

0

Bolryopera (1) 'aploslraca

0 0

0

0 0

-

L

0

Bolryopera (1) white'

0 0

0

0 0

0

L

0

Bofryoper. (1) .p. A

0

0

0
-

0 0

?

L

0

- 0 -

L

-

- 0 -

-

0
0

small value

large value

spedes~hology PR

IIpus

S.

0 0
0 -

Cryplogyrus trachylobus

Ce,.'ocyrlis stoermerl
C. 8p. aU. C. hl.tt/cosuso-presenceDR--

absenceS -L -notice that the skeletal structure of the groupis commonly characterized by possessing
another, horizontal or nearly horizontal connection of arches above PR, which may beparts of eucephalic lattice bars. This cannot
be formulated by Petrushevskaya's (1968)
method, but seems to be a significant skeleton
closely related to PRo Because of the similarity of the general shape to PR and the position in the skeletal structure, I call it DR in
this paper.
The subfamily or family assignment of thePseudocubus group has been unsettled. For
example, Petrushevskaya (1971 b, 1981) assigned Pseudocubus to the subfamily
Plagiacanthinae Hertwig, 1879, emend.
Petrushevskaya, 1971a, whereas Sanfilippo(1988) put it in the family Cannobotryidae
Haeckel, 1881, emend. Riedel, 1967. However, my investigations proved that it has aclose phylogenetic relationship to the subfamily Lophophaeninae Haeckel, 1881,
emend. Petrushevskaya, 1971 b, although thetypical 10phophaenin "ring" is not free fromthe shell wall and is actually not horizontal:
Their arches ap and pj or ap and pp when V
is absent have a certain common part justabove L, in spite of its various lengths. This
is notable because the part is considered to behomologous with L-R of the Pseudocubus
group (Figure 5). Moreover, a lopho-

950.

Lower and Middle Miocene lophophaenids

SI

shevskaya in Petrushevskaya and Kozlova,
1972, and the subfamily Ceratocyrtinae Petrushevskaya, 1981, are obviously junior synonyms of Lophophaenidae, based on theirskeletal structures. The precise subdivision
requires further detailed investigations infuture.
Genus Pseudocubus Haeckel, 1887,
emend. Petrushevskaya, 1971 b-=PR
Figure 5. Schematic illustration showing correspondence between Petrushevskaya's (1968) arches
and PR and L-R of this study.

phaenin skeletal structure is generally characterized by possessing prominent L' andoften also A' and D' (e.g. the genera Lophophaena Ehrenberg, 1847, and Pseudodictyophimus Petrushevskaya, 1971b, Fig. 23-2a3b). These features are undoubtedly in common with the Pseudocubus group, and two
species of the genus Ceratocyrtis Butschli,
1882, described in this paper have this type ofskeletal structure. Consequently, I consider
that the Pseudocubus group is a member ofLophophaeninae and that it would be betterto raise the subfamily to family rank.

Pseudocubus Haeckel, 1887, p. 101O.-Petrushevskaya, 1971b, p. 149.
Obeliscus Popofsky, 1913, p. 279.
Talariscus Loeblich and Tappan, 1961, p. 227.

Type species: P. obeliscus Haeckel, 1887
(designated by Campbell, 1954).
Pseudocubus obeliscus Haeckel
Figures 7-la--3b; 8-la--Ib
Pseudocubus obeliscus Haeckel, 1887, p. 1010, pI. 94,
fig. 11.--Petrushevskaya, 1971 b, p. ISO, figs. 76--1-6.
Plectophora triacantha Popofsky, 1908, p. 262, pI. 29,
fig. I, pI. 30, fig. I
Obeliscus peudocuboides Popofsky, 1913, p.280, pI.
29, figs. 4, S.

Description.- Test pyramidal in shape, having stout three-bladed scaffolds above PR,

Systematic description(b)

All specimens treated in this study aredeposited in the Department of Earth andPlanetary Sciences, School of Science,
Nagoya University (ESN).

~
A

DR'

V"

~~

o

: • Ax'--t;O L-R

MB(d)~V"(0)Class Radiolaria
Order N assellaria

DR

V"

Family Lophophaenidae Haeckel, 1881,
emend. Petrushevskaya, 1971 b
Remarks.- The diagnosis used in thisstudy follows Petrushevskaya ( 1971 b),
although she subdivided this taxon in her
later studies. For example, the family Lampromotridae Haeckel, 1881., emend. Petru-

..

I"

~~
0'
D

"--Lr

:Ax

--L-A

MB

Figure 6. Schematic illustrations of typicalskeletal structure of Pseudocubus obeliscus Haeckei.
(a): Dorsal view, (b): Right lateral view. (c):Basal view. (d): Right lateral view of gerontic form(?).

Kazuhiro Sugiyama

52

Figure 7. la-3b : PseudoclIblls obelisclIs Haeckel. la - lb: ESN 14650 I, TH-3. 2a, 2b: ESN
146502, TH-4. Arrow of 2a points to secondary arch. 3a, 3b: ESN 146503, TH-3. This specimen has
six longitudinal scaffolds, indicated by arrows, owing to the irregul ar bifurcation of primary skeleton,
which may also contribute to the incompletion of DR. Scale bars: l a, 2a and 3a=50,um ; others=20

,urn.

950.

Lower and Middle Miocene lophophaenids

53

Figure 8. la-ld: Pseudocubus obelisclls Haeckel , ESN 146541 , TH-4, la ; ventral view, Ib;
enlargement of oblique basal view, Ic ; enlargement of apical view, I d; enl argement of left lateral view.
Scale bars : I a = 50 ,u m; others = 20 ,u m.

usually lacking shell completely. Longitudinal scaffolds usually four in number, composed of A, two I" and V", tied by both PR
and DR, and tapering beyond DR. The
scaffolds sometimes bifurcate at the middle
part (Figure 7-3a). PR is either circular or
three-bladed in cross section and usually ovalin entirety. The minor diameter always liesin the sagittal plane. DR is three-bladed incross section . It has a circular outline when
one views it from the basal or apical side. A
side of DR is, however, always convex upward in lateral view, and has one verticil of
one or two conical spines on the top. In
adult specimens (?), the scaffolds havespinules above DR, and secondary arches may
be developed from the tips of them (Figure 72a). The secondary arches finally form

another horizontal connection, a side ofwhich is also convex upward in lateral viewand has a conical spine on the top. In suchcases the eucephalic wall is loosely definable
on the upper half of the test and some thickarches are vertically or obliquely developed
outside the eucephalic part (Figures 8-1 a- ld). Basic spicules located at the base of
the test, consisting of MB, A, D two L, V,
with or without Ax. A directed upward, circular in cross section in the proximal part,straight or slightly bent to dorsal side at thejunction of PR, and prolonged into a stout
three-bladed spine beyond it. One or two
apophyses usually present on the proximal
exterior side of A. In most specimens, D and
two L extending either laterally or slightly
downward, circular in cross section, more orK azuhiro Sugiyama

54

less tapering distally, and having several
small apophyses on the basal side. In gerontic specimens (?), however, two L become
longer and three-bladed, and have a verticil ofa few or several spinules radiating in all
directions. Then D' is well developed, but D
remains rodlike. L' always missing. Two I'
stout, commonly conical and proximallybladed, becoming longer and distally pronged
like two L in gerontic specimens (?). Two I"
extend upward from the corresponding points
to two I' . V rodlike, thin, directed obliquely
upward, extending to PR. V of some specimens not extending into V" and, in suchcases, two V" may be present in the specimens. Ax wartlike, sometimes undiscerned
from the basal apophyses on the D and two L.
TwoL-R long or short, sometimes unequal.
A' and V' missing. Each junction of DR
and A, V" and two I" bears one spine on theexterior side.
Remarks.- Variation of the skeletal structure is in rare cases observed with respect tothe intersecting position of V and PR, number of V" and length of L-R. Although

<oj

OR

Figure 9. Schematic ill ustrations of ty picalskeletal structure of Steganocubus subtilis gen. et sp.
nov. (a): Dorsal vIew. (b) : Right lateral view.
(c): Basal view.

Petrushevskaya (1971 b) mentioned that thedevelopment of V is capable of intensely
varying, all the examined specimens have V.
On the other hand , none have A' and V'.
Judging from the descriptions of previous
studies (Haeckel, 1887 ; Popofsky, 1908,
1913 ; Petrushevskaya, 1971b), however, it iscertain that this species is morphologicallyFigure 11. la, Ib: Steganocubus subtilis Sugiyama, gen. et sp. nov., ESN 146507, TH-2, paratype.
Scale bars =50 11m. This specimen has a quite irregular skeletal structure, so that the elements cannot be
named.
~
Figure 10. la-4b : Steganocubus subtilis Sugiyama, gen. et sp. nov. la- Id: ESN 146504, TH-4,
holotype. Arrow of Id points to vestigial D' . 2a, 2b: ESN 146505, TH-4, paratype. 3a, 3b : ESN
146403, same specimen as Sugiyama (1991 , pI. I, fig. 3), TH-I , paratype. 4a, 4b: ESN 146506, TH-2,
paratype. This specimen has reduced D and lacks V'. Scale bars: la, 2a, 3a and 4a = 50 11m ; others = 20

11m.

950.

Lower and Middle Miocene lophophaenids

55

56

Kazuhiro Sugiyama

much more variable than the result confirmedin this study.
Dimensions (in J.lm).-Based on measurement of 22 completely preserved specimens
examined by SEM. Height of test; 50-93.
Maximum width of test; 60-98. Maximum
diameter of PR; 18-30.
Occurrence.-TH-l,2,3,4 and 00-17,18(very rare in abundance).
Genus Steganocubus Sugiyama, gen. nov.

Etymology.-From the Greek steganos,
covered, and kybos, a cube; masculine gender.
Included species.-S. subtilis Sygiyama sp.
nov. (type species) and S. /ipus Sugiyama sp.
nov.
Diagnosis.-Large cephalis enclosing internal skeleton. A, two I", V" relatively long.
A', D, two L, two I' and V" commonly extending to the shell wall. They are, however,sometimes reduced or absent irregularly (Figures 1O-4b; 11-1 b). I" usually lies in abisectional position between D and L. D'
and L' vestigial (Figure 10-1 d) or completely
absent. V missing. L-R and Ax variable inlength. A few, short connector bars irregularly developed especially on the ventral side.
They connect PR and the shell wall or DR.
DR lies in a high position in the cephalis.
Y IX is less than 1. Thorax may be present.
Remarks.- This genus is distinguishedfrom Pseudocubus externally by the presenceof the shell wall, and internally by having
stout and long A', D and V', and by lacking
V completely. This genus was undoubtedly
derived from Pseudocubus by direct acquisition of the shell wall on the skeleton, accompanied by a certain modification of theskeletal structure. It is, therefore, clear thatthe segment above DR corresponds not to thewhole cephalis but to the upper part of the
eucephalic lobe.
Helotholus vema Hays, 1965, seems to nearly satisfy the generic diagnosis of this genus,
and one can be fairly certain that H. vema is
more or less related to the present new genus.
However, it often has V in the skeleton (e.g.
Petrushevskaya, 1971 b, figs. 24-2, 3 and 4;
Chen, 1975, pI. 16, fig. 4: Weaver, 1976, pI. 1,
fig. 11). Moreover, a personal preliminary
investigation, the results of which will be
published in the near future, revealed that the
typical form of H. vema has six longitudinal
scaffolds as illustrated by Petrushevskaya(1971 b, fig. 17-5) and another well known
form having bladed scaffolds including four
longitudinal ones and lacking Vas illustratedby Lazarus (1990, figs. 1-5) is possibly
conspecific with Pseudocubus warreni Goll,
1980. Furthermore, as mentioned in the
introductory section, Kellogg (1975) andWeaver (1983) concluded that the typicalform of H. vema was derived from a moreelongated ancestor H. praevema Weaver,
1983, by the sinking of the cephalis into thethorax and simultaneous increase of thoracic
width during a short interval. The latter
species may lack PR free from the shell wall
judging from Weaver's (1983) originaldescription. If their observations and conclusion are right, H. vema and two species ofSteganocubus gen. nov. are regarded as convergent forms having phylogenetically independent PR free from the shell wall.
Although such a situation is practicallysuspected, I cannot reject it completely atpresent. The assignment of H. vema to thisnew genus is reserved at this time based on theabove mentioned facts.

Steganocubus subtilis Sugiyama, sp. nov.

~ Figure 12.
la-4b: Steganocubus /ipus Sugiyama, gen. et sp. nov. la, Ib: ESN 146508, 00-27,
holotype. 2a, 2b: ESN 146509,00-28, paratype. 3a, 3b: ESN 146510,00-27, paratype. This specimen
has an irregular skeletal structure. 4a, 4b: ESN 146511,00-28, paratype. Arrows of Ib, 2b and 4b point
to I". Scale bars: la, 2a, 3a and 4a=50 Jim; others=20 Jim.

950.

Lower and Middle Miocene lophophaenids

57

Kazuhiro Sugiyama

58
Figures 10-1 a-4b; 11-1 a, I b

Helotholus vema Hays-Sugiyama, 1992, pI. I, fig. 3.

Description.-Cephalis hemispherical todome-shaped, porous, with or without short
external spines. It is composed of three lobesof which the eucephalic one is usually discernible externally by a certain stricture corresponding to the place of DR. Ante- andpostcephalic lobes are equal in size. Thorax
may be developed, but is short with an
obscure collar stricture. Wall thin, bearing
numerous, short by-spines on the upper half.
Pores are circular to oval, variable in size,
without regular arrangement. Skeletal construction as with genus: Ax wartlike orabsent. All of the elements of the skeleton
are circular in cross section.
Remarks.- This species is similar to
Helotholus vema Hays but is distinguishedfrom the latter species by its smaller test
dimensions, common development of 0, shorter Ax, and complete lack of V.
Dimensions (in jlm).-Based on measure-

F

I~:~~
L-R--f~
LI

L-R

Lr

DR

Figure 13. Schematic illustration of typicalskeletal structure of Botryopera (?) leptostraca sp.
nov. (a): Dorsal view. (b): Right lateral View.
(c): Basal view.

ment of 42 specimens examined by SEM.
Height of test; 50-93. Maximum width oftest; 55-85. Maximum diameter of PR:
15-30.
Holotype.-ESN 146504 (Figs. 1O-la-ld)
from TH-4.
Occurrence.-TH-l, 2, 3 and 4 (common inabundance ).
Etymology.-From the Latin subtilis, minute.

Steganocubus /ipus Sugiyama, sp. nov.
Figures 12-la-4b

Description.-Cephalis is composed internally of three lobes of ante-, eu- and postcephalis. They are, however, hardly recognized externally owing to the secondary development of the micro granular covering on theupper part of the cephalic wall. Thorax
relatively long, not closed distally, having amore or less distorted shape in cross section.
Collar stricture may be distinct. Pores are
circular to oval, deeply set in pore frames.
They are without regular arrangement on theupper part of the cephalic wall, but show
more or less longitudinal arrangement on thelower part of the cephalic and the wholethoracic wall. Numerous short by-spines
extending from the pore frames. Skeletal
structure is identical to that of S. subtilis sp.
nov. but the development of Ax is more
prominent.
Remarks.- This species is morphologicallydistinguished from S. subtilis by having amore elongated and thicker shell wall, moreor less regularly arranged pores, and distinct
Ax. However, the true phylogenetic relationship between the two species is at present
uncertain, and they possibly represent only
morphologic variants of one and the same
species.
Dimensions (in jlm).-Based on measure-~
Figure 14. la-4b: Botryopera (?) leptostraca Sugiyama sp. nov. la, Ib: ESN 146512, OD-27,
holotype. 2a, 2b: ESN 146513, OD-17, paratype. 3a, 3b: ESN 146514, OD-17, paratype. 4a, 4b: ESN
146515, OD-21, paratype. Arrows of la, 2a and 3a point to delicate meshwork between by-spines. Arrowsof Ib and 2b point to short I". Sa, 5b: Botryopera (?) whitei (Bj\'irklund), ESN 146516, OD-IS. Scalebars: la, 2a, 3a, 4a and 5a=50 pm; others = 20 pm.

950.

Lower and Middle Miocene lophophaenids

59

Kazuhiro Sugiyama

60

fore, the generic assignment of the following
three species remains tentative. In order tosolve this problem it is necessary to redescribe
the type species in detail, including its
skeletal structure.

Botryopera (?) leptostraca
Sugiyama, sp. nov.
Figures 14--1a-4b

Figure 15. Schematic illustrations of typicalskeletal structure of Botryopera (?) whitei (Bj¢rklund). (a): Dorsal view. (b): Right lateralview. (c): Basal view.

ment of 18 specimens examined by SEM.
Height of test; 65-123. Maximum diameterof test; 65-100. Maximum diameter of PR ;10-25.
Holotype.-ESN 146508 (Figures 12-1a,Ib) from 00-27.
Occurrence.-OD-17, 18,26,27 and 28(very rare in abundance).
Etymology.-From the Greek lipos, fat.

Genus Botryopera Haeckel, 1887
Botryopera Haeckel, 1887, p. 1108.
Trisulcus Popofsky, 1913, sensu
1971b, p. 140 (partim).

Petrushevskaya,

Type species.-B. cyrtoloba Haeckel, 1887
(designated by Petrushevskaya, 1975).
Remarks.- The generic concept used inthis study mainly follows that of Petrushevskaya (1975, 1981, 1986). This genus is easily
distinguished from Pseudocubus andSteganocubus gen. nov. by its large eucephalic lobe sitting on DR. This seems to beclosely related to the value of YjX, which ismuch larger than 1 in this genus. One canremark that the detailed skeletal structure isat the present time not well understood withrespect to every species in the genus. There-

Description.-Eucephalic wall pierced bynumerous circular to oval pores of relatively
small size, possessing a large number of byspines on the surface, rimmed by DR on thelower margin. Perforate, delicate shell walldevelops downward from DR or slightly
higher level than DR. It is cylindrical or
truncated conical in the lateral view, andcorresponds to ante- and postcephalic wall.
Delicate meshwork may also develop between
by-spines (Figs. 14-1a, 2a and 3a). DR and
PR are always connected by a part of A, two
I", V and two pillars developed near thejunctions of two L-R and PRo Further connector bars may be present between bothrings. A' well-developed. D and two L
have one verticil of one to three spinules ofwhich D' and L' are moderately developedand sometimes extend to the shell wall. Two
I' are proximally amalgamated with two short
1". V situated in the sagittal plane and may
penetrate the shell wall outside beyond thejunction of DR. V' and V" are completely
absent. All of the skeletal elements are circular in cross section.
Remarks.- This species is similar to
Botryopera (?) pseudoantarctissa Petrushevskaya in Petrushevskaya and Kozlova (1979),
but differs from the latter species in having V
in the sagittal plane, and rather delicate shellwall.
Dimensions (in ,um).-Based on measurement of 24 specimens examined by SEM.
Height of test; 65-95. Meximum width oftest; 65-73. Maximum diameter of PR:
15-25.
Holotype.-ESN 146512 (Figures 14-1a

950.

Lower and Middle Miocene lophophaenids

Figure 16. 1a-6b : Botryopera (?) whitei (Bj¢rklund). 1a,lb : ESN 146517,00-18. 2a,2b : ESN
146518, 00-18 . 3a, 3b : ESN 146519, 00-17. 4a, 4b : ESN 146520, 00-17. 5 : ESN 146521 , 00-18 .
6a, 6b : ESN 146522,00-17. Scale bars : l a, 2a, 3a, 4a, 5 and 6a = 50jim ; others = 20 jim.

61

62

Kazuhiro Sugiyama

950.

Lower and Middle Miocene lophophaenids-Ib) from 00-27.

Occurrence.-TH-4, 00-17,18,21,26,27
and 28 (very rate in abundance).
Etymology.-From the Greek, leptos, delicate, and ostracon, shell.

Botryopera (?) whitei (Bj¢rklund)
Figures 14-5a-5b; 16-la-6b; 17-la-3

Antarctissa whitei Bjprklund, 1976, p. 1125, pI. 13,
figs. 9-14, pI. 15, fig. 4.
Botryopera oceanica whitei (Bjprklund)-Petrushevskaya and Kozlova, 1979, p. 122, figs. 325-328.

Description.-Shell dome- to bullet-shaped,
entirely spongy due to the development of
secondary meshwork all around. Thick bristles may develop on the shell surface. Primary lattice work of eucephalis is, however,relatively rough, and the size of pores is thus
similar to Cryptogyrus trachylobus sp. nov.
rather than to B. (?) leptostraca sp. nov.
(Figure 16-6a). Shallow stricture rarely expressed on the shell wall. Fundamental
skeletal structure is common with that of B.
(?) leptostraca sp. nov. It is, however, much
more complicated by the development ofadditional connector bars between PR, DR
and the shell wall. PR is quite irregular inoutline. DR is usually embedded in theshell wall, so that the position is not clear
from the basal view. D' and L' well developed, sometimes extending outside the shellwall. V usually swerves from the sagittal
plane and makes an acute angle with Lr,
rarely with Ll. It is sometimes free from DR
and directly penetrates the shell wall beyond
the junction of PRo V' and V" may bepresent but are hardly distinguished fromother connector bars.
Remarks.-Although Petrushevskaya andKozlova (1979) treated this species as a subspecies of B. oceanica (Ehrenberg, 1872), suchspongy externality may be diagnostic for this~63

species group alone. BWrklund (1976)
mentioned that the occurrence of this species
in the Norwegian Sea is restricted to theUpper Miocene to Pliocene, but the Oidawara
Formation is undoubtedly lowest Middle
Miocene judging from all available geological and paleontological data. The age
difference suggests the immigration of it fromthe western Pacific to North Atlantic or thatthe occurrence was strongly affected by
paleooceanographic conditions. Further
examination is necessary to clarify the problem. B. (?) whitei differs from B. (?) leptostraca sp. nov. in its much spongier shell walland swerved V. Even if V lies near thesagittal plane (Figs. 16-3a, 3b), the former
species can be distinguished from the latter by
the development of D', L', V', and spongy
network on the shell wall. It is very difficult
to distinguish this species from Steganocubus/ipus sp. nov. when the specimens are laterally
set on a strewn slide.
Dimension (in J.lm).-Based on measurement of 47 specimens examined by SEM.
Height of test; 58-100. Maximum width of

(b)

(a)

DR

D~·
. ; "oo-;-..) oo-...}..
~~

/
PA'-

U

L·R

~~
V

L-R

Figure 18. Schematic illustrations of typicalskeletal structure of Cryptogyrus trachylobus gen. et
sp. nov. (a): Dorsal view. (b): Right lateralview. (c): basal view.

Figure 17. la-3: Botryopera (?) whitei (Bj9rklund). la, Ib: ESN 146523, OD-18. 2a, 2b: ESN
146524, OD-18. 3: ESN 146525, OD-17, transmitted light micrograph. Arrow points to swerved V.
4a-4d: Botryopera (?) sp. A, ESN 146526, TH-4. Arrows of 4b point to two 1". Scale bars: la, 2a, 3 and
4a=50 j.lm; others = 20 j.lm.·

64

K azuhiro Sugiyama

950.
test; 50-75.
15-30.

Lower and Middle Miocene lophophaenids

Maximum diameter of PR;

Occurrence.-OD-17, 18,21,26,27 and 28(rare in abundance).

Botryopera (?) sp. A
Figures 17--4a-4d

Description.-Eucephalic wall rugged,
pierced by numerous circular to oval pores ofrelatively small size, having many rodlike
by-spines on the upper half, edged by DR on
the bottom. Additional shell wall developed from slightly higher level than DR(Figure 17-4c). It is cylindrical in shape,
quite delicate, and incomplete in the proximal part. The skeletal structure is nearly
identical to that of B. (?) leptostraca sp. nov.,
but differs from the latter in having questionable V' which is proximally amalgamatedwith V and distally bends downward (Figures
17-4b-4d). V extends beyond the junctionof PR, and penetrates the shell wall. D'
vestigial. L' missing.
Remarks.- This species is distinguishedfrom B. (?) leptostraca and B. (?) whitei inhaving numerous by-spines on the upper partof the eucephalis and much thinner additional shell wall. The existence of questionable V' is also diagnostic of this species.
The proposal of a new name is postponed
because of the small number of available
individuals.
Dimensions (in ,um).-Based on two specimens examined by SEM. Height of test;
85-90. Maximum width of test; 60-63.
Maximum diameter of PR; 15-18.
Occurrence.- TH-4 (very rare in abundance).
Genus Cryptogyrus Sugiyama, gen. nov.

65

Etymology.-From the Greek cryptos, hidden, and gyros, circle; masculine gender.
Type species.-C. trachylobus Sugiyama
sp. nov. (by monotypy).
Diagnosis.- Test consisting of exclusively
large eucephalis, with or without loosely
defined ante- and postcephalic lattice shell,
and having a distinct ring structure in theskeleton. PR is thick, circular in cross section. DR is composed of four arches which
are convex upward and forms the lower
margin of a large eucephalic chamber. Y IX
is much larger than 1. A is short, usually notforming external prolongation. A' absent.
D and two L relatively thin. D' and L' may
be distinct. I' and V' missing. Two I" andV" are short. When one views from thebasal side, two I" always lie in a plane perpendicular to the sagittal plane. V is commonly present, but rarely absent as an
intraspecific variation. In the latter cases,
PR and DR are connected by five bars consisting of A, two I" and two pillars developed on
the ventral side (Figure 19-4b). There areno additional connector bars between PR andadditional lattice wall.
Remarks.- This genus differs from
Botryopera in lacking A', two I' and V', andhaving only four connector bars between PR
and DR in typical specimens. Further
remarks are given in the discussion of the typespecies.
Cryptogyrus trachylobus Sugiyama, sp. nov.
Figures 19-1a--5; 20-la--2b

Description.-Skeletal structure as withgenus: D and two L variable in length.
They are sometimes markedly long, and irregularly curved and bifurcated distally (Fig. 19-ld), and often have a few conical spinules+- Figure 19. la-5: Cryptogyrus trachylobus Sugiyama, gen. et sp. no~. 1a-~e: ESN. 146527, O~-27,
holotype. Arrows of I b point to the distal part of V and thick arch connectmg L With by-spme, respectively.
Arrow of Ie also points to the distal part of V. Arrow of Id points to the bifurcation of L. 2a, 2b: ESN
14652S, OD-17, paratype. D of this specimen is broken off. 3: ESN 146529, OD-IS, paratype. 4a-4b:
ESN 146530, OD-IS, paratype. Arrows of 4b point to two irregularly developed pillars between PR and
DR. 5: ESN 146531, OD-17, paratype. Scale bars: la, Ib, 2a, 3, 4a and 5=50jlm; others=20jlm.

66

K azuhiro Sugiyama

950.

Lower and Middle Miocene lophophaenids

near the junctions of A and L-R. D' and L'
may be recognizable among the spinules.
Big eucephalis supported by the skeleton,
pierced by large, oval to circular pores, having numerous by-spines on the surface.
Among the by-spines, those extending fromthe lower margin of the eucephalic lattice
shell, which are directed obliquely downward, are commonly much longer and thicker. Additional thick arches often growbetween the tips of them and distal parts of D
and two L (Figures 19-1 b, 20-2a). The
development of secondary network between
neighboring by-spines is also present in somespecimens (Figure 20-1 a). Adult specimens(?) (Figures 19-2a, 3a) have coarse latticewall which corresponds to ante- and postce-

67

Figure 21. Schematic illustrations of typicalskeletal structure of Ceratocyrtis sp. aff. C. histricosa (J¢rgensen). (a): Dorsal view. (b): Rightlateral view. (c): Basal view.

(a)

(b)
A

•....

,,

,,

v

- - - - - - - -L-R

~

,

I

:Ax

Lr

L'
(c)

(d)

Lr

•....

o

~

L'

Lr

o

LI

L'

LI

Figure 22. Schematic illustrations of skeletal variation of the genus Ceratocyrtis. (a) and (b); rightlateral view, (c) and (d); basal view. (a) and (c); extreme structure with L-R connecting Land PR, (b)and (d); another extreme structure with L-R connecting L' and PRo~
Figure 20. la-2b.: Cryptogyrus trachylobus Sugiyama, gen. et sp. nov. la, lb: ESN 146532,
00-18, paratype. Arrow of la points to secondary arch between by-spines. 2a, 2b: ESN 146533,00-18,
para type. Arrows of 2a point to well developed arches connecting two L with by-spines. 3a, 3b:
Ceratocyrtis sp. aff. C. histricosa (J¢rgensen), ESN 146534, TH-4. Arrow of 3b points to short L-R.
4a-6: Ceratocyrtis stoermeri Goll and 8j¢rklund. 4a, 4b: ESN 146535, 00-28. Arrow of 4b points tothe tip of Ax. Sa, 5b: ESN 146536, 00-26. Arrow of 5b points to short L-R. 6: ESN 146537, 00-26.
Scale bars: 1a, 2a, 3a, 4a, 4b, 5a and 6 = 50 Ii m; others = 20 Ii m.

68

K azuhiro Sugiyama

Figure 23. la, lb: Ceratocyrtis sp. aff. C. hislricosa (J¢rgensen), ESN 146538, TH-4. Arrows of 1b
point to L-R. 2a, 2b: Lophophaena sp., ESN 146842. White arrows of 2b point D', black one does
L-R. This specimen is from the Pliocene Nobori Formation, Shikoku, Southwest Japa n (corresponding
to the Spongaster pentas Zone of Riedel and Sanfilippo, 1978). Further information is given in Sugiyama
el al. (1992). 3a, 3b: Pseudodictyophimus lectari Caulet, ESN 146843. Black arrows of 3b point to D',
white ones to L-R. Locality same as for the preceding species. Scale bars: la, 2a a nd 3a = 50 j.lm;
others=20 j.lm.

phalic wall and is formed by further development of secondary network below the eucephalic chamber. The wall roughly encloses
the basic internal skeleton.
Remarks. - The development of additionallattice wall below the eucephalis has quite awide variational range in the population. It
is, however, continuous between individuals,
and may represent either ontogenetic stages or
phenotypic variation of the adult stage.
When the additional wall is indefinable, thisspecies is similar to the species of the genera
Peridium Haeckel , 1881 , Phormacantha J¢rgensen, 1905, and Plectacantha J¢rgen sen,
1905. The former species is, however, distin-

guished from them by having two thick I "arranged in a plane perpendicul~r to thesagittal plane, lacking a long extern"al prolongation of A, and having ante- and postcephalis in adult specimens (?) . This species
resembles Botryopera (?) leptostraca sp. nov. ,from which it differs in having larger pores on
the eucephalis and the features mentioned in
the generic remarks.
Dimension (in j.lm) .-Based on measurement of 22 specimens examined by SEM.
Height of test ; 60-93 . Maximum width oftest; 55-85. Maximum diameter of PR ;10-25.
H%type.-ESN 146527 (Figures 19- 1a

950.

Lower and Middle Miocene lophophaenids-Ie) from OD-27.

Occurrence.-TH-4,

OD-17, 18,21,26,27
and 28 (very rare in abundance).
Etymology.-From the Greek trachys,rough, and lobos, lobe.
Genus Ceratocyrtis Biitschli, 1882,
emend. Petrushevskaya, 197 I b
Ceratocyrtis BUtschli, 1882, p. 536.-Petrushevskaya,
1971b, p.98.
He/otho/us Jprgensen, 1905, p. 137.
Bath/ocalpis Clark and Campbell, 1942, p. 64.

Type

species.- Corn u tella ?

cucullaris

Ehrenberg, 1873 (designated by Petrushevskaya, 197 I b).
Remarks.- The generic concept including
the synonymy follows Petrushevskaya(l97Ib).

Ceratocyrtis stoermeri Golland BWrklund
Figures 20-4a-6
Ceratocyrlis stoermeri Goll and Bj¢rklund, 1989, p.
731, pI. 5, figs. 5-9.

Description.- Test large, two-segmented
form with ambiguous external stricture. The
first segment very small, pierced by small,circular to subcircular pores set in thickframes from which arise small by-spines.
The second segment large, campanulate, moreor less tapering distally, pierced by large,
subcircular to circular pores surrounded bycircular to polygonal-shaped thick frames:
the upper part is spinous; the middle part,
where the thorax is widest and the ridges ofthe pore frames are sharpest, has the largest
diameter of the pores; and the lower part israther smooth in outline. The terminationof the second segment is in line in lateralview, often accompanied by a delicate sieve
plate at the base. Two segments internallypartitioned by PR which is composed of
distinct arches but nearly horizontal. MB
relatively short. A makes a nearly right69

angle from the horizontal, and merges into
PRo D extends downward with a large angle
from the horizontal, and may have one, rarely
a few branches in the distal part. Two L
extending slightly downward, having distinct
L-R in the middle part, and may branch off inthe distal part like D. L' rarely present. V
connects with PRo Ax thick, long, usuallybifurcated in the distal part.
Remarks.-Morphology of this species isquite stable except for the development of thebasal sieve plate.
Dimensions (in ,um).-Based on I I specimens examined by SEM. Height of test;
120- 150. Maximum width of test; 105-140.
Maximum diameter of PR; 20-30.
Occurrence.-OD-17, 18 (very rare in abundance) and OD-26,27 and 28 (rare in abundance).

Ceratocyrtis sp. aff. C. histricosa(J¢rgensen)
Figures 20-3a, 3b; 23-1 a, I baff. Helotho/us histricosa J¢rgensen, 1905, p. 137, pI.
16, figs. 86-88.
Ceratocyrtis sp. aff. C. histricosus (Jprgensen)-Sugiyama, 1991, pI. I, fig. 9.

Description.- Test large, two-segmented bya distinct external stricture which corresponds
to the place of PRo The first segment hemispherical, small, pierced by circular to subcircular pores of small size, ornamented bynumerous, long by-spines arising from thepore frames, corresponding to upper part ofeucephalis. The second segment
campanulate, large, pierced by circular to
subcircular pores of large size, having longby-spines on the upper half. Skeletal structure as with C. stoermeri. A and V usuallypenetrate the shell wall to form conical to
rodlike spines which are longer and thicker
than by-spines. L' is commonly well developed, then L-R may connect the proximal
part of L' and PR (Fig. 22).
Remarks.- This species differs from C.
stoermeri in having distinct stricture between

Kazuhiro Sugiyama

70

the first and second segments, longer byspines on the shell surface, external prolongations of A and V, and the lack of prominentridges on the pore frames. It also differsfrom C. histricosa (J¢rgensen) in having amore spinous shell wall and longer second
segment.
Dimensions (in JLm).-Based on ten specimens examined by SEM. Height of test;
90-160. Maximum width of test; 80-160.
Maximum diameter of PR; 20-35.
Occurrence.- TH-l, 2, 3 and 4 (rare inabundance).

Acknowledgments
I wish to express my hearty thanks to Prof.
Junji Itoigawa of Nagoya University who
critically revised the manuscript and gave me
useful suggestions. I am very grateful to Dr.
Atsushi Takemura of Hyogo University ofTeacher Education for his helpful criticism of
the manuscript and generously providing
some Antarctic reference samples. I am alsoindebted to the following persons for many
encouraging discussions during the study:
Prof. Hiroshi Shibata, Drs. Tomowo Ozawa
and Shinobu Mori and Mr. Hiroaki Karasawa of Nagoya University and Dr. Hiroshi
Furutani of Museum of Nature and Human
Activities, Hyogo. Thanks are due to Dr.
Paulian Dumitrica, Institute of Geology andGeophysics, Romania, for his reading of thefirst version of manuscript. Sincere thanks
also go to anonymous reviewers who provided valuable comments on the manuscript.

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72

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Krasheninnikov, V.A., et al., Ibid., vol. 71, p.
667-686, pis. 1-6.

:Fffi!i'.Jll, Morozaki JiffiUffiJ, Oidawara iEf1R, Toyohama

~~.

,=

I:j:lHlH3 *""f$~I:j:l$l:j:lwr~,nii Lophophaenidae f4~mr9:!(7)'1N~~~ -:> \, \ -r : ~ffi J!f< §ffi
UffiJ~~~~~~(r$~~~)~la~~J!f<:Fffi!~~~iEf1R~~(~$~~~)l~~Gntc. 5 ~ 9 fjO) Lophophaenidae t+1.rzliJl.~!t~~O)'Fftll-:m~z.~W L 1':0 .:t GIi 7 fj~o)!J -1n7'v;:'*IH:5t~n, -f""'-C0)!J 17·v;:.:5t~~$c L-C:m:~U~;,hn~~~:k:m~iJ~~:g.)bGn~o 2~~4~fjz.m~Ltc.o~W~~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 169, pp. 73-96,7 Figs., April 30, 1993

951. A FOSSIL POPULATION OF GINKGO LEAVES FROMTHE XINGYUAN FORMATION, INNER MONGOLIA *LI-MING ZHAO
Institute of Botany, Academia Sinica,
Xiangshan, Beijing, P.R. ChinaandTAMIKO OHANA and TATSUAKI KIMURA
Institute of Natural History, 24-14-3 Takada,Toshima-ku, Tokyo, 171

Abstract. Fossil Ginkgo is one of the index genera characterizing the Late JurassicEarly Cretaceous Tetori-type and Siberian-type floras in Eastern Eurasia. Fossil Ginkgo
leaves are sometimes crowded in occurrence, thickly massed and appressed on the same
bedding plane, so representing a fossil population. One of us (Zhao) and his Chinese
coworkers collected a number of detached fossil Ginkgo leaves from such an occurrence fromthe Xingyuan Formation (=Jiufotang Formation) at the Xilutian Open Coal-Mine, near
Chifeng area, Zhaowudameng District, Inner Mongolia. These Ginkgo leaves are varied insize, form and mode of lamina-division. Nevertheless, they form a single fossil population,and thus belong to the same species, because their morphological features are similar and arethought to be within the range of variation and their cuticular features except for minor
points are the same. This paper deals with the description of these leaves, with geographicaland stratigraphical distributions of fossil Ginkgo leaves in the western part of Northeast
China, and with generic and specific revision of some previous works and makes a brief
comparison of the present population with those hitherto known. The present populationof fossil Ginkgo leaves is at least the same as those of Fuxin, Shahezi, Damoguaihe andDabao Formations in Northeast China. In the case of a small number of fossil Ginkgo
leaves, it is difficult to identify them down to species so far as they do not show their uniquemorphological or cuticular features. The present study shows that it is difficult to assess any
distinction between the genera Ginkgo and Baiera. The Ginkgo leaves comprising thepresent population is named by us as Ginkgo manchurica (Yabe et Oishi) Meng et Chen.
Key words. Fossil Ginkgo leaves, fossil Ginkgo population, younger Mesozoic,
Northeast China.

China. As to the geology of this area, noformal paper has been published except for abrief note by Shang and Zheng (1991).
According to them, the Mesozoic sedimentsin this area are divided into Xingyuan (ca.
380 m thick), Yuanbaoshan (ca. 186 m thick)and Shuiquan (ca. 245 m thick) Formations
in upward sequence. The Xingyuan Formation is equivalent to the liufotang Formation

Introduction

Fossiliferous younger Mesozoic sedimentsof nonmarine origin bearing rich coal-seams
are developed in the Chifeng area, Zhaowudameng District (roughly 41' 1T -4Y24'N
and 116'21'-120'59'£), Inner Mongolia,'Received August 20, 1992; accepted September 27,
1992
73

74

Li-Ming Zhao et al.

wihch is extensively distributed in West
Liaoning Province (Figure 1 and Table 2).
They listed the following plant taxa fromthe Xilutian Open Coal-Mine of the Pingzhuan Coal-Field (Figure 1) (*; nomen
nudum) :From the Xingyuan Formation (=Jiufotang Formation): Acanthopteris alata(Fontaine) Zhang, Coniopteris saportana
Heer, Nilssoniopteris chifengensis sp. nov. *,Nilssonia sinensis Yabe et Oishi, Ginkgo
manchurica (Yabe et Oishi) Meng et Chen,
G. tapkensis Doludenko et Rasskazova,
Sphenobaiera longifolia (Pomel) Florin, S.
nantianmenensis Wang, Czecanowskia dgida
Heer, Ixostrobus heed Prynada, Pityolepis
piceaoides sp. nov. *, P. pseudotsugaoides sp.
nov. *, Tsuga taxoides Tan et Zhu, T.
zhouiana sp. nov. *, Sphenolepis kurriana(Dunker) Schenk, Elatides curvifolia (Dunker) Nathorst, E. harrisii Zhou, Cephalotaxopsis asiatica HBDYS**, Sabinites neimongolica Tan et Zhu, Pagiophyllum cf. triangulare Prynada, Elatocladus smittiana (Heer)
Seward, E. yuanbaoshanensis sp. nov. *, E.
manchurica Yabe et Oishi, E. sp., Lindleycladus eichwaldii (Schimper), Scarburgia
triangularis Meng, Conites sp. (**; not theauthor's or authors' name but the name of theinstitution: HBDYS being after Huabei
Dizhi Yanjin Sou in Chinese = North China
Institute of Geosciences)
From the Yuanbaoshan Formation:
Acanthopteris onychioides (Vassilevskaja et
Kara-Mursa) Zhang, Arctopteris kolymensis
Samylina, Cladophlebis sp., Doratophyllum
nilssoniopteroides sp. nov. *, Ginkgo concinna
Heer, G. manchurica (Yabe et Oishi) Meng et
Chen, G. sibirica Heer, G. tapkensis Doludenko et Rasskazova, G. truncata Li,
Sphenobaiera biloba Prynada, Phoenicopsis
angustifolia Heer, Pityophyllum sp., Pityolepis
sp.
The fossil plant taxa listed above should becritically reexamined; for instance, thegeneric name Lindleycladus established byHarris (1979) would at present be valid only

for the former Podozamites lanceolatus (Lindley et Hutton) leafy shoots with longitudinally oriented stomata from the Yorkshire
Jurassic. However, at a glance from theabove list, the fossil plant assemblages of bothformations look to be of the Tetori type (e.g.
Kimura, 1987a, b), though the list shows
many coniferous taxa which are questionablein identification.
Recently, one of us (Zhao) and his Chinese
coworkers collected a number of fossil plantsfrom the Xingyuan Formation at the XilutianOpen Coal-Mine (Figure I) for the purpose of
making detailed palaeobotanical studies on
the Xingyuan plant assemblage (Xingyuan
flora). Previous to our studies, Zhao andTao (1991) described a new Ginkgo species
and illustrated several fossil plants collectedfrom the Xingyuan Formation.
In this paper, we describe Ginkgo leaves
from the Xingyuan Formation. These leaves
are crowded in occurrence, sometimes thickly
massed and appressed on the same bedding
plane. In addition, we revise and briefly
review the Ginkgo leaves hitherto describedor recorded from the younger Mesozoic
plant sites in the western part of Northeast
China, and make a brief comparison of thepresent population of Ginkgo leaves withthose hitherto known.
As to the geological age of the Xingyuan(or Jiufotang) Formation, there are twodifferent opinions among Chinese geologists
and palaeontologists; one regards it as latest
Jurassic and the other as earliest Cretaceous.
We are of the opinion that at present it isdifficult to make a precise age determinationof the Jiufotang Formation and its equivalent, because this formation is wholly of
non-marine origin and has not yielded any
fossils indicating its precise age.
Acknowledgements

One of us (Zhao) gives his sincere gratitude
to Assoc. Prof. J.R. Tao of the Institute ofBotany, Academia Sinica, Beijing for her

951.

A fossil population of Ginkgo leaves

75

the Ministry of Education, Science and Culture, Japan (given to Kimura; no. 02640615
in part and given to Kimura and Ohana' no.
04640725 in part).
'
Material and methods

20km

I0000.-_ _ _,,......_.......'

Figure 1. Location of the fossil plant site in thepresent work. A: Xilutian Open Coal-Mine. B:
Fossil plant site belonging to the Xingyuan Formation (= liufotang Formation), corresponding toLocality no. 22 in Figure 7 and Table 2. C: Pingzhuang Station. According to the informal
communication from K. Zhao, the Xingyuan(= liufotang) and Yuanbaoshan (= Shahai + Fuxin)
Formations are distributed in this area (see Table 2).

continuous guidance and encouragement andto Mr. P. Shang of Fuxin College of Mines
and C.X. He, a student of Assoc. Prof. Taofor their kind help in collecting the fossil
plants described herein. Zhao's thanks areextended to the Kambayashi ScholarshipFoundation and the Urban Land-System
Institution Co., Tokyo for their financial
support during his stay (for one year) inJapan. We are indebted to Dr. Shya
Chitaley of the Cleveland Museum of Natural
History, Ohio for going over the language ofthe manuscript. This study was supported
by Grant-in-Aid for Scientific Research from

Fossil Ginkgo leaves were collected fromthe Xingyuan (Jiufotang) Formation at the
Xilutian Open Coal-Mine (roughly Lat. 42°
OO'N; long. 119°12'E) near Pingzhuang,
Chifeng, Zhaowudameng, Inner Mongolia(Figure 1 ; indicated by '22' in Figure 7).
These leaves for cuticular study were peeled
off from tuffaceous shale and treated in Schulze's solution followed by diluted potassiumhydroxide (ca. 10%), stained with safranin,
and then washed in alcohol and xylene before
mounting in canada balsam. For the SEM
observations cuticles were sputter coated withgold and photographed with a Hitachi S-800,
20 kv.
The specimens, cuticular preparations andSEM stages examined and described in thispaper are kept in the Institute of Botany,
Academia Sinica, Xiangshan, Beijing.
Description
Ginkgoales
Genus Ginkgo Linnaeus
Ginkgo manchurica (Y a be et Oishi)
Meng et Chen
Figures 2-6
1991. Ginkgoites pingzhuangensis Zhao et Tao: Zhao
and Tao, p. 965, pI. 2, figs. 14-16.

Specimens examined: NCP 8259-8273,
8277-8278,8283,8288-8291,8332,8343,8345,
8348-8351,8353-8354,8367,8369.
Locality and horizon: Xilutian Open
Coal-Mine: Xingyuan (Jiufotang) Formation (possibly uppermost Jurassic or lowest
Cretaceous).
Occurrence: Very abundant.
Description of specimens: Obtained speci-

76

Li-Ming Zhao et al.

mens were all detached leaves; some withpreserved petiole. Cuticle is well preserved.
I) External morphology (Figures 2- 3) :Laminae are generally fan-shaped or semicircular in form, but remarkably varied in size,\.5-13 cm in radius (or high). Petioles arelong and slend er, 1- 2 mm thick; the longest
one is 3.8 cm long, but their abscission part ismissing in general. The lamina is fundamen-tally divided into two parts by a deep mediansinus reaching almost to the tip of the petiole
to form the most simple lamina. Each part isgenerally further dichotomously divided once
to three or more times into broader to fairly
slender segments or lobes by deeper or shallower sinuses. The final number of ultimate
segments or lobes is 24 in each part (Figures
21,31, 2m, 3m,) .

Figure 2. Ginkgo manchul'ica (Yabe et Oishi) Meng et Chen. Photographs ; a ll in natural size.
Detached leaves of vaned sizes, forms and modes of the lamina-division. A-type of leaves' 2a(NCP-8350), 2b (NCP-8348). B-type of leaves : 2c (NCP-8345), 2d (NCP-8332) . C-type of leaves: 2e(NCP-8347), 2f (NCP-8353), 2g (NCP-8290), 2h (NCP-8277), 2i (NCP-8343). D-type of leaves': 2j(NCP-8260), 2k (NCP-8359), 21 (NCP-8266). E-type of leaf: 2m (NCP-8265).

951.

A fossil population of Ginkgo leaves

We provi sion ally cl ass ified o ur leaves(lamin ae) into the follo w ing five types (A-E
types) :(1) A-type of lamin a (Figures 2a, 3a, 2b,
3b, 3n , 30): smaller in size (typi ca lly 1.5 cm
in radius) , di vided totally into 4- 6 segme nts.

2e(C)

77

(2) B-type of la mina (Figures 2c, 3c, 2d,
3d, 3p) : Rather larger in size (typically 2.5
cm in radi us), divided tota ll y into 4- 6 segments.
(3) C-type of la mina (Figures 2e, 3e, 2f,
3r, 2g, 3g, 2h, 3h , 2i, 3i): Medium-sized (typi-

78

Li-Ming Zhao et al.

cally 3 cm in radius), divided totally into 814 segments or lobes.
(4) D-type of lamina (Figures 2j, 3j, 2k,
3k, 21, 31): Rather much larger in size (typically 5 cm in radi us), divided totally into 1220 segments including lobes.
(5) E-type of lamina (Figures 2m, 3m) :Extremely large in size (10 cm in radius),
divided into at least 18 segments in this specimen.
The above classification is for the sake of

convenience and there are no strict morphological boundaries among the present
laminae; as a matter of fact, morphologicalvariation between A- and B-types, B- andC-types, C- and D-types and 0- and E-types
are gradual and continuous.
So far as our specimens are concerned, thefrequency of these types breaks down as follows: A-type (6%), B-type (14%), C-type(27%), D-type (40%) and E-type (3%). As
mentioned below, if these specimens belong

3m. Figure 3. Ginkgo manchuri~a (Yabe et Oishi) Meng et Chen. Line-drawings; all in natural size.
Figures 3a-m were drawn from Figures 2a-m respectively. 3n-p, additional drawings' 3n-o A-type ofleaves (NCP-8350). 3p, B-type of leaf (NCP-8369).
'
,

951.

A fossil population of Ginkgo leaves

to the same species, then the C- and D-typesof the leaves can be regarded as representativeof the species.
Both the right and left halves of a laminia
are normally almost bilaterally symmetricalin size, form and number of segments orlobes. The segments of lobes usually bearrounded or obtuse apex. In general, themore the number of segments or lobes, thehigher the lamina.
The vascular bundle appears to be single(?) in the petiole. It is then repeatedly
divided dichotomously into the lamina, ending at the distal margins of the ultimate seg-

79

ment or lobe, not converging at the apex, butsometimes coming close together there. No
thick veins along the margins of lamina arepresent as seen in extant or some fossil
Ginkgo leaves. The number of veins is 6-8(14-24 per cm in density) near the apex of theultimate segment or lobe.
2) Cuticular features (Figures 4-6): We
could observe cuticle in all the specimens
obtained. Cuticle is hypostomatic, but a few
occasional stomata are recognized on themarginal regions of segments or lobes on the
adaxial side. The upper (adaxial) cuticle ismostly 1-1.5/-lm thick and slightly thicker

80

Li-Ming Zhao et al.

than that of the lower (abaxial) one. The
upper cuticle consists mainly of vein-course
cells, 4-8 rows of elongated cells and interveinal ordinary cells, isodiametric andpolygonal in outline and irregularly arranged(Figurers 4d-e, Sa-A, 5b-d). Anticlinal andend walls of ordinary and vein-course cells
are mostly slightly sinuous. The sinuousity
is much clearer in the interveinal ordinary
cells than the vein-course cells. Each periclinal wall is markedly papillate at its centre(Figures 4d, Sa-A, 5b-c).
The lower (Abaxial) cuticle consists ofordinary cells and stomatal complexes. On
the vein course, cells are 5-10 rows of elongated cells. Their anticlinal walls are unevenly
thickened, but periclinal walls are smooth orslightly papillate (Figures 4a-b, 5e, 6b). The
outline of interveinal ordinary cells is usuallyindistinct under the light microscope but
much clearer under the phase-contrast andSEM microscopes (Figures 6d-e). On someordinary cells, a papilla 6-10 11m in diameteris present on each periclinal wall (Figures4b-c).
Stomatal complexes are scattered along theinterveinal areas, forming 2-3 rows. They
are irregularly orientated, but often longitudinal. Stomatal complex is haplocheilic andmonocyclic or occasionally amphicyclic.
Guard cells are sunken (Figures 4b, 4f, 4g, Sf).
Subsidiary cells are 5-8 in number, markedly
thickened near the aperture, and each with alarge-sized papilla overarched above theguard cells (Figures 4f, 4g, Sf). Figures 6d-f

show the inner view of stomata (SEM micrographs). Papilla is present on most of theordinary cells (Figure 6a) and Figure 6bshows inner view of the papillae on theordinary cells (SEM micrograph). Trichomeis not seen on both surfaces. Resin body isalso not observed between veins. Stem, twig
and reproductive organs are not known.
Discussion: In extant Ginkgo bi/oba Linnaeus, it is well known that the leaves arequite varied in size, form and mode of lamina
division, according to those on the longshoots, on the occasional off-shoots (outgrowths) and on the short shoots (e.g.
Seward, 1919, fig. 630; Sakisaka, 1958; Harris et al., 1974).
One of us (Kimura) cultivated several seedlings of Ginkgo bi/oba in the field garden
belonging to his institute under artificiallyworst conditions with regard to water supply
and amount of natural light for three years.
We found that the leaves sprouted every May,but did not grow in size. They remained
miniatures (about 2 cm in radius) until every
fall season. However, their cuticular features
did not differ from those of normally growing
leaves. If the detached leaves of extant
Ginkgo bi/oba were fossilized as impressions,then the above-mentioned miniatures ofGinkgo bi/oba leaves or those withmulti divided laminae on the occasional outgrowths would have been separated as
different species from the normal leaves.
The diagnostic features of fossil Ginkgo
and Baiera leaves depended only on their~
Figure 4. Ginkgo manchurica (Yabe et Oishi) Meng et Chen. Cuticular features (I). 4a, Lower(abaxial) cuticle, showing the vein (v) and interveinal (iv) areas. Surface view prepared from an E-type ofleaf (Figures 2m, 3m). The dark dots are stomata and small light dots are papillae in the interveinal area
(slide no. NCP-MI8265a). 4b, Lower (abaxial) cuticle, showing stomata (s) and papillae (p) in the interveinal area. Surface view. Enlarged partly from Figure 4a. 4c, Lower (abaxial) cuticle. Phase contrast
micrograph, showing vein (v) and interveinal areas. Surface view, prepared from one of the D-type leaves(NCP-8267) (slide no. NCP-MI8267a). 4d-e, Upper (adaxial) cuticle, showing vein (to the left in Figure 4d
and to the right in Figure 4e) and interveinal areas. Surface view. Figure 4d; phase contrast micrograph.
Figure 4e; normal light micrograph. Dark dots show the papillae prepared from a D-type leaf (shown in
Figures 2k, 3k; NCP-8359) (slide no. NCP-MI8359a). 4f-g, Stomatal complexes on the lower (abaxial)
cuticle, showing thickly cutinized and papillate subsidiary cells, seven (1-7) in Figure 4f and six (1-6) inFigure 4g. Inner view prepared from a B-type leaf (shown in Figures 2c, 3c; NCP-8345) (slide no. NCPMI8345a).

951.

v

400IJ m

L.....-_ _.....'

A fossil population of Ginkgo leaves

1...-_ _--',

81

200IJm82

Li-Ming Zhao et al.

40~m

[
200
~m

951.

A fossil population of Ginkgo leaves

mode of lamina division and number of veinsin the ultimate segment or lobe. We feel that
this much used distinction between the twogenera Ginkgo and Baiera is negligible.
The present Ginkgo leaves are all detachedand thickly massed and appressed on thesame bedding plane of the tuffaceous shale
bed. The leaves themselves can be peeled off
easily and their cuticle is seen to be wellpreserved. They differ from each other insize, form and mode of lamina division.
However, these features vary gradually andcontinuously and no morphological boundaries can be drawn amongst them. Also
their cuticular features are almost the same,
except for such minor points as sinuosity ofanticlinal walls of ordinary cells and developmental degree of papillae on the epidermal
cells. However, these features can vary fromlamina to lamina according to position.
Under the circumstances, we regard our fossil
Ginkgo leaves as belonging to the same fossil
population and showing only the variation ina single Ginkgo species.
About 70 Ginkgo (including Ginkgoites
and Baiera) 'species' or 'forms' (except
Sphenobaiera and Pseudotorellia) were reported from various localities and stratigraphicalhorizons of younger Mesozoic age in North
and Northeast China under the generic namesof Baiera, Ginkgo and Ginkgoites (Figure 7 ;Tables 1-2). It is very doubtful whether
such a large number of 'species' (or 'forms'
really existed.
Since the early 1900's pioneer studies of thefossil plants were made in western and northern Liaoning and southeast Songliao regions
mainly by Yokoyama (1906), Yabe (1908,1922), Yabe and Oishi (1933) and Oishi

83(1933). Table 2 shows the Jurassic-Lower
Cretaceous stratigraphical units bearing fossil
Ginkgo (including Baiera and Ginkgoites)
leaves (with asterisk) and Figure 7 shows the
localities of fossil Ginkgo (including Baiera
and Ginkgoites) leaves in Northeast China,
compiled by us.
1) In western Liaoning, fossil Ginkgo(including Baiera and Ginkgoites) leaves areabundant in the Beipiao, Haifanggou, Shahai
and Fuxin Formations. The fossil population of these leaves from the Fuxin Formation is similar to ours.
2) In north Lianing and southeast Songliao, fossil Ginkgo (including Baiera andGinkgoites) leaves are abundant in the Shahezi Formation. This fossil population is
also similar to ours.
3) In Daxinganling region (Inner Mongolia), fossil Ginkgo (including Baiera andGinkgoites) leaves are abundant in the Xingyuan Formation (=Jiufotang Formation;
the present one) and are rather abundant in
the Hongqi, Damoguaihe, Yuanbaoshan and
Yimin Formations. Of these, the fossil population of leaves from the Damoguaihe,
Yuanbaoshan and Yimin Formations is similar to ours.
4) The fossil population of Ginkgo (including Baiera and Ginkgoites) leaves in the
Dabao Formation (eastern Liaoning) is quite
similar to ours.
5) In eastern Jilin and eastern Heilongjiang, there are Jurassic and Lower
Cretaceous sediments mostly of non-marine
onglll. But the occurrence of fossil Ginkgo(including Baiera and Ginkgoites) leaves is so
sporadic that we are still ignorant about their
specific identification and the range of varia-~ Figure 5.
Ginkgo manchurica (Yabe et Oishi) Meng et Chen. Cuticular features (II). 5a, Wellpreserved upper (adaxial; A) and lower (abaxial; B) cuticles. Surface view. SEM-micrograph preparedfrom a lobe of the D-type leaf (shown in Figures 2k, 3k; NCP-8359) (stage no. NCP-8359SEM-2). 5b,
Upper (adaxial) cuticle, showing the vein (left side) and interveinal (right side) areas. Inner view.
SEM-micrograph prepared from the same lobe as Figure 5a-A (stage no. NCP-8359SEM-2). 5c, Vein area(vein course) of the upper (adaxial) cuticle, enlarged partly from the same lobe as Figure 5a-A. 5d,
Interveinal area of the upper (adaxial) cuticle, enlarged from the same lobe as Figure 5a-A. 5e, Lower(abaxial) cuticle, enlarged from Figure 5a-B. 5f, A stomatal complex, enlarged partly from Figure 5a-B.
Surface view, showing an aperture (ap) and papillate subsidiary cells (1-5).

84

Li-Ming Zhao et al.

20IJm

20IJm

951.

A fossil population of Ginkgo leaves

tion.
Under the circumstances, it is highly probable that the same population of Ginkgo
leaves was distributed at least in eastern Inner
Mongolia and eastern, western and northern
Liaoning in Jurassic and Early Cretaceoustime. The details are shown below.
Yabe and Oishi (l933) described 13'species' or 'forms' of Ginkgo (including
Baiera and Ginkgoites) from the following
localities (their current spellings are shown inparentheses) :Southwestern Jilin: Tao-chia-tun (Taojiatun) and Huo-shih-ling (Huoshiling).
Northeastern Liaoning: Pa-tao-ho (Badaohao; near Fuxin), Ta-tai-shan (Dataishan =Changtu; near Siping) and Sha-ho-tze (Shahezi = Changtu).
The above-mentioned localities belong to
the Shahai and Shahezi Formations (Table
2).
Central Liaoning: Wei-chia-pu-tzu (Weijiapuzi=Tianshifu; near Benxi), Nien-tzukou (Nianzigou) and Eru-tao-kou (Erdaogou).
The former two localities belong to the
Dabao Formation and the latter to the Sangeling Formation (Table 2).
The taxa described by Yabe and Oishi(1933) are: Ginkgoites ? crassinervis Yabe
(Taojiatun), G. digitata (Brongniart) var.
huttoni Seward (after Kryshtofovich, 1924;
without illustration) (Badaohao), G. cf. sibirica (Heer) [Shahezi, Taojiatun, Huoshiling,
Weijiapuzi, Nianzikou and Dataishan (after
Yokoyama, 1906)], Baiera cf. asadai Yabe et
Oishi (Erdaogou), Cf. B. concinna (Heer)(Shahezi), B. cf. gracilis Bunbury [Huoshiling, Nianzigou (after Yokoyama, 1906),
Taojiatun (after Yabe, 1908), Shahezi], B.

85

kidoi Yabe et Oishi, 1933 (= B. cf. angustiloba Yabe, 1908) (Taojiatun; after Yabe,
1908 H uoshiling), B. manchurica Yabe et
Oishi, 1933 [Shahezi, Dataishan (after Yokoyama, 1906), Taojiatun (after Yabe, 1908)
and H uoshilingJ, B. minima Yabe et Oishi,
1933 (Shahezi), B. orientalis Yabe et Oishi,
1933 (H uoshiling), B. cf. phillipsi N athorst(Huoshiling), B. sp. a (Huoshiling) and B.
sp. b (Shahezi).
We are of the opinion that so far as theexternal morphology of these Ginkgo (including Baiera and Ginkgoites) leaves illustrated
by Yabe and Oishi (l933) is concerned, mostof them are indistinguishable specificallyfrom the present Ginkgo leaves as shown
below:(I) Ginkgoites cf. sibirica (Heer); pI. 31,
fig. 8 (Shahezi)=our A-type, pI. 32, figs. 4-5,
8B (Taojiatun), fig. 6 (Huoshiling) and fig. 7(Weijiapuzi)=our B- and C-types.
(2) Cf. Baiera concinna (Heer); pI. 32,
fig. 6 (Shahezi) = our B-type.
(3) B. cf. gracilis Bunbury; pI. 32, fig.
13B (Shahezi; indeterminable), fig. 14 (Shahezi), fig. 15 (Huoshiling) and pI. 33, fig. 5(Shahezi) = our C-type.
(4) B. kidoi Yabe et Oishi, 1933; pI. 33,
fig. 3 (H uoshiling) = our D-type.
(5) B. manchurica Yabe et Oishi, 1933;
pI. 32, figs. 12, 13A and pI. 33, fig. 1 (Shahezi)=our C-type.
(6) B. minima Yabe et Oishi, 1933; pI.
32, fig. 11 (Shahezi) = our C-type.
(7) B. orientalis Yabe et Oishi, 1933; pI.
33, fig. 4 (H uoshiling) = our C-type.
(8) B. cf. phillipsi Nathorst; pI. 33, fig. 2(Huoshiling) = our C-type.
(9) B. sp. a (Yabe and Oishi, 1933); pI.
32, fig. 17 (Huoshiling) = our A-type, pI. 33,~
Figure 6. Ginkgo manchurica (Yabe et Oishi) Meng et Chen. Cuticular features (III). 6a, Lo.weT(abaxial) cuticle treated by KOH with much more time, showing the vein and interveinal areas. Inner view.
In the interveinal areas, many stomata (light ones) and papillae (small oval) are scattered. But it is difficult
to distinguish the stomatal complexes in such a low magnification. SEM-micrograph prepared from a
D-type leaf (shown in Figures 21, 31; NCP-8266) (stage no. NCP-8266SEM-4). 6b, Papillae, enlarged from
Figure 6a. Inner view. 6c-f, Stomatal complexes, prepared from an E-type leaf (shown in Figures 2m, 3m;
NCP-8265). Inner view. SEM-micrographs (stage no. NCP-8265SEM-l).

00

0>

_.i

I:'--<

~

j

;:::.
(}Q

~

I:)

D Markable citiesc~• JuraSsic-Early Cretaceous
Fossil plant sites yielding

I:)

:--..

leaves2 (compiledus).
Figure 7.by Localities
of known Ginkgo (including Baiera and Ginkgoites) leaves.

The numbers correspond to those shown in Table

951.
Table 1.

A fossil population of Ginkgo leaves

87

Baiera, Ginkgo and. Ginkgoites 'species' or 'forms' hitherto described or recorded by previous
authors from Jurassic-Early Cretaceous plant sites in Northeast China.
'; with description of external feature.
"; with description of external and cuticular features.
"'; without description and illustration (list only).
F.; formation, G.; group.

Baiera cf. angustiloba Heer : Yabe, 1908, Shahezi F.*~ asadai Yabe et Oishi, 1928 : Zhang et al., 1980, Guojiadian (Haifanggou) F.*
and Dabao F.*** : Zhang and Zheng, 1987, Haifanggou F.*** : Wang et al., 1989,
Haifanggou F_***

Baiera cf. asadai Yabe et Oishi, 1928 : Yabe and Oishi, 1933, Sangeling F.* and-----D-abao F.***
Baiera balejensis (Prynada) Zheng, 1980 : Zhang et al., 1980, Zhuanshanzi F.*
Baiera concinna (Heer) Kawasaki, 1925, Zhang et al., 1980, Guojiadian (Haifanggou)
F.* : Zhang and Zheng, 1987, Haifanggou ~*** : Wang et al., 1989, Haifanggou
F.***
Cf. Baiera concinna (Heer) Kawasaki, 1925 : Yabe and Oishi, 1933, Shahezi F.* : Oishi,
1933, Shahezi F.** : Sze ~ al., 1963, Shahezi F.** (Yabe and Oishi's and Oishi's
specimens, 1933) : Zhang and Zheng, 1987, Haifanggou F.***
Baiera czekanowskiana Heer, 1976 : Zhang and Zheng, 1987, Haifanggou.F.** : Wanget al., 1989, Haifanggou F.***
Baiera furcata (Lindley et Hutton) Braun, 1843 : Zhang et al., 1980, Damoguaihe F.*
and Beipiao F.* : Wang et al., 1985, Jubao F.*** and Jiufengshan F.*** : Zhang
and Zheng, 1987, Beipiao F.*** and Haifanggou F.*** : Wang et al., 1989, Beipiao
F.***
Baiera cf. furcata (Lindley et Hutton) Braun, 1843 : Wang et al., 1985, Jubao F.***
Baiera graciliS (Bean MS) Bunbury, 1851 : Yokoyama, 1906, Dabao F.* : Yabe, 1922,
Shahezi F.* : Sze et al., 1963, Fuxin F.***, Shahezi F.*** and Sangeling F.*(Yabe and Oishi's specimen, 1933) : Zhang et al., 1980, Guojiadian (Haifanggou)
F.* and Dabao F.*** : Wang et al., 1985, Dabao F.***, Beipiao F.***, Haifanggou
F.***, Wanbao F.*** and Manketouebo F.*** : Zhang and Zheng, 1987, Beipiao F.***and Haifanggou F.*** : Wang et al., 1989, Haifanggou F.***
Baiera cf. gracilis (Bean MS) Bunbury, 1851 : Yabe, 1908, Shahezi F.* : Yabe and Oishi,
1933, Shahezi F.* and Dabao F.*** : Oishi, 1933, Shahezi F.** : Sze et al., 1963,
Shahezi F.** (Yabe and Oishi's and Oishi's specimens, 1933) and Dabao F.*(Yokoyama's specimen, 1906) : Yang and Sun, 1982, Shahezi F.* and Yingcheng F.***: Wang et al., 1985, Hongqi F.*** and Beipiao F.*** : Zhang, 1987, Fuxin F.*** :Wang et al., 1989, Fuxin F.*** and Beipiao F.***~ cf. guilhaumati Zeil1er, 1903 : Sze, 1931, Fuxin F.***
Baiera kidoi Yabe et Oishi, 1933 : Yabe and Oishi, 1933, Shahezi F.* : Oishi, 1933,------Shahezi F.** : Sze et al., 1963, Shahezi F.** (Yabe and Oishi's and Oishi's
specimens, 1933) : Zhang et al., 1980, Shahezi F.*
Baiera manchurica Yabe et Oishi, 1933 : Yabe and Oishi, 1933, Shahezi F.* : Oishi,------1933 Shahezi F.** : Sze et al., 1963, Shahezi F.** (Yabe and Oishi's and Oishi's
specimens, 1933) : Zhang et al., 1980, Shahezi F.* (Yabe and Oishi's specimen,
1933) : Wang et al., 1985, Shahezi F.*** : Hao et al., 1986, Shahezi F.***
Baiera minima Yabe et Oishi, 1933 : Yabe and Oishi, 1933, Shahezi F.* : Oishi, 1933,------Shahezi P.** : Sze et al., 1963, Shahezi F.** (Yabe and Oishi's and Oishi's
specimen, 1933) : Zhang-et al., 1980, Shahezi F.* (Yabe and Oishi's specimen,
1933) and Anmin F.* : Hao et al., 1986, Fuxin F.***
Baiera orientalis Yabe et Oishi, 1933 : Yabe and Oishi, 1933, Shahezi F.*~ cf. phillipsi Nathorst, 1880 : Yabe and Oishi, 1933, Shahezi F.* : Oishi, 1933,
Shahezi F.**

88

Li-Ming Zhao et al.

Baiera pulchella Heer, 1876 : Toyoma and Oishi, 1935, Damoguaihe F.*
Oishi, 1933,
Baiera sp. a (Yabe et Oishi, 1933)
Yabe and Oishi, 1933, Shahezi F.*
Shahezi F.**
Oishi, 1933,
Yabe and Oishi, 1933, Shahezi F.*
Baiera sp. b (Yabe et Oishi, 1933)
Shahezi F.**
Baiera? sp. 3 (Sze et al., 1963) : Sze et al., 1963, Fuxin F.*~ sp. (Wang et al., 1985) : Wang et al., 1985, Wanbao F. *** and Changliangzi
F. ***
Ginkgo adiantoides (Unger) Heer, 1878 : Zhang et al., 1980, Yimin F.*
Ginkgo chilinensis (Lee) Zhang et al., 1980
Zhang et al., 1980, Shahezi F. * andDamoguaihe F.***
Ginkgo? crassinervis (Yabe et Oishi) Zhang et al., 1980 : Zhang et al., 1980, Shahezi
F.* (Yabe and Oishi's specimen, 1933)
Ginkgo curvata Chen et Meng, 1988 : Chen et al., 1988, Fuxin F.*
Ginkgo dayanensis Chang, 1980 : Zhang et al., 1980, Damoguaihe F.*
Ginkgo digitata (Brongniart) Heer, 1876 : Kryshtofovich, 1924, ? Fuxin F.*** : Szeet al., 1963, Damoguaihe F.* (Toyama and Oishi's specimens, 1935) and Fuxin
F:*** : Zhang et al., 1980, Damoguaihe F.*, Yimin F.*, Anmin F.***, Huolinhe
F.***, Dabao F.***, Beipiao F.*** and Shahezi F.*** : Wang et al., 1985, Didao
F.***, Anmin F.***, Jiufotang F.*** and Jiufengshan F.*** : Hao et al., 1986,
Changcai F.*** : Zhang and Zheng, 1987, Beipiao F.*** : Wang et al., 1989,
Beipiao F.***
Ginkgo cf. digitata (Brongniart) Heer, 1876 : Wang et al., 1989, Sunjiawan F.***
Ginkgo cf. hermelini (Nathorst) Hartz, 1896 : Sze, 1931, Beipiao F.*
Ginkgo huttoni (Sternberg) Heer, 1876 : Sze et al., 1963, Beipiao F.*** and Fuxin
F.*** : Zhang et al., 1980, Beipiao F.* and Shahezi F.* : Wang et al., 1985,
Xiaodonggou F.***, Anmin F.*** and Zhushan (Muling) F.*** : Hao et al., 1986,
Fuxin F.*** : Zhang and Zheng, 1987, Lanqi F.*** and Beipiao F.*** : Wang et al.,
1989, Fuxin F.***, Lanqi F.*** and Beipiao F.***
Ginkgo cf. huttoni (Sternberg) Heer, 1876 : Zhang, 1987, Fuxin F.*** and Shahai F.***
Ginkgo ingentiphylla Meng et Chen, 1988 : Chen et al., 1988, Fuxin F.**
Ginkgo lepida Heer, 1876 : Krasser, 1905, Shahezi F.* : Yokoyama, 1906, Dabao F.* :Yabe, 1908, Shahezi F.* : Zhang et al., 1980, Jiufotang F.* and Naizishan F.* :Wang et al., 1985, Dongshencun F.*** and Hongqi F.*** : Zhang and Zheng, 1987,
Haifanggou F.*** and Lanqi F.*** : Wang et al., 1989, Haifanggou F.*** andLanqi F.***
Ginkgo cf. lepida Heer, 1876 : Wang et al., 1989, Haifanggou F.***
Ginkgo manchurica (Yabe et Oishi) Meng et Chen, 1988 : Chen et al., 1988, Fuxin F.**and Xiaominganbei (Fuxin and Shahai)F. *- Ginkgo obrutschewi Seward, 1911 : Zhang and Zheng, 1987, Haifanggou F.*** : Wanget al., 1989, Haifanggou F.***
Ginkgo cf. obrutschewi Seward, 1911 : Zhang et al., 1980, Beipiao F.***, Haifanggou
F.*, Xiaoyingzi F.* and Houjiatun F.* : Zhang and Zheng, 1987, Haifanggou F.***
Wang et al., 1989, Haifanggou F.***
Ginkgo orientalis (Yabe et Oishi) Florin, 1936 : Zhang et al., Shahezi F.* (Yabe andOishi's specimen, 1933)
Ginkgo paradiantoides Samylina, 1967 : Chen et al., 1988, Shahai F.*
Ginkgo pluripartita (Schimper) Heer, 1881 : Chen et al., 1988, Xiaominganbei (Fuxin
and Shahai) F.*? Ginkgo pusilla Heer, 1876 : 'lIang et al., 1985, Peide F.***
Ginkgo schmidtiana Heer, 1876 : Krasser, 1905, Shahezi F.*
Ginkgo sibirica Heer, 1876 : Zhang et al., 1980, Damoguaihe F.*, Yimin F.*, Shahezi

951.

A fossil population of Ginkgo leaves

89

F.*, Yingcheng F.*** and Fuxin F.*** : Wang et al., 1985, Longzhaogou G.***,
Jubao F.*** and Anmin F.*** : Zhang and Zheng, 1987, Lanqi F.***, Haifanggou
F.*** and Beipiao F.*** : Chen et al., 1988, Fuxin F.* : Wang et al., 1989
Fuxin F.***, Shahai F.***, Haifanggou F.*** and Lanqi F.***-- -,Ginkgo truncata (Li) Chen et Meng, 1988 : Chen et al., 1988, Fuxin F.** andXiaominganbei (Fuxin) F.*-- -Ginkgo sp. (Yokoyama, 1906) : Yokoyama, 1906, Shahezi F.*
Ginkgo sp. (Wang et al., 1985) : Wang ~1 al., 1985, Ganhe F.***, Jiufengshan F.*"",
Hongqi F.*** and Manketouebo F.***
Ginkgo sp. 1 (Chen et al., 1988)
Chen et al., 1988, Fuxin F.*
Ginkgo sp. 2 (Chen et al., 1988)
Chen et al., 1988, Fuxin F.**
Ginkgoites chilinensis Lee, 1963
Sze et al., 1963, Shahezi F.** (Yabe and Oishi's
specimen, 1933)
Ginkgoites ? crassinervis Yabe et Oishi, 1933 : Yabe and Oishi, 1933, Shahezi F.* :Sze et al., 1963, Shahezi F.* (Yabe and Oishi's specimen, 1933) : Yang and Sun,
1982, Shahezi F.*
Ginkgoites digitata (Brongniart) Seward, 1919 : Toyama and Oishi, 1935, DamoguaiheF.*
Ginkgoites digitata (Brongniart) var. huttoni Seward: Yabe and Oishi, 1933,? Fuxin F.***
Ginkgoites ferganensis (Brick in Sixtel) Wang et al., 1985 : Wang et al., 1985,
Hongqi F.***
Ginkgoites fuxinensis Li, 1981 : Li, 1981, Haizhou (Fuxin) F.**
Ginkgoites cf. lepidus (Heer) Florin, 1936 : Sze et al., 1963, Beipiao F."** andDabao F.* (Yokoyama's specimen, 1906) : Wang et al., 1985, Beipiao F.***
Ginkgoites marginatus (Nathorst) Florin, 1936 : Wang et al., 1985, Hongqi F.***
Wang et al., 1989, Beipiao F.***
Ginkgoites obrutschewi Seward, 1919 : Wang et al., 1985, Xiaodonggou F.*"* :Hao et al., 1986, Huolinhe F.***
Ginkgoites orientalis (Yabe et Oishi) Florin, 1936 : Sze et al., 1963, Shahezi F.*(Yabe and Oishi's specimen, 1933) : Wang et al., 1985, Shahezi F.*** : Hao et al.,1986, Shahezi F.*** and Fuxin F.*** : Wang et al., 1989, Shahai F.***
Ginkgoites sibirica (Heer) Seward, 1919 or Ginkgoites sibiricus (Heer) Seward, 1919 :Toyama and Oishi, 1935, Damoguaihe F.* : Sze et al., 1963, Damoguaihe F.* (Toyama
and Oishi's specimens, 1935), Shahezi F.*** and Dabao F."** : Wang et al., 1985,
Hongqi F.***, Beipiao F.***, Anmin F.***, Jiufotang F.*** and Kuntouboluo
F.*** : Hao et al., 1986, Fuxin F.*** : Zhang, 1987, Shahai F.*** and Fuxin F.*"*
Ginkgoites cf. sibirica (Heer) Seward, 1919 or Ginkgoites cf. sibiricus (Heer) Seward
Yabe and Oishi, 1933, Shahezi F.* and Dabao F.* : Oishi, 1933, Shahezi F.** :Sze et al., 1963, Shahezi F.** (Yabe and Oishi's and Oishi's specimens, 1933)
and Dabao F.*** : Wang et al., 1985, Jiufotang F.***, Beipiao F.*** andand Taiyangling F.***
Ginkgoites cf. taeniatus (Braun) Harris, 1935 : Wang et al., 1985, Hongqi F.***
Ginkgoites truncatus Li, 1981 : Li, 1981, Haizhou (Fuxin) F.**
Ginkgoites wulungensis Li, 1981 : Li, 1981, Haizhou (Fuxin) F.**
Sze et al., 1963, Shahezi F.** (Yabe and Oishi's
Ginkgoites sp. 1 (Sze et al., 1963)
and Oishi's specimens, 1933)
Sze et al., 1963, Shahezi F.** (Yabe and Oishi's
Ginkgoites sp. 4 (Sze et al., 1963)
and Oishi's specimens,- 1933)
Ginkgoites sp. (Li, 1981) : Li, 1981, Haizhou (Fuxin) F.**
Ginkgoites spp. : Wang et al., 1985, Huoshiling F.***, Taiyangling F.***, Changliangzi
F.***, Xiaoling F.***, Houjiatun F.*"*, Jiuda F.*** and Changantun (Taiyangling)
F.*** : Hao et al., 1986, Sunjiawan F.***

Li-Ming Zhao et al.

90

Table 2.

Main Jurassic-Lower Cretaceous stratigraphical units in Northeast China (compiled by us) .
• : Occurrence of Ginkgo (including Baiera and Ginkgoites) leaves is known. F.; formation,
Gr.; gr.oup.
(I) The numbers correspond to those in Figure 7.
(2) The boundary between Upper Jurassic and Lower Cretaceous strata in this figure is still
controversial.

~
Stratigraphic
Units

I/)

::J
0

41
ttl
41
~
U

u

~

41
~
0

.... ...JWest
Liaoning
Sunjiawan F.
10·27 *

Daxinganling
North
Liaoning &East
Zhaowu- Liaoning
DongpoZhelimuSEXipo (W)dameng(E)
meng
Songliao
Quantou F.
Oayu F.

Banlashan F. Oenglouku F.
21·24·34? *10·27 *
Vingcheng F.
Fuxin F. *4·5·6·10·28
Shahai F. *4·5·6·10·27

21·23·24·25·26

Vimin F.*

Vimin F.*

VuanbaoHuolinhe F. shan F. *Oamoguai- Oamoguai- 7·8·9·28 *13·22
Shahezi F.*
he F.
* he F.
21·23·24·25·26
29·30·31
14·15·16 *Jiufotang F.
4·5·6·10 *
2nd
~

41

VixianQ. F. *
Q. 5·6·10

:J

1st

?

Huoshiling F.
21*------

Viliekede F. Ganhe F.
14·15 *

Shangkuli F.

?

Murui F.

- - - - -Jiufengshan F.*
14·15·16

0u;
I/)

~

....,
::J

=0

1J

~

Lanqi F. *Xingyuan F.
13·22 *?

Xiaoling F.*

Baoshi F.

2·3

Manitu F.
13 *Longjiang
Xiaodonggou
Fujiawazi F. ManketoueF.
F.
*
2·3- - - - - - - - - ~c:.~. J3_!

-

Sangeling F.
Baicheng F.

41

Lishugou F.

Baiyingaolao
F.

Tuchengzi F.
U

Nieerku F.

*

Jubao F. *7·8·9*
Oabao F. *
2·3

4·5·6
Haifanggou F.
4·5·6·10 *Wanbao F. *7·8·9

Zhuanshanzi
F.
2·3 *

Beipiao F.*
4·5·6
Hongqi F. *

~

41

~
...J

Xinglonggou F.

Kuntouboluo F.
*

7·8·9

Changliangzi

F.

*
2·3

951.
Table 2.

A fossil population of Ginkgo leaves(continued)

East

Jilin

Jilin region

Hunjiang region
ShirenJiaohe Sanyuan
Liuhe
pu

Jilin
Ouantou F.
Denglouku
F.

*

11

Liyan F.

Helongxishan

Baojiatun F.

Dalazi F.
Dongshan F.
Ouanshui- Ouanshuicun F.
cun F.
Changcai F. Changcai F.
Hengtongshan F.

*

Shiren F.

?

Xiahuapidianzi F.

*

Jiuda F.
11*
Deren F.
11
I ? *Houshigou F.

Sankeyushu F.

Naizi Cha- Naizi- Wulin
shan ng- shan F.
I--an F. F.
F. 11
32

Anmin F.
11·23

Yanji

E-Heilongjian,Jixi, BOIi,
Shuanyashan &WandashanregionLongjing F.

Heiweizi F.

Jinjiatun F. Moshilazi F.

*

91

Linzitou F. Baodaqiao

F.

Yingzuilazi

F.

.Sidaogou F.

Dashatan
F.

33

MUting F.
12·17·18·20**

Xishanping
F.

Chengzihe F.
12·17·18·20?

Shihebei F.

*~
12·17·18·19·20

Chaoyantun F.

Guosong F. Lamenzi F. ChangbaiF.

I

~

Q;

~
0

Taiyangling
F.

*
11

Houjiatun
F.
1

*

do
C)
~

0

01

0
III

.c

N
01

c
0

...J

Banshidingzi F.

Xiaoyingzi
F.
1*

Peide F.

..:

Dongshengcun
F.
12

*

92

Li-Ming Zhao et al.

fig. 6 (H uoshiling) = our C-type, pI. 33, fig. 10(Huoshiling) = indeterminable.
(10) B. sp. b (Yabe and Oishi, 1933); pI.
35, fig. 3A (Shahezi) = our C-type.
In addition to the above, identification of
the following forms seem to be doubtful.
(II) Ginkgoites ? crassinervis Yabe et
Oishi, 1933; pI. 32, fig. 8A (Taojiatun).
(12) G. digitata (Brongniart) var. huttoni
Seward; Kryshtofovich, 1924 (Badaohao;
without illustration).
(13) Baiera cf. asadai Yabe et Oishi,
1928; pI. 32, figs. 9-10 (Erdaogou).
In the same year, Oishi (1933) described the
cuticles of the following Ginkgoites andBaiera 'species' from Northeast China, of
which the external forms of laminae were
described by Yabe and Oishi (1933): Ginkgoites cf. sibirica (Heer), Cf. Baiera concinna(Heer), B. cf. gracilis Bunbury, B. kidoi Yabeet Oishi, B. manchurica (Heer), B. cf. gracilis
Bunbury, B. kidoi Yabe et Oishi, B. manchurica Yabe et Oishi, B. minima Yabe et
Oishi, B. cf. phillipsi N athorst, B. sp. a and B.
sp. b. So far our careful reexamination of
Oishi's published work does not reveal any
specifically diagnostic features amongst thepreceding. Their cuticles are quite similar to
ours.
Subsequently, many studies were made by
Chinese workers on the fossil Ginkgo (including Baiera and Ginkgoites) leaves from
Northeast China.
In the monograph made by Sze et al.
(1963), they established Ginkgoites chilinensis
Lee (from Huoshiling) as a new species andincluded Baiera cf. phillipsi Nathorst (described by Yabe and Oishi, 1933 and Oishi,1933) in their G. chilinensis. As mentioned
above, Baiera cf. phil/ipsi corresponds to ourC-type of leaf. The works of Yabe and Oishi(1933) and Oishi (1933) were followed byother workers without any critical reexamination.
Zhang et al. (1980) listed 21 Ginkgo (including Baiera and Ginkgoites) 'species' withbrief descriptions and illustrations known

from the Mesozoic plant sites in Northeast
China. They were collected from different
localities and horizons, but all of them were
illustrated only for their external forms. In
their monograph, they established Ginkgo
dayanensis Chang from the Damoguaihe
Formation (Table 2) together with G. sibirica
Heer and G. digitata (Brongniart) Heer. We
are of the opinion that the former two aresimilar to our C-type and the latter to ourA-type leaves.
Li (1981) originally described Ginkgoites
truncatus, G. wulungensis, G. fuxinensis andG. sp. on the basis of Ginkgo leaves withpreserved cuticles from the Haizhou Formation (= Fuxin Formation; Table 2), western
Liaoning. His Ginkgoites truncatus and G.
fuxinensis are like our B-type and C-type,respectively. It is easy to find out that there
is no difference in the cuticular features
between Ginkgoites fuxinensis and the present leaves. We feel that Li's Ginkgoites
species might be included in the same fossil
population as ours.
From the Fuxin and Tiefa Sedimentary
Basins (Figure 7), western Liaoning, Chen etal. (1988) described 9 Ginkgo species including their two new species, G. curvata and G.
ingentiphylla, and such species as G. paradiantoides Samylina, G. pluripartita(Schimper) Heer, G. sibirica Heer, G. truncata Li, G. sp. I and G. sp. 2. We observethat the leaves regarded by them as Ginkgo
curvata appear to be deformed in preservation and correspond to our B- and C- types.
The leaves regarded by them as Ginkgo ingentiphylla are much bigger than the others
and are just like our E-type. Their Ginkgo
pluripartita is very close in form to our B-type
and their G. truncata and G. paradiantoides
are our A-type. So far as the publishedfeatures of their leaves (including theirGinkgo manchurica) are concerned, we are
inclined to regard all their leaves as belonging
to the same population as ours.
If the above observations are correct, thenthere would be far fewer Ginkgo 'species' than951.

A fossil population of Ginkgo leaves

hitherto presumed during Jurassic and Early
Cretaceous time in Northeast China andadjacent regions. In addition, the genericdifference between Ginkgo (or Ginkgoites)
and Baiera becomes obscure.
Other Ginkgo populations: Heer (1876)
originally described Ginkgo sibirica, G. lepida, G. schmidtiana, G. jiabellata and G.
pusilla from the Jurassic of Ust Balej, Irkutsk
Basin. Doludenko and Rasskazova (l972)
used the generic name Ginkgoites after
Seward (1919) and regarded the species ofHeer mentioned above as conspecific andincluded them in Ginkgoites sibirica. Heer
(l876) also established Ginkgo concinna fromthe Jurassic of Ust Balej. According to
Doludenko and Rasskazova (1972), Beer's
Ginkgo concinna is distinct together with
their Ginkgoites tapkensis and G. heeri
known from the Irkutsk Basin.
On the other hand, Doludenko andLebedev (1972) also regarded Heer's Ginkgo
lepida, G. schmidtiana, G. jiabellata and G.
pusilla as synonyms of Heer's G. sibirica andregarded Heer's Ginkgo concinna and theirGinkgoites tylensis Lebedev from southern
Priokhotie, and Ginkgoites vachrameevi
Doludenko et Lebedev and Ginkgoites sp.
known from the Bureja Basin as distinct.
It is highly probable that there is anotherpopulation of Ginkgo leaves (Ginkgo sibirica
population) in the Irkutsk Basin, although as
far as we know, the detailed limit of morphological and cuticular variations for Beer's
species has not been shown beyond the worksof Doludenko and Rasskazova (1972) and
Doludenko and Lebedev (1972).
In fact, Ginkgo leaves corresponding to ourA-type and E-type have not been found fromthe Irkutsk Basin. In addition, we are ignorant as to the occurrence of Ginkgo leaves in
the Irkutsk Basin.
We suggest that the identification of fossil
Ginkgo (including Baiera and Ginkgoites)
leaves should be made not only from theirexternal morphology and cuticular features,
but also from their occurrence. It is quite93

difficult to make a precise identification in thecases of sparse occurrence or when preserved
cuticles are lacking, though cuticular featuresof Ginkgo (or Baiera) leaves, if present, arenot so effective and in some cases cannot help
in specific or even generic identification.
In the case of crowded occurrences of fossil
Ginkgo (or Baiera) leaves, we can treat themas a single population as in the present study.
The same results were arrived at as follows:(1) Kimura et al. (1983) on the basis ofimpressions of various types of Ginkgo (orBaiera) leaves from the Carnian Momonoki
Formation, Southwest Japan; they were
thickly massed and appressed in occurrence.
The Ginkgo (or Baiera) populations known
from the Norian Nariwa Group, Southwest
Japan (Kimura et al" 1983) and from theUpper Triassic Daedong Supergroup, Korea(Kimura and Kim, 1985) are the same ornearly the same as the Momonoki population.
(2) Ohana and Kimura (1986) describeddiminutive Ginkgo leaves with preserved cuticle from the uppermost Cretaceous (Maastrichtian) Omichidani Formation in the Inner
Zone of Central Japan, as a single Ginkgo
population. (3) Horiuchi and Kimura(1986) described Ginkgo leaves with nearly
entire or shallowly lobed margins and withpreserved cuticle from the Palaeogene Noda
Group, as a single Ginkgo population. (4)Zhou and Zhang (1989) described a Ginkgo
population consisting of various Ginkgo
leaves with preserved cuticle from the Middle
Jurassic Yima Formation, Henan, China.
These populations are different from eachother, according to their geographical andstratigraphical positions. As a matter ofcourse, difference of populations corresponds
to difference of species.
Nomenclature: We here adopt the genericname Ginkgo according to Harris et al.
(1974), instead of Ginkgoites (Seward, 1919).
In external morphology, our C- and Dtypes of leaves are clearly representative of thepresent population, and are close to Ginkgoites cf. sibirica, Baiera cf. gracilis, B. kidoi,

Li-Ming Zhao et al.

94

B. manchurica, B. minima, B. orientalis, B. cf.
phillipsi, B. sp. a and B. sp. b described by
Yabe and Oishi (1933). As mentioned
before in this paper, we feel that they areconspecific.
Chen et at. (1988) revised Baiera manchurica Yabe et Oishi as Ginkgo manchurica(Yabe et Oishi) Meng et Chen (comb. nov.)and included Baiera minima Yabe et Oishi,
B. orientalis Yabe et Oishi, Ginkgoites fuxinensis Li and G. wulungensis Li in Ginkgo
manchurica as being synonymous. Under
the circumstances, we give the leaves of thepresent population the name Ginkgo manchurica (Yabe et Oishi) Meng et Chen.
Ginkgoites pingzhuangensis described by
Zhao and Tao (1991) as a new species on thebasis of the specimen (NCP-8265) which isthe same specimen as that shown in the present Figures 2m, 3m, should rightly be withdrawn.

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Zhang, W. and Zheng, S.L., 1987: Early Mesozoic
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963-967, pis. 1-2. (in Chinese with English
abstract)
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221, p. 113-133, pis. 1-8.

Geographical and stratigraphical names: Japan: Momonoki tJ5/;;t, Nariwa Rl<;:fJ, Noda
!fEB, Omichidani :::k:i:![:fr. Tetori
Korea: Daedong:::kJj§J
China: Anmin * It, Badaohao /\ :i:![;bl, Baicheng B:I$;, Baiyingaolao B ~ jl1lj~, Banlashan ~ tv:
Banshidingzi f1Z:O r, Baodaqiao ~:::kBaojiatun * lif., Baoshi ~:0, Beijing ~t g, Beipiao ~t W-. Benxi *~, Binxian ~~. Boli ~ flJ, Changan * *.
Changantun **lif., Changbai * B, Changcai *lIt, Changchun *~. Changliangzi *~r. Changtu ~~, Chaoyang Ji!JlWi, Chanyangtun Ji!JlWilif.. Chengzihe :l$;rfoJ, Chifeng
iJf;~. Dabao :::k~. Dalazi :::ktv:r. Damoguaihe :::kJIWmfoJ. Dashatan :::k1:P1t. Dataishan:::kil"W. Daxinganling :::kJ!:!*~, Dayu :::k~. Donglouku ~:J;1!I!. Deren :fj'g(t:. Didao 1i1li:i:![, Dongpo
JEi:, Dongshan W. Dongshengcun 1m t1. Erdaogou =. :i:![ ¥Ill, Fujiawazi .f.\!j%O~r. Fuxin ~~. Ganhe i:f"foJ, Gannan i:f"l¥i, Guojiadian ~~%U5. Guosong
to, Haifanggou W ¥Ill. Haizhou W1+1. Hegang ~ ~. Heihe ~ foJ. Heilongj ian ~ ~IT. Helong fIJ~. Henan foJl¥i. Hengtongshan ='J=JiW. Hongqi U1i!t. Houjiatun f~*lif..
Houshigou f~:O¥lll. Hulun Nur Pff1jjji'lill. Hunjiang
Huolinhe li1*foJ. Huoshiling:k:o~. Jiaohe jB'Z foJ. Jilin E 1*, Jinjiatun ~ * lif.. Jiuda!A.:::k. Jiufengshan:1L ~ W.
Jiufotang :1L -Ml,¥:, Jixi ~"@. Jubao g~. Xingkai (Khanka)J!:!~. Kuntouboluo:J:$ AA l2Ji:*I. Lamenzi tv:r,r. Lanqi ~1i!t. Liaoning ~~, Liaoyuan ~~. Lingyuan &'i'~, Lin-

*1&

w.

*

m,

m

*

*

*

m

wIT.

*

Li-Ming Zhao et al.

96

zitou 1*.y rot Lishugou ~WrPJ, Liuhe 11JOi'PJ, Liyan l!t!6, Longjing lffi.*, Longzhaogou
lffi.JRrPJ, Manitu ~Jt,I!±., Manketouebo i1fi\j%M~tf, Moshilazi If!fE1i.v:.y, Muling ~~,Murui * J'ffij, Naizishan fjJ'J .y JlJ, Nenjiang itt IT, Nianzigou ~ .y rPJ, Nieerku i\lli 11= J!!!,
Peide ~:fi&" Pingzhuang f J±, Quanshuicun 7'R. 7.K ji, Quantou M, Sangeling =: 1ll~,
Sankeyushu =:t\I!:!mW, Shahai l,pw, Shahezi 1,pi'PJ.y, Shangkuli 1:.J!!!:1J, Shenyang i'fi~,
Shihebei :OjPJ~t, Shiren:O A, Shuanyashan .'lX."~ JlJ, Shuiquan
Sidaogou ~ :i:!!lilF,
Siping ~f, Songhuajiang 10::ttIT, Songliao 10i!, Sunjiawan f**~, Taiyangling:;t:~~, Taoan W~, Taojiatun 1lfi!J*11!:, Tianshifu B3 ffiljf$, Tiefa ~~, Tongliao )ii!, Tuchengzi ±:!1tZ.y, Ulanhot ,~llilif.5~, Wanbao ;i;~, Wandashan 7'CJ!JlJ, Weijiapuzi ~*$iIl.y, Wulin ,~1*, Xiahuapidianzi ~~Jt1E.J.y, Xiangshan ~JlJ, Xiaodonggou 'J'}l[rPJ,
Xiaoling 'J' li'it, Xiaorninganbei 'J' ~ '1i. Jill/-, Xiaoyingzi~.y, Xilutian g§ it 7(, Xingyuan 1!i'~, Xipo g§~, Xishanping g§JlJ:J5¥, Yanji ~E, Yiliekedc fj3-7U%~, Virna ~,w,;, Yirnin fj3-~, Yingcheng ~:!$;, Yingzuilazi 1ft~1i.v:.y, Yixian ~**, Yuanbaoshan 5C~ JlJ, Zalainor tL Ii!{ M5
Zhaowudarneng aB,~ ~ M, Zhelirnurneng tr £M, Zhuanshanzi f!JJlJ.y, Zhushan l$JlJ

*
**,

+

*

m,

~~I5"(J)...t.$~.3. "7 - T$tHI!*~1I eXingyuan) ~.ii1 T 3 '/~ili*~: :; "" 7 *[1,1&,MiJ'I?S!lU[I,ilrLltIlv::'iJ'vr-c, {~:O-1 =J- 3 ?~Ii, =>--7 :/Y}l[$G'):/"" ~ yml:toJ: lf~l1it~11ft!Jo/.JffG')~lJ8"Jt,d~RX:~*"ea; IJ, C ~ v::', i'lJ-:JtlljIij1:.v::'~f:l:rH, -c.il:t±:l-t" Go C G')t.:
lfP'J~"i5"G')1!i'~lIiJ'l?, J;J1:.G')J:? tJ:.il::tk~~-t"{~E-1 =J- 3 ?~iJ~9BJ!~.n, it.:19U7'J-tJ:-t.nI?G') cuticle iJ~J: i*ff~.n -C\"G o C.nI?G')-1 =J- 3 ?'ili*~~, ID, 'i:llrG')53-~
G'):tk~tJ: ~n.lb-CB£{~vc@;tsiJ\ -tG').il::tk, :to J: lftr'iG') cuticle G')~-t"~lJiJ~JRJ--c'a;
G C c iJ' I?, C.n I? G') -1 =J- 3 ? 'iIi, i'lJ-001*ffv::'~ L, iJ'''JJRJ-:f.iv::.~-t" G t G') c'iOUtfJT

<

<

c·

~.nGo

fL c' t Ii, ~Jt8"Jv::' C.n I? G') -1 =J- 3 ? 'i~ A - E G') 5 "JG')~v::'nvr, iJ'''J-C 9='OO}l[~t$,c <v::. i!?¥ 1ii' :to J: If"i5"~"i5"iJ' I? §G1X: ~ .ntdt:J 70 :f.i1ijJ:G') -1 =J- ? 'i c G').f;tt:<:~~d,..td5*,~ffifG')-1 =J- ?'iG') i:f.iJ G')~li, ~n.lb-Cj,'~-c'a;G C c, it.:C.nI?G'):f.ili, :Jt~8"J
i t.:ffif~8"Jv::., -t.n.:t.n, 1!m~G')lll1*ff~%RX: L -C I. 't.: C c iJ~~ I? iJ' c tJ: ··:d.: i t.: Ginkgo~ c Baiera ~ c G') Fa' G') ~JHiff:tE L tJ: I.' t G') c ~ ;t I?.n G
fL c· t Ii, P'J~"i5"}l[$:to J: lfi!?¥1ii'g§$:Jt~G') 1:.$:; "" 7 * - ~$ SlIE*iJ' I?%J I?.n-C
I.' G -1 =J- =< ? 'illl1*ffG')ltilRX::f.i c L -C, Ginkgo manchurica (Yabe et Oishi) Meng et Chen
3

3

o

0

~~~Lt.:o~li~.*::tt~.y.*ji~~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 169, pp. 97-128, 16 Figs., April 30, 1993

952.

DIMORPHISM OF LATE CRETACEOUS FALSE-PUZOSIINEAMMONITES, YOKOYAMAOCERAS WRIGHT AND
MATSUMOTO, 1954 AND NEOPUZOSIAMATSUMOTO, 1954*
HARUYOSHI MAEDA
Department of Geology and Mineralogy, Faculty of Science,
Kyoto University, Kyoto, 606-01

Abstract. Yokoyamaoceras jimboi Matsumoto, previously assigned to the Kossmaticeratidae, and Neopuzosia ishikawai (limbo), N.japonica (Spath), and N. haboroensis
Matsumoto and Inoma, previously assigned to the Puzosiinae, all co-occurring in Santonian
to lower Campanian deposits of the Upper Cretaceous Yezo Group in Hokkaido, Japan, andSakhalin, Russia, are· reinvestigated on the basis of many well-preserved specimens.
Observations on the stratigraphic occurrence, morphological variation, and shell ontogeny
show these four nominal species to be dimorphs of but a single biospecies. The first twomorphospecies represent the macroconch <M), and the latter two microconch <m) for which
the earlier available name is Yokoyamaoceras ishikawai (limbo, 1894), which is referred to
the Kossmaticeratidae. This is suggested by the mode of ontogenetic shell growth, particularly of the umbilicus.
A dimorphic lineage can be traced back to certain upper Turonian-Coniacian Kossmaticeras <M) and Yokoyamaoceras <m) species. The superficial morphological resemblance between the species and Mesopuzosia spp. is mere homeomorphy.
Key words. Dimorphism, Cretaceous ammonites, Yokoyamaoceras, Neopuzosia, Kossmaticeratidae, Puzosiinae

tine (Nullo et at., 1980), and Antarctica(Olivero, 1984). Species of the two genera
were described from Hokkaido and Sakhalinchiefly by Matsumoto (1954b, 1955, 1956).
Their generic and specific diagnoses were,
however, based on a limited number of specimens rather than population samples, and theextent of intraspecific variation could not be
confidently determined from previous studies.
I recently obtained many well-preserved
specimens of Yokoyamaoceras and Neopuzosia from the Upper Cretaceous Yezo
Group in Hokkaido and Sakhalin (Figure 1).
They suggest that the two genera, and particularly four coexisting nominal species: Yokoyamaoceras jimboi Matsumoto, 1955,
Neopuzosia ishikawai (limbo, 1894), N.

Introduction

Two desmoceratacean genera, Yokoyamaoceras Wright and Matsumoto, 1954,previously assigned to Kossmaticeratidae,
and Neopuzosia Matsumoto, 1954, previously
assigned to Puzosiinae, are very similar inexternal shell morphology and surface ornament, differing chiefly in the presence oftubercles in the latter. Both genera are indigenous to the late Cretaceous Pacific Realm,
and they are reliably known only from Hokkaido, Sakhalin, Koryak-Kamchatka, France(Kennedy, 1986), California, and British
Columbia (Haggart and Ward, 1989), Argen'Received October 23, 1992; accepted January 25,
1993
97

98

~_+-

Haruyoshi Maeda__---'V__---'!..--t----146·N

Figure 1. Index map of Hokkaido and South
Sakhalin showing distribution of the Cretaceous
Yezo Group (compiled from Matsumoto, 19S4a;
Tanaka, 1963; Geological Survey of Hokkaido,
1980; Zakharov et al., 1984). Locality names areabbreviated as follows. NB: Naiba (= Naibuchi :47·21'N, 142·32'E); TS: Teshio (44·47'N, 142·02'
E); AB: Abeshinai (= Saku: 44·42'N, 142·02'E);
EM: Embetsu (44·38'N, 141·S9'E); HB: Haboro(44· 16'N, 141·S4'E); KT: Kotambetsu (44·11'N,
141·S9'E); TP: Tappu (=Obira: 44·0S'N, 141·S6'
E); BI: Bibai (43·22'N, 142·02'E); IK: Ikushumbetsu (=Mikasa: 43·1S'N, 142·04'E); MJ: Manji(141·S7'E, 43·07'N); YU: Yubari (43·04'N, 141·S7'
E); OY: Oyubari (= Shuparo: 43·08'N, 142·0S'E); HO: Hobetsu (42·S2'N, 142·08'E); UK: Urakawa (42·11'N, 142·45'E).

japonica (Spath, 1922), and N. haboroensis
Matsumoto and Inoma, 1972, ranging from
Santonian to lower Campanian, are synonyms. This is based on the following observation, which suggests a wide range of morphologic variation, as well as dimorphism:
1) Nominal species show coincident distribution both in time and space.
2) Several morphological characters, previously regarded as criteria for discriminationof species, show continuous variation in large
samples.
3) Extreme similarity of immature stages.

4) A pair of large and small forms can berecognized (Figures 2, 3).
Following the works of Callomon (1955)
and Makowski (1963), numerous examples ofdimorphism in ammonoids have been reported; in the Paleozoic (e.g., Davis et af., 1969),
and in Mesozoic ammonites, particularly
from Jurassic ones (Callomon, 1963, 1985;
Westermann, 1964; Lehmann, 1966; Pal[raman, 1966,1967, etc.). These works
contributed greatly to systematic andphylogenetic reconsideration of ammonite
relationships. Dimorphism is also recognized among Cretaceous ammonites (Kennedy and Wright, 1985), and several taxa havebeen studied from this viewpoint, e.g., scaphitid heteromorphs (Cobban, 1969;
Tanabe, 1977a, 1977b, etc.), acanthoceratids(Wright and Kennedy, 1984; Kennedy andWright, 1985), placenticeratids (Kennedy andWright, 1983), and hoplitoidines (Reyment,
1988). However, many Cretaceous taxa
remain to be investigated more precisely.
In this paper, I first attempt to reconstruct
the dimorphism of the kossmaticeratid Yokoyamaoceras and puzosiine Neopuzosia.
Second, I discuss their taxonomic position at
the family level, based on the mode of earlyshell ontogeny compared with that of otherdesmocerataceans.
Repository of specimens.- The following
abbreviations are used to indicate the repository of specimens mentioned in this paper,
UMUT: University Museum, University ofTokyo, Tokyo 113; GK: Geological
Department, Kyushu University, Fukuoka
812. Material utilized here is shown in
Table 1.

Diagnoses of Yokoyamaoceras
and Neopuzosia
Yokoyamaoceras.- The genus Yokoyamaoceras was first established by Wrightand Matsumoto (1954), with Holcodiscus
kotoi Jimbo, 1894, from the Coniacian of the
Tappu area, Hokkaido, Japan, as typespecies. They defined Yokoyamaoceras as952.

Dimorphism of false puzosiines99

double collar ribbiconcave constriction
. rib
'.

...

r 0 s t rum .......... .,&!C.~,,~
lappet ........... ..

·.. ·periodic
constriction-.............. ....... ·parabolic

line'"
ventrolateral tubercle(Macroconch)(Microconch)

Figure 2. Schematic illustration of macro- and microconchs recognized in Neopuzosia and Yokoyamaoceras. The macroconchs (M> consist of two morphospecies: "ishikawai" and "japonica.""haboroensis" and "jimboi" comprise the microconchs (m>. The macro conchs are usually three times
larger than the microconchs. Scale bars: 10 mm long.

one of the tuberculate dwarf groups in thefamily Kossmaticeratidae, and regarded it as
an offshoot of Kossmaticeras in subsequentpu blications (Matsumoto, 1955, 1981). Kennedy (1986) recently placed this genus in thesubfamily Puzosiinae.
Species of this genus generally have small,moderate coiling to evolute, somewhat compressed shells with dense, fine, flexuous prorsiradiate ribs that are truncated somewhat
obliquely by periodic constrictions, as Kossmaticeras and some puzossines. Alternate
or opposite tubercles appear on ribs at eachventrolateral shoulder in the adult or near
adult stage. These tubercles may disappear
in the last 60 -150 volution of body chamber.
Tuberculation is variable, both in strength00and density, but was said to be the mostdistinct character by which the genus can bediscriminated from puzosiines. Umbilicaltubercles are not prominent even though they
appear in some doubtful forms. The sutureline is of kossmaticeratid type, simple, with asymmetrically tripartite lateral lobe (L) andmore weakly retracted umbilical elements
than in puzosiines (Wright and Matsumoto,
1954; Matsumoto, 1955,1956, 1981, 1991).
Six species of Yokoyamaoceras have been
described from the middle Turonian toSantonian. In this paper, I treat them asmorphospecies, with double quotation marks.
The morphospecies are as follows with brief
diagnoses and stratigraphic ranges as revised
by Obata et al. (1978) and my recent observa-

100

Haruyoshi Maeda

952.
Table 1.

Dimorphism of false puzosiines101

Mate.rial utilized in this paper including UMUT. and GK. type specimens.
localIty names are as same as in Figure I.
Morphosecies" Yokoyamaoceras kotoi""Y. jimboi""Y. ornatum""Y. (?) mysticum""Y. minimum""Neopuzosia japonica""N. ishikawai""N. haboroensis"
Desmoceras (Pseudouhligella) japonicum
Mesopuzosia pacifica
M. yubarensis
Jimboiceras planulatiforme
H auericeras (Gardeniceras) angustum
Kossmaticeras theobaldianum paucicostatum
Anapachydiscus sp.

No. of specimens

Abbreviations ofLocalities

5

TP, KT, HB

25
4

NB, HB, KT, TP, IK, HB
AB, HBUK

5

TP, IK, MJ

24

NB,TS,EM,AB,KT,TP, UK

30

NB, TS, AB, HB, KT, TP, IK, UK
NB,TS,AB,HB,KT,TP
AB, TP, IK

8
20
20
5

KT, TP, MJ, OY, HB
YU, OY, IK

1058

NB, TS, IK, UK
TP,IK

5

AB

TP

tions.

"Yokoyamaoceras jimboi Matsumoto, 1955""Yokoyamaoceras kotoi (Jimbo, 1894)"

Holotype.-UMUT. MM7625 (Figures 4-9,4-10), from the Saushi-sanushibe Valley,
Hobetsu (HO).
Diagnosis. - Morphospecies possessing fine,weak ribbing and with a brief, irregular tuberculate stage. Ribs finer (numbering about
30-40 per half a whorl) and weaker thanthose of" Y. kotoi." The ventrolateral tu bercles alternate, distributed with an incidence of
one tubercle per 3-4 ribs. Tubercles appear
at diameter of 20 mm or less, and extend forabout one-third to half a volution of the adultbody chamber. Adult shells range from 25
to 35 mm diameter. Lateral lappets are present in every specimen with a complete aper-

Holotype.-UMUT. MM7517 (Figures 415,4-16), from the Obirashibe (= Obirashibets) River, Tappu (TP in Figure 1).
Diagnosis.-Morphospecies showing many
distinct ribs and dense, extensive tuberculation over one whorl. The holotype has
about 25 ribs per half a whorl, and 23 of themdevelop tubercles at the ventrolateral shoulder. Tuberculation appears at a shell diameter of 15 mm or less, and continues for morethan a whorl. The shell attains at least 35
mm diameter. The apertural features of the
fully grown adult stage are unknown.
Occurrence.-Coniacian.
+-Figure 3. Adult macroconchs (1-4, X 0.67) and adult microconchs (5-8, X 1.5). All the specimens
are here assigned to Yokoyamaoceras ishikawai (limbo, 1894). 1,3, "Neopuzosia ishikawai (limbo)" <M),
UMUT. MM18341, from the lower Santonian of the Kotambetsu River (Loc. KT-5040). 2, 4, "N. japonica(Spath)" <M), UMUT. MM18337, from the Santonian of the Naiba River, South Sakhalin. 5,6, anintermediate form between "N. haboroensis Matsumoto and Inoma" and" Yokoyamaoceras jimboi Matsumoto, 1955" <m), UMUT. MM18385, from the uppermost lower Santonian of the Kotambetsu River (Loc.
KT-5302). 7,8," Yokoyamaoceras jimboi Matsumoto" <m), UMUT. MM18357, from the lower Santonianof the Obirashibe River (Loc. TP-022).

102

Haruyoshi Maeda

952.

Dimorphism of false puzosiines

ture (Figure 5).
Occurrence.- Lower Santonian except for
the uppermost part.
"Yokoyamaoceras ornatum Matsumoto, 1956"

Holotype.-GK. H521O, from the Abeshinai River (AB).
Diagnosis.- Morphospecies characterizedby much influated shell with subquadrate
cross section, and very coarse, flexuous distinct ribbing of about 20 ribs per half a whorl.
Occurrence.-Coniacian. Except fordifference in cross section, general features of"Y. ornatum" are quite similar to those ofcontemporaneous Y. kotoi, and there aresome intermediate forms between the twomorpho species.
"Yokoyamaoceras (?) mysticum Matsumoto,
1955"

Holotype.-GK. H3379, from Loc. U-147p,
Urakawa (UK).
Diagnosis.-Morphospecies with tubercles
at both ventrolateral shoulder and umbilicalmargin.
Occurrence.-Only the holotype is known
from the Santonian (Matsumoto, 1955, p.
156-157), and the identity of this species isstill uncertain.
"Yokoyamaoceras minimum Matsumoto, 1956"

Holotype.-GK. H4204, found in a pebble
in the Ikushumbetsu River (lK).
Diagnosis.- Morphospecies characterized
by very small shell size and irregular but
prominent tuberculation. The adult is 20
mm or less in shell diameter, with a pair oflappets.

103

Occurrence.-Upper Turonian to Coniacian.
"Yokoyamaoceras yokoii Matsumoto, 1991"
Holotype.- TMNH (= Toyohashi Museumof Natural History) 501, from Mj-k Member,Satonosawa Valley, Tappu (TP).
Diagnosis.- Morphospecies characterized
by small shell with subtrigonally shoulderedventer, in which asymmetrically approximated two rows of tubercles appear.
Occurrence.-Middle Turonian.
Neopuzosia.-Neopuzosia is characterized
by a moderate-sized, compressed (BjH:
about 0.6-0.7 in adult), somewhat widely
umbilicate Puzosia-like shell with slightly
convex sides, perpendicular umbilical wall,
6-8 periodic constrictions per whorl, and fine,
flexuous prorsiradiate ribs. The constrictions, associated with double collar ribs, areprojected forwards on periphery, and can beobserved in almost all ontogenetic stages. A
constriction usally truncates 2-4 ribs justbehind it. Distant, rather straight ribs may
appear suddenly on the living chamber ofsome adult shells, but flexuous ribbing still
remains in others. In contrast to other true
puzosiines in Hokkaido and Sakhalin, theshell never attains a large size, and the maximum diameter is usually less than 150 mill.
The type-species of Neopuzosia is based ona type specimen from the Urakawa area,
Hokkaido. It was first described as Desmoceras gaudama Forbes, 1846 by Yokoyama (1890). Subsequently, Spath (1922)
established the genus Kitchinites, based on~ Figure 4. Microconchs X 1.2. All the specimens, except 15, 16, are here assigned to Yokoyamaoceras
ishikawai (limbo, 1894). Arrows show successive change of forms. 1,2, an intermediate form between"Yokoyamaoceras jimboi Matsumoto" and "Neopuzosia haboroensis Matsumoto and Inoma," UMUT.
MM18351, from. the uppermost lower Santonian of the Kotambetsu River (Loc. KT-5302). 3,4, "N.
haboroensis Matsumoto and Inoma," holotype, GK. H5654, from the upper lower Santonian of theMiginosawa Valley (Loc. IA-I 564), Haboro. 5-12," Yokoyamaoceras jimboi Matsumoto." 5,6, GK.
H5201, exact locality uncertain, Ikushumbetsu. 7,8, a depressed form, UMUT. MM7623a, Kikumenzawa
Valley, Ikushumbetsu. The specimen was originally described as Yokoyamaoceras kotoi by Matsumoto(1955). 9,10, holotype, UMUT. MM7625a, from the Santonian of the Saushi-sanushibe Valley, Hobetsu.
11,12, UMUT. MM18359, from the lower Santonian of the Kumaoizawa Valley (Loc. IK-300), Ikushumbetsuo 13,14, an intermediate form between" Y. jimboi" and" Y. kotoi," UMUT. MM18350, from thelowermost lower Santonian of the Kotambetsu Valley (Loc. KT-5606). 15,16, Yokoyamaoceras kotoi(limbo), holotype, UMUT. MM7517, from the Coniacian of the Obirashibe River, Tappu.

Haruyoshi Maeda

104

omm

D

" sp.

c
.....
C

~

;:,...

MM18355

HB

MM18354

HB

MM7517<E> TP

MM111348

KT
TP

MM7823a

IK

MM7823b

IK

MM 18350.-

MM7824a

NB

MM7624b

NB:~

MM18358

IK'"....<::!

MM18359

IK

MM 18349

IK

C
..c:,

E:

<u

'-'

c

<::!

E:
<::!

;:...

c
~
c
;:,...

MM7825

KT

MM18345

KT

MM18357
MM18347

TP
KT

MM18351

KT

MM 18362KT

c:).2<::!

GK.H5854

....

<E>

HB

~

MM18353

AB

MM18338

EM

MM18341

KT

~

(M)

15

+

20

broken part.--s:::

30

D

40

uberculate stage
simple aperture

.~

~

(m)

KT

GK.H5855.~.~.~;:-;

HB

-s:::

c:;,

.

<E> HO

MM 18344

MM18352~:...

<B>: Holotype

lappeted aperturebiconcave constriction
aperture broken
tuberculate stage
tuberculate stage extending to inner whorl

Figure 5. Schematic diagram showing the diagnostic features and size in the adult stage for the fourmorpho species discussed plus" Yokoyamaoceras kotoi (limbo, 1894)." The latter species can be considered as an ancestral form of the microconchs.

952.

Dimorphism of false puzosiines

Holcodiscus pondicherryanus Kossmat, 1897
from India. He placed Yokoyama's specimen within his genus, and gave it a new
specific name: Kitchinites japonicus. Matsumoto (1954b) proposed Neopuzosia as anindependent puzosiid genus in which
flexuous ribbing still remains even in theadult stage compared with Kitchinites (Kitchinites), and this division has been followed
by several studies (Matsumoto, 1988, etc.).
On the other hand, Wright (1957) ranked
Neopuzosia as a subgenus of Kitchinites in thesubfamily Puzosiinae. Neopuzosia is currently regarded as transient from Mesopuzosiaof the Turonian-Coniacian to Kitch in ites
(Kitchinites) of the Campanian-Maastrichtian(Matsumoto, 1954b, 1981, 1988; Matsumoto
et al., 1972). Matsumoto (1954b, p.95)
interpreted" N. ishikawai" as a descendant of
Mesopuzosia yubarensis (limbo), and "N.
japonica" as derived from M. pacifica Matsumoto. Three Neopuzosia species have been
described from Hokkaido and Sakhalin, but I
treat them provisionally as morphospecies
with double quotation marks in this paper.
Details are as follows.
"Neopuzosia japonica (Spath, 1922)"

Lectotype.-Yokoyama (1890, pI. 19, figs.
5a, b) from the Urakawa area (UK; exact
locality unknown).
Diagnosis.-Morphospecies showing coarser and stronger ribbing than" N. ishikawai."
The rib density is 25-30 per half a whorl at
100 mm shell diameter. Some adult shells
may exhibit straight and somewhat stronger
prorsiradiate ribbing, but others still possess
flexuous ribbing as in the juvenile.
Occurrence.-Santonian to lower Campanian.
"Neopuzosia ishikawai (Jimbo, 1894)"

Lectotype.-UMUT. MM7507 (Figures 65, 6-6), from the Santonian of the ObirashibeRiver, Tappu (TP).
Diagnosis.-Morphospecies possessing fine,dense sigmoid ribbing (numbering about 3040 per half a whorl at 100 mm shell diameter).
The ribs are prorsiradiate, strongly projected105

forwards on periphery even in adult shells.
Occurrence.-Santonian to lower Campanian.
"Neopuzosia hahoroensis Matsumoto and Inoma,
1972"

Holotype.-GK. H5654 (Figures 4-3, 4-4)
from the Miginosawa Valley, Haboro (HB).
Diagnosis. - Morphospecies characterized
by a small, lappeted shell. The adult shellranges from 25 to 40 mm diameter.
Occurrence.-Uppermost lower Santonian
to lower Campanian.
I will concentrate my discussion mainly on
the four Santonian to lower Campanianmorphospecies: "N. ishikawai," "N. japonica," "N. haboroensis," and" Yokoyamaoceras jimboi" for they are abundant comparedwith other morphospecies from different horizons. For convenience, I adopt their specificepithets for the expression of these morphospecies, e.g., "ishikawai" for" Neopuzosia
ishikawai (limbo, 1894)" etc.
Mode of occurrence
The species of "Yokoyamaoceras" and"Neopuzosia" discussed here are found in thecentral slope basin facies and the western
shelf facies of the Yezo Group in Hokkaido(Matsumoto, 1942, 1954a; Okada, 1979,
1983). Figure 7 shows stratigraphic occurrence of Yokoyamaoceras and Neopuzosia in
the Turonian- Santonian sequence along the
Kotambetsu River, northwest Hokkaido.
Strict facies-dependent distributions are notseen.
The species of "Yokoyamaoceras" and"Neopuzosia" frequently co-occur in the sameplace (Table 2). Well-preserved examples
are usually obtained sporadically from calcareous concretions embedded in the muddy
deposits, in association with other ammonites, inoceramids, other benthic fossils, andnumerous plant remains. Most are found ina good state of preservation, except for thepartial collapse of the living chamber during
diagenesis (Maeda, 1987). Inner whorls with

106

Haruyoshi Ma eda

952.

Dimorphism of false puzosiines

protoconchs are usually well preserved, andamenable to ontogenetic study. A few specimens, larger than 100 mm in diameter, show
the slight dissolution of their inner whorls.
They are never concentrated into shell beds.
Adult features and variation
If we take no account of the barrier of
traditional classification, the coexisting fourmorphospecies can be easily grouped into two
categories: "macroconch" <M) and "microconch" <m) proposed by Callomon (1955,
1963) for dimorphic ammonites, in which
juvenile shell morphologies closely resemble
each other. The prerequisites for recognizing dimorphism (e.g., Callomon, 1963, 1969,
198 I ; Makowski, 1963 etc.) are discussed
below; Figure 2 illustrates the two forms
schematically. The macroconch consists of"ishikawai" (Figures 3- I, 3-3) and "japonica" (Figures 3-2, 3-4). "Haboroensis" (Figures 4-3,4-4) and "jimboi" (Figures 3-7, 3-8)
represent the microconchs. Both forms arenearly identical in the immature stage, but
exhibit remarkable dimorphic features in theadult; there are no intermediates. Macroconchs are three times (or maximum fourtimes) larger than the microconchs (Figure 5).
Macroconchs.-" Ishikawai" was distinguished from contemporaneous "japonica"byprevious authors (e.g., Matsumoto, I 954b) by
its weaker and finer ribbing. Plots of rib
density against shell diameter (Figure 8) showthat the ribs tend to coarsen as the shellreaches maturity. Rib density converges
finally to around 30 per half a whorl in both
macroconchs. Both macroconchs show awide continuous variation (Figure 6), and theranges of variation of the two "species" overlap. Besides ribbing, they share many com-107

mon morphologic characteristics. The
differences between the type specimen of"ishikawai" <M) and "japonica" <M) are,therefore, considered as within the range ofintraspecific variation, and I summarize both
macroconchs together as "ishikawai" after
this.
Fully grown shells comprise 8-9 whorls,
and generally reach 90- 140 mm diameter.
The body chamber extends for 240" to 280".
The general coiling pattern is slightly
modified towards the aperture of the adult, asa result of decrease of whorl expansion rate
(Figures 3-1, 3-2). At the same time, thewhorl becomes slightly compressed, and ornament may strengthen somewhat particularlyon the periphery. These features indicate
maturity, though approximation of the last
few septa and simplification of the last suture(Kennedy and Cobban, 1976) are not clearlyobserved. The aperture never develops peristomal modifications. The aperture line is a
simple prorsiradiate or concave pattern,which gently projects forwards on the periphery (Figure 2).
Microconchs. -" Jimboi" and "haboroensis" are nearly identical to macroconchs up to
15 or 20 mm shell diameter (Figure 9). Ribdensity of microconchs is the same as that ofmacroconchs (Figure 8). Fully grown shells
possess 12-14 septa per whorl, and the bodychamber length ranges from 240" to 280", as inmacroconchs. However, microconchs showquite different adult features from the macroconchs in shell size, mode of tuberculation, and peristomal modifications.
Fully grown shells, comprising 6- 7 whorls,
are no more than 25-40 mm diameter. In
later growth stages, alternate ventrolateral
tubercles appear in "jimboi." They appear
on the first half of adult body chamber, and~Figure 6. Macroconchs X 1.0. 1-3, "Neopuzosia japonica (Spath)," UMUT. MM18331, from theupper Santonian of the Omoshirubeshibetsu Valley (Loc. EM-404), Embetsu. 4, "N. japonica (Spath),"UMUT. MM6638, from the uppermost Santonian or lowermost Campanian of the Tannosawa Valley (Loc.
T-472), Abeshinai. 5,6, "N. ishikawai (Jimbo)," lectotype, UMUT. MM7507, from the Santonian of the
Obirashibe River, Tappu. 7, UMUT. MM18336; 8, UMUT. MM18386, "N. ishikawai (Jimbo)," from thelower Santonian of the Kaminosawa Valley (Loc. KT-5200), Kotambetsu.

Haruyoshi Maeda

108

Stratigraphy

Loc .
·5062
·5063

(Macro conch)(Microconch)

•

.5065
-5031

• 5033

-5034

.5035
'5036
.5037
.5211
.5039
• 5040
.5041

.5042

0

0

"N.

.0

E

....,

·5043
.5302
'5044
• 52320"N. japonica

!\:S

~

0

• 109

!\:S
~

• 5051• 022• 5056• 5057

..c:

CI)

0

• 5604• 5605• 5606• 5608• 5609• 5612"• 038• 038'• 5613• 5414

-5415
'5416

Y. minimum

Kossmaticeras sp.

• 5417·5419• 5420·5421• 5422SANDSTONE

ALT. OF SS. & MS.
• 5423• 5424

BIOTURBATEDMUDDY SS.

1100m

-

LEGENDBIOTURBATEDCOARSE MS.

~

PLANOLITES MS.

b~

SLUMP DEPOSITS=ACIDIC TUFF

Figure 7. Historical changes of macroconch "ishikawai" and "japonica," and microconch "jimboi"
and " haboroensis" lineage in the Turonian-Santonian sequence along the Kotambetsu River (KT). Solidcircle: occurrence of microconchs <m) ; Open circle: occurrence of macroconchs <M) . SS.: sandstone.
MS. : mudstone.

952.

Dimorphism of false puzosiines

109

Table 2. Fossil assemblages showing coexistence of macro- and microconchs at the same places. I, an
example housed in a calcareous nodule of 300 mm diameter, Ikushumbetsu (IK). 2, anotherexample housed in a calcareous nodule of 200 mm diameter, Teshio (TS). VA: very
abundant; A: abundant.
(I)

Locality: IK-300 (Lower Santonian)Species

Individuals<AMMONITES)

Neophylloceras ramosum (Meek)
Gaudryceras tenuiliratum Yabe
Tetragonites glabrus (limbo)
Damesites damesi (limbo)

I"Neopuzosia ishikawai (limbo)" <M)"Yokoyamaoceras jimboi Matsumoto" <m)Hauericeras (Gardeniceras) angustum Yabe
Polyptychoceras pseudogaultinum (Yokoyama)

2+
12+
3
4

3

5+

<BIVALVES)

Inoceramus (Inoceramus) amakusensis Nagao and Matsumoto
Inoceramus (Sphenoceramus) naumanni Yokoyama

3

60+<GASTROPOD)

Serrifusus sachalinensis (Nagao)<PLANT REMAINS)

VA

(2)

Locality: TS-8007 (Upper Santonian)Species

Individuals<AMMONITES)
Gaudryceras tenuiliratum Yabe
Damesites sugata (Forbes)"Neopuzosia ishikawai (limbo)" <M) and "N. japonica (Spath)" <M)"Neopuzosia haboroensis Matsumoto and Inoma" <M)
immature shells of "Neopuzosia sp." <M) or <m)Hauericeras (Gardeniceras) angustum Yabe
Polyptychoceras sp.

22<BIVALVES)
Inoceramus (Sphenoceramus) naumanni Yokoyama
immature bivalves gen. indet.

60+

3
2

9+
1+

3

<GASTROPODS)"Avellana" problematica Nagao
Tibia japonica (Nagao)<PLANT REMAINS)AHaruyoshi Maeda

110

No. of ribs/h.w.
60

o
~L
~

.... .... ·0

number of ribs
per half a whorl

50

v

o
0

40
®

®o

q,

®GO
®
®

®

D

00
®

®

®

30

0

1

10

<£>
1

50

0

·Y. jimboi"

®

~
20

v

•N. ishikawai"•N. japonica"•N. habroensis·0

ffi (1)0.• ®

<£> Lectotype
I

I

100

I

200

D
mm

Figure 8. Plots of rib density in relation to shell diameter (D) for medium-to-large specimens of fourmorpho species. The mode of ribbing shows a wide extent of variation even at the same shell size. It
varies continuously, and discrimination of "ishikawai" from "japonica" is impossible. Such variation inri bbing is regarded as intraspecific.

extend over 120°-240° of whorl. The tuberculation is absent in "haboroensis," however,
it is unsuitable for taxonomic discrimination
as stated below.
The tuberculation of the microconchs clearly changes chronologically both in densityand extent of appearance. Though there is acertain degree of variation even in the coexisting specimens, microconchs tend to lose theirtuberculation at progressively higher stratigraphic levels. Typical examples of microconchs from different stratigraphic levels
are shown in Figures 4 and 7. The lowermost Santonian forms (e.g., Figures 4-13, 4-14) are characterized by dense tuberculationon the adult body chamber. The specimen

ill ustrated possesses 18 ventrolateral tu bercles, appearing over about half a volution(170°) of the body chamber. It shows somewhat intermediate features between
Coniacian "kotoi" (Figures 4-15, 4-16) and"jimboi." Irregularly tuberculate specimens,
typical "jimboi," are dominant in the mainpart of the lower Santonian. The holotype(Figures 4-9, 4-10) has 8 tubercles extending
over one-third of a volution of the bodychamber. In the upper Santonian and lower
Campanian, microconchs are represented by
nontuberculate "haboroensis" (Figures 4-3,4-4), and tuberculate forms are unknown at
this level.
Recently, I collected several specimens

952.

DimOlphism 0/ /alse puzosiin es

III

Figure 9. Immature shell s of mac ro- and microconchs x 1.0. All the specimens are here assign ed to
Yokoyamaoceras ishikawai (limbo , 1894). 1- 3," Y. jimboi Matsum oto " ( m>, in a calcareous concretionfrom the lower Santonian of the Kotambetsu Valley (Loc. KT-5056). 1, UMUT. MM1 8346 ; 2, 3,
UMUT. MM 18344. 4- 12, coexistent" Neopuzosia species" in a calcareous concretion from the upper
Sa ntonian of the Kokuneppu Valley (Loc. TS-8007), Teshio . 4, UMUT. MM18380 ; 5, UMUT.
MM18381 , immature shells of "N. ishikawai (limbo)" ( M> or " N. haboroensis Matsumoto and Inoma"( m>. 6, 7, " N. ishikawai (limbo) " ( M>, UMUT. MM183 82. 8- 12, ontogenetic development of " N.
haboroensis Matsumoto and Inoma" ( m>, UMUT. MM18387.

intermediate between tuberculate and successive nontuberculate forms (e.g., Figures 4- 11 ,4- 12 via Figures 4- 5, 4-6 to Figures 4-1 , 4-2).
One of them, from the uppermost lower
Santonian (Figures 4-1 , 4- 2), is faintly tuberculate, with only 6 indistinct ventrolateral
tubercles on a limited portion (ca. 70· in
whorl volution) of the adult living chamber.
It exhibits features clearly transient between
those "jimboi" (Figures 4-5 , 4 - 6) and"haboroensis" (Figures 4-3, 4- 4). The two

microconch morphospecies are thus linked
together into a single lineage by such intermediate form s.
In the full -grown mature stage of ''jimboi''<m) and "haboroensis" <m), a characteristic
biconcave constriction develops at the end of
the body chamber, from which a pair oflateral lappets and a rostrum project ado rally(Figures 2,3-5-3-8,4). The biconcave constriction is adjacent to the last periodic constriction. It never appears at any otherHaruyoshi Maeda

112

growth stage, and is easily differentiated fromthe normal periodic constrictions by thedifference in curvature. The lappets are 2-5mm long, and have a simple tapering shape.
The distinctive apertural modifications are
regarded as diagnostic of maturity in microconchs (Kennedy and Cobban, 1976;
Callomon, 1981). "Jimboi" (m> and"haboroensis" (m> closely resemble eachother in size and apertural features at maturity (Figures 4, 5).
Besides these features, parabolic lines(= "demarcation lines" by Matsumoto et al.,
1972; Matsumoto, 1991; Figure 2) appear
repeatedly just in front of the periodic constriction in several specimens of microconch.
However, the appearance of parabolic lines
should be excluded from t~e criteria for recognition of microconchs, because they are notpeculiar to microconchs but are also fairly
common in macroconchs (Figure 3-1), andeven in huge puzosiines.
Ontogenetic shell growth
To confirm the dimorphic relationshipamong the four morphospecies discussed
above, I have paid much attention to the earlyshell morphology, which may possibly reflect
phylogeny as well as other diagnostic features. For the analysis of ontogenetic shell
growth, I prepared cross sections of more than30 specimens of the four morpho species andother desmocerataceans (Table 1) by polishing the shells along the center ::>f the
protoconch. The following parameters(abbreviations used in the text, figures, andtables are shown in parentheses) were measured at intervals of half a whorl (180') on
each sectioned specimen using a digital
micrometer (accuracy± 1 ,urn) attached to an
overhead profile projector (V-16D, Nikon);whorl radius (R), whorl breadth (B) andumbilical radius (C). The whorl volution isexpressed by 7l (= 180') scale from the centerof protoconch.
Figure 10 shows the median cross sectionsof early growth stage of "ishikawai" (M>,"haboroensis" (m>, and "jimboi" (m>compared with three other desmocerataceans.
The macroconchs are treated as "ishikawai."
The protoconchs of dimorphs are nearly identical, having an ellipsoid shape, 0.5 mm wide
and 0.3 mm diameter. They are enclosed by
the extremely depressed first whorl. In allmorpho species, the siphuncular tube lies atfirst in the center of the camera just as in
Nautilus. Thereafter, it moves towards theventer, and finally closes to the ventral shellwall at the 3-4 7l stage. The whorl isdepressed at first, but whorl height grows
more rapidly than breadth as the shell grows.
The umbilical shoulder appears at the 4-5 7lstage.
The profiles of" haboroensis" (m> (Figures
10-1), "ishikawai" (M> (Figures 10-2), and"jimboi" (m> (Figures 10-3) are almost identical. They share abrupt narrowing of umbilicus once at the second to fourth whorl (seealso Tanabe and Shigeta, 1987, fig. 4). It is
most conspicuous at 5-6 7l stage where umbilical shoulders project into the umbilical voidand form a remarkable "umbilical break."On the succeeding third to fourth whorl theumbilicus widens gradually, and finally theshell shows a somewhat evolute Puzosia-likemorphology. Kossmaticeras theobaldianum
paucicostatum (Figure 10-4) also shows aquite similar profile of juvenile shell shape.
In contrast, Mesopuzosia pacifica, which wasconsidered as the ancestor of "japonica" (M>,never exhibits such umbilical narrowing in
the early stage (Figure 10-6).
Early ontogenetic growth patterns of whorl
radius, breadth, and umbilical radius are
essentially similar among the three morphospecies (Figure 11). The growth of thewhorl radius (R) exhibits monophasic allometry at least for the early stage (2-12 7l
stage; D ~ 0.7-18 mm). That of whorl
breadth (B) of the three morphospecies showsdiphasic allometry, and the growth ratio
increases markedly at the 2-3 7l stage, which
may result from rapid whorl growth after the952.

Dimorphism of false puzosiines

7 lC'"

protoconch .......:::::.

2n'"

an'"

1

Figure 10. Median cross sections of early growth stage showing the ontogenetic change of whorl
shape and siphuncular position. 1, "Neopuzosia haboroensis Matsumoto and Inoma" (m>, UMUT.
MM18376, upper Santonian. 2, "N. ishikawai (Jimbo)" (M>, UMUT. MM18369, lower Santonian. 3,"Yokoyamaoceras jimboi Matsumoto" (m>, UMUT. MMI8372, lower Santonian. 4, Kossmaticeras
theobaldianum paucicostatum Matsumoto, UMUT. MM18374, Coniacian. 5, Desmoceras (Pseudouhligella) japonicum Yabe, UMUT. MM18379, middle Cenomanian. 6, Mesopuzosia pacifica Matsumoto, UMUT. MM18362, middle Turonian.

113

Haruyoshi Maeda

114

mm

mm

10

10.----------------------7~-------

0.1 1-------

0. N. ishikawai"'V. N. haboroensis·

0·Y. jimboi"

0.1

W.v.

W.v.
2

4

6

8

10

12

14

n 0.01

2

4

6

8

10

12

14

n

mm
10.--------------------------------

0.1 r----f'-1tf--------------------------

2

4

6

8

10

whorl volution

12

14

n

Figure 11. Ontogenetic allometry of radius vector (R), whorl breadth (8), and umbilical radius (C)in relation to whorl volution for three morphospecies in the early stage. Whorl volution of 2-12 7C
corresponds approximately to shell diameter of 0.7-18 mm.

952.
formationof the(=hatching).

primary

Dimorphism of false puzosiines
constriction

Double logarithmic plots of umbilicus versus shell diameter for selected specimens offour morphospecies show a similar growth
pattern throughout their ontogeny (Figure12). Remarkable reduction of umbilical

mm

liS

growth is present between 1-4 mm diameter,
and thereafter the umbilicus shows an
isometric growth pattern. The first flection
point at a diameter of 1.5 mm obviously
corresponds to the stage of abrupt umbilicalnarrowing in the second whorl. The second
flection point at a diameter of 3 mm reflects

u

50

o

~
....
~.......... ~®
.. @,'®

o®
®

2'()

o

10

..... <V

5

'<8>

2

o "N. ishikawai"e>"N. japonica"v"N. haboroensis"o "Y. jimboi"
0.5•

Mesopuzosia pacifica<8> Holotype<£> Lectotyp.e

0.2

mm
Figure 12. Allometric relationship between shell diameter (D) and umbilic~s (U) for t~e fo~rmorpho species and Mesopuzosia pacifica. Ontogenetic change of measurements In each specimen IS
linked by a line (same as in Figure 13).

Haruyoshi Maeda

116

its maximum decrease and formation of the"umbilical break" around the third whorl
stage (see also Figures 10, llC). The umbilical growth patterns of the four morpho species
markedly differ from that of Mesopuzosia
pacifica in the presence of growth reduction
at 1-4 or 5 mm diameter.
The ontogenetic changes of whorl shapeexpressed by breadth-height ratio (BjH) versus shell diameter (D) of the four morphospecies are also essentially similar up to
15-20 mm diameter, and show a gradualdecrease of this parameter with growth (Figure 13). At the stage less than I mm diame-ter, whorls are twice as wide as high, and the
BjH ratio then decreases rapidly. Rate 'Of
decrease reduces abruptly at a diameter of 4mm. In macroconchs, this ratio decreases
gradually after 15-20 mm diameter, and at
maturity it attains 0.6-0.7, which is somewhat
larger than that of Kitchinites (s.s.) species
from the Campanian to Maastrichtian (e.g.,
Howarth, 1965; Henderson, 1970). In fullgrown microconchs, the ratio is quite variable
as compared with macroconchs, and tuberculation appears coincidentally in "Y. jimhoi."
Contrarily, Mesopuzosia pacifica exhibits a

B/H
2.2

0

• N. ishikawai"<9• N. japonica·v• N. haboroensis·

0·Y. jimboi"•

Mesopuzosia pacifica

2.0<B> Holotype
0 Lectotype

1.5
\

\

\
I

\ I

o

\ I
\/

•

o

0

1.0

•

o(!)

~ (!) 00
(!)

(!)

0.6

,.

0.5

<0'
I

I

I

2

I
5

I

I

I

10

20

I

50

I
100

0

2ho D
mm

Figure 13. Ont?genetic changes of the breadth-height ratio (B/H) in relation to shell diameter (D)for four morphospecles plus Mesopuzosia pacifica.

952.

Dimorphism of false puzosiines

somewhat different pattern of decreasing BjH
ratio from those of the four morphospecies(Figure 13), particularly in the early growthstage in the presence of a sudden fall at 1.2
mm diameter and in the absence of the flection point at 4 mm diameter.
Dimorphism of four morphospecies
The following observations strongly suggest the presence of dimorphism among thefour morphospecies :I) They frequently co-occur in the
Santonian to lower Campanian in Hokkaido.
2) All morpho species share similar shellmorphology in the immature stage.
3) Their adult shell sizes are discretely
bimodal: the macroconch <M) (" ishikawai"and "japonica") has a simple aperture, andthe microconch <m) ("jimboi" and "haboroensis") has lappets.
4) Macroconchs may show slight numerical dominance, but great disparity in therelative frequency of the two froms is notobserved.
5) Some diagnostic features, such as rib
density and mode of tuberculation are unsuitable for taxonomic discrimination, because of
their wide range of variation.
However, the taxonomic barrier between
the Puzosiinae and Kossmaticeratidae has
prevented us from recognizing the dimorphism, and gave rise to somewhat complicated problems. Matsumoto et al. (1972)already commented on dimorphism in "Neopuzosia." They hypothesized that "ishikawai" <M) and lappeted "haboroensis" <m)were a dimorphic pair. On the other hand,
Matsumoto (1981,1991) also commented on
possible dimorphism between species of Yokoyamaoceras <m) and Kossmaticeras <M).
These hypotheses were, however, notconfirmed by these authors because of themarked disparity both in stratigraphic occurrence and frequency of presumed dimorphs.
For example, Y. jimboi <m) is quite common
in the lower Santonian of Hokkaido and117

Sakhalin, but Kossmaticeras spp., presumed<M), almost disappear In the upper
Coniacian.
Recently, Matsumoto (1988) described
dimorphism independently in every "Neopuzosia species." This is, however, quite
inconsistent with the situation in "N. haboroensis" which is represented only by lappeted
microconchs. Similarly, Alabushev andAlabusheva (1988) assumed sexual dimorphism within "Y. jimboi" by differences of
septation, shell ornament, and constrictions.
However, this interpretation should be rejected because both of their "dimorphs" areundoubtedly lappeted. Although the function of lappets is still obscure, this feature ispeculiar to the microconchs (Kennedy andCobban, 1976, etc.). The two forms in "jimboi" by Alabushev and Alabusheva (1988)
and" haboroensis" are, therefore, included in
the wide morphologic variation of microconch <m).
If we take no account of the taxonomic
artifice, but take "Neopuzosia" into consideration together with "Yokoyamaoceras" andKossma ticeras , differences in the range andthe occurrence of dimorphs disappear completely (Figure 7). Consequently, one "kossmaticeratid" and three "puzosiines" can beclassified as a single, dimorphic biospecies.
Higher taxonomic considerations
If one "kossmaticeratid" and three"puzosiines" can be lumped together under abiospecies, a problem naturally arises as to its
suprageneric pOSitIOn. Kennedy (1986)
presumed a dimorphic relationship between
Yokoyamaoceras and Neopuzosia, and placed
both genera together in Puzosiinae. On theother hand, morphological resemblance
between Kossmaticeras and Kitchinites (s.l.)including Neopuzosia was already pointed
out by previous authors (e.g., Kossmat, 1897 ;Yabe, 1927; Matsumoto, 1954b). Besides
the four morphospecies discussed here, there
are many puzzling examples as to whether

118

Haruyoshi Maeda

tuberculate and nontuberculate forms shouldbe placed in Puzosiinae or Kossmaticeratidae(Kennedy et'al" 1979, etc,). The ambiguityof diagnostic criteria between the two suprageneric groups is responsible for suchdoubts as to their classification (e.g., truncated ribbing by periodic constriction, andtuberculation).
As already admitted to some extent by
Matsumoto (1955, p. 155), the suturaldifference between "Yokoyamaoceras" and"Neopuzosia" is also ambiguous. "Ishikawai" <M>, "haboroensis" <m>, and "jimboi"<m> are not distinguishable from one anotherby their sutures at the same shell size, about
20 mm diameter. The sutures of kossmaticeratid "jimboi" <m> also show "puzosiine features," such as an asymmetrically tripartite L
and a retracted umbilical element (Matsumoto, 1954b, 1988; Figure 14).
In contrast, information on phylogeny is tobe found in early ontogeny, because ecological constraints may not so strongly influence
the juvenile shell morphology. Ontogenetic
changes of umbilicus/diameter ratio (U /0)
in "jimboi" <m>, "haboroensis" <m>, and"ishikawai" (including "japonica") <M> areshown in Figure 15 compared with those ofother desmocerataceans. The three morphospecies and Kossma ticeras theobaldianum paucicostatum (Kossmaticeratidae,
Coniacian) show a quite similar ontogenetic
pattern for this parameter with respect to aremarkable umbilical reduction in the 4-8 or
971 stage (about 1.3-7 mm shell diameter).
The minimum point in the graph corresponds
to the temporal formation of a sharp umbilical break at the 5-6 7l stage (about 2-3 mmshell diameter) on their profiles (Figure 10).
The temporal abrupt reduction of the umbilicus at an early stage seems to be a characteristic feature of the Kossmaticeras group,
Both Desmoceras (Pseudouhligella)japonicum (Oesmoceratidae, middle
Cenomanian; Figure 10-5) and Anapachydiscus sp. (Pachydiscidae, upper
Santonian) also possess similar patterns for

ontogenetic change of U /0 (Figure 15).
In contrast to this, true puzosiines such as
Mesopuzosia pacifica (Turonian; the direct
descendant of Puzosia), M. yubarensis(Coniacian), and limboiceras planulatiforme(Turonian) never show the narrowing of theumbilicus in the early growth stage. Theirumbilicus is still wide during the 4-9 7l stage(U /0 ~ 40%), and thereafter its umbilicus/
diameter ratio gradually decreases (U /0 ~35%). Hauericeras (Gardeniceras) angustum (Hauericeratinae, Santonian) also sharesa similar ontogenetic change of U /0 ratiowith Mesopuzosia. There are no intermediates between the Kossmaticeras and Mesopuzosia groups in this parameter. Neopuzosia

5mm
;

",

:
.i

",

__

.

!

2

3

:4
Figure 14. Suture lines of macro- and microconchs. 1, adult suture of "Neopuzosia ishikawai" <M) at about 140 mm diameter, UMUT.
MM6640, Santonian. 2-4, immature stu res atabout shell diameter of 20 mm; 2, "N. ishikawai"<M), UMUT. MM18380, upper Santonian; 3, "N.
haboroensis" <m), UMUT. MM18383, upper
Santonian; 4, "Yokoyamaoceras jimboi" <m),
UMUT. MMI8344, lower Santonian.

952.

UfD %

"

,
40

Dimorphism of false puzosiines

119

o

H· .

"

'. H

,,
,
\

,,

o

30

o "N. Ishikawa/"

\,
\,
\,

20

\l

\

0\

.
\

\
\

0 ... ·.. 0
10"N. haboroensis"o "Y. jimboi"/':, Kossmaticeras theobaldianum
paucicostatum• Mesopuzosia pacifica• Mesopuzosia yubarensis... Jimboiceras planulat/forme
o Desmoceras (P.J japon/cum

A Anapachydiscus sp.
H Hauericaras (G.J angustum
T

i

i

i

2

4

6

14
whorl volution

Figure 15. Ontogenetic changes of the umbilicus/diameter ratio (U/D) in relation to whorl volution
for three morpho species of" Yokoyamaoceras" and "Neopuzosia," and selected species of Kossmaticeras,
Desmoceras, Anapachydiscus, Mesopuzosia, and Hauericeras. This figure shows that the three morphospecies are clearly distinguished from the true puzosiines in the abrupt narrowing of umbilicus in thelimited stage of 4-9 7l stage (~ 1.3-7 mm shell diameter).

120

Haruyoshi Maeda

and Yokoyamaoceras obviously belong in
the Kossmaticeras group in terms of earlyshell shape. Thus the four morpho species inquestion should be placed in the family
Kossmaticeratidae.
Besides this, several morphological features
cast doubts on puzosiine origin of the fourmorphospecies. "Neopuzosia ishika wai"and "N. japonica" never exceed 150 mm
diameter, but true puzosiines, e.g., Mesopuzosia, commonly attain gigantic shell-size(500 mm or larger). In addition, the adultshells of Kossmaticeras (Collignon, 1955;
Kennedy and Klinger, 1985; Matsumoto,1991), Yokoyamaoceras, Neopuzosia (Figures
2,3-1,16-1,16-3), and Kitchinites (Kitchinites) (Henderson, 1970; Macellari, 1986)
similarly have a characteristic double collar
rib, interspaced with a sharp, deep furrow, on
the shell surface at the position of the periodic constriction on the internal mold. In
contrast, the collar feature of puzosiines isquite different. The shells of true puzosiines
have adapical collars only, and the furrow at
the constriction on the shell surface is obscure(Figure 16-2,16-4). These may also suppose the kossmaticeratid origin of "Neopuzosia" and Kitchinites. Comparable geographic distribution of Kitch in ites (Kitchinites) and diverse kossmaticeratids in thesouthern hemisphere during the lateCampanian to Maastrichtian (Henderson,
1970; Macellari, 1986; Bando et at., 1987
etc.) should also be noted.
The superficial resemblance of external
shell morphology of Mesopuzosia and Neopuzosia in the middle to late growth stages is ahomeomorphy, a phenomenon that is widespread among ammonites (Kennedy and Cobban, 1976).
The acceptance of kossmaticeratid affinitiesof the four morphospecies brings further
results. I agree with Matsumoto's presumption (Matsumoto, 1981, 1991) that the originof lineages with such dimorphs can be traced
back to the Coniacian, or further, to theTuronian (Figure 7) though examples from2

1

adapicaladapertural

....

<'

3

"

.... :: .. : ..

4

Figure 16. Comparison on collar rib of Yokoyamaoceras ishikawai (limbo) <M> with that of atrue puzosiine at the same shell size (D =approximately 100 mm). 3 and 4 are schematic profiles ofcollar ribs. 1,3, Y. ishikawai (limbo) <M>. 2,4,
Mesopuzosia pacifica Matsumoto. A sharp, deep
furrow between the collars on the shell surface is oneof the characteristic features of kossamaticeratids. It
does not develop in shells of true puzosiines.

lower stratigraphic horizons are rare.
"Neopuzosia haboroensis" (m) and "Yokoyamaoceras jimboi" (m) possibly originated in Yokoyamaoceras yokoii (m), Y. minimum (m) or Y. kotoi (m) (Figures 4-15,416) of Turonian-Coniacian age via someintermediate forms (Figures 4-13, 4-14).
On the other hand, the macroconch "Neopuzosia ishikawai" of Santonian-early
Campanian age can be traced likewise to thespecies of Turonian-Coniacian Kossm a ticeras (M), which is contemporaneous with theabove Yokoyamaoceras species (m). The
shell ornament may weaken simultaneously,in parallel, in both macroconchs and microconchs (Figure 7). Such simultaneouschange provides powerful arguments in support of dimorphism (Callomon, 1981),
though the morphologic change of the macroconch during the Santonian to lower
Campanian is slight.
Handling of such dimorphic "species"
involves several complicated nomenc1atorial952.

Dimorphism of false puzosiines

problems debated by some paleontologists(e.g., Westermann, 1969; Callomon, 1981).

I lump the four nominal species in a single
evolutionary biospecies belonging to the family Kossmaticeratidae. Thus the microconch
genus Yokoyamaoceras Wright and Matsumoto, 1954, and specific epithet of themacroconch: ishikawai of "Desmoceras ishikawai limbo, 1894," are adopted in therevised nomenclature of the species.
Conclusion

Three supposed puzosiines, Neopuzosia
ishikawai (limbo, 1894), N. japonica (Spath,1922), and N. haboroensis Matsumoto andInoma, 1972, and a kossmaticeratid Yokoyamaoceras jimboi Matsumoto, 1955, occurring from the Santonian to the lower
Campanian sequence of the Yezo Group in
Hokkaido, Sakhalin and KoryakKamchatka, are not independent biospecies
but dimorphs of a single species of Kossmaticeratidae, Yokoyamaoceras ishikawai(limbo, 1894) revised here (Figure 7). The
mode of ontogenetic shell growth, particularly the umbilicus, reveals a kossmaticeratid
origin for Y. ishikawai. Thus the dimorphic
lineage can be traced back to Kossmaticeras(M) and Yokoyamaoceras (m) of Coniacian-Turonian age.
Though little has been investigated about
dimorphism, there are several contemporaneous pairs of nontuberculate and tuberculate
genera in the family Kossmaticeratidae, e.g.,
M arshallites and Eomadrasites in the
Cenomanian, and Maorites and Jacobites of
the Campanian. The ancetral Kossmaticeras and the descendent Kitchinites should,
therefore, be more closely investigated todetermine the nature of their dimorphism.
Yokoyamaoceras ishikawai is a peculiar"false puzosiine" showing homeomorphy
with Mesopuzosia. On the other hand, true
puzosiines such as Puzosia and descendant
Mesopuzosia attain large shell size, and are
fairly abundant through the Albian to121

Coniacian of the Yezo Group in Hokkaido
and Sakhalin (Matsumoto, 1988; Maeda,
1991). However, they almost disappear in
the upper Coniacian. Y. ishikawai might
possibly have adapted to the vacated niche ofsuch true puzosiines during the Santonianand early Campanian, and succeeded in surviving in the Far East region of the northernhemisphere where Late Cretaceous kossmaticeratids are otherwise unknown.
In addtion, the umbilical ontogeny of
Mesopuzosia and Hauericeras is quite
different from that of other desmocerataceans(Figure 15). This casts doubt on the
puzosiine origin of Kitchinites, and may also
indicate a need for further phylogenetic reexamination of the Puzosiinae and Hauericeratinae in the large superfamily Desmocerataceae (Wright, 1981).
Acknowledgments

I wish to express my sincere thanks to Prof.
Itaru Hayami (University of Tokyo) for
encouragement and various suggestions
throughout this work. I am also muchindebted to Dr. William A. Cobban (U.S.
Geological Survey), Prof. Michael K. Howarth (British Museum), Prof. William 1. Kennedy (Oxford University), and Prof. Kazushige Tanabe (University of Tokyo) for valuable discussion about this work, and for
improvement of an earlier draft of this paper.
I thank Emeritus Prof. Tatsuro Matsumotoand Prof. luichi Yanagida (Kyushu University) who kindly provided me with many typespecimens from the University collections forthis study. Thanks are extended to Prof.
Kiyotaka Chinzei (Kyoto University), Prof.
Masayuki Tashiro (Kochi University), Prof.
Takashi Hamada, Mr. Takeo Ichikawa, Dr.
Tatsuo Oji (University of Tokyo), Yasunari
Shigeta (Mikasa City Museum), Drs.lkuwo
Obata, Tomoki Kase (National Science
Museum), Dr. Takashi Okamoto (Ehime University), Dr. Katsumi Abe (Shizuoka University), Prof. Hiromichi Hirano, and Dr. Hiro-Haruyoshi Maeda

122

shi Hayakawa (Waseda University) for theirhelpful discussion and cooperation both in
the field and laboratory.
Systematic paleontology

Superfamily Desmocerataceae Zittel, 1895
Family Kossmaticeratidae Spath, 1922
Subfamily Kossmaticeratinae Spath, 1922
Genus Yokoyamaoceras Wrightand Matsumoto, 1954
Type species.-Holcodiscus kotoi Jimbo,
1894, p. 33, pI. 5, figs. 2, 2a, b, by original
designation of Wright and Matsumoto (1954).
Synonym.-Neopuzosia Matsumoto, 1954
[type species: Kitchinites japonicus Spath,1922]. Yokoyamaoceras and Neopuzosia
were proposed in the same year, but the dateof publication for the former is July 31, whilethat of the latter is October 30.
Diagnosis.-A dimorphic genus of moderately involute or evolute, somewhat compressed shells with prorsiradiate ribs truncated somewhat obliquely by well-marked periodic constrictions, which are associated witha double collar rib. The cross section
exhibits temporal umbilical reduction, forming an umbilical break at the second to fo'urthwhorl stage. Macroconch three or four times
larger than lappeted microconch, but neverattains a large size. Alternate or opposite
ventrolateral tubercles appear in some adultmicroconchs.
Occurrence.-Middle Turonian to lower
Campanian of Hokkaido, Sakhalin, KoryakKamchatka, France, California, British
Columbia, Argentine, and Antarctica.

Yokoyamaoceras ishikawai (Jimbo, 1894)
Figures 3,4-1-4-14,6,9,10-1-10-3

Macroconch.Desmoceras gaudama (Forbes). Yokoyama, 1890, p.
184, pI. 19. figs. 5a, b.
Desmoceras ishikawai Jimbo, 1894, p. 28, pI. 17, figs.
5a, b.
Kitchinites japonicus Spath, 1922, p. 127.

Kitchinites ishikawai (Jimbo). Spath, 1922, p. 127.
Neopuzosia japonica (Spath). Matsumoto, 1954b, p.
91, pI. 17, figs. 1,2; pI. 18, figs. 1-4; pI. 23, fig.
1 ; Matsumoto, 1963, p.29, pI. 45, figs. 5a, b;
Matsumoto, 1988, p. 11 ; p.82, figs. 31 A-F, 32A,
33A-C.
Neopuzosia ishikawai (Jimbo). Matsumoto, 1954b,
p. 94, pI. 19, figs. 1-4; pI. 23, figs. 2a-d (see forcomplete synonymy); Matsumoto, 1963, p.43,
pI. 60, fig. 5; Vereshchagin, Kinasov, Paraketsoy, and Terekhova, 1965, p.50, pI. 53, figs. la,b; pI. 54, fig. I ; Matsumoto, 1988, p. II, figs.
32B-H,33I-K.
Kitchinites (Neopuzosia) japonicus Spath. Wright,
1957, p. 367; Haggart and Ward, 1989, p.220,
fig~ 3.6,5.9-5.12,~

Kitchinites (Neopuzosia) ishikawai (Jimbo). Wright,
1957, p. 367.
Microconch.Yokoyamaoceras kotoi Matsumoto, 1955, p. 152,
text-fig. 12.
Yokoyamaoceras jimboi Matsumoto, 1955, p.152153, fig. 13; pI. 9, figs. 4a, b, 5a, b (see for complete synonymy); Matsumoto, 1956, pI. 16, figs.
5a, b; Alabushev and Alabusheva, 1988, figs. 5,6; Matsumoto, 1991, p. 100, pI. 22, fig. 7; pI. 23,
figs. I, 2.
Kitchinites (Neopuzosia) haboroensis Matsumoto andInoma, 1972, p. 384, pI. 47, figs. 5, 6.
Kitchinites (Neopuzosia) aff. haboroensis (Matsumoto
and Inoma). Obata, Futakami, Kawashita, andTakahashi, 1978, p. 144, fig. 6, pI. I, figs.4a-c.
Kitchinites (Neopuzosia) cf. japonicus (Spath).
Obata, Futakami, Kawashita, and Takahashi,
1978, p. 143, pI. 2, figs. 2a-c.
Yokoyamaoceras aff. jimboi Matsumoto. Kennedy,
1986, p. 25, fig. 8-E; pI. I, figs. 6-8.

Diagnosis.- Yokoyamaoceras of compressed (BjH=0.6-0.7 in adult <M»),
Puzosia-like shell with very fine, weak,flexuous ribs numbering 30-50 per half awhorl, and periodic constrictions numbering
6-8 per whorl. A biconcave constriction
and a pair of lappets appear at the aperture ofadult microconch. The lower Santonian
microconch has impersistent, alternate
ventrolateral tubercles, numbering 6-18 per
70' -240' of a volution.
Lectotype.-UMUT. MM7507 (= lectotype
of Desmoceras ishikawai Jimbo, 1894, amacroconch of middle growth stage),designated by Matsumoto (1954, p. 94, as952.

Dimorphism of false puzosiines

123

Table 3. Dimensions (in mm) and ratios of Yokoyamaoceras ishikawai (limbo, 1984). I,
measurements of middle- to full-grown shells. 2, measurements of ontogenetic growthof UMUT. MMI8365 <M), from the upper Santonian of the Kotambetsu River (Loc.
KT-5031). - : calculated value; D: shell diameter; H: whorl height; B: whorl
breadth; U: diameter of umbilicus; Rib/HW: number of ribs per half a whorl;
Tbc/WV: number of ventrolateral tubercles per whorl volution; WV (7C): whorl
volution in 7C (= 180·) scale; R: whorl radius; c: umbilical radius; <L):
lectotype; (PC): protoconch.
(I)<Macroconch)

D

H

B

B/H

U

U/D%
32
32

54

Rib/HW

MM6642

26.8

0.90

37.7

9.8
15.5

8.8

MM7507 <L)

13.3

0.86

8.6
12.2

MM6637

54.0

23.0

18.0

0.78

17.1

32

45

MMI8340

80.2

30.2

23.2

0.77

28.0

MMI8337

94.7

37.1

25.1

0.68

31.0

35
33

26

41

MMI8341

-134

-50

30.7

-0.61

50.3

-38

32

<Microconch)

D

H

B

B/H

U

U/D%

Tbc/WV

MM18359

27.1

10.9

9.1

0.84

9.6

32.1

13.2

10.9

MM18351

37.2

15.5

11.3

0.83
0.73

10.8
11.9

35
34

11/130·MM18357
GK. H.5654

30.2

10.8

7.8

0.72

11.1

B

C

10/140·6/ 70·32
37(2)
WV (7C)

R

1

0.313

H

B/H

U

UjD%

0.0
16.6
31.5

0.328

0.585

(PC)

D

0.588

0.011

0.477

0.302

1.947

2

0.428

0.648

0.316

0.681

1.000

0.369

2.165
1.846

0.123

0.572

0.112
0.203

0.741

3
4

0.776

0.913

0.221

1.348

0.555

1.645

0.424

31.5

0.840

1.474

0.492

26.1

0.315

5

1.111

1.238

0.271

1.887

6

1.541

1.541

0.293

2.652

1.248

1.235

0.564

7

2.081

1.954

0.392

3.622

1.689

1.157

0.685

21.3
18.9

8

2.827

2.315

0.702

4.908

2.125

22.3

3.302

6.816

2.882

1.089
1.146

1.094

1.107

1.809

26.5

10

3.989
5.629

4.305

1.658

9.618

3.981

1.081

2.755

28.6

11

8.278

5.877

2.596

13.907

5.682

1.034

4.244

30.5

12

12.11918.1057.838
10.369

3.776
5.629

20.39730.2248.343
12.476

0.940
0.831

6.372
9.405

31.2
31.1

9

13

124

Haruyoshi Maeda"Neopuzosia ishikawai"), from the Santonianof the Obirashibe River, northwest Hokkaido,
Japan.
Material.-More than eighty specimensfrom the upper part of the Yezo Group at
various points in Hokkaido and Sakhalin.
UMUT. MM6624-6639, 6640-6644, 6675,
6883, 7507, 7623a, b, 7624a, b, 7625a, b, 7626,
7697,7698,7700,9137,18330-18353,1836518370,18376,18380-18387; GK. H3366,
5143,5201,5654-5656.
Dimensions.-Shown in Table 3.
Description.-Protoconch ellipsoidal, 0.5
mm wide and 0.3 mm diameter. Whorl globular at first. The umbilicus narrows temporally in the second to early fifth whorl stage(1.3- 7 mm shell diameter). In the third
whorl stage (0=2-3 mm; U/D = 20%), a
sharp umbilical break appears in the crosssection.
After the fifth whorl stage (0 ~ 9 mm), thecoiling becomes moderately evolute, with 40%of the previous whorl being covered. The
rather shallow umbilicus comprises 30-35% of
the diameter. The whorl section becomescompressed with growth, with the maximum
breadth towards the inner flank. Slightly
convex flanks converge to the roundedventrolateral shoulders.
Very fine, flexuous prorsiradiate ribs are
distinct from a diameter of 15 mm. The ribs
arise at the umbilical margin, being gently
flexed on the inner flank, and bend forwards
across the outer flank and ventrolateral shoulder to project over the venter in a fairly strong
convexity. These primary ribs bifurcate onceor twice on the mid-flank, and loop across theventer, while shorter ribs are also intercalated.
Narrow, deep, prorsiradiate periodic constrictions numbering 6-8 per a whorl, appear
at less than 5 mm diameter. They arestraight on the inner flank, but sweep forwards over the venter in a deep convexity.
Double collar ribs, interspaced with a deep
furrow, flank the constrictions and become
obvious in adults. The prominent adapicalcollar subdivides across the ventrolateralshoulder to give rise to two or four riblets
arranged in a virga toto me pattern. Parabolic lines commonly appear in front of theperiodic constrictions.
Phragmocones have 12-14 septa per whorl.
Body chamber length ranges from 240 to 280
of a volution. Sutures with a large tripartite
L and retracted umbilical elements, are variable. Macro- and microconchs are identicalin overall shell morphology up to 15-20 mm
diameter, and thereafter become dimorphic.
Macroconch <M>.-Fully grown shell
comprises 8-9 whorls (90-140 mm diameter).
The expansion rate decreases slightly in theadult body chamber, and the aperture
becomes narrower. The ribbing coarsens
somewhat and numbers 25-30 per half awhorl, strengthening particularly on the
periphery. Aperture simple. Tubercles
never develop.
Microconch <m>.-Fully grown shell, comprising 6-7 whorls, is 25-40 mm in diameter.
In lower Santonian forms, alternate
ventrolateral tubercles appear at 15-20 mm
diameter, and extend to the early half of theadult body chamber. They sometimes giverise to two or three fine ribs which loop acrossthe flank and converge at the umbilical margin. The upper Santonian to lower
Campanian forms have no tubercles. The
adapical collars are bullate on the venter insome microconchs. Finally, a biconcave
constriction, which is in close proximity to
the last normal constriction, paired lappets,
and a rostrum appear at the aperture. The
lappets are simple, tapering, and 2-5 mm
long.
Discussion.-UMUT. MM7623a (Figures
4-7,4-8) and MM7623b, which have adepressed body chamber (= larger B/H ratio;
Figure 13), were previously assigned to Y.
kotoi by Matsumoto (1955, p. 152, figs. 12,
13). However, these can be discriminatedfrom Y. kotoi by irregular, less extensive
tuberculation (about 160 in a volution), andthere are many intermediates between
depressed forms and typical "jimboi."
0

0

0

952.

Dimorphism of false puzosiines

Therefore, these can be included in the variation of the present species <m).
Y. ishikawai is very closely related toConiacian Kossma ticeras theobaldianum
paucicostatum Matsumoto, 1955 <M) andYokoyamaoceras kotoi (limbo, 1894) or Y.
minimum Matsumoto, 1956 <m) in many
respects. The species is, however, discriminated from such Coniacian forms by amore compressed shell form, finer, weakerribbing <M, m), and more irregular tuberculation <m). The macroconch of thisspecies also resembles the possible descendent
Kitchinites (Kitchinites), particularly K. (K.)
brevicostatus (Marshall) from the upper
Campanian to Maastrichtian of the Mata
Series, New Zealand in possessing fine,
flexuous prorsiradiate ribbing (Henderson,
1970, p. 32, pI. 13, fig. 4). The latter is, however, much more compressed than the present
species.
In addition, "Neopuzosia matsumotoi Collignon, 1961" from the Coniacian ofMadagascar (Collignon, 1961, p.54, pI. 23,
fig. 1) should be rather regarded as a true
puzosiine, Mesopuzosia as already stated by
Howarth (1965) and Matsumoto ( 1988),
because it never has double collar ribs.
Occurrence.-From the lower Santonian tolower Campanian in Hokkaido, Sakhalin,
Koryak-Kamchatka, and British Columbia,
California, and France. In Hokkaido and
Sakhalin, tuberculate microconchs (=" Y.
jimboi") are stratigraphically confined to themain part of the Inoceramus (Inoceramus)
amakusensis Zone in the lower Santonian.
Nontuberculate microconchs (= "N. haboroensis") are succeeded stratigraphically bytuberculate ones in the uppermost lower
Santonian to lower Campanian ( = uppermost
part of l. (I.) amakusensis Zone, and l.
(Platyceramus) japonicus and l. (Sphenoceramus) schmidti Zones).
Macroconch <M) (=" N. japonica") also
occurs from the Santonian to lower
Campanian in British Columbia and Washington (Haggart and Ward, 1989). Tuber-

125

culate microconch <m) (=" Y. jimboi") is
only known from the lower Campanian In
France (Kennedy, 1986).
References
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Dimorphism in the Late Cretaceous ammonite
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126

Haruyoshi Maeda

G.E.G. ed., Sexual dimorphism in fossil Metazoa
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952.

Dimorphism of false puzosiines- - , 1983: Collision orogenesis and sedimentationin Hokkaido, Japan. In, Hashimoto, M. andUyeda, S. eds., Accretion tectonics in the
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Abeshinai'tt'¥it!i'J, Bibai ~Ill'l., Embetsu JiJiU, Haboro :'flm, Hobetsu ~JiU, Hokkaido;ftW;J:ft Ikushumbetsu ~*JiU, Kaminosawa j:O)iR, Kikume(n)zawa *ilIiiiR, Kotambetsu"t!ftJiU, Kumaoizawa ~~~iR, Manji 77,*, Miginosawa ;;{:jO)iR, Mikasa -=~, Naibuchi!i'J{)1M, Obira +,¥, Obirashibe :t- 1::" '7 :/~, Oyubari *g5.&, Satonosawa {tc/!?iO)iR, Tannosawa
J#.O)iR, Tappu )t;:fJJ, Teshio ::R~, Urakawa ilIlror, Yezo ~~, Yubari g5.&.
l~r~*cf~MO)~:1'/.y7!iji7/~T1~, Yokoyamaoceras Wright and Matsumoto,
Neopuzosia Matsumoto, 1954 O)=~!Ji~: ;ftw:i!l: -'f'-lj- / ' 1) :/ 0) -lj-:/ =- 7 :/1954 S ~v:r- 7J :/ /~ =- 7 :/Il-M'T'illliJ> G [i'l]mB"Jvcit-t 0 "Neopuzosia ishikawai," "N. japonica," "N.
haboroensis" (7" '/ :/ 7 !IE M) c, "Yokoyamaoceras jimboi" (:1 ;t.. '< T -{ -t '7 ;t.. M) 0) 4%~fivC0~'-C, $~~;,tqC£..-j~5t~~:pJ~WL t..:. -to)*E:'1f1:, "4 fi" ft%t~I3t~O)$lii

Haruyoshi Maeda

128

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4-i-cm:m~.n-c ~t.:!I1J~Mt~1 ;f-Q)ge!l!.~Q)%Jt;IJ~7tM£*t;::~*-tJ:\,-':. c ;~ I?t;::, l)lPJml¥.Jt.(itl±l, 2) JIBil!:;IJ'I?Q)7il!:I~j(;:/ltQ)Mi{tl, 3) ~~Q)~1tt.( c";IJ'1? "4fm" fi
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129

PROCEEDINGS OF THE PALAEONTOLOGICALSOCIETY OF JAPANSome Cretaceous Radiolaria from Palawan, Philippines···············Fe P. Tumanda • Yolanda Maac-Aguilar
B *i!l~¥.o~~ 19931f.1f.~· ~~iI', I ~ 29-31 B I:::'

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standing nature of organisms, exemplified by recent.~Jfi(~;f.:/~~I?.il::±lLk • • ~.~~~~k::1

and fossil foraminifera ·· .. ···········Johann Hohenegger
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cp:Jtllcp~~~p'7Uff1:Jp=-.JE;;~1'fR£R~:lt............................................................ f:k 5C f!l Jf? 1 ::Ite Ban Sangao ~Te~ER.*I.J'i?1±l Lk He/eraphyllia (Anthozoa, Dividocorallia)............... fj.!.U'gf 1fl • :Iit*$':- • Nikorn Nakornsri

t reef builder ~~iJ!IC0\"-C ......................................................... cplif;~

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Holocene of Japan..................... Yukimitsu Tomita· Takahisa Goda
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Benthic Foraminifera from the type locality of the Jam-

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WII~~19ij·············································,j,¥r{g1;;-m-~.~~ '*'tTiiIf:h.iliill~m;;:t.t;::to~:Q ~R~~~~mft:o Schaubcylindrichnus ~=~JGfIi\'~:to J: U%ij\(;~~f~*~;;:t··········································~N.lE;fIl···························*~JIIlma~ • m.*oo:~ • "Ji.Jt3i§

Plagoniidae # (Nassellaria 111! §) ~7i"mt;:'"?\,'~

Jtfit<1'kJ,*,t;:iitr~;:t t.(\', ~ =- ~~mft:O·········~iR~­...................................................... ······:IlJ-JlI trm)IIII~* 1311~ J'I.~~fitll- -t~P"Ill1lfli\'~ c. t£$(-Fi1i~.~t;:.;z,..Gh:QtTU,1J 1 7'~~lljUIllJlE!~'g c.-t~·············································r,,'~~-· :<is:§ • .'iI:• • ~~~~M-r:Q=aJ'l.~~.rc:;C.1J7*/~-:7"'tRosiella ~ (.~) ~ *iiIf c. )ftft ....... ,. ·········:WOiR ~~v ~~~~~;6'G··················)!i~~~· fjt.1J-IEW'fi£::/;I( c. ~%1i.~;6, G.;z,..t.: 8!11!~=aJ'l. Nanonavis m~ir~~ - ~ t;: Nanonavis brevis a- '*',0 c. L ~ .•...••...•.•.•••...••••••••••••••••.•.•....•••..••••• •••••• JJQ~~

~flI'l....................................................··'j'MfrIj-.~ Stephanopyxis ~~~~~~*HIfIi\'~ c. 7i"m...................................................... ······7.f~lE;fIl
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1#=aJ'l.~::lp=-..f? ..............•.••••••........••.••..•. JJQ~~

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li~~ft:O;6,···············Mn:~J't· ~*~n:. s)II~I!i· i\1IjjJjJit~=:1Ji'I1l ~ i'I1l~Jtfit<1'kJ t;::to ~ :Q ijJUHI'~~~.............................................!lE~ 1ii.:m}J.j{~~~a~=aJ'l.m·····················.*.............................................!lE~ 1ii.~}J.j{ ~V/ -1:: Iv 7 ~~ C. Jtfit;j:§;6' G.;z,.. t.: i1!i:h.:i'lit;::to ~:Q *tT························:mllillm~ • B. Molfino • A. McIntyre
Polyptychoceras vancouverensis (Whiteaves), 1879 t;: '"?

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132

Palaeontological Society of Japan (PSJ) Council Actions
During its meeting on January 28, 1993, the PSJ Council enacted the followingchanges to its membership.
New members elected:
Naoko Egi,
Takeo Ichimura,Kazuya Kinoshita,
Takahisa Makiguchi,
Eiichi Nishimura,
Robert M. Ross,
Manabu Sugisawa,
Kazuhito Yamasaki,
Katsuhiko Yoshida,
Jie Yu.

Kiyofumi Hasegawa,N aoki Ikegami,
Katsuhiko Kishi,
Katsutomo Mano,
Hiroshi Oguri,
Kazuhisa Sasaki,
Kenshu Shimada,
Keiji Yamasaki,
Atsushi Yoshino,

Masashi Hirano,
Yasuhiro Itoh,
Naoki Kitamura,
Tsuneo Nakajima,
Hirokazu Ozawa,
Shinichi Sato,
Kazuo Tomita,
Tetsuji Yamasaki,
Ryuichiro Watanabe,

New Fellows approved:
Masao Futakami,
Yasufumi Iryu,
Yoshikazu Noda,
Seiichi Suzuki,

Yukio Gunji,
Akihiro Kano,
Kenichi Saiki,
Hitoshi Tanaka,

Hiroshi Hayakawa,Yasuo Kondo,
Takeshi Setoguchi,
Yukio Yanagisawa.

Resigned members:(Ordinary members) :Kunio Kanamori,Toshiaki Suyama,

Hiroaki Kobayashi,
Shuichi Yanai.

Mika Shigei,

Deceased members:(Fellows)
Y oshihisa Ohta,

Susumu Matsushita.

New Honarary members approved by the 1993 Annual Meeting of the Society heldon 29, January, 1993.
Keiji Nakazawa,
Katsura Oyama.

m~~~~(1993-1994)

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

April 30, 1993

New Series No . 169

CONTENTS
TRANSACTIONS
948.

Kuniteru Matsum aru , Myint Thein and Yujiro Ogawa: Early Miocene(Aquitanian) larger Foraminifera from the Shimizu Formation , A shizuri
Cape, Kochi Prefecture, Shikoku, Japan. . . .. . .. .. ... . ................ .

949.

Hisayo shi Igo , Katsumi

Ueno

and

KatsuoSashida : Lower Permianfu sulinaceans from Ban Phia, C hangwat Loei , northeastern Thailand.
950.

K azuhiro Sugiyama : Skeletal . structures of Lower and Midd le Miocene
lophophaenids (Radiolaria) from central Japan . . ....... . .. .. .. ... . .....

951 .

44

Li-Ming Zhao , T amiko Ohana and T atsuak i Kimura: A fossil population of

Ginkgo leaves from th e Xingyuan Formation , Inner Mongolia .
952.

15

73

H ar uyo shi Maeda: Dimorphism of Late Cretaceous false-pu zosiine ammo nites, Yokoyamaoceras Wright and M atsumoto , 1954 and Neopuzosia Matsumoto , 1954.

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

97

PROCEE DINGS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 129