ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No.

171

Pa laeo ntologica l Society of Ja pa n
September 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 University), Kei Mori (Tohoku University), Kenshiro Ogasawara (University of Tsukuba), Tsunemasa Saito (Tohoku University), Yokichi Takayanagi (Ishinomaki Senshu University), Kazushige Tanabe (University ofTokyo), 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),
Kazushige Tanabe (Planning), Hiroshi Noda (Membership; Co-Editor, Special Papers),
Hiromichi Hirano (Foreign Affairs), Itaru Hayami, Toshiyuki Yamaguchi (Editors,"Fossils"), Juichi Yanagida (Co-Editor, Special Papers), Tatsuaki Kimura (Friends ofFossils)

Secretaries: Katsumi Ueno, Shuko Adachi (General Affairs), Masanori Shimamoto (Editorialof TPPSJ), Akira Tsukagoshi (Planning), Katsuo Sashida (Membership), Tatsuo Oji(Editorial of "Fossils"), Takeshi Ishibashi, Akihiko Matsukuma (Editorial of Special
Papers), Tamiko Ohana (Friends of Fossils)
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 II cm)

All communication relating to this journal should be addressed to the

PALAEONTOLOGICAL SOCIETY OF JAPAN
c/o Business Center for Academic Societies,
Honkomagome 5-16-9, Bunkyo-ku, Tokyo 113, Japan

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 171, pp. 213-228,6 Figs., September 30, 1993

956.

UPPER CARBONIFEROUS FORAMINIFERS FROMBAN NA DIN DAM, CHANGWAT LOEI,NORTHEASTERN THAILAND*
KA TSUMI UENO AND HISA YOSHI IGO
Institute of Geoscience, University of Tsukuba, Ibaraki, 305

Abstract. White to pale gray, massive to thickly bedded limestone crops out in Ban Na
Din Dam, east of Loei, northeastern Thailand. This limestone belongs to the Lower
Permian Nam Mahoran Formation. We newly discriminated two important fusulinacean
species in this limestone; namely, Triticites (T.) samaricus Rauser-Chernoussova andJigulites grandis, sp. nov. These species strongly indicate a Gzhelian (latest Carboniferous)
age. The present discovery first confirms the presence ofthe Gzhelian fusulinacean fauna in
Thailand. Smaller foraminifers coexisting with these fusulinaceans are also listed andillustrated.
Key words.

Fusulinacean, Gzhelian, Loei, Thailand, Upper Carboniferous.

fusulinacean species which indicate aGzhelian age in the limestone belonging to
the Nam Mahoran Formation. The present
fossil locality is about 2 km east of Ban Na
Din Dam, Changwat Loei, northeastern
Thailand (Figure I). We describe these newly discriminated fusulinaceans and discuss thegeologic age of this fauna.

Introduction
The biostratigraphy of Carboniferous andPermian strata in Thailand was mainly established by means of fusulinaceans, and 20
almost continuously superposed zones werereported (Toriyama, 1984). Among them,zones indicating a Gzhelian (latest Carboniferous) age were uncertain, hence the boundary problem between the Carboniferous andPermian has never been fully discussed forthis country. Igo (1972) established theProtriticites tethydis Zone and Triticites
ozawai-Paraschwagerina yanagidai Zone in
the Loei-Wang Saphung area, northeastern
Thailand, and he concluded that the former
should be correlated with the uppermost
Myachkovian or lowermost Kasimovian andthe latter with the Asselian. Namely, heregarded that the uppermost Carboniferous
Gzhelian fusulinacean fauna is lacking in thisarea.
Recently, we discriminated interesting

Acknowledgments
The research was supported by a grant toH. Igo from the Monbusho International
Scientific Research Program, No. 02140210.
We thank Mr. Nikorn Nakornsri, Senior
Geologist of the Geological Survey Division,
Department of Mineral Resources, Thailand,for his field guidance. We also thank Drs.
Katsuo Sashida, Ken-ichiro Hisada and Miss
Apsorn Ampornmaha of the University ofTsukuba for their cooperation in the field.
Geologic Setting
Upper Paleozoic rocks distributed In the'Received June 17, 1992; accepted April 19, 1993
213

Katsumi Ueno and Hisayoshi [go214\

10 IOSI'E ,'"

\

,

I

I

,Ban Huai Muang

,,

,,

I
I

I
I

,
.
Ban Na Din Dam
I,--- .....

o
I

1

2km
I
I
I

Figure 1. Index map of Ban Na Din Dam, east of Loei, northeastern Thailand.

Loei area are divided into the Carboniferous
Wang Saphung Formation and the Permian
Ratburi (or Rat Buri) Group (Charoenpravat
et al., 1976). Although the Ratburi Group
has long been used as one of the most important Permian lithologic units all overThailand, it is in fact restricted to Permian
limestones distributed in PeninsularThailand, and the Saraburi Group (or Limestone) is commonly accepted to be in the Loei
Folded Belt (Buna')as, 1983). The lowest
division of the Saraburi Group is the Nam
Mahoran Formation that consists mainly ofwhite to pale gray, massive to thickly bedded
limestone. This limestone constitutes thecharacteristic topography of occasionally isolated high hills surrounded by steep cliffs,
with heavy tropical vegetation covering mostof them. One of these hills, located east of
Ban Na Din Dam, is partly exposed by recent

felling (Figure 1). Igo collected the present
materials from six different levels of thisoutcrop (Figure 2).
The lowest level, Loe-28, is white to pale
gray, thickly bedded limestone containing
abundant algae, fusulinaceans, and smaller
foraminifers. Microfacies of this limestone
is fusulinacean-algal packstone and grainstone.
The next higher level is Loe-29 and consistsof pale gray, thickly bedded limestone
containing hematitic matter and abundant
dasycladacean algae, fusulinaceans, and smaller foraminifers. Goniatites and brachiopods are also present but less common
compared with next higher levels.
The levels Loe-30, 31, and 32 are pale grayto white, thickly bedded limestone including
particularly abundant brachiopods,
bryozoans, goniatites, and crinoid columns

956.

Upper Carboniferous foraminifers from NE Thailand

215

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LO.2.! ~'•'40 .Loe31-J~
20 ,/~~_

I" • • Loe32

25 _~

Loe30

II

,

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II

II

II

II
II

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Loei

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Figure 2.

,
,. ,.
,.
,. II

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,
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50m

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Map showing fossil localities studied herein.

and less common gastropods, pelecypods, andtabulate corals. Brachiopod and goniatiteshells are frequently heaped in particular beds
as coquinoid limestone or sporadically embedded within wackestone. These limestone
beds do not yield any fusulinaceans but rarely
include smaller foraminifers.
The level Loe-33 represents the highest one
in this section and consists of a pale gray,partly dark brownish, thickly bedded limestone containing abundant fragments ofcrinoid columns, bryozoans, and algae (Tubiphytes sp.). Fusulinaceans are rare but smaller foraminifers are rather abundant in thislimestone. Microscopically, this is coarsegrained calcarenitic limestone.
The limestone exposed in this hill is lithologically very similar to that of the N am

Contour interval about 10 ffi.

Mahoran Formation typically exposed inTham Nam Mahoran, about 45 km south ofLoei from where Yanagida (1967) and Igo(1972, 1974) described Asselian brachiopods,
fusulinaceans, and conodonts.
Appraisal of foraminiferal faunas
Fusulinaceans and smaller formainifers
discriminated in the limestone exposed east of
Ban Na Din Dam are listed in Tables 1 and 2.
The level Loe-28 yields abundant Triticites(T.) samaricus Rauser-Chernoussova associated with smaller foraminifers. This
species was originally described from theSamara Bend of the Volga region by RauserChernoussova (1938) as a variety of secaUcus.
Rozovskaya (1958) redescribed this species

Katsumi Ueno and Hisayoshi /go

216

--

Table 1.

SpeciesSchubertella sp.

List of fusulinaceans from Loe-28, Loe-29, and Loe-33.

---Level

Biwaella ? sp.
Triticites (T.) samaricus Rauser-Chernoussova

Loe-28Loe-29

+
+
+++
+++

Jigulites grandis, sp. nov.
Rugosochusenella sp.

++

Rugosofusulina ? sp.
Nankinella spp.

Loe-33++++ + + : abundant + + : common +: rarefrom the Samara Bend and mentioned that therange of this species is restricted to CaB and
Cae. These units are correlated to the upper
Kasimovian to lower Gzhelian of the Russianstandard. Mikhaylova (1974) also reported
this species from the Triticites arcticus and T.
actus Zones (CaB) of the Pechora Urals.
Kanmera (1958) described this species fromthe Pseudoschwagerina Zone of the
Yayamadake Limestone, Kyushu, Japan.
Recently, Ozawa and Kobayashi (1990)
pointed out that this species is one of thediagnostic species of Zone AK30 (Daixina
robusta-"Pseudoschwagerina" minatoi Zone)of the Akiyoshi Limestone Group. They
considered that the zone should be correlatedwith the uppermost Gzhelian of the Russianstandard. According to Watanabe (1991),
this species occurs from the lower Asselian
Sphaeroschwagerina fusiformis Zone of the

Akiyoshi Limestone Group.
Chen and Wang (1983) described thisspecies as a subspecies of Triticites secalicus(Say) from the Sphaeroschwagerina sphaerica
gigas Zone of the Maping Limestone of
Yishan, Guangxi, South China.
Igo (1972) described Triticites samaricus
Rauser-Chernoussova from Ban Nam Lum,south of Phetchabun in central Thailand. In
Phetchabun this species is associated with
Quasifusulina tenuissima (Schellwien),
Triticites (T.) ozawai Kanmera, T. (T.) aff.
haydeni (Ozawa), Daixina lalaotuensis(Sheng), D. petchabunensis (Igo), and Paraschwagerina yanagidai Igo. He correlated
this fauna to those reported from the lower
Asselian of the Tethys region.
Among the above mentioned faunas, thoseof Yayamadake and Phetchabun should becorrelated with the uppermost Carboniferous~ Figure 3. 1-3: Spiroplectammina sp., longitudinal sections, IGUT-KU0380, IGUT-KU0381, IGUTKU0382. 4: Tuberitina bulbacea Galloway and Harlton, longitudinal section, IGUT-KU0383. 5:
Eotuberitina reitlingerae Miklukho-Maklay, longitudinal section, IGUT-KU0384. 6,7: Eotuberitina spp.,
longitudinal sections, IGUT-KU0385, IGUT-KU0386. 8: Diplosphaerina sp., longitudinal section, IGUTKU0387. 9,10: Globivalvulina spp., lateral sections, IGUT-KU0388, IGUT-KU0389. 11: Climacammina sp., longitudinal section, IGUT-KU0390. 12-14: Hemigordius sp. A, axial sections, IGUT-KU0391,
IGUT-KU0392, IGUT-KU0393. 15,16. Hemigordius? sp. B, 1~: axial section, IGUT-KU0394, 16:
oblique section, IGUT-KU0395. 17,18. Eolasiodiscus sp., 17: tangential section, IGUT-KU0396, 18:
sagittal section, IGUT-KU0397. 19: Endothyranella? sp., longitudinal section, IGUT-KU0398. 20:
Earlandia sp., longitudinal section, IGUT-KU0399. 21-23: Tetrataxis spp., axial sections, IGUTKU0400, IGUT-KU0401, IGUT-KU0402. 24,27,29. Bradyina spp., 24,27: axial sections, IGUTKU0403, IGUT-KU0406, 29: sagittal section, IGUT-KU0408. 25: Pachyphloia? sp., lateral section,
IGUT-KU0404. 26: Endothyra sp., axial section, IGUT-KU0405. 28: Pseudobradyina sp., sagittal section,IGUT-KU0407. 11,22, 29: x20, 1-10,12-16,19-21,23,24,27,28: x40, 17,18,25,26: X 100.

956.

Upper Carboniferous fo raminifers f rom NE Thailand''CS, G
..

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f.

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5

.

217

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,'

.5

6

7

9

3

12

13
15

19

25

10

16

21

20

26

27

28

Katsumi Ueno and Hisayoshi /go

218

Table 2.

List of smaller foraminifers from Loe-28 to Loe-33.

Species,Loe-28Loe-29Loe-31xx

x

x

Spiroplectammina sp.
Eotuberitina reitlingerae Miklukho-Maklay
E. spp.
Tuberitina bulbacea Galloway and Harlton
T. sp.
Diplosphaerina sp.
Earlandia sp.
Tetrataxis spp.
Bradyina spp.
Pseudobradyina sp.
Eolasiodiscus sp.
Globivalvulina spp.
Climacammina spp.
Endothyra spp.
Endothyranella ? sp.
Hemigordius sp. A
H.? sp. B
Pachyphloia? sp.

based upon the recent revised opinions concerning the Carboniferous and Permianboundary which were advocated by Ozawa
and Kobayashi (1990) and Watanabe (1991).
According to our consideration, the specimens previously described as Triticites samaricus by several authors do not belong to thesame species because these forms of Triticites
lack diagnostic features.
The level Loe-29 yields abundant Jigulites
gran dis , sp. nov. which is similar to Jigulites
longus (Rozovskaya) and J. magnus (Rozovskaya). These Russian species were describedfrom the Triticites (Rauserites) stuckenbergi
Zone (C 3 C), Jigulites jigulensis Zone (C 3 D),
and Daixina sokensis Zone (C 3 E) of theGzhelian. Daixina petchabunensis Igo described from a limestone exposed south ofPhetchabun is also similar to our Jigulites
gra n dis , but the former has more intensely
fluted septa and weakly developed chomata
than the latter. This level does not yield any
species similar to Carbonoschwagerina mori-

x
x
x

x
x

x
x

Loe-32Loe-33

x
x
x

x
x

x
x

x
x

x
x

x
x
x

x
x
x
x
x

x
x
x

x
x
x
x
x

kawai (Igo), "c." minatoi (Kanmera), andParasch wagerina shimodakensis Kanmera.
These species commonly occur in association
with Triticites (s.1.), Jigulites, and Daixina
and are regarded as the index species of theupper Hikawan (uppermost Carboniferous)of Japan (e.g., Ozawa and Kobayashi, 1990;
Watanabe, 1991). Fusulinaceans from thelevel Loe-29 are slightly older than thosereported from Ban Nam Lum. From theabove mentioned paleontological data, thislevel should be correlated with the middle
Gzhelian.
The next higher fusulinacean-bearing
level is Loe-33 and rarely yields Schubertellasp., Rugosochusenella sp, and Rugosofusulina ? sp. Although we obtained only one welloriented axial section, the occurrence of thegenus Rugosochusenella is worthy of note.
This genus was originally proposed from theWolfcampian (Lower Permian) of New Mexico, U.S.A. by Skinner and Wilde (1965).
The present Rugosochusenella sp. is more or956.

Upper Carboniferous foraminifers from NE Thailand

less similar to Rugosochusenella sp. A illustrated by Ozawa and Kobayashi (1990). The
latter unidentified species was introduced as adominant zonal species of Zone AK 30(uppermost Gzhelian) of the Akiyoshi Limestone Group. It is also similar to several
Rugosochusenella species reported from theuppermost Gzhelian to the "Schwagerina
Horizon" of various sections of Russia, such
as R. sangoneliensis Davydov, R. kalaikuchnensis Davydov, R. paragregaria (RauserChernoussova), R. hutiensis (Chen), andothers. Fossil contents are rather poor inlevel Loe-33, but it may be correlated with theDaixina sokensis Zone of the uppermost
Gzhelian (CsE).
As shown in Table 2 and Figure 3, we also
discriminated smaller foraminifers in eachlevel of the present section, but did not findany reliable species for detailed age assignment. Among them, Hemigordius sp. A and
H. ? sp. B are similar to the species which arerather common in the post-Sakmarian levelsof the Akiyoshi Limestone Group, but otherspecies are similar to forms reported hitherto
from the Carboniferous and Permian.
Systematic description

All specimens identified in this paper aredeposited in the paleontological collectionsof the Institute of Geoscience, University ofTsukuba (IGUT).
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea von Moller, 1878
Family Schubertellidae Skinner, 1931
Subfamily Schubertellinae Skinner, 1931
Genus Schubertella Staff
and Wedekind, 1910

Schubertella sp.
Figure 4-8

Description.-Shell small with 4 volutions,0.47 mm in length and 0.24 mm in width with219

a form ratio of 1.92. The first volution
coiled perpendicularly to outer ones.
Radius vectors of the first to fourth volution
0.03, 0.05, 0.08, and 0.13 mm, and form ratios
1.00, 1.22, 1.50, and 1.77, respectively. Outside diameter of pro loculus 0.030 mm. Septa
unfluted.
Remarks.- This species somewhat resembles Schubertella rara originally described by
Sheng (1963) from the lower part of the Qixia(Chihsia) Limestone of Guizhou (Kueichow)in its small shell size. The former, however,has less developed chomata than the latter.
Material studied.-Axial section; IGUTKU0416 from Loe-33.
Subfamily Biwaellinae Davydov, 1984
Genus Biwaella Morikawa
and Isomi, 1960

Biwaella ? sp.
Figures 4-1---4
The follwing description is entirely based on
one axial section illustrated on Figure 4-2.

Description.-Shell small and fusiform withbluntly pointed poles. Axial section of 4Yz
volutions 1.75 mm in length and 0.68 mm inwidth, giving a form ratio of 2.59. Shell
expands gradually throughout growth.
Radius vectors of the first to fourth, andfourth and a half volution 0.08,0.12,0.19,
0.29, and 0.39 mm, and form ratios 1.13, 1.50,
1.84,2.21, and 2.31, respectively. Proloculus
small and spherical, being 0.080 mm in outside diameter. Spirotheca thin and composed of a tectum and lower structurelesslayer in inner volutions, but of a tectum andfine alveolar keriotheca in outer ones.
Thickness of spirotheca of the first to fourth
volution 0.010,0.020,0.020, and 0.035 mm.
Septa weakly fluted only in extreme polar
regions. Chomata small and developed inall volutions except for the first one. Tunnel
angles of the third and fourth volutions 32
and 40 degrees.

Katsumi Veno and Hisayoshi /go

220

Remarks.- This unidentified species closely resembles some species of Biwaella, especially B.? tshelamtshiensis originally descri bed by Davydov (1984) from the Gzhelian
Jigulites altus Zone of southwestern Darvas.
The exact identification, however, is postponed until sufficient materials are accumulated.
Material studied .-Axia l sections ;IGUT-KU0409, IGUT-KU0410, IGUT-

KU0411. Sagittal section; IGUT-KU0412.
All specimens from Loe-28.
Family Schwagerinidae Dunbarand Henbest, 1930
Subfamily Chusenellinae Kahler
and Kahler, 1966
Genus Rugosochusenella Skinnerand Wilde, 1965

•
,

'.~

.

.

1

~•
.

~~
~~;

3

.~&
~ '~
.

7

' . ,'

6

5b

Figure 4. 1- 4. Biwaella? sp. , 1- 3: axial sections, IGUT-KU0409, IGUT-KU0410, IGUT-KU0411,
4: sagittal section, IGUT-KU04l2. 5, 6. Rugosochusenella sp., Sa: axia l section, IGUT-KU0413, 6:
sagitta l section, IGUT-KU0414, Sb: enlarged part of Sa. 7: Rugosofusulina? sp., axia l section , IGUTKU0415. 8: Schubertella sp. , axial section, IGUT-KU04l6. 9-19. Nankinella spp. , 9- 11,13- 19: axial
sections, IGUT-KU0417, IGUT-KU0418 , IGUT-KU04l9, IGUT-KU0421 , IGUT-KU0422, IGUT-KU0423,
IGUT-KU0424, IGUT-KU0425, IGUT-KU0426, IGUT-KU0427, 12: sagittal section, IGUT-KU0420. 7: X
10, Sa,6: x 15, 1- 4: x 20, 9- 19: x 30, Sb, 8: x 40.

956

···

Upper Carboniferous foraminifers from NE Thailand

19

221

~
20

Figure 5. 1- 20. Triticites (T.) samaricus Rauser-Chernoussova, 1- 4,7- 11,15- 17,19: axial sections,
IGUT-KU0428 , IGUT-KU0429, IGUT-KU0430, IGUT-KU0431 , IGUT-KU0434, IGUT-KU0435, IGUTKU0436, IGUT-KU0437, IGUT-KU0438, IGUT-KU0442, IGUT-KU0443, IGUT-KU0444, IGUT-KU0446, 5,
6,13,14: sagittal sections, IGUT-KU0432, IGUT-KU0433, IGUT-KU0440, IGUT-KU0441 , 12,20: slightly
oblique axial sections, IGUT-KU0439, IGUT-KU0447. All X 10.

Katsumi Ueno and Hisayoshi Igo

222

Rugosochusenella sp.
Figures 4-5-6

Description.-Shell small in schwagerinid
fusulinaceans and elongate fusiform withbluntly pointed polar ends. Axial section of
6Yz volutions (Figure 4-5a) 4.88 mm inlength and 1.63 mm in width, giving a form
ratio of 3.00. Early few volutions rather
tightly coiled. Radius vectors of the first to
sixth volution 0.09,0.12,0.20,0.31,0.49, and0.72 mm, and form ratios 1.55,2.25,2.38,3.09,
2.76, and 2.86, respectively. Proloculus
small and spherical, being 0.120 mm in outside diameter. Spirotheca thin and composed ofa tectum and keriotheca. Weak andirregular rugosity observed on surface of spirotheca. Thickness of spiro theca of the firstto fifth volution 0.010,0.020,0.035,0.050, and0.075 mm. Septa moderately fluted throughout shell length. Small chomata developedin all volutions. Tunnel angles of the second to sixth volution 36?, 28, 31, 37, and 27
degrees. Axial fillings restricted to polar
regions.
Remarks.- This unidentified species canbe distinguished from Rugosochusenella zelleri, the type species of the genus, describedby Skinner and Wilde (1965) from the Lower
Permian Horquilla Limestone in the Big
Hatchet Mountains of southwestern New
Mexico, in having a smaller shell, less tightly
coiled inner volutions and less developedrugosity on the surface of spirotheca.
Rugosochusenella sp. A (Ozawa andKobayashi, 1990) reported from Zone AK 30,
uppermost Carboniferous, of the Akiyoshi
Limestone Group, Japan is similar to ourpresent unidentified species but the former
has a larger shell than does the latter.
The present Rugosochusenella sp. is similar
to R. sangoneliensis Davydov, R. kalaikuchnensis Davydov, and R. paragregaria(Rauser-Chernoussova) reported from the
Dau:ina bosbytauensis-D. robusta Zone
(uppermost Gzhelian) of Darvas (Chuvashov
et al., 1986). Our present species has a smal-ler shell than these Russian species but moredetailed comparison is difficult because we
obtained only one axial section.
Material studied.-Axial section; IGUTKU0413. Sagittal section; IGUT-KU0414.
Both specimens from Loe-33.
Subfamily Schwagerininae Dunbarand Henbest, 1930
Genus Triticites Girty, 1904
Subgenus Triticites Girty, 1904

Triticites (Triticites) samaricus
Rauser-Chernoussova, 1938
Figures 5-1-20

Triticites secalicus var. samarica Ra userChernoussova, 1938, p. 112-113, 156, p1.4, figs.
1-2.

Triticites (Triticites) secalicus samaricus RauserChernoussova. Rozovskaya, 1958, p. 83, pI. 2,
figs. 7-8.
Triticites samaricus Rauser-Chernoussova. Kanmera,
1958, p. 168-171, pI. 26, figs. 1-13; Igo, 1972, p.
97-98, pI. 14, figs. 8-14; Ozawa and Kobayashi,
1990, pI. 5, fig. 5; Watanabe, 1991, figs. 4-6-13.
Triticites secalicus samaricus Rauser-Chernoussova.
Mikhay1ova, 1974, p.53-54, pI. 4, fig. 1 ; Chen
and Wang, 1983, p. 71, pI. 11, fig. 15.

Description.-Shell medium for genus andfusiform to elongate fusiform with bluntly
pointed polar regions and gently archedperiphery. Mature specimens having 5Yz to7, rarely 7Yz volutions, 5.03 to 6.88 mm inlength and 1.60 to 2.53 mm in width. Formratio ranges from 2.51 to 3.29, averaging 2.94
for 14 specimens.
Axis of coiling straight throughout growth.
Inner few volutions somewhat tightly coiled.
Radius vectors of the first to sixth volution ofwell oriented axial section (Figure 5-1) 0.15,
0.24, 0.38, 0.58, 0.82, and 1.09 mm, and form
ratios 1.87, 1.80, 1.74, 2.43, 2.65, and 2.95,
respectively.
Pro loculus small and spherical, sometimes
irregular in shape. Outside diameter ofproloculus ranges from 0.105 to 0.207 mm,
averaging 0.179 mm for 29 specimens.

956.
Table 3.

Upper Carboniferous foraminifers from NE Thailand

Measurements of Triticites (Triticites) samaricus Rauser-Chernoussova. (in mm)

Reg. no.
I
2

3
4
5
6
7
8
9
10

Figure

IGUT-KU0428
IGUT -KU0429
IGUT-KU0430
IGUT-KU0431
IGUT-KU0434
IGUT-KU0435
IGUT-KU0436
IGUT-KU0438
IGUT-KU0443
IGUT-KU0446

Length Width

5-1
5-2
5-3
5-4
5-7
5-8
5-9
5-11
5-16
5-19

6.43
6.88
6.45
6.00
6.43
6.53
5.03
6.63
5.78
5.53

2.35
2.33
1.98
2.05
1.95
2.20
1.75
2.53
1.98
2.10

F.R.
2.73
2.96
3.27
2.93
3.29
2.97
2.87
2.62
2.92
2.63

Radi us vectors
D.P.
0.195
0.125
0.180
0.130
0.205
0.180
0.140
0.195
0.210
0.140

Form ratio

I
2
3
4
5
6
7
8
9
10

1

2

3

4

5

6

0.15
0.11
0.15
0.11
0.16
0.15
0.12
0.16
0.16
0.13

0.24
0.18
0.24
0.17
0.28
0.22
0.19
0.24
0.27
0.20

0.38
0.27
0.38
0.27
0.46
0.39
0.28
0.35
0.42
0.30

0.58
0.49
0.56
0.43
0.56
0.59
0.44
0.53
0.58
0.46

0.82
0.81
0.81
0.65
0.91
0.85
0.67
0.76
0.79
0.63

1.09
1.13
1.05
0.98

7

1.17
0.92
1.06 1.35
1.05
0.92

Thickness of spirotheca
I

2

3

4

5

6

2.95
2.60
2.97
2.78

0.020

0.035
0.030
0.030
0.025
0.025

0.020
0.Q15

0.065
0.040
0.040
0.030
0.070
0.055
0.030
0.055
0.045
0.035

0.060
0.080
0.050
0.045
0.085
0.065
0.070
0.080
0.070
0.040

0.040
0.060
0.060
0.060

2.71

0.040
0.045
0.045
0.025
0.035
0.040
0.020
0.035
0.055
0.030

1

2

3

4

5

6

1.87
1.55
1.33
1.82

1.80
1.79
1.96
1.89
2.07
2.13
1.65
1.60
1.61
1.52

1.74
2.21
2.03
2.11
2.04
2.45
1.97
1.75
1.86
1.65

2.43
1.92
3.16
2.23
3.14
2.51
2.77
2.16
2.20
1.98

2.65
2.08
3.10
2.31
2.90
2.99
2.65
2.74
2.47
2.52

1.76
1.53
1.25
1.19
1.18
1.00

223

7
-

0.025
0.Q15
0.Q15

-

2.61
2.67
2.45

2.68

0.025
-

0.020
0.025
0.045
0.025

0.085
0.060
0.080
0.075
0.065

7

0.090

F.R.: Form ratio
D.P.: Diameter of pro1ocu1us
Tunnel angle (degrees)

1
2
3
4
5
6
7
8
9
10

1

2

3

24
32
21
21
25
25
21
25
22
27

21
29
34
24
25
21
25
28
26
24

36
51
30
40
31
25
29
30
28
27

4

5

39

43
34
76
39
41
46
42
36
42
36

33
33
35
38
35
34
31
33

6

7

36

-

47
78
-

36
52
45

37

224

Katsumi Ueno and Hisayoshi [goSpirotheca thin and consists of a tectumand keriotheca. Thickness of spirotheca of
the first to sixth volution of above-mentioned
axial section 0.020, 0.035, 0.040, 0.065, 0.060,
and 0.040 mm.
Septa thin and fluted only in extreme polar
regions. Septal counts of the first to sixth
vol uti on of one sagittal section (Figure 5-13)
10, 16, 19,21,22, and 22.
Chomata massive and developed in all
volutions except for the last one in somespecimens. Tunnel path narrow and almost
straight. Tunnel angles of the first to sixth
volution of well oriented specimen mentioned
above 24,21,36,39,43, and 36 degrees. No
axial fillings present.
Remarks.- This species is almost identicalwith Triticites (T.) samaricus originally described by Rauser-Chernoussova (1938) fromthe Upper Carboniferous of the Samara Bendin the Russian Platform. Subsequently,
Rozovskaya (1958) and Mikhaylova (1974)
also described this species from the RussianUpper Carboniferous. The present specimens, however, have a slightly larger shellthan those described in Russia.
In Thailand, Triticites (T.) samaricus wasreported by Igo (1972) from Ban Nam Lum,south of Phetchabun, where such
fusulinaceans as Quasifusulina tenuissima(Schellwien), Triticites (T.) ozawai Kanmera,
T. (T.) aff. haydeni (Ozawa), Daixina
lalaotuensis (Sheng), D. petchabunensis (Igo),
and Paraschwagerina yanagidai Igo are associated.
Triticites (T.) samaricus is closely similar
to T. (T.) noinskyi Rauser-Chernoussova but
differs from the latter in having a more tightly
coiled juvenarium.
The present species somewhat resembles
Triticites (T.) variabilis described by Rozovskaya (1950) in general shell shape. The for-mer, however, can be distinguished from thelatter in having more massive chomata.
Material studied.-Axial sections;
IGUT-KU0428, IGUT-KU0429, IGUTKU0430, IGUT-KU0431, IGUT-KU0434,
IGUT-KU0435, IGUT-KU0436, IGUTKU0437, IGUT-KU0438, IGUT-KU0442,
IGUT-KU0443, IGUT-KU0444, IGUTKU0446. Sagittal sections; IGUT-KU0432,
IGUT-KU0433, IGUT-KU0440, IGUTKU0441. Slightly oblique axial sections;
IGUT-KU0439, IGUT-KU0447. All specimens from Loe-28.
Measurements.-See Table 3.
Genus Jigulites Rozovskaya, 1948

Jigulites grandis Ueno and Igo, sp. nov.
Figures 6-1-16

Diagnosis.-Large Jigulites having afusiform to elongate fusiform shell, fluted
septa restricted to polar regions, well developed massive chomata and narrow tunnel
path.
Description.-Shell large for genus andfusiform to elongate fusiform with bluntly
pointed polar ends and almost straight lateralslopes. Mature specimens have 6 to 731;
volutions, 5.63 to 9.38 mm in length and 2.33
to 3.28 mm in width. Form ratio varies from2.32 to 3.05, averaging 2.51 for 13 specimens.
The holotype of 731; volutions 8.25 mm inlength and 3.20 mm in width with a form
ratio of 2.58.
Shell expands uniformly throughout
growth. Axis of coiling straight. Radius
vectors of the first to seventh volution of theholotype 0.l3, 0.21, 0.33, 0.53, 0.81,1.13, and1.48 mm, and form ratios 1.23, 1.55, 1.59, 1.64,
2.16, 2.35, and 2.69, respectively.
Prolocul us spherical and 0.130 to 0.230 mm~ Figure 6. 1-16. Jigulites grandis, sp. nov., 1: axial section of the holotype, IGUT-KU0448, 2-6,
9-12, 15, 16: axial sections of paratypes, IGUT-KU0449, IGUT-KU0450, IGUT-KU0451, IGUT-KU0452,
IGUT-KU0453, IGUT-KU0456, IGUT-KU0457, IGUT-KU0458, IGUT-KU0459, IGUT-KU0462, IGUTKU0463, 7,8, 13: sagittal sections of paratypes, IGUT-KU0454, IGUT-KU0455, IGUT-KU0460, 14: tangential section of paratype, IGUT-KU0461. All X 10.

956

Upper Carboniferous foraminifers from N£ Thailand
225

Katsumi Ueno and Hisayoshi /go

226
Table 4.

Measurements of Jigulites grandis, sp. nov. (in mm)
Radi us vectors

1
2
3
4
5
6
7
8
9
10

Reg. no.

Figure

IGUT-KU0448
IGUT-KU0449
IGUT-KU0450
IGUT-KU0451
IGUT-KU0452
IGUT-KU0453
IGUT-KU0456
IGUT-KU0458
IGUT-KU0462
IGUT-KU0463

6-1
6-2
6-3
6-4
6-5
6-6
6-9
6-11
6-15
6-16

Length Width
8.25
6.00
6.85
6.43
6.15
5.90
6.30
5.63
7.33
7.40

3.20
2.38
2.95
2.73
2.33
2.35
2.63
2.35
3.03
2.80

F.R.
2.58
2.53
2.32
2.36
2.65
2.51
2.40
2.39
2.42
2.64

D.P.
0.160
0.180
0.155
0.180
0.210
0.145
0.190
0.155
0.130
0.230

Form ratio

I
2
3
4
5
6
7
8
9
10

6
7
8
9
10

2

3

4

5

6

7

0.13
0.16
0.15
0.14
0.17
0.13
0.15
0.13
0.13
0.19

0.21
0.24
0.24
0.23
0.28
0.21
0.26
0.21
0.20
0.31

0.33
0.38
0.35
0.36
0.43
0.36
0.45
0.35
0.35
0.46

0.53
0.63
0.53
0.52
0.68
0.56
0.61
0.52
0.54
0.70

0.81
0.91
0.74
0.73
0.93
0.86
0.88
0.77
0.80
0.96

1.13
1.24
1.07
0.96
1.20
1.11
1.22
1.13
1.10
1.29

1.48

I

2

3

4

5

6

7

1

2

3

1.23
1.44
1.27
1.14
1.28
1.23
1.27
1.54
1.38
1.15

1.55
1.68
1.36
1.42
1.93
1.41
1.74
2.36
1.57
1.47

1.59
1.72
1.89
1.51
1.95
1.65
1.67
2.08
1.36
1.89

1.64
1.91
1.85
1.85
2.03
1.76
2.14
2.37
1.66
2.07

2.16
2.18
2.01
1.88
2.26
2.34
2.27
2.80
2.24
2.56

2.35
2.42
2.25
2.15
2.53
2.26
2.15
2.65
2.54

2.69

0.030
0.025
0.030
0.020
0.030
0.025
0.025
0.030
0.025
0.030

0.035
0.020
0.040
0.035
0.035
0.040
0.035
0.040
0.030
0.035

0.040
0.075

2.27
2.46

2.53

-

1

2

3

4

5

6

7

18
12

16
16
20
20
22

17
21
22
21
17
25
17
24
31
15

21
28
24
34
22
19
24
20
29
21

22
34
30
34
31
20
18
27
36
32

34
31
31
38
32
27
25
40
43
40

40

-

21
-

23
18
29

24

21
21
24
23
19

1.42
1.27

1.44

Thickness of spirotheca

Tunnel angle (degrees)

1
2
3
4
5

!

45
41

51

4

0.060
0.055
0.065
0.045 0.055
0.040 0.060
0.060 0.060
0.055 0.080?
0.045 0.070
0.045? 0.070
0.050 0.050

5

6

7

0.055
0.075
0.060
0.070
0.075
0.070
0.085
0.075
0.070
0.080

0.095
0.110
0.090
0.080
0.060
0.115
0.085
0.085
0.095
0.060

0.095
0.100
0.130

0.105

956.

Upper Carboniferous foraminifers from NE Thailand

in outside diameter, averaging 0.185 mm for
25 specimens.
Spirotheca thin and composed of a tectumand keriotheca. Thickness of spiro theca of
the first to seventh volution of the holotype
0.030,0.035,0.040,0.060, 0.055, 0.095, and0.095 mm.
Septa thin, numerous and fluted rather
intensely in polar regions, but with decreasing
intensity toward central part of shell. Septal
counts of the first to seventh vol ution in
typical sagittal section of paratype (Figure6-13) 10,13,20,28,28,32, and 307.
Chomata broad, massive and well developed in all volutions. Tunnel path narrow
and almost straight. Tunnel angles of thefirst to seventh volution of the holotype 18,
16, 17,21,22,34, and 40 degrees. Axial
fillings not developed.
Remarks.-Jigulites longus originally described by Rozovskaya (1950) is the closest to
the present new species. The former, however, can be distinguished from the latter inhaving more intensely fluted septa and smaller chomata.
The present new species is also related to
Jigulites magnus (Rozovskaya) but differsfrom the latter in having a larger form ratio,less fluted septa in the central part of the shelland more well developed chomata throughout growth.
Jigulites grandis, sp. nov. somewhat resembles Daixina petchabunensis described by Igo(1972) from Ban Nam Lum, south of Phetchabun, central Thailand. The former, however, is distinguished from the latter in having
less fluted septa and massive chomata even in
the outer volutions.
Etymology.- This specific name is derived
from the Latin gran dis , meaning grand, greator large.
Material studied.-Axial section of theholotype; IGUT-KU0448. Axial sectionsof paratypes; IGUT-KU0449, IGUTKU0450, IGUT-KU0451, IGUT-KU0452,
IGUT-KU0453, IGUT-KU0456, IGUTKU0457, IGUT-KU0458, IGUT-KU0459,

227

IGUT-KU0462, IGUT-KU0463. Sagittal
sections of paratypes; IGUT-KU0454,
IGUT-KU0455, IGUT-KU0460. Tangential
section of paratype; IGUT-KU0461. All
specimens from Loe-29.
Measurements.-See Table 4.
Genus Rugosofusulina RauserChernoussova, 1937

Rugosofusulina 7 sp.
Figure 4-7

Remarks.-Presence of rugosity in the spirotheca of this species seems to suggest that itis referable to the genus Rugosofusulina.
Material studied.-Axial section; IGUTKU0415 from Loe-33.
Family Staffellidae Miklukho-Maklay, 1949
Genus Nan kin ella Lee, 1934

Nankinella spp.
Figures 4-9-19

Remarks.-Several forms such as alenticular shell with angular periphery (e.g.,
Figure 4-10), a lenticular shell with shallow
umbilicus (e.g., Figure 4-11), and a thicklenticular shell with angular periphery (e.g.,
Figure 4-17) are identified, which are possibly split into several distinct species. In
this study, however, they are all treated as
Nankinella spp.
Material studied.-Axial sections
IGUT-KU0417, IGUT-KU0418, IGUTKU0419, IGUT-KU0421, IGUT-KU0422,
IGUT-KU0423, IGUT-KU0424, IGUTKU0425, IGUT-KU0426, IGUT-KU0427.
Sagittal section; IGUT-KU0420. All specimens from Loe-29.
References cited
Bunapas, S., 1983: Paleozoic succession in Thailand.
Proceedings oj the Workshop on Stratigraphic
Correlation oj Thailand and Malaysia, vol. I,

228

Katsumi Ueno and Hisayoshi [goTechnical Paper, p. 29-76.
Charopenpravat, A., Wongwanich, T., Tantiwanit,
W. and Theetiparivatra, V., 1976: Geological
Map of Changwat Loei, Sheet NE47-12, ser. 1 :250,000. Geol. Surv. Div., Dept. Min. Res.
Thailand.
Chen, X. and Wang, J.H., 1983: The fusulinids of theMaping Limestone of the Upper Carboniferous
from Yishan, Guangxi. Palaeont. Sinica, N.S.
B, no. 19, p. 1-139, pis. 1-25. (in Chinese withEnglish abstract)
Chuvashov, B.N., Leven, E.Ya. and Davydov, V.I.,
1986: Pogranichye otlozheniya karbona i permi
Urala, Priural'ya i Sredney Azii (biostratigrafiya
i korrelyatsiya). 15Ip., 32pls. Akad. Nauk SSSR,
Ural'skiy Nauchnyy Tsentr, Inst. geol. geokhim.
im akademika A.N. Zavaritskogo, Izdatel.
Nauka, Moskva. (in Russian)
Davydov, V.I., 1984: K voprosu 0 proiskhozhdenii
shvagerin. Paleont. Zhurnal, 1984, no. 4, p.316, pI. 1. (in Russian)Igo, H., 1972: Fusulinacean fossils from Thailand,Part VI. Fusulinacean fossils from northThailand. Geol. Palaeont. Southeast Asia, vol.
10, p. 63-116, pis. 9-19. Univ. Tokyo Press,
Tokyo.
- - , 1974: Lower Permian conodonts from northern
Thailand. Ibid., vol. 14, p. 1-6, pI. 1.
Kanmera, K., 1958: Fusulinids from the Yayamadake Limestone of the Hikawa Valley, Kumamoto
Prefecture, Kyushu, Japan, Part III. Fusulinidsof the Lower Permian. Mem. Fac. Sci., Kyushu
Univ., ser D, Geol., vol. 6, no. 3, p. 153-215, pis.
24-35.
Mikhaylova, Z.P., 1974: Fuzulinidy verkhnego karbona Pechorskogo Priural'ya. 116 p., 17 pis.
Akad, Nauk SSSR, Komi Filial, Inst. Geol.,Izdatel. Nauka, Leningrad. Otdel., Leningrad. (inRussian)
Ozawa, T. and Kobayashi, F., 1990: Carboniferous
to Permian Akiyoshi Limestone Group. In,

Organizing Committee Benthos '90 ed., Fossil
and recent benthic foraminifera in some selected
regions of Japan. Guidebook for field trips,
4th International Symposium on Benthic Foraminifera, Sendai, 1990, p. EI-E31, pis. 1-13.
Rauser-Chernoussova, D.M., 1938: Verkhnepaleozoyskie foraminifery Samarskoy luki i Zavolzh'
ya. Trudy Inst. Geol. Nauk, Akad. Nauk
SSSR, vol. 7, p.69-167, pis. 1-9. (in Russianwith English summary and description)
Rozovskaya, S.E., 1950: Rod Triticites, ego razvitie istratigraficheskoe znachenie. Trudy. Paleont.
Inst., Akad. Nauk SSSR, vol. 26, p. 1-78, pis. 110 (in Russian)- - , 1958: Fuzulinidy i biostratigraficheskoe raschlenenie verkhnekamennugol'nykh otlozheniySamarskoy luki. Trudy Geol. [nst., Akad.
Nauk SSSR, vol. 13, p.57-120, pis. 1-15. (inRussian)Sheng J.Z. (Sheng, J.c.), 1963: Permian fusulinids ofKwangsi, Kueichow and Szechuan. Palaeont.
Sinica, N.S.B, no. 10, p. 1-147, pis. 1-36.
Skinner, J.W. and Wilde, G.L., 1965: Lower Permian(Wolfcampian) fusulinids from the Big Hatchet
Mountains, southwestern New Mexico. Contr.
Cushman Found. Foram. Res., vol. 16, pt. 3, p.
95-104, pIs. 13-15.
Toriyama, R., 1984: Summary of the fusuline faunasin Thailand and Malaysia. Geol. Palaeont.
Southeast Asia, vol. 25, p. 137-146, Univ. Tokyo
Press, Tokyo.
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.
Yanagida, J., 1967: Early Permian brachiopods from
central Thailand. Geol. Palaeont. Southeast
Asia, vol. 3, p.46-97, pIs. 11-23. Univ. Tokyo
Press, Tokyo.

r-

Akiyoshi tkE', Guangxi($) gg, Guizhou (Kueichow) :Ji::N'I, Kyushu jlJIi, Maping.l?7 (.~) :>jZ, Qixia (Chihsia) ~ii, Yayamadake ~illffi, Yishan Jtill.
~ 11E~t*$Cl.I1~TT l' /~ .b.t-.t0)J:.$1:i~*ift1HL!H: !l-1 OO""(>ftilJ:.WEl!R*?'-Ii P 7 :di1c~t~~-t" ~Wi~!R.{r.Eft.: ,h:£ ""(>~1!f~.n,tJ:'-'o 4-@1 p ~ -1 #:ili: (7) rW......:.
Iv A*:t" A .... ;/< '7 '/ II.!::. ~.n~ EW<:;5;b' G, ?' -Ii P 7 '/ ~fl~l¥.n::.t~~-t" 7.> .!: . .7,..G.n ~Wi~$.:to J: V:/NW1f:fL$.{r.E~~~1j L t.:(7)·C~tU~.1!f-t" 7.>0 ±~tJ:~ft Triticites (T.) samaroo-nicus Rauser-Chernoussova .!::.~~(7) Jigulites grandis ""(>~~o 4-@I(7)j6J!tcJ: 1J!l-1 OO""(>~
El!R*.!::. """:'Iv A*(7)~~J'j!·rlmm;b~~ff:flltc~ L.G.n 7.> 1iJ~M:;b;~$ <tJ: -? t.::.o

J:.!flm~·~ml~~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 171, pp.229-236, 3 Figs., September 30, 1993

957. SPATHIAN AMMONOIDS METADAGNOCERAS AND
KEYSERLINGITES FROM THE OSAWA FORMATION
IN THE SOUTHERN KITAKAMI MASSIF,NORTHEAST JAPAN*
MASA YUKI EHIRO
Department of Earth Sciences, College of General Education,
Tohoku University, Sendai, 980

Abstract. Two ammonoid species, Metadagnoceras motoyoshiense, sp. nov. andKeyserlingites cf. K. middendorffi, are described from the middle part of the Osawa Formation of the Kitakami Massif. They are associated with Columbites and Subcolumbites.
Both Metadagnoceras and Keyserlingites are described herein from Japan for the first time.
They are index fossils of the Spath ian Stage (uppermost Lower Triassic) and support theprevious opinion that the Osawa Formation is correlative with the Spathian or upper
Olenekian. The significance of the ammonoid fauna of the Osawa Formation is also
discussed with its relationship to the zonal subdivision of the Spathian.
Key words.
Northeast Japan

Metadagnoceras, Keyserlingites, ammonoid, Triassic, Kitakami Massif,of Metadagnoceras and Keyserlingites fromthe Osawa Formation provides not only abasis for world-wide correlation of the formation but also data for reexamination of thezonal scheme of the Spathian.

Introduction
This paper reports the occurrence of ammonoid species belonging to the genera
Metadagnoceras and Keyserlingites in association with the Columbites-Subcolumbites
fauna from Lower Triassic strata in theSouthern Kitakami Massif, Northeast Japan.
In 1991, Miyuki Numakura, Atsushi Numakura and Makoto Kumagai collected someammonoid fossils, in which two specimens
are described herein as Metadagnoceras
motoyoshiense, sp. nov. and Keyserlingites cf.
K. middendorffi (Keyserling), from. theOsawa Formation exposed at Yamaya,
Motoyoshi District. One additional specimen belonging to the former and many specimens of some other ammonoid species werecollected later by the author. The discovery

The Osawa Formation andits ammonoid fauna
The Lower to Middle Triassic strata in theSouthern Kitakami Massif are called the Inai
Group. This group is divided into the Hiraiso and Osawa Formations of Scythian age,and the Fukkoshi and Isatomae Formationsof Anisian age, in ascending order. The
Osawa Formation, about 300 m in thickness,consists of dark gray, finely banded shales,
which are intercalated with medium- to finegrained sandstones. Two ammonoid zones
have been recognized in the formation: thelower Subcolumbites Zone and the upper
Arnautoceltites Zone (Bando and·Received December II, 1992; revised manuscript
accepted April 19, 1993
229

Masayuki Ehiro

230

Shimoyama, 1974). From the Subcolumbites
Zone, such ammonoid fossils as Preflorianites
aff. P. sulioticus (Arthaber), Columbites parisianus Hyatt and Smith, Subcolumbites perrinismithi (Arthaber), Eophyllites cf. E.
dieneri (Arthaber) and Leiophyllites sp. havebeen described (Bando and Shimoyama,1974). The Arnautoceltites Zone yields
Procarnites kokeni (Arthaber), Arnautoceltites sp., Isculitoides aff. 1. originis(Arthaber), Prenkites cf. P. timorensis Spath,
Eosturia towaensis Bando and Ehiro andothers (Bando and Shimoyama, 1974; Bandoand Ehiro, 1982). On the basis of theseammonoids, the Osawa Formation is correlated with the Columbitan to Prohungaritan or
the Spathian of the Scythian (Bando andShimoyama, 1974; Bando and Ehiro, 1982).
The ammonoid fossils, Metadagnoceras
motoyoshiense, sp. nov. and Keyserlingites cf.
K. middendorjJi, described herein were collected from shale beds in the middle part of
the Osawa Formation exposed at a location
to the northwest of Yamaya, about 1.5 km
northeast of Hiraiso Coast, which is the type
locality of the formation (Figure 1). The
present fossil horizon belongs to the upper
part of the Subcolumbites Zone of Bando andShimoyama (1974) and yields, in addition to
the above-mentioned two species, Columbites
parisian us, Subcolumbites perrin ismith i andsome other indeterminable species. Columbites is common, but Subcolumbites is rare.
On the genera Metadagnoceras
and Keyserlingites
Metadagnoceras motoyoshiense, sp. nov. isa stratigraphically important species, because
species belonging to the genus Metadagnoceras have been known only from Spathian
formations as was stressed by Tozer (1965).
They are reported from British Columbia inwestern Canada, Nevada in western UnitedStates, Primo rye in eastern Russia, Albania,
Chios, Central Iran, Timor and Guizhou
Province of China (Tozer, 1965, 1972; Kum-~3I
14"32"EI

I SOOm I~jw:~n 2

U1

Figure 1. Geologic map of the Motoyoshi
District in the Southern Kitakami Massif, Northeast
Japan (Compiled and simplified from various
sources).
I: Upper Permian Toyoma Formation, 2-5:
Lower-Middle Triassic Inai Group (2: Hiraiso Formation, 3: Osawa Formation, 4: Fukkoshi Formation, 5: Isatomae Formation), 6: Pleistocene terrace
deposits, 7: fossil locality.

mel, 1969; Wang, 1978). Metadagnoceras
is described herein for the first time fromJapan.
Species belonging to the genus Keyserlingites have been reported from Siberia, Spitsbergen, Arctic region of and British Columbia,
Canada, Idaho in the USA, Southern Primorye, Himalayas, and Timor (Kummel, 1969;
Zakharov, 1968). They are treated as index
fossils of Spathian or late Olenekian age(Tozer, 1965,1971). There are, however,different proposals with regard to the ages ofK. dieneri (Mojsisovics) from the Himalayas,
which was once described as Durgaites
dieneri by Diener (1905), and of K. angustecostatus Welter described from Timor.
For example, Kummel (1969) assigned themto the lower Anisian. These two "Durgaites" type species differ from other species
in having more prominent ribs across the957.

Metadagnoceras and Keyserlingites from Kitakami

venter. A recent discovery of K. dieneri andits related species in Qinghai Province ofChina (Wang, 1985) may indicate that the ageof Keyserlingites from these three districts(Himalayas, Timor and China) is early
Anisian rather than Spathian, because they
are associated with many Anisian ammonoids
in Qinghai. If we accept the opinion of
Wang (1985), the age of the Keyserlingites
faunas of the Boreal Province, includingSouthern Primorye, should be considered to
be Spathian, whereas that of the "Durgaites"
type faunas in the Tethys Province should beconsidered to be early Anisian.
No species of the genus Keyserlingites has
so far been described from Japan, although
Bando (1968) listed K. sp. from the Oro
Formation of the Yakuno Group in Kyoto
Prefecture. Keyserlingites middendorjJi, towhich the Kitakami specimen is compared, isknown from late Olenekian beds of Siberia in
association with K. subrobustus, which is anindex species of the K. subrobustus Zone, theuppermost ammonoid zone in the Spathian of
Canada (Tozer, 1965, 1967).
Thus, the present discovery ofMetadagnoceras and Keyserlingites enables acorrelation of the Osawa Formation directlywith the type Spathian Stage in Arctic
Canada (Tozer, 1967), another importantreference section of the Spathian beds inwestern Canada (Silberling and Tozer, 1968),
and the upper Olenekian (Kiparisova andPopov, 1964) or Olenekian (Vavilov andLozovsky, 1970) in Siberia and Primorye.

Problems of zonal correlationof the Spathian Stage
Silberling and Tozer (1968) and Tozer(1981) divided the Spathian Stage in western
North America into three biostratigraphic
units, the lower Columbites Beds, the middle
Subcolumbites Beds and the upper Neopopanoceras haugi Zone, and they correlated
the Kazakhstanites pilaticus (= Olenikites
pi/aticus) and Keyserlingites subrobustus

231

Zones in the Arctic region with the first andthe last unit, respectively. However, opinions differ with regard to the biostratigraphic
subdivision of the Spathian and correlationof subdivided units. Kummel (1969) divided
the Spath ian into only two zones, the lower
Columbites Zone and the upper Prohungarites Zone. He correlated the K. subrobustus
Zone of Canada with his Prohungarites zone,
which contains Subcolumbites. On the other
hand, Zakharov (1978) regarded the K. subrobustus Zone as older than the beds with
Subcolumbites.
Concerning the correlation of the Spathianammonoid zones, it must be remembered thatthe genera Columbites, Subcolumbites andKeyserlingites have never been found in aconsecutive sequence in North America.
Columbites parisianus, which is an index
species of the Columbites Zone (Smith, 1932)or Columbites and Tirolites Beds (Silberling
and Tozer, 1968), has been only reported
from southeastern Idaho, Subcolumbites only
from western Nevada, and Keyserlingites only
from British Columbia and the Arctic regionof Canada. Although Kummel (1969) regarded C. parisianus to occur only from Idaho,
the species came to be known from Mangyshlak in Kazakhstan (Shevyrev, 1968), from thePrimorye (Zakharov, 1968) and from theOsawa Formation in the Southern Kitakami
Massif (Bando and Shimoyama, 1974). In
the latter two districts, the above-mentioned
three ammonoid genera occur in a successive
sequence. In the Primorye, Columbites is
associated with Keyserlingites and occurs in ahorizon below that of Subcolumbites (Zakharov, 1968; Zharnikova, 1985). In the
Kitakami Massif, however, Bando andShimoyama (1974) already reported, though
not specifically discussing its significance,that Columbites and Subcolumbites co occur
at many stratigraphic horizons in the Osawa
Formation with the exception of its lowermost and uppermost parts. The occurrenceof Keyserlingites there overlaps with thestratigraphic ranges of Columbites and Sub-

Masayuki Ehiro

232

columbites. Therefore, the ammonoid zonal
scheme of the Spathian should be studied
further in detail.

a
Systematic Description
10mm

Specimens described in this paper are kept
in the Iwate Prefectural Museum, Morioka(IPPM).
Order Ceratitida Hyatt, 1884
Superfamily Dinaritaceae Mojsisovics, 1882
Family Dinaritidae Mojsisovics, 1882
Subfamily Khvalynitinae Shevyrev, 1968
Genus Metadagnoceras Tozer, 1965

Type species: Metadagnoceras
Tozer, 1965

pulcher~

Figure 2. Suture lines of Metadagnoceras andK eyserlingites.
a: Metadagnoceras motoyoshiense Ehiro, sp. nov.,
IPPM 60015
b: Keyserlingites cf. K. middendorffi (Keyserling),
IPPM 60013

Metadagnoceras motoyoshiense, sp. nov.
Figures 2a; 3-2a, 2b, 3Material: - Two specimens; IPPM 60015(holotype) collected by M. Kumagai in 1991
and IPPM 60032 collected by M. Ehiro, 1992.
Diagnosis: - Metadagnoceras of a large
size, with a small umbilicus. The surface isornamented with slightly sinuous growthlines and spiral lirae.
Description: - One specimen (lPPM
60015) is a right side of the inner mould.
The shell is laterally compressed due totectonic deformation and also slightly
deformed to make an ellipse. The bodychamber, apparently complete, is about twothird of a whorl. The conch is involute andlenticular in outline, with slightly convex
sides and acutely rounded venter. Maximum
whorl width lies near the centre. It attains adiameter of 185 mm, in the deformed state,
and at the adoral end its height and umbilical
diameter are about 101 mm and 7 mm, respectively. The side of the body chamber isornamented with faint, slightly sinuous
growth lines and indistinct, coarse spirallirae. Another specimen (lPPM 60032) is a
fragmented inner mould of the body chamber.
The suture is not perfectly preserved asshown in Figure 2a. It has a narrow androunded (?) first lateral saddle, a large laterallobe with prominent denticulations, and arounded second lateral saddle.
Comparison: - On account of its
lenticular shell form and possession of spirallirae, the present new species is somewhat
comparable with Metadagnoceras pulcher(Tozer, 1965, p. 29, plate I, figs. Ila-d), but it
differs from the latter in being of larger size,in having an acute venter and in having asmaller umbilicus. Other species ofMetadagnoceras are also distinguished easily
from the present species for the same reasonsand by having no spiral lirae.
Occurrence and geological age: - Dark
gray shale of the middle part of the Osawa~ Figure 3. la-b: Keyserlingites cf. K. middendorffi (Keyserling), IPPM 60013, lateral (a) and ventral(b) views, x 0.64. 2a-b, 3: Metadagnoceras motoyoshiense Ehiro, sp. nov. 2, holotype (IPPM 60015),
lateral view (a) and cross section (b), x 0.64. Arrow marks the position of the cross section. 3. IPPM
60032, lateral view, x 0.64.

957.

Metadagnoceras and Keyserlingites from Kitakami

,

,
\

,

\. . j2b

233

Masayuki Ehiro

234

Formation exposed at Yamaya, Motoyoshicho, Motoyoshi-gun, Miyagi Prefecture;
Spathian Stage of the Scythian.
Etymology: - This species is named forMotoyoshi District, its type locality.
Superfamily Ceratitaceae Mojsisovics, 1879
Family Keyserlingitidae Zakharov, 1970
Genus Keyserlingites Hyatt, 1900
Type species: Ceratites subrobustus Mojsisovics, 1885
Remarks: - Six Spathian species (K.
meridian us, K. middendorjfi, K. miroshnikovi,
K. stephansoni, K. subrobustus and K. tebenkovi) and seven Anisian species (K. angustecostatus, K. dieneri, K. pahari, K. pagoda,
K. planus, K. qinghaiensis and K. sinensis)
are presently recognized in this genus. The
Anisian species have more prominent ventral
ribs than the Spathian species.
K. bearlakensis Kummel and K. bearriverensis Kummel described from Idaho (Kummel, 1969) are not assignable to the genus
Keyserlingites, because they lack such characteristic features of the genus Keyserlingites as
bituberculation and deeply incised externallobe (Tozer, 1971). In the same way, K.
bearriverensis and K. sp. described from theHimalayan region of China by Wang and He(1976) do not belong to Keyserlingites. In
addition, as already pointed out by Guex
(1978), Keyserlingites sp. of Kumme1 (1968)
described from Afghanistan may also notbelong to this genus.
Keyserlingites cf. K. middendorjfi(Keyserling)
Figures 2b; 3-1 a, 1b

Material: - One specimen, collected by
M. Numakura and A. Numakura in 1991,
IPPM 60013.
Description: - One incomplete specimen
is examined. The conch is evolute and discoidal in outline, with convex sides andacutely rounded venter. The shell is thought

to be essentially a compressed form, though
the compression may be partially due to its
lateral deformation. The conch attains adiameter of at least 125 mm with an umbilicus
measuring about 2/5 of the diameter. The
lateral areas near the umbilical shoulder bear
widely spaced nodes. As the heightincreases, the nodes grow into radial ribs
which extend from the umbilical to the ventral shoulder. There are five nodes or ribs on
the half volution. On the ventral shoulders
there are occasional and less prominent small
tubercles whose arrangement alternates with
the nodes.
The suture is partly preserved as shown in
Figure 2b. It is ceratitic and consists ofrounded lateral saddles and denticulated lateral lobes. The ventral lobe is unknown.
The second lateral saddle is larger than theother lateral saddles.
Comparison: -The compressed shell formand shell ornamentation of the present specimen are similar to those of the specimens of
Ceratites middendorjfi (Keyserling, 1845, p.
170, pI. 1, fig. 1, pI. 2, figs. 1,3; Mojsisovics,
1885, p. 153, pI. 6, fig. 2; 1886, p. 38, pI. 2,
figs. 12, 13, pI. 3, figs. la-c, pI. 20, fig. 10 ;1886, p. 47, pI. 4, fig. 1 described as C. schrenki; 1888, p.6, pI. 1, figs. 12, 13 described as
C. nikitini), all collected from Siberia. However, it differs from the Siberian species in theform of the suture and, except for the lastmentioned one, in the possession of a wider
umbilicus. Some specimens of Keyserlingites middendorjfi illustrated by Zakharov(1978, pI. 17, figs. 1, 3, 6-7), however, have arather large umbilicus nearly the same size asthat of the Kitakami specimen.
K. miroshnikovi Burij and Zharnikova (Zakharov, 1968, p. 129, pI. 24, fig. 2, pI. 25, figs.
2,3, text-fig. 31-c), K. meridianus (Zakharov,
1968, p. 128, pI. 24, fig. 1, text-fig. 31-b) andK. tebenkovi (Zharnikova, 1985, p. 34, pI. 3,
figs.3a-b, text-fig. lc) described from Southern Primorye are similar to the present specimen in having a compressed shell form andlarger second lateral saddle. However, K.

957.

Metadagnoceras and Keyserlingites from Kitakami

miroshnikovi is distinguished from the Kitakami specimen in the furcated form of the ribs
and in having a slightly smaller umbilicus, K.
meridianus has a larger umbilicus and lessconspicuous ribs compared with the Kitakami specimen, and K. tebenkovi differs inhaving a less compressed shell form and more
numerous nodes.
Occurrence and geological age: - Sameas those for Metadagnoceras motoyoshiense,
sp. nov.
Acknowledgements

I would like to express my appreciation toProfessor Tsunemasa Saito of Tohoku University for his critical reading of the manuscript. Thanks are also expressed to Messrs.
Miyuki Numakura, Atsushi Numakura and
Makoto Kumagai for making available their
specimens to this study, and to Messrs.
Eishiro Shoji and Hiroe Kuriyagawa foraccess to important specimens.
This study was supported in part by the
Grant-in-Aid for Scientific Research from the
Ministry of Education, Science and Culture(no. 02452063), and by the Science Research
Fund from the Saito Gratitude Foundation.
References
Bando, Y., 1968: Stratigraphic problems concerning
the newly occurred Lower Triassic ammonites
from the Kitakami Massif and the Maizuru Zone.
Mem. Fac. Educ., Kagawa Univ., pt. 2, no. 174,
p.I-7. (in Japanese with English abstract)- - and Ehiro, M., 1982: On some Lower Triassic
ammonites from the Osawa Formation at
Asadanuki, Towa-cho, Tome-gun, Miyagi Prefecture, Northeast Japan. Trans. Proc. Palaeont.
Soc. Japan, N.S., no. 127, p. 375-385.
- - and Shim oyama, S., 1974: Late Scythian ammonoids from the Kitakami Massif. Trans.
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Diener, c., 1905: Entwurf einer Systematik der Ceratitiden des Muschelkalkes. Sitz. Akad. Wiss.
Wien, vol. 104, p. 765-806.
Guex, J., 1978: Le Trias inferieur des Salt Ranges(Pakistan). Eclog. Geol. Helv., vol. 71, p. 105-

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141.
Keyserling, A., 1845: Beschreibung einiger von Dr.
A.H.v. Middendorf mitgebrachten Ceratiten desarktischen Sibiriens. Bull. A cad. Imp. Sci. St.
Petersbourg, vol. 5, p. 161-174.
Kiparisova, L.D. and Popov, Yu. N., 1964: Projectof the dividing of the Lower Triassic into stages.

Intern. Geol. Congr. XXII Session, Reports of
the Soviet geologists, theme 160, p.91-99.
Kummel, B., 1968: Additional Scythian ammonoids
from Afghanistan. BulL Mus. Compo Zool., vol.
136, p.483-509.
--,1969: Ammonoids of the Late Scythian (Lower
Triassic). Bull. Mus. Compo Zool., vol. 137, p.
311-702.
Mojsisovics, E., 1885: Ober die Structur des Sipho
bei einigen triadischen Ammoneen. Neues
Jahrb. Min. GeoL Palaeont., vol. I, p. 151-162.
- - , 1886: Arktische Triasfaunen. Beitriige zur
palaeontologischen Charakteristik der ArktischPacifischen Triasprovinz. Mem. Acad. Imp. Sci.
St.-petersbourg, Ser. 7, vol. 33, p. 1-159.
- - , 1888: Ober einige arktische Trias-Ammoniten
des nordlichen Sibirien. Mem. A cad. Imp. Sci.
St. -Petersbourg, Ser. 7, vol. 36, p. 1-21.
Shevyrev, A.A., 1968: Triassic ammonoids of south
USSR. Akad. Nauk SSSR, Trudy Paleont.
Inst., vol. 119, p.5-272. (in Russian)Silberling, N.J. and Tozer, E.T., 1968: Biostratigraphic classification of the marine Triassic in North
America. Geol. Soc. America, Spec. Paper 110,
63 p.
Smith, J.P., 1932: Lower Triassic ammonoids ofNorth America. U.S. Geol. Surv., Prof Paper
167, p. 1-199.
Tozer, E.T., 1965: Latest Lower Triassic ammonoids
from Ellesmere Island and northeastern British
Columbia. Geol. Surv. Canada, Bull. 123, p. 145.
- - , 1967: A standard for Triassic Time. Geol.
Surv. Canada, Bull. 156, p. 1-103.
- - , 1971: Triassic Time and ammonoids: Pro\:1lems and proposals. Can ad. Jour. Earth Sci.,
vol. 8, p.989-1031.
- - , 1972: Triassic ammonoids and Daonella fromthe Nakhlak Group, Anarak Region, Central
Iran. Geol. Surv. Iran, Rept., no. 28, p. 29-69.
- - , 1981 : Triassic Ammonoidea: Geographic andstratigraphic distribution. In, House, M.R. and
Senior, J.R. eds., The Ammonoidea,' the evolu-tion, classification, mode of life and geologicalusefulness of a major fossil group, p.397-431.
Academic Press, London.
Vavilov, M.N. and Lozovsky, V.R., 1970: To thequestion of the stage division of the Lower
Triassic. Izvest. Akad. Nauk. SSSR, ser. geol.,no. 9, p.93-99. (in Russian)Masayuki Ehiro

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Wang, Yikang, 1978: Latest Early Triassic ammonoids of Ziyun, Guizhou - with notes on therelationship between Early and Middle Triassic
ammonoids. Acta Palaeont. Sinica, vol. 17, p.
151-177. (in Chinese with English abstract)- - , 1985: Remarks on the Scythian-Anisian boundary. Riv. Ital. Paleont. Strat., vol. 90, p.515544.
- - and He, G., 1976: Triassic ammonoids from theMount lolmo Lungma Region. In, Report of
Scientific Expedition in the Mount Jolmo Lungrna Region (l966-1968)-Palaeontology, Pt. 3-, p.

223-438. Science Press, Beijing. (in Chinese)
Zakharov, Yu. D., 1968: Lower Triassic biostratigraphy and ammonoids in Southern Primorye
Region. Akad. Nauk SSSR, Siberian Sec., Far
Eastern Geol. Inst., 172 p. (in Russian)- - , 1978: Lower Triassic ammonoids of East
USSR, 224 p. Nauka, Moskva. (in Russian)Zharnikova, N.K., 1985: Ammonoids of the Lower
Triassic Columbites zone in the Southern Primoryeo Palaeont. Zhurnal, 1985, no. 2, p. 31-37. (inRussian)

Fukkoshi )Jd.~, Hiraiso 3jllill, Inai flli#, Isatomae Wllliii}, Motoyoshi *~, Oro *8, Osawa*~, Yakuno ~~!I!Y, Yamaya lll~.

l' r Metadagnoceras cI'J J: rJ Keyserlingites :-1 f 2:ri, Metadagnoceras motoyoshiense, sp. nov. :to J: 'Cf Keyserlingites cf. K. middendorffi(Keyserling) ~~1(:-t Go jili.j,~ C ~ bii~OOii' I? fl.1JJ(I)~1(:"t' ~ Go ,::,.n I? f1. Columbites ~Subcolumbites C ~£: L, A /{ - AWl (iiiJWl=~*c*Wl) (I)~*1tE-c'~ IJ, .:t(l)£:/:l:Hl.*~J!ii'A /{ - A ilf ~ G ~ 'fl.:;t v.? ilfl::$t::.)r.j"Jt ~.n G c ~, ? vt*(I)~J! ~Ji:fif-t G ~(I)"t' ~ G ~t*~tg§$(I) A /{ - Ailffl., Tfft J: IJ, Columbites J!, Subcolumbites J!:to J: 'Cf
Neopopanoceras haugi 'Ill't::.lK7t ~.n, 1&1ltfl. 7J T p(~!1!!~(I) Keyserlingites subrobustust::.)r.j" J:t ~.n -n' G L ii' L, ~tl::lll:ttB"t'fl. Columbites c Subcolumbites (I) v :/:;:; fl.~:l c Iv
cmfl L, ~ I? t::. Keyserlingites (I)£:I±lJ!* ~ ,::,.n I? 2 ~(I) v:/:;:;1*1 t::.~ G (I)-C', l::~(I) A/{ - Ailf(l)7:/'f= / -1 r1tE*IK7tfl.:P}~~~-tGo
7kJA~Z~tJ:.JlttI!!0)*5RJ!.imA/~-AM7 /.,;: /i¥i$~tl::lll:ttB(I)*~:ttB~t::.7t;ffl-t G r$=~**~J!(I)*$ii' I?£:-t G 7:/ 'f= /

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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 171, pp. 237-248,4 Figs., September 30, 1993

958. EARLY EVOLUTION AND DISTRIBUTION OF THE GASTROPOD
GENUS NUCELLA, WITH SPECIAL REFERENCE TO
MIOCENE SPECIES FROM JAPAN*
KAZUT AKA AMANO
Department of Geoscience, Joetsu University of Education,
Joetsu, Niigata Prefecture, 943 JapanGEERAT J. VERMEIl
Department of Geology, University of California,
Davis, California 95616, USA
and

KEN NARITA
Education board of Shinshu-shinmachi,
Nagano Prefecture, 381-24 Japan

Abstract. Several well preserved specimens of the muricid gastropod genus Nucellafrom Miocene strata in Hokkaido, Japan, are classified into two species, N. tokudai (Yokoyama) and N. freycineti saitoi Hatai et Kotaka. N. tokudai first appeared in California
during the Early Miocene, and then spread westward to Japan and Kamchatka by the early
Middle Miocene. N. freycineti saitoi first appeared in Japan during the Middle Miocene,
and gave rise to a living species, N. f freycineti (Deshayes) during the latest Miocene or
Early Pliocene. A warm-water origin in the northeastern Pacific is postulated for the genus
Nucella, based on the early geographical distribution of the genus and on the presence of a.
thick denticulated outer lip in all early species. The distributional history of Nucella,characterized by a warm-water origin in western North America and subsequent dispersal toeastern Asia, is similar to that of many other shallow-water temperate North Pacific molluscs
and barnacles.
Key words.

Miocene, Nucella, Gastropoda, evolution, biogeography.

northern Japan, most authors recognize two
Recent species, N. freycineti (Deshayes) and
N. heyseana (Dunker) (e.g. Habe and Ito,
1965). Mitochondrial DNA sequences showthat there are two species among the genus
Nucella in northern Japan (Collins et al., in
preparation). The small N. freycineti is living in the middle and upper intertidal zone,
and the larger N. heyseana (Dunker) is in thelower intertidal and shallow sublittoral zones

Introduction

The modern muricid gastropod genus
Nucella is commonly known and widelydistributed on the shore of the North Pacificand North Atlantic Oceans. Its members are
predators of barnacles (Cirripedia), mussels(Mytilidae), and limpets (Acmaeidae). In*Received March 24, 1993; accepted June 21, 1993
237

238

Kazutaka Amano, Geerat J. Vermeij and Ken Narita(Collins et al., in preparation). It is possible that N. elongata Golikov et Kussakin issynonymous with N. heyseana, but the limitsof the species of Nucella in the northwestern
Pacific are not well understood.
Little is known about the early evolution
and geographical distribution of Nucella.
The genus first appeared in the Early Mioceneof western North America (Loel and Corey,1932). Some time later, Nucella reached thenorthwestern Pacific, but the scarcity of adequate fossil material has made it difficult totreat the history of the genus Nucella in Asia
in detail. The earliest appearance of Nucellain the Atlantic was in the Late Pliocene of theNorth Sea Basin in Europe (Vermeij, 1991a,
1993; Collins et al., in preparation).
Nucella is therefore one of a large number ofcold-water animal and plant genera that
invaded the North Atlantic from the North
Pacific after Bering Strait opened during
middle Pliocene time (Durham and MacNeil,
1967; Vermeij, 1991a).
Previous authors have figured only three
specimens of Nucella from Miocene strata in
Hokkaido. Yokoyama (1932) described
Coralliophila tokudai from the Okada Beds(=early Middle Miocene Horoshin Formation?) in the Vryu coal-field, Northwest
Hokkaido. Mizuno et al. (1969) illustrated
Nucella ishii Vozumi (MS) from the Kushiro
coal-field in eastern Hokkaido. Finally,
Amano (1983) treated one poorly preserved
specimen of Nucella from the Late Miocene
Vpper Togeshita fauna, but he was unable toassign the shell at the species level. These
three specimens were not compared with eachother, nor were they compared with fossil orliving species found elsewhere in the North
Pacific region.
We have had an opportunity to examine
the holotype of Coralliophila tokudai and tocollect additional well preserved materials
belonging to Nucella from several Miocene
formations in Hokkaido. Our purpose ofthis paper is to describe this Miocene material
and to discuss the early evolution and geo-

graphical distribution of Nucella in the North
Pacific.

Localities of the Miocene Nucellain Hokkaido
Two species of Nucella were distinguishedamong specimens from seven localities ofMiocene age in Hokkaido (Figure 1). These
localities, together with stratigraphical andfacies details, are described briefly below.
Loc. R-1. Small riverside cliff near
Kasugacho-danchi, Rumoi, northwest
Hokkaido; medium-grained sandstone;
upper part of the Togeshita Formation.
Loc. R-2. Bank of Rumoi River near Owada(Rumoi-shinkawa, Loc. T24 of Amano,1983); pebble-bearing muddy finegrained sandstone; upper part of theTogeshita Formation.
Loc. Col. Riverside cliff at about 1.1 km
upstream of Tanzan-zawa, a small tributary of Chepotsunai River, Tomamaecho, northwest Hokkaido; pebblebearing fine-grained sandstone; Chikubetsu Formation.
Loc. V-I. Bank of Tokushibetsu River near
Kotobuki Bridge, Vtanobori-cho, northeast Hokkaido; muddy fine-grainedsandstone; Tachikaraushinai Formation.
Loc. A-I. Roadside cliff at about 5.2 km
upstream of Sarusarube River, Vrahorocho, eastern Hokkaido; muddy finegrained sandstone; Ishiizawa Formation.
Loc. A-2. River bank at about 1.6 km upstream of Ishii-zawa; pebble-bearing
muddy fine-grained sandstone; Ishiizawa Formation.
Loc. A-3. Streamside cliff at about 1.2 km
upstream of Tanzan-no-sawa; pebblebearing medium-grained sandstone;
Ishiizawa Formation.

G:

90

~

~

'"
(5

.'

u

~

g"
::s
~

~

Hokka

:::::
''""'
~

"

\

I
'J--+----"\

42'

Figure 1. Localities of Mi ocene Nucella in Hokkaido (using the topog raphica l maps of " Rum o i", " Sankei ", "Occhube" , " Atsunai " a nd"Ombetsu," sca le 1 : 50,000 published by Geographical Survey Institute of Japa n) ,

'"

w

'D

240

Kazutaka Amano, Geerat J. Vermeij and Ken Narita
Systematic description

Family Muricidae Rafinesque, 1815
Subfamily Ocenebrinae Cossmann, 1903
Genus Nucella Roding, 1798

Nucella tokudai (Yokoyama, 1932)
Figure 2-2a-b, 4a-b, 5, 7a-b, 8

Coralliophila tokudai Yokoyama, 1932, p. 235-236,
pI. 2, fig. 1.
Thais (Stramonita) carrizoensis Loel and Corey,
1932, p. 249-250, pI. 47, figs. 2, 3a-b.
Nucella packi (Clark) var. talea Stewart, 1946, p. 102,
pI. 17, fig. II.
Thais sp., Lutz, 1951, p. 392, pI. 18, figs. 2, 5.
Thais lima (Gmelin). Hall, 1958, pI. 9, figs. 7,10.
Thais packi Clark. Addicott, 1965, fig. 3R.
Nucella ishii Uozumi (MS). Mizuno et al., 1969, pI.
28, fig. 2.
Thais (Nucella) packi Clark. Addicott, 1970, p.8485, pI. 9, figs. 1-4, 19.
Nucella packi (Clark). Addicott, 1980, pI. I, fig. 6;
Gladenkov and Sinelnikova, 1990, p. 126-127, pI.
19, figs. 4, 16.

Type locality.-Gengoro-sawa, Numatacho, Hokkaido. CM no. 26003 (Figure 2-8).
Materials.- Two specimens from the Togeshita Formation (Loc. R-I), one specimen
from the Chikubetsu Formation (Loc. C-I),
and seven specimens from the Ishiizawa Formation (Loc. A-I, 2, 3).
Description.-Shell small to medium,heavy, bucciniform, with very low spire.
Whorls more than four. Suture poorly
defined. Surface ornamented with flattopped spiral ribs and weak growth lines.
Spiral ribs 15 to 19 on body whorl, 4 to 8 on
penultimate. Aperture large, ovate. Siphonal canal very short and deep. Outer lip
thick with 13 to 16 teeth.

Measurements (in mm).ShellheightHeight ofaperture

BreadthNS*

Localities

25.8

20.3

18

A-I

27.0

21.5

19.6

16

A-I

57.7

41.3

33.7

19

R-I

49.3

33.1

30.4

17

R-I

16

C-I

12.7

8.5

*number of spiral cords on body whorl.

Remarks.- The present species was originally described by Yokoyama (1932, August)from the Middle Okada Beds in Hokkaido
under the genus name Coralliophila. The
species has subsequently been mentioned by
Hatai and Nisiyama (1952), Uozumi (1962),
Ohara (1966) and Ohara and Kanno (1973).
N. tokudai is characterized by very low spire,
thick outer lip with numerous small teeth andnumerous flat-topped spiral ribs of evenstrength, which are separated by rather deep
grooves.
Loel and Corey (1932, December) proposed
Thais (Stramonita) carrizoensis from the
Early Miocene Vaqueros Formation in
California. The holotype of this species has
11 flat-topped spiral ribs on the body whorl.
Although this number of ribs is smaller than
the typical N. tokudai, all other characters of
T. carrizoensis conform with those of N.
tokudai. Moreover, according to Addicott(1970) and our own observations, other specimens of T. carrizoensis have more numerous
ribs. We therefore regard Loel and Corey's
species as a junior subjective synonym of N.
tokudai.
Addicott (1970) considered Thais carrizoensis to be a junior subjective synonym of~
Figure 2. la-b,3, lla-b, 12: Nucella freycineti saitoi Hatai et Kotaka. la-b: X I, Loc. R-I,
Togeshita Formation, IGUT no. 11801. 3; x2, Loc. U-I, Tachikaraushinai Formation, JUE no. 15402.
lla-b: X I, Loc. R-2, Togeshita Formation, JUE no. 15403. 12; X I, Loc. R-2, Togeshita Formation,
IGUT no. 11803. 2a-b, 4a-b, Sa-b, 7a-b, 8: Nucella tokudai (Yokoyama). 2a-b; X 2, Loc. C-I, Chikubetsu Formation, JUE no. 15404. 4a-b, Sa-b; X I, Loc. A-I, Ishiizawa Formation, JUE no. 15405.
7a-b; X I, Loc. R-I, Togeshita Formation, IGUT no. 11802. 8; X 2, holotype (CM no. 26003). 6,9:
"Nucella" tokishiensis Itoigawa et Shibata. 6; X 1.5, paratype (MFM no. 10078). 9; X 1.5, holotype(MFM no. 10076). 10: Nucella freycineti (Deshayes), X I, Loc. Rumoi, Recent, JUE no. 15406.

958.

Early evolution of Nucella

241

242

Kazutaka Amano, Geerat J. Vermeij and Ken Nadta

Thais packi, which was described by Clark(1918) from the Late Oligocene or earliest
Miocene San Ramon Formation in California. T. packi, which was assigned to Nucellaby Stewart (1946), is very similar to T. carrizoensis in shell outline and sculpture. The
holotype of T. packi, however, possesses asmall labral tooth near the base of the outerlip. This feature is also preserved on several
other specimens from the San Ramon Formation. The presence of this labral projection
indicates that T. packi belongs to either
Acanthina or Acanthinucella. Because of
the position of the spine and the nature of theshell sculpture, we tentatively assign thespecies to the genus Acanthinucella (for a
discussion of distinctions among Nucella,Acanthina, A can thin ucella, see Vermeij,
1993). Specimens assigned to T. packi fromother Miocene strata in western North America and Kamchatka by Addicott (1965, 1970,1980) and Gladenkov and Sinelnikova (1990)
lack the labral projection. They can therefore be assigned to N. tokudai.
Polytropa ishii was listed by Uozumi (1962)
as a representative species of the Miocene

Atsunai-Togeshita fauna, but Uozumi neither
illustrated nor described the material uponwhich this name was based. In their surveyof the fauna of the Atsunai area in eastern
Hokkaido, Mizuno et al. (1969) figured one
specimen from the "Atsunai Formation" (=Ishiizawa Formation) as Nucella ishii
Uozumi (MS). In our examination of specimens from this formation, we were unable todetect differences between N. ishii and N.
tokudai. Therefore, we regard N. ishii as a
junior subjective synonym of N. tokudai.
Comparison.- The present species is similar to the northeastern Pacific Recent species
Nucella canaliculata (Duclos), especially, tothe form described by Dall (1915) as thevariety compressa. The latter form has a low
spire and numerous flat-topped spiral ribs
with narrow interspaces. However, N.
canaliculata differs from N. tokudai by having fewer spiral cords on the body whorl (12to 13 in N. canaliculata as compared to 15 to
19 in N. tokudai), and by having generallythin outer lip without denticles on the innersurface.
Several species of "Nucella" or "Polytropa"

Pacific Ocean• Early lIiocene
Alliddle lIiocene• Late Mioceneo'-----'
SOOkm
Figure 3.

Distribution of Nucella tokudai (Yokoyama).

958.

Early evolution of Nucellahave been recorded from early Middle
Miocene deposits in Honshu, but none
belongs to Nucella. Among them, "Nucella"
tokishiensis (Figure 2-6, 9) was describedfrom the Nataki Conglomerate and the Shukunohora Sandstone in Gifu Prefecture by
Itoigawa and Shibata (1976). This species
resembles N. tokudai in having 16 to 17
flat-topped spiral cords on the body whorl,
but it differs by being more slender, by thepresence of 9 to 12 low axial ribs on the
penultimate whorl, especially by the presenceof a knob at the posterior end of the inner lip(see also Horikoshi, 1983). The inner-lip
knob is absent in nearly all species of Nucella; it is present only in the Californian N.
emarginata (Deshayes), a highly derived
member of the genus (Vermeij, 1993; Collins
et al., in preparation).
Distribution.-Early Miocene Vaqueros
Formation and Jewett Sand, California;
early Middle Miocene Chikubetsu and Middle Okada Beds, Hokkaido, and Kakert Suite,
western Kamchatka; Middle Miocene Temblor Formation, Sobrante Sandstone, California, and Astoria Formation and Sandstone of
Floras Lake, Oregon, and Ishiizawa Formation, Hokkaido; Late Miocene upper partof the Togeshita Formation, Hokkaido, andCierbo Sandstone, California (Figure 3).

Nucella freycineti saitoi Hatai
et Kotaka, 1959
Figure 2-la-b, 3, Iia-b, 12.

Nucella freycincti [sic] (Deshayes) saitoi Hatai andKotaka, 1959, p. 9-10, figs. 2,5.
Nucella sp., Amano, 1983, p. 33, pI. 8, fig. 25.
Thais lima (Martyn). Gladenkov et al., 1984, p. 245,
pI. 62, figs.4a-b

Type locality.-Okamami-zawa,
Obanazawa-machi, Yamagata Prefecture.
lOPS no. 77797.
Materials.-Four specimens from the Togeshita Formation (Loc. R-l, 2) and one specimen from the Tachikaraushinai Formation(Loc. U-1)
Description.-Shell large for the genus,

243

solid, bucciniform, with low spire. Whorls
more than four. Surface ornamented withround-topped spiral cords, weak growthlines; low secondary spiral riblets in broadinterspaces between cords; 18 to 20 cords
and riblets on body whorl, 3 to 8 on penultimate whorl. Aperture large, ovate. Siphonal canal narrow and short. Fasciole
rather prominent. Inner lip smooth, covered
with thin callus. Outer lip thick, with 8
denticles on its inner surface. These denticles distinct on young specimens, obsolete on
adults.
Measurements (in mm).ShellheightHeight ofaperture

BreadthNS'

Localities

26.9

19.9

19.1

20

R-I

52.1

36.4

36.9

19

R-2

53.9

37.4

20

R-2

37.3

37.8
29.6

28.9

8.5

8.6

18
18

R-2

12.1

U-I

'number of spiral cords and riblets on body whorl.

Remarks and comparison.- The present
subspecies was proposed by Hatai and Kotaka (1959) for material from the Middle
Miocene Oinzan Formation in Yamagata
Prefecture, Northeast Honshu. In theirpaper, they misspelled the specific name, as
freycincti. N. f saitoi is distinguished fromthe Recent N. f freycineti (Figure 2-10) and
N. heyseana by the thicker outer lip, narrower aperture, and by the presence of 8 denticles on the inner side of the outer lip. As
pointed out by Hatai and Kotaka (1959), theshell length: breadth ratio of Recent adultspecimens is greater than that of the fossil N.
f saitoi. The fossil species lacks the axial
sculpture often observed in N. heyseana. In
other respects, however, N. f saitoi is very
similar to the Recent N. f freycineti.
Amano (1983) described and illustrated
one poorly preserved specimen of Nucellafrom the Late Miocene upper part of theTogeshita Formation in Rumoi, northwest
Hokkaido, but did not assign the shell at the244

Kazutaka Amano, Geerat J. Vermeij and Ken Nadta

species level. Several well preserved specimens have been collected since from the samesite (Loc. R-2=T24 of Amano, 1983). These
conform in characters of sculpture and outer
lip to N. f saitoi.
Gladenkov et al. (1984) recorded Thais
lima (Martyn) from the Middle Miocene
Etolon Suite in western Kamchatka. Judging from their description and illustration, we

believe the material from Kamchatka should
be referred to Nucella Jreycineti saitoi.
Gladenkov et al. (1984) described it as having
14 to 15 round-topped spiral cords with someinterstitial riblets, and possessing a thickouter lip with some crenulations. The
figured specimen has a lower spire than does
the type of N. f saitoi, but, as spire heighttends to be a highly variable character within

Okhotsk Sea

Pacific Ocean
Japan Sea.. Iiddle liocene• Late liocene

Figure 4.

Distribution of Nucel/a freycineti saitoi Hatai et Kotaka.

958.

Early evolution of Nucellaspecies of Nucella, it is therefore unreliablefor distinguishing among species. TypicalNucella lima (Gmelin) has a thin outer lip
without denticles.
Distribution.-Middle Miocene Ginzan andTachikaraushinai Formations, northern
Japan, and Etolon Suite, western Kamchatka; Late Miocene upper part of the Togeshita Formation, Hokkaido (Figure 4).

Early evolution and migration of
Nucella in Miocene
The evolutionary origins of Nucella remain
obscure, but the available evidence indicatesthat the genus arose in the warm-temperate
northeastern Pacific. The earliest record of
Nucella is of N. tokudai from the Vaqueros
Formation and Jewett Sand (Early Miocene)in California (Loel and Corey, 1932; Addicott, 1965, 1970). N. tokudai is extremely
similar to Clark's (1918) Thais packi fromthe San Ramon Formation (Late Oligocene
or earliest Miocene) in California. Thais
packi, which we here tentatively assign to
Acanthinucella, differs from N. tokudai onlyby the presence of a small labral projection.
In view of this similarity, it is likely that thetwo species share a common ancestry.
The earliest northwestern Pacific records ofNucella are those of N. tokudai from the early
Middle Miocene in Hokkaido and Kamchatka. These records indicate that N. tokudai
spread westward from California to Japan by
the early Middle Miocene, which corresponds
to the time of the so-called climatic optimumof the Miocene. Hokkaido at this time lay
in a mild- to cool-temperate zone (Ogasawara, 1988). Many warm-water species extended
quite far north in the Pacific at that time
(Marincovich, 1984). Exactly when and how
the dispersal of N. tokudai occurred is notknown. Miocene records of Nucella arelacking in Alaska, possibly because thewarm-water populations of early Nucellawere not adapted to the cold conditions
postulated to have existed in parts of Alaska

245

during the early Middle Miocene by Marincovich (1990).
Several other shallow-water gastropod genera had invasion histories similar to that of
Nucella. These include Littorina (Reid,
1989, 1990) and the buccinid Lirabuccinum(Vermeij, 1991 b). These genera had earlier
records in western North America than ineastern Asia, and evidently crossed the North
Pacific westward by early Middle Miocene
time. The same may be true for several
genera of barnacles, including Chirona andHesperibalanus (Zullo and Marincovich,1990). Large rugose mussels of the genera
Plicatomytilus and Tumidimytilus alsoachieved a broad amphi-Pacific distribution
during the early Middle Miocene (Allison
and Addicott, 1976; Kafanov, 1987;
Uozumi and Akamatsu, 1988). Thus, by the
Middle Miocene, many of the shallow-waterorganisms with which Nucella lives and uponwhich it feeds were widespread throughout
the North Pacific.
The second species of Nucella to appear in
the northwestern Pacific was N. freycineti
saitoi, which is known from the Middle
Miocene Tachikaraushinai and Ginzan Formations in Japan and the Etolon Suite in
Kamchatka. Shibata et al. (1981) assigned
an age of 13.7-13.8 Ma to the Tachikaraushinai Formation by Kj Ar dating method.
N. f. saitoi persisted into the Late Miocene
upper part of the Togeshita Formation, whichcontains some molluscs in common withthose from the Pliocene Sannohe Group in
Northeast Honshu and the Plio-Pleistocene
Omma-Manganji fauna in the Japan Sea
Borderland (Amano, 1986). The oldest N.f.
freycineti was described by Chinzei (1961)
from the Togawa Formation of the SannoheGroup. It is likely that this form evolved
from N. f. saitoi during latest Miocene or
Early Pliocene time.
The Miocene species of Nucella from Japandiffer from their living counterparts in thenorthwestern Pacific by having an exceptionally thick outer lip whose inner side is246

Kazutaka Amano, Geerat J. Vermeij and Ken Narita

denticulate. An outer lip of this kind iscommonly seen in tropical gastropods and issuspected to have an anti predatory function(Vermeij, 1987). It is likely that Nucellaarose and spread in relatively warm waters,
where predators are abundant. Subsequently, several lineages of Nucella became adapted
to cooler waters, where the outer lip became
thinner and the denticulation was lost, as inN. f Jreycineti, N. heyseana, N. lima, andmost populations of N. canaliculata. However, other lineages of Nucella retained or
reacquired thick denticulate lips. Thisapplies to the northeastern Pacific N. lamellosa (Gmelin) and the north Atlantic N.
lapillus (Linnaeus). The Californian N.
emarginata has secondarily adapted towarmer waters and, at least in populations
from sheltered sites, has evolved a very thick,strongly denticulate lip (Collins et al., in
preparation).
The early evolution of Nucella illustrates
the important point that many shallow-waterelements of the present-day temperate North
Pacific fauna arose in relatively warm waters
and soon thereafter spread throughout theNorth Pacific Basin. Because most previous
studies of the fossil faunas of this region havebeen restricted to particular areas, comparisons of specimens from different parts of theNorth Pacific have often not been made. As
a result, the number of groups like Nucellawhose geographical distributions came toencompass large parts of both the eastern andwestern North Pacific is probably much larger
than the available literature now indicates.
Many more examples will doubtless come tolight when detailed comparisons are madebetween faunas of western North America
and eastern Asia.
Acknowledgements
We are grateful to Prof. Hiroshi Noda of
the University of Tsukuba, Prof. David Lindberg of the University of California at Berkeley, Emeritus Prof. Junji Itoigawa of

Nagoya University, Prof. Itaru Hayami and
Dr. Minoru Utada of the University ofTokyo, Mr. Yoshitsugu Okumura of the
Mizunami Fossil Museum, for their help inenabling us to examine type material andother fossils in their care. Prof. Noda andEmeritus Prof. Saburo Kanno of the University of Tsukuba provided much usefuladvice and helpful suggestions. Finally, we
thank Dr. Janice Cooper and Dr. Edith
Zipser of the University of California at
Davis for technical assistance.

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- - , 1970: Miocene gastropods and biostratigraphyof the Kern River area, California. Ibid., 642, p.
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molluscan faunas from the Uryu coal-field ofCentral Hokkaido. Sci. Rep., Tohoku Univ.,
Spec. Vol., no. 6, p. 125-135.
Reid.D.G., 1989: The comparative morphology,
phylogeny and evolution of the gastropod family
Littorinidae. Phil. Trans. Roy. Soc. London,
vol. 324, B1220, p. 1-110.
- - , 1990: A cladistic phylogeny of the genus Littorina (Gastropoda): Implications for evolution of
reproductive strategies and for classification.
Hydrobiologia, 193, p. 1-19.
Shibata, K., Yamaguchi. S., Ishida, M. and Nemoto,T., 1981: Geochronology of the Desmostylusbearing formation from Utanobori, Hokkaido.
Bull. Geol. Surv. Japan, vol. 32, no. 10, p. 545-

549. (in Japanese with English abstract).
Stewart, R., 1946: Geology of Reef Ridge, Coalinga
district, California. U.S. Geol. Surv., Prof.
Paper 205-C, p. 81-115, pIs. 9-17.
Uozumi, S., 1962: Neogene molluscan fauna in Hokkaido. Pt. 1. Sequence and distribution of
Neogene molluscan faunas. Jour. Fac. Sci.,
Hokkaido Univ., Ser. 4, vol. II, no. 3, p.507544.
- - and Akamatsu, M., 1988: Notes on the four
diagnostic Miocene mytilids of Hokkaido, North
Japan. Saito Ho-on Kai, Spec. Pub., no. 2, p.
325-338, pis. 1-5.
Vermeij. G.1., 1987: Evolution and escalation; an
ecological history of life. 527p. Princeton Univ.
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- - , 1991a: Anatomy of invasion: the trans-Arctic
interchange. Paleobiology, vol. 17, no. 3, p. 281307.
- - , 1991 b: Generic identity and relationships ofthe northeastern Pacific buccinid gastropod Searlesia dira (Reeve). Veliger, vol. 34, no. 3, p.
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Kazutaka Amano, Geerat J. Vermeij and Ken Narita

248- - , 1993: Spinucella, new genus of Miocene toPleistocene muricid gastropod from the eastern
Atlantic. Contr. Tertiary Quaternary Geol., inpress.
Yokoyama, M., 1932: Tertiary Mollusca from thecoal-field of Uryu, Ishikari. Jour. Fac. Sci.,

Imp. Univ. Tokyo, sec. 2, vol. 3, pt. 6, p. 227-247,
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Zullo, V.A. and Marincovich, L.Jr., 1990: Balanoidbarnacles from the Miocene of the Alaska Peninsula, and their relevance to the extant barnacle
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*

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Palaeontological Society of Japan (PSJ) Council Actions
During its meeting on June 26, 1993, the PSJ Council enacted the followingchanges to its membership.
New members elected:
Yuichi Asano,Nobuharu Hori,
Kenji Kashiwagi,
Masahiko Konomatsu,
Takahiko Nonaka,
Masahiko Takata,Y oshihiro Yamasaki,

Vladimur I. Davydov,
Y oshiaki Ishida,
Kaoru Kitao,
Toshihiro Minami,
Y ong Ho Shin,
Yasufumi Tsuchiya,
Toshiyuki Wakida.

Christopher John Hollis,
Takuya Itaki,
Toshifumi Komatsu,
Tokuji Mitsugi,
Norimichi Souji,
Atsushi Yabe,

New patron member:Museum of Nature and Human Activities, Hyogo.
Resigned members:(Ordinary member)
Mika Sato.
(Fellow)
Toru Onoe,
Saburo Yoshida.

Seiji Sato,

Tatsuya Yamasaki.

(Patron member)Social Education Division, Itoigawa Municipal Board of Education.

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

September 30, 1993

New Series No. 171

CONTENTS
TRANSACTIONS
956.

Katsumi Ueno and Hisayoshi Igo: Upper Carboniferous foraminifers from
Ban Na Din Dam, Changwat Loei, northeastern Thailand . ........ . . ..... 213

957.

Masayuki Ehiro: Spathian ammonoids Metadagnoceras and Keyserlingites.

from the Osawa Formation in the so uthern Kitakami Massif, Northeast Japan
958.

229

Kazutaka Amano, Geerat J. Vermeij, and Ken Narita : Early evolution anddistribution of the gastropod genus Nucella, with special reference to Miocene
species from Japan . .......... ..... . ... .... .. . ... ... . .... . .. . ... ..... 237

PROCEEDINGS........ . .... ... .. .. ..... . .... .. .. .. . .. ... . .... . . . .. . ... .. 249