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The fossil on the cover is an adult example (T. TAKAHASHI colI.) of
Mikasaites orbicularis MATSUMOTO (subfamily Marshallitinae, family
Kossmaticeratidae) from the Lower Cenomani;m (Cretaceous) of theMikasa area, central Hokkaido. (photo by M. NODA, natural size)

All communications relating to this journal should be addressed to the
PALAEONTOLOGICAL SOCIETY OF JAPAN,
c/o Business Center for Academic Societies,
Yayoi 2-4-16, I311nkyo-ku, Tokyo 113, japan.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 126, pp. 303-326, pis. 49-53, June 30, 1982

743. MIOSPORES FROM THE EOCENE NANGGULAN FORMATION
IN THE YOGYAKARTA REGION, CENTRAL JAVA*KIYOSHI TAKAHASHI
Department of Geology, Nagasaki University, Nagasaki 852

Abstract. The author has discriminated and descrided 48 palynomorph types
which consist of 4 spores and 44 pollen grains from the Eocene Nanggulan Formationin the Yogyakarta region, central Java. 17 species of 48 palynomorph types arenew: Laevigatosporites javanicus n. sp., Smilacipites spinulifer n. sp., Magnoliaepolienites ellipticus n. sp., Quercoidites ellipsodeus n. sp., Tricolpopolienites
elongatus n. sp., Brevi tricolpites circularis n. sp., Tricolporopolienites javanensis
n. sp., T. marginatus n. sp., T. rasus n. sp., Retitricolporites protensus n. sp.,
Striatricolporites striolatus n. sp., Polygalacidites speciosus n. sp., Graminidites
punctulosus n. sp., Proteacidites matsuokae n. sp., Subtriporopolienites minutulus n.
sp., Subtriporopollis specialis n. sp. and Tiliaepollenites tropicus n. sp.
The Nanggulan palyno-assemblage is characterized by main pollen grains ofDicolpopollis malesianus, Polygalacidites speciosus, Striatricolporites spp., Proteacidites spp., Graminidites punctulosus etc. and a comparative study with some
Eocene palyno-assemblages in other areas is made.

Graminidites punetulosus etc., which occur
relatively abundantly or show morphologically characteristic features, is conspicuously different from the Eocene palynofloras of Japan and some other areas.

Introduction
This paper is a report on spores andpollen grains from a lignite seam of the
Eocene Nanggulan Formation in the Yogyakarta region, central Java.
The lignite material was provided by
K. Matsuoka, member of the Overseas
Field Research (grant no. 504308; project
leader T. Saito), who has collected it near
Kalisongo, west of Nanggulan in the Yogyakarta region, central Java.
The author has distinguished 17 new
species and described 48 palynomorph types
which consist of 4 spores and 44 pollen
grains. This palyno-assemblage is characterized by main pollen grains of Dieolpopollis malesianus, Polygalaeidites speeiosus,
Striatrieolporites spp., Proteaeidites spp.,

Acknowledgements
The author thanks Dr. K. Matsuoka,
Associate Professor of Department of
Geology, Nagasaki University, for hiskind offering of the lignite material of
the Nanggulan Formation.
Thanks are also due to Professor Dr.
T. Saito, Department of Earth Sciences,
Faculty of Science, Yamagata University,
who is a project leader of the Overseas
Field Research entitled "Micropaleontology, Paleomagnetology and lithostratigraphy of Tertiary rocks, central Java",
for his permission to publish this paper.

* Received July 13, 1981; read October 4,1981 at Hiroshima.

303

Kiyoshi TAKAHASHI

304

Miss Akiko Nishi has assisted in preparing the text-figure.
Material and method

The examined lignite material was collected by K. Matsuoka from the lowest
part of the Nanggulan Formation in the
Kali Puru section near Kalisongo, westof Nanggulan, central Java.
The material was processed by mechanical and chemical methods (maceration by KCIO s and cone. HNO s, treatmentby 15% KOH and then by acetolysis
method, centrifuging and washing in purewater after each step). The residues
were mounted in glycerine jelly. All
slides were sealed with a manicure.
All specimens illustrated in this paper
and the material from which they wereobtained are in the palynological collectionof the Department of Geology, Faculty of
Liberal Arts, Nagasaki University.
Paleontological notes and palynological assemblage

According to Kadar's report (1981), theNanggulan Formation, which is subdivided
into two members, namely the lower
Kalisongo and the upper Kaliseputih
Members, consists of four biostratigraphic
units of foraminifera and Axinea. The
lignite seam belongs to the lowest Axinea
zone of the Kalisongo Member, which
consists mostly of shallow-marine sedimentary rocks. Based on a recent studyof planktonic foraminifera, the Nanggulan
Formation ranges from Middle Eocene to
Oligocene.
Okada (1981) found well-preserved nannofossils in the middle and upper parts of
the Nanggulan Formation and confirmed
correlation with the Middle Eocene Nannotetrina quadrat a Zone (CP 13) and Reticulofenestra umbilica Zone (CP 14).

Matsuoka (1981) made a list of dinoflagellate cysts and Pediastrum from the
Nanggulan Formation and stated that thiscyst assemblage is characterized by a lackof Wetzeliaceae and Deflandreaceae.
The miospores recovered from the examined lignite material indicate 48 palynomorph types which appear to be importantfor characterizing the Nanggulan palynoassemblage.
The fossil genera and species found are
as follows.
Trilete spores:
Leiotriletes maxoides Krutzsch minoris
Krutzsch
Alsophilidites kerguelensis Cookson
Monolete spores:
Laevigatosporites javanicus n. sp.
?Laevigatosporites sp.
Pollen grains:
Inaperturate:
Smilacipites spinulifer n. sp.
Smilacipites echinatus Wodehouse
Classites capucinii Gonzalez Guzman
Monocolpate:
Clavapalmaedites sp.
Magnoliaepollenites ellipticus n. sp.
Dicolpate:
Dicolpopollis malesianus Muller
Tricolpate:
Quercoidites umiensis (Takahashi)
Takahashi
Quercoidites ellipsodeus n. sp.
Tricolpopollenites elongatus n. sp.
Cupuliferoidaepollenites jallax (Potonie)
Potonie
Tricolpites minutireticulosus Takahashi
Tricolpate-Tetracolpate :Brevitricolpites circularis n. sp.
Tricolporate:
Cupulijeroipollenites sp.
Tricolporopollenites javanensis n. sp.
Tricolporopollenites marginatus n. sp.
Tricolporopollenites rasus n. sp.
Tricolporopollenites consularis Takahashi
consularis
Tricolporopollenites ovatorotundus
Takahashi

743.

Miospores from the Eocene Nangg~[lan Formation

Tricolporopollenites sp. a
Tricolporopollenites sp. b
Tricolporopollenites sp. c
Tricolporopollenites sp. d
Cyrillaceaepollenites minor (Takahashi)
Takahashi
Retitricolporites protensus n. sp.
llexpollenites tertiarius (Takahashi)
Takahashi
Striatricolporites striolatus n. sp.
Striatricolporites agustinus Gonzalez
GuzmanStriatricolporites sp.
Euphorbiacites sp.
Tetracolporate:
Tetracolporopollenites obscurus Pflug &ThomsonStephanocolporate:
Polygalacidites speciosus n. sp.
Monoporate:
Graminidites punctulosus n. sp.
Triporate:
Proteacidites mollis Samoilovitch
Proteacidites cf. annularis Cookson
Proteacidites simi lis HarrisProteacidites matsuokae n. sp.
Subtriporopollenites minutulus n. sp.
Subtriporopollenites sp.
Subtriporopollis specialis n. sp.
Triporopollenites sp. a
Triporopollenites sp. b
Tiliaepollenites tropicus n. sp.
Tiliaepollenites cf. punctulosus Takahashi?Tiliaepollenites sp.

There are only a few papers on Tertiary palynology of Southeast Asia. Of
48 palyhomorph types, 6 species are
assigned to the miospores previously described from Paleocene-Eocene sedimentsof Southeast Asia and the Southern Hemisphere.
Alsophilidites kerguelensis Cookson was,hitherto, described and illustrated fromthe Tertiary brown coal of the Kerguelen
Island and from the bright coal seam of
the Eocene Yaw Series of Kalewa in
Burma.
Classites capucinii Gonzalez Guzman and305

Striatricolporites agustinus Gonzalez Guzman are known from the Eocene Mirador
Formation of the Tibu area in Colombia,
South America. Botanical affinity of theformer is unknown, but it may be a coniferous pollen type, judging from its morphological characteristics.
The latter
somewhat resembles the Anacardiaceae
type, according to Gonzalez Guzman's
opinion. The author is, however, ofopinion that the latter rather resembles
the Cucurbitaceae (Gymnostemma or
Actinostemma) type.
Muller (1968) described and illustrated
Dicolpopollis malesianus :'v[uller and Dicolpopollis elegans Muller from the Proxapertites Zone (Paleocene?) to Retitriporites
variabilis Zone (Eocene) of the Plateau
Sandstone Formation of Sarawak, Malaysia.
The Dicolpopollis specimens of the Nanggulan Formation of Java can be assigned
to the species D. malesianus of Sarawak.
This species is one of representatives of
the Nanggulan palyno-assemblage.
Proteacidites annularis Cookson is known
from the Paleocene-Miocene formations ofAustralia and from the Paleocene-Oligocene formations of New Zealand. Proteacidites similis Harris was first describedfrom the Middle Paleocene Pebble Point
Formation, Dilwyn Bay, Victoria, Australia. Both species occur very rarely inthe lignite seam of the Nanggulan Formation.
Some tricolpate, tricolporate and triporate pollen grains occur in common
with the Paleogene and Miocene of Japanand the Miocene of Korea: Quercoiditesumiensis (Takahashi) Takahashi, Cupuliferoidaepollenites fallax (Potonie) Potonie,
Tricolpites minutireticulosus Takahashi,
Tricolporopollenites consularis Takahashi
consularis, Tricolporopollenites ovatorotundus Takahashi, Cyrillaceaepollenites
minor (Takahashi) Takahashi, Ilexpollenites tertiarius (Takahashi) Takahashi

Kiyoshi TAKAHASHI

306

1100

10S0

1060

1120

1140

W-~--~--------+----------r--------~-----6°

J

o

A

200km

L'~_ _~~-,'

Text-fig. 1.

Map showing the locality of the lignite seam of the
Nanggulan Formation (x).

and Tiliaepollenites d. punctulosus Takahashi.
Smilacipites echinatus Wodehouse which
belongs doubtfully to the genus Smilax,
is an American species from the Eocene
Green River Formation, Colorado, U. S. A.
Leiotriletes maxoides Krutzsch minoris
Krutzsch, Cupulzjeroidaepollenites fallax(Potonie) Potonie, and Tetracolporopollenites obscurus Pflug & Thomson are
Tertiary European species.
Proteacidites mollis Samoilovitch is oneof the Maestrichtian-Danian Siberian
species.
The genus Polygalacidites was established by Sah and Dutta (1966) as a monotypic
genus with type species Polygalacidites
clarus Sah & Dutta from the Umsawmat
bed of the Cherra Formation (Eocene or?
Paleocene) in Assam, India. The new
species Polygalacidites speciosus is one of
the most remarkable species of the Nanggulan assemblage. This species shows
strong resemblance to pollen grains of
Polygala (Family Polygalaceae) which IS
herbs.
The Nanggulan palyno-assemblage isrelatively similar to the Kalewa palynoassemblage from the Eocene coal in
Burma, in spite of the different species
constituting the assemblage. The Kalewa
assemblage with Dicolpopollis kalewensis
as a main constituent and consisting of
tricolpate, tricolporate, triporate andTetracolporopollenites
possesses somecommon characteristics with the Nanggulan assemblage.
Germeraad, Hopping and Muller (1968)
reported on the palynological zonation of
Tertiary sediments from tropical areas,
namely Colombia, Venezuela, Trinidad,
Nigeria and Borneo. However, there isno common species between the Nanggulan assemblage and the Tertiary assemblages of the tropical areas above mentioned, excepting Dicolpopollis malesianus
Muller (1968) from the Proxapertites Zone
(Paleocene?) to the Retitriporites variabilis
Zone (Eocene) of the Plateau Sandstone
Formation in Sarawak, Malaysia.
Descriptive palynology

Anteturma Sporites Potonie 1893

743. Miospores from the Eocene Nanggulan Formation
Turma Triletes Reinsch 1881 emend.
Potonie & Kremp 1954
Subturma Azonotriletes Luber 1935
Infraturma Laevigati Bennie & Kidston
1886 emend.

Potonie 1956

Genus Leiotriletes Naumova 1939
emend. Potonie & Kremp 1954
Type species: Leiotriletes sphaerot riangulus (Loose 1932) Potonie& Kremp 1954.
Leiotriletes maxoides Krutzsch
minoris Krutzsch
PI. 49, Fig. 1.
1962. Leiotriletes maxoides Krutzsch minoris
Krutzsch, Atlas, Lief. I, 16, Taf. 1,
Fig. 2-8.

307

1929-1931, Rpts-ser. A, 2, pt. 8, 136, pI.
16, fig. 69.
1956. Alsophilidites kerguelensis
Cookson,Potonie, Beih. Ceol. lb., 23, 14, Taf. 1,
Fig. 3.
1960a. Alsophilidites kerguelensis Cookson,
Potonie, Senck. let h., 41, 1/6, 459-460,
Taf. 1, Fig. 20.

Dimensions: Grain size 45-61 pm inequatorial diameter; exine 1-1.5 pm thick,laevigate; trilete mark straight or slightly
undulate, reaching almost to the equator.
Occurrence: Few.
Comparison: These grains belong to
Alsophilidites llerguelensis Cookson fromthe Tertiary brown coal of the Kerguelen
Island and from the bright coal belonging
to the Eocene Yaw Series of Kalewa III
Burma.
Botanical affInity: Unknown.

Turma Monoletes Ibrahim 1933
Dimensions: Grain size 50 pm in diameter; amb broadly rounded triangularwith convex sides; exine 1.5 pm thick,laevigate; trilete mark not extending to
the equator.
Occurrence: Rare.
Comparison: The present grain seems
to belong to Leiotriletes maxoides Krutzsch
minoris Krutzsch from the Miocene andOligocene of Germany.
Botanical affinity: Unknown.

Subturma Azonomonoletes Luber 1935
Infraturma Laevigatomonoleti
Dybova & lachowicz 1957
Genus Laevigatosporites Ibrahim 1933
Type species: Laevigatosporites vulgaris(Ibrahim 1932) Ibrahim 1933.
Laevigatosporites javanicus n. sp.
PI. 49, Figs. 5-6.

Genus Alsophilidites Cookson 1947
ex Potonie 1956
Type species: Alsophilidites kergllensis
Cookson 1947.
.-llsophilidites kerguelensis Cookson
PI. 49, Figs. 2-3.
1947. Trilites (AlsoPhilidites) kerguelensis
Cookson, B. A. N. Z. Ant. Res. Exped.

Description: Monolete spore; reniform
or elliptical in lateral view. Exine smooth,0.6-1 pm thick. Dehiscence moderately
long, straight or slightly arc. Grain size
49-51 pm X 32.5-42 pm .
Holotype: PI. 49, Fig. 5; grain size 49x
34 pm; exine 1 pm thick, laevigate; slide
GN 4654.
Occurrence: Rare.
Comparison: The present species isKiyoshi TAKAHASHI

308

morphologically similar to the European
species Laevigatosporites nutidus(Mamczar)
Krutzsch nutidus (Krutzsch, 1967, 149-150,
Taf. 53, Fig. 4-12), but the former canbe distinguished from the latter by themuch thinner exine.
Botanical aJjznity: Polypodiaceae.

?Leavigatosporites sp.
PI. 49, Fig. 4.

Description: Monolete spore; reniform
or bean-shape in lateral view. Dehiscence
long, arc. Exine 1 pm thick, 1.5 pm thickin dehiscence side, laevigate except finely
punctate sculpture in dehiscence area.
Grain size 55.3 X 29.5 pm.
Occurrence: Very rare.
Remark: This specimen with the finely
punctate sculpture only in the dehiscencearea was very rarely found. This species
belongs doubtfully to the genus Laevigatosporites.
Botanical affinity: Polypodiaceae.

finely granulate or punctate sculptures andsharp conical spines somewhat densely orsparsely arranged, 0.5-1.5 pm in length,
apparently folded and deformed by pressure. Grain size 30-33 pm X 25-26 pm indiameter.
Holotype: PI. 49, Fig. 7; grain size 33x25.5 pm in diameter; exine thin, finely
punctate and sparsely echinate, 0.5-1 pm
in length; slide GN 4654.
Occurrence: Rare.
Comparison: Wodehouse (1933) established the genera Smilacipites and Peltandripites with large or small spines and/or
granules. The genus Smilacipites is muchsmaller in size than the Peltandripites.
The present specimens belong apparently
to the genus Smilacipites and are similar
to Smilacipites molloides W odehouse (1933,
p. 500, fig. 25) from the Eocene ParachuteCreek Member of the Green River Formation, Colorado, U. S. A., however this new
species is much smaller than S. molloides.
Botanical affinity: Smilax.
Smilacipites echinatus Wodehouse

Anteturma Pollenites Potonie 1931
PI. 49, Figs. 10-12.

Turma Aletes Ibrahim 1933
Subturma Azonaletes Luber 1935
emend. Potonie & Kremp 1954
Infraturma SUbpilonapiti Erdtman 1947
emend. Vimal 1952
Genus Smilacipites Wodehouse 1933
Type species: Smilacipites echinatus
W ode house 1933.
Smilacipites spinulifer n. sp.
PI. 49, Figs. 7, Sa-b.

Description: Inaperturate pollen grains;
ellipsoidal or originally spherical in form.
Exine thin, 0.5 pm± thick, provided with

1933. Smilacipites echinatus Wodehouse, Bull.
Torr. Bot. Club, 60, 500, fig. 27.

Dimensions: Grain size 22.5-27 pm X
16.6-21.5 pm in diameter; exine thin,smooth, provided with sharp spines irregularly arranged, varying from 1.7 pm
to 3.5 pm in length, from 1 pm to 2.5 pm
in width at base.
Occurrence: Few.
Remarks: The present specimens seemto belong to Smilacipites echinatus Wodehouse from the Eocene Parachute Creek
Member, Green River Formation, Colorado,
U. S. A. The grain of the Figure 10 possesses more numerous spines than othergrains. Wodehouse found two specimens
of Smilacipites echinatus and described

743.

Miospores from the Eocene Nanggulan Formatwnthat one of these shows the granularcharacter rather distinctly while in theother it is absent. The present specimensshow no granular character.
Botanical affinity: Belonging to thegenus Smilax is doubtful.

309

Genus Clavapalmaedites Rao& Ramanujam 1978
Type species: Clavapalmaedites
hammenii Rao & Ramanujam 1978.

Clavapalmaedites sp.
Infraturma Circumpollini (Pflug 1953)
Klaus 1960
Genus Classites Gonzalez Guzman 1967
Type species: Classites capucinii
Gonzalez Guzman 1967.

Classites capucinii Gonzalez Guzman
PI. 50, Figs. 1-3.
1967. Classites capucinii Gonzalez Guzman,
E. J. Brill, 62, pI. 30, figs. 2-2a.

Dimensions: Grain size 24.4-30 pm X
24-27 pm in diameter; size of ring 20(?)28 pm X 20(?)-23 pm in diameter; exine
0.5-0.8 pm thick; psilate; exine of ring
1 pm thick (ektexine: endexine=l : 1); a
reduced scar not visible on the proximalside.
Occurrence: Few.
Remarks: The grains seem to belong
to Classites capucinii Gonzalez Guzman
from the Eocene Mirador Formation (pollen zone IlIa) of the Tibu area, Colombia,
South America. A ring on one hemisphere of the grain exist distinctly, but
a reduced scar on one hemisphere is not
visible. The Figure 3 belongs very doubtfully to Classites capucinii.
Botanical affinity: Unknown.
Turma Plicates Naumova 1937
emend. Potonit~ 1960
Subturma Monocolpates Iversen& Troels-Smith 1950

PI. 49, Fig. 9.

Description: Monosulcate pollen grain;oval in distal polar view. Sulcus long,narrow, extending all along long axis ofgrain. Exine 1 pm thick; baculate andclavate, 1.3 pm in length. Grain size 25.5
X 22 pm in diameter.
Occurrence: Very rare.
Comparison: This grain is comparablewith that of Clavapalmaedites hammenii
Rao & Ramanujam (1978, p. 414) from the
Miocene strata of India. However, theformer differs from the latter in having
baculate and clavate sculptures.
Botanical affinity: ?Palmae.
Genus Magnoliaepollenites Nagy 1969
Type species: Magnoliaepollenites
simplex Nagy 1969.

Magnoliaepollenites ellipticus n. sp.
PI. 49, Figs. 13-15.

Description: Heteropolar, monocolpate
pollen grains; ellipsoidal to elongate ovalin distal polar view. Colpus long, strong,straight or somewhat arc. Exine 2-2.5
pm thick; ektexine: endexine=l : 1;
ektexine finely intrabaculate or intrarugulate, endexine chagrenate. On the surface
of exine very small pores relatively sparsely visible. Grain size 46.5-55 pm inlength; 25-32.5 pm in width.
Holotype: PI. 49, Figs. 15a-b; grain size
50 X 32.5 pm ; exine 2.5 pm thick (ektexine :endexine=l : 1); ektexine intrabaculate,Kiyoshi TAKAHASHI

310

endexine chagrenate; very small poresvisible on the surface of ektexine; slide
GN 4656.
Occurrence: Few.
Comparison: Nagy (1969) first established the new genus Magnoliaepollenites
and in the next year Krutzsch (1970) proposed the genus Magnolipollis. However,
the author recognizes that the latter is ajunior synonym of the former.
This new species differs from Magnoliaepollenites simplex Nagy (1969, p. 399,
pI. 41, figs. 1, 4) in form and structure ofexine. Magnoliaepollenites (al. Magnolipollis) magnolioides Krutzsch n. comb.,
Magnoliaepollenites (al. Magnolipollis)
micropunctatus Krutzsch n. comb., andMagnoliaepollenites (al. Magnolipollis)
neogenicus Krutzsch n. comb. neogenicus
differ in much thinner exine and structureof exine.
Botanical affinity: ?Magnoliaceae.
Subturma Dicolpates Erdtman 1947
Genus Dicolpopollis Pfianzl 1956
emend. Potonie 1966
Type species: Dicolpopollis lwckelii
Pfianzl 1956.

Dicolpopollis malesianus Muller
PI. 50, Figs. 4-10.
1968. Dicolpopollis malesianus Muller, Micropaleontology, 14, 1, p. 13, pI. 5, fig. 5.

Dimensions: Grain size 29-36 pm in·width, 20.5-27.5 pm in height; exine up
to 1.5 pm thick; columellae small, arranged in a reticulate pattern, forming
baculate to clavate muri; lumina variable
in size and shape, generally larger incentral zone, up to 1.5 pm.
Occurrence: Abundant.
Comments: Potonie (1960a) described

Disulcites kalewensis Potonie (464-466,
Taf. 2, Fig. 27-43; Abb. 3-4) from the
Eocene coal of Kalewa in Burma and he(1966) considered the genus Disulcites asa junior synonym of the genus Dicolpopollis and gave the emended diagnosis.
Dicolpopollis malesianus Muller is closely
similar to Dicolpopollis kalewensis(Potonie)
Potonie, but differs apparently in more
remarkable sculpture.
Muller (1968) found that Dicolpopollis
malesianus and Dicolpopollis elegans occur
from the Proxapertites Zone (Paleocene?)to the Retitriporites variabilis Zone
(Eocene) of the Plateau Sandstone Formation in Sarawak, Malaysia.
Botanical affinity: Palmae-Calamus.
Subturma Triptyches Naumova 1939
Genus Quercoidites Potonie, Thomson& Thiergart 1950 ex Potonie 1960
Type species: Quercoidites henrici(Potonie 1931) Potonie 1960.

Quercoidites umiensis (Takahashi)
Takahashi
PI. 50, Figs. 11-13, 30.
1957. Tricolpopollenites umiensis Takahashi,Mem. Fac. Sci., Kyushu Univ., Ser. D,
Geol., 5,4, 217, Taf. 38, Fig. 37-39;
Taf. 39, Fig. 31-32.
1979. Quercoidites umiensis (Takahashi) Takahashi, Takahashi & Kim, Palaeontographica, B, 170, 38, pI. 9, figs. 3-5, 24.

Dimensions: Grain size 27-34.2 pm X 1919.5 pm; exine 1-1.5 pm thick, intrabaculate; breadth/length ratio 0.56-D.72.
Occurrence: Rare.
Remarks: Morphologically the present
specimens appear to be closely comparable
to those of Quercoidites umiensis (Takahashi) Takahashi from the Paleogene and743. Miospores from the Eocene Nanggulan Formation
Miocene formations of Japan and from
the Miocene formations of Korea.
Potonie (1960a) bescribed Quercoiditessp. (grain size 20-30 pm), which is comparable with the present species, fromthe Eocene coal of Kalewa in Burma.
Botanical affinity: Cupuliferae.

311

form, size, and ornamentation of exine.
Botanical affinity: Cupuliferae.
Genus Cupulzjeroidaepollenites Potonie,
Thomson & Thiergart 1950
ex Potonie 1960

Quercoidites ellipsodeus n. sp.

Type species: Cupuli/eroidaepollenites
liblarensis (Thomson in Potonie,

PI. 50, Figs. 16-17.

Thomson & Thiergart 1950)
Potonie 1960.

Description: Tricolpate pollen grains;
elliptical to oval in equatorial view. Outline weakly crenate or finely wavy. Extremities of grain rounded or somewhat
broadly rounded. Three colpi slender,
more or less parallel, almost reachingpoles. Exine 1-2 pm thick, somewhat
thicker in the pole area, intrabaculate;ektexine: endexine=2 : 1. Grain size 3036 pm X 22-24 pm. Breadth/length ratio
0.67-0.74.
Holotype : PI. 50, Fig. 16; grain size
35.5x24 pm; exine 2 pm thick, intrabaculate; ektexine: endexine=2 : 1 ; surface
of exine uneven; breadth/length ratio
0.68; slide GN 4656.
Occurrence: Few.
Comparison: This new species can be
distinguished from Quercoidites henrici{Potonie) Potonie, Thomson & Thiergart(Potonie, 1931, p. 329, pI. 2, fig. 19;
Potonie, Thomson & Thiergart, 1950, p.
54, pI. B, figs. 22-23) in having slender
col pus and rough ornamentation of exine
surface and from Quercoidites microhenrici{Potonie) Potonie, Thomson & Thiergart(Potonie, 1931, p. 26, pI. 1, fig. V 19c;
Potonie, Thomson & Thiergart, 1950, p.
55, pI. B, figs. 24-25) in general form,size and ornamentation.
Quercoidites lusus Sah (1967, p. 51-52,
pI. 5, figs. 2-3) from the Upper Neogene
strata in Rusizi valley, Burundi, Africa,differs from the present species in general

Cupuli/eroidaepollenit'es /allax(Potonie) Potonie
PI. 50, Figs. 19-24.
1934. Pollenites fallax Potonie, Arb. Inst.
Palaeobot. Petrogr. Brennst., 4, p. 70,
fig. 10.
1953. TricoZpopolienites ZibZarensis (Thomson)
Thomson & Pflug fallax (Potonie)
Thomson & Pflug, PaZaeontographica,B, 94, S. 97, Taf. 11, Fig. 133-15l.
1960a. CupuZiferoidaepolienites fallax (Potonie) Potonie, Senck. Zeth., 41, 1/6, S.
468, Taf. 2, Fig. 64-65; Abb. 6.

Dimensions: Grain size 15-17.5 pmx8.5-10.5 pm; exine thin, 0.4-0.8 pm thick,laevigate or faintly chagrenate; Fig. 22exine 0.8 pm thick.
Occurrence: Common.
Remarks:
These specimens belong
clearly to Cupuli/eroidaepollenites /allax(Potonie) Potonie.
Botanical affinity: Cupuliferae.
Genus TricolpopoUenites Pflug& Thomson 1953
Type species:

Tricolpopollenites

pannularis (Potonie 1934) Thomson& Pflug 1953.
Tricolpopollenites elongatus n. sp.

Kiyoshi TAKAHASHI

312
PI. 50, Figs. 14-15.

Description: Tricolpate pollen grains;prolate or perprolate shape in equatorial
view. Meridional furrows (col pi) extending from pole to pole. Extremities ofgrain sharply rounded. Exine 1-1.5 pm
thick, chagrenate or/and punctate. Grainsize 38-43 pm X 16-21 pm. Breadth/length
ratio 0.38-0.55.
Holotype: PI. 50, Fig. 14; grain size
38.5 X 21 pm; exine 1 pm thick, chagrenate
to punctate; breadth/length ratio 0.54;
slide GN 4657.
Occurrence: Few.
Comparison: The present species isapparently similar to Quercoidites ellipsodeus Takahashi n. sp., but differs from thelatter in having larger size, more elongate
shape, and different structure of exine.
Botanical affinity: Unknown.

Genus Tricolpites Cookson 1947 ex
Couper 1953 emend. Belsky,
Boltenhagen & Potonie 1965
Type species: Tricolpites reticulatus
Cookson 1947 ex Couper 1953.
Tricolpites minutireticulosus Takahashi
PI. 50. Figs. 18, 25-27.
1979. Tricolpites minutireticulosus Takahashi,
Takahashi & Kim, Palaeontographica,B, 170, p. 40, pI. 10, figs. 5-8.

Dimensions: Grain size 15-21 pm X 12.5
-17.5 pm (Fig. 18-22.5 pm in diameter);exine 0.5-0.9 pm thick (fig. 18-1 pm thick),
intrabaculate; surface of exine very finely
reticulate (lumen less than 1 pm in diameter); breadth/length ratio 0.73-0.83.
Occurrence: Few.
Comparison: Morphologically the present specimens appear to be closely comparable to those of Tricolpites minutire-

ticulosus Takahashi from the Miocene
Changgi and Yonil Groups in the Yeoungill Bay district, Korea.
Botanical affinity: Salicaceae or Cruciferae.

Genus Brevitricolpites Gonzalez
Guzman 1967
Type species: Brevitricolpites variabilis
Gonzalez Guzman 1967.
Brevitricolpites circularis n. sp.
PI. 50, Figs. 28-29.

Description: Tricolpate or tetracolpate
pollen grains; spherical or spheroidal inpolar view. Three or four colpi very
short but distinct, 2-3 pm deep in polar
view, extending one third (or less) theradius. Exine thin, 1 pm or less thick,somewhat coarsely punctate. Grain size
15-17 pmX14-16 pm in diameter.
Holotype: PI. 50, Fig. 28; grain size 16.8
X 14.5 pm in diameter; colpi three, very
short; exine 1 pm thick, somewhat coarsely punctate; slide GN 4653.
Occurrence: Few.
Comparison: This new species can be
distinguished from Brevitricolpites variabilis Gonzalez Guzman with tricolpate totricolporate feature (1967, p. 35, pI. 12,
figs. 6-6b) from the pollen zones I-IV of
the Eocene Upper Los Cuervos and Mirador Formations, Colombia, in ornamentation of exine.
Tricolpites pachyexinus Couper withthree or four colpi (1953, p. 62, pI. 8, figs.
120-121) from the Upper Cretaceous Lower
Ohai Group, New Zealand, differs fromthe present species in having longer colpi
and smooth exine.
Tricolpites sp. A (Drugg, 1967, p. 49, pi.
7, fig. 37) from the Maestrichtian Marca
shale and the Danian Lower Dos Palos743.

Miospores from the Eocene Nanggulan Formation

shale at Escarpado Canyon in California
is similar to this new species, but theformer differs from the latter in having
larger size and smooth exine.
Botanical affinity: Unknown.

Subturma Ptychotriporines
Naumova 1939
Genus Tricolporopollenites
Pflug & Thomson 1953
Type species: Tricolporopollenites
dolium (Potonie 1931) Thomson& Pflug 1953.

Tricolporopollenites javanensis n. sp.
PI. 51, Figs. 1-5.

313

PI. 51, Figs. 7-9.
1961. Tricolporopollenites consularis Takahashi, Mem. Fac. Sci., Kyushu Univ., Ser.
0, Geol., 11, 3, p.323, pI. 24, figs. 55-56(pro parte).
1979. Tricolpolporopollenites consularis Takahashi subsp. consuiaris, Takahashi &Kim, Palaeontographica, B, 170, p. 41,
pI. 10, figs. 9-27; pI. 11, fig. 2.

Dimensions: Grain size 30-37 pmx2023 pm; exine intrabaculate or intrarugulate, 1-1.2 pm thick; breadth/length
ratio 0.56-0.66.
Occurrence: Few.
Comparison: These grains seem tobelong to those of Tricolporopollenites consularis Takahashi consularis from the
Paleogene and Miocene formations of WestJapan and from the Miocene Changgi and
Yonil Groups of Korea.
Botanical affinity: ?Cupuliferae.

Description: Tricolporate pollen grains;prolate to subprolate in equatorial view.
Three colpi extending from pole to pole.
Caverna relatively deep. Equatorial poresround, large, on a certain occasion withshort rugae. Exine smooth or weakly
chagrenate, 0.5-2 pm thick. Grain size
30-42 pm X 21-29 pm. Breadth/length ratio
0.55-0.78.
Holotype: PI. 51, Fig. 1; grain size 40.5
x22.5 pm; exine 1 pm thick, weakly chagrenate; pores round, large; breadth/length ratio 0.56; slide GN 4656.
Occurrence: Common.
Comparison: The present species issimilar to Tricolporopollenites (al. Cupuliferoipollenites) prolongatus (Takahashi) n.
comb. from the Miocene Cheonbug Conglomerate of the Y onil Group, Korea, butdiffers from the latter in size and poreshape.
Botanical afJinity: Unknown.

Description: Tricolporate pollen grains;
oval to subcircular in equatorial view;
circular to oval in polar view. Polar axis
longer than equatorial axis. Three colpi
parallel or converging to the poles, almost
reaching the poles; each with a somewhat
large and round pore. Exine punctate or
intrabaculate, 1-1.5 pm thick. Grain size
23-27 pm X 19-26 pm. Breadth/length ratio
0.83-0.96.
Holotype: PI. 51, Fig. 11; grain size 24
X 21.5 pm ; exine 1 pm thick, intrabaculate ;pore large, round; breadth/length ratio
0.895; slide GN 4655.
Occurrence: Common.

Tricolporopollenites consularis Takahashi subsp. consularis

Comparison: Tricolporopollenites marginatus is similar to Tricolporopollenites
emarginalis Takahashi (Takahashi & Kim,
1979, p. 41, pI. 11, figs. 11-22) from the

Tricolporopollenites marginatus n. sp.
PI. 51, figs. 10-14.

Kiyoshi TAKAHASHI

314

Miocene Changgi and Yoni! Groups of
Korea, but can be distinguished from thelatter in its comparatively larger poresand more spherical form.
Botanical affinity: Unknown.

Tricolporopollenites rasus n. sp.

Comparison: Only one specimen was
found. This specimen is comparable withthe grain of Tricolporopollenites ovatorotundus Takahashi from the lower coalbearing formation of the Changgi Group,
Korea.
Botanical affinity: Unknown.

PI. 51, Figs. IS-20.

Tricolporopollenites sp. a

Description:
Tricolporate
pollengrains; prolate to sub prolate in equatorial
view. Three colpi narrow, converging to
the poles, extending from pole to pole,with meridionally elongated pores. Exine
laevigate to faintly chagrenate, 0.5 fjm
thick. Grain size 15-19 fjm X 11-14 fjm.
Breadth/length ratio 0.71-0.77.
Holotype: PI. 51, Fig. 19; grain size 19
X 13.5 fjm; exine 0.5 fjm thick, laevigate;
pores meridionally elongated; breadth/length ratio 0.71; slide GN 4656.
Occurrence: Few.
Comparison: This new species is closely related to Tricolporopollenites (al.
Cupulzferoipollenites) pseudopusillus (Takahashi) Takahashi n. comb. (1979, p. 45, pI.
13, figs. 6-7) from the Middle Miocene
Pohang Formation of the Yoni! Group,
Korea, but the exine in the latter is thicker and its shape is much slender.
Botanical affinity: Unknown.

PI. 51, Figs. 15a·b.

Tri colporopollenites ovat orotundus
Takahashi
PI. 51, Fig. 24.
1979. Tricolporopollenites ovatorotllndlls Ta·
kahashi, Takahashi & Kim, Palaeonto.
graphica, B, 170, p. 42, pI. 12, fig. 2S32.

Dimensions: Grain size 22 fjm in diameter; exine 1.5 fjm thick, intrabaculate;equatorial pores small, round.
Occurrence: Very rare.

Description: Tricolporate pollen grains;subprolate in equatorial view. .Three
colpi comparatively narrow, extending
from pole to pole, with short rugae.
Cavern a not so deep. Exine intrabaculate,
1.5 fjm thick in the pole area, 0.8 fjm thickin equatorial area; ektexine: endexine=
2 : 1. Grain size 22.5x 17.5 fjm Breadth/length ratio 0.78.
.occurrence: Very rare.
Remarks: Only one specimen was
found.
Botanical affinity: Unknown.
Tricolporopollenites sp. b
PI. 51, Fig. 17.

Description: Tricolporate pollen grain;prolate in equatorial view. Three col piparallel, almost reaching the poles.
Exine chagrenate, 2 fjm thick, with somewhat meridionally elongated pores; Grainsize 24x 16 fjm. Breadth/length ratio 0.67.
Occurrence: Very rare.
Comparison: The present specimen issimilar to Tricolporopollenites (al. Cupuli/eroipollenites) ovuli/ormis (Takahashi) n.
comb. (Takahashi & Kim, 1979, p. 44, pI.
12, figs. 8-24) from the Miocene Changgi
and Yoni! Groups of Korea, but theformer differs from the latter in having
thicker exine and small rugae. However,
the author can not determine its species
name.

743.

Miospores from the Eocene Nanggulan Formation

Botanical affinity: Unknown.
Tricolporopollenites sp. c
PI. 51, Fig. 28.

Description: Tricolporate pollen grain;
spheroidal in equatorial view. Three colpi
parallel, not so long; caverna comparatively deep. Equatorial pores meridionallyelongated. Exine 1 pm thick, intrabaculate. Grain size 24x 21.5 pm. Breadth/length ratio 0.895.
Occurrence: Very rare.
Remarlzs: Only one specimen was
found.
Botaeical affinity: Unknown.

relatively narrow, almost reaching thepoles. Caverna not deep. Equatorial
pores large, equatorially elongated (rugae).
Exine 1.3 pm thick, smooth. Grain size
35.5 X 29 pm. Breadth/length ratio 0.82.
Occurrence: Very rare.
Remarl<s:
This specimen possesses
three large rugae and smooth exine. Accordingly, this belongs undoubtedly to thegenus Cupulzferoipollenites.
Botanical affinity: Unknown.
Genus Cyrillaceaepollenites Miirriger& Pflug 1951 ex Potonie 1960
Type species: Cyrillaceaepollenites
megaexactus (Potonie 1931)

Tricolporopollenites sp. d

Potonie 1960.

PI. 51, Fig. 16.

Cyrillaceaepollenites minor(Takahashi) Takahashi

Description: Tricolporate pollen grain;prolate in equatorial view. Three colpi
parallel, comparatively short, with large
and equatorially somewhat elongatedpores (equatorial diameter 4 pm, meridional diameter 3 pm). Exine 1 pm thick,finely intrabaculate. Grain size 28 X 20
pm. Breadth/length ratio 0.71.
Occurrence: Very rare.
Remarlzs: Only one specimen was
found.
Botanical affinity: ?Leguminosae.
Genus Cupuhferoipollenites Potonie
1951 ex Potonie 1960
Type species: Cupuliferoipollenit es
pusillus (Potonie 1934) Potonie 1960.

Cupuliferoipollenites sp.
PI. 51, Fig. 6.

Description: Tricolporate pollen grain;
subprolate in equatorial view. Three colpi

315-

PI. 51, Fig. 29.
1961. Tricolporopollenites minor Takahashi,Mem. Fac. Sci., Kyushu Univ., Ser. D,
Geol., 11, 3, p. 320-321, pI. 24, figs. 1831.
1979. Cyrillaceaepollenites minor (Takahashi)
Takahashi, Takahashi & Kim, Palaeontographica, B, 170, p. 46, pI. 13, figs.
32-36.

Dimensions: Grain size 13.5x 12.5 pm;
exine 0.5 pm thick, laevigate; rugae small ;breadth/length ratio 0.93.
Occurrence: Very rare.
Comparison: Only one specimen was
found. The present specimen is apparently identified with Cyrillaceaepollenites
minor (Takahashi) Takahashi from the
Paleogene and Miocene formations ofWest Japan and from the Miocene Yonil
Group of Korea.
Botanical affinity: Cyrillaceae.
Genus Euphorbiacites Sung & Lee 1976

Kiyoshi TAKAHASHI

316

Type species: Euphorbiacites wallensenensis (Pflug 1953) Sung & Lee 1976.

Euphorbiacites sp.
PI. 50, Fig. 31.

Description: Tricolporate pollen grain;prolate in equatorial view. Three colpi
more or less parallel, converging to thepoles, almost extending from pole to pole.
Caverna very deep. Pores large, elongating somewhat equatorially.
Exine isrelatively firm with sculpture which consists of closely spaced bacula of 2 pm
high in the polar area and 1.5 pm highin the equatorial area. Grain size 41 X
30 pm. Breadth/length ratio 0.73.
Occurrence: Very rare.
Comparison: Only one specimen was
found. The present specimen is similar
to Euphorbiacites sp. (Li, Sung & Li,
1978, p. 34, pI. 10, fig. 32) from the Eocene
and Oligocene formations of the YangtzeHan River plain, China, but the former
differs from the latter in having smallersize, thinner exine, and somewhat equatorially elongated pores (rugae).
Botanical affinity: Euphorbiaceae.
Genus Retitricolporites v. d. Hammen 1956
ex v. d. Hammen & Wijmstra 1964
Type species: Retitricolporites guianensis
v. d. Hammen & Wijmstra 1964.

Retitricolporites protensus n. sp.
PI. 51, Figs. 21-23.

Description: Tricolporate pollen grains;prolate in equatorial view. Three colpi
narrow, slender, converging to the poles,
extending from pole to pole with meridionally somewhat elongated pores. Exine
very finely reticulate. Lumen of reticulum
less than 1 pm in diameter. Muri baculate

to clavate, 0.5-0.7 pm high. Grain size 1617 pm X 9-12 pm.
Breadth/length ratio
0.56-0.71.
Holotype: PI. 51, Fig. 22; grain size 16
X 10.5 pm; muri baculate to clavate, 0.50.7 pm high. Breadth/length ratio 0.66;
slide GN 4653.
Occurrence: Few.
Comparisom: Retitricolporites protensus
is similar to the elongate form of Retitricolporites (al. Tricolporopollenites) microreticulatus (Pflug & Thomson) n. comb.,but differs from the latter in having
baculate and clavate muri.
Amongst the pollen of recent plants a
close similarity is seen with the pollen of
Hydrangea and it is quite likely that thepresent pollen grains are related to thefamily Saxifragaceae.
Botanical affinity: Saxifragaceae.
Genus Ilexpollenites Thiergart 1937
ex Potonie 1960
Type species: Ilexpollenites iliacus(Potonie 1931) Potonie 1960.

Ilexpollenites tertiarius (Takahashi)
Takahashi
PI. 51, Figs. 25-27.

1961. Tricolporopollenites tertiarius Takahashi, Mem. Fac. Sci., Kyushu Univ., Ser.
D, GeoI., 11, 3, p. 332, pI. 26, figs. 2933.
1963. Ilexpollenites tertiarius (Takahashi)
Takahashi, Mem. Fac. Sci., Kyushu
Univ., Ser. D, GeoI., 14, 2, p. 150, pI.
21, fig. 18.

Dimension: Grain size 18.5-27 pm X
15.5-19.5 pm; clavae 1.5-2 pm high, 1 pm
± in diameter; breadth/length ratio 0.570.92.
Occurrence: Few.
Comparison: The present specimens

74':1.

Miospores from the Eocene Nanggulan Formation

belong to Ilexpollenites tertiarius (Takahashi) Takahashi from the Paleogene andMiocene formations of West Japan andfrom the Miocene Changgi and Yonil
Groups of Korea. The Figures 26 and 27
with densely spaced clavae are closelycomparable with the grain of Ilexpollenites
tertiarius from the Hioki Group of Waku,
Yamaguchi Prefecture (Takahashi. 1963,
pI. 21, fig. 18).
Botanical affinity: Ilex.

Genus Striatricolporites v. d. Hammen
1956 ex Leidelmeyer 1966
Type species: Striatricolporites
pimulis Leidelmeyer 1966.
Striatricolporites striolatus n. sp.
PI. 51, Figs. 30-32.

Description: Tricolporate pollen grains;prolate in equatorial view. Three colpi
narrow, converging to the poles, extending from pole to pole, with meridionallymore or less elongated pores. Exine
finely striate; muri 0.9-1.5,um high. Striae
running parallel or more or less obliquely
to the col pus. Grain size 21-24 ,umx 1316,um. Breadth/length ratio 0.57-0.72.
Holotype: PI. 51, Fig. 32; grain size 23
X 13.2 ,um; exine finely striate; muri 0.9,um high; breadth/length ratio 0.57; slide
GN 4653.
Occurrence: Common.
Comparison: This new species is closely
similar to Striatricolporites augustinus
Gonzalez Guzman (1967, p. 39, pI. 12, figs.
I-H) from the Middle Eocene Mirador
Formation (pollen zones III-IV) of the
Tibu area, Colombia, but the former differs from the latter in possessing equatorially elongated pores (rugae), judging
from the Guzman's photographs.
The morphological characters of Stria-

317

tricolporites striolatus seem to be muchsimilar to that of the Gymnostemma-type,and it is quite likely that they might
represent that genus.
Botanical affinity: Cucurbitaceae-Gymnostemma.
Striatricolporites agustinus

Gonzalez Guzman
PI. 51, Figs. 33-34.
1967. Striatricolporites agustinus Gonzatez
Guzman, E. J. Brill, p. 39, pI. 12, figs.
1·1f.

Dimensions Grain size 21,um X 16-17.5,um; exine finely striate; muri l,um±
high; breadth/length ratio 0.67-0.83.
Occurrence: Few.
Remarl,s: Striatricolporites agustinus
is somewhat broader than Striatricolporites striolatus and possesses the equatorially elongated pores.
Botanical affinity: Cucurbitaceae-Gymnostemma or Actinostemma.
Striatricolporites sp.
PI. 51, Fig. 35.

Description: Tricolporate pollen grain;prolate in equatorial view. Three colpi
long, relatively narrow, longitudinal, converging to the poles, reaching to the poles,
with equatorial rugae. Exine finely striate;muri l,um high. Grain size 32.5 X 17 ,um.
Breadth/length ratio 0.52.
Occurrence: Very rare.
Botanical affinity: Unknown.

Subturma Ptychopolyporines Naumova
1937 emend. Potonie 1960
Genus Tetracolporopollenites
Pflug & Thomson 1953

Kiyoshi TAKAHASHI

318

Type species: Tetracolporopollenites
sapotoides Pflug & Thomson 1953.

Tetracolporopollenites obscurus
Pflug & Thomson
PI. 52, Figs. 1-3.
1953. Tetracolporopollenites obscurus Pflug &Thomson, Palaeontographica, B, 94, S.
108, Taf. 14, Fig. 86-99, 102-108.

Dimensions: Grain size 33-34,umx2527 ,um; exine 1-1.5,um thick, finely intrabaculate or faintly intrarugulate; pores
large, round or equatorially elongate.
Occurrence: Few.
Comparison: The grains seem to belong
to those of Tetracolporopollenites obscurus
Pflug' & Thomson from the Lower andMiddle Tertiary of West Germany,
Botanical affinity: Sapotaceae.
Genus Polygalacidites Sah & Dutta 1966
Type species: Polygalacidites clarus
Sah & Dutta 1966.

Polygalacidites speciosus n. sp.
PI. 52, figs. 4-14.

Description:

Stephanocolporate pollen

grains; suboblate to spheroidal with swelling in the equatorial area in equatorial
view; amb circular to subcircular in polar
view. Col pi numerous, generally six toeight in number, comparatively not solong, running more or less parallel meridionally, not reaching to the poles. Ora(pores) transversally parallel, connectedtransversally as ring or belt (synorate).
Exine 0.5-1.5,um thick, chagrenate. Grainsize 17-26 ,um X 20-27 ,um. Breadth/length
ratio 0.94-1.14.
Holotype: PI. 52, Figs. 5a-b; grain size
25x24,um; six colpi; synorate; exine
1,um thick, chagrenate; breadth/length
ratio 0.96; slide GN 4655.
Occurrence: Abundant.
Comparison: Hitherto, Polygalacidites
clarus Sah & Dutta (1966) has been described by Sah & Dutta (196b, 1968) fromthe Eocene (?Paleocene) Jaintia Series ofAssam. Polygalacidites speciosus differsfrom the Indian species Polygalacidites
clarus Sah & Dutta in number of col pus
and development of synora.
Botanical affinity: Polygalaceae-Polygala.
Turuma Poroses Naumova 1937
emend. Potonie 1960

Explanation of Plate 49(All figures x 1000)
Fig. 1. Leiotriletes maxoides Krutzsch subsp. minoris Krutzsch Slide GN 4655.
Figs. 2-3. Alsophilidites kerguelensis CooksonFig. 2: slide GN 4654; Fig. 3: slide GN 4655.
Fig. 4. ?Laevigatosporites sp. Slide GN 4653.
Figs. 5-6. .Laevigatosporites javanicus n. sp.
Fig. 5: holotype, slide GN 4654; Fig. 6: slide GN 4657.
Figs. 7-8. Smilacipites spinulifer n. sp. Slide GN 4654; Fig. 7: holotype.
Fig. 9. Clavapalmaedites sp. Slide GN 4654.
Figs. 10-12. Smilacipites echinatus Wodehouse
Fig. 10: slide GN 4655; Fig. 11: slide GN 4657; Fig. 12: slide GN 4653.
Figs. 13-15. Magnoliaepollenites ellipticus n. sp.
Figs. 13-14: slide GN 4657; Figs. 15a-b: holotype, slide GN 4656.

T AKAH ASH I:

Miosp07"eS from the Nanggulan F01"mation

Plate 49

743. Miospo1°es j1°om the Eocene Nanggulan F01°mation

319

Subturma Monoporines Naumova 1937
emend. Potonie 1960

Type species: Proteacidites adenanthoides Cookson 1950.

Genus Graminidites Cookson 1947

Proteacidites nwllis Samoilovitch

Type species: Graminidites media
Cookson 1947.

PI. 52, Figs. 20a·b; pI. 53, Figs. 1-2.

Graminidites punctulosus n. sp.
PI. 52, Figs. 15-19.

Monoporate pollen grains.
Figura circular to subcircular in outline.
Pore single, 3-4.7 pm in diameter, surrounded by large annulus of 3-4.5 pm inwidth. Exine finely punctate, 0.5-1 pm
thick, with secondary folds. Grain size
28-41 pm in diameter.
Holotype: PI. 52, Fig. 19; grain size 38
pm in diameter; pore single, 4.7 pm indiameter, surrounded by the annulus of
4 pm in width; exine 1 pm thick, very
finely punctate or intra punctate ; slide
GN 4656.
Occurrence: Common.
Comparison: This new species is closely comparable with Graminidites pseudogramineus Krutzsch (1970, S. 52, Taf. 1,
Fig. 6-11) from the Pliocene or Plio-Pleistocene sediments of Germany, but theformer differs from the latter in possessing much broader or stronger annulus
surrounding the pore.
Graminidites sp. from the Pohang Formation of the Yonil Group (Takahashi &Kim, 1979, p. 61, pI. 24, fig. 7) has only
a very weak annulus. surrounding the pore.
Botanical affinity: Gramineae.
Description:

1961. Proteacidites
mollisSamoilovitch,
Samoilovitch et aI., Trudy VNIGRI,
no. 117, p. 185, pI. 54, fig. 14; pI. 59,
figs. la·d; pI. 61, fig. 9.

Dimensions: Grain size 29-33 pm indiameter; triangular with convex sides inpolar view; exine punctate, 1.5-2pm thick.
Occurrence: Few.
Comparison: In general morphologicalcharacters the present specimens are considered to belong to Proteacidites mollis
Samoilovitch from the MaestrichtianDanian Upper Symsk Subseries, western
Siberian Lowland, USSR.
Botanical affinity: Proteaceae.

Proteacidites matsuokae n. sp.
PI. 53, Fig. 3-7.

Description: Triporate pollen grains;isopolar; amb triangular with somewhat
concave sides in polar view. Exine finely
punctate, 1.3-2.5 pm thick; ektexine: endexine= 1 : 2. Apertures 3-9 pm in diameter. Grain size 29-35 pm in equatorial
diameter.
Holotype: PI. 53, Figs. 5a-b; grain size
29 pm in equatorial diameter; exine finely
punctate, 1.5 pm thick; ektexine : endexine= 1 : 2; slide GN 4653.
Occurrence: Few.
Comparison:

Subturma Triporines Naumova 1939
emend. Potonie 1960
Genus Proteacidites Cookson 1950
ex Couper 1953

Proteacidites

matsuokae

can be distinguished from Proteacidites
scaboratus Couper (1960, p. 52, pI. 5, figs.
22-23) from the Upper Senonian or Maestrichtian to Danian strata of New Zealandin ornamentation and structure of exine.
The present species resembles Proteaci-Kiyoshi TAKAHASHI

320

dites incurvatus Cookson forma minor
Samoliovitch (Samoilovitch et aI., 1961, p.
188, pI. 50, figs. 3a-b; pI. 61, fig. 8) from
the Maestrichtian-Danian Upper Symsk
Subseries, western Siberian Lowland,
USSR, but the former differs from thelatter in having smaller grain size andaperture.
Botanical affinity: Proteaceae.
Proteacidites d. annularis Cookson

Range: Paleocene (Wangerip Group in
the Princetown area, South Australia);
Oligocene-Miocene (?) (brown coal, Moorlands, South Australia); Paleocene-Eocene(Lygistepollenites balmei-Nothofagidites
asperus Zone, Latrobe Group, Gippsland,
Australia); Paleocene (Waipawan)-Upper
Eocene (Runangan) (Dannevirke SeriesArnold Series, New Zealand); Middle
Oligocene (Waitakian, Landon Series, New
Zealand).
Botanical affinity: Proteaceae.

PI. 53, Fig. 8.
1950. Proteacidites annularis Cookson, Australian Jour. Sci., Res., Ser. B, BioI.
Sci., 3. 2, p. 170, pI. 1, fig. 15.

Dimensions: Grain size 27 pm in equatorial diameter; exine 2 pm thick (Ektexine: endexine=1 : 2), punctate; apertures 4-5 pm.
Occurrence: Very rare.
Comparison: EXcept for the comparatively thinner exine, the present specimen
appears to be almost identical with thegrain of Proteacidites annularis Cooksonfrom the Oligocene-Miocene(?) brown coal,South Australia.

Proteacidites similis Harris
PI. 53, Figs. 9a-b.
1965. Proteacidites simi lis Harris, Palaeontographica, B, 115, p. 94, pI. 29, figs. 1112.

Dimensions: Grain size 31.5 pm inequatorial diameter; ora circular 2 pm indiameter; exine 2.3 pm thick, scabrate
appearing minutely reticulate.
Occurrence: Very rare.
Comparison: The present specimen isvery closely similar to Proteacidites similis
Harris from the Middle Paleocene Pebble

Explanation of Plate 50(All figures x 1000)
Figs. 1-3. Classites capucinii Gonz31ez Guzman
Fig. 1: slide GN 4656; Fig. : slide GN 4653; Fig. 3: slide GN 4655.
Figs. 4-10. Dicolpopollis malesianus Muller
Figs. 4, 6, 10: slide GN 4654; Figs. 5a-b, 8, 9: slide GN 4653; Fig. 7: slide GN 4656.
Figs. 11-13, 30. Quercoidites umiensis (Takahashi) Takahashi
Fig. 11: slide GN 4654; Fig. 12: slide GN 4655; Figs. 13, 37: slide GN 4653.
Figs. 14-15. Tricolpopollenites elongatus n. sp. Slide GN 4657; Fig. 14: holotype.
Figs. 16-17. Quercoidites ellipsodeus n. sp.
Fig. 16: holotype, slide GN 4656; Fig. 17: slide GN 4654.
Figs. 18, 25-27. Tricolpites minutireticulosus Takahashi
Figs. 18, 27a-b: slide GN 4655; Figs. 25a-b: slide GN 4657; Figs. 26a-b: slide GN 4653.
Figs. 19-24. Cupuli/eroidaepollenites /allax (Potonie) Potonie
Figs. 19, 20: slide GN 4655; Fig. 21: slide GN 4654; Figs. 22-23: slide GN 4653.
Figs. 28-29. Brevitricolpites circularis n. sp. Slide GN 4653; Fig. 28: holotype.
Fig. 31. Euphorbiacites sp. Slide GN 4656.

TAKAH ASH I :Miospores from the Nanggulan Formation

~m"
'", .
~
.

:

to

.'
',~

~

' "

~

, .T/

'

.~ ..,.

.";J' t' -

28

.

•

29

30

Plate 50

743.

Miospores from the Eocene Nanggulan Formation

Point Formation, Dilwyn Bay, Victoria,
Australia.
Botanical affinity: Proteaceae.

Genus Subtriporopollenites
Pflug & Thomson 1953
Type species:

Subtriporopollenites

321

Subcircular or oval in equatorial outline.
Pores very small; two pores subequatorial
and one pore equatorial. Exine 1 pm thick,chagrenate, without atrium, labrum andannulus around the pore. Grain size 23.5
pm in equatorial diameter.
Occurrence: Very rare.
Botanical affinity: Unknown.

anulatus Pflug & Thomson 1953

Genus Triporopollenites Pflug &subsp. anlilatus

Thomson 1953 emend. Potonie 1960
Subtriporopollenites minutulus n. sp.

Type species:
PI. 53, Figs. 10-12, 13 (?), 18-19.

coryloides Pflug 1953.

Description: Triporate pollen grains.
Equatorial outline circular to subcircularor oval. Pores relatively large, circular,
2-2.5 pm in diameter; two or three pores
subequatorial. Exine 0.6-1.5 pm thick,finely punctate, with a weak annulus of
1 pm± in width around the pore. Grainsize 18-22 pm X 16-22 pm in equatorial
diameter.
Holotype: PI. 53, Fig. 10; grain size 20
X 19.5 pm in equatorial diameter; exine
1 pm thick, punctate; pores relatively
large, 2.5 pm in diameter; slide GN 4654.
Occurrence: Common.
Comparison: The present specimens
are comparable to Subtriporopollenites anulatus Pflug & Thomson subsp. nanus Pflug& Thomson (1953, S. 86, Taf. 9, Fig. 5455) and Subtriporopollenites firmus Pflug(1953, S. 86, Taf. 9, Fig. 62-63) from the
Tertiary deposits of Germany, but S.
anulatus nanus has intrabaculate to laevigate exine and S. jirmus has fossulate
sculpture of exine.
Botanical affinity: ?Juglandaceae.

Subtriporopollenites sp.
PI. 53, Fig. 15.

Description:

Triporate

pollenTriporopollenites

grain.

Triporopollenites sp. a
PI. 53, Fig. 16.

Description: Triporate pollen grain.
Triangular in equatorial outline. Threepores very small, planaperturate; twopores equatorial. Exine 1 pm thick, finely
punctate, with annulus around the pore.
Grain size 15 pm in equatorial diameter.
Occurrence: Very rare.
Remarks: This grain has three apertures (pores) and each is situated at the
midpoints of the sides of the triangularambo
Botanical affinity: Unknown.

Triporopollenites sp. b
PI. 53, Fig. 17.

Description: Triporate pollen grain.
Triangular with convex sides in equatorial
outline. Three pores round, equatorial.
Exine 2.5 pm thick, intrarugulate or finely
punctate, without labrum and annulus;
ektexine: endexine= 1 : 3. Grain size 32
pm in equatorial diameter.
Occurrence: Very rare.
Botanical affinity: Unknown.

Kiyoshi TAKAHASHI

322

Genus Subtriporopollis Sah 1967

Subtriporopollis
tenuis Sah 1967.

Type species:

Subtriporopollis specialis n. sp.
PI. 53, Figs. 20-21.

Description: Triporate pollen grains.
Amb rounded triangular or circular inpolar view. Pores circular, slightly protruded; two pores subequatorial. Exine
finely reticulate; lumina very small ;muri baculate, 0.5-0.7 fJ-m high; annulus
drop-shaped. Grain size 19-22 fJ-m X 17.518 fJ-m in equatorial diameter.
Holotype: PI. 53, Fig. 20; grain size 19
X 17.5 fJ-m in equatorial diameter; exine
reticulate; muri baculate, 0.7 fJ-m high;slide GN 4656.
Occurrence: Few.
Comparison: Sah (1967) established thegenus Subtriporopollis by the triporate
pollen grains with reticulum yielded fromthe Neogene strata, Rusizi valley, Burundi.
The present species differs from Subtriporopollis tenuis Sah (1967, p. 119-120, pI.
10, fig. 16) and Subtri poropollis rotzmdis
Sah (1967, p. 120-121, pI. 10, figs. 17-19)
in grain zize and pore size.
Botanical affinity: Rubiaceae.
Genus Tiliaepollenites Potonie 1931

Tiliaepollenites indubitabilis Potonie 1931.

Type species:

Tiliaepollenites tropicus n. sp.
PI. 53, Figs. 23-24.

Description: Triporate pollen grains.
Amb circular in polar view. Three germinals circular, relatively large, more orless protruded, with postvestibulum. Exine

Explanation of Plate 51(All figures x 1000)
Figs. 1-5. Tricolporopollenites javanensis n. sp.
Fig. 1: holotype, slide GN 4656; Fig. 2: slide GN 4653; Figs. 3-5: slide GN 4655.
Fig. 6. Cupuli/eroipollenites sp. Slide GN 4655.
Figs. 7-9. Tricolporopollenites consularis Takahashi subsp. consularis Slide GN 4654.
Figs. 10-14. Tricolporopollenites marginatus n. sp.
Figs. 10, 13: slide GN 4657; Figs. 11, 12: slide GN 4655; Fig. 11: holotype; Fig. 14: slide
GN 4654.
Figs. 15a-b. Tricolporopollenites sp. a Slide GN 4653.
Fig. 16. Tricolporopollenites sp. d Slide GN 4654.
Fig. 17. Tricolporopollenites sp. b Slide GN 4657.
Figs. 18-20. Tricolporopollenites rasus n. sp.
Figs. 18, 19: slide GN 4656; Fig. 19: holotype; Fig. 20: slide GN 4653.
Figs. 21-23. Retitricolporites protensus n. sp. Slide GN 4653; Fig. 22: holotype.
Fig. 24. Tricolporopollenites ovatorotundus Takahashi Slide GN 4656.
Figs. 25-27. llexpollenites tertiarius (Takahashi) Takahashi
Fig. 25: slide GN 4654; Fig. 26: slide GN 4657; Fig. 27: slide GN 4656.
Fig. 28. Tricolporopollenites sp. c Slide GN 4656.
Fig. 29. Cyrillaceaepollenites minor (Takahashi) Takahashi Slide GN 4653.
Figs. 30-32. Striatricolporites striolatus n. sp. Slide GN 4653; Fig. 32: holotype.
Figs. 33-34. Striatricolporites agustinus Gonzalez Guzman
Fig. 33: slide GN 4657; Figs. 34a-b: slide GN 4655.
Fig. 35. Striatricolporites sp. Slide GN 4654.

TAKAHASHI:

Miospores from the Nangg u lan Fonnation

Plate 51

10

11

' m'J

80

.
E)
~

17

1B

19

..

',

,

'~

,

'.

,

27

0.,
32

23a

743.

lVIiospores frorn the Eocene Nanggulan Formation

finely reticulate, 1 pm thick; lumina lessthan 1 pm. Grain size 28-31 pm X 27.5-30
pm in equatorial diameter.
Holotype: PI. 53, Figs. 24a-b ; grain size
28x27.5 pm in equatorial diameter; exine
finely reticulate, 1 pm thick, with postvestibulum around the pore; slide GN 4657.
Occurrence: Rare.
Comparison:
Tiliaepollenites tropicus
is comparable with Tiliaepollenites paucus
Sah (1967, p. 116-117, pI. 10, fig. 15) fromthe Neogene sediments of Rusizi valley,
Burundi, but the former differs from thelatter in having much smaller grain.
The general form, pore character andexine ornamentation of the grains strongly
suggest affinity with Tiliaceae.
Botanical affinity: Tiliaceae.

Tiliaepollenites ct. punctulosus Takahashi
PI. 53, Figs. 14a·b.

1979. Tiliaepollenites punctulosus Takahashi,
Takahashi & Kim, Palaeontographica,B, 170, p. 56, pI. 20, figs. 10-16.

Dimensions: Grain size 27 pm in eqatorial diameter; exine finely punctate,
1 pm thick, with postvestibulum aroundthe pore; germinals somewhat protruded,
equatorial.
Occurrence: Very rare.
Comparison: The present specimen isclosely similar to the grain of Tiliaepollenites punctulosus Takahashi from the~1iocene Changgi and Yonil Groups of
Korea, with exception of its smaller size.
Botanical affinity: Tiliaceae.
? Tiliaepollenites sp.
PI. 53, Fig. 22.

Description: Tetraporate pollen grain.
Equatorial contour circular. Pore circular,
relatively small, somewhat protruded;323

three pores subequatorial and one poreequatorial, with postvestibulum(?). Exine
laevigate, 0.5 pm thick. Breadth of porerim about 3 pm. Grain size 28 pm inequatorial diameter.
Occurrence: Very rare.
Remar!?s: Only one specimen was
found. This tetraporate grain belongs
doubtfully to the genus Tiliaepollenites.
Botanical affinity: ?Tiliaceae.
References
Belsky, C. Y., Boltenhagen, E. und Potonie,
R. (1965): Sporae dispersae der oberen
Kreide von Gabun, ,'\quatoriales Africa.
Palaont. Z., 39, 1/2, 72-83, Taf. 12-13.
Cookson, l. C. (1947): Plant microfossils fromthe lignites of Kerguelen Archipelago.
B. A. N. Z. Antarctic Res. Exped. 19291931 Repts. Ser. A., vol. 2, pt. 8, 127142, pIs. 13-17.
- - (1950): Fossil pollen grains of Proteaceous type from Tertiary deposits in Aus·
tralia. Australian Jour. Sci. Res., Ser.
B, Bio!. Sci., vol, 3, no. 2, 166-177, pIs.
1-3.
Couper, R. A. (1953): Upper Mesozoic andCainozoic spores and pollen grains fromNew Zealand. New Zealand Ceol. Surv.,
Paleont. Bull., 22, 5-77, pIs. 1-9.
- - (1960): New Zealand Mesozoic andCainozoic plant microfossils. New ZealandCeol. Surv., Paleont. Bull., 32, 1-87, pIs.
1-12.
Drugg, W. S. (1967): Palynology of the Upper
Moreno Formation (Late Cretaceous-Paleo·cene) Escarpado Canyon, California. Palaeontographica, B, 120, 1-71, pIs. 1-9.
Germeraad, J. H., Hopping, C. A. and Muller,
J. (1968): Palynology of Tertiary sediments from tropical areas. Rev. Palaeobot.
Paiyn., vol. 6, nos. 3-4, 189-348, pIs. 118, 19 figs.
Gonzalez Guzman, A. E. (1967): A palynological study on the Upper Los Cuervos andMirador Formations (Lower and Middle
Eocene; Tibu area, Colombia). 1-68, pIs.
1-30, E. J. Brill, Leiden.

324

Kiyoshi TAKAHASHI

Hammen, T. van der (1956): A palynologicalsystematic nomenclature. Bol. Ceol., vol.
4, nos. 2-3, 63-101, pis. 1-12.
- - and Wijmstra, T. A. (1964): A palynological study on the Tertiary and Upper
Cretaceous of British Guiana. Leidse Ceol.
Med., 30, 183-241, pis. 1-3.
Harris, W. K. (1965): Basal Tertiary microfloras from the Prince town area, Victoria,
Australia. Palaeontographica, B, 115, 75106, pis. 24-29.
Kadar, D. (1981): Planktonic foraminiferal
biostratigraphy of the Miocene-Pliocene
Sentolo Formation, Central Java, Indonesia.
35-47. in T. Saito (edit.): Micropaleontology, petrology and lithostratigraphy of
Cenozoic rocks of the Yogyakarta region,
Central Java. Yamagata Univ.
Krutzsch, W. (1962): Atlas der mittel- undjungtertiaren dispersen Sporen- und Pollensowie der Mikroplanktonformen des nordlichen Mitteleuropas. Lief. 1, 1-108, Taf.
1-46, Veb. deutsch. Verlag Wissensch.,
Berlin.
- - (1967): Atlas der mittel- und jungtertiaren dispersen Sporen- und Pollen-sowie
der Mikroplanktonformen des nordlichen
Mitteleuropas. Lief. 4 & 5, 1-232, Taf.
1-90, Veb. Gustav Fischer Verlag, Jena.
- - (1970): Atlas der mittel- und jungtertiaren dispersen Sporen- und Pollen-sowie
der Mikroplanktonformen des nordlichen
Mitteleuropas. Lief. 7, 1-175, Taf. 1-50,
Veb Gustav Fischer Verlag, Jena.
Leidelmeyer, P. (1966): The Paleocene andLower Eocene pollen flora of Guyana.
Leidse Ceol. Med., 38, 49-70, pis. 1-5.
Li, M. Y., Sung, T. C. and Li, Z. P. (1978):Some Cretaceous-Tertiary palynologicalassemblages from the Yangtze-han River
Plain. Mem. Nanjing Inst. Ceol. Palaeont.,
Academia Sinica, no. 9, 1-44, pis. 1-12.
Matsuoka, K. (1981):
Dinoflagellate cysts
and Pediatrulll from the Nanggulan and
Sentolo Formations in the middle. Java
Island, Indonesia. 48-51, pl. 1. in T. Saito(edit.) : Mrcropaleontology, petrology andlithostratigraphy of Cenozoic rocks of the
Yogyakarta region, central Java. Yamagata Univ.
Muller, J. (1968): Palynology of the Pedawan
and Plateau Sandstone Formations (Cretaceous-Eocene) in Sarawak, Malaysia. Micropaleontology, vol. 14, no. 1, 1-37, pis. 1-5.
Miirriger, F. und Pflug, H. (1951): Vber die
Altersstellung der Braunkohle von Burghasungen etc. Notizbl. Hess. L.-A. Bodenforsch., 4 Folge, Heft 2, 87-97.
Nagy, E. (1967): Palynological elaborations
the Miocene layers of the Mecsek Mountains. Ann. Hung. Ceol. Inst., \'ol. 52,
no. 2, 233-649, pis. 1-56.
Okada, H. (1981): Calcareous nannofossils of
Cenozoic formations in central Java. 2534, 6 tabs. in T. Saito (edit.): Micropaleontology, petrology and lithostratigraphy of Cenozoic rocks of the Yogyakarta region, central Java. Yamagata Univ.
Pflanzl, G. (1956): Das Alter der Braunkohlen
des Meissners, der Floze 2 und 3 des Hirschberges und eines benachbarten Kohlenlagers bei Laudenbach. Notizbl. Hess.
Landesamt Bodenforsch., 4, 232-244, Taf.
16-17.
Potonie, R. (1931):
Zur Mikroskopie derBraunkohle. Tertiare Bliitenstaubformen.

Explanation of Plate 52(All figures x 1000)
Figs. 1-3. Tetracolporopollenites obscurus Pflug & Thomson
Fig. 1: slide GN 4655; Fig. 2: slide GN 4657; Fig. 3: slide GN 4654.
Figs. 4-14. Polygalacidites speciosus n. sp.
Figs. 4-5: slide GN 4655; Figs. 5a-b: holotype; Figs. 6-9, 13-14: slide G='" 4654;
10-11: slide GN 4653; Fig. 12: slide GN 4656.
Figs. 15-19. Craminidites punctulosus n. sp.
Figs. 15-18; slide GN 4654; Fig. 19: holotype, slide GN 4656
Figs. 20a-b. Proteacidites mollis Samoilovitch Slide GN 4653.

Figs.

TAKAHASHI:

Miospores j?"om the Nanggulan F'o?"mcttion

Plate 52

4b

743.

Miospores from the Eocene Nanggulan Formation

Z. Bmunkohle, 16, 325-333, Taf, 1-2.
- - (1934): Zur Mikrobotanik des eozanen
Humodils des Geiseltals. Arb. Inst. Paiiiobot. Petrogr. Brennst. 4,25-125, TaL 1-6.
- - (1956):
Synopsis der Gattungen derSporae dispersae. I. Teil. Beih. Ceoi. lb.,
Heft 23, 1-103, Taf. 1-11.
- - (1960a): Sporologie der eozanen Kohle
von Kalewa in Burma. Senck. leth., 41,
1/6, 451-481, Taf. 1-2.
- - (1960b):
Synopsis der Gattungen derSporae dispersae. III. Teil. Beih. Ceol.
lb., Heft 39, 1-189, Taf. 1-9.
- - (1966):
Synopsis der Gattungen derSporae dispersae. IV. Teil. Beih. Ceol.
lb., Heft 72, 1-244, TaL 1-15.
- - , Thomson, P. W. und Thiergart, F.
(1950): Zur Nomenklatur und Klassifikation der neogenen Sporomorphae (Pollen
und Sporen). Ceoi. lb., 65, 35-70, Taf.
1-3, Abb. l.
Rao, K. P. and Ramanujam, C. G. K. (1978):
Palynology of the Neogene Quilon beds ofKerala State in south India I- spores of
Pteridophytes and pollen of monocotyledons. The Paiaeobotanist, vol. 25, 397427, pis. 1-5.
Sah, S. C. D. (1967): Palynology of an Upper
Neogene profile from Rusizi valley (Burundi). Musee Roy. ['Afr. Cent-Terv.,
Belg. Ann., ser. in 8°, Sci. Ceol., 57, 1173, pis. 1-13.
- - and Dutta, S. K. (1966): Palyno-stratigraphy of the sedimentary Formations ofAssam: l. Stratigraphical position of theCherra Formation. The Palaeobotanist,
vol. 15, nos. 1-2, 72-86, pis. 1-2.
- - and - - (1968): Palynostratigraphy of
the Tertiary sedimentary formations ofAssam: 2. Stratigraphic significance ofspores and pollen in the Tertiary succes-

sion of Assam. The Balaeobotanist, vol.
16, no. 2, 177-195, pis. 1-2.
Samoilovitch, S. R. et al. (1961): Pollen andspores of western Siberia, Jurassic-Paleocene. Trudy VNICRI, no. 177, 1-657, pis.
1-84 and I-LXV, text-fig. 1, tabs. 1-3.
Sung, T. C. and Lee, M. (1976): Mesozoic
and Early Paleogene sporo-pollen assemblages from Yunnan, China. Part II. Early
Upper Cretaceous assemblages from Lufengand Monding and late Upper CretaceousEarly Paleogene assemblage from Mengla,
Yunnan. 9-64, pis. 2-10. Sci. Publ. Co.
Takahashi, K. (1957):
Palynologisch-stratigraphische Untersuchung der tertiaren
Schichten im Kasuya- und Fukuoka-Kohlenfeld von Nordkyushu, Japan. Mem. Fac.
Sci., Kyushu Univ., Ser. D, Ceol., vol. 5,
no. 4, 199-221, TaL 38-39.
- - (1961): Pollen und Sporen des westjapanischen Alttertiars und Miozans. (IITeil). Mem. Fac. Sci., Kyushu Univ.,
Ser. D, Ceol., vol. 11, no. 3, 277-345, TaL
13-27.
- - (1963): Sporenpalaontologische Untersuchungen der Hioki-Schichtengruppe von\Vaku und Kiwado. Mem. Fac. Sci., Kyushu Univ., Ser. D, Ceol., vol. 14, no. 2,
143-157, textfigs. 1-5, tab. 1-2, TaL 20-22.
- - and Kim, B. K. (1979): Palynology of
the Miocene formations in the Yeo un gillbay district, Korea. Palaeontographica,B, 170,10-80, pis, 1-28.
Thomson, P. W. und Pflug, H. D. (1953): Pollen und Sporen des mitteleuropaischen
Tertiars. Paiaeontographica, B, 94, 1-138,
TaL 1-15.
Wodehouse, R. P. (1933): Tertiary pollen. II.
The oil shales of the Eocene Green River
Formation. Bull. Torr. Bot. Club, 60,
477-524, text-figs. 1-56.

9:1$:;'1'7, :;" :7:;-\7;/Jlvj1Jttr;l]O)~idfrIitT:/:7':7 :/ii'i'ij£Jli!l'fo:{Et}: *~ft, 9:ttff):;:;,,:7:; -v;/J IV j1 :Ii!!:1JO)!m~ill:T :/ Jl:7 :/I\TIO)!l£j;RiJ> I?, lli'l'f 4f!lU: :{U5}44 fi, 1fH
48fill~m~'iJIJ L
fiG;l&; L k: o 48fi9:tl7filliJ~~fill"(';5 0: Laevigatosporites javanicus n. sp.,
Smiiacipites spinulifer n. sp., Magnoiiaepollenites ellipticus n. sp., Quercoidites
ellipsodeus n. sp., Tricolpopollenites elongatus n. sp., Brevitricolpites circularis n.
sp., Tricolporopollenites javanensis n. sp., T. marginatus n. sp., T. rasus n. sp.,
Retitricolporites protensus n. sp., Striatricoiporites triolatus n. sp., Poiygaiacidites
speciosus n. sp., Craminidites punctulosus n. sp., Proteacidites matsuokae n. sp.,
-V 7,

325

326

Kiyoshi TAKAHASHI
Subtriporopollenites minutulus n. sp., Subtriporopollis specialis n. sp., Tiliaepollenites tropicus n. sp.
j- :/ Y '7 :/{t~)Hl'f~~t Dicolpopollis malesianus, Polygalacidites speciosus, Striatricolporites spp., Proteacidites spp., Graminidites punctulosus ~OX:t~{t:t'JI:' J: '0-C~.#~Gh-CU~, @~*.~~~~M~~{tmW~~~tt.~~~~~h~o~• •Explanation of Plate 53(All figures x 1000)
Figs. 1-2. Proteacidites mollis Samoilovitch
Fig. 1: slide GN 4655; Fig. 2: slide GN 4654.
Figs. 3-7. Proteacidites matsuokae n. sp.
Fig. 3: slide GN 4654; Fig. 4: slide GN 4657; Figs. 5a-b: holotype. slide GN 4653; Figs.
6-7: slide GN 4655.
Fig. 8. Proteacidites cf. annularis Cookson Slide GN 4655.
Figs. 9a-b. Proteacidites simi lis Harris Slide GN 4655.
Figs. 10-l3 (?), 18-19. Subtriporopollenites minutulus n. sp.
Figs. 10, 13: slide GN 4654; Fig. 10: holotype; Figs. 11, 18: slide GN 4653; Fig. 12:
slide GN 4657; Fig. 19: slide GN 4656.
Figs. 14a-b. Tiliaepollenites d. punctulosus Takahashi Slide GN 4654.
Fig. 15. Sub triporopollenites sp. Slide GN 4653.
Fig. 16. Triporopollenites sp. a Slide GN 4653.
Fig. 17. Triporopollenites sp. b Slide GN 4656.
Figs. 20-21. Subtriporopollis specialis n. sp.
Fig. 20: holotype, slide GN 4656; Fig. 21: slide GN 4654.
Fig. 22. ?Tiliaepollenites sp. Slide GN 4657.
Figs. 23-24. Tiliaepollenites tropicus n. sp.
Fig. 23: slide GN 4654; Figs. 24a-b: holotype, slide GN 4657

TAKAHASHI:

Miospores from the Nanggulan F01"mation

5aPlate 53

9a

10

"

~

• ','

j

O
"
I

~ ,p

,

7
6

•

11

...

,

fi
,~' ....

• " . :"

• I

'.

12

9b

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 126, pp. 327-333, June 30, 1982

744.

URANIUM-SERIES AGE OF THE "KAMETSU FORMATION",RIUKIU LIMESTONE ON THE TOKUNO-SHIMA,RYUKYU ISLANDS*AKIO OMURA
Department of Earth Sciences. Faculty of Science, Kanazawa University

Abstract. Uranium-series dating of some hermatypic corals (Acropora sp.)
reveals that the uppermost part of the Pleistocene series occurred on Tokuno-shima(Central Ryukyus) is likely to be Middle Pleistocene in age, at least older than 300,000
years. Although the coral ages obtained are almost near or beyond the limitation of
the 230Thj234U method of dating, the upper limit of the age is estimated from themean 23'Uj238U activity ratio to be approximately 700,000 years. This result is suggestive that reefy sediments formed during the last interglacial stage, 120,000 eyars
ago when sea level stood about 6 m higher than the present, is exposed nowhere on
the island. In addition, the absence of the Holocene Raised Coral Reef Limestone(Hanzawa, 1935) on the island may also imply that the vicinity of Tokuno-shima has
been locally subsided since the last 120,000 years.

The present paper deals with radiometricages estimated by the 23 0 Thj234U and
23 4 U/238U methods for hermatypic corals
from the uppermost part of the Pleistocene
series on Tokuno-shima, Central Ryukyu
Islands. These are the first uranium-series
dates from this island, which allow us tocorrelate the dated stratigraphic unit withthose on the other islands in Ryukyus
and to relate it to various Pleistocene
events in other areas.
Most of all islands in Central andSouthern Ryukyus are capped or rimmedwith Quaternary reefy limestone which
was divided into two stratigraphic units,
Pleistocene Riukiu Limestone and Holocene
Raised Coral Reef Limestone by Hanazawa{1935). Subsequently, the Riukiu Limestone
has been subdivided into a few unit onsome islands. For an example, Nakagawa(1967) redefined the Pleistocene series on
Tokuno-shima as the Ryukyu Group andstratigraphically subdivided into three,
Kametsu, Kinoko and Itokina Formations,in descending order, each of which wasmorphostratigraphically correlated among
subdivisions on the neighboring islands,
Okierabu-jima, Yoron-jima and Kikai-jima(Nakagawa, 1969). It presents, however,practical difficulty to determine the age ofeach unit by means of a radiometricdating such as 23°Th/ 234 U method, because
organic remains of calcium carbonate are
diagenetically altered in most cases
to stable phase (low Mg calcite) in
phreatic and/or vadose environments.
Moreover, morphostratigraphic techniquealso can be hardly employed to correlate
among subdivisions on respective islands,
because of the local variation in the rateof vertical displacement (Konishi et al.,* Received July 21, 1981; read Jan 24, 1981 at
Sendai.

327

328

Akio OMURA

1974). Those reasons have made it hardto assign chronologically the formationof Pleistocene series in Ryukyus, eventhough it was geologically investigatedin detail and stratigraphically subdivided
into some units.
The coral samples analyzed here are
only a genus Acropora collected at Isenzaki (27°39.4' N Lat., 128°56.3' E Long.),
the southernmost end of Tokuno-shima(Fig. 1), where a narrow (ca. 100 m inwidth) but cospicuous marine terrace isdeveloped over a distance of several
kilometers along the coast. The top of
the terrace is approximately 9 m inaltitude. As illustrated in Fig. 2, thelimestone unit which overlies a basal
conglomerate is about 7 m in the maximum
thickness. The uppermost part of thelimestone unit is abundant in coral heads(mostly of Acropora sp.), over 100 cm indiameter in growth position. During fivedays for an on-the-spot investigation,
fourty-eight coral heads were carefully
examined for their mineralogical natureby the staining method using Feigl's
solution in an area of about 20,000 m2 on
the surface of the terrace, and fifteen
samples were chosen for their appearanceof aragonitic nature and took back to thelaboratory.
The terrace sediments shown in Fig. 2dr--+_--,Amami-Ot>Tokuno-shi

128·

0

130·

c1'Oldnoerabu-jim"Yoron- j ima_........
Isen-zoki............5km

Fig. 1. Index map showing the locality where the fossil coral samples were collected.
(Dotted part is the distributing area of Pleistocene series on Tokuno-shima;
after Nakagawa, 1967).

744.

Radiometric age of Kametsn Formation

329

m

10

5

Fig. 2.

Simplified sketch of the terrace sediments at Isen·zaki, thesouthernmost end of Tokuno·shima.

come under a part of the Kametsu Formation of Nakagawa (1967) and are
regarded as the uppermost part of the
Pleistocene series on Tokuno-shima. The
HC dates of 5,800±230 (Mach ida et al"1976) through 43,700:':~Jgg years B. P. (seenin the table compiled by Pirazzoli, 1978)
have been reported for some corals whichwere collected at the same locality as the
sampling site in this study. The ResearchGroup for Active Faults (1980) seems toconsider that the limestone units formed
during two stages (120,000 and less than
100,000 years ago, respectively) areoccurred along Isen-zaki.
Results of analyses are summarized inTable 1.
X-ray powder diffraction patterns
revealed that low Mg calcite appeared
owing to the recrystallization of skeletal
aragonite was detectable in most samples.
It was only one sample numbered as 802-9-5 that was free of the secondary
calcite. In the present study, the analyses
were repeated for this and the other two
samples (Nos. 75-11-10-1 and 75-11-10-2)
in which only a few percent or less inweight of calcite was detected. Twosorts of 232U _228Th spikes, namely "Harwell"
spike allotted by the Uranium-Series
Intercomparison Project (Harmon et al.,1979) and "KU (Kanazawa Univ.)" spike
prepared by the author, were used asyield tracers in the repeated analyses,
separately,Consequently,
the high
reproducibility of the results attests to thereliability of each value listed in Table 1.
23 0Th/ 234 U ages were calculated by theequation;+("3'U _23 8 U)[Ao/(2o- }.,)]·[l-exp (2,t-A ot)]
where Ao and A., are decay constants of
23 0 Th and 23·'U, respectively. The quoted
errors are based on the counting statistics.
The samples containing five percent ormore of calcite cannot be anticipatedshowing the reliable uranium-series dates,
because a closed system respect to uranium
and thorium isotopes has not beenretained throughout their diagenetic
history. The upper six mTh/ 234 U ages
for three samples in Table 1, however,are reliable enough to use in the discussion below for the following evidences;(1) The specimens are entirely or almost
free of recrystallization, as shown by theabsence or an extremely small quantityof the secondary calcite. (2) The assumption of negligible initial 23 0Th is
supported by the observation that two
samples seem to be free of 23 2Th andthat 230Th/"32Th ratios in the other one

330

Akio OMURA
Table 1.

Isotopic composition and estimated ages of fossil corals
from the Kametsu Formation.
Activity Ratio

Isotope Concentraion

Estimat~d

CalciteSample No.

238U(wt %)

80-2-9-5

75-11-10-1

75-11-10-2

234 U

230Th

232Th
(ppm)

234 U/238 U

230 Th /232 Th 230 Th /234 U(dpm/g)230 Th Aga

(ka)

(ppm)

(dpm/g)3.07:':0.05

2.30:':0.04< 0.02

2.32:':0.04

1.01±0.02

1.01±0.02> 468

3.02:':0.05

2.33:':0.04< 0.02

2.29:':0.04

1.03=0.01

0.983=0.021

423_ 78

3.52:':0.08

2.70:':0.06< 0.02

2.70:':0.06

1.03=0.02

1.00=0.03

486_ 134

1.00:':0.03+00
529 -169

0< 1
3.41:':0.09

2.60:':0.07

3.34=0.04< 0.02+~+~

2.60:':0.06

1.02:':0.02

2.58:':0.03 0.0464:':0.0085

2.63:':0.06

1.04:':0.01

236 ± 43

1.02=0.03> 407

3.42±0.05

2.55±0.04 0.0257:':0.0034

2.46:':0 .03

1.02:':0.01

392 ± 52

0.943=0.018

301+ 35-27

2.50:':0.04

1.00:':0.01

1.04=0.0200

2.07:':0.04

1.01:':0.01

0.845:':0.01

201+13-12

380-2-8-3

5

3.22:':0.04

2.41:':0.03

80-2-9-2

5

3.26:':0.04

2.45:':0.03 0.0441:':0.0077

80-2-9-4

92.88=0.04

2.14:':0.03< 0.02

1.69=0.06

1.00=0.01

0.790:':0.02

169+16-13

75-11-10-4

21

2.83:':0.07

2.11:':0.05< 0.02

2.00:':0.08

1.00=0.02

0.948=0.045

320+ 220-70

75-11-10-543

2.59=0.06

1.93:':0.04

0.114:':0.016

2.25=0.06

1.00=0.02

1.17=0.04~< 0.02196 ± 34

82.3:':11. 8(For the upper three samples, the results in the upper and lower rows were obtained byuse of "Harwell" and "KU" spikes as a yield tracer, respectively.)(75-11-10-2) are very much higher thanthose in natural waters, which are commonly 1 to 3.
The 23 0Thj234U ratios for the upper
three samples in Table 1 range fromO.943±0.0l8 to 1.02±0.03. These values
are suggestive that 23 0Th has been nearlyattained equilibrium with its parent 234U.
For this reason, the limestone unit fromwhich the samples examined were derived
appears to be too old for the 230Thj234U
method of dating to be applicable, although the ages can be numericallycalculated by using the above equation,
as shown in the table. In other words,
the Kametsu Formation of Nakagawa(1967) is regarded as being more than
300,000 years old.
Nakagawa (1969) attempted once tocorrelate morphostratigraphicallytheKametsu Formation with his Wan For-mation on Kikai-jima. On the other hand,
the Pleistocene series on Kikai-jima was
subdivided into four units, Araki Limestone,
and Younger, Middle and Older Limestone
Members of Riukiu Limestone, in descending order by Konishi et at. (1974), whichhave been dated by the 230Thj234U and231Pa/ 235 U methods to be 35,000-45,000,
55,000-65,000, 80,000-100,000 and 120,000130,000 years, respectively.
The WanFormation of Nakagawa (1969) seems to
be matchable a suit from the Araki Limestone through the Middle Limestone
Member of Riukiu Limestone of Konishi
et at. (1974). In this viewpoint, it may
be said for the above-mentioned date of
the Kametsu Formation to be too oldcontrary to the expectations. However,
the reliability of an age of more than
300,000 years is supported by the averagevalue of 234U/23SU ratios in Table 1.

744.

331

Radiometric age of Kametsu Formation

The 23·'U/ 238 U actIvIty ratio can beused to estimate the age of biogenic
carbonates by the following equation,
provided that the initial activity ratio (Ro)is known and the sample has remained
closed to uranium and the daughters
between 238U and 23 4U in the uranium
decay series:
23 4U /238U = 1+(Ro-1)e-" 234 t

Because three values for Ro have beenreported up to present, the equation isplotted in Fig. 3 to show the decrease of
23·'U /238U ratio as a function of time forthe cases, each of which Ro is 1.15
(Thurber, 1962), 1.14 (Ku et at., 1977) and1.12 (Nikolayev et at., 1979), respectively.
The 234U /238U ratios for the threesamples are characterized by limited
ranges of 1.01±0.02 to 1.04±0.01, andthe mean is calculated to be 1.03±0.01.
From this mean 234U / 238 U ratio, the ageof the Kametsu Formation is estimated to
be 387,000 through 709,000 years. Finally,
it may be given as a conclusion here thatthe uppermost part of the Pleistocene
series on Tokuno-shima is 300,000 yearsof age or more, the upper limit of whichis estimated as being approximately
700,000 years.
It follows justly from a conclusion
stated above that a question arises as to
the reefy sediments which must havebeen formed around Tokuno-shima duringhigh stand of the sea 120,000 years ago.
Shoreline features standing about 6 m
above the present sea level have beenfound in many places in the world, andthey are thought to represent a eustatic
high sea stand that occurred 120,000[.[5-----IRo = 1.151
0[·[0

~

-------IRo=I.14/

>.

+.s;

~
=>

~

-----/Ro=1.I21
1.05

C\J

:s

<;)'

1'0
C\J

50

[00

Time in Units of 104 Years
Fig. 3. Decay curves of the excess 234U toward the secular equili·
brium with 2dBU for the initial 234U/23BU activity ratios of 1.15 (Thurber,
1962), 1.14 (Ku et ai., 1977) and 1.12 (Nikolayev et ai., 1979).

332

Akio OMURA

years B. P. (Bloom et at., 1974; and others).
The then reefy limestone ha ve beenreported on Kikai-jima, which is locatedapproximately 120 km NE of Tokuno-shima
and is the outermost island in the central
segment of the Ryukyu island arc, byKonishi et at. (1970, 1974). They argued
from the present elevation of such alimestone unit and the other units formed
during the subsequent interstadial phasesthat Kikai-jima has been rising at a rateof 1-2 m per 1,000 years since the last
120,000 years due to the neotectonic
interplay as plate convergence of theactive island arc and trench system. If
Tokuno-shima has not been moved
vertically during the past 120,000 years,
the island should be surrounded by thecoraliferous limestone unit which reaches
the maximum elevation of about 6 m andis dated as being 120,000 years. As
concluded above, even the uppermost
part of the Pleistocene series is thought
to be Middle Pleistocene in age and no
Late Pleistocene reefy limestone is foundon Tokuno-shima.
Consequently, no
occurrence of the reefy limestone formedin the last interglacial stage may besuggestive that Tokuno-shima locating
more backward than Kikai-jima in the
Ryukyu island arc, appears to have been
subsided during the past 120,000 years.
In addition, the absence of the Holocene
Raised Coral Reef Limestone (Hanzawa,1935) on Tokuno-shima may support thatthe vicinity of Tokuno-shima has been
locally subsided since the last 120,000
years.
The author is grateful to Prof. K.
Konishi, Department of Earth Sciences,
Faculty of Science, Kanazawa University,
for his continuous encouragement throughout this study.
This research was
supported in part by the Grant-in-Aid for
Scientic Research (Nos. 464265 and 56540481)
from the Ministry of Education, Science

and Culture.
References
Bloom, A. L., Broecker, W. S., Chappell,]. M. A., Matthews, R. K. and Mesolella,
K. J. (1974): Quaternary sea level fluctuations on a tectonic coast: New 230Th/
234U dates from the Huon Peninsula, New
Guinea. Quaternary Res., vol. 4, p. 185205.
Hanazawa, S. (1935): Topography and geology of the Riukiu Islands. Tohoku Univ.,
Sci. Rep., 2nd Ser. (Geol.), vol. 17, p. 161.
Harmon, R. S., Ku, T. L., Matthews, R. K. andSmart, P. L. (1979): Limits of U-series
analysis: Phase 1 results of the UraniumSeries Intercomparison Project. Ceology,
vol. 7, p. 405-409.
Konishi, K., Schlanger, S. O. and Omura, A.
(1970): Neotectonic rates in the Central
Ryukyu Islands derived from 230Th coralages. Marine Ceol., vol. 9, p. 225-240.
- - , Omura, A. and Nakamichi, O. (1974):
Radiometric coral ages and sea level records from the Late Quaternary reef complexes of the Ryukyu Islands. Proc. 2ndInt. Coral Reef Symp., vol., 2, p.595-613.
Ku, T. L., Knause, K. G. and Mathieu, G. G.
(1977): Uranium in open ocean: concentration and isotopic composition. DeepSea Res., vol. 24, p. 1005-1017.
Machida, H., Nakagawa, H. and Pirazzoli, P.
(1976): Preliminary study on the Holocene
sea levels in the Central Ryukyu Islands.
Rev. Ceomorphol. Dyn., vol. 2, p. 49-62.
Nakagawa, H. (1967): Geology of Tokunoshima, Okierabujima, Yoronto, and Kikaijima, Amami Gunto: Part 1.
TohokuUniv., Inst. Ceol. Pal., Contr., no. 63, p.
1-39. (in Japanese with English abstract)- - (1969): Geology of Tokunoshima, Ok ierabujima, Yoronto, and Kikaijima, Amami
Gunto: Part 2. Tohoku Univ., Inst. Ceol.
Pal., Contr., no. 68, p. 1-17. (in Japanesewith English abstract)
Nikolayev, D. S., Okunev, N. S., Drozhzhin,
V. M., Yefimova, Yeo I. and Belyayev,
B. N. (1979): 234U /238U isotopic rations in744.

Radiometric age of Kametsu Formationwaters of oceans, some seas and rivers.
Geokhimiya, no. 4, p. 586-597.
Pirazzoli, P. A. (1978): High stands of Holocene sea levels in the Northwest Pacific.
Quaternary Res., vol. 10, p. 1-29.
Research Group for Active Faults (1980):

333

Active Faults in Japan: Maps and Invetories. 363 pp., Univ. Tokyo Press, Tokyo.
Thurber, D. L. (1962): Anomalous 23·'Uj238U
in nature. Jour. Geophys. Res., vol. 67,
p. 4518-4520.

.~~u.~~nM.~~~ ~*M.~:m~ • • •~a~~~~~~~~ft~St~.h-n'~ff.l~~ (q:.)II,.t1967) ;I;'Gj3iHI~1...-t-=:"'T:/::t (Acropora sp.)

1~~1:::''01,'-C, 230Th
230Thj234U 1iZM.~ll!U~$~j@ifllp.&3'f;l;', ~~Ht.:t.h~~*-~;I;~~ G.h t-=:o ;l: t-=: 234U /238U 1iZM~gJtI:::.~--::i < 234U .~It 387,000~709,000.~~;J<otko .:t~t!i*,{if:[ (307].tJ~)
m~~.ht-=:o ~tJ:t.:)l?, ffli'!t~~.t~{[Ht307].tJ~, .:t 1...- -C.:t~~p.&lt~707].t t!iiltrn~.h~ 0 t!iJjjj, .~a~lt :r!!3'fatJ: t'-C·J! G.h~ :ifH,q:rs97Jc1tJl:to.l: If.:t~l&~~@I~lIJi:rsl*W1

1:::.%nJi: ~.ht-=: l\\(\!1.:!ifJ(4'k! (Konishi et ai., 1974) -'?7'C~ill:~ ull!H~"'T:/ ::z'l\(¥.~~:E-n (Han.
zawa, 1935, ~ Raised Coral Reef Limestone) ;I;~~jt:(EIlt~I:::.n!±l 1...- -n ,tJ:~,:: tit,!)dJ: < t {,~~127].Fs9It, .~aftJ5:;I;~tt~jlj!(cH:::'~~':: t ~~~1...- -C\"~o:;kttllJHlt<, ,::

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 126, pp. 334-340, pI. 54, June 30, 1982

745. A NEW NAUTILOID SPECIES FROM THE UPPERCRETACEOUS IZUMI GROUP OF SHIKOKU*
MITSUNOBU FURUICHI
Kagawa Natural Science Museum, Sakaide,
Kagawa Prefecture, 762

Abstract. A specimen from the Upper Cretaceous Izumi Group of the Sanuki
Mountains, Shikoku, is described as representing a new species of Eutrephoceras.
This is clearly different from E. dekay (Morton), the type·species from the North
American Atlantic Coast. But in the sutural pattern and shell form it possibly
resembles E. neubergicum (Redtenbacher) from the Upper Cretaceous of Europe andis somewhat similar to E. cookanum (Whitfield) from the Lower Tertiary of North
America. It is suggested that some characters of Nautilus, e. g. the less inflation offlanks and the slightly sinuous suture, are foreshadowed in this species.

The nautiloid fossil was collected froma nodule of the hard calcareous mudstone
in the massive black shale, about 150
meters above the base of the Izumi Group.
The location is shown in Text-fig. 1, andthe horizon is indicated as B in Textfig. 2. From the same horizon-B I havecollected Baculites subanceps pacificus
Matsumoto et Obata, Gaudryceras sp.,
Didymoceras sp., Acteocinidae ? gen. et sp.
indet., Aporrhaidae? gen. et sp. indet.,
Gyrodes ? sp., Naticidae ? gen. et sp. indet.,
Acila (Truncacila) hokkaidoensis (Nagao),
Anomia sp., Glycymeris sp., Inoceramus(Endocostea) balticus toyajoanus Nagao etMatsumoto, I. sp., Nanonavis elongatus(Nagao et Otatume), Parvamussium d.
awajiense (Ichikawa et Maeda), P. yubarense(Yabe et Nagao), Portlandia cuneistriata
Ichikawa et Maeda, Linthia sp., shark
teeth, fish scale and Populus sp., and fromthe horizon-A, about 150 meters lower
than B (see Text-fig. 2), Loxo japonica(Amano). Among these fossils, ammonoids
have been identified by Dr. T. Matsumoto,

Introduction

This paper is to report a new speciesof Eutrephoceras which was obtained
from the Upper Cretaceous Izumi Groupof the Sanuki Mountains of Shikoku andto consider its implications.
Geological Setting

The locality where the natutiloid specimen was found is the Tawa pit (long.
134 12'07"E, Lat. 34 10'44"N; Text-fig. 1)
at Tawa, Nagao-cho, Ookawa-gun, Kagawa
Prefecture. In this pit, the strata of
about 90 meters in thickness crop out.
They belong to the Nakato Shale, which
is a member in the lower part of the
Izumi Group, and overlies conformably
the basal member called the Korobiishi
Sandstone and Conglomerate (Nakano,
1953; Nakagawa, 1961).
00* Received July 22, 1981; read July 29, 1980,
at KochL

334

745.

Nautiloid from the Izumi Group

335

N

t

..
a

.<:

'"

--

~

0

'- a
0'"
Co a
Z

LEGENDg- --6

D.... 58.'. 40.. 3
l:b~1

1"--+-+1

2

X

10

/

9

.- 8

§

7
0

1

Text-fig. 1. Geological route map atand around the Tawa-pit. 1: granite, 2 :conglomerate, 3: coarse-grained sandstone,
4: medium-grained sandstone, 5: fine·grainedsandstone, 6: mudstone, 7: tuff, 8: nodule,
9: dip & strike, 10: locality of fossils, A &B: fossiliferous horizon; 1: basement rocks,
2-8: Izumi Groupgastropods by Mr. T. Kase, bivalves by
Drs. M. Tashiro and M. Noda, plant by
Dr. K. Matsuo, all of whom lowe much.
Now the horizon-B is characterized by
the common occurrence of Baculites.
This Baculites bed is traced to the west,
and it seems to be stratigraphicallyhigher than the Metapiacenticeras subtiiistriatum (Jimbo) bed (Matsumoto et ai.,1980) which is exposed in the west of thearea of Text-fig. 1. Consequently the
bed which yielded the present specimen
is regarded as Upper Campanian, because

Text·fig. 2. Geologic columnar section
at and around the Tawa·pit (Legend: sameas in Text-fig. 1)

M. subtilistriatum and B. subanceps pacificus
are the indices of the Upper Campanian(Matsumoto, 1977).
Palaeontological Description

Family Nautilidae de Blainville, 1825
Genus Eutrephoceras Hyatt, 1894
Type-species: Nautilus dekay Morton,
1834
Remarks: Cimomia and Eutrephoceras
are rather similar with respect to shellform and sutures, but in the typicalrepresentatives of them there is somedifference in the sutural pattern. Accord-

Mitsunobu FURUICHI

336

ing to Kummel (1956), in Eutrephoceras the
suture is essentially straight or only
slightly sinuous, but in some species ofCimomia it is so sinuous that it could
lead to the Hercoglossa type suture. In
some other species of Cimomia, it is transitional to the Eutrephoceras type.
Eutrephoceras is most intimately related
to Nautilus (s. s.). I expect that theancestor of living Nautilus could be foundin some Late Cretaceous species of
Eutrephoceras by way of the Tertiary
ones.
Occurrence: Upper Jurassic to Neogene.
World wide.

B

Eutrephoceras tawaense sp. nov.
PI. 54, Figs. 1-4; Text·figs. 1-4

Holotype: K-4-50 (in my collection
number), to be kept in the Kagawa
Natural Science Museum of Kagawa
Prefecture. It consists of a completely
preserved phragmocone (internal mould)
and the body chamber of one-third
volution. The shell is partly crushed, as
illustrated in Text-fig. 3 and PI. 54, Fig.
1. The surface of the whorl has partly
test and some epifauna. On the bodychamber weak breakage and cracks arerecognized and the conch is slightly
deformed, as illustrated in Text-figs. 3,
4-A and B and PI. 54, Fig. 2.
Diagnosis:
Shell nautiliconic, subglobular and involute. Umbilicus rather
small with rounded umbilical shoulder,

Text-fig. 3. Lateral view of the holotypeof E. tawaense sp. nov. A & B: Position ofconch section shown in Text-fig. 4 and themeasured points

and funnel like wall. Whorl nearly ashigh as broad or slightly higher thanbroad, with moderately convex flanks andmoderately rounded venter. Suture forms
very widely arcuate ventral saddle, very
shallow lateral lobe and a small andindistinct saddle on the umbilical wall.
Position of siphuncle unknown.
Remarks: Cross section of the whorl
is illustrated in Text-fig. 4-A and B. If
the un deformed shell is measured, theratio of the height to the breadth of thewhorl may be approximately 0.96. The
tangent line on the whorl side is almost

Measurments (in mm.) of Eutrephoceras tawaense sp. nov.
I
(A)

Preserved last part

I

I

(B)

Near the last septum---

Diameter
213.3
(100)
164.4
(100)

I

I
I

I

Umbilicus I
7. 7
(4)
(5)

I

I
I

I

Height
126.5
(59)
100.5
(61)

I

Breadth

B./H.

107.9
(51)
96.3
(59)

0.85
0.96

(A & B: shown in Text-fig. 3)

745.

Nautiloid from the Izumi Group

A

8

337

type-species, from the Upper Cretaceousof the Atlantic Coast of North America,
in its narrower whorl section and its
suture without ventral lobe.
With respect to the suture, the present
species is similar to the following species:

,

... ,

Text-fig. 4. Conch section and sutureof the holotype of E. tawaense sp. nov. A& B: shown in Text-fig. 3

parallel or slightly convergent. Venterand ventral shoulders are moderately
rounded_ There are about 5 camerae inan adoral quarter-volution. Umbilicus ismeasured about 7.7 mm. Shell surface ofthe preserved test as well as the internalmould is almost smooth without rursiradiate ribs of Cymatoceras type. The
lateral lobe of the last suture is moreconcave than the others, but it may havebeen secondarily modified as suggested bya crack. The restored original laterallobe is as shallow as the line shown in
Text-fig. 4-C. Eight septa are exposedon a part of the outer volution. The
septal distances on the mid-lateral of the
exposed part are measured in mm. inaboral order as follows:
13.3, 14.4, 15.5, 14.1, 21.7*, 18.2*, 17.5(*: on the crushed air-chamber)Comparison: Judging from the characterof the suture, shell form and the smoothsurface, the present species is assigned
to Eutrephoceras. It is clearly distinguishedfrom Eutrephoceras dekay (Morton)(Morton,
1834; see Miller and Garner, 1962), theEutrephoceras kobayashii :\1atsumoto, 1962,
Japan
E. kummeli Wiedmann, 1960, India, Madagascar and South Africa
E. meritenii Wiedmann, 1960, India, and Spain
E. montomollini (Pictet et Cam piche) , 1859,
England, France and Switzerland
E. neubergicum (Redtenbacher), 1873, Austria
and Germany
E. sp. Morozumi, 1979, Japan

E. kobayashii from the Upper Cretaceousof Hokkaido has a somewhat broader
whorl section, larger umbilicus and more
inflated flanks; E. kUl11meli (Wiedmann,
1960, p. 166-167, pI. 20, figs. B-D, pI. 25,
figs. 3, 6-7) has a more globular conch;
E. 71lonto71lollini (see Wiedmann, 1960, p.
164, pI. 19, fig. G, pI. 23, fig. H) has asubtriangular whorl section and moreglobular conch and E. meriteni (Wiedmann,
1960, p. 164-165, pI. 25, figs. 4-5, pI. 26,
figs. 5-6, text-fig. 7) an ovoidal whorl
section and more involute conch thanthe present species.
E. neubergicum seems to be very similar
to the present species in its restored shellform and pattern and density of septa.
The illustrated specimens from the Alps
(Hauer, 1858, p. 14, pI. 1, figs. 1-2;
Redtenbacher, 1873, p. 97-98, pI. 22, fig.
4) and Germany (SchlUter, 1876, p. 174175, pI. 48, figs. 3-5) are smaller andsomewhat deformed and the precise
comparison with ours is difficult.
Meanwhile, Wiedmann (1960) regarded"Nautilus" neubergicus from the Alps as
an example of Angulithes fieuriausianus(d' Orbigny) (Wiedmann, 1960, p. 183-185,
pI. 19, fig. A, pI. 20, figs. N, 0, pI. 21,

338

Mitsunobu FURUlCHI

figs. I, L, M, pI. 23, fig. 0, pI. 26, figs. 13, text-figs. 14-15) and transferred N.
neubergicus from Germany to Eutrephoceras darupense (Schluter) (Wiedmann,
1960, p. 157-158, pI. 21, figs. C (upper),D, pI. 23, figs. F, P, text-fig. 2). As isshown by Hauer's illustration (1858, pI. 1,
figs. 1-2), the Alpine specimen indeed
suggests a similarity to A. fleuriausianus,
but there still remain unsettled points. It
is doubtful whether the open umbilicus isreadily closed by the depression or not.
The illustrated syntypes of E. darupense(Sch1i.iter, 1876, p. 176, pI. 49, figs. 4-5)
have clearly broader septal interspaces
than E. neubergicum and a closed
umbilicus. One of the illustrated specimens of N. neubergicus by Schluter (1876,
pI. 48, fig. 5) shows a ventrally situatedsiphuncle like that of E. darupense, but
the siphuncle position of other specimens(Schluter, 1876, pI. 48, figs. 3-4) is notknown, as in the Alpine specimens. Fromthese points it is still premature to erase
the name E. neubergicum. Whether the
sub trigonal whorl section in the illustrated
type of "Nautilus" neubergicus Redtenbacher (Hauer, 1858, pI. 1, figs. 1-2) is as
a product of secondary deformation or theAngulithes type primary character is uncertain. The suture of Angulithes is, as arule, considerably sinuous. For the time
being, I follow Kummel (1956, p. 382), who
has assigned this species to Eutrephoceras.
(Prof. Matsumoto made an inquiry about
this point to Dr. Summersberger, but has
not yet received an answer.)E. sp., which was described by Morozumi(1979) from the Upper Cretaceous Izumi
Group of the Izumi Mountains, seems toresemble the present species, but the
specimen is too incompletely preserved
for the precise comparison.
In addition to these species, E. alsecence
Reeside (Reeside, 1927, p. 7, pI. 1-2, pI. 3,
figs. 1-5, pI. 5, figs. 1-2), from the Upper

Cretaceous of the Western Interior ofNorth America, should be taken into
comparison. I have excluded it from agroup of the above mentioned species forthe reason of its having a shallow ventral
lobe. Otherwise, this specise is especiallysimilar to the present species in the shellform, septal distance and size.
With respect to the sutural pattern, thepresent species may be compared alsowith the following Tertiary species of
Eutrephoceras:
Eutrephoceras bryani (Gabb), 1877, U. S. A.
E. cookanum (Whitfield), 1892, U. S. A.
E. japonicum (Shimizu), 1926, Japan
E. oregonense Miller, 1947, U. S. A.

Except for E. japonicum, the above
species have less inflated flanks, as in thepresent species. E. bryani (Miller, 1947,
p. 28-29, pI. 8, figs. 1-4, pI. 93, figs. 1-2),
however, is clearly distinguished by its
much broader umbilicus, broader spacingof suture and more compressed whorl
section at the adult stage.
It should, however, be taken into consideration that the septal distance may
change with growth. The measurements
shown in the preceding page suggestthat the septa may be somewhat more
widely spaced in immature stages thanin the adult stage of the present species.
Be that as it may, the present species canbe distinguished from E. bryani in otherpoints.
E. cook anum (Whitfield) (1892, p. 285286, text-fig. 2, pI. 48, fig. 1, pI. 49, figs.
4-5; Miller, 1947, p. 30-31, pI. 10, figs. 12, pI. 11, figs. 1-3, pI. 12, fig. 1) has amore rounded whorl section and somewhat larger umbilicus than the present
species. It has about 4 or 5 camerae in
a quarter volution, and is rather similar
to the present species as compared withthe others.
E. oregonense (Miller, 1947, p. 34, pI. 21,

745.

Nautiloid from the Izumi Groupfigs. 1-2) has a more or less subdiscoidal
conch and there are about 7 camerae in
the adoral quarter volution.
E. Japonicum (Shimizu, 1958, p. 25-27,
pI. 8, figs. 1-6; Kobayashi and Kamada,
1959, p. 109-110, text-figs. 2-3, pI. 9, figs.
1-2) is more globular in shell form withthe maximum breadth in the lower partof whorl section and there are only 3.5
camerae in a quarter volution, although its
sutural pattern closely resembles that ofthe present species.
Concluding Remarks

To sum up, the present new species
resembles certain Cretaceous and Early
Tertiary species of Eutreplwceras in shellform and suture. Especially it closely
resembles E. neubergicum from theUpper Cretaceous of Europe and somewhat similar to E. cookanum from theLower Tertiary of North America. It isalso suggested that it may foreshadowsome characters of Nautilus, such as theless inflation of flanks and the slightly
sinuous suture.
Acknowledgements

I am much indebted to Emeritus Professor Tatsuro Matsumoto of Kyushu
University and Professor Yuji Bando of
Kagawa University for their helpful
suggestions and critical reading of themanuscript. I also thank Drs. Ienori
Fujiyama, Masayuki Tashiro, Masayuki
Noda, Tomoki Kase, Hidekuni Matsuo,
Itaru Hayami and Yasuhiko Kamada for
their kind helps from their specialities.
I owe much to Mr. Tsutomu Ooyoshi,
chief of the Kagawa Natural Science
Museum, who has generously provided
facilities for the study.

339

References Cited
Gabb, W. M. (1877):
Notes on American
Cretaceous fossils with descriptions ofsome new species. Philadelphia Acad.
Nat. Sci. Proc., vol. 28, p. 276-324, pI. 5.
Hauer, F. von (1858): Die Cephalopoden der
Gosauschichten. Beitr. Palaeontographie
Osterreichs, vol. 1, no. 1, p.7-14, pI. 1-3,
Wien & OImtitz.
Kobayashi, T. and Kamada, Y. (1959): An
Eocene nautiloid from Kyushu, Japan.
Japan. Jour. Ceol. Ceogr., vol. 35, p. 105114, pI. 4.
Kummel, B. (1956): Post-Triassic nautiloid
Genera. Bull. Mus. Compo Zool., vol. 114,
no. 7, p. 324-494, pIs. 1-28.
Matsumoto, T., Hashimoto, H. and Furuichi,
M. (1980):
An interesting species of
Baculites (Ammonoidea) from the Cretaceous Izumi Group of Shikoku. Proc.
Japan Acad. vol. 56, ser. B, no. 7, p. 404413.
Matsumoto, T. (1967): A Cretaceous nautiloid
from Urakawa, Hokkaido. Japan. Jour.
Ceol. Ceogr., vol. 38, no. 2-4, p. 163-169,
pI. 3.
- - (1977): Zonal correlation of the Upper
Cretaceous in japan. Palaeont. Soc. Japan,
Special Papers, no. 21, p. 63-74.
Miller, A. K. (1947): Tertiary nautiloids of
the Americas. Ceol. Soc. Amer. Mem.,no. 23, p. 1-234, pIs. 1-100.
- - and Garner, H. F. (1962): Cretaceous
nautilides of New jersey. In Richards,
H. G. (ed.): the Cretaceous fossils of New
jersey, Part II. N. J. Bur. Ceol. Topogr.,
Palaeont. Ser., Bull. vol. 61, p. 101-111,
pIs. 65-67.
Morozumi, Y. (1979): A nautiloid from theUpper Cretaceous Izumi Group in the Izumi Mountains, Southwest japan. Bull.
Nat. Hist. Osaka Mus., no. 23, p. 11-18,
pI. 3.
Morton, S. G. (1834): Synopsis of the Organic
Remains of the Cretaceous Group of theUnited States. Philadelphia. p. 1-88, pIs.
1-19.
Nakagawa, C. (1961): The Upper Cretaceous
Izumi Group in Shikoku. Jour. Nat. Sci.

Mitsunobu FURUlCHI

340

Tokushima Univ. (Cakugei) , vol. 11, p.
77-124, figs. 1-8, tables 1-8.
Nakano, M. (1953): On the Izumi Group of
the central part of the Sanuki MountainRange, Japan. Ceol. Report, Hiroshima
Univ., no. 3, p. 1-14, (in Japanese withEnglish abstract).
Pictet, F. J. and Campiche, G. (1858-1860):
Description des fossiles du terrain cretacedes environs de Sainte-Croix. Geneve. p.
1-380, pis. 1-43.
Redtenbacher, A. (1873): Die Cephalopod enfauna der Gosauschichten in den nordostlichen Alpen. Abh. K. K. Ceol. Reichsanstalt. vol. 5, pt. 5, p. 91-140, pis. 2230.
Reeside, J. B. Jr. (1927): The cephalopods theEagle Sandstone and related formations in
the Western Interior of the United States.

U. S. Ceol. Surv. Prof. Paper. 151, p. 140, pis. 1-45.
SchlUter, C. (1876): Cephalopoden der oberen
Kreide. Paliiontgraphica, vol. 24, pt. 11,
p. 123-263 (1-144), pis. 36-55.
Shimizu, S. (1926): On two species of Nautiloidea from the Tertiary of Japan. Sci,
Rep. Tohoku Imp. Univ., ser. 2, vol. 9,
p. 26-27, pI. 8, figs. 1-6.
Whitfield, R. P. (1892): Gastropoda and Cephalopoda of the Raritan clays and Greensand marls. U. S. Ceol. Surv., Mon.
18, p. 1-402, pis. 1-50, (also issued as
N. J. Ceol. Surv., Palaeont., vol. 2).
Wiedmann, J. (1960): Zur systematik Jungmesozoisher Nautiliden (Unter besonderer
Berucksichtigung der iberischen Nautilinae
D'Orb.). Palaeontographica, vol. 115A,
p. 144-206, pis. 17-27.

Hokkaido: ~t$;@: Izumi:;frJ* Kagawa: 1ltJII Korobiishi: iJii;15 Nagao-cho: :!ltJ.T!1:IUJ Nakato:
IftJi Ookawa-gun: ::kJIIW Sanuki: ~JiJS( Shikoku: ggj;j!l Tawa:~;f!l Yezo: ~~ggj;j!lO)l:{fllBilli*;frJ*j.I~b> G£:/:I:llA.: *? A ;1]'1 ~{t15 (~fi): 1ltJlllliH;::7j-;(jl"tol:{fll Billi*;f!l*j.I~O) IftJiJi:!6-lIb> G ? A ;1]'1 ~{t15 b~£:/:I:l L t.:. ;:: n~ Eutrophoceras 0)
rfifie!:: L -ClBtI;lA.:. P.lill:11W~~-c-"'.:tb~'J\~ < illi~:(j(-c-;6o. ~fd\ljH1~o \"illl*Jll-c-)tli{fllt;::JA\,,~{flle!::{JtIJliit;::~< -CJAH(~{fll, "'.:t~t;::;6i IJ §:ll.t.:t.l:\"'J,~t.l:~{fll~t -?-C\"o.
~(J;.(O)~1r*!~~'0fie!:: L -c, Billi*c.~:MO) E. kobayashii, E. kummeli, E. meriteni,
E. montomollini, E. neubergicum e!::;f!l*JlJ~b>G¥jH!r~nt.:. E. sp., ~=:*c.fmmO) E.
bryani, E. cookanum, E. japonicum, E. oregonense b~;6 o. ;:: n G 0) ? t:> BlIUc.~:MO)
E. neubergicum (;:: EI Jltl;::~ Un'@;6 IJ) e!:: ~=:*c.fJJ:MO) E. cookanum b~~~O)7fJ:(j(t;::to\,,-C J: < ~{J;.( L-C\,'o.
t.l:to!JlJl~b~;6 i IJ ~n -Cto Gf ~1rjili[b~tI~t.l:b~ G l!U!l1l"to ;:: at Nautilus O)~:(j(",*nO)iifI~~~~~L-C\"oJ: ?I;::.~,?
£:/:I:lIIO):I1!!~n;fftI1 Baculites subanceps pacificus ~f:f-? ;:: e!:: e!:: .:tnb~*:I1!!ll.i1tO)i1!i1J"'eMetaplacenticeras subtilistriatum O)£:/:I:lII~O)l:(ft J: IJ £:/:I:l"t 0;:: e!:: b> Gl:{fll:IJ :/ /~.::. 7:/~.~~no.

~*~B

Explanation of Plate 54
Figs. 1-4. Eutrephoceras tawaense sp. nov.
1 & 3. Lateral views ...................................... x 2/5 (holotype, K-4-50)
2. Apartural view ........................................ x 2/5 (ditto)
4. Ventral view .......................................... x 2/5 (ditto)

FU R UICHI:

Nautiloid from the Izumi Group

Plate 54

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 126, pp. 341-355, pIs. 55, 56, June 30, 1982

746.

SOME CRETACEOUS ECHINOIDS FROM THE MONOBE
AREA, CENTRAL SHIKOKU*KEISAKU TANAKA
Geological Survey of Japan, Ibaraki 305

T AKESHI KOZAI
Odochi Primary School, Kochi 781-44

Abstract. This paper contains the systematic descriptions of some spatangoidechinoids from the Monobe area, Kochi Prefecture. The echinoids from the Bar·
remian Monobe·Yunoki Formation are assigned to Heteraster (3 species) and a new
genus (1 species), and a species from the Santonian part within the Kajisako Forma·tion to Epiaster. The Barremian species are common to those from other areas in
the Outer Zone of Southwest Japan. A new genus is established for a species charac·terized by having a single peripetalous fasciole. As an appendix an Albian species
belonging to this genus from another area in Japan is redescribed.

fossils, though in small numbers. All theechinoid specimens for the present study
were collected by the junior author (Kozai),and then their specific identification was
carried out by the senior author (Tanaka).
Thus, this paper is to present the resultof our study on these echinoids. Furthermore, appended to this paper by thesenior author in connexion with theestablishment of a new genus is thedescriptive remarks on a hitherto described
species concerned.
The repositories of the specimens
described here are as follows: OES=
Odochi Elementary School, Kochi; GSJ =Geological Survey of Japan, Tsukuba.
Before going further, we wish to
express our sincere gratitude to Professor
Emeritus Tatsuro Matsumoto of Kyushu
University for his valuable suggestions
and kindness in critical reading of theIntroduction

The Monobe area, Kochi Prefecture(Text·fig. 1) is known as one of theclassical and typical outcrop areas of theLower Cretaceous in Southwest Japan.
Recently a large number of molluscan
fossils were discovered at various horizons
within the Cretaceous strata of this area.
These fossils have been intensively studied
by specialists of respective fields. Con·
sequently, our knowledge about theCretaceous stratigraphy of the present
area have much increased. The up·todate stratigraphic context of the Monobe
Cretaceous has been described elsewhere(Tashiro, Kozai, Okamura and Katto, 1980 ;Tashiro, Kozai and Katto, 1980).
Echinoid fossils occur at several horizons
associated with abundant marine molluscan* Received August 3, 1981.

341

Keisaku TANAKA and Takeshi KOZAI

342

M-33

'&M-27

KAJISAKO

YunokiKa-05 X x
Ka-04

o,

Text-fig. 1.

2km
I

Localities of echinoid specimens in the Monobe area.

draft. We are also indebted to Dr. Masayuki Tashiro of Kochi University and
Dr. Ikuwo Obata of the National Science
Museum, Tokyo for their helpful advice,
to Professor Keiji Nakazawa of the
Department of Geology and Mineralogy,
Kyoto University for permission to examine
the collection, and to Mr. Yoshio Masaiof the Geological Survey of Japan forphotographing the specimens.

Localities of Echinoid
Specimens

The details of the biostratigraphy of
the Cretaceous strata in the Monobe area
have been published elsewhere (Tashiro,

Kozai, Okamura and Katto, 1980; Tashiro,
Kozai and Katto, 1980). The Lower Cretaceous of this area is divided in ascendingorder into the Monobe Formation (Lower
Barremian), the Yunoki Formation (Upper
Barremian) and the Hibihara Formation(Aptian-Albian), which are about 400 m,
750 m and 800 m thick respectively. Eachformation consists of sandstone, partly
conglomeratic, in the lower part andmudstone in the upper part. The Lower
Cretaceous echinoid examined came fromlocs. M-27 and M-33 of the Monobe
Formation and from locs. Ka-04 and Ka05 of the Yunoki Formation (Text-fig. 1).
All these localities are occupied bysiltstone or fine sandy siltstone. Loc. Ka04, among others, yields a number of746.

Cretaceous Echinoids from Monobe

echinoids. The Upper Cretaceous strata,
about 250 m thick, comprise the Nagase
Formation (Cenomanian) below and theKajisako Formation(Turonian-Lower
Campanian) above. The Nagase Formation
is dominated by sandstone, and theKajisako Formation by mudstone. Only{)ne Upper Cretaceous specimen available
for this study was obtained from loco M14 of the Upper Santonian part within
the Kajisako Formation (Text-fig. 1).
Locs. Ka-04 and Ka-05 are situated at
Kahoku-cho, Kami-gun, Kochi Prefecture,and locs. M-27, M-33 and M-14 at
Monobe-mura, Kami-gun.
Systematic Descriptions

Subclass Euechinoidea Bronn, 1860
Superorder Atelostomata Zittel, 1879
Order Spatangoida Claus, 1876
Suborder Toxasterina Fischer, 1966
Family Toxasteridae Lambert, 1920
Genus Epiaster d'Orbigny, 1853
Remarks:- The genus Epiaster has been
regarded as a synonym of the genus
Heteraster by Fischer (in Moore, ed.,1966). But, following Mortensen's (1950)
classification of the Echinoidea, we separate
Epiaster from Heteraster as a distinct
genus. Actually, Epiaster is characterized
by, among other features, the frontal
ambulacrum having rounded or commashaped pore pairs separated by a granuleand by the paired ambulacra which arepetaloid, generally depressed, and moreor less closed distally, with nearly equal
poriferous zones. On the other hand, in
Heteraster the frontal ambulacrum has
regularly or irregularly alternating larger
and smaller pores and the anterior paired
ambulacra are composed of unequal poriferous zones. Thus, the two genera aredearly distinguishable from each other.

343

Epiaster sp. aff. E. nobilis Stoliczka
PI. 55, Fig. 1
Compare:1873. Epiaster nobilis Stoliczka, Mem. Ceol.
Surv. India, Palaeont. Indica, ser. 8,
vol. 4, pt. 3, p. 20, pI. 3, figs. 7-8.

Material :-OES. 14007A, B (A, internalmould; B, imperfect external mould of
aboral surface), from loco M-14.
Description :- This is secondarily deformed. The test is medium-sized, andseems to have a rather roundly subovate
outline which is somewhat longer thanwide. The frontal sinus is rather shallow,
becoming much shallower near theambitus; thus the anterior notch, if any,
is probably indistinct. The apical system
is subcentral, and its structure is uncertain
because of the poor preservation.
The frontal ambulacrum is rather
shallowly sunken and subpetaloid; at the
middle of the petaliferous part the poriferous zones are somewhat narrower than
the interporiferous ones. Its pore pairs
are small, comma-shaped, circumflexed,
and are separated by a granule. The
paired ambulacra are wider than thefrontal ambulacrum, nearly straight,
shallowly sunken, petaloid, and more orless closed distally. The anterior paired
petals are slightly longer, slightly wider
and much more divergent than theposterior, extending about two-thirds theway to the margin. The pores of theanterior petals are transversely elongated,
and are somewhat longer in the posterior
poriferous zones than in the somewhat
narrower anterior; the pores of each pairare opposite and not conjugate. The
posterior petals extend about two-thirds
the way to the ambitus, and have pori ferous zones of nearly equal width, withsimilar elongate pore pairs.
The peristome is anterior and trans-

344

Keisaku TANAKA and Takeshi KOZAlversely oval. The plastron is amphisternous, probably protamphisternous. Periproct
is not observable. The aboral surface iscovered with numerous minute granules
and fine tubercles.
Peripetalous orlateroanal fasciole is not present.
Remarks :-Whether a subanal fascioleis present or lacking in the specimen
available can not be ascertained owing to
the poor preservation. Nevertheless, thepresent form is safely referable to Epiasteron the basis of the general features of
the test. This species closely resembles
Epiaster nobilis Stoliczka, from theSenonian (Arrialoor Group) of India, inmany respects. However, it differs fromthat species in that its paired petals are
more closed distally than in the Indianspecies. It is also to some extent similar
to an undescribed species of Epiasterobtained by the senior author from theAptian Hinagu Formation of Kyushu.
The Hinagu specimens, however, areconsidered as representing a distinct
species separated from the present species
by their paired ambulacra that are
narrower, more flexuous, less petaloid andmore open distally. Moreover, the present
form is easily distinguished from Epiasternutrix Lambert (1903, p. 86, pI. 3, figs. 35), from the Senonian of Madagascar, by
having nearly equal paired petals that are
longer, wider, less sunken and less closed
distally.
Occurrence :-A block at loco M-14,
middle part of the Kajisako Formation,siltstone inter laminated with fine sandstone, Upper Santonian because of theassociated occurrence of Inoceramus
japonicus Nagao et M<ltsumoto.
Genus Heteraster d'Orbigny, 185:1
Remarlzs :-The genus Paraheteraster
Nisiyama, 1968 (type-species: Washitaster(?) macroholcus Nisiyama, 1950, from theBarremian of Japan), is regarded as asynonym of Heteraster. This is to be
discussed in the later part of this paper.
Heteraster macroholcus (Nisiyama)
PI. 55, Fig. 21950. Washitaster (?) macroholcus Nisiyama,
Inst. Ceol. Palaeont.
Tohoku Univ.,
Short Papers, no. 1, p. 44, text-figs. 4-6.
1954. Washitaster macroholcus, Tanaka andOkubo, jour. Ceol. Soc. japan, vol. 60,
p. 221, pI. 7, figs. 1-2, text-figs. 5, 7.
1954. Washitaster barremicus, Tanaka andOkubo, Ibid., vol. 60, p. 220, pI. 7, fig_
3, text-fig. 7.
1965. Heteraster macroholcus, Tanaka, Trans_
Proc. Palaeont. Soc. japan, N. S., no_59, p. 133, pI. 16, figs. 3-4, text-fig. 4.
1968. Paraheteraster macroholcus, Nisiyama,
Palaeont. Soc. japan, Special Papers,
no. 13, p. 190, pI. 21, fig. 3; Ibid., no_II (1966), pI. 18, figs. 3, 12-13.
1968. Paraheteraster barremicus, Nisiyama,
Ibid., no. 13, p. 191.

Material :-OES. 04153,04177 and 04179.
from loco Ka-04, and probably also OES_
27006, from loco M-27.
Descriptive remarks :-In the present
species the frontal sinus is deep and very
long, with a distinct anterior notch, thefrontal ambulacrum shows an irregularalternation of several longer pores andone shorter pore in the outer row andsomewhat inward disposition of pores(slightly smaller than the normal roundto oval pores), opposite to shorter outerpores, in the inner row, the paired
ambulacra are flush, and the peristome isdeeply sunken. Furthermore, a broad,diffuse granular band or pseudo-fasciole
which is covered with streaks of granules
very slightly smaller than those covering
the rest of the aboral surface and decorated
with tubercles, as figured by Devries (1960,
pI. 16, figs. 5-6), surrounds the petals (Text-

746.

Cretaceo'us Echinoids from Monobe

4mm

L--...J

Text.fig. 2. Diagrammatic sketch of
granular band in Heteraster macroholcus(Nisiyama). GSJ. 6016, from the Arida
Formation (Lower Barremian), northeast of
Yuasa, Wakayama Prefecture.

fig. 2). It is about 6 mm wide in an about
70 mm long test. The largest (OES. 04153)
of the specimens available probably attains
about 70 mm in length.
Occurrence :-Loc. Ka-04, Yunoki Formation, fine sandy siltstone, Upper Barremian (OES. 04153, 04177,04179). Moreover, one specimen (OES. 27006) probably
identical with this species is found insandy siltstone of the Monobe Formation(Lower Barremian) at loco M-27.
Heteraster nexilis Nisiyama
pI. 55, Fig. 31950. Heteraster nexilis Nisiyama, Inst. Ceol.
Palaeont. Tohoku Univ., Short Papers,
no. 1, p. 42, text.figs. 1-3.
1968. Heteraster nexilis, Nisiyama, Palaeont.
Soc. Japan, Special Papers, no. 13, p.
186.

Material :-OES. 27001, 27002-1 and
27002-2, from loco M-27, and probably also
OES. 33012, from loco M-33.
Descriptive remarks:- This species ischaracterized by, among other features,the moderately deep frontal sinus, distinct
anterior notch, alternations of two to
three larger pores and one smaller pore345

at the middle of the frontal ambulacrum,and by the superficial paired ambulacra.
One (OES. 27001) of the specimens, thoughlongitudinally compressed secondarily, isprobably at least 27 mm long.
Occurrence :-Loc. M-27, Monobe Formation, siltstone, Lower Barremian (OES.
27001, 27002-1, 27002-2). Moreover, aspecimen (OES. 33012) probably identicalwith this species is found in fine sandy
siltstone of the Monobe Formation (Lower
Barremian) at loco M-33.

Heteraster sp.
PI. 55, Fig. 4

Material:- Two specimens, OES. 04154
and 04167, from loco Ka-04, and another
OES. 05057b, from loco Ka-05.
Description :-The specimens available
are poorly preserved and somewhat
deformed by the subsequent disfigurement.
The frontal sinus is rather shallow, becoming much shallower towards theambitus, thus the anterior notch, if any,
is probably indistinct.
The frontal
ambulacrum is rather shallowly sunken
and subpetaloid. Its poriferous zones
show regular alternations of a pair ofslit-shaped pores widely spaced and a pairof elongate oval pores close together, thepore pairs lining up in four files. However, in each specimen only a double pairof longer pores is placed opposite to eachother at the middle of the two poriferous
zones. The paired ambulacra are somewhat flexuous, slightly sunken and subpetaloid. The anterior paired ambulacra
consist of anterior poriferous zones withelongate oval pores close together andmuch wider posterior poriferous zones withslit-shaped pores widely set. The poriferous
zones of the posterior paired ambulacra
have slit-shaped pore pairs. The peristome
is transversely oval and shallowly sunken.

346

Keisaku TANAKA and Takeshi KOZAl

The plastron is mesamphisternous. The
periproct is longitudinal oval.
Remarks :-The present form is closely
similar in many respects to Heterasteryuasensis (Tanaka et Okubo) (1954, p.223,
pI. 7, fig. 6, text-fig. 7; redescribed by
Tanaka, 1965, p. 136. pI. 16, figs. 6-7,
text-fig. 6), from the Barremian of various
areas in the Outer Zone of Southwest
Japan. But the presence of double pairsof longer pores in the frontal ambulacrum
is not known in any specimens hitherto
referred to Heteraster yuasensis (instead,a double pair of smaller pores is exceptionally observed in either of the both
poriferous zones in some specimens), evenwhen the test is of normal size. Since
the present specimens are very small(probably about 12 mm long) and have no
more than twenty pore pairs in thepetaliferous part of the frontal ambulacrum,
they are considered as representing young
individuals. Therefore, it is suggested
that the presence of a double pair of
longer pores in the frontal ambulacrum
is not due to intraspecific variation in agiven species such as Heteraster yuasensis.
Double pairs of longer pores, if present,
may occur more commonly in the expectedlarger (adult) forms than in the smaller(young) forms of the species described
here. This, together with the slightly
depressed paired ambulacra, may remind
us of some similarity of the present form
to a Heteraster species, from the BarremianArida Formation of the Yuasa area,
Wakayama Prefecture (Tanaka and Okubo,
1954, p. 224, pI. 7, fig. 7, text-fig. 7),
which has been compared with Heterasterbohmi (Lorio!) (synonymous with H.
bravoensis (Bose) according to Cooke,1955) with some doubts. Anyhow, we
refrain from further taxonomic discussionof the present form, because of the verysmall size and poor preservation of thematerial available.

Occurrence :-Loc. Ka-04, Yunoki Formation, siltstone, Upper Barremian (OES.
04154, 04167); loc. Ka-05, Yunoki Formation, siltstone, Upper Barremian (OES.
05057b).

Suborder Hemiasterina Fischer, 1966
Family Hemiasteridae Clark, 1917
Genus Pseudowashitaster Tanaka nov.
Type-species: - Pseudowashitastermysticus Tanaka, sp. nov. (described below)
Generic diagnosis:- Test large or rather
large, oval in outline, with a very long,deep frontal sinus; anterior notch distinct.
Apical system far posterior, ethmophract,with four gonopores. Frontal ambulacrum
very long, deeply sunken; prre pairs
generally similar, not separated by agranule. Paired ambulacra flexuous, very
unequal in length and width; poriferous
zones unequal; anterior zones with minute,round pores; much wider posterior zones
with elongated pores. Anterior paired
ambulacra considerably narrow almost
throughout the way, much longer, muchnarrower, much less divergent than theposterior, very slightly sunken; posterior
paired ambulacra flush. Peristome faranterior; periproct high up on the posterior
truncated surface. Peripetalous fasciole
single, distinct; no other fascioles.
Occurrence :-So far as the material
available is concerned, this new genus
contains two species to be described
below, from the Barremian and theAlbian of Japan respectively. Moreover,an undescribed species probably referable
to this genus is found in the Aptian ofJapan.
Remarks:- This new genus is similar
to Washitaster in the general features of
the test. But the former differs from thelatter in that the peripetalous fasciole is746.

Cretaceous Echinoids from Monobe

not multiple but single and the pore pairsof the frontal ambulacrum are notseparated by a granule. Moreover, the
new genus has a much larger test thandoes Washitaster. On the other hand,
Pseudowashitaster resembles Heteraster inmany respects, but is clearly separated
from that genus by the presence of awell-defined, narrow, continuous peri petalous fasciole crowded by granules muchsmaller than those covering the rest of theaboral surface, and by the rather uniform
pore pairs of the frontal ambulacram. Heteraster has no more than a broad, diffuse
granular band or pseudo-fasciole decorated
with tubercles as figured by Devries (1960,
pi. 12, fig. 11; pI. 16, figs. 5-6). Regularly
or irregularly alternating larger andsmaller pores in the frontal ambulacrum
is usual in Heteraster, whereas in Pseudowashitaster somewhat inward placed outerpores (see Text-fig. 4), if present, are notcommon, their inconstant numbers andpositions being merely a matter of intraspecific variability. Furthermore, the considerable narrowness of the anterior paired
ambulacra also is one of the mostprominent characters of the new genus.
This feature, however, is not known in
any Heteraster species.
To sum up,
Pseudowashitaster is clearly distinguishable
either from Washitasterorfrom Heteraster.
The presence of a peripetalous fasciolealone suggests a close affinity of the new
genus to the hemiasterid echinoids. In
some of the specimens referable to thepresent new genus, however, the peripetalous fasciole is more or less poorly demarcated including very few granules orsmall tubercles in such a manner as
illustrated by Devries (1960, pI. 16, figs.
7-8), in very limited portions of the way.
Nevertheless, the peripetalous fasciole ofthe present new genus is far narrowerand far better defined than a broad,diffuse granular band or pseudo-fasciole347

as figured by Devries (1960, pI. 16, figs.
5-6), approaching to the genuine peri petalous fasciole as figured by him (1960, pI.
16, fig. 9) on the whole and is moreover
well traceable throughout the course. On
the other hand, so far as the generalfeatures of the frontal ambulacrum andpaired ambulacra are concerned, it isevident that Pseudowashitaster is moreclosely allied to Heteraster than toperipetalous fasciole-bearing hemiasterid
echinoids. Anyhow, attaching importanceto the presence of a peripetalous fasciolealone, one is strongly inclined to theconclusion that the present new genus is.
provisionally placed in the family
Hemiasteridae. On the other hand, apossibility can not be denied that the new
genus could be a specialized offshoot of
the Toxasteridae. Its exact taxonomic
position at family level will be reserved
for further study. Furthermore, it should
be noticed that Washitaster longisulcus(Adkins et Winton) (1920, p. 55, pI. 9, figs.
4, 8-10), from the Albian of Texas, whichappears similar to Pseudowashitasterspecies, has considerably narrow anterior
paired ambulacra. This may not bear agenetic relation to Pseudowashitaster but
may indicate a parallelism development
in different provinces.
Washitaster japonicus (Tanaka et Okubo)
which is now assigned to Pseudowashitasterwas placed by Nisiyama (1968) in thegenus Paraheteraster established by himself. But, according to Tanaka (1965),
Washitaster (?) macroholcus Nisiyama,
1950, the type-species of Paraheteraster,is characterized by, among other features,the pore arrangement of Heteraster (s. s.)
type (Devries, 1960) in the frontal ambulacrum and the lacking of genuine fasciole(instead, a broad, diffuse granular bandor pseudo-fasciole decorated with tubercles
surrounds the petals; see Text-fig. 2),
thus being referable to Heteraster. The

348

Keisaku TANAKA and Takeshi KOZAI

contrasting difference in width of theanterior paired ambulacra also makes
it impossible to place both Washitaster (?)
macroholcus and Washitaster japonicus in
one and the same genus. To sum up, it
is reasonable to regard Paraheteraster as
a synonym of Heteraster.
Pseudowashitaster mysticus
Tanaka, sp. nov.
PI. 55, Figs. 5-7; PI. 56, Figs. 1-2;
Text-figs. 3-4

1965. Washitaster (?) sp., Tanaka, Trans.
Proc. Palaeont. Soc. Japan, N. S., no. 59,
p. 138, pI. 16, figs. 8-9, text-fig. 7.

Type-specimens :-Holotype, GSJ. 6017,
represented by an internal mould, fromloco Ka-04, about 700 m south of Yunoki,
Kahoku-cho, Kami-gun, Kochi Prefecture,
Yunoki Formation, Upper Barremian;paratypes, GSJ. 6018 A, B (A, internalmould; B, external mould) and GSJ. 6019
(external mould), from the same locality.
Diagnosis :-Test large, oval, with thewidest point anterior to the midpoint, notgreatly constricted behind. Aboral surface
gently arched, highest near the apicalsystem. Frontal sinus very long, deep,
somewhat constricted and shallower
immediately behind the frontal margin,
nearly parallel-sided in the main part of
the way, extending to the peristome;
anterior notch deep. Apical system very
eccentric behind, ethmophract, with four
gonopores.
Frontal ambulacrum very long, deeply
sunken; poriferous zones broader thanhalf the width of the interporiferous zones.
Inner pores oval; outer pores elongatedsilt-shape, accuminated inwards, scarcely
or occasionally set more or less inwards;
pores of each pair opposite, rather close
together, not separated by a granule.
Paired ambulacra very unequal, flexuous,subpetaloid, open distally.
Anterior paired ambulacra very long,considerably narrow almost throughout theway, somewhat narrower than half thewidth of the frontal ambulacrum when

a

~
-~

.

b

-~ c
lOmm

L........J

Text-fig. 3. Suggested figure of Pseudowashitaster mysticus Tanaka, sp. nov.
Aboral view. Restored from the holotype
and paratypes.

Text-fig. 4. Pseudowashitaster mysticus
Tanaka, sp. nov., from loco Ka-04, Yunoki
Formation, Upper Barremian. a, Frontal
ambulacrum (Holotype, GSJ. 6017). b, Rightanterior paired ambulacra (Paratype, GSJ.
6019). c, Right posterior paired ambulacra(Paratype, GSJ. 6019).

7M.

Cretaceons Echinoids from Monobe

compared with each other near the midway between the apical system and theambitus, but rapidly widening near theambitus, very slightly sunken in thepetaliferous part extending about twothirds the way to the ambitus, diverging
at a smaller angle than do the posterior
paired ambulacra then turn forwards andlaterally, becoming almost straight, andthen diverge laterally and cross the ambitus
far anteriorly. Anterior poriferous zones
very narrow; pores minute, round, near
together. Posterior poriferous zones muchwider than the anterior; pores oval in
the inner row, elongate in the outer row;
pores of each pair opposite, rather close
together. Interporiferous zones wider
than the anterior poriferous zones.
narrower than the posterior poriferous
zones.
Posterior paired ambulacra shortest,wide, flush, diverging at a very large
angle and then converge laterally; petaliferous part extending about one-third theway to the margin. Anterior poriferous
zones much narrower than the posterior,composed of minute, round pores closely
spaced. Posterior poriferous zones consisting of oval inner pores and elongate
outer pores; pores of each pair opposite,rather near together.
Interporiferous
zones more or less wider than the posterior
poriferous zones.
Peristome far anterior, rather transversely oval. Periproct oval, verticallyelongate, high up on the posterior truncatedsurface. Tubercles on the aboral surface
small, of varying sizes, perforate, crenulate,scrobiculate; the largest scattered in theanterior interambulacral areas, becoming
smaller and closer along the frontal sinus
and near the ambitus; numerous granules
between tubercles. Peripetalous fasciole
crowded by much smaller granules single.
distinct, well defined, narrow, continuous,
running nearly in a straight line between

349

the anterior and posterior paired ambulacra, crossing the anterior paired
ambulacra at about one-fourth the way
from the ambitus; no other fascioles.
Measurements :-The holotype, thoughincompletely preserved, probably attains
70 mm or more in length. The peripetalous
fasciole is a little more than 1 mm widein the para types.
Remarks:- The three specimens available from the present area are incomplete
and secondarily depressed. Accordingly,
the specific diagnosis given here is basedon observations on several specimens(e. g. GS]. 6116, 6117) from the Upper
Neocomian (Barremian) Ishido Formationof the western Sanchu Graben, Nagano
Prefecture and one specimen from theBarremian Arida Formation of the Yuasa
area, Wakayama Prefecture (Text-fig. 5)
in addition to those from the Monobe
area. The Ishido specimens were described
previously as a species provisionallyreferred to Washitaster (Tanaka, 1965).
In the present study, the holotype is
not sufficiently well preserved and theparatypes are also fragmentary anddeformed by the subsequent disfigurement.

Text-fig. 5. Pseudowashitaster sp.
d. P. mysticus Tanaka, sp. nov. Diagrammatic sketch of a specimen of the Kyoto
Univ. Coll., from the Arida Formation(Lower Barremian), northeast of Yuasa,
Wakayama Prefecture. Aboral view.
x ca. 0.7.

350

Keisaku TANAKA and Takeshi KOZAI

Nevertheless, a suggested figure of thetest of the present new species, restoredfrom the holotype and para types, is shownin Text-fig. 3. Of course, a modificationof the horizontal outline of the test andthe position of the apical system, if
required, should be made, though probably
slightlY, when more specimens of goodpreservation are obtained. Moreover, wehave insufficient information about theside outline of the test, which is not
figured here. In a comparatively weakly
depressed specimen, registered GSJ. 6117,
from the Ishido Formation, the test
measures at least 13 mm high against thepresumed 30 mm length between the apicalsystem and the anterior margin. It
follows from this that the test is rather
low. A peripetalous fasciole, one of themost striking features of Pseudowashitaster,
is not preserved in the holotype becauseof the internal mould. It is, however,observable in the para types GSJ. 6018 B(Pl. 56, Fig. 2) and GSJ. 6019 (Pl. 56, Fig.
1) and also in an Ishido specimen (GSJ.
6117; Pl. 56, Fig. 3) and an Arida specimen (Text-fig. 5).
The outer pores in the frontal ambulacrum show intraspecific variation inarrangement. In some specimens (e. g.
holotype GSJ. 6017 and GSJ. 6117 fromthe Ishido Formation), one shorter poreis set more or less inwards every five
to seven longer pores near the middle
part of the poriferous zones and every
two to four longer pores near the distal
part. On the other hand, in a specimen(GSJ. 6116 from the Ishido Formation)
only one or two shorter pores are placed
somewhat inwards within a series ofabout 40 pores in the middle part of theporiferous zones.
This new species closely resembles
Pseudowashitaster japonicus (Tanaka etOkubo) (1954, p. 220, pl. 7, fig. 4, text-fig.
7) to be redescribed below, from theAlbian Yatsushiro Formation of Kyushu,
in the general features of the test. But
the peripetalous fasciole crosses theanterior paired ambulacra much lessdistally in the former species than in thelatter. Moreover, the outer pores in thefrontal ambulacrum are elongated slitshape and accuminated inwards in Pseudowashitaster nzysticus sp. nov., whereasin P. japonicus they are shorter andelongate oval.
In connexion with the occurrence ofPseudowashitaster in the Barremian andAlbian in Japan, it should be noticed thata specimen (GSJ. 6055) Qrovisionallyreferable to this genus was obtained by
the senior author from the Aptain Hinagu
Formation of Kyushu. This specimen (Pl.
55, Fig. 8) also is characterized by thelong, distinct frontal sinus, the frontal
ambulacrum composed of similar, elongate
pore pairs, and by the considerably narrowanterior paired ambulacra, although aperipetalous fasciole is not discernible
because of the internal mould. Furthermore, in the frontal ambulacrum three tofour shorter pores set somewhat inwards
are observed within a series of about 30
outer pores in the middle part.
The present species and Pseudowashitaster japonicus (Tanaka et Okubo) appear
similar to Washitaster longisulcus (Adkins
et Winton) (1920, p. 55, pl. 9, figs. 4, 8-10)
from the Albian of Texas, in many
respects. However, the formers have asingle peripetalous fasciole as against
multiple in the latter. In the above
Texas species the pore pairs of the frontal
ambulacrum are separated by a granule,but such is not the case with the Japanese
two species. Furthermore, a minor but
distinct difference is that the frontal
sinus is nearly parallel-sided in the mainpart of the way in the Japanese two
species, whereas in Washitaster longisulcus
it is biconvex-sided. It is also added here

746.

Cretaceous Echinoids from Monobe

that the test is larger in the formers thanin the latter.
Occurrence :-Loc. Ka-04, Yunoki Formation, siltstone, Upper Barremian (GSJ.
6017, 6018A, Band 6019).
Concluding Remarks

Summarizing the systematic descriptions,the following five species belonging tothree genera are identified on the material
from the Monobe area:
Epiaster sp. aff. E. nobilis Stoliczka
Heteraster macroholcus (Nisiyama)
Heteraster nexilis Nisiyama
Heteraster sp.
Pseudowashitaster mysticus Tanaka, sp.
nov.
Epiaster and Heteraster belong to the
Toxasteridae. Pseudowashitaster is a new
genus, assigned to the Hemiasteridae,
which is characterized by, among otherfeatures, a single peripetalous fascioleand considerably narrow anterior paired
ambulacra.
Epiaster sp. aff. E. nobilis was obtained
from the middle part (Upper Santonian)of the Kajisako Formation. Heteraster ct.
macroholcus and Heteraster nexilis camefrom the Monobe Formation (Lower
Barremian), and Heteraster macroholcus,
Heteraster sp. and Pseudowashitastermysticus from the Yunoki Formation(Upper Barremian). In addition to thespecies mentioned above, a regular echinoid
whose ambulacra are composed of trigeminate compound plates and a spatangoidechinoid were obtained from locs. M-51
and M- 03 (Text-fig. 1) respectively of theNagase Formation, but their genericdetermination is impossible because of thepoor preservation.
Furthermore, anechinoid fragment of unknown taxonomic
position is also found in the Hibihara
Formation.
Among the Lower Cretaceous echinoids,351

the two hitherto described species ofHeteraster are known to occur in the
Barremian of other areas in the Outer
Zones of Southwest Japan (Nisiyama,
1950; Tanaka and Okubo, 1954; Tanaka,
1965; Nisiyama, 1968). As mentioned
before, such is the case with Pseudowashitaster mysticus Tanaka, sp. nov. (e. g.
Tanaka, 1965). From the above facts it
is concluded that the correlation of theMonobe-Yunoki Formation based on themolluscan fanules (Tashiro, Kozai, Okamura
and Katto, 1980) and on the echinoid
fanules respectively are in harmony witheach other. However, Aphelaster serotinus
Tanaka et Shibata and Heteraster yuasensis(Tanaka et Okubo) which are known fromthe Barremian of various areas in theOuter Zone of Southwest Japan (Tanaka
and Okubo, 1954; Tanaka and Shibata,1961; Tanaka, 1965; Obata et al., 1979;
Hayashi et el., 1981), have not yet been
discovered in the Monobe area. The only
identified Upper Cretaceous echinoid isclosely allied to Epiaster nobilis Stoliczka
from the Senonian (Arrialoor Group) of
India. This is the first report of an
Epiaster species from Japan.
APPENDIX
Notes on Pseudowashitaster japonicus(Tanaka et Okubo)
by Keisaku Tanaka
This species originally referred co
Washitaster is represented by a single
specimen, from the Albian Yatsushiro
Formation of the Yatsushiro district,
Kyushu (Tanaka and Okubo, 1954, jour.
Ceol. Soc. japan, vol. 60, p. 220, pI. 7, fig.
4, text-fig. 7), which is the holotype bymono type. The original specific diagnosis
is mostly available, but it is too brief.
Accordingly, III connexion with the352

Keisaku TANAKA and Takeshi KOZAI

establishment of the new genus Pseudowashitaster and the description of its
type-species, the diagnosis of the present
species is improved in the lines to follow,
on the basis of the writer's subsequent
observations (PI. 56, Figs. 4-5; Text-fig.
6).
Material :-Holotype, GSJ, 6020, represented by an internal mould (A) andalso imperfect external moulds of theaboral (B) and oral surfaces (C). It was
obtained from the siltstone of the middle
part (Lower Albian) of the Yatsushiro
Formation.
Description:- The holotype is deformedby the subsequent disfigurement, and theoriginal size of the test is difficult toestimate. The test probably attains about

65 mm in length. Presumably, it hadoriginally an oval or elliptical outline.
The aboral surface is highest near the
apical system.
The frontal sinus is very long, deep,
somewhat constricted immediately behindthe frontal margin, and nearly parallelsided in the main part of the way,
forming a conspicuous notch in theanterior edge of the test then extends tothe peristome. The apical system is far
backwards and ethmophract, with four
gonopores, and the madreporite extends
as far back as to separate the posterior
genital plates, touching the posterior
occular plates.
The frontal ambulacrum is very longand deeply sunken, having poriferous

Explanation of Plate 55
Fig. 1. Epiaster sp. aff. E. nobilis Stoliczka
la, Aboral view, xL 1b, Oral view, xL OES. 14007 A, from loco M-14, middle part ofKajisako Formation, Upper Santonian.
Fig. 2. Heteraster macroholcus (Nisi yam a)
Aboral view, x 1. OES. 04153, external mould, from loco Ka-04, Yunoki Formation, Upper
Barremian.
Fig. 3. Heteraster nexilis Nisiyama
3a, Aboral view, x 1.2. 3b, Oral view, x 1.2. OES. 27001,from loco M-27, Monobe Formation, Lower Barremian.
Fig. 4. Heteraster sp.
4a, Aboral view, x 2. 4b, Oral view, x 2. OES. 04154, from loco Ka-04, Yunoki Formation,
Upper Barremian.
Fig. 5. Pseudowashitaster mysticus Tanaka, sp. nov.
5a, Aboral view, x 1. 5b, Oral view, x 1. Holotype, GSJ. 6017, from loco Ka-04, Yunoki
Formation, Upper Barremian.
Fig. 6. Pseudowashitaster mysticus Tanaka, sp. nov.
6a, Aboral view, xL 6b, Aboral view, x 1, external mould. Paratype, GSJ. 6018A, B,
from loco Ka-04, Yunoki Formation, Upper Barremian.
Fig. 7. Pseudowashitaster mystic us Tanaka, sp. nov.
Aboral view, x 1. Paratype, GSJ, 6019, external mould, from loco Ka-04, Yunoki Formation,
Upper Barremian.
Fig. 8. Pseudowashitaster(?) sp.
Aboral view, x 1.5. GSJ. 6055, from Yatsushiro district, Kumamoto Prefecture, upper partof Hinagu Formation, Aptian.
All specimens illustrated here are internal moulds, unless otherwise stated.
Photos by Y. Masai

TANAKA

and KOZAI:

Cretaceous Echinoids from Monobe

5b

40

Plate 55

746.

\~V

~a

-

"'-~

"

--

lmm

I

/

_II

~

~

I

'~1----~.
b'----'

Cretaceous Echinoids from Monobe

_____

I

/\'
'\

~

\~
,
d

~m

Text-fig. 6. Pseudowashitaster japonicus(Tanaka et Okubo), from the middle part
(Lower Albian) of the Yatsushiro Formation,
Yatsushiro district, Kumamoto Prefecture(Holotype, GSJ. 6020). a, Frontal ambulacrum. b, Right anterior paired ambulacra.
c, Right posterior paired ambulacra.

zones which are of about half the width of
the interporiferous zones. The poriferous
zones are composed of oval inner andelongate oval outer pores, and the poresof each pair are widely spaced without agranule in between. In the left poriferous
zone, five of the series of 75 outer poresare placed somewhat inwards with anirregular interval. The paired ambulacra
are very unequal in length and width,
flexuous, subpetaloid and open distally.
The anterior paired ambulacra are very
long and very slightly sunken in theirpetaliferous part, diverging at a rather
small angle.
They are considerably
narrow almost throughout the way andare much narrower than half the widthof the frontal ambulacrum when comparedwith each other near the midway between
the apical system and the ambitus. The
posterior paired ambulacra are very shortand flush, diverging at a very large angle.
In the both paired ambulacra, the anterior
poriferous zones are much narrower thanthe posterior, the former consisting ofminute, round pores close together and353

the latter of oval inner and elongate oval
outer pores each pore pair of which israther widely set.
The peristome is very eccentric in front
and poorly preserved. The plastron isamphisternous. The periproct is oval andvertically elongate. The tubercles are ofvarying sizes, perforate, crenulate andscrobiculate. They are rather widely
dotted on the aboral surface, becoming
smaller and closer along the frontal sinus
and towards the ambitus. The oral
surface is covered with rather closely
spaced tubercles which are somewhat
smaller than on the aboral surface. The
peripetalous fa sci ole crowded by granules
much smaller than those covering the restof the aboral surface is preserved in theanterior part of the test, and is single,
well defined and narrow (about 1 mmwide), crossing the anterior paired
ambulacra rather near the ambitus.
Remarks :-Whether peripetalous fasciole
is single or multiple (as in Washitaster)in the holotype can not be ascertained
because of the poor preservation. Nevertheless, this species is safely referable toPseudowashitaster on the basis of thegeneral features of the test.
The present species differs from
Pseudowashitaster mysticus Tanaka, sp.
nov. especially in course of the peripetalous fasciole and features of the pore pairs
in the frontal ambulacrum, as has been
described in the preceding main article.
Occurrence :-North of Tuzura, Sakamoto-mura, Yatsushiro-gun, Kumamoto
Prefecture, middle part of Yatsushiro
Formation, Lower Albian, siltstone (ColI.
T. Matsumoto and K. Kanmera).
References Cited
Adkins, W. S. and Winton, W. M. (1920):
Palaeontological correlation of the Frederiksburg and Washita formations in354

Keisaku TANAKA and Takeshi KOZAINorth Texas. Texas Univ. Bull., no.
1945, 84 p., 21 pIs.
Cooke, C. W. (1955): Some Cretaceous echinoids from the Americas. U. S. Ceol.
Surv., Prof. Paper, 264-E, p. 87-112, pIs.
18-29.
I'etudeDevries, A. (1960): Contribution
de quelques groupes d'echinides fossiles de
I' Algerie. Publ. Servo Carte Ceol. Algerie,nouv. serie, mem., no. 3, p. 1-278, pIs.
1-39.
Hayashi, T., Honda, M., Suzuki, T. andIwama, J. (1981):
Geological study ofthe lower Cretaceous Idaira Formation in
the northeastern area of Lake Hamana,
Central Japan. Bull. Aichi. Univ. Educ.,
vol. 30 (Nat. Sci.), p. 193-220, pIs. 1-8 (inJapanese with English abstract).
Lambert, J. (1903): Note sur quelques nouveaux Echinides cretaces de Madagascar.
Bull. Soc. Ceol. France, ser. 4, vol. 3, p.
75-88, pI. 3.
Moore, R. C. ed. (1966): Treatise on Invertebrate Paleontology, Part U, Echinodermata 3, vol. 2, 695 p.
Mortensen, Th. (1950): A Monograph of theEchinoidea. Vol. 5, pt. 1, Spatangoida I,a

432 p., 25 pIs. C. A. Reitel, Copenhagen.
Nisiyama, S. (1950): Two new species of the
Toxasteridae from Japan.
Inst. Ceol.
Palaeont. Tohoku Univ., Short Papers,
no. 1, p. 42-47.
- - (1966): The Echinoid fauna from Japanand adjacent regions. Part I, Palaeont.
Soc. Japan, Special Papers, no. 11, p. 1227, pIs. 1-18.
- - (1968): The echinoid fauna from Japanand adjacent regions. Part II, Ibid., no.
13, p. 1-491, pIs. 19-30.
Obata, I., Saka, Y., Matsukawa, M., Kase, T.
and Tanaka, K. (1979): Correlation of
the Cretaceous formations in the Shima
Peninsula. Mem. Natn. Sci. Mus., no. 12,
p. 73-82, pI. 9 (in Japanese with English
abstract) .
Stoliczka, F. (1873): Cretaceous faunas of thesouthern India. The Echinodermata. Mem.
Ceol. Surv. India, Palaeont. Indica, ser.
8, vol. 4, pt. 3, p. 71-127, pIs. 11-17.
Tanaka, K. (1965): Cretaceous echinoids fromthe Sanchu graben, central Japan. Trans.
Proc. Palaeont. Soc. Japan, N. S., no. 59,
p. 126-142, pIs. 15-16.
and Okubo, M. (1954): On some echi-

Explanation of Plate 56
Fig. 1. Pseudowashitaster mysticus Tanaka, sp. nov.
Aboral view, x3. Para type, GSJ. 6019, external mould, from loco Ka-04, Yunoki Formation,
Upper Barremian. A peripetalous fasciole is seen between two arrows.
Fig. 2. Pseudowashitaster mysticus Tanaka, sp. nov.
Aboral view, x3. Para type, GSJ. 6018B, external mould, from loco Ka-04, Yunoki Formation, Upper Barremian. A peripetalous fasciole is seen between two arrows.
Fig. 3. Pseudowashitaster mysticus Tanaka, sp. nov.
Aboral view, x 2. GSJ. 6117, external mould, from Ishido, Saku-cho, Minamisaku-gun, Nagano Prefecture, Ishido Formation, Barremian. A peripetalous fasciole is seen between two
arrows. Front to the right.
Fig. 4. Pseudowashitaster japonicus (Tanaka et Okubo)
4a, Aboral view. xL 4b, Oral view, xL Holotype, GSJ. 6020 A, internal mould, from
Yatsushiro district, Kumamoto Prefecture, Yatsushiro Formation, Lower Albian.
Fig. 5. Pseudowashitaster japonicus (Tanaka et Okubo)
Aboral view, x 3. Holotype, GSJ. 6020B, external mould, from Yatsushiro district, Kumamoto Prefecture, Yatsushiro Formation, Lower Albian. A peripetalous fasciole is seen between two arrows.
Photos by Y. Masai

TANAKA and KOZAI:C1"etaceous Echinoids j1"o?n Monobe

Plate 56

746. Cretaceous Echinoids from Monobe
noids from the Paleo-cretaceous of theYuasa district in the Kii Peninsula andthe Yatsushiro district in Kyushu. Jour.
Geol. Soc. Japan, vol. 60, p. 215-227, pI.
7 (in Japanese with English abstract).
and Shibata, M. (1961): A new speciesof Aphelaster from the Lower Cretaceousof Japan. Trans. Proc. Palaeont. Soc.
Japan, N. S., no. 42, p. 68-72, pI. 10.
Tashiro, M., Kozai, T. and Katto, J. (1980):
A biostratigraphical study of the Upper
Cretaceous formations at Odochi of Mono-

355

be, Shikoku. In Taira, A. and Tashiro,M., eds., Geology and Paleontology of the
Shimanto Belt-Selected Papers in Honor
of Prof. Jiro Katto, p. 83-94, pIs. 12-13,
Rinyakosaikai Press, Kochi (in Japanesewith English abstract).
- , - , Okamura, M. and Katto, J. (1980):
A biostratigraphical study of the Lower
Cretaceous formations of Monobe area,
Kochi Prefecture, Japan. Ibid., p. 71-82,
pIs. 10-11.

Arida 1Jm, Doiban ±F.S1ff, Hinagu B~R, Ishido :Q"~, Kahoku-cho 1if~tlDJ, Kajisako mfti:i;,
Kami-gun 1if~m, Kawanouchi JII / p;j, Monobe 4'kiflll, Odochi jeW, Sanchu IIp::p, YatsushiroAf\;, Yuasa ml:lt, Yunoki *rb/ *1!9004'kiflll:1ti!:f!ilZg[O) S:!IUc ? =- ft;p: f.Ii~lIr<*flIl, 4'kiflll:1ti!:f!ilZO)rflll s:iIl!*4'kiflll-*rb / *~g[O)?=-ft;Pt;::'0\,,-r Heteraster 311, wr~lwr1l~, J:flIlS:iIl!*mfti:i;.'1
I:j:rflll (-If':/ r =- 7:/) g[0) 'h O)t;:: '0 \" -r Epiaster 111~ OOUJIJ L. i;~4'ki~I¥.J~1lt~ IT"?
t.::.. wr1l1i.;j:!ll/*~ (/~1/~7:/J:flIl) ii'Gg[L, .:tLt;::~L-rwr~~:m:PIf,L, Pseudowashitaster mysticus t fl;15 Lt.::.. .: 0) 'h 0) Ii., I*l*JlJ:1ti!JlJI:j:r:1ti!lfIj*O):Q"~~~;fl:J~JlJlIr<ml:lt:1ti!)jO)1Jm~ (I. 'TtL 'h /~ 1/ ~ 7:/) ii' G {,g["1"7.l. wr~Ii., !M't;::.$-Q) .mJ:(E*klO)~~,ii,tJ:. I) .lii~I.'iltr~;!P:*, iltr;!P:*t;::;t,)~ 7.l~fLQ)tl.11{t;::.l: "? -r7i- Ji!.J:~f!:I."1" 7.l Washitaster ~Heteraster tli.IR~IJ~tL, Hemiasteridae fil-t;::~"1"7.l. mtEi<:Q)ijf, .:O)wr~Ii.~~1I(/~I/~ 7:/)t~~0)111ii'GtJ:.IJ,~Gt;::*~t;::;t,).:tG< /l'iJ:<E~tL7.l 111ii'~*lIr<Af\;:1ti!)jQ)B~R~(777-7:/) ii'G 'h.;r,..I.,t.:~tL7.l.
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June 30, 1982

New Series No. 126

CONTENTS
TRANSACTIONS
743. TAKAHASHI, Kiyoshi: Miospores from the Eocene Nanggulan Formation in
the Yogyakarta region, central lava .................................. 303
744. OMURA, Akio: Uranium-series age of the "Kametsu Formation", Riukiu
Limestone on the Tokuno-shima, Ryukyu Islands ...................... 327
745. FURUICHI, Mitsunobu: A new nautiloid species from the Upper Cretaceous
Izumi Group of Shikoku .............................................. 334
746. TANAKA, Keisaku and KOZAI, Takeshi: Some Cretaceous echinoids from theMonobe area, central Shikoku.......................................... 341