ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No.

152

Pa laeontologica l Society of Japan
December 30. 1988

Co-Editors

Hisayoshi lGo and Tsunemasa SAITO

Editorial Board: Yoshikazu HASEGAWA, Itam HAYAMI, Hisayoshi IGO (chief), Makoto KATO,
Itam KOIZUMI, Yasumochi MATOBA, Kei MORI, Hiroshi NODA, Kazunari TANABE,
Tsunemasa SAITO, Kazuhiko UEMURA, Akira Y AO
Officers for 1987-1988
President: Tatsuaki KIMURA
Honorary President: Teiichi KOBAYASHI
Councillors:

Kiyotaka CHINZEl, Takashi HAMADA, Yoshikazu HASEGAWA, Itam HAYAMI,

Hisayoshi IGO, Junji ITOIGAWA, Tadao KAMEl, Tatsuaki KIMURA, Tamio KOTAKA,Kei MORI, Ikuwo OBATA, Tsunemasa SAITO, Yokichi TAKAYANAGI, Kazushige
TANABE, Toshimasa TANAIMembers of Standing Committee: Kiyotaka CHINZEI (Membership), Takashi HAMADA (Foreign Affairs), Itam HAYAMI (Finance), Hisayoshi IGO (Co-Editor of Transactions),
Ikuwo OBATA (General Affairs), Sumio SAKAGAMI (Co-Editor of Special Paper),Kazushige TANABE (Planning), Yokichi TAKA YANAGI (Editor of "Fossils"), Juichi
YANAGIDA (Co-Editor of Special Paper)
Secretaries:

Katsumi ABE (Planning), Takeshi ISHIBASHI (Special Paper), Kunihiro ISHIZAKI("Fossils"), Tomoki KASE (General Affairs and Membership), Hiroshi NODA (Editorof Transactions), Temfumi OHNO (Membership), Tatsuo On (Planning), Kazuhiko
UEMURA (General Affairs), Toshiyuki YAMAGUCHI (Finance)
Auditor: Noriyuki IKEY A

The fossil on the cover is Vicarya yokoyamai Takeyama, an Early Middle
Miocene gastropod from the Kurosedani Formation at Kakehata, Yatsuo-cho,
Nei-gun, Toyama Prefecture, central Japan (Collected by T. Kotaka and K.
Ogasawara, IGPS No. 99075, photo by S. Ohtomo and Y. Kikuchi, xO.9).

All communication relating to this journal should be addressed to the

PALAEONTOLOGICAL SOCIETY OF JAPAN% Business Center for Academic Societies,
Yayoi 2-4-16, Bunkyo-ku, Tokyo 113, Japan

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 152, pp. 603-624, 9 Figs., December 30, 1988

867. NEW TAXA IN THE LATE TRIASSIC DAEDONG FLORA,
SOUTH KOREA. PART 1*
T A TSUAKI KIMURATokyo Gakugei University, Koganei, Tokyo 184

and
BONG-KYUN KIM
Seoul National University, Seoul, lSI, Korea

Abstract. From the Late Triassic 'Daedong System' of non-marine origin distributed in thesouthern part of the Korean Peninsula, we recognized a flora consisting of 79 species belonging
to 38 genera. Of these, eleven species belonging to nine genera are new to science and many planttax~ are newly found. This paper deals with the description of the following taxa recognized by
us In the Daedong flora: Annulariopsis bunkeiensis (Kobatake), n. comb., Diclyophyllul11 exile(Brauns) Nathorst, Hausl11annia ussuriensis Kryshtofovich, Cladophlebis l11ungyeongensis, sp. nov.,
Chiropleis coreanica, sp. nov., Anomozamites mungyeongensis, sp. nov. Otozal11ites l11icrauritus,
sp. nov. and Pterophyllum micraequale, sp. nov. In addition taxonomic description is further refinedin this paper on Coreanophyllum variisegmenlllll1 Kimura et Kim, a bennettitalean with bipinnate
habit of leaf.

Amisan, Jogyeri, Baegunsa and Seongjuri Formations and the Bansong Group in the Mungyeong
Coal-Field into the Buunryeong, Bolim, Dangi,
Dangog, Maseong, Bongmyeongsan andBongmyeongri Formations in upward sequence
respectively.
Unfortunately, stratigraphical correlation
among the above-mentioned coal-fields has notbeen made precisely because of being destitute ofguide fossils and appropriate key-beds.
The fossil plants of the 'Daedong System' werestudied by Yabe (1922), Yabe and Oishi (1929),
Kawasaki (1925, 1926, 1927, 1939), Oishi (1940),
Kon'no (1944), Kobatake (1954), Kimura et al.
(1982) and Kimura and Kim (1982).
Since 1974, we collected a number of fossil
plants from the new fossil-sites of the Amisan
Formation (Chungnam Coal-Field) andBongmyeongri Formation (Mungyeong Coal-Field)
and we made revision of the previous works on thebasis of specimens newly collected and recognized

Foreword

N on-marine sedimentary rocks yielding
abundant fossil plants are distributed in the
Daedong, Kimpo (Gimpo), Yeoncheon,
Chungnam, Mungyeong and Danyang Coal-Fields
in the Korean Peninsula (Kimura and Kim, 1984a,
1985).
These strata are called the Daedong Group in
the Daedong Coal-Field (North Korea), Kimpo
Group in the Kimpo and Yeoncheon Coal-Fields,
Nampo Group in the Chungnam Coal-Field andBansong Group in the Mungyeong and Danyang
Coal-Fields respectively, but these sedimentaryrocks have customarily been called the collective
and non-committal 'Daedong System' or 'Daedong
Supergroup' .
The Nampo Group is divided into the Hajo,* Received June 9, 1987: Revised manuscript accepted
November 9, 1988.
603

604

Tatsuaki KIMURA and Bong-Kyun KIM

79 species belonging to 38 genera (Kimura andKim, 1984a, 1985; Kim and Kimura, 1986). We callthe plant-assemblage known from the 'Daedong
System' the Daedong flora for the sake of
convenience.
Among them, the following taxa are new toscience: Cladophlebis mungyeongensis, Chiropteris
coreanica, Anomozamites mungyeongensis,
Otozamites micrauritus, Pterophyllum cheondaeriense, P. micraequale, Anthrophyopsis decurrens, Taeniopteris cheondaeriensis and T.
mungyeongensis. In addition there are Annulariopsis bunkeiensis (Kobatake), n. comb. and Corea nophyllum variisegmentum Kimura et Kim.
We described the above-mentioned new taxa in
our two serial papers, together with the description of the following characteristic taxa: Dictyophyllum exile (Brauns) Nathorst, Hausmannia
ussuriensis Kryshtofovich, Drepanozamites sp. A,
Podozamites ex gr. schenki Heer, P. sp. A,
Cycadocarpidium sp. cf. C. asaense Kon'no, C.
sp. cf. C. nagatoense Kon'no and C. sp. A.
All plant taxa from the Chungnam, Mungyeong
and Danyang Coal-Fields will be described in a
separate paper by J. H. Kim and Kimura in thenear future, together with a detailed comparisonof the Daedong flora with coeval floras and discussion of Late Triassic phytogeography in East Asia.
It becomes evident that the geological age of theAmisan and Bongmyeongri Formations is Late
Triassic on the basis of their floristic composition(Kimura and Kim, 1984b) and that of the Baegunsa
Formation is considered to be also Late Triassic
on the basis of the presence of Lobatannularia, oneof the important elements of the PermianCathaysia flora (Kim, J. H. and Kimura, 1988).
The age-determination mentioned above is supported partly by the occurrence of abundant andvaried conchostracans indicating the Late Triassic
age, from the Amisan Formation and Kimpo
Group (e.g. Kobayashi, 1975). However, theprecise ages of the Daedong Group, Seongjuri
Formation in the Chungnam Coal-Field and Bansong Group in the Danyang Coal-Field are still
uncertain. We are of the opinion that the geologicalage of the 'Daedong System' is mostly Late
Triassic except for the uppermost layers in mostcoal-fields, such as the Seongjuri Formation which
is barren of fossils.
The specimens here described are kept in the
National Science Museum of Korea, Daejeon City.
The localities of fossil plants are as follows:
Amisan Formation in the Chungnam Coalfield:
Jogaegog (or Jogaegol): Misan-myeon,
Boyeaong-gun, Chungcheongnam-do(roughly 126°40'00"E, 36°18'54"N).
Dobung Coal-Mine: Ditto (roughly
126°40'00"E, 36°17'48"N).
Seongjuri: Ditto (roughly 126°38'18"E,
36°20'00"N).
Cheondaeri (Cheondae or Yesan CoalMine): Changgog-myeon, Hongseonggun, Chungcheongnam-do (roughly
126°46'00"E, 36°31 '54"N).
Bongmyeongri Formation in the Mungyeong
Coal-Field:
Hanaeri: Maseong-myeon, Mungyeong-gun,
Chungcheongnam-do (roughly 128°05'
lO"E, 36°39'18"N).
We thank Dr. Kazuo Asama, National Science
Museum, Tokyo, Prof. Kiyoshi Takahashi,
Nagasaki University, Prof. Hisayoshi Igo, University of Tsukuba and Prof. Chang-Hi Cheong,Seoul National University, who cooperated with
us in collecting fossil plants. Our thanks are extended to Miss Tamiko Ohana, Tokyo Gakugei
University who made a clear line-drawing of thosespecimens here described. The present study was
supported by the Grant-in-Aids (Overseas Scientific Research Fund) of Ministry of Education,
Science and Culture; nos. 1974-7433,
1981-56041024 and 1982-57043021.
Systematic description

Equisetales
Genus Annulariopsis Zeiller, 1903
The relation between Lobatannularia Kawasaki,
1927 and Annulariopsis Zeiller, 1903 has been
discussed by such authors as Halle (1927), Yabe
and Koiwai (1928), Kawasaki (1934), Kon'no andAsama (1950) and Harris (1961). Annulariopsis has
been defined somewhat vaguely for whorls of small

867.

New Late Triassic plant-taxa from Korea J

lanceolate uni-nerved leaves borne in a terminal
position due probably to breaking off of a stemor bud above. According to Harris (1961), it isdistinguished from Phyllotheca, Schizoneura and
Lobatannularia by having leaves which are freeeven at their bases. It differs from the leafy twigs
of Neocalamites chiefly in that the Neocalamites
shoots do not originally break just above a leafywhorl but bear successive whorls.
However, we now think that it is close in form
to Annularia and Lobatannularia because of therecent discovery of shoots bearing successive
whorls as in Annulariopsis annularioides and A.
lobatannularioides from the Upper Triassic
Yanchang Formation (Huang and Zhou, 1980).
Annulariopsis differs from Annularia in its leaves
being free to their bases except for about 1 mm
at the base, instead of the leaves forming a basal
leaf-sheath as in Annularia. On the other hand,
Lobatannularia is easily distinguished fromAnnulariopsis by its whorl consisting of markedly
anisophyllous and largely fused leaves. Thus
Annulariopsis is a valid genus and diagnosis of thegenus will be emended as follows:
Emended diagnosis: Articulate plant. Stem
slender, marked with longitudinal ridges; internodes rather long. Leaf-whorls circular or oval,
often divided into two lobes, spread out in one
plane, obliquely attached to the stem and persistent. Leaves equal or sometimes unequal in size.
Adjacent leaves united for about 1 mm at the base,then free. Leaf linear, linear-lanceolate, or broadly
spatulate in form with a round or mucronate apex,
generally broadest at or near distal end anduninerved; margins entire or sinuous. Stem easy
to break off at the node; thus most of thespecimens showing the leaves in a terminal position on the shoot. (Fructification unknown.)

Annulariopsis bunkeiensis (Kobatake)
Kimura et Kim, comb. nov.
Figures la-b

Pseudolobatannularia? bunkeiensis Kobatake : 1954, p. 73,
figs. 5-6.
Annulariopsis inopinata Zeiller : Kawasaki, 1939, p. 8, pI.
I, fig. I.

605

Material: Lectotype: Reg. no. 81-303(Jogaegog). Paralectotype: Reg. no. 81-089(Dobung Coal-Mine). Other specimens: A numberof specimens were collected from Jogaegog andDobung Coal-Mine. Locus typicus: Jogaegog.
Stratum typicum : Lowest Member of the Amisan
Formation. Derivatio nominis: Bunkei-originallyadopted by Kobatake in Japanese means
Mungyeong in Korean.
Emended diagnosis: Stems slender, 1.5 mm
wide, its surface-ornamentation indistinct. Internode more than 2 cm long. Leaf-whorl oval, notdivided into two lobes, spread out in one plane,
obliquely attached to the stem and persistent;
smaller whorl, 1.2 cm (major) and 0.8 cm (minor),and larger one more than 2.6 cm (major) and 2.0
cm (minor) in diameter. Uninerved leaves morethan 10 in a whorl, unequal in size, broadly
spatulate in outline with mucronate apex andbroadest near the distal end; largest one 2.5 cm
long and 0.8 cm wide; margins entire or faintly undulated. Leaves mostly free but adjacent ones
united for about 1 mm at the base. Midnerve thick,persisting to the tip.
Distribution and occurrence: Annulariopsis
bunkeiensis is quite abundant but fragmented in
the lowest Member of Amisan Formation at
Jogaegog and Dobung Coal-Mine, where its
broken leafy stems coexist with several species ofconchostracans and rhizomes of Neocalamites
carrerei (Kimura et al., 1982). This species has beenrecorded by Kawasaki (1939) as Annulariopsis
inopinata from the Bansong Group at Yeonhari(Danyang Coal-Field) and by Kobatake (1954) asPseudolobatannularia? bunkeiensis from theDangi Formation at Gujeomri (Mungyeong CoalField). However, their precise locality is uncertain.
Discussion and comparison : Many specimens
were present at these localities but none of themwere complete. We could not show any
photographs of them, because they were allpreserved as faint impressions in a black shale.
This species was originally described by
Kobatake (1954) as Pseudolobatannularia ?
bunkeiensis as mentioned above. But his conceptof the genus Pseudolobatannularia on the basis of
detached leaf-whorls from the Norian Nariwa

606

Tatsuaki KIMURA and Bong-Kyun KIM

Group, Japan is said to be characterized by its leafwhorl consisting of more than 30 uninerved leaves
usually united with each other for most of theirlength. Thus, his Pseudofobatannufaria ?
bunkeiensis belongs to Annufariopsis of which aleaf-whorl consists usually of 10 uninerved leaves
united with each other only at their bases.
Our leafy-stems, detached leaf-whorls and asingle leaf-whorl described by Kawasaki (1939)
as Annufariopsis inopinata agree well' with
Kobatake's leaf-whorls in all the features. Thus,
we propose Annufariopsis bunkeiensis (Kobatake)as a newly combined name herein.
Since Zeiller (1902-3) established Annulariopsis
and described A. inopinata from the Upper
Triassic of Hongay Coal-Field, several species havebeen described mainly from older Mesozoic rocksof Eurasia.
Annulariopsis inopinata is distinguished fromA. bunkeiensis by its rather large-sized leaf-whorlwith linear or elongate-oblanceolate leaves. Among
our collection, there is no such a leaf-whorlreferable to Zeiller's species.
Oishi (1930, 1932a, 1940) and Oishi and
Takahasi (1936) described three imperfect leafwhorls as Annufariopsis inopinata ? from the
Nariwa Group and a single imperfect leaf-whorl
from the Carnian Momonoki Formation. Unfortunately, they are insufficient for definite
determination.
Annufariopsis inopinata described by Kiritchkova (1969) from the Upper Triassic of the
Cheliabinsk Basin, East Ural is also distinguishedfrom our specimens by its leafy stem bearing ra~herlarge-size leaf-whorls with linear leaves.
Annulariopsis inopinata described by Yabe
(1922) from the Upper Permian Gobangsan
Formation, Korea is now r~garded as Lobatannufaria ensifolia Halle (Kon'no and Asama, 1950).
Annufariopsis simpsoni (Phillips) Harris known
from the Middle Jurassic of Yorkshire (Harris,
1947, 1961) and Middle Asia (Sixtel et af., 1971)
is distinguished from the present species by its
markedly sinuous leaf-margins. However, the leafwhorls described by Jacob et al. (1955) as
Annulariopsis simpsoni from the Jurassic of
Afghanistan are rather similar in form to the pre-sent species because of their entire leaves.
Annulariopsis annularioides and A. lobatannularioides originally described by Huang andZhou (1980) are similar in form and size to the present species, but they are distinguished by the leafwhorls consisting of eight leaves with an acutely
pointed apex and by those divided into two lobes,
respectively.
Neocalamites annularioides originally describedby Sze (1956a) from the Upper Triassic in the
Junggaer (Dzungaria) Basin is similar in form andsize to the present species, but it is difficult to makea precise comparison by its published characters.
Hexaphyllum sinensis originally described by
Ngo (1956) from the Upper Triassic Xiaoping
Formation and later regarded as Annulariopsis ?
sinensis (Ngo) by Lee (in Sze et al., 1963) is easily
distinguished by its much larger leaf-whorls.
Annulariopsis oishii, the youngest recorded
genus, described by Kimura and Hirata (1975)
from the Lower Cretaceous Ryoseki Formation,
Japan differs from the present species in the leaves
bearing deeply sinuous or entire margins and an
acutely pointed apex.
Filicales
Family Dipteridaceae
Genus Dictyophyllum Lindley et Hutton, 1834
Dictyophyflum exile (Brauns) Nathorst
Figures 2a-h
Diclyophyl/um exile (Brauns) Nathorst, 1878b, p. 39, pI. 5,
fig. 7; Nathorst, 1906, p. 7, pis. 4-5, pI. 6, figs. 1-22,
pI. 7, figs. 2-11, text-figs. 1-3: Harris, 1926, p. 64,
pI. 1, figs. 1-2, text-fig. 7B: Harris, 1931, p. 80, pI.
13, figs. 15-16: Genkina, 1966, p. 62, pI. 10, fig. 7,
pI. 11, figs. 1-5, pI. 12, figs. 1-3: Zhou, 1978, p. 102,
pI. 18, figs. 2-3: Hsii el al., 1979, p. 23, pI. 27, figs.
2-6, pI. 28. For further references, see Nathorst, 1906
and Harris, 1931.

Material: Examined specimens; Reg. no.
A-5019 and 12 other specimens (Seongjuri) andReg. no. T -024 and eight other specimens(Cheondaeri).
Description : The petioles are represented onlyby their distal part each with two arms leaving

867.

New Late Triassic plant-taxa frol11 Korea 1

between both an oval space (Fig. 2b). They arepreserved presumably as their original state; bothdistal and proximal ends of each arm are missing.
Each arm bears more than 17 crowded pinnae (e.g.
Fig. 2a), more than 17.5 cm long. The pinnae are
slender, linear or linear-obI an ceo late in outline,
pinnatifid into triangular segments (or pinnules)
which are directed forwards, with a pointed apex
and with straight or shallowly undulated margins;
the incision is shallow and does not reach to a halfof the length of segments. The pinna rachis israther thick, 2 mm wide, and gradually narrows
proximally into a long and slender basal part withan extremely narrow sterile lamina, 1-2 mm wide
and 1-1.5 cm long, nearly entire at margin; theweb of the basal lamina of the adjacent pinnae isnarrow and sterile (Figs. 2c and 2e). There are aconsiderable number of isolated pinnae, varying
in size and form; the widest one is 4.5 cm (Fig.
2d). The venation is of the typical Dictyophyllul11type. The mid nerve of each segment is strong,straight, persists to the tip and gives off secondaries
at an angle of about 60° in proximal part and at
about a right angle near distal end. They bifurcate
twice or thrice in typical manner of sympodial
ramification and form polygonal meshes, with one
row of especially large-sized primary meshes oneach side of midnerve.
The fertile pinnae are similar in size and form
to sterile ones; their laminae appear to be thicker
than sterile ones. The sporangia are about
400-500 .urn in diameter, with 5-7 grouped into
a definitely uniseriated sorus, 1-1.5 mm indiameter (Fig. 2f) ; crowded sori are distributed
over the whole under surface of the lamina (Fig.
2d).
Distribution and occurrence: Locally abundant in
the Chungnam and Mungyeong Coal-Fields. Thisis the first occurrence of Dictyophyllum in the
Daedong flora.
Remarks: All the specimens collected fromvarious localities and horizons in the surveyed area,
are common in almost every important character,thus they belong to the same species and are identified to Dictyophyllum exile.
The present species is characterized by crowded
sori distributed over whole under surface of the607

pinna consisting of 5-7 sporangia and narrow
web of basal lamina of the adjacent pinnae.
The isolated pinnae of this species resemble
those of Dictyophyllum nilssoni, D. nathorsti, D.
falcatum and Camptopteris japonica.
Dictyophyllum nilssoni (Brongniart) Goeppert
is similar to pinna form to the present species, but
is distinguished by the shorter arms and thesporangia not arranged in sorus (Harris, 1931).
Dictyophyllum nathorsti Zeiller is also similar in
pinna form and soral character to the present
species but differs in its basal web of the pinnae
which are concrescent for a length of 5-8 cm, instead of nearly free as in the present species.
Dictyophyllum falcatus Naito (Kon'no, 1968)is also similar in pinna form and soral characterto the present species, but it differs in its shorterand small-sized arms.
Camptopteris japonica (Yokoyama) Kon'no(Kon'no, 1968) formerly defined as Dictyophyllum japonicum, is similar in pinna form andsoral character to this species, but it differs
essentially in its spirally twisted arms.
Genus Hausmannia Dunker, 1846

H ausmannia (Protorhipis) ussuriensis
Kryshtofovich
Figures 3a-b

Hausmannia ussuriensis Kryshtofovich, 1923, p. 295, fig. 4:
Kryshtofovich and Prynada, 1932, p. 367, pI. 2, figs.
4-5: Sze et al., 1963, p. 89, pI. 26, figs. 3-4, pI. 28,
fig. I.
Hausmannia cf. ussuriensis Kryshtofovich: Sze, 1933c,
p. 66, pI. 9, figs. 1-6.
Hausmannia nariwaensis Oishi, 1930, p. 52, pI. 7, figs. 2,
2a: Oishi, 1932a, p. 303, pI. 38, figs. 1-10 : Oishi andYamasita, 1936, p. 163: Oishi, 1940, p. 220, pI. 5, figs.
2-3.

Material: Examined specimens; Reg. no. TA-7
(Seongjuri) and T-OI05A-C (Cheondaeri).
Description: Four sterile leaf-fragments wereobtained. Two of them are petiolate; the petiole
is very slender, 1 mm wide, but its length isunknown. The lamina is broadly reniform inoutline and varies in diameter from 2.5 to 7.5 cm.
At the base, the lamina makes a deep and narrowsinus reaching to the centre. Both edges of the sinus

608

an d Bong-K y U n KIM
Tatsuakl. KMURA
I

2c

1a

~

l mm

~

2b

2f

.

867.

New Late Triassic plant-taxa from Korea 1

are convex outwards and overlap with each other.
Primary nerves are 12 in number, radiate throughthe lamina from the top of petiole, brancheddichotomously 2-4 times, and reach to the marginof lamina. The finer nerves originate nearly at aright angle or sometimes at narrower angles fromthe primary nerves, and are subdivided and united
to form small square or polygonal meshes. The
margin of lamina is irregularly, broadly andshallowly undulated.
Distribution and occurrence: The present
species is rare and is only known from the Amisan
Formation. This is the first occurrence in the
Daedong flora.
Remarks: Although Kryshtofovich's original
specimens were roughly drawn, we consider that
Hausmannia ussuriensis and H. nariwaensis belong
to the same species as already mentioned by Sze(1933c). The former name has priority. The present leaves are close to those of Hausmannia nariwaensis described by Oishi (1930, 1932a, 1940)
from the Nariwa Group.
Hausmannia ussuriensis resembles the following species in outline of lamina, but it isdistinguished by various reasons as briefly mentioned below:
Hausmannia crenala (Nathorst) Moeller, 1902, p. 50, pI. 5,
figs. 5-6: Lamina is semicircular with a deep mediansinus. Similar lamina was reported from the Liassic
Fuxian Formation (Ordos Basin) as Hausmannia
papilionaceae Liu (in Huang and Zhou, 1980).
Hausmannia denlala Oishi, 1932a, p. 306, pI. 21, figs. 1-4,
SA, pI. 35, figs. 2-3, text-fig. 2: A lamina with strongly
sinuate-dentate margin characterizes this species.
Hausmannia leeiana Sze, 1933b, p. 7, pI. 2, figs. 8-9: A
large-sized lamina (18-19 cm wide) is a characteristicof this species.

609

Unclassified fern
Form-genus Cladophlebis Brongniart, 1849

Cladophlebis mungyeongensis Kimura et Kim,
sp. nov.
Figure 4a-d

Cladophlebis nebbensis Brongniart: Kawasaki, 1939, p. 19,
pI. 5, figs. 18-19, pI. 6, fig. 20a, text-fig. 3a.

Material: Holotype ; Reg. no. 81-021 (A-D)(Hanaeri). Paratypes ; Reg. nos. 81-040 and 093(Hanaeri). Stratum typicum: Bongmyeongri
Formation. Locus typicus : Hanaeri (west-side).
Derivatio nominis: After Mungyeong Coal-Field
where the holotype was collected.
Diagnosis: Frond bipinnate, medium-sized, 50
cm long, lanceolate in outline and up to 20 cm
wide. Rachis up to 2 mm wide with wings, 0.5 mm
wide along its lateral sides. Pinnae set closely, alternate, mostly perpendicular or sometimes obliquely
at an angle of 45° to the rachis and nearly parallelsided for the most part; apex acuminate. The
longest pinnae on the middle portion of a frond
10 cm long and 1.5 cm wide at an interval of 1.5
em, overlapping each other laterally and bearing
16 pairs of katadromically arranged pinnules. Bothdistal and proximal pinnae set rather remotely at
an interval of 1.7-1.8 em, directed forwards andbackwards, respectively and bearing a reduced
number of pairs of pinnules. Pinnules set closely,
typically deltoid in form, sometimes falcate andattached to the pinna axis by their whole base atan angle of from 35° to 90°; margins finely serrate;
apex acute or sometimes rounded. Basiscopic basal
pinnule not specialized but slightly reduced in sizeand originating from the angle between rachis and~ Figures I (a-b) and 2 (a-f). 1. Annulariopsis bunkeiensis (Kobatake) Kimura et Kim, comb. nov.: la. An
incompletely preserved leaf-whorl consisting of uninerved, unequal-sized and broadly spatulate leaves, each of Whichpossessing a macronate apex (Lectotype, Reg. no. 81-303). Loc. Jogaegog. lb. A leaf-whorl and successive stem(paralectotype, Reg. no. 81-089). Loc. Dobung Coal-Mine. 2. Diclyophyl/um exile (Brauns) N athorst: 2a. A bendingarm; the web of the basal lamina of adjacent pinnae being sterile and narrow (Reg. no. B-7011). Loc. Seongjuri.
2b. Distal part of two arms (Reg. no. T-0009). Loc. Cheondaeri. 2c. A pinna most of which is fertile (Reg. no. A-5020).
Loc. Seongjuri. 2d. A large-sized fertile pinna (Reg. no. A-5040). Loc. Seongjuri. 2e. Drawn from 2a. 2f. A sorus
enlarged, but details of it uncertain, drawn from 2d.

610

Tatsuaki KIMURA and Bong-Kyun KIM

867.

New Late Triassic plant-taxa from Korea 1

pinna axis. Midnerve distinct, persisting to the tip
and typically sending off 6-7 pairs of once forked
lateral veins; each serration receiving a set of lateralvein. (Petiole and fructification not known).
Distribution and occurrence: The present
species is locally common in the Mungyeong CoalField.
Discussion and comparison: The holotype
represented by almost whole leaf except its petiole
is characterized by its main rachis with narrow
lateral wings and by deltoid pinnules with finely
serrate margins and typically with 6-7 pairs ofonce forked lateral veins.
Cladophlebis nebbensis described by Kawasaki(1939) from the Mungyeong Coal-Field is referable
to the present species in its form and venation of
pinnules.
The present species resembles Todites fukutomii
described in detail by Kimura and Ohana (1980),
but it is distinguished by pinnules having serrated
margins instead of the entire margins of the latter.
The present species is also distinguished from
Todites denticulatus (or Cladophlebis denticulata)
by its wholly deltoid pinnules.
Pteridospermales
Genus Chiropteris Kurr, 1845

Chiropteris coreanica Kimura et Kim, sp. nov.
Figures Sa-i

Material: Holotype; Reg. no. 81-009Ta(Cheondaeri). Paratypes; Reg. nos. T-0029b, d,
81-017Te, g (Cheondaeri). Examined specimens:
Reg. nos. 81-007T, 009Tf, 022Te, 032T, 035T,
T -0082 (Cheondaeri), 81-149, 242 (Hanaeri).
Stratum typicum: Amisan Formation. Locus
typicus: Cheondaeri (Cheondae Coal-Mine).
Derivatio nominis: After Korea.
Diagnosis: Leaf petiolate; petiole long andnarrow, more than 2 cm long and up to 2 mm

611

wide, consisting of a centrally projected and cylindrical part and flat lateral filmy fringes of about 0.5
mm wide. Lamina obdeltoid in form with broadly
rounded apical margin; basal angle 50° -90°;
lateral margin entire but apical margin shallowly
undulated. Leaves varying in size; laminae typically 2.5 cm high; the largest one 3 cm high and thesmallest one 0.5 cm high. Veins very fine, radiatingfrom the top of petiole, repeatedly forking
dichotomously at all levels, and reticulated into
polygonal or rhomboidal meshes; meshes rare inoccurrence near the base of lamina, elongated at
the middle region, 1.3 cm long and 0.5 mm wide,
but becoming shorter towards the apical margin,
where 2.5 mm long and 0.5 mm wide. (Cuticle notpreserved and fructification not known.)
Distribution and occurrence: The present
species is locally common at the Cheondae CoalMine and Hanaeri.
Discussion and comparison: Since Lilienstern(1931, 1932) reviewed ten species of the genus
Chiropteris known at that time, so far as we know,
eight additional species have been described under
this genus from Mesozoic plant-beds.
Most Chiropteris species hitherto known are
represented by digitate leaves in agreeing with theimportant diagnostic character of this genus given
by Schenk (1864) and Schimper (1869).
However, Chiropteris coreanica is characterized
by rather small-sized and fan-shaped leaves withentire or shallowly and irregularly undulated apicalmargin, and fine and equidistant veins. Accordingly, Chiropteris species with digitate leaves areeasily distinguishable from the present species.
The following Chiropteris species would becomparable with the present species in their entire
or shallowly undulated margin of leaf: Chiropteris
barrealensis Frenguelli, 1942, C. cuneata(Carruthers) Seward, 1903, C. kawasakii Kon'no,
1938, C. manasiensis Gu et Hu, 1979, C. reni-+-- Figures 2 (g-h)-4(a-b). 2. Dictyophy/!um exile (Brauns) Nathorst: 2g. A fertile branch (proximal part missing)(Reg. no. T-024). Loc. Cheondaeri. 2h. Main part of a sterile pinna (Reg. no. TA-T-001). Loc. Cheondaeri. 3.
Hausmannia (Protorhipis) ussuriensis Kryshtofovich: 3a. A broken sterile lamina (surface view) (Reg. no. T-OlOSA).
Loc. Cheondaeri. 3b. Drawn from 3a, showing the venation. 4. Cladophlebis II/ungyeongensis Kimura et Kim, sp.
nov.: 4a. A pinna with large-sized pinnules (paratype; Reg. no. T-OI16). Loc. Cheondaeri. 4b. Middle portion ofa leaf, drawn from 4c.

612

Tatsuaki KIMURA and Bong-Kyun KIM

Figures 4 (c-d)-S(a-c) (A ll natural size). 4. Cladophlebis l11ungyeongensis Kimura et Kim, sp. nov.: 4c-d .
A single steri le leaf (4d shows an upper part of 4c in organic connection) (Holotype; Reg. no. 8 1-093). Loc. Hanaeri.
5. Chiropleris coreanica Kimura et Kim, sp . nov.: Sa. A typical petiolate leaf (Holotype; Reg . no. 81-009Ta). Loc .
Cheondaeri. Sb. A small-sized leaf (paratype; Reg. no. 81-0 17Te). Loc. Cheondaeri. Sc. A laterally expanded leaf
(Reg. no. T-0082). Loc. Cheondaeri.

867.

New Late Triassic plant-taxa from Korea 1

613

5h
5f

5g

1/

form is Kawasaki, 1925 and C.? yuaniSze, 1956a.
Chiropteris barrealensis known from the Upper
Triassic of Argentina (Frenguelli, 1942) is easily
distinguishable by its longitudinally elongate leaf
with markedly auriculated base. Chiropteris
reniformis known from the Upper Permian
Gobangsan Formation, Korea (Kawasaki, 1925,
1931, 1934) and the Permian of China [Upper
Permian Shihezi (Shihhotse) Formation, Shanxi(Halle, 1927) and its equivalents in Hebei, Anhui
and Jiangxi, and Lower Permian Lower Shihezi
Formation (Gu and Zhi, 1974)], resembles the present species in leaf-form and venation, but it isdistinguished by its large-sized leaves (some exceeding 12 cm wide) and reniform laminae withdeeply cordate base (Kawasaki, 1931, pI. 72, fig.
201). In the present leaves, all the laminae bear anacute or wedge-shaped base and no lamina withcordate base is known. Chiropteris ? yuani fromthe Upper Triassic Keramayi Formation, Xinjiang(Sze, 1956b) resembles the present species in leafform and size, but it is distinguished by having
longer and wider vein-meshes.
Chiropteris manasiensis from the Keramayi
Formation (Gu and Hu, 1979) is similar to C. ?
yuani in leaf-form, size and venation, and it is also
distinguished from the present species by havingcoarser venation (up to 10 per cm at the middle
portion of lamina).
Chiropteris kawasakii from the Gobangsan

cm

2

]

Figures S (d-j): Chiropteris coreanica
Kimura et Kim, sp. nov.: All from Cheondaeri.
Sd. Drawn from Sa. Se. Lateral lamina (rightside) is partly folded (paratype; Reg. no.
T-0029b). Sf. (Reg. no. 81-022T). Sg.
(paratype; Reg. no. T-0029d). Sh. Drawn from
Sb. Si (Reg. no. 81-009Tf). Sj. An exceptionallysmall-sized leaf (para type; Reg. no. 81-0 17Tg).

Formation (Samcheog, Ganweondo and near
Pyeongyang City, North Korea: Kon'no, 1938) iseasily distinguishable from the present species by
having large-sized leaf (ca. 11 cm wide) with thickveins (or ribs).
Cyc!opteris cuneata originally described by
Carruthers (1872) from the Upper Triassic of
Queensland, and assigned later to Chiropteris
cuneata (Carruthers) by Seward (1903) together
with other leaves from the Upper Triassic of South
Africa, is most close to the present species in leafform, size and venation. However, it is difficult
to compare closely with the present species, because
this species from the Southern Hemisphere has notadequately been described.
Kon'no (1938) mentioned that no specimen ofwell-defined Chiropteris had ever been found in
the Late Triassic-Early Jurassic floras in Japan,
Korea and North China, despite its occurrence in
non-Asian regions. Now, we are certain that leaves
assignable to Chiropteris are present in the Late
Triassic floras of Korea (the present work) and of
China Chiropteris ginkgoformis Liu (MS) (in Fenget al., 1977; Xiaoping Formation, Guangdong), C.
manasiensis Gu et Hu (I. c.), c. taizihoensis Zhang(Zhang et al., 1980; Middle Triassic Linjia Formation, Liaoning), C. ? yuani Sze (/. c.).
Arthrophyopsis? sp. of Yokoyama (1905, p.
12, pI. 3, fig. 6) from the Nariwa Group is too
fragmented to compare closely with the present

614

Tatsuaki KIMUR A and Bong-Kyun KIM

Figures 6 (a-f). Anol11ozal11ileS l11ungyeongensis Kimura et Kim, sp. nov.: All from Hanaeri . 6a. Detachedleaves (paratype; Reg . no. KTM-46A). 6b. A typ ical leaf, its both ends are missing. Most segments (pinnae) are
truncated distall y (Holotype; Reg. no. KTM-15A). 6c. Drawn from 6b. 6d. An irregularly segmented leaf. Distal
margi n of segment (pinna) is occasionally rounded (paratype; Reg. no. KTB-1SB). 6e. Basal part of lamina (Reg.
no. KTM-1Sc). 6f. Occurrence of this speci es (Reg. no. KTM-46A).

867.

New Late Triassic plant-taxa fr om Korea I

61 5

Figures 7 (a-f). Coreanophy lluin variisegl1len{uln Kimura et Kim : Fro m Ha nae ri unless o th erwise menti o ned .
7a . A thick rac his still bea ring pin nae co nsidered to be im mat ure or abo rti ve o nes o n th eir la tera l sid es (H o lo type,Reg. no. 81·108). 7b. Ditto (para type; Reg. no . 8 1-253) . Arrows A a nd B show th e rachi ses. 7c. A full grow n pinna(para type; Reg. no. T 8 1-040). Loc . C heond aeri. 7d. Hairs o n th e rachi s (pa ra type; Reg. no . 8 1·1 28). 7c. A thickrachi s with num erous hair-scars o n its surface (Reg . no. 8 1·346). 7f. Draw n pa rtl y from Fig. 7d . All figures fro m
Kimura and Kim, 1982).

Tatsuaki KIMURA and Bong-Kyun KIM

616

7r

3cm
L..----'------''----''

7j

7p

7l'

0; ••

.:~~/.
7h..
'.::.\' 7h' x2

'

.

.".0

'

:.:/:::. Ijj)~~!~

..
r

::·.Y

~
7l"

)~

867.

New Late Triassic plant-taxa from Korea 1

species.
So far as we know, Chiropteris has not beenrecorded from the Jurassic plant-beds and therange of this genus is thought to be from Permianto Triassic in age.
Bennettitales
Genus Anomozamites Schimper, 1870

A nomozamites mungyeongensis
Kimura et Kim, sp. nov.
Figure 6a-f

Material: Holotype; Reg. no. KTM-15A(Hanaeri). Paratypes ; Reg. nos. KTM-15B, 46A.
Examined specimens; about 200 in number.
Stratum typicum: Bongmyeongri Formation.
Locus typicus: Hanaeri (west side). Derivatio
nominis: After Mungyeong, the largest town near
the locality.
Diagnosis: Leaves shortly petioled, long andnarrow, nearly parallel-sided for the most part,small-sized, up to 10 cm long and typically 1 cm
wide, noted range between 0.4 cm and 1.4 cm.
Lamina attached to the lateral sides of slenderrachis, suddenly narrowing near the base, regularly
divided into about 20 alternate pairs of nearlysquare or short-rectangular pinnae; sinus reaching
to the rachis, 1 mm wide; apex typically truncatedbut sometimes more or less rounded. Basal pairof pinnae extremely small. Pinna base sometimes
slightly expanded. Veins numerous, mostly simple,
arranged in parallel, typically 15-18 in each pinna(40-42 per cm). (Cuticle not preserved. Reproductive organs not known.)
Distribution and occurrence: The present new
species is only known from Hanaeri and is quite
abundant, sometimes occurring en masse with noadmixture of other taxa.
Discussion and comparison: Anomozamites

617

mungyeongensis sp. nov. is characterized by its
slender leaf of which lamina is divided into nearlysquare or short-rectangular pinnae ending truncated distal margin and each with 15-18 simple
and parallel veins. The pinnae are nearly uniformwith the exception of the small basal ones. The
typical length-to-width ratio of pinnae is 1.15,
noted range between 0.75 and 1.70.
One might get confused Anomozamites
mungyeongensis with Pterophyllum micraequale
and Coreanophyllum variisegmentum. In
Pterophyllum micraequale, however, most of
pinnae are longer, the typicallength-to-width ratiois 3.5, noted range between 1.3 and 4.5 and eachpinna bears 8-13 veins.
In Coreanophyllum variisegmentum, the typicalratio is 6.0, noted range between 1.5 and 15, andeach pinna possesses 3-6 veins. Thus,
Anomozamites mungyeongensis is clearly
distinguishable from these two species except forthe case represented only by the basal part of leaves
or pinnae.
The following species are similar to
Anomozamites mungyeongensis in leaf-form, butthey are distinguished from the present species by
respective reasons as briefly mentioned below:
Anol1lozamites gracilis Nathorst, IS76, IS7Sc: Distal
margin (or apex) of pinnae not truncated but mostly
rounded.
A. inconstans (Braun): Zeiller, 1902-3 and Sze et al., 1963:
Leaves twice or thrice as large as those of the present
specimens and distal margin of pinnae rounded.
A. loczYi Schenk: Schenk, ISS4 and Sze et al., 1963: Similarin size and form to A. inconstans. In the pinna base,lower veins decurrent and upper veins bending upwards.
A. millor Nathorst (non Kawasaki, 1926): Nathorst, IS7Sc(including A. gracilis), IS79; Harris, 1926, 1932b: Basalregion of leaves often entire like the lamina of
Taeniopteris. Most of recorded leaves twice as large as
the present specimens. Distal margin of pinnae rather
rounded.
A. nilssoni (Phillips): Harris, 1969: Leaf not parallel-sided~ Figures 7 (g-r). Coreanophyllul11 variisegmentum Kimura et Kim: From Hanaeri unless otherwise mentioned.
Natural size, unless otherwise indicated. 7g. A rachis-fragment still bearing two pinna bases. Small circles show hair-scars
on its surface (Reg. no. SI-346). 7h. Hair-scars (Reg. no. SI-346). 7h'. Detail of hair-scars enlarged partly from
7h. 7i. Drawn partly from 7a. 7j. Drawn partly from 7b. 7k. Rachis with pinna in organic connection (Reg. no.
SI-OSI). 71. Drawn from 7c.71'-1". Detailed venation drawn partly from 7c. 7m-q. Various pinna forms (7m,o-q; Reg. no. TSI-OOI, 7n; Reg. no. T75-250). Loc. Cheondaeri. 70'. Drawn partly from 7c. 7r. Occurrence ofdistorted and detached penultimate pinnae. All figures from Kimura and Kim, 1982 except for 7r.

61B

Tatsuaki KIM URA and Bong-Kyun KIM

3cm.. .... ... ..... ..

~. ;~'~-.-. ~.-. --.:.--.:.,-.••::..::.:==:\I,~

--

I: ;

1,---'-----8a

8a"

8b

F igures 7 (s- I}- 8 (a-b). Natural size un less oth erwise ind icated. 7. Corean ophy llul17 variisegmentul17 Kimuraet Kim : 7s (upper) . Distal pa rt of penultim ate pinn ae; ma rgins of segment s are reflexed (Reg. no. TBI- IOl ). 7s Gower).
Basal pa rt of a penu ltim a te pi nna. 71. A rac his bear ing hair-sca rs on its under sur face (Reg. no. Bl- 12B; enlarged
t wice). Loc. C heo nd aeri a nd Hanaeri respectively. 8. Otozamiles m icrauritus Kimura et Kim , sp . nov .: 8a, 8a ' , 8a".
Draw n fr o m 8e. 8b. Basa l pa rt of a leaf (pa ralype; Reg. no. Bl-153) . Loc. Ha naeri.

867.

New Late Triassic plant-taxa from Korea 1

for the most part but elongate-lanceolate in form, maximum width 4.5 cm. Number of veins fewer (5-10 at
pinna base). Distal margin of nearly all the pinnae finely
denticulated.
A. thomasi Harris, 1969: Leaf large-sized, attaining 20-30
cm long. Distal margin of some pinnae slightly waved
or hollowed.

Nilssonia formosa Vachrameev et Vasina (inGenkina, 1963) has pinnae of a similar form to
those of the present pinnae, but they are twice aslarge as the present pinnae.
Genus Coreanophyllum Kimura et Kim, 1982

Coreanophyllum variisegmentum
Kimura et Kim
Figures 7a-t

Coreanophyllum variisegmentum Kimura et Kim, 1982, p.
152, figs. 1-12.
A nomozamites cf. nathorsti (Schenk): Kawasaki, 1925, p.
41, pI. 40, fig. 111.
A nomozamites minor Kawasaki, 1926 (pars), p. 30, pI. 7,
fig. 22b (non 22a).
Pterophyllum nathorsti Schenk: Kawasaki, 1939, p. 25, 52,
pI. 9, figs. 38-40, text-fig. 4.

Examined specimens: About 1000 in number.
Distribution and occurrence: Detached pinna
fragments are quite abundant, sometimes thickly
massed and appressed in the Amisan andBongmyeongri Formations.
Comparison and remarks: The characteristic
bipinnate habit of this plant was already mentioned
in our previous paper (Kimura and Kim, 1982). We
included various specimens described by Kawasaki(1925, 1926, 1939) in our synonimic list. The
following species are represented by simply pinnate
leaves and are recorded from Japan, China, North
Viet Nam and East Siberia. They resemble orpartly resemble detached pinnae of CoreanophyUum variisegmentum, but they are distinguished from our species because of various
features as mentioned below:
Pterophyllum angustum (Braun) Gothan: Yabe and Oishi,
1929 (Upper Triassic Anyuan Formation, Hubei); Szeet al., 1963 (ditto); Oishi, 1935 (cf. P. angustum; Lower
Cretaceous Muling Formation, Heilongjiang); Zhang,
et al., 1980 (ditto); Oishi and Huzioka, 1938 (Upper
Triassic Nariwa Group); Oishi, 1940 (ditto); Hsii et al.,619

1979 (Upper Triassic Daqiaode Formation, Sichuan);Feng et al., 1977 (Upper Triassic Anyuan Formation
and Xiaoping Formation, Guangdong): Pinnae expanded at base and decurrent in some Chinese leaves,and pinnae broader in Japanese leaves.
P. bavieri Zeiller: Zeiller, 1902-3 (Upper Triassic of North
Viet Nam); Feng et al., 1977 (I. c.); Zhang, 1978 (Upper
Triassic of Guizhou); Zhou, 1978 (Upper Triassic
Wenbinshan Formation, Fujian); Hsii et al., 1979 (I. c.):Pinnae narrower and elongated.
P. contiguum Schenk: Schenk, 1883 (Lower Jurassic
Xiangxi Formation, Hubei); Hsii et al., 1979 (I. c.):Schenk's leaves were included in P. nathorsti by most
Chinese authors (see below). Other leaves from Sichuan
are larger in size and possess broader pinnae.
P. exhibens Lee: Lee (Li), 1964 (Upper Triassic Xujiahe
Formation, Sichuan); Zhou, 1978 (I. c.): Leaves larger
in size. Pinnae with decurrent base. Veins 5-12 in eachpinna.
P. firmi/olium Ye : In Wu et al., 1980 (Lower Jurassic
Xiangxi Formation, Hubei): Pinnae more elongated anddecurrent.
P. Jurcata Yang: In Chen et al., 1978 (Upper Triassic
Daqiaode Formation): Pinnae shorter and broader. Nine
veins in each pinna.
P. jixiense Chow et Wang: In Zhang et al., 1980 (Lower
Cretaceous Chengzihe Formation, Heilongjiang):
Pinnae longer, with expanded and decurrent base.
P. minutum Li et Tsao: Li et al., 1976 (Upper Triassic
Xiangyun and Yipinglang Groups, Yunnan); He et al.,1979 (Upper Triassic Jieza Group, Qinghai): Pinnae
narrower, basally expanded.
P. nathorsti Schenk: Schenk, 1883 (including Nilssonia
compta) (Lower Jurassic Xiangxi Formation, Hubei);Sze, 1931 (Lower Jurassic of Jiangxi); Sze, 1933a (Upper
Triassic-Lower Jurassic of Sichuan); Sze, 1949 (Lower
Jurassic Xiangxi Formation); HSii, 1954 (= Sze's
specimen); Sze et al., 1963 (ditto); Hsii el al., 1979 (I. c.);Wu el al., 1980 (Tyrmia nalhorsli; Lower Jurassic
Xiangxi Formation, Hubei): Pinnae rectangular for themost part of a leaf.
P. sinensis Lee: Lee (Li), 1964 (I. c.); Li el al., 1976 (I. c.);Feng el al., 1977 (I. c.); Zhou, 1978 (Upper Triassic
Dakeng Formation, Fujian); Chen el al., 1978 (Upper
Triassic Lamaya Formation, East Xizang); Wu el al.,1980 (I. c.): Very close to the present species. Pinnae
not varied but uniform, as judged from the position ofa leaf.
P. Iyrmense (Prynada) Krassilov: Krassilov, 1969 (JurassoCretaceous of the Bureja Basin): Rachis rather thick.
Pinnae elongate-rectangular. Basal pinnae not square
but elongated.
P. yingchenense Chang: In Zhang, 1980 (Lower Cretaceous
Yingchen Formation, Jilin): Leaf large, pinnae twiceas large as ours.
Tyrmia chaoyangensis Chang: In Zhang el al., 1980 (Middle

620

Talsuaki KIM URA and Bong-Ky un KIM

Figures S(c)-9(a-d). All natura l size. Sc . OlOzall1il eS lI1icraurilus Kimura et Kim , sp. nov.: (H o lo type; Reg.
no. 8 1·082) . Loc. Hanaeri. 9. Plerophy llll /l'l lll icraeqllale Kimura et Kim , sp. nov. : 9a. T wo detac hed leaves [H olotype(left); Reg. no. A-5066B]. 9b. (paratype; Reg. no. A-5066c). 9c. (parat ype; Reg. no. A-5066A). Loc. All from Seongjuri.
9d . Somewha t defo rm ed leaves (R eg. no. T-0060). Loc. Cheo nd ae ri.

867.

New Late Triassic plant-taxa from Korea 1

Jurassic of Liaoning): Pinnae set closely, uniform forthe most part of a leaf.
T. latior Ye: in Wu et al., 1980 (Lower Jurassic Xiangxi
Formation): No square or short-rectangular pinnae
recognized.
T. prinadae Vassilevskaja: Vassilevskaja and Abramova,
1966 (Lower Cretaceous of the Lena Basin): Rachis
thick. Pinnae shorter and rectangular.
Pseudocycas polynovii (Novopokrovsky) Krassilov:
Krassilov, 1969 (I. c.): Zhang et al., 1980 (Tyrmia
polynovii ; Lower Cretaceous Sunjiawan Formation,
Liaoning): Pinnae not varied but rather uniform.

As mentioned above, many allied leaves havebeen described from East Asia, but there is no
species unquestionably referable to Coreanophyllum variisegmentum. The difference between
Coreanophyllum and Ctenozamites (once called
Ctenopteris) was already discussed in our previous
paper (Kimura and Kim, 1982).
As no cuticle is preserved, some uncertainty still
exists whether Coreanophyllum is a bennettitaleanor not, but we are inclined to believe that it is mostprobably a bennettitalean because of its pinnae
being morphologically most close to the simply
pinnate leaves of Pterophyllum so far.
Genus Otozamites Braun, 1842

Otozamites micrauritus Kimura et Kim,
sp. nov.
Figures 8a-c

Material: Holotype; Reg. no. 81-082(Hanaeri). Paratype ; Reg. no. 81-153 (Hanaeri).
Stratum typicum: Bongmyeongri Formation.
Locus typicus : Hanaeri (west side). Derivatio
nominis: Small-sized auricle in each pinna.
Diagnosis: Leaves more than 20 cm long and11.5 cm wide at the middle. (Whole leaf unknown.)
Rachis 8 mm wide below and 4.5 mm above; surface longitudinally striated. Pinnae attachedperpendicularly to the upper sides of rachis. Pinnae
long and narrow, typical ones 5.4 cm long and 0.7
cm wide near the base, nearly parallel-sided, thengradually narrowing towards the acutely pointed
apex. Acroscopic basal margin slightly auriculated,auricle triangular and free, and basiscopic basal
margin slightly expanded. Margins entire. Veins
arising from the rachis, numbering 8-9 at the base621

of each pinna; 4-5 acroscopic veins forking at ornear the base, radiating towards the auricle andacroscopic margin, each auricle receiving one or
two sets of veins; 4-5 basiscopic veins forking
twice or thrice at all levels, one persisting to thetip and others ending at the basiscopic margin.
(Cuticle not preserved. Reproductive organs notknown.)
Distribution and occurrence: Only two leaffragments were obtained from the Bongmyeongri
Formation.
Discussion and comparison: Otozamites
micrauritus is characterized by its small-sized
triangular auricle. As shown in Fig. 8a" , each basal
half of acroscopic margin was covered by thebasiscopic part of upper adjacent pinna. Thus,
most of the basiscopic margins of pinnae broke off
when a rock was split for recovering plant fossils.
Accordingly, it is difficult to see the pinna-auricle
in this specimen without removing the matrix. Fig.
8b shows the basal portion of a leaf clearly exposing the auricles.
Otozamites micrauritus resembles o. bornholmensis Moller in pinna-form, small-sized auricle
and venation, but the former is distinguished by
its triangular auricles instead of semi-circular onesin the latter species.
According to our present knowledge, no
Otozamites leaves with such triangular auricles as
our leaves have been found in the literature.
Genus Pterophyllum Brongniart, 1828

Pterophyllum micraequale Kimura et Kim,
sp. nov.
Figures 9a-o

Anomozamites nilssoni (Phillips): Kawasaki, 1925, p. 40,
pI. 21, fig. 66.
A nomozamites minor Kawasaki, 1926 (pars), p. 30, pI. 7,
fig. 22b (non fig. 22a).
Pterophyllum aequale Brongniart: Kawasaki, 1926, p. 27,
pI. 6, figs. 24, 24a', 24a".

Material: Holotype; Reg. no. A-5066B (Seongjuri). Paratypes; Reg. nos. 81-293 (1-4), 203Z
(Hanaeri), A-5066A, C (Seongjuri). Examined
specimens; about 500. Stratum typicum: Amisan
Formation. Locus typicus : Seongjuri.

an d Bong-K y U n KIM
I
Tatsuak l. KMURA

622

3cm

99

9m'

9i'

X2 99'

! J~
9f

9h

867.

New Late Triassic plant-taxa from Korea I

Derivatio nominis: Leaf-form resembling thetypical Pterophy/lum aequale known from Europe
but usually smaller in size.
Diagnosis: Leaf petiolate. Lamina elongate-oval
or elongate-oblanceolate in outline, up to 10 cmlong and 3 cm wide at the widest portion, regularly
segmented. Rachis thick, some attaining 4 mm
wide, with longitudinal ridges on its surface, androunded below. Petiole more than 1.2 cm long.
(Very base unknown.) Pinnae consisting of 25-26
alternate pairs in fully developed leaf, varied inform and size according to the position of a leaf,
attached perpendicularly to the lateral margins ofrachis, but that angle of attachment becoming
oblique towards the apex. Division between pinnae
extending near to or up to the rachis. Pinnae on
the distal two-thirds of a leaf elongate-rectangularin form, typical ones being 1.0-1.4 cm long and3-3.5 mm wide; apex obliquely truncated orasymmetrically rounded. Pinnae on the basal onethird of a leaf short-rectangular, nearly square orsometimes deltoid; distal margin truncated orrounded. Veins parallel, numbering 8-13 at thebase of each pinna, about half of them forking
once near the base, but others simple. Terminal
pair of pinnae abruptly diminishing their size.
(Cuticle not preserved. Reproductive organs notknown.)
Distribution and occurrence: Leaf-fragments
are quite abundant, especially in several horizonsof the Nampo Group and Bansong Group in the
Mungyeong Coal-Field. They are also known fromthe Kimpo Group.
Discussion and comparison: Pterophyllum
micraequale is characterized by its small-sized leaf
with quadri-Iateral pinnae bearing simple or once
forked veins, numbering 8-13 at each base.
Those leaves which have been reported as
Pterophyllum aequale (Brongniart) Nathorst from~623

Europe (e. g. Nathorst, 1878a, b; Antevs, 1919;
Johansson, 1922) and from other regions (e. g.
Zeiller, 1902-3; Stanislavsky, 1971; Iminov, 1976)
resemble those of P. micraequale in the pinnaform, but the former differs especially in the leafsize being twice as large as those of P. micraequale.
The leaf-size of Pterophyllum micraequale isless than 10 cm in length and 3 cm in width.
Pterophyllum (Anomozamites) schenki described by Zeiller (1886, 1902-3) and reported bymany subsequent authors resembles P. micraequalein leaf-form and vein-density, but the former
differs in its large-sized leaf attaining 20-30 cm
in length and 5-12 cm in width.
According to Harris (1937), the leaf-cuticle ofJohansson's Pterophyllum aequale agree with thatof P. schenki from Greenland. Furthermore,
Lundblad (1950) instituted Pterophyllum compressum on the basis of the Swedish material, andincluded in its synonymy most Swedish
Pterophyllum leaves which had been regarded as
P. aequale and P. affine as well as Greenland
leaves which were reported as P. schenki. At any
rate, Pterophyllum micraequale is distinguishedfrom these European species by its small-sizedleaves.
Leaf-fragments of Pterophyllum micraequale
are locally quite abundant, often thickly massed
and appressed to form singly a distinct fossil
population. However, its leaves are sometimes
associated with those of Coreanophyllum
variisegmentum in some sparsely fossiliferous
horizons.
Pterophy/lum micraequale is distinguished from
Coreanophyllum variisegmentum by the following
features: 1) The lamina of a full-grown leaf of
Pterophyllum micraequa/e is divided into 25-26
alternate pairs of segments (or pinnae), whereasthat number in Coreanophyllum variisegmentum

Figures 9(e-o). Pterophyllum micraequa/e Kimura et Kim, sp. nov.: Natural size unless otherwise indicated.
ge. Occurrence of this species (Reg. no. 81-033). Loc. Hanaeri. 9f. Drawn from ge. 9g and 91. Drawn from 9a (91;
Holotype). 9g'. Venation drawn partly from 9g. 9h. Drawn from 9b. 9i. A typical leaf (paratype, Reg. no. 81-293-1).
Loc. Hanaeri. 9i'. Venation drawn partly from 9i. 9j. Leaf apex (Reg. no. A-SOOI). Loc. Seongjuri. 9k. A smallsized leaf with preserved petiole (Reg. no. T-0020). Loc. Cheondaeri. 9m. Basal part of a broken leaf (paratype;Reg. no. 81-293-4). Loc. Hanaeri. 9m'. Venation drawn partly from 9m. 9n. Basal part of a leaf with a thick rachis(paratype; Reg. no. 81-203Z). Loc. Hanaeri. 9n'. Venation drawn partly from 9n. 90. Basal part of a leaf (paratype;Reg. no. 81-293-2). Loc. Hanaeri.

624

Tatsuaki KIMURA and Bong-Kyun KIM

comes to 54-55. 2) The rachis of P. micraequale
is ornamented by several longitudinal striations on
its basal upper surface and is usually thicker thanthe pinna axis of C. variisegmentum with a medianfurrow above. 3) The pinnae of P. micraequale arebroader and typically quadri-lateral in form witha truncated apex, while those of C. variisegmentum
are narrower and typically possess a rounded or
obtusely pointed apex, although they are variablein form depending upon the position of
penultimate pinna. 4) Veins at the base of eachpinna are denser in P. micraequale (8-l3) thanin C. varriisegmentum (3-6). 5) The penultimate
pinna fragments of C. variisegmentum are also
locally quite abundant, often thickly massed andappressed to form a monospecific fossil population like that of P. micraequale. It would be worthy to note that in monospecific population of C.
variisegmentum, no P. micraequale leaf has so far
been recognized and that the same is true for the
population consisting singly of P. micraequale.
Therefore, we consider both species to be distinct
from each other and each species is thought to havemaintained its own exclusive community. It is,however, difficult to distinguish these two species
solely on the basis of small leaf- or pinnafragments in which veins are invisible.
In Japan and China, many bennettitalean leaffragments have been described under the name of
Pterophyllum aequale. However, we now consider

most of them to be not referable to P. aequale,
because they are far smaller in size than the leaves
identified as P. aequale in Europe and Middle
Asia. Among them, fossil plants in the following
records are similar in size, form and venation to
Pterophyllum micraequale:
Yabe and Oishi, 1929, p. 93, pI. 18, fig. 4, pI. 20, fig. 3:
Upper Triassic Anyuan Formation, Jiangxi, China.
Sze, 1931, p. II, pI. 2, fig. 5; Hsii in Sze and HSii, 1954,
p. 57, pI. 50, fig. 5: The same as the above.
Sze, 1933a, p. 20, pI. 4, figs. 2-7: Upper Triassic Xujiahe
Formation, Sichuan, China.
Oishi and Huzioka, 1938, p. 85, pI. 10, fig. 4, 4a; Oishi,
1940, p. 339 : Upper Triassic Nariwa Group: Somewhat
larger in leaf-size.
Sze et al., 1963, p. 152, pI. 60, figs. 5-6, pI. 69, figs. 7-8,
8a, pI. 71, fig. 9: Upper Triassic and Lower Jurassicof Sichuan and Guangdong: Leaf base is markedly
swollen.
Li et al., 1976, p. 118, pI. 33, figs. 8-10 : Upper Triassicof Yunnan: Veins in each pinna denser, 16-20 in
number.
Zhou, 1978, p. 108, pI. 21, fig. 5: Upper Triassic Wenbinshan Formation, Fujian (non his pI. 21, fig. 6).

Pterophyllum micraequale resembles P.
nathorsti described by Schenk (1883) from China,
but the former is distinguished by its typicallyquadri-lateral pinnae with a truncated apex.
Pterophyllum nathorsti has pinnae formed ofnarrower segments with a rounded apex.
(To be continued)

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 152, pp. 625-643, 16 Figs., December 30, 1988

868. GROWTH OF ANTLER IN THE SUBGENUS SIKA
(CERVID, MAMMAL) FROM THE PLEISTOCENE FORMATION
IN THE SETO INLAND SEA, WEST JAPAN*
HIROYUKIOTSUKA
Institute of Earth Sciences, Faculty of Science,
Kagoshima University, Kagoshima 890

Abstract. This paper presents the results of a paleontological study of fossil antlers of the
Pleistocene Sika-deer collected together with numerous specimens of Naumann's elephant(Palaeoloxodon naumannt) and many kinds of herbivorous mammals from the Bisan-seto area, of
the Seto Inland Sea, West Japan. To determine the tempo and mode of forking of fossil antlers,corresponding aspects of living Japanese deer were examined in detail, based on individuals whoseages were determined by the cementum layers of the teeth. This study con firmed that several formsof antler which had been identified as belonging to four fossil species (i.e. katokiyomasai, natsumei,
paleoezoensis and cf. greyi of the subgenus Sika) in previous studies, merely represent several different
growth stages of a single subspecies of the subgenus Sika. The subspecies Cervus (Sika) greyi
katokiyomasai Shikama and Hasegawa is selected as a valid name for the Pleistocene Sika-deerfrom the Seto Inland Sea examined here, according to the rules of zoological nomenclature. The
forking in C. greyi katokiyol71asai is, however, slighter and more slender than that of C. greyi
throughout the course of life. C. greyi katokiyomasai flourished in the lowland area of West Japanas a characteristic island form in association with other endemic deer species descended from earlier
migrants such as Nipponicervus and Elaphurus. Cervus nippon yesoensis of the Jomon Period of
Early Holocene age might have derived from C. greyi katokiyomasai in Late Pleistocene time andbecome distributed throughout the Japanese Islands.

Introduction

buffalos (Bison occidentalis), and several kinds ofCervidae (including Sinomegaceroides, ElaphUlllS,
Nipponicervus and Sika). Among them, Stegodon
orientalis was inferred to have derived from Early
Pleistocene deposits, while the Naumann's
elephant and other associated mammals arethought to have washed out from Middle to Late
Pleistocene formations, called the Bisanseto Group(Bando et a{., 1978). The stratigraphic position of
the mammalian fossils in this area will be discussed
in the next section.
Hasegawa (1972) regarded Naumann's elephantand its associated fauna to be a part of the"Pa{aeo{oxodon - Sinomegaceroides Complex"
which had a wide distribution in Middle to Late
Pleistocene formations over the Japanese Islands(Hasegawa, 1972). Among the mammalian

It is known that abundant mammalian fossilsof Pleistocene age occur on the floor of the Seto
Inland Sea, West Japan. These fossils are regarded
as derived fossils which were washed out from
Pleistocene formations flooring the sea bottom(Hasegawa, 1972; Otsuka and Shikama, 1977;
Bando et al., 1978). These mammalian fossils havebeen studied from a paleontological point of viewby many authors (Makiyama, 1938; Matsumoto,
1924; Naora, 1970; Shikama, 1941; Hasegawa,
1972; Otsuka, 1977; Otsuka and Shikama, 1977).
The fauna is mainly composed of elephants (suchas the genus Stegodon and Palaeoloxodon),*Received May 17, 1988; revised manuscript accepted
November 9, 1988.
625

626

Hiroyuki OTSUKA

figure 1. Localities of examined, living JapaneseSika-deer (A) and fossil Sika-deer in the Seto Inland Sea(B) in the Japanese Islands.

assemblages from the Seto Inland Sea, the deer
assemblage is predominated by Nipponicervus andSika, and is accompanied by a few species ofSinomegaceroides and Elaphurus. For this reason,
the deer assemblage from this area may be called
the Sika-Nipponicervus assemblage (Otsuka andShikama, 1977). This assemblage is distributed
over nearly the entire area of the Seto Inland Sea
from the bay-mouth area of Osaka Bay to thevicinity of Hiroshima City.
Cervus kazusensis of early Early Pleistocene age
is the first species of the subgenus Nipponicervus
to appear in north<!astern Asia and subsequently
spread from central to western Japan during
Middle and Late Pleistocene times (Otsuka andShikama, 1977). Continuous change in the antler
morphology can be observed from the Early
Pleistocene species, Cervus (Nipponicervus)
kazusensis to the Middle to Late Pleistocene
species, Cervus (Nipponicervus) praenipponicus.
These two species of Nipponicervus occur in thesame frequency from the Seto Inland Sea.

Therefore, the Sika-Nipponicervus assemblage in
the Seto Inland Sea seems to represent a transition
period from C. kazusensis to C. praenipponicus.
To date, the following species belonging to thesubgenus Sika have been recorded from the Seto
Inland Sea.
1. Cervus (Sika) yesoensis Heude (Shikama,
1941)
2. Cervus (Sika) nippon nippon Temminck(Shikama, 1941)
3. Cervus (Sika) cf. hortulorum Swinhoei(Naora, 1970)
4. Cervus (Sika) katokiyomasai (Shikama andHasegawa) (Shikama and Hasegawa, 1965;
Otsuka, 1977)
5. Cervus (Sika) natsumei Matsumoto (Otsuka
and Shikama, 1977)
6. Cervus (Sika) cf greyi (Zdansky) (Otsuka
and Shikama, 1977; Otsuka, 1977)
7. Cervus (Sika) paleoezoensis Otsuka andShikama (Otsuka and Shikama, 1977; Otsuka,
1977)
Among these species, the first three species wereidentified with the living species and their names
were merely listed without any paleontologicaldescription, whereas the other species were
proposed as new to science. The last three were
described on the basis of abundant antler
specimens in the Takao Collection, recovered fromthe sea bottom off Shodoshima (Otsuka andShikama, 1977). However, a systematic relationship among these species remains unresolved.
Recently, I have had an opportunity to study
many well-preserved fossil antler specimens ofcervids in the "Yamamoto" collection, gatheredfrom the Bisan-seto area about 38 km west of
Shodoshima by Mr. Keiichi Yamamoto, a juniorhigh school teacher. The collection includes
abundant specimens of elephants from the same
area (Otsuka, 1987).
Based on a statistical and comparative
morphological analysis of the fossil antlers of Sika
deer in the Yamamoto and Takao collections, I
reached the conclusion that those fossil antlers,thought to represent the four species, C. natsumei,
C. cfr. greyi, C. paleoezoensis and C.
katokiyomasai, represent different growth stages

868.

Growth of antler in Sikaof a single species. To demonstrate this, the antler
growth of living Japanese Sika-deers such as;
Cervus (Sika) nippon yesoensis, Cervus (Sika)
nippon centralis and Cervus (Sika) pulchellus; hasbeen examined in detail. The ages of the animals
were determined by the cementum layers of theirteeth by using Mitchell's method (1963). These
growth data clearly indicate that those several
forms of antlers previously identified with four
fossil species (katokiyomasai, natsumei, paleoezoensis and cfr. greYl) are merely different growthstages of antler of a single subspecies. A valid nameto represent the fossil Sika-deers examined here is
the subspecies name "greyi katokiyomasai". The
antler of this subspecies in the mature stage ischaracterized by a very wide angle in the firstforking with a much lyrated beam. This mode of
forking resembles that of Cervus (Sika) greyi
Zdansky in the Middle Pleistocene Choukoutien
Fauna of North China, but the Japanese subspecies
has a somewhat smaller and more slender antler.
C. (Sika) greyi katokiyomasai is probably derived
from C. (S.) greyi during the early Middle
Pleistocene when the latter species migrated to theJapanese Islands from the Asian Continent. The
subspecies flourished in the lowland area inassociation with such mammals of the forest
habitat as Naumann's elephant and other endemic
deers including Elaphurus and Nipponicervus.
Descriptive terms of antler
In this paper, descriptive terms andmeasurements of antler of the genus Sika follow
those proposed by Otsuka and Shikama (1976)(Fig. 2).
WL:
PL:
PD:
BD:
LTl:
LT2:
LT3:
LBI:

Whole length of antler
Minimum length of pedicle measured along theposterior border
Minimum width of pedicle
Side-to-side diameter of burr
Length of the first tine measured along its innersurface from the base of the fork to its tip.
Length of the second tine measured along its innersurface from the second fork to its tip
Length of the third tine, measured as in LTI and
LT2
Length of the beam between the bases of the first
and the second forks

Figure 2.

627

Terminology for antler of the subgenus

Sika.

LB2: Length of the beam between the bases of the second
to the third forksa I: Angle of the first fork
a 2: Angle of the second fork
a 3: Angle of the third fork
H:
Height of the first fork measured from the base of
the burr to the uppermost surface of the "webb"
DFI: Distance between inner surfaces of the base of thefirst forking of opposed antler
DF2: Ditto between the second forking
DF3: Ditto between the third forking

Growth of antler in the living Japanese Sika
1. Materials
The specimens used to study the growth ofantler of living Japanese Sika-deer are listed below.
The specimens of antler, with skull attached, wereborrowed from a number of universities andmuseums in Japan.
By using Mitchell's method (1967); Noriyuki
Ohtaishi, Hiroko Koike, and Mitsuharu Matsumoto; estimated the ages of these specimens fromthe cementum layers of the teeth.
2. Results
To determine the tempo and mode of forkingof antlers in life, the diameter of the burr (BD),the diameter and length of the pedicle (PD, PL),the whole length of the antler (WL), the length of
the beam between the first and the second forks(LBl) and the rate of lateral expansion of antler(DFl, DF2, DF3) were investigated.

Hiroyuki OTSUKA

628
Laboratory

Dept. of Anatomy,
Hokkaido Univ.

Specific Name

Number of
specimens

Locality

Nakanoshima in
Lake Toya, Hokkaido

C. nippon yesoensis

15

C. nippon centralis

11

Nara park

C. pulche/lus

11

Tsushima Is.

Tochigi
Prefectural Museum

C. nippon centalis

13

Nikko area

Lab. of Verterinary
Medicine, Kagoshima Univ.

C. pulche/lus

22

Tsushima Is.

1) Pedicle Index. Usually, deer antler is shedannually. When shedding occurs, a thin layer fromthe topmost part of the pedicle adheres to the antler
and is removed, causing the pedicle to become
shorted with every shedding. As the same time thepedicle diameter broadens continuously withgrowth. Therefore, it can be stated that a male witha long narrow pedicle is younger than one bearing
a short broad pedicle. For those specimens treatedin the present study, the pedicle widths of C.
nippon yesoensis, C. nippon centralis and C.
pulchellus broaden continuously with age, increasing from 2.5 to 3.0 mm per year in width,
while shortening at a rate ranging from 1.0 to 1.2
mm per year in C. nippon centralis and C. nippon
pulchellus, and 1.0 to 3.0 mm per year in C.
nippon yesoensis. Consequently, it can be assumed
that deers shedding larger antlers such as C. nippon
yesoensis tend to break larger pieces from the topof the pedicle than those shedding rather smaller
antlers, such as C. nippon centralis in Nara.
To determine an approximate age of male deer,a "Pedicle Index" is proposed. It is defined as [theratio between the diameter of pedicle (PO) and thelength of pedicle (PL)] x 100. As shown in Fig.
3, values of this index increase with the age ofanimal. An index value of 100 represents 2.5 yearsof age in C. nippon yesoensis, and about 3.5 yearsof age in C. nippon centralis in Nikko and C.
pulchellus. An Index value of 200 represents about
9 years of age in C. nippon yesoensis, 10 to 13 yearsof age in C. pulchellus and 14 years of age in C.

nippon centralis. Thus, the pedicle index seems to
be a useful method to estimate the age of deer ina single population using antlers. In this study, thismethod was applied to estimate the age of fossil
deer from the Seto Inland Sea which will be
discussed in the next section.
2) Diameter of Burr (Figs. 4, 5). In C. nippon
yesoensis, the burr increases in width from a
yearling to adult of eight years in age at a rate ofabout 4.8 mm per year, then after eight years old
it increases at only about 3.0 mm per year, andfinally settles at the rate of a seven year old
animals.
In the case of C. nippon centralis from Nikko
and C. pulchellus, for yearlings to eight year old
animals the burr diameter increases at an averagerate of 3.3 mm per year. Although antlers of fiveto ten year old adult C. nippon centralis from Nara
were not examined, they seem to show the sametendency as those of C. pulchellus (Fig. 5).
3) Length of Antler (Figs. 4, 5). The antlersof both C. nippon yesoensis and C. pulchellus havea maximum length (WL) at the age of seven; forinstance, the three-forked antler is 720 mm longin C. nippon yesoensis and 520 mm long in C.
pulchellus.
The length of the beam between the first andthe second forks (LBl) in C. nippon yesoensis has
a maximum value at the age of seven and it isabout 2.5 times its length at two years. For
juveniles, from four to five, this length varies considerably, as much as 130 mm, which is the largest

868.

Growth of antler in Sika

629

250

C. n. centra/is~/ Nikko----r----~~- _~~n=2--+-_+

0 200

0
.,...

C. pu/che/ /us.

..J

n=2

a.

1

• ~ ./ /1

><

j

=4.1

"-

;·r

Cl

a.

n=/

----~~--1----/f----~-+>< 150

(I)

1J

c::

(I)

.... ....-()/-/ f~:/ I

C.n. centra/is
Nara

1J
(I)

a.

100

50
9

10

11

12

13

14

15

16

Age (Years)
Figure 3.
Sika-deer.

Relationships between the Pedicle index (PD) and the age (years) of living Japanesevariation found during the life cycle. In C.
pulchellus, this length shows a considerable variation before the age of seven (the prime of life) andthe maximum value is reached in animals fromseven to ten years old (Fig. 6). In C. nippon
centralis from Nikko, this length increases in value
until eight years of age, but animals older than ten
years have almost the same value as for those ofeight years of age.
4) Height of the First Forking. In C. nippon

yesoensis and C. pulchellus, the height of the first
fork increases at a rate of 8 mm per year until theage of six years. The height is almost constant fromsix to eight years and after nine years it gradually
decreases, eventually reaching the value of a fouror five year old. Growth of the height of the first
fork of C. nippon centralis from the Nikko andNara areas shows a similar tendency as those ofnippon yesoensis and pulchellus.
5) Angle of the First Forking (Figs. 6, 7).

Hiroyuki OTSUKA

630

In C. pulchellus, the angle of the first forking inspecimens from two to six years of age ranges from55 to 85 degrees; from six to nine years of age, therange is from 65 to 80 degrees; and after ten yearsof age, it is less than 65 degrees.
The same angle for males of C. nippon yesoensis younger than eight years shows a small variation ranging from 65 to 75 degrees; for those older
than eight it tends to decrease. Finally, deer older
than ten years of age tend to have narrow fork
angles nearly equal to young males of about five.
The growth pattern of C. nippon centralis showsa similar tendency as that of C. pulchellus.
6) Lateral Expansion of Antlers. To understandthe growth related changes in the mode of lateralexpansion of antlers, the distance between the
forking points of opposite sides (DFI, DF2) was

examined. In C. nippon yesoensis, the distance
between the first forking points (DFl) increases
gradually at a rate of about 3 mm per year untileight years; thereafter it increases more slowly,
equal to that of a four or five year old. In theyoung of C. pulchellus, this distance increases
slowly. After nine years of age, the distance tends
to decrease.
In C. nippon yesoensis, the distance between thesecond forking points (DFl) at the juvenile stage,
from yearling to three years old, increases at a rateof 130 mm per year, and thereafter at a rate
ranging from 50 to 60 mm per year until eight yearsof age. After eight years, however, the increase is
only about 42 mm per year, a rate nearly the samea~ that of four or five years old.
In C. pulchellus, the distance between the550.......----r--.....,...--.....,...---.----,(mm)

7001--+---+---+---+-/(mm)

500f-+---+--+-

450

600

400

':J

::J

~

2500

....

Q) 350

Q.l

.....

.....

c

c

« 40

.....

« 300

a

.....
a

.c

.c

.....

'OJ 250

OJ

~ 300

~

CD

.:::!
E
Ctl

C
Q.l

-l

'+-

0

...

.J::

-l

Q)

200

CD
'+-

OJ

0

C

...

Q)

.J::

...J

OJ

C

Q)

...J

30

40

50

60

~mm)

Side-to-side Diameter of Burr(BDJ
Figure 4. Relationships between side-to-side
diameters of the burr (DB) and whole length of antler (WL)
(upper), and between side-to-side diameter of the burr andlength of the beam (LBI) (lower). of living Cervus (Sika)
nippon yesoensis Heude. collected from the Nakanoshima
in Lake Toya. Hokkaido.

35

40

45

50 (mm)Side-to-side Diameter of Burr(BD)Figure 5. Relationships between side-to-side
diameter of burr (DB) and length of the antler (WI) (upper).
and between side-to-side diameter of burr and length of
the beam (LBI) (lower) of Cervus (Sika) pulchellus
Imaizumi from Tsushima Island.

868.

Growth of antler in Sikasecond forking points (DF2) increases at a rate of
120 mm per year until the age of three years. For
the young males of from four to eight years old,
this distance increases irregularly. After eight yearsof age, however, the rate tends to decrease.

631

The rate of expansion of the antler of C. nippon
centralis from Nikko resembles that of C. nippon
yesoensis but it is smaller. That of C. pulchellus
is similar to C. nippon centralis with growth.
From these data, it is concluded that the antler

400r-----r-----.-----,-----.-.

(mm

250r--r-----r-----.-----.-----,(mm)

C.nippon yesoensis

C. pulche/lus... 10yl

350+-----4-----~----~-1

<t>9Y
~

CO 3010+------+-----1---,20'D-+---+-----~...0.1E:uQ)

CO 250't------+------t-~

o

..c

.....

o

0) 200+------1-__::.----

C

150+--+------'-.

o

Q)

--.J

2Y

o

150

100L-____~----~--~~--__:~~
50°
60°
7 °
80 0

2Y

100·~_4~----~--~±n----~,---~

500

Angle of Fi rst Fork

60 0

7 °

80 0

Angle of Fi rst For k

Figure 6. Relationship between the angle of the first fork (a 1) and the length of
the beam (LBI) in living Japanese Sika-deer. Cervus nippon yesoensis Heude Oeft) and Cervus
pulchellus Imaizumi (right).

C. pulchellus

C. nippon yesoensis~8if~----+-----+_----~----_t----_+----_i----~~_lo

1 4Y//~~~v

u.

,,7Y

+"'

~70)~----+------I------+-~~.

+sy

V ~+-----l~o

VJ"'10Y...-2Y~)o 6 ~--I------t-----+-----+----~-----t----t

1lY

\

Q)

01

~5~~----~----~----r-----r-----r-----r-----r--l
30

3

40

4

50

5

6

Side-to-side Diameter of Burr(BD)7(mm) 30

/

35

40

45

50 (mm)Side-to-side Diameter of Burr

Figure 7. Relationship between the angle of the first fork (a I) and side-to-side diameter of the burr (DB)of living Japanese Sika-deer. Cervus nippon yesoensis Heude (left) and Cervus pulchellus lmaizumi (right).

Hiroyuki OTSUKA

632

of Sika-deer shows a gradual increase in its total
length, angle of the first forking, and antler diversion until about eight years of age is reached.
Thereafter, all these values tend to decrease
eventually equalling those of four or five years old.
This may imply a low metabolic rate of older
Sika-deer.
Morphology and systematic position of thefossil Sika-deer from the Seto Inland Sea
Order Artiodactyla
Family Cervidae Gray, 1821
Genus Cervus Linnaeus, 1785
Subgenus Sika Sclatter, 1870
Type-species.- Cervus nippon Temminck, 1873

Cervus (Sika) greyi Zdansky subsp.
katokiyomasai Shikama and Hasegawa, 1965
Figures 14-16
cfr. Pseudaxis greyi Zdansky, Zdansky, 1925, Pal. Sinica,ser. C, vol. 2, fase. 3, p. 65-72, pI. 13, figs. 2-12.
Cervus (Pseudaxis) greyi Zdansky, Teilhard de Chard in andPei, 1941, Ibid., N.S. ser. C, no. 11, p. 76-80, pl. 2,
fig. 3.
Pseudaxis greyi var., Young, 1923, Ibid., ser. C, vol. 8,. fase. 2, p. 21-37.
Cervus ef. greyi Zdansky var. etva Teilhard de Chardin,
Teilhard de Chardin, 1936, Ibid., ser. C, vol. 7, fase.
4, p. 39-42.
Cervus (Sika) ef. greyi (Zdansky), Otsuka and Shikama,
1976, BUll. Nat. Sci. Mus., ser. C, Ceol., Paleont., vol.
12, no. 3, p. 1-40; Otsuka, 1977, Rep. Fac. Sci.
Kagoshima Univ., (Earth Sci., Bioi.), no. 10, p. 47-48.
Cervus (Rucervus?) katokiyomasai Shikama and Hasegawa,
1965, Sci. Rep. Yokohama Nat. Univ., ser. 2, no. 12,
p. 45-47; Otsuka, 1977, Rep. Fae. Sci. Kagoshima
Univ., no. 10, p. 48-51.
Cervus (Sika) natsumei Matsumoto, Otsuka and Shikama,
1977, BUll. Natn. Sci. Mus., ser. C (Ceol., Paleont.) ,vol. 3, no. 1, p. 17-19, pI. 1, figs. 4-7.
Cervus (Sika) paleoezoensis Otsuka and Shikama, 1977,
Ibid., vol. 3, no. 1, p. 19-24, pI. 2, figs. 2, pI. 2, figs.
1-12.

Type specimen: Right, shed immature antler
stored in the Department of Geology, Kumamoto
University. A plaster cast of the type specimen is
kept in the National Science Museum, Tokyo(NSM Reg. No. 6586) and in Kagoshima Univer-sity (ESK Reg. No. F-6052).
Referred specimens: 107 antler specimens
including 80 shed antlers are treated; 65 of thespecimens are right antlers and 42 specimens areleft antlers. These specimens were collected fromthe sea bottom in several areas of the Seto InlandSea, and are kept in the National Science Museumof Tokyo (Takao Collection) and the Natural
Science Museum of Kurashiki City (Yamamoto
Collection).
Collection localities: The Bisan-seto area of the
Seto Inland Sea and off Shodosima Island, in theeastern part of the Seto Inland Sea.
Horizon: The exact horizon is uncertain, butmany field data suggest that the deer fossils were
washed out from the Late Pleistocene Tsuchinoseto Formation (Mindel/Riss interglacia - Rissglacial age).
Subspecijic diagnosis: Sika-deer bearing a
medium to large antler, showing a greatmorphological variation in life. The antler in theadult stage is characterized by a much lyrated beamwith a long first tine and general form somewhat
resembling that of the subgenus Rucervus, but theantler of the young stage has almost the same shapeas that of many other species of the subgenus Sika.
Remarks: Among the 195 antler specimens ofSika-deer from the Bisan-seto sea and area off
Shodoshima Island, 31 specimens (about 16070) are
non-shed antler bearing a pedicle. Specimens
retaining the portion between the burr and second
forking point make up less than 12 percent of thetotal and most of the remaining specimens havelost the upper part of the antler. Although these
antler specimens are imperfectly preserved,
statistical analyses of the antler population strongly
suggest that they belong to a single species thatshows a large morphological variation in its growth
pattern.
Furthermore, there is a strong possibility thatseveral forms of antler previously reported fromthe Seto Inland Sea under such different specificnames as katokiyomasai (Shikama and Hasegawa,
1965; Otsuka, 1977), natsumei, cf. greyi andpaleoezoensis (Otsuka and Shikama, 1977; Otsuka,1977), represent different growth stages of a single
species. To prove this hypothesis, the following

868.

Growth oj antler in Sika

points regarding various characteristic of antlers
were examined.
1. Age of the fossil antler of Sika form the Seto
Inland Sea (estimation based on Pedicle Index).
1) Non-shed antler
For the 29 non-shed antler specimens, themaximum length (PL) and minimum width (PD)of pedicle were measured and each of the pedicle
indices was calculated. The indices for each antler
specimen were compared with those of living
Japanese deer (c. nippon yesoensis, C. nippon
centralis, C. pulchellus) and the ages of the 29 fossil
antler specimens were roughly estimated.
As already mentioned in the preceeding section,
the rate of shortening of the pedicle with age is fiveto eight percent larger in C. nippon yesoensis thanin C. nippon centralis or C. pulchellus. Therefore,
if the fossil antlers with pedicle attached arecompared with those of C. nippon yesoensis, theirages are likely to be estimated as being two or three
years younger than those for C. nippon centralis
or C. pulchellus..
The fossil antlers of Sika-deer from the Seto
Inland Sea are characterized as rather long andslender and their size nearly corresponds to thatof C. nippon centralis or pulchellus. The
shortening ratios of the pedicle of fossil aged deers
to those of juveniles, for examples, are 0.54, 0.58
and 0.61. These values correspond to those for a
12 to 13 year old C. nippon centralis and for a 10
to 11 year old C. nippon yesoensis. Accordingly,
if the fossil antlers are compared with those of C.
nippon yesoensis, age estimates are younger thanthose of C. nippon centralis or C. pulchellus by
two or three years.
2) Shed antler
As already mentioned, the burr diameter of theantler in a single population seems to be a usefulclue to the growth stage of antlers. To ascertain
the age of a shed antler, the angle of the first fork
and diameter of burr were plotted to show theirrelationship and are then compared with the nonshed antlers whose ages were estimated by thepedicle index (Fig. 10).
The general tendency shown in this figure
resembles the curve of the morphological growthchange of the antler of the living Japanese Sika-

633

deer shown in Fig. 7. In the case of C. nippon
centralis and C. pulchellus, the curves rise convexly
upward, showing the largest angle at six years ofage.
As stated earlier, antlers of Sika-deer from the
Seto Inland Sea are herein regarded as those of asingle subspecies Cervus (Sika) greyi katokiyomasai(Shikama and Hasegawa), representing growthstages of from two to eleven years based on theage data of living Sika-deer. Furthermore, thisantler asemblage is roughly classified into six ageclasses ranging from juvenile to adult males.
2. Changes in the beam length
Fig. 9 shows the relationship between the beam
length (LBI) and the diameter of burr, and Fig.
10 shows the relationship between beam length andthe angle of the first fork of C. (S.) greyi
katokiyomasai.
The rate of growth of the beam (LBI) of C. (S.)
greyi katokiyomasai resembles more closely thatof C. nippon yesoensis than C. nippon centralis.
Namely, that of C. greyi katokiyomasai shows arather wide variation of up to 200 mm for juvenile
to senescence. This value is almost equivalent tothat of C. nippon yesoensis, but is larger than thatof both C. nippon centralis and C. pulchellus. In
C. greyi katokiyomasai, the beam lengths between
the first and the second forks (LBI) from the agesof seven to nine, age estimated based on living C.
nippon centralis and C. pulchellus (or from fourto six years based on C. nippon yesoensis) arealmost 1.5 times longer than juveniles under five
(three) years old. Hereafter all age estimates are
given twice, the first time is based on C. nippon
centralis and C. pulchellus and the second, inparentheses, is based on C. nippon yesoensis.
Adult males, older than ten (seven), have stunted
antler corresponding in size to those of five (three)year old individuals. Thus, the beam length of C.
greyi katokiyomasai in life resembles that of C.
nipponyesoensis, however, their burr diameter ateight years of age does not exceed 50 mm andequals that of the young stage of C. nippon
yesoensis under four years of age.
3. On the re-examination of previously recorded
species on the subgenus Sika from the Seto Inland
SeaHiroyuki OTSUKA

634

400,-------,-------r-----~------~~----_r------,

(mm)

\,

~

~ 9l(6Y.)) , I,

A . "~

7Y.(4Y.)/~~ I~ ~6~ Y

35O+-----+------+r

y

~/

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§

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'+- 250+----

I

o

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Q)

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150

0
14479
/

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3y.

30

:566
810

I

14443-2

1~38

35

40

45

50 (mm)Side-to-side Diameter of Burr(BD)Figure 8. Relationship between the length of the beam (LBI) and side-to-side diameterof burr (DB) of Cervus greyi kalokiyoll1asai Shikama and Hasegawa from the Seto Inland Sea.
(mm.

__ 350r-+---~----~<-en.....J

E300r-r---+---1--COQ)

en,+-250T--r--~-­oFigure 9.
Relationship between the length of beam (LBI) and the angle of the first
fork (a 1) of Cervus greyi kalokiyol11asai Shikama and Hasegawa from the Seto Inland Sea.

868.

Growth of antler in Sika

635

1200
1100
1000
~
I-

0
LL
+oJ

90
800

(j)
I-

0,0: Takao collection.6.,0: Doi collection
40 ~°r-~------+------7---.: Yamamoto collection.... :holotype of C. katokiyomasai

0'44773010~--4-----~~--_/~~----~~----~------+-----~~~~
20
25
30

S ide-to-side
Figure 10. Relationship between the angle of the first fork (a I) and side-to-side
diameters of burr (DB) of Cervus greyi katokiyomasai Shikama and Hasegawa from the
Seto Inland Sea.

The dimensions of the antlers which had
previously been identified as the species
katokiyomasai, cf. greyi, natsumei, andpaleoezoensis are plotted in Figs. 8 to 10. As far
as these data are concerned, the type and referred
specimens of these species probably represent
several growth stages of a single species.
Cervus (Sika) natsumei Matsumoto (Otsuka
and Shikama, 1977) characterized by the tripleforked, small antler (NSM Reg. No. 14444-1, NSM
Reg. No. 14477) is interpreted to come from ayoung deer comparable to that of the living
Japanese Sika-deer undr three years of age.
The antler specimens described under the nameof Cervus (Sika) paleoezoensis Otsuka andShikama (Otsuka and Shikama, 1977) include

antlers of various growth stages older than three(two) years of age. The holotype (NSM Reg. No.
14476) characterized by a long first tine and wide
angle of the first forking represents antlers of midlife individuals, ranging from five (three) to seven(four) years of age, while the paratype (Reg. No.
NSMI4443-2) characterized by a short beam andshort but stout pedicle (pedicle index = 168) appears
to correspond to that of an aged male of C. nippon
centralis older than ten (seven) years.
The type specimen of C. (? Rucervus)
katokiyomasai (Shikama and Hasegawa, 1965) isrepresented by a right, shed, incomplete antler ofa young male lacking the upper half of the beam.
Based on the diameter of burr and rather wide
angles of the first fork, it is estimated to be middle

Hiroyuki OTSUKA

636,

under 3 years

3 -5 years

'I

"' I
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I,

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9 -10years

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Figure 11.
Growth series for antler of CervlIs (Sika) greyi katokiyomasai Shikama and Hasegawa. Approximate ages of each antlers were estimated from data on living CervlIs nippon centra/is Kishida from the Nikko andNara areas.

868.

Growth of antler in Sika

637

c. greyi katokiyomasai
120 C. greyi~...::::-+---+-

mnLD

1101--+---+---1-100
.Ott.

.....
0

90

U.

..(/)
.....

ao

u.

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(/)

Ql

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40'

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C. nippon yesoensis

nippon yesoensis

L4.ii:lJ=:=:= c. pulchel/us II1t.rr(Tor Ihama shell mound)

35

40

45

50

55

60

65 (mmSide-to-Side Diameter of Burr(BD)Figure 12. Relationship between the length of the beam (L B 1) and side-to-side diameter ofburr (BD) of living and fossil Sika-deer.
500r----------------r--.---.----,----r--,~(mm) C.greyi katokiyomasaidXl
C. greyi 'Ci"L:-=::':IIIlI

o450 C. nippon centra/is --0--0-0--0-

u.

"'0

-r....-.-r-4..i....~-L1

Cu

C.nippon yesoensis<ll

C. nippon yesoensis _ _

o 400

(f)

(Nakanoshima)__ _(Torihama shell mound)] 350,I--C_.-'-p_u_/--!c_h_e_I_lu_s.......,.~~""!"'~=..:!l:T't--"u.
c 3001-----r---t----+
<ll
<ll

....~

~ 2501----+----+---

E

ro

<ll

m 200~----+---~_~
'+--

o
..c
....

OJ

C 150

<ll

-l

35

40

45

50

55

60

65(mm)

Side-ta-Side Diameter af Burr (80)Figure 13. Relationship between the angle of the first fork (a 1) and side-to-side diameterof burr (BD) of the living- and fossil Sika-deer.

638

HiroYliki OTSUKA

868.

Growth of antler in Sika

aged between seven (four) and nine (five) years old.
Another, well-preserved antler of C. katokiyomasai was recovered from the sea bottom off
Tomogashima and this specimen was included in
the subgenus Sika (Otsuka, 1977). Judging fromthe pedicle index (Pd = 152), wide angle of the firstforking, and the diameter of the burr, this antler
specimen roughly corresponds to that of nine (five)
year old C. nippon centralis.
Cervus (Sika) cf. greyi, described from the sea
bottom off Shodoshima based on 33 incomplete
antler specimens (Otsuka and Shikama, 1977), wasconsidered to be one of the Sika-deers
characterized by "triple-forked, small- or
moderate-sized antler with short fore-tine and wide
angles of the first forking". This species was,however, descriminated from C. (S.) greyi
Zdansky, known in the Middle Pleistocene
Choukoutien Fauna in North China, by having
wider angles and a lower position of the first fork.
Furthermore, it was regarded to be a species
different from C. (S.) paleoezoensis by its "wider
angle of the first fork and rather short beam
between the first and the second forks" (Otsuka
and Shikama, 1977). So far as the present statisticalanalyses of abundant antler specimens from the
Seto Inland Sea are concerned, most of these antler
specimens of C. cf. greyi (Otsuka and Shikama,1976) seem to represent antlers of middle-aged
individuals probably between about four (three)and seven (four) years old based on comparisonswith C. nippon centralis and C. nippon yesoensis.
These morphological characters of the fossil
antlers as mentioned above strongly suggest thatspecimens identified as katokiyomasai, cf. greyi,
natsumei, and paleoezoensis do indeed represent
several different growth stages of a single speciesof the subgenus Sika. Among these four speciesof Sika-deer reported from the Seto Inland Sea,
katokiyomasai (Shikama and Hasegawa, 1965) is
selected as a specific name based on the rules ofzoological nomenclature. As will be discussed later,

639

katokiyomasai can be regarded as a subspecies ofC. (S.) greyi Zdansky (Zdansky, 1925) known from
China.
Cervus greyi (Zdansky) which is a representative
fossil deer in China, is mostly known in the Middle
Pleistocene Choukoutien Fauna. The typespecimen of C. greyi (Zdansky, 1925) is represented
by the basal part of a left antler with pedicle
attached and is characterized by a much Iyrated
beam forming wide angles of more than 95 degrees
with a long first tine. By the angles of the firstforking and the diameter of the burr, this specimen
is comparable to those of C. nippon yesoensis
between five and seven years of age. This typespecimen also resembles the antler of C. greyi
katokiyomasai of the same age range, but differs
slightly from the latter in its lower first forking.
Furthermore, the antlers of C. (Pseudaxis) greyi(Zdansky) (Teilhard de Chard in and Pei, 1941) and
Pseudaxis greyi var. Zdansky (Young, 1932) arecomparable to those of the adult stage of C. greyi,
probably older than eight years of age, on the basisof their pedicle indices. The antler of Pseudaxis
greyi var. elata (Teilhard de Chardin, 1936;
Teilhard de Chardin and Pei, 1941) is regarded asthat belonging to aged C. greyi older than elevenyears. This interpretation is supportive of a theory
that many types of antler of C. greyi (s.s.) from
Choukoutien do represent various growth stagesof a single species.
4. Comparisons and observations
On the whole, the Japanese endemic species, C.
greyi katokiyomasai, is closely allied to C. greyi(s.s.) in its forking mode, but the antler of theformer is more slender than the latter through its
entire lifespan. For instance, the thickness of burrof C. greyi katokiyomasai at the prime age is about
15 mm smaller than that of C. greyi from China.
Furthermore, it is inferred that C. greyi
katokiyomasai, a representative species of the
Middle Pleistocene island form, might have derived
from C. greyi of China.

+-- Figure 14. Shed antlers of Cervus greyi kalOkiyomasai Shikama and Hasegawa. All figures are outer views,
excepting 6, x 0.3. Estimated ages of each antler are as follows: 1-3: under three years old; 4-8: three to five
years of age; 9-11: five to seven years old. 1 (NSM Reg. No. 14444-5), 2 (NSM Reg. No. 14444-4),3 (NSM Reg.
No. 14477), 4 (NSM Reg. No. YM666), 5 (Reg. No. YM469), 6 (Reg. No. YM309), 7 (Reg. No. YM810), 8 (Reg.
No. YM714), 9 (Reg. No. HM185), 10 (NSM Reg. No. 14476), 11 (Reg. No. YM665).

640

HiroYlIki OTSUKA

868.

Growth of antler in Sika

Cervus (Sika) greyi katokiyomasai is also allied
to Cervus harbinensis excavated from the latest
Pleistocene fluvial deposits at Ku-hsian-tung near
Harbin, North China (Tokunaga and Naora, 1939)
by its long first tine and short beam, but isdistinguished from the latter by much lyrated beamin the adult stage.
Cervus greyi katokiyomasai is similar to C.
nippon yesoensis in the length of beam (LB1),
however, it is distinguished by its slender antler andwide angle of the first forking.
Numerous remains of hunted Sika-deer wereexcavated from the Torihama Shellmound of the
Early Jomon Period (5910±30 years B.P.
'" 5170±30 years B.P.) in Honshu (Education
Committee of Fukui Prefecture, 1981-1985).
Judging from the mode of the forking and the sizeof the antler, Sika-deers from this shellmound arereferred to the living Yezo-sika (C. nippon yesoensis) in Hokkaido. However, they had rather smaller
antlers than those of living C. nippon yesoensis.
C. greyi katokiyomasai resembles more closely the
Yezo-sika from Torihama than those from
Hokkaido because of its wide angles of the first
fork. However, C. greyi katokiyomasai isdistinguished from the Sika-deer from Torihama
by the long beam length between the first andsecond forks (LB1), and slender antler. C. greyi
katokiyomasai is clearly distinguishable from C.
nippon centralis by the large beam length between
the first and second forks (LB1) and the wide
angles of the first fork.
The phylogeny of the subgenus Sika may be inferred as follows: In the Middle Pleistocene, apopulation of C. greyi with a pair of slender andrather small antlers migrated to the JapaneseIslands, accompanied by Naumann's elephant(Palaeoloxodon naumannt) and "MayaSu-pu-hsian" (Elaphurus mayal)" . The species
flourished over in the lowland area of west Japanand gave rise to a characteristic, endemic, island
form - Cervus greyi katokiyomasai. C. nippon

641

yesoensis of the early Jomon Period might havederived from C. greyi katokiyomasai in the Late
Pleistocene and had dispersed throughout theJapanese Islands, including Hokkaido. After the
Early Holocene, the Yezo-sika (C. nippon yesoensis) diverged into the medium-sized Sika-deer in the
Kyushu and Honshu Islands, namely the Kyushusika (c. nippon nippon), Honshu-sika (C. nippon
centralis) and a few other island subspecies (c.
nippon yakushimae, C. pulchellus etc.).
Acknowledgements

Here I wish to express my indebtedness to thefollowing people: Mr. Sotoji Imamura, Professor
Emeritus of Hiroshima University who gave memuch valuable information about the deer fossils
in the Seto Inland Sea; Professor Yoshikazu
Hasegawa of Yokohama National University whogave me much helpful advice on the paleontological problems of fossil deer; Dr. Noriyuki
Ohtaishi of the Department of Anatomy, Facultyof Dental Medicine, Hokkaido University, Dr.
Hayao Nishinakagawa and Mr. Mitsuharu
Matsumoto of the Laboratory of Veterinary
Anatomy, Kagoshima University, Dr. Hiroko
Koike of Saitama University, and Mr. Masaaki
Koganezawa of the Tochigi Prefectural Museum,
who kindly gave me permission to measurespecimens of modern Sika-deer kept in theirlaboratories and who advised me on the biologicalproblems of deer. I am also greatly indebted to Mr.
Keiichi Yamamoto of Kurashiki City, Dr. Seijun
Hinoide of Uchimicho on Shodo-shima, Mr.
Kansho Doi in Fuke-cho of Osaka Prefecture, andDr. Tamotsu Narushima of Nantan-cho on Awajishima, who gave me permission to examine theircollection of fossil deer. Thanks are also expressed
to Dr. Brian Chisholm of the Department ofArcheology, Simon Fraser University for a criticalreading of the manuscript.

+- Fi~ure 15. Antlers of Cervus greyi kalokiyomasai Shikama and Hasegawa. All figures are outer views, x
D.3. EstImated ages are as follows. 1-2: seven to nine years of age; 3-6: nine to ten years of age. 1 (Holotype
of Cervus katokiyomasai Shikama and Hasegawa, 1965),2 (Reg. No. YM4D5, 3 (Reg. No. HM22D) 4 (Reg No..
YM559), 5 (Reg. No. YM665), 6 (Reg. No. YM616).
'

Hiroyuki OTSUKA

642

Figure 16. Antlers of Cervus (Sika) greyi karokiyol/wsai Shikama and Hasegawa. All figures are o ut er views,
x 0.3 . These are estimated as aged antler, older than eleven years of age. 1 (NSM Reg. No. 14443-2), 2 (Reg. No.
YM614), 3 (Reg. No. YM470) , 4 (NSM Reg. No. 14689) , 5 (Reg. No. YM789).

References cited
Bando , Y., Saito, M. and Takahashi, K., 1978: Geologicalstud y of the sea bottom and th e geology of Bisa n-setoarea in the Japanese Inland Sea (Seto Inland Sea).
Mem. Fae. Educ. Kagawa Univ., sec. 2, vol. 28, no .
I , pp . 9-41 (in Japanese wilh English summmy).
Education Committee of Fukui Prefecture, 1981-1985:
Torihama Shellmound. Fukui: Education Commitleeof Fukui Prefecture (in Japanese).
Koike, H. and Ohtaishi, N ., 1985: Prehistoric huntin gpressure est imated by th e age composition of ex-

cavated sika-d ee r (Cervus nippon) using th e annual
la yer of too th cem en t. Jour. Archaeol. Sci., No. 12 ,p.443-456.
Hasegawa, H ., 1972: The Naumann's elep ha nt , Palaeoloxodon naumanni (Makiyama) fr om the Late
Pleistocene off Shakagahana, Shodoshima Is., in Seto
Inland Sea, Japan. Bull. Nal. Sci. Mus., vol. 15 , pp .
513-591.
Honza, H ., Kagami, H . and Nasu , H. , 1970: Geology of
Bisan-Strait, Seto Inland Sea of Japan. Jour. Marine
Ceol., vo l. 6, no. I, pp. 12-32 (in Japanese withEnglish sUl'l1mmy) .

868.

Growth of antler in Sika

Imamura, S., 1974: Distribution, stratigraphy and paleogeography of fossil elephant, cervid and molluscs inthe western Seto-naikai (Seto Inland Sea). Jub. Pub.
Prof. Kusumi, pp. 107-117 (in Japanese).
Kaji, K., Koizumi, T. and Ohtaishi, N., 1980: Population
structure of Sika deer (Cervus nippon yesoensis
Heude) in the Nakanoshima Island of Lake Toya,
Hokkaido. Jour. Mamm. Soc. Japan, vol. 8, no. 5,
pp. 160-170.
Mitchell, B., 1963: Determination of age in Scottish red deer
from growth layers in dental cement. Nature, vol. 198,
no. 4878, pp. 350-351.
- - , 1967: Growth layers in dental cement for determining
the age of red deer. Jour. Animal Ecol., no. 36, pp.
279-293.
Naora, N., 1954: Old stone age in Japan. pp. 1-298 (inJapanese).
- - , 1970: Deer fossils found in the Inland Sea of Seto.
Sci. Rep. Inst. Min. Ind., Fac. Sci. Eng., Waseda
Univ., no. 26, pp. 55-61.
Otsuka, H., 1977: Studies on fossil deer of the Doi
Collection from the sea bottom off TomogashimaIslands in Osaka Bay (Pleistocene deer fauna in Seto
Inland Sea, West Japan - Part II). Rep. Fac. Sci.
Kagoshill1a Univ. (Earth Sci., Bioi.), no. 10, pp.
41-52.
- - , 1987: Middle- and Late Pleistocene Mammalianfaunae in the Japanese Islands with special reference
to the mammalian fauna of the Nishiyagi Formationin Akashi district. Bull. Natn. Mus. Jap. His!., vol.
13, pp. 275-287 (in Japanese with English abstract).
- - and Shikama, T., 1976: Studies on fossil deer of the
Takao Collection (Pleistocene deer fauna in the Seto
Inland Sea, West Japan - Part I). Bull. Natn. Mus.,
Ser. C, (Oeol.), vol. 3, no. I, pp. 11-42.
Ohtaishi, N., 1976: Developmental variation of the antlers

on Japanese deer at Nara park (Preliminary). Ann.
Rep. Nara Deer Res. Ass. 1975, pp. 107-122.
- - , 1980: Determination of sex, age and deathreason of recovered remains of Sika deer (Cervus
nippon) by jaw and tooth-cement. Kokogaku to
Shizenkagaku (Archaeology and Natural Sciences),no. 13, pp. 51-74 (in Japanese with English
summary).
Shikama, T., 1941: Fossil deer from Japan. Jubl. Pub.
Comm. Prof. Yabe's 60th Birthday, vol. 2, p.
1125-1170.
--and Hasegawa, Y., 1965: On a fossil cervid antler
from western Japan. Sci. Rep. Yokohama Natll.
Unil'., 2nd ser., no. 12. pp. 45-47.
Teilhard de Chardin, P., 1936: Fossil mammals from
Locality 9 of Choukoutien. Pal. Sinica, ser. C, vol.
7, fasc. 4, pp. 1-61.
- - and Pei, W. c., 1941: The fossil mammals fromlocality 13 of Choukoutien. Ibid., N. S. C, no. II,
pp.76-80.
- - and Trassaert M., 1937: The Pliocene Camelidae,
Girafidae and Cervidae of southeastern Shansi. Ibid.,
N. S. no. I, pp. 1-68.
Tokunaga, S. and Naora, N., 1939: Fossil remains
excavated at Ku-hsian-tung near Harbin,
Manchoukou. Rep. First Sci. Exped. Manchoukuo,Sec. 2, PI. 4, pp. 1-229.
Young, C. c., 1932: On the Aniodaclyla from the
Sinanthropus Site at Choukoutien, Pal. Sinica, ser.
C, vol. 8, fase. 2, pp. 1-100.
Zdansky, 0., 1925: Fossil hirsche China. Ibid., ser. C, vol.
2, fasc. 3, pp. 1-90.
- - , 1927: Weitere Bermerkungen uber fossile Cerviden
aus China. [bid., ser. C, vol. 5, fasc. I, pp. 1-90.
- - , 1928: Die Saugetiere der Quartarfauns von ChoK'ou-Tien. [bid., ser. C, vol. 5, fasc. 4, pp. 1-146.

iiU'''fKJiflf.O) ~~wc~Sika iJE~O)J$i;~ : iJ/iJiPfAJiro:i.p G fi, T '7"< /' :/ '7 ~(:t t~:>lJI!.f4~tjcHtJ: E
1.:.~-t7.:,IlfIl~Li!J¥!Jfl:1:'lll+iI~~'1iH.:.~-t0o .:.0)5t>, :::'.h"1-C·~~,Illt~.ht-=:iI>, l};7-. r ~.ht-=:
Sika iJE!1ll,fi n,U':'o)l:f I), *EiliXf..JiO) f:l: iI> I.:. , m'to).=. * :/ s/ 7J ~~'HI 0) '7 ;7-. l} - s/ 7J C [nJfil[O)fl:1:'l1.W."1-C'~"1.h Ll'0 C ~ht-=::::' U~iI;0o .i!I:¥, V1ll~ifJ!p$~O)ifalJ!EtJ>G51~mlfG.ht-=:W*!i!-B:;O) " v!J Y 3 :/9"I':'fi, 120.8H~-t Sika iJEl9l,O)jEj~I79~H~·~~Iv-c·1.'0o .7chGO)jEjO)~Bllf,,'i*I':' (:t1i k 0) )f;,Ii~ . ::*: ~ ~ 0) >6 0) iii iI; I), .7c O)%i=*tEfi~ - 0) Hi 0) jJIlt#l\lltfl: -C iI; 01lJ1iE:'rti:tJ~-C'-C~t-=:o643:::'h~lz:IDE-t0t-=:cli)I':', -t!-":/ rl~t&,WI':'J:,,?-C:¥lIK\iI~-jltJt~.h-CI.'7.;m[t.=.*:/

s/7J (::r.'/Y7J, *:/Y,,- '7 s/7J , '/Y"<S/7J) 0)3fil[O)jEjO))f;1l.~· :;!;:~~O)jJQlli\\lltil:I':'",,)I.'-C~llt L.t-=:o .7cO)~*, tt*~-"1 t-=:I;tI.' <-:::>iI>O)jEjo)~*~£i.:.:lJtPg ~.h t-=:ifJ!p F9;1l!~ 0) fl:1:'lSikaiJE~0)4fil[ (katokiyomasai· natsumei· d. greyi· paleozoensis) Ii, ~-O):flIlil;01.'li
iJE~F9O)~tJ: "? t-=::¥IIi\\MO)jEj~i~~ c L -C I.' 0 .:. c iI~1!fl Gil> c tJ: "? t-=:, ~t-=:I':',l?2,~~.h t-=:Sika ill!
~fi, .7c O)jEj O)!!?fj~H.:. J: "? -C, "r 00 0) 9"M~~ fll:O)}j';] Q 1i5i!J¥!Jllr-I.:.!!?f?W<i't.J tJ: ,7" v 1 EII.lli(Cervus
greyiH.:.ib!0i1~, 1~1!H.:.J:t...,z-C, .7cO)jEjli~~'H~',I-C·iI; I), .:t-O)!lEmic.7,.,tJ:~h, ilJ¥!JiD'iHlUt:J
1:, iJE:flIli<i katokiyomasai iI~~!R ~.h, 7J r '74'-3'7 -IT s/ 7J Cervus greyi katokiyomasai ~1Jg!'l,-t~o [nJiJE~~9"M~~~o).~K, T'7"<://'7~, ~~~~iI;~, ~~:flIlO)~7J~s/7Ji!I!~(Nipponicervus)tJ:C.'C C >61':', B*7!J~O)fJl.i1£fffH.:.~*Lt.:o;:k:j:'~ m~Trans. Proc. Palaeont. Soc. Japan, N.S., No. 152, pp. 644-653, 4 Figs., December 3D, 1988

869.

SOME ABERRANT FUSULINACEANS FROM THE
UPPER CARBONIFEROUS SEQUENCE OF THE
AKIYOSHI LIMESTONE GROUP, SOUTHWEST JAPAN*
KAZUYUKI MATSUSUE
Department of Geology, Faculty of Science,
Kyushu University, Fukuoka 812

Abstract. Some aberrant lenticular fusulinaceans are found from the Upper Carboniferous
sequence in the lower part of the Akiyoshi Limestone Group, Southwest Japan. These foraminifers
are characterized by two different stages of growth; planispirally coiled in the early stage of growth
and rectilinear in the last stage. The following species are described and discussed herein;
Rectomillerella okubensis, sp. nov., R. sp. A, R. sp. B, Palaeoreichelina disvoluta, sp. nov., and
P. ? sp. As far as the present knowledges are concerned, it is possible that Palaeoreichelina ispaleobiogeographically related to the Tethyan realm.

biostratigraphy of the lower part of the Akiyoshi
Limestone Group in the Okubo area, southern
margin of the Akiyoshi Limestone Plateau, largely
examining the smaller foraminifers and primitive
fusulinaceans (Matsusue, 1986). This paper deals
with the systematic paleontology of a part of these

Introduction

The Akiyoshi Limestone Group is exposed in
the western central part of Yamaguchi Prefecture,
Southwest Japan, and forms a large limestone
plateau measuring 17 km (NE-SW) by 8 km (NWSE). This limestone sequence spans a long timeinterval from the Early Carboniferous (Tournaisian) to late Middle Permian and has been
regarded as an organic reef complex developed on
the basaltic basement of an isolated seamount(Ota, 1968, 1977; Kanmera and Nishi, 1983;
Haikawa, 1986). The limestone is generally lightgray in color without showing any stratification,
except for the basal part which includes darkreddish tuffaceous matters. Therefore, thegeological structure of this limestone can beanalyzed mainly by discriminating biostratigraphical zones and by mapping them.
These biozones have been established on the basisof fusulinaceans, brachiopods, corals, andconodonts.
The present author has restudied a foraminiferal*Received June 4, 1988; revised manuscript accepted
November 5, 1988.

Figure 1. Map showing the location of the Okubo
area in the Akiyoshi Limestone Plateau.
644

869.

Aberrant fusulinaceans from Akiyoshi

foraminifers. The present author divided the lower
part of the Akiyoshi Limestone Group into thefollowing 10 foraminiferal zones in ascendingorder: Endothyra Zone, Endostaffella sp. A Zone,
Eostaffella mosquensis Zone, Eostaffella ikensis
Zone, Millerella yowarensis Zone, Pseudostaffella
minuta Zone, Pseudostaffella antiqua Zone, Profusulinella beppensis Zone, and Fusulinellabiconica Zone. These biozones range in age fromthe early Visean to Moscovian.
A large number of fusulinaceans have been
described from the Akiyoshi Limestone Group by
many micro paleontologists (Ozawa, 1925;
Toriyama, 1958; Ota, 1971; Sada, 1975, etc.).
However, very few fusulinaceans with an aberrant
form of coiling were reported until present.
Toriyama (1953) described an aberrant fusiform
genus Akiyoshiella from the Profusulinella Zone
in the northeastern part of the limestone plateau.

645

Yanagida et al. (1971) discriminated the
Akiyoshiella ozawai Zone between the Profusulinella beppensis and Fusulinella biconica
Zones in this area.
Although the present author could not discover
any species of Akiyoshiella in the Okubo area, he
has found some aberrant lenticular fusulinaceans.
They occur at seven localities which are
stratigraphically grouped into two levels. The lower
level (Locs. MF194 and ML133) is equivalent to
the middle part of the Millerella yowarensis Zone
(redefined by Matsusue, 1986, not Ota, 1977). The
limestones of this zone is distinguished by thedevelopment of a true-reef facies. The microfaunaof the zone is characterized by the first appearanceof the genus Mil/erella, namely Mil/erella yo warensis and M. toriyamai. This microfauna is considered to be earliest Late Carboniferous age. The
upper level (Locs. MF345, MF365, MF369, MF372

Figure 2. Map showing fossil localities in the Okubo area of the southern margin of the Akiyoshi Limestone
Plateau. Elevation contours in meters. Distributions of foraminiferal zones are after Matsusue (1986).

Kazuyuki MATSUSUE

646

and MF374) is slightly above the base of theFusu/inella biconica Zone, and the zone ischaracterized by an algal micrudite. The microfauna of the zone is represented by Fusulinellabiconica, Profusu/inella, Bradyina, C/imacammina, and staffellids, and is referable to theearly Moscovian.
The present author wishes to express his sincere
thanks to Professor luichi Yanagida of Kyushu
University for his kind advice and critical readingof the manuscript. Acknowledgment is also due to
Dr. Masamichi Ota, Director of the Kitakyushu
Museum of Natural History, and Messrs. T.
Haikawa and A. Sugimura, Curators of the
Akiyoshi-dai Museum of Natural History for theirsupports of the field survey.
Paleontological notes

Aberrant fusulinaceans with a deviated coiling
pattern at the last stage of growth are common in
the Upper Permian, such as Reichelina andCodonofusiella.
Ganelina et at. (1972) discussed a taxonomic
value of aberrant forms of some Paleozoic
foraminifers. They concluded that straightened
tests had a highly taxonomic significance, and thecharacter should be recognized as a generic one.
They considered that these essentially new
characters appeared in the advanced species of agenus clearly represented the qualitative change forthe new genus, whereas the increase in number ofchambers in the rectilinear part revealed only the

specific or subspecific character.
The aberrant forms are divided into two fundamentally different groups by Liem (1974). The firstgroup (Figures 3-A, B) is planispirally coiled except for the last volution, and then the last volution abruptly increases its height. In the transverse
sections, the last volution takes a fan-shape.
Foramina are basal throughout the growth. The
second one (Figures 3-C, D) is coiled in the inner
volutions, but then test becomes straightened at thelast stage of growth. Chambers of the straightenedpart are arranged uniserially and rectilinearly, andhave an almost constant dimension both in widthand height.
The term "uncoiling" has been used in manydifferent senses, representing a rapid increase inheight of volution, straightening, and even evolutecoiling. Liem (1974) proposed to use the term "uncoiled" (razvernutyi) in a restricted sense to definesuch a fan-shape as developed in the last volutionof Reiche/ina, and to use the term "rectilinear"(vypryamlennyi) to define a straightened shape in
the last stage of growth. He also proposed to use
these terms not only for fusulinaceans but also forendothyrids.
Leven (1970) established the genus
Pseudoreiche/ina, characterized by an aberrant
lenticular form, from the Lower Permian ofDarvaz of Pamir, USSR. In general, the form of
Pseudoreiche/ina is similar to that of Reiche/ina,
but the former genus is quite distinct from thelatter by having the staffellid wall. Some Early

A

Figure 3.
Schematic sections of the aberrant fusulinaceans(after Liem, 1974). A, B: uncoiled fusulinaceans. C, D: rectilinear
fusulinaceans.

D

869.

Aberrant fusulinaceans from Akiyoshi

Permian species described as Reichelina were included by Leven in the genus Pseudoreichelina. He
suggested that Pseudoreichelina evolved from astaffellid ancestor in the Early Permian, andReichelina, on the other hand, evolved from an
ozawainellid stock in the Late Permian.
Subsequently, Liem (1974) described anaberrant staffellid genus Palaeoreichelina from the
Moscovian of Vietnam, Southeast Asia. This genus
is lenticular in the early coiled part and rectilinear
in the last stage of growth.
At the same time, he established an aberrant
ozawainellid genus Rectomillerella which wasrectilinear in the last stage of growth.
Carboniferous foraminifers with a straightenedpart in the last stage of growth have been wellknown as Endothyranella since Harlton and
Galloway established it in 1930. This genus shares
the same character with Endothyra in the earlycoiled part, but the coiled part straightens out inthe last stage of growth. Generally, it is regarded
that Endothyranella is streptospirally coiled, andmost abundant in the early Late Carboniferous,
especially in the Moscovian interval (Armstrong
and Mamet, 1977, p. 53). Rectomillerella is similar
to Endothyranella in the general shape of the test.
But the former genus differs from the latter by its
planispiral coiling in the early stage of growth.
Davydov (1981) discussed the morphology andsystematics of two genera Pseudoreichelina andPalaeoreichelina. He described some aberrant
staffellids, from the Upper Carboniferous ofDarvaz, which are uncoiled in the last stage of
growth. Although the character of the last stageof them disagreed with Liem's original diagnosis,
Davydov conditionally included them with thegenus Palaeoreichelina. Furthermore, he remarked
that the Early Permian genus Pseudoreichelina also
included two types in the last stage of growth.
Therefore, this implies that the uncoiled andrectilinear types coexist in the Late Carboniferous
and in the Early Permian genera. He concluded
that Pseudoreichelina and Palaeoreichelina weredistinct from each other, since they were originatedfrom a different species and also they havemarkedly different stratigraphic ranges. RauzerChernousova (1985) noted that further details were647

required on these two genera to discuss theirgeneric in dependences from each other. The present author herein wants to regard them as independent genera as suggested by Davydov.
Paleobiogeographical significanceof aberrant fusulinaceans from Akiyoshi
The occurrence of aberrant lenticular fusulinaceans in the Late Carboniferous and the Early
Permian is scarce globally. The Early Permianaberrant fusulinacean, Pseudoreichelina has beenreported from Pamir of USSR (Leven, 1970),
Yugoslavia (Kochansky-Devide, 1966), Nevada ofU. S. A. (Douglass and Nestell, 1974), and South
China (Huang and Zeng, 1984). On the other
hand, Palaeoreichelina was originally describedfrom the lower Upper Carboniferous of Vietnam
and Thailand, Southeast Asia. Davydov (1981)
described some species, conditionally attributing
them to the genus Palaeoreichelina from the
Moscovian of Darvaz, Pamir. These two genera
are very close in their geographical distribution.
Although Pseudoreichelina has not been reported
from Japan, the occurrence of Palaeoreichelina in
the Akiyoshi Limestone Group strongly suggests
a close paleobiogeographic relationship between
the above mentioned areas and the Akiyoshi
Limestone Group.
Summarizing the Late Carboniferous aberrant
fusulinaceans, it is apparent that they arecharacteristic elements of the Tethyan realm. These
genera together with the Upper Permian aberrant
ones formed a part of important constituents of
the Tethyan fusulinacean fauna.
Systematic descriptions
All the described specimens are kept in the
Department of Geology, Faculty of Science,
Kyushu University whose abbreviation GK is theprefix for the specimens' repository numbers.
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinacea Moeller, 1878
Family Ozawainellidae Thompson and Foster,
1937

Kazuyuki MATSUSUE

648

Genus Rectomillerella Liem, 1974

Rectomillerella okubensis, sp. nov.

Type species:-Rectomillerella texasensis Liem,
1974, p. 25-26. = Millerella marblensis var.
Moore, 1964, p. 301-305, pI. 48, figs. 1-14,
19-21.

Figures 4- I, 2

Mifferella; Thompson, 1951, pI. 14, fig. 4 (only)
Mifferella; Moore, 1964, p. 301-305, pI. 48, figs. 1-14,

19-21.
Rectomillerella; Liem, 1974, p. 25-26.
Rectomifferella; Loeblich and Tappan, 1988, p. 253.

Diagnosis:-Test is composed of two parts. The
early coiled part is lenticular, planispirally coiled,involute in the inner volutions and evolute in thelast one. The straightened part of the last growthstage is rectilinear and uniserial. Wall is thin andconsists of a tectum, primatheca and upper
tectorium. Septa are straight. Foramina aresimple, basal in the coiled part, becoming medianin the rectilinear part. Secondary deposits are
weakly developed in the coiled part as upper
tectorium.
Remarks:-The type species of this genus was
originally described as a variant of Millerellamarblensis Thompson by Moore (1964). He
already regarded that this variant had a possibilityof being a new species of Millerella. In 1974, Liem
established the monotypic genus Rectomillerella
based on Moore's specimens, renaming it as R.
texasensis Liem.
The early coiled part of Rectomillerella is very
similar to that of Meillerella. But the former differsfrom the latter by having a rectilinear part in thelast stage of growth. The general test morphology
of Rectomillerella resembles Palaeoreichelina, butthey are quite distinct from each other by thedifference in wall structure. Rectomillerella is also
very similar to Endothyranella in the general test
form and has a similar stratigraphic range. The
difference between Rectomillerella and Endothyranella is only based on the mode of coiling;
the former is planispirally coiled and the latter iscoiled streptospirally.
Three species of Rectomillerella arediscriminated from the Akiyoshi LimestoneGroup.

Material studied:-Description is based on asagittal section (holotype, GK-D7001a) and atangential section of paratype which is obliquely
cut along the rectilinear part (GK-D7001b).
Description:-Test is small, composed of twoparts. Whole length of test in a direction parallelto the rectilinear part is 416 pm. The early partis planispirally coiled, lenticular. Peripherysubrounded in the inner two volutions andsubangular in the last one half volution. Coilingis two and a half volutions, involute in the inner
volutions and evolute in the last one, and expands
gradually and uniformly. The last volution contains 15 chambers. The later rectilinear part isuniserial, cylindrical, about 80 pm in width, 261
pm in length, and consists of 6 chambers. Septal
spacings are constantly disposed with the lengthof about 43 pm. Proloculus is small, 24 pm indiameter. Septa are almost straight or slightly
curved forward. Septal sutures are not seen.
Secondary deposits are weakly developed as inconstant pseudochomata in the coiled part.
Foramina are basal in the coiled part, but in therectilinear part they are not visible on account of
the obliquely oriented section. Wall is thin, dark,
and undifferentiated. Thickness of wall is 13 pmin the last volution and 8 pm in the rectilinear
part.
Remarks:-The present species has a rectilinear
part in its last stage of growth and develops
planispiral volutions in the coiled part with a thin,
microgranular, and undifferentiated wall.
Therefore, this species should be attributed to thegenus Rectomillerella. Rectomillerella okubensis
differs from R. texasensis Liem by its narrower
rectilinear part and more angular periphery in thelast volution.
Occurrence:-Loc. MF194 in the Millerella
yowarensis Zone (earliest Late Carboniferous).
Rectomillerella sp. A
Figures 4-3, 7, 9.

Material studied:-Description is based on a

869.

Aberrant fusulinaceans from Akiyoshi

tangential section (GK-D7002a) and two obliquesections (GK-D7002b and GK-D7007).
Description:-Test is composed of two parts.
The whole length of the test ranges from 540 to
637 11m. The early part' is lenticular withsubangular periphery, and is an almost planispiralcoil with a gradual rate of expansion. Number of
vol uti on is 2 to 3. Diameter of the coiled part is
between 360 and 502 11m. The later rectilinear partcomprises up to 4 chambers. Width of therectilinear part is the same as that of the coiled
part, measuring from 233 to 280 11m. Septal
spacings in the rectilinear part are constant andsimilar to those of the coiled part. Septa are slightly
convex forward throughout the growth. Septal
sutures are visible. Foramina are basal in the coiled
part, but central in the rectilinear part. Wall is thin,dark, and undifferentiated. Thickness of wall
ranges from 10 to 14 11m in the last volution andfrom 14 to 20 11m in the rectilinear part.
Secondary deposits are present as chomata in the
coiled part and also appear as thickenings of septa
around the foramen in the rectilinear part.
Remarks:- This species differs from other taxa
of Rectomillerella by its comparatively large test.
The presence of thickenings of septa around theforamen in the rectilinear part reminds the present
author of the apertural neck of Endothyranella
recta described by Rich (1980, p. 27). However,this species belongs to Rectomillerella because of
its planispirally coiled volutions in the early stageof growth.
Occurrence:-Loc. ML133 in the Millerella
yowarensis Zone (earliest Late Carboniferous), andLoc. MF345 in the Fusulinella biconica Zone
(Moscovian).

Rectomillerella sp. B
Figures 4-4, 5.

Material studied:-Description is based on two
tangential sections (GK-D7003 and GK-D7(06).
Description:- Test is composed of two parts.
Whole length of the test ranges from 820 to 899
11m. The early coiled part is lenticular, planispirally coiled, and periphery becomes sub angularin the last volution. So far as visible, diameter of649

the coiled part measures from 364 to 447 11m, andaxial length from 137 to 164 11m. The rectilinear
part is relatively long, being composed of 3 or 4
chambers. Width of the rectilinear part is almost
constant, between 85 and 137 11m, and narrower
than the axial length of last volution. Septal
spacings in the rectilinear part are relatively longin comparison with the width of chamber. Septa
are straight in the coiled part and convex forwardin the rectilinear part. Wall is thin, dark, and undifferentiated in the coiled part, and partiallydifferentiated with a tectum, primatheca and innertectorium in the rectilinear part. Thickness of wallis 12 11m in the last volution, and 13 11m in therectilinear part. Foramina are median in therectilinear part. Secondary deposits are weakly
developed as pseudochomata in the coiled part,and as thickenings of septa around the foramenin the rectilinear part.
Remarks:- This species differs from other taxaof this genus in having the long rectilinear partwhich shows a comparatively rapid expansion.
Occurrence:-Locs. MF365 and MF374, in the
Fusulinella biconica Zone (Moscovian).
Family Staffellidae Miklukho-Maklay, 1949
Genus Palaeoreichelina Liem, 1974

Type species:-Palaeoreichelina donghoiensis
Liem, 1974, p. 25, figs. 1-8.
Pseudoendothyra (part); Igo, 1972, p. 75, pI. 9, figs.
22-25.
Palaeoreichelina; Liem, 1974, p. 24-25.
Palaeoreichelina?; Davydov, 1981, p. 121-124.
Palaeoreichelina; Rauzer-Chernousova, 1985, p. 15.
Palaeoreichelina; Loeblich and Tappan, 1988, p. 286.

Diagnosis:- Test is composed of two parts;
the essentially lenticular, involute, planispirally
coiled part with sub angular periphery, and thelast straightened part with uniserially arranged
chambers. Wall is thick, undifferentiated in the
inner volutions, but becomes three-layered withluminotheca in the last volution and in therectilinear part. Septa are straight. Foramina aresimple, basal in the coiled part, and median in therectilinear part.

650

Kazuy uki MA TSUSUE

869.

Aberrant fusulinaceans from Akiyoshi

Remarks:-Previous authors all agreed toattribute this genus to the Staffellidae by its wall
structure, which is clearly distinguishable from thatof other fusulinaceans by the presence ofluminotheca in the last stage of growth.
However, there is a problem which the last stageof this genus is either uncoiled or rectilinear. Liem
(1974) noted that the last stage of growth of
Palaeoreichelina gave rise to the rectilinear form
with uniserial arranged chambers. He conditionally
included Pseudoendothyra sp. C of Igo (1972, pI.
9, figs. 4, 5, 22-24) in Palaeoreichelina. Davydov(1981) described three new species from the Upper
Carboniferous of Darvaz, namely Palaeo reichelina? igoi, P.? distincta, and P.? vozginensis.
He stated that all of them were uncoiled in the laststage of growth. In spite of his disagreement withthe original diagnosis given by Liem (1974),
Davydov conditionally attributed his three species
to Liem's Palaeoreichelina. He assigned thosespecimens figured by Igo (1972) to P.? igoi which
was described as having the uncoiled form in thelast stage of growth. Therefore, the genus
Palaeoreichelina now has two different types in thelast stage of growth. The Early Permian genus
Pseudoreichelina also has two types in the last stageof growth and is very similar to Palaeoreichelina.
However, they are quite different from each otherin their stratigraphic ranges.
It is probable that the uncoiled species belongs
to a new genus. This problem has not been solvedin this study, but a specimen from Akiyoshi shows
the rectilinear mode in the last stage of growth.
These two species are discriminated from the
Akiyoshi Limestone Group.
Palaeoreichelina disvoluta, sp. nov.
Figures 4-8, 10.

Material studied:-Description is based on a651

sagittal section (holotype, GK-D7004) and an
oblique section (GK-D7005a).
Description:- Test is small, planispirally coiled
with four volutions. Coiling expands moderately,
but bluntly increases its height at the last stage of
growth. Last two chambers straighten out, andbecome free from the previous volution. Septa arestraight or slightly convex and incline forward.
Sutural depressions are unrecognizable. Wall israther thick, but wall structure is indistinct due torecrystallization. Wall is undifferentiated, dark,
microgranular in the inner volutions anddifferentiated with luminotheca in the last stage of
growth. Whole length of test ranges between 862
and 894 p.m. Thickness of wall in the last volution is 23 p.m, and that of antetheca 22 pm.
Foramina are basal in the coiled part, and not
visible in the last stage of growth.
Remarks:- The recrystallized, indistinct wall
structure of this species well reveals character ofstaffellids. The aberrant stage of growth is poorly
developed in the described specimens. However,a figured specimen (Figure 4-8) shows a tendency
for the last few chambers to become free from theearlier portion of the test. Although foramina of
the rectilinear part are unclear, it is highly possiblethat the test of this species is rectilinear in the laststage of growth. These features are taken to assignthat this species to Palaeoreichelina.
Occurrence:-Locs. MF369 and MF372 in the
Fusulinella biconica Zone (Moscovian).

Palaeoreichelina? sp.
Figure 4-6.

Material studied:-Description is based on atangential section (GK-D7005b).
Descriptive remarks:-Only one tangential
section is available. Test is subspherical in the earlystage of growth, but in the last stage it becomes~ Figure 4-1-2. Rectomilfere/la okubensis, sp. nov., 1, sagittal section of the holotype, Loc. MF194, GK-D700Ia;
2, tangential section of the paratype which cuts the rectilinear part obliquely, Loc. MF194, GK-D7001 b. 3, 7, 9,
R. sp. A, 3, tangential section, Loc. MF345, GK-D7002a; 7, 9, sagittal sections, Loc. MLl33 and MF345, GK-D7007
and 7002b, respectively. 4, 5, R. sp. B, tangential sections, Loc. MF365 and MF374, GK-D7003 and 7006, respectively.
8,10, Palaeoreichelina disvoluta, sp. nov., 8, sagittal section of the holotype, Loc. MF369, GK-D7004; 10, obliquesection, Loc. MF372, GK-D7005a. 6, P.? sp., tangential section, Loc. MF372, GK-D7005b. All figures x 100.

Kazuyuki MATSUSUE

652

lenticular with prolonged, keeled periphery, concave lateral slopes, and broadly rounded umbilicalregions. Luminotheca is not visible, however, therecrystallized and obscure wall structure of thepresent species reminds the author of the characterof staffellids. Measurements are as follows: width,
602 ,urn; length, 261 ,urn; thickness of wall in thelast stage of growth, 13 pm. Although no sagittal
section of this species is known, a sudden increaseof height of volution at the last stage is interpreted
to indicate the uncoiling mode of growth of thisspecies. As mentioned above, this species is probably assignable to Palaeoreichelina. However, thepresent author could not conclude whether or not
the last stage of growth of this species reveals an
uncoiled or rectilinear type.
Occurrence:-Loc. MF372 in the Fusulinellabiconica Zone (Moscovian).
References
Annstrong, A. K. and Mamet, M. L., 1977: Carboniferous
microfacies, microfossils, and corals, Lisburne Group,
Arctic Alaska. U. S. Geol. Surv. Prof. Paper 849, p.
1-144, pis. 1-39.
Davydov, V. I., 1981: Novye vidy srednekamennougol'nykh paleoreichelin [New species of the Middle
Carboniferous genus Palaeoreichelina]. Paleont.
Zhurnal, 1981, no. 3, p. 120-124 (in Russian).
Douglass, R. C. and Nestell, M. K., 1974: The Pennianfusulinid genus Pseudoreichelina from the Pequop
Fonnation, Spruce Mountains, Elko County, Nevada.
Jour. Paleont., vol. 48, no. 6, p. I 170-1 173, pI. 1.
Ganelina, R. A., Grozdilova, L. P., Lebedeva, N. S. andSosnina, M. I., 1972: Taksonomicheskoe znachenie
priznaka vypryamleniya rakoviny paleozoiskikh
foraminifer [Taxonomic significance of the test' uncoiling foraminifera of Paleozoic]. Voprosy
Mikropaleont., vol. 15, p. 30-39, pis. 1-4. (inRussian).
Haikawa, T., 1986: Lower Carboniferous of the Okubo
area in the Akiyoshi Limestone Plateau, Southwest
Japan. Bull. Akiyoshi-dai Mus. Nat. Hist., no. 21,
p. 1-35, pis. 1-7 (in Japanese with English abstract).
Huang Zhixun and Zeng Xuelu, 1984: The early Early
Permian (Longlinian Stage) fusulinid fauna from
Longlin, Guangxi. Earth Sci.-Jour. Wuhan College
Geol., vol. 26, no. 3, p. 11-24, pis. 3-4 (in Chinesewith English abstract).
Igo, Hisayoshi, 1972: Fusulinacean fossils from Thailand.
part VI. Fusulinacean fossils from north Thailand.
(Contributions to the Geology and Palaeontology ofSoutheast Asia, XCVI). Geol. Palaeont. Southeast

Asia, vol. 10, p. 63-116, pis. 9-19.
Kanmera, K. and Nishi, H., 1983: Accreted oceanic reef
complex in Southwest Japan. p. 195-206. In :Hashimoto, M. and Uyeda, S. (eds.) , Accretion
Tectonic in the Circum Pacific Regions. Terra Sci.
Pub. Co., Tokyo.
Kochansky-Devide, V., 1966: Ober das geologische Alterder Gattung Reichelina und die Reichelinen Jugoslawiens. Eclogae Geolgicae Helvetiae, vol. 59, p.
39-45, pI. 1.
Leven, E. Ya., 1970: Novyi rod permskikh aberrantnykh
fusulinid [A new genus of aberrant Permian
fusulinids]. Paleont. Zhurnal, 1970, no. 4, p. 16-20,
pI. I. (in Russian).
Liem, N. V., 1974: 0 razvernutykh i vypryamlennykh
fuzulinidakh [On uncoiling and rectilinear fusulinids].
Voprosy Mikropaleont., vol. 17, p. 22-26, I pI. (inRussian with English abstract).
Loeblich, A. R., Jr. and Tappan, H., 1988: Foraminiferal
genera and their classification. 970 p., 847 pis., Van
Nostrand Reinhold Comp., New York.
Matsusue, K., 1986: Foraminiferal biostratigraphy of thelower part of the Akiyoshi Limestone Group. Sci.
Repts., Dept. Geol., Kyushu Univ., vol. 14, no. 4,
p. 163-185, pis. 2-7 (in Japanese with English
abstract).
Moore, W. L., 1964: Notes on the morphology andtaxonomic position of the fusulinid Millerella marblensis Thompson. Jour. Paleont., vol. 38, no. 2, p.
294-305, pis. 47-48.
Ota, M., 1968: The Akiyoshi Limestone Group: ageosynclinal organic reef complex. Bull. Akiyoshi-dai
Sci. Mus., no. 5, p. 1-44, pis. 1-31 (in Japanesewith English abstract).
- - , 1971: Faunas and correlation of the "Uzura" quarry
limestone of Akiyoshi, Southwest Japan, part 2.
fusulininan fauna. Ibid., no. 7, p. 65-74, pis. 12-13.
- - , 1977: Geological studies of Akiyoshi. Part I. Generalgeology of the Akiyoshi Limestone Group. Bull.
Akiyoshi-dai Mus. Nat. Hist., no. 12, p. 1-33, pis.
1-3.
Ozawa, Y., 1925: Palaeontological and stratigraphicalstudies on the Permo-Carboniferous Limestone of
Nagato, PI. 2, Paleontology. Jour. Coli. Sci. Imp.
Univ. Tokyo, vol. 45, no. 6, p. 1-90, pis. 1-14.
Rauzer-Chernousova, D. M., 1985: Sistematika semeistva
Staffellidae (Fusulinida) [Systematics of the family
StaffeIlidae (Fusulinida}J. Voprosy Mikropaleont.,
vol. 27, p. 5-23 (in Russian with English abstract).
Rich, M., 1980: Carboniferous calcareous Foraminiferafrom northeastern Alabama, south-central Tennessee,and northwestern Georgia. Cushman Found. Foram.
Res. Spec. Publ., no. 18, p. 1-62, pis. 1-22.
Rozovskaya, S. E., 1975: Sostav, sistema i filogeniya
otryada fuzulinida [Composition, phylogeny andsystem of the order Fusulinida]. Tr. Paleont. Inst.,
vol. 149, p. 1-267, pis. 1-35 (in Russian).

869.

Aberrant fusulinaceans from Akiyoshi

Sada, K., 1975: Early and Middle Pennsylvanian
Fusulinacea from Akiyoshi. Mem. Fac. Integrated
Arts and Sci., Hiroshima Univ., sec. 4, no. I, p.
1-29, pIs. 1-7.
Thompson, M. L., 1951: New genera of fusulinid
foraminifera. Contr. Cushman Found. Foram. Res.,
vol. 2, no. 4, p. 115-119, pIs. 13-14.
Toriyama, R., 1953: New peculiar fusulinid genus from the
Akiyoshi Limestone of southwestern Japan. Jour.

653

Pa!eont., vol. 27, no. 2, p. 251-256, pIs. 35-36.
- - , 1958: Geology of Akiyoshi. Part 3, fusulinids of
Akiyoshi. Mem. Fac. Sci. Kyushu Univ., ser. D, vol.
7, p. 1-264, pIs. 1-48.
Yanagida, 1., Ota, M., Sugimura, A. and Haikawa, T.,
1971: On the geology of the northeastern part of the
Akiyoshi limestone plateau. Sci. Repts., Dept. Geo!.,
Kyushu Univ., vol. II, no. I, p. 105-114, pIs. 7-10
(in Japanese with English abstract).

Akiyoshi tt~. Okubo*:z 1*

M~~~~~U~$~.*~~.~6R~~7~9~.:Wu~M~~~~~U~$~~~$~.*~~R~~7~9~.~~~Lk. =hG~R~~7~9~.~.~~~m~K~~OO~@lL. 1~JtIli.::.f;t~v~fij1Ht-ci![m.;jjCi=tJ:6. 4-@lRectomillerella okubensis. sp. nov .. R. sp. A,
R. sp. B. Palaeoreichelina disvoluta. sp. nov .. P. ? sp. ~ F,c JIll(: L t:.. mtE ~D G h -C l' 6
Palaeoreichelina~tl~4'kJ:tt!lJl~~:5:r,fIH;t T T ;<,:tt!l.l!!.X c I*Jjf'/iv;1$l '.
1"1-* 5flJ2.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 152, pp. 654-655, I Fig., December 30, 1988

SHORT NOTES21.

NEWLY DISCOVERED LATE TRIASSIC HEXACORALSFROM CENTRAL NEPAL*NOBUO YAMAGIW A

Department of Earth Science, Facuity of Education, Osaka Kyoiku University, Osaka 543

KAZUYUKI SHIRAISHINational Institute of Polar Research, Tokyo 173
andMITSUO YOSHIDA
Department of Geology, Geoscience Co. Ltd., Tokyo, 113

Some hexacoral specimens were collected by
Shiraishi in 1977 and Yoshida and Tokuoka in
1982 from limestone in the formation reported byBordet et al. (1975) as Rhaetian age at the hill slope
northeast of Jarsgeng River in Nyi-Shang area,
Central Nepal. According to Bordet et al., theformation is composed of sandstone, shale andlimestone. They collected pelecypods andfasciculate corals from the limestone, but the
species belonging to them (pelecypods andfasciculate corals) were not reported. Therefore,
the exact age of the formation was not clear.
The present authors found the following
hexacorals in a collection from the limestone.
I. "Thecosm ilia" sp. A
2. "T." sp. B3) from the Rhaetian in the Tarta Mountains inmany respects, but exact comparisons cannot bemade because of poor preservation of the material
in longitudinal section.
The second species (Fig. 1-2) resembles original
specimens of "Thecosmilia" norica, described byFrech (1890, p. 9, pI. I, figs. 14-24a; pI. 10, figs.
6, 6a) from the Upper Norian to Rhaetian
Zlambach beds but differs from the latter in having
smaller corallites.
The third species (Fig. 1-3) is closely allied tooriginal specimens of Thamnasteria (Astraeomorpha?) major Vinassa de Regny (1915, p. 103,
pI. 68 (6), figs. 13-15) from the Upper Triassic
in Timor in many important characters.
Judging from the evidence given above, the ageof the present hexacorals indicates Late Triassic(probably Late Norian to Rhaetian). They will be
described in detail on another occasion.
This short paper is the first report of a Late
Triassic hexacoral faunule found in Nepal.
The authors express their hearty thanks toProfessor Makoto Kato of Hokkaido University,
Professor Emeritus Keiji Nakazawa of Kyoto
University and Dr. Ienori Fujiyama of the National
Science Museum, Tokyo for their kind advice and3. Thamnastel ia (Astraeomorpha?) major
Vinassa de Regny
The first species (Fig. I-I) is similar to"Thecosm ilia" clathrata (Emmrich), described byFrech (1890, p. 15, pI. 4, figs. 1-5, 7-11) andZankl (1969, p. 29, pI. 9, fig. I) from the Upper
Norian and Rhaetian in the Austrian Alps and also
by Roniewicz (1974, p. 108, pI. 4, fig. I, pI. 5, fig.
*Received September 1, 1988; accepted October 20,1988.
654

Shorr Notes 21. Triassic H exacorals

655

suggest ions, and to Professor Takao Tokuoka of
Shimane Uni versity, who permitted them to study
th e present interesting materials.
References

1mm
'--'
Figure 1.
1. " Th ecosmilia" sp. A - Tran sve rse sec ti o n
2. " T." sp. B - Transverse sect ion
3. Thamnasferia (Asfra eo llw lph a?) lI1ajor Vin assa cle
Reg ny - Tran sve rse sect ion

Bo rcl et, P. , Colchen, M. & Le Fort, P ., 1975: Recherc hes
geo logiqu s clans L' Him araya clu Nepal, Region clu
Nyi-Shang. Edifions du Cel1fre Nafional de laBecherch e Scienfique, Paris p. 1- 138.
Frech, F. , 1890: Di e Kora ll enfa un a cl er Trias. 1. Di e
Kora llen cler lu vav isc hen Trias-Prov in z. Palaeol1fographica, vo l. 37, p . 1-11 6, pis. 1-2 1.
Ro ni ew icz, E. W. A. , 1974: Rh aeti an corals of the T arta
Mts. ACfa Geol. Pol., vo l. 24 , no. I, p. 97-1 16, pi s.
1- 10.
Vin assa cle Regny, P. , 19 15: Triadisc he A lge n, Spongien,
A nth ozoaen und Bryozoen aus Timor. Palaeo l1fol.
Tim or, vo l. 4, p. 5- 118, pis. 62- 72 ( 1- 10) .
Zankl, H. , 1969: Der Ho he Goli. Aufba u und Lebe nsbild
ein es Dachste ink alk-Riffes in cl er Obertrias cler
N6rd li chen Kalkalpen. Ahh. Senckenberg. Nafurforsch . Ges ., vo l. 519 , p. 1- 123, pis. 1- 15.

Trans. Proc. PaJaeont. Soc. Japan, N. S., No. 152, pp. 656-658, 2 Figs., December 3D, 1988

SHORT NOTES

22. A PERMIAN RUGOSE CORAL FROM KAMIKAGEMORI,
CHICHmU CITY, SAITAMA PREFECTURE, CENTRAL JAPAN;!
HISA YOSHI IGO
Institute of Geoscience, University of Tsukuba, Ibaraki, 305

Fossiliferous Upper Paleozoic limestones are
frequently intercalated in the so-called Chichibu
Paleozoic System in the Kanto Mountains. Theselimestones were interpreted as autochthonousinterbeds and were accepted as the most reliableindicators of geochronology of the Chichibu
System. Fossils described from these limestones aremostly fusulinaceans. Corals and other fossils arealso locally abundant, but these fossils have notyet been described. Many thin sections of corals
collected by Fujimoto and myself are now kept at
our institute and some of these specimens seem tobe important to discuss the similarity of coralfaunas reported from the various limestone blocks
in the Japanese Islands.
A Permian rugose coral, Yokoyamaefla(Yokoyamaefla) tertioseptata (Yokoyama) which
was collected at limestone quarry of
Kamikagemori, Chichibu City, Saitama Prefecture(Fig. 1) is described herein. This species has beenreported from the Taishaku (Yokoyama, 1960) andAtetsu Limestones (Yamagiwa, 1962) of southwest
Japan. Limestone of this quarry is pale gray,massive to thickly bedded, and it yields abundantfusulinaceans. Fujimoto (1936) described Triticites
simplex (Schellwien) and several species of
Pseudofusulina. Subsequently, Morikawa (1955)
added the occurrence of Schwagerina and otherspecies of Pseudofusulina. Some of the listed
species by them have need of taxonomic revision,
but they are characteristic representatives of theupper part of the Pseudoschwagerina Zone or theso-called Pseudofusulina Zone of Japan, and they
indicate a Sakmarian age.
Recently, Hisada (1983) studied the geology ofthis area, and he interpreted the mentioned
limestone as a large allochthonous block embedded
in his sheared strata of the Urayama Group. The
Urayama Group is now believed to be a Jurassic
olistostrome by Hisada (op.ciL).
Before continuing the paleontological description, I thank the late Professor Haruyoshi
Fujimoto for his offer of the present material. The
present study is supported by the Grant-in-Aid for
Scientific Research from the Ministry of Education, Science and Culture (No. 62540597).
Description of species

Family Waagenophyllidae Wang, 1950
Genus Yokoyamaefla Minato and Kato, 1965

Figure 1.

*Received September 30,1988; accepted October 20,1988.
656

Map showing fossil locality (x).

Short Notes 22. Permian coral

2a
Figure 2.
Yokoy al71aellalerrioseptala (Yoko ya ma)

657

2c

(Yokoyal11aella)a, Tangential section; b, tan gential sec tio n of the peri·
pheral ponion of co rallum ; c, lon gitudinal secti on.
x 3.8 .

Subgenus Yokoyamaella Minato and Kato,
1965

Yokoyamaella (Yokoyamaella) tertioseptata(Yokoyama, 1960)
Figures 2a-c
Slylidophy llum yokoyal11ai lerlioseplalUl11 Yokoyama,

1960, p. 245 , pI. 27, fig s. la-c; Yamagiwa, 1962, p. 91 ,pI. 4, figs. 4-6.
Yokoy amaella (Yokoya l11aella) lerr ioseptata; MinalO andKato, 1965, p. 139-140, pI. II, fi gs. 1-2.

Des-criplion .- Corallum compound, cerioid andattains more than 30 cm in diameter. Corallites
irregularly polygonal, but mostly five sided withstraight or slightly curved walls in transverse
section. Mature corallites about 7 to 10 mm indiameter. Periphery of corallum surrounded by acircular outer wall. Some corallites tend to circularin transverse section and partly fasciculate inperipheral portion of corallum. Walls originally athin dark layer but strengthened by thick mural
septa. Median layer of wall sometimes translucent.

Septa two orders, major and minor andstrengthened by a very thick coating in peripheral
zo ne and forming mural septa. They ha ve a darklayer in their middle part and dila ted bystereoplasmic deposits. Middle part of septa,
however, rarely provides a translucent layer insome corallites. Major and minor septa partly
interrupted by lonsdaleoid dissepiments where they
discontinuous and appear as a septal projection .

658

Hisayoshi 100

Major septa long, sinuous or more or less zigzag,
and reach nearly axial structure. Counter septum
directly unites with median plate of axial structure.
Minor septa alternate with major septa but slightly
shorter than the latter. Very short septa
sporadically developed between minor septa insome immature corallites.
Axial structure more or less solid and consistsof thick median plate conjoined with c~>unter
septum, and encircling axial tabellae. Septal
lamellae poorly developed and form loose spiderweb structure in some corallites.
Dissepiments mostly concentric and not so
numerous. Rather small vesicular lonsdaleoid
dissepiments partly developed in every corallite.
In longitudinal section peripheral dissepimentarium, very narrow tabularium and distinct
axial zone well differentiated and they dilated bystereoplasmic deposits. Dissepimentarium consistsof several rows of elongate and rather large
dissepiments facing inward. Axial column consistsof a thick median plate, steeply ascending crowded
tabellae and sporadically developed septal lamellae.
Tabulae very narrow, thin and nearly horizontal.
Remarks:- The type specimen of this species
was originally described by Yokoyama (1960) fromthe Taishaku Limestone of Hiroshima Prefecture.
Subsequently, Minato and Kato (1965) criticized
Yokoyama's original specimen, and they pointed
out that the holotype of Y. (Y.) tertioseptata does
not have any true tertiary septa. The present form
rarely has the so-called tertiary septa in some
immature corallites. As already suggested by
Minato and Kato (op. cit.) these short septa seemto be immature secondary septa. Translucent layer
which was also pointed out by Minato and Katoin the median part of septa and wall appears more
frequently than that of the original specimen.
Occurrence.-Collected from Loc. 12(Morikawa, 1955, Fig. 9) of limestone quarry of
the Showa Denko Co. LTD., Kamikagemori,
Chichibu City, Saitama Prefecture.
DepositolY. - Paleontological collection, Institute of Geoscience, University of Tsukuba, Reg.
No. IGUT 5400 (Collected by Haruyoshi Fujimotoin 1960).
References cited
Fujimoto, H. (1936): Stratigraphical and palaeontologicalstudies of the Titibu System of the Kwantomountainland, Part 2, Palaeontology. Sci. Rep.
Tokyo Bunrika Daigaku, sec. C, vol. 1, no. 2, p.
29-125, pis. 1-26.
Hisada, K. (1983): Jurassic olistostrome in the southern
Kanto Mountains, central Japan. Sci. Rep. Insl.
Oeosci., Univ. Tsukuba, sec. B, vol. 4, p. 99-119.
Minato, M. and Kato, M. (1965): Waagenophyllidae. Jour.
Fac. Sci., Ser. 4, Oeol. Mineral., vol. 12, nos. 3-4,
p. 1-241, pis. 1-20.
Morikawa, R. (1955): Schwagerininae in the vicinity of the
Shomaru Pass, eastern part of Kanto Mountain1and,central Japan. Sci. Rep., Sailama Univ., Ser. B, vol.
2, no. I, p. 45-114, pis. 5-15.
Yamagiwa, N. (1962): The Permo-Carboniferous corals
from the Atetsu and the coral faunas of the same age
in the southwest Japan. Jour. Osaka Univ., Liberal
Arts, no. 10, p. 77-114, pis. 1-8.
Yokoyama, T. (1960): Permian corals from the Taishaku
district, Hiroshima Prefecture, Japan. Trans. Proc.
Palaeolll. Soc. Japan, N. S., no. 38, p. 239-248, pis.
27-28.

659

SYSTEMATIC INDEX(New Series No. 145-No. 152)

PALAEOZOOLOGY
Protozoa
Page
Hori, Rie: Some characteristic radiolarians from Lower Jurassic bedded cherts of the Inuyama area,
southwest Japan ......................................................................... 543
Kobayashi, Fumio: Late Paleozoic foraminifers of the Ogawadani Formation, southern Kwanto Mountains,
Japan ................................................................................. .435
Kojima, Satoru and Mizutani, Shinjiro: Triassic and Jurassic Radiolaria from the Nadanhada Range,
northeast China ......................................................................... 256
Matsusue, Kazuyuki: Some aberrant fusulinaceans from the Upper Carboniferous sequence of the
Akiyoshi Limestone Group, southwest Japan ............................................... 644
Niko, Shuji: Early Carboniferous Eostajjelfa (primitive Fusulinacea) from the Ichinotani Formation,
Fukuji district, central Japan .............................................................. 117
Sashida, Katsuo and Tonishi, Keiji: Additional note on Upper Permian radiolarian fauna from Itsukaichi,
western part of Tokyo Prefecture, central Japan ............................................ 523

PoliferaIgo, Hisayoshi, 19o, Hisaharu and Adachi, Shuko: Permian sphictozoan sponges from the Ichinotani
Formation, Hida Massif, central Japan .................................................... .453

Coelenterata
19o, Hisayoshi: A Permian rugose coral from Kamikagemori, Chichibu City, Saitama Prefecture, central
Japan .................................................................................. 656
Yamagiwa, Nobuo, Shiraishi, Kazuyuki and Yoshida, Mitsuo: Newly discovered Late Triassic hexacorals
from central Nepal ...................................................................... 654
Yoshida, Yasushi, Okimura, Yuji and Kato, Makoto: Early Carboniferous corals from the Omi Limestone,
central Japan ........................................................................... 228

Mollusca
Hayami, ltaru: Functional and taxonomic implications of internal ribs of Propeamussium ........... .476
Hayami, ltaru: A Tethyan bivalve, Posidonotis dainellii, from the Lower Jurassic of west Japan ....... 564
Kase, Tomoki: Early Carboniferous (Visean) gastropods from the Hikoroichi Formation of the Kitakami
Mountains, northeast Japan .............................................................. 361
Kondo, Yasuo: Burrowing depth of infaunal bivalves - Observation of living species and its rleation toshell morphology ........................................................................ 306
Kozai, Takeshi: Cretaceous Caestocorbula (Bivalvia) from Japan .................................. 324
Maeda, Haruyoshi: Taphonomy of ammonites from the Cretaceous Yezo Group in the Tappu area,
northwestern Hokkaido, Japan ............................................................ 285

660
Majima, Ryuichi and Takahashi, Hirokazu: Miocene molluscs from the Kozono Formation, Saitama
Prefecture, central Japan ................................................................. 246
Matsukawa, Masaki: Early shell morphology of Karsteniceras (Ancyloceratid) from the Lower Cretaceous
Choshi Group, Japan and its significance to the phylogeny of Cretaceous heteromorphammonites .............................................................................. 346
Matsukawa, Masaki: Barremian ammonites from the Ishido Formation, Japan - Supplements and faunalanalysis - ............................................................................. 396
Matsumoto, Tatsuro: Notes on Forbesiceras (Ammonidea) from Hokkaido (Studies of Cretaceousammonites from Hokkaido - LX) ........................................................ 16
Matsumoto, Tatsuro, Asai, Akito and Hirano, Hiromichi: Some inoceramids (Bivalvia) from the
Cenomanian (Cretaceous) of Japan - II. Three species from Hokkaido, well known abroad but
hitherto undescribed in Japan ............................................................. 146
Matsumoto, Tatsuro, Asai, Akito, Hirano, Hiromichi and Noda, Masayuki: Some inoceramids (Bivalvia)
from the Cenomanian (Cretaceous) of Japan - III. Three species occurring commonly in the
North-West Pacific region ................................................................ 378
Matsumoto, Tatsuro and Morozumi, Yoshiro: A sphenodiscid ammonite acquired rarely from theCretaceous of Japan .................................................................... .465
Matsumoto, Tatsuro and Saito, Rinji: Little known ammonite Grandidiericeras from Hokkaido (Studies
of Cretaceous ammonites from Hokkaido - LVIII) ........................................... 1
Matsumoto, Tatsuro and Tanaka, Keisaku: Some inoceramids (Bivalvia) from the Cenomanian (Cretaceous)of Japan - IV. An interesting new species from Hokkaido ................................... 570
Matsumoto, Tatsuro and Yokoi Katsujoh: Tiny ammonite Worthoceras from Hokkaido (Studies ofCretaceous ammonites from Hokkaido - LXI) ............................................. .42
Matsuoka, Keiji: Pliocene freshwater bivalves (Lamprotula and Cuineopsis: Unionidae) from the Iga
Formation, Mie Prefecture, central Japan ................................................. .417
Niko, Shuji and Hamada, Takashi: Adnatoceras from Middle Carboniferous of the Ichinotani Formation,
Fukuji district, central Japan .............................................................. 223
Niko, Shuji and Nishida, Tamio: Early Permian cephalopods from the Mizuyagadani Formation,
Fukuji district, central Japan ............................................................... 35
Niko, Shuji, Nishida, Tamio and Kyurna, Yuko: Middle Carboniferous Orthocerataceae andPseudorthocerataceae (Mollusca: Cephalopoda) from the Akiyoshi Limestone, Yamaguchi Prefecture(Molluscan paleontology of the Akiyoshi Limestone Group - VIII) ............................ 335
Noda, Masayuki: A note on Inoceramus tenuistriatus Nagao et Matsumoto (Bivalvia) from the Upper
Turonian (Cretaceous) of Japan ........................................................... 582
Tanabe, Kazushige and Shigeta, Yasunari: Ontogenetic shell variation and streamlining of some Cretaceousammonites .............................................................................. 165
Tanabe, Kazushige and Zushi. Yoshiki: Larval paleoecology of five bivalve species from the Upper
Pliocene of southwest Japan ............................................................. .491
Tashiro, Masayuki: Cretaceous Eomiodon and Costocyrena (Bivalvia) from southwest Japan ........... 91

Arthropoda
Kobayashi, Teiichi and Hamada, Takashi: The third addition to the Silurian trilobite fauna of
Yokokurayama, Shikoku, Japan ........................................................... 109
Kobayashi, Teiichi and Hamada, Takashi: On the Silurian trilobite faunule of Hitoegane near Fukuji
in the Hida Plateau, Japan ............................................................... 131
Tsukagoshi, Akira and lkeya, Noriyuki: The ostracod genus Cyl/lere O. F. Muller, 1785 and its
species ................................................................................. 197
Yajima, Michiko: Pleistocene ostracoda from the Atsumi Peninsula, central Japan ................... .49

661

Brachiopoda
Endo, Kazuyoshi: Life habit and relative growth of some laqueid brachiopods from Japan ........... 180
Tazawa, Jun-ichi: Attenuatella (Brachipoda) from the Upper Permian of northeast Japan and its bipolar
distribution ............................................................................. 276
Tazawa, Jun-ichi and Takaizumi Yukihiro: Poikilosakos (Oldhaminidina, Brachiopoda) from the Permianof northeast Japan ........................................................................ 10

Mammalia
Otsuka, Hiroyuki: Growth of antler in the subgenus Sika (Cervid, Mammal) from the Pleistocene
Formation in the Seto Inland Sea, west Japan .............................................. 625

PALAEOBOTANY
Kimura, Tatsuaki and Kim, Bong-Kyun: New taxa in the Late Triassic Daedong flora, south Korea ... 603
Kimura, Tatsuaki, Ohana, Tamiko and Tsuji, Masanori: Early Jurassic plants in Japan. Part 8.
Supplementary description and concluding remarks .......................................... 501
Niko, Shuji and Ikebe, Nobuo: Fisherites, a receptaculitid from the Middle Ordovician of Pyongyang
Coal-field ............................................................................... 371
Ozaki, Kimihiko: Tetracentron leaves from the Neogene of Japan ................................... 77

662

INDEX OF GENERA AND SPECIES
Notes: Words listed are names of genus and species-groups which are described, discussed or illustrated
in the articles in Transactions and Proceedings of Palaeontological Society of Japan, New Series, No. 145 to
No. 152. Words in bold-type are names of newly proposed taxa.

A

Adonatoceras ............................. 224
Adonatoceras ichinotaniensis ......... 225, Fig. 3
A kiyoshiphyllum sty/ophorum ........ 233, Fig. 7
Amblysiphonella hidensis ........... .458, Fig. 3
Anchistrocheles hondai .......... . 60, Figs. 5, II
A nchistrocheles yamaguchii .... 62, Figs. 5, 4, IIA nnulariopsis . ............................ 604
A nnulariopsis bunkeiensis ........... 605, Fig. I
Anomozamites mungyeongensis ...... 617, Fig. 6
A Ilenuatella .............................. 279
A ttenuatella bandoi .............. 281, Figs. 3, 4
B

Barremites (Barremites) strellOSlOma . . .403, Fig. 8
Bay/ea yvanii ...................... 364, Fig. 3
Beedeina higoensis ................. .439, Fig. 4
Bitaunioceras .............................. 37
Bitaunioceras undulatum ......... 37, Figs. 2-3
Brachycyc/oceras .......................... 337
Brachycycloceras akiyoshiensis ....... 337, Fig. 2
Brachycycloceras sp. indet .................. 338
Bumastus g/omerosus ............ 110, Figs. I, 2
Bythoceratina angulata ........... 66, Figs. 5, II
Bythocythere alata ........... 65, Figs. 7, II, 12
Bythocythere ishizakii ........ 66, Figs. 7, II, 12
C

Caestocorbula ............................ 326
Caestocorbula (s. I.) antiqua ......... 326, Fig. 4
Caestocorbula (s. I.) ohtai ........... 329, Fig. 3
Caestocorbula (? Parmicorbula) minima ........ .
.................................. . 327, Fig. 4

CaeslOcorbula (? Parmicorbula) shikamai . ...... .

Callistocythere sp ........................... 64
Capnodoce traversi ................. 260, Fig. 3
Carcinophyllum ........................... 237
Carcinophyllum hasegawai ...... 237, Figs. 9-10
Cerauroides orientalis ............ 110, Figs. I, 2
Cervus (Sika) greyi katokiyomasai ............. .
.............................. 632, Figs. 9-16

Cheirurus hitoeganensis ............. 138, Fig. 2
Cheirurid, gen. et sp. indet. ......... 140, Fig. 2

Chion kiusiuensis ......................... 320
Chiropteris coreanica ............... 611, Fig. 5
Circe scripta . ............................. 318
Cladophlebis l1lungyeongensis ........ 609, Fig. 4
Clisiophyllum kurohimense .......... 231, Fig. 4
Cobanocythere ? pulchra ........ 70, Figs. 10, 12
Coniproetus .............................. 135
Copicyntra akikawaensis ............ 530, Fig. 7
Coquimba ishizakii ................. 64, Fig. 12
Coquimba poga ................. 64, Figs. 6, II
Coreanophyllul1l variisegmenlUm ..... 619, Fig. 7
Costocyrene hojiensis ............. 98, Figs. 3, 5
Costocyrena matsumotoi ............. 95, Fig. 5
Costocyrene mijunensis .............. 98, Fig. 2
Costocyrena minor .................. 96, Fig. 2
Costocyrena ohnishii .............. 99, Figs. 2, 4
Costocyrena otsukai obsoleta ..... 102, Figs. I, 6
Costocyrena otsukai otsukai ......... 100, Fig. I
Costocyrena radiolOstriata ............ 93, Fig. 2
Costocyrena sp. cf. C. peikangensis ... 103, Fig. 5
Crenulilimopsis oblonga ................... 318
Cuneopsis ................................ 423
Cuneopsis (Cuneopsis) gracilenta ..... 423, Fig. 5
Cuneopsis (Procuneopsis) okuyamai .. 425, Fig. 5
Cuneopsis (Tchangsinaia) praemaxima ......... .

.................................. . 327, Fig. 4.................................. 427, Fig. 4

Caestocorbula (? Parmicorbula) monobensis .....

Cuspidaria macrorhynchus ................. 322
Cye/ina sinensis ........................... 319
Cystauletes kingi ............ .456, Figs. 2, 4-6
Cythere .................................. 198
Cythere alveoli valva ............. 204, Figs. 4-6.................................. 328, Fig. 3

Caestocorbu/a (? Parl1licorbu/a) obso/eta ....... .
................................... 330, Fig. 4

Calliphylloceras sp .................. 399, Fig. 3

663

Cythere boreokurila ....................... 206
Cythere cronini .............. 206, Figs. 5, 6, 10
Cythere golikovi ............. 206, Figs. 5, 6, 10

Forbesiceras largilliertianum .......... 17, Fig. 2Forbesiceras beaumontianum .......... 19, Fig. 2Forbesiceras mikasaeme .......... 21, Figs. 1-7

Cythere hanaii .................. 208, Figs. 5, 7
Cythere kamikoaniensis .......... 210, Figs. 5, 7
Cythere lutea ................. 211, Figs. 4, 5, 8
Cythere maia ............................. 211
Cythere nishinipponica ........ 211, Figs. 4, 5, 8
Cythere nopporoensis ......... 212, Figs. 5, 8, 10
Cythere omotenipponica ....... 214, Figs. 4, 5, 9
Cythere sanrikuensis ........... 214, Figs. 4, 5, 9
Cythere uranipponica .......... 216, Figs. 4, 5, 9
Cythere urupensis .............. 217, Figs. 5, 10
Cythere valentinei .............. 217, Figs. 5, 10
Cythere sp ................................ 218
Cytheroma ? sj:> ••.•..•••••..•••••••.•••••.• 69

Geisonoceras sejunctus .............. 338, Fig. 3
Geisonocerina ............................ 340
Geisonocerina cribrella .............. 340, Fig. 4
Gigi ..................................... 559
Gigi sp. aff. G. Justis .. ............. 559, Fig. 9
Gleichenites ? sp. A ............. 503, Figs. 3, 18
Glycymeris rotunda ....................... 317
Gomphina veneriformis nelanaegis .......... 319
Grandidiericeras ............................ 1
Grandidiericeras nagaoi ............ 2, Figs. 1-4
Guexella nudata .................... 260, Fig. 4oH

Decoroscutellum (?) sp. indet ............... 133
Dicranopeltis tricomis ........... 115, Figs. 1, 2
Dictyophyllum exile ................. 606, Fig. 2
Dictyophyllum kotakiense .......... 504, Fig. 11
Dosinorbis japonicus ...................... 319

Hanaicythere .............................. 72
Hanaicythere nipponica ......... 74, Figs. 10, 12
Hausmannia (Protorhipis) ussuriensis .......... .
................................... 607, Fig. 3
Heteroceras (Argvethites) sp ..... .402, Figs. 3, 6
Hiroshimaphyllum simplex . ......... 239, Fig. 10

E
Encrinurus ............................... 140
Encrinurus Jimbriatus ........... 140, Figs. 2-4
Encrinurus cf. kitakamiensis ......... 143, Fig. 3
Encrinurus cf. similis ............... 144, Fig. 3
Encrinurus subtrigonalis ............. 110. Fig. 3
Entactinosphaera brevispinosa ........ 529, Fig. 7
Entactinosphaera sp ................. 529, Fig. 7
Eomiodon ................................ 104
EOllliodon ? kumamotoensis ................. 93
Eomiodon matsubasensis ............. 92, Fig.
Eomiodon matsumotoi ............... 92, Fig. 1
Eomiodon sakawanus ................ 91, Fig. 1
EostaJJella excavata ................. 120, Fig. 3
EostaJJella igoi ..................... 121, Fig. 4
EostaJJella kanmerai ................ 123, Fig. 4
EsotaJJella mosquensis .............. 123, Fig. 5
EostaJJella subulba ................. 126, Fig. 5
EostaJJella tenebrosa ............ 126, Figs. 4, 5
EostaJJella cf. postmosquensis ....... 125, Fig. 3
EostaJJella sp ...................... 127, Fig. 4
Eucyrtidiellum ptyctum .............. 260, Fig. 4
Eucyrtidiellum unumaense ........... 260, Fig. 4
F

Fisherites coreanicus ............. 373, Figs. 2, 3
Forbesiceras ............................... 17

G

Inoceramus . .............................. 147
Inoceramus ginterensis ........ 388, Figs. 5, 7-9
Inoceramus grandilis ............. 155, Figs. 4, 5
Inoceramus nodai ........ 571, Figs. 1-3, 5-13
Inoceramus pennatulus .......... 379, Figs. 1-3
Inoceramus reduncus ............ 383, Figs. 4-6
Inoceramus tenuis .................. 148, Fig. 1
Inoceramus tenuistriatus ......... 584, Figs. 5-9Inoceramus virgatus ............. 157, Figs. 6-8
J

Japonoscutellum tumidum ........... 109, Fig. 3
K

Karsteniceras ......................... 348, 399
Karsteniceras asiaticum .......... 399, Figs. 3, 4
Karstenicera obarai ............. 349, Figs. 1-4
Kars/enicera oba/ai .............. 401, Figs. 3, 5Katroma ................................. 551
Katroma kurusuensis ................ 553, Fig. 6
Katroma sp. N ..................... 554, Fig. 7
Kawanamia ............................... 366
Kawanamia onimaruensis ......... 366, Figs. 3, 4Kosovope/tis hidensis ............... 133, Fig. 1
Kosovopeltis hidensis forma striatum .133, Fig. 1

664

L

Lamprotula ............................. .420
Lamprotula sp. . ................... .423, Fig. 4
Lamprotula (Sinolamprotula) sp. A .. .420, Fig. 2
Lamprotula (Parunio) sp. B ........ .421, Fig. 3
Laqueus rubellus .......................... 181
Laternula Iimicola ......................... 322
Libycoceras awajiense ........... .466, Figs. 1, 2
Limopsis tajimae .......................... 317
Littorinides sp ...................... 367, Fig. 3
Livarella validus .......................... 260
M

Macoma con tabulata ...................... 321
Macoma incongrua ........................ 321
Mactra chinensis .......................... 320
Mactra veneriformis ....................... 320
Meretrix petechialis ........................ 318
Metaleiolichas ............................ 113
Metaleiolichas tuberculatus ....... 113, Figs. 1, 3N

Nemocardium samarangae ................. 318
Neoalbaillella pseudogrypus .......... 527, Fig. 9
Neonesidea hanaii ............... 60, Figs. 5, 11
Neoschwagerina craticulifera ..... .445, Figs. 6, 7
Neoschwagerina haydeni ............ .445, Fig. 7
Neoschwagerina simplex ............ .447, Fig. 6
Neoschwagerina sp. A ............. .449, Fig. 6
Nilssonia sp. B ................ 509, Figs. 17, 20
Nuttallia olivacea ......................... 321

oOblimopa japonica ........................ 318
Octatormentum 1 floriferum ......... 533, Fig. 5
Octatormentum 1 sp ....................... 536
Otozamites micrauritus .............. 621, Fig. 8
Otozamites sp. C ............... 511, Figs. 6,14

P
Palaeoreichelina ........................... 649
Palaeoreichelina disvoluta ........... 651, Fig. 4Palaeoreichelina 1 sp. . .............. 651, Fig. 4
Panopea japonica ......................... 321
Paracypris 1 sp ...................... 63, Fig. 6
Paracytheridea sp ................... 68, Fig. 12
Paracytherom sp. . ......................... 69
Paradoxostoma setosum .............. 68, Fig. 9
Paradoxostoma sp ................... 68, Fig. 8"Parafusulina" gigantea ........... .439, Fig. 4

Parafusulina hirayuensis ............ .441, Fig. 5
Paraleiolichas ............................. 113
Paraleiolichas globulus .......... 113, Figs. 1-3
Parahsuum (1) sp. A ................ 260, Fig. 3
Parunio ................................. .421
Pentactinocarpus sp. A .............. 261, Fig. 2
Phlebopteris sp. A .............. 503, Figs. 1, 10
Poikilosakos ............................... 11
Poikilosakos kamiyassensis ........... 12, Fig. 3
Posidonotis dainelli ................. 565, Fig. 1
Poulpus curvispinus ................. 261, Fig. 2
Procuneopsis ............................ .425
Propontocypris sp .......................... 64
Proetus .................................. 135
Proetus (Coniproetus) tenuiceps ... 136, Figs. 1, 3
Proetus (Coniproetus 1) subconicus ... 138, Fig. 1
Pseudoctenis nipponica ......... 509, Figs. 9, 16
Pseudoctenis 1 sp .............. 513, Figs. 5, 12
Pseudocythere sp. 1 .................. 67, Fig. 7
Pseudocythere sp. 2 .................. 67, Fig. 7
Pseudodoliolina pseudolepida ....... .443, Fig. 6
Pseudoorthoceratid, gen. et sp. indet .. 39, Fig. 3
Pseudostylosphaera ........................ 261
Pseudostylosphaera japonica ......... 261, Fig. 2
Pseudostylosphaera sp. cf. P. hellenica ........ .
.................................. 261, Fig. 2
Pseudostylosphaera sp. cf. P. tenue .. 263, Fig. 2
Pseudostylosphaera sp. A ........... 263, Fig. 2
Pseudostylosphaera sp. B ........... 263, Fig. 2
Pseudostylosphaera sp. C........... 263, Fig. 2
Pseudostylosphaera sp. D ........... 263, Fig. 2
Pseudostylosphaera sp. E . . . . . . . . . .. 263, Fig. 2
Pseudo<.jJgopleura (Stephano<.jJga) nishimurai ....
................................... 367, Fig. 5
Pterophyllum micraequale ........... 621, Fig. 9
Pterophyllum 1 sp .............. 509, Figs. 7, 15
Ptilophyllum shinadaniense ......... 507, Fig. 19
Ptilophyllum sp. cf. P. hsiangshanense ......... .
............................... 507, Figs. 8, 13
Ptilophyllum sp. A ........................ 507
R

Rectomillerella ............................ 648
Rectomillerella okuboensis .......... 648, Fig. 4
Rectomillerella sp. A ............... 648, Fig. 4
Rectomillerella sp. B ............... 649, Fig. 4
Reticycloceras (1) sp. indet .......... 343, Fig. 2
Ruditapes philippinarum ................... 320

sSclerochilus sp. . .................... 68, Fig. 9

665

Semicytherura sp. I ................. 68, Fig. 12
Semicytherura sp. 2 ................. 68, Fig. 12
Sinolamprotula .......................... .420
Solen strict us ............................. 321
Sphaerexochus sp. indet ............. 140, Fig. 2
Sphenopteris sp. G .................. 511, Fig. 4
Sphenopleris sp. H ................. 504, Fig. 2
Slichocapsa convexa ................ 263, Fig. 4
Straparollus (Euomphalus) asanoi ., .. 362, Fig. 2
Siraparollus (Euomphalus) sp ........ 364, Fig. 3
Siriacoceras (?) sp. indet. ........... 343, Fig. 2
Syringocapsa coliforme ... " ......... 556, Fig. 8
T

Taleiwaia laleiwai .................. 249, Fig. 3
Taleiwaia yamanarii ................ 25 I, Fig. 3
Tchangsinaia ............................ .426
Tegillarca granosa ......................... 3 17
Telracenlron masuzawaense ..... 78, Figs. 2, 3, 4
Telracenlron sp ...................... 8 I, Fig. 4
Tetratormentum aeutum ........ 532, Figs. 3, 10
Tetratormentum globiforme .. , ...... 533, Fig. 4
Thal11nasleria (Aslraeoll1orpha ?) lIIajor ....... .
................................. . 655, Fig. 1"Thecoslllilia" sp. A ............... 655, Fig. 1"Thecoslllilia" sp. B ................ 655, Fig. 1
Trachyleberis sp. . .......................... 64
Triaenosphaera ........................... 530
Triaenosphaera minutus ............. 530, Fig. 8
Triassocampe deweveri .............. 265, Fig. 3
Triassocampe sp. A ................. 265, Fig. 3

Tricolocapsa (7) JusiJormis .......... 265, Fig. 4
Tricolocapsa plicarum ............... 265, Fig. 4
Tricolocapsa lelragona .............. 267, Fig. 4
Tricolocapsa sp. cf. T. ruesli ........ 267, Fig. 4
Triplanospongos .......................... 536
Triplanospongos musashiensis ........ 536, Fig. 6

uUberinterna .............................. 521
Uberinterna virgispinosum ........ 531, Figs. 2, 8v
Valdedorsella (Pliezalpella) sp ....... .403, Fig. 3
Verbeekina verbeeki ............... .443, Fig. 5
Vicaryella allikoae .................. 251, Fig. 3
Vicaryella ishiiana .................. 253, Fig. 3w
Worlhoceras .............................. .43
Worthoeeras pacificum .......... .43, Figs. 1-4

xXiphichillis sp ....................... 69, Fig. 8yYeharaia annlllala .................. 267, Fig. 3
Yeharaia japonica .................. 267, Fig.
Yeharaia sp. A ..................... 267, Fig. 3
Yokoyalllaella (Yokoyalllaella) lerlioseptata ..... .
.................................. 657, Fig. 2

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 145- 152
1987 - 1988

Pala eo nt o logical Soc iet y of Japan

The heading in Japanese commemorates the handwriting of Prof. Matajiro YOKOYAMA, father of Japanesepalaeontology, who was a professor of stratigraphy and palaeontology at the Geological Institute, Imperial
University of Tokyo.
The fossil on the cover is Vicarya yokoyamai Takeyama, an Early Middle Miocene gastropod
from the Kurosedani Formation at Kakehata, Yatsuo-cho, Nei-gun, Toyama Prefecture,
central Japan (Collected by T. Kotaka and K. Ogasawara, IGPS No. 99075, photo by S.
Ohtomo and Y. Kikuchi, x 0.9).

CONTENTS
Number 145 (published April 30, 1987)
Transactions
ArticleP age

Figures

1-9

1-5

10-15

1-3

16-31

1-7

825. Matsumoto, Tatsuro and Saito, Rinji: Little known ammonites
Grandidiericeras from Hokkaido (Studies of Cretaceous ammonites
from Hokkaido - LVIII) ................................... .
826. Tazawa, Jun-ichi and Takaizumi, Yukihiro: Poikilosakos(Oldhaminidina, Brachiopoda) from the Permian of northeast

Japan .................................................... .
827. Matsumoto, Tatsuro: Notes on Forbesiceras (Ammonidea) from

Hokkaido (Studies of Cretaceous ammonites from Hokkaido LX) .................................................... .
Proceedings .........................................

32-34

Number 146 (published July 30, 1987)
Transactions
828. Niko, Shuji and Nishida, Tamio: Early Permian cephlopods from theMizuyagadani Formation, Fukuji district, central Japan ......... .

35-41

1-3

829. Matsumoto, Tatsuro and Yokoi, Katsujoh: Tiny ammonite Worthoceras
from Hokkaido (Studies of Cretaceous ammonites from Hokkaido- LXI) ....................................................

42-48

1-4

830. Yajima, Michiko: Pleistocene ostracoda from the Atsumi Peninsula,central Japan.............................. . ............... .
831. Ozaki, Kimihiro: Tetracentron leaves from the Neogene of

49-76

1-12

Japan ..................................................... .

77-87

1-4

Proceedings ....................................... .

88-89

Number 147 (published October 30, 1987)
Transactions
832. Tashiro, Masayuki: Cretaceous Eomiodon and Costocyrena (Bivalvia)
from southwest Japan....................................... 91-108
833. Kobayashi, Teiichi and Hamada, Takashi: The third addition to the
Silurian trilobite fauna of Yokokurayama, Shikoku, Japan ........ 109-116
834. Niko, Shuji: Early Carboniferous Eostajjella (primitive Fusulinacea)
from the Ichinotani Formation, Fukuji district, central Japan ..... 1l7-130

1-7
1-3
1-5

II

835. Kobayashi, Teiichi and Hamada, Takashi: On the Silurian trilobite
faunule of Hitoegane near Fukuji in the Hida Plateau, Japan ..... 131-145
836. Matsumoto, Tatsuro, Asai, Akito and Hirano, Hiromichi: Some
inoceramids (Bivalvia) from the Cenomanian (Cretaceous) of JapanII. Three species from Hokkaido, well known abroad but hitherto
undescribed in Japan ........................................ 146-164
837. Tanabe, Kazushige and Shigeta, Yasunari: Ontogenetic shell variationand streamlining of some Creta~eous ammonites ................. 165-179
838. Endo, Kazuyoshi: Life habit and relative growth of some laqueid
brachiopods from Japan ..................................... 180-194
Proceedings ........................................ 195-196

1-5

1-8
1-8
1-12

Number 148 (published December 30, 1987)
Transactions
840. Tsukagoshi, Akira and Ikeya, Noriyuki: The ostracod genus Cythere
O. F. Miiller, 1785 and its species ............................. 197-222
841. Niko, Shuji and Hamada, Takashi: Adnatoceras from Middle
Carboniferous of the Ichinotani Formation, Fukuji district, central
Japan ..................................................... 223-227
842. Yoshida, Yasushi, Okimura, Yuji and Kato, Makoto: Early
Carboniferous corals from the Omi Limestone, central Japan ..... 228-245
843. Majima, Ryuichi and Takahashi, Hirokazu: Miocene molluscs from theKozono Formation, Saitama Prefecture, central Japan ............ 246-255
844. Kojima, Satom and Mizutani, Shinjiro: Triassic and Jurassic Radiolaria
from the Nadanhada Range, northeast China ................... 256-275
845. Tazawa, Jun-ichi: Attenuatella (Brachipoda) from the Upper Permian ofnortheast Japan and its bipolar distribution ..................... 276-284
846. Maeda, Haruyoshi: Taphonomy 01 ammonites from the Cretaceous
Yezo Group in the Tappu area, northwestern Hokkaido, Japan .... 285-305
847. Kondo, Yasuo: Burrowing depth of in faunal bivalves - Observationof living species and its relation to shell morphology .............. 306-323
848. Kozai, Takeshi: Cretaceous Caestocorbula (Bivalvia) from Japan .. 324-334
849. Niko, Shuji, Nishida, Tamio and Kyuma, Yuko: Middle Carboniferous
Orthocerataceae and Pseudorthocerataceae (Mollusca: Cephalopoda)
from the Akiyoshi Limestone, Yamaguchi Prefecture (Molluscan
paleontology of the Akiyoshi Limestone Group - VIII) .......... 335-345
850. Matsukawa, Masaki: Early shell morphology of Karsteniceras(Ancyloceratid) from the Lower Cretaceous Choshi Group, Japan andits significance to the phylogeny of Cretaceous heteromorphammonites ................................................. 346-359

1-10

1-3
1-11
1-4
1-5
1-4
1-15
1-9
1-4

1-4

1-4

iii

Number 149 (published April 30, 1988)
Transactions
851. Kase, Tomoki: Early Carboniferous (Visean) gastropods from the
Hikoroichi Formation of the Kitakami Mountains, northeast
852.
853.

854.
855.

Japan ..................................................... 361-370
Niko, Shuji and Ikebe, Nobuo: Fisherites, a receptaculitid from theMiddle Ordovician of Pyongyang Coal-field ..................... 371-377
Matsumoto, Tatsuro, Asai, Akito, Hirano, Hiromichi and Noda
Masayuki: Some inoceramids (Bivalvia) from the Cenomanian(Cretaceous) of Japan - III, Three species occurring commonly in the
North-West Pacific region .................................... 378-395
Matsukawa, Masaki: Barremian ammonites from the Ishido Formation,
Japan - Supplements and faunal analysis - ................... 396-416
Matsuoka, Keiji: Pliocene freshwater bivalves (Lamprotula andCuineopsis: Unionidae) from the Iga Formation, Mie Prefecture,
central Japan ............................................... 417-429
Proceedings ........................................ 430-433

1-5
1-3

1-9
1-11

1-5

Number 150 (published June 30, 1988)
Transactions
856. Kobayashi, Fumio: Late Paleozoic foraminifers of the Ogawadani
Formation, southern Kwanto Mountains, Japan ................. 435-452
857. Igo, Hisayoshi, Igo, Hisaharu and Adachi, Shuko: Permian sphictozoan sponges from the Ichinotani Formation, Hida Massif,

central Japan ............................................... 453-464

1-7

1-7

858. Matsumoto, Tatsuro and Morozumi, Yoshiro: A sphenodiscidammonite acquired rarely from the Cretaceous of Japan .......... 465-475
859. Hayami, Haru: Functional and taxonomic implications of internal ribsof Propeamussium .......................................... 476-490
860. Tanabe, Kazushige and Zushi, Yoshiki: Larval paleoecology of five
bivalve species from the Upper Pliocene of southwest Japan ...... 491-500

1-4
1-7
1-3

Number 151 (published October 30, 1988)
Transactions
861. Kimura, Tatsuaki, Ohana, Tamiko and Tsuji, Masanori: Early Jurassic
plants in Japan. Part 8. Supplementary description and concluding
remarks .................................................... 501-622

1-21

iv

862. Sashida, Katsuo and Tonishi, Keiji: Additional note on Upper Permian
radiolarian fauna from Hsukaichi, western part of Tokyo Prefecture,
central Japan ............................................... 523-542 1-10
863. Hori, Rie: Some characteristic radiolarians from Lower Jurassic
bedded cherts of the Inuyama area, southwest Japan ............. 543-563
1-9864. Hayami, Haru: A Tethyan bivalve, Posidonotis dainellii from the Lower
Jurassic of west Japan ....................................... 564-569
865. Matsumoto, Tatsuro and Tanaka, Keisuku: Some inoceramids (Bivalvia)
from the Cenomanian (Cretaceous) of Japan - IV. An interesting new
species from Hokkaido ....................................... 570-581 1-15
866. Noda, Masayuki: A note on Inoceramus tenuistriatus Nagao etMatsumoto (Bivalvia) from the Upper Turonian (Cretaceous) of Japan 582-600 1-10
Proceedings ........................................ 601-602
Number 152 (published December 30, 1988)
Transactions
867. Kimura, Tatsuaki and Kim, Bong-Kyun: New taxa in the Late Triassic
Daedong flora, south Korea .................................. 603-624
868. Otsuka, Hiroyuki: Growth of antler in the subgenus Sika (Cervid, Mammal) from the Pleistocene Formation in the Seto Inland Sea,
west Japan ................................................. 625-643
869. Matsusue, Kazuyuki: Some aberrant fusulinaceans from the Upper
Carboniferous sequence of the Akiyoshi Limestone Group, southwest
Japan ..................................................... 644-653

1-9

1-16

1-4

Short Notes
21.
22.

Yamagiwa, Nobuo, Shiraishi, Kazuyuki and Yoshida, Mitsuo: Newly
discovered Late Triassic hexacorals from central Nepal ........... 654-655
Igo, Hisayoshi: A Permian rugose coral from Kamikagemori, Chichibu
City, Saitama Prefecture, central Japan ........................ 656-658
Systematic Index .................................... 659-661
Index of Genera and Species .......................... 662-665

1-2

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

New Series No. 152

December 30, 1988

CONTENTS

TRANSACTIONS
867. KIMURA, Tatsuaki and KIM, Bong-Kyun: New taxa in the Late Triassic Daedong
flora, south Korea ....................................................... 603
868. OTSUKA, Hiroyuki: Growth of antler in the subgenus Sika (Cervid, Mammal) fromthe Pleistocene Formation in the Seto Inland Sea, west Japan .................. 625
869. MATSUSUE, Kazuyuki: Some aberrant fusulinaceans from the Upper Carboniferous
sequence of the Akiyoshi Limestone Group, southwest Japan .................. 644
SHORT NOTES
21.

YAMAOIW A, Nobuo, SHIRAISHI, Kazuyuki and YOSHIDA, Mitsuo: Newly discovered

22.

100, Hisayoshi: A Permian rugose coral from Kamikagemori, Chichibu City,

Late Triassic hexacorals from central Nepal ................................. 654
Saitama Prefecture, central Japan .......................................... 656
SYSTEMATIC INDEX ........................................................ 659
INDEX OF GENERA AND SPECIES .......................................... 662