ISSN

0031-0204

fJ -lf~~i.~~~.~·;~fTransactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 124

Palaeontological Society of Japan
December 30, 1981

Itaru HAY AMI and Ikuwo OBA TA
Co-Editors
Associate editors Teruya UYENO and Tomowo OZAWA

Officers for 1981-1982
President:

Tetsuro HANAI

Councillors:

Kazuo ASAMA, Kiyotaka CHINZEI, Takashi HAMADA, Tetsuro HANAI,

Yoshikazu HASEGAWA, Itaru HAYAMI, Hisayoshi IGO, Junji ITOIGAWA, Tadao
KAMEl, Tatsuaki KIMURA, Ikuwo OBATA, Tsunemasa SAITO, Tsugio SHUTO,
Yokichi TAKAYANAGI, Toshimasa TANAIMembers of Standing Committee:

Kazuo ASAMA (Finance), Kiyotaka CHINZEI (General

Affairs), Takashi HAMADA (Membership), Itaru HA YAMI (Transactions), Hisayoshi IGO (Foreign Affairs),

Tatsuaki KIMURA (Planning),

Ikuwo

OBA T A

(Transactions), Tsugio SHUTO (Special Papers), Yokichi TAKA YANAGI ("Fossils")
Secretaries:

Kazuhiko UEMURA (Finance), Toshiyuki YAMAGUCHI (General Affairs),

Teruya UYENO and Tomowo OZAvVA (Transactions), Hisaharu IGO (Planning),
Ienori FUJIYAMA and Juichi YANAGIDA (Special Papers), Kunihiro ISHIZAI<l("Fossils")
Auditor:Kenji KURIHARA

The fossil on the cover is an adult example (1'. TAKAHASHI colI.) of

Mikasaites orbicularis MATSUMOTO (subfamily Marshallitinae, family
Kossmaticeratidae) from the Lower Cenomanian (Cretaceous) of theMikasa area, central Hokkaido. (photo by M. NODA, natural size)

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

Trans. Proc. Palaeont. Soc. japan, N. S., No. 124, pp. 187--207, pIs. 30-32, December 30, 1981

737.

EARLY JURASSIC PLANTS IN JAPAN.

PART 3*

T ATSUAKI KIMURA AND MASANORI TSUJIITokyo Gakugei University, Koganei, Tokyo 184

Abstract. This is the third of our serial papers on the early jurassic plants injapan. In this paper we describe Phlebopteris sp. A (Matoniaceae), Coniopteris
neiridaniensis sp. nov. (Dicksoniaceae), Clathropteris meniscoides, Dictyophyllum
Iwtakiense sp. nov., Thaumatopteris elongata, Hausmannia (Protorhipis) sp. A (Dipteridaceae) and the following sterile fern leaves: Cfr. Asterotheca oka/ujii, Cfr. Todites/ukutomii, Cfr. T. yamanoiensis, Sphenopteris sp. A, Cladophlebis kurumensis sp.
nov., C. pseudodelicatula, C. raciborskii, C. sp. A, C. sp. Band C. sp. C.

Systematic description
Introduction andacknowledgements

Family Matoniaceae
Genus Phlebopteris BRONGNIART,

In continuation of our previous papers(KIMURA and TsuJII, 1980a, b), we here
describe the fern leaves belonging to thefamilies Matoniaceae, Dicksoniaceae andDipteridaceae, and also other sterile fern
leaves from the Lower Jurassic Kuruma
Group and Iwamuro Formation. The
details of the stratigraphy of plantbearing formations and fossil localities
were given in our first paper (KIMURA
and Tsum, 1980a).
We express our sincere gratitude toProfessor Emeritus Thomas M. HARRIS,
F. R. S. of the University of Reading forhis helpful suggestions and critical readingover the present manuscript. Our thanks
are extended to Mr. Gumpei MORI, Mr.
Noriyuki SASAKI and Mr. Akihiro KOBAYASHI who helped us in collecting thefossil plants.

1836: 371
Phlebopteris sp. A
PI. 30, Fig. 1; Text-figs. la, bMaterial: NE-046, 123, 134, 146 (Neiridani).

Description.-Four detached pinnules
were obtained. They are long andnarrow, nearly parallel-sided, more than
5.5 cm long and 0.3-0.4 cm wide. The
midnerve is distinct but lateral veins areinvisible. Sori are on both side of themidnerve, 500 flm in diameter, consistingof 6 sporangia of about 200 flm in diameter around a placenta, about 75 flmin diameter. Details of sporangia areuncertain.
Distribution and occurrence.-Phlebopteris
sp. is rare and known only from the
Negoya Formation.
Remar!?s.-Our pinnules are ill-preserved
and specifically indeterminable. But they
remind us of those of Phlebopteris
polypodioides BRONGNIART and Matonia* Received june 13, 1981; Read October 11,
1980 at Toyama.

187

Tatsuaki KnvlURA and Masanori 'l'SUJII

188---------

1--=:::=======-=--=-=--::!::.~;c-::-.~:::~~ a..

~~~._~~~_~~~====~lb!:! ""• ....•.....
~
2k

,

·fII

2a

2f

If

21

2i

2h

~

II~

l~

r

~2m
O.Scm
L---...J2b2d

Text-figs. 1-2 (All natural size, unless otherwise indicated).
1. Phlebopteris sp. A. 1a: An incompletely preserved fertile pinnule, open circles indicate theposition of placenta (NE-146). 1b: Fertile pinnule (NE-123). In both specimens lateralveins are invisible.
2. Coniopteris neiridaniensis KI~mRA and TSUJII sp. nov. Sterile leaves (2a-f). 2a: Thelargest specimen bearing large-sized pinnules with deeply lobed margins (NNW -293, holotype; PI. 30, Fig. 2). 2b: A penultimate pinna with medium-sized pinnules (NEG-120,
paratype; PI. 30, Fig. 4). 2c: An ultimate pinna (fertile?; sori invisible) with large-sizedpinnules (NNW-590, paratype; PI. 30, Fig. 3). 2d: Ditto (NNW -777, paratype). 2e: A pinna
with medium-sized pinnules (NNW-671, paratype). 2f: Ditto (DAI-055, paratype; PI. 30,
Fig. 5). Fertile leaves (2g-m). 2g: (A-0480; sori not drawn). 2h: (NNW -005, paratype).
2i: (NNW -001, para type). 2j: A penultimate pinna, with small-sized sterile pinnules withentire margins (NNW-765, paratype). 2k: (NNW-561, paratype; PI. 30, Fig. 6). 21: (NE053, paratype; PI. 30, Fig. 6). 2m: A sorus and venation drawn from 2i.

braunii (GOEPPERT) (see HARRIS, 1980).
Family Dicksoniaceae
Genus Coniopteris BRONGNIART,
1849: 26
Dicksoniaceous ferns are rather rare in
the Lower Jurassic plant-beds of Japanand Coniopteris neiridaniensis is the only
one known.

Coniopteris neiridaniensis KIMURA
and TsuJI! sp. nov.
PI. 30, Figs. 2-7; Text-figs. 2a-m

Material: Holotype; NNW-292 (NishiNeiridani). Para types ; NNW-590, 671, 777,
NEG-120 (Negoya), DAI-055 (Dairagawa)(sterile leaves), NNW-001, 005, 561, 765, NE053 (Neiridani) (fertile leaves). Other specimens; NNW -293 and other 16 specimens, NEE307 (Higashi-Neiridani), NE-157, 160, DAI071, NEG-041, 110, 262, A-0480, 75064, 7207021(lwamuro). Stratum typicum: Negoya For-

737. Early Jurassic plants
mation, Kuruma Group. Locus typicus : Nishi·
Neiridani, a branch of the Dairagawa, Asahi·
machi, Shimo·Niikawa·gun, Toyama Prefecture.
Derivatio nominis: After the Neiridani Valley
where the hoiotype and most of paratypes
were collected.

Diagnosis.-Leaves at least tnpmnate,partly sterile and partly fertile. Rachis
rather slender, up to 2 mm wide, smooth,grooved above and rounded below. (Wholeleaf unknown.) Penultimate and ultimate
pinnae elongate-ovate in outline, attachedto the slender pinna rachis an angle of55-60 degrees. Pinnules katadromic inorder, varied in size and manner oflobing in different part of the leaf.
Pinnules elongate-rhomboid or elongateoblong in outline, asymmetrical, apex
obtuse or rounded; lamina decurrent toform a web along the pinna rachis.
Large-sized pinnules typically 1.5 cm longand 3.5 mm wide, deeply dissected into
lobes; lobes 5-6 pairs, directed forwards
and with rounded apex. Medium-sizedpinnules typically 0.5-0.7 cm long and 1.52.5 mm wide, and with shallowly lobed or
undulated margins; lobes or undulations
4 pairs, apices rounded directed forwards.
Basal pinnules not specialized. Smallsized pinnules elongate-triangular or rectangular, entire, 3 mm long and 0.5 mm
wide, apex obtusely pointed or rounded.
Venation appearing sympodially branched;lateral veins corresponding to lobes orundulations. Lateral veins generally simple in medium- and small-sized pinnules
but once forked in large pinnules.
Fertility varying from fully fertile tooccasionally fertile; fully fertile lamina
more or less reduced. Sori borne simply
on the apices of lobes, predominantly on
the apices of acroscopic lobes, semicircular, typically 550 p.m in diameter.
(Details of sorus unknown.)
Distribution and occurrence.-Coniopteris
neiridaniensis occurs rarely in the Negoya

189

Formation of Kuruma Group and the
Iwamuro Formation.
Discussion and comparison. - Unfortunately it is not clear whether the indusiumof Coniopteris neiridaniensis is cup-shapedor bivalvate.
Coniopteris neiridaniensis is characterized
by its sterile pinnules, especially by its
large- and medium-sized pinnules deeply
or shallowly divided into lobes with arounded apex and each lobe with asimple or once forked lateral vein.
Coniopteris neiridaniensis differs fromC. hymenophylloides (BRONGNIART) as
emended by HARRIS (1961) in its basal
pinnules. In Coniopteris hymenophylloides
the first basiscopic lobe of the basal
basiscopic pinnule develops as filiform
processes lying over the rachis, and thatof the opposite pinnule also lies over therachis. Instead, Coniopteris neiridaniensis
has unspecialized basal lobes.
Among the leaves having been regarded
as Dicksonia, Thyrsopteris and Coniopteris
hymenophylloides by the previous authors
from Japan, Korea and China, the following are rather like Coniopteris neiridaniensis:
1) The following leaves described by
YOKOYAMA (1889) from the Lower
Cretaceous Oguchi Formation. They
were later determined by OISHI (1940)
as Coniopteris hymenoplzylloides:
Dicksonia nephrocarpa BUNBURY; p.
25, pI. 1, figs. 1, 1a (non 1b).
Thyrsopteris prisca EICHWALD; p. 23,
pI. 1, figs. 3, 3a, 4: His sterilepinnules are similar in form to ours,
but their lareral veins are twice or
thrice forking in his fig. 3a.
T. murrayana BR01'GNIART; p. 22, pI.
12, fig. 5.
2) A sterile leaf described by YOKOYAMA (1906) from the Jurassic of
Fang-zi, Shandong, China as Coniopteris
hymenophylloides (p. 24, pI. 6, fig. 3),

190

Tatsuaki KIMURA and Masanori TsuJI!

venation not known.
3) Sterile leaves described by KAWASAKI
from the Daedong Group, Korea as
Coniopteris hymenophylloides (1925, p.
7, pI. 6, fig. 23; pI. 13, fig. 44, 44a:
1926, p. 14, pI. 4, fig. 13: 1939, p. 22,
pI. 2, fig. 11 ?).
KRASSILOV (1978) placed the weIl-known
species Coniopteris burejensis (ZALESSKY)SEW ARD in the genus Dicksonia, because
he found that their indusia were not cupshaped but bivalvate. He revived HEER'sname Dicksonia concinna for the leaves
formerly regarded as C. burejensis in the
Bureja Basin, because of its priority.
According to him, HEER's Adiantites
nymplzarum is con specific with Dicksonia
concin/la HEER.

Dicksonia concinna ('Coniopteris burejensis') is distinguished from C. neiridaniensis by its acutely pointed lobes.
Sterile pinnules of Coniopteris vachrameevii originaIlydescribed by VASSILEVSKAJA(1967) are similar in form to our largesized ones. But Coniopteris vachrameevii
is distinguished from ours by its lobes
with further minutely lobed apex.
CHOW and YEH (in SZE et aI., 1963)
instituted Coniopteris szeiana and included
SZE'S Coniopteris burejensis (1931, p. 43,
pI. 7, figs. 5-8) and C. sp. (? n. sp.)
(1933b, p. 79, pI. 11, figs. 4-13, 18, 19) intheir C. szeiana. Coniopteris szeiana ischaracterized by its pinnules deeply dissected into lobes with rounded apex.
However, this Chinese early Jurassic species is distinguished from ours by its
roundly rhomboid pinnules with 2-3 pairsof lobes, instead of elongate-rhomboid ones
with 4-6 pairs of lobes in ours.
The following leaves appear to berather similar in form to Coniopterisszeiana than to C. hymenophylloides andC. neiridaniensis : Thyrsopteris prisca HEER ;of KRASSER (1906) from the Jurassic of
NE-China. Coniopteris hymenophylloides(BRONGNIART) ; of Y ABE and OISHI (1933)
from the Jurassic of NE-China. Coniopteris sp.; of HE et ai. (1979) from theJurassic of Qinghai, NW-China.
Coniopteris embensis originaIly describedby PRYNADA (1938) from the Jurassic of
Emba Basin and later by BARANOV A etai. (1975) from the Jurassic of eastern
Caspian Basin shows the pinnules dissected
into lobes with rounded apex. But it
differs from ours in its smaIler numberof lobes (2-3 pairs) in each pinnule.
Coniopteris spectabilis BRICK illustrated
by SIXTEL and KHUDAlBERDYEV (1968, pI.
20; pI. 23, fig. 3) from the Middle Jurassicof Middle Asia resembles our C. neiridaniensis, but is also distinguished from ours
by its pinnules with smaIl number of lobes.
Family Dipteridaceae
Genus Clathropteris BRONGNIART,
1828: 62

Clathropteris 1I1eniscoides BRONGNIART
PI. 30, Fig. 8; Text-figs. 3a-e
Japanese specimens:

Clathropteris meniscoides BRONGl'IART: 01S111
and HUZIOKA, 1938, p. 78, pI. 8, fig. 4(Nariwa Group) : OISHI, 1940, p. 214, pI. 5,
fig. 4 (Neiridani, Kuruma Group).
Clathropteris obor-ata 01SlII: OISHI and HUZIOKA, 1938, p. 78, pI. 9, figs. 1-2 (Nariwa
Group) .
Chinese specimens:
Clathropteris meniscoides BRONGNIART (bysome spelt C. meniscioides): SZE, 1931,
p. 4, pI. 1, fig. 3 (Rhaeto-Liassic of
Jiangxi) : p. 33, pI. 5, fig. 3 (Jurassic of
Fang-zi) : 1949, p. 6, pI. 4, fig. 4 (non pI.
1, fig. 5 referable to C. obovata) (Lower
Jurassic of Hubei) : Hsc (in SZE and HsU,
1954) , p. 51, pI. 44, figs. 1-2 (Upper
Triassic of Yunnan) : SZE et a!., 1963, p.
85, pI. 26, fig. 5 (non pI. 27, fig. 4 referable to C. obovata) (Lower Jurassic of
Hubei) : LI, P. C., 1964, p. 108, pI. 2, figs.

737.

Early Jurassic plants

1, 2a, 3, 4; pI. 3, figs. 1-6; text-fig. 3(Upper Triassic of Sichuan): LI, P. C. eta\., 1976, p. 108, pI. 19, fig. 4 (Upper
Triassic of Yunnan): FEl'\G et a\., 1977,
p. 211, pI. 79, fig. 4 (Upper Triassic of
Guangdong): ZlIOU, T. S., 1978, p. 101,
pI. 18, figs. 7, 7a, 8 (Rhaeto·Liassic of
Fujian) : YA-;G, 1978, p. 485, pI. 168, fig.
1 (Upper Triassic of Sichuan) : HsC et a\.,1979, p. 22, pI. 20, fig. 7; pis. 21-25; pI.
26, figs. 1-6; pI. 27, fig. 1 (Upper Triassicof Sichuan) : HE et a\., 1979, p. 136, pI.
61, fig. 3 (Upper Triassic of Qinghai).
Specimens from other regions and further
references, see HARRIS, 1931, pp. 88-89
and VAKIIlC\/VIEEV et a\., 1980, p. 54.
Material: SHI-OOl, 057, 059 (Shinadani),
SAS-005, 014, 016-019 (Sasakomata), NEE153, 245 (Higashi-Neiridani) , NE-065, 084(Neiridani), Kr-007 (Tsuchizawa), A-2088(Iwamuro) .

Description.-Leaf-fragments are medium-sized, but the petiole and 'arms' of its
first branch were not found. Pinnae are
elongate-Ianceolate, gradually tapering
distally, and attaining a width of 3.5 em
in Text-fig. 3a (right side). The basal
lamina of adjacent pinnae is connected asshown in Text-fig. 3a. Margins areshallowly toothed, length of tooth andthat of its supporting lateral vein are, 1
em and 2.5 em (Text-fig. 3a), 1 em and 2.3
cm (Text-fig. 3d), 0.8 em and 3.7 em (Textfig. 3b; irregularly toothed), and 0.3 em
and 1.7 em (Text-fig. 3c). Marginal teeth
are short, triangular, acutely or obtusely
pointed and directed forwards. The pinnarachis is rather slender, 0.5 mm -1 mm wide,
lateral veins are distinct, arising at angles
of 40-50 degrees and at intervals of 10-12
mm. Tertiary veins are also distinct,
arising from lateral veins at about a rightangle, percurrent and slightly flexuous,by which the interspace between theadjacent secondaries are more or lessregularly divided into one series ofelongated rectangular meshes, typically 4

191

mm X 8 mm. The finest veins form a net
with rectangular meshes, typically 0.75
mm X 1.25 mm, usually with long axes
parallel to the lateral veins and withoccasional blind ending branches. Thesebranches are simple or bifurcate, or sometimes percurrent as shown in Text-fig.
3e. We have no fertile specimens.
Distribution and occurrence.-Clathropteris meniscoides is rather rare in the Kuruma Group and the Iwamuro Formation,but occurs widely in the Upper TriassicLower Jurassic of Europe, Central Asia
and China, and the Upper Triassic of SEAsia and Japan.
Discussion and comparison.-Our leaves,though they are all sterile, are characterized by clathrate meshes of the first orderformed by lateral and tertiary veins andclosely resemble Clathropteris meniscoides
BRONGNIART first described from Europe.
They resemble specimens of HARRIS (1931)
in venation.
Minor differences among the leaves ofthis species are given in detail by KON'NO(1968). According to him, in Clathropteris,the tertiary veins are distinguished bytheir manner of branching into thefollowing three types:
1) The tertiary veins are dichotomously
branched and percurrent clathratetertiaries are seldom developed as in
Clathropteris obovata.
2) The tertiary veins branch off in the
sympodial ramification, the clathratetertiaries are developed, but lessprominent, zigzag in course, giving
off alternately or subalternately
relatively thick quaternary veins.
3) The tertiary veins give off only verythin quaternaries in the manner ofmonopodial ramification, and theclathrate tertiaries are definitely
developed, thick, straight or slightly
arc-shaped in their course.
Thus our leaves may belong to his type-Tatsuaki KIMURA and Masanori TSUJI!

192

are close to his Clathropteris
meniseoides from the Upper Triassic ofsoutheastern border of Sarawak, Borneo.
As mentioned by KON'NO (1968), most ofleaves from Europe belong to his type-3.
So far as its venation is concerned,
Clathropteris obovata OISHI described by
OISHI and HUZIOKA (1938) from the
Nariwa Group is referable to C. meni2 andseoides.
Korean leaves regarded by Y ABE (1922)
as Clathropteris cfr. meniscoides and by
KA WASAKI (1925, '39) as C. meniseoides
are similar in venation to ours, but as
pointed out by HARRIS (1931), they havelarger marginal teeth than those ofEuropean leaves. From China, many leaves
were described under Clathropteris meniseoides (or meniseioides) as mentioned
before. They are all similar in venation
to ours, although according to KON'NO(1968) LI'S leaves from Sichuan are hardly
distinguishable from C. obovata.
The Tonkin leaves of ZEILLER (1903)
named Clathropteris platyphylla (GOEPPERT) are mostly like ours, but some (hispI. 30. figs. 3-8) have very broad pinnaeand might be distinct. SREBRODOLSKA]A(1961) included ZEILLER'S leaves (pI. 27,
fig. 2 and pI. 28, fig. 2) into her

Clathropteris mongugaiea.
Clathropteris meniseioides described by
STANISLAWSKI (1971) from the Upper
Triassic of Donbass is different from ours
in its small-sized leaf and non-clathrate
venation.
Genus Dietyophyllum LINDLEY
and HUTTON, 1834: 65

Dictyophyllum Iwtakiense Kill-lURA
and Tsu]II sp. nov.
PI. 30, Figs. 9-11; Text-figs. 4a-g
Dictyophyllum sp. : OISHI, 1940, p. 219, pI. 5,
fig. 6 (Neiridani, Kuruma Group). Dictyo-

phyllum ? sp.: KIMURA, 1959b, p. 68, pI.
2, fig. 4 (Kotaki Coal-Field, Kuruma
Group) .
Material: Holotype; KU-9002 (Kotaki
Coal-Field). Paratypes; NNW -223, 265 (NishiNeiridani), NEE-147, 225, 261 (Higashi-Neiridan i) , SHI-013 (Shinadani). Other specimens;
KU-9001, NEE-263 and other 10 specimens,
DAI-056 (Dairagawa), SHI-012 and other 4.
Stratum typicum: Hodonosawa Formation(KIMURA, 1959b after K. SASAI), Kuruma
Group (its corresponding formation in themain distribution of the group uncertain).
Locus typicus: Kotaki Coal-Field (now not in
production), Kotaki-mura, Nishi-Kubiki-gun,
Niigata Prefecture. Derivatio nominis: After
Kotaki where the holotype was collected by
K. SASAI in 1949.

Diagnosis. - (Petiole and 'arms' unknown.) Pinna rachis stout, 3.5-4 mm wide
below, 2.5-3 mm middle and 0.5 mm above.
Basal lamina undivided but broadly
undulated. Laminae usually developing
lateral segments, but incisions not to the
pinna rachis. Segments connected by a
web of lamina along the pinna rachis,becoming closer towards the pinna apex.
Segments in the middle region rectangular,
about 5.2 em long and 1.8 em wide, withbluntly pointed or rounded apex andattached perpendicularly to the pinnarachis. Apical segments attached obliquely,
triangular in form and with bluntly
pointed apex. Larger segments having
shallowly undulated margins or almost
entire; margins of smaller segments
entire.
Midrib of segment strongly
marked, lateral veins arising at intervalsof 7 mm at an angle of 55-90 degrees,making rather irregular polygonal meshes.
Smaller veins forming polygons, 1.5 mm
wide, enclosing blind vein-endings.
In fertile leaf, blind veins being theplacental bundles. Sori 1.1 mm in diameter, consisting generally of 6 sporangia
widespread but avoiding the region near

737.

Early Jurassic plants

4a

193

4b

Text·figs. 3-4 (a-b). (All natural size, unless otherwise indicated).
3. Clathropteris meniscoides BRO~G~L\RT. Sterile leaves (3a-d); Quaternary and finest veinsare excluded. 3a: (NE-065). 3b: (SAS-018; PI. 30, Fig. 8). 3c: (NEE-153). 3d: (SHI001). 3e: Final meshes drawn from 3b.
4. Dictyophyllum kotakiense KI~ICRA and TSUJI! sp. nov. 4a: Pinna base with a thick rachis(NNW -223, para type ; PI. 30, Fig. 10). 4b: Basal pinna with lateral segments (NEE-261;
para type) .

the midrib. Sporangia almost sessile,about 500 pm in diameter. Annules welldeveloped, probably complete and oblique.
Distribution and occurrence.-DictyophyllulIl lwtakiense is common in the
Negoya Formation, but rare in the
Shinadani and Hodonosawa Formations.
Discllssion and comparison.-Our material
is represented by a series of leaf-fragmentsof varied size and form. Our leaf-fragments look just like corresponding partsof the leaves figured as Dictyophyllum
nilssoni (BRONGNIART) GOEPPERT by
HARRIS (1931), OISHI (1932) and D.

spectabile NATHORST by OISHI (1932).
Their sori, however, are different. In
HARRIS' specimens at least the sporangia
are attached singly or in pairs on ultimate
veins, or at most in groups of three(according to his information given us in1981). In our leaf they are in groups ofsix or more and this is an importantdifference and precludes a specific identification and for this reason, we describe
our material as a new species.
OISHI's Dictyophyllum nilssoni and D.
spectabile from the Nariwa Group are all
Dictyorepresented by sterile leaves.

194

Tatsuaki KIlvIURA and Masanori TSUJI!

phyllum nilssoni illustrated by YANG
(1978) and D. cfr. nilssoni by SZE (1933a,'49), SZE et al. (1963) and FENG et al.
(1977) from China are also all represented
by sterile leaves. So it is difficult to makethe full comparison between ours andthese Asian sterile leaves which aresimilar in external form to ours.
Genus Thaumatopteris GOEPPERT,1841 : 2

Thaumatopteris elongata OISHI
PI. 30, Fig. 12; PI. 31, Figs. 1-2;
Text-figs. 5a-e
Thaumatopteris schenki NATIIORST: OISIII,
1931, p. 239, pI. 1, figs. 7, 7a, 8, 8a(Kuruma, Kuruma Group): 1932, p. 296,
pI. 17, figs. 3-4; pI. 18, fig. 1 (Nariwa
Group) .
Cfr. Thaumatopteris brauniana POI'I': OISIII,
1932, p. 297, pI. 35, fig. 4 (Nariwa Group).
Thaumatopteris elongata OISHI: OISIII, 1932,
p. 295, pI. 34, fig. 2; pI. 35, figs. 1-2(Nariwa Group): OISHI and YA~IAsITA,
1936, p. 149: OISHI, 1940, p. 223 (general
remarks): KI~IURA, 1959a, p. 10, pI. 2,
figs. 1-8; text-fig. 4 (Iwamuro Formation).
Thaumatopteris nipponica 015111: KI~ICRA,
1959a, p. 11, pI. 2, figs. 9-10 (Iwamuro
Formation) .
lvlaterial: NEE-268 (Higashi-Neiridani),
DAI-059 (Dairagawa), Kr-150 (Tsuchizawa),
A-2033 and other 16 specimens (Iwamuro).

Description.-Our material consists ofsterile pinna fragments of varied size andform but no petioles or pinna bases. The
pinna rachis attains a thickness of 2.5 mm(Text-fig. 5a) and bears pinnules withbluntly pointed or rounded apex, up to 1
em wide and more than 4.5 em long
(Text-figs. 5a-c). A short pinnule shownin Text-fig. 5d is possibly on the lower
part of pinna. The pinnules arise nearly
perpendicularly, closely or remotely set.

The margins are entire or broadly sinuous(Text-figs. 5c-e) but never lobed. At
their base, pinnules are not expanded(Text-fig. 5b), but in some specimens thelamina of adjacent pinnule is connectedby a web, 0.5 mm wide which runs along
the pinna rachis. The midrib is prominent,
but the other veins scarcely project fromthe lamina. The secondary nerves ariseata wide angle, then form polygonal meshes.
The first veins form rather elongatehexagonal meshes, 1.5 mm long and 0.5
mm wide (Text-figs. 5c-e). Blind veinendings sometimes occur in these meshes.
Distribution and occurrence.-Thaumatopteris elongata is possibly a local species,
and occurs rarely in the Iwamuro Formation. Kuruma and Nariwa Groups inJapan.
Remarks.-Our synonymy gives thehistory of Japanese specimens 'which we
determine as Thaumatopteris elongata.
Among our leaves, some pinnules withbroadly undulated margins remind us of
those of European Thaumatopteris schenki.
But Thaumatopteris elongata is distinguished by its pinnules with broadly undulated
margins instead of broadly lobed ones inT. schenki (HARRIS, 1931, fig. 35).
Our pinnules with entire margins remind
us also of the European Thaumatopteris
brauniana. But Thaumatopteris elongata
differs from T. brauniana in its pinnules
both with entire and broadly undulated
margins on the same pinna, instead ofpinnules unexceptionally with entire
margins in T. brauniana (HARRIS, 1931,
fig. 36).
Thaumatopteris nipponica OISHI is distinguished from T. elongata by its elongatetriangular or sometimes deltoid pinnules
with the secondary nerves occasionallybifurcating at a short distance from their
origin.

737.

Early Jurassic plants

6a lmm~/ [ t[j

3cm

49

195

4c

4e

5e

-&

O.5cm

~

5d

6btj
lcm

5b

5a

4t

Text·figs. 4 (c-g) -6. (All natural size, unless otherwise indicated).
4. Dictyophyllum kotakiense KI~ICRA and Tsc,I1I sp. nov. 4c: Middle region of a sterile pinna(NEE-147, para type ; PI. 30, Fig. 9). 4d: Ditto, with rectangular segment (NEE-255, paratype; PI. 30, Fig. 11). 4e: Upper region of a sterile pinna (SHI-013, paratype). 4f: A partof a fertile pinna, showing the distribution of sori (KU-9002, holotype). 4g: Sorus consistingof 6 sporangia, drawn from the holotype.
'5. Thaumatopteris elongata OISIlI. 5a: A part of sterile pinna with a rather thick rachis (A3016). 5b: Ditto (Kr-150; PI. 31, Fig. 2). 5c: An incompletely preserved sterile pinnule(NEE-215; PI. 30, Fig. 12). 5d: A small-sized sterile pinnule (A-3021). 5e: An incompletely
preserved sterile pinnule, showing the venation (DAI-059).
6. Hausmannia (Protorhipis) sp. A. 6a: An incompletely preserved fertile leaf (A-2041)(redrawn from KI~IURA, 1959a, p. 12, text-fig. 5). 6b: Detailed venation, drawn from 6a.
6c: Occurrence of sori, drawn from 6a.

Tatsuaki KIMURA and il1asanoTi TsuJI!

196

Genus Hausmannia DUNKER, 1846: 12
Hallsmannia (Protorhipis) sp. A
Text-fig. 6a-c

Hausmannia (Protorhipis) sp. : KIMURA, 1959a,p. 11, pI. 12, fig. 9; text-fig. 5 (Iwamuro
Formation) .
J"Iateriai: A-2041
leaf-fragment) .

(Iwamuro)(a single

Description.-Lamina about 4 cm indiameter, petiole not known.
Marginshallowly lobed; the number of lobes notknown. Primary veins radiating frompresumed top of petiole, branching dichotomously at least twice, reaching margin ofeach lobe. Primary veins slightly sinuous,
giving off finer secondary veins perpendicularly to form the first meshes withadjacent primary vein. Tertiary veins
uniting into numerous small secondary
meshes, typically about 1 mm square, but
occasionally ending blindly. Quaternaryveins simple or forking, mostly blindending in sterile part.
Sori circular, 0.5 mm in diameter, borneat the tip of a blind vein. Details ofsorus not visible.
Remarks.-No additional specimen has
been obtained after the one described by
KIMURA (1959a). We limit our specimen
to leaves described from East Asia incomparison. Of these Hausmannia crenata
by OISHI (1932) from the Nariwa Group is
distinguished by its secondary veins arisingobliquely and forming polygonal or sometimes rhomboidal meshes. Hausmannia
naritl'aensis described by OISHI (1930) fromthe Nariwa Group has almost entire or
broadly undulated margin as does H.
ussuriensis by KRYSHTOFOVICH (1923).
Hallsmanllia dentata described by OISHI(1932) also from the Nariwa Group
resembles ours in its lobed or dentate
margin of lamina. But it is distinguishedfrom ours by its generally unbranchedprimary veins and obliquely arised secondary veins. Hausmannia leeiana originallydescribed by SZE (1933a) from the LowerMiddle Jurassic of Shanxi is easily distinguished from ours by its entire or
shallowly undulated margin.

Sterile and unclassified
fern leaves
We have a large number of sterile fern
leaves and a few fertile ones withuncertain affinity.
Among them wedistinguished the following forms, depending on their external outline of pinnules
and venation.
Key to the sterile and unclassified ferns:
1. Pinnules small, less than 1 em long ..
. . . . . . . . . . . . . . .. Sphenopteris sp. A
1'. Pinnules often over 1 cm long .... 2
2. Pinnules less than twice as long as
broad ............ Cladophlebis sp. C
2'. Pinnles more than twice as broad .. 3
3. Pinnules with once forked lateralveins ............................ 4
3'. Pinnules with twice forked lateralveins ............................ 5
3/1. Pinnules with thrice forked lateralveins ........ Cladophlebis ku runzensis
4. Lateral veins up to 10 pairs ....... .
· ....... Cladophlebis pseudodelicatula
4'. Lateral veins 10-12 pairs ......... .
· ............. Cfr. Todites fllkutomii
5. Margins entire .................... 6
5'. Margins dentate at apical half ..... .
· . . . . . . . . . .. Cladophlebis raciborskii
6. Apex acute .. Cfr. Asterotheca okafujii
6'. Apex blunt. . Cfr. Todites yamanoiensis
This key omits some undeterminable
fragments including Cladophlebis sp. A(KIMURA, 1959a, p. 20, pI. 4, fig. 1) andC. sp. B (Ibid., pI. 3, fig. 6) from the
Iwamuro Formation.

737.

Early Jurassic plants

Genus Asterotheca PRESL in
CORDA, 1845: 81
Cfr. Asterotheca okajujii KIMURA
and OHANA
PI. 31, Figs. 3-4; Text-figs. 7a-d
Cladophlebis cfr. raciborskii ZEILLER: OISHI,
1931, p. 234, pI. 16, fig. 6, 6a; pI. 17, fig.
1 (Tsuchizawa, Kuruma Group).
Cladophlebis raciborskii ZEILLER forma integra
OISHI and TAKAI-IASI : OISHI and T AKAIlASI,
1936, p. 119 (Yamanoi, Habu Group):
OISHI, 1940, p. 283, pI. 19, fig. 1, la(~1omonoki, Mine Group) : KIMURA, 1959a,
p. 19, pI. 3, fig. 5; pI. 5, fig. 3 (Iwamuro
Formation). For further references, see
KI~ICRA and OIIA:"A, 1980, p. 74.
Comparable specimens: Asterotheca okafujii
KI~IL-RA and OHANA : KIMURA and OIlANA,
1980, p. 73, pI. 1, fig. 1; pI. 2, figs. 1-2;
pI. 3; pI. 5, fig. la; figs. 2a-f (Momonoki
Formation, Mine Group).

197

simple.
Distribution and occurrence.-This form
is common in the Kuruma Group andIwamuro Formation. Sterile leaves referable to this form are widely known in
the older Mesozoic plant-beds in East Asia.
Remarks.-Our leaves agree fully withthe sterile leaves of Asterotheca okajujii
originally described by KIMURA and OHANA(1980) from the Upper Triassic Momonoki
Formation, based on abundant leaves bothwith sterile and fertile parts. Similarsterile leaves had been determined by
OISHI and T AKAHASI (1936) as Cladophlebis raciborslcii forma integra.
Cfr. Asterotheca okajujii resembles
Cladophlebis raciborskii originally describedby ZEILLER (1903) from the Upper Triassicof North Viet-Nam, but it differs fromZEILLER'S species by its pinnules withentire margins.

Material: NE-034, 035, 042 (Neiridani),
NNW-061, 068, 234,428,511 (Nishi-Neiridani),
NEE-267 (Higashi-Neiridani), DAI-009, 014(Dairagawa), TOB-033, 046B (Tobiiwadani),
Kr-041, 074, 081 (Tsuchizawa), A-0318, 4065,
75039,75119 (Iwamuro).

Genus Todites SEWARD, 1900: 86

Description.-Frond is bipinnate andlarge-sized. Pinnae are nearly parallelsided for the most part, the largest (Kr081) is 6 cm wide.
Pinnules are set
closely, long and narrow, nearly parallelsided for the most part, then suddenly
narrowing towards the acutely pointed
apex, up to 2.9 cm long and 0.6 cm wide,
attached katadromically to the pinna axis
at an angle of 70-90 degrees, but theangle is reduced near pinna apex, apicalpinnules are often falcate. Margins are
entire, midvein is prominent, persisting to
the tip. typically sending off 15 pairs oflateral veins rather obliquely. Basal 8-9
pairs of lateral veins are usually forking
twice, others forking once or occasionally

Cladophlebis nebbensis (BRONC:"IART) NATHORTS: OISHI, 1931, p. 231, pI. 16, fig. 4, 4a(Tsuchizawa, Kuruma Group). For further
references, see KIMURA and OIIAl"A, 1980,
pp. 77-78.
Comparable specimens: Todites fukutomii
KI~lCRA and OHAKA: KI~It.;RA and OHAKA,
1980, p. 77, pI. 3, figs. 1-2; pI. 4, figs. 23; pI. 6, fig. 1; figs. 3a-c (Momonoki
Formation, Mine Group).

Cfr. Todites jll/mtomii KIMURA
and OHANA
PI. 31, Figs. 5-6; Text-figs. 8a-b

Mate/-ial: NE-011, 031 (Neiridani), NNW461, 660, 679 (Nishi-Neiridani), NEE-375?
(Higashi-Neiridani), TOB-027 (Tobiiwadani).

Description.-Frond is bipinnate, largesized. Rachis is rather slender, 1.5 mm
wide. Pinnules are subopposite, arisingat an angle of 80 degrees, straight and198

Tatsuaki KIMURA and Masanori TSUJI!

8b

Text-figs. 7-9. (All natural size).
7. efr. Asterotheca okaflljii KIMCRA and OllA;\;A. 7a: A pinna fragment with large-sizedpinnules (NNW-061; PI. 31, Fig. 3). 7b: (NEE-267). 7c: (NE-042). 7d: (DAI-009; PI. 31,
Fig. 4).
8. efr. Todites fllklltomii KJ:\IURA and OHA;\;A. Sa: The largest specimen with a rather slenderrachis (NE-031; PI. 31, Fig. 5). Sb: (NNW -679).
9. efr. Todites yamanoiensis (YOKOYAMA) KlJ"vlURA and OllA:-iA. 9a: (KU-4981202-1S). 9b: (A0405; PI. 32, Fig. 1).

probably almost uniformly wide up to theapex. Pinnules are set closely, commonly
arising at a wide angle, katadromic inorder, finger-shaped, with rounded or
obtusely pointed apex, typically 1.4 cm
long and 0.4-0.6 cm wide; margins are
entire. Basal basiscopic margin is often
slightly contracted but basal acroscopic
margin slightly expanded. Basal pinnules
are not specialized. Midvein is distinct,
persisting to the tip and sending off

typically 10-12 pairs of once forked lateralveins at a wide angle.
Distribution and occurrence.- This form
is occasionally found in the Shinadani and
Negoya Formations,
Kuruma Group.
Similar leaves are widely distributed in
the older Mesozoic plant-beds in Japan.
Remarks.-Our leaves agree in all features with the sterile leaves of Todites
fukutomii originally described by KIMURA
and OHANA (1980) from the Momonoki

737. Early Jurassic plants
Formation. Though we feel nearly surethat our present sterile leaves are sterile
Todites jukutomii, we prefer not to identify
them fully until we find fertile specimens
associated with them.
Similar sterile leaves from Japan havebeen described by OISHI (1931) as Cladophlebis nebbensis from Tsuchizawa.
Cfr. Todites yamanoiensis (YOKOYAMA)KIMURA and OHANA
PI. 32, Fig. 1; Text-figs. 9a-b
Cladophlebis haiburnensis(LINDLEY and
HUTTO:-:) : OISI-II, 1931, p. 237, pI. 2, fig.
2, 2a (Tsuchizawa): KIMURA, 1959b, p.
69, pI. 1, fig. 5; pI. 2, fig. 1 (Kotaki
Coal-Field) . For further references, see
KIMCRA and OI-IANA, 1980, p. 82.
Comparable specimens: Todites yamanoiensis(YOKOYAMA) KIMURA and OHANA: KI~IURA and OHANA, 1980, p. 82, pI. 4, fig.
1; pI. 5, fig. 1b; pI. 6, figs. 2-3; figs. 5ag (Momonoki Formation, Mine Group).
Material: NNW -498, 663 (Nishi-Neiridani),
SAS-008 (Sasakomata), 4981201-18 (Kotaki
Coal-Field), A-0405 (Iwamuro).

199

Remarks.-Our leaves agree in all features with the sterile leaves of Todites
yamanoiensis described by KIMURA and
OHANA (1980) from the Momonoki Formation. Sterile leaves resembling ours
had been determined as Cladophlebis
haiburnensis (LINDLEY and HUTTON) by
previous authors.
Our leaves are distinguished from Cladophlebis aktashensis TURTANOV A-KETO VA
by its shorter pinnules with fewer lateralvein pairs (7-11), instead of more elongatedpinnules with 12-15 (or more) pairs oflateral veins in C. aktashensis.
We consider the sterile leaves regarded
by OISHI (1931) as Cladophlebis haiburnensis
from Tsuchizawa and by KIMURA (1959b)from the Kotaki Coal-Field are like ours.
As previously mentioned by KIMURA
and OHANA (1980), our leaves resemble
those of Cladophlebis asiatica CHOW andYEH (in SZE et al., 1963) known from theLower-Middle Jurassic of China.

Form-genus Sphenopteris
STERNBERG, 1825: 15
Sphenopteris sp. A

Description.-Several detached sterile
pinnae were obtained. Pinnules are set
closely, elongate-triangular or elongaterectangular, apex obtusely pointed orrounded, and attached to the pinna rachis
at a wide angle; typically 2.2 cm longand 0.5 cm wide; margins are entire.
Basiscopic basal margin is sometimes
distinctly contracted and acroscopic basal
margin is usually expanded. Midvein isdistinct, persisting to the tip, typicallysending off 9-10 lateral veins usually
forking twice.
Distripution and occurrence.-Cfr. Todites ya11lanoiensis is occasionally found in
the Kuruma Group and rare in the Iwamuro
Formation. Similar leaves occur in other
older Mesozoic plant-beds in East Asia.

PI. 31, Figs. 7-8; Text-figs. lOa-c
Material: NNW -704,784,786-788, 830, 833.
835, 837, 839 (Nishi-Neiridani).

Description.-Leaf probably bipinnate,.
of small size, partly fertile and partly
sterile, but whole leaf unknown. Pinnules.
set rather remotely, oblong or elongatetriangular, typically 0.9 cm long and 0.4
cm wide, apex rounded, base distinctly
contracted.
Margins shallowly lobed(Text-fig. 10c); lobes 4-7 pairs. Midvein
distinct but not persisting to the tip,
sending off lateral veins not reaching themargin. Each lobe receiving a set oflateral vein, basal pair of lateral vein
forked once or twice, others mostly simple.

200

Tatsua1ci KIMURA and Masanori T suJI!

12a

~

. :w

lla

/' g~c~~Text-figs. 10- 14. (A ll na tura l size , unless otherwise indi cated) .
10. Sphenopteris sp . A. lOa: A part of fer til e leaf (NNW - 282; PI. 31, Fig. 7) . lOb: Twodetached fe rtil e pinna fragme nt s (NNvV - 784) . 10c: A pinnll le show in g th e ve nation a nd theposition of sori, draw n from lOa.
11. Cladophlebis kurum ensis KI MURA and T SG JII sp . nov . lla : A pinn a f ra g me nt beari ng large sized pinnules (TOB- 018B). llb: Two medium- sized pinnules, sh ow in g the ve nat ion (Kr - 068,
parat y pe). llc: A poorly prese r ve d large -sized pinnule (A - 0413) .
12. Cladophlebis pseudodelicatula 01 5 1-11. 12a: A part of a bipinnate frond (A - 4068; PI. 32, F ig.
3) (redrawn from KI ~WRA, 1959a , p. 12, te x t -fig. 6, w ith some e me ndation ) . 12b: A poorl yprese rved pinna , s howing th e outline of pinnu les (N EE- 099; PI. 32, Fig. 4). 12c: A pinna
fragm e nt bearing s mall-s ize d pinnllles (NE- 149; PI. 30 , Fig. 13) .
13. Cladophlebis raciborskii ZEILLER: (A- 3048).
14. Cladophlebis s p. C: (SI-I[ - 015; PI. 32, Fig. 10) .

In fertile pinnul es, each lateral vein terminated by a circular sorus, 0.5 mm in diameter (Text-figs. lOa , c). Detai ls of sorus
not visible.
Distribution and occurrence. - Sphenopteris sp. A is rare and occurs at a single
locality along the Nishi-Neiridani (Negoya
Formation ).

Remarlls.- Ow ing to our uncerta int y ofits soral character, we can onl y determin e
our leaves as SphenojJteris sp. A .

Form-genus CladojJhlebis
B RO NGN JART, 1849: 105

737.

Early Jurassic plants

Cladophlebis kurumensis KI:VlURA

and TsuJII sp. nov.
PI. 31, Fig. 9; PI. 32, Fig. 2;
Text-figs. 11a-c
Cladophlebis sp. a: OISHI, 1931, p. 238, pI. 17,

fig. 3 (Tsuchizawa).
Material: Holotype; Kr-052 (Tsuchizawa).
Paratypes; Kr-035, 041, 066, 068. Other specimens; TOB-018B (Tobiiwadani), A-0413 (Iwamuro).
Stratum typicum: Alternation ofsandstone and shale (Kr 2) , Kuruma Group
Locus(see KI1vIURA and TSUJII, 1980a).
typicus: Tsuchizawa, near Kuruma, KitaOtari-mura, Kita-Azumi-gun, Nagano Prefec·
ture. Derivatio nominis: After Kuruma, aclassical locality where the holotype and paratypes were collected.

Diagnosis.-Frond bipinnate. (Whole leaf
unknown.) Pinnules set closely, elongatetriangular in form, attached at an angleof 60-70 degrees to the slender pinnarachis, large-sized, typically 3.6 cm longand 1 cm wide at base, tapering gradually
towards the acutely pointed apex; sometimes falcate and expanded at acroscopic
base. Margins entire. Midvein rather
weak, sending off 12 pairs of lateralveins; the acroscopic ones directed forwards. Basal 4-5 pairs thrice forking
and others forking twice except apical 12 pairs forking once.
Distribution and occurrence. - Cladophlebis kurumensis occurs rarely at Tsuchizawa and Tobiiwadani (Kuruma Group)
and at Iwamuro (a single specimen).
Discussion and comparison.-Cladophlebis
kurumensis is characterized by its largesized and elongate-triangular pinnules withthrice forked lateral veins. The speciesof Cladophlebis known to us with thrice
forked lateral veins are mentioned below
for comparison.
The sterile leaf of Todites fastuosus(KIMURA) KIMURA and Tsu]II previously

201

described (KIMURA and TsuJII, 1980b)
from the Iwamuro Formation and Kuruma
Group, is distinguished from Cladophlebis
kurumensis by its triangular pinnules withfinely dentate margins.
Cladophlebis bitchuensis originally described by OISHI (1932) from the Nariwa
Group and recently by KIMURA and
OHANA (1980) from the Momonoki Formation, is distinguished by its elongateoblong or broadly linear pinnules with arounded or obtusely pointed apex.
Cladophlebis gigantea OISHI (OISHI, 1932)
from the Nariwa Group and C. gigantea
of P'AN (1936) from the Upper Triassicof China, and C. dr. gigantea of SZE(1956) and of SZE et al. (1963) from theUpper Triassic of China all have theirpinnule margins shallowly crenulated or
lobed.
Two ferns described from Malaysia by
KON'NO (1972), Cladophlebis dr. haiburnensis and C. ishiiana, look just like ours
but we refrain from identifying our leaves
with them, because in his C. dr. haiburnensis, pinnules are more narrower thanours and mostly directed forwards, andin his C. ishiiana, pinnules are deltoid or
triangular, rather resembling the sterilepinnules of Todites fastuosus.
An incomplete and deformed pinnule
illustrated in Text-fig.11c, probably belongs
to Cladophlebis kurumensis, though its
lateral veins are all thrice forked.
Cladophlebis pseudodelicatula OISHI
PI. 30, Fig. 13; PI. 31, Fig. 10; PI. 32,
Figs. 3-4; Text-figs. 12a-c
Cladophlebis sp. b: OISIII, 1931, p. 238, text-

fig. 2 (Tsuchizawa).
pseudodelicatula OISHI: OISHI,
1932, p. 288, pI. 11, fig. 2 (Nariwa Group) :1940, p. 281 (general remarks): SCHOROCHOVA and SREBRODOLSKAJA, 1979, p. 68,
pI. 11, figs. 1-3 (Upper Triassic of

Cladophlebis

202

Tatsuaki KIMURA and Masa,nori TSUjII

Primo rye) .
Cladophlebis clavatum KIMURA: KIMURA, 1959a,
p. 12, pI. 2, fig. 11; pI. 5, fig. 2; pI. 6,
figs. 1-2, 6, 9; text-fig. 6, 7 (lwamuro).
Material: NE-149 (Neiridani), NEE-047,
099 (Higashi-Neiridani), A-4063 and other S
specimens (lwamuro).

Emended diagnosis.-Frond bipinnate, rachis rather thick, 6 mm wide, sending off
pinnae suboppositely at intervals of 3 cm
at the middle portion of frond. Pinnae
arising at an angle of 40 degrees, thenbending outwards, linear lanceolate inoutline, tapering very gradually from thebase upwards to an acuminate apex.
Pinna rachis slender. Pinnules linear toelongate-Ianceolate, closely set katadromically, and attached by their whole baseto the pinna rachis, borne at a wide angle
or often perpendicular, typically 2 cm longand 0.5 cm wide, apex acute.
The first pair of pinnules larger thanothers. lVlidvein distinct, straight, persist-ing to the tip, and sending off typically
9-10 pairs of lateral veins at an angle of
45 degrees. Lateral veins mostly once
forking. branches forming a narrow angle.
Distribution and occurrence. - Cladophlebis pseudodelicatula occurs rarely in
the Nariwa and Kuruma Groups andIwamuro Formation. Similar leaves havebeen recorded from the Upper Triassicof Primo rye.
Discussion and comparison.-OlsHI (1932)
based this species on pinna fragmentsfrom the Nariwa Group and included his
Cladophlebis sp. b (1931) from Tsuchizawa
in it.
We have reexamined the specimens
described as Cladophlebis clavata (firstspelt 'clavatum') by KIrJlURA (1959a) andnow conclude that they are identical with
C. pseudodelicatula OISHI.
Additional leaves newly obtained fromthe Negoya Formation, though they arefragmental. also agree well with OISHI's
species.

Explanation of Plate 30
Fig. 1. Phlebopteris sp. A; a detached fertile pinnule (NE-123; Text-fig. Ib), x 2.
Figs. 2-7. Coniopteris neiridaniensis KIMURA and TSUJI! sp. nov.; all figures, x2. 2. Sterile
leaf-fragment with large-sized pinnules (NNW -293, holotype; Text-fig. 2a). 3. A sterile pinna
fragment with four large-sized pinnules (NNW -590, para type ; Text-fig. 2c). 4. Sterile leaffragment with medium-sized pinnules; a stem-like impression shown by a-b does not belong
to this leaf (NEG-120, paratype; Text-fig. 2b). 5. A sterile pinna fragment with mediumsized pinnules (OAI-055, paratype; Text-fig. 2f). 6. A fertile pinna fragment (NNW -561,
paratype; Text-fig. 2k). 7. A fertile leaf-fragment (NE-053, paratype; Text-fig. 21).
Fig. S. Clathropteris meniscoides BR001GNIART; a pinna fragment, showing clathrate meshes(SAS-OlS; Text-fig. 3b, c), xL
Figs. 9-11. Dictyophyllum kotakiense KI~ICRA and TSUJII sp. nov.; all figures, xL 9. A pinna
fragment (NEE-147, paratype; Text-fig. 4c). 10. Basal part of pinna base, with a thick
pinna rachis and undivided lamina (NNW -223, para type; Text-fig. 4a). 11. Two poorly
preserved lateral segments; an incision is not to the pinna rachis (NEE-225, para type ; Textfig. 4d).
Fig. 12. Thaumatopteris elongata OISHI; a detached sterile pinnule, with broadly sinuous margins(NEE-215; Text-fig. 5c), xL
Fig. 13. Cladophlebis pseudodelicatula OISHI; two detached pinna fragments (NE-149; Text-fig.
12c). xLKIMURA and TSUJII:

Early Jurassic plants

Plate 30

'73'7.

Early Jurassic plants

Cladophlebis pseudodelicatula looks rather
like C. delicatula of YABE and OISHI(1933) from the Jurassic of NE-China(possibly from the Lower Cretaceous
Yushugo:I Formation. according to therecent knowledge). It is also like Cladophlebis arglltula (HEER) from the JurassoCretaceous of East Siberia, but its pinnules
are more slender.
Cladophlebis racibors/n'i ZEILLER
PI. 31, Fig. 11, PI. 32, Figs. 5-9;
Text-fig. 13/'eciborskii ZEII.I.ER: ZE[LI.ER,
1903, p. 49, pI. 5, fig. 1 (Upper Triassicof North Viet-Nam) : KAW.,\SAK[, 1925, p.
15, pI. 7, figs. 25-29; pI. 34, fig. 94(Daedong Group, Korean
Peninsula):
OISHI (1932) (pars), p. 286, pI. 10, fig. 1(Nariwa Group; other leaves are referable
to Cfr. Asterotheca okafujii) : 1940, p. 282(general remarks) : LEE, 1955, p. 35, pI. 1,
figs. 2-8 (Middle Jurassic of N-Shaanxi):
SZE, 1956, pp. 20, 128, pI. 21, fig. 7; pI.
22, fig. 3, 3a; pI. 26, figs. 1-7; pI. 27,
figs. 1-5; pI. 53, fig. 3? (Upper Triassicof Shaanxi) : C[IOII' and CI[A";G, 1956, p.
55, pI. 1, figs. 6-8 (Upper Triassic of
Neimeng): K[MURA, 1959a (pars), p. 17,
pI. 4, fig. 7; pI. 5, figs. 4, 6; text-fig. 11(Iwamuro): 1959b, p. 70 (Kotaki CoalField): SZE et a!., 1963, p. 106, pI. 35,
fig. 3; pI. 36, fig. 3; pI. 37, fig. 3 (allreinserted from SZE, 1956): L[, P. C. eta!., 1976, p. 113, pI. 27, fig. 6, 6a; pI. 28,
figs. 3-5 (Upper Triassic of Yunnan):
HE et a!., 1979, p. 142, pI. 65, fig. 1, 1a(Upper Triassic of Qinghai): Hsc et a!.,1979, p. 33, pI. 16, figs. 7, 8; pI. 17, fig.
1, 1a (Upper Triassic of Sichuan) : Z[[OU,
H. Q. et a!., 1980, p. 78, pI. 2, figs. 1-2;
pI. 29, fig. 3; pI. 30, figs. 1-2 (Upper
Triassic of Shaanxi and Neimeng).

Cledophlebis

Material: A-0305, 0322, 0496 and other 7
specimens (Iwamuro), Kr-047, 069 (Tsuchizawa) , 4981103-2 and other 2 specimens(Kotaki Coal-Field) , NE-015 (Neiridani), NNW-

203650, 764 (Nishi-Neiridani), SHI-008 (Shinadani),
TOB-046A (Tobiiwadcni).

Distribution and occurrence. - Cladophlebis raciborskii is common in the
Iwamuro Formation but rather rare inThis species is
the Kuruma Group.
widely distributed in the older Mesozoic
plant-beds in East Asia.
Remar/?s. - The sterile pinnules ofAsterotheca oka/ujii are of similar sizeand shape but differ in having entire
margins in their distal halves (compare
Text-figs. 12c and 13).
Cladophlebis sp. described by SZE (1949)
from the Lower Jurassic of Hubei also has
pinnules of similar size, but their distal
margins have not been figured.
Among the leaves regarded by KIMURA(1959a) as Cladophlebis raciborskii fromthe Iwamuro Formation, some weremisidentified excluding pI. 2, fig. 12 andpI. 4, 7, 9 which we now determined as
Cfr. Todites denticulatus.
Cladophlebis sp. A
Cladophlebis sp. A: KI!o.[LRA, 1959a, p. 20, pI.

4, fig. 1 (Iwamuro).
Meterial: A-4073 (a single specimen).

Cladophlebis sp. B
Cladophlebis sp. B: K[MURA, 1959a, p. 20, pI.

3, fig. 6 (Iwamuro).
Material: A-3067 (a single specimen).

Cladophlebis sp. C
PI. 32, Fig. 10; Text-fig. 14
Material: SHI-015, 016 (Shinadani).

Descriptioll.- Two pinna fragments arepreserved on a sandstone slab. A partof one is shown in Text-fig. 14. Pinnules
closely set, varied in form, deltoid or204

Tatsuaki KIMURA and Masanori TSUJIl

oblong, 0.7-1.1 cm long and up to 6 mm
wide at base, with obtusely pointed orrounded apex; laminae connected atbase; margins entire. Midvein sinuous,
sending off 3-5 pairs of twice or once
forked lateral veins.
Distribution and occurrence. - Cladophlebis sp. C is rare and known only fromthe Shinadani Formation.
Remarks.-Our pinnules are characterized by their varied forms with a sinuous
mid vein and 3-5 pairs of twice or once
forked lateral veins, and prominently
contiguous laminae at their basal part.
Cladophlebis grabauniana and C. suniana
originally described by P' AN (1936) and
by SZE (1956) respectively from the Upper
Triassic of Shaanxi show the venation
similar to C. sp. C. But Cladophlebis
grabauniana is distinguished from C. sp.
C by its small oblong pinnules with astraight mid vein, and C. suniana by its
oblong pinnules with mostly simple lateralveins.
Sterile fern leaves described by TESLENKO (1970) as Cladophlebis nebbensis from
the Middle Jurassic of Southern Siberia
resemble ours in the form of pinnule andvenation.
But TESLENKO'S leaves areeasily distinguished by their pinnules

with finely denticulated margins.
Our leaves appear to be new to science,but here we reserve their specific identity
because of their fragmental state.

References
BARANOVA, Z. E., KlRITCHKOYA, A. I. and
ZACER, V. V. (1975): Stratigraphy andflora from the Jurassic of the easternpart along the Caspian Basin.
Bull.
V.N.I.G.R.I., vol. 332, p. 1-191, incl. pIs.
1-34 (in Russian).
CHo\\', T. Y. and CIIAl'G, S. J. (1956): On
the discovery of the Yenchang Formationin Alashan Region, Northeastern China.
Acta Palaeont. Sinica, vol. 4, no. 1, p.
53-60, pI. 1 (in Chinese).
FENG, S.N., ME;-;G, F.S., CHEN, G.X., XI,
Y. H., ZHANG, C. F. and Lw Y. A. (1977) :Palaeontological Atlas oj Middle-South
China. Part 3: Mesozoic-Cenozoic. p.
195-262, pIs. 70-107 (plants). Di-Zhi-ChuBan-She, Peking (in Chinese).
HARRIS, T. M. (1931): The fossil flora of
Scoresby Sound East Greenland. Medd.
om Gr1mland, Bd. 85, nr. 2, p. 1-104, pIs.
1-18.
- - (1961): The Yorkshire Jurassic flora.
I. Thallophyta-Pteridophyta. ix+212 p.
Brit. Mus. (Nat. Hist.).
- - (1980): The Yorkshire Jurassic fern

Explanation of Plate 31
Figs. 1-2., Thaumatopteris elongata OISlII; Poorly preserved pinnae, both x L I . Reinserted
from KIMURA, 1959a, pl. 2, fig. 2 (A-3034). 2. (Kr-150; Text-fig. 5b).
Figs. 3-4. Cfr. Asterotheca okajujii KI~I[;RA and OHANA; both xL 3. A sterile pinna fragmentwith large-sized pinnules (NNW -061; Text-fig. 7a). 4. Apical part of a sterile pinna, withsmall-sized pinnules (DAI-009; Text-fig. 7d).
Figs. 5-6. Cfr. Todites jukutomii Kl\ICRA and OHANA; both xL 5. A part of sterile leaf (NE031; Text-fig. 8a). 6. A pinna fragment (NNW -660).
Figs. 7-8. Sphenopteris sp. A; both x 2. 7. A part of fertile leaf (NNW -282; Text-fig. lOa).
8. Ditto (NNW -833).
Fig. 9. Cladophlebis kurumensis KI~IURA and TSUJI! sp. nov.; a pinna fragment (Kr-066, paratyp~), xL
Fig. 10. Cladophlebis pseudodelicatula OISHI; a pinna fragment (A-0309), xL
Fig. 11. C12dJ/)hI2bis racib~rskii ZEILLER; a pinna fragment (A-0496), xL

KIMURA and TSUjII:

Early Jurassic plants

Plate 31

737.

Early Jurassic plants

Phlebopteris braunii (GOEPPERT) and its
reference to Matonia R. BR. Bull. Brit.
Mus. (Nat. Hist.), Ceol., vol. 33, no. 5,
p. 295-311.
HE, Y.L., Wu, S. Y., Wu, S. W., LI, P.C., LI,
H. M. and Keo, S. X. (1979): Palaeontological Atlas of Northwest China. Volume
of Qinghai. Part 2. p. 129-167, pIs. 5082 (plants). Di-Zhi-Chu-Ban-She, Peking(in Chinese).
HsU, J., ZIIU, J. N., CHEN, Y., DUAN, S. Y.,
Hu, Y. F. and ZIIU, W_ Q. (1979): Late
Triassic Baoding flora of China. 130 p.,
75 pIs. Ke-Xue-Chu-Ban-She, Peking (inChinese) .
KA \\' ASAKI, S. (1925): Some older Mesozoic
plants in Korea. Bull. Ceol. Surv. Chosen(Korea), vol. 4, pt. 1, p. 1-71, pIs. 1-47.
- - (1926): Addition to the older Mesozoic
plants in Korea. Ibid., pt. 2, p. 1-35, pIs.
1-11.
- - (1939): Second addition to the older
Mesozoic plants in Korea. Ibid., pt. 3, p.
1-69, pIs. 1-16.
KI:\IURA, T. (1959a): Mesozoic plants fromthe Iwamuro Formation (Liassic), Tonegun, Gumma Prefecture, Japan.
Bull.
Sen. High Sch., Tokyo Univ. Educ., no.
3, p. 1-59, incl. pIs. 1-12.
- - (1959b): Mesozoic plants from the Kotaki
Coal-Field, the Kuruma Group, Central
Honshu, Japan. Ibid., p. 61-83, incl. pIs.
1-2.
KI:\IURA, T. and OIIA:"A, T. (1980): Somefossil ferns from the Middle Carnic
Momonoki Formation, Yamaguchi Prefecture, Japan.
Bull. Natn. Sci. Mus.
(Tokyo), ser. C, vol. 6, no. 3, p. 73-92,
pIs. 1-6.
KI:\IURA, T. and TSI:JII, M. (1980a): Early
Jurassic plants in Japan. Part 1. Trans.
Proc. Palaeont. Soc. Japan, N. S., no. 119,
p. 339-358, pIs. 38-40.
- - and - - (1980b): Ditto. Part 2. Ibid.,
no. 120, p. 449-465, pIs. 54-56.
KON'NO, E. (1968): Some Upper Triassic
species of Dipteridaceae from Japan and
Borneo. Journ. Linn. Soc. (Bot.), vol. 61,
no. 384, p. 93-105, pIs. 1-2.
- - (1972): Some late Triassic plants from205

the southeastern border of Sarawak,
East Malaysia. Ceol. Palaeont. SE-Asia(Tokyo Univ. Press), vol. 10, p. 125-178,
pIs. 24-32.
KRASSER, F. (1906): Fossile Pflanzen aus
Transbaikalien der Mongolei und Manchurei.
Akad. Wiss. Wien Denkschr., '-01. 78, p.
589-634, pIs. 1-4.
KRASSILOV, V. A. (1978): Mesozoic lycopods
and ferns from the Bureja Basin. Palaeontographica, Abt. B, Bd. 166, Lief. 1-3,
p. 16-29, pIs. 1-11.
KRYSHTOFOVICH, A. N. (1923): Pleuromeia
and Hausmannia in Eastern Siberia, withsummary of recent contribution to thepaleobotany of the region. Amer. Journ.
Sci., ser. 5, vol. 5, no. 27, p. 200-208,
figs. 1-7.
LEE, H. H. (LI, X. X.) (1955): On the age of
the Yunkang Series of the Tatung CoalField in North Shensi. Acta Palaeon!.
Sinica, vol. 3, no. 1, p. 25-46, pIs. 1-2 (inChinese) .
LEE, P. C. (LI, P. J.) (1964): Fossil plantsfrom the Hsuchiaho Series of Kwangyiian,Northern Szechuan.
lvIem. Inst. Ceol.
Palaeont., Acad. Sinica, no. 3, p. 101-178,
pIs. 1-20 (in Chinese).
LI, P.J., TSAO, C.Y. and We, S.C. (1976):
Mesozoic plants fromYunnan.
In'Mesozoic fossils of Yunnan, vol. 1'. p.
87-160, pIs. 1-47. Ke-Xue-Chu-Ban-She,Peking (in Chinese).
OISHI, S. (1930): Notes on some fossil plantsfrom the Upper Triassic Beds of Nariwa,
Provo Bitchu, Japan. Jap. Journ. Ceol.
Ceogr., vol. 7, no. 2, p. 49-58, pI. 7.
- - (1931): Mesozoic plants from Kita-Otari,
Provo Shinano, Japan. Journ. Fac. Sci.,
Hokkaido Imp. Univ., ser. 4, vol. 1, no.
2, p. 223-255, pIs. 16-18.
- - (1932): The Rhaetic plants from the
Nariwa District, Provo Bitchu (Okayama
Prefecture), Japan. Ibid., vol. 1, nos. 34, p. 257-380, pIs. 15-53.
- - (1940): The Mesozoic floras of Japan.
Ibid., vol. 5, nos. 2-4, p. 123-480, pis. 148.
OISHI, S. and HUZIOKA, K. (1938): Fossil
plants from Nariwa. A supplement. Ibid.,

206

Tatsuaki KIMURA and Masanori TSUJIIvol. 4, nos. 1-2, p. 69-101, pis. 7-12.
OISHI, S. and TAKAHASI, E. (1936): Rhaetic
plants from Provo Nagato, Japan. A supplement. Ibid., vol. 3, no. 2, p. 114-133, pI.
10.
OISHI, S. and YA~'IASITA, K. (1936): On thefossil Dipteridaceae. Ibid., vol. 3, no. 2,
p. 135-184.
P' AN, C. H. (1936): Older Mesozoic plantsfrom North Shensi. Palaeont. Sinica, ser.
A, vol. 4, fasc. 2, p. 1-49, pis. 1-15.
PRYKADA, B. (1938): Jurassic flora of the
Emba
Region.
Problems Palaeont.,
Moscow, vol. 4, p. 363-404, pis. 1-5 (inRussian).
SCHOROCHOVA, S.A. and SREI3RODOLSKAJA, LN.
(1979): New late Triassic plants from
Primorye. In 'Far-Eastern Palaeofloristics'.
Proc. Bioi. Pedol. Inst., Far-Eastern Sci.
Centre, Acad. Sci. USSR, N. S., vol. 53(156), p. 68-75, pis. 11-16 (in Russian).
SIXTEL, T. A. and KI-IUDAlI3ERDYEV, R. Kh.
(1968): Floras of the past of Middle
Asia. In 'The Palaeobotany of Uzbekistan
I'. Inst. Bot., Acad. Sci. Uzbek. SSR. p.
3-87, pis. 1-32. FAN, Tashkent.
STANISLAWSKI, F. A. (1971): Fossil flora andstratigraphy of the Upper Triassic of
Donbass. 139 p., 36 pis. Inst. Geol. Sci.,
Acad. Sci. USSR, Kiev (in Russian).
SREI3RODOLSKAJA, LN. (1961): On the Family
Dipteridaceae in the Middle Keuper floraof the South Primorye. Journ. Palaeont.,
Acad. Sci. USSR, 1961-1, p. 144-150, pis.

16-17 (in Russian).
SZE, H. C. (1931): Beitrage zur liassischen
Flora von China. Mem. Natll. Res. Ins/.
Ceol., Acad. Sinica, no. 12, p. 1-85, pis.
1-10.
- - (1933a): Beitrage zur mesozoischen
Flora von China. Palaeont. Sinica, ser.
A, vol. 4, fasc. 1, p. 1-68, pis. 1-12.
- - (1933b): Jurassic plants from Shensi.
Mem. Natn. Res. Ceol., Acad. Sinica, no.
13, p. 77-86, pis. 11-12.
- - (1949): Die mesozoischen Flora aus der
Hsiangchi kohlen Series in West Hupeh,
China. Palaeont. Sinica, N. S., ser. A,
no. 2, p. 1-71, pis. 1-15.
- - (1956): Older Mesozoic plants from the
Yenchang Formation, North Shensi. Ibid.,
vol. 5, p. 1-111 (Chinese part), 110-217(English part), pis. 1-56.
SZE, H. C. and HsU, J. (1954): The standard
fossils in China (plants). 99 p., 69 pis.
Di-Zhi-Chu-Ban-She, Peking (in Chinese).
SZE, H.C., LI, X.X., LI, P.J., Wu, S.C.,
ZHOU, T. Y., YE, M.N. and SI-IEN, G.L.
(1963): Fossil plants of China. Vol. 2.
Mesozoic plants of China. iv+429 p., 118
pis.
Ke-Xue-Chu-Ban-She, Peking (inChinese).
TESLE:-IKO, Y. V. (1970): Jurassic stratigraphy
and flora of Western and Southern Siberia
and Tuva. Bull. Sibr. Sci. Stud., Inst.
Ceol. Ceophys. Min. Res., ser. Palaeont.
Stratigr., vol. 42, p. 1-216, pis. 1-52 (inRussian) .

Explanation of Plate 32
Fig. 1. Cfr. Todites yamanoiensis (YOKOYAMA) KIMURA and OI-I.;\!\A; a sterile pinna fragment(A-0405; Text-fig. 9b), xL
Fig. 2. Cladophlebis kurumensis KIMURA and TSUJI[ sp. nov.; a pinna fragment (Kr-052, holotype), xL
Figs. 3-4. Cladophlebis pseudodelicatula OISIIl; both xL 3. A part of leaf, with a thick rachis,reinserted from KIMURA, 1959a, pI. 5, fig. 2 (A-4068; Text-fig. 12a). 4. Ditto (NEE-099;
Text-fig. 12b).
Figs. 5-9. Cladophlebis raciborskii ZEILLER; all xL 5. A part of leaf, reinserted from KIMURA,
1959a, pI. 5, fig. 4 (A-4038). 6-9. Pinna fragments. 6; (A-4006), 7; (A-4016, counterpart of
KI:\WRA, 1959a, pI. 5, fig. 6 regarded formerly as C. raciborskii forma integra), 8; (TOB046A), 9; (NE-015).
Fig. 10. Cladophlebis sp. C; a part of leaf (SHI-0l5; Text-fig. 14), xL

KIMURA and TsuJI!:

Early Jurassic plants

Plate 32

737.
V.\KHRA~[EE\"Early Jurassic plants

V. A., VL\Bl:\[[ROV[CH, V. P.,
DOBRCS[(['iA, I. A., KRASS[LOV, V. A.,
LE13EDEV, E. L. and SR[CURODOLS[(AJA,
I. N. (1980) : Mesozoic higher spore.bearing
plants in USSR. 122 p. Nauk, Moscow(in Russian).
VASS[LEVSKAJA, N. D. (1967): Early Cretace·ous ferns from the Lena·Olenek District,in the Lena Basin. Sci. Rep., Inst. Arct.
Ceol., lvIin. Ceol. USSR, Proc. Palaeont.
Biostratigr., vol. 17, p. 58-78, pis. 1-8(in Russian).
YA13E, H. (1922): Notes on some Mesozoic
plants from japan, Korea and China, inthe collection of the Institute of Geology
and Palaeontology of the Tohoku Imperial
University. Sci. Rep., Tohoku Imp. Univ.,
2nd. ser., vol. 7, no. 1, p. 1-28, pis. 1-4.
Y,\13E, H. and 0[5[[1, S. (1933): Mesozoic
plants from Manchuria. Ibid., vol. 12, no.
2B, p. 195-238, pis. 30-35.
Y A:'\G, X. H. (1978): Palaeontological Atlasof Southwest China. Volume of Sichuan.
Part 2. Division of plants. p. 469-684,

207

pis. 155-170. Di-Zhi-Chu-Ban-She, Peking(in Chinese).
Y OKo)' A~-IA, M. (1889): jurassic plan ts from
Kaga, Hida, and Echizen. JOUr/I. Coil.
Sci., Imp. Univ. Japan, vol. 3, pt. 1, p.
1-66, pis. 1-14.
---- (1906): Mesozoic plants from China.
Journ. Coil. Sci., Imp. Univ. Tokyo, vol.
21, art. 9, p. 1-39, pis. 1-12.
ZE[LLE[~, R. (1903):
Flore fossile des gites
de Charbon du Tonkin. viii +328 p., 56
pis. (Etud. Gites min. Fr.) Paris.
ZIIOC, H.Q., Qc, L.F. and ZIIA}\;G, W.P.
(Geol. Inst., Chinese Acad. Geol. Sci.)(1980): Mesozoic stratigraphy end fossils
in the Shaan-Can-Ning Basin. vol. 1. 212
p., 98 pis. Di-Zhi-Chu-Ban-She, Peking(in Chinese).
ZI[OC, T. S. (1978): On the Mesczoic coalhearing strata and fossil plants from
Fujian Province.
Prof. Pap. Stratigr.
Palaeont., Chinese Acad. Ceol. Sci., no.
4, p. 88-134, pis. 15-30 (in Chinese).

Geographical names in japan: Asahi·machi ijiJl ElIIIL
Dairagawa :*3JZJII,
I-ligashi-Neiridani
JlUJ!:A:)§,
Iwamuro ;(,\~,
Kita-Otari ~t'H'~,
Kotaki 'NiiL
Kuruma *Xl~,l\1omonoNagato *1"],
Negoya iJ;!))'i:fi,
Nariwa )Vi:;J~,
Neiridani ~'lA.fr,
Nishiki tJE/
Neiridani E§~A:)§,
Oguchi}{; I],
Sasakomata W+i'k,
Shinadani f,fc,~~,Tobiiwadani
11l'5:)§,
Tsuchizawa (Tsunzawa) ±iR,
Yamanoi ili!lrHI:;
Geographical names in China(with former expression in parenthesis) : Alashan rrnrrv:t.j£,
Baoding ~iJfl:,
Fang-zi (Fangtzi) to'f-,
Fujian 7ffi~,
Guangdong $]:1'[.
Guangyuan (Kwangytian) ).tJ[:,
Hubei(Hupeh) llj1]~t,~ jiangxi (Kiangsi) iIf1lj,
Neimeng I-".J~,
Qinghai Wiffi,
Shandong(Shandung) LU.!lL
Shaan-Gan-Ning f»\!tn~,
Shaanxi (Shensi) f»\!E§,
Shanxi (Shansi)
IlJE§,
Sichuan (Szechuan) jl9J11,
Xiangxi (Hsiangchi) ~£,Xujiahe (Hsuchiaho) ~H{
inf,
Yanchang (Yenchang) Ylf*,
Yungang (Yunkang) '~i~,Yunnan -,;slW,
Yushugoufiti<11itiil'j;
Geographical name in Korea (with former expression in parenthesis): Daedong(Taedong) :*FiU*,El *O)f}].Jtil:/--:1. =jff:~jl([!jfr.}itlio :f:0) 3 : mjf~il1V=-,,)-0~, *.J!l~J~.!ff.:ro J: lY1':·:'i.f~\hiil.p ~.-l")~0)
Co' ?(oo!jfr.}ltE~i(c'llJi: L 1.:.0"< l- ='- 7f-+ : Phlebopteris sp. A,
f~l.i'vGUf+: Coniopteris neiridaniensis KIMliRA
and TSUJI! sp. nov., -\",~;h/.J;-~ '5 !?f'iLH : Clathropteris meniseoides BRO:'\(;I'\[ART,
Dictyophyllum kotakiense K[~IClu\ and TSCJI! sp. nov., Thaumatopteris elongata O[SIII,
Hausmannia (Protorhipis) sp. A, fl:1l€ 1(, L <,t. ~rM1:O) (ft[i/fJl)j 0) ' / ",' : Cfr. Asterotheca okafujii KI:lluRA and OI-lA:,\A. Cfr. Todites fukutomii KI\IUL\. and OllA:'\A,
efr. Todites yamanoiensis (YO"OYA~[A) K[:I[CRA and OIlAI'\A, Sphenopteris sp. A,
Cladophlebis kurllmensis KI:I[LRA and TSuJII sp. nov., C. pseudodelicatula OISII[, C.
raehiborskii ZE[LLER, C. sp. A, C. sp. B, C. sp. C.

Co'fOO:!jfr.}O)Wi5X;/.J,!?, -=:'O)l1i'i!jfr.}ltr(t., l'vvP:Q, ~~;h/.J~~ '5 !?i'iLM!jfr.}lW-c·S:Q/.J~, t~/J'
hGLYf"rO)Co'",':roJ:U Ptilophyllum ("'*7'-7-"'1" f&j!t) ~{l;-t,'o-=:.O)U::ft., ,"311'J'O)jn'[!jfr.}.itl!J!f.!.1K 0) r.~:m ~ :15.~~-t :Q1: -C'1f(~i fA: ~N ,l::t,J:0;;t(t,t;! riJJ • iJ:Jl:IEHlJ
0Trans. Proc. Palaeont. Soc. Japan, N. S., No. 124, pp. 208-214, pIs. 33, 34, December 30, 1981

738.

A NEW PERMIAN RADIOLARIAN GENUS FROM THE
T AMBA BELT, SOUTHWEST J AP AN
A TSUSHI T AKEMURA
Department of Geology and Mineralogy, Faculty of Science,
Kyoto University, Kyoto 606, Japanand
KOJIRO NAKASEKO
Institute of Geological Sciences, College of General Education,
Osaka University, Toyonaka 560, Japan

Abstract. An Upper Permian radiolarian assemblage which includes more than
10 species has been found in the Tamba Belt, Southwest Japan. A new genus Neoalbaillella belonging to the family Albaillellidae DEFLAKDRE emend. HOLDSWORTH is
proposed from this assemblage. Genus Neoalbaillella is characterized by its curved
apical cone, winged pseudothorax and pseudoabdomen with windows and trabeculae.
4 species (of which 2 are new) belonging to this new genus are described.

new genera, Pseudoalbaillella and Parafollicucullus, and established nine radiolarian biozones for an interval from theUpper Devonian to Permian. In japan,
ISHIGA and IMOTO (1980) carried out adetailed biostratigraphic study on the
Pseudoalbaillella Assemblage (HOLDSWORTH and JONES, 1980) from a chert in
the Tamba Belt, Southwest japan.
In our own study, T AKEMURA found awell-preserved radiolarian assemblage including more than 10 species from theTamba Belt. Most of the AlbaiUellarians
which have been reported have imperforate shells. New radiolarians we reportin the present paper have shells withcuriously shaped wings and windows arranged in several transverse lines. We
propose for them a new name, Neoalbaillella which belongs to the family Albail-

Introduction
An interesting form was found by
DEFLANDRE (1952) from Carboniferous
deposits of Europe, which was described
as Albaillella. It is characterized by acone-shaped chamber with an open baseand a very marked bilateral symmetry of
the frame. Recently, research on radiolarian remains from Paleozoic strata has
increased a great deal in connection withbiostratigraphic study by means of radiolarians. Consequently, forms related to
Albaillella were found in several placesin the world.
The Permian radiolarian genus, Follicucullus was described by ORMISTON andBABCOCK (1979) from the Guadalupian
Lamar Limestone of West Texas. HOLDSWORTH and jOl'IES (1980) proposed two

208

738.

A New Permian Rad'iolarian Genus

lellidae DEFLANDRE, emend. HOLDSWORTH.
We wish to express our sincere thanks
to Professor Keiji NAKAZA WA of the
Department of Geology and Mineralogy,
Faculty of Science, Kyoto University andMiss Akiko NISHIMURA of the Instituteof Geological Sciences, College of General
Education, Osaka University for theirvaluable advice and kind assistance. We
also thank Dr. Nobuhiro IMOTO and Mr.
Hiroaki ISHIGA of the Department of Earth
Sciences, Kyoto University of Education
for their valuable assistance. We areparticularly grateful to Professor Tsunemasa SAITO of the Institute of Earth
Sciences, Faculty of Science, Yamagata
University for reading manuscript.

209

light gray bedded chert with rrany quartz
veins. One bed of the chert is about 10
cm thick and eight beds are exposed at
this outcrop.
We used a dilute hydrofluoric acid (about1%) and immersed samples in it for 12-24 hours. Delicate parts of radiolarians,
including wings or spines of Neoalbaillella,
would have been destroyed by a moreconcentrated acid. Residues obtained bythat process were sieved (200 mesh), picked
up by a thin brush and observed. We
also observed etched surfaces of the rocksamples (PI. 33, Figs. 4a, b). In this case,an acid of about 10% concentration wasused.

Radiolarian AEsemblage and its Age
Geological Setting, Materialand Method
The Tamba Belt is located in the Innerzone of Southwest Japan (Text-fig. 1) andconsists of Mesozoic and Paleozoic strata
which are mainly made up of sandstone,
shale. chert and basic volcanic rocks.
These strata have been dated as ranging
from late Paleozoic to Jurassic mostly
based on fusulinids occurring in a lenticallimestone, conodonts in a chert and radiolarians in chert and shale (SAKAGUCHI,
1961; Tamba Belt Research Group, 1979,
etc.).
T AKDIURA (1980) surveyed the southernpart of Kameoka City in Kyoto Prefecture,
located in the southern part of the Tamba
Belt. and recovered late Paleozoic to Jurassic radiolarians from 13 sites in the area.
The fossil locality which is discussed by
us in the present paper is AT26 situatedat about 300 m east of Mt. Kurogara-dake(Text-fig. 1). Locality A T26 is a small
outcrop within a 300-500 m thick, massiveor bedded chert sequence.
Samples from A T26 are composed of a

The radiolarian assemblage from A T26
includes more than 10 species, but mostof the species have structures which havenot been described so far. They are
identified as : Palaeoactinommids or Entactiniids, Angulobracchia(?) sp., Paronaella(?)
sp., Follicucullus ventricosus ORMISTON andBABCOCK, Follicucullus scholasticus ORMISTON and BABCOCK, Albaillellidae with
one spine, Neoalbaillella o rnitlwfo r17l is, n.
gen. n. sp., Neoalbaillella gracilis, n. gen.
n. sp. Neoalbaillella sp. A and Neoalbaillella sp. B.
Angulobracchia (?) sp. (PI. 34, Fig. 11)
as rrentioned above has two externalbean-.s along the axis of rayon both top
and bottom sides of each ray. Paronaella (?) sp. (PI. 34, Fig. 12) reserr bles
Paronaella sp. A of BAUMGARTNER (1980)
in shape, but rays are imperforate except
tips and have spines. Internal structuresof Angulobracchia(?) sp. and Paronaella(?)
sp. have not been observed.
Albaillellidae (PI. 34, Fig. 9-10) has one
spine on the ventral side and two rods
or flaps under its a~erture. The ventral
rod is weak and short, but the dorsal one

210

Atsushi TAKEMURA and Ko}iro NAKASEKO

is strong and platy. Its internal structure
is indeterminate.
From this site, no conodont has beenfound. This assemblage includes, however,Follicucullus ventricosus (PI. 34, Fig. 7) andFollicucullus scholasticus (PI. 34, Fig. 6)
and is devoid of Pseudoalbaillella andParafollicucullus. Following HOLDSWORTH
and JONES (1980), the Follicucullus Assemblage is dated to range from Wordian toupper Guadalupian and they state: "Pseudoalbaillella and Parafollicucullus probably
disappeared before latest Guadalupian
time." Also, in the Sasayama area of
ISHIGA and IMOTO (1980), the Follicucullus
Assemblage is younger than the Pseudoalbaillella Assemblage, although Neoalbailleila has not been reported there. So, theage of this "Neoalbaillella Assemblage"
is late Permian, probably Guadalupian oryounger.

Systematic Paleontology
Subclass RADIOLARIA MULLER, 1858
Order POL YCYSTINA EHREXBERG,
1838, emend. RIEDEL, 1967
Suborder ALBAILLELLARIA DEFLANDRE,
1953, emend. HOLDSWORTH, 1969
Family ALBAILLELLIDAE DEFLANDRE,
1952, emend. HOLDSWORTH, 1977
Genus Neoalbaillella T AKEMURA and
NAKASEKO, new genus

Text.fig. 1. Locality map (site plotted on Geographical Survey Instituteof Japan topographical map "Hoki" of scale 1: 25000).

738.

A New Penn:ian Radiolarian Genus

Type-species: Neoalbaillella omithofOrlnis T AKENlURA and NAKASEKO n. sp.
Diagnosis :-Nearly bilaterally symmetrical, conical or subcylindrical shells of
which upper part is winged and imperforate and of which lower part has windows.
The lower part of the shell possessing
trabeculae and windows arranged in transverse lines, but the upper part having an
unknown internal structure. Apical cone
bended in a bilaterally symmetrical plane.
Two longitudinal rods with three or morespines lying at the dorsal and ventral
sides of the lower shell part under thewings and protruding posteriorly fro:naround the aperture.
Remarhs:- The apical cone of the shellbelonging to this new genus is more orless curved. We consider the outer sideof this curve as a dorsal surface and theinner side as a ventral one. Each surface
has a wing and a longitudinal rod (Textfig. 2). The dorsal rod is more or lessstronger than the ventral one. When theapical cone can be observed at the upper
part of the shell with the cone curving
to the left, we are seeing the "left side"of the shell. If the cone curves to theright, we are seeing the "'right side".
In this new genus, there are neither
distinctive segmentations nor constrictions.
The upper part of the bo::ly, however, maycorrespond to a combination of an apicalcone and a pseudothorax of Pseudoalbaillella HOLDSWORTH and JONES, and thelower part to a pseudoabdomen.
The overall shape of the shell resembles that of Pseudoalbaillella HOLDSWORTH
and JONES, but the latter genus and other
Paleozoic genera belonging to Albaillellaria are not known to possess windows(DEFLANDRE, 1952; FORE:VIAN, 1963; ORMISTON and BABCOCK, 1979; HOLDSWORTH
and JONES, 1980). The trabeculae in thelower part of the shell of this genus (PI.
33, figs. 4a, b) may indicate affinity of this211

new genus to the Albaillellidae DEFLAKDRE
but we have seen no transverse cross bar.
Because of the possession of trabeculae
and windows, therefore,
Follicullidae
ORMISTON and BABCOCK is not suitable
to assign this genus.
These windows forming transvese lines
in the lower part of the shell seem tohave some similarity to the Triassic
Dictyomitrella and other Cyrtoids, Inorder to understand the taxonomy of
Paleozoic and Mesozoic radiolarians adequently, it is very important to elucidate
relationships of these resembling taxa.
Neoalbaillella ornithofonnis TAKE:vIURA

and NAKASEKO, n. sp.
PI. 33, Figs. 1-6; Text-fig. 2

Description :-Pseudothorax and pseudoabdomen cylindrical, with four or five,rarely more than six transyerse lines ofwindows.
Wings triangular in shape,
platy and bilaterally asymmetrically attached to the main body of the shell, but
two wings symmetrical to each other withrespect to a longitudinal two-fold axis
through the center of the body. Tworods, X-rod and V-rod, lying along theanterior sides of the two wings. The Xrod situated on the left side of the dorsal
wing and the right side of the ventral
one, and V-rod conversely the right sideof the dorsal wing and the left side of
the ventral one. Two wings bordered attheir upper edges by V-rods which areslightly curved posteriorly. X-rods extended in straight lines from the justside of Y-rods to a little belo\v Y-rods.
Trabeculae circular and lying on the innersurface of the shell wall. Dorsal andventral rods arising under wings.
Measurements :-Length of shell (exclusive of dorsal and ventral rods), 190-230p.m; Width of shell (including t\\'o wings),

Atsushi TAKEMURA and ]{ojiro NAKASEKO

212-'-:0\

x : ' '",,,\

v.w.

a.c.

d.w.

~oo~~-~

U P_ps_.t._. _
.
ps.a.

wd.

I.p.

v.r.

d.r.
10011

Text-fig. 2_

Neoalbaillella ornithoformis T AKEMURA and NAKASEKO n_ sp_(left side of the shell)
u. p. : the upper part of the shell
L p. : the lower part of the shella. c: apical cone
ps. t.: pseudothorax
ps. a. : pseudoabdomen
v. w. : ventral wing
d. w. : dorsal wing
x: X-rody: Y-rod
wd.: windows: spine
v. r. : ventral rod
d. r. : dorsal rod
a: aperture

Explanation of Plate 33(scale = 100 pm)
1-6. Neoalbaillella ornithoformis n. sp_
1. Holotype (left side) ATPMTB-ll03-1
2. Para type (left side) A TPMTB-ll03-2
3. Para type (left side) A TPMTB-1201-1
4a, b. Paratype (left side, stereophotograph) ATPMTB-llO~-1
5. Paratype (right side) ATPMTB-1201-2
6. Para type (right side) ATPMTB-1204-1
7-10. Neoalbaillella gracilis n. sp.
7. Holotype (right side) ATPMTB-1203-1
8. Paratype (left side) ATPMTB-1202-1
9. Para type (left side) ATPMTB-1203-2
10. Para type (right side) ATPMTB-1203-3

TAKEMURA and NAKASEKO:

A New Permian Radiolarian Genus

Plate 33

738.

A New Permian Radiolarian Genus

210-270 ,um; Diameter of the lower partof the shell. 80-95 ,urn.
Remarks:- This species differs fromother species of Neoalbaillella in that distinctive feature of the wings. Observation of a dissolved specimen seems toindicate that one wing comprises a combination of at least four rods including
X- and Y-rods. The species name, ornithoformis, is given because of the birdshaped appearance of this species.
Occurrence :-Upper Permian in theTamba Belt.

Neoalbaillella gracilis T AKEMURA

213

Description :-Apical cone inflated anddorm-shaped with apical horn. Pseudothorax and pseudoabdomen cylindrical,
possessing four to five transverse linesof windows. Wings bilaterally symmetrical, simple and slender. Dorsal rod
stronger than ventral one.
Remarks:- Though the wings of thisspecies are similar to that of Neoalbaillellagracilis n. sp., this species is distinguishedfrom other species by the shape of theapical cone. No species name is given
because of its poor state of preservationand scarcity.
Occurrence :-Upper Permian in theTamba Belt.

and NAKAsEKo, n. sp.
PI. 33, Figs. 7-10; PI. 34, Fig. 1

Neoalbaillella sp. B.

Description :-Apical cone slender, pseudothorax and pseudoabdomen long andcylindrical. Wings slender and possessing
X- and Y-rods, with both rods together
bordering the upper edge of the twowings; wings being bilaterally symmetrical and simple. Windows forming morethan five or six transverse lines but thoseof upper two or three lines being relict.
Dorsal rod stronger than ventral one.
Measurements :-Length of shell (exclusive of dorsal and ventral rods), 260280 ,urn; Width of shell (including two
wings), 180-200 pm; Diameter of the lower
part of the shell, 75-90 pm.
Remarks:- This species differs from
Neoalbaillella ornithoformis, n. sp. in having the bilaterally symmetrical wings andslender body and wings. The species
name, gracilis. means slender.
Occurrence :-Upper Permian in the
Ta:nba Belt.

PI. 34, Figs. 4-5

Neoalbaillella sp. A
PI. 34, Figs. 2-3

Description :-Pseudothorax trapezoidal,
pseudoabdomen trapezoidal or barrelshaped; body conical or spindle-shaped.
Windows forming more than six lines
and relict in upper two or three lines.
Platy wings and lower three or more
platy spines complexly connected withlongitudinal platy rods.
Remarks:- This species is distinguishedfrom other species of Neoalbaillella by its
complicated wings. The body shape of
this species resembles that of Mesozoic
Nasselarians, but at present there is noevidence to substantiate their relationship.
Occurrence :-Upper Permian in theTamba Belt.
References
P. O. (1980): Late Jurassic
Hagiastridae and Patulibracchiidae (Radio·
laria) from the Argolis Peninsula (Peloponnesus, Greece).
Micropaleontology,
vol. 26, no. 3, p. 274-322, pis. 1-12.
DEFLAl'ORE, G. (1952): Albaillella nov. gen.,Radiolaire fossile du Carbonifere inferieur,
BAu\IGART:-':ER,Atsnshi 1'AKEMURA and Kojiro NAKASEKO

214

type d'une lignee aberrante eteinte. Acad.
Sci. Paris, Comptes Rendus, vol. 234, p.
872-874, figs. 1-9.
FORE'VIA0:, I-I. P. (1963):
Upper Devonian
Radiolaria from the Huron member of the
Ohio shale. Micropaleontology, vol. 9, no.
3, p. 267-304, pIs. 1-9.
HOLDSWORTH, B. K. (1969): The relationshipbetween the genus Albaillella DEFLA~DRE
and the ceratoikiscid Radiolaria. Micropaleontology, vol. 15, no. 2, p. 230-236,
pI. l.
- - (1977): Paleozoic Radiolaria: Stratigraphic distribution in Atlantic Borderlands.
in Swain, F. M., ed., Stratigraphic micropaleontology of Atlantic Basin and borderlands: Amsterdam, Elsevier, p. 167184.
HOLDSWORTH, B. K. and JONES, D. L. (1980) :Preliminary radiolarian zonation for Late
Devonian through Permian time. Geology,
vol. 8, p. 281-285.
iSIIIGA, H. and INiOTo, N. (1980): Some Permian Radiolarians in the Tamba District,
Southwest japan. Earth Science (ChikYll

Kagakll) , vol. 34, no. 6, p. 333-345, pIs.
1-5.
OR~IISTON, A. and BABCOCK, L. (1979) : FolliCUCUlillS, new radiolarian genus from the
Guadalupian (Permian) Lamar Limestoneof the Delaware Basin. jOllrnci of Pcieontology, vol. 53, no. 2, p. 328-334, 1 pI., 2
text-figs.
SAKAGUCHI, S. (1961):
Stratigraphy andPalaeontology of the South Tamba District,
Part 1, Statigraphy. Mem. Osaka Gakllgei
Univ., B, vol. 10, p. 35-76.
TAKEl'vIURA, A. (1980): The Paleo-Mesozoic
system of the Tamba Belt in the southernpart of the Kameoka City, Kyoto Prefecture, japan. News of Osaka Micropaleontologists, no. 8, p. 22-31, pIs. 1-2. (injapanese)
Tamba Belt Research Group (1979): Paleozoic
and Mesozoic Systems in the Tamba Belt
(Part 5) -Permian and Triassic conodont
fossils in the northwestern hills of Kyoto
City. Earth Science (Chik)'ll Kagaku),
vol. 33, p. 247-254, pI. l. (in japanesewith English Abstract).

fttlil*t;'Gj£llnJ::~/vA*cjjlzllX£t!.{I::fiO)~~: g~Jf.ffUlIi\]m!.l'\Mffi t=- (1 7-'\' -~ t;;n';fp-to)"I't;'G Albaillellidae f3H=-~-r::5 NeoalbaillellaJi.&~~~~ct-::o .::.0)Ji.&11, ~rtjlCt-:: apical cone, JiO);/;::5 pseudothorax, window O);1,.G;/1,::5 pseudoabdomen ~t;f00)t;q','fj!!lc(;/;::5o :.:. --((1, *J;~t:.Jm-r::5 4 fill (I''], 2 :fllHUl'
f'!ll) ~Ri3JI~ Lt-::o'r'rHilj[lij. '-I'11I:il¥<1:0B
C,~/vA*cO)!&flX£t!.lWt:R~gjHI:,,-r::5oExplanation of Plate 34(scale=100 pm)
1. Neoalbaillella gracilis n. sp.
Paratype (left side) A TPMTB-1203-4
2-3. Neoalbaillella sp. A
2. A TPMTB-1203-5
3. A TPMTB-1204-2
4-5. N eoalbaillella sp. B
4. A TPMTB-1203-6
5. A TPMTB-1203-7
6. Follicllcullus scholasticus OR~IISTON and BABCOCK, 1979 A TPMTB-ll03-3
7. Follicuclllllls veniricoslls OR~IISTON and BABCOCK, 1979 ATPMTB-1202-2
8. Follicllculills d. ventricoslls OR~IISTON and BABCOCK, 1979 ATPMTB-1201-3
9-10. Albaillellidae gen. et sp. indet. ATPMTB-llOl-1,2
11. Anglllobracchia (?) sp. A TPMTB-1201-4
12. Paronella (?) sp. A TPMTB-1201-5

TAKEMURA and NAKASEKO:

A New Permian Radiolarian Genus

Plate 34

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 124, pp. 215-234, pIs. 35-38, December 30. 1981

739.

EARLY SHELL MORPHOLOGY IN SOME UPPERCRET ACEOUS HETEROMORPH AMMONITES*
KAZUSHIGE TANABE

Department of Earth Sciences, Ehime University, Matsuyama 790

IKUWO OBATA
Department of Paleontology, National Science Museum, Tokyo 160

andMASAO FUT AKAMI
Fuchu-nishi High School, Fuchu City, Tokyo 183

Abstract. This paper describes the early shell morphology in some Upper Cretaceous
heteromorph ammonites (five nostoceratids and three diplomoceratids) on the basis
of well-preserved specimens from Hokkaido. The ontogeny of the species examined
may be divided essentially into the planispiral ammonitella first stage and the uncoiled
or abnormally-coiled post-ammonitella second stage, both of which are bounded by anepionic constriction. The surface ornament first appears at some phase of the second
stage. These lines of evidence support the interpretation that an ammonitella wasformed within an egg capsule.
The three diplomoceratids are clearly distinguished from the five nostoceratids inthe presence of well-marked, prorsiradiate constrictions in the earliest post-ammonitellastage. The mode of coiling of the examined species is roughly classified into thethree types (planispiral, helical and repeating U-turn coiling types) by means of RAuP's(1966) parameters. We postulate that all of the examined species might have passed
a planktonic life for a limited period of the earliest post-ammonitella (= post-embryonal)
stage. However, a benthonic life can be inferred as to their middle to later growthstages, because of the loss of bilateral symmetry of the shells and other various
indirect evidence.

ammonites (TANABE et at., 1979, 1980a).
In those papers we stressed the significance of early internal shell structures tothe researches of major taxonomy andphylogeny of the Mesozoic Ammonoidea.
Most of the species examined in ourprevious papers belong to the Phylloceratina, Ammonitina and normally-coiled

Introduction
We formerly summarised the internal
shell structural characteristics in the early
growth-stages of some Upper Cretaceous* Received June 23, 1981; Contribution to
the early ontogeny of modern and fossilchambered Cephalopoda, III.

215

216

Kazushige TANABE et al.

Lytoceratina, besides selected scaphitids.
Although early ontogeny in the Cretaceous heteromorphs has been studied onsome genera (e. g. Ptychoceras, Hypoturrilites, Acrioceras, Ammonitoceras, Baculites, Scaphites and Otoscaphites) (BRANCO,1879-80; SMITH, 1901; BIRKELUND, 1967,
1979, 1980; BIRKELUND and HANSEN, 1968 ;ATABEKYAN and MIKHAYLOVA, 1976;
DRUSHCHITS et al., 1977; TANABE et al.,1979), details of the Nostoceratidae
and the Diplomoceratidae have not yet
been realized to our satisfaction. Speciesof the above two families are found abundantly in the particular sedimentary faciesof the Upper Cretaceous in Hokkaido, andthey have been already studied taxonomically by several authors, e. g., YOKOYAMA(1890), JIMBO (1894), YABE (1904), WRIGHT
and MATSUMOTO (1954), MATSUMOTO(1963, 1967, 1977), MATSUMOTO and MuRAMOTO (1967) and MATSUMOTO andKANIE (1967). Nevertheless, the characteristics of the early growth-stages ofthe species described previously have long
been uncertain, owing much to difficulty
in separating delicate young whorls fromhost rock.
Recently we have successfully observed
the external shell morphology in the earlystages of five nostoceratid and three
diplomoceratid species. This paper summarises the results of the observation andconsiders their implications for majortaxonomy and mode of life.
The species utilized were collected from
Turonian to Santonian rocks of Hokkaido.
Their locations, horizons etc. are described
elsewhere in the text. The repositoriesof the examined specimens are as follows:
Department of Geology, Kyushu University
for specimens with prefix, GK.; Department of Paleontology, National Science
Museum for specimens with prefix, NSM. ;Department of Earth Sciences, Ehime
University for specimens with prefix, EE.

Observations and tracing of external
shell morphology and sutures were madeusing a binocular microscope with a tracing mirror (Nikon Co., Model SMZ-10),
with magnifications of six to twenty
times.

Observation
SuperfamilY Turrilitaceae GILL, 1871
Family Nostoceratidae HYATT, 1894
Genus Eubostrychoceras
MATSUMOTO, 1967
Eubostrychoceras japonicum (Y ABE)
PI. 35, Figs. la-e; Text·fig. 1

Material :-One specimen, NSM. PM6593
from the Nakakinembets River, Obira area,
northwestern Hokkaido. Details for its
location and horizon are unknown. T.
MAEHARA coli.
Description :-The shell is made of aplanispiral ammonitella (=nepionic shell)and subsequent helically coiled free whorls
of more than four volutions. The ammonitella, about 2 mm in diameter, probablys consisting of a protoconch and afirst whorl, is involute with round venter.
It is followed by two widely open, slightly
helical whorls with large diameters in thenext growth stage. At the end of thesecond helical whorl the coiling axis shifts
abruptly at 180 degrees, and thereafter, thecoiling begins to change from dextral tosinistral. The sinistral stage is represented by two or more whorls with anarrow umbilical perforation, which arehelically coiled around the coiling axis.
The whorls of this stage are more elongated longitudinally than those of theearlier stage. The above ontogenetic
changes in mode of coiling can be easily
expressed by RAUP'S (1966) parameters

739.

Upper Cretaceous Heteromorph Ammonites

for shell geometry; i. e. T (whorl translation) tends to increase from the dextral
to the sinistral stages, in contrast to thegradual decrease in D (distance of generating curve from coiling axis). W (whorl
expansion rate) increases with growth ina constant rate.
The whorl of the ammonitella is smooth
without any trace of ornamentation, but
in the subsequent growth stage numerous
simple ribs with regular space begin toappear. The interspace is wider on theventral side than on the dorsal, and therib density increases with growth. Flaredribs are not present in the earlier stageof less than 3 mm in whorl height, but
afterwards they are common or frequent.
Constrictions without tubercle irregularly
occur along the flared ribs. Whorl section
is subcircular throughout the post-ammonitella stage.
The siphuncle is marginal at least in
the post-ammonitella stage, and is runningat or somewhat above the mid-flank.
Sutures in the early stages are characterized by bifid L, as in other nostoceratids.
Remarks:- The diagnosis of the present
species, except for the earliest stage, hasalready described in detail by MA TsmlOTO(1977). Our observation essentially coincides with that of MATSUMOTO. According to him, the present species shows afairly wide variation in the mode of coiling (especially in whorl translation) in theearly stage, and the dextral coiling ismore frequent than the sinistral in thelater stage.

Eubostrychoceras d. E. sa:wnicum(SCHLUTER)

Pl. 36, Figs. la-d; Text·flg. 2

Material :-One specimen, NSM. PM7127
from loco SN2001, upper stream of the217

Eubos trychoceras japonicum (Yabe)
NSM. PM 6593

/

/

B.

lmm

Text·fig. 1. Eubostrychoceras japonicum(Y ABE). Diagrammatic sketch of NSM. PM
6593, showing the mode of coiling andsutures in the early growth·stages. A.
lateral view, B. upper view.

Sannosawa, the tributary of the Horomui
River, Manji area, central Hokkaido (samespecimen as in fig. 1 on pI. 2 of OBATA
and FUTAKAMI, 1975). Upper Turonian.
M. FUT AKAMI colI.
Description:- The shell is rather small,consisting of two subparallel shafts in theearliest stage and succeeding at least threehelically coiled whorls in the later one.
A planispiral ammonitella is not preservedin the specimen examined, but it is probably small in size, judging from thedimension of the initial shaft (about 0.4
mm in the cross section), which might

218

Kazushige TANABE et al.

be connected with the ammonitella. The
initial shaft slowly increases its diameterwith growth, and by an abrupt U-curve
it is connected with a much longer second
shaft. The second shaft is never in contact with the first. It grows upward, andthen it passes to an abrupt bent, forming
an inverse J form. Subsequently the Jformed whorl changes to the succeeding
helically coiled whorls. The mode ofcoiling of the first helical whorl is partly
irregular with a slight deviation of thecoiling axis. The main helical whorls,
whose coiling axis is in parallel with thepreceding second shaft, are contiguous,
somewhat embracing each other. The
apical angle, about 90 degrees, does notchange with growth. The umbilicus israther wide, about 30% of basal diameterat the end of the third helical whorl.
Whorl section is more or less circularthroughout the whole stages preserved.
The initial shaft is smooth, but numerous fine ribs can be observed on thesecond shaft. The main helical whorls
are ornamented with numerous, oblique,gently flexuous ribs. The periodic constriction is well-marked (the earlier two
are especially conspicuous), and bordered
by a somewhat more raised rib. Its frequency is three or four per a helicalwhorl. From the end of the first helicalwhorl the ribs begin to change in the
direction of their obliquity at a constriction.
The siphuncle is located at the ventral
side at least in all stages preserved. It
is nearly at or somewhat above the midflank. Details of sutural ontogeny are
not observed in our specimen.
Discussion :-OBATA and FUTAKAMI(1975) compared the present species with
Heteroceras woodsi KITCHIN (1922, p. 49)
from the Upper Turonian Chalk Rock ofEngland in the change in the obliquity ofribs, the shape of helical whorls and the

Eubostrychoceras ct. saxonicum (SchlUter)
NSM.PM 7127

siphunde

'~

1

.

IUL E

1.A,...j-~Text·fig. 2. Eubostrychoceras d. saxonicum (SCHLUTER). Diagrammatic sketch ofNSM. PM 7127. Lateral (A) and upper (8)views, mode of coiling (C) and sutures (D)in the early growth-stages are illustrated.
Read PI. 36, Fig. ld for PI. 2, fig. ld.

absence of bifurcate ribs on the umbilicalmargin. H. woodsi is, however, a juniorsynonym of Turrilites saxonicus SCHLUTER(1875, p. 30), as WRIGHT (1979, p. 296)
has already pointed out.
WRIGHT(1979) included SCHLUTER'S saxonicus inDidymoceras, HYATT, 1894. Didymoceras
is a better defined helically coiled, "Bostrychoceras" like heteromorph with tworows of tubercles (MA TSUMOTO, 1977).
The simple ribs without tubercles andperiodic constrictions in WRIGHT'S (1979)
saxonicum seem to match well with diagnostic features of Eubostrychoceras MATSUMOTO, 1967. We, therefore, treat thepresent species under E. d. E. saxonicum(SCHLUTER), because the characters in theadult stage are not seen in our specimen.
It is similar to E. muramotoi MATSUMOTO(1967, p. 335, pI. 19, figs. 1-2) from theConiacian of Japan in the mode of coilingand surface ornament, but is easily distinguished in the larger height-breadthratio of the entire helical whorls, broader

739.

U1Jpe1' Cretaceous Heteromo1'ph Ammonites

umbilicus and the absence of bifurcate
ribs.
Genus Madagascarites COLLIGNON, 1966

Madagascarites ryu MATSUMOTO
et MURAMOTO
PI. 36, Figs. 2-3; Text·figs. 3-4

Material :-Two specimens, GK. H8086
from loco Ik 946, Katsurazawa, Ikushumbets area, central Hokkaido (Upper Turonian, T. MATSUMOTO coll.), and EE1013
from loco R4020F, the middle reach of the
Obirashibe River, about 400 m north fromthe junction with the Nakakinembets
River, Obira area (Upper Turonian, 1.
OBATA and others coll.). For the detailed
locations of both specimens see location
maps of MATSUMOTO (1965) and TANABE
et at. (1977).
Description:- The early-stages, except
for the ammonitella, are favourably preserved in the two specimens examined,
especially in GK. H8086. The ammon itella of this species is probably connected
directly with a long shaft, the preserved
earliest stage, as illustrated diagrammatically in Fig. 3. The shaft grows upward
to become slightly sigmoidal, or gently
curved, like an inverse J-form. In the
succeeding stage the initial shaft isencircled by two loose, helical whorls
through an abrupt U-form, which isvertical to the shaft. After this stage
the whorl is descending down with aloosely spiral twisting, and then repeatsa narrow U-turn at four times on a planenearly vertical to the coiling axis (seePl. 36, Fig. 3; Text-fig. 4). The coilingaxis in this stage, therefore, transforms
regularly at about 90 degrees, as if thepreceding whorls are surrounded by Uturns. The fourth U-shaped whorl isconnected with another more opened,

219

long U-turned whorl. The body-whorl
in the adult stage is not preserved in our
specimens. The first shaft is circular in
the cross section, but the succeeding
helical and U-shaped whorls are polygonalin the costal section and subcircular in
the intercostal section.
The initial shaft is ornamented withnumerous fine, annular ribs only. Quadrituberculated, flared major ribs anddense, fine minor ribs first appear in theearliest part of the first helical whorl,
and thereafter they occur regularly andfairly frequently. The latter are present
on the interspaces of and/or on theformer. The tubercles are sharp, spinose,
especially two on the ventrolateral portions.
The position of siphuncle is marginal in
all stages observed. It is running nearlyat or somewhat above the mid-flank.
Sutures are expressed by the formula,
E, L, U and 1. L and external saddles
are bipartite after the helically coiled
stage, and they tend to be deeply incisedin the late growth-stages.
Remarks:- The peculiar, but regularontogenetic changes in the mode ofcoiling in the present species have alreadydescribed by MATSUMOTO and MURAMOTO(1967). They interpreted that the peculiar
mode of shell growth is not of pathologic
nature, but a diagnosis of this species.
Our observation supports their interpretation. As MATSUMOTO and MURAMOTO(1967) have already pointed out, the modeof shell growth in the present species isfairly similar to that in Nipponites
17lirabilis YABE, 1904. The early stage
also resembles those of Muramotoceras(MATSUMOTO, 1977) and Eubostr)'c1lOceras
ct. E. saxonicum in the~ontogenetic change
from a straight form to a helical form.

Kazushige TANABE et al.

220

Madagascarites ryu Matsumotoet Muramoto
GK. H 8086

o

5mm
E

L

U

lJ\/V\ lmm UJL!L.
2

E

LUI
1

Text·fig. 3. Madagascarites ryu MATSUMOTO et MURA'dOTO. Diagrammatic sketchof GK. H 8086, showing the mode of coiling and sutures in the preserved earliest growthstage. s. siphuncle.

Genus Muramotoceras
MATSUMOTO, 1977

J\;fural7lotoceras yezoense MATSUMOTO
PI. 37, Figs. la-b

Jvlaterial :-One specimen, NSM. PM
7215 from a rollen nodule in the Sato-nosawa, the tributary of the Kamikinembets
River, Obira area. Middle ~ Turonian. T.
MURAMOTO colI.
Remarks :-The specimen examined is
one of the paratypes of the present
species, and has already described with
illustrations by MATSUMOTO (1977, pI. 54,
figs. la-b). As MATSUMOTO (1977) mentioned, during ontogeny the mode ofspiral growth of this species changes
from a s~raight or a slightly sin'.lo'Js form

to a free, helical coiling via an abrupt Jshaped whorl growth with a loosely
spiral twisting. In this respect thisspecies somewhat resembles Madagascarites ryu and Eubostrychoceras cf. E.
saxonicum, but differs in that the coilingaxis in the helical stage is not in parallelwith the initial straight arm, unlike thelatter two species.

Genus Neocrioceras SPATH, 1921

Neocrioceras spinigerum (jIMBO)
PI. 37, Fig~. 2a-b; Text-fig. 5

Material :-NSM. PM 9491 from theUpper Cretaceous of Hokkaido or south
Sakhalien. Details of its locality, horizonand collector are unknown.

739.

Upper Cretaceous Heterom01'ph Ammonites

221

x

c

B

D

10mm

E

F

X~'~';J'.
.

~

\

o

E

G

~

x

~B

\~'

-

10mm

Text·fig. 4. Madagascarites ryu MATSCMOTO et MCRAl\JOTO. Diagrammatic sketchof EE 1013. Six different views of the entire shell, showing the peculiar U-turn coilingin the early to middle growth-stages.

Description:- The specimen is of moderate size, about 44 mm in diameter. The
observable earliest portion is a part of awhorl with about 0.5 mm in height. The
imperfect whorl probably corresponds toa first planispiral whorl of an ammonitella,because of the presence of a conspicuous
nepionic constriction at its base (see PI.
37, Fig. 2b). The whorls following thenepionic constriction initially show a very
shallow, open helical, sinistral coiling,
but thereafter a loose coiling in a plane.
The initial whorl is circular in the crosssection, but later depressed to subcircularin the intercostal section or polygonal in
the costal section.
The initial whorl of an ammonitella issmooth. Prorsiradiate major ribs, which
are present in the post-ammonitella stage,

are dense, with regular intervals, associating lateral and ventrolateral tubercles.
Both tubercles occur periodically, but theformer ones are more frequent andweaker than the latter. Fine riblets areon the inters paces and/or on the majorribs, and they are looped even in theearly stage.
The siphuncular position is marginal(ventral) in all the stages observed.
Sutures are expressed as E, L, D and 1.
Lobes Land D, and other saddles arebifid throughout the post-ammonitellastage.
Discussion:- WRIGHT (1979, p. 293)
attributed Neocrioceras to the Anisoceratidae, but he did not mention the reason.
Neocrioceras, indeed, resembles Anisocerasof the Anisoceratidae in the almostKazushige TANABE et al.

~22

Neocrioceras spinigerum (J imbo)
A

NSM. PM 9491

c
10mm

B
2

I

U

L

E

flS1S1tlmm

Text-fig. 5. Neocrioceras spinigerum
Diagrammatic sketch of NSM.
PM 9491, showing the lateral (A) and ventral(8) views, and sutures (C) in the early
growth stages. nco nepionic constriction.
(jll-.!BO).

planispiral growth pattern of whorls withtubercles.
The surface ornament of
Neocrioceras is also similar to that ofHyphantoceras of the Nostoceratidae. In
connection with this problem, N. (?)
undulosum MATSUMOTO, described by
MATSUMOTO (1977, p. 343, pI. 47, fig. 1)
from the Turonian of the Obira area, isinteresting in that the mode of coilingand surface ornament suggest a transition
from Hyphantoceras to "typical" Neocrioceras. In this paper we follow MA TSUMOTO (1977), who referred Neocriocerasto the Hyphantoceras origin.
Family Diplomoceratidae SPATH, 1926
Genus Scala rites WRIGHT et
MA TSUMOTO, 1954

Scala rites scalaris (Y ABE)

PI. 38, Figs. 1a-b; Text-fig. 6

Material :-One specimen, EE 1010 fromloco R 6733a, the northern branch of the
Sannosawa, the tributary of the Shimokinembets River, Obira area (for its detailed
location see TANABE et at., 1977, fig. 8).
Middle Turonian. K. TANABE colI.
Description :-The specimen examinedis an imperfect, immature shell of about
38 mm in maximum size. The preserved
earliest stage is a straight arm of about
6 mm in length. It was probably connected directly with an ammonitella,judging from the dimension of the smallest
part (ca. 1 mm in diameter of the crosssection). The straight arm is followed
by later two whorls of broadly ellipticalopen coiling. The elliptical whorls are
not strictly in a plane, but form a very
low, variable helical coils with different
spiral axes. The whorls are circular or
subcircular in the cross section.
The initial arm is smooth with at least
two well-marked, prorsiradiate constrictions.
The succeeding whorls areornamented with simple annular ribs withregular intervals. Untuberculated flaredribs occur periodically in elliptical whorls
except for the earliest part.
The siphuncle occupies a marginal
position throughout all the stages observed.
The sutural formula is ELUI. All of theelements other than I are bifid. They
are deeply incised and narrowed at theirstems on the first helical whorl.
Remarks :-According to WRIGHT and
MATSUMOTO (1954), the present species
shows a considerable variation in themode of coiling in the early stage; but
in the later stage the loose, elliptical
whorls grow roughly in one plane.
The early stage of this species is fairly
similar to those of other diplomoceratids
such as Polyptychoceras, subptychoceras
and Ryugasella in the presence of an

7:]9.

Upper C1'etaceous Heteromorph Ammonites

22:3

Sea/arites sea/aris (Yabe)
EE 1 01 0

U

M

L

r- E
A

WW l
I

U

L

E

JWWl.fbase ofbody chamber
5mm

Text·fig. 6. Scalarites sealaris (YAIlE). Diagrammatic sketch of EE 1010, showing
the lateral view and sutures in the early growth·stages.

initial straight stage with well marked,
prorsiradiate constrictions (d. figs. 1-8
of WRIGHT and MATSUMOTO, 1954).

Scala rites mi/zoensis WRIGHTet MATSUMOTO
PI. 38, Fig. 3; Text·fig. 7

Material :-One specimen, EE 1011 froma rollen nodule, probably derived fromloc. R 2020, middle reaches of the
Obirashibe River (about 400 m north fromthe junction with the Nakakinembets
River). Upper Turonian. l. OBATA andothers coIl.
Description:- The shell is small, abol:t
23 mm in the preserved maximum dimension. A planispiral ammonitella is missingin the specimen examined, but it wasprobably followed by a straight arm ofat least 10 mm in length, which is theactually preserved earliest part. At thenext stage the straight arm is curved toform a U, and thereafter changes toelongated elliptical whorls in nearly one
plane. The whorl section is circular or
subcircular in all the stages observed.
The initial shaft is smooth, but withwell-marked,
somewhatprorsiradiate
The elongatedperiodic constrictions.
elliptical whorls are ornamented withnumerous narrow, annular ribs, which
are separated by broader inters paces.
The ribs are never flared at least in theearly stage. There are three weak constrictions on the elliptical whorls of thepreserved 1.5 volutions.
The siphuncle is marginal (ventral) in
all the stages observed. The suture is of
E LUI type. Except for the earliest
stage, E, Land U are bifid.
Re7l1arllS :-According to WRIGHT and
MATSUMOTO (1954, p. 118-119) this species
suggests a progression from the typical
Scala rites scalaris to Polyptychoceras inKazushige TANABE et al.

224

3.

Scalarites mihoensis
Wright et MatsumotoA

E E 1011...--- base ofbody chamber

E

LUI

2.~
..l!nm.

•/,"'- ...... "

I

\

....

__ ...

E LUI

\

I

1.

\

/

"

t0M
lmm

8
B

Text-fig. 7. Scalarites mihoensis WRIGIIT et MATSliMOTO. Diagrammatic sketch of EE
1011, showing the lateral view, cross-section of whorls along A-8 line, and sutures in theearly growth-stages.

the more elongated elliptical whorls withnearly straight arms and very infrequent
flared ribs. These characteristic featuresof the present species are already present
in our specimen of the immature stage.

Scala rites (?) aff. S. scalaris (Y ABE)
PI. 38, Figs. 2a-b; Text-fig. 8

Material ;-One specimen, EE 1012 fromloco R 4016F, middle reaches of the
Obirashibe River, near the junction with
the Nakakinembets River, Obira area.
For its detailed location see TANABE etal. (1977, fig. 9). Middle Turonian. 1.
OBATA and others colI.
Description;-The shell is small, about
18 mm in the maximum diameter, although
the body chamber is incomplete. An
ammonitella has already broken away
during fossilization, but it probably

existed and is followed by a long straightarm of the preserved earliest stage. In
the next stage the long arm forms a U
curve, and thereafter changes to later
whorls of loose and broadly ellipticallycoiling. In our specimen the elliptical
whorls are partly broken, but they show
very low, variable open helical coilings(see restored sketch of Text-fig. 8). The
whorls are circular or subcircular insection throughout the preserved stages.
The initial arm is smooth, but possesses
at least five, well-marked constrictions.
Simple, sharp-headed annular ribs first
appear at the beginning of the firstelliptical whorl. They occur regularly
and fairly freql.!ently, but their interspaces
tend to be broaden as the shell grows.
Ribs are never tuberculated. Flared ribs
and constrictions occur periodically on the
septate whorl, but are on the body-chamber.
The position of the siphuncle is marginal

739.

225

U1Jper Cretaceous HeteromO?'ph Ammonites(ventral) in all the stages observed. The
suture is expressed as E L U 1. Except
for the earliest stage, E, Land U arebifid.
Discussion :-Our specimen is probably
immature, because of its small size. It
is, therefore, rather difficult to determine
its taxonomic position exactly. However,the distinctly constricted straight arm in
the initial stage and the succeeding open,
elliptical whorls nearly in one plane seemto the diagnostic characters of the
Diplomoceratidae. The mode of coilingand surface ornament of this species
actually suggest us an affinity withspecies of Scala rites, especially S. scalaris.
In the typical form of S. scalaris thewhorls are more numerous and accordingly
the periodic constrictions and flared ribs
are more numerous than in this form.
Trianglites MATSUMOTO, 1977, whichis represented by a single species, T.
antiqus fvlATSUMOTO (1977, p. 350, pI. 58,
fig. 1; text-fig. 10), forms a roughly
triangular shape in one plane in the later
growth-stages. Its spirally coiled shellwith simple annular ribs in the earlier
stage are somewhat similar to those of
the present species in the same stage,
but the exact comparison is difficult,because the initial stage of Trianglites
remains uncertain. In this paper weprovisionally treat this species under
Scala rites (?) aff. S. scalaris.
Consideration
Taxonomic implications. - Phylogenetic
relationships of the Nostoceratidae andthe Diplomoceratidae have hitherto been
discussed by some authors (e. g. HYATT.
1894, 1900; SPATH, 1953; WRIGHT and
MATSUMOTO, 1954; WRIGHT, 1957, 1979;
WIEDMANN, 1962, 1969; HOWARTH, 1965;
MATSUMOTO, 1967, 1977; WARD, 1976).
Nevertheless, the schemes by previous

Sea/arites (?) sp. aff. S. sea/an's (Yabe)

U. ~L~:~5
W 7r1
~

W
U

L

EE 1012~ (last suture)

4
E 3

W?nf
I U L

E

!lSli1j 2
I U LEfVV\j 1lmm

5mm

Text-fig. 8.
Scala rites (?) aff. S.
scalaris (Y ABE). Diagrammatic sketch of
EE 1012, showing the lateral view (restored)
and sutures in the early growth-stages.

authors differ considerably from one
another with respect to the ancestordescendant relations at generic level.
The reason for this may be due to thewide inter- or intraspecific variation in
the mode of coiling and the frequentappearance of parallelism and homeomorphism among evidently different
lineages.
We discuss here the taxonomic valueof the morphologic characters observedin the early stages of the examined
species. The results of this work may
be summarized in Text-fig. 9. In all
species the suture is expressed by theformula, E, L, U and I: each element is
simple in the earliest stage, but thereafter E, Land U become to be bifid andflorid, being deeply and narrowly incisedat the stem. Thus, it is rather difficult
to classify the examined species above
the generic level on the basis of sutural
patterns only.

Kazushige TANABE et al.

226

FAMI~GENUS!lORMALLY-CO I LEO
SPECIESS. mihoensis
ScalarJ. tesS.

scaldr.is

5.7 sp. aff.
S.

E.

sClIlaris

jl!tponicu:-:

small ammoni t~l1,l?
(1st stage)
ditto?
(1st stage)oi tto?
(lst stage)large arr~"I1onitellaof ca. 2mm in dia::1.
(1st stage)

Eubos trychoceras£.

~

HadligascarJ.tes

M.

cf. suonJ.cullI
rl/u

ABNORMALLY-CO; LED

STAGE

small ammonitella?
(1st stage)

sm~i!ta:~~!fella?

-circularor subcircularin allstages

ditto
~
ditto
(2nd stage){3:::-d stagelloose, subcircular ~ helical ....·ith
low helical coiling narro ....· umbilc.
(3rd stage)

two parallel shafts~ contiguous(2nd stage)
helical coiling
Drd stage)circularor subcirc'..!larin all.stages(4th st.).. ·!.i . ·· ~ ··. .. · ] e
""">

0

""

.a
0

U

~.
~~

....

0

0'

.... t'"-

_0

::1=

"'~~
""'""
00
0

OM

polygonal~~" '" .a '"
"
-'C
c..
EI-o
0

5g~~g=~a:~---+lO~~~1~~!ica~ ~:i~~~;~ C~~~~l:~)(3rd stage)

g::~

~
~

'"o:

straight arm....,.loose, elliptical(2nd stage) nearly in a plane(3rd stagfJ)
ditto loose, subcircular(2nd stage) low helical coiling(3rd stage)(2nd stage)

"'e-

~

STAGE

c·_
c"

~

~

"'

~

"'

·

~,

. coiling transforms repeatedly at 90· :uCJ:u:,-a':r~h) ~n.
Hurulotocer..

H.

ye<Eoense

unknO'om

straight arm--+- loose helical coiling(2nd? stage)(3rd? stage!')r-----------r---------+-----------~r_------------------------t.c(~:r,~~~ur.=.~~~~) 8
Neocr1ocer.8

N. sp1n1gerum

small amlnonitella(1st stage)loose SUbci(~~~la~t~Og!l)ing in a plane

elliptic.
(2nd St.)

Text-fig. 9. Diagram showing the diagnostic characters in the early growth-stages of theexamined nostoceratids and diplomoceratids.

According to the early ontogenetic
study on some Upper Cretaceous ammonites
by TANABE et al. (1979) in the normallycoiled Lytoceratina and the heteromorph
Scaphitidae, the siphuncle attains to theventral position in the early half of thefirst whorl III the ammonitella stage,
while in the Ammonitina and Phylloceratin a its position gradually shifts fromsubcentral to ventral at some phase(between the end of the second whorl
and the middle of the third one) in the
post-ammonitella stage. In this respect,
the Nostoceratidae and the Diplomoceratidae are more similar to the coiled
Lytoceratina than the Phylloceratina andthe Ammonitina. We, however, do notintend to discuss the major taxonomic
positions of the above two families indetail, because of insufficient data forthis problem.
It seems important that the three
diplomoceratids studied (5. scalaris, S.
mihoensis, s. (?) aff. S. scalaris) havea straight or J-shaped arm with several
well-marked, prorsiradiate constrictions

the earliest post-ammonitella stage, as
in other diplomoceratid genera such as
Polyptyclwceras, Subptychoceras, Ryugasella, Solenoceras, etc. A straight arm or
shaft is also present in the early stageof some nostoceratids (e. g. Madagascarites
ryu and Eubostrychoceras d. saxonicum),but it is smooth without constriction. This
fact indicates that the above three
diplomoceratids are clearly distinguishedfrom the nostoceratids examined. We,
therefore, regard that the constrictedstraight shell in the early stage may be
a diagnostic feature of the Diplomoceratidae, as are the almost planispiralshell growth in the later stage and thepresence of sharp, annular nontuberculateribs.
According to MATSUMOTO'S (1967)
scheme for the phylogenetic relations of
the nostoceratid and diplomoceratid genera(here reproduced in Text-fig. 10, withrevision of MATSUMOTO, 1977), Scala rites,
the probable first-appeared genus of the
Diplomoceratidae, may have been derived
from Eubostrychoceras of the NostoIII

739.

Uppet· Cretaceous Heteromorph Ammonites

ceratidae in the early Turonian. Indeed,
the Middle Turonian S. scalaris is moreor less similar to E. japonicum of
the approximately same age in thepresence of annular, occasionally flaredsimple ribs in rather early stage, bJt thelatter differs from the former in theabsence of prorsiradiate constrictions in
the earliest post-ammonitella stage.
The early stage of the Upper TuronianE. ct. E. saxonicum is quite different fromthat of E. japonicum in the presence oftwo parallel shafts. This fact may indicate
the morphological plasticity of this genus.
Even in a single species, MATSUMOTO(1977) reported a remarkable variation in
the early-stages in a sample of E.
japonicum from the Obira area, northwestern Hokkaido. EubostrycllOceras is along-ranging genus from late Albian toCampanian, and has been regarded as oneof the main stocks of evolution, inparallel with another long-ranging genus,
Hyphantoceras (MATSUMOTO, 1967, 1977).
Taking all of the facts mentioned
above into account, it is postulated thatthe plasticity in Eubost rychoceras may
predict the characters of its offshooted
specialized genera such as Nipponites and/vluramotoceras.
It may be a hard problem for us todetermine whether the remarkable variation in mode of coiling in the Cretaceous
heteromorphs is taxonomically significant
or not. For examples, a peculiar U-turn
coiling is the common feature mlvladagascarites
ryu
andNipponites
mirabilis (MATSUMOTO and MURAMOTO,
1967).
The surface ornament of theformer species, however, q'Jite differsfrom that of the latter in having
ventral and ventrolateral tubercles, andin this respect it is more similar tothe species of Hyphantoceras. Based on
this evidence, MATSUMOTO (1967) interpreted that some tuberculated genera

227

including Madagascarites may have been
derived from Hyphantoceras while nontuberculated genera including Nipponites
from Eubostrychoceras. If this interpretation is correct, the mode of coiling may
be more variable in the Nostoceratidae
than the other characters.
In otherwords, the fairly large variation in modeof coiling in this family may have been
affected more strongly by an ecologicalfactor than a historical one.
Ontogenetic and ecologic implications.It has made clear from our study thatthe examined specimens of Ellbost r)'cl1oceras japonicum and Neocrioceras spin igerum have a planispiral ammonitella in
the earliest growth-stage. As has been
already described, in both species the ammonitella is smooth without any trace ofornament, terminating at the nepionicconstriction. It is followed by the succeeding straight or J-shaped arm and/or
loosely-coiled open whorls in the postJudging from theammonitella stage.
above fact as well as the dimensions of
the preserved earliest stage in all of theother specimens examined, it is presumed
that the shells of the Nostoceratidae andthe Diplomoceratidae are essentially divided into two stages, namely the "normal"ammonitella first stage and the uncoiled
or abnormally-coiled post-ammonitella second stage. In most species studied, ribs
and/or tubercles first appear at somephase of the post-ammonitella stage.
The presence of an ammonitella consisting of a protoconch and a first planispiral whorl has already been confirmed insome Jurassic and Cretaceous heteromorphammonites, i. e. Parapatoceras, Epistrenoceras, Spiroceras (DIETL, 1976, 1978), Acrioceras (BRANCO, 1879-80), Baculites (SMITH,
1901; BIRKELUND, 1979, 1980), Paraspiticeras, Leptoceras (WIEDMA]\l\, 1969), H),poturrilites (ATABEKYAN and MIKHA YLOVA,
1976), Ammonitoceras, Ptychoceras (DRUSH-

Kazushige TANABE et al.

228
ALBIANCENOMANIANTURONIANCONIACIAN

SANTONIAN

Text-fig. 10. Phylogenetic relationships of Cenomanian·Santonian genera belonging to the
Nostoceratidae and the Diplomoceratidae (adapted from l\hTSUMOTO, 1967, 1977 with his permission). Read Pseudhelicoceras for Pseudohelicocera>·.

CHITS et at., 1977), Scaphites (BIRKELUNDand HA~SEN, 1968, 1974; TANABE et at.,1979) and Otoscaphites (TANABE et at.,1979). As these genera belong to various
heteromorph families, it seems certainthat all heteromorph ammonites possessed
an ammonitella, as in other normallycoiled ones.
In conclusion. the presence of an ammonitella in the heteromorph ammonites
known to us seems to support the interpretation that an ammonitella was formed
within an egg capsule (BIRKELUND and
HA!'ISE);, 1968; DRUSHCHITS and KHIAMI,

1969; KULICKI, 1974; TANABE et at.,1980a; BIRKELUND, 1980). In all probability, a very young post-embryonal animalof heteromorph ammonites may havepassed a planktonic life at least for alimited period.
The species described are mostly distributed in the Upper Cretaceous of Northwestern Pacific province, especially Hokkaido and south Sakhalien. We know,
however, the evidence that in the Cretaceous heteromorphs certain species (e. g.
Turritites costatus LAMARCK in the Cenomanian) show a world-wide distribution

Explanation of Plate 35
Fig. l. Eubostrychoceras japonicum (YABE). NSM. PM 6593. Three lateral (a, c-d) and upper(b) views, and lateral view of the earliest growth·stage (e). a, c-d (x 1), b (x l.5), e (x 8).

TANABE et al.:Upper Cretaceous Heteromorph Ammonites

Plate 35

739.

Upper Cretaceous Heteromorph Ammonites

as guide fossils for international correlation (M.HSU:VIOTO, 1973, 1977; KENNEDY
and COBBAI\', 1976). Such differences in
geographic distributions among Cretaceous
heteromorphs are a result of one or a
combination of several factors; i. e. degrees in dispersal of eggs by ocean currents and post-embryonal planktonic or
nektonic life habits. The latter factor
might be correlated with the size of normally coiled stage. More concrete data on
the erliest growth-stage over various
heteromorph taxa are, however, needed
to ascertain the above hypothesis.
With respect to the examined nostoceratids and diplomoceratids in the later
growth-stages, a benthonic mode of life
can be inferred in view of their shellform with abandonment of bilateral symmetry and the characteristic mode of occurrence in the fine-grained bottom sediments, in which drifted plant and woodremains occur frequently (MATSUMOTO,
1977; TA:\"ABE, 1979).
The occasional
accompaniments of jaw apparatus withina rather long body-chamber in situ insome diplomoceratids (TANABE et al.,
198Gb and our unpublished data) may also
support the above interpretation.
As has been already pointed out in theprevious papers (e. g. MATSUMOTO and
OBATA, 1963; MATSUMOTO and OKADA,
1973; TANABE, 1979), in the Upper Cretaceous of Hokkaido the heteromorph ammonites occur commonly or abundantly
in a particular biofacies (so-called Baculites
facies or Scaphites facies), which suggests
an offshore, labile shelf paleoenvironment.
In all probability most nostoceratids anddiplomoceratids examined in the adultstage might have been adapted in various
niches as bottom dwellers.
The mode of coiling in the species examined is roughly classified into the three
types (loose planispiral, helical and repeating U-turn coiling types, (see Text-fig.

229

11», although the details for RAUP'S (1966)
parameters may be quite variable even ina sample of a single species. It seems
interesting that the range of variation in
the mode of coiling of Cretaceous heteromorph ammonites almost coincides withthat of modern gastropods. This suggests us an analogy in mode of life between the above two taxonomically different groups. As has been alreadydescribed, in the examined three helicallycoiled nostoceratids (Madagascarites ryu,
Eubostrychoceras japonicu11l and E. cf. E.
saxonicum), the preserved earliest stage
is encircled by the succeeding helicalwhorls, as if it is protected from destruction. This fact may lead to an interpretation that the siphuncle and camera werestill functional in the middle to adultstages. More concrete data on the modeof occurrence, functional morphology andcomparative anatomy are, however, required to discuss the ecological aspectsof Cretaceous heteromorph ammonites.

Summary
The early growth-stages of some Upper
Cretaceous heteromorph ammonites (fivenostoceratids and three diplomoceratids)
have been studied from the taxonomic,
ontogenetic and ecologic points of viewon the basis of well-preserved specimensfrom Hokkaido. The results of this study
are summarized as follows.
1. The shells of the species examined
essentially consist of the planispiral ammonitella and succeeding abnormally-coiled
whorls.
2. Their suture is expressed by theformula, ELUl. Each element is simple
in the observed earliest stage, but thereafter saddles and lobes, except for I, become
to be bifid and florid. In every species
the siphuncle position is marginal throughout the post-ammonitella stage, as inKazushige TANABE et al.

230
rf>9Vl""'E;..-----------------o Or tincrease of T, decrease ofW, D=const.

f COi{rC?I)Sf.

Z::;:;:~::d'" i G'0" Fi/I),",~Q :"01'
orr...

::I

E~"norma (

type

o

(])

3:
o
_

VI

,,'
.

Anisoceras
C r iocerat ites
Neocriocerasetc.

Dydymoceras
Emperoceras etc.
Hyphantoceras

o

o(])

Sealarites
G ryptoxoceras
Protancyloceras

L..

U
(])

-

etc.

helicoid forms

VI

U

.

-

C

II)

U

..:;. '0
Ryugasella ':0 7

( T)

(])

C

«,9-

TRANSLATION

o

o
(])

~~
.-;. p

(])

L..

(])

.....

Po

o

o(])

U

o

VI

~

L..

"0

(])

"0

o(])

o
II

I-

Scalarites

Baeulites
Sciponoceras

Spiroceratidae(Jurassic)

Polyptychoceras;
Subptychoceras
pseudoxybeloceras
. -

etc.

:,

increase of L decrease of 0, W = const.
Text-fig. 11. Three dimensional block diagram using RACP'S (1966) parameters for shellgeometry, showing the range of variation in the mode of coiling of the Cretaceous heteromorphammonites. Dotted lines in the diagram show the boundary between the whorl-overlapping andopen whorl fields. The range almost coincides with that of gastropods. Read Didymoceras for
Dydymoceras.

Explanation of Plate 36
Fig. 1. Eubostrychoceras d. saxonicum (SCIILUTER). NSM. PM 7127. Upper (a), lower (c)and lateral (b) views, and freehand cross section of the preserved earliest growth-stage (d).
a (x 6), b (x 4), c (x 7), d (x 10).
Fig. 2. Madagascarites ryu MATSUMOTO et MURA1'.IOTO. GK. H 8086. Axial section of theinitial helical coils (a), and upper view of the first helical whorl (b). Both x 7.
Fig. 3. Madagascarites ryu MATSUMOTO et MURAMOTO. EE 1013. Two different views of the
middle growth-stage. a (x 2), b (x 4).

TANABE et al.:Upper Cretaceous Heteromorph Ammonites

Plate 36

739.

UPPe1- C1-etaceous Heteromorph Ammonites

normally-coiled Lytoceratina.
3. The three diplomoceratids, Scala rites
mihoensis, S. scalaris and S. (?) aff. S.
scalaris are distinguished from the fivenostoceratids examined in the presence ofseveral well-marked constrictions on thestraight arm of the earliest post-ammon itell a stage. This characteristic featurein the early stage may be a diagnosis of
the Diplomoceratidae, because it is alsopresent in other genera of the same family.
4. The abrupt change from a normal
to an abnormal coiling at the nepionicconstriction, and the succeeding first appearance ornament and the increasing ofwhorl dimensions after the constriction
are observed in the specimens of E.
japonicUlIl and N. spinigerum. The samefeature has been reported in many generaof Jurassic and Cretaceous heteromorphs,
and furthermore, the presence of an ammonitella with similar internal structures
has been confirmed in all ammonites
known to us. All of these facts support
the previous interpretation that an ammonitella was formed within an egg
capsule (BIRKELUND and HANSEN, 1968;
DRUSHCHITS and KHIAMI, 1969; KULICKI,
1974; TANABE et al., 1980a; BIRKELUND,
1980). It is presumed that a very young,post-embryonal animal of heteromorphammonites might have passed a planktonic life at least for a limited period.
A benthonic mode of life can be inferred
as to their middle to later stages, becauseof the loss of bilateral symmetry of theshells and other indirect evidence.
Acknowledgement.-We are grateful to
Dr. Tatsuro MATSUMOTO, Professor
Emeritus of Kyushu University, for reviewing the first draft and for his valuable suggestions. Thanks are also due toProfessor Dr. Tove BIRKELUND of Copenhagen University for her discussion, andto Messrs.
Tatsuo MURAMOTO andToshiharu MAEHARA for providing us

231

with their interesting specimens. Messrs.
Tomoki KASE (National Science Museum)and Yuichiro MIYATA (Kyushu University)
kindly cooperated us in the field. Thisis a contribution of the cooperative research on evolutionary paleontology of theAmmonoidea, supported by the Science
Research Fund of Japanese Ministry of
Education, Science and Culture (Monbusho)(Project leader, I. OBATA, No. 56340041
for 1981).
References
ATABEKYA:-i, A. A. and MIKIIAYLOVA, I. A.
(1976): Development features of turrilitids
with reference to Hypoturrilites gravesianus d'ORB. Akad. Nauk SSSR Doki.,
vol. 231, no. 5, p. 206-209. (in Russian).
B1RKELUND, T. (1967): Submicroscopic shellstructures in early growth·stages ofMaastrichtian ammonites (Saghaiinites
and Scaphites). Medd. fra Dansk Ceoi.
Forening K¢benhavn, vol. 17, no. 1, p.
95-101, pIs. 1-4.
- - (1979): The last Maastrichtian ammonites. In B1RKELUND, T. and BROMLEY,
R. G. (eds.): Cretaceous-Tertiary Boundary Events. p. 51-57, Univ. of Copenhagen.
- - (1980): Ammonoid shell structure. In
HOUSE, M. R. and SE!\[OR, J. R. (eds.):
The Ammonoidea (Systmatics Assoc. Spec.
vol. No. 18). p. 177-214. Academic Press,Lond.
- - and HANSEN, J. (1968):
Early shell
growth and structures of the septa andthe siphuncular tube in some Maastrichtianammonites.
Medd. fra Dansk Ceoi.
Forening. K~benhavn, vol. 18, no. 1, p.
71-78, pIs. 1-4.
--- and - - (1974): Shell ultrastructure ofMaastrichtian Ammonoidea andsome
Coleoidea and their taxonomic implications.
Det Kong. Dansk Vidensk. Seis. BioI.
Skrift., vol. 20, no. 6, p. 1-34, pIs. 1-16.
BRANCO, W. (1879-1880):
Beitrage zur
Entwickelungsgeschichtederfossilen
Cephalopoden. PaIaeontographica, vol. 26(1879), p. 15-50, pIs. 4-13; vol. 27 (1880),
p. 17-81, pIs. 3-11.

232

Kazushige TANABE et al.

DIETL, G. (1976): Die entrollten Ammoniten
des Schwabischen Juras.
Stuttgarter
Beitr. Naturk., ser. C, no. 4, p. 36-43,
pI. l.
- - (1978): Die heteromorphen Ammoniten
des Dogger. Ibid., ser. C, no. 33, p. 197, pIs. 1-11.
DRUSIICHITS, V. V. and KIIIAMI, N. (1969):
Characteristics of early stages of ontogeny
in some early Cretaceous ammonites.
Moskov Obschch. Ispytateley Prirody
Byutl., Otdel. Ceol., Vol. 44, no. 2, p. 156157 (in Russian).
- - , DOGUZKAYEVA, L. A. and MIKIIAYLOVA,I. A. (1977):
The structure of theammonitella and the direct developmentof ammonites. Paleont. Jour., 1977, no.
2, p. 188-199, pIs. 5-6.
HOWARTH, M. K. (1965): Cretaceous ammonites and nautiloids from Angola. Butl.
Brit. Mus. (N. H.), Ceol., vol. 10, no. 10,
p. 337-412, pIs. 1-13.
HYATT, A. (1894): Phylogeny of an acquired
characteristic. Proc. Amer. Phil. Soc.
Philadelphia, vol. 32, p. 349-647, pIs. 1-14.
- - (1900): Cephalopoda. In ZITTEL, K. A.
von. 1896-1900: Textbook of Palaeontology,translated by EAsnIA:-.i, C. R. p. 502-604.
Lond.
jlMBO, K. (1894): Beitrage zur Kenntniss
der Fauna der Kreideformation von
Hokkaido. Palaont. Abh., Jena, vol. 6,
p. 149-194, pIs. 1-9.
KENNEDY, W.J. and COBBAN, W.A. (1976):
Aspects of ammonite biology, biogeography,
and biostratigraphy.
Spec. Paps. in
Palaeontology, no. 17, 94p. 11pls.
KITCHIN, F. L. (1922): Notes on Heteroceras
woodsi sp. nov., Appendix I. In SIIER-

LOCK, R. A. and NOBLE, A. H.: Ceologyof the Country round Beaconsfield. Mem.
Geol. Surv., U. K., Lond., sheet 255, 49p.
KL:LICKI, C. (1974): Remarks on the embryogeny and postembryonal development ofammonites. Acta Pal. Polon., vol. 19,
no. 2, p. 201-224.
MATSUMOTO, T. (1963): A survey of fossils
from japan illustrated in classical monographs(Primarilyanomenclatorial
revision). Pt. VII & X. Palaeont. Soc.
Japan, 25th Anniversary Vol., p. 27-32,
41-48.
- - (1965): A monograph of the Collignon iceratidae from Hokkaido. Pt. 1. Mem.
Fac. Sci., Kyushu Univ., ser. D, vol. 16,
no. I, p. 1-80, pIs. 1-18.
- - (1967): Evolution of the Nostoceratidae(Cretaceous
heteromorphammonites).
Ibid., vol. 18, no. 2, p. 331-347, pIs. 18-19.
- - (1973): Late Cretaceous Ammonoidea.
In HALLAIVI, A. (ed.): Atlas of PalaeobiogeograPhy.
p.
421-429.
Elsevier,
Amsterdam.
- - (1977): Some Cretaceous heteromorphammonites from the Cretaceous of Hokkaido. Mem. Fac. Sci., Kyushu Univ., ser.
D, vol. 23. no. 3, p. 303-366, pIs. 43-61.
- - and KANIE, Y. (1967): Ainoceras, a newheteromorph ammonoid genus from theUpper Cretaceous of Hokkaido. Ibid., vol.
18, no. 2, p. 349-359, pIs. 20-21.
- - and MURA!vIOTO, T. (1967): Two interesting heteromorph ammonoids from
Hokkaido. Ibid., vol. 18, no. 2, p. 361366, pIs. 22-24.
- - and OBATA, I. (1963): A monograph of
the Baculitidae from japan. Ibid., vol. 13,
no. 1, p. 1-116, pIs. 1-27.

Explanation of Plate 37
Fig. 1. Muramotoceras yezoense MATSU;vlOTO. NSM. PM 7215. Upper (a) and lateral (b) views.
Both x 1.5.
Fig. 2. Neocrioceras spinigerum (JI:-'1I30). NSM. PM 9491. Lateral views of the entire shell (a)and the preserved earliest growth-stage (b). a (x 2), b (x 6).
Fig. 3. Scala rites mihoensis WRIGIJT et MATSC:-'IOTo. EE 1011. Lateral view. x5.

TAN AB E et al. "Upper Cretaceous H eteromm'ph Ammonites

Plate 37

739.

Upper Cretaceous Heteromorph Aml1wnites- - and OKADA, H. (1973): Saku Formationof the Yezo geosyncline. Sci. Rept., Dept.
Ceol., Kyushu Univ., vol. 11, no. 2, p.
275-309 (in Japanese with Engl. abstract).
OBATA, I. and FUTAKAMI, M. (1975): Cretaceous stratigraphy of the Manji area,
Hokkaido. Bull. Natn. Sci. Mus., Tokyo,
ser. C, vol. 1, no. 3, p. 93-110, pIs. 1-2(in Japanese with Engl. abstract).
RAUP, D. M. (1966): Geometric analysis ofshell coiling: general problems.
jour.
Paleont., vol. 40, no. 5, p. 1178-1190.
SCHLUTER, C. (1875):
Uber die Gattung
Turrilites und die Verbreitung ihrer
Arten in der mittleren Kreide D~utschlands.
Sber. niederrhein. Ces., Nat.-u. Heilk.,
Bonn (in Verh. naturh. Ver. preuss.
Rheinl., Bonn), vol. 32, p. 27-31.
S:-!l TJ-! , J. P. (1901): The larval coil of
Baculites. Amer. Naturalist, vol. 35, no.
409, p. 39-49.
SPATH, L. F. (1953): The Upper Cretaceous
cephalopod fauna of Graham Land.
Falkland lsi. Dependencies Surv., Special
Rep., no. 3, p. 1-60, pIs. 1-l3.
TANABE, K. (1979): Palaeoecological analysisof ammonoid assemblages in the Turonian
Scaphites facies of Hokkaido, Japan.
Palaeontology, vol. 22, pt. 3, p. 609-630.
- - , FUKUDA, Y. and OBATA, I. (1980a):
Ontogenetic development and functional
morphology in the early growth·stages ofthree Cretaceous ammonites. Bull. Natn.
Sci. Mus., Tokyo, ser. C, vol. 6, no. I, p.
9-26, pIs. 1-5.
- - , HlRA:-':O, H. and KA)1IE, Y. (1980b):
The jaw apparatus of Scala rites mihoensis,
a late Cretaceous ammonite. Prof. SaburoKAN)10 Memorial Vol., p. 159-165, pI. 20.
- - , - - , ;'-.1AT5C:-'10TO, T. and MIYATA, Y.
(1977) :Stratigraphy of the Upper
Cretaceous deposits in the Obira area,

233

northwestern Hokkaido. Sci. Rep., Dept.
Ceol., Kyushu Univ., vol. 12, no. 3, p.
181-202 (in Japanese with Engl. abstract).
- - - , OBATA, I., FUKUDA, Y. and FLiTAKA:-'l!, M. (1979): Early shell growth insome Upper Cretaceous ammonites and its
implications to major taxonomy. Bull.
Natn. Sci. Mus., Tokyo, sec. C, vol. 5,
no. 4, p. 153-176, pIs. 1-6.
WARD, P. D. (1976):
Upper Cretaceousammonites (Santonian·Campanian) from
Orcas Island, Washington. jour. Paleont.,
vol. 50, no. 3, p. 454-461, pI. 1.
WIEDMANN, J. (1962): Ammoniten aus der
vascogotischen Kreide (Nordspanien). I.
Phylloceratina, Lytoceratina. Palaeontographica, vol. 118A, p. 119-237, pIs. 8-14.
- - (1969): The heteromorphs and ammonoid extinction. Bioi. Rev., vol. 44, p.
563-602, pIs. 1-3.
WRIGHT, C. W. (1957):
Superfamily Turrilitaceae MEEK, 1876. In MOORE, R. C.
(ed.): Treatise on Invertebrate Paleontology, Pt. L, Mollusca 4. p. L214-L228.
Geol. Soc. Amer. & Univ. Kansas Press.
- - (1979): The ammonites of the English
Chalk Rock (Upper Turonian).
Bull.
Brit. Mus. (N. H.), Ceol., vol. 31, no. 4,
p. 281-332, pIs. 1-7.
- - and MAT5U:-'10TO, T. (1954):
Some
doubtful Cretaceous ammonite genera
from Japan and Saghalien. Mem. Fac.
Sci., Kyushu Univ., ser. D, vol. 4, no. 2,
p. 107-134, pIs. 7-8.
YABE, H. (1904):
Cretaceous Cephalopoda
from the Hokkaido. Pt. 2. jOllr. Coll.
Sci., Imp. Univ. Tokyo, vol. 20. no. 2, p.
1-45, pIs. 1-6.
YOKOYAylA, M. (1890): Versteinerung aus
der japanischen Kreide. Palaeontogra.tJhica,
vol. 36, p. 159-202, pIs. 18-25.

Ikushumbets ~.J!!fF~lj,
Kamikinembets l:i\c~~Ij,
Horomui rm",],Manji 75"¥-,:"lakakinembets rl'~c;~JIiJ,Obira 'J':ijZ,
Obirashibe::::O)iR,
Sato-no-sawa (f;:~O)iR,
Shimokinembets rFic{t~1jKatsurazawa tiiR,
SannosawaKaz'ushige TANABE et al.

2:34

I&Wj(]ijE*~~'i;~Y.i:7 :/-'£:71 Hl&0)1JJ:WH't.l!:7rM&~: ~tw~O)J:imILITIE*i!iiO) Nostoceratidae,
Diplomoceratidae nr.jHv::.Jl&-t-7.> ~'~'2:7 :/-'£: 71 J- 5!~ 8 :fll!O)rR:ft:1JJWJO)ffi!:%!i,!l~~~lI& L~0)~~fu~J:0).~~~.L~.~%ft0)5~, a~.~~~ETO)&~~1JJWJ&~~t~

IE L c O),fi t 1llT1Hl;i:; ELUI -e, f&1Hl7H't.l!:{IlI]v::.ff:t£-t-7.>. i t-: Diplomoceratidae HO)
Scalarites3f!f (5~1,fift S.? sp.) flllX:ft:1JJJt1.lV::.0lt, prorsiradiate tJ:<rYn~{'f--:,f-:~
f$ltkO)j't.l!:~1'f-t-7.>,\',\-C:-, Y;iJJWV::.[j'ijt!fdJ: <rY,tLO)tJ:t, Nostoceratidae HO)~2: ftiR5JIj-c:-~ 7.>.
:: O)q~fj;!&(I'.JtJ:1JJ:Wj;'&ft Diplomoceratidae f40) (lliO)~IH::.~) ff:tE-t 7.> (WRIGIIT and MATSU~IOTO,

1954)::~iJ'G,[j'ijHO)~iiO)l'0~7,..tJ:-lt7.>. (tl!.O)%'l'{O)?~-C:U, !I~v::.IjlJti~O)~~niJ;~J?<r:::~ 7,..,I'fj-,fiO)ileJ1*9'e1=A'-c:-t fJH,fH::.~H:-t 7.>. ~O)~~*ffift~JUi&O)~nv::,.lt
rz-c:- ~, RAep (1966) 0) /~ '7 -" - 3' ~ Jflt, 7.> :: 2: f::' J: I) (1) :;JZ~m1 (2) ~@]m1 (3) lifE@]Wi~J~m1O) 3 'll'¥1::'iR;}-C:- ~ 7.>. LiJ' L itcJIlUill2: {l;Lt-: 'ffi ~ ~7G-t-~l'jJ;t5JIjO) *kVEO)~1::' t ff::tE-t-7.>. Ellbostrychoceras japonicum ~ Neocrioceras spinigerllm v::'[j'ijIELt-:.iIeJ{*-C:-ft, ::::<1JJWH::.ijJSffi!:U'.J 1 'ffi0) IjlJtiJ'Ah'! , ~:dJ: 7.> lEM"~O) 7 /' -'£: =- T '7 iJ~~5JI] ~nt-:. 7 /'-'£: =- T '71tlY'l~tJ: <Un (nepionic constriction) ~ljIl::.n'ffi~O)t!f:)l~H::.Mt <. {lliO)f!fv::.:tot'-C t, ill.%'Hi5*iJ' G[j'ijf*O) 7:/-'£: =- T '7 O)ff::tEiJ;jf-I~~,tL7.>. tJ::to~ITii~tnPll~~~j?:zffi1::'A"" -C I±lffl.-t7.>. tJJ:O)*~-\"'(lliO) lE1W{i; • ~~Y~~I::' t ~{J;!O) r"Itmfllfli~#i'0 7 :/-'£: =- T '7 iJ;~:g6b Gn -Ct, 7.> :: ~ iJ' G%" *-7.> 2:, 7 /' -'£: =- T '7 ft9~j:I'-e)t%lt ~n t-:nr1J§~tiJ;~1. '. :to~ G <~'ifi;~~O)
ifllfft[!UI:i.i:O) "IE'ii';mJ." Y;i]{*ft -nifi¥JtJ:i1-lEEij§~'8 7,.., tJI:i.i:Jl£~Em10)1:$1::.j@~r, L -C t, "" t-: 2:;Wf~~.h7.>.
Wlfill-)JJt· 'J-/:iHm1:· =J:jf;[~

Explanation of Plate 38
Fig_ 1.
Fig. 2.

Scala rites scalaris (Y ABE) _ EE 1010. Two lateral yie\\'s. a (x 3), b (x 6).
Scala rites (?) aff. S. scalaris (Y ABE). EE 1012. Two lateral views. a (x 3), b (x 7).

TAN AB E et al. :Upper Cretaceous Heteromorph Ammonites

Plate 38

235

PROCEEDINGS OF THE PALAEONTOLOGICALSOCIETY OF J AP ANO);{;tIt ............................ WJ#fii',(~<i.:.~;j{f;M It t ilmJ1l ......... .

~1fH;j"iitu- t

S *ilIE*r'f:~~ 128Ili1f'lll~ :r; J: 0' IMiitu~, r (I.. . .. .. .. .. .. .. .. ... 'hljldjll~' W:;t1iJ,5'~-jnJi]E*c;Jfwr&;I~O)xtltJ IJQ981:¥,lO}J 3, 4 R 1.:.1E.!ib:J($Jm${jfl-c:'lmm(~nt~o(:t?:f,t;'12714)

rr':Ji]l!*c~l;wJlltII'MO)MltJ................... .
· ... (lit.tf.A: *fti!fI)J • 18 {I::ILz. • f'L4>:~nr,)Vi;~ ..........................

ti:4>:ilin,;

i11't"JtH~ (l'IE;~O)±jHt~Eilt{lO) ~j~ ...... t.t:1J IJEt.jj
ItfJ,L i#l!jJtliU: O)I*ljlg(.:.J: 7.:>Mlt ........... .
· ....................... 'NjHill'E' 1:U/liHfft,\':ij;n~rO)I'lill!*c,JhhJJlltAno)~(lf:c ='--fX~ 7 ..

III!UEZ
l'il1.:o iili:ljli;~cO)IY,Jj!lH':'J: 7.:>x;J-lt ........... .
· ............ - III fl::.iEz • f'i:;j'~f!I; • IJ'~ {jill!:,l'ilt/!r'rH~f]l;V-~O)MjElJ'GO)::1 --':/ I· ....

H "U>-

0) ;Jhit}IJli;Hii;['.1 ~I~*O) ,lv;H;!IW!Wllf ............................ :i'IJI.'~:1,
ftffl#MJllt~O)M~~to)~~KJ:7.:>Mlt....

.... ti:;j,ji'EHI;' ;kIIl1~~::i, • IfI'I' J-t). Ill,Ns *0) ;JfiffifJJi:S il[i*c=t5<:~ (.m.II~f.'~~ i.:.:r;vt 7.:»~(1m¥f1) ....

· ....................... (~.jlJnr$tf· *EBfa-$·~m~n·mmW~·a:4>:~.·~. ~A :4>: iii.! )(;fll:ljl~ 0) 1::° A I· :/::1 7 0) i1, j¥t¥11Hl;E, ..

· ... , ..... , . . . . . . . . . . . . . . . . . . . . . .. ~JJR~=· ................................. tJ.:)t:%fll
On the Maastrichtian palynology- Tl'ipl'ojoctacites 7Et}) .................... r',':i,liffi it'i'
Mega phyto-fossils from the Lower Cretaceous of Kochi Prefecture ............. .

· ................... , ... tiJt:i&:J'11 '~iL!jtechnique of observation on internal
structure of microfossils with a scanningelectron microscope ............ NOMuRA, R.
;\Iotes on the genus Favocassidulina Loeblichand Tappan .................. NOMCRA, R.

{i III M~mlltj«1~fj<i].lllll,8: i':'Uvt 7.:> ;f - ~ :/!h 7(f-*f}ll~J~) rl'O):fiIJ('ft· fl'ft1'f&{t7-'i .... *trFll:;i}• vEjlll5tJ~] • 'i'\'!IU;1vl::::' 'h~R ~. :fiEBitWl,~Jj;f1~,~fJCI7j-I·,:g:I':'lm-t7.:> -.( A /. ~ I:: -/,7:/ 1&ft<;E,0):/t;{JJ ...... jll;j;ft Jl!;. *F''liW~' 1lJF"l9'1ifl

1!1,f.) '*!
Some new information on Late Mesozoic
non-marine faunas in Korea ........ !'.i!; !Ji:'~~
Ages of the Puchanghe and Jingking Formations in Yunnan with discussion on thecorrelation of the non-marine Cretaceousin East and Southeast Asia .. !llliiMf «(I::[bU
Mega-fossil floras in the Acid Rock Group(Upper Cretaceous-Lower Tertiary) ... ,iVf5EO)~ff1i ...... , .................

i\

7~7mMKU~7.:>Sm~~~.!WMc~0)~

It .............................. ·. *H.iiiJJJ
7 ~ n!U;j; '.:.Mt 7.:> 1~WJI'1'lji*c~IJli;flfi!ft!J1t'(, c-t,j'~I~(l1FiiI;5(lib 0) killfr Ut1'f&{t1::illf

A--j- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

71'1'1'1 (~im~IJBI'!~)

Neogene microfossils from Okushiri Island,
Hokkaido ............................... .
· .1"OCE, Y., AKIBA, F. and icHINOSEKI, T.

;jtiit}.;ulili)IIJlllJjO)~j;IIJI";l:jJfIJ'G B€-t7.:> 'Tii[ISTIE*Jjld&;E, ................ IYrJH ffL' 7j~*!jj
F~Mr~~"~W~·~*~·.X~O)::1~!ldt.fijlfUH':'-::>~'-C..................... .

.. }ji1l1tl;,,'J' ~~jrlfr~' ~(:t~~=. PEB'f1:Jtl!.
±(J.:i,'f7.kmO)~lfrl!t~J.JUj!Ht:fi~'f~(-'f¥f1) ....
.................................. ::::;<f>:ft=

fliil,':jM~)lll!IrO) cplfrmfft:i!f~&J'J.~1t:fi i':'-::>~'-C .................. /B4:)Jl(jf.IJBHII • ;\,ah~c
Molluscan fossils from the Atsunai Groupin the Kushiro coal field, eastern Hokkaido................................ HONDA, Y.
i:~j};II~,~*t}jHJjliO) "¥*!lff}~"Cl)=lxJUt:fi ....
.............. IFl{l::iEZ' tiFll~'::f • ~~ i\ltIiir4 fl ;¥t=IIU~*O)::::f1jJ'J. (Acanthotrigonia) 1':'
IYd-tQ~J5·~~ ............ 1TIf\;.iEZ· tiEB~T
Phylogeny of the superfamily Trigonioidacea(Non-marine Cretaceous bivalves) ..... .

236· . . . . . . . . . . .. . . . . . .. .. . .. . ..:l~ 1m 1'1' (f\':wJD

Java ...................... TAKAIIASHI, K.

Pseudogemmula, a new genus of the Tur-

li!J;jUr..)~Jitft'i¥J.)O)i'£dtE

ridae, Gastropoda .............. SHt;TO, T.
l:i'ij;(if:lff1Ttdti'I'O),(:jl!J\l (V!l;lvIH)'.7.t.. if 1 'lJio)
7J·J'J{,,*(I'~t6!ff;t ............ lJllli'liI-JlX: • "f-1f 11Early shell morphology in some Upper Cretaceous heteromorph ammonites ....... .
TANAI3E, K., OUATA, I. and Ft;TAKAMI, M.
i'~~iJ~F.1.m&O),P.Al:~EiP :/-1: 7-1 r ......... .

· ............... ~,+*· ....................... 1"~;j,;~nl;' Ep 1i*If:f![;
A new species of Acesta (Plicacesta) watanabei sp. nov .......................... .
· .. .. .. ... NAKANO, M. and OKA~IOTO, K.
"Megalodon" assemblage from Sambosan Belt
in Kyushu .................... T A:\lURA, M.
Miospores from the Eocene Nanggulan Formation in the Yogyakarta region, Central................... .
1lIIMf~· '/;';1"2:"i I) -1- (f\,:W'C)
j'j~4:~,~O)JO;(tJ[{#Jm:{::''0l 'L ........ ;fi~ff.ftgg!(r;Diagnostic features of "Cinko adiantoides"-type leaves .. HORIUCHI, J. and KIMURA, T.
Three Taxodiaceous conifers from the Neogene formations in Gumman Prefecture ..
· ............. KIMURA, T. and OKUBO, A.
Early Jurassic plants in Japan. Part 6 ....
· .. .. .. .. .. ... KIMURA, T. and TSUJII, M.
A New species of Pl'Otocedroxylon from theUpper Triassic Mine Group, Yamaguchi
Prefecture ...... OISIII, T. and NISHIDA, M.

fl'ifU ~ 7i7:*.~:Jl'¥ilfi:tJ{t{if::''0\''L, :to) 2· ....................... IU)lJ{,fM!~' ~1:r1:¥P]

CONSTITUTION
of the
PALAEONTOLOGICAL SOCIETY OF JAPAN(Jan. 20, 1978 amended)

1. The Society shall be known as the Palaeontological Society of Japan.
2. The object of the Society is to promote the study and popularization of palaeontology and related sciences.
Article 3. The Society, to execute Article 2, shall undertake the following business:
1. Issue the Society journal and other publications.
2. Hold or sponsor scientific lectures and meetings.
3. Popularize the science by field trips, scientific lectures and other projects.
4. Aid and encourage research work; award outstanding contributions to theSociety; carry out the objectives stated in Article 2.
Article 4. To attain the object of the Society, the Society may, by decision of the General
Meeting, establish within it research committees.
Article 5. The Society shall be composed of members who are active or interested in palaeontology or related sciences.
Article 6. The members shall be known as Regular Members, Fellows, Patron and Honorary
Members.
Article 7. Persons desiring membership in the Society are requested to fill out the necessary
application forms and receive the approval of the Council.
Article 8. Fellows are persons who have held Regular Membership in the Society for morethan ten years, have contributed to the science of palaeontology, have been
nominated by five Fellows and approved by the Council.
Article 9. Patrons are organizations supporting Article 2 and recommended by the Council.
Article 10. Honorary Members are persons of distinguished achievement in palaeontology.
They shall be recommended by the Council and approved by the General Meeting.
Article 11. The members of the Society shall be obliged to pay the annual dues stated in
ArticleArticle237

Article 12.

Article 13.
Article 14.
Article 15.

Article 16.

Article 17.

Article 18.

Article 19.

Article 20.
Article 21.

Article 22.
Article 23.
Article 24.
Article 25.

Article 12. Members shall enjoy the priyilege of receiving the Society journal andparticipating in the activities stated under Article 3.
The rates for annual dues shall be decided by the General Meeting. Rates forannual dues are: Regular Members, Yen 4,500; F:=llows, Yen 6,000; and Foreign
Members, Yen 6,000 or its equivalent in U. S. dollars; Patrons are organizations
donating more than a share (Yen to,OOO) annually: Honorary Members are freefrom obligations.
The budget of the Society shall be from membership dues, donations and bestowals.
The Society, by decision of the Council, may expel from membership persons who
have failed to pay the annual dues or those who have disgraced the Society.
The officers of the Society shall be composed of one President, fifteen Councillors
and several Executive members. The term of office is two years and they may be
eligible for re-election without limitation. The President may appoint several
persons who shall be Secretaries and Assistant Secretaries. An Executive Council
shall be nominated and approved by the Council. If necessary, several Executive
members, who must be Fellows, may be added with the Council's approval. Councillors shall be elected from Fellows by vote of returned mail unsigned ballot.
The President shall be a Fellow nominated and approved by the Council. The
President shall represent the Society and supervise the business affairs. The
President may appoint a Vice-President when he is unable to perform his duties.
The Society may have the Honorary President. The Honorary President shall berecommended by the Council and approved by the General Meeting. The Honorary
President may participate in the Council.
The Society shall hold regularly one General Meeting a year. The President shallbe Chairman and preside over the administrative affairs. The program for thr
General Meeting shall be decided by the Council. The President may call <iSpecial Meeting when he deems it nec~ssary. The General Meeting requires thE,attendance of more than one-t2n~:l of the members. The President shall call aSpecial Meeting at the written r.oquost of more than one-third of the members.
The request shall be granted only if the written statement fully explains thereasons for assembly and items for discussion.
Members unable to attend the General Meeting may give as attending member awritten statement signed by himself trusting the bearer with the decision of businessmatters. Only one attending member may represent one absentee.
The decision of the General IIIeeting shall be by majority vote. When the numberof votes is equal, the President shall cast the deciding vote.
The President and Councillors shall compose the Council. The decision of theGeneral Meeting concerning administration shall be considered and implemented
by the Council.
The Executive Council shall carry out the decisions of the Council.
An auditor shall be elected by the Council from Fellows excluding Councillors andSecretaries. The term of office is two years and he may be eligible for re-election.
The fiscal year of the Society shall begin on the first of January each year andend on the thirty first of December of the same year.
The amendments to the Constitution of the Society shall be decided at the General
Meeting and must be approved by more than two-thirds of those members who are
in attendance.

Addendum 1)

Voting in the Council shall be by unsigned ballot.

238
f3 *~:i.$J§t~~~IJ%O)mu~ • •PALAEONTOLOGICAL SOCIETY OF JAPAN, SPECIAL PAPERS, NUMBER 26 ~ 1983 ~I.!tK.
~fiLk<, ~~~.~~.Li~ • • ~~~.~~W~~~~, ~~.~K~h~~$~m~~~L, ~.~'!f L ~¥ffi;*- ~,T 812

mlNilnr:llm:'ffi~;h.'J'I'I*~llI\~$:tt!!~~:¥X~~tL

13 *tt'E¥o~~tHIj%f.W!t~ffil.i< (~1~1~f§.ili{;i(93) ~~;:::.$ L~Iv"'Cl~~l'.

tt'E*H;:::.Wi~J.,iffitr)(-C-, ~)(~t!f7JljllirJJ)U> ~h LH"J~~ Jb ~o I'\I-~*J~ § ~rv;:::.~Mll~r.riU)(~(1)~61:> k Jb ~(f9IJUi:/:/,f-;; 1/ A ~ 1V-:)(~riU)(~) -Coo Jb J:\, '. 7t!J'(~t?t*geri~!fIi ~1j"~-v;:::.KiH'tJ:Vj: f:f(lli~~ J., -: Co ~~ tJ 7i-b , I? UI1'ib:::£/il ~;/'LJ., Ji!.~~:l; J.,~~~;:::.~t, ~~4:1fflV;:::'A:. L- ~J:iic J: I) Jb 7t
m:b;~<~ J: l'.

r)cJ~·)(l¥:.!:: Jb v;: :. -j-7tt6tWm~7'~ljll: Lt.::I~li'~-C-,

r:njjlljf9T~;:::. {/(WJ! L ~lElimt,di!.flB ~J\(f±l -C ~ ~~tL~v;:::'.:5 J., -: .!::. $~;Jf.!::.!:: Jb f;:::.{6i'lij{m~'lJ- L Hltf±l L ~"f ~ l '. (mm~J:~tJg;tntZ L i~).
(3) Fr~nHl\;(Hlrn-C-~b;, {J~~'j~~~IJJlxL, ( J I)cJ~tl:Jif.~~f"?~r~I.'.
(a) '11 JZ,r.-n::14; f9TJm.fiI!:JRat..:lt.m~{tl9[·m,f,Iiflt~-. (4':k~A-C'.:5J.,-:U.
(b) iNcr,-t'r; ilriUx!illll. (frlilR~{-hiG~J., -: Uo(c) liJf'1\:Ij;J~~~\rr. (800~L 200 'f:fJJit, f\IX-C-IlJJo(d) IJ'ItJ:tJ: I? CH;:::,@:Xb;-j-)tt!it;Hin-C-.:5J., -: .!:: ~IDEIj!j. (fj:'mt;-~ 'f-~~~ L -Co Jb J: H.
(e ) ;;j<)(~:m:~ (f,IJJ: I) Ji!.~.m:liH t.::ltlJr:l¥.~-C-IV-:)( 3' -1 7'25{Jilif~J&j1r~iX~-t.::tc L, /q -j} -b';x..
V - I, 'r:{*b,~¥ffi;i1c~J., -:.!::); it":*)('j"'NI5* (8 ,f-,f:!I;z,...) 1;:::,1i'i!E~",,~$5J\.b;.:5~.!:: ~~t,~~~ J: ~~I)clilR (~JH;:::.)(;t~ J., /~ - -c :/ ~) ; tifilXl • ~~"* ~~ .!::hi'I!/:: I) 19[~N~; ~J1i.:IXI}:&:

( 2)

/(

~t5c~.

(f)

ftllt.pG~r:nrpljl'tIp,i::£f±l~tiJ..j6~:fT:f!i,t~~T~~,:iZ:lUiW. (~~Ji!.~~a1E(IJ]UJ:J.,;n~iUt.::~t~~'1J-L~~*- ~r~I.'J.

(g) ~~fth~~=$~.
(4) FfJ JZ,~lfIJRWoi1lttH$!;IJ

1982 if.. 5 Ji 31 EI CIZ,;~f). tf,'JffltWd*~ H~b~&:,'Ji!~;:::.A:. C v 7 '" V- .!:: m1983 ~ 1 Ji ~n;f.~H~-c-'liI'i'JiUJ!:!E~J:, rjlJ6j1?;'v;:::.I"]~~=Y!E-C-~. t.::tc L~~nlIit.::~tl~v:" ~1;Z,'Wi-.!:: 0)1m&15~,*iJj,:~, *.JI~~~ f"i.b· I? )}<61:> J., -: .!:: b:.:5 J., b' Jb Uti ~ Iv.
( 5 ) l:~c (f) ~ ftllb· I? 0) r:n !jlUtf1 '(::£ II\~ til.;~;z,..b: tJ: \, 'J#}f.-lt, 1983 1F-IJr~)( ia51£ifUfiWJliIi:<P. (i1iif:1EJill:*llJ1j-'mmllj)«kJ) ~IPf1j'I\,'t.:L"i-r~-C', J:x (2) ~~f'j:I.J;;z,...t.::~;/'L~\,'J.,~1rf;:::.~;z,...~[@:~h~~J:I)cJ!E L,

(6 )

i~.
ffifO::li.:b;"'UF:tJ:~1rvut, ;~lM'~~ -C-~i'k!El~-C- ~ J., t":ft1r! < r:n.ll'IJv;:::..!:: I) b'b' J., =-f!E-C-~. Xi~5~~lm
l!1JJJlt{G:~Fri\1flt, ~~b'G:fTtJ:l' (f91j~lt 11 Ji* fUrrf1j'i*<fiW), ~~l*1§. ~1ffltJ:Uk!Ef~r:njjlIH;:::..!::t) b·b, I) "i~.

~~~f.'~tX;mti.!:: ~it:J"jln;:::'J: I), ~~~~f:J\(liiIf9U-C-ft 10 Ji 20 13) i -CKfJJttlb;~h'f,t[\tJ:H-h':C fI1l!!.lJ4:~~Hb;*ll: ~;/'LJ., -:.!:: vctJ: "? ~\" i~.

(7)t17]Jj~}~,*.ft~~xi\ll C, Tlfjf9Ij~$~.!:: L~f'FJJltL~r~\". ;;l'Il)j~l~~ti~~~ffil.~v;:::.rj,'I.'~h~~r ~ \, '.

fif'J}tb;b,b, J., ~ -C, t~f7]IJtJ:~1r~lVt ~, 7]IJjjlIJftl'F~~i', *.II'IJ 25 $~~t;·v;:::.~*4~~ L
i~. ~;/'Lt),J:lt~A (f.§..Le;:~vutjliIJim ,!::\"5-:.!::~;:::.tJ:I)"i~. \"<'0b'~iffitr)(~~6I:>~Iffi}K~ J., .!:: ~v;:::.~t, 1i':,f,Ii/\,~~b' G1I'ijf; L ~, W[~J:~;F;WE-~tJ:\" J: 5 v;:::. L ~r~ \". r:n!jltl::~fi'i7f;!:Ij:~IJ;~U\. -C-~J., J: 5, ~f,ii;j~tr:;{::;fll;jflUJ· _/::-Fv;:::.+5J'-~13~c "?~,

O::li.:1i[5%, IH I:&:WJIlX:4:b;1.lJ I? ;/'LtJ:b' -? t.::~f.'f:'lt,

3' -1 f-c'i'f<jtfL~r~\".

/ill:&:~~}jf"" J., -: .!:: b~.:5 J., b· Jb L;/'L i ~ Iv.

------------,---~-~-._- - - - - - - ~/'ji{$i6ilYiW-----------~

1982~ 1 }J23, 24 S

1981~1l}] 23 S-------_ 1982'1'-_6_fj___ I
1982~ 4 }]----~ 129 Jill

\YiJ

i,

;jt

1m 3ti 7(

1982 ~~~ • ~~-C-':I::/ /;f-;/ '7.!,

i}'j:

_

r 7 / 7 A O)Ji!!'l~U:: J!i1J~J t.J':YiE ~n -n'G o (t!t~A:

Jl1jf:l3~W~ • tlLUft:K)~~l'jlj2,1G: T113 JfDjrl$::t~IR5!J;g: 2 - 4 -16

S *$§;~m-l! /?I -S *is"g:4'kJ$~IT.*

OS *is"g:4'kJ$~t!f.5]Jj-~. No. 24 [GO, Hisaharu: Permian Conodont Biostratigraphy of Japan(Dec. 15, 1981) t.J:,::O)vl~f:Ij):ti~niU~ (5Eirffi3,900FL r;5:jSf300F9)o ffili:A$i6I:tfHJj~
O)Itl!O)~tJq] l:< S*iS"g:4'kJ$~!~y.~Jj1!1·~*~ffiil.~fjlJJlX!JJ· :f1i/lI:B~,¥- (millIT.lm:Jlt!R$fii1f.tr-1LHI:k$f!R$i'ffi:!lli~$tf<~) (iZS~1G: -=:'fI:J~tHTmlili]3z:r,5ffjllHJjsl2r::l)~g 12172, i t~ 1:I::jJ.re ffQ~l~1lIT.I
19014) vc:tolm\,' L- i -to !illiZSvc J: 0 tJ:Hg~JJ&:JGI:I::JIt]j(7($~il1i1f9'E~.:jSfJm (i!*~%.ft) :to
J: rJ[!<lftH$f':.lt4'kJJm5tJm (IJJW*~%'fJ) -c-{, J&:flk -? -n, i-to
01lB'f1l53~*K)(fl/l11it.J::k$I*J1*~O)ta1J~m-C~DiIi L- k ":k~fflmx:t'J{*~~Jrn~1t< § t.~5:1U1

1*>" 0) f2i!f<: t.J: , S*$f;J\f~jj\!~t.J'0 r§~sefUl1'* *~PJfii1l~**i1: (B5' 452 '" - Y,
3,400 Fl, 9Bxffl: :fL~tI;A~fl) J t L- -Cfljrr~nt~o ;: 0)9'vCI:l:~1E0):k~t.J:*1i-tG §~5:~f,fi (!iJ4'kJ '1lii4'kJ' A~' is"g:4'kJ' !jffEij]h4'kJ) O)~*}i'?-~t.J:fl/lJlU~Jj· 5t~JJJjvc~~~n, 401O)*~iVG::Z v?v-" /1 9 8 1 ~ 12 }] 25 B
1 9 8 1 ~ 12 }] 30 B
ISSN(?i?iS"g:4'kJ~IUI*72) t.J:-fl/l~;wJ;AIJ-C-mm~n-c\"GoI'IJ.Ijlll% IT~*~ ~~ ~ ~8)( ]j( IR 5!J; g: 2-4-16

B*$~.l9J-l!/!1-P3(~ ff

0031-0204

(1[

;r,s

Q

*

~
Jii: 84780 '1=)
0 3 - 8 1 5 - 1 9 0 3)

B *is"g::1'kJ~~¥!iE'ij;C~'lfr i.i 124-!Y
2, a a a F9

r:p.Ill/j~* J?: m£it! /K 1l!I:~Uii~~l'IJlillli*A~:ft(1-[ g;S

.~

lS

EEl

3i ~t 2 / 13~

0 3 - 9 9 1 - 3 7 5 4)

Transactions and Proceedings of the PalaeontologicalSociety of Japan

December 30, 1981

New Series No. 124

CONTENTS
TRANSACTIONS
737. KIMURA, Tatsuaki and TSUJI!, Masanori: Early Jurassic Plants in Japan.
Part 3 ................................................................ 187
738. T AKEMURA, Atsushi and NAKASEKO, Kojiro: A New Permian Radiolarian
Genus from the Tamba Belt, Southwest Japan.. . . . . . . . . . . . . . . . . . . . . . . .. 208
739. TANABE, Kazushige, OBATA, Ikuwo and FUTAKAMI, Masao: Early Shell
Morphology in some Upper Cretaceous Heteromorph Ammonites . . . . . . .. 215
PROCEEDINGS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235