ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 105

Palaeontological Society of Japan
April 30, 1977

Itaru HAY AMI
Editor
Associate editors Ikuwo OBATA and Tomowo OZAWA

Officers for 1977 -1978
Honorary President: Teiichi KOBAYASHI
President: Ryuzo TORIY AMA
Councillors: Kazuo ASAMA, Kiyotaka CHINZEI, Takashi HAMADA, Tetsuro HAN AI,
Itaru HAY AMI, Syozo HAYASAKA, Hisayoshi IGO, Tadao KAMEl, Kametoshi
KANMERA, Tamio KOTAKA, Tatsuro MATSUMOTO, Tsugio SHUTO, Yokichi
TAKAYANAGI, Toshimasa TANAI, Hiroshi UJIIE
Members of Standing Committee: Kazuo ASAMA (Membership), Kiyotaka CHINZEI(General Affairs), Takashi HAMADA (Foreign Affairs), Tetsuro HANAI, Itaru HAYAMI (Transactions), Hisayoshi IGO (Planning), Kametoshi KANMERA(Special Papers), Yokichi TAKA YANAGI ("Fossils"), Hiroshi UJIIE (Finance),
Juichi YANAGIDA
Secretaries: Ienori FUJIYAMA (Special Papers), Kunihiro ISHIZAKI ("Fossils"),
Ikuwo OBATA and Tomowo OZAWA (Transactions), Kazuhiko UEMURA(Membership), Toshiyuki YAMAGUCHI (General Affairs)
Auditor: Tatsuaki KIMURA

The fossil on the cover:Original figure of Podozamites Reinii GEYLER,

1877, from the Tetori group. GEYLER'S description marked the onset ofmodern palaeontology in Japan.

All communications relating to this Journal should be addressed to the
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University of Tokyo Press, Hongo 7-3-1, Tokyo, Japan.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 105, pp. 1-16, pIs. 1, 2, April 30, 1977

SOME CONSIDERATIONS ON THE ANCESTOR OF
THE FAMILY VERBEEKINIDAE (FUSULINACEA)

672.

FUMIO KOBAYASHI
Institute of Geology and Mineralogy, Faculty of Science,
Tokyo University of Education, Tokyo 112<

Verbeekinidae C1)mJ\:;~t:::.I*I-t7.l;s=fC1)~~: vt*~ C1)i!ijj~£R.1i1f~~t:::..t IJ, Verbe·
ekinidae C1)tJlJ\:;~I:J:. Staffellidae c~;t i?:h -C ~t.:o LiJ' L LEVEN (1970) IU!fm ~'IV"~
7..t IJ Pamirina ~a-~P~L, OzawaineIlidae iJt Verbeekinidae C1)J\:;tJl,,(·~7.lc~;tt.:o*~C1)~~-t~~~j!l:j:1:J:.7i-1lI!:"(· alveolar keriotheca a-1f-t7.l:::' c,(X B~ t~ minerali·zation a-?I-t-C~,t~~,:::.c, "{'7::1,,-~a-tt.:t~~,:::,ct~c:·"(·~7.lo *~C1)m~~ P.
darvasica LEVEN I:J:."{ '7 ::I" - ~ a- ~ 11 t:::'A:ht~~' c mt fJ.a B~ t~ MiselUna ~, ~t:::. M.
dyhrenfurthi (DUTKEVICH) t:::.lifj;}.-t 7.liJt, ~~I:J:.iI 1* ge 1;.C1).t IJ 1JJMC1)J3!:i!M'''(' alveolar
keriotheca iJ t l±ll.l-t7.lo ¥~I:J:.S:JlI~Jlnl.J:tt!!C1)~:tt!!.t IJ *Hl,a-l*. L t.:C1)a-~ -:>iJ'l-tt:::.*Hl,=C1)*~ge~t~ i?

m:::. il1*ge1;.C1)J3!:i!M't:::.:to1-t 7.l ~@)~ff4::ii!rC1)~i!t:::. ,,?~, -C~~ L t.:o kC1)K6*,

m~~.t IJ t ~ i?t:::'r$c~;t i?:h7.l

Pamirina tethydis, P. leveni C1)=tfi~a-~clbt.:o :::.
Misellina ~c~~t~j;gf*iJt~ IJ, IJ'ij-Jl!Ht*:7UJ::t:::.&R--:5
I-t i?:h7.l:::. c a-~clb, OOlitt:::. OzawaineIIidae t:::.~-t7.l:::' C, Verbeekinidae C1)tJlJ\:;~I:J:.
OzawaineIIidae -(.~ 7.l :::. c a-lit~ L t.:o *~Ifli"(·I:J:.:::.:h i? a-,*,/C.'t:::.~~ L, Pamirina C1) 3 ~:hi?I:J:.~@)~ff4:~C1)*~~1;.J::,a-~~ L t.:o ~ i?t:::.vt*¥Ii:i!f~:ht.:~t:::."?~ '-C t ;s=fC1)~Wa-1J~;t t.:o'N;tlt7c

many previous authors treated that thefamily Staffellidae are the direct ascen·dants. In his detailed investigation on
the Verbeekinidae and Neoschwageri.
nidae, however, OZAWA (1970, p. 38, 44)
came to a conclusion that the Verbeeki.
nidae was not derived from the Staffel.
lidae but possibly from the Ozawainel.
lidae. In the same year, LEVEN (1970)
proposed a new genus Pamirina, P. dar.
vasica LEVEN as its type species, and also
described a primitive species of Misellina,
M. dyhrenfurthi (DUTKEVICH) from theArtinskian Safetdadonski Limestone,
southwest Darvas.
Furthermore, hepointed out that the Family Verbeekinidae
is the descendant of the Ozawainellidae.
CHOI (1972) discussed the classificationand phylogeny of the genus Misellina.
Although his conclusion is very inter.

Introduction

From the beginning of this century,
various classifications of the fusulinaceans
have been proposed by many authors.
Currently, fusulinacean foraminifers wereclassified into six families by Ross (1967).
Russian authorities, such as RAUSER.
CHERNOUSOVA (1963a, b) established sevenfamilies and ROZOVSKAY A (1969) recog.
nized eight ones, namely, Ozawainellidae,
Staffellidae, Fusulinidae, Schwagerinidae,
Verbeekinidae, N eoschwagerinidae, Schu.
bertellidae, and Boultonidae. Phylogene.
tic relationships among these families
were also discussed by these authors.
Concerning the family Verbeekinidae,* Received Feb. 23, 1976: read Jan. 26, 1975,
in Tokyo.

1

2

Fumio KOBAY ASH!

esting, it seems to be insufficient to settle
the origin of Misellina, because he treated
only single material from the Kitakami
Massif.
Recently, I have obtained three speciesof Pamirina from the Kwanto Mountainlands, Central Japan. In this paper, I
will describe these species and discuss
their ontogenetic and phylogenetic aspects
considering of several biocharacters andstratigraphic position of Pamirina. Furthermore, for clarifying the evolutionary
trend from the Ozawainellidae to theVerbeekinidae, phylogenetic and ontogenetic relationship between Pamirinaand Misellina is discussed. Three speciesof Misellina collected from the Kwanto
Mountainlands and the Akiyoshi LimestOne of west Japan, are also illustrated
in this paper.
All of the figured specimens are stored
in the Institute of Geology and Mineralogy, Tokyo University of Education.
Acknowledgements: I would like tothank Dr. Hisayoshi IGo, Associate Professor of the University of Tsukuba, for
his continuous guidance and criticalreading of this manuscript. Dr. T. KOIKE,
Associate Professor of Yokohama National University and Mr. K. WATANABE

Text-fig. 1. Index map showing thesample localities.

of Noda Senior Highschool helpfully
advised me. Mr. M. MATSUKAWA ofTokyo Gakugei University helped myfield survey and sampling. A part ofthe expense of this study was supportedby the Scientific Research Fund from
the Ministry of Education.

Geologic Setting of the
Localities of Pamirina
The present material was collectedfrom several localities and their brief
geologic setting are summarized below.
A. Itsukaichi: Upper Paleozoic andMesozoic strata are widely distributed
as several narrow belts trending northwest to southeast in Itsukaichi, southernpart of the Kwanto Mountainlands. Fromthe beginning of 1970, I have been engaged
in the geological survey of this area andbrief summary of geology was presented
by IGO and KOBAYASHI (1974). Pamirina
darvasica LEVEN was collected from Nishihosoo and Fukazawa, 4 km north and 2
km northwest of ltsukaichi, respectively.
This species occurred from the micritic
partly sparitic limestone pebbles of theTriassic intraformational conglomerate.
Pamirina darvasica is associated with
Misellina dyhrenfurthi (DUTKEVICH), Misellina sp. A and others. Accordingly,
its geologic age is approximately equivalent to early Artinskian of the Tethys
region.
B. Hanagiri: Hanagiri, near Kamikutsu, at the southeast foot of Shomaru
Pass (617 m), is situated about 60 km
northwest of Tokyo. According to MoRIKAWA (1955), the Upper Paleozoic formations are typically exposed at Kutsuzawa (valley) and subdivided into thefollowing seven members in descendingorder, 1: grey chert (more than 100 m
thick), 2: Kami-kutsu Limestone Conglo-

672.

Ancestor of the family Verbeekinidae

merate (about 30 m thick), 3: red chert(about 200 m thick), 4: Shimo-kutsu Conglomerate (about 40 m thick), 5: alternation of sandstone and slate (about 100
m thick), 6: Hanagiri Limestone (about
30 m thick), and 7: alternation of sandstone and slate (about 100 m thick).
The fourth Shimo-kutsu Conglomerate
and the sixth Hanagiri Limestone members bear some Lower Permian Schwagerinidae. The second Kami-kutsu Limestone Conglomerate yields some upper Middle Permian Neoschwagerinidae.
Based on my observation, the Hanagiri
Limestone Member attains about 40 min thickness at the type section and isalmost barren in megafossils except forcalcareous algae and small fragments ofcrinoid and shells. The lower part ofthis limestone frequently intercalates
thin red to brown tuff and is highly
crystalline. Fusulinaceans in this partare ill-preserved and deformed. The
middle part of the limestone is grey
micrudite and also frequently intercalates
red to brown tuff bands. It yields many
fusulinaceans, such as Pseudofusulina
fusiformis (SCHELLWIEN and DYHRENFURTH), P. spp., Biwaella sp., Toriyamaiasp., Schubertella sp., Pamirina tethydis
KOBAYASHI, sp. nov., P. leveni KOBA YASHI, sp. nov. and others. The upper
part of the limestone is massive, partly
bedded, mainly light grey to grey mic"
rudite and yields the similar fossils withthe middle part. The occu,rr~nce ofPamirina seems to be restricted to somepart of particular lithology in the limestone. The specimens are crowded in
the algal bio-sparudite, and they abruptly
disappear vertically as well as horizon.
tally. From the mentioned fusulinaceans,the Hanagiri Limestone Member is 'correlated to the upper Sakmarian and also
to the upper Sakamotozawan.'
C. Ohtaki: Carboniferous and Permian3

rocks composed of limestone, green rocks,
chert, sandstone, slate and conglomerate,
are distributed in the Ohtaki area, westof Chichi bu. These rocks are trending
WNW to ESE and bounded by the faults
with the Cretaceous formations at northern margin and with the Jurassic OhtakiGroup at the southern margin. FUJIMOTO(1936), ISHII (1962) and others contributed
to the geology and paleontology of thisarea. I collected some specimens ofPamirina sp." Misellina dr. dyhrenfurthi(DUTKEVICH), M. sp. A, M. spp., Nagatoella sp., Pseudofusulina sp. and Pseudoendothyra sp. from the limestone lenses
cropped out near Uzuradaira, Ohtaki
village. Geologic age of this limestone
is approximately early Artinskian in the
Tethys standard.
Previously described species
referable to the genus Pamirina
Paramillerella, Millerella, Staffella, Nan.
kinella, Pseudoendothyra and others frequently occur, from the Lower Permianrocks associated with many schwagerinids. These fusulinids have been ignoredby many authors because of their rareoccurrence, small size and rather longstratigraphic ranges.
The following
species reported from Japan are thought
to be assigned to the genus Pamirina.
(1) Paramillerella? sp.:- This unidentified species was described by T AKAOKA(1966, PI. 5,. Figs. 1-5) from the Futagoyama, Limestone, K wanto Mountainlands.
T AKAOKA did not describe any detailed
character of shell, but this species isapparently" assigned to' Pamirina. This
species coexists with Pseudofusulina kra/.
fti (SCHELLWIEN and DYHRENFURTH).
(2)Staffella sp. :- This unidentified spe 7
cieswas described by KANUMA (1960, 'Pl'.
12, Figs: 29-31) from the Lower .Permian
Shimadani Formation, Mino, ~untainc

4

Fumio KOBAY ASH!

lands, Central Japan. It has 4 to 5 volu.
tions, 0.34 to 0.40 mm in the length, 0.65
to 0.75 mm in the width, 0.032 mm in theoutside diameter of proloculus, low cho.
mata, thin spirotheca and scarcely reo
crystallized shell. Associated fusulinids
are Pseudofusulina sp. The above.men.
tioned characters of this species are
entirely identical with those of Pamirina.
(3) Pseudostaffella? tamanouchiensis SA.
KAGAMI:- This species was describedby SAKAGAMI (1956, PI. 37, Figs. 1-3)
from the limestone pebbles of the Tama.
nouchi Conglomerate in the vicinity ofItsukaichi, west of Tokyo. According to
SAKAGAMI, the important characters ofshell are as follows. Shell composed of 7
volutions, 0.67 mm in the axial length,1.11 mm in the median width and 1: 0.6
in the form ratio. The spirotheca com·posed of a tectum, diaphanotheca and athin lower tectorium. Associated fusuli.
naceans with this species are Nankinellasp., Staffella moelleri OZAWA, Mesoschu.
bertella thompsoni (SAKAGAMI), Pseudo.
fusulina vulgaris (SCHELL WIEN and DYH.
RENFURTH), P. vulgaris globosa (SCHEL.
L WIEN and DYHRENFURTH) and P. sp..
From the above·mentioned respects, SA.
KAGAMI inclined to propose a new genus
or new subgenus, but he postponed his
conclusion. In my opinion, diaphanothecaof this species is different from that of
the Fusulinidae and may represent rather
thick translucent layer corresponding to"protheca". Thus, Pseudostaffella? tama.
nouchiensis may be assigned to Pamirina.
(4) Millerella hataii SUYARI:- This spe.
cies was described from the Lower Per.
mian Kameiwa Formation, Kochi Prefec.
tUre, Shikoku, by SUY ARI (1962, PI. 1,
Figs. 5, 6).' The spirothecal structure,general shape of shell and number ofvolutions of this species are similar to
Pamir.ina .. ' However, this species shows-partly' evolute one or two whorls andhas rather larger shell than Pamirina.
From the above mentioned biocharactersof this species, I tentatively treat it asa member of Pamirina with a query.
In conclusion, at least the above men·tioned four species in addition to thethree species are referred to Pamirina
in Japan.
Theyare:Pamirina tamanouchiensis (SAKAGAMI),
by SAKAGAMI (1956)
Pamirina sp., by KANUMA (1960)
Pamirina (?) hataii (SUY ARI), by SUY A·
RI (1962)
Pamirina sp., by T AKAOKA (1966)
Pamirina tethydis
KOBAYASHI, sp. nov.
Pamirina leveniby thepresent
KOBA YASHI, sp. nov. author (1977)
Pamirina darvasica
LEVEN
Based on the associated fusulinaceans,stratigraphic occurrences of these speciesof Pamirina in Japan are from the upper
Pseudoschwagerina Zone to Misellina
Zone, or from the upper Sakamotozawan
to the lowest (?) Nabeyaman of Japan.
Pamirina darvasica is known from theArtinskian of Darvas. judging from ourup.to.date knowledge, this genus hascomparatively short stratigraphic ranges,
limited within Lower Permian, and it
represents an excellent stratigraphic
indicator.

1

I

Consideration of the Ancestorof the Verbeekinidae
There are many difficulties to discuss
the evolution of some groups of fusuli.
naceans which have less conspicuous
morphological differentiation and longstratigraphic ranges. Concerning theevolution and phylogeny among thegenera of the OzawaineIlidae, several bio·characters are utilized for the taxonomic

672. Ancestor of the family Verbeekinidae
subdivision, such as, the size. and shapeof shell, number of whorls, structure andthickness of spiro theca, coiling patternof shell, growth curve of shell and others.
Furthermore, the ontogenetical transitionof these biocharacters during the ontogeny and the biostratigraphic data ofthese fusulinids should be also considered.
In this chapter, I deal with the consideration of evolution from the Ozawainellidae to the Verbeekinidae withspecial reference to the spiro thecal structure and growth curve of Pamirina andalso certain species of Misellina.
[Spirothecal structure] As will bementioned in the later chapter, there
are two types of the spirothecal structure
in Pamirina. One type is recognized in
Pamirina tethydis and P. leveni, namely,
the proloculus and the first to third
volutions are composed of structurelesssingle layer. The fourth to last volutions
consist of a tectum and protheca. These
two species are morphologically moreprimitive and occupy a slightly lower
stratigraphic position than P. darvasica.
Pam;r;na {even; sp.nov.

Pam;r;na tethydis sp. nov.

Hanagiri

5

Hanagi ri Spec i mens

Specimens

0.60
0.70

0.50

0.60

OAO

(mm.)

Form ratio of Volution

ratio of
Volution

0.50

2

3

4

5(VoIJ

23

45 (Vol.)23

4

5 (VoL)2

3

45(VoIJ

Imm)
0.200

0.200
0.100
0.100

0

2

3

4

5 (Vol.)
(mm)

(mm.)

~

0.400

OAOO

0.300

0.200

~

0.300

0.200

0.100

0.100

vector2

3

4

5 (Vol.)Text-fig. 2. Variations in relative
growth of shells of Pamirina tethydis KoBAYASHI, sp. nov.

0

Variations in relative
Text-fig. 3.
growth of shells of Pamirina leveni KOBAYASHl, sp. nov.

Fumio KOBAY ASH!

6

tions consist of a tectum and fine alveolar
keriotheca. This species has morphologically more advanced biocharactersand occupies a slightly higher stratigraphic position than P. tethydis and P.
leveni.
The spirotheca of Misellina dyhrenfurthi and M. sp. A are composed of

The other type of spiro theca is recognizedin P. darvasica. The prolocular wall andsucceeding one to two and a half volutions are composed of structureless single
layer. The spiro theca consisting of atectum and protheca is developed in two
and a half to four and a half volutions.
Outer four and a half to the last volu-

Mise/lina dyhrenfurthi(Dutkevich) & Mise/tina sp.A

Itsukaichi&Ohtaki

Specimens

1.20-... ----------

_____

1.10

,I"
........,-,J.'-

,

1.00

"

/---"_.l.' ,---._._",1 - .. __ _ -"",-"'"

.-

(mm.)

0.700

/

0.80

" ~"

/~

0.70
0.600

,/

" "~

0.60

,It' ,,/ ~

0.500

0.400

0.700

0.600

~

,

0.200

0.400

,

I

I

,

,'

,

,, ,,
, "
"

I

t] / "/-------Microspheric

,

,'

I

I

I

,

Misellina sp.A

0.500

,,

I

----------- Mega.!ospheric

0.300

It' ,,I ~,'/ ,,' ~'trof Voiution

(mm.)

,

I

,-

I

I'

,

,,',,, /
"

",1

0.100

0.300

o

0.200

," " ",,"

0.100----Misellina dyhrenfurthi(Dutke.)

o

2

3

4

5

6

7

8

9 (Vol.)Text-fig. 4. Variations in relative growth of shells of Misellina dyhrenfurthi(DUTKEVICH) and Misellina sp. A.

672.

Ancestor of the family Verbeekinidae

7

Misellina claudiae(Deprat)

Akiyoshi Specimens
1.40
1.30

----_.- ..

1.20
1.10
1.00
(mm.

,,
) 0.90 4----\/

ra tio of Volution\\

1.400 0.80

\
\
\

\

1.300

/

/

0.70

\

\
\

\
\

1.200
1.100

0.60

/

0.50

.

I

I

(mm)

- - - Mega/ospheric--------- M i ero spheric

,
I

,I ,'

0.8001.100

II

0.7001.000

I

I

,
,

I

/

II
/

I

,

I

I

/

I
I
I

I

I

I

/

I

I

I

I

I

l

I

,

1.000

I

/

, ,
, I,
,
,

0.900

,

,

I

I

I
/

'

I"

I

'"

/ / t\\."
I/,' ~"

0.6000.900/

0.5000.800, ",/ " \.I?

,'/' / / '\.'~/,// ~"t;,,0.4000.700......

0.3000.600","".'

0.2000.500

o
0.200

o

2

3

4

5

6

7

8

9 (Vol.)Text·fig. 5. Variations in relative growth of megalospheric and microspheric shells of Misellina claudiae (DpPRAT).

8

Fumio KOBAY ASH!

lar keriotheca recognized in the third or
fourth to last volutions in the megalospheric specimens. However. the first
appearances of protheca imd keriothecastructureless single layer in proloculus
and inner onn or two volutions. A tectumand pro theca developed in the second to
fourth volutions. and a tectum and alveo1.40
1.30
1.20

Form ratio of Volution/ /"---- ---'"~(mm.)
1.400 1.00

.....

1.300
1.200
1.100

0.70

1.000

0.60

0.900

0.50

............•.
Misellina claudiae--------- fvficrospheric----fvfegalospheric

O.BOO
0.700

(mm.)
0.600 1.100

Miseltina sp.A
fvficrospheric- - - - - fvfegalo sph eric
0-----00

0.500 1.000
0.4000.900

0.3000.BOO
0.2000.700
0.1000.600

o
0.400
0.300
0.200

- - - Pamirina tethydis---------- Pamirina leveni

0.100

o

2
Text-fig. 6.

3

4

5

6

7

Average relative growth of shell.

·8

9 (Vol.)672. Ancestor of the family Verbeekinidae
started in later ontogenetic stage in the
microspheric specimens than those of
megalospheric form. These two speciesof Misellina are associated with Pamirina
darvasica in Itsukaichi and with NagatoeLLa sp. in Ohtaki.
The spirothecal structure of Misellina
claudiae from Kaerimizu of the Akiyoshi
Limestone is composed of structure lesssingle layer in proloculus and inner one
volution, tectum and protheca in one
and two volutions, and a tectum andthick layer of alveolar keriotheca inthree to the last volutions in the megalospheric specimen. However, as in thesame case of the above Misellina sp. A,
protheca and keriothe:a first appear alsoin later ontogenetic stage in the microspheric specimen. Associated fusulinaceans with M. claudiae are Nagatoellaorientis (OZAWA) and others. ParafusuLina
kaerimizensis (OZAWA) appears at stratigraphically several meters high fromthis level. Therefore, Misellina claudiae
is morphologically more advanced andoccurs from a slightly higher stratigraphic position than Misellina dyhrenfurthiand M. sp. A.
Summarizing the above-mentioned evidences, a keriotheca is not visible inPamirina tethydis and P. Leveni. The
first appearances of protheca and keriotheca are recognized in their earlier
ontogenetic stages in Misellina dyhrenfurthi and M. sp. A than Pamirina darvasica. The development of these layers
is the earliest in Misellina claudiae (Textfig. 8).
As discussed and noticed by many
fusulinacean paleontologists, the changeof spiro thecal structure through the
ontogeny and phylogeny is one of themost important phenomena to discuss
the evolution of this superfamily.
[Growth curve] Relative growth offusulinidshell is graphically expressed9by plotting the values of radius vector,
half length and form ratio at various
ontogenetic stages. All of the measured
values in Pamirina tethydis, P. Leveni,
Misellina dyhrenfurthi, M. sp. A and M.
claudiae are graphically shown in TextFigs. 2 to 5, and mean values of them
are also shown in Text-fig. 6. Pamirina
darvasica from Itsukaichi is omitted
because of its poor data. As evidently
shown in Text-fig. 6, the gradationalchanges of the growth curve of shellfrom Pamirina to Misellina are recognized.
In conclusion, the above considerations
on the spirothecal structure and growth
curve and on the spirothecal differentiation into two types in the PermianOzawainellidae, as later given in thediscussion on the diagnosis of Pamirina,
suggest that Pamirina and Misellina havethe same evolutionary series and that
the Verbeekinidae are the direct descendants of the Ozawainellidae (Text-fig. 7).

1

to Verbeekinidae
Neoschwagerini dae:advanced Misellina.A(M.claudia. etc.):. M. dyhrenfurthi ". M.otakiensis Group ."
'.

.

.

.. ::.,:-~

Pamirina darvaslca

1'.::1

P. tethydis
p. leveni.,.:....:.. ()F:\. .1

Ozawaine/l idae··i.)\'

Text-fig. 7. Phylogenetic position of
Pamirina; Dotted branches belong to the
Ozawainellidae, and crossed one is theVerbeekinidae.

Fumio KOBAY ASH!
P.tethy-P.dw,,-a Mdvhrenfurthh. M. sp.A.
dIS
sIca
. OJ'
IrMioalospheflc)
5.5.

5.5.

5.5.

I.w. n.. 5.5.

I.w. np.

M. sp.A(MicrosDheric)

5.5.

I.W. n.p.

10 .030

M. C'/audiae

M. claudiae

5.5.

5.5.

(MeoalosDheric) (Micros heric)::.'

1---H--1A-.:fr.. :12t .030 22 [I .069 26 \'..+-YLJJ

1--=-*-';"':"

I---tt--x-A ,~1Ji

I----II---().02715I)!

.065 25.020 12',:.04221.04514'; J.. .013 8::::.04618.;.:.025 13

I':itlf·::'i:...... ~ 014 5 IT 025 7 ;':I-----lf-""I~.~.: f-.:f-F!:."
R:':'l:'~ - ..y.J..... - - ~L; :013 5 ~.'~.~.- -I - - - - * - I ....

"':,:.~/

1'(: li.e:] -5.5.:5pirolheca! '5tructure- : Structureless

-

1)1

: .:

t.w.:thickne55 of wall:;:~1-+.-0-54-+-37--1.025 16 ""~ .050 291A .069 31

I I .019 10 ..L-l- .02511 11). -

A

.050?

::::.

.03419 ..!:+-.055 18

I.w. np.

I, ~::.:.

I---/l------+--+---I--+-+-f}.A:.::--+.05- 0-l-?-H+ .030 18 r.;~.
1--_-Il-_--I----r---+__"'<---+------1f.----1-1T1.063?

I.w. n.p.

);.057 26'..

.043 22"TT" .048 18

4 ~ .028 12 1')- .025 12.032 9 I:..'
9

-R.

-

1."...
'..,

r:;,_ -

4 ... .

- -:{\"' - : .

n.p.:number of parachomata~:Tectum+Protheca...........: Tectum + Keriotheca

Text-fig. 8. Diagram showing the transition of each spirothecal structure,
thickness of wall (in mm.) and number of parachomata at each ontogenetic stage.
The latter two biocharacters are shown with the mean values.

Description of Species
Family Ozawainellidae THOMPSON
and FOSTER, 1937
Genus Pamirina LEVEN, 1970
(Type species]: Pamirina darvasica
LEVEN, 1970
(Revised diagnosis]: Small and subspherical to thickly lenticular shell witharched to broadly arched periphery andinvolute volutions throughout growth,more or less depressed along the axisof coiling, a slight rotated coiling injuvenile stage in some specimens. Shell
not secondarily recrystallized, of 4 to 6
whorls, rarely 6.5 to 7 in mature stage.
Axial length ranges from 0.2 to 0.9 mm,
and median width from 0.3 to 1.2 mm,
giving form ratio of 0.4 to 1.0. Proloculus
spherical, small and its outside diameter
varies from 0.04 to 0.07 mm. Shell expands gradually. Spirotheca smooth, thin

(0.01 to 0.05 mm) and has two types intheir structure. One type is composedof structureless single layer in proloculus
and one to two and a half volutions,tectum and pro theca in two and ahalf Yolutions, and tectum and fine alveolar keriotheca in four and a half tothe last volutions, in advanced species,
such as Pamirina darvasica. The otheris composed of structureless single layer
in proloculus and succeeding one to three
volutions and tectum and protheca inthree to the last volutions, in such primitive species as P. tethydis and P. leveni.
Chomata weak but distinctly developed.
Tunnel low and the path rather regular.
Parachomata lacking.
(Discussion] : In typical Ozawainellidae, spirotheca is composed of thin
three layered structure, tectum, lower
and upper tectoria, as represented by
the most genera of the Carboniferous
Ozawainellidae. However, there are atleast two types of the spiro thecal structure in Permian species of the Ozawainel-

672.

Ancestor of the family Verbeekinidae

lidae. One consists of relatively thick,tectum and less dense structureless lower
layer which sometimes accompany finealveolar keriotheca, as clearly shown at
the mature stage of Toriyamaia KANMERA. It is also imagined that Leella
DUNBAR and SKINNER and Rauserella
DUNBAR may have similar spirothecal
structures to Toriyamaia. The other typeof spiro theca is found in Reichelina ERK.
According to my observation of the latest
Permian Reichelina cribroseptata ERK and
R. cfr. tennuissima M. MACLAY from
Itsukaichi (KOBAYASHI, 1975), the spirotheca is composed of three-layered structure as found in the Carboniferous
Ozawainellidae, and diaphanotheca isinvisible.
From the above mentioned, Pamirina
darvasica has the former type of spirothecal structure. However, no alveolar
structure is visible in P. tethydis and P.
leveni. Thus, I consider that the genus
Pamirina occupies the transitional stageof evolutionary lineage from the Ozawainellidae to the Verbeekinidae and alsothat P. tethydis and P. leveni are moreprimitive than P. darvasica in view of
the transition of spirothecal structure
and the stratigraphic position. Similartransition of spiro thecal structure is also
discriminated in the genus Triticites
GIRTY by ROZOVSKA YA (1948, 1949, 1950).
Pamirina tethydis KOBAYASHI,
sp. nov.
PI. 1, Figs. 1-12

Shell small, scarcely recrystallized,
subspherical in shape and with convex
lateral sides. Polar regions flat or withshallow umbilical depressions. Peripherybroadly arched throughout growth. Shellcoiled involute and planispiral throughout.
Mature shell with 4.5 to 5 whorls. Axial

11

length 0.338 to 0.425 mm and medianwidth 0.525 to 0.650 mm. Form ratioof mature specimens varies from 0.57 to
0.76 in nine specimens. Shell expands
gradually. Proloculus minute, sphericaland its outside diameter ranges from
0.038 to 0.060 mm in nine specimens.
Spirotheca thin, composed of structurelesssingle layer in the inner two and one
half to three and a half volutions,tectum and protheca in the succeeding
ones. Septa numerous, plane and un fluted
throughout. Septal counts from the firstto the fourth whorls 7, 10, 12 and 16,
respectively. Chomata distinct, weakly
developed, but indistinct or almost lacking in the outer whorls of some specimens. Tunnel low and its path regular.
Remarks:- This species is closely similar to Pamirina leveni KOBAYASHI in thesize of shell, number of whorls, spirothecal structure and others. However,the former has longer axis of coiling,
more broadly arched periphery andlarger form ratio of volutions throughout
the growth than the latter, and is easily
distinguished from Pamirina darvasica
LEVEN in the spirothecal structure andsize of shell.
Occurrence:- Abundant to common
from two levels in the Hanagiri Limestone, associated with P. leveni, Pseudofusulina fusiformis (SCHELLWIEN and
DYHRENFURTH), Pseudofusulina spp.,
Schubertella sp. and others.
Reg. nos.:- 2676-13 (Holotype), 2676-1
A, lB, 2A, 2B, 11 and 13, 2775-2A, 6A
and 7 (Paratypes).
Pamirina leveni KOBAYASHI, sp. nov.
PI. 1, Figs. 13-38

Shell small, scarcely recrystallized,
thickly lenticular in shape and convex
lateral sides. Polar regions flat or with

Fumio KOBAYASHI

12

Measurements of Pamirina tethydis
Half lengthReg. no.
of
specimen

L.

w-.

R.

P.

2775-2A
2676-1l.
2676-:-13:
2775-;7 :'.
2676;-2A
2775-6A
2676-1A
2676-2B
2676-1B.425.338.. 400.
.345
.235
.325
.263
.250
.238

.650
.588
.525
.575
.380
.450
.513
.400
.400

.65
.57
.76
.60
.62
.72
.51
.63
.59

· 045(?)
.060
.050
.050
.038
?
.038
.055
.050

slight umbilical depressions. Peripheryarched throughout. Shell coiled involute
throughout growth and slightly rotatedaxis of coiling observed in inner one or
two whorls. Mature shells with 4.5 to
5 volutions. Axial length ranges from
0.225 to 0.350 mm, median width from
0.538 to 0.650 mm, giving from ratio of

234
.035
.040
.035
.045
.030
.038
.038
.025
.030

.050
.070
.063
.055
.050
.063
.075
.055
.055

.100
.113
.155
.100
.075
.113
.105
.095
.088

.163
.163
.205
.138
.125
.175
.138
.130
.125

5

.225

.175

0.39 to 0.57 in eighteen mature specimens.
Height of volutions gradually increased.
Proloculus minute, spherical and its outside diameter varies from 0.045 to 0.070
mm. Spirotheca composed of structurelesssingle layer in inner two and a half
volutions, tectum and protheca in the
succeeding outer volutions. Septa numeMeasurements of Pamirina leveniReg. no.
of
specimen

L.

2676-2C
2676-2D
2676-42775-6B
2676-2E
2775-2B
2775-12775-2C
2676-9A
2775-3*2676-8A
2775-8A
2676-3*2676-12A*
2775-4A*
2676-2F*
2676-12B*
2775-4B*
2676-9B*
2676-6*.313.3500)
.300
.325
.270
. 225(?)
.325
.250
.275
.300
.230
.238
.238
.225
.200
.225
.110
.163
.180
.175

w.

Half length

R.

.650
.613
.625
.650
.600
.575
.638
.575
.538
?
.550

.48
.57
.48
.50
.45
.39
.51
.43
.51

.500
.439
.400
.313
.225
.300
.313
.388

.48
.51
.50
.72
.49
.54
.58
.45

*: immature specimen

.42

P.
.055
.045
.050
.055
.063
.055
.050
.045
.070
.050
.055
· 050(?).050.063· 050(?)
.050
.063
.050
.055

1

2

3

4

5

.038
.025
.025
.023
.035
.033
.030
.030
.025
.038
.025
.030
.038
.025
.038
.045
.025
.033
.038
.030

.060
.055
.035
.050
.060
.050
.050
.063
.050
.065
.045
.063
.050
.038
.050
.075
.050'
.063
.050
.060

.080
.095
.075
.075
.100
.088
.075
.095
.088
.113
.075
.088
.'088
.075
.088
.120
.063
.088
.088
.088

.125
.130
.125
.120
.130
.113
.100
.138
.145
.150
.113
.120
.125
.115

.163
. 175(?)
.163
.175

.170

672.

Ancestor of the family Verbeekinidae

13

KOBA YASHI, sp. nov. in mm.

Radius veoctor

Form ratio of vol.

1

2

3

4

5

.050
· 050
.050
· 045
.050
.050
· 050
. 050
.055

.088
. 095
.080
. 075
.075
.095
. 088
.080
.088

.155
. 155
.145
. 140
. 125
.165
. 138
.128
.145

.238
. 263
.225
'225
.213
.238
. 213
.213
.200

.350

. 300

.70
.80
.70
1.00
.60
.75
.75
.50
.55

rous, plane and unfluted throughout
growth. Septal counts of the first to
the fourth whorls 8, 11, 13 and 15, respectively. Chomata weakly developed or
almost lacking in the outer whorls ofsome specimens. Tunnel low and pathregular.
Remarks:- The present species is simi-

Fig. in PI.

2

3

4

5

.57
.74
.78
.73
.67
.66
.86
.69
.63

.65
.73
1. 07
.71
.60
.68
.76
.75
.60

.68
.62
.91
.61
.59
.74
.65
.61
.63

.64

.58

PI. 1, Fig. 1
PI. 1, Fig. 2
PI. 1, Fig. 3
PI. 1, Fig. 4
PI. 1, }?ig. 7
PI. 1, F!g.. 8.' PI. 1, Fig. 9.
.PI. 1, Fig. 10
PI. 1, Fig. 12

lar to famirina tethydis in the spirothecal structure, number of whorls,
chomata and others. P. leveni differsfrom the latter by more slender shell,
shorter axis of coiling and smaller
form ratio of volutions throughout thegrowth. It is also easily distinguishedfrom P. darvasica by the shape of shell

KOBA YASHI, sp. nov. in mm.

Radius vector

Form ratio of vol.
Fig. in PI.

1

2

3

4

5

1

2

3

4

5

.075
.055
.045
.063
.063
.055
.050
.050
.063
.070
.050
.050
.055
.050
.070
.063
.045
.063
.045
.070

.110
.088
.075
.100
.095
.095
.088
.080
.088
.100
.095
.088
.100
.095
.100
.100
.075
.113
.075
.095

.163
.138
.125
.150
.150
.155
.140
.145
.155
.170
.163
.150
.150
.150
.138
.170
.125
.170
.125
.163

.263
.225
.200
.245
.275
.245
.220
.250
.250
.275
.250
.225
.255
.238

.363
.345
.363
.375

.50
.45
.56
.37
.56
.59
.60
.60
.40
.54
.50
.60
.68
.50
.54
.72
.56
.52
.83
.43

.55
.63
.50
.50
.63
.53
.57
.. 78
.57
.65
.47
.71
.50
.39
.50
.75
.67
.56
.67
.63

.49
.69
.60
.50
.67
.56
.54
.66
.56
.64
.46
.58
.58
.50
.64
.71
.50
.51
.70
.54

.48
.58
.63
.49
.47
.46
.45
.55
.58
.55
.45
.53
.49
.48

.45
.51 (?)
.45
.47

.338

.50

PI. 1, Fig. 13
Pl. 1, Fig. 14
PI. 1, Fig. 15
PI. 1, Fig. 16
PI. 1, Fig. 18
PI. 1, Fig. 19
PI. 1, Fig. 20
PI. 1, Fig. 21
PI. 1, Fig. 22
PI. 1, Fig. 23
Pl. 1, Fig. 27
PI. 1, Fig. 28
PI. 1, Fig. 29
PI. 1, Fig. 30
PI. 1, Fig. 31
PI. 1, Fig. 32
PI. 1, Fig. 33
PI. 1, Fig. 34
PI. 1, Fig. 37
PI. 1, Fig. 38

Fumio KOBAY ASH]

14

and spirothecal structure. This species
resembles many species of Eostafiella,but it differs from them by the spirothecal structure, thicker spirotheca andslightly larger shell. It is also similar tosome species of Pseudoendothyra whichhave weakly recrystallized spirotheca andshell. The present genus is however,not subjected to the secondary mineralization. The specific name is dedicatedto E.Y. LEVEN who proposed the present
genus for his extensive paleontologicalstudy on fusulinaceans.
Occurrence:- Abundant in two levelsof the Hanagiri Limestone, associatedwith Pamirina tethydis. Rarely foundfrom Ohtaki associated with Misellina
dyhrenfurthi (DUTKEVICH), M. sp. A andothers.
Reg. nos.: 2676-2C (Holotype), 2676-2
D, 2E, 2F, 3, 4, 6, 8, 9A 9B and 12, 27751, 2B, 2C, 3, 4A, 4B, 6B and 8 (Paratypes).
Pamirina darvasica LEVEN
PI. 2, Figs. la, Ib, 2-3
1970. Pamirina darvasica LEVEN, Palaeont.
Jour., No.3, pp. 23-24, PI. 1, Figs. 112, 21-24.

Small shell with six whorls, subspherical in shape and with convex lateralsides, fiat or weakly depressed polar
regions and broadly arched periphery.
Rotation of axis of coiling slightly observed in juvenile stage. Axial length
reaches 0.825 mm, median width 1.088 mm
and giving a form ratio of 0.76. Shell
expands gradually. Spirotheca composedof structureless single layer in proloculus
and succeeding one to two and a half
volutions, tectum and protheca in two
and a half to four and a half volutions,and tectum and fine alveolar keriotheca
in four and a half to the last volutions.
Septa plain and unfiuted.
Chomataweakly developed. Tunnel low and pathregular.
Remarks:- The present material is specifically identical with the original specimens by LEVEN, but the former has
slightly larger shell and weaker chomata.
More detailed comparison with the original materials are postponed until more
numerous specimens are accumulated.
Occurrence:- Rarely found in the limestone pebbles of the Triassic intraformational conglomerate distributed in northof Itsukaichi, west of Tokyo. Associatedfusulinids are Misellina dyhrenfurthi

Explanation of Plate 1(All figures are x 65)
Figs. 1-12. Pamirina tethydis KOBAYASHI, sp. nov.
1-2, 4, 6-10, 12: Axial sections of the paratypes; 3: Axial section of the holotype; 5:
Sagittal section of the para type; 11: Tangential section of the paratype.
Loc.: All from the Hanagiri Limestone, Shomaru Pass. Reg. nos.: Figs. 1-12, 2775-2A,
2676-11, 2676-13, 2775-7, 2676-8B, 2775-2D, 2676-2A, 2775-6A, 2676-1A, 2676-2B, 2775-4C and2676-1B, respectively.
Figs. 13-38. Pamirina leveni KOBAYASHI, sp. nov.
13: Axial section of the holotype; 14-23, 27-34, 37-38: Axial sections of the paratypes;
24, 35-36: Sagittal sections of the para types; 25-26: Tangential sections of the paratypes.
Loc.: All from the Hanagiri Limestone, Shomaru Pass. Reg. nos.: Figs. 13-38. 2676-2C,
2676-2D, 2676-4, 2775-6B, 2676-2G, 2676-2E, 2775-2B, 2775-1, 2775-2C, 2676-9A, 2775-3,
2676-3A, 2775-8B, 2676-3B, 2676-8A, 2775-8A, 2676-3, 2676-12A, 2775-4A, 2676-2F, 2676-12B,
2775-4B, 2676:'10A, 2676-10B, 2676-9B and 2676-6, respectively.

.KOBAYASHI:

Ancestor of the Family Verbeekinidae

Plate 1

672. Ancestor of the family Verbeekinidae(DUTKEVICH), M. sp. A and others.

Reg. nos.: 2448-1, 2377C-1, 2448-4.
References
CHOl, D. R. (1972): Classification and phy.
logeny of Genus Miseltina with descriptionof some Misellina from the Lower Per·
mian in the Southern Kitakami Mountains,
Japan. Jour. Fac. Sci., Rokkaido Univ.,
ser. 4, Geol. and Mineral., vol. 15, nos.
3-4, p. 625-642, pI. 1.
FUJIMOTO, H. (1936): Stratigraphical andpaleontological studies of the Titibu
System of the Kwanto Mountainland.
Part 1, Sci. Rep., Tokyo Bunrika Daigaku,
sec. C. vol. 4, p. 157-188.
IGo, HISAYOSHI and KOBAYASHI, F. (1974):
Carboniferous conodonts from the Itsu·
kaichi district, Tokyo, Japan. Pal. Soc.
Japan, Trans. Proc., N. S. no. 96, p. 411426, pI. 56.
ISHII, A. (1962): Upper Paleozoic of the Nakatsu.gawa area, Okuchichibu, Kant6·
massif. Bull. Chichibu Museum, Natu.
Rist., no. 11, p. 1-21. (in Japanese)
KANUMA, M. (1960): Stratigraphical andpale:mtological studies of the southernpart of the Hida Plateau and north·easternpart of the Mino Mountainland, Central
Japan. Part 2, Paleontology, No.4,
Ozawainellinae, Schubertellinae, Verbe·
ekininae and Neoschwagerininae. Bull.
Tokyo Gakugei Univ., vol. 11, p. 55-73.
pis. 10-13.
KOBAYASHI, F. (1975): Palaeofusulina.Rei.
chetina fauna contained in the pebbles of
intraformational conglomerate distributedin the Okutama District, West Tokyo.
Pal. Soc. Japan, Trans. Proc., N. S. no.
100, p. 220-229, pis. 22-23.
LEVEN, E.Y. (1970): On the origin of theadvanced fllsulinid. Palaeont. Jour. Acad.
Sci. U. S. S. R., no. 3, p. 18-25, pis. 1-2.
(in Russian)
MORIKAWA, R. (1955): Schwagerininae in thevicinity of the Shomaru Pass, easternpart of Kwanto Mountainland, Central
Japan. Sci. Rep., Saitama Univ., vol. 2,

15

no. 1, p. 45-114, pis. 5-15.
OZAWA, T. (1970): Notes on the phylogeny
and classification of the superfamily
Verbeekinoidea (Studies of the Permianverbeekinoidean Foraminifera-I). M em.
Fac. Sci., Kyushu Univ., Ser. D, Geol.,
vol. 20, no. 1, p. 17-58, pis. 1-9.
RAUSER·CHERNOUSOVA, D.M. (1963a): Einige
Fragen zur Evolution der Fusulinideen. In
KOENIGSWALD, G. H. R., BUNING, W. L.
and WAGNER, C. W. (Editors), Evolutionary
trends in Foraminifera. p. 45-65, Elsevier.
- - (1963b): Historical development offusulinids and the boundary of stratigra.
phic subdivision. Problem of Miaopale.
ontology, vol. 17, p. 3-12. (in Russian)Ross, C.A. (1967): Development of fusulinid(Foraminiferida) faunal realms. Jour.
Pal., vol. 41, no.6, p. 1341-1354.
ROZOVSKAY A, S.E. (1948): Classification andsystematic characters of the genus Tri·ticites. Doklady Acad. Sci. U.S.S.R., vol.
59, no. 9, p. 1635-1648. (in Russian)- - (1949): On the evolution of the shellwall in the family Fusulinidae. Trans.
Palaeont. Inst. Acad. Sci. U.S.S.R., vol.
20, p. 345-348. (in Russian)- - (1950): Genus Triticites, its evolution
and stratigraphical importance.
ibid.,
vol. 26, p. 1-80, pis. 1-10. (in Russian)- - (1969): On the revision of the order
Fusulinida. Palaeont. Jour. Acad. Sci. U.
S.S.R., no. 3, p. 34-44. (in Russian)
SAKAGAMI, S. (1956): Fusulinids from thelimestone conglomerate in the Yagooki
valley, Tamanollchi, Hinode·mura, Nishi·
tama·gun, Tokyo·to, Japan.
Pal. Soc.
Japan, Trans. Proc., N. S., no. 24, p. 259265, pI. 37.
SUYARI, K. (1962): Geological and paleonto·
logical studies in central and eastern Shi.
koku, Japan. Part 2, Paleontology Jour.
Gakugei, Tokushima Univ., vol. 12, p. 163, pis. 1-12.
TAKAOKA, Y. (1966): Fusulinid from the
Mt. Tatoro, Mt. Kano, Mt. Futago and
Mt. Shiraishi areas of the Kwanto·massif,
Central Japan. Bull. Chichibu Museurr.,
Natu. Rist., no. 13, p. 39-70, pis. 5-16.

16

Fumio KOBAY ASH!

Akiyoshi :ekE,
Itsukaichi 11.13*,
Nishi-hosoo ggW~,
Fukazawa ~iR,
Hanagiri:re:w.J, Kami-kutsu J:~jj,
Shomaru :iE);{.,
Kutsu-zawa ~jjiR,
Shimo-kutsu """f~jj,
Ohtaki :kit,
Chichibu ~jC,
Uzaradaira ~3f,
Futagoyama rJlj,
Shimadani ,Ii!;~,Tamanouchi :KO)P:J,
Kameiwa 1I!.¥r,
Kaerimizu 1m!)*=

Explanation of Plate 2
Figs. 1-3. Pamirina darvasica LEVEN
1, 3: Axial (slightly tangential) sections; 2: Oblique section. Loc.: All from Itsukaichi.
la, 2-3: x 20, 1b (the same specimen as 1a): x 65. Reg. nos.: 1; 2448-1, 2: 2377C-1, 3:
2448-4.
Figs. 4-7, 12-13. Misellina dyhrenfurthi (DUTKEVICH)
4-7, 12: Axial sections; 13: Sagittal section. 4a, 5-6, 12-13: x 20, 7: x 25, 4b (the same
specimen as 4a): x 65. Loc.: All from Itsukaichi, except for 7 from Ohtaki. Reg. nos.:
1: 2448-3, 5: 2448-2, 6: 2448-5, 7: 2715-6, 12: 2448-7,13: 2448-12.
Figs. 8-11, 14. Misellina sp. A
8, 10-11, 14: Axial sections; 9: Sagittal section. 8a: x 25, 9-11, 14a: x 20, 8b, 14b (thesame specimens as 8a, 14a): x 65. Loc.: All from Itsukaichi, except for 8 from Ohtaki.
Reg. nos.: 8: 2715-4, 9: 2377C-5, 10: 2399-2-8, 11: 2399-2-9, 14: 2377d-1.
Figs. 15-16. Misellina claudiae (DEPRAT)
15: microspheric axial section; 16: megalospheric axial section, both are x 65. Loc.: Akiyoshi (Kaerimizu). Reg. nos.: 15: Ak-2, 16: Ak-3
Fig. 17. Thin section of limestone containing many specimens of Pamirina leveni KOBAYASHI,
sp. nov. and others. x 25. Reg. no.: 2676-2.

KOBAYASHI:

Ancestor of the Family Verbeekinidae

Plate 2

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 105, pp. 17-26, pis. 3, 4, April 30, 1977

673. MANCHUROCERAS FOUND IN SOUTH KOREA WITHNOTES ON THE MANCHUROCERATIDAE AND THE
MANCHUROCERAS PROVINCE
To the Memory of Takuji SHIRAKI, 1891-1971**

TEIICHI KOBA Y ASH!
Japan Academy, Tokyo

ifL«.Wi!E M anchuroceras

c M anchuroceras 1K:to J: Lf Manchuroceratidae ve:: . ,,?~, -C :**~=O)l*~rPoO):, He:::. Manchuroceras ~B'A..,-C:'~'00)-c:'gC.lI!Ii:-t00 :t:'O)t:pve:::. 2 ¥rfffiiJ:j;
0, *~;fSIHifi~O)~il"l;P]jl(!6-t:pO)~t-= t.d~$~7f;-t to) C ~x. Gn 0, Manchuroceratinae

It*~O)7i-v=- Kerkoceras, Parapiloceras :to J: Lf~ G < Coreanoceras ~§fft-t 0 0 Man·
churoceras Itrf:l1l~~*c.fi€:-¥v=-lt* 7 :; 7 iJ' G ~ -< =j -\" i -C:'51m L, 16 til!J;U.: ~~ L -C ~,00 ~v=- 31 A -< =- ne:::. 2 $iJ;j; I) ~~O)[l§::tff!fv=- %'i'ldJ''Jt.cliJ(JEM-C:'2b 00 .: c ~~[jj'j t:.
L -C, ~t*[l§f!fiJ' G~t*Ie:::.lt Piloceratidae iJ~JA 0'-;(PL. Piloceras, Allopiloceras It<

31:7.-<=-7· :;!--:7. r ~ n=-N..Lf.

<,:O)j:fu-c:'jilijliJ(JEMlKiJ~~il'L-C\,'t-=oIn South Korea Middle Ordovician
cephalopods are well represented, butlittle has been known of Lower Ordovician
ones, notwithstanding that piloceroids
are so common in the Hwangho basinthat the Wolungian was the so-calledpiloceroid age. Therefore I called attention to the presence of Manchurocerasin South Korea about ten years ago(KOBAY ASHI, 1966), because some specimens of the genus were contained in

n

'N*~-

SHIRAKI'S collection.
Although theirlocalities should be allocated in the Ordovician sequence exactly by field obser·
vation in future, it can be judged thatthey are somewhere in the middle partof the Jikdong (or Chiktong) limestoneor the Maggol (or Makkol) limestone s.
1., unless South Korea is an exception
in the Manchuroceras province. Because
Manchuroceras is a key to the Wolungian
fauna, the Mauchuroceras horizon must
be below the Maggol limestone s. str, or
the upper Jikdong limestone containing
Chazyan Sigmorthoceras and above theupper Tomkolian or upper Tremadocian
Protopliomerops zone yielding a few minute endoceroids and ellesmereoceroids.
It is probably higher than the Clarkellazone in the basal jikdong limestone whichapparently lacks cephalopods.
The Wolungian cephalopods found inSouth Korea are determined here as follows:
1. Manchuroceras d. platyventrum(GRABAU)* Received Jan. 13, 1977; read Jan 22, 1977,
in Tokyo.
** The late Takuji SHIRAKI was the first to
discover graptolites in Korea at Hwangji.
ri, Samcheok.gun, Kangweon.do. It was
during his field work for the graduation
thesis from the University of Tokyo, 1921.
Since then he has done distinguishedcontributions to the geology of the coal·
fields in Korea and their exploitation.
He was very helpful for me to provide
a large amount of Cambrian and Ordo·
vician fossils including those here de·scribed. Now I take a pleasure to dedicate
this paper to him.

17

18

Teiichi KOBAYASHI

2. lvlanchuroceras kobayashii OBATA
3. Manchuroceras tenuise sp. nov.
4. Manchuroceras hanense sp. nov.
5. Mancnuroceras? sp.
A1anchuroceras has long been thought
to be indigenous to the eastern part of
the Hwangho basin, namely in Hopei (orChihli), Shantung, Liaoning (South Manchuria) and North Korea. Now it is
known from the Kilienshan in the west.
To the south it is distributed as far as
the Himalayan and Tasman troughs
through the Yangtze basin (Central China
to Viet Nam). In short, it is a Wolungian
genus characteristic of the late or middle
Canadian cephalopod faunas on thewestern Pacific side, ignoring Liaotungoceras sp. listed in an Arenigian fauna ofArgentina by HARRINGTON and LEANZA(1957).
Beside this genus the family Manchuroceratidae comprises Kerkoceras, Parapiloceras and probably Coreanoceras. The
first of them is the closest ally to Alanchuroceras which was recently proposedwith Kerkoceras bicostatum from Central
China. The second is a highly specialized
genus known of monotypic Parapiloceras
shimizui from North China. The third
genus is represented by several species
in North Korea and North China (Hopei,
Shantung, CHEN, 1976). Recently it was
found in Central China where Coreanoceras trapezoidum and C. reniforme were
described from Szechuan (CHEN and Lru,
1974). In addition Coreanoceras lunatum
FLOWER, 1956 is known from the Canadianof Maryland, North America.
Of the genus iVlanchuroceras an isolate
occurrence is M. asiaticum BALASHOV,
1962 from the Krivolusky formation in thecentral Siberian platform which may bean early Middle Ordovician survivor. Its
cross section, however, reveals a different
outline of its endosiphocone. Emmonsoceras aristos (FLOWER, 1955) from Lake

Champlain, eastern North America may
be another Chazyan derivative from theManchuroceratidae stock as consideredby FLOWER (1958). Thus it is probablethat the related genera were survived in
Siberia and North America until theearly Middle Ordovician times. but in
the Canadian epoch the distribution of
the family was restricted to the western
Pacific regions exept for a stranger,
Corenoceras lunatum.
In the same epoch the Piloceratidae
were distributed extensively on the Atlantic side, from Spitzbergen to the Appalachian geosyncline through the Bearislands, SmqJla island of Norway, Scotland
and Newfoundland and further to thewest into Colorado and Texas throughthe Ozark hills. Allopilloceras baylorense
and Piloaras sp. in the EI Paso limestone,
Texas and several species of Kirkocerasand Utoceras in the Manitou limestone,
Colorado (ULRICH et al. 1943) are therepresentatives of the family in southwestern North America.
Furthermore, Pilocems tasmaniense TEICHERT, 1947 occurs in the upper Canadianof Tasmania in association with two
species of Manchuroceras beside Utoceras? sp. Allopiloceras calmus TEICHERT and
GLENISTER, 1954 is described from the
Emanuel limestone in Kimberley. western
Australia. The last is accompanied by
Hardmannoceras lobatum, while in the
Langkawi islands, Malay Discoceras(Hardmanoceras?) chrysanthimum KOBAYASHI,
1959 and Teiichispira kobayashii YOCHELSON and JONES, 1968 are contained in thesame limestone of Pulao Langgon, butneither piloceroid nor manchuroceroid.
Teiichispira is a very distinctive Macluritacean type of gastropod operculum
contained in limestones of upper Canadian
to lower Chazyan or basal Middle OrdoVICIan age. It is known now to be widelydistributed from Malay to Newfoundland

673.

Korean Manchuroceras; Manchuroceratidae

through northern Australia (Kimberley
and Toko range), Tasmania, Utah andAlabama (GILBERT-ToMLINSON, 1973).
On the basis of the above mentioned
facts it can be concluded that there were
two cephalopod provinces in the late
Lower Ordovician period. The Manchuroceras province on the western Pacificside was connected with the Piloceratidae
province at Tasmania. Additional evidences on Teiichispira and Allopiloceras
show that the connection between the
Atlantic and Pacific faunas were maintained further through northern Australia. Neither manchuroceroids nor piloceroids are, however, known from western
Asia, Europe exclusive of the Atlantic
margin and the Mediterranean region(TEICHERT, 1964). It is interesting to seethat the leading trend of evolution wasjust opposed to each other between thetwo groups. Namely, the manchuroceroids
are mostly orthoconic and invariably
circular to dorso-ventrally depressed,
whereas piloceroids are laterally compressed and commonly cyrtoconic.
Family Manchuroceratidae
KOBA Y ASHI, 1935

Diagnosis:- Siphuncle orthoconic, com~monly moderate in length, but rarely
very breviconic or strongly cyrtoconic,
large in comparison with conch andlocated marginally on ventral side, subcircular or more commonly dorso-ventrally depressed in section, but never compressed laterally; endocone thickened onventral side, accordingly endosiphocone
shifted dorsally; septa slanting dorsally;
septal neck shorter than septal distancewhich is relatively short. Apical. projection often present.
Remarks:- The camerate .portion of
the conch is largely destroyed probably

19

because the shell and septa were thinand camera had no deposit. Septa are,however, preserved in some specimens
near the siphuncle in J\!/anchuroceras andCoreanoceras with which the large size ofthe siphuncle and its marginal positionin the conch can be ascertained. Septal
sutures are completely or almost completely effaced on the ventral side insome forms of these two genera.
When I have instituted this family for
Manchuroceras, I pointed out as thedistinctions of the genus from Piloceras,
the siphuncle's straightness or weakcurvature, marginal position, dorso-ventral depression and its endocone's ventral
thickening where a median endosiphoblade exists.
Kel'koceras founded on Kerkoceras bicostatum CHEN and Lru, 1974 from theLower Ordovician of Szechuan is a very
close ally to Manchul'oceras, both having
the same kind of apical projection (KoBA Y ASHI, 1937). Kerkoceras' siphuncle isbrevi conic, dorsoventrally depressed,
straight" and flattened ventrally; its endocrine so· strongly thickened medially
to form a ventral ridge; septal suturestrongly slanting. dorsally. ,The second
species is Kerkoceras magnum CHEN,
1976 from Chingshueiho, Inner Mongolia.
Of Parapiloceras the apical end is unknown, but the genuS is well represented
by some siphuncles of mono typic Pafapiloceras shimizui which are breviconic
cyrtocones resembling. Piloceras and Cassinoceras. The cross section of Parapilocel'as' siphuncle is sub circular, ' butslightly depressed and its ,eIidcone isstrongly thickened on the concave 's.ide
as seen in figs, 1-2, pI. 17,: OBA'TA,1940.
Because Kerlwceras is. br.evicon.ic andbecam;e the cyrtoconic aspecl is 'seen
also in Manchuroceras ex .. M. manchuriensein fig. 14, pI. 33,ENDO, 1932, it. is "r,easonable to consider that Parapilocera$

20

Teiichi KOBAY ASH!

is a member of the Manchuroceratidae. rather than an isolate member of the
Piloceratidae.
Coreanoceras (KOBAYASHI, 1936) may
be an additional genus to the family. It
agrees with Manchurocems in the relatively long orthocone, depressed crosssection of the siphuncle, presense of
apical projection, the ventral thickeningof the endocone and the presence ofendosiphoblades, but it is distinct from
Manchuroceras in the longer apical projection, zigzag pattern of the septal sutureon the ventral flattening, the prominent
ventral ridge of the end ocone and paired
endoshiphoblades, instead of a median
one in Manchuroceras. A ventral alveolus
which appears in the siphuncle of the former genus in a certain stage of growth
is a matter of discussion, whether it isan original or a secondary structure. If
it be original, it shows the evolutionary
trend of siphuncular compartment which
introduced the Chihlioceratidae and the
Allotricoceratidae. Chihlioceras also has
an apical projection which is, however,quite diffferent from those of the Manchuroceratidae, if compared to GRABAU'S
model in his text-figures 4-5, 1922. Until
more will be known of Chihliocems any
further comparison is unable.
Finally, the apical projection is seenin Allopiloceras coarctum in figs. 1-2, pI.
18, though less prominent, and in Cassinoceras triton in figs. 1-2, pI. 5, thoughmuch less so. These are two exceptions
for the Piloceratidae in ULRICH andothers' monograph, 1943.
The Piloceratidae and Manchuroceratidae are two branches which were derived from an unknown common stock,and developed respectively on the Atlantic and Pacific side. The presence orabsence of the apical projection and thedepressed or compressed conch and siphuncle are two morphological differences

between the two families. The third
difference is the commonness of theorthocones or cyrtocones respectively inthe latter and former family. Because
Allopiloceras has the siphuncle nearlystraight and slowly tapering, in the
Piloceratidae, it may be the nearest to
the Manchuroceras.
Genus Manchuroceras (OZAKI),
KOBA YASHI, 1935
1927. Manchuroceras OZAKI, Jour. Geol. Soc.
Tokyo, v. 34, p. 45.
1935. Manchuroceras KOBAYASHI, Ibid. v. 42,
p. 743.

Diagnosis:- Breviconic orthoceracone;
siphuncle large, cylindrical, straight orslightly curved, ventrally marginal, subcircular, subelliptical or ovate and dorsoventrally depressed in various degrees
in cross section; its ventral side commonly flattened; septal neck long; septal
suture oblique to the axis of the siphuncle,
ascending from dorsum to venter; apicalpart of siphuncle conical, provided witha mammilary projection in some forms;
endocone thickened on ventral side toform moderate elevation where a medianendosiphoblade is often present; endosiphocone off-center, shifted to dorsum.
Type-species :-Piloceras wolungense KoBAYASHI, 1931.
Remarks:- When OZAKI (1927) proposed
Manchuroceras with a specimen fromYenchoucheng, east of Liaoyang, South
Manchuria, Northeast China, he describedno species of the new genus, although
he made its comparison with Chihlioceras.
Subsequently in 1935 when I examined
his specimen, I found his interpretationof its internal structure erroneous in that
his tubular alveolus is nothing but arecrystallized part in the ventral side ofthe endocone. Because his specimen was

673.

Korean Manchurocerae; jvlanchl~roceratidae

identifiable with Piloceras wolungense, I
wrote a new generic diagnosis on thebasis of this species. By the way theendosiphuncular segments were unusuallywell preserved in OZAKI's specimen. The
same kind of structure was, however, so
better preserved in Penhsioceras? sp. thatit showed the change of the endosiphucle
and the segments through growth (KoBAYASHI, 1947).
Independently from this type-revision
SHIMIZU and OBA TA (1935) proposed two
new names, Grabauoceras and Liaotungoceras assigning Piloceras platyventrum
GRABAU and Piloceras manchuriense ENDO
as the respective type-species. Theirgeneric description, however, did not appear until next year when they accepted
the synonymy of their Grabauoceras with
Manchuroceras. Of the distinction of
Liaotungoceras from Grabauoceras, i. e.
Manchuroceras, they noted its 0. e. Liao-

tungoceras') curved siphuncle and different
shape of endosiphocone in dorso-ventral
section. But the difference has since
been neither described nor illustrated bythem and no mention was given of Liaotungoceras in OBATA'S paper on the genus
Manchuroceras (1939).
ULRICH et al. (1943) on the other hand
accepted the validity of Liaotungoceras,
but saying that the siphuncle is straightor nearly so, instead of curved. They
pointed out its distinction from AlioPiloceras, but not from Manchuroceras andincluded Manchuroceras endoi in thatgenus. Precisely speaking, Piloceras manchuriense ENDO, 1932 comprises two distinct forms, namely one is straight andcircular in cross section and the othercurved and dorso-ventrally depressed in
section. Because the latter is the holotype, M anchuroceras endoi KOBAYASHI,
1935, must be its synonym, as pointed
out by ULRICH et al. (1943). This meansthat Liaotungoceras founded on Piloceras

21

manchuriense s. str. should be a synonymof lvlanchuroceras, because Piloceras manchuriense s. str. is inseparable from Manchuroceras compressum at least generically.
Distribution:- In adding GRABAU'S species of Piloceras, there were four valid
species in the genus as follows:
Manchuroceras platyventrum (GRABAU,
1922) Hopei, Shantung
lvlanchuroceras wolungense (KOBAYASHI, 1931) Liaoning
Ivlanchuroceras compressum (KOBAY ASHI, 1931) Liaoning
IVIanchuroceras manchuriense (ENDO,
1938) Liaoning
Later OBATA (1939) described five new
species of Manchuroceras (ozakii, ishidai,
yenchouchengense, kobayashii and katsunumai) with the specimens obtained fromthe Liangchia-shan and equivalent formations in Hopei and Liaoning.
CHANG and SHENG (1958) reported an
occurrence of Manchuroceras d. wolungense in the Hunghauyuan limestone on
the Szechuan-K weichou border and recently CHEN and LIU (1974) described
Manchuroceras badongense nov. from thesame limestone in Hupeh. In Northwest
China CHANG (1965) found a new species
Manchuroceras tochuanshanense in the
Duoquanshan limestone, south Kilienshan.
Recently CHEN (1975, 1976) found Manchuroceras sp. in the Lower Ordovicianof the Himalayas, and Manchuroceras
qingshueiheense in the Liangchiashan
formation, Inner Mongolia.
Incidentally, CHANG (1965) described
Wutinoceras shihuigoense,

Armenoceras

sp. and Alanchuroceras tochuanshanensefrom the upper part of the Lower Ordovician Tochuanshan limestone at Shihuigou, Chinghai. Because the first andsecond species look two Toufangian armenoceroids, as pointed out already (KoBAY ASHI, 1969) and because such armenoceroids have been unknown from theTeiichi KOBAY ASH!

22

Canadian or Lower Ordovician not onlyin Eastern Asia but also in the ArctoAmerican region or elsewhere, it is quite
probable that they were derived fromhigher part than the Manchuroceras horizon, and the part of the Tochuanshan
limestone must be already Middle Ordovician in age.
In North Korea I found Manchuroceras
sp. in Deogcheon district (KOBAYASHI,
1931a). In South Korea this genus is
now represented by four or more species
two of which are probably new species.
Thus the Korea-Chinese species attain
more than sixteen specie!;l.
Prior to these recent discoveries TEICHERT (1947) and TEICHERT and GLENISTER (1953) described two new speciesof Manchuroceras (steani and exaratum)from the upper Canadian of Tasmania,
while BALASHOV (1962) named Manchuroceras asiaticum for a new form from theearly Middle Ordovician Krivolusky formation in Central Siberia. The last would
be a solitary survivor of the genus, but
its endosiphocone shows a sharp angle
at the point of an endosiphoblade as seen
in its cross section in fig. 2b, pI. 10,
BALASHOV, 1962, the difference suggesting
the corresponding time-displacement.
Leaving this species for future confirmation, it can be concluded that Manchuroceras is widely distributed in the
Wolungian or the upper or upper-middle
Canadian formations in Korea, China andTasmania on the western Pacific side.

Manchurocems d. platyventru1l1(GRABAU, 1922)
Plate 3, Figures 1a-b
1922. d. Piloceras platyventrum GRABAU, Pal.
Sinica, ser. B, v. 1, fasc. 1, p. 42, pI. 4,
figs. 12a-c (a form), ? figs. 1a-c (bform), ? text-figs. la-e.

Description:- The Korean specimen isa straight siphuncle about 58 mm longand 36mm across, slowly tapering toward
the apex, subcircular in cross section,
but more or less broadly rounded on theventral than the other side. In the longitudinal section the breadth of the siph uncle increases from 28 mm to 33 mm
in the apical 41 mm. The endosiphocone
expands more abruptly, attaining 22 mm
in breadth at the adoral end. The siphuncular wall is almost the same inthickness all around it and composed ofnumerous very thin endosheathes. The
apical portion of the siphuncle is unpreserved. The adapical end of the specimen
is not polished through the endosiphuncle, but the ventral endosiphoblade canbe seen on the broken surface to crossthe endocone.
On the left side of the longitudinal
section from the observer 5 septa and 4
camerae are distributed in the length of
20 mm. The septum forms an angle ofabout 35 degrees with the siphuncularwall. Its funnel is geniculated at theneck of the preceding septum and invaginated into the latter's funnel for about
two-thirds or three-fourths the septal
distance.
This siphunde is almost identical withGRABAU'S type specimen in figs. 12a-c
on pI. 4. This endosiphocone looks somewhat more conical and slender, but sucha difference easily appears by the difference of orientation.
Discussion:- GRABAU (1922) proposed
this species with three specimens, but
for the third one in text-figs. la-e fromthe Peilintze limestone he stated that"the specimen in question may represent
a distinct species", because the annulations are finer, the cross section issubelliptical and the endosiphocone subcentral.
His diagnosis is evidently based on

673.

Korean Manchuroceras; Manchuroceratidae

the two other specimens from the Liangchiashan limestone. Assuming these two
to be the syntypes, the siphuncle isstraight and conical in apical part, but
it becomes sub cylindrical on the adoral
side; cross section sub circular with endosiphocone either circular and subcentral(a form) or excentric and remarkably
flattened ventrally (b form); mammilary
projection present (in b form).
Like M. wolungense it is known thatthe b form in figs. a-c, GRABAU, 1922
possesses a mammilary projection, but
it differs from M. wolungense in the crosssection. As OBATA (1939) excluded thisform from Al. platyventrum, it represents
a distinct subspecies. Accepting that aform in figs. 12a-c, GRABAU, 1922 as thelectotype, it is certainly remarkable that
the Korean specimen fits nicely with it.
Occurrences:- Geasan-chon, Sangjangmyeon, Samcheok-gun, Gangweon-do.

Manchuroceras kobayashii OBATA
Plate 4, Figures 1a-c
1939a. Manchuroceras kobayashii OBATA, Jour.
Shanghai Sci. Inst. vol. 9, p. 105, pI.
9, figs. 2, 3 & 5.
1965. Manchuroceras kobayashii CHAO, ChinKe et al. Chinese cephalopod fossils, p.
29, pI. 4, figs. 12-13.

Description:- A Korean specimen is anadoral part of an orthoconic sipnuncle
which is rapidly expanding with a similarrate as in the holotype. Its ventral side
is remarkably flattend and smooth; 4 to
5 annuli distributed in 20 mm graduallydescending dorsally from venter. Where
the two diameters of the siphuncle measure 28 mm and 38 mm, its wall is as
thick as 4 mm and 7.5 mm respectively
on the dorsal and ventral side, but noventral elevation is recognizable on this23

cross section.
In the ventral view of this specimen
the smooth thin shell of the conch isseen directly contact with the siphuncularwall along the broad flattened band whichis gradually broadened adorally. Startingfrom the lateral margin of this band,
septal annulations are inclined laterally.
They are fairly prominent and about 4mm apart from one another.
Comparison:- This species is well characterized by the straight siphuncle whoselateral diameter is expanding in a greaterrate than any other species of this genus.
The OZAKI'S holotype from Shihmenchai,
Hopei, China measures 133 mm in lengthand 76 mm and 34 mm in the major andminor diameters respectively at the adoralend, but the two diameters are almost
equal in the part 30 to 40 mm apartfrom the apex where they are 29 to 34
mm. In the apical 15 mm the siphuncle
expands very rapidly and the mammilary
projection is unknown. Septal annulations
are countable about 10 near the apex,
but 6 in the adoral part in a distance of
30 mm. They ascend from dorsum toventer gradually where in the latter theydie out. The smooth ventral flattening
attains 25 mm in width. The siphuncularwall is thickened and more swollen inwardly on the ventral side.
The Chinese and Korean forms resemble
M. compressum in the transverse section
so closely that the outlines of their siphuncles and endosiphocones look almost
identical, although M. compressum has
the ventral elevation and endosiphoblade.
Their distinction is quite evident in the
expansion of the siphuncle which is incomparably rapid in this species.
Occurrence :- Yongyeon-chon, Sangjangmyeon, Samcheok-gun, Gangweon-do.

Manchuroceras tenuise sp. nov.

TeiichiKOBAYASHI

24

A specimen from Guemdae-chon is astraight longiconic siphuncle regularly
tapering with the rate of 1 mm of breadthin every 5 mm or so of length; endosiphocone tapering. more abruptly to form
an angle of about 20 degrees at apex
and distinctly ovate and flattened on theventral side in cross section, although
the lateral diameter scarcely exceeds the
dorso-ventral one; endocone twice or
three times thicker on the ventral thanthe other side.; thickened part crossedby median endosiphoblade ; endosiphocone
located closer to the dorsal margin, subtriangular in cross section, but its ventral
wall is also somewhat angulated at the
middle point where the endosiphoblade
opens. Endosheathes are recrystallized
and camerate portion is unpreserved.
. This species can be distinguished fromother species of this genus by its slendersiphuncle and ovate and subtriangularcross sections respectively of the siphuncle and endosiphocone. The great thick·ening of the endocone on the ventral
side and the distinct endosiphoblade
which crosses this part are two characteristics commonly seen in this genus.
Occurrence:- Guemdae-chon, Sanjangmyeon, Samcheok-gun, Gangweon-do.

angulations and 3 shallow depressedintervals discernible in the length of 15mm; endosiphuncle excentric, located atthree-fifths the diameter from venter;endosiphocone expanding rapidly, with
an apical angle of 20 degrees; endocone
thickened more on ·the ventral side thanthe other,· forming a moderate medianelevation. The cross sections of thesiphuncle and endosiphuncle are subcircular at the adapical end, but theformer is elliptical at the adoral end.
This siphuncle is closely allied to thatof Manchuroceras compressum ih the rateof expansion and the ventral elevationof the endocone, but the endocone thickening: is not so much as .in that species,
as it is evident if compared in the samestage of growth. In this species thesiphuncle and endosiphuncle are subcircular at the adapical end, but theformer is ovate and the latter broad andarcuate in that species as seen in fig. 3
e, pI. 17, KOBAYASHI, 1931b. If comparedwith the section in fig. 3c, pI. 17, thecross section of the other end is quite
different in the outlines of the siphuncle
and endosiphocone and the thickeningof the endocone.
Occurrence:- Godoo-am, Gurae-ri, Sangdong-myeon, Yeongweol-gun, Gangweondo.

lvlanchuroceras hanense sp. nov.

Manchuroceras? sp. indet.

Plate 3, Figures 2a-c

Plate 3, Figures 3a-b

Siphuncle 6 cm long, tapering slowly,
approximate Imm in 5mm; 4 weak septal

An imperfect siphuncle resembles thatof Mani:huroceras tenuise sp. nov. A but

Plate 4, Figures 2a-b

Explanation of Plate 3
Figures la-b. Manchuroceras cf. platyventru·m (GRABAU)
Transverse (a) and longitudinaf (b) sections of siphuncle. xl. 5 Geasan-chon.
Figures 2a-c. Manchuroceras hanense sp. nov.
Transverse (a, c) and longitudinal (b) sections of siphuncle~ xl. 5 Godoo-am.
Figures 3a-b. Manchuroceras? sp. indet.
Transverse (a) and longitudinal (b) sections of siphuncle. xL 5 Godoo-am

KOBAYASHI:

Korean Manchuroceras; Manchuroceratidae

3a3bPlate 3

673. Korean Manchuroceras; Manchuroceratidae
tapers more rapidly. The apical anglesof this siphuncie and endosiphocone areabout 30 and 40 degrees respectively. In
cross section the siphuncie is circularand the endosiphocone also sub circularand apparently excentric, although it isdifficult to say the excentricity exactly.
The endocone has a low ventral elevation where an endosiphoblade exist~.
Occurrence;- Godoo-am, Gurae-ri, Sangdong-myeon, Yeongweol-gun, Gangweondo.
References
BALASHOV, Z. G. (1962): Ordovician nautiloids of the Siberian Platform. Leningrad
Univ. Press. 205 pp. 52 pIs.
CHANG, Jih-tung (1965): On some Lower
Ordovician nautiloids from Qilianshan,
Northwestern China. Acta Pal. Sinica,
vol. 13, no. 2, pp. 343-362, pIs. 1-2.
CHANG, M. S. and SHENG, S. F. (1958): The
Ordovician formation along the SzechuanKweichow border.
Acta Geol. Sinica,
vol. 38, no. 3, p. 326-342.
CHAO Chin-Ke, LIANG Hsi-ho, Tsou Hsi-p'ing,
LEI Ts-Kem and CHANG Jih·tung (1965):
Chinese Cephalopod Fossils. Each Groupof Fossils in China. 389 pp. 85 pIs.
CI"IEN, Jun-Yuan (1975): Nautiloidea. Tibet
Scientific Research Party: Reports of
Scientific :Research of Chumulangmafeng,
Paleontology, vol. 1, p. 267-294, 9 pIs.
- - (1976): Advances in the Ordovician
stratigraphy of North China with a brief
description of nautiloid fossils. Acta Pal.
Sinica, vol. 15, no. 1, pp. 55-74, pIs. 1-3.
- - and LIU, Ching-wu (1974): Ordovician
Nautiloidea. Nanking lnst. Geol. Pal.
Acad. Sinica: Handbook of Geology andPalaeontology of Southwest China. p.
134-143, pIs. 58-62.
ENDO, R. (1932): The Canadian and Ordovician formations and fossils of South
Manchuria. U. S. Nation. Mus. Bull. 164,
152 pp, 41 pIs.
FLOWER, R. H. (1955): Status of endoceroid

25

classification. Jour. Pal., vol. 29, no. 3,
p. 329-371, pIs. 32-35.
- - (1956): Cephalopods from the Canadianof Maryland. Ibid., vol. 30, no. 1, p.
75-96, pIs. 59-62.
- - (1958): Some Chazya and Mohawkian
Endoceratida. Ibid., vol. 32, no. 3, p.
433-458, pIs. 59-62.
GILBERT-ToMLINSON, J. (1973): The Lower
Ordovician gastropod Teiichispira in
Northern Australia. Bur. Min. Res. Geol.
Geophy., Canberra, Bull. 126, p. 65-88,
pIs. 29-34.
GRABAU, A. W. (1922): Ordovician fossils
from North China. Palaeont. Sinica, serB, v. 1, fasc. 1, 127 & 4 pp, 9 pIs.
HARRINGTON, H. J. and LEANZA, A. F. (1957) :Ordovician trilobites of Argentina. Dept.
Geol. Univ. Kansas, Sp. Publ. 1, 276 pp.
KOBAYASHI, T. (1931a): Studies on the Ordovician stratigraphy and palaeontology ofNorth Korea with notes on the Ordovician
fossils of Shantung and Liaotung. Bull.
Geol. Surv. Chosen (Korea), vol. 11, no.
1, 60 & 4 pp. 9 pIs.
- - (1931b): Studies on the stratigraphy
and palaeontology of the Cambro-Ordovician formation of Hualienchai andNiuhsintai, South Manchuria. Japan. Jour.
Geol. Geogr., vol. 8, p. 131-189, pIs. 16-22.
- - (1935): Restudy on Manchuroceras witha brief note on the classification of
endoceroids. Jour. Ge"ol. Soc. Japan, vol.
42, no. 506, p. 736-752, pIs. 3-4.
- - (1936). Coreanoceras, one of the most
specialized piloceroids and its benthonicadaptation. Japan. Jour. Oeoi. Geogr., vol.
13, p. 185-195, pIs. 22-23.
~ (1937): Contribution to the study of
the apical end of the Ordovician nautiloid.
Ibid., vol. 14, pO. 11-12, pIs. 1-2.
- - (1947): An observation of the endosiphuncular structure of an endoceroid.
Ibid,. vol, 20, p. 13-18, pI. 4.
- - (1959): On some Ordovician fossils from
Northern Malaya and her adjacence.
Jour. Fac. Sci. Univ. Tokyo, sec. 2, vol.
11, pt. 4, p. 387-407, pIs. 24-27.
- - (1966): The Chosen group of South
Korea. The Cambro-Ordovician forma-

26

Teiichi KOBAYASHI

tions and faunas of South Korea, Part 10,
Sec. A. Ibid., sec. 2, vol. 16, pt. 1, 84 pp.
- - (1969): The Ordovician of Eastern Asiaand other parts of the Continent. The
Cambro-Ordovician formations and faunasof South Korea, Part 10, Section D. Ibid.,sec. 2, vol. 17, pt. 2, p. 163-316.
OBATA, T, (1939a): Parapiloceras, a new
Asiatic piloceroid from North China.
Jubilee Publ, Comm. Prof. H. Yabe's 60
th Birthday, vol. 1, p. 255-260, pI. 18.
- - (1939b): Studies on the Ordovician
limestones in North China. The firstreport. Genus l\fanchuroceras.
Jour.
Shanghai Sci. Inst., vol. 9, p. 89-109,
pis. 7-10.
- - (1940): Studies on the Ordovician
limestones in North China. The second
and third reports. Ibid., vol. 9, p. 273286, 303-318, pis. 34-37.
OZAKI, K. (1927): On a new genus of Ordovician Cephalopoda from Manchuria. Jour.
Ceol. Soc. Tokyo, vol. 34, p. 43-51.
SHIMIZL", S. and OBATA, T. (1936): Studies
on the Palaeozoic cephalopods from Asia,
Part 1. Bull. Shanghai Sci. lust., vol. 5,

p. 85-146.
TEICHERT, C. (1947): Early Ordovician cephalopods from Adamsfield, Tasmania. Jour.
Pal., vol. 21, no. 5, p. 420-428, pI. 58.
- - (1964): Endoceratoidea. MOORE, R. C.
(ed.) Treatise on Invertebrate Paleontology, Part K, Mollusca 3, p. 160-189.
- - and GLENISTER, B. F. (1952): Fossil
nautiloid faunas from Australia. Jour.
Pal., vol. 26, no. 5, p. 730-752, pI. 108.
- - and - - (1953): Ordovician and Silurian
cephalopods from Tasmania, Australia.
Bull. Am. Pal., vol. 34, no. 1411, 66 pp,
6 pis.
- - and - - (1954): Early Ordovician cephalopod fauna from Northwestern Australia. Ibid., vol. 35, no. 150, p. 7-91, pis.
1-13.
ULRICH, E.O., FOERSTE, A. F. and MILLER,
A.K. (1943): Ozarkian and Canadian
Cephalopoda, Part 2, Brevicones. Ceol.
Soc. Am. Sp. Pap. no. 49, 240 pp. 69 pis.
YOCHELSON, E. L. and JOl'';ES, C. R. (1968):
Teiichispira, a new early Ordovician
gastropod genus. U.S. Ceol. Surv. Prof.
Pap. 613-B, p. 1-15, pis. 1-2.

Place Names in China (C) and Korea (K)Chinghai 'W'fIa (C), Chingshueiho m7J'(iDJ (C), Chumulangmafeng ~~Wn!ljJi! (C),
Deogcheon, Teokcheon ~11i (K),Gangweon-do, Kangweon-do lImO! (K),
Geasan-chon,
Kyesan-mal ~Ili;ft (K),
Geumdae-chon, Kuntae-chon ~-Er;ft {K),Godoo:am, Kotuoam
f.liiiJ(~ (K),
Gurae-ri, Kurae-ri :iIJIH[ (K),
Hualienshal jd!!~ (C),
Hunghuayuan l'J:
:re~ (C),Hwangji-ri ~iI!!.!! (K),Jikdong, Chiktong fililOJ (K),
Kilienshan, Qilienshan
1ilIl;il1li (C),Liangchia-shan ~*Ili (C),
Maggol, Makkol ~ilOJ (K),Niuhsintai 4'-'L'-Er(C),
Samcheok-gun =~m\ (K),
Sangdong-myeon l:JI:[oo (K),
Sangjang-myeon l:~oo(K),Shihmenchai Er'~ (C),Yenchoucheng ~1'1'llAlt (C),
Yeongweol-gun ~~IDI (K),
Yongyeon-chon ~1iIil1;ft (K)The author's thanks are due to Ass. Prof. 011 Geunwhan of Masan College of Education
for assistance in modern romanization of Korean place names.

Explanation of Plate 4
Figures la-c. Manchuroceras kobayashii OBATA
Longitudinal (a) and transverse (b) sections and ventro-Iateral view (c) of siphuncle.
xl.;) Yongyeon-chon.
Figures 2a-b. l\fanchuroceras tenuise sp. nov.
Transverse (a) and longitudinal (b) sections of siphuncle. xl. 5 Guemdae-chonKOBAYASHI:

Korean 11!lanchuroceras; Manchuroceratidae

Plate 4

2a

la

Ib

2b

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 105, pp. 27-54, pIs. 5, 6, April 30, 1977

674. NOTES ON THE MORPHOLOGY OF NEITHEA (CRETACEOUSPECTINIDS) WITH TAXONOMIC REVISION
OF JAPANESE SPECIES*IT ARU HAY AMI
University Museum, University of Tokyo, Tokyo

and
MASA YUKI NODA
Wasadahigashi Junior High School, Oita~:r!ll!*c=fj(~

Neithea O)M1L~c.*:ma:;fi0)7t~~(~:jl}~W: Neithea It!f#~i:!;7.:>!ju!Jjtl~

t.(1iJ(M!lJJVfl.~O) Pecten f::.lt~~n7.:>M:{j\-.a-~'l::>, 1iJ(M!lJJO)*WEi'I':1ge~c.:iOO~Mi~-.a-~~-t
7.:>J::f::~,*m!HI'~;f:t-c:'i:!; 7.:>0 i f;:., ~O)P3~ltlJl.~O) Pectinidae c.Ji'<t.( fJ 7"7 vEil> I?MPl!:~n''O ,t.: c.~;t I?n, jl'f]:wO)~ft.j:i1!!O);jHt-.a-U-t7.:>J::f::m.~C.~;t I?n7.:>o 13*~:It!!O)r

Neithea f::":)\"""(, 3-1" ". ... ~O);fic.O).lt~liJfJef::tC.vj\,'""(7t~~B~!f#-$O);fiO)7tM;G;O)qtTI"Hrt.(-:>t.: o tl:J.lK~£!iO)~}II~f::a:L..t.:~;fi N. alta
It:}O~ I? <*~~ttO);fi-C:', =":)O):;!- - ?/"- 0) 1iJ(M!IJJiI,7tft-t 7.:>~~-.a-ffi L..""(\,' 7.:> N. atava
Iti!t3'it(I".HQ1;: <7t:ffl-t 7.:> ;fi-C:', 13 *-C:'lt N. kanmerai c. IIflin ""(\, ,tdlllil'': nf::;f§~-t 7.:> ii',
~iII*7t!lf.<O)MtI~~ c. ;fOilflalJ!If.<O) 1'fEB~iI> I?!iWt.( ~*iI'~ I?nt.:o)-C:'N~.-t 7.:>0$~lII!*f::a:-t7.:>~-.a-N~WL..,

0

~7J(

Introduction

m· !IlfEBmtz

Through the efforts of some Japanese
stratigraphers and paleontologists thestratigraphic distribution and diagnostic
characters of several domestic speciesof Neithea have been much clarified inrecent years, but there are still many
doubts and unsolved problems as to theiraffinity and nomenclature. Evidently
much confusion has been arisen fromthe ambiguous definition of early described European species. For example,
D'ORBIGNY (1847) described many speciesof this genus with fine but highly synthetic illustrations, and his type specimens had long been unstudied. Consequently, many synonymous and doubt-

Neithea DROUET, 1825, is a well-known
pectinid genus for its lovely and readily
distinguishable morphology and its common occurrence in Cretaceous marine
beds. A large number of representatives
have been described from various areasof the world, and the known genericrange extends from the Berriasian to theMaastrichtian, although in eastern Asiaand northern Pacific region the occurrence seemS to be restricted to the Lower
Cretaceous.

* Received Feb. 4, 1977; read Jan. 22, 1977,
in Tokyo.
27

Itaru HAY AMI and Masayuki NODA

28

ful names were proposed and used.
Almost all the hitherto published works
on Neithea by Japanese authors, including HAYAMI (1965), followed the schemeof classification and nomenclature by
WOODS (1903) which is now regarded as
partly inadequate.
Recently DHONDT (1973) accomplished
a comprehensive revision on all theEuropean species of Neithea, and theclassification of this genus was much
improved. Since she observed many
original specimens of European species,
her revision seems to be generally reliable. She also discussed the taxonomic
position of Japanese specimens, suggesting that many of them are specifically
identifiable or comparable with formerly

Yamabe2 Miyako

3Ojika

4 Choshi
5 Sanchu
6Yuasa

7 Katsuura
8 Hagino
9 Susaki
10 Haidateyama
11 Yatsushiro+*•

N. akeloensis•

N. alava•

N. ficalhoi•

N. kochiensis

N.alta.... N. matsumotoi•

N. nipponica

T N. notabilis•

N. syriaca amanoi

Text·fig. 1. Map showing the occurrences of Early Cretaceous
species of Neithea in Japan.

674. Morphology oj Neithea
described European species.
Although the peculiarly designed morphology of Neithea is also interestingfrom other viewpoint than taxonomic,
little studies have actually been madeon its adaptive significance and paleoecology. In 1975, one of us (1. H.) had an
opportunity to observe numerous wellpreserved specimens of Neithea at the
British Museum (Natural History), Sedgwick Museum of Cambridge, Paleontologisk Institutionen of Uppsala University and Some other institutions inwestern Europe. The result of thiscomparative study, though already reflected in the systematic catalogue ofthe Mesozoic Bivalvia from Japan (HAYAMI, 1975, pp. 74-77), is shown in thispaper, and on this ground we intend tosummarize the classification and nomenclature of the Japanese species and todescribe some newly obtained material.
Two of us are responsible for the description of Neithea (Neithea) atava, andother discussions on the radial ribs, shellstructure, adaptive morphology and taxonomic revision are mostly written by
one of us (1. H.).

Previous studies on Japanese
species of Neithea
In Japan nine species of Neithea havebeen discriminated and described. Before
entering the description and discussion,
we are going to review the history oftaxonomic investigations on these species.
About 50 years ago, YABE and NAGAO
in YABE, NAGAO and SHIMIZU (1926) recorded the first occurrence of this genus
from the upper Neocomian Ishido Formation of central Honshu, describing two
small specimens under the name of Pecten(Neithea) cf. atavus ROEMER. Subsequently NAGAO (1934) described two species
from the Aptian-Albian Miyako Group29

of north Honshu and called them Pecten(Neithea) morrisi PICTET and RENEVIER
and Pecten (Neithea) sp.· An Aptian
bivalve fauna from the Hagino Formationof south Shikoku reported by AMANO(1957) also contains two species of Neithea, which were compared by him withPecten (Neithea) morrisi PICTET and
RENEVIER and Pecten (Neithea) quadricostatus SOWERBY.
One of us (HAY AMI, 1965) described
the majority of Early Cretaceous marine
bivalves from Japan with taxonomic
revision of hitherto known species. On
that occasion seven species of Neithea
were named and described, and a classification scheme of this genus, though nowadays partly invalid, was presented witha synoptic list of species in the world.
In the same work P. (N.) morrisi by
NAGAO (1934), P. (N.) sp. by NAGAO andP. (N.) cf. morrisi by. AMANO (1957)
were renamed Neithea (Neithea) jicalhoi(CHOFFAT), Neithea (Neithea) nipponica
sp. nov. and Neithea (Neithea) amanoi
sp. nov., respectively. P. (N.) cf. atavus[sensu WOODS, 1903, and YABE and NAGAO, 1926J was recorded also from theAptian of Miyako and regarded as amember of a newly proposed subgenus
Neithella HA YAMI, 1965 [type-species:
Janira wrightii SHUMARD, 1860J. In addition to these species Neithea (?) aketoensis from the Albian of Miyako,
Neithea (Neithea) matsumotoi from theAlbian of central Kyushu and Neithea(Neithea) kanmerai from the Aptian of
the same area were described as new
species. Owing to a misunderstandingof the type-species of Neithea, an invalid
subgeneric name, Aequineithea HA YAMI, 1965, was proposed in this paper:
it is actually an objective synonymof Neithea, as pointed out by Cox etal. (1969) and DHONDT (1973). HA YAMI in HAY AMI and KA w ASAWA (1967)

~oTable 1.
YABE andNAGAO 1926

NAGAO 1934

Synopsis of the classification of the Japanese species of Neithea

AMANO 1957

I-IAYAMI 1965

HAYAMI andS[[[KAMA andKAWASAWA 1967 SUZUKI 1972

DHONDT 1973

HAYAMI 1975
em. hereN. (N.)

N. (s.l.)aketoensis*

aketoensis

N. (N.)

N. (N.) atava

Iwnmerai*N. (N.) atava

~
~

"'I

~

P. (N.)

N. (N.)

N. (N.)

morrisificalhoi

alpinaficalhoi~"'0:::

N. (N.)

Ei::
.....

N. (N.)~

N. (NI.?)

kochiellsis*

Iwchiensis

N. (N.)
matsumotoi

N. (N.)§~

N. (N.)
matsumotoi*

matsumotoi

~
CI>
~

Os::!

llipponica*

P. (N.) cf.

atava

N. (NI.)
notabilis

N. (NI.)
notabilis

N. (N.)

N. (N.)

N. (N.)

amanoi*amalloisyriacaN. (N.) syriaca
amalloiN. (NI.) cf.
atava

N. (Nl.) cf.

P. (N.) cf.

morrisiatavusP. (N.) aff.

quadricostatus
P: Pecten

N: Neithea

N. (N.)
N. (N.)
quillquecostata lIippollica

N. (N.)

P. (N.) sp.

NI: Neithella* new species

N. sp.

~.........
~
t::1
~

674.

Morphology of Neithea

supplementarily described Neithea kochiensis sp. nov. and N. (N.) matsumotoi
from the basal part of the Shimantogawa
Group of south Shikoku. Furthermore,
several specimens comparable with someof these species were reported to occur
from the core of test boring in central
Taiwan (MATSUMOTO, HAY AMI and HASHIMOTO, 1965), the Choshi Group ofcentral Honshu (SHIKAMA and SUZUKI,
1972), the Lower (or Middle) Yezo Groupof central Hokkaido (MATSUMOTO andOKADA, 1973) and the Arida Formationof Kii peninsula (OBATA and OGAWA,
1976).
DHONDT'S (1973) monograph on theEuropean species of Neithea may also
exert a great influence on the classification and nomenclature of japanese
species, because there are some commonor closely related species between the tworegions. She is of the opinion that
Janira ficalhoi CHOFF AT, 1888, is a juniorsynonym of Janira alpina D'ORBIGNY,
1847, and the name of Neithea (Neithella)
notabilis (MUNSTER in GOLD FUSS) is
generally applicable for the small specimens which were interpreted as belonging to Neithea atava by WOODS (1903)
and japanese authors. As to japanese
specimens, N. (N.) nipponica HAY AMI,
1965, and N. (N.) kanmerai HAY AMI, 1965,
were regarded as synonymous with
Neithea quinquecostata (SOWER BY, 1814)
and Neithea atava (ROEMER, 1839), respectively. Pecten (Neithea) ct. morrisi
by AMANO (1957) was referred to Neithea
syriaca (CONRAD, 1852), while N. (N.)
amanoi HAY AMI, 1965, which is decidedly
conspecific with AMANO'S, was comparedwith Neithea coquandi (PERON, 1877).
Our opinion on her suggestions will befound in the taxonomic revision of thisarticle.
Because the synonymic and homonymic
relation among the taxonomic names in31

these works may be complicated, thesynopsis is shown in Table 1. Since thefull synonymic references were listed in
the separate paper (HAY AMI, 1975, pp.
74-77), they are not repeated here.

Nature of radial ornaments
in Neitlzea
The radial ornamentation on the disk,
which is commonly composed of ribsof two orders, has been regarded asimportant for the distinction and classification of the genus Neithea. Actually,
the number of primary radial ribs ofthis genus is invariably six, although thefirst and sixth ones are sometimes
obsolete in the subgenus Neithella. The
number of secondary ribs on five interspaces also seems to be nearly constantwithin one fossil population (or one
species), being commonly regarded as ataxonomic criterion. The left (or flat)
valves of Neithea have not been so carefully studied as the right (or convex)
valves in hitherto published taxonomic
works, but, properly speaking, the same
importance should be attached to thetwo valves.
In ordinary species of the Pectinidae
the radial ribs interlock with the corresponding grooves of the counter valve
at the ventral margin. It is the samewith Neithea (Neithea) aequicostata (LAMARCK, 1819) and a few other species of
Neithea which have radial ribs of almosta single order of prominence. In manyother species of Neithea with clearly
differentiated radial ribs, on the contrary,
the primary ribs of right valve commonlycorrrespond with the radial elevations(not depressions) of left valve to give
digitations or angulations to the ventral
margin, whereas the ribs of secondary
order always interlock with the groovesof counter valve. Such a mode of radial

32

Itaru HAY AMI and Masayuki NODA

ornaments is clearly observed in N. (N.)
atava [=N. (N.) kanmerai] and N. (N.)
matsumotoi as well as many other species
having primary ribs (Text-fig. 5). Yet, insuch species there is a narrow groove on
the top of each radial elevation in theleft valve. Consequently, six bipartite
ribs or six pairs of contiguous ribs arecommonly developed in the left valve. In
N. (N.) ficalhoi and N. (N.) syriaca amanoi
the surface of left valve is nearly flator broadly concave without conspicuousradial elevations, but the six pairs of
ribs are clearly distinguishable fromother ribs. In such a species the ventral
margin is usually not so strongly digitate.
The total number of radial ribs, therefore, can be expressed as follows: 6+5 n
for the right valve, and 12+5 (n-1) forthe left valve. Although there may be,of course, some minor modifications by
occasional irregular bifurcation or insertion of secondary ribs, the following
number of n appears to be basicallymaintained in the Japanese species.
n=1 Neithea (Neithea) alta sp. nov.
n=2 Neithea (Neithea) ficalhoi (CHOFFAT)
n=3 Neithea (Neithea) atava (ROEMER),
Neithea (Neithea) syriaca a11lanoi
HAYAMI
n=4 Neithea (Neithea) matsumotoi HAYAMI, Neithea (Neithea) nipponica
HA YAMI, Neithea (Neithea) aketoensis HAY AMI
We presume that this grouping by the
number of n also can be applied forvarious foreign species, since the numberof secondary ribs generally seems to beconstant within one species. It is interesting to see that Neithea (Neithea)
regularis (VON SCHLOTHEIM, 1813) fromthe Upper Cretaceous of western Europe
shows a considerable variation in the
differentiation of radial ribs into twoorders (DHONDT, 1973). Six primaries
are discriminated from secondaries insome specimens but not at all in otherones. Nevertheless, the total number ofradial ribs. is almost constant in thatspecies, and" N. (N.) regularis clearly
belongs to the group of n=3. It is suggested that the species, even if showing
phenotypically monotonous radial ribsof a single order, potentially have six
primary ribs. The radial ribbing of anearly species, N. (N.) alta sp. nov. also
indicates this potentiality. Anyhow, thesix primary ribs of Neithea are, actuallyor potentially, a quite stable character
which seems to have controlled by somephylogenetic (or historical) factor rather
than functional.
Such a regularity is scarcely knownin other Late Mesozoic and later pectinaceans. but somewhat similarly digitate ventral margin and corresponding
radial ribbing are seen in the genus
Amphijanira BITTNER, 1901, from theTriassic. The mode and number ofradial ribs in Neithea attract our evolutionary interest, because our knowledge
about the biological background andtaxonomic evaluation of such shellcharacters remains quite insufficient.

Origin and shell structure of Neithea
There are several different opinions
as to the ancestry of the genus Neithea
and its phylogenetic relation with othergenera of the Pectinidae. As reviewed
by DHONDT (1973, p. 9), certain authors
assumed iterative evolution of suchplano-convex pectinids from some stockof the family at different ages, but someothers expected its intimate relation to
Recent Pecten and also to the genus
Weyla BOEHM, 1920, which occurs commonly in the Lower Jurassic of SouthAmerica.

674.

Morphology of Neithea

In fact, the flat or rather concave left
valve, subequilateral outline and someother external characters of Weyla alata(VON BUCH, 1835) and its allies are considerably similar to those of Neithea,
though the radial ribs of Weyla arenever differentiated into two orders. It
is also interesting to see barely differentiated radial ribs and slightly digitate
ventral margin in an early species,
Neithea (Neithea) alta sp. nov., from theLowest Cretaceous of north Japan, because the morphology seems to be primitive for Neithea and possibly transitional between this genus and Weyla.
Yet, nothing is actually known about
the Middle and Late Jurassic representatives of the presumed lineage which
could link Neithea with Weyla.
On the other hand, Neithea is characterized by the development of a pairof strong isodont crural teeth, which is
DHONDTso far unknown in Weyla.
(1973) appears to have recognized thisdifference of hinge structure on thebasis of a silicified specimen of Weyla
taken out by dissolving the calcareous
matrix with some acid. She attachedimportance to this difference and almost
denied the possibility of direct relationship between the two genera. In thisconnection, however, some question
arises. In 1973, one of us (1. H.) joined
to a party of the Expedition of Chiba
University to the Middle Andes conducted by Professor MAEDA and collected
a lot of silicified specimens of Weyla
alata or its allied species from the
Liassic limestones (Pucara Group) in the
Tarma, Huancayo and Lircay areas ofcentral Peru. At many localities theumbonal areas of two valves are rather
rarely preserved, notwithstanding thatthe external surface of the middle andventral areas of disc is well exhibitedand scarcely water-worn. As the result

33

of dissolving the matrix with dilutedhydrochloric acid, several silicified shells
were completely extracted from thelimestone blocks. The shell so far aspreserved are always quite thin andfragile in the umbonal area but comparatively much heavier near the ventral
margin, provided with unusually strong"internal ribs ".
Somewhat similar phenomena havebeen known in various fossils from the
Eocene limestone of Hahajima, Boninislands (IWASAKI, 1975) and also fromthe Chalk of England (HALL and KENNEDY, 1967; CARTER, 1972), though they
are never silicified. As observed andinterpreted by IWASAKI, every specimenof Meotolima from Bonin, which isactually not a limid but a small spondylid, is represented only by the outer
foliated calcite layer, and the inner layerof original shell is not preserved at alL
During his stay in western Europe, oneof us (1. H.) could observe numerous
well preserved specimens of Spondylus
and Neithea from the Upper Cretaceousof Scania, Holland and England. In
almost all the specimens from the Chalkthe umbonal part is unusually thin andapt to be damaged, and only the outerlayer of their original test seems to havebeen fossilized. They have generally
well preserved surface ornamentation
but neither hinge teeth nor muscle impressions are shown on the internalsurface, as pointed out by CARTER (1972).
Unlike the shells of living pectinids the
thickness of the preserved test of Neithea
from the Chalk is nearly isometricallyincreased towards the ventral margin inproportion to the distance from the umbo.
Moreover, the radial ornament is too
sharply impressed on the internal surface even near the umbo to be interpreted as an original feature (Text-fig. 2).
In all the species of Recent pectinids,.

34

Itan~ HAY AMI and Masayuki NODAva lve of "Neilhea quadrico slala" (UMUT
849a, specimen for comparative studies).
Loc. Maastricht, Netherland. The th in anddama ged umbonal area and sharply impressed radial sculpture indicate the inner
layer of original shell was lost in the courseof fossilizat ion . [ x 1J

on the contrary, the shell is almost
entirel y composed of foliated calcite
except for a thin layer of myostracum(B0GG!LD, 1930; T AYLOR, KE NN EDY and
HALL, 1969, etc. ) ; the umbonal area ismuch thickened and radial ornament on
the internal surface is covered and obscured by successively deposited calcite.
Therefore, such a state of preservation
as seen in Neithea from the Chalk andW eyla from the Peruvian Lias would be
difficult to ex plain, if they were similar
to living pectin ids in the shell structure.
The Miyako Group of north Japan
yield various well preserved molluscan
fossils at numerous localities. In theupper part of the Hiraiga Formation(especially, the facies of Orbitolina Sandstone) and the Aketo Formation aragonitic shells are ·not necessarily preserved ,and some specimens of N eithea (Neithea)nipponica show also sharply scu lptured

Te x t -fig. 3. Interna l surface of a left
va lve of N eith ea ficalhoi [ GK. H6 281 J. Loc .
Hiraiga Format ion of the Mi yako Group at
south of Hiraiga, Tanohata-mura, I wate
Pref. The inner layer is, though has alread y been replaced by calcite, completel ypreser ved. Isodont teeth and adductor
muscle scar are observable. [ x 2Jinternal surface (HAY AM!, 1965, pI. 40,
figs. 3, 4). However, entire original
shells seem to be preserved in the lower
part of the Hiraiga Formation, althoughoriginally aragon itic shells and layers
seem to have been replaced by sparrycalcite. A left valve of Neithea (Ne ithea)
fic alhoi therefrom reveals notabl y smoothish internal surface and distinct crural
teeth (Text -fig. 3).
Anyhow, it is worth y of notice thatthe presence of inner aragonitic layer isstrongl y suggested both in W eyla andNei thea. In such cases as the Chalk
Neithea and the Peru vian W eyla it isprobably impossible to determine whether
isodont teeth are developed or not, be cause they, if originally present, had tobe composed of the inner layer. Thisis presumably the reason wh y the isodont
teeth and muscle impressions of N ei thea
are so rarely met with in European
specimens. If the material of Weyla
dealt with by DHONDT (1973) were of
the same state in preservation, the difference of hinge structure between
Nei thea and W eyla would not be firm ly

674.

Morphology of Neithea

conclusive.
Since any other type of shell structure
than foliated calcite and myostracumaragonite is scarcely known in the living
Pectinidae (exclusive of Propeamussium
and its relatives), the traditional referenceof Weyla and Neithea to this family is
open to discussion. So far as the presumed shell structure is concerned, thetwo genera are probably more similar
to Spondylus and Plicatula rather thanmany living genera of the Pectinidae.
Actually, somewhat similarly inequivalve
shell, radial sculpture of two orders andstrong isodont teeth are commonly seenin Spondylus, although the outline ofshell is considerably irregular probably
owing to the sessile life.
The shell structure of the Bivalvia is
generally evaluated as an importantand stable taxonomic character. Some
Late Paleozoic pectinaceans as the Aviculopectinidae are said to be different
from the living Pectinidae in having
inner layers of nacreous or crossedlamellar aragonite (NEWELL, 1938; etc.).
Yet, little has been known about themode of shell structure evolution fromthe aviculopectinids to pectinids, though
it is almost certain that the former is
as a whole ancestral to the latter.
B0GGILD (1930) actually studied the shellstructure of some Mesozoic pectinids:
Jurassic and Cretaceous Chlamys, "Aequipecten", Radulopecten and Entolium aswell as two species of Neithea from theUpper Cretaceous. He pointed out the presence of peculiar zigzag structure in manyof those species (but not in Neithea),
although it is open to doubt whether
the structure is original or not. Because
aragonitic shell is totally absent in the
Chalk and, if preserved, usually replacedby sparry calcite in other Mesozoic
rocks, the shell structure of Mesozoic
pectinids should be carefully restudied.

35

For the time being, Neithea and Weyla
are regarded as belonging to a particulargroup of the Pectinidae, for which atleast a subfamilial name, Neitheinae
SOBETSKI, 1960, is applicable.
The plano-convex outline of Neithea
is at a glance comparable with that ofliving Pecten, suggesting the same adaptive significance. However, the resemblance is probably superficial and does
not seem to indicate any direct phylogenetic relationship, since the shell structure, hinge apparatus and some otherimportant characters are quite different.
Adaptive morphology of Neithea
The peculiarly designed morphology
of Neithea arouses our interest in its
functional significance and mode of life.
Putting aside the unlikeliness of any
direct phylogenetic relation, the planoconvex form of Neithea is notably analogous to that of Pecten in the modern
seas. judging from the occurrences inJapan, western Europe and some otherregions, Neithea probably inhabited on
sandy or chalky substrate of shallow
clean seas and is commonly accompanied
by other pectinids, !imids, spondylids,
belemnites, brachiopods and echinoids.
Thus it possibly occupied a similar
ecological niche to Cenozoic Pecten (s. s.)
and Pecten (Notovola) which are decidedly good swimmers. Could Neithea
also swim well? Since this genus probably became extinct near the end of
Cretaceous without leaving any descendants, only iridirect evidences areavailable to solve this problem.
THAYER (1970) pointed out that thestriated (or quick) muscle of the adductor is considerably oblique to the surface of valves in swimming scallops andthat this is an adaptive structure toincreaee the velocity of clapping motion.

36

Itaru HAY AMI and Masayuki NODA

GOULD (1971) also studied the allometricchange of muscular size and position
through the growth of some living andfossil pectinids; the striated muscle is
proportionally enlarged and moves fromthe postero-dorsal portion to the central,so that the swimming ability is maintained against the accelerating. increaseof shell weight. The orientation andontogenetic allometry of muscles can beused as important criteria for theestimation of swimming ability in extinct
pectinids. Studies on the relative growthof shell thickness to shell size would bealso informative, since its negative allometry may be more advantageous for
swimming habit. Unfortunately, however,these characteristics are very difficult toobserve, chiefly because the inner layerof Neithea was frequently dissolved awayin the course of fossilization. Here only
some remarks will be given about the
paleoecology mainly on the basis of acomparative study with living pectinids.
Actively swimming pectinids in themodern seas seem in general to be characterized by 1) subequilateral shells with
subequal auricles, 2) shallow or obsolete
byssal notch in adult stage, 3) relatively
light shells, 4) considerably wide gapes
along the anterior and posterior margins
of disc to allow the ejection of propulsive
jet current, 5) large umbonal angle, and6) the development of a flange along thedorsal margin of right valve which iseffective to the force of oblique striatedmuscle.
Except for a few species of subgenus
Neithella, the two valves of Neithea arenearly equilateral and the byssal notch,
if present, is very shallow. This can be
regarded as a· suitable morphologicalcharacter for swimming straight ahead.
Yet, the external morphology of Neithea
does not show any of other attributes.
For instance, the umbonal angle of

Neithea (90-110" in LV) is somewhat
smaller than that of Pecten (110-120"),
Placopecten (ca. 120") and Amusium(115-140"). Anterior and posterior gapes
and a dorsal flange of right valve are
undevelopped in Neithea, so far as
several conjoined specimens from Europe
and Gulf Coast region were examined.
The lower-convex shells are not considered as a necessary condition for swimming, because such very actively swimming pectin ids as Placopecten and Amusium have biconvex or even upper-convex
shells. The strongly convex right valveof Pecten is rather regarded as a kindof gryphaeate morphology, which isadvantageous in preventing the commissure from sinking into soupy substrate, as interpreted by some authors(KAUFFMAN in Cox et a1., 1969; STANLEY,
1970; CARTER, 1972). The flat or evenconcave left valve must be effective tohold a layer of sediment grains, whichwere once suspended in the sea waterby means of jet current and settled on
its surface, and consequently to conceal
itself from predators. Very interestingis the fact that the surface of left valve
is frequently more squarrose than thatof right valve of the same individual(Text-fig. 4). It is owing to the more
erect growth lamellae in many speciesof Pecten and because of peculiarlymeshed ornaments in Mizuhopecten. Suchplano-convex shells and different sculpture of two valves must have been formedby certain selective advantage. Neithea
evidently shares such an inequivalve
morphology with Pecten, and similarliving position is strongly suggested. In
the newly obtained material of Neithea(Neithea) atava, serpulids attach themselves more frequently to the surface ofleft valve than right (PI. 5, Fig. 4b).
This phenomenon seems to be coincident
with the tendency in living shells of

674.

Jli[o?"phology of Neithea

37

Te x t-fig . 4. Difference of surface squarrosity between two valves (the same individual)of P ecten ( N otovola) alb ican s. Recent, Tsu ya zaki near Fukuoka . R: right va lve, L: left
va lve . [ x 2, under the same lighting]

Pecten.
Judging from the thickness of theouter layer so far preserved in Chalkand Gu lf Coast specimens as well as
Miyako ones, the original shells of Neithea are generally thicker than that ofliving Peclen. The silicified specimensof Weyla from the Peruvian Lias also
show proportionally much heavier outerlayer. Most species of Ne ith ea and also
Weyla, we s uppose, lived freely on soft
substrates, keeping their commissurenearly horizontal. They probably camouflaged themselves with sediments on
the flat valves like living Pecten, butthey were probably not so excellent
swimmers.
Repository of specimens

The following abbreviations areL!~ed
for the indication of th~ ipstitutes where
the specimens . studied are preserved :UMUT: UniverJ3ity Museum, Univer~sity of Tokyo
NSM: Department of Earth Science,
National Science Museum , Tokyo
GK: Department of Geo lo gy, Kyushu
University, Fukuoka
BMNH: Briti3h Museum (Natural History), London
Systematic descriptions ofJapanese species

In many respects DHONDT'S (1973)
suggestions on the taxonomic positionof Japanese specimens are pertinent andagreeable. However, as the result of
our comparative study between Japaneseand European specimens of each related
species, slight but clear differences areoften found in such morphological characters as the shell size, mode of radialribbing, outline and convexit y of twovalves. Curiously enough, the verticalranges of some common or closely
related species are not always coincident

38

Itaru HAY AMI and Masayuki NODA

between the two regions, notwithstanding that the chronology of the fossil
beds are generally reliable as documented
by ammonites and other indices. In
Europe, according to DHONDT (1973),
many species of Neithea survived from
Early to Late Cretaceous, whereas injapan all the Lower Cretaceous species
seem to have become extinct by the
beginning of Cenomanian. We have noLate Cretaceous representative of Neithea in japan in spite of other abundantmolluscan fossils. Common occurrenceof Aptian and Albian species of Neithea
has been known in Caucasus (DRUSCHIZA
and KUDRJ AVTSEVA, 1960), Lebanon(VOKES, 1947, etc.), Angola (DARTEVELLE
and FRENEIX, 1957, etc.) and some otherareas of Western Asia and Africa, butreliable information on the marine bivalve
faunas of this age in the midway areas
between japan and India is too deficient
to recognize the actual geographic distribution of each species. Moreover,
several species of Neithea from the
Comanche Group of the Gulf Coast region(ST ANTON, 1947, etc.) and its correlatives
in Mexico (BOSE, 1910; ALENCASTER,
1956) are, if not identical with, closely
related to some European and japanese
species. Therefore, it may be dangerous
to draw a hasty conclusion about theroute of migration.
Although the following taxonomic
treatment and consideration on the affinities are not always conclusive, in mostcases we are at least convincing thespecific discrimination and identity among
the fossil populations from japan in viewof .the morphology and distribution.

Neithea (Neithea) aketoensis

HA YAMI, 1965
Synonymy and type.-See HAYAMI, 1975,

p.77.
Remarks.-This is imperfectly known
species, because only one right valve
has ever been known. It was collectedfrom the basal part of the Aketo Formation of the Miyako Group together withseveral specimens of Neithea (Neithea)nipponica. As noted in the original description, the arrangement of radial ribs
appears at a glance to be quite irregular,
and primary ribs can hardly discriminated from secondaries. DHONDT (1973,
p. 13) pointed out the possibility thatthe present species may be synonymouswith Neithea (Neithea) aequicostata (LAMARCK, 1819), but the two must be unrelated, because the ribs of the present
specimen are generally much stronger
and more irregular in prominence. Owing to this irregularity this species wasreferred to Neithea with some doubt(HAYAMI, 1965, 1975). Nevertheless, if
we trace the weakly impressed growthlamellae on the surface at several ontogenetic stages, the ventral margin of
this right valve proves to be polygonallyangulated, as commonly seen in ordinary
six-ribbed species. Moreover, an unusually strong or bipartite ribs radiatesfrom the umbo to each angle of ventral
margin. If we interpret such an unusual
rib as primary and other ribs as secondary, the disposition of radial ribs is notessentially different from that of coexistent N. (N.) nipponica. Incidentally, in
an European species, Neithea (Neithea)
regularis (VON SCHLOTHEIM, 1813) fromthe Upper Cretaceous, according to
DHONDT (1973), there is a great variability in the extent of differentiation ofradial ribs into two orders: primary ribs
mayor may not be discriminated fromother ribs. At present, therefore, wecannot completely restrain the doubt if
the holotype of N. (N.) aketoensis might
be an aberrant individual of N. (N.) nip-

674.

Morphology of Neithea

ponica. Anyhow, it is needed to examine
further the intra populational variationof N. (N.) nipponica on the basis of more
plentiful material in order to confirm thetaxonomic validity of the present species.
Neithea (Neithea) alta HAY AMI, sp. nov.
Plate 5, Figures 1-3
1970. Neithea n. sp.: TAKIZAWA, Bull. Ceol.
Surv. Japan, vol. 21, no. 10, p. 6, listed.
1974. Neithea n. sp.: TAKIZAWA, ISSlllKI andKAT ADA, Geology of the Kinkazan
district, p. 21, listed.
1975. Neithea sp.: TAKIZAWA, Bull. Ceol.
Surv. Japan, vol. 26, no. 6, p. 18, listed.

Types.-Holotype (UMUT MM 5698),
right internal mould, and four paratypes(UMUT MM5699-5702), left externalmoulds, all collected by Takizawa fromthe Kobitawatashi Member of the Ayukawa Formation at Yokone, south coastof Ajishima island, Ojika-cho, Ojika-gun,
Miyagi Prefecture.
Diagnosis.-Small Weyla-like species of
Neithea, characterized by the moderately
convex right valve with 11 radial ribsof almost one order of prominence andremarkably concave left valve provided
with 12 angular and highly elevatedradial ribs forming six contiguous pairs.
Description.-Shell small for the genus,
scarcely exceeding 20 mm in height,
highly inequivalve, nearly equilateral, alittle higher than long.
Right valve
moderately convex, subtriangular withslightly concave antero- and posterodorsal margins of disc, ornamented with
Table 2.

UMUT MM5698
UMUT MM5699

39

11 highly raised and simple radial ribs
which are almost equal in prominence;
intercalary ribs seemingly absent. Leftvalve remarkably concave even at thevery early and later stages, provided
with 12 highly elevated and angularradial ribs which are at a glance similar
to those of Weyla but seem to form six
contiguous pairs; numerous concentricscales crossing the ribs and their interspaces. Posterior auricle of left valve
obtusely truncated and comparatively
small, while other auricles are notactually observable owing to the incomplete material. Apical angle of twovalves apparently small for Neithea,
though it may be due to the secondary
deformation.
Remarks.-Several years ago one ofus (I. H.) had an opportunity to observevarious Jurassic and Early Cretaceous
molluscan fossils from Ojika Peninsulaof north Honshu collected by Takizawaof the Geological Survey of Japan. Although many of them are secondarily
deformed and poorly preserved, several
specimens of a pectinid attracted ourattention, because it appeared to be oneof the earliest representatives of Neithea.
This species is now proposed on thebasis of five specimens from one andthe same locality: one is a right internalmould with partly preserved shell andfour are incomplete left external moulds.
The detailed surface characters of rightvalve and auricles are actually unknown.
Nevertheless, the present species isclearly distinguishable from hitherto
described species of Neithea by the

Measurements in mm [Neithea (Neithea?) alta sp. nov.]

Specimen

LengthHeightThickness

Right in. mould (holotype)Left ex. mould (paratype)

17.5

18.5
15.0+

ca. 4.5

14.0+

40

Itaru HAY AMI and Masayuki NODA

fewer and scarcely differentiated radialribs in right valve and remarkably concave left valve with well developed concentric scales and only six pairs ofangular ribs. Neithea (Neithea) aequicostata (LAMARCK, 1819) and a few otherspecies from the European Cretaceous
have also radial ribs of only a single
order, but the number of ribs. are much
more numerous. In the mode of radialribbing the present species is moresimilar to Neithea biangulata IMLA Y,
1940, from the Hauterivian of Mexico,
particularly to its young specimen. Although the left valve of that Mexica.n
species has not come to· science, theprincipal ribs of the right valve arealmost uniform in prominence, numbering 11 to 13. In N. biangulata, however,the shell becomes much larger ·and one
or two weaker riblets occur on eachinterspace after th.~ middle stage. Suchriblets are hardly expected in the present species, judging from the impression of ornaments on the right internalmould and the radial ribs impressed on
the left external moulds.
Although the radial ribs in the holotype (right internal mould) are apparently
equal in strength, the six contiguous
pairs of ribs on the left valve seem toforetell the characters of six-ribbed
species of Neithea (s. s.). Moreover, theorientation of concentric scales on theleft valve strongly suggests a more orless digitated ventral margin in accordance with the six pairs. The numberof radial ribs on the right valve, which
is 11, may be also meaningful, if one
regards the ribs as homologous with theprimary ribs plus median secondary ribs
in ordinary six-ribbed species. As noted
before, in almost all the species of Neithea(s. s.) radial ribs are basically composedof six primaries plus a multiple of fivefor the secondaries. Anyhow, the sur-face ornamentation of the present species
is interpreted as representing an earlystage of differentiation of radial ribs
into two orders. Taxonomically, thepresent species is tentatively referred to
Neithea (s. s.) with a query, since it possibly constitutes an infrageneric group
together with N. biangulata.
Occurrence.-This species is known
only from the above mentioned type
locality. All the specimens were obtained
from the middle part of the Ayukawa
Formation, where Berriasella sp. wasfound in association (SA TO and T AKIZAWAin T AKIZA w A, 1970). Most of theassociated species of Bivalvia, such as
Limatula akiyamae, Astarte cf. spitiensis,
Parallelodon (Cosmetodon) kesennumensis
and Parallelodon(Torinosucatella) kobayashii are the elements of the Late
Jurassic Kogoshio fauna, which actuallysurvived until the earliest Cretaceous(HAY AMI; 1961). As suggested by T AKIZA W A, the age of this fossil bed is almost
certainly Berriasian. According to DHOc
NDT (1973) the first appearance of Neithea
in Europe is also from the Berriasianbut it becomes common after the ValanginIan. In North America, the above
cited Hauterivian species, N. biangulata,
is probably the earliest. In Japan, asmall pectinid from the Upper Jurassic
Torinosu Group was once referred to
Neithea (KIMURA, 1951), but, as restudied
by TAMURA (1959), it probably belongs
to "Aequipecten". The present specimens,
though poorly preserved, seem to be aninteresting material for the consideration
about the origin and phylogeny of thegenus Neithea as well as the process ofdifferentiation of radial ribs into twoorders.

N.eithea (Neithea) atava (ROEMER, 1839)

674.
Table 3.

41

Morphology of Neithea

Measurements in mm and degrees [Neithea (Neithea) atavaJ

Specimen
NSM MM7348 Right in. mould
UMUT MM5703 Right in. mould
UMUT MM5704 Right ex. mo"Uld
UMUT MM5706 Right vaive
GK. H10295 Right in. mould
GK. H10298 Right in. mould
UMUT MM5703 Left in. mould
UMUT MM5705 Left ex. mould
UMUT MM5706 Left valve
GK. H10291 Left ex. mould
GK. H10292 Left in. mould
GK. H10294 Left ex. mould
GK. H10297 Left ex. mould

LengthHeightThickness

Hinge
lengthApical
angle

103.0
90.9
81. 0
92.4+
94.4
24.0
90.9
81. 0
92.4+
90.0
92.6
63.6
115.0

86.5+
93.8
88.9
96.0+
98.2
30.0
88.6
79.3
91. 0+
95.0
107.0
70.5
118.2

25.5+
21. 3
30.1
30.4
32.4
11. 6

53.0
56.5
?

81°
82°
77°
77°
75°
78°
104°
lOP
97°
102°
98°
105°
106°

Plate 5, Figures 4, 5; Plate 6,
Figures 1, 2
non 1926. Pecten (Neithea) d. atavus ROEMER:
YABE, NAGAO and SHIMIZU, Sci. Rept.
Tohoku Imp. Univ., ser. 2, vol. 9, no.
2, p. 61, pI. 13, figs. 18, 19. [=N.
(Neithella) notabilisJ1952. Neithea atava (ROMER): MATSUMOTO
and KANMERA, Guide books of thegeological excursions.
The lower
valley of the Kuma, p. 45 (non p. 49),
listed.
1954. Neithea atava (ROE1vIER): Cox, Jour.
Paleontology, vol. 28, no. 5, p. 627, pI.
64, figs. 1, 4.
1954. Neithea atava (ROEMER): FUJII, Jour.
Ceol. Soc. Japan, vol. 60, no. 707, p.
.U8, listed (pars).
1954. Neithea atava (ROMER): MATSUMOTO,
Cret. System, Japan. Islands, p. 113(non p. 114), listed.
1956. Neithea (Neitheops) atava (ROEMER):
ALENCASTER, Paleont. Mexicana, no. 2,
p. 12, pl. 2, figs. 5-7.
1960. Neithea atava ROEMER: DRUSCHIZA andKUDRJAVTSEVA, Low. Cret. Fauna from
Caucasus and Crimea, p. 190, pI. 11
figs. 4-6.
1964. Neithea d. quinquecostata (SOWER BY) :MATSUMOTO and KANMERA, Expl. Text.

?
?
56.5
?
?
40.8
40.0+

Geol. Map of Japan, Hina;;;u, p. 99,
listed.
non 1964. Neithea atava (ROMER): MATSUMOTO and KANMERA, Expl. Text, Geol.
Map of Japan, Hinagu, p. 106, listed.
[ = N. (N.) matsumotoi]
1965. Neithea (Neithea) kanmerai HAYANII,
Mem. File. Sci. Kyushu Univ., ser. D,
vol. 15, no. 2, p. 305, pI. 43, figs. 1-5,
pI. 52, fig. 3.
non 1965. Neithea (Neithella) sp. d. N. (N.)
atava (ROMER): HAYAMI, Mem. Fac.
Sci. Kyushu Univ., ser. D, vol. 15, no.
2, p. 307, pI. 44, figs. 1-4, pI. 52, fig.
4. [=N. (Neithella) notabilisJ
1970. Neithea (Neithea) sp.: TERAOKA, Rept.
Ceol. Surv. Japan, no. 237, p. 10, listed.
non 1972. Neithea (Neithella) sp. d. N. (N.)
atava ROMER: SWKAMA and SUZUKI,
Sci. Rept. Yokohama Nat. Univ., sec. 2,
no. 19, pI. 4, fig. 14. [=N. (Neithella)
notabilis]
1973. Neithea '(Neithea) atava (ROElVIER):
DHONDT, Mem. Inst. roy. Sci. Nat.
Belgique, no. 176, p. 40, pI. 1, figs. 2a,b [Complete synonymy and occurrences
about European specimens were given.]
1975. Neithea (Neithea) atava(ROMER) :HAY AMI, Bull. Univ. Mus. Univ. Tokyo,
no. 10, p. 74.

Itaru HAY AMI and Masayuki NODA

42

1976. Neithea kanmerai HAY AMI: OBATA ahd
OGAWA, Bull. Nat. Sci. Mus. [Tokyo],sec. C, vol. 2, no. 2, p. 101, listed, pI.
4, fig. II.

Material.-A specimen (NSM MM7348)
from the middle part of the Arida Formation at loco 1908 (see OBATA and OGAWA,
1976, p. 99), Hozudo, Yuasa-cho, Aridagun, Wakayama Prefecture (Y. OGAWA
coIl.) and ten specimens (UMUT MM
5703-5706, GK. H10291, 10292, 10294, 10295,
10297, 10298) from the lower member ofthe Haidateyama Formation at loc. Hk204
southern slope of Mt. Haidateyama,
Honjo-mura, Minamiamabe-gun, Oita Prefecture (M. NOD A coll.).
Description.-Shell very large, sometimes exceeding 100 mm in length andheight,plano-convex, subequilateral.
Right valve strongly inflated in young
stage but moderately in adult, higher
than long; antero- and postero-dorsal

margins of disc slightly concave; ventral
margin strongly digitate in accordancewith six primary ribs; apical angle about
80 degrees, gradually increased withgrowth; primary ribs invariably six innumber, strong, roof-shaped in transverse
section (see Text-fig. 5), commonly tripartite, more or less curved outwards;
1st and 6th primary ribs somewhat
weaker than other ribs; secondary ribs
variable in prominence, round-topped,commonly three on each interspace; bothlateral areas weakly striated; radialribs and inters paces marked with fine
growth-lamellae. Left valve nearly flator a little convex in adult stage, thoughmarkedly concave in early stage; anteroand postero·dorsal margins nearlystraight, forming an apical angle ofabout 100 degrees; primary ribs six innumber, bipartite, highly elevated, corresponding with the tripartite primary
Table 4.

Main diagnostic characters

speciessize(max. height)

radial ribs
(RV)

primary ribs
(RV)

primary ribs
(LV)

N. (N.) aketoensis

medium
51 mm

seemingly
irregular

not clearly
differentiated?

N. (N?) alta sp. nov.

very small
19mm

P-I-P

not clearly
differentiatedrecognized aspairs of ribs

N. (N.) atava

very large
120mm

P-3-P

roof-shaped
tripartite

highly elevatedbipartite

N. (N.) ficathoi

small
31 mm

P-2-P

round-topped
tripartite

flattenedbipartite

N. (N.) kochiensis

large
78mm

P-O-P

round-topped
simpleN. (N.) matsumotoi

small
29mm

P-4-P

round-topped
simpleelevated
bipartite

N. (N.) nipponica

medium42mm

P-4-P

not very strong
simplenot clearly
differentiated

N. (Nt.) notabilis

very small
15mm

P-O-P

strengthened
with growthstrongly
plicatedN. (N.) syriaca
amanoi

small
25mm

P-3-P

round-topped
simpleflattenedbipartite

674.

Morphology of Neithea

ribs of right valve; interspaces of primaries broadly concave, provided withweak secondary ribs which are normally
two in number; numerous concentric
growth lamellae cross the radials andinters paces, more squarrose than in rightvalve, indicating notably digitate ventral margin. Both auricles of two valvesacutely triangular, well developed; anterior auricle somewhat longer thanposterior one but distinctly narrower
with smaller apical angle; byssal notch
shallow throughout the growth; anterior
and posterior gaping, if present, very
narrow; hinge line moderately long,provided with numerous denticles; apair of strong and transversely striatedcrural teeth developed on the both sidesof resilifer, elongated along the boundary
between the disc and two auricles. Musculature unknown.
Remarks.-In Japan this giant species

43

has been called Neithea (Neithea) kanmerai, since one of us (HAY AMI, 1965)
proposed it as a new specific name.
DHONDT (1973) examined many hitherto
described specimens of Neithea from
Europe and clarified the diagnostic characters of N. atava which had been somewhat controversial among authors. At
the same time she properly pointed outthat N. kanmerai from Japan is synonymous with N. atava of original sense.
On the other hand, several well preserved
specimens were recently collected fromthe Haidateyama Formation of central
Kyushu and the Arida Formation of Kii
Peninsula.
The shell material is completely eroded
away except for an incomplete conjoined
specimen (UMUT MM5706), but the surface ornamentation of two valves arewell recognizable in other externalmoulds. The hinge structure is exhibitedof the Japanese species of Neithea
lateral areas(RV)

ventral marginauriclesconvexity(RV)

convexity(LV)

age(in Japan)

striatedpolygonal1y
angulated?

moderate?

low. Albiansmoothslightly
digitate

moderateconcave

Berriasianstriatedstrongly
digitate

sub equal

strong·moderateconcave--flator convex

Barremian
to Aptiansmoothslightly
digitatesubequalstrongweaklyconcave

up. Aptian

striated?

moderate?

Aptian orAlbianstriatedpolygonal1y
angulated

subequal

moderatenearly flat

Albianstriatedpolygonallyangulated?

moderatenearly flat

low. Albiansmoothstrongly
digitate

unequalan.>pos.

strongconvex-concave

Barremian
to Aptiansmoothroundedsub equal

moderatenearly flat

Aptian44

Itaru HAY AMI and Masayuki NODA

in the right internal mould (NSM MM
7348) and the auricles are completely
impressed on the left external mould(UMUT MM5703). The characteristic
shape of primary ribs, which are tripartite in the right valve and bipartite
in the left, is commonly seen, while thesecondary ribs of two valves seem tobe considerably variable in prominence.
In 1965, one of us proposed Neithea
kanmerai mainly on the basis of four
deformed right internal moulds andone incomplete left external moulds
from the Aptian Hinagu Formation ofcentral Kyushu. Because the originaldescription was based on such incompletematerial, the external features of rightvalve were not precisely known, and themode of radial ornament, especially the
relationship of surface relief between
two valves, was erroneously interpreted.
As the result of reexamination on the
Hinagu specimens, the primary ribs ofleft valve must be also six in number(instead of five) and distinctly bipartite.
Tripartition of primary ribs in rightvalve is hardly ascertained from theimpression of radials on the internalmoulds, but it is also indistinct in the
newly obtained internal moulds. Anyhow,
no essential difference can be found between the type specimens of N. kanmerai
and the present material.
On the other hand, as pointed out by
DHONDT (1973), the Hinagu specimens
as well as the present ones are quite
similar to the European specimens of
Neithea (Neithea) atava (ROEMER, 1839)
of proper sense in every diagnostic character. In addition to the occurrence inwestern Europe and north Africa, several
firmly identical specimens were recorded
from the Barremian or Aptian of Trinidad (Cox, 1954), Lower Cretaceous ofsoutheast Mexico (ALENCASTER, 1956)
and Hauterivian-Aptian of Crimea andCaucasus (DRUSCHIZA and KUDRJAVTSEVA, 1960). In every area N. (N.) atava
is distinguishable from other species by
the enormously large size and characteristic radial ribbing, and its geographic
variation in morphology, if present, seems
to be rather slight.
Occurrence.- The distribution of thepresent species seems to be world-wide,
and the vertical range extends from the
Berriasian to the Albian. In Japan thisspecies is also common in the Barremianand Aptian sandstones at the following
localities (see also Text-fig. 1). 1) Middle
part of the Hinagu Formation at loco Km.
3085c, south of Kobaru, Toyo-mura,
Yatsushiro-gun, Kumamoto Prefecture(type locality of N. kanmerai); 2) Middle
part of the Arida Formation at loco YS.
103, west of Kumai, and loco 1908, Hozudo,
both in Yuasa-cho, Arida-gun, Wakayama
Prefecture; 3) Lower part of the Hanoura Formation at loco Hy. 5004, north ofHiroyasu, Katsuura-cho, Katsuura-gun,
Tokushima Prefecture; 4) Ishido Formation at loco Hy. 4001, near Ichinose-bashi,
south of Kagahara, Nakazato-mura,Tano-gun, Gumma Prefecture; 5) Lower
member of the Haidateyama Formation
at loco Hk204 on the southern slope of
Mt. Haidateyama, and upper member ofthe same formation at loco Hk304 on thepath along the eastern side of a valley
from Kamikoshigoe to Mejiro-pass, bothin Honjo-mura, Minamiamabe-gun, Oita
Prefecture. At loco 1908 some Barremian ammonites were found in association (OBATA and OGAWA, 1976), andat loco Hk304 this species is accompanied by Ancyloceras (Ancyloceras)
vandenheckii (see NODA, 1977).
Neithea (Neithea) ficalhoi(CHOFFAT, 1888)

674.
5

4

3

2

R~

Morphology of Neithea
43,-~-.-....., ...."'"
~
2

-..;

6

~/

.....

R ....

..~

L ."...... ••.--_~

N. syriaca amanoi

N. notabi Iis

1
2
3
4
5
6
L~

N. ficalhoi
4 3

3 4

5

6

N. nipponica

7

6~1
R{.r
,
2

I

'.'••

123456123456

5

45

5

4

lcm

~

6

l-~

N. alta........

1

2

3

4

'H"'_'_~M_""

N. matsumotoi

N. atava

Text-fig. 5. Diagrammatic transyerse section of disk along the maximum length to show
the variability of radial ornaments among main Japanese species of Neitl!ea. R: right valve,
L: left valve, numerals 1 to 6: number of primary ribs from the anterior side, solid line:
outer surface, broken line: supposed internal surface.

Synoymy.-See HAY AMI (1965, p. 302;
1975, p. 75).
Remarks.-This species was originallydescribed from the (?) Albian of Angola.
In Japan specimens referable to thisspecies occur commonly from the Tanohata and Hiraiga Formations (upper
Aptian-lowest Albian) of the Miyako
Group. A specimen from a boulder ofsandstone at the upper stream of Onkonosawa in Yam abe area of central
Hokkaido (MATSUMOTO and OKADA, 1973,
p. 293) also certainly belongs to thesame species. This is easily distinguishable from other Japanese species of
Neithea by the combination of the following characters: relatively small size,
subequal auricles, feebly but regularly
tripartite primary ribs and invariably
two secondaries on each interspace inthe right valve, strong convexity of rightvalve and broadly concave left valve.
In these characters the Japanese specimens are probably identical with theoriginal specimens of Janira ficalhoi from
Angola (CHOFFAT and LORIOL, 1888).
As noted before (HAY AMI, 1965), thepresent species may be intimate to
Neithea alpina (D'ORBIGNY, 1847) fromthe Upper Cretaceous of western Europe.
DHONDT (1973) synonymized J. ficalhoi
with N. alpina together with Pecten
tricostatus BAYLE in FOURNEL, 1849, andJanira doumerci PERON, 1877. The present Japanese specimens are, however,not coincident with DHONDT'S descriptionof N. alpina in some characters. On thebasis of. the lectotype and other Upper
Cretaceous specimens of N. alpina, she
noted, "The right valves are not very

46

Itaru HAY AMI and Masayuki NODA

convex, particularly so on large specimens, and left valves are flattened: I
have not come across concave left valves.
.. On convex valves the [lateral] areas
are slightly curved outwardly and arecovered by a variable number of ribOn the contrary, the Miyako
lets."
specimens in question are characterized
by the comparatively strongly inflated
right valve and distinctly concave left
valve, though they are generally smaller
than European specimens of N. alpina.
One or two radial undulations are commonly seen on the lateral areas of theright valves in the Miyako specimens,
but they are too inconspicuous and weak
to be called riblets. Moreover, N. alpina
survived in Europe from the Albian to
the Maastrichtian (DHONDT, 1973) andhas generally been recognized as an
Upper Cretaceous species, while thestratigraphic occurrence of N. ficalhoi
in Japan is confined to the upper Aptian
and lowest Albian. Several species of
Neithea from the Lower Cretaceous
Comanche Series in the Gulf Coast region
are also characterized by two secondary
ribs on each interspace in right valve.
Among others Neithea occidentalis (CONRAD, 1855) [=Neithea irregularis (BOSE,1910)J from the Albian seems to sharemany common characters with the present specimens. Further comparative
studies between the two nominal species
are desirable. At present N. ficalhoi isregarded as specifically distinct from N.
alpina and N. occidentalis, but the threenominal species are certainly closely
related to one another and seem to constitute one and the same group of common phylogenetic origin.
Neithea (Neithea) kochiensis HAYAMIin HAY AMI and KA w ASA W A, 1967
Synonymy and type.-See HAY AMI, 1975,

p.76.
Remarks.-This is an imperfectly
known species founded on a solitaryinternal mould of right valve from theAlbian (if not Aptian) Doganaro Formation of the Shimantogawa Group in south
Shikoku.
Though nothing has been
known about the left valve, it seems to
be distinct from any other species of
Neithea from Japan by the relatively
large size, round-topped and comparatively narrow primary ribs, wide andflattened interspaces, weak and undeveloped secondary ribs and angularly
pointed umbonal area. The externalfeature is precisely unknown, but theinternal mould lacks any impressions of
seco'ndary ribs even near the ventral
margin but for numerous faint striae.
In the large shell size only Neithea(Neithea) atava is comparable with thepresent species, but the apical angle islarger, the primary ribs are more roofshaped in transverse section and thesecondary ribs are more developed in
the former. The holotype of N. (N.)
kochiensis occurred together with a specimen of Neithea (Neithea) matsumotoi,
but the morphological difference between
the two species is quite obvious. Among
various European species this may besomewhat similar to N eithea (Neithea)
striatocostatus (GOLDFUSS. 1833) (DHONDT,1973) from the Cenomanian-Maastrichtian, but the umbonal area is narrower
than in that species. The present species
was originally referred to subgenus
Neithella HAY AMI, 1965, with a query.
As renamed already (HAY AMI, 1975),
however, it seems to belong to Neithea(s. s.), since the weakness of secondary
ribs do not necessarily characterize thatsubgenus.
Neithea (Neithea) matsumotoi
HA YAMI, 1965

674.

Morphology of Neithea

Synonymy and type.-See HAY AMI, 1975,
p.76.
Remarks.-This species was originallydescribed from the Albian Yatsushiro
Formation of central Kyushu, and also
known from the Haidateyama Formationof central Kyushu (HAY AMI, 1965) andthe Doganaro Formation in south Shikoku(RA YAMI and KA w ASAWA, 1967). A left
valve of Neithea sp. from the core oftest boring at Peikan of west Taiwan(MATSUMOTO, HAYAMI and HASHIMOTO,
1965) probably belongs to the same
species. It is characterized by the small
size, small and subequal auricles, strong
and well discriminated primary ribs inthe right valve, commonly four weaksecondary ribs on each interspace andweakly striated lateral areas of the rightvalve, round-topped radial ribs on the
flattened left valve, of which six pairs
are notably heavier than other ribs, andrelatively prominent concentric growthlamellae on the left valve. As remarked
by HAY AMI (1965) and also DHONDT(1973), the present species actually resembles Neithea sexcostata (WOODWARD,
1833) in many essential characters. Yet,the resemblance is rather curious, as thegeologic ages of the two species arequite different. In various areas ofwestern Europe N. sexcostata commonlyoccurs in the Chalk and ranges from the
Cenomanian to the Maastrichtian, whereas in Japan and Taiwan N. matsumotoi
is known only from the Albian (? andAptian) non-calcareous deposits. As theresult of our comparative study with thespecimens of N. sexcostata from the
Chalk of England and Scania, it wasproved that the apical angle of rightvalve is a little larger, the convexity ofright valve is slightly weaker and thesix paired ribs of the left valve are distinctly heavier in the Japanese specimens.
N. matsumotoi is thus regarded as dis-47

tinct from N. sexcostata, but it remains
unsolved whether the morphologicalresemblance suggests some phylogenetic
relation between the two species or isonly due to convergence. The morphological similarity between N. matsumotoi and Neithea attockensis Cox, 1935,
from the Albian of India may offer
another taxonomic problem, but any
conclusion is now difficult, since thatspecies is represented by an poorly
preserved specimen.
Neithea (Neithea) nipponica
HAYAMI, 1965
Synonymy and type.-See HAY AMI, 1975,
p.76.
Remarks.-This species commonly occurs in the uppermost part of the Hiraiga
Formation and the Aketo Formation of
the Miyako Group. The age is certainly
lower Albian as indicated by associatedammonites. This appears to have suddenly substituted the pre-existing species,
N. (N.) jicalhoi, but the two species are
rarely but strictly associated with eachother at a few localities. There are no
individual showing intermediate morphology, and no direct phyletic relationship can be expected between the two
species. Rather it seems more reasonable
to consider that N. nipponica migrated
from somewhere to this area and usurped
within a short time the niche which had
been occupied by N. jicalhoi. N. nipponica
clearly differs from other Japanese
species of Neithea in the combination of
the following characters: medium size,
rather weak contrast of prominence
between the primary and secondaryribs, almost invariably four secondary
ribs on each interspace in the rightvalve, indistinctness of six pairs of ribsin the left valve, and fine radial ribletson the lateral areas of the right valve.

48

Itaru HAY AMI and Masayuki NODA

In many of these characters N. nippo.nica
resembles some specimens of Neithea
quinqllecostata (SOWERBY, 1814). DHONDT(1973) regarded the morphological differences between the two species statedby HAY AMI (1965) as insignificant andinterpreted N. nipponica as a juniorsynonym of N. quinquecostata. As theresult of further comparative studies
with the specimens of N. quinquecostata
preserved at the British Museum (Nat.
Hist.) and some other institutions inwestern Europe, however, one of us (1.
H.) maintains the taxonomic distinction
between the two species. In comparison
with the Upper Greensand and Chalkspecimens of N. quinquecostata including
the lectotype (BMNH no. 43324), theJapanese specimens have much narrower
umbonal area, more slender primaryribs and more rounded anterior andposterior extremities of right valve, lesspolygonally angulated ventral margin of
two valves, smaller size and probably
thinner test. These difference cannot
be attributed to the difference of ontogenetical stages between the materials,
though once assumed by DHONDT. At
the same time, however, it must be notedthat the morphological differences are
not so significant between the Japanese
specimens of N. nipponica and those of"N. quinquecostata" from the Lower
Greensand and Gault of England. Our
specimens are likewise similar to theillustrated specimens of N. qllinquecostatafrom the upper Albian of Crimea (DRUSCHIZA and KUDRJAVTSEVA, 1960), which
also show more rounded ventral margin
and weaker contrast between the radialribs of two orders. It is thus suggested
that somewhat significant morphologicalchange occurred in the stock of N.
quinqllecostata, and that the name of N.
(N.) nipponica is possibly applicable also
for the fossil populations from the Lower

Cretaceous of Europe. We think thenomenclatural translation of N. (N.)
nipponica to subspecies is not necessary,even if it proves to be phyletically related to N. quinqllecostata.

Neithea (Neithella) notabilis(MUNSTER in GOLDFUSS, 1833)

Synonymy.-See HAY AMI. 1975, p. 77.
Remarks.-In Japan this species occurs,though rarely, in the upper Neocomian
Ishido Formation of Kwanto mountains(Y ABE, NAGAO and SHIMIZU, 1926), theupper Aptian part of the Miyako Groupin north Honshu (HAY AMI, 1965) and theAptian (?) part of the Choshi Groupof east K wanto (SHIKAMA and SUZUKI,
1972). A few specimens from these areas
had been called Neithea (Neithella) d.
atava (ROMER), until in opposition to
WOODS' (1903) view DHONDT (1973) properly pointed out that the name of
Neithea (Neithea) atava should be applied for an unrelated species of muchlarger size. In comparison with many
European specimens referred by her to
Neithea (Neithella) notabilis, every essential character of two valves is infact so similar that no taxonomic distinction seems necessary for these Japa~
nese specimens. This species has small
shells, angular to round-topped primaryribs which become much broader throughthe individual growth, finely striated interspace without any ribs of secondary
order in the right valve, strong convexityof right valve and angularly plicated andstrongly digitate left valve.
As to the subgeneric diagnosis of thesubgenus Neithella, DHONDT (1973, p. 8)
wrote, "Neithella has been created forspecies with 4-6 asymmetrical ribs andhighly unequal auricles. The ribs of theleft valve are the reflection, and not the674.

Morphology of Neithea

negative impression, of those of the rightvalve". In fact, this subgenus is wellcharacterized by the very unequal "auricles: the anterior auricles is pointed
forward and much larger than the posterior, though in most Japanese specimens it is imperfectly preserved. In
some species of Neithella, as originally
noted, the 1st and 6th primary ribs aremuch weaker than others or so inconspicuous that they appear to have only
four or five ribs. Such a tendency is
scarcely known in Neithea (s. s.). We
also feel the necessity of some additional
explanation about the relation of radial
relief between two valves. In Neithea
and a few other Mesozoic genera of thePectinidae, unlike ordinary pectinids, theprimary ribs of right valve correspondwith the radial elevations of left valve
to give digitations to the ventral margin,
whereas the secondary ribs commonlyinterlock with the grooves of the countervalve. Such a feature is clearly observable in many digitate species of
Neithea (s. s.), and in this point Neithella
does not differ from Neithea (s. s.).
Neithea (Neithea) syriaca amanoi

HAY AMI, 1965
1957. Pecten (Neithea) cf. morrisi (PICTETand RENEVIER): AMANO, Kumamoto
Jour. Sci., ser. B, sec. 1, vol. 2, no. 2,
p. 88, pI. 1, figs. 17, 18, 20-25, 27-29.
1965. Neithea (Neithea) amanoi HAYAMI,
Mem. Pac. Sci. Kyushu Univ., ser. D,
vol. 15, no. 2, p. 299, text·fig. 4, pI. 41,
figs. 8-10, pI. 42, figs. 1-4.
1972. Neithea (Neithea) amanoi HAY AMI:
SHIKAMAand SUZUKI, Sci. Rept. Yoko·hama Nat. Univ., ser. 2, no. 19, pI. 4,
fig. 15.
1973. Neithea (Neithea) syriaca (CONRAD):DHONDT, Mem.
Inst. roy. Sci. nat.
Belgique, no. 176, p. 37 (pars).
1975. Neithea (Neithea) syriaca (CONRAD):49

Bull. Univ. Mus. UnitJ. Tokyo,
no. 10, p. 75.

HAYAMI,

Type.-Holotype of N. (N.) amanoi (GK.
H6267) from the Aptian Hagino Formation at Hagino, Kahoku·cho, Kochi Prefecture.
Remarks.-This subspecies is known
only from the type locality and the
Choshi Group of east Kwanto. It ischaracterized by the small size, almost
invariably three secondary ribs on eachinterspace in the right valve, fine radialribs of nearly one order of prominencein the left valve, almost smooth lateralareas, not strongly inflated right valve
and nearly flat left valve without marginal digitations. DHONDT (1973) regarded Pecten (Neithea) ct. morrisi by
AMANO (1957), which was described on
the basis of the same fossil population,
as belonging to Neithea (Neithea) syriaca(CONRAD, 1852). In fact, janira mOr1'isi
PICTET and RENE VIER, 1858, appears tobe synonymous with janira syriaca
CONRAD, 1852. According to VOKES(1947), the original specimen of j. syriaca
from Abeih of Lebanon was lost, but its
specific characters can be recognized insome "neosyntypes" from the type locality. The Japanese specimens actuallyresemble the topotypes of N. (N.) syriaca
and also some European specimens hitherto described under the name of N.
morrisi. The small size, relatively narrow
and rounded outline and smooth lateralareas of right valve are particularly
similar, while the radial ornaments oftwo valves are somewhat different. In
the Lebanon specimens there are commonly four secondary ribs on each interspace in the right valve and 26 to 28ribs in the left, whereas the Japanese
specimens usually possess three secondary ribs in the right valve and totally
22 to 24 ribs in the left. The six pairs

50

Itaru HAY AMI and Masayuki NODA

of ribs in the left valve, which correspond to the primaries of the right valve,are probably more prominently elevatedin the japanese specimens. Although
one of us (HA YAMI, 1975) once regarded
N. amanoi as a synonym of N. syriaca,
the above mentioned slight but clear
difference seems to deserve certain
taxonomic distinction. At present it isdifficult to say with documentation whether the difference represents the distinctness of species or is merely due togeographic variation, but the japanese
specimens are here tentatively treatedas a subspecies, considering the japaneseand Lebanon populations are almost contemporaneous. Most of European specimens of "N. morrisi" are more similar
to the Lebanon specimens than to japa.
nese ones, and they may be generally
referable to the nominate subspecies, N.
(N.) syriaca syriaca.
Various diagnostic characters of these
Japanese species and subspecies of
Neithea are coIlectively shown in Table
4. The taxonomic names here adopted
may be considerably different from
DHONDT'S (1973), but in many cases weagree with her in recognizing the morphological resemblance between European
and japanese specimens. The mode ofradial ribbing not only in the right valve
but also in the left seems to be important for the taxonomic distinction, andthe transverse sections of two valves inmain japanese species are diagrammatically shown in Text-fig. 5.
Summary1) The mode of radial ribbing in
Neithea is, because of its peculiarity andregularity, regarded as an interesting
character not only for the taxonomic
criterion but also for the general con-sideration about the differentiation ofradial ribs into two orders.
Manyspecies with strongly digitate ventral
margin show marked contrast of prominence between the primary and secondary ribs in the right valve and have
six pairs of contiguous ribs (or six
bipartite ribs) in the left valve. The
total number of radial ribs is basically
expressed as: 6+5n in the right valve.
and 12+5 (n-1) in the left valve (nvaries among species).
2) It is worthy of notice that thepresence of aragonitic inner layer isindirectly but strongly suggested in theoriginal shells of Neithea and Weyla.
Though any hasty conclusion on theirtaxonomic significance is avoided here,
it may bear something informative forthe consideration of shell structure evolu.
tion from the A viculopectinidae to thePectinidae as well as the relationshipbetween the Pectinidae and the Spondylidae.
3) Analogy with Pecten in the modern
seas leads us to suspect that the planoconvex shells of Neithea and HT eyla represent a kind of gryphaeate morphology
which may be advantageous in preventing the commissure from sinking into
soupy substrate, even though they were
not so excellent swimmers as Pecten.
Their nearly flat or even concave left
valve with frequently more squarrose
surface are probably quite effective tohold a layer of sediments in order toconceal themselves from predators.
4) All the species of Neithea, which
had been known from the Lower Cretaceous of japan, were taxonomically
restudied, and their specific names andaffinity were discussed in some detail on
the basis of a comparative study with
European specimens. As pointed out by
DHONDT (1973), there are some commonor closely related species between japan

674.

Morphology of Neithea

and Europe, though the actual geographic distribution and migration route ofeach species are difficult to say owing
to the deficiency of fossil records in the
midway areas.
5) Neithea kanmerai HAYAMI, 1965,
should be regarded as a synonym of
Neithea (Neithea) atava (ROEMER, 1839),
which actually shows very wide distribution. Detailed surface features of twovalves were known in the newly collectedmaterials from southwest Japan.
6) Neithea (Neithea?) alta sp. nov.
from the Lowest Cretaceous of north
Honshu is probably one of the earliest
species of this genus and appears toindicate an early stage of differentiationof radial ribs into two orders.

Acknow ledgments
It is a great pleasure to express our
sincere gratitude to Professor R. A.
REYMENT (Uppsala University) who
enables one of us (1. H.) to visit many
institutions of western Europe and tocompare Japanese specimens of Neithea
with European ones and to Professor
Tetsuro HANAI (University of Tokyo)
for his encouragements and reading of
this manuscript. We are much obliged
to Dr. Fuminori TAKIZAWA (Geological
Survey of Japan) and Mr. Yoshio OGAWA
who kindly offered their collections of
Neithea described in this article. Acknowledgments are also due to Professor
Tatsuro MATSUMOTO (Kyushu University), Drs. N. J. MORRIS and C. P. NUTTALL (British Museum of Natural History), Dr. Anthony HALLAM (Oxford
University), Dr. David PRICE (Sedgwick
Museum of Cambridge) and Professor G.
REGNELL (Lund University) for theirpermission to observe the specimens
preserved in respective institutions, andalso to Dr. Wolf-Ernst REIF (Tiibingen

51

University), Dr. A. V. DHONDT (RoyalInstitute of Natural Science, Belgique),Dr. W. J. KENNEDY (Oxford University)and Dr. Ikuwo OSATA (National Science
Museum, Tokyo) for their useful suggestions.

References
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52

Itaru HAYAMI and Masayuki NOD A

pp. Geol. Soc. America and Univ. Kansas.
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- - (1965): Lower Cretaceous marine pelecypods of Japan. Part I. Mem. Fac. Sci.
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MATSUMOTO, Tatsuro' (1954): The Cretaceous
system in the Japanese islands. Japan.

Explanation of Plate 5
Figs. 1-3. Neithea (Neithea?) alta HAYAMI, sp. nov.
Fig.!. Internal mould of a right valve, (UMUT MM5698), holotype, from the Kobitawatashi Member of the Ayukawa Formation at Yokone, Ajishima Island, Ojika-cho, Ojikagun, Miyagi Pref. (TAKIZAWA coll.) x2.
Fig. 2. Rubber cast of a fragmentary left external mould, (UMUT MM5700), paratype,from the same locality. (TAKIZAWA coil.) x2.
Fig. 3. Rubber cast of a fragmentary left external mould, (UMUT MM5699), paratype,from the same locality. (TAKIZAWA coIl.) x2.
Figs. 4, 5. Neithea (Neithea) atava (ROEMER)
Fig. 4a. Right internal mould (UMUT MM5703) from the lower part of the Haidateyama
Formation at loc. HK204 on the southern slope of Mt. Haidateyama, Honjo-mura, Minamiamabe-gun, Oita Pref. (NODA coil.) X 1. The impression of both wings is on the external mould of the counter valve.
Fig. 4b. Rubber cast of the left external mould of the same individual. xl.
Fig. 5. Plaster cast of a left external mould (GK. HI0297) from the same locality.
(NODA coil.) x 0.9.

HAYAM J and NODA:

M01'phology of Neithea

Plate 5Morphology of Neithea
Soc. Prom. Sci. Res., Tokyo. xiv +324
pp., 36 pis. [for 1953J- - , HAYAMI, Itaru and HASHIMOTO, Wataru(1965): Some molluscan fossils from theburied Cretaceous of west Taiwan. Petrol.
Ceol. Taiwan, vol. 4, p. 1-24 (2 pis. incl.).
- - and KANMERA, Kametoshi (1952) : Guide
book for geologic excursions: the lower
valley of the Kuma. Dept. Geol. Kyushu
Univ. 71 pp. 6 maps. [in JapaneseJ- - and - - (1964): Explanatory text ofthe geological map of Japan: Hinagu.
Geol. Surv. Japan. 147+27 pp. [in Japanese with English abstractJ- - and OKADA, Hakuyu (1973) : Saku formation of the Yezo geosyncline. Sci. Rept.
Dept. Ceol. Kyushu Univ., vol. 11, no. 2,
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abstractJ
NAGAO, Takumi (1934): Cretaceous Mollusca
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NEWELL, N. D. (1937-1938): Late Paleozoic
pelecypods: Pectinacea. Univ. Kansas,
State Ceol. Surv. Kansas, Publ., vol. 10,
p. 1-123 [1937J, pis. 1-20 [1938].
NODA, Masayuki (1977): A brief note on
Ancyloceras from the Haidateyama formation, Kyushu.
Trans. Proc. Pal. Soc.
Japan, n. s., no. 104, p. 418-423, pI. 44,
text-figs. 1-3.
OBATA, Ikuwo and OGAWA, Yoshio (1976):
Ammonites biostratigraphy of the Cretaceous Arida formation, Wakayama
Prefecture. Bull. Nat. Sci. Mus. [TokyoJ,ser. C, vol. 2, no. 2, p. 93-110, pis. 1-4.
D'ORBIGNY, Alcide (1843-1847) : Pateontologie
franr;aise. Terrain cretaces. III. LamelIibranches. Paris. 807 pp., pis. 237-489.
ROEMER, F. A. (1839): Die Versteinerungendes nord-deutschen Oolithengebirges.
Nachtrag. 47 pp., pis. 17-20.
SHIKAMA, Tokio and SUZUKI, Shigeki (1972) :Stratigraphy and tectonic development
mainly of Cretaceous formations of Choshi peninsula, Chiba Prefecture.
Sci.
Rept. Yokohama Nat. Univ., ser. 2, no.
19, p. 133-157, pis. 4-6. [in Japanesewith English abstractJ

53

STANLEY, S. M. (1970): Relation of shellform to life habits in the Bivalvia(Mollusca). Mem. Ceol. Soc. America,
125, p. 1-296 (pis. 1-40 incl.).
STANTON, T. W. (1947): Studies of someComanche pelecypods and gastropods.
U. S. Ceol. Surv., Prof. Paper, 211, p. 1256 (pis. 1-67 incl.).
TAKIZAWA, Fuminori (1970):
Ayukawa
formation of the Ojika Peninsula, Miyagi
Prefecture, northeast Japan. Bull. Ceol.
Surv. Japan, vol. 21, no. 10, p. 567-578,
pI. 1.
- - (1975): Lower Cretaceous sedimentationin the Oshika Peninsula, Miyagi Prefecture, northeast Japan.
Bull. -Ceol.
Surv. Japan, vol. 26, no. 6, p. 267-305,
pis. 10-16. [in Japanese with English
abstractJ- - , ISSI-IIKI, Naoki and KATADA, Masato(1974): Geology of the Kinkasan district.
Quadrangle series, Geol. Surv. Japan.
62+5 pp., 1 map. [in Japanese withEnglish abstractJ
TAMURA, Minoru (1959): Taxodonta andIsodonta from the Upper Jurassic Sakamoto formation in central Kyushu. Trans.
Proc. Pal. Soc. Japan, n. s., no. 34, p. 5365, pI. 6.
TAYLOR, J. D., KENNEDY, W. J. and HALL,
A. (1969): The shell structure andmineralogy of the Bivalvia. Introduction,
Nuculacea-Trigonacea. Bull. Brit. Mus.
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Ceol. Surv. Japan, no. 237, p. 1-84. [inJapanese with English abstract]
THAYER, C. W. (1970): Adaptive significanceof oblique adductor muscles in swimmingmonomyarian bivalves. Ceol. Soc. America,
Abstracts with programs, vol. 2, no. 7, p.
703.
VOKES, H. E. (1947): Contributions to thepaleontology of the Lebanon mountains,
Republic of Lebanon. Part 3. The pelecypod fauna of the "Olive locality"(Aptian) at Abeih. BuU. Amer. Mus.

54

Ita1'u HAY AMI and Masayuki NODA

Nat. Hist., vol. 87, p. 141-215, pis. 1-9.
WOODS, Henry (1902-1903): A monographof the Cretaceous Lamellibranchia of
England. Pectinidae. Palaeontogr. Soc.
London. p. 145-196, pis. 27-38 [1902J,
p. 197-232, pis. 39-52 [1903J.

YABE, Hisakatsu, NAGAO, Takumi and SHIMIZU, Saburo (1926):
Cretaceous Mollusca from the Sanchu graben in the
Kwanto mountainland, Japan. Sci. Rept.
Tohoku Imp. Univ., ser. 2, vol. 9, no. 2,
p. 33-76, pis. 12-15.

Ajishima m<J:tt!!.,Ili"
Ayukawa [MJII,
Choshi ~~T,
Hagino fJC!]lf,
Haidateyama -Mtmw,
Hiroyasu rt'J(,
Honjo *8:,
Hozudo i'!/!j!:p, Kagahara 1$ 'T 1*, Kamikoshigoe J:~MlS,
Katsuura ~im,
Kobaru 'J'I*,
Kobitawatashi 'H~:I!!t,Kumai fl~*,Mejiro f:I S,
Miyako 'gil,
Nakazato ,*,ll,Ojika !l±Jf[,
Sanchu W,*"
Susaki ~r.t,
Tanohata
EB'!!HIB,
Toyo *~,
Yamabe Wfftl,
Yatsushiro /\fI;';, Yokone t/!{i!'!,Yuasa ~i~.

Explanation of Plate 6
Figs. 1, 2. Neithea (Neithea) atava (ROE:VIER)
Fig. 1. Plaster cast of a right external mould (GK. H5002) from the lower member ofthe Haidateyama Formation at loc. Hk204 on the southern slope of Mt. Haidateyama,
Honjo·mura, Minamiamabe-gun, Oita Pref. (NODA coil.) xL
Fig. 2a. Internal mould of a right valve (NSM MM7348) from the middle prt of theArida Formation at loc. 1908, Hozudo, Yuasa-cho, Arida-gun, Wakayama Pref. (OGAWA
coil.) xl.
Fig. 2b. Rubber aast of the right external mould of the same individual. xL

H AYAM 1 and NODA:

Morphology of Neithea
Plate 6

55

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56
Some heteromorph ammonites from the Upper Cretaceous of Hokkaido············T. MATSUMOTOC P N S ................................................................................. ·································ii1Ij;(!iU if"'i5
Mid·Cretaceous Events ~ti1&~~~ ...................................................................,....... ~*:ita"

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Limestone, Chichi-Jima, Ogasawara Islands, Japan ....................................... K. MATSUMARL"On two new species of Nephrolepidina and Miogypsina from the Chichibu Basin,
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Mesozoic plap.ts from the Sebayashi Formation (Aptian), Kwanto Mountainland,
Gumma Prefecture, Outer zone of Japan •·· .... · .... ·· ............ ·T. KIMURA and M. MATSUKAWA
Addition to the Mesozoic plants from the Akaiwa Formation (Upper Neocomian),
Itoshiro Group, Central Honshu, Japan ······································· .. ·T. KIMURA and S. SEKIDO
Some conifer remains from the Eocene beds of Kuji, Iwate Prefecture, Japan················ .. ········ .. ··· ...... ·····T. KIMURA and J. HORIUCHI
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Palynology of the Lower Tertiary Concepcion Formation, Central Chile ............ K. TAKAHASHIGGIEf'lill#JlItJ'i?i£:f:I:lL..t.:-f / -i!:'7AAt::.,,?\,'-C .......................................................... "!fEB fftz
Note on some Cenomanian mollusks from Diego-Suarez, Madagascar ················ .. ······Y. KANIE
13 *i£:S9I!*,c Neithea ~;tM c i!i1:.Jrn ........................................................................... 3m7lc;f§

57
A new fossil Chlamys from the Miocene Otsutsumi Formation, Miyagi Prefecture···· .. ··········································K. MASUDA and Y. SATO
Colour pattern in fossil Umboniinae and its paleontologic significance .·········· ... ·······T. OZAWA:9311K~·/lI;J::$~~rr.CI)~.FI.ft:OCl):fll!r..'1*l1* ....................................................................+f.ii~;;I(

Nautilus pompilius CI)~m1~JH;::":)~'-C (T~) ...................................................... ······3jZ!Ilf~iR~P lifIity "- '7 *U '/ ~:r1

I- t;:: ":)~ '-c·············································3jZ!Ilf~iR· ::::J::.~ • 'gJlI31HM(T¥Il) .......................................... ~jJJ$ -nit

if:tEE1\\l~J::~t.:7 '/~:r1 I-mtCl)nIt:lltCl)j;!HIJjttc~~tt

Further notes on the Vascoceratids from Hokkaido, part 2······T. MATSUMOTO and M. NIl-lONG I
XK>Jl<"1!l p 7:r71-tf-~m\"t.:.FI.mt9='CI) Ca, P, Na Cl)f-IjjjtECI)~w································+tl!!
Manchuroceras in South Korea with notes on the Manchuroceratidae andManchuroceras province ...................................................... ···························T. KO!3AY ASHI
Upper Ordovician trilobites from the Langkawi Islands, Malaya·······································T. KOBAYASHI and T. HAMADA

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Quaternary Ostracoda from Kisarazu near Tokyo .... · ........ ······································M. YAJIMA................................................................................................ :o~ I;!!!I1!l............................................................................................. 1lI P ¥liz
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T~m~~IHJCI)*r$lifIitCl)iE~'7 =-i;::J: ~1tllHt:Oi;::":)\,'-C ··········································mm 71~
B*CI)9='~~WJ: f)iti'"~:r<"::ICI)1tJtft:o .................................... ,. ............................ ~~ ~~B*CI)V/t.-/t.-. 7m,/· :OlJJf*CI)::I / r''/ I-CI)~IifIFf ................................................... ~~~lj!jH:til'
B*CI)=~*CI)::I / F' '/ I- CI)~~rf ...................................................... ·····················~~~~i\~H:til,:=:~*J::t.t:Ow(:!iiIiflCl)::I / F'/ I- ~IifIFf ··········································l&i!1m~· lIJ*N4l.fIf,\ • 9='B1;ii5'B*CI)::::-I:*CI)::I / F' '/ I- CI)~~Ff ...................................................... ·····················,HI!l.~;;I(I:til,Jl[v:r#ii:CI)J3,~!/:i,0-;;(P~!l!IIMi;::~ I?n~tti'lill~

1916~1la *i!i~$J~~~::st1t~1t*~P.l::st::kt.tlJ.flt!!(t'Et: Thorium and protactinium isotopes in some present·day hermatypic corals andtheir implications to dating.
tables, 9 text.figs., 1976.

Trans. Proc. Palaeont. Soc. Japan, N. S. No. 101, pp. 271-290, 7*IDfUJ'Ot, Iml;!!!· +~W- • iWgIj~B1; • #~:N J:"(f =- "- -"/ ~ 7" '/it1 V-if" '/ ::I'CI) 1t * Y9C1) U· Th • Pa
iWJ(ftf;$: (238U, 234Th. 230Th, 231Pa) Cl)3jZ~~~:I:, iWgIj~/lI;it ~ 1-'1) 1 VCI) 238U Cl)1t*Y9~jj;:I:, t.c.l?"{ft;::iWJ!t/!J:\ilGCI)rii!¥*9='CI) 230Th/ 232 Th .Ita-, 1iVfECI)0-fJTi*i;::~=falt:N ~1Jo;t ~ -:. c i;:: J: "'? -C~;W!i;::0-;j:1T L,.f:-CI)*5:W:, ~ <c t iWgIj~/lI;it1 V-if"'/ ::I··CI)~;l!!:ft:O~*4t;::ML~m L?~, 230Th nIt:llt~~iJ!Jj~i*CI), :m~ 230Th i;::I*li'"~KiE~i'I/:r~iE~~1¥~I±lU.:tCl)-c·~~o-:. CI)~, m~1t*~*4C1)jf-i'" 230Th/ 232Th JUt, m,~J~~71(C1).f:-nJ: f) ~=ff.ii <I±l ~ -:. c ~J!1±l L,-:. CI)

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Cl)J:t!1JOi;:: J: ~ t CI) c;.lf;t t.: -:. cit, *IDfUJtCl)f.i5IDfU9=' t "'? c Ul!gJjB{)t.c.,~-c·~ ~ 0

t.c.:N, -:. CI) J: ? t;::ttIliE~nt.: 230Th ~<j1lt, iWJ-~*4C1) HC ttIliE~<j1 (t.: c ;tli SUESS, 1969) t;::)!i--j~, ttIliE~CI)*~tt~ffiRi'"~tCl)c5hn~o*IiJf~lt, -:. CI) J: ? t;::~Im*c~Wl-if" '/ ::I·~!i\CI)~~illJj~i;::~3l'JJt.c. 230Th nIt:llti*~, m-l!!:~*4i;::~mi'" ~~i;::r,,'~,~CI)-":) c t.c. ~~*4Y9C1):m~ 230Th :I:~;::~ L -C, ~';.If~~1Jo;t t.: t CI)-c', -:. Cl)nIt:W:ltiWJ(ftf;$:~u~~~CI)~J:\ilGi;::Mt~i±I3i'"~~A ifii:c~t.c.~n~oB*ti~~~:f:l:lt,

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58'N~,'Etlt 1955 if::iJ>1? 1966 if::v;::Mt-C±c L -C ~!lMc{&JtIl (J)7 :/-1:71 r (J)ilit¥J.J~A'~liif~~mrrL,~I;:: j"A-1:-e '7 AN· ~~ 4'- "- 1) 7" AN· ::z 1) 11 / =- -e '7 AHt.e c'(J) *~7H&~J'Bliif~v;:: ~..., -c, ftl!(J)liif~~ c :1J~il--1t-C*:m 7 :/ -1: 7 1 r liif~(J) ~~ 1/,,/v.A.,(J)5 I e.- J:H'v:.j1t ~ Lt.:o i t.:1itWv;::7\:;iJ t ft-C \" t;,!f.
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liif9'E~I;:: ta:1J Lt::.,*_(J)mlOXI;:: J: I) ~ I?:h QiJt, $-C' t 1970 if::U{&iJ, 1?*:m(J) ~.!!H*c'ffi-tlllJJ¥J.J1tE (J):l1!!'i'{
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0~~~lE'Et: ~.!!H*c:to J: a*'Titf\';(J)l¥iii1::fffL$.(J)liif9'El¥iii1::fffL$.(J)1tE~(ft~lt 1940 if::f\';v;::OO:(tL,*, 1950 if::f\';(J){&¥I;:::to1, '-C:7J 1) 7'ifiiiBW:-C'lt,~:hQ I;::~..., t::.o :::. (J) J: ? t.e1itWJi~MiPJ(J)r-C',:l1!!JgXH91:::.aJlH I?:h -Cl, ,t::.iJt;g~v;::~WiJt1Jnx.I?:ht::.*6
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B*'i!l~!IWJ~~~5!Ij~O)w.m~.

PALAEONTOLOGICAL SOCIETY OF JAPAN, SPECIAL PAPERS, NUMBER 23 ~ 1978 ~!llC-'t(J)~tJi~0~ 1... 'i -t 0 )j@~ t.e.,*~'i:i~ ;};f;fb (J)j]ft, iX(J)*JJH:::,i'I:b-tt-C $)?:,~~fF~ 1...,

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

April 30, 1977

New Series No. 105

CONTENTS
TRANSACTIONS672.

KOBAYASHI, Fumio: Some considerations on the ancestor of the family
Verbeekinidae (Fusulinacea) .. ........ . . .. .. . ... ....... . . .. ... . . . .....

673. KOBA YASHI, Teiichi:

1

Manchuroceras found in South Korea with notes on

the Manchuroceratidae and the Manc huroceras province.. . . . . . . . . . . ..
674. HAYAMI, Itaru and NODA, Masayuki:

17

Notes on the morphology of Nei thea

(Cretaceous pectinids) with taxonomic revision of Japanese species.. ..

p

PROCEEDINGS . . .. .. ..... . ..... . . . . . . . . ........ .. .. . .. . .. ... . .... . ...... . . . . ..

55