Transactions and Proceedingsof thePalaeontological Society of Japan
New Series

No. 84

Palaeontological Society of Japan
December 30, 1971

Editor: Takashi HAMADA
Associate editor: Yasuhide IwASAKI

Officers for 1971-1972
President : Tokio SHIKAMA
Councillors {* Executives) : Kiyoshi ASANO*, Kiyotaka CHINZEI*, Takashi HAMADA*,
Tetsuro HANAI*, Kotora HATAI, Itaru HAYAMI, Koichiro ICHIKAWA, Taro
KANAYA, Kametoshi KANMERA, Tamio KOTAKA, Tatsuro MATSUMOTO*, Hiroshi
OZAKI*, Tokio SHIKAMA*, Fuyuji TAKA!*, Yokichi TAKAYANAGISecretaries: Wataru HASHIMOTO, Saburo KANNOExecutive Committee
General Affairs: Tetsuro HAN AI, Naoaki AOKIMembership : Kiyotaka CHINZEI, Toshio KOIKE
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Publications
Transactions:Takashi HAMADA, Yasuhide IwASAKISpecial Papers: Tatsuro MATSUMOTO, Tomowo OzAWA" Fossils" : Kiyoshi ASANO, Toshiaki TAKAYAMA

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Trans. Proc. Palaeont. Soc. Japan, X.S., Ko. 84, pp. 179-189, pis. 22, 23, December 30, 1!:'·11586.

THE GENERA NEPHROLEPIDINA AND EULEPIDJNAFROM NEW ZEALAND
KUNITERU MATSUMARU
Department of Natural Science, Saitama University, Urawa, Japan:=. "" - : / - 7 / I;' iJ• ~'":> i!J( I i',·j· ;,s t- 7 ;::r v !::" ·T· 1 7 },'r:,!:: ::r..- v !::" -r· ·i 7 l,<:t\ :-=- =>-- ' / " - =;;/ I:·O):ft.\/j • ,;-~Ji:',jil,j",I,',';O)rjJti/[llt&)J:t!J.I•,'iiJ·GifUI', ~t: lepidocyclinas ~t0!NL t.:ij:,~!,J~. )}:v)·-·:f.ii@~ (·~"L·II'Ifill Nephrolepidina orakeiensis (KARRER), N. hornibrooki MATSUMARL',
sp. nov., N. howchini CIIAP:-.IA:" and CRESPIN, Eutepidina dilatata dilatata (MICIIE·
LOTTI) (,:' ~,'(\j]IJ L 7':' .:::. 0) ·j ·t,, Nephrolepidina h.rl, t::.IMJ L "( IJ:. II ;j;.:f; J::. eFT- A I· 7 ~ T
CO 'I' i-kl!tO)Jt!J.m ;J• •:.,
1',-j· ~:, N ephro!epidin a !:: lbli!d&~·]·l, f.!TfO) ;]:fili c, -& /.: :');)Ufil}~lf;f,~IVJG!r-:::-.lt.:,
K: )L If\ !lH,;·u

Recently, VAN DER VLERK (1959, 1963lproved biometrically that there is a tendency of the deuteroconch to enclose theprotoconch with advance of age, that is
to say, from the lower horizon to theyounger. This tendency was designatedas embryonic acceleration of the lepidocyclinas, and is important for studies on
the phylogeny of the lepidocyclinid species.
The writer examined some specimensof lepidocyclinas from the Otaian, top
Altonianjbasal Clifdenian and Waiauan
stages of New Zealand for evaluation of
the classification and evolution of thefamily Lepidocyclinidae. In this article,
the New Zealand Nephro/epidina and Eulepidina are described in the systematics.
and some species of the former genus
from Japan and Australia are comparedwith one another to interpret their phylogenetic interrelationship.

Introduction
The classification of the family Lepidocyclinidae has been established mainly on
the structure of the embryonic chambers.
Recently. with the progress of study on
the lepidocyclinas, the structure of theembryonic chambers has been classified
into the isolepidine to nephrolepidine
types or the nephrolepidine to trybliolepidine types, and their degree of variation within a species has also beenstudied. As the result, the lepidocyclinas
with the isolepidine. nephrolepidine or
trybliolepidine types of embryonic chambers should be reconsidered from thespecies level.
The stratigraphic distributions of thetwo genera, Lepidocyclina with isolepidine type and Nephrolepidina with nephrolepidine type and with trybliolepidine
type embryonic chambers. are known tohave restricted ranges, but with overlapping one another.

Acknowledgments
Deep appreciation is expressed to Dr.
N. de B. HORN !BROOK of the New Zealand.

Received Sept. 30, 1970; read Sept. 13, 1970
at Tokyo.

179

180

Kuniten~ JI!IATSUMARU

Geological Survey, for his kind offer of
New Zealand lepidocyclinas from theOtaian, top Altonian/basal Clifdenian andW aiauan stages and to Prof. Kotara
HA TAl of the Institute of Geology andPaleontology, Tohoku University, for hiskind helping in various ways and forreading of the manuscript, to Prof.
Emeritus Dr. Shoshiro HANZA wA, Prof.
Kiyoshi ASANO and Associate Prof. Tamio KOTAKA of the same Institute, Tohoku University, for their kind advice.and criti::i:om during the present study.

orakeiensis, sp. nov., Trybliolepidina aff.
howchini, Miogypsina cf. irregularis andAmphistegina lessoni. The planktonic
foraminifers from thi3 locality belong to
the range zone of Catapsydrax dissimilis.

o•
JAPAN

Localities, stratigraphic Positions
and Ages
The lepidocyclinas treated in this study
are from four localities (Text-fig. 1). The
descriptions of the larger foraminifers,
stratigraphic position and· age of each
locality are according to FINLAY (1947)
and HORNIBROOK (personal communication).
Locality 1. N18/569 (=GS733, same
locality, but different collection times),south head of Hokianga Harbour on thewest coast, north of Auckland, North
Island. The larger foraminifers fromthe "Orbitolite" bed at this locality are
Eulepidina sp., Nephrolepidina orakeiensis,
Miogypsina cf. irregularis and Amphistegina lessoni. This bed belongs to the
Otaian stage and has been correlatedwith the Aquitanian of the European·standard stages. The bed has yieldedsuch planktonic foraminifers as Globo.quadrina dehiscens, Catapsydrax dissimilis.and Globigerina woodi.
Locality 2. NS0/825, Whakoau Stream,
north of Gisborne on the east coast of
the North Island. The larger foraminifers from the sandstone at the base of
the massive mudstone (Otaian), at thislocality are Eulepidina sp., Nephrolepidina

o•

,,

0

S CAL E 1000 KM

Text-fig. 1. Index map showing the localities of New Zealand lepidocyclinas. 1:
Auckland, 2: Gisborne, 3: Greymouth, 4:
Pourere. The localities of Nephrolepidina
from Japan and Australia are shown bysolid dots.

.586.

Nephrolepiclina and E1depiclina

Locality 3. S44/559, Alexander St ..
Greymouth on the west coast of the South
Island. The larger foraminifers fromthe base of the Blue Bottom Formation
at this locality are Nephrolepidina sp.,
Trybliolepidina aff. batesfordensis. Heterostegina 2 spp., Cycloclypeus cf. posteidae.dodekaseptus, Gypsina cf. howchini andAmphistegina lessoni. The age of thisformation is at the boundary between
the New Zealand top Altonian and basal
Clifdenian stages, corresponding almost
to the base of the Langhian and at orjust below the earliest occurrence of·Globigerinoides glomerosus.
Locality 4. F5938, Pourerere. east
~oast of Dannevirke, North Island. The
larger foraminifers from the fine sandstone conglomerate and green sandstone-comprise Trybliolepidina n. sp. aff. rutteni.
Heterostegina 2 spp., Cycloclypeus indopacificus var. terhaari, Operculina cf. victoriensis 0. sp., 0. n. sp. aff. complanata:and Amphistegina lessoni. The planktonic foraminifer from this locality of
the W aiauan stage is Globorotalia mayeri
mayeri, and corresponds to the upper:stage of Indonesia.
Notes on the Geology
The marine Tertiary of New Zealandis distributed in the North Island in the
Auckland and Hamilton region, the east
-coast region of Gisborne, Hawke's bay
region and the southwest coast of
Wanganui. In the South Island, they
are mainly distributed in the east coast
region of Canterbury, the west coast
region of Greymouth and the south coast
region of Clifden.
Many stages have been proposed for
Tertiary rocks, and these comprise the·stratigraphical scale of New Zealand(FINLAY and MARWICK, 1940, 1947). Ac-cording to HORNIBROOK (1958), the New

181

Zealand lepidocyclinas occurred from theOtaian, Altonian. Clifdenian and W aiauan
stages.
JENKINS (1966) established planktonic
foraminiferal zones and judging fromhis zoning, the Otaian, Altonian, Clifdenian and Waiauan stages are represented by Globigerina woodi connecta,
Globigerinoides trilobus trilobus, Globigerinoides glomerosa curva and Globorotalia mayeri mayeri zones, respectively.
HOR:\IBROOK (1967, fig. 4) correlated the
New Zealand planktonic foraminiferal
zones of JENKINS with the Caribbean
planktonic foraminiferal zones of BoLLI
and BLOW. The New Zealand stages fromtheir respective planktonic foraminiferal
zones and occurrences of lepidocyclinas
are correlated with Zone N. 4, Zone N.
7, Zone N. 8 and Zone N. 12 (?)- to Zone
N. 13 of the proposed zonal series of
BLOW (1969).
Lepidocyclinas studied
The lepidocyclinas from the four localities stated above are of two types. One
type is a small shell, less than 3.3 mm
in diameter of the megalospheric form
and less than 5.4 mm in diameter of the
microspheric form, with less than 62
percent in the deuteroconch enclosureof the protoconch (=factor A). The
other type is a large shell, more than
6 mm in diameter and with more than70 percent in factor A value. The former
type belongs to the genus Nephrolepidina
and the latter to Eulepidina. The genus
Nephrolepidina is used in this paper,
when the structure of the embryonicchambers is the nephrolepidine type andthe number of specimens possessing such
type exceeds 80 percent of the total
number of specimens from one locality.
The following four species were identified from the four localities.

182

Kuniteru MATSUMARU

Locality 1. Neplzrolepidina orakeiensis(KARRER)
Eulepidina dilalata dilatata(MICHELOTTI)
Locality 2. Nephrolepidina orakeiensis(KARRER)
Locality 3. Neplzrolepidina hornibroolli
MATSUMARU, sp. nov.
Locality 4. Nephrolepidina howchini
CHAPMAN and CRESPIN
It is important to examine the embryonic acceleration of the specimens tointerpret the evolutionary developmentof the New Zealand lepidocyclinas. The
embryonic acceleration is represented byfactor A value and the biometrical analysis of the specimens is as follows.
1. Nephrolepidin a orakeiensis(KARRER)
Factor A (.%)=35.90; 44.19; 45.24.
3 specimens from locality 1 andlocality 2.
2. Eulepidina dilatata dilatata(MICHELOTTI)
Factor A (%)=71.05; 75.50; 74.68;
74.77; 75.00.
5 specimens from locality 1.
3. Nephrolepidina hornibrooki
MA TSUMARU, sp. nov.
Factor A(.%. m*±s**)=49.77±4.80.
16 specimens from locality 3.
4. Nephro!epidina howchini
CHAPMAN and CRESPIN
Factor A (%)=53.70; 55.38; 58.70;
61.36.
4 specimens from locality 4.
Though sufficient conclusions can not
be gained from these data, it appears
that the factor A values of Nephrolepidina oralleien sis. N. lwrnibrooki and
N. howchini show an increase in acceleration.
m*=mean, s**=standard deviation.

From the factor A data of Nephrolepidina howchini from Australia, and·
Nephro!epidina japonica and N. angulos([from many localities in Japan. the relationship of their embryonic acceleration
was examined. The factor A values ofNephrolepidina hornibrooki from New
Zealand and N. howchini from Australia
are judged to be of normal distribution:from the examination of X2 test (P*>0.05, :;**=2), and the statistical significance of the difference between the·means of factor A of those species is·tested by calculating the statistic t according to the formula (SIMPSON, RoE·.
and LEWONTIN, 1960).

N. lwrnibrooki:

N 1 =16, mds 1 =49.77±4.80..

N. howchini
N 2 =31. m 2 ±S 2 =51.64±8.43.
The difference is judged to be nonsignificant at the 5 percent level and 45degrees of freedom (J.J=N 1 +N 2 -2). Thismeans that the factor A of the populations of both species actually does notdiffer from one another. Thus, it isconcluded that the Nephrolepidina hornibrool<i bearing rock facies at the topAitonian/basal Clifdenian stages can becorrelated with the Nephrolef)idina howchini bearing limestone of the Batesfordian stage in Australia. And, Nephrolepidina howchini from Australia (MATSUMAIW, 1971) can be correlated \Vith thelower to middle part of the zoning of·
the three form groups based upon factor
A of the Japanese Nephrolepidina.
The shells of many generations ofNephrolepidina hornibrool<i from their·P*=probability, v**=degrees of freedom.

586.

Nephrolepidina and Eulep,idina·diameter (= Y) and thickness (=X) are
included in the following rejection ellipse(Text-fig. 2).

Phrolepidina, the allometric equation of
Y = bX" is calculated as follows (Textfig. 3) :Nephrolepidina ora!wiensis !KARRER):
Y=l.95X"" 9 ", n=17
Nephrolepidina hornibrool<i MATSUMARU, sp. nov.: Y =3.47X"·"', n=7
i\'ephrolepidina howchini CHAPl\'iA;\"
and Cl~ESPJN: Y =0.22X 1 " 10 , n=8

mm
3.0
>II

1-

.u

Cll

;:::-1200
111000

Cll

E

0

D

183

"":' BOO

1.0

a.ra; 600Ed 400
01.0 mm0

Thickness(=X)

e-O.SO)~(Y- 2.27} 1
0.81
0.27
N=23,

8= B0°21'

Text-tlg. 2. Rejection ellipse showing theposition and variation of Nephrolepidina
hornibrooki MATSl'Mi\Rli, sp. nov. based upon
the diameter and thickness parameters of
the shell.

Nephrolepidina howchini (MATSUMARU,
1971) fall in the following ellipse.

From the ellipses it is proved thathornibrooki is smaller than howchini andthat both species have essentially different shells. The species identification in
lepidocyclinas is by the size and ontogenetic development of the shell. The
characters of the shell through ontogeny
are represented by the half di:1meter andhalf thickness of the first to last voluti::>ns of the lateral chambers. From theme:tsurements of the New Zealand Ne-~ 200
150 "T-f---r--.-r-~-.
80 100200
400~

Half Thickness (=X)•: Nephrolepidina orakeiensis

v = 1.9 5 xo.99
o: Nephrolepidina hornibrooki
v =3.4 7 xo.94+: Nephrolepidina howchini
v =0.22 x1.4 o
Text-tlg. 3. Relationship between the half
diameter and half thickness of the shellthrough ontogeny of NejJhrolepid?:na orakeiensis, NejJhrolepidina hornibrooki and Nephrolepidina howchini.

Though some variation is noticed in
the characters stated above among theindividuals, in the c:tse of the japanese
Nephrolepidina, there are no remarkabledifferences in the relation between thevalue of the exponent a of many individuals and that of only one individual.
Though a of the three Nephro!epidina
species from New Zealand was not calculated for many individuals, it is considered that the data of NejJhrolepidina
ora!wiensis (cr=0.£'9), N. hornibrooki (a=Kunite1·u MATSUMARU

184

BLOW ~gi-ons
1969)
. ..

~~·one.

J'A PANNEW ZEALAND

AUSTRALIA

"'I

~

ZONE

Langhian.
N9~

Form Group~ .

T

M

Ill

~

-~

~

,j;;

.!

~

_g

0)

~

'V

::'

T 1 - -~g11

Burdigalian

:;~

§

::

1

.s

.s

o

.c:

0

0

-&~

-&.!):o1---------+-N 6-+----]_ -]_~~Aq u it ani an

Q.

.!!

N 4

~

~.

0

·-

-~~

~J
·a

_go

~

Q,

II?

.!j'

·

"""
II
c:~
c:
~-r- ~::::::= ~...:::::::::::.~ ....1....__
q-f.-- .'!?
.c:~ :~- .-f---g·~

Til

a.

~~
:: 11
~ \S

Ol

,.,

9
.c:
1 - - - - - - - - + - - - - , - - i - - - - f - II t---- II+-+---- ~
N8

en

'jj

--~

E

0

o
c:

g r-+-~

~

Q.

~

.c:

~

e-a
"'

-<=

-

-i

c:

'8

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.!::-

<:

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"'

4
~~~~--~~~;-r---~-~--~~-t~~======t:i~~r:+--<:~~-----L
N3<: -Chattian

Plank. F.= PI anktonic Foraminifera
Text-fig. 4.

Supposed phylogenetic trends of the genus NePhro!.epidina
from Japan, Australia and New Zealand.

0.94) and N. howchini (a=l.40) are representative of each species.
The evolutionary development of theembryonic acceleration of factor A andallometry through ontogeny of the individuals of Neph"rolepidina, and theirstratigraphic occurrences, the following
supposed phylogenetic tribes can beconstructed (Text-fig. 4).

1968. EulePidina dilatala dilatata(MICHEL-

lnaugura!.-Dissertation,
Ludwig-Maximi!.ians-Univ., p. 51.-55, pl.
3, fig. 1.

OTTI), LANGE,Pl. 22, figs. 28-38

Shell lenticular or elongate lenticular
in shape, outline polygonal, gene,rallywith pillars and pseudopillars distributedon general suaface of shell. Shell diameter 5.85* to 8.80 mm, thickness at
central portion 1.30 to 1.60 mm.
Embryonic chambers in equatorial section of eulepidine type, bilocular ; protoconch large. 505 to 760 microns in diameter, 430 to 715 microns in height, deuteroconch large 1035 to 1255 microns indiameter, 250 to 430 microns in height,
embraces protoconch along its outer

1861. Q;·bitoides di!.atata MICHELOTTI, Etudes
sur le mioc. inf., p. 17, pl. 1, figs. 1-2.

5.85* : This specimen is not a perfect form,
but a errosive one.

Description of Species
Family Lepidocyclinidae SCHEFFEN, 1932
Genus Eulepidina H. DOUVILLE, 1911
Eulepidina dilatata dilatata (MICHELOTTI)586.

Nephrol epidina and Eulepidina

circumference. Ratios of protoconch diameter to deuteroconch diameter 1.39 to
2.04. Factor A of degree of deuteroconch to enclose protoconch 71.1 to 75.0
percent in five specimens. Partition between two embryonic chambers 15 to 35
microns in thickness. Embryonic chambers ellipsoidal in general, extend towards periphery.
Equatorial chambers arranged in subcircular to polygonal rings in equatorial
section, vary ontogenically in shape fromarcuate through ogival to spatulate.
Lateral chambers large, subcircular to
subreniform in shape in transverse section, more or less spacious, especiallywide in tiers over embryonic chambers,
but narrow in tiers just above embryonicchambers.
Remarl<s: The present species resembles Eulepidina eodilatata H. DOUVILLE
in shell form. but differs in the larger
embryonic chambers and larger enclosureof deuteroconch to protoconch. Eulepidina dilatata dilatata differs in generalshell shape from Lepidocyclina (Eulepidina) ephippioides (jONES and CHAPMAN)
from the Kamanga grit, at the base of
the Tanakau group and from the basal
part of the Betilonga Limestone of
Guadalcanal, British Solomon islands(COLEMAN. 1963).
Depository: New Zealand Geological
Survey, Lower Hutt, New Zealand.

Genus Nephro/epidina H. DOUVILLE, 1911

Nephro/epidina orakeiensis (KARRER)
Pl. 22, figs. 1-27

1864. Orbitoides oralzeiensis KARRER, Novara
Exped., Ceo!. Theil, Yol. 1, p. 86, pl. 16,
fig. 21.
1926. Miogypsina orakeiensis KARRER, CHAPMAI', N.Z. Geol. Surv., Pal. Bull., no.

185

9, p. 94, pl. 1, fig. 21, pl. 18, figs. 6a-b,
pl. 20, fig. 2.

Shell of moderate size, measuring 2.0
to 3.2 mm in diameter a·nd 0.6 to 1.1 mm
in thickness. Shell lenticular to inflated.
outline polygonal or rounded, sometimes
with conical pillars on top of central
boss of inflated shell. and a narrow
peripheral portion composed . only ofequatorial chamber layer. Bilocular embryonic chambers of nephrolepidine type.
Factor A of degree of the deuteroconch
to enclose the protoconch varies from35.90 to 45.24 percent. Protoconch small,
245 to 255 microns in diameter and 160
to 220 microns in height, deuteroconch
large, 320 to 365 micr'ons in diameterand 120 to 185 microns in height. Ratiosof protoconch diameter to deuteroconch
diameter 1.29 to 1.42. Partition between
protoconch and deuteroconch solid andthin, 9 to 11 microns in thickness. Embryonic chambers ellipsoidal in generaland extend towards periphery. Allometryof half diameter to half thickness throughontogeny given in Text-fig. 3.
Equatorial chambers of a single layer,of arcuate form in nepionic stage, changing from ogival to hexagonal in neanic
stage through ontogeny, polygonal oroften circular in arrangement in equatorial section. gradually increasing thickness of roofs and floors of equatorialchambers from embryonic chambers toperiphery.
Lateral chambers large, subcircular to
subreniform in shape, characteristic pillars with one concentric ring in transverse section, more or less spacious,
arrangement of lateral chambers fromnarrow in tiers just above embryonicchambers to wide in tiers over embryonic chambers. Eight or nine layers oflateral chambers per tier over embryonicchambers in adult specimens.

186

Kuniteru MATSUMARU

Remarks: The species was originallydescribed on the specimens from Orakei
Bay, Auckland by KARRER in 1864. According to CHAPMAN (1926), KARRER's
original description of this species is asfollows : " The shell is 2 mm to 5 mm indiameter, circular in outline, somewhat
wavy on the tapering edge, which isusually broken; somewhat compressed,
convex towards the centre and generally
more so on one face; both sides ornamented with irregular excrescences
which cover the surface with a network".
CHAPMAN describes this species as "theshell shows a median layer of angularcell, which are apparently arranged withregularity one another; this layer o.:cupies about one tenth of the thickness ofthe shell at the centre; upon this thelayers of the greater portion of the shellhave cells longer than broad and 6 to 7
in number; they are apparently separated
by strong walls, of long rectangular form,
apparently dissimilar, and sloping upon
the axis of the median shell layer ; theshells of the median layer are of similarsize, rounded, or pentagonal and regular ;the embryonic cell is very much larger
than the remainder, and enclosed on one
side by two similarly larger, crescentic
cells". CHAPMAN more or less doubts
whether KARRER's species belongs to
Miogypsina (=the writer thinks to be
J\!Iiolepidocyclina) or to Lepidocyclina but
places his species in I'v!iogypsina.

FINLAY (1947) notes this species to bea lepidocyclinid. The writer puts thisspecies in the genus Nephrolepidina based
upon the nephrolepidine structure of theembryonic chambers and identified it with
Orbitoides orakeiensis.
Nephrolepidina orakeiensis resembles N.
morgani in shell form, but differs in the
allometry and the tendency of the deuteroconch to enclose the protoconch.
Depository: New Zealand Geological
Survey, Lower Hutt, New Zealand.

Nephrolepidina hornibrooki
MATSUMARU, sp. nov.
Pl. 23, figs. 12-37

Shell of moderate size, measuring 1.80
to 2.80 mm in diameter and 0.60 to 1.00
mm in thickness. Position and variationof diameter and thickness of shell given
in Text-filS. 2. Shell lenticular or sometimes inflated, outline steroid in megalospheric forms and polygonal or roundedin microspheric forms. narrow peripheral
portion composed only of equatorialchamber layer. Some conical pillars distributed on central part of megalosphericshell surface and stout conical pillars on
general surface of microspheric shellsurface.
Embryonic chambers of nephrolepidine
to trybliolepidine type of bilocular embryonic chambers. Factor A of degree

Explanation of Plate 22
Figs. 1-27. Nephrolepidina orakeiensis (KARRER).
1, 3, 5-6, 8, 10, 13, 15-16, 18-19, 21: equatorial view; 2, 4, 7, 9, 14, 17, 20, 22: yerticalview; 11: vertical section; 12: centered oblique section; 23-25: equatorial plane; 26:
nearly centered transverse section; 27: vertical section; 1-12: Locality 2; 13-27: Locality
1; 1-10: x7.5; 11: xl9.5; 12: x20.0; 13-26: x7.5; 27: x20.5.
Figs. 28-38. Eulepidina dilatata dilatata (MICHELOTTI).
28, 33, 35: equatorial view; 29, 34, 36: vertical view; 30-32, 37-38: equatorial plane; 2832: Locality 1; 33-38: Locality 2; 28-38: x 7.5.

MA TS UMARU.

M A TS UMAR U

Ne phrolepidina and Eulepidina

and K UMAGAI p hoto .

Plate 22

586.

Nephrolepidina and Eulepiclina

of deuteroconch to enclose protoconchranges from 43.08 to 60.42 percent, andmean value and standard deviation in 16
equatorial sections, 49.77±4.80. Protoconch small, 150 to 335 microns in diame·ter and 115 to 205 microns in height,
deuteroconch large, 265 to 420 microns
in diameter and 80 to 150 microns inheight. Ratios of protoconch diameterto deuteroconch diameter 1.15 to 1.90.
Partition between protoconch and deuteroconch solid and thin, 3 to 14 microns
in thickness. Embryonic chambers ellipsoidal, extend towards periphery, allometry of half diameter to half thickness
through ontogeny given in Text-fig. 3.
Equatorial chambers arcuate in nepionic stage near embryonic chambers,
through ogival to hexagonal form inneanic stage through ontogeny in equatorial section, polygonal in arrangementin equatorial section, increasing thicknessof roofs and floors of equatorial chambers
from embryonic chambers to peripheral
portion in vertical section.
Lateral chambers large, subcircular to
subreniform in shape in transverse section, more or less spacious, arrangementof lateral chambers from narrow in tiers
just above embryonic chambers to widein tiers over embryonic chambers. Number of lateral chambers per tier overembryonic chambers seven lateral layers
in adult.
Remarks: The present new species
resembles Nephrolepidina oralwiensis in
the allometry, but differs in the outlineof the megalospheric shell form and thetendency of the deuteroconch to enclose
the proto~onch. It resembles Nephro!epidina howchini in shell form, but differs from the latter in the allometry
and the tendency of the deuteroconch
to enclose the protoconch.
Depository: New Zealand Geological
Survey, Lower Hutt, New Zealand and187

IGPS* coli. cat. no. 91691 (Holotype).

Nephrolepidina howchini CHAPMAN
and CRESPIN
Pl. 23, figs. 1-11

1932. Lepidocyclina howchini CIIAPl\IA:" andCHESPI:--1, Canterbury ll1us., Rec., vol. 3,
p. 94, pl. 13, figs. 18-19.
1943. Lepidocyclina gippslandica CRESPI:--1,
Proc. Roy. Soc. Vic. (N.S.), vol. 55,
pt. 2, p. 165-168, pl. 3, figs. 1-7, pl. 6,
figs. 22-28.
1943. Lepidocyclina batesfordensis CRESPIN,
Ibid., p. 170-173.
1964. Lepidocychna howclzini CHAP:\IA:" andCHESPI:", CAHTER,
Ceo!. Surv. Vic.
mem. no. 23, p. 137-140, pl. 17, figs.
290-292.

Shell of moderate size, measuring 2.30
to 3.25 mm in diameter and 0.60 to 0.90
mm in thickness. Shell lenticular, outline polygonal or steroid. moderate peripheral portion composed only of equatorial chamber layer.
Embryonic chambers of nephrolepidine
to trybliolepidine type. Factor A values
from 53.70 to 61.36 percent. Protoconch
small, 220 to 365 microns in diameter and
170 to 295 microns in height. Ratios ofprotoconch diameter to deuteroconch diameter 1.74 to 2.23. Partition between
protoconch and deuteroconch solid andthin, 11 to 26 microns in thickness.
Embryonic chambers ellipsoidal extend
toward periphery. Allometry of half diameter to half thickness through ontogeny given in Text-fig. 3.
Equatorial chambers arcuate in nepionic stage near embryonic chambers,
through ogi val to hexagonal form in
IGPS*=Abbreviation for Institute of Geology and Paleontology, Faculty of Science,
Tohoku University, Sendai.

Kuniteru MATSUMARU

188

neanic stage through ontogeny, polygonal
or often circular in arrangement in equatorial section.
Lateral chambers, subcircular to subreniform in shape in transverse section.
more or less spacious, 8 layers per tierover embryonic chambers in adult.
Remarks: The New Zealand specimensshow more steroidal outline in shell shapethan the Australia specimens, but the
ontogenetic developments of diameter tothickness both in the former and thelatter agree with each other. Judging
from the factor A values, the former
may be from a horizon younger thanthe latter, although the former is based
upon the factor A values of only four
specimens.
Depository: New Zealand Geological
Survey, Lower Hutt, New Zealand.
Reference Cited
Br.ow, W.H. (1969): Late middle Eocene torecent planktonic foraminiferal biostratigraphy. In BRb~:-ii~IA)J, P. and RE~Z,
H.H. (Ed.J, Proceedings of the First International Conference on Planktonic Microfossils, vol. 1, p. 199-421, figs. 1-43,
pis. 1-54.
CIIAP:VI.\~, F. (1926): Cretaceous and Tertiary foraminifera of New Zealand. N.Z.
Ceo!. Surv., Pa!aeont. Bu!l., no. 11, p. 1119, pis. 1-22.
CIL\P~IA~, ?.]. (1963): Tertiary larger Foraminifera of the British Solomon Islands,

Southwest Pacific. Micropaleont., vol. 9,
no. 1, p. 1-38, figs. 1-3, pis. 1-9.
FINLAY, H.]. and MARWICK, ]. (1940): The
divisions of the upper Cretaceous andTertiary in New Zealand. Trans. Roy.
Soc. N.Z., vol. 70, pt. 1, p. 77:.135.
- - and - - (1947) : New divisions of the
New Zealand upper Cretaceous and Tertiary. N.Z. jour. Sci. Tee/mol., vol. 28,
no. 4, p. 228-236, tab. 1.
FINLAY, H.]. (1947): Foraminiferal evidence
for Tertiary trans-Tasman correlation.
Trans. Roy. Soc. N.Z., vol. 76, pt. 3; P327 -352, tab. 1..
HoRNIBROOK, N. DE B. (1958): New Zealand
upper Cretaceous and Tertiary foraminiferal zones and some overseas correla-·tions. Micropaleont., vol. 4, no. 1, p. 2538, pl. 1, tabs. 1-2.
- - (1967) : New Zealand Tertiary microfossil zonation, correlation and climate.
In I-IATAI, K. (Ed.J, Tertiary Correlations.
and Climatic Changes in the Pacific, P29-39, figs. 1-6.
]ENKINS, D.G. (1966a) : Planktonic foraminiferal zones and new taxa from the Danian
to lower Miocene of New Zealand. N.Z.
jour. Ceo!. Ceophys., vol. 8, no. 6, p. 10881126, figs. 1-15.
- - (1966b) : Planktonic foraminiferal datum
pianes in the Pacific and Trinidad Tertiary. Ibid., vol. 9, no. 4, p. 424-427, figs.
1-2.
MATSUMARU, K. (1971): Studies on the genus
Nephrolepidina in Japan. Tohoku Univ.,
Sci. Repts., 2nd ser. (Ceo!.), vol. 42, no.
2, p. 97-185, figs. 1-42, pis. 9-26.
SIMPSON, G.G., RoE, A. and LEWONTIN, R.C.

Explanation of Plate 23
Figs. 1-ll. Nephrol~~pidina howchii1.i CIIAP:viAN and CRESPIN.
1-2, 5: equatorial view; 3-4, 6: vertical view; 7-10: equatorial plane; 1.1: vertical section~
1-11: Locality 4; 1-10: x7.5; 1.1: x19.5.
Figs. 12-37. Nephrolepidina hornibrooki MATSUMARU, sp. nov.
12, 14, 16, 18, 21: equatorial view; 22, 24: equatorial view of microspheric form; 13, 15,
17, 19-20: vertical view; 23, 25: vertical view of microspheric form; 26, 28 (Holotype,
IGPS coil. cat. no. 91691), 29-31, 33-37: equatorial section; 32: equatorial section of
microspheric form; 27: vertical section; 12-37: Locality 3: 1-25: x 7.5; 26-37: x 20.0.

MATSUMARU:M A T SUMA R U

N ephTolepidina and E~depidina

and K uMAGA I photo .

Plate 23

586.

Nephrolepidina and Eulepidina(1960): Quantitative Zoology. Harcourt
Brace Co., New York, p. 1-440.
VLERK, I.M. VANDER (1959): Problems andprinciples of Tertiary and Quaternarystratigraphy. Geol. Soc, London, Quart.

189

jour., vol. 115, no. 457, p. 46-63, figs. 13, tab. 1.
- - (1963) : Biometric research on Lepido·
cyclina. MicroPaleont., 'I[Ol. 9, no. 4. P425-426, figs. 1-2, tab. 1.

Trans. ?roc. ?alaeont. Soc. Japan, N.S., No. 84, pp. 190-19;5, pl. 24, December 30, 1971

587.

THE DISCOVERY OF THE CYCAD-LIKE LEAFLETS WITH
TOOTHED MARGIN FROM THE LOWER CRETACEOUS
ITOSHIRO SUB-GROUP, THE TETORI GROUP,CENTRAL HONSHU, JAPANTATSUAKI KIMURA
Mejiro Gakuen _Woman's Junior College

and
SI-IINJI SEKIDO
Science Education Center of Ishikawa Prefecture

TllXRiTI!f:, li.li\.H3EIM·Hlr cr~·Msill!fl:) i;'Gri~**:~!Wli.;>)<liJ.;;, ~-r-;;~Jd~O)'JE5i!.: 1[4:'&0!t 1965 if'., 'fl!J[JIIxi1rW i>fHl-J Ill:imf:b/t/J.',-f" 0 ~t:ili:P~':· · l'l'i~i1Li',''i~'rci:0 N'/i.P G, ~U~v::. '-'' i?
t.::0L-v•JlU1io)&...>0~-r-;;~KZI~fl'll* 2 if8l~r%tl£U:, :nu=.~-r·;;~fir::..t.,v•-c,c<:::.v•t:>c:.;sL-v•JlU1iO)j;,{J),¥,ft Encephalartos -c·£b0 c 2':lc-cv•0;J:, ~*j'<O)foj){,j;, iJ,~i: Stangeria,

Bowenia, Zamia, Macrozamia ;t;J:.v: Microcycas ts.c·:::.tilU!!~t:lt£lf,~O)j;,0{,0")/;~?9iW!!-\J£llltifrt.~:<D 0 c l, :> t.:::rc~ ~ ~;'&JJIJ-J-.;;. tNfXI! c vitJJ:dJ:",,t- i.l· t- ft1::i ~ -r ·.'1 ~kJX·:-c·
iW!!~~Jill\I:JO)j;,0{7Uvi:J.dJ:*1t'f?O)rW\;;js:(;l:, 1962 '.j:, VAKHRAMEEV f:J:.-:> "C-t" ~- ·;; ?'f>l"i!i:OYT;';IIi~lill!*i;·G\"R'E · i\cUoctq1J: Neozamites ),'i:i::-)xi-0, :::.o)~iii!:liiiJ·!-Ih' v-tJII rjlif,Eft)i.)).~(!-'-f1'i11E3ili!*i.l·0 3 ftM:r-:1 Gh-c~,-, 0i.l:, :::.:::. i:i\cli1U- 0t,'\:of>:ii.:CO) c?.t c t ;;;J,cL-tJ:V•(7)-c", Neozamites elongata sp. nov.!::: t--crli-?'i"-.l-.;s, :::.cr)J(.ljli:/-"' ~ 7llfl170Hi:O)

<,

<,

* tt

:tw<t::~~c ~:lc-ct.-•0,;~

JljJ • WJ Ji it ;1:

as a new species under the name of Neozamiles originally established by V. A.
V AKIIRAMEEV (1962).
We should like to thank the staffs ofthe Komatsu City Museum, who co-operated for us in collecting material, and to
thank Dr. T. KOIWAI vvho kindly coworked with us in reading the articles
written in Russian. Our thanks are also
extended to Drs. T. l\1_ HARRIS. V. A.
VAKHRAMEEV, N.D. V ASSILEVSKAJA and
V. A. KRASSILOV, who sent us their valuable articles continuously, which havebeen indispensable to the present study.

In the past year, we and our co-workers
found two fragments of the cycad-likefrond ·be:J.ring pinnate leaflets with spe-cially toothed margin on their both sides,from the equivalent of the Lower Cretaceous Kuwashima sandstone-shale alternation member, the Itoshiro Sub-Group,
the Tetori Group, located along the uppercourse of the Mekkodani, Ozo, Oguchimura, Ishikawa-gun, Ishikawa Prefecture.
We here describe the above specimens
Received November 4, 1970; read }(ovember
7, 1965 at Chiba.

190

587.

Cycad-like Leaflets

The recent development of our knowledge regarding the living Cycads owes
much to the studies conducted by C. ].
CHAMBERLAIN, ]. SCHUSTER and P. GREGUSS. Among the above, for the studyof fossil members, more useful is GREGUSS's recent work (1968) with the detail
description in which most species of living Cycads were done respectively intheir xylotomy, leaf-morphology and also
leaf-epidermis.
So far as leaf-margin is concerned, it
has been known that spinose margin ischaracteristic feature of living Enceplw·
lartos, but is not always so; in most
species, margin is markedly spinose on
one or both sides and at the apex, but ina few species perfectly entire as in E.
humilis, E. eximius, E. lelzmanni and etc.
Besides this African genus Encephalartos, there are considerable numbersof species belonging to various genera
represented by the leaflets with toothed,spinose or serrate margin ; Stangeria
eriopus, Dioon spinulosum, Bozcen ia serrulata have toothed or serrate margin;
Bozcenia spectabilis, Stangeria paradoxa
both show deeply dissected margin ; most
species of Zamia and Macrozamia fawcettii show toothed or serrate margin in
the apical half or only at the apex oflamina; moreover in J\1icrocycas there
are some species with finely dentate
margin at the apex or whole around.
In case of fossil, these cycad-like leaflets with spinose or toothed margin being
encountered, they used to have beenchristened calling the name from Enceplzalartos based only upon the marginal
character.
FO:\TAINE described some cycad-likeleaflets with serrate margin and withCten is-type nervation forming meshes
under the name of Enceplzalartopsis nervosa (1889, p. 174, Pl. LXX, fig. 4; Pl. LXXI,
figs. 3-4; Pl. LXXII, figs. 3-4), which. he

191

considered to be an original type of therecent Enceplwlartos. As was stated by
SEwARD (1917. p. 508), however, its direct
relation not only to Ctenis but also torecent Encep!wlartos without net-worked
nervation, can not be made clear becauseof lacking the point of attachment to therachis. It seems to be impossibl.e to compare unexpectedly or directly between
the Cretaceous leaflets and the recent
cycadean genera depending only on having toothed or serrate margin.
Ctenis exilis originally described by
HARRIS (1964) exceptionally shows theleaflets with roughly toothed margin.

Genus Neozamites V AKHRAMEEV. 1962
In 1962, V. A. V AKHRAMEEV established
two new genera, Neozamites and Enceplzalartites, based on the cycad-like pinnaewith particularly toothed leaf-margin
from the Lower Cretaceous deposits near
Yakutsk. East Siberia, and also described
two new species in Neozamites and asingle new species in Encephalartites, they
were Neozamites verchojanensis (typespecies). N. lebedevii and Encep!wlartites
leipzigii. The following generic diagnosis
of Neozamites was given (freely translated) by V AKIII<AMEEV; ''frond pinnate;pinnae comparatively long and narrow,attached to the both sides of the uppersurface of rachis, narrowing gradually
towards the apex with several spinous
teeth; pinna base more or less auriculate
and sometimes cordate; margin spinously
toothed asymmetrically. each tooth directed forwards as well as the directionof apical teeth; nerves diverging dichotomously forking near the margin." Moreover based on the cuticles obtained fromsome specimen of N. lebedevii, he stated"epidermal cell walls conspicuously sinuous; stomata bennettitalean." It was also

192

Tatsuaki KIMURA and Shinji SEIGDO

mentioned by him that "among thesedescribed, both characters, toothed pinna
margin and cordate and asymmetrical
pinna base were distinct from those ofother cycad-like genera already knowri,
such as Sphenozamites and Otozamites."
Both of which are close in outline. especially in the nervation, to the present
genus. In Sphenozamites, pinnae are asymmetrical oval or rhomboidal in form
with contracted or cuneate bases. Sphenozamites, as was stated by SEWARD (1917,
p. 587), was establi3hed only on the outline of pinnae for the purpose of convenience in case of description. In Otozamites pinnae are varied oblong, ovate
or sword-like in shape, usually representing auriculate projection only at the upper basal margin. Among all species
belonging to both Sphenozamites and Otozamites, the toothed pinnae have not been
illustrated but S. belli SEWARD (1917, p.
588, fig. 629) in which the lower marginof pinna was drawn as loosely undulated
and some Otozamites from Primorie tobe stated later.
Another new genus Encephalartites wasdiagnosed by V AKHRAMEEV as well;
frond pinnate; pinnae long and narrow,ribbon-like with somewhat contracted

Text-figs. 1-2:

Encephalartites leipzigii

VAKHRAMEEV.

1: Toothed margin and the nervation ( x 1)
2: Mode of Attachment of leaflets ( x 1)
Both after V AKHRAMEEV (1962, p. 128, Fig.
la-b).

base and attached to both sides of uppersurface of rachis with their whole bases;
margin spinose on both sides ; nerves
being sent off not radially from a pointof attachment of leaflet as seen in Neozamites, but in parallel. about 12 in number at base. lateral nerves forking dichotomously, one of which reaching the
tip of each spines respectively, as shownin Text-figs. 1 and 2 (after V AKHRAMEEV).

Neozamites elongata KIMURA
et SEKIDO, sp. nov.
Pl. 24, figs. 1-4; Text-tigs. 3-4

Description of species: Frond pinnate,
more than 11.5 em long, 13.5 em or morewide at the widest, tapering gradually
towards the anterior ; pinnae long andnarrow, ribbon-like, the largest more than
7 em long, 1.1 em wide measured at 1 em
distance from its base, tapering gradually towards the apex, somewhat asymmetrical, slightly cordate at base, attached to both sides of upper surface of therachis with narrower central portion andsuboppositely with wide angle, getting
smaller in size towards the anterior and\Vith acute angle, about 50 degrees to therachis; margin asymmetrically toothed,having 5-6 pairs of conspicuous spines;
the apex of leaflet dissected into 3-5,
each spine sharply pointed, directed forward, the rest of margin loosely undulated ; nerves originate radially from thenarrower pinna base, then generally pareaile! to the direction of leaflet, about 12
in number at the portion of 1 em distancefrom the base, often dichotomously forking but not anastomosed, one at themedian part thick, looking like a midnerve, in posterior half or one-third, but
not appearing in smaller leaflets, outside
nerves reaching each spine. Any of epidermal cells have not been prepared be-587.

------a---3

Text-figs. 3-4: Neozamites elongata
KIMURA et SEKIDO sp. nov.
3:
4:

Model of traverse section of a frond ( x 1)
Model of the outline of a leaflet ( x 1)

ca·use of impression alone. Fructification
not known.
Description of specimens: Two fragments of frond have obtained. Pl. 24,
fig. 1 may show the middle portion ofa frond in which pinnae are attached toboth sides of upper surface of rachis
suboppositely with the central point ofcordate base as seen in recent speciesof Encephalartos, Ceratozamia and etc.
The distorted pose in the present figure
is not natural but due to the deformationin its preservation. Pl. 24, fig. 2 shows
the thickened median nerve and Textfigs. 3 and 4 show the models of traverse
section of a frond and the shape of leaflet respectively. Pl. 24, figs. 3-4 showrespectively an apical portion of a frond
and partly enlarged one in which the
thickened median nerve fail to notice.
judging from both Pl. 24, fig. 1 and fig.
3, a frond may have reached more than
30 em in length.
Remarks and comparison: Though, taking the recent knowledge regarding thecycadean leaflets into consideration, there
might remain some doubts in establishing some new genera only based upon
the leaflets with spinose or toothed margin, it is clear that the present specimens

Cycad-like Leaflets

193

in outline agree closely with the diagnosis of V AKHRAMEEV's genus Neozamites in representing spinose margin andsomewhat cordate base, the central partof which sti::ks to both sides of the upper surface of rachis, sending off nervesradially, not anastomosed.
At first the present specimens should
be compared with the type specimen, N.
verc!wjanensis described originally by
V AKHRAMEEV from the Lower Cretaceous
Batilifsk. Ekseniafsk form ations andtheir equivalents near Yakutsk and by
VASSILEVSKAJA (1966) from the equivalent of the above, East of the Lena. The
specimens derived from the bank of theRampeska (V AKHRAi\IEEV, 1962, Pl. XII,
fig. 1) resemble closely the present ones
in both outline and size, but the former
is more strongly asymmetrical and conspicuously toothed and each itself dissected more into 2-3 spines. Anotherillustrated specimens of N. verclwjanensis(Ibid., figs. 2-4; V ASSILEVSKAJA, 1966; Pl.
III, figs. 1-2) have short pinnae and abnormally dissected tips, from which thepresent ones are clearly distinguishable
in outline.
We have also been informed the occurrence of this species from the Lena basin
by KIRITCHKOVA but we couldn't referher article (1966) to the present study
for want of her paper.
The specimens derived from the Lower
Cretaceous at the bank of the Vilyuy
under the name of Neozamites lebedevii(V AKHRAMEEV, Pl. XIII, figs. 1-3), shouldbe, at second, compared with the present
ones. The former shows that the leafbases are conspicuously a uriculated, spinose margin also marked, very sharply
pointed at apices, and nerves are radialand evenly distributed, no midnerve-likethickening as seen in some leaflets in N.
verchojanensis (Ibid., Pl. XII. fig. 1) andin the present ones, can be noticed at all.

194

Tatsuaki KIMURA and Shinji SEKIDO

So N. lebedevii is clearly distinguishable
from both N. verchojanen sis and the present new species.
At third, KRYSHTOFOVICH and PRINADA(1932, p. 369) reported some specimens
under the name of Otozamiles denticulatus originally but not illustrated, fromthe Lower Cretaceous of Southern Primorie. V AKHRAMEEV (1962) renamed theabove Neozamites denticulatus (KRYSHTOFOVICH & PRI:'·~ADA), depending on theinformation by F. N. CHERNISHEV A who
had observed this specimen deposited at
the Leningrad Museum of Natural History. According to V AKHRAMEEV, N.
denticulatus shows that leaflets are very
short in length with symmetrical baseand with less toothed margin.
KRASSILOV (1967, p. 151. Pl. XL, fig. 1)
described the similar but more elongate
specimens from the Lower Cretaceous
Galenkov Series, Southern Primorie. His
single illustration shows pinnate habit
and the pinnae elongate-oblong in outline,
4 em long. 1 em wide at the widest withloosely undulated margin and with contracted base in both sides. These N.
denticulatus are easily distinguishable inoutline of leaflet from the other.
As is stated above, since the present
specimens might be distinct from thoseof any other species already described.
so we have hereby made them a new
species to give the name of Neozamites
elongata.
Of course. the present species is also

distinguishable from Encephalartites leipzigii in the nature of the leaf-base andits nervation. Besides the Potomac specimens by FONTAINE in question of their
attribution, spinose or toothed cycad-likeleaflets such as Neozamites and Encep/wlartites. seem to be limitted in their distribution geographically to Far East ofEurasia and in age to the Lower Cretaceous.
Locality and geological horizon: Equivalent of the Kuwashima sandstone-shale
alternation member (Lowest of Lo\ver
Cretaceous). the Itoshiro Sub-Group, theTetori Group, near Nijidaki (anothername; Benidaki), a branch of the Tetori,Ozo, Oguchi-mura, Ishikawa Prefecture.
Occurrence: Rare.
Reg. Nos.: Kl\1-N-1 (Pl. 24, fig. 1:
Holotype) and KM-N-2 (Pl. 24, fig. 3:
Paratype) ; Both type specimens havebeen deposited at the Komatsu City
Museum, Ishikawa Prefecture.

References Cited
CHAMBERLAIN, C.]. (1919): The Living Cycads. New York & London (Hajne1·•s reprint, 1965), pp. 1-172.
FoNTAINE, W.N. (1889) : The Potomac orYounger Mesozoic Flora. U.S. Geol. Surv.,
Mon. 15, pp. 1-377, 180 pis.
GREGUSS, P. (1955): Identification of Living
Gymnosperms on the basis of xylotomy.
Budapest. pp. 1-263, 350 pis., 8 tabs.
- - (1968) : Xylotomy of the living Cycads

Explanation of Plate 24
Figs. 1-4.

Neozamites e!ongata KIMURA et SEKIDO sp. nov.

Fig. 1:
Fig. 2:
Fig. 3:Fig. 4:

Holotype (Reg. No. KM-N-1) x 1
Enlarged (partly) of holotype, x 4
Paratype (Reg. No. KM-N-2) x 1
Enlarged (partly) of para type, x 3

KIMURA and SEKIDO:

Cycad-like L eaflets

Plate 24

587.

with a description of their leaves andepidermis. Ibid., pp. 1-260, 18;) pis., 80
text-figs.
HARRIS, T.M. (1964): The Yorkshire Jurassic Flora II. British Mus. (Nat. Hist.),London, pp. 1-191, 7 pis., 67 text-figs.
KRASSILOV, V.A. (1967): Lower Cretaceous
Flora of Southern Primorie and its stratigraphical significance (in Russian). Far
East Ceo!. Inst. Siber. Branch, Acad. Sci.
USSR, lvloskow, pp. 1-364, 93 pis., 33
text-figs.
KRYSHTOFOVICII, A. & PRI:"iAOA, V. (1932) :Contribution to the Mesozoic Flora of theUssuriland. Bull. U. Geol. Pros. Serv.
USSR, Bd. 51, fasc. 22, pp. 363-374, 2 pis.
SEWARD, A.C. (1917): Fossil Plants, Vol. III.
Cambridge, pp. 1-656.

VAKHRAMEEV, V.A. (1962): New Lower Cretaceous Cycadophyta from Yaktsk (inRussian). Jour. Paleont., No.3, Moskow,
pp. 123-129, 2 pis., 2 text-figs.
- - (1964) : Jurassic and Early Cretaceous
Floras of Eurasia and the palaeoftoristic
provinces of this period (in Russian).
Acad. Sci. USSR, Trans., Vol. 102, Moskow, pp. 1-261.
- - et al. (1970) : Palaeozoic and Mesozoic
Floras of Eurasia and phytogeography ofthis time (in Russian). Ibid., Vol. 208,
pp. 1-423.
V ASSILEVSKAJA, N.D. (1966) : Some Early
Cretaceous plants from Zhigansk district(in Russian).
Sci. Rep. Paleont. &Biostratigr., No. 15, Leningrad, pp. 49-76,
7 pis.

·-----------

Ishikawa-gun
Itoshiro
KomatsuKuwashima
Mekkodani

195

Cycad-like Leaflets.

1.i )II 'Iii\
11 @

s

;J'

~

~

tb

I]

rrt B'-

Nijidaki (Benidaki)Oguchi-mura
Ozo
Tetori·--·----~--~---U¥/ECW~)

M'i:
Jfrf.

1·

p

tJ

i""
.j ••

llX

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 84, pp. 196-204, December 30, 1971

588.

ON SOME BRYOZOA FROM NEAR NAMIOKA-CHO, MINAMITSUGARU-GUN, AOMORI PREFECTURE, JAPANTOMOKO HAY AMI
Institute of Geology and Paleontology, Faculty of Science,
Tohoku University, Sendai, Japanilf~~'?..i¥1~'-lWiN!li'ilii!Jfiifi:OJ;: 1:l"!hft.15:

iJ.l.iltr~Jtl!mlli'iJIIIJ'fj"ifi:OJ ::k~Rill!!l\lf i)' G c1:T.'ilft.
9 ~ 11 t.ill~~}JIJ L-ilcifi!G L.t.:, 11~~. ltl!~~:::. "?v''"C f;J:, f!f*O) ~UN,llffiJE~Jilt.150)1iJf'?E~*cfiiJG~/lL-t.~:Hi5ffifli~r.~L.t.:.
l2i 1.k fJi!: :Y15~1~f.:O)"t'¥1i'S-i" :Q.

Introduction and Acknowledgments

assistance in the field.

The fossil bryozoans described in thepresent article were collected by thewriter in September, 1970 from a single
locality of a roadside cliff near Aizawa,
southeast of Namioka-cho (Text-fig. 1).
These form the second fossil record of
the group from the younger Tertiary
rocks of Aomori Prefecture and contribute to our knowledge of the fossil bryozoan fauna of northeast Japan.
At this place the writer expresses her
thanks to Professor Kotara HAT AI of theInstitute of Geology and Paleontology,
Tohoku University, for his kindness andguidance throughout the course of thepresent work. Thanks are due to Professor Takehiko IWAI of the Departmentof Geology, Faculty of Education of theHirosaki University, for his kind guidance to the fossil locality and information on the geology of the area. Acknow.ledgments are also due to Miss Hiroko
UNO of the Hakodate School of the Hokkaido University of Education for her

NORTHEASTHONSHUText-fig. 1. Locality map of fossil Bryozoa.

Stratigraphic position and geologicalage of the fossil bryozoans

The locality of the fossil bryozoan specimens treated in this article is the roadside cliff near Aizawa, northeast of Namioka-cho (Text-fig. 1). At this locality,
there is exposed fossiliferous dark grayReceived January 14, 1971; read November.22, 1970, at the Hiroshima University, Hiro.shima.

196

588.

Bryozoa j)·om A.mnori

tuffaceous siltstone intercalated withrounded pebbles and cobbles of andesite
and soft rocks. The fossils are abundant
but most are rather fragile, and comprisebalanids, brachiopods, gastropods, fragments of spine and carapace of echinoids,
foraminifers, and bryozoans, etc. According to IwAr (1965a), this outcrop is one·Of the prolific fossil localities of the Daishaka Formation.
The Daishaka Formation is distributed
rather widely around the type locality
which is near the Daishaka Tunnel ofthe main Ou railway, Minami-TsugaruGun, Aomori Prefecture. The formationin the northeastern part of the Hirosaki
Basin has yielded many fossils such as
foraminifers (ASANO, 1938), molluscs(NOMURA and l-!ATAI, 1935; IWAI, 1960,
1962, 1965b), brachiopods (NOMURA andl-IATAI, 1935; l-!ATAI, 1940; KOTOH, 1957),·spines of echinoids, bryozoans (KAT AOKA, 1957), and calcareous algae, etc.
The stratigraphic position of the Dai.shaka Formation is shown in Text-fig. 2.
Age

Formation

Pliocene

Tsurugasaka Formation
Daishaka Formation
Otakizawa Member of Ka. Fm.
Kareizawa Formation

Miocene Toyamori Formation
Umanokamiyama Formation
PreTertiary
Text-fig. 2. The stratigraphic position of theDaishaka Formation, after IwAr (1965a).

The fossil bryozoans
The bryozoan specimens occur as broken stems and as branches or fragments

197

of encrusting forms. The bryozoans identified amount to 11 species distributed
among nine genera, and these are listed
in Table 1.
The bryozoan species listed in Table 1,
comprise several types, among which the
Cellariiform type represented by suchgenera as J'vficroporina, Diatosula andMyriozoum is the most abundant. The
next common is the Reteporiform typewhich is characterized by Reteporellina
and Schizoretepora, and only very few
incrusting zoaria of the Membraniporiform type are found in the fauna. The
fauna contains species that have beenrecorded living along the coast of Alaska, Aleutian Islands, Queen Charlotte
Island, and from Hokkaido, all in the
North Pacific.
In comparison with the fossil bryozoans
recorded by KATAOKA (1957) from thetype locality of the Daishaka Formation.
there are only two species in common,
namely, Microporina articulata and My-

riozoum suhgracile.
The fossil bryozoan fauna describedin this article comprise species which
are typically northern in distribution.
The fauna is characterized by the dominance of Cellariiform and Reteporiform
types and an assemblage of this kind oftypes is characteristic of rather shallow
water (STACH, 1936; LAGAAIJ and GAUTIER, 1965). The thermal conditions of
the sea in which the fauna once lived isthought to have been cold as is inferredfrom the distribution of the living counterpart species. The conditions were
probably similar to the present day east
coast of Hokkaido to northeast Aomori
Prefecture. This inference seems to be
in good agreement with the results obtained by ASANO (1938) from the foraminifers, by NOMURA and l-IATAI (1935)
from the molluscs, and by KATAOKA(1957) from his studies on the bryozoans

Tomoko HA YAM!

198
Table l.

Bryozoan species identified frcm 1\amioka-cho.
Distribution

Genus and species

Microporina articulata (F ABR ICil's)

Recent, Japan Sea; Off Torishima and Paramushir Island; California; Greenland andQueen Charlotte Island. Pliocene. Sado (Niigata Pref., Japan); Miocene, Ishikawa ?ref.,and Hokkaido.

Tricellaria sp.
Diatosula marion en se (Bus I<)

Recent, Prince Edward Island (80-150 fms);
Off Heard Island (75 fms) ; Off Marion Island(50-75 fms).

Schizobrachiella subhexagona (ORTMANN)

Recent, Sagami Bay.

Cystisella midwayanica CANU & BASSLER

Eocene, North America.

? Cystisella americana CANU & BASSLER

Recent, Gulf of Mexico (32 fms); Porto Rico.

Schizoretepora tumescens (ORTMANN)

Recent, Paramushir Island and Hokkaido ;·Sagami Bay (40 fms) and (200-230 fms).

Costazia rota (MAcGILLI\"RAY)

Recent, Juan Fernadez (35m); Australia;·
Cape Horn and East Africa; and Gulf of
Manaar.

Schismopora chrysalis CANU & BASSLER

Recent, Philippine Region (20 fms).

Myriozoum subgracile D'OR BIG NY

Recent, Japan (shallow water); Kara Sea(33-127 m) and Mourmane Sea (90 m) ; Paramushir Island; and Alaska (15-25 fms) and·(13-22 fms).

Myriozoum coarctatum (SARS)

Recent, Pacific Coast of Canada (Juneau
20 fms); Alaska and Off Washington; andQueen Charlotte Island .
..

of the Daishaka Formation.
The geological age of the Daishaka
Formation based upon the molluscs (NoMURA and HAT AI, lw AI). the brachiopods(NO;viURA and HATAI. HATAI), and thebryozoans (KATAOKA), is Early Pliocene.

Descriptions and Remarkson the Bryozoa
Order Cheilostomata BusK, 1852
Family Calpensiidae CANU
and BASSLER, 1923
Genus j'vficroporina LEVINSEN, 1909

Microporina articulata (FABRICIUS, 1821)
1905. Ce!!aria borealis (BusK), ROBERTSO!\,_____ - - - - - - - - - ·p. 287, pl. 14, fig. 86; pl. 16, fig. 102.
1929. Microporina japonica CANU and BAss-LER, p. 139, pl. 14, figs. 9-11.
1933. Cellaria borealis BusK, OKADA, p. 214.
1936. MicroPorina articulata (F ABRICit;s).
SAI<AI<URA, pp. 259-267, pl. 15, figs. 1-9 ..
1950. Microporina borealis (BusK), OsBURN,.
p. 106, pl. 11, fig. 2.
1956. Microporina articulata (FABRIC 1us),
MAWATARI, p. 120.
1958. Microporina articulata (F ABRicws),
ANDROSOVA, p. 125, text-fig. 40.
1965. Microporina articulata (FABRICics),
MAWATARI, p. 603, figs. 62a, b.
1970. Microporina articulata (F ABRICJcs),
HAYAMI, p. 325, pl. 35, figs. 1, 2.

Description: Zoarium erect, cylindrical,.
of jointed segments, branching formed of
9-10 rows of zooecia. Zooecia large, elongate, alternate, surrounded by a raised.

588.

Bryozoa .fro?n Ao1no1·i

199

Text-fig. 3. Bryoz oa from Na mioka- cho.

Dia lo su! a marion en se ( B us ~-: ) , x 30, show ing ordinar y an d b roke n ov ice ll e d zooecia.
? Cyslis e!L a americana C ANU and B ASSLEH, x 30.
.Schis mopora chrys alis C ANU a nd B ASSLE R, x 30, s howi ng a portion of large, mass i ve ,and multil a me ll ar zoar ium.
d . Tricellaria sp., x 30.

a.
b.
c.

thread; frontal very poro us. A pertur esemicircu lar. A vic ul aria distal, distribut ·ed at irregular inter va ls; rostrum triang ul a r , pointin g downward.
Measurements (in mm) : Zooecia: Lz= 0.72- 1.28, vV z = 0.28- 0.36.
DejJosilory: IGPS * co il. cat. no. 86809.
R ema rlzs: ROBERTSON (1908) noted that
J\;fi croporina articulata is a spec ies strict1y northern in distr ibution and statedthat it has not been reported south of
the Queen Charlotte Is lands.
Disl ribution: Recent, Param ushir Island a nd Hokkaido (t\'IAWATA RI , 1956) ;Param ush ir Is land (OKADA, 1933) ; Sagha lien 20.5- 66 m (ANDROSOV A, 1958) ; A laska
14 fms (OSBURN, 1950) ; Japan Sea (CANU
and BASS LER, 1929). Fossil, P liocene, Sado
(N iiga ta Pref.) (SAKAKURA, 1936); Miocene,
Hokka ido (HAYA iVIT, 1970).

Fam il y Scrupocellariidae LEVINSEN, 1909
IGPS*= abbrev iat ion for Insti t ute of Geolog y:a nd Paleontology, Tohoku Un i versit y, Sendai.

Genus Tricellaria FLEM ING, 1828

T rice LLaria sp.
Te x t-fig . 3- cl

DescrijJtion: On ly a fragmenta l zoa rium , uniserial (?). Zo oecia large, elongate;
opeci um oval larger than half as long as
front . Late r a l avicula ria la r ge. tri:mgular ro strum hooked at tip; front a l av icu la r i3. wa ntin g. Trace of two sp ines
at dista l part of mu r a l rim , and one ofscutum located a lmost at middie of la teral mura l rim.
Mea surements (in mm) : Zooecia: Lz= 0.60 - 0.72, Vh = OAO; Opec ia: lop = 0.40,
wop = 0.32.
DejJosilory : lG PS co iL cat. no. 86810.
R emarhs : As no comp lete zoa rium wasfound in the collect io n it cou ld not be
determined w hether it was uniserial orbiser ia l. However, the reference of t hepresent spec im ens to TriceLLw·ia seems to
be witho ut doubt, because there are nov ibracula or av icularia on the dorsa ls urface. This species may represent an

Torno leo HA YAM I

200

undescribed form, but n.aming is avoidedat present because of the lack of wellpreserved specimens.
Family Stomachetosellidae CANU
and BASSLER, 1917
Genus Diatosula CANU and BASSLER, 1927
Diatosula marionense (BusK, 1884)
Text-fig. 3-a
1884. Myriozoum marionense BusK, p. 171., pl.
23, fig. 6.
1929. Diatosula marionense (BusK), CANU andBASSLER, p. 293.
1.953. Diatosula marionense (BusK), BASSLER,
p. 200, text-fig. 1.50-'1.

Description: Zoaria free. branches subcylindrical. Zooecia immersed, lines ofseparation not visible; frontal wall withlarge irregular pores. Orifice rounded
above and lower margin possessing Ushaped sinus. A pair of small avicularia,
slightly prominent, present above orifice;ovicell when present occur on each sideof orifice. Ovicell subimmersed, large,
round. Interzooecial avicularia large. long
triangular. its tip pointed proximally.
Afeasurements (in mm): Orifice: lor=
0.20, wor=0.18; Interzooecial a vicularia:
lav=0.48, wav=0.28.
Depository: IGPS col!. cat. no. 86811.
Remarlzs: CANU and BASSLER (1929)
designated ivfyriozoum marionense BusK,
1884, as the genotype of Diatosula.
Distribution: Recent, Prince Edward
Island 80-150 fms, Marion Island 50-75
fms. Off Heard Island 75 fms (BUSK. 1884).
Fossil, this is the first record.

Family Schizoporellidae ]ULLIEN, 1903
Genus Schizobrachiella CANU
and BASSLER, 1920

Schizobrachiella subhexagona(ORTMANN, 1890)
1890. Schizoporel!a subhexagona ORTMANN, P51, pl. 4, fig. 3.
1957. Schizobrachiella subhexagona (ORTMANN), HARMER, p. i026, pl. 74, figs ..
17, 19, 24-26.
1965. Schizobrachiella subhexagona (ORTMANN), MAWATARI, p. 615, text-figs.
107a-c.

Description: Three fragmental specimens. Zoaria incrusting; z6oecia quincuncial, somewhat flat, short. Frontal
wall with many rather large pores which
extend around distal side of orifice. Orifice with a low, thin peristome; proximal
margin nearly straight at both ends, withwide and slightly rounded sinus. No·
a vicularia.
Measurements (in mm) : Zooecia: Lz=0.80. W z=0.68.
Depository: IGPS coil. cat. no. 86812.
Distribution: Recent, Sumbawa, 69-73m (HARMER. 1957); Japan (ORTMANN,
1890). Fossil. this is the first record.

Family Smittinidae LEVINSEN, 1909
Genus Cystisella CANU and BASSLER. 1927
C,)stise/la midwayan ica CAI':Uand BASSLER. 1920
1.920. Cystise!!a midwayanica CANe and BAss-LER, p. 479, pl. 8, figs. 5, 6.

Description: Only four fragmental zooecia; zoarium incrusting. Zooecia separation not visible. but interior subhexagonal ; frontal somewhat convex andfinely granulated. Peristome thin, little
salient in distal part; with 6 to 8 stout
spines. Two small cardelles present at
low position of peristome. Avicularia
form long chamber, median and conical;

588.

Bruozoa from Aomori

its orifice turned toward aperture.
Measurements (in mm): Zooecia (interior): Lz=0.32-0.40, Wz=0.30-0.36.
Depository: IGPS coli. cat. no. 86813.
Distribution: Recent, unknown. Fossil,
Eocene, North America (CANU and BASSLER, 1920).

? Cystisella americana CAI\U

and BASSLER, 1928
Text-fig. 3-bCompared with:
1928. Cystisella americana CAJ\U and BAssLER, p. 113, pl. 15, figs. 7, 8.
1940. Cystisella americana CANV and BAssLER, OsBCRN, p. 439.

Description: Zoarium incrusting, bilamellar or multilamellar. Zooecial separation not visible, but interior large andlong. Frontal fine granulated, two to
three pores present at marginal part.
Aperture large, semi elliptical. A vicularian chamber with a pair of glands, large
and long, almost occupying frontal. Avicularia very large, as large as aperture;
its shape somewhat elliptical, and narrower at distal part. Ovicell not found.
Measurements (in mm) : Zooecia (interior) : Lz=0.80-1.20, W z=0.44-0.46; Aperture: lap=0.32-0.36, wap=0.28-0.32.
Depository: IGPS coil. cat. no. 91692.
Remarl<s: The present species resembles C. americana CANU and BASSLE!{.
1928. in some aspects, but differs from it
in having a longer zooecium, in the orifice touching the avicularium. in larger
apertural dimensions, and by the presence of some marginal pores.
Distribution: Recent. Gulf of Mexico
32 fms (CA;\!U and BASSLER, 1928i ; Porto
Rico (OSBURN, 1940). Fossil, this is thefirst record.

201

Family Reteporidae SMITT, 1867
Genus Schizoretepora GREGORY, 1893
Schizoretepora tumescens(ORTMANN. 1890)
1890. Retepora tumescens OwrMAN:", -p. 34, pl.
2, fig. 20.
1956. Schizoretepora tumescens (OwrM AN!\),
MAWATARI, p. 131, figs. 14a-g.
1958. Retepora imperati var. tumescens (ORTl\-IANN), A:-mRosovA, p. 175, text-fig.
106.

Description : Fragmental zoaria, rather
thick. Zooecia facing one side, elongatesubhexagonal, convex, smooth with two
to six small pores. Orifice circular withshallow sinus. Frontal avicularia of two
kinds, one very large and prominent,
other small and not salient, sometimes
wanting. A pair of spines on distal corner of peristome. Dorsal vibices a little
salient, separated, elongate quadrate or
irregular. Dorsal avicularia small, somewhat acute, elliptical, variable in numbers.
Measurements (in mm) : Zooecia: Lz=0.80-0.84, Wz=0.28-0.40.
Depository: IGPS coil. cat. no. 91693.
Distribution: Recent, Kurile Islands
and Hokkaido (l\IIA WATARI, 1956) ; Sagami
Bay 40fms, 200-300fms (ORTMANN, 1890);
Saghalien (ANDIWSOVA. 1958). FossiL thisis the first record.

Famely Celleporidae BusK, 1852
Genus Costazia NEVIANI, 1895
Costazia rota (MACGILLIVRAY, 1885)
1929. Costazia rota (MA.cGILLIVRA Y),
and BASSLER, p. 431, fig. 3.

CA:--:uDescription: Only two fragmental zooecia ; zoarium incrusting. Zooecial some-

Tomoko HAYAMI

202

what elliptical. Frontal wall smooth withirregular pores at marginal part. Aperture round distally, and with round, widesinus proximally. Peristome thin notprominent. A pair of avicularia, prominent distally, rather large, in peristome
each side. Ovicell not found.
Measurements (in mm): Zooecia (interior) : Lz=0.76, W z=0.56; Aperture: lap=0.24, wap=0.20.
Depository: IGPS col!. cat. no. 91694.
Distribution: Recent, Philippine region
20 fms (CANU and BASSLER, 1929); Juan
Fernandez 35m and Bass Strait (MARCUS).
Fossil, this is the first record.
Genus Schismopora MACGILLIVRAY, 1888

Schismopora chrysalis CANU
and BASSLER, 1929
Text-fig. 3-c

1929. Schismopora chrysalis CANU and BAssLER, p. 430, pl. 62, figs. 11, 12.

Description: Zoarium very large, multi1amellar. Zooecia rather erect, orbicular;
frontal granular and surrounded by scattered areolar pores. Aperture round distally, and straight with wide, shallow
sinus proximally. Peristome thin; a pairof avicularia present at both sides, sometimes wanting. Interzooecial avicularia
sporadic, very few; its shape somewhat
elliptical. Ovicell not found.
Measurements (in mm): Zooecia: Lz=0.80, Wz=0.40; Peristome (with sinus):lper=0.24, wper=0.24.
Depository: IGPS col!. cat. no. 91695.
Distribution: Recent, Philippine region
20 fms lCANU and BASSLER, 1929). Fossil,
this is the first record.
Family Myriozoidae SMITT, 1867
Genus Myriozoum DONATI, 1750

Myriozoum subgracile D'ORBIGJ\Y, 1852
1908. Myriozoum subgracile D'0RBIGNY, RoBERTSON, p. 296, pl. 21, fig. 5.
1929. Myriozoum subgracile D'0RBIGNY, CANU
and BASSLER, p. 512, pl. 65, figs. 9-13.
1935. Myriozoum sub gracile D'OR BIGNY, SAKAKURA, p. 36.
1952. Myriozoum subgracile D'0RBIGNY, OsBURN, p. 514, pl. 64, figs. 3, 4.
1956. lv!yriozoum subgrac£/e D'0RBIGNY, MAWATARI, p. 132, fig. 15f-i.
1958. Leieschara subgracilis (D'0RBIGNY), ANDRosov A, p. 145, text-fig· 72.
1965. Leieschara subgracilis (D'0RBIGNY), MAw ATARI, p. 624, text-figs. 143a-c.

Description: Zoaria free, branches cylindrical. Zooecia immersed. its separation not visible. Frontal wall punctatewith rather large pores. Orifice higher
than broad, sometimes broader than high.
proximal margin straight with a wellmarked sinus. A pair of small aviculsria, occasionally only one avicularium,
lateral or above each zooecium, andmandible directed obliquely downward.
Measurements (in mm): Orifice: lop=
0.16, wop=0.12.
Depository: IGPS col!. cat. no. 91696.
Distribution: Recent, Puget Sound andBaffins Bay (ROBERTSON, 1908) ; Japan Sea
lCANU and BASSLER, 1929) ; Alaska 13-25
fms (OSBURN, 1952) ; Aleutian Islands andJapan (MAWATARI, 1956, 1965); AleutianIslands 10-100 m (ANDROSOVA, 1958). Fossil, Pleistocene, Chiba Pref. (Japan) (SAKAKUJ{A, 1935).
Myriozoum coarctatum (SARS, 1850)
1908. Myriozoum coarctatum (SARS), RoBERT·
SO:", p. 295, pi. 21, figs. 55-57.
1952. Myriozoum coarctatum (SARs), OsBURN,
p. 513, pl. 64, figs. 5, 6.

Description: Zoarium free, branches
cylindrical. Zooecia indistinct, no lines

588.

Bryozoa from Aonwriof separation; frontal a very thick tremocyst with large irregular pores. Orificelonger than wide, rounded distally,straight at sides. proximal border transverse with a narrow sinus. A vicularia
wanting.
l'vfeasurements (in mm): Orifice: lor=
·0.22, wor=0.20.
Depository: IGPS col!. cat. no. 91697.
Distribution: Recent, Off the coast ofOregon and Washington 152fms (OSBURN,
1952) ; Alaska (ROBERTSON, 1908). Fossil,
this is the first record.
References
A;-.;DRosov A, E.I. (1958) : Bryozoa of the older Cheilostomata of the north part of theJapan Sea (in Russian). Issledovaniya
Dalinevostochnykh Morei V, Akademiya
Nauk SSSR, pp. 90-204, 104 text-figs.
AsA:"fO, K. (1938) : On some Pliocene Foraminifera from the Setana Beds, Hokkaido.
japan. jour. Geol. Geogr., vol. 15, pp. 87103.
BASSLER, R.S. (1953) : Treatise on Invertebrate Paleontology, pt. G. Geol. Soc.
America and Univ. Kansas Press. pp. G
1-G253, 175 text-figs.
BusK, G. (1884) : Polyzoa I, The Cheilostomata. in Report on the scientific resultsof the voyage of H.M.S. Challenger, Zoo!.,vol. 10, pt. 30, pp. 1-216, pis. 1-36.
CA"iU, F. and BASSLER, R.S. (1920): North
American Early Tertiary Bryozoa. U.S.
Natl. Mus., Bull. 106, pp. 1-877, 162 pis.
- - and - - (1928) : Fossil and Recent Bryozoa of the Gulf of the Mexico Region.
Proc. U.S. Natl. Mus., vol. 72, pp. 1-165,
34 pis.
- - and - - (1929) : Contributions to the
biology of the Philippine Archipelago andadjacent regions; Bryozoa. U.S. Nat!.
Mus., Bull. 100, vol. 9, pp. 1-685, 94 pis.
HARC\!ER, S.F. (1957): The Polyzoa of the
Siboga Expedition, Pt. IV, pp. 642-1147,
pis. 42-74.
HAT AI, K. (1940): Cenozoic Brachiopoda from

20:3

Japan. Tohoku Imp. Univ., Sci. RejJts.,
2nd Ser., vol. 20, pp. 1-413.
HAYA!\11, T. (1970): Miocene Bryozoa fromSouthwest Hokkaido, Japan. Trans. Proc.
Palaeont. Soc. japan, N.S., no. 79, pp. 316336, 2 pis.
I\L\1, T. (1960): Pliocene Mollusca from the
Xishi-Tsugaru district, Aomori Prefecture, Japan. Saito Ho-on Kai Mus., Res.
Bull., no. 29, pp. 33-45, 1 pl.
- - (1962): The Pliocene deposits and molluscan fossils from the northeast marginof the Tsugaru Basin, Aomori Prefecture,
Japan. Ibid., no. 31, pp. 35-46.
- - (1965a) : The geological and paleontological studies in the marginal area of
the Tsugaru Basin, Aomori Prefecture,
Japan. Educ. Fac., Hirosaki Univ., Bull.,no. 14, pp. 83-155 (in Japanese).
- - (1965b): The geological and paleontological studies in the marginal area of
the Tsugaru Basin, Aomori Prefecture,
Japan. Ibid., no. 15, pp. 1-68.
KATAOKA, ]. (1957) : Bryozoa from the Daishaka Formation (Pliocene), Minami-Tsugaru-Gun, Aomori Prefecture, Japan.
Trans. Proc. Palaeont. Soc. japan, N.S.,
no. 28, pp. 143-153, 1 pl.
KOTOH, J. (1957) : Brachiopoda from the Daishaka and Tsurugasaka fossil zones, Minami-Tsugaru-Gun, Aomori Prefecture.
Ibid., no. 25, pp. 7-10.
LAGAAIJ, R. and GAUTIER, Y.V. (1965): Bryozoa assemblages from marine sedimentsof the Rhone delta, France. Micropaleont.,
vol. 11, no. 1, pp. 39-58, 34 figs., 1 chart.
MA\\'ATARI, S. (1956): Cheilostomatous Bryozoa from the Kurile Island and the neighbouring districts. Pac. Sci., \'OJ. 10, no.
2, pp. 113-135, 15 text-figs.
- - (1965) : Bryozoa: in New illustrated
encyclopedia of the Fauna of Japan (inJapanese). pp. 585-628. Holw ryu-Kan.
No~rt:RA, S. and HATAI, K. (1935) : On two
species of Brachiopoda from the Daishaka
Shell Beds of Daishaka, Aomori-Ken. Saito
Ho-on Kai Mus., Res. Bull., no. 5, pp. 5559.
- - and - - (1935) : Pliocene Mollusca fromthe Daishaka Shell-Beds in the vicinity of204

Tomoko HAYAMI

Daishaka, Aomori-Ken, northeast Honsyil,
Japan. Ibid., no. 6, pp. 83-142, 5 pis.
OKADA, Y. (1933): On a collection of Bryozoa from the Northern Kurile Expedition.
Biogeogr. Soc: japan, Bull., vol. 4, no. 3,
pp. 213-216.
ORTMAJ'\1\, A. (1890): Die Japanische Bryozoenfauna. Arch. f. Naturgesch., Bd. 1,
H. 1, pp. 1-74, 4 pis.
OsBuR:-.;, R.C. (1940) : Bryozoa of the Porto
Rico with a resume of the West Indian
Bryozoan fauna. New York Acad. Sci.,
Sci. Surv. Porto Rico and the Virgin Islands. voi. 16, pt. 3, pp. 321-486, 9 pis.
- - (1950) : Bryozoa of the Pacific coast of
America. pt. 1, Cheilostomatous-Anasca.

AizawaKareizawa
Otakizawa
Tsurugasaka

~'II

iR

.:E?t~.iR)( i~ iR
11!} 'r :!;&:

Allan Hancock Pacific Expedition, YO!. 14,
no. 1, pp. 1-269.
- - (1952) : Bryozoa of the Pacific coast of
America. pt. 2, Cheilostomatous-Ascophora. Ibid., vol. 14, no. 2, pp. 270-611.
SAKAKURA, K. (1935) : Pliocene and Pleistocene Bryozoa from the B6s6 Peninsula(I). jour. Fac. Sci., Imp. Univ. Tollyo,
Sec. 2, vol. A., pp. 1-48, 7 pis.
- - (1936) : On Microporina articulata (FABR ICIUS), a Cheilostomatous Bryozoa.
Jour. Geol. Soc. japan, vol. 43, no. 15, PP259-267, 1 pl.
STACH, L.W. (1936): Collection of zoarial
form with habitat. Jour. Ceo!., vol. 44,
no. 1, pp. 60-65.

Daishaka)( fR :illi!
Namioka-cho
if<! lli'i] 1!1J
Toyamori· =t'lli ~ ~Umanokamiyama .1m J :f~qiJ

Trans. Proc. Palaeont. Soc. Japan, 1\. S. No. 84, pp. 20:5-224, pis. 2:5, 26, December 30, 1911

589.

AMUSSIOPECTEN FROM NORTH AMERICA

AND NORTHERN SOUTH AMERICA*
KOICHIRO MASUDA
Department of Geology, Miyagi University of Education, Sendai, Japan~l:::;lj(:J":> J: Ol-/:1:;ljUI::'fi!lO) Amussiopecten:

Amussiopecten f<l: SAcco (1897)

t~n~~hL".t::).3lt, iltffi!.%JI!!O)ffi::::_:lfdi·G~< O)fllii.lq'llf='T~ht~"lt-'(-O)::(cJ:t£i.I;P,;JJc.:Y;t~~~"l-cv't~O)fC 'Lt1vGi~
f'J:.~<f~fp

c: J: ""·cliP l, ~l::*.i>J: u"f~{ij;;j~fi~G(Cox, 1942), jjf* Amussiopecten O)¥!l'r!;-"? f:_o~@]~~f'J:.,

Cox O)=Ff~11:;Jt;2;, ~lt;lj~ · rj:l;'j~ · ff·l:lii:O) Pectinidae \:::.-::>J,•L"~fl<l6!
Amussiopecten iJ; Late Oligocene fi' G Middle Miocene 1::: fl>!tL", G:i~l::
7 ;< U :b;;kl>ill
5J-1ITlL"v·t~z:: c '2::' TliWJ-90::::. C.iJ;--r::-f!;;t~" -,-t~h·f::,, r,;t9;11TillL'f'J:.:b V
7 11---.::.7 0) Pecten vanvlecki ARNOLD (1907), ]!1,! 1 :/ f'Rll'/t.~O) Pecten antiguensis
BRO\YN (1913), ::I :A !7 · U ;b 0) Pecten preglyptus OLSSON (1922), .....:: ;t- "X .J:.. 7 0) Pecten
churuguarensis F. and H. HODSON (1927) t~ C:'iJ;, AA G i.I·IC Amussiopecten v:::.Jm-9 ::s b0)'"(' <b ::S:::. bPJ'U-:> f-.=o t: G 1::::, Amussiopecten O);ffrf!_E'2::' 7" .J:.. tl.-- 1- • U ::I, 1- V .::. !7"-;; F':J";
J: V~.! ~'/::I iP G 3 t!l!J£Yt!.l t~" :::. 0) 'i', .! 4'- '/::I i.l' G 0) 1 fillliilmf*~i.l;yt~
i;f!:tf- "b 51~~·t l t.:f,~5R.

vc.JE<*<,

fJ B!.~=r---c· t~ v'O)'"C·t!ft!J!O)i\c~vi&J~;Z t~oAmussiopecten f'J:. Flabellipecten r:::.Jif,-9 0 "b O)'"(· <b 0 C. :-JS· ;Z. -c v' 0 "¥::%17.1; =r - r-r -;; / ~t~'.-'0)--r::·, .:t0)5nriO)i.ft!Ui£ l "'" l i.l·l, jiliJ ~fi®iY!.U: [!OJIJ ~ ;1"l:;;,-"' 2: t 0)vcayt~<--c· a; ::s c. m:(;l·fi~ ;c. -c ", z,"/j;:fi(,!))i:"C:'Ii Amussiopecten O)t;'fi~ '2::'W!~ l <iiatr r. t~ fliP, 'Jl'j(J;JJ~ c 0) fW)Jm, [lfJ:nHJl:t; J: V:'fJflifiO)fj['_ljU (: i'E:li:O)I'MJ)!, ;t; J: u: Amussiopecten O)~'J.¥!J·~f:I'I'J~:~'2::'~(,!) L. t.:"

W: 111 4 - t!i:

Introduction

able to Amussiopecten have been described under other genera or subgenera,such as Pecten, Amusium, or Aequipecten.
as will be pointed out below. There·
fore, Amussiopecten can be considered
to be one of the most significant andimportant pectinids among the Cenozoic
Pectinidae of the world, particularly
from the view point of its world wide
but locally limited geographical distribution, more or less restricted geologicalrange and intimate relationships withcertain other pectinids.
The name Amussiopecten has not beenused in describing any American species.
However, Cox (1942) has already pointed

Since Amussiopecten was establishedby SACCO 1897, it has been frequently
recorded from various localities of
Neogene and Paleogene formations inSouth and Central Europe, the Mediterranean Region, Iran, East Africa, Southeast Asia and East Asia, and believed
to represent a tropical or subtropical
pectinid. A.mussiopecten assumes suchvaried morphological characteristics thatit has been frequently confused withrelated pectinids. Some species refer-* Received January 24, 1971: read January
24, 1971 at Tokyo.
205

Koichiro JYIASU DA206

out that a well known Miocene form,
Pecten vanvlecki ARNOLD (1907) describedfrom the Vaqueros Formation in Central
to Southern Californi::l might be referable
to Amussiopecten, but this has not been
up to now confirmed. Moreover, it isreasonable to assume that some species
referable to Amussiopecten should havebeen distributed to the area extending
from the North America to SouthAmerica during the Tertiary period.
The writer has examined numerous
specimens preserved in the Departmentof Geology of Stanford University;
Museum of Paleontology, University ofCalifornia in Berkeley; U.S. Geological
Survey in ·Menlo Park ; U.S. Nationall'vl useum in Washington. D. C. ; California
Academy of Sciences in San Francisco ;Academy of Sciences of Philadelphia;
and Paleontological Research Institutionin Ithaca. The results of these examinations lead the writer to conclude thatseveral species described from the Miocene or Oligocene formations of the West
Coast of North America, Central America,
Caribbean Region, and northern SouthAmerica should be referred to the genus
Amussiopecten. Two species new to science are described in the present article.
In the present article the writer
wishes to describe the morphologicalcharacteristics of some species referable
to Amussiopecten and two new speciesof Amussiopecten, and to discuss therelationship among the species. Moreover, paleontological significance of thegenus Amussiopecten is given.

Acknowledgments

Acknowledgments are clue to Dr. A.
Myra KEEN, Professor Emeritus of the
Department of Geology, Stanford University, Dr. Leo G. HERTLEIN of the Cali-

fornia Academy of Sciences in San
Francisco, Dr. Wend ell P. WOODRING of
the Smithsonian Institution in Washington, D. C., Prof. ]. Wyatt DURHA1VI of theMuseum of Paleontology, University ofCalifornia in Berkeley and Dr. David
M. HOPKINS of the U.S. Geological
Survey in Menlo Park. for their continuous encouragement, criticisms andreading of the manuscript.
During the course of the present work
the writer received support from thefollowing persons to whom he takes
this opportunity to express his deep
gratitude: Drs. Harald A. REHDER andThomas R. WALLER of the Smithsonian
Institution. Drs. Norman F. SOHL and
Warren 0. ADDICOTT of the U.S. Geological Survey, Drs. Horace G. RICHARDS
and Charles GIVENS of the Academy of
Natural Sciences of Philadelphia. Dr.
Katharine V. W. PALMER of the Paleontological Research Institution and Mr.
Joseph H. PECK of the Museum of Paleontology, University of California.
Thanks are due to Mr. Kenji SAKAMOTO of the U.S. Geological Survey in
Menlo Park, who made photographic
work.
Acknowledgments are also due to the
Department of Geology of Stanford
University, Museum of Paleontology of
University of California in Berkeley,
California Academy of Sciences, U.S.
Geological Survey in Menlo Park,
Smithsonian Institution, Academy of
Sciences of Philadelphia and Paleontological Research Institution. for permission to study their collections.
Acknowledgments are also made to
the Ministry of Education of the Japanese
Government, for its permission for thewriter to study abroad.
Historical Review of Amussiopecten

589.

Amussiopecten

Amussiopecten was established by
SAcco in 1897 as a subgenus of Pecten
based upon Pecten burdigalensis LAMARCK, an important Miocene pectinidin Europe, and at the same time hedescribed two subspecies of P. burdigalensis. Subsequently, UGOLINI (1907, 1908)
raised Amussiopecten to generic rank, as
did many later authors, and he transferred eight species known from the
Miocene formations in Italy to thisgenus. Thenceforth, many species orsubspecies of Amussiopecten were described from Miocene formations inCentral and Southern Europe (SCHAFFER,
1910, COSSMAN and PEYROT, 1914,
TELEGD-ROTH, 1914, VON TEPPNER and
DREGER, 1918, EAMES and Cox, 1956, andothers). In 1922 VON TEPPNER summarized the classification of the Tertiary
Pectinidae in his ·' Fossilium Catalogus"
and included 18 species and seven subspecies of pectinids in the subgenus
Amussiopecten of the genus Philippia.
On the other hand, DEPERET and
ROMAN (1910, 1912) did not use theAmussiopecten as either genus or subgenus, but they raised Flabellipecten,
which was also established as a subgenus
of Pecten at the same time as Amussiopecten by SACCO (1897). to generic rank,
and moreover, they included the typespecies of Amussiopecten. Pecten (Amussiopecten) burdigalensis which was originally designated by SACCO, and the otherspecies of Amussiopecten, in the genus
Flabellipecten.
Subsequently, some authors in Europe(ROGER, 1939, VEIGA, 1954, ACCORD!, 1955.
v'
CSEPREGHY-l'v!EZ:-.JERICS, 1960, CTYROKY,
1969. etc.) followed the classification of
DEPERET and ROMAN, and they did not
use the taxon Amussiopecten but assigned
all known species and subspecies ofAmussiopecten to Flabellipecten.
From the Miocene formations de-207veloped in Iran, East Africa and Mediterranean Region several species ofAmussiopecten were described by Cox(1927, 1939), DOUGLAS (1928), and EAMES
and Cox (1956). Also from an Oligocene
formation in Iran Amussiopecten cL
labadyei (d' ARCHIAE and HAIME) wasrecorded by Cox (1936). In 1927 Cox
pointed out that V ala sing!zirensis andVolagendinganensis described by MARTIN(1909) from the Neogene formations in
Java, Indonesia, might also be referable
to Amussiopecten.
In the Far East part of Asia several
species referred to Amussiopecten were
described from the Neogene formationsin Japan and Taiwan (YOKOYAMA, 1922.
1928, NOMURA, 1933, 0TUKA, 1934, SHUTO.
1955. AKIYAMA, 1957, MASUDA. 1962a).
Recently, CHANG (1967) also reported theoccurrence of Amussiopecten sp. from
Early Miocene formation in Taiwan.
Among the species known from East
Asia, Amussiopecten praesignis (YOKOYAMA) has been frequently reported
from the Early Pliocene formations in
the Pacific Borderland of Southern
Japan and Taiwan (MASUDA, 1962b).
No species has hitherto been describedand recorded under AmussiojJecten either
a genus or subgenus from either North
America or South America. In 1969·
HERTLEIN alluded to the occurrence ofAmussiopecten in North and South America, but he did not give any details.
Thus, Amussiopecten in North and SouthAmerica has remained unknown.

Notes on Amussiopecten

In 1897 SACCO gave so simple diagnosis
to the subgenus Amussiopecten thatsome confusion arose between Amussiopecten and Flabellipecten, as mentioned
above.
Later COSSMAN and FEYR CT

208

Koichiro MASUDA(1914, p. 272) gave a better diagnosis of
the subgenus Amussiopecten, as follows:" Shell generally large, a little arched,with a little gape; both valves convex,
but left valve a little less convex thanright valve; auricles a little elevatedand rather long ; byssal notch not very
conspicuous ; ornamentation takes like
Pecten s. s., but more or less obsolete infull adult specimens (Translation)".
The following diagnosis of Amussio·
pecten as a subgenus of Pecten waspresented by Cox (1927) : "Shell orbi·
cular, equilateral. inequivalve, the rightvalve more inflated than the left; ears
fairly large, subequal, the sinus below
the right anterior one scarcely developed;
right valve sculptured with broad,depressed radial ribs, left valve with arather smaller number of much narrower
radial ribs; ribs frequently becoming
obsolate towards the ventral margin;
internal ornament sometimes consistingof broad ribs (corresponding in position
to the external interspaces) concave incross section, with prominently raised
margins, and sometimes of paired narrowribs (as in Amusium), each pair obviously
representing the margins of one of thebroader ribs, the central portion of whichis undeveloped ; ligament-pit deep; cardinal crura fairly well developed, three inthe right valve and two in the left;
auricular crura strong, tuberculiform ".
In 1956 EAMES and Cox raised Amussiopecten to generic rank and at the sametime they also used Flabellipecten as asubgenus of Pecten.
Although PHILIPPI (1900) concluded
that the genus Amusium may belong toa different lineage from that of Amussiopecten, it is considered that Amusium isrelated to Amussiopecten in generalfeatures except for the smooth surface,equally inflated valves, and rather thinshell. Also as pointed out by MACNEIL(1961), the present genus is closely related to the genus Pecten and is believed
to form a continuous lineage between
Pecten and Amusium.

Amussiopecten from Northand South America
So far as examined, among numerous
specimens collected from the Tertiary
strata in North America, Central America, West Indies and South America thefollowing species and subspecies whichhave been described under other generictaxa such as Pecten, Amusium andAequipecten can be considered to be referable to the genus Amussiopecten.
Also it appears that some species
referable to the present genus have frequently been recorded as Amusium fromthe Oligocene and Miocene formations.
For example: Pecten (Pecten) vanvlecki
ARNOLD (1907), Pecten (Amusiwn) antiguensis BROWN (1913), Pecten (Aequipecten) preg!yptus OLSSON (1922), Pecten(Pecten) hawleyi HERTLEIN (1925) and
Pecten antiguensis churuguarensis F. and
H. HODSON (1927). Among them Pecten(Pecten) hawleyi HERTLEIN can be considered to be a synonym of Amussiopecten vanv!ecki (ARNOLD) as discussed
later. Moreover, two new species from
Oligocene and Miocene formations in
Puerto Rico, Trinidad and Mexico, andone undescribed species from the Mioceneof Mexico have been found, but thelatter specimens are unfortunately sopoorly preserved that they are notnamed.
The following species of Amussiopecten
are confirmed by the writer from thegeological formations ranging from theLate Oligocene to Middle Miocene inNorth America, Central America, West
Indies and northern South America.

.589.

Am1tssiopecten

1. Amussiopecten vanvlecki (ARNOLD)
from the Early to Middle Miocene inSouthern to Central California, U.S. A.
2. Amussiopecten antiguensis (BROWN)
from the Late Oligocene of AntiguaIsland, West Indies.
3. Amussiopecten preglyptus (OLSSON)
from the Middle Miocene in Costa Rica.
4. Amussiopecten churuguarensis (F. and
H. HODSON) from the Late Oligocene in
Venezuela and Puerto Rico, West Indies.
5. Amussiopecten harrisi MASUDA, newspecies from the Late Oligocene in
Puerto Rico and Mexico.
6. Amussiopecten woodringi MASUDA,
new species from the Late Oligocene in
Puerto Rico and Early Miocene in
Trinidad.
7. Amussiopecten species from the Early
Miocene in Mexico.

Description
Family Pectinidae
Subfamily Amusiinae
Genus Amussiopecten SACCO, 1897
Amussiopecten SAcco, 1897, p. 53.

Type-species (original designation) :Pecten burdigalensis LAMARCK, 1809.
Miocene (Burdigalian), France.
Geological range :-Oligocene to Pliocene.
Geographical distribution :-Central andSouth Europe, Mediterranean, North andEast Africa, Iran, India, Indonesia,
Taiwan, Japan, West Indies, North
America, Central America and northern
South America.
Remarks :-This genus is characterized
by its medium to rather large, medium
thick, slightly inflated, nearly equivalved
shell; right valve always a little more
inflated than slightly inflated left valve;

209

rather large, angulated auricles withwide and shallow byssal notch ; surfaceof right valve with broad, low, square
sided or flatly rounded radial ribs whichusually tend to become obsolete towards
ventral and lateral margins; left valve
with rounded, low radial ribs and raised.
regularly spaced, fine incremental lines ;radial ribs usually narrower than theirinterspaces in younger stage but usuallytending to become broad and obsolete
towards ventral margin ; interior surface
with distinct paired internal ribs which
are more prominent near ventral margin ;hinge with simple cardinal crura having
a rather distinct provinculum and distinct
auricular crura which terminate distallyin a distinct oblong denticle at eachextremity.
The present genus can be distinguishedfrom Flabellipecten in having slightly
inflated, nearly equivalved shell, squarish
or flatly rounded, low radial ribs which
tend to become obsolete towards ventral
and lateral margins, and rather large,
angulate auricles in the right valve;left valve with rounded radial ribs
tending to become broad and obsolete
to\vards the ventral and lateral margins ;hinge with rather simple cardinal crura
provided with a rather distinct provinculum. Moreover, Amussiopecten is neverprovided with intercalary threads ineither valve but Flabellipecten usually
has intercalary threads in the left valve.
Although from the morphologicalcharacteristics DEPERET and ROMAN(1910) concluded that Amussiopecten
should be referred to Flabellipecten, thewriter believes that from the above
mentioned morphological characteristics
Amussiopecten should be ranked as agenus as did UGOLINI (1907) and EAMES
and Cox (1956). Also that Flabellipecten
should be placed as a subgenus of Pecten,
because Flabellipecten has characteristics

Koichi1·o MASUDA

210

very similar to those of the genus Pecten,
except for the greatly inflated rightvalve. fiat or sometimes concave left
valve, and rather smaller number ofdistinctly squarish. high radial ribs in
Pecten s. s.
Amussiopecten vanvlec!?i(ARNOLD, 1907)
Pl. 23, figs. la-b, 2, 3, 4
1907. Pecten (Pecten) vanvlecki ARNOLD,
Smiths. Misc. Coli., Vol. 50, p. 428, pl.
53, figs. 1, 2.
1907. Pecten (Pecten) vanvlecki ARNOLD,ARNOLD and ANDERSON, U.S. Ceo/.
Surv., Bull., 322, pl. 17, figs. 1, 2.
1925. Pecten (Pecten) hawleyi HERTLEIN,South. Calif. Acad. Sci., Bull., 24, pt.
2, p. 40, pl. 4, figs. 4, 5.
1932. Pecten (Pecten) hawleyi HERTLEIN,
LoEL and CoREY, Univ. Calif. Pub!.,
Bull. Dept. Ceo!. Sci., Vol. 22, no. 3, p.
194, pl. 20, fig, 1 (not fig. 2).
1932. Pecten (Pecten) vanvlecki ARNOLD,LoEL and COREY, Ibid., p. 193, pl. 20,
figs. 3, 5.
1932. Pecten vanvlecki ARNOLD, BRE?v!NER,
Santa Barbara Mus. Nat. Hist., Occ.
Paper No. 1, pl. 2, fig. 6.

Holotype :-U.S. Nat!. Mus., Coil. No.
165305.
Remar!?s :-From the study of many
specimens of A. vanvlec!?i the writerconsiders that the following features
must be added to the original description:
Shell slightly inflated, subequivalve,though right valve a little more convex
than slightly inflated left valve which
is sometimes nearly fiat in younger
stage but tends to become slightly inflated with growth; radial ribs of rightvalve flatly round-topped, more or lesssquarish in profile, rather distinctly
separated from their interspaces andvarying in number from about 13 to 18,

as pointed:.out by LoEL and COREY (1932).
Hinge with simple cardinal crura witha distinct provinculum and distinct
cardinal crura terminating distally inan oblong denticle. Paired internal ribs
well developed near ventral margin.
Comparison :-Pecten (Pecten) hawleyi
was described by HERTLEIN (1925) fromthe same formation containing P. vanvlecki, and was based upon one small
left valve and one· small. poorly preserved right valve. However, according
to the re-examination of type specimens(Stanford University Type Coli. Nos.
19, 20) it seems that P. hawleyi is asynonym of P. vanvlec!?i, that is to say,P. hawleyi has features characteristicof P. vanvlec!?i, such as slightly inflated
left valve with rounded radial ribs that
tend to become flatter and lower towards ventral margin. distinct, raised,
regularly spaced incremental lines anddistinct paired internal ribs. Thus, itappears that P. hawleyi represents animmature form of P. vanvleclli. Also P.
hawleyi was recorded by LOEL and
COREY (1932) from the Vaqueros Formation at Santa Ynez Mountains inSouthern California. However, according
to the writer's study of their typespecimens (Museum of Paleontology,
Univ. Calif. Berkeley, Coli. Nos. 31718,
31719), the left valve can be identified
with P. vanvlec!?i but the right valve
can be considered to be referable toAequipecten andersoni (ARNOLD).
Pecten vanvlec!?i has frequently been
confused with Pecten (Amusium) lompocensis ARNOLD (1906), described from theTemblor Miocene in Southern California,
as already pointed out by LOEL and
COREY (1932), but the former is easily
distinguishable from the latter by its
rather thick shell, squarish, low, flatlyround-topped radial ribs tending to be·come obsolete towards the ventral and589.

Amussiopecten

lateral margins, paired internal ribs
developed at lower part and larger andangulate auricles. The relations between
the shell height and hinge length are
shown in Text-fig. 1. However. becauseof the state of preservation sometimes
the two can not be distinguished fromone another.

211

near Paso Robles, Templeton quadrangle
and Nipomo quadrangle southwest
across the Paso Robles quadrangle tocentral western Adelaide quadrangle,
San Luis Obispo County ; Orange County :Early to Middle Miocene.
Amussiopecten antiguensis(BROWN, 1913)
Xmm

Pl. 26, figs. 4, 5
0

X

200

X
)<

X

X

X

X

xx
X

X

X

191.3. Pecten (Amusium) antiguensis BRm\·:-;,
Proc. Acad. Nat. Sci. Philadelphia, p.
613, pl. 18, figs. 1, 2, 3, 5.
1919. Pecten (Amusium) antiguensis BRm\·1',
CooKE, Carnegie Inst.
Washington
Publ., 291, p. 143, pl. 13, figs. 6, 7.

X
X
0

X

X
o lompocensis

X

X vanv lecki

X~--~10~----~~~---------S~Omm
HINGE LENGTH

Text-fig. 1. Graph showing the relationship between the height and hinge lengthof shell.

Dimensions (in mm):Valve R-L
R R
R
L
L
L
Height 95
105 9278120
80
50
Length 98
110 92 ca. 76
82
125
52
Depth 15*7
ca. 8 ca. 6 ca. 4*-thickness

Type locality :-Mouth of Ballard
Canyon, 2 miles south of Santa Ynez,
Santa Barbara County, California. Vaqueros Formation. Early Miocene.
Distribution :-Santa Y nez and Fox
Mountains, Santa Barbara County; Oak
Ridge, Ventura County; Santa Cruz
Island, Santa Lucia Mountains, andwestern foot hills from northern San
Luis Obispo quadrangle, railroad cutting

Holotype :- Acad. Nat. Sci. Philadelphia,
Coil. No. 1648.
Remarks:- This species was first described by BROWN (1913) from the Antigua
Limestone of Antigua Island, West
Indies, based upor" several poorly preserved specimens and then it was described from the type locality by CooKE(1919). Also it was reported by HARRIS(1926) from the Oligocene formation in
Trinidad.
The type specimens and the topotype
specimens (U.S. Nat!. Mus., Coil. No.
197139) are unfortunately not well preserved but this species is characterized
by the following features: Shell medium
in size and thickness, compressed; subequivalve, but right valve a little more
inflated than left valve, forming anangle of 120° or a little more at apex.
Right valve slightly inflated, with about
16 very low. flatly round-topped radialribs tending to become obsolete towards
ventral and lateral margins and fineincremental lines ; radial ribs. broader
than their interspaces; auricles rather
large, somewhat angulate, with fine concentric lines; anterior auricle with wide

212

Koichiro MASUDA

and shallow byssal notch. Left valve
slightly inflated, with very low, roundedradial ribs tending to become obsolete
and broad towards ventral margin, andregularly spaced, raised, fine incremental
lines. Hinge with simple cardinal crurawith fine provinculum; auricular crura
terminate distally in a distinct denticle
at each extremity. Interior surface with
distinct paired internal ribs near ventral
margin.
BROWN (1913) stated that the radialribs of this species are about 13 in
number but the writer finds that they
are about 16 in number.
According to the writer's study of the
hypotype specimen preserved in the
Paleontological Research Institution(Coil. No. 25058) which was illustrated
by HARRIS (1926) as Pecten antiguensis
BROWN from the Adivinanza quarry,east of Princes Town, Trinidad, it seems
that HARRIS's specimen can not be referred to P. antiguensis; that is to say,the specimen, consisting of a rightvalve, differs from the present species
in having a smaller apical angle andlarger number of more elevated radialribs, which are more rectangular in
profile and broader than those of P.
.antiguensis. Therefore, this can be considered to represent a new form ofAmussiopecten described in the present
paper.
Comparison :-Amussiopecten antiguensis differs from A. vanvlecki by its
smaller shell, larger apical angle andvery low radial ribs.
Dimensions (in mm) :Valve
R* L* L* L* R ** R** L **
64
63
55
40
Height 61
64
55 53 65
40
Lengthca.7 ca.4
Depth ca. 5 ca.5 ca.4*-cotype specimens,**-topotype specimens

Type locality :-Hodges Bay, AntiguaIsland, West Indies. Antigua Limestone.
Late Oligocene.
Distribution :-Known onLy from thetype locality.
This species has been reported fromthe Early Miocene of Venezuela by
HARRIS (1926, p. 107) but it has not been
. authenticated.
Amussiopecten preglyptus(OLSSON, 1922)
1922. Pecten (Aequipecten) preglyptus 0Lssoc-;, Bull. Amer. Paleont., Vol. 9, no.
39, p. 202, pl. 17, figs. 2,7.

Holotype :-Paleont. Res. Inst., Coil. No.
21140.
Remarks :-As this species was based
upon a poorly preserved left valve, its
detailed morphological features remained
unknown. It is characterized by its
rather small shell, slightly inflated left
valve, about 16 rounded, very low radialribs which tend to become obsolete towards ventral and lateral margins andnarrower than their interspaces inyounger stage but tend to become nearly
equal to their interspaces in breadthwith growth; fine, regularly spaced incremental lines; distinct auricular crura
terminating distally in an oblong denticle
at each extremity ; distinct paired internal
ribs near ventral margin ; and simple
cardinal crura.
This species was described under thesubgenus Aequipecten but from the above
mentioned morphological characteristics
it is evident that it can not be referredto that subgenus but to Amussiopecten.
Unfortunately the characteristics of theright valve remain unknown.
Comparison:- This species can be distinguished from A. vanvlecki by its
smaller shell, indistinct low radial ribs,
and smaller auricles.
Amussiopecten

.589.

A.mnssiopecten

preglyptus also is related to A. antiguensis, but it differs in the smaller apicalangle, the incremental lines not raised,and oblong denticles at each extremityof the hinge.
Dimensions (in mm) :-Height 50, length
50, depth ca. 5.
Type locality :-Upper Cocles Creek,
Limon Province, Costa Rica. Gatun Formation. Middle Miocene.
Distribution :-Known only from thetype locality.
Amussiopecten churuguarensis(F. and H. HODSON, 1927)
Pl. 26, figs. 6, 7
1927. Pecten antiguensis churuguarensis F. and
H. Hooso:'i, Bull. Amer. Paleont., Vol.
13, no. 49, p. 35, pl. 20, figs. 1, 3, 4,
pl. 22, fig. 2.

Holotype :-Paleont. Res. Inst., Coil. No.
21965.

Remar!zs :-The specimens studied en·able the presentation of the following
description: Shell rather small to medium
in size, thin, orbicular in outline, compressed, subequivalve but right valve
more inflated than left valve; valves·smooth but radiately ribbed and forming
an angle of about 110° at apex. Right
valve slightly inflated, with about 14
very low, flatly round-topped radial ribs
tending to become obsolete towards
ventral and lateral margins and regularly
spaced, fine incremental lines; radialribs broader than their interspaces ;:auricles extremely angulated, nearly
equal to each other, with wide andshallow byssal notch below anteriorauricle and with rugose, fine concentriclines. Left valve slightly inflated, withvery low, rounded radial ribs that arenearly equal to their interspaces inyounger stages, but tend to become.obsolete and broad towards ventral and213

lateral margins; and regularly spaced
raised incremental lines; auricles straight,
with distinct concentric lines. Hingewith simple cardinal crura and distinct
auricular crura. Interior surface withpaired internal ribs.
Although this species was originallydescribed as a subspecies of P. antiguensis, it is evident from the morphological characteristics above mentionedthat it should be raised to a species.
Comparison:-Amusium ocalanum(DALL) from the Eocene Ocala Limestone
in Florida (HARRIS, 1951) can be distinguished from the present one by its
oblique, thin shell, left valve little more
inflated than right valve, complicatedcardinal crura, faint radial ribs, rather
deep byssal notch and ctenolium. Amusium ocalanum is considered to be asignificant species in clarifying the relationship between Amusium and Amussiopecten. Need for a detailed study isevident.
This species can be distinguished fromother species of Amussiopecten by its
extremely angulate auricles.
Dimensions (in mm) :Valve
R* R
50
42
Height56
Length 45
6Depth ca.5*-holotype

R
35
38
4

R
L
28
45
45
30
3.5 5

L L
L
37
24 21
37
26 21
-ca. 2 2Type locality :-Districts of Petit, Miranda, Federacion and Buchivacoa, Stateof Falcon, Venezuela.· Churuguara Formation. Late Oligocene.
Distribution :-Churuguara Formation,
Venezuela; San Sebastian Formation,
Puerto Rico: Late Oligocene.
Amussiopecten harrisi
MASUDA, n. sp.
Pl. 26,

figs. la-b

214

Koichit·o MASUDA

Holotype :-U.S. Nat!. Mus., Col!. No.
646476.
Description :-Shell medium in size,
rather thin, compressed, subequivalve;right valve a little more inflated thanleft valve; valves radiately ribbed andforming an angle of about 1100 at apex.
Right valve slightly inflated, with about
16 low but rather distinct, perpendicularsided. flatly round-topped radial ribs
tending to become obsolete and broad
towards ventral and lateral margins,and faint, fine incremental lines; radialribs broader than their interspaces, which
are flatly round bottomed ; auricles somewhat angulate, with fine concentric lines
and faint, fine radial threads. Left valve
slightly inflated, with low, rather flatlyrounded radial ribs. which tend to become obsolete towards ventral and lateral
margins, crossed by regularly spaced,raised, faint, fine incremental lines;
radial ribs nearly equal to their interspaces in breadth at younger stage but
tending to become broader than theirinterspaces with growth; auricles
straight, with fine, faint radial threads
and concentric lines. Hinge with simple
cardinal crura provided with a distinct,
fine provinculum. wide and shallow
resilial pit, and distinct auricular crura
terminating distally in a distinct denticle
at each extremity. Interior surface with
distinct paired internal ribs near ventral
margin.
Dimensions (in mm) :-Holotype, height
57. thickness 12.5; paratype, right Yalve,height 28, length 29, depth, ca. 4; rightvalve, height 45, length 45.
Remarks:- This species is named inhonor of Late Prof. Gilbert D. HARRIS
who greatly contributed to the paleontological studies of Central and SouthAmerica and founded the PaleontologicalResearch Institution in Ithaca, New York.
Comparison :-Amussiopecten vanvlechi

can be distinguished from the present
new species by its larger shell, nearlyflat left valve in younger stage thattends to become slightly inflated withgrowth. the more distinct, high radialribs in right valve. and distinct radialribs narrower than their interspaces andraised, rugose. fine incremental lines in·
the left valve.
The present species also is related toA. antiguensis but it differs from the·latter in having distinct, perpendicularsided radial ribs in right valve and theleft valve with distinct, flatly rounded,broad radial ribs. It differs from A ..
preg!yptus in having left valve withflatly rounded radial ribs that are nearly
equal to their interspaces in younger
stage. auricles with radial threads anddistinct denticle at each extremity ofhinge.
Several poorly preserved specimensreferable to the present species are
known from an unnamed formation of
Late Oligocene in Mexico.
Type locality :-U.S. Geol. Surv .. Loc ..
No. 19780. San Sebastiar, quadrangle,
west of San Sebastian, Puerto Rico. San
Sebastian Formation. Late Oligocer..e.
Distribution :-San
SebastianFormation, Puerto Rico; Simojovel. Camino·
Carretero, Chiapas, Mexico (Univ. Calif.
Berkeley, Loc. No. B8267): Late Oligocene.

Amussiopecten woodringi

MASUDA, n. sp.
Pl. 26, figs. 2a~b, 3a~b
1926. Pecten antiguensis BRowr;, HARRIS,
johns Hopkins Univ., Stud. Geol., No.
7, p. 107, pl. 19, fig. 4 (non BROW:\',
1913).

Holotype :-U.S. Nat!. Mus .. Col!. No.
646474.

.589.

Arnussiopecten

Description :-Shell medium in size,
rather thin, compressed, orbicular inoutline. equilateral, subequivalve; rightvalve a little more inflated than left
valve; valves radiately ribbed andforming an angle of about 115° at apex.
Right valve slightly inflated, with about
21 very low, roundly flat-topped radialribs and fine incremental lines ; radialribs somewhat broader than their interspaces in younger stage but tend to be·come broader and obsolete towards
ventral and lateral margins; auricles
rather large, angulate, subequal to each
·Other, with somewhat raised, rugose.
fine concentric lines ; anterior auricle
with shallow and wide byssal notch.
Left valve slightly inflated, with very
low, rounded radial ribs tending to be·COme obsolete towards ventral and lateral
margins with growth and rather distinct, regularly spaced, fine incremental
lines ; radial ribs narrower than theirinterspaces in younger stage but tending
to become nearly equal to or a little
broader than their interspaces withgrowth ; auricles straight, with rugose,fine concentric lines and faint, fine radial
threads. Hinge with simple cardinal
crura provided with fine provinculum
and wide and shallow resilial pit; auricular crura distinct, terminate distallyin a distinct denticle at each extremity.
Interior surface with distinct paired
internal ribs near ventral margin.

and paleontological studies in theCentral America and South America.
Comparison :-Pecten antiguensis illustrated by HARRIS (1926) from the Ste.
Croix Formation in Trinidad can be referred to the present new species, because the type specimen (Paleont. Res.
Inst., Coli. No. 25058) has radial ribs
showing characteristics similar to thoseof the present species but it differs fromantiguensis in several particulars. Although HARRIS (1926) described thisspecimen as an Oligocene species, thelocality of this specimen at present is
considered to belong to the Early
Miocene Ste. Croix Formation.
This new species can be distinguishedfrom fl. anliguensis (BROW:t\) by its
smaller apical angle at the apex and thelarger number of very low radial ribs.
Also fl. harrisi MASUDA differs from thepresent one in having a smaller numberof perpendicular sided radial ribs in theright valve and distinctly broader radialribs in the left valve.
Type locality :-U.S. Geol. Surv .. Lac.
No. 19775. Central La Plata quadrangle,
northeast of Central La Plata, PuertoRico. Lares Limestone. Late Oligocene.
Distribution:- Lares Limestone. Puerto
Rico; Ste. Croix Formation, Trinidad:
Late Oligocene to Early Miocene.
.lmussiopecten sp.
Pl. 2:i, 1igs. 5a-c

Dimensions (in mm) :Valve
R-L
I~- L *
R
R
63
Height50
70
57
Length
52
70
63
Depth
10** ca. 11** ca.::J ca. 4*-holotype, **-thickness

215

L
<15
•!:i
ca.3

Remarlzs :-This species is named inhonor of Dr. Wendell P. WooDRING of
the Smithsonian Institution, who has
greatly contributed to the stratigraphicalThree rather poorly preserved specimens from the Early Miocene Tuxpan
Formation in Vera Cruz, Mexico are athand. They are in the collections ofSmithsonian Institution and in the
California Academy of Sciences. They
may characterized as follows: Shellrather large, medium in thickness, compressed, subequivalve; right valve a2Hi

Koichiro MASUDA

little more inflated than left valve;
valves radiately ribbed and forming anangle of about 115° at apex. Right
valve slightly inflated, with about 18
very low, perpendicular sided radialribs tending to become obsolete towards
ventral and lateral margins. and fineincremental lines; radial ribs broader
than their interspaces; auricles rather
large, somewhat angulate, subequal, withrugose. fine concentric lines. Left valve
slightly inflated, with flatly rounded,
low radial ribs and regularly spaced fineincremental lines ; radial ribs nearly
equal to their interspaces in breadth,tending to become obsolete towards
ventral and lateral margins ; auricles
straight, subequal, with rugose, fine concentric lines. Hinge with simple cardinal
crura and fine provinculum; auricularcrura distinct. Interior surface withpaired internal ribs.
Several fragmental specimens from the
Early Miocene Aguada Formation in
Puerto Rico (U.S. Geol. Surv., Loc. No.
19710) seem also to belong here, and areincluded with some hesitation.
These specimens apparently represent
an unnamed species, but naming iswithheld until better specimens arecollected.
From A. woodringi this form is distinguishable by its larger shell, more orless squarish and smaller number ofradial ribs in the right valve, and theleft valve with low radial ribs which
are nearly' equal to their interspaces inbreadth. A. harrisi also is related tothe present one but differs in having asmaller shell, rather distinct, squarish
radial ribs in the right valve, the left
valve with distinct radial ribs that arebroader than their interspaces, andauricles provided with radial threads
and concentric lines.
Dimensions (in mm) :-Height ca. 100,

Length 100, thickness 17 (U.S. Nat!.
Mus., Col!. No. 646478); length 100;height 42, length 45, thickness 10 (Calif.
Acad. Sci., Loc. No. 41035).
Locality:-" Hill A" in City of Tuxpan.
Vera Cruz, Mexico. Tuxpan Formation.
Early Miocene.
Distribution :-Tux pan Form at ion,
Mexico; Aguada Formation, Puerto Rico:
Early Miocene.
Paleontological SignificanceIn the present study the following
species referable to the genus Amussiopecten are recognized from the Late
Oligocene to Middle Miocene formationsin North America, Central America, West
Indies and northern South America ;Amussiopecten vanvlecki (ARNOLD). A.
antiguensis (BROWN), A. preglyptus (OLSSON), A. churuguarensis (F. and H.
HODSON), A. harrisi MASUDA, n. sp., A.
woodringi MASUDA, n. sp. and A. sp.
Judging from the descriptions andfigures the writer considers that thefollowing species and subspecies in
Europe and the Mediterranean Region
can be referred to the genus Amussiopecten.
Amussiopecten burdigalensis (LAMARCK.
1809) : Oligocene to Miocene
A. burdigalensis elongata (MYLIUS,
1891) : Miocene
A. burdigalensis spinosella (SACCO,
1897) : Miocene
A. burdigalensis minor (v. ROTH, 1914) :Oligocene
A. galloprovincialis (MA THERON, 1842) :MioceneA. jJasinni (lVIENEGHINI, 1857) : Miocene
A. koheni (FUCHS, 1876) : Miocene
A. vinassai (UGOLINI, 1899) : Miocene
A. ugolinii (DEPERET and ROMAN, 1912) :MioceneA. guebhardi (DEPERET and ROMAN,

589.

\

\

J

Amussiopecten

211

0

Text-fig. 2.

Distribution of Oligocene Amussiopecten.

1912) : Miocene
A. benoisti (COSSMAN and PEYROT,1914) : Miocene
The geological age of Amussiopectenin Europe and Mediterranean Region
ranges from the Late Oligocene to Middle
Miocene. However, in Europe there isso much confusion in the classificationof this group that a detailed study seems
necessary to clarify the biostratigraphicalrelations of Amussiopecten.
The following species are known
from the Late Oligocene to Early to
Middle Miocene formations in Iran, andEast and North Africa.
Amussiopecten burdigalensis (LAMARCK,1809) : Oligocene to Miocene
A. gregoryi (Cox, 1927) : Miocene
A. africanus (Cox, 1927) : Miocene
A. cf. labadyei (D'ARCHIAC and HAIME,
1854) : Oligocene (Cox, 1936)
A. sp., EAMES and Cox (1956) : Miocene
In Indonesia the following species
have been described from the Early
Miocene and Early Pliocene formations.
Amussiopecten singikirensis (~'IARTIK.

1909) : Miocene
A. gendinganensis (MARTIN, 1909) :Pliocene
From Japan and Taiwan the following
species are known from the Early Miocene
to Early Pliocene formations.
Amussiopecten praesignis (YOKOYAMA,
1922) : Pliocene
A. yabei (NOMURA, 1933) : Pliocene
A. iitomiensis (OTUKA, 1934) : Miocene
A. alziyamae MASUDA, 1962: Miocene
A. sp., CHANG (1967) : Miocene
From the species listed above it isevident that the geological range ofAmussiopecten is from the Late Oligocene
to Early Pliocene. The Oligocene species
are known from Europe (CSEPREGHYMEZNERICS, 1964, CTYROKY, 1969, etc.),
Iran (Cox, 1936), Puerto Rico and AntiguaIsland in West Indies, Venezuela and
Mexico, and also the Pliocene species
are known from Japan, Taiwan (MASUDA,
1962b) and Indonesia (MARTIN, 1909).
The other species are all known fromthe Early to Middle Miocene formations
in world wide distribution. That is toKoichiro MASUDA

218

Text-fig. 3.

Distribution of Miocene Amussiopecten.

say, it is evident that the Amussiopecten
group, except for a few species of East
Asia, became extinct at the end of
Middle Miocene time. The stratigraphi·Cal and geographical distributions of thegenus Amussiopecten is shown in Textfigs. 2-4. It is interesting to notice that{1) the genus Amussiopecten appeared in
the Late Oligocene and was dispersed
from Central Asia to Central America,(2) in the Early to Middle Miocene
Amussiopecten expanded its range toworld wide and many species appeared,and (3) in the Early Pliocene only a few
species lived in southern East Asia andthey became extinct at the end of Early
Pliocene.
All species of Amussiopecten in Europe
and America became extinct at the endof Middle Miocene, but in East Asia
three species survived to the end of
Early Pliocene. In general, the decreasein number of species with time can be
explained by the changes of oceanographic environmental conditions.
Thewater temperature generally became

gradually lower from the Late Oligocene
to the latest Neogene. Therefore, it canbe considered that as the result ofchanges in the oceanographic conditions
with time all species of Amussiopectenin Europe and America became extinct
at the end of Middle Miocene but inEast Asia three species were able tosurvive to the Early Pliocene. That is
to say, the environmental conditions inSouth East Asia did not change extremely
during the later Tertiary and have beenrather stable than those of the otherareas from the Early Miocene to Early
Pliocene.
From the accounts given above it appears that the distribution of Amussiopecten has been dependent upon thechanges of oceanographic conditions
progressively during its geological age.
Therefore, world wide occurrences ofAmussiopecten are considered to be verysignificant for interregional correlation.
The Late Oligocene to Middle Miocene
pectinid faunas in North America areusually composed of European elements

589.

Amussiopecten

219

(} ..

Text-fig· 4.

Distribution of Pliocene Amussiopecten.

such as Pecten, Flabellipecten, Amusium,

Amussiopecten, Argopecten, Lyropecten,
Aequipecten, etc., but the Late Miocene
to Pliocene pectinid fauna in the northernWest Coast of North America generally
contains a mixture of Asian elements
such as Swiftopecten, lv/izuhopecten, Yabepecten, Fortipecten and others (MASUDA
and ADDICOTT, 1970, MASUDA, 1971), thesurvivors of Miocene pectinids such as
Pecten, Argopecten, Lyropecten, Aequipecten and endemic genera such as
Patinopecten and Lituyapecten. But inthe southern West Coast of North
America, East Coast of North America,
and West Indies, the pectinid faunas
differ greatly from those in the northernWest Coast since the Late Miocene. In
the southern West Coast the Pliocene
pectinid fauna reveals a quite different:1spect from those of northern part:these faunal provinces indicate geographic differentiation. These facts can be
explained by changes of oceanographic
conditions as a result of paleogeographical changes in North and South America(OURHANI, 1961, WHITMOKE and STEWART,
1965, WoODRil'\G, 1966).
The North
American continent may have been connected with the South American continent
at least in the Late Miocene to Early
Pliocene.
In 1967 MACNEIL stated that most ofmolluscan stock in North America hasolder representatives in japan and theyin turn are immigrants from East Asia.
However, as far as now known, it isevident that the Amussiopecten groupand others such as Amusium, Argopecten.
Lyropecten, ilequipecten, etc. have moved
from the West Indies to North America.
Therefore, it can be considered that theolder stocks of Oligocene to Miocene
pectinids in North America should havebeen present in the Tethys Sea. Of
course, it is evident that Amu ssiopect enin North and South America did notcome from East Asia.
Furthermore, from the view point ofmorphological characteristics of Amussiopecten it can be stated that the older
the geological age, the smaller the shell.

220

Koichiro MASUDA

Thus, the latest species of Amussiopecten,
A. praesignis from the Early Pliocene ofJapan, has the largest shell (max. about
140 mm in length) ; but the Oligocene
species are always smaller (max. about
70 mm in length in A. woodringi), andmoreover, the Miocene species are
medium in size (max. 125 mm in lengthin A. vanvlecki). Therefore, it seems
quite possible and reasonable to determine the geological age of Amussiopecten
bearing formations and to correlate
geographically isolated geological formations based upon the morphologicalcharacteristics of this genus.
The life history of Amussiopecten is
unknown but judging from morphologicalcharacters such as auricular crura which
terminate distally in a very conspicuous,rounded, oblong denticle at each extremityof the hinge; the circular, equilateral,
compressed, rather polished, thin shell,and characteristic paired internal ribs,
it appears that Amussiopecten may havebeen adapted for active swimming habits.
Moreover, judging from the occurrencesof Amussiopecten,-they are usually foundin rather well sorted, very coarse to
medium-grained sandstone or impure
limestone-, it is considered that the life
environment of Amussiopecten may havebeen influenced by rather strong currents
on shallow sea bottoms.
References
AccoRD!, B., (1955): Stratigrafia e Palaeon·
tologia delle Formationi Oligo-Mioceniche del Trevigiano Orientale. Mem.
Jnst. Ceo/. Mineral., Univ. Padova, Vol.
19, pp. 3-64, tav. 1-5.
AKIYAMA, M., (1957): Amussiopecten iitomiensis (OTt:KA) and its allies from Japan.
Trans. Proc. Palaeont. Soc. japan, N.S.,
No. 25, pp. 31-39, pis. 6-7, 2 text-figs.
AR='IOLD, R., (1906) : Tertiary and Quaternary Pectens of California. U.S. Ceo/.

Surv. Prof. Paper 47, pp. 7-146, pis. 153, 1 text-fig.
- - , (1907) : New and characteristic species.
of fossil mollusks from the oil-bearing
Tertiary formations of Santa BarbaraCounty, California. Smiths. Misc. Coli.,Vol. 50, no. 4, pp. 419-447, pis. 50-58.
- - , and ANDERSON, R., (1907) : Geology
and oil resources of the Santa Maria
district, California. U.S. Ceo/. Surv.,
Bull., 322, pp. 1-161., pis. 12-16.
BREMNER, C.S.J., (1932): Geology of Santa
Cruz Island, Santa Barbara County,
California. Santa Barbara Mus. NatHis/., Occ. Papers, No. 1, pp. 5-33, pis.
1-4.
BROWN, A.P., (1913) : Notes on the geologyof the Island of Antigua. Proc. Acad.
Nat. Sci. Philadelphia, pp. 584-616, pls_
18-20.
- - , and PILSBRY, H.A., (1911) : Fauna of
the Gatun Formation, Isthmus of Panama_
Ibid., Vol. 63, pp. 336-374, pls. 22-29, 3
text-figs.
CHANG, L., (1967) : Tertiary Biostratigraphyof Taiwan and its Correlation. Tertiary
Correlation and Climatic Changes in the
Pacific. 11th Pacific Sci. Congr., Tokyo,
1966, pp. 57-65, 1 text-fig. 1 table.
CooKE, C.W., (1919): Tertiary mollusks
from the Leeward Islands and Cuba.
Carnegie Inst. Washington Pub!., 291, pp.
103-156, pis. 1-16.
- - , GARDNER, J., and WooDRI:'\G, W.P.,(1943) : Correlation of the Cenozoic formations of the Atlantic and Gulf Coastal
Plain and the Caribbean region. Ceo[.
Soc. Amer., Bull., Vol. 54, pp. 1713-1722,
1 pl.
CossMAK, M., and PEYROT, A., (1914) : Conchologie Neogenique de I•Aquitaine.
Tome II, Pelecypodes. Act. Soc. Linn.
Bordeaux, pp. 1-496, pl. 1-26.
Cox, L.R., (1927) : Neogene and Quaternary
Mollusca from the Zanzibar Protectorate,
in DAVIES, A.M., Cox, L.R., STOCKLEY,
G.M., STUBBLEFIELD, C.J., and WHITE,
E.I., Report on the Palaeontology of theZanzibar Protectorate. pp. 13-102, pls. 3-9.
- - , (1929) : Notes on the Post-Miocene

589.

A1nussiopecten

Ostreidae and Pectinidae of the Red Sea
region, with remarks on the geologicalsignificance of their distribution. Proc.
Malacol. Soc. London, Vol. 18, pp. 165-209,
pis. 11-13.
- - , (1936) : Fossil Mollusca from Southern
Persia (Iran) and Bahrein Island. Mem.
Geol. Surv. India, Palaeont. Indica, N.S.,
Vol. 22, no. 2, pp. 1-69, pis. 1-8.
- - , (1939) : Miocene, Pliocene and postPliocene Mollusca from Mozambique.
Dol. Serv. Indus!. Mineral. Geol. Mozambique, No.3, pp. 21-106, pl. 1-2, 1 text-fig.
- - , (1942) : Neogene aspect of Vaqueros
mollusks, in ScHEI\CK, H.G., and CHILDS,
T.S., Jr., Significance of Lepidocyclina(Lepidocyclina) californica, new species,in the Vaqueros Formation (Tertiary),
California. Stanford Univ. Publ., Geol.
Sci., Vol. 3, no. 2, pp. 34-35.
CsEPREGHY-MEZ:'\ERICs, J., (1960) : Pecti·
nides du Neogene de Ia Hongrie et leurImportance Biostratigraphique.
Mem.
Soc. Geol. France, N.S., 92, pp. 5-56, pis.
1-35, 1 text-fig., 3 tables.
- - , (1964) : L'Evolution de certains Pectinides Neogenes, Ia question du "Chattien"et Ia limite Oligo-Miocene. Inst. Lucas
Mallada, C.S. I.C. (Espana). Cursillos Y
Conferencias, IX, pp. 33-50.
CTYROKY, P., (1969) : The Family Pectinidae
in the Burdigalian of Czechoslovakia.
Sbor. Geol. Ved. Paleont., 10, pp. 7-66,
pis. 1-20, 12 text-figs.
DALL, W.H., (1898) : Contl"ibutions to the
Tertiary Fauna of Florida with specialreference to the Silex Beds of Tampa
and the Pliocene beds of the Caloosahatchie River. Trans. Wagne1· Free Inst.
Sci. Philadelphia, Vol. 3, pt. 4, pp. 571947, pis. 26-37.
DEPERET, Ch., and Ro~IA:'\, F., (1910): Monographie des Pectinides neogenes de!'Europe et des regions voisines. II. Genre
Flabellipecten. lVJem. Soc. Geol. France,
Paleontologie. Tom. 18, fasc. 2, pp. 105139, pis. 8-13, text-figs. 44-60.
- - and - - , (1912) : Monographies des
Pectinides neogenes de !•Europe et desregions voisines. II. Genre Flabellipecten.

221

Ibid., Tom. 19, fasc. 1, pp. 139-168, pis.
18-23, text-figs. 61-71.
- - and - - , (1928) : Monographie desPectinides neogenes de !'Europe et desregions voisines. II. Genre Flabellipecten.
Ibid., N.S., Tom. 4, fasc. 4, pp. 169-194,.
pis. 26-28, text-figs. 1-10.
DouGLAs, J.A., (1928) : Contributions toPersian Palaeontology III. pp. 1-19, pis.
8-15.
Dt:RHA~I,
J.W., (1961) :Paleogeographic
conclusions in light of biological Data.
Pacific Basin Biogeography. 1Oth PacificSci. Congr., pp. 355-365, 4 text-figs.
EA~IES, F.E., and Cox, L.R., (1956) : Some
Tertiary Pectinidae from East Africa,
Persia and the Mediterranean Region.
Proc. Malacol. Soc. London, Vol. 32, pp.
1-68, pis. 1-20.
ELDRIDGE, G.H., and ARNOLD, R., (1907) :The Santa Clara Valley, Puente Hills,
and Los Angeles Oil Districts, Southern
California. U.S. Geol. Surv., Bull., 309,
pp. 1-266, pis. 1-41, 17 text-figs.
GRANT, U.S., IV, and GALE, H.R., (1931) :Catalogue of the marine Pliocene andPleistocene Mollusca of California. Mem.
San Diego Soc. Nat. Hist., 1, pp. 1-1036,
pis. 1-32, text-figs. l-Ei.
HARRIS, G.D., (1926) : Notes on the Paleontology, in vVARING, G.A., The Geologyof the Island of Trinidad, B.W.I. johns
Hopkins Univ. Stud. Geol., No. 7, pp. 87112, pis. 16-20.
- - , (1951) : Preliminary Notes on Ocala
Bivalves. Bull. Amer. Paleont., Vol. 33,
no. 138, pp. 223-272, pis. 1-13.
HERTLEIN, L.G., (1925) : New species ofmarine Mollusca from western North
America. Bull. South. Calif. Acad. Sci.,
Vol. 24, pt. 2, pp. 39-46, pis. 3-4.
- - , (1969) : Family Pectinidae, in Treatiseon Invertebrate Paleontology.
(N)
Mollusca 6 (1 of 3), Geol. Soc. Amer.,
Univ. Kansas, N348-373, figs. C72-C94.
HoDSON, F., HoDsON, H.K., and HARRis,
G.D., (1927) : Some Venezuelan and
Caribbean mollusks. Bull. Amer. Paleont.,
Vol. 13, no. 49, pp. 1-160, pis. 1-40.
LoEL, W., and CoRr::Y, W.H., (1932) : The

222

Koichiro MASUDA

Vaqueros Formation, Lower Miocene of
California. I. Paleontology. Univ. Calif.
Pub!., Bull. Dept. Geol. Sci., Vol. 22, no.
3, pp. 31-410, pis. 4-65, 2 maps.
MACNEIL, F.S., (1961) : Lituyapecten (NewSubgenus of Patinopecten) from Alaska
and California. U.S. Geol. Surv., Prof.
Paper 354-J, pp. 225-239, pis. 35-46.
- - , (1967) : Cenozoic pectinids of Alaska,
Lceland, and other northern regions.
Ibid., 553, pp. 1-57, pis. 1-25.
MA"SFIELD, W.C., (1936) : Stratigraphic
Significance of Miocene, Pliocene andPleistocene Pectinidae in the Southeastern
United States. jour. Paleont., Vol. 10,
no. 3, pp. 168-192, pis. 22-23, 1 text-fig., 1table.
MARTIN, K., (1909) : Die Fossilen von Java.
Lamellibranchiata. Samml. Geol. ReichsMus. Leiden. N.F., 1, pp. 333-386, pis.
46-54.
MASUDA, K., (1962a) : Tertiary Pectinidae
of Japan. Sci. Rep., Tohoku Univ., Ser.
2 (Ceo!.), Vol. 33, no. 2, pp. 117-238,
pls. 18-27, text-figs. 1-11..
- - , (1962b) : Notes on the Tertiary Pectinidae of Japan. Ibid., Special Vol. No.

5, pp. 139-193, 9 text-figs., 9 tables.
- - , (1971) : On some Patinopecten from
North America. Trans. Proc. Palaeont.
Soc. japan, N.S., No. 83, pp. 166-178, pis.
19-21, 2 text-figs.
- - , and ADDICOTT, W.O., (1970) : On
Pecten (Amusium) condoni HERTLEIN
from the West Coast of North America.
The Velige1·, Vol. 13, no. 2, pp. 153-156,
1 pl.
No\Il"RA, S., (1933) : Catalogue of the
Tertiary and Quaternary l\'Ioll usca of
Taiwan. Sci. Rep., Tohoku Imp. Univ.,
Ser. 2 (Ceo!.), Vol. 16, pp. 1-108, pis. 1-4.
OLssoN, A.A., (1922) : The Miocene ofNorthern Costa Rica with notes on its
general stratigraphic relations.
Bull.
Amer, Paleont., Vol. 9, no. 39, pp. 1-288,
pls. 1-32.
OTUKA, Y., (1934) : On some fossils fromthe northern foot of Mount Minobu andfrom the Hayakawa Tuffite of Hakone.
Geol. Soc. japan, jour., Vol. 41, no. 492.
pp. 562-368, 1 text-fig.
PIIILIPPI, E., (1900) : Beitrage zur Morphologie unci Phylogenie der Lamellibranchier, II. Zur Stammesgeschichte derExplanation of Plate 25(Natural size)
Figs. la-b, 2, 3, 4. Amussiopecten vanvlecki (ARNOLD).
1a, right valve. 1b, left valve. U.S. Nat!. Mus., Coli. No. 646495. Loc. :-U.S. Geol.
Surv., Loc. No. M2418; about 1.5 miles north of Santa Barbara Canyon Ranch, Fox
Mountain Quadrangle, Santa Barbara County, California. Saltos Shale Member ofMonterey Shale. Middle Miocene. 2, hinge area of right valve. U.S. Nat!. Mus., Coli.
~o. 646498.
Loc. :-U.S. Geol. Surv., Loc. No. M2417; about 1..3 miles southwest of theDrill hole near Big Pine Road in Santa Barbara Canyon, Fox Mountain Quadrangle,
Santa Barbara County, California. Branch Canyon Formation. Middle Miocene. 3, left
valve. U.S. Nat!. Mus., Coli. No. 646496. Loc. :-U.S. Geol. Surv., Loc. No. M2458;
about 1.4 miles southwest of Willow Campground in Santa Barbara Canyon, Fox Mountain
Quadrangle, Santa Barbara County, California. Branch Canyon Formation. Middle
Miocene. 4, plaster cast of right valve. U.S. Nat!. Mus., Coli. No. 646497. Loc. :U.S. Geol. Surv., Loc. No. M3044; about 0.8 miles west of the Spring in Alamo Canyon,
Fox Mountain Quadrangle, Santa Barbara County, California. Branch Canyon Formation.
Middle Miocene.
Figs. 5a-c. Amussiopecten sp. a, right valve. b, left valve. c, upper view of 5a. Loc. :Calif. Acad. Sci., Loc. No. 41035; Tuxpan, Vera Cruz, Mexico. Tuxpan Formation.
Early Miocene.

MASUD A ·Amussiopecten
Plate 25

589.

Amussiopecten

Pectiniden.
Zeitsch. Deutsch.
Ceo!.
Cese!!., Bd. 52, pp. 64-117, 24 text-figs.
RoGER, ]., (1939): Le Genre Chlamys dans
Ies Formations Neogene de L'Europe.
Mem. Soc. Ceo!. France, pp. 5-294, pis.
1-35, 113 text-fig.
SAcco, D.F., (1897) : I Molluschi dei Terreni
Terziarii del Piemonte e della Liguria.
Part XXIV (Pectinidae). Torino, pp. 3116, pis. 1-21.
SCHAFFER, F.X., (1.910) : Das Miocan vonEggenberg.
I. Die Fauna.
Abhandl.
d.k.k. Ceo!. Reichsanstalt, 22, Bd. I. pp.
1.-126, taf. 1-48, 12 text-tigs.
SHuTo, T., (1955) : Amussiopecten from theMiyazaki Group, Miyazaki Prefecture,
Japan (Paleontological study of the
Tertiary Miyazaki Group). Trans. Proc.
Palaeont. Soc. japan, N.S., No. 20, pp.
101-110, pis. 16-17, 4 text-figs.
TELEGD·ROTH, K., (1914) : Eine Oberoligoziine Fauna aus Ungarn. Ceo!. Hungarica,Tom. 1, fasc. 1, pp. 5-77, pis. 1-6.
ToULA, F., (1908) : Eine jungtertiiire Fauna
von Gatun am Panama Kana!. jahrb.
Ceo!. Reichsanstalt, Bd. 58, pp. 673-760,
pis. 25-28, 15 text-figs.
VON TEPPNER, W., (1922) :Fossilium
Catalogus. I: Animalia. Lamellibranchiata tertiaria.
"Anisomyaria ". II.
pp. 77-296.
- - and DREGER, ]., (1918) : Neue Amussiopecten aus steirischen Tertiiirablagerungen.
Nebst e1mgen geologischen
Daten. jahrb. k.k. Ceo!. Reichsanstalt,
Bd. 67, pp. 481-502.

223

UGOLINI, R., (1907): Monografta dei Pettinidi
Neogenici della Sardegna. Parte Sconda.
Generi: Amussium, Amussiopecten. Palaeontgr. Italica, Vol. 13, pp. 233-242, 1 pl.
- - , (1908) : Monografia dei Pettinidi Neogenici della Sardegna. Part Terza ed
Ultima. Generi: Amussiopecten (cont.),
Flabe!lipecten, Pecten. Ibid., Vol. 14, pp.
191-224, pis. 22-25.
VEDDEil, ].G., (1968): Geologic map of Fox
Mountain Quadrangle, Santa BarbaraCounty, California. U.S. Ceo!. Surv.
VEIGA, 0., (1954) : Pectinideos do Miocenico
do Vale do Sado e da Serra da Arra.bida,
Comun. Serv. Ceo!. Portugal, Tom. 35,
pp. 155-186, 8 pis.
WEAVE!<, C.E., et a!. (1944): Correlation of
the marine Cenozoic formations ofwestern North America.
Ceo!. Soc.
Amer., Bull., Vol. 55, no. 5, pp. 569-598,
1 chart.
WHITMORE, F.C., Jr., and STE\YART, R.H.,(1965) : Miocene mammals and Central
American seaways. Sci., Vol. 148, pp.
180-185, 2 text-figs.
WooDRI"<G, W.P., (1966) : The Panama landbridge as a sea barrier. Amer. PM!os.
Soc., Proc., Vol. 110, no. 6, pp. 42S-433, 3
text-figs., 6 tables.
YoKOYAMA, M., (1922): On a new speciesof Pecten from the Neogene of Japan.
jour. Ceo!. Soc. Tokyo, Vol. 29, no. 530,
pp. 1- 2, pl. 5.
- - , (1928) : Mollusca from the Oil-Field of
the Island of Taiwan. Imp. Ceo!. Sun·.
japan, Rept., 101, pp. 1-107, pis. 1-18.

224

Koichiro MASUDA

Explanation of Plate 26(Nat ural size)
Figs. la-b. Amussiopecten harrisi MASUDA, n. sp. Holotype, U.S. Nat!. Mus., Coli. No. 646476.
a, right valve. b, left valve. Loc. :-U.S. Geol. Surv., Loc. No. 19780, San Sebastian
Quadrangle, Puerto Rico. San Sebastian Formation. Late Oligocene.
Figs. 2a-b, 3a-b. Amussiopecten woodringi MASUDA, n. sp. 2a, right valve. 2b, left valve.
Holotype, U.S. Nat!. Mus., Coil. No. 646474. 3a, right valve. 3b, left valve. Paratype,
U.S. Nat!. Mus., No. 646475. Loc. :-U.S. Geol. Surv., Loc. No. 19775, Central La Plata,
Puerto Rico. Lares Limestone. Late Oligocene.
Figs. 4, 5. Amussiopecten antiguensis (BROWN). 4, right valve. 5, left valve. Topotype,
U.S. Nat!. Mus., Col!. No. 167136. Loc. :-Hodges Bay, Antigua Island, West Indies.
Antigua Limestone. Late Oligocene.
Figs. 6, 7. Amussiopecten churuguarensis (F. and H. HoDsoN). Right valve, U.S. Nat!. Mus.,
Col!. No. 646477. Loc. :-U.S. Geol. Surv., Loc. No. 17253, Maca Quadrangle, PuertoRico. San Sebastian Formation. Late Oligocene.

MASUDA:

Amussiopecten

Plate 26

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 84, pp. 225-242, pis. 27, 28, December 30, 1871

590.

UPPER CRETACEOUS GLYCYMERIDS IN JAPANMASA YUKI TASHIRO
Shokey Highschool in Kumamoto

FJ:;$:0)l:~4lSilli*ii:?' !I-'\'-) !I 7-.IC"'?It''L:;:;h,i;"CICilcif;X~h'"C ~t.::;$:'f!lE3:!lE*i.lEO)

Glycymeris ~*Mf51"~~a~:i'l.!hjJ: IJ ~~ftiL., 4 Illi~ 9 f.]~~itl))l) Lt.:o Hanaia !lli~l"lr~~
S!lli*-J:iflls!lli*IC !j;f~~ "C<i!.> '.?, -'tO)i.!EiflJFviftl: f.t 1.: '"C 5ErfiJ~I~tJ: ~~~{l::.il~ '1!fL. <,-'tO)~{I::.v"lJ:~BS!lli*DO)fi]O)JlX::R;iti!!f!iHC.i>lt''"C~;ffl.~ ;/1,-Q o S":;$:1C.illt .Q Glycymerita Iffi~f"l, J:~B S!lli*'CifliJIIil!:~f~IC lfllJI. L. '"C v' .Q o l'i~:;$:}~0)lfll!ffii\Uf.i.!EO) Pseudoveletuceta, nov.
l"l, Hanaia J: IJ 5J"ip;h,f.:!f;ff,f.{a~tJ: glycymerid "C <!!.> .Qo l'i~":;$:~~0)1!~iill~ilf. tili~YiUil~F.fjlfl})j:O) Glycymeris amakusensis l"l, Glycymeris (s. str.) 0) Veletuceta group fcm; L
1J~::::_* 11EO)f)J!tc~W,i"" .Q o .i>.:C G <;ffl.J:i;!Hfi~*cot.: Veletuceta group O)~i/Jlla~tJ:~~O)-"? c ,ljil,:b;/1, .Q
133 ft .IE ;:t0

Introduction and ackowledgments
The family Glycymerididae is a wellknown marine bivalve family which·commonly occurs from the Lower Cretaceous to Recent. The Cretaceous·species have been described by many:authors from the several localities injapan: Y ABE and NAGAO (1928) and
NAGAO (1932) in Hokkaido, NAGAO (1934):and SAITO (1962) in Honshu, NAGAO(1930) and AMANO (1956, 1957) in Kyushu
and ICHIKAWA and MAEDA (1958) in
Shikoku. Recently HAY AMI (1965) established a subgenus Hanaia based upon
Glycymeris densilineata NAGAO and added
Glycymeris (Glycymerita ?) haipensis as a
new species from the Miyako group.
As I discovered many interesting facts.Qn morphological changes of glycymerids
through ages during my Upper Cretaceous fossil hunting in Kyushu, I
iurther visited several localities of Upper
Cretaceous glycymerids in Japan. I also
Received October 7, 1970; read November
29, 1969 at Kagoshima.

observed some glycymerids from Miyako,
Hokkaido and California at the Department of Geology of the Kyushu University (GKH). As a result of this study
the Upper Cretaceous Glycymeris in Japanis classified into four subgenera, Glycymeris (s. str.), Glycymerita FINLAY andMARWICK, Hanaia HAY AMI and Pseudoveletuceta. new subgenus. Hanaia is theoldest subgenus in Glycymeris and theother three subgenera may be derivativesof Hanaia. Glycymerita was prosperous
in the late Cretaceous in Japan. JapaneseUpper Cretaceous glycymerids are listed
below:
Glycymeris (Glycymeris) amalzusensis
NAGAO
Glycymeris (Glycymerita) japonica TASHIRO, new species
Glycymeris (Glycymerita) himenourensis
TASHIRO, new species
Glycymeris (Glycymerita) multicostata
NAGAO
Glycymeris (Hanaia) salida NAGAO
Glycymeris (Hanaia) matsumotoi TASHIRO, new species

225

Masayuki TASHIRO

226

Glycymeris (Hanaia) lwtsurazawensis
TASHIRO, new species
Glycymeris(Hanaia)hokkaidoensis(YABE and NAGAO)
Glycymeris (Pseudoveletuceta) mifunensis TASHIRO, new subgenus, newspecies
Limopsis kogata (ICHIKAWA and MAEDA)

Before proceeding further I wish to
express my sincere thanks to Emeritus
Prof. T. KOBA Y ASH! of the Universityof Tokyo for his reading the manuscript
and for criticism. I am also much indebted to Dr. M. TAMURA of the Kumamoto University for his kind guidance
and reading the manuscript. I thankProf. T. MATSUMOTO and Dr. I. HAYAMI of the Kyushu University for theircontinuing encouragement and supplyof glycymerid specimens. I am also
grateful to Prof. M. AMANO of the
Kumamoto University, Prof. K. ICHIKAWAof the Osaka City University, Dr. K.
MASUDA of the Miyagi University of
Education and Dr. I. OBATA of the
National Science Museum of Tokyo for
their kind help to my study.
The specimens here treated as KE
are kept in the Faculty of Education,
Kumamoto University.
Systematic descriptions
Family Glycymerididae NEWTON, 1922
Subfamily Glycymeridinae
NEWTON, 1922
Genus Glycymeris DA COSTA, 1778
Type-species :-Areaorbicularis DA
COSTA, 1778, (Area glycynzeris LINNE,
1758).

Subgenus Glycymeris DA COSTA, 1778
Several reports have been issued on
taxonomy of the Upper Cretaceous

glycymerids (FRE!\EIX, 1959; NICOL. 1945,.
1950; HAY AMI, 1965). Recently NEWELL(1969) in Treatise on Invertebrate Paleontology summarized above mentioned
works except for FRENEIX's. I agreewith NEWELL's general classification ofglycymerids. It seems. however, necessary for me to study further on the
Cretaceous glycymerid taxonomy. Veletuceta IREDALE is treated as a synonymof Glycymeris by NEWELL (1969). Although the discrimination between
V eletuceta and Glycymeris is difficult.
the Veletuceta group is distinct for its.
shape and prosperous in Upper Cretaceous and Palaeogene ages. The umbo of
Veletuceta is prosodetic and its beak is.
prosogyrate or orthogyrate. The umboof Glycymeris, on the other hand, is.
orthodetic or opisthodetic and its beakis opisthogyrate.
For the reasons.
mentioned above, I wish to leave Veletuceta as a valid group, if not as a subgenus. I have an opinion that recent
Glycymeris may be a derivative of theCretaceous or Tertiary Veletuceta.
Glycymeris (Glycymeris)amakusensis NAGAO
Plate 27, figs. 1-16, Text-figs. 1, 2
1930. Glycymeris amakusensis NAGAO, ]OUI".
Fa c. Sci., Hokkaido Imp. Univ., Ser. 4,
Vol. 1, No. 1, p. 15, pl. 2, figs. 4-7.
1957. Glycymeris sp., AMAKO, Kumamoto jour.
Sci., Sect. 1, Vol. 2, No. 2 p. 55, pl. 1,.
fig. 22.
1962. Glycymeris multicostata, SAITO, Bull.
Fac. Arts and Sci., Ibaraki Univ., Nat.
Sci., No. 13, pp. 62, 63, pl. 1, figs. 2, 3 ..

Observation and Remarks:- The specific name of amakusensis has been used
for the Upper Cretaceous glycymerids ofJapan. I collected about 300 specimens.
from many localities of the Himenouragroup in Kyushu and the Futaba group·590.

C1·etaceous Glycynw 1·ids

in F ukushima Pref. to confirm its shape.
The result of measurement is shown in
Tex t-fig. 1. The number of ve ntral
crenules (see Text-fig. 1, abbrev. C) is
about 25. The surface is ornamented
with numerous fiat-topped radial ribs,
each of wh ich is a bundle of 7 or 8
threads . The interspace between ever y
two ribs is very narrow. Generally f ully
matured specimens exceeding abo ut 20
mm. in len g th and the specimens fromthe sandstone at Locs. 14, 15 and 16 a r e

227

thick in test of shell. The radial ribsin these specimens are more elevatedthan in thin-shelled or yo unger specimens. Consequently growth lin es makesuch crenu lated appearance as describedby NAGAO (1930). In mature stage t wo
fairl y distinct patterns of shape areobservable as shown in Text-fig. 2. As
the intermediate fo rms are present,
these two g roups can not be distinguishedspecifically. UEDA and FURUKAWA (1959)
discriminated 3 variet ies of the species

H
mm

3 0

T
2 0

.

2 0

mm
Loc • 13

+ Loc. 11.
.

Loc • 140Loc.

x Loc.

12

Loc • 15

3

l 0

1 0

l 0

2 0

3 0

mm

L

Te xt -fig . 1. Scat t er diagrams of Hei g ht aga in st Length, and Th ickness against
Length of Gly cy meri s (s . st r. ) amallllsensis NAGAO (KE1799- KE1853) .
H: He ig ht, L: Length, T: Th ick ne ss, C: Nu mbe r of ve n t r a l crenules,© : Glycy meri s ( Glycymerila) him enou rensis T AS I-IIRO, n. sp . from Loc.
13 (for comparison). Positions of measur eme nt s a r e a lso s hown .

228

Nfasay~~ki

TASHIRO

10m m

c

Text-fig. 2. Glycymeris ( G!y cymeris) amakusensis N AGAO, A: Subt rian gu larform, B: Ligamenta! v iew of adult specimen, C: Subquad ra te form.

with no description. The umbon a l andmedian part of an immature valve isdark and othe r part encircling the above
mentioned pa rt is yello\:vish co lored.
T his dark part fades a wa y in mature
fo rm. Some specimens (Pl. 27, fig. 5)
show concentric color pattern as observable in some individuals of therecent species.
Occunence :- Locs. 1, 11- 16.
S ubgenus Glycymerita FINLAY
and MARWICK, 1937
TyjJe- sjJecies :-Gly cymeri s concava
MARSHALL, 1917.
This subgenus is chracterized asfollows: 1) The shell is subquadrate andwell inflated ; 2) the umbo is big andprominent; 3) both sides of the umbo
are strongl y depressed and rem arkab ly
shouldered; 4) the hinge plate is longand stout ; 5) the teeth on both lateralsides are long and extend horizontally;6) the radial ribs on the external surface
are strong and the radial threads are
weak or invisible.

appro x imatel y eq ual to each other,posterior marg in slig htl y s inuous, obliquely tr uncated, ventra l margin broadly
arched and gradua ll y changing into
anterior margin; umbo large, prominent,
and located almost medially ; beakorthog yrous, incurved a nd situateds lightly posterior to middle of dorsal
margin; both sides of umbo remarkably
shouldered and strongl y depressed;
surface ornamented with stout and flattopped radial ribs; ligame nt area narrow,triangular, provided with several chevrons ; hinge plate slightl y arched , very
long about four-fifths of length; about
24 teeth symmetricall y disposed , longand subhorizontal on both lateral sides,
short a nd diagonal below beak; inner
margin with crenules about 20 irr number.

Glycyme1-is (Glycymerita) japonica

TASHIRO, new species
Plate 27, figs. 17-22, Text-fig. 3

Description :- Shell small, well inflated,
subquadrate in outline, height and length

Te xt -fig. 3. G!ycy m eris (Glycymerita)japonica T ASH IRO, new species.

.590.

Cretaceous Glycymerids

229

Jvfeasurements (in mm.) :G. (Glycymerita) japonica
Specimen
LengthKE 1786 Right internal mould
27.0
KE 1787 Left internal mould
22.0
KE 1788 Left internal mould
22. 5
KE 1789 Left internal mould
21. 0
KE 1790 Right internal mould
25.5

Observation and Remarks :-In thisspecies the ratio of thickness/length ofshell is about 0.8. The crenulation of
the inner margin is frequently effacednear both lateral sides. The threads on
the radial ribs are almost invisible orvery delicate. The width of flat-topped
radial ribs is nearly equal to the widthof the inters paces in ventral side of adultspecimens. The width of interspaces in
the umbonal part of immature specimens,
however, is very narrow.
Comparison :-This species is similar
to the typical Clycymerita in generalfeatures except for its small size andnarrow ligament area. Clycymeris (Clycymerita) veatchii (GABB) from the Chicogroup in California (Pl. 27, Fig. 33) ismore inflated than this species and theradial ribs of veatchii are stronger thanthose in the present species. C. apletos
DAlLA Y and POPENOE (1964) from the
Jalama formation in California is somewhat akin to this species in its subquadrate and well inflated shell. C.
ap!etos, however, differs from thisspecies in the large shell and conspicuously incurved beak. C. apletos may be.a member of subgenus Clycymerita.
Occurrence :- Locs. 17 and 18.

Height

25.0
22.2
22. 4
18.5
25.5

Holotype
Para type
Ditto
Ditto
Ditto

Clycymeris (Clycymerita) himenouren sis

TASHIRO, new species
Plate 27, figs. 23-27, Text-fig. 4

Description :-Shell very small for
Clycymerita, orbicularly subquadrate,

higher than long, anterior and posterior
margins
obliquely
truncatedandshouldered at junctions with antero- andpostero-dorsal margins; ventral marginrounded; both sides of umbo depressed;
umbo moderately large, prominent,
located almost at mid-length; surface
ornamented with numerous and flattopped radial ribs which are broader
than their interspaces; ligament area
triangular,
narrow,
providedwithseveral chevrons; hinge plate long,moderately arched ; teeth symmetricallydisposed, short and diagonally bifurcated below beak, long and subhorizontal
in both sides of hinge plate; inner
ventral margin densely crenurated (about
30).
Obserualion and Remarlls :-The umbonal part of this species is black andshows the similar color pattern observedin the immature stage of Clycymeris (s.

1\feasurements (in mm.) :G. (Glycymerita) himenourensis

Specimen
KE 1779 Left valve
KE 1780 Left valve
KE 1781 Left valve
KE 1782 Right valve.KE 1784 Left valve

LengthHeight

13.0
12.3
7.0
10. 7
13.8

14.2
17.5
7. 7
11.2
15.2

Thickness

4.0
6.0
2.0

7.5
7.8

Holotype
Para type
Ditto.
Ditto .
Ditto.

Masayuki T ASH! RO

230

B

Locality Maps

D

0

2 km

D

X

9

0
SHiSHI -ISLAND

X X7

8

20km

590.

Cretaceous Glycyme1·ids

Table 1.
Mikasa sandstone
--

Loc.

------~----

1

Turon.

s.s.

About 500 m south of the Katsurazawa-dam,
Ikushunbetsu, Hokkaido.
The coast of the waterway near the Katsurazawa-dam, Ditto.

Upper Yezo Group2

Santon.

B
s.s.

The Sakainosawa river, Kyowa, Teshio prov.,
Hokkaido.
The Gakkonosawa river, Nakagawa-machi,
Teshio prov ., Ditto.

Loc. 20**c

Futaba Group
Loc.

3

Coniac.

silt s.

Goshonoura GroupThe Sakuradani river, Hirano-machi, Futaba Co.
Fukushima Pref.
D

s.s.

Loc.

4

Up. Alb.

Loc.

5

L.L. Cenoman. s.s.

Loc.

6

Enokuchi, Goshonoura-machi, Amakusa Co.
Kumamoto Pref.
Karakizaki, Ditto.
Shirahama, the Shishi-islancl, Izumi Co.
Kagoshima Pref.

Miyaji formation
Loc.

7

Up. Alb.?

Loc.

8

*

D

Turon.

9Tani, Miyaji-machi, Yatsushiro city, Kumamoto
Pref.

s.s.

Naraki, Koda-machi, Ditto.

Mifune Group
Loc.

D

Asanovabu, Mifune-machi, Kamimashiki Co.
Kumamoto Pref.

sh.

Nishiyama, Katashida-machi, Shimomashiki Co.

Loc. 10__1

Himenoura Group
Loc. 11

Fossil localities.

A

Loc- 19*

Loc.

231

Santon.

D
Okoshiki, Oda-machi, Uto city, Komamoto Pref.

sh.

Loc. 12

Wadanohana, Takado-machi, Amakusa Co., Ditto.

Loc. 13

The Kugu-island, Takaclo-machi, Ditto.
s.s.

Loc. 14

Karakizaki, Goshonoura-machi, Ditto.

Loc. 15The Mayu-island, Ditto.

Loc. 16

Oe, Amakusa-machi, Ditto.

Loc. 17
Loc. 18

Hongo, Kawaura-machi, Ditto.

Cam pan.
I

Kamihira, Ditto.

*Loc. 19: IK 981 of Kyushu Univ.

**Loc. 20: by HA.Yc\\11 of Kyushu Uni\-.

Masny~iki T ASHIRO

232

present species in hav ing the orbicularoutline and the smaller number of theradial ribs on the surface.
Occurrence :- Loc. 13.
Clycymeris (C lycy m e1·ita )multicostata NAGAO
P late 27, figs . 28- 32

1932. C!ycyme ri s hoklwidoe ns1:s
Tex t -fi g . 4. C!ycymeris ( C!ycymerita)
himenou rensis T AS I-II R O, ne w s pec ies .

str.) ama lcusensis.
The number ofchevrons at the ligam ent area changes
throu g h g ro wth and the y are about 6
in the adu lt stage.
Comparison :- This species is disting uished from Clycymeris (Clycym erita )jap onica from the upp er formation ofth e same gro up by the high and smaller
shell. C. umbonatus (SOWERB Y) from theGreensan d at Blackdo wn in Eng land issom ew hat akin to this species in the wellinflated and hig h shell. C. umbonalus
differs from this species in its small
umbo and its as ymmetrical hinge plate.
C. wnbonatus is probab ly a member ofsubgenus H an.aia. C. sachalinensis (Y A BE
and N AGAO) (1925) is somewha t simi larto this species in the stout ribs of th e
surface and the large and prominentumbo. C. sacha li n.en.sis diffe rs from theva r. mu!t icos ta.ta. NAGAO , j our. Fa c. S ci ., Ho!?!w ido
I mp . Univ., Ser. 4 , Vol. 2, p. 34, pl. 5,
figs . 10, 11.

Ob ser vation and R emarks :- On somead ult specimens the surface ornamentation is worn away but the subinternalar.d elevated radial ridg es can be seen.
The number of the chevrons in liga ment
area is about 8 in the adult stage. The
immature specimen is v ery similar tothe subtriang uler form of Clycymeris(s. str.) ama/zusensis in its outline, ex te rnal ornamentation and the structureof the hinge plate. The variegatedcolor patterns appear on th e umbonal
surface of an imm at ure shell. The
species is chara cteri zed by its orb-icul arly subquadrate and vvell inflated shell,la rge and prominent umbo , elo ngatedand stout hinge s tructure and flat-topped
radial ribs without threads on them .
These features are chracterist ics ofClycymerita .

M easurements (in mm.) :Specimen
KE 1858 Lef t va lve
K E 1861 Left va lve
KE 1860 Le f t va l ve
KE 1863 Left va l ve
KE 186'1 Left val ve
KE 1866 Righ t va lve
GKH 6990 Ri g h t val ve
GKH 6992 Le ft va lve
GKH 6993 Left va lve

C. (C!ycymeri ta ) mu!tico sta ta
Len gt h
Hei g h t

26.0
25.2
24.9
23 .1
20.8
15. 0
25. 8
20 . 8
24. 0

25. 5
25. 0
24.7
23. 0
19.5
14.0
zs. 8
19. 0
23 . 4Thickne ss

9. 8
9 ·6
9.0
8. 5
6. 5
4. 2
9. 5
5.2
9. 5

590 . . Cretaceous Glycymerids
Occurrence :-Locs. 2 and 19.

233

the subgenus Hanaia.
Glycymeris (Hanaia) salida NAGAO

Subgenus Hanaia HAY AMI, 1965
Type-species :-Glycymeris den silin eat a
NAGAO.
Hanaia is characterized with roundtopped radial ribs which are composedof a bundle of fine radial threads on
surface as already stated by HAY AMI(1965). The species of Hanaia changes
its shape of shell through geologicalages. This morphological change isshown in Text-fig. 5. This phylogenyof Hanaia from morphological standpoint is observed in the ontogeny oflower Lower Cenomanian species of G.
(H.) salida. On the basis of the morphological change, it seems possible that
I can estimate the age of species ofHanaia. NICOL (1950) included G. mallurensis (LEYMERIE) from the Neocomianof Europe and G. sublaevis (SOWERBY)
from the Aptian of England into subgenus Glycymerita. G. haipensis HAY AMI
from the Aptian of Japan was placed in
Glycymerita ? by HA YAMI (1965). Abovementioned Lower Cretaceous glycymerids, however, differ from the typical
Glycymerita in their small and pointed
umbo, asymmetrical hinge structure andthe distinct radial threads on the ribsof the external surface. It seems forme that these species may belong toPlate 28, figs. 1-8, Text-fig. 51930. Glycymeris amakusensis var. solida
NAGAO, jour. Fac. Sci., Hokkaido Imp.
Univ., Ser. 4, Vol. 1, pp. 16, 17, pl. 2,
figs. 13 and 14.
1958. Glycymeris aff. amakusensis, AMA:--:o,Dept. Geol., Fac. Sci., Kumamoto Univ.
pp. 68, 69, pl. 1, figs. 22-25.

Observation and Remarlls:-This
species was described as a variety of
Glycymeris amakusensis by NAGAO (1930)
because of its more inflated shell andits thicker test. In this species thechevrons on the ligament area are countable 6 or more at the adult stage. The
narrow and fine radial ridges appear
frequently on the internal surface near
the umbo but they are obscure on bothsides and in the ventral border. Radial
threads on a rib are about 8 in number.
Occurrence:- Locs. 4, 5 and 6.
Glycymeris (Hanaia) matsumotoi
TASHIRO, .new species
Plate 28, figs. 17-21

Description :-Shell small, asymmetrically subquadrate, longer than high,well inflated; anterior margin rounded
and passing gradually into arcuate

Measurements (in mm.) :G. (Hanaia) solida
Specimen
Length
28.5
KE 1760 Left internal mould
27.0
KE 1765 Left internal mould
23.5
KE 1761 Left internal mould
23.1
KE 1762 Left internal mould
18.0
KE 1768 Left internal mould
16.0
KE 1767 Left internal mould
13.0
KE 1763 Left internal mould

Height
25.0
23.2
21.1
18.0
17.0
14.3
12.9

Mas a yuki T ASH! RO

234

10 mm

Text-fig . 5. Gro wt h and geo logica l c han ges of subge nus Hanaia.
1- 5, The morpholo g ical chan ge t hroug h g r owt h of Clycymeris (Ha.naia ) so/ida.
NAGAO (Lo wer Cenomanian).
A- E, Ad ult shells of diffe ren t species in successi ve geo lo gical ages show ing
t he m orpholo g ica l cha n ge. A: C. (H. ) ha.ipensis H AY AM I, (Aptia n) (afte r
H AY AM I, 1965) . B: C. (H .) sublaevis (SOWERBY), (A pti an) (afte r W OODS, 18771913). C : C . (H. ) densilinea.ta N AGAO (Albio- Apt ian ), (a fter H AY AMI, 1965).
D: C. (H.) matsurnotoi T AS HIR O, n . sp . (Alb ian?) . E: C. (H. ) kat su ra.za. we nsis
T ASHI RO, n. sp . (Turoni an).

-ventral marg in; posterior ma rgin obli ·quely subtruncated; umbo small, situatedfairly posterior to mid-point of length,
beak orthog yrous, situated posterior tothe mid-point of dorsal margin; surface
ornamented with numerous round-topped
radial ribs each of which is a bundle of
7 or so threads; lig ament area na rrow,provided with several chevrons; hinge
plate asymmetrical, subhorizontal in.anterior half but oblique in posterior
half; teeth subvertical and small below
umbo, stout and weak ly hooked in bothlateral areas on the hinge p late; inner
ventral margin moderatl y crenu lated as

many as about 25 ; g ro wt h lin es on
surfac e weak.
Obse1'vation and R emar!?s :- MATSU MOTO (1953) listed Glycymeris elongata(MS) from the Loc. 8. The mor]:.>holog ical chan ge throug h growth is observable in this species as seen in C.
(H.) solida except for its last stage (Textfig. 5- 5). The n umber of liga ment
chevrons is about 5.
Comparison :- This species is distinguishab le from C. (H.) densilineata by
the more abundant crenules on the inner
ventra l marg in and the elongate out line.
G. (H.) haipensis is similar to this species;v!easurements (in mm .) :-c. (Ha.naia) matsumotoi
Specimen
KE 1869 Ri g ht exte rnal mo u ld
KE 1870 Right in te rn a l mould
KE 1871 Left in terna l mould
KE 1872 Left exte rn a l mould
KE 1873 Left in ter nal mould

Length
12. 4
10. 5
10.8
10. 0
13.2

Hei g ht
11. 0
9.9
9.2
9.0
12. 5

T hi ck ness
3.8

3.0

Ho lot y peParatype
Ditto.
Ditto .
Ditto.

.590.

235

Cretaceous Glycymerids

in the asymmetrical hinge plate and theasymmetrically subquadrate outline. In
G. (H.) haipensis the hinge teeth are,however, more stout and more primitive
than in this species.
Occurrence :-Locs. 7 and 8.
Glycymeris (Hanaia) katsurazawensis

TASHIRO, new species
Plate 25, figs. 9-15

Description :-Shell large for Hanaia,
well inflated, higher than long, orbicularly
subtriangular; anterior and ventral
margin rounded, posterior margin obliquely subtruncated; umbo prosogyrous,
prominent, located anterior to midlength; beak nearly orthogyrous, situateda little posterior to mid-point of dorsal
margin; surface ornamented with roundtopped radial ribs each of which bears
fine radial threads (about 9) ; a triangulardark colored area on posterior umbonal
part extends about 7 mm. from apex;
ligament area narrow and trianguler;
hinge plate fairly long, symmetricallyarched but angularly bending at bothlateral sides; teeth small and subvertical
below umbo stout and hooked to diagonal
on both lateral parts; inner margin

crenulated and 22 or so in number; upper
margin of hinge plate close to dorsal
margin of valve.
Obseruation and Remarks:- The umbonal part of this species resembles closely
shell characters of G. (H.) densilineata.
Consequently it seems for me that thisspecies was derived from G. (H.) densilineata. This species represents the laststage of morphological change of subgenus Hanaia.
Comparison :-This species is distinguished from G. (H.) ho!?i<aidoensis
with the anterior location of umbo to
the shell-length and the more abundant
threads on the ribs.
Occurrence :-Locs. 1 and 20.
Glycymeris (Hanaia) hohkaidoensis(YABE and NAGAO)
Plate 28, figs. 16a-16c
1928. Pectunculus hokkaidoensis YABE and::-JAGAO, Sci. Rept. Tohoku Imp. Univ.,
2nd Ser., Vol. 9, No. 3, pp. 82, 83, pl.
17, fig. 22.

Obseruation and Remar!?s :-The shellis asymmetrically subquadrate and theumbo is located posterior to the midlength. The round-topped radial ribs ofJ\1easurements (in mm.) :-G. (Hanaia) katsurazawensis
Specimens
KE 1854 Left valve
KE 1855 Left valve
KE 1856 Left valve
GKH 6980 Left valve
GKH 6981 Left valve
GKH 6982 Right valve

Length
24.0
25.0
23.5
25.8
24.0
21.0

Thickness
9.8
10.6
10.0
9.5
8.0

Height
24.8
26.7
26.0
25.9
24.0
21. 1

Holotype
Para type
Ditto
Ditto
Ditto
Ditto

G. (Hanaia) hokkaidoensis

Specimen
GKH 6984a Right valve
GKH 6984b Left valve

Length24. 5
25.0

Height23. 5
24.0

Thickness
8.6
9.0

Masayt~ki TASHIRO

236

this species are simi lar to those in C.
(H.) lwtsurazawensis except for thenumber of threads (abo ut 7 in thisspecies). C . (H.) matsumotoi is similar
to this species in the outline and theexternal sculpture, but differs from thisspecies in the sma ller size and numerous
ribs on the surface. The s ha pe of
middle aged stage of C. (H.) so/ida (3 and
4 in Text-fig. 5) is akin to this shape.
The surface is, ho wever , far stouter inthis species than in C. (!-!.) solida.
Occurrence :- Sandstone of the Saku
formation at the lower reaches of theKotosawa, Abesh inai district, Hokkaido(Loc. T - 453p, by MATSUMOTO).
Subgen us P seudoveletu ceta
TASHIRO, new s ubgenus
T yjJe-sjJecies :- Cly cymeris (Pseudoveletuceta) mifunensis T ASHTRO, new species.
Diagnosis :- Shell
small, orbicular,
more or less inflated, inequilateral ; umbo
small, prosogyrous, located anterior to
midd le; apex of ligamen t-triang le anterior to th e base; ligament area witha fe w che vro ns ; surface a lmost smooth
ex cept for fine rad ial threads; hinge
plate moderatel y arched with tax odont
teeth ; inner margin de nsely crenu lated.
R emaTI?s :- Th e characteristics of thissubgenus are as f ollows: 1) the ante riorlocation of beak, 2) the prosog yrous
umbo vvhich is located rem a rkabl y anterior to the mid-length, 3) the crowded

Text-fig. 6.

fine radial threads forming no bundle;
P ectunculina D'ORB IGNY is similar tothis subgenus in its prosogy rous umbo ,ventra l crenulation and hinge structure.
Pectuncu.lina, however, has a triangularligament pit in the centra l part of ligament.
This is of the Limopsidae_
Pro tarca STEPHENSON is similar to thissubgenus in havi ng a prosogy rous umbo
which is not detected in other generaof th e Gl yc ymeridinae refer red to NICOL(1945) and I EWELL (1969). I include thissubgenus into Cly cymeris, for it has
ventralcrenu lation s, archedhinge
structure and triang uler li ga ment areawith chevron grooves.
Clycymeris (P seu dovele tu ceta) mifunensis

T ASHIRO, new species
Plate 28, figs. 2'1- 30, Te x t-fi g . 6

D esc ription :- Shell ver y small, lon ger
than high, orb icu lar in outline, modelatel y inflated; anterior margin rounded;
posterior margin obliquel y expanded toposterior;v entral marg inbroadlyarched; umbo smal l and pointed, prosogy rous, located about two-fifths fromfront ; beak situated anteriorly; surface
ornamented with numerous fine radia lthreads; hinge plate rather narrow,moderatl y arcuated ; teeth subverticaland short below umbo , weakly hooked
to diagonal on both lateral sides; inner
margin crenu lated a bout 30 in number ;g rowth-lines weak.

G!y cy m zr is ( Pseu.doveletuceta) rmfunensis T AS HIRO,

new subge nus and new spe cies .

590.

Cretaceous Glycymerids!vfeasurements (in mm.) :G. (Pseudoveletucela) mifunensis
Specimen
LengthHeightThickness
KE 1774 Right internal mould
15.0
13.0
KE 1769 Left external mould
12.6
11. 0
3.4
KE 1776 Right internal mould
16. 0
13.0
KE 1777 Right internal mould
13.5
10.0
KE 1770 Left internal mould
12.8
9.0
KE 1771 Right internal mould
7.5
6.3

Observation and Remarks:- There areround-topped radial ribs in immature
specimens. and position of the beakand umbo is nearly at the middle oflength. Three chevron shaped ligamentgrooves in adult stage cross theirjunctions and make tubercles in somecases. In the valves longer than 12 mm.
in length the radial threads are crowded
and countable about 8 in number in the
distance of 1 mm.
Comparison :-This is similar to G. (s.
str.) amalwsensis in the outline and thehinge structure, but this differs fromG. (s. str.) ama!wsensis in the anteriorly
situated beak and the surface without
radial ribs. Round-topped radial ribs
and median location of umbo and beakare seen in immature specimens. Thesefeatures arech aracteristics in adult of
the species of Hanaia.
Occurrence :-Locs. 9 and 10.

Holotype
Para type
Ditto
Ditto
Ditto
Ditto

Plate 28, figs. 31-34
1958. G!ycymeris kogata ]CIIII<AWA and MAED:\, jour.
Inst. Po!ytech. Osaka C1:ty
Un£v., Ser. G. Vol. 4, p. 90-92, pl. 5,
figs. 4-7, 10.

Observation and Remarks:- Though
the specimens from the Himenoura groupare smaller than the specimens fromthe Izumi group of Awaji-island. theother morphological characters accord
between the specimens from the localities.
No precise description on ligament area
was given by ICHIKAWA and MAEDA(1958). The distinct ligament pit wasobserved by me in the specimens fromthe Izumi group (Topotype OCU 0211 bylCHIKA WA and MAEDA) and the Himenouragroup.
This species is referred toLimopsis by the mentioned characteristics.
Occurrence:- Lacs. 17 and 18.
References cited

Family Limopsidae DALL, 189;)
Genus LimojJsis SASSI. 1827
Type-species :-Area
1814.

aurita

BROCCHI.

Limopsis kogata (lCHIKA\VA

and MAEDA)

A~L\:-:o,M. (1956) : Some upper Cretaceous
fossils from southwestern Japan (Part 1).
Kumamoto Jour. Sci., Sect. 1 (Ceo/.) Vol.
], No. 1 pp. 63-86, pis. 1, 2.
___ (1957) : Upper Cretaceous molluscan
fossils from Shimo·Koshiki·iima, Kyushu.
Ibid., No. 2 pp. 51-75, pis. 1, 2.
DAILEY, D.H. and ?OPE:\OE, vV.P. (1964):

1\1easurements (in mm.) :-Limopsis koga ta
LengthSpecimen
5.0
KE 1877 Left internal mould
5.3
KE 1878 Right internal mould

Height
6.0
6.2

2:38

J]!Jasayuki T ASH! RO

Explanation of Plate 27.Clycym2r£s (Glycymeris) amakusens£s NAGAO
Fig. 1. Right valve, KE 1797, x 1, Loc.: Okoshiki at Oda-machi, Uto city, Kumamoto Pref.
Fig. 2. Left valve, KE 1798, x 1, Loc.: Ditto.
Fig. 3. Rubber cast of right valve, internal mould, KE 1813, x l, Loc.: Ditto.
Fig. 4. Right valve, KE 1815, x l, Loc.: Ditto.
Fig. 5. Immature right valve, KE 1805, x 3, Loc.: Ditto.
Fig. 6. Rubber cast of left internal mould, KE 1814, x 1, Loc.: Ditto.
Fig. 7. Umbonal view of right valve, rubber cast of internal mould, KE 1788, x Li, Loc.:
Ditto.
Fig. 8. Right valve, KE 1.836, x 1.5, Loc.: Wadanohana at Takado-machi, Amakusa Co.
Fig. 9. Right valve, KE 1835, x 1.5, Loc.: Ditto.
Figs. lOa, lOb. Lateral and dorsal views of left valve, KE 181.7, xl, Loc.: The northwest
beach of Kugu island at Takado-machi.
Fig. 11. Left valve, KE 1819, x 1, Loc.: Ditto.
Fig. 12. Left valve, KE 1.816, x 1, Loc. : Ditto.
Fig. 13. Left valve, KE 181.8, x 1, Loc. : Ditto.
Figs. 14a, 14b- Lateral and dorsal Yiew of right valve, KE 1822, x 1, Loc.: Ditto.
Fig. 15. Left valve, KE 1820, x 1.5, Loc. : Ditto.
Fig. 16. Left internal mould, KE 1852, x 1, Loc.: The Sakuradani valley at Hirono-machi,
Futaba Co. Fukushi rna Pref.
-Glycymeris (Glycymerita) japonica TASHIRO, new species
Fig. 17. Internal mould of right valve, KE 1786, x 1, I-Iolotype, Loc. : Hongo at Kawauramachi, Amakusa Co. Kumamoto Pref.
Fig. 18. Rubber cast of left internal mould, KE 1796, x 1, Loc. : Ditto.
Figs. 19a, 1.9b. Anterior and lateral views of right valve, rubber cast of external mould,
KE 1794, x 1, Loc.: Ditto.
Fig. 20. Rubber cast of right external mould, KE 1795, x 1, Loc. : Ditto.
Fig. 21. Internal mould of right valve, KE 1791, x 1, Loc.: Ditto.
Fig. 22. Internal mould of left valve, KE 1792, x 1, Loc.: Kamihira at Kawaura-machi,
Amakusa Co .
.Clycymeris (Glycymerita) himenourensis TASHIRO, new species
Figs. 23a, 23b. Lateral and dorsal views of right valve, KE 1779, x 1.5, Holotype, Loc.: The
north-west beach of the Kugu island at Takado-machi, Amakusa Co.
Fig. 24. Immature left valve, KE 1.781, x 3, Loc.: Ditto.
Figs. 25a, 25b. Lateral and dorsal views of imperfect left valve, KE 1.780, x 1.5, Loc. : Ditto.
Fig. 26. Umbonal view of right valve, KE 1782, x 1.5, Loc.: Ditto.
Fig. 27. Ligamenta! view of left valve, KE 1783, x 2, Loc.: Ditto.
Glycymeris (Glycymerita) multicostata NAGAO
Figs. 28a, 28b. Lateral and dorsal views of left valve, KE 1.854, x 1, Loc.: Sakainosawa,
Kyowa, Nakagawa-machi, Teshionakagawa Co., Hokkaido.
Fig. 30. Left valve, KE 1855, x 1, Loc.: Ditto.
Fig. 31. Immature right valve, KE 1863, x 1, Loc.: Ditto.
Figs. 29a, 29b. Lateral and dorsal views of left valve, GKH 6990, x 1, Loc.: Gakkonosawa at
Yasukawa, Nakagawa-machi, Hokkaido.
Figs. 32a, 32b. Lateral and internal views of left valve, GKH 6992, x 1, Loc. : Gakkonosawa
at Yasukawa, Nakagawa-machi, Hokkaido.

(to be continued)

TASHIRO :Cretaceous Glycyme1·ids-'

Plate 27

lS
28b

22

21

590.

Cretaceous Glycymerids

Mollusca from the Upper Cretaceous
Jalama formation, Santa Barbara county,
California. Univ. Calif. Publ., Geol. Sci.,
Vol. 65, pp. 1-27, pis. 1-4.
FINLAY, H.]. and MARWICK, ]. (1937): The
Wangaloan and associated molluscan
faunas of Kaitangata-Green island subdivision. /liew Zealand Ceo/. Surv., Palaeon[. Bull., 15, pp. 1-140, pis. 1-18.
FREl'\"EIX, S. (1959) : Lamellibranches duCretace superieur de France (Protobranches,Prionodon tes,Dysodon tes (pars)).
84• Congres des Soc. Savantes, pp. 175-248.
GoLDFcss, A. (1833-1840) : Petrefacta Germaniae. Theil 2, 512 pp., pis. 72-165,
Dusseldorf.
HABE, T. (1951-1953): Genera of the Japaneseshells. Pts. 1-4, 326 pp. (in Japanese).
HAY AI\!!, I. (1965) : Lower Cretaceous marine
pelecypods of Japan. Part I. Mem. Fa c.
Sci., Kyushu Univ., Ser. D. Ceo/., Vol. 15,
No. 2, pp. 221-349, pis. 27-52.
IcHIKAWA, K. and MAEDA, Y. (1958): Late
Cretaceous pelecypods from the Izumi
group. Part II, Orders Taxodontida,
Prionodontida, Dysodontida, Desmodontida and Adapedontida.
jour. Inst.
Polytech., Osaka City Univ., Ser. G, Vol.
4, pp. 71-122, pis. 3-7.
IREDALE, T. (1931) : Australian molluscan
notes, No. 1. Australian Mus. Rec., Vol.
1, No. 4, pp. 201-235, pis. 22-25.
MAKIYAMA, ]. (1957-1958): M. YoKOYAMA's
Tertiary fossils from various localities inJapan, Pts. 1-2. Palaeont. Soc. japan,
Spec. Paps., Nos. 3, 4.
MATSUMOTO, T. (1938) : The Geology ofGosyonoura Island, Amakusa. jour. Geol.
Soc. japan, Vol. 45, No. 532, pp. 1-46,

239

pis. 1-3.
- - (1939) : Geology of the Mifune district,
Kumamoto Prefecture. Ibid., Vol. 46,
No. 544, pp. 1-12, pl. 1 (in Japanese).
- - (1954 ed.) : The Cretaceous System in
the Japanese Island. xiv+324 pp., 36 pis.
japan Soc. Prom. Sci. Res., Tokyo.
- - , AMAKO, M., OKADA, H. and Occr<r, H.
(1960) :Graysonites (Cretaceous ammonites) from Kyushu with notes on
stratigraphy. Mem. Fac. Sci., Kyushu
Univ., Ser. D (Geol.), Vol. 10, No.1, pp.
41-58, pis. 6-8.
- - , and KAi'\MERA, K. (1964) : Explanatory
text of the geological map; Hinagu.
Geol. Surv. japan (in Japanese).
NAGAo, T. (1928) : Palaeocene Fossils of
the island of Kyushu, Japan, Part II.
Sci. Rept. Tohoku Imp. Univ., Ser. 4,
Vol. 9, No. 3, pp. 11-140, 17 pis.
- - (1930) : On some Cretaceous Mollusca
from the island of Amakusa, Kyushu,
Japan. jour. Fac. Sci., Hokkaido Imp.
Univ., Ser. 4, Vol. 1, No. 1, pp. 1-25,
pis. 1-3.
- - (1934) : Cretaceous Mollusca from the
Miyako district, Honshu, Japan. Ibid.,
Vol. 2, No. 3, pp. 177-277, pis. 23-39.
- - (1938) : Some molluscan fossils fromthe Cretaceous deposits of Hokkaido andJapanese Saghalien. Part 1, Lamellibranchiata and Scaphopoda. Ibid., Vol. 4,
Nos. 1-2, pp. 177-142; pl. 14.
NEWELL, N.D. (1969) : Superfamily Limopsacea, in Treaties on Invertebrate Paleontology (MooRE ed.) ,-(I) Mollusca 6-.
Ceo/. Soc. Amer., Univ. Kansas Press,
pp. N 264-N 269.
NICOL, D. (1945) : Genera and Subgenera of·--- -----------(continued from p. 238)

Glycymeris (Glycymerita) veatchii (GABB) (for comparison)*Figs. 33a, 33b. Lateral and dorsal views of both valves, 33a,: left vah·e, GKH 6983, x 1.
Glycymeris (Glycymerita) aff. apletos DAILAY and PoPENOE (for comparison)*Fig. 34. Right valve, GKH 6988, x 1.
Fig. 35. Internal view of right valve, GKH 6989, x 1.
* G. (G.) veatchii and G. (G.) aff. apletos are collected by T. MATSUMOTO from the Upper
Cretaceous Chico group in California, U.S.A.

24()

Masayuki TASHIRO
Explanation of Plate 28

Glycymeris (Hanaia) salida NAGAO
Fig. 1. Rubber cast of left internal mould, KE 1760, x 1, Loc. : Karakizaki of the Goshonoura
island, Amakusa Co., Kumamoto Pref.
Fig. 2. Rubber cast of left internal mould, KE 1765, x 1, Lac. : Shirahama of the Shishi
island, Izumi Co. Kagoshima Pref.
Fig. 3. Rubber cast of left internal mould, KE 1767, x 1, Lac.: Ditto.
Fig. 4. Rubber cast of left internal mould, KE 1768, x 1, Loc.: Ditto.
Fig. 5. Ditto; KE 1762, x 1, Lac.: Enokuchi of the Goshonoura island.
Fig. 6. Imperfect left valve, KE 1761, x 1, Lac.: Ditto.
Fig. 7. Rubber cast of internal mould, immature left valve, KE 1763, x 1, Lac.: Ditto.
Figs. Sa, Sb. Lateral and dorsal views of imperfect left valve, KE 1764, x 1, Lac.: Ditto.
Glycymeris (Hanaia) katsurazawensis T ASI-IIRO, new species
Figs. 9a, 9b. Lateral and dorsal views of right valve, GKH 6983, x 1, Lac.: The coast of thewaterway near the Katsurazawa dam, Ikushunbetsu, Hokkaido.
Fig. 9c. Umbonal view of GKH 6983, x 1..5.
Figs. lOa, lOb. Lateral and dorsal views of left valve, GKH 6981, x 1, Lac.: Ditto.
Fig. 11. Internal view of right valve, GKH 6982, x 1, Lac. : Ditto.
Fig. 12. Left valve, KE 1855, x 1, Holotype, Lac.: About 500 m. south of the Katsurazawa
dam, Ikushunbetsu.
Fig. 13. Internal view of left valve, KE 1855, x 1, Lac.: Ditto.
Fig. 14. Left valve, KE 1856, x 1, Lac.: Ditto.
Fig. 15. Surface ornamentation of left valve, KE 1857, x4, Lac.: Ditto.
Glycymeris (Hanaia) hokkaidoensis (YABE and NAGAO)
Figs. 16a, 16b. Lateral and dorsal views of right valve, GKH 6983, x 1, Lac. : The lower
reaches of the Kotozawa river, Abeshinai district, Hokkaido.
Fig. 16c, Surface ornamentation of GKH 6984, x 4.
Glycymeris (Hanaia) matsumotoi T ASI-IIRO, new species
Figs. 17a, 17b. Lateral and dorsal view of right valve, rubber cast of external mould, KE 1864,
x 2, Lac.: Tani of Miyaji-machi, Yatsushiro city Kumamoto Pref.
Fig. 17c. Surface ornamentation of KE 1864.
Fig. 18. Rubber cast of left internal mould, KE 1865, x2, Lac.: Ditto.
Fig. 19a. Ditto, KE 1870, x 2, Holotype, Lac.: Ditto.
Fig. 19b. Umbonal view of Halo type, x 4.
Fig. 20. Rubber cast of right internal mould, KE 1866, x 2, Loc. : Ditto.
Fig. 21. Rubber cast of left internal mould, KE 1869, x 2, Loc. : Ditto.
Glycymeris (Hanaia) dens£/ineata NAGAO (for comparison)Fig. 22. Right valve, GKH 6642a, x 1.5, Lac.: Hiraiga of Tanohata village, Iwate Pref.
(HAYA:VII collect.)Fig. 23. Left valve, GKH 6642b, x 1.5, Lac. : Ditto. (HAYA;'v!I collect.)
Glycymeris (Pseudoveletuceta) mifunensis T ASI-IIRO, new species, new subgenus.
Fig. 24a. Internal mould of left valve, KE 1769, x 2, Lac.: Nishiyama of Katashida-machi,
Shimomashiki Co. Kumamoto Pref.
Figs. 24a, 24b. Lateral and dorsal views of KE 1769, rubber cast, x 3.
Fig. 25. Internal mould of immature right valve, KE 1771, x3, Lac.: Ditto.
Fig. 26. Internal mould of right valve, KE 1777, x2, Lac.: Ditto.

(to be continued)

TASHIRO:

Cretaceous Glycymerids

Plate 23
8a

I

I 23

17c

30

24c
-- '

28a

29a

590.

Cretaceous Glycyrnerids

the pelecypod Family Glycymeridae. jour.
Paleont., Vol. 19, No. 6, pp. 616-621.
- - (1950) : Origin of the pelecypod Family
Glycymeridae. Ibid. Vol. 24, No. 1, pp.
89-98, 3 pis.
- - (1956) : Distribution of living glycymerids with a new species from Bermuda.
Nautilus, Vol. 70, No. 2, pp. 48-53, 2
pis.
0KA:\IOTO, K. and NAKANO, M. (1965):
Clycymeris and Cultellus from the
Tertiary Hioki (Ashiya) Group in the
Yuya-wan Area, Yamaguchi Prefecture,
Southwest Japan. jour. Sci. Hiroshima
Univ., Ser. C, Vol. 2, pp. 531-539, pl.
57.
SAITO, T. (1961) : The stratigraphic studiesof the Futaba-gun, Fukushima-prefecture,
Japan, Part 1. Bull. Arts and Sci., Ibaraki Univ., Nat. Sci., No. 11, pp. 108-113.
(in Japanese).
- - (1962) : The Upper Cretaceous System
of Ibaraki and Fukushima Prefecture,
Japan, Part 2. Ibid., No. 13, pp. 52-87,
8 pis.
SKAW ARKO, S.K. (1967) : Mesozoic Mollusca
from Australia and New Guinea. Bur.
Min. Resources Ceo/. Ceophys., Austratz:a,
Bull. no. 75, pp. 1-101, pis. 1-12.
STOLICZKA, F. (1871) : Cretaceous fanna ofsouthern India.
Vol. 3, Pelecypoda,
Palaeont. Indica, pp. 1-537, pis. 1-50.
TA:>.ICRA, M. and TASHIRO, M. (1966):

241

Upper Cretaceous System South of Kumamoto. Mem. Fac. Educ. Kumamoto Univ.,
No. 14, Sec. 1 (Nat. Sci.), pp. 24-34, pl. l.
- - and - - (1967) : Some problems on theage of the Mifune group (abstruct inJapanese). jour. Ceo!. Soc. Japan, Vol.
73, No. 2, p. 137.
- - , - - and MoTOJI:\L\, T. (1968): The
correlation of the Mifune Group with
the Upper formation of the Gosyonoura
Group, with description of some important pelecypods from the strata. Mem.
Fac. Educ., Kumamoto Univ., No. 16, Sec.
1 (Nat. Sci.), pp. 29--!2, pl. 1.
TA0:AKA, K. (1963) : A study on Cretaceous
sedimentation in Hokkaido, Japan. Ceol.
Surv. japan., pp. 1-119, 3 pis ..
UEDA, Y. and FuRUKA\YA, N. (1960): On
the Himenoura Group of the Amakusakamishima and adjacent islets, Kumamoto
Prefecture. Sci. Rept., Kyushu Univ.,
Ceol., Vol. 5, No. 1, pp. 14-34 (in Japan.).
- - (1962) : The Type Himenoura Group.
J'v!em. Fac. Sci., Kyushu Univ., Ser. D(Ceo!.), Vol. 12, No. 2, pp. 129-160.
Wooos, H. (1899-1913) : A monograph of theCretaceous Lamellibranchia of England.
Vol. 1 (1899-1903), xliii+227 pp., 42 pis.
YABE, H. and NAGAO, T. (1925): New orlittle-known Cretaceous Fossils from
North Saghalin (Lamellibranchiata and
Gastropoda).
Sci. Rept. Tohoku Imp.
Unit·., 2nd Ser., Vo1.7 , No. 4, pp. 111-

(continued from p. 240)
Fig. 27. External view of immature right valve, rubber cast, KE 1773, x 4, Loc.: Ditto.
Fig. 28a. Internal mould of right valve, KE 1776, x 1.5, Loc.: Asanoyabu of Mifune-machi,
Kamimashiki Co. Kumamoto Pref.
Fig. 28b. Rubber cast of KE 1776, x 3.
Fig. 29a. Internal mould of right valve, KE 1774, x 1.5, Holotype, Loc. : Ditto.
Fig. 29b. Ligamenta! view of Holotype, rubber cast, x 3.
Fig. 30. Surface ornamentation of right valve, rubber cast, KE 1775, x 7, Loc.: Ditto.

Limopsis kogata (Ici-IlKAWA and MAEDA)
Fig. 31. Internal mould of left valve, KE 1877, x 3, Loc.: Hongo at Kawaura-machi, Amakusa
Co. Kumamoto Pref.
Fig. 32. Rubber cast of imperfect right valve, ligamenta! view, KE 1880, x 5, Loc. Ditto.
Fig. 33. Rubber cast of imperfect right? valve, external mould, KE 1875, x7, Loc.: Ditto.
Fig. 34. Rubber cast of internal mould, OCU 0210, x 2, Loc.: The Awaji-island, Hyogo Pref.
(IcHIKAWA and MAEDA collect.)Masaynki TASHIRO

242

124, 2 pis ..
- a n d - (1928) : Cretaceous fossils from
Hokkaido; Annelida, Gastropoda and- -----

Lamellibranchiata. Ibid., Vol. 9, No. 3,
pp. 77-96, pls. 16-17.

------~-----

Abeshinai
Amakusa
Asanoyabu
EnokuchiFutabaGakkonosawa
Goshonoura
HironoHongo
Ikushunbetsu
Izumi
Kamihira
Kamimashiki
Karakizaki
Katashida
Katsurazawa
Kawaura
K6da
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243
PROCEEDINGS OF THE PALAEONTOLOGICAL SOCIETY
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Miyagi Prefecture ...... MURATA, M.,
BA!'>DO, Y., lSI-III, K. & NAKAZAWA, K.
On Utatsusaurus, a new genus of Ichthyosauria ................................. .
SHIKA:\1A, T., KAMEl, T. & MURATA, M .
A Problematica from the Mizuho TO of
Niigata Prefecture (ftij)C) ............. .
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Japan ........................... IEDA, K.
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. . . . . . . . . . . . NAKASEKO, K. & SUGA:-10, K.
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Name and surname; Title; Address; Speciality (main interests in researches, otherinterests) ; Present research projects expected to result in publications .
* ~~ ffij Dr. ]. P. CHEV ALI ER , Museum Nationa l d'Histoire Naturelle, Institute dePaleontologie, 8, Rue de Buffon, Paris-Ve, FRAN CE .
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Prof. K. V. Nikiforova (Chairman of the Organizing Committee), Geological Institute,
USSR Academy of Sciences, Pyzhevsk y per. 7, Moscow, USSR.

-r," List of Members" Issued Sept. 30, 1971
Page 1, Reight column, read *BANDO for BANDO
Page 12, Left column, SATO, Seiji; read Dept. Geol. & Mineral., Hokkaido Univ.,
Sapporo (060) for 4-1890, Kamitakaido, Suginami-ku, Tokyo (167).

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Transactions and Proceedings of the PalaeontologicalSociety of japan

December 30, 1971

New Series No. 84

CONTENTS
TRANSACTIONS
586.

MATSUMARU, Kuniteru: The genera NephTolepidina and Eulepidina
from New Zealand . . ............................... . ... . . ... . . . ....... 179

587.

KIMURA, Tatsuaki and SEKIDO, Shinji:

The discovery of the cycad ·likeleaflets with toothed margin from the Lower Cretaceous Itoshiro
Sub-group, the Tetori Group, Central Honshu , Japan.. .. ........... . 190

588.

HAY AMI, Tomoko: On some Bryozoa from near Namioka-cho, MinamiTsugaru-gun, Aomori Prefecture, Japan ... .... . ... ........... .. . ..... 196

589.

MASUDA, K6ichir6:

Amussiopecten from North America and northern

South America ......... . .. , .. . .... . ............. . . . ..... . ..... . . . ..... 205
590.

TASHIRO, Masayuki:

Upper Cretaceous glycymerids in Japan . .. ......... 225

PROCEEDINGS ....... ....... . . .. ...... .... .. . ..... .. ...... . .. ... . . .. .... . .. ... 243