ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 107

Palaeontological Society of Japan
September 15, 1977

~

.....

EditorItaru HAYAMI
Associate editors Ikuwo OBATA and Tomowo OZAWA

Officers for 1977 -1978
Honorary President: Teiichi KOBA YASIII
President:

Ryuzo TORIY AMA

Councillors: Kazuo ASAMA, Kiyotaka CHINZEI, Takashi HAMADA, Tetsuro HAN AI,
Itaru HAY AMI, Syozo HAY ASAKA, Hisayoshi IGo, Tadao KAMEl, Kametoshi
KANMERA, Tamio KOT AKA, Tatsuro MATSUMOTO, Tsugio SHUTO, Yokichi
TAKAYANAGl, Toshimasa TANAI, Hiroshi UJIIt
Members of Standing Committee: Kazuo ASAMA (Membership), Kiyotaka CHINZEl(General Affairs), Takashi HAMADA (Foreign Affairs), Tetsuro HANAI, Itaru HA YAM I (Transactions), Hisayoshi IGO (Planning), Kametoshi KANMERA(Special Papers), Yokichi TAKAYANAGl ("Fossils"), Hiroshi UJIIE (Finance),
Juichi YANAGIDA
Secretaries:

Ienori FUJIYAMA (Special Papers), Kunihiro ISHIZAKl ("Fossils"),

Ikuwo OBATA and Tomowo OZAWA (Transactions), Kazuhiko UEMURA(Membership), Toshiyuki YAMAGUCHI (General Affairs)
Auditor: Tatsuaki KIMURA

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

1877, from the Tetori group.

GEYLER's description marked the onset ofmodern palaeontology in Japan.

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

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 107, pp. 125-134, pis. 17-18, September 15, 1977

680.

AN OCCURRENCE OF ORDOSOCERAS IN JEHOL, NORTHEASTCHIN A, AND A NOTE ON THE POL YDESMIIDAE*TEIICHI KOBAYASHIJapan Academy, Tokyo~MiiJj£O) Ordosoceras 2:: Polydesmiidae f:''':J,'-C: Actinoceroid O)J'IltE0) kn 2:: ~n-C l ,t.::. Polydesmia I:' ":Jl '-CUltl]>"0 -C~~~ L t.::.:. b'J~;!; 0 n:, ~* 35 "fr,,":'g7 <O)*'T*~n;,*'S~n-Cl'00 M¥iiJj£O) Ordosoceras jeholense, nov. ~8C1lll:-t0f:.~t'::'"0-C, .;cO)iUl~~0)F.c1lll:0).;z,.tJ: Gf, Polydesmiidae 0)5t1Wl2:: 5t;(rj~~~ii;j" Lt.::.o .;cO)*ii:W,:IBffI;O) Chazyian
1ltrW], 4-13 0) Whiterock f:'vilt,ffi~-t 0:nLllJtJH:., *7:; 7 "CU7\:;0 Wutinoceras n:, (j(l ,"r:Polydesmia, Ordosoceras n::±rm Lt.::.o :.;h G2~'t~¥iiJ&:llMtj£-c:;!; 0 ~rftO)Meitanoceras
IU~ G < 1::. ..... '7 -V o)l 'nVP 0 Ordosoceras fLfi 2:: ;l1;1:.jff4~tJ: L -Co' 00'J,ttfi0

Polydesmia is an interesting example
in taxonomy that a gastropod genus
turned out a senior synonym of acephalopod genus. In 1931 I describedMaruyamaceras, nov. as an Ordovician
cephalopod genus (KOBAYASHI, 1931a),but later when I happened to see
LORENZ'S paper from the reverse side,r was strucked to find that his Polydesmiaof which he quoted that "Die Schnecken..
natur dieser Versteinerung ist unzwei ..
felhaft (1906, 5.103)", essentially coincidedwith the siphuncle of Maruyamaceras.
Merit to TEICHERT (1937) their synonymy
was subsequently warranted through
LORENZ'S type revision.
In 1940 r have studied available speci ..
rri\ens of Polydesmia including a ph rag ..
motone of the type .. species. As theresult I proposed a new family, Poly ..
desmiidae for Polydesmia (Polydesmia
and Maruyamacerina). Because of its
archaeism special attention was paid on
it for the evolution of the Actinoceratida* Received May 10, 1977; read June 18,
1977, at Shizuoka.

125(TEICHERT. 1933; FLOWER, 1957, 1968).
During these thirty .. seven years many
informations were added to our knowledge on this family including Ordosoceras
CHANG, 1959 and other new allied
species. It was long ago that TOKUDA
has submitted me a small collection of
Ordovician fossils from Jehol in theextreme southwestern part of Northeast
China (Manchuria).
At that time I
simply noted an inclusion of Polydesmia
by preliminary identification (KOBAYASHI
and CHOH. 1942), but recently I foundthe Jehol Polydesmia to be a new
species of Ordosoceras CHANG, 1959 byreexamination. Therefore Ordosoceras
jeholense is proposed for it and its
detailed description is presented in thispaper.
On this occasion I reconsidered thespeciality of the Polydesmiidae withadditional facts gathered since its proposal. The siphuncular segment whoselateral wall is composed of a medianannulation and a constriction on eachside is quite distinctive of this family
as well as the Meitanoceratidae, nov.

126

Teiichi J{OBAY ASH!

Compared to the former the segment isnarrower and taller, radial canals areextending from the central canal laterally and lamellar structure is undeveloped in siphuncular deposits in thelatter family. The Polydesmiidae comprise Polydesmia, }vIaruyamacerina andOrdosoceras among which the posterior
constriction is more and less developed
respectively in Ordosoceras and JV1aruyamacerina. The horn-shaped siphunculardeposits and oblique radial canals are
best shown in Polydesmia and somespecies of Ordosoceras. The septum is
ellipochoanoidal in Ordosoceras, but apparently holochoanoidal in the twoothers.
These three genera are typical of the
Maruyaman or early Toufangian faunaof the Hwangho basin. In the AlashanChaotzushan section in western Ordos
the Wutinoceras, Polydesmia, Ordosoceras
and Chisiloceras horizons are in ascending order (CHANG, 1959, 1960). They
are younger than the Wolungian
AI anchuroceras-Archaeoscyplzia fauna
whose age is late or middle-late Canadian. Therefore the Maruyaman fauna
is approximate to the early Chazyan orWhiterock age in North America andLlanvirnian of Europe.
While Manchuroceras was widely distributed in the western Pacific area
(KOBAYASHI, 1977), the family Polydesmiidae were endemic to the Hwanghobasin. lvleitanoceras subglobosum CHEN,
1975 occurs in central China in the early
Llandeilian Kuniutan Formation, but theHimalayan Ordosoceras species may be
a little older. They reveal anothergroup of endemic cephalopods in the
Himalayan-Yangtze area.
Before going further, I record mysincere thanks to Dr. N. KAMBE of theGeological Survey of Japan and Prof. I.
HAYAMI of the University Museum,

University of Tokyo for their kind assistances in preparation of this article.
Family Polydesmiidae KOBA YASH!, 1940
Diagnosis :-Shell medium sized, mostly longiconic, straight, but sometimes alittle curved or brevi conic, subcircularin cross section. Septum holochoanoidal
orellipochoanoidal,
cyrtochoanitic;
septal neck hook-shaped, but varying
from semicircular to abruptly recurved;
siphuncle subcentral to submarginal,
broad; siphuncular segments short; wallof the segment composed of a medianannulation and constrictions on its two
sides; radial canals highly oblique tosubrectangular to central canal; siphuncular deposits lamellar; stereo plasmic
deposit not well developed in camerae.
Remarks :-Polydesmia and Ordosoceras
are two typical genera of this family
which are well characterized by theannulated and somewhat shoulderedlateral projection and constrictions on
its two sides, highly oblique radial
canals and lamellar siphuncular~deposits.
Because the constrictions are reduced inlvfaruyamaceras peshanense, I have proposed klaruyamacerina for it as a subgenus of lvlaruyamaceras i. e. Polydesmia
in 1936. Yet the septal neck is hookshaped, though the hook is depressed
and the lamellar deposits and obliqueradial canals show its being a terminal
form of the family in parallel to ~heActinoceras-Armenoceras trend of specialization. Because no intermediate linkhas as yet been found between J'vlaruyamacerina and Polydesmia s. str., theformer would be better accepted for thetime being as· a distinct genus of thesame family.
CHAO et al. (1965) placed Discoactinoceras in this family, but judging fromthe septal form and siphuncular segment

•

/

680.

Ordosoceras in lehol

the genus was considered more allied to
the Huroniidae by SH!MANSKY (1962),
TEICHERT (1964) and others. In my
opinion it represents a new family, as
will be discussed in detail in near future.
Distribution:~ This is a small but very
distinct family in the early and (?)
middle Toufangian age in the Hwanghobasin.

Genus Polydesmia LORENZ, 1906
Synonym: Maruyamaceras KOBAYASHI, 1931

Diagnosis :~Conch straight or slightly
curved longicone or brevicone, subcircular in cross section; surface annulated in type-species; siphuncle large,
subcentral; septum apparently holochoanoidal; stereoplasmic deposit not welldeveloped in camerae. Siphuncular wallcomposed of a large annulation, medium
sized anterior hook or constriction andsmall posterior constriction triangle insection; radial canals strongly or moderately oblique; siphuncular deposit composed of radially disposed lamellae,
which grow typically from the proActinoceras stage to the Polydesmia
stage through the Actinoceras stage.
Type-species :~Polydes171ia canaliculata
LORENZ, 1906.
R emarlzs :~Shell is abruptly expanding
in P. shimamurai and slightly curved in
P. watanabei. In 1940 I have describedand discussed this genus in detail.
Since then several new species wereadded and new localities reported.
Beside the type-species the genus comprises now the followings including oneof provisional reference:
Armenoceras elegans ENDO, 1932
Liaoning
Nlaruyamaceras shimalllurai KOBAYASH!, 1931 North Korea

127

Afaruyamaceras watanabei KOBA YASH!,
1931 North Korea
lIlaruyamaceras sp. KOBA YASH!, 1931
North Korea
Polydesmia canaliculata magna KOBAYASH!, 1940 Liaoning
Polydesmia sp. KOBA YASH!, 1940 i. e.
Armenoceras elegans ENDO, (pars)
Liaoning
Polydesmia zuezshanensis CHANG, 1959
Ordos, Shantung (CHEN, 1976)
Polydesmia yilehetoensis CHANG, 1959
Ordos
Polydesmia 3 spp. A, B, C CHANG, 1959
Ordos
Polydesmia changkiuensis CHANG, 1960
Ordos
Polydesmia ? sp. CHEN, 1975
Qilianshan
Armenoceras elegans ENDO, 1932 (pars)in pI. 15, fig. 2 was identified with
Polydesmia canaliculata by KOBA YASH!
(1940). A specimen of this species from
Minshankou, near Penhsihu, Liaoning,Northeast China is a well preserved
phragmocone with camerate portion.
While they constitute a solid species,
P. canaliwlata manga is so isolated that
it is better recognized an independent
species.
SHIMIZU and OBAT A (1937)
reported Polydesmia from Hopei, but it
remained undescribed. Polydesmia listedfrom Jehol (KOBA YASH! and CHOH, 1942)is known now to be an Ordosoceras.
Polydesmia? sp. from a little known
locality in South Korea (KOBAY ASH!,
1966, p. 41) is here eliminated from thisgenus.
Distribution :~Insummarizingt.he
above occurrences, it can be said thatthe genus was widely distributed in
Eastern Asia in North Korea, Liaoning,
Shantung, Hopei?, Ordos and ? Kilienshan, all in the so-called Hwangho basinin the Ordovician palaeogeography.
Armelloceras elegans from the Middle

Teiichi KOBAY ASH!

128

Ordovician of east Siberia which apparently lacks constrictions on thesiphuncular segments and whose siphuncular deposits are not horn-shaped(BALASHOV, 1964), looks like an armenoceroid rather than a Polydesmia.
Genus Ordosoceras CHANG, 1959

Diagnosis :-Sim'ilar to Polydesmia, butdifferent principally in the siphuncularwall in longitudinal section which iscomposed of a large annulation completely separated from septa by two
constrictions the anterior and posterior
one of which are hook-shaped and linear
or subtriangular respectively.
Septa
apparently ellipochoanoidal ; camerae
with stereo plasmic deposits.
Type-species :-Ordosoceras splzaeriJorme
CHANG, 1959.
Remarks :-CHANG described beside theabove species the followings from Ordos :Ordosoceras quasilineatu11l CHANG, 1959
Ordosoceras sp. CHANG, 1959
Ordosoceras Izonguezense CHANG, 1960
Ordosoceras ordosense CHANG, 1960
Ordosoceras aff. splzaeriJorme CHANG,

Namely, the radial canals look nearlystraight and rectangular to the central
canal in the Himalayan ones. Theirsiphuncle is narrower and their siphuncular segments are taller in comparison with the breadth of the segment
at the constriction. In these aspects theHimalayan species are certainly closer
to Meitanoceras subglobosum from Central China (CHEN and Lru, 1974).
Whether or not they are congenericwith M. subglobosum is a question. I
think, however, that they are proper tobe included in the Meitanoceratidae
rather than the Polydesmiidae. None ofthem has the horn-shaped siphunculardeposits and oblique radial canals typicalof the Polydesmiidae.
Distribution :-Ordos, Shantung andJehol, western Liaoning, all in the
Hwangho basin of Eastern Asia, if theHimalayan forms are excluded.

Ordosoceras jeholense KOBAYASHI
sp. nov.
Plate i 7, Figures 1, 2; Plate 18,
Figures 2-4

1960

Ordosoceras splzaeriJorme okotiense
CHANG, 1960
Recently CHEN (1976) described O.
quasilineatum from Shantung, while
Ordosoceras jeholense, nov. is added tothem in this paper from Jehol.
CHEN (1975) described additional four
species from the Himalaya as follows:
Ordosoceras nyala11lense CHEN, 1975
Ordosoceras chiantsunense Tsou (MS)in CHEN, 1975
Ordosoceras contractum CHEN, 1975
Ordosoceras yaliense CHEN, 1975
These Himalayan species are similar
to the North Chinese species in the aspect of the siphuncular wall, but quite
different in the siphuncular structure.

Description :-Orthoconic phragmocone,
about 9 mm in length. in tan-coloured
limestone with scattered earthy patches.
Septal sutures on the weathered surfaceof the limestone are transversal, but
gently sinuated adapically in the medianpart of the ventral side, forming abroad ventral sinus like hyponomic sinus;
six sutures and five intervals in thelength of 20-22 mm. This camera waH
is broadly convex, but the convexity isstrengthened on lateral sides. Judging
from this curvature the conch seems to
be not elliptical, but suboval in crosssection having a fairly flattened venter.
The siphuncle is large, submarginal andapparently sub oval in cross section.

680.

01'dosoceras in Jehol

The phragmocone was cut longitudinally, but slightly oblique to the central
canal so that the canal disappears outof the section in the adoral half. In
the longitudinal section the conch appears to expand somewhat more rapidly
than the siphuncle. The central canalis located at about one-third the diameter of the siphuncular from thedorsal margin.
The stereoplasmic
deposit is well developed in the cameraeon the ventral side, although the outershell is gone. The camerate portion is
not preserved on the other side, because
the septa are thin and the camerae
were vacant. There the siphuncle was
exposed under erosion before the deposition of the limestone. The original
internal structure of the phragmocone
is, however, well preserved on theventral side.
The siphuncle, about 85 mm long, iscomposed of 19-20 siphuncular segments.
Three septa and two intervals arecountable in the length of 8 mm and 10
mm respectively in the adapical andadoral part. A siphuncular segment
consists of a median annulation andtwo constrictions on its both sides.
In the longitudinal section the septal
neck is hook-shaped, The connecting
ring describes therefrom almost threequarters of a circle and then straight
down obliquely, outlining a triangularcavity. The diameter of the circle ishalf as long as a septal interval. The
median annulation is protruded anterolaterally and isolated from the septa on
its two sides, but its distance from theseptum is shorter on the adoral sidethan the other, because the constriction
on the former side is smaller than on
the latter side.
The siphuncular deposit is quite different from the usual annulus.
It
encrusts largely the inside of the con-129

necting ring as far as the preceding
septal neck. The stereoplasmic deposition in this stage, however, cannot beanalyzed in detail with this specimen,
for growth lines are vanished by recrystallization. In the next stage thedeposit grows inward and upward i. e.
adorally to form an oblique horn which
is protruded as far as the level of thesucceeding septal neck. The growth of
this stage is clearly shown in section bya series of crescentic layers which
reveals the cone-on-cone growth of thehorn. In the adapical part of the section
it is seen that a horn is intercalated
between two horns showing the branching of radial canals.
In the camera the episeptal deposit ismore developed than the hyposeptal
one. The pseudoseptum terminates inward in juxtaposition to the end of aradial canal.
Comparison:- This species is allied to
Ordosoceras quasilineatum CHANG in thesiphuncular structure, particularly thedevelopment of the oblique horns, but
CHAl\G'S species is a much smaller form
which is quite distinct from this species
in the subcentral position of thesiphuncle, the conch expanding more
rapidly in comparison with the siphuncular expansion and the circular but
only slightly ovate cross section of theconch.
Occurrence :-Nantanchinkou area inthe southwestern part of the Nanpiao
coal-field, Jehol, Liaoning, Northeast
China. TOKuDA's collection from thearea contained Tojangocerina, Armenoceras, Orl7loceras (?), Polydes71lia andCal7leroceras' siphuncle beside Archaeosc),phia, Maclurites, Pagodispira andIsotelus or Isoteloides by preliminary
determina tion (KOBAYASHI and CHOB,
1942). They were procured from Ordovician limestones III some horizons.

Teiichi KOBAY ASH!

130

Among them the first to third cephalopods were already described under
the names of Tofangocerina nanpiaoense
KOBAYASHI and MATUMOTO, Jeholoceras
robustum KOBAYASHI and MATUMOTO
and Linormoceras centrale KOBA YASHI
and MATUMOTO (1942).
Jeholoceras is an orthoconic armenoceroid with a broad submarginal siphuncle
whose annuli are composed of verticallamellae protruding inward in different
length.
Linormoceras is an orthoconic ormoceroid with a large sub central siphuncle
whose stereo plasmic deposits consist ofannuli and endosipholinings. Linormoceras centrale minor was later describedby CHANG (1965) from the Yingou Groupof Yumen, Kansu.
These three cephalopods from Jehol
are presumably younger, but Archaeoscyphia must be older than the present
species.
Family Meitanoceratidae
KOBA YASHI, fam. nov.
Diagnosis :-Like Polydesmiidae siphuncular wall consists of middle annulation
and a constriction on each side, but
siphuncular segment is comparatively
taller and narrower; annulus has nohorn-shaped oblique projection; radial
canals extending from central canal subrectangularly. (See Meitanoceras subglobosum, pI. 18, fig. 1)
Remarks :-Four species of Ordosoceras
from the Chiatsun Group of the Himalayas (CHEN, 1975) belong probably tothis family, although they may begenerically distinguishable from Meitanoceras.
Distribution:- Llandeilian-Llanvirnian,Central China-Himalaya.

The age of the Maruyaman stage

When I proposed Marllyamaceras in1931, I considered it a late Wolungian
actinoceroid, because Afaruyamaceras
shimamurai, ]v!. watanabei and M. sp.
beside Cameroceras sp. were collectedfrom the Maruyama (Hwansan) limestone between the Onnoceras-bearing
Toufangian limestone and the CoreanoceJ'as-bearing Shorin Limestone in the
Kyeomip'o-Hwangju(Kenjiho-Koshu)
area, North Korea and because W ATANABE'S collection from Tungyiiehyuan,
Peshan-hsien, Shantung contained Marupamaceras peshanense, Stereoplasmoceras
d. machiakollense and ]v!anchuroceras
platyventrllm.
Subsequently SHIMIZU and OBATA(1937) pointed out that Maruyamaceras
did not occur together with Manchuroceras, but did the former in a higher
horizon than the latter in Peshan-hsien,
Shantung as well as the Liukiang coalfield, Hopei. They denominated theformation with the Wutinoceras zone
containing Wutinoceras, Maruyamaceras
and Armenoceras the Leichuang stage.
In the same year TEICHERT ascertained
the synonymy of Maruyamaceras with
Polydesmia.
Then I examined in 1940 the WATANABE collection again and confirmedthat the limestone slab containing
Manchuroceras platyventrum and Stereoplasmoceras d. machiakouense is different
from the Polydesmia peshanensis bearing
limestone in the lithological aspect.
Therefore I distinguished the Maruyaman
stage from the W olungian series and notedthat the stage is probably the equivalentof the Chazyan in North America ornear it, provided Armenoceras elegans
problem on which the reader is referred
to a later page.
Later in 1956 OBATA distinguished the680.

Orbosoceras in Jehol

upper or Wushan stage with Armenoceras(Paramenoceras) onukii and the lower orLeichuang stage s. str. with Polydesmia
kobayashi (nom. nud.) in the Leichuang
stage s. I. 1937 and correlated the two
combined to the Chazyan of North
America and the Llanvirnian of Europe.
In the mean time CHANG (1959, 1960)
found the following four cephalopod
zones in the Alashan-Chaotzushan area,
western Ordos.
II. Chaotzushan Limestone, 80-100 m
thick
3. Chisiloceras undlilatum zone in theupper part
2. Ordosoceras quasilineatum zone in
the lower part
1. Polydesnlia zuezshanensis zone III
the basal partI. Santaokan sandstone and limestone,
50 to 60 m thick
O. Wlltinoceras zone
In adding new facts CHEN (1976)
summarized the Ordovician cephalopod
biostratigraphy in North China in proposing Peianchuang suite for the unit
having Polydesmia and Ordosoceras asits keys. This unit is underlain by the
Liangchiashan Limestone
contallllllg
Coreanoceras and NJanchuroceras. The
former is the approximate correlative of
the Dawan suite and the latter that of
the Hunghuayuan in Central China where
the latter also yields Manchuroceras andCoreanoceras (CHEN and LIU, 1974). In
Central China the international correlation can be made on the basis of rich
graptolites, trilobites and others. There
the Hunghuayuan Limestone is dated at
the Arenigian exclusive of the Azygograptus sllecius zone and the Dawan
suite at the Llanvirnian in addition to
the suecius zone (Mu and CH'EN, 1962,
Lu, 1975).
Because the Peianchuang
suite is the Polydesmia-Ordosoceras age,131

the age determination of the Maruyaman
stage at the early Chazyan or theWhiterock is now thoroughly vindicatedby these studies.
Incidentally, FLOWER, (1976a, b) showsin his correlation table that the Whiterock of North America is coeval with
the Maruyaman, Wuting and Kangyao
Formations in descending order, but theHystricu1'Us faunule of the Wuting fauna
is Wolungian whereas some other Wuting
fossils are pre-Kangyao or the Maruyaman in age, because the Kangyao ispost-Wuting according to ENDO'S emended
sequence, 1953a, b.
Polydesmia and Ordosoceras constitutean early off-shoot of the Actinoceratida.
They are, however, not the oldest of
the group in Eastern Asia. As mentioned
above, Wutinoceras appeared before themin Ordos.
In Chinghai Wutinoceras
shihuigollense CHANG and Pararmenoceras
sp. 1 are reported together with Manchuroceras tochllanshanense CHANG fromthe Tochuanshan Limestone at Shihuigou(CHANG, 1965). The association of thesethree cephalopods indicates the transition
from the Wolungian manchuroceroid age
to the Toufangian actinoceroid age.
Now the age of Armenoceras elegans
is to be discussed. ENDO (1932) described
this species from the lower fossil horizonof the Ssuyen Formation at Penhsihu
and Niuhsintai. He correlated the formation to the Black River of North
America. This species was, however,referred to l\1aruyamaceras i. e. Polydesmia by TEICHERT (1933, 1937), KOBAY ASHI (1934, 1936, 1940) and SHIMIZU
and OBAT A (1936). Therefore its reference to the Polydesmiidae, if not Polydes17lia, is undeniable.
ENDO (1932) divided the Ordovician
rocks inclusive of Ozarkian ones in South
Manchuria into the Pingchou, Santao,
Kangyao, Wuting and Ssuyen Formation

132

Teiichi KOBAYASHI

in ascending order. The first and second
units correspond to the Wanwanian andW olugnian Series (KOBAYASHI, 1931a, b,1969).
He correlated the Kangyao,
Wuting and Ssuyen Formations to theStones River, Blount and Black River
respectively. Later (1953a, b), however,
he replaced the Wuting below the
Kangyao Formation by the reason that
the Wuting fauna includes some Canadian elements (KOBAYASHI, 1935, 1966b).
According to ENDO (1932) the Kangyao
fauna rich in Lophospira and othergastropods is older than the Ssuyen
fauna rich in Armenoceras and othercephalopods. The type-localities of the
Kangyao and Ssuyen Formations are,however, in separate basins of Yentai
and Penhsihu, this status making difficult
to verify the superposition of the latteron the former. On the contrary, thecoexistence of such cephalopods withvarious gastropods including LopllOspirais well known in the Machiakou Limestone in North China (GRABAU, 1922) aswell as the Unkaku bed at Sunghoni(Shokori) near P'yeongyang, North Korea(KOBA Y ASHI, 1929, 1930). Therefore I
think it quite probable that the difference of the faunal composition between
the Kangyao and Ssuyen Formations
depends much upon their facies. In
other words, they are heteropic frombut synchronous with each other at least
to a certain extent. Nevertheless theoccurrence of Armenoceras elegans in
the Kangyao-Ssuyen suite indicates its
being a post-Maruyaman survivor of the
Polydesmiidae.
Finally, a few words are added on

this occasion about Nybyoceras foerstei
ENDo, 1932, the type-species of Wutinoceras SHIMIZU and OBAT A, 1936, fromblack banded limestone of the so-called
Wuting Formation near Santaokangtzu
near Penhsihu.
I presume that theforestei's holotype was derived from alittle higher horizon than the earthy
limestone at San tao type-locality on thesecond railway tunnel of Santaokangtzu.
However, this forestei horizon must be
already in the Maruyaman stage, namely, the Peianchuang of North China, andthe Leichuang-Wushan suite of Hopei.
References
BALASIIOV, Z. G. (1964): Some nautiloids,
endoceroids and actinocerids from theOrdovician of Northeast of the USSR.
Sci. Inst. Ceol. Arctic Nat. Ceol. Comm.
USSR, Yol. 6, 71 pp, 6 pI.
CHANG Jih-tung (1959): Lower Ordovician
cephalopods from Chaotsushan, Ikechaomeng, Inner Mongolia. Acta Pal. Sinica,
vol. 7, no. 4, p. 259-283, pI. 1-6.
- - (1960): The stratigraphic relation of
Poiydesmia and its close relation with
Ordosoceras. Ibid., vol. 8, no. 1, p. 4153, 3 pI.
- - (1965): On some Lower Ordovician
nautiloids from Qilianshan, Northwestern
China, Ibid., vol. 13, no. 2, p. 343-357, 2
pl·
CHAO Chin.ke, LIANG Hsi-ho, Tsou Hsi.p'ing,
LEI Ts'ai-kem and CHA1'G Jih.tung (1965) :Chinese cephalopod fossils. 389 pp. 85 pIs.
Each Group of fossils in China.
CHEN Ju-yuan (1975): Nautiloidea. Tibet
Scientific Research Party: Reports of
Scientific Research of Chumulangmafeng,
Palaeontology, vol. 1, p. 267-294, 9 pIs.

Explanation of Plate 17
Ordosoceras jehoiense KOI3AYASHI, sp. nov.
Figures 1, 2. Two longitudinal polished sections, x 1.5.
Museum, University of Tokyo (PM389-1)

Holotype stored in University

KOBAY ASH!:

Ordosoceras in Jehol

Plate 17

680.

Ordosoceras in Jehol- - (1976): Advances in the Ordovician
stratigraphy of north China with a brief
description of nautiloid fossils.
ActaPal. Sinica, vol. 15, no. 1, p. 55-73, pI.
1-3.
- - and LIU Ching-wLl (1974): Ordovician
nautiloidea.
Nanking Inst. Geol. Pal.
Acad. Sinica: Handbook of Geology andPalaeontology of Southwest China, p.
138-143, pI. 58-62.
ENDo, R. (1930): The presence of Nybyocerasin South Manchuria. Denison Univ. Bull.
jour. Sci. Lab., vol. 30, p. 298, pI. 60.
- - (1932): The Canadian and Ordovicianof Southern Manchuria. U. S. Nat. Mus.
Bull., 164, 152 pp. 40 pI. map.
- - (1935):
Additional fossils from the
Canadian and Ordovician of the southernpart of Manchoukuo. Sci. Rep. Tohoku
Imp; Univ., Ser. 2, vol. 16, no. 4, p. 191223, 6 pIs.
--~ (1953a): A summary of the columnar
section in Manchuria.
Proc. Seventh
Pacif. Sci. Congr., vol. 2, Auckland &Christchurch, p. 316-326.
- - (1953b): A summary of the geologicalsection in Manchuria. Sci. Rep. Saitama
Univ., Sec. B, vol. 1, no. 2, p. 105-114.
FLOWER, R. H. (1957): Studies of the Actino·
ceratida, 1. The Ordovician developmentof the Actinoceratida.
New Mexico
Bureau Mines Min. Res., Mem. 12, 79 p.
6 p\.
- - (1968): The first expansion of theActinoceratida. Ibid., Mem. 19, pt. 1,
p. 1-16, pIs. 1-30 (pars).
- - (1976a): New American Wutinoceratidaewith review of actinocerid occurrence illeastern hemisphere. Ibid., Mem. 28, part1, p. 5-12, pIs. 1-5.
- - (1976b): Ordovician cephalopod faunas
and their role in correlation. BASSETT,
M. G. (ed.) The Ordovician System. Proc.
Palaeont. Ass. Symposium, Birmingham,
1974. p. 523-552.
GRABAU, A. W. (1922): Ordovician fossils
from North China. Palaeontol. Sinica,ser. B, vol. 1, fasc. 1, 100 pp. 9 pIs.
KOBAYASHI, T. (1929; ::\0):
Ordovician
Fossils from Corea and South Manchuria,

133

Parts 1 & 2. japan. jour. Ceol. Ceogr.,
vol. 5, p. 173-212, p\. 18-22, vol. 7, p.
75-100, p\. 8-11.
- - (1931a): Studies on the Ordovician
stratigraphy and palaeontology of north
Korea with notes on the Ordovician
fossils of Shantung and Liaotung. Bull.
Ceol. Soc. Chosen (Korea)", vol. 11, no. 1,
p. 60 pp. 9 pIs.
- - (1931b): Studies on the stratigraphy
and palaeontology of the Cambro·Ordo·
vician formation of Huo·Lien-Chai and
Niu.Hsin-Tai, South Man,churia, japan.
jour. Ceol. Ceogr., vo\. 8, p. 131-189, pI.
16-23.
- - (1934): The ·Cambro-OrdoYician For·mations and faunas of South Chosen,Part 1, Middle Ordovician Faunas. jour.
Fac. Sci. Imp. Univ. Tokyo, sec. 2, vol.
3, pt. 8, p. 329-519, pI. 1-44.
- - (1935): A question on the Wuting
fauna of Manchuria. jour. Ceol. Soc.
japan, vol. 42, p. 114-121.
- - (1936): An introduction to the classi·
fication of the primitive cephalopods.
Chikyu, the Clobe, vo\. 25, p. 11-27, 85103, 180-192, 271-281, 331-354.
- - (1940):
Polydesmia, an Ordovician
actinoceroid of Eastern Asia.
japan.
jour. Ceol. Ceogr., vol. 17, nos. 1-2, p.
27 -44, pI. 3-5.
- - (1966a): The Chosen Group of South
Korea. jour, Fac. Sci. Univ. Tokyo, sec.
2, vol. 16, pt. 1, p. 1-84.
- - (1966b): The Chosen Group of North
Korea and Northeast China. Ibid., vol.
16, pt. 2, p. 209-311.
. - (1969): The Ordovician of Eastern Asiaand other parts of the continent. Ibid.,vo\. 17, p. 163-316.
- - (1977): Manchuroceras found in South
Korea with notes on the Manchuroceratidae and the Manchuroceras province.
Trans. Proc. Pal. Soc. japan, N. S. no.
105, p. 17-26, p\. 3, 4.
- - and CHOH, R. (CIIA='iG Li-Sho) (1942):
Fossil zonation of the Cambro·Ordovician
Chosen Group in eastern Jehol. japan.
jour. Ceol. Ceogr. vo\. 18, no. 4, p. 213-224.
- - and MATUMOTO, T. (1942): Three new134

Teiichi KOBAYASHI

Toufangian nautiloids from eastern Jehol.
Ibid., vol. 18, no. 4, p.313-317, pI. 30-31.
LORENZ, T. (1906): Beitrage zur Geologie
und PaJaontologie von Ostasien unterbesonderer Beriicksichtung der Provinz
Schantung in China, II. Zeitschr. deutsch.
geo!. Gesell. Bd. 58, S. 67-122, Taf. 4-6,
Kartenskizzen.
Lu, Yen-hao (1975):
Ordovician trilobite
faunas of central and southwestern China.
Palaeontol. Sinca, N.S. B, no. 11, 463 pp,
50 pI.
Mu, A. T. and CH'EN Hsii (1962): Chinese
Graptolites. 171 pp. 21 pIs. Each groupof Chinese fossils.
OBATA, T. (1956):
Ordovician nautiloid
zones in North China.
Bull. Aichi
Gakugei Univ. (Nat. Sci.) vol. 5, p. 50-56.
SHIMIZU, S. and OBATA, T. (1936): Studies
on the Palaeozoic cephalopods from Asia,
Part 1. Jour. Shanghai Sci. Inst. sec. 2,
Yol. 2, p. 85-146.

- - and - - (1937): Ordovician stratigraphy
in eastern Hopei, China. Proc. Imp. Acad.
Tokyo, vol. 13, p. 107-109.
SHI~-1ANSKY, V. N. (1962): Actinoceratoidea.
p. 207-224, pI. 1-3 in ORLOV'S Principles
of Paleontology, tom 5.
TEICHERT, C. (1933): Der Bau der Actinoceroiden Cephalopoden. Palaeontogr. Bd.
78, Abt. A, p. 111-230, pIs. 8-15.
- - (1935):
Structure and phylogeny ofactinoceroid cephalopods. Am. Jour. Sei.
ser. 5, vol. 29, p. 1-23, 5 figs.
- - (1937): Polydesmia canaliculata LORENZ,
an Ordovician actinoceroid cephalopod.
Jour. Geo!. Soc. Japan, vol. 44, p. 709-713 ;Trans. Proc. Palaeont. Soc. Japan, no. 8,
p. 110-113.
- - (1964):
Actinoceratoidea.
In R. C.
MOORE (ed.) Treatise on Invertebrtae
Paleontology, Part K, Mollusca 3. p. 190216.

Place names in China (C) and Korea (K)Alashan ~iJMilJ (C),
Chaotzushan, Zuezushan t!/!-rilJ (C),
Chiatsun JtIti (C),ChinghaiWW (C),
Changkiu J$:j~ (C),
Dawan x'f91 (C),
Honguez fuI1B-r (C),
Hopei tilJ~t(C),
Hunghuayuan nTEIlm (C),Hwangho mtnr (C),Hwangju, Koshu jl1f1+[ (K),
Jehol l!MilJ (C),
Kansu i:tEk1 (C),Kanyao (![~ (C),
Kilienshan, Qilianshan jj[~:liliLU(C),
Kuniutan qtitfilli (C),Kyeomip'o, Kenjiho 3lit=lr1l (K),Leichuang 'uHf (C),Liangchiashan ?t~HU (C),
Liaoning ~$J (C),Liukiang WillI (C),
Machiakou .~*Ifij (C),Maruyama 1LilJ (K),Minshankou g}jilJlill' (C),Nantanchinkou MiJ\!l>'iZ~ (C),Nanpiao Wi;!]? (C),Niuhsintai tf-L.'iJ' (C),Nyalam 'it'l.* (C),Oketi ~Btt:£1iIT (C),
Ordos ~B ~f.~:I\Ii (C),Peianchuang ~tfftJ± (C),Peipiao ~t~ (C),Penhsihu *~¥1jI](C),
Pingchou 1'-1+[ (C),Peshan-hsien, Poshan jtlfilJ (C),P'yeongyang 1'-~ (K),Santaokan :=Jil:!X (C),Santaokangtzu :=Jilr.m-r (C),Shantung ilJJR (C),Shihuigou
EFR.iflf (C),Shorin-ri, Songnim-ni t~:ttm. (K),
Ssuyen!!B1lE! (C),Sunghoni, Shokokri!l1HIiflill. (K),Tochuanshan g-,jRilJ (C),
Toufangkou 5IJ511fij (C),
Tungyiiehyang JRHf
~ (C),
Unkaku-ri, Unhak-ni ~1'f.!HB. (K),Vieheto 1Jffj,[~~1 (C),
Wutingshan :n:fflilJ(C),
Wushan JiJ:;lll (C),Yali!!Eill. (C),Yentai jil/jiJ' (C),Yingou [1~i"li (C),
Yumen .=E["J (C)Explanation of Plate 18
Meitanoceras subglobosum CHEN and LID, 197~
Figure 1. Part of siphuncle in longitudinal section showing siphuncular sediments;
drawn from CHEN and LIlJ, 1974, pI. 60"fig. 7. x4.
Ordosoceras jeholense KOBAYASHI sp. nov.
Figure 2. Ventral view of the holotype, showing septal sutures. x 1.5.
Figures 3, 4. Part of longitudinal section of the holotype. (4: diagrammatic) x 3.

KOBA Y AS H! :Onlosoce?'as in J ehol

Plate 184

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 107, pp. 135-160, pis. 19-22, September 15, 1977

681.

TAXONOMIC STUDIES ON SOME FOSSIL AND
RECENT JAPANESE BALANOIDEA (PART 1)
TOSHIYUKI YAMAGUCHI

Geological Institute, Faculty of Science, University of Tokyo, Tokyo, 113<

13 ;$:o)l' --:>t;>O){r.1'jt; J: Lfm~ 7:; ';I -GmO)7tmctEt:Jlilf~ (-to) 1): {r.1j7:; ';I-Gm(;t 13;$:0) ~~=*,cJ.;JIlff:O)JtHl('")t;d?-Jtll1:r::.'iiltO)f'fG(f GK·(:tf;hJ: lilZJt V(.@:t±l-r-Qo -tn
G O)lilZJtt;> G ~;$:O):3t{$:(~uHE[JC c;n, m~~;$: 0) 7I-'ilIljf~~, 'iilttil\'mO)l:U~i:::' J: I) :fit;;/nJ
IE c; t:.o Lt;, Lm~ fil!O)vt*0)7t1~!H:::.(:t, :fit; J: Lf:fil*J O)7tmi:::.:;k ~ t.dlHLt;:;l;"? t:.o -tO)::ttoeJmEl:n:t1i1!t) TO) 7tmMt;;7f;)l~(j~zf:~O)~1!£t-:vti:::' i1i!;-:ll '-Ciilf~*O) ::tWH:::. J: "? -C 5Hil{c;n -C l ,t:.;:' i:::.;l; QO)~)C('(;t,13 ;$:0){r.1jt; J: Lf m~ 7 :; ';I -GmO)::t~toef.]M""(·;l;Q Balanus amphitrite !l' It.- - :it;;m~~¥o0)7}'ilIl7f;~, 'iilt;fi\\'mt; J: Lfik(;j;'ilIlO)7f;1i.~B~t#V&

c

n

c

i:::.JJ~*- -C,

0

JtH:fl!W~7t;ffj,c;:.

1:,,®,lliXtoe G Lfi:::.f4MII;!fWJtoe D:::.;Ij~-:ll' -C 4 1i1!i:::.7tm c; n t:.o it:.-t
c I:::' J: "? -C 1i1!7t{r.~ 13 ;$:"'0) lHI:O)II;'fJWI.i:n G O)n\.mM0)1tl'-;gC~-a:- J\"'}(ll.ctrl~i:::.iMg.];-r- Q ;:.
f!Jt~t c;n t:.oU~ r:J fiz

1968; 1970).
Studies on fossils were also started
by DARWIN (1851b, 1854b) and advanced
by SEGUENZA (1873-76), ALESSANDRI(1906, 1907a, b), PILSBRY (1918, 1924,
1930), WITHERS (1923, 1924) and others.
Recently, however, paleontologists havebecome increasingly interested in the
study of appendage morphology, anatomy, ecology, etc. Thus the boundary
between neontology and paleontology
has become more and more obscure.
NEWMAN, Ross, ZULLO, and others areadvancing the study of Balanoidea in
uniting neontological and paleontologicalstudies.
The japanese fossil Balanoidea havenot been studied in detail, though they
are commonly found in association withmolluscan fossils in the Neogene andQuaternary formations throughout japan.
In fact, "Balanus sPp.", found at theend of faunal list of fossil molluscs, isusually the only mention of the Balanus

1. Introduction

DARWIN'S (1851a, 1854a) two monographs on living cirripeds are classics
and are dependable even up to thepresent time. Following DARWI~, fromthe late 19th to the early 20th century.
the collections of biological expeditions
and other museum c(}llections werestudied by WEL TNER (1897), HOEK (1883.
1913), KRUGER (1911a, b), PILSBRY (1911,
1916, 1927), NILSSON-CANTELL (1921, 1932,
1934,1938), BROCH (1922, 1931), and others.
Most of these studies include taxonomic
descriptions of Balanoidea from japan
and adjacent seas. Yet, a systematic
description of Recent japanese Balanoidea
depends much upon a series of taxonomic publications made by UTINOMI (HIRO
prior to 1939) (HIRo, 1932a, b; 1933,
1935; 1937; 1938; 1939a, b; UTINOMI,
1949a, b; 1954; 1955; 1958; 1962; 1967;* Received May 26, 1977; read Jan. 31,
1976 at Kawatabi.
135

136

Toshiyulci YAMAGUCHIin stratigraphical or paleontologicalpapers.
Studies of Japanese fossil
Balanoidea which have been publishedare summarized in YAMAGUCHI (1971).
Species descriptions are frequently inadequate however so that it is difficult
to determine whether or not these Japanese species have been correctly
identified.
In this study, the writer attempts torefine the classification of some important species of Japanese living Balanoidea
on the basis of distribution, ecology,
habitat, biochemical characters andmechanism of reproductive isolation as
well as morphological features. That is,
the study has been conceived with interspecific and infraspecific relations between living populations. From thisvantage point it has been possible tomore realistically interpret the fossil
evidence.
2.

Methods of Study

Sampling and preparation of fossil specimens

Fossil barnacles are commonly foundas fragmented skeletal remains. However, they are often attached to bankforming oysters, other molluscan shells,
and to solid materials in mud and sandstone. Therefore, prior to preparing a
fossil specimen for study, the mode ofoccurrence should be observed in detail.
Besides general observations on thenature of the sediments, note of theassociated species is important for
paleoecological considerations.
This
holds especially true in the case of the
autochthonous occurrence of associatedspecies (cf. B. rostratus). Knowledge ofactual physical attachment between two
closely related species is also indispensable for demonstrating microgeographical sympatry between two species (cf.

Megabalanus rosa and M. volcano).

To obtain fossil samples from loose
sediments, they are sieved through a 1
mm or 2 mm mesh. The barnacles arethen picked up from sieved remains.
When articulated compartments are
obtained, the interior cavities shouldcarefully be examined to see if theopercular valves are present. If thespecimens are buried in a matrix ofhard sand or mudstone, they can beremoved with hand tools. Specimens
separated from a matrix can then becleaned ultrasonically. Besides ordinary
examination technique, X-ray photography can provide information oninternal microstructure of compartments.
Laboratory techniques for living specimens

For most field observations and laboratory techniques, the writer followed
those explained by NEWMAN, ZULLO andWITHERS (1969). In addition, for samples
to be prepared for the scanning electron
microscope (SEM), appendages weredissected in 70% alcohol. Each appendage with then washed in an ultrasoniccleaner and then put separately into asmall capsule with a net at each side.
The alcohol was then gently replacedby distilled water, the appendage frozen
in liquid N2 and freeze-dried in a vacuum
evaporator. Each appendage, placed on
the specimen holder, was then coatedwith carbon and gold-palladium (Au 60%,
Pd 40%), and observed under the SEM.
Depository, abbreviations and terminology

Specimens are deposited in the collection of the University Museum, University of Tokyo (UMUT). Registration
numbers are accompanied by prefix CA
for Cenozoic Arthropoda and RA for
Recent Arthropoda. Dissected appendages are mounted on glass slides, in theglycerin jelly and sealed with finger-nailpolish. Other living specimens are kept

681.

Taxonomic Studies on Japanese Balanoidea

in 70% alcohol. Morphological terms in
the present study are the same as thosein NEWMAN, ZULLO and WITHERS (1969),
HENRY and McLAUGHLIN (1975), and
NEWMAN and Ross (1976).
In the present study, the writer followed
a new classification of Balanomorpha
proposed by NEWMAN and Ross (1976)
based on comparative studies of surficial
morphology, internal wall structure,
trophi, chaetotaxis and intromittant
organ.
3.

Locality Register (Text-figure 1)

Living specimens:
A-Rocky shore (44°3.2' N., 144°16' E.) near
Futatsu-iwa, Abashiri City, Hokkaido.
B 1-3-Dredge samples; Funka Bay, Hokkaido: B 1-(42°6' N., 140°48.1' E.); gravel;
40 m in depth., B 2-(42°15.4' N., 1:W048.1'
E.) ; mud containing gravel and pumice;
74m in depth., B 3-(42°21.7' N., 1400~1.4'E.) ; mud; 53 m in depth.
C 1-3-Mutsu Bay, Aomori Pref. : C I-Dredge
samples (42°5' N., 140°49.4' E.) ; mud; 60
m in depth., C 2-Kanita (42°2.-1' N.,
140° 49.4' E.) ; on scallops Patinopecten yessoensis in the CUltivating cages. ColI.
Drs. K. ClIINZEI & Y. IWASAKI., C 3Rocky shore (45°54.2' N., 140°51.4' E.)
near Asamushi Marine Biological Station.
D-Dredge samples (39°20.6' N., 141°56.4' E.) ;Otsuchi Bay, Iwate Pref.; mud; 35 m indepth. Coll. Mr. E. HONZA.
E-Dredge samples off Tokai-mura, Ibaragi
Pref. ColI. Prof. N. NAsu.
F-Dredge samples, same as Loc. B-2 of
HORIKOSHI (1962), Tokyo Bay. ColI. Prof.
M. HORIKOSHI.
G-On the test pile (34°44.2' N., 137°38' E.)
submerged from Nov. 1973 to Aug. 1974,
Hamana Lake. ColI. Dr. T. KAZIHARA.
H-Rocky shore (33°40.6' N., 135°21.8' E.) at
Reisen bridge in the Tachigatani Inlet,
Tanabe Bay, Shirahama, Wakayama Pref.
Apr. 10, 1974.
.
I-Tanoshiro (34°34.5 1 N., 135°1.5' E.), Iwaya,
Awajishima Is., Tsuna-gun, Hyogo Pref.

137

samples: J I-Osaka Bay(34°29.5' N., 135°8' E.); mud; 38 m indepth.,
J 2-South of Tomogashima
Channel (34° 13' N., 134°57' E.); mud; 66
m in depth.

J 1-2-Dredge

Fossil specimens:
I-Horoshin-Tachibetsu: a cliff (43°49.7' N.,
141 °54.2' E.) of right bank of the HoroshinTachibetsu river, situated about 650 m
S600W of Matsuppu Station, Numata,Uryu-gun, Hokkaido; light gray fine sand;
Pliocene Horoshintachibetsu formation(OHARA, 1966) ; ColI. Dr. Y. I\\·ASAKI.
2a-d-Setana: a) a cliff (42° 44.5' N., 140°
24.9' E.) of right bank of the Kaigarasa wa
river, situated about 4.8 Km S68°W of
Mena Station, Ran-etsu, Isoya-gun, Hokkaido; conglomeratic coarse sand., b) a cliff
same as Loc. 12, YA\IAGUCHI, 1973., c) a
cliff (42°28.5' N., 140°11.8' E.) of left bankof the Toshibetsu river, situated about
950 m S48°W of Pirika Station, Pirika,
Setana-gun, Hokkaido; pumiceous coarse
sand., d) a cliff (-12°26.8'N., 140°4.6' E.)of left bank of the Tanekawa river,situated about 3.3 Km N5°E of Tanekawa
Station, Hikaridai, Set ana-gun ; coarse
arkose sand. a-c) The Pliocene Set ana
formation (IKEYA & UEMATSU, 1968). d)
The late Miocene Yakumo formation(IKEYA & UEMATSL-, 1968).
3a-f-Shimokita: a-d) cliffs same as Locs.
13a-d, YAMAGUCIII, 1973., e) a cliff(-H ° 11.0' N., 141 ° 16.6' E.) same as Loc. 6,
HATAI, et al., 1961; pumiceous silty sand.,f) a cliff (41°10.0' N., 14P16.7' E.) of leftbank of the mouth of the Maekawa
river, Nakanosawa, Mutsu City, Aomori
Pref.; pumiceous very coarse sand. The
Pliocene Hamada formation (YAMAGUCHI,
1970).
4a-c-Tsugaru: a, b) cliffs same as Locs.
11a, b, YA\lAGl'CHI, 1973., c) a cliff(45°55.4' N., 140°37.7' E.) of right bank of
the Rokumaibashi river, situated about 3
Km S77°W of Ushirogata Station, Rokumaibashi, Aomori City, Aomori Pref.; coarse
sand.
The Pliocene Kanita formation(UDll'RA, et al., 1959).

138

Toshiyuki YAMAGUCHI
w

°0

...

w

.,.,

...
45°'"

40°'"

• Holocene• Pleistocene... Pliocenew• l. Miocene

M

."

13~

• E.-M. Miocene

Text.fig. 1. Sampling localities of fossil and living Balanoidea of Japan.
5a-b-Daishaka: a) a cliff(40°49.8' N.,
140°38.5' E.) of right bank of the Amadai
river, situated about 2.9 Km N800W of
Tsugaru-Shinjo Station, Shinjo, Aomori
City, Aomori Pref.; medium sand., b) a
cliff same as Loc. 15, YA~IAGUCIIl, 1973.
The Pliocene Daishaka formation (I \\"A I,
1965) .
6a-c-Kamikita: a) a cliff (40° 43.5' N., 141 °17.3' E.) same as Loc. 6, IWAI & SIOBARA,
1969; massive mud., b) a cliff (40°43.6' N.,
141°16.1' E.) same as Loc. 9, IWAI &SIOBARA, 1969; massive mud., c) a cliff(40 043.4'N., 141°14.6' E.) same as Loc.Il,
I\\"AI & SIOBARA, 1969; massive mud.
The Pleistocene Numasaki mudstone.
7a-n-Mabechi river valley: a) a road-cut(40'25.3' N., 141° 16.8' E.) situated about681.

Taxonomic Studies on Japanese Balanoidea

3.1 Km N4°E of Sannohe Station, Sannohe·
gun, Aomori Pref.; massive medium sand·stone., b) a cliff (40°25.1' N., 141°17.7' E.)
situated about 1.4 Km N6SoW of Suwano·taira Station, Sannohe.gun; conglomeraticcoarse sandstone., c) a road·cut (40°24.9'
N., 141 ° 16.S' E.) situated about 2.4 Km
N6°E of Sannohe Station, Sannohe.gun;cross·laminated coarse sandstone contain·
ing gravel., d) a cliff (40°24.4' N., l·W
15.S' E.) of right bank of the Sarube
river, situated 1.9 Km N400W of Sannohe
Station, Sannohe.gun; barnacle limestone.,e) a cliff (40°24.1' N., 141"17.1' E.) ofright bank of the Mabechi river, situatedabout 1.1 Km N37°E of Sannohe Station,
Sannohe·gun; ill·sorted coarse sandstone.,f) a cliff (40022.S' N., 141' 15.5' E.) situatedabout 2 Km S34°W of Sannohe Station,
Sannohe·gun; conglomeratic coarse sandstone., g) a road-cut (40°22.7' N., 141 °16.3'
E.) situated about 1.7 Km S16°W of Sannohe Station, Sannohe-gun; shell limestone
containing gravel., h) a cliff (-!00 lS.l' N.,
141 ° lS.7' E.) of right bank of the Mabechi
river, situated about 2 Km S100E of Kintaichi Station, Ninohe·gun, Iwate Pref.;ill-sorted conglomeratic coarse sandstone.,i) a road-cut (40°17.7' N., 141°17.5' E.)
along the branch of the Jumonji river,situated about 1.6 Km N14°W of Tomai
Station, Ninohe-gun; conglomeratic coarseshell sandstone., j) a cliff (40°17.5' N.,
141"17.5' E.) of left bank of the Jumonji
river, situated about 1.3 Km N1SoW of
Tomai Station, Ninohe-gun; barnacle limestone and calcareous medium sandstone.,k) a cliff (40'7' N., 141 'S' E.) of left bankof the Mabechi river, situated about 300
m N53°E of Tomai Station, Ninohe-gun;
ill·sorted
silty
sandstone
containing
gravel., I) a cliff (40°16.1' N., 111°1S.2' E.)of right bank of the Mabechi river,situated about 1.7 Km N45°E of Kitafukuoka Station, Ninohe-gun; very coarse
sandstone., m) a cliff (40° 15.1' N., 141 °lS.6' E.) of left bank of the Shira tori
river, situated about 1.9 Km N5soE of
Kitafukuoka Station, Ninohe-Jun; illsorted conglomeratic silty sandstone., n)139

a road-cut (40° 14.3' N., 141 ° lS.4' E.) of theNamiuchi Pass, situated about 2.3 Km
S35°E of Kitafukuoka Station, Ninohe-gun;cross·laminated pumiceous sandstone. f)
The Pliocene Kamimetoki alternation(CHINZEI, 1966) of the Tomesaki formation. a) The late Miocene Miyasawa
sandstone (CIIINZEI, 1966) of the Tomesaki formation. d, g, i & j) The late
Miocene Metoki shell limestone (CHINZEI,1966) of the Tomesaki formation. b, c &e) The late Miocene Shinden sandstone(CHI:,\ZEI, 1966) of the Suenomatsuyama
formation.
k & n) The late Miocene
Maisawa sandstone (CHI]\"ZEI, 1966) of the
Suenomatsuyama formation. I) The late
Miocene Aikawa andesite (CHINZEI, 1966)of the Suenomatsuyama formation. h &m) The late Miocene Anaushi conglomerate (CHI:,\ZEI, 1966) of the Suenomatsuyama formation.
Sa-c-Oga: a) a coastal cliff (39°5S.3' N.,
139'51.1' E.) situated about 300 m SW ofAnden, Oga City, Akita Pref.; cross-laminated coarse to medium sand., b) a cliff(39°5S.2' N., 139°51' E.) of coast situatedabout 450 m SW of Anden; cross-laminated and ill·sorted conglomeratic sand.,c) a coastal cliff (39'5S.1' N., 139°50.9' E.)
situated about 550 m SW of Anden; coarseshell sand. a) The Pleistocene Anden
formation (KITAZATO, 1975). b & c) The
Pleistocene Shibikawa formation CRUZlOKA, 1959).
9a-d-Ohmagari: a) Shell quarry at Arasawa(39°24.3' N., 140°21.1' E.), Minami-Sotomura, Senpoku-gun, Akita Pref.; barnacle"coquina"., b) a cliff (39°21.6' N., 140'21.7'
E.) same as fossil locality at Asizawa
Pass of the Takinoue of OTUKA, 1936;
massive ill-sorted tuffaceous sand containing carbonaceous matters., c) a cliff (39°22.1' N., 140°21.7' E.) of left bank ofrivulet situated about 1 Km north of Loc.
9b; massive ill-sorted tuffaceous fine sandcontaining carbonaceous matters., d) a
cliff (39'21.7' N., 140°21.4' E.) of rightbank of riyulet, situated about 400 m westof Loc. 9b, Yasawagi, Riraga-gun, Akita
Pref.; massive ill-sorted tuffaceous fine140

Toshiyuki YAMAGUCHIsand containing carbonaceous matters.
The early to middle Miocene Sugota formation (KATAYAMA, 1941).
10-Ukibuta: a cliff (39°1S.S' N., 140°20.7' E.)
same as fossil locality at Ukibuta of
OTUKA, 1936, Higashi-Yuri-mura, Yuri-gun,
Akita Pref.; ill·sorted tuffaceous medium
sand; early to middle Miocene Sugota
formation (KATAYAMA, 1941).
ll-Ichinoseki: a cliff (3s055.7' N., 141°6.S'
E.) of left bank of the Iwai river, situated about 2.5 Km NSOoW of Ichinoseki
Station, Ichinoseki City, Iwate Pref.; illsorted medium sand containing many
pyroclastic rock fragments; middle tolate Miocene Shimokurosawa formation(ONODERA, 1957).
12a-b--Sendai: a) a cliff (3S0 15..:!' K., 140°
49.6' E.) same as Loc. 1 of the Tatsunokuchi formation, HA!'iZA\VA, et aI., 1953;
tuffaceous medium sand containing many
quartz grains; Pliocene Tatsunokuchi formation (HA~ZAWA, et at., 1953)., b) a
cliff (3S0 12.S' N., 140°47.5' E.) situatedabout 200 m south of the Oide bridge
across the Natori river, Moniwa, Sendai
City, Miyagi Pref.; ill-sorted calcareous
medium sand; early to middle Miocene
Moniwa formation (HANZAWA, et at.,1953).
13a-c-Sawane: a) a cliff (3s00.2' N., 13So
16.1' E.) situated about 1.1 Km NS5°W of
Sawane-cho,
Sawada-machi,
Sado-gun,Niigata Pref.; ill·sorted conglomeraticvery coarse sand., b) a cliff (3s00.9' N.,
13So 16.6' E.) situated about 1.4 Km N200W
of Sawane.cho; conglomeratic coarse shellsand., c) a cliff same as Loc. 17, YA~,lA­
GUCHI, 1973. The Pliocene Sawane for·mation.
14-Tanakura: a cliff (3701.5' N., 140°25.6' E.)
situated about 4.9 Km S57"W of IwakiTanakura Station, Tanakura, HigashiShiraka wa-gun, Fukushima Pref.; coarse
arkose sand; late Miocene Kamitoyo formation (IWASAKI, 1970).
15-Batoh: a cliff (35°45.1' N., 1400S.7' E.)
situated about 200 m east of the ShinNaka bridge across the Nakagawa river,Nasu-gun, Tochigi Pref.; cross· laminatedpumiceous conglomeratic coarse sand; late
Miocene Kobana formation (KAWADA,
1953) .
16-Tamatsukuri: a cliff (36°2.9' N., 140°27.1'
E.) situated about 7.5 Km S25°E of Tamatsukuri-machi Station, Fujii, Namekatagun, Ibaraki Pref.; fossiliferous fine sand;
Pleistocene Narita formation.
17a-c-Kioroshi: a) a cliff (35°49.9' N., 140°
9.6' E.) situated about 950 m S64°E of
Kioroshi Station, Inzai-cho, Inba-gun,Chiba Pref. ; fossiliferous well-sorted
medium sand., b) a cliff (35° 4S.5' N., 140°
12.2' E.) situated about 5.6 Km S5soE offossiliferous well-sorted medium sand., c)a cliff (35°47.1' K., 141°14.4' E.) situatedabout 9.S Km S55°E of Kioroshi Station;
fossiliferous well-sorted medium sand.
The Pleistocene Narita formation (KOZIMA,1955) .
18a-b-Tako: a, b) cliffs same as Locs. 18a,b, Y A~lAGCC[[I, 1973.
19a-c-Funabashi: a) Natsumidai-Kofun (anancient tomb) (35°41.S' N., 139°59.5' E.),
Natsumi·cho, Funabashi City, Chiba Pref. ~Kofun age (500-700 A. D.)., b) Miyamotodai-Kaizuka (a shell.mound) (35° 41.5' N.,
140°0.2' E.),
Miyamoto-cho,
Funabashi
City; late lomon age (3500 Y. B. P.)., c)
Ebigasaku-Kaizuka (a shell-mound) (3544.1' N., 140°2.5' E.), Ebigasaku, Oana-cho,
Funabashi City; middle Jomon age (4200
Y. B. P.). ColI. Miss. H. KOIKE.
20a-b-Semata: a, b) cliffs same as Locs.
19 a, b, Y.-\MAGlJCH1, 1973.
21-Taito: a coastal cliff (35°19.3' N., 140
24.7' Eo) situated about 2.4 Km NSOOEof Taito Station, Shimonohara, Isumigun, Chiba Pref.; poorly sorted conglomeratic mud containing fossil oysterbank (Crassostrea gigas) and autochthonous Mya arenaria oonogai; Holocene
Taito·zaki formation (lhlARA & TA1RA,
1974) .
22a-c-Kamiizumi: a-c) cliffs same as Locs.
20a-c, YA~lAGCCHI, 1973.
23a-b-Jizodo: a) a cliff (35°22' No, 140°6.2'Eo) same as Loc. 4, NAKAGA \\' A, 1960;
pumiceous and scoriaceous fossiliferous
medium sand., b) a cliff (35°21.4' N.,
0

0

681.

Taxonomic Studies on Japanese Balanoidea

140·5.4' E.) same as Loc. 5, NAKAGAWA,
1960; fossiliferous tuffaceous medium
sand. The Pleistocene Jizodo formation(NAKAGAWA, 1960).
24a-m-Kisarazu: a) a cliff (35·22.5' N.,
140·0.7' E.) same as Loc. 52, KOZIMA,
1966; cross-laminated well-sorted medium
sand., b) a cliff (35·23' N., 140·0.5' E.)
same as Loc. 55, KOZI:\lA, 1966; crosslaminated well-sorted medium sand., c) a
cliff (35·23.1' N., 140·0.5' E.) same as Loc.
56, KOZIMA, 1966; cross-laminated wellsorted medium sand., d) a cliff (35·23.1'
N., 139·58.5' E.) same as Loc. 60, KOZIMA,
1966; fossiliferous coarse shell sand., e)a cliff (35·23.2' N., 139·57.8' E.) same as
Loc. 63, KOZIMA, 1966; cross-laminatedwell-sorted medium sand., f) a cliff (35·22.7' N., 139·56.6' E.) same as Loc. 21a,
YAMAGUCl-ll, 1973., g) a cliff (35·22.6' N.,
139·56.6' E.) situated about 200 m southof Loc. 24f; fossiliferous pumiceous silty
medium sand., h) a cliff (35·22.5' N., 139·
56.5' E.) same as Loc. 69, KOZIl>IA, 1966;
fossiliferous pumiceous silty medium sand.,i) a cliff (35·22.5' N., 139·56.4' E.) sameas Loc. 68, KOZIl\IA, 1966; fossiliferous
pumiceous silty medium sand., j) a cliff(35·21.3' N., 139·55.1' E.) same as Loc. 78,
KOZIMA, 1966; fossiliferous pumiceous
silty medium sand., k) a cliff (35·21.3' N.,
139·55.3' E.) same as Loc. 21b, YAMAGUCHI, 1973., I) a cliff (35·21.1' N., 139·
54.7' E.) same as Loc. 80, KOZI~'IA, 1966;
fossiliferous pumiceous silty medium sand.,m) a cliff (35·20.8' N., 138·54.4' E.) sameas Loc. 81, KOZIMA, 1966; fossiliferous
pumiceous silty medium sand. a-k) The
Pleistocene Narita formation (AOKI &BABA, 1972). I-m) The Pleistocene Kamiizumi formation (AOKI & BABA, 1972).
25a-b-Futtsu: a) a cliff (35·20.4' N., 139·
52.8' E.) same as Loc. 85, KOZIMA, 1966;
coarse shell sand containing gravel; Pleistocene Jizodo formation (AOKI & BAllA,
1972)., b) a cliff (35·20.1' N., 139·52.2'
E.) same as Loc. 86, KOZIMA, 1966 ;coarse shell sand containing gravel; Pleistocene Naruto formation (AOKI & BABA,
1972).

141

26-Nishiyatsu: a cliff same as Loc. 22,
Y A~IAGUCl-ll, 1973.
27-Ichijuku: a cliff same as Loc. 23,
YAMAGUCHI, 1973.
28a-c-Tateyama: a) a cliff (34·58.1' N.,
139·49.4' E.) same as Loc. 1, YABE &SUGIY A:\IA, 1931; fossiliferous muddy sandoverlying reef-building corals., b) a cliff(34·58.1' N., 139·49.4' E.) situated about
150 m north of Loc. 28a; fossiliferous
muddy sand containing fragments ofcorals., c) a cliff (35·0.4' N., 139·53.2' E.)
same as Loc. 41, NOMURA, 1932; sandy
silt containing fossil oyster bank (Crassostrea nipponina). The Holocene Numa
formation (YOKOYAMA, 1911).
29-Yokohama: 8 m below the ground (35·25.9' N., 139·37.3' E.) same as Loc. 17,
MATSUSHll\IA, 1973; massive mud; Holocene Sakuragi-cho formation (5110 ± 125
Y. B. P.) (MATSUSl-lI:VIA, 1973).
30a-g-Totsuka: a) a cliff (35·24.7' N., 139·
32.-1/ E.) situated about 1.5 Km N10·E ofTotsuka Station, Totsuka-cho, Totsuka-ku,
Yokohama City, Kanagawa Pref.; mudcontaining bank forming oyster (Crassostrea
gigas)., b) a road-cut (35·23.4' N., 139·
31.7' E.) on the highway situated about 1
Km S45·W of Totsuka Station, Totsukacho, Totsuka-ku; mud containing bankforming oyster (Crassostrea gigas)., c) a
cliff (35·23.5' N., 139·32.7' E.) situatedabout 100 m west of Loc. 30b; mud containing bank forming oyster (Crassostrea
gigas)., d) a cliff (35·25' N., 139·32.9' E.)
situated about 2.3 Km N26·E of Totsuka
Station, Akiba-cho, Totsuka-ku; mud containing bank forming oyster (Crassostrea
gigas)., e) 6.2 m (A) (6550± 110 Y. B. P. ;MATSUSHIMA, 1971) and 10 m (B) below
the ground (35·22.8' N., 139·32.2 E.) sameas Locs. 2A and B of MATSUSl-lEVIA, 1972;
lignitic sandy silt., f) a cliff (32·23.2' N.,
139·30.9' E.) of rivulet situated about 2.4
Km S60·W of Totsuka Station, Kumisawacho, Totsuka-ku; mud containing bankforming oyster (Crassostrea gigas)., g) 0.5
m to 1 m below the Itachigawa river bed(35·21.7' N., 139·32.2' E.) situated about
1.2 Km N10·E of Ohfuna Station, Kasama-

142

ToshiY1(ki YAMAGUCHIcho, Totsuka-ku; shell sand_ e & g) The
Holocene Ohfuna shell bed (MATSUSllI:vIA,1972). a-d & f) The Pleistocene Naganuma formation (OTUKA, 1930).
31-Hitorizawa: a cliff (35'21.5' N., 139'36.6'
E.) of rivulet, situated about 2 Km S30'E
of Y6k6dai Station, Hitorizawa, Isogo-ku,
Yokohama City, Kanagawa Pref.; crosslaminated pumiceous medium to coarse
sand; Pleistocene Koshibaformation(OTUKA, 1930).
32a-b-Fujisawa: a) a cliff same as Loc.
25, YA:VIAGUCHI, 1973., b) a cliff (35'19.8'
N., 139'30.7' E.) of Kashio river, situatedabout 2.2 Km S73'E of Fujisa \Va Station,
Machiya, Fujisawa City, Kanagawa Pref. ;very coarse shell sand containing gravel.
The Holocene Ohfuna shell bed (MATSCSHlyIA, 1972). CoIl. Mr. Y. MATSCSHI~·IA.
33a-e-Kamakura: a) a cliff (35'20.6' N.,
139'33.4' E.) same as Loc. 321, SIIIKA:VIA& MASUJI~'IA, 1969; fossiliferous very
coarse sand., b) a cliff (35'20.6' N., 139'
33.2' E.) situated about 300 m west of Loc.
33a; pumiceous and scoriaceous coarse
sand., c) 5 m below the ground (2600±
80 Y. B. P., MATSUSHIMA; per. com.) (35'
18.4' N., 139'33' E.) of the Namerikawa
riverside, situated about 1 Km S20'W of
Kamakura Station, Kamakura City, Kanagawa Pref.; coarse sand; CoIl. Mr.
MATSUSHlyIA, d) 4 m below the ground(35' 19.3' N., 139'34' E.) situated about 1.2
Km N65'E of Kamakura Station, Yukinoshita-vVakaremichi, Kamakura City; coarse
sand containing gravel; CoIl. Mr. MATSL'SHlyIA, e) a cliff (35'21.5' N., 139'31..J' E.)of rivulet, situated about 1.5 Km S45'W
of Ohfuna Station, Okamoto-cho, Kamakura City; massive mud.
c, d) The
Holocene Kamakura shell bed (MATSUSHI\IA, 1974). e) The Holocene Ohfuna
shell bed (MATSUSHI.:vIA,1972). a, b) The
Pliocene Nojima formation (SHIKA\IA &MASUJIMA, 1969).
34-Higashi-Zushi: 5 m below the ground(35'17.7'N., 139'36.2'E.) same as Loc.
Z-1 (5520± 120 Y. B. P.; MATSUSHI\IA,1974) of MATSUSHIyIA, 1974, Zushi City,
Kanagawa Pref.; mud containing bankforming oyster (Crassostrea gigas) ; Holocene Zushi shell bed (MATSUSHIMA, 1974).
Coli: Dr. Y. KA,\IE.
35-Tsukuihama: a cliff same as Loc. 26,
YAyIAGUCHI, 1973.
36-Misaki: a cliff same as Loc. 27, YA~·IA­
GUCHI, 1973.
37a-b-Ninomiya: a) a cliff (3:5'18.7' N.,
139'15.1' E.) situated about 2Km N24'W of
Ninomiya Station, Uchikoshi, Ninomiya-cho,Naka-gun, Kanagawa Pref.; conglomeraticscoriaceous and pumiceous coarse shellsand., b) a cliff (35' 18.9' N., 139' 16.2' E.)
same as Loc. ms (Mushikubo), OTUKA,
1929, Oiso-machi, Naka-gun; scoriaceousandpumiceous medium sand.
The
Pleistocene Ninomi ya formation.
38-Kanogawa: below the ground (35'4' N.,
138'56.6' E.) situated about 500 m north ofBaraki Station, Nagasaki, Nirayama-cho,
Tagata-gun, Shizuoka Pref.; mud; Holocene. Coli: Mr. TAKAHASHI.
39-Nanao: a cliff (37'2.4' N., 139'57.4' E.)
same as Loc. 100, OTUKA, 1935, Nanao
City, Ishikawa Pref.; cross-laminated conglomeratic coarse shell sand containing
many bryozoans; early to middle Miocene
Nanao formation (OTUKA, 1935).
40a-f-Kanazawa: a) a cliff (36'31.5' N.,
136'41.2' E.) same as Loc. 31, KASE?,O &MATSUURA, 1965; coarse shell sand., b)a cliff (36'31.3' N., 136'42.3' E.) same as
Loc. 20, KASENO & MATSUURA, 1965;
massive fine sand., c) a cliff (36'32.8' N.,
136'42.5' E.) same as Loc. 16, KASENO &MATSUURA, 196:5; massive silty fine sand.,d) a cliff (36'33.6' N., 136'42.6' E.) sameas Loc. 9, KASEl'O & MATSCURA, 1965;
conglomeratic coarse sand., e) a cliff(36'34.1' N., 136',12.1' E.) of a branch of
the Kinppu river, situated about 100 m
SE of Higashi-Nagae, Kanazawa City,
Ishikawa Pref.; massive medium sand.,f) a cliff (36'34.7' N., 136'41.4' E.) sameas Loc. 2, KASENO & MATSUURA, 1965;
massive medium sand.
The Pliocene
Omma formation (KASE"IO & MATSUURA,
1965) .
41a-g-Mizunami: a) a cliff (35'24.9' N.,
137'19.3' E.) same as Loc. L15, ITOIGAWA,

681.

Taxonomic Studies on Japanese Balanoiclea

1960; massive medium sandstone., b) a
cliff (35°23.9' N., 137°H.8' E.) situatedabout 3.7 Km NI2°W of Mizunami Station,
Hongo, Mizunami City, Gifu Pref.; conglomeratic very coarse sa·ndstone., c) a
cliff (35°23.6' N., 137°16.9' E.) situatedabout 3.9 Km N38°E of Mizunami Station,Nataki, Mizunami City; conglomeraticvery coarse sandstone., d) cliffs (35°23.6'
N., 137° 16.6' E.) situated about 3.5 Km
N32°E of Mizunami Station, Dan, Mizunami City; conglomeratic very coarse
sandstone., e) a cliff (35°22.5' N., 137°
16.5' E.) same as Loc. S 45-5, ITOIGAWA,
1960; conglomeratic very coarse sandstone., f) a cliff (35°23.3' N., 137°12.8' E.)
situated about ·1.2 Km N37°E of Tokishi
Station, Shizuhora, Toki City, Gifu Pref. ;conglomeratic very coarse sandstone., g)a cliff (35°21' N., 137°12.7' E.) situatedabout 2.5 Km S72°E of Tokishi Station,
Nakahida, Toki City; conglomeratic very
coarse sandstone.
The late Miocene
Shukunohora sandstone of the Oidawara
formation (ITOIGAWA, 1960).
42a-e-Atsumi: a) a cliff (34°39' N., 137°
22' E.) same as Loc. 2, HAYASAKA, 1961;
mud containing bank forming oyster(Crassostrea gigas); Pleistocene Akasawa
siltstone (HAYASAKA, 1961)., b) a cliff(34°37.1' N., 137°14.7' E.) same as Loc.
139, I-IAYAs.-\KA, 1961; conglomeratic
medium sand., c) a cliff (34°37.1' N.,
13fD14.7' E.) same as Loc. 138, HAYASAKA,
1961 ; conglomeratic medium sand., d) a
cliff (34°37.1' N., 137°H.8' E.) same as
Loc. 137, HAYASAKA, 1961; conglomeratic
medium sand., e) a cliff (3el°H.8' N., 137°
H.8' E.) same as Loc. 28, YA:-IAGCCHI,
1973. b-e) The Pleistocene Toshima sand
(HAYASAKA, 1961).
43-Akashi: exact locality unknown, Akashi
City, Hyogo Pref.; Pleistocene Maikoshell bed (SIIIKA:-IA, 1936). Repository:
Osaka City Museum Natural History.
44-0kinawa-jima Is. : a road-cut (26°4004' N.,
127° 59.9' E.) situated about 500 m S200E
of Nakasone, Nakijin.son, Nago City,
Okina wa Pref.; calcareous fine sand;
Pleistocene Nakoshi sand.

4.

143

Evaluation of Morphological
Characters

Before entering into discussion ofsystematics, some morphological characters of balanoids are taxonomicallyevaluated. Taxonomy based on an adequate evaluation of morphological chraacters at the species level may provide
evidence of evolutionary processes, andcontribute to the determination of higher
ca tegories.

Compartments
External surface of shell wall:
That the nature of external surfaceof shell wall is often affected by the
microtopography of substratum has beenshown in the following examples; living
B. porcatus (=B. balanus), B. crenatus
and B. patellaris (=B. patelliformis)(DARWIN, 1854a); living B. rostratus
apertlls and B. balanus (PILSBRY. 1916);
B. shilohellsis (PILSBRY, 1930); living B.
ebunzells and fossil B. concavus (GREGG,
1948). etc. In fossil of B. rostratus
growing on Chlamys nipponensis andChirona (Chirona) evennanni on Chlamys
islandiClls, external surfaces of shellwalls have a fine relief which in effectis a replicate of ribs of the scallop. It
is well kno\vn that the external surfaceof shell wall is also influenced by other
ecological factors, such as population
density (K.HO, et aI., 1960a, b) andwave action.
Solidobalanus (Hesperibalanus) hesperius
nipponensis was originally distinguishedfrom S. (H.) hesperius s. s. by a smoothrather from ribbed external surface(PILSBRY,1916). However, living Japanese
specimens of S. (H.) hesperius growing
on the same substratum show a wide
range of individual variation; the external surface ranging from smooth to144

Toshiyuki YAMAGUCHI

ribbed (pI. 27*, figs. 4, 5). Thus thismorphological character can hardly be
regarded as useful in distinguishing S.
(H.) hesperius s. s. from S. (H.) h.
nipponensis.
In conclusion, since the external surface of the shell wall may be modified
by ecological factors, such as the natureof substratum, population density, etc.,
care must be taken in how this character is used in the classification of thebarnacles.
Radii:
In his revision of the Tetraclitidae, it
was emphasized by Ross (1969) that themode of shell growth is significant intaxonomy. Two modes of shell growth
were distinguished in one family as
follows: diametric shell growth (which
produces enlargement of radii) andmonometric shell growth (which does
not induce enlargement of radii).
The summits of radii both in Megabalanus rosa and 1\1. volcano as well asother members of "Megabalanus tintinnabulum" species group are nearlyparallel to the base (YAMAGUCHI, 1973).
But in a few species of Megabalanus,
the summits are oblique. Therefore,
this morphological character can't be a
sub generic character of this group.
In four living species of B. amphitrite
species group from Japan, the natureof summits of radii ranges from horizontal to oblique, and the degree ofinclination seems to reflect specificdifference.
In conclusion, the width and nature ofsummits of radii are important characterin taxonomy, but differences mayor
may not be useful above the species
level.

·pls. 23-27 to be published in part 2.

Internal structure of compartment
Longitudinal tubes, transverse septa andsubsidiary tubes:
PILSBRY (1916) interpreted the modeof distribution of transverse septa inlongitudinal tubes and the number of
longitudinal tubes in rostrum as useful
criteria in distinguishing subspecies ofB. rostratus. However, PILSBRY'S subspecific designations are not necessarily
acceptable in the light of modern species
concepts, because the entire range ofdiagnostic characters utilized are to befound within the population of Japanese
B. rostratus. On the other hand, living
Japanese B. amphitrite can be distinguished from three other species of B.
amphitrite group by the absence oftransverse septum in longitudinal tubes.
DARWIN (1854a) described that the longitudinal septa of B. amphitrite occasionally bifurcate at the base and makeirregular minute tubes (subsidiary tubes).
Subsequently, the presence of subsidiarytubes has been shown by following
authors: in B. amphitrite albicostatus(=B. albicostatlls) by PILSBRY (1916); in
B. pallidus pallidus and B. p. stutsburi
by HARDING (1962); in B. pallidus, B.
albicostatus albicostatus (=B. albicostatus)
and B. a. formosanus by UTINOMI (1967)
and in B. citerosum, B. suturaltus and B.
dentivarians of B. amphitrite-complex by
HENRY (1973).
Taxonomic evaluation on the mode of
distribution of transverse septa andnumber of longitudinal tubes should bemade through the examination of individual and geographical variations.
The absence of transverse septa and thepresence of subsidiary tubes are considered to be important morphologicalcharacters for classification; especiallythe latter may represent a close relationship among species.

681.

Taxonomic Studies on Japanese Balanoidea

Nature of cavity between sheath andinner lamina:
HENRY (1973) attached importance tothe presence of calcareous vesicules
filling the cavity between sheath andinner lamina, and pointed out that B.
citerosum, B. suturaltus, B. dentivarians,
B. crenatus, B. glandula, Semibalanus
cariosus and S. balanoides possess calcareous vesicules. In living Japanese B.
amphitrite species group, transverse septa
C=calcareous vesicules) are developed in
the cavity between sheath and innerlamina in B. albicostatus and B. kondakovi, but absent in B. amphitrite and B.
reticulatus.
This morphological character also
seems to be taxonomically important,although ecophenotypic variation needs
to be examined.
Opercular valves
Proportion of tergal and basal length of
scutum:Solidobalanus (Hesperibalanus) hesperius
nipponensis was distnguished from S.
(H.) hesperills s. s. by a difference in the
proportions of tergal and basal lengthof scuta (PILSBRY, 1916). The differenceis, however, intrapopulational in Japan.
Therefore, this character is not adopted
as a criterion for subspecific distinction.
On the other hand, in an examinationof shell growth of living Japanese B.
amphitrite, B. albicostatus and B. reticulatus, it was observed that growth of
the tergal and basal margins of the
scutum of B. albicostatus and B. reticlilatus is isometric, but that in B.
amphitrite it is allometric. Therefore,
there are cases where the shape of
scutum changes with age as well aswith ecological factors.
Ratio of the width of spur to the distance
between basiscutal angle and anterior
face of spur:

145

Ratios of the width of spur to thebasal margin have been noted in thesystematic descriptions by some authors.
But the extent of individual and geographical variations is rarely taken into
consideration. Closely related species
Megabalanus rosa and A1. volcano arebiometrically distinguished by the ratioof the width of spur and distance between basiscutal angle and anterior faceof spur (YAMAGUCHI, 1973). On theother hand in three living species fromJapan, B. amphitrite, B. albicostatus and
B. reticulatus of B. amphitrite group, theratio shows wide ranges of intra- andinterpopula tional varia tions which overlap each other.
In summary, these examples again
illustrate that a particular morphologicalcharacter may be important in somegroups but not in others. The value of
a c!1aracter must be judged in each case.
Outline of basal margin of tergum:
The nature of the basal margin of
tergum of Megabalanus rosa is clearly
different from that of Ai. volcano CY AMAGUCHI, 1973). HENRY (1973) regarded
the nature of basal margin of tergum
as a diagnostic character in the establishment of four new species in B. amphitritecomplex. In living B. amphitrite, B.
albicostatus, B. reticulatus and B. kondaIwvi from Japan, the nature of basal
margin is also important for the species
level classification.
Appendages
Number of segments in each cirrus:
NEWevlAN (1967) reported that Tetrachthamalus oblittertus [Chthamalidae]
collected from three localities shows anarrow range of individual variation but
a wide range of geographical variationin length and number of segments of
exopodite of cirrus III. However, it has
since been noted that the exopod of the

146

Toshiyuki YAMAGUCHI

third cirrus ranges between normal andantenniform on a seasonal basis in thesame population (NEWfI'IAN, pers. com.).
YAMAGUCHI (1973) demonstrated thatthe range of the number of segments ofcirri I to III in Megabalanus rosa and M.
volcano show narrow ranges of individual and geographical variations, but
those of cirri IV to VI show wide ranges
of individual variations; those in the
exopodite of cirrus I and in both rami
of cirrus III are distinctly different between two species; and these differences
aid in discriminating between species.
In four living species of B. amphitritegroup from japan, the numbers of segments in cirri I to III show narrow
ranges of individual variation, but they
overlap each other to the degree that it
is impossible to discriminate between
these four species by this morphologicalcharacter alone (Table 1).
Thus. taxonomical significance of thismorphological character must be evaluated in accordance with individual,
seasonal and geographical variations.
Mandible:
Ross (1971) noted the mandible of
Tetraclitella
Iwrandei[TetraclitidaeJ
differs from that of other species of thisgenus in the degree of development andnumber of subsidiary cusps of thesecond to fourth teeth. YAMAGUCHI(1973) demonstrated that the nature of
the fourth and fifth teeth of mandibleis clearly distinct between Megabalanus
rosa and M. volcano and represents a
specific character. On the other hand,four living species B. amphitrite groupfrom japan are difficult to separate bythis morphological character. In summary, it is again difficult to generalize
taxonomical importance of these morphological characters.
Applying the morphological evaluation

discussed above, it is possible to demonstrate various degrees of morphologicaldivergence between related forms inbalanoids of japanese area. When thedivergence is such that a significant gap
appears between species groups, subgenera and genera may be formed.
When the gap is recognizable, but not
fully defined or substantiated, the complex can be refined as a species group.
The following groups, occurring inJapanese waters, form basis of thepresent study.
"Balanus amphitrite" group (excluding
B. eburneus and B. improvisus) is chraacterized by the followings: Shell wall
externally with colored longitudinal
stripes and sometimes with colored horizontal lines. External surface of scutumsmooth and with weakly prominent
growth lines. Adductor ridge of scutum
weakly to strongly prominent but notvery long; and sometimes with shallow
pit below. Distal border of each proximal intermediate segment of cirrus III(frequently to VI) with several small
spinules. The group includes the following closely related species: B. amphitrite,
B. albicostatus, B. reticulatus and B.
Iwndakovi.
"Ivlega balanus tintinnablllllm" group isreadily distinguishable from the otherbalanoids by the following characters:
Parietes with longitudinal tubes andtransverse septa. Radii wide with tubes
and their summits usually parallel to thebase. Basis calcareous and tubiferous.
Scutum with beaked apex and withbroad basal border. Basal margin of
tergum roughly straight on both sidesof the spur. Intermediate segments ofcirri III and IV with short spinules on
the distal borders. The group includes
two species, AI. rosa and Jvl. volcano.
Solidobalanus (Hesperibalanlls) hesperius
is mainly characterized by the following

681.

Taxonomic Studies on Japanese Balanoidea
Number of segments in each right cirrus of B. amphitrite,
B. albicostatus, B. reticulatus and B. kondakovi.

Table 1.

rCI

.~
'-

147

rCm

rCll

rCV

rCIV

rCVI

end

exo

end

exo

end

exo

end

16

16

31

34

32

33

35

34

15

14

29

29

33

33

35

36

11

14

i-

-

35

37

36

37

11-13

14-16

14-16

29-31

29-34

32-35

33-37

35-36

34-37

12.0

15.0

14.7

30.0

31.5

33.3

34.3

35.3

35.7

26

29

24+

21+

2,1+

24+

26

28

28

29

25

24

exo

end

exo

end

RA827,j

23

13

16

13

RA8365

21

12

15

12

RA8370

21

12

U

11

21-23

12-13

14-16

21.7

12.3

15.0

exo

~ 1--------1-----------1---------------------,---------------------1---------~
N
33333322333
3
OR

RA8275

23

13

14

12

Hi

12

RA8366 117

13

13

13

16

12

I

7 ____3_8___ __RA8371 1_1_9____1_3___ 1_1_5_____
13___ 1._1_7____1_7___ 1__3_3____3_5___ 1__3__
1 39_____4_0___

N

3
13

3
13-15

X

19.7

13

11.012.716.313.728.330.730.033.532.032.0

3
16-17

3
12-17

3
26-33

3
28-35

2
29-38

2
24-40

3
17-23

3
12-13

2
28-32

2

OR

25-39

--~-------:'·----------I-----------I----------I-----------I----------l·---------36
32
33
36
37
RA8276i23
15
18
16
19
17
I 33
RA8367
RA8372
N
OR

x:

19
13
15
12
16
13
129
20
12
16
13
17
16
26+
----------1----------1---------3
3
3
3
3
[2
3
19-23 12-15 15-18 12-16 16-19 13-17 I 29-33
20.713.316.313.717.3
15.3 131. 0

RA8362

17

13

13

12

18

RA8363
RA83Gcl

19
17

10
12

14

8+

19

14+

13

N

OR

11~

-19
I 17.7

18

33

34

37
36

38
34

2
32

2
33

2
34-36

3
36-37

3
34-38

32.0

33.0

35.0

36.3

36.3

32
12+

~~

l-~-~-----~!---I.-~-.~----~-:-:---l-~-~----:-,~---

16

134

30

43

42

46

45

1 3

3
30-38

3
34-c14

1
42

3
35-47

3
40-47

33.7

40.3

42

'12.7

44.0

!----------

3
10-13

2
13-14

2
12-13

3
18-19

11.713.512.518.3

3

16-17130-34
16.331.7

N: number of individuals examined. OR: observed range. x: arithmetic mean. Locality:
B. amphitrite, B. albicostatus and B. reticulatus; Loc. H. B. kondakovi; Kasaoka Bay, Okayama
Pref., ColI. Dr. S. FUSE.

characters: Wall solid. Radii narrow
with oblique summits and denticulated
sutural edges. Scutum flat with weakgrowth ridges.
Faint callus-like projections observable between articularand adductor ridges in some specimens.
Tergum with wide spur. Crests fordepressor muscle strongly developed.
B. rostratus may be a Japanese re-presentative of B. balanus group of theAtlantic and is characterized by thefollowings: Shell with smooth externalsurface. Carinolaterals narrow. Rostrum
well developed. Radii narrow with oblique summits. Scutum with prominent
growth ridges and distinct longitudinal
striations.
Pit for adductor muscle
weakly developed or hardly visible.

Toshiyuki Y AMAGUCHl148

B. amphitrite inexpectus PILSBR1', 1916
B. amphitrite peruvianus PILSBRY, 1916
B. amphitrite forma hawaiiensis BROCH, 1922
B. amphitrite var. acutus WITHERS, 1924
B. amphitrite var. denticulata BROCH, 1927
B. amphitrite forma poecilosculpta BROCH,
1931
B. amphitrite kriigeri NILSSON·CANTELL,
1932
B. amphitrite raffiesi NILSSON·CAI'TELL,
1934
B. amphitrite forma formosanus HlRo, 1938
B. amphitrite var. jluminensis OLIVEIRA, 1941
B. amphitrite var. aeratus OLIVEIRA, 1941
B. amphitrite litoralis KOLOSVARY, 1948
B. amphitrite abundantus KOLOSy.ARY, 1948
B. amphitrite archi-inexpectatus KOLOSV.ARY,
1948
B. amphitrite hungaricus KOLOSV.ARY, 1948
B. amphitrite helenae KOLOSV.ARY, 1948
B. amphitrite var. vladivostokensis TARASOVand ZEVINA, 1957
B. amphitrite var. columnaris TARASOV andZEVINA, 1957
B. amphitrite var. kondakovi TARASOV andZEVINA, 1957
B. amphitrite karakumiensis KOLOSV .ARY,
1961
B. amphitrite merklini KOLOSV .ARY, 1962aB. amphitrite tongaensis KOLOSV ARY, 1962b

Tergum with wide and blunt spur.
B. crenatus may consist of several
geographic races or subspecies and ischaracterized by the following characters:
External surface of shell wall smooth.
Radii narrow with steeply oblique
summits and denticulated sutural edges.
Calcareous vesicules develop in thecavity between sheath and inner lamina.
Scutum with very weak growth ridges.
Articular ridge very prominent and reflexed. Adductor ridge very weak or
hardly visible. Pit for adductor muscle
deep.
In the following pages, the writerintends to evaluate the species levelclassification of some important groupsof Japanese barnacles.
5. Revision of Balanlls
amphitrite group

Necessity of a species revision of the B.
amphitrite group
B. amphitrite was originally describedby DARWIN (1854a). He divided it intonine varieties. However, his descriptions
are brief and imcomplete. The typespecimen, type-locality and "type" forvariety were not designated by DARWIN.
After that, the following varieties,
forma or subspecies were additionallydescribed as infraspecific taxa of B.
amphitrite.
B. amphitrite var. malayensis HOEK, 1913
B. amphitrite albicostatus PILSBRY, 1916

Recently, from a modern taxonomic
standpoint, HARDING (1962) reexamined
and redescribed the specimens which
had been studied by DARWIN. HARDING
allocated the taxonomic position of
DARWIN'S nine varieties of B. amphitrite
in accordance with the degree of morphological difference, as shown in Table 2.

Table 2(1854 a)HARDING (1962)
B. amphitrite var. communis ............................ B. amphitrite amphitrite DARWIN
B. amphitrite var. pallidus ... .......................... . B. pallidus pallidus DARWIN
B. amphitrite var. stutsburi ........................... . B. pallidus stutsburi DARWIN
B. amphitrite var. venustus ........................... . B. venustus venustus DARWII'
B. amphitrite var. niveus ............................. . B. venustus niveus DARWIN
B. amphitrite var. modestus ........................... . B. venustus modestus DARWIN
B. amphitrite var. obscurus ........................... . B. venustus obscurus DAR\vI:\!
B. amphitrite var. variegatus ............................ B. variegatus DARWIN
B. amphitrite var. cirratus ........................... . B. variegatus var. cirratus DARWINDARWIN

681.

Taxonomic St1~dies on Japanese Balanoidea

149

Table 3
HIRO

(1938)

UTI?-JOMI(1967)

B. a. forma communis DARWIN . . . . . . . . . . . . • . . . . • . . . . B. reticlllatlls UTI:\'Ol\1l
B. c. forma cirratus DARWIN . . . . . . . . . • . . . . . • • . . . . . . B. variegatus cirratus DAR\\"!:\,
B. a. forma poecilotheca KRUGER . . . . . . . . . . . . . . . • . . . • B. poecilotheca KRUGER
B. a. forma albicostatus PILSURY .•••..••.•..••••.... B. albicostatus albicostatus PILSURY
B. a. forma hawaiiensis BROCH • . . . . . . . • . . . . . . . . . . . . . B. amphitrite DAR\\'IN sensu HARDING
B. a. forma kriigeri NILSSON·CA:\'TELL •.....••••.•.. B. llliginoslls UTI:\'OMI
B. a. forma jormosanlls HIRO ...•...••.••.••••..•.... B. albicostatus jormosanus HIROUTINO~II (1967) also revised his (1938)
seven forms B. amphitrite from Japaneseand its adjacent waters, as shown in
Table 3.
HARDING (1962) and UTI NOM I (1967)
proposed their classifications for thisspecies group mainly on the basis ofdegree of morphological difference.
Other criteria, at the species level, werein general not utilized in the classification. Therefore, a reexamination of
B. amphitrite group from biologicalstandpoint could be of value.

Taxonomic problems concerning theJapanese B. amphitrite group
Excluding species from Formosa III
UTINOMI's (1967) list, there are six
species of B. amphitrite group describedfrom Japanese waters.
B. poecilotheca was described by
KRUGER (1911a) from Okinose Bank,
Sagami Bay. HIRO (1938) once placed
this species in B. amphitrite group andcalled B. a. forma poecilotheca, but inhis 1967's paper, he excluded this species
from his list of B. am,bhitrite complex.
Occurrence of B. amphitrite forma
cirratus from Iki Is. and Ariake Bay wasreported first by HIRO (1938). This form
was later called B. variegatus var. cirratus
by HARDING (1962) and B. variegatus
cirratus by UTINOMI (1967). Thus it
seems necessary to clarify the relation
between B. variegatus variegatus and B.

variegatus var. cirratus; are they mere
variations of one species or are they
representatives of two geographicallydefinable subspecies. Unfortunately thisspecies is too rare in Japanese waters
to ascertain its taxonomic status.
B. albicostatus and B. uliginosus (=B.
kondakovi) present no taxonomic problem(UTINOMI, 1970 ; HENRY and McLAUGHLIN,
1975).
In order to clarify the taxonomic
status of B. amphitrite and B. reticulatus,
it is necessary to examine descriptions,and illustrations of both DARWIN (1854a)and HARDING (1962) in some detail.
HARDIC\'G (1962) examined the specimens of B. amphitrite which was originally described by DARWIN (1854a). The
majority of DARWIN'S specimens areincompletely preserved, and their localities in many cases unknown. However,
HARDING found many complete individuals of communis and venustus
together on a single bamboo fragmentfrom Natal. These specimens are accompanied with handwriting of DARWIN
as B. amphitrite. In fact, as pointed by
HARDING (1962), DARWIN (1854a, p. 246)
had described B. a. var. venustus andB. a. var. communis from Natal. Amongthe communis on the bamboo, a complete
specimen was selected as the lectotypeof B. amphitrite by HARDIl\G, because
the type specimens of B. amphitrite had
not yet been designated by DARWIN and150

Toshiyuki YAMAGUCHI

later authors. Thus, B. a. var. communis
of DARWIN (1854a) was designated as B. a.
amphitrite by HARDING (1962).
UTINOMI (1967) interpreted that theillustratei specimen of B. a. var. communis (DAR,VIN, 1854a, pI. 5, figs. 2e, 2h
and 21) differed morphologically fromthe lectotype of B. a. amphitrite (HARDING, 1962, pI. 1, figs. a-h), and that thedifference deserves specific distinction.
Therefore, he gave a new specific name
B. reticulatus to the former.
Now it becomes necessary to reexamine
the figures and descriptions of bothDARWIN and HARDING to find out
whether B. a. var. comlllunis of DARWIN
and B. a. amphitrite described by
HARDING are conspecific with each otheror different species.
DARWIC\'S (1854a, p. 240) descriptionof the radii and tergum of B. a. var.
communis can be summarized as follows:
1) Summits of radii oblique or nearlyparallel to the base; 2) basal shape of
spur square or bluntly pointed. DARWIN
illustrated a specimen which is characterized by the oblique summits of radii
and the bluntly pointed basal shape of
spur (DAR,VI:--', 1854a, pI. 5, figs. 2e, 2h
and 21). DARWIN did not describe any
more details about radii and tergum.
HARDIC\G'S (1962, p. 277, 278) descriptionof the radii and tergum of B. a. amphitrite can also be summarized as follows:
1) Summits of radii in specimens including the lectotype (pl.1, figs. a-b) are almost
parallel to the base, but in some otherspecimens more oblique; 2) many specimens labelled by DARWIN as B. a. var.
comlllunis, which were later called B. a.
amphitrite by HARDING, have short andwider spur, but one or two individuals
have such narrow spur as DARWIN'S
figure (pI. 5, fig. 21).
Now it is obvious from the DARWIN'S
and HARDING'S descriptions that there

exist two distinct forms in one species.
One is characterized by the oblique
summits of radii and a narrow spur of
tergum, the other has the summits ofradii which are parallel to the base andwider spur of tergum.
Although DARWIN and HARDING interpreted these morphological difference as
infraspecific, UTINOlVII (1967) consideredthese differences as interspecific.
The writer interpreted from the DARWIN'S figures (pI. 5, figs. 2e, 2h and 21)that B. a. var. communis carries the following morphological characters in addition to the characters already describedby DARWIN and HARDING: Wider distance between anterior face of spur andbasiscutal angle than width of spur;slightly concave basal margin at carinalside of spur in tergum.
From the figures (HARDING, 1962, pI.
1, figs. e-f) of the lectotype of B. a.
amphitrite, the tergum of B. a. amphitrite
can be characterized by the followings:
Distance between anterior face of spurand basiscutal angle approximately half
width of spur; basal margin both sidesof spur straight. Therefore, the illustrated figures (DARWIN, 1854a, pI. 5, figs.
2e, 2h and 21) of B. a. var. communis
DARWIN is clearly different from those(pI. 1, figs. a-f) of HARDING'S lectotypeof B. a. amphitrite in the nature ofsummits of radii, width, shape andposition of spur of tergum.
Thus it appears that the two forms
are different species, as was pointed out
by UTlNOlVII (1967) and confirmed by
SOUTHW ARD (1975), by HENRY and McLAUGHLIN (1975), and by the present
investigation. The form characterized
by a oblique summits of radii and anarrov" spur is B. reticulatus.
Comparison of morphological characters of
the B. amphitrite group of japan

681.

Taxonomic Studies on Japanese Balanoidea

In order to evaluate the conclusions
based on the literature survey explainedin the foregoing lines, the morphologyof the Japanese specimens will be examined systematically.
Shell wall
1) Nature of external ornamentation, colorand stripes of shell wall
In B. amphitrite. B. reticulatus and B.
kondakovi, the external surface of shellwall is smooth with coloration of longitudinal stripes. Each stripe is always
found between two adjacent primarylongitudinal septa.
In B. amphitrite (pI. 19, figs. la-b,2a-h),
the ground color of the shell is dirty
white, and the color of stripes is grayish
blue to purplish blue or reddish purple.
The colored stripes are found on all theexternal surface except the central
trapezoidal areas of rostrum and lateralcompartments.
In B. reticulatus (pI. 19, figs. 5a-b,5g-n), the ground color of shell wall isdirty white to gray, light purplish redor purplish recl.
These longitudinalcolored stripes are intersected by whiteor colorecl horizontal stripes.
In B. lwndalwvi (pI. 19, figs. 6a-b,7a-h), the ground color of shell wall isdirty white.
The stripes are purple.
or reddish to bluish purple. These longitudinal colored stripes are occasionallyintersected by white horizontal stripes.
In B. albicostatus (pI. 19, figs. 3a-b,4a-h), the external surface of shell wall
has white longitudinal ribs. which arevariable in length and width and always
correspond to primary longitudinal septa.
But the septa clo not always correspondto the ribs. The ground color of shellis purple, reddish purple or violet.
2) Radii and their summits
In B. amphitrite, the radii are welldeveloped, and their summits are approximately parallel to the base. In B.

151

albicostatus and B. reticulatus, the radii
are moderately developed, but theirsummits are oblique to the base. In B.
kondakovi, the radii are narrow andtheir summits are steeply oblique to thebase.
3) Shell structurea) Longitudinal tubes (Text-fig. 2)
In B. amphitrite (pI. 20, figs. 5, 9a-d),
the longitudinal tubes have no transverse
septum. The longitudinal tubes in B.
albicostatus (pI. 20, figs. 6, lOa-d), B.
lwndakoL'i (pI. 20, figs. 8, 12a-d) and B.
reticulatus (pI. 20, figs. 7, lla-d) havetransverse septa which are parallel tothe base. However, the transverse septaof the first two species are arranged
regularly, while in the last they are irregularly spaced.
b) Subsidiary tubes (Text-fig. 2)
In B. albicostatus (pI. 20, fig. 2), each of
the longitudinal tubes is very often accompanied by small (subsidiary) tubes.
They are formed by secondary longitudinal septum which projects from theouter lamina and join to the wall of the
primary longitudinal septum.
In B. amphitrite (pI. 20, fig. 1), B.
reticlilatus (pI. 20, fig. 3) and B. lwndakoui(pI. 20, fig. 4), no subsidiary tubes arefound.
c) Cavity between sheath and inner lamina
In B. amphitrite (pI. 20, fig. 5) and B.
reticlilatlis (pI. 20, fig. 7), the cavity between sheath and inner lamina is filled
up by calcareous matter, while those of
B. albicostatus (pI. 20, fig. 6) and B.
kondakovi (pI. 20, fig. 8) have calcareous
vesicles.
Opercular ralz'e
1) Scutuma) Adductor ridge
The adductor ridge of B. amphitrite(pI. 19, fig. Id), B. reticulatlls (pI. 19, fig.
5d) and B. kondakovi (pI. 19, fig. 6d)

152

Toshiyuki YAMAGUCHI~: ", :-: '!'.'. " . ~B. amphitrite

B. albicostatus

B. reticulatus

B. kondakovi

Text.fig. 2. Schematic profiles of compartments of B. amphitrite,
B. albicostatus, B. reticulatus and B. kondakovi.

slopes towards the occludent margin.
However, that of B. albicostatus (pI. 19,
fig. 3d) projects subvertically from theinner surface of scutum.
2) Terguma) Spur
In B. amphitrite (pI. 19, figs. Ie-f), thespur is wide, and squarish at the end.
The spur of B. albicostatus (pI. 19, figs.
3e-f) is narrow and obliquely truncatedat the end. The spur of B. reticulatus(pI. 19, figs. 5e-f) is moderately wide but
becomes narrow toward the end androunded or obliquely truncated at theextremity. The spur of B. kondakovi(pI. 19, figs. 6e-f) is narrow, becomesnarrower toward the end and is sharply
pointed at the extremity.
b) Basal margin of tergum
In B. amphitrite, the basal margin at

the scutal side of spur is straight, andat the carinal side of spur it is straightor slightly concave. In B. albicostatus,
the basal margin at the carinal side ofspur is slightly concave. The basal
margin of both sides of spur of B.
reticulatus is approximately straight.
The basal margin at the carinal side ofspur of B. lwndakovi is deeply notched.
Animal parts
1) Cirria) Nature of anterior faces of cirri III
and IVIn B. amphitrite (pI. 21. figs. Ib-c), theexopodite of cirrus III is provided withmany short and blunt spines (hook-likespines) on the anterior face of eachsegment, but exopodite of cirrus IV has
a few short and fine spinules instead.

681.

Taxonomic Studies on Japanese Balanoidea

153

...
0

to

'"
I)

0

100

200

Km

I.'

26° N

.~

~"

III

o

100

200 KmB. kondakovi

Text-fig. 3. Distribution of B. amphitrite, B. albicostatus, B. reticulatus and B. kondakoL'iin the Recent sea. [Compiled from ARAKA\\A; 1973, HlRo; 1933, 1939a, b, lGA; 1973,
MAWATARI; 1967, MAWATARI and KOBAYASHI; 1954a, b, MIZUNO, HARADA and MIZCNO;
1964, TOYOSI:VIA and lRIE; 1962, UTINO~II; 1949, 1955, 1962, 1970, UTINO:VII and KIKUCHI;
1966, YAMAGUCHI; 1972MS, 1974MS, YAMAMURA; 1971, Y A~IA:VIURA, KU\\"ATANI and NISHII;
1969J. 1. Hamana Lake, 2. Mika\\"a Bay, 3. Tanabe Bay, 4. Kure, 5. Hiroshima Bay, 6.
Sasebo.

154

Toshiyuki YAMAGUCHIIn B. albicostatus (pI. 21, figs. 2b-c),
spinules on the anterior face of eachsegment of the exopodite of cirri III and
IV become obscure. The exopodite of
cirri III and IV of B. reticulatus (pI. 21,
figs. 3b-c) has dense short spinules at
the anterior face of each segment, while
in B. lzondakovi (pI. 21, figs. 4b-c) it has
a few short spinules near the anterodistal corner of each segment.
2) Labrum

The labrum of B. amphitrite (pI. 21,
fig. 1a) has 11 or more teeth, while thelabrum of B. albicostatus (pI. 21, fig. 2a),B. reticulatus (pI. 21, fig. 3a) and B.
kondalzovi (pI. 21, fig. 4a) has only 3 or 4teeth on both sides of the labrum notch.

In conclusion, four species discussed
above are morphologically distinct fromeach other. Degree of morphologicaldifference between the three species. B.
amphitrite, B. reticulatus and B. kondakovi, seems similar. B. albicostatus, however, seems different from the above
three species in having characteristic
ornamentation and shell structure.

Distribution
A conclusion is that in general in theJapanese waters, the four species occur
in macro geographically sympatric relationship. Now it become3 important
to examine whether or not this macrogeographical sympatry includes physicalcontact between species.
Members of the B. amphitrite group

Explanation of Plate 19
Fig. 1. Balanus amphitrite DARWI" (UMUT -RA8263) from Tanabe Bay (Loc. H). la-b. wholewall, x 2., lc-d. exterior and interior views of right scutum, x 6., Ie-f. exterior andinterior views of right tergum, x 6.
Fig. 2. Balanus amphitrite DARWIN (UMUT -RA8264) from Tanabe Bay (Loc. H). 2a-b. exterior and interior views of rostrum, x 2.S., 2c-d. exterior and interior views of leftlateral, x 2.S., 2e-f. exterior and interior views of left carinolateral, x 2.S., 2g-h. exterior and interior views of carina, x 2.S.
Fig. 3. Balanus albicostatus PILSBRY (UMUT -RA8265) from Tanabe Bay (Loc. H). 3a-b.
whole wall, x 2., 3c-d. exterior and interior views of right scutum, x S., 3e-f. exterior
and interior views of right tergum, x 5.
Fig. 4. Balanus albicostatus PILSBRY (UMUT -RA8266) from Tanabe Bay (Loc. H). da-b.
exterior and interior views of rostrum, x 2., ·le-d. exterior and interior views of leftlateral, x 2., ,Ie-f. exterior and interior views of carinolateral, x 2., .lg-h. exterior andinterior views of carina, x 2.
Fig. 5. Balanus reticulatus UTIl'o:\H (UMUT -RA8267) from Tanabe Bay (Loc. H). Sa-b.
whole wall, x 1.5., 5c-d. exterior and interior views of right scutum, x 4., 5e-f. extericr
and interior views of right tergum, x 4., 5g-h. exterior and interior views of rostrum,
x 1.5., 5i-j. exterior and interior views of left lateral, x 1.5., 5k-1. exterior and interior
views of left carinolateral, x 1.5., 5m-n. exterior and interior views of carina, x 1.5.
Fig. 6. Balanus kondakovi TARASOV and ZE\'I:\A (UMUT -RA8268) from Hamana Lake (Loc.
G). 6a-b. whole wall, x 1.5., 6c-d. exterior and interior views of right scutum, x 4.,
6e-f. exterior and interior views of right tergum, x 4.
Fig. 7. Balanus kondakovi TARASOY and ZEVI:\A (UMUT-RA8269) from Hamana Lake (Loc.
G). 7a-b. exterior and interior views of rostrum, x 1.5., 7c-d. exterior and interior
views of left carinolateral, x 1.5., 7g-h. exterior and interior views of carina, x 1.5.
[Descriptions to be published in Part 2J

YAMAGUCHi:

Japanese Balanoidea

Plate 19

681.

Taxonomic Studies on Japanese Balanoidea

are all warm water inhabitants. Theirgeographical range extends from southeast Asia to the Japanese coast.
The northern limit of distribution of
B. amphitrite, B. albicostatus and B. reticulatus is the Tsugaru Strait, but thatof B. kondalwvi is Tokyo Bay on the
Pacific side and Nakanoumi Lake on theJapan Sea side. The first three species
are widely distributed throughout theJapanese coast, but the last scattered.
B. amphitrite, B. albicostatus and B.
reticulatus have been reported from thesame general area at the following
localities: Ominato, Mutsu Bay (HIRO,
1938, 1939b); Aburatsubo Bay, Misaki,
Kanagawa Pref.; Mukaishima Is., Hiroshima Pref.; Kagoshima, Kagoshima
Pref.; Nomosaki Penin. (UTINOMI, 1962)

Tanabe155

and Sasebo Bay, Nagasaki Pref.; Hakata
Bay, Fukuoka Pref. (HIRo, 1938; UTINOMI, 1970). B. kondalwvi was reported,
together with the above three species,
from the same general area at thefollowing localities: Hamana _ Lake,
Shizuoka Pref.; Mikawa Bay, AichiPref.;Tanabe Bay, Shirahama (HIRo, 1937,
1938; UTINOwlI, 1967); Hiroshima Bay,
Hiroshima Pref. (ARAKAWA, 1973).
The distribution of this species groupso far known in Japan is summarized in
Text-figure 3.
Habitat
The writer had an opportunity, in
March to April, 1974, to observe thehabitat at Tanabe Bay (Text-figs. 4 and5), where four species, B. amphitrite, B.

Bay~

B. amphitrite
B. albicostatus~j

Hosonoo...

Tachi otani

Text-fig. 4.

2 Km, ---~----"

Distribution of B. amphitrite and B. albicostatus in Tanabe Bay.

Toshiyuki YAMAGUCHI

156

albicostatus, B. reticulatus and B. kondakovi had been reported to occur by
UTINOlVn (1967).
B. amphitrite and B. albicostatus were
found side by side and widely distributed
from the lower to upper levels ofintertidal zone in the inner bay. The
upper part of the B. amphitrite and B.
albicostatus zone overlaps the lower partof the Chthamalus challengeri zone of
the uppermost intertide.
They are,however, especially abundant in the
middle level. Less commonly, these two
species are also found 1) on the rocky
shore of the middle to lower tidal zone
facing the open sea, but not where
there is strong wave action, 2) in theestuarine regions temporarily influenced
by fresh water at the lower tide, and3) attached to buoys and other floating
objects in the inner bay.
B. reticulatus is mainly found on
floating objects, but sometimes in thelower level of intertidal zone of rocky
shores. On rocky shores, however, thisspecies is rather rare in comparison with
B. albicostatus and B. amphitrite. Therefore, in the lower intertidal zone of
Tanabe Bay, B. amphitrite, B. albicostatus
and B. reticulatus are found side byside, a sympatric relationship in microgeographical sense. The same relationship was detected in Yokohama and
Aburatsubo, Kanagawa Pref.; Heta andHamana Lake, Shizuoka Pref.; Mukaishima Is., Kure harbor and Kurahashijima Is., Hiroshima Pref.; Kagoshima
Bay, Kagoshima Pref.; Tomioka, Amakusa Is., Kumamoto Pref.; Sasebo Bay,
Nagasaki Pref.
Another species, B. lwndalwvi, wasreported from the Tachigatani Inlet in
the Tanabe Bay by UTINOMI (1967).
At Tachigatani Inlet, B. amphitrite, B.
albicostatus and B. l·eticulatus werecommon, but the writer was unable to

Explanation of Plate 20(Figs. 1-4. anterior view of rostrum)
Fig. 1. Balanus amphitrite DARWIN (UMUT -RAS264) from Tanabe Bay (Lac. H), x 5.
Fig. 2. Balanus albicostatus PILSBRY (UMUT -RAS266) from Tanabe Bay (Lac. H), x 5.
Fig. 3. Balanus reticulatus UTINOMI (UMUT -RAS267) from Tanabe Bay (Lac. H), x 2.
Fig. 4. Balanus kondakovi TARASO\' and ZEYINA (UMUT -RAS269) from Hamana Lake (Lac.
G), x4.
(Figs. 5-S. longitudinal section)
Fig. 5. Balanus amphitrite DAR\\'IN (UMUT-RAS270) from Tanabe Bay (Lac. H), rostrum, x6.
Fig. 6. Balanus albicostatus PILSBRY (UMUT -RAS271) from Tanabe Bay (Lac. H), right lateral,x5.
Ffg. 7. Balanus reticulatus UTIN01vII (UMUT -RAS272) from Tanabe Bay (Lac. H), right lateral,x4.
Fig. S. Balanus kondakovi TARASOV and ZEVINA (UMUT-RAS273) from Hamana Lake (Lac.
G), right lateral, x 6.
(Figs. 9-12. photo by soft x-ray. a. rostrum, b. right lateral, c. right carinolateral, d. carina)
Fig. 9. Balanus amphitrite DARWIN (UMUT-RAS274) from Tanabe Bay (Lac. H), x3.
Fig. 10. Balanus albicostatus PILSBRY (UMUT -RAS275) from Tanabe Bay (Lac. R), x 2.
Fig. 11. Balanus reticulatus UTI:\,OMI (UMUT-RAS276) from Tanabe Eay (Lac. H), x 1.5.
Fig. 12. Balanus kondakovi TARASOV and ZEVINA (UMUT -RAS269) from Hamana Lake (Loc.
G), x2.
[Description to be published in Part 2J

YAMAGUCHI:

Plate 20

Japanese Balanoidea

9b

9c

lOa

lOb

10c

11 a

11 b

llc

12 a

12 b

12 c

12 d

681.

Taxonomic Studies on Japanese Balanoidea

157

N

TanabeBay

iJ

High tide line
L ow tid e line
Sand beachText-fig. 5.

Distribution of B. reticulatus in Tanabe Bay.

find B. kondakovi on the rocky shore,
floating objects or piles there.
In Hiroshima Bay, four species, B.
amphitrite, B. albicostatus, B. reticulatus
and B. Iwndakovi, were found together
on the cultured oyster (Crassostrea gigas)(ARAKAWA, 1973). The writer found an
association between B. kondakovi and B.
albicostatus near Yanagawa, Saga Pref.,in Ariake Bay, and between B. kondakovi,
B. amphitrite and B. reticulatus in Hamana
Lake, in both cases on the stakes ofbamboo. B. kondakovi is generally notfound on rocky shores, but rather on
piles, oysters and stems of plants etc.,
in the innermost part of the bay.' B.
kondakovi may be ecologically different
from three other species.
In conclusion, the distribution of B.

kondakovi does not completely coincide
witj1 that of the other three species in
geographic sense, and the range of B.
reticula/us does not completely agreewith that of B. amphitrite and B. albicostatus in vertical sense. However, astrictly sympatric relation between thesefour species of B. amphitrite group canbe observed not only in the Tanabe Bay
but also at many other localities inJapan. Therefore, geographical variation (subspecies) and/or ecophenotypic
effect do not explain the morphologicaldifferences. It is clear these four taxa
are distinct at the species level.
Mechanisms of reproductive isolation
Because of the definition of species(MA YR, 1963), reproductive isolation is

158

Toshiyuki YAMAGUCHI

the most important criterion for thediscrimination at the species level.
Species of superfamily Balanoidea arehermaphrodites, with the exception offour species of commensal forms (HENRY
and McLAUGHLIN, 1967; McLAUGHLIN
and HENRY, 1972) which have complemental male. Though there are some evidencesof self-fertilization (CRISP, 1954; BARNES
and CRISP, 1956; BARNES and BARNES,
1958, etc), method of reproduction ofhermaphrodite is III general crossfertilization. Fertilized eggs hatch into
nauplius larvae and are kept in the
mantle cavity of adult individual until
the larvae reach a certain developmental
stage.
Therefore, the period whenmature eggs or nauplius larvae arefound in the mantle cavity of adults
may be regarded roughly as breeding
season.
Whether or not the eggs
attain the mature stage can be judged by
the darkness in color of eggs. The difference in breeding seasons of closely
related species 1vfegabalanus rosa andM. volcano, which had been previously
treated as two subspecies of M. tintinnabulum, were interpreted as a possible
mechanism of reproductive isolation(Y AMAGUCHI. 1973). In fact, the breeding season of M. rosa is in March to
May (Spring) when the water temperature begin to rise, while that of M.
volcano corresponds to the period of
maximum water temperature in Julyto October (Summer to Autumn).
Reproductive seasons of members of
the B. amphitrite group in Japan has
not yet been sufficiently examined.
However, nauplius and cyprid larvae of
B. amphitrite and B. albicostatus werereared in the following seasons: May
to August (B. albicostatus) at Misaki by
ISHIDA and Y ASUGI (1937), July to
September (E. amphitrite) at Akiho,
Yamaguchi Pref. by HUDINAGA andKASAHARA (1942).
Larval settlementsof B. amphitrite, B. albicostatus and B.
reticulatus were observed in the following seasons: April to December (E.
reticulatus) by MAw AT ARI and KOBAYASHI (1954a, b), May to November (B.
amphitrite and B. reticulatus) by Y AMAMURA, KUWATANI and NISHII (1969),
April to December by Y AMAMURA(1972). All the above three observations
made at Ago Bay agreed that thelarval settlements of three species

Explanation of Plate 21(Photo by scanning electron microscope (SEM).)
Fig. 1. Balanus amphitrite DARWIN from Tanabe Bay (Loc. H). la. labrum, av., x 100., lb.
8th segment of right cirrus III, Iv., x300., lc. 8th segment of right cirrus IV, Iv., x250.
Fig. 2. Balanus albicostatus PILSBRY from Tanabe Bay (Loc. H). 2a. labrum, av., x 100., 2b.
8th segment of right cirrus III, Iv., x 200., 2c. 10th segment of right cirrus IV, Iv.,
x 300.
Fig. 3. Balanus reticulatus UTINOMI from Tanabe Bay (Loc. H). 3a. labrum, av., x 100., 3b.
8th segment of right cirrus III, Iv., x200., 3c. 10th segment of right cirrus IV, Iv.,
x 300.
Fig. 4. Balanus kondakovi TARASOV and ZEVINA from Kasaoka Bay, Okayama Pref. 4a.
labrum, av., x 100., 4b. 12th segment of right cirrus III, Iv., x 250., 4c. 11th segment ofright cirrus IV, Iv., x 250.
[Description to be published in Part 2J

YAMA GUCHI :Plate 21

,Japanese Balanoidea

10

1b

1c

20

2b

2c

30

3b

3c

40

4b

4c

681.

Taxonomic Studies on Japanese Balanoidea

attain their maximum activity in june
to October. Unfortunately, the exact
breeding seasons of these species were
not mentioned by these authors. Someof these may breed all the year round.
However, the breeding seasons of thesethree species seem to be in summer
and roughly coincide with each other.
Morphological difference between thesethree species cannot be explained by polymorphism, because only one of them isrepresented at some places. Hybrids
have never been found. Therefore, it
is interpreted that these three species
are distinct from one another. If they
are not reproductively isolated by differences in breeding seasons, it must
be by some other unknown mechanism.

Stratigraphic distribution
The oldest fossil of B. amphitritegroup so far known from japan istraced back to the Pleistocene.
Fossil B. kondakovi has been foundattached to the bank forming oysters
in the Pleistocene and the Holocene
sediments at Locs. 24a, 30d, and 34.
They are, however, not associated withother species of B. amphitrite group.
Fossil oyster bank which is represented
by autochthonous Crassostrea gigas suggests an environment of the innermost
portion of the embayment. Thus, thehabitat of B. kondakovi has been unchanged since the Pleistocene.
Fossil B. albicostatus has been foundin association with B. reticulatus in the
Pleistocene and the Holocene sediments
at Locs. 21, 28a-b and 30a, g. Thus,
the sympatry of these two species was
already established before the Pleistocene.
Living B. amphitrite is found in association with B. albicostatus and/or B.
reticulatus in various bays of japan. B.
albicostatus and B. reticulatus are foundon the shells from the shell mounds of159

the jomon age (3500-4200 Y. B. P.) (Locs.
19b, c) and the Kofun age (500-700
A. D.) (Loc. 19a). Yet, fossil or evensemifossil B. amphitrite* has never beenfound in japan. Therefore, it is highly
probable that B. amphit1'ite is a newcomer to the japanese coasts sometimes
in the historical age, as was assumed
by HIRO (1938).
According to the trustworthy observation made by HIRO (1938) before theWorld War II, B. amphitrite hawaiiensis(=B. amphitrite) and B. amphitrite
communis (=B. reticulatus) live on the
shore below the low tide line, while B.
amphitrite albicostatus (=B. albicostatus)
are always found in the intertidal zone.
In detail, B. amphitrite is dominant andB. reticulatus is rare in such bays underlittle influence of oceanic water as Kureand Sasebo harbors. On the other hand,
B. amphitrite is extremely rare and B.
reticulatus is a common species in thebays faced on Pacific Ocean.
The recent observations made by thewriter in japanese embayments, including Kure and Sasebo harbors, clarified
the fact that B. amphitrite as well as
B. albicostatus becomes abundant in theintertidal zone. B. reticulatus, however,is still found to the lower level ofintertidal zone of shore or on floating
objects. In conclusion, B. amphitrite
seemingly extended its habitat into theintertidal zone in various bays of japan
during the past half century. The apparent shift in the vertical distributionof B. amphitrite may be interpreted asa result of the combination of two
factors, first, the increase in individual
number of B. amphitrite over the generalarea of B. albicostatus and B. reticulatus* OHARA (1969) reported on occurrence of
B. amphitrite from the Pleistocene Semata
formation; however, it seem that the identification is incorrect.

160

Toshiyuki YAMAGUCHIin the intertidal zone and the zone below the low tide, and second, the thinout of B. amphitrite in the zone below
the low tide, due to the competitive
advantage of B. reticulatus over B.

amphitrite.
This assumption seems
interesting but entrust its proof tofurther observation.
[to be continued]

Explanation of Plate 22
Figs. 1-5. Balanus albicostatus PILSBRY from the Holocene Numa Formation (Loc. 28a). 1ab. whole wall (UMUT -CA8313), x 2., 2a-b. exterior and interior views of right scutum(UMUT-CA8314), x5., 3a-b. exterior and interior views of right tergum (UMUT-CA
8315), x5., 4a-b. exterior and interior views of rostrum (UMUT-CA8316), x3., 5a-b.
exterior and interior views of left lateral (UMUT-CA8317), x3.
Figs. 6-10. Balanus reticulatus UTINO~H from the Holocene Numa Formation (Loc. 28a). 6ab. whole wall (UMUT -CA8318), x 2.5., 7a-b. exterior and interior views of left scutum(UMUT-CA8319), x 5., 8a-b. exterior and interior views of right tergum (UMUT-CA
8320), x5., 9a-b. exterior and interior views of rostrum (UMUT-CA8321), x3., lOa-b.
exterior and interior views of right lateral (UMUT -CA8322), x 3.
Fig. 11. Balanus reticulatus UTINOMI (UMUT -CA8323) on Cerithideopsilla dj adj ariensis (MARTIN)
from the Holocene Ofuna Shell Bed (Loc. 30e), xl.
Fig. 12. Balanus reticulatus UTINO:VIl (UMUT -CA8324) on coral from the Holocene Taito-zaki
Formation (Loc. 21), xl.
Figs. 13-18. Balanus kondakovi TARASOV and ZEVINA from the Pleistocene Akasawa Silt (Loc.
42a). 13a-b. whole wall (UMUT -CA8325), x 1.5., 14a-b. exterior and interior views ofright scutum (UMUT -CA8326), x 5., 15a-b. exterior and interior views of right tergum(UMUT -CA8327), x 5., 16a-b. exterior and interior views of left lateral (UMUT-CA8328),... x 2.5., 17a-b. exterior and interior views of left carinolateral (UMUT -CA8328), x 2.5.,
18. shell wall (UMUT -CA8329) on Crassostrea gigas (THUNBERG), x 1.
[Description to be published in Part 2J

YAMAGUCHI :Japanese Balanoidea

Plate 22

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

September 15, 1977

New Series No. 107

CONTENTS
TRANSACTIONS
Teiichi: An occurrence of Ordosoceras in Jehol, Northeast
China, and a note on the Polydesmiidae .............................. 125

680.

KOBAYASHI,

681.

YAMAGUCHI,

Toshiyuki: Taxonomic studies on some fossil and recent
Japanese Balanoidea (Part 1) •..•...•.....•..••.•.....•...•••.......••• 135