ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 108

Palaeontological Society of Japan
November 15, 1977

EditorItaru HA YAMI
Associate editors Ikuwo OBATA and Tomowo OZAWA

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

The fossil on the cover: Original figure of Podozamites Reinii GEYLER,
1877, from the Tetori group. GEYLER'S description marked the onset ofmodern palaeontology in Japan.

All communications relating to this Journal should be addressed to the
PALAEONTOLOGICAL SOCIETY OF JAPAN,
c/o Business Center for Academic Societies,
Yayoi 2-4-16, Bunkyo-ku, Tokyo 113, Japan.
Sole agent: University of Tokyo Press, Hongo 7-3-1, Tokyo, Japan.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 108, pp. 161-201, pIs. 23-27, November 15, 19'i'i682. TAXONOMIC STUDIES ON SOME FOSSIL AND
RECENT JAPANESE BALANOIDEA (PART 2)*
TOSHIYUKI YAMAGUCHI
Geological Institute, Faculty of Science, University of Tokyo, Tokyo, 113

B*m, «''"In'O){tE t-o J: rf m!t 7 Y '/ ;f-~O)?Hi'i~i¥JlVf~ (-f:.0) 2) : ;: O)~)(-Cli B*0)
m!t7 Y '/ ;f-~O) 5 ~ Balanus rostratlls, B. crenatus, Megabalanus
rosa, M. volcano t-o J: rf Solidobalanus (Hesperibalanus) hesperius ~~c~.., Go B.
rostratus t-oJ:rf S. (H.) hesperius 0) 2 f.i/H".!: ttE* , , «''"In'O):fi!!.l2J.TO)?tm1l:'ftC.~?t~fL"(I.'t:.o Ln, L-tfL '=>O):fi!!.l2J.TO)?tm1l:'fO)t¥!~ c ~ fLt:.%~i¥J!Mf~li B*0) m!t t-o J: rf{tEO) 1 ""JO)J.fu:Jj{llj\f*fFO) .m!*J:j:ttc.,5! '=>fL Go ttE"?"( -tfL '=> O):fi!!.l2J.T O)ttE*O)?tmli~~1ib '=> ;[LtJ: I.
L n'
L;: O)~fuilHi5.lIJO)%~i¥J!Mf~tc. J: "? "(jlIijf'Jln'-tfL-t'fL ~ '=>tc.J.fulH!i¥Jtc.~?t ~ fLG 1lJ~'~~N~
Ii LtJ:"o M. volcano 0) {tEnqi C:1ib"(i<I'f!I*,Il'I;O)]!~t!:!:tC.~,5!~fLt:.o ;: 0) {tEtc.Ii M.
rosa n;1tJt L"(t-o I), ]!~t!:!:tC.lijllij~0)!t71lii¥JF>~il$n'7tT L"(I. ,t:.;: C ~B(f:7f-'" Go B. crenatus t-oJ:rf S. (H.) hesperius liii-5O){tE~c~n;B*-CliJ:j:t~t!:!:f}Jl!Jji-Cl®lGfLGo jlIijf.ij!
c tm:tEi -cO)ll'ifr"l'j;t~i&JO)rs'tc.:fi!!~!Mf~-?rr G%1(tc.v'1 C Iv c:~{tn'~~1ib GfLtJ:1. '0 7 Y '/;f~O);: 0) J: 'j tJ:%~O):Il1:l!Jjrs'tc.ht:. G't:~'~li*m~n'-tO)~~HIU~", GWtflli~~;t~J"J~
GIl'if-5!t~,,¥=i¥Jtc.~,*f,f-I.,~f;tHll!-#!<'" Go
rltDj/(f-Z'±~tdtE t-o J: rf'0

6.

group, while B. r. apertus, B. r. dalli and
B. r. sutural is in the latter group. The
former group is further subdivided into
two subgroups by the nature of radiiand rostrum: one is B. rostratus s. s.
which has conspicuously sunken radii
and at the same time has about 14 tubes
in the rostrum. The other subgroup
was characterized by the slightly sunken
radii and about 18 or more tubes in the
rostrum, and includes B. r. alaskensis andB. 1'. heteropus. B. r. alaskensis was defined by the larger size with diameterof 60-70 mm and with four or five pairsof spines on each segment of the posterior cirri, while B. 1'. heteropus was
smaller in size with diameter of about
18 mm and with four pairs of spines on
segments of posterior cirri.
BROCH (1922) described B. r. forma
eurostratus from Departure Bay, Van-

Revision of Balanus I'ostl'atus

Historical Review
Balanus rostratus HOEK was originallydescribed by HOEK (1883) from Kobe,
Japan. PILSBRY (1911, 1916) proposed,
beside B. rostratus s. s., five subspecies:
B. r. apertus, B. r. alaskensis, B. r. dalli,
B. r. heteropus and B. r. sutural is. He
divided B. rostratus into two groups.
One has transverse septa of longitudinal
tubes extending near to the base, andthe other has transverse septa confinedto the upper half of longitudinal tubes.
B. rostratus s. s., B. r. alasl?ensis and B.
r. heteropus were included in the former* Received May 26, 1977; read Jan. 31,
1976, at Kawatabi.
Continued from Part 1 (Trans. Proc. Palaeont. Soc. Japan, N.S., No. 107, pp. 135-160,
pIs. 19-22)
161

162

Toshiyuki YAMAGUCHI

couver Island. NILSSON-CANTELL (1932)
described B. r. var. spiniferus from Kobe,
Japan, an area roughly corresponds tothe HOEK'S type-locality of B. rostratus.
Taxonomic treatment proposed by
PILSBRY (1911, 1916) has been adopted
by many authors, CORNWALL (1925), HIRO(1935, 1939b), HENRY (1940, 1942), T ARASOY and ZEVINA (1957) and UTINOMI(1958). HENRY (1940) pointed out thatB. r. heteropus and B. r. spiniferus aresynonyms of B. r. alaskensis, without
giving taxonomic remarks. Thus, the<:urrent classification of B. rostratus
seems to be an intact application or aslight modification of PILSBRY'S old classification explained in the precedinglines.
l\t/orphological Examination of Recent andFossil Japanese B. rostratus
Transverse septa.- The distribution pattern of transverse septa in the longitudinal tubes was regarded by PILSBRY

as one of the criteria for the subspecificdivision of B. rostratus. This character
will be examined on Recent Japanesematerials collected from Loc. C1 (Mutsu
Bay). They are composed of living
specimens and fragmental skeletal elements derived apparently from one
population. The distribution pattern oftransverse in the longitudinal tubes varies
between two extremes. In one extreme
the distribution of transverse septa
covers entirely from orifice to base, but
the other confined to the upper part oflongitudinal tubes (pI. 24, figs. 7a-g).
Radii.- Even in the specimens from thesame population. variably sunken radii
are found among compartments which
are of uniform size. Thus the degreeof sinking of radii does not related tothe shell size, that is perhaps to growthage.
Longitudinal tubes.- The number of longitudinal tubes in rostrum compartmentof B. rostratus are compared with the

1/1
III

.c

....::I·t·a.!: 20. . . .

~....::I

01
C

B

.- I

- .. ~

o

III

.c

E 15

z

..

.,

.

.

.. .

~

::I

.

.6- .

.

_--.--.-.

-. J . .. ..
.... .. .,. . . ..
"

5

.
10

(N=108)
15

20
Width

mm

Text.fig. 6. Relation between shell size (width) and number of longitudinal tubes
in rostrum of B. rostratus from Loc. C 1.

682.

Taxonomic Studies on Japanese Balanoidea

163

sampling of bottom sediments· at the
type-locality (Kobe, Japan; St. 233 of
Voyage of Challenger). It is found that
this species is no more alive in the areaof the type-locality off Kobe harbor because of the polution of sea in thepresent time. Therefore, three samples
from the neighborhood of the typelocality, that is Locs. I and ]1-2 were
examined. In these three samples, the
95% confidence limits of means (i±t·O'.. )for the number of longitudinal tubes

shell size (width) of the compartment
in text-fig. 6. The number of longitudinal tubes seems to be a character which
is not much influenced by width of thecompartment. It is also observable in
x-ray photograph that the number does
not change during shell growth, though
an increase or a decrease in number isdetectable in some traumatized specimens. In order to ascertain the numberof the tubes in topotype specimens of
B. rostratus, the writer made dredge

Number of longitudinal tubes
10

Lac
A

20

1S

Recenl specimens

,

B1

I,

(N=ll) I+(108) ICl(S 2) I

C2

I

1

C3

o

1

I

(14) ,

1+
I

I

(28)

I

1 (44) ,JII(27) I

J 2

I

28c
6b

8c

200
,

24 i

~

(64)

I I-----e,$-I-----''''-'''L-il

3S

I (S

6) ,

I
10

1S

20

Text.fig. 7. Mean, 95% confidence limits of the mean (x±t·u.. ) and twice standard
deviation of the mean for number of longitudinal tubes in rostrum of living and fossil
B. rostratus of Japan.

Toshiyuki YAMAGUCHI

164

Table 4. Statistic values of number of longitudinal tubes in rostrum of living andfossil B. rostratus HOEK. N: number of individuals examined, OR: observed range, x: mean, s: standard deviation, ax: standard error.

I
I

Living specimens

Loc.

N

OR

A

J2

11
170
108
52
14
40
65
28
21
44
27

12-18
10-20
11-20
11-20
11-21
11-23
12-22
13-22
11-22
14-25
14-21

28c
6c
8c
20a
24i
35

15
59
23
40
64
56

14-21
11-24
11-17
12-23
12-21
13-25

Bl
Cl
C2
C3
D
E

F
I

11

I
Fossil specimens

overlap with one another (Text-fig. 7).
Thus these three samples are not essentially different in number of longitudinal
tubes, and are derived from the same
popUlation.
In the same way, the means and the
95% confidence limits for the numberof longitudinal tubes in various Recent
specimens from several mutually distantJapanese localities were examined (Textfig. 7, Tab. 4). The sample means range
from 15.09 (Loc. A) to 17.11 (Loc. ]1).
Three samples from the different environments in tbe Mutsu Bay, that is,from the sea bottom (Loc. Cl). on cultivated scallops hanging from a raft
(Loc. C2) and on the rocky shore (Loc.
C3), are compared. The 95% confidence
limits of the means among three samples
from Mutsu Bay overlap with one
another (Te~,t-fig. 7). Therefore, so far
as this character is concerned, geo-

s

I

15.09
15.12
15.56
15.88
16.86
15.70
16.44
16.36
15.10
17.11
16.68

1. 92
1. 62
1. 93
2.05
2.77
2.10
2.09
2.24
2.66
2.36
2.03

17.67
16.02
14.43
16.68
15.53
16.52

2.09
2.54
1. 73
2.35
2.06
2.72

II

0.58
0.12
0.19
0.28
0.74
0.33
0.26
0.42
0.58
0.36
0.39

II

0.54
0.33
0.36
0.37
0.26
0.36

graphical and ecophenotypic effects are
regarded as insignificant.
The observed ranges and twice thestandard deviation of the mean, for thenumber of longitudinal tubes in Recent
samples from various localities of Japan,
greatly overlap and include 14 and 18
which were regarded by PILSBRY (1911,1916) as a subspecies criterion. In conclusion, it is not possible to separate B.
rostratus into subspecies or any otherinfraspecific taxa by this character.
The number of longitudinal tubes infossil specimens was also examined. theresult is shown in text-figure 7. The
same conclusion can be ascertained by
the Pleistocene and the Holocene specimens.
Distribution
The distributions of B. rostratus so far
known are summarized in text-figures 8

682.

Taxonomic Studies on Japanese Balanoidea

and 9. Depth distribution ranges fromOm to 168m.
From the text-figure 9, it is seen that,in Japanese waters, B. rostratus s. s. andB. r. eurostratus are distributed in amacrogeographically sympatric relationship. The distribution extends to Sagami
Bay on the Pacific side and to the
Tsushima Straits on the Japan Sea side,a branch further extends into the Inland
Sea of Japan through the Kanmon
Straits.
B. rostratus forma eurostratus was
originally described by BROCH (1922) from
Depature Bay, Nanaimo, Vancouver Is.
He mentioned that his specimens" quite
agree with the Japanese variants which
PILSBRY (1916) takes to be the typicalB. rostratus i. e. the forma eurostratus".

165

Therefore, B. r. eurostratus seems to becoincided with B. rostratus s. s.
NILSSON-CANTELL (1932) has described
B. r. var. spiniferus from Kobe, the typelocality of B. rostratus s. s. This variety
is embedded in sponges and characterized
by a few downwardly projecting spines,
and thus coincides with the PILSBRY'S
B. r. apertus (UTINOMI, 1958).
B. r. rostratus and B. r. apertus arefound by UTINOMI in the different dredges
at the same locality in Sagami Bay,central Japan. In fact. B. r. rostratus
with the smooth surface and with thespinose surface are found together inthe same dredged sample in Mutsu Bay(Loc. Cl). Thus B. r. var. spiniferus
which does not differ very much fromB. rostratus s. s. in morphology otherB. rostratus 5.5.
• B. r. a/askensiso B. r. apertus
A
B. r. do/Ii
x B. r.eurostratus• B. r.heteropus+ B. r. spiniferus """~.

A"::~·~ i"",.... 'X. ·i~"-.~---+------+------!-------!-----:"~ d;A}- • 'fjV"<.Y

Text-fig. 8. Distribution of B. rostratus and its" subspecies" in the Recent" sea.
[Compiled from CORNWALL; 1925, 1955, HENRY; 1940, 1942, HIRO; 1932b, 1933, 1935, 1939b,
HOEK; 1883, NILSSO:-.r·CANTELL; 1932, 1958, PILSBRY; 1911, 1916, TARASOV and ZEVINA;
1957, UTINOMI; 1955, 1970, YAMAGUCHI; 1972MS, 1974MS].

Toshiyuki YAMAGUCHI

166• Balanus rostratus s. s.

o B. r. aper tus
A B. r. dalli

X a ~ eurostratus+ B. r. spiniferus

Text-fig. 9. Distribution of B. rostratus and its" subspecies" in the Recent sea ofJapan. [Compiled from HIRO; 1932b, HOEK; 1883, KRUGER; 1911b, NILSSON-CANTELL;
1932, 1958, UTINOMI; 1955, 1970, YAMAGUCHI; 1972MS, 1974MSJ.

than spines, seems to be a mere phenotypic variation of B. l-ostratus.
Three subspecies, B. r. alaskensis, B. I".
apertus and B. r. dalli, has been foundin ,Puget Sound, west coast of British
Columbia, Alaska, Bering Sea, AleutianIslands, south coast of east Siberia,
Kamchatka, the Sea of Okhotsk and
Sakhalin. The localities of B. rostratus
and its subspecies described by PILSBRY(1911, 1916) are shown in text-figure 10.
The vertical distribution of three sub-

682.

Taxonomic Studies on Japanese Balanoidea

167+• B. r. a/askensiso B. r. apertus
I>.

B. r. da/Ii• B. r. heteropus..

bo 0 iIJ··A
00"
C

Text·fig. 10. Type and sampling localities of B. rostratus and its" subspecies"
described by PILSBRY (1911, 1916). a, b, c, d, and e: type localities of B. rostratus
s. s., B. r. apertus, B. r. dalli, B. r. alaskensis B. r. heteropus.

species is similarly restricted to shallow
seas less than 100 m in depth. They arefound in many places in macrogeographically sympatric relation. In fact, there
are records of the coexistence of certain
subspecies, such as B. r. alaskensis, B. r.
apertus, and B. r. dalli in Abachinsky
Bay, Kamchatka Peninsula (T ARASOV andZEVINA, 1957).
Further, B. r. dalli and B. r. eurostratus
occur together in Onagawa Bay, northern Honshu (HIRO, 1939b). B. r. rostratus
and B. r. apertus coexist in Sagami Bay(UTINOMI, 1958). And B. r. eurostratus
lives amidst the range of B. r. alaskensis
in the British Columbia area.
HENRY (1942) pointed out that the
number of longitudinal tubes in B. r.
alaskensis from Seymour Inlet, British
Columbia ranges from 15 to 20. This-rangeof variation of B. r. alaskensis from Seymour Inlet fall in that of Recent Japanese
B. rostratus s. s. HENRY also synonymized
B. r. heteropus and B. r. spiniferus with
B. r. alaskensis.
In conclusion, all the subspecies whichhave been proposed in B. rostratus do
not fit the current geographical definition of subspecies at least from themacrogeographical point of view.

Summary on Taxonomic Revision
The PILSBRY'S (1911, 1916) proposal of
five subspecies B. r. alaskensis, B. r. apertus, B. r. dalli, B. r. heteropus and B. r.
suturalis were not accompanied by any
biogeographical study of this species,
but based only on morphological differences. However; diagnostic charactershe employed for defining these' sub-

Toshiyuki YAMAGUCHI

168

species, such as the distribution patternof transverse septa, the nature of radii
and the number of longitudinal tubes inrostrum are found to be within the range
of intra populational variation of Recent
Japanese B. rostratus s. s. An analysis
of distribution demonstrated that thesesubspecies do not fit current geographical criteria for sUbspecies. Therefore
these subspecies are here regarded assynonymous with B. rostratus s. s. Although further study may prove theexistence of geographical subspecies in
B. rostratus, the proof needs an extensive biogeographical as well as morphological study on the material from outside of Japan.
Fossil Occurrence
Generally, fossil B. rostratus is found.'as fragmental skeletal elements whichshow allochthonous occurrence. But
autochthonous occurrence is often met
with in certain localities, such as Locs.
6b, 20a, 25b, 28c and 29. Three types
of autochthonous barnacle occurrences
can be recognized. 1) Barnacles which
are found on fossil bank-forming oysters(Crassostrea gigas at Locs. 6b and 25b;
Crassostrea nipponina at Loc. 28c). 2)
Barnacles which are embedded in massive mud as colonial aggregations ingrowth position (Loc. 29). 3) Barnacles
which are found in coarse sand (Loc. 20a).
Most of fossil barnacles found from
Loc. 20a are embedded in sand andattach to the external surface of Chlamys
nipponensis. Articulate valves of Chlamys
are occasionally found in the sand, ingrowth position with umbones directed

Jcm

Text-fig. 11. Mode of occurrence of fossil B. rostratus growing on autochthonous
Chlamys nipponensis (Loc. 20a, Pleistocene Semata Fm.).

682.

Taxonomic Studies on Japanese Balanoidea

downward and attached to other individuals or objects by byssus (Text-fig.
11; PI. 24, fig. 8; cf. PI. 24, fig. 9). In
such cases, fossil barnacles are foundon the external surface of both valves,often with a set of opercular valvespreserved in the shell cavity. Some
individuals may have been transported,but the distance must not have been
great.
The examination of 22 individuals of169

Chlamys, revealed that the number andmode of distribution of barnacles onthem was remarkably different between
left and right valves (Text-fig. 12; PI.
24, figs. 1-3). Namely, left-valve barnacles
are distributed on the entire surface,
while right-valve barnacles are restricted
to the posterior half of the externalsurface.
A similar trend was also observed on
the disarticulate valves of allochthonous..

...... .. e.- ..;. ..
•

• e•

..

I." . -:-: e... ..
·.....:;.
...... . -....-.. .
........... ...:.::......
... ..-. ,.. ., .
~..

.\I· •••

Autochthon

A

B

1em
~

..
.,. .. ..
... .. ......
·..
.... -.... :...
....·..... .. ....
.....; . .
•"

••• I

...... ..
...

••. .. ...
... .. .........
._...- . ..
.-::. . .. . ...-.
...:....
.:....
.. .. .-.:. ...

e• • • •_. ..

cAllochthonoText.fig. 12. Mode of distribution of fossil B. rostratus growing on Chlamys nip.
ponensis (Loc. 20a, Pleistocene Semata Fm.). A, B: left and right valves of 22 indivi·
duals of autochthonous Chlamys. C, D: left and right valves (11 specimens in eachvalve) of allochthonous Chlamys.

Toshiyuki YAMAGUCHI

170..:.. .
,-. -",.:. ..
.
... .: .
. .c:...: .
•

I

;.

.~.

~..

..
.........:...... .. .........
.... .. ...,:.....
.. ..:........ . .......
.
..... ...: .. ... -:...
'..-. .....
..'...
; .-,.....
. . ..
~:)f. ,~ ••:

.,.~

....:
... . ... ·.....
..
...

· ..
.... ........
·. ..

.-.: •• ,".

...

. ... .

..: ...
.-.:...
.... . "',,;.-.
.
:;... ., . :4'-'.. - , ."... :-..

1 em' Et--....J.
F'

Text-fig. 13. Mode of distribution of living B. trigonus growing on Chlamys nipponensis (Loc. C2). E, F: left and right valves of 10 individuals of living Chlamys.
x : B. rostratus, 6,: Megabalanus rosa. [Positions of barnacles on 10 individuals ofChlamys are plotted on the figure.]

Chlamys (Text-fig. 12). Thus, barnacles
found on disarticulate valves had grown
there during the life time of Chlamys.
Recent barnacles on living specimensof Chlamys nipponensis were examinedfrom Loc. C2. Most, on the 10 individuals of Chlamys, were Balanus trigonus
DARWIN, only a few individuals beingreferable to B. rostratus and Megabalanus
1'Osa. None the less, the mode of distribution of these barnacles is very
similar to that of the fossil barnaclesfrom Loc. 20a (Text-fig. 13). Therefore,
the distribution must be controlled by
the living position of Chlamys (pI. 24,
figs. 1-6, 8, 9).
Fossil barnacles at Loc. 20a are generally attached to Chlamys nipponensis,
but some of them are found on Anomia
lisekei, Area miyatensis, Patinopecten·
tokyoensis and Granulifusus musashiensis.

7. Revision of Solidobalanlls(Hesperibalanlls) hesperills
Solidobalanus (Hesperibalanus) hesperius
and its Subdivision
PILSBRY (1916) described S. (H.) hesperi us, the forma laevidomus and the subspecies nipponensis. The type specimenof S. (H.) hesperius s. s. is from the Bering Sea (Albatros St. 3483). S. (H.) h.
nipponensis was described from off Daikoku-saki, northern Japan (Albatros St.
3768), while S. (H.) h. forma laevidomus
are based on the specimens from San
Juan Is., Puget Sound.
The following characters were considered diagnostic by PILSBRY (1916). 1)
The external surface of S. (H.) hesperius
s. s. is typically ribbed but in some casessmooth; in S. (H.) h. forma laevidomus itis typically smooth but in a few cases
ribbed near the base; in S. (H.) h. nip-

682.

Taxonomic Studies on Japanese Balanoidea

ponensis it is always smooth. 2) The
scutum of S. (H.) hesperius s. s. has closeand sharp growth ridges; in S. (H.) h.
forma laevidomus, the ridges are very
low and more widely spaced growthridges; in S. (H.) h. nipponensis, they
are closely spaced. 3) The tergal marginof scutum is longer than that of basal
margin in S. (H.) hesperius s. s., but shorter in S. (H.) h. forma laevidomus; in S.
(H.) h. nipponensis the margin are nearly
equal in length. 4) The segment of posterior cirri has three or four paired
spines, the lower pair being minute in
S. (H.) hesperius s. s.; on the other hand
S. (H.) h. nipponensis has three paired
spines, the lower pair being very minuteor absent in some cases. PILSBRY did
not described this character for S. (H.) h.
forma laevidomus.
PILSBRY (1916) mentioned that thesemorphological characters were variable(p. 200), but thought it "desirable tosignalize by name the presence of amodified form" and proposed the newsubspecies, S. (H.) h. nipponensis.
japanese Living and Fossil S. (H.) hesperius
Many specimens of living S. (H.) hesperius were collected from Funka Bay,
Hokkaido, northern japan (Loc. B2),
attach to Turritella fortilirate SOWERBY(pI. 27, fig. 5), Macoma calcarea (GMELIN)(pI. 27, fig. 4), Panomya arctica (LAMARCK),
Mya truncata LAMARCK, Eunatica pila
(PILSBRY), Neptunea arthritica (BERNARDI), etc.
1) The external surface of shell wall
(pI. 27, figs. 1-5) show a wide range ofvariation from smooth type (nipponensistype) (pI. 27, figs. 2e-f, 4, 5) to strongly
ribbed type (hesperius s. s. type) (pI. 27,
figs. 3e-i, 4, 5). In the samples from
Funka Bay, large individuals (larger thanabout 5 mm in basal diameter) are generally ribbed, small individuals (smaller

171

than 3 mm in basal diameter) are usually smooth, and many of medium sizedindividuals are intermediate, having ribs
only near the base (laevidomus type).
Sometimes, however, a smooth externalsurface is observed in large individuals,
and ribbed surface are found in small
individuals.
2) The external surface of scutum injapanese living specimens shows a wide
range of variation, having either widely
spaced growth ridges (laevidomus type)(pI. 27, fig. 2a) or closely spaced growthridges (hesperius s. s. and nipponensis
types) (pI. 27, fig. 1a).
3) The proportion between basal andtergal length of scutum, in japaneseliving specimens, shows intrapopulationalvariation covering both the nipponensis(pI. 27, figs. 2a-b) and the hesperius s. s.
type (pI. 27, figs. 1a-b, 3a-b), with thelaevidomus type in between.
4) The median segments of cirri IV
to VI have two large paired spines andone to three short paired spines on theanterior face.
In the shell wall and scutum, two extreme forms are found in living specimens of Funka Bay. However, no significant difference can be detected in themorphology of animal parts between two
extreme forms. There are some specimens showing mosaic characters ofhesperius s. s. and nipponensis in which
the shell wall possesses strong ribs but
the basal length of scutum is longer
than the tergal length. In fossil specimens of S. (H.) hesperius, smooth andribbed individuals (pI. 27, figs. 10a-llb)
are associated.
In conclusion, intermediate forms withvarious combinations of characters between three types which are morphologically comparable with S. (H.) hesperius
s. s., S. (H.) h. forma laevidomus, and S.
(H.) h. nipponensis of PILSBRY'S senseToshiY1lki YAMAGUCHI

172

are found in living population of Funka
Bay. Therefore, it seems taxonomicallyirrational to classify S. (H.) hesperius
into these subspecies or for mae by thesemorphological characters.
8. Additional observation on
11fegabalanus tinti1l1wbul1l11l

group
Living ;vI. rosa and M. volcano, treatedas subspecies of M. tintinnabulum by
PILSBRY (1916) and later authors, aredistinct both in morphology, habitat andbiochemistry (composition of isozymes).
They are largely overlapping geographic
ranges. Where the ranges overlap, theyoccupy similar but in detail different
habitat; in general M. volcano lives on
rocky shores at the lower tide zone,
while bulk of M. rosa on floating objects.
However, it is occasionally observable
the physical attachment between the two
species. Therefore, they are sympatricin macro- and microgeographical sense.
Breeding season between these two
phena is completely separable. These
facts seems to furnish important proof
that two phena are regarded as distinct
at the species level (YAMAGUCHI, 1973).
In fossil specimens, macro geographicalsympatry may be assumed, if two or
more phena occur at the same horizon.
For example, fossils of lvI. volcano wererecently obtained together with lvI. rosa
from the same horizon of the same outcrop of the Pleistocene Nakoshi Sandstone of Okinawa Is. (Loc. 44). An individual of fossil lvI. rosa is attached tothe colony of M. volcano (pI. 26, fig. 13).
This observation strongly suggests the
microgeographical sympatry between thetwo species. Thus the speciation seems
to have been accomplished already in
the Pleistocene.

9. Stratigraphical distribution ofJapanese Balanoidea
Information on the stratigraphical distribution of japanese Balanoidea isscarce. Text-figure 14 summarizes thepresent knowledge on the distribution
in japan. Many species are found living
around japan. However, number of
species found in the Pleistocene is halfof that of living species. The species
limited to the Recent seem to belong
either to the rare species with limited
distribution or the immigrant species.
Chirona (Striatobalanus) kriigeri, Ogasa,vara Is.; Solidobalanus (S.) pseudauricoma, Ogasawara Is. and off Nagasaki
provide examples of species with restricted distributions. Most of symbiotic
species are too rare to be encounteredas fossils. B. eburneus, B. improvisus
and B. amphitrite are immigrant species.
The first two species arrived in thejapanese region after World War II,
while B. amphitrite arrived but in historical times, as explained in the preceding pages.
Around the Plio-Pleistocene boundary,
the occurrence of many warm water
species is noticable. It is interesting tonote, however, such warm water species
as B. concavus withdrew from japan atend of the Pliocene. On the other hand,
cold water species such as Chirona(Chirona) evermanni disappeared fromthe japanese region at the end of Pliocene or sometime in the Pleistocene, but
still persists in the northern part of the
Okhotsk Sea.
Not many species of Balanoidea canbe traced back into the Pliocene andMiocene. This is perhaps because of
the incompleteness of fossil record. Yet,
microgeographical sympatry can be detected between Megabalanus rosa andM. volcano even in the Pleistocene.

682.

Taxonomic Studies on Japanese Balanoidea

173

"c:

"
c:

Q)

c:
04

....c:
....>
-'

Ul

....

CJ

......,u u

Q)

D-

o

Of.

.....
..0
E

>,
<Jl

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c: III
CJ

Q.i
.....

E

>,u
<rl

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..oDE

Ul

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III

....."
u

Q)

DUl

cO

"

Ul

c:

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0

.....

Of;

.... .~"
III

Vl

o

G.i
~

4-1;'-

IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV
IV

C
C
C
C
C

Armatobalanus (A. ) cepa
Armatobalanus (A. ) quadrivittatus
Armatobalanus (A. ) allium
Conopea calceola
Conopea cymbiformis
Conopea navicula
Conopea cornuta
Conopea granulata
Balanus poecilotheca
Membranobalanus cuneiformis
Balanus eburneus
Balanus improvisus
Balanus amphitrite
Balanus albicostatus
Balanus reticulatus
Balanus kondakovi
Megabalanus volcano
Megabalanus rosa
Balanus trigonus
Chirona (Striatobalanus) tenuis
Chirona (Striatobalanus) amaryllis
Chi rona (Striatobalanus) krugeri
Solidobalanus (S.) hawaiensis
Solidobalanus (S. ) social is
Solidobalanus (S.) pseudauricoma
Solidobalanus (S . ) compressus
Balanus concavus
Balanus rostratus
Balanus crenatus
Chirona (C. ) evermanni
Semibalanus cariosus
Solidobalanus (Hesperibalanus) hesperius
lSalanus sp. aff. "amphItrite"
Chirona sp . A
Chirona sp. B
Solidobalanus? sp.
Megabalanus sp. A
Megabalanus sp. B

Q)u.....0
0

=

"u0

"u

c:

....:;;:0.....

.........0

..,CJ

"'-

0..

-'

..,
....
Ul
Q)

Q)

"u

"c:
"u

....:;;:0
:;;:
I

<rl
L1J

----

---

---

-- - r------ i - --- r---- - -- --- - --i - - ---

-

- -

-

'---

~

Text.fig. 14. Stratigraphical distribution of Japanese Balanoidea. W: warm water
species, C: cold water species.

Morphological stability of some species(B. crenatus and Solidobalanus (Hesperibalanus) hesperius) can trace back to theearly or middle Miocene.
B. crenatus is known from a few stations in northern Japan. Where extant
individuals are usually found attachedto the gastropod shells (Neptunea, Pugilina and Phalium) and the carapace of acrab (Paralithodes). Fossil occurrence
are also restricted to the northern Japan,
where usually fragmental skeletal elements are found in shelly sandstone of
Miocene age. When the compartments,

174

Toshiyuki YAMAGUCHI

scutum and tergum are recovered fromthe sediments, they clearly show thediagnostic characters as defined by theliving specimens. These characters include the nature of the external surface
and cavity between sheath and innerlamina of the compartments, growthridges, articular ridge, adductor ridgeand pit for adductor muscle of the scutum, and nature of spur fasciole, spurand basal margin of tergum. Thus, it
is evident that this species has survived
from the early or middle Miocene innorthern Japan without notable modification in morphology. Long-term stability
in morphology may furnish interesting
problems in paleobiology concerning themechanisms maintaining the status quoof a phenotype.

collections from various localities of theworld, and on thorough analysis of morphological characters of B. amphitrite
complex.
Although the works have been done
independently by HENRY and McLAUGHLIN (1975) and YAMAGUCHI (1974, MS),the classifications resemble strikingly
each other both in evaluation of morphological characters and in the conclusion. Therefore, to avoid overlapping ofsystematic descriptions, only a few supplementary descriptions are appended tothis paper. For full synonymies, diagnoses and detailed descriptions, refer tothe comprehensive work by HENRY and
McLAUGHLIN (1975).

Present article is based on YAMAGUCHI(1974, MS) which was submitted to the
Geological Institute, University of Tokyo
as a partial fulfilment of the requirements for doctorate degree in December. 25, 1974. YAMAGUCHI (1974, MS) has reexamined classification of Japanese B.
amphitrite group in a part of his taxonomic studies on Japanese fossil andRecent Balanoidea. The result is based
primarily on a taxonomic evaluation ofmorphological diversity of the group, but
is also supplemented by observations on
macro- and microgeographic distribution
and on seasons of larval settlements.
Shortly after my work, HENRY and
McLAUGHLIN. published a paper on theB. amphitrite complex in October, 1975.
Their work (1975) is based on extensive

Superfamily Balanoidea LEACH, 1817

10. Systematic Descriptions

Family Balanidae LEACH, 1817
Genus Balanus DA COSTA, 1778
Balanus amphitrite DARWIN, 1854
PI. 19, figs. 1-2; pI. 20, figs. 1, 5, 9;
pI. 21, fig. 1; text.fig. 15

B. amphitrite hawaiiensis BROCH, ROSELL,
1973, p. 82, figs. 5a-f and 6g-k.
B. amphitrite amphitrite DARWI", SOUTHWARD,
1975, p. 6, figs. 1-3; HENRY and McLAUGHLIN, 1975, p. 30, text-figs. 10, 11, 13; pI. 1 ;pI. 5, fig. 9, upper row right; pI. 9, figs.
b, c; NEWl\'iAN and Ross, 1976, p. 62.

Supplementary descriptions.-Shell smooth
and with several colored longitudinal
stripes. Generally, central trapezoidal
area of rostrum and lateral compartment

Text-fig. 15. Balanus amphitrite DARWIN from Tanabe Bay (Loc. H), Shirahama,
Wakayama Pref. 1-11: UMUT-RA8274. 1-6. right cirri I to VI, Iv., x23.3., 1: cirrus
I, 2: cirrus II, 3: cirrus III, 4: cirrus IV, 5: cirrus V, 6: cirrus VI., 7. 6th to 9th segments of exopodite of right cirrus III, Iv., x 46.5. 8. 8th to 12th segments of exopodite
of right cirrus V, Iv., x46.5. 9. labrum, av., x46.5. 10. right mandible, Iv., x46.5.
11. right maxilla I, Iv., x46.5.

682.

175

Taxonomic Studies on Japanese Balanoidea

~

8

9

10

11

Toshiyuki YAMAGUCHI

176

white and without colored longitudinal
stripe. Adductor ridge sloping toward
occludent margin. Exopodite of cirrus
III with many conic teeth and denticles
on anterior face of each intermediatesegment, but exopodite of cirrus IV
without conic tooth, but with a few
denticles.
Fossil record.-None in Japan.
Balanus albicostatus PILSBRY, 1916
PI. 19, figs. 3-4; pI. 20, figs. 2, 6, 10;
pI. 21, fig. 2; pI. 22, figs. 1-5;
text-fig. 16

B. albicostatus PILSBRY, HEKRY and McLAUGHLI", 1975, p. 108, text-figs. 20, 22a;
pI. 10, figs. h-k; pI. 11, figs. m, n.
B. albicostatus albicostatus PILSBRY, NEWMAN
and Ross, 1976, p. 62.

Supplementary descriptions. - Shell withwhite longitudinal ribs. Longitudinal
tubes with transverse septa. Adductor
ridge vertiGally projecting on the innersurface of scutum. Short spinules onanterior face of exopodite of cirri III
and IV obscure.

(in part), 1962, p. 274; ROSELL, 1973, p.
79, figs. 4a-g.
B. reticulatus UTINOMI, SOUTHWARD, 1975,
p. 11, pI. 1, figs. 4-15; HENRY and McLAUGHLIN, 1975, p. 88, text-figs. 11, 18;
pI. 7, fig. d; pI. 8; pI. 9, figs. a, d, e;
NEWMAN and Ross, 1976, p. 64.

Supplementary descriptions.-Shell smoothwith numerous longitudinal narrow
stripes which are intersected by whiteand colored horizontal stripes. Adductor
ridge sloping toward occludent margin.
Fossil record.-Pleistocene to Holocene.
Pleistocene: Loc. 33a., Holocene: Locs.
21, 28, 30e, g, 32a-b, 33d and 38.
Balanus kondakovi T ARASOVand ZEVINA, 1957
PI. 19, figs. 6-7; pI. 20, figs. 4, 8, 12;
pI. 21, fig. 4; pI. 22, figs. 13-18;
text-fig. 18

B. amphitrite kondakovi TARASOV and ZEVINA,
NEW~IAl'i and Ross, 1976, p. 63.
B. kondakovi TARASOV and ZEVINA, HENRY
and McLAUGHLIK, 1975, text-figs. 21, 22b,c, f; pI. 11, figs. a-m.

PI. 19, fig. 5; pI. 20, figs. 3, 7, 11; pI. 21,
fig. 3; pI. 22, figs. 6-12; text-fig. 17

Supplementary descriptions.-Shell smoothwith many longitudinal narrow stripes
which are often intersected by whitehorizontal stripes. Parietal tubes of Japanese specimens in single row. Adductor
ridge sloping toward the occludent margin. Exopodite of cirri III and IV witha few denticles on anterior face of eachsegment.

B. amphitrite amphitrite DAR\n::-;, HARDI::-;GFossil record.-Pleistocene and Holocene.

Fossil record.-Pleistocene to Holocene.
Pleistocene: Loc. 30b., Holocene: Locs.
21, 28a, b., 30g and 33d. With humanremains: Locs. 19a-c.
Balanus reticulatus UTINOMI, 1967

Text-fig. 16. Balanus albicostatus PILSBRY from Tanabe Bar (Loc. H), Shirahama,
Wakayama Pref. 1-11: UMUT-RA8366. 1-6. right cirri I to VI, Iv., x 19.3., 1: cirrus
I, 2: cirrus II, 3: cirrus III, 4: cirrus IV, 5: cirrus V, 6: cirrus VI., 7. 6th to '10th segments of exopodite of right cirrus III, Iv., x38.6. 8. 9th to 13th segments of exopodite
of right cirrus V, Iv., x38.6. 9. labrum, av., x38.6. 10. right mandible, Iv., x38.6.
11. right maxiIIa I, Iv., x 38.6.

682.

177

Taxonomic Studies on Japanese, Balanoidea

3

11

9

178

. Toshiyuki YAMAGUCHI

Pleistocene: Locs. 30d and 42a., Holocene:
Locs. 33e and 34.
Balanus rostratus HOEK, 1883
PI. 23, figs. 1-13; pI. 24, figs. 1-3, 7-8;
text-fig. 19

Balanus rostratus HOEK, 1883, p. 152, pI. 13,
figs. 16-22; GRUVEL, 1905, p. 239, fig. 267;
PILSBRY, 1911, p. 73, pI. 12, fig, 6; PILSBRY, 1916, p. 138, pI. 36, figs. 1-2a; HIRO,
1933, p. 71; HIRO, 1935, p. 217, pI. 10,
figs. 1-2; HATAI, 1938, p. 97; CORNWALL,
1955, p. 38, figs. 30-31; OZAKI, 1958, p.
175, pI. 10, figs. 15-16; OHARA, 1969, p.
126; Iw AI and SIOBARA, 1969, p. 5, pl. 3,
fig. 21; UTINOMI, 1970, p. 357; YA}.,IAGUCHI, 1971, p. 125, pI. 5, figs. 8-13; NEWMA!'I and Ross, 1976, p. 61.
Balanus rostratus rostratus HOEK, UTINOyIl,
1958, p. 294.
Balanus rostratus alaskensis PILSBRY 1916,
p. 141, pI. 38, figs. 4-5; CORN\\, ALL, 1925,
p . ..18,1, pI. 5, fig. 5; HENRY, 1940, p. 21,
pI. 2, figs. 9-12; HENRY, 1942, p. 117, pI. 3,
figs. 13-14; TARASOV and ZEVINA, 1957,
p. 200, fig. 79.
Balanus rostratus apertus PILSBRY, UTINOMI,
1958, p. 295, text-fig. 5.
Balanus rostratus dalU PILSBRY, HIRO, 1935,
p.218, pI. 10, figs. 1-2; HIRO, 1939, p. 211;KOLOSVARY, 1943, p. 89.
Balanus rostratus forma eurostratus BROCH
1922, p. 320; HlRo, 1932b, p. 550, text-fig.
4; HIRO, 1939b, p. 210; TARASOV andZEVLNA, 1957, p. 200.
Balanus rostratus heteropus PILS13RY 1916, p.
142, pI. 36, figs. 7-8.
Balanus rostratus var. spiniferus NILSSONCANTELL, 1932, p. 20, text-figs. 8-9.
Balanus sp. OZAKI, 1958, p. 176, pI. 22, fig.
23.

Diagnosis.-Shell conical to subcylindrical, with smooth external surface andwhite to dirty white in color_ Carinolateral narrow. Rostrum well developed_
Radii narrow, their summits oblique tothe base. Scutum with prominent growthridges which are divided into squarish
beads by longitudinal striations. Pit for
adductor muscle weakly developed or
hardly visible. Tergum with wide andblunt spur.
Descriptions.-General appearances: Shell
conical to sub cylindrical, rather thick
and white to dirty white in color, withsmooth external surface. Radii narrow,generally deeply sunken below surfaceof parietes, their summits oblique to thebase. Orifice ovate, variable in diameter,
often same as base in diameter, andslightly toothed.
Compartments: Parietes conical, smooth,with longitudinal tubes having transverse septa. Carinolateral narrow. Rostrum well developed. Radii narrow,generally deeply sunken below surfaceof parietes. Summits of radii oblique tothe base. Alae very narrow, and theirsummits oblique to the base. Sheath
extending lower than the half of thewall.
Opercular valves: Scutum concave externally, color white to dirty white,
external surface with very prominent
growth ridges, superimposed over veryfine growth lines; growth ridges imbricating downwardly, divided into squarish
beads by longitudinal striations, depression between ridges deepen toward basal
margin; articular ridge long, but not so

Text-fig. 17. Balanus reticulatus UTINOMI from Tanabe Bay (Loc. H), Shirahama,
Wakayama Pref. 1-11: UMUT-RA8367. 1-6. right cirri I to VI, Iv., x 10., 1: cirrus I,
2: cirrus II, 3: cirrus III, 4: cirrus IV, 5: cirrus V, 6: cirrus VI., 7. 7th to 9th segments of exopodite of right cirrus III, Iv., x 32.6. 8. 10th to 12th segments of exopodite of right cirrus V, Iv., x32.6. 9. labrum, av., x32.6. 10. right mandible, Iv.,
x32.6. 11. right maxilla, Iv., x32.6.

682. Taxonomic Studies on Japanese Balanoidea.. ;;S-:;;

A;- 2

-.,~
£
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, "..

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11

179

180

Toshiyuki YAMAGUCHI
Table 5.

~umber of segments in each right cirrus of B. rostmtus HOEK from Loc.

C3 (Mutsu Bay). N: number of individuals examined, OR: observed
range, x: mean, exo: exopodite, end: endopodite.

1 -RA8277
2 RA8278
3 RA8373
4 RA8374
5 RA8375
6 RA8375
N

OR

x

exo

end

exo

end

28
25
27
26
28
24

14
14
14
13
13
12

19
16
15
15
15
16

16
15
16

6
6
24-28 12-14
26.3 15.0

14

14
14

6
6
15-19 14-16
16.0 14.8

I

exo

end

exo

19
19
20
18
I
18
iI 16

16
15
15
15
15
14

34

I

6
6
16-20 14-16
18.3 15.0

prominent; articular furrow narrow andshallow; adductor ridge low, in somecases weak; adductor muscle pit weakly
developed or hardly visible; lateral depressor muscle pit rather deep, in somecases shallow. Tergum fiat, white todirty white in color, with weak growthridges and lines and faint longitudinal
striae; spur wide at base and obliquely
truncated at end; spur furrow absent;
articular ridge moderately developed;
ridge running from apex toward basiscutal angle; scutal margin concave;
crests for depressor muscle weak ordifficult to observe.
Base: Calcareous, thin and with radiating tubes.
Mouth parts: Labrum with two or threeminute teeth on both sides of notch andwith numerous very fine setules. Mandi-

rCVI

rCIV

rCm

rell

rCI

18+
18+
23+
32
28

I rCV
end
I exo end
34
22+
16+
31
33
29

3
4
28-34 29-34
31. 3 31. 8

37
27+
18+
31
29
33

37
18+
20+
21+
35
31

4
3
29-37 31-37
32.5 34.3

exo

end

38
18+

38
22+
8+
31
27+
31

lIt:

I

35
20+
35

3
3
35-38 31-38
36.0 33.3

ble with three large acute teeth and aminute but blunt fourth tooth, and withclosely set setae convering the narrow
stripe between the main part of mandible and tooth edge. Mandiblar palp
oval, with closely set setae. Maxilla I
with two large spines near upper distal
angle; an another large spine near lower
distal angle; one to three medium sized
spines at upper distal angle; five to eight
medium sized spines between upper andlower large spines; small, narrow andshallowly concave notch just below two
upper large spines; and with closely set
setae between upper and lower edges.
Maxilla II oval with closely set setae.
Cirri: Number of segments in each cirrus
is shown in table 5. Exopodite of cirrus
I approximately one and a half times aslong as endopodite; number of segments

Text-fig. 18. Balanus kondakovi TARASOV and ZEVI:-<A from Kasaoka Bay, Ok ayama Pref. Coil. Dr. S. FCSE. 1-11: UMUT-RA8362. 12, 13, 15 & 17: UMUT-RA8363.
14 & 16: UMUT -RA8364. 1-6. right cirri I to VI, Iv., x 16.3., 1: cirrus I, 2: cirrus II,
3: cirrus III, 4: cirrus IV, 5: cirrus V, 6: cirrus VI., 7: 7th to 9th segments of exopodite of right cirrus III, Iv., x 45.8. 8: 16th to 18th segments of exopodite of rightcirrus IV, Iv., x45.8. 9. labrum, av., x33.8. 10. right mandible, Iv., x33.8. 11. rightmaxilla I, Iv., x33.8. 12. penis, Iv., x 10.8. 13. labrum, av., x33.8. 14. labrum, av.,x33.8. 15. right mandible, Iv., x3.'l.8. 16. left mandible, Iv., x33.8. 17. right maxilla I,
Iv., x33.8.

682.

Taxonomic Studies on Japanese Balanoidea

11---1. . -'"~~.

-:::ic;~:.of......

15

16--'-='""~M:"-

17.----..:~

13

181

8

182

Toshiyuki YAMAGUCHI

nearly twice of that of endopodite. Anterior face of each segment in endopodite of cirrus I and both rami of cirrus
II remarkably protuberant. Exopodite
of cirri II, and of III, nearly equal to theendopodite in length, but exopodites havea few more segments. In cirri IV to VI
with slender segments, expodites nearly
equal to endopodites in length, and in
number of segments; median segmentswith three to five paired spines, generally four, which gradually shorten fromdistal to proximal side. Cirri IV and V
with many fine spinules in a row at
distal border of each segment.
Penis: Long, with slender dorsal point
and with very fine setae.
Fossil record.-Pliocene to Holocene. Pliocene: Locs. 2c; 3a, b, d, f; 4a-c; 5a, b;
13a-c; 31 and 40a-e., Pleistocene: Locs.
6a-c: 8a-c; 16; 17a-c; 18a; 20a, b; 22ac; 23a, b; 24a, c-f, h-m; 25a, b; 26; 35
and 43.. Holocene: Locs. 28c and 29.
Balanus crenatus BRUGUIERE, 1789
PI. 25, figs. 1-19; text.fig. 20

Balanus crenatus BRUGUIERE, 1789, p. 168;DARWIN, 1854a, p. 261, pI. 6, figs. 6a-g;
GRUVEL, 1905, p. 240, figs. 268-269; PILS.
BRY, 1911, p. 75, pI. 14, figs. 1-9; PILSBRY,
1916, p. 165, pI. 39, figs. 1-5, pI. 40, figs.
1-6; NILSSO:'-l·CANTELL, 1921, p. 326;
BRacH, 1924, p. 78, fig. 27, pI. 1, figs. 3-6,
pI. 2, fig. 14; CORNWALL, 1925, p. 476, fig.
2, pI. 4, figs. A-h; HIRO, 1935, p. 219,
figs. 2a-f, pI. 10, fig. 4; HE:'-IRY, 1940, p.
19, pI. 1, figs. 9-11, pI. 2, figs. 1-2; HENRY,
1942, p. 105, pI. 2, figs. 5-11; CORNWALL,
1951, p. 329, pI. 4, fig. F, pI. 5, fig. A;

CORNWj\LL, 1955, p. 28, figs. 21-22; TARA.
SOY and ZEVINA, 1957, p. 205, figs. 22, 82,
83; DA v ADIE, 1968, p. 63, pI. 34, figs. 1-7,
pI. 36, figs. 4-5; NE\HIA:,\ and Ross, 1976,
p. 60.

Diagnosis. - Shell white and smooth.
Parietes with longitudinal tubes. Radii
narrow with denticulated sutural edges;
summits steeply oblique to the base.
Alae well developed, with denticulated
sutural edges and summits slightly oblique to the base. Calcareous vesicules
develop in the cavity between sheathand inner lamina. Scutum warped externally, with widely spaced and very
weak growth ridges; articular ridge very
prominent and reflexed; adductor ridge
very weak or hardly visible; pit for
adductor muscle deep. Basal margin of
tergum comes near to the tip of thespur on both sides.
Descriptions.-General appearances: Shellconial to tubuloconical, rather thick andsmooth in general; color white. Radii
narrow, summit steeply oblique to thebase. Orifice large, rhombodial and irregularly toothed.
Compartments: Parietes smooth in general, but in rare cases slightly ribbed
near the base. Longitudinal tubes large
at base, and with transverse septa.
Calcareous vesicles develop in the cavity
between sheath and inner lamina. Suturaledges of radii slightly oblique and denticulated. Alae well developed, and theirsummit slightly oblique to the base.
Sutural edges of alae rather thick andslightly denticulated.
Opercular valves: Scutum warped exter-

Text.fig. 19. Balanus ,"ostratus HOEK from Asamushi (Loc. C3), Aomori Pref. 110: UMUT-RA8376. 1-6. right cirri I to VI, Iv., x 10., 1: cirrus I, 2: cirrus II, 3: cirrus
III, 4.: cirrus IV, 5: cirrus V, 6: cirrus VI., 7. 13th to 17th segments of exopodite ofright cirrus VI, Iv., x25. 8. labrum, av., x25. 9. right mandible, Iv., x25. 10. rightmaxilla I, Iv., x 25.

682.

183

Taxonomic Studies on Japanese Balanoidea

7

Toshiyuki YAMAGUCHI184
Table 6.

Number of segments of each right cirrus of B. crenatus BRUGUIERE.
rCI

RA8330

rClI

rCm

rCVI

rCV

rCIV

exo

end

exo

end

exo

end

exo

end

exo

end

exo

end

I 18

19

I 12

12

I 15

12

I 27

32

33

31

I 34

34

,-----

A specimen from Paramushir Is., Kurile Islands; ColI. Mr. K. KOBA.

nally, with widely spaced very weakgrowth ridges and faint longitudinal
striae; color white; articular ridge very
prominent and refiexed; articular furrow
narrow and deep; adductor ridge very
weak or hardly visible; adductor muscle pit deep and narrow; lateral depressor muscle pit small and shallow, but
in some cases variable in size and depth.
Tergum fiat with weak growth lines andfaint longitudinal striae; color white;spur fasciole wide; spur wide, short androunded at end; width of spur approximately equal to one-third of basal
margin of tergum ; distance between
basiscutal angle and anterior face of
spur nearly equal to width of spur;crests of depressor muscle well developed; basal margin near tip of the spuron scutal side and away from on carinalside.
Base: Calcareous, thin and solid.
ivlouth parts: Labrum with three teeth
On both sides of notch. Mandible withthree acute and large teeth and minutebut bluntly rounded fourth and fifth
teeth. Mandiblar palp oval, with closely
set setae. Maxilla I with two large
spines at distal angle; five medium sized
spines between upper and lower large
spines; a small, narrow and slightly concave notch just below the two upperlarge spines; and with closely set setae
between upper and lower edges. Maxilla
II oval with closely set setae.
Cirri: Number of segments in each cirriis shown in table 6. Exopodite of cirrus
I approximately one and a half times aslong as endopodite, having twice as many
segments as endopodite; anterior face ofboth rami protuberant. Cirri II to VI,
exopodites nearly equal to endopodites
in length and number of segments.
Median segments of cirri IV to VI withtwo paired long and three or four paired
short spines on anterior face.
Penis: Nearly equal to or longer thancirri; with short a blunt dorsal point
and very fine setae.

Fossil record.-Early or middle Miocene
to Holocene. Early or middle Miocene:
Locs. 9a-d ; 10; 12b and 39., Late Miocene:
Locs. 7a, c-e, g-n; 11; 14; 15 and 41a,c-e, g., Pliocene: Locs. 2b; 3a, e-f; 4b-c ;5a-b; 7f and 12a., Pleistocene: Locs. 7f;
8a-c and 23b.
Recent distribution.-This species is widely distributed in the northern Pacific andAtlantic Oceans, and the Arctic Ocean,as shown in text-figure 21.
Genus Megabalanus HOEK, 1913

Megabalanus rosa (PILSBRY, 1916)
PI. 26, figs. 11-13
Balanus (Megabalanus) rosa PILSBRY, YAMAGUCHI, 1973, p. 130, pI. 6, figs. la- j, 3, 6,
7; pI. 7, figs. 3a-4b; pI. 8, figs. 1-2b; textfig. 10.
Megabalanus rosa (PILSBRY), NE\\"~IAN andRoss, 1976, p. 68.

Additional remarks.-For synonymy, description and fossil localities see YAMAGUCHI (1973). The following localities of682. Taxonomic Studies on Japanese Balanoidea

9

185

10

Text-fig. 20. Balanus crenatus BRUGUIERE from Murakami Bay, Paramushir Is.,
Kurile IsIs. Coli. Mr. K. KOBA. 1-10: UMUT-RA8330. 1-6. right cirri I to VI, Iv.,
x 20., 1: cirrus I, 2: cirrus II, 3: cirrus III, 4: cirrus IV, 5: cirrus V, 6: cirrus VI.,
7. 15th to 17th segments ot' expodite of right cirrus IV, Iv., x33.4. 8. labrum, av.,
x 33.4. 9. right mandible, Iv., x 33.4. 10. right maxilla I, Iv., x 33.4.

186

Toshiyuki YAMAGUCHI.. ~.tt~~;.: .. ' ...;i-'''':.

. . :Ii. ... - ......... ...

.....~Ii

Text-fig. 21. Distribution of B. crenatus in the Recent sea. [Compiled from BROCH;
1924, CORi\WALL; 1925, 1951, 1955, DARWIN; 1854a, HE:"RY; 1940, 1942, HIRO; 1935,
NILSSON-CANTELL; 1921, PILSBRY; 1911, 1916, 1921, STEPllENSEN; 1938, TARASOV; 1936,
1937, TARAsov and ZEVINA; 1957, UTINOMI; 1955, 1968, 1970, YAMAGUCHI; 1972MS,
1974MSJ.

fossils are newly added: Locs. 13a-b;
17a-c; 23-a-e, I, m; 25b; 28a-c; 30g; 31;
33c; 37a-b and 44. The writer foundsome errors in the previous descriptionof M. rosa (YAMAGUCHI, 1973). One of
the errors arose from misidentification:
the fossil specimens from Loc. 16 is not
M. rosa. Therefore, Loc. 16 should be

excluded from the fossil localities ofthis species. The geological age of the
Misaki Formation at Loc. 27 (YAMAGUCHI, 1973) is not Miocene but probably
Pliocene. Thus, the straigraphical rangeof M. rosa is rectified from Pliocene to
Recent.

682.

Taxonomic Studies on Japanese Balanoidea

Megabalanus volcano (PILSBRY, 1916)
PI. 26, figs. 1-10, 13

Balanus (Megabalanus) volcano PILSBRY, YAMAGUCHI, 1973, p. 133, pI. 6, figs. 2a-j, 4,5; pI. 7, figs. 1a-2b; pI. 8, figs. 1-2b; textfig. 2.
Megabalanus volcano (PILSBRY), NEWMAN andRoss, 1976, p. 69.

Additional remarks.-For synonymy anddescription see YAMAGUCHI (1973). Nothing had been known about the fossil
record of this species. However, undoubted specimens of M. volcano were
newly obtained from the Pleistocene
Nakoshi Sandstone of Okinawa Is. (Loc.
44). Thus, the Pleistocene should be
added to the stratigraphical range ofM. volcano.

Family Archaeobalanidae NEWMAN
and Ross, 1976
Genus Solidobalanus HOEK, 1913
Subgenus Hesperibalanus PILSBRY,1913
Solidobalanus (Hesperibalanus)
hesperius (PILSBRY, 1916)
PI. 27, figs. 1-18; text-fig. 22

Balanus (Hesperibalanus) hesperius PILSBRY,
1916, p. 193, figs. 1-ld, 7-8; HIRO, 1935,
p. 225, fig. 5; TARASOV and ZEVINA, 1957,
p. 228, fig. 89.
Balanus (Hesperibalanus) hesperius forma
laevidomus PILSBRY, 1916, p. 196, pI. 49,
figs. 2-5; pI. 50, figs. 12b.
Balanus (Hesperibalanus) hesperius nipponellsis
PILSBRY, 1916, p. 199, pI. 49, fig. 6.
Balanus hesperius PILSBRY, HIRO, 1939b, p.
212; CORNWALL, 1955, p. 35, fig. 27.
Balanus hesperius laevidomus PILSBRY, HE:"RY,
1940, p. 31, pI. 3, figs. 1-4.
Balanus (Solidobalanus) hesperius PILSBRY,187

HENRY and McLAUGHLIK, 1967, p. 47.
Balanus (Solidobalanus) hesperius laevidomus
PILSBRY, HENRY and McLAUGHLIN, 1967,
p. 47.
Balanus (Solido balanus) hesperius nipponensis
PILSBRY, HENRY and McLAUGHLIN, 1967,
p. 47.
Solidobalanus (Hesperibalanus) hesperius hesperius (PILSBRY), NEWMAN and Ross, 1976,
p. 51.

Diagnosis.-Shell white and smooth tostrongly ribbed. Parietes without longitudinal tube. Ra4ii narrow with denticulated sutural edge, summits slightly
oblique to the base. Alae wide andnearly parallel to the base. Orificeslightly toothed. Scutum flat, with weakand closely spaced growth ridges ingeneral; articular ridge very short; faintcallus-like projections observable between
articular and adductor ridges in someindividuals; articular furrow wide anddeep. Tergum with wide spur; crestsfor depressor muscle strongly developed.
Descriptions.-General appearances: Shell
conical to sub cylindrical, smooth tostrongly ribbed, and white in color. Radii
commonly narrow, but fairly wide insome sub cylindrical individuals, summits
oblique to the base. Orifice rhomboidal
and slightly toothed.
Compartments: Parietes smooth to strongly ribbed, without longitudinal tube. Alaerather wide and their summits approximately parallel to the base. Suturaledges of radii slightly denticulated, but
those of alae simple.
Opercular valves: Scutum flat, with closely to widely spaced weak growth ridges
and faint longitudinal striations on external surface; occludent and tergal
margins straight; articular ridge very
short, very prominent, reflexed andpointed at end; articular furrow wideand deep; adductor ridge very short andprominently to weakly projected; faintToshiyuki YAMAGUCHI

188

7

, ,

",

\

?~~.T'" 'C.
. ~--

9

10

Text.fig. 22. Solidobalallus (Hesperibalall11s) hesperius (PILSBRY) off Kafuran, west
Kamchatka. ColI. Mr. K. KOBA. 1-10: UMUT-RA8368. 1-6. right cirri I to VI, Iv.,
x 30., 1: cirrus I, 2: cirrus II, 3: cirrus III, "': cirrus IV, 5: cirrus V, 6: cirrus VI.,
7. 8th to 10th segments of exopodite of right cirrus VI, Iv., x50. 8. labrum, av., x50.
9. right mandible, h'., x 50. 10, right maxilla I, Iv., x 50.

Taxonomic Studies on Japanese Balanoidea

682.

callus-like projections observable between
articular and adductor ridges in someindividuals; adductor muscle pit narrowand deep. Tergum fiat, with weakgrowth lines and faint longitudinal striations; spur short, broad and rounded atend; width of spur approximately equal
to one third length of basal margin of
tergum ; distance between basiscutal
angle and anterior face of spur extremely short; spur facio Ie wide; articularridge very prominent, and running from
apex toward basiscutal angle; articularfurrow wide and deep; crests for depressor muscle well developed.
Base: Base calcareous, thin and solid.
Mouth parts: Labrum with three teeth
on both sides of notch. Mandible with
Table 7.

rCIl

rCIV

rCIIl

rCVI

rCV

exo

end

exo

end

exo

end

exo

end

exo

end

I 13

9

I 11

10

I 11

10

I 15

19

I 17

23

RA8295 ! 17
RA8296 : 12,
RA8297914RA8377
RA837812
12RA8379RA838013
RA8381
13
RA8382
9RA8383
16
I
RA8384 ! 17

9
7
6
9
8
7
8
7
7
8
8

9
I 9
8
10
8
10
I 10
8
7
12
10+

12
9
8
11
9
9
10
9
8
12
12

12
10
I
8
10
9
I
10
I 10
I
9
8
I
12
I
11

11

I 19

20
18
15
21
14+
21
23
17
17
21
23

N
OR
x

11

10
7-12
9.1

11
8-12
9.9

*RA8368---_.-

three large teeth, minute but bluntly
rounded fourth and fifth teeth, and withclosely set setae between upper andlower edges. Mandibular palp oval withclosely set setae. Maxilla I with twolarge spines at upper distal angle; oneor two large spines near lower distal
angle; three to six medium sized spines
between upper and lower large spines;
small, narrow and shallow concave notch
just below two upper large spines; andwith closely set setae between upperand lower edges. Maxilla II oval withclosely set setae.
Cirri: Number of segments in each cirrus
is shown in table 7. Exopodite of cirrus
I slightly longer than endopodite, andsegments slightly protuberant. Exopodite

Number of segments in each right cirrus of Solidobalanus (Hesperibalanus)
hesperius (PILSBRY).
rCI

1
2
3
4
5
6
7
8
9
10
11

189

11
9-17
13.1

6-9
7.7

I
I

i

11
8-12
9.9

9
7
10
9
9
10
8
9
13
11
11
7-13
9.7

I

15
13
17
15+
22
15+
14
13
19
24

9
10
13-24 15-23
17.3 19.6

22
24
18
26
20+
13+
25
20
19
26
28

25
23
22

I exo
I 32

end

i

25
25
23
23
23
24
14+
22
16
29
30

I

27 I
19+
19
13+
20
21
25
30

9
9
18-28 19-30
23.1
23.6

26

l 17+
22
25
23
19
14+
20
14
30
28

21+

10
9
14-30 16-30
23.0 24.0- - - - ----

* A specimen (RA8368) from off Kafuran, west Kamchatka; CoIl. Mr. K. KOBA.
Other specimens (1-11) from Funka Bay (Loc. B3). 3, 4, 7, 9 & 10: Shell wall is ribbed.
1, 5, 6 & 8: Shell wall is smooth. 2 & 11: Shell wall is entirely smooth, having ribs only
near the base.
N: number of individuals examined, OR: observed range, x: mean.

190

Toshiyuki Y AAJAGUCHI

Text.fig. 23. Distribution of Solidobalanus (Hesperibalanus) hesperius in the Recent
sea. [Compiled from CORNWALL; 1955, HIRO; 1935, 1939b, HENRY; 1940, 1942, KOLos.
VARY; 1943, KOLOSVARY and WAGNER; 1941, NILSSO",,·CANTELL; 1927, PILSBRY; 1916,
1921, TARASOV and ZEVINA; 1957, UTIXO~Il; 1955, 1970, YA~IAGl'CI-Il; 1972MS, 1974MS].

of cirri II, and of III, nearly equal toendopodite in length and in number ofsegments. Each ramus of cirri IV to VI
elongate; exopodite smaller than endopodite in length and in number of segments; median segments with two largepaired spines and one to three shortpaired spines on anterior face.
Penis: Longer than cirri with a shortand blunt dorsal point and with veryfine setae.
Fossil record: Early or middle Miocene
to Holocene. Early or middle Miocene:
Locs. 9b-c and 10., Late Miocene: Locs.
2d; 7a. d, g, i, k, nand 14.. Pliocene:
Locs. 1; 2a; 3a, d-e; 4a-c; 5a; 7f and
40a, c-f.. Pleistocene: Locs. 8a-c; 23b;
24a-c, e, i-j, I; 26 and 3l.
Recent distribution.-Records of this species are so far restricted to the northernpart of the Pacific as shown in textfigures 23.

11. Conclusion

The following species and species
groups of the fossil and Recent Balanoidea of Japan are studied taxonomically;
B. amphitrite group, B. rostratus, Solidobalanus (Hesperibalanus) hesperius, Megabalanus tintinnabulum group and B.
crenatus.
In the previous works B. amphitrite
was subdivided into various infraspecifictaxa according to the degree of morphological differences. In this articles,
however, they are treated as distinct
species, for the following reasons. The
morphological differences between them
are clearly recognizable. Their macro·
geographical distributions overlap. Where
the ranges overlap, they occupy similar
but in detail different habitats. B. amphitrite and B. albicostatus usually liveside by side in the mid-intertidal zoneof rocky shore. B. reticulatus is, in gen-

682.

Taxonomic Studies on Japanese Balanoidea

eral, found attached to the floating objects, but sometimes found on the lower
intertidal zone of rocky shore. B. kondakovi is not commonly encountered, but
lives on the brushwoods or the stems
of marine plants, etc. in the innermost
area of the embayments. Yet, thesedifferences in habitats are not mutually
exclusive. Where the habitats overlap,
the barnacles are found to live side byside in several localities of Japan. The
breeding seasons of both B. amphitrite
and B. albicostatus are almost all theyear round, and coincide with eachother. Therefore, some other isolating
mechanisms other than displacement ofbreeding season and/or habitat must be
operative.
Fossil B. albicostatus, B. reticulatus andB. kondakovi were found from the
Pleistocene and from shell mounds ofhistorical age, but B. amphitrite has
never been found as fossil or even semifossil. The absence of B. amphitrite in
the fossil record supports HIRO'S (1938)prediction that this species may havearrived on the Japanese coasts duringhistorical age. Comparisons of descriptions and illustrations of DARWIN (1854a)and HARDING (1962) strongly indicatesthat B. amphitrite var. communis of
DARWIN (1854a) includes two distinct
species, B. amphitrite DARWIN sensu
HARDING and B. reticulatus UTINOMI, as
noted by UTINOMI (1967).
B. rostratus had been previously subdivided into five subspecies, B. r. alaskensis, B. r. apertus, B. r. dalli, B. r.
heteropus and B. r. suturalis, based solely
on the morphological characters. The
diagnostic characters include the distribution pattern of transverse septa inlongitudinal tubes, number of longitudinal tubes in rostrum compartment, anddegree of sinking of radii (PILSBRY, 1911,1916). However, the morphological dif-191

ferences employed to discriminate thesesubspecies seems to fall within the limits
of individual variation of one sample
from one population. Therefore, it isunreasonable to classify B. rostratus into
subspecies by using these morphologicalcharacters, but this does not deny a
possibility of presence of subspecies in
B. rostratus. However, the sub specificname hitherto proposed are presently
placed in synonymy with B. l'ostratus s. s.
until the specimens from entire range ofdistribution are studied in detail. B. r.
forma eurostratus described from Vancover Is. by BROCH (1922) and B. r. var.
spiniferus from Kobe by NILSSONCANTELL (1932) are definitely synonymsof B. rostratus s. s.
Megabalanus tintinnabulum group wasstudied in detail by the writer. Living
M. rosa and M. volcano are distinct bothin morphology and biochemistry (composition of isozymes), are sympatric inmacro- as well as microgeopraphicalsense, and they are reproductively isolated (YAMAGUCHI, 1973). Occurrence offossil M. volcano together with M. rosa
from the Nakoshi Sandstone indicatesthat the speciation had been completed
and then the micro geographical sympatry
had also been established before the
Pleistocene. The two species had been
distinct also in morphology since thattimes.
Solidobalanus (Hesperibalanus) hesperius
had been divided into forma laevidomus
and subspecies nipponensis by morphological characters such as nature of external surface of shell wall, proportion
between basal and tergal length of scutum, number and nature of spines onanterior face of segments of cirri IV to
VI, etc. (PILSBRY, 1916). However, themorphological characters separatingthese two taxa are to be found within
the limits of the individual variation of

Toshiyuki YAMAGUCHI

192

of living or fossil populations. Therefore, for the same reasons utilized insynonymizing the subspecies of B. I'ostratus, it seems appropriate to do so here.
B. crenatus as well as S. (H.) hesperius
can be traced back to the early or middle Miocene without any detectablechanges in morphology. The stabilityof morphology extending over such along stratigraphical range may become
one of the interesting subjects of paleobiological studies.
Acknowledgements
The author wishes to thank Professor
William A. NEWMAN, Scripps Institutionof Oceanography, who kindly read themanuscript and provided invaluablecomments. Thanks are also due toEmeritus Professor Huzio UTINOMI of
the Kyoto University who also offeredvaluable advice.
The present study was carried out
under the direction of Professor Tetsuro
HANAI, the University of Tokyo. The
writer is much indebted to him forvaluable advice and encouragement. I
am also grateful to Drs. Itaru HAY AMI,
Kiyotaka CHINZEI and Takashi HAMADAof the University of Tokyo, Dr. Yasuhide
IWASAKI of the Kumamoto University
and Dr. Noriyuki IKEYA of the Shizuoka
University for their suggestions and discussions throughout the course of thisstudy. The sampling of living specimens
were made by the R/V Tanseimaru of
the Ocean Research Institute, Universityof Tokyo, the R/V Shunpumaru of theKobe Marine Observatory and R/VExplanation of Plate 23
Fig. 1. Balanus rostratus HOEK (UMUT-RA8277) from Asamushi, Mutsu Bay (Loc. C3). la-b.
whole wall, xL, lc-d. exterior and interior views of rostrum, x 1., Ie-f. exterior andinterior views of right lateral, x 1., 19-h. exterior and interior views of right carinolateral, xL, li-j. exterior and interior views of carina, xL
Fig. 2. Balanus rostratus HOEK (UMUT -RA8278) from Asamushi, Mutsu Bay (Loc. C3). 2ab. exterior and interior views of right scutum, x 2., 2c-d. exterior and interior views ofright tergum, x 2.
Fig. 3. Balanus rostratus HOEK (UMUT-CA8279) from the Holocene Numa Formation (Loc.
28c). 3a. whole wall, x 1.5., 3b-c. exterior and interior views of left scutum, x 3.5., 3d-e.
exterior and interior views of left tergum, x 3.5.
Fig. 4. Balanus rostratus HOEK (UMUT -CA8280) on Crassostrea nipponina from the Holocene
Numa Formation (Loc. 28c) , x 0.6.
Fig. 5. Balanus rostratus HOEK (UMUT -CA8290) from the Pleistocene Semata Formation (Loc.
20a). 5a-b. exterior and interior views of right scutum, x 2., 5c-d. exterior and interior
views of right tergum, x 2.
Fig. 6. Balanus rostratus HOEK (UMUT-CA828l) on Granulifusus 11lusashiensis from the Pleistocene Semata Formation (Loc. 20a) , x 1.5.
Fig. 7. Balanus rostratus HOEK (UMUT-CA8282) on Area miyatensis from the Pleistocene
Semata Formation (Loc. 20a) , x 1.
Figs. 8-13. Balanus rostratus HOEK from the Pliocene Hamada Formation (Loc. 3d). 8a-b. exterior and interior views of right scutum (UMUT-CA8283), x 4., 9a-b. exterior and in.
terior views of right tergum (UMUT-CA8284), x 4., lOa-b. exterior and interior view'sof rostrum (UMUT -CA8285), x 1.5., lla-b. exterior and interior views of left lateral(UMUT-CA8286), x1.5., l2a-b. exterior and interior views of left carinolateral (UMUTCA8287), x 1.5., l3a-b. exterior and interior views of carina (UMUT -CA8288), x 1.5.

YAMAGUCHI:

Japanese Balanoidea

Plate 23

682.

Taxonomic Studies on Japanese Balanoidea

Onokoro of the Kobe University. Special
thanks are extended to Professor Noriyuki NASU of the Ocean Research Institute, University of Tokyo and to Dr.
Yukito ENOMOTO of the Kobe Universityof permission to JOlll their survey.
Thanks are also due to Mr. Yoshiaki
MATSUSHIMA of the Kanagawa Prefectural Museum, Professor Masuoki
HORIKOSHI and Dr. Takeshi KAJIHARA of
the Ocean Research Institute, Universityof Tokyo, Dr .. Junji ITOIGAWA of theInstitute of Each Sciences, Nagoya University, and Miss Hiroko KOIKE of the
Department of Anthropology, Universityof Tokyo who kindly provided specimensof living and fossil barnacles.

References
ALESSANDRI, G. (1906) : Studi monografici sui
Cirripedi fossili d'Italia. Palaeontogr.
Italica, vol. 12, p. 207-324.
- - (1907a) : Osservazioni sopra a1cum Cir·ripedi fossili della Francia. Soc. Italiana
Sci. Nat. Milano, Atti, vol. 45, p. 251291.
- - (1907b) : Obsevations sur les Cirri pedes
fossiles de la France. Feuille Jeunes
Natural. Paris, vol. 37, p. 169-176, 193197.
AOKI, N. anq BABA, K. (1972): Compilationof the stratigraphy and molluscan fossil
assemblage of the Pleistocene ShimosaGroup, eastern part of Kwanto Plain,central Japan. Jour. Ceol. Soc. Japan,
vol. 79, p. 453-464. [in Japanese with Eng.
lish abstract].
ARAKAWA, K. Y. (1973): Handbook to prevention and extermination of foulingorganisms attached to cultured oysters.
Rept. Hiroshima Fish. Exper. Stat., 28 p.
[in Japanese].
BARNES, H. and BARNES, M. (1958) : Further
observations on self-fertilization in Chtha·
malus sp. Ecol., vol. 39, p. 550.
- - and CRISP, D. J. (1956): Evidence of selffertilization in certain species of barna-

193

cles. Jour. Mar. BioI. Assoc. U.K., vo!.
35, p. 631-639.
BROCH, H. (1922): Papers from Dr. Th. MORTENSEN'S Pacific expedition 1914-16. X.
Studies on Pacific Cirri peds. Vedensk.
Meddel. Dansk Naturhist., Foren. KifJbenhavn, vol. 73, p. 215-258.
- - (1924): Cirripedia Thoracica von Norwegen und dem norwegischen Nordmeere_
Eine systematische und biologischtiergeographische Studies. Vidensk. Kristiania,
Mat.-Naturw. Kl., Skri/t., vol. 2, p. 1131.
- - (1927): Report on the Crustacea Cirridedia. Trans. Zool. Soc., pt. 2, p. 133138.
- - (1931): Paper from Dr. Th. MORTEXSEN'S Pacific Expedition 1914-16. LVIIndomalayan Cirripedia. Vidensk. Meddel.
Naturhist., Foren., vol. 93, p. 1-146. .
CIIINZEI, K. (1966) : Younger Tertiary geologyof the Mabechi river valley, northeast
Honshu, Japan. Jour. Fac. Sci., Univ_
Tokyo, sec. 2, vol. 16, p. 161-208.
CORNW ALL, 1. E. (1925): A review of the Cirripedia of the coast of British Columbia,
with glossary, and key to genera andspecies. Contrib. Canad. BioI. Fish., n. s.,
vol. 2, p. 469-502.
- - (1951): The barnacles of California(Cirripedia). Wasmann Jour. BioI., vo!.
9, p. 311-346.
- - (1955) : Canadian Pacific Fauna. 10. Arthropoda. 10e. Cirripedia. Fish. Reser.
Board Canada, p. 1-49.
CRISP, D. J. (1954) : The breeding of Balanus
porcatus (DA COSTA) in the Irish sea. Jour.
Mar. Bioi. Assoc. U.K., vol. 33, p. 473496.
DARWIN, C. (1851a): A monograph on the
subclass Cirripedia, with figures of all
the species. The Lepadidae; or pedunculated Cirripeds. Ray Soc. Publ. (London), 400 p.
- - (1851b): A monograph on the fossil
Lepadidae, or pedunculated Cirri peds ofGreat Britain. Palaeontogr. Soc., vol. 6,
p. 1-88.
- - (1854a): A monograph on the. subclass
Cirripedla, with figures of all the species.

194

Toshiyuki YAMAGUCHIRay Sec. Publ. (L.ond.on), 684 p.
- - (1854b): A m.on.ograph .of the f.ossil
Balanidae and Verrucidae .of Great Britain. Palaeontogr. Soc., vel. 8, p. 1-44.
DAVADlE, C. (1963): Systematique et Structure des Balanus f.ossiles d'Eur.ope etd'Afrique. Edit. Centre Nat!. de la Reserch$ci., p. 1-146.
GREGG, J. H. (1948): Replicati.on .of substrata
detail by barnacles and s.ome .other marine .organisms. Bioi. Bull., vel. 94, p.
161-168.
GRUVEL, J. A. (1905): M.on.ographie des Cirrhipedes .ou Thec.ostraces. Mass.on et Cie(Paris), 472 p.
HANZAWA, S., HATAI, K., !WAI, J., KITAMURA, N. and SHIBATA, T. (1953): The
ge.ol.ogy .of Sendai and its envir.ons. Sci.
Rept., Tohoku Univ., ser. 2, vel. 25, p. 150.
HARDI:-.IG, J. P. (1962): DARWIN'S type specimens .of varieties .of Bal.anus amphitrite.
Bull. British Mus. (Nat. Hist.), Zool. ser.,vel. 9, p. 273-296.
HATAI, K.M. (1938): A review .of the f.ossil
Cirripedia and shark's teeth fr.om the regi.on .of n.ortheast H.onshu, Japan. Bull.
Biogeogr. Soc. Japan, vel. 8, p. 95-102.
HATAI, K., MASUDA, K. and SUZUKI, K.
(1961) : A n.ote .on the Pli.ocene megaf.ossil
fauna. fr.om the Shim.okita Peninsula,
A.om.ori Prefecture, n.ortheast H.onshu,
Japan. Saito Ho-on Kai Mus., Res. Bull.,

n.o. 30, p. 18-38.
HAYASAKA, S. (1961): The ge.ol.ogy and pale.ont.ol.ogy .of the Atsumi Peninsula, Aichi
Prefecture, Japan. Sci. Rept., Tohoku
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HENRY, D. P. (1940): The Cirripedia .of Puget
S.ound with a key t.o the species. Univ.
Washington Publ. Oceanogr., vel. 4, p. 148.
- - (1942) : Studies .on the sessile Cirripedia.of the Pacific c.oast .of N.orth America.
Univ. Washington Publ. Oceanogr., vel. 4,
p. 95-134.
- - (1973) : Descripti.ons .of f.our new species.of the Balanus amphitrite-c.omplex (Cirripedia, Th.oracica). Bull. Mar. Sci., vel.
23, p. 964-1001.
- - and McLAUGHLIN, P.A. (1967): A revisi.on .of the subgenus Solidobalanus HOEK
(Cirripedia Th.oracica) including a descripti.on .of a new species with c.omplemental males. Crustaceana, vel. 12, p.
43-58.
- - and - - (1975): The barnacles .of the
Balanus amphitrite c.omplex (Cirripedia,
Th.oracica). Zool. Verhandl., n.o. 141,
254 p.
HIRO, F. (1932a): Rep.ort .on the Japanese
species .of the genus Calantica (Cirripedia).
Annot. Zool. Japan, vel. 13, p. 467-482.
- - (1932b) : Rep.ort .of the bi.ol.ogical survey.of Mutsu Bay. 25. Cirripedia. Sci. Rept.,
Tohoku Univ., ser. 4, vel. 7, p. 545-552.

Explanation of Plate 24
Figs. ·1-3. Balanus rostratus HOEK .on Chlamys nipponensis fr.om the Pleist.ocene Semata F.ormati.on (L.oc. 20a). Aut.ochth.on. la-b. B. rostratus (UMUT-CA8289) .on left and rightvalves .of Chlamys, x 0.6., 2a-b. B. rostratus (UMUT-CA8290) .on left and right valves .ofChlamys, x 0.6., 3a-b. B. rostratus (UMUT-CA8291) .on left and right valves .of Chlamys,
xO.6.
Figs. 4-6. Balanus trigonus DARWIN .on living Chlamys nipponensis fr.om Kanita, Mutsu Bay(L.oc. C2). 4a-b. B. trigonus (UMUT-RA8292) .on left and right valves .of Chlamys, x 0.6.,
5a-b. B. trigonus (UMUT-RA8293) .on left and right valves .of Chlamys, x 0.6., 6:J-b. B.
trigonus (UMUT-RA8294) .on left and right valves .of Chlamys, x 0.6.
Fig. 7. Balanus rostratus HOEK fr.om Mutsu Bay (L.oc. Cl). living. Ph.ot.o by s.oft x-ray. 7a-g.
r.ostrum, x 1.5.
Fig. 8. M.ode.of .occurrence .of Balanus rostratus HOEK .on aut.ochth.on.ous Chlayms nipponellsis
fr.om the Pleist.ocene Semata F.ormati.on (L.oc. 20a).
Fig. 9. M.ode.of life .of living Chlamys Ilipponensis .on gravel at Asamushi, Mutsu Bay.

YAM AGUCHI :Japanese Balanoidea

Plate 24

682.

Taxonomic Studies on Japanese Balanoidea- - (1933): Report on the Cirripedia collet·
ed by the surveying ships of the Imperial
Fisheries Experimental Station on thecontinental shelf bordering Japan. Rec.
Oceanogr. Works Japan, vol. 5, p. 11-84.
- - (1935): The faua of Akkeshi Bay. II.
Cirripedia (Studies on the cirripedian
fauna of Japan. I). Jour. Pac. Sci., Hokkaido Imp. Univ., ser. 4, vol. 4, p. 213229.
- - (1937) : Studies on the cirripedian faunaof Japan. II. Cirripeds found in thevicinity of the Seto Marine Biological
Laboratory. Mem. Coil. Sci., Kyoto Imp.
Univ., ser. B, vol. 12, p. 385-478.
- - (1938) : On the Japanese forms of Balanus amphitrite DARWIN. Zool. Mag. (Japan), vol. 50, p. 299-313. [in Japanesewith English abstract]_.- (1939a) : Studies on the cirripedian faunaof Japan. IV. Cirripeds of Formosa (Taiwan), with some geological and ecological remarks on the littoral forms. Mem.
Coil. Sci., Kyoto Imp. Univ., ser. B, vol.
15, p. 245-284.
- - (1939b) : Studies on the cirripedian faunaof Japan. V. Cirripeds of· the northernpart of Honshu. Sci. Rept., Tohoku Imp.
Univ., ser. 4, vol. U, p. 201-218.
HOEK, P. P. C. (1883) : Report on the Cirripedia
collected by H. M. S. Challenger during
the years 1873-76. Results Explor. Voyage
H.M.S. Challenger, Zoo!., vol. 8, p. 1-169.
- - (191?): The Cirripedia of the SibogaExpedition. B. The Cirripedia sessilia.
Siboge-Expeditie, 31b, p. 129-275.
HORIKOSIII, M. (1962) : Distribution of benthic
organisms and their remains at the entrance of Tokyo Bay (Co-operative survey
at the entrance of Tokyo Bay, in 1959part 7). Jour. Oceanogr. Soc. Japan, 20th
Anniv. vol., p. 146-154. [in Japanese withEnglish abstract]
HUDINAGA, M. and KASAHARA, H. (1942): 9n
the rearing and metamorphosis of Balanus
amphitrite hawaiiensis BROCH. Zool. Mag.
(Japan), vol. 54, p. 108-118. [in Japanesewith English abstract]
HUZIOKA, K. (1959): Geological map, Togaand Funakawa (scale 1: 50,000). Geol.

195

Surv. Japan, 61 p. [in Japanese with English abstract]
IGA, T. (1973): Barnacles of Lake Shinji-ko,
Naka-umi and Jinzai-ko. San-in Bunka
Kenkyu Kiyo, no. 13, p. 59-69. [in Japanese]
IKEYA, N. and UEMATSU, K. (1968): The
stratigraphic relation between the Kuromatsunai andSetanaFormations.-Geologyof the Setana-Imagane-Yakumo area in
the Oshima Peninsula, southwestern
Hokkaido-. Jour. Ceol. Soc. Japan, vol.
74, p. 21-36. [in Japanese with English
abstract]
ISHIDA, S. and Y ASLJGI, R. (1973) : Free-swimming stage of Balanus amphitrite albicostatus. Plant Animal (Japan), vol. 5,
p. 1659-1666. [in Japanese]
ITOIGAWA, ]. (1960): Paleoecological studiesof the Miocene Mizunami Group, central
Japan. Jour. Earth Sci., Nagoya Univ.,
vol. 8, p. 246-300.
IWAI, T. (1965) : The geological and paleontological studies in the marginal area of
the Tsugaru basin, Aomori Prefecture,
Japan. Bull. Educ. Pac., Hirosaki Univ.,
no. 15, p. 1-68.
- - and SIOBARA, T. (1969): Pleistocene
mollusca from Kamikita-gun, Aomori
Prefecture, Japan. Bull. Educ. Pac., Hirosaki Univ., no. 20B, p. 1-7.
IWASAKI, Y. (1970): The Shiobara-type molluscan fauna. An ecological analysis offossil molluscs. Jour. Pac. Sci., Univ.
Tokyo, ser. 2, vol. 17, p. 351-444.
KASENO, Y. and MATSUURA, N. (1965) : Pliocene shells from the Omma formation
around Kanazawa City, Japan. Sci. Rept.,
Kanazawa Univ., vol. 10, p. 27-62.
KATAYAMA, M. (1941): Study on the geological structure of the Japanese oil-fields;on the Tertiary formations between Honjo
and Yokote. Jour. Japanese Petrol. Technol., vol. 9, p. 93-105. [in Japanese]
KATO, M., HAYASAKA, K .. and MATSUDA, T.
(1960a): Ecological studies on the morphological v~ri·a'fion of a sessiJ'e barnacles, Chtllamalus challengeri. 1. Changesof the external appearance introduced by
the population density. Bull. Mal·. Bioi.

196

Toshiyuki YAMAGUCHI

Stat. Asamushi, vol. 10, p. 1-7.
- - , - - and - - (1960b) : Ecological studies
on the morphological variation of a sessile barnacles, Chthamalus challengeri.
III. Variation of the shell shape and of
the inner anatomical feature introducedby the population density. Bull. Mar.
Bioi. Stat. Asamushi, vol. 10, p. 19-25.
KAWADA, K. (1953): Geological studies on
the Yamizo, Torinoko and Toriashi mountain blocks and their neighbourhood in
the northeastern Kwanto district. Sci.
Rept., Tokyo Bunrika Daigaku, ser. C,
vol. 2, p. 217-307.
KITAZATO, H. (1975): Geology and geochronology of the younger Cenozoic of Oga
Peninsula. Tohoku Univ., Inst. Geo!. Pal.,
Contr., no. 75, p. 17-49. [in Japanese withEnglish abstract]
KOLOSV ARY, G. (1943) : Cirripedia von Thora-

cica in Sammulung des ungarischen National Museums. Hist. Nat. Mus. Nat.
Hungarici, Ann., Zool., vol. 36, p. 67-120.
- - (1948) : New balanids from the middleMiocene of Varpalota in Hungary. Foldt.
Kozl., vol. 78, p. 101-112.
- - (1961): Einige interessante BalanusFunde aus der Burdigalienstufe von
Turkmenistan. Acta Bioi. Szeged, n. s.,
vol. 7, p. 99-102.
- - (1962a) : New data to the Balanida-faunaof the Burdigalien of Karakum. Acta
BioI. Szeged, n. s., vol. 8, p. 199-202.
- - (1962b) : New micro-balanids from Tongatabu. Acta Bioi. Szeged, n. s., vol. 8,
p. 193-197.
KOJIMA, N. (1958): Geological study of theKioroshi district, Chiba Prefecture, Japan.-The studies on the Narita group(1) -. Jour. Geol. Soc. Japan, vol. 64,

Explanation of Plate 25
Fig. 1. Balanus crenatus BRUGUIERE (UMUT-RA8330) from Paramushir Is., Kurile Isis. Coli.
Mr. K. KOBA. la-b. whole wall, x 1.5., Ie-d. exterior and interior views of right scutum,
x 5., Ie-f. exterior and interior views of tergum, x 5., Ig-h. exterior and interior viewsof rostrum, x 2., Ii-j. exterior and interior views of right lateral, x 2., lk-l. exterior andinterior views of right carinolateral, x 2., 1m-n. exterior and interior views of carina,
x2.
Figs. 2-7. Balanus crena/us BRCGCIERE from the Pliocene Daishaka Formation (Loc. 5b). 2ab. exterior and interior views of right scutum (UMUT-CA8311), x 5., 3a-b. exterior andinterior views of right tergum (UMUT-CA8332), x 5., 4a-b. exterior and interior viewsof rostrum (UMUT-CA8333), x 2., 5a-b. exterior and interior views of right lateral(UMUT-CA8334), x2., 6a-b. exterior and interior views of right carinolateral (UMUTCA8335), x 2., 7a-b. exterior and interior views of carina (UMUT-CA8336), x 2.
Figs. 8-13. Balanus crenatus BRCGUlERE from the late Miocene Oidawara Formation (Loc.
41e). 8a-b. exterior and interior views of left scutum (UMUT-CA8337), x 5., 9a-b. exterior and interior views of left tergum (UMUT-CA8338), x 5., lOa-b. exterior and interior views of rostrum (UMUT-CA8339), x 2., lla-b. exterior and interior views ofright lateral (UMUT-CA8340), x 2., 12a-b. exterior and interior views of right carinolateral (UMUT-CA8341), x 2., 13a-b. exterior and interior views of carina (UMUTCA8342), x 2.
Figs. 14-18. Balanlls crena/us BRCGCIERE from the early to middle Miocene Moniwa Formation (Loc. 12b). 14a-b. exterior and interior views of left scutum (UMUT -CA8343),
x 4., 15a-b. exterior and interior views of right tergum (UMUT -CA8344), x 4., 16a-b.
exterior and interior views of right lateral (UMUT-CA8345), x 2., 17a-b. exterior andinterior views of left carinolateral (UMUT-CA8346), x 2., 18a-b. exterior and interior
views of carina (UMUT -CA834 7), x 2.
Fig. 19. Balanus crenatus BRUGClERE (UMUT-CA8348) from the late Miocene Shimokurosawa
Formation (Loc. 11), x 1.5.

YAM AGUCHI:

Japanese Balanoidea

Plate 25

682.

Taxonomic Studies on Japanese Balanoidea

p. 165-171. [in Japanese with English
abstract]- - (1966) : Geological structure of the southeast coast of Tokyo Bay, Japan.-Studies
on the Narita group (6) -. jour. Ceol.
Soc. jaban, vol. 72, p. 205-212. [in Japanese with English abstract]
KROGER, P. (1911a) : Beitrage zur Cirrpedienfauna Ostasiens. Adh. d. II. Kl. d. K.
Ak. d. Wiss. II. Suppl., Bd. 6, p. 1-72.
--. (1911b) : Zur Cirripedienfauna Ostasiens.
Zool. Anzeig., vol. 38, p. 459-464.
MATSUSI-IlMA, Y. (197l): HC age and molluscan assemblage of the Of una shell bed
in Yokohama City. Bull. Kanagawa Pref.
Mus., Nat. Sci., vol. 1, p. 61-72. [in Japanese with English abstract]. - (1972): Molluscan assemblages of theinner part of Paleo-Ofuna Bay, central
Kanagawa Prefecture. Bull. Kanagawa
Pref. Mus., Nat. Sci., vol. 1, p. 31-43. [inJapanese with English abstract]--. (1973): Preliminary report on the molluscan assemblages from the alluvial deposits in Yokohama City. Bull. Kanagawa
Pref. Mus., Nat. Sci., no. 6, p. 7-19. [inJapanese with English abstract]- - (1974) : Geology of Kanagawa Prefecture.
no. 1. Res. Rept., Kanagawa Pref. Mus.,
Nat. Hist., no. 5, 40 p. [in Japanese]
MA\\-ATARI, S. (1967): Biological studies on
fouling in Japanese Horbors I. General
concept of researches. Misc. Rept. Res.
Inst. Nat. Res., no. 69, p. 87-114. [inJapanese]- - and KOBAYASHI, S. (1954a): Seasonalsettlement of animal fouling organisms
in Ago Bay, middle part of Japan I. Misc.
Rept. Rec. Inst. Nat. Res., no. 34, p. 3747.
- - and - - (1954b): Seasonal settlementof animal fouling organisms in Ago Bay,
middle part of Japan II. Misc. Rept. Res.
Inst. Nat. Res., no. 36, p. 1-8.
MAYR, E. (1963): Animal species and evolution.
Belknap Press, Harvard Univ.
Press, Cambridge, Mass., 797 p.
McLAUGI-ILI:-.1, P.A. and HENRY, D.P. (1972):
Comparative morphology of complemental
males in four species of Balanus (Cir-197

ripedia Thoracia). Crustaceana, vol. 22,
p. 13-30.
MIZUNO, T., HARADA, E. and MIZUNO, N.
(1964) : Seasonal fluctuation and vertical
distribution of larval of fouling organisms around Sakai Harbor in Osaka Bay.
Mem. Osaka Cakugei Univ., B, no. 13,
p. 127-135.
NAKAGAWA, H. (1960): On the stratigraphyof Jizodo and Yabu formations. jour.
Ceol. Soc. japan, vol. 66, p. 305-310. [inJapanese with English abstract]
NEWMAN, W.A. (1967): A new genus of
Chthamalidae (Cirripedia, Balanomorpha)
from the Red Sea and Indian Ocean. jour.
Zool. (London), vol. 153, p. 423-435.
- - , ZULLO, V. A. and WITHERS, T. H. (1969) :Cirripedia. (in Treatise on Invertebrate
Paleontology, part R, Arthropoda 4, vol.
1, ed. MOORE, R. C.). Geol. Surv. Amer.
and Univ. Kansas, p. 206-295.
- - and Ross, A. (1976): Revision of thebalanomorph barnacles; including a catalog of the species. San Diego Soc., Nat.
Hist., Mem., vol. 9, p. 105.
NILSSO:-i-CANTELL, C. A. (1921): CirripedienStudien zur Kenntnis der Biologie, Anatomie und Systematik dieser Gruppe.
Zool. Bidrag, Uppsala, Bd. 7, p. 75-390.
- - (1932) : Cirripedien aus Japan gesammelt
von Dr. SMITH, Dr. HABERER und Dr.
HILGENDORF', in dem Berliner Museum
aufbewahrt. Ark. Zool., vol. 24A, p. 130.
- - (1934): Indo-Malayan cirripeds in the
Raffles Museum, Singapore. Bull. RajJIes
Mus., no. 9, p. 42-73.
- - (1938) : Cirri pedes from the Indian Ocean
in the collection of the Indian Museum,Calcutta. Mem. Indian Mus., vol. 13, pt.
1, p. 1-81.
NOMURA, S. (1932) : Mollusca from the raisedbeach deposits of the Kwanto region.
Sci. Rept., Tohoku Imp. Univ., (Ceol.) ,vol. 15, p. 65-144.
OHARA, S. (1966): Stratigraphy and geologic
structures of the Tertiary deposits in the
Uryu coal-field, Hokkaido, Japan. jOllr.
Call. Art. Sci., Chiba Univ., vol. 4, p. 617630.

~98

Toshiy~~ki YAMAGUCHI(1969): The type Semata formation.
jour. Coil. Art. Sci., Chiba Univ., Nat.
Sci. ser., vol. 5, p. 303-318. [in Japanesewith English abstract]- - and TAIRA, K. (1974): Molluscan remains from the Taito-zaki formation.
jour .. Coil. Arts Sci., Chiba Univ., B-7,
p. 43-53.
OLIVEIRA, L. P. H. de (1941) : Contribui~ao aoconhencimento dos crustaceos do Rio deJaneiro. Sub-ordem "Balanomorpha" (Cirripedia; Thoracica). Mem. Inst. Oswaldo
Cruz, vol. 36, p. 1-31.
ONODERA, S. (1957): A new occurrence ofDesmostylus from Ichinoseki City, Iwate
Prefecture, with reference to the geologyof the locality. jour. Geol. Soc. japan,
vol. 63, p. 238-253. [in Japanese with English abstract]
OTUKA, Y. (1930): On the stratigraphy of thenorthern part of the Miura Peninsula,with special reference to the variationof the shore-line in the latest geologicalage. jour. Geol. Soc. Tokyo, vol. 37, p.
343-386. [in Japanese]- - (1935) : Stratigraphy of the northeasternpart of the Oti Graben, Ichikawa Prefecture. jour. Geol. Soc. japan, vol. 42, p.
483-510. [in Japanese]- - (1936): On the Takase-gawa tuff beds
in Yuri district, Akita Prefecture. jour.

Geol. Soc. japa1l, vol. 43, p. 697-706. [inJapanese]
OZAKI, H. (1958): Stratigraphical and paleontological studies on the Neogene andPleistocene formations of the Tyosi District. Bull. Nat. Sci. Mus., vol. 4, no. 1(no. 42), p. 1-182.
PILSBRY, H.A. (1907): Cirripedia from the
Pacific Coast of North America. Bull.
U.S. Bur. Fish., vol. 26, p. 191-204.
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Sea. Bull. U.S. Bur. Fish., vol. 29, p. 5984.
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contained in the collections of the U.S.
National Museum: including a monographof the American species. Bull. U.S. Nat!.
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- - (1918): Contributions to the geology andpaleontology of the Canel Zone, Panama,and geologically related areas in central
America and the West Indies. Cirripedia
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- - (1921): Barnacles of the San Juan Islands, Washington. Proc. U.S. Nat!. Mus.,vol. 59, p. 111-11;5.
--- (1924): Miocene and Pleistocene Cirripedia from Haiti. Proc. U.S. Nat!. Mus.,vol. 65, p. 1-3.
- - (1927) : Littoral barnacles of the Hawai-

Explanation of Plate 26
Fig. 1. Megabalanus volcano (PILSBRY) (UMUT-CA8349) from the Pleistocene Nakoshi Sandstone (Loc. 44). la-b. crowded individuals, x 0.8.
.
Figs. 2-10. Megabalanus volcano (PILSBRY) from the Pleistocene Nakoshi Sandstone (Loc. 44).
2a-b. exterior and interior views of right scutum (UMUT-CA8350), x3., 3a-b. exterior
and interior views of left scutum (UMUT-CA8351), x 5., 4a-b. exterior and interior viewsof left scutum (UMUT-CA8352), x 5., 5a-b. exterior and interior views of right scutum(UMUT-CA8353), x5., 6a-b. exterior and interior views of left scutum (UMUT-CA8354),
x 5., 7a-b. exterior and interior views of right tergum (UMUT-CA8355), x 5., 8a-b. exterior and interior views of left tergum (UMUT-CA8356), x5., 9a-b. exterior and interior
views of left tergum (UMUT-CA8357), x 5., lOa-b. exterior and interior views of righttergum (UMUT -CA8358), x 5.
Figs. 11-12. Megabalanus rosa (PILSBRY) from the Pleistocene Nakoshi Sandstone (Loc. 44).
11a-b. exterior and interior views of left scutum (UMUT -CA8359), x 5., 12a-b. exterior
and. in~erior views of left tergum (UMUT -CA8360), x 5.
Fig. 13. Association of Megabalanus volcano (PILSBRY) and M. rosa (PILSBRY) from the Pleistocene Nakoshi Sandstone (Loc. 44). (UMUT -CA8361). x 1. v: volcano, r: rosa.

YAMAGUCHI:

Japanese Balanoidea

Plate 26

682.

Taxonomic Studies on Japanese Balanoidea

ian Islands and Japan. Proc. Acad. Nat.
Sci. Phi/ad., vol. 79, p. 305-317.
- - (1930): Cirripedia (Balanus) from the
Miocene of New Jersey. Proc. Acad. Nat.
Sci. Philad., vol. 82, p. 429-433.
ROSELL, N. C. (1973) : Some thoracic barnacles[Crustacea: CirripediaJ of Manila Bay.
Kalikansan Philipp. Jour. Bioi., vol. 2,
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SEGUENZA, G. (1973-1876): Richerche paleontologiche intorno ai cirripedi terziariidella Provincia di Messina. Con append ice
intorno ai cirripedi viventi nel Mediterrano, e sui fossili terziarii dell'Italia
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[in JapaneseJ- - and MASUJINIA, A. (1969) : Quantitative
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SOUTHWARD, A. J. (1975) : Intertidal and shallow water Cirripedia of the Caribbean.
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STEPHENSEN, K. H. (1938): Cirripedia (incl.
Rhizocephala). (in Zoology of Iceland,
vol. 3, pts. 30-31). Copenhagen, Levin andMunksgaard, p. 1-11.
T ARASOV, N. E. (1936): On the Arctic faunaof Cirripedia Thoracica II. Arctic Ins/.
Leningrad, Trans., vol. 33, p. 45-49. [inRussian]- - (1937): Contribution to the fauna of
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III. Arctic Inst., Leningrad, Traits., vol.
50, p. 35-59. [in RussianJ- - and ZEVINA, G. B. (1957): Cirridedia

199

Thoracica of the sea of the U.S.S.R. Zool.
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200

Toshiyuki YAMAGUCHI

Bioi. Lab., vol. 17, p. 339-372.
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WEL TNER, W. (1897) : Verzeichnis der bisher
beschriebenen recenten Cirripedienarten.
Archiv. fur Naturg., vol. 1, p. 227-280.
WITHERS, T. H. (1923): Die Cirripedian der
Kreide Rugerns. Univ. Greifswald, Geol.
Paleont. Inst., Abhandl., vol. 3, p. 1-54.

Explanation of Plate 27
Fig. 1. Solido balanus (Hesperibalallus) hesperius (PILSBRY) (UMUT -RA8295) from Funka Bay(Loc. B3). la-b. exterior and interior views of right scutum, x 4., 2a-b. exterior andinterior views of right tergum, x 4., Ie-f. exterior and interior views of rostrum, x 1.5.,
19-h. exterior and interior views of right lateral, x 1.5., 1i-j. exterior and interior viewsof carina, x 1.5.
Fig. 2. Solidobalanus (Hesperibalallus) hesperius (PILSBRY) (UMUT -RA8296) from Funka Bay(Loc. B3). 2a-b. exterior and interior views of right scutum, x 8., 2c-d. exterior andinterior views of right tergum, x 8., 2e-f. exterior and interior views of rostrum, x 3.,
2g-h. exterior and interior views of right lateral, x 3., 2i-j. exterior and interior viewsof carina, x 3.
Fig. 3. Solidobalallus (Hesperibalallus) hesperius (PILSBRY) (UMUT -RA8297) from Funka Bay(Loc. B3). 3a-b. exterior and interior views of right scutum, x 5., 3c-d. exterior andinterior views of right tergum, x 5., 3e-f. exterior and interior views of rostrum, x 2.,
3g-h. exterior and interior views of right lateral, x2., 3i-j. exterior and interior viewsof carina, x 2.
Fig. 4. Solidobalanus (Hesperibalanus) hesperius (PILSBRY) (UMUT -RA8298) on Macoma tokyoensis from Funka Bay (Loc. B2), x 0.75 (smooth and ribbed forms).
Fig. 5. Solido balanus (HesperibalallUs) hesperius (PILSBRY) (UMUT -RA8299) on TUiTitella fortilirata from Funka Bay (Loc. B3), x 0.75.
Fig. 6. Solidobalanus (Hesperibalanus) hesperius (PILSBRY) (UMUT -CA8300) from the Pleistocene Nishiyatsu Formation (Loc. 26), x 2.
Figs. 7-9. Solidobalanus (Hesperibalanus) hesperius (PILSBRY) from the Pleistocene Narita
Formation (Loc. 24i). 7a-b. exterior and interior views of left scutum (UMUT -CA8301) ,x 7., 8a-b. exterior and interior views of left tergum (UMUT-CA8302), x 7., 9a-b. exterior
and interior views of rostrum (UMUT-CA8303), x 3.
Figs. 10-11. Solidobalanus (Hesperibalanus) liesperius (PILSBRY) from the Pleistocene Shibikawa
Formation (Loc. 8c). lOa-b. exterior and interior views of rostrum (UMUT -CA8304) ,x 3., 11a-b. exterior and interior views of left lateral (UMUT-CA8305), x 3.
Figs. 12-14. Solidabalanus (Hesperibalallus) hesperius (PILSBRY) from the Pliocene Horoshintachibetsu Formation (Loc. 1). 12. whole wall (UMUT-CA8306), x 2.5., 13. (UMUTCA8307) ; 13a. whole wall, x 2.5, 13b-c. exterior and interior views of left scutum, x 7,
13d-e. exterior and interior views of right tergum, x7., 14a-b. exterior .and interiorview s of rostrum (UMUT -CA8308), x 2.
Figs. 15-16. Solidobalanus (Hesperibalanus) hesperius (PILSBRY) from the early to middle Miocene Sugota Tuffaceous Silty Sand (Loc. 9b). 15a-b. exterior and interior views ofrostrum (UMUT-CA8309), x 3., 16a-b. exterior and interior views of left lateral (UMUTCA8310) , x 3.
Fig. 17. Solidobalanus (Hesperibalanus) liesperius (PILSBRY) (UMUT-CA8311) from the late
Miocene Maisawa sandstone of the Suenomatsuyama Formation (Loc. 7k), x 2.
Fig. 18. Solidobalanus (Hesperibalallus) hesperius (PILSBRY) (UMUT-CA8312) on Placopecten
setanensis from the late Miocene Yakumo Formation (Loc. 2d), x 0.6.

YAMAGUCHI:

Japanese Balanoidea

Plate 27

682.

Taxonomic Studies on Japanese Balanoidea- - (192-1): The fossil cirri pedes of New
Zealand. New Zealand Ceol. Surv., Paleont., Bull., no. 10, p. 1-47.
YABE, H. and SUGIYAMA, T. (1931) : A studyof recent and semi-fossil corals of Japan.
Sci. Rept., Tohoku Imp. Univ., (Ceol.) ,vol. 14, p. 119-133.
YAMAGUCHI, T. (1970): The Neogene of thenortl:eastern part of the Shimokita Peninsula, northeast Honshu, Japan.-On thestratigraphic relation of the Tomari, theGamanosawa and the " Sunagomata "formations-. jour. Ceol. Soc. japan,
vol. 76, p. 185-197. [in Japanese withEnglish abstract]- - (1971) : Fossil barnacles from the Pleistocene Miyata formation. Sci. Rept., Yokosuka City Mus., no. 18, p. 122-129.
- - (1972) : Fossil barnacles in Japan. Master
Thesis, Univ. Tokyo (MS).
- - (1973): On Megabalanus (Cirripedia,
Thoracica) of Japan. Publ. Seto Mar.
Bioi. Lab., vol. 21, p. 115-140.
- - (1974a): Taxanomic studies on somefossil and Recent Balanus in Japan. Dissertation Thesis. Ceo!. Inst., Univ. Tokyo(MS).
- - (1974b): On some fossil Balanus from201

the Miocene Mizunami group. Bull. Mizunami Fossil Mus., no. 1, p. 215-220. [inJapanese]
YA:VIA:VIURA, Y. (1972): Ecological studies on
marine fouling communities in pearl culture ground. II. Seasonal changes in theconstitution of marine fouling communities at various depths in Ago Bay.
Bull. Nat!. Pearl Res. Lab., vol. 16, p.
2038-2051. [in Japanese]- - , KUWATAl'\I, Y. and NISHII, T. (1969) :Ecological studies of Marine foulingcommunities in pearl culture ground. I.
Seasonal changes in the constitution ofmarine fouling communities at a pearl
cultivating depth in Ago Bay. Bull. Nat!.
Pearl Res. Lab., vol. 14, p. 1836-1861. [inJapanese]
YOKOY A:VIA, M. (1911): Climatic changes inJapan since the Pliocene epoch. jour.
Coli. Sci., Tokyo Imp. Univ., vol. 32, p.
1-16.
ZEVI;-':A, G. E. and TARASOV, N. I. (1963) : The
cirripedian fauna (Cirripedia, Thoracica)along the continental coast of southeast
Asia. Trudy Inst. Oceanol., vol. 70, p. 76100. [in Russian]Trans. Proc. Palaeont. Soc. Japan, N.S., No. 108, pp. 202-221, pis. 28-30, November 15,)977

683. SOME LATE TRIASSIC ElV ALVIA AND GASTROPODAFROM THE DOMEYKO RANGE OF NORTH CHILE*ITARU HAY AMI
University Museum, University of TokyoSHIRO MAEDA
Department of Earth Sciences, Faculty of Science, Chiba University

andCARLOS RUIZ FULLER
Instituto de Investigacions Geologicas, Antofagasta

T !J -~~fijl ~. -" 1 :1 ilJttBQ)=::I:,fi[!f3tM=:tlcJU·o.J:. V'~'@'1Ji: 1970fFt::. T~*~ C 'T !J -ill!.
1.l1EP.JTc7)~jrC1Tt.ch;/'tt.: ~. -" 1 :1 ilJ:lfuQ):lfu.ii~~~fpJ~1tQ)~~£~~~t Lt.:iti6,*, 7;"~ 77 jJ:A!J Q)f.<J 180 km ~*Q)Ii'J;.:OER;'Eh~' i?~i?;/'tt.:{t:Ollf.pt::. Limidae Q) 2 ~f.!I!~~
tr=:tlcft 13 f.!I! C ~'@'1Ji 3 f.!I!~~J.jIj, ii~ Lt.:Q) -C1ic~-f" Q o :::.. Q) 5 b Antiquilima ataca.
mensis It 9=>- :7;f,(!t::.i!!:.ff.i¥Jt::.~* Lt.: Ctenostreon Q)~1f{~ tvlll:L fpJJMQ)~~~~*T Q
J:t::.JI!i!,*;t.,Qf.!I!-c;t.,Qo :::"Q){t:Oi!f.lt.-{Jv-.pfijlQ)-l!o' r· . . ~:A:1:lfu~i/'i?~i?;/'t-Ct'Q=:
1i:,fi[!f3tM (/ !J 7;"-) Q)iIMj;;lIW~f.ft::'~lmTQm~$ < ~;z,.., .gj:;'6Q)'I1:.t1Jit£tTQQ)-C, .-{Jv-t::'Jh <it;ffi"T Q Pucara JWffc7)~~i/;:::" Q)ilJ:lfut::.~ L -Ct, Q:::" C a-~ L -Ct':6 o =:&'&Q)
j'~Ji,!Ht-~'c.J:. <*-t¥ ~;/'t -Ct, Qi/;, i/:A, i?t, 7:7 v:O-C-C g -Ct ,t.: c,'6th;/'tQ !19mlt Uf L
Ifm:k L -Ct, -C, ~Q)mii!I~~~~p'xQ)bi/;t't::..J:."? -C~J.jIji¥Jtdt:o{ti/;;z,..i?;/'tQo {t:Olt$~c tl§Ht1'Fm~~ft, ~~l~I!I-::..J:. "?-c~.~~*-cg Q~~i/;;t.,Qo Jt{tlt~J.jIji¥Jtdt:O{tQ)f3t-C~~ ;/'ttdf~Q) !19j'~Q)~JWi/' i? pgfijlt::.ri'ih -Cjfijrr Lt.: c~:t i?;/'t Q o:ilill./<.

~. mr83gg!l~ • C. R. FULLER

cions Geologicas (Santiago and Antofagasta) with the support of the Overseas
Scientific Research Funds, Ministry of
Education, Government of Japan. The
present article is a part of its outcome,particularly about the newly discovered
Triassic molluscan fossils from this area.
The Domeyko Range is located in thearid area of northern Chile (Provincesof Antofagasta and Copiapo), extending
meridionally, ranging from 3,800 m to
4,900 m in altitude. Between this range

Introduction
In October and November, 1970, a geological and paleontological reconnaissance survey on the Mesozoic terrainof the Domeyko Range of Chile wascarried out in cooperation between themembers of the Chiba University Palaeontological Expedition to the Andes andthe staff of the Instituto de Investiga-* Received April 22, 1977; read June 18,
1977, at Shizlloka.

202

683.

T1"iassic Molluscs from Chileand the still higher Andean Cordillera
on the east there are several salt lakes,
namely, Salar de Atacama, Salar de
Imilac and Salar de Punta Negra from
north to south.
The fossils described in the present
article were obtained at two localities(GCH 26 and GCH 85), both of which aresituated on the eastern slope of thisrange near Salar de Punta Negra, about
180 km southeast of Antofagasta (Textfigs. 1, 2). The stratigraphy and geologic
structure of the Mesozoic terrain in thisrange have not been sufficiently clarified,although, as shown in the geologic mapof Chile (published from the Instituto deInvestigacions Geologicas in 1968), it has
generally been regarded as jurassic andpartly as Cretaceous. Actually, un doubt-

400km

Text.fig. 1. Index map.
C: Cerro de Pasco area, D: Domeyko
Range (locality of the present material)

203

ed Lower jurassic fossils occur at sever··al localities on the western slope of thisrange. The occurrence of various fossils
at the present localities was first foundby Dr. CHONG, but they remain underscribed except for a species of Thecosmilia (hexacoral) in another paper related to the same expedition (MAEDA,YAMAGIWA, FULLER and CHONG. 1974).
The fossiliferous beds are greyish(brownish if weathered) impure limestones of 1-3 meters' thickness whichalternate with black shales. They contain abundant molluscan shells which
are mostly fragmentary but sometimes
well preserved. The strata near thetwo localities dip steeply to the east or
are nearly vertical, being strongly folded.
Because of the much flattened topography and insufficient exposure, it isdifficult to recognize the detailed stratigraphic sequence in this area, but thevarious molluscan fossils, as described
here, clearly indicate an upper Triassic
age instead of jurassic. Actually thisfauna has a marked resemblance to theUpper Triassic ones of northern andcentral Peru, which were described by
j AWORSKI (1922), KORNER (1937), Cox(1949) and HAAS (1953). Like the fossils
from the Pucara Group of Peru, themolluscan shells are considerably silicified, offering interesting material not
only for the taxonomic studies but also
for the consideration of fossilization andsilification.
Acknowledgements. - We express our
sincere thanks to Drs. Guillermo CHONG
D. and jose CORVALAN D. (Instituto deInvestigacions Geologicas, Santiago), Dr.
Giovanni CECIONI (Universidad de Chile),
Professor Takeshi CHISAKA (Chiba University), Professor Tatsuaki KIMURA(Tokyo Gakugei University), Dr. Takashi
HAMADA (University of Tokyo) and Mr.
Haruo . T AZUKE (Education Center of204

Itaru HAYAMI, Shiro MAEDA and C. R. FULLER

Text.fig. 2. Fossil localities in the Domeyko Range (GCH 26 and GCH 85)

Chiba Prefecture) for their close cooperation in collecting the material, to Dr.
Hideo KATSUKI (President of the Chiba
University), Professor Emeritus Teiichi
KOBAYASHI (Japan Academy) and thestaff of the Department of Earth Sciences, Chiba University for their generous support to this project, and also to
the members of the Mitsubishi Mining
Co. and Nissan Motors Co. and Mr.
Toshio KUROSAKI of Japan External
Trade Organization for their kind assistance in the actual field survey.
Thanks are also due to Professor Tetsuro
HANAI (University of Tokyo) and Pro-

fessor Kenji KONISHI (Kanazawa University) for their discussion about thefossilization of this material.
Presesvation and silicificationof fossils
Most of the fossils here studied wereobtained from the weathered surface of
impure limestone in situ at the two
localities. The calcium carbonate oforiginal shells have been commonly replaced by silica, and the fossils seem to
be more resistant to chemical and mechanical weathering than the matrix. The

683.

T1'iassic Molluscs from Chile

actual extent of silicification, however,is considerably variable among species
and individuals. Because selective fossilization of particular shell layers is
occasionally seen, careful observation
and adequate interpretation on the stateof preservation are required for thetaxonomic study on this material.
Chemical preparation of fossils by acidis really effective to take out the silicified shells from the limestone blocks.
If the calcareous cement is completely
dissolved, the rock becomes quite loose,looking as if it were Neogene or Quaternary sediment. The residue is wholly
composed of fine to medium-grained sandand numerous fragments of silicified
fossils, occupying 15 to 40 percent of theoriginal block in weight. It may be,
therefore, the best way for preparationof material to remove mechanically theloose residue after soaking the blocks
in diluted HCI for several days. For
large blocks we repeated the combination of the chemical and mechanicalprocedures for several times. The silicified shells are, however, quite fragile
and apt to be damaged, especially inthin-shelled species. In some bivalves
the silification is not completed or only
the outer layer is selectively preserved,
and, even if they had originally thicktests, this method is hardly applicable.
In such cases it is partically effective
to coat some part of the surface withparaffin before soaking the blocks inacid.
As the result of this preparatory work,
13 species of the Bivalvia and 2 speciesof the Gastropoda came to our knowledge as listed in Table 1. In the sametable the actual state of preservation isshown in contrast with the presumed
original shell structure and mineralogy
which are, though not directly evidenced,
generally inferable from the data by

205

TAYLOR, KENNEDY and HALL (1969, 1973)
and others.
Although the process and cause of theselective fossilization and silicification
still remain unsolved, we tentatively
summarize the result of our observation and interpretation in the following
items:
1) In most species of the Bivalvia theouter layer is better preserved than the
inner. The detailed ornaments andgrowth increments on the external surface are generally well preserved, whereas the hinge teeth and musculature areobservable only in a few species.
2) Regardless of the difference of shell
fabrics, fossils originally composed ofcalcite seem to be well preserved. The
outer layers of the shells of Modiolus
sp., Pinna sp., Pseudolimea chongi sp.
nov. and Antiquilima atacamensis sp. nov.
and also the almost whole shells ofPlicatula sp. and Gryphaea sp. are theseexamples. They are generally silicified,
but the silicification is often incompletein the interior of the foliated layers ofPlicatula sp. and Gryphaea sp. In every
specimen of Pinna sp. original simple
prismatic structure is well preserved,
notwithstanding that each prism has
been completely replaced by pseudomorphous silica (see Plate 28, Fig. Ib).
The preserved shells of Antiquilima
atakamensis are very thin (less than 0.4
mm in maximum thickness) and probably
produced by silicification after selective
fossilization (see Text-fig. 3).
3) The inner layer of crossed-lamellar
or nacreous aragonite in many of above
mentioned species was dissolved at allwithout any trace. The space originallyoccupied by the inner layer seems tohave been closed by compaction. This
state of preservation is apparently similar to that of EUropean Chalk which wasstudied by KENNEDY (1969), CARTER

Itar1,£ HAYAMI, Shiro MAEDA and C. R. FULLER

206

Table 1. Late Triassic Bivalvia and Gastropoda from the Demeyko
Range of north Chile and the state of preservation
Presumed original shellstructure and mineralogy

Actual state of preservationand silicification

Modiolus sp.

o : finely prismatic, calcite

o : finely prismatic, silicified

I : nacreous, aragonite

I : not preserved

Pinna sp.

o : simply prismatic, calciteo : simply prismatic, silicified

I : sheet nacreous, aragonite

I : apparently not preserved

Eopecten sp.

0: foliated?, calcite
I : not inferable

o : apparently structureless,

Plicatula sp.

o : foliated, calcite

o : foliated, not completely

Speciessilicified
I : not preserved

I : crossed· lamellar,
aragonite

silicified
I : apparently structureless,silicified

o : finel y foliated, calciteo : apparently structure less,I : crossed· lamellar, aragonite

silicified
I : not preserved

Antiquilima atacamensis
sp. nov.

o : finely foliated, calciteo : apparently structureless,I : crossed.lamellar, aragonite

silicified
I : not preserved

Gryphaea sp.

o : foliated, calcite

0: foliated, not completely
silicified
I : foliated, not completely
silicified
probably only outer layer preserved, structureless, silicified

Pseudolimea chongi sp. nov.

I : foliated, calcite

Unionites sp.

not inferable

Myophorigonia sp. aff.
M. paucicostata (JAWORSKI)o : prismatic, aragonite

Schafhaeutlia americana Cox

composite prismatic,
0: apparently structureless,silicified
aragonite
M: crossed-lamellar, aragonite M: apparently not preserved
I : complex crossed-lamellar, I : not preserved
aragoniteo : crossed-lamellar, aragonite 0: apparently structureless,I : complex crossed-lamellar,
silicified
aragonite
I : apparently structureless,incompletely silicified

Palaeocardita peruviana Cox

Septocardia peruviana (Cox)

I : lenticular, nacreous,aragoniteo : apparently structureless,silicified
I : not preserved0o : crossed-lamellar, aragonite o : apparently structureless,silicified
I : apparently structureless,silicifiedo : crossed-lamellar, aragonite o : apparently structureless,I : homogeneous or complex
incompletely silicified
crossed-lamellar, aragoni te I : apparently structureless,incompletely silicified
I : complex crossed-lamellar,
aragonite

I sopristes sp.

..

Planosplrzna sp.

not inferable

Preserved shell apparently
I structureless,silicified

Chartronella pacifica

not inferable"Outer layer" : silicified"Middle layer": not silicified
but for small quartz crystals"Inner layer" : silicified(JAWORSKI)

0: outer layer, M: middle layer, I: inner layer

683.

Triassic Molluscs from Chile(1972) and some others. However, thevanishing of aragonitic shells in thepresent material is not so exhaustive.
The shells of the Trigonioida and Veneroida, which are believed to have beenoriginally aragonitic, are often well preserved and silicified, though the original
shell fabrics are hardly observable. The
inner layer is generally silicified as wellas the outer in Septocardia peruvianaand sometimes also in Palaeocardita
peruviana. In such a case the hinge
structure and other internal characteristics are clearly observable by etching.
Judging from the deeply excavated adductor muscle scars, the myostracumaragonite probably vanished before thesilicification. On the other hand, theinner layer seems almost totally absent
in the specimens of Myoplwrigonia sp.
aff. M. paucicostata and Schafhaeutlia
americana. In some specimens of Palaeocardita peruviana and Isopristes sp. only
the external and internal surface issilicified like sandwich, and, when etched,
the middle" layer" of shell commonly
becomes hollow, indicating incomplete
silicifica tion. In this case the preservation does not seem to be controlled by
the original nature of shell. If theactual state of preservation is compared with the presumed original shellstructure, it may be said the prismatic
aragonite is commonly preserved andthat crossed-lamellar and complex
crossed-lamellar aragonite mayor may
not have been fossilized. Nacre seemf.
to be totally absent in the present material without any trace.
4) In a large number of specimens ofChartronella pacifica the outer and innersurface of shell is well silicified, while
the middle" layer" is quite incompletely.
By etching the middle" layer" becomesa cavity in which many microscopic
crystals of druse-fashioned quartz are

207

observed. This state of preservation
seems quite similar to the case of thegastropod fossils from the Pucara groupin the Cerro de Pasco area of central
Peru (HAAS, 1953, p. 9). Since the gastropod belongs to an extinct group,
their original shell structure and mineralogy are hardly inferable. HAAS (1953)
appears to have regarded the three" layers" of gastropod shells as original.
We do not think, however, they represent any original shell structure, because
the thickness of the cavity corresponding to the middle "layer" is quite irregular. Moreover, as noted before,
essentially similar phenomena are also
observed in some etched specimens of
Gryphaea sp., Palaeocardita peruviana andIsopristes sp. It is here suggested thatthe silicification proceeded from the external and internal surface and has notbeen completed in the middle part ofthick shells (see Text-fig. 3).
5) Generally speaking, the selective
preservation of fossils at the present
localities is believed to be primarily controlled by the structure and compositionof original shells, while the extent ofsilicification seems to vary in accordancewith local environmental condition.
Prismatic and foliated structures of calcite, regardless of the extent of silicification, are commonly well exhibited, butsuch microscopic original shell structuresof aragonite as crossed-lamellar are obliterated, even if the shells are preserved.
Since skeletal aragonite in permeablecarbonate rocks is said to have been
vanished or more commonly converted
to calcite (HALL and KENNEDY, 1967;
etc.), the obliteration may have occurredat the time of the conversion before thesilicification. The sandwich-like layered
structure of silicification in some molluscan shells are decidedly regarded ashaving nothing to do with the originalItaru HAY AMI. Shiro MAEDA and C. R. FULLER

208
OFC

IICA

~
SElECTIVE

PRESERVATIONCONVERSION TO CALCI TE. . .. ..... . ':','::.:. .. .....

.......................

','

D
Sill CIFICATION(NOT COMPLETED)SILICIFICATION
,'"

" "

.. ': . . ',' . : .,·········P>i
Th in-shelled Pterioids

Th ick-shelled Veneroids

Text.fig. 3. Inferred process of preservation for the bivalve fossils from theDomeyko Range. Left: complete silicification after the vanishing of inner layer [OFC:
outer foliated (or prismatic) layer of calcite, ICA: inner crossed·lamellar (or nacreous)layer of aragonite]. Right: incomplete silicification after the recrystallization of total
shell [OCA: outer crossed·lamellar layer of aragonite', ICA: inner complex crossedlamellar layer of aragonite]. These are not sketches but based on the actual state ofpreservation in the shells of Alltiquilima aiacamensis and Isopristes sp.

shell structure and mineralogy.

Modiolus sp. indet.

Repository

Plate 28, Figure 2

All the specimens described in thispaper are preserved in the Departmentof Earth Sciences, Faculty of Science,
Chiba University.

This species is represented by two
specimens: one is a left valve (Gc. 1001,
64.9 mm long, 42.3 mm high, 13.6 mm
thick), and the other is a broken conjoined specimen (GC. 1002). The nearlysmooth surface is divided into two subequal areas by a distinct diagonal carina.
The umbo is not terminal, the anterior
lobe being wide but not much inflated.
Judging from the general external characters, the present specimen is betterreferred to Modiolus than to Falcimytilus,
though specifically indeterminable.

Systematic description[by Itaru R"'YA~II and Shiro MAEDA]

Class Bivalvia
Order Mytiloida
Family Mytilidae RAFINESQUE, 1815
Genus i'vlodiolus LAMARCK. 1799

683. Triassic Molluscs from Chile
Occurrence.-Limestone at loco GCH 85.
Family Pinnidae LEACH, 1819
Genus Pinna LINNE, 1758
Subgenus Pinna LINNE, 1758
Pinna (Pinna) sp. indet.
Plate 28, Figures la, b

Six specimens (GC. 1003-GC. 1008) areavailable for the present study. All ofthem are conjoined but fragmentary.
The shells are completely silicified but
show simple prismatic structure along
the fracture, which is essentially thesame as that of the living species ofPinna. The apical angle is as small as
30 degrees. The median ridge is distinct
but becomes somewhat rounded in theadult stage. The transverse sections ofconjoined specimens, therefore, are subrhomboidal near the umbo and thickly
lenticular near the postero-ventral periphery. The largest specimen (GC. 1003)
exceeding 85 mm in width, provided withsome 20 flexuous weak radial riblets on
the dorsal slope of surface. The ventral slope is nearly smooth except forfine incremental lines, which indicate
rectangularly truncated and widely
gaped postero-ventral margin. The present species is undoubtedly referable toPinna (s. str.), but the incomplete material prevents us from determining its
specific name.
Occurrence.--Limestone at Loc. GCH 85.
Order Pterioida? Family Pectinidae RAFINESQUE, 1815

Genus Eopecten DOUVILLE, 1897
Eopecten sp. indet.
Plate 28, Figure 3

209

The present species is represented only
by an incomplete specimen (GC. 1009).
It is probably a left valve of an unnamed species of Eopecten, jUdging fromthe indistinctly differentiated anteriorauricle and mode of radial ribbing. The
radial ribs on the disk are as numerous
as 90 and irregular in prominence andbreadth, often increasing their number
by irregular insertion. The preserved
test is extremely thin.
Cox (1952) is of the opinion that some
Triassic species hitherto referred to
Eopecten [= Velata, Velopecten auct.]
should be assigned to Leptochondria
BITTNER, 1891. In fact, the present
species is somewhat similar to Leptochondria pascoensis Cox, 1949, from theUpper Triassic of Peru, but the shellsize is much larger and the radial ribs
are more numerous in the present species. In Europe Eopecten seems to havefirst appeared in the Lower Jurassic,
but in Japan an undoubted species ofthis genus has been known from theUpper Triassic. Eopecten has been traditionally regarded as belonging to thePectinidae, but actually there is somedoubt as to the family reference. In
fact, the radial ornamentation and deep
byssal notch with distinct ctenolium insome Jurassic species may remind ones
those of the Chlamydinae. Yet, theradial ribs of two orders in the left
valve of some other species, as well as
the plano-convex outline and indistinctly demarcated anterior wing of left valve,
seem to suggest some relationship to the
Aviculopectinidae and the Oxytomidae.
Therefore, this genus together with
Leptochondria may be an importantgroup for the consideration on the ancestry of the Pectinidae. Since the, preserved shell of the present specimen isunusually thin and may represent OI~YY
the outer layer, its original inner layer

210

Itaru HAY AMI, Shiro MAEDA and C. R. FULLER

was possibly composed of other material
than foliated calcite. Incidentally, theinner layer of a left valve is said to becomposed of nacre in the Aviculopectinidae and of crossed-lamellar" calcite"in the Oxytomidae (Cox et al., 1969;
ICHIKAWA, 1958). Because the present
species may be more or less deviatedfrom the type-species of Eopecten fromthe Jurassic, further studies on typical
species of this genus~ especially on theirshell structure, are needed to solve thistaxonomic problem.
Occurrence.-Limestone at Loc. GCH 85.
Family Plicatulidae WATSON, 1930
Genus Plicatula LAMARCK, 1801
Plicatula sp. indet.
Plate 28, Figure 4

An incomplete specimen (GC. 1010,
29.2 mm long, 31.8 mm+ high) is regarded
as a left valve of Plicatula owing to thecharacteristic hinge structure and muscle impression. Although its externalsurface was damaged during the etching,radial ribbing seems to be inconspicuous.
Occw-rence.-Limestone at Loc. GCH 85.
Family Limidae RAFINESQUE, 1815
Genus Pseudolimea ARKELL in DOUGLAS
and ARKELL, 1932
Pseudolimea chongi HAY AMI
and MAEDA, sp. nov.
Table 2.

Plate 28, Figures 5-8, 9a-c

Material. - Holotype: GC. 1011, left
valve. Paratypes: GC. 1012, conjoined
specimen; GC. 1013 - Gc. 1016, rightvalves; GC. 1017-GC. 1019, left valves.
Some other fragmentary specimens.
Diagnosis. - Medium-sized species of
Pseudolimea characterized by the noncarinate, gibbose and obliquely ovate
outline, not clearly demarcated auricles
and about 19 angular radial ribs with
one rib let of secondary strength on eachinterspace.
Description. - Shell equivalve, inequilateral, obliquely ovate, higher than long,strongly inflated; preserved test thin;anterior umbonal ridge undeveloped; nolunule; umbo highly protruded above
the dorsal margin; both auricles moderate in size, not much flattened, notclearly demarcated; byssal gape, is present, narrow; primary radial ribs as
many as 18 to 20, commonly roof-shaped
but sometimes bi-angulated; their interspaces provided with one much weakerriblet respectively, though such riblets
are very weak or invisible on the anterior auricle; internal structure unknown.
Remarks.-More than ten specimens
are available for the study. Their outerlayer is well silicified, but the peripheral
part is more or less broken in all the
specimens. The characteristic ornamentation composed of radial ribs of twoorders is well observable in every specimen, and the almost closed anterior

Measurements in mm [Pseudolimea cho7lgi sp. nov.]

Specimen
Left valve (GC. 1011), holotype
Conjoined valves (GC. 1012), paratype
Right valve (GC. 1013), paratype
Left valve (GC. 1017), para type

LengthHeightThickness

24.3
17.8
20.0+
29.6

27.2
20.1
26.2
31.1

9.8
13.3
8.4
9.9

683.

Triassic Mollt£scs from Chile

commissure and the ornamentation on
the anterior wing are recognized in theparatype (GC. 1012). The inner layer is
not preserved at all. Though the hinge
structure is not revealed, the present
species seems to be an early representative of Pselldolimea. It is in fact similar
to Pselldolimea hettangiensis (TERQUEM,
1855) from the lower Lias of western
Europe (DECHASEAUX, 1936; Cox, 1944),
but the present species has weakersecondary riblets and more obliquelyelongated shell. Some Carnian limids
from the St. Cassian beds of Alps described by BITTNER (1895) seem also tobelong to Pselldolimea, as were treatedby Cox (1~44), but the present species
is probably distinct from them, because
the shell-size is much larger.
Occurrence.-Limestone at Loc. GCH 85.
Genus Antiquilima Cox, 1943
Antiqllilima atacamensis HA YAM!
and MAEDA, sp. nov.
Plate 28, Figures 10-12

Material. - Holotype: GC. 1020, left
valve. Paratypes: GC. 1021, conjoined
specimen; GC. 1022, GC. 1023, left valves;
GC. 1024, GC. 1025, right valves. Someother fragmentary specimens.
Diagnosis. - Medium-sized species of
Antiqllilima characterized by the strong
incremental lines on the acutely triangular right anterior auricle, wide byssal
gape and about 18 radial ribs whichbear numerous fine radial threads andbecome squamose or knotty towards theventral periphery.

211

Description.-Shell of medium-sized forthe genus, slightly inequivalve, inequilateral, subovate, much higher than long,
not strongly inflated; preserved test
very thin; anterior umbonal ridge undeveloped, observed only near the umbo;
lunule not delimitted; umbo low, situatednear the mid-point of hinge-line; anteriorauricle comparatively large, obtusetriangular and rather flattened in theleft valve, whereas it is acutely triangular, well inflated and marked with erect
incremental lamellae in the right; byssal
gape wide in the right valve but almost
absent in the left; posterior auricle
moderate in breadth, obtusely truncatedin each valve; surface of disc ornamented with about 18 round-topped
radial ribs, which are simple on the umbonal and middle part but become squamose or even knotty near the ventral
margin of adult shell; radial ribs andinterspaces marked with numerous fineradial threads and fine incremental lines,
looking cancellate; fine radial ribs observable also on the posterior auricle;
ligament and internal structure unknown,
because the inner layer is not preservedat all.
Remarks.-The holotype reveals nearlycomplete outline, but other specimens
are not complete or even fragmentary.
The knotty radial are observed in theholotype and some of the para types andremind us strongly of the ornamentation of Ctenostreon, but other essential
characters are similar to those of Antiquilima. The inequality of anterior auricles between two valves is recognizedin the paratype (GC. 1021). In the an-Table 3. Measurements in mm [Antiquilima atacamensis sp. nov.]
Specimen
Left valve (GC. 1020), holotype
Right valve (GC. 1021), paratype-15.7·10.4+HeightThickness

59.1
54.8+

7.6
9.2

212

Itaru HAYAMI, Shiro MAEDA and C. R. FULLER

terior view of this specimen the byssal
gape is quite asymmetrical, that is tosay, the anterior margin is profoundly
concave in the right valve but only
slightly in the left. Although such a
feature has not been recorded in any
described species of Antiquilima, we presume .that it may be a common tendency
in this genus, because the anteriorauricle is acute-angled only in the rightvalve also in Antiquilima cubiferens(WHIDBORNE, 1883), as figured by Cox(1943). Acutely triangular anterior auricle with similarly erect incremental
lamellae is known in the right valvesof early representatives of Ctenostreon
such as C. japonicum HAYAMI, 1959, fromthe Lower Jurassic of west Japan.
The present species is anyhow morphologically transitional between Antiquilima and Ctenostreon. Although thedata are not necessarily sufficient to reconstruct the evolutionary lineage, it ispossible to consider that the species ofJurassic Ctenostreon are actually descendants from Some Upper Triassic
species of Antiquilima. Although Cox
HERTLEIN in Cox et al. (1969) noted that
Antiquilima ranges from Lias to Bajocian, KIPARISOV A et al. (1966) recorded
an undoubted species of this genus fromthe Norian·-Rhaetian of east Siberia.
BITTNER (1900, p. 207) suggested thepossible derivation of Ctenostreon from
Mysidioptera, but the former seems to
be more closely related to Antiquilima
than to the latter.
The unequality of the anterior auriclesof the present species suggests a byssate
sedentary life and subparallel or obliqueliving position to the surface of stable
substrate.
Occurrence.-Limestone at Locs. GCH
85 and GCH 26.

Family Gryphaeidae VYALOV, 1936
Genus Gryphaea LAMARCK, 1801
Gryphaea sp. indet.
Plate 29, Figures 1, 2This species is represented by three
specimens (GC. 1026-GC. 1028). They are
incompletely silicified, showing foliatedshell structure. One of them (GC. 1026,
34.0 mm long, 51.8 mm high, 24.5 mm
thick) is conjoined and better preserved
than the other specimens. The left valveof this specimen is strongly inflated,
provided with incoiled umbo, fairly wide
attachment area truncating the umbonal
region and a distinct posterior lobe
which is delimitted by a deep sulcus.
However, such a sulcus is undevelopped
in other left valves. No radial ornament is seen on the su"rface of twovalves. This is probably an undescribed
species of Gryphaea, but it is no difficult
to find any definite specific criterion,
because of the great morphological variability. In South America Gryphaea
darwini FORBES and a few other speciesof Gryphaea have been known from theLower Jurassic, but, so far as we areaware, there is no Triassic gryphaeid
comparable with the present species inthis continent. It has been said thatthe homeland of Gryplwea was the Arctic
region, because the occurrence of Triassic species was scarcely known fromother regions (STENZEL in Cox et al.,1971, p. NI070; etc.). If the present species is a true early representative of
Gryphaea, it may bear something informative to the long-pending question
about the origin and migration route ofthis genus.
Occurrence.-Limestone at Loc. GCH 85
and GCH 26.

683. Triassic Molluscs from Chile

213

Order Unionoida

Order Trigonioida

Family Pachycardiidae Cox, 1961

Family Trigoniidae LAMARCK, 1819

Genus Unionites WISSMANN, 1841

Genus Myophorigonia Cox, 1952

Unionites sp. indet.

Myophorigonia sp. aff. M. paucicostata

Plate 29, Figures 3a, b, 4

(JAWORSKI)

There are four conjoined specimens(GC. 1029-GC. 1032), the tests of which
are partly preserved but commonly
weathered or broken. Two specimensshow oblong and weakly carinate outline and closed posterior margin (GC.
1029, 53.6 mm long, 29.5 mm high, 24.0mm thick; GC. 1030, 55.2 mm+ long,
34.6 mm high, 23.6 mm thick). In an incomplete specimen (GC. 1031) the ventral
margin is slightly sinuated at the middle
and the test (only the outer layer) withfine incremental lines is well preserved.
Two specimens (Gc. 1029, GC. 1032)
shows the opisthodetic and external
position of ligament. The umbo is placedat about one-fifth of total length fromthe anterior end. Because of the absenceof inner layer, the hinge and muscle
structures are unknown. Although thegeneral outline may remind one of somespecies of Pachymya and certain pholadomyid genera, the completely closed
posterior margin, absence of radiallydisposed pustules, the position of ligament and smooth surface indicate thatthe present species should be referredto the Pachycardiidae. The anterior
located umbo and moderately large
shell-size may be comparable with thoseof some species of Trigonodus, but theoblong and weakly carinated outline andother external features seem to indicatethat the present species belongs toUnionites.
Occurrence. - Limestone at Loc. GCH
85.

Plate 29, Figures 5, 6
Compare.1922. Myophoria paucicostata JAWORSKI, Neues
Jahrb. J. Min. usw., Beil.·Bd. 47, p. 126,
pI. 5, figs. 9-11.
1929. MyopllOria paucicostata JAWORSKI:
STEIN:VIANN, Geologie von Peru, p. 63,
figs. 69A-E.
1937. Myophoria paucicostata JAWORSKI:
KORNER, Paleontographica, Bd. 86, Abt.
A, p. 184, pI. 12, fig. 4.
1949. Myophoria paucicostata JAWORSKI: Cox,
Bol. Inst. Ceol. Peru, no. 12, p. 25, pI. 1,
fig. 11.
1952. Myophorigonia paucicostata (JA\'VORSKI) :Cox, Proc. Malac. Soc. London, vol. 29,
pts. 2-3, p. 52, pI. 3, fig. 3.
1969. Myophorigonia paucicostata (JAWORSKI) :Cox in Cox et al., Treatise on Inver·tebrate Paleontology, Part N, p. N485,
fig. D71-3.

The present spec.ies is represented bythree fragmentary specimens (GC. 1033GC. 1035): one is a left valve and two
are right. They show similarly eight or
so tuberculated and highly elevated radial ribs on the anterior slope of discas typical specimens of lvlyophorigonia
pallcicostata from the Upper Triassic ofPeru. In comparison with the Peruvian
specimens illustrated by JAWORSKI (1922)
and Cox (1949), however, the marginal
sulcus is narrower and merely as wide
as the interspaces between other radialribs, and the shell size seems to be muchlarger. The height of those Peruvian
specimens scarcely exceeds 20 mm inlength and height, whereas the largest

214

Itaru HAYAMI, Shiro MAEDA and C. R. FULLER

specimen (GC. 1033) in the present collection, though its original outline cannot be reconstructed, is evidently taller
than 45 mm. In these respects the present specimens are more similar toanother Peruvian specimen figured by
KORNER (1937) and probably specificallydistinct from JAWORSKI'S. The materialis, however, too insufficient to propose
a new taxon.
Occurrence.-Limestone at Loc. GCH 85.
Order Veneroida
Family Fimbriidae NICOL, 1950
Genus Schafhaeutlia COSSMANN. 1897
Schafhaeutlia americana Cox
Plate 29, Figure 7
1949. Schafhaeutlia americana Cox, Bol. Inst.
Geol. Peru, no. 12, p. 31, pI. 2, figs. 1, 2.

This species is represented only by aleft valve (GC. 1036, 49.1 mm long, 48.2
mm high, 20.2 mm thick). The preserved
test is unusually thin and probably represents only the outer layer. In viewof the rounded outline, concentric ornamentation and strong convexity thepresent specimen is assignable to Schafhaeutlia americana Cox 1949, which was
originally described from the Upper
Triassic of Peru, though the hinge andother internal structures are unknownat all.
Occurrence.-Limestone at Loc. GCH 85.
Family Carditidae FLEMING, 1828
Genus Palaeocardita CONRAD, 1868
Palaeocal'dita peruviana Cox
Plate 30, Figures la, b, 2-4, 5a-c
?1937. Cardita aft. beneckei BITTi\'ER: KORNER,

Palaeontographica, Bd. 86, Abt. A, p. 192,

pI. 12, fig. 7.
1949. Palaeocardita peruviana Cox, Bot. Inst.
Geol. Peru, no. 12, p. 31, pI. 2, fig. 4.

Material. - GC. 1037-GC. 1039, rightvalves. GC. 1040-GC. 1042, left valves.
Many other incomplete specimens.
Description.-Shell of medium or large
size for the genus, sometimes exceeding
45 mm in length, equivalve, highly inequilateral, trapezoidal to oblong, about
1.4 times longer than high, strongly inflated; test moderate in thickness; antero-dorsal margin short, never concave,sloping down into the rounded anterior
margin; postero-dorsal margin long,nearly straight or feebly convex, passing
gradually into the posterior; posteriorcarina extending from the umbo to thepostero-ventral corner where the posterior margin forms a nearly right angle
with the ventral; maximum convexity
lying on the carina; umbo prominent,
angular, slightly opisthogyrous, placedat about one-fourth of shell from theanterior end; lunule and escutcheon notdiscriminated; surface ornamented with
20-22 roof-shaped radial ribs; one ofwhich coincides with the carina andabout eight of which are distributed on
the posterior area behind the carina;
interspaces of radials also angular, never
flattened; somewhat scaly incremental
lines crossing the radials; hinge plate
moderate in breadth, provided with carditid-type teeth as formulated: 3a 3b PI/
2 4b PH; 3a elongated and curved along
the pre-umbonal margin; 2 very opisthocline, narrow; 3b acline, triangular, neverbifid, massive and highly elevated; 4brather thin; PI and PII strong but short,
remote from the cardinals; anterior adductor scar deeply impressed, borderedposteriorly by a buttress; inner ventral
margin coarsely crenulated in accordance with the external radial ribs.

683.
Table 4.

Triassic Molluscs from Chile

215

Measurements in mm [Palaeocardita peruviana Cox]

Specimen

LengthHeightThickness

Right valve (GC. 1037)
Right valve (GC. 1038)
Right valve (GC. 1039)
Left valve (GC. 1040)
Left valve (GC. 1041)

26.6+
28.1
46.0
30.1
30.0

21. 8
19.9
33.9
21. 9
20.1+

9.6
9.5
16.8
9.7
9.6

Remarks.-More than 20 silicified specimens are available for the study, but
few of them are complete. In every
specimen the surface ornaments are wellobservable, but the inner layer mayor
may not be preserved. The hinge structure is best revealed in two specimens(GC. 1037, GC. 1040), which were takenout from blocks of limestone by diluted
HC!. The present specimens seem to bereferable to Palaeocardita peruviana Cox,
1949, originally described from the Upper
Triassic of Peru, because all the essential
characters are the same. Although theradial ribs are apparently more widely
spaced on the posterior area in the Peruvian specimen (Cox, 1949, pI. 2, fig. 4), thedifference may to due to variation within
one and the same species. KORNER (1937)
figured a similar carditid specimen also
from Peru under the name of Cardita
aff. beneckei. Cox (1949) suggested that
it is conspecific with P. peruviana, but
the number of radial ribs, 29 according
to KORNER, is too numerous and theumbo is placed very anteriorly for thisspecies, jUdging from the individual
variation of the present material. KORNER(1937) proposed at the same time thegenus Schizocardita for an aberrant
carditid also from the Upper Triassic
of Peru, which possesses a rostrum-likeacuminate postero - ventral projection.
Although the taxonomic evaluation ofthis character is not yet clear, it is interesting that the hinge and other essential characters of the type-species ofthat genus: Schizocardita cristata KORNER, 1937, are fairly similar to those of
the present species. Such a projectionis, however, not recognized from theincremental lines of any specimen in thepresent collection.
Occurrence.-Limestone at Loc. GCH 85
and GCH 26.
Family Cardiidae LAMARCK, 1809
Genus Septocardia HALL and
WHITFIELD, 1887
Septocardia peruviana (Cox)Plate 29, Figures 8a, b, 9a-c, 10
1949. Pascoella peruviana Cox, Bol. Inst. Ceol.
Peru, no. 12, p. 35, pI. 1, figs. 9, 12-15,?pI. 2, fig. 5.
1969. Septocardia typica HALL and WHITFIELD:
KEEN in Cox et al., Treatise on Invertebrate Paleontology, Part N, p. N586,
figs. E85-1a-f (non 19, h) [non S. typica
HALL and WHITFIELD, 1877].

Material.-GC. 1043-GC. 1045, left valves.
Description.-Shell inequilateral, suborbicular to gibbose, nearly as long as
high, strongly inflated; test heavy; antero- and postero-dorsal margins short,
gently arcuate, passing gradually into
venter; umbo subcentral, highly salient,slightly but distinctly opisthogyrous;
lunule and escutcheon not delimitted;
surface ornamented with about 19 roofshaped scaly radial ribs which are comparatively wide on the anterior-middle

216

Itaru HAYAMI, Shiro MAEDA and C. R. FULLER
Table 5. Measurements in mm [Septocardia peruviana (Cox)]
Specimen

LengthHeightThickness

Left valve (GC. 1043)
Left valve (GC. 1044)

24.3
22.2

24.7
21.1

12.6
10.5

surface and densely spaced on the posterior area; bottom of the interspace
also angular and never flattened; ligament short, inserted in an oblique furrow
bordered by the nymph, hinge plate
wide, provided with two cardinal teeth(2, 4b) and one posterior lateral tooth(PH) in the left valve; cardinal 2 very
opisthocline, elongated, highly elevated;
4b also stout, prosocline; 3a and 3b
probably connected below the umbo,
judging from the shape of their sockets
in the left valve; PII tusk-like, well developed; no anterior lateral tooth; anterior adductor scar subovate, deeply
excavated, bordered posteriorly by aprominent buttress, while the posterior

one is obscure; inner ventral margin
strongly crenulated in accordance withthe external radials.
Remarks.-In this collection the present
species is represented by three silicified
left valves which were obtained byetching. Two of them (GC. 1043, GC.
1044) are well preserved and nearlycomplete, showing both the external andinternal characters. The other specimen(GC. 1045) is incompletely silicified, andits hinge plate and inner layer were
mostly dissolved. They are quite similarin every essential character to the specimens from the Norian at several localities in central Peru, which were described by Cox (1949) under the name

Explanation of Plate 28
Fig. 1. Pinna sp. indet.
1a: Right valve of a conjoined specimen (GC. 1003) xl; 1b: part of fracture profile in
the postero.dorsal area of the same specimen, revealing simple prismatic structure, x 4.
Fig. 2. Modiolus sp. indet.
2: Left valve (GC. 1001) xL
Fig. 3. Eopecten sp. indet.
3: Left valve (GC. 1009) xl.
Fig. 4. Plicatula sp. indet.
4: Internal view of a left valve (etched specimen) (GC. 1010) xL
Figs. 5-9. Pseudolimea chongi HAYA:\-I1 and MAEDA, sp. nov.
5: Right valve (GC. 1014), paratype, x 1.5.
6: Left valve (GC. 1011), holotype, x 1.5.
7: Right valve (GC. 1016), paratype, x 1.5.
8: Right valve (GC. 1013), para type, X 1.5.
9: Conjoined valves (GC. 1012), paratype, x 1.5. 9a: right view, 9b: left view, 9c: anterior view.
Figs. 10-12. Antiquilima atacamensis HAYAMI and MAEDA, sp. nov.
10: Right valve of a conjoined specimen (GC. 1021), paratype, xL
11 : Left valve (GC. 1022), paratype, xL
12 : Left valve (GC. 1020), holotype, xl.
All specimens from Loc. GCH 85. Photo by HAYAMI with whitening.

HAY AMI, MAEDA and FULLER:

Triassic Mollusca from Chile

Plate 28

683.

Triassic Molluscs from Chile217

of Pasco ella peruviana. Although theaverage shell-size may be somewhat
smaller than in Cox' specimens, the hinge
structure, outline and mode of radialribbing are quite identical. Keen in Cox
et al. (1969) refigured Cox's specimens
under the specific name of Septocardia
typica HALL and WHITFIELD, which was
originally based on the material fromthe Upper Triassic of Nevada. This
treatment, however, does not go downwith us, because the original specimensof S. typica, as illustrated by HALL and
WHITFIELD, show more inequilateral andrectangular outline, thinner test and lessangular radial ribs and interspaces. We
regard here Pasco ella peruviana as a
distinct species of Septocardia. Cardita
n. sp. ex aff. giimbeli Pichler-pichleri
BITTNER described by KORNER (1937)
may be also another representative of
this genus, though the shell-size is muchsmaller than the present species.
Occurrence.--Limestone at Loc. GCH 85.

ventral rim of each imbrication suggest
the presence of such radial ornaments.
The hinge structure and wide Cardinialike pseudolunule are, though obscurely,
exposed in an incomplete right valve(GC. 1048), which was obtained by etching. JUdging from these characteristics,
the present species is certainly referable
to the genus Isopristes, which was orignally founded on a species from theUpper Triassic of Peru. In comparison
with Isopristes crassus NICOL and ALLEN,
1953, however, the shell of the present
species seems to be more globose, andthe concentric lamellae are more densely spaced.
Occurrence.-Limestone at Loc. GCH 85.

Family Cardiniidae ZITTEL, 1881

Planospirina sp. indet.

Genus Isopristes NICOL and ALLEN, 1953

Plate 30, Figures Sa, b

Isopristes sp. indet.

Only a single silicified specimen (GC.
1050, 29.4 mm in max. diameter, 20.8 mm
in min. diameter, 22.1 mm in height) was
obtained by etching. It shows low spire,descending suture near the aperture,somewhat irregularly undulated surface
of upper whorl and gibbose aperture
with feebly concave inner lip and inductura of moderate breadth, although theexternal surface and the marginal partof outer lip are_ incomplete. These
characteristics seem to indicate that thepresent species belongs to the Naticopsinae, probably the genus Planospirina.
In comparison with Nerita esinensis
STOPP ANI, 1858, from the Ladinian of
Tyrol, the type-species of Planospil'ina,
many characters are quite similar, but

Plate 29, Figures 11a-c, 12This species is represented by four
specimens; two (GC. 1046, 35.0 mm long,
35.2 mm high, 26.1 mm thick; GC. 1047,
35.3 mm long, 32.6 mm high, 26.3 mm
thick) are conjoined valves, one (GC.
1047) is a left valve and one (GC. 1048)
is a fragmentary right valve. They are
incompletely silicified and generally poorly preserved, but equidistantly spaced
and somewhat imbricated concentriclamellae are clearly observed on theanterior part of a conjoined specimen(GC. 1046). Small radial ribs of Isopristes
type are actually invisible on the surface, but numerous fine crenules on the

Glass Gastropoda
Order Archaeogastropoda
Family Neritopsidae GRAY, 1847
Genus Planospirina KITTL, 1899

218

Itaru HAY AMI, Shiro MAEDA and C. R. FULLER

the aperture is semicircular and higher
than broad in the present specimen.
Occurrence.-Limestone at Loc. GCH 85.
Family Paraturbinidae COSSMANN, 1916
Genus Chartronella COSSMANN, 1902
Chartronella pacifica (JAWORSKI)
Plate 30, Figures 9-11

1922. EucyclllS pacificus JAWORSKI, Neues
Jahrb. j. Min. usw., Beil.·Bd. 47, p. 140,
pI. 4, fig. 11.
1949. Eucyclus hartisoni Cox, Bol. Illst. Ceol.
Peru, no. 12, p. 37, pI. 2, figs. 17, 18.
1953. Chartroniella pacifica (JAWORSKI) [sic]:
HAAS, Bull. Amer. Mus. Nat. Hist. vol.
101, p. 81, pI. 5, figs. 31-41, 45-47, 54.

Several incomplete specimens are available for the study. Two of the illustrated
specimens (GC. 1051, 28.6 mm in max.
diameter, 26.1 mm in min. diameter, 40.3
mm in height; GC. 1052, 28.0 mm in max.
diameter, 27.1 mm in min. diameter, 38.2mm in height) show nearly complete
outline, though their outer lips are partly
broken. They are incompletely silicified
and were obtained by partial etchingfrom limestone blocks. Judging fromthe high-turbinate shell, obliquely growing last whorl with unusually deep suturenear the aperture, gradually weakened
and sometimes obscurely serrated keel,
well marked growth lamellae on thebase, obscure spiral ribs on the upper
whorl and very prosoclinal outer marginof aperture seem to be essentially thesame with those of Chartronella pacifica(JAWORSKI) from the Upper Triassic ofPeru. If compared with many silicified
specimens of various ontogenetic stages,
which were fully described by HAAS
(1953) on the material from the Cerrode Pasco area, the mode of allometric
growth, especially the weakened keelwith growth and obliquely descending
suture of the last whorl, is quite similar,
endorsing the present specific identification.

Explanation of Plate 29
Figs. 1, 2. Cryphaea sp. indet.
1 : Left valve of a conjoined specimen (GC. 1026) xl.
2: Left valve (GC. 1027) xl.
Figs. 3, 4. Unionites sp. indet.
3: Conjoined valves (GC. 1029) xl. 3a: left view, 3b: right view.
4: Right valve of a conjoined specimen (GC. 1031) xl.
Figs. 5, 6. Myophorigonia sp. aft. M. paucicostata (JAWORSKI)
5: Left valve (GC. 1034) xl.
6: Right valve (GC. 1033) xl.
Fig. 7. Schajhaelltlia americana Cox
7: Left valve (GC. 1036) xl.
Figs. 8-10. Septocardia peruviana (Cox)
8: Left valve (etched specimen) (GC. 1044) x 1.5. 8a: external view, 8b: internal view.
9: Left valve (etched specimen) (GC. 1043) x 1.5. 9a: external view, 9b: internal view,
9c: upper view.
10: Left valve (etched specimen) (GC. 1045) x 1.5.
Figs. 11, 12. Isopristes sp. indet.
11: Conjoined valves (GC.1046) xl. 11a: left view, llb: right view, 11c: anterior view.
12: Internal view of a fragmentary right valve (GC. 1048) x 1.5.
AIl specimens from Loc. GCH 85. Photo by HAY AMI with Whitening.

HAY AMI, MAEDA and PULLER:

Triassic Mollusca from Chile

Plate 29

683.

T1"iassic Molluscs from Chile

Occurrence.-Limestone at Loc. GCH 85.

Incertae sedis
Gastropod genus and species indet.
Plate 30, Figures 6, 7a, b

Two incomplete specimens of a gastropod (GC. 1059, 1060) are at a glance
similar to the specimens of the preced-·
ing species, but the spire seems to bemuch lower and the base is much flatter.
In the trochiform outline they resemble
some specimens of Chartronella wortheniaeformis (Cox, 1949) from the Upper
Triassic of central Peru (HAAS, 1953),
but the spiral ribs on the upper whorl
and base are much stronger.
Occurrence.-Limestone at Loc. GCH 85.
Concluding remarks

1) As a result of the geological andpaleontological survey to the south Andes
performed in 1970, a molluscan faunule,
which was found in the limestones exposed on the eastern slope of the Domeyko
range about 180 km southeast of Antofagasta, was described in this paper.
The bivalves consisting 13 species include two new limids : Pseudolimea chongi
sp. nov. and Antiquilima atacamensis sp.
nov. The latter species appears to foretell some characteristics of Ctenostreon.
The internal characters of Palaeocardita
peruviana and Septocardia peruviana were
well observed on the basis of somesilicified specimens taken out from limestone blocks by etching.
2) In some bivalve specimens, particularly in the Pteroida, the outer layer(originally composed of calcite) is wellpreserved, whereas the inner layer (originally composed of aragqnite) is totallyabsent without any trace. The vanishing of originally aragonitic layer is,219

however, not exhaustive, since the testsof the Veneroida and the outer layer of
the Trigonioida are often well preserved.
The secondary silicification of molluscan
fossils does not seem to be related tothe selective fossilization, because themiddle "layer" of some bivalve andgastropod shells becomes hollow byetching regardless of the difference ofinferred original shell structure andmineralogy.
3) The present molluscan faunule shows
marked similarity to the Upper Triassic(mainly Norian) fauna of the PucaraGroup in Cerro de Pasco area of Central
Peru. In addition to the above mentioned Peruvian species, Schafhaeutlia
americana and Chartronella pacifica arecertainly common elements, and there
are some species belonging to Myophorigonia and Isopristes which were also
originally proposed on Peruvian materials. hi Chile some Middle Triassic(mainly Anisian) molluscs were describedfrom the environs of Alto de Carmen
in Province of Atacama (ZEIL, 1958;
BARTHEL, 1958) and some Anisian and
Norian-Rhaetian sequences were known
in the coastal region of Province ofCoquimbo and Aconcagua (CECIONI and
WESTERMANN, 1968), but the occurrenceof Pucara type fauna has not been recorded. Taking also the resemblance oflithology into consideration, it is reasonable to regard the present fossiliferous
limestones as belonging to the southern
extension of the Pucara Group.
References
K. W. (1958) : Eine marine Faunule
aus der mittleren Trias von Chile. Neues
Jahrb. f. Ceol. Paliiont., Abh., vol. 106,
p. 352-382, pis. 19, 20.
BITTNER, A. (1895) : Lamellibranchiaten deraipinen Trias. 1. Revision der Lameilibranchiaten von St. Cassian. Abh. k. k.
BARTHEL,

220

Itart£ HAYAMI, Shiro MAEDA and C. R. FULLER

geol. Reichsanst., vol. 18, no. 1, p. 1-236,
pIs. 1-2-1.
- - (1900) : Ueber nachtriadische Verwandte
der Gattung Mysidioptera. Verh. k. k. geol.
Reichsanst., 1900, p. 207-208.
CARTER, R.M. (1972) : Adaptations of British
Chalk Bivalvia. Jour. Paleontology, vol.
46, no. 3, p. 325-340, pIs. 1-3.
CECIONI, G. and WESTER:l:IANN, G.E.G. (1968) :The Triassic/Jurassic marine transitionof coastal central Chile. Pacific Geology,
vol. 1, p. 45-71.
Cox, L. R. (1943): The English Upper Liasand Inferior Oolite species of Lima. Proc.
Malacol. Soc. London, vol. 25, nos. 5-6,
p. 151-187, pIs. 6-29.
- - (19-1-1): On Pseudolimea Arkell. Ibid.,
vol. 26, nos. 2-3, p. 74-88, pIs. 2, 3.
- - (1949) : Upper Triassic Mollusca from
Peru. Bol. Inst. Geol. Peru, no. 12, p. 150, pIs. 1, 2.
- - (1952a): The Jurassic lamellibranchfauna of Cutch (Kachh). No.3. Palaeont.
Indica, ser. 9, vol. 3, no. 4, p. 1-128, pIs.
1-12.
- - (1952b) : Notes on the Trigoniidae, withoutlines of a classification of the family.
Proc. iHalacol. Soc. London, vol. 29, nos.
2-3, p. 45-70, pIs. 3, 4.
- - et al. (1969, 1971): Treatise on inverte-

brate paleontology. Part. N. Bivalvia (3vols.). Vols. 1 and 2, 952 pp. [1969J; Vol.
3 (oysters by STENZEL, H. B.), 272 pp.
[1971J. Geol. Soc. America and Kansas
Univ.
DECHASEAUX, C. (1936): Limides jurassiquesde Pest du bassin de Paris. Mem. Mus.
Roy. Hist. Nat. Belgique, ser. 2, vol. 8,
p. 1-58, pIs. 1-3.
HAAS, O. (1953) : Mesozoic invertebrate faunasof Peru. Part. 1. General introduction,
Part. 2. Late Triassic gastropods from
central Peru. Bull. Amer. Mus. Nat. Hist.,
vol. 101, p. 1-328, pIs. 1-18.
HALL, A. and KENlXEDY, W. J. (1967) : Aragonite in fossils. Proc. Roy. Soc. ser. B,
vol. 168, p. 377-412.
HAYA:l:II, 1. (1959): Lower Liassic lamellibranch fauna of the Higashinagano formation in west Japan. Jour. Fac. Sci.
Univ. Tokyo, sec. 2, vol. 12, pt. 1, p.3184, pIs. 5-8.
ICHIKAWA, K. (1958): Zur Taxionomie und
Phylogenie der triadischen "Pteriidae"(Lamellibranch.) mit besonderer Beriicksichtigung der Gattungen Claraia, Eumorphotis, Oxytoma und Monotis. Palaeontographica, vol. 111, pt. A, p. 131-212, pIs.
21-2-1.
JAWORSKI, E. (1922): Die marine Trias in

Explanation of Plate 30
Figs. 1-5. Palaeocardita peruvian a Cox
1: Right valve (etched specimen) (GC.1037) x 1.5. 1a: external view, 1b: internal view.
2: Left valve (GC. 1042) x 1.5.
3: Left valve (etched specimen) (GC. 1041) x 1.5.
4: Right valve (GC. 1038) x 1.5.
5: Left valve (etched specimen) (GC. 10-10) x 1.5. 5a: external view, 5b: internal view,
5c: upper view.
Figs. 6, 7. Gastropod genus and species indet.
6: Upper view of an incomplete etched specimen (GC. 1060) x 1.5.
7: Apertual view of an incomplete specimen (GC. 1059) x 1.5.
Fig. 8. Planospirina sp. indet.
8: Partly etched specimen (GC. 1050) x 1.5. 8a: apertural view, 8b: upper view.
Figs. 9-11. Chartronella pacifica (JAWORSKI)
9: Apertural view of an incomplete specimen (GC. 1053) x 1.5.
10: Adapertural view of a partly etched specimen (GC. 1052) x 1.5.
11 : Apertural view of a partly etched specimen (GC. 1051) x 1.5
All specimens from Loc. GCH 85. Photo by HAYAJ.II with whitening.

HAY AMI, MAEDA and FULLER:

Triassic Mollusca from Chile

Plate 30

683.

T1'iassic Molluscs from Chile

Siidamerika. Neues Jahrb. f. Min. usw.,
Beil.-Bd., vol. 47, p. 93-200, pIs. 4-6.
KENNEDY, W. J. (1969): The correlation of
the Lower Chalk of southeast England.
Proc. Geologists' Assoc., vol. 80, pt. 4, p.
459-560, pIs. 15-22.
KIPARISOVA, L.D., BYCHKov, J.M. and POLVBOTKO, I. V. (1966) : [Late Triassic bivalve
Mollusca from east Siberia, USSR.] 230
pp., 40 pIs. All Soviet Union Geol. Inst.
(VSEGEI). [in Russian]
KITTL, E. (1899) : Die Gastropoden den Esinokalke, nebst einer Revision de Gastropoden der Marmolatakalke.
Ann. k. k.
Naturhist. Hofmus. Vienna, vol. 14, p. 1237, pIs. 1-18. [ not seen]
KORNER, K. (1937) : Marine (Cassianer-Raibler)
Trias am Nevado de Acrotambo (NordPeru). Palaeontographica, vol. 86, pt. A,
p. 145-237, pIs. 10-14.
MAEDA, S., YAMAGIWA, N., FULLER, C.R.
and CHONG, D. G. (1974) : Thecosmilia sp.
from Cordillera Domeyko, Chele.
In

221

MAEDA, S. (ed.) : Palaeontological study
to the Andes, p. 23-27. Geol. Lab. Chiba
Univ., Chiba.
NICOL, D. and ALLEN, W. T. (1953) : A new
pelecypod genus from Upper Triassic
strata in Peru. Jour. Washington Acad.
Sci., vol. 43, no. 11, p. 344-346.
STEINMANN, G. (1929) : Geologie von Peru. 448
pp., 9 pIs. Heidelberg.
TAYLOR, ]. D., KENNEDY, W. J. and HALL, A.
(1969) : The shell strllcture and mineralogy of the Bivalvia. Introduction, Nuculacea-Trigonacea. Bull. Brit. Mus.
Nat. Hist., Zoology, suppl. 3, p. 1-125, pIs.
1-29.
- - , - - and - - (1973) : The shell structure and mineralogy of the Bivalvia.
Lucinacea - Clavagellacea, Conclusions.
Ibid., vol. 22, no. 9, p.253-294, pIs. 1-15.
ZEIL, W. (1958): Marine mittlere Trias der
Hochkordillera, Provo Atacama. Neues
Jahrb. Geol. Piilaont., Ablz., vol. 106, p.
339-351, pIs. 17, 18.

222

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Upper Jurassic and lower Cretaceous radiolarians from the Shimanto group in
Tokushima Prefecture, Japan ...................................................... ··················NAKASEKO, K.
Cryptocephalic and Cryptothracic Nassellaria from the Shimanto group in Toku·shima Prefecture, Japan (Part 1) ····································NISHIMURA, A. and NAKASEKO, K.
m;l£fl1LER~(I)7.tfjt~;:: ,,?~, -C ...................................................... ····································.HW1.\!.(~-(I):a-II (E"{#t. =~*) (l)r$.t f)~Lt-:li'Y'i!ffl1L!E.c;T,..GnQ{tE"~;::0l'-C·········~ii{~T· ~~7-.~-(I):a-II (E"{#t. =~*) (I){tE"mrr:~(I)¥J~tlt ................................. ~~7-.~. ~~7-.~. ~ii~T
f.KaE"EJ(~pj1l:f~ Heterophyllidae ~;:: ,,?~, -c ...................................................... ···············j1<.i1ltt~
fJCaE" ~EJ(F.i1l:fr$m (l)1£f§ c ~f§~;:: ,,?~, -C ..................................................................... :llt#~1 ;f-=f~::r·~Jt~rI:I~u,.. GnQ1&=f1 ;(;{1jij)(;c.:t(l)~ij)(;~;i:'J;::"?~'-C························ri1liE-· /NI~'&I£
On Siphonofusus and Euthria of the Indo·West Pacific ·····.·······································SHljTO, T.
Miocene molluscs from the Macasilao and Malabago formations, Negros Islands,
the Philippines ...................................................... ········.·······································SI-IUTO, T.
A new species of Bankia in Paleogene Matsuiwa, Amakusa, Kumamoto Prefecture·········*~~~
1illJP.JTrnilffi1l:f(l)~Jt{tE", C «~;:: "Cerithium" pyramidaeforme NAGAO, 1930 ~;::"?~'-C ...

...............................................................................................................

~J/Jl!t'*~.

:!Ii'i*~:g:A new species of Pterotrigonia from Upper Cretaceous Onogawa group, Japan ············mtl ~'"7!/"J] A"J] Jv/i!i; ~ 1""jfu~~1 / -I! '7 J...A ...................................................... ···!l!fm~z • ~iIEln't
l'h~Q Mediterranean Province ~;:::t-oltQ Aptian (I)':::'j!jJt~t;::"?~'-C(l)~~ ··················$!l!fJ'ttt
Some biometric aspects of Nautilus marcomphalus ···························HIRANO, H. and OBATA, 1.
Pulse and locomotion of Nautilus macromphalus in captivity······························...................................................... ·····················MIKA;\I1, S., HIRANO, H. and HAMADA, T.
Nautilus izumoensis Yokoyama (I):fr~~B':J~~t··········································!I!f;ft1$7C • tj\~W-~Imf!~
A Danian nautiloid from Majunga, northwest Madagascar ···············OBATA, 1. and KAl'IE, Y.
Discoactinoceras and the Discoactinoceratidae fam. nov. ·· .. ·.······························KOBAYASHI, T.
One some ammonoids from the Aohama formation of Kitakyushu City, Japan ······NISHIDA, T.
~l:;~~J:$B!lli*.t f) ~i" Q Apiotrigonia hetonaiana sp. nov., Senis japonica sp. nov.
(I) =itiffl t;:: ,,?~, -C ...................................................................................................... mf\;iEz
illciIW-AiliJ!Gl£::r 1/ p ~ 3 :"-$(1)7 :"-"",1"1 r t;::"?l'-C ...................................................... t.t*iif!~~l:;~~~~l:;J:$itifm.:::.*~)j3JtJ(lt~ 3 itifji, Arakia, Okabea :t-o.t O~ Oshitea (8 itiffla-~ts) .........................................................................................................................../J\;j:j;j!g-

Jv-7W~f~iPi:a-fli" Q i=3~lt~itiff4 Japexiaceae (~~l:;J:$itifm.:::.*~, 1 itifji 1 itiffl) ·········/Ni;j!gComparison between paleotemperature by Emiliani et aI. (1974), change of solar
radiation and paleoclimate based on palynological analyses of samples from
Lake Biwa ...................................................... ·································FU]I, N. and HORIE, S.
Palynological study of the deposits in Lake Biwa, central Japan. II. Paleoclimatebased on pollen analyses of the upper 60 m of the 200 m core············FuJI, N. and HORIE, S.
"Pseudoschizaea" from the Quaternary Sediments of the Ariake sea area·········TAKAHASHI, K.
1iii#Jjjljl:;$(I)$l/1Tt!:t;fl4t!J1l:ft;::"?~, -C ................................................................................. JEJ:~
~1li{ittc;fl4t!JlffK~tI:! L k*~&lf~f*(I)itiffjIH;:: ,,?~, -C ...................................................... ···t.t~3'5;f1l
A revision of genus Conulariopsis Sugiyama, 1942 ················································MURATA, M.
A fossil Amphineura from the Permian Akasaka Limestone, Gifu Prefecture,
Japan ............................................................................................................ ···MURATA, lVT.

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:5!f~71jfil;Q)JU~~!\\ft::. Cytherelloidea ~t::."?\,'-C·······························································!l!fJjJ(iVl~
Permo.Carboniferous trilobites from Thailand and Malaysia···KoBAYASHI, T. and HAMADA, T.
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Transactions and Proceedings of the PalaeontologicalSociety of Japan

November 15, 1977

New Series No. 108

CONTENTS
TRANSACTIONS
682. YAMAGUCHI, Toshiyuki: Taxonomic studies on some fossil and recent
Japanese Balanoidea (Part II) ........................................ 161
683. HAYAMI, Itaru, Shiro MAEDA and Carlos Ruiz FULLER: Some Late Triassic
Bivalvia and Gastropoda from the Domeyko Range of North Chile .... 202
PROCEEDINGS ...................................................•............ 222