ISSN

0031-0204

fJ ~.. ~i.~~/.t'~~·!-l.t
Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 125

Pala eontologica l Society of Ja pan
Apr il 15, 1982

Itaru HAY AMI and Ikuwo OBAT A
Co-Editors
Associate editors Teruya UYENO and Tomowo OZAWA

Officers for 1981--1982
President:

Tetsuro HANAI

Councillors:

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

Yoshikazu HASEGAWA, Itaru HAYAMI, Hisayoshi IGO, junji ITOIGAWA, Tadao
KAMEl, Tatsuaki KIMURA, Ikuwo OBA T A, Tsunemasa SAITO, Tsugio SHUTO .
Yokichi TAKAYANAGI, Toshimasa TANAIMembers of Standing Committee:

Kazuo ASAMA (Finance), Kiyotaka CHINZEI (General

Affairs), Takashi HAMADA (Membership), Itaru HAY AMI (Transactions), Hisayoshi IGO (Foreign Affairs),

Tatsuaki KIMURA (Planning),

Ikuwo

OBATA

(Transactions), Tsugio SHUTO (Special Papers), Yokichi T AKAYANAGI ("Fossils")
Secretaries:

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

Teruya UYENO and Tomowo OZAWA (Transactions), Hisaharu IGO (Planning),
Ienori FUJIYAMA and juichi YANAGIDA (Special Papers), Kunihiro ISHIZAKI("Fossils")
Auditor:Kenji KURIHARA

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

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

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

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 125, pp. 239-258, pIs. 39, 40, April 15, 1982CRETACEOUS TETRAGONITID AMMONITE JAWS:
A COMPARISON WITH MODERN NAUTILUS JAWS*

740.

YASUMITSU KANIE
Yokosuka City Museum, Yokosuka 238

Abstract. The morphology and mineral composition of jaw apparatuses in
Tetragonites glabrus, Gaudryceras denseplcatum, G. tenuiliratum and G. sp. aredescribed. The studied material is well·preserved specimens in some calcareous
nodules from the Turonian·Maastrichtian of Hokkaido and Sakhalin. The upper andlower jaws of these species consist of black outer lamella, reduced short innerlamella and covering rostra. The two lamellae are composed of carbonate apatite, andthe rostra are calcareous covering. The lamellae and rostra may have been diageneticallyreplaced from chitin and aragonite respectively, judging from the comparison withmodern cephalopod jaws. The calcareous rostra of the upper and lower jaws areanalogous to rhyncholite (upper jaw) and conchorhynch (lower jaw) of modern
Nautilus instead of coleoids in the internal microstructure, and mineral and chemicalcomposition. These evidences suggest an analogy in feeding habits between tetragonitid
ammonites and Nautilus, although the ammonites have larger and stronger jaws.

of ammonites in Jurassic examples(Lehmann, 1967, 1970, 1971a, 1972, 1976,
1978, 1979; Lehmann & Weitschat, 1973;
Kaiser & Lehmann, 1971). Recently
Zakharov (1979) described the jaw apparatuses of Cretaceous Tetragonites.
Lehmann et at. (1980) described the jaw
apparatuses of Lytoceratacea. On theother hand, rhyncholite and conchorhynch
were interpreted as calcified elements ofnautiloid jaws (d'Orbigny, 1825; Foord,
1891; Rutte, 1962; Teichert et at., 1964;
Gasiorowski, 1973; Shimansky, 1962 ;Miiller, 1974; etc.) or coleoid jaws (Biguet,
1819; Gaillardot, 1824; de Blainville, 1827;etc.), but some were considered to belong
to ammonoids (Schmidt-Effing, 1972;
Mundlos, 1973; Tanabe et at., 1980; etc.).
Almost all cephalopods have jaw apparatuses and radulae. The taxonomicand ecological implications of jaw apparatuses in modern coleoids were studied

Introduction
Anaptychi and aptychi are often preserved in ammonoid body chambers.
They have been interpreted as ammonoid
opercula (e.g. Retowski, 1891; Schmidt,
1928; Trauth, 1927-1936; Nagao, 1931a-c,
1932; Fischer & Fay, 1953; Arkell, 1957;
Schindewolf, 1958) or jaw apparatuses(e.g. Meek & Hayden, 1864; Kennedy &Cobban, 1976). Recently, Closs (1967a, b)
discovered several examples of an aptychus(upper jaw)-radula associations within
the living chambers of an Upper Carboniferous ammonoid, Eoasianites (Ctaphyrites) rionegrensis Closs, and Lehmann(1967, 1971b, 1979) reported in situ aptychi
or anaptychi and radulae in some Jurassic
ammonites. Thus, aptychi and anaptychi
have been confirmed as the lower jaws

* Received October 22, 1980; revised manuscript received November 18, 1981.

239

Yasumitsu KANIE

240

by Clarke (1962), Nixon (1969) and others,
and those in Nautilus by Okutani &Mikami (1977) and Saunders et al. (1978).
Most previous works on ammonoid jaws
have been done mainly on the basis of
the Jurassic materials from Europe.
However, Nagao (1931a-c, 1932) reported
Cretaceous aptychi and anaptychi from
Hokkaido and south Sakhalin, and Trauth(1935) redescribed Nagao's Neoanaptychus.
Kanie et al. (1978) preliminarily reported
the jaw apparatuses of some Late Cretaceous ammonites from Hokkaido and
Sakhalin. They recognized in some specimens that a pair of lower and upper jaws
are preserved in the body chamber; calcareous coverings similar to rhyncholiteand conchorhynch exist in the rostra of

44°'--+-----i

lower and upper jaws; the ammonite
jaws are biomineralogically comparablewith those of extant Nautilus. Furthermore, the tetragonitid jaws were interpreted to be analogous to those of Nautilus
owing to similar feeding habit (Kanie &Tanabe, 1979; Tanabe et al., 1980).
This paper presents a description o.
the general morphology of the teragonitid
ammonite jaws from the Upper Cretaceousof Hokkaido and Sakhalin (Text-fig. 1),
and compares them with the jaw apparatuses of modern Nautilus by some mineralogical analysis. On the basis of theresults obtained, the tetragonitid jaws arerestored.
The depositories of the specimens examined are as follows (abbreviations areshown in parentheses):
Yokosuka City Museum, Yokosuka 238 (YCM)
Department of Geology, Kyushu University,
Fukuoka 812 (GK)
Department of Geology and Mineralogy,
Hokkaido University, Sapporo 060 (UH)
Japanese Expert Consultation on Living
Nautilus. c/o Yokosuka City Museum(JECOLN)
Y. Kawashita's private collection, 2 AI-5,
Tomatsu-Chiyoda-cho, Mikasa 068-22
M. Nihongi's private collection, 8S, 22W, Chuo,
Sapporo 064 (MN)
T. Takahashi's private collection, 28-109,
Hanazono-cho, Mikasa 068-22

General morphology
Comparison with modern cephalopods

Text-fig. 1. Locations of the specimens
studied and exposures of Cretaceous deposits(solid area). N: Naibuchi, K: Tombetsu, R:
Obira, As: Ashibetsu, Ik: Ikushumbetsu, Y:
Oyubari, Kd: Furenai, and U: Urakawa
areas.

Identification of lower and upper jaws.The lower jaw is distinguished from theupper one by several characters (Textfigs. 2, 3), namely, the larger size oflamellae (compared with the height andwidth of living chambers), the broader
lamella angle (angle between left andright sides of the outer lamellae) and less

7 W.

C?"et aceo'Us Tet?"agoniti d Ammonite J aws

u P P E R J A VI

241

L 0 VI E R

J A 11w
y

Vampyromorpha
Va mpyr o t ~ ut his in f~ r n a lishOctopoda

Teuthoidea

Sepioidea
S ~p ia (P ia tys ~p ia) ~ scu (~n to

Text ·fig . 2. Va ri ous ki nds of bas ic mo r pholog ies in mode rn co leo id jaws. Up per jaw of
the Vampyromo r pha cited from Iverso n & Pi nkas (1971) and lower jaw from Clarke (1980) .
Octopoda, T eut hoi dea and Sepioidea cited from Tanabe et al . (1980) . T erminolog ies are
based on Cla rke (1962) . w: wi ng, c: cres t , Iw: latera l wal l, h : hood.

extended hood area (Clarke, 1962 forcoleoids; Oku tani & Mikam i, 1977 andSaunders et al., 1978 fo r Nautilus ).
NJorph%gical comparison with modern
cejJhalojJods.- T he mor pholog ical featuresof the tetragoni t id jaws (Text-figs. 4, 7)
can be correlated w ith t hose of li vin g
Nau tilus rather than w ith those of modern
coleoides.
These amm oni te jaws arecharacteri zed by the presence of rhyncholites and conchorhynchs at the anterior
part of the jaws (T anabe et al., 1980),
T he Nautilus jaw is also covered wi thrhynch olite and con chorhynch (Text-fig.
3; Okutan i & Mikami , 1977; Saunders etal., 1978). Those of col eoids, however,lack calcareous coverin gs v isible w ith the
naked eye (T ext-fig. 2). The lower jawof the Tetragon itidae is morp hologica ll ysimilar to those of Nautilus an d 11ampyroteulhis infernalis in the scar cely extended crest and lateral wall. In the
T etragoni t idae the upper jaw more orless resembles t he lower jaw except forthe more exten ded w in gs cover ing the
reduced short late ra l wa lls an d crest,
whereas in these li ving ce phalopods theupper and lower jaws are morpholog icallyqu ite diffe rent from each other .

Description of jaw ajJparaluses
T he jaw spec imens examin ed are att r ibuted to Caudr)'ceras and T etragonites(Tetragon iti dae : Lytoceratin a). They werepreviously named Neoanapt ychus by NagaoYas~~?nitsu KANJE

242

u P PER
'"o
H

'"W

f-

V)

o

CL

0<

o

H

'"
W
f-

Z

<l:

teral

wall

(VENT RAL)
rostrum

L 0 HE R J AH

'"o
H

'"

W·

f-

Z

<l:

'"o
'"
H

W
f-

V)

o

CL

Text.fig. 3. Basic morphology of mode rn Nautilus jaw. L: length of latera l wa ll ,W : leng t h of wi ng, Hd : length of hood, Hi : heig ht, B: breadt h, IX: wing (la teral wall )ang le, f3: jaw a ngle. Terminologies are most ly based on Okutan i & Mika mi (1977) and
Sau nd ers et ai. (1978) . Solid a rea is covered with a ca lcareous cove rin g.

(1931a) and Anaptychus by Trauth (1935).

Jaws of Gaudryceras s pp.

Family T etragonitidae Hyatt, 1900
Subfamily Gaudrycerat inae Spath, 1927
Genus Gaudryceras Grossouvre, 1894

General characters: The lower andup per jaws are of Anaptychus type. They
are broader than high. Lower jaw islarger and broader than upper one. Both

740. CTetaceous Tetmgonitid A 1nmonite Jaws

243

UPPER JAH
U2
0::

....o
0::
W

......
V")

o

a..

0::

o
....
0::
W

......
Z«(DORSAL)i(VENTRAL)LOHER JAH

0::

....o

0::

w

......
z«

T ex t-fig. 4. Restoration of Caudryceras jaw and t he meas ureme nts. W: lengt h of
w in g , Hi: heig h t, B: breadth , Hd: leng th of hood , IX : w in g ang le, {3 : jaw ang le . Solidarea is co vered by a ca lcareous co ve ring. U1 a nd L3 (Lehman n et al ., 1980).

lower and upper jaws are constructed bywing, hood and ros trum with calcareous
covering and crest plus minor latera l wall

beneath them. Wing a ng le (ex, angle between left and right sides of the wi ngs;
T ext-fig . 7) is w ide in the lower jaw.

Yasumitsu KANIE

244

Jaw angle ([3) is obtuse; the curve between rostrum and wing margin is gentle
without notch. The black lamellae are
made up of apatitic mineral and covered
with a thin calcite layer.
Remar/,s: The specific distinction ofthese jaws in accordance with the classification of Gaudryceras is difficult at
present.
Upper jaw (Pl. 39, Figs. 1-3,5; Text-fig. 4-U)
Previously described material: The following specimens are attributable to the upper
jaws of Caudryceras spp.
1931c. Neoanaptychus sp. form 7, Nagao, p.
216, pl. 15, fig. 6.
1935. Anaptychus
tenuiliratus(Nagao),Trauth, pp. 452-454.
Material: Attributed to Caudryceras denseplicatum (limbo) in situ. GK. HSOS7. Loc.
R4041p2, the upper part of the ObirashibeRiver, Obira area, northwestern Hokkaido.
Lower Santonian (H. Hirano & K. Tanabecoil.) ; to C. tenuiliratum Yabe in situ. UH3191.
Loc. the Ikushumbetsu River about 200 m east
from the junction with the Kikumezawa,
Ikushumbetsu area, central Hokkaido. Santonian(R. Saito coll.); to ? C. tenuiliratum Yabe.
GK. H2262. Loc. N146, the lower part of the
Santan, Naibuchi area, south Sakhalin. Santonian (T. Matsumoto coil.) ; to ? C. tenuiliratum
Yabe. Loc. Kawakami, the Obirashibe River,Obira area. Sahtonian ? (T. Takahashi coil.) ;Table 1.

to ? C. sp. Loc. Y5251a, the lower part of
the Hakkinzawa,
Oyubari area, central
Hokkaido. Turonian (T. Matsumoto, H. Hirano& K. Tanabe coil.) ; to ? C. sp. Loc. R4040b,
the upper part of the Obirashibe River, Obiraarea, northwestern Hokkaido. Upper Coniacian(H. Hirano & K. Tanabe coil.) ; to ? C. sp.
MN6, S3c. Loc. the upper part of the Yubarigoenosawa, Ikushumbetsu area. Coniacian ?
(T. Nihongi coll.); to ? C. sp. (=Neoanaptychus form 7) UH3193. Loc. Kikumezawa,
Ikushumbetsu area. Santonian ? (R. Saito
coil.) ; to ? C. sp. GK. H3220. Loc. Ur512Y4,
the Ikandai Valley, Urakawa area, southerncentral Hokkaido. Lower Campanian (T. Matsumoto coil.) ; to ? C. sp. GK. HSOSS. Loc. K57,
Heitarozawa, Tombetsu area, northern Hokkaido. Upper Maastrichtian (T. Matsumoto, H.
Okada, I. Orita & T. Muramoto coll.).

Description: The wings are triangularand make up the greater part of theupper jaw. Wing surface is marked byconcentric costae and numerous growthlines of variable prominence. The wings
are slightly compressed towards the flankof the living chamber, and combine a hoodat the antero-dorsal part, being narrowly
flat at the dorsal part; growth lines gently
concave toward anterior direction. In
some specimens (e.g., PI. 39, Fig. Ib),
folds or a groove are found; wings posteriorly projected in later growth stage.
Crest and lateral wall are small. Wing

Measurements (in mm) of upper jaws of Caudryceras spp.

Specimen

W

Hd

Hi

B

a

~

UH3193
MN6
GK. HSOS6
R4040b
GK.H3220
GK.HSOS7
GK. H2262
UH3191

56.9
5S.S+
34.7
33.4+
2S.7
23.7
16.5
12.7

3S.4
41. 5
31. 2
30.7
24.3
lS.7
14.9
10.9

37.0
29.2
25.7
27.4+
25.0+
23.1+
lS.S
11. 6

5S.0
40.0
30.9*
27.0*
32.9

60°
49°
51°
65°
4So

14.0
14.4+

47°
sr*

165°
155°
170°
165°
165°
135°
160°
170°

Lh

(Hi/LH)

29.2(0.64)

lS.9(0.61)

W: wing, Hd: hood, Hi: height, B: breadth, a: wing angle, ~: jaw angle, Lh: height ofliving chamber, *: deformed specimen.

740.

Cretaceous Tetragonitid Ammonite Jaws

angle is 47-65°.
The rostrum projects toward the ventral
side and is covered with a calcite layer.
In specimen MN6 (PI. 39, Fig. 1), thepoorly preserved rostral tip fills inside of
the rostrum. The rostrum, originallythickness
sharp-topped, decreases intoward the wings; the outer surface of
the wings is covered with a light-colored,thin (approximately 500 flm thick) calcite
layer. The jaw angle is generally obtuse(135-170"), without a notch (Table. 1).
Remarks: In comparison with Gaudryceras, the upper jaw of Nautilus has lessextended wings, larger lateral wall andnarrower jaw angle (118-119°). The upper
jaw of Tetragonites is clearly distinguishedfrom that of Gaudryceras in the higher,
narrower and more rounded outline.

245

Y1664a, the upper part of the Shiyubari River,Oyubari area, central Hokkaido. Santonian (Y.
Haraguchi coll.); to G. sp. GK. HS04S. Loc.
Kd905, Furenai area, central Hokkaido.
Campanian (T. Matsumoto coll.); to? G.
denseplicatum (Jimbo). YCM. Ur307002. Loc.
U307p, the Ikandai Valley, Urakawa area,
southern-central Hokkaido. Lower Campanian(Y. Kanie colI.) ; to ? G. sp. GK. HS045. Loc.
As1140, the lower part of the Souashibetsu
River, Ashibetsu area, central Hokkaido.
Coniacian (T. Matsumoto coll.).
Description: Wings are triangular andmake up the greater part of the lower
jaw. Wing surface is marked by concentric costae and numerous growth lines.
Wings are slightly compressed toward
the side of living chamber; the hoodcombines wings at the antero-ventral part,being flat, occasionally folded at the ventral
part (Text-fig. 4-L2); growth lines gently
concave toward the aperture. Wing angle
is 80-87". Wings are made up of carbonate apatite in the specimen YCM.
Ur307002.
Lateral wall is small. Thick calcareous
substance (rhyncholite) covers entire lateralwall (Text-fig. 5). Lateral wall continues
to rostrum at the tip of beak. The rostrum
is covered with a calcareous layer forming a beak which consists of an alveolus
structure of 2 mm wide at the dorsal part,though decreasing in thickness to 0.5 mm
toward posterior part of the wing andhood areas (Text-fig. 6-1).
Remarks: The general morphology ofthe lower jaw in Gaudryceras is similar
to that of Nautilus except for the longer
wing, partly flattened hood, and larger

Lower jaw (PI. 39, Figs. 4-6; Text·figs.
4-1, 5, 6-1)
Previously described material: The following specimens are attributable to the lower
jaws of Gaudryceras spp.
1931a. Neoanaptychus tenuiliratum Nagao,
pp. 106-109, fig. 1.
1931c. Neoanaptychus tenuiliratus Nagao, p.
216, pI. 15, fig. 1.
1935. Anaptychus
tenuiliratus(Nagao),Trauth, pp. 452-454.
Material: Attributed to Gaudryceras denseplicatum (Jimbo). GK. HS054. Loc. R2101p, theupper part of the Obirashibe River, Obira area,
northwestern Hokkaido. Lower Turonian (T.
Matsumoto & H. Hirano coll.); to G. tenuiliratum Yabe. GK. HS049. Loc. R2251b, themiddle part of the Kamikinenbetsu River,Obira area. Upper Turonian (T. Matsumoto& H. Hirano colI.) ; to G. sp. GK. HS059. L,)c.

Table 2. Measuremenls (in mm) of lower jaw of Gaudrycers denseplicatum (?).
Specimen

w

Hd

Hi

B

a

YCM. ur
307002

37.S

25.0+

34.0*

51.0+

SO°

Symbols as in Table 1. * deformed specimen.

Lh
140°

(Hi/Lh)Yasumitsu KANIE

246

T ext-fig. 5. Crest and lateral walls tructure (a rrow) of the lower jaw probabl yattributable to Gaudry ceras sp., GK . H8045,
from loco As1l40, the lower part of theSouashibetsu River , Ashibetsu area . Coniacian . x 2.3. Photo by K . Tanabe .

broader than high. It is slightly larger
than upper jaw. The breadth of upper
jaw is approximately equal to or smaller
than height. The jaw plate consists ofouter lamellae (wing and hood) and innerlamellae (minor crest plus lateral wall).
Wing angle (a ) is wide in lower jaw.
Jaw angle ((3) is obtuse to rectangular;
the curve between rostrum and wing
margin is gentle without notch. The
wing is mainly composed of apatitic
mineral with a thin coverin g of calcite
layer.
Upper jaw (PI. 40, Fig s. 1-3 ; T ext -fig . 7-U)
Ma terial: AttributedtoT etrago ni tes
g labrus Oimbo) in sit u. GK. H2073 . Loc . N143a,
the lo we r part of th e San tan, Naibu chi area,
south Sakhalin. Lower Santonian (T. Matsumoto colI. ) ; to T . gl abrus Oimbo) in situ.
GK423 . Loc . Sakhalin . Upper Cretaceo us (collec tor unknow n) ; to ? T. sp. Loc. t he low erpart of th e Kaneobe tsu, Oyubari a rea , central
Hokkaido . Turonian (Y. Kaw as hi ta coll. ) .

structureofT ext-fig. 6. Alveolus
Gaudryce ras
denseplicatum(1) , YCM .
Ur307002 , from loc o U307p , th e Ikandai. Valle y, Urakawa area . Lower Campanian(Tanabe et al ., 1980) and Na u tilus macromphal us (2), jECOLN .M5, from Noum ea , New
Ca ledonia . Both are th e lower jaws (dorsal
v iews) . Photos by Kanie .

jaw angle. The lower jaw of T etragonites
is clearly distinguished from that ofGaudryceras in the greater height, narrower wing angle, and larger jaw angle.
Subfamily Tetragonitinae Hyatt, 1900
Genus T etrag onites KQssmat, 1895
Jaws of T etrag onites glabrus(jimbo, 1894)
Generalcharacters :

Lower

jaw

is

Description: The wings are tri angularand make up the greater part of theupper jaw. Wing surface is marked byconcentric costae and numerous growthlines. Wings are slightly compressed
toward the side of living chamber; thehood is flat or waved in dorsal part;
growth lines concave toward the anterior
part. Crest and lateral wall unclear.
Wing angle is approximately 55 °.
The rostrum is sharply projected toward
the ventral side and covered with a thickcalcareous layer (rhyncholite), decreasing
the thickness toward the win g ; outersurface of wings is covered with a thin(ca. 500/-lm thick) calcite layer (PI. 40,
Fig. 2). The jaw angle is approximately
55° in Kawashita's specimen, wi thout anotch (Table 3).
R emm-ks : The lateral wall and crestof T etrag'onites are reduced to a short

740.

C1"etaceous Tetragonitid Ammonite Jaws

247

u P PER
0<

o>-<
0<
W

.....
o
V)

c..

0<

o

>-<
0<
W

.....
Z
«

(DORSAL)(VENTRAL)

LOl1ER JAH

0< .

o

>-<
0<
W

.....
o
c..
V)

Text·fig . 7. Restoration of Tetragonites jaw and the me3.surements.
show n in text·figure 4. Ul a nd L3 (Lehmann et al., 1980) .

inner lamella as compared with tho3e ofthe upper jaw of Nautilus.
Lower jaw (PI. 40, Figs. 3-5; Text ·fig.7-L)
Previously described material : The follow ·Legends areing specim ens are attr ib utab le to th " lower
jaws of Tetragonites glabrlls .
1931a . Neoanaptychlls tenuiliratll1n Nagao, pp .
106- 109, fig . 2.
1931c. Neoanaptychus sp. form 5, Nagao, p.
216, 'PI. 15, fig . 4.

Yasumitsu KANlE

248

Measurements (in mm) of upper jaw of Tetragonites glabrus.

Table 3.

W

Hd

B

Hi

a

{3

32.3

27.9

20.0*

28.0

55°

100°

17.3+

13.8+

Specimen
Kawashita's
colI.
GK423

Symbols as in Table 1.

GK. H8043
GK. H8049
UH3194
MN83a
GK. H2073
UMUT.MM
6999(Hi/Lh)38**(0.48)* deformed specimen; ** restored.

Measurements (in mm) of lower jaws of Tetragonites glabrus.

Table 4.
Specimen

Lh

W

Hd

B

Hi

a

28.9
26.2
24.6
16.6
10.0

23.2
19.3
10.4
10.3

18.9*
20.0+
13.4
11. 0*
7.2*

23.0*
30.9*
30.5+
27.0+
25.0*

85°
95°
95°
70°

14.6

13.4

7.8

21. 0

90°

Symbols as in Table 1.

{3

Lh(Hi/Lh)37**(0.36*)

135°
115°
100°
150°
110°

* deformed specimen; ** restored.

1931c. Neoanaptychus sp. form 1, Nagao, p.
216, pI. 15, fig. 2.
1931c. Neoanaptychus sp. form 4, Nagao, p.
216, pI. 15, fig. 3.
1935. Anaptychus
tenuiliratus(Nagao),Trauth, pp. 452-454, pI. 14, fig. 6.
1935. Anaptychus
tenuiliratus var. lata
Trauth, pp. 455-456, pI. 14, figs. 8, 9.
1979. Anaptychus sakhalinensis Yu. Zakharov,
p. 87, pI. 20, fig. 2.
Material: Attributed
toTetragonites
glabrus (Jimbo). GK. H8049. Loc. R2251b, themiddle part of the Kamikinenbetsu River,Obira area, northwestern Hokkaido. Upper
Turonian (H. Hirano & K. Tanabe colI.) ; toT. glabrus (Jimbo) in situ. GK. H2073. Loc.
N143a, the lower part of the Santan, Naibuchi
area, south Sakhalin. Santonian (T. Matsumoto
colI.) ; T. glabrus (Jimbo) in situ. GK. H8043.
Loc. N382, the luhachirinpan-Ichinosawa,
Naibuchi area. Santonian (T. Matsumoto colI.) ;to ? T. sp. MN83a. Loc. Takishita-Nakakinenbetsuzawa, the middle part of the ObirashibeRiver, Obira area. Upper Turonian (M. NihongicolI.) ; to ? T. sp. (=Neoanaptychus form 5)
UH3194. Loc. Kikumezawa, just above thejunction with the Ikushumbetsu River, Ikushumbetsu area, central Hokkaido. Santonian(R. Saito colI.) ; to ? T. sp. UMUT. MM6999.
Loc. Na & H-Cr738, the second eastern tributaryof the Naibuchi River, Naibuchi area. Santonian(M. Kawada coll.).

Description: The wings are quadrate
to triangular and make up the greaterpart of the lower jaw. Wing surface ismarked by concentric costae and numerous
growth lines. Wings are gently compressed toward the side of living chamber;the hood is fiat and sometimes folded atthe ventral part (Nagao, 1931c, pI. 15, fig_4); growth lines gently concave toward
the anterior part. Wing angle is 90-95°(Table 4). Crest and lateral wall aresmall, existing as inner lamella. Thincalcite layer covers the greater part ofouter surface of the wings. Wings arecontinuous to the rostrum at the tip ofthe beak (PI. 40, Fig. 5c). The rostrum
is sharp-topped, but dentic1es as cuttingedge are not preserved; rostrum is covered
with a calcite layer, the thickness of which
decreases toward the posterior part ofwings.

740.
Table 5.

Cretaceous Tetragonitid Ammonite Jaws

249

Measurements (in mm) of jaw appoffaratuses of adult Nautilus macromphalus.
The specimens are from of Noumea, New Caledonia.

Specimen

W (L)

Hd

B

Hi

a

f3

up. (31. 3)
JECOLN.
M4
low. 28.9

19.0

17.0

25.3

33°

119°

19.5

33.0

24.5

52°

102°

up. (34.0)
JECOLN.
M5
low. 29.0

21. 5

27.2

25.4

31°

118°

Lh

(Hi/Lh)(0.27)

94(0.26)(0.27)
93

18.3

30.6

23.7

51°

102°

(0.26)

W: length of wing (lower jaw), L: length of lateral wall (upper jaw), Hd: length ofhood, B: breadth, Hi: height, a: wing (lower jaw) or lateral wall (upper jaw) angle, [3:jaw angle, Lh: height of living chamber, as illustrated in Text-figure 3.
Table 6.

Comparison of jaw size of tetragonitid ammonites and modern Nautilus.

Genus

Upper

Lower

Gaudrycaras

Hi/Lh 0.64-.61(2.4-2.3)
W/Lh 0.81-.67(2.3-2.0)
0.48(1. 8)
Hi/Lh
0.27 (0.8)
W/Lh
0.27
(1. 0)Hi/Lh
WL/Lh 0.36-.33 (1. 0)

0.47 (1. 5-1. 3)
0.19*(-)
0.26 (1. 0)
0.36-.31 (1. 0)

Tetragonites
Nautilus

Hi: height, W: length of wing, L: length of lateral wall, Lh: length of living chamber.
The number in parentheses is the ratio with that of Nautilus. *: deformed specimen.

Remarks: The general outline of thelower jaw of Tetragonites is closely
similar to that of Nautilus except for the
narrower wing angle (51-52°) and smaller
size (Table 5).
Discussion: The upper jaw plates oftetragonitid ammonites are morphologicallymore similar to those of octo pods thandecapods, and they resemble those of
Nautilus more closely than octo pods (Textfigs. 2, 3). Namely, the crest and lateralwall of the lower jaw of Nautilus areshortly reduced as in tetragonitid ammonites. Meanwhile, the upper jaw ofmodern Nautilus consists of wide innerlamellae (lateral wall and crest) and shortly
reduced outer lamellae (hood and wing).
The outer and inner lamellae are morphologically similar between the upperand lower jaws in tetragonitid ammonites.
The ratio of jaw height to living chamberheight is less than 100 percent; for ex-

ample, the upper jaw of Gaudryceras is61-64%; the upper jaw of Tetragonites is
48% and lower jow 47% ; the upper jaw
of Nautilus macromphalus is 27% andlower jaw 26% (see Tables 5, 6).
Table 6 gives the size of the jaw apparatuses of tetragonitid ammonites inrelation to the living chambers. The sizeof the jaw apparatuses in Gaudryceras is
1.7-2.4 times and in Tetragonites is 1.31.8 times as large as that of adult Nautilus
macromphalus.
Sharp denticles are present on thealveolus structure in living Nautilus, but
unknown in the Tetragonitidae. However,it may be due to the matter of fossilization (Text-fig. 6-2).
Mineral composition

The mineral composition of the jaw
plates of tetragonitid ammonites wasYasumits~~ J{A N IE

250

10'

20

JO

so

·: 0

60

2

20

JO

"

"

60

70'

Text-fig. 8. X-ra y powder diffraction
patterns of the lo wer jaw attributed to
Gaudryceras dens eplicatum. 1. Carbonate
apatite, 2. calcite. The specimen examined
is preserved in a calcareous nodule, from loco
U307p, Ikandai Va lle y, Urakawa area, south-e rn-central Hokkaido.
All pattens are
obtained by Cu / Ni radiation. Chart speed
20 mm / min. Scanning speed 2° / min. Analyzed by A. Kato and S. Matsubara.

analyzed by an X-ray powder diffraction
method. The patterns are compared withthose of modern coleoids by the same
method. The specimens examined are
the lower jaws of Gaudrycems sp., T etragonites glabrus, Nautilus macromphalus
and Sepia sp. It was found that thecalcareous covering of the rostrum in
Gaudryceras is made up of calcite (Textfig. 8-2), whereas the black outer lamella(wing) is composed mainly of well-crystallized carbonate apatite (Text-fig. 8- 1).
The whole surface of the outer lamellae
in Gaudrycems (PI. 39, Fig. lb) and T etragonites (PI. 40, Fig. 2) is covered witha calcite layer about 500 flm thicl<. The
layers grades into the calcareous coveringof the rostrum. The lower jaw plate of
N autilus consists of outer and innerl·amellae, calcareous rostrum, and denticles.
The former two are made up of chitin(Saunders et al. , 1978), but the X-ray
powder diffraction studies show that
almost all materials display broad bands
near d =4.4 A only as represented by
Text-figure 9-2, also that of Sepia (Text-

4

5
JOO

1<1'

20

JO

"

°c , 30 I':I ln . heated

"

"

,<1'

T ex t-fig . 9. X-ra y powder diffraction
patterns of th e lower jaws attr:buted to two
kinds of modern cep halopods . 1, 2. Nearly
amorphous materials with broad bands abou t
d =4.4 A only , 3. calcite, '1. aragonite, 5.
ara gonite and calcite (heated by Y. Fukuda).
Sepia sp. from Mexico Bay. Nautilus macromphalus from Noumea, New Caledonia.
Patterns were taken under th e same recording conditions as those in Text-fig ure 8.

fig. 9- 1). The calcareous covering of therostrum is composed of aragonite (Textfig. 9- 4), but the denticles are of calcite(Text-fig. 9- 3). Another test showed thata pair of the aragonitic rostra changed
into calcite by heating at 300 °C in half
an hour (Text-fig. 9- 5). The w ing surfaceof the jaw of Nautilus macromphalus
shows similarities with that of T etmgonites, a prismatic layer covers the outersurface of the wings of N autilus 111acrom-

74C.

Cretaceous Tetragonitid Ammonite Jaws

phalus (Fukuda, 1978).
Discussion: The wing surface of theupper and lower jaws of tetragonitid
ammonites are covered with a calcite
layer, approximately 500 fJ-m in thickness.
HOlder (1958) reported that a thin calcareous layer covers the outer surface ofa Jurassic anaptychus, probably attributable to Psiloceras torus d'Orbigny. Recently, Zakharov (1979) described thecalcareous covering, 100-200 fJ-m thick, on
the lamella of the lower jaw in Tetragonites popetellsis Yabe. The outer lamellar
surface (wing) of the lower jaw of modern
Nautilus is underlain by an aragonite
layer, 50-75 fJ-m thick (Fukuda, personalcomm.). Its thickness is about 1/10 ofthat of tetragonitids. The aragonite layer
covers the inner side of broadly extended
lamellae. Therefore, the upper jaw ofmodern Nautilus is mostly occupied by the
inner lamellae (crest and lateral wall). The
outer lamellae of the upper jaw in modern
cephalopods are generally reduced inlength. Both the upper and lower jawsof tetragonitid ammonites are characterizedby similar morphology. The calcified denticles of the tetragonitid jaws were originally made of calcite as in Nautilus, therostrum and the outer surface of wings
consisted of aragonite, and the wings weremade up of chitin. During the course offossilization, the rostrum and outer surface
may have changed into calcite and thewings into carbonate apatite. This suggests that aragonite has changed into
calcite during diagenesis. Such a conversion of aragonite into calcite is also indicated in the septa of Subptychoceras sp.
(specimen YCM. Ur307001) preserved in
the same nodule.

Restoration of jaws

The restoration of tetragonitid ammonite
jaws is undertaken on the basis of the251

above mentioned observations and someanatomical studies of living Nautilus.
Saunders et al. (1978) discussed thefunction of the calcareous coverings ofthe rostral part, rhyncholite (upper jaw)
and conchorhynch (lower jaw). in modern
Nautilus and the extinct Nautilida. Tanabe
et al. (1980) clarified some of the previously known fossil rhyncholites andconchorhynchs belong to such Cretaceousammonites as Gaudryceras, Tetragollites,
Zelalldites and Neophylloceras. Morphologically the tetragonitid ammonite jaws
are most similar to those of living Nautilus. However, the analogous internal
structure is observed between upper andlower jaws of the ammonites in contrast
to the great difference in Nautilus. The
outer lamella (wings) of the upper jaws
of Nautilus is reduced in length and theinner lamella (lateral wall) extends broadly.
The lower jaw of modern coleoids iscomposed of lateral wall instead of wing.
Text-fig. 10 shows the articulation
system of the upper and lower jaws of
Nautilus pompilius in a buccal mass inthe anterior part of the soft body. The
calcareous covering, namely rhyncholite(rh) and conchorhynch (co), is shown as
solid on soft X-ray photographs (Text-fig.
10), whereas the lamellae mostly made ofchitin are faintly shadowed. Text-fig.
10-2 shows the articulated upper andlower jaws of Nautilus macromphalus.
The muscle system between the upperand lower jaws of the present two speciesof Nautilus is designed in Text-figure 11
on the basis of data above mentioned andsome anatomical studies. Namely, thewing and lateral wall are distinguishedfrom the attachment muscle system on
the lamellae and the existence of the calcareous covering; the outer surface of
the wing is covered with a calcareous
layer and the muscle attaches inside; theinner surface of the lateral wall is overlain

252

Yasumitsu KANIE
I

1.
I
/
/
/
/

-------....-,- ./Text-fig. 10. Th e art iculation of the upper and lowe r ja ws of modern Nautilus . rh :rhyncho li te, co: co nc hor hy nch. Photos by Kani e. 1. Soft X-ray photog rap h of the jaw ap paratus of Nau tilu s l)o11lpilius L inn aeus (nat ura l size). Specime n ] ECOLN. P 2 ~ . Loc. - 150
m, off Bind oy, Negros Or ie nt al, Phi lippines. Condition: 30 seconds fo r an integ ral time, 40
kV-6 rnA fo r a n e ne rgy leve l pe r channel w ith Sof tex·EMB. 2. Latera l view of an a rti cu la ted
jaw of Nautilu s 11lacro11lphalus Sowerby (natural size). Specimen ] ECOLN. M5 ~ . Loc. off
Noumea, New Caledonia.

by a calcareous layer and under lies themuscle attachment.
It is suggested t ha t the articulation andmuscle system of both upper and lower
jaws of tetragonitids are quite different
from those of li ving cephalopods. T he
attachment areas of the muscles of tetragon itids are inferred from the morpho-

logica l feat ures of the muscle-scar, though
there is no informat ion on t he jaw musclescar. The muscle a ttachments of thelower jaw in tetragonitids might be insideof the w ing (Text-fig. 12), because of theexistence of a calcareous layer on the
w in g surface. The muscle of the upper
jaw also attaches inside of the wings

740.

Cretnceous Tetrngoniticl Ammonite Jnws

253

Text·fig.11. A nter ior view of articu lated
upper and lower jaws of mod ern Nautilus
species with jaw musc les (horizontal lin es) .
U: upp er jaw, L: lower jaw.

Bf1p----::....

----

Text-fig . 13 . Resto red jaw apparatus of
tetragonitid ammonite w ithin a buccal mass.
The apparatus is covered with buccal mem brance and papi ll ae (BM P) . Discussed w ith
Y. Fukuda and K. Tanabe.

Text-fig. 12 . A nter ior view of articulated
upper and lower jaws of a tetrago nitid am monite . Horizontal lines show the pres um ed
jaw muscu lar system. U: upper ja w, L:
lower jaw. Disc ussed with Y . Fukuda andK. Tanabe.

based on the s imilar lamellar structure.
Therefore, the buccal mass system andmorphological feature of the upper andlower jaws suggest an articu lat ion systemof the tetragon itid ammonite jaws asshown in Text-figure 13.
The feeding habits of tetragonitid am monites which have a conchorhynch at
the rostral part of the lower jaw arepart ly inferred from those of Nautilus
with simi lar conchorhynch. Dean (1901),
Bidder (1962), Haven (1972), Mikami &Okutani (1977), Kanie & Tanabe (1979),

and others observed the feeding habits ofliving Nautilus. In their views, jYautilus
mostly scavenges an imals on the sea floor
and rarely feed on slow moving benthos.
The function of the rhyncholites andconchorhynchs to bite and cut up a preyhas already demonstrated by T anabe etal. (1980) through the observat ions of crop
content remains. Naut ilus is a slow moving an ima l, its maximum swimming speed
being 25 em /sec (Ward et al., 1977), andusually 2.8 em / sec (Mikami et al. , 1980).
The tetragon itid animals with similarshells to Nautilus are probably also slow
swimmers.

Conclusions

The general morpho logy of the jaw
apparatuses of Late Cretaceous Tetragonites and Caudryceras is described .
They are of anaptychus-type, and are

254

Yasumitsu KANIE

often preserved in situ within the bodychambers. The upper and lower jaws
are distinguished from each other by differences of morphology, size, internal
structure and others. The tetragonitid
ammonite jaws are more similar to thoseof modern Nautilus than to those ofcoleoids especially in the existence of
calcareous coverings at the rostral part.
The jaw lamellae of tetragonitid ammonites are composed of carbonate apatite,and the rostra grade into wing surface
layer consisting of calcite, whereas thelamellae of modern cephalopods consist ofchitin. The rostra and denticles of Nautilus jaws are made up of aragonite andcalcite, respectively. The outer lamellaeof modern Nautilus are overlain by verythin aragonite layers. Therefore, theammonite jaws originally seem to havebeen analogous to modern cephalopod jaws
in regard to mineral composition and internal structure.
Based on the similarities of jaws andshell morphology, tetragonitid ammonites
are interpreted to be analogous to Nautilus
in living habits. The jaw apparatuses oftetragonitid ammonites are relatively large
and probably stronger in comparison withthose of Nautilus.
Acknowledgements. I am deeply indebted
to Dr. K. Tanabe, Mr. Y. Fukuda, Dr. I,
Obata, Dr. H. Hirano and Dr. T. Okutani
for instructive discussion and advice on
the cephal oped jaws. I thank Prof. Dr.
U. Lehmann and Prof. C. Teichert for
helpful criticism on the manuscript, Prof.
Emeritus T. Matsumoto and the membersof the Japanese Expert Consultation on
Living Nautilus (JECOLN) for constantencouragement, and Dr. A. Kato and Mr.
S. Matsubara for the X-ray powder diffraction studies of the specimens, The
work was helped by Dr. A. C. Alcala,Messrs. M. Nihongi, T. Takahashi and Y.

Kawashita, and Drs. M. Kato, N. Minouraand H. Okamoto. This study was supported in part by Grant-in-Aid from the
Ministry of Education, Science and Culture(Kanie, no. 264263 for 1977; Takayanagi,
no. 334043 for Cooperative Research in
1978-1980). Contribution 53 of the JECOLN.

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Yasumitsu KANIE

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Explanation of Plate 39
Figs. 1-6. Jaw apparatuses of Caudryceras species.
la, b. Upper jaw probably attributed to Caudryceras sp. a: ventral, b: lateral views. xL
MN6. Loc. the upper part of the Yubarigoenosawa, the Ikushumbetsu River, Ikushumbetsu
area, central Hokkaido. Upper Yezo Group (Coniacian ?). M. Nihongi coil. (Kanie et al.,1978) 2. Lateral view of the upper jaw attributed to Caudryceras sp. xL UH3193. Loc.
the Kikumezawa, the Ikushmbetsu River, Ikushumbetsu area. Upper Yezo Group (Santonian?). R. Saito coil. 3. Mode of occurrence of the upper jaw in Caudryceras denseplicatum(Jimbo). xL GK. H8087. Loc. R4041p2, the upper part of Obirashibe River, Obira area,
nortwestern Hokkaido. Upper Yezo Group (Lower Santonian). H. Hirano and K. Tanabecoil. 4. Upper view of the deformed lower jaw attributed to Caudryceras sp. x 2. GK. H8059.
Loc. the upper part of the Kamikinenbetsu River, Obira area. Middle Yezo Group (Upper
Turonian). T. Matsumoto coil. 5a-c. Upper jaw of Caudryceras tenuiliratum Yabe. a: mode ofoccurrence, b: lateral, c: anterior views. a (x 1), b, c (x 2). UH3191. Loc. the Ikushumbetsu
River, 200 m east from the junction with the Kikumezawa, Ikushumbetsu area. Upper Yezo
Group (Santonian). R. Saito coil. (Nagao, 1931a, c; Tanabe et al., 1980). 6a, b. Lower jaw
probably attributed to Caudryceras denseplicatum (Jimbo). a: dorsal, b: lateral views, co:conchorhynch. x 1 YCM. Ur307002. Loc. U307p, Ikandai Va11ey, Urakawa area, southerncentral Hokkaido. Upper Yezo Group (Lower Campanian). Y. Kanie co11. (Kanie et al.,
1978; Lehmann et al., 1980)
Photos, without whitening, by Kanie (1-3, 5, 6) and K. Tanabe (4).

KAN IE:

Plate 39

Cretaceous tetragonitid ammonite Jaws

1b

2

3

4

60

50

5c

6b

740.

Cretaceous Tetragonitid Arnrnonite Jaws

S. 49-55, Taf. II.
Shimanskii, V. N. (1962): Subclass Ectocochlia(in Russian; English ed., 1974). In Orlov,
Yu. A. Chief ed. Fundamentals of Palaeontology,
vol.
5,
p. 18-32. Izdatel'Academii Nauk.
Tanabe, K., Fukuda, Y., Kanie, Y. and Lehmann, U. (1980): Rhyncholites and conchorhynchs as calcified jaw elements insome Cretaceous ammonites.
Lethaia,vol. 13, no. 2, p. 155-166.
- - , Hirano, H. and Kanie, Y. (1980): The
jaw apparatus of Scalarites mihoensis, alate Cretaceous ammonite. Prof. S. Kanno
Mem. Vol., p. 141-147, pI. 20.
Teichert, C., Moore, R. C. and Zeller, D. E. N.
(1964): Rhyncholites. In Moore, R. C. ed.
Treatise on Invertebrate Paleontology, p.
K567-584. Geol. Soc. Amer. & Univ. Kansas
Press.
Trauth, F. (1927-1936): Aptychenstudien. I.
Ann. Naturhist. Mus. TVien, Bd. 41, S.

257

171-259 (1927); II. Bd. 42, S. 121-193, Taf.
3-5 (1928) ; III-V. Bd. 44, S. 329-411, Taf.
3-5 (1930); VI-VII. Bd. 45, S. 17-136, Taf.
1 (1932); VIII. Bd. 47, S. 127-145, Taf. 3(1936) .
- - (1935):
Anaptychi und anaptychusahnliche Aptychi der Kreide. N. Jb.
Mineral. Ceol. Palaeont., Abt. B, Bd. 74,
S. 448-468, Taf. 14.
Ward, P., Stone, R., Westermann, G. andMartin, A. (1977):
Notes on animal
weight, cameral fluids, swimming speed,color polymorphism of the cephalopod
Nautilus pompilius in the Fiji Islands.
Paleobiology, vol. 3, no. 4, p. 377-388.
Zakharov, Yu. D. (1979): New finds of rhyncholites, anaptychi, aptychi, and remainsof cephalopod radula at USSR territory(in Russian). In Gramm, M. N. ed. Fossil
Invertebrates of Far East, p. 80-91, 99100, pis. 19-20. Vladiostok.

Ashibetsu p~lj, Furenai ~i*J, Hakkinzawa g1i2i,R, Ikandai *~i1, Ikushumbetsu ~~~Ij,
Juhachirinpan-Ichinosawa 18tt.fLE-O)iR, Kamikinenbetsu l::*t~lJlj, Kaneobetsu ~~lJlj, Kikumezawa ~ITiiiR, Naibuchi i*JilHL Nakakinenbetsu qo*t~~Ij, Obira 'PP:, Obirashibe IJ\f~,Oyubari :;!;:!7ryR, Santan =:ft, Shiyubari ::t!75f, Souashibetsu ~p~lj, Takishita i1i1T, Tombetsu ij{lJlj, Urakawa lmi"J, Yubarigoenosawa !7ijft~O)iR
m.~jin'@1m~lHlif e 0) j:t~liIf7t:~:::. J: 0 T r 7 ::l.=. T A H Y :/ -'£- -r 1 r O)';'[i~i} : ~tiffiJ1J • :f'1ii:t:nP.=. Y :/~"? - A r Y 1:: '/ Y :/1lii'ili':O)E)J(J'{ /:; "- - It-- 'I' ,:::. f* ff- ~t:.
Tetragonites glabrus, Caudryceras denseplicatum, C. tenuiliratum, C. sp. 0)1rii*0)7f~
1~ e iil~mJlX:>2:"iitmx: L t:.o :::.
f? O)~i*':t, :71-7 -" 7 • bt "Ht L t:. i*J 7 -" 7 e E)J(J'{O)~:lf$J:I)fllfJJlt~n-n'Qc Ji1I:'jtj1:t:n-:ti:t-1 rY/{!J1 r, It.i::'jtjI:t:nIt---lT11-J:I)t.C I), m~jij'{0) EI!lli*t+ "- -n,@mO)m~eft~.~~Q:::,e~J:~~,~n~n~+:/eY7~-r1rJ:~~.~nke~;tf?n0 0 E)J(J'{~:If'il!lI:t, i*J'il!ltmm·iil~, 1t~mJlX:il'f?, m~;t'7.b.:n"1mO)Y :/:171
I- (l::'lJJD·::J:/:1 Y :/!J (T~JD ~:::.J!ili(bL~ 0 0 :::. n f? O)~*,H:::.£--:)t,~, T r 7 ::J".=. T A f-'r
Y:/-c-r1 rO)i\'iilife lJ:EjZO)-)if,>2:"!Jl5CLt:. o :::.nf?O)y:/-,£--r1 rO)'g:n{,:~':t, ;t'7.L\:n'1m
J: l)iJ;A.,;t-C', tt5fgf,0):;!;:~~~:::.M~0i\'iil!i0):;!;:~~0)j:t':tJ: ~:;!;:~t'o :::'O)JJI~':t, T r 7:1"'=' T Af;'fY:/-'£- -r 1 r iJ;;t '7 l,,:f.l"1 me mfbLO)1t~$>2:" L ~t,t:.:::' e >2:"~1Jtr-r00

MLCHlH't258

Yasumitsu KANIE

Explanation of Plate 40
Figs. 1-5. Jaw apparatuses of Tetragonites glabrus (Jimbo).
la, b. Mode of occurrence (a) and lateral views (b) of upper jaw. a(x1), b(x2). GK423.
Loc. Sakhalin. Horizon unknown (Upper Cretaceous). collector unknown. 2. Lateral viewof upper jaw; white calcareous covering is observable. x 1.5. Loc. the lower part of the
Kaneobetsu, Oyubari area, central Hokkaido. Middle Yezo Group (Turonian). Y. Kawashita
coIl. (Tanabe et al., 1980). 3a, b. Mode of occurrence of lower and upper jaws (a), dorsal
view of lower jaw (b). Ij: lower jaw, uj: rostral part of upper jaw. a(x3/4), b(x2).
GK. H2073. Loc. N143a, the lower part of the San tan, Naibuchi area, south Sakhalin. Upper
part of the Miho Group (Lower Santonian). T. Matsumoto coIl. (Kanie et at., 1978; Tanabe
et al., 1980; Lehmann et at., 1980). 4a, b. Lateral (a) and deformed dorsal (b) views of thelower jaw. x 2. GK. H8049. Loc. R2251b, the middle part of the Nakakinenbetsu River,Obira area, northwestern Hokkaido. Saku Formation (Upper Turonian). H. Hirano and K.
Tanabe coIl. Sa-c. Mode of occurrence (a), lateral (b) and dorsal (c) views of the lower
jaw. a (x 1), b, c (x 2). GK. H8043. Loc. N382, the Juhachirinpan-Ichinosawa, the Naibuchi
River, Naibuchi area. Upper part of the Miho Group (Santonian). T. Matsumoto coIl.
(Tanabe et al., 1980).
Photos, without whitening, by K. Tanabe (lb, 3-5), 1. Obata (la) and Kanie (2).

KANIE:

Cretaceous tetragonitid ammonite jaws

4b50

Plate .405cTrans. Proc. Palaeont. Soc. japan, N. S., No. 125, pp. 259-276, pIs. 41-43, April 15, 1982

741.

EARLY JURASSIC PLANTS IN JAPAN.

PART 4*

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

Abstract. This is the fourth of our serial papers on the early jurassic plants
in japan. In this paper we describe Ctenozamites sarrani belonging to Pteridospermales,and Anomozamites sp. A, Otozamites jujimotoi, O. neiridaniensis sp. nov., Ptero·
phyllum ex gr. propinquum, P. sp. A, Ptilophyllum cfr. cutchense, P. nipponicum
sp. nov., P. shinadaniense sp. nov., P. sp. A. and P. sp. B, all belonging to Ben·
nettitales.
The occurrence of Ctenozamites and Ptilophyllum is the first record not onlyin the early jurassic flora but also in the older Mesozoic floras in japan.

Introduction and acknowledgements

Systematic description

In continuation of our previous papers(Kimura and Tsujii, 1980a, b, '81), we
here describe the fossil plants belonging to the Pteridospermales andBennettitales from the Lower Jurassic
Kuruma Group and Iwamuro Formation.
The details of stratigraphy of plant-beaing
formations and localities were given in
our first paper (Kimura and Tsujii,
1980a).
We express our sincere gratitude toProfessor Emeritus Thomas M. Harris,
F. R. S. of the University of Readingfor his helpful suggestions and [criticalreading over the present manuscript.
We also give our thanks to Mr. Gumpei
Mori, Mr.
Noriyuki Sasaki and Mr.
Akihiro Kobayashi for their kind help III
collecting the fossils here described.

Pteridospermales
Genus Ctenozamites Nathorst 1886: 122
Ctellozamites sarrani (Zeiller) Harris
PI. 41, Fig. 1; Text-fig. 1
Ctenopteris sarrani Zeiller: Zeiller, 1903, p.
53, pIs. 6-7, fig. 1, la; pI. 8, figs. 1-2(Hongay Coal-Field, North Viet-Nam) ; p.
292, pI. 44, figs. 3-4 (Tai-pin-tchang,North Yunnan; corresponding possibly to
the Yipinglang Group) : Sze, 1956, pp. 39,
146, pI. 35, figs. 3, 3a, 4 (Upper Triassic
Yanchang Group, N-China).
Ctenozamites sarrani (Zeiller) Harris: Harris,
1961, p. 170 (name): Sze et a!., 1963, p.
198, pI. 58, fig. 1 (reinserted from Sze, 1956,
pI. 35, fig. 3a) ; pI. 59, figs. 2, 3 (from
Sze, 1956, pI. 35, figs. 3, 4): Li P. C.,
1976, p. 126, pI. 40, figs. 5, 6 (Upper
Triassic Yipinglang Group).
Material: NNW -146,(Nishi-Neiridani) .

846(counterpart)

Description: PI. 41, Fig. 1 shows two
pinna fragments. Leaf-outline and forkingof main rachis are unknown. The pinnae
are more than 14 cm long and 5.2 cm wide,
nearly parallel-sided and with a thick* Received june 13, 1981; Read jan. 23, 1981
at Sendai.

259

260

Tatsuaki KIMURA and Masanori TSUJI!

pinna rachis, 4 mm wide with longitudinalribs on its surface. Segments (or pinnules)
are typically rhomboidal, 2.9 cm long and1.7 cm wide, and arising on the upper sideof pinna rachis at a wide angle. The
apex of segments is obtuse or rounded,
its margins are entire and are flat but
apical margins are sometimes reflexed.
The bases of segments are not contracted.
Veins are 10-11 in number at base, thenforking once or twice, but the marginalones are mostly simple. They are nearlyparallel and end in the outer margin.
Text-fig. 1 shows the form of typicalpinnules and venation. The cuticle is notpreserved.
Distribution and occurrence: In the
Kuruma Group, a single specimen was
obtained from the Negoya Formation.
This species has been known from theUpper Triassic of North Viet-Nam andof North and Middle China.
Remarks: Our leaf corresponds in sizeand outline of segments (pinnules) toCtenozamites sarrani. But it is somewhat
different from Zeiller's original specimensof Ctenopteris sarrani. Differnces are:In the original specimens, the veins are
more crowded and generally forking once,
and the segments arise at reduced angle
and are usually directed forwards, insteadof arising at wide angle in ours.
Our leaf resembles a well-known European species, Ctenozamites cycadea (Berger) Schenk (references, see Harris, 1961,
p. 169; 1964, p. 95). But our leaf differsfrom Ctenozamites cycadea in its largesized segments with denser veins forking
twice, instead of medium-sized segments(2 cm long and 1 cm wide, according toHarris, 1961, '64) with coarser veins(usually 6) forking once-twice in C. cycadea.
Ctenozamites cycadea finely illustrated
by Delle (1967) from the Middle Jurassic
of Tkvarchelian Coal-Basin (Transcau-casia) and Ctenopteris cycadea (Brongniart)
by Yang (1978) from the Upper Triassic
Daqiaode Formation, Sichuan, are alsosimilar in form of segments to ours, butthey are distinguished from ours by theirmedium-sized segments with coarser veinsforking once or twice. Ctenopteris megaphylla described by Yang (1978) fromthe Upper Triassic Daqiaode Formation
resembles Ctenozamites sarrani in form
but smaller.
Ctenozamites usnadzei originally described by Doludenko and Svanidze (1969)
from the Callovian of Georgia and laterby Barnard and Miller (1976) from theMiddle Jurassic of Iran, is distinguishedfrom ours by its rhomboidal segments
typically with a free acroscopic marginand decurrent basiscopic margin, andfurther by its small-sized segments with
1-3 small teeth at apex.
Recently 6 new Ctenozamites species
were described from the Upper Triassicof China by Hsii et al. (1975) and Chen
and Duan (1979a, b). Among them
Ctenozamites otoeis described by Chen
and Duan (1979a) from the Upper Triassicof Yanbian, has segments of similar size
to our specimen of C. sarrani, but they
are longer and have a decurrent base.
The veins are more crowded.
This is the first record of the genus
Ctenozamites in Japan.
Bennettitales

Genus Anomozamites Schimper, 1870: 140
Anomozamites sp.

A

Anomozamites sp.: Kimura, 1959a, p. 22, pI.
8, fig. 3 (Iwamuro).
Material:

A-0558 (a single specimen).

Remarks: Our leaf-fragment resembles
Anomozamites major (Brongniart) Nathorst, and also Pterophyllum pinnatifidum
Harris. Further material is needed for741.

Early Jurassic Plants

specific determination.
Genus Otozamites Braun, 1842
Otozamites jujimotoi Kimura
Text-figs. 2a-b
Otozamites fujimotoi Kimura: Kimura, 1959a,
p. 23, pI. 12, figs. 2-4, 6; text-figs. 12-13([wamuro) .
Material: Lectotype; A-4090 ([wamuro).
Paraiectotypes; A-2037, 2054, 208l. Other
specimens; A-0432, 0489, 0491, 2053, 2064-2066,
2068, 2088, 2091, 4091, 4094. Stratum typicum:
Iwamuro Formation. Locus typicus: Iwamuro,
Shirasawa-mura, Tone-gun, Gumma Prefecture.
Derivatio nominis: After Professor Emeritus
H. Fujimoto of the Tokyo University of
Education.

Emended diagnosis: (Length and apexof leaf unknown.) In middle region rachis
2 mm wide, lamina 6 cm wide but narrower in basal part of leaf. Pinnae variedin form according to their position on aleaf, attached perpendicularly to the upperside of rachis by the lower half of base,thus the upper half free, forming a
distinct angle entirely covering therachis and the lower part of opposite
pinna base. Pinnae mostly elongated but
the proximal ones broadly deltoid or circular in form. Elongated pinnae 3 cmlong or more, nearly parallel-sided, typically 1 cm wide. Deltoid or circularpinnae 8-20 mm long or more and 1014 mm wide, with pointed apex. Circularpinnae with rounded apex restricted tobasal part of a leaf. Veins on the lower
half of pinna mostly simple or once forked
and parallel, but those on free angle radiating, simple or once forked. Concentration of veins about 20 per cm. (Cuticle
not preserved.)
Distribution and occurrence: Otozamites
jujimotoi is occasional in the Iwamuro
Formation, and seems to be local.
Comparison and remarks: Since Kimura

261(1959a) described this species, only a few
additional specimens were obtained. Otozamites jujimotoi is characterized by its
varied pinnae in form, with distinct
acroscopic basal angle.
Otozamites gissaricus illustrated by
Iminov (1976) from the Middle Jurassicof Central Asia, is similar in form toours, but O. gissaricus is a smaller leaf
with narrower pinnae, especially on theproximal part of a leaf.
Specimens determined by Iminov (1976)
from the Upper Triassic of Central Asia
as Otozamites pamiricus Prynada are
distinguished from O. jujimotoi by its
pinnae with rounded or truncated apex.
Otozamites ptilophylloides originally described by Barnard and Miller (1976)
from the Middle Jurassic of Iran is similarin pinna form to O. jujimotoi, but it ismacroscopically distinguished from ours
by its pinnae mostly with rounded apex
and smaller number of veins in eachpinna.
Although we still do not know the formof the whole leaf of Otozamites jujimotoi,
we feel sure it is distinct for its varied
and characteristic pinna forms with distinct acroscopic angle.
Otozamites

neiridaniensis Kimura andTsujii sp. nov.

PI. 42, Figs. 1-5; PI. 43, Figs. 1-2;
Text-figs. 3a-f
Otozamites molinianus Zigno: Oishi, 1940, p.
333, pI. 31, figs. 3, 3a, 4 (Neiridani).
Otozamites sp.: Oishi, 1940, p. 336, pI. 30,
fig. 5 (Neiridani).
Material:
Hoiotype; NEE-095 (HigashiNeiridani). Para types ; NEE-065, 174, TOB013A, B, 050B, 050B', 050C (Tobiiwadani),
NNW -084 (Nishi-Neiridani), NEG-251 (Negoya) . Other specimens; NE-026 and other9 specimens (Neiridani), NEE-007 and other64, NNW -002 and other 57, NEG-006 and other49, DAI-004 and other 38 (Dairagawa), TOB-

262

Tatsuaki KIMURA and Masanori TSUjII

2b'

2b

------...
,,--,

2a

3b "'"
3c

741.

Early Jurassic Plants

004 and other 21.
Stratum typicum: Negoya Formation, Kururna Group. Locus typic us : Higashi-Neiridani,
Asahi-machi,
Shimo-Niikawa-gun, Toyama
Prefecture (see Kimura and Tsujii, 1980a, p.
342, fig. 2). Derivatio nominis: After the
Neiridani where the holotype and many otherspecimens were collected.

Diagnosis: Leaf rather large, long andnarrow, linear; length estimated at 30 cm
or more, width over the middle leaf typically about 2-2.5 cm, occasionally 3.7 cm.
Leaf tapering gradually to both ends.
Rachis rather thin, 1 mm wide, usuallyconcealed. Pinnae oval or short-rectangular in form, typically 1.2 cm long and 0.8
cm wide, with rounded or obtusely pointed
apex, attached on the upper side of rachis
by the lower two-thirds or three-fourthsof their width (accordingly upper onethird or one-fourth of acroscopic basal
part free), at an angle of 60-90 degrees.
Acroscopic basal corner slightly auriculated but seldom covering the pinna on
the opposite side of the rachis. Basiscopic
basal angle slightly contracted or occasionally straight but not decurrent. Near theleaf base, pinnae becoming shorter andmore deltoid but still broad, and near theleaf apex, pinnae becoming narrower.
Veins arising from proximal two-thirds
or three-fourths of pinna base and radiating into the acroscopic corner and therest of pinna, branching at all levels, at

1.

2.

3.

263

a concentration of 15 per cm at base.
(Petiole not known and cuticle not preserved.)
Distribution and occurrence: Otozamites
neiridaniensis is abundant in the Negoya
Formation and rather rare in the Shinadani
Formation. A doubtful leaf-fragment is
known from Neiridani.
Comparison and discussion: Otozamites
neiridaniensis is characterized by its ovalor short-rectangular pinnae with a rounded or obtusely pointed apex. Its
auricle is developed slightly if at all andits veins are crowded.
Oishi (1940) described several leaf-fragments from Neiridani (exact locality notclear) as Otozamites molinianus. But ourobservation of his figures shows thatthey do not belong to Zigno's species
but to Otozalllites neiridalliellsis, because in his pinnae auricles are scarcely
developed.
In Otozalllites 77l0linianus,
auricles are distinctly developed (Zigno,
1883; Moller, 1903). Oishi (1940) also
described a single leaf-fragment as Otozamites sp. from Neiridani. This specimen
agrees possibly with the basal pinnae of
Otozamites neiridaniensis.
Among 160 Otozamites species hitherto
known from the Mesozoic plant-beds, thefollowing species resemble ours in rectangular or triangular pinnae with rounded or obtusely pointed apex, and withnot or only slightly expanded auricle:

Text-figs. 1-3. (All natural size)
Ctenozamites sarrani (Zeiller) Harris; Outline of pinnules and venation (NNW -146;
counterpart of NNW -846 shown in PI. 41, Fig. 1).
Otozamites jujimotoi Kimura. 2a: A leaf-fragment with elongated and auriculated pinnae(A-4091; counterpart of A-4090 shown in Kimura, 1959a, text-fig. 12e). 2b-b"': Deltoid
and auriculate pinnae at the basal part of a leaf CA-2054; redrawn from Kimura, 1959a,
text-fig. 12b, a, d, c respectively).
Otozamites neiridaniensis Kimura and Tsujii sp. nov. 3a: A medium sized leaf (l\EE-095,
holotype; PI. 42, Fig. 1). 3b: Ditto, with apex (TOB-050B, para type) . 3c: A large-sizedleaf (TOB-050B', para type ; PI. 42, Fig. 2). 3d: Apical part of a broken leaf (TOB-013B,
para type). 3e-e': Pinnae, showing the venation (NEE-255). 3f: A large-sized pinnae,
showing the venation (NNW -084, para type ; PI. 43, Fig. 1).

Tatsuaki KIMURA and Masanori TSUJII

264

Otoza71lites falsus Harris: Harris, 1949,'69; Middle Jurassic of Yorkshire. O.
indosinensis Zeiller: Zeiller, 1903; Upper
Triassic of North Viet-Nam. O. kondoi
Oishi: Oishi, 1940; Upper part of Middle
Jurassic MOll(~ Formation, NE-Japan. O.
parallelus Phillips: Harris, 1969; Middle

Jurassic of Yorkshire. O. reglei (Brongniart) Saporta: Brongniart, 1828; Bartholin, 1894; Moller, 1903 (dr. reg lei) ;Lower Jurassic of Bornholm. O. simpsoni
Harris: Harris, 1949 (regarded as O.
feistmanteli Zigno), '69; Middle Jurassicof Yorkshire.

4e

4a

4f

~I--4i

o

2cm
,

4j

Text-fig. 4. Pterophyllum ex gr. propinquum Goeppert; varied leaf-forms and venation.
4a: (NNW -459). 4b: (NNW -666). 4c: (NNW -670). 4d: (NNW -S50). 4e: (NNW -531). 4f:(NNW-743). 4g: (NNW-SID). 4h: (NNW-539). 4i: (NNW-743). 4j: (NNW-356; PI. 43, fig.
5). 4k: (NNW -657). 41: (NNW -326). 4m: (NNW -314). 4n: (NNW -009). (All natural size)

741.

Early Jurassic Plants

These differ as follows: Otozamites
falsus has mostly rhomboid pinnae withan acute apex. O. indosinensis, O. parallelus and O. simpsoni have more elongate
pinnae. Ptilophyllum khargaense (Kilpper,
1966) from the Middle Jurassic (?) of
Egypt has pinnae of rather similar shape
but the lower ones differ in being imbricate. The closest to our species is
Otozamites Iwndoi but its pinnae have anearly symmetrical base and its veins
which are coarser, only fork near their
origin.
Genus Pterophyllum Brongniart, 1828: 25
Pterophyllum jaegeri Brongniart described by Oishi (1931) from Tsuchizawa
is not Pterophyllum but referable to
Nilssonia sp. A to be described later.
Pterophyllum ex gr. propinquum Goeppert
Pl. 41, fig. 2; Pl. 42, figs. 6-7; Pl. 43,
figs. 3-6; Text-figs. 4a-n
SPecimens from East Asia:
Pterophyllum propinquum Goeppert: Oishi,
1931, p. 245, pl. 18, figs. 1-2 (Tsuchizawa,
Kuruma Group) : Yabe and Oishi, 1933, p.
227, pl. 34, fig. 2 (Lower-Middle Jurassicof Liaoning, NE-China): Oishi, 1940, p.
344 (remarks) : Kimura, 1959b, p. 70, pl.
2, fig. 2 (Kotaki Coal·Field, Kuruma
Group): Sze et al., 1963, p. 157, pl. 61,
fig. 8 (reinserted from Yabe and Oishi,
1933, pl. 34, fig. 2).
Pterophyllum aff. propinquum Goeppert: Yabe
and Oishi, 1929, p. 91, pl. 18, fig. 6(Daedong Group, N-Korea).
For Europeaw references, see Antevs, 1919,
p. 28.
Material: NNW-009 and other 32 specimens(Nishi-Neiridani), DAI-027 (Dairagawa), SHI035,040 (Shinadani), Kr-087, 090 (Tsuchizawa),
4962113 (Kotaki Coal·Field), A-7535, 751124(Iwamuro) .

Description: Leaves are varied in sizeand form, but are mostly medium-sized265

and oblanceolate in outline. Leaves taper
gradually towards the proximal end, aremore than 6.5 cm long and 4 cm wide at
the widest portion, and possibly contract
suddenly at the apex. The rachis israther thin, 2 mm wide. The petiole is
not known. The pinnae arise at a wide
angle from the lateral margins of rachis.
They are rectangular, up to 2 cm long
and are shorter towards the base of theleaf. The width of pinnae is variable
but often about 1.2 cm; pinnae near theleaf apex are narrow. Margins of adjacent pinnae are mostly apart but arealways connected to one another near
the rachis by a web of lamina. The
apex of pinnae is truncate in lower partof leaf, but rounded in the middle ofleaf. Veins are parallel and simple oroften fork once. Their concentration istypically 25 per cm at the middle ofpinna. The cuticle is not preserved.
Text-figs. 4a-n show varied pinna forms
and venation.
Distribution and occurrence: This species is locally very abundant in the
Negoya Formation, but rather rare in the
Shinadani and Iwamuro Formations. In
East Asia, this species occurs in theLower-Middle Jurassic of NE-China andthe Upper Triassic (?) of N-Korea.
Remarks: Pterophyllum ex gr. propinquum is characterized by its rectangularpinnae with an expanded base and truncated or rounded apex. Our leaves areindistinguishable from Pterophyllum propinquum illustrated in detail by Antevs(1919) from the Liassic Hor Sandstone,
Sweden, although in our large collection,
leaves with elongated pinnae such as
Antevs' pI. 3, figs. 10-12 have not beenfound.
The leaves regarded by Oishi (1931) as
Pterophyllum propinquum from Tsuchizawa and by Yabe and Oishi (1929) as
P. aff. propinquum from N-Korea areTats~[aki KIMURA and Masanori

266

TsuJI!

6a

6b
5b

5b'

~
a.5cm
'-------'

,
\''''

4cm

....... ---~-

741.

Early Jurassic Plants

indistinguishable from ours, although the
Korean leaf is about twice as large as
ours. A leaf-fragment regarded by Yabe
and Oishi (1933) as Pterophyllum propinguum possibly from the Beipiao Group(Lower-Middle Jurassic), NE-China issimilar in pinna form to ours.
Pterophyllum leaves with similar pinna
form to ours have been described asfollows:
Pterophyllum pinnatifidum Harris: Harris, 1932; Rhaetic of East Greenland. P.
kochi Harris: Harris, 1926, '32; Ditto:
Lundblad, 1950; Rhaeto-Liassic of Sweden.
P. abnorme Etheridge: Walkom, 1917:
Jones and Jersey, 1947; Upper Triassic of
Queensland. P. sensinovianum Heer : Heer,
1876; Jurasso-Cretaceous of the upper
Amur: Vakhrameev and Doludenko, 1961;
Upper Jurassic of Bureja Basin.
They are distinguished from ours as
mentioned below. In Pterophyllum pinnatifidu17l, pinna apices are often acuminate. In Pterophyllw1Z kochi, pinnabases are not contiguous except in theapical part of leaf and are square in the
middle of leaf, and slightly contracted in
the proximal part of leaf. In ours, pinnabases are usually contiguous from one
pinna to the next.
A little known Pterophyllum abnorme
is distinguished from ours by its usually
twice forked veins instead of simple or

5.

6.

267

once forked ones in ours. In Pterophyllum
sensinovianum, its pinnae are more elongated than those of ours.
Pterophyllum propinquum illustrated by
Iminov (1976) from the Upper Triassic
of Pamir is, in our opinion, rather close
to P. tsetzei Schenk or to P. subaequale
Hartz because of its large-sized leaves
with more elongated pinnae.
PterophylluJJl sp.
A illustrated byWeber (1968) from the Rhaeto-Liassic of
Bayreuth resembles P. propinquum. Pterophyllum exhibens originally describedby Li, P. C. (1964) from the Upper
Triassic Xujiahe Formation, Sichuan andlater by Zhou, T. S. (1978) from theUpper Triassic Wen bins han Formation,
Fujian, China, is similar in its expanded
pinna base, but it is distinguished by its
more elongated pinnae with smaller
number of veins (5-12) in each pinna.

Pterophyllu71l sp. A
Pterophyllu11l sp. A and sp. B: Kimura, 1959a,
p. 24, pI. 9, figs. 2-3 (Iwamuro).

Remarks: Kimura (1959a) illustrated
two Pterophyllum leaf-fragments as P.
sp. A and sp. B. Our study convinced
us that they are con specific but the
specimens are too poorly preserved todetermine. Their cuticle is not preserved.

Text·figs. 5-6 (a-f). (All natural size, unless otherwise indicated)
Ptilophyllum cfr. cutchense Morris. 5a: A leaf-fragment, showing the venation (TE-001;
PI. 43, Fig. 7). 5b: Ditto, with apex (TE-003; PI. 43, Fig. 8). 5b': Venation drawn from
5b.
Ptilophyllum nipponicum Kimura and Tsujii sp. nov. 6a: A typical leaf (NEG-720, holotype; PI. 43, Fig. 9). 6a': Venation drawn from 6a. 6b: Lower part of a leaf (NNW -806,
PI. 41, Fig. 3). 6b': Possibly representing the basal part of 6b. 6c: Basal part of a leaf,
bearing broader pinnae with truncated distal margin (NE-067, paratype). 6d: Apical partof a medium-sized leaf bearing rather remotely set pinnae (NNW -353, para type; PI. 41,
Fig. 4). 6d': Venation drawn from 6d. 6e: A leaf-fragment bearing rectangular pinnae(NEE-074; PI. 42, Fig. 8). 6e': Venation drawn from 6e. 6f: A leaf-fragment bearing
broader and remotely set pinnae (NNW -745).

Tatsuaki KIMURA and Masanori TSUJII

268

Genus Ptilophyllum Morris, 1840: 327
About 60 Mesozoic species have been
described. Many of them have similar
pinnae. Thus except the leaves with
distinct morphological features, it is getting difficult to identify them depending
only on their macroscopical forms. Our~91 ib o.scmiknowledge of the epidermis of this genus
owes much to the studies of jacob andjacob (1954), Kilpper (1965, '66), Harris(1941, '42, '46, '49, '53, '69), Doludenko(1963), Bose and Kasat (1972) and Barnard
and Miller (1976).
We recognized five forms in the Lower
jurassic of japan. This is the first record

741.

Early Jurassic Plantsof this genus in the older Mesozoic floras
in Japan. The phytogeographical implication on the occurrence of this genus in269

the Japanese Lower Jurassic will be mentioned in our later paper treating theconcluding chapter of this series.

Key to the Japanese Lower Jurassic species of Ptilophyllum
Leaf large-sized, widest part over 5.4 cm ....................... . P. nipponicum sp. nov.
I'. Leaf medium-sized about 3 cm wide .................................................. 2
I". Leaf small-sized width less than 1 cm ................................................ 3
2. Pinnae elongate-rectangular ..................... " .......... . P. shinadaniense sp. nov.
2'. Pinnae elongate-oval or rectangular ......................................... . P. sp. A
3. Pinnae elongate-triangular ... " ...................................... . P. dr. cutchense
3'. Pinnae rhombic ....... " ...................................................... . P. sp. B
1.

Ptiiophyllum dr. cutchense Morris
PI. 43, Figs. 7-8; Text-figs. 5a-b

Comparable specimens:
Ptilophyllum cutchense Morris: Bose andKasat, 1972, p. 118, pI. 1, figs. 7-10; pI.
2, figs. 14-17 (Upper Jurassic of Rajmahal
Hills and Lower Cretaceous Jabalpur Series,
India) .
Material: TE-OOl, 003 (Teradani, Negoya
Formation) .

Description: Two small-sized leaffragments were obtained. The leaves are
0.8-1 cm wide with an acuminate apex
as shown in PI. 43, Fig. 7. The rachis
which is partially exposed, is about 0.5
mm wide. Pinnae are attached on uppersurface of rachis, closely set or imbricate,mostly attached by entire base, arising

6.

7.

8.
9.

at an angle of about 60-90 degrees. Pinnae
are as a whole linear, elongate-triangular, sometimes rhomboidal, sublate orovate in form; margins are straight.
Apices are usually obtuse or rounded andthe basiscopic basal margin straight,
slightly contracted, rounded or decurrent.
Veins arise from the whole base. They
are parallel, mostly simple, 5 in number
on each pinna. Cuticle is not preserved.
Distribution and occurrence: This form
is rare and known only from Teradani,
Negoya Formation.
Remarks: Our leaves reaves resemble
closely some smaller leaves illustrated by
Bose and Kasat (1972) as Ptilophyllum
cutchense. Ptilophyllum horridum Roy
and P. institacallum Bose are allied forms
to P. cutchense. But both are distinguished from our leaves by their radiating

Text-figs. 6 (g-k) -9. (All natural size, unless otherwise indicated)
Ptilophyllum nipponicum Kimura and Tsujii sp. nov. 6g: A leaf-fragment bearing
elongated pinnae with decurrent base and contracted acroscopic base (NNW -241, paratype). 6g': Vanation drawn from 6g. 6h: Small-sized leaf-fragment (NNW-353, paratype).
6i: Large-sized leaf-fragment bearing broader pinnae with slightly decurrent base (NNW241, paratype). 6j: Apical part of a leaf (NE-169). 6k: Basal part of a leaf with thickpetiole of which base is expanded. Pinnae are small-sized, and with straight or slightly
contracted base (NNW -111, paratype; PI. 43, Fig. 10).
Ptilophyllum shinadaniense Kimura and Tsujii sp. nov. 7a: A part of leaf (SHI-007,
holotype; PI. 41, Fig. 5). 7b: A leaf-fragement, showing the venation (SHI-024, paratype).
7c: Ditto (SHI-019, paratype; PI. 41, Fig. 6).
Ptilophyllum sp. A; a part of leaf (SHI-030; PI. 43, Fig. 11). 8': Venation drawn from 8.
Ptilophyllum sp. B. 9a: A part of leaf (NEG-51; PI. 43, Fig. 12). 9b: Ditto (A-0379).
9b': Venation drawn from 9b.

270

Tatsuaki KIMURA and Masanori TSUJII

and forked veins instead of mostly simple
and parallel ones.
Ptilophyllum cantherifera originally described by Douglas (1969) from the Lower
Cretaceous of Victoria (Australia) is similar in form, but is distinguished by its
pinnae attached by central basal area
instead of by the entire base in ours.
Thus our leaves are most like the smallsized Ptilophyllum cutchense, but we here
refrain from fully identifying them withthis Indian species, because our leaves
without cuticle are only two andincomplete.

Ptilophyllum nipponicum Kimura andTsujii sp. nov.
PI. 41, Figs. 3-4; PI. 42, Fig. 8; PI. 43;
Figs. 9-10; Text·figs. 6a-k
Material: Holotype; NEG-270 (Negoya).
Para types ; NNW -111, 241, 353, 697, 806 (NishiNeiridani), NE-067 (Neiridani). Other specimens; NEG-On and other 3 specimens, NNW004 and other 6, NEE-074 (Higashi-Neiridani),
NE-169, DAI-086 (Dairagawa), IN-OOl, 002(Inumatadani) .
Stratum typicum: Negoya Formation, Kururna Group. Locus typicus: The Negoya, Asahimachi, Shimo-Niikawa-gun, Toyama Prefecture.
Derivatio nominis: After Nippon meaning
Japan in Japanese.

Diagnosis: Leaf large, oblong or obi anceolate in form; the widest part 5.4 cm.
(Whole length and leaf apex unknown.)
Rachis slender 2 mm wide at the middleof leaf but 3 mm wide at the basal part.
Petiole about 5 cm long with striatedsurface, base distinctly expanded. Pinnae
varied in form according to their position
on a leaf, attached on the upper side ofrachis at an angle of 50-90 degrees byentire base, set closely but sometimes
remotely. Middle pinnae elongate-rectangular, typically 3.5-4 cm long and 0.5-1 cm

wide. Apical pinnae similar in form to
those of middle ones but smaller in size.
Basal pinnae typically deltoid in form,
1.5 cm long and 0.5-0.8 cm wide at base,but smaller towards the base of leaf.
Acroscopic basal margin mostly rounded
and basiscopic basal margin straight,
slightly contracted or slightly decurrent.
Apex acutely or obtusely pointed. Veins
numerous, divergent, forking at all levels,
28-30 per cm in density at the middle ofpinna. (Cuticle not preserved.)
Distribution and occurrence: Ptilophyllum nipponicunz is common only in the
Negoya Formation.
Comparison and discussion: It is clearthat our leaves belong to Ptilophyllum
redefined by Harris (1969), because ouracroscopic pinna base is not auriculated
and the basiscopic pinna base is typicallydecurrent_
Ptilophyllum nipponicunz is characterized
by its large-sized leaf with elongatedpinnae above and deltoid pinnae below
with divergent and forked veins, and its
thick petiole and expanded base.
Several Ptilophyllum species with large
leaves have been described. Ptiloph'yllum
nipponicum closely resembles P. hirsutwn
Thomas and Bancroft described in detailby Harris (1949, '69) from the Middle
Jurassic of Yorkshire. But the pinnae of
Ptilophyllum hirsutum do not have adecurrent basal margin.
Ptilophyllum harrisianum described byKilpper (1968) from the Liassic of N-Iran
has pinnae of similar size but with anauriculate acroscopic base. Kilpper's Ptilophyllum zirabense (1968) has more elongated pinnae.
Ptilophyllum shinadaniense Kimura
and Tsujii sp. nov.
PI_ 41, Figs. 5-6; Text-figs. 7a-c
Material:

Holotype; SHI-007 (Shinadani).

741.

Early Jurassic Plants

Para types ; SHI-019, 024.
Stratum typicum:
Shinadani Formation,
Kuruma Group (see Kimura and Tsujii, 1980a,
p. 342, fig. 2). Locus typicus: The Shinadani(near Agero Village), a branch of the Dairagawa, Omi-machi, Nishi-Kubiki-gun, Niigata
Prefecture.
Derivatio nominis: After the
locality, the Shinadani where the holotype
and para types were collected.

Diagnosis: Leaf medium-sized, 3 cm
wide. (Whole length unknown.) Pinnae
ansmg at an angle of 65-80 degrees tothe rachis, elongate-rectangular in form,
typically 1.8 cm long and 3.5-4 mm wide.
Width of pinnae about one-fifth its length.
Pinnae straight or very slightly falcate,apex obtusely pointed, basiscopic basal
angle normally decurrent but often concealed by the pinna below. Veins arisingfrom whole base of pinna, nearly parallel,
often forking once at the middle of pinna,
8-10 in number in each pinna. (Cuticle
not preserved.)
Distribution and occurrence: Ptilophyllum shinadaniense is rare and known only
from the Shinadani Formation.
Discussion and comparison: Ptilophyllum
shinadaniense is characterized by its medium-sized elongate-rectangular pinnaewith parallel veins often forking once.
Ptilophylll1711 contiguum originally described by Sze (1949) and later by Wu
et al. (1980) from the Lower Jurassic
Xiangxi (formerly Hsiangchi) Formationin W-Hubei, China has pinnae of similar
form. But it is distinguished by its smallsized leaf (13.5 cm long and 1.6 cm wide)and fewer veins (4-6 in each pinna).
Ptilophyllum basekgabense originallydescribed by Barnard and Miller (1976)
from the Middle Jurassic of N-Iran has
smaller pinnae (8-15 mm long and up to
3 mm wide) with fewer veins (4-5 ineach pinna).
Some leaves of Ptilophyllum caucasicum
Doludenko and Svanidze from the Upper

271

Jurassic of Georgia resemble P_ shinadaniense, but others have very long andnarrow pinnae (Doludenko and Svanidze,
1969).
Ptilophyllum boreale (Heer) (Seward,
1917; Zamites borealis by Heer, 1874)
from the Lower Cretaceous of Greenland
is distinguished by its stout rachis andfewer veins (3-5 in number in eachpinna).
Ptilophyllum sokalense described by
Doludenko (1963) from the Jurassic of WUkraine has pinnae of similar form, but
their basiscopic margin is mostly straightor slightly contracted.
Ptilophyllum sp. A
PI. 43, Fig. 11; Text-fig. 8, 8'Material:

SHI-030 (Shinadani).

Description: Leaf medium-sized, elongated, 2.5 cm wide over the most part;neither end known. Pinnae elongate-ovalor rectangular, attached by entire baseon the upper side of rachis (2 mm wide)at an angle of 60-90 degrees, 1.4 cm longand 0.5 cm wide. Acroscopic basal angles
mostly rounded or slightly decurrent,
apices rounded or obtusely pointed. Veins
indistinct, possibly simple and parallel,
not radiating, 9 in each pinna. (Cuticle
not preserved.)
Distribution and occurrence: A single
specimen was obtained from the Shinadani
Formation.
Remarks: Our specimen is characterized
by its elongate-oval or rectangular pinnae.
Our specimen is distinguished from
Ptilophyllum shinadaniense from the same
locality by its shorter pinnae_
Some leaves regarded as Ptilophllum
caucasicum Doludenko and Svanidze (Doludenko and Svanidze, 1969; Doludenko
and Orlovskaja, 1976) from the Upper
Jurassic of Georgia and Kazakhstan are
similar in pinna form and venation toTatsuaki KIMURA and Masanori TSUJII

272

ours, but others have very long andnarrow pinnae.

Ptilophyllum sp. B
PI. 41, Fig. 7; PI. 43, Fig. 12; Text-figs. 9a-b'
Material: A-0379, 0456, 7564 (Iwamuro),
NEG-051 (Negoya).

Description: Leaves small-sized, longand narrow and nearly parallel-sided,
4 mm wide. Rachis 1 mm wide but mostly
concealed. Pinnae rhombic, attached byentire base at a wide angle on the upperside of rachis, closely set or imbricate,apices usually obtuse, acroscopic basal
margin straight or rounded, basiscopic
basal margin mostly straight but sometimes slightly contracted. Veins indistinct
arising from the whole base, parallel:
probably simple, about 5 in each pinna.
(Cuticle not preserved.)
Distribution and occurrence: This form
occurs sparsely in the Iwamuro and
Negoya Formations.
Remarks: Judging from the mode ofpinna attachment and venation, our leaves
may belong to Ptilophyllum. So far as
we know, such small-sized leaves as ours
have been described as Ptilophyllum or
Otoza71lites as follows:
Ptilophyllum
bengalense (Oldham) Schimper: Kilpper,
1968; Liassic of N-Iran. P. elongatum

Douglas: Douglas, 1969; Lower Cretaceousof Victoria (Australia), P. kazachstanicum
Genkina: Genkina, 1963; Middle Jurassicof Eastern Ural. P. marginatu7ll Prosvirjakova: Prosvirjakova, 1966; Middle
Jurassic of Mangwishlak. P. minor Kilpper: Kilpper, 1968, Liassic of N-Iran. Otozamites abreviatus Feistmantel: Feistmantel, 1879 (pars): Jurassic of Madras
Coast. O. bengalensis (Oldham and Morris)
Schimper: Oldham and Morris, 1862 (regarded by them as Palaeozamia bengalensis); Jurassic of India. O. bunburyanus
Zigno: Zigno, 1881; Middle Jurassic of
Italy. O. contiguus Feistmantel: Feistmantel, 1876; Jurassic of India. O. kilpperiana Barnard and Miller: Barnard andMiller, 1976; Middle Jurassic of N-Iran.
O. tenuatus (Leckenby) Harris: 1969; Middle
Jurassic of Yorkshire. O. vemavaramensis
Bose and Jain: Bose and Jain, 1967; Upper
Jurassic (?) of India.
Among these, our leaves are most like
Ptilophyllu71l bengalensis in pinna form.
But we refrain from determining it specifically because our leaves are all incomplete.
References
Antevs, E. (1919): Die liassische Flora desHorsandsteins.
K. Svensk. Vet.-Akad.
Handl., Bd. 59, no. 8, p. 1-71, pis. 1-6.
Barnard, P. D. E. and Miller, J. C. (1976):

Explanation of Plate 41
Fig. 1.

•Ctenozamites

sarrani (Zeiller) Harris'
A part ,of leaf WI'th t wo pmna.
f ragments.

J~dgmg from their wedge-shaped arrangement, they may represent the pinnae forkin~dichotomously below (NNW-846; counterpart of NNW-146 partly shown in Text-fig 1)
1
x L ' , X.
Figs. 3-4. Ptzlophyllum nipponicum Kimura and Tsujii sp. nov.; both, xL 3. Lower half of al~af (NNW -806, para type ; Text-fig. 6b). 4. Apical part of a leaf, with rather remotely set
pmnae (NNW -353; Text-fig. 6d, d').
Figs. 5-6. Ptilophyllum shinadaniense Kimura and Tsujii sp. nov.; both, xL 5. (SHI-007,
holotype; Text-fig. 7a). 6. (SHI-OI9, para type ; Text-fig. 7c).
Fig. 7. Ptilophyllum sp. B; a leaf-fragment (A-7564), x 2.

F~g. 2. Ptero~hyllum ex gr. propinquum Goeppert; (NNW -695),

KIMURAand TsujI!:

Early Jurassic plants

Plate 41

741.

Early Jurassic Plants

Flora of the Shemshak Formation (Elburz,
Iran). Part 3: Middle Jurassic (Dogger)
plants from Katumbargah, Vesek Gah andImam Manak. Palaeontographica, Abt. B,
Bd. 155, p. 31-117, pis. 1-15.
Bartholin, C. T. (1892): Nogle i den bornholmske Juraformation fore kommende
Planteforsteninger. Bot. Tidskr., Kjtbenhavn, Bd. 19, p. 87-115, pis. 1-6.
Bose, M. N. and Jain, K. P. (1967): Otozamites
vemavaramensis sp. nov. from the Upper
Gondwana of the east coast of India.
Palaeobotanist, vol. 15, no. 3, p. 314-315,
pI. 1.
Bose, M. N. and Kasat, M. L. (1972): The
genus Ptilophyllum in India. Ibid., vol.
19, no. 2, p. 115-145, pis. 1-14.
Brongniart, A. (1828-1838):
Histoire desvegetaux fossiles ou recherches botaniques
et geologiques sur les vegetaux renfennes
dans les diverses couches du globe. 1(1828-1837). xii+488 p., 171 pis., 2 (18371838). 72 p., 28 pis. Paris.
Chen, Y., Duan, S. Y. and Zhang, Y. C. (1979a) :New species of the late Triassic plantsfrom Yanbian, Sichuan, I. Acta Bot.
Sinica, vol. 21, no. 1, p. 57-63, pis. 1-3(in Chinese).
-, and (1979b): Ditto, III. Ibid.,
vol. 21, no. 3, p. 269-273, pis. 1-3 (inChinese) .
Delle, G. V. (1967): The Middle Jurassic floraof the Tkvarchelian Coal-Basin (TransCaucasia). In 'Palaeobotanika VI'. Komarov
Bot. Inst., Acad. Sci. USSR, Acta, ser.
8, p. 51-132, pis. 1-25 (in Russian).
Doludenko, M. P. (1963): New species of
Ptilophyllum from the Jurassic of thewestern Ukraine. Journ. Bot., Acad. Sci.
USSR, T. 48, no. 6, p. 796-805, pis. 1-4(in Russian).
Doludenko, M. P. and Orlovskaja, E. R. (1976) :Jurassic flora of the Karatau. Acad. Sci.
USSR, Trans., vol. 284, p. 1-262, incl.
pis. 1-89 (in Russian).
Doludenko, M. P. and Svanidze, Ts. I. (1969):
The late Jurassic flora of Georgia. Ceol.
Inst., Acad. Sci. USSR, Trans., vol. 178,
p. 1-118, pis. 1-81 (in Russian).
Douglas, J. G. (1969): The Mesozoic floras of273

Victoria. Pts. 1-2. Ceol. Surv. Vici.)ria,
Mem. 28, p. 1-310, incl. pis. 1-51.
Feistmantel, O. (1876): Jurassic (Oolitic) floraof Kach. In 'Fossil flora of the Gondwana
System, Vol. 2'. Mem. Ceol. Surv. India,
Palaeont. Indica, ser. 11, pts. 1-2, p. 180, pis. 1-12.
- - (1879): Fossil flora of the Upper Gondwanas on the Madras Coast. In 'The
fossil flora of the Upper Gondwanas'.
Ibid., vol. 1, no. 4, p. 191-233, pIs. 1-16.
Genkina, R. Z. (1963): Fossil flora of the
Mesozoic coal-bearing deposits of the East
Ural-field of Oesk Coal-Basin Inst. Ceol.
Exploit. Foss. Fuel, Acad. Sci. USSR, p.
1-115, pIs. 1-51 (in Russian).
Harris, T. M. (1926): The Rhaetic flora of
Scoresby Sound East Greenland. Medd.
om Cnmland, Bd. 68, nr. 2, p. 44-148,
pis. 1-13.
- - (1932): Ditto. Pt. 3: Caytoniales andBennettitales. Ibid., Bd. 85, nr. 5, p. 1133, pis. 1-18.
- - (1941): On Ptilophyllum gracile sp. n.
Ann. Mag. Nat. Hist., London, ser. 11,
vol. 8, p. 132-140.
- - (1942): Notes on the Jurassic flora ofYorkshire. 1-3. 1. Ptilophyllum caytonense
sp. n.; 2. Deltolepis crepidota gen. et sp.
n. ; 3. On Nilssonia campta and its reference to Beania gracilis. Ibid., ser. 11, vol.
9, p. 586-587.
(1946): Ditto. 31-33. 31. Ptilophyllum
pectinoides (Phillips) Morris; 32. Coniopteris arguta (L. & H.) Seward; 33.
Stenopteris nan a sp. n. Ibid., ser. 11, vol.
13, p. 392-411.
- - (1949): Ditto. 40-42. 40. Otozamites
anglica (Seward) n. comb.; 41. The narrow-leaved Otozamites species; 42. Ptilophyllum hirsutum Thomas & Bancroft andits differentiation from P. pecten (Phillips).
Ibid., ser. 12, vol. 2, p. 275-299.
- - (1953): Ditto. 58-60. 58. Bennettitaleanscale-leaves; 59. Williamsonia himas sp.
n.; 60. Williamsonia setosa Nathorst.
Ibid., ser. 12, vol. 6, p. 33-52.
- - (1961): The form and structure of
Ctenozamites cycadea. Bull. Brit. Mus.
(Nat. Hist.), Ceol., vol. 6, no. 6, p. 159-

274

Tatsuaki KIMURA and Masanori TSUjII

173, pis. 31-32.
- - (1964): The Yorkshire jurassic flora.
I I. Caytoniales, Cycadales and Pteridosperms. viii + 191 p., 7 pis. Brit. Mus.
(Nat. Hist.).
- - (1969): Ditto. III. Bennettitales. v+186
p., 7 pis. Ibid.
Heer, O. (1874): Die Kreide-Flora der arctischen Zone. K. Svensk. Vet.-Akad. Handl.,
Bd. 12, no. 6, p. 1-138, pis. 1-38.
- - (1876): Beitrage zur Jura-Flora Ostsibiriens und des Amurlandes. Acad. Imp.
Sci., St.-Petersb., Mem., T. 22, p. 1-122,
pIs. 1-31.
HSii, J., Chu, C. N., Chen, Y., Hu, Y. F. andTuan, S. Y. (1975): New genera andspecies of the late Triassic from Yungjen,
Yunnan, Ii. Acta Bot. Sinica, vol. 17, no.
1, p. 70-76, pis. 1-6 (in Chinese).
Iminov, Y. C. (1968): New Jurassic plantsfrom Middle Asia and North Afghanistan.
In 'The Palaeobotany of Uzbekistan 1'. p.
95-104, pIs. 35-37. FAN, Tashkent.
- - (1976): Cycadophytes of the late Triassic
and jurassic of Middle Asia and theirstratigraphic significance. 72 p., 48 pis.
Ibid.
Jacob, K. and Jacob, C. (1954): Cuticularstudy of Indian Ptilophyllum fronds from
Cutch and Jubbulpore. Mem. Ceol. Surv ..
India, Palaeont. Indica, N. S., vol. 33, no.
1, p. 1-34, pis. 1-10.
Jones, O. A. and Jersey, N. J. de (1947): Theflora of the Ipswich Coal Measures-Morphology and floral succession.
Univ.
Queensl. Pap., Dept. Ceol., N. S., vol. 3,
nos. 3-4, p. 1-88, pIs. 1-10.
Kilpper, K. (1965): Nomenklatur und Ges-

chichte mesozoischer Pflanzen. 1. Die
Gattung Ptilophyllum Morris (in Grant).
N. jb. Ceol. Paliiont. Mh., Bd. 4, p. 193205.
(1966): Ptilophyllum kharganense n. sp.
aus der Kharga-Oase, westliche Wiiste,
Agypten (Mittel Jura ?).
Palaeontographica, Abt. B, Bd. 117, p. 75-82, pI.
30.
- - (1968): Einige Bennettiteen-Blatter aus
dem Lias von Karmozd-Zirab (N-Iran). j.
Linn. Soc. (Bot.), vol. 61, no. 384, p. 129135, pIs. 1-3.
Kimura, T. (1959a): Mesozoic plants fromthe Iwamuro Formation (Liassic), Tonegun, Gumma Prefecture, Japan. Bull.
Sen. High Sch., Tokyo Univ. Educ., no.
3, p. 1-59, incl. pIs. 1-12.
- - (1959b): Mesozoic plants from the Kotaki
Coal-Field, the Kuruma Group, Central
Honshu, Japan. Ibid., p. 61-83, incl. pis.
1-2.
Kimura, T. and Tsujii, M. (1980a): Early
Jurassic plants in Japan, Part 1. Trans.
Proc. Palaeont. Soc. japan, N. S. no., 119,
p. 339-358, pis. 38-40.
- - and - - (1980b): Ditto. Part 2. Ibid.,
120, p. 449-465, pis. 54-56.
- - and - - (1981): Ditto. Part 3. Ibid., no.
no. 124, p. 187-207, pIs. 30-32.
Li, P. C. (1964): Fossil plants from the Hsuchiaho Series of K wangyiian, Northern
Szechuan. Mem. Inst. Ceol. Palaeont.,
Acad. Sinica, no. 3, p. lOl-178, pIs. 1-20
(in Chinese with English description).
Li, P.C., Tsao, C.Y. and Wu, S.C. (1976):
Mesozoic plants from Yunnan. In 'Mesozoic fossils of Yunnan, Vol. 1'. p. 87-

Explanation of Plate 42
Figs. 1-5. Otozamites neiridaniensis Kimura and Tsujii sp. nov.; all figures, xLI, 3. Leavesof typical size. 1; (NNE-095, holotype; Text-fig. 3a), 3; (NE-046). 2. A large-sized leaf
(TOB-050B', para type ; Text-fig. 3c). 4. Basal part of a leaf (NEE-250). 5. Ditto (NEE073).
Figs. 6-7. Pterophyllum ex gr. propinquum Goeppert; both, x 1. Leaf-fragments. 6; (NNW073), 7; (NNW -600) .
Fig. 8. Ptilophyllum nipponicum Kimura and Tsujii sp. nov.; apical part of a leaf (NEE-074;
Text-fig. 6e), xL

KIMURAand TsuJI!:

Early Jurassic plants

Plate .42

741. Early Jurassic Plants
160, pIs. 1-47. Ke-Xue-Chu-B3.n-She, Peking(in Chinese).
Lundblad, A. B. (1950): Studies in the RhaetoLiassic floras of Sweden. 1. Pteridophyta,
Pteridospermae, and Cycadophyta from the
Mining District of NW Scania. K. Svensk.
Vet.-Akad. Handl., N. S., Bd. 1, no. 8, p,
1-82, pIs. 1-13.
Moller, H. (1903): Bidrag till Bornholms
Fossila Flora (Rhat och Lias). Gymnospermer. Ibid., Bd. 36, no. 6, p. 3-48, pIs.
1-7.
Oishi, S. (1931): Mesozoic plants from KitaOtari, Prov. Shinano, Japan. Journ. Fac.
Sci., Hokkaido Imp. Univ., ser. 4, vol. 1,
no. 2, p. 223-255, pIs. 16-18.
- - (1940): The Mesozoic floras of Japan.
Ibid., vol. 5, nos. 2-4, p. 123-480, pis. 1-48.
Prosvirjakova, Z. P. (1966): Jurassic flora of
Mangwishlak and its significance forstratigraphy. 174 p., 35 pis. VSEGEI,
Acad. Sci. USSR (in Russian).
Oldham, T. and Morris, J. (1862): Fossil floraof the Rajmahal Series in the Rajmahal
Hills. In 'Fossil flora of the Gondwana
System, Vol. 1, pt. 1'. Mem. Ceol. Surv.
India, Palaeont. Indica, ser. 2 (1-4), p.
1-52, pIs. 1-35.
Seward, A.C. (1917): Fossil plants. Vol. 3.
xviii+656 p. Cambridge.
Sze, H. C. (1949) : Die mesozoischen Flora ausder Hsiangchi kohlen Series in West Hupeh,
China. Palaeont. Sinica, N. S., ser. A,
no. 2, p. 1-71, pis. 1-15.
- - (1956): Older Mesozoic plants from the
Yenchang Formation, Northern Shensi.
Ibid., no. 5 (139), p. 1-110 (Chinese part),
111-217 (English part), pis. 1-56.
Sze, H. C., Li, X. X., Li, P. C., Wu, S. C., Zhoh,
T. Y., Ye, M. N. and Shen, G. L.. (1963):
Fossil plants of China, Vol. 2. Mesozoic
plants of China. iv+429 p., 118 pis. KeXue-Chu-Ban-She, Peking (in Chinese).
Vakhrameev, V. A. and Doludenko, M. P.
(1961): Late Jurassic and early Cretaceous
floras from the Bureja Basin and its significance for stratigraphy. Ceol. Inst.,
Acad. Sci. USSR, Trans., vol. 54, p. 1-

275136, pis. 1-60 (in Russian).
Walkom, A. B. (1917): ~1esozoic floras ofQueensland. Part I.-concluded. The floraof the Ipswich and Walloon Series. (d) - (f) .
Queensl. Ceol. Surv. Publ., no. 259, p. 148, incl. pis. 1-9.
Weber, R. (1968): Die fossile Flora der RhatLias-VbergangschichtenvonBayreuth(Oberfranken) under besonderer Berucksichtigung der Coenologie. Erlanger geol.
Abh., Heft 72, p. 1-73, pis. 1-17.
Wu, S. Q., Ye, M. N. and Li, B. X. (1980):
Upper Triassic and Lower and Middle
Jurassic plants from the Hsiangchi Group,
Western Hubei. Mem. Nanjing Inst. Ceol.
Palaeont., Acad. Sinica, no. 14, p. 63~131,
pIs. 1-39 (in Chinese wi th English abstract).
Yabe, H. and Oishi, S. (1929): Notes on somefossil plants from Korea and China belonging to the genera Nilssonia and Pterophyllum. Jap. Journ. Ceol. Ceogr., vol. 6,
nos. 3-4, p. 85-101, pis. 18-20.
- - and - - (1933): Mesozoic plants from
Manchuria. Sci. Rep., Tohoku Imp. Univ.,
sec. ser., vol. 12, no. 2B, p. 195-238, pIs.
30-35, 1 tab.
Yang, X. H. (1978): Palaeontological atlas ofSouthwest China. Division of Sichuan.
Part 2. p. 469-684, pis. 155-190. Di-ZhiChu-Ban-She, Peking (in Chinese).
Zeilkr, R. (1903): Flore fossile des gites deCharbon du Tonkin (Etudes des gites
mineraux de Ia France). 328 p., 56 pis.
Paris.
Zhou, T. S. (1978): On the Mesozoic coalbeanng strata and fossil plants from Fujian
Province. Prof. Pap. Stratigr. Palaeont.,
no. 4, p. 88-134, pis. 15-30 (in Chinese).
Zigno, A. de (1856-1885): Flora Fossilis
Formationis Oolithicae. Le Piante Fossili
dell'Oolite. 2 vols. Padova. Vol. 1, Livr.
1, p. 1-32, pIs. 1~6 (1856) ; Livr. 2, p. 3364, pIs. 7-12 (1858) ; Livr. 3-5, p. 65-223,
pIs. 13-25 (1867) ; Vol. 2, Livr. 1, p. 1-48,
pis. 26~29 (1873); Livr. 2-3, p. 49-120,
pIs. 30-37 (1881); Livr. 4-5, p. 121-203,
pIs. 38-42 (1885).

276

Tatsuaki KIMURA and Masanori TSUJI!

Geographical names in Japan: Agero J:lMt, Dairagawa 7cfJI[, Higashi-Neiridani *'ilA.'fr,
Inumatadani ::kf!R'fr, Iwamuro !5~, Kita-Otari ~t'J''fr, Kotaki 'Nil!:, Kuruma *.~, Mone ~tH, Nariwa illt;r:], Negoya i);(J.7E'fr, Neiridani 'ilA.'fr, Nishi-Neiridani itS'ilA.'fr, Shinadani tIi't'fr,
Teradani ~'fr, Tobiiwadani ~!5-'fr, Tsuchizawa (Tunzawa) ±tR, Geographical names in
China (with former expression in parentheses): Beipiao (Peipiao) ~t~, Daqiaode 7cjliijJfu,
Fujian liii}t, Hubei (Hupeh) mJ~t, Liaoning ji,¥, Sichuan (Szechuan) ~JI[, Taipingchang(Tai-ping-tchang) :;tf~, Wengbinshan )(J(Ll!, Xujiahe (Hsuchiaho) ~*i'PJ, Xiangxi (Hsiangchi) tt~, Yanbian :l£ill, Yanchang (Yenchang) ~:R:, Yipinglang -f¥,N, Yongren(Yungjen) 7k1=, Yunnan ~i¥i, Geographical name in Korea (with former expression) : Daedong(Taedong) 7ci'U0) '/13*0)tJ]1t!l:).o.7*cim1mfr.:fio -to) 4: llfjijfrn1::'''::J--::i~, *.~J\~H'f;fo'J:rY!5~~JiliJ'G"::J~'
jI'fiJ(f;fo' J: V' ~;1< 7-7 7. @H::.~-t 0 im1mfr.:fi:a:-r.cll U~o'/ jI"filii" § : Ctenozamites sarrani (Zeiller) Harris~;1< 7- 7 7. § : Anomozamites sp. A, Otozamites fujimotoi Kimura, O. neiridaniensis
Kimura and Tsujii sp. nov., Pterophyllum ex gr. propinquum Goeppert, P. sp. A,
Ptilophyllum dr. cutchense Morris, P. nipponicum Kimura and Tsujii sp. nov., P.
shinadaniense Kimura and Tsujii sp. nov., P. sp. A, P. sp. B.
tJJ:O) 5 i?, Ctenozamites ~ft, 13*-C':i&tJ]O)fflli-c'j;) I), Ptilophyllum ~ft, 13*O)tlJW9:t~WiJ'GO)iIHJJO)~lli-c'j;)00

C <1::'1&~0)~llift, I3*O)tJ]JW:).o. 7 *cim4&.rllf 0) tl

im1m~~~&~:a:-~~-t0J:'C.~~.Mc~0o*M~~·ft*~~

Explanation of Plate 43
Figs. 1-2. Otozamites neiridaniensis Kimura and Tsujii sp. nov.; both, xl. l. A large-sizedleaf-fragment with elongated pinnae (NNW -084, para type ; partly shown in Text-fig. 3f).
2. Basal part of a leaf (TOB-032).
Figs. 3-6. Pterophyllum ex gr. propinquum Goeppert; all figures, x l. Leaf-fragments. 3;(NNW -240), 4; (NNW -356B), 5; (NNW -356; Text-fig. 4j), 6; (NNW -014).
Figs. 7-8. Ptilophyllum cfr. cutchense Morris; both, x l. Unsatisfactorily preserved leaf-fragments. 7; (TE-OOl; Text-fig. 5a), 8; (TE-003; Text-fig. 5b).
Figs. 9-10. Ptilophyllum nipponicum Kimura and Tsujii sp. nov; both, xl. 9. A leaf lacking
both apical and basal portions (NEG-270, holotype; Text-fig. 6a, a/). 10. Basal part of aleaf with petiole of which base is expanded; upper arrow indicates a thick rachis and lower
one an expanded petiole-base (NNW -111, para type; Text-fig. 6k).
Fig. 1l. Ptilophyllum sp. A; a leaf-fragment (SHI-030; Text-fig. 8), xl.
Fig. 12. Ptilophyllu11l sp. B; a leaf-fragment (NEG-051; Text-fig. 9a), x 2.

KIMURA and TSUjIl:

Early Jurassic plants

Plate 43

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 125, pp. 277-295, pis. 44-48, April 15, 1982

JAPANESE MIOCENE CIRRIPED BALANUS SENDAICUS:
A COMPARISON WITH TETHYANBALANUS CONCA VUS G RODP*

7-12.

TOSHIYUKI YAMAGUCHI
Geological Institute, Faculty of Science, University of Tokyo, Tokyo 113

Abstract. Balanus sendaicus was described as a new species by Hatai, Masuda
and Noda (1976) based on a single scutum from the Miocene Moniwa Formation,
northern Honshu, Japan. However, the scutum deposited in the Saito Ho·on Kai
Museum, Sendai, Japan (Hatai et al., 1976) could not be found. Tergum and shellwall, which with scutum comprise the skeletal elements, had not been described.
According to my investigation of topotype materials and of materials illustrated by
Hatai et al. (1976), the shell walls described and illustrated under the name of B.
rostratus by Hatai et al. (1976) belong to B. sendaicus. B. sendaicus, including tergum
undescribed by Hatai et al. (1976), is therefore defined in this paper. B. sendaicus
is distinct from the already known taxa, as mentioned by Hatai et al. (1976). However, B. sendaicus is closely related to the extinct B. concavus sensu de Alessandri(1906), which flourished in the Mediterranean region (Tethys) from the Oligocene toPleistocene. B. sendaicus belongs to the B. concavus species group represented by
the nominal species based on having transverse septa in longitudinal (parietal) tubesof shell wall and the characteristic ornamentation of scutum. Disappearance of theB. concavus group from the Atlantic and the Mediterranean regions in Pleistocene ismysterious as mentioned by Pilsbry (1916), though "four" species belonging to theB. concavus group are living in the eastern Pacific (Newman, 1979). B. sendaicus isrestricted to the Miocene, is the first fossil record of the B. concavus group from theeast Asia, and indicates the presence of a connection between the Tethys and theeast Asia.

Four new species including B. sendaicus
were based on scuta and three new species,
B. rostratZis and B. sp. based on the shellwalls. No tergum was described.
Fossil specimens are usually found as
fragmental skeletal elements of shell walland opercular valves, and therefore it isdifficult to reconstruct the exoskeleton ofa species from the mixture of fragmentsof several extinct species. Under thesecircumstances, each fragmental skeletal
element, such as shell wall, scutum or
tergum, has been frequently chosen as

Introduction
Balanus sendaicZls was proposed as a
new species by Hatai, Masuda and Noda
(1976), based on a single scutum from the
Miocene Moniwa Formation of the Sendai
area of northeast Honshu, Japan. Seven
new species including B. sendaicus, analready known species of B. rostratus,
and an in determined species of Balanus
were described from the same locality.

* Received June 30, 1981; read Jan. 24, 1981
at Sendai.

277

Toshiyuki YAMAGUCHI

278

representative of a new species. Thus insome cases, an individual consisting of
three kinds of skeletal elements, maypossesses three different species names.
Therefore, biological study of fossil
specimens of extinct species is faced withdifficulties unless basic work, revealing
the precise combination of skeletal elements, is done first. When a complete
shell wall is discovered, it is necessary to
determine whether or not opercular valves
are preserved in the shell cavity.
In this paper the following points are
considered: 1) Description of shell walland tergum of B. sendaicus, whose originaldescription was based on a single scutum,2) The status of B. sendaicus as a previously unknown taxon, as discussed by
Hatai et al. (1976), 3) The affinity of B.
sendaicus with known species and species
groups, and 4) The stratigraphic andgeologic distributions in Japan and thepaleoecological significance of B. smdaicus.

Balanus sendaiclls, type-locality
and formation

The type-locality of this species is an
outcrop (Loc. 4) of the Miocene Moniwa
Formation situated about 200m south of the
Oide Bridge across the Natori River, near
Moniwa, Sendai City, Miyagi Prefecture,northeast Honshu, Japan. The Miocene
Moniwa Formation overlies the Miocene
Takadate Andesite with a clino-unconformity. B. sendaicus was collected fromthe basal part of the Moniwa Formation
which consists of gravelly calcareous
medium-grained sandstone containing very
abundant well preserved marine fossils.
Reports on brachiopods, shark teeth, fossil
problematica, and on pelecypods havealready been published (Hatai et al., 1973,
1974a, 1974b, 1974c, respectively). A
report on cirripeds was the fifth of thisseries from the same locality. Besides
B. sendaicus, shell walls but no terga ofB. rostratus, B. miyagiensis, B. tolzokuensis,
B. rikuzenensis, and B. sp. and scuta of
B. oidensis, B. moniwaensis, and B. natorianus were described. Except for B.
rostratus and B. sp., all were proposed as
new species.

Combination of skeletal elements of

B. sendaiclls
The writer has collected many wellpreserved fossil cirri peds from the above
locality. Four species by shell walls, fivespecies by scuta, and two species by terga
were distinguished. However, the combination of these exoskeleton elements
was not clear. Judging from shell size
and the frequency of occurrence of eachelement, it was presumed that shell wallof B. rostratus and scutum of B. sendaicus,
as described by Hatai et al. (1976) and apreviously undescribed tergum probably
constituted skeletal elements of one species.
This is also supported by the fact thatthe following morphological characteristics
of shell walls of B. rostratlls identified
by Hatai et al. (1976) are clearly different
from the true B. rostratus proposed by
Pilsbry (1916); narrow radii and oblique
summits of radii. In order to confirm theabove conjecture additional material wassought.
Recently some shell walls collected fromthe same locality, identical with B. rostrat liS of Hatai et al. (1976), were foundto contain paired scuta and terga within
their shell cavity. The specimen of B.
rostratus illustrated by Hatai et al. (1976,
pI. 2, fig. 2, left, and PI. 44. Figs. 2a-c,
left, in this paper) was also found tocontain four opercular valves in its shellcavity (PI. 44, Figs. 2d-g, in this paper).
It is important to note that morphologically742.

Miocene Cirriped Balanus sendaicus

these scuta are identical with the holotype scutum of B. sendaicus (Hatai et al.
1976, pI. 2, figs. 9 and 10). It can, therefore, be concluded that B. sendaicus iscon specific with B. rostratus of Hatai etal. (1976). However, B. rostratus of Hatai
et al. (1976) is not the true B. rostratus
in the senS3 of Pilsbry (1916) and thename B. sendaicus should be used. As aresult of the confirmation of the correct
combination of skeletal elements, thetaxonomic status and the affinity of B.
sendaicus can be discussed.
In the description of B. sendaicus, Hatai
et al. (1976) pointed out that "In the external sculpture, the present scutum resembles that of Balanus concavus Bronn(Pilsbry, 1916, p. 100, pI. 21, fig. 1), a
Pliocene species of the British Red Cragespecially in cancellated sculpture, bifurcation of the longitudinal striae, generalshape of scutum, ... ". Thus, owing to distinct growth ridges and longitudinal ridges
or striations frequently bifurcated on external surface, scutum of B. sendaicus
possesses common morphological characteristics found in scuta of all membersof the B. concavus species group. The
tergum of B. sendaicus described here
also has a long spur and partially closed
spur furrow, characteristics found in somemembers of the B. concavus group. Thesemorphological characteristics found in B.
sendaicus are also present in members of
the B. perforatus species group. However,in members of the B. concavus group, theexternal surface of scutum is heavily
ornamented, the aperture is small, andthe carina strongly curved. Therefore,
B. sendaicus is regarded as a member ofthe B. concavus group.

279

Morphological comparisons of B.
sendaicus with the living taxaof the B. COllcaVllS group
According to Newman and Ross (1976),
the B. concavus group contains the following "five" living taxa, B. aquila (=B.
gregarius (Conrad), 1856; a senior synonymof B. aquila Pilsbry, 1907, see Zullo,
1964: 360, 1969: 6, 1979: 4), B. concavus
concavus, B. c. mexicanus, B. eyerdami(=B. panamensis Rogers, 1948; a seniorsynonym of B. eyerdami Henry, 1960, D.
P. Henry, pers. comm., in Newman andRoss, 1976: 102), and B. regalis. B. COI1cavus concavus of Newman and Ross (1976)
includes the two already known "B. c.
indicus" and B. c. sinensis as synonyms of
B. c. concavus (Newman and Ross, 1976).
B. c. mexicanus lacking the transverse
septa. in the parietal tubes of shell wall
does not belong to the B. concavus groupin the strict sense, but is close to B.
pacificus of the B. perforatus group
These four
(Newman. pers. comm.).
extant members exclusive of B. c. mexicanus are living in Indo-Pacific, B. c. concavus in Dar-es-Salaam of Tanzania, thePersian Gulf and Hong Kong, and theother three species on the eastern Pacific.
In the following paragraph, B. sendaicus
is compared with these four "living" taxaof the B. concavus group on the b::lsis ofmorphology.
B. gregarius (Conrad), 1856 was originally described as Tamiosoma gregaria,
a new genus and species of a rudistid
pelecypod by having the peculiar, elongate,vesiculose basis. This species is the mostwidespread and commonly encounteredbarnacle in the Miocene and Pliocene
formations of central and southern California and Baja California regions (Zullo,
1964, 1969, 1979). "It was not until theopercular valves were described by Woodring", in Woodring et al. (1940), "that any

280

Toshiyuki YAMAGUCHI
Table 1. List of members of the B, concavus group and geographic and stratigraphic
ranges quoted from Newman and Ross (1976) except for B. concavus concavus
and "B. concavus indicus". [Oligo.: Oligocene, Mio.: Miocene, Plio.: Pliocene,
Pleisto. : Pleistocene]

Balanus bloxhamensis Weisbord, 1966 :48, fossil, (Florida; Mio.)B.
concavus concavus Bronn sensu de Alessanri, 1906: 295, fossil (Europe; Oligo.-Pleisto.)B.
c. alloplax Pilsbry and Olsson, 1951 :200, fossil, (Ecuador; Oligo.)B.
c. chesapeakensis Pilsbry, 1916: 103, fossil, (Maryland; Mio.)B.
c. coosensis DaIl, 1909: 138, fossil, (Oregon; Mio.)B.
c. dal/onii Davadie-Suaudeau, 1952: 20, fossil, (Algeria; Plio.)B.
c. eseptatus Pilsbry, 1924: 1, fossil, (Haiti; Mio.)B.
c. finchii Lea, 1833 :211, fossil, (Maryland; Mio.)B.
c. glyptopoma Pilsbry, 1916: 102, fossil, (east USA; Mio., Panama &east Mexico; Plio.)B.
"c. indicus" Nilsson-Cantell, 1932 :2, living, [Dar-es-Salaam, Tanzania]B.
c. oligoseptatus Kolosvary, 1961: 149, fossil, (USSR; Oligo.)B.
c. proteus Conrad, 1834: 134, fossil, (east USA; Mio.-Plio.)B.
c. raphanoides Moroni-Ruggieri, 1952 :71, fossil, (Italy; Plio.)B.
c. rariseptatus Pilsbry, 1918: 186, fossil, (Panama; Mio.)B.
c. rubescense Seguenza, 1876 :450, fossil, (Italy; Tertiary)B.
c. scutorum Seguenza, 1876 :74, fossil, (Italy; Plio.)B.
gregarius (Conrad), 1856 :315, living, [California, 0-18 m], fossil, (central, southern &Baja California; Mio.-Plio.)
B.
indicus Withers, 1923: 291. fossil, (Pakistan; Mio.)B.
panamensis Rogers, 1948 :95, living, [Gulf of California to Pacific coast of Panama]B.
polyporus Pilsbry, 1924 :2, fossil, (Haiti; Mio.)B.
regalis Pilsbry, 1916: 108, living, [southern & Baja California]B.
sendaicus Hatai, Masuda and Noda, 1976: 13, fossil, (Japan; Mio.)B.
talquinensis Weisbord, 1966: 37, fossil, (Florida; Mio.)B.
vadaszi Kolosvary, 1949 :2, fossil, (Europe; Mio.)

attempt could be made to determine theaffinities of this unique species." "W oodriing concluded that B. gregarius was related to the B. concavus Bronn group.""Comparison of well preserved opercularvalves with those of the extant Pacificcoast species B. aquila reveals no apparent
differences. The only major distinctionin the two species would appear to be atendency for the formation of elongate
cellular basis in the fossils. However,this unique basis has now been discovered
in extant specimens of B. aquila taken by

the Allan Hancock Foundation off Santa
Rosa Island, California. It is probablethat the prevalence of elongate basis in
fossils is related to past environmental
conditions (warm, shallow embaymentswith high sedimentation rates) which are
not widespread along the Pacific coast
today." (Zullo, 1964: 360). The morphological characteristics of this species, suchas deeply sunken radii, growth ridge of
scutum divided into beads by striations,and the rather narrow spur of tergum
considerably close to the basiscutal angle,

742.

Miocene Cirriped Balanus sendaicus

Table 2. Morphological comparisons of B.
sendaicus with the extinct B. concavus
concavus sensu de Alessandri (1906) andthe living taxa of the B. concavus group.
0: agreed; x : disagreed. Informations
on parietal tubes of shell wall in fossil
European B. c. concavus and living B.
gregarius are by courtesy of Newman(in press, pers. comm.).

'"
;:!
;:,
'"
'"
l':!
.:;
'"
'"
'";:! '";:! '"'" '"l':!
;:, .;:: ;:!
;:, '" .~
!::
'-'
-",

;:!

~

~

.~

.
'" ....
~

l':!

B. sendaicus

(;j
~

Qi
..c::

::1

~

'"'-'

.;::,.

'"....

<:;~I radii widelOx x x x

summits of radii
oblique

10 x OOx
I: surface strongly
E I sculptured

u
Cfl

~

~

'-'

l':!

surface smoothparietal tubes with
septa

::l

~

~ ~ ~ ~~
i 0 x OOx

Cfl

B

'-'

~

Ix 0000

10 x x x xI

adductor ridge weak

lOx
x 0 x
I

spur furrow closed

I x OOx 0
I§ I spur shortlOx x x x
~ I basal margin straight Ix x 000
bj)

are different from those of B. sendaicus.
B. pana71lensis Rogers, 1948 [Holotype,
Calif. Acad. SCi. Dept. Paleo. Type ColI.
9434; paratypes, 9435-9437J was collectedfrom C. A. S. Loc. 32532 of Chame Point,
Canal Zone, Panama. Nature of shellform, toothed orifice, internal and basal
margins of scutum, external surface oftergum, and basal margin of tergum foundin this species are similar to those of B.
sendaicus. Relatively narrow radii in comparison with B. sendaicus, weakly scul-

281

ptured external surface of scutum, andshallow and opened spur furrow of tergum
found in this species are apparently distinct from those of B. sendaicus. Nevertheless, among the American living taxa
B. pana71lensis resembles the fossil Japanese B. sendaicus.
B. regalis Pilsbry, 1916 [Holotype,
USNM 43485J was described from Point
Abreojos, west coast of Lower California
based on shell wall without opercularvalve. Rugged shell with irregularly arranged ribs of various size, extremely
narrow radii, and moderately projecting
and short adductor ridge of B. regalis
distinguishes it from B. sendaicus. Morphological observation of opercular valves
is based on samples obtained through thecourtesy of Professor Newman. This
species resembles B. sendaicus in the natureof spur furrow, parietal tubes of shellwall with transverse septa, and basal
margin of tergum.
As mentioned above, the three living
taxa distributed along the west coast of
USA and central America are morphologically distinct from the fossil Japanese
B. sendaicus. The morphological difference
suggests the species level distinction between B. sendaicus and the three livingtaxa. Among the three living taxa, B.
pana71lensis has more characters in common
with B. sendaicus than the other two
species. Thus it can be inferred that,among the living American species of theB. concavus group, B. pana71lensis is related
to extinct B. sendaicus.
In Indo-Pacific area, survival of the infraspecific taxa of B. concavus is indicatedby only three reports: Broch (1931), Nilsson-Cantell (1932) and Utinomi (1969).
Their taxa were included in the B. c.
concavus of Newman and Ross (1976).
According to Newman and Ross (1976),
studies on B. c. concavus except for thethree reports mentioned above are con-

282

Toshiyuki YAMAGUCHI

cerned with fossil materials.
In Broch's type specimen of B. c.
sinensis from Hong Kong, Newman andSouthward "have distinguished two
species, B. reticulatus and B. variegatus".
"They have concluded that B. c. sinensis
was a junior synonym of B. variegatus
and that B. reticulatus of Utinomi remains
valid (Newman and Southward, pers.
comm.)" (Yamaguchi, 1980: 1088).
Based on reexamination of Utinomi(1969),s materials by Newman, B. c. concavus from the Persian Gulf reported by
Utinomi (1969) is Chi rona d. amaryllis
previously called B. amaryllis, because it
does not have the parietal tubes in shellwall and has denticular ribs in inner baseof parietes (Newman, pers. comm.).
Therefore, the decision whether B. c. concavus of Newman and Ross (1976) is extant
or not should be determined by reexamination of "B. c. indicus".
"B. concavus indicus" Nilsson-Cantell,
1932 was described based on only one
specimen collected from a buoy in Dar-esSalaam, Tanzania and has been preservedin the Berlin Museum. However, the typespecimen can not be found in the Museum(Newman, pers. comm.). Live "B. c. indicus" has not been reported since 1932.
Name indicus is preoccupied by B. indicus
Withers, 1932. Therefore "B. c. indicus"
is a homonym of B. indicus. However,both are apparently different from eachother in morphology, so a replacement
name is proposed by Newman (in press,
pers comm.). Therefore," B. c. indicus" oftaxonomically invalid name is used in thispaper.
Before entering to morphological comparison of B. c. concavus with "B. c. indicus", it will be historically reviewed on
taxonomy of B. c. concavus. B. concavus
was described by Bronn (1831) from the
Pliocene of Andona Valley, near Asti,
Piedmont, Italy (Pilsbry, 1916). However,the substance of B. concavus was notalways clear until de Alessandri (1906)'s
redescription of topotype materials.
Darwin (1854a) reported three different
forms of terga under the name of B. concavus: 1) terga of pI. 1, figs. 4g from
Coralline Crag, England (fossil) and of pI.
1, fig. 40 from Turin, Italy (fossil) having
a narrow spur furrow and basal margin
slightly depressed; 2) tergum of pI. 1, fig.
4k from Maryland, USA (fossil) having avery wide spur furrow and a straightbasal margin; and 3) tergum of pI. 1, fig.
4m from Portugal (fossil, recent ?) having
certainly opened spur furrow and basal
margin slightly projected. These morphological differences are usually regarded
to represent distinction of species level,
therefore, it is not appropriate that those
three forms of terga are summarized to a
species of B. concavus. Darwin (1854b)
also reported two different forms in terga
under the name of B. concavus. Specimens
of pI. 4, figs. 4d and 4e and specimen of
pI. 4, fig. 4c in Darwin (1854b) are morphologically identical with those of pI. 1,
figs. 4g and 40 and that of pI. 1, fig. 4m
in Darwin (1854a) respectively.
After Darwin (1854a, b), studies of B.
concavus have been subjected to his great
influence, and three distinct forms haveprobably been treated as a species of B.
concavus. This fact seems to be a maincause for the taxonomic confusion of B.
concavus.
Revision of topotypes (pI. 16, figs. 2125; pI. 17, figs. 1a-4b) of B. concavus
Bronn by de Alessandri (1906) seems to
be found a clue to solve the taxonomic
confusion and pointed out that terga of
the real B. concavus are identical withthose of pI. 1, figs. 4g and 40 in Darwin(1854a) and of pI. 4, figs. 4d and 4e in
Darwin (1854b). Therefore, B. concavus
redescribed by de Alessandri (1906) iscalled B. c. concavus sensu de Alessandri

742. Miocene Cirriped Balanus sendaicus(1906) in this paper. It is interpreted that
B. c. concavus of Newman and Ross (1976)
contains five or more species of B. c.
concavus sensu de Alessandri, "B. c. indicus", B. c. sinensis (=B. amphitrite s.
1.), and at least two species of Balanus(?) eliminated B. c. concavus sensu de
Alessandri from Darwin (1854a)'s B.
concavus.
In this paper, B. concavus in the following references is considered to be identicalwith B. concavus concavus Bronn in thesense of de Alessandri (1906): Darwin,
1854a, p. 17 (in part), figs. 4g and 40;
Darwin, 1854b, p. 235 (in part), pI. 4, figs.
4d and 4e; de Alessandri, 1895, p. 282, pI.
2, figs. 10a-l0e; Pilsbry, 1916, p. 100, pI.
21, figs. l-lc; Davadie, 1963, p. 52, pI. 24,
figs. 8-9, pI. 26, figs. 1-4, pI. 28, figs. la2b; and Menesini, 1965, p. 110, pI. 28, figs.
2-6, pI. 29, figs. 1-9, pI. 30, figs. 1-4, pI.
34, figs. 7-8, pI. 35, figs. 1-2, pI. 41, figs.
3-8, pI. 42, figs. 1-8, pI. 43, figs. 1-3.
"B. c. indicus" and B. c. concavus sensu
de Alessandri (1906) have common morphological characteristics, such as conicalshell form, rhomboidal orifice, oblique
summits of radii, external scutal surface
with longitudinal striations, relative spur
width for basal length of tergum, relative
position of spur, shape of basal marginof tergum, etc. However, the following
morphological differences are present:
shell surface is smooth in "B. c. indicus",
but costated, sulcated, festooned, or withlongitudinal ribs in B. c. concavus. Parietal
tubes of shell wall have not transverse
septa in "B. c. indicus", but have transverse septa in B. c. concavus (Informationon this shell structure is by courtesy ofProfessor Newman, pers. comm.). The
adductor ridge is very low and short in"B. c. indicus", but prominent, long, clear
and slightly curved in B. c. concavus.
Tergal surface has longitudinal striae on
carinal side of "B. c. indicus", but is283

longitudinally and transversally sulcatedin B. c. concavus. Spur furrow is open
in "B. c. indicus", but closed in B. c. concavus. The morphological resemblances
show close relationship between them,
however, the morphological distinctions
are usually sufficient to recognize themas different species. As mentioned above,
B. c. concavus sensu de Alessandri (1906)is neither an extant species nor "B. c.
indiclls", therefore, B. c. concavus is an
extinct species.
B. sendaicus and B. c. concavus sensu de
Alessandri (1906) closely resemble eachother in their morphology. The common
characteristics are conical shell shape,rhomboidal and toothed orifice, havingtransverse septa in parietal tubes of shellwall, striated external surface of scutum,
etc. In B. sendaicus, shell wall is generally smooth, but occasionally costated in
carina and carinolateral. This character
resembles that of B. c. concavus. The two
species, however, differ morphologicallyin the following characters: high, longand clear adductor ridge of scutum in B.
c. COllcavus, but obscure in B. sendaicus;
long spur length in B. c. concavus, butmoderately long in B. sendaicus; and surface of tergum longitudinally and transverselly sulcated in B. c. concavus, but
having only weak growth lines in B.
sendaicus. These morphological differences
are sufficient enough to distinguish themas two distinct species. Morphologicalresemblance, however, suggests their close
relationship.
Furthermore, when B. sendaicus and"B. c. indicus" compared on the basis ofmorphology, they are found to resemble
each other. The conical form, rhomboidal
and moderately wide orifice, smooth shellsurface (in B. sendaicus, carina and carinolateral are often costated, but generallysmooth), scutum with well developed
growth lines and strong striae on external

284

Toshiyuki YAMAGUCHI

surface, and crenulation of basal marginof scutum are their common character.
However, they have four different characteristics: parietal tubes of shell wall
have not transverse septa in "B. c. indicus", but have the septa at upper halfof shell wall in B. sendaicus, though usually secondarily filled up with calcite as
result of fossilization; the adductor ridgeof scutum is very low and short in "B. c.
indicus", but obscure in B. sendaicus; distance from basiscutal angle to spur of
tergum approximately is equal to spur
width in "B. c. indicus" (Henry and
McLaughlin, 1975), but is 2/3 in B. sendaicus (average 25 individuals examined);and the spur furrow is open in "B. c. indicus", but partially closed in B. sendaicus.
Thus, morphological distinction is clear
and sufficient enough to separate theminto two distinct species.
B. sendaicus was described by Hatai et al.
(1976) based on only one scutum (Holotype).
However, the shell walls of B. sendaicus
were described under the name of B.
rostratus Hoek, and the tergum wasundescribed. Thus, B. sendaicus of Hatai
tit al. (1976) clearly does not agree with
B. sendaicus redescribed here. However,as mentioned above, B. sendaicus redefined
here is not identical with the alreadyknown living species of the B. concavus
group.

Comparison with the extinct taxaof the B. conc(wus group

B. bloxhamensis Weisbord, 1966 [Holotype, PRI (Paleontological Research Institution) 27408J was described from thelate Miocene Choctawhatchee Formation
at Jackson Bluff in Leon County, Florida.
This species is based on complete shells
without opercular valve, and "the V-shaped
cut at the summit of the radii, the longitudinal slit dividing the radii, and thevertical ridges transversing the radii, arethe distinctive characters of the shell".
These characters are distinct from thoseof B. sendaicus.
B. concavus alloplax Pilsbry and Olsson,
1951 [Holotype, ANSP (Academy of Natural Sciences of Philadelphia) 18869J isrepresented only by opercular valves fromthe Oligocene age of Playa Rica, Rio
Santiago, Ecuador. Shell walls without
opercular valves were described fromanother fossil locality, Posorja, Ecuador.
It is difficult whether these shell walls
belong to the same subspecies, until more
fossil specimens from the type locality
are available. The following morphological characteristics of this subspecies
are distinct from those of B. sendaicus;
scutum strongly wrinkled; tergum with "adecidedly sunken spur fasciole·'; spur
removed "from basiscutal angle by nearly
twice width of spur".
B. concavus chesapeakensis Pilsbry, 1916
[Holotype, ANSP 1143J is based on acomplete shell having scutum and tergum
from the Miocene beds of Maryland. Along and high adductor ridge in its middle
part, and lateral depressor muscle pit
having a strongly prominent bordering
ridge, parallel to adductor ridge in B. c.
chesapealwnsis distinguish it from B. sendaicus.
B. concavus coosensis Dall, 1909 [Holotype, USNM 153960J was originally described as a variety of B. tintinnabuluJn,based on four specimens of shell wallwithout opercular valves from the Mioceneof Coos Bay, Oregon. As Pilsbry (1916)
poined out, B. concavus coosensis having
oblique summit radii and no porous radiiis not related to B. tintinnabulum. Verylarge specimen attaining 50 mm in greatest diameter and the relatively narrow
rostrum typical of B. concavus coosensis is
not found in B. sendaicus.
B. concavus dallonii Davadie-Suaudeau,

742.

Miocene Cirri]Jed Balanus sendaicus

1952 is based on a specimen of shell wallfrom the Astian (late Pliocene) of AIr
Lalla Barta, Algeria. The shell surface
having ribs and striations and large orificedistinguishes it from those of B. sendaicus.
B. concavus eseptatus Pilsbry, 1924 [Holotype, USNM 352257J was described basedon complete shell wall without opercularvalve from the Miocene of Haiti. In B.
concavus eseptatus, the radii are narrower
and the orifice is larger than those of B.
sendaicus.
B. concavus finchii Lea, 1833 was originally described as B. finchii Lea from
the Miocene of St. Marys, Maryland.
Pilsbry (1930: 432) redescribed and illustrated topotype specimens. This subspecies is distinguished from B. sendaicus
by large orifice, weak growth ridges anda long, moderately well developed adductor ridge in scutum.
B. concavus glyptopoma Pilsbry, 1916
[Syntype, ANSP 1140J was describedfrom the Pliocene Caloosahatchie Formation, Florida. The rather numerous small
ribs, sometimes weak or nearly obsoleteon the surface of parietes, a strong but
rather short adductor ridge, and a short,widely obliquely truncated spur of thissubspecies distinguish it from B. sendaicus.
B. concavus oligoseptatus Kolosvary, 1961
[Holotype preserved in ColI. Mus. Inst.
Syst. Zool. Univ. SzegedJ from the upper
Oligocene of Kizil-Kum, USSR has levelsummits radii and large orifice, which are
different characteristics from B. sendaicus.
B. concavus proteus Conrad, 1834 wasdescribed based on specimens from JamesRiver, Suffolk, and Yorktown, Virginia.
This subspecies is characterized by strong
ribs on the external shell wall, and slightly
well developed adductor ridge. Thesemorphological characteristics are not present in B. sendaicus.
B. concavus raphanoides Moroni-Ruggieri,
1952 [Holotype, Cirr. 342J from the Pli-

285

ocene of Bagni of Casciana, Italy is represented by an internal mould of shellwall. The characteristic cup-shaped base,cylindrical form and wide orifice of thissubspecies are not present in B. sendaicus.
B. concavus rariseptatus Pilsbry, 1918
[Holotype, USNM 324292J is based on one
cluster of shell walls without opercularvalves from the Miocene Gatum Formation of Panama Canal Zone. The slightly
longitudinally roughened walls of characteristic of this subspecies are not present
in B. sendaicus.
B. concavus rubescens Seguenza, 1874
was originally described as a variety ofB. concavus from the Tertiary formationof the province of Messina, Italy. In B.
concavus rubescens, the growth ridges ofscutum are scarcely detectable and thelongitudinal striations on external surfaceof scutum are weak. These morphologicalcharacteristics are distinct from those ofB. sendaicus.
B. concavus scutorum Seguenza, 1876 wasdescribed as a distinct species B. scutorum
from the province of Messina, Italy. The
following morphological characteristics ofthis subspecies are different from thoseof B. sendaicus: surface of scutum withfine longitudinal striations; spur widthnearly equal to 1/3 basal length of tergum;
narrow spur furrow; distinctly prominent
adductor ridge.
B. indicus Withers, 1923 [Holotype, In.
20233J was reported from the Miocene of
Karachi, Mouth of the Indus. This species
is distinguished from B. sendaicus by rather
deeply sunken radii, narrowly spaced andweakly prominent growth ridges on
scutum, and spur fasciole in tergum.
B. polyporus Pilsbry, 1924 [Holotype,
USNM 352258J from the Miocene of Haiti
is represented by only one shell wall
specimen. The somewhat roughened wall
surface and rostrum notably longer thanthe other plates distinguish it from B.

286

Toshiyuki Y AMAGUCH I

sendaicus.
B. talquinensis Weisbord, 1966 [Holotype, PRI 27374J was described from the
Miocene Choctawhatchee Formation atJackson Bluff in Leon County, Florida.
The following morphological characteristics of B. talquinensis are distinct fromthose of B. sendaicus: small beads or
nodules are found at the intercepts of theradial riblets with the growth ridges ofscutum; adductor ridge of scutum is highand long; spur furrow of tergum is shallow and moderately wide.
B. vadaszi Kolosvary, 1949 was reported
from the early Miocene of Fels6tarkany,
Hungary. Compartments with narrow
radii and coarse white ridges on surface
and scutum without longitudinal striations
on external surface of B. vadaszi are notfound in B. sendaicus.
In conclusion, as mentioned above, B.
sendaiclls is not identical with any of thealready known living and extinct taxa of
the B. concavus group, and is surely adistinct species as pointed by Hatai et al.
(1976). However, because B. sendaicus

based on a single scutum of Hatai et al.
(1976) is not entirely identical with thatof this paper based on complete specimens
and description given by Hatai et al. (1976)
is too brief, B. sendaicus is redescribed
here.
Systematic Paleontology

Superfamily Balanoidea Leach, 1817
Family Balanidae Leach, 1817
Genus Balanus Da Costa, 1778
Balanus sendaicus Hatai, Masuda
and Noda, 1976
[PI. 44, Figs. 1-6, PI. 45, Figs. 1-9,
PI. 46, Figs. 1-22, PI. 47, Figs. 1-22,
PI. 48, Figs. 1-3, Text·fig. 1JBalanus concavus Bronn: Yamaguchi, 1974, p.
218, pI. 67, figs. 9-10.
Balanus rostratus Hoek: Hatai, Masuda andNoda, 1976, p. 14, pI. 2, figs. 1, 2, 13-17,
19-21.
Balanus sendaicus Hatai, Masuda and Noda,
1976, p. 13, pI. 2, figs. 1 (same specimen
as fig. 20), 9 (same specimen as fig. 10,
Holotype, missing), 10, 14-17, 19, and 20.

Explanation of Plate 44
Balanus sendaicus Hatai, Masuda and Noda, 1976
from the Miocene Moniwa Formation
All specimens were illustrated under the name of Balanus rostratus Hoek in plate 2 of Hatai
et al. (1976) and were deposited in the Saito Ho·on Kai Museum, Sendai, Japan.
[Hatai, et al.'s specimens are not accompanied with registration number.J
Fig. 1. 1a-b, shell wall (same specimen as pI. 2, figs. 1 and 20 of Hatai et al.) on gravel, xL
Fig. 2. 2a-c, shell walls (same specimens as pI. 2, fig. 2 of Hatai et al.) on a brachiopod shellof Coptothyris grayi miyagiensis Hatai, Masuda and Noda, 2a, xL 2b-c, x 2. 2d-k, opercularvalves preserved in the shell cavity of shall (left) [These opercular valves were not foundby Hatai et al.), x 5; 2d-e, right scutum. 2f-g, right tergum. 2h-i, left scutum [morphologically identical with Holotype (missing), pI. 2, figs. 9 and 10 of Hatai et al.J, 2j-k, left
tergum.
Fig. 3. 3a-b, shell wall (same specimen as pI. 2, fig. 14 of Hatai et at.) on a brachiopod shell of
Coptothyris grayi miyagiensis, 3a, xL 3b, x 2.
Fig. 4. 4a-b, shell wall (same specimen as pI. 2, fig. 15 of Hatai et al.) on a brachiopod shell of
Coptothyris grayi miyagiensis, 4a, xL 4b, x 2.
Fig. 5. 5a-b, shell walls (same specimens as pI. 2, fig. 19 of Hatai et al.) on a molluscan shellof Anomia sp., 5a-b, xL
Fig. 6. 6a-b, shell walls (same specimens as pI. 2, figs. 16 and 17 of Hatai et al.) on gravel, xL

YAMAGUCHI:

Miocene Cirriped Balanus sendaicus

Plate 44

742.

Miocene Cirriped Balanus sendaicus

Description. -General appearance. -Shell
conical and smooth often with weak longitudinal ribs. Color of shell ivory yellow,
frequently with very fine striations ofpale reddish purple or tinted with palereddish purple near orifice or on the shellwall. Orifice toothed.
Compartments.-Parietes very wide in
rostrum and lateral. Radii wide in rostrum
and lateral, and moderately wide in carinolateral and carina; summits oblique.
Sheath solid. Parietal (longitudinal) tubes
with transverse septa in upper half of shellwall. Parietal tubes usually secondarily
filled up with calcite as the result of fossilization. Those transverse septa usually
not found by soft X radiographs (PI. 48,
Figs. 15-19), although, they observable in
the longitudinal thin section of shell wall
(Text-fig. 1, PI. 48, Figs. 1-3).
Base. -Calcareous.
Opercular plates.-Scutum slightly concave externally with narrowly inflected
tergal segment; basal angle obliquely
truncated; external surface with prominentand crenulate growth ridges and moderately prominent longitudinal ridges frequently bifurcated by the weak longitudinal striae; nodes observable at the connection between the ridges; these ridges
circumscribing square depressions ("rather
coarsely latticed sculpture" as pointed by
Pilsbry, 1916: 101); color ivory yellow;
inner surface smooth on upper part; articular ridge moderately projected, reflexedand greater than l/Z length of tergal
margin; articular furrow narrow andshallow; adductor ridge weak or hardly
visible; pit for adductor muscle wide anddeep; pit for lateral depressor muscle wideand deep; inner surface of basal margin
faintly crenulated.
Tergum flat, moderately wide with aweakly developed beak (PI. 44, Figs. Zfg); external surface smooth with fine
growth lines; color ivory yellow, tinted

287

oText·fig. 1. Shell structures in longitudinal sections of a rostrum (UMUT
CA9686) from the Miocene Moniwa Formation (Loc. 4). Photos for Text-figs. la-b
are shown in PI. 48, Figs. 1-2. la, a parietal
tube with transverse septa in a longitudinal. section parallel to the parietal (longitudinal)
tube. lb, parietal tubes with transverse
septa in an oblique longitudinal section. I:
inner lamina, Lr: longitudinal rib, Ls: longitudinal septum, 0: outer lamina, P: parietal
tube, S: sheath, T: transverse septum.

with pale reddish purple near apex between spur furrow and carinal margin;
basal margin of both sides of spur approximately straight; spur furrow extremely
narrow in young specimens or partiallyclosed in large specimens and slightly
sunken; spur rounded at end and withseveral very weak projections on innersurface of basal margin; spur width approximately 1/3 length of bz.sal margin288

Toshiyuki YAMAGUCHIor less; distance between basiscutal angle
and anterior side of spur nearly equal to
1/3 length of spur width; crests for depressor muscle well developed; rest of
inner surface smooth.
Stratigraphic and geographic ranges,
definition of B. c. concaVllS andhabitat

The straigraphic range of each memberof the 'E. concavus group summarized byNewman and Ross, 1976 is shown in
Table 1. The oldest record is the Oligocene of Nurri, Sardinia Is. and Capo S.
Elia, Cagliari, Italy concerning B. c. concavus (de Alessandri, 1906), of Pomaz andGod, Hungary concerning B. c. concavus(Kolosvary, 1943), of Ecuador concerning
B. c. allopax (Kolos vary, 1943), of Budafork concerning B. c. chesapealcensis andB. c. glyptopoma (Kolos vary, 1943), andof Kizil-Kum, USSR concerning B. c.
oligoseptatus (Kolos vary, 1943).
The
youngest record is the Pleistocene ofMonte Mario, Roma; Civitavecchia, Palo;and Sciacca, Sicilia Is., Italy concerning

B. c. concavus (de Alessandri, 1895, 1906),
of Pisa province, Italy concerning B. c.
concavus (Menesini, 1965), and of Monastir,
Tunisia concerning B. c. concavus (Davadie
1963), etc. However, most of the fossils
were obtained from the Miocene andPliocene.
There are two critical and important
problems of stratigraphic distribution of
the B. concavus group:1) Most of B. c. concavus reported in
the literature appear only as listed occurrence of fossil and/or have only briefmorphological description without illustration. Are most of these specimens
identical with B. c. concavus in the senseof de Alessandri (1906, pI. 16, figs. 21-25;
pI. 17, figs. 1a-4b), based on reexamination of topotypes of Bronn (1831)'s B.
concavus? The writer discussed precisely
on B. concavus of Darwin (1854a, b) andof de Alessandri (1906) in the precedingline.
2) Geologic age-determination in Neogene and Quaternary were advanced bybiostratigraphy based on planktonic microfossils since 1950's and by magnetostra-

Explanation of Plate 45
Balanus sendaicus Hatai, Masuda and Noda, 1976
All specimens are topotypes from Loc. 4 of the early to middle Miocene Moniwa Formation andpreserved in University Museum, University of Tokyo (UMUT).
Fig. 1. a complete shell (UMUT CA964l). la-b, a shell wall (conical form), xL lc, four opercularvalves articulated, x 2.
Fig. 2. a complete shell (UMUT CA9642). 2a-b, a shell wall (cylindrical form), x 1.5. 2c-j, four
opercular valves, x 2.5; 2c-d, right scutum. 2e-f, left scutum. 2g-h, right tergum. 2i-j, left
tergum.
Fig. 3. a complete shell (UMUT CA9643). 3a-c, a shell wall (cylindrical form), x 1.5. 3d-k, four
opercular valves, x 2.5; 3d-e, right scutum. 3f-g, left scutum. 3h-i, right tergum. 3j-k, left
tergum.
Fig. 4. a shell wall (UMUT CA9644). 4a-b, x 1.5 (a single right scutum preserved in shell cavity).
Fig. 5. an aggregation of shell walls (UMUT CA9645), x 1.5.
Fig. 6. shell walls (UMUT CA9646). 6a-b, x 1.5.
Fig. 7. a shell wall (UMUT CA9688) on fragment of brachiopod shell. 7a-b, x 1.5.
Fig. 8. a shell wall (UMUT CA9647). 8a-b, xL
Fig. 9. a shell wall (UMUT CA9648). 9a-b, xL

YAMAGUCHI:

Miocene Cirriped Balanus sendaicus

Plate 45

742.

Miocene Cirriped Balanus sendaicus

tigraphy based on paleomagnetic polarity
in sediments since 1970's. Determinationof the stratigraphic range of B. concavus
requires knowledge of the precise geologic
age of the sampling horizons. However,many previous studies of B. concavus did
not incorporate the current stratigraphic
method mentioned above in age-determination, because they were published before the recent advance of age-determination.
Stratigraphy of Italy, based on modern
geologic views, was established after theWorld War II and thus the geologic ages
published before the World War II will
have to be compared with the advanced
stratigraphy and geologic age.
Therefore, in case where these two
important problems of identication or definition of species and geologic age are
not concurrently solved, discussion on
stratigraphic and evolutionary change ofB. c. concavus or other members of theB. concavus group should be restricteduntil the precise range of the B. concavus
group in Mediterranean region is summarized.
The fossils of this group were foundin areas surrounding the Mediterranean,corresponding to the western part ofTethys Sea (Oligocene-Pleistocene), England (Pliocene), west Atlantic coast of
North and South America (Pliocene), theeast Pacific coast from California to Mexico (Miocene-Pliocene), and east Pacificcoast of Peru (Pleistocene). One noteworthy result is, as pointed by Pilsbry(1916), that this group had flourished from
the Miocene to Pliocene and had lived in
the Atlantic and the Mediterranean region
until Pleistocene, but is extinct in thisregion ever since, though its members
continuously live on the east Pacific coastof California and Baja Cailfornia at present. The disappearance of this groupfrom the Atlantic and the Mediterranean289

region in Pleistocene is a mystery.
In the east Pacific region, after the
closure of Panama isthmus in the Miocene,
three species: B. gregarius (= B. aquila,
a junior synonym of B. gregarius), B.
panamensis, and B. regalis, live in the
California and Oregonian Provinces from
California to Peru-Chile. They are relictsof the Tethys (Newman, 1979). B. gregarius is endemic to the narrow Transition Zone (about 4 in latitude) in between
California and Oregonian Provinces (Newman, 1979). The nature of this "so-called
4 short-range endemism" is discussed indetail by Newman (1979).
Japanese fossil B. sendaicus is the firstrecord of this species group from thewestern Pacific area. B. sendaiclls werecollected from several localities of northern Honshu, Japan. The geologic age
is restricted to the Miocene.
B. sendaiclls occurs in association withextinct pectinid Chlamys kaneharai, brachiopods and bryozoans from the type
locality of this species (Loc. 4). Brachiopods are well preserved, and the majorityof them are articulated. Bryozoans havefine and reticulated structure, and arealso well preserved. Shells of B. sendaiclls
are chiefly found as fragmental, skeletal
elements, however, some of shells arecomplete shell walls and in some cases
opercular valves are preserved in their shellcavities. These facts seem to indicatethat those fossils were not transported so
far away from their habitat. Those fossils
associated indicate the paleoenvironmentof moderately strong water agitation.
The habitat of B. sendaicus is also presumed from the nature of fossil 2.ssemblage and sediments at the other fossil
localities of B. sendaicus.
00Conclusion
The description of Balanus sendaicus

290

Toshiyuki YAMAGUCHI

Hatai, Masuda and Noda (1976) was basedon a single scutum, but it could not foundout in the Saito Ho-on Kai Museum,Sendai. Study of topotype materials suggests that B. sendaicus is conspecific withshell walls incorrectly identified as B.
rostratus Hoek by Hatai et aZ. (1976). B.
sendaicus, based on scuta, terga and shellwalls, is distinct from the known taxa,
and is a good species as proposed byby Hatai et at. (1976). B. sendaicus is redescribed on the basis of skeletal elementsof the topotypes.
B. sendaicus belongs to the B. concavus
group based on having transverse septa
in parietal tubes of shell wall and thecharacteristic ornamentation of scutum
and seems to have derived from theclosely related B. concavus concavus Bronn
sensu de Alessandri (1906) which has
mainly lived in the Mediterranean regionof Italy and its adjacent areas from the
Oligocene to the Pleistocene.
Darwin (1854a, b)'s B. concavus includes
two or more taxa which are morphologically distinct from true B. concavus concavus. Newman (in press) reexaminedbased on the materials of Darwin (1854a,b), and recognized three taxa includingtrue B. concavus concavus (Newman, pers.
comm.). It is difficult to decide whether
or not extinct infraspecific taxa of B.
concavus are distinct from the other taxa,
judging from their brief descriptions andtheir incomplete assemblage of skeletal
elements. Their taxonomic positions will
be valid by the Zoological Nomenclature,
however, their holotypes and topotypes
are necessary to be reexamined from themodern taxonomic point of view.

Acknowledgments
The author wishes to thank Professor
William A. Newman, Scripps Institutionof Oceanography, who kindly read themanuscript, and provided with invaluablecomments and living American species of
the B. concavus group for comparative
study. The author is also indebted toProfessor Elena Menesini for providing
invaluable informations of the European

B. concavus.
Thanks are due to Professor Tetsuro
Hanai, University of Tokyo for reading
the manuscript and giving valuable advice.
I am greateful to Dr. Paul Frydl of ourInstitute for the reading of the draft

Explanation of Plate 46
Balanus sendaicus Hatai, Masuda and Noda, 1976
All specimens are to po types from Loc. 4 of the early to middld Miocene Moniwa Formation andpreserved in University Museum, University of Tokyo (UMUT).
Figs. 1-4. rostrum, x 1.2. 1a-b, (UMUT CA9649). 2a-b, (UMUT CA9650). 3a-b, (UMUT CA9651).
4a-b, (UMUT CA9652).
Figs. 5-S. lateral, x 1.2. 5a-b, left, (UMUT CA9653). 6a-b, left, (UMUT CA9654). 7a-b, left,(UMUT CA9655). Sa-b, right, (UMUT CA9656).
Figs. 9-12. carinolateral, x 1.2. 9a-b, left, (UMUT CA9657). lOa-b, left, (UMUT CA965S). lla-b,
left, (UMUT CA9659). 12a-b, right, (UMUT CA9660).
Figs. 13-16. carina, x 1.2. 13a-b, (UMUT CA9661). 14a-b, (UMUT CA9662). 15a-b, (UMUT
CA9663). 16a-b, (UMUT CA9664).
Figs. 17-19. scuta. 17a-b, right, (UMUT CA9665), x 3. lSa-b, left, (UMUT CA9666), x 2.5.
19a-b, left, (UMUT CA9667), x 2.5.
Figs. 20-22. terga, x 2.5. 20a-b, right, (UMUT CA966S). 21a-b, right, (UMUT CA9669). 22a-b,
left, (UMUT CA9670).

YAMAGUCHI:

Miocene Cirriped Balanus sendaicus

Plate 46

742.

Miocene Cirriped Balanus sendaicus

291

Locality

Text-fig. 2. Sampling localities of fossil
Balanus sendaicus. Details for each site
are given in the section Locality.

manuscript. I am indebted to Dr. Koichiro
Masuda of Miyagi University of Education and Dr. Sadako Takeuchi of the Saito
Ho-on Kai Museum for lending the typematerials of Hatai, Masuda and Noda (1976).
Thanks are also due to Dr. Junji Itoigawa
of Nagoya University and Mizunami Fossil
Museum for providing fossil specimens of
the Mizunami area. This study was supported in part by Grant-in-Aid for Scientific Research from the Ministry of Education, Science and Culture, project no. 434042.

1a-b.-Obuchi: a) cliff (40 05S'55.1"N, 141°
21'42.9/1E) along the Oippe river, situatedabout 2.S5 km N26°W of Obuchi, Rokkashomura, Kamikita-gun, Aomori Pref., crosslaminated coarse shell sand ("coquina")., b)
cliff (40 05S'55.1/1N, 141°20049.3/1E) along the
Oippe river, situated about 3.55 km N44°W of
Obuchi; cross-laminated coarse shell sandstone("conquina")., a-b) late Miocene Takahoko
Formation.
2a-c.-Ohmagari: a) shell quarry at Arasawa(39°24'16.2/1N,
140 021'4.2/1E), Minami-Sotomura, Senpoku-gun, Akita Pref. ; barnacle shellsand ("coquina")., b) cliff (39°21'34.l"N,140021'42.1/1E) at Asizawa Pass of Takinoue,
Ohmori-machi, Hiraga-gun, Akita Pref.; massive poorly sorted tuffaceous sandstone containing carbonaceous matter., c) cliff (39°
22'6.5"N, 140021'42.1"E) of left bank of creaksituated about 1 km north of Loc. 2b; massive
poorly sorted tuffaceous fine sandstone containing carbonaceous matter., a-c) late Miocene
Sugota Formation.
3.-Ukibuta: cliff (39°1S'47.S"N, 140°20'41.1/1E) situated about 100 m east of Ukibuta Tunnel,
Ukibuta, Higashi-Yuri-mura, Yuri-gun, Akita
Pref.; poorly sorted, tuffaceous, coarse sandstone; late Miocene Sugota Formation.
4.-Sendai: cliff (type locality of B. sendaicus) (3S012'45.4"N, 140 047'26.9"E) situatedabout 200 m south of the Oide Bridge acrossthe Natori river, l\1oniwa, Sendai City, Miyagi
Pref.; poorly sorted, gravelly, calcareous,
medium. shell sandstone; early to middle
Miocene Moniwa Formation.
5.-Nanao: cliff (37"2'23.5"N, 136°57'23.4"E)situated about 0.95 km NS5°W of Nanao Station, Iwaya-cho, Nanao City, Ishikawa Pref.;cross-laminated conglomeratic coarse shellsandstone containing many bryozoans; late
Miocene Nanao Calcareous Sandstone.
6a-b.-Mizunami: a) cliffs (35°23'32.5N",
137"16'35.6"E) situated about 3.5 km 1\32°E of
Mizunami Station. Dan, Mizunami City, Gifu
Pref.; conglomeratic very coarse sandstone.,b) cliff (35°22'25.7"N. 137°16'2S.6E) situatedabout 1.95 km N61°E of Mizunami Station,
Sakurado. Mizunami City., a-b) late Miocene

Toshiyuki YAMAGUCHI292
Oidawara Formation.

References
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Balanus sendaicus Hatai, Masuda and Noda, 1976
All specimens are from various localities of the Miocene and preserved in University Museum,
University of Tokyo (UMUT).
Fig. 1. Specimen from the late Miocene Takahoko Formation (Loc. 1). 1a-b, left scutum (UMUT
CA9671), x 2.5.
Fig. 2. Specimen from the late Miocene Sugota Formation (Loc. 2a). 2a-b, left scutum (UMUT
CA9672), x 2.5.
Figs. 3-5. Specimens from the late Miocene Sugota Formation (Loc. 2b). 3a-b, left scutum (UMUT
CA9673), x 4. 4a-b, left scutum (UMUT CA9674), x 4. 5a-b, right scutum (UMUT CA9675),
x4.
Figs. 6-10. Specimens from the late Miocene Sugota Formation (Loc. 2c). 6a-b, left scutum(UMUT CA9676), x 3. 7a-b, right scutum (UMUT CA9677), x 4. 8a-b, left tergum (UMUT
CA91378), x 4. 9a-b, left tergum (UMUT CA9679), x 4. 10a-b, right tergum (UMUT CA9680),
x4.
Figs. 11, 12. Specimens from the late Miocene Nanao Calcareous Sandstone (Loc. 5). 11a-b, rightscutum (UMUT CA9681), x 4. 12a-b, left scutum (UMUT CA9682), x 4.
Figs. 13, 14. Specimens from the late Miocene Oidawara Formation (Loc. 6b). ColI. Mizunami
Fossil Museum, Mizunami, Gifu Pref. 13a-b, left scutum (UMUT CA9683), x 2.5. 14a-b, rightscutum (UMUT CA9684), x 3.
Figs. 15-19. X-radiographs of to po type specimens from the Miocene Moniwa Formation (Loc. 4),
xL 15, carina (UMUT CA9661). 16, rostrum (UMUT CA9649). 17, rostrum (UMUT CA9651).
18, rostrum (UMUT CA9652). 19, left lateral (UMUT CA9653).
Figs. 20-22. Interlaminate figures observed in thin section parallel to the base of shell wall (Loc.
4). 20, carina (UMUT CA9685). 21, rostrum (UMUT CA9686). 22, left lateral (UMUT CA9687).

YAMAGUCHI:

Miocene Cirriped Balanus sendaicus

Plate 47

21

22

20

742.

Miocene Cirriped Balanus sendaicus

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293

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[in Japanese].
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Balanus amphitrite Darwin group (Cirripedia, Balanomorpha) from the late
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Bioi. Bull., vol. 127, no. 2, p. 360.
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Diego Formation, San Diego County,
California. Contrib. Sci. Los Angeles Cty.
Mus., no. 159, p. 1-25.

Explanation of Plate 48
Shell structure in longitudinal sections of a rostrum (UMUT CA9686) from the iVliocene
Moniwa Formation (Loc. 4).
Fig. 1. a parietal tube with transverse septa in a longitudinal section parallel to the parietal(longitudinal) tube. la, x 8.4. 1b, x 10.2. 1c, x 42.
Fig. 2. parietal (longitudinal) tubes with transverse septa in an oblique longitudial section (fourparietal tubes are found). 2a, x 5.8. 2b, x 10.2. 2c, x 42.
Fig. 3. parietal (longitudinal) tubes with transverse septa in an oblique longitudinal section. 3a,photos by an optical microscope of Replica film (cellulose acetate film), x 10.2. 3b-e, external
and internal walls (correspond to the outer and inner laminae) of parietal tubes and transverse
septa partially left the original shell structure. Interspace of the parietal tube is secondarily
filled up with the calcite which contrasts with the calcite composing the transyerse septa ingrain size, shape and arrangement. Photos by scanning electron microscope (SEM) of theetched surface. 3b, enlargement of top of fig. 3a, x 42. 3c, ditto, x 128. 3d, ditto, x 290. 3e,ditto, x 728.

YAMAGUCHI:" • , .

• 0

. /,11 . ,.

Miocene Cirriped Balanus sendaicus

Plate 48

74'2.
-

Miocene Cirriped Balanus sendaicus(1979): Thoracican Cirripedia of theLower Pliocene Pancho Rico Formation,

295

Salinas Valley, Montery County, California.
Ditto, no. 303, p. 1-l3.

Obuchi j'i§,'lbt, Oippe oIttm, Takahoko I:~, Ohmagari :leB±!, Arasawa m/R, Asizawa F/R,
Takinoue i"ri1 / j:, Sugota ®i~EE)' Ukibuta i'¥~, Sendai {wii', Oide ~I±l, Natori 1;jfz, Moniwa
rJ1:&!, Nanao -Gj'i§, Mizunami Jlffi1.f.!, Dan B):, Sakurado :ft<¥:. Oidawara ~~Epjs:O)<=p~til:§e)]1$m Balanus sendaicus Hatai, Masuda and Noda 1:''?\''l:-77- ;7..(0) Balanus concavus flM c JtiB( L l : - : Balanus sendaicus t:t fwii'ftiBJ=-:B-::(P-t0<=P~til:a)}!~iJ'i?~i?nt-:%\!:0)-tm:B-0) 1 '?O)ttllJR (scutum) 1:'~--:)\"l::I:IH#. :f:\)iEB. !!!.fEB(1976) I:.J: I)~flc Ll:gc~~nt..:o ..:c0)f,\'?;;jq11wii'0)J?r~¥jt'6?,:ftj-\\t#Jtrnf:.f!i';:"i1f~nl:\"0,:t--rt-=iJ~ . ..:c':'I:'f:tJil.'?vti?ntJ:iJ'...,t-: o ..:cO)1;lltro){, cl:.ffi!:O)(iliO)tm:B-. 1rJR (tergum):toJ:lf)l!(J%\!: (shell wall). 'Hcf(~nt.:iJ'...,t..:o ]jj(iE~f,\'?* (to po type) :toJ:lf:I:IH#i? (1976)
1:.lZIlF~nt-:f,\'?*~1E-c-. :I:IH#i?tJ: B. rostratus c Ll:Bc~ ·lZIlFLt-:)l!(J%\!:I:t B. sendaicus
I:.~-to ..:cnt1:>;t. *~~Jt-ef:t:I:IH#i?I:'J: ...,l:~c~~nt.:tJ'...,t-:1rJR~1!;col: B. sendaicus~OOIi'{H:' -t 0 0 B. sendaicus f:t:I:IH# i? 0) j'1'i:f1lli 0) c ? I) J;t9;o0):B-mMtJ, i?!JJ.: 0 0 B. sendaicus
f:tiWi~lJttJ, i? J':~1Jt1:'~<=P$~~7 - 7- ;7..$v:,~* U-: de Alessandri (1906) O)~,*{n '-c0)~~:fH! B. concavus CJli~-C'~00 B. sendaicus ,:t)l!(J%\!:0)~"i1f (longitudinal it..:l:tparietal tubes) 1:.jj~E~ffiU~~ (transverse septa) ~M''?.:. c:to J: lfttll*-&O)*~BJtdHm~
tiffiv:.J:...,l: B. concavus v:.ft~~n01iIiMv:.~-to .:.O)flMv:.Jm-t "4 1ll!" tJ:*tm:t::qzi$l=~}&!, Ll:\"0 (Newman, 1979) vtnc'{":lel1§I'F. ~<=P$~~iJ'i?O) B. concavus flMO)r~~f:t Pilsbry (1916) 0) j'1'i:f1lli 0) J: ? I:. ~ ;7.. 7 Y 7;7.. -C'~00 {tE'O) B *0) B. sendaicus
l:t. <=P~til:v:.llNi?n. B. concavus :fH!MO)*7:;7tJ'i?0).ft:m0){tE'~cttct.:I) • ..:cLl:7- 7-;7..$ c *7:; 7 O)rs'lO)'?t.:tJ~ I) O)ff-tEHI'i:f1lli-t 0 0
LlJ J:J ffZ

Postscript: After submitted, Saito Ho-on Kai Museum of Natural History (SHM) let me know on registration numbers of illustrated specimens
in the plate (pI. 2) of Hatai et al. (1976). The registration numbers are as
follows.
species namefigure inspecies name inRegistration numberin this paper
Hatai et al.
Hatai et al.
SHM, Reg. No. 22069
figs. 1 & 20
Balanus rostratus
B. sendaicus
B. oidensis (Holotype)Not studied
22070
figs. 3 & 4
22071
22072
22073
22074
22075
22076
22077
22078
22079
22080
22081
22082
22083

figs. 5 & 6
figs. 7 & 8
figs. 11 & 22
fig. 12
fig. 18
fig. 23
fig. 2
figs. 13 & 21
fig. 14
fig. 15
figs. 16 & 17
fig. 19
figs. 9 & 10

B. moniwaensis (Ditto)
B. natorianus (Ditto)
B. miyagiensis (Ditto)
B. tohokuensis (Ditto)
B. rikuzenensis (Ditto)
B. sp.
B. rostratus
B. rostratus
B. rostratus
B. rostratus
B. rostratus
B. rostratus
B. sendaicus (Holotype,missing)

Not studied
Not studied
Not studied
Not studied
Not studied
Not studied
B. sendaicus
B. crenatus
B. sendaicus
B. sendaicus
B. sendaicus
B. sendaicus
B. sendaicus

296

Palaeontological Society of Japan Special Papers No. 24
Permian Conodont Biostratigraphy in Japan
By Hisaharu IGO
Issued December 15, 1981, 50 pp., 12 pis.

Price (postage and handling incl.) U. S. $24.00.
Clark and Behenken (1971) summarized "Conodont and biostratigraphy of thePermian" in Symposium on Conodont Biostratigraphy edited by Sweet and Bergstrom.
In this paper, they mentioned that Permian System represents the "last frontier" forconodont research. Permian conodont study was the same situation in japan.
About ten years ago, Hisaharu Igo initiated to study Permian conodonts and hesucceeded to recover numerous specimens from the various sections which were dated
by fusulinaceans mainly in central part of japan. The author clarified stratigraphic aswell as geographic distribution of many important species and compared with the rangeof the previously described species in North America and Europe. Conodonts collectedfrom chert and compound-type are mostly excluded in this volume, but he proposed
two conodont faunas and two assemblage zones in the japanese Permian and described
44 species and subspecies from 14 genera, among which 10 species and 1 subspecies
are new to science.
This volume would be indispensable for geological and paleontological libraries aswell as workers who may have interests in conodontology and later Paleozoic to early
Mesozoic historical geology.
Orders must be accompanied by remittance made payable to Dr. Tsugio Shuto,
Editor of the Special Papers, c/o Department of Geology, Faculty of Science, Kyushu
University, Hakozaki, Fukuoka 812, japan.

297

PROCEEDINGS OF THE PALAEONTOLOGICAL SOCIETY
OF JAPAN

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Subcommission on Devonian Stratigraphy (7116-7129, New York State) ........ ~lTI~&±
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MEBI2B&~· :W:~lil:;R:

Bi!E* • ~~':::*1t~~J't , .....

.............. re7klt5C· *~*1!Jj. WJ1*~7k

Mesozoic floras in South America"" .. ,.,.
,., ,.,'.,., ... ,',." .. ,"""" ,Kimura, T.
*~~B.illi:~M~~m*~m1:M~cO)ft~R*4 ................................ l!§lTI ~7- Y 1'm00~':::*c Nothojagus &;;!t1t:P0)~~IU::~I:U~$... ,"",., ..... ,.", .... ,Wl11=~~i::"7mffll31 ~/, (Tarija) ~~O)~I2B*clTflJ:JL!!1J!J&J1t::r:I O)~:l* .,', .. ,"""',."." ...
:'G~*~. ;f'ii;lt-~=::. • Arozqueta, B.

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298!J ~ /'1t:ltMeiJillUlli~Lm{t::fi ............ ~#~=q,i¥i*0){t::fifl.mVc::.~-t0=· -=:O)r.~m!

..... .

.................................. ± ~Jg-

Jurassic trigoniids from Peru ............. .
... . Maeda, S., Kawabe, T. & Rangel, C. Z.
Some Jurassic Bivalvia from the Domeyko
Range of North Chile ................... .
· ... Hayami, I., Maeda, S. & Chong, G. D.
Stereostylus maedai, a new Permian coralfrom Colq uencha, Bolivia .......... Yamagiwa, N., Mukumoto, M. & Urdininea, M.
Some interesting Carboniferous fossils from
Huancavelica, Central Peru ............. .
Yamagiwa, N., Ishii, K. & Villavicencio, E.
7~TAq,$~~0)~·q,~WO)~~~~~~~~-~~~]t¥a~--..................... .
· ......... :tlLl:.i1t1<: • tmImff- • :I.mitliR~ •:IHJ:!¥:- • Rangel, C. Z.• Urdininea, M. R.
Copacabana ~llf (=~*) O):!lWfffivc::.-:>l'L ..
.................................. :Ilt#!¥:7 ~ T A q, $ Abancay ~t 15 ~ ~ LihuircoQuisuar )v- rO)(:~£E.{t::fi~~ .... :I;&:J:i1t:;!;:",,)v • ;f ~ 1:0 7 £Lower Permian JliiiJEJ\fjVc::.
-:>l'L ............................ iliP III ff-

Paleoscenidiidae O)7tm~tLJi: c NassellariaO)il{tVc::.~-t0~~ .. ¥r;fi~n]. q,t!!:~¥?XI'l~
Stylosphaerids O)7Dl:i\'.\'m:Vc::.-:>l'L ........... .
· ................... jffi;fillJl'f· q,1!l:J!1¥?XI'l~~]IHlJ~~~)(:k;tf"<!J ~O)f,',&{t::fi c~~Bi'j-fI::· ..... , ....... mlIllmlJ3· mtiR Dt· fti:~ IE-O):fr~ (::fiER-=~*) ~J:$O):fffLEI;{t::{5~~O)~HIll ............... . 'ti:~~~T· ~IDIl7-.~
On the Upper Permian Palaeojusulina-Colaniella fauna from Khao Doi Pha Phlung,North Thailand .. Sakagami, S. & Hatta, A.
m£!;~'jPfm~tjffi$O) JEiR~O) J:$B!lli:7forU'H1:fffLEI;llf ................ ~III~T • *~'ffiIE
Biostratigraphy of the Oligocene-Miocene
sequence along Kali Oyo, east of Yogyakarta, central Java ........ ; ............ :· ..... Saito, T., Kadar, D., Niitsuma, N.,
Matsuoka, K., Kotaka, T., Tanaka, H.,
Harada, K., Sudijono, A. F. & Hasibuan, F.
Paleogene Foraminifera from Hokkaido,
Japan. . . . . . . . . . . . . . . . . . . . . . . . .. Kaiho, K.
)(~ O)!lillfiTt!!:i'¥~i1:fffL£E. .. il1iitmIl$t.' • 'ti: III j\!ij1i:)(01; 0) !lillfiTt!!::k%:ffiLEI; .... :f'~1L ffilJl.lil.· j'SjiliPI$t.')(£!;0)!lillfiTt!!:::fiW(~7 ~ / 7' '7 ~" r ~..... .
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Wall ultrastructure of cassiduline Foraminifera ............................ Nomura, R .
Internal structure of cassiduline Foraminifera. ............................... Nomura, R.
J=l*1iaO)~t!!:JI£~:fffLEI;llf~O) %'l<c.:cO) il:9iJjJ\. ................................. i'I'J:fi!1;f*~~~~~:ffiL£E.mO)f,',&'J\~r",17t;:fj1vc::. -:>l 'L ....
· ...................... '" ~U! i$· ::fiJI I
7J • 'H'rFa* • ~*!¥:14fj • ±Il! *.~!Jlf "jl=Remarks on ostracodes from the Ananai
Formation ...................... Ishizaki, K.
Some Neogene ostracodes from the Ryukyus. ............................... Nohara, T.
Ultrastructure of the ostracod carapace, VI.
Parabolic pattern ................ Okada, Y.
7:; -;;;fm Tetraclita squamosa O)-=: "!lli:f!J!"
O)f!J!rl3'~1* ('f¥a) .................. Il! PffZ
Outline of the Permian trilobites in Japan.............. Kobayashi, T. & Hamada, T.
Genetic control of septal numbers and species
problem in a solitary scleractinian coral· . . . . . . . . . . . . . . . .. Mori, K. & Minoura, K.
Alcyonarian spiculite Vc::.-:>l'L ..... .IH1!lOO=~W£!;O) • • ::{5w(~~~£WLk.~~~~o):IPlM:¥fI:: .......................... :k;fiW3Mt'7 '7 ~*7~:¥fl::i~.I15E*0) Fa~i1Ic::.-:>l'L-USIP
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W~~70)~~KU~05E@~~~~~70).

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· . . . . . . . . . . . . . . . . . . . . . .. :f'~*JtI'l~. t.'t'~~~Jg
Growth rate of Leioceras opalinum, a Jurassic
ammonite ., .. Hirano, H. & Kennedy, W. J.

Nautilus macromphalus~{t~~~iM'O)g~7Dl:O)i\'.\'m:c.:cO)............ 3f!Jlf~J!! • fbJlII1¥~~r~-/~~~~ff10)~.~.~~0).~%~Vc::.-:>\"L.............. mlIl1¥~· 3f!Jlf~ill~3"ff1mO)%~0)~J'!ll.~~Ac~~~~~.................................. J!7Jc
Selected bivalve fossils from the Cretaceousin the Sakawa Area, Shikoku ........... .

jt

299
................ Tashiro, M. & Matsuda, T.
Nipponopecten akihoensis (Matsumoto) IC--:>l'L ..
· " ............................... {tcji:~~
alE Jll\l~Ft 11:150) rS1:.4ta~B'9 Wf5E III-Placopecten nomurai Masuda IC--:>l'L ....... .
· ................................. {tcji:~~--:>l'L .............. fi t.lHYijU (Gu Zugang)alE~~J'l.1I:EO) rS1:.4ta~i¥JliIf'JEII -~*~~.~a.~~~WLk~~~MAAO)W
fffiifiI.J4ta1l:E .. :ttfi JII ~,qp • 1t#-1Y"fl~ • 1JQifJi~~
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ji[. Ej31f.f mNm

A squalodont whale from the Miocene Ashiya
Group, Kyushu ................ Okazaki, Y.
rrJ'r~~~ Mesocena matuyamai n. sp. 0)1'Iii:

i:tz.S<J5f:;(Plc--:>l'L ....................... .
.............. 'J,;f*r!,.~§f:I· 'HHfI~ . rSlTIi3l:x
Coccolithus pelagicus (Wallich) Schiller in
Plio-Pleistocene time ............ Nishida, S.

..................... .
· ............. 'J'Jx 1'&. iJ'!l1;)i3l:g: • 'i!i'ffiHJf-=*rlli(J\lIillf4tO)Iti%llf~

Fossil pollen grains from the Miocene
Takaya plant-bed in Noto Peninsula,
Central Japan ........ Fuji, N. & Kawai, A.
~l:ifiJJf,iJIHfIl 0);1:JT~ -= * 0) fEflj ~ If ...... {tcji:Wt< i'fj
Review of the Pleistocene Shiobara Flora in
Tochigi Prefecture ................ Onoe, T.
EI ;'f·(~;1:JTi!to) 7>j-~0)5fmlc --:>l 'L .... i¥J*~g:
The past distributional history of the genus
Acer in the northern Pacific region ..... .
................................ Tanai, T.

1lD~::&~~ : Early

Cretaceous Gastropoda fromthe Choshi District, Chiba Prefecture,
Central Japan. Trans. Proc. Palaeont. Soc.
Japan, N. S., No. 118, pp. 291-324, pis. 3436, 1980.
*~~X'l T'!.(H'F,~~Y~~O) rtfil~~*}JHJ!llf~~~LkD~m~EK~~~2~0)~.~K~~
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~~~~~k~O)mR~%.
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iltWB<Jts:.tfl.illY~'G,

5fm~B<JWf5E~j1f;~,..
· . . . . . . . . . . . . . . . . . . . . . .. #JJj#-~m· Mttfpg:
III I] !!f-~1ifJ'l'Iii:llf~ " (J -If-mO)-;1:JTlI ......... .
· . . . . . . . . . . . . . . . . . . . .. FI;Jji:i&(~Ur>· i%'imXt.(t:\!1I:j;§*IPJ~llf4to) Neocalamites (iFffl'im) f,:n~· .. .. .. .. .. ... *ti~§f:I· ~ ill\:t:';J. *1tR:yNerithopsidae 0) 1 ~~ 2 ;1:JTlI~iffW:;/; ~ Aporrhaidae 0) 3 ~11~ (3trll~12:fil!0) ni1fEm1tEO)

Coreanophylium variifolium gen. et sp. nov.
-a Mesozoic bipinnate "Pterophylium",from the Daedong Group, Korea ....... .
· ................. Kimura, T. & Kim, B. K.

~±wc LLl'~~;,

IllCI".Itil~~~*ltIl;f*~~O)#1I:Elc--:>l'L-C';/; ~ c*:fIl~-=*~O) 7 :;/~ :/T~0)7;,-mtC--:>l'L....
· ............. *15 iIiZ· ggm IDI<. ElJ4t:fll$~jJ.Ht*1:.:/ 31"0)*, Cyathocaulis Ogura 0)
5fm .............................. gs Ill¥ilx~UIi.iJ:l[~.--... ::{H*1:.:/ !/O);1:JTJm Protosphaeropteris 0)*m7C~i¥J;0:!& .............. gsElJ¥ilX
Protocedroxylon triassicum Yamazaki et
Tsunada O)*tfil,*WJijJ(lc --:>l 'L ........... .

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A) Ocean floor, continental margins and exotic terrains, B) Palaeogeography of sedimentarybasins, C) Eustatic events and their correlations, D) Ocean currents and climatic events, E)
Floral biostratigraphy, G) Macrofaunal biostratigraphy, H) Supra-regional standard zones, I)
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Transactions and Proceedings of the PalaeontologicalSociety of Japan

April 15, 1982

New Series No. 125

CONTENTS
TRANSACTIONS
740. KANIE, Yasumitsu: Cretaceous tetragonitid ammonite jaws: a comparison
with modern Nautilus jaws ............................................ 239
741. KIMURA, Tatsuaki and TSUJII, Masanori: Early Jurassic plants in Japan
Part 4 ................................................................ 259
742. YAMAGUCHI, Toshiyuki: Japanese Miocene cirriped Balanus sendaicus : a comparison with Tethyan Balanus concavus group...................... . . .. 277
PROCEEDINGS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 297