ISSN 0031-0204

fJ~'~i.~~~~:k·!c.fTransactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 128

Palaeontological Society of Japan
December 30, 1982

ltaru HAY AMI and lkuwo 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, lkuwo OBA TA, Tsunemasa SAITO, Tsugio SHU TO.
Yokichi TAKAYANAGI, Toshimasa TANAIMembers of Standing Committee: Kazuo ASAMA (Finance), Kiyotaka CHINZEI (General
Affairs), Takashi HAMADA (Membership), ltaru HAYAMI (Transactions), Hisayoshi IGO (Foreign Affairs), Tatsuaki KIMURA (Planning), Ikuwo OBATA(Transactions), Tsugio SHUTO (Special Papers), Yokichi TAKAYANAGI ("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. 128, pp. 421-432, pI. 66, December 30, 1982

A STORY OF THE BELEMNITE-DEVELOPMENTAL
APPROACH TO FOSSIL MORPHOLOGY*(Presidential address, Palaeontological Society of Japan, 1981)
TETSURO HANAI
Geological Institute, University of Tokyo, Tokyo 113

The addresses that the past presidentsof the society have delivered were, inmany cases, elegant reviews of taxonomic
and phylogenetic studies on a certain taxon,
directed toward explaining the relation
between their own work and the advancement of knowledge on that taxon. Kobayashi (1960) presented a review of phylogenetic studies on trilobites, and Matsumoto (1975) on ammonites. However, because I am not in position at present todiscuss the phylogenetics on any taxon, I
would like to reminisce about my ontogenetic studies on a belemnite.
I started my study of paleontology withfossils from the Cretaceous deposits of
Miyako district, Iwate Prefecture. Since
then, more than 30 years have been passed.
Of course, I did not devote all of theseyears to the studies of this local fauna.
Yet, I reflect upon the considerable timethat I have spent toward understanding
this paleontological problem. The generalidea at that time was to make an accurate
and precise geological survey and to collectas many fossils as possible. It seems tome that the general trend in thinking atthat time was that the more fossils collected, the better the scientific results.
Specimens, thus collected during one fieldseason, were too much to cope with in
one year of laboratory work and untreatedspecimens had to be left behind year afteryear. It was an odd time when researchfunds tended to be given to those who
would remain in the field the longest.
However, I had to describe these specimens in one way or another, and decided
to commence my study with the description of belemnites. But I soon noticed
that it was an extraordinary, if not impossible task, to describe all of these specimens, which included foraminifers, bivalves, ammonites, gastropods, corals,echinoids, crinoids etc. I was forced tochange my approach. Instead of starting
the descriptive study, I let my thoughts
become confounded about the magnitudeof the project and about what type of a
product would result from such a descriptive effort. Even if I was lucky enoughto find a species that moderately changed
its morphology during the time of thedepositional sequence, and could trace thechronological variation in that species, I
still wondered if we might not be far from
elucidation of the paleontological problem.
As Makiyama mentioned in his study on
the phylogeny of Umbonium, boasting
about one's phylogenetic tree is similar to
boasting of the shape of a tree in his
garden, others are at a loss about what
to do, except for just praising it (Makiyama, 1935, p. 253).

* This is the English translation of the presidential address delivered at the Annual
Meeting of the Society, held at Tohoku
University, Sendai, on 23 January, 1981.

421

422

Tetsuro HAN AI

When I look back upon those days, I
think that the general belief was thatgeneralizations will surface if enough solid
facts are accumulated. It was this approach used by most paleontologists that
Kimura ironically criticized. He concluded
that they are virtually the same 2.S theperson who, when standing at a street
corner and looking at the moving car
passing by before him, believes he canunderstand what mechanisms make the
car run (Kimura, 1958, p. 124). It was along time later before I understood thatthe old inductive philosophy of Francis
Bacon, which underlay the thought of
paleontology at that time, had led to myproblems.
When I was reading Carr's "What .is
history?", I encountered a situation similar
to that of paleontology in the explanationof historical philosophy in 19th century.
That was "the belief of fetishism of facts",
which might be expressed as "facts speak
for themselves (Carr, 1961, p. 16)". He
explained that it is this belief "which during the past hundred years has had suchdevastating effects on the modern historian, producing in Germany, in GreatBritain, and in the United States, a vastand growing mass of dry-as-dust factual
histories, of minutely specialized monographs of would-be historians knowing
more and more about less and less, sunkwithout trace in an ocean of facts (Carr,
1961, p. 15)". Replacing the words "historians" with paleontologists and "factual
histories" with descriptions of species, I
was shocked to find the situation quite
similar to what I was about to do withbelemnites and ostracods. Therefore, in
the introduction for the checklist of Japanese ostracods (Hanai, 1977, p. 1), I emphasized the necessity for reflection onthat part of Carr's passage quoted above.
Last year, when Gould (1980, p. 96)
discussed the promise of paleobiology as"a nomothetic science based upon evolutionary theory", one of the impediments
to paleobiology that he pointed out was
overdominance of inductive philosophy inpaleontology. Another was easy-going
extrapolation ism in which biological theory,founded in neontology, is extended rashly
into the past to reexemplify the theory
utilizing imperfect fossil records, and thenconclude that the same rules hold true
there. In connection with the latter impediment, I also pointed out that "it isrelatively easy to extrapolate biologicaltheories (e.g. the theory of speciation) speculatively into the geological past, but it isdifficult, if not impossible, to elucidate withfossil evidence the paleobioiogically significant historical events, which may becomplicated, but which may interpreted interms of biological theories (Hanai, 1977,
p. 85)". At the same time, I was at aloss as to what approach we should taketo find paleobiologically significant events.
Then, I realized that the relation between the facts and the historians explained by Carr is quite similar to thatbetween paleontological facts and paleontologists. On this relation in human history,
Carr mentioned that "the historian starts
with a provisional interpretation in thelight of which selection has been madeby others as well as by himself. As he
works, both the interpretation and theselection and ordering of facts undergo
subtle and perhaps partly unconscious
change, through the reciprocal action of
one or the other (Carr, 1961, p. 29, 30)",and drew the conclusion on "What is
history?" that "it is a continuous processof interaction between the historian andhis facts, an unending dialogue between
the present and the past (Carr, 1961, p.
30)".
Further, Carr, explaining Croce's "contemporary history", argued that "history
consists essentially in seeing the pastA story oj the belernnite
through the eyes of the present and in
the li ght of its problem , and tha t the main
work of the historian is not to record,
but to evaluate; for, if he does not evaluate how can he know what is worth
recording? (Carr, 1961, p. 21 )" . From thisargument, it seems clear that an interpretation by a historian stands on theevaluation of the facts us ing all his knowledge. Therefore, it occurred to me thatthe task of paleontologists might be started
with the interpretation of fossi l data
through the present eyes of the paleontologist himself. However, my knowledgethen did not give me confidence to proceedwith this approach.
At fi rst, I took up belemnites for study,
but because of their characterless, bulletshaped morphology, soon the morphologicalcharacteristics to describe them were exhausted. Further and fortunately , there
was only one species that I could identify
with satisfaction. If I had taken up ataxon other than belemnites, which includes
many species with variety of morphology,
I might have been busily occupied w ith
the description of the species. I completed
the description of a species and I gave it
a new name, Neohibolites miya/wensis.
One of the specimens used at that time isshown in PI. 66, Fig. 1. Its only conspicuous
character that can be described is a ventral furrow. How far the furrow extends
adapically is considered to be a diagnostic
character of species and genus. I fe lt it
rather strange to give a character of
unknown biological significance an important taxonomic value without somethought. So, a question, what is the biological meaning of the ventral furrow ,occupied my mind . In this case, the needto eva luate the facts became pressing, as
I had only anthropocentric knowledge
based on my own experience; for examp le,
the electric wires and gas pipes indispensable for our modern life are to be423

protected in a furrow, an acute spine issuitable for thrusting, and a long timestanding at attention is quite uncomfortab le ... etc. Thus, I was satisfied w ith therestoration of a belemnite by Stevens(1965, p.48) where he positioned an arteryand a vein in the ventral furrow. It wasan anthropocentric and unsophisticated interpretation.
However, if that is the case, the furrowalone is sufficient to protect the veins.
Yet, when the belemnite is split into twopieces longitudinally along the medianplane, a smooth and fiat surface is foundbetween the alveolus and ventral furrow,
whereas the rema inin g surface is rough .
The presence of the commissure or s lit
between the planes under the ventral furrow concerned me as I interpreted thebiological meaning of the furrow . The
commissure was first described by Shvetsov(1913, p. 65) and because it was observable
in Neohibolites from Mi yako, a description
was given, leav ing aside the question ofits biological significance (Text-fig. 1).
One can give a description of the specimen
in his own way, even if he knows nothing
about it.

T ex t-fig . l. Sli t plane of Ne oh ibo lites
miyallOens is Hanai , 1953. x 3. MM2541

424

Tetsuro HANAl

After some effort had been made to
overcome Stevens' interpretation, I began
to make interpretations based on theknowledge of the living organisms thathave a close relationship to the fossils.
In other words, it was an effort to havea critical eye for the evaluation of thefacts. But, because the neon to logicalknowledge has not always been accumulated for understanding of the ancientorganisms, many of the results of neontological studies are little use to paleontologists, or paleontologists are often incapable of utilizing neontological knowledge.
Consequently, paleontologists began tomake observations and experiment withliving organisms, and began to study theneontological literature on pertinent subjects, in order to obtain knowledge andability applicable to their own purposes.
It may be important to emphasize that
paleontologists normally have never looked
about for fossil characters with an eye toapplying or enforcing known neontologicaltheories.
Gustomesov (1975, p. 320), who observeda thin and low ridge running longitudinally on the ventral surface of the squid's
mantle, considered it a remnant of a cariniform fin, and further related this knowledge with the facts of the past, that is,the ventral furrow of the belemnite
overlying the commissure. The explanation
was that the slit between the planes wasfilled successively as the rostrum grew,
and a deposit was secreted by the thinprocess of the mantle extending into theslit. It has been well established since
Naef (1922, p. 213-221) that the lateralfurrows of the belemnite are the scars of
the attachment or insertion of the muscleof the lateral fins. This interpretation
was thus extended into that of the ventral as well as dorsal furrows, the latterof which has also been reported in somebelemnites. The restored belemnite has,

therefore, a tail which looks something
like the posterior half of a dirigible inhaving horizontal as well as vertical tailplanes. Nevertheless, this interpretation
did not give us answer to the question of
why only the ventral furrow has the commissure and other furrows have no particular structure under them.
In this way, hypotheses are derived fromindividual experiences, biological knowledge or anything else. The source of thetheories is of infinite variety, and it matters not from where we have obtained or
as stated by Hempel (1966, p. 15) "invented", the theories. Moreover we "donot believe that we .. · start with observations and try to derive our theories fromthem (Popper, 1957, p. 134)". Then thetheories must be tested as to whether or
not they can explain the results of observations
accumulated
intentionally.
Theories that are falsified are eliminatedand among the remaining competing theories, the best-tested one should be preferred. Under the influence of the aforementioned ideas of Popper, Gould (1980,
p. 102) explained metaphorically that thismethod, when applied to paleontology,
seems like intelligent fishing in the poolof evolutionary biology. If so, I would
like to consider where the fish are.
I was not at all satisfied with thosehypotheses that had been proposed to explain the morphology of belemnite, because
they remain fixed, as if set in concrete.
There are some differences among theauthors, but since Seilacher's work (1970,
p. 393-396), the morphology of fossils has
usually been approached in terms of a
combination of phylogenetic, adaptational,
and architectural aspects. Sophisticatedcombinations of these approaches are ofcourse useful in morphological studies.
The phylogenetic approach is, however,merely an enumeration of the facts following the order of time. unless its ex-

A story of the belemnite
planation is accompanied with genetic anddevelopmental mechanisms. The adaptational approach is also a static comparison
between the morphology and the optimal
design, because with only this approach
we can not make clear the dynamic mechanisms or the genesis of the morphology.
The architectural approach may be mechanically dynamic but it is biologicallystatic in itself. In comparison with thesestatic approaches, the developmental approach is the most direct and dynamic
way to understand morphology throughunderstanding the mechanisms of its
formation. I look upon this approach as
the only way, because I hold a belief thatevery organism has its own unique processof pattern formation. Otherwise it can
not attain its existence. However, ourpresent knowledge on pattern formationis still far from sufficient for providing avehicle for an interpretation of the development of squid.
In the case of belemnites, we fortunately
have certain ontogenetic facts in that theearly part of the shell growth is preserved,
enclosed by the successive growth of theadult shell. In the case of modern squids,
their development has been studied intensively; fishery science in Japan provides us with the results of many excellent
studies. The first task which I undertook
was a study of the ontogeny of belemnites,that is the enumeration of the constituent
parts of a belemnite in order of theirformation without considering their developmental mechanisms. When I started
to describe Neohibolites from Miyako, there
were only a very few papers on the internal structure that utilized a polishedsection of the belemnite. But, as the
Paleozoic nautiloids were in most cases
observed from making polished sections,
there seemed to be no reason why thesame technical methods were not applicable to the study of belemnites. Then the425

description of the internal structure of abelemnite was undertaken from observations of both polished and thin sections.
A structure, named the intermediate
camera was found between the protoconch
and the primordial rostrum, and a bowlin-bowl structure was detected in theprimordial rostrum. Further, the commissure seems not to extend into the primordial
rostrum. The primordial rostrum has long
been thought to be merely a beginningpart of the rostrum. Thus the question,
which was the earlier construction, protoconch or primordial rostrum, disturbed
me in developing a sequential descriptionof the ontogeny of belemnite (PI. 66, Fig.
2). Finding of new structure, however,changed this view on the primordial
rostrum to that it is a construction quite
different from the true rostrum. Thus,
in my earlier paper (Hanai, 1953, p. 66), I
argued that the primordial rostrum wasconstructed first, according to the viewthat smaller construction is formed earlier.
The bowl-in-bowl structure of primordial
rostrum has a smallest bowl in its apex.
But in the later papar (Hanai, 1956, p. 1571),
I did not designate which construction isthe earlier. The only conclusion that canbe drawn is that Neohibolites experienced
two stages. The earlier one is the stage
that includes constructions of protoconch,
intermediate camera, primordial rostrum
and perhaps a few chambers. In the laterstage, a slender true rostrum with pointed
apex and the rest of the chambers wereformed. In applying the methods of microscopic observation of thin sections inpetrology, this sequence is determinablefrom the fact that the true rostrum shaved
off the side of the primordial rostrum (d.
Hanai, 1953, pI. 7, figs. 1-4). For a difference of opinion on the details of formation of the primordial rostrum refer toJeletzky (1966, p. 129, 130; 1972, p. 169,170) and Barskov (1972, p. 493).

Tetsuro HANAI

426

bf---l 1 /10 mm

Text-fig. 2.

The cleavage of Todarodes pacificus Steenstrup, 1888.

A study on the development of themodern squid was started at the eleventhhour in order to interpret the ontogenyof belemnites in terms of the developmental mechanisms. Text-fig. 2 shows
the cleavage of Todarodes studied by Soeda(1952, p. 10-15). Fertilized egg and first
to fourth cleavage stages are shown in
Fig. 2a and Figs. 2b-e, respectively. Afterthe micromere appears in the central area,
the blastomere covers the northern hemisphere of the egg and a blastodiscus isformed (Figs. 2f-h). Text-figure 3 shows
the median sections of the blastodisc us,

Text-fig. 3. Longitudinal sections of anadvanced cleavage stage (a) and a blastula
stage (b) of Todarodes pacificus Steenstrup,
1888. (After Soeda, 1952.)(After Soeda, 1952.)

where a single layer of blastomer (Fig.
3a) becomes double along the marginal
area of the blastodiscus (Fig. 3b). Thus
the blastodiscus grows into the blastula.
Text-figs. 4 and 5 are the illustrations

Text-fig. 4.
Longitudinal sections ofyounger embryos of Loligo vulgaris Lamarck,
1799. (After Korschelt, 1892.) c. g. cerebral
ganglion; ect. ectoderm; end. endoderm; fg.
fore-gut; m. mouth; rna. mantle; meso mesoderm; mg. mid-gut; sh. s. shell gland; y. m.
yolk membrane.

A story of the belemnite427c
Text-fig. 5. Dorsal (a) and ventral (b, c) views of younger embryos of Loligo
vulgaris Lamarck, 1799. (After Korschelt, 1892.) a. info anterior infundibulum; an.
anus; ar. arm; e. eye; fi. fin; gi. gill; info infundibulum; m. mouth; rna. mantle;
o. v. otolithic vesicle; p. info posterior infundibulum; sh. S. shell gland; y. S. yolk sac.

of the development of Loligo from
Korschelt (1892, p. 349; pI. 36, fig. 2). As
the endoderm appears on the inside andthe mesoderm in the medial part, theectoderm in the central area of the blastulasubsides to form a shallow invagination,
the rudimentary shell gland. Then the
rudiment of the mantle extends to close
the opening of the invagination in forming the shell sac (Text-fig. 5). According to Korschelt (1892, pI. 37, figs. 23, 24),
when the large eyes, rudiments of the fin,connected digestive organs and ganglions
are formed, the shell sac is enclosed completely in the dorsal part of the mantle
and extends the full length of the mantle(Text-fig. 6). The animal consumes theyolk and takes a shape close to the adult.
This is shown in the Text-fig. 7, which
is taken from Choe and Oshima (1961, fig.
3.26) who illustrated a young form of
Sepioteuthis just after hatching by dissolving the shell membrane by using enzymes.
These observations on the developmentof various species of squids are well summarized by Ishikawa (1975, p. 379-381)
in Japanese. Recently, the study on theembryonic and post-embryonic development of squids has been advanced verymuch by Boletzky and Boletzky (1973, p.
141-157) and Bandel and Boletzky (1979,
p. 324-335).

Provided that the belemnite shell, including the phragmocone and rostrum,
was formed in the shell sac closely similar
in function to that of the present day
squid, an interpretation that the ventral
furrow underlying the commissure corresponds to the concrescence line of the
opening of rudimentary shell sac seemed
reasonable. Then, it naturally follows
that the variously shaped rostra, from
slender to sturdy, from sharp and pointed
to clavate with a nearly round apical end,and often with many aberrant forms, are
all formed in the blind part of the shell sac.
In the second step, an attempt to test
this interpretation has been made throughTetsuro HANAl

428

~Ai.

Y.ni.

a

y:m.

b

Text-fig. 6. Longitudinal sections of older embryos of Loligo vulgaris Lamarck,
1799. (After Korschelt, 1892.) an. anus; ar. arm; c. g. cerebral ganglion; ect.
ectoderm; fg. fore-gut; fi. fin; inf. infundibulum; i. s. ink·sac; m. mouth; rna.
mantle; meso mesoderm; mg. mid-gut; o. v. otolithic vesicle; p. g. pedal ganglion;r. s. radula sac; s. gl. salivary gland; sh. s. shell sac; st. stomach; v. g. visceralganglion; y. m. yolk membrane.

comparative study of the developmental
process. The comparison in paleontology
consists of two aspects. One is to simplify
the complex state by finding out the common aspects between the two objects intended to compare to each other, and to
give an explanation to the complexity byminimum interpretation. And the other is
to search for the paleobiologically unique
historical events through the eliminationof the determined common aspects.
The protoconch of Neohibolites is approximately 0.3 mm and opening of shellsac of Loligo is approximately 0.2 mm indiameter. The protoconch was probably
formed as the outer shell under the rel-

atively protected condition in the egg
capsule. It is not necessary to infer thatthe shell sac was formed by closing its
opening at the stage of protoconch formation, but at the stage of the formation of
the intermediate camera, the epithelium,
which has extended covering the outerface of the protoconch, is perhaps fused
to from closed shell sac. The primordial
rostrum has a bowl-in-bowl structure of
crescent-shaped, solid plates, and its apex
has not yet been pointed. Perhaps theanimal is still in the egg capsule. Theintermediate camera might be filled withfluid, because it does not possess an opening. After hatching, the true rostrum ofA I3tu'ry of the belemnite

429This lattice-like construction is somewhat
similar to that of the pillar structure observable in Sepia. It is a belemnite witha spongy rostrum; it is lighter than that
we have supposed_ Consequently, Barskov

,

T
\

\
\
I

\
"

t/

I

mg
to

;nr1
Text-fig. 7. A young form of Sepiateuthis lessaniana Lesson, 1830, just after
hatching.

(After Choe and Oshima, 1961.)

the belemnite starts its excessive relative
growth, shaving off the side of the primordial rostrum and growing into a
pointed and slender rostrum with a length
ten times as long as its diameter. Passing
this stage, the young rostrum increases
its thickness to become a stout adult_
Barskov (1972, p. 493) pointed out thatthe layer that forms the protoconch isprismatic and the layer, which depositedthe adapical side of the protoconch, leaving an intermediate camera between the
protoconch and the layer, is nacreous_
Spaeth et at. (1971, p. 3149) proved thatthe innermost layer of the rostrum cavum
consists at least in part of aragonite. The
bowls of the primordial rostrum seem topreserve a structure of perpendicularly
disposed acicular crystals presumably ofaragonite. Following the pioneer work of
Muller-Stoll (1936, p. 172-174), recently
Barskov (1970, p. 559), Spaeth (1971, p. 34,35) and Spaeth et at. (1971, p. 3148) clarified that the true rostrum is primarily
composed of the alternation of the compact nacreous layer with tabular crystals
and a layer of prismatic crystals having
the later fillings as a result of diagenesis.

e~d

11\

WL.· . l·;:~~k::::::i.::/!)I....... ·I···

1'\/\
a

myo

b

Text-fig. 8. Scheme of dorsoventral
sections of marginal portion of nautiloid shell(a) and an idealized belemnite (b). co. areaof mantle epithelium for secretion of conothecal and septal layers; ect. area of mantle
epithelium for secretion of ectostracum; end.
area of mantle epithelium for secretion ofendostracum; gl. growth lines; mg. mantle
groove in which periostracum originate at its
base; p. periostracum; myo. area of mantle
epithelium for secretion of myostracum.

430

Tetsuro HANAI(1972, p. 498, 499; 1973b, p. 292) pointed
out correctly that the rostral layers homologously correspond to the nacreous layers
or endostracum of Nautilus. Where thenshould the layers corresponding to theperiostracum and ectostracum of Nautilus
be found in belemnite? He concluded thatperhaps they were lost.
However, it
seemed to me that in the process of thedevelopment, the layers were pushed away
toward the ventral area of the rostrum.
The layers correspond to the concrescencezone of the shell sac opening. This opening of the epithelium will have been
stretched longitudinally in the process ofdevelopment as the belemnite grew rapidly
towards a direction slightly dorsal to thecentral axis of the development (Text-fig.
8). The ventral furrow should be expected
to be formed in the area where the epithelium for secretion of periostracum andectostracum extend longitudinally (Textfig. 9). Certainly, the concept of homology
is a driving force and fundamental to comparative biology.
Now, it is clear th1t if this is the case,then a shell structure different from thatof the rest of the rostrum is expected to
be found in this area. The cross sectionof the ada pica I part of the rostrum isshown in PI. 66, Fig. 3, where the radial
structure with concentric growth lines isfound. But the section crossing the ventral furrow shows, in spite of the continuation of growth lines across the commissure, sharp discontinuity of the crystal
structure between the calcite which fills
the commissure and the calcite whichforms the main part of the rostrum (PI.
66, Fig. 4). Thus the ventral furrow is a
structurally weak part of the belemnite
body, and the belemnite body coils veryloosely as if it held the weak part in its
venter. This interpretation may enable
us to gain a preferable understanding of
the ventral furrow of belemnites.

Text-fig. 9. Explanation of homologous
relation of mantle epitherium between nautiloid and belemnite. Idealized mantle epitherium for secretion of nautiloid shell
Figures for explanation oflayers (a).
homologous relation (b-e). Idealized mantle
epitherium for secretion of belemnite shelllayers (f). ect. area of mantle epitheriumfor secretion of ectostracum; end. area ofmantle epitherium for secretion of endostracum; myo-co. area of mantle epitheriumfor secretion of either myostracal, conothecal
or septal layers. Broken line represents the
dorso-ven tral plane.

When one studies the paleontologicalfacts, it seems relevant to fish in the poolof the mechanisms by which the organisms
build up the facts. Then, I venture to
say that the fact of the ventral commissure of belemnites may turn into a paleontologically significant historical event
which may have occurred in the rudimentary shell sac in the early stage ofthe development, and gave rise to theformation of the ventral commissure. The
conclusion seems to emerge through giving the developmental explanation to theenumeration of the constituent parts of abelemnite following the order of theirformation. The developmental explanation
was obtained by comparing this enumer-

A story of the belemnite
ation with the development of the modern
organisms closely related to the belemnites.
Carr (1961, p. 57) explained the similarity
between historians' and scientists' tasks,
quoting a modern physicists' statementthat what they investigate is not facts
but events. What we as paleontologists
search for may be the interpretations
called events of the past, and we may besatisfied with knowing what was happening in the past.
References cited
Bandel, K. and Boletzky, S. V. 1979. A com·parative study of the structure, develop·
ment and morphological relationships ofchambered cephalopod shells. Veliger, v.
21, no. 3, p. 313-354, 6 pis., 27 figs.
Barskov, I. S. 1970. Structure of the belemnitid
rostrum. Paleontol. J., no. 4, p. 557-559,
2 figs.
- - . 1972. Microstructure of the skeletal
layers of belemnites compared with external shell layers of other molluscs.
Paleontol. J., no. 6, p. 492-500, pl. 6, 3 figs.
- - . 1973a. Structure of the protoconch andontogeny of the belemnites (Coleoidea,
Cephalopoda). Dokl. Akad. Naul? SSSR,
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- - . 1973b. Microstructure of the skeletal
layers of Sepia and Spisula compared withshell layers of other mollusks. Paleontol.
]., no. 3, p. 3-13, pl. 1, 4 figs.
Boletzky, S. v. and Boletzky, M. V. v. 1973.
Observations on the embryonic and earlypost-embryonic development of Rossia
macrosoma(Mollusca,
Cephalopoda) .
HelgolCinder wiss. Meeresunters., v. 25,
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Carr, E. H. 1961. What is history? The George
Macaulay Lectures delivered in the University of Cambridge January-March 1961.
Macmillan, London, 155 p.
Choe, S. and Oshima Y. 1961. On the embryonal development and the growth of thesquid, Sepiotheuthis lessoniana Lesson [inJapanese]. Venus, v.21, no. 4, p. 462-476,
5 figs., 3 tables.

431

Gould, S. J. 1980. The promise of paleobiology
as a nomothetic, evolutionary discipline.
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Gustomesov, V. A. 1975. The furrows, commissure and fissure on belemnoid rostra
and an interpretation of their biologicalsignificance. Paleontol. J., no. 3, p. 316322, 1 fig.
Hanai T. 1953. Lower Cretaceous belemnites
from Miyako district, Japan. Jap. J. Ceol.
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- - . 1956. Interpretation of the ontogeny ofa Lower Cretaceous belemnite (Abstract).
Bull. geol. Soc. Am., v. 66, p. 1571.
- - . 1977. Introduction (p. 1) and Further
problems for systematic study of Japanese
Ostracoda (p. 84-86). In T. Hanai et at.
Checklist of Ostracoda from Japan and its
adjacent seas. Univ. Mus., Univ. Tokyo,
Bull. no. 12, llO p., 4 pis., 2 figs.
Hempel, C. G. 1966. Philosophy of naturalscience. Foundations of philosophy series.
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Jersey. ll6 p.
Ishikawa, M. 1957. Cephalopoda (p. 376-384,
14 figs.). In M. Kume and K. Dan [eds.]
Invertebrate embryology [in Japanese].
Baifukan, Tokyo. 436 p.
Jeletzky, ]. A. 1966. Comparative morphology,
phylogeny, and classification of fossil
Coleoidea. Paleontol. Contrib., Univ. Kans.,
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Kimura, M. 1958. The causes of evolution
and statistic consideration on evolutionary
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Tokyo. 330 p.
Kobayashi, T. 1960. Studies on trilobites inlast three decades. Trans. Proc. palaeontol.
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Korschelt, E. 1892. Beitrage zur Entwicklungsgeschichte der Cephalopoden.
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Entstehung des Darmkanals und Nervensystems in Beziehung zur Keimblatterfrage.
Festschr. zum 70. Geturtstag R. Leuckarts.

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Tetsuro HANAIEngelmann, Leipzig, p. 347-373, pIs. 36,
37, 9 figs.
Makiyama, J. 1935. Morphogenetic history of
the Umboniidae [in Japanese]. Venus, v.
5, no. 5, p. 241-255, 2 figs.
Matsumoto, T. 1975. Ammonitology in JapanA historical review. Trans. Proc. palaeontol. Soc. jap., N. S., no. 97, p. 1-6.
Muller-Stoll, H. 1936. Beitrage zur Anatomie
der Belemnoidea. Nova Acta Leopoldina,
Hall/Saale, new ser. 4, no. 20, p. 160-226,
pI. 1-14, 5 figs.
Naef, A. 1922. Die fossilen Tintenfische. Eine
palaontologische Monographie. Verlag von
Gustav Fischer, Jena, 322 p., 100 figs.
Popper, K. R. 1957. The poverty of Historicism.
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Seilacher, A. 1970. Arbeitskonzept zur Konstruktions-Morphologie. Lethaia, v. 3, p.
393-396, 1 fig.
Shvetsov, M. S. 1913.
Lower Cretaceous
belemnites of Abkhazia [in Russian].
Yezhegodnik po Ceol. i Mineral. Rossii.
(Annuaire Geologique et Mineralogique dela Russie), v. 15, nos. 2/3, p. 43-74, pI. 2-5.

Soeda, J. 1952. On the artificial inseminationand the early cleavage of the ovumin
oegopsid cephalopod, Ommastrephes sloani
pacificus, Surume-ika [in Japanese]. Bull.
Hokkai.do reg. Fish. Res. Lab., no. 5, p.
1-15, pI. 1-4.
Spaeth, C. 1971. Aragonitische und calcitischePrimarstrukturen im Schalenbau eines
Belemniten aus der englischen Unterkeide.
Paliiontol. Z., v. 45, nos. 1/2, p. 33-40,
pI. 2-4, 1 fig.
- - , Hoefs, ]. and Vetter, U. 1971. Some
aspects of isotopic composition of belemnites and related paleotemperatures. Ceol.
Soc. Am. Bull., v. 82, p. 3139-3150, 13 figs.
Stevens, G. R. 1965. The Jurassic and Cretaceous belemnites of New Zealand and areview of the Jurassic and Cretaceous
belemnites of the Indo-Pacific region. New
Zealand geol. Surv., paleontol. Bull. 36,
283 p., 25 pIs., 43 figs.
(In preparing this English translation of theaddress, the addition of text-figures 8 and9 has been made to further explain thecontents.)

Explanation of Plate 66
Figs. 1-4. Neohibolites miyakoensis Hanai, 1953
la, b. Ventral and lateral views of rostrum, MM2535, loco Hn-0220, northern cliff of Hiraiga
cove, Tanohata-mura, Tanohata Formation, xL
2. Dorso-ventral section of the apex of phragmocone, showing bowl-in-bowl structure of theprimordial rostrum, intermediate camera and protoconch, MM6995, loco Ks-2005, artificial
channel east of Ebisudana, Hideshima, Miyako-shi, Tanohata Formation, x 70.
3a, b. Transverse section of adapical region of rostrum, MM6996, loco Hn-031O, northern
cliff of Hiraiga cove, Tanohata-mura, Hiraiga Formation, x 17 (a, transmitted light, b,
crossed nicols).
4a, b, C. Transverse section of rostrum cavum, showing sharp discontinuity of crystal structure between commissure and the rest of rostrum, in spite of the continuous nature of
the growth lines across the two areas, MM6997, loco OJ-5549, northern cliff of Raga
cove, Tanohata-mura, Hiraiga Formation, Orbitolina facies, x 17 (a, transmitted light, b,
crossed nicols, c, same as 4b except stage rotated 45°).

HAN A f :A story of the belemnite

Plate 66

Trans. Proc. Palaeont. Soc. Japan, N. S.. No. 128, pp. 433-443, pI. 67-68, December 30, 1982

753. FORMATION AND FUNCTION OF THE SIPHUNCLE-SEPTALNECK STRUCTURES IN TWO MESOZOIC AMMONITES*
KAZUSHIGE T ANAJ:jE
Department of Earth-Sciences, Ehime University, Matsuyama 790, JapanYOSHIO FUKUDA
Chiba Prefectural Institute of Public Health, Chiba 280, JapanandIKUWO OBATA
Department of Paleontology, National Science Museum, Tokyo 160, Japan

Abstract. SEM observation on the siphuncle-septal neck structures in two Mesozoic ammonites, Eleganticeras elegantulum and Reesidites minimus is described andcompared with modern chambered cephalopods. The state of development of theporous and nonporous structures within the chambers of the two ammonites is very
similar to that of modern Nautilus; and furthermore, a thin conchiolin membrane(pellicle) covers the greater portion of the siphuncle and the inner chamber surface
as in Nautilus. Based on these observations and the knowledge on modern Nautilus
biology a model is proposed for a new chamber formation in ammonites. This work
also postulates that in ammonites the cameral liquid initially filling up the late-formedchambers was primarily drained into the siphuncular cord osmotically via a wettable
pellicle and porous septal prismatic zone within the septal neck region.

Nautilus, Sepia and Spirula, have chambered shells preserved in the modern
fauna.
Owing much to the works of Denton
and Gilpin-Brown (1961a, b, 1966, 1971,1973) and others (Denton et al. 1961, 1967;
Collins and Minton 1967), our knowledge
about buoyancy control and shell formationin modern chambered cephalopods has
greatly increased: i. e. these works demonstrated that (1) living animals possess

Introduction
It is well known that many extinct
cephalopods had chambered shells consisting of a shell wall, septa and siphuncle.
Among modern cephalopods, chambered
shells have been completely lost or reduced
to chitinous gladii in most coleoids. Onlyin three taxonomically different genera,* Received March 12, 1982.

433

434

Kazushige TANABE et al.

low-pressure gas and various amounts ofliquid (cameral liquid) within their chambers in order to approximate their densities to that of seawater, and that (2) they
can migrate vertically by pumping cameral
liquid osmotically into or out of theirchambers through the siphuncle.
The above model for buoyancy control
by Denton and Gilpin-Brown has subsequently been applied to many extinct
chambered cephalopods including ammonites (e. g. Heptonstall 1970).
Recent advances in modern Nautilus
functional morphology and physiology(Ward and Martin 1978; Ward et a!. 1980;
Collins et al. 1980), however, suggest that
Denton and Gilpin-Brown's model cannot
be entirely accepted for all chamberedcephalopods.
In the previous paper (Obata et al. 1980),
we have proposed hypothetical models
showing possible pathways of cameral
liquid removal in ammonites and modern
chambered cephalopods. That paper was
based mainly upon the comparative ultrastructural observations on the siphuncularmembranes and septal prismatic zone
within the septal neck region; but little
was described for a thin organic membrane
covering the siphuncle and inner side of
the camera of all modern chamberedcephalopods. This membrane, named thepellicle by Denton and Gilpin-Brown (1966),
has also been recognized in several exceptionally well-preserved ammonites (Mutvei
1967; Erben and Reid 1971; Bayer 1978;
Bandel 1981). Nevertheless, details of its
ultrastructure and relations with otherfine stuructures such as the de-coupling
space and the septal prismatic zone haveremain unsolved.
In this paper we describe the ultrastructural relations of these fine stuructures,
discussing their possible function for a
new chamber formation and cameral liquid
removal.

Materials and Methods
Materials: - Specimens of one earlyJurassic Eleganticeras elegantulum (Younget Bird) and two late Turonian (Cretaceous)Reesidites minimus (Hayasaka et Fukada)
were examined in this study. The former
was found in a derived nodule of late
Pleistocene glacial deposits near Hamburg,
Germany, and the latter two were collectedfrom a nodule in the sandstone bed (Mikasa
Formation) at loco M 180 in the Sannosawa
Valley, Manji area, central Hokkaido(Japan).
Specimens of modern chamberd cephalopods utilized for comparison are: a shellof Nautilus pompilius Linnaeus, captured
live from off Bindoy, Negros Oriental,
the Philippines, a shell of Spirula spirula
Lamarck stranded on a beach near Noumea,
New Caledonia, and cuttlebones of two
Sepia esculenta Hoyle captured live fromoff Tsuyazaki, northern Kyushu (Japan).
Methods :-Ammonite specimens wereprepared for SEM observation by sectioning along the median plane. Subsequently
the sectioned surface was etched with 4%hydrochloric acid for a few minutes, andthereafter the etched surface was coatedwith gold using an iOIl coater. The septal
neck region was removed from the shellsof Recent species and coated with gold.
They were not etched. Our observation
was made by means of a SEM (HitachiCo., H-450 type). All specimens examined
are stored in Ehime University without
registered number.

Observations
Siphuncular wall: - The siphunculartubes of E. elegantulum and R. minimus
are both composed of multi-layered, concentric membranes of unporous material(PI. 67, Figs. 1-3, 5). In the examined two

753.

Ammonite Siphuncular System

specimens of R. nzinimus decalcified siphuncular membranes, each of which isabout 2-3 flm in thickness, retain wellpreserved conchiolin, consiting of lace-like,tuberculate fibrils (PI. 67, Figs. 1-2). The
above characteristic features of siphuncularmembranes have already described in someother late Cretaceous ammonites by Obataet aI. (1980). Obata and others compared
the ammonite siphuncular wall with theinner horny layer of modern Nautilus
siphuncular wall, because of the ultrastructural similarity of conchiolin (see TEM
photographs of Gregoire, 1962).
Such conchiolinic siphuncular membranes
have already found in many Mesozoic ammonites belonging to various taxa (Schindewolf 1967; Erben et aI. 1969; Erben andReid 1971; Westermann 1971; Kulicki
1979; Obata et aI. 1980; Bandel 1981). We,
therefore, believe that the siphuncularwall in all Mesozoic ammonites was madeof conchiolinic material only.
Pellicle :-A thin membrane covering theammonite siphuncle is clearly distinguishedfrom the siphuncular membranes by its
different ultrastructure. As shown on PI.
67, Figs. 2-3, in R. minimus it is formedof unorientated microfibrils of about 0.1
to 0.2 flm in diameter. Ultrastructurally,
the membrane is apparently very similar
to those of modern Nautilus and Spirula,which are present on the inner chamber
surface (Gregoire 1962, 1973; Denton and
Gilpin-Brown 1966, 1973). We, therefore,call this membrane the pellicle. The microfibrils forming the pellicle of R. minimus
are, however, coarser than those in Nautilus(see Gregoire, 1962, figs. 43-44, 58, 59-60).
Jeunniaux (1963) using a selective enzymatic method, showed that the pellicleof Nautilus consists mostly of chitin.
From these lines of evidence, the pellicleof ammonites was probably made of chitin.
In E. elegantulum the pellicle separates
from the siphuncular tube in the anterior

435

portion of a chamber, where it forms anarrow space (here named the de-coupling
space), together with the siphuncular tube
and septal layer (PI. 67, Fig. 5). Then it
extends to the inner chamber surface (PI.
67, Fig. 6).
As the presence of a pellicle has alreadybeen confirmed in some well-preserved
Jurassic and Cretaceous ammonite specimens (Branco 1880; B6hmers 1936; Erben
and Reid 1971; Westermann 1971; Bayer
1975, 1978; Bandel 1981), it was probably
present in all Mesozoic ammonites. According to the above authors the pellicle
covering the greater portion of the siphuncle is fixed to the ada pica I surfaceof the septum and ventral shell wall, where
it forms a semi-conical space, formerly
termed the de-coupling room by Bandel
and Boletzky (1979). These previous observations thus correlate well with ourdata. It should be noted that in bothNautilus and ammonites the pellicle covers
the greater part of the siphuncular tube
in contr11st with the absence on the siphucnular surface in Spirula (Dauphin
1976; Bandel and Boletzky 1979).
Septal prismatic zone :-In the specimenof E. elegantulw1l a thin, porous ridge-likeprismatic zone, is favorably preserved
under the nacreous layer within the septal
neck region (PI. 67, Fig. 5). Also it extends to the ada pica I side of the septa(PI. 67, Fig. 6). Near the septal neck theprismatic zone is laterally covered withthe concentric siphuncular membranes,
which are, in part, absent within the prochoani tic septal neck.
The presence of a septal prismatic zone
has been ascertained in some Paleozoic
and Mesozoic ammonites (Mutvei 1967;
Birkelund and Hansen 1968; Erben andReid 1971; Drushchits et aI. 1976; Kulicki
1979; Obata et aI. 1980; Bandel 1981).
As compared with the examples of previous reports, the septal prismatic zone in436

Kazushige TANABE et al.

E. elegantulum is more widely developed.
The septal prismatic zone of ammonites
is apparently homologous with those ofother modern and extinct chambred cephalopods, as Bandel and Boletzky (1979) and
Obata et al. (1980) have already pointed
out. The state of development of thiszone, however, varies greatly among ammonites and modern chambered cephalopods. Namely, the zone of Spirula spirula
is characterized by its highly porous structure consisting of pillar-like prisms, which
are widely distributed within the long
septal neck region (PI. 68, Figs. 3-4). Such
pillar-like prisms are also present within
the cuttlebone of Sepia esculenta in theearly stage (PI. 68, Fig. 5), but they gradually change to the platy prisms as the
cuttlebone grows (PI. 68, Fig. 6).
In Nautilus the septal prismatic zone isrestricted to the adoral side of a septum
and within the short retrochoanitic septal
neck region (PI. 68, Fig. 1). The zone of
Nautilus is made of pillar-like prisms, as
in Sepia and Spirula. In the adoral sideof the septal neck the prisms are covered
with the aggregated spherulitic aragonite
needles of the middle chalky layer (Denton
and Gilpin-Brown, 1966) of the siphuncularwall (PI. 68, Fig. 2). Although the chalkylayer is completely lost in ammonites, thestate of development of the septal prismaticzone in ammonites is, thus, comparable
in morphology to that of Nautilus in its
restriction to the region of the septum.

New Chamber Formationin Ammonites
Mutvei (1967, text-fig. 4) proposed amodel showing the successive stages of a
new chamber formation in the earlyJurassic ammonite, Promicroceras sp. According to his model the septal layer issecreted earlier than the conchiolinic siphuncular tube. Mutvei's interpretationis, however, incorrect at least for theformation of a septum and siphuncle.
Kulicki (1979, pI. 40, fig. 1) actually figured
a median-sectioned specimen of the Jurassic Holcophylloceras zignodianum, in which
the siphuncular tube in its full length isalready present before the secretion of a
new septum.
Earlier formation of the siphuncular tube
than the septum is also observed in modern
Nautilus (data from observations of N.
pompilius specimens captured from the
Tailon Strait, Philippines). Recent works
on modern Nautilus functional morphology
have given an important information for
a new chamber formation of ammonites.
Denton and Gilpin-Brown (1966, 1973),
Collins et al. (1980) and Ward et al. (1981)
have demonstrated that in modern Nautilus
and Spirula the latest formed chamber is
initialy filled with liquid (=cameral liquid).
The liquid is subsequently drained into
the siphuncular cord after the complete
secretion of a new septum. Collins et al.
(1980) suggested that during the septum
formation the cameral liquid in a living
Nautilus functions to support an incomplete septum from the hydrostatic pressure. The experiments by Collins andMinton (1967) and Raup and Takahashi(1967) provided another interesting factthat the siphuncular tube of Nautilus is
imploded more easily than the other partsof the shell. To sum up, an earlier formation of the siphuncular tube than the
septum is absolutely necessary for a living
Nautilus in a deep sea to support thismechanically weak structure from the highhydrostatic pressure. Such situation mightpresent in ammonites.
There are two different views as to the
new septum formation in ammonites. One
was proposed by Seilacher (1975) and Bayer(1978), who stressed the pellicle was formed
before the nacreous septum could formon it. According to their interpretation,

753.

Ammonite Siphuncular System

the septum of ammonites represents themineralized elastic membrane of a stretchedmantle epithelium (Seilacher's "pull-off"model).
The other opinion is that the pellicle asbase for mineral deposits is not present
when septum secretion starts (Bandel1981).
As described before, in the specimen of
E. elegantulum examined the pellicle covers
both the siphuncular tube and inner chamber surafce. Furthermore, in the adapicalside of the septum the septal prismaticzone is intercalated between the pellicle
and the nacreous septal layer. These observations support Seilacher's "pull-off"model.
The results of this work and the knowledge on modern Nautilus chamber formation may lead the following generalized
models for a new chamber formation inammonites.
1. The primary epithelium of the adorally "withdrawing" mantle first secretesa pellicle (Text-fig. 1-2).
2. In the second stage the adoral endof the epithelial cells secrets the porous
septal prismatic zone, and simultaneously
concentric siphuncular membranes become
thick (Text-fig. 1-3).
3. In the third stage the calcium-secreting cells of the mantle in the adoral sideof the septal prismatic zone begin to add
the nacreous layers of the septum (Textfig. 1-3'""4).
4. In the last stage the septum attains
to a full thickness, and thereafter thecameral liquid filling up the new chamber
is gradually pumped out into the siphuncular cord (Text-fig. 1-4).
Denton and Gilpin-Brown (1973) suggested that in modern chambered cephalopods, cameral liquid removal for a newchamber formation and buoyancy control
is caused by the transportation of saltions and the consequent osmotic flow of437

the liquid through the liquid permeablesiphuncular wall and/or epithelium. Indeed,
the siphuncular epithelia of Nautilus andSepia. are histologically compared with thedeep, basal infoldings of proximal renal
tubes of advanced animals (Denton and
Gilpin-Brown 1961a, 1966, 1973 ; Bassot andMartoja 1966; Fukuda et al. 1981). Fukuda
et al. (1981) also realized that in thealmost mature animal of N. pompilius the
cube, calcium-secreting cells along theadoral side of the last septum are adapically displaced to the high columnar siphuncular epithelial cells with distinct oval
cavities. These lines of histological evidence well explain the osmotic removalof cameral liquid under the high hydrostatic pressure.
Judging from the similarities of fundamental internal shell structures to modern
Nautilus, the above osmotic removal modelby Denton and Gilpin-Brown may be applied to the cameral liquid drainage for a
new chamber formation in ammonites.
If this presumption is correct, the pellicle
covering the siphuncular tube and septal
surface might have functioned as a transmitting agent of the cameral liquid, if not
in direct contact with the siphuncle, outof chamber.
In ammonites the greater portion of thesiphuncular tube is made of unporous conchiolinic material, and the porous structure
is restricted to the septal prismatic zone
within the septal neck region. These
structural characteristics support our previous interpretations that most cameral
liquid in ammonites was primarily transported through the narrow space of the
septal prismatic zone, and that the liquid
was not moved directly through the thick,conchiolinic siphuncular membranes (Obataet al. 1980). Furthermore, it is postulated
that the de-coupling space in several lateformed chambers was probably filled withliquid for maintaining dampness of theKazushige TANABE et al.

438

r ventra Ism.... .. ".
"

sm

se

2'i·::~i1~:·;._~~sm
1.

_J-sm

:::: ... : /spe

anterior

ldorsa I

Text-fig. 1. Model showing the successive stages of a new chamber formationin ammonites. See the text for details of this figure. n: nacreous layers of septum,
sp: septal prismatic zone, p: pellicle, ds: de-coupling space, sm: siphuncular membranes, pe: primary epithelium of mantle, se: calciumsecreting epithelium of mantle,
spe: siphuncular epithelium. Possible pathways of osmotic cameral liquid drainage
are shown by arrows.

pellicle. This possible case is illustrated
in Text-fig. 1.
Remarks on the Mode of Lifein Ammonites

We previously showed the diagrams of
the siphuncular systems of selected modern
and fossil chambered cephalopods including ammonites (Obata et al. 1980, fig. 3).

Text-fig. 2 summarizes the revised diagrams of ammonites and modern cephalopods, in which some our own new observations are added. As shown in thisfigure, the siphuncle-septal neck structuresin ammonites and modern species differ
considerably from one another in the
degree of development of porous and/or
nonporous layers. This may be importantin estimating the mode of life of am-

753.

Ammonite Siphuncular Systemmonites, because the rate of cameral liquid
removal is affected by the surface area
and the permeability coefficient of theporous layer which contacts the siphuncular epithelium (Heptonstall 1970; Chamberlain 1978).
Obata et aL (1980) and Bandel (1981)
emphasized that the conchiolinic siphuncular membranes in ammonites and theinner and horny layer of Nautilus
siphuncle both seem to be unpermeable toliquid and salts, because of their un porous
ultrasturcuture. The conchiolinic membranes are, therefore, regarded as the chief
inhibitor of the liquid transport, and causes
the liquid permeability coefficient of the
Nautilus siphuncle to have a low value(ca. 24 microdarcies, Chamberlain 1978; 7
microdarcies, Moore 1981). Recently Wardand Martin (1978) measured a maximuJT'

rate of cameral liquid removal of 1.0 mIl
day Ichamber in four live N. macromphalus
living at 0 to 25 m in water depth. This
pumping capacity of living animals is
about 30 times slower than the liquid flow
rates through the tube wall (e. g. Collins
and Minton 1967; Chamberlain 1978). The
osmotic pumping ability of the siphuncularepithelium in living Nautilus, accordingly,
limits the maximum flow rate to about
1 ml/day/chamber. Collins et al. (1980)
showed that the cameral liquid in Nautilus
is not used for vertical migration but
functions as ballast to maintain slightnegative buoyancy in the sea. The cameral
liquid is, therefore, always pumped out
slowly to compensate for both episodic andlong-term growth of the shell and tissue.
In contrast to the situation in Nautilus,
the siphuncular epithelium of Sepia and
DORSALDORSAL

A

439

n
'"
,(:I

septal,L;{,~~necksp~:~:""""""'!""l!"'"l!l!.\':'~'VEN TRA L

VENTRALoc
DORSAL

VENTRAL

VENTRAL

Text-fig. 2. Diagrams of the siphuncle-septal neck structures in ammonites and modern
chambered cephalopods (adapted from Obata et al. 1980, fig. 3). A: ammonites, B: Nautilus,
C: Spirula, D: Sepia. c: conchiolinic membranes, c': pellicle, sp: septal prismatic zone,
ds: de-coupling space, spr: spherulitic prismatic layer, p: prismatic layer of shell wall, n:nacreous layer. Arrows indicate adoral direction. Siphuncular tissue is dotted.

440

Kazushige TANABE et al.

Spirula is covered with the wide-developed,
pillar-like septal prismatic zone and/or the
spherulitic prismatic layer of the siphuncular wall, without the intercalation of aconchiolinic layer. Sepia undergoes daily
vertical migration by changing theirweight in the sea with the successive
pumping of cameral liquid into or out of
siphuncle, and the rate of liquid removal
measured in S. officinalis is about 4 to 4.5
m1/0.5 day, which is much greater thanthat measured in Nautilus (Denton and
Gilpin-Brown 1961a, b).
Although the mode of life in Spirula
remains uncertain, it lives in mid-waterof the deep sea (Clarke 1970; Denton and
Gilpin-Brown 1973). Clarke (1970) showedthat living Spirula is found at depths
around 600 to 700 m during the day.
Because of the characteristic internal shellstructures and the presence of the lowpressure gas within the camerae, it has
been generally accepted that Spirula cancontrol its buoyancy by pumping cameral
liquid osmotically like Sepia (Denton et al.
1967; Denton and Gilpin-Brown 1971, 1973).
This hypothesis is also supported by thefact that the siphuncular tube wall of

Spirula is permeable to liquids (Denton etal. 1967).
The septal neck-siphuncle structures ofammonites are apparently very similar to
those of modern Nautilus in the state ofdevelopment of porous and unporous layers
within the chambers (Text-fig. 2). This
fact suggests us an analogy in way oflife between the two taxonomically different groups.
Some previous authors emphasized thatammonites, like modern chambered cephalopods, could control their position in thewater column via the osmotic flow ofcameral liquid through the conchiolinic
siphuncular wall (Mutvei 1967 ; Heptonstall
1970; Mutvei and Reyment 1973; Ward
1979; Ward et al. 1980) or through theseptal prismatic zone (Bandel and Boletzky1979). We, however, do not agree withthis interpretation because of the following
reason.
As was demonstrated mathematically by
Chamberlain (1978), the dimensions of thesiphuncle must increase more rapidly than
the increase of shell size in order tomaintain an uniform fluid transport for agiven pressure, fluid viscosity and siphuncle

Explanation of Plate 67
Figs. 1-6. Siphuncle·septal neck structures in ammonites.
Fig. 1. Ultrastructure of siphuncular wall in Reesidites minimus (Hayasaka et Fukada),
showing the lace-like coarse fibrils of conchiolin. Specimen A from loco M 180, Manjiarea, Hokkaido. K. Tanabe colI.
Fig. 2. Outermost surface of siphuncular tube in R. minimus, showing the thin pellicle
membrane covering the siphuncular wall. Specimen B from the same locality as that inFig. 1. K. Tanabe colI.
Fig. 3. Part of Fig. 2, showing the unorientated microfibrils of conchiolin.
Fig. 4. Septal neck structure of Eleganticeras elegantulum (Young et Bird) in the mediansection. Arrow indicates adoral direction. Specimen from a derived gravel in the late
Quarternary glacial deposits near Hamburg. U. Lehmann colI.
Fig. 5. Part of Fig. 4, showing the de-coupling space (d) in the adapical side of the septum.
p: pellicle, h: horny siphuncular membranes.
Fig. 6. Part of Fig. 4, showing the porous septal prismatic zone (pr) between the pellicle(P) and the nacreous septal layer (n).
Photos by Y. Fukuda under 20 KV for an acceleration voltage.

TANABE, FUKUDA

and OBATA:

Ammonite Siph~mcular System

Plate 67

753.

Ammonite Siph'nncular Systempermeability. The growth of siphuncle
dimensions (tube diameter and tube wall
thickness) versus shell radius in modern
Nautilus and most ammonites is strongly
negatively allometric (Trueman 1941 ;Westermann 1971; Chamberlain 1978;
Tanabe et al. 1979), and this indicates adecrease in the capacity of cameral liquid
removal during life, if this is the controlling factor. From these indirect evidence it is suggested that like modern
Nautilus the cameral liquid removal inammonites was undoubtedly very slow,
at least in the late growth stage. It seems
likely that the liquid in adult ammonites
was not used for vertical migration as in
Nautilus.
Acknowledgement

We thank Drs. J.A. Chamberlain Jr.
(Brooklyn College of the City Universityof New York), C. F. Roper (Smithsonian
Institution), W. J. Kennedy (University of
Oxford) and T. Birkelund (Copenhagen
University) for reviewing the earlier darft,and Drs. T. Matsumoto (c/o Kyushu University), U. Lehmann (Hamburg University), C. Gregoire (U~ge University) and
K. Bandel (Erlangen-Niirnberg University)
for their valuable discussions. Studies ofmodern cephalopods were made possible
by facilities of the member of oversea
research project in 1981 (leader Dr. S.
Hayasaka of Kagoshima University) andthe staff of the Marine Biological Stationof the Faculty of Agriculture of Kyushu
University at Tsuyazaki, Kyushu. U.
Lehmann and Dr. S. Mikami also provided
specimens of Eleganticeras and Spirula
for this study. This work was supportedby the Science Research Fund of the Japanese Ministry of Education, Science andCulture (Tanabe, oversea research andObata, Tanabe and Fukuda, No. 56340041
for 1981).

441

References
Bandel, K. (1981): The structure and formation of the siphuncular tube of Quenstedtoceras compared with that of Nautilus(Cephalopoda). N. lb. Ceol. Paliiont. Abh.,161, 153-17l.
- - and Boletzky, S. v. (1979): A comparative
study of the structure, development andmorphological relationships of the chambered cephalopod shells. The Veliger 21,
313-354.
Bassot, J. M. and Martoja, M. (1966): Histologie
et fonction du siphon chez Ie Nautile.
C. R. Acad. Sci. Paris, Ser. D, 263, 980-982.
Bayer, U. (1975): Organische Tapeten im
Ammoniten-Phragmokon und ihr Einfluss
auf die Fossilisation. N. lb. Ceol. Min.
Mon. 1975 (1), 12-25.
(1978): The impossibiliy of inverted
suture lines in ammonites. Lethaia 11,
307-313.
Birkelund, T. and Hansen, J. (1968): Early
shell growth and structures of the septa
and the siphuncular tube in some Maastrichtian ammonites. Medd. fra. Dansk
Ceol. Forening. K¢benhavn 18, 71-78. pIs.
1-4.
B6hmers, J. C. A. (1936): Bau und Struktur
von Shale und Sipho bei Permischen Ammonoidea. Drukkerij Univ. Apeldoorn.
125 p.
Branco, W. (1880): Beitrage zur Entwichelungsgeschichte der fossil en Cephalopoden.
Teil. 2. Palaeontographica 27, 17-81, pIs.
3-1l.
Chamberlain, J. A. Jr. (1978): Permeability ofthe siphuncular tube of Nautilus: its
ecologic and paleoecologic implications.
N. lb. Ceol. PalCiont. Mh.1978 (3), 129-142.
Clarke, M. R. (1970): Growth and developmentof Spirula spirula. l. mar. bioI. Assoc.
U. K. 50, 53-64.
Collins, D. H. and Minton, P. (1967): Siphuncular tube of Nautilus. Nature 216,916-917.
- - , Ward, P. D. and Westermann, G. E. G.
(1980): Function of cameral water in
Nautilus. Paleobiology 6, 168-172.
Dauphin, Y. (1976): Microstructure des coquilles de Cephalopodes. 1. Spirula spirula L.

442

Kazushige TANABE et al.

(Dibranchiata, Decapoda). Bull. Mus. Nat.
Hist. nat. (3) 54 (382), 197-238.
Denton, E.]. and Gilpin-Brown, J. B. (1961a):
The buoyancy of the cuttlefish Sepia officinalis (L.). J. mar. bioi. Assoc. U. K.
41, 319-342.
--- and - - (1961b): The effect of light on
the buoyancy of the cuttlefish. Ibid., 41,
343-350.
- - and - - (1966): On the buoyancy of thePearly Nautilus. Ibid., 46, 723-759.
- - and - - (1971): Further observations on
the buoyancy of Spirula. Ibid., 51, 363-373.
- - and - - (1973): Floatation mechanisms
in modern and fossil cephalopods. Adv.
Mar. Bioi. 11, 197-268. Academic Press,Lond.
- - , - - and Howarth, J. V. (1961): Tile
osmotic mechanisms of the cuttlebone. J.
mar. bioi. Assoc. U. K. 41, 351-364.
- - , - - and - - (1967): On the buoyancy
of Spirula Spirula. Ibid., 47, 181-191.
Druschchits, V. V., Bogoslovskaya, M. F. andDoguzhayeva, L. A. (1976): Evolution of
septal necks in the Ammonoidea. Paleont.
Jour. 1976 (1), 37-50.
Erben, H. K., Flajs, G. and Siehl, A. (1969):
Die Friihontogenetische Entwicklung der
Schalenstruktur ectocochleater Cephalo-

poden. Palaeontographica 132A, 1-54, pis.
1-15.
- - and Reid, R. H. (1971): Ultrastructure ofshell, origin of conellae and siphuncularmembranes in an ammonite. Biomineralisation 3, 22-31, pis. 1-2.
Fukuda, Y., Tanabe, K. and Obata, I. (1981):
Histology of the siphuncular epithelium of
Nautilus pompilius and its functionalsignificance. Fossil Club Bull. 14, 29-40(in Japanese with English abstract).
Gregoire, C. (1962): On sUbmicroscopic structure of the Nautilus shell. Bull. Inst. Roy.
Soc. Nat. Berg. 38, 1-71.
- - (1973): On the sUbmicroscopic structureof the organic components of the siphon
in the Nautilus shell. Archives Internat.
Physiol. Biochim. 81, 299-316, pis. 1-14.
Heptonstall, W. B. (1970): Buoyancy control
in ammonites. Lethaia 3, 317-328.
Jeunniaux, C. (1963): Chitine et chitinolyse.
Masson edit., Paris, 181 p.
Kulicki, C. (1979): The ammonite shell: its
structure, development and biological significance. Pal. Polon. (39), 97-142, pis. 24-48.
Moore, W.A. Jr. (1981): The permeability ofthe siphuncular tube of Nautilus and its
relation to buoyancy control. Cephalopod
Newslettter 5, 13-14.

Explanation of Plate 68
Figs. 1-6. Septal neck structures in modern chambered cephalopods.
Fig. 1. Highly porous septal prismatic zone (pr) in the adoral side of the septum in Nautilus
pompilius Linnaeus. h, c: inner horny (h) and middle chalky (c) layers of the siphuncularwall, n: nacreous septal layer. Specimen from off Bindoy, Negros Oriental, the Philippines. S. Hayasaka and others colI. in 1981.
Fig. 2. Part of Fig. 1, showing the loosely packed, unoridntated aragonite needles of the
middle chalky layer, which are adapically replaced to the septal prismatic zone.
Fig. 3. Septal prismatic zone (pr) in Spirula spirula Lamarck. sn: long septal neck, s:
adapical surface of the septum. Specimen stranded on a beach near Noumea, New Caledonia. S. Mikami colI.
Fig. 4. Details of the septal prismatic zone (pr) of S. spirula, showing the continuous relation with the chalky (= spherulitic prismatic) layer (c) of siphunclar tube. Same specimen as that of Fig. 3.
Fig. 5. Pillar-like prisms (pr) forming the septal prismatic zone of Sepia esculenta Hoyle in
the early stage. sn: nacreous septal layer. Specimen from off Tsuyazaki, Kyushu.
Fig. 6. Strongly modified platy prisms of the septal prismatic zone in S. esculenta in thelate-growth stage. Same specimen as that in Fig. 5.
Photos by Y. Fukuda under 20 KV for an acceleration voltage.

TANABE, FUKUDA

and OBATA,'

Ammonite Siphuncular System

Plate 68

753.

Ammonite Siphuncular System

Mutvei, H. (1967): On the sUbmicroscopic
shell structure in some Jurassic ammonoids.
N. Jb. Ceol. PalCiont. Abh. 129, 157-166,
pI. 14.
- - and Reyment, R. A. (1973): Buoyancy
control and siphuncle function in am·
monoids. Palaeontology 16, 623-636.
Obata, I., Tanabe, K. and Fukuda, Y. (1980):
The ammonite siphncular wall: its micro·
structure and functional significance. Bull.
Natn. Sci. Mus. Tokyo, C6, 59-72, pIs. 1-3.
Raup, D. M. and Takahashi, T. (1967): Experiments on strength of cephalopod shells.
Abstr. Progm. geol. Soc. Amer. 172, 173.
Schindewolf, O. H. (1967): Analyse eines Ammoniten-Gehauses. Abh. Akad. Wiss. Lit.
math.-naturw. Kl. Jg. 1967 (8), 138-188.
Seilacher, A. (1975): Mechanische Simulation
und funktionelle Evolution des AmmonitenSeptums. PalCiont. Z. 49, 268-286.
Tanabe, K., Obata, I., Fukuda, Y. and Futakami, M. (1979): Early shell growth insome Upper Cretaceous ammonites and its

443

implications to major taxonomy. Bull.
Natn. Sci. Mus. Tokyo, C5, 153-176, pIs.
1-6.
Trueman, A. E. (1941): The ammonite bodychamber, with special reference to thebuoyancy and mode of life of the living
ammonite. Q. J. Ceol. Soc. Lond. 96, 339383.
Ward, P. D. (1979): Cameral liquid in Nautilus
and ammonites. Paleobiology 5, 40-49.
- - , Greenwald, L. and Greenwald, O. E.
(1980): The buoyancy of the chambered
Nautilus. Sci. Amer. 243, 190-203.
- - , - - and Magnier, Y. (1981): The
chamber formation cycle in Nautilus
macromphalus. Paleobiology 7, 481-493.
- - and Martin, A. W. (1978): On the buoyancy of the pearly Nautilus. J. Exp. Zool.
205, 5-12.
Westermann, G. E. G. (1971): Form, structureand function of shell and siphuncle incoiled Mesozoic ammonoids. Life Sci.
Contr. R. Onto Mus. 78, 1-39.

*1'. ~ 7 :/ -1:- 7"1 r ~ 2 f!fi0) Jl~~~-~~~f14~O) %nlt C:. ~1Jg: ;; "" 7 *,c.ii11M Eleganticeras elegantulum :to J: lfB!IE*-cf~M Reesidites minimus t;:Ii'iJ)E~:h7.>f*ffO) J: \,,~(.:t:h.:ftL r 1 -:;, ,,:/h"v!;f, :to J: lf~t#Jj;m:7J*:tt!!.fl.i1G.ilE) t;:"0\',-e, ltl!:{;.jq1;jfflHf.!:~z. SEM
z. ffl\"-e~WL., m~~ltl!:l!AfE~O) >b 0) C:. .It~L-t.: • .:to)f.551t J:8a 2 f!fiO)Jl~~~-~~~
m~It.:;t ? A 711 0) >b 0) C:. J: <~ftl"t Q .: UM-yiJ' --:d.:. ~*f.S*:to J: If:;t ? A 711 0)~3(t~*%nltt;:M"t Q liIf~nlt*iI' I?, "ltl!:Si:" O)%nltt;:~;l: IJ ~~~~}jO)7t7£,t;:~H;)Q~3(t~%nlto)-1:- T Iv (ilf! 1~) iI~i!lil'tLQ. m1'.:;t ry A 711 ~~ r !1" p :z ry 1 -j} -("It., %nltfi[f~O)~3(t~It.
lfn.~O)nlt7tt;:ftlt.:~{2f;: (cameral liquid) t;:: J: "? -ejjl!1jt.: ~tL -e:to IJ, ~~O)jCnltf~.:tO)~{2f;:It.
)1f&lft;:Jl~~~ J:&,:t;:j;JHl:l ~tLQ • .: 0)': C:. ii, I?, :to.:t I? <7 :/-1:-7"1 r ~I;::to\',-e >b~3(t~1t.~{2f;:-c:-jjl!1j~tL -e\"t.: C:. m)E~tLQ. .: O)~{2f;:IH~I;: "ltl!:&':" H!!E13 L., ~~~ffJ!iO)~:fL'i!t:tttt}jz.Jill. "? -e3(t~71-A..C:. j;J~1±l ~ tLt.: -c';5)J ? 7 :/-1:-7"1 r ~ C:. ftlt.:ltl!:{2f;:m~z.~"0m1'.:;try A -j}°1 O)~1li§%Jmil'I?~*-Q C:., 7 :/-1:-7"1 r ~O)3(t~~{2f;:It.m~~O)%nlto)~I;:Mf* L.,1'.{2f;:0)f'¥7tl;:It.f*l~ L.tJ:il' "? t.: C:. ,1[',;btLQ.
tJIIIl"$-nlt· mE8%'1'. • 'N~Hm~.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 128, p. 444, December 30, 1980

SHORT NOTES

20. HAYAMINA, A NEW NAME FOR NEUMAYRIAOHTA 1981, NON DE STEFANI, 1877*
YOSHIHISA OHTA
Department of Earth Sciences, Fukuoka University of Education,
Munakata, Fukuoka 811-41

The name Neumayria, recently proposed
as a genus of early Cretaceous corbiculids(Ohta, 1981), is a junior homonym of
Neumayria de Stefani, 1877 (p. 322), a
Pliocene gastropod from Italia. I hereby
propose the name Hayamina to replace
Neumayria Ohta, 1981 (p. 117).
I am indebted to Emeritus Professor
Tatsuro Matsumoto of Kyushu University
and Prof. Makoto Kato of Hokkaido University for bringing the homonymy to myattention, and for advice on nomenclature.

References
Ohta, Y., 1981: Some Lower Cretaceous Corbiculidae and Neomiodontidae (Bivalvia)
from Japan. Bull. Fukuoka Univ. Education, 31, [3J, 103-134, pis. 1-8.
Stefani, C. de, 1877: Molluschi Continentali
Fina ad ora Notati in Italia. Nei Terreni
Pliocenici, ed Ordinamento di Questi Ultini.
Atti Soc. Toscana Sci. Naturali, 3, 274325, pis. 17, 18.

*83 (1981, p. 117) It, E*O)T(fllSil!i:*iJ,~.i!i[I±li"Q Corbiculids O)-mH;::~~c VC
Neumayria ~!Ii!~l..,t.::iJ~, ;:o)~~lti""CK de Stefani (1877, p. 322) I;::;: ryilj~~h-c,-,t.::O)-r:, -t-hl;::~*-Q "to)c l..,-Crfit.::l;:: Hayamina ~!Ii!~i"Qo*83liR* Received September 29, 1982.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 128, pp. 445-451, pI. 69, December 30, 1982

754. OLDHAMINA FROM THE UPPER PERMIAN OF THE
KITAKAMI MOUNTAINS, JAPAN AND ITS TETHYANPROVINCE DISTRIBUTION*
JUN-ICHI T AZA WA
Institute of Geology and Paleontology, Tohoku University, Sendai 980

Abstract. Oldhamina kitakamiensis sp. nov. is described from the Upper Permian
Toyoma Formation in Kanayashiki, southern Kitakami Mountains, northeast Honshu,
Japan. This new species is close to Oldhamina anshunensis Huang, 1932 from theUpper Permian Lungtan and Dalong Formations of south China.
The genus Oldhamina is now known from the Permian of Armenia, Pakistan,
Burma, Thailand, China and Japan, all in the Tethyan Province.

Triassic Systems, 1975; Murata and Shimoyama, 1979) assigned the Kanayashiki
Sandstone Member to the lower part of
the Toyoma Formation (=Abadehian ofcentral Iran, after Murata and Shimoyama,1979) based mainly on molluscan fossils.
However, I consider that the member
belongs to the upper Toyoma Formation

Introduction
The purpose of this paper is to indicate
the presence of the aberrant strophomenide
Oldhamina in the Upper Permian of Japan.
The specimen described as Oldhamina
kitakamiensis sp. nov. was discovered in1973 by myself and Dr. N. Minoura fromsandstone float obtained in the lower
stream of Komasunaizawa valley, Kanayashiki, Yahagi-cho, Rikuzentakata City,
Iwate Prefecture, northeast Honshu, Japan(Text-fig. 1).
In the Yahagi district sandstone andconglomerate of the Kanayashiki Sandstone
Member (60 m thick in Kanayashiki) of
the Upper Permian Toyoma Formation are
well exposed along the Komasunaizawa
valley (Ehiro, 1974, 1977; Murata and Shimoyama, 1979). There is no doubt thatthe block of grey, coarse-grained sandstone
containing the brachiopod fossil was derived from this member.
Murata and his collaborators (Murata,
1969; Working Group on the Permian-t.6. Mt. Matsukurao!

* Received March 18, 1982.

Text-fig. 1.

445

2 km
,

Map showing the fossil locality.

Jun-ichi TAZAWA

446

and is correlated with sandstone beds ofthe latest Permian Nabekoshian age(=Changxingian of south China, after
Tazawa, 1975; Ishii et aI., 1975; Minato
et aI., 1978), which crop out on the southeastern slope of Mt. Nabekoshi, Kesennuma, 10 km south of Kanayashiki. This
result is based on the following stratigraphical and paleontological data: (1) thesandstones of both areas are lithologicallylike each other and they rest on thickshales probably from lower to middle partsof the Toyoma Formation (see Ehiro, 1977,
figs. 6, 10; Tazawa, 1976. fig. 1); (2) bothsandstones yield Eolyttonia d. E. nakazawai Shimizu (Kamada, 1979, p. 738;
Tazawa, 1975, p. 637), an element of theuppermost Permian fauna of the Kitakami
Mountains; and (3) the sandstone in Kanayashiki carries Oldhamina kitakamiensis
sp. nov. which is close to Oldhamina
anshunensis Huang, 1932 from the Upper
Permian Lungtan and Dalong Formationsof south China; the latter is equivalent to
the Changxing Formation (e.g. Zhao, Liang
Table 1.

and Zheng, 1978).
Distribution of Oldhamina

The genus Oldhamina is distributed in
the Permian of the Soviet Armenia, west
Pakistan, Burma, Thailand, China andJapan. There is no report of this genus
outside the Tethyan Province. The following 14 species are known, most ofthem occur in the Upper Permian Wujaping to Changxing Stages of south China(Table 1):
Oldhamina decipiens (de Koninck, 1863).
Middle to Upper Permian (Wargal Limestone and Chhidru Formation) of the Salt
Range, west Pakistan (de Koninck, 1863a,
1863b; Waagen, 1883; Noetling, 1904, 1905;
Reed, 1944), Upper Permian (Lungtan and
Dalong Formations) of Guizhou, south
China (Huang, 1932; Wang, 1955; Fengand Jiang, 1978).
Oldham ina d. O. decipiens (de Koninck,
1863). Middle to Upper Permian ("Fusulina
Limestone"=Wargal Limestone to Chhidru

Distribution of the genus Oldhamina.
Permian

11 ~I!

Region--.~-------------------:

O. decipiens
O. cf. decipiens
O. aff. decipiens
O. transkaukasica
O. squamosa
O. anshunensis
O. d. ansnunensis
O. regularis
O. grandis
O. hoshanensis
O. transversa
O. jiaozishanensis
O. subsquamosa
O. kitakamiensis

+
+

I

+
+

+
+
+
+
+
+
+
+
+
+
+
+

Pakistan (Salt Range), China (Guizhou)
Burma (Shan State)
Thailand (Thum Nam Maholan)
U.S.S.R. (Armenia)
China (Guizhou, Hubei, Sichuan, Fujian)China (Guizhou, Sichuan, Guangdong)
China (Anhui)
China (Hubei, Guangdong)
China (Guizhou, Guangdong)
China (Guangxi)
China (Qinghai)
China (Guizhou)
China (Guizhou), Japan (Kitakami Mountains)

751,.

Oldhamina from Japan

Formation of the Salt Range) of Shan
State, east Burma (Diener, 1911).
Oldhamina aff. O. decipiens (de Koninck,
1863). Lower Permian (Pseudoschwagerina
Zone to the lower Parafusulina Zone=
Sakmarian s.l.) of Thurn Nam Maholan,
central-north Thailand (Yanagida, 1967).
Oldhamina transkaukasica (Stoyanow,1916). Upper Permian (Dzhulfian s. s.) of
Armenia, U. S. S. R. (Stoyanow, 1916;
Ruzhentzev and Sarycheva, 1965).
Oldhamina squamosa Huang, 1932. Upper
Permian (Lungtan and Dalong Formations)of Guizhou, Hubei, Sichuan and Fujian,south China (Huang, 1932; Jin, Liao andFang, 1974; Zhang and Jin, 1976; Yang,
Ni, Chang and Zhao, 1977; Tong, 1978;
Feng and Jiang, 1978; Liao, 1979, 1980a).
Oldhamina anshunensis Huang, 1932. Upper Permian (Lungtan and Dalong Formations) of Guizhou, Sichuan and Guangdong,south China (Huang, 1932, 1933; Wang,
1955; Zhang and Jin, 1976; Feng andJiang, 1978; Liao, 1979, 1980a).
Oldhamina d. O. anshunensis Huang, 1932.
Upper Permian (Lungtan Formation) of
Anhui, south China (Zhang and Jin, 1961).
Oldhamina regularis Huang, 1932. Upper
Permian (Lungtan and Dalong Formations)of Hubei and Guangdong, south China(Huang, 1932; Yang, Ni, Chang and Zhao,
1977).
Oldhamina grandis Huang, 1932. Upper
Permian (Lungtan and Dalong Formations)of Guizhou and Guangdong, south China(Huang, 1932; Wang, 1955; Yang, Ni,
Chang and Zhao, 1977; Feng and Jiang,1978; Liao, 1980a).
Oldham ina hoshanensis Huang, 1936.
Upper Permian (Hoshan Formation) of
Guangxi, south China (Huang, 1936).
Oldhamina transversa Jin and Ye, 1979.
Upper Permian of Qinghai, northwest
China (jin, Ye, Xu and Sun, 1979).
Oldhamina jiaozishanensis Liao, 1980.
Upper Permian (Lungtan Formation) of

447

Guizhou, south China (Liao, 1980a).
Oldhamina subsquamosa Liao, 1980. Upper
Permian (Lungtan and Dalong Formations)of Guizhou, south China (Liao, 1979, 1980a,
1980b).
Oldhamina kitakamiensis Tazawa, sp.
nov. Upper Permian (Kanayashiki Sandstone Member, upper part of the Toyoma
Formation) of the Kitakami Mountains,
northeast Honshu, Japan.
Systematic descriptions

Order Strophomenida Opik, 1934
Suborder Oldhaminidina Williams, 1953
Superfamily Lyttoniacea Waagen, 1883
Family Lyttoniidae Waagen, 1883
Genus Oldhamina Waagen, 1883
Type species:- BelleropllOn decipiens deKoninck, 1863, from the Middle- and Upper
Productus Limestone (Wargal Limestone
and Chhidru Formation) of the Salt Range(de Koninck, 1863a, p. 8, pI. 3, figs. 1, 1a),by original designation of Waagen (1883,
p. 403).
Diagnosis :-Shell medium to large insize. Pedicle valve strongly convex toflattened, with many pairs of lateral septa
which are high, sharp and dipping to thefront.
Remarks: - The genus Oldhamina ischaracterized by its inward-thinning andforward-dipping lateral septa. This feature
was demonstrated by Noetling (1904, fig.
3; 1905, fig. 1) and subsequently pointed
to by Huang (1932, p. 73) as the genericdiagnosis.
The type species of Oldhamina has avery strongly convex pedicle valve andthis feature was often regarded as a
general character of this genus (e. g.
Waagen, 1883; Noetling, 1905; Muir-WoodJun-ichi T AZA W A

448

and Williams, 1965). However, there aresome species with weakly inflated or
almost flattened pedicle valve, such as O.
anshunensis Huang, 1932, O. hoshanensis
Huang, 1936 and O. jiaozishanensis Liao,
1980. The new species described herein
is a form of the less-convex type.
Oldhamina kitakamiensis sp. nov.
PI. 69, Figs. 1-6; Text·fig. 2.

Material: -One specimen (Holotype),
partial internal cast and mould of a pedicle
valve, lacking the anterior and posterior
portions, IGPS colI. cat. no. 97715.
Description :-Shell large in size, elongate subtrigonal in outline; length morethan 60 mm, width about 70 mm. Pedicle
valve slightly convex in lateral and anterior
profiles.
Interior of pedicle valve with regularly
and symmetrically arranged lateral septa
on both sides of median septum. In thepresent material 13 septa are counted on
one side of the pedicle valve. Lateral
septa thin, with acute crests, inclined
toward the front at angles of 60 -65
near the median septum, almost straightnear the lateral margins of the valve anddipping to the front at angles of 40 -50
in lateral profile (see Text-fig. 2; PI. 69,
Fig. 6). The spaces between them 3.5-4.2
mm in width. Median septum low andobscure.
Remarks:- This specimen is safely assigned to the genus Oldhamina on thebasis of its sharp and forward-dipping

.,
S

t S e

0

0

0

0

C~.?. .?_S
1 em

Text-fig. 2. Lateral septa of pedicle
valve of Oldhamina kitakamiensis sp. nov.,
longitudinal section.

lateral septa.
Oldhamina kitakamiensis seems closest to
Oldhamina anshunensis Huang, describedand figured by Huang (1932, p. 77, pI. 6,
figs. 6, 7; pI. 7, figs. 1-4) as Oldhamina
squamosa var. anshunensis from the upper
part of the Permian coal-bearing formation(Dalong Formation) of Guizhou, but thelateral septa of the Chinese species are
more convex forward than those of theKitakami species.
Oldhamina hoshanensis Huang from fossiliferous black limestone (Hoshan Formation) of Hoshan, Guangxi (Huang, 1936,
p. 491, pI. 1, figs. 2-4; text-fig. 2) resembles
O. kitakamiensis in having slightly convex
pedicle valve, but the former is distinguished from the latter by its lateral septa
which are more or less at right angles tothe median septum.
Oldhamina jiaozishanensis Liao, describedand figured by Liao (1980a, p. 262, pI. 4,
fig. 30) from the Lungtan Formation of
Guizhou somewhat resembles O. kitakamiensis in size, outline and curvature ofpedicle valve, but the Chinese species
differs from the present form in having
26 lateral septa on each side of the pedicle
valve.
Oldhamina decipiens (de Koninck) can beeasily distinguished from the present form
by its highly inflated pedicle valve.
Ehiro (1974, p. 460; 1977, p. 9) listed
Leptodus richthofeni Kayser from the
Kanayashiki Sandstone Member in thetype locality. Ehiro's specimen stored in
the Institute of Geology and Paleontology,
Tohoku University is represented by apartial internal mould of a pedicle valve.
The feature of its lateral septa strongly
suggests that the specimen can be assigned
to O. kitakamiensis.
Acknow Iedgements
I wish to thank Dr. Nachio Minoura of754.

Oldhamina from Japan

Hokkaido University for helpful discussion
on stratigraphy of the Yahagi district.
I am grateful to Professor Paul Copper ofLaurentian University, and Professor Kei
Mori of Tohoku University for criticalreading of the manuscript, and to Mr.
Shohei Otomo of Tohoku University fortaking photos.
References
Diener, C. (1911): Anthracolithic fossils of
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Ehiro, M. (1974): Geological and structural
studies of the area along the HizumeKesennuma Tectonic Line, in southern
Kitakami Massif. Jour. geol. Soc. Japan,
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Contr. no. 77, p. 1-37. (in Japanese)Feng, R. and Jiang, Z. (1978) : Phylum Brachiopoda. in Geological and PalaeontologicalTeam of Guizhou (ed.), PalaeontologicalAtlas of southwest China; Guizhou, Part
2. Carboniferous to Quaternary. p. 231-305,
pis. 85-108, Geological Publishing House,
Beijing. (in Chinese)
Huang, T. K. (1932): Late Permian Brachiopoda of southwestern China. Palaeont.
Sinica, Ser. B, vol. 9, fasc. 1, p. 1-138,
pis. 1-9.
- - (1933): Late Permian Brachiopoda ofsouthwestern China, Part 2. Ibid., vol. 9,fasc. 2, p. 1-172, pis. 1-11.
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Mountains of Texas with notes on lytton ids
from southwestern China. Bull. geol. Soc.
China, vol. 15, no. 1, p. 489-493, pI. 1.
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City Univ., vol. 19, art 6, p. 107-138, pis.
1-4.
Jin, Y., Liao, Z. and Fang, B. (1974): Permian

449

Brachihpoda. in Nanjing Institute ofGeology and Palaeontology (ed.), Palaeontological Atlas of southwest China. p. 308313, pis. 162-165, Science Press, Beijing.
(in Chinese)
Jin, Y., Ye, S., ,Xu, H. and Sun, D. (1979):
Phylum Brachiopoda. in Nanjing Instituteof Geology and Palaeontology and Geological
Institute of Qinghai (ed.), PalaeontologicalAtlas of northwest China; Qinghai, Part
1. p. 60-217, pis. 18-57, Geological Publishing House, Beijing. (in Chinese)
Kamada, K. (1979): The Triassic Inai Groupin the Karakuwa area, southern Kitakami
Mountains, Japan (Part 1) -Stratigraphy
and Paleogeography. Jour. geol. Soc. Japan,
vol. 85, no. 12, p. 737-751. (in Japanese)
Koninck, L. de (1863a): Descriptions of somefossils from India, discovered by Dr. A.
Fleming, of Edinburgh. Quart. Jour. geol.
Soc., London, vol. 19, pt. 1, p. 1-19, pis. 1-8.
- - (1863b): Memoire sur les fossiles paleozoiques recueillis dans l'Inde par M. Ie
Docteur Fleming. 44 p., 11 pis., Liege.
Liao, Z. (1979): Brachiopod assemblage zone
of Changhsing Stage and brachiopods from
Permo-Triassic boundary beds in China.
Acta Stratigr. Sinica, vol. 3, no. 3, p. 200207, pI. 1. (in Chinese)- - (1980a): Upper Permian brachiopods fromwestern Guizhou. in Nanjing Institute ofGeology and Palaeontology, Academia
Sinica (ed.), Stratigraphy and Palaeontology of Upper Permian coal-bearing formations in western Guizhou and eastern
Yunnan. p. 241-277, pis. 1-9, Science Press,
Beijing. (in Chinese)- - (1980b): Brachiopod assemblages fromthe Upper Permian and Permian-Triassic
boundary beds, south China. Can. Jour.
Earth Sci., vol. 17, no. 2, p. 289-295.
Minato, M., Kato, M., Nakamura, K., Hasegawa,Y., Choi, D. R. and Tazawa, ]. (1978):
Biostratigraphy and correlation of thePermian of Japan. Jour. Fac. Sci., Hokkaido Univ., Ser. 4, vol. 18, nos. 1-2, p.
11-47.
Muir-Wood, H. M. and Williams, A. (1965):
Strophomenida. in Moore, R. C. (ed.) ,Treatise 011 Invertebrate Paleontology,

450

Jun-ichi TAZAWA

Part H (Brachiopoda 1). p. H361-H521,
Geol. Soc. Amer., New York and Univ.
Kansas Press, Lawrence.
Murata, M. (1969): Molluscan fauna of the
Toyoma Formation (Late Permian). Saito
Ho-on Kai Mus. Nat. Hist., Res. Bull.,no. 38, p. 1-22, pis. 1-4.
- - and Shimoyama, S. (1979): Stratigraphynear the Permian-Triassic boundary andpre-Triassic unconformity in the Kitakami
Massif, northeast japan. Kumamoto Jour.
Sci. Ceol., vol. 11, no. 2, p. 11-31. (injapanese)
Noetling, F. (1904): Ueber den Bau und die
Organisation der Lyttoniidae Waagen.
Verhandl. d. Deutsch. Zool. Cesellschaft(1904), p. 103-122.
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Lyttoniidae Waag. emend.
Noetling.
Palaeontographica, Bd. 51, Lief. 4, p.
129-154, pis. 15-18.
Opik, A. A. (1934) : Uber Klitamboniten. Tartu
Univ. (Dorpat) , Acta Comm., Ser. A, vol.
26, no. 3, p. 1-239, pis. 1-48.
Reed, F. R. C. (1944): Brachiopoda and Mollusca from the Productus Limestones of
the Salt Range. Palaeont. Indica, N. S.,
vol. 23, no. 2, p. 1-678, pis. 1-65.
Ruzhentzev, V. E. and Sarycheva, T. G. (1965) :Evolution and change of marine organisms
at the boundary between the Palaeozoic
and Mesozoic. Trudy Paleont. Inst., vol.
108, p. 1-431, pis. 1-58. (in Russian)
Stoyanow, A. (1916):
On some Permian
Brachiopoda of Armenia. Trudy Ceol.
Kom., N. S., vol. Ill, p. 1-95, pis. 1-6. (inRussian and English)
Tazawa, j. (1975): Uppermost Permian fossils

from the southern Kitakami Mountains,
northeast japan. Jour. geol. Soc. Japan,
vol. 81, no. 10, p. 629-640, pis. 1-3.
- - (1976): The Permian of Kesennuma,
Kitakami Mountains: A preliminary report.
"Earth Science" (Chikyu Kagaku) , vol. 30,
no. 3, p. 175-185, pis. 1-3.
Tong, Z. (1978): Carboniferous and Permian
Brachiopoda. in Geological Institute ofsouthwest China (ed.), PalaeontologicalAtlas of southwest China; Sichuan, Part 2.
Carboniferous to Mesozoic. p. 210-267, pis.
77-92, Geological Publishing House, Beijing.
(in Chinese)
Waagen, W. (1883): Productus-Limestone fossils. Palaeont. Indica, ser. 13, vol. I, pt.
4, fasc. 2, p. 391-546, pis. 29-49.
Wang, Y. (1955): Brachiopoda. in Yang, K.
and Wang, Y., Index fossils of China;
Invertebrate, Part 2. p. 109-171, pis. 61103, Geological Publishing House, Beijing.
(in Chinese)
Williams, A. (1953): The morphology andclassification of the oldhaminid brachiopods.
Jour. Washington Acad. Sci., vol. 43, no.
9, p. 279-287.
Working Group on the Permian-Triassic Systems(Bando, Y., Ichikawa, K., Ishii, K., Kambe,
N., Kanmera, K., Minato, M., Murata, M.,
Nakazawa, K., Shimizu, D. and Toriyama,
R.) (1975): Stratigraphy near the PermianTriassic boundary in japan and its correlation. Jour. geol. Soc. Japan, vol. 81, no.
3, p. 165-184. (in japanese)
Yanagida, J. (1967): Early Permian brachiopods from north-central Thailand. in
Kobayashi, T. and Toriyama, R. (ed.),Ceology and Palaeontology of southeast

Explanation of Plate 69
Oldhamina kitakamiensis sp. nov.
Holotype, from sandstone float of the Kanayashiki Sandstone Member of the Toyoma Formation, lower stream of Komasunaizawa valley, Kanayashiki, Yahagi-cho, Rikuzentakata City, Iwate
Prefecture, northeast Honshu, Japan, IGPS coil. cat. no. 97715.
Figs. 1-3. Ventral, side and anterior views of internal mould of pedicle valve (xl).
Fig. 4. Latex cast of pedicle valve interior (x 1).
Fig. 5. Natural cast of pedicle valve interior (x 1).
Fig. 6. Longitudinal section of the latex cast of pedicle valve interior, showing lateral septaof the valve (x 3).

l'AZA WA,'

Oldhamina from Japan
Plate 695

754.

Oldhamina from Japan

Asia, vol. 3, p. 46-97, pIs. 11-23, Univ.
Tokyo Press, Tokyo,
Yang, D., Ni, S., Chang, M. and Zhao, R.
(1977): Phylum Brachiopoda. in Geological
Institute of Hubei et al. (ed.). Palaeontological Atlas of south-central China. Part
2. Late Palaeozoic. p. 303-470. pIs. 129-188.
Geological Publishing House. Beijing. (inChinese)Zhang, S. and Jin, Y. (1976): Late Paleozoic
brachiopods from the Mount Jolmo Lungma
region. in Tibetan Scientific Expeditional
Team. Academia Sinica (ed.). A report of451

scientific expedition in the Mount Jolmo
Lungma region (1966-1968); Palaeontology. fasc. 2, p. 159-242, pIs. 1-19. Science
Press. Beijing. (in Chinese)
Zhang, Y. and Jin, Y. (1961): An Upper
Permian brachiopod fauna from Jingxian.
Anhui Province. Acta Palaeont. Sinica,
vol. 9, no. 4, p. 401-417, pIs. 1-4. (inChinese)Zhao, J .• Liang. X. and Zheng. Z. (1978):
Late Permian cephalopods of south China.
Palaeont. Sinica, N.S., no. 12. p. 1-194,
pIs. 1-34. (in Chinese)

Iwate :!5-'f. Kanayashiki ~~iJ&. Kesennuma ~fl~t?:l. Kitakami ;ltl::,i*liR. Mt. Nabekoshi ~~ill. Rikuzentakata JliliMi%81, Toyoma

Komasunaizawa !N~

1"r*. Yahagi~('f-;ltl::Jli:lllil::tm-"')v .b. *il€lJliU[.~ Oldhamina t; J: U'II'U~O) T 7- .A:llli~i*l 7J;{ff: mlifl;ltl::llJ:Lill. :E'-'f!Jil<Jlilifltriit 81m ~fFII1J ~1'l9.(0) 'Hifi*liRl-:::.t;l'"C, l::tm-"')v.b. *~1'liJ.&Iij):!5tm~ (If*~l::tm) I-:::.Eb*-t7.JIij):!5O)ljiiEJ: I). JlJilJe.~ Oldhamina O)ftEb~ H/I!If*9BJiI.~.:ht~. ;:.:h'1c
Oldhamina kitakamiensis C -6lri5 L. ~cjrll(-t 7.J. *fllHt,*,~mtm, vcHI· gg)11 • $*~~O)
l::tm-"')v.b. *, Jei1-Wm t; J: U' :kflimb, G ¥liE • itc1l'lI( ~.:h"Cl' 7.J Oldhamina anshunensis
Huang, 1932 I-:::.:i&~ J: <{l;,("Cl'7.J.
Oldhamina a?E*T 7- .A:Lill~0) 7)v -" .=. 7, /{=\'-.A!':/. t." )V-7, !' -1, ,*,~O)-"')v.b.
*b' G 13fl1):b;ffi G.:h"Cl' t~b:, ;: 0) t~U' B *O)-"')v.b. *b' G¥fit~l-:::' 1 :f!M:1.m:t G.:ht~o81iRMl-Trans. Proc. Palaeont. Soc. Japan, N. S., No. 128, pp. 452-464, pis. 70-73, December 30, 1982

755. TWO NEW JURASSIC GENERA OF FAMILYP ALAEOSCENIDIIDAE (RADIOLARIA)*
A TSUSHI T AKEMURA
Department of Geology and Mineralogy, Faculty of Science,
Kyoto University, Kyoto 606, Japan
andKO]IRO NAKASEKO
Institute of Geological Sciences, College of General Education,
Osaka University, Toyonaka 560, Japan

Abstract. Exceedingly well-preserved Jurassic radiolarians have been obtained
from manganese ore deposits in the Mino Belt, central Japan. Among them, two
species of family Palaeoscenidiidae, Hilarisirex quadrangularis n. gen., n. sp. andDiceratigalea hemisphaera n. gen., n. sp., which have a median bar, two apical spinesand four basal spines, are described. They also belong to Nassellaria, because theirtests are composed of cephal is and thorax, and possess cephalic skeletal elements.
The new subfamily Hilarisisirecinae has been created for these two new genera.
Consequently, taxonomic position of family Palaeoscenidiidae must be re-examinned.
This family may be regarded as an ancestor of Nassellaria. Also, it is possible that
Nassellaria in the current classification is a polyphyletic taxon. The definition ofNassellaria must be re-examined. A lineage of "Paleozoic Palaeoscenidiids->Hilarisirex
quadrangularis->Diceratigalea hemisphaera" is proposed. If this process of evolution
is correct, the most important taxonomic criterion of Nassellaria is the pattern of
cephalic skeletal structure.

idae to suborder Spumellaria based on thesimilarity of skeletons between Palaeoscenidium and Haplentactinia.
Triassic
forms of Palaeoscenidiidae were describedby Dumitrica (1978). They possess one
apical spine and four basal spines within
spherical lattice shells. Furutani (1982)
carried out a detailed study on Paleozoic
palaeoscenidiids and revealed the processof evolution from Palaeoscenidium to forms
with spherical shells (Genus Tlecerina
Furutani etc.).
Two forms of Palaeoscenidiidae with

Introduction
Palaeoscenidiids are primarily Paleozoic
spicular forms with two to four apicalspines and three to four basal spines, reported from Silurian to Carboniferous(Deflandre, 1953; Foreman, 1963; Riedel,
1967; Holdsworth, 1977).
Holdsworth (1977) proposed systematics
of Paleozoic radiolarians, in which hequestionably assigned family Palaeoscenidi-* Received May 20, 1982.
452

755. New Jurassic Paleoscenidiidae

KYOTO

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140"
138"
134"

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Index map. Inuyama area (Text-Fig. 2) and Gujo-Hachiman area (Text-Fig. 3)

454

Atsushi TAKEMURA and Kojiro NAKASEKO

Text-Fig. 2. Locality map (Inuyama area). (site plotted on Geographical Survey Instituteof Japan top:>gtaphical map "Inuyama" of scale 1: 25000)

755. New J'/,~mssic Paleoscenidiidae455

T ex t·Fi g . 3. Loca lit y ma p (Gujo -Hachiman a rea ) . (s ite plotted on Geograph ical Sur vey
Institute of Japan topog r a phica l ma p "H achim a n" of sca le 1: 50000)

two apical spines and four basal spines
occurred in the radiolarian assemblage frommanganese ore deposits in the Mino Belt,
central Japan (Text-fig. 1- 3; Takemura
and Nakaseko, 1982a). Th is assemblage isnamed as Unu111 a echinatus Assemblage
and regarded as middle Jurassic (Yao,
Matsuda and Isozaki, 1980; Mizutani et at. ,1981).
These two forms a lso possess cephal is,456

Atsushi TAKEMURA and Kojiro NAKASEKO

thorax and cephalic skeletal elements,
same as Nassellaria. We regard them as
two new genera, Hilarisirex and Diceratigalea, of family Palaeoscenidiidae on thebasis of the pattern of basic skeletal
structure.
We wish to express our sincere thanks
to Professor Keiji Nakazawa, Professor
Tadao Kamei and Dr. Niichi Nishiwaki of
Department of Geology and Mineralogy,
Faculty of Science, Kyoto University for
their valuable advice. Special thanks are
due to Mr. Koji Wakita of Geological
Survey of Japan for his guidance in thefield.
Problems on phylogeny andclassification of Nassellaria
and Palaeoscenidiidae

In the present paper, we describe two
species, Hilarisirex quadrangularis n. gen.,n. sp. (Pis. 70-71 ; Text-fig. 4) and Diceratigalea hemisphaera n. gen., n. sp. (PI. 72 ;PI. 73, Fig. 1). These two species belong
to the family Palaeoscenidiidae, and at thesame time, possess characteristics of Nassellaria. Namely, the pattern of basic
skeletal structure accords with that of
Palaeoscenidiids (Text-figs. 4-5), and on
the . other hand, the external shapes ofshells, which are composed of cephalis
and thorax, are similar to those of nassellarians.
Deflandre (1953) and Foreman (1963) reported spicular form with flat lamellae,
named Palaeoscenidium cladophorum Deflandre.
Holdsworth (1977) illustrated
Silurian Palaeoscenidium composed of only
skeletons. Such various forms with spherical shells as Pentactinocarpus, Pactarentinia, Tlecerina and so on were describedby Dumitrica (1978) and Furutani (1982).
In the present paper, we regard Hilarisirex quadranqularis and Diceratigalea
hemisphaera as descendants of Paleozoic

Palaeoscenidium, based on the accordanceof the pattern of their basic skeletons withother Palaeoscenidiids. If not, only relying
upon the shell shape, it is very difficult
to clarify the phylogeny of other Palaeoscenidiids mentioned above. When the pattern of internal basic skeletal structure isadopted as a criterion, we can easily explain the phylogeny.
Hilarisirex quadrangularis is more similar to Paleozoic Palaeoscenidium in shapethan Diceratigalea hemisphaera. Furthermore, the wall structure and the wholeshape of D. hemisphaera resemble Mesozoic
Nassellaria such as Napora (PI. 73, Fig. 2).
Hence, the lineage of "Paleozoic Palaeoscenidiids --> Hilarisirex quadrangularis-->Diceratigalea hemisphaera" can be roughly
presumed. This lineage may be anotherone from that of Triassic Pentactinocarpinae.
From these two new genera and thislineage, three problems on phylogeny andclassification of Nassellaria and Palaeoscenidiidae have arised. They are 1. ontaxonomic position of family Palaeoscenidiidae, 2. on phylogeny of Nassellaria and3. on taxonomic criteria of Nassellaria.
1. On taxonomic position (If family

Palaeoscenidiidae
Holdsworth (1977) tentatively assigned
family Palaeoscenidiidae to suborder
Spumellaria. Dumitrica (1978) and Furutani (1982) support this view because offorms with spherical shells.
However, Hilarisirex quadrangularis andDiceratigalea hemisphaera possess cephalothorax and cephalic skeletal elements.
These two species obviously belong toNassellaria (Ehrenberg, 1875). Therefore,
the assignment of Palaeoscenidiidae to
Spumellaria must be re-examined.
2. On phylogeny of Nassellaria
Dumitrica, Kozur and Mostler (1980)

755.

New Jurassic Paleoscenidiidae

described Anisian or Ladinian (Triassic)
radiolarians, which are the oldest identified radiolarians in the Mesozoic Era. This
assemblage includes multi-cyrtoids, Pylentonemids and so on, as Nassellaria. De
Wever (1981) assigned Poulpus and Saitoum
to Pylentonemidae. But on the other hand,Holdsworth (1977) regarded family Pylentonemidae as Spumellaria.
The origin of Mesozoic multi-cyrtoids is
uncertain. Takemura and Nakaseko (1981)
described Permian Neoalbaillella and pointed out its similarity of internal structures
to Mesozoic multi-cyrtoids.
Although Hilarisirex quadrangularis andDiceratigalea hemisphaera belong to Nassellaria, they are regarded as descendantsof Palaeoscenidiids. Therefore, family
Palaeoscenidiidae may possibly be an origin
of Nassellaria.
But, as mentioned above, there areseveral possible forerunners of Nassellaria.
In this case, Nassellaria is a polyphyletictaxon and the definition of Nassellaria
must be changed. Furthermore, all current classification of Nassellaria must be
re-examined. In order to solve this problem, detailed studies on internal structures of Paleozoic and Mesozoic radiolarians
are required.
3. On taxonomic criteria of Nassellaria
Pessagno actively studied various Mesozoic nassellarians (Pessagno, 1969; 70; 76;
77a; 77b; Pessagno, Finch and Abott,1979). But he regarded shell structures ofpost cephalic chambers as a criterion offamily-grade classification.
Empson-Morin (1981) described Campanian radiolarians and stated that since
cephalic skeletal elements of most di- to
multi-cyrtoid Nassellaria
show only
minor variation, the taxonomic usefulnessof it is questionable. She maintained theview that an observational error by "optical
illusion" had been made with Amphipyndax.

457

Takemura and Nakaseko (1982b), however,showed the existence of Amphipyndacids
among Jurassic radiolarians by using SEM.
If the lineage of "Palaeoscenidiids-> H.
quadrangularis->D. hemisphaera" is correct,
the external shell has been markedly
changed through the process of evolution,
while the pattern of cephalic structure
remained constant. This fact indicatesthat cephalic skeletal structure is the mostimportant taxonomic criterion of Nassellaria and shell structure is secondary.
Systt'matic Paleontology

Subclass RADIOLARIA Muller, 1858
Order POL YCYSTINA Ehrenberg, 1838,
emend. Riedel, 1967
Family P ALAEOSCENIDIIDAE Riedel,
1967, emend. Holdsworth, 1977
Subfamily Hilaricirecinae Takemura
and Nakaseko, n. subfam.
Type genus :-Hilarisirex Takemura andNakaseko, n. gen.
Definition :-Palaeoscenidiidae with two
apical spines and four basal spines, possessing poreless and spherical cephalis in
the apical hemisphere and thorax in thebasal hemisphere. Median bar, two apicalspines and four basal spines composing
cephalic skeletal elements.
Remarks :-Forms of this new subfamily
possess characteristics of Nassellaria. But,
because of their pattern of basic skeletal
structures, it is clear that they belong tofamily Palaeoscenidiidae. Although formsof this subfamily are regarded as descendants of Paleozoic palaeoscenidiids (see theprevious section), they may possibly be on
another lineage from Triassic Pentactinocarpinae (Dumitrica, 1978).
Cephalic skeletal elements of this sub-

458

Atsushi TAKEMURA and Koji1"O NAKASEKO

family are composed of seven elements(Text-figs. 4-5). They may correspond to
those of Nassellaria; they are median bar(MB), apical spine (A), vertical spine (v),two lateral spines (L) and two secondary
lateral spines (1). Dorsal spine (D) do notexist. Four basal spines (two (L) and two(1)) arise primarily laterally from bothends of median bar. This pattern is notreported from other palaeoscenidiids.
This subfamily differs from other palaeoscenidiids in possession of cephalo-thorax,
and from other Nassellaria in its patternof basic skeletal structure.
Genus Hilarisirex Takemura
and Nakaseko, n. gen.

Type species :-Hilarisirex quadrangularis Takemura and Nakaseko, n. sp.
Diagnosis: - Cephal is possessing two
equal triradiate apical spines. Thorax, inshape a frustum of a quadrangular pyramid,
having thin and irregularly scattered walls
between feet. Four triradiate feet according with edges of the thoracic frustum
and protruding below aperture.
Remarks:-This genus is distinguishedfrom Diceratigalea Takemura and Nakaseko, n. gen. by its shapes of both two
apical spines and thorax. The genus namemeans hilarious king.

Hilarisirex quadrangularis Takemura
and Nakaseko, n. sp.
PI. 70, Figs. 1-2; PI. 71, Figs. 1-2;
Text-fig. 4

Description:-Test quadrangularly pyramidal with two apical spines and four
feet. Cephalis spherical, pore less with
uneven surface, from which two strong
and triradiate apical spines arise obliquely.
Each spine possessing three ridges, ofwhich two are on the apical side and theother one arise from near collar portion.
On the lower surface of cephalis, fourridges arising at the terminal point of
inner four basal spines, and lying obliquely
downward to terminate at four verticesof thoracic frustum. In some specimens,(PI. 70, Fig. 2a), many cephalic ridges lying
on cephalic surface of collar portion.
Thorax in shape a frustum of quadrangular pyramid of which four ridges
accord with four feet. At the apical endof thorax, near collar portion, four transverse ridges lying with small nodes at
each end, from which four feet arise below.
Four feet triradiate, possessing three ridgesof which the outer one accords with theedge of thoracic frustum. The other two
inner ridges on the inner side of thethoracic wall, are connected with those ofadjacent feet in arch-shape at the upper

Explanation of Plate 70(Abbreviations are same as Text-fig. 4.)
Figs. 1-2. Hilarisirex quadrangulalaris TAKEMuRA and NAKASEKO, n. sp.
Fig. 1. Holotype, A TJRMN-ll01-1, !NY -101
I-a, b. lateral view, stereophotographs (scale = 100 fl.)1-c, d. basal view, stereographs (scale=100 fl.)1-e, f. cephalic skeletal elements, stereophotographs (scale=lO II)
Fig. 2. Paratype, ATJRMN-ll03-1, TKN-105
2-a.
lateral view (scale= 100 fl.)2-b, c. cephalic skeletal elements, stereophotographs (scale =: 10 II)

TAKEMURA and NAKASEKO:

New Jurassic Paleoscenidiidae

Pla.te 70

l-c

l-d

2-b

2-c

755.

New Jurassic Paleoscenidiidae

459

a

c

t

Text·Fig. 4. Hilarisirex quadrangularis T AKEMURA and NAKASEKO n. gen., n. sp.
a: apical spine of Palaeoscenidiidae, b: basal spine of Palaeoscenidiidae, MB: median bar,
A: apical spine of Nassellaria, V: vertical spine, L: lateral spine, I: secondary lateralspine, c: cephalis, t: thorax, w: wall of thorax, p: pore, ap: aperture, f: foot.

Atsushi TAKEMURA and Kojiro NAKASEKO

460

A-~~:~

r ' G--v~)- ---L(1)r"'

A--Q .,.o--V{?)".
,
Q A{?)-9roH- --v

D--:.~:

I

J

I

~.\

--L

...

1MB

~. -

//1

I

J

J

.....

--L(1)

1(1) MB

A--

---L

.

.

;.

--v"O'. : t:\'~'/I \V~

0-"-

I'... ----L

.

J

,'.

I"'

I '

I

I

I

I

I

I

I
I

I MB

---V(1)---L(1)1(1) MB

Text-fig. 5. Cephalic skeletal structuresof Napora sp. (left) and Hilarisirex quadranqularis T AKEMURA and NAKASEKO n.
gen., n. sp. (right)
above: view toward side with vertical spine,
middle: side view from position perpendicular
to sagittal plane, below: view from apex,
D: dorsal spine. (The other abbreviations
are same as Text-fig. 4)

part of thorax (PI. 71, Figs. 2-b, c). At
the lowermost part of thorax, inner ridges
extend transversely to be connected withthe adjacent ridges, forming a square
framework in which aperture is enclosedwith a circular skeleton. A thin wall
scattered with circular pores connects boththe square frame and the circular skeletonaround aperture.
Four lateral sides of thoracic frustum,
where is surrounded by inner ridges offour feet, are covered by generally thin,swelled walls with irregularly distributed,
circular or elliptical pores. Four triradiate
feet protrude below aperture.
Cephalis possess seven skeletal elements,
a median bar two weaker apical spinesand four stronger basal spines (Text-fig. 5).
The median bar is shifted slightly fromthe center of collar plate. Two apicalspines arise obliquely upward from bothends of median bar, terminating at cephalicwall to be connected with outer triradiatespines of cephalis. Four basal spines run
laterally to cephalic wall, jointed with four
cephalic ridges on the wall surface andconnected with four feet.
Measurements: - Length of shell (exclusive of apical spines and feet), 100200 f-t ; maximum width of thorax, 90-110 f-t.
Remarks :-Because the median bar isshifted from the center of the collar plate,
one of two apical spines of inner cephalicskeletal elements is longer than the other.
Longer apical spine may correspond to

Explanation of Plate 71
Figs. 1-2. Hilarisirex quadrangularis T AKEMURA and NAKASEKO, n. sp.
Fig. 1. Paratype, ATJRMN-ll02-1, !NY-lOl
I-a.
lateral view (scale = 100 p)
1-b, c. basal view, stereophotographs (scale=50 p)
Fig. 2. Paratype, ATJRMN-ll04-1, TKN-105
2-a.
lateral view (scale=100 p)
2-b, c. basal view, stereophotographs (scale= 100 p)
2-d, e. cephalic skeletal elements, stereophotographs (scale = 10 p)

TAKEMURA and NAKASEKO:l-a

New Jurassic Paleoscenidiidae

Plate 71

l-b

l-c

2-b

2-c

2-d

2 -e

2-a

755.

New Jurassic Paleoscenidiidae

vertical spine (V) of Nassellaria, andshorter one to apical spine (A) of Nassellaria. Then, a pair of basal spines on theside of vertical spine (V) are lateral spines(L), and another pair on the opposite side
(side of apical spine (A)) are secondary
lateral spines (1) (Text-figs. 4-5).
Total shape of this new species is similar
to Paleozoic Palaeoscenidium. Probably,
this new species may have been evolved
from Paleozoic spicule-form palaeoscenidiids, by complication of skeletons and formation of cephalo-thorax.
Occurrence: - Jurassic manganese ore
deposits in the Mino Belt.
Genus Diceratigalea Takemura
and Nakaseko, n. gen.
Type species :-Diceratigalea hemisphaera
Takemura and Nakaseko, n. sp.
Diagnosis: - Cephalis possessing two
triradiate apical spines, of which one isweaker. Thorax hemispherical in shape,
possessing latticed shell. . Four feet triradiate, protruding below from aroundcircular aperture.
Remarks: - This genus differs from
Hilarisirex Takemura and Nakaseko, n.
gen. in possession of both an odd pair ofapical spines and hemispherical thorax.
The genus name means helmet with twohorns.
Diceratigalea hemisphaera Takemura

and Nakaseko, n. sp.
PI. 72, Figs. 1-2; PI. 73, Fig. 1

Description:- Test hemispherical with
two apical spines and four feet. Cephalis
spherical, pore less with uneven surface,from which two triradiate apical spinesarise. Each spine possessing three ridges,of which two are situated on the apicalside. One of two apical spines, strong

461

and long, arising obliquely laterally andcurving slightly downward. Another one
weak, short and straight, arising more
apically.
Thorax hemispherical with thick latticed wall. Pores generally circular or
elliptical, of which sizes and disposition
are irregular. Aperture at the base ofthorax, circular and large. Four feettriradiate and strong, extending obliquely
outward.
Each foot possessing threeridges, of which one is situated on theouter side. This outer ridges arising fromthe collar portion, where inner four basal
spines terminate, running downward within
the thoracic wall and protruding outward
at the lower part of thorax to form four
feet (PI. 72, Fig. 2-a). Inner two ridges offour feet arising from around aperture.
Cephalic skeletal elements, of which
combination and disposition are same as
those of Hilarisirex quadrangularis Takemura and Nakaseko, n. sp. (Text-fig. 5),
are composed of median bar, two apicalspines and four basal spines. Longer apicalspine is connected with outer stronger
triradiate one, and shorter with outer
weaker and straight one: Four basal
spines are connected with the outer ridges
of four feet at collar portion of cephalicsurface.
Measurements: - Length of shell (exclusive of apical spines and feet), 90-100 fl ;maximum width of thorax, 90-100 fl.
Remarks :-Correlation of cephalic skeletal elements of this new species withthose of Nassellaria is same as that ofHilarisirex quadrangularis Takemura andNakaseko, n. sp. (Text-figs. 4-5).
Although the total shape of this species
resembles that of Mesozoic tripod nassellarians such as Napora, their skeletal
structures are quite different (Text-fig. 5).
Napora possesses median bar (MB), apicalspine (A), vertical spine (V), dorsal spine(D), two lateral spines (L) and two second-462

Atsushi TAKEMURA and Ko}iro NAKASEKO

ary lateral spines (1). In Napora, prolongations of dorsal spine (D) and two lateralspines (L) construct three feet. Diceratigalea hemisphaera never possesses dorsal
spine (D) and prolongations of two lateralspines (L) and two secondary lateral spines(1) construct four feet. One specimen ofNapora (PI. 73, Fig. 2) has four feet, ofwhich three are prolongations of dorsal
spine (D) and two lateral spines (L). The
other one arise from around aperture independently of cephalic skeletal elements.
Total shape of external shell of Diceratigalea hemisphaera is more similar to
Mesozoic nassellarians than Hilarisirex
quadrangularis, of which shape resembles
Paleozoic Palaeoscenidium.
Therefore,
Hilarisirex may be an intermediate form
between Palaeoscenidium and Diceratigalea,
and D. hemisphaera may have been evolved
from Hilarisirex.
Occurrence: - Jurassic manganese ore
deposits in the Mino Belt.
Refe:-ences
Deflandre, G. (1953): Radiolaires fossiles. in
Crasse, P. P. ed., Traite de Zoologie. Paris,
Masson, vol. 1, pt. 2, p. 389-436.
De Wever, P. (1981): Une nouvelle sousfamille, les Poulpinae et quatre nouvelles
especes de Saitoum, Radiolaires mesozoiques tethysiens. Ceobios, no. 14, fasc. 1,

Lyon, p. 5-15.
Dumitrica, P. (1978): Triassic Palaeoscenidiidae and Entactiniidae from the Vincentinian
Alps (Italy) and Eastern Carpathians(Romania). D.S. Inst. Ceol. Ceojiz., vol.
64, pt. 3, paleont., p. 39-54.
- - , Kozur, H. and Mostler, H. (1980): Contribution to the radiolarian fauna of the
Middle Triassic of the Southern Alps. Ceol.
PalCiont. Mitt. Innsbruck. Bd. 10, no. 1,
p. 1-46.
Ehrenberg, C. G. (1875):
Fortsetzung der
mikrogeologischen Studien als GesammtUebersicht der mikroskopischen Palaontologie gleichartig analysirter Gebirgsarten
der Erde, mit specieller Rucksicht auf den
Polycystinen-Mergel von Barbados. Abh.
preuss. Akad. Wiss., Jahrg. 1875, p. 1-226,
pis. 1-30.
Empson-Morin, K. (1981): Campanian Radiolaria from DSDP Site 313, Mid-Pacific
Mountains. Micropaleont., vol. 27, no. 3,
p. 249-292, pis. 1-13.
Foreman, H. P. (1963): Upper Devonian Radiolaria from the Huron member of the Ohio
shale. Micropaleont., vol. 9, no. 3, p. 267304, pis. 1-9.
Furutani, H. (in press): Middle Paleozoic
Palaeoscenidiidae (Radiolaria) from Mt.
Yokokura, Shikoku, Japan. Part I.
Holdsworth, B. K. (1977): Paleozoic Radiolaria;
Stratigraphic distribution in Atlantic borderlands. in Swain, F. M. ed., Stratigraphic
micropaleontology of Atlantic basin andborderlands, Amsterdam, Elsevier, p. 167184.

Explanation of Plate 72(Abbreviations are same as Text·fig. 4.)
Figs. 1-2. Diceratigaiea hemisphaera TAKEMuRA and NAKASEKO, n. sp.
Fig. 1. Holotype, ATJRMN-ll03-2, TKN-105
I-a, b. lateral view, stereophotographs (scale = 100 p)
I-c.
weaker apical spine (scale=lO p)
I-d.
basal view (scale=100 p)
Fig. 2. Para type, A TJRMN-ll04-2, TKN-105
lateral view (scale = 100 p)
2-a.
2-b, c. basal view, stereophotographs (scale= 100 p)
2-d, e. cephalic skeletal elements, stereophotographs (scale = 10 p)

TAJ(EMURA and NAJ(ASEJ(O:New Jurassic Paleoscenidiidae

l-a

l-b

Plate 72

l-c

2-b

2-d

l-d

2-c

2-e

755.

New Jurassic Paleoscenidiidae

Mizutani, S., Hattori, I., Adachi, M., Wakita,K., Okamuta, Y., Kido, S., Kawaguchi, I.
and Kojima, S. (1981) : Jurassic Formationsin the Mino Area, Central Japan. Proc.
Japan Acad., vol. 57, ser. B, no. 6, p. 194199.
Pessagno, E. A. Jr. (1969): The Neosciadiocapsidae, a new family of Upper CretaceousRadiolaria. Bull. Amer. Pal., vol. 58, no.
253, p. 377-439, pis. 23-38.
.(1970): The Rotaformidae, a new familyof Upper Cretaceous Nassellaria (Radiolaria)
from the Great Valley Sequence, California
Coast Ranges. Bull. Amer. Pal., vol. 58,
no. 257, p. 5-32, pis. 1-9.
- - (1976): Radiolarian zonation and stratigraphy of the Upper Cretaceous portion of
the Great Valley Sequence, California Coast
Ranges. Micropaleont. Spec. Publ., no. 2,
p. 1-95, pis, 1-14.
- - (1977a): Upper Jurassic Radiolaria andradiolarian biostratigraphy of the California
Coast Ranges. Micropaleont., vol. 23, no.
1, p. 56-113, pis. 1-12.
- - (1977b): Lower Cretaceous radiolarianbiostratigraphy of the Great Valley Sequence and Franciscan Complex, California
Coast Ranges. Cushman Found. Foram.
Res., Spec. Paper, no. 15, p. 1-87, pis. 1-12.

- - , Finch, W. and Abott, P. L. (1979): Upper
Triassic Radiolaria from the San Hipoolito
Formation, Baja California. Micropaleont.,
vol. 25, no. 2, p. 160-197, pis. 1-9.
Riedel, W. R. (1967):
Protozoa (subclass
Radiolaria). in Harland, W. B. et al., eds.,
The fossil record. Geol. Soc. London, p.
291-298.
Takemura, A. and Nakaseko, K. (1981): A
new Permian radiolarian genus from theTamba Belt, Southwest Japan. Trans. Proc .
Palaeont. Soc. Japan, N. S., no. 124, p.
208-214, pis. 33-34.
- - and - - (1982a): Jurassic Palaeoscenidiids from manganese ore deposits in the
Mino Belt and their significance on evolution and classification of Nassellaria.
JRS 81 OSAKA, Proceedings of the firstJapanese Radiolarian Symposium. Newsof Osaka Mictopaleontologists, Spec. vol.
no. 5. (in Japanese with English abstract)- - and - - (1982b): On cephalic structuresof Jurassic Nassellaria. ibid. (in Japanesewith English abstract)
Yao, A., Matsuda, T. and Isozaki, Y. (1980):
Triassic and Jurassic radiolarians from the
Inuyama area, central Japan. Jour. Geosci.,
Osaka City Univ., vol. 23, p. 135-154, pis.
1-3.

:; "" '7 *'['. Palaeoscenidiidae H:1.&Iit.li!.O) 2 ~ji" :fULJ:I!l!.f¥IG&UmJ:J\.~:I!l!.f¥IGO)~I#Hff4t·~~~4tK~~~~~~ff~.*b·G~, :;,,"'7e4tMO)~O)t~~Gh~, ~b~~.~O)
J:\":1.&Iilc.li!.!lU~-z.iliHI:l~~. -to)4tb'G, Palaeoscenidiidae HI;:'ji,~~ Hilarisirex quadrangularis n. gen., n. sp. tt. GUI;:', Diceratigalea hemisphaera n. gen., n. sp. 0) 2 fi-z.
;: h G 2 ji,1:t., Nassellaria!ll! § O)l1:Jt-z. ~b-lt ~ '0t::.~, ~!II!H Hilarisirecinae-z.~P~ L. t::.. -to)~*, Palaeoscenidiidae HO)~MB9(ft~&U Nassellaria O)~~jH;:' '0\,,~:pf~iM~~~'~i.J~~ t:t::.. Nassellaria 1:t.~*tiIC-e~~ RTfjgl1:b~~ IJ, -to)~~, 1t01(0)~M-z.t&*i.J·GJ!@:~tt.fthfitt.Gtt.\',. "it::., ;:hG Palaeoscenidiidae f40)*tiIC:fe~-z.~~~t, Nassellaria O)~M£i\I!t L.~I:t., cephalis O)1t,UJ~b~J'&~lI!~-e~~;: ci.J~1f-~~h~~ L. t::..

~.

463

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Atsushi T AKEMURA and Koiiro NAKASEKO

Explanation of Plate 73(Abbreviations are same as Text·fig. 5.)
Fig. l. Diceratigalea hemisphaera T AKEMURA and NAKASEKO, n. sp. Para type, ATJRMN·1l04·3,
TKN·105
I-a.
lateral view (scale = 100 /1)
1-b, c. cephalic skeletal elements, stereophotographs (scale = 10 /1)
1-d, e. basal view (scale=100 /1)
Fig. 2. Napora sp. (possessing four feet) ATJRMN·1l04·4, TKN·105
2-a, b. lateral view, stereophotographs (scale = 100 /1)
2-c, d. cephalic skeletal elements, stereophotogtaphs (scale= 10 /1)2-e, f. basal view, stereophotographs (scale=50 /1)

TAKEMURA and NAKASEKO:

New Jurassic Paleoscenidiidae

I-b

2-c

I-d

2-e

Plate 73

l-c

2-d

l-e

2-£

465
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Bivalve fossils from the Miyanohara Formation (Lower Cenomanian), Sakawa area,
Shikoku ...................... Matsuda, T.
-gj1!UIf<1IllR~WO) Monotis 0) {tE1il' %w'll'!:{tcr:p:i.f~ Nasselaria ml1Rilt£E.O)jiHtc *~I:'''::>\''-C .................. ¥TN JJ PJ • r:piltii"$l'XHf>.................................. ~Ji:"1if~!/foipGO) Cassidulinidae (fl".:fL£E.) .... !l!fNWx

Record of an ammonite from the Shimanto
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..... .. ... Matsumoto, T. and Tamura, M.
Neomphaloceras Matsumoto et Obata (:I&:)1iW
p~ U-:7 ,/-1::~1 ~ O)~Jm) I:'''::>\''-C .. t'l*;i¥Hf>
Cretaceous nautiloids from Hokkaido-IV. A
new nautiloid species from the Cenomanianof Hokkaido
A. Taxonomy ....................... .
. ... Matsumoto, T. and Takahashi, T .
B. Ontogeny .. Obata, r. and Futakami, M.
Buccal mass structure of Gaudryceras (Cretaceous Ammonoidea) ................... .
................ Tanabe, K. and Fukuda, Y.
The jaw apparatuses of some Cretaceous
desmoceratid ammonites ........ Tanabe, K.
Internal shell structure of the Cretaceous
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Crassostrea gigas (Thunberg) from themarin Pleistocene of the west coast areaof Ise Bay, Southwest Japan ........... .
................ Yamada, J. and Honda, Y.
Paleontological significance of Neverita(Neverita) asagaiensis (Makiyama, 1934)(Gastropoda: Naticidae); a revised nameof Ampullina asagaiensis ............... .
· .. ... Honda, Y. and Marincovich, L. N. jr.

Two interesting ammonoid species from the
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. ............... Nishida, T. and Kyuma, Y.

A new genus Pentactinosphaera from the
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Nishimura, A., Nagata, K. and Nakaseko, K.
Some radiolarian fossils from the Upper Cretaceous formations in the Shimonto Belt.......... Yamauchi, M. and Nakaseko, K.
Radiolarians from the chert member of the
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I..." .. " ........... Kimura, T. and Aiba, H.

466
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SYSTEMATIC INDEX(New Series No. 121-No. 128)

PALAEOZOOLOGY
Protozoa
Page
Takemura, Atsushi and Nakaseko, Kojiro: A new Permian radiolarian genus from the Tamba
belt, Southwest Japan .................................................................. 208
Takemura, Atsushi and Nakaseko, Kojiro: Two new Jurassic genera of family Palaeoscenidiidae (Radiolaria) .................................................................... 452

Coelenterata
Igo, Hisayoshi and Adachi, Shuko: Three species of Lithostrotion from the Ichinotani Formation (Upper Paleozoic corals from Fukuji, southeastern part of the Hida massif,
part 5) ................................................................................ 179
Kato, Makoto: Mazaphyllum (Rugosa) from the Silurian of Japan ........................ 386
Nakai, Hitoshi: Silurian Corals from the Yokokurayama Formation in the Mt. Yokokura
region, Kochi Prefecture, Southwest Japan-Part 1. Halysitidae ........................ 139
Omura, Akio: Uranium-series age of the "Kametsu Formation", Riukiu Limestone on the
Tokuno-shima, Ryukyu Islands .......................................................... 327

Brachiopoda
Copper, Paul, Tazawa, Jun-ichi, Mori, Kei and Kato, Makoto: Atrypa (Devonian Brachiopoda) from Japan ...................................................................... 368
Tazawa, Junichi: Oldhamina from the Upper Permian of the Kitakami mountains, Japan andits Tethyan province distribution ...................................................... 445

Mollusca
Bando, Yuji and Ehiro, Masayuki: On some Lower Triassic ammonites from the Osawa
Formation at Asadanuki, Towa-cho, Tome-gun, Miyagi Prefecture, Northeast Japan .... 375
Furuichi, Mitsunobu: A new nautiloid species from the Upper Cretaceous Izumi Group of
Shikoku ................................................................................ 334
Hanai, Tetsuro: A story of the belemnite-developmental approach to fossil morphology .. 421
Hayami, [taru and Kase, Tomoki: Cenomanian molluscs in a sandstone block from the sea
bottom off the southern coast of Kuji, Northeast Japan ................................ 29
Honda, Yutaka: Corbiculid Mollusca from the Urahoro Group, Kushiro coal field, eastern
Hokkaido .............................................................................. 14
Honda, Yutaka: A new Clinacardium from the Omagari Formation of the Ombetsu Group,
Kushiro coal field, eastern Hokkaido .................................................... 127

467

468

Systematic IndexKanie, Yasumitsu: Cretaceous tetragonitid ammonite jaws: a comparison with modern
Nautilus jaws ........................... " ............................................. 239
Matsumoto, Tatsuro, Morozumi, Yoshiro, Bando, Yuji, Hashimoto, Hisao and Matsuoka,
Atsushi: Note on Pravitoceras sigmoidale Yabe (Cretaceous heteromorph ammonite) .. 168
Matsumoto, Tatsuro, Muramoto, Kikuo, Hirano, Hiromichi and Takahashi, Takemi: Some
Coniacian ammonites from Hokkaido.................................................... 51
Nakano, Mitsuo and Okamoto, Kazuo: A new species Acesta (Plicflcesta) watanabei from
the Miocene in Southwest japan ........................................................ 357
Ohta, Yoshihisa: Hayamina, a new name for Neumayria Ohta 1981, non De Stefani 1877 .. 444
Tanabe, Kazushige, Fukuda, Yoshio and Obata, Ikuwo: Formation and function of thesiphuncle-septal neck structures in two Mesozoic ammonites .......................... 433
Tanabe, Kazushige, Obata, Ikuwo and Futakami, Masao: Early shell morphology in someUpper Cretaceous heteromorph ammonites .......... , ................................... 215
Tashiro, Masayuki and Matsuda, Tomoko: The bivalve fossils from the Cretaceous Fukigoshi
Formation of the Monobe area, Shikoku ................................................ 393

Arthropoda
Hu, Chung-Hung: Studies on the ontogenic and phylogenic development of two Upper Cambrian trilobites from South Dakota .................................................... 159
Nohara, Tomohide: Notes on three Cytherelloidea ostracodes from the Ryukyus ............ 122
Nohara, Tomohide and Nakasone, Noriko: Sexual dimorphism of the paleocopid ostracodegenus Manawa from Okinawa-jima. " ................................................... 364
Yamaguchi, Toshiyuki: japanese Miocene cirriped Balanus sendaicus: a comparison with
Tethyan Balanus concavus group ...................................................... 277

Echinoderma ta
Tanaka, Keisaku and Kozai, Takeshi: Some Cretaceous echinoids from the Monobe area,
central Shikoku ........................................................................ 341

PALAEOBOTANY
Kimura, Tatsuaki and Tsujii, Masanori: Early jurassic plants in japan. Part 3 ............ 187
Kimura, Tatsuaki and Tsujii, Masanori: Early jurassic plants in japan. Part 4 ............ 259
Kimura, Tatsuaki, Yoshiyama, Hiroshi and Ohana, Tamiko: Fossil plants from the Tamaand Azuyama Hills, southern Kwanto, japan............................................ 87
Ling, Hsin Yi: Crassicorbisema, a new silicoflagellate genus, from the southern oceans andPaleocene silicoflagellate zonation ......................................................
1
Takahashi, Kiyoshi: Miospores from the Eocene Nanggulan Formation in the Yogyakarta
region, central java .................................................................... 303
Takahashi, Kiyoshi and Matsuoka, Kazumi: Neogene microfossils of Chlorophyceae, Prasinophyceae and Acritarchs from Niigata, central japan .................................... 105

469

INDEX OF GENERA AND SPECIES
Notes: Words listed are names of genus- and species-groups, which are described, discussed
or illustrated in the articles in Transactions and Proceedings of the Palaeontological Society ofJapan, New Series, Nos. 121-128. Words in bold-faced type are names of newly proposed taxa.

A
Acanthohalysites kuraokensis ............ 152
Acant hotrigonia mashikensis ............ 408
Acila (Trullcacila) mOllobellsis .. 397, pI. 62
Acila (Truncacila) sp. aff. A. demessa .. 397

Acesta (Plicacesta) watallabei ....... .
............................ 360, pIs. 57, 58
Acesta (Plicacesta) sp. d. A. (P.)
watanabei ...................... 361, pI. 57
Alnus d. japonica .................. 95, pI. 9
Alsophilidites kerguelensis ........ 307, pI. 49
Anaptychus sakhalinensis ................ 248
Anaptychus tenuiliratus ........ 244, 245. 248
Anaptychus tenuiliratus var. lata ........ 248
Angulobracchia (?) sp ................. pI. 34
Anomia sp. aff. Anomia foldia . ... 405, pI. 65
Anomoza11lites sp. A .................... 260
Anthonya sp. d. A. apicalis .... 38, pI. 4, 409
Anthonya 11lifunensis .. , .............. '" 409
Anthonya sp. d. Anthonya 11lifunensis ..
................................ 409, pI. 65
Arca (Eonavicula) sp ............. 402, pI. 64
Astarte sp ..................... , ......... 410
Cf. Asterotheca okafujii .......... 197, pI. 31
Atrypa (Planatrypa) .................... 370
Atrypa (Planatrypa) japonica .... 370, pI. 59
Atrypa desqua11lata mut. japonica . .... '" 370
Atrypa pallciplicata . ............. , ....... 370
Avellana sp ......................... 43, pI. 5
B
Balanus concavus ........................ 286
Balanus rostratus ...................... 286
Balanus senaaicus ............ 286, pis. 44-48
Baltisphael'idium llakajoellse .. 118, pI. 14
Baltisphaeridium sphaeroides .. 117, pI. 14
Barroisiceras ............................ 60
Barroisiceras (Barroisiceras) onilahyense.................................. 60, pI. 8
Barroisiceras onilahyense ................ 60

Batissa sitakaraensis .................... 18
Birostrina nipponica ...................... 32
Breviarca unisulcata .................... 400
Brevitricolpites circularis ...... 312, pI. 50
Buxus japonica .......................... 96
Buxus 11licrophylla var. japonica ....... .
. ............................. 96, pis. 9. 11c"Callista" (Pseuda11liantis) crenulatus ....
.................................... 39, 410
Circe tolwdai ............................ 20
Cladophlebis clavatum .................. 202
Cladophlebis haiburnensis ................ 199
Cladophlebis kurumellsis .. 201, pIs. 31, 32
Cladophlebis nebbensis .................. 197
Cladophlebis pseudodelicatula .. 201, pIs. 30-32
Cladophlebis raciborskii ...... 203, pIs. 31, 32
Cladophlebis raciborskii f. integra ...... 197
Cladophlebis d. raciborskii . . . . . . . . . . . . .. 197
Cladophlebis sp. a ...................... 201
Cladophlebis sp. A ...................... 203
Cladophlebis sp. b ...................... 201
Cladophlebis sp. B ...................... 203
Cladophlebis sp. C ................ 203, pI. 32
Clathropteris 11leniscoides ........ 190, pI. 30
Clathropteris obovata .................... 190
Classites capucinni . ............. ,. 309, pI. 50
Clavapal11laedites sp ............... 309, pI. 49
Clillocardium omagariellse...... 130, pI. 15
COlliopteris lleiridalliellsis . ..... 188, pI. 30
Corbicula (Batissa) sitakaraensis . ... 18, pI. 2
Corbicula (Corbicula) kotakai .... 22, pI. 2
Corbicula (Corbicula) tokudai ...... 20, pI. 2
Corbicula atrata tokudai .................. 20
Corbicula sitakaraensis .................. 18
Corbicula tokudai ........................ 20
Corbisema apiculata var. iner11lis .......... 2
Corbisema constricta ................ 4, pI. 1
Corbise11la cuspis .................... 3, pI. 1
Corbisema disym11letrica angulata .......... 3

Index of Genera and Species

470

Corbisema disymmetrica communis ........ 4
Corbisema disymmetrica disymmetriea ... .
· ................................... 6, pI. 1
Corbisema geometrica . ..................... 2
Corbisema inermis ........................ 5
Corbisema inermis crenulata .............. 5
Corbisema inermis disymmetrica .......... 6
Corbisema inermis inermis .......... 2, pI. 1
Corbisema navicula constricta .............. 4
Corbisema navicula navicula .............. 3
Corbisema naviculoidea .............. 3, pI.
Crassicorbisema .......................... 4
Crassicorbisema disymmetrica .............. 5
Crassicorbisema disymmetrica disymmetrica· ................................... 5, pI. 7
Crassicorbisema disymmetrica
dumitricae ............................ 5
Crassostrea kawauchidensis ...... 405, pI. 63
Crassostrea ct. kawauchidensis .......... 405
Ctenopteris sarrani ... " ............. , ... 259
Ctenozamites sarrani ............ 259, pI. 41
Cupuliferoidae pollenites fallax .. 311, pI. 50
Cymatiosphaera pulchella ...... 118, pI. 14
Cymbophora sp. . ................... 39, pI. 5
Cymbophora ? sp. .. .............. 409, pI. 65
Cyrena sp. . ............................. 20
Cyrillaceaepollenites minor . ....... 315, pI. 15
Cytherelloidea .......................... 122
Cytherelloidea asatoensis ................ 123
Cytherelloidea hanaii .................... 123
Cytherelloidea munechikai .............. 123
Cytherelloidea senkakuensis . ............. 123

D
Diceratigalea
461
Diceratigalea hemisphaera ........... .
· ........................... 461, pIs. 72, 73
Dicolpopollis malesianus .......... 310, pI. 50
Dictyocha navicula ... ..................... 3
Dictyocha naviculoidea .................... 3
Dictyocha apiculata var. inermis .......... 4
Dictyocha triacantha var. inermis .......... 2
Dietyocha triacantha var. inermis f.
inermis .............................. 2, 5
Dictyophyllum kotakiense ...... 192, pI. 30
Dictyophyllum sp. . ..................... 192
Drepanochilus minimus .......... 42, pI. 5
E
Elegant iceras elegantulum ........ 434, pI. 67

Entolium sp. ct. Entolium obovatum ....
.. .. .. .. .. .. .. .. .. . ... 34, pI. 4, 403, pI. 65
Eomiodon matsubasensis .......... 410, pI. 65
Eosturia ................................ 382
Eosturia towaensis .............. 382, pI. 60
Epiaster ................................ 343
Epiaster nobilis . ......................... 343
Epiaster sp. aft. E. nobilis ...... 343, pI. 55
Eubostrychoceras japonicum ...... 216, pI. 35
Eubostrychoceras ct. E. saxonieum................................ 217, pI. 36
Euphorbiacites sp ................. 316, pI. 50
Eutrephoceras .......................... 335
Elltrephoceras tawaense ........ 336, pI. 54

F
Falsicatenipora .......................... 144
Falsicatenipora shikokuensis .. 144, pIs. 16, 17
Follicucullus scholasticus . ............. pI. 34
Follicucullus ventricosus .............. pI. 34
Follicucullus ct. ventricosus .......... pI. 34
G

Gaudryceras spp. . ............... 242, pI. 39
Gleditsis horrida ........................ 96
Gleditsia japonica ........................ 96
Gleditsia ct. japonica .............. 96, pI. 9
Glycymeris (Glycymerita) sp. aft. G. (G.)
hokkaidoensis .................... 31, pI. 4
Glycymeris (Pseudoveletuceta) sp. ct.
11lifunensis .................... 402, pI. 62
Gonio11lya (Gonio11lya) subarchiaci ........ 41
Gonio11lya (Gonio11lya) sp ........... 41, pI. 5
Goshoraia crenulata .... 39, pI. 5, 410, pI. 65
Graminidites pllnctulosus ...... 319, pI. 52

H
Hausmannia (Protorhipis) sp ............. 196
Hausmannia (Protorhipis) sp. A ........ 196
Hayamina .............................. 444
Heteraster .............................. 344
Heteraster 11lacroholcus .......... 344, pI. 55
Heteraster nexilis ................ 345, pI. 55
Heteraster sp .. " " ............... 345, pI. 55
Halysites ................................ 149
Halysites catenularius ... ..... 150, pIs. 16, 19
Halysites kuraokensis ............ 152, pI. 18
Halysites suessmilchi ........ 151, pIs. 17, 18
Halysites sp. A .................. 154, pI. 19
Halysites sp. B .............. 155, pIs. 18, 19

Index of Genera and Species
Hilarisirex ............................ 458
Hilarisirex quadrangularis .. 458, pIs. 70,71
I!lex corn uta ...................... 98, pI. 11!lexpollenites tertiarus . ........... 316, pI. 51
Inoceramus (Birostrina) concentricus
nipponicus . .................. " ......... 32
Inoceramus (Birostrina) sp. d. I. (B.)
concentricus nipponicus .......... 32, pI. 4
Inoceramus concentricus nipponicus ...... 32
Inoceramus concentricus var. nipponicus .. 32
Inoceramus sp. . ......................... 32

J
Jugl1ns cinerea .......................... 94
Juglans megacinerea ... ................... 94
Juglans cinerea var. megacinerea .. 94, pI. 9
L

Laevigatosporites javanicus .... 307, pI. 49
Laevigatosporites (?) sp ........... 308, pI. 49
Lancettopsis sp ................... 117, pI. 13
Legumen ? sp ..................... 411, pI. 65
Leiosphaeridia d. fastigatirugosa . . 115, pI. 13
Leiosphaeridia grandiformis . ... 116, pI. 13
Leiosphaeridia minuscula ...... 116, pI. 13
Leiotriletes maxoides minoris .... 307, pI. 49
Lima (Acesta) aff. amaxensis ............ 360
Lima (Acesta) d. smithi ................ 360
Lima sp ................................. 360
Lithostrotion (Lithostrotion) decipiens ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 180, pI. 26
Lithostrotion (Siphonodendron) hidense .. 183
Lithostrotion (Siphonodendron)nipponalpinum ................ 182, pI. 27
Lithostrotion asiatica . .. " ............... 183
Lithostrotion decipiens .................. 180
Lithostrotion heptagonale ......... ....... 180
Lithostrotion portlocki .................. 180
Lithostrotion (?) proliferum ............ 182

M
Madagascarites ryu .............. 219, pI. 39
Magnoliaepollenites ellipticus .. 309, pI. 49
Manawa ................................ 365
Manawa konishii ........................ 365
Margarita funiculata .................... 41
Margarites ? funiculatus .......... 41, pI. 5
Matsumotoa unisulcata ... ......... 400, pI. 62

471

Matsumotoa unisulcata densestriata ...... 400
Mazaphyllum . " ......................... 387
Mazaphyllum mirum ... ......... 390, pI. 61
Mesosaccella mifunensis .......... 398, pI. 62
Metasequoia d. glyptostroboides .... 91, pI. 9
Micrhystridium ariakense ........ 117, pI. 14
Muramotoceras yezoense .......... 220, pI. 37
Myrtea (s. I.) sp. . ............... 408, pI. 65
N

Nanonavis pseudocarinata .. 400, pIs. 62-64
Navicula sp ............................. 400
Neoalbaillella .......................... 210
Neoalbaillella gracilis ...... 213, pIs. 33, 34
Neoalbaillella ornithoformis .... 211, pI. 33
Neoalbaillella sp ................. 213, pI. 34
Neoanaptychus tenuiliratum ........ 245, 247
Neoanaptychus sp ................... 244, 247
Neocrioceras spinigerum . ......... 220, pI. 37
Neohibolites miyakoensis .......... 423, pI. 66
Neumayria .............................. 444
Nippononectes kozaii ... ........... 404, pI. 63
Nucula (Pectinucula) kochiensis ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 396, pI. 62oOldhamina
447
Oldhamina kitakamiensis ...... 448, pI. 69
Ostrea sp. . ............................. 405
Otozamites fujimotoi .................... 261
Otozamites molinianus .................. 261
Otozamites neiridaniensis .. 261, pIs. 42, 43
Otozamites sp ........................... 261
pPalambages sp. . ................. 114, pI. 14
Paliurus ? sp ................ " ........... 98
Parapopanoceras kokeni .................. 380
Parobolinoides contractus ........ 160, pI. 20
Paronella (?) sp ....................... pI. 34
Parvamussium cowperi yubarense ........ 404
Pecten (Propeamussium) cowperi var.
yubarensis ............................ 404
Pecten (Syncyclonema) d. obovatus ...... 34
Pecten (Syncyclomena ?) aff. obovatus .... 34
Peroniceras .............................. 52
Peroniceras latum ............ 52, pIs. 6, 7
Peroniceras yubarense ........ 55, pIs. 6, 7
Petalaxis portlocki ...................... 180
Phlebopteris sp. A ................ 187, pI. 30

472

Index of Genera and Species

Picea maximowiczii ...................... 90
Picea d. maximowiczii ............ 90, pI. 9
Pinus fujiii ........................ 91, pI. 9
Pollenites fallax ... ............... 311, pI. 50
Polygalacidites speciosus ...... 318, pI. 52
Portlandia (s.l.) nagaseana .... 399, pI. 62
Pravitoceras sigmoidale ...... 169, pis. 22-26
Procarnites . ............................. 379
Procarnites acutus ...................... 380
Procarnites kokeni . ............... 380, pI. 60
Procarnites aff. kokeni .................. 380
Propeamussium cowperi yubarense ...... 404
Proteacidites d. annularis ........ 320, pI. 53
Proteacidites matsuokae ...... 319, pI. 53
Proteacidites mollis .............. 319, pI. 52
Proteacidites similis .............. 320, pI. 53
Pseudamiantis crenulatus ............ 39, 410
Pseudamiantis ? erenulata . ........... 40, 411

Pseudowashitaster .................... 346
Pseudowashitaster iaponieus ...... 351, pI. 56
Pseudowashitaster mysticus ......... .
............................ 348, pis. 55, 56
Pseudowashitaster (?) sp. . ........... pI. 55
Pterophyllum propinquum . ............... 265
Pterophyllum aff. propinquum .......... 265
Pterophyllum ex gr. propinquum ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . .. 265, pis. 41-43
Pterophyllum sp. A ...................... 267
Pterosper11lella pterina . ........... 114, pI. 14
Pterotrigonia (Acant hotrigonia) higoensis........................................ 36
Pterotrigonia (Aeanthotrigonia) longiloba........................................ 36
Pterotrigonia (Acanthotrigonia) sp. d.
P. (A.) longiloba ................ 36, pI. 4
Pterotrigonia (Acanthotrigonia)
11lashikensis ........................ 36, 408
Pterotrigonia (Aeant hotrigonia ?)11lashikensis .................... 408, pI. 64
Pterotrigonia (Acanthotrigonia) sp ....... 37
Pterotrigonia (Pterotrigonia) brevicula ..
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34, pI. 4
Pterotrigonia (Scabrotrigonia) imanishii .. 406
Pterotrigonia (Scabrotrigonia ?) imanishii................................ 406, pI. 64
Pterotrigonia (Scabrotrigonia) monobeana....................................... 407
Pterotrigonia (Scabrotrigonia ?)11lonobeana ................ 407, pis. 64, 65
Ptilophyllu11l ............................ 268

Ptilophyllum cutehense .................. 269
Ptilophyllum d. cutehense ........ 269, pI. 43
Ptilophyllum nipponicum .. 270, pis. 41-43
Ptilophyllum shinadaniense .... 270, pI. 41
Ptilophyllum sp. A .............. 271, pI. 43
Ptilophyllu11l sp. B .......... 272, pis. 41, 43
Q

Quercoidites ellipsodeus ........ 311, pI. 50
Quercoidites umiensis ............ 310, pI. 50
Quercus sp ......................... 95, pI. 9
R
Reesidites mini11lus ............. . 434, pi. 67
Retricolporites protensus
316, pI. 51

S
Salix sp ........................... 94, pI. 10
Salix d. integra .................. 92, pI. 10
Seabrotrigonia i11lanishii ................ 406
Sealarites mihoensis .............. 223, pI. 38
Scala rites sea/aris ................ 222, pI. 38
Scala rites (?) aff. S. sealaris .... 224, pI. 38
Schedohalysites .......................... 146
Sehedohalysites kitakamiensis.. 146, pis. 17, 19
Schedohalysites sp ....................... 146
Schloenbachia (Barraisiceras)
haberfellneri .......................... 60
Septifer d. 11lifunensis .......... 403, pI. 63
Siphonodendron hidense .................. 183
S11lilaeipites eehinatus ............ 308, pI. 49
Smilacipites spinulifer ... ....... 308, pI. 49
Sornayceras .............................. 57
Sornayceras proteus ..... ................. 57
Sornayeeras sp. d. S. proteus ...... 57, pI. 7
Sphenopteris sp. A .............. 199, pI. 31
Striatricolporites agustinus ...... 317, pI. 51
Striatricolporites striolatus .... 317, pI. 51
Striatricolporites sp ............... 317, pI. 51
Stylaxis portlocki . ....................... 180
Subtriporopollenites minutulus .. 321, pI. 53
Subtriporopollenites sp ........... 321, pI. 53
Subtriporopollis specialis ...... 322, pI. 53

T
Taenicephalus shu11lardi . . . . . . . . .. 162, pI. 21
Tetraeolporopollenites obscurus .. 318, pI. 52
Tetragonites glabrus . ............. 246, pI. 40
Thau11latopteris braun ian a .............. 194
Thaumatopteris elongata .... 194, pis. 30, 31

Index of Genera and Species
Thaumatopteris nipponica ................ 194
Thaumatopteris schenki . .............. '" 194
Thetironia affinis ........................ 409
Thetironia affinis japonica ................ 37
Thetironia affinis var. japonica .......... 37
Thetis japonica ............. '" 37, pI. 5, 409
Thetis sp. aff. Thetis japonica .... 409, pI. 65
Tiliaepollenites d. punctulosus .... 323, pI. 53
Tiliaepollenites tropicus . ....... 322, pI. 53
Tiliaepollenites (?) sp. . ......... 323, pI. 53
d. Todites fukutomii ............ 197, pI. 31
d. Todites yamanoiensis. . . . . . . . .. 199, pI. 32
Trapa bicerata .......................... 100
Trapa macropoda .................. 99, pI. 9
Trapa maximowiczii .............. 100, pI. 9
Tricolpites minutireticulosus . ..... 312, pI. 50
Tricolpopollenites elongatus . ... 311, pI. 50
Tricolpopollenites liblarensis ............ 311
Tricolporopollenites consularis consularis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 313, pI. 51
Tricolporopollenites javanensis . ...... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 313, pI. 51

473

Tricolporopollenites marginatus· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 313, pI. 51
Tricolporopollenites minor .............. 315
Tricolporopollenites ovatorotundus· .............. " .. " .. " ....... 314, pI. 51
Tricolporopollenites rasus . ..... 314, pI. 51
Tricolporopollenites tertiarius .......... 316
Tricolporopollenites umiensis ............ 310
Tricolporopollenites sp. a314, pI. 51
Tricolporopollenites sp. b314, pI. 51
Tricolporopollenites sp. c
315, pI. 51
Tricolporopollenites sp. d315, pI. 51
Trigonia brevicula ...................... 34
Trigonia pocilliformis .................. 406
Trilites (AlsoPhilidites) kerguelensis .... 307
Triporopollenites sp. a .......... 321, pI. 53
Triporopollenites sp. b .......... 321, pI. 53
Tytthodiscus densiporosus.. 113, pIs. 12, 13
Tytthodiscus densiporosus
densiporosus .................. 113, pI. 12
Tytthodiscus densiporosus minus . .... .
· ....... , ................... 114, pIs. 12, 13