ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 111

Palaeontological Society of Japan
Sept. 30, 1978

Itar u HA YAM I
Editor
Associate edito'rs Ts unemasa SAITO and Tom owo OZAWA

Officers for 1977 - 1978
Honorary President:

Teiichi KOBAYASHI

President : Ryuzo TORIY AMA
Councillors:

Kazuo AS AMA, Kiyotaka CHI NZEI, Takashi HAM ADA, Tetsuro HAN AI,

Itaru HAY AMI, Syozo HAY ASAKA, Hisayos hi lGO, Tadao KAMEl, Kametoshi
KANMERA, Tamio KOTAKA, Tatsuro MATSUMOTO, T s ugio SHU TO, Yokichi
TAKAYANAG I, Toshimasa TANAI, Hiroshi UJIIt
Members of Standing Committee:

Kaz uo ASAMA (Finance), Kiyotaka CHI NZEI

(General Affairs), Takashi HAMADA (Foreign Affairs), Tetsuro HANA I, It aru HAY AMI (Transactions), Hisayoshi IGO (Planning), Kametoshi KANIVIERA(Special

Papers),

Ikuwo

OBATA (Membership),Yo kichi

T AKA YANAG I(" Fossils"), Hiroshi UJIIt, Juichi YANAG ID A
Secretaries:

Ienori FUJlY AMA (S pecial Papers), Kunihiro ISHIZAKI ("Fossils"),

Tsunemasa SAITO

andTomowo OZAWA (Transactions),

Hiromichi

HIRANO and Yasumitsu KANlE (Membership), Toshiyuki YAMAGUCHI(General Affairs)
Auditor:

Tatsuaki KIMUR A

The fossil on th e co ve r:

Original figure of Podozamites R einii GEYLER ,1877, from the Tetori group.

GEY LER'S description marked the onset ofmodern palaeontology in Japan.

All communications relating to this Journ a l s hould be addressed to the
PALAEONTOLOGICAL SOCIETY OF JAPAN,
c/ o Business Center for Academic Societies,
Yayoi 2-4-16, Bunkyo-ku, To kyo 113, Japan.
Sole agent:

Uni versity of Tokyo Press, I-Ion go 7-3-1, Tokyo , Japan.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. Ill, pp. 333-347, pis. 45, 46, Sept. 30, 1978

691. ONTOGENETIC VARIATION OF TRIGONIOIDES (S.S.)PAUCISULCATUS (CRETACEOUS NON-MARINE BIVALVIA)*SEONG YOUNG YANG
Department of Earth Science, Kyungpook National University
Daegu, 635, Korea.

Abstract. Trigonioides (s. s.) paucisulcatus shows positively the ontogenetic
variation on its pseudocardinal teeth and V-shaped surface ornamentation. The
number of the pseudocardinal teeth increases with growth, that is, from two on
both valves (5, 3, PIIl/4, 2, PH, PlY) in immature specimens, to four or five on
both valves (5,3, la, Ib, Ic, PlH/(6), 4, 2, I'a, I'b, PlI, PlY) in adult specimens.
The V-shaped ornamentation is gradually effaced with growth. From the comparison with T. (s. s.) kodairai from the lower stratigraphic horizon, it can besaid that the phylogenetic trend is read at least on three characters; 1) The
angle of the V-ribs on the surface decreases. 2) The surface ornamentation iseffaced. 3) The number of the pseudocardinal teeth increases.

pelecypod genera in the upper Mesozoic
of Asia. However, its evolutionary trend,
phylogeny in one sense, has not beenclearly understood yet. This deficiency
has come firstly from that the relative
stratigraphic positions of the fossil localities have not been confirmed, which is inturn because of their separate occurrencesin dispersed basins, and secondly fromthat their genetic characters and theirvariations have not been grasped.
The two species, T. (s. s.) kodairai andT. (s. s.) paucisulcatus, are found fromsuccessively exposed Gyeongsang Group,
Korea; the former is from the lower partand the latter from near the uppermost
horizon. The comparison between thf"
two, therefore, is quite important forunderstanding the evolutionary trend of
the present genus. Recently I have foundseveral new localities of T. (s. s.) paucisulcatus near its type-locality and made afairly good collection. Present work is toreport some of the results of the study

Introduction
Trigonioides (s. s.) paucisulcatus was proposed by SUZUKI (1940) as a subspeciesof T. kodairai KOBAYASHI and SUZUKI(1936) based on the specimens from Baegandong, Yeongcheon-gun, Korea. At
that time, the type-specimens were sostrongly deformed and fragmental thatits internal structures, especially of thehinge teeth, could not be completely described.
Subsequently, OTA (1959a) has reported
its occurrence from Goshonoura (Cenomanian), Japan and raised it up to an independent species. But the Japanese specimen is also poorly preserved, so OTA
himself left its hinge teeth undescribed.
As previously mentioned (YANG, 1974),
the genus Trigonioides is one of the mostinteresting and important non-marine* Received September 30, 1977; read January 30, 1976, at Naruko.

333

s. Y. YANG

334

- - - - y +-----~~_I_36°

B:BUSAN

0: DAEGU

G: GYEONGJUU:ULSANY : YEONGCHEONo!
Text·figure 1.

20!

Index map.

40

60Km

691.

Ontogenetic variation of Trigonioides paucisulcatuson the new acquisitions, putting special
stress on the ontogenetic variation of thehinge structure and other characters.
Acknowledgements :-1 would like to express my sincere gratitude to Emeritus
Professor Tatsuro MATSUMOTO of Kyushu
University for his continuous encouragement on my study and critical readingof the typescript of this paper, Professors
Ki Hong CHANG and Yeong Gi KIM of
Kyungpook National University for theirintroducing new fossil localities. My sincere thanks are also due to Mr. Gilsu
LEE of the latter University for his assistance in field and laboratory works.

Notes on Stratigraphy
Since T ATEIWA (1929) first mapped the335

Gyeongsang Group, the area around thefossil localities has been surveyed fromtime to time by several geologists (SONet. aI., 1969, CHANG, 1970, PARK et. aI.,
1973). However, the stratigraphy of thearea in the section of Waegwan-DaeguYeongcheon-Gyeongju has not been concretely established yet. This is becauseof lack of a pertinent key bed and little
paleontological studies about the group.
I have briefly investigated the geology
around the fossil localities. It is illustrated in a simplified form in Text-figure
2. In order to show the relative stratigraphic horizons of the two species, ascheme of subdivision of Gyeongsang
Group is presented in Table 1.
The Sinla Subgroup was correlated by
SUZUKI (1940) to the Gyliakian (Ceno-

Table 1. Subdivision of Gyeongsang Group.
JAYANGSAN FM.
P-.

~

GEOCHEONRI FM.

0

JAIN FM ....... T. (s. s.) paucisulcatus

l'J
rl:l

SONGNAEDONG FM.

~

P-.

~

0

~

l'J
0

Z
<G
(f)

~

(f)

<G

....l

(f)

SINLA CONGLOMERATE FM.

Z
~

><

0

BANYA WEOL FM.
HAMAN FM.

....l......

0
0

DAEGU FM.**P-.
0~

CHILGOG FM.

ZOJINJU FM.*

0 0
0rl:l
<G~

HASANDONG FM ... ···· T. (s. s.) kodairai

O~

Z(f)

NAGDONG FM.*

*CI-IANG (1966) proposed Yeonhwadong and Dongmyeong formations for Nagdong (Naktong) and Jinju (Shinshu) formations, respectively, which were named by TATEIWA (1929),
on account of remote distance from the type-section Waegwan-Daegu-Gyeongju. Recently
CHANG (1975) withdrew his proposal by reason of priority.
**Daegu formation (by TATEIWA, 1929) is subdivided into Haman, Banyaweol andSongnaedong formations in ascending order.
The porphyrite formations, that is, the so-called Hagbong (Kakubo), Chaeyagsan (Saiyakusan) and Jusasan (Shushazan), are excluded from the stratigraphy because of theirintrusive relation to the adjacent formations. However, the extrusive features also arefound here and there. And it has been in dispute that the porphyrite formations must be
considered of stratigraphic units or not.

s. Y. YANG

336

manian-Turonian) Mifune Group in Japan
based on the common occurrence ofThiara (Siragimelania) tateiwai. The fossiliferous Lower Mifune is now referredto Middle to Upper Cenomonian (TAMURA
and MATSUMURA, 1974; MATSUMOTO,
1977). And, as pointed out above, T. (s. s.)
paucisulcatus also was reported by OTA(1959a) from the Cenomanian Upper Goshonoura Group (see MATSUMOTO, 1977).
Thus, in the present state, the Sinla Subgroup (strictly speaking, the fossil bed ofJain Formation, upper part of Sinla Subgroup) can be roughly correlated to Upper
Cretaceous (Cenomanian-Turonian).

35"55'

w'-

-"~"

,-

_ _ _ _ _ _-\ v,",=,-=~,,>\~;

'c::'

:v

v

V
-', V

v

v

V

v
V

'I

v

',---------,,'==,~y'-,,-~,,,,---~"-'~_Qliv

GE NCHEONAr -Text.figure 2. Geological map around the fossil localities.
1. Quaternary (Alluvium), 2. Bulgugsa granite. 3. porphyrite.
4. Jain Formation. 5. Songnaedong Formation.

Material
Among the fossil localities, Loc. 6 isthe type locality of T. (s. s.) paucisulcatus
designated by SUZUKI (1940) and theothers are new. I have visited several
times the type-locality, but I have foundno molluscan fossils except a gastropod,

Thiara (Siragimelania) tateiwai. The occurrence of fossils at the six localities isreviewed in Table 2.
As shown in Table 2, the fossils arecommonly found in siltstone bed and theoccurrence is not even but varies withlocalities. The present species occurs
gregariously at Loc. 2 and 5, but has not691. Ontogenetic variation of Trigonioides paucisttlcatus

337

Table 2. Fossil occurrences and lithology at six fossil localities.
Loc.

Lithology

1light grey siltstone

T. (s. s.) paucisulcatus

abundant2

silicified light grey
siltstone

T. (s. s.) paucisulcatus
Thiara (Siragimelania) tateiwai

gregarious
abundant

3

light grey siltstone

T. (s. s.) paucisulcatus
T. (5.) tateiwai

rarefrequent4

grey sil tstone
dark grey siltstone

T. (5.) tateiwai
Estheria sp.

rare
frequent

5

light grey siltstone

T. (s. s.) paucisulcatus
T. (5.) tateiwai

gregarious
rare

6

light grey siltstone

T. (s. s.) paucisulcatus
T (5.) tateiwai

rare, not foundgregarious

Fossils

been found at Loc. 4 and 6. The siltstone
beds yielding fossils seem to be of nearly
the same stratigraphic horizon of lower
Jain Formation.
More than 500 specimens of T. (s. s.)
paucisulcatus have been collected fromthe above four localities (Loc. 1, 2, 3 and
5). The specimens are deposited at the
Department of Earth Science, Kyungpook
National University, Daegu, Korea.

Observation and Recognition of
Ontogenetic Variation.
Since the fossiliferous beds are considered of nearly the same stratigraphic
horizon, as mentioned above, and no specific difference can be confirmed among
the samples from the different localities,
it is evident that all of the trigonioidid
samples fall within the same species.
The typical trigonioidid V-sculpture
looks fairly variable at a glance. In
general, the immature specimens (smaller
than 2 cm in length) are ornamented allover the valve with the sculpture, while
the adult specimens (larger than 4 cm)
show complete effacement on the valve
except weak and tough radial ribs on the

Abundance

posterior side. However, it can be safely
pointed out that these features are only
ontogenetic, for the intermediate specimens (from 2 to 4 cm) shows the transitional state of the sculptur!=! between theabove two. On the angle of the Vsculpture, no marked variation has been
observed.
Most of the specimens possess wellpreserved test except several of natural
mould state and are secondarily deformed.
The internal structures have been observedon the 55 specimens from which the tests
were dissolved out by dilute hydrochloric
acid. The muscle scars, i. e., adductor
scars and pedal scars, and the ventral
crenulation are observed constantly. The
anterior adductor scar is more prominent,
but smaller than the posterior one. The
ventral crenulation changes its characterwith its position on the ventral margin.
That is, on the posterior ventral margin
it is fairly distinct while it becomes
weaker towards the anterior and ultimately
effaced on the anterior end. The crenulation is fine on the anterior side and roughand coarse on the posterior side, reflecting the coarseness of the radial ribs on
the surface. However, no difference has

338

S. Y. YANG

been observed on these internal structures
between immature and adult ones.
On the other hand, fairly large variation of the hinge structure has been observed in accordance with the size of thespecimens as shown in figures of Plate 46.
On the smaller ones, the hinge structure
is considerably simpler than the larger
ones which possess a larger number ofhinge teeth. When the specimens arearranged in order of size, these features
show a gradual change which must be
regarded as ontogenetic variation.
Ontogeny is generally understood as alife history of an organism and necessarily
includes the development from the larvalstage to the senility. And the recognition of the larval stage is most importantin biological sense. However, in molluscan
fossils, of which the initial stage. i. e.,trochophore plus veliger, is not usuallypreserved, the ontogeny must be inevitably
considered as partially after the deposition of the prodissoconch. Moreover,when the hinge structures of the Bivalvia
are concerned, which cannot be observed
even on the prodissoconch, the term
'ontogenetic' may be rather inappropriate
in a strict sense. However, in such acase, the ontogeny is practically used ina broad sense as a continuous change withgrowth.
On the Dental Formulae.- The well-known
BERNARD'S scheme designed to give a
concise expression on the bivalve hinge
teeth and their disposition has been considered to be fairly suitable in regard to
the taxonomy and phylogeny in some ofthe heterodonts based on the homologous
dental structure. The adoption of thescheme to trigonioidids was made first by
Cox (1955). Subsequently, its usage had
been current among the authors of trigonioidids. HAYAMI in HAYAMI and ICHIKAWA (1965) maintained that the BERNARD'S
scheme for the heterodonts should not be

used for the trigonioidids. However, afterthat it has been also used for trigonioidids
without examination of the homology
between the heterodonts and the nonheterodonts besides a little modification.
As pointed out also by BOYD and NEWELL(1969), 'the scheme is difficult to apply
widely even to heterodonts as well as
non-heterodonts because homologies commonly cannot be readily recognized among
dissimilar bivalves'. In BERNARD'S formula, the cardinal teeth 1, 2 and 3 aremeant to be ontogenetically originatedfrom the anterior lateral teeth AI, All
and AlII, respectively. But in trigonioidids
without any anterior lateral teeth, there
is no yardstick on deciding which is 1 or
2, etc. Consequently, the formulae may
be changeable due to which to decide
arbitrarily. In order to reduce such a
confusion, OTA (1975) proposed to nominate
and fix the pseudocardinal tooth as 3
which is associated with the pedal scar inright valve and to arrange other symbols
in order, that is, 5, 3, 1a and 1b fromanterior extremity toward ventral on rightvalve and 6. 4, 2, l'a and l'b on left
valve.
In some of the present specimens (seePI. 46, figs. 14c, 15b), there appear obviously three pseudocardinals under the
Ota's 3 in right valve. In such a case,it seems to be needed that another symbol
1c should be provided for presenting the
new tooth ventrally found. The symbols
6, 5, l'a and l'b are considered quite
strange as well as 1c in usual BERNARD'S
notation. The 'a' and 'b' are commonly
provided for the teeth emerging from one
cardinal divided anteriorly and posteriorly,
respectively, while in trigonioidids theymean only order. In short, the current
method of naming teeth in trigonioidids
is quite peculiar in comparison to that inheterodonts.
BoYD and NEWELL (1969) proposed a

691.

Ontogenetic variation of Trigonioides paucisulcatus

new method of recording hinge teeth with
1 and 0. And they presented several
merits in their new system of being freefrom implication of uncertain homologies
by avoiding specification of homologues
for poorly understood group and of being
able to be converted readily version of
BERNARD'S system whenever homologies
are secure. However, there are somedemerits also in their method. When all
the hinge teeth and the sockets are recorded only with 1 and 0, it is difficult topoint a certain one among the teeth,
especially in trigonioidids of which hinge
teeth are variable both with ontogeny andphylogeny. And as in their formulae thedisposition is reversed anterior to right,
posterior to left in contrast to BERNARD'S
scheme where anterior is to left, posterior
to right, the abrupt adoption to trigonioidids may give rise to another confusion. It is rather convenient to use theconventional BERNARD'S scheme withpointing out that the usage in trigonioidids
has not relation to homologues of theheterodonts and is independent fromphylogenetic relativity between.
In this paper, the both schemes aretentatively used for comparison.

To return to the main subject: the hinge
structure of the present species can be
expressed as Table 3. The left column isdue to the modified conventional BERNARD'S
scheme in trigonioidids and the rightcolumn due to the new system of BOYD
and NEWELL.
The length of the hinge teeth and thecrenulation on the both of anterior andposterior sides are illustrated in Textfigure 3. As shown in the dental formulae
and the Text-figure 3, it is evident thatthe posterior lateral teeth PII, PIlI andPIV are constant, whereas the pseudocardinal teeth are considerably variable.
The hinge structure has been generally
regarded as one of the most constantcharacters of the Bivalvia and used as abasis of high rank taxonomy. So at the
beginning, I considered the hinge variety
only apparent due to the state of preservation. However, through the careful examination of more than 50 specimens ofwide range in size, I have ascertainedthat the variation must be regarded as
ontogenetic. So far as I have observed,
the specimens smaller than 20 mm inlength all possess only two pseudocardinals
on each valve. On the other hand theTable 3. Variation of hinge structure of Trigonioides (s. s.) paucisulcatus.

PIlI

5 3
4 2

PI!

5 3 (1)4 2

PH

5 3 1a
(l'a)
4 2

PH

(lb)
5 3 1a4 2 l'a
5 3 1a 1b
4 2 l'a (l'b)

PlV
PIlI
PlV
PIlI
PlV
PIlI

PI!

PlV
PIlI

PI!

PlV

(lc)
PIlI
5 3 1a 1b
4 2 l'a l'b
PI!
PlV
5 3 1a 1b 1c(6) 4 2 l'a l'b

PIlI
PH

339

PlV

010
0101 RV
101
1010 LV
010------(1)0101 RV
101-- 0 1010 LV
010-- 0 10101 RV
101--(1)01010 LV
010----(1)010101 RV
101---- 0 101010 LV
010--- 0 1010101 RV
101-(1)0101010 LV
010--(1)01010101 RV
101-- 0 10101010 LV
010----101010101 0 RV
101----010101010 (1) LV

S. Y. YANG

340

RIGHT VALVES
5 3 10 1b 1c Pilia.p. a. p. a. p. a. p. a. p.

KPE 1709~

KPE 1703~

KPE 1715

1

KPE 1704~~

KPE1698~

I

KPE1890i,

KP.E 1700

t

KPE16B2~I,*

I i i:

KPE 16B1~ I ~KPE1853~

KPE1687

KPE1695

KPE1707t~

KPE1689

KPE1690

1
l

!!l

~ i

KPE1691

i

:I

I

a
~Ii t

I!i

a.p. a.p.

l I KPE 1684
~ I KPE1688

;I;

:I

:I

-i=1= 3:

11

KPET702=1~ t KPE 1713

I ~

I_

l j

~i ~

*l

~ t KPE1681~ :t= ~l I

~ ~ :I:

I I
~ i

ji
_t~

::I

em
{

iI

KPE1699
KPE 1701

tj;$

KPE1697
KPE1686
KPE1BB7
KPE16B5

~I ~

~

!i t
'1 I 1:1:*i Il

a.p. a.p. a.p' a.p. a.p.

i i
I ~

KPE 1714

KPE'j879

a. p.

LEFT VALVES
6 4 2 10 1'b PII PIVKPE1694~ f KPE1705

I:

1: a. p.

2: KPE 0000

3:1

4:

I

5:1

6:

t

7:1

8:...L

i

1
-

691.

Ontogenetic variation of Trigonioides paucisulcatusspecimens larger than 35 mm possess fouror five cardinals, and the other ones ofintermediate size three or four. The toothindicated in parentheses is conventionallyimplied incipient presence as a point. As
illustrated in Text-figure 3, this incipient
hinge tooth becomes gradually prominent
with growth.
Having a prejudice that the hinge teeth
are constant, the previous authors paid
little careful attention to the hinge variation. But according to my observation(YANG, 1974), the specific difference must
be present at least between T. (Kumamotoa)
suzukii and T. (Kumamotoa) mifunensis.
Even though the hinge structure has notbeen investigated yet on a viewpoint of
ontogenetic variation among other trigonioidids, this possibility cannot be completely eliminated. The hinge teeth areconsidered not to be always constantamong the samples of T. (Kumamotoa)
matsumotoi, T. (Kumamotoa) mifunensis
and Wakinoa wakinoensis. According to
the descriptions of the previous authors(TAMURA, 1970; OTA, 1975, the dental
formulae are as follows;
TAMURA (1970)
T. (K.) mifunensis(PIlI)
5 3 laIb(6) 4 2 l'a l'b PII
PlVWakinoa
5 3 (1)PIlI
PV(6) 4 2 (1') PIIPlVOTA (1975a, b)
Wakinoa wakinoensis
5 3 1
(PI)
PIlI
(6) 4 2 (1')
PII
(PlV)

341

T. (K.) suzukii
5 3 la(lb)
PIlI(6) 4 2 l'a
PII
PlV
T. (K.) matsumotoi & T. (K.) mifunensis
5 3 laIb
PIlI(6) 4 2 l'a (l'b) PII
PlV
Hoffetrigonia kobayashi
5 3 laIb
4 2 l'a (l'b)

PIlI
PII
PlV
The teeth, 6, 1', Ib and l'b expressed inparentheses may be brought to light as ofontogentic variation, if the characters be
examined on a larger number of large
samples of a wider range in size.
Systematic description

Superfamily Unionacea
Family Trigonioididae Cox, 1952
Genus Trigonioid.es KOBA Y ASH! andSUZUKI, 1936
Trigonioides (s. s.) paucisulcatus SUZUKI,
1940 (emend)
PI. 45, figs. 1-16; PI. 46, figs. 1-26.

1940. Trigonioides kodairai paucisulcatus SuZUKI, Japan.
Jour. Ceol. Ceogr., 17,(3-4), p. 229, pI. 3, figs. 1-4, text·figs.
18-20.
1959. Trigonioides paucisulcatus SUZUKI, OTA,Trans. Proc. Palaeont. Soc. Japan, N. S.,
no. 34, p. 101, fig. 16.
1968. "Trigonioides" paucisulcatus SUZUKI,
KOBAYASHI, Ceol. Palaeont. S. E. Asia,4, p. 128, pI. 23, fig. 4.

Text-figure 3. Crenulation and relative length of the hinge teeth.
2. Specimen number preserved at Kyungpook National
University. 3. Hinge tooth with crenulation on both sides illustrated by short horizontal lines
and of length expressed by vertical heavy line. 4. Posterior lateral tooth with fine crenualtion. 5. Hinge tooth with crenulation on both sides, but the crenulation on anterior side notobserved. 6, Tooth not observed. 7. Hinge tooth of the length not measured exactly, may
be longer than the length illustrated. 8. Incipient presence of subumbonal pseudocardinal tooth.
1. a; anterior side, p; posterior side.

342

s. Y. YANG

Description.-Shell medium in size, commonly about 30-45 mm in length, ratio of
L/H about 1.3, subequilateral and equivalve ; anterior margin well rounded, ventral
one broadly arcuated, posterior margin
rather straight or subtruncated, generallytransversely subovate in outline; moderately inflated; umbo slightly prosogyrous,
placed about two-fifths of the shell lengthfrom the anterior extremity; escutcheon
and lunule narrow. The shells less than
20 mm in length, probably of immature
stage, commonly covered with the characteristic V-sculptures, while those oflater stages over 40 mm in length are ornamented only on the posterior area withrough carinae. On the intermediate ones
the umbonal and the posterior parts bothcovered with the ornaments. The carinae
on the postero-ventral part number commonly four or five. The angles of the
V-ribs about 7° _10° and the radial ribs
and grooves on the umbonal part fairly
fine.
Hinge plate moderate in breadth, provided with the opisthocline pseudocardinalteeth and the posterior lateral teeth; theposterior lateral teeth one on right valve,
two on left valve; the pseudocardinals
two on each valve in immature ones;
forming the dental formula as follows:
5 3
PIlI
4 2 PHPIV
but the number of the pseudocardinals
increases successively with growth to fiveon right valve and four or five on left
valve; forming the dental formula asfollows:
5 3 IaIb
Ic
PIlI(6) 4 2 I'a I'b
PH
PIV
The crenulation of the hinge teeth welldeveloped; it is finer on the posterior
lateral teeth than on the pseudocardinals.
Adductor scars subequal in size and form,
situated close to the outer ventral endsof the pseudocardinal teeth and the pos-terior lateral ones; anterior one subcircularand more strongly impressed, accompaniedwith a distinct minute pedal scar; posterior one also subcircular but not so distinct. Ventral margin crenulated internally; the crenulation stout and strongon the posterior part, becoming weakerand finer towards the anterior and finallycompletely effaced at the anterior end.
Umbonal cavity moderately deep.
Comparison.- The present species is similar to T. (s. s.) kodairai in surface ornamentation, but not identical in its confinement of the ornamentation to theposterior and the umbonal areas and in
the angle of the V-ribs. In this respect,
it is rather nearer to T. (Kumamotoa)
mi/unensis. The radial ribs on the umbo
are finer and the angle of the V-ribs issmaller than in any other trigonioidid
species.
The number of the hinge teeth of thepresent species is quite similar to T.
(Kumamotoa) matsumotoi and T. (K.)
mi/unensis except possessing Ic. However,with respect to other features of the hinge
structure, the present species is similar toT. (s. s.) kodairai rather than to T. (Kumamotoa) spp. The crenulation of thepseudocardinals in the present species are
as fine as in T. (s. s.) kodairai, and not soprominently stout and strong as in T.
(Kumamotoa). The relative length of thepseudocardinals is also rather nearer tothe former. And the hinge plate also isidentical to that of T. (s. s.) kodairai. For
this comparison, the hinge structure ofT. (Kumamotoa) mi/unensis is presented
in Plate 46 (fig. 27).
Phylogenetical remarks.-From the comparison of T. (s. s.) kodairai with T. (s. s.)
paucisulcatus, it can be safely said that atleast three characters are phylogeneticallyvaried. The first is that the surface
ornamentation is gradually effaced withupward stratigraphic sequence. It is in-691. Ontogenetic variation of Trigonioides paucisulcatus
Table 4.
Specimen
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE
KPE

1651 (L)1652 (L)1653 (L)1654 (L)1656 (L)1657 (R)1658 (L)1659 (L)1660 (L)1662 (L)1664 (R)
1666 (R)
1668 (R)
1672 (L)1673 (L)1674 (L)1675 (L)1676 (L)1678 (L)1680 (L)1684 (L)1694 (L)343

Measurements of Trigonioides (s. s.) paucisulcatus
L

H

D

64.5
42.2
42.6
55.2
25.6
55.2
44.6
41. 9
30.9+
42.2
26. 7
36.8
37.8
39.4
37.8
23.2
39.5
31. 8
20.8
38.6
29.9
42.0+

50.9
32.5
32.6
45.1
19.2
43. 7
32. 7
32.3
24.4
30.4+
20.9
29.8
31. 5
29.5
26.9
17.1
28.8
23.8
15.2
28.2
21. 8+
33.4

28.8
16.8
15. 7
19.5
8.8
18.8
17.5
16.2
14.6
16.9
9.3
12.2
13.9
14.3
15. 1
7.9
17.5
11. 1
8. 7
12.6
11. 0
20.2

16. 1
11. 0
9.8
14.2
13.0
9.9
10.0

9.8
9.8

8.5

L/H

D/L

IlL

1. 27
1. 30
1. 31
1. 22
1. 33
1. 26
1. 36
1. 30
1. 27
1. 39
1. 28
1. 23
1. 20
1. 34
1. 41
1. 36
1. 37
1. 34
1. 37
1. 37
1. 37
1. 26

0.45
0.40
0.37
0.35
0.34
0.34
0.39
0.39
0.47
0.40
0.35
0.33
0.37
0.36
0.40
0.34
0.44
0.35
0.42
0.33
0.37
0.48

0.25
0.26
0.23
0.26
0.24
0.24
0.32
0.27
0.26

0.22

L: length, H: height, D: distance between umbo and anterior extremity, I: inflation,(L) : left valve, (R): right valve. Mean of L/H: 1.31; standard deviation of L/H:
0.058, mean of D/L: 0.38; standard deviation of D/L: 0.044. (Based on 22 specimens,
exclusive of the strongly deformed or broken specimens).

teresting to note that a similar feature
was recognized also on the genus Wakinoa,
that is, W. wakinoensis- W. intermediaW. obsoleta (HASE, 1960). The second is
the angle of the V-ribs on the mediansurface becomes also gradually smaller.
It can be more prominently grasped when
the Koreanaia is concerned. From thispoint, the Wakinoa wakinoensis is considered to be put phylogenetically between
Koreanaia and Trigonioides (s. s.). The
third is that the number of pseudocardinalhinge teeth is increased also with upward
stratigraphic sequence.

References cited
BOYD, D. W. and NEWELL, N. D. (1969): Ad·
dendum; Limitations of BERNARD and
MUNIER-CHALMAS system for bivalve
hinge notation. Treatise on invertebrate
paleontology, part N, Mollusca 6, Bivalvia,
vols. 1 and 2. Lawrence, Geol. Soc.
America and Univ. Kansas Press, p. N

908-913.
CIIAl'\G, K. H. (1970): Geology of upper Mesozoic strata, N. Gyeongsang Province,
Southern Korea(I). Jour. Ceol. Soc. Korea,
vol. 6, no. 1, p. 1-12.
Cox, L. R. (1955): On the affinities of theCretaceous Lamellibranch genera Trigonioides and Hoffetrigonia. Ceol. Mag., vol.
92, no. 4, p. 345-349.

344

S. Y. YANG

HASE, A. (1960): The late Mesozoic formations and their molluscan fossils in west
Chugoku and north Kyushu. jour. Sci.,
Hiroshima Univ., Ser. C, vol. 3, no. 2, p.
281-342, pis. 31-39.
HAYAMI, 1. and icHIKAWA T. (1965): Occurrence of Nippononaia ryosekiana from the
Sanchu area, Japan. Trans. Proc. Palaeonto Soc. japan, N.S., no. 60, p. 145-155,
pI. 17.
KOBAYASHI, T. (1956): On the dentition of
Trigonioides and its relation to similarpelecypod genera. japan. jour. Geol.
Geogr., vol. 27, no. 1, p. 79-94, pI. 5.
- - (1968): The Cretaceous non-marine pelecypods from the Nam Phung Dam site in
the northern part of the Khorat Plateau,
Thailand with a note on the Trigonioididae. Geol. Palaeont. S. E. Asia, vol. 4,
p. 109-138, pis. 20-23.
- - and SUZUKI, K. (1936): Nonmarine
shells of the Naktong-Wakino series.
japan. jour. Geol. Geogr., vol. 13, nos.
3-4, p. 243-257, pis. 27-29.

MATSUMOTO, T. (1977): Zonal correlationof the upper Cretaceous in Japan. Palaeonto Soc. japan. Spec. Pap., no. 21, p. 6374.
OTA, Y. (1959a): Trigonioides and its classification (Studies on the molluscan faunaof the Cretaceous Inkstone series, part2). Trans. Proc. Palaeont. Soc. japan,
N.S., no. 34, p. 97-104.
- - (1959b): On the" Nippononaia" from theCretaceous Wakino subgroup, north Kyushu, Japan. Ibid., p. 105-110, pis. 10,11.
- - (1975): Notes on the genus Trigonioides(Bivalvia). (in Japanese). Bull. Fukuoka
Univ. Educ., vol. 24, pt. 3, p. 79-98.
PARK, B. S. and YUN, Y. Y. (1973): Geologicalmap of Korea, Jain sheet. Geol. Surv.
Korea, Seoul.
S01', C. M. et. al (1969):
A study on thesedimentary environments and geologic
structure of Korea. (in Korean). Res.
Rept. Minist. Sci. Tech., 50pp.
SUZUKI, K. (1940): Non-marine molluscan
faunule of the Siragi series in South

Explanation of Plate 45
Figs. 1-16. Trigonioides (s. s.) paucisulcatus (SUZUKI, 1940)
1. Conjoined valves, immature specimen. (KPE 1665) 1a: right valve, 1b: left valve,
showing radial ribs covering almost the whole shell.
2. Left valve (KPE 1656), immature specimen, showing radial ribs on the posterior part.
3. Conjoined valves (KPE 1669), opened ventrally, showing fine radial ribs.
4. Left valve (KPE 1660), showing fine radial ribs on umbo.
5. Left valve (KPE 1676), showing rough radial ribs on posterior half.
6. Right valve (KPE 1664), showing rough radial ribs on postero-ventral area.
7. Right valve (KPE 1661), showing radial ribs on posterior half.
8. Left valve (KPE 1659).
9. Left valve (KPE 1658), showing radial ribs on posterior half.
10. Right valve (KPE 1663), showing radial ribs on posterior half.
11. Right valve (KPE 1667), showing rough radial ribs on postero-ventral area.
12. Conjoined valves (KPE 1655), showing radial ribs on umbo and posterior half. 12a:left side view, 12b: right side view, 12c: dorsal view.
13. Left valve (KPE 1653), showing radial ribs on umbo.
14. Left valve (KPE 1652), showing radial ribs on posterior half. x 0.9.
15. Left valve (KPE 1651), showing rough radial ribs on posterior half. x 0.86.
16. Left valve (KPE 1654), showing rough radial ribs on posterior half.
All figures are of natural size unless otherwise stated. Loc.: at a point 1.5 Km northeast
from Geoncheon-ri, Seo-myeon, Weolseong-gun, Gyeongsangbug-do, Korea. All of the above specimens are kept in the Department of Earth Science, Kyungpook National University, Daegu,
Korea.

YANG,'

Trigonioides paucisulcatus

Plate 45
'I ,

691.

Ontogenetic variation of Trigonioides paucisulcatus

Tyosen. japan. jour. Ceol. Ceogr., vol.
17, nos. 3-4, p. 215-231, pIs. 22-24.
TAMURA, M. (1970): The hinge structure of
Trigonioides, with description of Trigonioides mijunensis, sp. nov., from upper
Cretaceous Mifune group, Kumamoto
Pref., Japan. Mem. Fac. Educ., Kumamoto
Univ., no. 18, sec. 1, p. 38-52, pIs. 1,2.
- - and MATSU1".lURA, M. (1974): On theage of the Mifune group, Central Kyushu,
Japan, with a description of ammonite
from the group by MATSUMOTO, T. Mem.
Fac. Educ. Kumamoto Univ., no. 23, ser.

345

1, p. 47-56, pl. 1.
TATEm'A, I. (1929): Geological atlas of Korea, no. 10, Keishu, Eisen, Taikyu, andiVakwan sheets, Ceol. Surv. Korea, Seoul.
YANG,
S. Y. (1974): Note on the genus
Trigonioides (Bivalvia).
Trans. Proc.
Palaeont. Soc. japan, N. S., no. 95, p. 395408, pIs. 54,55.
- - (1976): On the non-marine molluscan
fauna from the upper Mesozoic Myogog
formation, Korea. Ibid., no. 102, p. 317333, pIs. 33,34.

Ahwa-ri IlPfkll, Baegan-dong s*rlOJ, Banyaweol -¥-1~fI, Chaeyagsan ~~ilI, Chilgog
Daegog-ri *1:l-lI,
Daegu*Ji'II,
Dongmyeong JRilA,
Geoncheon-ri ljil:JII.!1!,
Gyeongju m!1i'I,
Gyeongsang ~1'J;j,Hagbong '~~If,:,
Haman mt.*,
Hasandong ~ilI1JOJ,
Jain ~t:, Jayangsan ~~llJ, Jinju -Wr1+1, Jusasan ~(1i'.PllJ,
Nagdong m-JR, Seo-myeon
ggji'i:j,Sinla ~,\ifL Songnae-dong i".M"J~Il], UIsan ;1{ilI, Waegwan ~~, Weolseong-gun
flilile'fli), Yeongcheon j}cJII, Yeongcheon·gun :7l<JIIn~, Yeonhwa-dong ll:t~1JOJ~1:l-,

Trigonioides (s. s.) paucisulcatus O){lI!If,f;:!ll(;:lH': J: 1.> ~~I': '0~, L :- T. paucisulcatus
G1!r <0) 1l*~~t:R l- L, .:to)I:"JflIHf/f~, ~ji'i:j~tlIijtJ: c'HJi\l"·'f.: *li*~¥IH5-t 1.> 0 T. (s. s.) paucisulcatus 0) pseudocardinal teeth t.::.ltl]jJ GtJ'I':~ l- ~ '~!!O~tJ;it{(1!b I?n
1.>0 .:nlt ontogenetic variation cmfR~n1.>o -ttd?i? pseudocardinals It immature
specimens -Cltjj~J)ll!:c U.: ::::''0-:)'0l-tJ'tJ:~'O)I':, n\i;~ I.:J: "JLiXtj) 1.:1!r< tJ:"JL adultspecimens -CIt 5, 3, la, Ib, Ie, PIII/(6), 4, 2, I'a, I'b, PIl, PlV t.:tJ:1.>o l\'!lifJ'l9ICf(ft1\rili~tJ'Gj£:H:\-t1.> T. (s.s.) kodairai cJ:t~Udt.*, *k!Cjfg{I::.O)~5'\c l-L='0O)tf}~
1.:~{l::.tJ~;t.o .: "J t.:': c tJ; ilA G tJ,I': tJ: "J t.:o
L ~JHiO)V*~~~tJo)V*O)fiJJtlt/J-~ <tJ:1.>.
2. ~JHio)V*~~!llHtt.:A, t':A,r~9<:-t 1.>0
3. pseudocardinal teeth O)1&lt~ <tJ: 1.>00) m;5'\}JBit]!itJ ,346

s. Y. YANG
Explanation of Plate 46

Figs. 1-26. Trigonioides (s. s.) paucisulcatus (SUZUKI, 1940)
1. Right valve (KPE 1703), internal mould showing weak crenulation on pseudocardinalteeth.
2. Left valve (KPE 1683), internal mould showing muscle scars and weak crenulation
on pseudocardinal teeth.
3. Right valve (KPE 1704), 3a: internal mould showing crenulation on pseudocardinalteeth, 3b: rubber cast of 3a, S; crenulated on ventral side only, 3; crenulated on bothsides, la; incipiently presented, PIlI; not crenulated, but lamellar.
4. Left valve (KPE 1688), 4a: internal mould showing crenulation on both pseudocardinaland postero-lateral teeth, 4b: rubber cast of 4a, 4; crenulated on both sides, 2; crenulatedon dorsal side only, PII, PlY; crenulated only one side, dorsal and ventral side, respectively.
S. Left valve (KPE 1701), Sa: internal mould showing crenulation on pseudocardinalteeth, Sb: rubber cast of Sa, 4; crenulated on both sides, 2; crenulated on both sides, situated vestigially between 4 and l'a, l'a; crenulated on dorsal side only. Thiara (Siragimelania) tateiwai associated with immediately below the umbo.
6. Right valve (KPE 1710), internal mould showing crenulated pseudocardinal teeth andmuscle scars on anterior side.
7. Right valve, rubber cast of KPE 1709, showing pseudocardinal teeth, S; not crenulated, 3; crenulated on both sides, la; presented incipiently.
8. Right valve (KPE 17l4) , 8a: internal mould showing crenulated pseudocardinal teeth,
8b: rubber cast of 8a, S; crenulated on ventral side only, 3 and 1a; crenulated on bothsides, 1b; crenulated on antero-dorsal side.
9. Conjoined valves (KPE 1682), internal mould ventrally opened, showing crenulated
pseudocardinal teeth, 9a: right side view, 9b: left side view, 9c: rubber cast of 9a, S;
crenulated on ventral side only, 3; crenulated on both sides, 1a; crenulated on anteriorside only.
10. Right valve, rubber cast of KPE 1698, showing crenulated hinge teeth, S; crenulated
on ventral side only, 3; crenulated on both sides, la, 1b; not preserved, PIlI; crenulatedon both sides.
II. Right valve (KPE 1879), lIa: internal mould, lIb: rubber cast of lIa, S; crenulated
on ventral side only, 3, 1a and 1b; crenulated on both sides, 1a closely associated with 3.
12. Right valve, rubber cast of KPE 1700, showing crenulated hinge teeth, 5; crenulated
on ventral side only, 3; crenulated on both sides, 1a; crenulated on dorsal side, PIlI;
crenulated on both sides.
13. Right valve, rubber cast of KPE 1689, showing crenulated hinge teeth, 5; crenulated
on ventral side only, 3 and 1a; crenulated on both sides, 1b; not observed, PIlI; partly
preserved, crenulated on both sides.
14. Right valve (KPE 1687), 14a: anterior side view, internal mould showing crenulated
pseudocardinal teeth, 14b: right side view, the umbo cut out showing crenulated hinge
teeth, the specimen strongly compressed antero-posteriorly, 14c: rubber cast of 14b,
showing hinge teeth, S; not preserved, 3, 1a and 1b; erenulated on both sides, Ie; crenulated on antero-dorsal side, PIlI; crenulated on both sides.
15. Right valve (KPE 1690), lSa: internal mould, 15b: rubber cast of 15a, showing crenulated hinge teeth, 5; crenulated on ventral side only, 3, 1a and 1b; crenulated on bothsides, Ie; crenulated on anterior side, PIlI; crenulated on both sides.
16. Left valve (KPE 1684), 16a: internal mould showing erenulated hinge teeth, 16b:
rubber cast of 16a, 4; crenulated on both sides, 2; erenulated on dorsal side, l'a; incipient
presence, PIl, PlY; crenulated on upper and lower side, respectively.

YANG:

Trigonioides paucisulcatus

Plate 46

691. Ontogenetic variation of Trigonioides paucisulcatus347

17. Left valve (KPE 1697), 17a: internal mould showing crenulated hinge teeth andventral crenulation, 17b: rubber cast of 17a, 4, 2 and l'a; crenulated on both sides, 2
presented vestigially between 4 and l'a, l'b; incipient presence, PIl, PIV,; crenulated on
upper and lower side, respectively.
18. Left valve, rubber cast of KPE 1713, showing crenulated hinge teeth, 4 and 2; crenulated on both sides, l'a; incipient presence, PIl, PIV; weakly crenulated on upper andlower side, respectively.
19. Left valve, rubber cast of KPE 1702, showing crenulated hinge teeth, 4 and 2; crenulated on both sides, l'a; crenulated on anterior side, l'b; not observed, PIl and PIV;
crenulated on upper and lower side, respectively.
20. Left valve (KPE 1887), 20a: internal mould showing crenulated hinge teeth, 20b:
rubber cast of 20a, 4 and 2; crenulated on both sides, l'a; incipient presence, PIl, andPIV; crenulated on upper and lower side, respectively.
21. Left valve (KPE 1685), internal mould showing crenulated hinge teeth.
22. Conjoined valves (KPE 1681), internal mould showing crenulated hinge teeth.
23. Left valve (KPE 1686), 23a: internal mould showing crenulated hinge teeth andventral crenulation, 23b: rubber cast of 23a, 4; crenulated on both sides, 2; crenulated
on antero-dorsal side, l'a; incipient presence, PIl and PIV; crenulated on upper andlower side, respectively.
24. Right valve, rubber cast of KPE 1695, showing crenulated hinge teeth and posteroventral crenulation, 5, 3 and 1a; crenulated on both sides, 1b; not observed, PIlI; crenulated on both sides.
25. Conjoined valves (KPE 1706), opened ventrally about at right angle, internal mould,
showing crenulated hinge teeth and the muscle scars.
26. Right valve (KPE 1696), internal mould, compressed antero-posteriorly, showing ventral crenulation.
Loc.: at a point 1.5 Km northeast from Geoncheon-ri, Seo-myeon, Weolseong-gun, Korea.
Figure 27_ Trigonioides (Kumamotoa) mifunensis TAMuRA, 1970.
Rubber cast of KPE 1023, showing strong and stout pseudocardinal hinge teeth.
Loc.: See Tamura, 1970.
All figures are of natural size.
The specimens illustrated above are kept in the Department of Earth Science, Kyungpook
National University, Daegu, Korea.

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 111, pp. 348-357, pis. 47, 48, Sept. 30, 1978

692. ONTOGENIES OF FOUR UPPER CAMBRIAN TRILOBITES FROMTHE BONNETERRE DOLOMITE, MISSOURI*CHUNG-HUNG HU
Department of Physics, Taiwan Normal University, Taiwan
Abstract. As one of the series of papers dealing with the ontogenic changesof various Trilobita the present study clarified the ontogenic development of four
Upper Cambrian species from the Bonneterre Dolomite in Missouri, i. e., Coosella
prolifera, Holcacephalus tenurus, Welleraspis missouriensis and Meteoraspis globosa.
Among these four species, Coos ella prolifera was investigated in great detail todistinguish the metaprotaspid, paraprotaspid, early meraspid and late meraspid
stages. Moreover, two different morphologic groups were clearly shown in the
Coosella prolifera and Holcacephalus tenurus samples respectively throughout the
ontogenic changes. These bimodal phenomena were supposed to implicate a sexual
dimorphism of the trilobites.

frontal furrow, and narrower preglabellar
field, and the pygidium has a narrower
marginal border; the second group has abroader glabella, indistinct frontal furrow,broader preglabellar field, and the pygidiumThe
has a broader marginal border.
species Holcaceplwlus tenllrus (WALCOTT)
shows also two morphologic groups: thefirst has a narrow preglabellar field anda broader glabella; and the second isbroader. and the glabella is slender. Thesebimodal phenomena are postulated as
sexual dimorphism within each species
population.
The morphogenesis of Coosella prolifica
and Meteoraspis globosa are closely similar
to those of Syspacheilus dunoirensis(MILLER), Coosella convexa TASCH, WilsOl1ella pel1nsylvanica Hu, Crepiceplwl1ls
deadwoodensis Hu, and Coosia albertel1sis
RESSER. The ontogenic development of
Welleraspis missouriel1sis resembles W.
loc/zmal1ae Hu, and Pemphigaspis bullata
HALL; and the instars of Holcacephalus{enurus are identical with those of material
found from South Boulder Creek, Madison

Introduction
The present report describes the ontogenic development of Coosella prolifica
LacHMAN, Welleraspis missouriensis (LaCHMAN), Holcacephalus tenurlls (WALCOTT)
and Meteroraspis globosa (MILLER). The
studied material was collected by Dr. Ch.
LacHMAN-BALK, New Mexico Institute of
Mining and Technology, from the Bonneterre Dolomite, Upper Cambrian, Cedaria
Zone, Saline Creek, 1.2 miles southeast of
the School House at Avon, Ste GenevieveCo., Missouri, in 1940. Among the described species, Coosella prolifica is represented by four ontogenic stages: i. e.,
the metaprotaspid, paraprotaspid, earlymeraspid, and late meraspid stages, andthose of the others are rather poorly
known. They are either lack the earlier
stages or show discontinuity of the growthsequence. Coosella prolifica shows two
different morphologic groups: the first is
characterized by longer glabella, deep

* Received Nov. 18, 1977; read June 3,1978
at Tsukuba.

348

692. Ontogenies of Cambrian t1"ilobites
Co., Montana by Hu (1971). These ontogenic similarities within each groupcertainly indicate close phylogenic relationships.
Acknowledgements.- The author wishes
to express his thanks to Dr. Ch. LaCHMANBALK, New Mexico Institute of Mining
and Technology, New Mexico for permission to describe her collections. Thanks
are also due to Dr. Kenneth E. CASTER,
University of Cincinnati, Ohio for readingover the present manuscript. The figured
specimens are all stored in the Geology
Museum, University of Cincinnati (UCGM),
Ohio.
Systematic Pa!eontology

Family Coosellaidae PALMER, 1954
Genus Coosella LacHMAN, 1934
Coosella prolifica LacHMAN
PI. 47, figs. 1-34
Coosella prolifica LOCHMAN, 1936, p. 39, pI. 9,
figs. 4-11 ; Lachman & Duncan, 1944, p. 83,
pI. 12, figs. 35, 36.
Syspacheilus camurus LOCHMAN, 1940, p. 42,
pI. 3, figs. 21-25.

Remarks.- The present species is represented by two different morphologic
groups: the first group has the cranidiumwith longer glabella (figs. 18,22,24), deep
frontal furrow, and narrower preglabellar
field, and the pygidium has a narrower
marginal border (figs. 29, 31, 34); and theother one has a cranidium with broader
glabella, indistinct frontal furrow, andbroader preglabellar field (figs. 20, 21, 23),and the pygidium has broader marginalborder (figs. 30, 32, 33). The author is of
the opinion that these differences areminor and do not pertain to generic or
specific level differences (as some think),
since, both forms have a similar shaped

349

cranidium, glabella and pygidium, andsimilar depth of the dorsal furrow. They
are probably instead represent sexual
dimorphism within the same species population. The first group would be "female"and the second "male" forms. If thisinterpretation is tenable, then "Coosella
prolifica" of LacHMAN must be viewed as
the female form, of LacHMAN'S (1940)"Syspacheilus camurus" which is based onmale specimens of the same species.
Coosella prolifica LacHMAN, ontogeny
Metaprotaspid stage (PI. 47, figs. 1, 2).The shield is rounded in outline, convex,
and is about 0.30-0AOmm in sagitallength ;the axial and pleural lobes are not distinctly divided, except for a longitudinalfurrow; a few glabellar lobes are rather
faintly marked along the central axis;
the frontal lobe is convex, and demarked
with a pair of distinct pits laterally; thesuperciloid ridges are extended from theanterior lateral lobe, and curving posterolaterally for a short distance; the small
occipital node is faintly elevated at theposterior shield margin; the pleural lobe
is broader than the axis, and slopes downward around the margin. The skeletal
surface is faintly covered by granules.
Parapl'otaspid stage (PI. 47, figs. 3-·6).The small shield is subrounded, convex,
about 0.36-0.50 mm in length (sag.); theaxial lobe is faintly delimited by dorsal
furrow, slenderly cylindrical; no distinct
glabellar segments are recognizable, except
for a few faintly depressed impressions
marked along the dorsal furrows, whichsuggest the presence of the axial segments; the frontal lobe is not distinctly
elevated, and except for a pair of pits, isunmarked at the anterolateral margin;
the paired superciloid ridges are short,and connected by a pair of elevated andwell delimited palpebral lobes ; the occipital

350

Chung-Hung Huring is transversely oval, highly convex,
and well separated by a furrow; thefixigena is convex along the outer margin,
and is wider than the axis; the posterior
fixigenal border is about the same width
as the occipital ring; the narrow lenticular
protopygidium is divided by one to threesegments, it slopes downward steeply; theskeletal surface is faintly granulated.
The morphogenesis of the skeleton during this stage saw the glabella developed
from a cylindrical to slightly forwardly
expanded; the frontal pit becomes shallower; the posterior fixigena increases inwidth; and the protopygidium increases
in number of segments and from lenticular
to semicircular in outline.
Early meraspid stage (PI. 47, figs. 719).-The cranidium is trapezoidal in out'line, convex, about 0.60-1.0 mm in length(sag.); the slender cylindrical glabella isdeeply defined by a dorsal furrow, andwithout distinct glabellar segments; theoccipital ring is convex, bearing a minutemedian tubercle, and deeply separated byan occipital furrow; the narrow anteriorborder arches forwardly, and is welldelimited by a frontal furrow; the fixigena
is convex, broadly triangular, with the narrow medium-sized palpebral ridge locatedin front of the mid-line (tr.) of the glabella; the anterior facial suture is shortand convex, and the posterior one is divergently convex; the posterior fixigenal
border is convex, elevated from the broadand deep border furrow, and is about thesame width as the occipital ring.
The morphogenesis of the instars during this stage saw the slightly forwardly
expanded glabella become cylindrical, theanterior border increased in width; thepalpebral lobe moved backward from theanterior border; the fixigena reduced thewidth from wide to narrower than theglabella. The surface of the skeleton iscovered by faint granules and a few

coarse ones.

Late meraspid stage (PI. 47, figs. 1114).-The cranidium is trapezoidal in outline, convex, about 1.0-1.8 mm in segittal
length; the glabella is subquadrate, slightly
tapering anteriorly and with a round
anterior margin; the glabella furrows are
not recognizable, except for the very faintoccipital pair; the dorsal furrow is narrow and deep; the convex occipital ringis crescentic, bearing a medium sized
occipital tubercle; the narrow preglabellar
field is present, gently convex, and welldefined by a broad anterior furrow; theanterior border is broad, convex, andarches forwardly; the convex fixigena is
about one-third the width of glabella, has
the medium-sized palpebral lobe situatedon the mid-length of the glabella; theposterior fixigenal border is narrower thanthe occipital ring (tr.), and is deeply differentiated by a border furrow. The
anterior facial suture line is short, slightly
divergent-convex, the posterior one isdivergent laterally and convex. The
surface of· the skeleton is covered by bothfaint and coarse granules; irregular ridges
are elevated along the anterior border.
The skeletal morphogenesis during thisstage sees the preglabellar field increases
in width; the glabella becomes subquadrateand slightly deepens forwardly; the posterior fixigena grows narrower as thepalpebral lobe moves backward from the
mid-line of the shield (tr.).
The smallest pygidium is about 0.5 mm
in length (sag.), it is convex and consistsof 6-7 segments; the anterior thoracic
segments are ankylosed with the terminal
plate-the rudmentary pygidium- andshow free pleural spines; the axial lobe isconical, tapering posteriorly, and reaches
less than the full length of the shield.
The surface is minutely granulated, and a
few coarse nodes are present along thepleural bands.

692.

Ontogenies of Cambrian tr'ilobites

351

The morphogenesis of the pygidium
during these growth stages are: the rudimentary pygidium or the ankylosed posterior plate is changed from a rounded
posterior margin to an inwardly directed
bend; the surface granules on the pygidial
surface are reduced.
Remarks.-The earlier ontogenic developmental stages of the present species
are closely similar to those of Syspacheilus
dunoirensis (MiLLER) (Hu, 1972), Coosella
convexa TASCH, Wilsonella pennsylvanica
Hu (Hu, 1968), Crepicephalus deadwoodensis
Hu (1972), Coosia albertensis RESSER(Hu, 1975), and Meteoraspis globosa (MILLER) (LaCHMAN & Hu, 1961, and presenttext). Thus, these genera possibly haveclose phylogenetic relationship.
Figured specimens. - UCGM. 43267,
43267a-43267z, 43267a' -g'.

unknown in the present specimens.
Two incompletely preserved specimens,
i. e., a cranidium and a pygidium are
figured by LacHMAN (1940) and referred
to Prochuangia? berryi LacHMAN. The
cranidium has a subquadrate glabella,convex but faint glabellar furrows, and anarrow fixigena. It is possibly a memberof the genus Nisonella or Genel'ievella;
however, the specimen shows an incomplete postfixigena. The associated pygidium is semicircular in outline, withnumerous marginal spines. These arecertainly not the generic features of thegenus Prochuangia, because Prochuangia
has a pygidium with a pair of large caudal
spines and a transverse subquadrate outline (KaBA YSHI, 1935). This pygidium
belongs more likely to Meteoraspis, i. e.,
M. globosa (MILLER).

Family Tricrepicephalidae PALMER, 1954

Meteoraspis globosa (MILLER), ontogeny

Genus Meteoraspis RESSER, 1935

The ontogeny of the present species isincompletely known. The smallest instars
are assigned to the early meraspid stage
and are succeeded by late meraspid andholaspid stages, thus completing the postparaprotaspid growth sequence. These
are described as follows.
Early meraspid stage (PI. 48, figs. 23,
24).-The cranidium is about 0.50-0.75 mm
in sagittal length, convex, trapezoidal totruncatotriangular in outline, with narrow
convex anterior border which arches
slightly forward; the anterior border iswell defined by a frontal furrow, and has
a median niche toward the anterior gla·
bellar margin; the glabella is cylindrical,
convex, without distinct glabellar segments or furrows; the fixigena is convex,
and is of about the same width as theglabella; the medium-sized palpebral lobe
is a narrow, elevated ridge, well definedby a distinct palpebral furrow; the palpebral lobe is located in front of the mid-

Meteoraspis globosa (MILLER)
PI. 48, figs. 23-36
Blountia globosa B. B. MILLER, 1936, p. 25-27,
pI. 6, figs. 22-24.
Meteoraspis globosa (MILLER) in Lacm.IA:VI &Hu, 1961, p. 141, pI. 28, figs. 7-19.
Prochuangia? berryi LacHMAN, 1940, p. 39, pI.
4, fig. 17 only (not 18-20).

Remarks.- The present species is represented by more than 30 small andlarge sized cranidia. The smallest cranidium, assigned as early meraspis, is about
0.65 mm in sagittal length, and the largest
adult cranidium measures 6.0 mm long.
They show a continuous growth sequence.
The morphogenesis and the adult morphologic characteristics are identical withmaterial found in the Wind River Mountains, Wyoming, reported by LacHMAN &Hu (1961), except that the librigena is352

Chung-Hung Hutransverse line of the cranidium; the wellelevated palpebral ridge arches anterolaterally from the anterolateral margin of
the glabella and terminates in front of
the palpebral lobe; the occipital ring isconvex, lenticular, and bears a mediantubercle; the posterior fixigenal border iselevated, and deeply separated by a borderfurrow, and is broader than the occipitalring; the dorsal surface of the cranidium
is deeply demarked by an axial furrow;a few coarse granules are sparsely scattered among the finer ones.
The earliest instar of the present stage
is known by LOCHMAN & Hu (1961, pI.
28, fig. 12) from the Wind River Mtns.,
Wyoming. It is about 0.5 mm in sagittal
length, and shows a well defined dorsal
furrow, and a cylindrical glabella, without
a glabellar segmental furrow. This 1961
material possibly represents one stepearlier than the material here illustrated(pI. 48, fig. 23).
Late meraspid stage (PI. 48, figs. 2533).-The cranidium measures 0.70-1.0mm
in sagittal length; it is triangular in outline, convex, with an arched anteriorborder, and deep dorsal furrows; theanterior border is broad, convex, and distinctly separated by a frontal furrow; anarrow preglabellar field is present. It isnarrow in the smaller instars but becomes
broader and about the same width as
frontal border in the larger ones; theglabella is cylindrical to slightly tapering
forward from the deeply marked occipital
furrow; no distinct glabellar segments areseen; the convex occipital ring is lenticular, arches posterioriy, and bears aminute median node. The fixigena isslightly narrower than the glabella, convex,
and has the medium-sized palpebral lobe
located on the mid-transverse line of the
cranidium; the posterior fixigena is about
equal in width to the occipital ring, anddeeply marked by a broader furrow; theposterior fixigenal border is convex; theanterior facial suture line is convergently
convex, and the posterior one is divergent
laterally. The skeletal surface is marked
by coarse and medium-sized granules.
The morphogenesis of the cranidium
during the present stage sees the preglabellar field change from narrow to
broader, the glabella from cylindrical toconical, the fixigena from broad to narrower, and the palpebral lobe moves
backward from near the anterior borderto the mid-transverse line of the cranidium.
The present stage is correlated with what
was reported as a "series of small holaspid
cranidia" by LOCHMAN & Hu (1961, pI. 28,
figs. 13-17) from the Wind River Mtns.,
Wyoming.
When the cranidium attained the holaspid
stage (pI. 2, figs. 34-36) the glabella becomes oval, the fixigena narrower, twodistinct pits are impressed in the frontal
furrow, and the skeletal surface has fewer
but coarse granules.
Remarks.- The ontogenic developmentof the present species is rather similar tothat of Coosella prolifica LOCHMAN, exceptthat in M eteoraspis globosa (MILLER) theskeletal surface is coarsely granular.
They are possibly closely related.
Figured specimens. - UCGM. 43270,
43270a -43270m.
Family Lonchocephalidae HUPF:, 1953
Genus Welleraspis KOBA YASH!, 1935
Welleraspis missouriensis (LOCHMAN)
PI. 48, figs. 1-11
Avonaspis missouriensis LaCHMAN, 1940, p. 41,
pI. 3, figs. 10-13.

Diagnosis. - Cranidium trapezoidal in
outline, strongly convex; glabella subquadrate to oval, marked by pair of

692. Ontogenies oj' Cambrian trilobites
traceable glabellar furrows; occipital ring
bearing a stout median spine and a minutemedian tubercle; preglabellar field very
narrow; anterior border narrow, convex,
distinctly separated by frontal furrow;fixigena triangular, bearing small palpberallobe. Librigena elongate with mediumsized genal spine. Pygidium roundly subtriangular in outline, convex, conical, axis
divided by shallow furrows into four ormore rings and a terminal portion; narrow marginal border horizontally convex.
Skeletal surface covered by medium-sized
granules.
Description. - The cranidium is trapezoidal in outline, strongly convex; theglabella is broadly subquadrate to oval,
highly convex, above the fixigena, andhas two pairs of traceable glabellar furrows; the glabellar furrows are faintly
impressed laterally to the dorsal furrows;
the dorsal furrow is V-shaped, deeply surrounding the glabellar margin; the occipital ring is convex, triangular, narrow
laterally and broadening to the central
portion and then drawn out into a stout
occipital spine; a transverse, elongate,
minute tubercle is born on the center ofthe occipital ring; the preglabellar fieldis very narrow, convex, and slightly depressed in front of the glabella; theanterior border is narrowly crescentic,
arching forward, and is distinctly delimitedby the frontal furrow. The fixigena isnarrowly triangular, convex along thefree margin, and about one-fourth thewidth of the glabella between the palpebral
lobe and the dorsal furrows; the medium
to small-sized palpebral lobe is distinctly
defined by a palpebral furrow, elevated,
and located in front of the mid-line (tr.)
of the glabella; the elevated palpebral
ring is oblique toward the anterior lateral
margin of the glabella; the anterior branchof the facial suture is convergently convex,
and the posterior one is divergent pos-353

terolaterally and convex; the posterior
fixigenal border is narrower than the occipital ring (tr.) and deeply impressed by
the border furrow.
The librigena is narrowly elongate andthe ocular platform is about one and half
the width of the lateral border, convex,
and well defined by a broad lateral furtow; the ocular ring is medium-sized; thegenal spine is medium-short, and has abroad base; no posterior border in known.
The pygidium is subtriangular in outline, convex; the axial lobe is conical,
convex above the pleural lobe, tapers
posteriorly, and is divided into five or
more convex rings by furrows; theterminal portion extends to the posterior
end of the pygidial margin; the pleurallobe is convex below the axis, about thesame width as the axis, and divided into
four deep furrows and faint interpleuralgrooves; the marginal border is slightly
broader than the anterolateral marginthe facet region, and narrows posteriorly
to the posterior terminal portion, and bears
a few pairs of serrate nodes.
The skeletal surface is covered bymedium-sized granules, and parallel ridges
along the cephalic border, i. e., anteriorborder of the cranidium and lateral borderof the librigena.
Remarks.-The present species differsfrom the type species Welleraspis jerseyensis (WELLER, 1900), W. swartzi (TASCH,
1951) (RASETTI, 1954; Hu, 1968), W.
lochmanae Hu, 1969, W. lata HOWELL,
1957 (Hu, 1964), by the strongly convex,
broader glabella, narrower preglabellar
field, and round and less numerous pygidial
segments.
Welleraspis missouriensis (LOCHMAN),
ontogeny

The ontogenic development of the present species is comparable to that of W.

354

Chung-H'wng Huswartzi (TAscH) (Hu, 1968; RASETTI, 1954),
W. lata HOWELL (Hu, 1964), and W.
lochmanae Hu, 1969 although the ontogenyof W. missouriensis is incompletely known.
Two growth stages are recognizable in
the ontogenic development, i. e., the earlymeraspid and the late meraspid stages.
Early meraspid stage (PI. 48, figs. 7, 8).The cranidium is trapezoidal to roundedin outline, convex, about 0.45-0.65 mm inlength (sag.); the glabella is expanded
forward from the occipital furrow, convex
above the fixigena, and marked by pairsof poorly separated glabellar segments;
the triangular occipital ring is convex,
and bears a medium-sized occipital spine;
the anterolateral glabellar pits are welldefined; a pair of short superciloid ridges
extends from the anterolateral glabellarpits, and ends in front of the elevatedpalpebral lobe. The triangular fixigena isconvex, broad, and has the narrow andwell defined palpebral ridge located at thefirst glabellar furrow (tr.); the rear border
is elevated, distinctly demarked by aborder furrow, and is about one and ahalf the width of the occipital ring (tr.).
The skeletal surface is covered by mediumsized granules and scattered with sparse
nodes.
Late meraspid stage (PI. 48, fig. 6).The cranidium is trapezoidal in outline,
highly convex, and is about 1.0-1.5 mm inlength (sag.); the subquadrate glabella isconvex, with both anterior and posterior
margins rounded; the dorsal furrow is
deep and v-shaped; three pairs of glabellar
furrows are faintly impressed laterally tothe dorsal furrows; the crescentic convex
occipital ring is well defined by a deep

Explanation of Plate 47
Figures 1-34. Coosela prolifica LOCHMAl\"
1, 2, metaprotaspides, showing the indistinctly developed axial and pleural lobes. 1, x50,
UCGM. 43267; 2, x 43, UCGM. 43267a.
3-6, a few paraprotaspides, showing the well differentiated axial and pleural lobes andthe presence of the protopygidium; 3, x 39, UCGM. 43267b; 4, x 42, UCGM. 43267c; 5,
x 42, UCGM. 43267d; 6, x 30, UCGM. 43267e.
7-10, a few early meraspides, showing the presence of the anterior cranidial border; 7,x17, UCGM. 43267f; 8, x19, UCGM. 43267g; 9, x21, UCGM. 43267h; 10, x17, UCGM.
43267i.
11-14, a few late meraspides, showing the presence of the preglabellar field and the subquadrate glabella; 11, x 13, UCGM. 43267j; 12, x 12, UCGM. 43267k; 13, x 10, UCGM.
432671; 14, x 14, UCGM. 43267m.
16, 25, two nearly complete Iibrigenae; 16, x 9, UCGM. 43267w; 25, x 10, UCGM. 43267x.
15, 19, 20, 21, 23, a few "male" cranidia, showing the short broad glabella and the wider
preglabellar field; 15, x 8.6, UCGM. 43267n; 19, x 5.4, UCGM. 43267q; 20, x 6, UCGM.
43267r; 21, x7, UCGM. 43267s; 21, x7, UCGM. 43267t; 23, xlO, UCGM. 43267u.
17, 18, 22, 24, a few "female" cranidia, showing the narrow and longer glabella, narrower
preglabellar field and the distinct frontal furrow; 17, x 6, UCGM. 432670; 18, x 11, UCGM.
43267p; 22, x4.7, UCGM 43267t; 24, x3.4, UCGM. 43267v.
26-28, three immature pygidia, notice the thoracic segments and the well ankylosed
pygidial plate; 26, x17, UCGM. 43267y; 27, x10, UCGM. 43267z; 28, x 11.5, UCGM.
43267a'. 30, 32, 33, three "male" pygidia, showing the round pygidial margin; 30, x 2.3,
UCGM. 43267c'; 32, x3.2, UCGM. 43267e'; 33, x5.6, UCGM. 43267f'.
29, 31, 34, three "female" pygidia, showing the narrower pygidial margin; 29, x5, UCGM.
43267b'; 31, x4, UCGM. 43267d'; 34, x5, UCGM. 43267g'.

Hu:

Trilobite ontogenies

2

Plate 47

3

11

4

10

14

6

8

9

15

16

5

7

13

12

21

22

24

23

31

30

692.

Ontogenies of Cambrian trilobites

occipital furrow; the occipital furrow isnarrow across the sagittal line, and wider
laterally to show a transverse W-shape;
the broadly based, short occipital spine isextended posteriorly; the narrow, convex
anterior border arches anteriorly; thetriangular fixigena is convex and slopes
downward along the lateral margin, andhas the small-sized, well elevated palpebral
lobe located in front of the glabella,. between the anterior first and second glabellar furrows; the elevated palpebral
ridge extends continuously from the palpebral lobe and ends at the anterior firstglabellar furrow (tr.); the posterior fix igena is about the same width or slightly
narrower than the occipital ring (tr.); theanterior facial suture line is convergently
convex, arid the posterior one is divergent
and posterolaterally convex. The surfaceof the carapace is covered by mediumsized granules and bears scarce coarse
ones.
The meraspid pygidium (PI. 48, figs. 9,10).-Two well preserved instar pygidia
are assigned to the meraspid stage without any definitive differences separatingthem. They are about 0.45-0.55 mm insagittal length, convex, and have 4-7
segments; the axis is conical, tapers possteriorly, and is divided by segmental
furrows; the pleural lobe is either thesame width as the axis or slightly broader;each of the pleural bands ends in a pairof short spines; the pleural spines areturned upward to reveal serrate nodes.
Figured specimens. - UCGM. 43268,
43268a -43268 j.
Family Norwoodiidae WALCOTT, 1916
Genus Holcacephalus RESSER, 1938
Holcacephalus tenu1'Us (WALCOTT)
PI. 48, figs. 12-22

355

NorllJoodia tenurus WALCOTT, 1Y16, p. 172,
pI. 28, figs. 2-2 g.
NorllJoodina tenurus (WALCOTT) in LOCHMAN& DUNCAN, 1944, p. 137, pI. 13, figs. 15-17.
Holcacephalus tenurus (WALCOTT), LOCHMAN& Hu, 1960, p. 823, pI. 98, figs. 1, 2,4-7,
14-23, 28-32, 40-44, 52 only; Hu, 1971, p.
176, pI. 3, figs. 1-32, and text. fig. 4a-k.
Holcacephalus ct. tenurus (WALCOTT), PALMER, 1954, p. 742, pI. 84, fig. 3.

Remarks.- The present species is represented by a few adult skeletons andseveral earlier instars. The morphogenesisof these carapaces is closely comparable
to that of a collection from the Pilgrim
Formation, Upper Cambrian, Montana,reported by Hu (1971). The adult cranidia
are separated into two different morphologic groups: one has the cranidium withshort, broad and cylindrical glabella, andnarrow preglabellar field, while the otherhas an elongate, narrow, conical glabella.
and broader preglabellar field. The generalmorphogenesis of the early instars shows
the axial and the pleural lobes to beindistinct, but well separated laterally;
the glabella expands forwardly, and becomes cylindrical to conical later; thepreglabellar field changes from narrow to
broad, the palpebral lobe moves from theanterior shield margin to the midline of
the cranidium, and the posterior fixigena
increases in width; the freely articulated
pygidial segments are ankylosed.
Figured specimens. - UCGM. 43269,
43269a-43269i.

References
Hu, C. H. (1964): The ontogeny and dimorphism of Welleraspis lata HOWELL (Trilobite). Jour. Paleont., vol. 38, no. 1, p.
95·97, 1pl.
- - (1968): Notes on the ontogeny and sexual dimorphism of Upper Cambrian trilobites of the Welleraspis faunule Pennsyl·
vania. Nanyang University Jour., vol. 2,

356

Ch'Lmg-Hung Hu

p. 321-357, 5pls.
- - (1969): Ontogeny and sexual dimorphismof three Upper Cambrian trilobites. Nan.
yang University jour., vol. 3, p. 438-463,
3pls.
- - (1971): Ontogeny and sexual dimorphismof Lower Paleozoic Trilobita. Palaeollto.
graphica, Americana, vol. 7, no. 44, p. 31155, 25pls.
- - (1972): Ontogeny of three Cedaria zone
trilobites from Upper Cambrian, Montana.
Trans. Proc. Paleont, Soc. japan, N. S.,
no. 85, p. 245-259, pis. 29, 30.
- - (1975): Ontogenies of three Late Cam·brian trilobites from the Deadwood For·mation, northern Black Hills, South Da·
ktda. Bull. America Paleont., vol. 67, no.

287, p. 251-273, 3pls.
Hu, C. H., and Li 1. L. (1971): Ontogeny offour proparian trilobites from Upper Cam·brian, Montana, U. S. A. (With a descrip.
tion of Vernaculina lush/hi, n. sp.). Proc.
Ceol. Soc. Chilla, no. 14, p. 165-185, 4pls.
5figs.
KOBAYASHI, T. (1935): The Cambro·Ordo·
vician formation and faunas of South
Chosen. Univ. Tokyo, Fac. Sci. jour., sec.
2, vol. 4, pt. 2, p. 49-344, 26pls.
LacHMAN, C. (1936): New trilobites fromthe Bonneterre dolomite (Upper Cambrian)of Missouri. jour. Paleont., vol. 10, no.
1, p. 35-43, 1pl.
- - (1940): Faunas of the Basal Bonneterre
dolomite (Upper Cambrian) of south·

Explanation of Plate 48
Figures l-ll. Welleraspis missouriensis (LacHMAN)
1-4, few cranidia, showing the strongly convex glabella; 1, x 8.6, UCGl\,1. 43268; 2, x 10,
UCGM. 43268a; 3, x8.8, UCGM. 43268b; 4, x9, UCGM. 43268c.
5, a nearly complete librigena; x 7, UCGM. 43268d.
6, a late meraspid cranidium; notice the quadrate glabella; x 17, UCGM. 43268e.
7, 8, two meraspid cranidia; notice the forwardly expanded glabella; 7, x 22, UCGM.
43268f; 8, x 21, UCGM. 43268g.
9, 10, two meraspid pygidia; 9, x 24, UCGM. 43268h; 10, x 12, UCGM. 43268i.
11, a small nearly complete holaspid pygidium; x 16, UCGM. 43268j.
Figures 12-22. Holcacephalus tenurus (WALCOTT)
12, a complete anaprotaspis; notice the faint furrow along the central axis; x 37, UCGM.
43269.
13, 14, two metaprotaspides, showing the well differentiated axial and pleural lobes; 13,
x 32, UCGM. 43269a; 14, x 31, UCGM. 43269b.
15-17, three paraprotaspides, showing the presence of the protopygidium; 15, x33, UCGM.
43269c; 16, x33, UCGM. 43269d; 17, x32, UCGM. 4326ge.
19, a broken cranidium, showing the quadrate glabella; x 7, UCGM. 43269f.
18, 20, 21, three nearly complete cranidia, showing the narrower and elongate glabella;
18, x 2.2, UCGM. 43269g; 20, x 8, UCGM. 43269h; 21, x 8.5, UCGM. 43269i.
22, a complete librigena; x 12.5, UCGM. 43269j.
Figures 23-36. Meteoraspis globosa (MILLER)
23, 24, two early meraspid cranidia; notice the presence of the anterior border; 23, x 20,
UCGM. 43270; 24, x 20, UCGM. 43270a.
25-33, a growth series of late meraspid cranidia, showing the presence of the preglabel·lar field and the broaden of the glabella; 25, x17.5, UCGM. 43270b; 26, x17, UCGM.
43270c; 27, x 16, UCGM. 43270d; 28, x 15, UCGM. 43270e; 29, x 14, UCGM. 43270f; 30, x 15,
UCGM. 43270g; 31, x 15, UCGM. 43270h; 32, x 12, UCGM. 43270i; 33, x 10, UCGM. 43270j.
34-36, three holaspides, showing the oval, strongly convex glabella, and two pits infrontal furrow; 34, x5.8, UCGM. 43270k; 35, x2.1, UCGM. 432701; 36, x4.4, UCGM.
43270m.

Hu:

Plate 48

Trilobite ontogenies

4

3

2

9

5

13

7

12

6

11

10

8

17

16

15

24

25

26

28

27

14
18

19

23

22

21

29
31

32

33

34

35

36

692. Ontogenies of Cambrian trilobites
eastern Missouri. jour. Paleollt., vol. 14,
no. 1, p. 1-53, 5pls.
LacH~IA;'!, C. and DUNCAN, D. (1944): Early
Upper Cambrian faunas of central Montana. Ceol. Soc. America, Sp. Paper., no.
54, 181pp, 19p1s.
LacHMAN, C. and Hu, C. H. (1960): Upper
Cambrian faunas from northwest Wind
River Mountains, Wyoming (Part I).
jour. Paleont., vol. 34, no. 5, p. 793-834,
pis. 95-100.
- - and - - (1961): Upper Cambrian faunas
from the northwest Wind River Mountains, Wyoming (Part II). jour. Paleont.,357

vol. 35, no. 1, p. 125-146, pis. 26-30.
- - and - - (1962): Upper Cambrian faunas
from the northwest Wind River Mountains, Wyoming (Part III). jour. Paleont.,
vol. 36, no. 1, p. 1-28, 7pls.
PALMER, A. R. (1954): The faunas of the
Riley Formation in central Texas. jour.
Paleont., vol. 25, no. 6, p. 710-786, pis.
76-92, 3 tables.
RASETTI, F. (1954): Phylogeny of the Cambrian trilobites family Catillocephalidae
and the ontogeny of Welleraspis. jour.
Paleont., vol. 25, p. 559-612, pI. 65, 4 textfigs.

*ilfnix""C"lt. Coosella prolifera, Holcacephalus tenurus, Welleraspis missouriensis:}.; J: (J Meteoraspis globosa ()) 7J '/ 7' Y 7*c~Wl'::::~!H. 4 fifi(})Mt4>5ej:.~ c I) '&.;-?tJ,·->f;:.oJJre9=' Coosella prolifera (})5ej:.'I~ff:'il~I:f7'GjtlcfJJ3l?tJ'IC -t J.> ;:. c tJ,-C' ~ t.::.tJ', {t!!.(}).::::fifilc-?\" -C liT71tJ:~t/-li:f4l?.ntJ:tJ, ") t.::.o LtJ'L, Coosella prolifera tJ: I? (JIC Holcacephalus
tenurus ()) 2 fifilc~ L -C Ii, 5ej:.~'-Plc=*j1rnm~~5JIj-t J.> ;:. c tJ~i'ifli-g-C''&.; I), .:t.n~lffJfifiJ'<~.""C"'&.;J.>c~~~timmTrans. Proc. Palaeont. Soc. Japan, N. S., No. 111, pp. 358-369, Sept. 30, 1978.

693.

ON THE GENERA SIPHONOFUSUS AND EUTHRIA OF
THE INDO-WEST PACIFIC*TSUGIO SHUTO
Department of Geology, Kyushu University, Fukuoka 812

Abstract. The taxonomic and stratigraphic relation of three gastropod genera
Buecinulum, Euthria and Siphol!ojusus is reexamined. The result shows that Bue·
cinulum and Siphol!ojuslls derived respectively in the Early and Middle Miocene
from Euthria which appeared in the Eocene. Sipholl0jUSUS, an Indo·West Pacificelement, reveals a morphoseries of enlarging shell and weakening sculpture in. each of its three lineages.

I.

later recognized by him (1921) as a synonym of Buccinulum. POWELL (1927) accepted IREDALE'S idea and gave a comprehensive revision of Buccinulum of New
Zealand. On the other hand, THIELE(1931) regarded M. comeus and B. lineatum as congeneric. When KURODA andHABE established (1952) and redescribed(1954) Sip/wno/usus, they did not make acomparison of that genus to any of theabove noted taxa, but compared it with
Manaria SMITH, 1906, from Arabian Sea.
As to the southeast Asian species inquestion, MARTIN (1895) and WANNER and
HAHN (1935) described some species witha columellar plait under the genericname of Siphonalia. While MARTIN (1914)
and OOSTINGH (1939) introduced Eulhria
djocdjocartae and E. bantamensis, respectively, AL TENA (1950) described Buccinulum oostinghi. BEETS (1942, 1944 and 1952)
described several species under Buccinulum and made a revision (1942) of therelated species from the Indonesian region.
MACNEIL (1960) referred both
BEETS' and ALTENA'S species together
with the species from Okinawa and Taiwan to A/er CONRAD, 1858 (type-species:

Taxonomic status of Siphollo(usus and related taxa

A group of morphologically very characteristic species, both fossil and living,have been described under various genericnames of Buccinulum, Euthria and Siphono/usus from the Indo-West Pacific region.
Euthria J. E. GRAY (in M. E. GRAY),
1850, with the type-species Murex comeus
LINNAEUS (0. d.), Buccinulum DESHA YES,
1830, with the type-species Buccinum linealum CHENINITZ=B. linea MARTYN (s. d.,
IREDALE, 1921) and Siphono/usus KURODA
and HABE, 1952, with the type-species
Siphonalia lubrica DALL (0. d.) are so
closely related with one another that
their taxonomic distinction is debatable.
IREDALE (1918) recognized the genericseparation of Alurex comeus from Buccinum linea tum on the basis of criticaldifference in shell form and discordance
in radular feature and placed them respectively in Euthria and Evame H. and
A. ADAMS, 1853. The latter genus was* Received Dec. 7, 1977; read Oct. 16, 1977
at Kumamoto.

358

693. On the genera Siphonof~~sus and Euthria
Fusus a/er: LAMARCK=Murex a/er GMELIN), and transferred, on that occasion,
A/er into the Buccinidae from the Vasidae, although THIELE (1931), Wenz (1943)
and many other authors treated thatgenus in the Vasidae. A/er is, however,featured by the low spire, laterally expanded body whorl and two columellar
teeth and is clearly distinguished not only
from those described by BEETS and ALTENA, but also from those described by
MACNEIL himself. SHUTO (1969) did notrecognized any distinction between Siphono/usus and Euthria and treated them assynonymous. It is not easy to give adefinite answer to the question whether
these taxa really represent respectively
one genus or not.
The type species of the above taxa
show some important common morphological features.
Common characteristics of Siphono/usus,
Euthria and Bueeinulum: Shell is solid,
moderate in size and fusiform with highspire and remarkably long snout. Axial
sculpture shows retrogressive development. Aperture is oval and acuminatedboth posteriorly to the well defined gutter
and anteriorly to the long and narrowcanal. Labrum is sharp at the marginand provided with threads inside. Inner

Fig. 1. Type-species of Buccinulum,
Siphonofusus and Euthl-ia. 1: Buccinulum
linea (MARTY:'-I), 2: Siphonofusus lubrica(DALL) and 3: Euthria cornea (Llf\:-;AEUS)359

lip is arcuately concave, covered by callus
and has a parietal ridge, which demarcatesthe gutter. Columella is somewhat twisted
and provided with a more or less distinct
fold at the entrance of the canal. Operculum is ovate with nucleus at theacuminated anterior end.
Distinction: Bueeinulum is devoid of
the axial sculpture of the teleoconch except on the very early whorls and provided with a somewhat shorter snout
than the others as illustrated in the typespecies (Fig. 1). The type species of Euthria and Siphono/usus show closely similar morphology in their early stage of
growth. In that stage, they have distinct
axial plicae reaching to both the upperand lower sutures. The axials become
weaker in the later stages. However,
weakening of plicae with growth is much
more exaggerated in Euthl'ia so that thepenultimate and body whorls, at least, ofadult shell of Euthria are devoid of theaxials. In addition to the difference inaxials, spiral sculpture of Euthria is much
weaker than Sip/wno/usus. Comparisonof the rhachidian radular teeth of Bueeinulum (Bueeinulum linea: THIELE, 1931)
and Siphono/usus (Siphono/usus lubrieus: KURODA and HABE, 1954) clarifies
remarkable difference (Fig. 2). The central tooth of S. lubricus has three small
cusps gathering at the central part of its
arcuately protruded frontal margin. The
lateral tooth has three stout cusps. While
Bueeinulum linea has triangular central
tooth with rather large three cusps. Its
lateral teeth are stouter and shorter thanthose of Siphono/llsus. Radular teeth,
particularly the central tooth of Euthria(E. queketti SMITH, BARNARD, 1959) is decidedly closer to Bueeinulum than to Siphono/usus.
Living species belonging to Buccinulum,
Ellthria and Siphono/usus are distributed
respectively in New Zealand-Tasmanian

360

Tsugio SHU TO

Fig. 2. Radulae of Siphollofusus andthe related taxa. 1: Siphollofusus lubrica(DALL) [KURODA and HABE, 1954J, 2: Siphonalia fusoides (REEVE) [KURODA and HABE,
1954J, 3: Euthria quaketti SMITH [BARNARD,
1959J and 4: Buccillulum linea (MARTY"')[THIELE, 1931J.

seas, Mediterranean-African seas andIndo-West Pacific_ Buccinulum dates fromthe Earliest Miocene in its home areaand some doubtful species (Euthria bantamensis OOSTINGH) has been known fromthe Pliocene of Java, Indonesia. The first
appearance of Euthria in the Indo-West
Pacific is the Middle Eocene of Java
(E. djocdjocartae MARTIN). COSSMANN(1901) listed three Eocene species andthree Oligocene ones of Euthria from theMediterranean region. Siphono/usus was
widely distributed in the Middle Mioceneof the Indo-Malayan region, but is unknown in Early Miocene and earlier age.
Its geographical range was expanded
northward in Late Miocene and later age.
Fusus contorta GRA TELOUP, which CosSMANN referred to Euthria, from theMediterranean Burdigalian may be classified to Siphono/usus. Anyway the oldest known occurrence of Siphono/usus isin the Middle Miocene. Paleobiogeographical distribution of these taxa, particularly the known first occurrence of thetaxa, suggests that Euthria is the groupof the oldest origin among them.
Euthria djocdjocartae is very suggestive
concerning the origin and phylogeny of
Buccinulum which appeared in the Earliest Miocene of New Zealand. E. djocdjocartae has a peripheral angulation anddistinct axial ribs on the first to fourth
whorls and then becomes smooth so thatthe body whorl is roundly curved at sideand provided with distinct growth lines
and a few faint spiral lines on the lateralside, otherwise it is almost smooth (Fig.
3). The snout is rather short. That is
to say, it shows an intermediate featureof Euthria and Buccinulum. It is strongly
suggested that New Zealand-Tasmanian
Buccinulum might have evolved from E.
djocdjocartae.
Siphono/usus may be another derivativefrom Euthria. The former appeared in
Middle Miocene both in the Indo-West
Pacific and Mediterranean regions as mentioned above, but its post-Burdigalian history is not clear in the Mediterranean.
It seems, at least, Siphono/usus did not
succeed in growing up to a major groupin the Mediterranean region. On thecontrary it flourished in the Indo-West
Pacific region. Geologic events in the
Middle East must have been concerned
EOCENE

OLIGOCENE

MIOCENE

PLIOCENE OUATERNARY...._ _ _ _ _ _ _ __EUlhria==~:----,_MEDITERRANEAN INDO-MALAYANMEDITERRANEAN-W. AFRICAN

Siphono(usus _ _ _ _ __INDO-W. PACIFIC

Fig. 3. Taxonomic relation and geographic distribution of the subgenera Euthria,
Buccil!ulum and Siphonofusus.

693. On the genera Siphonofusus and Euth1'ia
in the above history of the taxon. Geological history of the Middle East involves
events of closing Suez breach in Oligocene
time and only a limited interchange ofthe Mediterranean and Indian marine
faunas was probable mainly in Medial
Miocene time through Iranian-Turkey
channel. Larval type of Siphono/usus
must be lecithotrophic as suggested byits paucispiral and globose protoconch andin consequence dispersion into the Mediterranean beyond the ephemenal IranianTurkey channel must be very difficult.
Siphono/usus, therefore, is naturally regarded as a typical Indo-West Pacificelement which evolved successfully after
the Suez breach was closed. Not a small
genetic difference worthy, at least, ofsubgeneric separation may be postulated
between Euthria and Siphono/usus. In
conclusion, the present author prefers torecognize subgeneric separation among
Euthria, Buccinulum and Siphono/usus.
SMITH (1906) established Manaria with thetype-species Manaria thurstoni SMITH as agenus in the family Fasciolariidae. It has
fusiform shell consisting of more than eight
convex whorls sculptured by fiat axial ribs
and spiral lines. Its aperture is provided
with six to seven denticles inside the labrum
and an oblique fold on the columellar lip.
Accordingly it is morphologically very
close to Siphono/usus, but is distinguishedfrom the latter in having decidedly shorter canal. KURODA and HABE (1954)
reported a Japanese species which closely
resembles M. thurstoni in typical featuresof the shell. According to them, thisspecies, Manaria sp*., has a rhachidian
tooth, which is curved, laterally elongate
rectangular and provided with three small* KURODA and HAI3I, (1954, p. 230) cited
this species as Marania sp., but this citation
is simply an error for Manaria sp. as under·stood from their text.

361

cusps gathering at the middle of the anterior margin. Its lateral tooth has longand stout cusps at both ends and a minuteone between. The fact about the radular
teeth reveals firstly that Manaria shouldbe transferred from Fasciolariidae to Buccinidae and secondly that Manaria, thoughclosely allied to, is clearly distinguishedfrom any of Siphono/usus, Euthria andBuccinulum.
II. Evolutional lineages ofSiplwllOfllSllS

Species in question of East and South
Asia may be sorted into four groups.
They are Sip/wno/usus orangensis stock,
S. martini stock, S. chinensis stock andEuthria group.
Siphono/usus orangensis stock: This
stock includes S. wanneri (BEETS), S.
orangensis (BEETS), S. dinglensis (SHUTO),
S. oostinghi (AL TENA) and S. longicanalis(NOMURA and ZINBO). These species are
together featured by the solid shell withthe distinct spiral threads all over theshell surface and the prominent axial
plicae running from suture to suture.
Teleoconch whorls, at first, is concavoconvex in profile with distinct subsutural
band and convex lateral side, between
which is a narrow but distinctively concave band. Suture is somewhat cluspingwith distinct subsutural thread(s). Spiralsculpture is of adapical development in
ontogeny (SHUTO, 1969 p. 47).
The
spirals generally consist of almost equally
spaced and equally distinct five primary
threads on the first whorl. The uppermost one (Pl) is the distinct subsutural one, the second upper (P2) is on
the concave band and the third to
fifth threads (P3-P5) are on the convex
lateral surface. :Pl becomes broader andmay be split into two threads on thelater whorls. P2 abruptly becomes weak-TS1£gio SHUTO

362

er. Position of P3 is the peripheral angulation of the later whorls. The sixth
primary thread (P6) and so on appear
successively from the lower suture. Axial
plicae are counted about eight (rarely
seven or nine) for the first whorl andthey are increased in number graduallyto ten-eleven on the last whorl of theadult shell. They extend from suture to
suture not only on the early whorls but
also on the later ones. Growth lines arevery gently opisthocyrt with its prosocline
upper arm, slightly opisthocline lower
arm and the apex of the very shallow
sinus on the peripheral angulation. Species of this stock show a tendency of
weakening of plicae in later growth stage.
This tendency is more clearly exhibited

2

3

Fig. 4. 1: Euthria djocdjocartae MARTIN from Kali Poeroe, Central Java; Reg_No. St. 9173, Geo!. Reichsmus. Leiden; 2:
protoconch of the preceding specimen; 3:
protoconch of Euthria cornea (LINNAEUS)[COSS","A);N, 1901J.

in the species of later geologic age. Sizeof adult shell is larger in the species oflater age. The following species havebeen known hitherto.
Siphono/usus wanneri (BEETS, 1942), Middle
Miocene, Rembang, Java. Indonesia.
S. orallgensis (BEETS, 1942) , lower Upper Miocene, Kutei, Kalimantan.
S. dinglensis (SI-lUTO, 1969), Upper Miocene,
Panay, Philippines.
S. oostinghi (ALTENA, 1950), Lower Pliocene,
Java, Indonesia.
S. longicanalis (Nmll':RA et ZINBO, 1934), Middle Pleistocene, Kikai-Jima, Japan.

Siphonofusus martini stock: S. kelletiiformis (VREDENBURG), S. martini (WANNER and HAHN), S. indicus (DEY), S. macneili
sp. nov., S. japonicus sp. nov. and S.
lubricus (DALL) consititute this stock_
Species of the S. martini stock havecharacteristic shell with the whorls moreor less distinctly angulated peripherally
and with the axial sculpture being strongon the peripheral angulation and denuded
above and below it. Profile of the first
whorl of the teleoconch is rhomboid withsharp median angulation and the surfaces
above and below the angulation are fiat.
Development of the spiral sculpture isadapical. Basic pattern of the spiral sculpture is represented by a strong subsutural
thread (PI), P3 on the peripheral angulation and P4 between the angulation andthe lower suture. A weaker primaryspiral (P2) may be ready to appear fromthe lower suture in some species. Axial
plicae are seven or eight on the first
whorl and increased in number ontogenetically to nine or ten of the adult bodywhorl. The axials become weaker, particularly near the sutures. on the latterwhorls, while the spiral threads persist
their distinctness throughout growthstages. Weakening of the axial plicae ismore exaggerated on the species of693. On the genera Siphonofusus and E-uthria
youner geologic age. Growth lines are
similar to those of the foregoing stock,
but the opisthocyrt sinus is slightly deeper
and the lower arm is more protruded
opisthoclinally than in S. orangensis stock.
S. martini stock is distinguished, furthermore, from S. orangensis stock in being
provided with sharper peripheral angulation and weaker axials particularly near
the suture. The following species belong
to this stock.
Siphonofusus martini(WANi\:ER et HAHN, 1935),
Middle Miocene, Rembang, Java, Indo.
nesia.
S. kelletiifonnis (VREDE:-IBURG, 1924), Middle
Miocene, Myaukmigon and Thanga,
Lower Burma.
S. indicus (DEY, 1962), Middle Miocene, Qui.
lon, South India.
S. macneili sp. nov., Upper Miocene, Okinawa,
Japan.
S. japonicus sp. nov., Pliocene, Kyushu, Japan.
S. lubricus (DALL, 1918), Recent, Southwest
Japan.

Siphonolusus chinensis stock: Speciesof Siphonolusus chinensis stock exhibit
sharply angulate profile of the spirewhorls. Above this angulation is a gently
sloped ramp, on which the axial sculpture
is extremely weak. Body whorl is distinctly biangulated with a peripheral anda basal angulation. Axial plicae are about
six or seven on the first whorl and they
are increased in number to eighteen on
the adult body whorl. The spiral sculpture
is close set and weak above the angulation, distinct and subequal on the lateralsurface and unequal with weak secondaries
intercalated on the basal surface. The
snout is long and markedly recurved.
The species belonging to this stock show
some resemblance to Aler with defined
shoulder slope and sharp peripheral angulation, but have definitely higher spire andrelatively shorter snout than the latter.
Three species are referred to this stock.

363

Siphonofusus sp. Middle Miocene, Sedan, Java,
Indonesia.
S. dentifera (MARTIN, 1895), Upper Miocene,
Tji·Odeng, Java, Indonesia.
S. chinensis (MACNEIL, 1960), Pliocene, Oki.
nawa and Recent, Southwest Japan

Euthria teschi stock: Three species are
known to belong to this stock. They are
E. djocdjocartae MARTIN, E. teschi (BEETS)
and E. ovennanae (BEETS).
On E. djocdjocartae the spiral sculpture
consists of the subsutural thread (PI), a
thread on the peripheral angulation (P2)and a third one (P3) on the lateral surface.
Profile of the whorl is sharply angulate
at the lower one fifth of the whorl height.
The surface between the distinct subsutural band and the peripheral angulation is steeply sloped and very slightly
concavo-convex with an obsolete depression below the suture. The surface below
the angulation is remarkably receded.
Axial plicae are nine, eight and ten respectively on the first, second and third
whorl. They are strong on and below
the peripheral angulation but abruptly
weakened above and do not reach to theupper suture. The peripheral angulation
and axials persist on the early four
whorls, but on the fifth whorl they disappear abruptly. Spirals on the later
whorls are small in number, irregular in
spacing and extremely weak.
E. teschi and E. overmanae are closely
similar in the general morphology. Their
spires are high, acute and rather straightsided consisting of very slightly convex
whorls. Whorls are provided with finespiral threads. Growth-lines show shallow
sinus. The most obvious difference between them is suggested by the axial
sculpture. Morphology and developmental
pattern of the axials on the early whorlsof E. teschi are similar to those of E.
djocdjocartae, but the axials disappear
earlier in the former. E. overmanae has

Tsugio SHUTO

364
geological rancJe
species011'JO·I-::-E-'.'r.!~~:.;·r:-I••--1 P.

Q.

Siphono::'al'S:Jall'wr':: (Sel!tsl
ol'anile1:':i.;; Ulel!t&J
d::"a!cmll!..a t5huto)
ova~i.r",,;;i.

(Altcnol)

Z,m,Ji.<:.:lII:l!:''; (SOIIIUr4 et ::inbo)
l";':J'~::1!i (fl..:mer at 1Ia.'lnl
kcZZct;i~:':;,,,,,::,;; (Vradenburg)(Dey);'Uircll.S,-.aar4i.::: :>1'. nov.
J~pr:m::I,lI<,J sp. nov.
lllbJ"!<l'Wf (call)

dmlt;i:"".;;r.l (Martini

ahir... na:'s (Mact:cil)':'~jthria

tee:!;::' (l'eet.!llOl.'eJ ....aH;;;C IBeets)
djDI!d$Cr!r:ll·ta.C xartin
na:uiurensiu (Beets)bant~.:::r;8~.c (Oostingr.)

1Fig. 5. Geological range of species of
Euthria group and three lineages of Siphono/usus from Indo-West Pacific region.
Horizontal and vertical lines in the columns
respectively indicate the geological range
and relative shell-size of each species.

only weak axials on the earliest whorl.
In other words, the lineage of E. djocdjocartae, E. teschi and E. overmanae exhibits
an evolutional trend of earlier disappearance ofaxials in later species.
Diagnosis of E. madurPnsis is not clear
because of poor preservation of the specimens, and its phylogenetic relation to any
other species is not presumable.
E.
bantamensis is provided with coarse spiralthreads on all the whorls and with rather
distinct axials on early whorls so that it
may be regarded as an intermediate form
between Buccinulum and Siphono/usus.
III.

Biological aspect of Eutllria
and Siphollofusus

According to NORDSIECK (1968), Euthria
cornea (LINNAEUS), the type-species of
Euthria, dwells under stones of the rocky
littoral zone or on the vegetated rocky
bottom of nine to twenty-seven meters
depth, while BARNARD (1959) noted thatthe living and dead shells of South African
Euthria had been collected from the depth
range of twenty-fifty fathoms. The latter

author did not refer to the substrate.
Siphono/usus dinglensis, Late Miocene
species of the Philippines, is a minor element of the Philindophos dijki (MARTIN)Dendropoma javana (MARTIN) associationof muddy sand bottom of deeper shelf(SHUTO, 1969). S. japonicus from theUpper Pliocene of South Japan associateswith Anadara (Scapharca) castellata (YoKOY AMA)-Glycymeris rotunda (DUNKER)Amussiopecten praesignis (YOKOYAMA) association in the muddy sandstone, which
was presumably deposited on the shallow
shelf (SHUTO, 1962). MACNEIL (1960) mentioned that the Shinzato tuff member
and Yonabaru clay member, from which
Siplwno/usus chinensis and S. macneili had
been obtained, of the Shimajiri Group of
Okinawa might have deposited respectively
in bathyal depth of 200-400 fathoms and150-300 fathoms. S. chinensis was dredged
alive at the depth of fifty-nine fathoms
in the Korean strait.
Concerning bathymetric range of the
taxa in question, an idea may be acceptable that Siphono/usus is a dweller ofsand or sandy silt bottom of the middle
shelf and deeper zone in contrast to
Euthria, at least a part of Euthria, whichdwells on the rocky bottom of shallower
depth under higher energy environment,
although the data are not comprehensive.
This is concordant with the fact that all
the fossil species of Southeast Asian
Euthria, E. djocdjocartae, E. teschi, E.
madurensis, E. overmanae and E. bantamensis, lost their protoconchs and eventhe earliest part of the teleoconchs and areprovided with a septum at the top of thepreserved earliest whorl, although protoconchs of Siphollo/usus species are generally preserved in more or less observable condition. These facts apparently
suggest that Euthria and Siplwno/usus
differs from each other in their ecological
adaptation.

693.

On the genera Siphonoj'Us'Us and E'Uthria

Protoconch of Siphono!usus is paucispiral, consisting of about two (one and ahalf to two and one-third) smooth volutions. The nucleus is small, depressedand somewhat oblique, and the second
volution is globose. The boundary between the protoconch and the teleoconch
is indicated in most cases by abrupt appearance of the spiral and axial sculpture
or a curved, oblique and smooth boundary
riblet. Euthria has a quite similar protoconch to Siphono!usus judging from thefigures (COSSMANN, 1901 and BARNARD,
1959). According to Lo BLIANCO (1899),
the egg mass of Euthria cornea consists
about fourty semitransparent egg-capsulesof eight-nine millimeters long and four
millimeters wide, which contain respectively twenty-four ova on an average.
This feature is comparable to the eggs ofBabylonia and comparison leads to the
supposition that ova of E. cornea must belarge with a diameter of about 0.5 mm or
more, although he did not state the sizeof ova. In other words, Euthria cornea
must be a species of lecithotrophic larval
type and is potentially a species of quick
evolution and limited geographical distribution. Situation must be similar for otherspecies of Euthria and also Siphono!usus
and Buccinulum. This supposition is endorsed by the fact that many closely allied
species have been differentiated in a single
biogeographic province.
IV.

Species reference list

Siphonofusus wanneri (BEETS)
Siphonalia martini WAj\;1\ER et HAl-IN [pars],
1935, 2eitschr. Deut. Geol. Gesel., 87, (4),p. 222, 234, pI. 18, fs. 14 and 15 [excl. fs.
16-20].
Buccinulum wanneri BEETS, 1942, Leid. Geol.
Meded., 13, (1), p. 228, pI. 24, f. 4.
Siphonofusus orangensis (BEETS)
Buccinulum orangense Beets, 1942, Leid. Geol.
Meded, 13, (1), p. 225-226, pI. 24, fs. 10-13.

365

Siphonofusus dinglensis (5HUTO)
Buccinulum (Euthria) dinglense 5I-1UTO, 1969,
Mem. Fac. Sci., Kyushu Univ., 5er. D. Geol.
19, (1), p. 122-124, pI. 9, fs. 19, 22-24.
Siphonofusus oostinghi (AL TENA)
Buccinulum oostinghi AL TENA, 1950, Leid.
Geol. Meded. 15, p. 231, f. 19.
Siphonofusus longicanalis (NOMURA et Zlj\;(30)Siphonalia longicanalis NOMURA et Zlj\;130,
1934, Sci. Rep. Tohoku Imp. Univ., 2nd 5er.
Geol. 16, (2), p. 159, pI. 5, f. 26.
Siphonofusus martini (WANNER et HAHN)
Siphonalia martini WANNER et HAl-IN [parsJ,
1935, Zeitschr. Deut. Geol. Gesel. 87, (4), p.
222, 234, pI. 18, fs. 16-20 [excl. fs. 14-15J.
Buccinulum martini BEETS, 1942, Leid. Geol.
Meded., 13, (1), p. 222-225, pI. 24, fs. 1-3(and 5-6?).
Siphonofusus kelletiiformis (VREDENBURG)
Siphonalia (Kelletia) kelletiiformis VREDENBURG, 1924, Rec. Geol. Surv. India, 55, (1),
p. 66-67, pl. 2, f. 11.
Siphonofusus indicus (DEY)
Siphonalia (Kelletia) indica DEY, 1962, Palaeont. Indica, N.5., 36, p. 76, pI. 4, fs. 6-8.
Siphonofusus macneili sp. nov.
Afer aft. A. oostinghi MACNEIL, 1960, Prof.
Pap. U. S. Geol. Surv., (339), p. 76, pI. 3,
fs. 24, 25, 27, 28 and 33.
Siphollofusus japonicus sp. nov.
Siphonalia spadicea fuscolineata: 51-IUTO,.
1962, Mem. Fac. Sci., Kyushu Univ., Ser. D.
Geol. 12, (1), p. 36, pI. 7, fs. 5 and 9.
Siphonofusus lubricus (DALL)
Siphonalia lubrica DALL, 1918, Proc. U. S.
Nat. Mus. 54, (2134), p. 230.
Siphonofusus lubricus: KURODA et HABE, 1954,.
Venus, 18, (2), p. 95-97, fs. 16 and 20.
Siphonofusus sp.
2 specimens, Reg. No. Ge M. 556 with name"Buccinulum sedanense BErns" MS, Museum
Geol. Inst., Univ. Amsterdam
Siphonofusus dentifera (MARTIN)
Siphonalia dentifera MARTIN, 1895, Samml.
Geol. Reichsmus. Leiden, N. F., 1, (1), p. 96,
pI. 15, fs. 215-217.
Siphonofusus chinensis (MACNEIL)
Afer chinensis MACNEIL, 1960, Prof. Pap.
U. S. Geol. Surv., (339), p. 76, pI. 8, f. 19
and ? pI. 15., fs. 23-24.

366

Tsugio SHU TO

Euthria djocdjocartae MARTIN
Euthriajogjacartensis MARTIN, 1914, Samml.
Ceol. Reichsmus. Leiden, N. F., 2, (4), p. 142,
pI. 3, fs. 85, 85a, b· and c.
Pusio djocdjocartae MARTIN, 1883-87, Samml.
Ceol. Reichsmus. Leiden, 3, p. 104, pI. 6, f.l05.
Euthria teschi (BEETS)
Buccinulum teschi BEETS, 1944, Ceol. Mijnb.
N.S. 6, p. 14-16, fs. 1-6.
Euthria madurensis (BEETS)
Buccillulum madurensis BEETS, 1944, Ceol.
Mijnb., N. S. 6, p. 16, fs. 7 and 8.
Euthria overmanae (BEETS)
Buccinulum overmanae BEETS, 1942, Leid.
Ceol. Meded., 13, (1), p. 226-227, pI. 24, fs.
7-9.
Euthria bantamensis OOSTINGH
Euthria bantamellsis OOSTINGH, 1939, Illgell.
Ned. -Indi'e, (8), p. 116, pI. 14, fs. 242a and b.

v.

Description of species

Siphono!usus macneili sp. nov.
Afer aff. A.oostinghi: MACNEIL, 1960, Prof.
Pap. U. S. Geoi. Surv., (329), p. 76, pI. 3,
fs. 24, 25, 27, 28 and 33.

Type-specimens: holotype, USN M 562702(pl. 3, f. 33) from loco 17449, Kakazu,
about 900m south of Madama-bridge, Naha,
Okinawa: paratypes, USNM 562700 (pl. 3,
fs. 24 and 25) and USNM 562701 (pl. 3,
fs. 27 and 28) from loco 17451, north ofIwa, Kochinda, Okinawa, Japan. The
holotype is an adult specimen and theparatypes are juvenile.
Description: Shell broadly fusiform andmoderately small, attaining about 35 mm
in height in the adult stage. Protoconch
large and paucispiral, consisting of depressed first volution and broadly convex
second one. Teleoconch whorls about
seven, being bluntly angulated at the
periphery, which is slightly anterior tothe middle of the whorl. Axials prominent, from suture to suture and slightly
oblique to the axis of coiling in the young

stage, but weakened above the peripheral
angulation and becoming more vertical in
the adolescent and later stages. Spiralthreads close and moderately weak. Aperture rhomboidal in shape with distinct
posterior notch and long, narrow andstraight canal. Labrum teeth, parietal
ridge and columellar plait distinct.
Comparison,' The present species shows
diagnostic charact.eristics of Siphono!usus.
MACNEIL compared the above noted
specimens with Siphono!usus oostinghi(ALTENA), but the former is readily distinguished from the latter in its one-halfsmaller size, abruptly narrowed entranceof the canal, sharper peripheral nodules
and weaker spiral threads. Furthermore,
axials of S. macneili are weakened indefinitely earlier stage than S. oostinghi.
S. macneili is similar to S. japonicus sp.
nov. from the Pliocene of south Kyushu,
Japan. The former is, however, distinguishable from the latter in having a more
contracted base, shorter aperture and 7-8
degrees wider apical angle than the latter.
Horizon:Upper Miocene Yonobaru
Clay Member of the Shimajiri Group.

Siphono!usus japonicus sp. nov.
Siphonalia spadicea fuscolineata: SHUTO, 1962,
Mem. Fac. Sci., Kyushu Univ., Ser. D,
GeoI., 12, (1), p. 36, pI. 7, fs. 5 and 9.

Type-specimen: holotype, GK-L 4891 (pl.
7, fs. 5 and 9) from loco MI 6351, Hagenoshita, Takanabe-cho, Koyu-gun, Miyazaki
Prefecture, Japan. Depository, Department of Geology, Kyushu University.
Description: Shell moderately small andbroadly fusiform. Protoconch not observed.
Teleoconch whorls more than seven,moderately angulate at the median peri phery and slightly concave below the suture.
Axials prominent from suture to suturein the young and weakened in the later693. On the genera Siphonojusus and Euthria
stage except on the periphery. Spirals
close and moderately weak. Apertureelongately rhomboidal in shape, withoblique anterior canal and distinct andsmall posterior notch. Blunt columellar
plait and sharp parietal ridge defining theentrance of anterior canal and posterior
notch, respectively. Growth-lines sinuousand converging opisthocyrtly with upper
suture by an angle of 45-50 degrees.
Comparison:
Although the present
species is close to the preceding one, it isclearly separated from the latter as
mentioned above. Comparing with S.
kelletii/onnis (VREDENBURG), S. japonicus
has a less contracted base, weaker axials,
more sharply narrowed entrance of thecanal and more deeply sinuous growthlines. S. indicus (DEY) has distinctly
stronger axials and more oblique canalthan the present species. S. lubricus(DALL) has a longer canal, weaker parietal
ridge, and more defined infrasutural depression than S. japonicus.
Horizon: Lower part of the Takanabe
Member of the Miyazaki Group. Pliocene.
Siphono/usus sp.
Specimen: Ge M. 556, Museum Geol.
Inst., Univ. Amsterdam.
Locality: Environs of Sedan, Java, Indonesia.
Description: Shell thick, fusiform andmoderately small (32.4 mm in height).
Whorls angulated at the periphery andconcave below suture. Peripheral angulation blunt on early whorls and sharper onlater whorls. Sutures clusping. Axial
plicae 7-8 for a whorl, wider than interspaces, reaching to the lower suture andweakened above the angulation. Spirals
weak, dense and equidimensional above
the peripheral angulation and distinct andalternate below it. Body whorl large,
about three-fourths of the total height of

367

Fig. 6. Siphonofusus sp.
Museum Geol. Inst, Univ. Amsterdam, Reg.
No. Ge M556 (labeled a2 Buccinulum sedan.
ensis) .

the shell, sharply angulate peripherally,
contracted at the base to the very long,
obliquely bent and tapered snout. Lateralsurface below the peripheral angulation
provided with 10 sets of distinct andslightly weaker threads. Aperture rhomboidal in shape with sharp posterior notchbordered by a parietal ridge at the entrance and with long and narrow canalanteriorly. Labrum sharp with thread-liketeeth inside. Inner lip arcuate, covered
with distinct but narrow callus. Columellar plait sharp at the lower part ofthe inner "lip.
Comparison: The present species ischaracterized by its long, oblique andtapering canal, sharp peripheral angulation,
pronounced infrasutural depression andprominent plicae. In this sense, it resembles S. chinensis (MACNEIL), but theformer is readily distinguished from thelatter in having much stouter plicae andmore depressed shoulder slope. The present species differs from S. denti/era

Tsugio SHU TO

368(MARTIN), which has a distinct basal
angulation.
Horizon: Rembangian (Lower Middle
Miocene).

Works Cited
ALTE:"A, C. O. van Regteren (1950): The
marine Mollusca of the Kendeng beds
(East Java). Gastropoda, Pt. 5. Leid.
Geol. Meded., vol. 15, p. 205-240.
BARNARD, K. H. (1959): Contributions to theknowledge of South African marine Mol·lusca. Pt. 2, Gastropoda: Prosobran·chiata; Rhachiglossa. Ann. SO:J.th AfricanMus., vol. 45, (1), p. 1-237.
BEETS, C. (1942): Mollusken aus der Tertiar
des Ostindischen Archipels. A: Die Gas·
tropoden·Gattung Buccinulum im Ostindis·chen Archipel. Leid. Geol. Meded., vol.
13, (1), p. 218-229, pl. 24.
- - (1944): Die Gattung Buccinulum im
Altmiozan den Insel Madura (Ost Indie).
Geol. Mijnb., N. S., 6e Jrg., (1-2), p. 1416.
- - (1952): Reconsideration of the so·called
Oligocene fauna in the asphaltic deposits
of Buton (Malay Archipelago). 1, Mio·
Pliocene Mollusca. Leid. Geol. Meded.
vol. 17, p. 237-258, pI. l.
CONRAD, T. A. (1858): Descriptions of two
new genera of shells. Proc. Acad. Nat.
Sci. Phi/ad., vol. 8, p. 165-166.
COSSMANN, M. (1901): Essasis de Paleoconcho.
Logie Comparee. Tom. 4, p. 1-293, pis. 110. Paris.
DALL, W. H. (1918): Notes on Chrysodomus
and other mollusks from the North Pacific
Ocean. Proc. U. S. Nat. Mus., vol. 54,(2134), p. 207-234.
DESHAYES, P. G. (1830): Histoire naturelle
des vers et mollusques, p. 1-256. Paris.
(not consul ted) .
DEY, A. K. (1962): The Miocene Mollusca
from Quilon, Kerala (India). Palaeont.
Indica, N. S., vol. 36, p. 1-129, pI. 1-10.
EA:VIES, F.E. and SAVAGE, R.J.G. (1975):
Tertiary Faunas (revised), Vol. 2, p. 1447, London.

GRAY, M. E. (1850): Figures of molluscous
animals. vol. 4. London. (not consulted).
IREDALE, T. (1918): Molluscan nomenclaturalproblems and their solution, 1. Proc.
Malac. Soc. London, vol. 13, p. 28-40.
(1921): Ditto, II. ibid., vol. 14, p. 198208.
KURODA, T. and HABE, T. (1952): Checklist and bibliography of the Recent marine
Mollusca of Japan. p. 1-210, Tokyo.
- - and - - (1954) : New genera of Japanesemarine gartropods. Venus, vol. 18, (2),p. 84-96.
Lo BLiANCO, S. (1899): Notizie biologischeriguardante special mente il periodo dimaturita sessuale degli animali del golfo
di Napoli. Mitt. Zool. Stat. Neapel, vol.
13, p. 448-573.
MACNEIL, F. S. (1960): Tertiary and Quater.
nary Gastropoda of Okinawa. U. S. Geol.
Surv. Prof. Pap. 339, p. 1-148, pI. 1-19.
MARTIN, K. (1883-87): Palaontologische Er·
gebnisse von Tiefbohrungen auf Java,
nebst allgemeinen Studien liber das Ter·
tiar von Java, Timor, und einigen anderer
Inseln. Samml. Geol. Reichsmus. Leiden,
Bd. 3, (1), S. 1-42, Taf. 1-3; (2-3), S. 43184, Taf. 4-9; (4-5), S. 185-304, Taf. 1014 und (6), S. 305-381, Taf. 15.
- - (1895): Die Fossilien von Java. ibid.
N. F., Bd. 1, (1), S. 1-132, Taf. 1-20.
- - (1914): Die Fauna des Obereocans vonNanggulan, auf Java. ibid. Bd. 2, (4), S.
105-178, Taf. 1-6.
NOMURA, S. and ZINBO, N. (1934): Marine
Mollusca from the "Ryukyu Limestone"of Kikai.zima, Ryukyu Group. Sci. Rep.,
Tohoku Imp. Univ., 2nd Ser. (Geol.), vol.
16, (2), p. 109-164, pis. 1-5.
NORDSIECK, F. (1968):
Die europiiischnMeeres.Gehiiuseschnecken (Prosobranchia)
vom Eismeer bis Kapverden und Millelmeer.
S. 1-273 inc I. 31 Taf. Stuttgart.
OOSTINGH, C. H. (1939): Die Mollusken desPliocans von Slid·Bantam in Java. Ingen.
Nederlandsch.lndi"e, IV, Mijnb. Geol. Jrg.
6, (8), p. 103-119, Taf. 13 und 14.
POWELL, A. W. B. (1927): The recent andTertiary species of the genus Buccinulum
in New Zealand with a review of related

693. On the genera Siphonojusus and Euthria
genera and families. Trans. Proc. N. Z.
Inst., vol. 60, p. 57-98, pI. 1-4.
SI-IUTO, T. (1961): Palaeontological study of
the Miyazaki Group-A general accountof the faunas. Mem. Fac. Sci., Kyushu
Univ., Ser. D, Geol., vol. 10, (2), p. 72206, pI. 11-13.
- - (1962): Buccinacean and Volutacean
gastropods from the Miyazaki Group.
ibid., ser. D, Geol., vol. 12, (1), p. 27-85,
pI. 6-13.
- - (1969): Neogene gastropods from Panay
Island, the Philippines. ibid., vol. 19, (1),p. 1-250, pI. 1-24.
SMITH, E. A. (1906): Natural history notesfrom R. 1. M. S. Investigator. Series III,No. 10, On Mollusca from the Bay of
Bengal and the Arabian Sea. Ann. Mag.
Nat. Rist., ser. 7, vol. 18, (105), p. 157-369

175 and (106), p. 245-264.
THIELE, J. (1931): Handbuch der Weich tierkunde. Bd. 2, p. 1-778, 783 fs. Jena.
VREDENBURG, E. (1924): Indian Tertiary
Gastropoda, No.5. Fusidae, Turbinellidae, Chrysodomidae, Strepturidae, Buccinidae, Nassidae, Columbellidae, withshort diagnoses of new species. Rec.
Ceol. Surv. India, vol. 55, (1), p. 52-77,
pI. 1-5.
WANJ\ER, J. und HAHN, E. (1935): Miocane
Mollusken aus der Landschaft Rembang(Java). Zeitschr. Deut. Ceol. Cesel., Bd.
87, (4), p. 222-273, 5 taf.
WENZ, W. (1943): Gastropoda im O. H.
SClll :-<DEWOLF (herausgegeb.) Handbuch
der Paliiozoologie, Bd. 6, I, Teil 6, p.
-1201-1506, Berlin.

Buccinulum, Euthria :to.t LJ Siphonofusus 0)7t~J::0)~f*~f!f~jf-j J..,. -th 1?71'riiJJmO)!!I!~~f;;*i" G ;:: c. Mtfkllt71' I?~ I?.hG Euthria 71' I? Buccinulum 71'*~1ltfJJjtFlt.::.
Siphonofusus 71.*~1lt*jtFJl'::i*~ U.:;:: C ~!!JlI?71,t.:: U.:. -1:/ l':;t~¥$IRO)*-J.i'l&i¥JtJ:~*"C3.;G Siphonofusus 113 '?O)fiIl:*7"UI'.::7t71'.h. 4}*7"IJ·(,11ID!:0)*Jfntc~~~JO)~~ftc ~'?
Jfj~*7"U71'~/Jb I?.h G.

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

Sept. 30, 1978

New Series No. 111

CONTENTS
TRANSACTIONS

691. YANG, Seong Young: Ontogenetic Variation of Trigonioides (s. s.) paucisulcatus(Cretaceous Non-Marine Bivalvia)...... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 333
692. Hu, Chung-Hung: Ontogenies of Four Upper Cambrian Trilobites from theBonneterre Dolomite, Missouri ........................................ 348
693. SHUTO, Tsugio: On the Genera Siphono!usus and Euthria of the Indo-West
Pacific . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 358